Phylogenetics and systematics of North-African Geckos Tarentola Von der Fakultät für Lebenswissenschaften der Technischen Universität Carolo-Wilhelmina zu Braunschweig zur Erlangung des Grades eines Doktors der Naturwissenschaften (Dr. rer. Nat.) genehmigte Dissertation Von Ismail Mustafa Bshaena aus Tajura / Libyen 1. Referent: apl. Professor Dr. Ulrich Joger 2. Referent: Professor Dr. Miguel Vences Eingereicht am: 21.03.2011 Mündliche Prüfung (Disputation) am: 14.06.2011 Druckjahr 2011 Vorveröffentlichungen der Dissertation Teilergebnisse aus dieser Arbeit wurden mit Genehmigung der Fakultät für Lebenswissenschaften, vertreten durch den Mentor der Arbeit, in folgenden Beiträgen vorab veröffentlicht: Publikationen Joger, U., Bshaena, I. : A new Tarentola subspecies (Reptilia: Gekkonidae) endemic to Tunisia. Bonn Zoological Bulletin 57 (2): 267-247 (2010). Tagungsbeiträge Bshaena, I & Joger, U. : Phylogeny and systematics of North African Tarentola. (Vortrag). Deutsche Gesellschaft für Herpetologie und Terrarienkunde (DGHT) Nachzuchttagung Deutscher Herpetologentag, September (2009). A First of all I wish to express my sincere gratefulness to Allah, without his help this work would not been done. C I would like to express my gratitude to many individuals who assisted me K throughout the course of my doctoral research. First, I would like to express my appreciation to my supervisor, Prof. Dr. Ulrich Joger for his kind supervision, N sincere help, fruitful discussion, an open ear and for the extraordinary advice, caring, encouragement, and affection he bestowed on me. O My sincere thanks go to Prof. Dr. Miguel Vences for the opportunity of participating in this project, to work in his laboratory, and offered numerous W critical comments which contributed to improve the study. L I am also very grateful to Prof. Dr. Dirk Selmar, who kindly agreed to become a member of the dissertation committee. E I am much obliged to several people who contributed in sample material or D helped to make sample material available for me. I am grateful to all people in Libya who help me in field work for catching Lizards. G Many thanks go to all our laboratory technicians group who help me in the first period in the laboratory work. E In addition, I would like to thank all my colleagues for his kindly suggestions and M discussions. My special thanks go to my wife Asya Swiddan for her patient love and my pretty E kids Mustafa, Nada and Shahin for their lovely smiles that can relieve any kind of tiredness. N I am greatly thankful to my mother for her heartily feelings, unconditional support T and constant prayers. My huge thanks go to my brothers, especially Usama for his support and S managing our business in Libya during my PhD study Braunschweig, 16.03.2011 Ismail Mustafa Bshaena i Contents Chapter 1: General Introduction………………................................... 1 1.1. Introduction…………………………………………………………. 1 1.2. Family Gekkonidae………………………………………………… 6 1.3. Genus Tarentola……………………………………………………. 9 1.4. Area of study……………………………………………………….. 28 1.5. Previous studies……………………………………………………. 38 1.6. Aims of work……………………………………………………….... 41 Chapter 2: Morphological Study……………………………………...... 43 2.1. Introduction…………………………………………………………... 43 2.2. Material and methods………………………………………………. 46 2.3. Results……………………………………………………………….. 59 Chapter 3: Molecular Phylogenetic Study………………………………. 103 3.1. Introduction…………………………………………………………... 103 3.2. Material and methods……………………………………………….. 104 3.3. Results ……………………………………………………………….. 118 Chapter 4: General Discussion…………………………………………… 155 4.1. Morphological comparisons………………………………………… 155 4.2. Molecular phylogenetic affinities…………………………………… 160 4.3. Genetic distances……………………………………………………. 166 ______________________________________________________________________________ Phylogenetics and systematics of North-African Geckos Tarentola ii 4.4. Definition of Species ………………………………………………... 167 4.5. Conclusion of morphological and molecular study………………. 169 4.6. Taxonomic conclusions……………………………………………... 175 Summary:………………………………………………………………… 202 References:…………………………………………………………………. 206 Appendices:………………………………………………………………... 223 ______________________________________________________________________________ Phylogenetics and systematics of North-African Geckos Tarentola iii List of Figures 1- An adult T. annularis in Acacia tree holding its prey………………………….... 7 2- General distribution range of genus Tarentola…………………………………..10 3- Photograph of an adult T. mauritanica, with map showing its distribution range…………………………………………………………………………………14 4- Photograph of an adult T. m. fascicularis, with map showing its distribution range…………………………………………………………………………………15 5- Photograph of an adult T. m. juliae, with map showing its distribution range…………………………………………………………………………………16 6- Photograph of an adult T. m. pallida, with map showing its distribution range……………………………………………………………………………...