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Biodiversity of the Snakes in Northern and Western Mountains of Iran, With

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Biodiversity of the Snakes in Northern and Western Mountains of Iran, With Biodiversity of the Snakes in Northern and Western Mountains of Iran, with Special Emphasis on Biodiversity in Colubroids Mahdi Rajabizadeh Thesis submitted to obtain the degree of Academiejaar 2013 - 2014 Doctor in Science (Biology) Rector: Prof. Dr. Ann Depaepe Proefschrift voorgedragen tot het Dean: Prof. Dr. Herwig Dejonghe Bekomen van de graad van Doctor in de Wetenschappen (Biologie) Promotor : Prof. Dr. Dominique Adriaens Cover illustration: A conceptual inference from the ability of modern science to give a proper picture of the complexity within the living organisms. Reading* and Examination Committee Prof. Anthony Herrel (MNHN, Paris + WE11)* Dr. Danny Meirte (Africa Museum, Belgium)* Dr. Zoltan Nagy (KBIN, Belgium)* Prof. Luc Lens (Ghent University, Belgium) Prof. Dominique Adriaens (Ghent University, Belgium) Prof. Frank Pasmans (Ghent University, Belgium) Dr. Tom Geerinckx (Ghent University, Belgium) I dedicate my dissertation to Dr. Mahmoud Latifi his great book, snakes of Iran, inspired me to study about biodiversity of the snakes in Iran Acknowledgement The success and final result of this thesis required a lot of guidance and assistance from many people and I am extremely fortunate to have got this all along the completion of my research course. Whatever I have done is only due to such guidance and assistance and I would not forget to thank them. I take this opportunity to express my profound gratitude and deep regards to my promoter Prof. Dr. Dominique Adriaens, for his exemplary guidance, supports and constant encouragement throughout the course of this thesis. Without him and his kind supports I could never accomplished this thesis. His great guidance, statistical comments, very detailed scientific discussions and patience to read and correct my drafts enabled me to complete my PhD dissertation. One simply could not wish for a better or friendlier supervisor. Thank you my dear professor. I am obliged to the research staff of the International Center for Science, High Technology and Environmental Science (ICST). Most of this research were supported by ICST research grants. I am very thankful to the president of ICST, Prof. Dr. MJ. Khanjani and official managers, Dr. Naseri, Dr. Alizadeh, Dr. Yaghubi, Dr. Bordbar and Dr. Kalantary. My sincere thanks go to all the personnel of ICST specially my friend and colleague, A. Yazdanpanah. I am grateful for their cooperation during the period of my assignment. I owe sincere and earnest thankfulness to my scientific colleagues, Prof. Dr. Nasrullah Rastegar-Pouyani, prof. Dr. Hamid Reza Esmaeili, Prof. Dr. Göran Nilson, Josef Schmidtler, Dr. Nikolai Orlov, Prof. Dr. Natalia Ananjeva, Dr. Nikolaus Stümpel, Prof. Dr. Ulrich Joger, Dr. Bahram Kiabi, Dr. Majid Kaboli, Dr. Haji Gholi Kami and Dr. Daniel Melnikov. I am also deeply indebted to my colleagues and friends in the field expeditions, specially my dearest friends Dr. Hiva Faizi and Dr. Roman Nazarov and my colleagues, Fariborz Heidary, Amin Ghadami, Soheila Javanmardi, Ruzbeh Behruz, Masoud Yousefi, Bahman Zangi and Mohammad Jahan. My sincere thanks go to my colleagues in the Evolutionary Morphology of Vertebrates research group, especially Tom Geerinckx, Barbara De Kegel, Joachim Christiaens and Hilde Van Wynsberge for their kind cooperation. I am profoundly indebted to the members of my PhD jury: Prof. Dr. Anthony Herrel, Dr. Danny Meirte, Dr. Zoltan Nagy, Prof. Dr. Luc Lens, Prof. Dr. Frank Pasmans and Dr. Tom Geerinckx. I gratefully acknowledge the kind cooperation of the following scientists for loan of the specimens or providing valuable data about specimens out of Iran: Michael Franzenand and Frank Glawin in ZSM, Patrick Campbell and Barry Clarke in BMNH, Roman Nazarov and Valentina Orlova in ZMMU, Raffaeil Ernst in MTKD, Aziz Avci in Adnan Menderes Üniversitesi, Boris Tuniyev in Sochi National Park Scientific Department and Konstantin Milito in ZISP. Last but not the least, I would like to appreciate my family for their unending support, encouragement and prayers. Really words fail me to express my deepest appreciation to my Mother. I am grateful to my Financee, Jaleh, and thanks to her family for supporting me. Through preparing this thesis, Jaleh was always beside me and motivated me all the time. Finally, I would like to thank everybody who was important to the successful realization of this thesis, as well as expressing my apology that I could not mention them personally one by one. Table of contents Chapter 1, Introduction 1-1- General context 1 1-1-1- Evolution of snakes 1 1-1-2- Snake phylogeny and