Balsaminaceae in Madagascar and the Comoro Islands, Fourteen Species of Impatiens (Balsaminaceae) Are Described As New (I

Balsaminaceae in Madagascar and the Comoro Islands: Systematics, Evolution, and Biogeography by Marie Elisette Rahelivololona Cover: Impatiens nomenyae Eb.Fisch. & Raheliv. (© E. Fischer) Balsaminaceae in Madagascar and the Comoro Islands: Systematics, Evolution, and Biogeography by Marie Elisette Rahelivololona from Mahajanga Submitted Dissertation thesis for the partial fulfillment of the requirements for a Doctor of Natural Sciences Fachbereich 3: Mathematik/Naturwissenschaften Universität Koblenz-Landau December 2018 1. referee: Prof. Dr. Eberhard Fischer 2. referee: Prof. Dr. Wilhelm Barthlott Date of defence: 06.02.2019 Head of the comission: Prof. Dr. Rainer Graafen Additional Referees: Prof. Dr. Sylvain Razafimandimbison Dr. Steven Janssens Table of Contents Table of Contents Chapter 1. Introduction 1 Chapter 2. New taxa of Impatiens (Balsaminaceae) from Madagascar I 13 Chapter 3. New taxa of Impatiens (Balsaminaceae) from Madagascar II. 42 A collection from Masoala Peninsula Chapter 4. Balsaminaceae. In: The Natural History of Madagascar 60 Chapter 5. New taxa of Impatiens (Balsaminaceae) from Madagascar III 69 Chapter 6. A new epiphytic species of Impatiens (Balsaminaceae) from 90 the Comoro Islands Chapter 7. New taxa of Impatiens (Balsaminaceae) from Madagascar IV 94 Chapter 8. New taxa of Impatiens from Madagascar V. New species of 152 Impatiens from Masoala Peninsula Chapter 9. New taxa of Impatiens (Balsaminaceae) from Madagascar VI. 171 Impatiens otto-eleonorae, a new species from Masoala Peninsula, and notes on the taxonomic relationships of Impatiens firmula and I. hildebrandtii Chapter 10. New taxa of Impatiens (Balsaminaceae) from Madagascar 181 VII. Two new species of Impatiens from Mt. Marojejy, Madagascar Chapter 11. New taxa of Impatiens (Balsaminaceae) from Madagascar 192 VIII. Impatiens max-huberi, a new species from Marojejy and Anjanaharibe-Sud Chapter 12. New taxa of Impatiens (Balsaminaceae) from Madagascar 198 IX. Impatiens lutzii, a new species from Montagne d’Ambre National Park Table of Contents Chapter 13. Impatiens galactica (Balsaminaceae), a new spurless 207 species of section Trimorphopetalum from Madagascar Chapter 14. Impatiens sielmannii (Balsaminaceae), a new epiphytic 216 species from Madagascar Chapter 15. Impatiens stefan-vogeli (Balsaminaceae), a new species 226 from Madagascar Chapter 16. Phylogeny and biogeography of Balsaminaceae inferred 236 from ITS sequences Chapter 17. Phylogeny, infrageneric classification and species 266 delimitation in the Malagasy Impatiens (Balsaminaceae) Chapter 18. Conclusions 288 Chapter 19. Zusammenfassung 292 Acknowledgements 296 References 298 Declaration on the own contributions on the submitted papers 311 Curriculum Vitae 314 Declaration Chapter 1 — Introduction Chapter 1 Introduction General introduction The family Balsaminaceae includes only two genera, the monospecific Hydrocera Blume ex Wight & Arn. and its species-rich sister Impatiens. The latter genus contains more than 1000 species. Plants are more or less succulent, annual, or perennial with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves, helophytic or mesophytic, with solitary flowers, or aggregated in cymose inflorescences (Grey-Wilson 1980a, Song 2006). The two genera are easily distinguished from each other: Hydrocera has free petals and an indehiscent ‘berry- like’ fruit, whereas Impatiens has the petals variously united, and a characteristic, explosive, dehiscent capsule. Other generic names such as Petalonema Peter, Semeiocardium Zoll., Trimorphopetalum Baker and Impatientella H.Perrier, are synonyms of Impatiens (Grey-Wilson 1980a; Rao et al. 1986). Hydrocera is restricted to the Indo- Malesian countries, while Impatiens occurs in Asia, Africa, Madagascar, India, Europe and North America, and is absent from Australia and South America. Its hotspots of diversification are in tropical and subtropical montane regions of the Old World. Most traditional classifications based on morphology place Balsaminaceae in Geraniales and suggest that it is closely related to Tropaeolaceae and Geraniaceae (Cronquist 1981, Thorne 2000), while some important taxonomists treated Balsaminaceae as an order of its own: the Balsaminales (Dahlgren 1989, Takhtajan 1997). However, recent molecular analyses indicate that Balsaminaceae belong to the order Ericales (Morton et al. 1996, 1997, Soltis et al. 2000, Albach et al. 2001, Anderberg et al. 2002, Bremer et al. 2002, Geuten et al. 2004). Both plastid (rbcL) and nuclear (18S rDNA) or combined sequence data suggest a close relationship between Balsaminaceae and Tetrameristaceae, Marcgraviaceae, and Fouquieriaceae (Song 1 Chapter 1 — Introduction 2006). This knowledge is helpful to