Climate change and its implications for the terrestrial vertebrate fauna

10 Pages·1994·5.3 MB·English

by  Philip Stott

M
J)wi>w:itt»n at to$&ri Softrty oj'S. Aust (199-Ji. LISllt. $9-i&
CLIMATE CHANGE AND ITS IMPLICATIONS FOR
THE TERRESTRIAL VERTEBRATE FAUNA
by Philip Srorr
Summary
Sum. P. (1994) Uimuie change and its implications tor the terrestrial vt?r1ebiate Iuuhu (IW4). ftaiis. r? .W1.
$ JteSl 118(1), 59-66. M May. 1994,
A limited number|)fstudiesotthebiology ofbvarietyofspeciesot terrestrial vertebrate* isusedtospeculate
Ht*>uiiheu respondsloclimatechangeuspredictedby globalcirculationmodels. Dramaticchangesinthedistribution
and abundance ol animals in Austodia that has already occurred consequent to European seUleiiieni h noted
Speculationaboutthe impactol"climatechangeonther^Uilivcabundanceofmammalsandreptiles, rangechanges
ofkanuaroo-., oibbtiv.inaiidareas,1oodsecurityoftheSpimfcx HoppingMouse, eompeiiiionbetweentwospecies
ofskinks. anddiseasetransmission is included. Nest-site selection by tortoisesand social structureoftbxe.s are
given astwoexamples where behavioural plasticity demonstrates somecapacity tocope in xitu with me effects
ofclimate change, but the ability ot must terrestrial vertebrates to track rapid climate change aerify* different
NUhstraies isquestioned. Reservations are expressed about the knowledge bnseupon which the speculations arc
based. Curwiy ttWspeciesisfisuneofstudies availabletocomparedetaileddataondistributionandabundance
with climateover a wide geographical ranged backed up with biological information sufficient to explain the
mechanisms by winch the species niterael with their environments,
KES WORDS:Climatechange.mammals, repnlt-s,Testudines.abundance,distribution, rainlall.thought.disease.
MtitTffius ftt&msfus, Mai ropusfuliginoxut, OnctwQjmi amkvh^
Introduction The sentiments expressed in this paper arc
complementary tothoseexpressed inessays by Arnold
Some 6000yearsago. within thespanof100 years, (1988). Busby (1988). Graetz U9K8), Mam(1988), and
theNegeualionat Cold WaterCave. Iowa, changed from Fossingham (1993). The paper augments previous
tvircs*toprairie (Ooralec/<//. 1992)- Thechangeswere contributions by exploring some examples of the
associated with atemperaturerise ofc ^°C,and with mechanisms by which climate change might affeel
such a complete change in vegetation structure must vertebrates It isa speculative paper; ihe scenariosarc
havecome aprofound change in the vertebrate (anna presented with the intent of illustrating the level of
ol the locality. It may be. during thenext century, that complexity, rather than making confident predictions
similardramatic changes in biota will result Iroman about the outcome. The predictions 3re weakened
anthmpogenieully-eiihanecd "greenhouse"effect The becauseasingle study ofalpCdl populationoverashort
terrestrial vertebrate fauna of Australia has already period oftime is not necessarily representative ofthe
undergone profound changes as a consequence of biology oi a species during alL seasons and over the
fcuropean settlement (Reeher & Lun 1990); climate whole of its range (see Kemper ft G& 1987). and the
change would compound the impacts of introduced range and habitat ofa species as observed to date do
competitors and predators, destruction and not necessarily represent accurately the realised
fragmentation ofhabitat, altered fire regimes, hunting, environmental nicheofthespecies, northis in turn its
and diseases. fundamentalnichet,Possinghain 1993). Inaddition, the
Realistic* predictions about the impact oi' ehmate predictionsofglobal climatemodels are noteonsrdered
chanceon thedistribution and abundance Ofterrestrial tobe rcliahleatthe regional level (Gordonctat. 1992)
vertebratespeciesrequire a foundationot information As the numberoflogical steps increases, sodie etrors
from several sources. Essential are detailed forecasts are summed.
m
Itomeltmaiologists; predictionsabout changes the The paper reviews those climatic predictions of
distribution, composition, and productivity or plant particular importance toanimals, uses someexamples
communities and other elements ol food webs, and tu explore the mechanisms by which climate change
detailed information about the present distribution, might affect terrestrial vertebrates, thencxarninesthe
abundance, physiology, andecologyofvertebrates. The capacity ofanimalstocopewiththepredictedchanges.
