The Leaf Anatomy of Apocynaceae in East Africa - a Monograph of Pleiocarpinae
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ro c no o £>\ A CONTRIBUTION TO THE LEAF ANATOMY AND TAXONOMY OF APOCYNACEAE IN AFRICA - The leaf anatomy of Apocynaceae in East Africa - A monograph of Pleiocarpinae/ Elizabeth Omino Propositions (Stellingen) 1. We come to know our own minds only by explaining ourselves to others. Until we have to defend our opinions in public, they remain half-formed convictions based on random impressions. Jean Bethke Elshtain, Democracy on Trial. 2. Apocynaceae leaves exhibit a wide range of variability in anatomical characters, some of which can be confidently used to distinguish the genera and the species. This thesis. 3. The genus Pleiocarpa has proved to be extremely variable as a whole, and one cannot merely unravel this mystery based on morphology alone. This thesis. 4. Pairs of related genera occurring in different continents will often prove to be congeneric. F.J. Breteler, The boundary between Amherstieae and Detarieae (Caesalpinioideae). In: Adv. in Legume Syst. 7: 53-61, 1995. 5. Although the species may be defined as a group of similar organisms that interbreed or share a common lineage of descent, there is no universal agreement on how to define a species. V.H. Heywood. In: Global Biodiversity assessment, UNEP 1995. 6. The measure of a truly great man is the courtesy with which he treats a little man. 7. After all the outcry on human rights and equality for women leading to the Beijing conference, the British High Commission cannot, surely, in this time and age equate the Kenyan women with children. Recent requirement for all Kenyan women travelling to Britain to obtain written permission from their husbands when applying for a visa. 8. No matter how carefully we may strive to classify various entities of life, there will always be the odd man out. 9. Nothing is as easy as it looks, everything takes longer than you expect. And if anything can go wrong it will, at the worst possible moment. Adapted from Murphy's Law. Propositions belonging to the thesis entitled "A contribution to the leaf anatomy and taxonomy of Apocynaceae in Africa" by E. Omino. Wageningen, April 22, 1996. A contribution to the leaf anatomy and taxonomy of Apocynaceae in Africa The leaf anatomy of Apocynaceae in East Africa A monograph of Pleiocarpinae Promotor: Dr.Ir.L .J .G .va n derMaesen , hoogleraar in de plantentaxonomie Co-promotor:Dr.A.J.M.Leeuwenberg, universitair hoofd-docent, vakgroep Plantentaxonomie A contribution to the leaf anatomy and taxonomy of Apocynaceae in Africa - The leaf anatomy of Apocynaceae in East Africa - A monograph of Pleiocarpinae (Series of revisions of Apocynaceae XLI) Elizabeth Omino PROEFSCHRIFT ter verkrijging van de graad van doctor in de landbouw- en milieuwetenschappen opgeza g vand erecto rmagnifies , DrC.M.Karssen , inhe topenbaa rt e verdedigen op maandag 22 april 1996de s namiddags te vier uuri n de Aula van de Landbouwuniversiteit teWageninge n CIP-gegevens Koninklijke Bibliotheek, Den Haag Omino, Elizabeth A contribution to the leaf anatomy and taxonomy of Apocynaceae in Africa : The leaf anatomy of Apocynaceae in East Africa :A monograp h of Pleiocarpinae (Series of revisions of Apocynaceae XLI)/ Elizabeth Omino. -Leide n : Backhuys Publishers. - 111. Thesis Landbouwuniversiteit Wageningen. -Wit h index, ref. - With summary in Dutch. ISBN 90-73348-54-4 NUGI 835 Subject headings: leaf anatomy / Apocynaceae ;taxonomy . TABLE OF CONTENTS General Summary i Samenvatting . .. 2 i Leaf anatomy of Apocynaceae in East Africa 3 1.1 Summary 3 1.2 Introduction 3 1.3 Material and Methods 4 1.4 Observations 4 1.5 Characteristics of the genera 10 5.1 Acokanthera G.Don 10 5.2 Adenium R.B r 11 5.3 Alafia Thouars 12 5.4 Ancylobotrys Pierre 13 5.5 Baissea A.D C 14 5.6 Carissa L 15 5.7 Carvalhoa K.Schu m 16 5.8 Clitandra Benth 17 5.9 Dictyophleba Pierre 18 5.10 DiplorhynchusWdw. ex Fie. & Hiern 18 5.11 Funtumia Stapf 19 5.12 Holarrhena R.B r 20 5.13 Hunteria Roxb 21 5.14 Landolphia P.Beau v 22 5.15 Mascarenhasia A.D C 23 5.16 Motandra A.D C 24 5.17 Oncinotis Benth 25 5.18 Picralima Pierre 26 5.19 Pleiocarpa Benth 26 5.20 Rauvolfia L 28 5.21 Saba (Pichon) Pichon 29 5.22 Schizozygia Baill 30 5.23 Stephanostema K.Schu m 30 5.24 Strophanthus DC 31 5.25 Tabernaemontana L 32 5.26 VoacangaThouars 33 1.6 General discussion 34 1.7 Conclusions 37 1.8 Figures and tables 42 1.9 Anatomy references 76 2 A monograph of Pleiocarpinae (Apocynaceae-Plumerioideae-Carisseae); Series of revisions of Apocynaceae XLI 81 2.1 Summary 8i 2.2 Introduction 81 2.3 General part 82 2.3.1 History of the subtribe Pleiocarpinae 82 2.3.2 History of the genera Hunteria Roxb.,Pleiocarpa Benth . and PicralimaPierr e 82 2.3.3 Geography and distribution 83 2.3.4 Morphology 