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Apocynaceae, Apocynoideae) Systematics of the tribe Echiteae and the genus Prestonia (Apocynaceae, Apocynoideae) Dissertation zur Erlangung des Doktorgrades Dr. rer. nat Eingereicht an der Fakultät für Biologie, Chemie und Geowissenschaften J. Francisco Morales Bayreuth Die vorliegende Arbeit wurde von Oktober 2013 bis Januar 2016 in Bayreuth am Lehrstuhl Pflanzensystematik unter Betreuung von Prof. Dr. Sigrid Liede-Schumann und Dr. Mary Endress (Institute of Systematic and Evolutionary Botany, University of Zurich, Switzerland) angefertigt. Vollständiger Abdruck der von der Fakultät für Biologie, Chemie und Geowissenschaften der Universität Bayreuth genehmigten Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.). Dissertation eingereicht am: 31.01.2017 Zulassung durch die Promotionskommission: 15.02.2017 Wissenschaftliches Kolloquium: 27.04.2017 Amtierender Dekan: Prof. Dr. Stefan Schuster Prüfungsausschuss: Prof. Dr. Sigrid Liede-Schumann (Erstgutachterin) Prof. Dr. Carl Beierkuhnlein (Zweitgutachter) Prof. Dr. Bettina Engelbrecht (Vorsitz) PD. Dr. Ulrich Meve This dissertation is submitted as a “Cumulative Thesis” that includes four publications: one published, two accepted, and one in preparation for publication. List of Publications 1. Morales, J.F. & S. Liede-Schumann. 2016. The genus Prestonia (Apocynaceae) in Colombia. Phytotaxa 265: 204–224. 2. Morales, J.F., M. Endress & S. Liede-Schumann. Sex, drugs and pupusas: Disentangling relationships in Echiteae (Apocynaceae). Accepted, Taxon. 3. Morales, J.F., M. Endress & S. Liede-Schumann. A phylogenetic study of the genus Prestonia (Apocynaceae). Accepted, Annals of the Missouri Botanical Garden. 4. Morales, J.F. & M. Endress. A monograph of the genus Prestonia (Apocynaceae, Echiteae). To be submitted to Annals of the Missouri Botanical Garden or Phytotaxa. 4. Declaration of contribution to publications The thesis contains three research articles for which most parts were carried out by myself, under the supervision of Dr. Mary Endress, University of Zürich and Prof. Dr. Sigrid Liede-Schumann, University of Bayreuth. Field work for the DNA samples was conducted in Brazil, Colombia, Costa Rica, Panama and Peru from 2010 to 2015, supported by a private donor, the Instituto para la Investigación científica para la Amazonia de Colombia (SINCHI), the Graduate School of Bayreuth University and the DeutscheAkademische Austauschdienst (DAAD). Molecular work was done at the University of Bayreuth, under the supervision of Angelika Täuber and Margit Gebauer. Herbarium work was supported partially by the Missouri Botanical Garden, St. Louis, Missouri and the University of South Florida, Florida. Prof. Dr. Sigrid Liede-Schumann and J. Francisco Morales chose the DNA markers used in the phylogenetic studies. Papers were prepared under consideration of the comments and suggestions of all coauthors. 1st publication Morales, J.F. & S. Liede-Schumann. 2016. The genus Prestonia (Apocynaceae) in Colombia. Phytotaxa 265: 204–224. Authors’ contribution: Fieldwork in several Colombian departments (Antioquia, Cundinamarca, Guaviare, Guainia, and Tolima), the visit of different herbaria in Colombia and United States, and the morphological analysis of the herbarium samples were done by J. Francisco Morales, as well as the idea and structure of the manuscript, including distribution maps and line drawings. Photographs for this paper were taken in Colombia during 2013. J. Francisco Morales drafted the paper, Dr. Mary Endress reviewed a preliminary version and Prof. Dr. Sigrid Liede-Schumann added some suggestions and comments, which were incorporated in the final version. 2nd publication Morales, J.F., M. Endress & S. Liede-Schumann. Sex, drugs and pupusas: Disentangling relationships in Echiteae (Apocynaceae) (Accepted: Taxon) Authors’ contribution: Fieldwork for collecting the DNA samples in Latin America, as well as the visit to different herbaria with the same purpose was carried out by J. Francisco Morales from 2010 to 2015. J. Francisco Morales did the molecular work alone under the assistance of Angelika Täuber and Margit Gebauer at the Department of Plant Systematics, University of Bayreuth. Sequence alignments, matrices, and data analysis were carried out under the supervision of Prof. Dr. Sigrid Liede-Schumann. J. Francisco Morales wrote the paper, conducted the final data analysis, and drew the figures. Dr. Mary Endress corrected and reviewed the draft. Prof. Dr. Sigrid Liede-Schumann reviewed the final version of the manuscript before sending it for publication. 