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Genus Darevskia

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Genus Darevskia Phylogeny and morphological variation in the rock lizards of the genus Darevskia Mariam Gabelaia The doctoral thesis is submitted to the School of Natural Sciences and Medicine, Ilia State University and the Faculty of Sciences, Ghent University to obtain the degree of Ph.D in Life sciences and the degree of Doctor of Science: Biology School of Natural Sciences and Medicine Faculty of Sciences Supervisors: Prof. Dr. David Tarkhnishvili and Prof. Dr. Dominique Adriaens Ilia State University and Ghent University Tbilisi 2019 Statement As an author of this doctoral thesis, I confirm that this compilation thesis is my original work and does not include the materials of other authors that have already been published, accepted for publication or submitted for a degree, which have not been cited in an accepted manner. Mariam Gabelaia -------------------------------------------------------------- i აბსტრაქტი Darevskia გვარის კავკასიური კლდის ხვლიკები მცირე ზომის ხვლიკების მრავალფეროვანი ჯგუფია, რომლებიც ძირითადად დიდსა და მცირე კავკასიონზე ცხოვრობენ. Darevskia-ს სახეობების საზღვრების დადგენა რთულია, მათი გარეგნული მსგავსებისა და სახეობებს შორის გადაფარვადი მორფოლოგიური ნიშნების გამო. ამასთან ერთად, ისინი ხშირად ჰიბრიდიზირებენ და შედეგად ხშირია შუალედური მორფოლოგიისა და გენეტიკის მქონე ინდივიდები, რომელთა რომელიმე ცალკეული სახეობისთვის მიკუთვნება ძალიან რთულია. ეს განსაკუთრებით ვლინდება თვისობრივი მორფომეტრიული მეთოდებით შეფასებისას, რომელიც სხეულისა და ქერცლების სხვადასხვა განაზომებს ეფუძნება. ამ კვლევაში, ჩვენი მიზანია ვიპოვოთ თანხვედრა ფილოგენეტიკურ კავშირებსა და მორფოლოგიურ მსგავსებებს შორის Darevskia სახეობებში და გამოვიყენოთ გენეტიკური და მორფოლოგიური მეთოდები, ან მათი კომბინაცია სახეობების საზღვრების დასადგენად. კვლევისას ძირითადად ვიყენებთ მიტოქონდრიული დნმ- ის ხაზსა და გეომეტრიულ მორფომეტრიულ მეთოდებს (ფორმის ორ- განზომილებიანი ფურიეს კონტურული ანალიზი და სამ-განზომილებიან ლენდმარკებზე დაფუძნებული ანალიზი, ‘ტრადიციული’ მორფომეტრიის საპირისპიროდ) და ვცდილობთ გამოვავლინოთ ის მორფოლოგიური ნიშნები, რომლებიც ცალკეული სახეობებისთვისაა დამახასიათებელი და კარგად აღწერს სახეობათა დივერგენციას. კვლევის შედეგად დადგინდა, რომ Darevskia სახეობების თავისა და ზოგიერთი ქერცლების ფორმაზე გეომეტრიულ მორფომეტრიული ანალიზის გამოყენებით შესაძლებელია მათი ფილოგენეტიკური კავშირების მსგავსი სურათის მიღება, ან სახეობათა საზღვრების დადგენა, თუმცა უფრო ეფექტურია როდესაც გენეტიკურ (მიტოქონდრიული დნმ) ანალიზთან ვაერთიანებთ. ძირითადი საძიებო სიტყვები: Darevskia , ფილოგენია, გეომეტრიული მორფომეტრია. ii Abstract Caucasian rock lizards of the genus Darevskia are a diverse group of small lizards mainly living in the Greater and the Lesser Caucasus. It is difficult to delimit Darevskia species due to a high degree of morphological similarity and overlapping identification characters. Additionally, they also strongly hybridize and thus individuals with intermediate morphology and genetic characters are found. As a result, there are extreme difficulties in assigning found individuals to a distinct species. This is particularly true for the traditional morphological methods such as qualitative morphometrics, where multiple measurements of the body and scales are compared with each other. In the present project, we aimed to find the links between the phylogenetic relationships and morphological similarities in Darevskia, and apply genetic and morphological methods, or their combination, for delimiting the species boundaries. We mainly focus on the matrilineal phylogeny and geometric morphometric techniques (two-dimensional Fourier outline shape analysis and three-dimensional landmark-based analysis in comparison with ‘traditional’ morphometrics). More specifically we aim to reveal the morphological features which are species-specific and follow the species evolutionary divergence pattern. In conclusion to our study, the head shape and even some scalation features of Darevskia species can be suitable markers for depicting the species phylogenetic relationships or delimiting the species boundaries when assessed with geometric morphometric techniques, but perform the best if integrated with genetic (mtDNA) analysis. Keywoords: Darevskia, phylogeny, geometric morphometrics. Overzicht Kaukasische rotshagedissen van het geslacht Darevskia vormen een diverse groep van kleine hagedissen die hoofdzakelijk voorkomen in de Grote en de Kleine Kaukasus. Darevskia iii soorten zijn echter zeer moeilijk van elkaar te onderscheiden als gevolg van hun grote mate van morfologische gelijkenissen en de overlap in kenmerken die gebruikt worden om de soorten te identificeren. Bovendien vertonen ze intensieve hybridisatie, waardoor individuen met een intermediaire morfologie en genetische merkers worden teruggevonden. Dit maakt het extreem