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Evolution and Biogeography of Buxus L. (Buxaceae) in Cuba and the Caribbean

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Evolution and Biogeography of Buxus L. (Buxaceae) in Cuba and the Caribbean Evolution and biogeography of Buxus L. (Buxaceae) in Cuba and the Caribbean Dissertation zur Erlangung des akademischen Grades des Doktors der Naturwissenschaften (Dr. rer. nat.) eingereicht im Fachbereich Biologie, Chemie, Pharmazie der Freien Universität Berlin vorgelegt von Pedro Alejandro González Gutiérrez aus Gibara, Kuba Berlin, Juni 2014 This work was carried out between 2009 and 2014 under the supervision of Prof. Dr. Thomas Borsch in Institut für Biologie, Systematische Botanik und Pflanzengeographie der Freien Universität Berlin. 1st Reviewer: Prof. Dr. Thomas Borsch 2nd Reviewer: Prof. Dr. Kurt Zoglauer Date of defense: 17. Juli 2014 II Acknowledgements Herewith I would like to thank Prof. Dr. Thomas Borsch and the institution led by him, the Botanical Garden and Botanical Museum Berlin-Dahlem (BGBM), for the support during the last five years. Prof. Borsch accepted me as doctoral student in 2009 and from that date to the present I have always had his support and advice. Without his constant support and that of the BGBM this thesis would have remained a dream. Special thanks to the members of my dissertation committee for generously offering their time, for their attention and patience. For their encouragement and various kinds of help I thank Dr. Rosa Rankin Rodríguez (National Botanical Garden of Cuba), Prof. Dr. Egon Köhler (Humboldt University, HU) and Prof. Dr. Kurt Zoglauer (HU). Dr. Rankin provided valuable information about the cultivated species of Buxus in the National Botanical Garden of Cuba and about herbarium specimens. Dr. Köhler has shared with me his rich knowledge about the Cuban Buxus and Dr. Zoglauer suggested me to make my doctoral thesis on Buxus during my first visit to Berlin in 2005. I am deeply indebted to Dr. Susy Fuentes Bazan (BGBM), who kindly and patiently explained to me every detail regarding the work in the laboratory and who got me started with data analysis, for sharing with me her knowledge. Also I am grateful for her encouragement and her support in the organization of my last visit to Berlin. Further I thank Dr. Andrea Rupps (HU) for her help in organizing my previous visits to Berlin. My warmest thanks are extended to Dr. Angela T. Leyva Sánchez and the institution led by her, the National Botanical Garden of Cuba, for giving me the possibility of working with the living collection of Cuban Buxus and for her valuable support in many ways. I appreciate the advice of Prof. Dr. Werner Greuter on nomenclatural issues of the chapter 3 of this thesis. My thanks also go to the German Service of Academic Exchange (DAAD) for supporting me with scholarships (December/2009-May/2010, March/2011-June/2011). I III am grateful also to the Society of Friends of the Botanischer Garten und Botanisches Museum Berlin-Dahlem for supporting the fieldwork in Cuba and my stays in Berlin (2009-2010, 2010-2011, 2012 and 2013-2014). My thanks extend to the Cuban and Panamanian environmental authorities for providing permissions to collect samples included in this study. I am very grateful to the following people and institutions for collaborating with this work with samples, pictures and valuable information: Lic. Carmen Galdames (Smithsonian Tropical Research Institute, Panama), Dr. James C. Salomon (MO, USA), Dr. Lilian Ferrufino (TEFH, Honduras), Dr. Alberto Areces (Puerto Rico), Dr. Miguel Cházaro (University of Guadalajara, México), M. Sc. Sergio Avendaño (Institute of Xalapa, México), M. Sc. Yocupitzia Ramírez (Institute of Ecology, Michoacán, México), Dr. John Clark (University of Alabama, USA). To the curators and staff of the herbaria B, HAJB, IEB, MO, NY, TEFH, US, XAL. For their collaboration in the field, thanks go to Dr. Jorge Gutiérrez (National Botanical Garden of Cuba), Dr. Nils Köster (BGBM), M. Sc. Wilder Carmenate (Botanical Garden of Holguín, Cuba), M. Sc. Orestes Méndez (University of Las Villas, Cuba), M. Sc. Idelfonso Castañeda (Botanical Garden of Santa Clara, Cuba). For their advice and valuable support: Dr. Federico Luebert (University of Bonn, Germany), Dr. Marc Appelhans (University of Göttingen, Germany), Dr. Angélica Cervantes (México), Dr. Natalia Filipowicz (Gdansk, Poland), Dr. Rocío Santos (University of Sevilla, Spain). For his support and supervision in the laboratory I appreciate the collaboration of Ing. Kim Govers. Also for their support and collaboration in the laboratory I thank Bettina Giesicke (Institute for Biology of the Free University of Berlin, IB-FU), Dipl. Biol. Doreen Weigel (FU) and Dr. Markus Ackermann. Ramona Henkel (HU) and her supervisor Prof. Dr. Gert Dudel (Technic University of Dresden) helped with the determination of nickel concentrations. Lic. David Lambert identified the species of insects seen in the flowers of Cuban Buxus. I thank Dr. Nadja Korotkova (BGBM), Dr. Sasha Ismail (BGBM), Dr. Lars