Skip to main content

Full text of "Transactions of the Royal Society of South Australia, Incorporated"

See other formats

VOL. 118, PARTS 1 & 2 
31 MAY, 1994 

Transactions of the 

Royal Society of South 



Williams, W. D. Climate change and its implications for South Australia — introductory 

remarks ----------- l 

Schwerdtfeger, P. A brief overview of climate research ------ 3 

Allan, R. J. Modelling climatic change and variability ------ 9 

Williams, M. A. J. Some implications of past climatic changes in Australia 17 

Fitzpatrick, R. W. & Wright, M. J. Climate change and its implications for South 

Australian soils ---------27 

Bates, B. W., Charles, S. P., Sumner, N. R. & Fleming, P. M. Climate change and its 

hydrological implications for South Australia 35 
Harvey, N. & Belperio, A. P. Implications of climate change for the South Australian 

coastline _____--___ 45 

Tyler, M. J. Climatic change and its implications for the amphibian fauna 53 

Stott, P. Climate change and its implications for the terrestrial vertebrate fauna - 59 

Boardman, R. Some possible effects of climate change on vegetation 69 

Burns, M. E. & Walsh, C. The economic implications of climate change - - 83 
McMichael, A. J. & Beers, M. Y. Climate change and human population health: global 

and South Australian perspectives ------ 91 

Thumlert, T. A. & Austin, A. D. Biology of Phylacteophagafroggatti Riek (Hymenoptera: 

Pergidae) and its parasitoids in South Australia 99 
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 115 
Lange, R. T., Lay, B. G. & Tynan, R. W. Evaluation of extensive arid rangelands: the 

land condition index (LCI) -------125 

Prideaux, G. J. A small sthenurine kangaroo from a Pleistocene cave deposit, Nullarbor 

Plain, Western Australia -------- 133 

Zbik, M. The Cook 007 meteorite: a new H4 chondrite from South Australia - - 139 

Brief Communication: 

Read, J. L. A major range extension and new ecological data on Oxyuranus microlepidotus 

(Reptilia: Elapidae) ---------143 

Walker, S. J. Growth in the Australian burrowing frog, Cyclorana australis (Gray) (Anura: 

Leptodactylidae) --------- 147 






VOL. 118, PART 1 

Proceedings of a symposium entitled 

held on II November, 1993. 


CONTENTS, VOL. 118, 1994 

PARTS 1 &2, 31 MAY, 1994 

Williams, \V. D. Climate change and ils implications tor South Australia - introductory remarks 1 

Schwcrdtfeger, P. A brief overview of climate research „_..-. 3 

Allan, R. J. Modelling climatic change and variability - - - 9 

Williams, M. A. J. Some implications of past climatic changes in Australia 13 
Fitzpatrick, R. W. & Wright, M. J. Climate change and its implications for South Australian 

soils ------------ 27 

Bates, B. W, Charles, S. P., Sumner, N. R. & Fleming, P. M. Climate change and its 

hydrological implications for South Australia ----- 35 

Harvey, N. & Belperio, A. P. Implications of climate change for the South Australian coastline 45 

Tyler, M. J. Climatic change and its implications for the amphibian fauna 53 

Stutt, P. Climate change and its implications for the terrestrial vertebrate fauna - - 59 

Boardman, R. Some possible effects of climate change on vegetation - - - - 69 

Burns, M. E. & Walsh, C. The economic implications of climate change 83 
McMichael, A. J. & Beers, M. Y. Climate change and human population health: global and 

South Australian perspectives - 91 

Thumlert, T. A. & Austin, A. D. Biology of Phylacteophaga froggatti Riek (Hymenoptcra: 

Pergidae) and its parasitoids in South Australia 99 
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 - - 115 
Lange, R. T., Lay, B. G. & Tynan, R. W. Evaluation of extensive arid rangelands: the land 

condition index (LCI) - ------ 125 

Prideaux, G. J. A small sthenurine kangaroo from a Pleistocene cave deposit, Nullarbor Plain, 

Western Australia --------- [33 

Zbik, M. The Cook 007 meteorite: a new H4 chondrite from South Australia - 139 

Brief Communications: 

Read, J. L. A major range extension and new ecological data on Oxyurunus mic role pi dolus 

(Reptilia: Elapidae) - - 143 

Walker, S. J. Growth in the Australian burrowing frog, Cyclorana ausnalis (Gray) (Anura: 

Leptodactylidae) - - - 147 

PARTS 3 & 4, 30 NOVEMBER, 1994 

Davies, M. & Watson, G. F. Morphology and reproductive biology of Limnodynastes salmini, 
L. convexiusculus and Megistolotis Ugnarius (Anura: Leptodactylidae: 
Limnodynastinae) _________ [49 

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 --------- 171 

Pamment, D., Beveridge, I. & Gasser, R. B. The distribution of nematode parasites within 

the stomach of the western grey kangaroo, Macropus fuliginosus - 187 

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 - - - 197 

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 ---------- 207 

Hutchinson, M. N., Milne, T. & Croft, T. Redescription and ecological notes on the pygmy 

bluetongue, Tiliqua adelaidensis (Squamata: Scincidae) - - - 217 

Fuller, M. K. & Jenkins, R. J. F. Moorowipora chamberensis, a coral from the early Cambrian 

Moorowie Formation, Flinders Ranges, South Australia - - - 227 

Peterson, M., Shea, G. M., Johnston, G. R. & Miller, B. Notes on the morphology and 
biology of Ctenophorus mckenziei (Storr, 1981) (Squamata: 
Agamidae) ----------- 237 

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-" - - - - - 245 

Olsen, A. M. The history of the development of the Pacific oyster, Crassostrea gigas (Thunberg) 

industry in South Australia -------- 253 

Bird, A. F. & Yeates, G. W. Studies on Aprutides guidettii (Nematoda: Seinuridae) isolated 

from soil at Northfield, South Australia ------ 261 

Brief Communication: 

Tyler, M. J., Davies, M., Watson, G. F. The hylid frog Litoria alboguttata (Giinther) in 

the Northern Territory --------- 267 

Insert to Transactions of the Royal Society of South Australia. Vol. 118, parti 3 & 4. }Q November, 1994 




ByW. D. Williams* 


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. I 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. 

Jhmwtitm of the famtl Stittfto •>* S. i/»,w (WMl IIX<k 10. 




1 1 is my privilege, as l*re.sKk-ntot the Royal Society 
of Soulh Auslralia, to introduce dais important 
symposium on climate change and its implications for 
Soulh Ausitaha. 

In doing no, I wish lusi !o welcome all participant* 
anil I extend * particular welcome lo Sir Kijtk 
Qliphant, an Honorary bellow of our Soeiety. Our 
piitmn. Her Excellency the Governor. Dame Roma 
Mitchell, has indicated lhai Vice-Regal commitments 
preclude her attendance I trust that alt panic tpams. 
sitidemv disiinguixhed scientists, or mere mortals like 
me. will find the symposium interesting, useful and 
a constructive addition To local debate on this mosl 
important mailer. 

The Royal Soeiety of South Australia held its fust 
meeting m November 18811, shortly after f,>ueen 
Victoria assented to the use of the wold Royal The 
Society, however, succeeded ihe Adelaide 
PliiKs-mphieal Society lonned much .'arlici, and indeed 
noi long .titer Ihe loundation of the Slutc of South 
Australia It was formed in 1853. Since those early 
years, ihe Society has mel regularly, published 
SClctitifk papers, maintained an extensive libnuy. 
awarded granls for research or awards lor meritorious 
contiibuiions to scienee. and its Fellows have included 
most of South Australia's best known scientist* 

The overall objective of the Society is the promotion 
and diffusion of scientific knowledge, and it is in 
support of ihis objective that the present symposium 
has been sponsored. I need hardly remind litis audience 
that any change to the climate of Soulh Australia will 
have profound, significant, and comprehensive effects 
on the South Australian environment, economy, social 
structure and public health - lo name but ihe most 
obvious features thai could be affected. It is to orntg 
1 his to Ihe allention ot Ihe community m general a;al 
its decision makers in particular thai Ihe present, 
symposium is being held. Such an aim tails squarels 
within tlic overall objective of ihe Society. To promote 
farther ihe views expressed, the papers presented arc 
heing published in this part of the rmnsifcttor)< tifihe 
RiAul $\KWty of Soulh Australia and are also available 
separately as a book, 

Ol course, Ihe symposium is only one of many thai 
haw been and are planned to be held lo discuss cl mate 
chrnigc At the international level, numerous meterings 
have been held, several .scientific journals specifically 
cartel icsearxh on clonaie change, and I here C5 .tn 

l)v|t:Mltnenl nf /nulo»y. University of Adelaide. Soulh 
Austndta 5005. 

international commniec which reports regularly (IPtX- 
Intergovernmental Panel on Climate Change). 
Likewise, at the national level, considerable activity 
prevails. Indeed. Ihe Australian Academy of Science 
is currently sponsoring a meeting in Canberra on the 
subject of climate change, The National Greenhouse 
Response Strategy was finalised in WV2. and at the 
Suae level, various governmert agencies maintain at 
least a watching brief -an Ihe subject, a number •>♦" 
natural and medical scientists, economists and others 
arc aclively interested in ihe subject, there is a Climate 
Change Committee (serviced by the Department iM 
Environment and Natural Resources and including 
representatives (mm a wide variety it\ government 
departments) which reports regularly to Cabinet and 
has published several important documents (e.g. South 
Australian Climate Change Committee IWO, W\) and, 
just a tew years ago ( WK#K a wide ranging conference 
was held in Adelaide on Ihe subject of climate change: 
1 refer lo Greenhouse '#o'; Planning for Climate 
Change &kl(&dfe Conference (Dendy 1989). f-'inally, 
I note the publication in November RJ93 of South 
Australian Greenhouse News , Vol. I. No 1, the first 
o\' what is iniended to be a regular series of newsletters 
with particular emphasis on South Australia. It is 
published by the South Australian Department ol 
Environment and Natural Resources with the support 
i>f the Office of hncrgy. 

Against this flurry of recent and ongoing activity one 
might ask, why should the Royal Society sponsor yet 
one more meeting to address the subject? Whal 
possible good can arise from another ''mkfpxfi 

The answer is simple. Fusu researvh and views on 
this subject pmceeds apace and there is a constant need 
io provide an opportunity for Mich research to be 
regularly aited tor die benefit of the wide? community. 
h is indeed half a decade since the lasi major meeting 
took place in Adelaide to discuss this matter. 

Second, ihe Royal Soeiety is quite independent of 
government departments, research institutions and 
universities, and k therefore in a unique position - 
indeed, has a special responsibility - to provide an 
opportunity for views to he aired which may not 
necessarily conform to the pam line, euiiem ideology 
oi generally accepted scientific views As will become 
obvious in some of the papers to be given, there ate 
views held by some which do not agree with 
widcspicad views. | Sir Mark Ohphani, in a comment 
at the conclusion of the Symposium potnicd to a rveeui 
ariiele in Nature, hmrt which pointed to the lack «il' 
any evidence of change in the present climate,) 


Ft 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 to plan ahead tor possible 
change, which may. in any es'ent, be more rapid than 
predicted. A recent article in the New Scientist (Nielsen 
1093) 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 tew degrees warmer than now may 
change very suddenly to become either significantly 
colder or warmer. In other words, we may be forcing 

the present climate into an unsteady state when large 
natural changes in climate could be triggered by 
relatively smaJl 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 to 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 !he 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 Margaret 
Davics, 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- 


Lroncri , I (1993) Who will underwrite the Hurricane? New 

Scientist, 7 August 1993. 29-33. 
Zit.LMAN, J. f 1993) Memorial lecture. University of 

Melbourne, 3 September 1993. (Unpublished) R. H. (1993) Chill warnings from Greenland. New 

Scientist. 28 August 1991 29-33. 
Df.nt.iv, T. (Bet.) (1989) "Greenhouse '88: Planning for 

Climatic Change Adelaide Conference Proceedings" (South 

Australian Department of Environment & Planning, 

South Australian Climate Change Committee (1990) 

"Implications of Climate Change tor South Australia*' First 

report of the Climate Change Committee. August 1990. 

(Department of Environment & Planning. Adelaide) . 
_____ (1991) "The Greenhouse Strategy for South Australia" 

(Department of Environment & Planning, AdeJaide). 


By Peter Schwerdtfeger* 


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. 


by Plicr ScHWERpTfhunR* 


SfHWCKPTniotK, P. |RKW) A brief overview of climate research, Tnins. ft, Sot: S >lu\i IIX(l), J-7 31 Mh\\ 19M4 
The problems liiLing those attempting ir> generate reliable prognostic climate model* is formidable Good estimates 
ol trends in conccntrwiion of q|1 radiating atmospheric gases are necessary and these must he entered into a touted 
numerical model incorporating all of the important feedback processes. Included in these are tlie parallel processes 
nev'v'rrmu in Ihe oceans The xiirtplesi text of the prognostic prowess of a model is ID run it backward:, in lime 
a simple teM ul' veracity that nm noi supported any modern long-term climate prediction model, 

KEY WORDS: climate change, models, radiating atmospheric pases, fncteomiogv. 


Ovet a third of a century ago. I found myself as a 
freezing, fledgling gco-scicntisf, upprehensively 
prodding the fro/en crust of Arctic sea ice bcluw, in 
order to render advice on our mutual safety to my Red 
Indian companion, who gingerly guided the hungry 
dogs that, from meal to meal, dragged the sled which 
held our food and my bare-bones scientific equipment. 
We knew thai we were venturing close lo lite brink 
uf cbacn when the supporting ice became less than .six 
inches thick My yelling out "150 mm" would only base 
generated dangerous confusion. The metcorologieaJ 
parameter ol greatest concern to me then was the 
profile of temperature from the surface downwards, 
and for (he following ten yeans, i remained interested 
in ice-bound temperatures in the Antarctic as well, and 
pondered on how lltese arose, At the time. ihc;>c weic 
genet ally tegatded as activities unbecoming of 
atmospheric scientists, but with the growing relevance 
attached to the new sctcncc of climate, the perceived 
significance of the extent of polar ice, and its thermal 
as well as physical content has been dramatically 

During the last 25 year*, 1 aarted to raise m> head 
.and examine the way in which Ihe radiant energy 
incident on the Earth's surface was. transferred not only 
lo depths below but also to the atmosphere above. In 
the 1960s, many "real" meteorologies referred 
indulgently to the "boundary layer boys' ta phrase that 
the well known Princeton University theoretical 
meteorologist Joseph Smagorinsky used in a tdk in 
Melbourne, when he unveiled the then excitingly new 
results of numerical 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 
transformation of energy from the Sun inlo other 

• Institute for Atmospheric and Marine Science. Hinders 
University (if South Australia, GPO Box 2100, Adelaide 
SA MKIl ' 

radiative, conducted, conveeiive, and evaporative iorms 
takes place at the Earths surface. Apart fiotU the 
conducted heat which involves the material below the 
surface only, the ihree remaining fluxes drive l hut all 
important distribution tottc. the diurnal boundary layer. 
the thickness of which can vary from millimetres lo 
kilometres, depending on the season and time of day. 
Without the critical knowledge of the flows of energy 
into ihi> boundary layer, resolved for a sufficiently 
finely spaced grid coveting the area being modelled. 
every numerical meteorologist would he marooned. 
Therefore for some lime. I continued to feel happy with 
the relevance of my work, delving in the lowest 10m 
of aimosphyrv blanketing the surface of the Larth. 

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 group. I became persuaded thai there was 
more 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 atmosphere, especially when the 
possibility ol both making and viewing the icsults of 
actual physical measurements in real time allows 
important physical connections to be grasped. More 
than ever now. my heart bleeds for those meteorologists 
who spend all of their windowlcss lives hypnotically 
hunched in from of iheir computer screen.\. 

The science of the wather was, in earlier years;, 
often conveniently divided into metWW$Vgy% which 
involves statements about auv.Osphcrie conditions ill 
various given times, including tlic future, when 
concepts of fonuwititf are involved, and c!imaiofog\\ 
which is concerned with quintessential summaries or 
meteorological conditions over specified places and 
periods, Applied sciences arc strongly driven by public 
perceptions and wishes, in which matters, of course. 
sctenltsts arc increasingly not averse, tl not forced, to 
offer their guiding influence, The last decade flirt seen 
the more geographical science of climatology fall into 
relative disfavour as the persuasive value ol the study 


ul'i Itnuui . in Ifwing linked with Ihc (Itnmn of inexorable 
tiends and mystery Ul extremely lunc-ienn lurcvasis. 
has tKSCtl hcr.iklcO u> an essential focus or serious 

Newspaper and tcleviMon have popularised Ihc 
iub*,ea nf meteorology throueh I he regular presentation 
ofweathei forecasts, and satellite images of ihe haul) 
have helped « ready in gcneraling an appreciation of 
ilic global naUiie ol metcorol"gical processes reflected 
in vivtblc cloud structures. lheie is no ihuibl (hat an 
advance knowledge of details of nieicon-ilogiuil 
conditions N important in many areas ol human 
endeavour Over relatively short terms and f^ir specific 
ar<*as such as landing grounds, aviation safety depends 
toll it. ftl*W.a>tnig '^ so uiflueoees many other 
lommcrcial operations, including those connected with 
agriculture, the nWUluctUpfl Ol icecream and drinks. 
ihc selection Of routes tor shipping as well as personal 
dunces regarding clothing and recreational activities 
A; a result, the concepts nf nwtcorolo^y and jhtvvustifif* 
have become alino.M synonymous in the pnhlic imnd 

However, more general cliinntological summaries 
also help lo guide the public, particularly travellers, 
even though a little caution is teefuircd when !■ h » little 
detail is ollered. For example the halt! statement that 
Adelaide's mean January temperature i^ 23*C could 
easily pfove deceptive to a irme visitor, uhen ii is 
known that (lie mean maximum lempciatuie is M) Q C 
and rhc mean minimum ib°C wiih extreme values or' 
these two parameters of almost 48 °V and 7%^ ha* nig 
heen ^corded, fcquipped with this additional 
iriloimation. rhc intending traveller might obtain some 
idea twto whai types of clothing to pat.k An even mure 
useless piece til ".sianti -alone" cliniulologieal 
information,, cvcepl perhaps to wine-makers needing 
lo be assured ihnl the teniperatutcs mainlaincd in deep 
cellais me beitcfiuuJ to (he maturation trf their 
products is thai Adelaide's mean annual tcmpcialurc 
is |7% Kcsputisiblc and useful elimatnlogical 
compilations usually aun to provide envelopes for 
ranges of values wuhin which parameters of interest 
might be expected to lie, The choice of these 
paiaoieters as well as rhc time scale will depend -on 
the purpose ol the application. For example, ftw 
agricultural decisions inforuiation about mitildl/ 
U'tti{HT\>\nt*, Miiif nt<1iatttitt and uintltnrss l.uidudmv' 
likely extremes) is important. In tlte liylit of the present 
knovvlcdee of the Earth's regional elnualcs, it is easy 
10 overlook Ihc achievement \)l colonial planner such 
as Colonel William Jaghi. the founder ol' Adelaide, 
who within six weeks o! artival. barely in advance of 
die lirsl Furttpeiin sellTers. was able to assure himseli 
is lo the general naliire ttf Adelaide's chaiate and n» 
icalisc (I ml tl was the only location near the coast of 
whai had been designated a.s South Australia where 
j city based \m l l )th century technology cxitiUI be 

The need for information on weather in general 
terms is readily accepted for .seasonal I'orecaMs for 
agricultural and water resources planning Inevitably 
because ol the historically obseived link between 
protracted cstremos in meteorological conditions and 
the economy, the possibility Ml reliably usccrtammv 
lone -term trends tsan attractive proposition, Ihcrciu 
lie the origins of the clamour for. and the justificaliun 
of, modern climate research- 

Meteorological pioces^es occur on many difieivnt 
lime-scales. The Iwo most familiar of these arc ihc 
variations in insolation which resuit in the annual c>\k- 
ll|' seasons and ihe daily risme and sctiina of ihc sun 
VMh are associated with systematic insoiaiirtn chariec^ 
v\IiilIi resull in every other meicorological paratnelei 
experiencing cui^cc|ucni variutious. rimnucs on otlict 
irnic-scjIcH are fat le*.s piediciahle. At the hieh 
Ir-tiucncy c\m\. luilnik-nu- is Km dcHtibed by nsiii>: 
stalislical expressions- Low frequency meteorological 
changes h;r.e been discovered in geological studies of 
ihe histoiy nl the liarth with dislinct climatic events 
being separated hy t'ron) thousands lo inillions of years 
On Iffts >^ale, geologists and glacioloj:tsts have Itnv 
led meiconMogists with the intensity ol rheir interest 

Systematic peritxiic mejciMlojiical changes can hi* 
summarised by the chmaloloey ol the periods uudei 
consideration. Prior to the development of modern 
methods of recording meteorological values, ihe 
descriptions of historians, ineludine the authors of the 
Old Ti'sutnwnt olfcr mvesltgators ol past climates a 
wealth of information, hor examjile, dui'mg his yeais 
in f^ypt, by warning of the seven years of famine !♦• 
follow years oj plenty. Joseph issue*] his Pharaoh with 
the first recorded successful long range svcathci 
toiecust. \Uc Litr/f Ac A$H of Ihe Middle Ages is a 
well rccoaled historical fact, easily verifiublc througf. 
Mudtcs \A the adVanCC and tetieat ol Alpine elatiei>. 
t )nly in Ihe 20ilt ceniurv have nune-shauV last worked 
sornt: fitK) years ago. hectmie visible aeam with the 
retrdat »tf these glaciers which be»an in me 19th 
cerau.v vloilern visitors to the Belvcdore Hotcf 1 
^hicli. a hundred years a^u dranun,eidlv Mverin-»keU 
Uic Khoitc Glacier in Svvit/ei'land. may well ponder 
on how long powerful binoculars will sutlicc to oiler 
a glimpse of the glacier's snout liillowinjj its drammi*. 
recession. It was this same "Middle A^es" ice age 
which contributed, substantially lo the elimination of 
ihe ViKinji colonies from Greenland at nhout the tftgTtt 
time that Christopher Columbus was busily divertinj' 
buropean atieution to warmer parts of the New World. 
Urysoti & Murray (1^77/ offer accounts of elimaU 
changes and the consequences thereof to various 
ovihsaiions over several thousand yeats ol documented 

There isabsohikly m*dnubi ihat the climate ttf Hit 
fctrtii is and ha-; always been changing (Brooks l°J») 
fluwoer. wltuL has also been dunging during the lasi 


two decades is the public perception Of ihc nature oi 
climate changes and concern thai the contribution to 
these by human aciivuics may *• really overshadow 
natural random trends. Quick |o respond 10 such 
concerns, many atmospheric scientists have soughi to 
quantify and offer physical explanations fin* both 
pnrn.iry ohscivaltons and assumed consequential 
changes. Changes of climatic pammelers littCJShrttlg 
"naturally", are on seemingly random tunc scales 
because of ihe dil'icrent rates governing the physical 
feedback' systems which link most, if not all. 
professes in Hie atmnsphcic. I he vast r;tn£e or these 
processes, and their intcrrMioris. orlei Ihe pet feet 
recipe for chaos. 

I he subjection of pasl climatic data to IreqUency 
analysis in (he hi>pc of obtaining cyclically tepetittvc 
messages has been in vain. My Flinders University 
colleagues, John B)'C. Roland Bvmn-Scoti and .\dinm 
Goidon furvc demonstrate*! UtH phenomenon of 
randomness ill an unusually direct way. The) ha^e 
translated random levels ol net- insolation, by means 
ol a simple, physical computer model in which 
atmospheric and oceanic momentum arc plausibly 
coupled, into time series nf air ami sea temperature 
'.•.Inch over a period oi' several simulated centuries 
display an uncanny superficial resemblance to actual 

We a* the harth to be a simple, heat conducting 
hollow shell, vimilai perhaps to early artificial 
satellites, bill not even containing a suttenngdug, tlicn 
titaeorotogy would be simple, the surface temperature 
would be the only reportable factor, and it could be 
».dctil:iled from a knowledge of the Softft ron.Vii/n 
I which in facl is not i|tiile constant! and the measurable 
radiative properties ol the satellite surface The resuli 
is commonly referred to as the plitnetary icmpetvture 
and the Iwo important planetary tadiativc properties 
111 which this temperature is related are the alhedo or 
surface reflectivity toward the wavelengths of solar 
radiation, which arc predorn inanity visible and the 
planet's emissivify, a factor which determines the 
ellleicney ol the export of radiative energy, mainly in 
the infra-red range -J' wavelengths, out to space 
Oiherwise, this loss of radianr enemy k determined 
only by the tcmpciaturc a fact discovered by Newton 
three centurie* ago and quantiiicd by Stefan and 
Boll/mann late last century. This relative 
meteomtngical sirnpheity is further enhanced by its 
insiaituweiiy: a thin hollow planetary *he.ll cannot store 
any heal energy and the coupling of temperature and 
rudiational events occurs without any delay. 

By virtue of its atmosphere, not to mention its solid, 
heat storing mass, out real harth has made this simple 
business far more complicated. For example, the 
diurnal delay between maximum insolaiion at the local, 
ueoetaphieal noon and the maximum air lemperaiure. 
which may occur 2,5 houis later is almost entirely 


ascribiible to Ihe thermal admittance or conductive- 
rapacity ot the ground. I he Earth's surface ilscifdoes 
not have a uniform albedo; this varies fnnn 3?4 9? over 
large pads o| the ucean to SO- 90% rttfer freshly &Wen 
snrAv The level of heat storage depends .in the heal 
capacity and thermal conductivity of the marcrul 
beneath the l:arlhs surface, but these conductive 
processes cannot compete with the ctliuency of both 
horizontal and vertical encr£\ transfer mechanisms 
within the Earth's dynamic atmosphere Viewed from 
sp3cc. the clouds which result from rhc presence of 
water vapour are the most dominating and reflective 
feature. Clouds are largely responsible lor the Earths 
average albedo being about 54%. A.-* in the case ot the 
hypothetical, hc>llow. atmospherelcs^ plrtnetdi>cus>ed 
carlieT. the 6h% of the incident solar energy which 

accepted by the Earth can also be used as the basis 
foi the calculation of a steady state pltincrars 
temperature, in tact ibis turns out to be ribout 22°C 
Although this figure has real meaning m ictnis of the 
Earth's temperature when observed from outer space. 
it is difficult, unless resting on a polar ice floe, to 
appreciate its signilicaucC back on Patth uselr where 
ihe aieally integrated mean surface temperature is about 
I5°C The corresponding figure in upper nirncspheTic 
lewis which still emit, significant amounts of infra-led 
radiation is about 4$PC. It is esscntiu) TO realise that 
the planeiaiy temperature must be regarded as a 
radiativcly weighted mean over die emuc ihirknexs or 

1 he atmosphere. 

The atmosphere, beciiusc- it contains polyatomic 
Sascv which pariiaJly absorb and re-emit ihor 
characterising band* ol mfta-red radiation which the 
Earths surface transforms from absorbed incident sola! 
radiation, greatly modifies the simple, ^-dimensional 
piciute of planetary temperature, Simpson (1927. 1928, 
W2 U ) was one of Ihe first to quantify, in terms ot 
spectrally selective absorption, the Greenhouse Ejjcvi 
winch is essentially responsible lor the mean 
temperature at the surface of the Earth bcinx 
uppro\im<iiely 37°C warmer than the planetary 
temperature. The over-whelmingly imporwm gas in 
this radiative process is water vapour, H : 
Unde islanding how this process contributes to the 
distribution nf temperature vvithin the aimo'-phere- is 
difficult enough in itself, because Ihe concentration of 
this vital vapour varies with both locution and urue 
Its concentration and temperauitc in turn determine 
the existence of cloucK, which increase the reflection 
*M in-coining solar radiation, thereby generating the 
fundamentally important feedback process of ttic 
F ; .arth's >urfaee hccT.mitng shaded below, thereby 
ledueing its lemi'kenilure and oDtiscquenil y atsd the level 
of evaporation. All of these linked steps occur neither 
instantaneously nor at rales which can be uniquely 
--.peeified The physical clarity ot the eiilitv pfOt.w- 
which is most readily comprehended in a single. 


vertical, dimension, becomes obscured hy large scale 
horizontal advective processes, ol which the: winds near 
the giuund and j Movement of clouds above are Ihe most 
obvious' manifestations The variability ol H.O in our 
atmosphere results from the f&CI that it readily changes 
Irnm \upour lo liquid and even .solid phases vvilhin ihe 
range of temperatures encountered on Enrth,. all forms 
having strikingly dilfcrcnl mobilities. 

I here are other yases which c»»mrihulc to the 
atmospheric Grct'nhottse fffi'ir, which il should be 
emphasised, rs significantly dillcrem Iron) the market 
garden type ihe latter having enclosing impermeable 
membranes to inhibit Mb collective nml water vapour 
losses fmm Ihe system, The additional «;im-s arc also 
puly-amniic.aluit is ihey have three or moie atoms per 
ivmlevolc and are present [n tonceni rations which ate 
Miiall compared to that Of water vapour In the absence 
of strongly localised sources ol product inn and ol sinks. 
which absoin Item, these gases OK considered to be 
urntnrrnh mixed wilh ihe niher. dominating gases of 
(lie ainiospheie. such as wcygen and nitrcigcn. All o( 
the early radial inn model* nf the atmosphere in 
focussing K)n the importance of water vapour, deigned 
to consider only cirhon dioxide, CO-. US being ol even 
secondary importance in ihcii scliemes. Other 
members of Ihe atmospheric family of poly-atomic 
eases, including methane, oxides of nitrogen and ozone 
wore regarded as negligible in inlluencv. 'Ihe 
importance accorded these eases chanced dramatically 
following the first lew years nf observations, when 
ihcse became available during the W7£k I nam a global 
uuwork nt mnimonng siauons which delected steady 
tm. reuses in the menu annual concentrations .if 
particularly COj and L\i A in Ihe atmosphere. (e.g. 
M.I.I i*/7)) Those ate readily explicable in terms or 
the ever increasing comhustion ol fossil fuels, the 
tolling and binning o( the world's remorselessly felled 
remnant forests, and the flnlnlence nf ihe world's 
huiveomng herds nf tame runiinuuiv Jt is reasonable 
|0 accept (hat with significantly greater concentration 
levely these gases should ot included in the accounting 
ol niditUive energy transfer and heating. The success 
H ith which this can be done also depends' on the ability 
lo ideulily Ihe nature and magnitude ol all of the 
leedbikk processes 

With the acknowledged increase ol the "lesser'* 
ladiutively important gases, arises the question as to 
the means thereby the atmosphere js also <|l>lc 10 
continuously shed part of the concentration, Thai this 
nnol always have been so is clear since CO-, and Of, 
have both been released Hi the air at least since life 
began on hurtli. even m limes when their mean 
concentrations were relatively srubte On Ihe basis nf 
current estimates of the global roles o^ COi 

'Jr-s^i-,, m 1 I) M%2>; PtrP, Thesis. MtfcnMup 
Depaiunent Urnvetsaiv ot Melbourne L'npnhl 

production and knowledge ol its actual concentration 
in the atmosphere, u is a Mmplc matter to calculate 
the rare nl one of ihe Karth.s leedhaik systems 
is dealing wjth the "fis-iru/* of ever increasing loads 
of this gas by, iot example pmduung cnrbonales as 
U resull nl Ihe dissolving Oj the CO- in water This 
ha> long bren a fertile field ol" investigation lor 
geologists. However, it is lift* apparent that this 
process is not coping with all of the CO, curreoil> 
bcuie produced. so ihat its atmospheric eonceulfatioii 
is continuing to increase, 

In summary, the pnablcms fating Ihnsc aiming lo 
generate a reliable prognostic climate model are quite 
formidable. It is necessary lo have good estimates of 
ihe trends in concentration ot all radiating atmospheric 
gases ami lo ensure lhal these arc entered into a tested 
numerical model which incorporates all ol Ihe 
miportam feedback processes made even more comple\ 
by the appreciation in recent years that models of global 
atmospheric circulation cannot function without 
consideration of (ftp parallel pn»eesses incurring in the 
oceans. The simpler test ol Ihe prognostic pinwess 
ot u model is to run it backwards in lime. When Uwe 
Radok and Dick lenssen developed Australia's lirsl 
numerical short-term weather forecasting model al 
Melbourne University m the lale 1950s. (Jcnsscn 
bJrO'l. they used (his simple and honest procedure, 
io the best of m\ knowledge, no modern long-term 
■climate prediction model has yet withstood Ihis .simple 
lesi ul veracity. 

in spite of the Fact ihut one stroke of a IS 
presidential pen can have greater consequences for ihe 
Austrnlian agricultural economy than even some of thr 
moie advehc predieiions olleaxl by the proponents ol 
climate models ciiTremly in vo^ue, I believe thai 
climate research is tinpmtunt I also believe that some 
of the recommendations ot those whose tailh in currcni 
model prcvlictiiins is clnse lo absolute, in which u 
radical reduction in the global scale of combustion is 
urged, deserve substantial endorsement, parlienlaily 
w here this may lead to adecellcration of the senseless 
de.siriic-t#onof the world's ti-ttesrs- Others may applaud 
the development ot niore vegetarian habits by the 
increasingly carnivorous buniiin race, whvnher or nor 
a reduction in global cattle numbers has any ultimate 
significant impact on climate uvnds. I do. however. 
find it rcgivllablc, thai the slroitger proponents ol 
climate research have disproportionately triggered a 
wide range o! political uiccluuiisius by means * ■! 
potentially alarming information, without proper 
\uhslantiuliou, apparently lo generate and maintain high 
levels of public attention. So far the main resuli appears 
to have been to gencraie public confusion between the 
concepts underlying the Greenhouse tffcvt and the 
Oztme Hole I well recall iwo remarkably different 
lectuio held within ihe space of $ single decade 
Id tlie very <ii)ie I'niveiMts ol Adelaide, by an 


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 tor 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 impacit 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. 


Brooks, C. E. P. (1949) "Climate through the ages: a stud) 

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-tiTi (1971) "Inadvertent climate modification (Report of the 
study of man's impact on climate)". (Massachusetts FnsL 
of Technology Press, Cambridge). 

Simpson. G. C, (1927) Mem. Row Meteowl. Soc. 2, 16. 

($28) Ibiii. 3, 21. 

(1929) Ibid, 3, 23, 


By Robert J. Allan* 


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. 


by RORFBT J. Al ! AV 


At t an K -I (W l M) Modelling climate t)h&ngp and v inability. Vmtf.v. ft $/£ .V. .<!«>/, I Hfl l). SH8, J] May, P$4 
Recent imeiesi Hi climatic chunks toto engenders considerable debate uhoui itic enhanced greenhouse effect 
;ttid its possible global impacts. Numerical computer models of the climate system inv important tools in the 
m iewific assessment of the enhanced greenhoase ellceL This paper briefly reviews the model development and 
approaches U*ffd t»t simulate the nature of anthropogenic changes and natural viu lability m the eh mute system. 
Research in Australia is given particulai emphasis 

KEV WORDS: climate- eham*e. numerical computer models 


lhe global climate system involves closely linked 
imctactions between the atn'osphere, oceans, 
c-ryosphcrc and the biosphere. This system is driven 
hv energy derived Irom sitfut radiation, with the nei 
incoming energy being balanced by that which is lost 
to space, The nonlmeai and highly dynamic '.'Innate 
system is fl consequence of the redistribution ol this 
enc^y by thermodynamics and the toiccs of motion 
dot ived from The planet's rotation Any lluciuatious or 
changes in ihe climate system can be brought about 
by external forcing, or by processes and readjustments 
intrinsic to such a closely coupled nonlinear .system. 
Since lite svstem is finely balanced, what might appear 
en he minor changes in any pari ol die system can cause 
large changes in its character. 

Until the latter part ol this ccntuiy. attempts to 
underhand the climate system, arid the variations and 
changes in jl, were pursued through observational 
studies and the development of physical theories ol the 
interactions and motions governing it. With the advent 
of, and improvements in, computer technology, ei forts 
to produce computer models of the climate system rcSVC 
expanded rapidly. Much initial modelling work locuscil 
on the ^^<1 to improve day lo day weather forecasts 
uMti£ models capable of resolving syn»ipii_- scale 
features in the atmosphere- However, the longer term 
envelope of weather events iluit constitutes the climate 
has now become a significant area in numerical 
modelling. Concern* about the enhanced grve-nhouM: 
effect and a need lo understand natural fluctuations in 
ihe elinUUc system have seen a strong locus on ihe 
development ol models able to capture Ihe processes 
governing interactions inherent in the climate system. 
Such interactions are shown schematically m r ig. 1. 

This paper piovides uh overview of the important 
characteristics of numerical models that have been used 
to improve Ihe scientific understanding of changes and 
fluctuations in the climate system. II focuses on both 

Climate Impact Group. CSIRO Division of Atmr»phenc 
Ke>ti3rch. PMB No. 1. Moalialioe 3195. Victoria." 

international efforts and the progress being made in 
Australia to examine potential climate problems using 
numerical computer models. 

L*m order models 

Some models have been developed lo improve 
understanding of particular features that dominate the 
climate over regions ot the globe and for various 
periods ol lime. Such models use what are often called 
"primitive" forms of the basic physical dynamics that 
are necessary to encompass the processes underlying 
climatic phenomena. These computer models contain 
sets of mathematical equations desenbiug. the most 
important physical interactions at each point on 
horizontal and vertical grids of points that cover the 
region in which the climatic feature occuis. All the 
equations governing the physical properties o( the 
phenomenon are solved at time steps indicative ol the 
observed nature of the processes heme- examined, lit 
this way. a model of the real system is built up. 
However, such models arc only as good as the 
mathematical resolution of the dynamics of the feature 
being modelled. Some real processes ate relatively 
easier to describe mathematically and Iheit mieiactitwis 
are known. Others are difficult to capture fully, or the 
present physical undemanding of Iheni may be limited. 
Jn general, low order models nave focused on lvsolving 
the important atmospheric and/or oceanic character of 
a particular climatic phenomenon. 

Perhaps the most coneeniryied emphasis in this area 
hasheen in the construction OffjOw order models aimed 
at capturing the essence vf bl Nino- Southern 
Oscillation (l:NSO) events. The FaNSO phenomenon 
is a large-scaJe orcan-atmosphere interaction that 
occurs irregularly and is centred in the Indo-Pacilic 
basin. It involves a close coupling of the important 
features o! the climate system across this region and 
has significant impacts in the marine and terrestrial 
environments. Some of the tnauJ variables and 
feedbacks known lo operaie during fcNS'O events arc 
shown schematically in Fig. 2. and a review of the 
phenomenon can be found in Allan r|OQ|). 




Changes of 
solar radiation 



HjA N^. °2. C0 2< °J elc • 




Atmosphere-land coupling v 

Atmosphere-ice coupling 


Wind slress 

Changes of 
atmospheric composition 

Changes ol land features, 

orography, vegetation, 

albedo, etc. 

Changes of ocean basin 
shape, salinity, elc. 







Musi low order models of ENSO have located on 
the Pacific Ocean region bounded by the suhtroptcs 
tn both hemispheres. Such models are very simple, 
with minima! physical representation of the amu.cvphere 
Mten tolfy w*> layers in the vertical) and a construction 
designed lo resolve ihe basic dynamics ofthe ENSO 
phenomenon alone. The building of models in this 
context has required substantial resources and 
particular recourse to observational studies and 
dynamical theories which provide a conceptual 
framework ofthe physical processes likely to underlie 
rhe phenomenon. At present, there is no firm theory 
describing all aspects of ENSO events, hut rather a 
dynamically consistent understanding of Seme of ihe 
important processes governing its behaviour. Current 
models of ENSO have been built under ihis constraint. 
More recenl developments in low order coupled ocean* 
atmosphere models have seen improvements m Ihe 
ability of ibcsciools ro predict ENSO behaviour The 
most comprehensive review of lire current \talas of 
ENSO modelling is Liven in Neclin ero/> (1942). Jn 
Australia . such work is perhaps besi characterised hv 
ihe low ordci model developments at the Bureau oi 
Meicoiolrvy Research Centre iBMRC)iKlecman BM, 
|99?5 Further progress can be expected ns ihe 
underlying dynamic* powering ENSO episodes 
become belter understood, and the capabilities of 
models to represent such complex processes develop 

General Circulation Modelling 

More detailed numerical models for climatic 
research are the General Circulation Models (GCMsi, 
which are three dimensional models of the global 
climate system. These models icpreseni interacuons 
between features of the climate system ihrough a series 
of mathematical equations on a spherical grid. The 
erid, representing a latitude by longitude array, is 
repeated in the vertical plane over the desired number 
of levels or layers ihrough Ihe depth of the atmosphere. 
This structure is shown in Pig 3, and provides the 
Iramework within which the model simulates Ihe 
dominant processes of the climate system over time. 
Physical processes and interacuons arc calculated at 
each grid point simultaneously in die ddisc model. 
In addition, in some models important oceanic links 
to the GCM atmosphere are mimicked by including 

a "slab ocean" in the model. This is a 50 m deep layei 
covering the Earth's oceans that performs like a well- 
mixed oceanic *near surface" layer. However, sea 
surface temperatures simulated by this appioach mnsi 
he continuously corrected to take account of the heat 
transport by horizontal currents and deep ocean 
processes. This is termed a Q-flu\ correction to the 
GCM. Using this technique and given the constraints 
mentioned, an evolving simulation of the Lorih's 
climate is produced by the model that captures numv 
observed aspects of the climate system Detailed 
specifications ot the CSIRO Division of Atmospheric 
Research (DAR) and the BMRC GCMs that have been 
developed in Australia, arc given in MrXlregur rr a/ 
(1993b) and Hart et ul. UOTTO. 

Despite the relatively sophisticated nature of GCMs. 
ihey are coarse in rheir spatial resolution with 
gndpoints spaced al around 500 km apart and vertical 
levels representing the atmospheric structure usually 
limited to between 10-20 layers As n result ot these 
spatial restrictions imposed by limilatiomof computing 
resources, many smaller features of the climate system 
such as tropical cyclones and deep cyclonic Nvsrcm* 
cannot be resolve^ by GCMs, Another important 
problem that stems from the spatial limitations is that 
some physical processes such as those relating lo 
aspects of cloud physics, cannot he fully resolved and 
arc parameter i!u:d in these ntcJclv These factors limit 
the current capabilities ot' GCMs to capture the full 
intricacies pf 'global clirnale At present, for example. 
GCMs arc unable to produce realistic ENSO events. 
Nevertheless, it must be realised that these are early 
generation models and that many of ihe large-scale 
aspects of the teal climate are seen in the current GCM 

An important advance in the development of GCMs 
is the coupling of atmospheric and oceanic modcK 
(formulated similarly to the GCMs) in oider to capture 
more ofthe essential elements responsible for observed 
climatic patterns. Currently, such coupled models 
require correction for fluxes of energy momentum and 
moisture exchange between "he atmosphere and ihe 
oceans if ihcy arc to resolve full ocean-aimo\phere 
interactions. Most simulations ot enhanced gieenhouse 
eftect conditions with current coupled ocean 
alTtioiphere GCMs have used flux correction. 
However, as noted in the section on low order models, 
coupled models provide the basis for a more realisiic 

pl( I TOP. A simple schematic view of UK major interactions in the global climate system. 

fig 2. BOTTOM. A schematic diagram of P.NSO interactions involving major oceanic .and atmospheric variables in ihe 
PiicifiL Hu.\in. The two response* (fiwi and slow) in the ocean rvfrr tn Jiriricm feedbacks reselling from Oic aatnru o( 
(lie dynamical waves generated by unpieal ocean-atmosphere interactions inherent in the phenomenon 



on the surface 

third dimension 

Fig 3. A schematic representation of a CjCM as it divides the earth into a regular three-dimensional grid. 









9 0.6 


0.2 - 


! t 




— <x 


X C 











i i 





Fig. 4. Pattern correlation for seasonal values ol model 
simulated and observed mean sea 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 

representation of the real world. Oceanic GCM.s 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 are slower than in the atmosphere. Ir 
must also be remembered that models of the 
atmosphere owe much to a long 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 due to sparse observational data coverage tor the 
oceans, 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 coupied model 
development at the Australian Bureau of Meteorology 
is given in Power et al. (1993). Similar development 
at CSfRO's Division of Atmospheric Research is in a 
rather more advanced state. 

In using GCMs for studying variations or changes 
in the global climate, one important prerequisite is thai 
the models can perform an acceptable simulation of 
current climatic elements. A number of 
intereomparisons between various GCM simulations 
of climatic fields and observed climatic parameters 



have now been performed \$f research gtoup^ around 
the world (Houghton e) a! [990, (992), Specific 
intcruimpnrisons For CCM simulations over Ihe 
Australian region arc detailed in Whcuon & Pmoek 
(1901. J993t and Whetton et at. (1994) (Fig. 4i. The 
must encouraging aspect ol these checks is Ihal ihey 
have objecuvety revealed a general improvement mi 
CiCM simulations a.s model resolution and physics have 
been refined 

The procedure employed to se* up a model run id 
priKluct ""oorilmT or present climate conditions usually 
involves an expei uncut wheie the CCM is run forward 
in lime under ihe current atmospheric CO : 
concentration with observed sea sutface temperatures 
and initial atmospheric input Holds. The aim ntrhis 
type of experiment is to obtain a sample of simulated 
climate lung enough for suilisticully valid analyse In 
ptaciice. thi>. normally requites between 10-30 year* 
of simulntcd climatic variables. The various llehk 
generated ean then be examined ufitg careful suuslicul 
tccfmin,ues that chccV the pattern of correlations 
between each of the GCM control output data 
distributions with the- appropriate observed field 
patterns. Simulated fields can also be examined tor grid 
pi*inl and total spatial field significance. If this prints 
to be satisfactory, then the GCM can be used to address 
the possible change to ebmule dial could resuh imm 
the enhanced greenhouse ef feel 

Two types of experiments arc commonly conducted 
to assess possible climatic change due to increasing 
C<Jj. The first, and most widely employed technique 
r*> ilffls in which the experiment is pcrlornied by simply 
miming the GCM with doubled CO : These 
equilibrium simulations require about a 10-year time 
penud foi the GCM to adjust to the now CO, limit 
alter it is instantaneously imposed, and then a vimple 
of ihe next 20-30 years of data is taken to rcpiesent 
(he enhanced greenhouse climate slaiisiics A doubling 
10 C<Ji is a standard experiment used as a benchmark 
against which all GCMs can be compared It does not 
relcr lo any particular dale in the future and implicitly 
Includes the radiative effect of increases in other 
greenhouse gases, which are expressed a* 
COyequivalents, The more physically realistic 
approach , which is being undertaken as coupled ivcan- 
atmosphere models become more w idely available und 
computing capabilities improve, is to perform a 
transient experiment. In this type ol" Simulation the 
CO, m the model atmosphere is gradually increased 
in a manner which loosely approximate* the projected 
increases in this gas, |o a lime when Ihe level has 

International evaluations of elinmre change research 
and GCM simulations and intercompansons a*e 
compiled in Ihe reports ol the Intergovernmental Panel 
on Climate Cfwttpe fTPCC) (Wriughliin n at i ut >C 
1992) In Australia GCM research with "in house" 

models is concentrated at BMRC and CSIRO Division 
of Atmospheric Research. Both groups have produced 
doubled CO. equilibrium runs- using unproved versions 
of their GCMs. and are developing GCMs lor transient 
experiments iTower c/ al. IW3) and to simulate UNSO 
and natural elunahe variability. .At the CSIRO Division 
nl' Atmospheric Research, results of enhanced 
greenhouse effect experiments have been used a* part 
Of me input into regional climate change scenarios for 
Australia, Important ojuuibuiionw 10 IPCC evaluations 
have been made by t'SIROs Division ol Atmospheric 
Research through studies examining the resolution of 
possible changes in rainfall amounts and intensities in 
doubled CO : simulations (Gordon ei al, !992>. This 
group also It4s reported on Ihe nature of variability 
within the climutz system simulated hv dtiuhled CO z 
experiments (Gordon & HflW 1994) Climate variability 
studies a1 CSIRO include ihn'-e using the recent 
Atmospheric Model Intercompanson Project (AMIP) 
re.\ult> lo highlight ifit impact ol chaoiic influence* on 
the atmosphere (Dix & Hum 1994) and an ongoing 
experiment simulating 500 years of climate 

Limited Area and nested modelling 

The need to rucus on likely changes in regional 
climates, together with the poor spatial resolution of 
GCMs, means dial mm lets ihal ean roolve smaller scale 
piocesses arc required Limited area models (LAMs) 
have finer spatial resolutions than the GCMs and can 
resolve leatuics of orography and geography that arc 
smoothed in coarser global models (Gtorgj 199(3; Giorgi 
rt a I I990< 1992) Typical horizontal grid resolutions 
thai have been achieved by LAMs in Australia aiC as 
low as 60 km wuh IX levels in the atmosphere 
(McGregor & Walsh 1993, McGregor fit ui 1903a). 
LAMs ait constructed to cover limited regions of Ihe 
Earth's surface and to tbcus on specific features of ihe 
climate system while beint computationally economical. 
Such models can be used to examine how a particular 
synoptic system, such as a tropical cyclone, may have 
behaved if it were Ifl have been uilUicnccd by diOcTeiit 
environmental o^ndirions (Evans 1993; Evans eral. in 
presst- For enhanced ereenliousc experiments this mxy 
involve increasing sco surlate lernpcraturei to mimic 
doubled CO : conditions and then testntg the sensilivit) 
of die tropical cydone and the amount ol rain ii pnxluces 
to such a change in boundary conditions. These nnxicls 
are particularly useful in assessing the sensitivity of hxral 
preeiptotion-producing systems to a changed 
environment. Joint icscaicli b> BMRC and DAR has 
concentrated on the simulation ol "east-coasf lows or 
what are orten termed 'cut-off" lows that can produce 
copious rainfall over southeastern Australia. This co- 
Operative study is examining the effect that increased 
sea surface temperatuics may have on (he pieeipiiatu-U 
n^jirne of synoptic* systems ( MvTnnes ti mL 1992) 


(E t At I AM 

• • 






'i * 
* * -l 

Lis, S Schematic shewing the nesting of the LAM grid within 
The CSIRO GCM urid over Ausiralui. 

An important new development using both GCMs 
and LA Ms is (he ion of ihe two in what is 
termed a "nested" model, The GCM generate.* the 
inlbntiaiion from a full global simulation ami provtdc* 
the boundary conditions tor the LAM that is embedded 
or nested within tile larger, coarser model. This is 
illustrated in Tig. 5. The nested modelling approach 
is capable of resolving features as small as 
60 x 60 to- Recent research at the CSIRO Division 
of Atmospheric Research has focused on the use oi 
this approach to produce finer scale regional resolution 
of climatic features and their sensitivity to changed 
environmental conditions. So far. LAMs have only 
been nested in equilibrium boundary conditions 
produced by GCMs with a simple Q-flux corrected 
ocean model, As such; these LAMs are limited in their 
peifoimance by the accuracy of the GCM boundary 

Other types of models are currently being used or 
developed internationally and in Australia. One 
interesting application is the use of tracer transport 
models to examine the atmospheric CO_, budget 
Studies at CSIRO DAR using such models arc 
described in tinting tt al, (1993), Such work is 
particularly important in helping to estimote'thc global 
carbon budget . 


Over the last five years, progress in developing 
numerical computer models of the climate system has 
been substantial, mainly as a result of the stimulus 
provided by concern about the enhanced gieenhouse 
effect and the need to investigate its possible global 
impacts, Today a suite of numerical models is being 
developed to lacMe various problems relating to the 
climate system. Specialised low order models focusing 
on particular features of the climate system are best 
exemplified by models ot the ENSO phenomenon. 
Such research has already been responsible for 
significant improvements in the forecasting uf ENSO 
phases. Larger atmospheric and oceanic GCMs arc 
used to simulate the global climate and likely changes 
and fluctuation*; in climatic patterns due to the 
enhanced greenhouse effect and minimi variability. 
Hf forts are also progressing with the full coupling ot 
the ocean-atmosphere system and the addition of 
realistic and interactive cryospheric and biospheric 
elements. Finer-scale resolution m the numciicaJ 
modelling of climate has been attempted using LAMs 
that have been nested in the coarser GCMs, or used 
as *'stand alone" models eupable ol resolving specific 
synoptic features and their sensitivity to potential 
changes m the elimafe 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 in 
improve observational data records and understanding 
of climate dynamics, so thai model results can be 
assessed in their proper context. 


] would like to thank Dr Chris Mitchell and Messrs 
Barrie Hunt and Kevin Hennessy, CSIRO Divison of 
Atmospheric Research and Dr Janette Lindcsuy. 
Department of Geography, Australian National 
University for discussions and then constructive 
comments on uhe initial manuscript of this paper. This 
work contributes to the CSIRO Climate Change 
Research Program and is part funded through 
Australia's National Greenhouse Research Program. 


Allan. R. J. (1991) Australasia pp. 73-120. In GlanU. M.. 
KM/, R. & Nicholls, N, (Eds) "ENSO Teleconnectjon* 
Liriltfog Worldwide Climate Anomalies Scientific Basis 
and Societal Impacts" iCawhridge Llniveisiiy Press. 
Cambridge. UK). 

Knttno, T Ci_, TRroiKcrft, C M , FRASjerv, R ,1 A 
GKANtK. H. (1993) Synthesis inversion of atmospheric 
CDa using ihc GLSS tracer transport model. CSIRO 
Diu.uon <>/ s\mto\phttH Hescanh, lhh> t\ip, No. 2ft 
Melbourne 1-44 



E vans, J. L, (1993) Sensitivity of tropical cyclone intensity 
to sea surface temperature. J- Climate 6, 1133-1140 

_. Ryan-, B. R &. McGregor. J. L. (In press) A 

numerical exploration of the sensitivity of tR'pical cyclone 
rainfall intensity to sea surface temperature. Ibid. 

Giorgl F. (1990) Simulation of regional climate using a 
limited area model nested in a general circulation model 
Ibid. 3, 941 963. 

, Makinucci, M. R, & VlSCONT!, G. 0990) Use of 

a limited area model nested in a general circulation model 
for regional climate simulation iwer Europe- J. Geophvs, 
fife 95, 18413-18431. 

„ _. &c (1992). A 2 x CO : climate 

change scenario over Europe generated using a limited area 
model nested in a general circulation model 2: Climate 
change scenario. Ibid. 97, 100U-10028. 

Gordon, H. B. & Hum, 8, G. (1994) Climate variability 
Within an enhanced Greenhouse simulation. Climate Dm. 
9. 195-212. 

. Whetton, P, H , Pmxx'K, A. B. v Fowler. A. M, 
& Hayi.ucK- M. R. U992) Simulated changes in daily 
rainfall intensity due to enhanced greenhouse effect: 
Implications for extreme rainfall events. Ibid. 8. 83-102. 

Hart, T. L., Bovrkh, "W., McAvaney. B. J., Forgan, 

B. W. & McGregor, I L. (1990) Atmospheric general 

circulation simulations with the BRMC global spectral 

model: The impact of revised physical paramctcrisations 

J. Climate J. 436-459. 

Houghton, J. T.. Jenkins, G. J. & Kphraums, J. J. (1990) 
"Climate Change: The IPCC Scientific Assessment" 
(Cambridge University Press, Cambridge). 

. __, CALLANDhK. B. A. & Varnfy. S. K (1992) 

"Climate Change 1992' The Supplementary Report to the 
IPCC Scientific Assessment" (Cambridge University Press, 

KubbMAN, R. (1991) A simple model of the atmospheric 
response lit F.NSO sea surface temperature anomalies. / 
Azmos. Set. 48, 3-18. 

(1993) On the dependence of hindcast skill on ocean 

thermodynamics in a coupled ccean-atmosphcre model. 
./. Climate 6 t 2012-2033. 

McGregor. J. L. & Walsh, K. (1993) Nested simulations 
of perpetual January climate over the Australian reeinn. 
I Qeoplm. Res 98. 23283-23290. 

_ Walsh, K. & Katzfev. J. J. (1993a) Nested 

modelling for regional climate studies, la pp. 367-386 
Jakeman, A. I. Beck. M. B. & McAleer, M. J. (fids) 
"Modelling Change m Environmental Systems"* (John Wiley 
& Sons LLd.. Chichester). 

Gordon, H. B.. Watterson. L G., Di\\ M. R & 

Rotstayn. L. EX (1993b) The CSIR0 9-I<evel atmospheric 
general circulation model CSIRO Division of Atmospheric 
Research t Tech. Pap. No. 26. 1-89. 

McInnls. K. L., Leslie. L. M. &. McBriuh, I I. (1992) 
Numerical simulation of cut-off lows on the Australian east 
coast: Sensitivitv u> sea-surface temperatures. Inx. V. 
Oimatoi 12, 783-795. 

Neelin. J. D,, Lath*. M.. Aii-AAR'i, M. A. P.. Cane. M. 

A., CUBASCH, V., GaTE.S. W. L., GbNI, P. R, s Ghii , M.. 

Gordon, C\, Lac, \-C. Mlchoso, C. R.. Mfthi., G, 
A.. Ohkruluek, J. M,. Philander, S. G. H., Scbopl. 
P. S. t Sperber. K. R , Steri , A-j Tokiok^, T., Tkibbia, 
J. & Ze&iak* S. H. (1992) Tropical air-sea interaction in 
general circulation models. Clim: Dyn. 7, 73-104. 

Power., S. B.. Colman, R. A.. McAvvNtv, B, J,. Dahni, 
R R.. Moore, A. M & Smith. N. R. (1993) The BMRC 
coupled atinosphere/ocean/sea-ice Model. BMRC Res Rep 
No. 37 (Bureau of Meteorology, Melbourne). 

Whetton, P. K. & Pittock, A. B. (1991) Australian region 
intercompanson of the results of some general circulation 
models used in enhanced greenhouse experiments. CSIRO 
Division of Atmospheric Research t 'lech Pap. A'o. 21, L73. 

& (1993) The growing consensus in simulated 

regional climate for enhanced greenhouse conditions: Is 
there a palaeoclimatic analogue? Quaternars Australasia, 
11. 1. 81-87. 

, Ravner, P. L. Pjtiock. A. B. & Haylock. M R. 

(1994) An assessment of possible climate change in the 
Australian region based on an intercompanson of general 
circulation modelling results. ./ Climate. 7. 441-463. 



ByM. A. /. Williams* 


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. 

Ihwwiitm of the Row! Stuictv oj S. jfrv; .I<KU|. llSill IWH 


by M A. J. Williams* 


Williams. M, A. J, (1994) Some implications of past climatic changes in Australia, Traps, /?. 5oc. A. Mst, U81O, 
17-25, 31 May; 1994 

An important question,, in the context ol possible human impact on global climaTt\ is whether past climatic 
history can oiler insights into possible fuiim? climatic change, fcquallv critical is the likely response of the physical 
environment to any future climatic changes, [{violence trom pollen analysis, lake fluctuations, desert dunes and 
coastal plains from Holoeene deposits in Queensland. Victoria, New South Wales and Northern Territory, 
demonstrates that the response of different elements ol the Australian landscape to geologically-recent changes 
in temperature and precipitation was often timc-transgressivc. Any attempt lo use palacocnviromncnial data lo 
predict possible future change must therefore take due account of the varying response times of different constitueiiLN 
til the Australian landscape to external disturbance. A synchronous response to climatic change is more likely 
with relatively simple biophysical systems such as small closed lake basins pi somcc-hordciing dunes than with 
more complex systems such a> tropical rainforests and tropical coastal plains. 

kLV WGKDS Climatic change, Aiuimlian landscape, past climates. Queensland rainforest. Victorian lakes, 
Sew South Wales Junes. Northern Territory coasta. plains, response limes. 


There is increasing concern thai the accelerating 
impact of human activities upon our natural resources 
of land, air. water, plants and animate is seriously 
damaging many of our more fragile ecosystems, 
culminating m irreversible losses of genetic as well as 
cultural diversity. Human action* may also he 
contributing to possible changes in world climate 
through a combination of burning of fossil fuels and 
changing land use (most noiably deforestation), both 
ol which reduced the terrestrial store of carbon by about 
6.0 ± 0.5 and 1.6 1 1.0 PgC in 1990, respectively 
{Houghlon ct ai 1992). 

Although the increase tn the atmospheric 
concentration of carbon dioxide, methane, nitrous 
oxide and other "greenhouse" gases since the start of 
the industrial revolution is well documented, the likely 
impact upon global climate and ecosystems remains 
unclear (Pcannan 1988; Houghlon ei at. 1990. 
Dunnette & O Bnen 1992). Notwithstanding the 
unavoidable scientific uncertainties over the magnitude 
and frequency of nature climatic fluctuations, there 
appears to be a significant measure of agreement that 
the increase in anthropogenic aerosols and trace gases 
will enhance the greenhouse effect, culminating in 
global warming of the lower atmosphere, particularly 
in middle to high latitudes (Houghton ft al 19^0; 
1992). However. There is far less agreement on how 
the global pattern of precipitation might respond lo 
enhanced greenhouse warming, prompting a number 
oi researchers to look to tile past as a source Of possible 

MawTM'in {.traduate Centre foi KnvironnicHial Studies. 
L'niversity of Adelaide. South Australia 500?. 

analogues for future global warming (De Dcckker et 
al 1988; Peiii-Mairetfw/. I99J; Street-Permit 1994) 
The aim of this paper is to consider some of the ways 
in which a study of geologically recent changes in the 
Australian environment can offer insights into how the 
various elements of our natural environment are likely 
lo respond to future climatic change. 

Past climatic rh*tng«s 

Will {-/(marie hixtorx ttptav irself 

The seaich tor past clirnafic analogues of a warmer 
future planet earth is based on a number of explicit 
and implicit assumptions. A major bul unproven 
assumption is the notion of cyclic change. *as iti the 
past, so, fail, in the future". Linked to this assumption 
of recurrent climatic changes is Lhe equally unproven 
assumption of similar recurrent boundary conditions. 
Boundary conditions in (his context means the global 
distribution of land, sea and ice; the global distribution 
of major vegetation /ones; and (he global albedo 
pattern. Insofar as global fluctuations in suifaec 
insolation and m ice volume arc linked to cyclical 
changes in lhe orbital geometry ol the earth at timc- 
scalet. of l()M0 ? years, certain climatic changes arc 
indeed cyclic, at least when set against (he last two 
million years of lhe Quaternary period (Williams ft 
at. 1993). But what of time-scales of shorter duration, 
more obviously relevant to present human pre- 
occupations? Here it is important lo distinguish 
beiwccn climatic change and climatic variability. 

Climatic change and climatic variability 

Much confusion over climatic variability and 
climatic change stems from a failure to specify the time 



scale involved. Fig. 1 illustrates climaxic variability a( 
different time scales during the last 0.9 million years 
(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. riiis 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. 
It 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 

® Utile lea Age 
® Last Glacial 
©Last Interglacial 

pig, L Climatic variability at different scales in time during 
the last 0.9 million years (from Williams etal. I993, adapted 
from Australian Academy of Science 1976). 

temperatures between about 1940 and 1960 (Fig. la). 
None of these global temperature changes seem to be 
linked in any obvious way to the steady exponential 
increase in the concentration of atmospheric carbon 
dioxide - a point conveniently forgotten by over 
enthusiastic advocates of enhanced greenhouse 

Vie reconstruction of past climatic events 

The reconstruction of past climatic events is based 
on many independent sources of proxy data^ These 
"natural 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 arc discussed 
in derail by Bradley (1985, 1990) and by Williams et 
al (1993)." 

Limitations of palaeodinuxtic enquiry 

Certain elementary precautions are essential when 
using any particular type of evidence to reconstruct 
past environments and climates. Each may be useful 
for a particular purpose and for a particular spatial and 
temporal time-scale. However, as Williams etai (1993) 
note: "Difficulties anse immediately when we use- 
Procrustean tactics to force the data to yield 
palaeoenvironmcntal information at particular scales 
in space or time tor 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" (op tit., 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 palacaclimatie reconstruction. (After Bradley 1990, Table L) 
* Minimum sampling interval in most cases- T * temperature: H — humidity or precipitation; C = chemical campus it inn 
nj air (CJ, Miter (C H ) or soil (Cj: B = hiomass and vegetation patterns: V = volcanic eruptions: M ~ penrnaBnetic field 
variations; L - sea lewis: S — soteir activity. 

Information derived 

T. H, B, V, M, L, S 
T.H C a , B, V, M. S 
T, H, C w , B, V, M 
T, H f C a , B> V, M, S 
T. H. B 
H, B. M 
T, C w , B, M 

T. H r C 5 , V 
T. H. V, L 
H- EL V. M. L 

Best Temporal 




Range (yr) 

Historical records 



Tree rings 



Lake sediments 

tr to 20 yr 

K> J -10 ft 

lee corev 


10 s 


100 vr 



100 yr 


Ocean «.<-»res 

1000 yr 

10 7 



to 4 


100 yr 

to 5 



100 yr 

10 7 




1Q 1 



oi a natural archive usually provides direct information 
about some component of the natural environment. 
buch as a river, lake or dune. To inter climate (or some 
particular attribute of climate, such as temperature or 
precipilation) from a particular element of ihe 
landscape is a far more circuitous and difficult 
procedure, with tar greater scope for interpretative 
Consider, for instance a lake. The outstanding work 
by Gasse, Fontes. Street-Perrott and co-workers on 
reconstructing the climatic history oflakes in Africa 
has demonstrated the need for calibration using preseni- 
day chemical, physical and biological data, as well as 
the need for taking due account of local hydrologies! 
factors arid of extreme events when deducing 
palncohydrology from palaeolimnology (Fig. 2) (Street- 
Perroti '& Roberts 1983; Fontes et at. 1985; Gassed 
«/. 1987; Sueet-Perrott 1991). Only then is it possible 
to attempt a reconstruction of palaeoclimate. From 
reconstruction of local palacoclimates to regional or 
even global palaeoclimalic modelling is yet another step 
removed from the original field data. Provided that the 
limitations of palaeoclimatic enquiry are clearly 
recognised, and the appropriate steps are followed in 
interpreting past climate from proxy data, useful 
insights are possible when using past environmental 

Font environmental analogues 

A decade has elapsed since Piuoek & Salinger (1982, 
1983) suggested that the early Holocene climate of 
Australia towards 9000 - 7000 years ago may be a 
suitable analogue for the continent in a C0 2 -wa;med 
earth Granted that many early Holocene inke and 



(lakos and dvfin;) 

calibration using 
modem toteroticvs 


(wr»H?'. sails an<3 solutes and stable 
li=otopo oataeotudrjtsts) 

^ffxvts ot local 




(eslimatton or palaso-prsrlpiiailoh, 
-uvaporoliin, -tornperature, etc ) 


Fig. 2. lake and river fluctuations as plaeoclimatic indicators 
(Source; K Gasse 1980, unpublished seminar paper to 
IX-|witnicnl of Geography, Mouash Umvci'NJTV), 

poUen sites in Australia and Papua New Guinea do 
seem to indicate a warmer and/or wetter climate at this 
lime {Williams 1984), just how reliable is the evidence'/ 
Can it be quantified? Is ihe "Climatic Optimum" or 
"Hypsithermal 5 ' 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 attempl to answer some of these questions, it 
is appropriate to consider ihe evidence from a variety 
of different Holocene sites across Australia. In the 
ensuing discussion we consider four distinct types ot 
Holocene site in four widely separated localities: the 
rainforests of northeast Queensland, "maar" lakes ol 
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) palynolo&ieal studies of the late 

Pleistocene and Holocene vegetation history at four 

pollen bearing lake or swamp sites in the Alherton 

Tableland of northeast Queensland have revealed 

that the period of maximum rainforest expansion 

lasted from 6000-8000 (o 3CO0 years B.P. (Fig. 3). 


Quincan I Biomtteld 
Crater J, Swamp 

Lynch 's 

A ' 


n 5 


2 7 







Time ot arrival 
of rainforest 

Peiiofl ol 



1400 1600 1600 2000 2200 2400 2600 

Present mean annual raintall (mm) 
hg. 3 VcgctaUon changes in northeast Queensland deduced 
from pollen analysis of Holocene naler lake sediments. 
(From De Deckker cr at, 1988, adapted from Kershaw 



However, as Figure 3 clearly shows, the response of 
the rainforest to increasing warnuh and increasing 
effective precipitation was strongly time-transgressive. 
Rainforest appears in the Bromfield Swamp pollen 
recrird towards 8500 years BR, at Lake Quincan 
Crater not until after 70(H) years B.P. The demise of 
the rainforest at all ibur sites was roughly synchronous 
(3000 years B.P.), indicating that the rainforest 
responded slowly to climatic amelioration (wanner and 
wetter conditions) but rapidly to climatic deterioration 
(cooler and drier conditions). 

Application of the CSIRO hioelimalte prediction 
system developed by H A , Nix to Kershaw's 
Queensland rainforest pollen data enabled Kershaw & 
Nix (1989) to derive more quantitative estimates of the 
Holocene climates. Their analysis revealed that 
Holocene temperature maxima were not achieved in 
the Atherlon Tableland until 5000 years B.R, with 
mean annual temperatures up to 3.5° higher than today 
persisting until about 3500 years B.R Mean annual 
precipitation at that time appears to have been at least 
300 mm higher than today, and most probably 
500-800 mm higher (Kershaw & Nix N89; De 
Dcckker<7 al. 1988). 

Onr obvious conclusion to be drawn from the 

Queensland pollen record is that the interval 7000-9000 
years B.R was not the lime of "Climatic Optimum" loi 
northeast Queensland. The wettest and warmest period 
came considerably later, and on present evidence seems 
to have lasted from 5000 to 3500 years B.R If this is 
true tor the tropica) northeast of Australia with its 
monsoonal summer rainfall regime is it true also for 
the temperate southeast of the continent, where the 
rainfall comes mainly from the westerly airmasse.< 
which pass across the southern margins of the continent 
most persistently in winter, when the Antarctic 
convergence is in its most northerly position? 

The Holocene crater lakes of western Victoria 

Lake Kcilambetc in Victoria ts perhaps the bcsl 
studied Holocene lake in Australia (Dodson 1974; 
Bowler W8I; De Deckker 1982, Chivas 'let al 198.V 
1986), lake other volcanic explosion-crater or ""maar* 
lakes in Victoria, it occupies a small closed basin and 
is highly sensitive to changes in precipitation and 
evaporation over its catchment. Five "maar" lakes in 
western Victoria . including Lake Keilambete. have 
yielded useful information about Holocene changes in 
the balance between precipitation and evaporation (D.-* 

P/E Estimates for 5 Victorian Maar takes for the Holocene 
Keilambete Gnotuk Bullenmerri West Basin East Basin 













Medium and 
becoming low 

Low - Medium 

Low - Medium 








Medium ? 

Low and 






A —, 



Medium - Low 

Low - Medium 



< — 













High - Medfurf* 




Medium » LOW 










1U """ 

< 50 km 

Fig. 4. Changes, in Holocene preeipitalioil/evaporulion (P/£) ratios tor five volcanic "maar" lakes in western Victoria. <Fror 
De Deckker ei ai 1988 » 



Dcckkcr 1982; Oe Deckker M al. 1988)- Highest 
precipitation/ evaporation (P/E) ratios, peak laVe levels 
mid lake salinity minima were all within (he interval 
7000 lo 5000 years B. P From 5000 to 3000 B P (when 
northeast Queensland w;vs warmer and welter than 
today; lake levels were lower and salinity values were 
higher than in the preceding 2000 years or so (Fig. 4). 
One possible inference is that winter rainfall may 
have been lower towarvM. 5000-3000 years B.I 1 . Equally 
plausibly, lolal annual ramlall (including both summer 
and waiter precipitation! may have been reduced al lha! 
time. We cannot, as yel, ehoose between these two 
possibilities. What docs appear certain is that the lime 
of greatest effective precipitation (7000-5000 years 
B.P.) was several thousand years earlier than the time 
nf maximum eflcctive precipitation in northeast 
Queensland. It is tempting to speculate that we are 
seeing the effects ot two distinctive climatic systems: 
one tin northeast Queensland) controlled by the tropical 
summer monsoon, the other (in western Vicloua) 
controlled by the winter westerlies. If so. the response 
of both systems 10 postglacial warming was not 
sy til hrunous along the easiern third of Australia. Nor 
may it be synchronous m the race ot any future global 
wunmng. We tuio now to the semi-arid inland areas 
of New South Wales 

The Holocent somre-bornVrin*: dunes of 
central western New South Wales 

Holocene palaeiKiunauc data are exceedingly scarve 
for the semi-arid and arid areas of Austialia which 
together compi ise 75% of our present land area The 
late Pleistocene fluctuations in the Willandra t.ake\ 
region or western New South Wales so carefully dated 
and elucidated by Bowler (I970 1 , W83) do not yield 
a Holocene signal and in any event have far more to 
do with runoff from the Eastern Highlands via the 
Pleistocene l.acnlan-Willandra river system than with 
local changes in rainfall and evaporation (Williams ci 
al, 1986). 

One climatically-sensitive area capable ol providing 
useful informal ion aboul 1<huI hydrologieal events is 
the desert margin system of central western New South 
Wales studied by Watson <t975 2 . 1976) and by 
Williams vt ul. (1991). The study area occupies about 
SO 000 krrr in the semi-arid region bounded by the 
Darling River lo the north and wesl, and the Willandra 

'Bowler. J M (L*?70) l-ate Quaternary environments: a 
study of lak*?s diid associated sediments in boutheostern 
Australia. Ph.D. thesis, Australian National University, 

^Wasson. R. J. (1975; Evolution ot alluvial tans in two ateas 
of .southeastern Australia Ph.D. Ilicsi*. Mm-quaiie 
1 tnivwMiy, UnpubL 

Creek distributary of the Lachlan River to the south 
(Fig. 5). There are no perennial rivers in lite entire- 
area. The ephemeral stream channel known locally as 
Crowl Creek of Sandy Creek (Fig. 5) Mows 
intermittently during very wet years and Ihe West to 
east trending linear dunes aie tptltfj vegetated and 

Fig. 5. Map of central western Ne* South Wales showing 
location of Sandy Creek. The shaded areas are i>oln(e«i 
Tallies over 200 in in elevation (Source: Williams el al. 

The Holocene climatic history' of thi* vast area is 
poorly understood, but ihc pattern of local 
environmental chaises is now reasonably welt 
documented (Figs 0, 7V Soutce-bordcring dunes were 
actively forming from channel sands ferried in by 
Sandy Creek between about 5500 to roughly 600 years 
B.P. Both before and after thai interval Ihe dunes were 
inactive, vegetated and stable (Williams et al. 1991) 
Source-bordering dunes are dunes which develop 
immediately downwind ot a patent source of sand, such 
as the sandy bed of a river, a sandy lake beach or a 
sandy alluvial tan. There ate three prerequisites tor 
the formation ot source-bordering dunes: 
(I) A regular replenishment of the sand supply (lor 

instance, from a seasonally-flowing sand-bed 

t2) Strong unidirectional winds for at least part of the 

year; and 
(3) A sparse or limited vegetation cover adjacent to 

the sand source. 

Any interpretation of the Holocene climatic histoty 

ol this region must take into account the requirements 

for source-bordering, dune formation, the apparent 

absence of early lo middle I lolocene dune deposits in 




C Si fs cs a 

§12A: Don's Gully 

h?£?o\%X horizontal 

^Oqo5) bedding 

? Top ot calcic unit ? 

§7: Dillon Valley 


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 
Fit;. 5). The block diaeram shows Sandv Creek, the source-bordering dunes and the wind direction. (Source: Williams 
el al, I991). 

Years Before Present 


Aftuvta] and aeciian satafc 

Pedogenic cartiona'j? 

Alluvial and aeoiian sands 

pDdrjgenic carbonate (6) 

Alluvial one Sooliar ?arid^ 

Aboriginal f"res psi 

Higli lake levels tgi 
Calcareous duot 

Gypseous lurelte (i 
Calcareous etust 

Fig. 7. Generalised late Quaternary piedmont stratigraphy in 
the centraJ western New South Wales study area. The 
numbers in brackets denote numbers of samples dated, and 
includes both radiocarbon and thermoluminescence dates 
(Source: Williams et al. I99L, revised to show additional 
dates. ) 


this area (FJg. 7). and the fact that the present-day 
dunes are vegetated and stable. A tentative 
palaeochmatic interpretation, based mainly upon the 
geomorphic evidenec and needing to be tested by future 
independent work, is as follows: 

• 10,000-5500 yeans BP: 

Rainfall more uniform 

Wind velocities low 

Surface well vegetated 

Erosion and deposition very slow 

• 5500-600 years BP: 

Higher winter rainfall 
Seasonal channel How 
Summers very hot, dry and windy 
Summer deflation 

• 600 years B.P to present: 

Hot summers, cold winters 

Rainfall more uniform 

Wind velocities low and/or 

Denser vegetation 
If ihis interpretation is correct, any change to a welter 
climate or to a more seasonal distribution of rainfall, 
with more runoff during the winter months, could lead 
to a renewed phase of source-bordering dune formation 
provided the summers remained dry and windy and 
the riparian vegetation relatively sparse. Should the 
future climate in this area become both warmer and 
wetter, the somewhat paradoxical outcome could be 
a replenished sand supply and reactivation of the linear 
source-bordering dunes. 

Arnhem Land Plateau 
(Middle Prolerozoic 
Sandstones & Dolerites) 

Sandstone Outlier 

Wooded Lowlands 

Paperbark Swamps 

Shelly Sand Ridge 
(Holocene Cheniers) 


Uranium bearing 
Lower Proterozoic 

Strike Ridge 

Tidal River 

Coastal Plains 

Late Catnozoic 
Sands & Gravels 

Late Quaternary 
Marine , Estuarine 
& Freshwater Clays 

Fig. 8. Block diagram showing die Arnhem Land plateau of Northern Territory and the geologically youthful coastal plains 
to the nonh. (Source: Williams 1991.) 



Tht* HuliK-Ttie constat plain* of the 
MligatM* Rivers ana, Northern lerrilor) 

The Iroptciil Ofttiftl plains or Northern Territory 
stand in almost diametric contrast to the ephemeral 
streams and desert dimes ol senu-nrid western New 
South Wales. Their significance stems tnnn the lati 
ihat the HoJoeene coastal plains which extend loi some 
thousands ol kilometres from the far northwest ot* 
Western Australia io the tar northeast ol Queensland 
are very dynamic landforms, and highly sensitive: to 
even minor changes in sea level, sediment ti\id. salinity 
and wave climate (Williams 1991). They arc also host 
to a unique and abundant fauna and Mora (Haynes et 
at, 1991 >, In addition, the coastal plains west of Darw in 
abut onto the Arnheni Land plateau (Tig 8). wilh its 
abundant galleries of Aboriginal rock art. and its record 
nl over 50 000 years of prehistoric Aboriginal 
settlement (Roberts et at. 1990), 

With I he tnclline of" I he laic Pleistocene ire sheets 
of North America and Europe, world sea levels lose 
from their last glacial maximum level of aboul -135 m 
(at 18 00U years B.Pl to their present levels lOwanK 
MKMI-71KX) "years R.P. (Williams et uL (90& 

The initial rise was rapid, and vast areas of the 
continental shell oil northern Australia were 
submerged at a rote of over 20 metres a year or roughly 
40 cni a week. Once this rise had stowed down, which 
it did in the last few thousand years, coastal mangroves 
heean to colonise the micrtidal muds_ allowing muddy 
■sediment* to accumulate on rhc old late Pleistocene 
land surface inland of the coastal mangrove fringe. I h>s 
process accelerated once the sea attained ty presem 
level, and widespread mangrove swutups developed 
<ktos,x (he present area of the fOfetfll plains to mid- 

llolocene times, from about 5000 to 2000 years B.P. 
(Woodtoile et al 1985). These mangrove swamps 
proved to be highly efficient sediment traps, and wett 
eventually buried by estuarine and. ultimately, by 
Ireshwater muds (Woodtoile vt at, 1986; Williams 1991; 
Wasson 1992) The present-day coastal plains arc thus 
a relatively youthful rcaiure, and arc gene rail v less Ihar 
2000 years old. 

How these plains might respond to sea level change* 
will depend upon a variety oi factors, including lire 
rate of sea level rise; the impact of any climatic change 
upon runoff and sediment yield in the litlal rivers; the 
magnitude and frequency of future cyclones, and the 
relative duration of the wet and dry seasons. Since none 
of these variables is accurately ptedietable, speculation 
seems unwarranted heyond noting that the coastal 
plains arc likely to remain a geomorphically dynamic 
and aclocly developing feature, of (he landscape, jus! 
as in the past 5000 years. 


Provided certain common-sense pieeautions are 
observed, an appreciation of Holoccne climatic and 
other environmental changes in Australia can be a 
useful guide to possible future landscape responses Io 
glotal wanning whatever its ultimate causes. The 
Australian landscape is a palimpsest of landforms, soils 
and plant associations, all of which will respond to 
lulure climatic change in a variety of ways. The way 
in which our rainforests, lakes, dunes and coastal plains 
have responded to Holocene climatic changes often 
us some guide to their possible future responses, It is 
highly unlikely that these will be either simple or 


AlSTKMJAK ACADEMY Ol 1 SCSI NCI (1976) Kcpotl of :| 
Committee on Climate diuime" (Australian A. udemy pit 
Science. Canberra). 

BoWLfcft, t- M. (t^SIi Au*tiuliun sail l.iku*: 1.1 
prflactihydmhtgieul approach Hyilmiholngin 82, 411-44, 

_____ (19K3) Lunettes as indite* ol livdmlngh'ttl change: 
A review of Australian evidence, hot, R. .W. \'m. 95. 
147 hN 

BRADirv, R «N. (IWt-Quaternurv PaJeiK-lnitatology" (Allen 
&. Umvm. London). 

iLd.) tJyyt.n "Global Changes ut the Past "" (| CAR 


1,1986) Magnesium content ol 

OMt<e tor Fnlerjisciplinary Earth Studies, Roukter. 

CliAi't'LLL. J. M. A, Sl Ghindkod. A. (1983) "Pn»Leedings 
of Ihe Kirst ("MM AN/ Conference. Howmans<uft l&B. 
VmIh ncs I and 2? (Dopl ot Biotieography & Gcotnor- 
(ilioloay, Australian National University. Canberra! 

Chtva.s A R..Df Dpckki-r P&Snnirv.r M C. [WAS) 
Strontium content <.>( oslracods indicates lacutlrinc 
paJaensalimiv, /Vuwrv 31ft. 25I-2&. 

non marine o.stracod shells: u new palemalinmnerer and 
palueotheimomete-i. I'alttvowgr. y'aluevi hr/mtul. 
PatatWt'f. 54. 43 hi. 

Dh DECKKfcK. P. (J982) Holocene ostratods. other 
invertebrates and fish remains from core* or tout moor lfl1«H 
in southeusteni Austral ia, ^c', R. ,V'c, Via V4. 193-2211 

, Kfrskaw, A, P & Wti iiaMS, M. A. I (8*881 Pa^ 

environmental analogues, pp. 473-488. /// CJ. I. Pearman 
(kd.) "(jiwnhousc: Plfttimny, tor etimaie change" (H. .1 
Brill. Leiden & CSIRO. Melbourne). 

0« ios< >N„ S, H974) Vegetation and climatic history near Lake 
Keilamhete, western Victoria. Auxt. ./, Hot. 22, 704-7t7 

DnNMi-l ih, D. A. St O'Bkii'n. K. J, (hds) (1992) "The 
Science ot Global Change: The Impact ol Human AeTTVftiea 
on the eitvironnifnc" (Anienean Chemical Society. 

FosTrs, J-C Gasm . K, Caiioi. Y.. Pi.A/tA.1. J.-C. 
Cahjiom=u P. DiMht'fLi'. P. A. & Kac/makska. I. 
1,1985) I resrrvvarer and nurme-hke environiDcnifi trom 
I mI-kltil L.r.i -■ in Nonhern Sahara. N<nun- 317, r>08-6lo 



Kac/marsK-\, 1.. Dp, Deckker, P.. Soelie-Maksche. I,, 
Caelot. Y & Puptojple, P- A (1987) Biological remains 
geochemistry and stable isotopes for the reconstruction of 
environmental and hydrological changes in ihe Hotocene 
lakes from North Sahara. Palaeovcogr. Palacr.chmatot. . 
Palaeoecol. 60, M6. 

Ha^Es. C. D, Ridpath. M. G. & Wni.iAMS, M. A. J. 
(EUs) (19911 " Momoonal Australia." (Batkema. Rotterdam) 

Houghton, J- T., Callander, li A. & Varne'i, S. ft. 
(Eds) (1992) '■Climate Change 1992. The Supplementary 
Report to the 1PCC Scientific Assessment " (Cambridge 
University Press. Cambridge). 

Jenkins, G, J. & Ephraumv J. J. (Eds) (1990) 

"Climate Change, The 1PCC Scientific Assessment'. 
(Cambridge University Press. Cambridge). 

Kershaw. A, P- (19S.i) The Vegetation record from 
northeastern Australia; 7^2 KA.'pp 100-101, Vol I and 
I-ia 67. Vol.2. In Chappell J. M, A & Grindrod. J. (Eds) 
"'Proceedings of the Hirst CLiMAN/ Conference. 
Howmuns Gap. 1981 . Volumes 1 and 2," (Depu ol 
Biogeography & Geomoi phology. Australian National 
University, Canberra). 

& Nix, H A 1 1989 > The use of bioelimati: 

envelopes for estimation of quantitative palaeoclunati. 
values. Proc. CUMANZ 3 Swift. . 78-85. (CSIRO. Division 
of Water Resources, Canberra) 

Petit Marie, N . KoNTUiNr, M. & Row am>. C. (I99U 
Atmospheric methane ratio and environmental changes in 
the Sahara and Sahel during the lust 130 Kyrs. J\ilaco\>foy,r. 
Palufoetimatol. Paheoeaif. 86. 197-204. 

Pittock, A. B. & Salinger, M J <l*>82) Toward regional 
scenarios for a CO^-wanned earth. Climate Change 4, 

. & (19H3) The climatic optimum and a 

CO^ warmed earth: the Australasian Region, pp. 122-125. 
In Chapped J. M. A & Grindrod, J. (Eds) "Proceedings 
of the First CL1MANZ Conference. How mans Gap, 19SE 
Volumes 1 and 2 ** (Dept. of Biogeography & 
Geomoiphologv. Australian National Universitv. Canberra) 

Roberts. R. G.. Jones, R. & Smith. M' A. (1990) 
Thermoluminescents dating of a M> 000-year-old human 
occupation site tn northern Australia. Natun 345. 153-150, 
Street -Perrott, F. A. (1991) General circulation tGCM) 

modelling of palaeoclimate.s: a critique. Vie Hol^eem* 1. 

fl994> Palaeo-perspcctives; Changes in terrestrial 

ecosystems. Amhio 23(1). 37-43. 

& Roberts, K, (1983) FI actuations in elused-basin 

lakes as an indicator of past atmospheric circulation models, 
pp. 331-345. fa Street-Perron, F A.. Beran, M. & Katelifle. 
R. (Kds) "Variations in the Global Water Budget" (D. 
Reidel. Dordrecht). 

Wassov R, J. (1976) Holoccne aeolian lamlfonn.v in the 
Belarabon area. S.W. ol Cobar. N.S.W, Journal Pnn: H. 
Sac. NSMi m 91-101. 

(1992) "Modern Sedimentation and Late Quaternary 

Evolution of the Magcla Creek Plain" (Research Repoit 
6. Supervising Scientists for the Alligator Rivers Region, 
Australian Government Publishing Service. Canberra), 

Wit.t.iAMS. M. A, J. (1984) Palaeocl "mates and 
palaeoenvironments: (a) Quaternary environments, pp. 
42-47 In Veevers, J. J. (Ed } "Phancrozoie harth History 
of Australia." (Clarendon Press, Oxford). 

_ (1991) Evolution of the landscape, pp. 207-221 In 

Hayncs, C. D.. Ridpath. M, G, & Williams, M A. J. (Eds) 
"Monsoonal Australia - '. (Balkema.. Rotterdam). 

, Adamson, D. A & Baxter, J. T U98h) Ute 

Quaternary environments in the Nile and Darling basins 
Australian Geogruphicul Studies 24. 128-144. 

_____ . De Deckkek. P. Adamson. D. A &. Talbot. 

M. R. (1991) Episodic ttuviatile, lacustrine and aeolian 
sedimentation in a late Quaternary desert margin system, 
central western New South Wales, pp. 258-287 In Williams. 
M.AJ, De Deckkei. P. & Kershaw, A. P. (Kds) The 
Cainozoie in Australia: A Re- Appraisal of the F.vidence". 
(Geolosieal Society of Australia Special Publication No. 
18. Sydney). 

__, . DonKErley, D. 1... Dr Deckkbr, P.. Kershaw. 

A. P. & Stokes. T. 0993) "Quaternary Environments'*. 
(Edward Arnold, London). 

Woodrofee, C. I>.. Tjiom, B. G. & Chappeil, J. (1985) 
Development of widespread mangrove swamps in mid- 
Holocene times in Northern Australia. Nature 317, 711-713. 

. Chappfi t . 1. M. A.. Thom, B. G. & Wallensky. 

h. tf98o) "Geomorphological dynamics and evolution of 
(he South Alligator tidal river and plains. Northern 
Territory". (Australian National University, North Australia 
Research Unit. Mangrove Monograph No. 3, Darwin) 


ByR. W. FiTZPATMCK*f & M. J. Wright* 


Fitzpatrick, R. W. & Wright, M. J. (1994) Climate change and its implications for 
South Australian soils. Trans. R. Soc. S. 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 puipose 
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 (ii) 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. 

Jwnxaaums of the R/tyat Society of S. Au\L (1994), 1W(1) 27-34 


by R W. FuzPATRiCK"-*t Si M. J. Wright* 


fiT/^ATPjCK, K. W. & Wku.iii, VI. J. (1994) Climate change and its implication* for South Aii*ir«l>tm suu> tfrmS 
/?. Soc £ 4»t< U»(1). 27-34, 31 May. 1994 

The nature and distribution of dominant soils in South Australia is briefly reviewed with particular reference 
to major must relating to climate change. For this purpose the Stale hay been divided into two [t&H)&K (•) the 
southern leeion <_<r agriculturally developed area which lies south of latitude &2°S and is mainly used [or dryland 
cereal/sheep production and (,ii) the northern region or semi-arid and avid areas which mostly lie north of Iwtiuide 
.12°S and are mainly used lor low intensity giuiing of natural langeland A large proportion of South Australia, 
including many texture contrast soils of rhe high rainfall areas, has dispersive soils Mibject to sodicily and which 
are highly prone to waterlogging and wliniiy throughout a significant proportion of the profile. 

i his paper attempts to forecast the most likely consequences of global chmaie change on the dominant noIIs 
of South Australia. The- direct influence of increasing winter temperatures and decreasing vvimer rainfall on Ihe 
wide range o\' soil types (hat occur in South Australia is a> ycl unclear, However, the magnitude and extent of 
supposed degradation or enhancement of particular soil morphological propenie*. and decline or increase ol 
sail site properties leg. salinity or en.>sioni are mailers ol speculation, hi tact, these changes can affect the processes 
of global climate change by affecting production of greenhouse gases or causing chatties in vegetation. Mow 
vails modify under Ihe influence of changing climate will depend principally on soil type topography and changes 
of vegctarion 

South Australia's soils will respond to global climate change through changes in *oil motM-ire. FtftJ temperature 
and soil organic matter content. Decline in winter rainfall m Ihe high raintall regions occurring to (he *outh ol 
UtiiuOc 32°S will have ,< beneficial effect by substantially reducing seasonal waterlogging and formation of aquic 
iOils (including non-tidal acid sulJak* soils). In contrast, dryland salinity may continue to expand with a corresponding 
decline in stream water quality. 

It is not yet possible to accutalcly quantify regional soil changes resulting I mm climate cha igq given ihe present 
uncertainty about the amounts and rates of global climate change, and particularly concerning regional patterns 
of temperature, precipitation and coastal gcomotphic change*. The cuneuily increasing rate of land and watct 
exploitation in South Australia will likely have a greater impact on soils, adverse or bcnclicial, than the effects 
of climate change. 

Kav Wokos: climate change, soils. South AusUaliu. fioil moisiim:, si>il tcmpcraimc, sod organic content 
■ialinity. »odieily, soil acidification. 

Introduction Has been estimated over the lasl 100 years, eshtnaies 

Ctt warrfling up to 5°C over the next 100 years have 

The ^teeidiouse effect is the warming of the earth been made (Bouwman J990), A corresponding 

resuming from increases tn Ihe concentration of carbon warming of the oceans and rise of sea level of up to 

dioxide (C(> 7 ) and other radiatively active gases 0.5 mare also forecast. Changes of this magnitude and 

including methane (CH 4 ), nitrous o\ide <N 3 ()). o/one rale represent a significant change in our environment 

(O-,) and the chlorofluorocarbons (CFC's; thai reduce thai would have a profound effect on the Earth's 

ihe loss of outgoing infrared radiation (i.e. limit heat ecosystems and human activi'.ics (Bolin et «/. 1986. 

looses from earth into space; Tabic I). A global Wild 1993). 

increase in CO, has been reliably documented during Much of the emphasis in reecm discussions of the 

the UK) years and although this is due mainly to m ,p a ct of greenhouse induced climate change in 

the burning of fossil fuels there is an appreciable flux Australia has focusscd on the atmosphere and on 

of CO* from ihe oxidation of soil organic matter and agricultural and forestry production (Pcarman 19JSX). 

the burning of forests (Table 1). CO, levels of twice However, the effects df 'climate change on Australian 

the present axe forecast by the year 2050 (Peatman ^ \\^ j s a t s0 u fundamental issue Given the present 

1988; Houghton 1990: van Brccmen & Feijtel 1990). statc f scvcrc so j| degradation in Australia (e.g. 

Soil is not only a source of CO, but also of CH A and Chartres et ai 1992) the possibility of further 

N,0 (Table 1). Atmospheric warming of about 0.5°C degradation should be of national concern. 

- - — ■ South Australia is a region which is currently 

- CSIRO Division of Soils, Private Bag No. 7. Glen Osmond. ft-Rucnced imin | v bv Wliue ~ r rainfall bul where the 
South Australia 5064. ... ' .* * 1 . 

t Co-operative Rcs.ea.tih Cent* ri>i Soil aiut Umi ciinvm ramlall vanes grcaUy from north to south 

Management Hence, lot discussion purposes, we have divided South 



"a c 

"5 ■** 


3 5 


5 B 

-1 -^ 

S *3 

*1 2 

- eg 

c o o 

— ODL-Q 

p. E3. 

c e 







s g 



cr r i 

oc 1 

1 OB 

q. c- 

i 1 

J 5' 


"3 S 

C C ^ 

— O r-i ■— -t — f> 


C* r-l o> 

OC' — — E*1 — 

,5Ji O 'T — o c; QJ 3 
* u c 1 j§ — * 3 


Q Co 



— — "", — in *C fN t f3 "53 


a — ^- .— 

s 3 

in CL bB 

C t* B| 5 S 
■""■sic &*s 

c --- 

lis t 
1 §£? 
















Australia into the following two regions' (i) the 
southern region the agriculturally developed area 
which lies south of latitude 32°*S and is mainly used 
for dryland cereal/sheep production and (ti) the 
northern region - the semi-arid and and areas which 
mostly lie north of latitude 32°S and are mainly used 
for low intensity grazing of natural rangeland. I he 
greenhouse effect is likely to influence the soils in each 
region differently. The southern region is considered 
likely to suffer a drying climate due to a reduction in 
winter rainfall whereas the northern region is more 
likely to receive enhanced summer rainfall. The 
objectives of this paper are to; (i) summarise existing 
information on soils in South Australia, (ii) highlight 
the most likely consequences of global climate change 
on the range of South Australian soils and (iii) identify 
future priorities for research on global climate change 
on South Australian soils. 


Soils are complex systems which arc strongly 
influenced by processes occurring in the atmosphere, 
biosphere, hydrosphere and lithosphere (Fig. 1). 

In general, soils develop slowly over thousands ol' 
years, the absolute rates depending to a large extent 
on past climates. Their component parts also differ in 
their rates of development For example ( soU biotic 
processes resulting from die interaction of the 
atmosphere and biosphere are much more rapid than 
soil weathering processes resulting from the interaction 
of the hydrosphere and lithosphere. Thus, in response 
to greenhouse induced climate change, the more 
environmentally sensitive soil biotic paicesses would 
be expected to respond more rapidly (Bouwman 1990: 
VaraUyay 1990; Wild 1993). In coastal landscapes, 
many soils would also suffer relatively rapid water 
erosion, waterlogging and salinisation as sea levels rise 
with global wanning. 





Fig. 1. Simplified diagrammatic representation illustrating that 
soils are complex systems which are influenced by processes 
from the atmosphere (climate)- biosphere Ivegelation ;-ind 
fauna), hydrosphere (hydrology) and lithosphenr (geology 
and topography). 


hillucttce of had dealing 

An important factor in assessing greenhouse effect 
is the change already brought ahoul lo Australian soils 
tl o result of European settlement, Land clearing. 
grazing and cultivation generally have resulted in a 
rapid decline of soil otgunic matter, and organism* and 
consequently a deterioration o! soil structure and 
increasing susceptibility lo erosion. A marked 
reduction in the diversity of fauna and flora of most 
ecosystems has also accompanied European settlement 
and this has made our soils and landscapes less resilient 
to change 

The effects of greenhouse induced climate change, 
therefore, must he assessed for soils and landscapes 
already degraded: pusline condilions are virtually non- 
exisieni. Given this scenario, there is a significant 
environmental question to be answered; If there is 
greenhouse induced climate change of the magnitude 
forecast, what will be its effects on South Australian 
soils and landscapes"' 

Soil Carbon and the CO^ cycle 

The amount of carbon stored in the world's soils as 
fresh organic mailer, stable humus and charcoal, is two 
to three times higher than carbon stored m natural 
vegetal ion and crops (Table I, Rouwman 1990; 
Standing committee on international programs oflhe 
International Society of Soil Science 1990; Wild 1993). 

Distribution and major properties ofsoih in 
South Australia 

Generally speaking, South Australian soil landscapes 
are extremely variable ftnd complex due. in pan. io 
the great age ol much ol the continent- A signilicant 
proporiiou (>609() of soils in South Australia are 
>alme, or sodic and/or texture contrast (i.e. duplet* — 
have bleached sandy 10 sandy clay loam topsoil 
honmtis overlying clayey subsoil hori/ons lhai may. 
or may not be mottled.) (Table 2) Sodic duplex sous 
are inherently subject hi waterlogging because of water 

perching on the more impermeable clayey subsoil 
hon/on and consequently saturating the lopsoif 
hon/ons, especially where there is an increased mpui 
of water (bllowing removal of native vegetation, A 
similar effect can be caused, in the short term, by 
excessive irrigation. Therefore, to proceed from a more- 
general understanding of the processes of waterlogging 
and dryland salinisation to a more definite 
understanding of these processes in an individual 
catchment; in order to predict the effects of 
management or climate change, is extremely difficult. 
The central »nd southern Ml Lofty Ranges area is 
luithei complicated by highly variable ecology and 
weathering patterns of the soils in those landscapes. 
Il is apparent that in adjacent landscape positions one 
is confronted with deeply weathered soils Much 
contain ancient stored salt, juxtaposed with very 
youthful soils on partly weathered rocks which arc 
generating salt as a result of contemporary weathering 

The higher rainfall areas (i.e. greater than 500 mm 
average annual rainfall) which lie south ol latitude 32°S 
are restricted to coastal and sub coastal plains and 
ranges. A high proportion or both regions has soils 
which arc sodic throughout or through a significant 
proportion of the profile. Table 3 Itsts the estimated 
areas of sodic soils in South Australia, south of latitude 
32 °S. Tftc 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 fof reproduction here. 
The semi- and arid regions north of this l.iiilude conlain 
even more significant areas (if saline and sodic soils. 
particularly the desert loams and associated red clays. 
The lable was prepared by re-interpreting maps 
published by Norlhcolc (I960) and Northeote et al, 
(1968), Sheets 1 and 10. respectively, erf the Atlas of 
Australian Soils; die original scale of which was 

|£ WW 00ft 

Tabic ? I attribution ofMtline u/ui sodic satis in relation m rainfall m South Australia {a/cor Nttrilitwtu- & Skene 1972 


(run it 




Percentage area oi each map unit within anniwl rainfall /ones 
Alkaline strongly '-nciie Non-ulkulinc noetic 

Gradations) Duplex Neutral Acid 

profile duplex duplex 


Lex lure 






450.2 W 




25 y 

2 3 

6 y 
J 5 

<* ! 

22 4 


24 .2 

5 9 

i qui voj,uuo 
-■ — - - - 

■ All wnhin northern region of South Australia. 

** Largely within northern reeion hut ;i significant in'Ca of southern region incliHlcxI. 

*** All wnhin MjintiOin region c*i_epi lor small aavi ol northern reeion in 250-350 nine. 

2 8 






On an area basis two broad groups of sodic soils 
arc prominent in the southern region, the £rey-hmwn 
calcareous loams (36$ of the area) and the duplex .soils 
with alkaline soil reaction trend <I8%), thai is. soils 
v»ilh pH generally increasing with ileplh and alkaline 
in the subsoil. These two soil groups include the 
majority of the cereal and sheep producing land in the 
State Although of much smaller area, die sodic duplex 
soils of southern byte Peninsula. Yorke Peninsula, die 
Mid-North, eastern Mt Lofty Ranges, Kangaroo Island 
and the South-East are critically important t>ecause they 
have high rainfall and a high production potential. They 
are also important because they indicate the regions 
where waterlogging and secondary salinisation is a 
current or polenoal problem. 

I ",\H!.fc 3. Lstimuteif orats of Suite in South 4nsftaUo SVftlh 
nfiohwde JPS (modified after Nauht et al. W&j, 



Sods tlwt are >odic ihrcuighout: 
al Clayey soils, uniform texture 


Grey-brown calcareous loams-, 

umtonn and giadational texture* 


Alkaline duplex ".oils 

Sun- lota! 
Soils that have sodic supm.hIs 
(I) Alkaline duplex soils 
ci Neutral duplex suite 
Acidic duplev *oils 






Total sodic soils 
Nun sodie sr*lK, 


Alkaline duplex yuift 

Neutral duplex soils 

A: ulie duplex soils 

Clayey soils.. unifi>rm i.eunre 


Grey -brown calcareous I'umis. 

uniform and gradattonal wxtute 














12 M 










*ndy soil* inciKlimj fM>U>ok 

Tout non Addic m-iI 
LacwJ area south df latitude 32°S 









Projections for climate- change 

Forecasts of greenhouse induced climate change are 
based on global models th;it incorporate the. circulation 
of the Earths atmosphere and ocean* and the albedo 
effect of its polar tec caps. Present models vary in their 
forecasts of global CO? and temperature changes 
although a commonly acceprcd view is rhal a doubling 
oi C0 2 will result in a temperature rise ot 4°C 
(.FVnrman 198X; Wild ls*>3>. These models uIm> predict, 
to varying degrees, a consequent mcre.*s-e in 

evaporation and precipitation i.e. an intensification of 
the hydrological cycle with increased rainfall crosivity. 
Greater uneenaimies exist m the prediction of regional 
climate change arid ils consequences within continents. 

In Australia, a v best guess* scenario has been 
developed by Pittoek & Nix (I9S6) based on historic 
records and a global model from which a rise Oi 
between 2 and4,7°C in mean annual temperature by 
the year 2030 is predicted (see also Pearman I4SK). 
In summary, temperature increases are predicted to be 
greatest in southern Australia, particularly in winter; 
summer rainfall ureas will receive an mcicasc in 
precipitation of up to 50% t and the monsoonal 
mltuence will extend larthe.r south; the winter rainfall 
areas of southern Australia will receive less 
precipitation, particularly in winler. 

The climatic models vary even more widely in their 
prediction ot the moisture slaius pf Australian soils 
under doubled atmospheric CO, Only a broad 
generalisation i)l increased soil moisture in the mirth 
and a decrease in the south can be made. 

The uncertainties in predictions ol climalc change 
into the next century are considerable. Nevertheless, 
it is clear thai even a modest greenhouse effect will 
significantly change the regional climatic regimes of 
Australia. In the southern pans of the continent, 
particularly, the climate change forecast will create 
drier soil moisture conditions and impose further 
limitations for a range of land uses. When this scenario 
is superimposed on ihe present pattern of soil 
degradation in South Australia, it i* apparent that large 
areas of land could be at risk bf further degradation 

Not all the predicted consequences oflhe greenhouse 
effect, however, arc negative. Toroeast increased 
precipitation in the northern part of South Australia 
would have the potential to increase soil moisture stores 
with a positive outcome for soil stability and for die 
pastoral and agricultural industries. In contrast. 
decreased precipitation in the higher rainfall areas of 
southern South Australia would reduce seasonal 
waterlogging (sec beluw). 

Impact of past climate change 

Because soils and landscapes develop over long 
periods, many are found to contain information about 
past climate change Among the most significant 
examples are the soils and associated sediments of 
coastal flood plains about Australia (Division of Soils, 
CSIRO 19837. Al ,nc P eak °*' mc worltTs last glaciatinii 
20,000 years ago, sea levels were about 140 in below 
the present and the Australian coastline wa.< 
consequently much more extensive. At the end of 
glaciation, global seas rose rapidly in reach their 
present level about 6000 years ago. During this rise. 
ocean waters invaded large atcas of coastal land add 
embayed all coasiul rivers. The rivers deposited 



.sediment in marine and esiuarinc environments so tnur. 
today, their coastal flood plains contain a cominiioiis 
record of sedimentation. Around Australia, high walu 
tables, high salinity, and sulphates characterise ihe soils 
of these Hood plains. 

Taw i 4, Possible negative modifications of fund use due to 
rfianys- in wmperatun\ rainfall ami wtttdines* in South 


Nisittx sea levels. 

• wave erosion 

• flooding of consul flood plains 

• salimsation ol coastal Hood plains 

Northern Region (Arid ami ,omi ariil zone* winch 
mostly lie mirth of laliimle vT°S) 

bn reused rainfall and crosieilw sh^hlh higher 


• increased risk of landslides 

• ini.reased risk ot filter erosion 

• (louding of lowlands 

,Sou(hi'rn Region (High mtnlall /ones which lit suUiti Ol 
latitude 52?S) 

Cha/iyd rainjidl im ulrni r an n a\rj riosn'tty; 
higher lempe'aiurr.s 

• lower soil moisture 

• decreased vegetative covet 

• low of soil biotic component* and organic mallei 
■ ilcr^notarum of soil structure 

• increased risk ol water ciosion 
» increased salimsation 

• cliMeonraluitl cA Mreani wuter quality. 

• increased winu erosion 

• dufctter atmosphere 

lrt such coastal landscapes, the late geological pas; 
provides an accurate guide for predicting the 
consequences of sea level rise resulting from the 
greenhouse effect. These are increased Hooding ami 
an elevation ol water tables, and increased salimsation 
aad encroachment of marine and cstuanne 
environments, accompanied by a build up of sulphidic 
sediments in expanded mangrove swamps (Tables 4, 
3L This will lead to the development of greatet area.s 
of acid sulphate soils. The area ol coastal Hood plain 
actually affected, particularly in the gulf regions of 
Sourh Australia^ would depend on the magnitude O'' 
the sea level rise; a special study would be required 
tor reliable estimates. 

Rises in sea level also cause widespread wave erosion 
of coastal soils, near shorelines, as well as 
desmhilisation of coastal dune landscapes (Table 4) 
Much of the evidence lor these effects in the late 
geological past have been covered by the ocean or 
obi iterated. Nevertheless, a significant risk to coasdincs 
around South Australia is apparent (Harvey ci Relperio 

this volunie). 

T\ui i 5- Possible fit nil Ifa nwdifi-ianons in Umduse due to 
Ju/nge.s in temperature, ramftdl and windinrss in South 

I "oast line 

Rising seu level* 

• flooding and formation ol" coastal mangrove swamps 

Northern Region (Arid and semi-aiid /ones which 
mostly lie north of latitude 32°S) 

hn erased rainfall: slightly higher temperatures 

• increased soil moisture 

• increased vegetative cover 

• i educed Water ero.sion 

• increased simi nrgiimc mutter 

Southern Region iHmh rainfall rones which lit south of 
latitude 32«Sj 

Ihtri-awa 1 rainfall: higher temperatures 

• towei wider tables — ground and perched I reduced 
incidence of seasonal walcrtoceinjjt 

• decreased soil m idiftcatinn 

The usefulness ol the geological record for predicting 
(he effect of greenhouse induced climate change on 
other Australian .soils and landscapes is much less 
certain. This is because much of the stratigraphic and 
geomorphie evidence tor climate change during the last 
20.0CX) years relates to world-wide (Pleistocene) glacial 
climates. In Australia, a large part of Tasmania and 
a small area of the mainland were glaciated, inland 
take levels were high, river discharges were higher than 
the present, forge areas of hill slope land were 
crosionully unstable and eroded sediment filled local 
valley floors In central Australia, the dry phases of 
the late Pleistocene were characterised by extensive 
movement of sand dunes and the mobilisation of large 
quanlilics ol dust which was deposited as a clayey 
mantle over landscapes in eastern Australia These 
unstable soil and landscape conditions changed to 
relative stability as climate warmed during the last 
10.000 years (Holoeenef 

If we use the past as an analogy, the predicted 
greenhouse warming should be seen as leading to soil 
formation and. therefore, uot cApceted to result in 
widespread instability and degradation of soils and 
landscapes in non-coastal Australia. However, two 
factors make the present different from the late 
geological past. First, the rate of el i mate 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 soils and 
landscapes have already been modified following 
Kuropean settlement in such a way as to make them 
more vulnerable to rapid environmental change. In 
many parts of South Australia, thresholds of stability 
have been exceeded, resulting in decelerated 
waterlogging, salimsation and erosion (fLupairtck ei 
at [992, (993). If present land use is not responsive 


m greenhouse induced climate change, these and other 
♦uriit^ til soil degradation are likely to increase, 
particularly In the southern region where decreased 
soil moisture would make present land use marginal 

in auine ureas 

In this connection, Pituxk & Nix (1986) have 
modelled changes in plant productively as a result oi' 
projected greenhouse induced climate change in 
Australia. In northern Australia, ihc positive outcome 
Qf ihe greenhouse effect m increased precipitation and 
vegetative cover could mean a shift towards an increase 
in the stahilily of sods to water erosion. As elsewhere, 
however, much would depend on land use practices, 
A decrease in plant production is anticipated in the 
southern region of South Australia, signalling a 
decrease in soil vegetative cover and soil biotie activity 
li is suggested that rhesc cffc-eis would he relatively 
rapid with a corresponding decrease in soil structural 
stability and an increased susceptibility to erosion In 
sourbem Australia, therefore, the ycai 2050could see 
a return towards conditions that prevailed in the lale 
Pleistocene given The tipper (worst J greenhouse 

Prajeclions frvr Soils and landscapes 

Pmm the foregoing, it is possible to outline some 
it| the mam consequences of induced 
donate change in South Australian sods and 
landscapes. Although these can only he hroadly staled. 
they have a potential value in identifying like areas of 
positive and negative outcomes These are summarised 
in Tables 4 and 5. It may be argued that some of ihese 
projeciion& are alarmist. Hovvrvcr, walcrh tggttg- 
sail ihisal ion. acidification, wind and water 0(06(00 are 
already major problems in South Australia and there 
is no evidence that ihey are under control (Fitzpatriek 
?l A 1W2. 1993. 1994: Naidu et oi 1991). Even the 
con.scrvative "middle scenarios' of greenhouse induced 
climate change would exacerbate these problems in 
many areas. 

Soti eroitort 

Somh Auviraha has a high proportion of sodic and 
saline soils (Norihcote & Skene 1972; Naidu el al. 
I9°3). The >ttdic:sulmc ratio of approximately 5*1 
ranges between 4 and ID limes that reported tor olher 
continents (Szaboles I989| and is consistent with (he 
high proportion of sodium present in soil solutions and 
groundwaters. In Sooth Australia, mosi sodium 
affected soils are the resuti of past inundations by 
braekiwh water supplemented possibly by cyclic salt 
Thus, m MibsoiN, chloride is the dominant anion and 
exchangeable Mg/Ca ratios arc high. The incidence 
of sodicitv often coincides with the spatial distribution 
oi duplex- soils. This associauuu indicates a pattern of 
environmental hazard which appears to at least broadly 

coincide with many areas under agneuhuie in the 
southern region, and substantially limits its 
productivity. Saline, sodic and sodic duplex soils are 
often predisposed to land degradation (e.g. Naidu ft 
al. 1993. Isbcll ft at. 19X3) and, significantly, covet 
>oO% oi the South Australian land mass. The effects 
of adsorbed sodium on clay dispersion are most 
pronounced in dense alkaline subsoils which comprise 
over K6% of Australian sodic soils i,Northcoic & Skene 

The impact of soil sodicity on the environment is: 
a most important land degradation issue in South 
Australia Both primary and secondary sodiftcalionof 
soils causes undesirable changes in soil structure. 
severe hillslopc erosion, waterlogging and erosion oi 
downstream watercourses. The associated increase in 
colloid and nutrient loading of streams also contributes 
to sedimentation and die consequent loss of reservoir 
storage capacity, together wiih serious water quality 
problems. H(AVc*ver. it is the contact of sodium-afieeu'd 
soils with water in the form of ram splash, surface 
runotf. thmugh How. natural ground water flow, or that 
pumped tor irrigation from other catchments, which 
provides Ihe expanded scale on which environmental 
problems and hazards become most conspicuous 
(Fiupatnck et al. BBM). 

In South Australia generally, climate change is likely 
to incieasc the tncideoce of water emsinn (Table 4), 
Although lower precipitation in die south will generally 
reduce the rate of waiter erosion, rbi-s is counterbalanced 
by the less intensive sod conservation inlluence of poor 
vegetative gntwth due ID an inadequate moisture supply 
for plants (exacerbated by an increase in Temperature, 
as well). Lower precipitation and higher temperatures 
will ftlttfi lead tu increased wind erosion, especially 
where sandy soils arc dominant tie in the semi ai id 
northern portions of the southern region). In the 
northern region, increased sumrneT rainfall is likely 
to be of higher intensity than at present and is thus 
expected to present a higher erosion risk there loo, 
particularly to the widespread saline aind sodic desert 
loams and led clays ot the stony tablelands. 

