A Revision of the Genus Allium L. (Liliaceae) in Africa

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A Revision of the Genus Allium L. (Liliaceae) in Africa A REVISION OF THE GENUS ALLIUM L. (LILIACEAE) IN AFRICA CENTRALE LANDBOUWCATALOGUS 0000 0086 9517 Dit proefschrift met stellingen van BRIGITTA EMMA ELISABETH DE WILDE-DUYFJES biologisch doctorandus, geboren te Bandung, Indonesia, op 29jun i 1936,i sgoedgekeur d door depromoto r Dr. H. C. D. de Wit, hoogleraar ind ealgemen eplantensystematie k en -geografie eni nhe t bijzonder die van de tropen end e subtropen. De Rector Magnificus vande Landbouwhogeschool, J. P. H. VAN DER WANT Wageningen,1 8novembe r 1976 Viri^zot 6^3 582.572.225(6) BRIGITTA E. E. DE WILDE-DUYFJES A REVISION OF THE GENUS ALLIUM L. (LILIACEAE) IN AFRICA PROEFSCHRIFT TER VERKRIJGING VAN DE GRAAD VAN DOCTOR IN DE LANDBOUWWETENSCHAPPEN OP GEZAG VAN DE RECTOR MAGNIFICUS, DR. IR. J. P. H. VAN DER WANT, HOOGLERAAR IN DE VIROLOGIE, IN HET OPENBAAR TE VERDEDIGEN OP WOENSDAG 26 JANUARI 1977 DES NAMIDDAGS TE VIER UUR IN DE AULA VAN DE LANDBOUWHOGESCHOOL TE WAGENINGEN Date of publication 22 December 1976 H. VEENMAN & ZONEN B.V. - WAGENINGEN - 1976 Thisthesi si sals opublishe d asMededelinge n Landbouwhogeschool Wageningen76-1 1(1976 ) (Communications Agricultural University, Wageningen, The Netherlands) /f 92°f £73 STELLINGEN I De Linneaanse naam Allium ascalonicum moet geaccepteerd worden; de Linneaanse naam Alliummagicum moet verworpen worden. LINNAEUS, C. [Strand]. 1756. Flora Palaestina. LINNAEUS, C. 1759.Syst . Nat. LINNAEUS, C. 1762.Spec . Plant, ed.2 . STEARN, W. T. 1960. Bull. Brit. Mus. (Nat. Hist.) Vol. 2,no .6 : p. 182. Int. Code Bot. Norn. 1972,Art . 69, 70. II Bij hetaanwijze n vanhe ttyp e vanLinneaans e soortnamen waarvan geen authentiekedoo r Linnaeus geziene exemplaren aanwezig zijn, kan een bij een literatuur-citaat uit de protoloog behorend herbariumexemplaar als lecto- type aangewezen worden. DEWILDE-DUYFJES , B.E .E . 1973. Taxon 22(1):p . 57-91. Ill Eendefinitiev e indelingi nsubgener aen/o f sectiesva nhe tgenu sAllium moet wachten totdi tgenu s als geheel ina l zijn aspecten evenwichtig bestudeerdis ; hieraan lijkt ookd ebou w vand ehoningkliere n ind esept a vanhe t vrucht- beginsel eenwaardevoll e bijdrage te kunnen leveren. IV Bijhe taanneme n van gegevensomtren td eouderdo m van taxa uit fossielen, en hettrekke n vanconclusie s daaruit, wordt niet zelden derege l veronacht- zaamd dat uit fossielen slechts een minimum ouderdom kan worden vast- gesteld. Voorts verdient het persen vanbotanisch e gegevens in het keurslijf van geofysische hypothesen geen aanbeveling. RAVEN, P. H. & T. E. RAVEN. 1976. The genus Epilobium in Australasia. New Zealand D.S.I.R. Bull. 216:i n het bijzon- der p. 68-73. SCHUSTER, R. 1976. Plate tectonics andit sbearin g onth egeo ­ graphical origin anddispersa l ofAngiosperms . In: Originan d Early Evolution of Angiosperms. ed. C. B. Back. Columbia Press: p.48-138 , 45fig., i nhe tbijzonde r p. 53, 54,58 , 69, 76, 119, 121. V Bij taxonomische studies voor hetMiddellandse-Zeegebie d moeten onder- zoekersdi ewerke nvoo r Europa rekeninghoude n metd emogelijkhei d dat ook soorten metee nklei n areaal eveneens kunnen voorkomen in Afrika, envic e versa; dit geldt vooral voor dezuidelijk e Griekse eilanden end e Cyrenaica, envoo rZuid-Spanj e en hete rtegenove rliggend egedeelt eva n Noord-Afrika. SANDWITH, N. Y. & N. D. SIMPSON. 