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Nicole Höcherl Nesting behaviour of the paper wasp Polistes dominula – with special focus on thermoregulatory mechanisms ………… Nistverhalten der Feldwespe Polistes dominula – mit besonderem Augenmerk auf thermoregulatorische Mechanismen Doctoral thesis for a doctoral degree at the Graduate School of Life Sciences, Julius-Maximilians-Universität Würzburg Section: Integrative Biology submitted by Nicole Höcherl from Wörth an der Donau Würzburg 2015 Submitted on: ........................................................................................... Office stamp Members of the Promotionskomitee: Chairperson: Prof. Dr. Thomas Dandekar Primary Supervisor: Prof. Dr. Jürgen Tautz Supervisor (Second): PD Dr. Johannes Spaethe Supervisor (Third): Prof. Dr. Stefano Turillazzi Date of Public Defence: …………………………………………..……… Date of receipt of Certificates: ………………………………………..….. Affidavit I hereby confirm that my thesis entitled ‘Nesting behaviour of Polistes dominula – with special focus on thermoregulatory mechanisms’ is the result of my own work. I did not receive any help or support from commercial consultants. All sources and/or materials applied are listed and specified in the thesis. Furthermore, I confirm that this thesis is has not yet been submitted as part of another examination process neither in identical nor similar form. Würzburg, 22.12.2015………………………………………… Eidesstattliche Erklärung Hiermit erkläre ich an Eides statt, die Dissertation ‘Nesting behaviour of Polistes dominula – with special focus on thermoregulatory mechanisms’ eigenständig, d.h. insbesondere selbstständig und ohne Hilfe eines kommerziellen Promotionsberaters, angefertigt und keine anderen als die von mir angegebenen Quellen und Hilfsmittel verwendet zu haben. Ich erkläre außerdem, dass die Dissertation weder in gleicher noch in ähnlicher Form bereits in einem anderen Prüfungsverfahren vorgelegen hat. Würzburg, 22.12.2015………………………………………… The Wasp to the Butterfly: “Non qui fuerimus, sed qui nunc simus vide!“ “Do not consider what we were, but what we are!“ „Schau nicht nach dem, was wir waren, sondern auf das was wir sind!“ from Phaedrus, Liber Fabularum, Appendix Perottina 31: Papilio et Vespa (The Butterfly and the Wasp; Der Schmetterling und die Wespe) TABLE OF CONTENT LIST OF USED ABBREVIATIONS AND SYMBOLS ........................................................ 8 SUMMARY ............................................................................................................................. 9 ZUSAMMENFASSUNG ....................................................................................................... 12 1. INTRODUCTION ............................................................................................................ 15 1.1 POLISTES DOMINULA – A MODEL ORGANISM FOR THE EVOLUTION OF SOCIALITY ............... 15 1.2 NESTING BEHAVIOUR OF POLISTES DOMINULA .................................................................. 17 1.3 THERMOREGULATION OF INDIVIDUAL PAPER WASPS ......................................................... 19 1.4 THERMOREGULATION OF POLISTES DOMINULA NESTS ....................................................... 21 1.5 THE AIMS OF THIS WORK ..................................................................................................... 24 2. MATERIALS ..................................................................................................................... 26 2.1 EQUIPMENT ................................................................................................................................... 26 2.2 SPECIAL SOFTWARE ........................................................................................................................ 27 3. METHODS ........................................................................................................................ 28 3.1 GENERAL REMARKS............................................................................................................. 28 3.1.1 Selection of suitable nesting sites .............................................................................................................................. 28 3.1.2 Selection of suitable Polistes dominula nests for thermographic studies ............................................................... 28 3.2 INVESTIGATION OF THE NESTING BEHAVIOUR OF P. DOMINULA ....................................... 29 3.2.1 Screening of abandoned Polistes dominula nests ....................................................................................................... 29 3.2.2 Observation of nesting season and nest sizes of newly founded Polistes dominula nests .................................. 30 3.2.3 Recording of foundress associations ........................................................................................................................ 30 3.2.4 Recording of survival rate and productivity ............................................................................................................ 30 3.3 INVESTIGATION OF THERMOREGULATORY MECHANISMS OF INDIVIDUAL POLISTES DOMINULA WASPS ................................................................................................ 31 3.3.1 Challenge of Polistes dominula individuals with stressors ........................................................................................ 31 3.3.2 Observation of dominance battles ............................................................................................................................ 32 3.4 INVESTIGATION OF THERMOREGULATORY MECHANISMS OF POLISTES DOMINULA NESTS .. 33 3.4.1 Measurement of the influence of adult paper wasps on the nest temperature ................................................. 33 3.4.1.1 Recording of the nest temperature with the help of thermography .......................................................... 33 3.4.1.2 Recording of the efficiency of cooling the nests by fanning and water evaporation ................................ 35 3.4.2 Measurement of the influence of the nest size on the nest temperature ........................................................... 35 3.4.3 Measurement of the influence of the site on the nest temperature .................................................................... 36 5 3.5 DATA EVALUATION AND STATISTICAL ANALYSES ................................................................. 37 3.5.1 General remarks ........................................................................................................................................................... 37 3.5.2 Analysis of the nesting behaviour of Polistes dominula ............................................................................................ 38 3.5.3 Analysis of the thermoregulation of individual Polistes dominula wasps............................................................... 38 3.5.4 Analysis of the thermoregulation of Polistes dominula nest ..................................................................................... 39 4. RESULTS ........................................................................................................................... 40 4.1 THE NESTING BEHAVIOUR OF POLISTES DOMINULA ........................................................... 40 4.1.1 Screening of abandoned nests ................................................................................................................................... 40 4.1.2 Length of nesting season and nest sizes .................................................................................................................. 42 4.1.3 Foundress associations ................................................................................................................................................ 47 4.1.4 Survival rate of Polistes dominula nests ....................................................................................................................... 47 4.1.5 Productivity of Polistes dominula nests ........................................................................................................................ 49 4.2 THERMOREGULATORY MECHANISMS OF INDIVIDUAL ADULT PAPER WASPS ......................... 51 4.2.1 The examination of different artificial stressors for investigating the thermal behaviour of P. dominula wasps .......................................................................................................................................................... 51 4.2.2 Response of Polistes dominula to the different artificial stressors smoke and CO2............................................. 53 4.2.3 The thermal behaviour of dominant and subordinate Polistes dominula queens during dominance battles ........................................................................................................................................................ 58 4.3 THERMOREGULATORY MECHANISMS IN POLISTES DOMINULA NESTS ................................. 60 4.3.1 The comparison of nest temperature between active and non-active nests ...................................................... 60 4.3.1.1. Single thermographic shots of an active nest revealed first insights of nest temperatures ....................... 60 4.3.1.2 Thermographic shots over 24h reveal differences in nest temperature between active and non-active nests ......................................................................................................................................... 62 4.3.1.3 Simultaneous monitoring of active and non-active nests at outside temperatures
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