SYSTEMATICS AND BIOGEOGRAPHY OF SOUTHEAST ASIAN SENECIONEAE (ASTERACEAE): REVISION OF CISSAMPELOPSIS AND GYNURA AND HYBRIDISATION IN THE INTRODUCED CRASSOCEPHALUM Dissertation Zur Erlangung des Grades Doktor der Naturwissenschaften Am Fachbereich Biologie Der Johannes Gutenberg-Universität Mainz Ongkarn Vanijajiva geb. am 24 März 1977 in Krabi, Thailand Mainz, 2008 Tag der mündlichen Prüfung: 23 September 2008 Gynura siamensis Vanijajiva & Kadereit CONTENTS SUMMARY...............................................................................................................................1 ZUSAMMENFASSUNG.........................................................................................................2 1. GENERAL INTRODUCTION..................................................................................3 1.1 The Flora of Southeast Asia………………………………………………………...…...3 1.2 Tribe Senecioneae Cass……………………………………………………………….....4 1.3 Genus Cissampelopsis (DC.) Miq...................................................................................6 1.4 Genus Gynura Cass...........................................................................................................6 1.5 Genus Crassocephalum Moench.......................................................................................7 1.6 Statement of research…………………………………………………………..………...7 2. A REVISION OF CISSAMPELOPSIS (ASTERACEAE: SENECONEAE).................9 2.1 Introduction.......................................................................................................................9 2.1.1 Taxonomic history………………………………………..…………….........………9 2.1.2 The relationships of Cissampelopsis……………………………………….………10 2.2 Material and Methods…………......................................................................................10 2.3 Results and Discussion....................................................................................................12 3. A REVISION OF GYNURA (ASTERACEAE: SENECONEAE) ...............................32 3.1 Introduction.................................................................................................................... 32 3.1.1 Taxonomic history…………………………………………………………………32 3.1.2 The relationships of Gynura…………………………………………..……………34 3.3 Material and methods……………………………………..............................................34 3.4 Results and Discussion……………................................................................................34 4. MORPHOLOGICAL AND MORLECULAR EVIDENCE FOR INTERSPECIFIC HYBRIDISATION IN THE INTRODUCED AFRICAN GENUS CRASSOCEPHALUM (ASTERACEAE: SENECONEAE) IN ASIA.........................107 4.1 Introduction………………...........................................................................................107 4.2 Material and methods………………………………....................................................108 4.2.1 Plant material………………………………………………………………….….108 4.2.2 Pollen and seed fertility…………………………………………………….…….109 4.2.3 Chromosome counts…………………………..………………………………..…110 4.2.4 DNA isolation, PCR amplification and sequencing……………………………...110 4.4.5 Sequence alignment…………………………………………………...…….……111 4.3 Results…………….......................................................................................................112 4.3.1 Geographical distribution and ecology, morphology, taxonomy……………....…112 4.3.2 Taxonomy of Crassocephalum in Asia………………………………………...…112 4.3.3 Pollen and seed fertility………………………………….…………………….....119 4.4.4 Chromosome numbers……………………………………….…………………...119 4.4.4 Nuclear ribosomal DNA ITS sequences……………………………...……….….120 4.4.6 Chloroplast DNA trnL-3′-trnF sequences……………………………..………….121 4.4 Discussion……………..………………………………………………………....…...122 4.4.1 Taxonomy and chromosome numbers of Crassocephalum in Asia……………122 4.4.2 Introduction history of Crassocephalum in Asia……………………….……122 4.4.3 The intermediate specimens………………………………………………….…...123 5. REFERENCES……………………………………………………...…………………124 6. ACKNOWLEDGEMENTS ……………………..………...….………..………….….130 7. CURRICULUM VITAE………………………………………………..…………..…131 1 SUMMARY This dissertation includes two parts. The first contains an introduction to the flora of Southeast Asia and the objects of this study, Asteraceae-Senecioneae Cass. with Cissampelopsis (DC.) Miq., Gynura Cass. and Crassocephalum Moench (Chapter 1). The second part comprises three manuscripts based on original research (Chapter 2- 4). In chapter 2 a revision of the Asian genus Cissampelopsis is provided. Two sections with ten species and two varieties are recognized. These are sect. Buimalia C. Jeffrey & Y. L. Chen with C. buimalia (Buch.