Oleanders Nerium L and The Oleander Cultivars Ser-Wageningen University and Research 290321 PDF

SERIES OF REVISIONS OF APOCYNACEAE
PARTXX
OLEANDERS
NERIUM L.ANDTHEOLEANDER CULTIVARS
F. J.J. PAGEN
Department ofPlant Taxonomy
Agricultural University Wageningen, The Netherlands
Wageningen mm A g r i c u l t u r a l U n i v e r s i t y
BIBLIOTHEEK
LANDBOUWUNIVERSITEIT,
WAGEiNINGEN
i LANDBOUWCATALOQUS
00000307 7829 \^n 2.02,^3

Cip-data Koninklijke Bibliotheek, Den Haag
ISBN906754 1079
© Agricultural University Wageningen, TheNetherlands, 1988.
No part of this publication, apart from abstract, bibliographic and brief quota-
tions embodied in critical reviews, may be reproduced, re-corded or published
in any form including print, photocopy, microform, elektronic or elektromag-
neticrecord without written permission from thepublisher Agricultural Univer-
sity,P.O.Box 9101, 6700HBWageningen, the Netherlands.
Printed intheNetherlands by Drukkery Veenman B.V., Wageningen

CONTENTS
INTRODUCTION 3
PART ONE:NERIUM OLEANDER 5

Nerium L 5
Nerium oleander L 5
Description 7
Geographical distribution 10
Ecology 10
Flowering and fruiting periods 12
Vernacular names 12
Specimensexamined 14
Nomen nudum 18
Excluded species 18
Pollination of theoleander 19
Locating the stigmatic area on the pistil in Nerium oleander L. by
means offluorescence microscopy 19
Pollinating mechanism 22
Apoisonous plant 24
Medicinal uses 25
Other uses 25
PART TWO: PHYTOCHEMISTRY OFNERIUM L. (N. G. BISSET) 27

Cardiacglycosides 28
Other constituents 33
Conclusion 36
References 37
PART THREE: THE OLEANDER CULTIVARS 39

Tentative Checklist ofOleanderCultivars 39
History ofthecultivated oleander 43
The oleander incultivation today 46
ACKNOWLEDGEMENTS 49
REFERENCES 51
APPENDIX 55
Agric. Univ. WageningenPapers87-2 (1987)

INTRODUCTION
The present paper, devoted to the genus Nerium L. covers taxonomie and
other aspectsofthewild oleander andoftheoleander cultivars.
The first section deals with thewild oleander andincludes a revision ofthe
genus.
A precursor of the revision of Nerium L. waspublished by LEEUWENBERG
in 1984andhasbeen completed bythepresent author under LEEUWENBERG's
supervision. Thestudy is based on several hundreds of herbarium specimens,
most ofthem wellprepared andrichly provided with flowers and/or fruits.The
genus Nerium L.isconsidered tobemonotypic, itssingle species being Nerium
oleanderL.
Adistribution map ofthenaturally occurring Nerium oleanderL.isincluded.
The flower ofNerium oleanderL.features apistil with ahighly differentiated
pistil head. An experiment executed to pinpoint the location of the stigmatic
area isdescribed andthepollination mechanism isdiscussed.
Oleanders areknown aspoisonous andmedicinal plants. BISSETdiscussesthe
phytochemistry of Nerium L.in thesecond section.
The third section dealswith thecultivated oleander.
An inventory ofthe oleander cultivars hasbeen compiled bythepresent au-
thor, resultingintheTentative Checklist ofOleander Cultivars.
Oleander cultivars varyinform, colour andscentoftheflowers, andinhabit.
Severalcultivars havevariegated leaves.
Oleanders arevery popular ornamental plants and commonly cultivated in
tropical and subtropical countries as a garden shrub and elsewhere as a pot
plant.The history ofthe oleander incultivation isdiscussed.
The phytochemistry chapter was written by N. G. BISSET; F. J. J. PAGEN au-
thored theother chapters.
Agric. Univ.WageningenPapers87-2 (1987)

PART ONE
N E R I U M OLEANDER
N E R I U M L.
The genus was established by TOURNEFORT in 1700 as 'Nerion' and was

adopted by LINNAEUS,whoin 1737named thegenus 'Nerium'.
The name 'Nerium' isderived from the Greek 'nerion', used by DIOSCORIDES
toindicate theoleander. Thisname supposedly refers totheGreek sea-god Ner-
eus and his fifty daughters, the Nereides. The ancient Greek maintained holy
forests, exclusively planted with oleanders, and garnished altars to honour the
Nereides,whowereconsidered to beinfallible guides (DRAPIEZ 1835).It is often
suggested also,that 'nerion' isderived from the Greek 'neros',meaning 'moist',
alludingto thewet placeswheretheoleander grows wild.
The genusNerium contains only onespecies,Nerium oleanderL.,and belongs
to the tribe Nerieae of the subfamily Apocynoideae (Echitoideae) of the family
Apocynaceae.
The most closely related genusisAdenium (PLAIZIER 1980).Inflorescence and
flower (corolla,corona, pistil,and especiallythestamenswithanthers extending
into long hairy spirally twisted appendages) of Adenium and Nerium are quite
similar.Adenium differs from Neriummainlyinhavingsucculentstems,alternate
leavesconfined totheapicesofthebranchlets,colletersonthebranchlets, slight-
lyzygomorphic flowers, corolla lobes shorter than the corolla tube, obcordate
and lessprominent corona lobes,reflexed follicles, and glabrous seedswith tufts
ofhairateach end.
N E R I U M O L E A N D E R L.
The binomial Nerium oleanderwas given by LINNAEUS in his Species Plantar-

