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Phylogenetics of Myrmica Ants and Their Social Parasites Phylogenetics of Myrmica ants and their social parasites Gunther Jansen Department of Biological and Environmental Sciences Faculty of Biosciences University of Helsinki Finland Academic Dissertation To be presented, with permission of the Faculty of Biosciences of the University of Helsinki, for public criticism in Auditorium 1041, Biokeskus 2 (Viikinkaari 2) on the 26th of June 2009 at 12 o'clock noon Helsinki 2009 Phylogenetics of Myrmica ants and their social parasites c 2009 Gunther Jansen (Summary, Chapter II, III, IV) c 2009 Wiley-Blackwell (Chapter I) Author's address: Department of Biological and Environmental Sciences P.O. Box 65 00014 University of Helsinki Finland e-mail: gunther.jansen@helsinki.fi [email protected] ISBN 978-952-92-5568-9 (paperback) ISBN 978-952-10-5577-5 (PDF) http://ethesis.helsinki.fi/ 2 Phylogenetics of Myrmica ants and their social parasites \Denn nur im Irrtum, nur durch den Irrtum, in den er unentrinnbar hineingehal- ten ist, wird der Mensch zum Suchenden, der er ist, der suchende Mensch; denn der Mensch braucht die Erkenntnis der Vergeblichkeit, er muß ihren Schrecken, den Schrecken jeden Irrtums auf sich nehmen und, ihn erkennend, bis zur Neige auskosten, er muß des Schreckens inne werden, nicht aus Selbstqual, wohl aber weil nur in solch erkennenden Innewerden der Schrecken zu ¨uberwinden ist, weil nur dann es m¨oglich wird, durch des Schreckens h¨oherePforte hindurch ins Sein zu gelangen." | Hermann Broch, Der Tod des Vergil 3 Phylogenetics of Myrmica ants and their social parasites This thesis is based on the following articles, which will be referred to by their Roman numerals. I Jansen, G., R. Savolainen and K. Veps¨al¨ainen(In press). Barcoding as a heuristic tool for classifying Nearctic Myrmica ants. Zoologica Scripta. II Jansen, G., R. Savolainen and K. Veps¨al¨ainen(Manuscript). Phylogeny, social parasite - host relationships, divergence time estimation and bio- geography of the Holarctic ant genus Myrmica. III Jansen, G. and R. Savolainen (In press). Molecular phylogeny of the ant tribe Myrmicini (Hymenoptera: Formicidae). Zoological Journal of the Linnean Society. IV Jansen, G. and R. Savolainen (Manuscript). Phylogenetic test of parasite-host associations between Maculinea butterflies and Myrmica ants. 4 Phylogenetics of Myrmica ants and their social parasites Table of contributions I II III IV Original idea RS, GJ RS, KV RS RS, GJ Field work GJ, RS, KV RS, KV, GJ RS see (II), gb Lab work GJ GJ GJ GJ Data analysis GJ GJ GJ GJ Manuscript GJ, RS, KV GJ, RS, KV GJ, RS GJ, RS preparation GJ: Gunther Jansen; RS: Riitta Savolainen; KV: Kari Veps¨al¨ainen;additional samples have been obtained from colleagues as acknowledged in the individual papers; gb: sequences obtained from Genbank Supervised by: Dr. Riitta Savolainen Dr. Kari Veps¨al¨ainen Department of Biological and Environmental Sciences University of Helsinki Finland Reviewed by: Dr. Niklas Wahlberg University of Turku Finland Dr. Duur Aanen University of Wageningen The Netherlands Opponent: Dr. Se´anBrady Smithsonian Institution Laboratories of Analytical Biology MD, USA 5 Phylogenetics of Myrmica ants and their social parasites Contents 1 Introduction 7 1.1 The art of knowing what surrounds us . 7 1.2 Systematics . 7 1.2.1 Taxonomy of systematics . 7 1.2.2 Molecular taxonomy . 8 1.3 Systematics in motion: ants . 9 1.3.1 The family Formicidae . 9 1.3.2 The genus Myrmica ....................... 11 1.4 Social parasitism . 11 1.4.1 Emery's rule . 12 1.4.2 Maculinea: socially parasitic butterflies . 13 1.5 Outline of the thesis . 13 2 Materials & Methods 14 2.1 Samples . 14 2.2 Selection of molecular markers . 14 2.3 Phylogenetic analyses . 15 2.4 Divergence time estimation . 17 2.5 Coevolutionary analyses . 17 3 Results and Discussion 18 3.1 Systematics . 18 3.1.1 Molecular taxonomy of Nearctic Myrmica . 18 3.1.2 Phylogeny and systematics of Myrmica . 19 3.1.3 The tribe Myrmicini . 20 3.2 Biogeography . 21 3.2.1 Myrmicini . 21 3.2.2 Diversification and dispersal of Myrmica . 21 3.3 Social parasitism . 23 3.3.1 Satellite genera . 23 3.3.2 Applicability of Emery's rule . 23 3.3.3 Testing cophylogeny between parasitic Maculinea butterflies and Myrmica hosts . 24 4 Conclusions and future directions 26 5 Acknowledgements 28 6 Contents Phylogenetics of Myrmica ants and their social parasites 1 Introduction have been developed for objects ranging from elementary particles to mathema- 1.1 The art of knowing what tical theories and from minerals to orga- surrounds us nisms. The field of biology called syste- matics may be seen as a crystallisation of \Nur der Satz hat Sinn; nur im our continuous and universal ambition Zusammenhang des Satzes hat ein to know and interpret the world. It thus Name Bedeutung." forms the corner stone of biological in- | L. Wittgenstein, Tractatus formation; any other biological research logico-philosophicus, 3.3 field benefits from systematics through One of the major hominid innovations the retrieval and accumulation of