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Rubiaceae, Ixoreae SYSTEMATICS OF THE PHILIPPINE ENDEMIC IXORA L. (RUBIACEAE, IXOREAE) Dissertation zur Erlangung des Doktorgrades Dr. rer. nat. an der Fakultät Biologie/Chemie/Geowissenschaften der Universität Bayreuth vorgelegt von Cecilia I. Banag Bayreuth, 2014 Die vorliegende Arbeit wurde in der Zeit von Juli 2012 bis September 2014 in Bayreuth am Lehrstuhl Pflanzensystematik unter Betreuung von Frau Prof. Dr. Sigrid Liede-Schumann und Herrn PD Dr. Ulrich Meve 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: 11.09.2014 Zulassung durch die Promotionskommission: 17.09.2014 Wissenschaftliches Kolloquium: 10.12.2014 Amtierender Dekan: Prof. Dr. Rhett Kempe Prüfungsausschuss: Prof. Dr. Sigrid Liede-Schumann (Erstgutachter) PD Dr. Gregor Aas (Zweitgutachter) Prof. Dr. Gerhard Gebauer (Vorsitz) Prof. Dr. Carl Beierkuhnlein This dissertation is submitted as a 'Cumulative Thesis' that includes four publications: three submitted articles and one article in preparation for submission. List of Publications Submitted (under review): 1) Banag C.I., Mouly A., Alejandro G.J.D., Meve U. & Liede-Schumann S.: Molecular phylogeny and biogeography of Philippine Ixora L. (Rubiaceae). Submitted to Taxon, TAXON-D-14-00139. 2) Banag C.I., Thrippleton T., Alejandro G.J.D., Reineking B. & Liede-Schumann S.: Bioclimatic niches of endemic Ixora species on the Philippines: potential threats by climate change. Submitted to Plant Ecology, VEGE-D-14-00279. 3) Banag C.I., Tandang D., Meve U. & Liede-Schumann S.: Two new species of Ixora (Ixoroideae, Rubiaceae) endemic to the Philippines. Submitted to Phytotaxa, 4646. In preparation for submission to Phytotaxa: 4) Banag C.I., De Block, P., Alejandro, G.J.D., Liede-Schumann S. & Meve U.: The Philippine species of Ixora (Rubiaceae). Publications not included into this thesis: 5) Banag C.I., Manalastas N.M.B. & Alejandro, G.J.D. 2014.: Synonymy of Philippine Ixora (Ixoreae, Rubiaceae) and Ixora silagoensis sp. nov. Nordic Journal of Botany doi: 10.1111/njb.00411. 6) Banag C.I. & Tandang D.: Taxonomic notes on three widespread endemic Ixora (Rubiaceae) in the Philippines. Submitted to National Museum of the Philippines Journal of Natural History (under review). Declaration of contribution to publications This thesis contains four research articles for which most of the work presented was carried out by myself. Molecular and most of the analytical work was done at the University of Bayreuth with the assistance and support of Prof. Dr. Sigrid Liede-Schumann, PD Dr. Ulrich Meve and Prof. Dr. Grecebio Jonathan Alejandro. Sample collections were conducted in the Philippines from 2010 to 2013 with the help of the Thomasian Angiosperm Phylogeny and Barcoding group from the University of Santo Tomas (UST), Manila, Philippines and Mr. Danilo Tandang and Dr. Arvin Diesmos from the National Museum of the Philippines (NMP). I prepared all manuscripts with considerations to the suggestions and comments of all co-authors. 1st Publication Banag C.I., Mouly A., Alejandro G.J.D., Meve U. & Liede-Schumann S.: Molecular phylogeny and biogeography of Philippine Ixora L. (Rubiaceae). Submitted to Taxon, TAXON-D-14-00139. Fieldwork was done by myself with the help of students and researchers from the University of Santo Tomas and National Museum of the Philippines. I did the molecular work alone with assistance of Mrs. Angelika Täuber at the Department of Plant Systematics, University of Bayreuth. Data analysis was done by myself with guidance from Prof. Dr. Sigrid Liede Schumann and Prof. Dr. Arnaud Mouly. I have written the manuscript with considerations to the comments of all co-authors. 2nd Publication Banag C.I., Thrippleton T., Alejandro G.J.D., Reineking B. & Liede-Schumann S.: Bioclimatic niches of endemic Ixora species on the Philippines: potential threats by climate change. Submitted to Plant Ecology, VEGE-D-14-00279. Georeferencing of all localities found in the herbarium specimens of Ixora was done by myself. Mr. Timothy Thrippleton (ETH, Zurich) helped me run the modelling in MAXENT using the R script prepared by Prof. Dr. Björn Reineking. Data analysis was done by myself with the assistance of Mr. Thrippleton and Prof. Dr. Sigrid Liede-Schumann who had the idea for this research. I have written the manuscript with considerations to the comments from all co-authors. 3rd Publication Banag C.I., Tandang D., Meve U. & Liede-Schumann S.: Two new species of Ixora (Ixoroideae, Rubiaceae) endemic to the Philippines. Submitted to Phytotaxa, 4646. Fieldwork in Samar was done by myself for the collection of I. reynaldoi and the fieldwork in Palawan was done by D. Tandang (NMP, Philippines) and Prof. Grecebio Jonathan D. Alejandro (UST, Philippines) for the collection of I. alejandroi. Morphological analysis of the two new species was done by myself. Botanical illustrations were drawn by Arianne Zacarias (UST) and Nemesio B. Diego, Jr. (NMP). I prepared the manuscript with considerations to the comments from all co-authors. 