83 2.3.5 General taxonomic references 86 2.4 Taxonomic part 87 2.4.1 The subtribe Pleiocarpinae 87 2.4.2 Key to the genera of the subtribe Pleiocarpinae 87 2.5 HunteriaRox b 88 2.5.1 Hunteriaballayi Hu a 90 2.5.2 Hunteriacamerunensis K .Schum . ex Hall 03 2.5.3 Hunteriacon golana Pichon 96 2.5.4 Hunteriadensiflora Pichon 99 2.5.5 Hunteriaghanensis Hall &Leeuwenber g 102 2.5.6 Hunteriahexaloba (Pichon ) Ominocomb , nov 104 2.5.7 Hunteriamacrosiphon Omin o sp.no v 107 2.5.8 Hunteriamyriantha Omin o sp.no v 109 2.5.9 Hunteriaoxyantha Omin o sp.no v 112 2.5.10 Hunteriasimii (Stapf) H.Huber 115 2.5.11 Hunteria umbellata(K .Schum. ) Hall 118 2.5.12 Hunteriazeylanica (Retz. ) Gard.exTh w 122 2.6 PicralimaPierr e 128 2.6.1 Picralimanitida (Stapf)Th. &H.Dur . 128 2.7 Pleiocarpa Benth 134 2.7.1 Pleiocarpabicarpellata Stapf 135 2.7.2 Pleiocarpabrevistyla Omin o sp.no v 139 2.7.3 Pleiocarpamutica Bent h 142 2.7.4 Pleiocarpapycnantha (K.Schum. ) Stapf 146 2.7.5 Pleiocarpa rostrata Benth 152 2.8 Doubtful species 156 2.9 Excluded species 157 2.10Inde x of exsiccatae 158 Index of scientific names 169 Curriculum Vitae 177 Acknowledgements 177 VI GENERAL SUMMARY This publication comprises two main parts. Part one(Chapte r i) dealswit h thelea f anatomy of Apocynaceae inEas t Africa, wherethi sfamil y isrepresente d by7 7specie s in3 0genera .Ou to fthese ,3 7specie s in 26gener a areexamine d and these exhibit awid erang e of anatomical variations. Many species can be separated on thebasi s of their leaf anatomy and many genera are confirmed to be closely related.Th e most outstanding anatomical characters includeth edorsiventra l leaves,presenc eo f laticifers, andbicollatera l vascularbun dles with intraxylary phloem. Parttw o(Chapte r 2)i sa monograp h ofth eapocynaceou s subtribe Pleiocarpinae- Plumeriodeae-Carisseae, which comprises of three genera, Hunteria (12 spp.); Pleiocarpa( 5spp. )an dPicralima (1sp.).Th e subtribePleiocarpina e isrestricte d to Africa, althoughon especies ,Hunteria zeylanica, extendst oAsia .I ti seasil y distin guished fromth eothe rmember so fth etrib eCarissea eb yth epresenc eo fcompletel y apocarpous ovaries, with 2-5 carpels, which are a great exception in the family Apocynaceae. Thegenu sPicralima wa sfirst describe db yPierr ei n 1896an dha ssinc eremaine d monotypic.It sonl y species,Picralima nitida, i sno tonl y exploited for itswood ,bu t also used medicinally in its present distribution area and employed as well as an arrow and afish poison . In this publication, three new species are described in the genus Hunteria (Roxburgh, 1814, 1824)an don e new combination is made. The genus Pleiocarpa (Bentham, 1876)ha s proved to be aver y difficult subject due to insufficient collections and lack of sufficient information from thefield. A t present it has 5 species and one doubtful species of which the collections are all either sterile oronl y bearing fruits. One new species is described. Parallel to the investigations reported here a taxonomic treatment was prepared for the Flora of Tropical East Africa.Thi s contribution, covering all the mentioned 30gener a and 77 species, will be submitted to the editors in Kew shortly. WageningenAgricultural UniversityPapers 96-1 (1996) SAMENVATTING Deze publicatiebestaa t voomamelijk uit twee delen. Hoofdstuk i behandelt de bladanatomie van vertegenwoordigers van de in Oost Afrika voorkomendeApocynaceae ,ee nfamili e diedaa r7 7soorte ni n3 0gener atelt . Hiervan werden 37 soorten van 36 genera bestudeerd en deze vertonen een grote variatieaa nanatomisch ekenmerkstaten .Vee lsoorte nkunne nworde nonderscheide n aand ehan dva nd ekenmerke n vand ebladanatomie ,e nd enauw e verwantschappen tussen vele genera kunnen er mee worden bevestigd. De meest opvallende anatomische kenmerken zijn de dorsiventrale bladeren, de aanwezigheid van melksapvaten, en bicollaterale vaatbundels met intraxylairfloeem. Hoofdstuk 2 is een monografie van de subtribus Pleiocarpinae, welke uit drie genera bestaat: Hunteria(me t 12soorten) ;Pleiocarpa ( 5 soorten) en Picralima( 1 soort). De subtribus Pleiocarpinae van de onderfamilie Plumerioideae, tribus Carisseae, is voomamelijk beperkt tot Afrika, maar een soort, Hunteriazeylanica, komt 00k in Azie voor. De subtribus verschilt duidelijk van andere in de tribus Carisseae door de aanwezigheid van geheel apocarpe vrachtbeginsels met 2 tot 5 vruchtbladen, hetgeen een grote uitzondering is in de familie Apocynaceae. Het genus Picralima werd door Pierre in 1896 beschreven en is sindsdien monotypisch gebleven.D eenig esoort ,Picralima nitida, wordtnie tallee n vanwege het houtgeexploiteerd , maar 00k medicinaal benute nbovendie n zowelvoo r pijlgif als voor visvergift. InHunteria (Roxburgh , 1814,1824) worden drienieuw e soorten beschreven, en er is een nieuwe combinatie gemaakt.