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Demography in Eusocial Hymenoptera Demography in eusocial Hymenoptera Dissertation Zur Erlangung des Grades Doktor der Naturwissenschaften Am Fachbereich Biologie Der Johannes Gutenberg-Universitӓt Mainz Boris Hendrik Kramer geb. am 1. März 1978 in Essen Dekan: Prof. Dr. come on 1. Berichterstatter: Prof. Dr. black for a reason 2. Berichterstatter: Prof. Dr. stop trying Tag der mündlichen Prüfung: 19. Juni 2013 2 "If you want to be incrementally better, be competitive. If you want to be exponentially better, be cooperative." Unknown Internet source 3 This thesis is based on the following four research articles, which are presented as chapters I to IV and are referred to in the introduction, the summary and the general discussion by their respective numbers: Chapter I Kramer, B.H ., Schaible, R. (2013): Life Span Evolution in Eusocial Workers—A Theoretical Approach to Understanding the Effects of Extrinsic Mortality in a Hierarchical System. PLoS ONE 8(4). Chapter II Kramer, B.H ., Scheuerlein, A.: Worker quality is determined by colony stage and not social environment in Lasius niger . Under review at Myrmecological News. Chapter III Kramer, B. H ., Schaible, R. (2013): Colony size explains the life span differences between queens and workers in eusocial Hymenoptera. Biological Journal of the Linnean Society, 109 (3): 710-724. Chapter IV Kramer, B.H., Scharf, I., Foitzik, S.: Colony size and per-capita productivity in ants with single foragers. Under review at Behavioral Ecology and Sociobiology. 4 Summary In my doctoral thesis I investigated the evolution of demographic traits within eusocial Hymenoptera. In the social bees, wasps and ants, eusociality has a unique effect on life span evolution as female larvae with the same genetic background can develop through phenotypic plasticity to a queen or a worker with vastly diverging life- history traits. Ant queens belong to the longest-lived insect species, while workers in most species live only a fraction of the queen’s life span. The average colony size of a species is positively correlated with social complexity, division of labor and diverging morphological female phenotypes all of which also affect life span. Therefore the demographic traits of interest in this thesis were life span and colony size. To understand the evolution of worker life span I applied a trade-off model that includes both hierarchical levels important in eusocial systems, namely the colony- and the individual-level. I showed that the evolution of worker life span may be an adaptive trait on the colony level to optimize resource allocation and therefore fitness in response to different levels of extrinsic mortality. A shorter worker life span as a result of reduced resource investments under high levels of extrinsic mortality increases colony fitness. In a further study I showed that Lasius niger colonies produce different aging phenotypes throughout colony development. Smaller colonies which apply a different foraging strategy than larger colonies produced smaller workers, which in turn have a longer life span as compared to larger workers produced in larger colonies. With the switch to cooperative foraging in growing colonies individual workers become less important for the colony caused by their increasing redundancy. Alternatively a trade of between growth and life span may lead to the results found in this study. A further comparative analysis to study the effect of colony size on life span showed a correlation between queen and worker life span when colony size is taken into account. While neither worker nor queen life span was associated with colony size, the differences between queen and worker life span increase with larger average colony sizes across all eusocial Hymenoptera. As colony size affects both queen and worker life span, I aimed to understand which factors lead to the small colony sizes displayed by some ant species. I therefore analyzed per-capita productivity at different colony sizes of eight cavity dwelling ant species. Most colonies of the study species grew larger than optimal productivity predicted. Larger colony size was shown to increase colony homeostasis, the predictability of future productivity and in turn the survival probability of the colony. I also showed that species that deploy an individual foraging mode may circumvent the density dependent decline in foraging success by splitting the colony to several nest sites. 