temporal scale of possible changes should also be
considered. Predicted climate changes
* MUaonpiageomfentf.rnvRiorsoenwroaretnhtyalCaSmcpiuensceof tahnedUniRvueirnsiictlyanodf Evidence from climate models indicates that
Adelaide. South Australia *>0('S emissions of "greenhouse" *ascs into the atmosphere
DO P STOTT
will cause global warming of0.3 (ranee 0.2-0.5) °C the easternand western Pacific Ocean, ENSO would
per delude, and associated changes in patterns of be weakened (Rind 1991) ihus tending to reduce
precipitation (Houghton**/at,ls>92). Whilst thegeneral elimaijt. variability ineastern Australia and couutennv
predictions *>f the models are broadly accepted, the to some extent the predictions outlined above
issue Ucomplicated by the lack til clearobservational
evidence ofchanges attributable with certainty CO an
rnhaneed "greenhouse"clfc'cl. On examining records Direct effects
of meteorological observations, Nicholls A: Lavery
t1992t mill ItOI able to identify any clear trench Ln Direct effects ofeiincite on vertebrates should Ik
more readily elucidated than would oethe case with
rAauisntfraallliaatupreluintaiblleI9mSeXt.eoArlotlhooguigchalBustrattuiwonss&inSt.aSpoluetsh indireci effects. Twoexamples are presented ol ditect
()9ylJ note a WflfRUCf trend in South Australia since oeffftecetmspeorfactluirmea-tdeeopennidcepnttilcsse.xTdheteerlmuisntaitsiotnh,e parnodceusVs
bWo.sutnI,dsihocfypcaastuteixopneerdietnhcaei".thMeotdrecilusi awraesno"tVlcoosncsilsetetnhte second is the influence of temperarurc on the
distribution oficstudines (turtles and tortoises) Hoih
in theirpredictionsorseasonalchanges in temperature
patterns forSouth Australia(see Boerflal 1992; Gales examples demon.sltatc that the knowledge base is
W «/. 1992); al the present stale of refinement of the inadequateeven inthis more straightforwardutccgnry.
generalcirculationmodelssuch uncertainties inliming,
magnitude, and regional patterns are well recognised h'mi'i'ruiim-dvptndeni sex determination
iHnii^tiuui el Hi 1902).
Temperature-dependentsex determination (TDSD)
Rainfall has a major influence on food supply for has been demonstrated in many species ol lesludmes
venehraie species, particularly inanduicas llaarstna leg Mrowsky etaL J984) croenditiansfe.g. Webb
el at. fl9*>3j predict a global increase in tropical atil 1983) ar.'ngamid (Charfmer 1966 videBull 1980).
disturbances. At present, some i-I tltc^e distutbances und a eekkonid (Wagner 1980). Slight (<2°C)
extend into northern South Australia hs tropical- departures fium pivotal incubation lemperalures may
extraitopical cloudbands (TbCBs). and as a result result in eniircly male or entirely female sexual
heavy rain falls on an average of nine days per year phenoiypes, even in some of the species which have
tKuluiel 1990). However, the distribution and extern heiemmorphic s?x chromosomes (Servanffi\) WR9)
oftherainfall iserratic. Gordonrtal. (1993)cautiously Some species have twotha'sholds. with males abutting
report the results ol simulations which indicaicchanges from intermediatettnd femalesfromextremeintubation
indaily rainfall intensity across Australia, particularly temperatures (Ynlema 1976: Webb & Smith 19X4).
in summer, in the form ofincreases in the frequency TD.SDhas been demonstratedtooccur inan Australian
ofheavy <12.X-25.6mm. r. 31%: >25.6 mm. r, 95*) crocodilian (Webb ft al. I9X.V). an Australian
rainfall days, an c. \5% decrease in the frequency of frcshwriter-'cstuarinetesmdincfWebheral I9K6), tutti
light I <rt.4 nuni rainfall days, and a decrease in the Australian marine tcstudines iLimpus etat IWS). In
numberoframdays. Whettonetal. tinpressj review contrast, it does not oeeui in several Australian
the predictionsoffive global circulation models; lour trr-shwatertcstudines. includingspecies found in South
predict increasing, summerrainfall over the wholeof Australia (Georges I9«8;' Thompson I98«aj.
South Australia ull live predict increasing summer Preliminary studies suggest that TDSD occurs in at
raintall over much ofthe far north). <ind four predict least four Australian agamids (G. Johnston pfif3
a decrease in winter raintall over most ol" South n»nm
).
Australia,
The influence ol environmental temperature on sex
A weakness of the models is thai they do not take ratios is a mechanism by which climate change can
intoaccount all majorknown influencesonclimate.The directly affect thedistributionofreptiles. Specieswith
F.I Nino Southern Oscillation (FNSO; phenomenon is TDSD would be more susceptible to rapid climate
notcoupled with the majorjjiobql circulation models change than species without TDSD Webb & Smith
[Includingthe CS1R09 model) despite itsinfluenceon (1984) noted variation tn sex ratios of Crovothius
vaiiat'iluy of rainfall in Australia iWliertun <; al. in jt'hmttwi halchlmgs in the field, and speculated thai
prcsst fcNSO is of particular importance to die it was due to the interaction hetween TDSD and
Kcoiiimcnt of many .arid zone species |Austin & geographic differences in mean air tcjupcrarures in
Williams I9XH. Gractx el al. SWfe. The influence of ii warmerclim<uc.theeffect might be to skew- thesex
ENSO needs io be eonsidered in addition lo chamres ratio completely, blocking reproduction, and thus
predicteda>accompany ihcenhanced greenhouseeffect leading lo the local extinction of the specres. Such a
iWalkerer*//. I9&9) The issueis lurthercomplicated scctiarto has. been advanced by Pieau 11982) as a
beetiUse HNSO itself may be affected, [f increased possible reason lor the cxtinclion ol many \1c\o/nic
CO-, works to equalise lempcrauncs in the w&ttifiC of rep4iles
IMPI ICATIONS OK CI INTAIE CKANGJ* ON i! RRfSTRIAL VERTEBRA'!'!* B\l»NA <«l
Distribution of w.\(ut(me>\ Relator ufmmhnce ofmammals und rattle*
And Australia alteady has a more diverse and
TcsUidines, which require external sources ofheat abundant reptile fauna than arid areas in Noon
for metabolic activity, are limited in latitudinal America and Africa (Pianfca IMS?). In part, the
*Jisttibuuon by temperature However, mean annual diversity andabundance^t the reptiles isattrihuledto
temperature is far roncrudeameasuretopredict limits the high variabilityorrainfall which isa featurenl the
EOdistiiblltion. Lcnglhoithe growingpcfttfMfes been Australian inland fMorton & James 198$) Proposed
generally accepted 05 ft limiting factor, since al the changes inclimate might therefore heexpected tolead
latitudinal limits ol distribution adults may not have toan inctease inthediversity and abundanceofrepiilcs.