um (1:209. 1753).The epithet 'oleander' isderived from the Latin 'olea', mean-
ing 'olive tree' and the Greek 'dendron': 'tree' or 'shrub', as oleander and olive
leaveslook similar.
The oleander with odourless singleflowers, asitisgrowinginthe Mediterran-
ean region, was the only kind known in Europe until about 1683, when VAN
RHEEDE TOTDRAKESTEIN introduced inHolland an oleander from India (aculti-
vated plant), which attracted attention because of the powerful scent of its
flowers. At the same time a double-flowered form of this 'Sweet-scented
Oleander' wasintroduced into theDutch gardens from Ceylon by BEVERNINGK.
Thiswasalso acultivated plant.
Agric. Univ. WageningenPapers87-2 (1987) 5
The Indian oleander, asit grows wild in Iran, Afghanistan, Pakistan, North-
ern India, Nepal and China, differs from the Mediterranean kind not only in
having fragrant flowers: the flowers are larger and borne in much heavier clus-
ters; the flowering period is much longer; the plants are less robust (needing
more warmth and shelter in cultivation); the segments of the corona are longer
and morefinely divided,usuallyirregularly,infour totenfiliformlobes,whereas
the corona lobes of the Mediterranean oleander are much less prominent: the
segments of the corona are divided in three (sometimes four) lobes, the middle
one short and triangular, with one linear lobe on either side (sometimes two
on one side);the appendages of the anthers areprotruding (scarcely protruding
in the Mediterranean kind); the calyx lobes are erect (versus apically recurved
intheMediterranean oleander);finally,theleavesarelonger,narrower and more
widely spaced.
LEEUWENBERG also inspected specimens ofoleanders,collected more recently
intheMiddle East,with smalland fragrant flowers (Leeuwenberg, pers.comm.).
In 1737 the oleander of the Indian kind was taken up by LINNAEUS, in his
Hortus Cliffortianus, as a variety of the common Mediterranean oleander. In
his Species Plantarum (1753) LINNAEUS did no longer consider it as a separate
variety, but united both plants as one and the same species: Nerium oleander
L. In the thirteenth edition of Systema Vegetabilium (1774) he admitted the
Indian oleander tothevarietalstatusand listeditasvariety ßofNerium oleander
L.Thedouble-flowered form waslisted asvariety y.
The Indian oleander was first distinguished as a species by MILLER (1768).
The plant was named Nerium indicum Mill., typified by a specimen in Herb.
Miller: 'Nerium foliis lineari-lanceolatis rigidis acutis,Nerium indicum angusti-
folium, floribus odoratis,simplicibus. H.L.'.
Apart from that, MILLER also distinguished the double-flowered form as a
separate species: Nerium latifolium Mill.. Later authors generally rejected this
viewandconsidered thedouble-flowered plantasavarietyofthe single-flowered
Indian oleander.
Itisworth mentioningthat Nerium oleanderL.istypified byaspecimen (Phot.
1-2) cultivated in the Hortus Cliffortianus, which in fact is an oleander of the
Indian kind: BM-Herb. Cliff, p. 76: 'Nerium indicum, angustifolium floribus
odoratis simplicibus.H.L.' (lectotypedesignated by STEARN IN DAVIS, 1978).
Inaddition tothespeciesmentioned above,severalotherNerium specieshave

been distinguished. However, from comparative studies of several hundreds of
herbarium specimens including the type specimens cited below and many living
plants in gardens, LEEUWENBERG (1984)concludes that they all should beconsi-
dered as belonging to a single species: Nerium oleander L., a conclusion which
makesthegenusNerium monotypic.

DESCRIPTION
LEEUWENBERG'S description of Nerium oleander L.,including synonyms and

typification, isreproduced belowwithhiskind permission.
Nerium oleander L., Sp. PI. 1:209. 1753;Stearn in Davis, Fl. Turkey 6:159.
1978.
Type: cultivated, Netherlands, near Haarlem, deHartecamp = Hortus Clif-
fortianus (BM-Herb. Cliff, p.76,designated byStearn) (Phot. 1-2). Homotypic
synonyms: N.floridum Salisb., Prod. 147.1796.N. lauriforme Lam.,Fl. Fr.2:
299. 1805.
Heterotypic synonyms:N. indicumMill., Gard. Diet.ed.8no.2. 1768.Type:
Herb. Miller, 'Nerium foliis lineari-lanceolatis rigidis acutis, Nerium indicum
angustifolium, floribus odoratissimplicibus.H.L.'(BM,holotype). Homotypic
synonym: N. oleander var. indicum (Mill.) Degener & Greenwell, Fl. Hawai.
Fam. 305(7/25). 1952.
N. latifolium Mill., op.cit.no.3. 1768. Type: Herb. Miller, 'Nerium (latifo-
lium)foliis lanceolatis longioribus flaccidis' (BM,holotype).Homotypic synon-
ym:N. odorumSalisb.,Prod. 147.1796(notofSoland.).
PHOT. 1. The lectotype specimen of Nerium PHOT.2.Labelofthelectotypespecimen of Ner-

oleander L., BM-Herb. Cliff, p. 76, designated ium oleander L., BM-Herb. Cliff, p. 76: "Ner-
byStearn (phot. W.A. Brandenburg). ium indicum, angustifolium floribus odoratis
simpicibus.H.L."(phot.W.A.Brandenburg).

