Studies on Systematics, Morphology and Taxonomy of Caiophora and Reproductive Biology of Loasaceae and Mimulus (Phrymaceae)
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Studies on systematics, morphology and taxonomy of Caiophora and reproductive biology of Loasaceae and Mimulus (Phrymaceae) 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 Markus Ackermann aus Buchen (Odenwald) Berlin 2011 This work was carried out between 2002 and 2011 under the supervision of Prof. Dr. Maximilian Weigend Institut für Biologie of the Freie Universität Berlin. 1st Reviewer: Univ.- Prof. Dr. Maximilian Weigend 2nd Reviewer: Univ.- Prof. Dr. Hartmut H. Hilger Date of defence: 12.07.2012 Contents Introduction 11 1. Introduction 12 1.1. The family Loasaceae 12 1.2. The genus Caiophora 15 1.3. The Uplift of the Andes and the diversification of species 27 1.4. Pollination biology of Loasoideae (Loasaceae) and Mimulus (Phrymaceae) 28 1.5. Hybridization in Caiophora and Mimulus 29 1.6. Working hypothesis 31 1.7. Objectives of this study 31 1.8. Overview of the dissertation 32 1.9. Literature 33 Taxonomy and Systematics 46 2. A preliminary phylogeny of Loasaceae subfam. Loasoideae (Angiospermae: Cornales) based on trnL(UAA) sequence data, with consequences for systematics and historical biogeography* 47 2.1. Abstract 47 2.2. Introduction 48 2.3. Material and methods 54 2.4. Results 54 2.5. Discussion 63 Systematics and plausibility of the molecular analyses 63 Historical biogeography 67 2.6. Acknowledgements 68 2.7. Literature 68 3. Notes on the genus Caiophora (Loasoideae, Loasaceae) in Chile and neighbouring countries* 71 3.1. Abstract 71 3.2. Introduction 72 3.3. Material and Methods 73 3.4. Results and discussion 74 3.5. Acknowledgements 98 3.6. Literature 98 3.7. Index of collectors 100 4. Catalogue of the Vascular Plants of the Southern Cone (Argentina, Southern 6 Brazil, Chile, Paraguay, and Uruguay): Loasaceae* 103 4.1. Blumenbachia Schrad. 104 4.2. Caiophora C. Presl 108 4.3. Huidobria Gay 116 4.4. Loasa Adans. 117 4.5. Mentzelia L. 130 4.6. Nasa Weigend 133 4.7. Scyphanthus Sweet 134 4.8. Literature 135 4.9. Abbreviations 137 5. Catalogue of the Vascular Plants of Bolivia: Loasaceae* 138 5.1. Loasaceae Juss. 138 5.2. Caiophora C. Presl 138 5.3. Gronovia L. 143 5.4. Klaprothia Kunth in Humb. 143 5.5. Mentzelia L. 144 5.6. Nasa Weigend 145 5.7. Literature 146 5.8. List of Synonyms: 147 Anatomy and Morphology 150 6. The seeds of Loasaceae subfam. Loasoideae (Cornales) II: Seed morphology of „South Andean Loasas“ (Loasa, Caiophora, Scyphanthus and Blumenbachia)* 151 6.1. Abstract 151 6.2. Introduction 152 6.3. Material and methods 155 Seed material 155 Storage 156 Microscopy 156 Descriptions 156 6.4. Results 157 Blumenbachia sect. Angulatae 160 Blumenbachia sect. Blumenbachia 160 Blumenbachia sect. Gripidea 163 Loasa ser. Loasa (syn. Loasa ser. Acanthifoliae) and Loasa ser. Macrospermae 163 Loasa ser. Deserticolae 164 Loasa ser. Floribundae 164 7 Loasa ser. Volubiles 168 Scyphanthus 170 Loasa ser. Acaules 172 Loasa ser. Pinnatae 172 Caiophora 175 6.5. Discussion 176 6.6. Acknowledgements 179 6.7. Literature 180 Floral ecology and hybridization 182 7. Hybridization and crossability in Caiophora (Loasaceae subfam. Loasoideae): Are interfertile species and inbred populations results of a recent radiation?