1941. Additions to the flora of the Cyrenaica. Journ. Bot. Vol. LXXIX: p. 33-44, in het bijzonder p. 34. RECHINGER, K. H. 1950.Grundziigede r Pflanzenverbreitung in derAegai sI .Vegetati o Vol.II : p.55-120 . GREUTER, W. 1971.Betrachtunge n zur Pflanzengeographie der Sudaegais(ed . Arne Strid). Opera Bot. No. 30:p . 22-24. VI Door Love& Lov eword td e cytolooggetypeerd ,nie td e taxonoom. LOVE, A. & D. LOVE. 1974.Cytotaxonomica l Atlas of the Slovenian Flora (in het bijzonder p.XI) . VII Het isvoorbari g om opgron d van verschillen ind evroeg e embryonale ontwikkeling vand eAnura e n de Urodela tebesluite n tot een diphyletische oorsprong van derecent eAmphibia. NIEUWKOOP, P. D. &LIE N A. SUTASURYA. 1976. Embryological evidencefo ra possibl epolyphyleti c origin ofth erecen t amphib­ ians.J . Embryol.exp .Morph . Vol.35,1 : p. 159-167. VIII Bij hetintroducere n vanchemisch e bestrijdingsmiddelen in bepaalde niet- Westerse gebieden, geeft men zich dikwijls onvoldoende rekenschap van het feit dat, als gevolg vansocio-culturel e omstandigheden, bedreigende vormen vanmisbrui k voord egezondhei d vanmen se nmilie uvoo rd ehan d liggen. IX Het iszee r waarschijnlijk dat Orang Utans voor hun voortbestaan grote arealen primair, niet uitgekapt, (laagland-)regenbos nodig hebben; hetzelfde geldt, mutatis mutandis, voor Rafflesia en vele andere plante-e ndiersoorten . Tegen: WILSON, C. C. & W. L. WILSON. 1975. The influence of selective logging on Primates and some other animals inEas t Kalimantan. Folia Primatol. 23: 245-274. X Omd e recreatieve ennatuurwetenschappelijk e functie van het Groene Hart van Holland tebehouden , moet o.a. het jaarlijks toegestane accres van wo- ningen ind ealdaa r gelegen dorpen, drastisch verminderd worden. XI In weerwil van de mening van Steam, was Rabelais bekend met de narco- tiserende werking van hennep. STEARN, W. T. 1974. Typification ofCannabi s sativa L. Bot. Mus. Leafl. Harvard Un.Vol .23 ,No .9 :p .325-336 . XII Biologen zijn de ergsten. (Uitspraak van verscheidene boswachters). A revision of the genus Allium L.(Liliaceae) in Africa Proefschrift BRIGITTA E. E. DE WILDE-DUYFJES Wageningen, 26januar i 1977. Aan de nagedachtenis van mijn Vader Aan mijn Moeder CONTENTS Page INTRODUCTION 1 SYSTEMATIC POSITION OF ALLIUM 2 THE CHARACTERS IN ALLIUM 3 Habit and growth 3 Bulb and leaves 3 Outer bulbcoat-leaves 4 Increase bulbs 4 Leaf blades 4 Scape 6 Inflorescence 6 Spathe 6 Flowers 6 Stamens 7 Pistil 7 Nectaries 8 Fruits 8 Seeds 8 SEEDLINGS 9 ANATOMY 11 PALYNOLOGY 13 CARYOLOGY 14 DISTRIBUTION 15 ECOLOGY AND FLOWER BIOLOGY 16 USES AND PHYTOCHEMISTRY 17 TYPIFICATION 18 ADDITIONAL REMARKS ON THE PRESENTATION OF DATA 19 SUBDIVISION INTO SECTIONS 20 DESCRIPTION OF THE GENUS ALLIUM 23 KEY TO THE SECTIONS AND SPECIES 25 DESCRIPTIONS OF SECTIONS AND SPECIES 29 NAMES EXCLUDED FOR AFRICA 212 GENERAL BIBLIOGRAPHY 213 SUMMARY (English) 222 (Dutch) 223 ACKNOWLEDGEMENTS 225 INDEX OF NAMES 226 ALPHABETICAL INDEX OF ACCEPTED SECTION- AND SPECIES NAMES Page Section Allium 29 Briseis 113 Codonoprasum 91 Melanocrommyum 188 Molium 120 Schoenoprasum 83 Allium ampeloprasum 63 ascalonicum 54 baeticum 78 blomfleldianum 185 cepa 84 chamaemoly 153 crameri 208 cupani 109 dregeanum 75 erdelii 121 fistulosum 87 guttatum 34 longanum 143 mareoticum 41 massaessylum 165 neapolitanum 158 nigrum 193 ssp. multibulbosum 195 ssp. nigrum 196 