Molecular phylogeny, phylogeography and taxonomic revision of species of the genus Perisesarma De Man, 1895 (Crustacea: Decapoda: Brachyura: Sesarmidae) DISSERTATION ZUR ERLANGUNG DES DOKTORGRADES DER NATURWISSENSCHAFTEN (DR. RER. NAT.) DER FAKULTÄT FÜR BIOLOGIE UND VORKLINISCHE MEDIZIN DER UNIVERSITÄT REGENSBURG vorgelegt von Adnan Shahdadi aus Gevin, Hormozgan, Iran im Jahr 2017 i Das Promotionsgesuch wurde eingereicht am: 25.04.2017 Die Arbeit wurde angeleitet von: Prof. Dr. Christoph Schubart Unterschrift: ii This work is cordially dedicated to: My family; My parents, who devoted their life to their children, My patient wife and my lovely son for making a sweet home, My brothers and my sisters for their continuous countless supports iii Abstract of the Dissertation Molecular phylogeny, phylogeography and taxonomic revision of species of the genus Perisesarma De Man, 1895 (Crustacea: Decapoda: Brachyura: Sesarmidae) by Adnan Shahdadi University of Regensburg The present work forms part of an ongoing revision of sesarmid genera. The conspicuous and speciose genus Perisesarma De Man, 1895 is among the most taxonomically complex and problematic taxa of the family. This genus was suspected to be a polyphyletic taxon, comprising morphologically heterogeneous species. Moreover, some members of the genus have a close phylogenetic relationship with the sister genus Parasesarma De Man, 1895 and show many morphological similarities, except for the existence of an epibranchial tooth in Perisesarma. Therefore, the present study intends to evaluate the monophyly of Perisesarma and reconstruct phylogenetic relationships in comparison to Parasesarma by examining a variety of morphological characters and comparing them with results from molecular markers. Up to four genetic marker with different evolutionary mutation rates were used for different parts of this study. These include the mitochondrial genes Cox1, 16S, ND1 and the nuclear gene NaK. A comparative morphological analysis reveals that there is no unequivocal separation between species of Parasesarma and Perisesarma, because of intermediate conformations of the epibranchial tooth. In our molecular analysis, most species of Perisesarma cluster solidly with species of Parasesarma, but without being reciprocally monophyletic. Morphology and genetics also indicate that the West African species of Perisesarma and P. fasciatum are markedly different from all other species of the genus. Therefore, we here suggest with robust double support, a new classification, transferring most species of Perisesarma to Parasesarma and the iv three West African representatives and P. fasciatum into two new genera, thus restricting the genus Perisesarma to the type species P. dusumieri. This study also uncovers some un-described forms of the former genus Perisesarma and presents new insight in to the phylogeography of some phylogenetic clades. Perisesarma n. sp1. is described from mangroves of southern Vietnam, differing most significantly from congeners by the tuberculation pattern of the chelar dactylus and its unique G1 morphology. Genetically, Perisesarma n. sp1. is markedly divergent from other congeneric species, both in mitochondrial and nuclear DNA. This study helps to reveal marine biogeographic barriers with restricted gene flow, among them a putative barrier between the northern Australian coastline and adjacent areas of South East Asia and the South Pacific. In agreement with these findings, we provide evidence for genetic uniqueness of representatives of the mangrove crab genus Parasesarma from northern Australian mangroves, based on three mitochondrial and one nuclear DNA marker. This distinct taxon is here described as a new species. Morphologically the new species is very similar to P. lividum from the southwest Pacific and P. samawati from East Africa. Genetically, however, it is significantly distinct from all other congeners. The same genetic disjunction is found between the two very closely related species P. semperi from S.E Asia and P. longicristatum from Australia. P. indiarum, was originally described from Ambon, Indonesia, and is assumed to be distributed all over S. E. Asian Island. But here presented genetic and morphological evidences reveal a distinct separation of Malay Peninsular populations from the types and therefore these are herein described as a new species. v Acknowledgments I express my sincere appreciation to my advisor, Prof. Dr. Christoph Schubart, for his cordiality, support and guidance thorough my graduate studies. Further thanks are due to my PhD mentors, Prof. Dr. Christoph Oberprieler (University of Regensburg) and Prof. Dr. Roland Melzer (LMU Munich), for their comments and beneficial advice in course of my PhD. My thanks also go to Prof. Jürgen