Wulc Hogging and dryland .utlhury 

Extensive deforestation which followed Eumpcait 
settlement in South Australia hies contributed to a 
detrimental shift in the hydrologie balance ul ruauy 
catchments. This has resulted in a rise of perched ami 
saline groundwater tables causing waterlogging and 
dryland ■ alinii •. fc>bc a major land degradation problem 
in South Australia Over 220,000 ha of land arc 
estimated to be affevied by dryland salinity throughout 
ihe major agricultural districts al an annual cost to this 
stale of approximately S25 million in losi agricultural 
production. The problem is worsening. In particular, 
there i.s a growing concern by property holders in the 
Ml Lofty Ranges over the rapid mcrcase m saline. 



structureless acid sulphate soils lhat arc* waterlogged 
and nighlv panic lu water erosion i Fit zpatrick ei al 
1992, 1993 > 

Desphc an obvious reduction in seasonal 
waterlogging due lo j decline in winler rainfall in high 
rainfall region .south of latitude 32 C S, dryland salinity 
may continue to expand with d corresponding decline 
in stream water quality (Tables 4 and 5). According 
to Sadler ct ai (W8H), ihe. processes involved in such 
"saline" sod ic soil environments, due to reduced winter 
precipitation, arc further complicated by the balance 
between rales of leachmg and techarge. Overall, the 
consequent reduction of water quality and land 
capability are, therefore, only producls ftr A complex 
set of interactions which can occur between the 
following factors in Mediterranean climates sodium- 
affected soils, ambient levels of soluble sails and the 
temporal !low pattcius, spatial distribution, relative 
contributions and quality of surface water, and ihnuigh 
flow and ground vvaier. In particular, the leach a it' of 
saline godjc soils with good quality waier ne. 
ratnwaler) also poses a threat lo structural stability in 
buildings, bridges. earth dams and embankments 
(Rtzpatrick et ol 1994) 

Soil acidification 

tu the southern region, a reduction in winter 
precipitation may decrease downward infiltration and 
leach u g ol water and hence lower the rale of soil 
acidification. Similarly, a decrease in winter 
precipitation and increase in winter temperatures will 
discourage heath vegetation which produces acidi lying 
litter (Table 4). In contrast, in those presently high 
raimall regions where climate change will cause an 
increase in summer precipitation, soil acidification may 
accelerate dac lo *i ranger leaching and chemical 
reactiviiv (eg carbonate dissolution may occur? 

the northern regions, bionws. and hence charcoal will 
increase, whereas in Ihe southern region reduced 
rainfall will lead to a generally reduced biomass and 
consequent reduction in the amount ol charcoal 
produced, even though lire frequency could increase 
because of higher temperatures and lowered moisture 


Changing agricultural practices (that lead to erosion, 
saliiusatioii, sodi heal ion, acidification and 
waterlogging) make it difficult to monitor the effects 
of climate change in individual catchments. Special 
research that takes account of these complications is 
needed on specific South Australian soil types lo 
quantify' regional effects of climate change and to plan 
strategies co ciroe with the changes because of 
implications for the well being of our communities and 
the environment, It would appeat thai the dotmuantly 
pastoral activities of the northern region stand to benefit 
by reason of increased summer rainfall at a time when 
temperatures are suitable tor major growth of herbage 
Salinity of surlace soils, at least, should aKo be reduced 
by the increased precipitation (Brinkman & Brarnmer 
1990). In ihe southern region, on the oihei hand- 
agricultural pursuits could be adversely affected by 
reduced rainfalL although increased temperatures will 
improve winter growth of cereal crops. Agricultural 
activities may also be able to expand somewhat in the 
northern marginal fringe to take advantage of increased 
rainfall lliere. Finally, at a time when the demand lor 
good quality water is escalating, the limited 
understanding i\i the factors and interrelationships 
imyjlved in soil and water management may well be 
further complicated by a changing global chmaie 
induccd by the "greenhouse effect" 

bioltigicul changes including the nutrieni flfg/itlfi 
OraeU et at. (19$$ in discussing the implications 

ol'climate ehangi lor and zone vegetation .stresses Ihe 
importance of the spatial patterns of soil types, 
furthermore, given the importance of this spanal 
dimenMon. there ts a tequirement, as yet not met. ior 
models of vegetation response to plant-available 
moisture and available nutrients lhat arc specific to the 
major soil types. Three requirements were identified 
for forecasling vegetation change. The first was that. 
because the spatial patterns of soil lype arc slill 
expected to determine the distribution of vegetation, 
it is essentia! that the forecasts of future climatic 
conditions have a spatial resolution equivalent to that 
of the mapped soil landscapes. 

We consider that much of the carbon stored in South 
Australian soils is in the torm ol charcoal due to a long 
history of burning. With climate change in ihe 
directions discussed, it is reasonable to expect that in 


The authors wish to acknowledge Dr P. H. Walker. 
formally of CS1RO Division of Soils, who contributed 
significantly !o this paper by providing espert advice 
in the torm of .i draft review ;hai formed an original 
framework for it 


Bkinkmain, R. & Bkammkk, H. (WO) The influence r>f a 

changing climate on Soil prupenies. /m/M. 14th tni, Ctfntth 

Soil Sri,, tytfta \99il 383-2ffi 
BdUfc, B.. DuQfc B R & Dt MotH. J. <Cds ) (WR6) The 

Greenhouse effect, climate change, and ecosystems". (Wiley. 

tlodWMAis. A. Pi (Pd.J I W) "Soils and the Grcenlmirse 

Efrecf. (Wiley, Chichester). 



Chartres, C J.. Helyar. K. Wy Fitzpatrick. K. Yk & 
Williams, J. (1992) Land Degradation as a result of 
European Settlement of Australia and its Influence on Soil 
Properties, pp. 3-33. In Gilford. R.M. & Barson, M.M. 
(Eds) 'Australia's Renewable Resources Sustainability and 
Global Change". International Geosphere-Biosphere 
Programme Australia Planning Workshop, Oct 2-4 1990, 
Bureau of Rural Resources Proceedings No. 14 v (AGPS, 

Division or- Soils, CS1RO (1983) "Soils; an Australian 
viewpoint" (CSIRO: Melbourne/ Academic Press, 

FITZPATRICK, R, W . NAtDi;, R. & Snu. P. Q. (1992) Iron 
deposits and micro organisms occurring in saline sulfidie 
soils with altered soil water regime in the Mt. Lofty Ranees, 
South Australia, pp. 263-286 in Skinner. H. C. W? & 
Fitzpatrick, R. W. (Eds) "Biomineralization Processes of 
Iron and Manganese - Modern and Ancient 
Environments". Catena Supplement No. 21. 

, Hldnall, W. H., Self, P. G, & Naidu, K (1993) 

Origin and properties of inland and lidal saline acid sulfate 
soils in South Australia, pp. 71-80. In Dent, D. L. & van 
Mensvoort, M. E. F. (Eds) "Selected papers of the Ho Chi 
Minh City Symposium on Acid Sulfate Soils" International 
Inst. Land Reclamation and Development Publication 53. 

, Boucher, S., Naidu, R. & Fritsch, E (1994) 

Environmental consequences of soil sodicity. Aim. J. Soil 
fa& 32 (In Press). 

Gr*etz ; R. G.. Walker. B. H., & Walker, P. A. (1988) 
The consequences of climatic change for seventy percent 
of Australia. In Pearman. G. I. (Ed.) "Greenhouse: planning 
for climate change" (CSIRO, Australia). 

Houghton, J. T. , Jenkins, G. J, & Ephrauvts, J. .1. (Eds) 
(1990) "Inter governmental panel on Climate Change." 
(Cambridge University Press. Cambridge), 

Iskell. R. F j Reeve, R., & Hurras;, J T. (1983) Salt and 
sodicity. pp. 107-11. In "Soils: an Australian viewpoint". 
(Division of Soils, CSIRO Melbourne/Acadcimc Press. 

Naidu. R., Merry. R. H., Churchman, G. I, Wright, M. 

1, Murray, R. S., Fitzpatrick, R. W. & Zarcinas, B. 

A. (1993) Sodicity in South Australia; - a review. Ausi. 

J Soil Res., 31. 911-929. 
Northcote. K. H. & Skene, J. K. (1972) Australian soils 

with .\alme and sodic properties. CSIRO Soil Pub, No. 27. 
Pearman, G. I. (Ed.) (1988) "Greenhouse: planning for 

climate change". (CSIRO Australia). 
Pittock, A. B. & Nix. H. A. (1986) The effect o! changing 

climate on Australian biomass production - a preliminary 

studv. Climatic Change 8. 243-255. 
Sadler, B. S., Mauoer, G W. & Stokes, R. A. (1988) The 

water resource implications of a drying climate in south- 
west Western Australia, pp. 296-31! In Pearman, G. E 

(Ed.) "Greenhouse - planning for climate change'. (CSIRO, 

S/abolcs. I, (1989) "Salt affected soils'. (CRC Press, Boca 

Raton, Florida). 
Standing Committee On International Programmes Of 

The International Society Of Soil Science (ISSSi 

(1990) "At global change . Do soils Iptta? 1 (do ISRJC, 

P.O. Box 353, 6700 AJ Wagerungen, Netherlands), 
Van Breemen. N. & Feijtel, T. C. .1. (1990) Soil processes 

and properties involved m 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 ol climate changes 

induced in soil degradation processes. Trans. 14th Int. 

Congr. Soil Sci.. Kyoto, 1990. 265-270. 
Wild, A. (1993) SoUs and the environment: an introduction. 

Chapter IE (Cambridge University Press, Cambridge). 



ByB. C. Bates*, S. P. Charles*, N. R. Sumner*, & P. M. FLEMiNGf 


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. 

tnmsththm t (l rftr X«\al Soaw $& Aval HWk Kftftli 35-45 


hy B. C. Bates* S. P, Charles*, N. R t Sumnu**, & R M. FfcEMiNtil 


Hrvii-i. B. C, CiiAhi.LS, S. "P., SuMrsbK, N. R.. & t*i pmisc. P M (W94) Climate change and its hydrologie implication*, 
(or South Austria Iruns, H, Sot: S- Aust. 08(1), .15-43. 31 May. 1994. 

The possible effects of doubled atmosphciie concentrations of carbon dioxide on die North Para River ai Penriee culehnieni 
arc evaluated (isfrig results ftruti »» general circulation model, a stochastic weather generator and u conceptual water balance 
mtulel, Although results show only a marginal decrease in median monthly runoff during winter months, large increases 
in monthly I WWfl EBftMUM are indicated tor August and September due to large increases in extreme monthly raiutall Modest 
incrfatcr. in evapotranspiration were also indicated loi these months 

krv Wohos: Climate change, hydrology. South Australia,, water vield. 


Global climate change caused by rising atmospheric 
concentrations ol carbon dioxide (CO : ) and other 
trace gases may have a significant impact on regional 
water icsources. Recent research suggests that plausible 
climatic changes will affect the timing and magnitude 
ol runofl and soil moisture, change lake levels, and 
increase evapotranspiration (Cohen 1986; l.ellenmaier 
& (Jan 1990: Allen el ul 1991, Lcttcnmaier & Sheer 
1991- Mimikou 4t n\ 1991; Nash & Gleiek 1991; 
Pana.eoulia J992) Such scenarios have important 
irr»|>Ik"alit>r^ lor future watci resources planning and 
management, the environment and the national 

Most hydiologic scenarios arc based on the climatic 
predictions of numerical rni»dels of the general 
circulation of the atmosphere. General Circulation 
ModeK iGCMVi can produce long-term simulations 
Of the energy and water fluxes in the atmosphere and 
'. nil : anil ocean surfaces on a global computational gt id 
of cells and a number of vertical layers. Predictions 
of changes in climatic variables such a> precipitation 
and temperature arc considered to he more reliable than 
those tor runoff and soil moisture (Gleiek 1989)- The 
predictions are provided as spatial averages over areas 
of the order of Kr to UP krrr due to the limitations 
of present-day computers. Current GCMs perform 
reasonably well in simulating the present climate with 
respect to annual or seasonal averages at this spatial 
scale However the direct use of GCM outputs to drive 
hvdrologic models is considered lo be impnjper due 
to the coarse (relative to river basin scalei re-solution 

* Diviwion of Water Resource*, CSIRO, Private Bag PO , 

Wemblcv. Western Ausltalia 6014 
t Division of Water Resources. CSIRO. GPO Bo< 1666: 

Canberra, Australian Capital Territory 260! 

ol the spatial gfids used by GCMs and the simplified 
GCM representations of land surface processes, energy 
transfer within oceans, and subgrid-^cale atmospheric 
processes such as connective storms. Moreover, GCM 
based assessments of climate change are based on 
steady-slate simulations of currcnr climate and the 
climate associated with a doubling of current 
atmospheric concentrations of CO., 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 in increasing 
almospheiic concentrations of trace gases 

Consequently, assessments of the impact of climate 
change on hydrologic systems frequently use doubled 
C<) 2 scenarios consisting of u spectrum of uniform 
shifts to historical temperature series and sealings of 
historical precipitation series, based on GCM trends. 
The size of the adjustment may vary from month to 
month to relied seasonality in the assumed changes. 
ristimates of potential evapotranspiration for ihc 
changed climate scenarios are usually obtained by using 
simple scaling factors, sometimes varying seasonally, 
and by taking qualitative account of precipitation and 
temperature trends. The historical series and changed 
climate scenarios arc used as input to mathematical 
models or hydrologic pmcesses and Ihc model outputs 
are evaluated to discern possible changes in soil 
moisture and water yield for a given catchment. 

It is frequently argued that working with hypothetical 
scenarios suits the purpose of a sensitivity analysts ol 
WZU&T resources and that results are not intended to be 
a prediction oi changes. However, this approach sutlers 
from three major limitations. Firsl, it ignores any 
changes in the distribution and frequency of 
precipitation events and any changes in the nature and 
variability ol temperature semes. This may be 
regardless of whether such cnanees art* indicated K 



GCM predictions Second, the use of hypothetic ill 
•ccnarios based on an arbitrary number of temperature 
and precipitation perturbations and stationary wind and 
relative humidity series, say, is not realistic and may 
tuck, the internal consistency of GCM simulations of 
tuliirc climates. Third, concurrent historical climatic 
seties arc relatively short in length which may 
compromise the evaluation of the response of a 
hydrologte system to climate variability as well as 
climate change. 

An alternative 10 the conventional approach is to use 
t| stochastic model representing daily weather 
vai lalions at a location. The parameters o! such a model 
characterise the behaviour of the present day climate, 
Changed climate sequences can be produced by 
adiuslmg the model parameters in a manner eonsiNtetit 
with GCM trends. The model is then used to generate 
long-term sequences of synthetic daily weather records 
winch in turn are used to drive a hydrologic model 
tWilks 1992; Bates et al 1993; Charles et al. 1993) 
Such an approach allows consideration of changes in 
the distribution and frequency ol precipitation events 
ami changes in the variability ot other climatic 
variahles It also preserves the infernal consistency of 
GCM simulations of future climates 

In Ihis paper, an attempt is made to assess the impact 
of a doubling of current atmospheric concentrations 
ol CO* on the North Para River al Pen rice catchment 
in Sooth Australia., Historical climatic and hydrologic 
series and results Ironi a single GCM. a stochastic 
weather generator, and o conceptual wafer balance 
mode! have been used. ConcluMons jre drawn on 
possible changes in water yield as a result of climate 


of GCM rainfalls by Charles et al. revealed a general 
trend towards increasing rainlall amounts and changes 
m the rainfall occurrence process under doubled CO> 
conditions. The latter is consistent with the work Ol 
Gordon el al. (1992) who found marked changes \\\ 
the magnitude and frequency of extreme rainfall evenK 
when comparing results from equilibrium experiments 
with the CSW04GCM 

Stochastic uvathrr generator 

The stochastic weather generator used in this study 
is based on the WCtliN generator described b> 
Kiehatdson & Wright (1984). The daily climatic 
variables simulated by WGEN are precipitation 
recurrence and amount, maximum and minimum 
temperature, and global solar radiation {#,}. 

Precipitation occurrence is described by a Iwo-sink 
(wel or dry day), tirst-order Maikos chain. The 
transition probabilities lor a given location are allowed 
lo vary thtough an annual cycle by defining separate 
probabilitics for eus'h of the 12 calendar months'. 

h - & '■'■ 

h i - Fl 

where //„ = probability that a day in state I will be 
followed by a day in stale/ (;\y = denotes a dry day; 
/'. j = I deuoies a wet day): and ./„ ^indicator 
variable denoting the presence or absence ol 
precipitation on day /; The variation ol ptectpilation 
amounts on wet days is characterised using either a 
gamma or mixed exponential distribution. 

The temperature and solar radiation components are 
represented as 

itctwral limitation model 

The GCM used m this study. CSIRU9. has beeu 
developed by the CS1RO Division of Atmospheiic 
Research. The model operates with nine veitieal levels 
in the atmosphere and a horizontal resolution ol about 
MH) km y! 600 km, A computational lime step ol 3U 
minutes is used, The simulated climate data come from 
equilibrium (constant CO : concentration) runs lot- 
present day (control) and future (doubled COi) 
climates. A complete description of die model is .jo en 
by McGregor ct al (1993). 

Charles rt ul, (1993) reported distinct differences 
between the monthly rainlall patterns found m 
Australian meteorological station records and those in 
the LSINU9 control run. the differences were with 
respect to rainlall amount and its geographical 
distribution. Maximum daily gud cell rainfalls for 
CSIR09 control runs were also round lo he between 
% and ': of (he maximum daily rainfalls recorded on 
inesosealc svaletsheds. Nevertheless, the comparison 

x*(t) = A\*(r-h + Bern 

where x = (3 k \) vector n[ climatic variable^ 
i — (3 x 1) random forcing vector consisting of three 
independent standard normal voi tares; A and 
B - (J x 3) mali ices obtained fiom the lag-U and 
lag-l correlation matrices for the components of v 
(Malalas N67); and the asterisk denotes 

jf**(1 - *$) - tHiO) tnfr) 

in which k — J. 2. 3 and y - 0. Ir ft^O) = mean 
for climatic variable k and state j, and 
0\lt) = corresponding standard deviation. The annual 
cycles of i^i and o Ai ure. modelled by single Kiuner 
harmonics with fixed phase angle* 


tfov&r f+ttl - 7rH'l-Ki)'(M) 

investigation of historical Australian deify maximum 
and minimum temperature scries conditioned on wet 
and dry days has shown thill the use or' a single 
harmonic to describe annual cycles of means alul 
standard deviations is often unjustified Higher 
hat monies are present in the mean series and the 
standard deviation series are more realist ically 
represented by fixed monthly value* isee alijO 
McCjsUII 199:!). Thus a menu temperature varies f D 
may he written as 


where r = unconditional probability of 8 wet day 

h = P{\\H\ '/^i Pn)\'- <*- $ = sha r e and SL ' a,t -* 

parameters o*' (he gamrva distribution, and 
rf*— Pij-pui ls u "'leasure 6J persistence. For 
simulating doubled CO, conditions, wc construct (he 
ratios (Wilks 1992): 

Wl = N 

gJ^^iM |2:/rr (/ + #) f 3051 * I 

where fe * I, 2 / - 0. I: / = Julian dale. 
u — constant; wi<3; R f -amplitude of the rth 
harmonic, tt = 3.14 in this case; 4> = phase angle. 
and i, - flh residual. 

Similarly, Investigation of Australian solar radiation 
series has indicated lhai ihe titling ot single harmonics, 
the use Of a fixed phase angle and the use ot the 
generation scheme defined above is an inadequate 
approach (see Bates ct al. 1993) Our approach involves 
the calculation qf the upper envelope lor clear day 
conditions, ff ( .. (A documented FORTRAN-77 
computer program tor calculating the upper envelope 
can be ohtamed from the authors). This theoretical 
maximum is based on geographical location and 
average clear day atmospheric conditions For each 
calendar month, a generalised beta distribution is fitted 
to ihe daily residuals (the difference between the 
theoretical maximum and recorded daut) for wet and 
dry d4>*s. 

Ctinnuc sequences for doubled CO : conditions 

Wilks (1992) presented a method for the adaption 
of stochastic daily weather models fitted to current 
climatic series ro the generation ol synthetic series tor 
future climates. The adjustments to the model 
parameters were made in a manner consistent with the 
changes in monthly statistics derived from comparisons 
nl GCM runs for control and doubled CO ; 
conditions This approach was based on the notion that 
GCM results are often available in terms of monthly 
rather than daily values or even monthly means and 
variances, The C$IR(WGCM runs provide daily values 
lor 3d climatic variables tor 30 year periods- The 
variables include: precipitation, humidity at level I: 
maximum and minimum screen tcmperaiurv; 
temperature at level I; and net solar radial ion at ground 
level l/c\„) Consequently, our method for making 
parameter adjustments to stochastic daily weather 
models uses this information. 

Let P - monthly precipitation for a month 
comprised of A'duys, The mean of P and ns variance 
arc delincd hv | Wilks !W2> 

(i[P') _ irVfJ' 
tf.P) rap 

tW 3 [1 Ho' (H r ')il + </') < (!-</' H 
raff 1 [1 4-o(W>U ^ d) i {\-d')\ 

where the prime denotes doubled CO : conditions. 
The solution of the above equations requires two 
addiiional constraints, For catchments in the arid and 
semi-arid regions of Australia where tt < 5. we set 
tt' = -ttx./ttj and d' - dd : i </, in which the 
subscripts I and 2 denote GCM values for a nearby 
GCM grid cell tor control and doubled C(K 
conditions, respectively. 

For mean doubled CCK daily muximum and 
minimum temperatures, we assume: 

r k JLt) - T Lt {t) + %V) 


where the prime denotes doubted CO. conditions; 
T == hannorucs fitted lo the observed mean series: and 
7\ 7? -harmonics fitted to screen leinpeiatutes 
from the control and doubled CO : GCM runs 
respectively. Standard deviations are assumed to be 

Clear day solar radiation is affected by prectpitable 
water content and CO, concentration, Here we derive 
daily level ! vapour pressures using the temperatures 
and relative humidity at level I and hence upper 
envelopes lor (he control and doubled CO> GCM 
runs. The GCM solar radiation is R s = R xr , f (I a) 
where a is Ihe albedo assigned to the GCM grid cell. 
Generalised beta distributions arc fitted to the 
corresponding residuals (R„-Rj 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 each month are scaled by the 
ratios of the corresponding doubted COj and control 
means and variances, respectively. The uppei envelope 
for doubled CO, conditions is obtained by scaling 
historical dailv vapour pressuics by the ratio of derived 
doubled CO. and control GCM vapour pressures. 



RfeW kahmte model 

\ modi Tied version of Boughton's (1984) SFB model 
was used for the study (Fig. 1), The model uses daily 
rainfall and potential evapotranspiration data us input 
to estimate monthly Kirearnflow. The original model 
has three parameters requiring calibration; the surface 
storage capacity [&). the daily infiltration capacity (F) 
controlling the movement of water from Lhe surface 
store to lower .store; and the bascflow lactor \R) 
determining the portion of the daily depletion ot water 
in the lower store that appears as bascflow (0^-6<l). 
The four remaining parameters arc fixed: the fraction 
of the surtace storage capacity that does not drain to 
the lower store [NDC = 0.5); the maximum limiting 
rate of cvapotranspi ration (E„ nn = 8.9 mm d '); the 
lower store depletion factor iDPF = 0,005); anil a 
bascflow threshold for the lower store 
(SDR,,, iLi - 25 mm) defining the depth of water in the 
lov*et store at which bascflow will cease. 

Dr a it i age 
= (1-MDC)S 

Men -drainage 

r^M o., 



= (I H1DPF5S 
l>'ij? I Structure ot the SFB modcK 

The model operations may be summarised as 
lollows. Rainfall begins to fill the surface store and 
any water in excess of the non-drainage component of 
thai store infiltrates into the lower store at a maximum 
daily rate of F mm d 1 . Surface runoff {Qj occurs 
when the drainage component of the surface store is 
full and mav be written as 


F tanh {P/F) 

where P = rainfall excess remaining after the surface 
store is filled. The surface store contents are depleted 
by evapotranspiration which occurs at the potential rate 

(£ ,) when the non-drainage component 


Otherwise, the actual evapotranspiration rate (E IT ) is 
given by 

£ = min (/■;,„„..> / (NDC.S): E M _ 

where j = depth of water in the non-drainage 
component of the surlacc store. The lower store is 
depleted by deep percolation (XJJ and baseflow \Q h ) 
which are defined by 


Q h ~ B.DPFiSS-SDK ) 

where SS — depth of water in the lower store. 

In this study, estimates of daily potential 
evapotranspiration in mm d were obtained using the 
Priestley-Taylor equation (Priestley & Taylor 1972); 

L £,,„ - art,, ftrU + y) 

where /. = latent heat oi' vaporisation in MJ kg ', u 
- 1.3 in this study. A = slope of the vapour pressure 
curve in kPa°C ', H„ ~ net radiation in MJ m z d ' , 
and y — psyehrometric constant in kPa°C l . 

Although the SFB model parameters have some 
physical basis, they cannot be readily determined by 
physical measurement. Thus parameter estimates must 
be obtained by fitting computed to observed monthly 
streamflow hydrographs. Formal optimisation 
techniques can be used to facilitate the estimation 
process. These techniques use a subjectively chosen 
criterion (the 'objective function") to quantify 
discrepancies between the computed and observed 
hydrographs for a given set ot* parameter value*- The 
estimates of the model 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 ihis study, the estimates for the SFB model 
parameters were obtained using a simulated annealing 
algorithm (Kirkpatrick emi 1983; Press er al. 1992), 
Simulated annealing is a stochastic, multivariate 
optimisation technique which seeks the global or near 
global minimum of a user-defined objective function 
without gelling trapped in a local minimum. The 
function need not be smooth or even continuous in its 
domain. The method can be considered us a biased 
random walk that samples the full parameter space and 
provides a solution that is independent of the starting 



point h can accept a 'move' ibai increases llic value 
ot Ihe objective function as a part of a full scries ijf 
moves for which Ihc general trend is lo decrease ihc 
llincliou value. Details of ihe algoriihin used will be 
published elsewhere, 

The objective function used in this study may he 
wtiucu as 

Estimates of daily potential evapotranspiration were 
obtained using the guidelines described by Smith 
(1491), clear day radiation and day length estimates, 
and temperature and sunshine hours data from the 
Nuriootpa Viricuhural climate station (34° TSFSL, 139° 
O'P.) which is located some 10 km from ihc catchment 

9 - H 

where the model parameters vector B = {S. F r B, NtX, 
B m t Dpft KO/L,,,)' in which Ihe prime denotes the 
transpose Of a vector, // = number of months in the 
observed streamflow hydrograph record; k = number 
of months in ihc 'warm-up' period which is excluded 
Ip'ih the calculation of ihc objective function value. 
and Ihe disturbance - is defined by 

X, K 

~, - <(?, + nei - 1ft + Rj) %i *»*0\ Xt>0 

in which Q = observed streamtlow. Q = computed 
streamtlow (g = Q h H- QX &VA Ki« ^ ~ 
Iranslormation constants which can be estimated by 
trial and error (Bates & Walts I9XS). Generally, Ihe 
best 1ms are ohtamed with \, < I and X> = 

Case Study 

Dt'MriptUm of ctihhtnem a*ut dam 

The /ibove methodology was applied in die North 
Para River at Pennce ealchmcni m South Australia, 
fhc catchment comprises an urcu of 118 keif nnd is 
located within Lhe Gawler Rivei basin some 50 km 
north cast of Adelaide ft r fec$ If™ 1 "* KW m lo 500 m 
AHD and has duplex soils and a rni\ed ground 
cover'. Land use includes horticulture, viticulture, 
grazine. and arable farming- 
Daily streamflow at the Penrice gauging Million 
(34° 2S'S. 139° 4*b) and daily rainfalls recorded at 
Angaston Post Office (34° WS. 139° 3'E) and 
Kcynetun(34 D 33'S. 139° 8'F) for ihe period January 
1978 to December 1989 were used in Ihe study Over 
this penod, the mean annual runoff was 6.200 Ml 
(50 mm) and the mean annual rainfall (65% of which 
falls in the period May to September) was 550 mm. 
The average daily maximum winter and summer 
temperatures were 13°C and 29°C, respectively (Chiew 
& McMahon 1993') 

| Cuirw, V H, S- & McMahon. T A. (1993) Complete >ci 
of daily rainfall, potential evap* rtranspi rati un and strcamJInw 
Jau for 28 uurctuliucd AiiMmlian catchments. C'enlir for 
I Applied Hydrology. Uni versus- m 
M**Hntnmc. S3 pp. (unpubl ) 

Moth'i cahhrutUw 

Preliminary investigations revealed lhat the 
perlbrmauce of the 3-parameier SFB model was 
inadequate. However, satisfactory Model fits were 
obtained by %elring A, =0.6. £„ M , = S.9 mm d' 1 . 
ADC = 5. SDR lti/i} = mm, and allow me the 
paiamcters if, ft B. and OFF to vary during ertlibrahon 
runs The final parameter estimates obtained were: 
S = 165 mm; F - 0.9 mm d ': B = 0.414: -md 
OFF - 12 The 5 and OFF estimates indicate that 
ihere is a significant loss of water fiom ihc catchment 
10 the regional groundwater. 

An important assumption in this study is lhat Ihc SFB 
model is able to simulate runoff ftOItl the North Para 
catchment under climatic conditions that are different 
lo those for which the model has been calibrated. For 
example, the efleci of possible changes m plan! 
transpiration rates and vegetative cover due IP CO, 
doubling Op the model parameters is ignored. This was 
considered to be a reasonable assumption glV6fl the 
current level of unecmontv regarding the nature and 
magnitude of these changes. 


To simplify the analysis of the results, the behaviours 
nt three variables under control (1 > CCK) and 
doubled CO; (2 « CO : ) conditions were examined' 
ill monthly rainfall: (2) monthly cvapoiranspiralion: 
and t3) monthly total runoff. Synthetic climatic 
sequences of 1,000 years duration were generated for 
control and doubled CO* conditions and used to drive 
the SFB model. Lxploralory analyses of lhe data 
revealed that the variable distributions were highly 
skewed, Thus Ihc median rather than the mean was 
adopted as the measure of* central tendency 

Fig. 2 compares lhe distributions o\ simulated 
monthly rainfall lor control and doubled CO, 
conditions. Three features arc worthy of note under 
doubled CO, 7 conditions: (1) there is an increase in 
minimum monthly ryinfall for the period from .tune 
to AugusL and a decrease lor Ihc months of May, 
September and October: (1) median rainfalls lor lhe 
period from March to November are greater than or 
equal to those for present day conditions: and (3) there 
are marked increases in extreme (high) rainfalls tor 
the month oi'January and the period from August to 

Fig 3 compares Ihe distributions of modelled 
monthly evapotranspiration for control und doubled 


Table 1. Seasonality and distribution of monthly runoff for control and doubled C0 2 conditions. 


Monthly Runoff (mm 




























1 7 










(0 4j 





















































III. 7) 




































































Values in parentheses are lor doubled CO : conditions. 

1 xC02 


1 S 



o - 


a s ■ 

- s 


£ U *-* LJ u 

m = 


: B 


o - 





Fig. 2. Box pjofe of modelled monthly rainfall for control (1 X C0 2 ) and doubted CO? (2 x C0 2 ) conditions. [Edges of 
boxes mark upper and lower quartiles, and horizontal blank line within each box depicts the median. Distance between 
the quartilcs is the interquartile range (1QR). End points of whiskers attached to boxes denote either data extremes (no 
adjacent horizontal hnes) or adjacent valued defined by upper quartile plus 1.5 X IQR and lower quartile minus 1.5 X IQR. 
Horizontal lines mark data points that lie beyond adjacent values.] 



1 xC02 
















Fig. 3. 

Box plots of modelled monthly evapotranspiration for control (1 X C0 2 ) and doubled CO : (2 x C0 2 ) conditions. 

1 x C02 2 x C02 


■sr - 

1 "" 


CM - 





o - 

1 — 












o - 










o _ 













o - 






o - 

H H H W 







R H 





Fig. 4. Box plots of modelled monthly runoff for control (I x C0 2 ) and doubled C0 2 (2 X C0 2 ) conditions. 



C0_ conditions. Theic appears to be a general 
increase in cwipotranspiration for the period from 
March It) November under doubled CO : condition*. 
Furthermore, a review of evapouauspi ration values 
greater than the 75th percentile point revealed an 
increase in high evapotranspnalion values over [he 
entire year. The st?e of the increase is quite large for 
the months of January. February and November 

Tnhle I lists various percentile points for simulated 
monthly runotf lor control and doubled CO, 
conditions, The distributions of the runoff data are also 
illustrated in Fig. 4. Overall, there is little evidence 
of changes in the median and upper and lower quartile^ 
Of monthly runotf. However, it is evident that the 
extremes (high runoff values) for the doubled COg 
climate are greater than those for present day conditions 
foi the period from July to October. There are also 
indications of reductions in the 10th. 30th and 50th 
percentile points for September. 

Fig 5 shows an empirical quantile-quanlilc plot of 
the annual maxima vf monthly iunoff. The annual 
maxima for both conditions are very lor 
maxima below their medians (about 11 mm;. Above 
the medians, the maxima are higher for the doubled 
CO. climate lhail for the present day and the 
differences between the series appears to grow with 
increasing percentile rank. Thus there is a suggestion 
of an increase in Hood risk A review of (he maxima 
revealed thai the extreme event* were due to the marked 
increases in extreme rainfalls and modest increases in 
cvapotranspiralion for August and September under 
doubled CO? conditions. 

Percentile ranks for 1 x C02 
50 so =~ 



*t - 


CU - 

» ■ 




& - 


... ........ 


c - 

D 20 40 60 80 100 

Annual maximum runorl for I X C02 (mm) 

Hi* 5. Hmpirical quamilc-qu-jntife plot of annual maxima of 
monthly iiiriuft tot cowrol - CO;) and doubled CO ? 
[2 -' GO-0 vomlitiuiis 


The most prominent effect of doubled atmospheric 
concentrations i>( carbon dioxide v>n the water yield 
of the Noah Para River catchment. South Australia, 
is a si/table increase in the annual maxima of monthly 
runoff. This may indicate an increase in flood risk. 
However, there appears to be little impact on the 
seasonality and magnitude of the median and uppe. 
and lower quart iles of monthly runoff. 

The above results highlight the benefits of usinjt 
.stochastic weather generators in assessments of tht 
impact ol climate change on hydrologie systems. The 
estimation of extreme hydrologieal events based oil 
original and perturbed historical records are not 
reliable when the return periods of these events are 
of the order ot the original record length, An impasse 
arises when the return periods of inteiesi arc greater 
than the period of record. Thus the length of the 
historical record limits what can be said about tin- 
changes in the tails of the distributions of hydrologk 
variables that may be caused by C0 2 doubling, Jn 
contrast, our approach is capable of pnxlucing climatic 
and hydiologie series pf arbitrary length. This enables 
a more detailed investigation of changes in interannual 

The results presented heroin were obtained using one 
GCM {CSfR09). a stochastic weather generator and 
a water balance model (SFH). Thus the quality of the 
results rely heavily on the accuracy and relevance of 
these models and the implicit assumption that the long- 
term 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 OCMj are fundamentally similar and that 
they differ principally with respect to their 
parameterisation ot certain processes such as cloud 
formation and rainlall Thus the use of one rather than 
a suite of GCMs should not seriously comptomisc the 
results Of this study 


J his work contributes to the CSIRO Climate Change 
Research Program and is part funded through 
Australia's National Greenhouse Research Program. 
We are indebted to F.H.S. Chicw and T.A. McMahon 
(Centre for Environmental Applied Hydrology, 
University of Melbourne), the Bureau Of Meteorology, 
and the Engineering and Water Supply Department ol 
South Australia for permission to use their data sets. 
Special thanks are due to the CSIRO Division of 
Atmospheric Rcse.uch for supplying CSIR09 GCM 




Aur\. R G, Gichlki, F N. &. Roshnzwhig. C. (1991) 

C0 2 -mduced climatic changes unci irrigation-water 

requirements. J! Water Rcsour Plug- M$>mt, Amer Sue. 

Civ. Eng. 117, 157-178. 
Bails. B C, Charles. S- P. & Fleming, P. M. (1993) 

Simulation of daily climatic series lor the assessment of 

climate change impacts on water resources pp 67-72- In 

C. Y. Kuo (Ed.) .'"Engineering Hydrology" (Amer. Sol 

Civ. Ung., Mew York). 
Bates. D M & Watts, D r G. 0988) "Nonlinear Regression 

Analysis and Its Applications." (J. Wiley. New York). 
Houghton, W. C. (1984) A simple model for estimating the 

water yield ot ungauged catchments. Civ. Eng. Trans, hist. 

Eng. Ausr. CE26, 83-88. 
Charit*. S, R, Helming, P. M- & Bails. B. C (1993) 

Problems of simulation of daily precipitation and other input 

time series for hydrological climate change models. Proc. 

Hydrvl. & Water Rcsour. Synjp, Inst. Big: Aust. Nat, Conf. 

Pub/, tfa 9J/M. 469-477. 
Cohen. S J. < 1986; Climatic change, population growth, and 

their effects on Great Lakes water supplies. Prof. Geog. 

38, 317-323. 
GLLtCK, P. H (1989) Climare change, hvdrology, and water 

resources. Rev. Geophys. 27. 329-344. 
Gordon. H. B.. Whetton. PH,. Pittock, A. B., Foavelr, 

A. M. & Haylock, M. R. (1992) Simulated changes in 

daily rainfall intensity due to the enhanced greenhouse 

effect: implications for extreme rainfall events. Climate 

ftymtW'A 8. 83-102 
KlRKPATRTCK- S., Gelatt, C D. Jr. Si. VtCCHI. M. P. (1983) 

Optimization by simulated annealing. Science 220, 671-680. 
LeTTENMAIER. D. P, & G\K, T. Y, (1990) Hydrologie 

sensitivities of the Sacramento-San Joaquin River basin, 

California, to global warming- Writer Resaur, Res, 2fi. 

& Sheer, D. P. (1991) Climatic sensitivity of 

California water resources, / Water Rexnur: Pfnt>. Mgrnt. 

Amer. Soc. Civ. Eng. U7, 108-125. 

Maialas, N. C. (19671 Mathematical assessment of synthetic 

hydrology. Winer Resour. Res, 3. 937 945. 
McCas-kill, M. R. (1992) Generation of long-term weather 

data sets for Charters Towers. Dvpicat Agronomy Tech 

Mem. No. 72, CSIRO Div. Tropical Crops and Pastures. 

Brisbane, Australia, 24pp 
McGregor. J. L .. Gordon, H. B . Wattlrson, I. G-, Dix. 

M. R. & Rotstayn. L. D. (1993) The CSIRO 9- level 

atmospheric circulation model. Te.(h. Pfiper 26, CSIRO Div, 

Atmos. Res. 89pp. 
MlMIK-OU. M.. Kouvqfqllos, Y. . Cavadias. G. & 

Vayianos, N, (1991) Regional hydrological effects of 

climate change. / Ihdrol. 123, 119-146." 
Nash. L. L & Gllick. P. U. (1991) Sensitivity of 

streamtlow in the Colorado Basin to climatic changes, find. 

125. 221-241. 
Panagoliua. D (1992) Impacts of GrSS-modellcd climate 

changes on catchment hydrology. Hydro!. Sci. J. 37. 141-163. 
Press. W H., Teukolsky. S.A , Vetterung. W. T. & 

FlaNNORY, B, R (1992) "Numerical Recipes in Fori RAN; 

The Art of Scientific Computing." 2nd ed- . (Cambridge 

Univ. Press. Cambridge.) 
Priestify, C. H. B. & Taylor, R. L (1972) On the 

assessment of surface heat flux and evaporation using large 

scale parameters. Mori- Weativer Rev. 100. 81-92. 
Richardson, C. W. & Wright, D. A. (1984) WGHN; A 

model for generating daily weather variables. Rept No. 

ARS-8. US. Dcpt. of Agriculture. Agricultural Res. 

Service, 83pp. 
Smith, M. (1991) "Report on the Expert Consultation on 

Procedures for Revision of FAO Guidelines for Prediction 

of Crop Watei Requirements." (L-and & Water Development 

Div. Food & Agricultural Organization of the United 

Nations, Rome). 
Wilks, D. S. (1992) Adapting stochastic weather generation 

algorithms for climate change studies. Climam Change 22. 



By Nick Harvey* & Tony BELPERiof 


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. 

hmi.MV'iioftx of ihr AW Jtown Mf & At*sl. \$9jh IWoi. & A 


by Nkk Harvev* & Tony BEiretunt 


H.arvpy, N, & Hi i ftfcWO, T. (1993) Implications 01 climate ejjang* for the South Australian coasilim- Tram, 
H W S. flfc* 0994 > UK(i). 15 52. 31 May. 1994. 

Recent attention has been focused on the effect of climatic change on sea level and the liflWtS i>l J r)M|(Q Wfl 
Icwl on dmsuiI environments. However, the variation in physical ajid geological processes which am responsible 
Tor sea level change is "I'cn ovcilowfced or underestimated. This paper presents recent geological ami ftde gauge 
.i.dii itoin South Australia to demonstrate that ncotcctonie and anihmpogcwe influences have rented in a general 
ivvctcstiniatujn of the current ntte of sen level rise, apftrl front two sites where the reveise is true. The pupei 
i.tineludes that all tide gauge .sites must be corrected lor vertical ciustal movements before any BCinclllfcilWtS arc 
drawn regarding local or global sea level cluuige. The implication* of this for South Australian pOWIS are that 
.liliutatnenis to sch level trends should be made before any vulnerability ahxexMucitts me conducted 

Krv Wowns: sea level change, coastal environments, climate change, vertical crustal movements. Soulh 


The recent tfteenuiouse debate has focused attention 
on the effects ofclimaiic change on sea level I Wat rick 
ft al. I993j and also The effects Of a rising sea level 
on coaxial environments {Bird 109}; Toolty & 
Jelgersma 1992). In addition* ihe work of the 
Intergovernmental Panel on Climate Change (IPCC) 
has produced scientific assessments of climatic chance 
( Houghton ?t al. 1991 . Houghton vt ol. 1992). together 
with the IPCC Common Methodology for assessing 
the vulnerability ol coastal areas in sea level use (IPCC 
199ft However, there has been some criticism ol the 
applicability of this IPCC Common Methodology to 
the Austtalian region (Kay er al 1902; Woodrofle & 
Mclean 1993 f 

Before discussing Ihe implications ot* this recent 
research tonne South Australian coast, il ti important 
lo note thai theie is u great variation in physical und 
geological pnx.vsses which are responsible lor sea level 
change. These variations, which have been categorised 
by Pugb (1993) in terms of their spatial and temporal 
inlluence. can vaiy considerably from short term wind 
^ftves with periods of about 10 seconds and an extent 
of tens of metres, up lo global changes in sea level 
related to sea floor spreading with time pcriuds or 
hundreds of millions of years. In addition, the process 
of redistribution of mass over the earth resulting from 
dcglaciution. addition of meltwater to the oceans and 
transgression and regression over the continental 
shelves, itself results in a variety Bf isostntie responses 
of the crust to the changing loads. The resultant lack 
of uniformity in global sea level change is often 
overlooked or underestimated. 

* Mawsun Graduate Centre for Cnvirrmmenial Studies. 

t'niwrsny ot Adelaide, Sntllll Australia 5005. 
i South Austiahan Department • «! Mines god ^ncrev. IM H.iV, 

I5t Eastwood. 8. Ausl. 5UM 

Perhaps the most studied geological period in terms 
■of sea level change is the Quaternary (Williams vt uL 
1993) where the effect of climatic change has resulted 
ut numerous fluctuations of sea level in response to 
the waxing and waning erf continental ice sheets. 
Consequently (here has been alternate Hooding and 
exposure ol continental margins together with periods 
nl erosion and sedimentation 

The most recent time when climatic conditions and 
sea level were similar to the present was daring the 
last imerglacial period art around 125,000 years before 
present (BP^ when sea level in the Australian region 
was between 2 and tf m higher than tivjay (Chnppcll 
1987). Since then* sea level lluetualions have always 
been lower than present with evidence from the 
Australian region of a low sea level of between 130 
and 165 m lower iChappcIl 1987) * 18,000 years BP, 
after which it rose al a rate of between 6 and 12 mm 
vr 1 prior to reaching its present level between 6000 
and 7000 years BP. 

It is in the record of the last 600** years titat scientists 
have focused attention on finding evidence for any long 
term trends in sea level either 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 indicate that subtle 
differences in sea level behaviour are the norm even 
for the Soulh Australian coast, long regarded us stable 
•and uniform (Lamheek & Nakada 1990). 

There has also been direct measurement of sea level 
and detailed analysis of recent tide gauge records to 
extrapolate relative trends (Gomitz 1993; Piraz/oli 
1991). As noted by Bird (1993) there are numerous 
laetors affecting relative sea level change Apart from 
eustatic sea level change there is the leetotitc response 
of the land and the isostaitc response of the continental 
margins relating to changing volumes of ice. water or 



sediment. In addition, human activities such as 
groundwater or hyjmcarhnn extraction land 
reclamation, artificial coastal ^ruelurcs, dredging anil 
pumping of sediment can affect local sea level change. 
Pira//nh (19S9) suggested that local secular tide gauge 
Jala are dominated h> neolectonic and anthropogenic 
effects, resulting in an over-estimation of gluM sea 
level rise by 2 in 3 (rmes when these (actors are ignored 

South Australian polio of coast protection 
and new development 

The current South Australian policy on coast 
protection ami new toastal development was prepared 
by the Coast Protection Board and endorsed by the 
Snuili Australian Government in May 1991. I he policy, 
whii h K described jn detail elsewhere (Coast 
Pioleetiun Bnaid $91}. relies m part on local records 
of coastal erosion, flooding and sea level rise but tnpit 
imporumly Kas incorporated the IPCC /Miniates ol 
greenhouse induced euslatie sea level rise. These 
estimates predict a veil level rise to the year 2UM) ol 
approximately 0.65 in (range 0.33 Itt to 1. 10 m) for a 
"business as usual" scenario (Houghton et tit. 1991). 
Given these estimates the Coast Protection Board used 
the "precautionary principle" in preparing its policy, 
The precautionary principle which was adopted by aJI 
Australian governments stales (hat "where there arc 
threats of serious ot irreversible environmental damage. 
lack of full scientific certainty should not he used as 
a reason lor postponing measures to prevent 
environmental degradation" i Intergovernmental 
Agreement on the environment 1992, para 3.5.1). 

In accordance with (his principle the Coast 
Protection Board has adopted the policy that any new 
coastal development should be capable of being 
reasonably protected from a I m sea level nse by the 
year 2100. The policy establishes the 100 year average 
return interval (ARfj water level as a standard rbr 
coastal development in South Antral ia. It recommends 
lHal site and building levels should he determined by 
Aiding 0.3 m to the 100 year ARI water level and 
(where appropriate) making an adjustment lor localised 
subsidence or uplift. Floor levels of buildings should 
be an additional 0.25 m above this leveL and buildings 
should not be approved unless they are capable nf being 
protected Of raised to withstand a furthci 0.7 m of scd 
level rise (e.g. by means of a bund wall). In the cose 
of flood protected sites, the calculation of the lOOyear 
ARI design Hood level must incorporale the extreme 
tide (plus surge) and storm water events, together with 
wave eftects wuhm the development. 

The poliey also makes & general recommendation 
lor an erosion setback distance. This is to he based 
upon 100 years ol erosion at a site, allowing for local 
uft&Ufcl processes and ti sea level rise of 3 hi to the 
year 7050, and taking account ol storm eiiMon liont 

a series of severe storms. For major coastal 
development u is suggested lhaL calculations .ire hasexl 
upon 2'H.i years of elusion. 

The policy is less specific about the protection oi 
existing property although it reaffirms an earlier 
Government poliey not to protect private property 
All hough pail of the Coast Protection Board's duties 
are to protect the coast, most coast protection woifcs 
arc carried by local councils. Accoidmg to the 1991 
policy statement, the Coast Protection Roar<j provides 
councils with giants of up lo 80$ ol the cost ot 
approved coast protection works and up to the same 
amount tor storrn damage repairs the issue of cost 
sharing between State and local government cs currently 
an issue of debate but the underlying question for any 
protection works is essentially a decision wheihci to 
poilcct or relocate. This i.% complicated by the level 
of public or private involvement and the relative 
responsibilities ol Slate and local government. K>r tlm 
reason the Coast Protection Hoard and local 
government determine issues on a case by case 

CiUibal sea level rise climates 

A major problem in identifying the current rate ol 
euslanc sea level chance from tide gauge data is I he 
influence of ncutectome. isostaiic and anthropogenic 
effects, compounded by a geographical bit*s in the 
distribution of reliable tide gauge data (Wai rick l'>93; 
Gornit? 1993: Aubrey & Emery '99i; Wondwortli 
1993). These problems create uncertainly Ml 
exuapolating the custatic component of sea level change 
and has led authors such as Aubrey & bmcry (1993) 
to expressed caution in attempting lo extrapolate actual 
sea level changes fi um ihe data. They suggest that the 
apparent post I930 accelerated sea level nse may be 
related In factors other than human induced factors such 
as a delayed response to climatic wuirmng, following 
the Little Ice Age. oeeanographre factors, or perhaps 
may nol even be statistically significant 

Oihvr auihors such as Oornit? |W9J) suggest rrut 
after extraction of long-term trends and data averaging, 
that it is possible lo obtain a true picture of sea level 
rise Gornitz presents evidence based on 16 tide gauge 
data studies u> suggest that estimates of global sea level 
rise over the last 100 years has been between 0-5 and 
3 mm yr 1 , with most estimates in the range of 1 to 
2 mm yr 1 iGorniu 1093), 

til addition lo studio attempting lo identify custatic 
sea level changes based on analysis ol tide gauge data, 
there are also II number nf studies on sea level rise 
projections related to climate change. The key study 
has been the IPCC *ea level rise predictions wiih u 
besi estimate of ji 0j65 m rise in the year 2100 
<Houyhtoti er ai. 1990) upon which the South 
Ausiialian coastal poliey has been based. The IPCC 



report provided a significant downwards revision of 
earlier sea leveJ rise predictions 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-caleulation (0.46 m by the year 2100, 
Wigley & Raper 1993). 

Hie South Australian coast: Seen far 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 mvnad 

of factors affecting relative sea level behaviour at each 
tide gauge site. Although there is a variation in the 
quality of tidal records available, the NTF analysis of 
tidal data from gauges with an acceptable datum 
stability indicate an Australian average of 
1,51 ±0.18 mm yr"' at the 95% confidence level 
(Mitchell 1991) This figure is in agreement with global 
analyses of 1.0-2.0 ram yr 1 (Gomitz 1993), 
notwithstanding the caution expressed about the 
validity of these figures (Aubrey & Emery 1993). 

South Australian sea level trends 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 I). 

Fig. 1. Location of Ode gauges in South Australia. 



Souih Australia - Neotectonic contribution 
to sea level change 

Australia is frequently, and incorrectly, quoted as 
a stable continent from which absolute sea levels can 
be measured. Neoieetonie movements caused by 
Structural geoidal. and isostalic pioccsses. together 
with factors such as sedimem compaction. affect 
Australia at various spatial and temporal scales. This 
complexity of underlying factors that control relative 
sea level change ha* bee" demonstrated by the 
reconstruction of palaeo-sea level histories- from 
numerous sites around Australia and South Australia. 

'I he sea reached its present level around the South 
Australian coast between 7<)00 and O(XM) yr HP. South 
Australia is in the "far field" in its response to global 
deglactahon. and is affected by subtle, ongoing tsostatic 
adjustment of shelf and coast. This is manifested as 
an apparent highsiand of the 6000 yr BP shoreline (Fig. 
2), the height of which vaties systematically and 
predictably around the coast In particular, the height 

RADIOCARBON YEARS ■ BP (thousand years) 

Fig. 2. Palaco *eu levels relative to present sea level* ;tt vm ions 
tites in South /\usirah;i 

of the fughstand increases up the two gulfs with 
increasing distance from the continental margin, Sea 
level change over the past few thousand years is 
dominated by this regression, which increases in 
magnitude from I m or less along Eyre Peninsula W 
3.0 m at the head of Gulf Si Vincent and 4.5 m in 
Upper Spencer Gulf. Isostatic adjustments thus vary 
from 01 mm yr 1 to 0.8 mm yr' averaged over the^e 
time scales. This has caused slow bul obvious coastal 
regression, particularly at the beads of both gulfs, 

Superimposed on this geographically variyhlc 
Holoccnc isosialie warping arc longer term tectonic 
movements, Tectonic effects arc most noticeable along 
the South Rast coastal plain, between Lake Alexandriua 
and Mi Gatnbier. where Quaternary volcanism has 
resulted hi ongoing uplift and tilting of the coastal plain 
The scale and variability of this upwarp can be 
illuslraled by the changing elevation of the lust 
imcrglacial shoreline (Fig. 3). This 125.000 year old 
shoreline rises progressively southwards, from 3 in 
above present sea level at Salt Creek, to in excess ol 
IS m near Port Macdonnell. Uplift rates in the Foti 
Macdonnell region are a minimum of 0.2 mm yr 1 if 
averaged our over this entire time period. 

Another under-rated effect associated with cilies is 
sediment compaction and land subsidence associated 
with coastal reclamation and withdrawal ot 
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 local record trom PUrt Adelaide elcorly 
illustrates these effects and the inherent danger of using 
tide gauge data without adequate ncolectouic correction 
(Belperto 1989. 1993). Data from tide gauge*. 
mangrove migration patterns and trom dated subsurface 
strata all indicate a contemporary relative rise in sua 
level within the Port Adelaide estuary. The geographu. 
restriction of these effects to the Port Adelaide region. 
together with preliminary geodetic evidence, indicate 
that the apparent rise in sea level is 3 kval 
phenomenon, resulting chiefly from subsidence of the 
land. Belpcrio (1993) concluded that up to 1.0 rri ot 
surficial compaction and land lowering hail occurred 
in association with wetland reclamation, acid sulphate 
soil development, increasing urban and industrial 
development and groundwater withdrawal- Highly 
variable rales of land subsidence, between 1.8 and 
10 mm yr 1 . were estimated to be occurring over 
different parts ot this regioti. More significantly, some 
three quarters of the secular rise of sea level indicated 
by the Port Adelaide and Outer Harbor tide gauge data 
could be attributed to land subsidence over the last 50 
vears at this location. 

Corrections to tide gauge derived sea level trend 
daw should he made tor these various ncotcctonic 



contributions. Preliminary corrections have been made 
for South Australian tide gauge sires incorporating 
known neolectonic variations (Table 1). The tide gauge 
data from Port Adelaide and Outer Harbor have been 
used in global and Australian sea level rise averages 
without adequate local neotcctonic correction. Many 
of the world's tide gauges are similarly biased by land 
subsidence effects, indicating the importance of making 
such local neotcctonic corrections to all tide gauge data 
before interring local or global sea level changes. 

Table \. Sea tff&t trends calculated from South Australian 
titli- £(tltg£ data. 


years of 




mm yr ' 

trend 1 
mm yr ' 

Port Adelaide (Outer Hh) 




Port Adelaide (inner Hh) 

.35. 1 



Fart Lincoln 




fori Macdonell 





Port Piric 

-0.20 : 


T haven a rd 




Victor Harbor 


I 17 


*Source: [Mitchell 1991. p. 355) except : Mitchell 1993. pere. 
comm.) 3 this paper. 

Vuftierabifiry ofihe South Australian (.oust 
to sea level rise 

A number erf papers presented at the Australian 
"Greenhouse 87 Conference? 1 discussed the general 
coastal impacts of a greenhouse sea level rise around 
the Australian coast (sec Pcarman 1985). However, 
very little work has been conducted on the vulnerability 
of the South Australian coast to current erosion 

processes, or on the effects of an accelerated sea level 
rise. Harvey (1993) provides data on the sensitivity of 
selected South Australian coastal environments to 
development. Fothcringham & Caton (1989) give a 
broad overview of potential impacts of a greenhouse 
sea level rise on the 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 including a variety of coastal landfbrms, 
including ciiftcd coasts, rocky outcrops, mangroves, 
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 se3 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 Tertiary 
limestone rocky coasts. The actual rate of chll or bluff 
retreat on these coasts will vary whh factors such as 
rock 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 movement, and 
sediment size variability. The high energy beaches of 

J 12 1 


J 10H 



L si 




2 -> 





l-ig. 3>. Changing elevation of the last interglacial shoreline (I250OO yr BP) in South Australia. 



the Younghushand and Sir Richard peninsula for 
example art: backed by an extensive dune coastal barrier 
system which would be vulnerable to increased storm 
attack wilh elevated sea levels. Coastline retreat and the 
development of dune blowouts is likely ro cause a 
migration nf ihe barrier ttrwaaK the Coomng, Av thi.% 
occurs underlying e-Meretc unci back barrier muds would 
become exposed causing variable rates of rvtreat In 
addjuon raised vvater levels will Impact on the Coorong. 

In contrast, [he metropolitan sandy coast lacks the 
extensive backing dune barriers of the vmdj-ea\t. Urban 
encroachment actus* the frontal dune togetlter wich 
exien.ove protective works have necessitaied a sand 
replenishment piogram to maintain tlie beaches. Elevated 
sea level* dre likely hi have greatest financial impact 
in this area where storm protection will need to be 
upgraded together with an increased sand rcplem.dimcnt 
programme, if the metropolitan beaches are lo be 
oiainramed. To the nonh of metropolitan Adelaide, tund 
subsidence has already been noted for die Port Adelaide 
diea. ElCVfitC<l sea levels will exacerbate the rate of 
relative sea level rise causing mangrove* to advance 
further intend and changes to ecological conations of 
rhc intcr-iidal and supra-tidal biota Jn some place* 
mangrove advance m,ry be restricted hy artificial 
embankments resulting in ntangriA'e die back. 

In »he guli regions, similar displacement. o( ecological 
communities such a* seagrasses, mangivivvs and 
samphrres would be pronounced along the low gradient 
Loasts. This rapid coastal retteat would be associated 
with reactivation of tidal swamps, localised flooding (^ 
the coastal plains and erosion of beach ridge >ysteois. 

In other parts of the coast, an elevated sea level is 
likely to cause flooding or low lying land, cnlaigemenr 
ut Luasial lakes and/«»r connection of some lakes to the 
sea. raised groundwater levels and alteration locsruarine 
emit onnicrtLs. South Australia has few cstuurics 
although there could be major implications foi urhan 
development adjacent to c&hjaries such as the 
Onkapannga and the Porl River In the case of the Port 
Rivei estuary, approximately 25% of the new by iirhon 
development is cm reutlv below high water, Elsewhere, 
potential impacts on the Murray River residual estuary 
umlicially constrained by die construction of barrages) 
may he less significant for urban areas hut ccmld have 
major implications Ibi the operating levels of the 
hnrr«igc.% and affect the management of the Murray 
Mouth and lower Murray Lakes region iHarvey I9SK) 

appropriate national response strategies and 
implementation requirements for coasts vulnerable in 
sea level rise. The Common Methodology lor assessing 
vulnerability comprises the following basic steps: 

1 Delineation of case study area and specification ol 
accelerated sea level rise and climate Change 
boundary conditions, 

2 Inventory of study aiea characteristics. 

9 Projection of relevant development factors, 

4 Assessment of physical changes und natural 
t espouses. 

5 Formulation (If response strategies and assessment 
o\ their costs and effects. 

6 The assessment of the vulnerability profile und 
inlerpreialion ol results. 

7 Identification of actions to develop | lonj> term 
coastal z^mc management plan. 

The assessment process* using these steps, is 
explained in detail together with tables and checklists 
oo assist ui die compdmion ol data in a consistent manner 
(IPCC 1991). 

The Common Methodology approach was found to 
be deficienr by Kay el id. (1992) in iheir Western 
Australian case study They suggest that there are 
problems in the biophysical structure of the assessment 
and also in the engineering dominated approach lo cost 
benefit response mechanisms They also indicate -that 
there is an essential step between the assessment of 
impacts and the formulation of a policy response to impacts, In Western Australia, the policy 
formulation includes inter- and inlra-govcimitenl liaison, 
public consultation, find political consultation, eventually 
leading to strategic coastaJ zone management. However, 
the fact that strategic coastal engineering decisions are 
part of wider State and regional planning issues, means 
that preference nviy be given to reactive coast protection 
strategics hti sliort term erosion problems rather than 
long-term coastal strategics related to sea level rise This 
type of prohtcin highlights some ni the ddficultics in 
developing a common approach strategy encompassing 
biophysical. admimMialoe and legislative factors iKay 
ct of #92}. 

In .South Auslrali.*, the 1PCC Common Melhtxlolopy 
has yet to be tested bun it is likely that bureaucratic and 
political problems are em rend) compounded In 
uncertainty with the fevieu of the 20 year old Cwk 
Pntii'dion Aft, debate over exist sharing tor coastal 
management between State and local "ovemntenj. 

In late 1991 the Coastal Zone Management Sub-gioup 
of the IPCC released its "Common Methodology" for 
the assessment of vulnerability of coastal areas tn sea 
level use (IPCC W9I) Anadvisotv group comprising 
the r - 'nited "Natrons Environment Program (lINIiP) and 
12 nations initialed a series of case studies to examine 


Appropriateness o/\«//ev<r $00} ^tspaUav priirv 

Initial estimates wf the current rue of sea level r **e 
obtained bv averaging ode gauge data from nround the 
world have been reducing. The principle reason i\ the 
recognition that hind level changes need to he renvivexl 



from the tide gauge dutffl before they can be used for 
ihiv purpose. Underlying, unrecognised neoicenmic 
effects remain the mam reason tor the geographic 
variability in secular tide gauge trends A gloM sea 
level rise cannot be expected to be detected uiKil 
adequate corrections arc made for these effects at each 
tide gfiUgt site. With consensus estimates of predicted 
sea level rise now down to 0,65 m over the next 100 
years, these neotectonJc effects will be determining 
factors in overall local sea level behaviour. In the gulfs, 
iiny rj$c of .sea level will be mitigated by ongoing 
isostaiic upwarp of up to OS mm yr 1 . In the 
South -Each tectonic uplift will be a mitigating 
circumstance In major towns, particularly where 
wetland reclamation has occurred, oi excessive 
groundwater withdrawal is taking place, any 
greenhouse sea level rise will be exacerbated by land 
.subsidence effects. 

In the absence o( accurate geodetic or uJnmcliic data, 
Ihc geologic record can he used to obtain first order 
estimates of land level changes- In South Australia, the 
coastal record has produced some useful results thai 
go some way towards explaining the variability itl yeti 
level change documented by ride gauges. However, 
there remains a pressing need tor local and globa' 
crustaJ scale geodetic control of all tide gauge sites 
In addition, a geographically suitable spread of tide 
gauge sites is required to account far the scale and 
variability of neotectonic processes. In South Austral ia. 
these lide gauge sites arc required within the gulfs as 
well as along Ihc oceanic coasdmc and ot (shore islands 

The implications pf tMs for South Australia's coastal 
policy arc twofold, hirst, it highlights the need for 
accurate local sea level data which are then corrected 
for ncoleetonic and anthropogenic factors. The 
preliminary corrected sea level trend figures presented 
in this paper indicate that the current rate of sea levc' 
ris<? in South Australia has generally been overestimated 
apart horn two sites where the reverse is true Second. 
Ihis South Australian case study provides support lot 

Piray/oifs concerns thai the global averages lor current 
cusutie sea level rise may also be overestimated- The 
compounding effect of uncorrected global sea level rise 
trends added to uncorrected local trend* will produce 
inaccuracies in the sea level trend data upon which the 
policies are based. 

In addition the uncertainties surrounding climate 
change predictions and 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 have major cost 
implications lor coastal development, it is unlikely that 
a greater precision foi climate chance models and scu 
level response will be reached in the near future. 
H < >we\ er, i i is possible to reduce some of the 
uncertainties of current sea level measurements by 
Correcting sea level data for neotectonic and 
anthropogenic influences. 


The data presented in this paper indicate the need 
for taking neotectonic and anthropogenic factors into 
account for calculating current rates of sea level rise. 
The secular trend obtained from tide gauge data is only 
a relative sea level trend miicc Ihc tide gauge cannot 
distinguish between a real rise of sea level and crustal 
.vubsidence at the site. This |\aper indicates ihal the 
current rate of sea level rise based on South Australia 
lide gauge data has generally been nvcresli mated a port 
from two sites where the reverse is true- li is apparent 
from this paper that fill tide gauge sites must he 
corrected for vertical crustal movements before any 
conclusions are drawn regarding local or global sea 
level change. The implications of tins foi South 
Australian coasts are thai adjustments to sea level trends 
should be made before any vulnerability assessments 
are conducted It may also be possible to revise policy 
guidelines based on the revised sea level trend data. 


AiiRfcrY. l> Ci & F>ifeRY. K. O. (ls>su> Rtfeenl global sea 
levels and land levels pp. 45-56 /// Warrick, R A. , Batnw, 
I- M & Wiglev, T M t. (Rn\> "Climate Jinit sen eve! 
<li«nge. observations, projections and implications^ 
(Cambridge tmversity Press, Cmnhiidge). 

Bili'fiuo. A P iNHSJ) The greenhouse debate and ruhffi sea 
levels: geological and oilier factors eontrollinu sea !cvel 
ebartgp pp. 77-80 U Dcndy. T. (Ed.) "Greenhorn* '88 
planning for climate change" (Department of Fnvironniem 
and Planning, South Australia). 

(IW) I -and subsidence and sea level use in Ihe IW\ 

Adelaide estuary; implications for monitoring the 
greenhouse effect. Auwalitin Journal oj h/tth Stirm r 40 

Biko. E. C. R 1 1993) "SiibmergVng coasts; the erteei* oi' at 

rising sea level on coastal environment:/ iJohn Wiley &, 

Sons. Chichester j. 
Cuasjai. Frotix i ion BoAKo(l992t Coastal erosion, ilmjchni; 

and sea level flue standards and nwneetioH policy Cna^lhu' 

Davis. G. H. (1987) Land sabytlence and sea level rise on 

Ihe Atlantic coast plain tjivir><ttrncat>tl Gtv/ntn *wtl Water 

Sdwtcv 10. 67-XO. 

Fothmunoham, D. & CArox. iW89) South Australian 
coastal lamlforms response to a greenhouse vea level rise 
pp. 77-Ht) In Dendy. T. <&tf.l 'Greenhouse '88; planning 
tor chmmc change" i Department of Jlmimnmcnt and 
Planning. South Australia!, 



Gornitz, V_ (1993) Mean sea level changes in the recent past 

pp. 25-44 In Warrick, R. A., Barrow. E. M. &. Wigley, 

T. M L. (Eds) 'Climate and sea level change: observations*. 

projections and implications"* (Cambridge University Press. 

H \ftVPY. N. (1988) Coastal management issues tor the mouth 

of the River Murray South Australia Cottstal Management 

16(2), 139-149. 
(1993) Sensitivity of South Australian environments 

W marina construction pp. 327-342. In Thomas. D. S G. 

& Allison, R, J. (Eds) "Landscape sensitivity" (John Wiley 

& Sons. Chichester). 


(1992) ''Climatic Change 1992. The supplementary report 
to the IPCC assessment" (Cambridge University Kress. 
Cam bridge) . 

Houghton. J. T.. Jenkins. G. J. & Ephraums, J. J, (1991) 
"Climatic Change. The IPCC scientific assessment" 
(Cambridge University Press, Cambridge) 

IPCC (1991) "Common melhodologv for assessing 
vulnerability to sea-levd rise" prepared by the IPCC Coastal 
Zone Management Subgroup (Ministry of Transport &. 
Public Works. The Hague. Netherlands), 

Kay, K. C, Eliot. 1. & Klem. G. (1992) 'Analysis of the 
IPCC sea-level rise vulnerability assessmeni methodology 
using Geographe Bay SW Western Australia as a case study 
in Loastal risk management" (Report to the Department of 
Arts. Sports. Environment and Territories) 

MrmtHLU B. (1991) Sea level and climate change, pp. 
327-342. Proceedings of the Second Australasian 
H>drographie Symposium Special Publication 27 
(Australian Hydrographic Society, Sydney). 

PeaRMAN. G. 1 (1988) ""Greenhouse '88 planning tor climate 
change" (CSIRO Division of Atmospheric Research. 

PIRA770) I, P. A. (1986) Secular Trends of relative sea level 
(rsl) changes indicated by tide-gauge records Journal of 
Coastal Research 1. 1-26. 

(1989) Preseni and near-future global sea level 

change, Poleogeogf'iphx, Pulroclhnutologx, Paleovcolngy 
(Ghbal ami Planetary Change) 75. 241-258. 

PVOH, D. 7- (1993) Improving sea level data pp. 57-71 hi 
Warrick, R. A.. Barrow, E. M. & Wigley, T. M. L. (Eds) 
"Climate and sea level change: observations, projections 
and implications" (Cambridge University Press. 

Toou-y, M. J. &. Jeegeksma. S. (1992) "Impacts of sea-level 
rise on European coastal lowlands" (Blackwell Publishers, 

Warrick, R, A. (1993) Climate and sea level change; a 
synlhesis pp. 3-21 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) 

. Barrow. E. M. & Wigley. T. M. L. (Eds) (19931 

"Climate and sea level change: observations, projections 
and implications' 1 (Cambridge University Press, 
Cambridge), f. M. L. & Rapdr, S. C. B. (1993) Future changes 
in global mean temperature and sea level pp. 111-135 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) . 

Williams, M. A. L. Dcnkerley, D. L Dr. Decker, P., 
Kershaw. A. P. & Stokes, T. (1993) "Quaternary 
environments" (Edward Arnold, London) 

Woodroefe, C. D. & McLean, R. K (1993) "Cocos 
(Keeling) Islands: vulnerability to sea level rise" 
(Department of Environmeni. Sport and Territories- 

WooDWOKi'H. P. L. (1993) Sea level changes pp. 379-391 In 
Warrick, R. A.. Barrow, E M. & Wigley, T M L. (Eds) 
"Climate and sea level change: observations, projections 
and implications 1 ' (Cambridge University Press. 

Wvsisfc. A. R. (1989) Implications for coastal erosion and 
flooding in South Australia pp. 77-80 In Dendy. T (Ed.) 
"Greenhouse '88: planning for climate change" (Department 
of Environment and Planning. South Australia). 



By Michael J. Tyler* 


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. 

Tramtutious t& thr Royal Society of S, Aust, (1944), 118(0. 53-57. 


by Michael I Tyler* 


Tyler, M .1. IW4) CI i ma lie change and its implications flu the amphibian fauna, Trans. R Sot: $ Ami. 118(1), 
53-57. 31 May. 1994 

The dependence of frogs upon moisture make* them highly sensitive to environmental conditions. The South 
Australian frog fauna includes 28 speues which collectively and individually experiences a wide temporal range 
of temperature and availahle moisture. Any climatic change involving warmer and moisler conditions is likely 
tu enhance their distribution and abundance 

Krv Words; Fro^s. moisture, distribution, South Australia. 


The relevance o\' the study of frogs many evaluation 
of the impacts of climatic change hinges upon 
recognition of the dependence of these animals upon 
moisture. Nevertheless ihe survival of frogs throughout 
the period lhat witnessed the entire evolution of ad 
olhct terrestrial vertebrates, demonstrates the capacity 
of frogs to survive massive environmental changes. 

The complete extent of the diversity ot the modern 
frog fauna is unknown: Duellman (1933) estimates a 
total of 3967 species at 3Lxii 9U and numerous species 
have heen described subsequently. In Australia the 
current total is 203, but many more await description. 

Despite Ihe success and longevity of frogs as an 
evolutionary lineage they remain dependent upon 
moisture because of the relative permeability of the 
skin and (in most species) the need to deposit eggs in 
free bodies of water. 

Il follows lhat frogs arc highly sensitive indicators 
of environmcnial pollution and, equally, that they 
contribute an early warning system in terms of 
detecting environmental changes. 

Any climatic change m South Australia at a regional 
oi total level is likely lo impact upon the distribution 
of species, and upon the number within this Stale. 

South Australian frog fauna 

Currently 28 species of frogs have been reported 
trom South Australia. The largest number of species 
in any area (11) is to be found in the lower southeast 
which is also the area of highest rainfall. However 
comparable numbers arc found in the arid northwest 
(9) and northcasr (10) (Table 1). 

The northwest and the northeast arc also the portions 
of South Australia where numerically and 
proportionately there axe the most species not shared 
with other areas (Table 1). In each instance these 
"unique" species arc known within South Australia 

•' Department of /oology. University of Adelaide, South 
Australia 5005. 

from fewer than six localities, and each species is more 
widely distributed 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 

T.J»mr I: Geographic <!ntrti<1cn\iii\ 

"j i' South Australian fw$ 

Numoc-r uf 

Geographic Area 






flinders Ramies 


F.vre Peninsula 


Yorke Penmsuia 


Ml Lofty Ranges 


Murray Vallev 


Kangaroo Island 


Lower southeast 


Nullarbor Plain 


Species Known from 
sronlmed to Ices than ua 
lhat area l«alitlfi 












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 latitudinal gradient. The one area in Australia in 
which this requirement is met is the Northern Territory, 
wherc there is a progressive reduction in annual rainfall 
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 


M. J. 1YI.LR 




:;:•.**:':•:•: i 







hig I Northern Jcrntory showing latitude^ isohyels and major centres, 









1 1 1 [ ' 1 1— 

1200 1000 AX> 600 50Q 400 300 200 

RflMfill (mm) 


Fig. 2 Number of species plotted against isohyets {r — .750) 

Tyler & Davies (1986) there is a distinct association 
between rainfall and the number Off 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 involves 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 Adehtus brevis). To examine any 
geographic trends in size I have used a median 






48 50 

Mean of Medians (S-V) 

Fig. J, 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). 
Ihe second between 12° and 26°S (r = .975). 


M I IYI.11R 

measurement, bcmg ihe midpoint between the- We of 
the smallest udull male and largest adult lemale, The 
first of tllcse analyses followed ihe standard 1° 
latitudinal divisions (Fig. 3). The second involved 
similar latitudinal divisions but separated at the 
intermediate O..S° divisions (Fig, 4). Essentially the 
results dre similar small species are associated with 
high levels ol moisture, and in arid areas larger species 




-• ' 





'.. .i'r, -.1 sa i'e * 

Mwy, ft) MfdiiW! 



I'lii 4, Number of species occurring within particular latitudes 
(boundaries at ha] (degree positions) plotted against median", 
of snout to vent length (S-V). Regression line r ?= _S»2t 

The overall lifestyle of frogs also is reflected along 
these north-south transects of decreasing available 
moisture. In terms of the size of the individuals there 
is a shift in the predominance of small species to larpsr 
ones associated with rainfall. Equally there is adi.stincl 
change in the overall lifestyle of these same species, 
such that the terrestrial/aquatic mode shifts from more 
than 50 r v in the extreme north of the Nonhern 
Territory to sew ul 2\° latitude south 

r.vtviing geographic trends in South Australia 

Within South Australia there is adinxt .ivsricTutinn 
between the number (and diversity) ot species and 
geographic area, but without the latitudinal decrease 
from the and north to the moist south. Instead there 
are pockets of species, each ol' which has its own 
unique features. 

For example any gradient is destroyed by the 
presence of a wet refuge like that of ihe Coongie Lakes 
system in the northeast of this State - an intrusion 
of un area wholly atypical within a broad /.oneot low 
and irregular rainfall. The Coongie Lakes Irog fauna 
is particularly rich, but not permanent because ol the 
period ol" drought. It has been suggested thai the 
northeast of the Stale presents a dynamic situation in 
which there is contraction and expansion ul populations 
according |0 the availability of moisture and repeated 

iransportatum of species from southern Queensland 
i "Tyler mi): Fig. 3). 

The River Murray presents a further dimension 
because it constitutes a route for the introduction of 
species from the cast, litoriu pmmi and Criuiu 
parin\i\>nifera enter the State b> this means, and are 
confined to its vicinity. 

The toadlet Psnidophrync ovadvnutlh has an 
extensive disiribulion in Western Australia, whereas 
in South Australia it has been found only at Victory 
Well in the Evcruxd Ranges in the northwest. The 
existence ot species within such restricted areas is ol 
significance to palacoclimatic interpretation. Became 
of the small geographic area to which it is confined, 
the dependancc of the species upon moisture, and the 
existence of more arid conditions around the area, it 
can be inferred that since the time of dispersal to that 
site, it has not been more arid there than at present. 

With the exception of Cnmu ripan'a which is wholly 
confined to the Flinders Ranges, every species in South 
Australia has a more extensive distribution beyond tin 
State boundaries, In fact the percentage of endcmism 
m South Australia is less than m any other Australian 

The implications of climatic change 

The climatic variables that influence ilic geographic 
disiribulion ot Australian species of frogs arc x^ry 
poorly known. Bntftstrom (1^70) was Ihe first In 
demonstrale an association in montane species between 
alhtudinal location and tolerance to temperature 
fluctuations. In particular he indicated that ihe species 
occupying the cooler habitats had the least capacily 
to adapt to a change in thermal regime. 

Unfortunately the climatic variables that have 
particular influence upon the spatial distribution ot 
frogs are, at besl, inferred ralher than demonstrated 
For example, what is the limiting factor of ihe 
distribution of the tree frog Litotiu rulullu which 
extends as far south as Wilpena Creek in the Flinders 


This species ranges across the entire northern hall 
ol the cotuineni, principally within the area of summer 
rains <Fig. 5). A commensal species, it is clearly 
capable of adapting to changing environments It can 
he assumed that a climatic change producing warmer 
and moister conditions will enhance the geographic 
range of /,. rubella. 

Perhaps the most significant influence of climatic 
change will be the creation ol more aquatic breeding 
sites and the persistence o\' these sites for longer 
periods. Together these taetors will modify habitats in 
a manner that is advantageous to frogs. Hence the 
Coongie Lakes are likely to become more persistent, 
and the fauna thete more stable in a temporal sense 
than at present. 


A Conclusion 

2 , ., 

Any climatic change that results in a moister and 

wanner 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 

I \y '/ '/]' V •• / x \ temperatures experienced already by South Australian 

V^ I / \ species, and the fact that a warmer and moist regime 

~/_ y% > /V will ameliorate the existing seasonally harsh 

I V /{y _ -'A environmental conditions. 

Vja rv v -^ / Acknowledgments 

V^,^^ I would like to express my deep gratitude to the 

> „ University of Adelaide and the South Australian 

V ""} Museum, for 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 

Fig. 5. Geographic distribution of Litoria rubella Fennel and Veronica Ward for preparing the figures. 


Brattstrom, B. H. (1970) Thermal acclimation in Australian In Tyler, M. J., Twidale, C. R., Davie-s, M. & Wells, C. 

amphibians. Camp. Biochem. Physiol. 35, 69-103. B. (Eds) "Natural History of the North East Deserts" (RoyaJ 

Dukllman, W. E. (1993) Amphibian species of the world; Society of South Australia, Adelaide). 

additions and corrections. Univ. Kansas Mas. Nat. Hist & DAVIES, M. (1986) "Frogs of the Northern 

Spec Publ. (21), 1-372. Territory". (Conservation Commission of the Northern 

Tyler. M. J. (1990) Biogcography. Chap. 20. pp. 223-226. Territory, Alice Springs). 


By Philip Stott 


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 is 
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 

J)wi>w:itt»n at M to$&ri Softrty oj'S. Aust (199-Ji. LISllt. $9-i& 


by Philip Srorr 


Sum. P. (1994) Uimuie change and its implications tor the terrestrial vt?r1ebiate Iuuhu (IW4). ftaiis. r? .W 1 . 
$ JteSl 118(1), 59-66. M May. 1994, 

A limited number |)f studies ot the biology of b variety of species ot terrestrial vertebrate* is used to speculate 
Ht*>ui iheu responds lo climate change us predicted by global circulation models. Dramatic changes in the distribution 
and abundance ol animals in Austodia that has already occurred consequent to European seUleiiieni h noted 
Speculation about the impact ol" climate change on the r^Uilivc abundance of mammals and reptiles, range changes 
of kanuaroo-., oibbtiv. in ai id areas, 1ood security of the Spimfcx Hopping Mouse, eompe iiiion between two species 
of skinks. and disease transmission is included. Nest-site selection by tortoises and social structure of tbxe.s are 
given as two examples where behavioural plasticity demonstrates some capacity to cope in xitu with me effects 
of climate change, but the ability ot must terrestrial vertebrates to track rapid climate change aerify* different 
NUhstraies is questioned. Reservations are expressed about the knowledge bnse upon which the speculations arc 
based. Cur wiy ttW species is fi sune of studies available to compare detailed data on distribution and abundance 
with climate over a wide geographical ranged backed up with biological information sufficient to explain the 
mechanisms by winch the species niterael with their environments, 

KES WORDS: Climate change. mammals, repnlt-s, Testudines. abundance, distribution, rainlall. thought. disease. 
MtitTffius ftt&msfus, Mai ropus fuliginoxut, OnctwQjmi amkvh^ 


Some 6000 years ago. within the span of 100 years, 
the Negeualion at Cold Water Cave. Iowa, changed from 
tvircs* to prairie (Oorale c/<//. 1992)- The changes were 
associated with a temperature rise of c ^°C, and with 
such a complete change in vegetation structure must 
have come a profound change in the vertebrate (anna 
ol the locality. It may be. during the next century, that 
similar dramatic changes in biota will result Iroman 
anthmpogenieully-eiihanecd "greenhouse" effect The 
terrestrial vertebrate fauna of Australia has already 
undergone profound changes as a consequence of 
fcuropean settlement (Reeher & Lun 1990); climate 
change would compound the impacts of introduced 
competitors and predators, destruction and 
fragmentation of habitat, altered fire regimes, hunting, 
and diseases. 

Realistic* predictions about the impact oi' eh mate 
chance on the distribution and abundance Of terrestrial 
vertebrate species require a foundation ot information 
from several sources. Essential are detailed forecasts 
Itom eltmaiologists; predictions about changes m the 
distribution, composition, and productivity or plant 
communities and other elements ol food webs, and 
detailed information about the present distribution, 
abundance, physiology, and ecology of vertebrates. The 
temporal scale of possible changes should also be 

* Uopi of frnvironraental Science and Ruinicland 
Management. Roseworthy Campus of the University of 
Adelaide. South Australia *>0('S 

The sentiments expressed in this paper arc 
complementary to those expressed in essays by Arnold 
(1988). Busby (1988). Graetz U9K8), Mam (1988), and 
Fossingham (1993). The paper augments previous 
contributions by exploring some examples of the 
mechanisms by which climate change might affeel 
vertebrates It is a speculative paper; ihe scenarios arc 
presented with the intent of illustrating the level of 
complexity, rather than making confident predictions 
about the outcome. The predictions 3re weakened 
because a single study of a lpCdl population over a short 
period of time is not necessarily representative of the 
biology oi a species during alL seasons and over the 
whole of its range (see Kemper ft G& 1987). and the 
range and habitat of a species as observed to date do 
not necessarily represent accurately the realised 
environmental niche of the species, nor this in turn its 
fundamental niche t,Possinghain 1993). In addition, the 
predictions of global climate models are not eonsrdered 
to be rcliahle at the regional level (Gordon ct at. 1992) 
As the number of logical steps increases, so die etrors 
are summed. 

The paper reviews those climatic predictions of 
particular importance to animals, uses some examples 
tu explore the mechanisms by which climate change 
might affect terrestrial vertebrates, thencxarnines the 
capacity of animals to cope with the predicted changes. 

Predicted climate changes 

Evidence from climate models indicates that 
emissions of "greenhouse" *ascs into the atmosphere 



will cause global warming of 0.3 (ranee 0.2-0.5) °C 
per delude, and associated changes in patterns of 
precipitation (Houghton**/ at, ls>92). Whilst the general 
predictions *>f the models are broadly accepted, the 
issue U complicated by the lack til clear observational 
evidence of changes attributable with certainty CO an 
rnhaneed "greenhouse" clfc'cl. On examining records 
of meteorological observations, Nicholls A: Lavery 
t1992t mill ItOI able to identify any clear trench Ln 
rainfall at reliable meteorological stations in .South 
Australia up until I9SX. Although Burtuws & Staples 
()9ylJ note a WflfRUCf trend in South Australia since I, ihcy cautioned thai the trciui was "Vlosc le the 
bounds of past experience". Models are not consistent 
in their predictions or seasonal changes in temperature 
patterns for South Australia (see Boer fl al 1992; Gales 
W «/. 1992); al the present stale of refinement of the 
general circulation models such uncertainties in liming, 
magnitude, and regional patterns are well recognised 
iHnii^tiuui el Hi 1902). 

Rainfall has a major influence on food supply for 
venehraie species, particularly in and uicas llaarstna 
el at. fl9*>3j predict a global increase in tropical 
disturbances. At present, some i -I tl tc^e distutbances 
extend into northern South Australia hs tropical- 
extrai topical cloudbands (TbCBs). and as a result 
heavy rain falls on an average of nine days per year 
tKuluiel 1990). However, the distribution and extern 
of the rainfall is erratic. Gordon rtal. (1993) cautiously 
report the results ol simulations which indicaic changes 
in daily rainfall intensity across Australia, particularly 
in summer, in the form of increases in the frequency 
of heavy <12.X-25.6mm. r. 31%: >25.6 mm. r, 95*) 
rainfall days, an c. \5% decrease in the frequency of 
light I <rt.4 nun i rainfall days, and a decrease in the 
number of ram days. Whetton etal. tin press j review 
the predictions of five global circulation models; lour 
predict increasing, summer rainfall over the whole of 
South Australia ull live predict increasing summer 
raintall over much of the far north). <ind four predict 
a decrease in winter raintall over most ol" South 

A weakness of the models is thai they do not take 
into account all major known influences on climate. The 
F.I Nino Southern Oscillation (FNSO; phenomenon is 
not coupled with the major jjiobql circulation models 
[Including the CS1R09 model) despite its influence on 
vaiiat'iluy of rainfall in Australia iWliertun <■; al. in 
prcsst fcNSO is of particular importance to die 
Kcoiiimcnt of many .arid zone species | Austin & 
Williams I9XH. Gractx el al. SWfe. The influence of 
ENSO needs io be eonsidered in addition lo chamres 
predicted a > accompany ihc enhanced greenhouse effect 
iWalkerer *//. I9&9) The issue is lurther complicated 
beetiUse HNSO itself may be affected, [f increased 
CO-, works to equalise lempcrauncs in the w&ttifiC of 

the eastern and western Pacific Ocean, ENSO would 
be weakened (Rind 1991) ihus tending to reduce 
elimaijt. variability in eastern Australia and couutennv 
to some extent the predictions outlined above 

Direct effects 

Direct effects of ei incite on vertebrates should Ik 
more readily elucidated than would oe the case with 
indireci effects. Two examples are presented ol ditect 
effects of climate on icptilcs. The lust is the process 
of temperature-dependent sex determination, and uV 
second is the influence of temper arurc on the 
distribution of icstudines (turtles and tortoises) Hoih 
examples demon.sltatc that the knowledge base is 
inadequate even in this more straightforward utccgnry. 

h'mi'i'ruiim-dvptndeni sex determination 

Temperature-dependent sex determination (TDSD) 
has been demonstrated in many species ol lesludmes 
leg Mrowsky etaL J984) f croenditiansfe.g. Webb 
a til 1983) ar.'ngamid (Charmer 1966 vide Bull 1980). 
und a eekkonid (Wagner 1980). Slight (<2°C) 
departures fium pivotal incubation lemperalures may 
result in eniircly male or entirely female sexual 
phenoiypes, even in some of the species which have 
heiemmorphic s?x chromosomes (Servan ff i\) WR9) 
Some species have two tha'sholds. with males abutting 
from intermediate ttnd females from extreme intubation 
temperatures (Ynlema 1976: Webb & Smith 19X4). 
TD.SD has been demonstrated to occur in an Australian 
crocodilian (Webb ft al. I9X.V). an Australian 
frcshwriter-'cstuarine tesmdinc fWebh er al I9K6), tutti 
Australian marine tcstudines iLimpus etat IWS). In 
contrast, it does not oeeui in several Australian 
trr-shwater tcstudines. including species found in South 
Australia (Georges I9«8;' Thompson I98«aj. 
Preliminary studies suggest that TDSD occurs in at 
least four Australian agamids (G. Johnston pfif3 
n»nm ). 

The influence ol environmental temperature on sex 
ratios is a mechanism by which climate change can 
directly affect the distribution of reptiles. Species with 
TDSD would be more susceptible to rapid climate 
change than species without TDSD Webb & Smith 
(1984) noted variation tn sex ratios of Crovothius 
jt'hmttwi halchlmgs in the field, and speculated thai 
it was due to the interaction hetween TDSD and 
geographic differences in mean air tcjupcrarures in 
ii warmer clim<uc. the effect might be to skew- the sex 
ratio completely, blocking reproduction, and thus 
leading lo the local extinction of the specres. Such a 
scctiarto has. been advanced by Pieau 11982) as a 
possible reason lor the cxtinclion ol many \1c\o/nic 



Distribution of w.\(ut(me>\ 

TcsUidines, which require external sources of heat 
for metabolic activity, are limited in latitudinal 
*Jistt ibuuon by temperature However, mean annual 
temperature is far ron crude a measure to predict limits 
EO disti iblltion. Lcnglh oi the growing pcfttfM fes been 
generally accepted 05 ft limiting factor, since al the 
latitudinal limits ol distribution adults may not have 
enough tunc ts) accumulate sufficient energy reserves 
|ii survive the winter (MncCultoch At Seeoy 1483). 
Alternatively, distribution may be limited by the ability 
riff hatch! ings tp survive overwintering m the nest 
I Breitenbach */«//. 19X4; Congdon ei at 1987). Ohburd 
& Brooks (1987) suggest thai a critical factor i> 
temperature durioy the maturation Of ova, which 
requires the accumulation of heal units over spring 
sufficient tor successful reproduction. [ suggest another 
possibilkv; that distribution i; limited by the probability 
of ambicni temperatures being sufficiently high during 
the nesting season VQ permit the energy expenditure 
necessary in dieting the nest ehamher. an activit*. 
reeoenisod as being energetically demanding (Con«dnu 
& Galicn m**). 

To predict the effect that climate change would have 
on a species requites detailed knowledge both of the 
culture of the change in climate and of the mechanisms 
by which climate change would affect a species tn 
lestudmcs. we need In know which ot the abovc 
hypotheses is acceptable before we know whether iq 
-.Moviine flumbci of days between threshold 
temperatures. nadir winter temperatures, number of 
spring days above sft temperature threshold, or the 
probability of occurrence in spring of spot tenineniUires 
anovc a tJite>hold. 

Indirect effects 

teftects mediated by other factors -ire inherently more 
complex than direct ^ffoets. Attempts at predicting 
changes m animal distributions bused on detailed 
analyses of complex mechanisms winch include 
consideration of matters such as physiology, population 
dynamics, inlet specific intei actions, hchavinurjl 
changes, and nucrohubitui conditions arc fraught with 
potential tor error. An alternative is to identity a 
Miuiller suite of influences which drive the system and 
determine the end result. N'ix 09H2) saw climate as 
Ihe major determinant of the distribution of terrestrial 
urgaiusms. and several authors have used various 
climatic indices to explain the diversity and abundance 
of particular Australian hiol3. Of particular value are 
those lafe studies which corn-pate detailed dan on 
distribution and abundance with climate over a wide 
geographical range Both mechanistic and deterministic 
explohirions |o]|«w, 

Relator ufmmhnce of mammals und rattle* 

And Australia alteady has a more diverse and 
abundant reptile fauna than arid areas in Noon 
America and Africa (Pianfca IMS?). In part, the 
diversity and abundance ^t the reptiles is attrihuled to 
the high variability or rainfall which isa feature nl the 
Australian inland fMorton & James 198$) Proposed 
changes in climate might therefore he expected to lead 
to an inctease in the diversity and abundance of repiilcs. 
relative to mammals. 

Not all mammals would necessarily be adversely 
affected. Fni the large arid areas of Australia, ncl 
annual productivity and hence the carrying capacity 
expressed as loial biomass or* venebrates is related to 
annual rainfall (Burhidge & McKcn/ic I9S9)_ 
However, the species composition ol the lolal biomass 
is largely determined by the predictability and 
distribution of the rainlall, rather than its amount. 
Patchy rainfall favours birds, bats, and mobile large 
mammals such as kangaronv (Burhidgc &. .McKen/ie 
1989) which ate physically capable of moving long 
distances to environments made favourable by receni 
heavy rains. Irregular rainfall kvuurs reptiles, which 
have very low field metabolic rales relative to mammals 
( Nagy 19871 and can survive for long periods without 
food (Morion & James |983i. A change in rainfall 
patterns to fewer days with rain, a lower probability 
of light rainfall, and an increase in the frequency of 
heavy rainfall would not have a great impact on mobile 
animals, but would favour reptiles over small 
mammals. Small mammalian eellulose-depctidcnc 
herbivores would be particularly disadvantaged; they 
are vulnerable because their energy intake iv limned 
by Ibeu cut si/e. This represents a similar proportion 
of the size of the individual as in larger animals, but 
the energy expenditure for maintamance oi body 
temperature must be relatively higher than for larger 
mammals which have a lower surface area lo body 
mass latio (Morton 1990V they are also limited by 
their restricted mobility in their ability to exploit a 
patchy environment; and they are most vulnerable to 
competition from rabbits (Burbulgc & McKvn/.ic 1989. 
Morton 1990). 

It could be argued that Ihe balance between the 
diversity o\ reptilian and mammalian species as al the 
time of European settlement was determined in some 
prior, more severe period of aridity (such periods :uv 
known from prehistoric tunes- see Singh I9KJ), and 
theieloic the balance would not be affccled by a lurifier 
increase in temperature and in the variability of rainfall. 
Further, any mammals which might have been affected 
are already extinct as a consequence of buropcan 
settlement. The counter argument is that effective 
aridity in the future may be more extreme than in recent 
evolutionary time. Climatic aridity t wherein increased 
evaporation is in excess of increased rainfall; may be 
compounded by "emulated aridity" because nt die 


? srorr 

consequences of the removal of primary productivity 
from the arid system in the tbrm of livestock and 
livestock products [BujfWdgO & McKcii/ic NK'J), and 
rhe sequestration within the arid system of primary 
productivity and nutrients in (he tissues of livestock 
and rabbits. Thus the resources available to native 
vertebrates would be sigmltcanlly diminished, 
particularly during the resource "bottlenecks" of 
droughts, and in drought refuges (see Mortem 1990). 

Range changes oj kangaroos 

The responses of kangaroos can perhaps be predicted 
with a I ill le more confidence than those ol other 
vertcbiates. Many studies of kangaroos have been 
undertaken, including thorough studies of their 
distributions (Fig. la) as part of the basis for managing 
populations which are commercially harvested. 

Caughley tt al. (IW7) have demonstrated that the 
distribution of three species of kangaroos is. m the 
major pan. determined by elttnaie. Win 1st 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 (Marmpus giganfcit\) and (he Western 
Grey Kangaroo (A/. JhliyjinoMt.s), are closely associated 
with the seasonality of rainfall; they overlap in areas 
nf uniform seasonality of rainfall, but M. i>igartifu.\ 
occurs in areas where summer rainfall predominates, 
and fa', faliginosas occurs in areas where winter rainfall 
predominates. The Eastern Grey tolerates higher 
seasonal temperatures than the Western Grey provided 

that there is summer rainfall. Both require a 
heterogenous habitat with shelter being an important 
component (see Htll 1981. Cannes tt al. WV1). The 
distributional data have been used by Walker (JWOt 
to develop an integrated modelling and mapping system 
which could be used to predict and map changes in 
distribution consequent to climatic change. Caughley 
ct al. 0987) suggest that climate change in the past 
bus influenced the distribution of mucropods: it is 
therefore reasonable to use their conclusions to predid 
the distributional responses of these three species* - lu 
future climate change. 

If. as predicted (above), the winter ramtall zone 
contracts to the south and temperatures rise, the 
distribution of the Western Grey Kangaroo would also 
contract to the south (Fig. lb). 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 South 
Australia, but these are minor projections into tins State 
of a distribution whose western boundary runs alony, 
or jum to the east of the States eastern borders 
(Caughley et al 1984). M. gigantcas could extend a 
considerable distance to the west of its present 
distribution, and hence across the north of South 
Australia to occupy suitable habitats in the northern 
pari ol the present range ol M\. il ceitutu 
conditions arc met. They are that summer rain becomes 
more common in norlhcrn South Australia, that reliable 
water is provided by increased frequency oi' heavy 
raudall and/or livestock water supplies, and that habitat 
heterogeneity persists m the face of climate Lhnnyr 

\J* 2-r/- 1 "-* 

M gigantet;s 

(a) Present distributions ol grey kangaroos 
(Density > 0,1 per square Km,) 

(b) Possible future distributions of grey kangaroos 

Pig, I mi Pre&eiv distributions of grey kangaroos. <bl Possible future distributions ol grey kanaguroos. Adapted bttmC&enCi 
ft tit. HWIt, Cdi.fchtcy & Ciriag (IvSlj, Cautihltry H at tN83). Cjnj-htcy ft at 0984), .Shoo vt nl (WK.4). 


BbWftl/ >>fnthhir< in tifitl mvm 

In and Australia, the Rabbit Or\vtotoi;u* 
i i^i/i i////y is decreasing die probability of survival of 
CTiiflM perennial plums during dnitighu and having a 
profound effect on trie reciuitmcnl of sonic spevtes, 
sin i idem in unic to eliminate them from foe landscape 
U.ailge it Graham $83: COWC 19K7) The response 
of the rabbit to elmrahJ change is therefore of particular 

Historical records show thai rubbiis in and areas have 
been severely reduced in lumbers during pasi dmughb 
(Griffin & Friedcl WN.V and may under divught 
conditions, become oxt'mei over large areas iMvcrs A 
barker r»75i. kceruitmont is most unlikely under 
draught conditions (King / af\ WN3). and prolonged 
dnnishls such us Ihc fciSCtf and J92.V38 droughts 
ai Alice Springs (Griffin Si Friedcl |VKS| km ihe 
h»n«eviiv of ihc species (see e.g. Dunsiuorc 1974). 
hven so. a lew rabbits survive in rehires The quality 
of the refuges is determined by (hen ability W harvest 
and siore water and nutrients flowing from larger arras 
of the landscape (Morton 1990) such that run-on horn 
ligln ramlall is sulficicnl to stinUiiute some plant growth 
during the drought penud (l.udwig 1987}. At Wilehitie. 
South Austrulia. Cooke fJ982i noted thai a sharp (all 
of liule more than Stum 01 tain might he sufficient to 
yield run-off which, if concentrated along drainage 
lines, would ensure ilutl succulent food in the form of 
chenopod shrubs would he available to rabbits living 
01 warrens along those drainage lines Once heavy raui 
lalls (>2G mm near Carnarvon, Western Australia. 
King tt ul, WH3). rabbits begin to breed, and cUn 
expand from the refuges to reccilonise the bulk fit the 

Heavy rainfalls Jre rare in the Austialiuu and /one. 
whole years may pass without a rainfall event ^ 
12.5 mm (Slaltord Smith & Morion WW). I.i^ht falls 
arc more common, hut it has been predicted (sec a>nei 
that the heavy falls would become more common, and 
light falls somewhat less eununon Hence, drought? 
ure likely to be shorter in duration, but the refuges 
which sustain the residual rabbit population during 
dnmghls would he k httle tes** reliable. With a 
coincident rise in lenipcraiure exacerbating the severe 
physiological stress experienced bv rabbits under 
present summer conditions iHayward 1061;. local 
extinction becomes more likely during droughts, hut. 
v\i|h decreased return times lor heavy ram. plagues be cxpvvicd more fi'cquenilv in those areas where 
rabbits survive 

Vegeurion changes consequent Id lite increased 
variability of precipitation might not favour rabhils. 
Rabbiis are found in chenopod sfU'liblands, but the 
majority of the teed is provided by Hie short grasses 
and Sorbs between the shrubs; Ihe chenopods an eaten 
only during droughts (Hall it «/. 1964; Gnfliu & 
Inedel WIS), Under conditions rif increased wlinialic 


variahihly perennial plants would DC favoured tf^st 
ephemeral plants (Stafford Smith &. Nonon 1990). so 
rabbit populations could be expected to become less 
dense unless palatable perennial gravies Mich & 
Ihettwda sp replaced annual uruvsoand lorbs. Further 
exploration ol this sccnai to would need to take account 
of a potentnil southern extension. i( the dominance oi 
Cj, over C^ grasses (Henderson c*f i>/ in pres<), the 
relative importance of C } and C grasses |o rabbils. 
and Ihe miplit'auons for rahbits of a change in the 
seasonal distribution of r<tin awards summer rainfall 
in arid area* of South Australia. 

J-oiiJ laurih jor Spinifcx Hopping M<uor 

.Vorowy* ii/<*v/V. the Spimfcx Hopping Mouse, is 
widely disinbttted throti^h sandy area-, ol uonhcrri 
South Australia, mainly in association with spinfffcx 
grasses (Walls & Aslin fVSll A major cornponent oi 
its diet ts seed (Finluyson 1940). and Irom the 
carboliydrate in seed it is able to denvc sufficient 
metabolic water to ^ur\ ive indefinitely; one female is 
known to have reared a single young without 
supplementary water (Havcisiock & NVatts W5) Hence 
ihe regularity of seed produciion would inOuencc 
suivivahiht) of ,V. tttefe in northern areas of the State- 
Seed produciion may be influenced b>' soil moisluivx 
lenipcraiure. and CCK levels, and two mechanisms by 
which N. tj/r.\is miglu be advantaged are exploied 

Soil n>oisture is one of Ihe mosl iiiipoitani 
climatically determined variables for grasslands 
[PiltOCfc L993) and he(K«- lor species such ns S. itkxis 
which depend in laree mcasuie on grasses (MucMillan 
Sl Lee 1969;. Walker *i al CWKM| anticipate that mean 
soil moisiure is likely to diminish in northern South 
Australia, although at present the reliability of 
predictions is questioned (Vinnikov W\\ Pillock 1993), 
More important for seed produeliou aie episodes of 
highei soil moisiure following heavier rainfalls, which 
arc predicted (above) lo become more rrcqucm. Thus 
pulses of seed .->uppl> may become p>ore frequent, and 
support denser populations ol .V. t.r/f.u>. 

The rate ol' ^nnvih and ihe speed at which seed 
development occurs following rains may be 
accelerated. Imai ti ul, (|985i observed increased Seed 
vield per plant for noe gmwn under enhanced 
greviihou&c conditions. Giflord (1970. I9K8) predicted 
thai wheat yields in areas with more strongly seasonal 
rainfall would increase as a lesult ol the enhanced 
"gieenuouse" effect, and that some grain growth would 
become possible under conditions of dtfidiiy which 
cuTrenily preclude any yield. The faelor influencing 
yjetd was stimulation to plant growth by boih incicased 
CCV. and warmei temperamres, which would result in 
a shorter grov* inc season such that the giam was more 
likely to he filling under a favourable soil moistuie 
regime. Ifnhanced efficiency 9f water use would also 
occur vlue to partial stomatal closure (Chaves Si Percim 


K Mori 

1992V Thus droughts as perceived by AA al&is may 
be ameliorated by the supply <>f some seed when none 
would be otherwise bt available. 
Australian arid-?one \oils are generally infertile 

(Morton i99to. find nutrient lltnlt&fl la) •.-ounter 

growth-stiniuUling mechanisms Although seed 
produciiou may be limned by the availability erf 
phosphorus, nitrogen is less hkcl) lo be limiting lot 
C. plants* hot a likely corollary is that the protein 
content of their seeds would be lower |see Conwy 
1992 !. A/, a/cu'v would not be disadvantaged, &1 it is 
more likely to survive drought on low protein diets 
which obviate the need to expend water to dispose of 
waste nitrogen (MacMillan & lee 1969)- 

( tMiftt'Hlion between Ctcnolus species 

Many of the 70 species in the seine id genus Cromr/cs 
9ft> associated with spinilex (Cogger 19*)?). and it is 
HOI unusual lor several species to be synu-pic, 
suggesting fine niche separation between Ihem. II 
eJimaie influence*, the niche sepatation. climate change 
may affect rhe hulancc between the species 

Chtunns heletmr and C. pantherinus are two species 
occurring synipatrically in the lar noiih-east of South 
Australia: Pianka (1969) noted that they shared similar 
niches, and suggested that C. pamhenm<s would be 
excluded by C helcnue but tor iu reproductive 
capacity, lames (J99ia) (bund a high degree of dietary 
overlap between the two species, and noted that dietary 
overlap in Qtnotits was highest during the driest period 
of his study. There is evidence to suggest that Ihe 
separation between these two species is raised on then 
thermal responses: Pianka (N8<S» found C. pantherinus 
to h;oe <i lower mean body temperature than C 
hefetuie. and .fames (1991 b) speculated thai C. 
pantherinus tand C. brooksi) can be active at winter 
Icinperaturcs which preclude activity by £ IwIvmic 
(and other Ctetwtus species), This permits C 
pantherinus and C hroftksi to begin reproduction 
earlier than the olbei species. Jf activity at different 
temperatures Is critical either for maintaining stable 
niche separation or tor sustaining a mechanism oi' 
oscillating disequilibrium between the specie*, an 
increase jn temperature during winter may /csull in 
competitive exclusion ol C. pantherinus by C hvlenae 
in those areas where I hey are sympalric 


In stable ecosystems, there is generally a significant 
level of accommodation between hosi population* and 
disease -causing agents, particularly if they have co- 
cvoJved. However, tiansnussion ol infectious disease 
is a dynamic process, and in many cases is dependent 
on ihe capacity of the infectious agent to survive outside 
ol tlic host. Helminth parasites oltcn have obi igatoiy 
larval stages which may sursive for long periods on 
the ground, and thus be susceptible to climatic 

influences. Should ihe donate change, the 
accomodation between the host and the paravilc may 
be disturbed- 

Amongst the parasites erf livestock there are examples 
ot species whose transtnissibility is known to be 
affected by climate. The larvae ot Hacmonehus 
tr>/uvv-wv, j gjAint-iniesiinal parasite of sheep, requiTc 
mean temperatures >|S°C for normal development, 
whereas the development C/fOtfertagfa circtuncincta 
larvae is suppressed above I5.5°C. As both require 
moisture, the former is an (organism of summer rainroll 
■uica-s. and the latter of winter rainfall aieas (Southcotl 
N al 1976), 

There is less known about the parasites of Australian 
narive vertebrates, and most of the published 
investigations have been Laxonomic (v g Reveridge & 
Durene-Dcsset I992|. Arundel et at (1990) undertook 
one or the lew epidemiological studies, which 
demonstrated that helminth parasites can cause 
considerable mortality in Eastern Grey Kangaroos, and 
concluded that development of frec-hvinc larvae ls 
influenced by climate In North America, the Moose 
Ahc.s ahe.s can C*.ist synipatncally with White-tailed 
Deer (klocoitens viryiniamts only in those areas when* 
circumstances do nor favour persistence ol infective 
larvae ol the. meningeal worm Purelaj)hostn)ni>\lu\ 
tenuis (Gilbert 19421. Hence, climate change may 
indirectly inlluencj the distributions of terrestrial 
vertebrates through its effect on the probability of 
disease transmission. 

Mechanisms for coping with elimatc change 

FVcssmghaoi (W*)3) recognised that (here are ihn-e 
means by which a speuo might survive dimute change 
range change to track shifting clirwitc ?oncs. 
tolerance of the change, and/or microevolutionary 
change F.xarnples are presented which demonstrate 
that lolerancv in the form ot behavioural plasticity may 
counter climate change, but tracking appears 
implausible for many small terrestrial species 


The Red Fox {Vulpcs wipes) has a complex social 
siruciure which can be modified to cope with 
environmental change. Zabel & Taggart (1989) have 
demonstrate*.! an effect by the FJ Nino phenomenon 
on the food supply ot a population of toxes on Round 
Island, Alaska. Increased water temperatures in the 
Bering Sea were associated with widespread nesting 
failure in the .seahjrd species which comprise most ot 
the summer diet of the foxex Resorption and 
prcimplanlauon Joss arc known to occur in pregnant 
vixens (Ryan 1976). a common cause of which ts 
nutritional stress (sec Moustgaard 1969). Hence, il the 
available tood was uniformly distributed amongst the 
foxes, total rcpttidnctive failure m the fox population 



may well have occurred However, on Round Island, 
iliftary change^ m smaller, less common, and less 
accessible seabird species were associated with changes 
in »hc social viruciure of the foxes. Polygyny, the 
trpioduetive mode prioi to the dietary change, wns 
supplanted by monogamy. The tnWS help i\ essential 
loi euptui uig and delivering prcv to a Incusing female 
and her liiler (Klennan 1977): thus ihe change in the 
social siructuic meant that assistance provided by The 
mule lit\ was 1ocu>sed on fewer cubs ai a time when 
it would have been more difficult tor the males 10 
pnxure SmkI Individual reproductive pujecess. (in terms 
of cubs reared to sexual maturity) of the reduced 
nuiilher of breeding lemalo was noi significantly 
affected hy the LJ Nino phenomenon. Hence, a 
llcmpurjry ) climate change which lead to total 
icpiuducnve failure of the two seabird speciev nmu 
prominent in the did. ol ihe foxes did not in turn lead 
to Ma! reproductive failure in the tbxes. 

The Eastern Long necked tortoise xChckulhuk 
l<mxnvlli*\ has simpler behavioural patterns than ihe 
Red Fox. hut still has \ome plaMiciiy. Ii appear*, to 
adjust its selection ol nesoug sites to Like account of 
meteorological parameters likely to affect incubation 
temperature. At Armidale. New South Wales, the 
species digs nesting chambers in unshaded areas, whicti 
increases insolation and hence egg lempeTaiurc, and 
sltonen.N iiicubaiion (Parmcntcr 1976). The same 
species at Rnsewonhy, S«»uth Australia, digs about two- 
ihinJs of its nests in sites shaded for more than half 
of the day (Sloll 1987, 1988). Nest temperatures were 
not recorded at either site, hut mean daily temperatures 
during the incubation period aie highet at Rosewoithv 
iX^°C higher in January) and cloud coveT is les> 
frequent (0.7 oktas less in January)- Thompson (!9SSbl 
has demonstrated thai unshaded nests of Etmlnut 
ftktcquarii at ftnrmcra* 5?outh Australia can he 2 h°C 
warmer than shaded nests, and attributed deaths in 
some unshaded nests to excessive heal. Thus it is 
reasonable to speculate that C /v>wv>o//ia. like the 
species of testudines considered Isy Bull et at. (I982J 
and Schwarzkopf & Brooks 11987). positions its nests 
relative in shade to ohuin opiimum subsuilace 
temneratuies fen incubation. 

Dorale ct til. (1992) dated laic Hiilotcne vcgcfctliou 
chances at two sites Li Towa which correspond in a 
rale of reireal ol prairie of 300-h00m pei annum. 
Huwevct. ihe anticipated t'atc of anthropogenic climate 
change is much greater than in the past (FVissutgham 
I993) Wiih the tow relief c>f the inland plains of 
m»nhern South Australia, mean thermal giadients are 
Slight, and a typical distance between isotbems 
corresponding to the predicted annual rare of 
temperature change of 0.03 °C is 20(H) m Because 
unnvy bird species oi the Australian arid /one are 

nomadic (VVVcns 199]|, tracking climate change is 
physically and behaviourally possible. However, 
>cdcntaty species may have behavioural difficulties 
Few datll are jvatUhle on the dispersal capability ol 
small lerrcstnal wrtcbrato, but the longest dispersal 
movement recorded by James (I99le) for any individual 
Of five species of Crctioms xvas 605m. indicating thai 
uuasMsted dispersal is most unlikel) lo be able lo track 
ehniaie change ai the predicted rate. 

Possingham (1993) points out that comparisons of 
dispersal capahilily with die rale of climate change may 
be simplistic. The comparison is useful to identify 
species which are physical ly incapable o1 tracking 
climate, but cannot hy itself determine competence. 
There must be suhH^uenl stages in the process ol 
identifying species at mk, such as consideration, of 
physical hanier> and iiitcr-relationships between 
species Even if Crwotus spp. were physically capable 
of tracking climate change, Ihcrc is a close and 
presumably obligatory association between many 
species of GwiorVM and Iriotlhi and PI vein;* line '-pp. 
(spinitex grasses) and their attendant termites. These 
are primarily dislTihuted in infertile, sandy soils 
(GraeU ft at. I9S&). which indicates that sprnifex- 
dependent species ol erenow* which me ievs tolerant 
oi increased temperatures would be unable to track 
climate change acmss elunges in soil renihty and type. 
1-or vertebrate*- in the north of South Australia, tracking 
temperature change> means a generally soudiern 
extension in range twith or without a northern 
contraction, a separate issue which would depend on 
the upper limits of tolerance), but for C pdMhttinm 
there would be constraints because long distance 
dispersal or even local spread of Tri(ial& and 
PteanuUiH' seem lit require considerable tunc (Jacobs 
B82J. Also, these grasses would not extend imo heavy 
clay soils and limestone plains, die laiici being 
geueially south of the present distribution of C 

The stccpnes5 of climatic gradient hi moui'.taittOUs 
areas is much greater than on plains, and thus 
alUtudinid tracking of climate elwngr. is much more 
feasible than latitudinal tracking tor small leiresLnal 
vertebrates. Generally, a short chmb in altitude 
corresponds to a maior shdl in latitude 'Peters A: 
IWluig 198?). Over a distance of about 15 km in die 
Adelaide I hlls, a 5G0m increase in altitude is associated 
with a fall in January mean maximum temperature of 
about 5°C. and a rise in mean annual rdinridl of about 
600 mm, However, whilst the climate as ntie 
component oi a species" environment may track up a 
mountain, other compuneiils of the e-nvmmmenl may 
be fixed; for example, substrate structure and futility 
usually change with altitude 

To allow Hacking requires the linking of aieas 
managed primarily tor conservation along latitudinal 
and alliludiual gradients (Norton I°u0t. Ihe teviesv 



or nature rexcrvex in souih-ensiern New South Wales hy Macules & Stein (#89) confirmed that 
a single, todgt mirmw. rectangular reserve aligned 
..'Ic-nu jn ultituiJinul gradient would be the configuration 
which would most patsinionionsi) ivieet ihe dominant 
environmental requirements ol temperature, rainfall, 
and substrain for 26 canopy tree species which occur 
in the region: Macke) ei at. (IsWX) note ihc necessity 
ttnd Hguc the validity of using vegetation daw us 
surrogates for data on fauna habitat in the present 
circumstances or paucity of the preferred primary daia. 
and advocate u Incus on ecological gradients in oaler 
i" poiviije a tnaryin flf safety in assessing areas for 
conservation value HosvtYer. because of Ihe low reliel 
of much ol South Ausirnlia. only latitudinal gradients 
arc po.ssihlc in most areas. 


