Taxonomic and phylogenetic relationships of freshwater crabs of the genus Potamon (Crustacea: Brachyura: Potamidae) from Iran DISSERTATION ZUR ERLANGUNG DES DOKTORGRADES DER NATURWISSENSCHAFTEN (DR. RER. NAT.) DER FAKULTÄT FÜR BIOLOGIE UND VORKLINISCHE MEDIZIN DER UNIVERSITÄT REGENSBURG vorgelegt von Alireza Keikhosravi aus Sabzevar, Iran im Jahr 2013 Das Promotionsgesuch wurde eingereicht am: 30.10.2013 Die Arbeit wurde angeleitet von: PD Dr. Christoph Schubart Unterschrift: ii Dedication This work is dedicated to: My wife, Maryam for encouragement, patience and continuous support My little, lovely daughter, Ariana My parents, who their love and sacrifices made possible for me to accomplish the education My advisor, PD Dr. Christoph Schubart for his fatherly support iii Abstract of the Dissertation Taxonomic and phylogenetic relationships of freshwater crabs of the genus Potamon (Crustacea: Brachyura: Potamidae) from Iran by Alireza Keikhosravi University of Regensburg In an era of climate change and habitat destruction, assessing current biodiversity as soon as possible is of crucial importance. To uncover the real biodiversity, taxonomic problems in species complexes and phylogenetic delimitations of species need to be clarified. Potamon is a well-known freshwater crab genus with a distribution from the Middle East through southern Europe and the far north of Africa. Phylogeny of Potamon has been a source of disagreement among specialists over the last few decades. In order to understand the evolution of the group it is crucial to resolve the phylogenetic relationships among the species of Potamon and to discover intraspecific endemic units using different tools. The purposes of the current study were: 1) to reconstruct the phylogenetic history of different species of the genus Potamon in Iran, 2) to find out, if there is consistency of morphological with molecular characters, 3) to determine intraspecific molecular variability among populations of Potamon elbursi in order to understand initial steps of regional differentiation as well as importance of endemic forms for conservation issues. In the course of the study two mitochondrial (16S rRNA and Cox1) and one nuclear (28S rRNA) genes were partially amplified and sequenced. A significant divergence was shown among three distinct lineages previously considered to belong to Potamon persicum complex and associated with their geographical distributions. One lineage corresponds to a formerly synonymised species, P. elbursi, which is here revalidated and redescribed. Another lineage (western Iran) still must be considered a cryptic lineage, since it is genetically distinct, but the current morphological evidence cannot separate it from P. persicum. Consistent molecular and morphological divergence was also recorded from the westernmost slopes of the Zagros Mountains where Potamon ilam, a sister species to Potamon mesopotamicum, is described as a new species to science. Overall, our results show that Potamon radiated at different time intervals in the Middle East, leaving a genetic footprint of historical iv differentiation processes and following successive isolation within different watersheds. The current locally confined and unique biological diversity of Iran can be explained mainly by different topography and climate regimes resulting from mountains uplifting. Genetic assessment of populations of P. elbursi elucidates a significant genetic differentiation among populations without any linkage with geographical dispersal. Some populations show high endemicity, resulting from strong genetic drift or geographic barriers. Endangered aquatic ecosystems and occurrence of genetically distinct endemic and isolated populations of P. elbursi, call for considering the Alborz Mountains for possible future conservation plans. v Acknowledgments I express my sincere appreciation to my advisor, PD Dr. Christoph Schubart, for his endless patience, and fatherly support and guidance thorough my four and half years of graduate studies. Further thanks are due to my PhD mentors, Prof. Dr Christoph Oberprieler and Dr. Sara Fratini, for their comments and beneficial advice in course of my PhD. My thank also goes to Prof. Jürgen Heinze (University of Regensburg) and his staff for the use of the necessary facilities and continuing support. My visit to Natural History Museum, Wien (NHMW); Natural History Museum, London (NHM) was funded by the “Synthesis” Project of the European Union, which is greatly appreciated. I am grateful to Dr. Jørgen Olesen (ZMUC), Dr. Paul Clark (NHM), Prof. Michael Türkay (SMF), Prof. Peter K. L. Ng (ZRC), Dr. Peter Dworschak and Dr. Alireza Sari (ZUTC) for loaning me material from their collections in course of my PhD. My thanks also go to my uncle Mr. Esmaeil Keikhosravi for his great help in collecting specimens. It also goes to other people who accompanied me with sampling: Fatollah Houshmand, Behzad Fathinia, Morteza Zarei and Madjid Moradmand. Dr. Mehdi Abedi deserves special praise for hosting me several months and providing much needed support, help and advice. Dr. Reza Naderloo, Dr. Catia Bartilotti, Dr. Ivana Silva, Nicolas Thiercelin, Jürgen Trettin for all sort of support, including troubleshooting, brainstorming and friendship. Warm thanks also go to Peter Koller, Nicole Rivera, Adnan Shahdadi, Claudia Laurenzano, Masaki Suefuji, Marion Füßl and Manuel Stemmer. I express my gratitude to Mrs. Christiane Wanke and Doris Rothgaenger, for helping to make a comfortable situation for my family while they lived here. I am deeply indebted to my parents-in-law, Mr. Yadollah Houshmand and Mrs. Kobra Keikhosravi for supporting my family. I also wish to thank my brother and sisters for their support, their understanding, their power to arm me with new resources at each of my visits. I appreciate the Akademisches Auslandsamt of the University of Regensburg for financial support. vi Table of Contents Chapter 1 .................................................................................................................................. 1 General Introduction .............................................................................................................. 1 What are freshwater crabs? .................................................................................................... 1 Behavioral Ecology of freshwater crabs ................................................................................ 2 Freshwater crabs and human life ........................................................................................... 5 Conservation perspective of freshwater crabs ....................................................................... 5 Biogeographic importance of freshwater crabs ..................................................................... 6 Taxonomy and diversity of freshwater crabs ......................................................................... 7 Origin of freshwater crabs ................................................................................................... 10 Genus Potamon .................................................................................................................... 11 The aim of the present work ................................................................................................ 12 Chapter 2 ................................................................................................................................ 14 Introduction ........................................................................................................................ 15 Material and Methods ......................................................................................................... 15 Taxonomy ............................................................................................................................ 18 Molecular results ................................................................................................................. 24 Discussion ............................................................................................................................ 30 Acknowledgements ............................................................................................................. 32 Chapter 3 ................................................................................................................................ 33 Introduction ........................................................................................................................ 34 Experimental Procedures .................................................................................................... 35 Morphology ........................................................................................................................ 38 Systematic Account ............................................................................................................ 39 Molecular results ................................................................................................................. 46 Discussion ........................................................................................................................... 48 Acknowledgements ............................................................................................................ 50 Chapter 4 ................................................................................................................................ 51 Introduction ........................................................................................................................