Phylogeny, character evolution and species diversity in Crambinae, Heliothelinae and Scopariinae DISSERTATION zur Erlangung des akademischen Grades DOCTOR RERUM NATURALIUM (Dr. rer. nat.) vorgelegt dem Bereich Mathematik und Naturwissenschaften der Technischen Universit¨at Dresden von M. Sc. Th´eo Antonin Baptiste L´eger geboren am 02.01.1989 in Lausanne, Schweiz eingereicht am 10. Januar 2020 Die Dissertation wurde in der Zeit von 09/2014 bis 12/2019 an den Senckenberg Naturhistorischen Sammlungen Dresden angefertigt. ii Charles Darwin, letter to T. H. Huxley, 26 September 1857 Darwin Correspondence Project, “Letter no. 2143”, https://www.darwinproject.ac.uk/letter/DCP-LETT-2143.xml 1. Gutachter 2. Gutachter Prof. Dr. Christoph Neinhuis Prof. Dr. Niklas Wahlberg Lehrstuhl f¨ur Botanik, Systematic Biology Group Fakult¨at Mathematik und Faculty of Science Naturwissenschaften Lund University Technische Universit¨at Dresden S¨olvegatan 37, Lund Dresden, Deutschland Schweden Declaration Erkl¨arung gem¨aß § 5.1.5 der Promotionsordnung Hiermit versichere ich, dass ich die vorliegende Arbeit ohne unzul¨assigeHilfe Dritter und ohne Benutzung anderer als der angegebenen Hilfsmittel angefertigt habe; die aus fremden Quellen direkt oder indirektubernommenen ¨ Gedanken sind als solche kenntlich gemacht. Die Arbeit wurde bisher weder im Inland noch im Ausland in gleicher oder ¨ahnlicher Form einer anderen Pr¨ufungsbeh¨orde vorgelegt. Berlin, 14. Januar 2020 Th´eo L´eger iii iv Acknowledgements This work would not have been possible without the help and support from various people. I want to express my sincere gratitude to Matthias Nuss and Bernard Landry for introducing me to the fabulous group that represent Pyraloidea and to the thrilling field of research that is systematics. Matthias Nuss supervised me for five years and suggested most of the projects presented here. I appreciated the liberty I was given in the realisation of my projects and enjoyed our discussions that sharpened my scientific reasoning. Huge thanks goes to Bernard Landry from the Mus´eum d’histoire naturelle de Gen`eve (MHNG) for suggesting and co-supervising my phylogenetic work on the Crambinae and Scopariinae. I am very thankful to my thesis director Christoph Neinhuis for accepting to supervise my work and guide me through the administrative standards of the Technische Universit¨atDresden (TUD). The supportive and entertaining atmosphere at the Senckenberg Museum f¨ur Tierkunde (SMTD) greatly contributed to the accomplishment of my work. As my closest colleague, I am thankful to Richard Mally who co-authors two of the papers presented here and set up the protocols in the lab for the Lepidoptera section during his time at the SMTD. Also, having him alongside when collecting, setting moths and escaping earthquakes during our fieldtrip to Mount Kinabalu was a much appreciated and memorable experience. Special thanks are given to Manuela Bartel and Christian Kehlmaier who provided great help with lab work. Furthermore, I am indebted to Christian Schmidt, Francesca Vegliante, Franziska Bauer and Klaus Dieter-Klass for our en- tomological discussions and to Loreen Kn¨obel and Melita Vamberger for our entertaining lunch breaks. For their help concerning field expeditions, I am grateful to Dr. Stephen Sutton, Prof. Charles Vairappan (Institute for Tropical Biology and Conservation), Ching Y. Chung (Sabah Biodiver- sity Centre), the staff of the Sabah Parks and Peter Achleiner (Borneo, Malaysia); Clemente, InocˆencioJos´e, Miguel Bungo (Universidade Kimpa Vita, Ndalatando), Jos´e Lau (Universidade Kimpa Vita, U´ıge),Christoph Neinhuis, Thea Lautenschl¨ager,Christin Heinze (TUD) (Angola); Ashley Field, Daren Crayn, David Rentz and the staff of the Australian Tropical Herbarium (James Cook University, Cairns), Christoph Neinhuis and Sarah Wagner (TUD) (Queensland, Australia). David Lees and Geoff Martin from the Natural History Museum (London, UK), as well as my friends Claire Monari and Jonty Forrester are thanked for the help and entertainment graciously offered during my tight-budget stay in London. Moreover, I am really indebted to all the people who provided material for molecular investigations. I am thankful to the Senckenberg Gesellschaft f¨urNaturforschung, the Mus´eumd’histoire na- turelle de Gen`eve and the Museum f¨urNaturkunde Berlin for providing funds, as well as all other temporary employers that helped bridging the gaps. Also, I want to thank Patrick Schmitz who, on a night of May 2010, gave a presentation on endemic aquatic moths of Hawaii at “la Datcha”, the student local at the University of Geneva. That talk left me bewildered and with one thing on my mind: “That’s what I want to do!”