The Evolution of Social Traits and Biodiversity in the Ants
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The evolution of social traits and biodiversity in the ants. Thesis submitted by Henry Ferguson-Gow For the examination of the degree of Doctor of Philosophy University of East Anglia September 2014 This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and that use of any information derived there from must be in accordance with current UK Copyright Law. In addition, any quotation or extract must include full attribution. i 1 Abstract Cooperation has shaped the evolution of life on Earth. The ants are the most numerically diverse of the eusocial Hymenoptera, and display wide variation in social complexity. This positions the ants as an ideal taxon in which to study social evolution in a comparative framework. Social evolution theory has generated many hypotheses that are testable in ants, however the lack of comprehensive or complete phylogenies, and the decentralised and scattered nature of trait data, has been an obstacle to these types of study. In this thesis I construct a large species-level, and a complete genus-level, phylogeny of the ants, and draw together a large dataset of social traits from the literature in order to test hypotheses concerning the evolution of social traits in the ants. I find evidence that the earliest ant was large bodied, and lived in small highly related colonies. I show that group size is a significant trait in the evolution of sociality in ants, predicting the probability of a species having polymorphic workers, or of being polyandrous. I also show that the change in these traits is correlated between ancestral nodes on the phylogeny. Furthermore, in the Attini, colony size correlates closely with non-reproductive and reproductive division of labour. Together these results cement group size as a driving force of social evolution in the ants, and this has interesting implications for social evolution in general. Finally, I report the first evidence that intermediate colony sizes, the presence of discrete worker castes and polygyny are associated with increased diversification rates in ants. This thesis provides a valuable tool for the study of comparative hypotheses in the ants in the form of a complete genus-level phylogeny, and offers significant evidence to support several key hypotheses in social evolution. Furthermore, these results generate hypotheses regarding the evolution of social traits for future research. ii 2 Table of Contents 1 Abstract ................................................................................................................... ii 2 Table of Contents .................................................................................................. iii 3 List of tables .......................................................................................................... vi 4 List of figures ......................................................................................................... ix 5 Acknowledgements .............................................................................................. xii 6 Declaration ........................................................................................................... xiii 1 Introduction ............................................................................................................. 1 1.1 The ants ............................................................................................................................. 1 1.2 The importance of social evolution .................................................................................. 2 1.3 Phylogenies and comparative biology .............................................................................. 4 1.4 Thesis overview ................................................................................................................. 4 2 A supertree phylogeny and a trait database for the Formicidae ......................... 7 2.1 Abstract ............................................................................................................................. 7 2.2 Introduction ...................................................................................................................... 7 2.2.1 The study of macroevolution .................................................................................... 7 2.2.2 Ant macroevolution .................................................................................................. 8 2.2.3 The history of ant systematics and phylogenetics .................................................... 8 2.2.4 Building large or complete phylogenies ..................................................................11 2.2.5 Trait data .................................................................................................................14 2.3 Methods ..........................................................................................................................14 2.3.1 Phylogenetic reconstruction ...................................................................................14 2.3.2 Trait database .........................................................................................................20 2.4 Results .............................................................................................................................25 2.4.1 Phylogenetic reconstruction ...................................................................................25 2.4.2 Trait data .................................................................................................................31 2.5 Discussion ........................................................................................................................34 2.5.1 Phylogenetic reconstruction ...................................................................................34 2.5.2 Database .................................................................................................................36 3 The phenotype of the ancestral ant ..................................................................... 38 3.1 Abstract ...........................................................................................................................38 3.2 Introduction ....................................................................................................................38 3.3 Method ...........................................................................................................................44 3.3.1 Trait Data.................................................................................................................44 3.3.2 Supertree construction ...........................................................................................45 iii 3.3.3 Ancestral state reconstruction ................................................................................45 3.3.4 Rates of worker head-width evolution ...................................................................47 3.4 Results .............................................................................................................................48 3.4.1 Ancestral state reconstructions ..............................................................................48 3.4.2 Rate shifts in worker head-width evolution ...........................................................50 3.5 Discussion ........................................................................................................................53 4 Social trait evolution and coevolution in the ants. ............................................. 56 4.1 Abstract ...........................................................................................................................56 4.2 Introduction ....................................................................................................................56 4.2.1 The evolution of social traits ...................................................................................57 4.2.2 Correlated evolution in social traits ........................................................................58 4.3 Method ...........................................................................................................................60 4.3.1 Trait data .................................................................................................................60 4.3.2 Supertree construction ...........................................................................................61 4.3.3 Ancestral state reconstructions ..............................................................................61 4.3.4 Correlated evolution ...............................................................................................61 4.4 Results .............................................................................................................................62 4.4.1 Ancestral state reconstructions ..............................................................................62 4.4.2 Correlated evolution ...............................................................................................62 4.5 Discussion ........................................................................................................................66 5 Colony size predicts division of labour in Attine ants ....................................... 69 5.1 Abstract ...........................................................................................................................69 5.2