moeilijk om de gevonden specimens toe te wijzen aan gekende soorten. Dit is vooral zo indien gebruik wordt gemaakt van traditionele morfologische onderzoeksmethodes, zoals kwalitatieve morfometrie, waarbij meerdere lengtemetingen worden uitgevoerd op het lichaam en de schubben, om dan onderling te vergelijken. In dit project trachtten we de verbanden te achterhalen tussen de fylogenetische relaties en de morfologische gelijkenissen in Darevskia, waarbij op basis van genetische en morfologische methods (afzonderlijk en gecombineerd) gebruikt worden om soorten af te bakenen. We richten ons hier vooral op de matrilineaire fylogenie en geometrisch morfometrische technieken (twee-dimensionale Fourier analyse om de vorm op basis van contouren te analyseren, en drie-dimensionale vorm analyses op basis van landmarks, en dit vergeleken met meer ‘traditionele’ morphometrie). Meer specifiek is het doel om de morfologische eigenschappen te achterhalen die soort-specifiek zijn, en zodoende de evolutionaire divergentiepatronen van de soorten te achterhalen. Een conclusie van dit onderzoek is dat zowel de kopvorm, maar ook bepaalde eigenschappen in het schubpatroon bij de Darevskia species bruikbare indicatoren zijn om fylogenetische verwantschappen te achterhalen, of voor het identificeren en afbakenen van soorten. Betrouwbare resultaten worden echter enkel verkregen indien de data geïntegreerd werd met de genetische (mtDNA) analyses. Keywoorden: Darevskia, fylogenie, geometrisch morfometrie. iv Acknowledgements First of all, I would like to thank my supervisors David Tarkhnishvili and Dominique Adriaens for their professional and moral support throughout my doctorate studies. Their contributions to my dissertation are immense. I also want to thank Natia Barateli, Giorgi Iankoshvili, Levan Ninua, David Dekanoidze, and Aleksandre Gavashelishvili for their help in field work, Mari Murtskhvaladze, Tamar Beridze, Gocha Golubiani and Ana Avaliani for their help in laboratory work, Mikheil Elashvili and Giorgi Datunashvili for technical assistance, and Levan Mumladze, Wolfgang Böhme and Cort Anderson for their contributions to my doctorate research. I would also like to thank the researchers and coworkers at the Institute of Ecology, Ilia State University and Evolutionary morphology research group, Ghent University for their professional suggestions, support and friendly atmosphere. This doctorate research has been supported financially by Erasmus Mundus Action 2 EUROEAST Partnership, Shota Rustaveli National Science Foundation (Awards #GNSF/ST07/6-217, #217478 and #PhDF2016_79), internal budget of the Institute of Ecology (Ilia State University) and Alexander von Humboldt Foundation. Finally, I would like to thank Martin Piazena and Charlotte Cailliau, who made this joint collaboration possible, Ia Gurchiani, Elene Zhuravliova, Nana Rekhviashvili and UGent Joint Phd Team for their huge help in every administrational aspect and Ilia State University and Ghent University for providing the necessary infrastructure and shelter for my research. v Table of Contents Table of Contents .......................................................................................................................... vi List of Tables ................................................................................................................................. vii List of Figures .............................................................................................................................. viii List of Appendices ....................................................................................................................... xiv Abbreviations ............................................................................................................................... xv Chapter 1: Phylogenetic relationships and species’ boundaries in Darevskia ............................ 1 1.1 General Introduction ............................................................................................................ 1 1.2 Methodology ......................................................................................................................... 4 1.2.1 Sample collection ........................................................................................................... 4 1.2.2 DNA analysis .................................................................................................................. 8 1.2.3 ‘Traditional’ morphometric analysis ............................................................................. 9 1.2.4 Geometric morphometric analysis .............................................................................. 10 1.3 General Discussion ...........................................................................................................
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