Thermal Biology of the Emu, Dromaius Novaehollandiae
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UNIVERSITY OF NEW SOUTH WALES Thesis/Project Report Sheet SumameorFamilyname: ........... Jfl,.i;l].oueY, .............................................................................................................................................................. First name: ........9.~~ ................................................................. Othername/s: ...... J);~Y.ID................................................................................ Abbre vtation.or· · & d egreeasgtvenm· · theU ruvemtyca· · I e nd ar: ...................................Ph.D. ............................................................................................................ Schoo); .......f\i.gJgg;hg,~J.. ...?.s;;h~n9.~ .................................... Faculty: .. J?.Jg,lgg:i,g.~.l .. liiDR. .. ~.. .a:v.iour.al .. Sr::iences..... TI&: .•.. ....•.• ~~.~~ .•.~~.<?..~~ ...9-:f. ..."!:h~ ... ~ ....(Dranaiu s novaehollandi ae)......................................................... .. .................................................................................................................................................................................................................................... Abstract 350 words maximum: (PLEASE TYPE) The emu, Ort?mBillS MI·'*Mllendiae, is a large ( 40 kg) flightless bird that inhabits areas as diverse in temperature regimes as the sno'w' country of the Great Dividing Range and the arid interior of the Australian continent. Several aspects of the emu's physiology suggest it is geared to life in the heat. The basal metabolic rate ( BMR) is lo'w'er than expected for a bird of its size. This may be a phylogenetic attribute, but it helps life in the heat by reducing energy needs and the endogenous heat load . The BMR and body temperature of resting males are lo'w'er than that of resting females . This may be a result of selection operating to reduce the energy needs of the male during his eight 'w'eek incubation fast in 'w'inter. In hot conditions the emu maintains thermal balance by increasing evaporative 'w'ater loss, about 30% of'w'hich is cutaneous in origin. It avoids respiratory alkalosis 'w'hen panting by reducing tidal volume under mild heat stress. It is facilitated in its ability to remain active on hot, sunny days by the structural and optical properties of the plumage . less than 10% of solar radiation incident on the emu acts as a heat load at skin level. When 'w'i nd speed increases to 6 m/s the heat load at skin level is reduced to less than 1.5% of solar radiation. The heat load from radiation may be reduced further by the presence of a sub-cutaneous fat layer on the back in summer. When 'w'ater intake is restricted, osmoregulation is facilitated by a reduction i n the normal evaporative response to heat challenge. life in the co 1d is facilitated by the emu 's ability to i nc rease its oxygen extraction rate at 1O'w' ambient temperatures (T a), and by recuperative heat exchange in the nasal turbinates, 'w'hich allo'w's the emu to exhale air considerably cooler than body temperature at lo'w' Tas . These t'w'o factors acting in concert amount to savings of up to 20% of the energy consumption required to maintain body temperature at lo'w' ambient temperatures.· Declaration relating to disposition of project report/thesis I am fully aware of the policy of the University relating to the retention and use ofhigherdegreeprojectreports and theses. namely that the University retains the copies submitted for examination and is free to allow them to be consulted or borrowed. Subject to the provisions of the Copyright Act 1968, the University may issue a project report or thesis in whole orin part, in photostate ormicrofilrn orothercopyingmedium. I~r~=T~,~~=~~~·t.•oct::~r:::•motioo&~::':~w1~:'~'. m m The University recognises that there maybe exceptional circum tances requiring restrictions on copying or conditions on use. Requestsforrestriction for a period of up to 2 years must be made in writing to the Registrar. Reques oralongerperiod of restriction may be considered in exceptional circumstances ifa ccompanied by a Jetter of support from the Supervisor or Head of School. Such requests must be submitted with the thesis/project report. FOR OFFICE USE ONLY DateofcompletionofrequirementsforAward: /'S. I I · (/'!, I I s ReKistrara!llDeputyPrincipal TillS SHEET IS TO BE GLUED TO THE INSIDE FRONT COVER OF THE THESIS MBT 598.53041 1 Thermal Biology of the Emu (Dromaius novaehollandiae) by Shane Kevin Maloney Thesis submitted for the degree of Doctor of Philosophy to the University of New South Wales, Australia July 1993 UNIVERSITY OF N.S.W. 2 o m::c 1993 LIBRARIES UNNERSITY OF NEW SOUTH WALES Thesis/Project Report Sheet SumameorFamilyname: ............ ~.ialQUe.¥, ............................................................................................................................................................. First name: ........~~~ ................................................................. Othername/s: ...... J$§Y..;i;!.L .............................................................................. Abbreviationfordegreeas given in the University calendar: ......~!':.:.!?..~ ............................................................................................................................ School: .......l?.~g.:J.gg;hg§:f.. ... P.f1:1W.9.~ .................................... Faculty: .. J2• .:i.glgg;;hg,?,.,l ... ?aP.Q, .. J?.OO~Y.iQW:.9..l .. SQieuc.es..... nt1e: ...........~~.~~ ...~~S?.~9.9.¥. ...9.t ..t.h~ ... ~ ....mt.8U§i~~. noy~ehollancliaeL ....................................................... Abstract 350 words maximum: (PLEASE TYPE) The emu~ f)rQIMiU$ MI•'&?M/1Bndi8tl1 is a large ( 40 kg) flightless bird that inhabits areas as diverse in temperature regimes as the snow country of the Great Dividing Range and the arid interior of the Australian continent. Several aspects of the emu~s physiology suggest it is geared to life in the heat. The basal metabolic rate ( BMR) is lower than expected for a bird of its si2e. This may be a phylogenetic attribute, but it helps life in the heat by reducing energy needs and the endogenous heat load. The BMR and body temperature of resting males are lower than that of resting females. This may be a result of selection operating to reduce the energy needs of the male during his eight week i ncu bati on fast in winter. In hot conditions the emu maintains thermal balance by increasing evaporative water loss, about 30% of which is cutaneous in origin. It avoids respiratory alkalosis when panting by reducing tidal volume under mild heat stress. It is facilitated in its ability to remain active on hot, sunny days by the structural and optical properties of the plumage. Less than 1 0% of solar radiation incident on the emu acts as a heat load at skin level. When wind speed increases to 6 m/s the heat load at skin level is reduced to less than 1.5% of solar radiation. The heat load from radiation may be reduced further by the presence of a sub-cutaneous fat layer on the back in summer. When water intake is restricted, osmoregulation is facilitated by a reduction in the normal evaporative response to heat challenge. Life in the cold is facilitated by the emu's ability to increase its oxygen extraction rate·at low ambient temperatures (T a), and by recuperative heat exchange in the nasal turbinates, whtch allows the emu to exhale air considerably cooler than body temperature at low Tas. These two factors acting in concert amount to savings of up to 20% of the energy consumption required to maintain body temperature at low ambient temperatures. Declaration relating to disposition of project report/thesis I am fully aware of the policy of the University relating to the retention and use ofhigberdegree project reports and theses, namely that the University retains the copies submitted for examination andls free to allow them to be consulted or borrowed. Subject to the provisions ofthe Copyright Act 1968, the Universitymay issue a projectreportorthesis in whole orin part, in photostate ormicrofilm or other copying medium. ''""l~JJnl-icyMlaofilm•of o3S04nJ•""'"'inDh~"'lionA""""'"'"""tlonol(opp1Woblo "'""'J•••only), ......... : "...... : .. :;:: .:.... !.............. ... 1 I 1............. w,;;;;; ......... !................. .. ...........?.!2/I:;.~:;; ....................... The Universityrecognises that there may be exceptionalcircumftances requiring restrictions on copying or conditions on use, Requests for restriction for a period of up to 2 years must be made in writing to the Registrar. Reque;tJ;ralongerperiod ofrestriction may be considered in exceptional circumstances ifaccompanied by aletter ofsupport from the Supervisor or Head ofSchool. Such requests must be submitted with the thesis/project report. FOR OFFlCE USE ONLY DateofcompletionofrequirementsforAward: /J. (I· li'S Ige~straranaiJCuty JPri.l nCJpn THIS SHBBTIS TO BEOLUBDTOTHBINSIDBFRONT COVBROFTHBTHBSIS 2 Table of Contents TABLE OF CONTENTS 2 LIST OF FIGURES 7 LIST OF TABLES 12 DECLARATION OF ORIGINALITY 1 3 ACKNO~DGEMENTS 14 ABSTRACT 16 Chapter One: Thermoregulation in Birds: a Review 1 8 1.1 What is Thermoregulation? 1 8 1.2 Why Thermoregulate? 1 9 1.3 How is Thermoregulation Achieved? 21 1.4 Metabolism 22 1.5 Insulation 27 1.6 The Respiratory System 3 2 1.7 In the Cold 38 1.8 In the Heat 41 1.9 Control 4 6 1.10 Dehydration 4 8 1.11 Brain Temperature 48 1.12 The Case in Point, The Emu 50 Chapter 2: Basal Metabolism and Body Temperature 53 2.1 Introduction