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Chronoethological assessment of well-being and husbandry in captive koalas, Phascolarctos cinereus, Goldfuss 1817 [Elektronische Ressource] / von Annette Rabea Benesch

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Chronoethologicalassessmentofwell-beingandhusbandryincaptivekoalasPhascolarctoscinereus,Goldfuss1817DissertationzurErlangungdesDoktorgradesderNaturwissenschaftenvorgelegtbeimFachbereichBiowissenschaftenderJohann-Wolfgang-Goethe-UniversitätinFrankfurtamMainvonAnnetteRabeaBeneschausGroß-UmstadtFrankfurt,Februar2007vomFachbereichFB15BiowissenschaftenderJohann-Wolfgang-Goethe-UniversitätalsDissertationangenommenDekan:..............................................................................Gutachter:...........................................................................DatumderDisputation:.............................................................Formymother,AndreaBeneschIn the heat of the day in an average zoo,What's a koala most likely to do?In the fork of a tree, in a featureless heap.It closes its eyes and endevours to sleep.And the visiting children and parents, they bothSay, "Oh, what a lazy marsupial sloth!It's far to lethargic to romp or to toil,Perhaps it is drunk on the eukalypt oil."But that isn't fair; it's improper to mockA creature that works on a different clock.The schedule koalas are destined to keepIs to deal with their business while we are asleep.And, up in the treetops, throughout the long night,They climb and they feed and make love, or they fight.And if they took notice of me or of you,They'd probably think we're a sluggardly crew.

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Published 01 January 2007
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Chronoethologicalassessmentof
well-beingandhusbandryincaptivekoalas
Phascolarctoscinereus,Goldfuss1817
Dissertation
zurErlangungdesDoktorgrades
derNaturwissenschaften
vorgelegtbeimFachbereichBiowissenschaften
derJohann-Wolfgang-Goethe-Universität
inFrankfurtamMain
von
AnnetteRabeaBenesch
ausGroß-Umstadt
Frankfurt,Februar2007vomFachbereichFB15Biowissenschaftender
Johann-Wolfgang-Goethe-UniversitätalsDissertationangenommen
Dekan:..............................................................................
Gutachter:...........................................................................
DatumderDisputation:.............................................................Formymother,AndreaBeneschIn the heat of the day in an average zoo,
What's a koala most likely to do?
In the fork of a tree, in a featureless heap.
It closes its eyes and endevours to sleep.
And the visiting children and parents, they both
Say, "Oh, what a lazy marsupial sloth!
It's far to lethargic to romp or to toil,
Perhaps it is drunk on the eukalypt oil."
But that isn't fair; it's improper to mock
A creature that works on a different clock.
The schedule koalas are destined to keep
Is to deal with their business while we are asleep.
And, up in the treetops, throughout the long night,
They climb and they feed and make love, or they fight.
And if they took notice of me or of you,
They'd probably think we're a sluggardly crew.
Ronald Strahan, "The Incomplete Book
of Australian Mammals”Contents
1 Introduction 7
1.1 Chronobiologyanditsapplicationincaptivemanagement . . . . . . . . . . 7
1.1.1 Circadian rhythmsandtheinternalclock . . . . . . . . . . . . . . . . 7
1.1.2 Zeitgeber,entrainmentandmasking . . . . . . . . . . . . . . . . . . . 8
1.1.3 Seasonalchangesinperiodicity . . . . . . . . . . . . . . . . . . . . . . 10
1.1.4 Animalwell-being–aproblemofmeasurement . . . . . . . . . . . . 10
1.1.5 Chronoethology–atooltomeasureanimalwell-being?. . . . . . . . 12
1.2 TheKoala, Phascolarctos cinereus,Goldfuss1817 . . . . . . . . . . . . . . . . . 13
1.2.1 Taxonomyandanatomy . . . . . . . . . . . . . . . . . . . . . . . . . . 13
1.2.2 Habitat,distributionandstatus . . . . . . . . . . . . . . . . . . . . . . 16
1.2.3 Nutritionanddigestion . . . . . . . . . . . . . . . . . . . . . . . . . . 17
1.2.4 Socialstructure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
1.2.5 Chronobiologyandtimebudget . . . . . . . . . . . . . . . . . . . . . 20
1.2.6 Reproductivebiology . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
1.2.7 Koalasincaptivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
1.3 Aimsofthisstudy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2 Materials andMethods 31
2.1 ZoosandObservedAnimals . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
2.1.1 TarongaZoo,Sydney,Australia . . . . . . . . . . . . . . . . . . . . . . 31
2.1.2 ZooDuisburg,Germany . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.1.3 TiergartenSchönbrunn,Vienna,Austria . . . . . . . . . . . . . . . . . 41
2.2 VideoObservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
2.3 Analysisofvideotapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
2.3.1 Behaviouralcategories . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
2.3.2 Places. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
2.4 Analysisofdata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
vii2.4.1 Generaldataanalysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