N. odorum Soland. in Aiton, Hort. Kew. ed. 1. 1: 297. 1789; Willd., Sp. PI.
2. 1: 1235. 1797. Type: Belutta-areli Rheed., Malab. 9: 3,t.2. 1689 (lectotype).
Homotypic synonyms: N. odoratum Lam., Enc. 3: 456. 1792. N. verecundum
Salisb.,Prod. 147.1796.
N. carneum Hort, ex Dum.-Cours., Bot. Cult. ed. 2. 3:268. 1811 (published
with thisbinomial asvariety ofN. oleander,typeapparently not preserved).
N.flavescens Di Spino,Jard. de St. Sebastien 1812,ex Roem. &Schult., Syst.
4: 410. 1819. Type not yet traced. Homotypic synonym: N. luteum Nois. ex
Steud.,Nom.ed. 1:553. 1821.
N. splendensHort, exPaxt., Mag. Bot. 3:73. 1837.Type apparently not pre-
served.
N. thyrsiflorum Paxt., loc.cit.,withplate (platedesignated lectotype here).
N. mascatense A. D C , Prod. 8:421. 1844.Type: Muscat, Aucher-Eloy 4925
(G-DC,holotype (microfiche seen);isotypes seen:BM,FI, FI-W, K, P).
N. kotschyi Boiss.,Diagn. Ser. 1.7:21.1846.Type:Iran: Kuh-Delu, Kotschy
558(G, holotype,not yetseen;isotypes seen: BM, BP,C, FI, FI-W, K, P, UPS,
W,WAG). Homotypic synonyms:N. odorum Soland. var. kotschyi (Boiss.) Bo-
iss., Fl. Or. 4: 48. 1875. N. indicum subsp. kotschyi (Boiss.) Rech. f. Fl. Iran.
103:3.1974.
N. oleandersubsp. kurdicum Rech, f., op.cit. 2.Type:Yugoslavia: Dalmatia,
Rechinger 12141(W, holotype).
Shrub or small tree 1-6 m high. Trunk 2-15 cm in diam. or more (?); bark
smooth, pale to dark grey or pale grey-brown. Branches flexible, bearing a ter-
minal inflorescence subtended by 3 equal branchlets, therefore candelabrum-
shaped (modelof LEEUWENBERG asdescribedby HALLE&OLDEMAN, 1970,Essai
sur l'architecture et la dynamique de croissance des arbres tropicaux, Paris);
branchlets slightly triangular at the apex, furthermore terete, green, glabrous
or obscurely puberulous at the apex, 3-6 mm in diam., with clear sticky sap.
Leaves ternate (exceptionally some opposite or alternate), with a single row of
colleters in the axils, those of a whorl equal, shortly petiolate; petiole 3-10 mm
long, glabrous or puberulous; blade (often thickly) coriaceous, very narrowly
elliptic, 4-8 x as long as wide, 5-21 x 1-3.5 cm, acuminate or acute at the
apex, cuneate at the base or decurrent into the petiole, smooth when fresh, dull
and with a more or less granular structure of impressed veins above when dry,
revoluteattheentiremargin,glabrousonboth sidesorsometimespartly puberu-
lousbeneath; numerous straight rather obscure lateralveins.
Inflorescence terminal or in the forks, more or less thyrsoid, lax, especially
in the first branchings, variable in size, often long-pedunculate. Peduncle,
branches and pedicels angular, pubescent or glabrous. Bracts sepal-like,
1-2 x aslongasthem, with colletersintheaxils,outside pubescent or glabrous.
Flowers 5-merous, actinomorphic except for the subequal sepals, fragrant or
not. Sepals pale green when corolla white or pale-coloured, suffused with dark
red when corolla dark-coloured, free, narrowly triangular to narrowly ovate,
1.5-4 x as long aswide, 3-10 x 1-3 mm, acuminate, pubescent outside, inside
8 Agric. Univ. WageningenPapers87-2 (1987)
pubescent at the apex and with a single continuous row of colleters
(0.5-0.8 x 0.15mm),imbricateinbud,erector apicallyrecurved. Corollawhite,
palepink,dark pink,wine-red,paleyelloworsalmon, often withdarker longitu-
dinallinesinthroat (doubleonlyseenincultivated specimens),glabrousoutside,
insidewitha(basallyinterrupted)beltoflongrecurvedhairsfrom 3-7mm above
thebaseto theinsertion ofthestamens and from there to theapex of the corona
sparsely pubescent (with ordinary and/or glandular hairs); tube 1.9—4.5x as
long as the calyx, 0.8-1.3 x as long as the lobes, 12-22 mm long, infundibuli-
form, at the base 2-4 mm and at the throat 8-12 mm wide, abruptly widened
from the insertion of the stamens, lobes in the bud overlapping to the right,
obovate or nearly so, 1.2-2 x as long aswide, 13-30 x 8-25 mm, rounded, en-
tire,moreor lessspreading;corona inthecorolla mouth composed of 5truncate
epipetalous parts each bearing 3-8 unequal or less often equal, in many cases
partly united linear or triangular lobes (linear and triangular lobes often on
a single corona part). Stamens barely included, inserted at 7-13 mm from the
corolla base;filaments short, 1-2 mm long,with recurved long hairsinside,gla-
brous outside;anthersnarrowly triangular, fertile at thesagittate base,introrse,
4-6 x 1.2-1.7 mm (except for appendages), with stiff upcurved hairs outside,
glabrousinside,apicallywitha filiform denselyhirsutulous 7-9 mm long appen-
dage (the appendages of the 5anthers usually twisted around each other); cells
2, dehiscent throughout with a longitudinal slit. Pistil 9-15 mm long; ovary
broadly ovoid, often laterally compressed, 1.5-2 x 1.2-2 x 1-1.5 mm, retuse
at the apex, pubescent, composed of 2 fused carpels, with 2 indented lines of
dehiscence; style cylindrical, glabrous, about 0.5 mm in diam.; pistil head
1.2 x 1.2 mm composed of a basal ring (0.4mm high and conical),a cylindrical
0.5 x 0.6 mm central part topped by 2 oblong erect 0.3 mm long lobes; apical
lobes 0.4 mm long. The head adnate to or coherent with the connectives of the
anthers by which the style and pistil head are shed with the corolla. Disk none.
Ovulesineachcellmany.Fruit abicarpellate follicle subtended bythe persisting
calyx, medium brown, narrowly or more frequently very narrowly oblong,
(5.5)7.5-17.5 x 1-1.3 x 1-1.2 cm,glabrous,longitudinallystriate,obtuse at the
apex,narrowed towardsbothends,bivalved,septicidal,many-seeded;wall fairly
thick, woody. Seed medium brown, oblong, 4-7 x 1.5-2 x 1mm, with a 9-12
mm long coma at the truncate apex (directed towards the apex of the fruit),
acute at the base, tomentellous with upcurved hairs grading into the coma; en-
dosperm about 0.3mmthick, starchy,creamy,surrounding theembryo; embryo
straight, spathulate; cotyledons about 1.5 x as long as the rootlet and about
3.5 x as long as wide, oblong, obtuse at the apex, truncate at the base (in a
seed 6.5 x 1.5 x 1mm: embryo 6 mm long; cotyledons 3.5 x 1 mm; rootlet
2.5 x0.6mm).