The paper has loeused ono number of Minties which 
have relevance to the issue of climate change On 
reviewing Ihe topic, it is apparent ihax fhe zoological 
hase from which changes may be predicted is 
imperfect. Reliable information on the present 
distribution, abundance, population dynamics, and 

interspecific relationships of Australian vertebrates is 
limited (Norton 1990), but there is suflicienl 
information in indicate that climatic influences on the 
distribution or many animals operate through 
mechanisms which are subtle and as \ct poorly 
understood, and sufficient information 1o wurrnm a 
conclusion that climate change of any magnitude is 
quire likely to a fleet the distributions of many species 
ol terrestrial vertebrates. 

Conclusions about the fate of individual species arv 
at present speculative. Deterministic studies such as 
those undertaken on kangaroo peculations &xe less 
speculative than mechanistic studies hcvausc of the 
complexity of the means by which ehmalo influences 
the biota, but mechanistic examinations arc 
complementary in thut they nu\ rcveaJ critical aspects 
of detail not apparent to deterministic considerations 


Thanks are extended to Susan Canhcw who provided 
constructive criticism of the manuscript fti Megan 
Lewis who prepared lor presentation and publication 
the figures on the distribution of kangaroos, and to 
Kathleen Williams who cared tor the temple of the soul 


UAoi o. S W (19HXI Possible elteets rrt climate change on 
wildlife in Western Australia, pp. 375-386 tn 
fVarman, U. 1. iVA.) " Planning Ibrdimjte 
chnm>" (C'StKO. Melbourne) 

Awmji.i , H-. DrAirsiPK. K. I. Hakmioan, K. E.& Black, 
K. IW90) EpKletllidifTgU'i)) nhserwriions nn the helminth 

pnaaiEoti ol whvf&ohh c'\'"'"c/o Shaw m Viwcirta ftwt 

H-)AJ/_ fa 17, 3ft*. 
AlsTi.s, M P 3c Wiihams. O B (19XN) Influence of 

climale and community composition on the population 

demography i»l pastuic species in semi arid. Australia 

l&gctatkt 77. 43-49. 
R^vi.hsIock, ft k. A WAns, C H ( I97.*> ) Water balance of 

mi tall foliating todcnls I, Ad libitum watei intakes wd effects 

of water restriction on growth of young. Comp. Bitniwrn. 

P/m/of 50A. 8N-825. 
Brvrft.iDC,r_, I & Di'Rrni'-Di.ssl.'i, M C ()W2) A new 

speues of tnchustmnpvloKl nematode. Ottilia huinov. trom 

i native rodent, Rttttus fuscipes (Watcrhousel Irons. R. 

jm> \ -Utsi lift. I2M2R. 
BeHt. G. J ( . .V!rh,\Ki.ANt. N. A, & LA/Akh. M. OW; 

<irvt*nhoase s>as-md»Jced climate change simulated with the 

CCC second-eenctation General Circulation Model J. 

CIMnfli 5, 1045-1077. 
Rkiiri.vuACH, G L, Cosguon, J. D. &. SLLi R. C (rtHte) 

Wuitet tenipeniiurcs of Chrysrtny\ picta nests in Mictn^un 

Bf|0Ct<i on h*tchnmi survival titqiftttliiyjru 40, 7h Kl 
Btfll I j. 1 1980) Se* determination in reputes Q fov 3n>i 

55. 3-21 

. . Voot, R. C. & McCo\. C. J. (|U»2) Sex 

detenu mi mi icinpuiiiiores in turtles: A geographic 

vxmipinisun Fvnlutt*m .4ft 32M3J?- 
tiiHWimi, A A A. Mt K*-N/in ( N I- (I'JH9) Hiltcrnsni ihe 

modern decliro. of Wislern Australius venchr-iie launa; 

t.'.iUNCs and conservation imptiCJiions, fttot. Qttl\Qn< -50. 


Hi RMflM. K. R A Stai-i i ;;, M, .1. tm\) 'Trends in South 

AiiAir.'iti.'in tniiximunt Icmpctalurc records"" IS. A 

Department of nnvironment ami Planning, Adelaide) 
BlsliV J. K (IWI) HniL-liiu: n hioelitualiL -innlvsis :nu| 

prt'dietion svstern. Plant Pronrt. Quort A. 8-9. 
fMRNS S C 'Popm . A, R. & Gtuoc, G. C, HW> PftWlRl 

distributions and hahiut assoeiati«m> ot ked Kangar»o'-. 

Mtumptj\ tufits. and Western Grey Kanyiiioos, MtiCWpH* 

luli^in(>stis. in the South Australian pastoral ZfKift. Atm. 

Wildi, Res 18, 377 402 
CfcVGHfcEV. G.GRioo. G, C &SHOMI.I (1983) How mam 

kangarooi^ Ssattii (Sydney) 14, 151-152. 
__, Brown, B., Dosn.Nr,, P. & Giuo . U. U9«4j The 

gre> kangaroo oveiktp /one. AbfSl WtldL Res. 11. J-10 
SttoRT. J.. Ghioo. G. C & Ni\. H. (I987> 

Kanyanxis and climate; An analysis ol dislnhulion, / Attitit. 

Ecoi. 56, 751-761. 
Chakni£r ; M (1966) Action 4c la temperature no 1 la sex 

ratio che2 ttfrnbryoit iiA^unut itgamit 4A£amidac. 

L.accrtilien). So<\ Bwl, tie Lbmi Afhtnitt \M), 620-622. 
Ckavfv M, M. & Pur-ir-iUA, .1, S. M992) Water stiess, CO : . 

and climate change. J, r\p. Hot, 4.V IDI 1139 
CuotitK. H G. (1992) "Reptiles and amphibians ol 

Australia". 3rd Ldrn (Reed Books, Sydney). 
CnsiOOON, J D A GaTTPM. K F flWM Mitveinrnfs and 

euerj:etJe> ol* nesting Chr\scm\s pivto, fk'rptttt>tvHh'Cl 45, 

Bui-nrNiJACii, G. I„, .SiiLN, H C, A. Tinki i., D. W, 

(I0K7) Reproilneiion and iiesimg ei-ojogy i»l Snappirg 

Turtles (Cht'hiJra serpentina) in southeastern Michigan. 

/fttf 43. 39-S4, 
G<JNR(>y. J. (1^92) Intluenee a( elevated atmospheric CO 

iioneenti'aUoas oil plani nuiniuui. Au\t ./. Bt>t 40, 445-45K 
Coowr. B H (1982) A shortage ol 'water in natural pjOorev 

Ms a lector Inniluuj *i population "1* rahbits. OfwIvLtfRHl 

amiculus (U. \, in -irid, rmnri-eastem South Australia. Aus: 

WiML M,\, 9. 4<)5 476 



(19871 The rllvcV. ul rabbit grflTtAg on resene-aiion 

of ihcoaks, Alh'Citiitiirtnti n'rr;dtfmt<* uml saltwater ii-rrec> 
Kh'lali'tita ftuhHomrehitH. in theCourony National ftuK. 
South Australia rftfltf. / Btfl/ 13. U-CQt 

DoRv.r I A , Gon'/ai lis. 1.. A Ri-aoas M K..Hm.mii 
D. A.. MuRREl.t- M, T. A: Bam-.h. R. G. (T992) A high 
resolution record of Holwcuccliirate chain* in spelcoihem 
VtfMc from CVM Waref Cave, northeast Iowa .Scrrtnv 258, 

M NSvr.fci;, J S I|974| the mbbil m suoalpiiie souMl-eaMcin 
Australia I. Population Mruclurc and produciiMly. ,4uat. 
miiH Rr*. I- I ir>. 

Hnlavsaoj H , H ( 194ti) On the central Au«-Tnlian mammals 
I. The Tr*m\ R St» . S. im, 64, 125*136 

Oaif.s. W, {. , . MiTttiru., J. F. ( B(»kk. G. J., Ci.hasch, 0, 
8i Miiisiikm V P 13992) Climate modelling climate 
nrodillntn and model validation pp. 97-134 hi Houghton, 
1 T., Callander. B A & Variicv, S K. (tds> "Climate 
ehamje (442: The supplementary report to (he 1PCC 
Scientific A«c>snicnr (Cambridge Univcrsaly Pres>, 
Cambridge i 

r.h<m<.fs, A. I.I98X) Ses determination 14 independent o| 
hiLUhmion lempciaiiire m another Chclid turtle, OitioJtna 
tmfjiatftL t\>/vfa 1988. 24*-254. 

GtiniKD. R, M. (3V79) Giowth mid >icld ul CO,-cnnthed 
when undei water limited conditions, ,-lti.*/ J Ptanr 
Physiol- 6. 367-378 

(|9Mt Direct effecs of higher ojrbon dioxide 

L'onecmraiions on vegetation pp. 506- M9 to Pcarman. 0. 
I- (Cii ) "Greenhouse: Planning for clinute change 1 ' 
{CS1KM. Melbourne). 

Gn m ffl F. I, W9921 Retroduclivc logic and Hie Dffecli of 
meningeal wi'.rmv A comment ./ Witttt Mottop, 5ft. 

Gordon, H. B . WHernjN. P II Prime*. A B.. Fi/a*» i^i, 
A M £: HvuOtk, W K fffV?) Simulated ch,im.'es In 
daily rainfall imensrty due >.o the enhanced crccnhousc 
I'lfevi Implications thi extreme rainfall events Cfmutic 

Ommfst 8, H.vio:. 

Graft.*. R. P. Waikpr. ft. H & Wa.kfp. P A (P'H.m 
The e'lnsiujuenees ill chfinit change for wveni\ pcrvL'iii 
<■! AUsUuli;t. pp. .^9-420 hi Pcm nan. I ltd.) 
•Xj-ivcnhouse, Planning for clinuie change". (CSIRO, 

DRIf-r-i^. 0. K A FRIED©.; M. H. <W85i ObOOptliAOuS 
Lhanee in eemral AuMralu: Some implication* ol major 
ee-tliitfiuil esenN far land nianauement ./ Mul ffrlffriff 
ft 63-Wl, 

Ma^hsma. R .1 ( Mm inn. i F & SiiNOH . C A rlQW) 
Tropical diuurKwivsniaOCM. Climate Dxn- H, 24/-157; 

H.mi. G A.. SPETHT* R. I ^R>Rr/in.'C M i1%4) 
Re^cncralion of the vegctaiion mi Koouainore Vcticujlion 
Reseive. l92^-1962. Au\L J 8t>r, 12. 2n.S-264 

(■Kvwaho. I S. (Wf>l) Thi ( abiltiy trf (lie rabbil W Mirviw 
tondinons ol water resinction. CSIHO H'tldl, /u'i. 6, 
160 175. 

Ht'M'LKMi^, S.. HAI (hHM.tV . P., Vi in CAf.MMbKhK, 5. & 

Osmund, B. fin press) Are C 4 pathway plants threatened 
In glt»hnl climnie change? fa Hvopbysiolcipy nf 
photosvmliesis 1(K) Eeolo^'ieaJ studiex Heidelberg. Spnngcj 
Ve.-Lie Cited in PITTOCK, A B. ilWl. A climate chanet 
l>er>pective on urxssliindx }*Ttu XVtl hst Gn.Wii Cdltgr- 

Hill C. J. r!98L) OiVtrihutmn of the grey kangaroo* in 
south, inland Queensland. Ai*a. fo/nge. 1 3. 5R-<i6. 

H'»ouifit)N, .1. T.. Caiiasdlr, B. A & , S, K, 
HW2) *Climaie change 1992 The supplemeniiin IftpOQ 
U'the IK'C ScicrUiric Asses sin cnl". (Cambridp-. Uruversiiy 
?nt«. Cambridge). 

Imai. K.. Coleman. U K & Y^naoiswa. T (t9KY) Uiereiwc 

in atmospheric partial pre<*ure of carhtm diosidc «nd 

prowth and \ lekl of nee {Ory^t \ t jlna I., J ^t3/>. ../. tYt>/t 

5«rt. 54. 4I5^4LS. 
Jaivjus. S. W. (W831 RvliUiunships dismhuinni acid 

t-Mflution erf fhodta and PJMrmchtty tGraminae) pp 

2^7-1^0 //t Barker. Wi R. A Grcenslndti, P. l.iHds. 

"rivolution ol the Hon and lauru of arid AuslrahV. (Peacock 

Publishers. Adelaide*. 
,UMt:s, C- 1) <!991a) Tcmpoul vjrutiun itulieb ami trophic 

partitioning by coe\i c iinc lipids [devout* Sctneidae) u» 

central Australia. Otxtifogih 85. .*>53-y>l 
i|99ih) Annunl variation in reproductive eyclel o\' 

seincid lizards {Cu-noiuit m central Austmloi G>fwia 1991 


tl L '9lc) EVtpulalton dynamics, democruph>, (|ov] lift- 

lnsti>ry of <i_vmpatric seincid lizard*- lCuvicm>) in central 

Australia, iterpeiofaxica 41- 194-210 

KlMPPR. C M., Kirt'HI.NTK, D J., HoMHIRtV-S. W F-- 
Hmw, R. a... Bi<Uiit>. A l Hi ScitMfa. t, H. (1087) 
The ilenioaraphyjnd physiology of tWnrnsfp. (Rjcdenlia: 
Mtiridaet in the Mitchell dfx,\ Ko-nb^rky. Western 
A.uHtralia. J, Zool.. L/wd. 212. 535 562 

KlNr.,, D. R., Whi', S H . &. Scnstnri. G L (!i®31 
Population fluctuations and repatdueitoo i)| rjhhtlv in h 
pastoral arcs on the coast north of Carnarvon. WA, Aum. 
WikU. lies, ID. 97-104 

Kihiman, D G (1977) Monogamy in mammal.v Q Rev. 
Bhl. 52. 39-69, 

Kiiunm., I. (1990) Tropical-extratropical eloudband* m dte 
Aoslralian region, ttu J Chtttai 10, 341-364 

Uvnce. R. T. & Gkaham. C. R. (1983) Rabbits and the 
failure ol regeneration m Australian arid /one Arua/i Au&. 
I Fcol, K 377-3HI 

Ljmcus; C, i., Ri:u>, P. C & Milll*. I D. W985) 
Temperature dependent sex determination in Queenshmil 
soil turtles* Jntraspecific variation in Curetrti ttWi'ftii. pp 
541-^31 ft) Grigy;, G. T .Shine. R A, Hhirnom H- (Cdsi 
"Biology of Australian Fro«s and Reptiles*', fSurrcy Bcaity 
& Sons, Sydney) 

Li:nwiu, J. A- (1987) Primary productivity in arid lund> 
Vtvths ftnjj renhlles. J tr*/(/ ^"'"i/ 13. 1-7. 

NtActti loch, R. D. & Stcci , D. \1, 1 1983) Detnograptiy. 
growih, and food of West eni PdinM Turtle*. C7/rwtY»v.> 
pick' beilii (Gmv). from snuthcrn Saskaichewjin. CWw ./. 
Z/«./ 61 1449-1509 

Ma< KtV. B- G., Nt>. H. K, HlTCHINsoS. VI I- . 
MACIvUhUK, .1 R & 1'LtMiso. P, M. (T9K8i A^iessiny 
represenia)ivene\\ cif plat es fut cmiscrvaiton tt->ervation 
and heritage listing. Enytnm. Mrf/tagfi 12. 501-514 

MvcMu* fcNiX, R & I rr-. ^ K- (NtSV) Water nieyhidism 
of AuMnlian hopping mice Comp- Btochtm* Fktafytt, 28. 

Main, A. R. Il988i Climate change and its impact on nature 
con<c*r\aiton in Australia pp 361-374 ki Brarman. tk L 
(Ed.. i ^Greenhouse L*tnnoing iur diituio crMngc** (CSIRO. 

MARta i.r-s, C R & Stns, ). L. (19891 Patfems in the 
distributions of ipevies und |he>jelet.1ioti of naloie fe^erves 
An example from Eiuahprui, Ibresis in scnilh-eaiderri Ncv> 
StiiiTh Wale. Hm( O.mvr. SO, 2I9 23H 

Mimotos, .S. R. and Jamls. C. D. 0988' Tbe divurxity ami 
ahundancoof hvards in and Australia: A r^ippraisal . Am. 
N,u 132. 23? 25K 

(1990) The impact *.<f tuiopcan sctiirinc-it '«r. in. 
vertehrdle animaK ol ai jd Australia A coni-eptual model 
Pr<K. t'cvl. Sot: Ausl 16. 201-213. 

MorviGAARi). J. (1969) Nutritive rnlUlBfKBl upon 
reproduLtion, pp. 4R9 5lti M Cole. H H. & Cupps. P. T. 
(lids) (1969) "kepnuluciuin m domestic animals". 
(Academic Precis. New VorV). 



Mkonovnky N 5 L>t rrcttfc P H &\Vhitm(h<( C P. (IffiWI 
Sex ratios of two species ol sea lurtW nesting in Surinamc. 
dm .1 Z«>i hi, 2227 223V. 

Mvin-v;, K, & Pakm'K, B, S, 0975) A SWcfJ of me biology 
of the wild rabbit in climatically different regions in eastern 
Australia VI Changes in numbers and distribution related 
U> climate ami land systems in scmiarid notlh-weMern Nov 
South Wales, flfcfn WtUtL rV< 3, 

Nvo, K A fl$j87j I teld metabolic rule and fond 
requirement wiling in ummmalsand birds. &.*«\ Klotun-r 

N:r mil is, N. & I.avlky. U 0992) Australian lamfdll ircitds 
during the twentieth century. /w 7. CUmatnl U. 153-162 

M\. M. A [tfW2) Finviit.iiiiieiilrtl determinants • »t 
bmueographv and evoluoon in Terra Austrulis, In Barker. 
W R Ik Greensfade. P I iEJsi "'H'.oluLion utrhc Moid 
and laurta of" arid Australia" (Peacock Press. Adelaide). 

NonoN. T tl<WO) Mainlaminc hiohjpcu' diversity pp, 
■17 53 hi Norton, '1' , & Williams, Q 'bosl "Climate change 
in Australia: b.nvironrnetiial. mciocvonomic and political 
considerations" (Centre foi Resource and Fnvrronmental 
Studies, A.N.U, Canberra?. 

OnnARU, M. B. & Bfc&Ofc*. R J rlS)^7J Prediction ni uV 
unwet of the annual nesting season ol ihe dim n ion Snapping 
Turtle. Chvhttrrt Si-rfterififui //rr/i<V(f/fl($PU : 41 324-328. 

P\h\ir.MTFft. C J 0976) The udluiHl history ol rhe Auviah.n'i 
frrdiwatct turtle. ChvMimt hmwufti* Shaw 0>slndioitLt, 
Clielidac) PhD ihesis} of New hnuland 

Pfclfcks.. R. L, & DAPtiNO. J. I). (14X51 The gieenhouxe 
effect «nd nature le.sexve.s Biosnitto 35. 707-717. 

Punka, E. R. l ><>9) Svmpalrv ol desert li/ards " C /<vk»''I,m 
Hi Western Australia, kevhus 50. 1012-1030. 

(1985) Some intercontinental comparisons (il desert 

lizards. Mr/. Qfrijgfc AVv. I, 490-504. 
Pit At . C 1 >K2) Modalities of the action of temperature on 

sexual differentiation in field-developing cwbryos rtf 1h« 

Kniopcai'i Pond Turtle, £mvs orbtothtm (F.mvdidae). ,/, 

eyi. iftrf 2-20, 353-360 
Pint h k, A, B (l*W3) A elirnale change persretive on 

.masslands. Pruv, VI7/ fm Gmwl Ctmt>t. Hit IfrOfe) 
Pool., W P. (1V60) Breeding oi the wild ranbil. 

(/KXil&kMai vummhii (L, I *n relation (o the emoounietit. 

C5WW WW/. «r*. 5. JJ^Ui 
P(y;siNt,nxM, H. I* (feSOj linpaci ol* elexated aimosphene 
C*(^ uii hii>diver<it> : Mechanistic populniii>n-dvnaiiiie 
jjcispaiive iuti t Bet Jt 1121 

Kr» in I-:. II K& Lim L il^'iO) A review of cuireni fdciis 
ivf I he Efcltnotipii- '(/nscrVjliun ;md iiuiia^eniriil of 
AuM»ali:c. lcirestrilil seitcburtc fauna, ftw ^t-,-/ V(« 
Atf 16. 2M7-30I, 

Hivn. D (1991) Chtnme ViUTWbll!^ and cliinale change, pp. 
£9-7N }n Sclilesmger, M K ^Fd.) *tireenlnuwe-yjv 
imlaeed chmaltc ehan^c: A t nucat ol .simulations 
•iiid observations" tUlsevier. New ttufci. 

ttv.\N> U IE t I47bi Obscnmions on the tvpr-Klucbon and age 
structure oi the fnx. I4///'i v vn//tv L. in New South Wulcv 

am WihiL A'e.v. A H--20, 

Seinv-xk/knt-r, I . Hl Brooks. K J. {ViXl) Kest-sirc ficlOcfWH 
Jiid ttftspring ses ralio in Rtiitled Turtlek, Chcy^ wys j»u ta 
Gtfpmu 1V87, 53-61, " 

Stmr.. j\. ZakoknK! P l)ok|//i, M. Hi Pll-Al . C fWVj 
I cmale biased sex ralio in udults nf ihe inrtlt hnys 
fithituittns al the northern limit ot its disliibuuon in Fftmtc! 
A fjioV>able tonsetiuunct of the interaction fifWrffMrairjtt 
U'tlh licnoivpiu leu dctcruiiuuiion Can J &>*■•} 67. 
I27M I2S4 

Singh. G, (IVXt). I.ate Quatern.u^ pollen res'ords an.t 

seasonal pataeocliuiates ot Lake Frotne. South Australui 

ttKiUfhtoli^m 82, 419-430, 
SlMl'OKU Smiiii, L>, M. &. Mokion. 5 K- tl'JVO) A 

I'ramework for the eeolopv ol'atid Australia. 7. Arid htm*. 

18, 755- 27S. 
Siori, K (IVK/j Terrestrial movements of the freshwater 

tortoise Ou'I'kHw hfi^tVaSi^ Shavs us ntnnitoied with y 

spool tracking device. Mtst. Wiltll. jffi 14, 559-567 
(I9KK) Terrestrial inovenients of the freshwaier 

tortoise Cfwliklinn lon,i>itelli,s M Sc, thesis. Uni\ersity of 

Add Hide 
S<a mcott, W. H.. Maiok, fi, W. &. Bahocw, 1. A. (197m 

Seasonal pasture contamination and avadabllits t.f 

nematodes lor ynuine ^heep. Aw,r. J. Ay,rH\ Res 27. 


rn£)MP'.so\. M. B t IMX8m> liiflueiite of ineubation 

lempeiatute and wnler potential on sex deiermin.iiion m 

kmytlum mo i $(BtfXt\ ' I Test ud I ties : Pleti rodi r.j ) 

Hrrpt'ififtJ^ira 44. K6-90 
(!9SKb» Wed temperjrures in rhe Pleurodiran turtle. 

h'mwtuw nvtiifUftn'r V<ipvU> !9XK ( 906- 100(1 
VlNKIMA , K V il99|t .Soil moisture toiilcnl with ^lo(l,,l 

warmtiui. ppt 261-268 In Sthlesinger. M. E. <Fd,t 

"Trrvculioiise g;is-md|ico.l l I. unite chalice: A Ci'lLic*! 

appraisal ol sinmlalimiv and ubstMvaii..ins" (KUevier 

Amsterdam l 
WmiM'K. E. 1 1980) Temperature dependent sex 

iletL-niimnlioii iri/i^ukk. li/jr-J AppcitdiV /p,3ll to Bull, 

J. J tJsJKOi Sex deiertnuialiou in reptiles Q Hc\, tfi<>t, 55. 

Vr SLKtK. B H., Yoi \o, M. D.. Pakm.ow, J. S.. To. KS.K 

L>. Fumisx:.. K M.. W**G] u.s C R & I.ANie.JUKo. J. 

J. 11989) Global clirnatC change and Australia: Kfleein on 

renewable natural rey oiroek pp. 13^)3 /» Global climate 

elianye issues loi Australia Pipers presented ai the IW 

meeting bf ihe Prime Minister s Science C'ouncir 

tCoiiJHioiiweiiltli of Australia fajihertaj 
Wmkik. P A 0990) Modelling wildldedistrihuuons usinp 

a geographic information system: Kan&imos in relation m 

climate 7, Itiofrvx. 17, 279-289. 
Wads, C. H (W70) The (*)<>ds eatcn_by <nme Australian 

desert I'odents S Ait\t. Not 44. 71-/4 
& Asttv. 11 J. tt9»l) The rodents ol Australia 

tAnyus & Robe-oSon, Sydney,). 
WbHB. C. J,. Bno;wdfmi ( R a; MMsitit .is, S C_ (I9XH 

0»if,v/v/»,v tt.'tm\n.*m in ihe MeKmlay River. NT, VI 

Nesrtng btofogy- /l//\t. Wthjl. fa 10.' o07-d37. 
& Smiiu, A. M. i!9S4t Sev o»(io and survicurvhip 

in ihe Atislt Lilian IresliWfner crocodile CnttvtJ\itis tohlWiiflti 

Symp. :ooL Sue. Iwirf- 52. 319 355. 

CHOOUnM'J'T. O- ^t WHtTfJIFAn. P J t|9H(i) Ncstx, 

eggs, and embryonic developiuenl o] GUwttPGtWllS 
itiscttfftto (Chelonu: Cuiettochelidacl from northern 
Australia. 7. Zoai, . UvuL (Hi 1. 52L-5S11 

Win Hun. P. M.. Fowitw, A, M , Mvuuo;. M. R. 3t 
Pi i tot n , A. B. (in press) Impticaiions of climate ^han^e 
due to llic enhanced e.reciihnu'-c cltect Bd Hoods and 
dmugbiv in Austrjli.i Chniuh thntHNUt. 

Wiuvs. J A. (I99|) EcoloL'ieal simtlaiiiy ni shtub-deseo 
avilaunns ol Australia and Noiili America. FcpIuvv 72. 

YMF-MA, C L (1976) Elft^lsiil iiieniiatinii temjieraiure on 
v.-xual ditierciiiiaiion in die lurtle. Chvlvrint serpentina, 
1 Motph. L50, 453-462. 

Lmx. C J. & fAftftirr. .S. J. f|0«0i Shift hi Red PMi 
iH/pcn vutpes. mating system associated with CJ Nmo in 
the Bering Sea. Aftim. Behw. 38. 830 838. 



By Robert Boardman* 


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. 

rnmsactions ,tf the Howl ,W/,-fy "/£ *tffc <l l >94), liHOK ftMtt. 


bV ROBliR'i BuARDMaN* 


Hviskuman, R. T, ,IWt Sorra possible tffcltf ft clhiMtc- change on vegetation. Draft K W S Havf. JlSdr 
ISSMJI, 31 May, IW4 

One of the inmi difficult tasks for manager* of land which gruws nmg-Mved plant species, (specie* that dominate 
ecosystems), is how to assess that change in environmental conditions is actually ol si^nihemice 10 the presem 
ecosystems, and what to do to ensure that the ecosystem, rather than the species Themselves tan be maintained. things to do Which will make adaptation possible jre limited b)' the indefinite nature of nme frames. 
Areas of the Suit** north ot Kangaroo Island extending to the noilhcrn Stale boundary, ire the parts most sensitive 
tn changes in vegetation. 11 is likely that the nmsi strongly threatened species m the long-term arc Ihe perennial 
species which dominate woodland formations. Low scictophytl woodland and mallec fonnutions appear tu be 
particularly at risk. These formations occur til the /one with an average growing season, hi the best possible 
conditions, of from .V5 to 5.5 months. A IflTgC proportion of species in the semi arid and arid zones arc uee* 
and shiubs which are classed as "sensitive" by dendroelimaiotogists and dendioehirmoloeists. The first visual 
changes to vegetation will result from changes in physiological processes that are mediated ChrOQjiE plan' meiJinolism. 
Physiological processes of plants and microflora arc more sensitive to temperature than ecological processes, 
but this sensitivity Id temperature change will Iv observed through the symptoms which will be Mfttajpcfl in 
natuie and reflect the impacts through interactions within and between species- It should be possible to use 
tjutccotogtcat information to help track the pathways to new synecological combinations of species There is a 
need and an opportunity to identify "indicator species" and particular leatures fur measurement. Alternative 
approaches to the question of minimum water icqiiirvtneni tor a woodland without "gaps", and the extent ■ il Mingc 
covei in the m'envood and whole system, are reported. The optimum covet in the ovet vsood of 32±57c should 
be adequate to sustain low woodland on run-on sites in the 20(KHX) mm nonfall /one of S.A. The ecological 
significance pi Goyder's line and its potential usefulness are discussed. Features whtch permit adaptation include 
planting botanivally closely-related land races (provenances) and species, in five distinct croups of Encahptm 
that have been matched to three climatic models The aim has been to provide a neat-egg of genes, allowing 
adaptation lo ocvur whilst retaining ecosystem structures associated with particalai land-uses Provision Of gene 
banks and corridors of tend able lo pcrmii native species h ► migrate, are still ^een 10 be the best option tit the interim 

Ktv Wont>s: and /one, vegetaium. climate change impact. Goyder's line Fucahpnts- 

ltHfodncdon Climate change scenarios for the Australian region 

(Climate Impact Group 1992) are now less definite than 

South Australian vegeiaiion is strongly associated the original single emissions scenario (1PCC H90l 

wilh climates winch arc dominated by cool wet winters from which some of us worked in 1988 (Greenwood 

and warm to hen dry summers Rainfall and potential & Boaidman 1089; Boardrnan \W*) Global warming 

evapoiranspiration occui al levels which impart a range models are less simplistic and now include a range of 

from sub-humid to arid climate types The vegetation gases, responses in cloudiness, absorption of heal by 

has hcen classified by ils lorm (Specht W75; Boomsma the oceans and a degree of climate "Hcnshivky"', not 

ftLcwiv 1*376; Alias of South Australia 1986). These included previously "Ihe CSIRO GCM model has 

forms show marked correlation with isohyets and lines provided plausible ranges of IocjiI temperature wanning 

of equal potential evaporation. and changes in rainlall per decree of global warming 

A larve proportion of species- tn the semi-arid and overall. 

arid /ones arc trees and shrubs which ate clawed -Sub-regions have hetn added in the rainlall change 

as "sensitive*- by dendrocl.malolo-isls and scenarios which, so fat as S.A. vegemuoo is 

dcndtochronologists (Fritts 1976). Tree rings represent concerned, largely reflect writer findings. Areas ol the 

the end point of ihe lung-lived perennial plants Slate north of Kangaiw Island are placed in sub-region 

allocation o( carbohydrate resources poxlucxd by * ^ Clunale Change Group (WQ2) and this extends 

photosynthesis. This link emphasises the reliance of "lost « f *e way to the northern boundary. The 

semi-arid and and zone vegetation on o tolerable reduction in winter turn, despite a trend towards a 

infreiiuenev of favourable growing seasons when ^cater proportion of annual nunM in summer 

conditions cnahle species Lo maintain their place in together with a marked rise in potential 

ecosystems of the region. evapoiranspiration has been modelled, and lejds to 

greater droughiinc^s and diminution of growing season 

* Department of Prfmarv Industries, SA. Forestry. 135 described for Ihis port of Ihe Slate (ApjnMidix 1) 

Waymouth Street, Adelaide. South Australia 5000 (Boardman W9R l Q 92|. 



Artus of the State souih ot FIcuricu Peninsula art* 
placed in sub-region R: (his is a sub-region unlikely 
hi suffer marked change in gjttes rainfall and may nol 
even see a decrease m winter rainfall- This fus in with 
earlier progni^cx already deduced for this partol ihe 
Sltei Impacts on vegetation seem likely to be mlrumul 
and povsibly will see a more favourable sei o( "growing 
conditions. DendnmieUicul dala on nalive forest in the 
I*jwcr South-east (Ruitcr 1964) indicate that the nalive 
tree species arc better adapted to a summer rainfall 
climate lhan the Mediterranean type which has 
persisted since the last glacial epochs That is. most 
of their evolutionary adaptation look place in a climate 
similar to that now present m the eastern pan of NSW 
(Board man I9jj6), 

The impact of ENSO {El Nino - Southern 
Oscillation} hits not been included in GCM models 
Thi* appears only Ui affect Mtb-region A of the Stale 
which is bisected by it (Allen 1988) The boundary 
between ENSO affected areas and those dominated by 
the Antarctic polar circulation systems lies along a NW 
Sh direction which crosses the northern Hmdets 
Ranges. The 'Settled areas" of the State SW of ihc 
boundary /one are not directly affected by liNSO. and 
only indirectly influenced by the alternate system, 
called La Nina. 

I< is the time-scale of changes rather than their degree 
which is different, compared v* ilh the early model, and 
il has heen extended. A range of temperature change 
is now given miner than an average. However the 
median lemper.iture change (or the ?070 scenarios is 
similar to that used in r$88L and these will be tised 
specifically here. Physiological processes ot plants and 
nuemflnra are marc sensitive in tempc-fulute ihan 
ecological processes, bui ibis sensitivily in Leroperaiure 
change will be observed through the symptoms which 
will he ecological in nanire -und Tcilcct the impacts 
through interactions within and between species. They 
will be the first changes fcv attract attention We expect 
to find features which can be measured and which will 
indicate the nature nl" change^ taking place. 

What is the nature of changes 
that may b* expected? 

The scquencv of reviews adopted brv the organisers 
of this meeting implies that the mosi obvious etlccis 
ol climale change indicated tor South Australia will 
be noticed in soils (which f think is unlikely! and in 
ftf£8£ we recognise as wetlands: namely, low coastlines 
and coastal swamps, swales between sand-dunes, lakes 
and lunettes. Most indications of the impact of seasonal 
changes in rainfall distribution, both in quantity and 
intensity, and a warmer climate indicate that it will be 
topographically low-lying areas which will be altected 
early during (he process ot change It seems reasonable 
KO suggest (hat it wilt he the vegetation in nreas nios'l 

sensitive to direei and intense climate change which 
shows the visible signs first rather than the animals 
and Ihc soils. However, it may be tempting to suggest 
that il will be the microflora and microlaunii whkh 
will be most sensitive to climale change until is it is 
realised (hot both JrC more closely linked to the 
dominant higher plant species ot an ecosystem ami tin 
weather, than to climate i Howard l967;Lewin. 1985: 
Simpson l%7>. 

II appears likely thai lire first visual changes l>> 
vcgciaiion will result from changes in physiological 
processes that are mediated rh rough plani metabolism 
In this respect, the tallest elements )0 ecosystems. 
especially trees, arc likely to show the ef feels lust. 
Trees are more likely 10 suffer donate change effects 
lhan shrubs, (he result ol their abilily to pioduee a bole, 
or trunk. As Ihe bole lengthens and its diameter 
becomes brgci. a greater amount of tissue Heeds 
sustenance which is unnecessary forlcwr plants. A 
sheath ot new tissue is added each growing season 
comprising extra phloem, or husi, Kark ±nA mensem 
tissue, the cambium It is a cost Ini keeping the tree 
in a lofty position in ihc ecosystem There is no 
difficulty with maintaining ihis structure in a stable 
climate: allocation ol resources is attuned to the 
situation: wood growth is laid down ill a predictable 
fashion including well into old age VVuring |I9S8) and 
franklin vt ol. f 1988 1 have di%cussed the nature ol 
changes which may result in tree death. There i a 
complex series ol allcmatc pathways which lead to a 
single result. The first of these is a reduction in food 
M.ibsionce available for new wood production. 

Different environmental stresses affect the major 
components of the resource -budget allocation. There 
is a hierarchy Rf normal allocation of citrbohydralt* 
with a live, Stem growth only occurs once the tesouiec 
demands of new leaves, new roots and an intrinsically 
regulated reveisc allocation into storage (*'recuvciy 
insurance") has been accommodated. Hence, the time 
when physiological stale is most suitable for assessment 
is just before a flush in new leaves becomes apparent. 
Flowers are ni>i produced annually ami when they do 
depends on the amount of stored resources 
accumulated. At this lime carbohydrate and nutriem 
reserves in iwigs, older foliage, large diamcicr roois 
and ihc stem itself are at their greatest. The arbiter 
of this is the amount of new wood produced per unit 
of foliage 

Ft ills (1v*76| has produced model diagrams which 
illustrate the factor* which interact and lead Ui 
formation of li narrow grmvth ring in the trunk Hi. 
model has twv parts which can be related to the S.A. 
situalion. currently and as it may develop over ihc next 
century. Pan A of his model contains relationship-. 
associated with low precipitation and high temperature 
<7//rwg the #n)win# \ctmort that lead lo ihc formation 
of a narrow ring in irees on dry sues. Pan B is a 



(H.s>jf)le precursory extension to part A . time* when 
low p recipiiation and hii.ii temperature oecur/?ryo'" W 
thf <t.'h'//;,ij s?us<m and mid to xht* amount ol stress. 
The complexity of these rea* (ions is indicted by ihe 
two parts which each have 24 possible stupes. When 
bold patty oicct thev aei additoely and directly on the 
cambium, bul only combine turlheJast three Mages 
Combination of pans A ami B is not an uncommon 
situation in S.A. when dry winter precedes a droughty 
growing season The net result is reduced rales ol cell 
division in die cambium and so fewer wood jells 
become differentiated, as' a consequence <ff low 
production of growth regulatory substances (plant 
hormones! and Ibod resource. Several other common 
factors affect the cambium when a narrow ring is 
formed including years uf intense flower and seed 
production (mast years) and attack by pests, mninly 
defoliators bul also including hark beetles and sapwood 
borer*. DeudiiKlmmologistshave attemplcd to identify 
the range of causes when a narrow ring is farmed and 
some models exist for these situations (Fritts i t t%), 

fthtcoccologkal plant Studies have shown i>cles ol 
change have alicndy been experienced by the gcueia 
and species still present in Australia. Species may be 
adapied 10 the scenario in different ways or ma\ be 
tolerant tn different decree*. A sustained trend away 
from not nudity a, ill have an effect proportional to any 
dormiint evolutionary adaptation already incorporated 
imo iht genes. Wc hnsc no ready way uf knowing what 
these mifctfu. be in specific ca*e> This position is 
aggravated by the apparent lag ol many centuries Ivfore 
consequent vegetation changes occurred hi lire past. 
The current climate change scenarios preclude this 
significant lag period apparent in the past- It a trend 
in the raie of change Lit climate becomes apparent then 
the stress impacts are likely to accumulate m ways not 
seen before. The impact is then likely to be Similar 
to one oi'n range of possible impacts such as illustrated 
hv Barker ft989] looking at different recurrent periods 
between fires in At'tnia woodland. 

Periods of extended drought arc not particularly 
harmful if tree* have well-established mot system*- and 
foliage canopies, partly because drought is reckoned 
in relative terms to the normal climate. Where there 
has been natural election by dnnighi, trees rarely die 
frnrn drought. In the short term, adaptation may help 
retain reserves by temporary reduced demand Trees, 
such as those in S.A. are probably more acclimatised 
10 chrome drought than most high forest liee species. 
They already have features which reduce the impact 
fit drought: low leaf area indices » leaf area per unit 
of ground area), smaller, thicket leaves, stfap-likc 
petioles leading to panpbolomcTrie idroopmgi leaf 
posture; leal oils and Wax coating. Some species 
readily drop older foliage ro reduce leaf area. The 
adjustments can mean thai wood pioduction is 
computable- with thai on more mellow sites hecause 

respiration load is less A considerable vegetation 
inenta occurs with existing, mature tree-dominated 
ecosystems heeause Irees influence microclimate .SO 
significantly, especially in the availability ol sunlight 
|)|)tj t- »p-Hiil mouture They can also alicr the character 
of sptl ehemisUy as Lhcv SEOffi and retransloeate scarce 
nutrients in tissues awjry from recycling. 

Consideration of mienKlimaies emphasises that the 
regeneration phase, when seedlings become 
established, is possibly the moM ctitical phase ill the 
life-cycle of higher plants The game is one of numbers 
and nf resource reserves in the seed. Most species in 
S.A. hayc adapted so that they produce large numbers 
uf seeds with .*mull or negligible food reserves, so 
regeneration becomes a maner of numbers Vege- 
utional change associated with climate change will 
depend therefore, on the reproductive vigour of invasive 
and endemic specie* and whether mkfoebmates. and 
provision of soil conditions are equally-suited to the 
potential invader 

Persistent drought however, halls photosynthesis. 
Jeads in depletion of carbohydtate teserves, and 
depletion of defensive compounds. Individual species 
within a plant community will vary in the rate of 
reaction, Air turbulence and wind shear, especially ol 
siiong. hot. dry winds, as well as chilling winds will 
aggravate drought and damage loliuge, taller trees are 
more at risk than short ones. ShaJlow-rooicd trees 
growing over stony materials allocate more resources 
to root growth than to shoots. t-ven so. they ate 
susceptible to infrequent intense drought and arc liable 
to die oil in whole groups. Damght will inhibit 
microbial activity and reduce mincrahsaliuii of 
nutrients from liner; higher tempcrarurcs in times of 
ram will accelerate mineralisation rates hut also 
accelerate leaching losses - both conditions in a 
changed climate will limit the quantity ol nutrienls 
available. Nearly any stress, if it becomes sustained. 
leads to forest and woodland decline; it reduces Icaly 
canopy, photosytuhetic activity, stored reserves and 
delcnsive factors throughout trees mote so than in 
lesser vegetation. Consequently the balance between 
species and individuals will alter and. in ecological 
terms, a sc/al change will be initialed. 

Scientific investigation.** since 1988 into impacts of 
laciurs in the climate change situation on vegetation 
mainly have been concentrated upon the effect of elev- 
ated atmosphenc carbon dioxide concentrations on 
physiological and meiabolie processes. This has been 
discussed frequently as a "fertilizer effect" and evidence 
in the Held as well aa numerous laboratory studies have 
been made lo adduce its impact. The subject wa.% re- 
viewed in Australia by the Ecological Society and 
Socictv of Plant Phvsiologisis iWM2. I*W; Gilford 

The impact on biodiversity has beca given much less 
attention. It has been reviewed by Possingham (1M93> 



at the species level in terrestrial ecosystems. He took 
a mechanism, appmaeh la an implied rapidly-changing 
physical environment and concluded thai, in the short- 
term, extinctions will occur ihmugb direct interactions 
of species with changes in the climate til" Iheir 
environment. A second set Of species face extinction 
either because ol opportunities availahle to disease- 
causing species or to loss uf mutual support (eg. loss 
of shade ) W unsustainable symbiotic relationships 
Patucular "key: *lone" species m the association may 
become extinct and so change the essence of that 
system. Direct inrei -specific competition is considered 
to be the least likely cause of species loss Irojn a 
present-day community. FVissingham (1993) argue* that 
each species will be affected by one. or any 
combination 8! Jour basic pathways, change in 
geographical position to follow favourable eltmartlcally- 
cnrtfntlled features uf the current range; .select inn of 
tolerance to climate change withitnhe current /onaiton. 
niiao-cvoliioonitn change, and extinction He suggests 
lliat ir should be possible to use auteeolngtcal 
infoimation to help truck the pathways 10 new 
^yiioeoIit£ieaJ combinations ol species. This supports 
the idea ihat there is j need ami an opportunity hi 
Identify 'indicator species' and partieulai features of 
these for measurement 

South Anstralian vegetation 
in Ibt bit 20th < enlurv 

The foregoing discussion has described gCQCrtf] 

trends and effects. Once tine wishes to review the local 
situation, one has to observe its irregularities. The 
Flinders Ranges, for example, cause the southern 
climate patterns to extend northwards but any change 
is modified by a reduction in maritime conditions 
(Boardman 1992) This is reflected, for example, in 
the direction taken by GoydetS Line between hudundu 
gnd Crystal Brook. When climate change U considered, 
however, it has lo be noted thai rainlall, lempeiatares 
and parltciilai ly maritime condition*, arc modified by 
the topographically-high region: 

Natural vegetation in rurul South Australia has been 
affected ai low altitudes by the direcL influence of 
changes ui sea levels associated with Plcistoevne 
glaciation. A. significant part has been ufrected b> 
marked climate changes associated with the Jce Ages. 
Collectively there have been both cdaphic and 
geographic barriers lo plant migration. Both of these 
barriers have mean! that a considerable, bui 
unknowable, number (it native species has the capacity 
U) grow in S. A .to colonise sites and cnmpeie strongly 
with local species, hut these have heen unable to reach 
it by natural means. On the other hand, the species 
•which are present have suffered genetic depletion 
through intense cnvimnmermil selection pressures in 
the last two million years in rhcir rcftigin. and have 

limited intrinsic adaptability to change in directions 
not already encountered (Boardman 19861 

Much of the vegetation outside the settled areas and 
mlluenced by Aboriginal peoples, can be classed as 
ancient secondary vegetation, A large pmptiruon ol 
(he remnant vegetation, including conservation reserves 
in the settled areas should be regarded as modem 
secondary ur tertiary vegetation (Barker !W> 
Boardman 19861- These Ia*l two categories were lirvi 
affected by European migration purposefully, and 
.secondly, but without direction, from escapes oi 
introduced domestic animals and plants which have 
become frrjl Kloot (1983. IW5. 1987 ) has assessed 
plant introductions in great detail and has identified 
these from plant migrants and soioc native specie- 
Klool (1987) jlso evinced evidence that intense huntinp 
especially on isolated land, like Kangaroo Island very 
catty in the period ol settlement by migrants, decimated 
native grazing species of kangaroo and wallaby, 
transforming the native plant ecosystems rapidly and 
dramatically. Fire frequency and species life-span 
interact and thi.v affects the stability or vulnerability 
of ecosystems (Barker 1989). Such information is used 
|o manage range in the pastoral /.one Freuiienr 
levitating fires in the first half of this century, as the 
population increased, and tire protection measures in 
the latter half, also have bad a strong visible impact 
on partially cleared and uncleared native ecosystems 
in the areas south of the 32nd Parallel (Grandison 

The studies of KJoot (1987) arc- a reminder that weeds 
ate likely lo be the banc ol any attempt by native species 
jt colonisation of new habitat, even it the habitat is 
an amenable one. Many of the weed species introduced 
from overseas are much more adaptable (notice Kloofs 
method of separating natural migration from garden 
escape) and often they have a much more aggressive 
behaviour. Many ;ire natural pioneer species. Two 
examples of weed species likely to be affected in the 
climate change sccnano and able lo migrate southwards 
and compere with native species needing to do the 
same to survive, are Montpcilicr broom (Genista 
fnottspes.uilana) and Salvation Jane ischium 

It is likely that the most strongly threatened species 
m the long-term are the perennial species which 
dominate woodland formations Lou sclcrophyll 
woodland and malice formations appear to he 
particularly at risk. I hese formations occur in the wnc 
with an average growing season, in the best possible 
conditions, oi' 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 it is through different 
levels of pollen deposits, whose abundance .speaks for 
itself or through macro-fossils On the -other hand. 
many of the perenninl herb and Tow shrub layer species 



£tt Jneal ecosystems are more ubiquitous than the 

ecosystem-dominants, and frequently less constrained 
by particular sile lacu>rv This is partly because 01" 
similarity m the microclimates they generate, and 
partly, because tree-dominated ecosystems provide 
amelioration of the weather from shelter and shade 
The huge demand tor moisture by trees in locfi) 
ecosystems means that tolerance of chronic drought 
by imdcrstorey species is normal. I his is the case when 
thC undeistorey of the malice woodlands is comp^reo 
with Ihai ol loresl ecosystems in the Mt Lofty Ranges 
showing that many species have it continuity in 
distribution acros> a wide nui^e in rainfall Irom neai 
IU250 inin :n the Murray Malice to around WOO aim 
art Clcland Conservation Park (MiCann J989; Dashorsi 
&. kssop t°-90). 

Investigations nf Wttndlund productivity 
in the S.A. sLiht-arid 2011c In relation to cunwiU' 

Trees, as long-lived perennial plant species 
frequently dominate all ot major ports of the 
ecosystems that contain them They accumulate a majnr 
part of ecosystem solid dry matter in their wood and 
bar* (issues, Wood and hark arc that part of Ihc 
perrenial planus which sequester CO. and which helps 
In maintain stability of this gas in Ihe ainiospbea'. The 
Kcscarch Branch of the former Woods and Forests 
Department has investigated the amounts and trends 
in wood accumulation by trees growing in stands, h 
has been recorded on Hn unit area basts, cither as 
volume pci hectare (preferred by foresters because rt 
is bulkwcss whkh is of interest) or as dry weight per 
hectare. The overall effect over a long time determines 

Studies in S,A. were stimulated by apprehension 
about fossil fuel supplies, future costs of fuel, and the 
need to assess renewable fuel resources in the mid-7()*s 
(Kiddle ef ot, $85] I his work was expanded upon 
when the Htectntily Trust of South Australia (HTSAl 
wished id assess the opportunity to sequester GOj 
produetd from their power staitons (Boardmaii el al 
JV'^2V Productiviiy of twenty iree species growing in 
plantations over a range ol sties, soils and climate, has 
been measured, mostly as single species, but also *s 
mixtures of specie*. About one-third have been 
remeasured which means we have been able to establish 
trends with increasing age. The use uf plantations ts 
useful as both the age and the density of trees in the 
stand, or level of occupancy of site are known factors. 
It also means thai we were able to extend methods Irorn 
existing forestry science and use them to extrapolate 
i Hidings over time and space, to other localities. Wc 
have compared our result with general models d forest 
productivity, to see how local stands compare with their 
counteipaits on a global-scale. 

World plant life-zone, classification <Holdndgc l l J74) 

indicates lhal sustainable tree-dominated ecosystems 
arc unlikely to exist on land in S A receiving less Ifbui 
250 mm rainfall. Consequently our reseaiuh set nut 
to include all well-established plantations in a WW with 
average rainfall between 250 mm and 600 mm. hi Ihe 
evem, mine ol the plantations suitable for the, study 
were found in areas which received rainfall of lesft iliau 
340 mm. Dryland cereal cropping, illicit lequircs 
220 mm o^ winter veason tamlall to produce an 
econumic crop of obu/aere |540L/ha|_ rarely extends 
below the 280 mm i.sohyet (French Otf&J, However. 
HTSA asked that poientiut for trev-grownifci Hi the 200 
— 300 nun rainfall zone should be given special 
consideration. What we found is ol interest in the 
present eonte\L. 

At an ecosystem level, soil moisture replenishment 
and capacity to store water are critical factors which 
determine whether trees will grow or not, In S. A. there 
is a surprisingly frequent iiweisc cot relation which 
exists between soil depih and fertility; a targe number 
ol shallow soils, which have low capacity lo Store 
moisture and physically restrict root development an: 
fertile and an even larger proportion of deep soils, able 
to absorb all the rainfall reaching the ground and 
provide unrestricted physical capacity for root 
development are infertile in the lupsoil where £5% 
or more of roots arc nuiniatly found In both cases 
productivity is markedly behvw rhe level expected when 
conditions are Ihe btSt that can be cApcvied, 

Malice woodland gives way to lull shruhtond with 
Ataxia specie* within the 200-300 mm rainfall /ooe. 
There arc exceptions and an hiatus is present as in 
western byre Peninsula, where it is associated soils 
conditions that are unsuited to either lite-fonnarion. 
Average rauilall in a poor delimiting factor because it 
is the critical periods and intensity of rainfall which 
count the most. The annual lamfall in diought years 
is often as little as half that of medinn years Jn this 
/one, also, infrequent long periods ol dnmglt'. i>crsist. 
These considerations have loci to the concept and 
definition of "intlucntiuJ ruinftjl" The quantity uf 
moisture available to prevent willing, expressed as us 
equivalent in rainfall, is 100 mm and it has to he present 
in the upper root zone The tactors which determine 
in ii Lin a i: ri rainfall have been assessed for dryland cereal 
cropping and sites with natural vegetation, potentially 
suitable for cereals. Influential rain, on average is 
needed for 3-4 5 months in 12 The minimum amount 
of influential rainfall to produce 2j0dry t/hu/y of WtxxJ 
exceeds 200 mm. That js to say, in this zone sites 
cafxible of producing wood have lo receive water in 
excess of that received from uaitiraJ rainfall directly 
In other words, they have to be "run-on*' cites which 
receive water a> run-offer percolated as drainage in tin 
areas of higher elevauon Ihc minimum amount of Ihe 
total water which has to be available o- found to be the 
equivalent Of 300 mm, In months when there is a full 



of rain sufficient to promote* growth ( ie a surplus over 
respiratory nccdsi, u needs to be supplemented by ;ti 
least 25%. Seasonal V9*4fillOt1 is critical tor vegetation 
suivival in this 7one, The frequency distribution Clf 
sustained leugtha Iff ram)' season for a number ol 
representative weather stations has been investigated 
and is summaiised in Tabic I (Boardman KffiOL Ii 
emphasises thai growth rings in trees uiu) not be an 
anmwl sequence itf rings but of adequate rain events. 

Taw l I. 77h* frrqnvnn >/Vf tcnutn of win j$*nfar& f >ju gfttffl 

ilmvtktH A* ,w7ev rn tfte 200 mm r^ AX) ww .vivr-i^e nuftfitll 
:mv iff South Australia. 

Avenge rainfall 3G0 mm 250 mm 200 mm 

Rainy season 


Al least 5 months 17 years 

At iL'nst ^ months 40 years 

At least 3 months 70 year* 

Al least 2 months 90 years 

At least 1 month % years 

5 years I year 

L~ years 5 years 

35 years 16 years 

63 years 33 years 

8?» years 65 years 

Trurnblc (1937, 1939), considering dryland cereals 
requirements, showed thai 200 mm gross annual 
rainiall will provide annual herbaceous vegetation with 
wih-rrcc conditions for up to 3.6 momh.<- Trees in 
woodland, however, intercept a proportion of the cross 
rainfall wilh iheir leafy canopies, It the trees are mature 
and spaced so as to absorb all the capacity which is 
available for ihern to lay down new wood, ie the site 
Is fully occupied by the smallcM number ul trees able 
ro do w), then Ihroughfall, rain able to wet the soil, 
is reduced to 160 mm. This effective accession will 
only pro 1 , ide for |Va months growing season, which 
is quite insufficient to support woodland At 300 mm 
rainfall, the is likely in he sufficient for 
4 5 months. An actual growing season of four months, 
ire minimum likely to carry a suslainahle stand of 
woodland, calls for a gross rainfall equivalent of 
320 mm in Upper Eyre Peninsula and for 3111 mm in 
the Mutt ay Malice, hollo win-: Trumble's argument, the 
"run-on" requirement from these evins ide rat ions k tor 
about 150 mm water, but it must be delivered in the 
Winier rainfall season ro be "inlluentiar 

Alternative approaches to the question of minimum 
water requirement for a woodland without "gaps" bus 
h«id provided by Spccht (1973, 1975, 1983) and Walker 
ct al ilOTrt. I9SH|. Both have dealt with natural 
vegetation of the kind being discussed here. Spccht 
worked from the direction of evaporative demand and 
defined the tolerable limits bO cvapoUanspualion slress. 
Evergreen, perennial plant communities, under 
conditions which are the best to be expected, need a 
minimum net monthly supply of soil moisture 
determined by rain that falls and any accession or loss 
i*l water in the soil. We called Ihis the "soil available 

water" requirement and it ranges from 37.5 io 
62.5 mm. Sues able lo sustain woodland are 
characterised by a need lor additional water which is 
equivalent to 71% of average rainfall for the month. 
In turn, this implies thai there is adequate storage 
available in the soil within reach of imHs, 1'uktng 
canopy inlerccplion into consideration indicates thai 
run-on component needs to be not Jess than 47 *¥ of 
rainfall at a 200 mm sjte. again showing that aboul 
300 mm of rainfall equivalent is needed to sustain 
woodland formations. 

The stand structure of woodland which will optimise 
net dry matter production from lite site factors available 
can be deduced from ihe reseanh of Spccht and 
Walker. Spccht has found strong telauonships between 
l he foliage projected area (IPC) of the tallest plant 
structural layer (overwood — FPCo). and of the whole 
ecosystem fFI'Cl), to his evapotrauspiralion coefficient, 
and hased his findings on a very wide sample of matuie 
Australian native ecosystems. Mature low woodland 
in the 200-300 nun rainfall zone has an overwood cover 
ol 32 lo 12% and total ecosystem foliage cover of 
75—6%. k. there will be about 20% bait giuund. 
Walker's group took a different view of foliage eowr 
which recognised its fragmentary structure 
corresponding to about B0& of FPCo. The oplumim 
cover hased on their work, and adjusted to FPCo terms, 
sue^ests that cover LiF32± 5% should be adequate to 
siisuin low woodland on nin-on sites in the 
200-300 mm i.i i ida 1 1 zone ol S.A Their models allow 
one to assess stocking rates m young-age and juvenile 
stands as well as mature ones. 

Thus, the effectiveness of wy<cr supply can be 
associated with net primary production and ihe length 
of the growing season. These have both been mapped 
by tor present -day climate and Ihe median scenarios 
by Greenwood & Boardman. (1989) and Boardman. 
(IM**2). Net primary production is likely to change by 
183: for each incremental gain or loss of 10 mm m 
average annual rainfall These, preliminary' findings, 
now likely tu be extended further ahead in time, are 
suirlcicm to indicate, the level at changes which can 
be expected and where the changes arc- more likely to 
utcur. I he maps show die "best that can be expected" 
and until the sub-optimal standing of existing vegctatioti 
can be assessed, they form a working basis lor taking 
adaptive action Table 2 lists the tree and shruh species 
whose survival and distribution in the /.ones most 
sensitive to change, air the ones UkcJy to be influenced 
by threats and opportunities presented by the climate 
chants scenario. This is hased on Chippendale & Wolf 
11981); Boomsma (I98h, BoornsmaA Lewis (1976) aixl 
Coslcrmanns (1983). Table 3 shovwx the cucalypt 
species sorted into their botanical affinities fPryor Sc 
Johnston, 1971 > in relation to changes iu ceulogical 
slalus that are possible. 






1=1 w 
<2 | 

< < w 






v\\\v\\v\\\ nwwvwww 


1/3 Cfl <S\ IS) t/i UO CO 

t/5 i/i w on **. i/i on vi v) i!; 




rg CQ < 



J* 5 





a. g 













= -a * 

cfij S 

= ja — c 

o s £3 a 

h 3 £P ^ E 

"— l Q|i - U L. 

G c/i u -J ** 



§ § g 

K ? r ° "a Sp' a * 5 

^S?3 3 ? ? 3 s 

is gi 3 

£ £ u$ S u-S.t?*£ sTzl o 


C — a 

as « 


- 1 




C P £3 M 

xj ^ " n 2 




22 |i tfcs-s 8 

) w_ 13 n» r. _ 
g*S | & 6 

S «J CQ ^ 

S c c 

3 C 

_c "^ S- 

3 P 5 

1 3 "3 

o a 5 
CQ c« OS 

< a 52 w* ^ 

■g^: =13 p a e 3 5 3 ? ? § 

?v _-B K K si 9* »» •» -s# -3 - ■* S^^ 




Uj Uj 

'3 S f " s J g 

3 » a 5 ae 5: a 

§.§.§,§.§, &^ 

*? "5 "^ "5 "5 r -j -a 

r- g g 


*< r* n« b- t^ a -^. a 

a id 


"C ?c = 

S -S 5 a h 3 

a ^ '-<-=: 

3 S 

5 5 5 5 3 


=3 3 g 

&. Ei. 

ii ^ 5: 


"S "S ^ "S 3 a, 

uj a 5 ^ & & 



\VV\\\\\V\\\\\\\\\\\\\ \\\\\\ 


Qui tu 8 | X X w X w * 

on co cfl So cfi 

— S-i 
-_ -U C 


s 6 




O <D £ 


JS E^_ 

X) ^ 

IS ^' 


o y o oi>.= cj . 

w b _j c S 

XI 0- 


p x JS _ * o 

X 3 

6 § g.J5 ^ .a i= « - c 

^ X f - (= G 

o B "S ^Px 

5 3 

u .^ u O U 

X oj 1) W ■= zz. ra 


1 1 | « § g £ 



Emu ) 
Kopi i 



1 s 

1 1 

a § 

t« — 

o ^ 

-r-, •& 

~ -S, 



bftTa y o 

1 &£§ 

ftwij <L> *£ ^3 _9 

3^ £ > w S .= 
> ^ L0 23 Z, CQ 

























173 CJ 

£ % 


a -s^a 

y i 

iu o 3 

ST 3 S 

El (- 

P s 





\ \ \ 

\ V 

V V 


\ V \ \ 

t=&oHHH t UXXN 


*trt oq 

*-* *yi 

2 -a 


qj y 

S ^ 


3 S 

E e 

g 1 *3 jy 

_ 3 i_ — fi fl 

j* -M eg S S 

'. ^ 

3 b « c 3 -^ 


3 fi 

c »« a-S a^e cr~ 

D D y ,, i) 


&j> £ p a 1 2 



.. PL, 

o o 

pq z 







SlQ S 

SS Sal 

53 Pu P J ^ 

S - 

— ' *-• U ,■'-■ '- J - ■ - ■ '..' 

co ^ c/: kJcj^jOUj I*; t/3 w c/3 

< . CO 3 


-2 3 

s u a ^ £ 

eS! k. tri Oo^3 

x as a 

SI -Q 

5 f -> zz 

3 P £ 9 

3 « ■? ~^ 

W-W M ff « «« 

FX &,.&, &, 


H 5 H 2 5^ 

go £r 

3 p"B"5"^~5"3"q"5^ 

■J v U si 'O !j tj tj \l --j 

f o L; G K f o 5 Lj 

a r a ^ p ,£S .a ^ 

ti ci TT L& 



l \ 

1 \ 

Is § la a 3 

m en 

\ \\ 

< ffi < CG < 

< < cd a x 

UJ LJ _J W .U 

2 z g z z 

D D -J -' zj 
lt: u*. 'J". yr> -si 

x x >• 









3 a 


_i U 



r O 


U _j 

rn < 
l_ Z 

»j — : m 


c Z "c a; 

, ~ iu -. -c < 

M.= Si 

c u F S3 S 

£ -- oj •_> -c 

,t- „ aj B ,fc 


* If. 2 

cZ^< £ 5 pof2>-S ^ > I 
a r * !2 * - £ B >*. ■< ^ c * *: ". 

E ^? -- T - ? ff « -5 pi ui 

t^ > 13 "«fi «S 

'K&fc £5£ 

r( hJ 'rt l " -'"'"' 

s:^ ^S.?- 

The ecological significance of Goyder's I.ine 

The practical limits io sustained primaiy production 
tor land-use historically have been contentious and are 
likely to remain so, G. W, Goyder was put on his mettle 
to define such a boundary, which, all things considered, 
has been largely vindicated by time. He surveyed his 
line ntf demarcation for sustainable land use options 
in 1865 (MeGowan 199(f). He rode on horseback over 
30<X) miles (almost 5000 km) to define it. He 
associated the boundary with repeatedly observable 
(acts related to the southernmost extension of drought 
as indicated by vegetation and soil features. These were 
the "chewed remains of saltbushes and other lt>w 
shrubs"' and "light soils which were susceptible to wind 
erosion" when dry, indicating stlty-sandy textures and 
low contents of organic matter in the top soil. It has 
been alleged that he only surveyed the Upper- and Mid- 
north Regions but eurypolated his criteria to Eyre 
Peninsula and the Murray Matlee regions without 
specific attention being given to them. However, the 
boundary line in the central section, from near Mt.KKila 
in the west to Eudunda in the east, is only 350 miles 
long. His extensive distance travelled tor the survey 
demanded by the Government suggests he ventured 
much further afield to justify his boundary. 

The Line, recognised as a broad limit, has come to 
be associated with three features, an average Winter 
rainfall Of 220 mm, a rainfall/potential evaporation 
ratio of 0,26, (Trench 1989). and the southern extent 
of ecosystems dominated by saltbushes species: in 
particular Maireana pynumdata and M. sedifolia. 
None of these features alone is adequate to define the 
sustainable land-use criteria for the 20ih century. Of 
particular concern here is whether its original joint- 
cnteria will be assessable in lime to serve in the latter 
half of the 21st century. 

1-reneh (1989) showed thai if the Winter (April- 
October) rainfall/evaporation ratio was to be 
maintained, the boundary associated with Goyder's 
Une would have to be located where the winter rainfall 
isohyet is currently 285 mm. i.e. shift some 70 kin 
south (\\' its present location on the plains of Eyre 
Peninsula and the Murray Malice. Greenwood & 
Buardman (1989) assessed changes in growing season 
and net primary production in native ecosystems. The 
northern limit of low woodland, defined as ecosystems 
which increment new growth, under adequate 
conditions, of less than 2.2 dry t/ha/y would move 
south by about 100 km. These Iwo independent 
estimates, considering ihe uncertainty, ate in reasonable 
agreemenf. There is likely to be a drastic impact on 
vegetation. To the east of the Flinders Ranges the 
change indicates there would be a much more intense 
gradient of change from the hills to the plains than is 
presently the case. 



Vlaptariw* to greenhouse-inclined 
climate change 

One oi ibe mnvi difrleuli lasks hir managers ol land 
which grows long-lived plain species, trees and shrubs. 
especially where these species dominate ecosystems, 
is how to .assess (hat change in environmental 
conditions is actually of significance to the proem 
ecosystems, and what lo do to ensure that the 
ecosystem, lalhei lhan the species themselves, can be 
maintained Practical things to do which will make 
adaptation possible are limited by the indefinite naiure 
oi' (he lime frames. It is difficult to make a positive 
contribution over such I protracted period without 
ctfnii.s being conlonnded by normal variation in 
weather and climate year by year. One positive 
Suggesting heme widely canvassed is mainly applicable 
to commcivial iree species subject lo improvement 
through hreeding This has been to progeny-test In 
areas which have 8 climate akin to the changes expected 
hut current l> outside the regions of economic viability. 

The soil Criteria adopted by CJoyder eart "be 
ascertained on the basis <*f lexlural classification and 
decree of degradation associated with historic land- 
use and Ihe rainfall quantity and evaporation trends 
should be discernible through use of running averages. 
Consequently, a lulure equivalent of Goydefs Line 
could be used to highlight the progress of climate 
change, should it eventuate- 
Better definition of the boundary could be achieved 
with the selection of "indicator' plants, such as Maimm 
pyrisrmdifolui^ indicative of change in favourable or 
unfavourable environmental conditions, unci good 
definition of the ecological facDfts which determine 
their limitations or provide ior stability is needed- 
Ability of indicator species to colonise sites south and 
wl*M uf Ihe current distribution should be priority 
Mibjetis for ecology studies. 

Another approach which has been adopted by 
I'umary Industiit^ SA-Rirvstty. lhat recogmse> 
and woodland have multiple-uses, has been installed 
(in 1992; near Ciiimeracha in a Demonstration Forest. 
This investigation has been designed to integrate both 
long term and short-term benefits. Uses included are 
wiiter supply catchment protection, agio- forestry and 
potential commercial forestry. Botanically closely- 
related land-races (provenances) and species, in live 
distinct groups of Fucatyptw similar in fashion to Table 
St have been matched to three climatic models. Trees 

are being gmwn from seed collected ft) in good quality 

local populations in S.A. tit) I mm central Victoria 
where the rainfall distribution pattern is like the 
postulated change, bur the temperatures do not change, 
and (iii) and populations in ihe central part of eastern 
NSW where rainfall distribution with a tendency to 
mi miner maxima and wanner temperatures occur. The 
(lowering, seed set and natural regeneration phases of 
ecosystems, as mentioned above, are likely to be the 
most sensitive scral stages to significant climate change 
The Gumeracha investigation aims to lover this 
eventuality by providing genes in the dominant species 
which contain adaptation to climate in the directions 
in which it \a very likely to change, The trial is 
providing a bank or nest egg nt genes lo allow 
adaputtion to occur whilst still likely to retain ecosystem 
structure associated with land-use 

A Forest Reserve at Whyte-Yarrnwie- in the zone cast 
of the southern Flinders ranges, which our studies 
indicate will suffer the most intense changes and 
increases in stress, ha*, been teserved tor scientific 
studies One option we are considering is to plant a 
suite wf 'indicator species', in range from those suited 
well at present to others crowing in regions akin to 
the future climates of the scenario. Planting hi a set 
design at ten-year interval lor the ne.xt century would 
be immensely valuable in an ecological sense. Plaining 
a bracket of three consecutive years ar the decadal mark 
would help to reduce the effects of annual variation 
in weather. Such a programme calls for dedication. 
conscientious adherence ro ihe plan and consistency 
to be adopted by managers, and lasting over several 
generations of ecologisls. It may be loo much lo ask. 

A second question concerns conservation of the 
current species composition of extwysicms which are 
valued and preserved in Conservation and National 
IViks. Greenwood & Bourdmau (1989) reviewed ihe 
impact of the median climate change scenario on the 
repiesentalivencss of Parks m S. A. There has been no 
significant addition to options suggested m 198K. 
Provision of gene banks and corridors of land able lo 
permit species migration, mainly southwards is still 
seen to be the best option in the interim. In the absence 
of physical evidence to justify the need or conservation 
practices, sadly little has been added beyond icvision 
of the Native Vegetation Clearance Act. to strengthen 
its preservation provisions, and to positively protcci 
roadside native vegetation, There is still a need to 
include and emphasise whole-ecosystem conservation, 
and to modify current preference and emphasis on rare 
and endangered species. 



Appendix I. Average annual temperature, annual rainfall and growing season est i mutes far recent times and the late 21st 
Centura adapting the IPCC 199/ median estimates of '■■Change in temperature and rainjatl with latitude. Growing season length 
has been esttnuited iv///i die De Munontic Drought index pi mi mean mtmthly rainfall and temperature data and estimates 
far reeeni times and the late 21st Century respeitively jsee Bisardnum, 19X91, 





Growing Season 







in G.S. 


Late 21C 


Late 21C 


Late 21C 

% of year 

Eyre Pen. 







2 5 


































Pf Lincoln 

16 35 








16 55 















Streaky Bay 








Yorke Pen. 


16 6 

20 05 
















19 35 
















































Snow town 

















Raseworthy Coll. 











IS. 45 



































La memo 
















Mt Lofty Radges 

i Ml Crawford 







+ 16 









Upper .SB 



IS. 35 














lower SE 

















Mt Burr 









AlllN. R. J. (198H) El Nino southern oscillation influences 
in the Australasian region, frog, in Phvs. Gog. 12, 

Al'SMAP (1989a) Natural vegetation about 1780. Map 
CSU89/055 (Surveying and Land Information Group. Dept 
of Administrative Services, Canberra). 

( 1989b) Present vegetation in Australia. Map 

CSL'89/054. (Surveying and Land Information Group. 
Dept of Administrative- Services. Canberra), 

Bapkpr. S. (19K9) Natural ecosystems in the Greenhouse, 
pp. 1 05- I OK In Dendy , T . (Ed.) "Adelaide Greenhouse *88 
Conference", fDept of Environment & Planning. 

Boardman, R. (1980) Perspectives on managed ecosystems 
for the semi-arid /one of South Australia pp. 1-16 fti 
Section 13- Agriculture and Forestry. Proceedings of the 
50ih Jubilee ANZAAS Congress, Adelaide fWoods & 
Forests Department mimeo). 

0986) The history and evolution of South Australia's 

forests and woodlands, pp. 16-31 In Wallace, H. R. fhd.) 
'The ecology of the forests and woodlands of South 
Australia" (iS.A, Govt Printer. Adelaide). 

__(19X9 The impact of the postulated enhanced 
Greenhouse Effect In Dendy. T <F.d.) -Adelaide 
Greenhouse "88 Conference*" (Dept of Environment & 
Planning. Adelaide). 

(1992) A study of the growth and yield of selected 

plantations of native trees growing in the semi-arid #>ne 
of South Australia and their capacity to .sequester carbon 
dioxide (Dept of Primary Industries, S A.; Woods and 
Forests, Adelaide?, 

Boomsma, C. D. fl98h Native trees of South Australia. 
2nd edtn Bulletin 19, 1-228 (Woods & Forests Department, 

& Lrwis, N. B. (1976) The native forest and 

woodland vegetation of South Australia. Bulletin 25, 1-313 
(Woods and Forests Department. Adelaide). 

Chippendalei, G. M. & WcjLr, L, (1981) Natural 
distribution of Euealyptus in Australia Special Publication 
No. 6 ( Aust. Nat. Parks and Wildlife Serv.. Canberra) 

Ci.TMATn Impact Group (1992) "'Climate change scenarios 
for the Australian Region*'. (CSIRO Division ot 
Atmospheric Research. Mordialloc). 



Costermans. I.. (1983) "Native trees and shrubs of south 

eastern Australia 4 ( Weldon Publishing. WHlctughby. NSW). 
Da.shoRst. D. R. M & Jessof, J. R (1990) "Plants of the 

Adelaide plains and hills". (Kangaroo Press. Kenthuxst, 

Fkanmin, J. r.. Shi.'gaki, H, H. & Harmon. M. E. (1987) 

Tree death as an ecological process. Hiosartice, 37, 550-50. 
French, R. J. (1989)" The Greenhouse Effect and its 

implications Ji>r prmiarv pniduction in South Australia, 

pp. 137-142. In Dendy, f . (Ed.) "Adelaide. Greenhouse '88 

Conference." iDept of Environment & Planning. .Adelaide). 
Friths. H. C. (1976) "Tree rings and climate'' (Academic 

Press, New York). 
Gmiord. R. M. (1993) Review Responses of vegetation to 

elevated atmospheric concentration of CO*. Climate 

Change Newsletter 5(2). 10- 1 L 

Implications of climate change for South Australia First 

Report. (Tiie Government ot South Australia. Department 

of Environment & Planning,. Adelaide). 
GranDIsov. R. (1983) The relocation of fifteen sites painted 

hy S. T.G ill whilst accompanying the HoiToek's expedition 

of 1846. Row G&mr, Soc Australasia. S,A. Branch. Proc, 

83. 12-21, 
Greknwooo. G= & Boakdman. R. (1989) Climate change 

and -some possible effects upon the terrestrial eeologv of 

South Australia, pp. 112 132. In Dendy, T. (Ed.rAdclaide 

Greenhouse "88 Conference". (Dept ot Bnvironmem &, 

Planning. Adelaide). 
Griehn. T. & MiCasmi i : M. (Eds) (1986) "Atlas of South 

Australia (S.A Govt Printing Division & Wakefield Press, 

Hoi dridgi ■ L. R (1974) "Life zone ecology". (Tropical 

Science Center, San Jose, Costa Rica). 
Howard. G. W (1967) Arthropods inhabiting pine titter in 

the South-east of South Australia. M.Ag.Se. Thesis. 

University of Adelaide Unpubl. 
Kiddle, G,,, R., Archer, G. K. & Vav Dfr 

Sommen. F. (1987,1 A study of the growth and 

characteristics of woodlot and amenity tree plantings in 

semi-arid rural South Australia. (Wood*. & Forests 

Department, Adelaide). 
Kioot, P M. (1980) Dr. Richard Schombunik's 'Naturalised 

weeds"' (1879). / .Adelaide Bot- Card, 2. 195-220. 
. (1983) hailv records of alien plants naturalised in 

South Australia, 'ibid. 6. 93-131. 
(1985) Plant introductions to South Australia priot 

to 1840. Unci, 7, 217-231. 

11987,1 The naturalised tlura uf South Australia. 4. 

Irs manner of introduction. Ibid. 10, 223-240. 
LfiWrN, R, (1985) Plant communities resist climatic change. 

Seietur 228. 165-166. 
McCansj. I. R, (1989) "The Malice in llowc-r (The Vietoriaji 

National Parks Association, Melbourne). 
McGowan, R (1990) Govder's Line. Heritage Australia. 

Sprint*, 1990. 1-7. 
PENRluGE, L, K. <fc W.ALKER, J r (1986) F.ffeU o! 

neighbourmft trees on eucalypt growth in a semi-arid 

woodland in Australia. J. Eart. 74, 925-936. 
PosSIN'gham, H. P. (1993) Impact of elevated atmospheric 

CO-, on biodiversity: mechanistic population-dynamic 

perspective. Auxi. J, Bat. 41. 11-21. 
Pkvor, L. D. & Johnston, L. A, S. (1971) A classification 

of the eucalypts (AND Press. Canberra), 
Rliter. 1 H. (1964) Firms ntdiutu D.Don in the lower south- 
east of South Australia. M.Sc. thesis University of Adelaide. 

Simpson, J. A. (1967) Mycological aspects of decomposition 

of pine litter M Ag.Sc. thesis. University of Adelaide. 

Sparrow, A (1989) Malice vecetation in South Australia 

pp. 109-124 hi Noble, J. d A Bradstock. R. A. (Eds) 

"Mediterranean landscapes in Australia. Mallee ecosystems 

and then management"' (CSJRO. Melbourne). 
Souhi> Oh PLAN! Physiologist (1993) Aust. J Boh 41. 
SpEcht, R. L, (1973) Water use by perennial evergreen plant 

communities in Australia and New Guinea. Au±t, J Bar, 

20, 273-299. 
1)975) "The vegetation of South Australia" (Govt 

Printer, Adelaide), 
__ (1983) Foliage projective cover of overslorey and 

uniierstorey strata of mature vegetation in Australia. Aust. 

J. Ecol 8. 433-439. 
Tru.mri.h, H. C- 1 1937 ) The climate control of agriciiltuic 

in South Australia. Trams. R. Soc. S. Aust. 61, 41 62. 
(1939) Climatic taeiors in relation to the agricultural 

regions of southern Australia. Ibid. 63. 36 43, 
Walker. J , Sharpe. P L t Penkidge, L. K. & HsrN-l Wv 

(1986) Competitive interaction between individuals of 

different sue; the concept of ecological fields. CSIRO, 

Institute ^f Biological Resources. Division of Witter and 

Laud Resources. Tech. Mem, 86/11, 
. Crapper. P. F. & Penridce, L. K, (1988 > The 

crown gap ratio <C) and crown cover: The held study /inst. 

J. Eeoi & I01-I0S. 
Waring. R. H. (1987) Characteristics of trees predisposed 

to die. Biosaente 37. 569*574. 


By Michael E. Burns* & Cliff Walsht 


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 

Key Words: Climate change, economics, greenhouse gas emissions, anthropogenic 
influence, economic models. 

Tmnsih:ti<*n.\ nj the Jttmt Society of S. Auu tWfy 118t.l). 83-H1* 


by Michael E. Burns* & Cuff Walsh t 


Hvkns, M. E. &, C. il*W) The economic implications of Oiinaic change, jfwvt. /?. Soi. S -tusf. liSto 
83-?* 31 May, 1004 

Feedback effect* frprfl the rest of ihe world on the economic structure and well being of Souilt Australia aic 
likely tu be more significant than direct economic impacts of climate chanpe m South Australia or even Australia 
as a whole, tconomist* can help to identify climate sensitive sectors of the economy and analyse how change 
in them will affect other sectors. Economists can also influence anthmpogeme changes by prescribing appropriate 
incentive xttueiure to redirect human actions in the common interest. Because of the nature of the data upon 
which economic models ate built, their predictive capacity is restricted to short period;. i>^ 1-3 years It is postulated 
that areas such as agriculture, forestry. HtiBfttifi and hunting would be highly sensitive, to elimaie change whilst 
electricity, gas and water, construction, a-crealional and other personal services logethei with ownership wtfilwel hugs 
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 pas emissions 
in jn economic framework are controversial and difficult to resolve in a global context and the significance- of 
eolhihnraiive HpproaLhes to these and other problems cannot be overemphasised, 

KFY WORDS climate change, economics, eax emissions, anthropogenic mihiciKC. economic models 


An important feature of the symposium fo>T which 
ibis p^per initially was prepared was the hringme 
together ul at least a little more information abom the 
likely regional and local effects of climale change 
ibeyond very tentative material available fiom the 
Greenhouse 'KS Adelaide workshop and elsewhere). 
The contribution economic can make doesn't depend 
totally on this information, however Like other 
sciences (social us well as physical), economics can 
contribute lo discussion <md debate through increasing 
understanding and application ot its way of thinking 
about issues and problems We are not here referring 
to the common view that economists insist on reducing 
everything to dollars and cents. Rather, we mean the 
more general contribution that economics can make 
through its development of frameworks for evaluating 
public policy oplions which 
» emphasise costs as well as benefits of policy action; 

* pay patlicular attention to incentive structures m 
policy design; 

* TCCttgmsc how the actions of one individual, 
tmterprise. region or nation impact on others; and 

* titnwirliMandlng the usual presumption that 
economises think that markets work perfectly, 
acknowledge cases where "the invisible hand" of the 
market is either arthritic or non-existent, such as 
when l here -ire spillover effects, poorly defined 
propeny rights., mgh uansacnons costs, or the need 

School <»f KcHiomics, Flinders- University o\' South 
Australia, OPO Box 2100, Adelaide, South AuM.tihn soot 
South Australian Centre lor Fconornic Studies, The 
Lm varsity tM Adelaide. South Australia 5005. 

to impute "existence values" (eg. of life on earth) 

exists, and design policies to moderate their 


Moreover, like those physical scientists who arc 
deeply engaged in attempting tn understand climate 
chance and the role of greenhouse gases in il, the 
economists' framework is general equilibrium in nature 
and, to the greatest extent possible and necessary foe 
the problem at hand, it h$ global in its recognition of 
impacts. To be more specific, the economists' 
modelling includes fecdhack effects between markets. 
including from wotld markets. 

This observation leads lo our firsl, simple* and yet 
profoundly significant point about the economic 
implications of climate change. That is, lor a small 
region like S.A., with a not highly unusual economic 
and physical structure, teedback effects on our 
economic structure and well being from the rest ol 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 (X modelling of various sccnanos 
produced in late 1980's, estimates of the loss of World 
output and income (often extrapolations from specific 
country studies, especially the USA) ranged from 
\%-b% of GOV. for temperature increases in the range 
2-10 degrees Celsius These are very large output and 
income changes in absolute terms, and their effects on 
gaiwth of demand for South Australian products would 
be veiy substantial and likely to outweigh any direct 
adverse effects of climate change in South Australia 
on South Australian production and incomes. 

The estimates of climate change, however, remain 
highly uncertain and h«/ve tended to be progressively 



reduced in more recent years. The uncertainties about 
whether, when and how climate change will OCCUI 
moreover; are compounded by Ihe lael thai. Ihcv arc 
based on a presumption ifun policies will noi change 
soon enough, or sufficiently, to alter outcome**. 
Predicting policy responses is as important, in the end. 
as predicting climate change ilsclf 

Lncertaiittfcs in eliuiutc prediction: 
ihv policy feedback, issu<* 

Others are hetier qualified than us |n explain the 
nature ol the models heme Used to generate clmtace 
piedietmiis based on observed trends in greenhouse 
w.i\ emissions, and the uncertainties that xunound 
them. Whai we might usefully focus sonic attention 
on. however, is the uuci-rc-luted lssUi? of policy 

The economic pn'jeclions ol the consequence pf 
climare change that currently exisl .ire ha>ed im a 
"worst case" policy scenario of no significant change 
)ft policy hi response to emerging, or unlteipaied. 
clinialc changes. Thr^ dcurcc ol pessimism may be 
unwarranted, but climate change certainly involves 
sotne characteristics which give pessimistic outcomes 
a higher than usual weight in probability distributions 
delined over likely responses. 

Policy responses lo climate change have 
characteristics similar to what economists, in other 
contexts, call public goods. For example, in ihe lanuhai 
case ol a system of national defence, once u unit of 
"protect ion" is provided lor one person 11 is 
simultaneously provided tor till others, and those 
unwilling lo contribute lo the co\l.% cannot be excluded 
J'roo) benefits. In largo numbers contest, individuals 
see themselves as having negligible impact on 
quantities produced whether chcy offer to contribute 
to ct^ts or not. and therefore ihe rational response is 
lo-uUctupt to free ride. Oven though all others may do 
likewise, and nothing t^ pr*>duccd as a result, it Mill 
will not pay to offer lo contribute because whether you 
do or not ecncrjljy wul not aftect decisions of others 
This is a ■ Uitvsjc case ot market failure, and a case 
where governuieui provision, funded throuoh 
compulsory taxes, is inevitable and desnahle. 

It should he noted, nonetheless, that even m the 
relatively simple case ol defence, giving governments 
the power lo coerce" docs not ensuic that an optima! 
quanhty ot detente will be produced. Voting, lobbying 
und other means by' which people signal ihcu 
picleiences in the political market have peculiar 
characteristics, and the incentive structures facing 
political and bureaucratic decision-makers have a 
tendency 10 encouoge oversuttpty of public seeloi 
goods and set vices. 

Clearly, policies to control climate change have the 
same characteristics as defence: ellective action will 
benefit all nations, whether or not the) contribute to 
the cosi (financially and/or their own policy actions). 
They also lace the added complexity, of course, thai 
it government does not exist that can coerce .nation 
slates to pursue policy changes, We have lo rely on 
international cooperation, a notoriously ran. 
phenomenon, pursued through complex procedures, 
which is likely to see countries and gioups holding out 
(free t'idingj at others expense - even when 
perceptions of the likely costs 0!' lailurc to obtain 
agreement are broadly agreed to be high. The deferred 
nature of the likely costs of glohal warming, even d" 
agreed lo be inevitable, add lurlhei difliculties ol 
getting cooperation 

This i% nol intended lo be loudly pessimistic - just 
to reinforce what common experience (it noi common 
sense) predicts about global policy change- (ioeii 
scientific uncertainties, watchful caution aboui glohal 
action is desirable anyway. The classic example ol 
i'hoiuas Malthus" 18th Century prediction that global 
population would grow exponentially but fond 
production only arithmetically, and the Jailed Club of 
Rome predictions in the ]%0s. serve as eloquent 
warnings against precipitate policy responses. On a 
more optimistic note, there is some evidence thai 1rec- 
rider behaviour gives way to greater cooperation when 
the stakes are agreed to be very high, it only tor leat 
that failure to participate or contribute will result in 
retaliatory aclion. 

1n all likelihood, the less developed wot Id (l.DCsi 
will be less willing to bear the cosl ol policy change*. 
and more likely l<» slay outside any agreements CWCli 
though they also are more likely lo use technologies 
which arc inefficient from the viewpoint of greenhouse 
gas (Gfi) emissions One consequence of this is thai 
(he "price" the developed world. South Australia and 
Ausiralia included, has to pay might be not just (he 
cosl ol icducmg emissions m then own areas, but also 
substantial resource Irausteis to I DCs to buy their 
paitictpation Those costs are likely 10 he high even 
without transfer-* to I. DCs: it has been suggested, foi 
example, that reducing greenhouse gas emissions by 
Ihe order proposed under the Commonwealth's #90 
interim planning target would be cost 1-3% ol 
AusinuVs GDP fIC IW>. 

Despite the lack of accurate and detailed predictions 
abc-in chmaie change, about which elements are due 
to human actions, and in what ways policy-makers 
world-wide are likely to respond, there are still a 
number of ways in which economists can usefully 
contribute to the debute. Tor one thing, they can help 
to identify climate sensitive sectors of the economy 
and analyse how changes in litem will allevl Mllliei 


Taw , I Breakdown of Sou] h Australian g«9Jto state prodmi and employment by se/iswvity w ctimatt change. /*W-W 


Slate fotai 
High SeiiHittvit) 

Agiiv'ultuiv, forestry, fishing and hunting 
Morlvrak* stiiMtivitv 

Electricity, gas and water 


Recreation. pcCTUHa) and other services 

Ownership of dwellings 

Negligible sensitivity 



Wholesale and retail trade 

Transport, storage ami communication 

Public Administration and Community Service* 

Finance, property and business services 



% of 


















37 1 























35. 8 










Identity tag sensitive areas and 
relevant flow hid effects 

fceonomists have their own range of "global models" 
aimed al capturing ihe interactions between markets 
within and across counties. As in other areas of 
modelling, (here are well-understood limitations 
associated with the use of models. For one thing, tor 
opera/tonal reasons (tractahiluy of solutions, etc.). 
there are trade-offs between the level of disaggregation 
and detail, the richness of the dynamic specification 
and feedback mechanisms, and the complexity of the 
functional forms use to describe decision-making 
hchaviour. For another, not only are data limited, but 
laboratory experimentation and replication ftf 
experiments are not generally possible. Moreover. 
because of technological, climatic, political and 
population changes and so on. many of the parameters 
of an economic model also will change over itnie: 
depending on the variables in question, useful 
predictions may only be made for the very short-run 
(1-5 years ahead). 

Subject to Ihe above reservations, however, it |S 
possible both to identify qualitatively those parts of 
the economy (and of other countries' economics) which 
ate most sensitive to climatic change and to discuss 
gencmlly how changes in ihe affected sectors might 
How -on through the economy as it whole A simple 
approach, illustrated in Table I is to consider a 
breakdown of the economic activity of a region such 
as South Australia into the output and employment 
contribution of different industry sectors, noting the 
sensitivity of each sector to climatic change. (We follow 
the classification of sectors adopted by W 1991). 

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 into more 
specifics. For example, we could track the impact on 
other sectors which supply inputs to agriculture of a 
climate-induced change in agricultural output through 
the use of input-output (or, preferably, computable 
general equilibrium) models, it is important to note that 
the relevant analysis must lake into account the global 
economy. To illustrate this, while agriculture may be 
hurt ( it may also become more pnxluctive) in Australia . 
because it is hurt elsewhere such as China. India or the 
USA, Australian exports may become metre valuable and 
Australians become better off, oven though total 
Australian prtKtuclion is reduced. By the same token, 
however, it other countries become poorer, the value 
of our exports could diminish. Sorting out these issues 
clearly is vital to understanding the consequences ot 
climate change for us, and globally (since we assume 
South Australians have the well-being oi others as part 
of their concerns). 

The need to take into account the global international 
trade interactions has already been the subject of analysis 
and research by economists in North .America and 
Europe At the most recent meetings of the prestigious 
American Economic Association, an entire session was 
given over to Greenhouse Warming considerations. At 
this session. Redly & Holimann (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 A. Hohmann, through its 
analysis of the interactions of global regions (North 



America. European Community, tormer Soviet Union, 
northern Europe. China, Japan. Australia, Argentina, 
Brazil and "Rest of World"), gives powerftil illustration 
of rhe complexity ol climatic change effects 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 
fin line with international prices). 

Evaluate particular scenarias 

As was done hy CSJRO (MS and revised 1992) 
specific (and moderately plausible) climate change 
scenarios can be considered- The major characteristics 
Of these scenarios for the Australian region are set out 
in Table 2. 

Using the most recent of these- a group of economics 
graduates (Hutchins el al 1993) in a project for South 
Australian Department of Environment and 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 ol 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 
ORANJ, to examine the consequences of reduced 
wheat yields, increased forestry and fishery 
pioductivily and so on. Second, there have been 
evaluations of some of the more extreme sources of 
costs due to flooding droughts and cyclones (an 
evaluation ol costs, of course, opens upon 
considerations of costs of ''abatement"). 

The first of these approaches, embedding the 
predicted effects of climatic change into supply shocks 
in the agricultural sector Or (national and global) 
economic models, has been Widely used. Comparison 
of the results of 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 
being considered from one siudv 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 general 
scenario, which involves temperature increases of 2°C 
in the tropics and in the range 4°C to 12°C for the polar 
regions, has agrieulturuJ productivity increasing in the 
North of the (former) Soviet Union. Canada and 
Europe, but reducing in the United States and most 
of the rest of Kurope due to drought (see Tobev a ui 

In addition lo incorporating economic feedback 
mechanisms and international trade linkages, some 
attempt has also been made to extend the interface 
between the economy and the environment by 
embodying possible agricultural sector responses. 
Questions ol "adaptation potential" and "adaptation 
capability" are considered to bear- respectively, upon 
crop substitutions that arc potentially available and 
desirable due to climatic change, and upon the 
constraints lo these substitutions due to poorly 
developed markets lor crop inputs and a range of 
infrastructure considerations, including the skill level 
of the agricultural labour lorce. 

Taw r 2. CSJRO tfimau tfrHtfri S&Htitim Inland J992. &totf/ftti fur war 2030, Sounrs. Ptwnmti itQXH) am! Climau- 
Chunxv Gnmp (J992)* 

19 K7 






-2 M 4°C 

-50% in north 
-20 V; iii si >i ill i 

20-10% \ncrcnw 

+ 20 to 140 cm 

Extend 200-400 km farther south 
Frequency 30-60$ 

Rise 1(X) m per l°C warming 

Decrease 20% North ol 3ft°S 
fnOrfcEAt' south of 36°S 

+0.? to 2.5 D C 

Summer lo -207; 
Winter SA to lOfl 
SE AUst -10 to I 10'!; 

General increase 

-5 to 35 ecu 


Rise (00 m per 1°C warming 

Strengthening of monsoon westerlies 
in nonh Aust. & me SE trade winds 
in summer. 



Insights iotii (he occurrence ttf 
anthropogenic changes 

N^>ncoflru*anal\ses considered so for distinguishes 
between changes due t* * human acnons anil change 1 , 
which hu v c oihei causes. It appears 10 be true that, 
by accident or otherwise, human impacts on cUrruiU*. 
so lar tanil tor some lime ahead), may have made dungs 
heller lor humans than would otherwise haw hcvn the 
disc. There is also the RflCJ that, \o some extent, (he 
living world adapts to new conditions 

Ihcse possibilities apart, it is well understood by 
economists Itiat individuals will sometimes impact on 
Iheir cnvimnrncnl in a way thin reduces the common 
irood. through, for example: 

■ externalities (spillover cffccisl; 

• inadequately defined property rights 

■ transaction costs 

• failure 10 recognise "existence values" (eg. ol tile 

on earth}. 

For such case*, economists alvo are able In presenile 
the appropriate incentive stiueturc tu redirect human 
actions in the common interest. 

Lxternahlies occur where the act tons ol one 
decision-maker impose costs tor bestow benefits) upon 
other individuals. Problem*, iieeur when the rplfcWtfil 
decision-maker is not required to take into account 
Ihese external cn.Os (or henelils), so that an 
inappropriate level of an activity is undertaken. As an 
example, a manulaclurer, in ihe course of generating 
income, tifoaj use a process that involves signiovanl UG 
ciiiissioiis. Suppose thai these emissions lead ((Act 
lime) to a elinulic change which reduces global 
agricultural productivity and results in a range of price 
increase* of basic commodities. Because- the 
manufacturers derision whether to produce or not does 
not have to take into account die (external) costs 
imposed upon others, a siltnition can arise where the 
value o! a manufacturer's output, over and aboSe die 
(internal) costs of production, is less than the costs 
imposed upon the rest of the community through the 
impacts upon agriculture A tax on emissions would 
encourage the inaiiufaeiuici to behave in a socially 
preferred way. 

Whether the presence of externalities is a major 
problem in practice Ircxjuetiltv depends upon how well 
denned are ihe property rights of a particular resource. 
Ownership of resources such a* air, water and other 
facets of the natural environment, lor a variety of 
reasons, often is poorly defined If il is mil defined 
in law who owns a particular rcsotiiec. theiuonsumei\ 
of the resource arc withoul legal recourse when ihc 
quality and quantity of iJil resource is reduced dtie In 
the jctions of others With regard to the earlier 
example, agricultural producer^ are not in position to 
sue the iiiauutaclurcr(s) tor a loss in pmductoiU 
brought al>»'ut by increased GO emissions. 

It might seem that externality problems can be 
handled by the simple act ot ensuring drat die 
ownership of resources (such as air and water. ere) 
is fully deiined under the law, Leaving aside for the 
moment obvious legal complexities, if. lor example. 
the agricultural sector was given property rights over 
the atmosphere, lhal sector could (m llieory) sue the 
manufacturer for the costs due to GG emissions. Even 
if ihc manufacturer was granted ptopcity rigtusover 
the atmosphere- then (in theory) a better outcome could 
he obtained as the agricultural pioduecfs could decide 
to pay the manufacturer to decrease produclion or to 
inve\i in a cleaner pmccss. t'l his may seem "unlair", 
but it would result in moie socially efficient outcomes*. 

h is. perhaps, useful lo remember here lhat 
economists do not prescribe ?ero emission of GG's or, 
in general, zcai levels of any other form of TpoHmiori*. 
Most anthropogenic aclivitiev are "polluting" or have. 
externalities to some degree and economists see the 
problem as halaneing (on the margin) ihe full costs 
(including externalities! of an aetivit) with the benctits 
of thai activity The assignment of property rights can 
go some way to seeing that decisions aboui activity 
levels lake some account of externalities. 

This question of identifying and taking appnipriatc 
account all relevant costs is ceutial to t)ie economist's 
trade. It also leads directly to a further complication 
in the treatment ol cxlernalily producing activities. 
F.vcn if property rights could be clearly defined, there 
may be substantial transactions costs involved itj 
obtaining the desired outcome. With regard to ihc 
emissions ol GG's, which manufacturers arc 
responsible tor which proportions of the emissions, 
and how much arc individual agricultural producers 
fiftected (and by whom)? At the very least, there OOUld 
be very vubslantial monitoring costs involved, and the^e 
would be "compounded" by further legal and 
administrative costs As a resuli. (he costs ol'idenhfvmg 
and achieving appropriate levels ul GG emissions may 
be greater lhan any benefits obtained Fconomists must 
design policies that Like into account the costs of 
administering ihc policy. Il is fur litis reason that a range 
of othei policy-measures arc often vousidc ted, such 
as "pollution" taxes or dclemml fines. The proposed 
hydro-carbon lax. considered in more detail below, is 
one such example. 

Consideration of factors such as these give, insight 
as to why ordinary decent citizens of planet Earth may 
make decisions that arc rational for iheir individual 
>elves, but contrary to the globed good. Understanding 
these factors also is necessary for policy design, but 
unfortunately may nol he sulficienl. Policy parameters 
can only be set on the basis of agreed valuation's of 
current and future states of the world. Such estimates 
must include valuations ol quite complex items, such 
4s environmental resources or even "tile on earth* The 
difficulties here include the absence of, or irrelevance 


M I- Hl'KNS £ ( WALSH 

of. a simple market value and the question ol' how the 
[uitiriB should be valued relative lo the present lime (the 
■ Ii.mlo of u discount rate). 

The difficulties in evaluating assets such as the (ileal 
Baincr Reel tyl Ajitatctica are well understood The 
novate au<. lion of a communal good will always ignore 
the values held by individuals vvjjfl will he excluded 
(nun nseof the asset; the imputation of value on the 
haxis ol euirent use ol a resource ignores bolh llie 
option valued of those who would value ihc 
oppottnmiv lo possibly use a resource in future and 
the "existence values" held by those, lor example, who 
will nevrr use a icsouree in any way hut who simply 
value 'Is presence tflld preservation 

Astahsis nf alternate policy measures 

Insights titlo the reasons Why human aclim-o arc 
likely in lead to undesirable lewis ol greenhouse gas 
omission and associated climatic change, such as those 
discussed above, enable economists to prescribe 
incentive structures whteh will rcdirevl hilh'ian actions 
toward the common interest There arc obvious 
m lot mammal obstacles \o Hie detenninatuin iff such 
incentive structures, as can be illuslralod by brief 
consideration ot an ^ideaf data situation. 

<Vjirly. it would be useful d" it was known exited v : 
li'-w die emissions associated withdilYererii production 
prvKY>ses and consumption activities impacted upon 
lutute pioduetion and consumption, the "taste" 
par-ulneiers ol the population that describe the 
relationships between consumer choice and prices: 
and other taclors, such as the structures of the JiOercnt 
markets m the economy and Lhe policy objectives ni 
domestic and foreign governments. If all of this was 
known, then, as in any other closed equilibrium system 
whose descriptive parameters are known, an optimal 
intervention xitaieey ootiW be conceptualised and 


Fn practice, ihe Itamcwork within which we have 
to work has a! best only some stable parameters and 
a ranne of stochastic elements, HirrhctuMrc. whatever 
stable parameters may exist are not known with 
certainly but must be estimated, and the precision of 
such estimates is itsc 1 1 dependent upon out 
understanding of the nature of the framewnrV itself 
Of" .nurse, no less is Irue regarding the estimation ot 
the parameters of systems in lite physical sciences. 

The impact ol all ol litis uncertainty is that economic 
policy prescription often bc-a cart he thought i<f as being 
a two slage process, Pirsl, largely on lhe baxis t>( 
Theoretical considerations*, tl is possible to make 
qualitative prescriplions thai will be nabusi with respect 
to a number of alternative models and parameters 
de*ctihuie teal wot Id behaviour. The general 
su*^esttou, teJet icd to above, that M hydro-carbon u\ 

would reduce the incentive for participation in activities 
which contribute significantly to ©3 emissions and 
climatic change, is an example ol a reasonably robust 
qualitative policy prescription. 

f he second stage is the more difficult. Ii <s one thing 
Ur be aware, thai a hydro-car bun tax can impact upon 
behaviour ma desirable manner, hut it is quite anoihei 
to know what st/c lax should he used and on what 
particular commodities U is 10 obtain insights into ilits 
latter question that mueh current ceonomie policy 
research is directed. At the recent American 
Kconomics Association meclme metiliotied earlier, two 
examples were provided of this kind of research 
activity Not surprisingly, bolh papers use models 
embodying gross simplification of the glolvtl economic 
and climatic interlace. 

tu Nordhaus (IW), dynamic integrated cbmttie- 
economy IDIOT.) models are extended Tt . by 
integrating the econotrue costs and henefits ol OH(i 
reduetions with a simple dynamic representation ol the 
scienltlk links ot emissions, concentrations :ind elimaio 
change" (Nordhaus 1993 pp. 313% Conditional upon 
;in estimated loss tu overall N'attonal Product ol W< 
for every 3°C warming. Nordhaus' study suggests thai 
the optimal carbon-tax policy would involve an initial 
tax of just over S5 per ton, i islny sleaddy over tune 
lo reach about S20 per ton (based on turrenl prices i 
hv the end of the next ecnlury. It is estimated thai the 
net beneftt.s. globally, of such a policy 'ompared to 
doine nothing would be of the order of Sto billion 
annually. More xigntlieant is the conclusion thai fhr 
popularly recommended immediate cut ol G(i 
emissions (iclalive to 1990) would require an milial 
Utx of $5Ci per ion and overall, compared to a uo-Lonlrol 
situation, would actually involve a global cost of over 
$750 billion annually 

Feonoruic policy, whether to tlo with OU emissiou>. 
or otherwise, is in itself a complicated and controversial 
issue. There has been much argument that public 
bodies arc inelUeient m their own activities and thai 
la.x revenues are badly used It is also widely accepted 
(hat ntosi taxes, including many existing ones, cause 
inefficiencies in the economy as a whole by preventing 
lhe price system from signalling the "true" values ot 
commodities and resoURvs. With respect to issues such 
as these. Nordhaus makes two uiteresliug observations 
with respect to ihc imposition of even the "optimal" 
rate of tox on appropriate commodities. Hist, tt the 
lax revenues are used with a phmsiblc level ot 
inefficiency in the public sector, all bencltts due t0 
GHG emission reOucnnti may be totally outweighed. 
Second, however, il these lax revenues could he used 
to eliminate existing laxes winch were distoiung the 
pi ice mechanism in a eostly manner, the annualiscd 
global benefits nimbi mueiise irom Slfi trillion to over 
^250 billion. 



The above is useful in that it gives a flavour ol iiow 
economists approach the integrated analysis of climate 
mid the economy over lime. It also indicates thai no 
single economic policy can he considered in isolation 
from other government interventions in the economy 
Wfuii the discussion fails to convey, however, is the 
scale ol the uncertainties which economists admit to 
in this aiea of investigation. 

Guskins & Weyant (1093) offer a vivid, if perhaps 
ciUremc. illustration of tins uncertainty, In tiieir 
eom|xarjson of 14 "DICE-type H models and 13 climatic 
scenarios., estimates ol the average carbon tax required 
I'vci lire period 2000-2020 to achieve, for example, u 
20 % reduction in carbon emissions {compared to 1**90) 
\nry fnuii $50 per ton to $330 per ton. In fairness. 
extreme estimates tend 10. flow fiom extreme 
assumption underlying alternative models and 
lundamental aspect ol mot let development is to identily 
the potentially sensitive areas and to refine the accuracy 
and level of agreement regarding these area 1 *. 

Otlter policy evaluation exercises, in many ways 
simpler are undertaken by economists, including Ihe 
pievmusly cited approaches using the OKANI model 
This model is a highly disaggregated general 
equilibrium model o\' output, employment and 
eon.sumplion in the many diffcicnt sectors and regions 
of the Australian cconomv. It would not "be 
operationally feasible to construct a DICE model at 
this loci ol detail, but what ORAM can do is predict. 
lor a given well-defined policy initiative, the likely 
miero-ellecls across Ihe regions ol the economy, To 
know in advance, for example, that a particular section 
ot the Adelaide metropolitan region is likely to 
experience significant unemployment because of GG 
policy measures, ctin be of significant use to social 


The capacity ol economists to provide clear 
predictions of the costs and benefits of climate change, 
or of policies to modify likely fulure climate changes 
is limited by u number of factors, not least the 
uncertainties surrounding predictions ol climate 
change. The work dime so far is slight and highly 
speculative, but it has grown - and in collaboration 
with our colleagues in the physical and health sciences, 
ibr example, it could further expand in extern and value 
to the policy process - especially as the world works 
its way out of the problems of recession thai have 
thoroughly captured the attention of policy -makers in 
recent years. 

The significance ol collaborative approaches cannot 
be over-emphasised Especially in areas vvheie extant 
knowledge is very limited, and subject to a very high 
degree of uncertainty, it can ensure that research and 
modelling investments by different disciplines are 
mutually supportive Because of the nature of their 
'"business", economists bring not only analytical and 
modelling skills to the consideration of issues such as 
climate change, but also a sharp focus on what is 
"policy-relevant" information and analysis. 

Contrary to the belief of many, K not most, non- 
economists, economists do not believe that private 
sector markets always work welt and provide the right 
signals and incentives But their scepticism about 
private markets applies equally to political markets 
where lobbying has a powerful role, and where 
regulations can become captured by those they are 
supposed to regulate. An early keen eye lo these issues 
can result in policy debate and design being locussed 
on achieving better ultimate policy outcomes at lower 


Climate Cuvscr GRotr* (I992> "Chinee change scenario:, 

for tile Australian rcgi>w,< CNIRO Division of AiinoxphcriV 

Research. Mordialloc). 
<J\skins. O W ft WrvAvr. J. P. (1993) Model Comparisons 

of the Costs of Reducing CO : Emissions Anu'titalt 

honomii' Rt'vit'w 83(2), 318-323 

HOCOIHON. J. T , .IkSKlKS, G. J & Et'HR.V_MS, f, I. (Ms) 

(i*J90) "Climate Change, The 1PCC Scientific Assessment" 
(Cambridge Umversily Press. New York). 

HmuiiNs, D,, Gross, R , Wif.sov. B. A Stirpm«o. ?{#??) 
"The economic unpad of climate chance on ugflculfune 

in South AiislraJiii. (Department 0! Environment and I .and 
Management . Adelaide) 

(C (Inihsiuv Commission) (1001) "Costs and benefits of 

reducing kirceutiouse yus emissions" (AGPS,: Canberru). 
Noudikus. W. D. (1993) Optimal greenhouse pas reductions 

and tu* police in the 'DICE" model hut-tit an Lamttmtt 

Review te\2i 313-317. 
Piarman. G I. (WK) "Greenhouse - planning lor elimUW 

change" (CSIRO Division of Atmospheric Research. E, J. 

Brilf Melhourne & Leiden). 
Rullv, J. & Hoiimans N. ( [993 1 CliiiiJic rtian-ie ;»ld 

agriculture: 'fhe rote of inlcrnational (rude itiH'tieun 

Economn Review 83 (2j. 306-112 
|omI:T. J.. REEiXfc J. & KANt. 5 (1992) Economic 

implications ol* global change for world agriculture. Mwnuil 

ttj Agrlt ultuial title! Re\i>urt>:\ F.n>tnft?Tti\ 17, 195 205 


By A. J. McMiCHAEL*t & M. Y. Beers*± 


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 injury 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. 

thltmtttwm <>/ lht fanvt Sitarry >>f S, .<Imav. (1994). IWiO. 91 y& 


by A. I McMichaix*-! & M, Y Beers** 


MrMiemn , A. I & Blcrs. M, V. 1 19941 Climate change and human population health: global anil Suuth 
Australian perspectives. Trews. R Soc S. Auxt UH(1.) 91-98. 31 Mh><. 1994. 

Public health scientists now need 10 think within an ecological framework because of planetary overload by 
llie human species, Climate change must impact on human health Predicted huiilth btumds include increases 
>u thermal eoiemes which impact particularly on the very old and the wry young. Natural disasters such an 
cyclones, floods and ocean storm surges wilt increase rates of uijuty ami death, Inxect and otiici vectors tin intecnous 
dist.ta.scs will t hange geographic distribution resulting m changes in the demography of diseases such as malaria, 
yellow (ever, dengue, encephalitis and cholera. Changes to agricultural productivity will >*ltet food availability 
M Hirtoy regions Rising seas would, amongst problems, disrupt sewage disposal causing diarrhocal diseases, 
t'lmiate-inducvd environmental disruptions would create "ecologieal refugees' leading to spread 01 'infectious diseases 
and social disorganisation. Ozone depletion with increased cK|Hthuic to ultraviolet radiation will lead to an increase 
mi skin CiiiieeiN. eye disorders and immune suppression. The significant environmental degradation in South Australia. 
together with climate change could lead to significant inrreakct» in water pollution, outbreaks ot Ross River virus, 
Murray Valley encephalitis, dengue fever, rick borne dtemes and possibly Hania viru>c< together with jrastrointcstiiiaJ 

Ki.Y Wouos: global environmental change, human ecology, human population healili. 


During its quarter million years ul existence. Homo 
wpietis has adjusted lo seveial shifts »n the prevailing 
paililc of environmental health hazards. Each shift has 
coincided with culturally -driven chatigcs tn (he social 
organisation and ecological relationships of the human 
species: the advent of agriculture, urban sertlemeni. 
industrialisation, and the .spread of *'alilttent" Uvmg. 
Today, another eatcgoiy of environmental health ha mo.1 
is emerging that is qualitatively dishnci From earlier 
ha?ards and may well have wider-ranging health 
consequences. This hazard arises From the plantctary 
overload resulting from the cumulative impact ol the 
human species, via population growth, land pressurcs v 
energy-intensive technology and a high output of waste 
gases - including changing the gaseous eontposilion 
ot [ftti lower atmosphere and, hence, its heat-trapping 
propenies (McMichacl 1993) 

Appraisal rjC the health hazards of these incipient 
global environmental changes requires us to think 
within an ecological framework Public health 
scientists have not heeded this framework previously 
because, until recently, humans were (appai Cully) 

Department of Community Medicine. University of 

Adelaide, South 'Australia 5005. 

Cunent address: Department ol" Epidemiology and 

Population Sciences". London School ot Hygjeoe and 

Tropical Medicine, Keppel Street. London. WOE 7HT. 


Current sddrcv Communicable Diseases Control Uqfli 

Epidemiology Branch, South Ausltalhtn Health 

Onmmivjmi, Adelaide. 5001. 

living within the biosphere's carrying capacity. Now. 
howevvr, then W* signs thai our aggregate impact is 
reducing Ihe ^lability and productivity ol various of 
baith's natural life-supporting systems. Each of these 
individual pmblem areas is already a Familiar subject 
of en 1 , ipjnmcntal research and debate: greenhouse gas 
accumulation* stratospheric ozone loss, laud 
degradation, aquifer drawdown, depletion of ocean 
fisheries, loss of biodiversity, and the ecological and 
social consequences of rapid urbanisation Bui their 
wider, collective, rcolog'uul signilieancc lot human 
population health has been rarely considered 
There is much that is uncertain - and 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 ol gteenhouse gas accumulation upon 
the world's climate. The consequent difficulty for 
health researcher's is knowing whether and how lo reap 
further uncertainties about health outcomes upon the 
underlying uncertainties about global -change 
processes. Nevertheless, since sustained overload of 
the biosphere must, logically and eventually, entail a 
reduction of its life-supporting ability, it is a reasonable, 
and prudent, inference that this poses a fundamental 
hazard to human health. Estimations of health unpad 
must therefore be attempted 

The best -documented and most discussed aspects nf 
global environmental change are those peiiaituug to 
climate change. "Climate change" has Keen widely 
taken to include both greenhouse enhancement and 
(WCme depletion. Purists may argue* that 'climate* is 
a iroposphei ic phenomenon, atld has to do wiih 


\ J, MrMKIIU -.1. & M V BEtfttS 

temperature, precipitation, humidity ;*nd winds; since 
nn>ne depletion i 1 * occurring in the stratosphere, and 
its main anticipated consequence i*> an ine reave in 
surface-level ultraviolet irradtatium it is iuiI pan of ihe 
climate change However, the demarcation is not 
clearcul Ihe re arc various inteiyctive and olhct 
relations between diese two processes, For example. 
chloiofluoroearbons (CFCfi) contribute m holh 
processes; tropospherie warming may enhance 
Mtatospherie COOllflg; and ozotte depletion may affect 
the radiative forcing properties of Ihe airnosphcre 
overall, in thts papei we use the inclusive approach. 

Two olher ^ttatHyiflg comments are needed here. 
First, specifying Ihe adverse healih effects ol climate 
change rests largely un extrapolation and. in some 
Ljsex. reasonable conjecture iJvlcMichael 1093. 
Maskeli 1993) Unlike the diavilv toxicologic*! 
hazards posed by localised environmental pollutants, 
this category ol healih hazard does not readily permit 
the empirical observation of health impact in one 
population as a basis \kw estimating the risk in other 
populations. As moiv information about these complex 
systems accrues, the modelling ol population health 
impact* vvill improve 

Second, it is generally diflicull In make predictions 
Of localised effects ol climate change upon ihe health 
of South Auslialiaits hot certain, simpler, processes 
— such as skin cancer risks due to incieascd ultraviolet 
irradiation (llVRj and the change in mortality due to 
increased frequency of heatwaves local estimates 
can be made, but for many other processes fc g effects 
upon regional and global food production) the health 
of South Australians will reflect their sharing of ihe 
experiences of wider populations. Therefoie. this pujier 
concentrates initially upon the general possibilities of 
population health impact, alter which Mime more 
specific comments are made about the South Australian 

Greenhouse ^ases and climate change 

Vmious natural 'greenhouse' gases in the lower 

iilniosphcre uhsorb much of the heat that is m radiated 
Irvm the Earths surface. Fossil fuel tomhustion and 
accelerated forest clearance since INLlO has increased 
the concentration of carbon dioxide Irorn 275 ppm to 
36(1 ppm. Concentrations of other anthropogenic 
greenhouse gases, especially methane and the 
chlorofluorocarhons iCHL's). aie aJso increasing (1PCC 
W2: F.pslein 1992). 