2.4.2 Chrono-ethograms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
2.4.3 Periodlengthsandpowerspectrum . . . . . . . . . . . . . . . . . . . 46
2.4.4 Activityprofiles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
2.4.5 Timebudgetandday:nightratio . . . . . . . . . . . . . . . . . . . . . 47
2.4.6 PSTH-graphs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
2.5 Statisticalanalysisanderrorestimation . . . . . . . . . . . . . . . . . . . . . 48
3 Results 49
3.1 SydneyKoalaWalkabout(TarongaZoo) . . . . . . . . . . . . . . . . . . . . . 49
3.1.1 Chronoethogramoftotalactivity . . . . . . . . . . . . . . . . . . . . . 49
3.1.2 Timebudgetsandday:nightratiosofbehaviour . . . . . . . . . . . . 55
3.1.3 Seasonalvariations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
3.1.4 Feedingbouts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
3.1.5 Reference-Ethograms. . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
3.2 Duisburg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
3.2.1 Chronoethogramandactivityprofiles . . . . . . . . . . . . . . . . . . 68
3.3 TiergartenSchönbrunn,Vienna . . . . . . . . . . . . . . . . . . . . . . . . . . 86
3.3.1 Chronoethogramandactivityprofiles . . . . . . . . . . . . . . . . . . 86
3.4 Comparison Koala Walkabout (Sydney) – Duisburg Zoo – Tiergarten
SchönbrunnVienna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
3.4.1 Activitypattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
3.4.2 TimeBudget . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
3.4.3 Day:nightratio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
3.4.4 Feedingbouts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
3.5 Influenceoflight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
3.6 Influencesofkeepersandhandling . . . . . . . . . . . . . . . . . . . . . . . . 106
3.6.1 SydneyKoalaWalkabout(TarongaZoo) . . . . . . . . . . . . . . . . . 106
3.6.2 Duisburg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
3.6.3 TiergartenSchönbrunnVienna . . . . . . . . . . . . . . . . . . . . . . 114
3.6.4 Influenceoffixedfeedingtimes . . . . . . . . . . . . . . . . . . . . . . 116
3.7 Theinfluenceofvisitors: SydneyKoalaEncounter(TarongaZoo) . . . . . . 129
3.7.1 Reference-chronoethograms . . . . . . . . . . . . . . . . . . . . . . . . 129
3.7.2 ComparisontoKoalaWalkabout . . . . . . . . . . . . . . . . . . . . . 134
3.7.3 Feedingbouts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
3.7.4 Externalinfluences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1383.7.5 Changesinbehaviour: Locomotoractivity . . . . . . . . . . . . . . . 141
3.8 SocialbehaviouratKoalaWalkabout . . . . . . . . . . . . . . . . . . . . . . . 143
3.9 Location . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
3.9.1 TarongaZoo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
3.9.2 Duisburg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
4 Discussion 151
4.1 KoalaWalkabout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
4.1.1 GeneralactivitypatternandactingZeitgebers . . . . . . . . . . . . . 151
4.1.2 Entrainmentandmasking–competingstimuli . . . . . . . . . . . . . 152
4.1.3 Timebudget-comparisontofree-rangestudiesandseasonalvaria-
tions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
4.2 Comparisonofzoos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
4.2.1 Activitypatterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
4.2.2 TimeBudget . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
4.3 Zeitgeberandmaskinginindoorenclosures . . . . . . . . . . . . . . . . . . 162
4.4 Influenceofkeeperandhandling . . . . . . . . . . . . . . . . . . . . . . . . . 164
4.4.1 Cleaning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
4.4.2 Foodpresentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
4.4.3 Weighing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
4.4.4 ChangeofweighingtimeatViennaZoo . . . . . . . . . . . . . . . . . 167
4.5 KoalaEncounter-Influenceofvisitors . . . . . . . . . . . . . . . . . . . . . . 168
4.5.1 Generalactivitypattern . . . . . . . . . . . . . . . . . . . . . . . . . . 168
4.5.2 Influenceofvisitors . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
4.6 Increasedlocomotoractivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172
4.7 Chronoethology–atooltoevaluatewell-being? . . . . . . . . . . . . . . . . 173
4.8 Restinglocation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
5 Suggestedchangesinkoalahusbandry 177
6 Summary 179
A EthogramsatKoalaWalkabout,TarongaZoo,Sydney 183
A.1 Totalactivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
A.2 Feeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
A.3 Locomotoractivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
A.4 Presenceonground . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197A.5 Presenceincanopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
A.6 Presenceintreeunderneaththecanopy . . . . . . . . . . . . . . . . . . . . . 209
B EthogramsatKoalaEncounter,TarongaZoo,Sydney 215
C Individualtimebudgetsandday:nightratiosforallobservedkoalas 221
D MeteorologicaldataforSydney 233
E TimesofphotosessionsatKoalaEncounter 235
F MammalspeciesatKoalaWalkabout 237
G Detailsofstatistical analysis 239
Acknowledgements 245
References 249