Life History and Mating Behaviour of the Scorpionfly Panorpa Alpina Rambur (Mecoptera): Choice and Strategy with Remarks on the Methods of Causal Modeling

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Life History and Mating Behaviour of the Scorpionfly Panorpa Alpina Rambur (Mecoptera): Choice and Strategy with Remarks on the Methods of Causal Modeling Life History and Mating Behaviour of the Scorpionfly Panorpa alpina Rambur (Mecoptera): Choice and Strategy With Remarks on the Methods of Causal Modeling Lebensgeschichte und Paarungsverhalten der Skorpionsfliege Panorpa alpina Rambur (Mecoptera): Wahl und Strategie Mit Anmerkungen zu den Methoden der kausalen Modellierung Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von Gerhild Bornemann aus Melle Bonn 2012 Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen-Friedrich-Wilhelms-Universität Bonn. 1. Gutachter: Prof. Dr. K. P. Sauer 2. Gutachter: Prof. Dr. W. Alt Tag der Promotion: 17. 12. 2012 Erscheinungsjahr: 2013 Abstract Zusammenfassung LLLife history theory bases on the fundamental DDDie Theorie der Lebensgeschichte basiert auf assumption that the behaviour of living things at der grundsätzlichen Annahme, dass das any time of their lives aims at the maximisation of Verhalten von Lebewesen zu jedem Zeitpunkt their reproductive success. The present thesis ihres Lebens darauf ausgerichtet ist, den eigenen adopts this assumption, which is in accordance Reproduktionserfolg zu maximieren. Diese with the flexibility of the condition-dependent Annahme ist auch die Basis der vorliegenden mating behaviour of scorpionfly males. Their Arbeit, da sie sich mit der bekannten Flexibilität mating strategy can be composed of two or three des konditions-abhängigen Paarungsverhaltens different tactics, one of which involves the energy von Skorpionsfliegenmännchen gut in Überein- consuming production of salivary secretions as stimmung bringen lässt. Ihre Paarungsstrategie nuptial gifts. The choice of the actual tactic is an kann sich aus bis zu drei verschiedenen Taktiken adaptation of the behaviour to the current zusammensetzen, von denen eine die energie- situation. aufwändige Produktion von Sekreten als As the mating behaviour of the scorpionfly Paarungsgeschenk beinhaltet. Die Wahl der Panorpa alpina was yet unknown the study Taktik ist eine Anpassung des Verhaltens an die started with lifetime observations of test aktuelle Situation. populations that were assigned to different Da zum Paarungsverhalten der nutrient regimes. They showed that this species is Skorpionsfliege Panorpa alpina noch keine Daten mainly diurnal and both sexes mate repeatedly. vorlagen, begann die Untersuchung mit Lebens- Besides, the results indicated that males and zeitbeobachtungen von Versuchspopulationen females are cryptically choosy and that females unter verschiedenen Nahrungsbedingungen. Es also exert precopulatory choice. zeigte sich, dass diese Art hauptsächlich bei Tage During observation only two of the known aktiv ist und beide Geschlechter sich mehrfach tactics of scorpionfly males were recorded. A verpaaren. Die Ergebnisse lieferten außerdem mate choice experiment, in which differently Hinweise auf ein kryptisches Wahlverhalten bei nourished individuals were mated in all possible Männchen und Weibchen, sowie präkopu- combinations, revealed that the males apply a latorische Wahl der Weibchen. third tactic, which has not been described before. Während der Beobachtung wurden nur For the analysis of the experimental data, in zwei der drei bei Skorpionsfliegenmännchen addition to conventional methods like ANOVA and bekannten Paarungstaktiken festgestellt. In einem ANCOVA, structural equation models have been Partnerwahlexperiment, in dem verschieden used to compare different hypotheses about ernährte Individuen in allen Kombinationen choice and strategy. Both kinds of methods miteinander verpaart wurden, stellte sich jedoch unanimously produce the result that precopulatory heraus, dass die Männchen eine weitere, bislang and cryptic choice occur in males as well as noch nicht beschriebene Taktik anwenden. Bei females. Also both sexes behave strategically, i.e. der Auswertung des Experimentes kamen neben they adjust their behaviour to their own body den üblichen Methoden wie ANOVA und condition. It can be gathered from the results that ANCOVA auch Strukturmodelle zur Anwendung, male behaviour changes along a gradient of mit denen verschiedene Hypothesen über Wahl nutritional states. Behaviour ranges from und Strategie vergleichend überprüft werden indiscrimination shown by males of very low konnten. Beide Verfahrensweisen führten nutritional state and cryptic choice in the literal übereinstimmend zu dem Ergebnis, dass sowohl sense shown by males of average nutritional Männchen wie auch Weibchen präkopulatorisch state, to the newly discovered tactic called und kryptisch wählen. Außerdem verhalten sich adjusted paternal effort, which is shown by males beide Geschlechter