The United Nations intergovernmental Kanel <»n 
Cliiinate Change (1PCC) estimates lhal, in consequence 
ol the increase in radiative forcing, the average global 
temperature will rise by around 2-3 °C by die year 2100 
(1PCC 19^2) While this estimate is acknowledged to 
he uncertain, the evidence is firming that huiua.ii 
activity ls now warming rhe Earth's surface (Lncis & 

Carlson 1992;. A rapid global temperature increase 
en* several degrees would be without precedent in all 
ol human hiMoiy. and would pose a langc of risks 10 
human healih. both by direct and (probably more 
importanll indirect mechanisms <!.asl I99J). Ihe 4-5 °C 
increase in temperature after the more recent ice age 
which iscvcral smaller II actual ions aside | oeeurrcdover 
several thousand years, caused major ecological 
changes - including the spread of forests and. together 
with intensified human predation. the extinction ol 
many laigc mammals: ii also presumably contributed 
to Ihe onscl of agriculture 

The most likely impacts of greenhouse-induced 
climate change arc summarised in Table 1. 

Taih i. !. Muin [tirtiotioi tjffetty «•/ (Ifibttl dimate change tiii 
pupuluHon health. 

Direct Increased r^uency -i) heatwaves __ deaths. 

jllnew. injury, 

Wespiratotv dicers of moisture, duns., pollenv 

Climuu. instability _ dtwiMcts (eyeluuev 

floods, tires). 
Indirect Altered habitat and li^n<ini.vflOri id veetm 

home infectious diseases O.dher contapmus 

disease* obm. 

Impaired crop production I %oi I temperature. 


Sea-loci rise — inundation, janiuuion. 

suliuii>. CtC. 

Demographic disruption. pnvirnranftrilill 


The "lain direct hazard to health from a temperature 
increase comes fiom iheimal extremes. It is predicted 
thai rhe frequency of heatwaves in temperate and suh- 
tropical climates will increase (Brasseur 1991) - mr 
example, a doubling in the annual number ol days over 
1#°C in Washington, DC, is predicted over the next 
half century (Leaf 1989). Il vs not unusual m temperate 
/ones, sueh as South Australia, tor three to four d;tys 
each summer to record temperatures ervatcr than ^X°t'\ 
However, with an average annual teiiiperatute increase 
of 2-3°C. the nurruVt of days in excess, til 3S°C is 
predicted to rise (Hwan et at. 1991). 

In icsponse, death rates among the most susceptible, 
particularly the very old and very young will rise 
(McFarlanc 1978: Kilbourne 1992) Causes of death 
to rise would include heat-stroke, exhaustion, 
cerebrovascular stroke and. probably, accidents and 
acts of violence. On the other hand , winter deaths from 
inllueu/.a and Irom hypothermia - particularly among 
the homeless poor fWMfl DcHtl to London to New Ybqrk 
— could be expected t0 decrease. 

Weather patterns ate likely to he destabilised, with 
more frequent cyclones. Hands and oceun stonn surges. 
These "natural 1 ' disasters would increase rates of injur) 
and death They would also disrupt agticultu re. local 
transport and sanitary engineering. hringe dwcllcr\ in 



Oimsv Ijtnwii^g or exposed co rivers mid oceans. tt* in 
coastal Bangladesh, will be the most vulnerable 
(Houghton 4 oi > W901. In Australia, il is predicted 
that cyclonic activity would extend down the eastern 
coast ftoin !l littlWrt? bound ol 27°.S l« M°S (Australian 
Bureau of Statistics W&Z) However there we unlikely 
Ui he consequences I* tr Australia's southern coastline 

Various insect, vertebrates ami other vectors for 
infections disease will change their geographic 
distribution (Cook 1902; Shope Wl). As highci- 
latiiuck: loeattons become wanner and wpften 
mosnuitocs' will tend to spread ihcic. As wvll as 
extending Iheu spread and seasonality in tropical 
counti ics. mosquito-borne infections (malaria, yellow 
lever, dengue, and various tonus of encephalitis) may 
remit) itKlirrcnlly unaffected lemperale areas. (Ancient 
Rome and eajflj fcuiope weie malarious; so, earlier 
this century, were puns bribe Uniied States, L.ngland. 
Italy and northern Australia.) Vector-borne diseases 
spiead by flies, water-snails and vertehrnic animals are 
also Itkelv io change their distributions. Recent 
evidence iliat cholera is *voetgr>b(trn£ by raking rvluge 
under the mucinous t niter coat of aquatic i:lgac 
imeludiuy species found Ifl both fre<h and coastal 
waters!, suggest thai its spiead may also be assisted 
by warmer oceans i.fcpstcia 1992) 

Ijing-tenm increases in background temperature mav 
reduce the yield of some of the world's most productive 
gram-growing temperate regions the American 
praiites. western Luinpc, the Ukraine and couvul 
Australia while increasing (lie agricultural 

productivity iYt other regions (northern Kurope and 
Canada* {IWTC 1990). Crops, that have been 
selectively bred for specific cn\ ironmeuis. may fail to 
germinate. Climaic change niay also alter habitats lor 
ihcne insects, fungi and micro-organisms thai cause 
diseases" oi grains, fruits and vegetables. While 
increased levels of carbon dioxide in ihc atmosphere 
may enhance plant growth (especially the evolutionary 
oldet "€}" plants), most current evidence indicates 
lhal. on balance, this would nor be a significant eJleci, 
Further, while the overall net effect ol climate change 
upon world agueuhure is uncertain (Parry 1990*. 
fmsson 1989) at a regional level rood security may 
be seriously threatened - especially in pomvr 
countries m the semi-arid and humid tropics Climatic 
change would also perturb the growth and productivity 
ol livestock and I'ish. local fuclwood supplies and the 
,n liability of freshwalei all important determinants 
ct health. 

A rise in sea levels is predicted from thermal 
expansion of the ocean. The IPCC estimates a rise of 
approAimately 0.65 metres by The year 2lltf\ with 
serious c fleets upon low-lying coastal communities. 
The moM vulnerable settlements arc those in crastnl 
hnvltftds such us in Bangladesh, China and Egypt, and 
m wiall island nations. Inundation and storm surges 

could lead to significant displacemem ol people 
iMaskell fftiL IW: Worresl & Grant 19K9). Risiue 
ijcas would also disrupt sewage disposal (thus causing 
diarfhoeal diseases), cuwe increa.scd salinity ol cuaslal 
freshwater estuaries and farmlands, and harm wetlands 
and the breeding grounds of marrv otxsin fish (Waincs 
& Fuchs 19911 

Many ol these climate-based environmental 
disruptions would etcuie "ecological refugees" 
(Jacobsoii 1989* Dispossessed people from aieas 
damaged by floods, cyclones, using seas oc droughts 
would migrate to the relative safety of ewes. Shanty 
towns would proliferate and so would infectious 
diseases and social disorganisation. It is conceivable 
if climatic and agricultural and dentogiaphie circum- 
stances deteriorate significantly in the coining century, 
that Australia will havean uilliu - voluntarily or even 
uncontrollably - of refugees from Southeast Asia. 

D/otu* layer depletion 

Stratospheric ACTOR which resides at IU-20 km 
altitude, accumulated over several billion years 
principally as. a result ol Ihc evolution ol oxygen- 
releasing photosynthesis. This o/onc "layer" provides 
life on r-arth with substantial protection from 
biologically -damaging I'VR (particularly the higher- 
energy UV V which is entirely abxwhed and the 
medium-energy UV-B which is largely absorbed), 
lodav. the stratospheric o/one is being damaged by 
human-made cltloroiluoiocarbons <CFC*>) and oiher 
industrial gaseous emissions. 

At mid-northern latitudes (30-50°N) the o/onc layer 
thinned progressively by around 0.5% per year during 
ihc 19KUs and caily 1990s (Stolarski 'ft ai 1991; 
Blti&scur 1991: L'NHP 1991). Overall, during lhal period 
(here was an accumulated 8% loss in winter and x\2% 
loss in summer Meanwhile, in the ^lutlicm 
hemisphere, significant cumulative ozone loss ocx-urred 
during the 1980s from the pole "down" to latitude 30°S, 
which includes the lower parts of Australia arid ol 
South America, There now appear to be well- 
established, strengthening trends in ozone depletion 
in both hemispheres The anticipated increases m UV 
irramance at ground-level would be greatest JH the 
SOUthcrti pails o\' Africa, Australia and South America 
and in the mid-latitudes (30-oO°N) in Hurope. Asiaarul 
North America (Madionich I992). 

The predicted direct elfecls ol increased human 
exposure to L'VR include inereusev in skin cancers, 
eye disorders, and suppress! \c cflfefCS upon the immune 
system. The UN Environment Pmgramme csltmatcs 
tluit for every lv<' dectease in ozone, there is an 
approximately L$S uierease mcvposure to ultiavtolci- 
B (UV-B) radiation at rhe Earth's surface, in turn, each 
sustained \% increase in UV-B would cause an 



estimated 2-3% increase in the incidence of skin 
cancers, predt.miina.nil> non-melanoma cancers. 
Increased UY-fi exposure is also predicted to increase 
the occurrence of cataracts and of pterygia (tissue 
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). Although there has been 
frequent reference to these various predicted health 
outcomes, the estimated increases in incidence are 
based on rather sparse data (Armstrong in press). 

During ISW, an attempt was made in Australia to 
predict (he number of extra I. VR-induced cases of skin 
cancer — basal and squamous cell carcinomas and 
malignant melanoma - and of ocular cataracts and 
pterygia that would occur over the next tour decades 
(Frascr et ai 1993). The calculation entailed, first, 
estimating the dose-rcsponsc relationship between level 
of UVR exposure and the incidence from existing 
epidemiological data. Then, from knowledge of trends 
in ozone depletion and ground-level UVR during the 

preceding decade, and from estimates of future trends 
in ozone damage, predictions were made of changes 
in ground-level UVR over coming decades. From these 
estimates, and appropriate amplification factors, the 
numbers of ex tru incident cases were predicted. The 
predictions for South Australia are shown in Fig. I. 
(Note., however, that the techniques for making such 
predictions arc still crude, and, in this particular case, 
there was inadequate allowance for minimal duration 
of exposure and for cancer latency periods. These 
predictions are therefore illustrative, not definitive!). 
Increased I'V-B exposure would also have 
deleterious effects on the world's biota, including 
impairing crop yields (Worrcst & Grant 1989). The 
estimated danger to marine organisms could be 
significant, since increased UV-B irradiation of the 
ocean's surface could damage the photosynlhetic 
phyioplankton population the basis of the aquatic 
food chain (Smith fi a! 1992). (A quarter of the protein 
supply in the human diet comes from the sea, as does 
much of the protein ted to livestock.) 

amplification factor 


w^*sp «*» j*^ *M0'ka# c 





1 7 













3 1 



i I 


y A 

6L m 


% MS 





Fi^i, I, Estimylcd annual extra numbers of cases oi skin cancer and ocular lesions due to increased grnund-level exposure 
to ultraviolet radiation, consequent upon stratospheric o/nne depletion. Numbers are expressed relative to the (background) 
incidence of ihese conditions in I99U (from; Eraser « nl. 1993. Note also the caveats m the accompanying text.) 



South Australia 

Hcutheote 1)990) ItiU purioscJ dial climate change 
should no< be considered in isolation from climate 
variability 4v the taller may prvrvuJe valii-hk- 
iufoimation on which lo base estimates of change. An 
examination ol (he climate and ecology ol South 
Australia provides particularly interesting perspectives 
ol such variation with which to consider the possible 
human health effects of climate change. 

Australia as a whole and South Australia in particular 
has out* ol (he most vanable climates on earth. Tins 
is no new phenomenon but has been (he case tor 
hundreds ol thousands trf years (Ntcholls 1992; Cook 
1992}. This predictable but irregular variabilis has 
icsuUed in the adaptation of the flora and fauna ol the 
State, to suit such changeable conditions. 

Covering one eighth of the Australian continent, 
South Australia lies wholly in the temperate /.one. The 
land is low. with the inland areas being largely covered 
by plains, sand and gibber desetts. X()% of the State 
is less than 300 metres above sea level, The climatic 
consequence is lhai apart Iroin lite coastal areas, thcic 
arc no sij*mlteant mountain range:* lo encourage 
precipitation evenly throughout the State. Much of SA. 
therefore, is arid or semi-arid (Gardner I99t). 

Weather never replicates itself exactly from year io 
>ea! (Voice tt al. 1993). Australia has l>een found to 
h**rve the highest interseasonal rainfall variability and 
runoll m the world (McMahorr tt al. 1987) with 
drought, being a frequent occurrence when riefimd bk 
annual rainfall in the lowest 10% of records (Gibbv 
& Mahet 1967). SA js the driest Stale with fuut-fmlLS 
Of tbC State normally receiving less than 250 mn 6| 
ram annually while only Kim: thud ol Australia a*, 
a whole receives less than 250 mm The basic features 
of the SA climate are hot. dry summers with relatively 
niUd nights and cool but not severe winters, where most 
of the rain falls between the months ol May lo Atn/usl 
(Gardner 1993), 

tt is clear thai changes to the rainlall pattern alone 
could diaiiiatieaily ultei the ecology oi this State. 

Not only is tiie geography of the land and natural 
fluctuations influential in climate variability in SA hut 
a phenomenon known as JiNSO, of llie til Nino 
Southern Owittaijon plays a significant role in the 
climate of South Australia The Southern Oscillation 
is a major shift in air pressure between Asia and eastern 
Pacific regions which ellects changes to trade winds, 
cloud amounts and rainfall over the tropical Pacific 
through to Eastern Australia f Voice et al 1953: 
Nicholls 1992). extending its influence as far west as 
the River Nile (Quinn 1992 j. The Southern Oscillation 
is measured by an index Of the difference in air pressure 
between Tahiti and Darwin, A posnivc index (low 
prcssuie at Darwin) means that tiade winds blowing across the Pacific teed moisture into ttie 

monsoons of Asm and Australia. A negative index (high 
pressure al Darwin) means thai llie tiade winds are 
weak or even reversed . 

A significant wanning of the sea surface in the 
eastern equatorial Pacific occurring at the same time 
as negative index measurements ot the Southern 
Oscillation, is known as El Nino The reverse effect, 
known as La Nina, takes place when the Southern 
Oscillation index is positive and the waters of the 
eastern Pacific are cold. The combination of the 
Southern Oscillation and l : .l Nino is known as the Ul 
Nino- Southern Oscillation or UNSO 

This is noL a recent effect. Palewliuialie recoids have 
uncovered evidence of the PNSO as far back as 1000 
years (Cook 1992). It is predicted that UNSO in 
unlikely to change dramatically or vanish under 
conditions brought about by the level of climate change 
current!) predicted (Voice «•/ al, 199T), However, it 
is not yet known whether b'NSO may become variable 
or more frequent, destabilising the present 12 month 

At piccJit, r.NSO accounts tot about 90% of utajoi 
droughts- in Eastern Australia Further, this same 
phenomenon is responsible lor appioximatcly 20% ol 
all rainfall variations in eastern Australia including 
floiMJuig. wc! seasons lasting sex to nine months as well 
as less ijtamatic events (Voice et al 199^), tt is 
therefore a significant contributor lo the extreme 
variability in the Australian, and South Australian, 

Over the pusi 200 years ol white settlement, human 
activity in adapting and munipulatinj: environments to 
provide for hotter living standards may have increased 
human vulnerability to ttieettccls ot sigrnhcant chmattc 
and environmental variation (Hcathcote 1990). In other 
words, human adaptation has been in the form ol 
modifying the external environment as opposed to the 
lunger evolution ol native Mora and fauna which modify 
then own physical chaiactenstics and systems. 

Human attempts at environmental modification m 
Soulli Australia haVe included extensive land clearance 
resulting in desertification and increased salinity of soil 
and water. Deforestation ol eaiehmem areas has led 
Bp Hooding and erosion, and agricultural piaeitees 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 Mows, In recent 
years, significantly lower water levels and reduced How 
in this State, combined with pollution with phosphates 
and nitrates, have resulted in extensive algal blooms 
(Soong 1993) The Murray has a fall of only 20 metres 
across South Australia (Gardner 1993) and U therefore 
not a last- Mowing mvr, aiiUVnng it MjvtcpUtrfc to algal 



Within the conic* 1 & SVlch significant environmental 
ifegradalton, attempts must be made to ptcdiet ami 
ptepaie for human health needs in a climate change 
scenario where avviage global surface temperatures arc 
predicted *> rise b\ Mound ? 3°C by the year 2100 
■ 1PCC 1992). 

(>vet thousands of years of climatic variability. 
Australia's flora and tauna have evolved lo aceonumxlatc 
the climate, and indeed to flourish. NichoUs (1992) 
describes adaptations in Ihe behaviour and breeding 
patterns of the Red Kangaroo, the long-haired rax and 
other Australian binis, inlets and plants, All have 
developed survival patterns of breeding in plague 
proportions in wet seasons when food is plentiful and 
a virtual shut duwn m tinier of extended drought. 

More, permanenl increases in summer rainfall and 
higher minimum temperatures in SA as predicted by 
climate models (Heathcotc 1990) could lead lo plagues 
unhampered ty the naiurai controls of diougbt and cold. 
In die past 12 months, excessive rainfall and milder than 
usual minnriuni temperatures have led lo plagues ot 
uuce T locusts, various bird species, wasps and 
mosquitoes. Floods have created disaster and death in 
human populations. Water supplies have been cunflami 
rutted with human and animal sewage (Walters hW>. 

The human health consequences of such 
meteorological variation have included a SfSUth 
Australian outbreak o( Ross River Vims (RRVi ol 
epidemic pnipurtrorrv Tlteie weic in excess of 800 CfiSCS 
of RRV in the period mid-September 1992 to Mid -May 
rW iCaniemn I99.V). By comparison, previous lesser 
cpidcmKS vmctc in I9K4, 136 cases and 198ft U$ cases 
Oflfeinsdfcw IWl.i Tlve underlying annual mean number 
of case?, over the last ten years (excluding epidemic 
years) is le.v* lhan )0 cases per year (Bell rt id. W9I). 
The I992.'V3 epidemic occurred as & result of high 
rainfall ami nuld letrtperalurcs rmm August 1992 to 
February 1993 as reported by the South Australian 
Regional Office of the Bureau of Meteorology (1993) 
which provided perfect brrolijn' conditions lor trtosquito 
vectors Atdex \\$ila.x and Cuiex imnulirnstris. 

'W.uihKs, R. Q99QJ Personal eoiimtunieaiion, Stale Wmrr 
Ubonitary, Sot*h Awtralin. 

-C*Mfc*os. S. (199.1) Unpublished data Commtimeabtc 
Discxves. Control Unit. Epidemiology Branch. Public ami 
i-titironntcnuU Health .Service South Australian Health 
( :ommi*MCHi. 

'Sithiumt, R W (I9W) the Ukely Impact ol nim.ite 
Ctunge on Medical Mvtor* H*pet presented *• the Australian 
Tropical Health and Nutrition Conference. Brisbane. 

'tUiwy, R. D,, Hudson, H. J., .Shai'kiin, U. K. & Wilis, 
M C 0993) Evidence tor mi liyfigcwwnj form d i.ynw* 
BtitmtiBBfe in Australia. Riper presented at tin* AusImUinii 
tropical Health and Nutrition Conl'urenec. Brisbane. 

\>owswako. K (1993) Disaster management. Report to the 
7iii Standing Coturmtlce on Industry, Science. lecrtnotagy. 
Transport, Communication and tnrraslruttun: Wtktinft* , SA 
Wed 13th October 

Diseases predicted lo uicreasc include Mutray Valley 
Encephalitis (MVE), also known as Australian 
encephalitis, Caused by a mosquito home Muviviras 
MVE was last recorded in South Australia in 1974 
P.xeessive rains as a result of an MNSO event had led 
to an extended breeding season for the heron population, 
the natural reservoir of infection lor MVP., The viruv 
is transmitted via die mosquito vector Attics aegypM 
and has a case fatality nae from 0..V& to HY% (Henenson 
19901 and is often around 30% (Manson-Bahr N87i. 
Nicholls (1986) subsequently devised a method, based 
on historical research, of picdicting outbrealcs of MVE 

using ,r * Pwvw mean hit pnvfturt aTtnc Suuthcn 


Oilier predicted intpaet* ol cluuaur change mi South 
Australia include the move south of dengue t'cvr lkr3c 
borne diseases such as Lyme disease, and even malaria 
(Sulhertf W92\ Kwznrtul. I993| In Ministry 1993, 
local transmission of falciparum malaria is reported *> 
hiive occurred in Oueeflsland, snutb of the nineteenth 
parallel (MunaySnulti A ^i-insteio ]993). Although 
Australia was certified iicv of endemic Malaria by the 
World Health Or#misahott in WKI, there ate over 700 
cases of imported Malaria annually, The common 
\-ectors for malaria. Ihe Anopiirlt's jaruuti s.L and An, 
punctukitus mosquito are considered lo exist only above 
the 19th parallel, known as Australia's malana receptive 
zone. However. An. jaruuti in known to range- south ol 
the 21st parallel, while An. anruillpes, previously im- 
plicated in local transmission ol malana in Victoria and 
New South Wales (Ford 1950). ranges as &r south as 
Tasmania (Russel l l )90). 

The implication lor Stuilh Auslr.dia is tl«l widt 
climatic change. Ett least An. amwtipes is cjipuble pf 
estabhshiiii? a niche here, becomine itilecled Vr$ biling 
cases which are at the gaxnetocytaeovic slage or" the 
disease and transmitting the disease locally. 

The spread of Lyme disease eould occur because ol 
increased vegetation due to highei rainfall in South 
Australia Although no vector for the anaerobic 
sptriK-liactes of the Boncllac parasites, the causativv 
agent for Lyme disease, has yel been identified in 
Australia (Russel cr aJ. IW3), there is suggestiw 
evidence from serology reports from patients with Lyme 
type symptoms, thai indigenous Lyme borreliosis (LHl 
is widespivad in Australia and thai a number ot strain* 
of the disease are present (Barry rt nl. 1993**). 

Fethaps more fiequent mouse plagues would bring 
with them new and serious diseases such as the Hanla 
Virus which emerged in the US (his year (Leduc e/ at 
1993; Motel \W$). McUkkIs of eoutioHiugsueh plagues 
are ad hoc at present, expensive (SI.6rn.) and largely 
inappropriate for long-term use, The use of poisons, 
such as strychnine in the 1993 South Australian mouse 
plague (Downward I993 1 ) is not a viable long-term 
response K> :h i tilet ns diat liave llieli otiguis t» ocologlcal 
and cJimAfic disruption. 



Coritaimnation of water supplies has led to anecdotal 
reports of increased incidence of diarrhoea! and 
gastrointestinal diseases, '('here have been 1250 eases 
Of Hepatitis A or infectious Hepatitis in South Australia 
since 198*. with MO of these occurring in I9K6 (Hell 
tt al. 1991). Hepatitis A is a disease of public health 
importance in flood situation* or other disasters which 
jffect infrastructure, Water-borne diseases such as 
yiurtliasis (caused by the pioto/ouri tiiardia kunblia), 
Cdwpxhdntnt't jejimi (found in milk. wiiIct, meat) and 
amoebiasis {EntamiH'hu histtttyticu) could also pose 
problems in a wetter SA, while increased temperature^ 
may lead to more infection with Lfyjonclki and fotid 
borne disease such as Salmonella and Yersitwia. the 
latter begftfl already on the increase and ol significant 
public health concern (Bell et ol |99?J Ltgkwvlfa 
bacteria survives belter m water aerosol which contains 
Wuc-grccn algae (O'Brien & Bbopal 1993), this having 
significant impliealions lor the transmission of the 
disease in a warmer climate scenario. 

II is predicted that with global surface warming of 
2-3°C the main direct hazard to health front a 
temperature increase comes from ihenna! extremes. 
South Australia endured temperatures in excess of 4i)°C 
in February 1993, provoking media headlines as fai 
away as Brisbane of "Heatwave kills && hundreds in 
hospital" fTJw Courier Mail 1993), while in Adelaide 
it was "Heatwave drama as toll rises" (The Advcrttscr 
1993). Children, the elderly and Hie unwell weic the 
main groups affected. 

Finally, SA currently enjoys up to 3,51X1 hours ol 
sunshine annua My, out of the approximately 4,380 
totift possible (Ganinei 1993). Increasing levels of 
UVR pose n worst case scenario of not only 
significantly ttteteasmg skin cancer rates, but also oi 
initiating immunosuppression, rendering the population 

more vulnerable to various infectious agents. Evidence 
from transplant patients, the irnmum compromised and 
AIDS patients illustrate well die range of opportunistic 
infections which may result from sweeping population- 
based immunosuppression. The efficacy of 
immunisation may be reduced, and diseases whieh have 
been "controlled" in the past may therefore re -cmcigc. 


One result of the increasing impact wHIonm sapiens 
on the world's natural systems is that our focus oi 
environmental health concern must move beyond the 
realm ol polluted local environments ami into I he ivahu 
ol disrupted natural systems Anticipating effects on 
population health is made difficult by the combination 
of urilamiliarity, uncertainty and tmuny (Ihe 
unacccptability of an empirical "^wait-and-see" 
approach). The risks of cancer and cataracts from 
ozone depletion can be easily appreciated; likewise the 
health hazards ftom heatwaves. More difficult to 
Cam ephialisc but of potentially much greater impact 

are the anticipated indirect health consequences ot 
climatic effects op food production and on the spread 
of infections., and, perhaps, immune suppression b> 
ultraviolet radiation. 

Some extreme, and extremely costly, technological 
adaptations may he possible - such as erecting 
ultraviolet barriers over living spaces or investing 
massively in seawater desalination. However, in 
general, human population he-aim cannot be sustained 
if the Earth's natural systems are not maintained Tor 
the first time in human history, there is evidence that 
various of those natural systems are becoming over 
loaded at a global level This portends a frameshift 
in how we must think about "environmental health* 4 . 


Akmsthong, B. K. (in press) Implications ot incnsxmi-wlur 
II VB for cancer incidence //(Chimin M-l-thl I The Role 
of the Stratosphere in Global Change' 1 NATO Advanced 
Study Institute Scries- (Springer wafitt. Heidelberg) 

Australian BtmcAi. Oi -Statistics 0992) Auslruti&V 
!;nvironrrtcnl. fcttlex and fettn. AUSU4. Canberra 

Bin, 1, t'AMiKON, S.. P.stikman, A. McCaiil, K. . 
Pfta-.sT, K * Wi iNSTrm, P <190i) Soiith Austral im Kcalrfn 
Statistics t'hartbouk. Supplement 2; lidectioiiH and 
Notifiable l>iscascs (ttptdcminlogy Branch, South 
Australian Health Coiimil*h|oii, Adelaide). 

BtNtNsON. A S (Ed.t (1990) "Control of Coimniiiiiciiblc 
Diseases in Man'. (American Public Health Association. 

BkAssi:uH, G. P. (199t» Ozone depletion. A deepening, 
broadening trend. Nututr 352. 66M 

COO&i f" R (1992) I King |ftt? rinjts to sludy pijst HI 
Nino' Southern Oscillation influences on climate, pp, 
20t 214 In Diaz, H R & Markpraf. V. (lids) "El Nino 
HistoiHrtl and Aloof limaiic Aspen ot the Southern 
Oseillaiiorf (Camhridge I'nivemiiy Pro*. Canihiidiv) 

Cook. O. C, (1992) Effect ol' gJOtml wantimn on the 
distribution of parasitic and other infectious diseases; a 
review. J Ho\ Sol. Mai »5. 68«-G91. 

( kosson, R M989) Greenhouse warming and climate 
change, why should We calf' tinul I'nUt y 14. 10'Mltf. 

LrsniN, ft (ivvi) Cholera and the environment f Letter > 

Intuit 3.19, IU»7tllHH. 
KWVJ, C, RfcV/vKT T # CMVtftT, I) (1991) HCHMh 

Implications of Long Term Climatic Change CAQrS 


Poro. K (19V)) The Malar In piohlmi m Australia and die 
AuMrjiUiiu PitediL TcinToiit's ,\Uui .1, Afl&r. 1, 749-760. 

Ffcvse-K. P.. BwVant V _. & P.w-sm. C (1993) Rsumyiion of 
the health costs, to 2030 AD. of enhanced ultraviolet 
radiation due to climatic change Pinal Report to the 
Ocprtitiiierif of Hcahh HohvHMj, ifiul C'onimunin' Services 
and PcparOpnii of'ihc Atts, S^wnt, ||ie tnvtfonmeni 3nd 
Territories, Canberra. 

(fAKDNiR, K M (199.^) Souih Australian Vcarhook No 27 
(AuHlirtlmii Htnrau ol St;MlM|rs, Soilltl Austiit|l:i|i Ollur, 


\ I \I;MK HA HI * M V BM kS 

Gibus. \V. J & Mahi*.k. J. V. (Nti7» Rainfall denies .is 
drought trulic.itiMs f*(imiitoM\vv-;ihlt Buruuu ol Mctcopijojiv. 
Bull. No. iK 

lUlNfis A & Pi i MS. C. <199I) Potential on health 
nl atmospheric thnmie Mmtnul of Ptthtif tirohh Mutlfciitt 
13, 69- m 

HiMtKiui. L (l&KII Historical Experience tif Climatic 
Change in Australia: Cc-Nsons Erom Analogy h BtookbcU 
II & Ootibc, I- (Eds) "Global Change Hv H»m:m 
dimension^ Academy of the Social Sdfintfsa m \uslraha 
JSt Research School c»l Pacific Studies Canberra] 

lltjl k.hitin. J. E. JlnMns, G. J. & Ehhkai ms, J I (Eds) 
(|W<b "Chilian.- Change, (he IPCC Scientdk Assessment' 
(Cambridge University Press, Cambi idgei 

I Nil IW-OVIKSiMI'VlAl PANfcl, ON ClJSMVfc CllANiU (IK < | 

(19V2) (Submission I'rorp Workiny Group 1) 'W2 IPCC 
Supplement, Scientific Assessment nj Climate CTtaBjtP" 
(World Meteorological Organization and Hurled Nauon:-. 
Environment Programme, penevaj. 

0990) "Wlicsmaicert SUirtmarj <■' the Potential 
Impact! of Climate Change, Report prepared for the IK'C 

by Woiloii£ thoijp If (World Meteorological (beam/alum 
and United Nations Environment Programme, Geneva .md 

1 \* tinsos, ,1 I, il l W>) Abandoning homelands . pp V'->, />i 
Brown L. R- el ul (Eds) "State of the V\4ir!d Iwr (New 
York, N»Mt«<to 

Ku not, ant, K M (IVV2) Illness due l>i thermal cuiemi-*, 
pp 4 1 >I M)l to Last, .1. M. A Wallace, R. B (EdsrMaw- 
Roscnau l^ast. Public Health and Preventive Medicine" f Uth 
edition). (Appleton &. Cange, Norwalk, Connecticut) 

l..vr>, (V, A, & Cam.son. B, E. (19M2) (iiobal warming. 
Kccpinjr tlte Son oi proportion NtiWre 360. 2*)7 

Last, J M <W)> Global Environment. Health , anJ Healllt 
Services, pp. 677 684. In East. ,|. M. & Wallace. K H 
(Edsi "Maxvy Rosertau I nst Pubhi- Health and Pievyniiu- 
Medicine" lApplctnu & Lanec. Norwalfc, Connecticut). 

LtAt , A. t!988] Potential health effecLs ol eloba) climatic and 
environmental changes. Nt j \r I'nyjami J» mmul of 'Medicine 
321. 1577 158.1 

U lit i\ J. W. Uiiihm. .1. 1-,, Glass, ft E £ W\rsos. 

A, J, trWl Hantaan < Korean Hemorrhagic Kvcr) and Related 
Rodent Zoonoses, pp. 149 158. hi Morse, S, S. (J;ct. i 
'ImriphV Viliisi-s" (Opioid I'nivrisiiv P/Clft, Nr\v \iitk? 

Mii#ktfNii m, S (l*W?> iinplt-Miiorr ol recent lolal 
alrnosphetu: cuxou measurenitnls t<u bioJiigicalU acliee 
ravlianon a-acbin}' ihe l^uih's sut &w <*/■< ij ih\\>inil Hrsi<tt\l< 
t stars HI, 57-40 

MaS-min -FJaiih, R F Ai Bi 1 1 . U R (l:ds) (l«>H7) rfyfun^n^ 
Iio|>hhI IliM-Msrs" NiMckvnib HdUton (Rdlirrc loidall, 

M* I aki an» , A, UV>78> l>Jily nn>ruht> jnd environment in^lish DnjliirtwUtfWS 2. Death:. dttHQfl suinMiei hot sprllh 
inCirratci l_.*!ndon tfnvtriwtwttMl Ki:u-/mh 15. 333-341 

MrMrtixtN. T A , Hini avsun, R I., HMtots, A & 
Srik.'.mmvv R (I987t Runol'l variability, a global 
perspective In The Influence of C'linum Change and 
C'louaiK Variability in the Hydioln He^llTW and Wmri 
Resources. IMP llrhernationul Hyilroloj)icat Pro^'funum i] 
Pub No |6K. 

McMk-iim'.i , A. I. <W*>.t) lilohal environoiem tb:iti;*c ;ind 
liuiiiau [io|iul.Uton lu';d|ti A i»Hirt*ptii>i! -oul Miriildu 
challenge hi ttideraiyipfiy tni J t:pu!>»tt"l 22, l*& 

(W>V) "Ptorietmv Ovcilo:*d Global f-'m noinnt'iil 

Change and the Health n| the Mutnan Species , *(Canibridj : *i: 
UmvcrMty Pa*s%. Cambridge}, 

MAWvJr-iL, K.. M'Ni/tK, I M Ac Cau^nixk, B. A fTO 'v 
Basic Science of Climate Change, Utncri 342, 102? KVM. 

MrsTn. K (199,1) Rout biinjjs diseaM- and dcaih, ,\>u 
Scientist IKHft. b 

MOHISON. W I . (l!W9) Hth\tS OJ ultOiVioK-l <;mIi;Vioii ott U . 

immune system in humans. Fht4*H'favwiitfTy and 
rtu*thu*toiz\ 5ft. ff|5>924 

Mr k>'AVSMiio. S & WhiNMhtN. P. (1993> A time bomb n 
North Queensland: Case Report q£ Introduced Mjii.n. 
Soiilli o< Hie ntiirtceiiih parallel. Cnrntnutftfe/bh Pritwsiv 
Inlellwrnc 17(10). 211 213 

NichoU-S, N. (l%b) A nicthcHl tor predicting Murray Val|.-\ 
Enceph.iltns in Southeast Australia using the .Sou them 
Oscillation Anst. J. fUp. fttof. Med. .Vc>. 64(6) 587 5«>4 

IW02) Historical 1 EI Nino/Southern Oscillation 

variability in the Ausfinl.isi.m region pp.l5M72. fn Dm/. 
Hi V & Markeral. V. (Eds) M BI Nino HislorKal and 
Pateoelimalic Aspects ot the Southern Oseilluiion" 
iX'dmhlid^e Univetsiiv Press. Camhridgei. 
ryOHitw, N.J. A; Hikn'M, K. s {['Wii I .i^ionmure's dt^Msr 
the iidecbvv dose paradox Uitutt 342{SKb2). 5-6, 

Pmoo . M. 11990) "Climate Chance and World Aericuhurc' 
(l.aiihse;iu Publieaiionh, roiidon) 

Oiri".. W H. (K$2) A study oJ Southern Oseillation 
iliinatie aclivily Itn AD>b22 WO tncorporatine Nile 
River Hood ifciu hi Mia/ ( H E A Mu^ial. V (fdh)"ld 
Nino tlr*:ioiiral and ftdeutiitnalic Aspects ollhe Soutlicrn 
OsciUat^cT (Cambridge t'niversuy Ptcss, Carnbnttg#) 

Ri ssr-i . R (MW) Mosijuitovh and mo^|vnio-borue disease 
in soiiihca'-lcrn Au.-.lraba. (.Arbovirus Disease Control 
I'ornmimv, Nvw South Wah-s nt-pannu-ui *-t H.-nltb) 

Smith, h C , Pwf /i i IN, H. B., BAKtft. K. S., BlDlOM*E. R 
R., BoicHhH, N. P. C-uiK I., Kaioniv.. I > . 
MacInjviu, S. Maiikk, H A, Min/ii-';. I). 
Oiww si-K, M. Was, /., & Waii us, K. .1 (l l W) Omne 
dcjilclion: Ijhraviolci fadiatum and phytoplankton biology 
in Antarctic Waters .SV/f/fM 2S5, ( ;s}-*JV) 

Smok, K iIOO?} Global climate charuie and infeclious 
diseases. r.nvirorwK-nuif Health PcjupacftvtliiG, 171 174. 

S<*>v,, R S (IW^ Al|*nl HkHiins hi tla- Rivet Murray, /n 
! wan. C t', . Bryant. C. A.. Calvert. G. D. & Garrkk, 
J. A il.dsi 'HiMlth in the Grcvnhousi-. Iht- Mt-dHiil hm.1 
EuvHonnicntal Eilccts of Global CtimaLe Change' (AGPS*. 

S<n, 'IH AtSlfrAltAN Rfc^liUNAt Ol'MU. BoM:AO Or 

MiTt,ok!»iu/;v MtMthh Wnilhrt Hivtrw. Mav V) 1 )? h\ 

May 1W 
Slia-AKSKl K \, Hir«.MiOlh, P, Mr Pl'ilk.*-., R I) A 

HrkM.vfs, r, R |N')|) Total o/otie livnds deduced lavti 

Nimbus TOMS data. Geophysical Rf.yrutvh Data L<iit •• 

18. II MS -1018 
iv.iou H R FWg?) .Th» biological clleeiH of LfV B on the 

rye Phntwhintix/r) atui f > JuH/tt/tntri<>Y 50. 489-493 
lllli ApVI.KftSI « Adclflldo, Elid;iy, hVbtujnv \ 1903 
IH! I "*ih K*tk Mmi. Brisbane. Eriday, Pcbruary 5, 1993. 
L'N Enviki/nmi -ni Pk(Kar\MMi (1991) "EnvinmmentJl 

(-.llrusolO^melVplriion I0'» t i|M.Ue"(lNrP. Ntiirobo. 

Vol '. M.. Bki ; \vsri-«. A & SWNNlift, C fl^Mi El Nifid 
Southern Oscillation: Climatic Variability and ClinuiiL 
Change, hi Ewan, ( E , Bryant, b. A., Calvert, ii I). 
*V (iainck, J. A. (l-.ds> 'Health in llu- Cin-t'iihoithc Thi- and Envifonmentiil Elleel^ ot Global Climate 
Chance" tAOPS. Canberra). 

Wlinsh in, p il99l) Arbovirus Notifications. South 
Australia, 1<W| tin 7 May) CommttnuuNc DryttfieJ 
hiwthwncc 15(11). |& 

Wokio-si. R i. x Gpant i- n. yoSM Biftfecw pj 

uliMviolc-i H iwflflUon «^ii iriicsinal plains arid inoiine 
(»rpani»nis, Dp 197-206. In Ruisell Jones. R. & Wicles. 
T iLdsl 'Ozone Deplelion: flealth and Ensironmemul 
L onsL'tniencL-s (John Wiley •< Scum, t'lhehestfU 




VOL. 118, PART 2 




By T. A. Thumlert & A. D. Austin* 


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, Chalcididae, Elasmidae, Encyrtidae, Eulophidae, 
Eupelmidae, Pteromalidae. 

Tnmuu-ium tftfti MW Snrwn o/ S, Aust (WW, IIX(.m. W-lh' 


by I. A. Thlmi RRX & A. D. Austin* 


Thimuki, T. A. & Ai-SUN, A D. (1094) Biologv oi Phvluitrafjham Jht^uttl Riek iHymenoptera: Pcrjiidae) 
*nd it> paraMtoidi in South Australia. Trans, R. S&. S. Aim. IlSOt "910. 3! May IW4. 

Information is presented on the biology of FYrWui reopha^ifrny^ani Riek (ihe leal-blister sawlly f, art emerging 
pest nf plantation and ornamental eucalypts in south-eastern Australia. The range of cucalypl species attacked, 
the extent Ell tree damage, seasonality, fecundity and longevity of P. frng%ntt\ art* investigated tor populations 
in the Adelaide region. The hyrnenoptemn parabitoid* associated with P. fiitggatli arc reviewed. An illustrated 
key to the 17 *;occic\ encountered in the Adelaide region is presented, along with notes on rJlinf hioloy.y, relationship 
with thru host, ntid species recorded from other pans of Australia, 

Ktv Words* Ph\)aiU'{}i>hti}infrofWnnt. IVrgidae, parasitosis, hyperparashoids. Bracotndae, khncumoiudac, 
Chalcididae, nUtsmtdae. Fncvrlidae, tulophidac t Cupchrtidac, IVtontnlidae 


Phylacieaphaga fwggatli Riek, the leaf-blister 
savvlly, is a native leal-mining species that is generally 
restricted in distribution to the south-eastern and south- 
western parts of the conl inept (Riek 1955; Benson 
1963; Farrell & New 1980; Curry 1981). Larvae feed 
on a range of euealypl species by mining the mesophyl! 
layer between the upper and lower epidermis of large 
leaves. This pattern of feeding results in characteristic 
oval or efrmgate, brown blisters, which are nost 
common on the lower branches ot young trees (Farrell 
A New 1980; Nuttall 1985). Heavy infestations of P 
ftQggQtlt cause substantial loss of photosynthetic area, 
resulting in stunted growth or death ol trees. This 
damage is of most concern where large numbers of 
saplings exist in a confined area, such as euculypt 
plantations, ornamental eucalypts in parks and gardens, 
and potted trees in outside nurseries. 

lu 1985 i* fruggarii : was accidentally introduced into 
New Zealand where n has become a serious pest of 
euealypt forests and ornamental trees (Nuttall I9S5; 
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 (Auslin & 
Faulds 1989; Faulds 1990), 

fjiven the pest status of P. jroxgatti, relatively 1 ttfc 
work has been undertaken on its biology oi that 0] jts 
parasiloids Other than the work of Fartvll A New 
{ I9K0), who provide a detailed account of some aspects 
of the biology of P. frogj&tfti in the Melbourne area. 
the species has received only anecdotal attention 
(Froggatt 1899; anon. 1950; Riek 1955, 1970; Moore 
I960; Curry 1981; Naumann [983; Bnngey WS6). 
Information on its parasiloids is even more limited. 

Dquuimcii! ofCrBp Protection, Waile Campus, Univoity 
of Adelaide. Glen Osmond. S. Ausr S064 

Farrel! & New' (1980) record some observations on the 
development and mode of parasitism tor two species. 
Bracon sp. and Cirrospilu,s sp. in the Melbourne area, 
while other workers have simply listed the species 
reared (Moore 1966: Curry 1981; Boueek 1988). In 
these studies no more than six species have been 
recorded and, with the exception of Boueek (1988l 
the accuracy of species identifications is questionable. 
More recently. Austin & Faulds (1989) have described 
two species oi' Bracon that have been introduced into 
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 P froggatii, focusing on aspects 
that have been largely neglected by previous worker* 
At the same lime, detailed information is presented on 
the large parasitoid complex associated with P frnggatti 
in the Adelaide region. An illustrated key to their 
indenufieation. 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 ihc ecology and possible 
control measures of this emerging eucalypt forest pest. 

Materials and Methods 

b'U'Ut .uic\ 

Huealypl trees infested with larvae and pupae of ft 
froggani were monitored at several sites in the Adelaide 
region, viz., from ttie western coastal suburbs (Dover 
Gardens), city parklands and adjacent suburbs (Stum. 
and LobethalAVoodside area (Adelaide Hills), between 
August 1990 and July 1991. The first two sites comprise 
mostly ornamental trees (miniatures o\ numerous 
species) in parks and along roadsides, while the latter 
site is semi rural and comprises both ornamental trees 



and natural!) growing trees in paddocks and along 
roadside verge* (mostly Eucalyptus nmutltluh'mi\, f 
chuUnalyx and t. Irwrcu/fv/), The climate ol the Inst 
two sites is very similar with average monthly 
maximum and minimum temperatures ol 2&°C and 
l3*& while me Lohahal-Woodsidc site has average 
monthlv maxima and minima of 2l°C and 8*^0. The 
average annual rainfall for these two areas is 
substantially different (Adelaide 5K5mm; LobeihaJ 


(aborafan retiring 

luiested blanches, mostly of E latnahlitltmis, were 
collected from the field, placed in plastic hags and 
returned to the laboratory. They were placed in 
'.oniainers ol watct al room temperature or constant 
20°C and TlfropgatU allowed Ed develop. Leaves with 
larvae, pupae and/or developing parusitoids were back 
lit with an optical fibre light to determine the exact 
stages present and observe their development Most 
parasitised and unparasitised pupae were separated into 
5 cm dia. plastic containers and allowed to complex 
development. Once emerge, adult P, fktggfiM and 
parasitoids were kept alive by supplying ihem with a 
smear of honey and water via a saturated cotton roll 
poshed through the lid ol a small glass vial. Virgin 
female R frotwatli were sometimes isolated from males 
as soon as they emerged lo provide individuals (or 
laboratory experiments. 

f'tuututitv ami hmgtvity 

Virgin female P. Jhwgorri which had emerged over 
a 14 hour period weie held al MHva.ini ?0°C with honey 
and wilier supplied. Ten individuals, randomly selected 
each day, were measured tor body length, tolled in 70S 
EtQUi si juashed on a microscope slide and (he number 
ol lulls developed 6gg$ counted. 

To determine the longevity of /* fro^atti adult 
females were kept m 20 cm diameter plastic cages and 
their survival monitored twice daily until all wasps had 
died. Longevity was compared tor I) virgin or mated 
females. 2\ with or without water and honey, and M 
al live ic-mpc-ouuicH (Iff 15, 20, 25, 30°C) Urn females 
dial had emerged within a 12 hour peiiod were 
randomly selected ftOfla stock cages and liunsletrcd into 
experimental containers. Three replicates were run lor 
each of the above 20 treatments Honey or water were 
not supplied during mating to those which went into 
Ihe "no lood" ireatments. All cages were held al 12.12 h 
photoperiod in temperature cabinets wilh M).5 t* 
tempeialure ranges. The mean survival time for each 
treatment was calculated and plotted against 
temperature, and ihe regressions analysed using a 
2-tailed F-test 

Tavuinitfv onii \rfjfmin$ electron mt'croxcopY 

Terminology I»m motpliologtcal structUies generally 
lollow Ciauld (1984) and Oauld & Bofeui 11088) fiw 

iehncumonid and braconid wusps and Boucck N98H.I 
for chalcidoids. Where necessary, morphological 
structures referred to in the key are indicated on the 
figures. Voucher material of all species have btxn 
lodged in the Waite Campus insect collection. 
University of Adelaide, 

Specimens tor SEM examination were killed fllttl 
washed in a sirong detergent solution before being 
dehydrated it) 100% EtOH and critical point dried in 
;m Kmscope CPD 250- They were coated twice with 
carboii and gold/palladium before being examined 
under a Phillips 505 Scanning Electron Microscope 

Results and Observations 

TdXQIf&ffiy (jf Pbylactenphaga 

The taxonomy ol Phylacteopluiga species is contused 
and requites urgent study. Nominally, only one species, 
P eucalypti Eroggalt, is recognised which has been 
divided into four subspecies (R e, eucalypti, P e 
fro^arti Rick.. /' C tasmnnica Riekand P. e, otcidtnH 
Benson) based on colout and si£e (Benson 1963). At two of these subspecies [P. e, fraggatti and P. «* 
nccidcris) occur sympatrically and may therefore 
represent distinct biological species (Naumann 198'3). 
However, of the several hundred specimens reared from 
leaf mines by us during 1990-91 all belong to the one 
colour form, that of P. e, fvggM Rtefc (i-C antennal 
scape and pedicel and sculellum of female pale yellow 
tit orange-yellow). Furthermore, mating of field- 
collected material in the laboratory was always 
successful- supporting Ihe ptcmisc dial eithei only one 
species is present or one is dominant in the Adelaide 
region. Because all reeeni auihors (ca Parrel I & New 
1980; Curry 1981. haulds 1990) have adopted species 
level status for the subspecies P, *\ fm^arti, this 
approach will he used here lo save confusion. 


In the field, males were often observed perched on 
the dotsat sutlacc ol leaves with their body held at 45° 
above the surface, in Ihe vicinity of ovipositing, or 
testing females, or Hying in small swarms of 5-10 
individuals, close lo leaves where mating pairs were 
perched Copulation is strophnndous, i.e. male and 
female a/e (omed end-to-end with the male genital,.* 
twisted through 180° (Gould & Bolton 1988) In the 
laboratory mating took 2-3 minutes. During copulation 
the female stands with her wings folded at rest, while 
males have their wings spread at about 45° and 
peiiodieally fan them. After uncoupling, the female 
immediately moves away and the male remains in 
place, tapping its abdomen scvend times on the leaf 
surtace betoie Hying oil, 


II . I 

fecundity *W o^positim 

The number of e^*,\ cart ted l>y vitgiu females ranged 
(hum 5? lo 87. while Cgg number within Ihese limits 
was positively correlated with both age and size of 
lemales [i 0.76 and r -0.66, P<O,05t On inridun/,, 
iem.ile P fi**fti&ttti move sideways over the suifaceof 
a leal perpendicular CD die mid vctn. with this 
movement continuing until ati oviposttton site is 
chosen. The female saws into the leaf tissue far J 10 
sot. then remains stalionary lot IM nun wftile 
oviposilmy, At dns time the bpdy and antennae are held 
at an angle of aboul 45° to the leal and to (he mid vein, 
possibly id position I he egv m> that the larva hatches 
in the direction towards the apex of the leaf. Once 
completed, thr female tubs her abdomen over the 
position of the 6§g spot three or tour tunes, poss-.bly 
Id seal ihe bole, and then moves away, either Staying 
on the same leaf oi moving to another leal to oviposit 
[vfijL's are generally laid close to the mid vein attd along 
the length oi the leaf, but not tn the distal ?0 30 mm 
The small egg spots, aboul 0.X mm In diameter, lorm 
raised jdohuJous Mtuctures < egg-galls) on the surface 
Of the leaf. The number of egg spots per leal ranecd 
ftom 7-170, with numbers over about W rcpresen me. 
the oppositions ol more than one Jciuule. 

Ihe morphology, coloul add size of the egg. iaival 
instars and pupa of P fras%atti are virtually identical 
to those described by l-'arrell & New (1980). Briefly. 
the egg is flattened and oval in shape, measmm), 
()./> s 0.5 mm. There are five larval instars in the 
female and (bur in males. Tor the Adelaide population 
females bad mean head capsule widths of 0.49 (I). &62 
(II), 0.80 ill!), JIM (IV) and I 25 mm (V) <n > # 
in each case), with malts beiny virtually identical to 
Jeuiale tusiats 1-IV. These sizes are almost identical 
to ihOSC measured for the Melbourne population bv 
Farrell & New (WHO), with the exception of female 
inslar V which averaeed I .OK mitt m the hitler study. 
Ibis discrepancy may be due to our selection of larvae 
from non-overlapping mines, where there was no 
competition for lood between individuals. At room 
temperature in the laboratory during January and 
February (daily avctaec about 23°C) larval 
development to cocoon initiation (t.c including Ihe 
prepupal period) look 15-18 days and ihe complete Irfe- 
cyclc about 30-35 days. 

Pupal ion is preceded by a prepupal stage, 
distinguished from the last larval instax by a lighter 
coloured head capsule. The picpupa moves awav f W\ 
the light perimeters of the mine into a mote cetitiul 
position where it begins to spin a thin ovoid cocoon, 
Silk is laid down ftom mandibular glands as it swings 
its head from side to side. The cocoon js attached to 
Ihe mine flix>r and lite leaf 8UI facft abcry* by silk, and 
forms an annulu> of creases ut llic blistci tissue. The 

period from cocoon imitation to adult emergence took 
about eight days m the laboratory tor berth sexes Two 
to three days after cocoon iormaiion the prepupal 
cuticle was shed revealing the eye spots and appendages 
of the developing pupa By the fifth day llic pupa 
hecame pigmented, at which stage the .sex of 
individuals could be determined by the colour ot (he 
ihotaxaud picsencc of the female Ovipositor, ihe thorax 
of males being black and that of females ontnge, The 
pupa moulted on day .wen into the pteadult stave. 
which is chatactetiscd by having all the features of a 
fully formed adult. The body is completely pigmented 
and the wings which begin to unfold, are frequendy 
lanned whhin the mine to help them dry On the eiehih 
day an oval-shaped hole is c ut in the lop ol lite nunc, 
and the wasp voids a gtecn-ttcy meconium just belore 
eating or immediately alieiwards In the Jabotatory 
approximately 25ft of adults died after (ailing >o 
emerge from their mine. This mortality was highest 
lor leaves containing multiple individuals 

l.on(te\it\ and (wnvittlmny, 

To dcietmtrie tnc optimal holding conditions for P 
ffi)g£&Tfi in the laboratory and determine the potential 
for the species to overwinter as adults, longevity ol 
wasps was measured at vaiious tcmpciatuics ^u\d 
combinations of mated orunmaled female*, with (hex) 
and water supplied or not. The te$*resMoiis ot mean 
surv ival time against temperature were significant tor 
the four combinations \M virgin and maled lemales, 
with iuid without food and walet supplied (fig. 1) Ihe 
slope of the regression lor females with lood was 
significantly different from those wiibom food ami 
water for both virptn IF = 6.X5;d,l = 2, n; P<U0S 

fig. laj and mated females if 474, d.f. - 2, 5; 
P<0.05 - Fig. lb), but mating did not afffct the 
survival of Ihe lernulcs lor citbet of the treatments with 
if 4.5, d.f. 2. 5; P>0.05l or without (bod and 
water (F ^ I 64; <1 f. = 2, n; P >0.<)5). In figure lb 
the data lor mated females with food and water supplied 
a! 10°C arc excluded because of tile breakdown of the 
temperature cabinet pan way through Ihe experiment. 
The longevity oJ iemales was increased when held at 
lower temperatures and when water and honey were 
supplied, but was not affected by mating. It ts not 
known whether horlt water and honey were responsible 
for the increase in the survival rate ol the tetnalcs, as 
these factors were not tested separately. Possibly only 
water is required to prevent females from desiccating, 
and lood is ununportam for longevity, though it may 
he. necessary for increased fecundity. 

The longWl 'hat adult females survived in the 
laboratory at tempo atutcs below 10°C was '28 days 
Hence, il seems unlikely that f* fibgfttftti overwinteis 
only as adults A small number of occupied mines was 
found on irees in coastal suburbs of Adelaide during 
June and July, hut at no other siles. It seems likely. 



therefore, thai some pupae continue developing in 
refuge areas until about mid winter and the subsequent 
adults remain quiescent for Ihe rest of winter before 
ovipositing into leaves in late August or early 
September (sec Fig. 2). 

jltifta? timfopment 

First and 2nd instar larvae first mine towards the 
distal end of the leaf following Ihe direction of (he 
venation, bid as they grow with each subsequent moult, 
the mine becomes progressively larger and circular or 
oval in shape. As observed by previous authors (Farrcll 
& New 1980; Nulla]] 1985: 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 
I hat almost the whole leaf can become blotched. The 
largest number of larvae observed to complete 
development on a single large leaf was 21, so thai any 
remaining eggs from the large numbers of ovipositions 
(see above) either died beforehand or were eaten by 
the first emerged larvae. Leaves supporting more than 
about Hi larvae often resulted in smaller adult wasps 
compared with leaves supporting fewer larvae. Other 
authors (Farrell & New 1980; Curry 198JJ have 
reported Ph\lavieophaga as confining its development 
only W the lop surface of a leaf- This was confirmed 
here lor cucalypt species that have differently textured 


20 25 

Temperature < ac > 

13 20 

Tornoeraiure ( : C) 

Pig- I- Longevity vi' Q Ph\'lucieophtt,i>tt Jht^uni measured 
as rhr mean survival tune for three replicates of 10 WUKJta 
(i:'S.E.) at different temperatures, ia) tor virgin 9 9 Vtflft 
[pod ami Wdter supplied (■) and without food und water 
I I, and trO for mated 9 9 with food and water supplied 
<•) and Without tbod and water ( :) 

dorsal and ventral surfaces, but tor species with 
isoluteru! (eaves (i.e. identical surfaces on leaver 
hanging vertically), mining was carried oul on either 
side, sometimes resulting in individual larvae being 
separated only h\ the internal venation of the leaf. 

Damage w iTttss 

The levels of damage varied between infested (roes 
and between individual sites, possibly because of 
differences in the tree species present, their size, age 
and health Quantifying damage for these variables was 
beyond the scope of this study but casual observations 
In the field aliW for some useful generalisations to 
be made. Tree.s which supported large numbers of P 
pfiggUtU mines were similar in several respects, they 
were usually small <<4 m tall), were often situated 
on roadsides, carparks, parklands or isolated groups 
in paddocks, and were usually sheltered to some degree 
from westerly winds. Possibly sheltered localities 
prevent adult wasps from being blown away from their 
host plan I, since lhe> are not strong fliers. I rees that 
became heavily damaged appeared healthy in the early 
stages ol attack, in that they had dense lush foliage and 
did not appear io be stressed As reported hv FantrJI 
& New ( 1980), damage is more severe towards the base 
of trees, particularly on low hanging branches The 
apex region is usually not attacked, especially tor trees 
above 4 5 m in height. In some suburban areas, where 
there were many young trees ol the same species (1-2 tn 
in height), damage often reached very high levels. 
causing leaf shedding and sometimes death of saplings. 
Well-established trees OK m high), were far les. 
frequently attached and, if so, were usually in clo%e 
proximity to heavily infested young trees. Mines on 
such trees were generally confined to low hanging 
branches with semi-mature leaves. 

Hoa nmge 

The range of tree species attacked by Phyktcicaphasa 
spp. was compiled by reviewing previous studies, as 
well as surveying a wide range of trees in the Adelaide 
region during J990- 4 )! Trees in open situations, 
nurseries and gardens were examined, with those in 
the latter two areas being used to confirm specks 
indeniilicalion. The data presented in Table I show 
that at least 27 eueulypt species arc susceptible to attack 
both in Australia and New Zealand, with most pi these 
records being attributable to P. fr(%i>atii. In Australia 
the most widely infested species arc Ettealypmy 
hotryoifies, £ c<in>alrinUttsis. F, • itriodnnt, f 
rlatlocatw, E. ficift>lia, t. $\obtttHt and £. xranafo 
Also, several non-eucalypt species have been reprrtod 
as hosts, but there is some doubt as to the accuracy 
of at least some of these accounts as they have mostly 
not been confirmed since the initial tegorta 



Patvsiroid species 

Over the period of this study 17 species of 
hymenopteran parasitoids were associated with P 
froggatti- 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 
hyperparasitoids {Elasmus ausira lien sis Girault, 
Cirrospilus ma rg is cute 1 1 urn (Girault) and Eupelmus 
sp.), while three have not had their biology conformed 
yLanopltagus sp. . Pediobius sp. and Chrysononnnyia 
sp. 2). The species in this latter group were not 
positively reared fxoxd P. froggatti. but rather emerged 
into rearing containers with euealypt 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, it is likely, therefore, that these three species 
are parasitoids of P. fioggatti, but further study is 
required to confirm this. 

Seasonality of Phylaeteophagu froggatti and its 

Fortnightly field excursions from August 1990 to July 
1991 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 
preadull 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 ol 
infested leaves and trees that P. froggatti is most 

TABLE 1. List of tree species which have been recorded in the literature and during this study as susceptible to damage 
hv Phylacteophaga spp. (sources of information are as follows: I, Anon. 11950/ and 2. Moore fI966f, coastal N S.W: 3, 
Riek /I955jand 4. Earrell & New/!9H0f, Melbourne reghm, Vh\{ X this project 11990-91}, Adelaide region , S.A.; 6, Curry 
{1981}, Perth region, WA. , 7, Nuttal J19H5J and H, Ka\ I19H6J, New Zealand: sources J 5, 7 and 8 are far Phy lacleophaga 

Heal Plant 

Recorded Phylacteophagn damage 
N.S.W. Vic. S.A. W.A. 

2 3 4 5 6 

N Z 

.uhptus botr\oide.\. 
camaldulensis . . . 


cttriodora .... 


diversieotor ...... 

ficifolia , 


vamphocephala . . . 

grandis , 

lehmannii 1 1 


U-mnxvtori rosea . , 

longlfolia . . . 

nun utata. 



melliodora ... 








sideroxyton ....... 



Agonis flexuosa 

Bciula (Birch) 

Utphostemon ennferuts (TriKlania) 

Quercus palustris , 

Q. robur * 




Table 2. Summary of the relationship between Phylacteophaga 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 







Gregarious (G) 

Ectoparasitoid (C) 

Hyperparasitoid (H) 

Braeon confusus 


larva; pupa 





phylacta yphagus 


larva; pupa 




ParaphvlcLX sp. 






Brachymeria sp. 






Elasmus australiensis 




P; H 

Apleurolropis sp. 






Chrysonotomyia sp. 1 






Chrysonotomyia sp. 2 









larva; pupa; pa 



P; H 

Cirrospilus oi dpi t is 






Cirrospilus sp. 3 



S; G 



Cirrospilus sp. 4 






Cirrospilus sp. 5 






Diaulomorpha sp. 


larva; pupa 




Pedlohius sp. 






Eupchnus sp. 





P; H 

Lcuiophagus sp. 





1990 1991 

S . O . N . D JFMA 

Phylacteophaga froggatti 
Adelaide Plains 
Adelaide Hills 



Bracon confusus — ^^_ 

Bracon phylacteophagus ^^_— ^^^—^— __^___^^_^^^^^^___^^_ 

Paraphylax sp. • 

Brachymeria sp. — 

Elasmus australiensis •^— 

Apleurotropis sp. ■ 

Chrysonotomyia sp. 1 ■ 

Chrysonotomyia sp. 2 • 

Cirrospilus margiscutellum ' 

Cirrospilus occipitis - 

Cirrospilus sp. 3 _ _ 

Cirrospilus sp. 4 i i 

Cirrospilus sp. 5 » 

Diaulomorpha sp. -^ ^—^— 

Pediohius sp. • 

Eupelmus sp. ^ — 

lariophagus sp. • 

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. 

rm'ijcTor.omux* ntotwrri and ra i*\kA$noii;s 


numerous at sites on the Adelaide Plains from about 
early December io late March < Rig. 2). unci thai (hero 
are three, possibly four, peaks in numbets over the 
perunl September to March. In the Lobethul-WooUstde 
aiea {Adelaide Hills), largest numbers occurred slightly 
later, from about early January to late March, with $gg 
spots and mine initiation not appearing unol later 
November. presumably because of the coolei average 
iciupciutures there compared with the Adelaide Plants. 
Of the U species of parasitoids, six were encountered 
over a period of Iwo months or more (i.e. Bma>tt 
phvhnit'ophugus Austin. CiiTDSpilus. nuitgis* rurvlhtti , 
C bccipfft-S Girault and Cirrospilus sp. 4, f:itprltntt\ 
sp. ami Dinulomorptw sp. 1 (Fig. 2). and Of these (ft) 
descending order) C tmirt*ixvi4teUttm . ft 
pln'kutcophngits. Eupebmts sp. and Diaulomorpht; sp 
were reared in the largest numbers. No parasitised 
hosts oi parasitoid pupae were lound in mines during 
July and August, though two udult female ft 
ph\'{tiiffiipha\\us ami one (.'. muryJMiut/Uon were 
located in old mines during this lime, indicating thai 
these :md possibly all parasitoids in the /' fn/^uUi 
complex oveiwmtet quiescently as adults. 

Lewis of patusttism 

Collections of mined larvae during January and 
February (991 enabled assessment to be made of the 
levels of parasitism in the field at three separate sues 
vi/ , I obethal t Adelaide Hills), Slurl and I>.>ver 
Gatdens (Adelaide Plainst (Table 3). These data show 
a forge range in the level o\ parasitism o( P. fnni-.uut' 
(15,0 to 60.3%), and in all cases C ?U4f$SCUt€ltUM W& 
by far the most important parasitoid. Other parasiii id S 
contributed to \\.5% of total parasitism, ol which 
H. phyhuifopha^us was most common, These data. 
however, represent a minimum estimate rjf percentage 
patasihsm. particularly tor Ihe Stmt sample. wheic P 
twggsm were mostly represenred by early life-history 
stages when collected, so thai those sampled could have 
been paiasitiscd if left in the field. Because C 
ftuiftfiuutelltim is capable ot paralysing and Icediug 
on ail stages of P. fraggpm from early instars to the 
prcadult stage, art estimate of the maximum potential 
parasitism was obtained by adding the measured level 
ol parasitism with the proportion of hosts still viable. 

Bnrfn#y of ptinisitvuJ spent-* 

Family lehneumonidae 

Puraphvlax sp. 

FIG. 3 

There arc more than 50 recognised specie- ol 
Puntphsinx from Australia, the majority of which arc 
undescribcd (Ciauld l9H4i. Where their hiologv ift 
known, I hey have been recorded mostly a% prhrtaQ 

ectnparasitoids arid hyperparasitoids ot lepidoptcran 
hosts. The single male specimen reared here in March 
1'V'M is an cetoparsitoid of P fw^am larvae. It belongs 
to the iovu.x species-group, and is probably the same 
species as thai recorded by Gauld (I u 84) from P. 
ftfigftutti in south -eastern Australia, Being Ihe only 
ichncutnonid in this parasitoid complex, Paritph\la\ 
sp can be easily identified l>y lis characteristic wing 

Family Bnieoniduc 

ftnuon &J1$UU& Austin $ ft phyfocteophagus Austin 

bICS 3, 4. 6 

Both these species aic solttaiy pinnaiy 
cctopurasitoids of third iostar larvae to early stage 
pupae ol P J'roxxatii liable 2). B. fikylacteopfotgttS was 
the more commonly encountered species of the two, 
ami was found associated with us host for the same 
period thai occupied mines were present in the field 
(September to May). R o>nfusus was collected for only 
a short period during December-January. The biology 
pjf these species has been discussed by Austin & faulds 
(WS9) and Faulds (1990) and only observations 
complementary to those given by these authors are 
presented here. Pnoi to WHV, inferences to Bruam 
associated with PhyttfCtcophagfl did nor recognise the 
presence of two syitipatiica'ly occurring species. 

B. fim/t4sus and B. j»}i\kuit\'phu^u.\ oviposit one egg, 
rarely two. in a mine, either next In or onto the surface 
ol the host. The host larva is paralysed at opposition, 
thereby allowing the secure attachment of a newly 
emerged Bruam larva in the absence of an active host 
which normally whips its abdomen about when 
disturbed. Late stage Branm larvae are easily 
distinguished from other parasiloids by their large sue. 
dorsal ampullae and covering of long hairs, but they 
are identical to each oihcr. Contrary to the findings 
of Far rell & New (1980). most laivae which we gently 
dislodged from n host with a fine brush did not die, 
but rather successfully reattached themselves- When 
lecding is completed. Bmcon larvae normally move 
away from their host before commencing cocoon 
( urisintction but occasionally pupate inside their host's 
pupal cocoon When spun jway from h host pupa, 
Bfaam coioons often have loose frass and Mptfltfcd 
cuticles mcoiporaled into their silk matrix. They are 
white in colour, ovoid in shape and measure about 
7.U "X li3 mm. The confined prcpupa deposits a 
meconium at one end of its cocoon anil then bunches 
into the opposite end so that II occupies only about 
h.ih of the internal space Adults emerged in the 
laboratory 11-12 days after the commencement of 
cocoon eonsUucUou. Adult B. p,hxfat'Wt>pfui$u.s held 
at constant I5°C were still alive after ON davs and 
individuals were observed to feed on ihe honey and 
water provided 



Along with PamphylaK sp. t ft Cattfiisks and B. 
phvlucivophagus can be easily recognised by their 
relatively complete venation and large size I <3 inmi, 
and the species separated by their colour and 
differences in pilosity. 

Family Chalcididae 
Bnuhymti tii sp. 
FIG 7 
This genus is a moderately large genus in Australia 
comprising primary parasitoids and obligatory 
hypcrparasitoids o\' the pupae of a great ranee of 
holometabolous insects, but particularly I.cpidoptera 
(Boucek 1^88). The species recorded here was 
reared from several pupae ol P. fro^atti during one 
week in December IW). Brtuitwurta sp, is easily 
identified by its small robust form, enlarged hind 
femur, elongate posimarginal vein, and densely 
punctate dorsal mesosoma. 

Family Eupchnidac 
Eupelmus sp. 
FTC. 8 
This species was generally reared as a solitaiy 
primary parasitoid from the pupal stage of its host, but 
on several occasions was also found to be 
hyperparasitic on smaller conspeeific larvae and those 
of several other undeniilied parasitoids. Mature larvae 
can be identified from the other common parasitoid-. 
in mines by the presence of dorsal ampullae and dense, 
long hairs on the thoracic segments. Enpelmus sp. was 
reared from P fioggafU from mid summer to mid 
autumn. Adults can be separated from the other species 
in the complex by the presence vf a greatly enlarged 
mesopleuron. exposed ovipositor, and form of the wing 

Family Flasmidae 

klasmus uit.strtiliensis Girault 

FIG. 9 

This species was reared either as a solitary primary 

ecioparasiloid oi hyperparasitoid of P. froggaili during 

December and January, Previous reports (Rick 1%7; 

Austin & Allen 1989; Allen 1990) have recorded this 

species as being gregariously hyperparasitic on 

ichncumomd pupae associated with the lepidopteran 

defoliator (hvba lumens Walker tt can be readily 

identified by Us large disc-shaped hind coxa, cream 

coloured legs with black hairs and spines, and 

distinctive criss-cross pattern ol black hairs on the hind 

tibia. See Rick (1967) for additional taxonomic 

information and list of synonyms. 

Family Pieromalidae 
txiriophiiiilts sp. 
FIGS 10, II 
According to Boucek (1988). this small genus is 
n presented in Australia only by L. distinguvruitts 
Uoersteil, a cosmopoliian parasitoid of beetles 
associated with stored grain. The single specimen 
reared here was removed in early February l°Vl from 
a P. ftuggaiti mine that did not contain any other 
parasitoids or other potential hosls, Given (he paucity 
of biological miomiatiou available on this genus we 
are unable to speculate further on this species 
However, given that a dead P fttyfaQlti larva was the 
only other occupant ol 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 u( the antenna. 

Figs 3. 4_ Dorsal view ol the "I V flftnwi -aip,. S tfntcon phxkutcnphuxus Austin, A, tiratvn confiwis Austin 
Scales ^ 250 um>- 



l<miily Hulopbidae 
f'x!*ii'jtnus ^p 

fig. 12 

Members ot this genus arc known lo be primary and 
hyperpara.siunds of eggs, pupae and sometimes larvae 
of various inseei-,. mainly Of l.cpidoptrra. Coleopteru. 
Diptera and other Hymenoptera. with several species 
being reported as attacking leal-miners (Boueek 
IS>88> An unidentified Pfdiofritts specica IS 
hyperparasitic on U. likens ^ Austin & Allen 1989; 
Allen 1990), the latter occasionally appearing on (lie 
same host plants as P fruy^ani- However, variation 
in sculpturing on the scutellum indicates that the 
Pafiobiits associated with these two hosts are drfftrefl 
species, Previously. P. bruchickla (Rondani.i a 
gregarious species (Boueek. 1988). has been found as 
a hyperparasitoid on B. ph\tucteopfuti>u.s hi New 
Zealand (FauJds H90J, and this species resembles 
closely the single specimen recorded here Fftftnbjub 
sp. was reared m a container from a numbei fff leaves 
mined in late Decemher 1990 Inspection of these leaves 
revealed the presence o\' no other potential hosts. I ike 
Aplettwtntpis sp. and Dnm/ttmorphu sp.: this species 
can be identified from others in the complex by the 
presence of several long hairs on the scutum and 
scuielluiu, urtd the posterior expansion ni the 
scutellurri- in addition to the characters in the key. it 
can be separated i'mm AplmttvtrapL\xp. by the absence 
of a posffrro-mcdijl suIu_n on the scutum and from 
Diiittlomorphit sp by the very distinctive colour ol (he 
latter species 

Aplcuredopis sp. 
FIG 13 

This genus has previously been recorded .is erm*rijjn}! 
I mm P. ffiUggOUi mines (Rick 1955. Farrell & New 
1980), with Boueek (1988) listing A mmotipt ?vu\ 
(Girauit) as parasitisine P. imulyplt in the Melbourne 
area. Material reared over a three week period in 
December 1990 here confirms that this specif, i-, 
gregarious, with the Jarvae probably being 
endoparasitic, as holes were found in the bodv at die 
host next to the parasitoid pupae. A single mine of a 
laie stage P fro^ntti larva was found to contain 12 
parasitoid larvae and pupae, of which mosi pofnjil £ted 
development m the laboratory- Compared wtftl 
Pfdiobius sp.- this species can be easily recognised 
by the presence of a large postero- medial sul*. »j> on 
the scutum (see above) 

Duuilomorpha sp. 

FiGS 14, 15 

This gregarious primary ecloparasitoid attacks the 

laivae and pupae of P. frdggaHi with up In 1(1 

parasitoids being iceoidcd from ,j single mine At loom 

lempcrature itiOtf individuals hi a Single mine emerged 
within 13 days after the commencement of pupation. 
Mature Urvac can be easily recognised from the other 
common parasitoids in the complex because of their 
gregarious hehavimir and ubKenee of dorsal ampullae- 
Adults can be recognised by their metallic green- gold 
tolour. while leys and light brown antennae Nee 
comments nndei Ptvfiuhhn sp.) An apparently 
uudesenbed species belonging to this genus has been 
recorded fnim PhvUuU-opnd^a mines in Western 
Australia and elsewhere in Australia (Boueek 1988), 
and this may be the same species as reared here. 

Cfiwsotwtofnyw -p. i and *p - 
WC5 lb. 17 

According to Boueek (.1988), members of this 
medium m/vU t.'viais ol adout 40 described Australian 
species, develop in the eggs or young larvae of leaf- 
mining and gull-forming Diptem and Lepidoptera. 
Virtually no biological information is available for any 
Australian species, eScepl lor one reared from Penhuki 
^Ixpltopa Common (jairah leaf miner i in the warmer 
legions of the Darling Ranges and coastal regions of 
south-west Western Australia (Ma/anec 1988). It is a 
solitary endoparasitoid most frequently ol the 2nd and 
3rd ucsiar larvae. 

Two species belonging to this genus have been 
ass'teuited with P. fhtg$ktti in the Adelaide region. 
OirysoHotutnxia sp ! was reared as a gregarious 
primary ecloparasitoid of a single late stage larva ot 
." fitig^tH : collected at Lobethal in March I99|, while 
two specimen;-, ol Chn.spno'otnyuj sp. 2 were reared 
only from mined leaves in February 1991 collected at 
die Dover Gaidens site, and therefore arc only 
tentatively assumed to be parasitic on this host. 
Although very small in sue (about 1 mm in length). 
these two species can be separated from the other 
colophiil genera in the complex by the ante.nnal funicle 
being only 2 -segmented and the scutellum lacking a 
submedial groove. They can be separated from each 
other by the form of the wine venation and their colour. 

t'itr,>sfnti<.s rthirxiM uft'lltim (Giraull) 

FIGS 18. 19, 22 

Cim>\p]his is p large and taxonomically difficult 
genus of small often brightly patterned wasps which 
are well-known primary and hypcrparasitoids of 
Mijc.ilypt leaf -mining insects. Two described species 
have previously rw-n remed from PhvhifTrfpfniga spp.. 
C. mctrutsiviriiaw 'tuuiult) from Western 
Australia and C. oca'pitis Girauk from N.S.W. 
(Boueek 1988). while undetermined species have been 
reared fmm P. fto£gum in Melbourne by Rick (1955) 
and Farrell & New (1980) arid in Western Australia by 



rmucrowpHAGA fkoggatti and its parasitoids 


TAB] I J. Mortality <>/'Phvh<:teophaga I'roggaUi m wealed In collections made during January and February 1991, from 

hdnthul where trees were tri relatively early stages cf ttltaLk., and from Sturt and Du\er Gardens where tri'ts Hud been 

hea\d\ attacked during the precceding weeks (— ~ information unavailable), 

Collection Sites 



Dover Gardens 

Total number ot minus examined 
% already emerged , , . . 
Viable P. fmggatti 
% larvae, pupae & pn*adul(>. . . 
Causa of Death Unknown 

f v larvae and pupae 

% preadults ... 

% roLti 


% parasitized by 

Cirnupilus mur^iwutellum 
% parasitized by Bracon spp. 

% parasitized by others 

%■ Tola) parasitism . . . , 

% max. potential parasitism , , , 











I 2 






Curry (19811. Clearly, at least sortie Cirrospilus spp. 
arc widespread in Australia. Here we record C 
margiscuitdlum from the Adelaide region and C. 
<}Ciipiti\ from Perth (material in ANIC) tor the- first 
time, indicating that these species are distributed across 
the southern part of the continent, al least. 

C ttwrgt battel 'I ton was by far the most commonly 
reared parasitotd associated with P. froggatti in \he 
Adelaide region (Table 3), and was present in mines 
for nearly the whole rime that host larvae and pupae 
were available (late September to late May) As shown 
by Farrell & New (1980), laboratory observations 
confirm il to develop as a solitary primary parasitoid 
of third insiar larvae to late stage pupae, as well being 
hyperparasitie, in the Adelaide region, on the larvae 
of Bracot i spp., LitpchmtA sp.. Dtuulomorpha sp. and 
small conspecifie larvae occurring in the host mine. 
When multiple P fwggam were found in mines, with 
some parasitised by Bntcori spp.. the iatter appeared 
to be preferentially parasitised by C margiscutellum 
rather than unparasiliscd primary hosts. 

Usually only one Cirrospilus larva was (bund feeding 
on a host and generally il did not confine its feeding 
to one feeding site, as indicated by numerous dark 
melanjsed spots found over the host's integument. In 
the laboratory. C. tnargiscutelluni held at room 
temperature took 16-17 days to complete development^ 
once emerged from the egg. Mature Cirrospilus larvae 

could be recognised from those ol the other common 
parasitoids In the complex by the absence of long scute 
over the body and the presence of ampullae on both 
the dorsal and ventral surfaces. The larvae of different 
species of Cirrospilus could not be separated. Adults 
of Cirrospilus spp. can be distinguished from other 
eulophids in the complex by the presence of a 
submedial groove on the scutellum, stibmarginal vein 
with at least two bristles, and their metallic and'or 
striking colour patterns. C. margiscutellutn is the only 
member of the germs in this complex to have the face 
with transverse black bands. 

C. (/capitis Girault & Cirrospilus sp. 3-5 
FIGS 20, 21 23-26 
Four other species of Cirrospilus were reared from 
P. froggatd mines, all of which are solitary primary 
ectoparasitotds. with the exception ol* Cirrospilus sp. 
3 which was found to be gregarious. C. occipitis and 
sp. 5 were observed to feed on host pupae, and sp. 

3 and sp. 4 on Ihe larval stages. C. ocdpitis and sp. 

4 were found in mines over the summer months, early 
December to late March, and early January to late 
March, respectively, while sp. 3 and .sp. 5 were 
recorded from only a tew samples m one week in early 
March and late March, respectively. The live species 
encountered here can be separated relatively easily by 
their distinctive colour patterns. 

hig* 5-9. 5-6. Fore and hind wings. 5. Pomphylax sp., 6. Brat on rnnfusiis Austin, y ; 7-9. Lateral view of whole body: 
7, Bntchymt'ria sp.; 8, Eupelmussp., 9, Elasmus austratiensis OirauJt. Scales: Figs 5. 6 — 1.0 mm; Figs 7-9 = 0.5 mm. 
Abbreviations: bu = bullae: he = hind coxa: hf == hind femur: nip 5 mesopleuron; pp = prepectus, pv = pustmaiginul 
vein; sv - sligrnul vein. 



Figs 10-17. 10-11, Lariophagus sp.: 10, antenna; II, hind leg; 12, Pediobius sp., dorsal view of rnesosoma and metasoma; 
13, Apleuwtropis sp., dorsal view of scutellum, propodeum and mctasoma; 14 15, Diaulomorpha sp,: 14, antenna; 15, 
hind leg; 16-17, Fore wings: 16, Chrysonotumvia sp. 1; 17, Chrxsonotomyia sp. 2. Scales: Figs 10, 14 = 0.5 mm; Figs 
II, 15 = 0.5 mm; Fig. 12 = 0.5 mm; Fig. 13 =250 /xm; Fig. 16 = 200 /im; Fig. 17 = 0.5 mm. Abbreviations: cl = clava; 
ev =t cubital vein; f = funicular segments; pe ■= pedicel; sc = scape; smv - submarginal vein. 



Figs 18-26. 18-19, Cirrospilus rmrgisvutelti&n (Girault): 18, Tore wmg; 19, anterior view ot head showing transverse banding; 
20-21, antennae: 20. Cirrospilus occiputs Giraull; 21, Cirrospilus sp. 4; 22-25. Cirrospilus spp., dorsal view of mcsosoma 
showing colour patiern: 22, Cirrospilus margiscutellum (Girault); 23, Cirrospilus occipitis Giraull; 24. Cirrospilus sp. 
3: 25. Cirrospilus sp. 4; 26. Cirrospilus sp. 5. dorsal view of mcsosoma and metasoma showing colour pattern. Scales: 
Figs 18. 19 * 0.5 mm: Figs 20, 21 = 200 *mi; Figs 22. 23 = 0.5 mm: Figs 24. 25 = 250 ,mi; Fig. 26 = 0.5 mm. 
Abbreviation; sg - submedial groove on scutellum. 



Key to the parasilnirls of Phylaaeophaga froggatti 
ht Ihe Adelaide region (based on females! 

J Forr win*: with more than one enclosed cell (Figs 5, 6j; 

prepectus absent . . . . . . ... 2 

I-oit wine with no more than one enclosed cell (P|g% 
7, 16-18); prepceius usually present fFigp 8 sM sometimes 
abggfil (Fift. 7i iChaleidoidcu) 4 

2 Fore wing with vein 2mcu present: hind wing 
wiih vein i rn nieeiine R> After Rs diverging Irom 
Sc * R (Fie swlthneumonidae) |tVne wing with two 
bullae im 2m-cu (Fie, 5). mesosculum smo»Mh and 

unscutpluredl ftfityl^tto* sp. 

Fore iVjng wiih vein 2meu absenl. hind wing with V81H 
r-m meeting K^ before Rs diverging fnirn Sc + R 
(Fig. ti' (Bmconidae) . ..... ...3 

5, Vertex, occiput and temples rnosiry hairless except 

lor row or ^nori hairs around margin w t eyC5 and 

posterior vertex (F»x "*> propodeum blftclt 

— Bnir,m phxhtnropfweut Austin 

Vertex oeuput ana temple-- sparsely covered *olh 
ICmiK hair. ( f if. 4^i propodeum orange-yellow 
Braam \im[u\u\ Austin 

4 , Rind lemur en I arged and w if h teeth on ventral edge ( I ; ie 
7); prtpectus very small. \1nuaJI3 absent (Chuleulidaei 
I malar solute dinner, posimarginal vein longer than 
stigrnal vem (Fig. 7): propoedum with rough alveolate 

a ulpmringj _ _ fir,irU\i>\i'fkt sp. 

Hind femur not enlarged and without teeth (k : ivs 8, II), 
prepeetus distinct <Fi^ s, 9) - -5 

.V Mcsopleuron greatly enlarged, convex, developed into 
.( Iflrge undivided shield (Ftg S) [posirna'fiiual vein 
Mightly shorter lhan sligmal Vtsinr ovipositor protruding 
Trom posterior meiasoina, sheaths black wiih noddle 
thud while iFie- 81] . . . , , . liufwhnus sp. 
Mtsopleumn not greatly enlurgcd (Fig. H) b 

6, Hmd eo\a developed as a large flat disc t.Fig. 9), hind 
tibia with stue fanning distinct criu cross pattern ; fore 
wing narrow, posimarginal vein much longer than stigmal 
vein (Fie- 9) (El» mi elite) [bod> dark, tcgUta UoAleg! 
pale except for hind coxae|/\/av*ao anittmssus^ Girauii 
Hind not a.* a large flat disc (Fig. Iljj hind tibial 
hflfrs not toiMiing Lns>-L'ni_ss pattern; lore w»in£ relatively 
tm.;.d (Figs (MS I i 

I Hind tarsi >segnienied (Fig_ II i_ antenna aunoui <hsimct 
lumeular segmenls and eiava (Fie, 10) (I'lcromalidae) 
(head round m anterior view basal third bl tore Wffltf 

\iauall>' devoul ttf hairs] . , .tMrfephogui sp. 

Hind larsi 4-sejrmented (Fig, 1?), antennq wuh dUtinci 
funicular segmem.v Ifld cUva (Hgs 14 20 ?h 
t! ulophniac) .. . X 

X. L>ftr>;il me>OMvna with long stout hair. |f-i^_ U); 
Mii-i;iM_>mit distinctly petinlabr (Fig,s 12, I3», hodv hiacK 

HTid shiny. . - ... . . , , . , ,9 

Ifiklrtal mebosoma >vith hairs then mostly line utd siwrt; 
Mu-Jasoniij scHsde, or peintle less conspicuous (Fip, 26), 
tn.»dy colour variable , , , , l(J 

o Siuullum longiuidinally idii^ose with exception HI 
longitudinal OiOdiat tine which is smooth lF»g ID; Tl 
ot'metasoma with ant^riot tlange (Fig 12); fcg&till bfetpK 
except pniximal 3 tunal segments of mid and hind lees 
which are while . . , , , . . , .PttlioNm >.p 
Seuiellum with curiaccttu.s sculpturing (Fi^. 13); Tl of 
oieUistMTw wiihiint Range (Fig. l3);aHlegsetivtaJ kiCQfluie 
pale in colour ApUwanhjtw sp 

10 ImUmlIc d! anlctmue 3->.egmented (Fig. 14) [body metallic 
gret-n-gnld in colour all legs distal to coxae while, beefy 
Ab(M 2 rnm m longjbl DtmtfomvTpbn .sp 

Funk-lc Hi antennae 2 veflmemtcd [Pt& J(' 2l,i , it 

II. Seutelliiin without suhmedial grooves: submarginal vein 
Will! 2 dorsal bristles (Figs 16, 17): body less than I mm 

in length [ChrvM/wttftmyw A^hmead) 12 

Seutelium with M/hmedial grooves (Figs 22. 23>; 
submarginal vein with more irun 2 dorsal bristles (Fiji 
ISl bod> iiiejter than I mm in length iCirmspiht.s 
Vfetwood) |3 

H- Cubital vein ot lore wing indicated by a xmgh 
miy, Dl'h^rS (Tig- lo); sttgnial vein distinct and intuseatt; 
an»iind distal end '.Fig. lot; me.sosoma and head metalhi 

gTccn . , , Chtvsimautms'ui sp. v 1 

Cuhilrd Vein of-Wlftg not mdtcjted (Fig. 17); stigmal vein 
"i<"i and not int'usca(e (Fig. 17); mesosoma and head 
ntm-rnetallic and darV in eolourC/fo'V''/">'<wv/t< sp. 1 

13 riijva o( antenna eoniinuous wilh funicular segments 

(Fig. 20] ,, , - ..._____ _.M 

Clava sepaiittcd Irofii funicular xegments by distinct 
constriction between .segments (Fig 21).. . . |fr 

14 Stigmal vein int'uscate around distal end, siuuetirnev onl*. 
tiiinlly (Fig, 18); vertex with short sMul black hair* 

(Fig. J0> .. h 

Stiginal vein without intuseationdistally vetiex with >ml> 
fine pale hairs |mesosoma with yellow and metallic eieen 
markings ll-'ig. 2^\\ , , Cirmspilus octipif# Girault 

15 Dorsy_l mesosoma metallic green-blue with slight yellow 
markings in posletior itait' (Fig. 21), surface with 
rein -iiUre sculpturing; face with transverse durK 
sinpCK (Fig. 19); pronotum shorter than seuium 

(Fig 22 1 

CifitmpiiUS marntsrutftttittt fGiraulO 

Pnmotum yellow with 3 longitudinal black bands, resi 
^' mesosoma orange with black marking* (l-ig- 24) 
surface Willi eoreaeious sculpturing', pmnotum nearly 

as long as sculum ... „ Cirrrnplfus sp. 1 

16 MeUsoma rounded in dorsal view; pronotum yellow 
oranee. rtM ot dorsal mesosoma orange with black 

marking.-. (Fig, 2o,i . Cint>spilus ftp, S 

Metasoma broadly elongate, pointed posteriorly , 
dwrsal mesosoma yellow with black markings (Fig. 
25) Cuwspifus sp. 4 


This study confirms a number ol aspects of the 
biology ol t- \froggaui ftrsi reported hy Farrcll Sc New 
(1980), 45 well as documenting the fecundity and 
longevity ot adults, and details of~its parasitotd fotnplex 
for Ihe First time, deviously, this insect had been 
recorded only as a sporadic pest of eucalypts in the 
Adelaide region and then mostly on ornamental tnics 
However, the status of P. ftoggaiti as an emerging 
serious pest both in Australia and New Zealand has 
increased dramatically over the last lew years and (his 
is correlated with a corresponding expansion in native 
woodlot plantings in south-eastern Australia. Indeed 
Ihe Engineering &. Water Supply Department of Soutb 
Australia has rated P. froggolli as one of the most 
important pests at its Bolivar woodlot (pers, eomm. 
S Shaw). 

Dunng this study we recorded nearly three times 
the number of parasitoid species associated with P 
ftoggatti as any reported by previous authors (Riek 
1955; Farrcll & New WSOfCurry !9R|). The M/e Oj 
the parasitoid complex in the Adelaide region, howevei, 
is unlikely to he significantly larger ihait elsewhere in 



Austialia. Rather, the concentration of this stud\ on 
rearing parasitosis and its longer duration has mean* 
lhal several rarer species have r>een collected. AJmj, 
the taxonomy tff the gamps involved is now heller 
known (sec in particular Boueek 1988 and Austin & 
PanKis 1989), so that species can be more easily and 
reliably identified. However, the makeup of the 
parasitoid complex and relative ahundanee of species 
does seem to differ between regions. For instance, in 
the Adelaide region Ctrro.spihu sp. I was far roorc 
aKindani than B, phyhicteophuxus, while in Molhoome 
the latter species was more abundant tFarrell & \ew 
|9H<Ji. In Western Australia Curry (WHO reared six 
paranoids from Phslacteopha?,u (three Cirrosptfus 
spp. , Elastics sp. . Enpelwus sp. , an unnamed culophid 
and Apnntch's sp. > bul no firm on, Aushn & Faulds 
(1989) have pointed out that the reeoitf of Apaitteh:* 
sp. rnusi he erroneous as members ol this genus and 
all microgasirine hraconids are end\ypaiasitoid i - ol 
lepiilopieran larvae The only major parasitoid tason 
not iccordcd in this study is isopluioides \\vsiruh\n>us 
Ginmlt, a pieromalid, which has been reared from P 
froifWttt only in Western Australia I Boucek 19881. 
Parrel I <te New (19X01 suspected the Cirmxpihn sp 
reared by them in Melbourne to be polyphaguu* and 
to use alternative hosts at different limes or the vear 
to EWttd continuously. Although we did nnl rear 

parasitoids from other euc:,Iypt leal rmnim/.s In ihe 
Adelaide region, the hid the multiple Ctnosfnhis 
species have often been recorded from other hosts does 
provide some indirect evidence that metubets uf litis 
paiasitotd genus are polypfmgous (pers. comm. N, 
Gough. (. D. Kallmann), 

This study should provide a solid basis lot more 
detailed work on die ecology of R /feggQft/i 
particularly ihe factors that influence its abundance, 
means of overwintering, and the susceptibility cjT its 
eucalypt species. These and other lactors will be 
important in understanding how K jro^uttt develops 
as a pest and how best it can be controlled- Programs 
aimed at limiting the damage caused by this sawfly 
will need DO take into account the mortality caused by 
the above parasitoids. In this respect future research 
might profitably examine both seasonal and regional 
differences in rates of parasilism and the species 


We lhauk Vr John LaSalle, International InMiluteol 
Entomology. London for confirming the identity of the 
ehalcidoid parasiunds, and Mr I'aul Daugetfield fot 
|bC hne drawings, electa^ micrographs and hi* 

cvmtneuls on the manuscript 


Am'N i 19501 The leaf bhsici sawUy * Phyiaaeophaw 
tucahpu. The AgfttiiltUml Gareitc 470 47t. 

All IN, <J. R. (1V9UI Uruba lupins Walker (Lepidnpt/ra: 
Novtuidac) larval survival and parasitoid hiolngy in the 
field in South Australia. J AW t'tit Sot 29,^01-312 

At sun. A. D. & Au.r.K. G. R. 1,1989) Parasitoids of Vtaba 
fuf>t>m Watker '.Lcpidoplcra NoUmdae) hi South Australia 
with description ol two new species of Braeorudae. Tmn.\. 
K Stn S AUXt 113, I69-|«4 

&. Pai'I.i>s W. (1980) Two new Australian sprues 

ol Bruroti I* (Hvmvnoptcta Bracouidac) parasitic on 
PkyktcfcvBbagappa, {llymenopteni: fcrguiaet. ./. Aum *nt 
Sue 28. 207 211 

BfcNSON. R. B. (1963) .Some new Western Australian sawflies 
ot ihe .ind Phvliictcoptiauinai'' (HyMCfW *i:i 
Pergidael. / Rty Soe\ W Auxt. 46. SI 84, 

BiNOi-v, R. (I9H6) Insect pest*, pp. 233 24) /" Wallace. 
If R rf«l \. "The KCcfojE) of rlu ftwc&Uf and Woodlands 
iif South Australia," tGovl. Printer. Adelaide). 

U>hC>k. 7 M988) "Australasian Chalcidoidea 
iHymcnoptera). A UiasvNtL-mutic Revision ol (jenem of 
Fourteen families, with a Reclassification ol S\\n w.\" K'AR 
lnieiii:ilunial, W*!il|iii):l'»iu'l. 

Cl'ftHY, -I. S, tt9KI) Niilivi- wasps hit inirtxiuceJ Hw pvst 7 
V< # 4«af 22. 61-62. 

r-VKklLL. O. S. &. Ni.w. I R. U980) Some axpCCli M the 
biology ol the eucalypt mining sawlly PhylaijunphQftti 
JmxnulU Kick (KyiticHoplera: f'ei]gid3e). Aust / Ami J8, 

F\ti r*. W. jfiPStn rntixxluction into New Zealand of BntctHt 
fthviacii-opfutsu.y. a bioeontml agent pf Phytat ie ophugn 
jroMiW, huahplus leal iiiliimy \nwl1\ N / / ffoftil 
,SV I 211. ^4-64 ' 

Frijggatt. AV- W. tl899i A new genus and species of suwflv. 

Proc. Linn. Stn: NS.W U, ttO-134. 
i !ai i f>. |. I"J, U t )K4| "An introduction to the Ichneumon idae 

of Australia"' iBntish Museum (Natural History] London). 
& Boiton. B. (Eilsi (1988) "The Hymenoptera; 

(British Museum (Natural H<sbiry|. London and Oxlonj 

I'nivcrsity Press. 0,\tord). 
Kav. M K {l l >X6i Phvttniropha&i Jfo^uii Ru-k 

(Ilvmenoptfra: Per^idae) f:ural\ptu.< lc;if tnmitie sywdv 

Htrr\f utul /tml/tt fnstcte in N&w /auttirui No M 
Mv/ASl-i', / (\ l )HX) Immature stages arid lltC '"\loiy ol 

Chrysoiiujotnyia sp (Hymcrioptera. HuhtphtUae). a 

porasilticnl ol rhc (arrah It^fntiner, / Autr em $i* 27 

279 2M2 
M'>)Ri, K M 0Q66J Observalions on some Austiulnm 

forest insects 22 Notes on some Australian leaf-miners. 

Ami frvl tf. 303-34?, 

NaI'M'VNN. I. U. |W83» A new g^nu-. t-t Phylueteophaginae 

from northern Australia with a key to the genera oi the 

suhfamily (Hvmenoptci*a TenthrL-dinoidCii Pcrgidaei. 7. 

Aust enr. Sat, 11. 237 242 
NuiiALL, M. J (1°S5> New insect pest atmcLs eucalypt* 

Phyfoctrophaqo frog$attj (Hymenopiers: Pcrpidyef The 

\'i-w ?*uthtu) thtrtwr 106, 11*4-5. 
RihK. E f (1955) Australian leaf ntininu sawUies ol the 

genus PhyjactcOpliasa (Hymenoptera: Tenthredinoidea) 

Au\( ./. '/<n>i \ V5-9» 
<l%7) Ausltalian Hymenoptera C'lutlvuKuvlci* l;«ntily 

Dulophidae, subfamily Elasmidac. JbiJ, i5, |45*W. 
_ (l 4 >7t)i A rede rin it i on of the roihf»»mily 

Phyiaeteophagmue wiih (K-st nplum nt u new genua and 

species (Hymenoptera- Svmpnvta fVreidae). / Mist em. 

W. *>. 215^218. 





ByD. E. Bayliss* 


Bayliss, D. E. (1994) Description of three new barnacles of the genus Elminius 
( Cirri pedia : Thoracica) from South Australia, with a key to the Elmmiinae. Trans. R. 
Soc. S. 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 S, Aust. (1994), 118(2), 115-124. 



by D. E. Bayliss* 


Bayi.iss, 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), 115-124, 31 May, 1994. 

Three new species of intcrtidal barnacles from South Australia are described. Elminius flmdersi 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 Words: Cirripedia, Elminius flmdersi sp. nov., Elminius placidus sp. nov., Elminius erubescens sp. nov., 
Elminius modestus, Elminius adehudae, Elminius covertus, intertidal, South Australia, taxonomy. 


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 Popes (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, E. 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 Finniss Street, North 

Adelaide, South Australia 5006 
Current address: 7 Thomas Street, Kingsgrove, N.S.W. 2208 

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 Elminius lacking 
a spur. 

This paper describes three new species of Elminius 
which, with E. modestus and E. adelaidae, brings to 
live the number of species of this genus found in South 


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 




2 mm 

Fig. 1. Etminius jHndersi sp. nov. holotype. A. External view ot shell; B. inner view of shell; C. seutum, internal view: 
D. lergum, internal view; E. scutum, external view; F. tergum. external view; G, body; H. labrum; I. mandible; J. maxillule; 
KM. cirri Mil; N. middle segment, posterior ramus, cirrus VI. 



Paratypes: SAM C4243, WAM 121-92; D. Ba> -"tss, 
21.vii.m92, same locality as holoiype. 

Desrripfion oflu>hiype\ Shell (Figs IA. IB): conical, 
while (b grey, opaque. Purieties folded. Orifice 
pentagonal in outline, width H length, cartnal sides 
almost twice as long as lateral sides, rostral ->ide 
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 

Opcrcula (Figs 1C. ID, IE, IF). Solid white to grey 
in colour. Scutum longer than high, basal marcin 
slightly convex with slight upward turning at tergal 
comer, cresUs for depressor muscles absent, articular 
ridge, articular furrow nuxlerately developed, adductor 
ridge not apparent, adductor pit taint, externally growth 
ridges present. 

Tergum hatchet shaped, vertical articular ridge 
folding inward to form centrally limited ibid, articular 
margin genllv curving in apical region from eaitna! 
margin, becoming parallel with basal margin to form 
long narrow handle, basal margin curving sharply, 
almost at right angles, to Conn projection with cat trial 
margin, projection with prominent, deeply carved 
crests lor tergal depressor muscles, spur confluent with 
basiscutal angle. 

Body (Fig. 1G). prosotna white, broad, rounded 
Cirri light reddish brown in living specimens, cirri 1, 
II and III slightly darker than cirri IV, V and Vf, colour 
lost on preservation. 

Mouthparts (figs IH-IJ): Labrtim with three teeth 
and setulae, on each side ot central notch, Mandible 
with live teeth, lower short pectinate edge terminating 
in short curved spine. Maxillule with two large spines 
above notch, live smaller spines in notch, tour large 
spines below notch, smaller spines at lowei an^le. 

Cirri (Figs IK-IN) Cirrus I with anterior ramus 
x 1.5 length ol posterior ramus, segments broad. 
Slightly protuberant. Anterior ramus of cirrus II sjigbtly 
longer than posterior ramus, segments slightly 
protuberant anteriorly. Cirrus III with anterior ramus 
slightly longer than posterior ramus, segments slightly 
protuberant anteriorly; setae on both rami extremely 
long, stout pectinate setae on six distal segments of 
posterior ramus. Cirri IV to VI all subequal in length, 
segments with three large pairs, two small pairs o1 setae 
on anterior lace, small proximal pair oJ setae between 
segments. Number of segments in rami of cirri t-VI 
shown in Table I- 

Penis: Penis as long as cirrus VF setose, baMuVi sal 
point absent. 

timation: Shell can be tubular, conical or flattened, 
Parieties can he smooth or undulating with variable 
number ol longitudinal folds. Maximum basal diameter 
17mm. Colour varies from white to light grey. 

Eiymotoxy: The species name refers to the 
FHndersian biogeographic region. 

Compuristm with other species: Elminius jlinderst 
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, 

£ flindem cannot be reliably distinguished from E. 
modestus on the basis of external shell appearance. The 
opercular plates are. however, quite different, The 
tergum (Fig. ID, IF) is harchet shaped with a long 
handle and an articular furrow which is restricted to 
the central portion. 

E moderns (Fig. 2B, 2D) has a deep articular fold 
running from the apical end to almost the basiscutal 
angle. The articular margin, unless worn, is straight. 
The scutum (Fig 2Cj has a grey bund, but this feature 
is not always apparent. 

7 \bl_t 1- Ctrrut counts. Elmirnus Hinders] j/A <rt0w \imterior 
nmiux Jlrsi), 

Basal diameter 





in iv 

V V| 

\[){\ (holotypei 



12.11 22.20 

24.25 26.27 




12.11 18.17 

21.20 23.22 




!?.!> 21.20 

23,24 26.25 




M.I3 23-,23 

27,27 21 JH 



11.10 22.10 

24,23 23,23 





-. i 


Fig. 2. Ehmnius m<Hh\lus Darwin. A r scutum, internal view; 
B. tergum, internal view. C. seutum, external view; D. 
tcrgutri. external view 


t> t k-\> • |&5 

h. tnatipiw also can be distinguished from other 
member- ot' the gemis in lacking peuinare i i. tl or. Hi. 
prfsteritfr ramus of cirrus [IJ The npcreutai flstpd arc 

pUtc wrvte with a grey hand at the iostral end and a 
"i. ill orange Jot centrally fi. flinders' has ;i dark 
ho»wn hand a! the rostral end and another brown b.mO 
centrally 1 on Haps which arc dull while. 

B. tulelttidtie tBcOyUss IMHK Fig 2B) ha* a very small 
.mi* ular furrow, which may not be visible il the tergum 
is viewed direciiy frOW above Tfic Sftetl tS JUO^l 
thinner. translucent, and is light brown The opercular 
flaps are, however, too close to £ lltndersi fa 
apptarahee to be useful lor identification. 

ffohirm, IJminiusflimU'rsi is found in the tpttltlvial 
/one iia waters sheltered from direct wave action It 
Is found in habitats with stronger water How and rtwre 
turhulcnec than other members of the Genus in South 
Australia. In Ihe £ull re^iorr, where w.oe impact is 
diminished it ocurs on exposed rocks, If ^rows p 

lirgg m/cs on jetties jlthough il is not found un surfaces 
facng directly into waves at more exposed localities. 
Outside the jmlf regions it ts found in habitats prcHectCd 
from oceanic waves such as in boa! enclosures behind 


Settlement occurs on a wide variety of surfaces 
including roekv cement, wood, metal and plastic. Il 
iv rarely found in maneroves where water flow is 
gentle. At Port Pirie it can be found DP fl>c|tfi in the 

strongly flowing sections uf'the lidal river, but nol m 
nearby mangroves 

Disltihutiou: Elminhi.s flindersi is widespread til 
■South viMi.fhi hum Kangaroo Island toCedunaand 
the western Byre Peninsula, and occurs in both gdlft 
(r% 3} In the Spencer Gulf it can be lound at fori 
w.nsu indicating thai il can tolerate the wide salinity 
and temperature range found in .South Ausrrabun 

F%. \ Lhstrihuln'tiol / ~ht,!ni,4: ffwhvM pjtftef||, t'Mmtty '('/W.//>/.w H:t_vliv-. . Mmmm\ jlimUtsi \p 0<w h'lmhuus /u'tu ntn\ 

sy tins jntl lUttiimu* a^btsvvm *-P nov in Stfufti Australia, 



Elminius ptucidus Ip, nov 
FIGS 3. 4 

ShUWHi MftfetffW Womerslcy Sc Edmonds J 1(58: 
24.1 Thomas & Fdmonds IM79; 161, Hutchings At 
Rcchcr l ( iK2: 95, 

llnlolspe: SAM C4244. on branches of AvtWWifl 
w/WM/mCYvwcli, South Australia. 33**8'S. IftWL: 
6. Kayliss. 6ii.|092: dissected ipartially). 

Paratopes; SAM C4245, WAM 728-92, D. Ba> ! iss, 
6 n M82, same locality as holotype. 

D?\(,riph<>tt (tfhttjttnpr: Shell (Fig. 4A, 4&V up 
conical, prey to while with thin brown and dh] k gf^y 
bands parallel to bt'9£ Paricties gently folded. Shell 
thin and nearly translucent. Orifice large, pentagonal 
in outline, rostral side concave and broad, eannal sides 
longer than lateral sides, width % length. Basal outline 
sinuous. Alae wider than radii with less oblique 
summits. Radii with oblique summits, narrow: radii 
and alae with prominent growth lines. Shell 7mm in 
basal diameter. 5mm in width, 5mm in height. 

Opcrcula (Figs 4R-F): thin, translucent, white in 
colour, grey margin near articulating margins cl 
internal surfaces of terguiu and scutum. Scutum longer 
than high, basal margin slightly convex with prominent 
depression tor depressor muscles, no crests apparent, 
articular ridge, articular furrow miKierately developed, 
adductor ridge, adducior pit not apparent. external iy 
growth ridges present. 

Tergurn wedge shaped, articular margin and basal 
margin gently curving from apical end. ctwveryirre On 
basiscutal angle, articular furrow shallow, apical portion 
large with small, feeble crests lor tergal depressor 
muscles; spur confluent with basiscutal angle. 

Body (Fig. 4G); Prosoiha white, narrow, elongated, 
palps and chti I lo III very dark brown, cirri IV to 
VI light brown with dark brown pigment along and m i 
(vice ">f segments, cirrus T covering oral cone. 

Mmiihparis (Figs 4H .1*: Labium with three tefitfl 
and setulae \m each side of central notch. Mandible 
with bve teeth, lower pectinate edge terminating in 
-4i"n 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 4K-N»: Cinris I with amenor ramus %\ 1 
length of poster inr ramus, anterior ramus with broad 
proximal segments, narrow distal segments, with long 
setae, segments slightly protuberant- Anterior Tamus 
of cirrus IJ slightly longer than posterior ramus, 
segments slightly protuberant anteriorly. Cirrus 111 with 
anterior ramus *:J.25 length of posterior ramus. 
segments slightly protuberant anteriorly, setae on rami 
very long, stout pectinate setae on six distal segments 
of posterior ramus. Cirri IV to VI all subequol in 
length, segment with four large pairs, two small an:- 
OfSCtiLC Oti .interior face, small proximal pair of setae 
bclween segments, anterior face With dark brown 

pigment. Number of segment* in rami of cirri 1 10 VJ 
^hrjwfl in Tabic 2. 

T\)it-i:2, Cufal totmiA, F.liniiiuis pKK-i'Jus >/' " m biohn.n 
titmtiA Jtrxt), 

Basal diameter 



111 JV 

\ I 

raiMatypH) 14.H HU0 13. ti 27.24 socfjmP 

4 14 f, 9JH 12 12 24.21 26.25 31.29 

■i.o 12.6 10.10 114 1 &33 2%2* 30,26 

J 14,,' I'.),'* |4il.l 23*23 27 27 2B.27 

BJ1 l-i.K 10.10 14. M 25.24 28 23 36,21 

Ferns: Perns as fnflg as cirrus VI, setose, basidorsal 

point Hbscnt 

fariati&in Shell Is usually upright conical or tubular 
and rarely flattened. Shell may have visible banding 
but X It) magnification fcs otu u required to see the dark 
and light bands. On mangroves the barnacles nav 
appear solidly erey. Some specimens collected from 
Toeks have dark grey bands and ihe shell may appear 
almost bluish Thomas tc Fdmonds (l979jj described 
ihe shell as heuig 'bluish-green' Some specimens have 
a slight pink flngjg towards the top of rhe punches. 

The tergum is cKirernely thin near the basal margin 
and wear may alter the shape. In some specimens the 
tergum is virtually triangular with the basal margin 
u-iy gently curving from the tergal crcsis to the 
basiscutal angie, On unworn specimens the external 
sudace of the tergum is grey with a white area at the 

apical i?nd. 

The scutum in many specimens has a nearly straight 
basal margin and the articular margin is almost at right 
angles forming a right angle triangle shape, The 
depression lor the depressor muscle is usually well 
formed with the shell being extremely thin in this area 

hjynmlo^w The species name is derived from the 
Limn ptuiuiu.\ meaning calm, tranquil with reference 
to the habitat of this species. 

Comparison with olhcr sprats: The shell is thin and 
translucent wilh narrow dark bands. Other specie- afC 
umfoim in colour and lack the alternating light and 
dark banding. The opercular plates are thin with grey 
margins internally alotuj the articulating margins. 

The iergum is distinctive i \ sh;ipe. The apical reeu-p 
i> lar^e with vety leeble crests The articular margin 
and the basal margin curve gently to the basiscutal 
angle forming a triangular or wedge shape. 

The prosoma is narrow and elongated us arc the c irri. 
The dark colour of the cirri contrasts with the white 
prosoma. The coloration survives preservation. 

The opercular Haps arc cream with a dark black band 
at the rostral end and another black band centrally. This 
enables it to be distinguished Irom t, modestHS, bul 
it cannot be readily distinguished from other South 
Australian species which have die same pattern of dark 
bands: on iighter coloured Haps. 




AB . 

CDEF v_ __, 

G . 
KLM . 

— ' 

HIJ , 

1 mm 


Fig. 4. Elminius placidus sp. nov. holotype. A. External view or 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. maxillulc; 
KM. cirri T-1U; N. middle segment, posterior ramus, cirrus VI. 






' ' 


u . 


1 mm 

D 2 mm 

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. labium; I. mandible; J. maxillule; 
K-M, cirri Mil: N. middle segment, posterior ramus, cirrus VI. 


at- IJAYM&S 

Uahihit E h'ttnih i pfaGtdUS (5 Ml internal species 
which is Iburtd in sheltered hatmats It is highly 
abundant in mangroves in Spencet Gull It settles nn 
maiuliesandpncumatophoies, hut is larely found flfl 
leaves, It is not restricted to mangroves hut also seltFs 
un rucks, cement. wood, nietal and other surfaces 

lh\tnhiiti<>t<\ f-'hfiiuitts }>la< >>lus is widespread, befflg 

found un Kangaroo Island Vorke Peninsula, Spcncvi 
Gull and Hya Peninsula (Pig V). It is nol lound in 
the mangroves north ol Adelaide which are occupied 
hy / tuit'louluc. 

Elminius etuhescens sp no\ 

PIGS 3. 5 

lilmim'us covering Foster 1982; 26. 

Holotypc: SAM C424o. collected on juhhci tytvs 
al QjcilClg boat ramp, Adelaide, SoUfll Ausindi,, 
34 tM io'S, 138°36'F, D. Raybss. 19.v i.WI. dissected 

Paratypcs: SAM C4247. \VAM 729-M2. D Bayliss, 
&«J tWt ffolfl the same locality ag holotvpc. 

Dru tipit'on of ihe holotypc Shell (Figs 5A. 58): 
flattened conical, translucent with red coloration, 
appearing datk purple before ihe body amoved, colour 
vrevish purple when preserved Parieties gently folded 
Orilice small, pentagonal in outline, width 34 length. 
lustral side short and sliaight. othei sides almost equal 
in length Basal outline >inuous, A lac wider than radii, 
wtlh lev* ohl n(iie summits. Radii with oblique summits, 
narrow. Shell 8mm in nasal dratnater, 7mm in width. 
3mm in height 

Opeivula /Figs 5C-F): Thin translucent, reddish 
ere\ ish purple with white areas when preserved 
Scutum longer lhau high, hasal margin slightly come\ 
with upward turning al tergal corner, articular ridge 
articular furrow moderately developed, apex rellecied 
outwards, externally gtowth ridges apparent. 

Tcrgum wilh articular ridge folding inwards to form 
well developed lurrow, articular margin concave, hasal 
margin curving sharply to form projection with carinal 
nurgin, well developed L resis lor tergal derpe.ssor 
mij.clcs. spur confluent with basiseutal angle 

Body (Fig. 50»: Prosoma light bruwn. palps and 
cirri I and II with some dark hpiwn pigment, cirri 111 
to VI clear partially light brown preserved material 
with body and cirri almost uniformly light hrown, 
prosoma and cirri elongated, cirrus I overtopping orjM 

Moulhparis digs 5H-J)* Lahrum with three tetrih 
and selulae pn each side of central uoMi Mandible 
Willi live teelh. lower short pectinate edge terminating 
in short uirvcd spine Maxillule with two Lure spirit 
helow nnleh. smaller spines al lower angle 

Cirri {Figs 5K-Ni Cirrus I wilh anterior ramus -1-5 
h-ngih of posterior ramus, segment* sliently 
(..oMUi.inr Ant.-riur ramus of cirrus 11 slightly lOOgfcJ 

i i posterior ramus, cements slightly protuberant 
anteriorly Cirrus III wilh anterior ramus slightly longer 
than posterior ramus, segments slightly protuberant 
amei ioi Jv . setae oi\ both uimi long, stout pectinate setae 
on six distal segments of posterior ramus. Cirrus IV 
to VI subequal in length- segments with three large 
p.iirs, tWO '-mall pairs of setae on anterior face- small 
proximal pair of setae between segments. Number of 
segments in rami of ciori i to VI shown in Table 3, 

T vlil I 3 (Jrral ctmnL\: Elminius crubeseens sj>. rttn 
tanwrwr minus fir.v.t. diameter 





H.O (huloivpiM 12.7 9.9 11,10 21.24 25 24 2$, 30 

7 14.6 0.0 I2J2 25.28 2921 32.31 

/ fl IMS 10,8 11,11 23,20 25 24 25.24 

7 s I3,fi 1 1. 10 12.11 20, 19 23,23 25.2s 

9-0 t2,ti 9,9 13.12 2,3,23 25.26 30.27 

Penis: Penis as long as cirrus VI. seiose. hasidorsal 
point absent. 