4th Publication Banag C.I., Alejandro, De Block, P., G.J.D., Liede-Schumann S. & Meve U.: The Philippine species of Ixora (Rubiaceae). In preparation for submission to Phytotaxa. Fieldwork in the Philippines was done by myself with the help of students and researchers from the University of Santo Tomas and National Museum of the Philippines. All herbarium specimens of Ixora from A, C, L, NY, and US was sent on loan to UBT (curator: PD Dr. Ulrich Meve). Research visits to BK, BR, C, CAHUP, K, P, PNH, PPC, SAN and morphological analysis of all herbarium specimens was done by myself. Botanical illustrations were drawn by Arianne Zacarias (UST), Joseph Capino (UST) and Nemesio B. Diego, Jr. (NMP). I prepared the manuscript with considerations to the comments from all co-authors. CONTENTS Summary 1 Zusammenfassung 3 1. General Introduction 6 2. Aims of Research 10 3. Synopsis 3.1 Materials and Methods 11 3.2 Results and Discussion 17 3.3 Conclusion and Perspectives 21 3.4 References 23 Publications 1. "Molecular phylogeny and biogeography of Philippine Ixora L. (Rubiaceae)" 29 2. "Bioclimatic niches of endemic Ixora species on the Philippines: potential threats by climate change" 56 3. "Two new species of Ixora (Ixoroideae, Rubiaceae) endemic to the Philippines" 84 4. "The Philippine species of Ixora (Rubiaceae)" 95 Acknowledgements 229 Eidesstattliche Erklärung 232 Summary Summary Ixora L. belonging to the tribe Ixoreae (Rubiaceae, subfamily Ixoroideae), is a large pantropical rain forest genus of shrubs and small trees. It comprises ca. 530 species that is presently subject to intensive investigations leading to far-reaching systematic changes. Recently, based on molecular evidence, Ixora was further enlarge to accommodate several genera (Captaincookia N.Hallé, Doricera Verdc., Hitoa Nadeaud, Myonima Comm. ex Juss., Thouarsiora Homolle ex Arénes and Versteegia Valeton). In consequence, tribe Ixoreae has become monogeneric. Although there have been a number of revisions focusing on specific geographical regions, e.g. in Africa, Madagascar, Marquesas Islands and Australia, the continental Asian taxa have never been fully revised and no clear picture of their diversity is available. In the Philippines, the only available account of Ixora is an enumeration by Merrill published in 1923 which is more than 85 years old and outdated due to numerous classification changes in the family Rubiaceae. To date, species diversity of Ixora in the Philippines remains poorly known and basic information on species distribution and ecology, taxonomic keys and diagnostic illustrations are limited. With this work, I want to fill a gap in our knowledge of the genus by assessing the phylogenetic relationships of Philippine representatives of Ixora: 1) using morphological and molecular data, 2) derive information on environmental requirements of Ixora species endemic to the Philippines using Maxent, and 3) publish a comprehensive taxonomic revision of the Philippine Ixora including nomenclatural treatments (lectotypifications) and new records, full illustrations, distribution maps and keys of the recognized taxa. An important first result of the present research was the finding that Philippine Ixora is polyphyletic, with representatives from at least three independent lineages. In both the parsimony and Bayesian analyses of the combined chloroplast markers (rps16 and trnT–F) and nuclear markers (ITS and ETS), two main lineages of Ixora were recovered, an Asian- Pacific lineage and an Asian-African-Malagasy-Mascarene-Neotropical lineage. Also, the results of the Statistical Dispersal-Vicariance Analysis (S-DIVA) revealed a complex biogeographical history for Ixora with several dispersal and vicariance events that have led to its current distribution. Furthermore, Philippines are shown to constitute an overlay zone into which species from both major lineages of Ixora, the Pacific and the African-Asian one, have immigrated and subsequently radiated. 1 Summary The results presented also revealed for the first time the bioclimatic relationships of endemic Philippine Ixora and showed that the species preferred areas with lower amount of annual rainfall and regions with higher annual precipitation variance (standard deviation). This may indicate the species’ potential to adapt to the changing climate or an adaptation of the species to monsoon
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