Facultative butterfly-ant interactions – the role of variation in composition of nectar secretions Facultative butterfly-ant interactions – the role of variation in composition of nectar secretions Dissertation zur Erlangung des Grades eines Doktors der Naturwissenschaften – Dr. rer. nat. – der Fakultät Biologie / Chemie / Geowissenschaften der Universität Bayreuth Vorgelegt von Holger Daniels Bayreuth, Mai 2004 Die vorliegende Arbeit wurde am Lehrstuhl Tierökologie I der Universität Bayreuth in der Arbeitsgruppe von Herrn Prof. Dr. Konrad Fiedler angefertigt. Gefördert durch die Deutsche Forschungsgemeinschaft im Rahmen des Graduiertenkollegs 678: „Ökologische Bedeutung von Wirk- und Signalstoffen bei Insekten - von der Struktur zur Funktion“. Vollständiger Abdruck der von der Fakultät für Biologie, Chemie und Geowissenschaften der Universität Bayreuth genehmigten Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.). Tag der Einreichung: 04. Mai 2004 Tag des Promotionskolloquiums: 21. Juli 2004 Prüfungsausschuß: Prof. Dr. K. Fiedler (Erstgutachter) Prof. Dr. G. Rambold (Zweitgutachter) Prof. Dr. E. Komor (Vorsitzender) Prof. Dr. K. Dettner Prof. Dr. K. Seifert Dissertation als pdf-Datei und Bilder aus den Farbtafeln in höherer Auflösung auf Nachfrage verfügbar. Thesis available as pdf-file, likewise single pictures out of the colour plates in high resolution. Kontakt / contact: [email protected] Sie geben um zu leben Khalil Gibran (Vom Geben) Gewidmet dem Andenken an Walter Peter 1 INTRODUCTION .................................................................................................................4 2 GENERAL LIFE-HISTORY TRAITS OF THE SPECIES INVESTIGATED.....................13 2.1 Lycaenid butterflies ........................................................................................................13 2.2 Food plants .....................................................................................................................15 2.3 Ants ................................................................................................................................16 3 DEVELOPMENT OF SAMPLING METHODS..................................................................26 3.1 Abstract..........................................................................................................................26 3.2 Introduction....................................................................................................................26 3.3 Obtaining an artificial second generation of P. coridon...................................................27 3.3.1 Material & methods ................................................................................................... 27 3.3.2 Results...................................................................................................................... 29 3.3.3 Discussion ................................................................................................................ 30 3.4 Stimulation of the dorsal nectar organ............................................................................31 3.4.1 Techniques tested...................................................................................................... 32 3.4.2 Discussion ................................................................................................................ 36 3.5 Construction of an ”artificial ant“ ..................................................................................38 3.5.1 Material & methods ................................................................................................... 38 3.5.2 Results...................................................................................................................... 39 3.5.3 Discussion ................................................................................................................ 41 3.5.4 Acknowledgements ................................................................................................... 42 3.6 References.......................................................................................................................42 4 CHEMICAL COMPOSITION OF LARVAL NECTAR SECRETIONS FROM THREE SPECIES OF MYRMECOPHILOUS LYCAENID BUTTERFLIES..........................................46 4.1 Abstract..........................................................................................................................46 4.2 Introduction....................................................................................................................47 4.3 Material & Methods .......................................................................................................48 4.3.1 Insect collection and rearing....................................................................................... 48 4.3.2 Sampling and processing of nectar secretions and hemolymph ..................................... 50 4.3.3 Statistical analysis ..................................................................................................... 51 4.4 Results ............................................................................................................................51 4.4.1 Sugar composition and concentration.......................................................................... 51 4.4.2 Amino acid composition and concentration ................................................................. 54 4.4.3 Differences between the species ................................................................................. 59 4.4.4 The influence of larval food ....................................................................................... 61 4.4.5 Comparison of the amino acid profiles........................................................................ 62 4.5 Discussion.......................................................................................................................65 4.6 Acknowledgements .........................................................................................................71 4.7 References.......................................................................................................................71 5 A TASTE FOR BLUES? – BIOASSAYS WITH ANTS FROM DIFFERENT SUBFAMILIES ON COMPONENTS AND ANALOGA OF LYCAENID CATERPILLAR SECRETIONS...........................................................................................................................77 5.1 Abstract..........................................................................................................................77 5.2 Introduction....................................................................................................................77 5.3 Material & Methods .......................................................................................................79 5.4 Results ............................................................................................................................82 5.5 Discussion.......................................................................................................................93 5.6 Acknowledgements .........................................................................................................98 5.7 References.......................................................................................................................98 6 BENEFITS DERIVED FROM LARVAL NECTAR SECRETIONS OF FACULTATIVELY ANT-ASSOCIATED LYCAE NID BUTTERFLIES – DOES TENDING PAY FOR ANTS? ...103 6.1 Abstract........................................................................................................................103 6.2 Introduction..................................................................................................................103 6.3 Model............................................................................................................................105 6.4 Discussion.....................................................................................................................108 6.5 References.....................................................................................................................114 7 SYNOPSIS.......................................................................................................................119 8 SUMMARY ......................................................................................................................135 9 ZUSAMMENFASSUNG (GERMAN SUMMARY)...........................................................137 10 DARSTELLUNG DES EIGENANTEILS.....................................................................141 11 ACKNOWLEDGEMENTS...........................................................................................142 12 APPENDIX...................................................................................................................144 12.1 “Artificial ant” (chapter 3.5).........................................................................................144 12.2 Semi-synthetic diet formulation (“artificial food”) (chapter 4)......................................144 12.3 Erklärung .....................................................................................................................146 1 Introduction H. Daniels Department of Animal Ecology I, University of Bayreuth, D-95440 Bayreuth, Germany Interspecific mutualisms are ubiquitous and fundamentally influential at all levels of biological organization