GEOGRAPHICAL DISTRIBUTION
The distribution map of Nerium oleander(Map 1)is prepared by the present

author using the locality data of the several hundreds of herbarium specimens
cited below.
The localitydata ofthespecimenspresentinthefollowing herbaria have been
assembled by LEEUWENBERG: AAU, BP, BR, C, E, FI, FI-W, G, L, MO, NY,
P, S, UPS, W, WAG, Z. The present author examined the specimens of the
herbaria BM and K.
The oleander is widely cultivated and naturalizes very easily. Therefore, in
many areas the occurrence of Nerium oleander is only subspontaneous. When
a herbarium specimen was evidently or presumably not of truly wild origin,
itsdata werenot processed.
Distribution: Libya, Niger, Tunisia, Algeria, Morocco, Portugal, Spain,

France, Italy, Malta, Yugoslavia, Albania, Greece, Cyprus, Turkey, Syria, Le-
banon, Israel,Jordan, Ethiopia, Oman, United Arab Emirates, Iraq, Iran, Pak-
istan,Afghanistan, India (Kashmir, Punjab), Nepal and China (Yunnan).
ECOLOGY
Nerium oleanderis generally found in a wet habitat on sites well exposed to

thesun.Itoccursespeciallyalongthebanksofstreamsandinthebedsofstreams
and wadis that carry water during at least a part of the year, less often on lake
or seashores; on sand, clay, among stones and in rock crevices.Alt.: from -100
m(near theDead Sea)toabout 2500m (in theAtlas Mountains).
The oleander is a very tolerant and adaptible plant, able to survive floods
aswellaslongerperiods of drought.
It can grow very wellwith its roots immersed. The flexible branches with the
tenacious, narrow and smooth leaves and the extensive root system enable the
plant to survive floods.
Onthe other hand, Nerium oleanderhascharacteristics typicalfor non-succu-
lentxerophytes.Theleafistoughandleathery,coveredwithathickcuticle(scler-
ophyll).The stomata are sunk in depressions on the lower surface of the leaves
(several stomates in each pit) to prevent excessive water loss in drying winds.
Microscopic hairs in these pits reduce air movement around the stomata, and
thustranspiration, evenmore.Thelongrootsenabletheplant totapwater from
deep sources.
All parts of the plant are poisonous. Livestock, especially goats, usually ap-
pear to beaware ofthis and avoid the plant.

MAP 1. Geographical distribution ofnaturally occurring Nerium oleander L.

FLOWERING AND FRUITING PERIODS
Nerium oleander,awide-ranging species,showsavariability inflowering time

throughout itsrange.In thisrespectfivezonescanbe distinguished.
Flowering period
1. Southern France and northern Italy: June-Sept.

2. Portugal, Spain, southern Italy, Malta, Yugoslavia,
Albania, Greece,Turkey and Cyprus: (Apr.) May-Sept.
3. Morocco,Algeria, Tunisia,Niger and Libya: Mar.-Oct.
4. Syria, Lebanon, Israel,Jordan and Iraq: Apr.-Nov.
5. Oman, United Arab Emirates, Iran, Afghanistan,
Pakistan, India, Nepal and China: almost year round
Fruits can be found on the plants at any time of the year in all regions. In
springand summer onlydry fruits are present.
VERNACULAR NAMES
English: Oleander, Rose Bay (USA; another plant called Rose