Variation. Minimus vrubtscens is usually flattened 
although tubular and upright conical forms can occur. 
Hie shell often has baud longitudinal folds which vary 
in number between specimens. Maximum basal 
diameter l2mn). 

h\ttmlo£\- The specific name is derived from Ihe 
Latin enthescerc, to blush with modesty, in reference 
i.i it$ redness. 

Cl >mpunst>n wtfjb wr spcacs: Llmimtt.\ emhescens 
can be easily distinguished from other species in South 
Australia by the rvd coloration of the translucent shell 
which, in rhe field, appears dark purple. The shell is 
Otherwise- close to £ tttodtStMS, It lacks the ridges seen 
m fi cttvenus, although worn specimens of the iwo 
species aa- similar in appearance. The tergoseuial Haps 
fire straw to pale yellow with two dark brown to black 
hands. A small portion cd the Haps between the tostial 
end and the first dark band are while, h. covertus has 
six pairs uf dark spots on white tergoseutal flaps. 

Hhbttdt: Elminius eruh€JfcctjS is common in sheltered 
waters in the Adelaide region although ii is not found 
m mangroves. It settles on a wide variety of surfaces 
including rOCk, a'nicul. wood, metal and mhher. If 
ft Ihe highcsl barnacle in the inlcrtidal 7.onc. There is 
hrtlc overlap with /•". modzftnis which forms a tone 
tielow it On aides it is usually found on surluces which 
are overlapping ot do not face directly into the 
afternoon sun. although it can stand exposure to 
sunlight for pari of 'he day- E adfhiidnv avoids sunlight 
and is found under nuks which are lower it) the 
iriMUdal zone, 

Otsrrihution. This species is very common in the 
Adelaide region- bul was not found elsewhere in South 
Australia (Fig. 3 ». Its distribution outside o\' South 
Australia is unknown. 



Key to species of the Elminiinae 

Shdl Willi Ih.II 

comparuncntal plates . . , 

Shell wiili xi \ 

compart mo nta I plates ........ 

Teryuni with spV/ 
confluent with baMsculal 

ungk . , . 

Tergum with distinct spur 

Shell white or xr&y . 
Shell not white 01 grey 
Shell while, u-rgurn with 
straight articukn margin, 
deep articular furrow . 

Shell greyish white, 
tergum hatchet shaped. 

concave articular niaruin. 
centrally Idealised furrow 

Shell grey; narrow daik 
hands alternating with 
white Icrgum wedge 
shaped i weak crests. 

shallow furrow 

Shell light brown, tergum 
with small lurrow 
restricted lo apical end . 

Shell translucent with ted 
coloration purple in 
licld, IcigOrn wilt 
concave articular margin, 
deep furrow 

Shell bull red With CftAftl 

ridges, (erg urn hatchet 
shaped . 

Shell pale brown with 
reddish bmwri bands 
between low ribs, tergum 
with spur longer ihan 
bauscutal angle 

Shell greyish cream wilh 
darker radial bands, 
tergum with spur not 
longer than h<tMxcuU«l 

Ehmnius kitisii Gray 

(South Aniern.ii i 



E/nwuu\ nunUwfus 
Darwin (Australia. hK W 
Zealand. Kumpe, Snub 

Ijfwnnt.* flintier \i >p nnv 
(South Australia! 

Etminitt.s phirUhiS sp uov 

(South Australia) 

Klmmius aJcta'tdac 
Bayliss (South Australia) 

Etmtttius vnihcMttis sp 
nov. (South Australia) 

Ehuinius c<wrtu\ rosier 
l NSW. Western Austutno 

Hexaminuis fblionwt 
Anderson, Anderson & 
ton (NSW) 

Hf\4irriiniu\ juiptintui 
Fusk-f (NSW) 


The diversity of extant species, its well as tonsil 
evidence, suggests that the F.lmiminae originated itt 
south-eastern Australia (Buckeridge pR2, 1984). The 
species present In South Australia are, with the 
exception of Elminius modestus, not found itt NSW 
The genus Hexatmnhts is not represented in South 
Australia and Ehnmius an'frrus is also absent. 

Souih Australian barnacle populations are 
geographically isolated from the eastern States bv a 
long expanse of 'coastline, from Robe lo Cape Otway 
in Victoria, in which mtcrtidal species are virtually 
absent ( Wor iters ley A Fdmonds 1958). In South 
Australia there are extensive ateas of coastline wfuch 
are protected from oceanic waves in the gulfs which 
Contain ideal liablGUS for EJmmius. Speciation may have 
occurred in the variety of sheltered habitats which are 

It is also possible that species from elsewhere along 
the southern coastline of Australia have been 
intnKluced. At present the distribation of Elminius 
species in other Stales has not been determined. The 
presence ol F. vowrtus in Western Australia (Foster 
1982) has been confirmed bv Jones (1990). Il is possible 
that it was introduced from eastern Australia. E, 
modestus may have been introduced from New Zealand 
(poster 1982; Flowerdew 1984). but an elect rophoretic 
i omparison involving South Australian forms lias yet 
to be done. 

Al present there is a large scale 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 Ehmnhts are extremely abundant in 
mangroves in South Australia, but their distributions 
do not overlap, E udtfunktr is found in the mangroves 
north of Adelaide whereas E. pltutdus is found m 
mangroves in the Spencer Gull The two gulfs have 
considerable differences in ihcit marine invertebrate 
fauna (Shepherd &83jj 

Only E. adelmdac utilises the leaves as well as the 
branches and pncumalophorcs. Anderson et al. (1988) 
have described a species ih.xamimusfoliorum, which 
is specialised Ecu living on leaves. The adaptations they 
list as important for this species, which include thin 
shell, thick basal memhrane long elongated cirri and 
rapid cirral beating, are also found in E. adeluiduc. 
Nevertheless. £ culekudae grows to a much larger size, 
is found in other habitats and has a larger variety of 
cirral beating patterns, including ihe ability to hold the 
cirral fan fully extended. J5 phtadus has a mure 
restricted ranee of cirral activity and lacks the ability 
lo beat rapidly. The basal memhrane is also thinner. 
Its thin shell and elongated body suggest a Bpecfies 
adapted for very calm habitats, bur not necessarily 

Jn New Zealand £ modesuts is reported lo live in 
mangroves (Moore 1944; Morton & Miller 1968). The 
present author has observed several spat falls in 
mangroves near .Adelaide, but they failed lo persist. 
It Ls uncommon for E. flinders/' to be found in 
mangO'ves .w^\ no E r ndwstms were found in 
mangroves despite its abundance on rocks in the 
Adelaide region. 




t would like lo thank Dr A. J. Butler for providing 
use of microscopes for examining specimens and for 
reading a draft of the manuscript. 


Andkrson, D. T. : Andkrson, J. T, & Egan, F..A, (1988) 
Bala no id barnacles of the genus Hexaminius 
(Archaeobalanidae: Elminiinae) from mangroves of New 
South Wales, including a description of a new species. Rc<; 
Aust. Mtts. 40, 205-223. 

Bavliss, D. E. (1982) Switching by Lepsiel/a vinosu 
(Gastropoda) in South Australian mangroves. Oecohgia 
(Bert) 54, 2(2-226. 

____ (1988) A new intertidal barnacle of the genus Elmintus 
(Cirripedia: Thoraeica) from South Australia. Truns. R. 
Soc. S. Aust, 112(2). 75-79. 

Buckeridge. J. S. (1982) The barnacle subfamily 
Elminiinae- Two new subgenera and a new Miocene speeies 
from Victoria. J. R. Soc. N.Z. 12(4). 353-357, 

(1984) A new species of Elmintus from Pomahaka 

River. Souihland. New Zealand. A'.Z. j. qeoi xeophxs. 27. 

Darwin, C. (1854) "A monograph on the subclass Cirripedia, 
with figures of all the species. The Balanidae, the 
Verrueidae. etc" (Ray Society, London). 

Egan. E. A. & Anderson, D. f. (1985) Larval development 
o( Elminius cowrtus Foster and Hexaminius popeiana 
Foster (Cirripedia: Archaeobalanidae: Elminiinae) reared 
in the laboralory. Aust, J. Mar Frestm: Res. 36, 383-404. 

Fi.owr.RDEW. M. W. (1984) Electrophoretic comparison of 
the anlipodean cirripede, Elminius modestxts. with 
immigrant European populations. J. mar. biol. Ass. U.K. 
64, 625-635, 

Foster, B. A. (1978) The marine fauna ot New Zealand: 

Barnacles (Cirripedia: Thoraeica). Mem. N.Z. Oeeanogr. 

Inst. 69. 1-160 
(1980) Biogcographic implications of re-examination 

of some common shore barnacles of Australia and New. 

Zealand. Proc hit. S\mp. Mar. Biogeographx & Evolution 

in the Southern Hemisphere. 613-623. N.Z. DSIR 

Information Ser. 137. 
(1982) Two new intertidal balanoid barnacles from 

eastern Australia. Proc. Linn. Soc. N.SM 106(1), 2L32. 
Hutchings. P. A. & RncrtBR, H. F. (1982) The fauna o( 

Australian mangroves Ihid 106(1). 83-121. 
JONES, D. S. (1990) The shallow-water barnacles (Cirripedia: 

Lepadomorpha, Balanomorpha) of southern Western 

Australia. In Wells F. E., Walker D. L, 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). 
MooKb, L. B. (1944) Some inlertidal sessile barnacles of 

New Zealand. Trans. R. Soc. N.Z. 73, 315-334. 
Morton, J. E. &. Miller, M. C. (1968) "The New Zealand 

Sea Shore" (Collins, London. Auckland). 
Pope, H. C. (1945) A simplified key to the sessile barnacles 

Ibund on the Ricks, boats, wharf piles and other installations 

in Port Jackson and adjacent waters. Rec. .4usr. Mtts. 21, 

Shepherd. S. A. (1983) Bcnthic communities of upper 

Spencer Gulf, South Australia. Trans. R. Soc S, Au\L 

107(2), 69-85. 
Thomas, 1. M. & Edmonds. S. J. (1979) Intertidal 

invertebrates, pp. 155-166, ///Tyler, M. J., Twidalc. C. R.. 

and Ling, J, K. (Eds) "Natural History of Kangaroo Island" 

(R(tyal Sociely of S. Aust.. Adelaide). 
Womersley. H. B. S. & Edmonds, S. J. (1958) A general 

account of the intertidal ecology of South Australian coasts 

Aust, J. Mar. Ereshw. Res. 9. 217-260. 


By Robert T. Lange*, Brendan G. LAYf & Rodger W. TrNANf 


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. 

5&MM<»ftM oj'tfir to.w/.Wm Of S $ftf. fl99-t). H8lTi. U5-I31 


by RonnRT I. Langc* Brendan c. L&Yt & Rodoer W. Tvn\n-- 


Lanoc R T , I ay. B, G. & ft nan- R. W. (1994) Evaluation $ extensive arid ran^taniN: the land condition 
index (LCI). Trywtt- /? Swe & Ms! 10*12).. 125 13 f 31 May, |yv4. 

Social and historical circumstances thai have given rise to timorous new ran^eland manager i cut legislation ifl 
Snulh Australia arc outlined- A program specified fy uV legislation is explained and Jescnhe.l Whittl will determine 
tho condition of the 40 million hectares of arid rangeland in South Australia rclmive tncnierw aUmt susiamanility 
and prevention and rectification of degradation An oh|eetive assessment ot averse ranecland condition, the Land 
Condition IndoM (J«C{) is introduced and results obtained by its use arc illustrated *hl\ appraised 

Ki-.y WfrJU*i> And /one, ranjHands. Australia, vegetation, assessment, technique 


The Pastoral Land Management and Conservation 
Act (1989i came into operation tn South Australia in 
March. 1990, alter years of public contention. It 
challenged the then Out buck Management Branch Of 
the South Australian Lands Department Uts the 
responsible ngency) with an immense, mandatory and 
assessment task within an 8-year deadline- In Its 
combination ol 'difficult features and its emphasis OB 
sustainabiltty, this (ask may have parallels in other arid 
parts of the world, where our procedures mi^fn. 
therefore, be applicable. 

The particular combination of challenging features 
is a vast rangclaud area (the sire ol fraq ot Somalia 
fbl evample) poorly documented and difficult of access, 
lo be assessed for the integrity flj its vegetation and 
soils in deiaif according to characteristics mil 
accessible by remote sensing, using minimal time, staff 
and money, 

Wc faced Ihc further difficulty (given the limited 
amount of resouires relative to the immense size ol 
the task) that our assessments even of small tracts ol 
land (a lew hundred ha or so) would have to be capable 
of withstanding challenge and close scrutiny in a court 
of law. 

This paper outlines (a) the historical and .social 
context ol the Act, (b) an approach (including a new 

index ol land condition) which we have devised to help 
achieve what the Act requires and (c) some initial 
msulls obtained by application ol the index, 

Nalional Key Centre for Teaching and Research in Dry land 
Agriculture and Land Use Systems The Univcisity of 
Adelaide Roscworthy Campus, Roseworthv. South Australia 

Current address: Bo\ 330. Whyulla. South Australia jt>0<). 
Pastoral Management Branch. Resource Conservation 
Di vision, Department of Environment and Natural 
Resources. CPU Box 1047. Adelaide. South Australia 5fl0l 

Australian Background 

Whether native inland Australian vegetation can 
ultimately withstand the impac' of ungulate flocks and 
herds is yet to be determined. Since it evolved in total 
isolation from ungulates, it may prose unavoidably 
susceptible to the effects of large numbers of them 
From prehistoric lime until die mid -Nth century, the 
inland arid regions of S.A. sustained only sparse 
Aboriginal nomadic hunter-gatherers. Due to laek of 
surface waters most vegetation probably experienced 
only very light grazing pressure, on average, by 
kangaroos alone. That legime changed "teata- 
elysmically" t Adamson & Fox 1982 1 when Europeans 
colonised much ol the country lapped ground waters 
and introduced millions of ungulate stock and various 
feral animals, creating a sedentary grazing industry 
which now occupies 40 m ha (Fig. I) and is mostly 
in the early part ol its second century of operation. 

The saying "Out of sight, out of rnmd" aptly 
summarises public and governmental attitudes to 
pastoral zone landcare throughout much of the 
industry's history iLange 1983). Nearly all of South 
Australia's I m people dwell in the arable coastal 
regions and of these the vast majority live in and about 
Adelaide. Only a lew hundred at any one time have 
had extensive pastoral zone experience. City-dwellers 
have only recently become better aware and more vocal 
about the and zone, as tourism, wildlife protection 
recreation and mining have increased. 

Sheep Stocking enterprises typical of the more 
southerly parts of the zone are pmccctcd from predatory 
dingoes by a special fence (the Dog Fence), featuies 
include the subdivision ^i .he rangeland into wire- 
fenced paddocks with water available for stock at fixed 
points, approximately fixed flock size year in, year out 
and a system of vehicular access tracks. This leads to 
the development o\ repealed dnnk-eenlred patterns of 
unequal llocktirne distribution (piosphere pallern, 
Langc 1969. 1985: Andrew 1988) in which flocklime 



Soil Conservation Districts in the pastoral zone 

Fig. I. The pastoral lease lands of South Australia, shawlrtg also rhcir 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 on a broader scale, with most 
pastoral properties there without internal fences 

Scientific and genenil literature, 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; Watte 1896; Ralclil'le 
1936; Pick 1944: Lay 1979). This was ot public concern 
particularly in the 1930s when the issue of soil erosion 
was topical worldwide (Jacks & Whyte 19V}) and has 
subsequently remained a problem. Haifa century Utter 
in the 1980s, government administration again publicly 
conceded that some pastoralists were still I ailing to 
meet landcare obligations and that land degradation 
from overstocking and feral animal pests was 
extensive 1 - In an attempt to accommodate all 
interested parlies, the Pastoral Land Management and 
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 landcarv 
legislation tin some ways setting scarcely-attainable 
ideals of landcare perfection) but it also provides 
provisions for grazing industry protection and appeal 
How these opposiies 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 must be carried out by 
the Pastoral Management Branch. Of these tasks, 
this paper refers to the requirement lor vegetation 
assessment. This immense task, which the Act requires 

'South Australian Government. Lands Department (ls»8l> 
The administration, management and lenurt- of South 
Austi alias P;iNti>rdl Lands. Interdepartmental Kepmi 
(Chairman: J. Viekcry. Lands Department). 

no in 


lij he earned QUI within iggltt years am! at 14-year 

mtel vuls tlicreulfer. relates 10 40 W ha otitic rangekuids 
rind is to specify the condition pi the land according 
to criteria derived from the Objects of the new Aei 

These cth tl"tv£lj n quire thai the a 

area he diteclly inspected by teams of trained assessors 
i»n the ground. As is usual where a larve arid 
eomplieaied lask must he accomplished piecemeal by 
JiUcrcnl teams rjfjet iJVCi sWlldapdi&ed procedure* 
need lo he adopted lo ensure thai the findings ol any 
assessment :ik- independent of the parttculai team 
employed The development ol ihc index and training 
plO&dUfGj described below i> an attempt ul -.ucn 

In Soulli Australia's non arable inland regions a 
pastoral lease is the only farm of tenure than can he 
planted over Crown (government! land that w |o 6? 
used 1'T long-term grazing purposes a pastoral lease 
cannot be granted or extended without an assessment 
ol die condition ot the land 1'irsi having been made 

(PLMC Act, 1989, See, 2a 23), This is *hj rhe 

assessment |ir,ig(ain is urgent and crucial. I "he pa&fttl 

fndwrfry r* aOTiiing ihc pouring ol new lease*. 

specify in*; new stacking limits and other conditions 
consequent upon these assessments. 

Section 6 of Ihe PLMC Act Males that such 
assessment of the condition ol the land mm b< 
thorough, miisi include an assessment C*f the cap tcit^ 
M tlie land to carry stock, must be conducted in 
accordance with recognised scientific principles and 
must he curried out by persons who are qualified and 
experienced in land assessment technique 

The tuilher meaiune ol assessment derives tlirc*crl> 
I mm relevant Objects ot the Ft MC Act (See. 4) which 
are: (a) to ensure that all pastoral land in the Sta.e is 
well-managed and utilised prudently so thai its 
amcwablc resources arc maintained and its yield 
sustained and (b) to provide for (1) the elTecove 
monitoring of the condition of pastoral land, (2i the 
prevention of degradation of the land and its indigent »u ■ 
plant and animal life and (3) the rehabilitation pf Ihe 
land in eases of damage. The Minister is expressly 
chained (PLMC Aei Siv S) with the duty ol acting 
coosistenilv with and in furtherance ot ihose Objects. 
Assessment pursuant to ihe Act thus is the pn>ce,s ot 
obtaining scientifically based answer* to the Obiccu 
posed as questions, viz.. h this pastoral tand well- 
managud and utilised prudently so that us renewable 
resources are being maintained and its yield sustained'* 
Is degradation of the land and ol its indigenous plant 
and animal life being prevented' 1 Is there degradation 
which requires rehabilitation*' Has effective mnnika-ing 
ufthc condition of the land been provided' What N 
theeap."-iv i .it the land to cam' s *ocJ < '■asistentlv wilh 
answers to ihe foregoing quest tons'-' 

Since 'degradation' o. Jet med t PLMC to Sec 3) 
as a decline in die quality bf the natural resources ol 
the land resulting from human activities, s-ornc ol Ihc 
implications atfffiftlg from die Objects are wnmediaiety 
apparent- For instance, ihe conventional rangcJands 
dug ma lhal the desirable condition of the vegelation 
is that which feeds most stock is potentially denied by 
Chis legislation. The optimal condition instead is that 
where indigenous ptant and animal life retain their 
prepastoral integrity while a? far as possible sustaining 
a viable pastoral industry. Appreciation of this 
difference is crucial 10 developing the condition index 
outlined below and |l will be furthei examined in I lie- 

That the answers to these assessment questions are 
meant to he taken seriously is made plain in many pan* 
ell the PLMC Act iiictudme Sec, 43 for example, a 
Section excluded from the appeal provisions (See 54 
>>). Section 43 makes il clear lhal the Pastoral Board 
shOUld require the removal ttf stuck from an area nol 
only if its believes thai the land has been damaged but 
also il the Board considers ihar rhe land is likely to 
suffer damage. 

Essential Preliminaries to Assessment 

Il Should be noted here that the assessment program 
as developed consists of two pans viz . ihc lease based 
Land Condition Index determination iihc subject ot this 
paper) and a network o) paddock-based pftoto- 
mon iron tig sites. These lattet sites, with vegetation and 
soils data collected, form ihe base line for the 
jssessment ol future trc-nds in CttCjl pa.ld.n_k 

Any thorough assessment & a station or lease in 
.accordance with recognised scientific principles tSec 
6 of the PLMC Act.i should employ many independent 
samples in an effort to achieve a picture which is as 
balanced and equitable as is practicable. In practice, 
sampling falls somewhere between systematic and 
random, as in the way the industry itself informs 
potential buyci> about the contents of a bale of wool, 
for example, where the sample is trom an arbitrary 
grab. In sampling station condition, reliance is usually 
placed upon the pastoralist's system of vehicle access 
tracks to spread samples throughout the overall area 
rhpjj to avoid possible observer bias, samples are 
dr.Avn hy Stfifl msiricfed-iundotu processes bom 
what is track accessible, there are some obvious 
Lonsequences i>1 such sampling but they are more 
academic than practical and there is no evidence that 
they significantly affect comparisons. All well 
developed pastoral land has adequate watering points 
and a network of access tracks linking them. Thus there- 
is a higher duui random likelihood of sampling, ClftftN 
to vvaltr, and a correspondingly reduced likelihood ai 
interpiufintjf ranges, than would result on random 




Km frorr water 
3 W 6 AS 

10 5 


S - 

1 = 3.6 km 





i 1 


5 = 3.6 km 


| 20 


-in - 

S£ corner, McDouall Peak Station 


Km from water 
S 4.5 s 7,5 




f J - 


l.i KjT 


10 — 

x = 4.2 Km 

SE cornor, Millers Creek Station 

on the LCI 
sampling route 

route relocated 
without reference 
to water 

Fig - Histograms showing the percentage of sanipling points at various distances from water points on the access tracks 
(above) and when the same track-grid is offset trom water-points (below). 

tracks. This effect is consistent from station to station 
(Fig. 2) and can always be taken tnto account. 

Recvgnirion of Vegentnon Jypi- 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 type> 
is represented and. of that type, which of the various 
sub-types tailed components. In practice this ability 
comes from instruction manuals and Held 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 a time, thus reducing the 
immediate diversity to be handled, The assessment 
teams then attack the task by completing assessments 
pi nne District before it-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 criteria to distinguish condition classes (see 
below) and we illustrate the resulting manuals with 
extensive colour photo-guides. All teams arc then 
trained en bloc tin extended field calibration exercises) 
to achieve a uniform, consistent interpretation of all 

All main diagnostic criteria refer to perennial plants, 
rather than to ephemerals which appear only after 
significant rainfall. The main reason for reference Co 
perennials is that they embody the drought fodder 
reserves of the rangetand as well as being the reliable 
buffer against wind erosion. There arc a tew places, 
particularly in northern cattle rangelands, that lack 
perennial vegetation sufficient for present purposes; 
these will be treated in separate ways. 

TaHu: !. Vi'^elaiunt tyfte and ivmputwnt uwtl in LCI 



Chenopod shrublands 

Woodlands with grassy 

Arriplcx ivsicurm Maifiww 
astrvtruha treeless plains 
Mucin utwura- A. ramulosiv 
grass sp. on deep sands. 

The level ar which types and components in general 
are distinguished is indicated by the examples giving 
the names of predominant species in Table 1 

C ond\ j ion Ckt.sses 

Arguments about non-linear **state and transition" 
models of rangelands dynamics notwithstanding 
(Westoby ei 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 l he original stock palatable perennial species 
still 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 

1 ess- palatable perennial species including weeds or by 
ephemerals and Class 1. 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 thai 
deterioration to Class I goes hand in hand with the 
onset and rapid acceleration of soil erosion (Ratcliffe 
1930; Jacks & Whytc 1939; Pick 1944; Lay 1979). 

The advantage of distinguishing only three classes 
is that only two sets of separation-criteria are then 
required i.e. separating Class 3 ftom Class 2 and Class 

2 from Class I Obviously, it is easier to train all 

Tin-: k.i 


assessment teams in absolute: consistency when they have 
to apply only two sets of separation criteria to describe 
the condition of some given vcgclational component 
instead ot three, lour or more sets. Contrary to what 
at first sight might seem to he the case^ this restriction 
to just three condition classes in no way limits the 
genera! sensitivity of inter-lease comparisons. When the 
condition of two leases is to be compared, the sensitivity 
of the index Of comparison depends instead on a 
sufficient number ol samples plus the use Of weighted 
averages, as will be demonstrated. So, in specifying 
arbitrary but consistent criteria for the separation of 
condition classes, counts can he avoided in favour of 
absolute differences. Regarding the /I vcsicorhi-M 
a\m>tncha component of the chenopod shruhlands. for 
example, the main criterion for separating condition 
Class } from condition Class 2 is the elimination o\A. 
vcsiiuriu (a grazing-susceptible species of the original 
palatable perennial component*. Condition Class I is 
distinguished from Class 2 by loss of /If tisrrofriihn, 
a palatable but more graztng-rcsistant perennial, lakaig 
die example of the Acacia attcum-A. /wwf/tei/-gmss sp 
component of the woodlands with grassy uTidersk'iev, 
condition Class 2 involves loss of all ,1 an*'unt 
regeneration, appearance of a distinct browse line on 
old trees, elimination of the most palatable perennial 
$ r,is ses ( $ j 1 , MdMi iiaiher para Jo U2 ) and se ve t e 
pruning, at least, of palatable shrubs (e.g. Eremophiia 
hiirnbei). Similar sets of criteria have been adopled |o 
distinguish condition classes of all vcgclational 
components of all types in the rangelands, supported 
by photographic examples of these classes, 


An essential for condition assessment conducted 
piecemeal by diiferent teams, using tins method is a 
lull scale combined Oeld-tiaining exercise. The Soil 
Conservation District to be assessed it traversed lo 
intersect all vegetation types and components. At >top 
after stop on this traverse, samples are assessed bv the 
party as a whole, according to the manual, until all 
questions have been resolved on the spot, guaranteeing 
uniformity of interpretation. Only then ;ire the different 
teams allocated their particular shares of the district 
workload and Ihe actual assessments conducted 

Prcrictcrminaiitm o/ Sampling Paints 

Iking, available maps of station tracks, the 
8$SC5smCnl sampling route is pre-selectcd for best 
systematic coverage and the total length involved is 
calculated using an opisomeler That length is then 
divided into 100 equal parts within each ci( which a 
precise stopping-point ts drawn at random. The results 
tire tHwi programmed into a computerised trip-meter 
(Halda Instrument Co) mounted in the access vehicle, 
which then signals the stops. This guarantees restricted- 
random sampling of the route. 

Assessment and LCI Calculation 

Field teams comprising a scientific office! and a 
technical officer operate self-contained 4- wheel drive 
expeditionary vehicles equipped 1or remote area 
operations, including radio communications. 
Navigation is by reference to available maps and 
satellite imagery using a vehicle-mounted OPS unit. 
At each sample site (of which the trip meter sounds 
prior warning and displays a count down to 10 no. the 
reference is made to the 50 m-square plot located 
immediately beyond a line parallel to the direction ot 
vehicular travel. 10 m offset from the left side of the 
vehicle. The sample within this plot is raied according 
to vegetation type. comp< nenl and condition, is 
photographed and lite data are teeorded. At the 
conclusion of sampling, the station land condition 
(LCD is calculated as in the hypothetical example in 
Table 2, 

lABLt :, Sample oj cuicuhtitm Of LCI istawm Uitui 

Condition Percentage Multiplier 
Class of samples 









Note that this index has limits of 1.00 (the whole 
station in "degraded" condition) and 3.00 (the whole 
station in "imdeeradecT condition). Actual stations have 
values between these limits. Note also that the index, 
once calculated, is not to be interpreted mechanistically 
but instead is used for first-order sorting of stations 
according to condition. Other evidence and data 
collected independently about each station arc then 
taken into account. Stations can be compared directly 
by the t.Cf only where they involve similar paslurc 
types and components in similai proportions. 

The Nature of Initial Results 

Assessment learns obtained first results during 1991 
from the Kingoonya Soil Conservation District of South 
Australia using 19 stations. Fig. 3 makes a comparison 
between the LCI scores ot lands making up the stations 
in this district and the- ma* 'mum stocking permitted 
on ihem by- the provisions of existing lease documents. 
As can be seen, no station attains the ideal ot the Act 
(all samples in Class V). Instead, stations range very 
widely in condition, from as low as 160 up to 270, with 
an average of aK>ut J00. This low average score would 
imply that maximum stocking allowances will have to 
be reduced, on average, to enable some progress with 



the rehabilitative process prescribed by the PLMC Act. 
As can also be seen, there is no consistent relationship 
between the present condition of stations and 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 it can be seen that the 
LCI provides a comparative assessment ol stations on 
standardised criteria derived specifically from the 
Objects of the Pastoral Land Management and 
Conservation Act. synthesising an overall picture from 
data provided by means of teams of which none has 
seen more than a tew of the total stations. It thus fulfils 
its design purpose and can deliver defensible 
comparative assessments to the Pastoral Board, which 
has the task of finalising judgments affecting stocking. 

hig. 3 shows the crucial dillerence between 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 tlats with inedible samphires 

(Chenopodiaceae tribe Salicorneae) vegetation might 
score a perfect 3.00 on the LCI scale, for completely 
satisfying the landcare Objects of the Act, while being 
intrinsically useless for pastoral production whereas 
overgrazed and denuded lands (e.g. chenopod 
shrublands reduced to ephemeral grasslands) of very 
low LCI SCQre might cam the pastoralist substantial 
profits as a result of high animal production. The aim 
of the Act where LCI scores are low is to arrest decline 
and then reverse it into a long-term trend of ascending 
LCI scores as a first priority. Any stocking of the lands 
has to be commensurate with that and be a secondary 

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 as can be achieved, the data 
yield estimates of the proportions of the district that 
arc 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 vegctational component can be specified, 


Land Condition Index (LCI) 

160 200 


6 12 

Stock maximum sheep/km 2 

I u\ i I he prevailing itiiMiifiiching between Mocking allowances amongst leases of the Kinsnnnyri Soil District. South Australia. 

IHr IX ! 


revealing ihe relative degree to which I he ditierem IJ pc- 
have been degraded. This js important management 
nii« 'filiation It is upon Ihe basis- of data such as these 
that the Pastoral Land Management and Conservation 
Aei must operate and without which its admimstrj ton 
cannot address itfi legally-stipulated imperatives. 


Wei I -reasoned technical prescription-, lor l. ( nd L j.rc 
hi ihe Slate's pastoral zone have been available loi 
almost a century. Waite (18Vo) aimed fox a balanve 
hetween offtake and -attainability via a well-reasoned 
ecological argument involving land atid flock 
subdivision, wateipoini multiplication* nutrition of 
lactating ewes, the distances sheep walk, drought 
strategy, deferred crazing, spelling and economies. 
Wattes key principles have been thinly linked v»ith. 
good landcarc by Lay (i979) on the basts oriori£-lerm 
studies. The most recent managerial preset ip-ou 
(Stafford -Smith & Mortmi WOi is Mule different in 
principle from Ihe first (Waitc 1S96) sU tl is nol 
defioicncy ol management advice that explains a 
century of ongoing degradation. 

f torn initial experience witL* ihe |Cl tt now seems 
probable thai most stations m South Australia will fall 
• h..rt of Uh_ land ^'-ndioon ideal of the new Act. Mm, 
means that reclamalory. i>Jmbj|itator> action will 
predictably be required which involves 
recommends ions for reduced stocking levels. We 
believe that those stations managed in closest accord 
with the Act (highest 1 CI) should retain for the present 
Ihttr existing slocking allowances and act as 
betichmatks foe other stations with similar vegetal ton 
types but lower indices. These other stations should 
be given lowet slocking limits while rehabilitation 
programs are initiated. One reason for not 
recommending slocking allowances higher than ai 
present in any instance (even tin stations with highest 
LCI) is clue lo a feature of the index itselF, In order 
lo guard against likely accusations that the assessment 
scoring procedures are W*\ severe, they were structured 
in favour of high LCI scores rather than otherwise Pi -r 
example, downgrading a saltbusb (Atrip/ex \v\uxh'i(*\ 
shruhland from condition Class 3 to 2 is forcM; 
while just a scattering o\ salthush remain, even ibi igji 
the ungra/ed density was many thousand/ha ttilB 
where even the best-preserved vegetation in an area 
5C0TfcS less than the ideal, there is no full compliance 
with the suNlainability notion central to (be Act 

In its considerations of the Mrs! leases assc-scO 
by this method in the Kingoonya Soil Conservation 
District, the Pastoral Board has been reluctant to 
.id just leases with low LCI scores unless these scorn 
can be dfofc th ItHbUtftbk '" CUItenl management 

k.itl«-r it has indicated it would await an indication 
ol trend Mom permanent photo-monitoring sites set up 
n the time ihe LCI is determined 

Of the many further points that might be made, the 
one warranting most emphasis here in that the LCI and 
associated procedures have to be taken against the 
backdrop of the stringencies, pressures and urgencies 
[hat attend it. Those who mighi regard is as hasty 
(compared with what has been attempted in some other 
Australian States) should recall the timeframe The 
whole ran^e lands (40 m ha) must he by law assessed 
in eight years; this is a formidable and costly task. 

Lhirnatelv. however landcare in Ihe zone will 
depend not on the assessments but on Ihe determination 
will- winch executive government uses the assessment 
infmriiation lo ensure that more sustainable land 
management practices are adopted in our .arid 

Kcfm nee* 

A.UAMSON, D & l(i\, M. "1982) Change in Australian 

v. ^j 1 4nce European .sciiiemciu pp l(W i4< ( In Sniith, 

1. fFd | "A Historv '•! AlUtlnifuynn SellltTncnr t MeCJraw 
Hill. Sydney) 
AM'HIW, M. H <WHH) £jffl2l«g i-iipnt' in rdjin... tt) 
h.L.lock watering points, Vratttl* m bctfi tttw Eyol, 3, 

Ulv.N, S lISV?. TM dlpfU Of stMlenienl p\t) 
nrctlpatioit m Australia urnm the imli*cnou> vegetation 
TrttHtti H. ftw s. ha/ LL W5-20& 

hos, Li V ft WHVff. R 0- O'MO) The Rape ol the 

1 jfth A World Survey III Soil Kr-.-M-ai" (Faher& Faber, 

London ) 
LANfiU, K t |W69> Tiiu pjosphciv ihe&p truck- and dung 

patterns. L Raftx? Mtt«fi£ 72 t 596-400 
(1983) Arid zone lunuYar* Education and Polities. 

pp. 112-129 Jft 'What fillMrr li.r Australia's Ao<l | arnl - 1 ' 

(Ausi. Cnnncrv. bound., Vie Aust.). 
I19K.M <listnbuiiofri of :aockiiit,' intciisil) 1 

pnntuced by sheep flocks grazing Australian chenorxxi 

tOtrublantlB. .1hm$ ft. Sm S Aim 10*, Ifi? 174. 
i A H G (Pl^) Shrub population dynamics undo gruging 
.i tplttj tenti rtuth pp. I07-I2y //* (jiaer/. R. i Howe.v 

K. t Rtjsi '*Sludie>i;J the Ausiralian Arid iftOHC IV Cheno|>)d 

Snrubl.inds" iTSIRC), Aust.). 
i'i. k, J. H (1944) 'Australia 1 * Dying Heart' (Melbourne 

Cnivcrsitv Press. Oxford University Prtss).. 
Rvit | hi . , ft N. (V)Mi) Soil Drift in "lie Arid PaUorot Artas 

111 Suulh Au*.Ii:jUj f'StK. (Aiixt.) Pamphk-L No. 64. 
South Aostkalian Oovt-.RNMtNr 0989) Paifiantem, PaMoral 

l.untt Mjiia^cnienl jud Const rvaiion Att (Govt. Printer, 

SiAi-hOHO Smum, U VI, & Morion, S. R. tlW) A 

framework for the BCptojtt Ol arid Australia. / of H/vJ. 

hn~ 18, 255 Z7& 
W \ni« ( P tlS*)6) Pl^lural ikvLlopaient - past ami prcv.-tit 

' Ofv..:rvr,-- Sept 26 IS»fr (S. AtlSt I 

W>s.oHv. M.. Walm:r. B. A n.a Mmk. i, tW)) 
Opportunistic management tor r&n^eUnds not at 




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, 

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 3 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. 

Impactions of the Royal Society of S. (1994), 118(2). 133-138. 


by G. J. Prjdeaux* 


pRinr.AUX, G- J, (1994) A small sthenunne kangaroo from a Pleistocene cave deposit, Nullarbor Plain, Western 
Australia. Trans. R. Soc. S. Aust. 118(2), 133-138. 31 May, 1994. 

Simosthenurus maddocki mdiorhorensis ssp. ntfv, 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 smaJler. The P is small, 
narrow and inflated posteriorly. Geographic d watting due to resource limitations is a possible explanation for 
the small sr/e of this Nullarbor subspecies. 

Ke\ Words sthenurine kangaroo, Simosthenurus maddocki mdkirhorensis ssp. nov., Simosthenurus maddocki 
maddocki, Lindsay Hall Cave. Nullarbor Plain, Pleistocene, geographic dwarfing. 


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 

* Sehix>l of Biological Sciences, Flinders University of South 
Australia, GPC) Box 2100, Adelaide. S. Aust. 5001 

Plain (Fig. I). 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. 







Fig. I. Deposits yielding Simosthenurus maddocki in southeastern Australia. 



Material and Methods 

The material is housed in the Western Australian 
Museum, Perth (WAM). Mensuration, dental 
terminology and nomenclature follow Ted ford (1966). 
As the homology of premolar cusps is currently being 
re-examined by Prof. David Ride (pers. eomm.), 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. 


Order: DIPROTODONTIA Owen, 1866 

Suborder: PHALANGERIDA Aphn & Archer. 1987 

Superfamily: MACROPODOIDEA Gray. 1821 

Family: MACROPOD1DAE Gray. 1821 

Subfamily: STHENURINAE (Glauerl, 1926) 

Genus: SIMOSTHENURUS Ted ford, 1966 

Simosthenurus maddocki maddocki 

Wells & Murray, 1979 


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 
(Flanncry & 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 (31°35'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 3 very small, 
narrow to tapered anteriorly, but inflated posteriorly. 

Fig. 2. Stereopair of Simosthenurus maddocki nullarborensis ssp. nov. lefl maxilla (WAM 92.9.8. holotype) in occlusal 



Vlg. 3 SleiViipaiiS nf Sutio\lttitU4ru\ <r«uhlink> nufluht*" >im,\ ssp i)i.v_ left P\ tup, uiid P\ Nitloni |WAM 92 *JR. Intlu(>pe) 
m occlusal view. 

Dt'Mription of holotypc: Maxilla (Fig, 2). 
Fragmentary nature ol specimen has icsullcd in 
preservation of very few non-denial characters suitable 
lor description or comparison. Palatine vacuities 
appear to have extended anteriorly to level of dP 1 
anterior loph. < nils base of masseteric process 
preserved in hololype making an estimation of sifce 
difficult. Posterior aspect pf huceiuatoi muscle scat 
is laterally wide 

Dentition (Figs 2-5, Table I) includes p\ dP\ 
M 1 \ excavated P\ with Leeth exhibiting only 
beginning wear. 

P 7 : Verv small with relatively high labial crest 
containing prominent anterior labial cusp and small 
cusptilcs posteriorly Transverse ridge joining labial and 
lingual posterior cusps encloses posterior basin, Fine 
transverse ndgelets occupy longitudinal basin, with 
anterionnosi slightly larger and dividing <>ll small 
anterior hasim 

iip j ; Completely molariform. widei across 
posterior loph than anterior loph. and smaller than 
succeeding molars, 

P*. Posterior third of [*■ intlated both lahially and 
lingually. Posierolabial stylar cusp is well developed 
but does not attain height of posterior labial cusp 
Labial crest divided into three cuspules bordered al 
both extremes by prominent anterior and posterior 
cusps. Transverse ridge leading halfway into 
longitudinal basin from labial crest slightly overlaps 
with small ridge descending across Jrom lingual L'fesfl 


Fig. 4, drawing nf ihe holotypc- P^ ot SlmtiSthetlUtUS 

fnaddocki ntilUirbor?n\t\ v.p mtv A ;iiiLenitr huMn: P. 
|K>stcnur basin: L. longitudinal hasm. |.;«. labial crest; Li. 
lingual crest: PLSC, stvlar cusp, (length = 


Ci l PRIDHW a 

[«i fortn anterior border ttl poslenoi ba&m, Small 
anierior basin separated by transverse | [djgc dost ending 
hugiuilly from prominent anterolahial cusp to lingual 
inunterpart, Ridge appear Ip have formed fftoftl 
uiiifnaiion rif two smaller ridges descending 
transversely from each anterior cusp. 

M Mi Molars increase in length and width 1mm M ( 
to M 1 (M 4 not preserved) Respective widths across 
pmtnloph and tnetaloph in M 1 arc identical. In M-\ 
proroloph wider than metaloph. and in M A wider 
again. Molars low crowned with lophs notably convex. 
anteriorly. Very line erenulations on molars apfl 
extensive, slightly coarser on posterior side >-i 
mcialoph. Anterior cingulum well developed, hut not 
especially broad. It extends anteriorly from up of 
p-iraeone, then labially across almost entire width "t 
ptomloph. Low, weak poslprotocrisia extends 
posterolubially from promcone and meets with very 
61JK0] crest originating from metaloph. forming a crista 
obliqua. kirigUftl 10 this structure, a deep fossette is 
present in median valley. Labially, a notch is loaned 
between well developed postpara-and premeracristat 
Pi rSlCI [Or eingul urn broad and formed by fine 
poslhypocnsla descending posictolabrally, ihen labially 
across back of molar, and overlapping wtth less 
pioiimietil postmetacrisi.i 

Ftiwtyp'" vvam ^. i. : 7 a Mii^ui.a. unasdt tarted P' 

i Lindsay Hall Cave Collected in April 1992 b> 
Lindsay Hatcher, Western Australian Plane Caving 

tttffotitW- It is only possible to get an impression 
of variation withm £ m. rtt<llurf><»<tLsis by comparing 
the P 3 of WAM 92,9.8 and WAM 92.12.7. They differ 
sightly m si/e. namely length and posterior width, but 
more noticeably in morphology. Whereas the posterior 
third ot the P' in WAM 92.9.X is inflated both labially 
and lingually, Iherc in only a lingual inflation in WAM 
92.12.7 (fig. 4i. This ls due largely to Ihe reduced 
development of the postcmlabial stylarcusp in WAM 
92.12.7 compared to tin- huUypc- This is an especially 
variable character in other slhenunncs, including S 
tn. mw&dockl and S. tndilcnhilis* and probably also 
represents iiUrasubspccilie variation in S IW. 
null-irhni, usis Other notable differences between the 
premolars are Ihe lateral tapering present in WAM 
92.9,8. where Ihe tooth becomes more nanow 
anteriorly than WAM 92.12 7. and the incipient 
development of the anterior cingulum in WAM 92.12.7 
compared to WAM 92.9.8. Iritrasoccilic variation in 
premolar morphology is common in all slhenurine 
species far which numerous individuals are known 
(pers observation). 

f,\>tnpari,\t>n with oihct taut: Clearly, the skull fA 
S tn tiuf/arhi/rrn.m was very small even considering 

he. S Stt:reop.m a! Smcwffwf/Wf^ mffixft nufidrfyi nfal* -vsp nov, P , top i WAM 92 12.7. puratyrx). and ivpical 

SimnMhrnuru; nuuhtmh mdtt.iin'ki P* hoflOtU (SAM P27752), .n .x.ctusdl view. 



ihe difficulty involved with interpreting skull si/e from 
fragmentary juvenile specimens. Ihe preserved 
maxillary region is smaller in every respect than 
similarly-aged individuals of 5. #////, previously 
recognised as the most diminutive extinct sthenurine, 
II is much smaller than similarly-aged S. m. nuiddocki. 
Anterior extension of Ihe palatine vacuities and ihe 
morphology of the buccinator muscle scar in 5. m, 
nullarborensis are both similar to S. tn. maddocki 
The P : and P* of S m. nullarborensis are 
considerably smaller than S. m. maddocki (Table I). 
Although only two P 1 specimens were available to 
conduct one-tailed t -tests, both length (t=3.3, P=0.09) 
and anterior width (t=30, P=O.OI) for S. m. 
nullarborensis fell significantly outside of the known 
range of S. m. maddocki. As with the maxilla, they 
are smaller in size than those of any sthenurine. Based 
on the comparison of mean length and width 

measurements, they are 24% smaller than S. tn. 

S> m. nullarborensis and 5. m. nuiddocki differ in 
several morphological features of the P\ The 
poslerolabial stylar cusp is less prominent in S. m. 
nullarborensis . especially in WAM 92.12.7, Neither 
specimen possesses a stylar cusp which attains the 
height of the posterior labial cusp, contrasting S. m 
maddotki in which the height of the stylar cusp 
consistently exceeds that of the posterior labial cusp, 
fn S. m. maddocki the labial crest is divided into either 
two or three cuspules between the larger anterior and 
posterior labial cusps. Both S. m. nullarborensis 
specimens possess three cuspules but a similar 
variation in number could also occur in this subspecies. 
In the holotype P 5 the anterior basin is well formed 
and separated Irom the longitudinal basin by a 
transverse ridge (Fig. 4). This structure is not present 

T\bLt 1. Check teeth dimensions of Smiosthcnurus maddocki nullarborensis ss/t now, S. m, maddocki and S, p\U 

(mean iMtimhinJ iU'viuannt lohu-'nti range if 











ttt. nulla d)orcrt\L\ 








m. maddocki 

9.2 (fci5) 


6.2 rt>.S3J 

7.3 i0.5i) 

16,7-8.1 1 

6.3 uu'O 


6.2 0'.5vi 
1 5.7-0 y| 




10.0 duo 

\9h Ht5| 

7 4.<On 

|6'7 5| 


1 8-6-9 l| 

6.5 (0.23) 
\t&A o'»| 

6.7 (0i36l 


dP A 


m. nullarborensis 








ni- maddocki 




|« * M,l|| 

|8.(I 4.0| 

SO 1031 








9.2 \Q.7b) 

!8 9 •»,(«! 

9,6 (0.-V) 


4.9 (UMJ 

'4 3-5,31 

5 2 i(J031 





m. nullarborensis 


£8 .0 14) 




6.5 turi' 

(0.0 7.01 

5.9 {0-21. 



m. maddocki 

I6.U 10391 
|I5 3I7,0| 

8.1 (050 

10.1 (0 5ft) 
1« 3-0.1] 

8.3 taxo 






16.1 (O.OJt 


9.2 (0 54i 


11.4 at72\ 


9.8 BMtfll 

|H.O HUt| 

9.8 lOKN' 




15.5 (0,53) 

7.7 kuk) 


9.5 MUK) 

9.4 (U4f») 


9.4 iO-ok) 

|H1-I0 5| 


M 1 


m. nullarborensis 








W. maddocki 

10.8 WW 


10.1 (0.24J 

10.0 |0 Mi 
|M,4 t(l,g| 

5 3nkP\ 

1 4.8 5.7| 

5-2 (0 in) 

14 A (>.!') 




10.3 (0.51) 

10.2 (ii..ui 

|Mh t0,7| 

10.0 (0 (3| 

1^.4 I0.5| 

5,3 tftMj 







m, nullorboirnsis 








m. mcuUloi'ki 

11.2 to:i) 

|I0.9 ll.5| 

[itta n. d| 

10,3 Hn-ii 

|4.0 |II7| 

5.5 hum 


5.5 mmt, 






10.5 (0 44. 


10,2 ( 36) 

LW io x| 

6.1 fU.39) 


6.6.0 33. 
H.2 7JH 




m. miliar horensis 








m. maddocki 

11.4 m.yh 


J I.I (0.45) 

10. 1 (0,48) 
1 9.5-107 1 

5.7 uu7> 

150-6 21 

5.5 (0 3K) 





110.8 I2.2| 

10.7 (a-*. 

IllVtl || ( (.| 

103 (0.50) 


6-5 I06H. 



a j prii?! 

in V m maddocU when* inward curving "I [fi£ antoriof 
eMjeincs of the Jubiul and lingual crests occurs (Fig 
5). These Oft<Jp do not meet and result lp ftjl unrci i< «r 
basin which is very poorly designated •■( completely 
i rscni A mare intermediate condition is obwrved m 
W'AVI M2.12.7. 

Complete molansnlion of the dl J ' fa .. ti.n.i..te. 
of all sthenurincs I ikewise, the incrca%c in tnttlui 
length and relative changes in vvidllis aloiijj the tOOth 
row m S. tti n<titwhtnchs)\ are typical of most sp» \\ 
Molar morphology is very similar to ,V Wl mttikltH kl 
making them readily separable from the siTttibflg-firattl 
S gifli. which possesses a larger ensta oblaaua and 
less extensive line enamel cienulations, Therelhre. 
-ippc r molars llf die subspecies are separable un JUZC 
■ !- with measurements "I tbC s ' " J nnlhirh,>nvt\i\ 
molars approMTtiaiely I37J smaller than mean values 
lor S. m maitJacU. 


Althitu^h some doubt exiskd nimally »>vei which 
taxoikumc rank should apply |o the Niillarhor form, 
morphological similarity lo nomtnoiypic S fnaddo* ftj 
precluded a specific separation. Designation nf ImsmI 
subspecies has previously been made in relcicrne Ul 
dwarfing macropod lineages (Marshall & I ■•■ " I 
WTO; Dawson & Flimnery IMS) I'liev are- An effetfiyti 
torm of recognising cleat ent temporal Wl geographical 
\; ( 'iants within a species- Size redan .>n Ml F imd 
molar dimensions (.24% and 1396 respectively i < I 
^ r// maddncki to S HI mtllarborcnsis is computable 
W that observed by Marshall & Comiceini for other 
iiiudiUJiito large si/ol macropods from the Pleistocene 
lo I loloeene. they concluded thai resource limit- i | 
pritb.iblv accounted for dwarfing in these muciopods 
.aid Surct)philus htirrisii. Dwarfing ts commonly 
. \\\\ in island populations couicmpoianeous with 
large niatuiuul populations on the mainland (1 omoloiu 

1985) \ umhip effect between mainland regions 
caused by climatic unpredictability, winch today 
typifies tlie Ntillarbor Plain, may also have been 
significant in the Pleistocene environment, perhaps 
Icadmc- tO si/e reduction m $ m. titill<tthon'»xi\ 

lnltrestingly> analysis of si?.e variation \f\ fhylmituc- 
cynoccfihaius from Ac Nidlarbor Plain tl owry 1972) 
did not statistically separate las a subspecies) the 
population sample from larger T <_\nocc(>hiitu\. even 
though several small individuals were present 
Unlorunately, the laik of 7 S. m. rudlnrbtnvn.\i.\ malenaJ 
at this stage prevents a more thorough statistical 
hiJv-.i- [hnn thai COWlUCWd for Ule permanent 
prenic-lar However, dimensions of most measurements 
lall well oiiistde the known range of ,V. ni madiocki 
support mg us current designation as a Separate 
snn,p^ ■ 

A piobablv late Pleistocene age is attributed to the 
Lnid'.av Hall C«VE deposit based on the similarity of 
J W fntlltirhi>rcn\is to & tn, tnatidodit, and (he 
u'-(,cvunenec offtftOtbOT widespread, but undescribed 
sthernuaie I tic bone-bearing strata of Lindsay Hall 
C ave may equate wilh unit 2 in Madura Cave 
il.ond.Jiu, 1965; Lundclms & Turnbull IU89) which 
..., ■-. ( ;m. yielded 'his undescribed species Support lor 
Hits would icly on a detailed straligiapluc .nveMigation 
of cave deposits in the atv.i 


I 'h.mks tO Pr Rod Wills (blinders University), Dr 
John Lt>ng (WA. Museum) and an anonymous 
reviewer lor ollci mu comments Oft the manuscript. Dr 
Long also provided Fig 4. and with Dr Alex Baynes, 
was responsible tot hum ol the tosstl niatenaJ. I am 
giateful to the Western Australian Plane Caving Group 
lor their efforts, parlieulatly Wendy Bmks, Kalhcnne 
( *t isp and Lindsay Hatchci who discovered the deposit 
and collected t"o,sil material from the rate. 

KviV re nets 

D/.'A'MtN. I. ii PlANHERV. V \r jWHS) IWtllOtntl Hid 
phylogcnctic status nf living a«u foniil feaqgati^s iittd 
w;ill;ibi(-'; ol the gMUS Mocrttplti Shaw iMacrupo<licl:ic 
Marsupiiila) with (I mc*a >ub\ietieri»: rume for the I n 
walliihic-s, Ansl >■ tort 53. 473-498 

la MVNCttY 1 i & \i()v\ ,1 H, (I5M33) OccorttMWt* ol "a- 

exiinct inaeiojitMliiJ muRUplBl SlUUMtfymUM vrA,/ in 

1 ,--.v Smith Walex Att*t MaulWtil <». ' /- * 1 '. 
LctMiuasn, fift, V il'^K?) Body si/lm>I itmiiiuittK on tsluii.t- 
Ihc Inland Ruil i i.-MMiiincd \mcr Niti 125. MO 3MJ 

l.owtA- J- W. J. "i'>72i rhtt taxpnonili statist I iiuall R 

liiviii. mii's (Vt.jf,iipi,iia. Thylacinidae) frtsm Wesicm 

, -i\n I (f *•>, IV Aust $9, IV- >H 
I | Ml i . I • ( h, ||9W) Verlebralc rrtit.uiir, 'mhi. the 

Nullarhor < aws, Western Australia, fbht 46 J$ WJ 
feTntwnwj W n IWS9J.Tht mamrrailiiin 

of Mudma Cavfr. Wrsii-.r. Anio.iN.. I'o; VII: 

Mut-mrHiUt.lj.-. raScnuriuae. MHCfVpt I H .vid.-iu h-n 

111 Ihe nur\upij| rn^Uon ol 0K IttUWfl ftcLtuinn. GeOh 

|7im). I /I. 
Mr^Hii 1 ti ^CimHicciM, R. S. U l >78) Variatnlity. 

..voliiiKmurv ty'es. atld iillomelry in dwarrttlg KllC^gCS 

hiU.'hioiovs 4. tlll-ltv. 
I'iiim.i, N S (WHO) Mj< luptnlid skeletons, including: 

'>irth".ihi :tum\ iLilfonl, frOOl jii UttUSLial 'JmsMicU tSQtcT 

iteptJMl ii* ihc -.ouiheaHt ol "South Atranilij. AYr. .S. AuM. 

ft/(W, 18 131-141. 
1 ntfo. R. H ilUfioi A review >-t tlie macropodid genuj 

memmti thus Cuhj Fabl C»n>t, StL 57. 1-72. 
fl -. R T A. Mcuhan. K I. MUA'i A new sltienurinc 

kangaroo (MarMiplala, MacrnpnUidsc) (rum southeastern 

Sauti 4oictratia. Trnnx, R. Sac S, Auxl 105, 213-21*). 
MrtWAierv, k. & vvutiAMs r> l. g. (1984) Tlte 

luvat vciiL*hr;jicdeposiLMif VieUtHa Kossil Ca\c Naract^me: 
an iNlntUuctifi! It. ince'C-i'ln^y ;tml I;iimi;( /1m/ /%•/ 21 


ByM. Zbik* 


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 (Fa 195±03 n = 30), orthopyroxene (Fs 175±12 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. 

Ihrnsitciuins <>\ il,< MnvW W/cO .//\ Alin. (1994). 1180) B9 M r 


by M. Zbik* 

/niK, m (.1994) I he Cook 007 meteorite: a rrew Hi ehondrite fconi South Australia, fttuuf, A' Jri? S \hm 
118(2). 139-142, 3j May. >V*)4 

The- CwuJl »HT7 meteorite, a single stoiu: ol over KK) kg in weight, wad fottfid a' COOM "i I9HH II has been classified 
as .in R4 ehondrite of shock Pacies S5-6 and wnlal/te olivine u-.. (l) , rj)H n 30), otthopyroxcnciK ,.i . n - Ify 
clinopyroxcne maskclynitc. nickel-iron and troilile Mineral compositions and textures oidicaic thai Cook 007 
was a metamorphosed pun of the H-meleorile parent body and was very strongly shocked before reaching the Earth, 

Km Won ox: Cuok, meteorite, chondrilc. 


A single mass of the Cook 007 meteorite weighing 
over 100 kg, was found at Cook on the South Australian 
part ot the Nullarbor Plain (Fig. 1), in 1989. The 
approximate co-ordinates ol lite locality are 30°37'S 
I30°25TE. The meteorite was collected illegally and 
exported to the United Stales ot America. The South 
Australian Museum obtained a piece from an American 
meteorite dealer, Mr Allan Lang in April of GW1. 

In recent years the Nullarbor Plain has proved to be 
a productive area for the recovery of meteorites (K< '.an 

1992; Bevan & Binns 1989a. I989b» and m the last icw 
years has altracled the attention of illegal meteorite 
collectors. Under legislation enacted by I he 
Government ol' South Australia, all meteorites found 
in the Stale ate the property of the South Australian 
Museum An unfortunate consequence ot the illegal 
trade in meteorites is the loss of important information 
«m the exact date and location of the find. In accordance 
with the guidelines on the nomenclature ot meteorites 
from the South Australian Nullarbor (Bevan & Pring 
1993), the meteorite has been named Cook 007, being 
the seventh meteorite to be recorded from the Cook 

1 is 




* Fuel 

Kg. I. Map 0v South Australia showing the approximate 
location of the Cook 007 meteorite. 

* Polish Academy 
Warsaw. Poland 

of Sciences, Space Reseaich Centre. 

Physical description 

The piece of meteorite obtained from the United 
Slates weighs 26.2 kg. It is irregular in shape and is 
about 35 em in length, 50 cm in width, 4-11 em thick 
and polished on one side. I he specimen is currently 
on display in the meteorite exhibition in the South 
Australian Museum. The piece has a dark brown 1 mm 
thick weathering crust covering (he outer surface. The 
interior of the stone shows no sign of weathering, but 
the heavy weathered crusts covering the surface 
indicate that the meteorite had been exposed to the 
elements for many years. A number of quartz grains, 
probably 0l BCOliajl 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 arc partly 
reeryslalltsed 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 lo 1 cm in diameter. Metal and 
troilite occur as finely disseminated grains throughout 
the matrix. A slice was cut and a polished thin section 
was prepared and used for petrographtc examination 
and electron microprobc analyse-.. 



Fig. 2. rhotnmicrograph ol'the (thin) section ofihe Cook 007 
meteorite (length t tf the section *« cm) showing large dark 
(staining) area and granular structure lull of chondrules 
and chondrule fragments. 


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 fxm. Mineral analyses 
are presented in Table I. 

Well-defined chondrules and chondrule fragments 
are composed predominantly of olivine and 
orthopyroxene. "Barred" chondrules arc 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 lurbid lamellae of glass. The glass has a bytownitc 
Composition, and is probably maskelynite. Some large 
and strongly shocked olivine grains that occur as bright 

Fig. 3. photomicrograph tit the Conk 007 meteorite in thm 
section, showing fragment of barred olivine ehomirulL 
(about 1 mm in diameter). 

purple crystals under crossed nicols are unusual and 
need further investigation. Chondrules composed of 
radial pyroxene crystals arc also present {Fig. 4). 
Several of these display B cryptocrystalline structure 
(Fig. 5) with strong wavy extinction. A number of 
granular olivinc-pyroxenc chondrules and porphyrilic 
pyroxene chondrules arc present. They contain some 
coarse, cuhcdral olivine grains and have a poikililic 
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 stale recrvstallised areas and abundant 

TaBI t I. Average vhvmical compositions of major minerals in Ihv Cook 007 meteorite. 



orthopyroxene wt% 
core rim 




n - . 


TiO : ... 


Feb . . . T . . . 


MgO . . . . . .... 


Na0 2 

K 7 o' 

c'rfl, . 




















i 1 






II h 









18 9 










■ ).n 














96 4 






En 80 7 

E«74 4 

En 5Sft 

Am ? . , 


Wo 9: , 

Wo, b , 



Fig. 4. Photomicrograph of the Cook 007 meteorite in thin 
section, showing a radial pyroxene chondrulc (about 
0.5 mm in diameter). 

* . 

Fig. 5. Photomicrograph of the Cook 007 meteorite in thin 
section, showing a cryptocrystalline pyroxene chondrule 
(about I 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 Fa, 

n = 30. The oithopyroxene shows 

only a small variation in chemical composition with 

a mean ferrosilite content of Fs t 

n = 15 and a 

wollastonite content of 1 mol%. Some pyroxene 
phenocrysts contain calcium-rich rims. These rims 
have a composition of Fs 15J with a wollastonite 
content of Wo 95 mol%. The composition of the 
clinopyroxene needles within the glass is Wo 3GU En 556 
Fs 84 (30 analyses) (Fig. 7). The maskelynite glass has 
yielded a composition similar to bytownite (11 
analyses). The poor quality of microanalyses is a direct 
result of the high degree of shock metamorphism. 

The pyroxene geothermometers of Wells (1977) and 
Lindslcy (1983) suggest that the Cook 007 meteorite 
was heated to temperatures of between 600°C and 
700°C during metamorphism while the meteorite was 

Fig. 6. Photomicrograph of the Cook 007 meteorite in thin 
section, showing planar fractures in olivine crystal (about 
0.2 mm in diameter). 

Fig. 7. Silicate mineral chemistry in Cook 007 meteorite. 
Isotherms show temperature calculations based on the 
calcium content of orthopyroxenes and clinopyroxenes 
(Lindsley 1983). 


M /.Blh 

Mill pari of the large H type of the meteorite parent 
body. After this the meteorite was probably remelted 
in a major pari by impact and heated to a temperature 
between II00°C and 1200°C, as indicated by ihe 
chemical zoning of pyroxene grains. 


feldspar altered to muskelynite, occurrence of 
mechanical polysynthelic twinning in orlhopymxene 
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 Stuffier ct al. (I99|). 

The Cook 007 meteorite has been classified as an 
H4 ehondritc- The olivine composition (Fa W5+)M ) is 
within the range of the H chondrites (Keil & Frediksson 
|%4>. 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 
wollastonile content of the orthopyroxene is similar to 
that found in other H4 chondrites (Scott el al. 1986). 
The presence of undulatory extinction, the strong 
planar fracture formation in olivine, plagioelase 


The author wishes to thank Mr Huw Rosser ol 
CHMMSA. 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. 


Hi-v<\n. A. W R. (1992) Australian Meteorites. Rec. Aust. 

Mas. Supplement 15. 1-27 
& Binns, R. A. 0989a) Meteorites from the 

Nullarbor Region, Western Australia: I. A review of past 

recoveries and a procedure for naming new finds. 

Meteoritics 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. Ibid. 24. 135-140. 
& Pking, A. (1993) Guidelines for the nomenclature 

ol meteorites from the South Australian Nullarbor Ibid. 

28, 600-602. 
Hhymann. D. (1967) On the origin of hypersthenic 

chondrites: Ages and shock effects of black chondrites. 

Icarus 6. 189-221 
Keil, K. & Frp.driksson, K. (1964) The iron, magnesium 

and calcium distributions in coexisting olivines and rhombic 

pyroxenes of chondrites. J. Gcoph, Res. 69. 3487-3515. 
Lindsley, D. H. (1983) Pyroxene thermometry. Am. Mar. 

68. 477-493, 
Scott, E. R. D.. Taylor, G. J. & Kljl. K. (1986) 

Accretion, and brecciation of ordinary 

chondrites: evidence from pctrologic studies of meteorites 

from Roosevelt Countv. New Mexico. Proc. Lun. Plan. 

St. Conf, 17. EH5-EI23. 
Sioekler, D., Keil, K. & Scott, E. R. D. (1991) Shock 

metamorphism of ordinary chondrites, Gcoch, Cosm. Attn 

55, 3845-3867. 
Van Schmus. W R & Wood, J. A. (1967) A chemical- 

petrologic classification for the ehondritc meteorites 

Gench. Cosm, Acta 31, 747-765. 
W6U-S, P. R. A. (19771 Pyroxene thermometry in simple and 

complex systems. Cottrr. Min. Peir. 62, 129-139. 
Zbik, M. (1982) Pore Spaces in Stonv Meteorites. Bull, 

Aeaii. Pol. Sc Terrc 30. $9-65, 



Brief Communication 


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 1 . 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 2 . 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 2 . O. microlepidotus is a rarely seen snake because most 
of its life is spent in rat burrows 3 . 



The Inland Taipan \0\\nnmus mitrolejridoWs) is .i I Jigv 
elapid snake endemic lo Australia. Considering hotb venom 
toxic iiv and average venom yield per bite. (7 tnicrttlcptdixtis 
is the world's most dangerous snake 1 . Despite its size and 
medical find scientific signi ftcancc. the status and distribution 
of this snake, have been difficult to ascertain. Alter hcin;' 
described in IS*N. (X mumfapidmis was not loimd again until 
|074 J _ Following its rediscovery, ti has been recorded from 
the channel country of the Cooper Creek and Georgia and 
Dutmantina Rivers of south- western Queensland and north- 
eastern South Australia 1 0. tnirrolrpid*na* i> a rarely ttOfl 
snake because most ot its lift IS ^peni in rai burrows 

|n April 19V2, u large elapid was collected tut the Coober 
Pedy to William Creek Road (29°Q3'S, I3$TO'B] (Fie. 1,1 
by local contractor Jeff Bolaiid- He recognised thai the suak L 
was different from the Western Brown iP.svudotMlit rwchafis) 
and Mulea Snake il'si'ihhfhiy uustrahs} with winch he was 
verv familiar. Jeff Boland subsequently collected a slouched 
skin Irotn the Moon Plain (2X°52'S. iMAft'E) (Fig. I) and 
sent both the specimen and the slough to Ute author, 

/* Mt Barry / 

- V 


\. ^ -> 


V K / .v - 

X. / r^U*- j _m- __ 

\ Jt **s. 

Coober Pedy ^<v """ 

\ F -S 


* 0. mizralepdohis lotaltiies 

\ \ 

Fig. I, New records of OvvuntrtHS mi* nih-(n,itthi\ Di On- 
Coober Pedy region. 

Ihc dorsal scales o! the specimen were uniformly dark 
brown with the head nearly black. The specimen and the 
slough had 23 mid-body scale rows and a single anal .-.ealc 
These characteristics enabled the snake and (he slough to be 
confidently identified us (X mu rnkpidntus. The sealation oi 
the Coober Pedy specimen closely matched that ol O 
niicrnicptdoim specimens from Moomha except thai rather 
than the lower primary temporal scale extending to the lip 
between the >lh and 6th labials, this scale had fused with the 
6th labial in the Coober Pedy specmien- Inspection ol 'other 
specimens in the .South Australian Museum indicates that the 
status of the lower primary temporal scale is considerably 
plastic m p. fHwrnU'pidiHUs 

O. mnroicpuLmts can be further distinguished from the 
syrnpalnc. and highly variable/ 3 nm halts m having a longer 
head, smaller and more numerous nuchal scales, a pronounced 
canihus and plain ventral scales as distinct from spoiled or 
dark ettged ventral scales in P tuwludis. In addition, mosi 
P. ntuhiths from within the range cjf.lhe O. microlepidotus 
evhihii irregular black spots ot black erossbands. whereas 
these bands wen. absent from the Coober Paly and M'*omba 
specimens ol the 0. micndipiilutus. 

In trie winter of 1992, two road-killed O. nufrolcpt dolus 
were found on the Coober Pedv to Ml Barry Road |28°34'S, 
(34°.54'& 28°33's. \W3Sl fSAM R4()4t)4|). a live 
specimen was sighted on (he Dingo Fence (our kilonteties 
south of the onuma! locality f2V>°U5'S, I3SWE) and a farUicr 
specimen wan "killed at lorn Cat Hill [29*00 f & I34°45'n.) 
in Coober Pedy (Hg I). A further road killed specimen was 
located on the Coober Pedv to Mt Barry Road (2S°57'S. 
)M°47'F ISAM R424K4]) n September \m\ South 
Australian Museum recoids Are denoted by the prefix SAM , 
On September b. W$ d live fi 'utcrtMcptdotus was captured 
by Jctl Boland on the Moon Plan, adiaeent to the Breakaways 
Reserve. The following week a ruad killed spa'imen 13VWS, 
Lv5°IOTj was collected a|iproxmiateU. 40 km east of Coober 

The live specimen was eaptured in the rain with a strong 
wind blowing and a recorded temperature at Coober Pedy 
ot only \5°C- This snake was uniformly black on the doisal 
surface with a white, unmarked ventral surface. Ope month 
alter if* capture the snake sloughed, revealing a yellowtsh 
belly and a dark brown dorsal surluee wirh black edges io 
some wcales producing a shghi herring bone pattern The head 
remained a glossy black colour following the slouch 

The O. minvlt'pulofus from Cnoher Pedy region were about 
500 km from the nearest known O. tmirolvpidutus locality 
at Goyder Lagoon., m noitb-eastern South Australia. The 
Coober Pedy population of O. nu'cn'h'puhtu.s is separated 
from that or the channel country by the huge salina of Lake 
Eyre and ihe dunelields of the Simpson Desert. The discovery 
of O, nrivroU'pulotus near Coober Pedy is highly significant 
as it represents the first known* eturrenee of the species near 
a considerable population centre and also raises the possibility 
of u much more expanded range than previously recognised 
lor this tnrportanl snake 


The ('oobei IVily specimen* of O, p/.'nioicpuhtus were 
found Irom. April September, Interestingly, iheGoyder Lagoon 
population ,j| O i>!fcn>ft-{yhioiit\ has been recorded 
predominantly in Match and April whereas CovucevieV 
report* that f! tnnttdi'pnU>tu\ only SIDCiycR lor 1-3 weeks 
iri lale -w inter -carb tytjog, Although Covacevtch? indicate 
that (> m/i n'lt'fndotiis ate piidomrnantly active on siill days, 
Mirtsehin* collected several individuals on windy Jays. The 
Lupntit: ot the specimen nennhc Hreakuways Kesetvv on g 
cold, wet, winUv day indicates dial these snakes may be 
riKOnnteied .ir any tunc ihmiij'hour aulumn, wmti*f and 
spring. 0. micridcpUhius arc possibly also active in cracks 

and mammal botes during iii«' wiinni'i months but vc rmj 

recorded because lltcv do not need to come lo the surface 
to bu.vk 

O tun n>tf{tit!nJu\ recoid.s funn flit- (Wht Pedy region 
weie IoiiihJ in IwoddlWctU habitats HlVC ri| the records were 
from cracking I'ypseouM.otls. lour of records were from 
a lao/c au-a ol g)pst sous soils known is the Mi Kin Plain. These 
nuiK JuppOrl little or no perennial vegetation. Co\et and 
spines diversity ot annual vegetation val\ dramatically 
depending on Ihc season. The dominant, albeit sparse. 
vrjtcMltiin dm nig Orlober 1992 consisted ol Alriph \ 
spon$i<>\ii. SUlxoLi kali , Ifalipu-rum jhmbutulum. huiUumu 
/rfhtjrf>i\ Arui'uirlin nnuuriium and the grasses f&tHQQflpflfftt 
pid Kfthsllux . l'/irtn>{ um decnmpiniturn , .4ti,\{idrr 
amttoMHtfuutit'S, AmixWo pc-ti'muui. &Cgtoxtft xttifaUa and 
t' dirldr Vegetation envcT ww. pealest in the sniall inn-on 
ureas in the undulating plain. These low lying mc^ were also 
the: most heavily cracked. 

Two other records ^ere fiom gihhei count) y dominates! 
by the Oodnadatta Saltbu.ah irriplri nummidurta. Other 
common plants include*.! Srh'mdifthj iMfhtitii, Atrtpk'S 
\{>tn\::ti>M\, Snrcuaientu and Snlsoia kult Small gibsn 
»U-pirssnms Wilhol this ltahir;tl i .munricd a.u klpj i lay iilt.t 

wi_re vegttatcd wiih several graw and daisy -.pecies. 

The plains country around Coober Ivdy b. very 
hctcn^cucous and ihc two hahiwu* dcscj Fbed dbuvc uftpfl loi tu 
a inrfsnu with the gibbet more '.-onimon on hic.h ground and 
die vvpseous soils usually m low lying country, Therefore 
snakes m this re^.ton may depend upiin, or prcrtrr, etthei hiihii,-)! 
and yet he recorded crossing between arrjl&iif lavoured habitat 

The iruikuif' fvpscous .soil habitat is very similar lo the 
preferred habitat described for G m'h rofvpnimus at Clifton 
Hills, South Australia 1 HM "t QUCCftfrKM* .dthoajih lhc> 
have also heen recorded from gibber plains and '»vul dunes . 
I v -e cracking plains therefore uppew to be the key habitat 
for the O, mtcrofepufauis The cracking gypseous plains are 
qune widespread near the south-M-est margin of the I ..ike I'.y re 
Hasm and ure often asKOtialed with runoff /one* bom hir«k 
iHMjQf rountry These plains are often interconnected by creeks 
which |>ossib!y also provide :rppropna.te habital 

Inrerestingly the Moon Plain and surrounding ^ypstous 
TO&tOfp! were nol identihed as ptttential siIl-s kn () 
rrturtf/epjdtHij.s based on climatic indices' . Habitat atid 
biol'>gieal indices, ntthcr than climatic variables may tftca-Ibtv 
hit more important than climate in determining the range of 
(.' ffttt rolfpidtxus, 

The eastern population erf O. mirrotfpidiHu.\ fteds 
v'\lri!Mvely i>n tht: Long-haired Rat tHaltu-x vittmiyitnus) wiTti 
winch its ecology and distribution have been;ibl\ 
linked' 1 he cvolalton ot a laie~r si/*-, n rapid snap-release 
bite and t.\tremely potent venom in (I mnrvlepiti*)tus is 

believed to be- a tcsjumsc lo their prcdatiou on uiamnials ftflltt I 
Pill defend tllcmscKes h> bmtlj* sava/elv' IhC'if-r- 
particular emphasis vvav placed on Ncarchmg lor R 
\'itht\i\hnux in ultlei niiiiiI^ii si/ed mammats in the Coobt-- 
1'r.iy *-.'mon. io predict the boltmttSl ranee "I u 
mil mh'puloim , 

Field surveys were eoniuicied dunny B92 W % fnc rm kir* : 
cla\ regions adjacetu to the fl, »iicr<tlfpuii)His records on Ml 
iJarry, Annw Creek. Hdta KoJfOI ain't SrurtH I'tcek SWftons 
(< i .iscettam wbvthcr (\ m'u ruicpbktfUS, <»r their pa'y spcci'-s. 
ixeupied a more extensive range, The striking feature of the 
Cincked gypseous soil localities was the diversity ot muriimaK 
and paucitv of reptiles. Of particular tiole was thu presence 
cjI Plains Kat.s if'\n«ti.'rrl\,\ nuxtndis) Although A' r///<j,a\:w.. 
reached the CiMher Pedv region in (473-71 (1 Botand K 
tirvenhuliJ pei% eomnt.i they do not appear to inhabit tlv 
region nonnallv .md were not recorded irt litis sun'ev. }' 
tin\troh\, FrtrrCfttlB mice {iry^adinu (4>in\\U), Uescrt Mk, 
tpXtUifotmS drsevUH), House Mice (Ww.v doimsfn NJi I, 
pHucidou' l J t;migatcs iWiiniwdt' e'A'V''') and Pumratl*- 
(SmmThi'pMimncmui'a, s crtt^fiCaudaxa) wew Iftmiecl thtt 
a Wide range, M3iutnal densities wcic highei III the eta-. \ n 
gypseoii*. soil*, ihan tn other habitats in the rey.ion H , While 
rf|C .alnll O wirtolfpidnfHS jnohaf)t> Iced picdommantly <x 
P. utistmiis, L jorrcm and the occasional binl. juvenile snake* 
Ltuilit teed on r° yjirxii and M d<tttn-\iirn\ 

BctiAUgti Ogfa iiinfinnrtl uviads now Wist liuin ditt»u-til 
localities, initial suspicions that the specimen-, represented 
snakes thai were artificially tianslocated from then bftginftl 
ranee can o(i»tjat»ly he discounted However, H Ls not Kn n wn 
wheiher this apparent rsnzc expansion is a recent 
phenomenon, facilitated by mobility of ihc tiorrh eastern 
populaliou, or whether O, wUndcpidodis have occupied Ihc 
plflins orouftd I'oobet Pedy for a long periiMl CummiK-uh!' 
ihc de]x:udetice on Ihc R. vilfosismms tn the channel country, 
a potential sienatio is thai die i> MJi rtdi>pkhin\ tMlKWcd 
the rats Irom north eastern South Austnelia through to Coober 
Medy dunmi ihc m( piaphe ot l-)7b It is mihkHy, lntwcvr,, 
that O. nrit MtrpidflbUi could have expanded its range by over 
•MM! km dumig ihc romsc of 8 single rat ptaglic A nft?N 
tcasibk- ex|il;malion is that O, tifiiT'ttrpidofH* liaw ah\:tv 
inhabitul the gypseous plains around the south western margfjj 
ot ihc luk* Eyie basin bm the sparscucss of the human 
population combined with ihe prvdomtnaiitly underground 
Qa^U t'l the snake have accounted COj the paucity of tccords 
These same factors pn)bably explain why O. torn /\*lrptdotus 
eluded discovery in (he ftirdsMlie rceton lor so long. Increased 
snake number- as a reMitt of targe mammal populations winch 
huve K\s|>t'jn1ei! io wood seasons, combiritjd with increased 
vigilance ,ind awareness oj total lesulents probabls explain 
why the <). hticMtqpiilotus was discovered in the region in 
I9ft2< A edsninued intciest in this species is predicted to result 
in the disiuvory ol (I tw< rflrpidoJttx over a broader njfi^r 
in 'tacky ^vpveous couutiv t" ihc sinilli and west of Lake 

This papei is dcdirate.t lo Jell Boland whose interest ;"ul 
awatCTic.s rcsnllcd in the discovery of O, rmcrok'pido\us neat 
C<viber Pedv Thanks also \$> to Mick and Den Hvans. Grce 
SUiglviuer. Peter Paisley. Btc Mussared. Katherine Moseb\ 
Steve .ind Jama Green, John Fewster and 2ofs Kowen tir»f 
assisiam v witli behl woik and io Murk Huu hins»in. icaneitc 
Covacevich and Peter Mirtschin for help in lite preparation 
of this paper, Frank Badman a:;stste<l with plant identification. 


'Broad, A. J., Sutherland, S. K., & Coulter, A. R. (1979} 

Toxicon. 17, 661-664. 
2 Covacevich, J., Peam, J. & White, J. (1988) Venoms 

and Victims. Amphion Press and Queensland Museum: 

3 Covacevich, J. (1993) Journal of the International 

Herpctological Symposium. 

4 Mirtschin, P. J. & Reid, R. B. (1982) Trans. R. Soc. S. 

Aust. 106, 213-214. 
5 Mirtschin, P. (1981) Herpetofauna 13, 20-23. 
6 Shine, R. & Covacevich, J. (1983) J. Herpetol. 17, 60-69. 
7 Longmore, R. (1986) Australian flora and fauna series 7. 

Bureau of flora and fauna, Canberra. 
8 Read, J. L. (1992) Rangel. J. 14, 145-156. 

J. L. READ, 3D Eyre Court, Roxby Downs, S. Aust. 5725 




Brief Communication 


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 13 . Hence it is now used extensively as a 
diagnostic tool 1 3 . 

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 1,2,4 . 

Ihin.uiiiion* of thr f<u\al Soncty of S (IW4), 118(2), 147 I4H. 



The majority ol known Australian fossil hogs have been 
identified by the characteristics of the ilium, a distinctive bone 
in whiih the morphology vanes greatly between family, genu*, 
and species' 1 , Hence it is now used extensively as a 
diagnostic tool . 

The physical characteristics of the frog can be identified 
Irani ihe si/e and points lor muscular allachmenl on the ilium . 
These features represent adaptations to the environment, For 
example a short dial shaft is characteristic of a burrowing 
frog which does not make long jumps 1 "'*. 

To permit extrapolation of the size of a frog from the length 
ot Ihe dium requires an understanding of ihe relationship 
between Ihe ilium length (ID and snout to vent length (S-V). 

It is generally considered that Ihe dium grows linearly in 
relation to the S-V of the donor animal*, 

However, the data from which these assumptions are made 
have been based on small sample sizes, and little is known 
of Carl) ontogenetic changes 1 . ll is also important to note 
that it is common in the Animal Kingdom for a change in 
si/e lo icsull in u change of" the proportions of the hotly- A 
simple example Of such allometrie growth can be seen during 
Ihe development of human beings, in which a large change 
in the proportions of the limbs and the head, in relation ro 
trunk si/e. can be seen when children are compared with 

This study was an attempt to determine the nature o! growth, 
ln»m juvenile lo adult form, ot the burrowing frog Cycforamt 
austndts (Gray. 1842) via the relationship between ilial length 
and snout vent length. 

The ilia examined were dissected from 24 preserved 
specimens in the collections held at the Department of 
/oology, t niversitv ot Adelaide. Before dissection the S-V 
of each specimen was measured with a pair of NSK electronic 
digtial callipers The pelvis was then removed, and soaked 
in bleach to loosen muscle, before being transferred to a V°C 
oven to dehvdrale f*»~ 24 hours. The IL, tor each ilium was 
measured using electronic digital callipers, with Ihe aid of 


1-ig I Left lateral aspect of Cvrhivtta austmlis pelvic girdle 
UL - 30.6 mm. S-V - K2.1 mm). 

Abhtevitiihms. D, acel cxp_dorsal acetabular expansion, 

I) pmm,_dorsat prominence; p. prolub. dorsal 

protuberance; pre acel. /one pre-acelahular /one. 

a Wild M3 dissecting microscope. Ilial length was expressed 
as the distance between the lip of the dorsal acetabular 
expansion, and the end of the dial shaft 1 

The characteristics of the ilium of the specimens examined 
did nol differ significantly from those described (see Jig, 
It. The S-V ranged 20.0 mm to 83,1 mm. and IL 6.1 mm 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 
ilium, i.e. the ilium grows linearly in relation lo the body 
length throughout ontogeny (see Pig. 2). 

The linear growth of C oushntis permits a very accurate 
method for estimating the size of an individual from a 
disarticulated ilium. 

I am greatly indebted to Prof. M. J. Tyler lor the 
opportunity lo undertake this research and for constructive 
criticism of the early drafts:, and I am extremely grateful lor 
(he assislance from K Maurice-Jones in the production ot 
the figures. 

1.90 - 

lag tn V - 0_n£>?H + K372 log 1D X 
r* = 0.9517. n*24 

0.9 11 1,3 1.5 

log l0 Ilial length (mm) 

Pig. 2. Regression line of ilium length of Cwlonmo uustml'ts 
Willi snout- vent length For •: = 18-30 mm,y 51.25 mm 
(95% confidence limits =49.36 53*23). 


Tyler, M. J. (1976) Trans. R. Soc. S. Aust. 100(1), 3-14. 

2 Tyler, M. J. (1982) pp. 159-168 In Rich, P. V. & Thompson, 
E. M. (Eds) "The Fossil Vertebrate Record of Australasia" 
(Monash Univ. Press, Clayton). 

3 Tyler, M. J. (1990) Mem. Qld. Mus. 28(2), 779-784. 

4 Trueb, L. (1973) pp. 107-108 In Vial, J. L. (Ed.) 

"Evolutionary Biology of the Anurans. Contemporary 

Research on Major Problems" (University of Missouri Press, 


5 Tyler, M. J. (1993) Pers. comm. 

6 Bogin, B. (1988) "Patterns of human growth" (Cambridge 

University Press, Cambridge). 

STEVEN J. WALKER, 12 Preston St., Blair Athol, S. Aust. 5084 

VOL. 118, PARTS 3 & 4 
30 NOVEMBER, 1994 

Transactions of the 

Royal Society of South 



Davies, M. & Watson, G. F- Morphology and reproductive biology of Limnodynastes 
salmini, L. convexiusculus and Megistolotis lignarius (Anura: 
Leptodactylidae: Limnodynastinae) ------ 149 

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 --------171 

Pamment, D., Beveridge, I. & Gasser, R. B. The distribution of nematode parasites within 

the stomach of the western grey kangaroo, Macropus fuliginosus - 187 

Ainslie, R. C, Johnston, Dl 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 - - 197 

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 - 207 

Hutchinson, M. N., Milne, T. & Croft, T. Redescription and ecological notes on the 

pygmy bluetongue, Tiliqua adelaidensis (Squamata: Scincidae) - 217 

Fuller, M. K. & Jenkins, R. J. F. Moorowipora chamberensis, a coral from the early 

Cambrian Moorowie Formation, Flinders Ranges, South Australia - 227 

Peterson, M., Shea, G. M., Johnston, G. R. & Miller, B. Notes on the morphology 
and biology of Ctenophorus mckenziei (Storr, 1981) (Squamata: 
Agamidae) - ____ 237 

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 ----------- 245 

Olsen, A. M. The history of the development of the Pacific oyster, Crassostrea gigas 

(Thunberg) industry in South Australia - 253 

Bird, A. F. & \eates, G. W. Studies on Aprutides guidettii (Nematoda: Seinuridae) isolated 

from soil at Northfield, South Australia ----- 261 

Brief Communication: 

Tyler, M. J., Davies, M., Watson, G. F. The hylid frog Litoria alboguttata (Giinther) 

in the Northern Territory -------- 267 






VOL. 118, PART 3 





By Margaret Davies* & GraemeF. WATSONf 


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 

},<in\<iai<>ii,\»f:I tl Kf\\-tl St*dit\ v) 5 finxt, tt^Ur liffifl, 149- 16&. 






D'WIIn. M< & W'aison, C5, I. (1904) Morphology and reproductive biology of t./mmniynftMiw ^tl'tntu, I 
t otnvsittMithis and MvMhlvktiSS liwtwius (Anura Lcplodactylidac. LiiiinodynuMiiwci. rftftfJ. tf. .W. V AuM 
iWi-M. I4 l > l<>9 M) November, IW. 

I Alcrnal morphology, osicologv, siruciuiv ot ihi larval ehondroerainum, development and call cbaoiffipi l&tcs 
ol ihe limnodviiasime frogs Unmodyiut>ws sufnUfd and A. tnrivtxtusivhts arc described and compared with snnilai 
data derived from McgtviftfolH U\>hnrm,s. These tiiUd are pre.*iemed lo coninbtilo u> ftur undeManding ot ihe biology 
ol ihe anurjjn luima us well us Ui a wider analysis o! ihe genera Linmnd\tia\(r\ and A/re/v/i'/'J/n being undertaken 
lo examine the hypothesis that M. fii^nanus is the sister (axon lo I- iimwuimuius and that /,. \<ilmmt is Ihe 
eldest relative to this species pair, 

KI-'Y WORDS. Lttnti'hlwasU's c<>niv\utsailus, bnnuhfauistts snimini, Mei'tsiolotis lignanus, Irogs. calls, larvae, 
lite history, osteology, morphology, larvul ehondrocianhi 


Uttokuhnasii's Fit/mger comprises fl species (Tyler 
Ip92), generally ueeepted to tall into three species 
groups lite L. immaniensis group, the/., rforsalis 
group and the L- omuius group (Tyler vt ai 1979, 
Koherts&Maxson 1986; Mahony & Robinson I9H6) 

Ihe composition of three groups anil the 
complement ol the genus has hecn challenged by 
Hutchinson & Maxson (1987) who proposed, on the 
basis of data derived from comparisons ol albumin 
using the immunological technique of* micro- 
complement fixation, that Limno<i\nastv\ is 
paraphyletic and thai Mcy.istoltms hgnanus is the 
closest relative of L iomexiusvulus. Uinnotlxnttst< J \ 
sahnini is proposed a.s the closest relative oi these nvo 
species, all three tailing in the L. lasmtmUmsis species 

Tins contention has not been widely accepted 
(Coggct 1992; Tyler 1992. Liuicjohn a at. IW3-: 
Rohcrts h Watson 1993) and the question will nol be 
resolved until data are provided lor a broader analysis 
ol ihe genus. Such data should include morphology, 
osteology and biological data such as calls and 
developmental information. Although the recognition 
ol these three Laxa is not in dispute, availability 
data \anes hetween them. 

lVpurimcnt ol Xoolngv, University of Adelaide, S uOl 
Australia 5005 

IX'purtmeni ol /oolouv, Unoersiiy bT Melbourne. 
Purkulle. Victorm JmL 

Parker (1940) provided morphological descriptions 
of (IlO&t species of Ummulynusuw and included some 
information on ihe osteology of the ctfrpajia ds well 
as the nature ol the hyoid. In man) cases he had limited 
material available to him. Moore { 1901 ) pmvidcd some 
further morphological data lor selected species together 
with distributional data whilst Lynch (1971) diagnosed 
the eenus on the basis ol osteologieal characters derived 
from tour .species, not the laxa under consideration 
here. Mflfifisniforti H$tuinn.\ was not available, ot 
course, to any of these worker*. That genus and species 
were descrihed very comprehensively by lylcr tit ui. 
|1979». However, Ihe description did nol include a 
formal description of the skull or of post cranial 
elements, although the skull s\as illustraied. None ol 
these authors examined the chondmcrania Of larval 
forms of the taxa 

As part of a wider study examining the phylogenelic 
relationships oi and Megistolotts and 
of ongoing studies ot the hiology of Austialopapuan 
frogs*, we provide data on tin* morphology (including 
data from tfu- LbondiociarUiiu) and biology Of £, 
Vi>nyc tiu.sittlu.s , /„ sulrwni and M. li^/uiriits and 
consider these data in the light o\ published literature. 
Although Parker i W40i provided relatively 
comprehensive descriptions im ibe external morphiilogv 
of/., saimini and L omvexiu\ti(lu\, his sample size 
was necessarily MUall. v> v\e provide lurlher 
descriptions including measurements in a standard 
lormat to allow for direct comparison belween taxa. 
t'or the same reason, we include a furthei lUustration 
of ttic skull of M lixnatius ui a ciunpositc llgure ol 
the skulls of all ta\a 



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 at, (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 lluctuating field 
conditions; on return to the laboratory, they were reared 
at 30°±1°C in glass aquaria (25 X 25 X 8 cm) 
containing aerated dechlorinated tap water. Larvae 
were fed boiled, organically-grown mignonette- lettuce 
leaves supplemented with Sera Bioflakcs 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 Sll (L. convexiuscuhts 
and M. Ugnarius) 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 1 ms for temporal aspects, and less than 
40 Hz for dominant frequency. Three calls of each 
individual were analysed. Calls of M, Ugnarius are 
those of the holotype and paratype described by Tyler 
et at. (1979) and reanalysed here using more modern 

equipment to facilitate comparisons between the three 

Limnodynastes salmini Steindachner, 1867 
FIGS 1-14 

Limnodynastes salmini Steindachner. F (1867) Amphibien 
in "Reise der ostereiehischen Frigate Novara von die Erde 
in den Jahrcn 1857, 1858, 1859" Zoologie 1(4), 1-70. (State 
Printer, Vienna). 


NHMW 14849 (two syntypes) from Cape York Qld 
(as Australia) (Cogger et al. 1983). 


it ' 

Fig. 1. A. Limnodynastes salmini, B. L conve.xiusculus and 
C. Megistolotis Ugnarius in life. 




A Urge species (males 43-7o mm, lemalcs 61 mm) 
characterised by yellowish dorsolateral skin Folds, a 
relatively distinct tympanum, glandular nuptial 
excrescences in the male, poorly Hanged ringers in rhe 
female, first finger longer than the second, male call 
a resonant "unk", deposition of eggs in a foam nest. 
Description (based on SAM R 4X969) 

Head depressed; slight!) longer than broad (HL/HW 
I.()7l slightly more than % of snout-vent length (HI 'S- 
V 0.35). Snout prominent, ovoid when viewed from 
above and rounded in profile (Pig. 1) Nures dorsal in 
position, then distance from end of snout less than that 
from eye. Eye to naris distance less than internyrial 
span (E N/IN 0.92). Cantluis rostralis well defined and 
straight. Lorcal region straight, sloping ventrolateral^. 
Lye moderate in diameter Tympanum distinct, 0.6 of 
eye diameter (Fig. 1) Vomerine teeth m long horizontal 
rows, meeting in midline, posterior to small lateral 
choanae. but not contacting thern. Tongue broad Vocal 
sac slits lateral to tongue. 

Fingers eyJindrical. unfringed. basally webbed (Figs 
2. 3); subartieular and palmar tubercles well 
developed. Supernumerary tubercles at junction of first 
and second fingers and second ami third lingers. Inner 
palmar tubercles large, outer divided Nuptial 
excrescences not detectable Tinners in order of length 
3>J>2 = 4. 

Hind legs moderately short (TL/S-V 0.40), Toes 
long, cylindrical with narrow lateral fringes and basal 
webbing ie\cept between iocs 4 and 5J (Fig. 3). 
Subaniculai tubercles large, conical. Large prominent 
inner metatarsal tubercle; rounded thickening Of skin. 

but no outer metatarsal tubercle. Iocs in ordei of length 

Dorsum with well-developed, elevated, pigmented 
glands Supralabial glands elongate and prominent 
(Fig. 1) Skin not covered by spines. Ventral skin 
smooth, pigmented. Canthui stripe through eye and 
tympanum to axilla. Two pale (cream! stripes laterally 
from 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 

The second finger and the thumb mi their medial 
sides are fiinged in 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 
it is wide (HL-HW. mean = 1.09. range L00-I.I8). 
Head length is about *A 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 internal ial span tE- 
N/IN, mean = 0.87, range 0.70-1.07). 
Material examined 

QUI: QNPWS N17095. Bundaberg, nr Mon Repos, 
N2X20O, N28228. N28251. Lake Nugga Niigga, AS5I. 
Gladstone, A6, Southwood N.P. (nr MoonieL SAM 
R4I%0-7(1 UA7 B17MX AI729. BI726 nr Ban Ban 
Osto'Io^x (based on AUZ AI61) (Fig. 4). 

Skull moderately well ossified. Sphencihmoid poorl\ 
ossified, not in bony conlaci with nasals, extending 

Fig, 2 Palmar view ot the hand ot A. feruatc i-imtm,h<rnistc\ >y>ti\'e\tuscufns (SAM R4l l J«3) B. RSmflie / sahutut (SAM 
R4I969) ;tnd C faflBje Mtytsu^mis tfmwftn (SAM K'1t986> Scale bar - 5 iuin_ 



■':■■■■;■-. v- 

Fiu. 3. Palmar view of hand and plantar view of fool of A- Limnodynastes convexiuscuius {SAM R41983) and B. L. salmini 
"(SAM R41969). Scale bars -5 mm. 



Fig. 4. A. Dorsal and B. ventral views of the skull of Lmituklynasies convexiusculus (UAZ A 233); C. Dorsal and D. ventral 
view trf the skull of L. sohnini (UAX AL6I); E. dorsal and K ventral views of the skull of Megistototis lixnarius (UAZ 
AI742> Scale bar = 5 mm. 



'A length Of orbit in ventral View. Prootie and 
exoecipital nut fused, Exoecipital not ossified dorso- 
or ventro-medially. Crista parotica short and stocky* 
not arliculaiing laterally with short expanded otic ramus 
Of squamosal, frontoparietal fontanel lc poorly 
exposed. Frontoparietals well ossified, anterior 
extremities extending S A length of orbiL Orbital edges 
of frontoparietals straight, not angled posterolateral!). 
Anterior margin of frontoparietal fontanelle formed by 
sphenetlimoid at a level about l A anteriorly along length 
of orbit. Posterior margin undefined because of 
liixtapostiion of frontoparietals medially and lack pj 
medial ossification of exoecipitals. Nasals poorly 
ossified, maxillary process elongate and moderately 
broad, in contact with well-developed preorbital 
process of pars laeialis of maxilla. Palatines broad, 
expanded laterally, running under deniigerous 
proicsst's of vomer to overlie edges of ventral 
sphcnelhinoid medially. Parasphenoid robust Cultri- 
forni process elongate, broad almost reaching medial 
extremities of palatines. Alae hroad. moderately long, 
expanded laterally, at right angles to eullriform process. 

Pterygoid robust. Anterior ramus in short contact 
with poorly developed pterygoid process of palatal shell 
of maxilla. Medial ramus slender overlying alae of 
parasphenoid Posterioi ramus very robust and 
elongate Junction of three rami very rohust. 

Quadtatojugal robust and entire. Squamosal robust 
with moderately long zygomatic ramus and short 
expanded otic ramus Maxilla and premaxilla dentate. 

Pars facialis of maxilla deep with well-developed 
preorbital process. 

Alary processes of premaxilla broad, slightly 
bifurcated dorsally. directed posteriorly- Pterygoid 
processes ot palatal shelf poorly developed. Vomers 
reduced medially with extremely elongate horizontal 
dentigerous processes. Columella bony, sigmoid in 

Pectoral girdle areiferal and robust. Slender 
omosternum, broad vipbisternum. Siernum 
cartilaginous. Clavicles slender, closely applied 
medially Coracoids very robust, widely separated 
medially. Bicapitate scapula robust. Supraseapub about 
l /2 ossified. Antcroproximal crest of humerus 
moderately well developed. 

Carpus of five elements (Pig. 5) Lateral process on 
medial surface of first metacarpal. Sesamoids absenl 
at junctions of metacarpals and/or phalanges. 

Seven precocious noil-imbricate presacral vertebrae. 
Vertebrae 1 and II fused. Relative width of transverse 

ill>IV>U = SD>V>VI>V1I>VIII 

Sacral diapophyses poorly expanded. Ilia extending 
very slightly anteriorly to sacral diapophyses. Urostyle 
crest approx. % length of urostyle. 

No ilial crest. Dorsal prominence prominent (Fig. 
6). Dorsal protuberance ovoid and lateral. Pubis 
calcified Three tarsal elements in fool Prehallux large. 
hastate, cartilaginous dorsally with bony base. 

Fig. S Dorsal view of the carpus of A. Limn>> j .\ convcsiasculus. B. L sttlmim and C. Mcgtsiolotis Uwarius, Scale 
bars = 5 mm 





Fig. 6. Lateral views of the ilia of A, Limnodynastes safmint, Fig. 7. Ventral views of the hyoicl in A. Limnodvtuistcs saimim. 
B. L. convexiusciiius and C. Megistolotis liynurius. Seale B. L. amvexiusculus and C Megistolutis lignahus. Scale 

bars = 5 mm. bars = 5 mm. 







4 6 8 



> 4 




UJ 2 





TIME (ms) 







TIME (ms) 

Fig. 8. Power spectrum, waveform and sonagram (sampling frequency 59 Hz) of the advertisement call oiLimrwdynustes 
salmini. Note that the ordinate for the waveform display is not labelled because it depicts a relative linear scale in volts. 



Hyoid plate wider than long; posterior processes 
slender, asymmetrical in length. Anterior processes 
short, blunt perforated. Short, broadly based 
anteromedia! processes on anterior hyale. Posterior 
cornua ossified (big, 7). 


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 almost 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 facialis of the maxilla 
in the very large female UAZ AI729_ 

In two of the specimens examined, the exoceipital 
and prootie are confluent dorsally, but not vent rally. 

but again such ossification docs not appear to be size 

The other features of the cranium do not appear to 
be variable. 
Material examined 

UAZ A1729. female. Ban Ban Springs, B1730. male, 
same locality, B1726, male, same locality. A16I, sex 
not known, Brisbane, B1736, Stage 33 larvae, B 1737. 
Stage 35 larvae, QNPWS 17095. ?male, Mon Repos 
nr Bundaberg. 

Li tnnodxt tastes salmini calls from Hooded 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 lo 
be a possibility, but we have no direct evidence of this 
Advertisement call 

The advertisement call of/., salmini is a well-tuned. 
single note (mean duration 67.4 ms. Table 1) repeated 
regularly in long calling sequences. Call repetition rate 

Pig. 9. Distribution of Limnodynastes salmini I open squares) and L. eomexiieseidus closed circles. Symbols may represent 
more than tine close locality. 

Morphology anh reproductive biol<x;y 


Tabi £ I. Durutiotts ("is) of flit- mllx <>f Limnodynasles 
salmini. I ., convexiusculus and Meeistolotr- liunanuv Mrun\ 
and ranges ishnvn in parentheses) based on atudxsis a/three 
mils from eaeh individual listed 


Nil recorded Cull duration 

t.tmnodynasies salmini 3 

I nnnoilxnash's ronvesiusctthts 6 

M<\i:t\t<>lo!i.\ h^nartus 







for the three individuals reported here was 
approximately 0.91 ealls/s (at an effective temperature 
of 25.2°C). Waveform and spectographic displays of 
a single note and a power speetrum are shown in Fig. 8. 

LinmodymtsU's salmini has an eastern Australian 
distribution, the most northerly record being St 
Lawrence (Moore 1961). The most southerly record 
is near West Wyalong, N S.W. (Fig_ 9). These records 
cast doubt upon [foe interpretation ol Conger '' oi 
U983) that the type locality is on Cape York Peninsula. 
Natural history 

The species is predatory on oiher species or lings. 
One was collected with a L. antatus head first down 
its throat in the process ol being swallowed. 
Life history 

Eggs ure laid in a large, foamy egg mass. Two such 
spawn clumps contained approximately 1630 and 2410 
eggs respectively. These clumps were collected soon 
after deposition at 1120 on 2LU99I near Ban Ban 
Springs. SF. QUI. A sample of ten eggs hud a mean 
capsular diameter o\' 1.55 mm (range 1.44-1.68 mm) 
and a mean egg diameter Ctf 1.27 mm ( range 
1. 18-1.34 mm). There were two jelly layers surrounding 
pigmented eggs which hatched at stage 19 about 24 h 
after collection (Fig- 10). 

Just prior lo hatching (slill al stage 19). the 
stomodaeum was a pit anteriorly and a pair ot well* 
developed pigmented adhesive glands projected 
vcntrally. Two pairs of external gills were apparent, 
the first with two and the second with six short, broad 

At hatching, a depression in the presumptive eye 
region was apparent. 

At 0850 *>n 24.U99I. embryos were at stage 21 but 
the external gills had disappeared. The cornea was 

transparent The mouth had perforated and 
ketiHim/ation had commenced upon the upper beak 
The adhesive organs remained extremely prominent- 
elevated and slightly pigmented (Pig, 10), The nare.s 
were perforated, as was (he anus as a median aperture. 

Twenly-I'our hours later <>n 25.1.1991 embryos were 
at stage 24, The operculum was not fully formed The 
horny beak was keratinized and some ot the tooth rows 
were partially delectable. All ol these had keratinized 
by stage 26. 

Stage 27 was reached by 27 i. 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 al 
this stage. Larvae were at stage 29 on 31 i. 1991. Stage 
35 was reached on 2.ii.|99|. 

A larVa at stage 34 is illustrated n> Fig. iL The hody 
is ovoid and widesL al the level of the eyes. The snout 
is evenly rounded in dorsal view and slightly truncated 

Fig. 10. A. Embryo ul t.hnm>d\nastes salmini ui stage 20, 
B. at Mane 21 and C at Mane 19. Scale bars = I mm. 

' 2 

Fm II A, Lateral and B. dorsal view> of a larva of Umnodxtutstes salmini al stage 34. Scale bar = HI mm. 



ill lateral vicvs with an anteriorly projecting oral disc. 
The nares are dorsolateral and nol raised- Moderately 
huge c>cs are dorsolateral. The spiracle is sinistral and 
attached to the body wall along its medial edge. It is 
short, opening slightly dorsally and visible when 
viewed from above- Its diameter is constant along its 

The anal lube is broad and opens medially. The tail 
tins are gently arched and rounded terminally. The 
dorsal lui commences posteriorly to the body and is 
deepest about halfway along its length. The anal tube 
is broad and opens medially- The ventral tin 
commences posteriorly to the body and is deepest 
about halt 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 lores) region and under the eye (second sequence). 

Tf)C mouth is anteioventral. The oral disc comprises 
a horny beak of moderate proportion and there are tour 

to five upper and three lower rows ol labial teeth (Tig. 
13). The most anterior upper row and the two most 
posterior lower rows are undivided. Labial papillae 
arc interrupted anteromediall) but extend laterally and 
posteriorly around the oral disc. 

The dorsal surface of the body and the tail 
musculature and fins are heavily suffused with 

Larvae reached stage 36 by 7.h.l99I and stage 3° 
by I0.ii. 1991, Stage 41 was reached by I2iU99l and 
the species had fully metamorphosed by 5,iiU99L 43 
days after spawning. 

Measurements oi~ developmental stages are given in 
Table 2, 

Chondral milium (Stage 34 larva, based on VAZ 
B1736) (Fig. 14) 

Neurocranium approximately T-shaped box 
comprising anterior braincasc and posterolateral 
spherical otic capsules. Large ovoid frontoparietal 
foniatldk exposes braincasc dorsally bounded by 
orbital cartilages laterally and by tectum synoticum 

Fig. 12 A. Lateral and B dnrsul views of a larva -<t Litnm<tl\nu\its SuJtnttoi ji «afiC 34 shmving location ol ihe ueuiomiiNi 
tells of the lateral hue System. ScaJu bar - _s min 

I iv IV A. Oral disr of larva ot UtfititnhtiuMcs i<>tnrunMt//n\ 31 \\w ttJttuJB '»' / Wwm/ai stage 34. Scale bar - I 




posteriorly Narrow ethmoid plate forming anterior 
margin of frontoparietal fonlanelle, Ethmoid plate 
projecting anteriorly from neurocranium as two 
anterolateral!) diverging moderately hroad linger-like 
eornua trabcculac. Floor of neurocranium comprising 
basts cranii and planum basale. Basis cranii perforated 
by paired foramina caroticum primarum. Posteriorly 
planum becoming confluent with otoccipitaJ arch and 
perforated by notochord. 

Massive approximately L-shaped palatoquadrate. 
Processus descendens (attachment to neurocranium) 
widening laterally to form arcus subocularis separated 
from basis cranii by oval subocular fenestra. Medial 
processus ascendens overlain dorsally to about half iis 
width by anterior extremities of otic capsules. Arcus 
subocularis curving laterally while sloping vemrally. 
Anterolateral edges bearing low crest, becoming 
progressively higher and confluent with broad 

processus museularis quadrati. Arcus subocularis 
attached anteriorly 10 neurocranium by eomimssura 
quadratocranialis anterior. Finger-like projections 
extending vcntronicdially into subocular foramen. 

Processus museularis quadrati inclined medially, 
bound to laminorbitonasalis by non-ehondrified 
ligamentum (cartilago) tectum forming tunnel with 
commissura quadratocranialis anterior through which 
pass M. levator mandibulae posterior and M. 1. in, 

Processus articularis quadrati of palatoquadrate 
extending anteriorly from processus museularis 
quadrati and articulating ventrolateral!} 1 with slightly 
curved Meckel's cartilage 

Ligamentum quadrati ethmoidale attaching laterally 
about half-way along eornua trabcculac. Dorsomedial 
edges of Meckel's cartilages attaching with small 
shallow mfrarosiral cartilages which form shallow 

Tvhii 2. Mi-n\U( dntu <>n tlnrtnjiinx I.nnnodynusics sulmini. // = number nfiiuHvidiHils, \ = ttivott, 

Age. days and 


Body length \ 

range in 


Total length v 

range in 


25. i. 199 1 




5-20. ii. 1991 




3-15.ii 1991 

43 56 
2-15. ii. 1991 


2-18. ii, 1991 




2-m.ii m\ 



l5 20.ii.N91 




55 M 

I2 20.ii.!99| 










(4 96-7.04) 

(6.2 8.16) 

7 93 





(8.2-14. 2) 






(14.7 17.4) 

1 16 3 19.9) 

1 13.9-18.5) 

< 13. 1-19.9) 









111.6 16.4) 
























(23.9 40.3) 












42 92 


( 39.9-48.5 1 






(39.4 47.4) 





imcilcdU in ventral wew wiih hroad non-chorkJrified 
symphysis, elements joining to form lower jaw a) 

Supra rostral curtilages forming upper jaw ol "tadpole 
ptoieclmg anteriorly ventral toeorriua traheculae and 
lo both Meckel s curtilages and intrarostrals dorsal 1 v. 
Ccultal corpus of suprarosuul lying anteriorly in 
extremities o| coinua trabeculae with non-chon<Jrified 
medial symphysis, Alae extend to level ol Meckel's 
cartilages laterally. 

/ .imwxlynaslc* convexiusculus 

iMaeleay. JX77) 

l : 1GS 17. \ n \s 

ttimiVih'rnm\\\ius%'utns MueteuA. W tlK77i f&V Lw»i Rm 

\ A' If 2. 135-138 \\if,\. 

LimtUhhiH^'i^ tJih'UflHtt Dtf Vis (1844) r>W, Sac 

X SM. i*. 65 -M (66i. 

Wuiiunitit:, noxat^iiiiu-iit' van Kampcn. F* Nf. ilsKNt Mhw 


Holotype. presumed losl, from Mavvalta. Binaturi 
River near Dafli. Papua New Guinea (as Katowi 
ICOUpr i7 0?. 1983). 

A moderately large species t males 44 54 mm S-V, 
females 44-ti4 mm S-V) characterised by a prominent 
sacral hump, a mid-vertcbrai stripe, a rugose d- rsum 
wiih longitudinal skin folds, an indistinct tympanuin. 
nuptial excrescences in I he form ol line spines mi the 
maJc. females with well-developed llanges on Ifrsl and 
second fingers, First finger shorter or equal to second 
in length, male call a resonant "unk'l deposition ot'e^s 
in ;i loam npftl. 
Dfsihptitm t based m SAM R41983) 

Head high, slightly longer than broad (HI 'HW M4j< 
less than hall sinful to vent length (HL/S-V 1X4 h. Snout 
not prominent rounded when viewed from above and 
in profile (lie. I) Nares dorsal in position, then 
distance from cud of snout less than that from eye. 
luternanal spun approximately equal to eye to nans 
distance (K-N'IN 0.96 j. Canthus rostralis poorly 
defined and straight rye moderate, diameter greater 
Ihan eye to nans distance. Pupil shape vertical. 
Tympanum relatively smalL not clearly delined. about 
O.K eye diameter (Fig. IJ. Vomerine tcelh IN two 
elongate series, curved across the palate posterior to 
t'hoanae; not meeting medially and extending laterally 
in i-ytrcrnitics ot small ehoanae. 

Jougue bmad. fingers cylindrical Second RflgCf 
••lielitly fringed medial to thumb trigs 2. 3), unwebbed. 
Subarticular tubercles moderately developed, rounded. 
Inner palmar tubercle very prominent, raised: outer 
divided. Nuptial excrescences in form ol large ovoid 
group of fine spines on medial surface of thumb with 
small group on dorsal surface of basi: nf second finger. 
Interdict Ul webbing absent. 


I inyers in ordet oi length 3>2>! 4 Hind legs 
rmtdcutely short (Tl S \ r 046). Toes long and 
cylindrical with very uanow lateral fringes. 
Suhaniculut tubercles pprtinnl and prominent: elongate 

ovoid inner metatarsal tubercle: outer metatarsal 
tubercle absent. Hies unwebhed other than basal 
fringing fPife 3) In ordet ol length 4^>5>2;>! 

Dorsum rugose with pale mid vertebral stripe. Pelvic 
region humped (rig. It- Chocolate-coloured, well 
defined markings of elongate or small circular shapes 
trending as dorsolateiat anteroposterior patterns. 

Body and limbs with numerous very small conical 
black spines, nb! prominent on lateral side of head arid 
on eyelids; on mandible, posterior portion of body. 
Literal surface of forearms, dorsal and lateral surfaces 
o^ legs and plantar surlacc ol loot, 

Ventral surface smooth, /ream; throat speckled with 
pigment. Well-developed suprulabial gland (big. 

The hind legs are moderately short, I L S V mean 
043, range 0.38-0.47. The head is longer than broad 
in most specimens (HI 'HW mean = I I L ranjir 
0.98-1.23) and is less than hail the snout to vent length 
(HL'S-V mean = 0.38'. r<nge 0.33-0.41). Hye to naris 
distance is usually less than interuarial spun (L-N, IN 
mean - 0.93, range 0.77-1 I4> 

The small group ol fine spines on the dorsal surlacc 
6f the base of die second finger associated with the 
nuptial excrescence in the described specimen was not 
observed b any other material, ihe nuptial excrescences 
being confined to the medial surface ol the thumb. 
rcniulcs have well-dou-loped flanges on oppose 
suifaces of the thutfib uxid Second fingers (Fig. 2), 

The dorsum varies, in tuhercularity and can be more 
or less tubercular than described. A degree of 
regularity m hack patlctiaug between specimens is 
evident The conical spine* on the dorsum are less 
developed in some specimens and the inner palmat 
tubercle is extremely well developed in some male 
Materia/ examined 

Northern Territory: UAZ 81738-° Jahiru. SAM 
R4I979 RaraitI Ck nr .labiru SAM R4W73 Darwin 
(northern suburbs*. DAZ B1773 Radon Ck Rd tni Mt 
Brockmanl. SAM R4HJ74 \1 km K Ten Roper River 
Rd/ Stuart Hwy. Darwin. SAM R4W7I:. UAZ BI72^ 
Snake Bay Swamp. Melville Island. 

Western Australia: SAM R41*>&0-83, Mitchell Plateau 
tupstream from Crusher sit;), SAM R4b)S4 -5, Mitchell 
Plateau campsite. SAM R41 n 78, Kununuvra. SAM 
R4W77 Parry Ck Rd Kuriunurra. 

