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Contribution to the Biosystematics of Celtis L. (Celtidaceae) with Special Emphasis on the African Species

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Contribution to the Biosystematics of Celtis L. (Celtidaceae) with Special Emphasis on the African Species Contribution to the biosystematics of Celtis L. (Celtidaceae) with special emphasis on the African species Ali Sattarian I Promotor: Prof. Dr. Ir. L.J.G. van der Maesen Hoogleraar Plantentaxonomie Wageningen Universiteit Co-promotor Dr. F.T. Bakker Universitair Docent, leerstoelgroep Biosystematiek Wageningen Universiteit Overige leden: Prof. Dr. E. Robbrecht, Universiteit van Antwerpen en Nationale Plantentuin, Meise, België Prof. Dr. E. Smets Universiteit Leiden Prof. Dr. L.H.W. van der Plas Wageningen Universiteit Prof. Dr. A.M. Cleef Wageningen Universiteit Dr. Ir. R.H.M.J. Lemmens Plant Resources of Tropical Africa, WUR Dit onderzoek is uitgevoerd binnen de onderzoekschool Biodiversiteit. II Contribution to the biosystematics of Celtis L. (Celtidaceae) with special emphasis on the African species Ali Sattarian Proefschrift ter verkrijging van de graad van doctor op gezag van rector magnificus van Wageningen Universiteit Prof. Dr. M.J. Kropff in het openbaar te verdedigen op maandag 26 juni 2006 des namiddags te 16.00 uur in de Aula III Sattarian, A. (2006) PhD thesis Wageningen University, Wageningen ISBN 90-8504-445-6 Key words: Taxonomy of Celti s, morphology, micromorphology, phylogeny, molecular systematics, Ulmaceae and Celtidaceae, revision of African Celtis This study was carried out at the NHN-Wageningen, Biosystematics Group, (Generaal Foulkesweg 37, 6700 ED Wageningen), Department of Plant Sciences, Wageningen University, the Netherlands. IV To my parents my wife (Forogh) and my children (Mohammad Reza, Mobina) V VI Contents ——————————— Chapter 1 - General Introduction ....................................................................................................... 1 Chapter 2 - Evolutionary Relationships of Celtidaceae ..................................................................... 7 R. VAN VELZEN; F.T. BAKKER; A. SATTARIAN & L.J.G. VAN DER MAESEN Chapter 3 - Phylogenetic Relationships of African Celtis (Celtidaceae) ......................................... 31 A. SATTARIAN & L.J.G. VAN DER MAESEN Chapter 4 - Macro-morphology of African Celtis (Celtidaceae) ..................................................... 45 A. SATTARIAN & L.J.G. VAN DER MAESEN Chapter 5 - Pollen morphology of African Celtis (Celtidaceae) ...................................................... 51 A. SATTARIAN; R.G. VAN DEN BERG & L.J.G. VAN DER MAESEN Published in Feddes Repertorium 117 (2006) 34-40 Chapter 6 - Endocarp morphology of African Celtis (Celtidaceae) ................................................. 57 A. SATTARIAN & L.J.G. VAN DER MAESEN Published in Blumea 51: 2 (2006). Chapter 7 - Revision of African Celtis (Celtidaceae) ...................................................................... 65 A. SATTARIAN; L.J.G. VAN DER MAESEN & J.J. WIERINGA Two new species have been described in Blumea 50 (2005) 499-503 Chapter 8 - The Celtis philippensis complex ................................................................................... 93 A. SATTARIAN & J.J. WIERINGA Chapter 9 - Conspectus of Celtis (Celtidaceae) species worldwide ................................................. 97 A. SATTARIAN & L.J.G. VAN DER MAESEN Chapter 10 - Summary and Conclusions ........................................................................................ 115 A. SATTARIAN & L.J.G. VAN DER MAESEN References ...................................................................................................................................... 119 Appendix 1 ..................................................................................................................................... 126 Appendix 2 ..................................................................................................................................... 130 Appendix 3 ..................................................................................................................................... 134 Samenvatting .................................................................................................................................. 137 Summary ........................................................................................................................................ 139 Acknowledgements ........................................................................................................................ 140 Curriculum vitae; Publications ....................................................................................................... 141 Summary in Farsi ........................................................................................................................... 