. I also thank my friends from Dresden, Geneva and elsewhere for the leisure time beside PhD work, v vi eventhough my interest for tiny dull moths remain obscure for most of them up until now. Last but not least, I thank my parents and my sister for always being supportive of my passions and endeavours. Abbreviations Institutions BMNH: British Museum of Natural History, London BORN: Borneensis, Institute for Tropical Biology and Conservation, Kota Kinabalu, Malaysia ITBC: Institute for Tropical Biology and Conservation, Kota Kinabalu, Malaysia MHNG: Mus´eum d’histoire naturelle de Gen`eve, Switzerland SMTD: Senckenberg Museum f¨ur Tierkunde Dresden TUD: Technische Universit¨at Dresden Scientific terms BI: Bayesian Inference bp: base pair BS: Bootstrap support ESS: Effective Sample Size GTR: General Time Reversible MCMC: Markov Chain Monte Carlo ML: Maximum likelihood pp: posterior probability Phylogenetic groups CAMMSS clade: clade comprising the Crambinae, Acentropinae, Midilinae, Musotiminae, Schoenobinae and Scopariinae, as of Regier et al. (2012) OG clade: clade comprising the Odontiinae and Glaphyriinae, as of Regier et al. (2012). PS clade: clade comprising the Pyraustinae and Spilomelinae, as of Regier et al. (2012). non-PS clade: clade comprising the Odontiinae, Glaphyriinae, Crambinae, Acentropinae, Midili- nae, Musotiminae, Schoenobinae and Scopariinae as of Regier et al. (2012). Genes CAD: carbamoyl-phosphate synthetase 2, aspartate transcarbamylase, and dihydroorotase COI: cytochrome oxydase subunit I DDC: dopa decarboxylase EF-1a: Elongation Factor I alpha GAPDH: glyceraldehyde-3-phosphate dehydrogenase IDH: isocitrate dehydrogenase MDH: cytosolic malate dehydrogenase rps5: ribosomal protein S5 vii viii Abstract Pyraloidea is one of the largest superfamilies of Lepidoptera comprising 15 955 described species. The monophyly of the group was suggested by the presence of a tympanal organ in the first ab- dominal segment; it has been confirmed later by molecular data. This group is divided into Pyralidae (5750 species in 5 subfamilies) and Crambidae (10 205 species in 14 subfamilies). The work presented here provides new insights on the phylogeny and evolution of morphological and ecological characters of the Crambinae and Scopariinae, as well as the first phylogenetic place- ment, host plant record, and taxonomic work on the Heliothelinae. Crambinae (2,047 species) and Scopariinae (577 species) were previously recovered as sister groups in a molecular phylogenetic analysis of three and two species respectively, while Heliothe- linae (50 species) have never been investigated in a phylogenetic framework. The morphology- based classification divided Scopariinae into 24 genera and Crambinae into 9 tribes and 176 genera, while Heliothelinae comprised the two tribes Heliothelini and Hoploscopini. Analyses of a six-gene dataset including 110 taxa with Maximum Likelihood and Bayesian Inference confirm the monophyly of Crambinae as well as Scopariinae, while no support was found for the posi- tion of Heliothelinae. Phylogenetic results demonstrate that Scopariinae were oversplit at the generic level by traditional classification, with five genera found nested within, and consequently synonymized with the two species-rich genera Eudonia and Scoparia. Anarpia, recovered as the basal-most lineage of Scopariinae, occurs in the Mediterranean region, in contrast to the remain- ing Scopariinae found in temperate regions and wet tropical montane forests. In Crambinae, our revision of the tribal classification leads to two synonymizations, one reinstatement as well as the description of a new tribe. Furthermore, several genera are confidently assigned to a tribe for the first time and two are synonymized with Microcrambus. Ancestral character reconstructions for 27 morphological characters provide new apomorphies for several clades and reveal several mod- ifications of the female oviscapt associated with the behavior of laying adhesive or non-adhesive eggs, the latter being associated with the use of Pooidae as host plants. Inclusion of a greater number of taxa and genes comprising all subfamilies of Crambidae sup- ports the polyphyly of Heliothelinae. Heliothelinae s. str. are sister to Scopariinae, while Hoploscopinae are sister to (Heliothelinae+Scopariinae)+Crambinae. Heliothelinae s. str. occur in xerothermic habitats and larvae of Heliothela wulfeniana are recorded as miners in Violaceae. In contrast, Hoploscopinae occur in montane rainforests of the Oriental and Austral-Asian re- gions. Field observations on Mount Kinabalu and subsequent studies of the material collected provided the first host plant record for Hoploscopa and highlighted a high proportion of unde- scribed species in this genus. Following an iterative approach combining morphology and DNA barcoding from museum