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POLYGALARUM AFRICANARUM ET MADAGASCARIENSIUM PRODROMUS ATQUE GERONTOGÆI GENERIS HETEROSAMARA Kuntze, a GENERE POLYGALA L. SEGRE

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POLYGALARUM AFRICANARUM ET MADAGASCARIENSIUM PRODROMUS ATQUE GERONTOGÆI GENERIS HETEROSAMARA Kuntze, a GENERE POLYGALA L. SEGRE PAIV_CUB.QXD 02/04/2008 17:50 PÆgina 1 POLYGALARUM AFRICANARUM ET MADAGASCARIENSIUM PRODROMUS ATQUE GERONTOGÆI GENERIS HETEROSAMARA Kuntze, A GENERE POLYGALA L. SEGREGATI ET A NOBIS DENUO RECEPTI, SYNOPSIS MONOGRAPHICA JORGE AMÉRICO RODRIGUES PAIVA FONTQUERIA L MADRID 1998 PAIV_CUB.QXD 02/04/2008 17:50 PÆgina 2 FONTQUERIA is a series of botanical publications without administrative binding. It publishes original works in botany, especially those that are of interest to the editors. Its publications are in any language, the only limitation being the capacity of the team of editors. Accredited with the International Asociation for Plant Taxonomy for the purpose of registration of new non-fungal plant names. PRODUCTION AND DISTRIBUTION: Cyanus, S. L. Camino de los Vinateros, 87-1ºC E-28030 Madrid Database consultant: Guillermo GONZÁLEZ GARCÍA Composition: Ulpiano SOUTO MANDELOS & Ambrosio VALTAJEROS POBAR Screen operator: Samuel FARENA SUBENULLS, Emilio NESTARES SANTAINÉS Preimpression: Sonja MALDÍ RESTREPO, Demetrio ONCALA VILLARRASO Editor F. Javier FERNÁNDEZ CASAS Real Jardín Botánico. CSIC. E-28014 Madrid Joint editors André CHARPIN (French texts) Conservatoire botanique, Genève. Suiza Manfred DITTRICH (German texts) Conservatoire botanique, Genève. Suiza Cirilo H. NELSON SUTHERLAND (Latin texts) Universidad Nacional Autónoma de Honduras. Tegucigalpa Antonio PRETEL MARTÍNEZ (Russian texts) Unidad de Genética. Facultad de Ciencias. Universidad de Málaga Antonio M. REGUEIRO Y GONZÁLEZ-BARROS (English texts) Virgen del Pilar, 9. E-28230 Las Rozas, Madrid Editing advisors Consuelo CEBOLLA LOZANO. Facultad de Biología, Universidad Antónoma. E-28049 Madrid José María MONTSERRAT MARTÍ. Institut Botànic de Barcelona María Antonia RIVAS PONCE. Facultad de Biología, Universidad Antónoma. E-28049 Madrid Alfonso SUSANNA DE LA SERNA. Institut Botànic de Barcelona Madrid, June 1998 ISSN: 0212-0623 Depósito legal: M-29282-1982 Volume XLVIII was distributed on 15-I-1998 Volume XLIX was distributed on 17-XII-1977 PAIV_000.QXD 02/04/2008 16:53 PÆgina i POLYGALARUM AFRICANARUM ET MADAGASCARIENSIUM PRODROMUS ATQUE GERONTOGÆI GENERIS HETEROSAMARA Kuntze, A GENERE POLYGALA L. SEGREGATI ET A NOBIS DENUO RECEPTI, SYNOPSIS MONOGRAPHICA JORJE AMÉRICO RODRIGUES PAIVA REAL JARDÍN BOTÁNICO Madrid, 1998 PAIV_000.QXD 02/04/2008 16:53 PÆgina ii PAIVA, J. A. R. (1998). Polygalarum africanarum et madagascariensium prodromus atque geron- togæi generis Heterosamara Kuntze, a genere Polygala L. segregati et a nobis denuo recepti, synopsis monographica. Fontqueria 50: vi+346. Keywords: Polygala L., Heterosamara Kunze (Polygalaceæ), Anatomy, Cladistics, Morphology, Palinology, Phytochemistry, Phytogeography, Taxonomy, Africa, Asia, Madagascar. Résumé. Révision des espèces africaines et malgaches du genre Polygala L. (Polygalaceæ) et synopsis du genre Heterosamara Kunze, consideré comme distinct du précédent et qui se ren- contre en Afrique et en Asie. Histoire résumée de la famille et du genre. Caractères généraux, étude des caractères morfolo- giques et anatomiques des différents organes, aspects ultrastructuraux des trichomes. Étude détaillée des graines et de leur adaptation à la myrmécochorie et l´ornitochorie, de la diversité bio- logique et écologique. Le genre Heterosamara, caracterisé par son pollen hétéropolaire, est réha- bilité. Le genre Polygala est restreint aux espèces à grains de pollen isopolaires et polyzonocol- porés. Les aspects phytogéographiques et la phylogénie de la famille, spécialement du genre Polygala, sont discutés. Une analyse cladistique est également présentée. En complément de la partie descriptive, sont ajoutés une carte, huit tableaux, quatre phénogrammes, trois cladogram- mes et 52 planches. Une liste des nomina excludenda et des nomina rejicienda vel dubia est éga- lement établie, ainsi qu´une liste des références bibliographiques mentionées dans le texte. —. ˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛ ˛˛˛˛ Polygala L. (Polygalaceæ); ˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛ ˛˛ ˛˛˛˛ ˛˛˛˛ Heterosamara Kuntze ˛ ˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛ ˛ ˛˛˛˛. ˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛. ˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛ ˛˛˛˛ ˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛. ˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛ ˛˛˛˛˛, ˛˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛; ˛˛˛˛˛˛˛˛ ˛˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛˛. ˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛. —˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛ Heterosamara, ˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛. ˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛ ˛˛˛a˛˛˛˛ ˛ Polygala. ˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛ ˛˛˛˛˛˛˛˛˛, ˛ ˛˛˛˛˛˛˛˛˛˛˛ ˛. Polygala. ˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛. ˛ ˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛ ˛ ˛˛˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛, ˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛ ˛˛˛˛˛, ˛˛˛˛˛˛ ˛˛˛˛˛˛, ˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛, ˛˛˛ ˛˛˛˛˛˛˛˛˛˛˛ ˛ 52 ˛˛˛˛˛˛˛. ˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛ ˛˛˛˛˛˛˛˛ nomi- na excludenda ˛ nomina rejicenda vel dubia ˛ ˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛ ˛˛˛˛˛˛˛˛˛˛. Zusammenfassung. Vom Autor wird eine Revision der afrikanischen und madagassischen Arten der Gattung Polygala L. (Polygalaceæ), sowie eine Synopsis der afrikanischen und asiatischen, von Polygala abgetrennten Gattung Heterosamara Kuntze vorgelegt. Nach einem historischen Überblick über Familie und Gattung werden die wichtigsten morpho- logischen und anatomischen Merkmale, sowie die Ultrastruktur ihrer besonderen Behaarung dar- gestellt. Eine spezielle Studie ist der biotypischen und ökologischen Variabilität der Samen und ihre Anpassung an die Verbreitung durch Ameisen und Vögel gewidmet. In einer palynologischen Studie werden morphologische und ultrastrukturelle Merkmale ermittelt. Dabei hat sich gezeigt, daß bei Polygala isopolare und polyzonokolporate und bei Heterosamara nur heteropolare Pollenkörner vorkommen. Die Verbreitung und Phylogenie der Familie und der Gattung Polygala werden besprochen; die Gattung wird auch kladistisch bearbeit. Die vorliegende Arbeit wird durch 1 Karte, 8 Tabellen, 3 Kladogramme, 4 Phänogramme und 52 Abbildungen ergänzt; eine Liste mit den Nomina excludenda und Nomina rejicienda vel dubia, sowie ein Literaturverzeichnis schließen die Arbeit ab. Summary. A revision of the African and Malagassy species of the genus Polygala L. (Polygalaceæ), and a synopsis of the genus Heterosamara Kuntze, segregated from the former one and extending throughout Africa and Asia. A historical summary of the family and the genus. General characterization and study of the morphologic and anatomic characters of all its organs, with ultrastructural aspects of its charac- teristic tricomes. A special study of the seed and their adaptation to mirmecochory of ornitho- chory, and of the biotipic and ecological variability. A special study of the morphologic and ultras- PAIV_000.QXD 02/04/2008 16:53 PÆgina iii J. A. R. PAIVA - POLYGALARUM AFRICANARUM ET MADAGASCARIENSIUM... III tructural characters of the pollen grain. The genus Heterosamara, with heteropolar pollen, is reha- bilitated, keeping in Polygala the species with isopolar and polyzonocolporate grains. The phy- togeographic aspects ant the philogeny of the family are discussed, particularly of , with an atta- ched cladistic analysis. As a supplement to the descriptive part, 1 map, 8 tables, 4 phenograms, 3 cladograms and 52 plates are added. A list of nomina excludenda and nomina rejicenda vel dubia are also included, together with the list of bibliographic references mentioned in the text. Resumen. Revisión de las especies africanas y malgaches del género Polygala L. (Polygalaceæ), más sinopsis del género Heterosamara Kuntze, segregado de aquél y que se extiende por África y Asia. Resumen histórico de la familia y del género. Caracterización general y estudio de los caracte- res morfológicos y anatómicos de todos sus órganos, más aspectos ultrastructurales de sus carac- terísticos tricomas. Estudio especial de las semillas y de sus adaptaciones a la mirmecocoria u ornitocoria, de la diversidad biotípica y ecológica. Estudio especial de los caracteres morfológi- cos y ultrastructurales del grano de polen. Se rehabilita el género Heterosamara, de polen hete- ropolar, manteniendo en Polyga las especies con granos isopolares y polizonocolporados. Se dis- cuten los aspectos fitogeográficos y la filogenia de la familia, particularmente de Polygala, acom- pañada de análisis cladístico. Como complemento de la parte descriptiva, se añade un mapa, ocho cuadros, cuatro fenogramas, tres cladogramas y 52 láminas. Se acompaña también una lista de los nomina excludenda y nomina rejicienda vel dubia, más la lista de referencias bibliográficas men- cionadas en el texto. INDÍCULO Ficha bibliográfica y resúmenes ............................................................................................. [ii] Indículo..................................................................................................................................... iii Agradecimientos .................................................................................................................... vi Prólogo ...................................................................................................................................... 1 INTRODUCCIÓN ....................................................................................................................1 MATERIAL Y MÉTODOS ...................................................................................................... 8 MATERIAL ................................................................................................................................8 MÉTODOS ................................................................................................................................
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