classification 2 1-1-3- Global matters about snake biodiversity 4 1-1-4- Species concept 5 1-1-5- Subspecies concept 7 1-1-6- Species delimitation and integrative taxonomy 7 1-1-7- Cryptic biodiversity in snakes 8 1-1-8- Biodiversity of snakes in Iran 9 1-2- Objectives and thesis outlines 12 1-2-1- General objectives 12 1-2-2- Outline of the thesis 13 1-2-2-1- chapters three and four 13 1-2-2-2- chapter five 13 1-2-2-3- chapters six, seven and eight 13 1-2-2-4- The general discussion 14 1-3- Background information about Iran 14 1-3-1- Physical geography of the northern and western mountains of 15 Iran 1-3-2- Geology of Iran 16 1-3-3- Climate of Iran 17 1-3-4- Vegetation of Iran 20 Chapter 2, Materials and Methods 2-1- Study area 22 2-2- Samplings and collections examination 23 2-3- Taxonomic studies 24 I 2-4- Analyses 26 2-4-1- Traditional biometry and morphological analysis 26 2-4-2- Geometric morphometric and geometric analysis 30 2-4-3- Molecular analysis 31 2-4-4- Selection of operational taxonomic units 32 2-4-5- Criteria used for species delimitation 32 References of both chapters 1 and 2: 34 Chapter 3, Analysis of Dorsal Color Pattern Variation in the Montivipera raddei Species Complex Abstract 43 3-1- Introduction 44 3-1-1- Distribution and taxa within the Montivipera raddei species 45 complex 3-1-2- Environmental factors 46 3-2- Materials and Methods 47 3-2-1- Voucher specimens 47 3-2-3- Environmental factors 52 3-3- Analysis 55 3-4- Results 55 3-5- Discussion 56 References 59 Chapter 4, Review of Taxonomy and Distribution of the Montivipera raddei Species Complex Abstract 63 4-1- Introduction 64 4-2- Materials and Methods 64 4-3- Results 66 4-3-1- Multivariate and univariate comparison of OTUs 68 4-4- Discussion 75 II 4-4-1- Montivipera kuhrangica (Rajabizadeh, et al., 2011) 78 4-4-1-1- Holotype 78 4-4-1-2- Terra typica 78 4-4-1-3- Diagnosis 79 4-4-1-4- Description of holotype 79 4-4-1-5- Additional specimen 83 4-4-1-6- Habitat 83 4-4-1-7- Conservation 84 References 85 Chapter 5, Geographic Variation, Distribution and Habitat of the Dice Snake (Natrix tessellata) in Iran Abstract 87 5-1- Introduction 88 5-2- Materials and Methods 89 5-3- Results 90 5-3-1- Construction of Operational Taxonomic Units (OTUs) 91 5-3-2- Multivariate and univariate comparison of OTUs 93 5-3-3- Intra-OTUs variation 99 5-3-4- Distribution 101 5-3-6- Habitat 103 5-4- Discussion 106 5-4-1- Distribution of Natrix tessellata in Iran 106 5-4-2- Habitat of Natrix tessellata in Iran 107 5-4-3- Geographic Variation of Natrix tessellata in Iran 108 5-4-4- Intra-OTU variation 109 5-4-5- Taxonomic comments 110 5-4-6- Biogeographic aspect 112 References 114 III Chapter 6, Taxonomy, Distribution and Geographic Variation of Eirenis punctatolineatus Abstract 118 6-1- Introduction 119 6-2- Materials and Methods 121 6-3- Results 122 6-3-1- Color and pattern 122 6-3-2- Construction of Operational Taxonomic Units (OTUs) 125 6-3-3- Multivariate and univariate comparison of OTUs 125 6-4- Discussion 131 6-4-1- Distribution 131 6-4-2- Taxonomical and geographical variations 132 6-4-3- Eirenis (Pediophis) punctatolineatus punctatolineatus (Boettger, 132 1892) 6-4-4- Eirenis (Pediophis) punctatolineatus condoni (Boulenger, 1920) 133 new combination References 135 Chapter 7, Review of Taxonomy and Distribution of the Eirenis medus Species Complex Abstract 138 7-1- Introduction 139 7-2- Materials and Methods 139 7-3- Results 143 7-3-1- Distribution and morphology of the Eirenis medus species 143 complex 7-3-2- Morphological comparison 143 7-3-3- Taxonomic account 146 7-4- Eirenis (Pediophis) kermanensis (Rajabizadeh et al., 2012) 146 7-4-1- Holotype 146 7-4-2- Paratype 146 IV 7-4-3- Diagnosis 146 7-4-4- Description of holotype 147 7-4-5- Variation 148 7-4-6- Etymology 148 7-4-7- Habitat 148 7-4-8- Comparison and relationships 149 References 150 Chapter 8, Taxonomy of Eirenis (Pseudocyclophis) persicus Abstract 154 8-1- Introduction 155 8-2- Materials and Methods 157 8-3- Results 163 8-3-1 Construction of Operational Taxonomic Units (OTUs) 163 8-3-2- Multivariate and univariate comparison of OTUs 164 8-3-3- Intra-OTU variation 167 8-3-4- Geometric morphometric analysis 173 8-3-5- Molecular analysis 176 8-4- Discussion 178 References 181 Chapter 9, General Discussion 9-1- Taxonomy of the examined species of the genera Eirenis, Montivipera 184 and Natrix in Iran. 9-1-1- Taxonomy of Montivipera raddei species complex 185 9-1-2- Taxonomy of Natrix tessellata 186 9-1-3- Taxonomy of examined species of genus Eirenis 187 9-1-3-1- Taxonomy of Eirenis medus species complex 188 9-1-3-2- Taxonomy of Eirenis punctatolineatus 189 9-1-3-3- Taxonomy of Eirenis persicus species complex 190 V 9-2- Biogeography of the examined species of genera Eirenis, Montivipera 191 and Natrix in Iran.
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