BayisEpilobium angustifolium), Rose Laurel.
Dutch: Laurierroos, Lauwerroos, Oleander, Oleanderboom.
German: Lorbeerrose, Oleander, Rosenlorbeer, Unholdenkraut.
French: Belladonna (Corsica), Fleur de Saint-Joseph, Laurelle,
Laurier, Laurier-Blanc, Laurier des Jardins, Laurier-
Rose, Laurier-Tropical, Laurose, Nérier à Feuilles de
Laurier, Nérion, Oléandre, Rosage, Rosagine.
Spanish: Adelfa, Alendro, Baladre, Eleandro, Eloendro, Laurel
Rosa, Oleandro, Rosa Francesa.
Catalan: Baladra, Baladre, Ballestera, Biva, Diva, Llorer Rosa,
Oleandre, Sanet.
Basque: Epriotz-orri, Eriotz-orri, Eroitzorri, Erroitzorri.
Galician: Adelfo, Loendreiro, Loendro, Nerio.
Portuguese: Adelfa, Aloendro, Espirradeira, Landro, Loendreira,
Loendro, Loureiro Rosa, Nério, Oleandro, Rosa
Laura, Rosa deCeilao, Sevadilha.
Italian: Alessandrina, Alloro d'India, Alloro Indiano, Alloro
Rosa, Ammazza l'Asino ('donkey killer'), Amnaya
d'Assino, Erba da Rogna, Lauri Rosa (Sicily), Lauro
Indiano, Lauro Rosa, Lauro Roseo, Leandro, Mazza

di San Giuseppe, Nerio, Oleandro, Rosagine, Rosa
Lauro.
Albanian: Rhododaphne.
Greek: Agriodaphne, Nerion, Pikrodaphne, Rhododaphne,
Rhododendron, Sphaka (Crete).
Cyprian: Orodaphne, Pikrodaphne, Rhododaphne.
Syriac: Orodafni.
Hebrew: Ardaf, Harduf.
Arabic: Dafla, Defla, Diffla, Difla, Difli, Dufla, Somm el-him-
ar, Sumelhimar, Summ Al-himar, Tiffla, Tofla, Ward
el-himar.
Kurdish: Zhala.
Persian: Khar Zahr ('donkey poison', 'asses bane'), Kharzah-
rah.
Sanskrit: Asvamaraka, Chandata, Hayamaraka, Karavira, Pra-
tihasa, Virahuha, Vishavrykshanka.
Hindi: Kanel,Kaner, Karber, Kuruvira.
Tamil: Agam,Alari,Alarida,Aralee,Arali,Asuvabari, Irattai-
chivappalari, Kanaviram, Karaviram, Katturepatta,
Kaviram, Kayiram, Sevvalari, Trattaichegappayalari,
Urattaivellaiyalari, Vellalari, Vellaiyalari.
Bengali: Karabi, Lal-kharubee, Pudma-kurubee.
Baluchistan: Gandeli,Jaur, Jowari.
BolanHills: Jaur.
Bombay: Ganira, Kanher, Kanhera, Kanir.
Canarese: Biligampuganagile, Biliganagile, Kanagile, Karavira,
Kempuganagile, Paddali.
Deccan: Kanirkejur.
Fort Sandeman: Ganderae.
Gujerati: Kanera, Karen, Lalkaren.
Khaisar: Jaur.
Kumaon: Kanur, Kanyur.
Las Bela: Kuran.
Malayalam: Alari, Arali, Chuvannarali, Kanaviram, Karaviram,
Raktapushpam, Satraprasam, Veluttarali.
Mamu: Jaur.
Marathi: Kaneri, Kanher.
Mundari: Kanaili.
Punjabi: Ganera, Ganhira, Kaner, Kanira.
Pushtu: Ganderai.
Ranikhet: Kanpuri.
Sadani: Kaner.
Santal: Rajbaka.
Sharing: Gander.
Sibi: Jaur.
Tang Masezai: Gandher.
Telugu: Ettaganneru, Ganneru, Karaviram, Kasturipatta.
Urdu: Kanir.
Uriya: Konero, Konioro, Koniyoro, Korobe, Korobiro, Kor-
obyo.
Chinese: Kap chukt'o ('mingle- bamboo- peach blossom'), Kia
Tchou T'ao.
Japanese: Kjotikto, Kjotjeksto.
Sundanese (W.Java): Djoerè, Jure.
PhilippineIs.: Adelfa, Baladre.
Tagalog: Baladri.
FijiIs.: Vasa.
Tonga Is.: Lolie.
Hawaiian: Oleana, Oliana, Oliwa.
West Indies: South Sea Rose.
Dominican Republic: Flor del Peru, Laurel Rosado, Martinica, Piruli, Rosa
del Peru.
Puerto Rico: Adelfa, Alelia, Alheli Extranjero, Laurel, Laurel Rosa-
do, Oleander, Sea-side Rose.
Antilles: Rosa Francesa.
Mexico: Laurel, Laurel Rosa, Oleanda, Rosa Laurel.
Guatemala: Narciso.
Venezuela: Rosa de Berberia, Berberia.
Colombia: Azucena de la Habana, Flor de la Habana, Laurel de
Judea.
Argentina: Laurel Rosa.
SPECIMENS EXAMINED
Aselection ofthespecimensseenby LEEUWENBERGand/or PAGEN
LIBYA:Derna (fl.,fr.) Gregory 23.VII.1908 (BM,MO);ibid.(fl. May)Pampanini&Pichi-Sermolli