strategisch, d.h. sie passen of very good nutritional state. Such well nourished ihr Verhalten an ihren eigenen Zustand an. Aus males the more reduce the volume of the salivary den Ergebnissen lässt sich ableiten, dass sich das secretions that they produce and feed to the Verhalten der Männchen entlang eines female during copulation, the better nourished the Gradienten des Ernährungszustandes verändert. respective female is. This tactic is closely Die Spannweite reicht von Wahllosigkeit bei sehr connected to the behaviour of the females that schlecht ernährten Männchen, über kryptische grow increasingly indiscriminate with increasing Wahl im klassischen Sinne bei durchschnittlich nutritional state. Moreover, when mating with well ernährten Männchen, bis hin zu der bisher noch fed males, females generally are less restrictive. nicht beschriebenen Taktik, dem angepassten Thus, well fed males can expect a high pay off väterlichen Aufwand, bei sehr gut ernährten despite their reduced investment. Männchen. Diese verringern das Volumen der von The results indicate that the pay off of a specified ihnen produzierten und den Weibchen behaviour depends strongly on the interplay of präsentierten Geschenke umso mehr, je besser male and female behaviour. Therefore, it is ernährt das Weibchen ist, mit dem sie sich important to include both sexes when studying paaren. Diese Taktik ist eng mit dem Verhalten choice mechanisms. As the behaviour of both der Weibchen verknüpft, bei denen der Grad der sexes is governed by their own state and the state kryptischen Wählerischkeit mit einer of the partner and as these are possibly both Verbesserung ihres Ernährungszustandes assessed by multiple cues, it is imperative to abnimmt. Zudem sind sie bei Zusammentreffen conduct detailed analyses to understand mating mit sehr gut ernährten Männchen generell behaviour. With regard to that the methods of weniger restriktiv, so dass ein sehr gut ernährtes statistical modelling are beneficial because the Männchen trotz verringerter Investition eine hohe flexible models allow applying more realistic Auszahlung erwarten kann. model structures that can be tested statistically in Die Auszahlung, die ein Verhalten addition to the tests addressing the explanatory einbringt, ist also stark von den value. Wechselwirkungen zwischen Männchen- und Weibchenverhalten bestimmt. Daher ist es wichtig, bei der Untersuchung von Wahlmechanismen beide Seiten mit einzubeziehen. Da das gezeigte Verhalten außerdem bei beiden Geschlechtern vom eigenen sowie dem Zustand des Partners abhängt, die möglicherweise beide anhand von mehreren Merkmalen eingeschätzt werden, sind detaillierte Analysen für das Verständnis des Wahlverhaltens unumgänglich. In dieser Hinsicht sind die Methoden der kausalen Modellierung vorteilhaft, weil die flexible Modellstruktur wirklichkeitsnähere Modellierungen erlaubt und sie, neben dem Erklärungswert der Variablen, auch die angenommene Kausalstruktur einer statistischen Überprüfung zugänglich machen. “We shall pick up an existence by its frogs” Charles Fort (1931): Lo! I Contents Inhalt Section I Introduction Einleitung Chapter I Theoretical Background Theoretische Grundlagen Life Histories and Lifetime Observations Lebensgeschichten und Lebenszeitbeobachtungen..................................................................... 17 Research Questions Fragestellungen............................................................................................................................ 18 The Knowledge about P. vulgaris as Guideline for the Work on the Life History of P. alpina Das Wissen über P. vulgaris als Leitlinie für die Arbeit über die Lebensgeschichte von P. alpina ................................................................ 19 Chapter II An Introduction to Statistical Modeling Einführung in die statistische Modellierung Correlation Korrelation..................................................................................................................................... 20 Path Analysis and Structural Equation Modeling: Methods and Development in their historical Context Pfadanalysen und Strukturmodelle: Methoden und Entwicklung im historischen Kontext..................................................................... 21 Box 1: Path Analysis Box 1: Pfadanalyse....................................................................................................................... 23 Box 2: Structural Equation Modeling Box 2: Strukturmodelle.................................................................................................................. 27 Box 3: Directional Separation Box 3: Direktionale Separation...................................................................................................... 30 Terminology Terminologie................................................................................................................................
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