enoughtuncts) accumulate sufficient energy reserves relative to mammals.
|Ailitesrunravtiivveelyt,hediwsitnritbeurti(oMnnmcaCyulbteolcihmitAtedSbeyeothyea1b4i8l3i)t.y Not all mammals would necessarily be adversely
riff hatch!ings tp survive overwintering m the nest aafnfneucatled.proFdnuicttihveitylaragnedahreindceartehaes coafrrAyuisntgracliaap,acintcyl
t&IBermepBiertroeaontkbuasrceh(*1d/9«u8/r7/i.)oy1s9uXtg4hg;eeCsotmnagtthudaroiantaieoincartiOtfi1c9a8ol7v)a.,faOchwtobhruircidh> eaxnpnrueaslsedraaisnfallolial (bBiuormhaisdsgoer*v&eneMbrcaKtcens/iiscrelaIt9eSd9)t_o
However, the speciescomposition ol thelolal biomass
requires the accumulation of heal units over spring
sufficient torsuccessful reproduction. [suggestanother disistlrairbguetliyondoeftetrheminraeidnlalbly, rtahteherprtehdaicntaibtisliatmyounatn.d
possibilkv; thatdistribution i; limited bythe probability Patchy rainfall favours birds, bats, and mobile large
ofambicni temperatures being sufficiently highduring mammals such as kangaronv (Burhidgc &. .McKen/ie
the nesting season VQ permit the energy expenditure
1989) which ate physically capable of moving long
necessary in dieting the nest ehamher. an activit*. distances to environments made favourable by receni
reeoenisod as beingenergetically demanding (Con«dnu
& Galicn m**). heavy rains. Irregular rainfall kvuurs reptiles, which
have very low field metabolic rales relativetomammals
To predict theeffect thatclimatechange would have (Nagy 19871 andcansurvive forlong periodswithout
on a species requites detailed knowledge both of the food (Morion & James |983i. A change in rainfall
culture ofthechange inclimate andofthe mechanisms patterns to fewerdays with rain, a lower probability
by which climate change would affect a species tn oflight rainfall, and an increase in the frequency of
lestudmcs. we need In know which ot the abovc heavy rainfall would not haveagreatimpacton mobile
hypotheses is acceptable before we know whether iq animals, but would favour reptiles over small
-.Moviine flumbci of days between threshold mammals. Small mammalian eellulose-depctidcnc
temperatures. nadir winter temperatures, number of herbivores would beparticularly disadvantaged; they
spring days above sft temperature threshold, or the are vulnerable because their energy intake iv limned
probability of occurrencein springofspottenineniUires by Ibeu cutsi/e.Thisrepresentsa similarproportion
anovc a tJite>hold. ofthe sizeofthe individual as in larger animals, but
the energy expenditure for maintamance oi body
temperature must be relatively higher than for larger
Indirect effects mammals which have a lower surface area lo body
mass latio (Morton 1990V they are also limited by
teftectsmediatedby other factors-ire inherently more their restricted mobility in their ability to exploit a
complex than direct ^ffoets. Attempts at predicting patchy environment; and they are most vulnerable to
changes m animal distributions bused on detailed competition from rabbits (Burbulgc& McKvn/.ic 1989.
analyses of complex mechanisms winch include Morton 1990).
consideration of matters such as physiology, population It could be argued that Ihe balance between the
dynamics, inletspecific inteiactions, hchavinurjl diversity o\ reptilian andmammalian speciesasal the
changes, and nucrohubitui conditions arc fraught with time ofEuropean settlement was determined in some
potential tor error. An alternative is to identity a prior, more severe periodofaridity (suchperiods :uv
Miuiller suite ofinfluenceswhich drive thesystem and known from prehistoric tunes- see Singh I9KJ), and
determine the end result. N'ix 09H2) saw climate as theieloicthe balance would notbeaffccledby a lurifier
Ihemajordeterminant ofthedistributionofterrestrial increaseintemperatureand inthe variability ofrainfall.
urgaiusms. and several authors have used various Further,any mammalswhich might havebeen affected
climatic indicestoexplain thediversity and abundance are already extinct as a consequence of buropcan
ofparticular Australian hiol3. Ofparticular value are settlement. The counter argument is that effective
those lafe studies which corn-pate detailed dan on aridity in thefuturemaybemoreextremethan in recent
distribution and abundance with climate over a wide evolutionary time. Climaticaridity twherein increased
geographical range Both mechanisticanddeterministic evaporation is in excess ofincreased rainfall; may be
explohirions |o]|«w, compounded by "emulated aridity" because nt die
ta ? srorr
consequencesof the removal ofprimary productivity that there is summer rainfall. Both require a
from the arid system in the tbrm of livestock and heterogenous habitat with shelter being an important
livestock products [BujfWdgO & McKcii/ic NK'J), and component (see Htll 1981. Cannes tt al. WV1). The
rhe sequestration within the arid system of primary distributional data have been used by Walker (JWOt
productivity and nutrients in (he tissues of livestock todevelop an integrated modelling and mapping system
and rabbits. Thus the resources available to native which could be used to predict and map changes in
vertebrates would be sigmltcanlly diminished, distribution consequent to climatic change. Caughley
particularly during the resource "bottlenecks" of ct al. 0987) suggest that climate change in the past
droughts, and in drought refuges (see Mortem 1990). bus influenced the distribution of mucropods: it is
therefore reasonableto use theirconclusionsto predid
Range changes oj kangaroos the distributional responses of these three species* lu
-
Theresponsesofkangarooscan perhaps be predicted future climate change.
