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Diversity of the Genus Euphorbia (Euphorbiaceae) in SW Asia

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Diversity of the genus Euphorbia (Euphorbiaceae) in SW Asia Dissertation zur Erlangung des Doktorgrades Dr. rer. nat. an der Fakultät Biologie/Chemie/Geowissenschaften der Universität Bayreuth Amir Hossein Pahlevani Aus dem Iran, Tehran Bayreuth, 2017 Die vorliegende Arbeit wurde von April 2012 bis Dezember 2016 am Lehrstuhl Pflanzensystematik der Universität Bayreuth unter Betreuung von Frau Prof. Dr. Sigrid Liede-Schumann und Herr Prof. Dr. Hossein Akhani angefertig. Vollständiger Abdruck der von der Fakultät für Biologie, Chemie und Geowissenschaften der Universität Bayreuth genehmigten Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.). Dissertation eingereicht am: 15.12.2016 Zulassung durch die Prüfungskommission: 11.01.2017 Wissenschaftliches Kolloquium: 20.03.2017 Amtierender Dekan: Prof. Dr. Stefan Schuster Prüfungsausschuss: Prof. Dr. Sigrid Liede-Schumann (Erstgutachter) PD Dr. Gregor Aas (Zweitgutachter) Prof. Dr. Gerhard Gebauer (Vorsitz) Prof. Dr. Carl Beierkuhnlein This dissertation is submitted as a ‘Cumulative Thesis’ that includes four publications: Three published and one submitted. List of Publications 1. Pahlevani AH, Akhani H, Liede-Schumann S. Diversity, endemism, distribution and conservation status of Euphorbia (Euphorbiaceae) in SW Asia. Submitted to the Botanical Journal of the Linnean Society. (Revision under review). 2. Pahlevani AH, Liede-Schumann S, Akhani H. 2015. Seed and capsule morphology of Iranian perennial species of Euphorbia (Euphorbiaceae) and its phylogenetic application. Botanical Journal of the Linnean Society 177: 335–377. 3. Pahlevani AH, Feulner M, Weig A, Liede-Schumann S. 2017. Molecular and morphological studies disentangle species complex in Euphorbia sect. Esula (Euphorbiaceae) from Iran, including two new species. Plant Systematics and Evolution 4. Pahlevani AH, Riina R. 2014. Synopsis of Euphorbia subgen. Esula sect. Helioscopia (Euphorbiaceae) in Iran with the description of Euphorbia mazandaranica sp. nov. Nordic Journal of Botany 32: 257–278. Declaration of contribution to publications The thesis contains four research articles. The major parts of the underlying studies were carried out by myself at the University of Bayreuth and partly in the Department of Botany at Iranian Research Institute of Plant Protection. Most phylogenetic and statistics analyses were carried out with support of Prof. Dr. Sigrid Liede-Schumann. All manuscripts in this thesis have been prepared under consideration of the comments of all coauthors. 1st publication Pahlevani AH, Akhani H, Liede-Schumann S. Diversity, endemism, distribution and conservation status of Euphorbia (Euphorbiaceae) in SW Asia. Submitted to the Botanical Journal of the Linnean Society. Authors’ contribution: The idea and concept of the manuscript were developed by the first author with guidance of Prof. Dr. H. Akhani and Prof. Dr. S. Liede-Schumann. About 22,000 specimens in several herbaria (B, BM, E, FUMH, HSBU, IRAN, K, M, MSB, PR, SAV, TARI, TUH, UBT, W, WU) have been studied by the first author. Coordinates of all studied specimens have been extracted and transferred to the dataset by the first author. Several periods of field work have been carried out in Iran by the first author. Drawing The production of 243 maps, analysis of species richness, evaluation of conservation status of the endemic and subendemic species were exclusively performed by A.H. Pahlevani. Four new species, 13 new records, three synonyms were reported and described by A.H. Pahlevani. The manuscript was written by the A.H. Pahlevani taking into account the useful comments of Prof. Dr. S. Liede-Schumann and Prof. Dr. H. Akhani. 2nd publication Pahlevani AH, Liede-Schumann S, Akhani H. 2015. Seed and capsule morphology of Iranian perennial species of Euphorbia (Euphorbiaceae) and its phylogenetic application. Botanical Journal of the Linnean Society 177: 335–377. Authors’ contribution: The idea and concept of the manuscript were developed by the first author. Most parts of this manuscript were developed by A.H. Pahlevani, including identification and selection of the specimens used for the work, selection of the characters to be studied, photographing all seeds and capsules, carrying out all measurements, lab work, developing a key based on solely seed and capsule morphology both for the sections and all species, as well as describing two new species. Phylogenetic analysis was completed by the first author under the profound support of Prof. Dr. S. Liede-Schumann. SEM photography was supported by Prof. Dr. H. Akhani. The manuscript was prepared by A.H. Pahlevani taking into account the useful comments of Prof. Dr. S. Liede- Schumann and Prof. Dr. H. Akhani. 