3rd publication Morales, J.F., M. Endress & S. Liede-Schumann. A phylogenetic study of the genus Prestonia (Apocynaceae) (Accepted: Annals of the Missouri Botanical Garden). Authors’ contribution: The DNA material was obtained by J. Francisco Morales from 2010 to 2015 through fieldwork in Latin America or the visit to different herbaria. Molecular laboratory work, sequence alignments, and matrices assembling were done by J. Francisco Morales. Dr. Sigrid Liede-Schumann reviewed the final versions of the assembled DNA sequence matrices. The manuscript was prepared by J. Francisco Morales with the support of Dr. Mary Endress, who also reviewed the paper and made several comments and suggestions. The final version was reviewed by Prof. Dr. Sigrid Liede-Schumann and all comments were incorporated in the final version of the manuscript. 4th publication Morales, J.F. & M. Endress. A monograph of the genus Prestonia (Apocynaceae, Echiteae, Prestoniinae) (In preparation, to be submitted March 2017) Authors’ contribution: The first author conducted the study of herbarium specimens and field work in several neotropical countries, drafted the paper, descriptions, illustrations, maps, and took most of the pictures included in the paper. Dr. Mary Endress improved the English style and the structure of the paper and made several comments on the original manuscript, included in the final version to be submitted. The paper is almost ready to be sent for publishing and only minor formatting issues are still lacking. CONTENT Summary 1 Zusammenfassung 3 1. General Introduction 5 2. Aims of Research 9 3. Synopsis 10 3.1 Materials and Methods 10 3.2 Results and Discussion 13 3.3 Conclusion and emerging research challenges 18 3.4 References 20 Publications 1. The genus Prestonia (Apocynaceae) in Colombia. 27 2. Sex, drugs and pupusas: Disentangling relationships in Echiteae 75 (Apocynaceae). 3. A phylogenetic study of the genus Prestonia (Apocynaceae). 132 4. A monograph of the genus Prestonia (Apocynaceae, Echiteae, 183 Prestoniinae) Acknowledgements 494 (Eidesstattliche) Versicherungen und Erklärungen 495 Summary Summary Of the 9 tribes in Apocynaceae-Apocynoideae, the Echiteae is a group of lianas (rarely erect herbs) and comprises 19 genera and about 200 species, of which 16 genera are restricted to the Neotropics. The tribe Echiteae was proposed more than 120 years ago, but its composition has varied through time. The results of some phylogenetic studies have suggested that the tribe is non-monophyletic, a condition also applying to of its genera in the present circumscription. In this work, we provide the first phylogenetic study of the Echiteae, using nuclear (ITS) and chloroplast markers, in order to test the monophyly of the tribe and its subtribes, analyzing the largest dataset ever compiled. The results show that Echiteae and four of its subtribes (Echitinae, Parsonsiinae, Peltastinae and Prestoniinae) as previously circumscribed are non-monophyletic. The fifth subtribe, Pentalinoninae, though monophyletic, does not belong to Echiteae; Prestonia and Temnadenia are resolved as polyphyletic, whereas Fernaldia is nested within Echites and Peltastes is nested in Macropharynx, and therefore, these two genera are reduced to synonymy. Fourteen genera are maintained and a new subtribe Laubertinae is proposed. In the ancestors of Echiteae, we hypothesize that an evolutionary shift took place in which steroidal alkaloids and/or cardenolides, characteristic for apocynoids, were replaced by parsonsine type pyrrolizidine alkaloids as the henceforth predominant chemical defense compounds. Prestonia is the second largest genus of Echiteae, with ca. 56 species distributed in Tropical America and the West Indies. The last monograph was published in 1936, but it is out of date due the number of species described since then. Only synopsis for some areas and flora treatments are available. Preliminary phylogenetic studies have suggested that the genus is not monophyletic. At the same time, four infrageneric classifications proposals have been published, but never have been tested with molecular data. Therefore, the circumscription of Prestonia and the relationships between the species remained unresolved. Two phylogenetic analyses using chloroplast (trnL intron and trnL-trnF intergenic spacer, rpl16 intron, rps16 intron, matK and 3′ / 5′ trnK intron) and nuclear data (ITS) were 1 Summary conducted for Prestonia with three aims: 1) to determine if the genus is monophyletic and to establish its relationships with the others genera in the tribe Echiteae, 2) to evaluate the current infrageneric classification proposals, as well as to determine the interspecific relationship in Prestonia, 3) to provide the necessary nomenclatural and / or taxonomic changes. The results of the present thesis show that Prestonia in the previous circumscription was polyphyletic, because P. riedelii is sister to Rhodocalyx and Temnadenia ornata is nested
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