with a Iillle more confidence than those ol other If. as predicted (above), the winter ramtall zone
vertcbiates. Many studies of kangaroos have been contracts to the south and temperatures rise, the
undertaken, including thorough studies of their distribution ofthe Western Grey Kangaroo would also
distributions (Fig. la) aspartofthe basis for managing contract to the south (Fig. lb). Perhaps more
populations which are commercially harvested. remarkable might bechangestothedistribution ofthe
Caughley tt al. (IW7) have demonstrated that the EasternGrey Kangaroo. Atpresentthe speciesoccupies
m
distribution of three species of kangaroos is. the two small and widely separated areas in South
major pan. determined by elttnaie. Win1st the Australia, but theseare minorprojectionsinto tinsState
distributions are directly determined by land use and ofa distribution whose western boundary runsalony,
the availability of food, water, and shelter, these or jum to the east of the States eastern borders
attributesareinturngreatly influenced by climate. The (Caughley et al 1984). M. gigantcas could extend a
distributions ofthe two grey kangaroos, the Eastern considerable distance to the west of its present
Grey Kangaroo (Marmpusgiganfcit\) and (he Western distribution, and hence across the north of South
Grey Kangaroo(A/.JhliyjinoMt.s), arecloselyassociated Australia to occupy suitableMhabitats in the northern
with the seasonality ofrainfall; they overlap in areas pari ol the present range ol Jultgino.su.\. il ceitutu
nfuniform seasonality of rainfall, but M. i>igartifu.\ conditionsarcmet. Theyarethat summerrainbecomes
occurs in areas where summer rainfall predominates, morecommon innorlhcrnSouth Australia, that reliable
andfa',faliginosasoccurs inareas wherewinter rainfall water is provided by increased frequency oi' heavy
predominates. The Eastern Grey tolerates higher raudall and/or livestock watersupplies, and that habitat
seasonal temperatures thantheWesternGrey provided heterogeneity persists m the face ofclimate Lhnnyr
\J*2-r/-1"-*
M
gigantet;s
(a) Presentdistributionsolgreykangaroos (b) Possible future distributions ofgrey kangaroos
(Density > 0,1 persquare Km,)
Pigf,ttIit.miHWPrIet&,eiCvdid.ifscthrticbyut&ionCsiroifaggre(IyvSklajn,gaCraouotish.lt<rbyl PHosastiblteN8fu3t)u.reCdjinsjt-rhitbcuytifotnsaotl g0r9e8y4)k,an.aSghuorooosvt.nAlda(pWtKe.d4)b.ttmC&enCi
iMPMCAnONSOrCIUVTAlK HANtil* ON IERRFSTRIAI VKklFHRAI> rAl MA
I fcl
BbWftl/ >>fnthhir< in tifitl mvm variahihly perennial plants would DC favoured tf^st
Inand Australia, the r.umpc.in RabbitOr\vtotoi;u* ephemeral plants(Stafford Smith &. Nonon 1990). so
i i^i/i i////y isdecreasing die probability of survival of rabbit populations could be expected to become le&ss
CTiiflM perennial plums duringdnitighu and having a dense unless palatable perennial gravies Mich
profound effect on trie reciuitmcnl of sonic spevtes, Ihettwda sp replaced annual uruvsoand lorbs. Further
sin idem in unictoeliminatethem from foe landscape explorationol thissccnaito would need totakeaccount
U.aiilge it Graham $83: COWC 19K7) The response ofa potentnil southernextension.i( thedominance oi
ofthe rabbit toelmrahJ change isthereforeofparticular Cj, over C^ grasses (Henderson c*f i>/ in pres<), the
importance relative importance ofC and C grasses |o rabbils.
}
Historicalrecordsshowthai rubbiis inandareashave and Ihe miplit'auons for rahbits of a change in the
been severely reduced inlumbersduring pasidmughb seasonal distribution ofr<tin awards summer rainfall
(Griffin & Friedcl WN.V and may under divught in arid area* of South Australia.