-Ham. ex D. Don) C. Jeffrey & Y. L. Chen, C. erythrochaeta C. Jeffrey & Y. L. Chen and C. calcadensis (Ramasw.) C. Jeffrey & Y. L. Chen and sect. Cissampelopsis with C. glandulosa C. Jeffrey & Y. L. Chen, C. walkeri (Arn.) C. Jeffrey & Y. L. Chen with var. walkeri and var. floccosa Vanijajiva & Kadereit (var. nov.), C. corifolia C. Jeffrey & Y. L. Chen, C. volubilis (Bl.) Miq, C. ansteadii (Tadul. & Jacob) C. Jeffrey & Y. L. Chen, C. spelaeicola (Van.) C. Jeffrey & Y. L. Chen and C. corymbosa (Wall. ex DC.) C. Jeffrey & Y. L. Chen. Keys, descriptions, photographs of floral characters and distribution maps are provided. Chapter 3 is a revision of the palaeotropical genus Gynura. Forty-four species are recognized of which three are described as new. These are G. davisii Vanijajiva & Kadereit, G. siamensis Vanijajiva & Kadereit and G. villosus Vanijajiva & Kadereit. Gynura dissecta (F. G. Davies) Vanijajiva & Kadereit, G. annua (F. G. Davies) Vanijajiva & Kadereit and G. aurantiaca (Bl) DC. subsp. parviflora (F. G. Davies) Vanijajiva & Kadereit are new combinations. An identification key, descriptions and distribution maps are provided. In chapter 4, an analysis of the genus Crassocephalum in Asia, introduced there from Africa, is presented. This analysis is based on extensive field work, herbarium studies, analyses of pollen and seed fertility and chromosome numbers as well as ITS and trnL-F sequences. We found that Crassocephalum in Asia comprises two species and their interspecific hybrid. The two species are C. crepidioides (Benth.) S. Moore and C. rubens (Juss. ex Jacq.) S. Moore, of which the latter is a new record for Asia. The hybrid between these two species originated from a cross C. crepidioides (2n=40) as female and C. rubens (2n=40) as male parent. 2 ZUSAMMENFASSUNG Die vorgelegte Dissertation enthält zwei Teile. Der erste beinhaltet eine Einführung in die Flora Südostasiens und die Untersuchungsgruppe Asteraceae- Senecioneae Cass. mit den Gattungen Cissampelopsis (DC.) Miq., Gynura Cass. und Crassocephalum Moench (Kapitel 1). Der zweite Teil besteht aus drei Manuskripten, die auf originalen Forschungsergebnissen basieren (Kapitel 2-4). In Kapitel 2 wird eine Revision der asiatischen Gattung Cissampelopsis vorgelegt. Die folgenden zwei Sektionen mit zehn Arten und zwei Varietäten werden anerkannt: sect. Buimalia C. Jeffrey & Y. L. Chen mit C. buimalia (Buch.-Ham. ex D. Don) C. Jeffrey & Y. L. Chen, C. erythrochaeta C. Jeffrey & Y. L. Chen und C. calcadensis (Ramasw.) C. Jeffrey & Y. L. Chen sowie sect. Cissampelopsis mit C. glandulosa C. Jeffrey & Y. L. Chen, C. walkeri (Arn.) C. Jeffrey & Y. L. Chen mit var. walkeri und var. floccosa Vanijajiva & Kadereit (var. nov.), C. corifolia C. Jeffrey & Y. L. Chen, C. volubilis (Bl.) Miq, C. ansteadii (Tadul. & Jacob) C. Jeffrey & Y. L. Chen, C. spelaeicola (Van.) C. Jeffrey & Y. L. Chen und C. corymbosa (Wall. ex DC.) C. Jeffrey & Y. L. Chen. Schlüssel, Artbeschreibungen, Fotographien von Blütenmerkmalen und Verbreitungskarten werden präsentiert. Kapitel 3 beinhaltet die Revision der paläotropischen Gattung Gynura. Vierundvierzig Arten werden anerkannt, darunter die folgenden drei Neubeschreibungen: G. davisii Vanijajiva & Kadereit, G. siamensis Vanijajiva & Kadereit und G. villosus Vanijajiva & Kadereit. Gynura dissecta (F. G. Davies) Vanijajiva & Kadereit, G. annua (F. G. Davies) Vanijajiva & Kadereit und G. aurantiaca (Bl) DC. subsp. parviflora (F. G. Davies) Vanijajiva & Kadereit sind Neukombinationen. Ein Schlüssel, Artbeschreibungen und Verbreitungskarten werden vorgelegt. In Kapitel 4 wird eine Analyse von Crassocephalum in Asien, einer aus Afrika eingeschleppten Gattung, präsentiert. Diese Untersuchung basiert auf umfangreicher Feldarbeit, Herbarstudien, Analysen der Pollen- und Samenfertilität, Chromosomenzählungen sowie ITS- und trnL-F-Sequenzen. Die Studie ergab, dass Crassocephalum in Asien mit zwei Arten und deren Hybrid vertreten ist. Die zwei Arten sind C. crepidioides (Benth.) S. Moore und C. rubens (Juss. ex Jacq.) S. Moore, wobei letztere einen Neufund für Asien darstellt. Der Hybrid aus diesen beiden Arten resultiert aus einer Kreuzung von C. crepidioides (2n=40) als weiblichem und C. rubens (2n=40) als männlichem Elter. 1. INTRODUCTION 3 1. GENERAL INTRODUCTION 1.1 The Flora of Southeast Asia Southeast Asia (Fig. 1) consists of two geographic regions: the Asian mainland and island arcs and archipelagoes to the east and southeast. The mainland part consists of Cambodia, Laos, Myanmar, Vietnam and Thailand. The maritime part consists of Singapore, Brunei, East Timor, Indonesia, Malaysia and the Philippines. The area extends more than 3,300 km from north to south and 5,600 km from east to west, stretching from 30° northern to 11° southern latitude and from 92° to 142° eastern longitude (Holloway & Hall, 1998). The climate of Southeast Asia is essentially hot-tropical all year round. Precipitation is high, but