orientate 200 paniculatum 95 papillare 147 roseum 169 ruhmerianum 182 sativum 59 var. ophioscorodon 62 var. sativum 60 schoenoprasum 89 schubertii 188 sphaerocephalum 44 ssp. curtum 52 ssp. durandoi 50 ssp. sphaerocephalum 46 subhirsutum 126 ssp. spathaceum 142 ssp. subhirsutum 131 ssp. subvillosum 137 tourneuxii 150 trichocnemis 92 triquetrum 113 vineale 29 INTRODUCTION Being well acquainted with theLaborator y for Plant Taxonomy and Plant Geography of the Agricultural University at Wageningen after nearly 4 years of botanical activities in tropical Africa, andafte r having received the degree of M.Sc. at the University ofLeyde n in 1966,1 was glad tostart , atth e instiga­ tion ofProf .dr .H .C .D . DE WIT, arevisio n ofth e genus Allium forAfrica .Th e rich assortment of Allium taxa grown in the botanical garden had drawn my attention andle dtoward s this decision. During the several years ofth e present study on Allium still more specimens from Europe and North Africa were taken into cultivation, which, in addition to the rich herbarium collections examined in the major European and North African herbaria, proved very helpful fora goo d understanding of the species. Allium inAfric a islargel yconfine d to the Mediterranean region which implied that the North African taxa could not be satisfactorily delimited and named without an accompanying study of the taxa occurring in the northern and eastern parts ofth e Mediterranean vegetation. All more recent treatments of North African Allium were inloca l floras, e.g. PAMPANINI (1931) for the Cyre- naica, CUENOD (1954) for Tunisia, QUEZEL & SANTA (1962) for Algeria, and TACKHOLM & DRAR (1954) forEgypt , apart from the several precursory papers by MAIREc.s. ,resultin g in MAIRE'Streatmen t ofth e genusfo r thewhol e of North Africa in the 'Flore d'Afrique du Nord' (1958). Besides these works, several recent papers by FEINBRUN and KOLLMANN for theNea r East, mainly Israel, and by VON BOTHMER and EKBERG for theAegea n area were of great impor­ tance. The above mentioned earlier botanists dealing with African Allium hadpai d but little attention to the taxa described from Europe and the students of European or eastern Mediterranean Allium hardly took notice of those from Africa. The present study, including the examination of a great part ofth e European-Mediterranean material, forced me to accept a rather broad species concept, andtherefor e brought about several name changes andth e reduction into synonymy of many names ascompare d to MAIRE'S treatment. Nomenclature, first ofall , demanded that the Linnean binomials for Allium speciespossibl y found inAfric a betypified , which led toa previou s publication by me on the subject: 'Typification of 23Allium species described by LINNAEUS and possibly occurring in Africa', (Primitiae Africanae 9), in Taxon 22(1), 1973, p. 57-91. This article forms the first part of my present doctor's thesis. Meded.Landbouwhogeschool Wageningen 76-11 (1976) SYSTEMATIC POSITION OF ALLIUM The genus Allium occupies, together with a few related genera, as there are the South American genus Nothoscordum KUNTH, and many other small South American genera including Ipheion RAFINBSQUE, the SW. Asian genus Nectaroscordum LINDLEY, and possibly the South African genus Tulbaghia L., a rather intermediate position between Liliaceae and Amaryllidaceae.
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