Evolution, Biogeography, and Monographic Treatment of Coccinia (Cucurbitaceae)
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Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität München Evolution, Biogeography, and Monographic Treatment of Coccinia (Cucurbitaceae) Norbert Holstein Munich, April 2012 ii Gutachter: 1. Gutachter: Prof. Dr. Susanne S. Renner 2. Gutachter: Prof. Dr. Günther Heubl 3. Gutachter: Prof. Dr. Gert Wörheide 4. Gutachter: Prof. Dr. John Parsch Dissertation eingereicht am: 30. April 2012 Mündliche Prüfung am: 18. Juli 2012 iii Erklärung Diese Dissertation wurde im Sinne von §12 der Promotionsordnung von Prof. Dr. Susanne S. Renner betreut. Ich erkläre hiermit, dass die Dissertation nicht einer anderen Prüfungskommission vorgelegt worden ist und dass ich mich nicht anderweitig einer Doktorprüfung ohne Erfolg unterzogen habe. Ehrenwörtliche Versicherung Ich versichere hiermit ehrenwörtlich, dass die vorgelegte Dissertation von mir selbständig, ohne unerlaubte Hilfe angefertigt wurde. Norbert Holstein iv Note In this thesis, I present the results of my doctoral research, which took place from September 1, 2007 to April 30, 2012, under the supervision of Prof. Dr. Susanne S. Renner. The results of my research led to four manuscripts of which three have been published (chapters 1 to 3), while a fourth (chapter 4) has yet to be submitted. I carried out all data collection and analyses, if not otherwise indicated in the corresponding acknowledgements. Writing and discussion of these manuscripts involved collaboration with Prof. Dr. S. S. Renner. List of Publications Papers Holstein, Norbert and S.S. Renner. 2010 [published 2011]. Coccinia (Cucurbitaceae) gains two new species from East Africa, three new synonyms, and one new combination. Kew Bulletin 65(3): 435–441. doi:10.1007/s12225-010-9229-9 Holstein, Norbert and S.S. Renner. 2011. A dated phylogeny and collection records reveal repeated biome shifts in the African genus Coccinia (Cucurbitaceae). BMC Evolutionary Biology 11: 28. doi:10.1186/1471-2148-11-28 Holstein, Norbert and S.S. Renner. 2011. Coccinia intermedia – a new species from West Africa. Phytokeys 7: 27–36. doi: 10.3897/phytokeys.7.2032 Holstein, Norbert. Monograph of Coccinia (Cucurbitaceae). Submitted to Phytokeys (April 25, 2012). Posters Holstein, Norbert and S.S. Renner. Neogene biome shifts in Africa as a driver of speciation in Coccinia (Cucurbitaceae)? Systematics 2009, 7th Biennial Conference of the Systematics Association, August 10–14, 2009, Leiden, Netherlands v Talks Presenting author is given in bold. Holstein, Norbert. February 16, 2012. How climate change in Africa over the past 7 million years affected diversification in the African cucurbit genus Coccinia. Seminar at the NCB Naturalis, Leiden, Netherlands Holstein, Norbert. October 12, 2011. How climate change in Africa over the past 7 million years affected diversification in the African cucurbit genus Coccinia. EES conference 2011, Munich, Germany Holstein, Norbert and S.S. Renner. February 25, 2011. Rapid filling of climatic niches in the African cucurbit Coccinia. Biosystematics 2011, Berlin, Germany Holstein, Norbert and S.S. Renner. September 18, 2011. African late Neogene climatic oscillations and speciation in Coccinia. 19th International Symposium “Biodiversity and Evolutionary Biology” of the German Botanical Society (DBG), Vienna, Austria Holstein, Norbert and S.S. Renner. November 4, 2009. Evolution of species ranges and habitats in Coccinia (Cucurbitaceae). Seminar of the Institute of Systematic Botany and Mycology, LMU Munich, Germany Holstein, Norbert and S.S. Renner. August 7, 2009. Evolution of Coccinia (Cucurbitaceae) and its sex chromosomes — first results. University of Dar es Salaam, Tanzania Holstein, Norbert and S.S. Renner. September 14–19, 2008. Evolution of sex chromosomes in Coccinia (Cucurbitaceae). EES Summer School “Evolution of sex chromosomes”, Frauenchiemsee, Germany Holstein, Norbert and S.S. Renner. December 3, 2008. Evolution of sex chromosomes in Coccinia (Cucurbitaceae) — a first report. Seminar of the Institute of Systematic Botany and Mycology, LMU Munich, Germany vi Funding My doctoral research was funded by grants to Prof. S. S. Renner from the Deutsche Forschungsgemeinschaft (RE 603/6-1 and 603/6-2). Travel to the herbaria in Paris, Brussels, and Vienna was supported by the Munich Graduate School for Evolution, Ecology and Systematics (EES). Travel to the herbarium of Leiden was supported by the SYNTHESYS project of the European Community Research Infrastructure Action, which is part of Framework Program 7. vii Index of Abbreviations ! = seen a.m. = ante meridiem (forenoon) bp = base pairs CA = California Cat. Print. = Catalogue du Printemps CVH = China Virtual Herbarium (http://www.cvh.org.cn/cms/) comm. = communicated by cpDNA = plastid DNA df = degrees of freedom DNA = desoxyribose nucleic acid doi = digital object identifier E = eastern e.g. = exempli gratia, for example ESS = effective sample size ex cult. = from cultivation F81 = Felsenstein 81 Fig. / fig. = figure fl = flowering fr = fruiting ft = feet G = goodness-of-fit value GTR = general time reversal Herb. = herbarium HPD = highest posterior density ibid. = ibidem (at the same place) ICBN = International Code of Botanical Nomenclature i.e. = id est, that is IICT = Instituto de Investigação Científica Tropical Inc. = Incorporated ITS = internal transcribed spacer IL = Illinois JPS = JStor Plant Science LB medium = Lysogeny broth medium l.c. = locus citatus LFY = LEAFY (gene homolog) L/W/H = length/width/height viii Ma = million years matK = maturase K gene MCMC = Markov chain Monte Carlo MI = Michigan ML = Maximum Likelihood mls = miles Monogr. Phan. = Monographiæ Phanerogamarum Ms. = manuscript Mss. = manuscripts Mt = Mount mt. = mountain Mts = Mountains Mus. Zeyl. = Musæum zeylanicum n = set of haploid chromosomes, such as in 2n = 24 N = northern ndhF = nicotinate hydroxylase subunit F gene nom. illeg. = nomen illegitimum nom. nud. = nomen nudum nov. = new n.v. = non vidi (not seen) P = P-value (significance value) PCR = polymerase chain reaction P × E = polar by equatorial diameter pers. comm. = personal communication p.p. = pro parte Prodr. Pl. Cap. = Prodromus Plantarum Capensium psbA = gene for coding the D1 polypeptide of the photosystem II reaction center rbcL = gene for ribulose-1,5-bisphosphate carboxylase oxygenase large subunit rpl = ribosomal protein large subunit gene rps = ribosomal protein small subunit gene S = southern s.coll. = without collector SD = standard deviation s.dat. = without date s.det.loc. = without detailed location (only country or province given) ser. = series s.loc. = without location ix s.n. = sine numero (without number) SNNPR: Southern Nations, Nationalities, and People’s Region sp. = species [singular] spec. nov. = species novus (new species) sphalm. = sphalmate (by mistake) spp. = species [plural] stat. nov. = status novus (new status, e.g. as subspecies instead of species) subsp. = subspecies Tab. / tab. = table trn = gene of a transcribed ribose nucleid acid (followed by the corresponding letter of the transferred amino acid) TVM = transversion model USA = United States of America UTM = Universal Transverse Mercator var. = variety W = western WGS84 = World Geodetic System from 1984 Herbarium abbreviations follow Index Herbariorum (http://sciweb.nybg.org/science2/IndexHerbariorum.asp), except CVH and JPS (see above). Abbreviations of author names follow International Plant Names Index (IPNI; http://www.ipni.org/). x xi Contents Erklärung ............................................................................................................................... iii Ehrenwörtliche Versicherung ................................................................................................. iii Note ...................................................................................................................................... iv List of Publications ................................................................................................................ iv Papers ............................................................................................................................... iv Posters .............................................................................................................................. iv Talks ................................................................................................................................... v Funding ................................................................................................................................. vi Index of Abbreviations .......................................................................................................... vii Contents ................................................................................................................................ xi Zusammenfassung ............................................................................................................ 1 Summary ............................................................................................................................... 2 General Introduction .............................................................................................................. 3 Introduction to the Study Group ......................................................................................... 3 Evolution of Habitat Preferences ........................................................................................ 6 Evolution of Sex Chromosomes ........................................................................................