142 VII VIII Chapter 1 ——————————— GENERAL INTRODUCTION Summary The research questions of the project are introduced. In this thesis, the results are presented of biosystematic studies in Celtis, particularly African Celtis, the type genus of Celtidaceae, formerly Ulmaceae-Celtidoideae (Elias 1970; Grudzinskaya 1967; Judd et al. 1994; Omori & Terabayashi 1991, 1993; Ueda et al. 1997; Song et al. 2001). The African species of Celtis have been revised. We introduce the history of the Ulmaceae family (morphology and phylogeny). 1.1 Phylogeny During the last decade, our knowledge and understanding of phylogenetic relationships within the Angiosperm has improved, this has been accomplished largely based on various studies (e.g. Chase et al. 1993; Chase and Cox 1998; Soltis et al. 1998; Hilu et al. 2003). The classifications of the Angiosperm phylogeny group (APG 1998; APG II 2003) have been on the basis of these molecular and phylogeny studies, leading to the recognition of a series of monophyletic orders and families. Despite of advances in understanding phylogenetic patterns, there is a need for more studies addressing the phylogeny and relationships between morphology and phylogeny based on DNA sequences (Endress et al. 2000; Nandi et al. 1998; Doyle & Endress 2000; Zanis et al. 2003; Walter et al. 2004). What are the tools available today? Phylogenetic methods and data used: DNA sequences, cpDNA, MP, ML, Bayes. There are large advantages of using molecular data in phylogeny, the main ones are the large number and reliability of characters, or data power (e.g. Chase 1993). Moreover, homology assessment is easier for DNA sequence data than for morphology in angiosperm plants. Recent molecular-phylogenetic research has shown that relationships of the Urticales are different from what was based on morphological characters; here we briefly introduce some part of the phylogeny of Ulmaceae. Molecular data show the place of Urticalean families within a well- defined and supported Rosales (sensu APG 1998; including Barbeyaceae, Dirachmaceae, Eleagnaceae, Ramnaceae, and Rosaceae (Chase et al. 1993; Gunter, Kochert, & Giannasi 1994; Soltis et al. 1997; Qiu et al. 1998) and also shows that Celtis, Humulus, Trema , and Boehmeria in an Urticalean clade with Morus and Ficus and Urticalean clade is placed in their Rosid I clade, closely related to Rhamnaceae and Rosaceae (Chase et al. 1993). Molecular studies within Urticalean rosids included a restriction site mapping analysis of plastid DNA that indicated Ulmaceae were sister to the rest (Wiegrefe et al. 1998; Sytsma & Guries 1998). A preliminary rbcL analysis of the Ulmaceae and Celtidaceae (11 genera) and four genera representative of other Urticalean families provided strong support for a monophyletic Ulmaceae sister to a broadly paraphyletic Celtidaceae including Urticaceae, Moraceae and Cannabaceae (Ueda et al. 1997). The three-gene analysis of Soltis et al. (2000) also provided strong support for the separation of Ulmaceae and Celtidaceae. A small-scale matk analysis of Celtidaceae (5 genera of Celtidaceae and 8 other genera of Urticalean rosids) placed Cannabaceae solidly within a portion of the Celtidaceae (Song et al. 2001). 1 Chapter 1. General Introduction Table 1.1. Main history of Ulmaceae Mirbel. Author Date Family Genera Mirbel 1815 Ulmaceae Mirb. s.l. Ulmus, Holoptelea, Phyllostylon, Planera, Hemiptelea, Zelkova, Lozanella, Ampelocera, Celtis, Pteroceltis, Trema, Parasponia, Aphanthe, Gironniera, Chaetachme, Plagioceltis, Chaetoptelea, Barbeya. Mirandaceltis Link 1831 Ulmaceae Mirb. s.s. Ulmus, Holoptelea, Phyloptelea, Phyllostylon, Planera, Hemiptelea, Zelkova. Celtidaceae Link Lozanella, Ampelocera, Celtis, Pteroceltis, Trema, Parasponia, Aphananthe, Gironniera, Chaetachme Engler 1907 Ulmoideae Ulmus, Holoptelea, Phyloptelea, Phyllostylon, Planerar, Hemiptelea, Zelkova. Celtidoideae Lozanella, Ampelocera, Celtis, Pteroceltis, Trema, Parasponia, Aphanathe, Gironniera, Chaetachme, Mirandaceltis Grudzinskaya 1965 Ulmaceae Mirb. s.s. Ulmus, Holoptelea, Hemiptelea, Zelkova, Planera, Phyllostylon. Celtidaceae Link Celtis, Pteroceltis, Trema, Parasponia, Lozanella, Aphananthe, Gironniera, Chaetachme, Ampelocera Hutchinson 1967 Ulmeae Ulmus, Phyllostylon, Planera, Holoptelea, Celteae Celtis, Mirandaceltis, Pteroceltis, Ampelocera, Zelkova, Hemiptelea, Trema, Parasponia, Aphananthe, Gironniera, Chaetachme, Lozanella Sweitzer 1971 Ulmoideae Ulmus, Holoptelea, Phyllostylon, Planera, Hemiptelea, Soepadmo 1977 Zelkova, Ulloa 1995 Celtidoideae Lozanella, Ampelocera, Celtis, Pteroceltis, Trema, Parasponia, Aphanathe, Gironniera, Chaetachme Ulmaceae and Celtidaceae are separate families (Sytsma et al. 2002) while Cannabaceae and Celtidaceae should be merged. The origin of Cannabaceae within a clade of Celtidaceae is supported by ultrastructure, chromosome number, and cpDNA restriction site (Wiegrefe et
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