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Social Life-Styles in Caterpillars: Behavioral Mechanisms and Ecological Consequences Claudia

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Social Life-Styles in Caterpillars: Behavioral Mechanisms and Ecological Consequences Claudia Social Life-Styles in Caterpillars: Behavioral Mechanisms and Ecological Consequences Claudia Ruf Social Life-Styles in Caterpillars: Behavioral Mechanisms and Ecological Consequences Dissertation zur Erlangung des Grades eines Doktors der Naturwissenschaften – Dr. rer. nat. – der Fakultät Biologie / Chemie / Geowissenschaften der Universität Bayreuth Vorgelegt von Claudia Ruf Bayreuth, Dezember 2002 Die vorliegende Arbeit wurde am Lehrstuhl Tierökologie I der Universität Bayreuth in der Arbeitsgruppe von Herrn Prof. Dr. Konrad Fiedler angefertigt. Gefördert durch ein Stipendium der Bayerischen Graduiertenförderung und durch die Deutsche Forschungsgemeinschaft (Fi 547/8-1). Vollständiger Abdruck der von der Fakultät für Biologie, Chemie und Geowissenschaften der Universität Bayreuth genehmigten Dissertation zur Erlangung des Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) Tag der Einreichung: 03. Dezember 2002 Tag des Promotionskolloquiums: 28. Februar 2003 Prüfungsausschuß: Prof. Dr. K. Fiedler (Erstgutachter) PD Dr. B. Stadler (Zweitgutachter) Prof. Dr. K. Dettner (Vorsitzender) Prof. Dr. D. von Holst Prof. Dr. Th. Foken Dissertation als PDF-Datei verfügbar PhD thesis available as PDF file Kontakt / contact: [email protected] PRO CAPTV LECTORIS HABENT SVA FATA LIBELLI TERENTIANVS MAVRVS Meinen Eltern gewidmet Contents Preface i Chapter one: General introduction 1 Chapter two: General life-history traits of the species investigated 11 Chapter three: Physical basics of thermoregulation 27 Chapter four: Thermoregulation by endogenous means: Metabolic heat production 35 Chapter five: Tent-based thermoregulation: Behavioral mechanisms and physical features of the tent 45 Chapter six: Plasticity in foraging patterns of larval colonies of Eriogaster lanestris 67 Chapter seven: Trail-based communication in social caterpillars of Eriogaster lanestris 93 Chapter eight: The mechanisms of trail-based communication: Trail marking and recruitment 111 Chapter nine: Mechanisms for and consequences of behavioral synchronicity in Eriogaster lanestris 135 Chapter ten: Colony survivorship of social caterpillars in the field: A case study 159 Chapter eleven: Larval sociality in three species of central-place foraging (Lasiocampidae): A comparative survey 179 Chapter twelve: Implications of the gregarious life-style on the development of Araschnia levana 207 Chapter thirteen: Group size influences larval survival of Araschnia levana 227 Chapter fourteen: Synopsis 257 v Contents Summary 265 Zusammenfassung (German Summary) 267 List of publications 273 Appendix 275 Erklärung (Statement, in German) 283 vi Preface Some remarks in advance… This doctoral thesis follows the widespread international standard of PhD theses and is structured in independent chapters which together build a closed unity, yet can be read separately without having read preceding chapters (cumulative design). Nevertheless, accompanying chapters shall help the reader to gain further concomitant information and give the whole thesis coherence. In Chapter one I give a general introduction on social caterpillars and a first overview of the present state of research as well as an outline of the thesis including the formulation of the questions and hypotheses addressed. Chapters two and three mainly include literature basics and show only few original data. They give an overview of the species investigated and some additional information on physical details of thermoregulation which might be helpful for those readers who have not yet dealt with this subject. Chapters four to thirteen all show original data and follow the general appearance of scientific papers since the majority of these chapters has been published yet (Chapters 4,5,6,7) or are presently under review (Chapters 8,10,11). I decided to accompany each chapter with its own references. Although this means partial redundancy of references, readers who are only interested in partial aspects of this thesis benefit from the integrity of each chapter. In a last chapter, the synopsis, I merge the contents of all chapters to provide general discussion and conclusions about the significance of the data for caterpillar sociality. Acknowledgements First of all I want to express my gratitude to three persons who made an important contribution for the success of this PhD thesis as they acted as my advisors or helped me during a long time during the last three years: I acknowledge the support of my doctoral advisor PROF. DR. KONRAD FIEDLER at any time of my work, especially regarding statistical analyses and his steady readiness for discussions. I owe special thanks to BERND KORNMAIER for technical