infor- was undoubtedly the ability to use ab- mation. It is no coincidence that the stract language. It allows individuals to National Science Foundation is coordi- refer, to describe and to exchange infor- nating a tremendous effort to catalogue mation. Efficient communication is im- and annotate all life on Earth in the possible without a framework of words, form of the Tree of Life { systematics is phrases and concepts, shared between the cement of biology. the two partners, which guarantees that they refer to the same reality, and mu- 1.2 Systematics tually understand the outcome of the verbal exchange. All references pertai- 1.2.1 Taxonomy of systematics ning to one object are assembled un- der a common name, which can thus be Systematics may generally be divided considered as the fundamental building into two subdisciplines: taxonomy and block of information. Classes are erected phylogenetics. While the former is to store properties that describe more mainly concerned with the description than one object. of new species, classification and iden- As knowledge becomes more com- tification (focused on morphology), the plex, the need for abstraction grows, latter aims to infer the evolutionary re- necessitating generalised classes that lationships among organisms. themselves consist of classes. This, in Taxonomy is by far the older of the turn, goes hand in hand with the adop- two systematic disciplines; species have tion of hierarchies and the introduction been described since the earliest times. of classification rules to frame new data The first formal rules were established within the existing scheme. With such by Carolus Linnaeus, who introduced multi-layered knowledge systems, infor- the binomial nomenclature which is still mation can be efficiently retrieved, ex- in use today. In the 18th and 19th changed, and updated. The omnipres- century many more specific and subspe- ence of multilevel organisation in our cific variations were discovered, which daily perceptions and actions { think of led to additional, cumbersome taxono- libraries or catalogues { testifies the cen- mic terms such as subspecies and vari- tral role of conceptual arrangements in eties. In the 20th century many of these the human mind. forms were synonymised, leading to a Complex organisational schemes history of obscurity and disagreements. Introduction 7 Phylogenetics of Myrmica ants and their social parasites Traditional taxonomy is a time con- 1.2.2 Molecular taxonomy suming process, consisting of the orde- ring of large samples according to si- Tautz et al. (2003) introduced DNA ta- milarity, and the identification of con- xonomy as a possible solution to the sistent, defining characters and/or mea- taxonomic crisis. This methodology is surements. Subsequent corroboration of a new approach to taxonomy in which the initial idea is then sought in other the traditional methods for species dis- data (geography, ecology, physiology...) covery, description and identification are and by comparison to reference material. complemented and eventually replaced This may lead to new descriptions, revi- by a scheme, in which a short DNA frag- sions or synonymisations. Finally, newly ment is extracted, amplified and com- described species need to be deposited in pared against a reference database for a publicly accessible reference collection; identification. Hebert et al. (2003a) type specimens thus form the core of ta- called this process DNA barcoding. xonomy. These need to be managed and The proponents of barcoding invoke stored in museums, which is impractical several arguments to explain the need and incurs a significant cost. for alternative methods of species dis- Although taxonomy was the primary covery (Hebert et al., 2004, 2003b; Smith form of systematics for a long time, in- et al., 2005; Tautz et al., 2003). Tra- terest in it has been declining during ditionally defined taxa are the pro- the 20th century. There are many rea- duct of the opinion of one or a few sons for this situation: (i) a dusty image; specialists, and thus highly subjective. (ii) unpopularity of morphological work The similarity of morphological charac- and species description; (iii) loss of spe- ters can be misleading owing to paral- cialised taxonomic knowledge after re- lelism, convergence, analogy or mimicry. tirement of taxonomists; (iv) the rise Conversely, intraspecific variability, es- of molecular data; (v) funding difficul- pecially in morphometrics, may make ties; (vi) lack of scientific appreciation species delineations unstable. Diagnos- for species descriptions and publication tic characters may be unavailable for in idiosyncratic journals. cryptic species, and morphologically dis- Paradoxically, in the age of uncon- tinct life stages cannot be identified trollable, anthropogenically induced loss using adult morphology (Hebert et al., of diversity { a glance at the IUCN Red 2004; Monaghan et al., 2005). list for 2008 reveals that no less than Barcoding, the proponents argue, 38% of living beings on earth are cri- overcomes these obstacles.
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