Nauheimer (IB-FU), Dr. Ludo Muller (IB-FU) and Dipl. Biol. Virginia Duwe for their useful commentaries on early versions of chapters and summaries of this thesis. Thanks go also to Mrs Margitta Hielscher (HU), Michael Rodewald (BGBM) and Tamara Freyer-Dohlus for their assistance in the microscopy and the preparation of the IV illustrations, respectively. I thank Dipl. Geogr. Bastian Stößel for his help with editing of maps. I also appreciate the assistance and support of my colleagues in Berlin: Dipl. Biol. Vanessa Di Vincenzo and Christina Grüber; and of my colleagues of Centro de Investigaciones y Servicios Ambientales y Tecnológicos de Holguín (CISAT): Elier, Tony, Carmen, Sara Ilse, Sara, Alejandro, Carlos, Sergio, José Alberto, Elena, Frank, David, Yanier, Beatriz, Jacobo and Osvaldo, for their constant support. Special thanks go to the departments of International collaboration of CISAT, of the Delegation of Ministry of Sciences, Technology and Environment in Holguín and to Lucia Pérez in the office of protocol of CITMA in Havana for their valuable help and support. I am endlessly grateful to my family, in a wide sense, for their support in all ways. I specially thank my mother Miriam and my sister María, for their constant effort and love which helped me to achieve this goal. My deep gratitude goes to my wife Zaharaí and my children Carlitos and Sandra for being my inspiration and the most important reason to continue. To my family in Berlin, Gila and Rolf Hamacher for adopting me as a true member of their family and for their valuable help and sincere friendship in all circumstances. I thank to Marco and Tanja Hamacher, and Prof. Dr. Max Steller for their support. Thanks go also to my sister- in- law Lynn Woolfrey for helping me with corrections to my English and to my brother Omar for his support. I am indebted to my friends Octavio and Martina, Mandy and Yudith, Claribel and Rene, Elke and Dörthe for nice moments in Berlin and for their valuable support. Thanks go to Lucy and Maikel for their support in Havana, to Lino Céspedes and Nelida Abarca for their valuable help in decisive moments. I am grateful to Cuba and its nature for being an endless source of inspiration in my work. Final thanks go to my professor Dr. Jorge Sierra Calzado for showing me the way to Botany through his wonderful lectures on Systematic Botany in 1994-1995 at the University of Oriente in Santiago de Cuba, and to my father Pedro Emilio for encouraging me to continue my studies to obtain my doctorate. To my professor and friend Jorge Sierra and to my father Pedro to whom I dedicate this work. V Summary González Gutiérrez, Pedro Alejandro. 2014. Evolution and biogeography of Buxus L. (Buxaceae) in Cuba and the Caribbean. Doctoral thesis, Fachbereich Biologie, Chemie, Pharmazie. Institut für Biologie/Botanik, Freie Universität Berlin, Germany. Buxus L. is the largest genus of the family Buxaceae, with c. 100 species distributed in all continents except Australia and Antarctica. The centres of morphological and ecological diversity of Buxus are the Caribbean, East Asia and Africa including Madagascar. Cuba is the territory with the highest number of Buxus taxa in the world. In Cuba a total of 37 species and seven subspecies of this genus occur, 95% of which are endemic. Most Cuban Buxus are endemic to the serpentine outcrops in east, central and west Cuba while a smaller number of species inhabits limestone areas of the island. About 50% of the Cuban Buxus accumulate or hyperaccumulate nickel (Ni), being the only members of the family Buxaceae with such ability. The adaptation to growth on serpentines and their capacity to accumulate or hyperaccumulate Ni appeared among the factors triggering the evolution of Buxus and other groups in the Cuban flora. For the reasons exposed above the Cuban and Caribbean Buxus are an appealing group for biogeographical and evolutionary studies. This study uses them as a model to address the following questions: (1) Are the Cuban and the other Caribbean Buxus a monophyletic group? (2) When did Buxus arrive in the Caribbean and when occurred the diversification of the genus on the islands? (3) What are now the closest relatives and were then the ancestors of Cuban and Caribbean Buxus and where did they occur? (4) Are there specific migration routes of Buxus in the Caribbean? and (5) Which factors have triggered the speciation of Buxus in Cuba? Following a detailed introduction that summarizes our current state of knowledge and reviews the existing literature, Chapter 2 of this thesis focuses on the phylogeny and biogeography of Buxus including a total of 53 species of Buxus, where 34 species and 5 subspecies are from Cuba (c. 90% of taxa known), several also from different localities, and also species from the Bahamas, Hispaniola, Jamaica, Mexico, Panama and Puerto Rico. Buxus representatives from Eurasia and Africa were also included. The outgroup includes all other genera of Buxales and representatives of VI Trochodendrales, Proteales, Sabiales and Ranunculales.
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