conditions, becomeoxt'meiover largeareas iMvcrsA
barker r»75i. kceruitmont is most unlikely under J-oiiJ laurih jor Spinifcx Hopping M<uor
draught conditions (King / af\ WN3). and prolonged .Vorowy* ii/<*v/V. the Spimfcx Hopping Mouse, is
dnnishls such us Ihc fciSCtf and J92.V38 droughts widely disinbttted throti^h sandy area-, ol uonhcrri
ai Alice Springs (Griffin Si Friedcl |VKS| km ihe South Australia, mainly in association with spinfffcx
h»n«eviiv of ihc species (see e.g. Dunsiuorc 1974). grasses (Walls & Aslin fVSll A major cornponent oi
hven so. a lew rabbits survive in rehires The quality its diet ts seed (Finluyson 1940). and Irom the
ofthe refuges is determined by (hen ability W harvest carboliydrate in seed it is able to denvc sufficient
and siorewater and nutrients flowing from largerarras metabolic waterto ^ur\ive indefinitely; one female is
ofthe landscape(Morton 1990) such thatrun-on horn known to have reared a single young without
ligln ramlall is sulficicnltostinUiiutesomeplantgrowth supplementary water (Havcisiock&NVatts W5) Hence
during thedrought penud (l.udwig 1987}. AtWilehitie. ihe regularity of seed produciion would inOuencc
South Austrulia. CookefJ982i noted thai a sharp (all suivivahiht) of,V. tttefe in northern areasofthe State-
ofliule morethan Stum 01 tain might he sufficient to Seed produciion may be influenced b>' soil moisluivx
yield run-off which, if concentrated along drainage lenipcraiure. and CCK levels, and two mechanisms by
lines, wouldensure ilutl succulent food in the formof which N. tj/r.\is miglu be advantaged are exploied
chenopod shrubs would heavailable to rabbits living Soil n>oisture is one of Ihe mosl iiiipoitani
01 warrensalongthosedrainage lines Onceheavy raui climatically determined variables for grasslands
lalls (>2G mm near Carnarvon, Western Australia. [PiltOCfc L993) and he(K«- lor species such ns S. itkxis
King tt ul, WH3). rabbits begin to breed, and cUn whichdepend inlaree mcasuieongrasses (MucMillan
expand from therefuges to reccilonise the bulkfit the Sl Lee 1969;.Walker*ial CWKM| anticipate that mean
Itindscnpe. soil moisiure is likely to diminish in northern South
Heavy rainfallsJre rare inthe Austialiuu and /one. Australia, although at present the reliability of
whole years may pass without a rainfall event ^ predictions isquestioned (Vinnikov W\\ Pillock 1993),
12.5 mm (Slaltord Smith& Morion WW). I.i^ht falls More important for seed produeliou aie episodes of
arcmorecommon, hut ithasbeenpredicted(seca>nei highei soil moisiurefollowing heavier rainfalls, which
thattheheavy falls wouldbecome morecommon, and arcpredicted (above) lo become more rrcqucm. Thus
light falls somewhat less eununon Hence, drought? pulsesofseed .->uppl> may become p>orefrequent,and
ure likely to be shorter in duration, but the refuges support denser populations ol .V. t.r/f.u>.
which sustain the residual rabbit population during The rate ol' ^nnvih and ihe speed at which seed
dnmghls would he k httle tes** reliable. With a development occurs following rains may be
coincident rise inlenipcraiureexacerbatingthesevere accelerated. Imai tiul, (|985iobserved increased Seed
physiological stress experienced bv rabbits under vield per plant for noe gmwn under enhanced
present summer conditions iHayward 1061;. local greviihou&cconditions. Giflord (1970. I9K8) predicted
extinctionbecomesmore likely duringdroughts, hut. thai wheatyields inareas with morestrongly seasonal
v\i|h decreased return times lor heavy ram. plagues rainfall would increase as a lesult ol the enhanced
iiug.nl becxpvvicd more fi'cquenilv in thoseareaswhere "gieenuouse"effect, andthat somegrain growth would
rabbits survive become possible under conditions of dtfidiiy which
Vegeurion changes consequent Id lite increased cuTrenily preclude any yield. The faelor influencing
variability of precipitation might not favour rabhils. yjetd wasstimulation toplant growth by boih incicased
Rabbiis are found in chenopod sfU'liblands, but the CCV. and warmei temperamres, which would result in
majority of the teed is provided by Hie short grasses ashortergrov*incseason such thatthe giam wasmore
and Sorbsbetweenthe shrubs; Ihechenopodsan eaten likely to he filling under a favourable soil moistuie
only during droughts (Hall it «/. 1964; Gnfliu & regime. Ifnhanced efficiency 9fwater use would also
Inedel WIS), Underconditionsrifincreased wlinialic occur vluetopartial stomatal closure (ChavesSiPercim
M
K Mori
1992V Thus droughts as perceived by AA al&is may influences. Should ihe donate change, the
beameliorated by the supply <>fsomeseed when none accomodation between the host and the paravilc may
would be otherwise bt available. be disturbed-
Australian arid-?one \oils are generally infertile Amongsttheparasiteserflivestockthereareexamples
(Morton i99to. find nutrient lltnlt&fl la) •.-ounter ot species whose transtnissibility is known to be
growth-stiniuUling mechanisms Although seed affected by climate. The larvae ot Hacmonehus
produciiou may be limned by the availability erf tr>/uvv-wv,jgjAint-iniesiinal parasiteofsheep, requiTc
phosphorus, nitrogen is less hkcl) lo be limiting lot mean temperatures >|S°C for normal development,
C. plants* hot a likely corollary is that the protein whereas the development C/fOtfertagfa circtuncincta
content of their seeds would be lower |see Conwy larvae is suppressed above I5.5°C. As both require
1992!. A/, a/cu'v would not be disadvantaged, &1 it is moisture, the formeris an(organismofsummerrainroll
more likely to survive drought on low protein diets uica-s. and the latterofwinter rainfall aieas (Southcotl
which obviate the need to expend water todisposeof N al 1976),
waste nitrogen (MacMillan & lee 1969)- There isless knownabouttheparasitesofAustralian
narive vertebrates, and most of the published
( tMiftt'Hlion between Ctcnolus species investigations havebeen Laxonomic (v g Reveridge&
Manyofthe 70species intheseineid genusCromr/cs Durene-Dcsset I992|. Arundeletat (1990)undertook
9ft> associated with spinilex (Cogger 19*)?). and it is one or the lew epidemiological studies, which
HOI unusual lor several species to be synu-pic, demonstrated that helminth parasites can cause
suggesting fine niche separation between Ihem. II considerable mortality inEasternGrey Kangaroos,and
eJimaie influence*, the niche sepatation. climate change concluded that development of frec-hvinc larvae ls
may affect rhe hulancc between the species influenced by climate In North America, the Moose
Chtunnsheletmrand C. pantherinusare twospecies Ahc.s ahe.s can C*.ist synipatncally with White-tailed
occurring synipatrically in the lar noiih-eastofSouth Deer (klocoitens viryiniamtsonly inthoseareas when*
Australia: Pianka(1969) noted that they shared similar circumstances do nor favour persistence ol infective
niches, and suggested that C. pamhenm<s would be larvae ol the. meningeal worm Purelaj)hostn)ni>\lu\
excluded by C helcnue but tor iu reproductive tenuis (Gilbert 19421. Hence, climate change may
capacity, lames (J99ia) (bundahighdegreeofdietary indirectly inlluencj the distributions of terrestrial
overlapbetween the two species,andnotedthatdietary vertebrates through its effect on the probability of
overlapinQtnotits washighestduringthedriestperiod disease transmission.