Queensland; SAM R4l975-b Hdward River 
Township, QNPWS a.^3 Sugttf Cane Creek. Tully 
Mission Beach Rd. A39f--b Kurraminc. Murderim* 
Poim. N264I-2. N2659 Buiuaga (Cape York). N2b3> 

I i" 14. Dorsal alio" VWllrdj view* ui the Tb^nilnunraa nt A, |J \h-yi.\7i>lotii /a,'/iur(/o. wqgjt .'?; (.', IV UttWtftixHWito ■ 
tymwJtfKilfittS Siulv >0: and F., P. U sahnini \h.\' M Stair bai - I Hn»t 



Somerset nip Cape ViKfcJ, UAZ AI774, NIMOX V. 
N2S030 uwtffvrlle 'ii^n C now, N33923 Km 

River Brt\ nr Wcipa 

OMcttfoXY (based on UAZ AI65. BI72*i |f^ 4] 
Skull Well ossified Sphenethmoid moderately 

iv I fed, in bonv contact with nasals emending fiaU 
length nt orbit in ventral view, Pmohc fused wilh 
cxoecipilal- Exoecipnai not ossified dorso- or ventro- 
medial t>. Ossification of otic region reduced ventrally. 
Crista parolica short and sleeky, not articulating laterally 

vs. nh expanded otic ramus d sqduittQte) t mntoparictaJ 
fontanel It \>ery pearly exposed. Frontoparietal well 
ossified; anterior extremities extending -A length ol orbii 
(about half length ossified portion dorsal sphenethmoid* 
Oibitai edges of frontoparietals straight, not angled 
postciolutemllv, Anterior margins of fmnAO^licia] 
lontanelle rbnned by sphenethmoid about halfway along 
length of nrbti Kisteriof margin masked by juxtaposition 

ol medial margins ot frontoparietals- Nasab moderately 
vsell ossified; crescent ie anteriorly, overlying 
sphenethmoid along posteromedial extremities and with 
well developed broad maxillary process in long contact 
with well-developed prtorbital process of pars facialis 
of maxilla. Palatines hroad, complete, running bent-nli 
dcntigcrous. processes of vomers and overlying ventral 
sptuueihinoid medially. Rirasphcnokl robust. Ciiltrifbrm 
process urn shaped, modeiatclv broad extending about 
% length of orbit. Alae broad, expanded slightlv 
laterally and angled slightly posteiolaterally 

Ptctvgold robust. Anterior ramus long, in short 
contact with moderately well-developed pterygoid 
process of palaial shelf of maxUlii. Medial ramus 
moOerately long, subaeumma'e, not overlying alac of 
parasphenoid. Posterior ramus extremely lobusl, 
Junction of duvc rami extremely robust. 

wiwdraiojugal robust and entire. Squamosals 
modciulcly robuM wilh uiodeialcly long /sgoina'u 
ramus and short, expanded otic ramus. Maxilla and 
pivmaxilla dentate, Pars facialis of maxilla deep with 
well-developed picoibitol process 

Alaiy pmcess ol pivmuxillu broad, bifurcate 
dorsally. directed posteriorly. Posterioi proecs-. <>l 
hi r maxilla moderately long, straight along 
antciomedial edges, then curved posicromediallv. 
Palatal shelf well developed, with moderately well 
developed pterygoid process. Vomers redueed medially 
with exfreuielv \QQg dentigerous processes. Columella 
caved and bony. 

Pectoral girdle arciferul and robust. Slender bony 
iini.tsTVrnum wad stalked knobbed cartilaginous 
aulciior extension, Broad MpJnslerncmi Sternum 
cartil.tcinous, Clavicles slender, curved, closely applied 
1 1 1. d lally Coracoids robust , moderately separated 
medially. Scapula bicapiiate. robust, SuptascapuJa 
about : '\ ossified, Anleroprmiiiml eiesi of humerus 
moderately well developed. 

Carpus of five elements. Pans of sesamoids at 
lunetions of metacarpals ana proximal phalanges and 
other luncfion. 1 . beiwecn phalanges except terminal ones, 
Lateral flange on first metacarpal (Pig. 5). 

Seven precocious non imbricate presacral vertebrae 
Vertebrae I and II fused, Relative widths of transverse 


Sacral diapophyses poorly expanded. Ilia extend 
slightly .interiorly to saeial diapophyses, L'rostylc cgftSI 
approximately half length of urostylc. 

Slender ihal crest. Dorsal piwnineuce prominent (Pig. 
6)_ Dorsal protuberance lateral. Three tarsal elements 
Irl tool. Prehallux large, wedge-shaped with bony biov 
and narrow dorsal cartilaginous edge 

Hyoid plate wider than long. Posterior processes 
broad and truncated Anterior processes slendei. 
expanded and truncated distally Short anteromedial 
pn*cesses of anterior hvale. Posterior eornua ossified 
(Fig. 7). 
lit nut nw 

Variation exists in the degree ot ossification ot the 
sphenethmoid which can be slightly greater or less than 
tllustraied. In one specimen (UAZ BI7281. a large male, 
the posteromedial pnicess of the uasal is more extensive. 
overlying the sphenethmoid and almost reaching th« 
anterior extremities ol the frontoparietals. 

The length of the deiiugemus pmeesxes of die vomeis 
may be slightly longer than those illustrated 
Mitlvruii examined 

UAZ A165 no data. AJ33 sex unknown Kununurra. 
B1728 male MclviLle hland. HI738 male Jabiru B17.W 
male Jabtru. B1230 stage 34 larva. B1504 swge 30 laiva. 
BJ773 Radon Creek Rd nr Mt Broekman, A1774 
ftiwnsville Town Couunon, 

Lmmudxnastt'S lomcxtusatlus calls from smaJI 
hollows, cub holes, among long grass and dense 
vegclalion in grassland, culverts and swamps <Tylere' 
ql 1VS3: Davies_ Martin fyler and Watson unpubl 

U/i> nnrtHt'Ht Oil! 

The advertisement call of/., iwivcxi tm'itlus is a well- 
tuned, single note (mean duration 74.7 ins. Tabic fl 
repcatOil regularly in long cjllin.e-scquenccs. Call 
rcpetilinn rate lor the six individuals reported here was 
2 1-4 (range 1 5°~2.57) caJIs/s (at an effective temperature 
of 2K.0°CJ. Waveform and speetrographic displays oi 
a single note and a power spectrum are shown in Pig. 15 

ti'tuujdxtutstes ty»>\VAt't\( uhi.s has u dixtribution 
across the north "I Australia in the wet/dry tropics and 
including southern New Guinea (Pig. 9). The species 
\> cunbnt-d Lottie Kimbcrtey a*gion in Western Australia 
and occurs as fat south as 34 km S Delamea* 
Highway/ Victoria Kighway.)cn in the Northern Territory 
and F.urimbulah N.P. near Mil lamvale in Queensland. 



Life history 

Hggs are pigmented and laid in a foamy egg mass. 
Tyler ei ai. (N83) described the tadpoles of L 
convt'AiuscuIus 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 disc is 
surrounded by a single row of labial papillae which 
are interrupted anteriorly {Fig. 13). The lull life history 
of this species has not been described. 
Chandrpcnmium (stage 33 larva, based on UAZ BI53I) 
ilig. 14) 

Neurocranium approximates T-shaped box 
comprising anterior braincasc and posterolateral 
spherical otic capsules. Large ovoid frontoparietal 
fontanelle exposes braincase dorsally, bounded by 
orbital cartilages laterally and by tectum synotieum 
posteriorly. Narrow ethmoid plate forming anterior 
margin of frontoparietal fontanelle. Bthmoid plate 
projecting anteriorly from neurocranium as two 
antcrolaterally diverging, moderately broad, finger-like 

cornua trabeculae. Floor of ncurocramium comprising 
basis cranii and planum basalc. Basis cranii perforated 
by paired foramina carotieum primarum Posteriorly 
planum becoming confluent with otoccipilal arch and 
perforated by notochord, 

Massive approximately L-shapcd palatoquadrale. 
Processus descendens (attachment to neurocraniuml 
widening laterally to form areas subocularis, separated 
from basis cranii by oval suboeular fenestra. Medial 
processus ascendens slightly overlain dorsally by 
anterior extremities of otic capsules. Areas subocularis 
curving laterally and sloping ventrally. Anterolateral 
edges bearing low crest, becoming progressively higher 
and confluent with broad processus muscularis 
quadrati. Areus subocularis attached anteriorly (o 
neurocranium by commissura quadratocranialis 
anterior Finger- 1 ike projections extending 
vcntromedially into suboeular foramen 

Processus muscularis quadrati inclined medially, 
bound to lamina orbitonasalis by nonchondrificd 
Hgamentum (eartilago) lectuni. tonning tunnel with 






i r 


l r 




4 - 

TIME (ms) 





TIME (ms) 

fig. 15, Powei spectrum, waveform and sonagram (sampling frequency 59 H/) oJ the advertisement call of t.mmaJymisws 
c&tyexlusculus. Note thai the ordinaic litr the waveform display is not labelled because ii depicts a relative linear scale in volts. 



commissure quadrato cranial is anterior through which 
pass M levator ujandihulae posterior ami M.l.m. 

Processus arucularis quadrati of pulatoqttadrale 
extending anlenoi Iv ftOftl processus muscularis quadrati 
and articulating veiitrolaterally with slightly curved 
Meckel's cartilage. 

Ligamenlum quadrati ethmoidale attaching laterally 
midway along eorrtua trtibeculae. Dorsomedial edges 
of Meckel's cartilages attaching with int'rarostra! 
cartilages. Infruroxtral cartilages small. Ionium; shallow 
inverted U in ventral view with broad non-chondr tiled 
symphysis, elements joining to form lower jaw ol 
lad pole. 

Supran>stral cartilages feoiung upper jaw of tadpole 
projecting anteriorly venlral to cornua liabceulac and 
dorsal 60 bcrtfi Meckel's cartilage and mlramstnils 
Central corpus ol suprarosiral tytfig umet tor to 
cxiietnities of coruua trabcculae giving appearance Of 
pain** I structure with non-chonclnlied medial symphysis. 
Lateral alae extending to level of Meckel's cartilage 

Megistototis lignarius 

Tyler, Martin & Davies, 197 u 

FIGS 1-7. 9. (4, 16 

Mt r i;i\it>ft>us lifiruttrits Tyler. M- J- Martin, A fc- & Navies. 
M 1,107V! Aitsr. I jhfit 27 135-150. 


WAM R5H2W Lake Argyle -Kununurra Rd. 6.5 Km 
N Labi Argyle Tounsi Village. Lake Argyle. Kimherley 
Division, W. A. 

A moderate-sized species (males 43-62 mm S V 
Icmales 47-61 mm S-Vj characterised hy q large and 
pnwninent tympanum (apptoximalely the si/e of the eye* 
trig. It. male nuptial excrescences consisting ol' small 
clusters of latee conical black spines, Umiaks with 
moderately developed flange on (he medial side ol the 
second finger, first finger longer than .second, 
advertisement call consisting; of a soft regularly-repeated 
"lap", eggs deposited in a foam nest 

We have little to adit It) the comprehensive description 
o1 l\|ci c/ ut (197^1. However in five females retired 
ftprjfl the original spawn clump we recorded moderately 
develop**.! flanges on the medial .surface of the second 
linger, a ehantctcr absent from the paratype materia' 
iFig 3), 
Mrcanfiw (based on LAZ A 1742) (Fig. 4i 

Skull moderately well ossified, in bony contact with 
nasals dnd extending hall length of orbit in ventral view. 
PppOtk fused with exoecipitaf Pxoeeipiial nol tuscd 
dorso or venlro laterally Ossification ol pn»ottc reduced 
ventrolaterally. Crista paroliea moderalely long and 
slender, nol articulating laterally with short, very poorly- 
expanded otic ramus of squamosal r-iontopancial 
fontanel le moderalely to well exposed expending abpui 

half lenglh ol orbit anlenorly, Orbital edges of Ironlo 
parictal.s straighl . nol angled [josterolatetally Anterior 
margins ol frontoparietal lontanelle formed by spheneth 
moid about hallway along length ol orbit, Posterior 
margin undefined- Nasals >lender_ pooily ossified. 
Maxillary process acuminale. widely separated Irom 
miKJc*rately-de\'elopcd preorbital process of pars facials 
of maxilla. Palatines broad, not expanded laterally over- 
lain about hallway along length by lateral 'i ol 
denligerous process of vomer. Parasphenoid robust. 
Cultriform process moderalely bmad. alac ran quite, nut expanded laterally. 

Pltrygoid robusl, not in conlael with |xioiiy-deveioped 
pterygoid process of maxilla. Medial ramus long, 
moderately slender, acuminate: posterior ramus 
moderately stocky j hi mes expanded at junction of three 

Qnadratojugal robust and complete. Squamosal 
moderately robust with short slender zygomatic ramus 
and Short Slightly expanded otic ramus Maxilla and pre 
maxilla dentate, teeth extending past pterygoid process 
of palatal shelf, Pars facialis of maxilla moderately deep 
wilh moderately well developed preorbital process 

Alary processes ol preihaxilla broad al base, slender 
laterally, directed posteriorly. Palatine pnvesscs widely 
sepaialed. slightly angled postetornedmlly. Palalal shell 
poorly developed. Vomers entire, widely .separated 
medially. Denligerous process elongate, cre.sceniu 
behind ehoauae, Columella bony, cuived. 

Pectoral girdle arciteral and mbust. Well-developed 
omoslcrmim and xiphisicrnum. Clavicles poorly 
separated medially Coracoids widely separated 
medially. Suprascapula ty ossified. Well developed 
humeral crest. 

Phalangeal formula of hand 2.2.3,3. Carpatia appear 
Itl be fused with Os centrale posraxiale Very bony pre- 
pollex with spine antcrorncdially. Metacarpal I elongate 
with mediolateral flange (Fie. 5). Pairs o\ -.esamokls 
at junctions of metacarpals ami proximal phalanges and 
proximal phalanges and next phalanges. 

Seven procoelous mm- imbricate presacral vertebrae. 
Relative widths ul transverse processes: 


f'rostyle crest extending -•_. lenglh of clement. Ilium 
with slum slender crcsr. Prominent .dorsal prominence 
(Fig. bi. posterolateral protuberance. Phalangeal formula 
of foot 2.2,3,4,3. Two tarsal elements. Terminal 
phalanges knobbed. 

Uyoid plale wider than long. Posterior pnx-essc-. 
slender and rounded distally. Anterior pnvevscs slendt'r 
slightly expanded distutly Amerornedial pna. esses crt 
aulci lor hyalc expanded and broadly truncated distally 
H ig. 1). Pisterior cornua short and ossified 

Varialion occurs in the ossification of life* 
sphenerhmoid mul fosia pannna in the matcri-il 
m icd Maximal ossification of Kith these elements 



is as illustrated although slight ossification of the nasal 
capsules was noted in UAZ A174I, In this specimen 
also, the gap between the extremities of the dorsal crista 
parotic a and the otic ramus of the squamosal was 
slightly less than shown. 
Material examined 

Northern Territory: UAZ A228 Cannon HiJl, A58S 
male, B1236 larvae stage 27, B1237 larvae stage 37 

Western Australia: UAZ A248. A1768 sub adult. 
AI769, A398-400 males. B428 B17771-72 females 
laboratory reared from spawn from tvpe locality. 
B1741Q, B1742 males no data. 

Wc have encountered a number of M. ligtiarius since 
the species was described and all have been true to 
the habitat description of Tyler et aL (1979). 

The species is confined to the escarpment country 
of the Kimberley and northern Northern Territory as 
defined by Tyler et aL (1979). 





20 - 

Advertisement call 

The advertisement call of M. liqtiarius is a relatively 
poorly-tuned, single note (mean duration 26.6 wis, 
Table 1) repeated regularly in long calling sequences. 
Call repetition rale tor 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 history 

Tyler et at, (1979) described the complete life history 
of this species. Tadpoles are lotic with mouthparts 
adapted to last llowing water and are characteristically 
heavily pigmented. They actively select areas of 
streams in which water is last-flowing over riffles and 
avoid the calmer pond areas (Davies. Watson and 
Williams unpubl. data). There are live 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 
disc and are interrupted anteromedially. 


TIME (ms) 







TIME (ms) 

Fig. lfi. Power spectrum, waveform and sonogram (sampling frequency 59 H/» of the advertisement call of Mvgistohtis 
(ignarius. Note that the ordinate for the waveform display is noi labelled because it depicts a relative linear scale in volts. 



Chomi roc ran turn (Stage 27 larva, based on UAZ 
B1236) (Fig, 14) 

Neurocranium approximates T-shaped box 
comprising anterior brainease and posterolateral 
spherical otic capsules. Large ovoid frontoparietal 
fontanellc exposes brainease dorsally. bounded laterally 
by Orbital cartilages and posteriorly by tectum 
synolicum. Narrow ethmoid plate forming anterior 
border of frontoparietal fontanelle. Ethmoid plate 
projecting anteriorly from neurocranium as two 
elongate anterolaterally diverging, finger-like, cornua 

floor oi neurocranium comprises basis cranii and 
planum basale. Basis cranii perforated by paired 
foramina carot'tcum primarum. Planum becomes 
confluent posteriorly with otoccipital arch and is 
perforated by notochord. 

Massive approximately L shaped palatoquadrate. 
Slender processus ascendens (attaching palaloquadralc 
to neurocranium) widening laterally to form arcus 
subocularis. separated from basis cranii by oval 
subocularis fenestra. Medial processus ascendens 

extensively overlain dorsally b> anterior extremities of 
otic capsules. Arcus subocularis curving laterally while 
sloping venlrallv. Anterolateral edge bearing low crest. 
becoming progressively higher and confluent with 
broad processus muscularis quadrati. Arcus 
subocularis attached anteriorly lo neurocranium by 
commissuta quadratoeranialis anterior No finger-like 
projections of element extending ventromedial^ into 
subocular foramen. 

Processus muscularis quadrali inclined medially, 
bound to lamina orhitonasalis by nonchondrified 
ligamenlum (cartilagoj tectum, lorming tunnel with 
commissure! quadratocranialis anterior through which 
pass M. levator mandibulac posterior and M.l.m. 

Processus articularis quadrat] of palatoquadrate 
extends anteriorly from processus muscularis quadrati 
and articulates ventrolateral^ with slightly curved 
Meckel's cartilage. Anterolaterally. ligamenlum 
quadrati lethmotdale attaches laterally to cornua 
trabecule about one third along its length. 

I '\m.r 3. Variation in morphological, osteolotficaf. developmental and behavioural characteristics »j/Ximnodynasies salmini, 
L. eonvexiuseulus and Megistololis lignarius Karvotxpic data are from Mahonv <& Robinson (J986). 


t sahniw 

JL nmvcxiuseulus 

A/, liviuirius 


Nuptial pad. spiny 

Spines on skin 

Glandular dorsum 


Planned fingers in female 

Relative finger lengths 

Karyotype (2ni 

NOR. (chromosome number) 


Nasal* appro*, iriangular 

Frontoparietal I'onlanellc exposure 

Double tooth row on vomers 

Rel, ol pars facialis with maxillary 

process of nasals 

Robust icn oi pterygoid rami 

Ret. anterior nanus ol pterygoid 

and maxilla 

Mod. slender cultrilorm process 

Hastate piefhillu* 

Carpal ia 

Double sesamoids on fingers 


Highly melaiiic larvae 

Heavily pigmented eggs 


Larval mouth 

Cornua irabeeubie extending over 

mouth cartilages 

Calling site 
Well tuned call 
Riflle-seekint; larvae 










mod. distinct 


very disLinel 


well developed 


3>!>2 4 

3>2> 1-4 








































2,4/4: 3 

Roeky stream 



DorsomcdiaJ edge of each Meckel's cartilage attaches 
with infrarostral cartilages which form shallow U in 
ventral view with conjoined broad, non-chondnfied 
symphysis. Cartilages form lower jaws of tadpoles. 

Supramstral cartilage lorm upper jaw and project 
venirally to level of anterior extremities o\' cornua 
Irabeeulae, posteriorly do not reach level of Meckel's 
cartilages dorsally. Central corpus of suprarosttai lies 
between extremities of cornua trabcculac: appears lo 
be paired. namrw medial non -ehondrified symphysis. 

Lateral alae just e sending to level u>t Meckel's 
cartilage laterally. 


Table 3 consolidates those ehaiacters shO-Wirtg 
variation between the three taxa. Llowever, this is a 
phenelic comparison and as such does not clarify the 
relationships between the laxa. which await a cladistic 
study of the genus in Uttt) with appropriate outgr<>ups 
(Dav tcs in prep.). 

lulormation on call characteristics generally have not 
been included in phylogenetic analyses, although ihese 
data ate ol particular interest. The advertisement calls 
of all three species described here appear to be of 
sinnkii harmonic structure (figs 8. 15, 16). II is nol 
clear how calling frogs produce such (requeues speura. 
although most interpretations .suggest thai (lie \v>eal 
coids generate a carrier frequency which is mod Tie-* I 
by secondare pulsing (amplitude modulation) that 
results in a pattern of side bands of (he dominant 
frequency (Walkins 1967; Ryan 1988). Alternatively. 
males may produce a lundamental frequency with 
associated harmonies, with one or more of fcsc 
harmonic frequencies being emphasised by the 
resonance characteristics ot the body cavities. In either 
case, well-timed calls produce a distinctive spectral 
structure, and this pattern is particularly clear in the 
calls t>\' I. cntiirsmscufux and L. sulmini, where ihe 
fotirA frequency band contains most energy (Table 4, 
ligs K, 15). Despite the overall structural similarity ot 
the calls pf 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 1988/ and. in tlus case, are 
consistent with the overall trend in male body size in 
the species with L Htlmi/ti being generally larger than 
L, convexiusenhts. 

fhe mode of production ot ihe advertisement call 
of M. ligUCtMUS is equivocal. While sharing basic 
spectral similarities with the calls of the other species, 
the call is less well- tuned and trie envelope shape is 
markedly different, having a rapid iise-tirne that is 
likely to generate a transient wide spread of frequencies 
(Rossing 1982/ and to obscure the harmonic frequency 
spectrum (Table 4. Fig. 10). There is also a cleai 
difference in the call characteristics of the two recorded 
individuals (Table 4}. Individual #1 (SAM R16229) was 
calling in a rock crevice in a stream that, based on 
many subsequent observations of this species in (he 
field (M. J. Tyler. M. Davics & G. F. Watson unpubf ). 
is u typical calling position. In contrast the call of 
individual #2. the holotype CWAM R58229), was 
recorded while the male was moving in ihe open, a 
situation that has never again been encountered by us. 
Because of the behaviour of the holotype at Ihe time 
when its call was recorded ii is more likely that the 
i all of individual #1 represents a typical call of this 

Major developmental differences between the three 
taxa tic in the iotic nature of the larvae of ' M. Iii>tit}ritts 
as opposed to the lentic larvae ot £ cmvcxhtxcidUi 
and t, sulmini. Lohc adaptations in M. Ii\>}mri}(\ 
include a flattened body, strong tail musculature with 
relatively narrow lad tins and a relatively large ventral 
suctorial mouth disc. This is of a less elaborate nature 
than found m some lotie species (Davics 1989a) 

Time to metamorphosis in V7. h&uiritis is longer than 
in L, snf nt'm i {nox known in L. K"mx\\iitscultts) being 
about 65 days as opposed In $3L 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 amirs. 

T\HI t 4. invrihutuiit of Jnym'fUhs tHu \>UhiK the mils of "Limnodynask-s salmim. L. convcxiusculus tiir/J Megistnlnds 
ligrffirtUK, Vw fftdiMttCy with Most vtt»-t$\ >ttit dominant jhyiwn<\) i$ .urulrrlinrd. 

L. sutmini 

/ rt'nvtixitKSVUltts 

M. Iii»niirim 











































1 %w 




















M- IMVII-S g 11 l' WA'ISON 

N*>ik- of the taxa appear *f> have exceptional adaptations 

iii i;tpu! larval development, again reftetfine w« k ' NS 

ephemeral nature of Ihc water bodies in whiCTi they 

Tht* lulif nature olthe SfytyTtf&lOik knvae |$ also 
tetlcaed in ihc chondrocnmiurn The leneth Cfl the 
eornua IrahfittJfeie anil their relaliotiship with ihc rfotitrul 

cartilages tl i Ff«j rs In these lotie and lenlic species. I In* 
annua trabecule alt elongate in lotie M, liputrtus 
and relatively shorter in lenlie /.. utlnurtt and L 
it >nvi'\:'nt.\citlti t \ This observation does not lit the 
generality of Sokol (1981) for lotie sp&Cyfes of othei 
luiiiilies 1)1 Which shortening of the eornua trabecule 
is correlated with's of the MOtltfl a way from j 
viihieiininal position iri lotie larvae, 

descriptions ol choiidnxrama of limnodynaslinL 
lava viie unavailable frf comparison, the only published 
dala on Australian species being tliOSC ftOTft the- 
myohalruLhinr species Ufit'rnlcm lillumunln (Davie 
titffJh) and two species iff PsetuJopfitvru' (.hn onsen 
LQft&t. none r)f U/hfch feu 'otic species 

Ossification of ihe skull is negatively convlaied Willi 
sire- The large /.. salmmi has die poorest ossirimtion 
and die sniatlcsi species L canvcMuuvltis has Ihe 
fjicjtest. Intuitively one would expect largei species 
to have >lmnger, more ossified skull 1 -' than smaller one L 
However, the aduli skeleton is Ihe producl of both 
mitogenetic and phyloscnciic constraints (hueb & 
\ibcrch J9S5: DaVJCS I98*)b; Hanken I^Oi and .such 
a simple correlahon rarely holds. 

I .ynch ( Ipfl 1 provided a diagnosis fal UnittndvnaMrs 
based on examination ol the skulls ol L dorsalis. L 
lUtchcn, L. fM-ronf and L ta\tttcm!c»\^ Data from 
ihe three taxa investigated here do not coufoim ItrttriS 
diagnosis in a number o\ features The nasals arc very 
anal! in A/cc/.s/o/m/v rather ihan being relatively small 
although I, SdltylW Jnd ft toinexiiiuttius Hi the 
diagnosis. The nasals are not Jfl broad eonui.t wjtfi the 
rnox iliac in Mr^isntlotts and the frontoparietal 
lonianelle is pbfil ly exposed in both /- sahnint and /_ 
a>n\r-\iu,\twtt<> as opposed to thfl moderate exposure 
defined Ibr the genus. Lynch describes the /ygomatk 
Kiinii- as being as long as the otic ramus in 
J.itntiodxnuMc* bill the former is longer than the latier 
in both I -'dlmmi and L crmwxmsathis. The palatines 
arc noi narrowly separated medially m Mcyivrototix 
and the sphcnerhmoid does not extend unlet Ull ly ED the 
centre- o\' the nasals in / . \<ilnnnt. The alae ot the 
parasphenoid are not broadly overlapped laterally by 
ihc median rami OftflO pterygoids in any ol (be three 
iaxa .md die anterior ramus of ihe pieiygoid dOc$ noi 
conform Pp the generic deseriplion m either 

Parker (I'MOt discussed (he. relative lengths of 
metacarpal I in tfiWIrtrf.VWW mul reported a trend 
io^juJ- reduction til 'he phalanges ollhe first finger. 

Internally, Ihe metacarpal bone is disproportionately 
long The irend was noticed in L jlvnhrri f- pirnta 
<.ilMl lo a lesser extent in /.. xahmni Parker sugyesicd 
that il is significant that these species do not have 
nuptial excrescences, He went on to argue thai species 
thai spawn in water, such as Lintfwtlvttttsics. need 
nuptial cm. ivseencLs io jid m clasping the female and 
lhat lack of Ihe same must be compensated tor in some 
way, i.e., by changes in the relative length o\ the 
finders. The same trend is: ivcordcd in A/. Ii^nnhus 
hut this species has well-developed spinous nuptial 
escrescericcs, mi does no| support 'he assurilpiion. 
I urlher. we rceord lierc ihc presence of glaiidulaj 
nuptiul excrescences m /.. submnt contrary lo the 
observations of Parker (JsMO). 

Parker M940) provided X-rays ol Ihe hands i^ /. 
ta\tn<wtt'ttsi.\> L ptvnm, L fletcheri and /.. sahnnu 
to illustrate the nature o\' the metacarpals and digils 
The man-rial in this >»udy for which comparable daia 
arc-available ate all males and all have ;i lateral flange 
on Ihe medial surface of the' first metacarpal 
t.xainination of Parkers material shows (he presence 
i)\ ihi> structure onty in male /.. pcront but nol IN 
female i MlnUtltWU} mtlMe t, flwhert Wheiher this 
i-, a sexually dimorpliK character remains in doubt as 
ihc \-rays are exlremely granular and may not be of 
sulTiuiera quality to detect the slructure. 

Limnoilyntisti's suhnitn and /,. ct>m{>xitt\cttlu,s shafc 
similar habitats and. except for the difference in 
dominant Irequeney, have very similar calls. Ihe 
distributions of these two species have a /one of overlap 
between Miriamvale and St Lawrence in Oueemtand 
Identification by way v\ call from these areas must 
therefore he regarded as dubious when compiling 
distribution data. 

The data recorded here conlnbule lo our knowledge 
ol the morphology and reproductive biology ot ihrec 
Ausiralopapuau anuran.s and also will contribute to a 
wider analysis ollhe phylo^entic relationships, within 
Ijnmoiixnusles b'it/inger. 

AeKuow Icdgments 

This study was supported by an Australian Research 
Grants Council Grain to the first author h\m\ Michael 
J. Tyler. We have also drawn upon material eotleued 
together wiih our colleagues Angus A. Martin and 
Mitphacl f. Tyler whose support and company wv 
gralelully acknowlfdee. I. eeunnc Seller and Michael 
J. Tvlcr assisted with tadpole rearing. Keith MeDonaU 
provided distributional data for Queensland. Michael 
lylet c riticatly read the nianuseripl anil we lhank htm 
for his comments. We thank Angus A.. Martin toi 
allowing the analysis of original recordings o! the taxa 



(_*H,(ii.M. H. G. (1992) *Rcpliteii and Amphibians ol 
Australia" 3rd cdtn (Reed. Sydney-). 

_, 0\Ml-,K"V K. F. & CotiUtk. H. M. (B&lj 
/ooiogKal Catalogue oX Australia. V*i| !. Amphibia and 

Rcptilia" t Australian Government Publishing Service, 

Davii.s. M. 1 1089a) Descriptions of the tadpoles of HHTIC 

Auslraliun limnodvnasline leptotlactvlkl frogs. Trans. K 

Sot, S. Aust. 115/6776. 

_ (1989b) Ontogeny oi' bone and the role ol 

heterochrony in the myobatrachine genera Upvtokut, Cnn>a 

ami Pseud flphrvrte I Anum: Leptodaciylidae: 

M\ohatrachinae). I Morphoi, 200. 269-30Q 
Dam's. D. D. & Guri, V. R. (1947) Clearing and staining 

small vertebrates. Ftvldumn lechtftquts 4. 1-16. 
DinqHRKUS. G & I-'mixh. L D. U977) hn/yme bearing wl 

aleian blue slatned small whole vertebrates lor 

demonstration ol' cartilage. Statu Tt\hm>l. 5i, 229-232, 
GOSNEfU K. L. (1960) A simplified tabic tor staging anuran 

embryos and larvae with notes on identification, 

ttitpehiloy.un 16, 183-190. 
Ha Mis UN, £ (IWZl Adaptation of bone growth to 

miniaturization of body size, pp. 70-104 /// Hall, B K (Bdj 

'"Bone" Vol. 7 Bone Growth- B. (CRC Press. Boca Ralon. 

Hi rutlNSON. M. N. & Ma\s«jn. L. R. (1986) Albumin 

evolution, paraphyly and the relationships of the Australian 

m»g Mei>t.\u>hnis hummus (Anura- LimntHlvnastmae) Sjtff, 

zW. 36, 387-414 
J a< *.usi sj, C M. (I968i The developmeni ol the 

chondtocranium in two species of the Australian anuran 

genus Emulophrxnv Fil/ingcr. Aust / S'V? 16. I IS. 

I.IMI IJilUN. M. 1. ROHKKIS. J. I)., WATSON, G. F. & 

nwii-s. M. (1003) Chapter 7. Family Myobairaehidac. pp 
41-57 fn Ross. Ci. A (Ed.) "Fauna ol Australia. Vol 2A 
Amphibia & RcptiTu" (Aust Govt Publ Service. 

Lynch, J. D. ifO?D Evolutionary relationships, o.steoloyy 
and ?oogeographv o! leptodaetyloid frogs. Vtiiw K>nn<t.s 
Mus AW- Hist, Ocuis ftthl No. 53. 1-238. 

MaO&Ny; M- J. & RimiN.snN, R- S- tl9K6i Nikleolar 
organiser region (NOR) in karyotypes ol Australian g'tuuul 
frogs (Family Myobairacliidiic). (itnttuu 68. 110-127. 

Moow J. A (1961 1 The Irons ol eastern New South Wale* 
Butt 4m .A/ma iV«/ Hist' 121(3), 155-385. Plates 27 45 

PAkKKR, W M (1040) The Australasian frogs ol the family 
Leptodactvlidae, Mm'/- Zn>loi>teac 42. 1-106. 

Rrmi-Ris\ L D. & Maxmjn, L. R. 11086) Phyiogeiletic 
relationships of the genus linnuuiynustf,\ (Anura; 
Mvobatracfudae): u molecular perspective. Aust. J, Ztml 
34, 561 573. 

m & Waison. G, E (1903) Chapter 6. Biogeography 

and phytogeny of the Anurti. pp. 35 40 In Ross. G. A 
I Ed.) "Fauna ol Australia Vol. 2A. Amphibia & ReptHia. 1 * 
i Aust, Govt Publ. Service. Canberra). 35-40, 

R.jsmm., T. D. U082) "The science of sound" tAddison- 
Wesley Publishing Co., Reading Massachusetts}.. 

Ryan, M J. (I98K) Constraints and patterns in the evolution 
ol anuran acoustic communication, pp. 637-677 fu Pntyscti. 
B.. Ryan. M J . Wileyski. W, Hetherington. T, H. & 
Walkowiak \N. (Eds) "The evolution of Ihe amphibian 
auditory system." (John Wiley & Sons. New York). 

SThm.ipi. K. H. (10K8) Peripheral and central meeahnisms in 
vocalisation, pp, 537-558 In Fril/sch. B.. Ry;in. M. J., 
Wileyski. W.. Hetherington. T. E. & WalWowiak. W. ;P.ds) 
"The evolution of the amphibian auditory system." (John 
Wiley & Sons. New York). 

Sdmh , O- M. (I0S!) The larval ihondrocranium o\ Pch'ihlcs 
putntants, with a review of tadpole chondrocrania. ./. 
Morphtd 169. 161 183. 

T«1 Ufi, L. & AtBtM h, R (10X5) Mmiaturi/alion and the 
anui'un skull: aeasestudv of hcicrochronv pp. 113-121 In 
Dvmcker. H. R. .fc Fieische. G. (Eflx) "Functional 
morphology of vertebrates" (Gustav Fischer Vfcrlru] 

hi ik. M 1. (I062i On the preservation of anuran ladpoitto 
Aust. ./. .SV/. 25. 222. 

(1002) 'Encyclopedia ol Australian animals. Frogs" 

(Annus &. Robertson, Svdn-n J 

Ckoiik. G. A. .^'DaviVs, M. (1083) Reproductive 

bioloiiv of the froes o\~ the Magela Creek system. Northern 
Territitrv Rvi. S.Aiht. Mus. 18. 415-140. 

Makiis. A. A & P-AVlfiS. \L (1070) Biology and 

systematies of a new lininodynastine genus (Anura 
Leptodactvlidae) from northwestern Australia. Aust. I. 
7.oot. 27. '135 150. 

Wairins. W A. (1067) The harmonic interval, faei Of 
artifact in spectral analysis of pulse trains, pp. 15-43 hi 
Travolga. W. N. (Ed.) "Marine bio acoustics, Voluiie 2" 
(Pergarnon Press, Oxford). 




ByD. E. Rounsevell*, D. Ziegeler*, P. B. Brown*, Margaret DAViEsf 



Rounsevell, D. R, 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 1 1 species. 
Key Words: Anura, Bryobatrachus nimbus gen. et. sp. nov., Tasmania, new genus, 
new species, morphology, osteology, advertisement call, development, habitat, 
cryptozoic, distribution. 

Tritn\' t ti(ttw\ u\ ifa kttuil .Vx'jV/v of S. Aits', (1994*. IIKMi, 171 R$5 


bv 0. E. Rounsevell* D. Zilcellr* R B. Browne Makgakm DavilsI 



Rut'wwM , D. E - ZiH.hLhR, D , Brown. P. B. . D.wifcs. M. & Litti Moi'V M .1 11994) A new genu* Qdid 
species of frogfAnura: Lcplodaclylidac: MyohairHehiliae) from southern TjMtUtt&i /r<//(s ft. ,Vo». V. *t/f\/. JlSOl. 
171 185. 30 November. 1994. 

BryvtMtmi'hu.s nimbus sen. el sp nov_ isdesoihed front mois! rddl hahiiat in southern Tasmania. The genus 
is distinguishable by iLs direct development, lusnn frf vent-brar VII *"'d VIII Wiih Ibg sacrum, dentate 
maxillary arch, absence of vomerine teeth and the presence ol a columella Tb? •■pet ie\ is small finals IW71 
lermdes 25-30 mm S-Vl with distinctive dark patterns on the douoam and an advertisement call that is B xene> 
ol'Hoks" with a pulse repetition rate of 5.J-6.1 pulses.'s\ Eggs are laid m cavities in mo*s in groups DJ 9-14 and 
hatch as tour-legged, lailed froglcls. & nimbus grn. ei sp. nov. is cryptuzoie in vegetation al poorly drained sues 
•in wet peat, or in edaphic moss in implicate raintbtest and subalpme moorland at alrfctUdci Irotn near >cu level 
la I.KK) m. The yenus and specie* are endemic to Tasmania and bring ihe fipjc fauna nf ihe island lo I! specie 

Kh\ Woros A mint. Rtvubaimttuty nimbus een. ei sp. nov,. LiMnama. new genus, new species, morphology 
osteology, advertisement call, development, habitat, cryptoyoic, distribution. 

ln I rod net ion 

The troy fauna 6f Tasmania is an element of flit 
Bassian Province o\' south-eastern Australia (LiUlcjohi'i 
ft Watson 1985), As well as species with wide ranging 
distributions throughout eastern Australia (e.g.. 
Htwuh'llu siswieru Girard. 1858 and Liwmtdvnasles 
nisnntriiensis Giinther, IH5S | Leptoclaetylidae. 
Myobutraehmae and Limuodynastinael), there are two 
endemic species, Litoriu bttnowsae (Seoli. 1942) 
(fh lidae) and RunidelLt Uismam'ensis (Giinther. 1864; 
(Lcplodactylidae; Myobatrachinae). 

Ten tana are recognised in Tasmania and aspects Of 
their biology are relatively well known (Martin & 
Littlejohn 1082). With a greater appreciation of ihe 
need lo document the biodiversity oj the continent, 
efforts have been directed toward surveys and the 
mapping o\' distribution patterns of local fauna, During 
one such project aimed at providing an alias of the I roes 
o( Tasmania, one of us (D.Z.) recalled having neard 
at several high altitude sites, a distinctive frog call that 
could not be attributed lo any described Tasm.mian 

In late November 1992 calls were recorded at the 
Had/ Mountain National Park and specimens were 
collected (Robertson IW; Rnunsevell &. Swain 1993 > 

* Parks and Wildlife Serviee. Department of Environment 
and Land Management. GPO Box44A. Hobart, las. 7001, 

+ Department oF/oologv, University ol Adelaide S Ausi 

4 Department of /oology. Universilv of .Melbourne. 
Parkville, Vie, 3052 

1 ANPfckiitN, M. L, 097K> The eomparuuve myology im] 
osteology nf the carpus jhJ tarsus ol selected anurans. PhD 
DisM-Ttafion, Dept ol Sy^lt'inalics and Ecology. DniVi-i'Miy 
of Kansas (I'npubl.). 

P.vaminalion of the collection at ihe Tasmainan 
Museum and Ait Gallery. Hobart. revealed further 

The rfOgS could not be referred to any known ^enus. 
and so a new genus is erected W accommodate rhem 
here, We describe Ihe species and provide information 
on its bioloyy and distribution 

Materials and Methods 

Material reported here is deposited in the Tasmanian 
Museum and Art Gallery. Hobart (TMAG), South 
Australian Museum. Adelaide tSAMi, Museum of 
Victoria, Melbourne (NMVh Lhe Australian Museum. 
Sydney (AM > and the Department of Zoology. 
University of Adelaide (UAZ>. 

Measurements were taken usine dial calipers reading 
to 0.05 Tiiol- or with an eye-piece micrometer 
Vleahiirernenls (mint were, cyediamelcr iL): eye-to- 
naris distance (I*, N). internarial span ilN'j: snoul-veni 
length (S-V) and ribia lctuth (I L). and for selected 
specimens, head lenetb (HL) and head width (HW)> 
The methods ufnieasoreiiKnl follow Tyler (I96S) and 
interpretation ol data follows Tyler U97N). Uata au 
presented as means with ranges m pareniheses. The 
tympana ol many specimens are indistinct or noi 
visible, and hence accurace measurements oi head 
width ami head length were not always possible 

Osleologieal data were obtained from .specimerK 
cleared and stained with alizarin Red S for bone aftet 
the method of Davis & Gore (1947b and with ali/ann 
Red S for bone and aleian ^lue lor cartilaee after the 
method of Dingcrkus & Uhler (I977L Osleologieal 
descriptions follow Trueb (1979), and Andetsen 
(197S 1 ) for the carpus and tarsus. 


> l KlUiNSKVrU, ft /li Ql ! I k. P H IIKOWN, M DAVthS & M. .1. LI 171 fJOUN 

I me drawings wetc made with the aid ofa Wild M8 
sletcoseopic dissecting microscope with attached 
camera luuda 

Most Hold ohsei vutions were made in (he spring ami 
Mimrnt; ol 1993, at the type baility Wl Hail/ Mountains I'itk. and during a Held survey in Ocioher 
and November 1993 (Xiegeler 1 994/. Au temperature 
and relative humidity were measured with a "Zeal ' 

h\fl\tig psyctironietcr (&$ 2X42/66) 

Rcvordiugs nj the advertisement calls nf une 
iQilivfdunl wcte made with a Sony WVLLm Pro- 
Walk nun cassette i t1 p,. reeonlet and a Btytf foMJB 
vardtonl dynamic microphone by G, F Watson til the* 
t\pe locality on 5 D© laiibci 1992 at about I330 hours 
L ST. The wet -bulb air temperature was 8.5°C and 
a dt>-hulb air-temperature was' 9.2 Q C in the \ieimiv 
of Ihc calling mate. 

Recordings ol advertisement calls of several 
individual* al.WJ were made at the type locality hy 
j* ft Brown on 30 November 1992 U5tng a Ma»aoi/ 
Supei scope C-205 cassette recorder and a Sennheisei 
Ml, ttf) clccirel microphone: temperatures wen- nifl 

I he calls wete replayed on a Nakamichi Dragon 
easseMe recorder . with the line output directed to a ka\ 

i l -1,-k-s DSP-5500 digital Soua-Graph 0A\n of 

suflie<cnl intensity thai did not overlap those of 
neighbours were analysed. The number of pulses in 
a call (determined hy inspection), jind measuteiiicMs 
Of vail duiatiim mm and pulse repetition rale las 
tmIscvs, Irom Ihc peak of (he first pulse to the peak 
ol the last pulse) were determined imm the wave toim 
display ol each clear call- 
Tor the si*; clear calls obtained by G F Watson, 

d< u.-nt |rct|ueucae.-. (H/) were determined from the 

power spectrum for the complete call, with the 
oi,i\imum and ucm highest peaks hcing measured 
Pulse durariou, and alack and decay times of tlv 
middle pulses ,dso wefe estimated io the nearest 
millisecond (ms) from the waVe-form displays, The 
repetition rate of the calls lutf calls, nun) also was 
duennmed from litis sequence. 

I i^'liuvii dills bom the tccording sequence ohtamed 
In P. B. Brown were analysed, and duration, number 
of pulses jihI pulse rate determined. As calls of several 
individuals are included, the values are treated .is a 


Lout ami ran families are native to Australia H\ lidae, 
Ramdac MiomIk lut.n: mid Leptodjciyl idae 
<M>ob:nr.ujhidac ot m-.ns authors;. The new frog was 

' Owns M ilS»M7» Taxonomy and s\Mcrn:iiivs Ui iRt! acrfO l 
i'fh'tnhw <St av (Altura Leptuduclytidae). PhD Ttu:.-,».- 
Ut'pariment pi Zoology. University rtf Adut-ink- tinpoM 

identified as belonging to the family LeptodaetylitJae. 
sub- family Myobatrachinae, on the basts ««l: lU 
terminal phalanges knobbed; (2) intercalary elements 
arisen!' (3i apical element ol M. ihWrmntuhhularis 
abseni; [4) pectoral girdle arcderal. O) palatines 
discrete: [6j pivpbai vn^eal lolds absent. (7) alan 
processes ol hyoid plait moderately hroad and wing- 
like: (8) cricoid cartilage divided vcntrally: t f >) 
intervertebral discs not fused to centra in adults; (10) 
attachment of M. iHWrmantiihaiiins. upon M, 
snhmtJimhs .thsent 

Ihc first three eharacreis exclude the species from 
the Hybdae, and the fourth excludes it from the 
Kanidac. wtiiisl the llrsl. Fourth, fifth and sixth exclude 
it fmrn Ihc Microhyhdae. The remaining character 
are definitive of ihe leptodactvlid subtarniK 
Myobatrachinae (Parker 1940; Lvueh I97|; 1 v let 1072 
Da vies 19X7 » 

Litdcjohn ei at, tl993i report 9-12 cenera as beiiv- 
variously recognised within the M_voh;itraeluriae 1 he 
new species cannot be placed in any of these genera 
on the bas'is of the following combination of chanicieiv 
an apparent autapomorphy of the vertebral column ol 
fusion ol die ptx'sacral vertebrae VII and VIII with the 
sacrum, presence of a dentate maxillary arch, the lack 
of vomerine teeth, ptesetice ofa columella, and direct 
deve|i>pincnl o\ the young with a four legged, tailed 
faciei hatching from the Egg membranes. 

Flic Froe is exchided specifically from Ihe 
myobatrachine eenera as follows: together with the 
vcitebta) fusion. (J) trom Assa by the absence oi 
parental care by the male, by the absence of vt»mei me 
teeth ahd oy the complete phalangeal formula. (2) front 
Crinui isvnsu Blake 1973) by the absence of vomerine 
teeth and the absence of a frcc-swimmine. tadpole: (3i 
from ('n'ttauuahy the absence of vomerine teeth and 
ofa projecting dorsal flange on the anterior ramus ol 
the pterygoid, by ihe protecting snout and by the 
granular ventral surface: (4) from Runitfallsl by ihe 
width of the bases of the alary processes of the hvoid 
by the absence of free-swimming tadpoles and by llu- 
uatureol the omostemuni: (5) from (atuiacrytus by ihc 
absence of T-shaped terminal phalanges mi the diyils, 
by the absence oi vomerine teelh and by the lack nl 
free-swimming tadpoles: (Ti) from Upefflteia by the 
absence \^ hypertrophied dermal glands and o\ two 
raised compressed metatarsal tubercles, and by the 
absence of a tree-swimming tadpole: (7) fttifti 
Pst'ttdtiphrvn*-: hy the presence of a columella and ot 
teeth on the maxillary arch, and the absence ola Ircc 
•-.wiHimmg tadpole; (8) from Aretiophrxtw by the 
presence of a columella and ot teeth on the nmillaty 
arch, and by the absence of moditied phalangeal 
formulae on the hands and feci. f9) from Myobai ravhus 
by the presence of teeth on the maxillary arch. h\ the 
absence of a inodifieel phalangeal lormula of the h.<ud> 
;jjkJ by the absence oi reinforcement ot ihe anterioi 


locality as holotvpc. coll. P B. Brown & I"). /iegeler. 
29-^-1^)2: NMV DO73I0. CT, same locality as 
holotvpc coll. A. & J. E. Wapslra, 29.xi.f992 

rjtutiftri of (he skull hy anterior placement ol Ihe n-isals. 
{{()) from Mt'Untiriut by the presence ol teeth on the 
maxillary arch, by a large omosternum, by the lack 
of extensive reduction of the palatines laterally, hv diC 
less extensive nasal*, ami by the projecting snout and 
shape ol Ihe head. 

Genus Bryohatrachus yen- nov. 
Type species; Bryobatntchus tiimhu.\ 
Di<t$nosiie dtjhtmon 

I. Alary processes of hyoid plate broad and wing- 
like 2. Cricoid curtilage divided ventrally 3. 
Intervertebral discs untitled in adults 4. Attachment 
of Al iniemuottlibularis on M. submentals absent. 5. 
Hypcrtrophicd dennal glands ahsent. ft Small flalieucd 
inner metatarsal tubercle. 7 Tiny, or no outer 
metatarsal tubercle. 8. Snout projecting when viewed 
from above. 9. Maxilhuy arch toothed. 10. Vomers 
reduced to one or two fragments at edge of choanac. 
II. Columella present. 12. Palatines slightly reduced 
laterally. 13 Nasals not located anteriorly on skull 14. 
Omoslernum large, mushroom-shaped wall narrow 
stalk. 15 Phalangeal formula of hand 2,2,3,3, 16. 
Phalangeal formula of foot 2.2,3.4,3. 17. Terminal 
phalanges knobbed. IS. Vertebrae VII and VIII lused 
with sacrum. 20. Development direct lour -legged 
tailed Irogiet hatching from egg membranes. 

Derived from the Greek bryun (— moss) and 

butraviuKs ( - hug), alluding to ihe habit of breeding 
in moss or moss-like vegetation. 

BryobatmehuK nimbus sp. now 

rgs i io 

H<>i<>i\f>c; TMAG CI012, an adult male. 3(X) m north 
^)( Lake Espcranee <146°46'fc. 43°I3'30"SK HarU 
Mountains National Park, 920 m, collected by D. fc. 
Rounsevell & D. Ziegcler on I&X.I993. 

Paraiypcs: 9 Ct &, 3 9 9 and 2 s,a ! TMAG 
C10O9, o\ same data as holotvpc. except collected 
1x4993: TMAG CIOK), u'. same data as holotypc; 
TMAG CI013. s.a.. Mt Spivnt (M5 58'£a 42°47'30"S), 
coll. D. Ziegcler, 21.X.N93; TMAG C1024 Of, 
lopotype, coll. P. B. Brown. 2H.mU993, I 'MAG 
0102$ 9 . same data C1024 and C1025 in amplexav 
I MAG C350. (2 cr cr). Mt La Perouse Base Camp. 
ANZSLS Expedition Jan 1984; TMAG C345. s.a.. 
same data as C350; TMAG C869, ry . Ml La Perouse 
<I46°44'30"T:'. 43°30'20 "S) s rocks above Pigsty Ponds 
on Ml La Perouse Irack. 1.7 km NW of summit, coll 
M. N. Hutehmson & S. Hudson, 9.ii.l990; SAM 
R4367I (cleared and stained), o*. same locality as 
holotvpc. coll D. E Rounsevell. 25.xi.l992, SAM 
R43072, ■ /. same data: AM R143566 (cleared and 
stained), 9, same locality as holotvpc. coll. D. E. 
Rounsevell. 25 xi. 1992: AM R143565. a, same 


A small species (males 19-27 mm SA'. females 
25 30 mm S -V) characterised by a looihcd maxillary 
arch, presence ofa columella, lack ol vomerine lecih, 
unmodified phalangeal formulae, knobbed terminal 
phalanges, widely exposed frontoparietal fonlanclle, 
presacral vertebrae VII and VIII fused with the sacrum, 
iintringed fingers and toes, granular ventral surlacc. 
dorsum consistently marked with dark chevron-shaped 
mark between eyes, pair of parallel dark lines from 
shoulder along anterior portion of hack and pair of dark 
patches in coccygeal icgion. eggs large and 
unpigmented. four-legged tailed froglet hatches from 
egg membranes, advertisement call a series of "toks" 
with a pulse repetition rate ol 5.3 6.1 pulses/s. 

DtMriftiitm ttf hi)h>i\i>e 

Maxillary teeth piesem: vomerine Iceth absent. 
tongue .ival, free behind; tympanum obscure- Snotil 
short, projecting and slightly iruncated when viewed 
Irom above, overshot and sloping posteriorly when 
viewed in profile (Ptg. IA). Eye lo naris distance les- 

Eig. I. tfry<>innnn hu*. ntmhtix gwi- a sp. nflV A. laicrul view 
n( the hc;id; B palmar view o| hand: unit C. phmtiti vie* 
nf fom iMolutype. TMAG IOI2i. Scale bar ?> Ittffl- 


f> r KOliNSLVEIl I IX /ii r.rlEft. P. |i brown, m twins & M, J, littlhohn 

tftwi inu-riiarial span (E-N'IN - 0,7b, Nures located 
laterally on snout, directed dorsal Iv. Canthus totalis 
sUatt-'hi, loreal region concave, facial shell 
conspicuous. Hve promiticin. pupil horizontal when 
constricted. Fingers short, unwebbed with flattened 
subarbcular tubercles (Fig. IB), Terminal discs 
undiluted. Palmar tubercles flattened but conspicuous. 
Several supernumerary tubercles present: fingers in 
order rrj length 3>4>2>L 

Hiiidlimbs short (TL/S-V 0.3b), lues relatively 
short, un fringed and unwebbed (Fig. 1C); in order of 
length 4>}>5>2>\_ Small flattened inner 
metatarsal tubercle, and liny rounded outer 
tubcrelc. Subartieular tubercles small and noi 

Dorsal surface sparsely tubercular tubercles linear 
along length. Prominent tubercles on dorsal surface 
ol legs Ventral surface granular with well -developed 

Fig. 2. Ri Yt>b<ttr,hiw.\- nimbus gen Cl Rp. nov A. cJtti sttt.ilcml 
and B. ventral views in lite (fVrtiivpe TJVJAC HHN) ff'M" 
J. VbM) 

coarsely granular pelvic patch. Small bifurcated 
unfirnbriatcd cloacal Hap. 

Dorsum dark tan in preservative with ehevron- 
shaped black mark between eyes, paired clongaied 
black markings from scapula region, and paired 
moderately elongate black patches in the coccygeal 
region. Canthal stripe anterior to nans, through loreal 
region and eye to axilla. Pale patch beneath eye (Fit's 
I, 2) 

Ventral coloration chocolate with cream speckling 
(Pig. 2): dark chocolate suffusion on throat. Paired 
vocal slits pCWfcriOfiy al angle of jaw. 

Colour in life 

Dorsal surface shades of dark brown. Darkest brown 
markings distinctive varying in intensity and occurring 
bilaterally in pairs m the coccygeal region and m 
association with scapulae Coccygeal pan lie within 
region of paler brown or "ground" colour, not hidden 
when colour intensity changes. Larger patch of dark 
biMwn occurs between scapulae and anieromedially 
which, when at darkest, can conceal upper pair ol 
bilateral markings. Chevron-shaped mark of d^rt 
colour between eyes. Small white patch above cloaca 
Limbs barred with dark brown. Other broad dark 
hrown patches occur along paler brown Hanks. Dorsal 
surface pi snout and inner thighs unmarked and paler 
brown or "ground" colour. Paired canthal stripes bom 
ttari.s to (lank broadest in tympanal ivgion Dark brown 
canthal stripe passes through mosL ol the eye eveepl 
uppermost part of iris. Iris dark brown below anil 
iridescent gold above facial stripe Lower stripe i)\' 
white or cream, commencing between eye and nans 
and varying in width being narrowest, or with lower 
edge notched, below eye and including neither lower 
eyelid nor upper lip. Ventral surface dark brown 
covered with irregularly-shaped fine white spots oi 
similar sue extending over the limbs and sometimes 
to lower throat. Chin sometimes paler and less spotted. 
Pale lemon-yellow tint on paler parts of rhroal and 

Dimensions (mm): S-V 22.6; TL 8.2: E-N 1.7; IN t.4, 
t 2 7. 


There is little variation in external morphology, other 
than colour pattern. The protruding snour is not as 
pronounced in all paratypes; in those in which it is 
most developed, it appears to be thickened and whitish. 
The legs are uniformly short (TL'S V = 0.35 
|0.3I-0.38|) T and the head usually longer than width 
(HI /HW - J.UG 10.96-1.23. ri - 9|>. there is 
variability in the relationship between cyc-to-naris 
distance and iutcrnarial span (L-N/IN -- 0.g«3 
|07l I.L2|L The tympanum is obscure or indistinct 



(Figs I, 2). Toes ate uniformly unhinged, but there 
is variability in development oi ihe put mar tubercles; 
in TMAG CIU09. these are particularly pronounced. 
When present* the outer metatarsal tubercle is tiny ivA 
the inner mclaiarsal tubercle usually llaliened A tarsal 
fold is present in TMAG ClOtO. Rugositv of the dorsutn 
varies bill is not conspicuous- Ventral granularity of 
the belly is variable. 

The ground colour of the dorsum and the extern 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 arc often iyrate am' can 
vary in length, occasionally almost coalescing wilh the 
coccygeal pair 

Dorsal colour varies from very dark brown to groy- 
brown or Ian, and, in darker specimens the amerior 
pair of markings becomes obscure and often merges 
with the suprascapular pair, f he coccygeal pair always 
contrasts with the general bodv colour and is distinctive 
(fig. 2A>. 

A pale or tan nhd-vertebral stripe is presenl in several 
specimens, and occasionally a pale medial ventral 
stripe is present in paler specimens. The ventral surface 
can be dusky grey with white spots, or pale with dark 
spots. The throat is always pigmented, but added dark 
suffusions arc present in calling males- The throat has 
a salmon iridescent hue in lite, and a lemon colonition 
often occurs in the axillae. Reddish patches occur along 
Ihe lower Hanks ami in the inguinal region. A pale spot 
often occurs above the cloaca. 

t'hc pale stripe beneath the dark camhal stripe of 
(he head varies in extent, maximal development being 
sliuwn in Fig, 2, 

0\u>o!o s -\ (based on SAM M3G7I) 

Skull poorly ossified- Sphenethmoid ossified 
medially, with ossification extending anteriorly 
between the nasals dorsalK and venirally. Cartilage-- 
of nasal capsules arc considerably calcified ventrallv 
(Fig. 3). Proolie and cvoceipital superficially fused by 
extensive calcification giving short and stock)' crista 
parotica. nxoccipuul calcified dorsomedially and 
vcntromedially. Crista parotica not articulating with 
elongate unexpanded otic ramus of squamosal. Carotid 
canal roofed on frontoparietals medial lo well 
developed cpiotic eminences. Occipital condyles 
widely separated. Froniopurieial fontunetle widely 
exposed for approximate length oi orbit- Anterioi 
extremities of frontoparietals extend anteriorly to 
anterior margin of frontoparietal tontanellc. Orbnal 
edges of frontoparietals slightly curved and angled 
slightly posterolateral!)' Nasals moderately well 
ossified, approximately triangular and widely separated 
medially- Maxillary process short, and widely 
separated from well-developed preorhital process of 
pars facialis of maxilla. 

Palatines moderately iobust, reduced slightly 
latctally and curving pcslerornedially to overlie 
sphenethmoid at anterior extremities of orbit 
Kirasphenoid moderately robust with moderately 
broad, deeply erenale euliriform process extemlinji 
approximately .4 length of orbit in ventral view. Alae 
moderately short, relatively broad, extending slightlv 
poste ml ate rally, not overlain by medial ramus ol 
pterygoid, Pterygoid robust; anterior ramus in lony 
contact with maxilla anteriorly, and with base ot 
squamosal shaft and with cartilaginous quadrate 
posteriorly. Squamosal robust, with short zygomatic 

fig. X Iint'hiUfih1iu\ uufthus^cn ct sp. tun.: A dpucaj JtiJ P vvnirjl views of the skull o1 SAM R4367I. Scale bai = ? mm 


n i- uolnsi:vfi.i , [>, ZikGtLHR. p h known M. davii.s a m_j. LjTTLfirOHN 

ramus and long unexpended (Jut! lamux not overlying 
crista parotica [Fg, 3). 

Maxilla and prcinaXiiiu dentate, Palatal shell deep 
wiih well-developed palatine processes of pivniavillae 
not abutting medially Well developed pterygoid 
process of maxilla. Alary processes Of picuiaxillac 
broad al hase with narrower poster* >dorsal projections, 
slightly concave. Vomerine (raiments present on edge 
of choanae on right hand side, Bony columella present 
(% 3» 

Pectoral girdle arciferal and rohusl (hie, 4A). 
Omosternuni large, muslmn'm shaped with an elongate 
narrow stalk; xiphisternum also large and mushroom 
shaped, with a short broad stalk. Some calcification 
oC xiphisternum and cpieoracoid curtilages. Sternum 
cartilaginous. Clavicles moderately slender, curved, 
poorly separated medially: cor-u oids robust. 
moderately widely separated medially Scapula 
bicapitale, longer than clavicles Mipiascapula about 

1 1 ossified, 

Bight non- imbricate proeoofoUfl presacral vertebrae. 
Vertebra VIII (used with sacrum and with vertehra VII. 
Vertebra VII fused with vertebra VIII and vulh sacrum 
(Pig. 5). Transverse processes of vertebra IV with 
bilateral anomally (Fig. 5). Sacral diapophyses poorly 
expanded Relative widths of transverse processes: 

medioproximal side ol base ofO. metatarsus II. Second 
element appears to be result of fusion of two elements 
and lies ut base and slightly lateral to O. metatarsus 
II, and articulates with hase of O. metatarsus I and () 
ceniralc prehallucis. Distal preliallical element small 
and knobbed, calcified. 

Hyoid plate longer than broad (Pig. 4C). Base of 
alary processes occupying '/J to '/;• of lateral edges ol 
hyoid plate; not pedunculate. Anleromedlal processes 
of .mterioi hyalc long and slender, Posterolateral 
processes of hyoid plate irregularly shaped. moderately 
long Posteuor cornua ossified. 


Umstyle bicondylar with dorsal crest extending 
appro*. ' h length. Small rounded dorsal prominent r 
on ilium (Fig. 5). Dorsal protuberance not prominent 
llial crest absent, Pubis calcified, 

Humerus with strongly-developed deltoid crest 
anleroprovimally. Phalangeal formula ol hand 2.2,3,3. 
Carpus of six elements exhibiting moderate torsion. 
O. radinle and O ulnare present, O, utdiale larger Of 
the two. Both elements articulate with O. radioulna 
proximally and with each other posteromedial l\. 
Dislally. both articulate with large transversely 
elongated O ccntiale poslaxialc which articulates 
disi;illv with buses of O. mclacarpi III, IV and V. 
Moderately well-developed flange extends tmiu 
lalemproximal corner. Small calcified palmar sesamoid 
pM'VJiualiy on ventral surface (Fig 6>. O. eentrale 
prcaxiale arliculates laterally with O, radiale. dislally 
with 0- cent rale posiaxiale and with carpal element 
ol O. distale carpale 2 and O. dtslale carpale 3 and 
laterally with basal prepolhcal element. 

Carpal element of O. distale carpale 2 articulates 
with carpal element of© distale carpale 3. Distal tips 
of terminal phalanges knobbed. 

Phalangeal formula o\' foot 2.2,3.4,3. O tibiale and 
O. fibulare elongate and fused at either end. Bones of 
approximately equal length. Two distal tarsal elements 
present. Lateral elements largest, lying at base ol (X 
metatarsus III and extending laterally to articulate with 

liy 4. Hrsi'hatnu/uty ttimhuA gen. et up, no\ . A- doisal vm;\\ 
ol [\w pectoral girdle B lateral view ol the ilium: C. ventral 
\il-w ..I the hvoid (SAM R43b7lj Scale bars - t mrn 



Fig. 5. Bryohatrachus nimbus gen. et sp. nov. : A. dorsal view of the vertebral column B. ventral view of vertebrae VI. 
VII and VIII and the sacrum (AM RI43566) and C. ventral view of the vertebral column (SAM R43671). Scale bar = 5 mm. 


Fig. 6. Bryohatrachus nimbus gen. et sp. nov.: A. dorsal and 
B. ventral views of the carpus (SAM R43671). Scale 
bar = 5 mm. 


Paratype (AM R143566) 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 
omostemum, 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. 



.Advertisement call 

Males call from ihe ground surface beneath dense 
vegetation, and from prepared breeding chambers 
within cushions ol sphagnum oredaphic lichen. Male 
advertisement calls are heard in loud chorus, diurnallv, 
in spring and early summer. 

The following description of (he advertisement call 
(Fig, 7) is based on the sequence recorded by G. F. 
Watson. I "he call is a single quasi-periodic pulse train 
with a duration ot 10O9-J2S1 "is (mean = 1157), and 
consists i>f ;i series of 7-8 (mean = 7.5) short pulses 
(duration; range = 7.2-10.4 ms; mean = 8.74 ms) with 
rise limes (attack) of about 1,0 3.5 ms and fall tunes 
(decay) of about 4.0-5.V ms. The pulse rates range from 
5.3 to 6.1 pulses s (mean = 5.74) (Table li. The calls 
are repeated at a rate of 2.65 calls/min. Although there 
is a wide spread of spectral energy (as a consequence 
of the short rise-times oi the pulses), there are two 
dominant Ircquencies of equal energy at 2100 2140 and 
2f>8Q My in three of the calls, there is only one peak 
at 2140 H/ in two calls, and in the remaining call there 
are two peaks at 2120 and 2740 Hz (with the latter 
being lower by 2 dB). 

The values obtained from each of the 18 calls 
recorded by P. B. Brown arc 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 eail 
duration, number o\' 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 i960 Hz. 

Ctilling period 

The calling period in alpine habitat is seasonal. Calls 
in chorus were heard al the type locality from early 
October to late December. Earliest calling heard was 
in the Hart./ Mts on 1.x. 1993. and the latest at Mt 
Norold (146°LV40"E, 43°15'30"S), on 2. hi. 1994. 
Chorusing occurred in ram and whilst snow fell. 

Fn early spring, calling was interrupted frequently 
by heavy snow talis, and recommenced when thaw set 
in. During summer, calls ceased during the warmest 
pail of the day at temperatures above 15°C and as the 

-^/V— "-n-— — r* 

200 400 600 800 





Fie 7, A wave lotm display of an advertisement call of a male of tirvahatravhus nimbus, gen. et sp. nov. recorded Ji ii 
wet butb air temperature of 8.5 a C, in the Hart/ Mountains. The depicted call is the first of the series recorded by 
Ci. F Watson. (See Table I for mure information.) 

\'\\\\ i. I, Values for si\ aclveriiscment vails of a male of Bryobatraehus nimbus i>eti. et sp run' at the typr lotutirs af a 
wet-bulb air temperature of K5 A C on 5, Kit, b)92. 


Pulse rule 

Pulse duration 

Dominant f 

equencies (Hzj 

Ddferenee (dRl 





lower (DFI) 

higher |DF2) 




5 33 




1 063 









8 5fl 





5 56 




















IABI.H 2. IWt/t v/V AS udvrrlt\cmt>tt( tolls tij si-Vtrtil nnli-i 
of Bryobatrachus nimbus #i'n. el sjk now recorded h\ P. ft 
Hrnwn of ttie hpf hteidfty on 30. \iJ992. 



Pulse rale 






(H/i * 




4 20 


































4 4d 












1 1 

































5 M 







tmdpoint ilf band: ** second peak at 1820 Hz 

vegetation dried oul. Individuals called in cool 
conditions during the early morning and in the evening 
(19:00-21:00) at Ml La Perousc on LiUttft (S. 
Corbcti pers. cumin, i. Ftugs were no| heard at night 
Ziegcter < 1994) observed culling in air temperatures 
of 4 5-12°C and reiauve himiidities ol 62 -94'£. The 
frequency ***** calls beard increased when there was 


The species is crypto/oic, and was eolleeted during 
.spring and summer. Calling naies were eolleeted trum 
the surface of the peat beneath coarse, low vegetation, 
from within breeding chambers 01 nests, in cushions 
ol sphagnum or oilier simiiai plants, hc-neath rocks 
(Ziegeler I994). or under branches lying on or 
amongst vegetation, Females were found in breeding 
chambers with a mule or. when males were chorusing 
in spring, on ihe surface of the vegetation. Males 
continued to call even when leuiales occupied the same 

At the type locality where a huge population exists, 
individual frogs arc regularly spaced, remaining 
hidden. They appear not to aggregate or to use "pen 

Fig X. A. Ncsl ol Brvohalroihus nimbus gen. C{ sp_ <io\_; B. subalpinc moorland habitat al Harl/ Mts; type localUv in 
middle distance viewed horn the track (Photo J E. «& A. Wapstral; C subalpine moorland habitat al 800- 1000 tn aliunde 
at Ml Hesperus, I). Implicate ramloresi habitat at over 750 m altitude at Mi Hobs. 



hty, *■). Hi v.ihufuhhus nrnthitt gt»n. cl Sp n*»v A dUI$Ul view 
C*| ggja; B, dermal view of cmbryu at stage 5 I Fownsemi 

vV: Stewart |V>XS). C , ilnrvjil VVfiWV "» ImgliMs 

surlace water. None was collected during autumn or 

Locomotion is by crawling or walking In (he open, 
frogs usually crouched and leniained Mill until 
disturbed and Ihcn sought cover. They jumped or swam 
only when forced. In captivity, both sexes displayed 
thigmotropism by forming individual cavities in which 
to shelter in loose sphagnum. They entered the 
sphagnum backwards. 


The species breeds in spring, laying eggs in nests 
in moss or lichen- Nests >ne concealed spherical 
cavities (3.5-4.0 cm in diameter) in clumps of moss 
approximately 2-5 cm below the surface (Tig. SA) 
They can be exposed by parting the stems of moss. 
Single males, male-female pairs (sometimes in 
amplcxus), eggs, or froglets were found in ten nests 
in a small area Of the lype locality on three visits 
t24.\ii.lW3, II. i.1994 and"28.i.lW4) On the fust visit, 
live nests were Ibund. One contained 6-10 eggs, anolher 
contained five froglets (Fig- 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 
Visit three nests containing eggs were found. Two 
contained 12 and 14 eggs. Those in the remaining nesi 
could not be readily counted as the jelly surrounding 
Ihem had begun to merge. The last visit produced two 
new nests, one containing W eggs, and another 
containing 5-6 froglets. Bach nest was found in a 
separate cushion of rnoss. Other empty and possibly 
disused nests were round in adjacent patches ot moss 
in the same area. 

Eggs containing embryos (Fig- 9B) at stage 5 of 
Townscnd & Stewart (1985) were located on I0.xii.9j* 
bin not in nests. Two groups of eggs were found on 
the surface ol the vegetation and contained three ;ind 
six eggs respectively. The latter included four dead 
embryos mlcetcd with fungus. 

The mean diameter of lour of the newly laid eggs 
round on ll.i/M was 3.49 (nit) (3-33-3.65)i and the 
rm-an capsule diameter was J3.57 mm (13.02 -14. 13). 
They were surrounded by a single jelly membrane (fig 
°A) These eggs cannot be ascribed to a stage as for 
ihe direct developing FJeitthertu/ttcty/us roqui 
flownse.nd & Stewart ls>85) 

The mean S-V length Of fiVfc melamorphs was. 
6,0 mm (5,4-6.35) and the mean total length. 14.51 mm 
(13.97-N.05). fail length varied considerably {Fig. 9C). 
Tail muscle is well developed and, tail fins narrow, 
equal in width dorsally and vcntrally. Tail fins do not 
appear to be vascularised The body is heavily 
pigmented; that on tail muscle being liner and thai on 
tail fins patchy. In lite the body and tail are coveted 
with fine while dots. There is no eloacal tail piece 
The mouth extends beyond the eye and the fingers and 
toes appear lo be fringed, 




flrvv that melius nimbus is Ibund in subatpinc 
moorland (Fig- HO) and implicate ramlorcsi m southern 
Tasmania (Fig, HO). It is icstricted io poorl> -drained 
sites from lowland to subalpint* localities t/iegelcr 
1994). The soil type is peal overlying .sandy or stony 
substrate i/iegcler 1994). It occurs on Pre-Cantbnan 
inetaniorphies. Upper Carboniferous-Permian 
sedimentary deposits, Trtassie sandstone and 
Pleistocene glacial deposits. 

The subalpinc moorland comprises <» diverse range 
of plant communities. Five recorded communities are 
IfKh ris set pili folia- ttnpad'tsma mnuis-Gleichcnia 
alpina lernland. Lepn>spenman uitidum-Galmia 
yrandis-E. minus tail shrubland (both occur at Hart/ 
Mis [fag, 8B|>. £ M-rpilifi'lhiRirhvu \coparin low 
shrubland with Asiefia alpbnt, G. alpina and t. nanus 
i Adutiisons Peak |146°49'|-, 4^°2rS|). E. serpilifolia 
Melaleuca squamea open heath (Ml Sprenu ami 
Carpha alpina Isophvsis tasnuitica herbtield (Ml 
Sprenl. Mt Hesperus |146°l4'ri, 43°07'2O"S] [Fig. 
XCj) (Zfogefer 1994)- 

The impheale rainlorest communities aie tloristtcally 
complex. The shrub layers are dense and species 
diverse at all altitudes Ai higher altitudes ihe dominant 
irecs are NothojaiiHs eunnin^hamii, liuaypltia 
ntiWxatiii, Irhtutaiis \elai>iin>itles, Naihofit^us gUimH 
and Phxliocludtts uspleniifalius and the ground cover 
is dense moss andor/l.v/W/V/ alpina (Fig. 8D; 750 m. 
Ml Bobs M6*36'E. 43°18'Sh At low altitudes A' 
i unnintfhamii , Melaleuca squarrosa and R 
aspteniifo!iu\ are dominant and ground eovei is dense 
moss and Bh( latitat watisii (Alexander Creek 
I4r>°05'b. 4^:6 , S> l7iegtfer 1994). 

I he climate is m the perhumid cool *onc of Gcntilli 
(1972) but is cool and consistently wet. The mean 
annual precipitation is 25(10 mm in large parts of this 
region and is received as rain, snow. hail. fog. mtst 
and frost evenly distributed across the seasons t Bureau 
Of Meteorology 1991), 

Distribution and abumkniee 

Bryobafrutlms nanbus occurs in mountains in 
southern Tasmania south ol 42°48' latitude \¥Q 10). 
It is known from 15 localities, mostly described by 
Ziegeler (1994). that range from sea level to 000 m 
in an area approximately HO kin < 50 km. The 
localities arc bounded by Ml Sprenl in the north-west. 
Haru Mts in the north-east. Ml La Reroute Hi the 
south-east and Bathursl Harbour area in the south west 
(Fig. 10). All localities arc south ol Ihe Muon \h\er 
and Serpentine River and none is over 40 km from the 
sea. At the northern and eastern edges of this 
disiribuiion. B. nimbus is confined to sues above S00 in 
altitude whilst in the south-west, its ranue descends 
to near sea level. 

Hundreds Of males were heard in chorus at the i> pc 
locality during spring 1VMJ. Other large population.s 
occur ai known locations (Zieeeler 1994), Allhough 
>2ei>graphically restricted, the species is abundant 
within available habitat, and calling males wcic 
estimated to occur at densities ot O.t-I.O'irr- 

Seafches conducted further north at Philps Peats. 
t'yndall Range, Ml Field. Mt Anne. Mi Weld and Ml 
Picion lailed to locate the species despite ihe presence 
ol' suilabte habitat and weather conditions (/tegeler 

Consetration status 

The species is secure. All locations where it has been 
recorded are in reserves, as is most of Ihe suitable 
known habitat. Us habitai shows evidence of no lire 
oi only very low frequencies of lire. 

From the Latin nimbus (= rain cloud) 
reference to the habitat o\' the species. 


(\nnnt<tn name 

The name "moss froglef has been coined and used 
for this species (/aegeier 1994 > , 

1 46' 00' 

1 Mt Sprenl 

2 HarVMt 

3 MtUtPf-iOufee 
A Ml Hssperus 

5 Mt Naroid 

B F^Uerelrar pi 

8 AOamsunfi a k 

9 'Moonlight tucigr 
10 PreapilnuR BluM 

n ironbound Ra 
12 Loui&a R. 
i3 Balhucat Ra. 
14 Mi Mslaleucu 
'5 Alexander Ck 

Fig- 10, Distribution ol ESryobanxivhus nimbus gw*. el sp no* 
m southern Tasmania A, Southeastern Australia, and ft. 
localised distribution in Tasmania. Closed entice an* 
localities IVnm whim ^wviruenv li;tve been collected; open 
(numbered) circles mv cull records. 


I) I- KOUNSJ-VM1 li /Ami I I R f 1 B UROWN. M. DAY II :S & M I J ITTI.KKjHN 

Comparison wuh ttther spct'iex 

Br yt thai melius ttimhu.s is distinguished Irom other 
mvobaLrachine frogs in Tasmania in the following ways. 
The species U not associated with bodies of op(.n 
surface wafer. || is most likely in be- confused u tih 
sympaii ic Ranaicha lasmamensis from which il can 
he distinguished by relatively-short, unfringed toes jnd 
hv the lack of brilliant carmine patches usually present 
Utl the flanks and the concealed surface of the thighs. 
The i. all oi' R. tasmunietisis is a qu.neiing "bleat 
slowly and irregularly repeated lUtllejohn 1970) ami 
its iniciohabiiat includes vegetation cm the margins E3C 
open and running water. R. sigHijera has relatively 
lone, fringed toes and is found in the Vcg&atvOfl on 
Itic margins of permanent or temporary water bodies, 
The call is a series of short rapidly repeated notes 
"ciick crick crick crick" (Martin & Littlejohn 1982) 
(iax-vinia lacvts has a rounded snout with a smooth 
ceiuntl skin which is usually boldly marbled with dark 
brown markings. The species is found in dry 
sclcrophvll (open/ forests at low altitudes (Martin & 
l.i(llc|ohn IP82) These authors describe lite call as 
El luirsh uniting Vra-a-a-a-aek cra-a-a-a-ack, crack- 
crack, crack" The other myobatracbine species in 
Tasmania is Psftui<>{>hr\ne setrttntanaorata which has 
a smooth ventral surface strongly marbled in bhtck and 
wlme. The head is rounded and the undersurfacc tit 
the thighs are bright yellowish orange to orange in life: 
(he call is a short, harsh "creek" repealed slowly and 
irregularly (Martin & Littlejohn 1982) 

CamparuUvt' material examined 

4rt*nophme roruaS- UA/ B53K B54I, K762, 

AS79 81. A760-I. Assa dariinzUmi: UA/ AGi Cnntu 
^•ot^iana: UAZ B754. AI3406. Gencritiia laevis- SAM 
R4260B. R89K2A.fc. G Icai SAM R5787A. C. 
vifturkina: SAM R9425 |2), Mytthatraefws %f*Jldih 
UA/ B4'll. P534-5. B757. AJ5& Puna rima fMwtli 
UAZ B750. Ranidellahilingua: LA/ B1927, BW5, fc 
(frscnn-oUL UAZ nisM(V3L R s>(auerti. UA/ A»l 
R. ilHttantfoW? UA/ AL92. AH>5, Bs»2s». A*. 
pariiisiyjipra: UA/ AI85. BIW.U R, raw>t<i: UA/ 
flft2fl-% R. riparia: UAZ A184. AiS'A ANX R 
siKhfftrq: UAZ AW3, A 197, B84X o?k A" 
suhtnsixnijera. UAZ A 190, R. tasmarnvnsi.s : UA/ 
A 186 Pseudophnne wmanarmonua: UA/ R63J5 P 
htbtvm: UAZ A577. AI72. B532. B5-KX A' vtnhivea, 
LA/ A20U B537. B764. Z 3 ^ueruhcrr. UAZ AlW 
053 9. B765. £ accaUntalis: SAM RJ7522 
h)u<itiit\iu.\ uattitosms; UA/ B75'J. A205-f>. A' 
lanzcttvnsts: UA2 A392, K ty^pflfbtt; UA/ A775-6. 
B7>2. tipervh-'iu idiissmut; SAM R40I46. U. arcvaa/a, 
SAM RI7347. K fl^Jptt UAZ AS72-K t_( hareahs- 
UA/ ftfea. BK62, A22o U atpintiakL SAM R29580 
t/. r^w: UAZ A869-70. B87I. B4S_>. B4S6. // /avc//. 
SAM K29603, K29606-7 U ^atuhdvsa: SAM 
R2 70S2- tl inwulata; UA/. ASKS 82b. BW. 1M79 (>. 

hievi K oni: UA/ AotH, RSI7 U hik^mrnh UA2 
A7f>7-«, B8%. B8I2, A79*>-80t). R iUtkiohm; LA/ 
AI712, AI7I7. RI7U-I7I6. M fNtfrfiirJV NMV D23Mft 
U ;;:/cao>;^m-' UAZ AI722. 6 minmla SAW 
R29642-3, R29M5-6. K mi<>hrr^. UAZ A562 
AS80 9. t ; . nKv» l7 C'AZ 1012. A 1013, AS 16, B8U I 
talpa v\.\'A \59U4, BS64 f/ imrfiyh>rm UAZ 
A62J. AK92-4. A39>-»'>. /./, /v/r/v: NMV D236.H9. SAM 
R29659 R29652 


firyohai radius nimhas exhibits a number of unusual 
(retires in its morphology And tcproUuctivc biology, 
Many t»l the characters coritnfmltng to the recognition 
of the genus and species arc influenced by hclcrocfmmv 
in other nwohaiiachine genera tDavies I9K9)- The 
jireseiiee of teeth on the niaxillajy arch, reduction of 
the vomers -tiui jhsence nl vomerine teeth, presence 
o) a cotumellit and reduction of the patalincv iatcrall>. 
are all presumed labile features that van 
inttaeenctically in e.g.. fft/ntfrnt and Crmht isfusu 
Mc-vcr el al. I982K Howl-vct. none ot these Icdiun-s 
either sinely or in c«>nibinalior» is delintlive 0\ 
Rn'olmt ixtchNs. All ate coupled with the unusual 
feature of fusion of the posterior portion of the vertebral 
cnlumu with the sacrum. 

Vertebral fusions lend to he in an anterior/posterior 
sequence (Trueb |973), ami lusinns ot presacral:-. VI | 
and VIII with the sacrum are unusual. The nature ot 
the fusion cauntu be described as a presacral shield. 
as known in a lew la*a including Bravhyc&phittld 
t'ppi(ihhan tBrachycephalidaei. ami thoueht W be 
prv*tecti\e on the basis ot" terrestriahsm exhibited by 
such taxa (1'rucb \Wfh)\ 

Fusions apptpat lunj: that shown hv B. mtnhus h;i\c 
been recorded in the- Butomdac ie .Jj , Dtt}\namipi{> 
seeO|tindist»u J9Sl| ut IhttJt-thaOs {Trufib 1973). and 
in the Pipidac; allhouv-h I he fusions reported in 
fSrtu/hvt'ir'nu hints and HYnienoihirus by Canfi;iteH;i 
A Irueb 1 19X8 ) are mteiied. and not identified by 
temnant vertehrac oi tccorttod spinal foramina, 
vertebral fusion hi Vtpa trtycr\i more closely 
approximates that in fJidynampis (Truth 1°K4L f$)f!t 
ol these faoulics is natoc uy Australia. 

Although contorminp to tlie nmibairaclune patreni. 
the width of the junction of the alary processes wittl 
(he hyoid plate is not as broad as irt most other genera 
and approaches that shown by the enigmatic gcniiN 
Rhc'ohatnuhus <Davie> &. Burti>n 1482, Mahony rial 
IVXat, fhc siihfumilml ptaccnieol ot wlii»:h reman- 
contentious tTyler IV8& l-itttcjohn ?t ai, |*W34i 

rerrc-stnaf eee deposition arid direct development 
have evolved a number of nines in Australian frogs 
(Martin l*:>67f Reproductive modes include complete 
nUraeapsulat development ic.i.', in the nucrohslids 
Sphi'tu>phryvf ami Cophhaltis) , through hatching al 



a relatively Ifttc Wtt&P in otuoy.cny leg., the 
myobaltachinc I'setufophnne} u> non-swimming, non 
(ceding larvae lhal are subject 10 bt/.uuc forms Of 

parental oars (c g . the myobatrachine >iww and ihe 

rhcobatrachine Rheobalraehus). AJI of these gradations 
ol direct development art fcUftJ wilhin the 
Myobatracliiuac. and reproductive mode can 
sometimes vary uilraycncncally. In P\CU(itfpfo w«\ £ 
th.ntyfusi differs from congeners in laying eggs in water 
ami ihuv not rely iny upon Hooding ol nests to natnuc 
IVnchlng of larvae ul later stages {Main N»4; Bradford 
& Seymour 19K.S), whilst in (U'(>erinja> tj. rosea and 
<7 laiva do no) have free swinimitv: larval stages (Main 
fl at. Y&}\ Mam 1%J). 

Intracapsulai development is known for the 
monotypic y.enera Mxobairaehus and Arvna^layne 
(Roberta 1981 1 1984). nn<l is interred for Mwairinh; 
(Main et al. 1959). The larvae ol Geoettniu rosea are 
inactive in broken down egg capsules which till 
shallow depressions in damp soil or in the iottcn 
ecntres of hollow logs (Main er al 1959) Larval Iff* 
ol d. lutea resembles that ol G rosea (Mum l°M,i 
Sir h larvae are highly modi Tied, having no mouth disc 
(I large yolk sac, and an elongate tail (Watson A: Martin 
1473). The sliueluie i)J Hie tail lcpoitcd fol C rfctfld 
is similar to thai observed in H nimbus I'ro.vlcMs, 

Allhouyh details of Ihc lift history ol It nimlno have 
not yet been dcsciibed, our limited data indicate thftl 
development is intracapsular until Ihc linal Mayes ol 
metamorphosis, and that the froglclx stay in the nest 
with (he broken iluwn egg capsules until the tad is fully 
re.sorbed and inetarnorphie climax is reached. 

The diarnelers ol ihe egg$ al V \ &,7 mm arc smaller 
thaJi ovarian diameters fccoided fol Ateiu/phtvttc 
rt'itohJa and tfgg diameters in M\u that an -has ecuhta' 
jfjbflltetls I98t, I984h but within the range resided 
lor P&eudtfpnrpw spp. and lor (re<>etitiia vieforiatta 
and 6' A/em (see Tyler EJS9 for compilation of fetflj 
All ol these species exlnbil hums ol doni 
development Capsule diameter is pailieularly luiy.e. 
being almost double (hat recorded in the field lor 
Myobalntehtts y*iwl\lii and l ( 5 Umes that ol eggs 
Ii yd ruled in the laboratory (Rohens |»)R|) Chileli si;-f 
compares closely with those or othei direct-devclopine 
species dvlei JW.)). 

hoin ot.liet (asiiuimart specict [Rdtydtifa Ufjrtlfv-Kh 
ft. fastHitniensis, (iciHrinia Uteris, I'snulophrxne 
sttniniarworoto) have advert isepk*nl calls composed 
ott ruins oi pulses. Hie short pulse trains <if S northus 
are similai to those ol A*. \h>nijera, but die call 
repetition raU-N and pulse repetition rates ami dominant 
ttc-quetkK^ aie hi.ehcr m this tuttet species tl inkjtihn 
iy<V1, N70). Ihc* call ol H. iasnia/nenst.s is to*»rc 
complex. heinf composed ol a e.roup ol pulse Iruins, 
oi notes, of hi£h ;»nd regular pulse rates (littlejohn 
IM7t)j. so thur if has .j bleating qualit\ The i all of G 
tuevi\ consists ol a scties nf pulse trains ill which the 

pulse intervals dei rease ihrouj'h each note (latllc|ohn 
& VUtriin 1964; Hatitson& Lmlejohn 1985) Theu.ll 
^ P wnimamiiwitu consists ol a single complex pulse 
iiuji). usually with biphaslc structure (McDonnell et 
al 1978) Henee, the advLtliscruenl cull ol H. nttt)l>t>.\ 
ta -.uHicioU^ unlike those ol'all other spceies of ami ran 
occurring in 'lasmania lo lie a reliable uuheutot lot 
Klenlibcahon h»ssibl>. the most siniilat advert ise men' 
cull ol a southern tuyobalrael'ud to that of 8 mtnbto 
is that at' Ranidelfa slattern of south western Western 
Australia, vvlneh consists ol u slow pulse train and very 
short pulse durations (I Jtilejobn 19-V), KirSI; M .1 
Lillleiohn & K ti ( l.tlUejolm unpubl.). 

Hryohatruehus nanhus is most suiulai in external 
nictrphology *ind cranial osteology io Ranitiella 
lasnum'ienxis tDavies unpubl.). in its teptoduetive 
biology to (ieoerinia rosea and G. lulea, and mi the 
.structure of the hyoid to Rhtobattadius, It is not 
possible, thetvlore. 1o nJcmily die sislei lu\on to the 
gcoufi on the basis of the phenetie eompurison 
presented here, Such an itlenlilicalton musi avvau a 
detailed analysts of. at least, the subfamily 
Myobatrachmae. incorporating all available data. 

ii. nimbus appears to be conlmcd to southern 
Tasmania despite Ihc presence ol apparently suitable 
habhut in highlands beyond It has been sought north 
ul the* Seipcutme and Huon River systems withoti! 
success The distribution might be explained by the 
extent ol Pleistocene jjjaciuiiou ui the centra) highlands 
of the island, and the severity of the cold, dry climate 
in ice Iree areas (Galloway l9Sfi) making die central 
highlands unsuitable for ii in ihe past, coupled with 
a lack of northward dispersal since. 

The region currently occupied by Ihe species has a 
Ciinlmually wel equitable elimaic pmduccd by year 
round high rainhdl within T^e perhumid cool climafi, 
/one (CicnUlh 1972). All localities arc less than M) km 
From Ihc sea. and arc suhjeii to coastal dirmtk' 
iiiiluences, parlieulatly increased precipitation Imin 
orographic interaction with moist prevailing west In 
south west winds. The direct development ol' the 
juvenile stages recjuiies a eludate lice Iront the 
extremes of desiccation ll is most unlikely that the 
known habhni types could support the species until a 
vers long tunc altei a wildlnc l Brown A l\«dgei 15^2) 
H nimbus can be icgatded as behiu^uig to luc 
inlolerant communities, and the occurrence ol lire 
represents ihe irreutest potential lineal to lis survival. 

During the present vv»>rk several clunhes of 
developing eggs weie touud oti ihe sin lace ot moss at 
Hart/ Mountains. The stte had been covered b> a 
Nnowdrift lor some time F.gglaying beneath the snow 
rather than in nest could account; tor then location arid 
lalct exposure alter the thaw. The embryos would be 
prone io desiccation before completing iheir 
development Altenifltivelv. stepping on patches of 
moss when walking across the site could eject cgg> 


0, ) aOUNSEVttL. I) /It.Gbl LK, P B BROWN M DAVIKS # M I, lITTIXJOIIN nest chambers, Whx hover K the case, caution 
ts advisable when working al breeding ftit ;itums 


Di 'irjeme Watson recorded the call used in the 
an&tysia Or Dale Rubens located ihc first nesi chamber 
and helped to clarify the breeding biology Dr Roy 

Swain and Mr .1. 1-irn provided assistance in the 
laboratory, and Ntkkt Mitchell helped in the field. 
Barbara and Pelcr Wilson . .layne Baliner, Sib Corbetl 
,tnd Michael Askey-Doran reported the calls Irotn 
Ml NoroUI. Balhurst Range Moonlight Ridge and 
Ml Melaleuca. We thank stall of die Parks and Wildhle 
Service who conltiouied to this study Jnd Assoc. Pn a- 
A. A. Martin and M.J, Tyler for constructive reviews 
Of the muiubcripi. 

Kt'l'i rentes 

In ,\Kt V J. D (|°73i J\t\onomy and rctauonsnips ci| 
in>ob;iimc tunc [JTQJftfc t Lepioditctvlitlue » j numeric J I 
appuui.h ,w,v/, / Ami 21. I|U-UHJ 

BfcATWP, P. F & .Si.M.HK, R, S, (lOS.s) Knetyy 
I 'itnserv: ( (nin dunne the dclavcd hatching period in (h> f"-L* 

l\<u<U'i>>t>\>u btbtmi Phys'tfri ?tfol 58(M, 491490, 
Uu<M\ M, i cV V<m:,\-» » 13 (V/H2) I lori*ri« and firt 
• iMiinte^ of a vegetation sequence from scdgcliimfhciUh In 
i;tii)l"iesl at BathurM H.irhour, tasmania, Attst, J B*u 
Jftfft] 659-074 

t \ns \. D, C $ I ku H I.. tlOKXi l.vuhiuun ot pipoid 

hoc-* morphology and pliyloyeuetie relationships in 
f\iut<Hnvu'ih>iiurns, J fhrpcf") 33(4). 43^456 
IMwis \1. iI9K<)j Onh^eny ol hone and the rule ol 
lieiOMK'linuiy in the myobatraehinc genera Ifpprfltefo, Crinitt 
and f J SL'u<lophf\nt ( Anura: Lcpiodaerylidae 
Mvohauaehinuei. 7. Morphol, 20ft. 269-WD. 

L v Hi Kiitr, T C, il l >K2| Ckicnlngv and myotomy h 
the gastric brooding rrtig Rhcohairachus Wh*<* (Anilfii: 
I .cphkUtylidaei 40ft 7. /W. 30, 503-521 

Davis, |> I). & CioHt . V. K, ff947) Clearing nd stamiim 

sou-til icrtennncs. Fictiliumt Trcfiniams 4, Vlft. 
Di'^.fkM', G. & L k. L D. (1977) Rn/yn.e clc.urm t < ol 

akian Mue Mat tied whole small vertebrates for 

eieiuoii-.dution n) cartilage Stain H'thti>>(. 52. 22 c > 33 1 , 
(,ui:tu\\ K W ll^Rhl Australian snowbelds pa^t and 

present, in Barlow. H. A. (P.d.t "flora and I auita ut Alpine 
^I'ii.tiij ago and origins." (C'S-IRG, Melbourne). 
CiiviHU. J. {1^721 Australian climate patterns " (Nel\on. 

trBAMtM^>V. A. (... C. (WH1) Miirf ihi ttoijy jml! ^liy loj;rtifl k 

P»siiion ot the WeM African uhtyrvtmipM yuexUttli 

AnuVrsson, l*>()3 i Anura. Hutoaitlaei, Mituii, ruci fUitiunu 
NS SuppL XV- No. IJ. 1K7 :i_\ 

IUkkimi^j PA & I || it uohn, M J- (I^K?) I)iphai-v in Hie 
adveiti^emc-nt ealK of Geocrhua l<u')i.\ (Anura: 
tvptodumhdaei Vfefiftl rt-sponsc o( ftialCK .liinna tietd 
pi^baCK e^pcTiinenls tttfun: h\o( SwiOblvl 18, 6773 

llni. w W R. |>AI '.h.k.v e II fcMAttBm-.l R < I^H2l 
Syste'iiiahL resolutiiui «| the genera ol ihe Crtnfa eoniplLv 
i \ .r.phibia. Aniirii Mvohatrttbuluei, Pfitu Hwl Set 
MwO,m ,,,,„•, V5, 423 427. 

I ihi iohn. M- I. |W50| C.ill tliltLTL-iiiuiiuin in a voinpk-\ 
ol seven spcevjs of Crinia lAruiru. LLpbidaLlvliUai'r 

Kxyhtifim 13. 452-46ft 

I l l »6t,i Age and origin oi -suiik- Mjuth\\r.stern AuMnikin 
\pi'»M:s itl ' Cnnia (Anuru: LqttodjL-fy lidiier pp. 51_S S3ft. 
in Plau, W. V (lut M "VfHchrak Sprcialion" (t 'rmtrsjlv 

oi T'cva.-. Pres.N. AuMim. 

(1964) Gcopraphii isolation and matin? call 

ilitierentiaiiuri in Crinia m^iuftro Ew>fU9irttt 18 2t-2-2ti(> 

_____ (I97t.n Crinia UwntwiivsiM tAniir* Lcptodaetyiidaer 

pfipgraptuc dearibuiioii, maiing. tall structure and 

r*ianftrHhips, /Vf^( tJnn, .V/(, NnvSaniii Hfoto\ 94. 119 |27. 

& Makiin, A, A, ii%4» Ihe Crinia tWYl\ eomplei 

i A n ii id Leptodactvlidue) in xouihdislum AusUjiIim- -f/o' 
,/ /auA \Z 70-83. 

RoHttios I D-. W-mson. G. F & llAVJI.R, \t 

tl*W3j Family Myohatruchiduf. pp. tV.57. hi Ros^, OL A 
lEd.J 'Tauua o[ Australia. Vol, 2A Amf>lnhia and keplilu 
iAGPS. Criiibcna). 

_ & Watson. G. K tl9H5» Pattern* ot distrihuiioii 

.preiation and vicarianee hio^eo^iapliv DJ' southeastern 
Australian amphibians, pp Vl-97. /// Griyg, G. . Sfflnc R 
t^ Khmann. II. il-.dM'The bioUfgy ol Au$tro|fl$lfin ttfify 
jnd rt-'ptilcv; (Smrey Healty &. Sitns, Chippine Norton) 
I writ, J 1) (l°7l) Pvolijiioniiry relalionships, osieolopy, 
and /oo^'tographv of leptouaetyloid Iro^-,, I'mw Kvn$a\ 
Mus- Nm. fJiM- <A.v/\_ Puhf. '53. I-23X. 

M smo^v. M-. Ty4-T_, M. J. i& Davh-.s. M (WM) A new 
vpt'ciL.-> ot thC liL'iiU:, Hh tt /lm It m hn\ ( Anin.i 
I.eplodaLlylidar) Innii Qiit^nshind 7WTV R. Sih S, \h\, 
I08f3). 155-162 

Mais. A, R (l%3l A new speetcs of Cn/ua lAnur.. 
L eptoaaciylidae i from National Kirk, Nornatup W, Ah.m. 
\<af X, 143-144. 

il'>64l A new speeics ol Psnulophixtie (Arnna 

l.epK»daetvlidae» from norih-weste-rn Au^lralia fhnt. 9. 
fi6 72_ 

, iii .itN M ! 'VI" \ K (1959) Ewluy 

ol Australian Iroys, pp. 3%-41l. hi Kcasi. A.. Croek-f 
K, I tV Christian. C, S. Hldsj "Bio^eoi-rapln antlpCT>10jy 
in Austriilin" Monograph, fttaf. VIII iW Junk I Rfi 
MARTIN A, A. (1967) Australian anuian lite histories; sonic 
evolutionary and ccolo^icjl aspect:;, pp, 175-IMI hi 
Wradiui ly A W (Ed.) 'Auslodiyn mbnd wiitt-oj ;in<J their 
tiiuiid ' ' Atislriiliari Nalionul l_ nivtrsily Hivsw. ('aiibcrni) 
, & Ll'l iLhiOHN, Vt J (NM2) -Tasriiariian amphibians" 
Paun:j irf laHmania handhuok No 6 (f'niveiMty o| 

l.iMnania. Itobart)- 
M<IX)NNiii I J,, G\kisu>i: D K & Lrni.iJoiiN, M". J 

d^THi Analysis ol a narniw hyhnd /one between Iwli 

species ui Pwtiilapfvyiw (Anura; Leptodaetvlidaei iii SOUth 

L'tislem AuMialta, Kvnlntitm 32, $02 tfl2 
I'ahmh, H- W (l ( M0) The Australasian \r&&0\ the ijind-. 

Leptodaetyhdae flftpfr. Zt»o/. 42. 1-106. 
ROtitlffS. f. pi (I'ASti IorvcMnat breodiny m the Au^todtan 

Ic-ptOdavtybd lio^ M\<A'ti/mihn\ x^aniilii (Grayt /h(,^ 

K// ( //_ AV\. K, 4.SI-JM 

I I9H4 ) 1 tr i c s I f i a 1 egg dcp< »$il n m and direct 

development in irtnnphryne rotunda Tyler, a m.vohatruetud 
frog homeoiisial sand dunes al Shark Bay. W.A. thift> II. 
l ( d-200. 

R.i-uk.nhn, S, tl993j On lhc iraek of the unforgettable- CTDiifc 
ra.smani.o, new trot', tbffi 15(2 ». Ifi^l7. 



Rounsevfjx, D. E. & Swain, R. (1993) Current issues in 
the conservation of the terrestrial herpetofauna of Tasmania, 
pp. 71-74. In Lunncy, D. & Ayers, D. (Eds) "Herpetology 
in Australia - a diverse discipline." (Surrey Beatty & Sons, 
Chipping Norton). 

Townsf.nd, D. S. & Stewart, M. M. (1985) Direct 
development in Eleutherodaetxlus coqui (Anura: 
Leptodaetylidae): a staging table. Cope'ta 1985 (3), 423-436. 

Trljeb. L. (1973) Bones, frogs and evolution, pp. 65-132. hi 
Vial, J. L. (Ed.) "Evolutionary biology of the anurans: 
contemporary research on major problems' (University of 
Missouri Press, Columbia). 

_ (1979) Leptodaetylid frogs of the genus Telmatobius 

in Ecuador with description of a new species. Copeia 1979 
(4), 714-733. 

(1984) Description of a new species of ' Pipa (Anura; 

Pipidae) from Panama. Herpetologica 40(3), 225-234. 

TYLER, M. J. (1968) Papuan hvlid frogs of the genus Hxia. 
Zvol. Verh. Rtjksnms. Nat. 'Hist, Leiden No. 96, 1-203. 

(1972) Superficial mandibular musculature, vocal sacs 

and the phylogeny of Australo-Papuan leptodaetylid frogs. 
Rec. S. Aust. Mus. 16(9), 1-20, 

_____ (1978) "Amphibians of South Australia" (Govt Printer, 


(1989) "Australian frogs" (Viking O'Neil, Melbourne). 

Watson, G. E & Martin. A. A. (1973) Life history, larval 
morphology and relationships of Australian leptodaetvlid 
frogs. Trans. R. Soc. S. Aust- 97, 33-45. 

Zeigeler., D. (1994) "Survey of the distribution and habitat 
of the moss froglet (Genus nova, species nova) in south- 
west Tasmania" Wildlife Report 94/2, Parks & Wildlife 
Service. Hobart. 




ByD. Pamment, L Beveridge & R. B. Gasser 


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 

h<muHt«^ <>? tin Row! Sou.-iv •>( s Auto. (1994), mQl K7-496 


by 1> PammenT, I. BEVl.RUXil $ R. B. Gassi K 


I^mmi-M- D. Blvi KMK.r. I, & Gvwi-k. R B, ( 19*>4 1 The distribution '"•! nematode parasites ttUhtf HM ttiimiitih 
ol the western grey fcmgiirpu. Mutropusfii^moMts. Trwty K Sof S Attst 118 ,(.$) IH7-lo u , 30 November. N'M 

The distribution ol'stron»yhd nematodes in the -aomaehs often western grey kangaHfOs {MiK'fOpUi /h//cw"MI.v), 
lioni Haltah, Victoria indicated that eatfi gfiHUS encountered, {hucinu, K/kv/'Wou, hibto\inmxvln.s, 
PpfVMlStr<tf)%yluti and Ftlunntnui. occupied u specific region wiihin the Ntoiiiaelv Ih'tr di-anhution bore no 
relationship to histologically defined i^miis of yasiriL mucosa, and IliC pathological changes detected in the mucosa 
were related more closely with mucosal type than with local nematode densities. 1 he anatomical and histological 
leaturcs of the Momueh q\ M !uh\>tni>\n,\ w described. 

Kl s Words Neniatnda Strongyloidea. M«> roptis. Mitcrornididac, dislribuiion. gastric attalomv, 


The differential localisation of multiple, closely 
related parasitic nematode species within the 
gastrointestinal tract of their hosts has been investigated 
extensively in the ease of the oxyuroid parasites of 
tortoises (Petter L963, 1966; Sehad 1963). Niche 
diversilleaiiun lias been demonstrated on ihe basis of 
differences in the linear and radial distribution of 
nematode species and in their (ceding behaviour inglu 
H96K) suggested that because of the complex anatomy 
of their stomachs and the diversity of nematode speeips 
harboured within them, the kangaroos represented dn 
equally suitable group of hosts (or studying the 
coexistence of congeneric and coufanulial nematode 
species within the same host Studies undertaken to 
date support Inglis' (1968) suggestion, but have been 
limited to the red kangaroo, Mittropus rufu.s (see 
Mykytowycz 1964, Dud/inski & MyVytowye/ 1965; 
Arundel CI ul 1979) and Ihe tammar wallaby. Mtureptts 
ntgenii (see Smales & Mawsmi 197Ku). In (his study. 
the distribution of nematodes within Ihe stomach of 
the western grey kangaroo. Mtumpux Juli$>btth\m. was 
investigated and relationships between this distiihulion 
and gastric mucosal histology examined. Pathological 
changes in the gastric mucosa were also "investigated. 
Hxamination of M. fidigittosus allowed analysis of the 
relationships ul lour confamihal genera, and in ihe case 
of one genus, Ckunitui, the comparative distributions 
of several congeneric species. 


('t)lhi-iiim <»/ muti'ruil 

Stomachs were obtained from ten western grey 
kangaroos. Mcnropu.\ fttli^inoMty, culled at Hattah 
Lakes National Park <34 45'S, |42°1>T.). Victoria in 
November I l >90. Animals were shot, weighed and sex 
and body measurements were recorded Ihe stomach 
was removed rapidly from the carcass, the oesophagus 

Department of Veterinary Science. Imiversit\ i>r Melbourne. 
Puikville, Vie W2 

Fit, I Slomaeh of Af,n n'pmjuh^im'Mty unopened Lines 
and numbers indicate ihe positions ol ligaimvs mid the ordci 
m ;vliidi they were applied 

Anatomical landmarks shown art ihe oesophagus loi, the 
proximal diverticulum oh. lite gastric pouch tg) and the 
pylorus (p) Scale bar 5 CTft. 

fig.' 2 Slomaeh i)f Afefrgfttfl fnhy,Hi<>.ui.\, opened to -June 
distribution ol Ihe principal types t>\~ mucosal suifaccv 
delineated by while broken tines, the glandular mueos:iol 
the proximal diverticulum (di, the squamous epithelium 
ol the saccular and tuhutai foiestoncuh isi, proximal and 
distal to the oesophagus In) and the gastric sulcus (gs), ihe 
glandular mucosa of the iiibular forcsiomaeli (mi. the 
ntuio.saol the gAStric pouch (g) and pylorus tpi Scale Itai 
5 em. 



and pylorus were nod with string, numerous small 
punctures were made in the stomach wall and (he entire 
stomach was immersed in 5D litres of neutral bullcred 
LV>f lormol saline (r>^ formaldehyde!. 

In the laboratory, stomachs, were washed in water 
to remove formalin, photographed and weighed, then 
subdivided with ligatures and opened. Anatomical 
nomenclature for the regions of the stomach lollows 
Hume (1982). The ligatures were placed in the 
following order dig I)- at the level o| the oesophageal 
opening (I), midway between the oesophageal opening 
and the extremity of the sacculm forestomaeh i2> ui 
the junction of the tubular lorestomaeh with the gastric 
pouch (3) and at its junction with the pyloric antrum 
14). Subsequently, the tubular IbresLoinach was 
>ubdividcd by three ligatures (5-7) placed equidistantly. 
The content from each section was removed and 
weighed. Within eaeh section of the stomach, a 
transverse strip of tissue was removed, embedded in 
paraffin, sectioned at a thickness pf ,^/m, and stained 
with haemaunylin and eosm for histological 
examination- The extent of the different epithelial 
regions of the stomach was determined by curling out 
the entire squamous, mucus-secreting, glandular 
(proximal diverticulum), gastric and pyloric regions 
and weighing them Ihe .ivetage weight per enr ot 
eaeh mucosal type was determined by weighing 2 cm 
portions of each mueosal type, and the area GCfiipted 
in each stomach was calculated by dividing the two, 

As a control lor histopathologic^! examination, a 
single adult M. (ttlti;mostts irom Healesville Sanctuary 
killed foi other reasons, was examined. This animal 
had been treated regularly with anthelmintics lo remove 
nematodes prior lo death by intravenous injection of 
a barbilurate Portions of gastric mucosa were fined 
immediately m 10 V? neutral builered lormol saline (4% 
formaldehyde) and were processed tor histological 
examinations as described above. 

f'ara\iu *hn>U:al <>hs< nttnftotf 

The total number ol nematodes in each section of 
Ihe itn stomachs was estimated by a dilution technique 

(Clark el at. 1971} and the number \)i' eaeh species 
present determined by clearing all nematodes in the 
appropriate subsample in lactophenol and identifying 
them to species under a compound microscope. 

To establish whether or nor parasite distribution was 
affected by this method til' fixation, two additional 
stomachs collected were ligated with string at gift points 
along their length before being immersed in 
formaldehyde solution. 

Representative specimens of eaeh nematode spectes 
have been deposited in the South Australian Museum 
(SAM). Adelaide, Nomenclature of species of the 
genera LahiOftirotyyU® and Chnteitut is currently undei 
revision, For this study, the species names applied arc 
the same as those used b\ Bevendge & Arundel 1 1°79> 


(jrttss anatomy of the stomach 

The gross anatomy nf the stomach Of Af, juli^ittosu.s 
(Figs I. 2) resembles that of M yjyanteus (Langer et 
al WHO. Del low 1982 i Wet weight of stomach 
contents (Table Ij in individual sections demonstrated 
that Sections I and 2 (= the saccular torcsiomach 
I Lunger et al, I9H0]) constituted 22.3% o\' total 
stomach weight, sections 3 6 constituted 73.59; of total 
stomach weight ( - tubular forvstomach), and sections 
7 and H constituted 4.3% ot total stomach weight (— 
hind stomachy J'hc gastric sulcus was prominent and 
extended halfway along the tubular lorestomaeh, 

Relative surface areas occupied by different epithelial 
types (Table 2) demonstrated that squamous ane 
mucus secreting cpithelin were the two predominant 
ivpes. The proximal diverticulum *>f the succulat 
forestomach was lined by a distinctive, glandular 
epithelium. The remainder o! section I and section 3 
of the stomach is lined by squamous epithelium. Apart 
from the gastric sulcus which is covered with a 
squamous epithelium, sections J lo 6 are lined with 
cardiac epithelium Section 7 has an acid-secreting 
lundic epithelium, while section 8 is lined by a 
distinctive pyloric epithelium 

T.aui i i Wet u-rh^v ,>f ttmh-rU of eight dtflerctu teutons at ihe sumach of wn Mucmpus lulieiTtosus///<m fhiihth lakes 
\:)ri"ttnt i'urk, ii\i<.»ta. 

Proportion of 





Total Wumhl 


Section No, 

weiehi (to 




1 2 

LpiuVhul 1 Vfv 





s (siHi.illliillst 





IK 1 




toy -34* 



tn (ntucouvi 





21. B 










71 774 









g gastric) 






p (pylorui 


2 1 5ti 




- t or key io stomach section*; ..-> Fly 1 

UlSfklDirriON n\- Nl MAiouirs IN K\N<.AKUi'S 

TAB1 E - Ami, SS, u prtrctiruxi i»/' the u-ttif, c.vN/w'//" 
dtffvrtJl) rjHlkriuji swtfWJ in tfu \u>?tutrh <<\ nn Mucmruts 
f hliginosih. ,/ftWM Ihtfttlf) htkti NtifiotMt fltfA. I'nti/tVi. 


l"hil S.'t»mith 
\irn On uf'hil 






Mtiuts <ccreline 





UiliWtnr gastric 









HiMflo.KUiil fivmn's. $f en.v/nV ephhciui 
Squamous epithelium 

This epithelium ranged from 80-330 ftm in 
thickness. Basal cells in this region were irregularly 
cuhoidul in shape, nuclei WBlfi relatively small 
basophilic aiul had a prominent dark nucleolus The 
cytoplasm was compact, eosinophilic and eel! margins 
were indistinct- In the mid-region of the epithelium, 
cells were enlarged, cell margins were readily 
distinguishable, irregularly euboidal or polygonal in 
shape with an enlarged, pale nucleus and prommcnl 
nucleolus. Marginatum of chromatin was evident in 
most nuclei. Towards the lumen of the stomach, celts 
liecame squamous and the cytoplasm more densely 
eosinophilic. The nuclei were indistinct. On ihe surface 
itself, flattened keiatinised cells were visible sloughing 
into ihe lumen, and in several sections, a distinct layer 
ol adherent bacteria was evident, closely applied to 
the superficial keratimsed layer ]*hc squamous 
I -pnhHium was folded, with tnierdigitauons of the 
lanuiia propria protecting into the base of the folds 

Cardiac epithelium 

This epithelium was up to 600 ;uu thick ami w\*s 
composed of elongate, parallel glands 45 ;un in width. 
The cells lining the glands were euboidal to columnar, 
25 /tm in length and 10 jtm wide. The glands consisted 
ol two distinct cellular components. The cytoplasm of 
ec-lh at the hase of die glands was family eosinophil!' 
and had a foamy appearance; cell boundaries were 
distinct. Nuclei were situated at Ihe bases cd 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 eosinophilic and the cytoplasm had 
a granular appearance. Nuclei were large and rounded 
with a prominent nucleolus and punctate, emargirmte 
chromatin Surface epithelial cells wete longei and 

more slender than cells within the glands. The lamina 
propria was very narrow and was bounded internally 
by a broad lamina muscularis mucosae 

Epithelium o\ the proximal diverticulum 

This epithelium of the proximal diverticulum of itn 
sacciform loresUmtaeh Oil feed Irom that of the cualUk 
region (Pig. 3). The epithelium was. extremely thick, 
up to LS mm, and was composed of elongate glands, 
up to 1.55 mm long and 0.4 mm wide, which became 
suiiinus towards the base. Cells lining the glands were 
euboidal in shape, approximately 15/mi by: Ktym in sue 
with a highly eosinophilic cytoplasm. Tile nuclei werc 
small and were situated at the hase of the cell -with 
emargination of chromatin or with chromatin 
distributed in a punctate pattern, and a small nucleolus. 
The lumina of glands were dilated and contained a 
highly eosinophilic fluid presumably secreted by ihe 

Fundie epithelium 

This epithelium was ) 55 mm thick and was 
composed of parallel, elongate glands. The histological 
appearance of the glands w-tt* similar to that found in 
other mammals, with elongate surface cells, euboidal 
mucous neck cells at ihe anterior ends of Ihe glands, 
and chief cells and parietal tells towards the base. 

Pyloric epithelium 

This epithelium was 1.2 mm thick and consisted ol 
groups i»f very long slender glands curving tnvvard 
towards the stomach lumen, with slightly sinuous 
raises, Cells at the basal region were euboidal to 
columnar, with pale, foamy, eosinophilic cytoplasm, 
prominent nuclei wnh emargination of chromatin and 
a large nucleolus. In the mid-region of the glands, cells 
tended to be low euboidal in shape with a more 
eosinophilic cytoplasm than cells at the base. The 
lamina propria was narrow and a prominent lamina 
muscularis mucosae was present immediately below 
the base of the glands. 

Histopadrologieal changes in the eprthelia 

In spile of the large numbers of nematode parasite?, 
present in ihe stomachs of kangamos no ;'!<<* 
Pathological changes were observed in the mucosa 
The squamous epithelium demonstrated few 
histopaihologieal changes. Occasional small clusters 
of lymphocytes and macrophages were evident in the 
mid- region of the epithelium, or in the lamina propria 
tf'ig 4). Eosinophils were occasionally prominent in 
the lamina propria. Accumulations ol inflammatory 
cells were visible, most frequently close to the junction 
of the squamous and cardiac epilhclia. 

