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Phylogeny of the Symphyla (Myriapoda)

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Phylogeny of the Symphyla (Myriapoda) Phylogeny of the Symphyla (Myriapoda) Inaugural-Dissertation to obtain the academic degree Doctor rerum naturalium (Dr. rer. nat.) submitted to the Department of Biology, Chemistry and Pharmacy of Freie Universität Berlin by Miguel Domínguez Camacho from Madrid (Spain) December 2009 Time period: October 2006 - December 2009 Supervision by Dr. Markus Koch Institute of Biology, Freie Universität Berlin 1. Gutachter: Prof. Dr. Klaus Hausmann 2. Gutachter: Prof. Dr. Thomas Bartolomaeus Date of defence: February 12th 2010 Scolopendrellopsis (Symphylellopsis) subnuda Acknowledgment I would like to thank everybody who has made it possible to carry out this project. I am grateful for the generous supply of specimens and soil samples (as appears in the list in the first chapter). Antoni Serra (UB), Eduardo Mateos (UB) and David Carreras (Observatori Socioambiental de Menorca) helped with valuable recommendations on sampling sites in Catalonia and Minorca. Queralt Castañares and Jónathan Pérez provided technical support during the field trips. Discus- sion with María Teresa Domínguez and Ulf Scheller aided the identification of specimens. Ulf Scheller also helped with critical reviews and kindly providing literature. Scientific support in Madrid by Carlos Simón (UAM) and Carmen Gutiérrez (CSIC) is also acknowledged. I would like to extend my thanks to Antoni Serra for scientific support in Barcelona and multiple valu- able collaborations. Thanks are also due to Gerald Beatty, my English teacher, for corrections and linguistic help during the writing of the manuscript. I am also grateful for their scientific support in Berlin to Prof. Dr. Thomas Bartolomaeus, Dr. Thomas Stach and Prof. Dr. Klaus Hausmann. All the people of the AG15 “Systematik und Evolution der Tiere” of the FU-Berlin have helped me in countless situations throughout the investigation. They have kindly assisted me always when I required them in the laboratory, with software and with paperwork. Moral support has been also decisive during this research period. For this reason, I would like to thank all my friends and family for their support. Special thanks are due to my parents, Fernando and Irene, and to my brother, Marcos, who have always believed in my passion for Zoology. I have shared the marvellous experience of living these three years in Berlin with Belén Benito, a very special person for me. I extend my gratitude to the promoters of this project, Dr. Markus Koch and Prof. Dr. Thomas Bartolomaeus, without whom this project would not has been possible. Above all, I would sincerely like to thank Dr. Markus Koch, who has been my supervisor during the whole investigation. He has taught me all the methodology very patiently and given me a masterful introduction to the fantastic world of the evolution of the arthropods. This project was financed by the Fundación “la Caixa” and the DAAD (German Academic Exchange Service). Financial support for the field trips was also provided by the FU-Berlin. 5 Contents General introduction .....................................................................................7 Chapter 1. Contribution to the taxonomy of the European Symphyla ..........15 Introduction .................................................................................................................15 Material and methods 1. Material studied ...................................................................................................16 2. Field trips and soil fauna extraction .....................................................................17 3. Microscopy ..........................................................................................................18 4. Terminology ........................................................................................................19 Results and discussion 1. Resolution of different microscopy methods ........................................................19 2. List of species with remarks on the taxonomy .....................................................19 3. Key to the European genera of Symphyla ............................................................45 Literature ....................................................................................................................46 Chapter 2. Comparative skeletomuscular anatomy of the head ....................49 Introduction .................................................................................................................49 Material and methods ..................................................................................................51 Results ........................................................................................................................51 Discussion 1. The tentorial complex 1.1. Common pattern and variations within Symphyla .......................................85 1.2. Clarification of misunderstandings in the literature .....................................86 1.3. True “fultural sclerites” in Symphyla ..........................................................86 1.4. Tentorial mechanisms and phylogenetic implications ..................................87 2. Musculature of the head 2.1. More primitive musculature in Scolopendrellidae .......................................89 2.2. Outgroup comparison ..................................................................................91 2.3. Mandibular mechanisms .............................................................................94 Literature ....................................................................................................................94 6 Chapter 3. Morphological phylogenetics of the Symphyla ..........................97 Introduction .................................................................................................................97 Material and methods 1. Examined taxa ...................................................................................................100 2. Morphological characters ..................................................................................101 3. Cladistic analysis ...............................................................................................102 Results ......................................................................................................................103 Discussion 1. Incongruence between phylogeny and the traditional classification ...................107 2. Character evolution 2.1. Number of the scuta ..................................................................................109 2.2. Morphology of scuta .................................................................................111 2.3. First pair of legs ........................................................................................112 2.4 Spiracles and tracheae ................................................................................113 2.5. Mouthparts and feeding mechanisms ........................................................113 Conclusions: Evolution of the Symphyla 1. Phylogenetic interrelationships of Symphyla .....................................................115 2. Biogeographical aspects in dating clades ...........................................................115 3. Taxon sampling and the symphylan ground pattern ............................................116 4. Evolutionary history of the Symphyla ................................................................116 Literature ..................................................................................................................117 Apendix I ..................................................................................................123 Apendix II .................................................................................................137 Summary ...................................................................................................139 Zusammenfassung .....................................................................................141 Curriculum Vitae .......................................................................................143 7 General Introduction The Symphyla are the smallest subgroup of the Myriapoda. They are minute centipede-like animals (with an average length of 1-8 mm), representatives of the soil fauna. Their soft, flexible and unpigmented body is suited to subterranean life in soil (Verhoeff 1934). They also present other typical adaptations to subterranean life habits, such as the lack of eyes, the presence of long monoliform antennae and numerous mechanoreceptors, chemoreceptors and hygroreceptors. However, they can leave soils, and sometimes occur in leaf litter, under bark and moss or inclusively on trees in tropical regions (Adis & Scheller 1984). Moisture seems to be the most important factor determining their habitat (Waterhouse 1968). They are presumably a geologically old group (Scheller & Adis 2002). Their minimal age has been dated from the mid Silurian, about 430 million years (Edgecombe 2004), although the only fossil symphylans are known from amber with an age of 25-50 million years (Poinar & Edwards 1995, Scheller & Wunderlich 2004). Their biogeography agrees with their old estimated age, since they have a worldwide distribution which varies among the different genera. Some seem to have a typical Gondwanan disjunctive distribution (table 1). Although the fact that symphylans can be found throughout the world, they have been relatively neglected in biogeographical studies, and one of the main reasons is
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