* 183 7.1. Abstract 183 7.2. Introduction 184 7.3. Materials and methods 188 Sampling 188 Experimental crosses 189 Fruit set 192 Seed masses 193 Seed germination 193 7.4. Results 193 Fruit set 193 Seed masses 196 Germination 196 7.5. Discussion 198 Possibility of interspecific hybridization 198 Premating isolating mechanism and the causes of hybridization in nature 199 7.6. Conclusion 200 7.7. Acknowledgement 201 7.8. Literature 201 8. Nectar, Floral Morphology and Pollination Syndrome in Loasaceae subfam. Loasoideae (Cornales)* 207 8.1. Abstract: 207 8.2. Introduction 208 8.3. Materials and Methods 211 Plant material 211 Total nectar amount of individual flowers 214 8 Pollinator observations 217 Correlates of nectar production 217 8.4. Results 217 Total nectar amount 217 Altitudinal distribution, floral morphology and pollinators 219 8.5. Discussion 220 Overall nectar and sugar production in relation to pollination syndrome 220 Pollination, nectar and elevation 221 Evolution of nectar characteristics and pollination syndromes 221 8.6. Acknowledgements 222 8.7. Literature 222 9. Reloading the revolver – male fitness as a simple explanation for complex reward partitioning in Nasa macrothyrsa (Loasaceae, Cornales)* 228 9.1. Abstract 228 9.2. Introduction 229 9.3. Material and methods 230 Plant material and cultivation 230 Glasshouse experiments 232 Field observations 233 Statistics 233 9.4. Results 233 Flower longevity and self-pollination 233 Nectar replenishment 234 Thigmonastic stamen movement 235 Flower visits in nature 236 9.5. Discussion 236 9.6. Acknowledgements 238 9.7. Literature 238 10. Chilean Mimulus (Phrymaceae) – premating isolating mechanisms and floral ecology of flashy coloured Monkey flowers* 241 10.1. Introduction 241 10.2. Materials and methods 245 UV-patterns 245 Nectar production and floral size 245 Pollen number and Pollen-ovule ratio (P/O-ratio) 247 Experimental pollination 247 10.3. Results 248 Display size, visible and UV-patterns 248 9 Nectar production 248 P/O-ratio 249 Experimental pollination 251 10.4. Discussion 251 UV-patterns 251 Nectar production 252 P/O-ratio 254 Experimental pollination 254 10.5. Conclusions 256 10.6. Literature 257 11. Conclusions 264 11.1. Morphology, Taxonomy and Systematics 264 11.2. Floral ecology and hybridization 265 11.3. Literature 269 Summary 273 Zusammenfassung 276 Contribution to Chapters 279 Curriculum Vitae 281 Publication List 286 Congress Contributions 288 Appendix 289 Appendix A. (Seed morphology of „South Andean Loasas”) 289 Appendix B (Hybridization and crossability in Caiophora) 292 Appendix C (Pollination syndrome in Loasoideae) 294 Introduction Cover illustration: The cover inset shows a balloon-shaped funnel revolver flower ofCaiophora chuquitensis from the High Andes. Nectar characters and floral morphology indicate it is hummingbird- pollinated. Within the monophyletic Loasa-Caiophora clade, switches from short-tongued bee- to hummingbird-pollination appear to have taken place over time. See Ackermann M, Weigend M. 2006. Annals of Botany 98(3): 503–514 http://aob.oxfordjournals.org/content/98/3.cover- expansion (Chapter 8). Chapter 1 — Introduction 12 1. Introduction 1.1. The family Loasaceae The family Loasaceae is largely restricted to the Neotropics. The first taxa were named by Linné (1753) in “Species Plantarum” as Gronovia scandens (1753: 202) and Mentzelia aspera (1753: 516). In 1763 Adanson described the eponymic genus Loasa (1763). Several years later De Jussieu (1804) described the new family