Heinze (University of Regensburg) and his staff for the use of the necessary facilities and continuing support. Dr. Alireza Keikhosravi (University of Sabzevar, Iran) deserves special praise for his great support, help and advice. I deeply thank my friends Theodor Poettinger, Dr. Nicolas Thiercelin and Dr. Katharina von Wyschetzki for all sort of support, including troubleshooting and brainstorming. I am indebted to many institutions and colleagues which/who facilitated specimens, allowed access to samples or helped in the field. Among those our special thanks go to Peter J.F. Davie from the Queensland Museum in Brisbane (Australia); to Peter K.L. Ng, Lee Bee Yan, Chen Zijian Paul, Jose Christopher E. Mendoza and Muhammad Dzaki Bin Safaruan from the National University of Singapore (Singapore); to the late Michael Türkay, Moritz Sonnewald and Bianca Trautwein from the Senckenberg Museum Frankfurt (Germany); to Charles Fransen, Karen van Dorp and Kees Van Den Berg from the Naturalis Museum in Leiden (Netherlands); to Paul Clark, Miranda Lowe and Harry Taylor from the Natural History Museum in London (UK); to Sara Fratini and Gianna Innocenti from the University of Florence (Italy); to Tohru Naruse from the Ryukyu University Museum, Fujukan (Japan); to Laure Corbari from the Muséum National d’Histoire Naturelle, Paris (France); Tomoyuki Komai from the Natural History Museum and Institute Chiba (Japan); to Stephen Keable, Helen Stoddart, Alex Hegedus and Sue Lindsay from the Australian Museum in Sydney; and to Dwi Listyo Rahayu, Sivasothi, Estelle Roux, Pierre Armand Mvogo-Ndongo, Peter Koller and Niko Ramisch for collecting additional material. vi My visits to the Naturalis Museum in Leiden (Netherlands) (NL-TAF), Natural History Museum in London (U.K.) (GB-TAF), Muséum National d’Histoire Naturelle, Paris (France) (FR-TAF) received support from the SYNTHESYS Project (http://www.synthesys.info/), which is financed by European Community, Research Infrastructure Action under the FP7 “Capacities” and my visits to the Australian Museum in Sydney and Queensland Museum in Brisbane received support from the international PhD program at Universität Regensburg (iPUR). Both institutions are greatly appreciated. I express my gratitude to Dr. Narjes Javadi Motaghi, Dr. Heda Hosaini and his kind wife, Nikoo, to Dr. Marion Füßl, Linda Dangler and her friendly parents, for helping to make a comfortable situation for me and my family while living in Regensburg. Warm thanks also go to Dr. Catia Bartilotti, Dr. Ivana Silva, Dr. Jürgen Trettin, Dr. Abel Bernadou, Dr. Tomer Czaczkes, Andereas Trindl, Christiane Wanke, Nathalie-Nida Moske, Katharina Gryska, Michael Roehrl and Elahe Parvizi for help and friendship. I am being sponsored by the Iranian Ministry of Science, Research and Technology during my PhD studies at the University of Regensburg. vii According to INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE (ICZN), Article 8.3, here I state that: "This work is not issued for the purpose of zoological nomenclature" viii Table of Contents Chapter 1 General Introduction .......................................................................................................................1 Chapter 2 Evaluating the consistency and taxonomic importance of cheliped and other morphological characters that potentially allow identification of species of the genus Perisesarma De Man, 1895 (Brachyura, Sesarmidae) ......................................................................................................17 Abstrat............................................................................................................................................17 Introduction ...................................................................................................................................18 Material and Methods ...................................................................................................................19 Results and Discussion .................................................................................................................23 Conclusions ...................................................................................................................................25 Chapter 3 Perisesarma n. sp1., a new species of mangrove crab from Vietnam (Decapoda, Brachyura, Sesarmidae), with an assessment of its phylogenetic relationships...............................................32 Abstract .........................................................................................................................................32 Introduction ...................................................................................................................................33 Material and Methods