of his study. There is evidence to suggest that Ihe
separation betweenthese twospecies is raisedonthen Mechanisms for coping with elimatc change
thermal responses: Pianka(N8<S» found C. pantherinus
C
to h;oe <i lower mean body temperature than FVcssmghaoi (W*)3) recognised that (here are ihn-e
hefetuie. and .fames (1991b) speculated thai C. means by which aspeuo mightsurvive dimutechange
pantherinus tand C. brooksi) can be active at winter range change to track shifting clirwitc ?oncs.
Icinperaturcs which preclude activity by £ IwIvmic tolerance of the change, and/or microevolutionary
C
(and other Ctetwtus species), This permits change F.xarnples are presented which demonstrate
pantherinus and C hroftksi to begin reproduction that lolerancv in the form ot behavioural plasticity may
earlier than the olbei species. Jfactivity at different counter climate change, but tracking appears
temperatures Is critical either for maintaining stable implausible for many small terrestrial species
niche separation or tor sustaining a mechanism oi'
oscillating disequilibrium between the specie*, an Ttdcnwcc
increase jn temperature during winter may /csull in The Red Fox {Vulpcs wipes) has a complex social
C
competitiveexclusionol C.pantherinusby hvlenae siruciure which can be modified to cope with
in those areas where Ihey are sympalric environmental change. Zabel & Taggart (1989) have
demonstrate*.! an effect by the FJ Nino phenomenon
Bpicltrmhfio^y on the food supply ot a population oftoxes on Round
Instableecosystems, there is generally asignificant Island, Alaska. Increased water temperatures in the
level ofaccommodation between hosi population*and Bering Sea were associated with widespread nesting
disease-causing agents, particularly if they have co- failurein the.seahjrd species which comprise most ot
cvoJved. However, tiansnussion ol infectious disease the summer diet of the foxex Resorption and
isadynamic process, and in many cases isdependent prcimplanlauon Joss arc known to occur in pregnant
onihecapacity ofthe infectiousagent tosurviveoutside vixens (Ryan 1976). a common cause of which ts
ol tlic host. Helminth parasites oltcn have obiigatoiy nutritional stress(sec Moustgaard 1969). Hence, il the
larval stages which may sursive for long periods on available tood was uniformly distributed amongst the
the ground, and thus be susceptible to climatic foxes, total rcpttidnctive failure m the fox population
JMW U AlifiYS O! C[ IMATP. CHAVGF. ON TERRESTRIAL VERTEBRATE FAUNA 65
may well have occurred However, on Round Island, nomadic (VVVcns 199]|, tracking climate change is
iliftary change^ m smaller, less common, and less physically and behaviourally possible. However,
accessibleseabirdspecies wereassociated withchanges >cdcntaty species may have behavioural difficulties
in »hc social viruciure of the foxes. Polygyny, the Few datll are jvatUhle on the dispersal capability ol
trpioduetive mode prioi to the dietary change, wns small lerrcstnal wrtcbrato, but the longestdispersal
supplantedby monogamy. The tnWShelpi\essential movement recordedby James (I99le) forany individual
loi euptuiuiganddelivering prcv toaIncusing female Offive species ofCrctioms xvas 605m. indicating thai
and her liiler (Klennan 1977): thus ihe change in the uuasMsted dispersal is most unlikel) lobeablelotrack
social siructuic meantthatassistance provided by The ehniaie change ai the predicted rate.
mule lit\ was 1ocu>sed on fewercubs ai a time when Possingham (1993) points out that comparisons of
it would have been more difficult tor the males 10 dispersalcapahilily with dieraleofclimate changemay
pnxure SmkI Individual reproductivepujecess.(in terms be simplistic. The comparison is useful to identify
of cubs reared to sexual maturity) of the reduced species which are physically incapable o1 tracking
nuiilher of breeding lemalo was noi significantly climate, but cannot hy itself determine competence.
affected hy the LJ Nino phenomenon. Hence, a There must be suhH^uenl stages in the process ol
llcmpurjry ) climate change which lead to total identifying species at mk, such as consideration, of
icpiuducnve failure of the two seabird speciev nmu physical hanier> and iiitcr-relationships between
prominent in the did.ol ihe foxes did not in turn lead species Even ifCrwotus spp. were physicallycapable
to Ma! reproductive failure in the tbxes. of tracking climate change, Ihcrc is a close and
The Eastern Long necked tortoise xChckulhuk presumably obligatory association between many
l<mxnvlli*\ has simpler behavioural patterns than ihe speciesofGwiorVM and Iriotlhi and PIvein;*line '-pp.