Loasaceae to unite the genera Mentzelia and Loasa. Since then, several new species and genera have been described. In 1900 Urban and Gilg divided the Loasaceae into three subfamilies: Gronovioideae, Mentzelioideae and Loasoideae. Several molecular studies (e.g. Moody et al., 2001; Hufford et al., 2003; Schenk and Hufford, 2010) largely confirm this subfamilial classification (compare current classification; tab 1.1). In their monumental “Monographia Loasacearum” Urban and Gilg (1900) described more than 80 new species, and their work is still fundamental for taxonomic studies in Loasaceae. Since then, several new genera have been described: Aosa Weigend, Chichicaste Weigend, Fuertesia Urb., Nasa Weigend, Plakothira J.Florence, Presliophytum (Urb. & Gilg) Weigend, Schismocarpus S.F.Blake and Xylopodia Weigend (Urban, 1910; Blake, 1918; Florence, 1985; Weigend, 1996, 1997c, 2006; Weigend et al., 2006). When Weigend (1996, 1997a, b, c) began with his studies about Loasaceae, he revised numerous herbarium specimens and conducted several field trips to South America, that led to collections of unknown species. He and his co-authors described more than 60 new taxa and recombined some 70 taxa, most of them belonging in Loasaceae subfam. Loasoideae (e.g. Weigend, 1996, 1997a, c, 1998, 1999, 2000, 2001a, 2002a, 2004b, 2006, 2007; Weigend et al., 1998, 2003; Dostert and Weigend, 1999; Weigend and Ackermann, 2003; Rodríguez, 2008; Henning and Weigend, 2009, 2011; Henning et al., 2011). Today, Loasaceae comprises 20 genera with more than 300 species (Weigend, 2004c, 2010a). The distribution of the family is largely restricted to the New World. The northernmost species are found in the genus Mentzelia (subfamily Mentzelioideae), ranging from Argentina/ Chile to the northern United States, whereas the southern distribution limit is formed by species of the genera Loasa and Blumenbachia (Fig. 1.1 C), distributed from Brazil to Patagonia (Chile and Argentina). The tribe Loaseae (subfam. Loasoideae) is restricted to Central and South America (Fig. 1.1 B) and the tribe Klaprothieae from northern Bolivia up to central Mexico (Fig. 1.1 A). There are only two genera outside of the Americas: Kissenia with two species in Africa (Dandy, 1926) and Plakothira with three species on Marquesas Islands (Florence, 1985). The subfamily Loasoideae, with 13 genera and more than 220 species (compare tab. Chapter 1 — Introduction 13 Subfamily Tribe Genus number of species Gronovioideae Cevallia Lag. 1 Fuertesia Urb. 1 Gronovia L. 2 Loasoideae Klaprothieae Klaprothia Kunth 2 Plakothira J.Florence 3 Xylopodia Weigend 1 Loaseae Aosa Weigend 7 Blumenbachia Schrad. 12 Caiophora C.Presl ca. 50 Chichicaste Weigend 1 Huidobria Gay 2 Kissenia R.Br. ex T.Anderson 2 Loasa Adans. 36 Nasa Weigend >100 Presliophytum (Urb. & Gilg) Weigend 3 Scyphanthus D.Don 1-2 Mentzelioideae Eucnide Zucc. 13 Mentzelia L. 80 Schismocarpus S.F.Blake 1 Petalonychoideae Petalonyx A.Gray 5 Table 1.1: Infrafamilial classification of Loasaceae (compiled from Hufford et al., 2003, 2005; Weigend, 1997c, 2004c, 2006; Weigend et al., 2004) 1.1), is the species-richest group within the family, with its centre of diversity in the Andes of Peru (Weigend, 2002b). Nasa, the largest genus within the subfamily, is most diverse in the Amotape-Huancabamba Zone (northern Peru: Weigend, 2002b; Weigend et al., 2004; Weigend and Gottschling, 2006; Henning et al., 2011).