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Balsaminaceae in Madagascar and Comoro Islands: Systematics, Evolution, and Biogeography

  • In this thesis, I present the results of my studies on taxonomy, systematics, and biogeography of Impatiens (Balsaminaceae) in Madagascar and the Comoro islands. In Chapter 1 I reviewed the literature on taxonomy and classification of Balsaminaceae, on habitat, world distribution, morphology, molecular phylogenetics and infrageneric classification of the genus Impatiens. In Chapters 2-15 (Fischer & Rahelivololona 2002, 2003, 2004, 2007, 2015a, 2015b, 2015c, 2016, Fischer et al. 2004. 2017, 2018a, b submitted, Rahelivololona et al. 2003) I presented the first results of a revision of Balsaminaceae of Madagascar and the Comoro islands including the description of 78 new species. In Chapter 16 (Yuan et al. 2004) we worked on the phylogeny and biogeography of Balsaminaceae inferred from ITS sequences using combined results from molecular phylogenetic and morphological analyses. In Chapter 17 (Rahelivololona et al. 2018) we conducted a phylogeny and assessment of the infrageneric classification of species in the Malagasy Impatiens (Balsaminaceae) with a particular emphasis on taxa collected from Marojejy. Below I summarise the most important findings of each chapter and provide an outlook for future studies. How many species of Impatiens occur in Madagascar and the Comoro islands? To provide additional information on the taxonomic revision of Impatiens in Madagascar and the Comoro islands, the identification of already described species as well as the description of new species was conducted. Based on herbarium specimens from BR, G, K, NEU, P, TAN and on living plants collected during several field trips, 78 new species and 6 nomina nova have been published and another 70 new taxa are already identified. Actually more than 260 species occur in Madagascar and the Comoro islands and all of them are endemic. For each species, a description of the morphology, phenology, ecology and known distribution range was provided. Apart from new taxa, the delimitation of already described species like Impatiens firmula Baker and Impatiens hildebrandtii Baill. could be clarified by studying the types and by observing the variability in the field. Are the groups of Impatiens in Madagascar monophyletic, and what is the systematic position of Trimorphopetalum? Yuan & al. (2004) conducted a molecular phylogenetic study to examine the morphological and karyological evolution, and the historical biogeography of the Balsaminaceae family by using nucleotide sequence data of internal transcribed spacer regions of nuclear ribosomal DNA. The results support the monophyly of the Malagasy endemic section Trimorphopetalum and show that the cleistogamous Impatiens inaperta should be included in the sect. Trimorphopetalum which is the most derived within Impatiens. Therefore, the section Preimpatiens proposed by Perrier de la Bâthie (1934) is paraphyletic. Rahelivololona & al. (2018) provided a phylogenetic study focused on three subdivisions (based on macromorphological characters) proposed by Perrier de la Bâthie (1934). The analysis was done using two nuclear AP3/DEF homologues (ImpDEF1 and ImpDEF2) and the plastid atpB-rbcL spacer to reassess or assess the monophyly of the Malagasy Impatiens, of the sections Preimpatiens (Humblotianae and Vulgare groups) and Trimorphopetalum. A focus was on the species of Impatiens from the Marojejy National Park and of the morphologically variable species I. elatostemmoides, I. “hammarbyoides”, I. inaperta and I. manaharensis, using monophyly as the primary criterion. As results the Malagasy Impatiens are paraphyletic and the section Preimpatiens sensu Perrier de la Bâthie (1934) (= subgen. Impatiens sensu Fischer & Rahelivololona 2002) was not resolved as a monophyletic group. The section Trimorphopetalum sensu Perrier de la Bâthie (1934) (= subgen. Trimorphopetalum sensu Fischer & Rahelivololona 2002), however, was strongly confirmed as a monophyletic lineage (BS: 92; BPP: 1). Neither the Humblotianae group nor the Vulgare group was supported as monophyletic. None of the morphologically variable species appeared to be monophyletic and the sampled species of Impatiens from the Marojejy National Park do also not form a monophyletic group. What are the biogeographical position and the distribution patterns of Impatiens in Madagascar and the Comoro islands? Investigation of the geographical affinities and species distribution of section Impatiens (including Humblotianae group and Vulgare group) and section Trimorphopetalum were conducted and the origin and evolution as well as species richness and endemism were discussed. The isolation, the climate and the complex topography of Madagascar have generated the microhabitats and ecological niches favourable to the diversification of Impatiens species. Impatiens of Madagascar with 260 endemic species is actually the largest genus in Madagascar. Therefore, Madagascar and the Comoro islands are among the most species-rich regions in the world for Impatiens. Future studies In Impatiens on Madagascar, there remain numerous unresolved questions that need to be adressed: • A further study based on a much larger molecular data set and sampling from the entire geographic ranges of Impatiens in Madagascar is needed to retest the monophyly of the different subgenera and sections, as well as a molecular dating of the Malagasy Impatiens. • The study of pollinators as a key for understanding the radiation and species richness is required: Within Impatiens the different shapes of spur are related to pollinators (bees, birds, butterflies and moths). Therefore pollinator observation of specific species need to be done to understand the radiation of species by adaptation and coevolution with these pollinators. A pollination study with a large number of species within section Trimorphopetalum will help to understand the mechanism of complete disappearance of the spur, the shift of pollinators and the evolution of species richness. • The destruction of the natural habitats of Impatiens and the subsequent reduction of humidity in logged area constitute a severe threat for the survival of many species. The conservation and reforestation of vulnerable areas such as Ankaratra, Daraina, Mandraka and Col des Tapia near Antsirabe is required. • In terms of conservation and to mitigate the threat on the genus, a study on the ex-situ-conservation of Malagasy Impatiens species is very important as long as some species are suitable for horticultural purposes (e.g. Impatiens mayae-valeriae, Impatiens emiliae and species with broad red spur). • Finally, the publication of the revision of Impatiens of Madagascar and the Comoro islands will help other botanists to identify the species and will thus increase our knowledge on the group.

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Author:Marie Elisette Rahelivololona
Referee:Eberhard Fischer, Wilhelm Barthlott
Advisor:Eberhard Fischer
Document Type:Doctoral Thesis
Date of completion:2019/02/07
Date of publication:2019/02/08
Publishing institution:Universität Koblenz-Landau, Universitätsbibliothek
Granting institution:Universität Koblenz-Landau, Campus Koblenz, Fachbereich 3
Date of final exam:2019/02/06
Release Date:2019/02/08
Number of pages:314
Institutes:Fachbereich 3 / Institut für Integrierte Naturwissenschaften / Institut für Integrierte Naturwissenschaften, Abt. Biologie
Licence (German):License LogoEs gilt das deutsche Urheberrecht: § 53 UrhG