Red Fox. hut still has \ome plaMiciiy. Ii appear*, to (spinitex grasses) and their attendant termites. These
adjust its selection ol nesoug sites to Like account of are primarily dislTihuted in infertile, sandy soils
meteorological parameters likely to affect incubation (GraeU ft at. I9S&). which indicates that sprnifex-
temperature. At Armidale. New South Wales, the dependentspeciesol erenow* which me ievstolerant
speciesdigs nestingchambers in unshadedareas, whicti oi increased temperatures would be unable to track
increases insolation and hence egg lempeTaiurc, and climatechangeacmsselunges insoil renihty andtype.
sltonen.N iiicubaiion (Parmcntcr 1976). The same 1-orvertebrate*- inthenorth ofSouth Australia, tracking
speciesat Rnsewonhy, S«»uth Australia, digsabout two- temperature change> means a generally soudiern
ihinJs of its nests in sites shaded for more than half extension in range twith or without a northern
ofthe day (Sloll 1987, 1988). Nest temperatures were contraction, aseparate issue which would depend on
not recorded ateithersite,hut meandaily temperatures theupper limitsoftolerance), but forC pdMhttinm
during the incubationperiod aie highet at Rosewoithv there would be constraints because long distance
iX^°C higher in January) and cloud coveT is les> dispersal or even local spread of Tri(ial& and
frequent (0.7 oktas less inJanuary)- Thompson(!9SSbl PteanuUiH' seem lit requireconsiderable tunc (Jacobs
has demonstrated thai unshaded nests of Etmlnut B82J. Also, thesegrasseswould notextend imoheavy
ftktcquariiat ftnrmcra* 5?outh Australia can he 2h°C clay soils and limestone plains, die laiici being
warmer than shaded nests, and attributed deaths in geueially south of the present distribution of C
some unshaded nests to excessive heal. Thus it is ptmthvrirjut;
C
reasonable to speculate that /v>wv>o//ia. like the The stccpnes5ofclimaticgradient hi moui'.taittOUs
species of testudines considered Isy Bull etat. (I982J areas is much greater than on plains, and thus
and Schwarzkopf& Brooks 11987). positions its nests alUtudinid tracking of climate elwngr. is much more
relative in shade to ohuin opiimum subsuilace feasible than latitudinal tracking tor small leiresLnal
temneratuies fen incubation. vertebrates. Generally, a short chmb in altitude
corresponds to a maior shdl in latitude 'Peters A:
IWluig 198?). Overa distanceofabout 15 km in die
Dorale ct til. (1992) dated laic Hiilotcne vcgcfctliou Adelaide Ihlls, a 5G0m increaseinaltitudeisassociated
chances at two sites Li Towa which correspond in a with a fall in January mean maximum temperature of
rale of reireal ol prairie of 300-h00m pei annum. about5°C. andarise inmeanannual rdinridlofabout
Huwevct. iheanticipated t'atcofanthropogenicclimate 600 mm, However, whilst the climate as ntie
change is much greater than in the past (FVissutgham component oi a species" environment may track upa
I993) Wiih the tow relief c>f the inland plains of mountain, other compuneiils of thee-nvmmmenl may
m»nhernSouthAustralia, meanthermal giadientsare be fixed; forexample, substratestructure and futility
Slight, and a typical distance between isotbems usually change with altitude
corresponding to the predicted annual rare of To allow Hacking requires the linking of aieas
temperature change of 0.03°C is 20(H) m Because managed primarily tor conservation along latitudinal
unnvy bird species oi the Australian arid /one are and alliludiual gradients (Norton I°u0t. Ihe teviesv
A
6fT P STOTT
or nature rexcrvex in souih-ensiern New South Wales interspecific relationships ofAustralian vertebrates is
uiiJcriuis.cn hy Macules& Stein (#89)confirmed that limited (Norton 1990), but there is suflicienl
a single, todgt mirmw. rectangular reserve aligned informationin indicatethat climatic influenceson the
..'Ic-nujn ultituiJinul gradient would be theconfiguration distribution or many animals operate through
which would most patsinionionsi) ivieet ihedominant mechanisms which are subtle and as \ct poorly
environmental requirements ol temperature, rainfall, understood, and sufficient information 1o wurrnm a
and substrain for 26 canopy tree species which occur conclusion that climate change of any magnitude is
in the region: Macke) ei at. (IsWX) note ihc necessity quirelikely toafleet thedistributionsofmany species
ttnd Hguc the validity of using vegetation daw us ol terrestrial vertebrates.