….17 7- Photograph of an adult T. deserti, with map showing its distribution range………………………………………………………………………………….18 8- Photograph of an adult T. boehmei, with map showing its distribution range………………………………………………………………………………….19 9- Photograph of an adult T. mindiae, with map showing its distribution range………………………………………………………………………………….20 10- Photograph of an adult T. ephippiata, with map showing its distribution range………………………………………………………………………………….22 11- Photograph of an adult T. parvicarinata, with map showing its distribution range………………………………………………………………………………….24 12- Photograph of an adult T. a. annularis, with map showing its distribution range………………………………………………………………………………….24 13- Photograph of an adult T. neglecta, with map showing its distribution range………………………………………………………………………………….26 14- Map shows the topography of Libya……………………………………………..33 15- A satellite image of the African Sahara desert………………………………….36 ______________________________________________________________________________ Phylogenetics and systematics of North-African Geckos Tarentola iv 16- Distribution map of genus Tarentola in North-Africa…………………………...44 17- Map of Libya showing the main study sites……………………………………..47 18- Foot underside of gecko Tarentola………………………………………………53 19- Ventral view of Tarentola head……………………………………………………53 20- Map showing the initial groups used to define character variation……...…....60 21- CDF plots for Tarentola sp-Tunisia, Tarentola deserti-North Africa, Tarentola sp-complex-Libya, and Tarentola deserti-Libya…………………………………78 22- CDF plots for T. mauritanica from (Tunisia, Algeria, and Morocco), T. m. fascicularis-Libya, NW- African T. deserti, and T. deserti-Libya……………….79 23- CDF plots for T. sp-complex-Libya, T. mauritanica from (Tunisia, Algeria, and Morocco), and T. deserti-Libya……………………………………………………80 24- CDF plots for T. mauritanica (Tunisia, Algeria, and Morocco), T. deserti-Libya, T. sp-Ras Lanuf-Libya, and T. sp-sister group Ras Lanuf)………………….…81 25- CDF plots for T. sp-Sabha-South Libya, T. mauritanica (Tunisia, Algeria, and Morocco), T. deserti-Libya, , and T. sp-Suirt-sister group Sabha……………..85 26- CDF plots for T. mauritanica (Tunisia, Algeria, and Morocco), T. sp-Suirt- sister group Sabha, T. deserti-Libya,T. m. fascicularis-Egypt, T. m. fascicularis- Libya, T. mauritanica-Tripolitania, and T. sp-Sabha-South Libya……………..86 27- CDF plots for T. sp-complex1-Libya, T. sp-complex2-Libya, T. sp-Suirt-sister group Sabha, T. deserti-Libya, T. sp-Ras Lanuf-Libya, T. sp-sister group Ras Lanuf, T. m. fascicularis-Libya, T. neglecta-South Libya, and T. sp-Sabha- South Libya………………………………………………………………………….87 28- PCA for specimens T. deserti-Libya, T. sp-complex-Libya, T. sp-Tunisia, and T. deserti-North Africa………………………………………………………………92 29- PCA for specimens T. deserti-North Africa, T. deserti-Libya, T. m. fascicularis- Libya, T. mauritanica (Tunisia, Algeria, and Morocco)………………………...93 30- PCA for specimens T. deserti-Libya, T. sp-Ras Lanuf-Libya, T. sp-sister group Ras Lanuf-Libya, and T. mauritanica (Tunisia, Algeria, and Morocco). ..…….94 31- PCA for specimens T. deserti-Libya, T. sp-Suirt-Libya, T. sp-Sabha-South Libya, T. mauritanica (Tunisia, Algeria, and Morocco)…………….…………100 ______________________________________________________________________________ Phylogenetics and systematics of North-African Geckos Tarentola v 32- PCA for specimens T. deserti-Libya, T. m. fascicularis-Libya, T. sp-Suirt- Libya, T. sp-Sabha-South Libya, T. mauritanica-Tripolitania, T. mauritanica- Egypt, T. mauritanica (Tunisia, Algeria, and Morocco)………………………..101 33- PCA for Libyan specimens, T. deserti, T. m. fascicularis, T. sp-complex, T. sp-Ras Lanuf, T. sp-sister group Ras Lanuf, T. sp-Suirt, T. sp-Sabha, and T. neglecta- Gaber-Aown Oasis Lake………………………………………..…102 34- Neighbor-Joining phylogenetic tree, based on partial 12srRNA sequence 363 bp of 440 specimens, depicting the relationships among helotypes…………121 35- 50% majority-rule consensus obtained from Bayesian MCMC analysis. Based on partial 12srRNA sequence 382 bp………………………………………...…126 36- Tree derived from ML analysis, based on partial 12srRNA sequence 382 bp………………………………………………………………………………127 37- 50% majority-rule consensus obtained from Bayesian MCMC analysis. Based on partial 16srRNA-1 sequence 447 bp………………………………………...133 38- Tree derived from ML analysis, based on partial 16srRNA-1 sequence 447 bp…………………………………………..……………………………………....134 39- 50% majority-rule consensus obtained from Bayesian MCMC analysis. Based on partial 16srRNA-2 sequence 604 bp………………………………………...135 40- Tree derived from ML analysis, based on