choose suitable outgroups for the phylogenetic analyses of Balsaminaceae. General Ecology and Distribution Impatiens is nearly always associated with wet and humid places, frequently growing along the sides of rivers and streams, in grassy marshlands, or in the spray zones of waterfalls and cascades. It is rare to find Impatiens growing by stagnant waters. Most species are terrestrial and a few are epiphytic. It is a large genus with over 1000 species scattered throughout in the highlands and mountains of the tropical and subtropical regions of the Old World. It has five conspicuous diversity hotspots: tropical Africa (ca. 120 spp. Grey-Wilson 1980a); Madagascar (ca. 260 spp. Fischer & Rahelivololona, 2002, 2003, 2004, 2007, Fischer et al. 2003, 2015a, 2015b, 2015c, 2016, 2017, 2018 submitted); southern India and Sri Lanka (ca. 220 spp., Bhaskar & Razi, 1981); the eastern Himalayas (ca. 120 spp.), and Southeast Asia in its broad sense (including Myanmar, Thailand, southwest China, the Indochina peninsula, and the Malesian archipelago, ca. 250 spp.). There are 7 native species in North America with two species reaching Central America (Mexico: Impatiens mexicana Rydb., Costa Rica: Impatiens turrialbana J.D.Sm.) 1 native species in Europe (Impatiens noli- tangere L.) and 1 native species in boreal Eurasia (Impatiens parviflora DC.). Numerous invasive species have been introduced to Europe, including Impatiens capensis Meerb. from North America, Impatiens parviflora L. from Boreal Eurasia, and Impatiens glandulifera Royle from the Himalayas. There are no native species in South America or Australia. However, Impatiens walleriana Hook.f. from East Africa is now a neophytic invasive weed in tropical America. Morphology and Character Evolution of Impatiens Members of Impatiens are primarily annual or perennial herbs. The stems are typically rather thick and fleshy and often semi-translucent, only rarely becoming thickened and woody. Some species are rhizomatous or tuberous. While most species in Africa and Madagascar are perennials (Grey Wilson 1980, Perrier de la Bâthie 1934), most Asian species are annuals (Ruchisansakun et al. in press). Tubers occur in African taxa (Impatiens etindensis Cheek & Eb.Fisch. 1999) and in Madagascar (e.g. Impatiens 2 Chapter 1 — Introduction tuberosa H.Perrier, I. barthlottii Eb.Fisch. & Raheliv., I. loki-schmidtiae Eb.Fisch. & Raheliv.). While the majority of Impatiens species have spirally arranged leaves, only few taxa dispose of opposite or verticillate leaves. The leaves, which are generally simple with a dentate margin often have short hair-like appendages at the leaf base that are functional extrafloral nectaries. Whereas the teeth become less distinct when reaching the leaf base, the appendages become slightly larger often changing into glandular structures on the leaf petiole (Janssens 2008). Most of the Impatiens species possess a long leaf petiole, yet this can sometimes be absent. Leaf shape varies from linear- lanceolate to elliptic and sometimes peltate. The leaves are generally becoming membranaceous, often almost transparent when dried. The genus Impatiens has remarkable flowers that show a fascinating diversity in shape and colours. They are monosymmetric and usually resupinate through twisting of the pedicel. In most species each flower has three sepals, two of which are usually small- sized lateral sepals and one petaloid lower sepal that is modified into a nectary-tipped spur. In most Impatiens, there are only 2 lateral sepals (one pair) which are narrow- lanceolate as in I. rubrostriata Hook.f. or broadly ovate as in I. purpurea Hand.-Mazz. Some species have 4 lateral sepals (2 pairs), and usually the inner pair is linear, smaller and more narrow than the out pair. The occurrence of 4 lateral sepals is not common in Impatiens but exists in different regions: 7 African Impatiens species out of c. 120 (Grey-Wilson 1980b), and 2 species in Madagascar (Fischer & Rahelivololona, 2003, i.e. I. kuepferi Eb.Fisch. & Raheliv. and I. wohlhauseri Eb.Fisch. & Raheliv.). The number and the form of lateral sepals is a key character for the classification in Impatiens. The lower sepals of Impatiens exhibits an extraordinary wide range of variation in form and size: they typically present a shallow naviculate lower sepal constricted into a slender filiform or short cylindrical obtuse spur, and on the other hand the bird pollinated taxa assigned by Perrier de la Bâthie (1934) to the Humblotianae-group have a deeply navicular, large bucciniform or saccate lower sepal, much longer and gradually merging into, or rather abruptly extending into the spur. These bird-pollinated taxa are represented in Africa (e.g. Impatiens niamniamensis Gilg, Grey-Wilson 1980a), Madagascar (e.g. Impatiens humblotiana Baill.,

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