surrogates for data on fauna habitat in the present Conclusionsabout the fateofindividual speciesarv
circumstancesorpaucity ofthepreferred primary daia. at present speculative. Deterministic studies such as
and advocate u Incus on ecological gradients in oaler those undertaken on kangaroo peculations &xe less
i" poiviije a tnaryin flf safety in assessing areas for speculative than mechanistic studies hcvausc of the
conservation value HosvtYer. because ofIhe lowreliel complexityofthe means by which ehmalo influences
ofmuch ol South Ausirnlia. only latitudinalgradients the biota, but mechanistic examinations arc
arc po.ssihlc in most areas. complementary inthutthey nu\ rcveaJcriticalaspects
ofdetail notapparenttodeterministicconsiderations
CfWClusion
Ackuowledgmettf-s
Thepaperhas loeusedono numberofMinties which
have relevance to the issue of climate change On ThanksareextendedtoSusanCanhcw whoprovided
reviewing Ihe topic, it isapparent ihax fhe zoological constructive criticism of the manuscript fti Megan
hase from which changes may be predicted is Lewis whoprepared lor presentationand publication
imperfect. Reliable information on the present the figures on the distribution of kangaroos, and to
distribution, abundance, population dynamics, and KathleenWilliamswhocared tor thetempleofthesoul
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GorAdoMn, H£:.HBv.uWOHtekr,njWN.KP IfIffVP?r)iSmiem*u.laAtedB.c.h,Fiim/.a'*e»si^Ii,n Tsoeimlpteurrtaltesu*reJndterapsepnedceinfitcsveaxridaettieornmiinnaCtuiroentrtiin QttuWie'eftnisi.hmpipl
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Melbourne) pick'beilii (Gmv). from snuthcrnSaskaichewjin. CWw ./.
DRIf-r-i^. 0. K A FRIED©.; M. H. <W85i ObOOptliAOuS Z/«./ 61 1449-1509
eLeh-atlnieitefiuiinleeesmernaNl fAaurMrlaanldu:niSaonamueemiemnptlic.a/tiMounl* oflfrlmffarjifofr Ma<MAKCtIVv.UhBU-K,G..1, RNt&>.1'HL.tMisKo,. PH,lMT.CHI(TN9sKo8Si.A^iVIessiIn-y
ft 63-Wl, represenia)ivene\\ cifplates futcmiscrvaiton tt->ervation
MH.aTm^riho.spimcGaa.lAdR.i.u.1uSrP(EKMTwHmiTv*sinRn.ni.aIOiCM^F.R>&CRlriSm/iaiitNenO.DHx'.nC-CHM,A24/ri-l11Q%5W47)); Mva4ocf0n3MdA-uu5hM*1en4rilftciaNagienX,lhioRsptipn&ign.gIErmnri-y.cten^m.CKo-Mmrp(f-/NttaSBgVtf)oicWha1tt2m.e*r50Fnk1it-ea5fy1yh4tit,di2s8m.
(KtRRvoeewns^aedchiniovcne.ro.anlsIilo9Son2.l^o-(f1wW9aft6t>h2lee.)rvTAerhgueic\st(iLanaicbJitioilnto8intmy>.ri,trCKf1So2(oI.liuHea2Oinra.nbSoHb'-rit2ell6d4lVW,cti/Muci'uirj.lviio6w,n Maci(MoEnend,l.<b.coiA*r.u^\rGaRnri.eet)eoI.nnlh9io8n8uisAeCusltiLr*matalntinaeoipcnghpainu3gr6ed1i-ai3nt7du4iiotksicirBmMrpngaaccr*tm*aon(n.CSntaItkRuOrL.e
Ht'1M6'0LK17M5i.^, S.. HAI(hHM.tV. P., Viin CAf.MMbKhK, 5. & MAdRitstariib.ru-ts,ionCsofRipe&vieSstunnsd,|he).>jeLl.et.(1i1o9t8i9o1f Pnaaltofieemfse^ienrvtehse
Osmund, B. finpress)AreC 4pathway plants threatened Anexample fromEiuahprui, Ibresis inscnilh-eaiderriNcv>
In glt»hnl climnie change? fa Hvopbysiolcipy nf StiiiTh Wale. Hm( O.mvr. SO, 2I9 23H
pVhe.o-tLoisevmCliiteesdisin1(PK)IETeTolOoC^'Ki,eaJAstBu.diielxWlHe.idAelcbleirmga.teSpcnhnagncejt Mimaohtuonsd,an.Sc.oRo.fahvnadrJdsamilnsa.ndC.AuDs.tr0al9i8a8:'ATbr^eipdpirvauirsxailtyAamm.i
l>er>pectiveon urxssliindx }*Ttu XVtlhst Gn.Wii Cdltgr- N,u 132. 23? 25K .
Hill C. J. r!98L) OiVtrihutmn of the grey kangaroo* in verteh(r1d9l9e0a)nTihmeaKiomlpaaictjd*A.<ufsttruailoipacaAncsocntii-ireipntcu-ailt m'«or.deiln.
south, inland Queensland. Ai*a. fo/nge. 1 3. 5R-<i6. Pr<K. t'cvl. Sot: Ausl 16. 201-213.
H'»HouWi2fi)t)N*,Cl.i1.maTi.e.cChaainigeas1d9l9r2, TBh.eAsup&pleVmaern.iisiun, ISf,tpKO,Q MorrevpirGodAuALRtiio)n., pJp.. 4(R199659l)tiMNuCtorliet.ivHe Hr.nl&UlBCfuKpBpls.uP.poTn.
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?nt«. Cambridge). (Academic Precis. New VorV).
)
f* P. SIOPJ
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dSemx ratiosoftwospeciesol sealurtWnestinginSurinamc. seasonal pataeocliuiatesot Lake Frotne. South Australui
.1 Z«>i hi, 2227 223V. ttKiUfhtoli^m 82, 419-430,
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(1985) Some intercontinental comparisons(il desert J. 11989) GlobalclirnatCchangeand Australia: Kfleeinon
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