The glands of the cardiac region were intact histo 
logically and no developing -icmaitKles were seen within 
glands. The lamina propna. however, was diffusely 



r /' .1=11 



V.* *-V : 

> ♦■ 

; # , 


> • 

® s 


# S 

G '% 









**m *-' 




inlihnjicd with mononuclear cells and granulocytes. 
principally eosinophils, with occasional large, local 
accumulations ol 'celts {Ftg£ 5, *5) The submucosa was 
also uniformly infiltrated with the same inflammatory 

The proximal diverticulum invariably exhibited 
prominent infiltrations oi' the lamina propria with 
mononuclear cells, predominantly plasma cells, and 
lymphocytes, with a few eosinophils, sometimes 10 the 
extent that inflammatory cell accumulations obliterated 
individual glands il-ig 3l. In spile of die sometimes 
intense but chronic inflammatory reaction, very few 
nematodes wen: seen within glands Those encountered 
were within 01 just below the epithelium and were 
surrounded by large accumulations of inflammatory 
cells. The limdie mucosa exhibited 130 signil leant 
pathological changes. The pyloric mucosa was 
histologically normal in most animals, but in several 
kangaroos, the megalo-schi/onts of a eoccidium 
developing towards the base of the glands obliterated 
numerous glands m the vicinity As common as the 
schi/onts were areas of necrosis and inllammalon, cell 
debris indicating a site where a schi?on( had ruptua-d 
to release mero/oites. An intense inflammatory 
reaction surrounded megalo schi/onts, characterised 
by large numbers oi neutrophils and eosinophils, 

NufllbfKS dud distribution bf nematodes 

The numbers of nematode parasites tound in each 
o\ the 10 kangaroos examined are shown in Table 3. 
The distribuiion of nematode species ,m\ genera in 
terms of absolute numbers in each section are shown 
tn Figs 7-K. Hxprcssmg the results as densities (worms 
per g of stomach content) produced similar results, with 
the sole exception tif section 1 of the stomach, in which 
densities of Cfoutimi spp. were higher than m othei 
sections. Distributions of (he genera differed. Cl/mitUi 
spp. occupied the anterior sections ( l-4i of the stomach, 
lMhi<t.str<>ii\>\lus kiittgl and /. cl. hiftapillosus isensu 
Bcvendgc & Arundel I97$J occupied a relatively 
restricted region in the third segmcnl. and R. uusimlis 
and P pearsoni occurred m seclions 3-6 of the tubular 
lorestomueh, Very few nematodes were detected in 
regions 5 6 of the tubular forestomaehs. In the gastric 
pouch (section 7), the only nematode species 
encountered was hfunncmu twsirtifis. This is a new 
host record E ttusiralis was found in I wo animals, with 
iwo nematodes in each animal No nematodes were 
found in section S. Comparison of data from these 10 
kangaroos with those from the two animals whose 
stomachs had been ligated prior to fixation revealed 
no differences m the distribution of parasites within 
l he stomach. 

T\bu 3. \>nnihcrs of helminth parasites in the \unntuli\ often Matropus fulu?iiiosii.v l>><m Ittittuh htkr,\ \dtiottu( I'ml-, 




Kangaroo No, 

CUmami spp. 







37 iWO 




M 000 



i ,400 


80 870 






































2 Its 


S3, 330 


Standard Krror 

Qf Mean 





Figi 3-6. Hislopalholngieal changes in dilteienl legions ol ihe stomach ot MaCt'npUS filliginmus associated wiih nematode 
parasitism. 3. glandular epithelium of the punelal diverticulum show ing dilated lamina ( I ) ( filled with eosinophilic secretion, 
and massive infiltrations of lyuipltocyles and plasma cells between glands; 4, squamous epithelium (s) tit Ihc sueeulat 
forestomaeh with minimal inflammatory change: 5. base ol glands of cardiac mucosa from tubular liireshimaeh showing 
diffuse inllltralion of lamina propria with mononui.lL.ii ( i lis im, museulans mucosae): 6. distal region ol glands o I cardiac 
mucosa showing intili ration of lamina propua v ilh mononuclear celts. Sealc hais: I ig. 3. I0ain:>ig. 4, 100 ppu 

Figs S. tS. 50 M|ft_ 



Cloacina spp. 

Labiostrongylus spp. 


| 700 

5 600 

6 500 
2 400 


1 2 3 4 5 6 7 

Stomach section 

2 3 4 5 6 7 

Stomach section 

Popovastrongylus pearsoni 

Rugopharynx australis 


12 3 4 5 6 7 

Stomach section 



<2 60000 

O 50000 
«• 40000 




12 3 4 5 6 7 

Stomach section 


7. Distribution of genera of strongyloid nematodes in eight regions of the stomaehs of ten Macropus fuliginosus from 
attah, Victoria. (Bars represent standard errors of means.) 



Cloacina expansa 

12 3 4 5 6 

Stomach section 
Cloacina cf. elegans 

1 2 3 4 5 6 

Stomach section 
Cloacina cf. magnipapillata 


Stomach section 

1 2 3 4 5 6 

Stomach section 
Cloacina obtusa 


100" -r 

Stomach section 
Cloacina sp. 



E 30 

d 20- 

12 3 4 5 6 

Stomach section 

Fig. 8. Distribution of species of Cloacitw in the eight divisions of the stomach of ten Macropus fulig'mosus from Hattah, 
Victoria. (Bars represent standard errors of means.) 


p. PAMMI M | w vi KIOGI: & R. fi l.a.sM I-: 


Tin gastric jiudirrn i>1 nuerupndids was lev !>• wo I 
by Hiime IWSSj The anatomy and histology q| the 
stomach ofthi: eastern giev kangaroo, W, nJguutctih 
has been described previously (Scbak-i jfi Williams 
lX7o Lyngei ei ui IS8Q; Dcllow & Hume NM) 
However then.' is no comparable infoimution lor ihc 
closely lelatcd species Af Ju/i^ino\u\. In addition, 
whil.- the histological mid nlUasutKUiiLiI features u! 
ihc stomach C)| ihc Umuiiir wallaby, Mot ropus ntxenii 
lu\\c been described in detail (Genimcll & fln^efhardl 
W*?7i. ClaUl lor ihc eastern grey kangaroo llPC 
limited. Wc examined pathological changes in 
(he mucosa associated with nematode parasitism and 
the distribution of nematodes in mlainmship fo 
epithelial [yppa anil therefore an examination of both 
the cross and microscopic leaiuresot the stomach of 
A! fulfgjntfteit was necessary. In Ibis study. Ihc gross 
.mammy pi the stomach of A/. juilgintl&HS did not differ 
significantly from thai ol M fii^attteu.s- Proportions 
!n wvijtfti or ihc three principal stomach regions ifl M 
fulttitKwt: -i' UlaJ '<<a-stoittaeh lubular fotestomach 
and hilid *tftftacll f22% , H'% and k v A j, resembled ihai 
ol M v.ixamvus lIoslK iM-;. /tri and 7^a The 
historical results 1mm this study provide additional 
information on gaslru anatomy of grey kangaroos, rhc 
iruisi - ^iMt'kanl difference from earlier works is Ihc 
recognition thai lhe mucosa of the proximal 
diverticulum of A/, fallgftoffiun is disimcily different 
histologically from that ol the cardiac region and 
■ imimms o| elongate parallel glands producing an 
eosinophilic secretion. The vitpe r. Hue for M, 
\>\uMnt<us, although lhe histological differences have 
mil been reported m ihc literatuic and both CtqnuUih 

- "us have been considered to have an identical 
mucosal type (flume 19ft!), a conclusion which would 
appear to be erroneous, Although anatomically 
distinctive the function ol (his isolated <UVt< of 
glandular mucosa is unclear 

Anatomical lemis applied to lhe slotnachs of 
kangaroo*- arc not consislent (see Hume I9K2) The 
system of nomenclature suggesied hy Kichunkon fJ^XO) 
as conforming most < loscly til Nummn Amwmiho 
lit'titutrsa is suitable for descriptive purposes, bin is 
nol wvll minted to nii!*_nonal studies. The terminology 
-i.ii bj Hume-:i"Sri ... which the stomach is divided 
ifltO uceukii lofc-iommh, tubular forcslomaeh and 
iMiuhi'irriuch > N more appropriate when dealing with 
the distribution of neouitiHles and ol epithelial rype% 
and has therefore hcen ufthsed in this studv 

neicrnnnittg lhe position ol netnalotlL"-, wiUiut ;ui 
otjMti clearly leqoires very rapid fixation or 
immobilisation o! rhc neni^iodc.-; fnllnwinjj »iv. deaiK 
ol lhe NW VVith small host :>pecies t this \y.\s bcCfl 
adoL'ved UHittg rapid rrtY.?m<: i-uhad I0p>?| <%: 
Holmes I'JNo.tA.r mp.d immersion in lo, > i '■.'ofute 

NH'.H, Willi large hosts .such as kangaroos, the technical 
difficulties ate grajitfit The methods used in the past 
Ituve been ligation ol the stomach immediately 
following the death ol the host in lhe laboratory 
(Smales A: Mawson l*?78b) or ligation tollowm^' host 
death in the field iDud/mski Ai Mykyiovvye/ Mtr)5; 
Anmdcl fl ul W7v>| Ditficulliev occur in accurately 
idL-nt'tvinv .maromical "kuidmarks" and ptacuii', 
lioaiures on stomachs under field conditions and 
therefore ihc approach taken here was to rapidly 

nefse the enure stomach m a large volume ol 

fortnaJdehyde solution and carry out its subdivision m 
the Uio"rat*'i> Thai this did produce an accurate 
rcprvsertuilion of the longitudinal distribulion ot 
nematodes was confirmed by comparison with data 
from the stomachs of two kangaroos which were limited 
in the field prior to fixation. 

In spue of technical limitations, each of the nematodt- 
eenera exhibited fl restricted localisation within lhe 
-.loniitch. Data have been presented (Fig. 7) as absolua 
numbers of wonns tn each stomach section lather than 
nimibci of worms per gram of stomach couteni 
(density) 'lhe only uenus lor which the disttibutiou 
pattcT'!1 is allcred in comparing density ralhcr than 
attsulute numbers is Cioadna because of the high 
ilensilv ol nematodes m section I of the stomach 
Chmanu 5pp. were located primarily in the saccular 
lorestoiu.cch with maximum densities in Section I 
htht(f:\fr"tt^\lu,\ spp. were also presenl in lhe saccular 
fore-stomach, hut maximutii numbers occurred in 
secliun 3 at the anterioi end of the tubular foresmmach. 
Both Raiiopharyia austraii.s and P<fpo\asirtm^\lu i 
ptitrsoiii occurred principally in the tubuU< 
forestomach and all stnuigyloid nemaUKies were abseni 
from the hindstornacti (sections 7-H). an area of the 
stomach charactctiscd iu macropodids by a low pfl 
(Dud/mski & Mykytowycz l%5; .Smales and Mawson 
V)lHh), and inhabited ofllji by the Utehoslrongyloid 
ecnu.s t'tUuiin'tnu. 

1'hi d-sirihuiion •(! uematudes within the siomath 
ol M fh/iv>ttK.'SU\ is similar to that reported for other 
kangaroo ^X-des. Smales •& Mawson (1978b) lound 
that ChKtiim spp. vvciv most abundant in the saccular 
forcstomach of M. ruxenti while 8, aiLsttufis was ntosi 
abuudani in the lobular foresiomach. Likewise, 
Aiundel fi ai. (1979) lound that maximum mnnbi -r. 
of H, tiitsmih* occurred in lhe tubular lorcshnnueh ol 
M ruJH\, The distribution of hthi< >\m <ti,i> vIh,s spp. was 
reslncied in M. fiiliiimosus (mainly in seciiiHi 3) when 
Compared with data from nflici hosts iDud/rnski & 
Mykytowycz !9fO, Smales & Mawson 1978b) bin this 
mas be related in parasite density. The numhet ot 
Lubnf\m/iti>\ins spp. in the piesenl study was low 
'mean - 120. lahle ?>\, Dud/inski & Mykytow>c/ 
(1465 1 divided their specimens of A/ rufiis inlected 
Witt /- ton^tsph:uiaris into "heavily infected" ( >20(J 
worms) and "lightly infected" (<5(»0 wonns) hosts. 



They showed lhat the maMtuum numhe* C»l i, 
h'twlsfntttlitfi\ always occurred in the tubular 
Ion-stomach but that with increasing abundance, the 
numbers of nematodes in |W succulut forestomai h 
rose. It similar iiiechanrsms operated in A/ (ult^iwmt\. 
ihcil higher parasite densities would he cspected in 
result in a Mendicant increase in parasite iiumhcis in 
sections 1 anil 2 ut the stomach P. pWftmi in our 
study occurred predominantly m the Ulbifbrip 
loresiomaeh (sec-nous 4 and 5) but in M *'uvftiii lite 
same species occurs primatdy in the sacciform 
loresiomaeh tSmalcs & Mawson ffl9H\ The reasons 
(or this difference are no! known, iHtt It mn\ r> n.\ i 
diltcrcuccs in the anatomy ami physiology ul the 
stomachs ol the etcy kangaroos (M. gi$amrt{.\ and M 
fuftyjtvt.sus) when compared With thai Off M vi*>:t'nii 
(see Kiehaidson KWQi I-anyer .7 ,//. I9S&; Dcllow 
b)X2; Dcliow A Hume 1982) ot may be a ftmClttW &/ 
host associations /' prnntmi is a common and 
abundant pa i us He in M. tltg&tti (see Smales & Mawson 
LSTBfc), Its distribution in A/. ju/ii' is uuichmoie 
restricted, bctuy kntmn only from Kangaroo Klaud, 
Sonlh Australia tHevcndee $86). KershnH.k. South 
Australia (Wiesner. unpublished 1 ) and HaUah. 
Vtctoita. Ii was not encou rite red in a survey ol the 
parasites of 40 Sf tufn>in<>.<u,\ by BcNcndpc & Aiundct 
(IW/U). A7. fnh\itt/it t stf\ m;t\ N- an abnormal or unusual 
host lor the parasite and bus may be reflected to •' 
differing distribution of the nematode in the stomach 

The distribution ol couecitenc*putasite specie-, was 
not investigated in (be case o\ the two species ul 
Luhtastrort.Kyho because "I me if relatively restricted 
distribution within the stomach. Ul the sevetal sp a 
ol Chuultta ptesent. onlv three. C t'\puN\ti, (" 
hvtfnformis and C. vi. eU-^nns oecuired ut sufficiently 
high densities !o permit comparisonv- tn each case. 
there was no evidence o! dillei my dtsti that ions Eui h 
parastie had maximum density in Ihe blind sac ol 
the saccular loiestomach. One Clutuifta Specie* 
occurred in sections I to 6ol the stomach, luil tn vci v 
low numbers only The data therclbre suj^pesi that there 
is no or very lillle tceional scpuomoM wilhin the 
stomach when several congeneric spec teg arc piv-enl. 

The lack ol dillercuces tn the distribution of 
t 'otijvnerte species ol Ctouafiu contrasts wtdi studies 
i»ti the oxvmroid nematodes ol tortoises fSchad R?W; 
Pellet I%M in which each nematode species was 
shown to have a resit icted niche. I he results obUined 
here, with no apparent ntche see.tvcvition. aiec\'ustslenl 
with the hypotheses proposed by Rohde (19801 for 
mouoevnean jiaiasites ol lisli. m which lestucted 

' D. Wiesner HKl t.'ompanson nt hi-hnmth parasite 
burdens in wcjtcrrj vrcv kai^aiy^ Wfl 1 ><>pny(tilis , -' h '- yus 
iDcMii.iresl, !K2J» grwing tiiwuml. inipiTveiJ UTIU de^r.ided 
|jjisiuie' ( K".v\M"ilu Avi)(.u!iuivl College. utlpuWiofrcd 

niches were absetii but allegations oi tonspeettic 
paiusiies weie assumed to tKcur tn onJcy to laeilime 
reproduction iKohde WD. Competition W|$ nert 
considered to be a tJgnifi«im eliunenl in dislnhution- 
iis niches were ^ilipct abundant and wete not saturated 
In the spcchueits of M, fttfiyjm^tis examined, the 
numbers ol nematodes ptesent ^m\ kangaroo were low 
when eoutpatcd with previous studies {Wevendec &. 
Arundel W^: Wicsnet. unpublished') in which tolal 
nematode numbers ranging from ."UHMXIU u» 500X100 
were encountered in some kangaroos. These data imply 
thai the numbers oi niches iivailablc loi most species 
ot v;istnc nematodes m Af. firflgtHPSUS are large ami 
it this is the case, competition would not he expected 
it) be a major factor affecting nemaUvJe distributions. 
A far larjzer sample of kangaroos should be examined 
lo test for evidence of imeravlions between component 
parasite taxa lHosle& Cabaret 1992). However, littir 
or no evidence o\' competition has been found in 
stomach-inhabiting nematodes in M ntfu\ t M 
,if/,et//Uews or Pi'troyate spp (see Hosle &. Bevendge 

l'W J 0. 

Histop.ahoK>eical changes in the gastric mucosa wete 
restricted h> ddfuse infiltrations of mononuclear cells 
in ihe lamina ptopria between glands. Changes in the 
sc| miliums epithelium were ne^Hj'ible. The 
mouonucleat cellular mltltratuuis were diffuse ami 
there was no apparent tclatiouslup with nematode 
distribution. Cellular changes m ihe lamirui propria 
were most marked m the luhulat torestomaeh. and ii 
was here that the greatest numbers of nejnalodes wen. 
found. However in the saeeulat toieslomueh where 
(loadiui spp. were dominant. Ihe marked changes jii 
the glandular epithelium compared with the lack ol 
change in the adjacent squamous epilhelium suggested 
that the type of epilhelium present siptilicanth 
inlluenees the extent ol pathological L'hflngM seen. In 
Ihe luudus. an mea esscniiaJly devoid of nematodes. 
no significant patholoj'ical chauves were lound. 

In the pyknic antrum, focal lesions detected were 
caused by selu/oiiis of a species of Etmvrta Although 
a number of species ol this t'entis are known to occur 
m M. fuliytnostn (see Bui ker ft ol. I989J, the sprrn •. 
present in the p\ lt>rus et>uld nt»l be identified, because 
ontv the oocysts excreted m lijeeeshavc beendesctibcd. 

In the study of the pathological changes induced in 
the stomach ol At yjytitih'iis by nematodes. Arunde! 
t'( at, (I'WO) eotumented on macroscopic lesions caused 
by Uthtt'.\{rr>nKyln\ >pp. . by SthtH\;\tofilt;s sp. and by 
KitvttpfutrYtiA tt:\%'mim<w\ None ol tlie lesions thev 
reported was found in M. fuli^iiu*su.s ul Mallah. 
although u_ w evident that the presence of t Tnatiha spp. 
and rV. anstralis in the stomach lvsulted in a diffuse 
ga.slritis similar lo ihe itiliamtnalory cbauyes induced 
hi strnnjfyloid nemalcnfes in the l;irec inicsoncs *a 
eqiiids .Marker $ van Dreumel \^\ 



Hit mcun intensity of inlcelion of kangaroos with 
r»aslrir nematodes ut HafUth (Tabic 3) (mean 87.2(H) 
nematodes) was stmiJ;jr in that (45,2001 reported Irorn 
40 iivtMMnging M; MltftiHOSltf collected in Wcitefin 
ViiUmu and in South Australia by Bevendge & 
Arundel (1979)'- Given the high density of kangaroos 
at Haliah al the lime ol the collect ion ol 78 per -.tjuaic 
kilometre (Morgan 1490), it WftB considered possible 
that parasite abundance- might also be h&htfT. its occurs 
in populations: of A/, wwuflwts (Arundel pi Cff }990), 
bin IhlS was not Ihe case. A more details! study ol 
ihe epidemiology ol nematode inlccliuns m M 
f«Hxm<>sti,\ in Wi stern Victoria is ivipiin-d before the 
significance ol ihe cut rent data can be assessed. 


VW wish to thank the Victorian Department ol 
Cnn.-.L-tnitmn and huvioumteiu lor the JiivilutuM to 
Lvtllect kangaroos al Hatlah and tor assistance wiih the 
Mitlcuuui We are paiuculairy indebted to B. Wallet 1 - 
and (1 O.ulson tor their help 


AtiiMiti, J H . ttrvrwinoi I X Pwisiimno, I' I ^ 
(t974) Parasite;, Tind paihoiop.ical findings mi-nclosndaml 
tree ranyan^ populations ol Xhntupits tufas (I h-suuncst) 
(Miirsupialial at Menindtc. New South Walcv Au.v. WflJi, 
fo\. 6. 361 .W 

. Dl MCSU K. K I, H\WK1.,AN K r \Hi.\tK.R 

(1990) Ppidemioloiikai observations On Ihff helminth 
\,t\\\tiKi\\ Mn< tt>f>u\ i;ii!iinh>is Shaw in Vieloria. Ihitl 17. 

39 'I 

Barkih, I k.- O'Cai i viiiAN, M 0. & Ut vi tou.i , I, 
i|0H*)| Host-parasite assoeiations ut fclmiftll spp. 
( Apicomplexa; Riinerudae) in kangaroos and wallabies ut 
t he Mnu^MdcrrtpdA (VUusupialtu: Ytueropodidack hti V. 

J\l'nt9fitA W, 24I-2M. 

& V\n Dun vim . \ A (NHS, |hf Alimentary 

svml-ih pp, 1-2*7. Vffl, 2, In .lubb. k. V l- , Kennedy, B 
C. & PaluiL-i- N it-:ds) "Pathology ol Domestic Animals 1 '. 
3rd lido. (Academic Press., londoni 

Bi-vmuuck I. (WXCo New specie and new retards ol 
f{tfnntn/nm\:\lu,\ Muww<(i, 1977 iNernaloda: Cloaeininae* 
Irom Australian marsupials. Buff. Mux nut. HtxL nut 

Hirh, rfwnt j v& 8. 257- 2<o. 

iV A,ftl NOM , J. B (197Q) Helminth parasiles of gjcj 
kangaroo?;, Mom'/Mis gifiCtrtf&Wi Shaw and Al futiginosu.\ 
il.Vvmaresli. m eastern Australia. AUHH Wikfl. $X fi_fW77_ 

bl sh (W 0. & rl>JLMWl I C I WSftl ticlmmUisol 
levstr '.taup ducks- pnitcrn-. vi! assoLUlii.n- CVtfi, / /<W 
fi4 13: Ut 

Ci uk, C- I- ^ ' M-fe, A M. & TiikTOV.. J. A. (|U7H 
BtfUiplinj! leetini'.)Uf tot i^tinintinp round worm hurilctTi 
t)| vlicofl and caule /.\/», Porusiwl, 30. 181 t^6. 

Djilow, l> W ils»82) Studies on the nuliinun of 
mat-rnpttdiiie inai>apiats. III. The flow ol "dipesty llinniyh 
itie sloiiiaeh uixl inieslinc p| liiucio^dines ,\r\\\ shc-ep. tiu&t 
7. Siw/. 30, 75 1 76S 

<V Hi Mr, I Li ti&jQ.) Studies on die nutriiiiin ol 
niaLTorxxline marsupials IV Digestion in (he storruRli and 
the intestine ol XUu repus gtgatfft'US, Thyn)gah* ih<ti\ «nd 
Mutnnnt\ in^tiiii Ihid 30, 767777 
Dl o/tNSKi. M I (ft MYK^TWWVC/. K (l%5) Disuibiiiioii 
of ihe iieiiuiode Luhu}\troni>vlu\ hntiii\fiitul(iri\ (Wood) 
within Ihe ^lomaeti of the red kangaroo ftfi'gmlpht rujii 

GiMMtn , R T, & l-NLHiufcrvi W V. (IV77) The structure 
ol Ihe cells lining ihe stomach of the tammai w.illahv 
(Mtitnyw.s cuyimi) J AtiuL 123. 723-73$- 

HttsiK. H. A; BlAlRloor, I 1 199_"> J lnterspeLific ami 
ififcerjlcTtrric rrlatioiwlups between nematodes parfliilic iti 
the stomai tis o! kanv'aroo.s. haitx ft. Soi S. Um Jl7. 

C V C \\t\Ht i ,f. (ITCt Inic-ryeueric lel.ittons between 
iiemutodex ol liie dit;e.slive trael in lambs: u mullivariaif 
.ippio.ich. fat J Pi)'\tsiu>f 12. W 1 * IKo 

Hiimi-. I. I> llyH^i ' v nij!esli\e pliy^ioloyv and nutrition ol 
mil ii fiaU Monoyruph^ on Marsupial Liioloyv " 
(Cumnndge C'ruver^iiv Mre&s. rambrid^et. 

I\ra is, W G. ilMnXi The yeographiejl and evoIulionai> 
rclaiionstiips of Australian irieliosuonpylnid purasiies and 
(heir hosts. A fj«fl, S<n ff)<1l J 47 52T-347 

I.AM.r -b. P Hi mow, D W, & HI mi , |, IX ]Wty S(oi,i,uli 
slrueuire and funetion in three speeies of mactopodine 
marsupials. ,t//.v/_ A ?jsof 2K. I -IK. 

Mohoan. 6. Q. (WO) Hunan tsulkyne - Mournpull Block 
ItJngflWO Hopul.ilioiis. Nowmhei IVVO. Report to the 
Division ot Na-Oonul Parks and Wildlife Department ot 
Conservation and Lnvuonmenl. Victoria. Centre lot 
houram Evaluation, Instmue oMidueation, UmveiMt.v of 

Mvk> 1 1 mm. /. R. (l%4l \ survey of Ihe eud<»parasiies oi [\\t 
red kunui.roo Xhyik'iti rujo liicsmaresii hh'ttsftoh^ 54, 

Pl'TTlR A h il%3> rujuilipre tle.i espcecs dans lev 
populations de nemaUntes parasiles du colon des lorlui's 
Icricsttcs, t_ r flchil St'anr- Aaid &'/., ft/nc 257. 
:i5.' 2154 

(106m |x|uilthrc des especes dan^ les populations dc 
neruatodes ptirasites du colon des lonues terrestres, Mitt/, 
X/u\. nut- Hist nut., htti\ Sent* A. 2uoL 3M 1-252. 

RioiAKosov K. V (WHO) The structure and radiographie 
anatomy ot the alimentary naet ot the ra/timSr wallahy 
\1um<pu\ *tii>t/tii ( MurxupialiaJ. I. The slomaeli. Au\t J 
A»>i 28. 367-574 

RohijI-, K il l »77i A nouetimpeiitive mcelianisiii icsponsiblc 
for rcslneimi: niches /,tol in:. 1W. It4 172 

&)W) Comparative studies on mknihahitai utilisation 

by teiopariisites<»l SOlTlO marine fishes troni the North SCO 
and Papua New (iuinca. ///»/ 204, 27h4. 

SiiiM>. G A. (t%S) Niche divcrsifiealion in a parustii-. 
species (lock Nuttttv (Londoni 198. 404 400. 

St hah K. H, \ (ft Williams, D J (IH76i On the struciuu- 
ol the mucitus uiemhrancs i\\ ihe slontach in ihe kangaroo.-. 
Pnn\ /jhfl S«/f, iothl, !87r.. 105-177 

Svm.i.s. R I.. & Mawson, P M. 0^7Ha| NVmntodr 
I'ausitx's ot ttiL- Kait^'arito Islam.) Walluhy. \Ut< n>pu\ fn^rnr 
(t^esmarcst). I. Seasonal und gprjgrapbieaj distnt>uiion 
Itvtty H Sot 5, Atot, 102. $U|5, 

Si i!U7Mb) Nematode and orhei ptuasitus ijl 

lIVC Kunuaroo Island Wallaby. Mum/pus t-timun 
(DesmarcHi). 2, Sire seleetion within the stomach. Ihni 
102. 7s»-83 





ByR. C. Ainslie, D. A. Johnston & E. W. Offler* 


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. 

'nwisttcihvo »! tin tiwt Spurts r<fS. Aitu <1W), 118(3), invito 



by R, C. Ainsuu. D. A, Johnston ik E. W. OFM.tR* 

Summary R- C.j Iohnskin. \X A.. & Umt-K, F W. I IW|. C}f t»wih o1 the P> tsuhmift KtntWti dtfvbt'UjgC 
< r Kuo at locations near u>; find remote from, a powet stalkm thermal outfall in northern Spender Gull. South 

Australia, ffzm/ir ft Sfo Si Mh H8(3K 197 ?ih. .VI November. 1094, 

"I he e^ttvtji nl 'he seagra^ frnuh'tua m/ih"mi was monitored m northern Spencer Gulf Soulh Australia, between 
hue ItfSO and l990 at a sue within ihe influence ot the thermal Uiseharge li'mi (he Northern Power Station, and 
ai a Gull 'ambient sue, P uttuew growth was also monitored in Pi Paterson. a large shallow bay surrounded 
by extensive mudflats, immediately io the south of the power station, but beyond the influence nl ihe thetmal plume 

P sinmt.Mi meadows adjacent to the power station show relatively minor reductions fit growth characteristics 
(leal blade niomass. pmdueiiyity, and feat" gflpwtbj compared io lh-^c ju Gulf -ambient condiimns, despite the 
jaet ihai summer water temperatures adjacent io the discharges arc consistency slightly lusher than Gull ambient 
flflfl have reached 28*0, a temperature comparable io the highest held temperatures previously avoided lor Pwdtmui. 

However, in the naturally warm waters of Km Paterson. where average summei water lemncmtures are marginally 
lusher than those within the influence of the thermal plume und where inieinattent peak temperatures exceed 
ty°C. P simuna \v.\s significantly reduced productivity 1 , standing hioniass. blade length and blade gtowih iate. 
typical of Posichmnt species in marginal environments. 

Despite The minimal effects oflhfi current dis'.haree on the seugntsses, (he e\ (dertce from Pt Paterson suggests in northern Spencer Oulf where summer temperatures are more typical oi a subtropical ihan a temperate 
marine environment P sitiu<>Sii is near die upper limns of its tenipcmiuie tolerance during ihe holiest lime of 
die ycai Should there be localised increases in maximum water temncratun-s in the Gull from future thermal 
discharges) >.V)°C'K (here is Ihe fH»tentuil for more widespread occurrence of -.lunied scugr;<sses such as limud 
m the shallow waters ol Pt Paterson, One possibl; consequent vol ihis could be localised increase in the movement 
(i| sediments on the sloping banks of the Gull channel. 

Ki \ Wukos: scagrass grovyth. waiei lemperaiurc. Spcneet Gull 


Spencer Gull. South Australia, extends, ahoin 200 
km inland to the and mid north of South Australia (Figs 
I). Northern Spencer Gulf is defined as that portion 
ol the Gulf nortli j# 33°S. The waters lit the northern 
Ciull art* characterised by high summer salinities, 48 
(Nunes Si Lcnnon 1986), and summer temper-aluies 
in the nhd to high 20s°C (Ainslie ci ai 1989), 

Shepherd (J983) found that the subtidal bcnlhic 
lommunilies were "impoverished in terms ol the overall 
sjiv-i h-s richness" and concluded thai this may be 
iiitlicutoc ot" a stressed hypersaline environment. He 
.suggests that it is critical (hat the component parts of 
(lie biological system u\ ihe norlhcrn Gulf "should 
receive very detailed study io determine its capacity 
to receive additional stresses'*. 

Environment 6c Technology Oepartmrnt. I he hleetncuv 
Trusi of South Australia, PO Box 11 Adelaide $ Ausi 3G(fi 

Haiis, I. K. (19H2) Siihmutine geology, sediment 
transport, hydrodynamics and quuierary hislory ol rtorlheni 
Spenevr (julf. South Australia. Seminar on the Research 
Needs for Management or the Soulh Australian Gulf**. 
Aiisiralian Marine Sciences and Tcelirioiop.ies romniUi-e, 
Adelaide, 10 Nowrrlbci I9ft3, 45-H. UnpilHl. 

The extensive seayrass meadows are an important 
component ot the biological system of the gulf, as u 
habitat and ntiisvry rcuton. Ibr their intrinsic 
conservation value, and also !'«.r the role they play in 
stabilising scdimenls, particularly Ihe sloping banks 
ol the Gull channel 1 . 

In 1955 a 90 Megawati (MWHhcrmal power station. 
Playford Posver Station, was established on the eastern 
shore of northern Spencer Gull, soulh off the city of 
Port Augusta (Big, U: by the I9c«0s the capacity had 
been increased to 330 \]\V. The cooling water 
discharge from ibis development resulted in surface 
water ttmpcralures about 0°C above ambieni near the 
pCiWcr .station, with only occasional incui"sions of the 
water to the interlidal regions (Ainslie el al. 1989), 

tn the late 1970s the decision was made to proceed 
wtih the development ol a new power station oi up to 
a pOS5jh)c 750 MW (Fig. I), with the ultimate potential 
to discharge 4 million cubic metres of warmed seawater 
a day to the Gull, more than doubling Ihe volume Of 
water discharged at 6°C above Gull ambient (Ainslie 
fi al. 1989), and with predictions of localised but 
noticeable increases in temperatures of gull waters 




Deep Limit' of 


Inshore Seaqrass 
Verge JS^* 2 

60 50 40 30 20 10 



discharge: site 1 


Coohnq water Inlet 
and outlet 






Fig. I. Northern Spencer Gull 'South Australia, Seagrams monitoring sites; (1) power suit ion thermal discharge (2) Gull 'ambient 
site remote From the discharge, and (3) Pt Paterson. Inset: seagrass distribution From low water mark to the Gulf channel 
at site I 



Under some tidal conditions short term maxima wefe 
predicted lo exceed M^C m the vicinity p| tJv [JrtWW 

station tuufalls J . 

The reported eKects of thermal discharges on 
seagrasses range from large scale devastation resulting 
Irooi temperature increases only a tew degrees 
Centigrade ahoVfc summer ambient (Thorhaug fl <//. 
I°78). to reduction in seagrass density (Robinson rVJflJ 
and changes in specific growth charade] islics such as 
leaf thickness and biomass (Vicente 1977;. 
Cireumstantial evidence suggeststhat relalively :;mall 
changes in temperature of inshore waters may ais" have 
indirect effects on seagrasses through increased 
bacteria! and luugul attack (Rasmussen ls>77) 

Once seagrass meadows he.gin to delenorate. 
erosion, sillaOon arul reduction in water clarity may 
lead ip a decline in the seagrasses beyond the direct 
influence of the discharge (Shepherd I986r Other 
secondary effects such as increased impacts of grazing 
organisms may also accelerate the decline vi seag lasses 
under stress | Shepherd ft ui. WX9h 

Both Pt>sitli>tiia austntiis and P sittttusti occur in 
northern Spencer Gu If bur the latter species is dominant 
in the inshore meadows near the power stations. 

This paper describes the result of a monitoring 
programme conducted to assess the effects of the 
increased thermal effluent from the first 500 MW 
development of Northern Power Station on (lie growth 
of the seagrass P, simtOMi, in the light ol predictions 
<y( localised water temperature extremes appmai hing. 
or even exceeding, llic upper known field icniperatutes 
foi this speeies'- 

Materials unci Methods 

Preliminary field work was undertaken in the 
10X()-j08? Milliliter season to establish sampling 
techniques. The results of the seagrass monitoring 
programme for the period |^S7-19B? are reported in 
this paper 

Water temperatures and growth ol \ Pt'Sidoruu siiiuosa 
wcie measured al three sites tit northern Spcncei Ciult 
Uig. t). The Guij sties v^eiv established on Mansri i 
lines, Ihc clevalion and location of which were 
cotifirrned relative to an established fVrt Augusta 
Power Station datum as part of a contiguous study of 
the hrnihic wtauna of the northern Gulf jAinshe er 
at. I9K9). Site I is within 200 m of the power station 
and within the influence of the thermal plume ot (tie 
new Northern Power Station. Sile 2 is on the eastern 
side of the Gulf, JJJ km south ol the power station 

LlcUneity Trust of South Australia (WH5i Northern tVuwr 
Station 1-tu Imparl Statement. Angnsi !*)X* 

Prepared i»v fCdnhiJJ Steams, t'npuM 

anil beytind the predicted influence ol Die dischatgt- 
plume 1 . Site 3 is in Port Paterson. a large shallow- 
emhayment, also beyond the predicted influence of the 
thermal plume. The naturally high water temperatures 
which have been recorded in Port Palcisoir can be 
attributed to the insolation of the extensive surrounding 

20 m transects were established a\ each site about 
5 m seaward ol, and parade! to the inshore seaerass 
verge, all at a depth of about 03-0.5 m below I.WP 
(a depth ot up to 15 m during the highest tides). 

Al each site in sift< water temperatures weie avoided 
within the leal blade canopy with portable data locgei-. 
in custom-built PVC waterproof housings, and lilted 
with 15 k thermistor detectors. 

Air temperatures were recorded at the South 
Australian Bureau of Meteorology weather station 
located on the Northern Power Station sue. 

Our object was to estimate several measure- vt Hie 
biomass and growth rale of seagrass. 

A wide range ot sampling t)iiadrai areas is ciicti 111 
the literature fot seagrass studies, depending 6(1 I&C 
leaf blade density and the need to ensure that ftft 
adequate nuinbci of led blades is tagged tor 
measurement. Shepherd (1983 1 used sub samples vA 
50 leaf blades for detailed measurements oi' Ptis'ul^nid 
ttHsmids leal blade lengths and widths in Spencer Gult 
More recently Pollard iV Greenway ( used 
samples of between 20 jr^i 60 shoots in a leaf marking 
study o; the productivity of three species of seagrasses 
in the warm waters ol Curns Harbour. Queensland 
In this study 15 cm- steel framed quadrats were used 
to define measurement amas for each sampling site 
Preliminary collections in the summers of I9H5 and 
l9Ku \ielded information on the leaf blade numbers 
lor the quadrat area. Stxtv nine quadrats, random Is 
placed along the transect lines, were sampled al all 
locations; the mean number of leaf blades quad r,n was 
34.5 + 17. A decision was made to proceed with the 
Held studies using live quadrats per sile (averaging 170 
blades'site) The sampling a tea sue was I}2fj cm'. 
comparable ret that ol a number of other productivity 
studies of sca^iiivi • with a similar growth habit 
i/iemau #74; TJiUrWfi tf ui. 197K: Walker & Me 
Comb 1988). 

VVithm the quadrats. all 6Cdgl k$S 1>IaU03 WCM 'agged 
and harvested using methods oullined in Pieman (1974). 
Harvesting was earned out at high rifle, appimunately 
every si\ weeks depending "fi weathei conditions and 
underwater visibility. 

Total blade lengths, incremental growth (length} wkI 
dry biomassol wliolc blatles and incieinenlal growlti 
(productivity) were reconled lor each harveM. 
Numbers of new shoots, i c shoots which grew 
subsequent u> the uutial tagging, and their gaiwth and 
dry biomass were also recorded A simple "shoot 
index" is derived which is the number of hflft 



shoors/nr (of scafloor area)/day expressed as a 
percentage of the original number of leaves lagged/nr 
tor each sampling site. Blade widths were also recorded 
but were only used as a verification of the species 

Dry hiomass was determined by drying freshly 
harvested blades to constant weight at IU5 D C, after 
removal of epiphytes with a stainless steel sciaperand 
treatment in a 5% hydrochloric acid solution. 


Figure 2 presents a comparison of the Port Augusta 
summer air temperature and water temperatures at sites 
I to 3 during the summer of 1987-W88 with a 300 MW 
power station operating. 

In the shallow seagrass beds of northern Spencer 
Gulf, 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 term 
fluctuations are most pronounced at site 3 in the 
shallow bay of Pi Palerson. and least evident at the 
Gulf ambient site 2. Overall, summci water 
temperatures are marginally higher al site I neat the 
thermal discharge than at the Gulf ambient site, but 
consistently highest al site 3 in Pi Palerson (Fig. 2). 

Al all sites, despite the higli summer temperatures, 
seasonal temperature variation is typical of temperate 
regions with winter water temperature dropping below 
I5°C, even at site I adjacent to ihc power staliou 

The summer relativity of the waier temperatures 
between the sites floes not persisl throughout ihe year 
During the cooler months, Gulf ambient temperatures 
fall significantly below those of Port Palerson, which 
in turn is cooler than site 1 adjacent to the power siaiion 

Productivity (mean growth, dry wcighl/rrr/d) ol 
Positional at sites I to 3, from November l l >86. 
ro August 199tij is shown on Fig. 3. 

At ihe- Gulf ambient site 2, and al site I near die 
power station cooling water outfall productivity shows 
a pronounced seasonal cycle with peaks between 
October and April. Although ihe seasonal cycle is less 
obvious in the seagrasses of the naturally-warmed site 
in Port Paterson, highest productivity values at this site 
were recorded in summer (Fig. 3). 

Productivity was consistently highest at the Gull 
ambient site 2 and least at site 3 in Port Paterson. with 
highest recorded productivity being 37 ± 1.1 g/trr/d, 
3.1 ± ().*) g/nr/d, and 1.6 t 0.3 g/nr/d at sites 2, 1_ 
and 3 respectively (Fig. 2), 











- - - -- 

Vr ■ . 










■15 ■». j . , » --.■ „ „,■■,■■,■ i ■ ■ ■ , . . . , 

: f||nife : fwl :: ; ::l::::: 







Kig 2 


I. J si 

Maximum daily air temperatures al the S.A. Bureau oi Meteorology Station Port AuiiUsLi, and ihree hourly maximum 
r temperatures in the seagrass canopy tit siieh i, 2 jmU 1 inset: water temperatures, lint's ol best lit (polynomial 
SqLiarc$ regression) at .sites I u» J. 



Productivity data for each site were compared, no 
a priori attempt being made to discern seasonal "cut- 
offs" Variances were not homogeneous (Bartlctfs test, 
P<0.01), and analysis of variance was therefore not 
used. However Kruskal-Wallis analysis showed 
significant differences between all three sites (P<0.01). 

the greatest difference being between sites 3 and 1 and 
between sites 3 and 2 (Fig. 3). 

Maximum mean standing biomass also occurs in 
summer {Fig. 4). As with productivity, the seasonal 
cycle is most pronounced at the cool Gulf ambient site 
2, and least pronounced at site 3. The highest standing 

Jan-87 Jun-87 Nov-87 Apr-88 Sep-88 Feb-89 Jul-89 Dec-89 May-90 

Site 1 

Site 2 

Site 3 

Fie. 3. Seagrass productivity, leaf blades, g irr d (dry weight) at sites 1, 2 and 3. 


E 300 






i i i 

I i 

Dec-88 Feb-89 Apr-89 Jun-89 Aug-89 Oct-89 Dec-89 Feb-90 Apr-90 Jun-90 Aug-90 

Site 1 

Site 2 

Site 3 

Fig. 4. Seasonal variation in mean standing biomass, leaf blades, g/jrr (dry weight) at sites I, 2 and 1 



biomass wa?i recorded at sue 2 in November 1985, 
352 <:nr. The highesi summer mean standing 
biomass recorded at site 1 nearest the outfall was 231 
g/tn*, m November [98?, while the mean standing 
biomass is consistently lower al sile 3 in Port Palerson 
than at the other two sites with u summer maximum 
of 91 g/m- fFig. 4). m November 1988. Analysis of 
variance Using transformed data llog |M t indicated 
significant differences between the standing biomass 
data ol each of Ihe three transects (P<0.0l). An A 
pi'Mcn'on jnultiple comparison test (Sokal $ Rohlf 
1969) indicated that although the differences between 
sites 3 and 2 and 3 and I were significant, sites 1 and 

2 were not significantly different. 

Comparisons were made of total h|ade lengths and 
blade growl h rales for sites I, 2 and 3 after combining 
Ihe data collected lor these measurements for January 
1983 and January 1989. 

Mean blade lengths were 438.9 + lol .3 mm, 316.0 
i 95.11 mm, and 1355 ± 50.3 mm at sites 2, I and 

3 respectively (significantly different, ANOVA, 

F = 367,6, F,*,., m ~ 3.0).' 

Mean summei growth rates (incremental blade 
lengih.i were 4.7 4 2.8 nim/'d, 3.6 ± K8 mm/d and 
I 9 fc 11 mm/d at sites 2. I and 3 respectively As 
with blade lengths there were significant differences 
between Ihe sites (ANOVA. F - 254.0, 




Host, W. M. (19773 Marine Biological Studies in Relation 
in the Operation tit the Torrens Island Power Station. MSC. 
'Ilicsts. Adelaide University Unpubl. 

Shoot production occurs all year round; the shooi 
index is highest at all sites during summei Shooi 
indices are similar at sites I and 2, while the shoot 
index at site 3 is consistently much higher (about 2 
to 3 limes as higlu than al the other two sites (Fig 5) 

Although no measurements were taken. Held 
observations suggest thai the epiphytic growth was 
consistently higher al site I near the outfall than at the 
gulf-ambient sile 2, and higher again at site 3 in Pt 
Paterson. This was particularly evident lor Ihe serpulid 
worm hilospira awvexis, a species which has 
previously been demonstrated to thrive in the thermal 
outfall al Torrens Island Power Station in ihe Pt. River 
estuary, .South Australia*. 


A number of studies have indicated the susceptibility 
v\ seagrasses in subtropical regions 10 walct 
temperature increases above ambient (Thorhaug el at. 
197S; GFSAMP 1984). tri particular, there is evidence 
that many organisms, including seagrasses, are living 
close 10 the upper limits of their thermal tolerance 
during the warmest pari of the year, and even short 
periods above the summer maxima can have sfgrrificanj 
adverse impacts (/ieman 1974; GHSAMP 1984), 

Larkum ik Den Hartog U989J hypothesise that the 
current distribution of species of the genus Pi>\iilonta 
in temperate regions, and the lack of evidence that anv 
RiSldOniQ species has ever occurred in tropical 
conditions in the past, may indicate thai species of this 
genus have more limited temperature tolerance lhan 
species which occur in the tropics. 

Dec Mar Jun Sep Dec Mar Jun Sep Dec Mar Jun Sep Dec Mar Jun 
86 87 87 87 87 88 88 88 88 89 89 89 89 90 90 

Site 1 

Site 2 

Site 3 

I ig ft Seasonal variation in shoot index | number of new snootsmrU ,t\ U percentage Of Ihe Original number Of blades 
pci in ai the time of lagging) at siles I. 2 and 3. 

Sl-AOKASS I.RilWiH IN SPI 'M<TiR 014.1- £t» I 1 H At STKAIIA 


Id northern Spencer Gulf, although .seasonal wulet 
u-mpcmturcs range from as low as I IX m winter lit 
the iukI u» high 20* during summer (Amshe ft ui 
WJStyi (he high summer water temperatures are more 
typical of subtropical conditions than those normally 
associated with temperate coaslaJ waters ol the wulhem 
Australian coastline. (Suhi topical conditions ute 
defined as those where water leiTipcratuies ecnctallY 
ranee from 20 25°C. and do not exceed M) Q C 

During rind summer, when air lemperaliires range 
to 45°C. the water temperature til lite scaea asses ai (he 
outtall (site I) is consistency about 5 1.5°C higher 
ihan at dull' ambienl sale 2 unset Fij 2). The difficulty 
m dclttune local ambient, water temperature is. 
howevei, ilfusttuted b> the dtffctcme between site 2 
ami she 1 on the southern shores ol (he lame shallow 
cmhayiueni ot Port Paterson. The latter she is also 
he\i>nd the inllne-nce ol ihe (hernial plume, but exhibits 
consistently high avcunje summer tempera Mi res limvt 
He ?,)■ up lo 2°C warmer ihan dull ambient (nisei 
He, 2) 

Comparison ol suiuinet tcnipeialuie regimes 
between siles is complicated by (he shop term 
occurrence cif extreme temperatures, mufti lemi 
maximum temperatures at site I i28°C on two 
occasions. Fig. 2) arc about l°C higher than those 
ice uded at the Gulf ambient she. and as hieh as the 
maximum field temperatures recorded ptioi u> llu> 
siudy for Australian PosUUmia species (Walker & 
McComb I98S». At site 3 which has (lie nmsi 
pronounced diurnal variations the short lerm uia\u:ut 
iccontcd during summer exceeded 30 a C on a number 
of occasions, more than 3°C higher Ihan the peak 
temperature at the Gull ambienl site 2 (Fig 2A» Hrtd 
(he highest field temperatures at which t'osidonta 
species have been rceordeti. 

Although seasonal data have been unavailable lor 
V<isid<m'm sinu>iui, mUumaiiou pti the ptoduclivity u| 
a number ol different species ol scaerasses has been 
reviewed ty Walker & McComb 09KS) and Millman 
el al. t IVK^h These aulhois lound that seauuiss species 
in lempeiate waleis in general lend to show a 
pronounced seasonal variation with adisluici LMovwim 
si-ason in summer However Walker & McComb 'NK8) 
lound no clear seasonal pattern in the producib Uy of 
f'outhtnm uustrults in the semi-enclosed buyy of Shark 
Bay, Western Australia over a one year period. The 
water tcmpeiatuivs in Shark Bay ranged from about 
I8' J C to more ihan 26°C. and maximum temperatures 
up to about 2S°C have been recorded in Lhu urea 
{Walker & McComb !<>«*), Despite the fact that .a all 
i hive sites in the present study summer water 
temperatures resembled Ihe semi tropical condition-* 
i)| 'Shark Bay, ihe E xinut> , <ti entwine al Gulf ambienl 
(sire 2) ami in Ihe marginally warmer conditions 
adjacent tp the power station thermal discharge (site 

ll. show seasonal Oucluation m both productivity and 
slat id me biomass. with niauma lor both measutc. n 
the wanner summer mouths, hvcji al site 3 with the 
most extreme conditions, .summer 'peaks a a* evideiH 
in ihese features, although seasonal patterns are not 
;r> pnMjouiiecd as at the other twe siu-s. Hits persistence 
of summer peaks in productivity and vumdine biom.txs 
l»attieuJai]y at .sites I and ? su^htsIs thai the /' situtiKyt 
ol northern Speueei Gulf may be more tolerant ol 
extreme summer conditions than, loi example- the t\ 
tiitslrulis ot Shark Bay. 

At all sues thcie was sonic ^nation between 1hv 
values rvcooJed from veai to ycai toi suumiet peaks 
ol both productivity and standing binmass This 
vanaiiou was, however, no nn»a' pronounced Ihan 
natural year lo year variations recotded in fl number 
ol recent siudies ol a rauw.e o\ iempctaie Ausliabau 
scay i asses (Wul kei & McCofiib 10SS; llillman A tit, 
l L JS^). AHrUffigh Ihe erowth ol Iheseac'rass/'* \inu<K\tt 
ddlers betueen ihe three sites, ihe inter annual 
i comparisons indicate thai- at any eiveit site, ihe jjiowth 
is comparable Irum \ear to year. There is no evioence 
ol "deteriorrtlian* ol site I adjacent to the power stahou. 
or at either of the two cnntml sues, durine the ctfUr«t 
of the current study. 

Although nuriieo'u- stiuties hflVC documented (he 
range of effects (hat arliliciallv imposed leiupcraJuie 
regimes may have on seagrasses. lew have attempted 
lo describe Ihe progiessrve ellecis on gKMVtil 
characteristics which occur with incremental 
temperature increases atunc ambienl. The cueption 
is that of'Thorhauy ct til ffSTW) who rect>rdfd delaik\l 
changes in ^lowth of leal blades, productivity and 
siandinj', hiomass islanding crop) in nopicaJ arkl 
subtM>pt».ul Unii'twin mju'mIs ftlWcclcd to inerenient.d 
iiHTcases in water lem|terature above ambicul, 

Accepting thai dillereiti species may have tpnie 
different (emnerature tolerances, h is neverlhehss ol 
interest lo examine die snmlanly between Th$t$$$i<\ 
species wilh a subtropical distribution, and /' sirutosn 
subjected to above -arnbienl te i ripe ratu res in an 
environment where summci teniperamro are jlrvatlv 
high in terms of the uet^graphic disiribuiion oi (his 
tcui|iefatc species. 

Thorhau;) ri al (I07S) report that, with increases 
as hide as \.y°C above sunimci ambient watpr 
lempcralurc, v.rowih pel blade, pioduclivily. an.: 
sUnduie, biontass ol THoktSilci declined io 04'v , (>(P* 
and 82 r M tcspectivcly. of that at ambient tcmperuttire^ 
and conbriued to decline sharply with incremental 
inuvase in above ambienl temperature Al ^C above 
atubient ihe sea^riuss 'disappeared" (mm the arca 
(Thorhatif e/ al I47KJ. 

ihe inaMiniim tucaii pPXhlC^lD fWfrfcS '"' f 
sintiOHt at site I. with aveia^c suuniKf svjkt 
temperatures 0.^-1. 3°C above Chill ambient and short 
term peak temperatures up lo l°C higher Ihan short 



lerm Gull 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 
I, mean growth per blade being only about 77% of 
that of blades at site 2 during the warmest time of the 
year. Despite the tact that over the period December 
1988 to August 1990 standing biomass of P, sinuosa 
at site 1 was nol statistically significantly lower than 
at site 2, the maximum mean standing biomass 
recorded during this period at site 1 was only about 
10% 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 
an^ustifolia stand at a depth of 11-12 rn, 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 1 and 2 ranged from about 125 
g/nr to 175 g/nr, while the winter standing biomass 
at site 3 in Pt Paterson was about 50 g/nr (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 1 and site 

2 are at 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. angnstifolia 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 I, 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 1 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/? sinuosa is 
not well documented, P. australis survives in a wide 
range of estuarine salinities up to 57 (Cambridge & 
Kuo 1979). Tyerman et ai (1984) have shown that the 
salinity tolerance 6TB 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 o\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 1'halassia 
Thorhaug et ai (1978) suggest tentatively that increases 
above ambient water temperatures of about 1.5°C may 
be considered "rational" with respect to the limited 
impact on seagrass and associated communities. In the 
current study, the maximum summer temperatures at 
site 1. near the thermal discharge, fall within this 



category. The results df this study also support the 
contention that the overall change in P. vinuosa growth 
is relatively minor with this magnitude ol tt'inpeiuture 
increase above ambient, less than the order of change 
considered acceptable by Thorhaug et ah (197Hi for 
Ttuilassia. In Pt Puterson, inconsistently higher above 
iinihicnl summer water temperatures than recorded 
near ihe thermal outfall, P. siituosa. although exhibiting 
stunted growth characteristics of this species in 
marginal environments (Cambridge 1974; Cambridge 
At Kuo N79), persists with no indication of ongoing 
decline, in terms ol'biornass, productivity, leal blade 
growth and length, or areal extent ol' local distribution, 

II has been proposed that the sediment stabilising 
role o! seagrasses may be particularly important in 
maintaining the integrity of the sloping banks of the 
channel of northern Spencer Gulf'. While the stunted 
seagrasses on the relatively protected shores of Pt 
Paterson survive wiihout any evidence ol ongoing 

decline, il the seagrasses in the more exposed Gulf 
channel (lor example adjacent to the thermal outfall) 
were to be subjected to further, small summer watci 
temperature increases, resullant stunted growth could 
lead to increased localised mobilisation of the shoreline 
sediments. This, in turn, could lead lo longer term 
effects on the spatial distribution of seagrasses in this 
urea. Given this possibility, any consideration 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. 


This work was funded and supported by The 
Electricity Trust of South Australia as part of fa 
ongoing programme of environmental research, We are 
graieful lo Dr. Alan Butler lor reviewing tne- 


Ajnv u - ( R, C tl9R0> Polenta! Gieenhouse Impacts on Ihe 
mologv ol Ihe Shallow Waters Df the South Australian 
f lulls, and otl the Grey Mangrove Ario'H/tnt tTuirina pp 
03 V0 tn Dendy. T iftd.l "Greenhouse SX Planning for 
Climate Change" (Department of Hnviroiunenl and 
Planning Adelaide). 

ItJHNSTON'i E A-. & Oritik. B W i&$9) 

Intertidal communities ol northern Spencer Gulf. South 
Australia, trans. H. &r, S Avst- 10, 69-*a 

CftMnkiix.K. M L, (1974) Seagrasses of souilv western 
Australia with special reference to ihe ecology ol fttbidon'to 
ouMiafis Hook F. in a pollute environment Aqual. B*rt. I. 

& Kcft, ). (1979) Two new species ol ft&£ra$ift£ 

from Australia Pvstclutun sinuosa and P atigu>fifnh>t 
iPostdoruaeeact. thid 6. 3U?-32ji 

Cit,s\MiMl984» lMr>/K\CvLlN'LSCo'WMi-».'V.'Hn/|A( \f\jNjVtfl& 
Joint Group ot E xperts on the Scientific Aspects o\' Marine 
foliation, Thermal Discharges in the Marine environment. 
&p, Stud Gl.SAMP, 24, 

HiMWK'k. K., A. & Black, N, D_ {l L )Kh) **G rowing Medu 
lor Oniarnenial PUms and Turf." (New South W:*Icn 
Umversiiv (jreMt. Sviiney). 

Mit, K.. Waikkr. D: I . I.amki.'M. A. W. D.. & 
Mi/comb, A. .1 (I9h'9) Productivity and Nutrient 
I iiiiilalioTL pp 635-668 In LarUwM, A. W. D., MeComb. 
A J and Shepherd, S A, (Eds) "Biology ot Seagrasses" 
ibUevicr, Amsterdam) 

Lakklm. A W. I> (1976) toojojtry of Bolany Bay, 1. Growth 
of h^:Jf>/iui ttuxrrutis < Brown i Hook, f. in Rnianv Bj\ and 
other havs of the Sydney Basin. Anaf. X Mar. k'r?$h» Kt's 
27. 117 i27, 

& den H.\fcToo. C. M989) hvoluiion and 

Riogeography oJ Seaprasscs pp. L12-445 in Laikum, 

A W DL, McCoinb, A. J- & Shepherd. S. A (fids') "Bioloj^ 
ol Seaytasso'" (ELsevier, Amsterdam), 

NtvhKMSMS. V P U9KKj Responses of a I'ostdomu 
community to prolonged reduction tn h^hf. Aquoi Bin 31. 

Ni Nis. R. A & Llanon, C_i. W liVSb) PhV>ic*d1 properlv 
distributions and seasonal trends in Spencer Gull, South 
Australia: and inverse &(wvy. Aust J, Mot: thwfn-i R*\ 
37(1). 39 53, 

Ptii.i ARU. P. C. & CiKhrNHAV, M. HW) Photos^mhctie 
characlerisiies ol seagrasse.f {Cynunlorta xfrruhua. 
flttilaSstil Hcmjmchn. and Zttficm capnvami) m a low liylu 
environment, with a comparison of leal tnatkinu ami 
I-jcunal gas measurements of productivity. Ibid. 44dt_ 

Rasmu:>SiiN, B- {Wf'h The uA&tfftg disease ol eelgrass 
(£&Mffli imrma] and ils elTtels on environmental lactois 
and fauna np- 1-44 //( McRov, C. P (ft Helfferich, C. {Hd.M 
"5pagr4SS Eeosvslems. A Scientific Perspective" (Marcel 
DekKer. NfiJW Yolk;. 

Robinson. K. I. M (I9^7| mi'ects orthermal power station 
eltlucnt on the seagrass henthic eonmiunitics oJ lake 
Maequane, a New South Wales Coastal Lagoon. W'thimls 
7(1), t 12 

Snii'HLRD l S. A. (WS3J Benthie eomrnuriioes ot upper 
Spencer OuM, South Australia. Tnms. ft. Soi' S. Aust. 
107(2). 6^-85, 

iNHCO Coast.d waters pp. _U6 3*fi hi Nance, C &. 

Speight. D. L. (Kdsl "EmtronmeniHl Chanues m S.A* 
(Longman and Cheshire, Melbourne) 

, McComh, A.J , Bii.rini5, D- A Nkvi,w\( skas. 

V., SintrssoN, D. A. & Wtsi. R. J. <l^8*>> Decline of 
Se.tgrasser- pp. 211-225 in Larkum, A. W. U, McComh. 
A. J & Shepherd, S. A. l£ds) "Biolopv ot Scagrtsst-- " 
(Rlseviei. Arnnicrdani). 



Sokal, R. R. & Rohlf, F. J. (1969) "Biometry" (W. H. 
Freeman & Co., San Francisco). 

Thorhaug, A., Blake, N. & Schroeder, P. B. (1978) The 
effect of heated effluents from power plants on seagrass 
(Tlialassia) communities, quantitatively comparing 
estuaries in the subtropics to the tropics. Mar. Poll. Bull 
9. 181-187. 

Tyerman, S. D., Hatcher, A. I., West, R. J., & Larkum, 
A. W. D. (1984) Posidonia ausiralis growing in altered 
salinities: leaf growth, regulation of turgor and the 
development of osmotic gradients. Aust. J. Plant Physiol. 
11, 35-47. 

Vicente, V. R (1979) Impact of heated effluents from a 

power plant on seagrass beds. Proc. Assoc, lsl. Mar. Lab. 

Carrib. 1979, 33. 
Walker, D. I. & Mccomb, A. J. (1988) Seasonal variation 

in the production, biomass and nutrient status of 

Amphibolus antarctica (Labill.) Sonder ex Aschers., and 

Posidonia australis Hook. F. in Shark Bay, Western 

Australia. Aquat. Bot. 31, 259-275. 
West. R. J. & Larkum, A. W. D. (1979) Leaf productivity 

of the seagrass Posidonia australis, in eastern Australian 

waters. Ibid. 7, 57-65. 
Zieman, J. C. (1974) Seasonal variation of turtle grass 

Tlialassia testudinum Konig, with reference to temperature 

and salinity effects. Ibid. 1, 107-123. 




VOL. 118, PART 4 






ByRobynL. Molsher*, Michael C. Geddes* & David C. Paton* 


Molsher, R. L., Geddes, M. C. & Paton, D. C. (1994) Population and reproductive 
ecology of the small-mouthed hardyhead Atherinosoma microstoma (Gunther) 
(Pisces: Atherinidae) along a salinity gradient in the Coorong, South Australia. Trans. 
R. Soc. S. Aust. 1 18(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 36^15 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 m November. The numbers of large hardyheads (>35 
mm) in samples declined in December and January, reflecting post-breeding 

Key Words: Atherinosoma microstoma, Coorong, hardyhead, salinity, reproduction, 
population biology, fecundity. 

lhwttntlumx of ihr fioytl Sorim of S AuK (1994). 118l4l. 20/-2I& 




by Robvn L. Moism-R*. Michael C. Glddls* & David C, Paton* 

Sum mar v 

Mwisher, R t L , Gr.nnr.5, M. C. & Pa ton. D C \\W4\ Population and reproductive ecology of ihc small 
mouthed hardyhead AthvnmiMmiu murosttunu (TJunther) i Pisces: Alhcnnidaei along a salinity gradient in irte 
Coorong, Scrtlth Australia. Trans, ff. Sor S V,-m. M8(4)< ?07 2I6 1 30 November iiW4 

Athfrw\rni(t r/uavMOtHtl ill iht Coorong exhibited a one -year hie cycle will. multiple spawning oVet a four 
rnonlh breeding season from September to December Large- numbers of larval and juvenile lo.h (5-t5 Wlffl long) 
appeared in samples during October and November and grew rapidly over summer, mint reaching length,'- oi 
2o- *5 mm by attlumn and 36-45 mm by the following spring. Only one ovary developed in females and this began 
10 enlarge dining August when batches of eggs began maturing. GonosornaUV indices also began to increase ai 
this lime Ivmale fish with spent ovaries were first caught tt» November. The numbers ol large haidybvad* ( >35 nmij 
in samples declined in December and January, reflecting post-breeding mortality. 

Salinities ranged from 9 lo 67 g L ' at Noonameena, pie most norihcil> sampling Sitt and Prflti) 35 to w-t g 
L ' nt Tea Tret Crossing in ilk- southern end of the G>orong lagoon system. High salinities did not have a marked 
elfect on the population ecology or reproductive potential ol \ Athtritwst'tna mtirosionui. Hardyfteads vvetc caught 
ni all sites on all sampling occasions atid no marked differences were found in llsrt si/e, gtov-th lute, condition 
\$\ rHalive hatch fecundity lor lish caught from different localities along the salinity gradient Hardyheads were 
coping well with the high and fluctuating salinities that exist in the Coorong. Significant differences in batch 
leeimdiiy were found between years, with those- m spring WO being about hall those ol tW. perhaps rcflettjog 
diflcrcnces in food uvailahilily The possible effects of future proposed icductnms m -.alinity lor the Coorong 
on the biology ol \'A. mtrrt'Stoma are discussed. Changes in salinity arc unlikely to limn die distribution ol hardy heads 
BW!Cfl| ihroucb possible influences on ihcir food supply. 


■itht'tiH'.>sonui mnr t K\toma. Coorong. hardybead. salinity, reproduction population biology. 


The Alherinidac I hardy heads) is a widespread family 
nl small fishes that are commonly found in calm, 
shallow waters and often have short life cycles (e.g. 
Gon & Bcn-Tuvia 1983: Prince €t tli Wl, Prince and 
Potter 1983, Potter ei of. W83. IV86). Aihctinids mc 
often coiybalinc but high salinity may affect their 
population and reproductive biology. Hor example, 
suspected dwarfing in Jjie Mediterranean atherinid 
Arht'rina hoxeri, may be associated with high salinities 
(Gon & Rerr-Tuvia [983). The North American 
alhetinid. AtheuHops uffini.s ujjinis. spawned in the 
field at salinities of 72 p.p.t. but the young died within 
four month:* (Carpelan 1955) Mosl fish species hi the 
Laguua Madrc of Texas fup to SO p.p.t. 1 do nol spawn 
at salinities greater than 4j» p.p.t (Hcdgpcth W67j 
cietictally, high salinities and fluctuations in .sajtmty 
restrict reproduction in many -icjuabc animals, affecting 
both the number of offspring produced and the liniing 
and length of the breeding season (Kinnc t%4i. 

* Department of 7.oology, Uiiivtn'MlV of Adelaide. Solid) 

Aus|r;d..i SWS- 
1 IvAMSOl-) , V (t v '?X) Taxononiie and systematic review ot 

the Australiati fish species of the family Athcrinidat v>)\h 

references to icluted species ot the oUt world l'npob!» W 

PhD Thesis, Matquane L'niAeisitv. 

The small -mouthed hardyhead. Atlurinotomct 
microstoma (Giinthct). is found in abundance over a 
wide range ot saluiitics including estuaries, marine 
ciiibayiiiems and hypetmarine lagi>ons in south-eastcni 
Australia from the Tuggerah Lakes in New South 
Wales, southwards and westwards it) Spencer Gulf in 
South Australia Uvantsoff N78 1 ; McPowall I9HU; R. 
Connolly &.G. K. Jones pets, comm ) The maximum 
sizes approach 00 firm (total length) in Tasmania and 
80 mm in Victoria (Cadwalladei & Backhouse NS^i 
The life cycle of A. mtcr<h\ti>mt! has been sludied in 
Dec Why Lagoon NSW., where saliniO'es ranged 
ftpm 3 'o [3 p.p.t. (Potter cl at. Wotj)- In these 
estuarinc conditions, t mi> nrstomu exhibited a one 
year life cycle with u four. month breeding season 
during spring (August lo NoVt-mbet ). Prv entcied ih. 
trappable pt)pidaiion ill October, while larger adulb 
tapidly declined in abundance after November. Growth 
effectively ceased over autumn and winter (April io 
August ) 

A. mitrosioma has been found thonigbotit the 
Cp< "ong where cttndiftons range from esluarine in ihc 
north In hypcrsaltne in the south (Geddes <Sc flullei 
J0S4). Although hnrdyheads have been caughl in ihc 
Coorong in excess of 11)0 p.p.t. IDS ttoud disstdved 
,olids). ei|uivulcni to a salinity Uf QJ (rp.t.. Geddes 
< 1987) has suggested that cxtreniely high sat hi I lie* 



<>100 p.p.i. IDS) during late dimmer cind autumn 
ina> restrict the southerly distribution ol these fish in 
(he Coorong m some years, This study takes advantage 
ot the longitudinal gradient of salinity that exists in 
the Coorong to study the effect oi high sahnil) on ihe 
reproductive performance and population structure ot 
A. miirostoma. 

Materials and Methods 

the population structure and reproductive biology 
of hardvheads were studied at five sites within the 
(ooiong: Noonamecna. Villa dei Yumpa, Polieemans 
Priint, Salt Creek and Tea Tree Crossing (Fig. I). These 
sites spanned a 65 km length of die Coorong lagoon 
system, with Noonameena in the north usually 
experiencing a lower range of salinities than the other 
four sues that were farther south (Gcrides 1987) The 
southernmost site. Tea Tree Crossing, was at the 

5. Tea Tree Crossing 
^mi Coorong Lagoon 

Fig. I The C*»oiong U*goun-.lkiwimi the five MnptfTTH Mies, 

southern extremity of permanent water in the Coorong 
where die highest salinities in the Coorong had been 

The salinities and temperatures of the water at each 
ot the sues were measured al monthly intervals between 
September two and January 1992. Water temperatures 
were recorded at a depth of 40 em between 0900 ti 
and 1800 h, Watci samples, collected Irom a depth ol 
40 cm, were returned to the laboratory and salinities 
were estimated by measuring elect Heal conductivities 
(conduemity meter C'I)M3i and converting these 
values to salinities usinu tables from Williams (1986), 
Samples W'tth conductivities o\cr 100 mS cm ' were 
diluted and the calculated salinities multiplied by The 
dilution factor. 

Hath month, fish were caught from each site usm^ 
a 5,5 m long, IJ5 m deep seine net with a mesh si/.e 
of 1.9 mnt. All seining was conducted at a depth of 
0.2-0.8 in with hauls over a distance ot 40 m. On mosi 
occasions a single haul was taken but when the sample 
contained low numbers of fish ( <50h additional seines 
were undertaken. Successive semes in the one area 
showed no sigmficanl difference in the mean length 
of fish caught (t - 1,348. N - 200. P - 0.179). Fish 
traps (63 by 36 by 36 em, mesh size 1.9 mm) baited 
with meat (usually chicken) and set overnight in water 
0.4-0.8 m deep, were used in die first six months of 
the study. Traps were eNpected to caieh larger fish, 
whose superior swimming ability may have enabled 
them to escape the seine net_ and thus provided 
information on the larger adults in the population They 
also provided additional fish tor assessing reprmluclive 
condition. In addition, a plankton net (330 ;<m mesh. 
60 cm diameter) was hauled through ihe water i, depth 
0.2-0.6 m) for two minutes lo check lor The present, c 
of larval A tmtrcsUnna, Fish were immediately 
preserved m 10'v buffered formalin and returned to 
(he laboratory where their length and reproductive 
condition were measured. 

'The lota! length (TL-tip oS snout to ericl ol caudal 
fin) of each fish Irom both ihe seme and trap samples 
was recorded to the nearest millimetre using dial 
calipers. Where the number ol individuals was large. 
a random subsample of approximately 150 individuals 
was measured The siandard length iSL-tip of snout 
to postetior ed^: of ihe last lateral line scale) of a 
subsample of fish was measured to determine the 
relationship between TU and Si,, thereby allowing 
comparisons with other studies. Lengths and body 
weights nl" hardvheads caught in October 1990. 
Deeembet 1990 and March 1991 at each of three sites 
i Aoonameena. Foliccmaris ftiint and Tea Tree 
Crossinei were measured .so that length weighr 
regressions could he calculated. Fish condition was 
assessed by comparing these ienglh- weigh! regressions 
between si lev All length and weight measurements 
were obtained Irom fish that had been preserved in KT# 



buffered formalin. Sex ration were calculated for up 
fep 50 haalyheads {>M) mm length) caught in each 
seine sample from October 1990 to May 199). The sex 
of the fish wa.x determined by macroscopic examination 
of the gonads. Gonads in fish <3fl mm had not 
ddierenlialed 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 thai exhibited this coloration was determined. 

Seasonal patterns in the reproductive cycle of 
hardy beads were determined from changes in 
gonosomatic indices (GSI) and the examination of 
oocytes in the ovaries of female fish. Male fish were 
not investigated as preliminary examinations showed 
no marked changes in gonadal weight The 
gonosomatic index (GS1) was calculated by expressing 
gottad weight as a percentage o! body weight (l)e 
Vlaming ft ai 1982). Gonad weight and body weight 
were measured to Ihe nearesl milligram for up lo 20 
female fish <TL>40 mm) in each sample. Only female 
fish that were at least 40 mm in length were used in 
the analysis as the ovaries of smaller fish did not 
contain maturing oocytes. Ovaries o\ female fish from 
three sites (Noonamecna. Policemans Point and Tea 
Tree Crossing) were excised and presetved in modified 
Colon's fluid (Puckridge 1988- > These ovanes were 
then teased apart, shaken vigorously, and stored for 
two weeks to separate the oocytes from the ovarian 
tissue. Preliminary microscopic examination revealed 
ihree different egg types, which parallels the situation 
found tor M'tiidxt menuiiti (fonovcr 1985). The ihree 
general egg types were classified by size (oocytr 
diameter) and appearance as follows: 
I. immature oocytes: 0.05-0.70 mm in diameter The 

smaller oocytes in this group (0.05-0.22 mm) had 

a clear cytoplasm and large nucleus, while the larger 

ooevtes (0.23-0.70 mm) were white, opaque and 

ollen irregular in shape. 
2 maturing oocytes: 0.71- 1.60 mm in diameter 

Spherical and dark yellow in colour. 
3. ripe oocytes; 1.61-2.50 mm in diameter Spherical 
and hydratcd with a yellowish yolk centre. 

Diameters of oocytes were determined from a single 
measurement on a random orientation basis iWcsi 
1990) using a microscope-video attachment and a 
digittser pad downloaded to a computer, Subsequent 
categorizations of oocytes were based only on 
appeaiance. Numbers of maturing and ripe oocytes in 
ovaries from samples over the two breeding seasons 
were counted using a stereo-dissecting microscope; 
immature oocytes were difficult to count and were only 
counted for samples of fish caught between September 

J PlifKKloGK J. T. (1988) The hfe history ol a gizzaru shad, 
the bony bream, Nennthilosn evibi (Guntbcr) 
tDomsomatinae. Teleosu) in the Lower River Murray, 
South Australia; MSe Thesis University ot Adol.ialr. 
I inpubl. 

au - 

Immature fish 

im mala re 

c/j 40 - oocytes 

Q) r- 


% n 


8 30- 




ai 20- r 





2 10 - 

n . . 

1 rin 

U T\ 

111 | 1 1 1 1 1 1 1 1 1 | 1 1 1 T | 1 1 1 1 

0.5 1.0 1.5 2.0 2.5 


cr\ _ ooevtes 

DU , — 

■ 1 

Mature fish 

w 40- 


4— ' 




S 30- 

**— r 



il 20- 







2 10 - 

j ! maturing 


UN Hnnnll 

_D T 

U | i ; i i | < f i i | i i i i | i i i i | i i i i | 

0.5 1.0 1.5 2.0 2.5 

c;n — 


Ripe fish 

40 - 



fi ii 


8 30 " r- 






£ 20 - 

;| 1 maturing 


i oocytes 


n ' r 


| 10 -n 

r 1 

n — , L 

(1 nflflU 

rutin nRnrinnnr 


tJ 1 1 

T 1 |"|"l 1 T'l | ! 1 1 1 | 1 T' 1 1 | 1 I 1 1 

0.5 1.0 

1.5 2.0 2 


Oocyte diameter (mm) 

pig. 2. Frequency of oocyte diameters from (he yirtlads of an 
immature fish, a mature fish and a ripe 1Kb. 



I'M) and January 1991. Fish were classified into (bur 
sWgcs depending on the type of oocytes present (Fig, 2); 
I immature fish: immature oocytes only, firm ovary 
I mature fish: immature and maturing oocyte 
3. ripe fish; immature, maturing and ripe oocytes. 
4 spent fish: maturing or ripe oocytes absent, flutckj 


B;uch fecundity" was detmed as die numbet ol 
nuuuiing .k.lvics in the ovary ami presumably 
represented | he maximum number ot oocytes that could 
he ripened and subsequently spawned al one lirne. 
"Relative batch fecundity" was defined as the numbci 

tfffrraiufiflg otHjytes pei grttfliJofcWT-fittfeocty wcjgbi 

(Conover 1985) Differences in the fecundity of temalc 
lisli at dillerenl sites in the Coorong were assessed by 
comparing icbrivj batch leeundities. 


PhyXital Charm tenstic \ of the VoarQrtg - September 
l$Q to I'twntn IW2 

A longitudinal gradient m salinity persisted m the 
tortrmg with salinities increasing from Noonarneena 
in die North Lagoon to Tea "free Crossing at the 
southernmost end ol the Cooroug (Fig. 3.» Hypersalme 
conditions ( >3S g I, ') were maintained at all sites in 
Hie Smith Lagoon, except briefly at Villa del Yumpa 
in September I99|_ At Noonarneena, conditions were 
esuianne during winter and spring (9 32 g L J and 
generally hypermarine during summer. Seasonal trends 
were also evident at .ill silos with salinities high in 
summer lading m autumn and rising again in spring, 
In Seprcmhei logo, salinities ranged from <2 gL l at 



a 40 

:Tea tree Crossing 

•salt Cf«sk 

5 Poli^emans Point 

* Villa dei Yumpa „ 

• Nnnnamaana 

| p i t 1 i — T 





! i J 

S O N tl 


J F M A M 

T — T 

-J J A S O M II 

19.90 f991 1992 

Seasonal fluctuuinws in sabmfy and lornperaiun? Bl 
Sires in the CnOTTing Lagoon. SalliUWBS irre slftwwl ft*! 

i site anil l*-mpcritiirt>.irt; \huwnax ihe tueun jik! raBgC 

iIil live <iiC5 ^v-nihinal. 

Noonarneena to 57 g L ' *tl Tea tn*g Crossing, wbifc 
in February 1991 salinities had risen to 60 g L ' and 
94 g L" 1 nr these two sites respectively. Seasonal 
changes in salinity were similar in the two year-. 
However. salinities reached their minima earlier in (991 
( August/Septembei ) and were tower than in Ihe 
previous year Water temperature varied seasonal!', 
vvuh maximum water temperatures in excess of 24°(* 
recorded dUriUg lWC summer ami minimum 
temperatures Of II°C in winter (Fig. 3). 

Distribution of tish tn the Coontnx 

Hardyheads were collected from all sites throughout 
the 17 month period. The highest salinity at which they 
were found was 94 g L' (Tea Tree Crossing. 
February 1001). Five other species of tish weie enughi 
commonly; yclloweye mullet (Aldriehvtkt forsteri) 
congolli iPsrtidophritts urvilli), river garfish 
{Hvporhantphus ive/'/f/m >, greenback flounder 
{Riwmhnsnlcu ut pinna) and blue spot gnbj 
{Psi'udoy.ohius olonmi) Alt species were caught at 
Noonarneena and Villa dei Yumpa where salinities 
were lower. However, yeltmveye mullet was the Ofllj 
species caught with A wit tostomu at Tea Tret* 
Crossing, and then only during winter and spmu , 
Yclloweye mullet, congolli and blue spot goby were 
caught at salinities up to 64, 83 and 87 g L' 
respectively. Flounder and garfish were caught only 
al relatively low Salinities' |<36g L 1 ) and only on 
three occasions. A. mierostomu tar outnumbered othci 
lish species in each sample. 

Population Sinn run- ^/Atherinosoma microstoma in 
the Cooronx 

I he abundance offish collected at the five ales on 
the 17 sampling occasions was analysed by two way 
analysis of variance. The number of hardyheads caught 
in the first seine tor each sample was i\^d in the 
analysis There was no significant difference between 
sites" (F - 0.87. d.f, - 79. p>0.05» but there was a 
highly significant difference between sample dates 
tl"- 6.03, d,f, = 79, p<0.001). Greater numbers ol 
hardyheads were caught during spring and summer 
j'he length frequencies of fish in each sample were 
inspected and no consistent differences between sues 
were noted so the sites were pooled tor length 
ffequciirs analysis. The sizes ot hardyheads Were 
highly vaiiable al any one lime in the year, bul there 
were distinct seasonal patterns (Fig 4). Larvae ynd 
try (5-15 mm; length class I ol Fig 4) were prominent 
in October, November and December (spring-early 
summer) e,i 199(1 and 199L At thi.s time of the year, 
length-frequency distributions were often distinctly 
Wttiodal, t-onsistmg nf suuil fish i c25 mm) arid largef 
lish l>3? mm), The 1990 cohort o\ smaller fish 
showed a gradual increase in length during summer 
reaching lengths of 26-35 mm (length class 3) by M;r. 



1991 and 36-45 mm (length class 4) by the following 
spring (September 1991) at all sites (Fig, 4i. 

The length-weight relationships for fish collected 
from Noonameena, Policeman* Point and Tea Tree 
Crossing are shown in Table I. Statistical comparisons 
of the slopes by ANCOVA revealed non-significant 
differences between the sites in December 1990 and 
March 1991 (Tukey HSD p>0.05). ]n October 1990. 
significant differences were tbund between each of the 
three sites (Tukey HSD, p<0.05), however the 
assumption of homogeneity of slopes was violated in 
(his month (F - 5.66. d.f. = 271, p - 0.004). The 
.slopes of the regressions, which indicate relative 




1 ■« 

- ) ■•; i > s S ■'■ 

v « g ( :: d 

- 5 r, I s .7 


Fig. 4 Relative length-class frequencies of Atlierinosomo 
microstoma caught in .seine ne' hauls from October 1990 
until January 1992 (two samples 14 days apart Wert; taken 
in October 1991). Catches from the five sample sites have 
been pooled as there were no consistent differences be! ween 
sites in length class frequencies. Ihe number ol fish 
measured in each sample is indicated at Ihe top lefl of each 

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 Off male and female fish in 
the trap and seine samples from all sites combined 
taken m October 1990. December 1990. March 1991 
and May 1991 are shown in Fig. S, Females were 
significantly larger than males (independent i-tests. 
P<0.05) in each month. The largest lemale fish caught 
it] the trap samples was 85 mm (November 1990) while 
males were below 67 mm (although a single male lish 
of 75 mm 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 (\~ =37.82, N - 413. P<0.01). Orange 
coloration of the eyes arid mid lateral line was only 




3 o 


a 20 


§ 15 



Ocl90 20 1 





-• 1 1 — i r 

M9r 91 

?n=S1 15 



n Pi 

1 1 — 1 1 I T ' 

*i en ■«* m -*t Ol o 
v -<T IS} in id n> t-- 

6 ip 6 tA 6 ir» *' 
■a -4 io ifi © id 

i r 


:'j in 

oj •? -3) ■<* o) o 

«■ in if> to ic r- 

o ih o »i o io o in 

n n v t in m id iB 

Length (mm) 

□ Female □ Male 

Fig. 5, Relative length class frequencies lor lemale and male 
Athcrinoxoma mivivstonut in Ociobet 1990. December 
1990. March 1991 and May 1991. For each month fish lium 
irap and scinc-ncl samples were combined. 

Tabu-: I. Length Weight regressions t)J 'Athcrinosoma microstoma from Niumamevna (NM). Policemans Point fPP) and Tea 
iter Crossing (HCJ in Octobrr 1990. December 1990 ami March 1991 

Regressions are of the form Y - for where: Y = weight, X — length, a = intercept ami b — slept 

<lfttrminniton. n - sample sire ami * 


'efficient of 

Sample date 


October 1990 

December l«>90 







9.12 x UV 
I 55 X 10 ' 

3.54 x 10' 

4 36 X 10* 

5.75 X 10* 
8.13 X 10° 





0.89 1*** 







March UMl 


T" C 

6.03 x 10° 
■S.32 X 10 ft 
?;Q8 x LO* 6 



2 9H 








found in male fish over 36 mm TL and only during 
ihe breeding season {September-January). indicating 
lhat it is related lo reproductive behaviour. 

The relationship between total length (TL) and 
standard length (SL) wa.s expressed by the following 

(r_0.99, N=34, P< 0.001). 

Jrirtiufl Rvprotluitivp Cyrle of Atherinosoma 
microstoma in the Coorong 

Only one ovary developed in female A, microstoma 
and this remained small {<2% of body weight or 
GSI<2) through autumn and winter in the Coorong 
{Fig. 6). In spring. Itic ovaries showed a marked 
increase in si7c 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 in reproduction was also 
reflected in die proportion ol immature, mature, ripe 

A — ' 

E 4 


I \ * 

PoilCBTnans PtaKil 

^ ■' . 

• • - i , 

Sail Creek 


|Tea Tree crossing 



1 ', 


— I 1 1 r t — i 1 — * — ' — i — 1 T ' ■ t t ; . 





fig 6. -Seasonal changes in the gonnsoniatic index of female 
Atherinosoma microstoma ai live sites in the Coorong 
LflgooTI Points represent mean values fit up lo 20 fish and 
vertical lines art one standard deviation each side oT the 
mean, tVo standard deviation is given when .sample size 
uus less than HI. 

and spent fish in the samples (Fig. 7). Female fish 
possessed only immature oocytes from May lo July. 
Maturing oocytes were first detected in August with 
large numbers of mature and ripe fish present 
throughout spring. Spent fish were first detected in 
November. Some immature eggs remained in the ovary 
indicating that not all oocytes were matured -and shed 

Batch ft'CMuiiiy of Alhennosoma microstoma in tht 

The numbers of oocytes, classified as immature, 
maturing or ripe rtl the ovary of individual fish collected 
from the two breeding seasons are shown in Fig. 8. 
The large variation in numhers of immature oocytes 
may relate lo fish si/c and lo losses due to maturation. 
There were smaller numbers of maturing and ripe 
oocytes with most fish having from 5 to 40 maturing 
oocytes and 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 1991 at the time of peak 
gonad weight {Fig 9a). Relative batch fecundity 
(numbers of maturing eggs per grain of ovary free body 
weight) lor fish wa.s independent of ovary free body 
weight (r = 0.04, n - 101) (Fig. 9b), indicating lhat 





















! 1 I 

§ C £L 











* s s 





JH Spent □ Mature 
|H Ripe ■ Immature 

Fig. 7. Relative proportion of female Atherinosonnt 
mimntomtt characterised by the most advanced clutch 
present in the ovary. Data from all five sites combined, 



fish of different sizes were allocating a similar 
proportion of (heir resourees to egg maturation. 
Relative batch fecundities (Fig. 10) were not 
significantly different between sites in spring 1990 
(F - 0.16. d.f. = 2,45, P - 0.851) or 1991 (p = 3.23. 
dX - 2,98, P>0.05). However, significant differences 
were found between years, with relative hatch 
fecundities in 1990 being about half those o( 1991 
(F = 5I.A7, d.r. = 1,147, P = 0.001). The number o\' 
maturing eggs per ovary tended to decline after the peak 
spawning period in October J09L 


Influence oj Salinity on Distribution, Growth, Size and 
Condition of " Aihennosoma microstoma 

Over the summers of 1990-1991 and 1991-1992 the 
distribution of Atherinosoma microstoma in the 
Coorong was not restricted by salinity. All other fish 
species appeared to be limited by salinity during the 
high-salinity summer season in a manner similar lo 
that reported in Gcddes (1987) and Geddcs & Hall 
(1990). The highest salinity at which A. microstoma 
was collected was 94 g L '. This record exceeds the 
maximum Held salinity for this species from coastal 
salt ponds in Victoria (82 p.p.t.. Chessman & Williams 
1974). In March 19X5, a few individuals of A, 
microstoma were trapped at 149 p.p.t. TDS (salinity 
approximately 130 g L ') al Tea Tree Crossing in the 
Coorong (Gcddes 1987; D. C. Paton unpublished). A. 
microstoma ranks along with Cxprttwdan varie^atus. 

which has been recorded al 142 p.p.t. TDS (Simpson 
& Gunter 1956), as one of the most salt-tolerant fish 
species in the world. This ability allows A. mtcrostoma 
to survive at the southern end of the Coorong during 
extreme hypersaline conditions. Abundance data 
suggested that hardy head numbers at the southern sites 


1 2 3 

Ovary free weight (g 

o J 

10 i 

d - 

fipe oocytes 

I; li \\ ii i i 

; -i - — r 



| maturing oocytes 



20 - 



o -I 

1 1 111 I Hi H.i fl : 


1 1 1 1 I I I I I 

i i i i I i i i i i i 


inillll 1 1 ill 

immalure oocytes t .I -i i inn: "] "i 

I > i i ' ! " 
fOO 300 

T 1 1 


Number of oocytes 

Fig. X. Frequency distributions of the numbers of immature. 
maturing and ripe oocytes in the ovaries of female 
Athcrirut.sttmu microstoma caught in the Coorong during 
the 1990 and 1991 breeding seasons. 

1 1 1 1 1 r 

12 3 4 

Ovary free weight (g) 

Fig. 9. Relationships between (a) batch fecundity and ovary 
free body weight (r = 0.68. n = 101 ) and {h) relative 
batch fecundity and ovary tree body weight (r = 0.<U. 
n = 101) for female Aiherinusoma microstoma. Points 
represent fish collected in September. October and 
November 1991. 



wric not ulfcetcd bill high variability, perhaps 
associated with schooling behaviour of trie fish, and 
|0H numbers ol samples were confounding factors in 
llns analysis. 

No major diflcrcttces were found between sites in 
growth, size or condition ol hardy heads in the 
i "<>omng. Growlh rales implied from the progression 
<»t length modes suggest ih&iA, mnmstoma can achieve 
a total length of 35 mm in lour months, followed by 
a period of little growth over winter, reaching lengths 
of 45 mm the following spring, and thereby becoming 
sexually mature within the first year of lite- There was 
\h) indication of dwarfism related to high salinity in 
this population, Maximum sizes lor both sexes Were 
similar at all sites in the Coorong and comparable to 
those lound in the Dee Why Lagoon population (Potter 
i'i al, l u S6) and lor this species in general (Cadwulluder 
Si. Backhouse 10X3). Maximum total lengths recorded 
lor males were 67 mm (present study) and 66,5 mm 
(Dee Why Lagoon) and for females 85 mm (present 
study! and 8M mm (Dee Why Lagoon), Fish 
condition in the Coorong did not differ signifitantlv 
between sites in March when .salinities were high and 
ranged from 58 at Noonameena to 92 g L ' at Tea 
Tree Crossing, 

Ufi fii\ii>r\\ Annual Cvrfe dtnl Pitpulattoii Structure 
of Atherinosoma microstoma 

The post-breeding decline in larger fish at all litest 
from December indicates a one-year I lie cycle. The 
population showed a numerical dominance of females, 
and mules were also significantly smaller than females 
Similar patterns of life history and sexual dimorphism 

have been found in the population at Dee Why Lagoon 
New South Wales (Potter c/ .//. J9S6) and in atherinids 
from the Northern Hemisphere (e.g. Gon & BcnTuvia 
1983), The proportion o\ female fish in this population 
appears to decline over the breeding season and a sex 
ratio of close to one is reached in December. This mav 
be due to differential mmtalitv during the breeding 
season, or to an increase in the numbers ol male fish 
reaching maturity ( >M) 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 m si/.e between sexes, ihr 
prolonged breeding season and the stunt life cycle of 
A. microstoma may be indicative of a species where 
sex is determined after conception by environmental 
factors, such as temperature (Conover 1984). Conovct 
& Kynard (19X1) conclusively showed that temperature 
exerted a direct influence on primary sex 
differential fan, rather than causing sex specific 
moriality in the athetinid, Mcmcliu memd'ni Thus, if 
eggs spawned in the cooler spring waters develop into 
females, and eggs spawned in the warmer waters of 
late summer develop into males, then females would 
have a longer growing season which would account 
for their significantly larger size. 

Spawning occurred from September to December 
giving a protracted four month spawning season. This 
ia supported by the large numbers of larvae collected 
from October to December from all sites. The absence 
v\i larvae in the samples, the small GSIs, and the 
absence of maturing and ripe oocytes in the ovaries 
during autumn and winter BKfl, indicates that this 

3 ! 

a* o 


■ i 

S e 

a> — 



• Noonameena 

: Pollcemans Point 


u Tea Tree Crossing 



w „ 



i f r 











Tig ID, Mrun relative ba\ch fecundity ot temnlL- Atlurituntwitt irwrnsiomo from three sites in ihe Coorong Lagoon OVei 
ihc |$9QJ>| .in,i i**vi-'/2 breeding seasons Point* represent mean values of up to 10 fish and vertical lines are one standard 
deviation each side "I Hie mean No standard deviation is given when the sample si/e was less than 10. Only fuh with 
Maturing Vt ripe oocytes were included 111 the analysis. 

1 1 ui.uu\ or mtmtttmm wcmvrawa fn nin owrunc 


species lias only one breeding season pel year. Mean 
gonosomaric indices and relative batch fecundities were 
grCatcSI in November in 1900 and October in 1991. 
indicating lhal spawning probably pesJwd during these 
months. The later onset ol "spawning in 1990 fertflCrfad 
by the later drop ill salinities in that v-ai Thus a 
rc*Ji*v I ion in salinity may heoneofibc envininm.mial 
variables involved in triggering spawning. 

A protracted spawning season may rdleet repeated 
u batches' Ufpggji hcinp spawned 01 .1 lack ol pooiifiiiui 
svnehmny in gonadal development (DeVlwnine NHfi 
Analvvas Of Cgg types shiAv.s that oocyte develop 1 
in ,1, microstoma 1^ "'eouip sviu hrum.ns" x- at lens* 
rwo size groups of oocytes were present 11I the ovary 
of an individual at some tune during die - reprodu J e.c 
eyele (DeVlaming 1983,1 This type fjf &th»U 
development implies that the whole clinch of ripe 
oocytes will be shed over a short period (West WO). 
Multimodal frequency distributions of oocyte 
diameters, as tound 0$r A microstoma in the CpDTbqg. 
are characteristic ol multiple spawning fishes t.Hcmpel 
1979). The presence ol several modes of developing 
oocytes, the presence of remnant ripe pggti in some 
ovaries and the decline in the number of maturing eggs 
per Ush as the spawning season progressed in.Ii u 
thu) .4. microstoma is .1 muiupk-spavwioig risn 
Multiple batches ol egg£ .panned -.iicccssively within 
one spawning period have been lound in other 
athennids including Mcnttlm menitiiu (Cnnovjrf 19*4; 
and M otulrns (Hubbs 1976). Protracted .spawning is 
common among athciimds in Australia (e.g. Jvantsoi! 
1V7K; Prince ■& l'oltei [983; Potter ft tjl ttgftl and 
in Ihe Northern Hemisphere (e g. C'aipelan |955, Oon 
& Hen -Tuviu IOK3; (.amove/ 1984; Middauch c^ 
Hemmer 1992). A four month protracted breeding 
season over spring parallels that found for this sfWr&i 
from Dee Why Lagoon, New South Wales whefC 
spiiwiiinj: occurred from August to November (Toller 
ft ol I98t.i. 

influence oj Suitnih on lite Rejourn,*?*^ Pcrfi >)>?,:■;>>_ 
of Alherinosoma microstoma 

A. mnrasiomo is the only iccorded Australian 
aiherimd to reproduce in hypeisaline waters, with the 
possible exception ol Croh'OMVf>luilus fniiHirod'niih 
(Lcfl&ntOn 1977)- The gradient in salinities alone the 
Conning did not reduce the length ol the Npaufture- 
season in A. microstoma as fish were- in spawning 
condition 1rom September to December in both 1990 
and 1991 a.t all sues Salinities during these pcooj- 
1 aue,ed from 32 g I ' tit Noonamecna in September 
|9')l> in 74 >: l ' at lea 'free Crossing in December 
1990. Salinities arc usually at their lOWfisi ie the 
( 'i tuning during spring which may account for I he lack 
ol any cleat influence ni salinity on reproduction m 
the present study. 

Gonae developmer.i commences in late wiiiler>eatlv 
spring di a lime when day length and temperature art 
increasing and these arc liki l\ 'o be the cm 1 mume ma! 
cues for gonad recrudescence. Brcedme in spring nmy 
he tuned to lake advantage of seasonal peaks in food 
ftVtlilttb'litf In ritB Coonme_ hurdyhcads feed mainlv 
on /ooplankton, ill particular uslracods and copepods. 
and these are most abundant dm me winter and spring 
in the Coorong when salinities ate relatively love 
iGeddes 198?) hemalc hardyheads only develop a 
ponton of their eijgs atone time, suggest uuj that either 
a physiological OrteOlOgWA] foefor (e.g. insufficient 
Fo.rtd) limirs hatch lecundiLv. Relative batch fecundities 
and gonosomuih. indices were lower in 1990 when the 
ejowth ami porturmanLC ai~ Httpplli luhcrosa, a key 
aquatic plant in the *i1*lhe*l CoOMngi w '^ reduce.) 
compared with 1 C )9I ( 0"her resources mcludiii.e 
/ooplankton may also have been lower. Mulliph 
.p.twnin^ is a common reproductive strategy amone 
>mall fish species and maMinhcv the numbers of ege. 
.i small fish can produce (Potter pcrs. eOmim). lit 
jdduion. A. mnn>\tomu may be able to adjust batch 
M/e m response to environmental conditions. Such a 
stiiirouy WOUld allow haidyheads trtevploit txith longer 
reproductive seasons and b c itci '|Ualitv seasons. Thiv 
suaicev WWlFd require the oiittaLion of more oocyle*- 
than would be evpecied to be shed m musi yejrs. 

ll<t fiffle ^f AtherinosniTta microstoma in itn Coonmy 
^IheTtmiJ^ a.-, -ja impurunt con ( poneni of ill-, 
bioniass ol small lish Ol mkiiiv Jtjuane ccosvsterns, 
uieinding Bardawil Lagoon (Hen -P.ivia ifel, t.a^uiia 
Madre iHedgpelh I a f.t7)- Hamelm Pool (Lenantou 
1077k iVel Harvey esliiarv iRirierer///. W83uiudthe 
SwaiVAotn Kivci systca- fPrmec & ftmer 1^83). A 
uiictv.siotnit is $ prorninenl component ot the C0OfO0g 
ou.vstem. in pankular us a major lood item for 
•.elected piscivorous water birds (Patou ls*X2t. Hivb 
and fluctuating salmincs along the Coorony did not 
U'aw a narked elteel on the population ceoloyy or 
reproductive behaviour Of A. >nnro\totnu ^\6 no 
SiglllffcHnl difti*-r«-ne^s in reloiive batch fecundities were 
tounj between sites thai iffteied in salinity within a 
vt m. ri*as. 4_ ,/uto^stfino appears m be well adapted 
to hvpetsalme cundioeais and lumre small changes m 
saliinl> vvuiild tun be expected to arted. haalyhcad.s in 
the Cooron«. Howevr,. tUeo hxtd -apply may be 
affected by salinity. Future studies should be directed 
tuward the diet oi A. Nttvrotovrtta ill the Ctldmng and 
the idc!ur.s that mijiht limit these resources. 

The agricultural areas ol the Upper Se»uth bast of 
South Ausimha suffer (row dryland salini/alion arid 
sLasonal inundation tjf lawlying areas with surplus 
i 1 r.rcc water. Theri. arc proposals to drain some of 
this sur'ace water and possil.K ^lunndwaier into Lhc 
Soulti Lagoon of the C'outuny (tipper Soutti t-aa 
Dryland Salinity and FlOOd Maiiauemeui Hlai. 5t( « on 



Committee. I99.V). Such inputs of fresh or brackish 
water may disrupt the seasonal fluctuations in salinity 
which are an important feature of the Coorune and 
which may act as a partial cue to spawning in 4 
wiiinMema. Oiher consequences of ilie inputs, such 
as an increase in water depth and consequent reduction 
in the exposure ♦>!" the highly productive ntud Mats. 01 
a rise in nutrienl or heavy metal concentrations mas 
also affect the food chains in the Cooronj* supporting 
4. mii mwma and m-turn piscivorous birds. 
Consideration needs to be eiven to the role cvi 
hardy heads in the Coorong ecosystem in Inline 
management proposals. 


We thank Damon Oliver. Colin Molsher, Alex 
Weissnuinn and David Pcakc-Jones for field assistant-l- 
and Di Keith Jones tor helpful comments on the 
manuscript. This research was supported b\ a grant 
loi die Murray Darling Basin Commission The 
Department of Zoology in the University Of Adelaide 
provided laboratory space and logistical support. 


Hi \ irvh. A U9S4f Some aspects of ihe miinagenieni of 
Haidautl Lagoon General fisheries CiKtiuH /or ihe 
Mediterranean Si. 529 -540. 

t. \pwAi i \pi ft, 1* I. & Bai.khoi.isk, G. N. ( 19H3 » *A yuide 
in ihe freshwater fafi of Victoria" I Victorian Government 
Printing Office. Melbourne). 

( aiu', L H (1955) Tolerance ol the Sun rrancis..o 
topstnelt, Arherinops ajjinis ujluv.w in conditions in fflJl.1 
producing ponds bordering Snn Francisco Bav. California 
Filth and Game 40, 279-284, 

C'ihnsvhn. li, O A WnriAMs. W VL <!M74j Distribution nl 
ttsli in inland saline waters m Victoria. Australia. 
/ Mar f-'reshWiiter Rss 25. in 7 172. 

O'Ntft'i'-M. IX O. (l°S4i Adaptfvr^ipnifiL-itncLM-i teiiiperarurv 
dependent sew determination in a fish The Atnerieaa 
Xanavlisi U.\ 297 lQ 

<19K5| HicM and laboraloty aists^piBnl o\ pattcoi- 

l(l fecundity of j multiple spawning lish: the Atlantic 
Stlvv-i^itlc Meiihhn nwnidta. fishery Bulletin SLV .W4I. 

& Kvnako. B. E. <l98lt Environmental bCH 

determination: Interaction ol" temperature and genotvpc Ul 
a fish, *etem>- 213. 577-57« 
Dm amino. V. (1983) Oocvte development patterns and 
hormonal im'ul.vmems ailtorm tc!eosi:<_ pp 176 1 99. hi 
Rankin. J. C. t Piicticr, T. J, £c Duggan. R. (Bis) "Conlrol 
processes in t"e-h physiology' (Croom Helm London) 

Gkus.sman. G. & Oiai'man, I 1 . <I982.) On the use 

of the vs>nosoiiiaiK index t 'tnttft Han h,trt f-'fnstoL 73A 
M 59. 

Uppei South East Dryland Salinnv ami Flootl Management. 
Plan Steering t 'ommiftcc (I993t lippcr South East Dryland 
Salinitv and Rood Management Plan. IVah l*iiviromnc-utal 
Impact Statement Natural Resources Council of South 

GhuoKs, M. C. (|0S7» Changes in salinitv and in ifn- 
JiVTinuiionof ma^ophyies, macrobeniho* and bsh m ihe 
Cooruiip Lagoons, Souih Australia. tolUvwine a peiiod ol 
River Murray flaw. Trans R Sin S. Au\t «1. H} ISI 

_ & Bi'llf.R. A. J. (I9S4) PhysicochcMiicai .aid 
bio)oj» ( i,;il studies on Ihe Cooivrtg lagouns, South AusUtiUji 
iind the etfeei ol vtlinitv <»n the JUlribllliDH of ihe 
rnacrohenihiis. /hit/ 108. 51 ')2. 

& Hail. D. (199$) the Murrav Mouth and Cwtanmjj 

PP 20I-2I4. hi Maekay. N, <V Ha.sihurn. D, tCds) Tin- 
Murray" i Murray Darting Basin t'ttinmission. C'anbeirar 

Qnfc, O. &; Bpn h.'via. A. ilvSAj The biology of BtfVe* f 
Sandsmcli Aiherina htftvri Kisso in the Bardawd l.aizoon 
on Ihe Mcdiierrjneii" c<>i«s| yf Siftai, 7 /'o'?. n'f'V. 22 

537 >47, J. W_ (1967) Kcolnpieal aspect^ ot Ihe Laguria 
Madjc. A hypersaline esiuarv pp, 41)8-4(9, hi LjuII, (J. H 
<Hd k 'Ks'Lurtcs" (American Academy for Ihe Advanct- 
nienf of Science. Publication 8J. Washington! 

HiMhii. (., il9?9) "l-arly lite history o! marine fish' 
lUniversjiy ot Washington Press. SealtJcl. 

Hi nos C (197M fhediel reproductive pattern and Iccnndih 
of ,\ff<ni,iit: ifudens. Capeia 2, ^B6*3H& 

Kinnl. O. (1964) The ellecis ol lempefiUurc and salinitv mi 
marine and brackish water annuals F'arl 2 Salinity und 
temperature, salinity combinations. Chtann,ur Mar Biol 
Ann Ktv. 2 2M1-339. 

I ! vwTo-w, R, C -I. 11977) lashes from the hypersaliiu- 
waters ol "ilv* str-inirilohte /one of Shark Bav. W.A. Li,'f>em 
2, 397-55l> 

MrDoWMt. R. M. (I9K0) ^Freshw.iier ffshej ot "South 
extern Austral ia' (A H & A. W Reed Ply L(d. Sydn-'y). 

Mioo-v (ai. D. P. &. HrMMi:k, M, J. (1992) Rcproduciivr 
Bttofegy of the inland silvciside. Memdiu benliina. (Pisces: 
Alherinidaei ironi Dlaekwaier Bav. Hc-rida. Coptux 11 


P\nis, P *)982) "Bioia of Ihe ( oorone" (Souih Aasiulian 
Dcparrmenl nl'Huviroiirncnt and Manning. Publication No 
55, Adelaide) 

Polit-J/., I. C. LOMikAOAN, \ \< LhNAMViN, R. C. .1 , 
t'HHVSIAl. H, J iV (iK^Al, (' I ( n>S 1 ) AhntKhu, ■ 

dlslribulion and itec sttucturc e^i lish populations in a 
Western Australian estuary, ./. //.W, Ltmd 2<MX 2J 5U 

KamsoU- W. ( \Mtkory. R. & MlWMAKp. 1 

il9Sf>j I ite cycles and disinbutii^ri t »l .uherimds in lhfi 
marine and esiuanne wulers of Aimthern Ausitaha 
llyih-t>hiah.>gia 139, 25 40. 

Princc, J. D. & Poi TfiE. f C fWfcJ) Ule-cycie durition. 
growth and spawnine tunes of live species of Alheriiiidae 
ffeleostei) found in a Western Ausiniliyn estuarv Httli) J. 
Mat, t-'reshwi Res, 34. 2*7 JU>! 

, Li.NAN'fON, R CI. &: I.OWUM.^-., N. R 

< 1982 > Segregation yrjd teedmy ot alhenmd Specie 
-lleleosiij m ; .ouUi wesicrr. Au^lralian eMuanes. ibht 3J 


Snu-sos. I> G. t V GfM-i-Ji. C (K5iS) Ni>tes .m huhitat-, 
kystpinatic character and )de histories ot Texas s/ill water 
cypnnodonies- tkiftH* SfuJn \ 7*u>L 4M). 115-134. 

Wnsr. O. 1 1990 J Mdhtwis nl !>M's^iie, ovui ian developmeni 
in fishes - a review Ausi J- Mar Freslin: Res, 41. 199 222 

Wiiiiams, W, l> < 1986V Conduetivitv and salinitv of 
Australian Salt Ute that 37, 177 1X2 




By Mark N. Hutchinson*, TiuMiLNEf & TimCroft$ 


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 
Key Words: Tiliqua, Scincidae, lizards, morphology, natural history, conservation. 

}mmtuti<>n\ *>] thr Rmut Society of S. glSfftfc flMli 217 >2<> 


by Mark n. Hutchinson*. Tim MiLNift & TrM Croft* 


Hi.'TCHINsOn. M. N,. MlLNC, T- & Croft. T. U994) Redescnption and ecological notes on the pygmy bluetongue. 
lUiquo wJeteidemis (Squumata, Scincidae). Trans. R Sm. R /Jw.vr 118(4), 217-226. 30 November. IW4. 

A preliminary study of a population of the pygmy blueiongue, Wiqua adekudensi'; (Peters, 1863) lias enabled 
us 119 redesvribc the species and give a preliminary account of lis naajral history. Life colouring, intrapopulation 
variation., sexual dimorphism and general morphology of! he skull and mandible arc described. Fvpmy bluelongucs 
at 1he study site are diurnal inhabitants of open tussock grassland and use spider holes lor shelter, Males had 
enlarged turgid testes during spring and a female examined at this lime had yolked ovarian follicles. Males were 
more active and trappable than teuiates during spring, but both sexes were sedentary during laU summer-autumn. 
Utters ol" 1-4 live young were born in the maternal burrow during February-March. We suggest that the lack 
prf sightings oH T adelaidensis this century has been due partly to its specialised ecology- and partly to a real 
decline attributable in habitat destruction. 

Ki i Wokus: V/itftui, Scmeidac, lizards, morphology, natural hnlory. conscnanon. 


'kontmt tittr fiitf wtuiigem, xttinigem terrain w?r" 
(Richard Schomburgk. quoted by Peters, I8t>3) 

Schomburgk's remark ("found only in sandy, stony 
terrain") is the only published first -hand information 
available on the ecology ol' TiHqua adelaUU ,t mi.\. the 
pygmy bluctongue lizard, u speeies which has been 
regarded as one of the most seriously endangered ot 
Australia's reptile speeies, if not actually extinct 
(Cogger 1992, Fhmunn 1992; Hutchinson |992h The 
discovery of a population of the species near Burnt. 
S.A. (Armstrong & Reid 1993: Armstrong naL 1993} i 
following the first sighting ot the species for 33 years. 
has presented an opportunity tor urgently needed study 
of the speeies which had previously been known from 
only 20 museum specimens, mostly collected last 
cenlury (Ehmann 1982, Shea 1092* 

The original description (Peters 1863) was brief, 
based on two syntypes probably collected in Lhe vicinily 
ot'Gawler, S.A. Mitchell (1950) tedescribed die species 
based on SA Museum specimens Wrf figured the head 
shields and whole animal for the first time. No fuither 
formal descnplions have appeared m print, save tor 
those of Cogger (e.g. 1975, 1992). based on the old 

' Mark Hutchinson. South Australian Museum, PfJ Box 234. 

Adelaide. SA. 5001. 
t Tim Milne, School of Biological Sciences, Fbmh-t- 

University, GPO Boh 2 MX). Adelaide, SA, 5001. 
j Till] Croft. Biological Conservation Branch, Department 

of Environment and Natural Resources. GPO Box no 7. 

Adelaide. SA. SflQl. 

1 Suiu, G, M. t!992> The systematic* and reproduction ,n 
bluc(onj;ue li/ards ot tin.- genus IHiqu.i I Si]uamata: 
Suncidae). Unpub. PhD ihcsis. IVpi ot Vcie'innry 
Anaiomy, University of Sydney. 

and faded museum material. Shea (|990i described a 
number oT scalation and osieological character staler 
lor T. ml e Undents in order to establish the validity 
of the genera Ti/iqmt and Cycfodftmorpin.s. Shea's 
unpublished thesis f 1992' > gives a thorough 
description o\' the species" scalation. morphometries, 
colour pattern and osteology based on the twenty 
specimens then extant. Shea & Hutchinson (1992) 
illustrated and commented on the denutry and dentition 
of T- adeiuicfensts. 

Rhmann (1982} summarised available data on diet 
and external morphology and drew together the meagrt 
data bcurmg on the provenance of the specimens known 
lo him. He also attempted to reconstruct the species 
ecology, using analogies with related or physically 
similar reptile species. These speculations were eited 
several times subsequently, sometimes in such a way 
that it was not clear that there were no direct 
observational data on the subject (fchmann 1992). 

Field work has begun, aimed at determining the best 
methods for locating T. udcluidaisis, gathering 
preliminary data on us ecology and making a first 
attempt to determine the number and size of surviving 
populations. This article gives a summary of the 
morphological variation that we have observed within 
a single population of /. iukkudvri*is and provides a 
preliminary account Of its natural history ai this site. 


Current research on the biology and conservation 
of this species is concentrated on one site located in 
the Burra area (33°41'S, 13S°?6'Lj. approximately 
160 km north ol Adelaide. South Australia. The site 
was the fust of several found to support T adcluidensis. 
following the initiation ol lieldwotk in Oelohci, 1992 


M N HI l< HINKOV l \f)LM & I . KOI-. 

( Armslntrig tt at IQ93) I lit* daui picscnted lien' stem 
hom the first season ol field w<»i4 mu » | n -. jt< ■pui.H t m, 

i arriod "" f (tow nriu Oiacitoar J99: innuutMay rm 

Specimen ColStuunn 

Live A <uhl<iiJi'tt\i\ l..Iu.Ji.\1 fcn 'hrci 

Pilfell imps. ^(1 metre stretches fli iU vm«. 
!• ricijjg were eree'ed, along which were placed si* 
25 em lengths of PVC pipe dug vertically into the 
*dn>»*nd. Four haphnes were KOI at the study ole 
the period 4 Nuvembcr-20 December W2 An 
additional line was in place durmg 30 Noveniber-2u 
I v.. ember and a lurthet two were set on 4 December. 
I he five most successful spring rraplincs were rfc$C| 
during 2-21 February W93, Total trap njgftts by fotS 
method were 1,956, 

The other two DWttflXft "lied on the species" hole- 
dwellinu habits (see fhihiiut section) 

Hand collection. This method relied on opportunism 
stublmgs of free ranging animals, ot on surpi [xjhg 
amiualsal the entrances to burrows Most individuals 
caught by hund were juveniles which were exliaclcd 
from burrow entrances usnle, Ion- lorccps or clamps. 

We !|lso used ,i method we termed *lishmg~ BlIiCHlg 
l^arcK to -,ci/e an insect bail lied to a cotton thread 
A grasshopper on a piece of COtton tied to the cud '.I 
a three metre bamboo pole was held .niKidi |fo 
ttitiaoce of th* 1 livard's Ihhiow; lizards, seizing Ih*. 
eiasstiupper could be pulled into the air and captured 
as limy dropped lo (he ground and tried to letuin to 
tin- burrow (Strong cr al 1W describe a snmlai 
collection lecluiiquc)- I i/ards had to be lured 
completely out ol' the burrow before heme, allowed to 
si fee (he grasshopper, miuc the lizard's hind limbs 
could tiold the burrow run. preventing hsdislodemenl. 
Iticir extreme wariness made the use ol a long pol: 
necessary to distance the collector from ihc lizard. 

Trapping using baited (peanut butter or undines, 
aluminium "Sherman 1 ' traps was tried but without 
success (approximately 2,500 Hap nights/. 

The time of day oi collection W0$ noted, .is was Uic 
dads MiaMiiM.m luoperaiure. Lizards were measured 
and sex was determined by the presence of bulges di 
tile base of the Iml and/or Itemipcncs in males- with 
a lack ol these Icatures indicating a Kittle Ihc I . . i 
were then marked by toe clipping, photographed and 
teleased. Only one or two uwf- wen. kim.-,. d I 
limited number in COtTJUDCliotl weh flour paliein 
VariWbui^ being ftlflftuiCfll to iUenlily each individual 

1 united use was » of radiotracknlg using a small 
eM.etuul UUll supplied by Transceiver IServi. es 
Adelaide, attached using superglue iind Surgical tape 
i |hl lizard's shouldct Ihc device WUH intended rot 
shnn term use. and was run successfully on a tirnglc 
lizard located every two houis. fol six day-.. A SdCQTUJ 

'ml resulted irt the death of the lizard within the fitM 
two-bout period and use of this approach was 

In the des\ riptive section, scale Icatures occurring 
bilaterally weic counlcd on both sides. Paravertebral 
scales were counted using Greer's (1982) method. 

