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Phylogeography of the Scorpion Genus Buthus in the Maghreb Region

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Phylogeography of the Scorpion Genus Buthus in the Maghreb Region Faculdade de Ciências da Universidade do Porto Tese de Mestrado em Phylogeography of the scorpion genus Buthus in the Maghreb region por Diana Pedroso Mestrado em Biodiversidade, Genética e Evolução Junho de 2012 Todas as correções determinadas pelo júri, e só essas, foram efetuadas. O Presidente do Júri, Porto, ______/______/_________ Assinatura: _______________________________________________________________________ Diana de Oliveira Gomes Pedroso Phylogeography of the scorpion genus Buthus in the Maghreb region Thesis submitted to the Faculty of sciences of the University of Porto in the fulfilment of the requirements for the degree of Master of Science Supervisor: Dr.Arie van der Meijden CIBIO, Research center in Biodiversity and Genetic Resources of the University of Porto Co-supervisor: Dr. D. James Harris CIBIO, Research center in Biodiversity and Genetic Resources of the University of Porto AN ESSAY ON SCORPION by Victor Fet 1. One-twelfth of you were born under this sign. A vile tail-biting beast, ’tis also mine Subject of study. In the times of yore When alchemists were to bypass the ore To get the gold—the scorpion, alas, Was just too bad to be ignored, and thus Hundreds of rare specimens, no doubt, Were ground down in bulk by some imposing lout, Or burnt alive with lashing metasomas Amidst sulphuric mist and vile aromas. Be as it may, Linnaeus comes to mind— Who, in his System, every type and kind, Variety and species, form and race Positioned well in God-appointed place. Six kinds of scorpion—the whole distinguished lot— Were given names in their generic slot, Named Scorpio by Carolus the Great, In year seventeen and fifty-eight. Type A—for buthids. Mighty is their race, Found in all continents, in nearly every place, In deserts or in tropics, high and low, They everywhere in black light brightly glow. With an immensely potent venom, those Beasts, as a rule, have thin and weaker claws. … Victor Fet Euscorpius — 2003, No. 11 Acknowledgements I would like to thank to the many people that helped me all way, during in my master’s thesis and also in the writing of this thesis; To Dr.Arie van der Meijden for receiving me as his master student in this project, for helping me all the way, motivating me when things were not going as smoothly as they I planned, for teaching me and still giving me wings to fly; To Pedro Coelho for all the help in laboratory, for all the suggestions in the thesis and for all the fun moments at the ecology lab; To Pedro Sousa for his help suggestions and patience in the writing of this thesis and through all the last year; To Dr.James Harris for receiving me in the Integrative Biogeography, Ecology and Evolution group and helping me with his suggestions; To all my colleagues in the ecology lab: Iolanda, Beatriz, Luís, João Maia and Fátima Jorge for all the fun moments during my master’s. Last but not least, to my mother and to Bruno, for all the love, comprehension and loads of patience with me. Table of contents Abstract ............................................................................................................................ 3 Resumo ............................................................................................................................. 5 List of Figure .................................................................................................................... 7 List of Tables .................................................................................................................... 11 Objectives ......................................................................................................................... 13 Chapter I Introduction ........................................................................................................ 15 Paleontology and terrestrialization of scorpions ................................. 15 Fluorescence : Strategy or side effect? ............................................... 21 Venom and toxicity ........................................................................... 22 Genetic markers ................................................................................ 24 The genus Buthus ............................................................................................... 27 Previous studies .................................................................................................. 31 Chapter II Methodology ...................................................................................................... 35 Sampling ............................................................................................ 35 DNA extraction, amplification and sequencing ................................... 39 Nuclear markers ................................................................................. 40 Mitochondrial markers ....................................................................... 43 Phylogenetic analysis ......................................................................... 44 Chapter III Results................................................................................................................ 47 Chapter IV Discussion .......................................................................................................... 63 Conclusion ......................................................................................................... 71 Appendix ............................................................................................................ 73 1 2 Abstract The genus Buthus is one of the most widespread scorpion genus in the Maghreb region, and even though several phylogenetic studies were recently performed upon it, the phylogenetic relationships within the genus are still complex and remain partially unsolved. In total, 59 specimens from several Buthus species were used in this work and their phylogenetic relationships investigated with different phylogenetic inference approaches as the Maximum Likelihood (ML) and Bayesian Inference (BI) methods, based on mitochondrial sequence data of three mitochondrial markers. This study was the first to include multiple mitochondrial markers in order to obtain further resolution within the genus The result of the phylogenetic analysis uncovered 5 well supported clades in the Maghreb region: three exclusive to Morocco, one with its distribution core in Morocco but also shared with Algeria and one last clade, unreported until now, from Tunisia. The final trees resulting from the ML and BI analysis do not present basal resolution and still the relationships between the clades lacked support. However, the support for each of the formed clade is high, holding significant levels of genetic diversity within the clade and between clades. The clades with its distribution centre in Morocco present higher genetic distances within each clade, especially the most widespread clade in Morocco (clade 3) in opposition to the single clade present in Tunisia, with the lowest pairwise genetic distance values. There seems to be a positive correlation between genetic distances between clades and geographical distances, which also may point towards a recent expansion of Buthus towards eastern region of the Maghreb, more precisely into Tunisia. This study corroborated the results from previous studies, like Sousa et al. (2012,) and found even more diversity within Buthus, for instance in Tunisia, which had never been studied in such detail before. The genetic distances among the clades also corroborate the theory that the centre of diversity of Buthus is in Morocco, more specifically in the Atlas- mountains that can act a reservoir of diversity for Buthus due to its role as a refugia during the glaciations. The mountains can also act as a geographical barrier to gene flow, impeding it, and at the same time, favouring processes of speciation and endemism. 3 4 Resumo O género Buthus é um dos géneros de escorpiões mais difundidos na região do Magreb e apesar de ter sido alvo de recentes estudos filogenéticos, as relações filogenéticas dentro do géneros ainda permanecem complexas e parcialmente não resolvidas. No total, 59 espécimes de diferentes espécies de Buthus foram utilizados neste estudo e as suas relações filogenéticas investigadas através de diferentes abordagens filogenéticas como o método de “Maximum Likelihood” (ML) e “Bayesian Inference” (BI) baseando em sequências mitocondriais de três marcadores mitocondriais distintos. Este estudo foi o primeiro a incluir marcadores mitocondriais múltiplos com vista na obtenção uma melhor resolução dentro do género. Os resultados da análise filogenética revelaram 5 clades bem suportados na região do Magreb: três clades exclusivos de Marrocos, um exclusivo da Argélia, um outro que possui o seu centro de distribuição em Marrocos mas também está presente na Argélia e um último clade desconhecido até agora, presente na Tunísia. As árvores finais resultantes da análise de ML e BI não apresentam resolução basal e as relações entre os clades têm carência de suporte. Contudo, o suporte para cada um dos clades formados é elevado, com altos níveis significativos de diversidade genética dentro dos clades e entre clades. Os clades com o seu centro de distribuição em Marrocos apresentam maiores distâncias genéticas dentro de cada clade, especialmente no clade mais difundido em Marrocos (clade 3) em oposição com o único clade presente na Tunísia, que apresenta os valores mais baixos de distância genética. Parece existir uma relação positiva entre as distâncias genéticas e a distância
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