6142 (FI, K); ibid, (fl., fr.) Palma 10.IV.1926 (FI); ibid, (fl., fr. Mar.) Davis 50372 (K); 24 kms.
W. ofDerna (fr. Jan.) Loufty Boulos 1204(S); ElBarcat (fr. Mar.) Corti 20,21,22 (FI).
NIGER:Araka (fl., fr. Mar.) Meniertzhagen 176(K).
TUNISIA: Qabis (buds, fr. Mar.) Pitard 190 (K, MO, WAG); ibid. (fl.) Kralik 9.V.1854 (FI-W);
BouArada (fl., fr.) LeTestu7.IX.1898 (BM);Zaghwan (fl.)Kralik 3.VIII.1854(FI-W);AIHamma-
mat (fr. Oct.) Gandoger 106 (K, MO); Nabul (fr. Oct.) Gandoger 106(C, K, Z); Tunis (fl.) Aberg
anno 1848 (S);Jazirat Zambrah (buds) Ochsenius IX.1865 (Z);Tabarqah (fl., fr. July) Jansen 620
(WAG):ibid.(fl. May)Davis&Lamond D.57848 (BM).
ALGERIA: Annaba (fl., fr.) Decaisne 7145 (BR); ibid. (fl., fr.) Dukerley s.n. (BR); between El
Aouana and Jijel (fl. May) Davis 52792 (BM); Alger (fl., fr.) Gandoger 155 (BM, C, FI); ibid.
(fl.) Woelmont s.n. (BR); Hammam Melouane (fr. Oct.) Kramer 5321 (Z); Blida (fl., fr.) Bellot
14.VIII.1869 (FI); ibid. (fl.) Chabert VI.1871 (FI); ibid. (fl. May, fr. Nov.) Jamin 59 (FI-W, K,