5 Zusammenfassung In dieser Doktorarbeit habe ich die Evolution von demographischen Eigenschaften bei eusozialen Hautflüglern (Ameisen, Bienen und Wespen) untersucht. Eusozialität hat einen starken Einfluss auf die Evolution der Lebenspanne, aus einem weiblichen Ei können sich bei den sozialen Hautflüglern sowohl Königinnen als auch Arbeiterinnen Phänotypen entwickeln, die sich jedoch stark in ihrer Life-history unterscheiden. Ameisenköniginnen gehören zu den langlebigsten Insekten, aber leben Arbeiterinnen nur einen Bruchteil dieser Lebenspanne. Die durchschnittliche Koloniegröße ist hierfür ein bedeutender Faktor, denn sie steht in einem positiven Zusammenhang mit der Arbeitsteilung im Insektenstaat und den daraus resultierenden morphologischen Unterschieden zwischen den unterschiedlichen Kasten, die wiederum die Lebenspanne beeinflussen. Dementsprechend waren die Lebensspanne und die Koloniegröße die demographischen Eigenschaften auf die ich mich in dieser Doktorarbeit fokussiert habe. Um die Evolution der Lebenspanne der Arbeiterinnen zu untersuchen habe ich ein Trade-off Modell entwickelt, welches die hierarchische Struktur, die man bei sozialen Insekten findet, berücksichtigt. Ich konnte dadurch zeigen, dass die Investitionen in Arbeiterinnen bei hohem extrinsischem Sterberisiko reduziert werden sollten, um die Fitness der Kolonie zu optimieren. Interessanterweise ist in diesem Falle eine kürzere Arbeiterinnenlebenspanne mit einer höheren Fitness verbunden. In einer weiteren Studie habe ich gezeigt, dass während des Wachstums einer Lasius niger Kolonie die Investitionen in einzelne Arbeiterinnen variieren. Kleine Kolonien produzieren kleinere Arbeiterinnen mit einer längeren Lebensspanne als größere Kolonien. Die Unterschiede in der Lebensspanne können entweder durch den Größenunterschied oder aus den veränderten Anforderungen an die Arbeiterinnen resultieren, die mit der Änderung der furagier Strategie bei größeren Kolonien zusammenhängen. In einer weiteren, vergleichenden Studie habe ich dann den Effekt der Koloniegröße unterschiedlicher eusozialer Hautflügler auf die Lebensspanne untersucht. Während weder die Lebensspanne der Königinnen noch die der Arbeiterinnen mit der Koloniegröße im Bezug stehen, zeigt der Unterschied in der Lebenspanne zwischen diesen beiden Phänotypen einer Art eine positive Beziehung zur Koloniegröße. Um zu erklären, warum verschiedene Ameisenarten unterschiedlich große Kolonien bilden, habe ich bei acht Ameisenarten, die kleine Kolonien bilden, die Produktivität pro Arbeiterin untersucht. Es zeigte sich, dass diese pro-Kopf Produktivität, welche direkt die Fitness der Kolonie bestimmt, bei keiner der untersuchten Arten mit der Koloniegröße ansteigt. Eine Möglichkeit, die Reduktion der pro-Kopf Produktivität zu umgehen, ist das Aufteilen der Kolonie auf mehrere Nester, damit die gleiche Anzahl Arbeiterinnen auf eine größere Fläche verteilt ist. 6 Table of content Demography in eusocial Hymenoptera .................................................................................................................... 1 Summary .................................................................................................................................................................. 5 Zusammenfassung .................................................................................................................................................... 6 Table of content ....................................................................................................................................................... 7 General Introduction ................................................................................................................................................ 9 Demography .............................................................................................................................................. 9 Eusociality ............................................................................................................................................... 10 The evolution of life span within social insects ........................................................................................ 11 Division of labor ...................................................................................................................................... 15 Colony size ............................................................................................................................................... 18 Goals of this thesis ................................................................................................................................... 20 References ................................................................................................................................................ 23 Chapter I : Life span evolution in eusocial workers – a theoretical approach to understanding the effects of extrinsic mortality in a hierarchical system. ............................................................................................. 31 Abstract ...................................................................................................................................................
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