Vegetation was quantitatively assessed aiouud the 
most successful pitfall \hk using two methods: 

O) We recorded all plant species in a 30 m >: 30 m 
quadrat, 15 m either side of each pitfall line, taking 
.olleetions tit plants in cases where identilv was tn.i 
clear. I hese collections were subsequently identified 
and Indeed With the State Herbarium. 

b) vve assessed vegetative cover withtn the 
10 m -• 30 ra quadrats by the simple transect methyl 
known as "Step-point" (Cunningham (9J5). At e\ci> 
sicp in set directions ai.ross the quadrat we recorded 
(he plant sp+xtes encountered at the tip of the shoe. 
scoring approximately 300 points in the quadrat. This 
method gives estimates ot percentage of ground cover 
of plant species, bare ground, ruck and litter. 

The behavioural data vve present stein mainly Irom 
licld observations, but we obtained confirmatory dalu 
in many case* from observations of captive specimens 
ai Adelaide /«>o if Morlcv and R. Ainsley pers 

cumrn i 


Spmtto m 

The rollovvitig descriptive sectinn is Mtteudcd '■» -liuv. 
variation witnm a $fl\gte population. K.arber auihors 
(fhiuaun i'>83; Mitchell 1950) give additional data 
b.wu some ol the oldet specimens, and nil of the 
available data on Ihjs older material is discussed in 
Shea's thesis. 

A total of 63 specimens was collected during Bw 
mm \-r\ pefTiod. and an additional lour were horn tn 
captivity. Six ol Ihexe specimens vvere collected dead. 
tilled by Ibc clapid snake tkBUtfohaja fi^rifh. 
specimens) or birds ol nivy (two specimens). A 
seventh, R4U83S dieii dnrmg li iah ..| an external radio 
transmitter. FflUl ^CCfmcflS, an .mmature. an adull 
matt- atld two adult lemales. were retained and are on 
loan lu Adelaide Zoo. The remainder were marked ami 
rtlcaftcd The success of collection methods varied 
according to lime of year f'ittall rtapptne wa- 
successful only during November-December, while 
hand collection became much more successful in 
February -April ( fable 1). 

All prcdaior killed specimens are damaged, although 
in two the damage was minor and confined to restricted 
arens of Ihe body R4CK87 and R40744 had some 
ui... [Vitic patches on the dorsal a/id ventral body surfac 
R41M387 had a sagittal trauure ot ihe skull running from 
the trwtrtl sl.iIl lo ihc level of the eye. R40689 il«» n* icek and posleo'lalcral region ol tht 
skull dicpl\ ,i:on>:cd atid punly removed and a deep 



wound in the right side trf the neck. R4072X lacked 
(he head and right forehmb. as well as ihe liver, heart. 
lungs and stomach. R40738 and R40745 were partly 
digested, with Ihe skin on the body sloughing away 
and much of the internal soli anatomy missing. Two 
skulls and associated mandibles were prepared Irom 
snake killed specimens, one articulated fSAMA 
R40738) and one partly disarticulated (R40745) 

Tm<i it |. Rftuinr siiffcx* of capture icrhnhfucs. 


Number caught 
November- rebruary 

December May 




not used 


"All but one neonates 

Gitnrttl Appearance 

Tiliqua adeiauknsis is a moderate sized skink with 
short extremities, a relatively heavy body ^nd large 
head. The body and tail are soft and flexible, but Lhc 
head is heavily armoured by the well developed head 
shield osteoderms. The toes are shon. the third and 
fourth toes of the hind foot being equal in length. The 
tail tapers rapidly from the base and is thin and slightly 
lalerally compressed over ihe distal two-thirds 


The dorsal surface of the head, body, limbs and tail 
is light grey brown, yellowish brown, orange, tan or 
Choeblalfi brown, the distal portions of the extremities, 
especially the forelimbs, being a paler yellowish hue. 
Dorsal and lateral scales usually have narrow darkei 
edges producing tine, longitudinal lines along the back 
and sides. The lower lateral surfaces are pale greyish 
becoming oi'f-whilc ventrally, The dorsal surface is 
unmarked, or shows variable development of blackish 
spotting, including a vertebia! series of irregular small 
blotches (which may coalesce into a ragged vertebral 
stripe) from the nape to the base of the tail, and several 
laterodorsal and upper lateral scries of small hlack 
flecks; these may be crudely aligned to form weak 
transverse bars. The midlateral region often lias 
scattered grey -white flecks. The venter is immaculate. 
or with slightly greyer margins lo the scales forming 
narrow longitudinal lines The iris of the eye is bright 
orange. The tongue is pale rose pink, with no trace 
of melanic pigmentation. The roof of the mouth and 
buccal commissures are mauve. The abdominal 
peritoneum is black. Juveniles are consistently greenish 
grey to mid-brown, becoming reddish tan on the Lail 
and limbs. The range of variation in black pigmental ion 
is similar to that seen in adults, hut many juveniles 
have more extensive and obvious white spotting on the 

Ip 5 .;* I ■■' 

Fig- [. Head shields of Tittqua adelwJensts (SAM A R.40838). A. dorsu, view H, left luicral \ lew C , detail of chin sbi< M- 
D. detail «»f right side showing asymmetric variation in circumoeular scalation. Abbreviations for cjreumocttlur seah*: 
po. poslocular- prsb. presuboctilar; psb, posisuboeular*. sh, suboculars; sc, Mipraciliaries. Scale bur - 10 mm. 



RgDESCRlPTION AMD \'(»iks on fiiftjt'1 MHfiint-ssts 



This is based on eight specimens, four held in the 
S.A Museum <K4(J687. R4Gfjg9, R4tl744and K4(IS^i, 
plus ,i live adult male, two adult females and subadull 
field at Adelaide /oo. The head shields ate shown in 

IV. i 

The species hits (lie scalaiion chaucteristics listed 
by Shea (IWO.) for Tduftut. The last is 
horizontally divided, and the temporal scalahon 
posteiiur u> the secondary temporals is irregular and 
hide ddlercniialcd (mm the body sculalion The 
supraciliary scale count is reduced (mode Si A 
complete subocular scale row. consisting ol a laree 
piesuhocular. a largei anterior and smaller posterior 
subocidar and one or two poslsuhoculars. runs from 
the anicrioi supraciliary lo the postoeular. Nuchals are 
absent or a single slightly enlarged pair may be present , 
ohen separated by a median occipital or tnternuchal 

Dorsal scales smooth, in 3.V36 (mean U.h) rows 
at nndbodv, paravertebral scales 77-86 tinciai XvOi, 
paravertebral scales between parietal and the level ol 
aiiletioredge ol hind hmt> 70-81 (mean 77rM; SUfcfcllgflal 
lamellae under fourth toe unpaired. 10-13 (mean 11.6,1. 
nasals in point contact or narrowly separated (0-2); 
strong postnarial jiroove present; prefrontals in broad 
contact; frontoparietals paired; interparietal about same 
si/c as frontal, much larger than frontoparietals and 
separating parietals; each parietal in contact 
aiUctolateraliv with the postoeular and posterior 
supraciliary and bordered posterolateral!)' by four or 
live enlarged scales: a median occipital scale present 
or absent <0 4» posterior to interparietal: supraoculars 
3/3, the first iwo contacting the frontal: supraciliaries 
5/5 f5rtj iti one specimen), the second the longest; 
complete suboeular scale row present; supralabials H. 
the eighth only half as high as the seventh: infralubials 
7 ( > (mean 7 ty). a single primary temporal is followed 
hv f hive secondary temporals; post mental contact first 
two infraluhials, a single large, quadrangular anterior 
car lobule. 

The dorsal head shields, frontonasal, prefrontals. 
Itoutaf frontoparietals and interparietal, are thickened, 
wan weakly corrugated surfaces and dcepU irifiscd 
siiitit<-s this ni^usitv increases wirh si/e In laJ£C 
males the subticular scales also become thickened and 
shghlly overhang Ihe suborbital supralabials. 

Skull ntul tnandihk- 

The characteristics noted for the species by Shea 
<W0; 1992') are confirmed in the SAM A specimens 
The species has ivh derate I y naivow separation o( the 
pre- and posilronLals on the medial orbilal marrin 
binger-like nasal processes of the fromals arc prcsenl 
tti give a W-shaped frontal-nasal eonlaci. The jugal is 
broad and flattened and contacts the post frontal. \ 
lacrimal bone is absent, There is a well-developed 
medial palatine ptocess of the ccropterygoid which 
excludes the pterygoid from the edge ot the infraorbital 
vacuity The coronoid process ot the dentary is enlarged 
and swept hack to cover the lateral face of the dorsaJ 
process ot : the coronoid. Dentition is heterodont, wilh 
the cheek teeth markedly larger than the anterior teeth. 

In general aspect (Fig. 2j. the skull and mandible 
i»l /.' aJcianh'H^i,\ are very like those 0! other Jiltifito 
species. The proportions of the snout, tapering and 
pointed, rather than blunt im\ rounded, and the parietal 
legion, constricted, rather than laterally expanded- 
resemble those of adults of the other spet (CS ol Tilhjw. 
The dorsal head shield osteodentins are ultimately luscd 
wiih the bones of the skull roof, to a greater degree 
than is usual in mosi other Ilhauti. Thus, even thoueh 
E udeliiulemts in as small as or smaller than 
congeneric neonates, it iSontogenelically advanced iri 
its proportions and degree of ossiftcaiion In these 
features /' udelauienMs is pmgenetic. not neotenic as 
is frequently the case in miniaturised lizards (Rieppel 
l c >84>. Lven so, 1, udclatdcnsis aJso shows sonic 
neotenic. features in retaining rclalively large upper 
temporal fenestrac ami in some aspects ol braim ase 
anatomy (Shea pers. comm.). 

Unique or unusual features of the skull are few but 
include the very closely apposed, almost parallel 
palatal rami ol the pterygoid's In most other Tifufim 
iL $i$Q& is* an exception! the pterygoid margins are 
more widely separated and diverging. The closely 
apposed pterygoids may be correlated with a narrowing 
of the skull posteriorly as /. ttdehudensis has the 
greatest skull width across the jugals, rather than at 
the level of (he ijuadraies. The maxillary process of 
the jugal tapers rapidly and fails to contact the 
prefrontal- This character state is shared with 7" 
unthifUst-'ttiu) and most /. tu-tiftindis, whereas OiflOf 
Filn{na species have the jugul eMendme untenorly to 
contact the prefronlal adjacent to the lact imal loiamen. 

Rj£ 2 Skull and rmtnilihlc ot T\tU)ua iuietauJcti.sis (SAMA K4(THX|. A. skull, ri^'lit lateral view, showing conuci between (juy and r»i]u;iUi»>s»il (si|l h»>ncs, B, sKail, dorsj) view, showing sioit nusilljiy pru<.evs of the iuiial (p. mas > and 
QpCli hUprat_eni|Kiral feneNlrau (slfi. Nme pe^islcnl head shield i>.\tei>cJefTUs v\ Inch tjhscure sutures t>t Ltnd puslfmnial 
botVH! olhci ikuH mi>i sutures life visible ihroiuh the tuie^dcrms <\ ventral view, shovvmp vtuUivuuilum of ihc palatal 
complex ice. ccutpterv^MKi: pi palatine: pt£, pu.TV^oitf I - D. rsgjft niandihuhtr nirtfU*, tahtai view. Khuwnu vers well devvi"peJ 
L*qrOrtt*kl prDCi&y Of we dentals (p. wk ) E, Kfl uiuruhbutar rUnUW, view Scale lur |o ntm, 


The upper temporal fcnestiue are relatively larger than 
in adults of other Tilitjua species, in svhich (hey are 
slii-like or absent. 


Based on (>1 specimens. Snout-vcnt length (SVL) 
38-107 mm tmean 732). Axilla-groin length lAGL) 
2.V67 mm ( mean 44.0. n = 58). Snout-axilla. distance 
(SADt 16-41 mm (mean 28.3, n = 58). Totviimb length 
11-21 mm Hind limb length 11-21 mm. Tail length I TL) 
22-79 mm, 

Sexual Dimorphism 

Adult males and females (arbitrarily defined as 
>85 mm SVL) differ markedly in head and body 
proportions Adull males on average arc shorter than 
females (SVL males range 87-106, mean g 3.4, n - 17; 
females range 88407. mean 98.3. n — 14) but head size 
does no) overlap (SAD/SVI. males 0,36-0.4"'. mean 
0.38, n - 17; females 0.30-0.35, mean 33. n s 14). 
I he relatively large head size of males is a trequeni 
phenomenon in skmks fe.g, Sunboiwc 1985, 
Hutchinson et ai 1989; Hutchinson & Donnellan 1992) 
but is rarely as marked as it is in large adult male J 
nckiaUtensis. Male combal has been recorded m other 
Dliqun species (reviewed by Greer |989) ? a selective 
pressure which might lead to their large head to body 

Ecological Notes 


The main study site lies at an elevalion ol abou! 
500 m on undulating terrain cut by small, nueiiuiltcnt 
stream courses. The ground is stony in places, with 
underlying shale and sandstone bedrock just exposed 
0J1 the surface. The soil is hard-packing clay-sandy 
loam (red-brown earth. French ft ai 1968). The 
vegetation of the main trap site is characteristic ol 
degraded remnant native grassland, wilh rhc area 
around the Site being bare of trees and shrubs (Table 
2) A lull list of plant species is provided in the 
Appendix The exact original nature of this grassland 
is unclear, as copper mining and farming near Ihe site 

T ski i 2 OronmJ jxvrr <// Tilitjua adehmlepsr. vtulv we 

# Cover 




Native Annua) 










Bare G round 




i ttwt 


since 1845 may have resulted in ihe removal ol trees 
and shrwhs for both industtial and domes-lie use. 
Similar hilly areas m the district support AltacaMUitvui 
vi-niftlfata (drooping she oak) low open woodland 
over similar ground cover of native grasses iStipa spp.. 
Dtw.ttu/niti spp. i and mat-rush or "ironvrass 
[Umtaruira spp. L as recorded in the survey area 
Jessup f 1948) concluded thai at least some of the area 
had probably been essentially treeless prior to 
European seiliement. 

The relative abundance and species corngoSttJQn 01 
native grass species varies within the study area. The 
immediate vicinity of the most productive traphnc was 
dominated by one species of spear glass {Stipa, 
tentatively identified as S. etTWftf>hi(<n. but elsewheie 
in ihe same paddock, pygmy bluetongues were found 
where 5. nodosa was Ihe common spear grass, and 
other grasses, notably wallaby grasses. Damhonia 
spp., and wire grass,'Ja hehriona, were locally 
COftUflOft, Thus the piecise species composition of the 
underslorey may be less important for Ihe survival of 
pygmy bluetongues than the tussocky Structure which 
provides ground cover throughout the year Othct 
areas surveyed in nearby paddocks which do not 
appear to support pygmy bluetongues showed 
increased ground cover by introduced plant species. 
and hence a decrease in the amount of covet during 
Jate summer -autumn r.vcn at the least distuibed pari 
of Ihe study site, only 50% of the ground c»vei is 
perennial vegetation, so that the surface of the ground 
is tar more exposed in autumn than in spring. Adjacent 
areas dial have been ploughed at any stage show 
iiunimal reeolomsaiion by native plants, in particular 
hmnmdni spp 

Aside from a single luvcnilc found under a stone, 
the only mtcrohabitats in whiten 7 adrlnidrnsis have 
been found sheltering are vertical or near vertical holes. 
We suggest dial the lizards arc using spider holes, not 
digging iheir own burrows. The holes are* perfectly 
circular up to about 20 mm in diameter, lack any sign 
of excavated sod at the entrances and are 
indistinguishable from holes at the study she inhabited 
by lycosid and mygalomorph spiders In at least one 
case a h/ard inhabited a hole to which the lid of a 
trapdoor spider was still attached. Two large species 
Of wo|t spiders. Lyt>*sa slirlititfi, which makes a lid tor 
its burrow, and Lycosa pjlbcrta, which docs not, have 
been identified at Ihe study silt as has a species of the 
trapdoor spider genus Bluktstotiia (probably B aurcai 
Lycosids and their burrows aie very common in the 
area, The lizards appear lo make no obvious external 
modifications lo the holes, save for a slight bevellmu 
ol the- edges caused by their coming ;ind gOlUg 
f/nmr mux? and rtU'VfftU-nt.s 

We obtained lew recaptures, and made only limited 
use ol radio-tracking, so (hat our result* arc 


The urea covered by a male that was nidiotracked 
durme spring encompassed 70nr and two butiow\ 
over a period of six days Home range overlap occur;, 
as there were at least Iwo other occupied butiows 
within (he area rhuii the \u:\n\ covered A second. 
smaller male was fitted With the Hacker, hut wheu 
relocated after the first iwo hour period ii was 
moribund, appuieully ditough exhaustion as it 
struggled to Ibrce itself into a holt* but wift prevented 
by (he bulge of the transmitter. It wn$ striking that, 
although shelter m die form of dense glass and 
Lumatidut ttwSOcks was nil around, the lizard put all 
of its ellons inio finding, shelter in a hole. 

The same male successfully (racked in November 
was caught again during March. It was still within (he 
same aica as IflK previous spring hut in yet another 
hole. In another case, a burrow containing a female 
and young was abandoned and the female was" 
discovered in another holt some 5 m i'rotu Ihc original. 

(July one ot 22 captures during November 
December was an adult female. AH animals captured 
during spring/early summer were caught eilher by 
pitfall trapping or by hand, and so required the 
individual to be actively moving away hom Oil tun row. 
However, of the adults captured during February- April 
one fa female) was captured by hand and 14 by "fishing" 
but pit till I trapping caught Tame, despite in excess ot 
thicc weeks intensive piUalltng m areas known to 
support a significant number ol pygmy bluetnngues 
Of the adults caught during sumuiei/auiuuin. 12 wctc 
Inmates, and three were males, a reversal ol the 'remt 
shown in spring/early summer 

The lop-sided sex ratio in our spring sample tonly 
a single adult female caught) suggests much greater 
levels oi male activity during, the spring mating season 
This is supported by the six predator kills collected 
during October November, ol which five were sexually 
mature males, while only one was an adult female- 
Males ot other species ot litiifuu have increased 
activity levels relative to females during ihi> time of 
year (Bull vtuL 1991). l he tesults so fat indicate thai 
both mates and females may have relatively hunted 
home ranges throughout most of the year, but ai spring 
mates appear to wander inoie widely (or at least. inoie 

During Novcmber-Deeembei I'W, lajufai) was 
above average and daily maxima ranged between 1*> W (* 
and 36 P C Days when lizards were caught had maxima 
between 2()°C and 36%'. and die time ot caplute varied 
bom 08:15 to W:M. All of the days on which pygmy 
bluetongues were caught were samny at some slip. . 
and so ground teiiipciaiures often would have exceeded 
the air temperatures. Traps were monitored alter dusk 
on warm nights to cheek for noclurnal activity, but no 
lizards were captured ai this time Diurnal behaviour 
was also observed In captives, which showed ipl&tgll 
ol noeiurual activity 

ON JlKtfl •' U^tAtDKSSIS 22 l 


K40687 contained a huge lepidopietan latva (a 
hadenine nocmid, probably f\-r\oi*iw<i, G Drown 
pets, eottun.); K40f?8M Itad the icuuuus ol an 
apparently identical larva, plus several leaves unit 
flowcis of the hetb Mt-iiutwo- R40744 contained rim ■< 
acridoid grasshoppers and the temains oi a small 
beetle, fchmawi < l$82), after examining spec imens then 
in the SAMA collection, tecotded cockroaches, ants. 
a spider, grasshopper and beetle and some plants 
\LMwulla seed, possible chenopod mateiiaJ). Wild 
li/ accepted grasshoppers ottered .is bait, and 
captive animals eat an omnivorous diet, including 
mealworms, crickets, chopped fruit and vegetables ane 
law egg. Thus the evidence to dale indicates that '/' 
mft'ltiithtnt* eats a wide variety i*( invertebrate animals, 
but also includes plants in its diet, 

The relative lack of movement away Irom (he 
burrows, at least in late summer-autumn, suggests thai 
ai this time ol yeai /! luh'taiiicnvis is probably a sit 
and wait forager. Burrow emranees are used as vantage- 
points from which li/ards would be able lo make shoh 
forays after any prey detected nearby The presence ol 
leaves and Jlowets m the diet suggests wider foraging 
al least in spring. The study site suffered an outbreak 
of plague locusts \Ch* *t(f*n tics {crminiftm) duiuig tl|C 
period of observation which may have influenced the 
lizards' behaviour The sedentary beha\ tout may also 
be related to avoiding predation at the time of year 
wheu ciouud eovei is sparsest. 


All r»t the predator killed males (eollecled between 
14/10/92 aud 30/11/92) had lesles iliac were enlarged 
and turgid or starling to regress One ^' the trapped 
males had strings of dried seminal matettal protruding 
Irom the vent, The single dead female (R40744. 
collected 7/11/921 had lout enJaiged ovaiian follicles 
(iwo left, two rii'.ht). Together these, dam indicate a 
spring testicular maximum, with spcinuogenesis and 
inalmg behavtout coinciding with ovulation. Oilier 
t'llttfmi sjkvics synloptc with T. tuletu'uictisis, f, nt^asn 
and 7 scitit t>n1cs, are known to he spring breeders 
(Bourne <7 «//. 1080: Shea IW) 

Neonates were first observed ft] The Held site on 28 
Pi'bruarv |99$, when fonryoiing were ohserved in the 
but row of an adult temate. Inspection o| other burrows 
known to harbour females revealed olher lilters, litter 
size ranging l-i irnean 2 J). Ail juveniles esanuned a? 
this tune had raw pmk umbilical .seais uidieatutg very 
reeenl birth Two temales which pave birih at Atlelaidi' 
Zoo iku'h\£ mid-March had liner sues of tlnec and tmc 
WiUnn tvvo weeks ot their^eiy. juverules and 
sometimes adult-s abandoned the birth burrow Juveniles 
were observed from late March alone in holes. 
sometimes much smaller than those used by adults 

The smallest animals caught m November were 
06-70 mm SVL. while die laigesl young with disiinca 



umbilical >cars in May were 56-60 mm SVL This 
implies * hat the 7, aticIaUknsi\ bom at ihc en J ol 

summer reach about 70 mm SVL by the end ot spr jng 
ol that year. This SVL is considerably smaller than 
the smallest breeding female recorded (SVl- % mm), 
so that first breeding must occur HO earlier than (he 
saontl spring (appro*. 20 months or age). 

Tihqua luh-ki'uiensn appears to be extremely 
sensitive Iti both movement and nose, making u 

difficult to observe lizard* busking uvteidc (heir 

burrows. Lizards bask with the back leys or lip ol (he- 
tail remaining in the entrance of the burrow From this 
position, they can back rapidly into their burrows it 
disturbed- Onee inside, the li/ards increase their 
security by turning the head sideways to force the snoui 
and occiput against the sides t>f the burrow. Thus 
wedged in place, the lizards present only the armoured 
head to an intrude! and seem almost invulnerable, any 
creature small enough to enter the hole is unlikely io 
Ik Strong enough to harm or dislodge me bzatd. 'I lie 
significance of the heavy osleoderma] armour on (he 
head seems to be associated with the use of the head 
as a combination anchor and doorstop. 

Several behavioural attributes of captive lizards 
suggest why detection ot the species tti Ihe field is 
difficult. The Itrsi is crypsis, When disturbed the 
lizards freeze - such stationary animals are well 
Camouflaged against the reddish brown soil of the area, 
making Ihem hard to see. 1 he second is a leluetaiv v 
to emerge into open spaces. In captive conditions Ihe 
lizards spend almost all of Iheir active time in holes 
oi among Inter. Thirdly, the lizards have a well 
developed .md unusual ability to mtA'e in conttned 
•paces. Both in the Held and in captivity Uiey have been 
observed moving directly backwards into cover or 
dovvn a hurrow. avoiding a L-tuiii with its concomitant 
greater level of disturbance. The small, thm tails may 
facilitate this manoeuvre by not getting in the wav u $ 
they niight if longer or more massive. The bode f* 
unusually flexible:, un attribute which (uoluhU assists 
in negotiating the confined spaces Ol bunows 01 
tussocks. On several occasions, an was known 
to have entered a hole head first, turned around within 
the hole and emerged head first. 

When disturbed or handled, this species has not w 
been observed to exhibit Ihe exaggerated defensive 
display employed b> its larger relatives (Oupcutei & 
Murphv 1978; Greer K>«9|, When handled the I r/ard 
will twist svitti great sijengih and agility, often Rapine 
and endeavouring to bite in a similar fashion to othei 
rm diuiu-si/ed skmks ie,». E^crnia spp i 

Possible Reasons (to Decline 

Wt'jua aMaoUnsis remained undetected ill a well 
Irequcttted pan ot South Australia f^t ovct three 

decades in spite of diligent searching by hcipctologisfs 
Based on our experience with the species, we -aiggcsi 
thai two factors combined to hamper searcher- tack 
of information on its habits and habitat, and a probable 
real decline in the numher of populations. 

Ehrnann (1982). based on his interpretation of the 
specimens and historical data, speculated dial 7" 
iideUtulensis had inhabited a limestone-ehenopod- 
malice association, White the speculative nature ot this 
assessment was noted by Ehmann himself, it 
nevertheless evened a bias on many of the attempts 
to locate the species. Ihe grassland habitat in which 
we have found lite species is difficult to search casually 
for leptik-s, and consequently has received littlc 
nltenlion. This is especially so because most grassy 
terrains in this pan a\ South Australia arc assumed tQ 
be heavily disturbed, supporting only the most wide 
ranging and ecologically tolerant reptiles. The 
distinction between native versus introduced gf&£|{10%jt> 
has been important in locating populations ol this 

Lven when nut- knows the eoneet habitat, the shy 
nature ot the lizards makes them very dillicult to litid 
unless tt is known exactly how and where to §e&rcb. 
The crucial piece of information that allowed us to 
locate additional colonies was the discovery of their 
dependence on spider boles, information that was only 
acquired after we hail 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 d warning of human appniach and 
is out o\ sight by Ihe time the hole has been noticed, 
intensive field work had been going on tor six weeks 
at the site before we observed any /. adeiithlemh under 
natural conditions. However, having discovered the 
combination ol habitat and miemhabitat ■ Wt hHVC bwfl 
able to locate rurther populations ol pygmy biuctongues 
ui nearby areas. The species is now known tti occur 
at six sites running Irom south of Burra to north ol 
Halleti. the extremes separated by about 50 km. 

The apparent rarity, caused by lack of attention lu 
grasslands as a habitat, is probably coupled to a drastic 
reduction in abundance caused by pasture improvement 
and cropping. Native grassland similar to that ill which 
the Spccjcfl occurs at Hurra once extended south on 
to the Adelaide plains, bill & prune agricultural bud. 
was one of the first major habitats in South Australia 
io be clc-aied and ploughed. Ploughing permanently 
alters ihe vegeturion and ground cover, converting a 
native, largely perennial flora into an introduced, 
largely annual one. In addition, ploughing would 
destroy the burrows, killing lizards directly and" leaving 
the survivors viihoui spelter and at the mercy ot 
predators The fact that most specimens were collected 
last century may simply be correlated with the tact thai 
this was when most of then habitat was being ^onwnni 
If rjiiculiure. 

RI.UI.Sr'klPllON and Norr.soN thjqua ADKLitOMSIS 


Remaining native grassland is now extremely patchy. 
and the remnants have generally been heaviK invaded 
by introduced annuals. However, populations oi pygmy 
bltichmgues have now been found in some fjf these 
remaining poekets, and provided lhal land use is nut 
Changed, these colonies may be relatively secure. 
Further work is focussing on Uridine the extent ol the 
current range of the pygmy bluetongue, approximating 
population size within ihese areas, and confirming 
many pi the subjective ideas developed during the 
COUfSC <jflhis first SC^SOrtV RfeW work. This will lead 
to uu accurate assessment ol the slutUN of the spc ics. 


We are very grateful |u property owners, parlicularly 
Harry and Judith hberhard, lor allowing access bo their 
land, and to Sieve Vogei lor arranging accommodation. 

This work was made possihle by grant support from 
the Endangered Species Programme. Australian Nature 
Conservation Agency and we thank Si. Stephens 
B, Male and J. Hicks nf ANCA lor then support. 
P. Copley of the Biological Conservation Branch, Dept 
i»l Environment and Natural Resources, assisted with 
advice and ideas, and DENT? 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 Niejaikeand A. Yates. P Lang, Native Vegetation 
Management Branch, helped with plant, especially 
grass, identification, and R. Davies and M. Hyde 
kindlv shared their data on sites where native grassland 
remnams still occuired. D. Hirst (SAMA) identilk'd 
spiders and provided background information on 
them. We thank D M. Bull and G. M. Shea for 
their comments which considerably improved the 


AKMom\<., li, At R»-ui, J .;IW> The n;dis,ovcT) "I Itvc 
Adelaide* pvgmv hlueiongue, Ttiiqua Oiblauivtists (Kier>, 
\thXi. Ut-rjtcfitfaimi 12, 3 6. 

I Km*. .I ft. w &. i-h rcinw.v M. N. (10*9$) 

discovery ol a insulation of the rare scinetd h/ald CilUfuo 
UiklaMttak (fetciM fid Sbitth Au.\t \-lus 26. 5;- 5S 

Kohhm A R - Tauok. .1 I . Hi Wmson\ T. rj. » lOHoi 
Annual cycles of plasma and testicular androgens ih the 
h/iiid Ilfiifwj (Trui-hitlfxnurust ni^osu. Otn i <>m}> 
liutorriuot §fl 27* »J8fi 

fiiiU. C. M. MiNUuv. A, & Dntus. Ci £1090 
Asyrvritooous seasonal activity of mate and female sleepy 
li.'-irds fjUtfW VJ&tXU «t ttttptftfl 25, 436-441, 

('ARflMriR. C C & MliRPItV. J B iN7S| TongUc display 
by (lie LomnKm bluctt>neue, TititfUt/ s. .inctwUv* (Reptile*, 
UceniUa, Seineiduo. ihui 12. 428-42^, 

C"(,f;iu, H. G. (1975) "Reptiles and Amphibians ol 
Aiisitaliu"iA. >l. & A. W- Reed Sydney). (Fifth edition. 

1. 1 \Nisv.tiAM, Q M (!3751 Modified step poiminj; a 
rapid method of assessing vegetative cover, Soil flwti ' 
if, 256-2K. 

I-iimann. k. 09113) Tlif natuial history and cnri&ei\Jiion 
slalilH of the Adelaide pygmy bluetongue lizard Ttltyun 
nJi UtuU'mh Ht'ffH-i'ifbtitin 14, 61-76, 

(1992) * fciicyclopuedia ttl Australian Wddlite. 

Reptiles'" t Angus & ffcibertson' National Photographic Irak* 
ol AuNirati.m Wildlife. Sydney). 

I hi n<, n. R, J., M-vmrsoN. W, h. & Ci arm-., A. L. [J9681 
Solis and agriculture nf ihe northern and Yort.e Peninsula 
unions ijf South Australia. SpFiiAi Huld-nn Nth f,68, 
[>cptUtnuru i)f Itfr/cu/ttvr. South -iustnilii:, (QutfomtnEffl 
Prtnur, Adciatdc). 

OhltH. A. V- (I^S2) A new species oi Isiolnptwut 
(Laccrtilia: Scincidaei trom Wcsiein Australia, with HOtCi 
ou ilu biology and relationships ol other Australian -vtveic-. 
ffrr Au\t. Mit) J4, S4*> ?7^ 

HutCjflNfiOS, M. N, (J99£) "Mittalcncd reptiles in Soum 
Australia. Section 7 //i "lav S-K Ihd.) Thitaiened speeics 
and liabifuts in Sauth Auslralia. A cuuilysl for eunnnunity 
action." (South Australian Advisory Committee on 
Threatened Species Adelaide) 

& DuVNM l W S. C (1^92) Taxonomy and geneln 

variation in the Australian li/.ards oi the genus PsQU&tftiiiia 
iSeiueidae" Ly^osoininae) /, A'wr. Mtf, 26 21^-264 

Rimii*r*so,s. P. ^ Rmvi.ins^n. P A- (19891 

Tuxoruruie revision and eeologv of (be endemic PaMnaniun 
.selficid lizards Ltiuli'niyntti rrticrt'lcpuk'tittn and L 
[»trtt'>iUiti hip. hoc K. Sot. KtsHhwUi 12.1 257-274. 

Jn&Ofr, J. P tl9K*)) A lis! i>f 'he \ascular plants of Soutb 
Australia tcOiiion 31 / Atltfoidv Bin, Gimi. \1. \Ab\ 

Jbsst p. R \V. l'l l HSi A vt^ctaULtn iiild pasture survey o\ 
Omniics Pyre. rJurra and Kimbcrle\. South Austialirt 
front, ti. St*\ S, .hew. 72. ?S-B8. 

PirtKS, VS (IM(»M Eine IJhersietit der von Hrft- Richard 
Sehomtturgk an das /ooloyisehe Museum cin^csandten 
Amphibien. ,iuv liiiehslelde bei Adelaide in Siidaustralien 
Mh:r K h-t'uss JiittL HV'.s-. Bcfl W& 228-236. 

Sut'A. G. M. (1991)) The genera Ttliatut and C\i1otUmu>rphn> 
(Liieemlia: SetflcidaC).: generic uiagnnses and systematic 
relabonshtpv Mrm Q,i \ffa, IV^MS-Wi 

i l*W3 ) The mule rcpn tducOve welc <M the e.istcrn blue- 

tonpued liAtrd TtlttfHii sontxHdgs scincvidw (.Squamata 
SLineidaei. pp. 357^403. In l.unney, IX & Ayies, D. thds) 
"IJerpe'Mli-ey in Ausirjilwi: A t>Jvejse Diseiptine", 
ZihiIo^icjI Society of New South Wales with Surrey BcaUy 
& Sons. Sydney! 

& Mi.tchins-ov M. N 0^2) A new .species oi 
lizard {Titiquu) lioiti the Miocene of Riversleiirh 
gueen-slunil. Mem Q,J Mu?. 32, 303-310. 

SlMHiinvt-, M. P I19S5) Se.vuj) dimorphism ano 
TCpr^de>Ct|on r>t Ottfiproph-ifts $nu henofi (I-'jeerliljn- 
Setncidacl. pn IHfv hi Orjggi O t < Shine, R. & 
f.hinann. H tPds), "The Biology of Proys 
jnd Re-uilus". (Surri:y Beatly & Sons. Sydney! 

Stkono. D,_ Li MHiiRMAN, B. it BuATTsrrkjiM. R H (l*Wi 
Two new Simple methods for Catching small fast li/.ards 
Mfrft liVv 24. 22 2* 



Appendix I 

Plant dixersttx at Tiliqua adelaidensis StUdy site. Species noted during the general search but not recorded during step- pointing are indicated 
M (*t. Taxonomy modified from Jessop (WHty. 


Common Name 



NatiW .species 

Aihplex semibaceant 

Encitxtaena totnentosa ....... 

Maireana apbylla 

M. enchylaen<nde\ 

M. trichaptem 

Salsola kali .... - 


Ptilotus spafhulatns 


O.xalis perermans 


Erodium crinitum 


Euphorbia dmmmondii 


Crxptandra amara 


Pttnetett micrantha 


Aspenda cottferta . . 


Convolvulus erubesietiK 


Withlenbergia haeola 


Cioodenia pumatifida 


Leptorhxnchos ufuatnaitts 

Motunu lepiophvlla 

Vitiadinia euneata ........... 

V i<rat Hi* . . . . , 


Lomandra efjusn 

L. multipara . - 


Juncus bujonius 

J. krauss'H 


Arisiida behriana , 

Danihonm caespitosa , 

D pilosa 

D. racemosa 

Stipa blackii . . . , . . 

5. eretnophila (possihly .V. puberula) 

S, nodosa 

lliemeda triandra 


Hianex dianosus ...,,. 


Medicago Uttoralis . i . j 

M. minima . . 

Trifolium angusufolium 

/* arvrnse 


Ecniian plnntagweum . - - 

Neatosiema apulum 


Marrubium vulgare 

Salvia verbenaca 


Arctotbeca calentlula 

Canhamus lanaius 

Hypoehoeris glabra 

Sonchus oleraceus , . 


Gynandrirts setifolia 


Avena barbata . 

Brachxpodhan distachyan 

Bromus rubens . 

Hordeum glaucum 

U'tiam pvrenne 

V'ufpia nmralis 

berry saltbush 

ruby saltbush . . , 


wingless bluebush 

malice bluebush ...__.. 

roly-poly , 

pUNSVlJll , . , 

native .sorrel 

blue storks bill 

caustic weed 

long- flower cryptandra 

silky r tee-flower 

common woodruff 

Australian bindweed 

bluebell , . 

cut-leal goodenia ,..,..,....., 

scaly buttons 

minnie daisy 

New Holland daisy ... 
woolly New Holland daisy 

scented mat rush 

stiff mat-rush 

toad rush 

sea rush 

brush wire-grass 

white-top ... 

velvet wallaby grass , . . . 

wallaby grass , . 

eresteu spear grass 

desert spear grass 

spear grass. ................. 

kangaroo grass 

wiry dock 

strand medic ...__............ 

woolly bun medic 

narrow -leaf clover 

hares loot clover . . 

salvation Jane 

hairy sheepweed 


wild sage 

Cape weed . . . . . 

sallron thistle r , . - . , .... 

smooth caisear . ... 

common sow thistle 

thread iris . 

bearded oat 

lalsc brome 

red brome , 

northern barley grass , 

perennial ryegrass , 

rats-tail fescue , 













































- 1 






By Margaret K. Fuller* & Richard J. F. Jenkins* 


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. 


by Margaret K. Fuller* & Richard J. F. Jenkins* 


Fui.lhr, M. K. & Jenkins, R_ I F. (1994) M</orowipora 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 Liehenaria 
Winchell & Schuchert first appeared in the Early Ordovician. However, it is possible that Moonmipora chamberensis 
is an early representative of the tabulates, extending their time range down to the Lower Cambrian. The tabulate- 
like characteristics of Moorowipora cliamberensis 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 bowrnani Lafuste, and Tabulaconus kordae Handfield, but is unlike any other 
previously described Cambrian coralomorph. 

Kf.y Words: Moorowipora chamberensis. Early Cambrian, Moorowie Formation, tabulate coral. Flinders 
Ranges, South Australia. 


The Early Cambrian coral Moorowipora 
chamberensis gen. et sp. nov., is found in slumped 
reetal blocks within a megabreccia forming part of the 
Moorowie Formation in the eastern Flinders Ranges 
of South Australia. Moorowipora chamberensis occurs 
with Flindersipora bowrnani 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, Liehenaria 
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 (I962)| 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. 

1 Mount, T. J. (1970) Geology of the Mount Chambers 
Gorge region. B.Sc. Hons. thesis The University of 
Adelaide (unpubl). 

2 Hari, J. (1989) Lower Cambrian corals from 
Archaeocyathan-i/J/'f/jy/fw clasts within the Moorowie 
Formation megabreccia, northern Flinders Ranges, South 
Australia. B.Sc. Hons thesis. The University of Adelaide 

and siltslones, 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 1 ; Hart 1989 2 ; Lafuste et al. 
1991; Savarese et al. 1993). This stratigraphic level 
represents part o\ a suggested third transgress! ve/ 
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 et 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 10 m in maximum 
dimension. Within individual blocks, the organisms 
are commonly preserved in lite position. The dominant 
fauna! 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 tore-reef) probably 
protected the carbonate frameworks from vadose 

M K VVU.VH& R .». »- ll-NKINS 

diagctiesis. resulting m tt»c* remarkably pristine 
preservation of the fauna. The corals commonly form 
encrusting [g upright, nil colonies, the Ijuici' up lu 
00-70 cm high. Individual colonics (end to he widely 

Ftitidt'r\ifu>n! howmutii lias been found in biohcrms 
in the lower Oraparinna Shale al Ten Mile Creek (Fig 
I), as well as near the Moorowie Mine (Lafuste pi al. 
1991) At Ten Mile Creek, trtlobite and eehinoderm 
tiagmenis form hash beds in (he Oraparinna Shale, and 
associated aichaeocyalhs have been correlated wilh 
I aurml Assemblage 9 (Daily 1956). winch equates wilh 
the Pantn/ui ]tmi'iit> Zimc of Jc\\ (1990; Lafuste flal. 
1991). This slraligraphic correlation indicates dial the 
Moomwic f-oriuaiion is Botomiun in age 


Although complete reervstalli/ution of the colonics 
has CiCCUTrcd, an indication ol ihe primary micro 

structure ol [he skeleton can be seen in some patchy 
domains, lodges ol' the possible primary 
phuelcls arc dLsiinguished hy erenaie to wavy lines ol 
minute inclusions within the much coarser crystals 
comprising (he recryslallised fabric. Narrow, lath-like 
zones showing irregular extinction under polarised lighi 
occur at some places approximaU-Iv it:uisversc to 
skeletal elements where .secondary carbonate crystals 
penetrate the coral skeleton. This suggests a residual 
overprint of the original mineralogy disturbing Ihe 
optical continuity of the subsequent recrystallisation 
Apart from reerystalliscd .par. cavities surrounding 
00ralHtC$ and within ihe caliccs may be Tilled with very 
line sand or silt. It appears that during life, part or 
whole colonies may have been temporarily covered 
with a thin layer o( sediment causing the death of some 
/ooids The. survivors rejuvenated new parts ol the 
corallum, Some colonies seem to have been eroded 
by rapid energetic influxes ot coarse sand (also noted 
on enletmicrabe enerustaiions and arehacocyaths), 
allowing only a lew eoralhtes to continue their growili 
liactuics posi date growth and arc often filled with 
very tine sediment t Figs 2 and 3CJ. 

Fig l_ Location ituip showing fossil occurrence near ihe 
Mu<»rowic Mine and ilic distribution ol hariy ami Middle 
Cambrian outcrops in Ihe Hinders Rani^s ul Soinli 

Re, 2 [tilerpretive sketch (loneiiudimd section) nfhnloivpc 
SAM P3-II6^ illusinjtinjj cone in ran shaped colony: mi 
innii of individual corallilcs. prouvorullile (a); tabulae (hi 
septa (c); caliee fd): lateral im. tease [top centre unit right) 
and pcnpheinl iFitrae.uln.ular increase (\ 2). 

Fiy,, 1 U.njiiuidmal tectum-, ol hoMiypt- SAM P34J55; dluxiruiin^ parte ol' ihe colony (see Hg. J), A & B Top of ih»* 
i ttlnnj with vviilK extending atMivc [he Linitlliirn, rrieifmdy of increase and corallue birucuirc; C - base of colony U 4); 
.mil D Higher .iii^ m Ht_ niiou <i| <B) showing normal ami ihickcned tabulae (lower led); and oblique projections ol the 
outer wall (centre, right) (\ 15) » 






Fig. 4. Transverse sections and interpretive sketches ofholinype SAM P34I65: (A & B) teratoid lx 10) and (C & D) fasciculate 
(\ 10.5) areas of Ilie coralluni- Diflerenccs can he seen in corallile shape. Septa, and new walls across coraHites formed 
during ot'lset formation, Rejuvenation of corullites (C '&. O) is evident Lop nehl and bottom centre. 



The xerm 'platelet' (Lafustc ct iti. Wl). is used in 
describe relic mierosirucUital elements pF (he skeleton. 

Systematic palaeontology 


Class: AN 1 HO/OA 

Subclass: 7TABULATA 

Family: uncertain 
Genu* Mooronipora ecu. rtov. 
Type species: \ r foi>rt)\\ipttnt chumhetvusis so nov 

Erwuiiojiy: For the type locality near the Moorowie 
Mine in the eastern Flinders Ranees. of South Australia, 

OiiWttttsis, Corallum small, varying from massive 
eerioid to fasciculate, comprising polygonal, ova) or 
rounded corallites. Coralliles arc long, luberoid to 
irregularly cylindrical. Walls are thin, wavy to crenale, 
rarely strain*. Tabulae are irregularly spaced, mostly 
complete* concave upwards to undulating horizontal. 
Septa absent, or number up to 20 in each coiallile. 
Wliere present, sepia are randomly spaced, short and 
form wedge to spine-like projections inlo coralliles, 
arising from inward angulations of the wail. Pores 
appear to be absent, 

Moorowipora chamberensis sp. nov, 
FIGS 2 7 

U\mt>U*8\: For nearby Mt. Chambers. 
Oiatim^h; As for genus, 

Txpe specimens; The specimens described in this paper 
are held at the South Australian Museum (SAM), 
ilolotype SAM P34I65, lour thin sections. SAM 
P3416M; SAM P.U]b5-2; Paratypes SAM 34166-1: 
SAM P 34166-2, 

Material: The hotutype, paratypes as well as several 
other colonies come from one rock sample measuring 
270 mm long, 230 mm wide and 120 mm taken 
from a large boulder v/nhin the Moorowie Formation, 
neat the Moorowie Mine in ihe eastern Fltudcis 

Descriptifm: In transverse section (Figs 4. Mt eoratlttes 
show gradation into two distinct habits, massive eerioid 
and fasciculate dendroid. In fasciculate habit, coralliles 
rarely touch, are circular lo slightly oval in shape and 
vary in diameter front 0.95 to 3.75 mm. Coralliles with 
massive habit are rather irregularly shaped 5 or 6 sided 
polygons, sometimes oval or rectangular, rarely 
circular; Ihcy vary individually in diameter rffdjp 0.77 
to 13 mm. 

Walls arc thin, varying between 0.1 mm and 0.15 mm 
tn thickness and show a relic fibrous structure (Figs 

A midline is apparent between many adjoining 
coralliles. Walls are wavy to slightly crenate. being 
rounded in isolation or adjacent to small spaces in the 
corallum. with u tendency to become suaighter and 
less distinct where Ihcy merge wiih the walls of 
adjoining coralliles. 

In longitudinal section {Figs 2. 3). each colony is 
generally small, numbering from 2 or 3 to about \h 
coralliles. Colonies are either cone- to fan shaped, 
diverging outward from a single protocorallite. or more 
rectangular where Ihey appear to arise from several 
adjacent corallites. Increase is both lateral and 
peripheral intracalicular producing I, 2 or moic offsets 
l Figs 2, 3A). 

Individual coralliles are tuberoid to irregularly 
cylindrical in shape and vary greatly in width and 
length up to 19.5 mini, prior to increase (formation 
of a new eorallile) The base is rounded and blunt, the 
protocorallite producing 3 or 4 short septa prior to ihe 
formation of an initial tabula. Thecal ice is prominent 
extending between 2.5 and 4.75 mm past the last tabula 
I Figs 3D. 3D). 

Tabulae arc mostly complete, mainly convex-upward 
Cn undulating horizontal, often down- turned where thev 
meet the wall <Figs2. 3), They are irregularly spaced, 
but commonly occur at the same level in adjaoejl/l 
coralliles. The distance between them is highly variable 
(0.35 to 2.1 mm), while ihe thickness of tabulae varies 
from 0.002 mm (o 1X01 mm. 

Septa number up to 20 in fasciculate corallites: arc 
very short (0.01-0.2 mm), generally equal in length, 
triangular to wedge-shaped, often indistinct, They form 
protrusions or ihe wail of the corallite ai sites of inward 
creasing (Figs 4, 5). In massive, eerioid corallites in 
the main body oi the corallum, there may be up to 10 
septa or septa may be absent Septa are randomly 
spaced; long and short septa may alternate, or only 
long or short septa may be present Sepia arc wedge 
to spine-like in shape. Sepia are generally longer than 
in the fasciculate corallites. In longitudinal section septa 
are observed as continuous vertical laminar f4ftt£S 
mterseciim; normally with tahulac. 

Micraswuiurc; The miscmstructure was studied A 
magnifications up to x 200, and photographs wen. 
taken under polarised light. 

In transverse section at low magnification (Figs 4, 
5i relic fibrous elements which ibrm the selerenchymc 
and apparently represent indications of original 
biocryMals appear as continuous Itneae across the wall 
and into the septa At higher magnification (Fig. b) 
the interlocking fibrous elements form triangular to 
rectangular bundles, composed pf narrow parallel sided 



and blade IBM geniculate structures up to 164 flin bug 
and 37 /mi wide These are tingled towards and awa\ 
from thi' centre ol each corallum. The bundles have 
ihfl appearance df crossing, or being slacked over 
Mi»ilrrl\ iujj layers. Near the midline ot walls the fibres 
art often less oblique and have a slightly different 
orientation, appearing to be broader and more 
randomlv oriented 

In longitudinal section at magnifications of > 100 
tO H J(K), the midplane ol the wall seems to have been 
tomposed of irregularly shaped, eremite, interlocking 
platelets Which individually represent (he librous hneae 
of transverse i. tits. Platelets may fx: almost rectangular. 
vertical lu slightly inclined, occasionally almost 
horizontal in the middle of the wait. They commonly 
occur diverging outward towards the top of eorallites 
(Fig. /), Wall platelets are more elongate and wider 
than the fibre bundles seen hi transverse section,, being 
up tu I'M) /t ni in length and I3K /mi in width. .in- LuiiiinuoiiN with the inner edge of the 
wall, which converges slightly around them. The 
•arm miv ot'ihe tabulae is similar to the sepia and wall 

in transverse section, with bundles of fibres being 
mainly triangular, or irregularly shaped Triangular 
bundles of fibres are up to 360 fort in length, and 
308 /mi in width. In tabulae of normal thickness, 
adjoining triangular bundles interlock forming a 

Rltj } Imiixverxc section ol holoivpe SAM PJ4lfc$; purl oi" 

t.orailom showing both cericnti and fasciculate ureas \ \ \ \ i. 

I lu (>. Tiansvcrso section and interpretive sketch of holoiypc 
SAM P34165i illustrating injiigularaiid icetaiitfulai bundles 
of fihrcs extendinu utnm ihe vvalls ol the coral lite •. \\ 
and y-yl define boundaries ol illusrrauuii, in is iimlliiir 
ul ttie wall (\ 44). 


upper and lower surface. A more complicated 
mlerloeking pattern ts formed in thickened tabulae, 

Kt'tmtrh: Mooro^tttotv chatufn'remis is dimorphic m 
hoth the morphology ol the eolony and the method ol' 
iueicase. Dimorphism often occurs in tabulate-; and 
may be shown as differences in corallite si/i\ shape 
and internal structures (Oliver l%8. 1**75) Many 
iriira.speeine variations arc probably environmentally 
controlled, being inriuenced by taetors including thfi 
adiaeeni sediment, and the position ol corallites wilfuri 
ii colony (Oliver IWitt) The two differem 
morphological fornix observed in M. citmnbernisi.\ 
probably reflect (lie mode ol increase, which also 
appears to be dimorphic, being related to the position 
of individual corallites within Ihecorallum. Corallites 
within the fasciculate area, which generally occurs at 
the outei edges of the colony, appear lo have resulted 
Irom lateral increase. Offsets branch away from the 
main colony producing isolaled t-orallites which rarely 
touclt and are therefore unnaffecied by crowding (Figs 
2, 5) Such corallites are thus round IfeJ slightly oval 




V..— - 

hijj 7 Longitudinal section i cartoon sketch) showing large 
instills n( uV nvrystalliNJimn fabric fa) incorporating the 
WUll <>( WDlpou/lJ Cor:jlltfes_ <bf Relic biot'iy-slal dhivs 
diverge towards the coralluni, 

in transverse section. IVnphcrat inirncahcular incTease 
is mnsi common in the massive, cerioid pans, and 
where a solitary corallite htW become established ifv^- 
2, }), Usually one, I wo or more offset* ate produced 
at the same time, with new walls growing from sties 
6f septal insertion across the culice. Both methods of 
increase commonly cum a' djffwrtf stage .■» ol growtb 
within the same corallde. and are probably lelated U} 
the amount of space surrounding it, 

The variable distance between tabulae and Eh? 
picsence or absence of septa not appe.ii lo be linked 
to any particular stage of gioswh a characteristic which 
ttUS been suggested as possibly occurring in some (Hill iVKI). Sepia prim. inly occur in the 
protoeorallite and immature corallites. while at othei 
stages of growth they may or may not be presenl. 

Discussion and Conclusion 

When compared with other Cambrian Gtftalf 
suggested to have labulalc affinities (Sainton W). 
Moftov iptttii rhattibcrcusis is closest in th 
morpho'ogy tori iahnkii'imtis kanlae Hand Held IMrWJ. 
from the Early Cambnan iBotomian) of east central 
Alaska and British Columbia, fn vertical seelion, M 
\hitinbcn-nsis and T h>nhtt differ in the shape ul the 
corallites. which ait more tubular in appearance in the 
l< >rmer Height and width vary, with mature corallites 
being up to 19.5 mm long and \U mm Wide m M 
ihamberensis. while corallites of T, kort/tw ;ia' up lo 
65 mm long and 27 mm m width m the colonial Ibrm 
(Uebrenne et ai- (W7), The tabulae also differ, being 
undulating horizontal to concave upward in M 
\ haaibervash and either jioit/ontul or slightly concav* 
downward in 7_ kontae. Incomplete tabulae are rnoic 
dissepiment like and walls are generally thicker in the 
latter (Handfield IOfW>)_ The microsirueturc of both 
corals i,*. significantly dillerent in iiansverse y$fttinn, 
being geniculate fibres in M, iltumbintiM*. and 
concentric, light and dark wavy lamina! tons in 1 
kt/hUw Tabulae also differ, being formed from bundles 
of fibres extending upward and downward from a 
medial line in the former, uniike the two layeied light 
and dark /ones of T, kenluv Howevei. plaieletv 
• longitudinal section* in the walls ol M- rhatnbcrcnM- 
arc of similar shape, hut generally larger M, 
chantbt'rcn,\t,\ may belong in the family Tabukicomdae. 
but. the above differences, likely preclude it from llus 

A'/, (huwherctnis is distinguished irom Lip<>fh>ni 
km and L Junta Jell & Jell (976, bom 'in- K.rK 
Mutdle C ambnan of western New .South Wales, by lh<- 
piesence ol tabulae and the shape and atran^enienl ol 
septa, although rhe coiallitesof A. Hwa arc ol' similar 
length and width. Cumbrotnpa intituaiwn\i\ I'm/ L V 
Howell 1955. from the Middle Cambrian of finish 


M K M LLER & R. J. f. JilNMNS 

Columbia, is distinguished from M rhtimhttensis tijf 
the niorc slender cor<illiics and the absence (A iahulat 
and sepia. Most ol the Early Cumbrian I "r.iitiniorphs 
described by Korde (1963. 1984a. b. i98<). WOi. art 
parilv synonyms of already deseribed kh.i-„<k>iiJs CM 
Indioeonu/oans and partly nonnna tJubnt or nufln 
(/huravlev W oi 1993, p. 369). 

A'/, clutmberi'ttsis- also differs Irom Fhtuirrsipnru 
htiwnumi Lafuste 1991. although there are some 
similarities in mierostrtkture. M < lutrnberettsis is 
v.noid in fasciculate and lias wedge lo spine -shaped 
sepia up to O.Z mm in lenclh arising Irom continuous 
wills 0.1-0.15 mm thick Iti contrast F. bownumi is 
meaiidmid to cer ioid, and has 6-16 strongly developed, 
straight lo si ightly curved septa up to 0.8 mm in length. 
with rhe eddies of septa bOAffiflg very short blunt -.pines. 
WalK form very short segments between the sepia and 
are 0.13-0.25 mm in thickness (Lafuste et al. N9I) 
In /*.' tfitwmuiti tabulae are mostly concave-downward 
and closely spaced 1 0.2-0.3 mm), but are undulating 
hotvonlal to concave and more regularly spaced in M 
iiuimhrtensis- The mode pf increase is by longitudinal 
liNMnu in die former while both lateral and intacalieular 
peripheral increase occurs in the latter. 

In transverse section, the mierostrueture of bodi 
corals is sumlai, with walls consisting of geniculate 
fibres which diverge and converge in two direcinms. 
In vertical section platelets in M. cfiambermsis are less 
elongate and broader when compared with E bimnumt. 

\\ is considered that Ihe genus Ucht'ttoha which 
has a time range Irom the basal to the early/late 
Ordovjcian is the most ancient tabulate coral. Us 
colonial form is ecrioid, it has a simple morphology 

and tabulae, is ascptate. and may have rare mural pores 
(Bassler 1950: Flower 1961. MeLeod 1979; Scruilou 
1984. Latin 19841. M. chamberensiv has structural 
characteristics which demonstrate us affinity lo lite 
tabulates (including septa, which are not present in 
Iiehenands) These are (!) the eerioid lo fasciculate 
form of ihe polony; (2) the spine like to wedge-shaped 
septa; (3) its mode of increase; (4) the generally 
complete well-formed tabulae. The obseived relic 
ink mstructure appears to be similar lo Ihe pmnatelv 
(clinogonallv) librous structure of some tabulates (see 
Hill 1981. p. Pt52). including Iiehenands, though the 
extent of the modifying influence of diagenesis K 


M. cluimbemms with its tabulate like characicnstics 
may he eithet an early representative of the tabulates. 
or belong to u new group of corals with convergent 
evolution contributing to their similarities. These 
alternatives have been suggested by Lalusle ei al. (199! "i 
lor A bowmaniy whereas Scruiton (1992). regards the 
latter possibility as most likely The addition of hi. 
cfiainbcrcnsis lo the group of known early 
coralomorphs, provides further evidence that tabulate 
enrals may have their origin in the Raily Cambrian. 

Viknou lodgments 

The authors gratefully acknowledge Brent Bowman 
and Jt>hfl Hart who collected the studied material. Dr 
David Gravestoek is thanked for additional information 
and his constructive criticism ol a draft of this 
manuscript- We would also like t0 thank Mt OitW 
S'nidero for his photographic work 


tfv.suK, R S. U^50> Fauna! lists and dewciiptwn ot 
Palaeozoic Cural.v Cn'ot, Sen; 4m. Mt-tu, 44. Htf 

I>aiii. H (19561 The Cambrian in South Auwillla 20th 
Jno-rn Gtot Con#n'x\ Mexico 2, 91 147. 

(Im <,\Kr*«t, C R iW64t Lower Cambrian stratigraphy of the 
Hinders Ranges, thins. H, Sue S, fim KK. W 144. 

& Johns* »v J t U%2) Cumbrian sequence ol" the 

western rhmJets Ranges, Quart swC twtex, Ceot Sm 
S Aum 4 _* S 

f)iKwtN-ui-. T M, GANi.t.Mf) . «, A. & 1.\ksii . J, G 
(I'TS/j hhuim otitis Hand field Miemslruciure and ilv 
implication at the (tuononiv of pnmilive cnra|>. four. Put 
fil. I ■'» 

I-I.OWIH. K, \\ (Wfijj Khmtoya and rckiled colonud enrols 
Ntn> <Ur< ht\ Uih Mem, 7. |-97, 

I-kmv, M, A, A H-iwiii 8. E (1955) An Ipper Camhrmn 
c--rj| iri-iii Vtdiitana Jour. Pai. 29. 181 183. 

I ■!>■ ■.', i-M<M k, D. I, & Uibbiikt. ). r- |©915 Jjequeocc 
antigraphy of 1hc EiRlferq OftKei and Arrows- Biisins: 
J li.iim-wtnk rot t'iMiil'run oil seulrh- Al y t 4 Jour 31, 


II ssonhiu, R. C. \VH$\ Ear(> Camhriun com Mike fosijls 

IVom the northern Cordillera of western Caxbdu. Cat) 

Jour, hank .SV 6. 7H2-7HS 
HlU D (NSt) Ruposaand PahUlato. pp. 1-762. fnTcicUcri, 

C ifivl ) "Treatise nf invertebrate palaeontology Pj<i K 

CuclCQttfmtil. Supplement t ' fGeoI. Soc. Am. A Utu 

Kanhab, Buuld^r. CVilormJi*.) 
J,\Mt.s, K. V. A GKAVl.STfJCK, D. I- (ly^l)j Luwer Carnhrun 

shell and shell niavyin buddups, f-'lnidcrs Riineey. South 

Australia. SeflitvenU'fogv 37, 455-480. 
Ifi i f A. {\99Qft Bjostmligr-Jphy. t/t liengWon, S , ComvaV 

Morris. S. Cooper. H. I. Jell, P A A: Runnegar. B N 

ihdS) Earl) Caiiihrtan fOssits from South Au'stmli;*- is&?C' 

AusH/i, Pal Mem. 9 t J4-W. 
(Si Jr.i i. J. S < I97oi t'.ailv Middle Camtmart eorals 

from western New South Wales. AUtunnga I. t81-l u 5 
K'H-Di- K. B. lj£5?) Problematic fossils Irom the Carnbnait 

deptiMTS ot the south-eastern Siberian Ptattiimi- Tnmsl , 

HoU A^iti: Sm. OSSR^ Fjolh Sa. Sat. 125, 358-360 
! *M Mt-. J . Di-ufdiNNFi. K. Cjanoin. A- A: GKAVbS'iueK, D 

(1991) The oldest Uihutate eoral and the associated 

ArehJeoeyatha. hwvr Cambrian, Flinders Ranges, -Stiutli 

Air^ulla (, 24. 697-7IK. 



Lawb, R. S. (1984) Lichenaria Winchell & Schuchert 1895? 
Lamoiiia Raymond 1924, and the early history of the 
tabulate corals. IV International symposium on fossil 
Cnidana. Palvonlographica Americana 54, 159-163. 

McLeud, J. D. (1979) A Lower Ordovician (Canadian) 
liehenarid coral from the Ozark uplift area. Jour. Pal 53, 

Oliver, W. A. (Jr.) (1968) Some aspects of colony 
development in corals. Ibid. 2, Suppl. 2. 16-34. 

( 1975) Dimorphism in two genera of Devonian tabulate 

corals. U.S. GcoL Sun: Prof. Pap. 743-D, 1-11. 

Savaresr, M., Mount, J. F, Sorauf, J. F., & Bucklin, L. 
(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 
groups." {Academic Press, London & New York). 

(1984) Origin and early evolution of tabulate corals. 

IV International symposium on fossil Cnidaria. 
Paleoniographica Amer. 54. 110-118. 

(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. 

graptolites and corals from the Lower Silurian of 

Minnesota: Geol. Minn. 3. pt. I, p. 83. 
Zhuravlev, A. Y.. Debrenne. F. & Lafuste, J. (1993) 

Early Cambrian microstructural diversification of Cnidaria. 

Cour. Forsch. Inst. Senckenberg 164, 365-372. 




ByM. Peterson*, G. M. SHEAf, G. R. Johnston^ & B. Miller§ 


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. 

Tninxuth t >tnofth<>RoyatSoae<vnfS, Ausi (1994), I1H(4K 2>7 .>44. 


by M. Peterson* G. M. SheaI. G. R. Johnston^: & B. MiLLbR§ 


PhrbitsoN, M-. Sht*. G. M., Jowssion, O. R. 'V Mu u.r. B. iW94) Notes on the morphology and biology ol 
Ctenophnrttt >nckat:.iei (Slorr, lOKI) (Squamata: Agamidae). Tranx. fi Sm\ S, )usi IIHMl 237-244, November, 

The systematic status of Ctetiuphonis tiKkcuntfl is reviewed, on (be hasis ol "additional muleiial Irom Western 
Australia and Soulh Australia- Habitai duiu ft* kiiown localities, stomach contents, one record of clinch sjwi, 
and behavioural observations and thermal preferences of eapiivc individuals are provided. C, nukt'iwt is 
morphologjcallv very similar to C\ Mtuulaiit.-. differing mainly in adult si/.e and subtle niodificuttori-s ot colour 
pattern, C mckftirtei and C scutula/HS are pbeneiieully most similar to C. trixuint.s. 

Kiv Wohds: Squamata, Agarmdac. C/ctit.pht>/ti.\, morphology, biology, habitat, dicl. thermal biology. 


The agamid Amphibolurns ntckemici was described 
(rum two specimens (Western Australian Museum 
|WAMI R59753-54), collected at 5 km and 8 km SW 
of Ponier Rock, Western Australiii (Slorr 1981). The 
species was subsequently transferred, with -a number 
of 'other agamid species, to Ctenophontx (Storr 1982). 
Additional specimens of C. mckenzivi have since been 
reported from Western Australia and South Australia 
by Sehwaner& Tyler (1985). MeKen/iert a/. (1987). 
Anon, (1988), Wilson & Knowles (1988). Armstrong 
(1992) and Ehrmann (1992). This additional material. 
previously uneharaelerised. considerably extends our 
knowlcdgc of the morphology, distribution and habitat 
preferences for this species and provided (he basis for 
this papei. 

Materials and Methods 

distance) NE'-Colona", 3I°31'S 132°09'E. 13.xi.l9K4: 
SAM R32264-65. R32268-69. 0.5 km S Mitcherie 
Rockhole, Yumburra Conservalion Park. 3I°27'S 
132°49'E, 17.x. 1987; SAM R32266-67, 5 km S 
Mitcherie Rockhole, Vuntburra Conservation Park. 
31°29'S I32°50'E. 18.x. 1987 (Fig. I). 

Measurements taken on preserved material follow 
those used by Shea &. Peterson (1985). Log-transformed 
morphometry data were statistically analysed with 
analysis of covariance. using the SYS TAT statistical 

package (Wilkinson 1987). 

Dietary data were obtained by examining stomach 
contents of six specimens: SAM R2832. R25572. 
R2nlu0-02. WAM RJ59753. 

Four specimens were maintained alive for some time 
following collection. The lizards were housed in a 
vivarium with a heat lamp and fluorescent lighting, 
both turned on at 0630 h and off at 1930 h. 

Eivc specimens from Western Australia, additional 
to the types, have been collected: South Australian 
Museum (SAM) R2832, Western Australia; WAM 
R9I764-65, 9 km E Kilidwcnma Granite Rock. 
32°04'S 124°00'K. 29.iii. 1984: WAM R91842, R9I852, 
15 km E Kilidwerinia Granite Rock. 32°03'S 
124°05'E. 5.x. 1984. Eleven specimens have been 
collected from South Australia: SAM R25572. R20I62. 
12.6 km (air distance) NE"Colona". 3I°31'S I32WE: 
SAM R26160M, 16.2 km (air distance) NE 'Colona'; 
31°3I'S I32°II'E. SAM R268W, 15.2 km (road 

+ o9 Alvah Street. Si James, Western Australia 6102. 
t Department ol" Veterinary Anuiomv, University of Sydney, 

New Soulh Wales 2006." 
$ School of Biological Sciences, Flinders University, Bcdthrd 

Park, South Australia 5042. 
^ Department ol* Animal Services. Adelaide University, Soulh 

Australia $005, 

'J i 

,"...... ■ ! ■ ■ .. ■ ', i ... t. | ...'.,"";....., 


*-■•'.■ . •' J 1 ', 

1 IflUi *." .■■' ■ ■ ' -■■■ . . 

' I 




,,,.. . J i . ., .. .,- ■ . ..... ,, ....,,..,-,_ .„-(. ,, . ,| . ...,.' 

Fig. U Distribution Of C. mi.kcurjf't. 



corresponding lit the local natural phOWjStWti at lhat 
lime Water, mealworms and chopped fruit were 
available ail libitum However, one individual ( fe-l226ff) 
became emaciated and was euthunased \\ 1800 h oil 
ll.\iU9tf7 (i.c , 54-55 d post capture) the remaining 
three li/ards were placed in a thermal gradient 
(X-60°C>. and allowed to acclimate. Calibrated copper 
Consmntan thermocouples were placed at least 20 mm 
into the cloaca and held in position wi»h surgical tape 
Cloaca I temperature* were electronically recorded 
every 10 niin. for each animal from 1130 h on 
l2.\iU9H7to 1130 hpn I3.xh.|987, AnytemperatureA 
taken while li/ards were tangled in the thermocouples 
were excluded from the analysts. 

Thermal data were analysed using a one-way 
ANOVA to test Tor differences in thermal preferences 
between individuals. In comparing thermal preferences 
during scotophase and photophusc. data from all 
individuals weie pooled. 



The South Australian sample (snout-vent length 
|SVI_| 41 76 mm, x - 615 rnm, n = \\f\$ similar in 
morphology and colouration to the Western Australian 
sample (SVL 31.5-71,5 mm. x - 5X.4 mm, n = 7) 
Tliereare minor, but snuisticaHy significant diflerenccs 
heiween the two (Table I) in the relationship between 
tail length (TL) and snout venr length t slopes 
r ,- = 1477, p < 0.05; overall. IL'SVL 
2!5,0-252.9#, x - 233.5'.$. n - II vs 206.3-236.9% 
x - 222-5%. n = 6 respectively) arid between head 
depth iHD) and head length (HL) (slopes: 
F J|W - 9.900, p <_ 0.01: overalirHD/HL 577-67.9':; 
v - 61.4%, n - 11 v.v 62.0-67.5^. x - 64.0 f v , 
n — 7 respectively), with the eastern juveniles having 
a longer tail and the eastern adults a more depressed 
head It is possible ihai the apparent long tail on eastern 
juveniles may he an artefact due to the lack of very 
small juveniles in the sample. 

The affinities of C tnrkncJvi have Wot previously 
been adequately defined. Slorr (1981) assumed a close 
affinity with C sauulatus (Stirling & /eiu. 1893). 
bul provided no argument for this. The C m<knt:U-i 
material now at hand clearly indicates that this species 
is very close to C scuta lotus \ Head, HtNvy and limb 
proportions are similar, I he arrangement Of spine 
clusters, rows of enlarged sealesand skin folds on the 
head, neek and body is identical, and most elements 
of (he colour pattern are shared. Both specie* the 
following colouration elements (Fig. 1\ 

1. a pale vertebral stripe, bordered bv 

2. a dark- paravertehml stripe, bordered tatctally h 

1 a longitudinal scries of pale blotches, coalescing 
anteriorly and often posteriorly to form an irreguhi 
stripe from hchind eye. ovei the ear ihm\ upper 
posiaiirieuiar spines, to rhe tail base, where (he two 
stripes ollen fuse_ This clement is hoidered laterally 

4 an upper lateral /one from below ihe eye, through 
the ear. over the forelirnh and hindbmh to the 
proximal part of the tail, hordered vent rally by 

5. a broad pale midlateral stripe from axilla to groin, 
then along hind edge of thigh along ihe proximal 
pail ol the tail, dark margined ventrally; 

6. on nape and fotebody, a series of dark transversa 
bars connecting rhe dark paravertebral stripes and 
interrupting ihe pale vertebral stripe:,; 

7. narrow pale caudal rings usually present, overlying 
other elements ol t.^il pa'tern; 

8 dark ventral markings in males and some females 

Consisting o| at least a broad median stript from 

chin to eular fold, bmadesi iu>t rostial to the gular 

(old. and a rbomboidal to kite-shaped dark breast 

patch, extending eaudully a variable distance, ut 

seme cases t«» the groin Some specimens of both 

species have additional dark ventral markings 

including streaks and spots laterally tin the throat. 

and dark anterior margins tii the thighs (Fig. 3: 

StotT (1981) differentiated C. imkt'n:ivi from C 

\iutulaius by four characters; smaller size, deeper 

bead, fewer subdigital lamellae on the fourth toe and 

colouration (back blackish brown with whitish 

vertebral sinpe and transverse lines is pale brawn with 

paired dark brown blotches merging tm forebuck 

to form erossbunds). The additional niarerial 

now available supports' the si/e diffeieuce 

(&VL - 31.5-76.5 mm vv 31-115 mm; Sum n aL I983J 

The smallest male C ttnkcuzia wtrii diluted, wax-filled 

femoral and preanal pores and turgid testes has 

SVL - 48 mm (SAM R25572). while a female of llle 

same size (SAM K26890) has well-developed ovaries 

and oviducls. One female (SAM R2K32. 

SVL = 71.5 rnm) is gravid, with three ovtducal eggs 

However, the remaining three characters employed are 

non-diaenovrk or require qualification 

The purportedly deeper head o( C m<kctuii'> is noi 
apparent in Qui measurements (fable I), although it 
ts clear Irom the very much lower values given by Srorr 
(I98li that he used different measurement;-, u, obtain 
a he.v.) depth/head length ratio (possibly head deplh 
as defined k Witteu 1 19*51), We have measured head 
depth at ihe highest poini of the skull, over the parietal 
eye, and head length from tip of snout to rostral margin 
ol ear As noted above, ihe eastern sample of adult C 
nukat:Jet had a shallower head than the western 
sample A difference <a .-.miliar magnitude was also 
present between the eastern sample and f. \ritutlatu\ 
(slopes. L, ,, - &$4, p <• 0.(H) 



*^ T "' "~ : ' 

....— -..■■■■ .■—-.•- ^^„,.>., > 

«ts*" : 

Fig. 2. Dorsolateral views of A. male C mckenziei (15.2 km NE "Colona" SA) and B. male C. scutuhuus (Hospital Rocks, 
WA) in lite (photographs M. Peterson). 



The lower number of subdigilal lamellae in C 
fruken:iti is not an absolute difference, and is 
geographically variable. Western C mcki'ncjei had 
counts of 25-30 (x - 27.4, mode - 27, n - 14) while 
eastern C nu'kctizU'i had 25-32 i x - 29.6, mode =s 30, 
n = 18) Thfi C svuudatus sample we examined had 
28 41 (x = 34.9, n - 80) snbdigital lamellae, only 
7,55? ffiWer plan 32. although Storr et al. (1983) give 
a range of 31-44. All means are significantly different 
(pairwise t-tesls. eastern r.v western C. mtkonzivi , 
i M) — 3.36, p < 0.05; western C. mckt'nrjci VS C. 
.\i-utttititus, u t2 - 10.88, p < 0.005. eastern C 
trukenztei v.s C. scululalus, U >0 ^ 8.33, p < 0,005). 

As noted above, the colour pattern is composed of 
the same elements in each species, The difference noted 
by Storr (1981) is due to paling ftf the dark dorsal 
markings in C. .scuttiUitus^ particularly the centres "1 
such markings, and enlargement oi' the pale dorsal 
spots I lie dark crossbands on the forebaek of C 
scHtulatHs are also present in C, mckenziei, though 
narrower, less contrasting and often fewer. 

Sympalry is not yet known between C mckt'trzift 
and C sanulatits to confirm a species-level difference, 

although the two species arc known to approach to 
within 85 km of each other (WAM R9l7o4-65 v.s 
R65538, 0.5 km S Buningonia Spring, respectively) 
However, the lower number of subdigital lamellae in 
western C Jtickr/rJci, which arc geographically closest 
to C scvtulofu.s, together wilh the much smaller adult 
size, and consistent evenly dark upper lateral zone ( fj 
usually light centred) suggest that C mckenzici is 
specifically distinct. 


The two Western Australian localities listed above, 
and the two South Australian localities I2.ft km Nfc 
and lfi.2 km NE *Col»>na" are respectively sites BA2B. 
BA5, K02 and K04 of McKen/.ie & Robinson f 1987.1. 
while ihc localities 0.5 km S and 5 km S Mucherie 
Rockhole are sites MI5I and MF2J of Copley & Kemper 
(1992). From the combination of photographs of the 
habitat and floristie lists at each locality provided by 
MeKcnzic & Robinson (1987) anil Copley & Kemper 
(1992). specimen collection data from both museums 
and observations by the senior author, it appears that 
the primary floristie components o\ the C. jnckeitzjei 

Fig, V typical uu'lanin ventral patterns ut A C mckai:ici and B, C. scuuihiius Scale bars - 2 eili 



habitats are h'ucahptus trfposii f'v.M ex Miq. (giant 
malice). Acacia tisntildii K.v.M. (umbrella wattle) and 
A. papxntcarpa Benth. (western myall) over a 
chenopod understorey ol \ Atriplcx vesicaria Heward ex 
Benth. (saltbushK Mairvam spp. (bluebush). Rlw&tdia 
sptnexcens R.Br, and Sclcmhtena spp. The ehenopod 
understorey is common 10 all localities. The substrate 
type at sites BA2B, BA5, K02, K04 and ML5I 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 
fO the substrate recorded from site MI2I. 

At sites BA2B, BA5, KG2. K04 and 15.2 km NF. 
"Cnlona". C. mcketiziei was sympatric with C. pains 
(Peters 1866). and at the former three sites it was also 
.sympatric wilh ihc agamid Tympattocryptis lineata 
Peters 1864. At site MI2I, it was the only agamid 
recorded, while at site M151, C. fordi (Storr 1965) and 

Pogona minor (Slernlekl 1919) were also recorded. The 
latter species was also found at 15.2 km Nf'Colona". 


I he dominant prey items in all specimens examined 
were ants, particularly small Iridomymtc.x. although 
some larger ants (Gtntpanotus, Mehtphorns and other 
unidentified genera) were present. Other prey items 
recorded were Hcmiptcm (single individuals ol 
Povcihmteris sp. in R22835, R2832. two pentatomids 
m R2832, and a small unidentified bug in R26I62), 
Coleoptera (one small cuiculumid prothorax. and a 
large scarab larva in R2K32; three unidentified elytra 
in R25572), Meeoptera (a large abdomen in R2832), 
Diptera (one small lly wing in R2832), Homoplera 
(one small wing in R25572). Orthopiera (one small 
grasshopper head in R25572), Hymenoptera (one small 
bee head in R25572) and flower parts (R26161). 

T\UI v 1_ Allometiii t-tftuilions and calm luted vtthtvs Jitr cratdul taut somui'ti pmportiort.s m C. nicken/iei {mcki, C. xcululutus 
{sCU) and C. crislatus icri), Eastern unit western samples oj ' C. mcken/iei are kept separate for tail length and head depth, 
but pooled elsewhere, us the equations were not significant I \ different, Values u and h M/h'e the equation y = /«". s.e, ts 
Mandurd error nf u, r - lorrchttinn n /efficient, uL - docttion iff significant ailonwlry, C' r , C 7f} and C} e, arc calculated 
percentage proportions at SVL - 32. 76 and 105 mm {approximate minimum sire for all spei ies. maximum sire ofC mcken/iei 
and approximate large udult size for hath C. scutuUtlus and C cnsLtULs) 








( --m 

































46 17 





















1 1582 


1 104 




2 14 
















































40 S3 


































1 .234 









I 206 

942 8 






























.93 1 2 


















Two lypes Crt circumduction and iwu lypes <»f "head 
hob were seen. The circumduction types s«cm |H 
correspond 10 the challenge wave and submissive wave 
tk^nlxJ by BraUstrom (K>7I) in Po^omi hrtrhtirn 
(C uvicr |JS2 L ')- The bead -bobs differed in cadence and 
degav m which the, head was moved aided by 
extension of die Inrebmhs. In u dominant male the head 
was turned a greater distance below and above die 

norntbl ptano^o! - bstci rase ihun ii wag in a subordinate 

male "i |wi) lemulc**- 

Two eaptive mules were observed in an agonistic 
display I poll being plaeed m ihe enclosure lor 
acclimation, they head-bobbed and circumducted 
several bines be to re presenting !■» each Otto •'' a 
distance ol about IS em, They were lacing in Ihe siiiut 
diivaion and continued to bead- bob As the display 
intensified they coiled iheir Lails loosely and both tlid 
SCVCTai hind-leg push-ups, similar \o diusc described 
lor [ftc ( . tli-< n.sii complex by Gibbons U'W). The 
display ended Whcil one lizard biithe other on Ihe nape- 
II uy rolled violent ly ;ibt>ut (he cage and then 

■„ pnratftd 

tymp) ralitrc preferences 

The three C tmkt'tt:tei sludied maintained body 
1. ...permutes between ll.l°C and 43.4°C i \ - 34,7 C. 
\M\ ill a thermal gradient over a 24 h period 
• l.ibh \\ t iK-ic were btgnifiCflflt differences in thcoiul 
pieii-n-nces between individual lizards (p = 0.04). This 
nifty love been due n» the low vaj iflnsc pf R322W with 
respect to the other two individuals Differences 
between Itightl U and daytime hody temperatures 
uyerc barely significantly dilicTcut fp - 0.05). 
However there was a large difference in volunlary 
minimum body temperature daring photopruise(Hl n O 
und scoiophase t2K.7°C). This may relied greater 

J MH I 1. t ompttMt&n oj ffwrvtttl /»*■/<'• u.v. iinuttiy, tht<\ 
i i. m i /ii'' nti, in .i iluinuil ihiJi'tit Mi • i\un-irnnl\ ,m 

I i/ard 







3? 3 

34. ,' 

25 .: \<t g 
M4 II t JWJ 

4.05 n 0-43 4 

3 M 

II 1-43 4 

.'- jn J*2fiti7. /.• 0.IW 

1 Muoia. -S m iImmoi '"phvlogenclic »meJ hitfc 
Nhigfi^ruphiuil reluiinnsliips ut tliL gonon in lire tnnnly 
AglilllidHC (Replllili l-acerliliui- (PhD thesis t'tuvriMly 

, r Mil ■iti^jno 
: vvi. i C j. (J962) "t. iHiiputanvr ruornTwIugy and 

kurynlogv of the Xusiiiilir-n niCtnhOTS mi \iu_ liiiinly 
\".;tiT,u[ ( j. i ,'hylavvrtetir irnpli^ition^ (fhDllurvi.-.. 

activity during die day- Inierestingly, Ihe Vidnntarv 
maximum body teinperalure (43.'1°C) was achieved 
during the night, indicating that some nocturnal activity 


Mnrpholtigy mui wlatiomhtys 

The LitOmties of C. ftickenrici and C. sviuulwus to 

othef t(M remain uncertain, PtMktii d^u stated 

wilhoul providing evidence. Ihal C. scutuluius and ( 
i ri\ratu.\ (Ciuy IS4D were "obviously rather eloselv 
relaBttl". Moody (1980)' added the C ttunticutcu^ 
group to the Inliet complex, diagnosing it on die basis 
Of&hap&Ofthc nuxh.-il ptiK'rss.if ihe pivanicul.irboni-, 
limb length! body size, and presence ola nuchal crest 
c\tK\ keeled vertebral scale line. Storr 114X2). fa 
resurrecting Cwtiophonis, tlid nol place ( . 
t fjitfii' n>iin\, (\ | n\hiu<\, ( »h -krtijft or ( 
Mtittttatus m any ot the species-groups within the 
genus, Subsequently, Siort et til (1**83) placed all lour 
species Needier with the C ictnulains SpCOKS-gTOUp 
o( Storr (IM66I and C dpcresii species gTOUfi til 
Houstitn (I47S) in H single expanded (', Jf<n\ 
species group, noting thai this was merely an 
assemblage^ and sssociartcd C mokruzici and ( 
uittuluius in an undiagnosed informal subgroup, linked 
by inference (Storr ft at 1983: 32) with C VtiSkltM 
Wiifen (MX2\ I9S_S) placed ( '_ ^riimiiittis with the ( 
ihih nitidis gp.nip and C. t ri$tufu.\ with C vatulii iiultu 
in a C ihstuius group. Both groups shared dill k \entnil 
markings aiul posrenor inlcrscalar lemoral and prcanal 
poic v The < . i ii\min; group was differentiated from 
the C. ttuuulittus group by the more widely spaced 
pores, and smooth (v.\ usunlly keeled) venlral scales. 
II ic piesence ol a vct'iebrnl scale ridge and a nuchal 
ridge vwis considered diagnostic Tor the C <ristafits 
group, although the presence of holh dark ventr.J 
palleiii and a vertebral scale ridge in C. smtnluius was 
considered primitive within the C. nuuuhan\ group 
A vascuJar tissue block deep lo the vertebral and nuchal 
ridges, possibly functioning in civst erection. w;is noled 
for C cvisttUHS ;ind C i-antlich\ctus. but was nol found 
m flic v . W[Ji UkfM group members examined (which 

did nol include C. scuttiltittis), Citsi dtctical was noted 
for C rrixkiTWkt but not ford vaut/inm iw. We h 
observed nuehul cresi erection in both C VQHtitciHi M.\ 
(Cj. M.S.) and C scumhuus (M.P.: Storr et nf. 1083 
Plate 2 1 However, Wttren (1982) considered this 
sasculur tissue block to he plesiomorphic within 


Body, head and limb pioporlions are similar in C 
rri\iiVti,\ l C, i and C S£ utuhUtts (Table hand 
,dl sluue similar male ventral pattern, a nuchal vtftsA 
lund a vencbiiil line of enlarged strongly keeled seales. 
the latter unlike memhers of the t'. mueulatu\ species 
group- Consequently, we believe that the nffinities a\ 



G mckenziei and C. scittidafus are with C cristatus, 
although it is clear that a more rigorous claclistic 
analysis of the phylogenetic relationships within the 
Australian agamic! radiation is sorely needed. 


The ant-dommated diet of C. mckenziei is similar 
to that reported for other small Ctenophorus species 
(Pianka 1986; Baverstock 1979; Mitchell 1973). 
Although hidomyrmex spp. were the most commonly 
eaten ants, their dominance may simply reflect 
availability rather than any selection by the lizards (E. 
Matthews, pcrs. comm.). 


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 G decresii complex. The 
observations presented here show that this is clearly 
not the case. Whether this behaviour is homologous 
in C. mckenziei and the C. decresii group is unclear. 

Thermal preferences 

The mean body temperature of C. mckenziei reported 
here is lower than that reported for C. scuttdatus in 
a laboratory gradient by Licht eral. (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 et ah (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. 
mckenziei 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 1991; G.R.J., G.M.S., 
pers. obs.). A constant source of heat in a thermal 
gradient 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 scuudatus: SAM R1459a-j, R3024a-b, R48I4a-l 
(syntypes), between Fraser Range and Queen Victoria 
Springs; WAM R1235, R1761, Laverton; R2841, Gutha; 
R5306, Wadgingarra; R81 70-71, "Yuin"; R9352. 
Malcolm; R9510, Morowa; R11236, 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 Malice Hen Rocks. 


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 is 
thanked for field assistance. 


Anonymous (1988) City intrigued to discover thai some mice 
are still rare. Enpkm (23), 15. 

Armstrong, G. P. (1992) Reptiles pp. 163-179. In Copley, 
P B. & Kemper, C. M. (Eds) "A biological survey of the 
Yellabinna region South Australia in October 1987" (Dept 
of Environment & Planning, S. Aust.). 

Baverstock, P. R. (1979) A three year study of the mammals 
and lizards of Billiat Conservation Park in the Murray 
malice. South Australia. S. Aust, Nat. S3, 52-58. 

Bedeord, G. (1991) Additional nocturnal sightings of three 
species of dragon lizard in nothern Australia. Herpelofauna 
21, 30. 

Brattstrom, B. H. (1971) Social and thermoregulatory 
behaviour of the bearded d ration, Amphibohmts harhatus. 
Copeia 1971, 484-497. 

Copley, P. B. & Kemper, C. M. (1992) "A biological survey 
of the Yellabinna region South Australia in October 1987" 
I Dept of Environment & Planning, Adelaide). 

Cuvier, G. L. C. F, D. (1829) "Le Regne Animal" 
tDeterville, Paris). 

Ehmann, H. (1992) "Encyclopedia of Australian Animals, 
Reptiles" (Angus & Robertson. Sydney). 

Fyfe, G. (1981) Nocturnal sightings of two dragon species. 
Hcrpetofauna 13. 34. 

Gibbons, J. R. H. (1979) The hind leg pushup display of the 
Amphiholurus decresii species complex (Lacertilia; 
Agamidae). Copeia 1979. 29-40. 

Gray, J. E. (184!) Descriptions of some new species and tour 
new genera of reptiles from Western Australia, discovered 
by John Gould, Esq. Ann. Mag. Nat. Hist. 7, 86-91. 

Houston, T. F. (1978) "Dragon Lizards and Goannas of 
South Australia" (South Australian Museum, Adelaide)- 

Licht, P., Dawson, W. R., Shoemaker, V. H. & 
Main, A. R. (1966) Observations on the thermal relations 
of Western Australian lizards, Copeia 1966, 97-110. 

McKen/je, N L., Roif-e, J. K. & Carter, D. (1987) 
Reptiles pp. 179-210. In McKenzie, N. L & Robinson, 
A. C (Eds) "A biological survey of the Nullarbor region 
South and Western Australia in 1984" (South Australian 
Department of Environment and Planning, Western 
Australian Department of Conservation and Land 
Management and Australian National Parks and Wildlife 



& RoniNsoN, A. C. (1987) Ibid. 

Mitchell, F. J. (1973) Studies on the ecology of the agamid 

lizard Amphibolurus maculosus (Mitchell). Trans. R. Sac. 

S. Aust. 97. 47-76, 
Morley. T. P. <& Morley, P. T. (1985) An inventory of the 

reptiles eff Dan^gali Conservation Park- Herpetojauna 15, 

Peters, W. (1864) Ubersieht der von Hrn. Richard 

Schornburgk an das zoologische Museum eingesandlen 

Amphihien, aus Buchsfclde bci Adelaide in Sudaustralien. 

Mber. K. Preuss. Akad. Wiss. Berlin 1863, 228-236. 
Peters, W. (1866) Eine Mitteilung iiber neue Amphihien 

(Amphibolurus. Lygosoma, Cyclodus, Masticophis, 

Crouiphopeltis) und Fische {Diagrammu : Hapalo^enys) des 

Kgl. zoologi.schen Museums. Ibid. 1866, 86-96. 
Pianka. B. R. (1971) Notes on the biology of Amphibolurus 

cristatus and Amphibolurus scutulatus. West. AkSt. Nat, 

12, 36-41, 
(1986) "Ecology and natural history of desert lizards. 

Analyses of the ecological niche and community structure" 

(Princeton University Press, Princeton, New Jersey). 
Schwaner, T. D. & Tyler, M. J. (1985) Reptiles and 

amphibians pp. 56-67. In Aslin, H. J. (Ed.) "A list of the 

vertebrates of South Australia". 
Shea. G. M. & Peterson, M. (1985) The Blue Mountains 

Water Skink, Sphenomorphus leuraensis: a redescription. 

with notes on its natural history. Proc. Unn. Hoc. N.H.W, 

108. 141- J48. 

Sterneelu. R. (1919) Neue Schlangen und Echsen aus 
Zentralaustralien. Senckenbergiana I, 76-83. 

Stirling, E. C. & A. (1893) Verfebrata. Trans. R 
Sac. S. Aust. 16, 154-176. 

Storr. G. M. (1965) The Amphibolurus maculatus species- 
group (Laecrtilia: Agamidae) in Western Australia. J. R. 
Soc, West. Aust. 48, 45-34. 

(1966) The Amphibolurus retiadatus species-group 

(Lacertilia: Agamidae) in Western Australia. Ibid. 49, 17-25. 

(1981) Three new agamid lizards from Western 

Australia. Rec. West. Aust. Mus. 8, 599-607, 

(1982) Revision of the Bearded Dragons (Lacertilia: 

Agamidae) of Western Australia with notes on the 

dismemberment of the senus Amphibolurus. Ibid, 10. 

Smith, L. A. & Johnstone, R. E. (1983) "Lizards 

of Western Australia, II. Dragons and Monitors." (Western 

Australian Museum. Perth). 
Wilkinson, L. (1987) SYSTAT: the system for statistics. 

(SYSTAT Inc., Evanston). 
Wilson. S. K. & Knowees, D. G. (1988) "Australia's 

Reptiles. A photographic reference to the terrestrial reptiles 

of Australia." (Collins, Sydney). 
Wittrn. G. J. (1985) Relative growth in Auslralian agamid 

lizards: adaptation and evolution. Aust. J. Zool. 33. 





By Stephen C. McKillup* & RuthV. McKillup* 


McKillup, S. C. & McKillup, R. V. (1994) Reproduction and growth of the smooth 
pebble crab Philyra laevis (Bell 1855) at two sites m 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. 

ttomtrtfcm »! ito IfyM Scteitf* »l £ #&. iiw4». UJttf). 2*5-2*1 


by stvphrn C. McKjllip* & Ruth v, McKtu lp 


Ma.n un\ SX\ & M.KiLLtf, R, V, U994) Reproduction and growth ul Ihc- .smooth pebble crab Phihru hf\U 
(Hell hW> al two sites in South Australia during W90-91. 7/wjy. J?, -Sor. S-Ausi.. 11N|4/, 245-25K JQNlfttfctbw, 

The growth and lite history ol the smooth pebble crab Wftprt l"<v>.\ (Be!!! wns interred irnm collection* rmidt! 
bi-iwecn July l$9fl and December |99] from Sultana Point ant] Coohowie. South Australia, fi feitfifJ appeared 
to be scmelp.irous and to reproduce twice a vc-ar at both sites. At Sultana hunt ovieeroiis females wen; common 
in winter {June-July! and from late spring to late summer (November February' PswS WStfM* * efc " common from 
late winter lo early sprints and also in early summer. Distinct coIhkls or new recruits were found in early spnne 
and again ip earlv summer. Recruits gre* i.ipidlv, reaching sexuaJ maturity within lour months. The mean sue 
of jutllll females and the proportion which were ovigeroas liiffered amongst populations A tabontfory experiment 
showed that food availability affected growth and whether females became oviperous. We postulate that differences 
in size and ovisery amonesi populations of Phdyra tuteis may be- caused by differences in the availability of food, 

Krv Wciro.s Phthra. pebble crab, life history, intenidul, food, sandrlat 


The smooth pebble crab. Philyra laevis (Bell 1855 K 
IS common on sheltered intertidal sandflafs in southern 
Western Australia. South Australia, Tasmania and 
Victoria (Phillips*^/. 19S4> Hale (1976) describes 
the feeding and courtship behaviour of P faevi\, but 
iixilc else is known about this crab. Tasmanun 
populations hfffi laevis have been reported to host a 
ucmciteun (genus Caranonemertes) (Bell & Hickman 
1985) and the trcmatode Microphallus para^rapsi 
Smith. 198-MBell I98S). From W90-9I wc sHidjed the 
feeding behaviour of Philyra laevis. finding that 
although individuals often fed upon other dead or 
damaged animals, many were unwilling to teed upon 
members of their own species and that Water borne 
cues from damaged P laevis inhibited feed me 
(McKillup & McKillup 1992). During the study, we 
collected P laevis over an J8-month period from two 
sites 10 kilometres apart in South Australia, and also 
sampled 10 other southern Australian sites. These data. 
logelhei with results of a laboratory feeding 
experiment, are used lo infer the lite history o\' P laevis. 


Sampli/ix of Philyra laevis /row Sultana P>int and 

Samples, of Philvra laevis were collected from the 
mtcrlidal sandflat at Sultana Point, lower Yorkc 
Peninsula, South Australia <35.0X°S 137 44°P.> in July 
I9'M). and then approximately every four weeks from 
September 1990 to August V991 and in Decembei 1991. 

* Department of Zoology, LruverMlv OS Adelaide, South 
Australia, 5005 Australia, 

* Present uddre^ Depart meni of Biologs, Central 
()ueen.dund Universitv. Rockhampton, Queensland 4-702 

I he sandllat at Coohowie Nay, 10 km north of Sultana 
Point, was sampled in the same way but less frequently 
(November 1990 and January. Maah. April. August 
and December 1991) 

P. laevis is active whilst the BOIKfflAI fc covered by 
water during ebb and rising tides, bul remains buried 
in the substratum ul high water or when the sandflai 
ts completely exposed (Hale 1976). At least 30 and 
usually more than 100 individuals were collected from 
within the interltdal zone where P, laevis occurred 
(from low water to about mid tidal level! between the 
time of low water and when the sandllat was mundated 
by The nstmj tide, At each site at least 160m : of 
substratum, consisting of .several haphazardly chosen 
0.5 metre wide strips from the waters edge lo mid tide 
level, were hand searched to a depth of 3.5 cm, P. lathis 
was found no tteeper than 3.0 em in the substratum 
(McKillup & McKillup unpubl.j. Crabs were frozen 
and later examined tor sex. whether females were 
ovujerous. and the carapace width of all individuals 
was measured to the nearest 005 mm. 

Reproductive tottddion and average size oj adid; 
females at additional sites 

During January and February 1991, at least 73 
Philvra laevis were collected from each of 11 sites m 
South Australia and one in Victoria (Swan Bay within 
Port Philip Bay), and examined tor sex. sue and 
reproductive state as described previously. 

hihoratnry e.vperitnent on sexual development and 

Abdominal morphology in the Crustacea can usually 
be useo to determine sex: adult males often have a 
relatively narrow, concave sided and tapered abdomen. 
whilst leiualcs have a hroader and often circulat 
abdomen almost as wide as the carapace (Hannoll 


-. ' McKH.l |JP (& R V M.hlLLUf 

I u- I. ArxJomirwl im>rplu>tugy <i! Phi(xr ( i /<vo7a, from left 
h. rljshi; tiiMc intcnscdiatcj = juvenile remalt- 1 **n*l .uiuli 

I982i I 'us was the case lor all i'ltilvnt W'7,, wilh 
carapaces wider than US mm, hul three morphological 
type;-, were found in crabs 13,5 mm wide or smaller 
Ihc two previously described forms, plus individuals 
with a convex sided abdomen which, ai its' widest 
point was approximately hall'the width of the carapace 
if ig I) The last dewiibcel individuals were named 
'intermediates , 

The growth of 20 females, 20 unci mediates and 20 
males all of carapace width 10.5 mm or less, wa- 
Observed in the laboratory Considering thai vw also 
tumid differences m ilk- si/c and proportion of adull 
lemales which were ovigcrous amongst sites, and thai 
differences in adull si/e amongst iunspcedledi 
lUjj a) i lies were also reported by Hule (1976), 
observation* were made as pail of a manipulative 
laboratory experiment designed !o examine the effects 
ol lutid availability upon growl h and egg production 
of Mfilvtit /tun:,. Crabs were placed individually in 
SO \ 50 v _s() mm deep plastic dishes, caeh tilled ft ith 
10 ml n\ seawaiti, All individuals were numbered on 
the dorsal side or their carapace with non tovu. 
vvaicrproof ink- Six dishes, containing two mules, two 
females and two intermediates were placed within each 
of 10 lidded 280 x 3K0 :•: ilO mm deep p|.,-ai.. trays 
containing seawatcr to a depth ol 10 mm. Aeration was 
not needed The seawaler in the trays maintained a high 
level nf humidity whn. h reduced evaporation I'rom the 
dishes and also provided a marine environment for the 
i, W i r.ihs which ClTirUtPd out during the cxpeiimcnl. 
Trays were kepi at mum temperature and ualurul day 

rive traV$ were assigned randomly to a "high food" 
treatment and the M) crabs within these were fed every 
working das (from Monday lo Friday), whilst the 30 
nabs in the oilier five" (rays were led wcekh. on 
Wednesday-, .is .. "hiw ffati* treatment. Dil'IcrciU 
frequencies of ieeding provide different levels of food 
ilVdlkblllty I" mverlcbajiest.;,^. Calow 1973; Monarly 
W7S), For each Ieeding the ^ix crabs in a tray were 
removed placed in sis separate dishes used for feeding 
only BJld offered od libitum ciushed cockle. Knleh'sio 
\ifihtrhui Lamarck from Sultana Pomi P fut'viv is 
often found feeding on this bivalve in (he field 
INK K.llupcv- McMlup 1992) All leO fot 12 min 
or less and wen:- returned (o their permanent dishes 

after 15 min. This method ot feeding prevented the 
scawaiet in ihe permanent dishes from hecomlne 
fouled; it was replaced lorinighlly. Crabs wen* 
inspecled on every working day. individuals which had 
moulted were examined and sexed as either male 
lemule or intermediates, .md ,my which had i limbed 
out were replaced m their dishes. 

The experiment began on April 2nd 1991 using new 
recruits from Ihe December 1990 January 1991 cdhpfl 
ill Coohowie. and continued until all crabs had moulted 
ai leasl once. 


$atnp/itix $( Philyra laevis fnun Sultana hunt cW 

( rHfhnutr 

the frfe& structure of Ihc population a(Pli!l\ra hirvi\ 
al Sultana Point from July 1990 to December 1991 i- 
illusiraied in Figs 2. 3. In midwinter (July 1990), all 
males were at leasl 8 5 mm wide, all lemales :it least 
15 mm wide jihI 93% tit tire killer were ovigcrous, 
By early sprine (September 1990), dead males and 
females wen. extremely common and large live 
individuals were not: almost half of the live males 
collected were smaller than 8 5 mm and appeareti to 
be ,1 dimmer cohort of recent recruits. Only iwo live 
adull females were found; both were more than 75 mm 
wide and ovigcrous. The rcmamdei of the sample 
consisted of "intermediates" 7.5 mm W'idc or smaller 
which were no! present in the July sample. A 
lahoraiory experimenl showed lhat intermediates wen 
juvenile lemales (see below). In mid-spring (October 
I990J only one cohort of males was present, and thesv 
malcis were large: than the recruits first seen m fhc 
previous month The juvenile females were also larger 
and some relatively small adull females were preseni. 
bul none whs ovigerotis, In late spring (Novcrnhci 
1990), uo juvenile females were found; the sample 
consisted ol aduh females al leasi 9,5 mm wide. 89$ 
of which were ovigcrous, plus males from 4,5 |o 
18,5 mm wide. In early summer (Dccembci WOi 
anolher cohort of relatively small males was present 
together with a cohort of luvemle females up to 9:3 mm 
wide. Dead adults were again common, but live adults 
were also found and 7KV of adult females were 
ovigcrous^ The small nudes and juvenile lemales 
continued to grow through summer and autumn 
(January lo April 1991) and the number of ovigcrous 
adult females declined (30aV-v. in January. 14% m 
February and 5Vr m March). Prom mid to late- 
autumn (April and May 1991.1 no juvenile females were 
found and oniy 1.5% and 2% respectively of aduh 
females were ovigcrous, but in early wintet (June I99l}« 
the percentage ol ovigcrous females had risen lo 6X r , 
and further increased lw 94% by late winter (July 1991) 
The early spring (August 1991) sample was very similm 
lo dial of September 1990; dead adult males and 








— ^ 









13th July 90 
M Hill ll , 



19th Jan '91 

■kUUfrl 1 ITTTI T 1 

,4 5 

27 b C 


7fhSep , 90 




14 5 27 5 


6th Oct '90 


27 5 



8th Dec '90 

Ll ill 


ISfri June '9 

* ■ i i ' ■ < i i i 


3U ' 

23rd Feb '91 




■ ■ i i ,■ r ■ ■ . ■ , 



23rd July '9" 

M >'.>■■ , ;■ rrr . 

14 5 27 5 

3th Apr '9 


,9th Dec '91 



n J , P . >'.i ; r , ■ 

27 5 14 5 27.5 

25th May '91 


27 5 

I 1 1 n n r i i i i i 
14 5 27.5 

Carapace width (mm) 

Fig. 2. The size structure of female Philvra laevis collected from Sultana Point, South Australia on 14 occasions between 
July 1990 and December 1991. Solid bars indicate juveniles; open bars, adults. 

















"0 t , 30 

27 5 

27 5 


17th Sep '90 

,j L.j i,i 


27 5 




8th Dec '90 



27 5 

Carapace \A/idth (mm) 

Fig. 3. The size structure of male Phihra laevis collected from Sultana Point, South Australia on 14 occasions between 
Julv 1990 and December 1991. 



(emales were common and the sample oflivc Philxra 
Uu'Vts consisted entirely of juvenile females less than 
K.5 rum wide and males which were almost all smaller 
than those collected during the previous month. The 
December 1991 sample was similar to the one ftom 
the previous December, consisting of adult males plus 
adult and some juvenile females, with h$% of adult 
females ovigerous. Dead adults were again common 
OTI the sandtlat at Sultana Point in December 1991 

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 lemales 
were also present and dead males and adult females 
were common. The average carapace width of the 

cohort of juvenile females present in early autumn 
(March 17th 1991) was larger than in January and most 
appeared to have moulted to adults by April 13th 1991, 
In late winter (August 18th 1991) dead adults were 
common and juvenile females plus small males were 
present. Jn early summer (December I9lh 1991) dead 
adults were also found, very lew 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 females 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. 

TahlI: I. Comparison of the mean carapace width (in mm) of adult female Philyra lacvis /row Coobowie and Sultana Point 
tin November ctdicelions were made 17 dcivs apart; rth the other 5 accnstotis crabs were Ci>liected front both sites ^>n tin 
same day, \ — mean, s ~ standard deviation and n — somple size 

Dale collected 



Sultana Point 

November 1990 
19 Junuurv 1991 

17 March 1991 
13 April 1991 

18 August ISBN 

19 December 1991 












1 73 



1 42 













none louml 







Tabu; 2. Hie number of adult female Philyra lacvis and those twigerous in samples atlleeted from Coobowie and Sultana 
Point on 6 occasnm,\ during the same month. The Nttvemhtr colleetnms were made 17 days apart; on the other 5 tteeastons 
crabs were collected from both sites on the same day. 

Date collected 


Coobowie Sultana Point 

Collected Ovigerous Collected Ovigerous 

November 1990 
19 January 1991 

17 March 1991 
13 April 1991 

18 August 1991 

19 December 1991 
















1 abu 3. 77u' percentage ofovfgerous adult female Philyra lacvis at 12 sites sampled during January and February 199J, 
together with the mean carapace width of adult females from each site. 

Date sampled 


Number rff 
mature lemales 


Number and 

Mean carapace 

width of 
females (mm) 

19 January 

19 January 

(9 January 
!9 January 

19 January 

20 January 
20 January 
29 January 
29 January 
2 Fehruary 
2 February 
13 February 


Edithburgh Bay 
Sultana Point 
Hickey's Point 
Pmnl Turum 
Rogues Point 
James Well 
Pine Poml 
Foul Buy 

Stun Bay 
Swan Bay (Vic. 


















14 35 


























\ r Mi K It I i"- v U V Mr KJI.LUP 

Rcprofhifiin- condition and avenj^ Kite rffoftfttlts iff 
addifiomd tfWJ 

Data for the numbei of adult females c* 'I Iceted and 
the number and percentage whiih weiv OVigafOOg lb) 
12 populations .sampled in mid lo lute summer 1991 
are in Table 5; The percentage of ovigerous females 
decreased at Suliana Point from January to February, 
...onlv samples collected on ihc 19- 20th January 1991 
woe compared statistically- Nevertheless, the 
proportion ovigerous difleicd significantly amongst the 
seven sites sampled on lower Yorkc Peninsula \Z& ' 
contingency tabic comparison, di * — 6. Chi squared 
StttlSti* - 122$ ?<> (MlllSj The :• igMt'u. ..." 
heterogeneity amongst sites was largely due to Ihc 
greater proportion of ovigerous lemales at Sultana Ppmi 
fiable 11 c'uiv-.dering all sites sampled, the highest 
percentage ol ovigerous aduh females (KV* > was at 
Swan Hay, Vm*h iu dimng earlv Fchruaiy (when only 
I If were omgcjous at Sultana Poinl, see earlier 
discussion oi Figure 1). The Swan Ray population also 
contained ihc largest Phi/yru (oca. lotind 
Furthermore t-u the seven site-, sampled Horn the 
19 20th )anuary (excluding Point Turton where only 
three lemales wen? collected), ovigerous females were 
lound onlv at sues where the average carapace widlJi 
.»! Imiales was greater than LV00 mm (Table 3)- 

hthontutn vyu ntneor on wxml dwe\tipw?m 

Male, adult lemale and ■intermediate** Phdxm Uu>vt\ 
ottered food on live of seven days per week, moulted 
SOOncr ttalfl (hose only \h\ once per week ( Fable 4j, 
At then first moult in the laboratory all males moulted 
in males, all lemales lo females and all mlennediaLes 
to females, except for the two smallest which remained 
as the intermediate form unril they moulted again. In 
all cases the variance of days elapsing before moulting 
was grealci in the low food treatment, and by 
inspection the distributions in ihis treatment w«nc 
.skewed lo the fight. None of the adult females in ihc 
low lood rrentment, but all in the high food treatment 
were ovigerous by July ls>9l 


Pbiivni fai'vts reproduced twice a year at Sultana 
Point ami death of most adults during the breeding 
moudis suggests Otis species is largely sciiielptirous 
kLvniit;. found in caily spring (September 1990 » 

reached sevual maturity and reproduced from late 
spring to laic summer (November 1990 lo February 
1991). while those first lound in early .umnier 
(IXxcmbct I'WO) reached sexual maturity by nud 
autumn (April 1991) and reproduced until early winter 
(June l L )91> Data from Coobowie were consistent with 
this pattern of recruitment, but suggest that many adult 
P ldfvi\ at Coobowie did not reproduce in the summer 
of 1990-01, although daUi were scanty, being onl v ft 
November 1990 and January 1991 

Recruits were found one month after reproducing 
females were present in laic spring and two momh 
alter they wvre present in early wuitei. Considering 
that Ihe sandflal was only sampled monthly, that the 
smallest crab found was 2.8 mm (McKillup & 
McKillup unpubl.) and that smaller individuals Were 
likely tii be overlooked amongst sand grains ami 
detritus, the larval stage o\ R iacvi,^ is likely to b t - of 
relatively short duration (perhaps only 1-2 weeks) 
Another member of the same genus, this purse crab 
Philvra globnsa (Fabricius), has a larval stage lastim- 
II Jays in (he laboratory at an average temperature ot 
2K°C (Knshnan &. Kannupandi 1990). 

The moult from juvenile to adult form in females 
appears to coincide with sexual maturity, since only 
iwo females with juvenile abdomens of more than M'H) 
examined were "vigerous (McKillup & McKillup 
unpubl.). A relative Cnon allomelrie 1 ') increase in 
abdomen compared to carapace width during the moult 
lo adulthood is common in braehyurans (Hartnoll 

For sites sampled I mm the l9-20lh January 1991. 
a greater proportion of ovigerous adult females was 
present where the mean carapace width of females was 
relatively large Also, there was a greater proportion 
of ovigerous lemales at Sultana Point than Coobowie 
on all occasions when comparison was possible. 
Differences amongst sites were nut caused simply bv 
larger lemales being more likely lo he ovigerous; 
examination of the data used to compile Tabic 2 showed 
thai in November, 60 of n9 adult females between 8 
and P mm wide were ovigerous at .Sultana Point. b..i 
none ot 19 collected on the same date and within the 
same size range was ovigerous al Coobowie 
I -Lirthcnnore- on January l n th KJ9L none of the 17 aduh 
temates collected from Coobowie was ovnierous. eve-n 
though the carapace widths of these individuals were 

IAHI K fl Tbt tumuli Ju\\ clufv.itii: bedm- Ph>l\ia laevfii fir\i muulnxl /" hfah (tttd htwjboit trsunneW^ i "tntt\ 
v, sumdurd Jwuttion. n = Mitn/iic Ytee 


[•iii;ile i 
h'i.. r0ttJifllr$ 


1 !.„,J 


Treat men 


I.-w Fund 



>A 10 

' ' (in 

7 38 

ill 79 



3s> 10 
45 20 

3H 10 




within the si/e range of the 15 ovigerous females 
collected from Sultana Poim. Similarly, on December 
17th M)l, none ol o adult females from Coobovvie was 
<>\ Igtja&USj even 'hough mk were within the va;- range 
of ovigerous females rrom .Suliana Point. 

Results of the laboratory experiment were consistent 
with food supply allcctmg the frequency oi inoulimp. 
A sample tfl crabs will contain individuals at different 
Mages of the moult cycle In the low lood group, crabs 
aboul to moult would have done so soon after the 
experiment began, hut those which had moulted shortly 
before being collected would have lo prepare to moult 
again under the laboratory conditions of low linni 
availability, In contrast, in the high lood group. crab> 
prepared to moult would do so. and others which had 
recently moulted would have adequate resources 
available to jjrow and moult again This can explain 
the greater mean, variance and positive skew m the 
number o! days before moulting in the low compared 
io the high food irealnicni iTahlc 4i. 

We suggest, for the following reasons, that 
differenees in si/c and the proportion oi ovigerous adult 
females amongst sites were caused by differences m 
food availability Firstly, in (he laboratory, crabs ;n lite 
high food treatment moulted sooner (and iherelure 
iirew faster) than those in the low looJ treatment and 
only females in the high food treatment pnxJuced vgjjv 
Secondly, adull female P. lacvis al Coobovvie were 
consistently .smaller than those at Sultana Point The 
inlcrtida! scavenger NffltSttiiUS fnmpa'tiitts (Lamarck). 
a prosohranch snail which occupies a similar niehe and 
is often found feeding with Philyru Uwvi.s, is also 
smaller (and hungrier) at Coobovvie than at Sultana 
Point's and we have postulated there is less food 

available lo \. paupciuius at Coobowic than al .Suliana 
Point (MeK. Hup & B u ,| cr 1979, l l JS3: McKillup lOKet) 
P'urthermore. we have wore recently postulated that 
intenidal scavengers may generall> be short of food 
(McKillup & McKillup |W)- All Suuih Aiistialun 
populations o\ P liwvis sampled contained snialici 
females on average ihan the population al Swan Ray 
Victoria, .suggesiing that food availability may bi 
limiting the growth and reproduction of man} 
populations, of this scavenger Numbly, lentales i>| 
another leucosiid crab. Hbuiia Javvi.s (Bell, I855] wen 
only found oAigerous during Decemhet. Januaiv. May 
and August at Wellington, New Zealand, but records 
from other localities suggest the duration Of ibC 
reproductive season vanes amongsl siLes (Wear k 
Fielder WS5J. 

The hypotheses that differences in food avuilabilav 
a,rO limiting the duration of the reproductive period t$ 
Pfuhra Uh'Vi> and that individuals in natural 
populations of this species ate short of h)ud could be 
tested b> frequently feeding marked individuals in the 
field and comparing their growth and reproductive 
output with the icM of the population, the results ol 
these experiments may explain vvny Philxni /t/ev/.v fob 
a relatively short lifespan and two seasonally opposed 
breeding seasons in South Australia. 


We wish to thank Alan Bullet and Keith Walker 
cement during the study, ~ 
es, n<-M p|oltta W\ Mk 
onsliuclive comments on a diall 0| Ihe 

We wish to tiianK Alan tsuiiei and Kciin walker for 
(heir encouragement during the study, and Alan Btuler 
Michael f'Vih'i. tVin kd*iOi*i tuui a I u -t • Wolk for ilioii 

uien ciiLuuiaucuiirm um mii inc suiuv. uqg man otuiei , 
Michael Coates, Hon Heidei and Allee Wells for their 
critical and eonsliuclive coiituteiU.s on a diall 0| I he 


til i i , t* J, (| l >8Xj A siucly nl the lilc-histnry or .U/. /-,•'/<', <>///< \ 
pan/^m/ni Smith N8J ilremaiorlu: MierophalluJaei )\m 
Vi<>< R"\. Six lasm- \ll lt<M25. 

& HickM-w i L. [W5j ot>siT\uii.»!<x rm 

f rifi j mm erne rit j s ( Neme-'i