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Thesis)) Opengesneden En Bestudeerd Onder De Microscoop UvA-DARE (Digital Academic Repository) Evolutionary biology of Brunellia (Brunelliaceae, Oxalidales) Orozco Pardo, I.C. Publication date 2001 Document Version Final published version Link to publication Citation for published version (APA): Orozco Pardo, I. C. (2001). Evolutionary biology of Brunellia (Brunelliaceae, Oxalidales). Universiteit van Amsterdam. General rights It is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), other than for strictly personal, individual use, unless the work is under an open content license (like Creative Commons). Disclaimer/Complaints regulations If you believe that digital publication of certain material infringes any of your rights or (privacy) interests, please let the Library know, stating your reasons. In case of a legitimate complaint, the Library will make the material inaccessible and/or remove it from the website. Please Ask the Library: https://uba.uva.nl/en/contact, or a letter to: Library of the University of Amsterdam, Secretariat, Singel 425, 1012 WP Amsterdam, The Netherlands. You will be contacted as soon as possible. UvA-DARE is a service provided by the library of the University of Amsterdam (https://dare.uva.nl) Download date:11 Oct 2021 I I Evolutionaryy Biology of Brunellia ^i^ Pa#h (Brunelliaceaeff Oxalidalesl w w rozeoPar r c-% c % Evolutionaryy Biology of Brunellia Ruiz & Pavón (Brunelliaceae,, Oxalidales) --- +-_- Ai Evolutionaryy biology of Brunellia (Brunelliaceae,, Oxalidales) ACADEMISCHH PROEFSCHRIFT terr verkrijging van de graad van doctor aan de Universiteit van Amsterdam opp gezag van de Rector Magnificus Proff dr. J.J.M Franse tenn overstaan van een door het college voor promoties ingestelde commissie, inn het openbaar te verdedigen in de Aula der Universiteit opp dinsdag 4 december 2001 te 12.00 uur door r Claraa Inés O roze o Pardo Geborenn te Bogota (Colombia) Promotiecommissiee : Promotores:: Prof. Dr. H. Hooghiemstra and Prof. Dr. P. J. M. Maas i^^ Overige leden: Prof.. Dr. A. M. Cleef, Universiteit van Amsterdam Dr.. Ferry Bouman, Universiteit van Amsterdam Dr.. P. van Welzen, Leiden University Dr.. R. van der Ham, Leiden University Prof.. Dr. J. Woif. Universiteit van Amsterdam Prof.. Dr. F. Weberling, Ulm University, Germany Faculteitt Natuurwetenschappen, Wiskundee en Informatica Publisher:: Universiteit van Mmaterdam, Amsterdam Layy out : MCAV Diseho y Producción Grafica Pre-Press:: Punto Grafico Digital. Bogoia-Colombia Printing:: Tipografia y Litografia ïriana Thee research in this publication was made possible by financial support of the Universidad Nacional Colombia.. Bogota- Colombia, partially supported by the Comité de invesiigación v DesarroSlo Cien- tificoo (CINDEC) of the Universidad Nacional de Colombia and also partially supoorted Dy the Smithsoniann Institution and Amsterdam University. NoNo pari of this book may be reproduced in any format by print, photocopy, microfilm or by artoihe meansmeans -without the written permission from the publisher / author. ISBN:: 90-76894-07-S Claraa Ines Orozco P 2001 Too Dr. Thomas Van der Hammen andd my family r r «A* * Too the memory of Dr. José Cuatrecasas "1 1 1j j 1 —— - -- -^" - CONTENTS CONTENTS SUMMARY Y RESUMEN N CHAPTERR 1 Introduction Introduction CHAPTERR 2 SobreSobre la position sistemética de Brunellia Ruiz & Pavón CHAPTERR 3 LeafLeaf anatomy in Brunellia Ruiz & Pavón 32 CHAPTER4 4 AA comparative study of inflorescences in Brunellia Ruiz & Pavón andand related taxa 53 CHAPTERR 5 AA study of floral morphology in Brunellia Ruiz & Pavón ?2 CHAPTERR 6 PollenPollen morphology of Brunellia Ruiz & Pavón and related taxa 90 CHAPTERR 7 AA cladistic analysis of Brunellia Ruiz & Pavón based on morphological characterscharacters ^7 CHAPTERR 8 PhytogeographyPhytogeography 134 CHAPTERR 9 GeneralGeneral discussion and conclusions 159 fc-ft fc-ft SAMENVATTING G BruneiliaBruneilia is een geslacht van boomsoorten met een wijde neotropische verbreiding, in de tropische Andes vooraff m Colombia. Ook zijn er soorten m Centraal Amerika en de Caraïbische eilanden Vierenvijftig soortenn worden in deze studie formeel erkend. De algemene doelstelling van deze studie is het in termen vann evolutie verklaren van de waargenomen kenmerken in BruneUia, alsmede de patronen van variatie (Brunelliaceae).. Om dit doel te bereiken zijn verschillende disciplines van de biologie gebruikt die niet eerderr werden toegepast of tot de nu bereikte resolutie in de kennis van de groep leidden. Twee hoofdstukken betreffendee de fylogenie zijn ontwikkeld in deze studie: een met het presenteren van een hypothese betreffendee de systematische positie van Bruneltia, gebaseerd op morfologische kenmerken {hoofdstuk 2).. en een betreffende de evolutionaire