consulting throughout the course of my work. Without him most electronical measurements would not have been possible. I also thank him for providing three privately owned computers which were essential for measuring activity patterns. Furthermore, I thank him for his incredible patience in our private life which was absolutely dominated by caterpillars for many months during the last years. In addition I am much obliged to JÖRG HAGER who helped establish experiments and maintaining colonies. In particular I thank him for providing help with vii Preface collecting and cleaning the cocoons of Eriogaster despite intense problems with erucism in the second year. Besides, I am grateful to a number of persons who provided helpful comments during the course of the preparation of some experiments and parts of this thesis. I am grateful to PROF. DR. JAMES T. COSTA, Western Carolina University, for his contribution to the discussion of Chapter 7 as well as some hints to maintaining colonies and relevant literature. PROF. DR. THOMAS FOKEN, Dept of Micrometeorology, gave helpful evidence for temperature measurements and good ideas with respect to the physical features of the tent (Ch. 5). Data from the meteorological station in the Botanical Garden of the University of Bayreuth were provided by JÖRG GERCHAU. STEFAN FORERO, Dept of Experimental Physics II, conducted the spectrophoto- metrical analyses (Ch. 5). MARIELUISE OBERMAYER, Univ. of Würzburg, made the histological studies from Chapter 8. ANJA FREESE established contact with the E. catax project which made the studies from Chapter 11 possible. I also thank my collegues notably HOLGER DANIELS, DIRK SÜßENBACH and DR. KLAUS FISCHER for several small favors that made life easier now and then. The last three years have been overshadowed by severe health problems. I thank JÖRG HAGER, FRANK RAMMING and BERND KORNMAIER for numerous rides to Erlangen which enabled me to avoid further surgery before the end of the doctoral thesis. This work was supported by a grant from the BAYERISCHE GRADUIERTENFÖRDERUNG and the DEUTSCHE FORSCHUNGSGEMEINSCHAFT (Fi 547/8-1). Last but not least I am obliged to my parents who gave me moral and financial support during the years of my studies. viii Chapter 1 General Introduction Present state of research Butterflies and moths (i.e. the order Lepidoptera) probably represent the best known taxon among the phytophagous insects. However, there is a deep gap in our knowledge about the ecology of the predominant number of species which is not only true for tropical species, but even for many of the most common species of our local fauna. This particularly holds for the larval stages of most Lepidoptera. Hence, the larval ecology is only well understood for a rather small number of species which are of economic interest as they are forest or agricultural pests. The larval stages of the majority of the about 160,000 described species of butterflies and moths show solitary life styles. In contrast, only about 300 species are known so far which exhibit gregarious or ‘social’ life-styles for at least part of their larval development (for a general discussion of terms see: Costa & Fitzgerald 1996, Costa & Pierce 1997). Even though this list is certainly incomplete because of the nearly complete lack of knowledge of life-history traits of most tropical species it seems obvious that evolution favors solitary life- styles in most cases investigated because of the high costs related to social life. The costs of larval societies are obvious: Caterpillars are slow moving insects and suffer great losses from predators and parasitoids which could be even more pronounced in groups of larvae that are more conspicuous to their enemies (Costa 1993, Knapp & Casey 1986, Stamp & Bowers 1988). Besides, groups of caterpillars have a higher risk of being infected by pathogens (Hochberg 1991) and may rapidly overexploit their larval resources because of competition for food within a colony which may result in the inability to finish development and sometimes the death of the whole offspring of a female (Tsubaki 1995). 1 Chapter one: General introduction Five factors acting singly or in combination are commonly regarded to be responsible for the fact that despite strong selective pressures against larval communities such systems may evolve and persist over time: 1. The inability of the imago to ingest food favors egg clustering Comparative investigations revealed that a reduction of the proboscis in imagines frequently makes for clustering eggs in large clutches because of energetic constraints (Miller 1996, Tammaru & Haukioja 1996, Hebert 1983). Caterpillars hatching from an egg cluster immediately leave their siblings in most species. Nevertheless, egg clustering is obligatory for evolving social life- styles in caterpillars and gregarious larvae are significantly more frequent in species
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