WAG); ibid. (fl. July) Salle 129 (C, P, WAG); ibid. (fr.) Bommer s.n. (BR); between Blida and
Chiffa (fl.)Thoren III. 1881(S);Médéa (fr.) Chabert 5.X.1871 (FI);Berrouaghia (fl.) Luizet6.X.1886
(MO); Boghar (fl.) Debaux VI.1856 (NY); ibid. (fr.) Debaux X.1856 (NY); ibid. (fl.) Debaux
2.VII.1856 (BM, BR, FI, S, WAG); ibid. (fr.) Debaux 25.XII.1855 (BM, BR, FI, S, WAG); ibid.
(fl., fr.) Gandoger anno 1906-1907 (NY); Cherchell (fl.) Hafstrom 10.V.1925 (S);Theniet El Had
(fr.) Chabert 24.IX.1871 (FI); Djebel Maïz (fl.) Uggla 18.IV.1936 (S); Kristel (fl., fr.) Debaux
4.VIII.1881 (S); Oran (fr.) Debaux anno 1892 (FI); ibid. (fr.) Gandoger anno 1909 (FI, K, MO);
ibid. (fr., fr.) Romain VII. 1849 (BM); Sig (fl., fr.) Durando anno 1850 (FI-W); Lauriers Roses
(fl.) Faure 20.VI.1912 (BM, FI, Z); Sidi Bei Abbes (fl., fr.) Warioy 1.VII.1876 (FI); Tlemcen (fl.)
Faure 25.VIII.1932 (S); between Rhoufi and Biskra (fl.May)Davis 52377(BM);Biskra(fl.) Balansa
IV.1853 (FI-W); ibid. (fr.) Chevallier VI.1897 (FI, WAG); ibid. (fl.) de Geer V. 1891 (S, UPS);
ibid. (fl.) Rouseau V.1927 (BR); between Laghouat and Aflou (fr. May) Urwi 62 (Z); Am Sefra
(fl. May) Chevallier 342(FI, WAG); ibid. (fl. May) Hochreutinger 523(Z);Beni Ounif (fl.) Sander-
mann Olsen 25.V.51 (C); Béchar (fl.) Andreânszky 22.IV.1928 (BP); ibid. (fl. July) Harshberger
1127 (NY); Ahaggar (veg. July) James 11 (BM); Guelta Afilale (fl., fr. Aug.) Van Impe 32 (BR);
Garet Rouba (fl., fr.) Faure 2.VIII.1933 (BR, MO); 72 kms. N.W. of Tamanrasset (fr.) Croston
6.II.1978(MO);
MOROCCO: 25 kms. N. of Agadir (fl., fr. Mar.) Miller et al. 408 (BM, MO); Tamri (fl., fr. Mar.)
Davis 48461 (BM, NY); Oued Tamri (fl) Bannerman 2.III.1952 (BM);Tachdirt (fl.) Andreânszky
27.VI.1930 (BP); Oumenast (fl. Aug.) Williamson 519 (BM);Tislit Valley (fl., fr. Aug.) Newbould
184(BM);Jemaa de Mrirt (fl. July) Davis 55161 (BM, MO);Ifrane (fl. May) Jahandiez 447 (BM);
Oued Kroumane (fl. July) Woodeil 105(BM); Bab Bou Idir (fl. Aug.) Fry 42 (BM);Taza (fl.) An-
dreânszky 17.V. 1930(BP);Targuist (fl.June)Font Quer 486(BM);AIHoceima (fl., fr. Aug.) Heath
378(BM,K);CapdesTrois Fourches (fl.) Gandoger IV. 1908 (MO);Melilla(fl.) Sennen&Mauricio
18.V.1931 (BM); Sebkha Bou Areg (veg.) Gandoger IV.1909 (NY); Berkane (bud Apr.) Wilczek
etal.873(MO);Taforalt (veg.)Faure 2.V.1930(FI).
PORTUGAL: Mora (fl. June) Raiuha 3603 (S); between Mertola and Pomarao (fl. June) Moller
1286 (COI, Z);Tavira (fl. May) Welwitsch 229 (FI-W, K);Faro (fl. July) Bourgeau 1952(BP, BR,
C,FI-W, K, WAG).
SPAIN: Alcala de los Gazules (fl. June) Bourgeau 326 (BM, FI-W, K, P, WAG, Z); Algeciras
(fl., fr.) Winkler 12.V.1873 (BP); ibid. (fl.) Reverchon 26.VI.1987 (BM); ibid. (fl. Aug.) Ellman &
Hubbard 395 (K); San Roque (fr.) Regneil 27.XII.1938 (S); Ronda (fl.) Reverchon 20.VII.1889
(BM); Ojén (fl.) D.B.L. 5.V.57 (BM); Monda (veg.) Thornberg 30.IV.1968 (C); Malaga (fl. Apr.)
Brandt 950 (S); ibid. (fl.) Cyrén 4.V.1924 (S); ibid. (fl.) Erlandsson 23.VII.1952 (S); ibid. (fl., fr.)
Lange 1.1872 (C, UPS); between Canillas de Aceituna and Sedella (fl. Aug.) Goyder & Jury 452
(BM); Almunécar (fl.) Roivainen 25.IV.1952 (UPS); Almeria (fl. Apr.) Glanville 197 (BM); ibid.
(fl. Apr.) Stace 141A(BM); ibid. (fl. June) Ballet al. 222(BM);4 kms. W. of Almeria (fl., fr. Apr.)
Gardner 1694 (BM); Alcolea del Rio (fl.) Anonymus 1810 (FI); between Cordoba and Sevilla (fl.)
Garnett V.1932 (BM); between La Carlota and Cordoba (fl. May) Dandy 1473 (BM); between
Arquillos and Canena (fl. May) Pedersen 10363 (C); Murcia (fr.) Lange 19.XI.1851 (C);Alcoy (fr.
May) Dechamps & Doutrelepont 2188 (BM); Chiva (fl.) St.-Lager 15.VI.1890 (K, NY); Valencia
(fl., fr.) Bicknell 1.IX.1905(BM); Segorbe (fl.) Pau 2.VII.1887 (BP, FI); ibid. (fl. Aug.) Willkomm
484 (BM, C); Benicarló (fr. Jan.) Sennen 3984 (BM);ibid. (fl. Aug.) Gédéon 4229 (BM); Roquetas
(fl. May) Nüssen 539 (BM, UPS); Tranquera (fl., fr.) Galathea Exp. 1845-1847 (C); Barcelona (fr.
Dec.) de Sequenza 274 (FI); ibid. (fl.) Sennen 19.VI.1929 (BM); Ibiza; San Antonio Abad (fl., fr.)
Lindahl 21.VI.1870 (S,UPS);Santa Eulalia del Rio (fl. June)Welti-Hug 694(Z);EsCana (fl. June)
Cannon 3287(BM).
FRANCE: Toulon (bud, fr. Sept.) Koehler 4926 (BR);ibid. (fl.) Loiseleur s.n. (S);ibid. (fl.) Tholin
7.VI.1880 (Z);La Londe-les-Maures (fl., fr. June) Coraze 1358(BR);ibid. (fl.) Mercurin 6.VI.1934
(BR); La Garde-Freinet (fr.) Desplantes VIII.1926 (S); Roquebrune (fl.) Bertrand VI.1899 (FI);
Agay (fl.) de Maupassant 24.VI.1885 (Z);ibid. (fl.) Vidal 3.VI.1888 (S);ibid. (fr.) Vidal 24.X.1888
(S); Corsica: St. Florent (fl.) Hafstrom 10.VI.1930 (S);ibid. (fl., fr.) Autheman 11.1.1874(BP);ibid.
(fl. Aug.) Gabriel 1358 (BM, Z); ibid. (fl., fr. Aug.) Kralik 692 (FI-W, K, WAG); ibid. (fl., fr.)
Levier 3.VII.1880 (FI); ibid. (fl. July, fr. Oct.) Mabille 158 (BM, BR, FI, K); ibid. (fl.) Pelgrims
Agric. Univ. WageningenPapers87-2 (1987') 15