3rd publication Pahlevani AH, Feulner M, Weig A, Liede-Schumann S. 2017. Molecular and morphological studies disentangle species complex in Euphorbia sect. Esula (Euphorbiaceae) from Iran, including two new species. Plant Systematics and Evolution Authors’ contribution: The idea and concept of the manuscript were developed by the A.H. Pahlevani under guidance of Prof. Dr. S. Liede-Schumann. The collection of plant material was conducted by A.H. Pahlevani. ISSR Laboratory work was conducted in Dr. A. Weig laboratory. ISSR data analysis was done by A.H. Pahlevani and Dr. M. Feulner under supervision of Dr. A. Weig and Prof. Dr. Liede-Schumann. Phenetic analysis (PCA) was carried out by A.H. Pahlevani. Preparation of a key to the species, typification and taxonomic evaluation, as well as the description of two new species were carried out by A.H. Pahlevani. The manuscript was prepared by A.H. Pahlevani under recognition of the comments of all coauthors, especially the useful comments of Prof. Dr. S. Liede- Schumann. 4th publication Pahlevani AH, Riina R. 2014. Synopsis of Euphorbia subgen. Esula sect. Helioscopia (Euphorbiaceae) in Iran with the description of Euphorbia mazandaranica sp. nov. Nordic Journal of Botany 32: 257–278. Authors’ contribution: The idea and concept of the manuscript were developed by A.H. Pahlevani. Field work, visits to the herbaria housing the studied specimens, all taxonomic treatments, key construction and description of new species, have been undertaken by the first author. Typification was carried out by A.H. Pahlevani and R. Riina. Lastly, the paper was written by A.H. Pahlevani with the useful comments of R. Riina. Content Summary 1 Zusammenfassung 3 1. General Introduction 5 2. Synopsis 14 2.1. Material and Methods 14 2.2. Results and Discussion 21 2.3. Conclusion and Perspectives 32 2.4. References 34 Publications 1. Diversity, endemism, distribution and conservation status of Euphorbia (Euphorbiaceae) in SW Asia 43 2. Seed and capsule morphology of Iranian perennial species of Euphorbia (Euphorbiaceae) and its phylogenetic application 44 3. Molecular and morphological studies disentangle species complex in Euphorbia sect. Esula (Euphorbiaceae) from Iran, including two new species 45 4. Synopsis of Euphorbia subgen. Esula sect. Helioscopia (Euphorbiaceae) in Iran with the description of Euphorbia mazandaranica sp. nov. 46 Acknowledgements 47 Eidesstattliche Erklärung 48 Summary Summary Euphorbia is one of the three most species-rich genera of flowering plants with more than 2000 species occurring in all temperate and tropical regions. They occupy a wide range of habitats and exhibit great diversity in growth forms. In habit, the species vary from small ephemerals to various forms of herbaceous annuals or perennials, cushion-forming subshrubs, large shrubs, small trees and cactus-like succulents. After recent molecular studies, Euphorbia has been divided into four monophyletic subgenera including, Athymalus (c. 150 species), Chamaesyce (c. 600 species), Esula (c. 500 species) and Euphorbia (c. 800 species). Taxonomy of Euphorbia is extremely difficult due to the species richness accompanied by a cosmopolitan distribution, the extreme morphological plasticity among certain species and convergent evolution of certain morphological characters. As a consequence, the circumscription of the Euphorbia species especially in SW Asia remained unresolved. To resolve phylogenetic and taxonomic complex groups and uncertainties among the species of the genus Euphorbia, this thesis aims 1) to complete phylogenetic and phylogeographic study of the genus (subgen. Esula) in SW Asia, by integrating hitherto unanalyzed species in the existing phylogeny using the markers ITS and ndhF, 2) to study seed and capsule morphology of the Iranian species of Euphorbia and study their evolution by plotting them on the molecular phylogenetic tree, 3) to evaluate the genotypic diversity of some complex, rare or endemic Iranian Euphorbia species (section Esula) using ISSR DNA fingerprinting, 4) to trace diversification, endemism, distribution and conservation status of the genus Euphorbia in SW Asia, 5) to update the taxonomic treatment of the two largest sections, Helioscopia and Esula of the subgenus Esula in Iran. The phylogenetic relationships of Iranian species based on internal transcribed spacer (ITS) nuclear and ndhF plastid regions are updated and used for the characterization of the synapomorphies of each clade in subgenus Esula. It is confirmed that there are 13 sections in subgenus Esula in Iran. The presence or absence of granulate elements on seed surfaces represents a phylogenetically important trait for section delimitation. The capsule surface is synapomorphic for several sections, including Helioscopia (tuberculate-verrucose),
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