relaties tussen de soorten van Brunellia {hoofdstuk 7). Patronen vann geografische verbreiding (hoofdstuk 8), gebieden van speciatie en diversificatie zijn vastgesteld op basiss van de fylogenetische studie van Brunellia in relatie met andere taxa en geologische gegevens van dee Andes, Centraal Amerika en het Caraibtsche gebied. Hernieuwd onderzoek van kenmerkende systemen omvattee de anatomie {hoofdstuk 3), patronen van bloeiwijzen {hoofdstuk 4), bloem morfologie (hoofdstuk 5) enn pollenmorfologie (hoofdstuk 6) om te worden gebruikt voor de studie van de evolutionaire relaties van de soortenn van het genus (hoofdstuk 7), De dieptestudie van deze systemen van kenmerken leverden zeer belangrijkee gegevens op vanuit het gezichtspunt van ontwikkeling en gedrag' in relatie met andere taxa gerelateerdd aan Brunellia. Inn Hoofdstuk 2, uitgaand van de hypothese dat Brunellia een genus van de Cunoniaceae is en niet het enigee geslacht in de familie van de Brunelliaceae, is de systematische positie van Bnjnetlia onderzocht waarbijj de fylogenetische relaties van dit taxon, alsmede 20 morfologische kenmerken vergelijkenderwijs geanalyseerdd zijn met de elf geslachten van de Cunoniaceae. Twaalf taxa maken deel uit van de mgroup. Hett programma Hennig86 gebaseerd op de methode van parsimonie werd gebruikt voor de selectie van de hypothesenn van relaties De gegevens van de ingroup werden eerst geanalyseerd met vier outgroups: Connams,Connams, Fothergilla. Davidsonia en Quercus, vervolgens met drie met uitsluiting van Quercus, en tenslotte mett elk van deze genera apart; in het laatste geval werden de kenmerken gepolariseerd De resultaten latenn zien dat Brunellia een taxon is dat verschillend is van de Cunoniaceae en dat de Cunoniaceae een niet-monophyletischeniet-monophyletische groep is. In deze studie wordt voorgesteld Brunellia, Spiraeanthemum en Acsmithia tee plaatsen in een familie verschillend van de Cunoniaceae. Echter, Bradford (pers. med), op basis van de SAMENVATTING SAMENVATTING resultatenn van moleculair onderzoek signaleert Bruneüia ooft als een natuurlijke eenheid onafhankelijk van SpiraesnthemumSpiraesnthemum en Acsmithia Meer onderzoek is nodig om de hypothese betreffende de relaties van lezeleze taxa te ondersteunen Ook wordt de monofyietische oogsprong van de Cunoniaceae ter discussie gesteldd door de aanwezigheid van Davidsonia binnen de interne groep De nauwe verwantschap tussen de basaiee Rosidae met de lagere Hamamelidae wordt bevestigd. De apomorfïe en plesiomorfie van enkele kenmerkenn wordt bediscussieerd. Van geen enkel tribus van Engler kon de monofyiie worden bevestigd. Eenn groot deel van de informatie die gebruikt is in de analyse van de evolutionaire verwantschappen van BruneüiaBruneüia (Hoofdstuk 7) is gebaseerd op de resultaten van de anatomische studie {Hoofdstuk 3). In d.t hoofdstukk wordt de bladanatomische studie van 24 soorten van Brunettta gepresenteerd, inclusief van stomataa en nodale anatomie. De nodaie anatomie van Brunetlia is unilocular Anatomische kenmerken van dee bladsteel en de bladschijf hebben taxonomisch belang De vaatcilinder van de bladsteel is gewoonlijk continuee in het midden en distale deel van de bladsteel, terwijl deze meestal discontinue is m het deel dicht bijj de knoop. Hett adaxiale deel van de vaatciiinder van de bladsteel is samengedrukt en vormen in wemtg soorten als het waree bogen. Schorsbundels zijn frequent in het distale en middendeel van de bladsteel. terwijl mergbundels alleenn in S.. foreroi aangetroffen worden. Onder de meest bruikbare kenmerken van de bladschijf zijn de aanwezigheid vann holtes of golvingen en de aanwezigheid van een hypodermis De reductie blaadjes om opvallende of weinigg opvallenae stipulae op de bladsteel te maken is in veel soorten met de verschijningsvorm van een enkell blad waargenomen. Het onderzoeksresultaat geeft aan dat veel soorten ten onrechte met enkelvoudig bladd werden beschouwd in feite hebben alle soorten van Bruneüia samengesteld bladeren De stomata zijn actinocytisch,, behalve in S. cufervens/s met anemocytische stomata Zoalss anatomische kenmerken zijn ook kenmerken van de archtectuur van bloeiwijzen {Hoofdstuk 4) gebruiktt voor de analyse van de evolutionaire verwantschappen van Bruneüia. Dit hoofdstuk omvat het onderzoekk aan 35 soorten, die tweederde van de soorten van deze monogenerische familie vertegenwoordigenn en representatief zijn voor de totale variatie van de bloeiwtjzen in dit genus. Het onderzoek werdd gerealiseerd om de bloeiwijzemorfologie te begrijpen en deze informatie te gebruiken voor taxonomische doeleinden,, als ook om de verwantschapsrelaties tussen de soorten te leren kennen. Voor de interpretatie vann de bloeiwijzen zijn de concepten van de school van Troll (1964) gevolgd
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