VII.1956 (BR);ibid. (fl.) Rauch 14.VIII.1922 (Z);ibid. (fl. July) Segal 232(WAG); ibid.(fl.) Schinz
27.IX.1922(Z).
ITALY:Tignale(fl.) Rigo 12.VII. 1878 (BR,UPS,Z);Dolceacqua (fl.) Hesselman 18.V. 1896(UPS);
Ventimiglia(fl.June,fr. Oct.)Pollini 2318(BM, BP,FI,K,Z);between Bordighera and Ventimiglia
(fl.) Barla VI.1847 (FI); Bordighera (fl.) Bicknell 11.VII.1892 (BR, FI, NY); ibid. (fl., fr.) Bicknell
19.VI.1890 (AAU, S, UPS); ibid. (fl.) Seiffert 16.VII.1890 (BM, BP);ibid. (fl.) Bicknell 12.VI.1906
(S); ibid.(fl.)Bicknell30.VII.1890(BM); Imperia (fl.)CorradiVIII.1938(FI);Andora (fl„ fr.) Lacai-
ta 27.VIII.1882 (BM); between Andora and Capo Mêle (fl.) Piccone 1.VI.1873 (FI); Genova (fl.)
Woods 540/1 (BM); Pisa (fl.) Van Heurck anno 1868 (BR); Sansepolcro (fl.) Pichi-Sermolli
17.VII.1932 (FI);Antignano (fl.) TaniVII.1930 (FI);Istia(fl.) Marchesetti 2.VII.1866(FI); Tortora
(fl.) F.Q. 24.VI.1917 (BM); Nova Siri (fr. Oct.) Gavioli 28291 (FI); Trebisacce (fl.) St.-Lager
4.VII.1891 (BM, NY); Gallipoli (fl.) Bradford 1926 (BM); Neto R. (fl.) Fiori 26.VII.1883 (FI);
Palizzi (fl.) Biondi 5.V., 4.VI., 7.VI.1877 (FI); Reggio di Calabria (veg.) Caruel 25.X.1883 (FI);
Capraia (fl.) Bavazzano 23.VI.1969 (FI); ibid. (fr.) Bavazzano 30.III.1969 (FI); ibid. (fr.) Béguinot
& Donia 6.1.1898 (FI); ibid. (fr.) Requien 28.X.1847 (FI-W); ibid. (fl., fr.) Sommier 23-29.V.1910
(FI); IsoladelGiglio(fl.,fr.) FlaruGulio 26.IX.1894(FI);ibid.(fl.)Tonina anno 1894(FI); Sardinia:
Orune (fl.) Martelli 7.VI.1899 (FI); Dorgali (fl.) Martelli 19.VI.1895 (FI); Arbatax (fl.) Martelli
26.V.1895 (FI, S); Iglesias (fl. July) Müller 1828-4 (BP, BR, K); ibid. (lg. Mar.) Fiori 120 (BM);
Cagliari(fl.) Sommier 12.V. 1872(FI);Alcamo(fl.) Fleming5.X.1843(BM);Mondello (fl.) Sommier
3.VI.1873 (FI); ibid. (fl.) Strollz 4.VII.1873 (BP); Palermo (fl.) Todaro anno 1849 (S); ibid. (fl.,
fr. July) Todaro 472 (BM, BP, BR, FI, K, S); ibid. (fr.) Todaro 2.XII.1921 (BM); 15 kms. E. of
Cefalü (fl. May) Davis & Sutton 64032 (BM); Mazzarra (fr.) Borzi XI.1875 (FI); Mandanici (fl.)
Aiuti V.1872 (FI); Messina (fl.) Sequenza IV.1865 (FI);Alcantara R. (fr.) Karpati 12.X.1961 (BP);
10 kms. N.E. of Taormina (fl., fr. July) Akeroyd et al. 3585 (BM); Taormina (fl.) Borzi VI.1881
(FI); ibid. (fl.) Borzi 6.VII.1884 (BP); ibid. (fl.) Groves 29.V.1926 (BM); Brucoli (fl. Apr.) Larsen
et al. 35721 (AAU); Ragusa (fl. June) Kundise 629(AAU); Camastra (fl., fr.) Martelli 11.VII.1906
(BP).
MALTA: Malta (fl.) Pariatore 15.IX.1834(FI).
YUGOSLAVIA: Split (fl.) Prior VI.1843 (K); Cannosa (fl.) Vetter 28.V.1926 (S);Ombla (fr.) Ado-
movic VII.1907 (BP); ibid. (veg.) Lengyel 13.IV.1909 (BP);Dubrovnik (fl.) Barrat VIII-420 (NY);
ibid. (fl.) Pichler V.1872 (BP, K);ibid. (fl.) Schlüters9.VI.1880 (AAU, BM, S,UPS,WAG);4kms.
E.ofMorinje (fl., fr. Aug.) Forst-Olsen 1444(AAU); HercegNovi (fl.) Lenander 16.VI.1938(S).
ALBANIA:Vlore(fl., fr. Sept.)Aiston &Sandwith 2779(BM, K);ibid. (fl.) Baldacci 1902(BM).
GREECE: Kefallinia (fr.) Schimper & Wiest X-XI.1834 (FI, K); Kalavrita (fl.) Bornmüller 1105
(Z); Patras (fl. May) Mattfeld 141 (K, MO); Dhiakopton (fl., fr.) St.-Lager 20.VII.1899 (BM, K,
NY); Argos (fl.) St.-Lager 26.VII.1899 (C, G); Porös (fl.) Heard 1924 (BM); Mt. Imittos (fl. June,
fr. Sept.) De Heldreich 960 (BM, C, FI, K, P, S, UPS, Z); ibid. (fl. Aug., fr. Oct.) Atchley 368
(K); Marathon (fl.) Walker 5.VI.1882 (BM); ibid. (fl.) Rogers 433 (K); Kimi (fr. Aug.) Rechinger
37867 (AAU); Skiros (fl. Apr.) Rechinger 744 (BM); Athos (fl.) Trivaleszky 731 (BP); Samothraki
(fl.) Cyrén 13.VI.1932 (S); ibid. (fl. June) Rechinger 9935 (BM); Khios (fl. May) Rechinger 5353
(BM); ibid.(fl.July)Platt 338(K);Sifnos (fl., fr.) Royal Liberty School22.VIII.1963(K); Karpathos
(fl. June) Greuter 5819 (Z);ibid. (fl. June) Rechinger 8112 (BM); Tilos (fr.) Desio 14.X.1922 (FI);
Crete: Kastelli (fl. June) Baldacci 186 (BM, MO); Platanias (fl.) Reverchon 4.VII.1883 (BP, FI,
K, UPS);Gavdhos (fl.June)Rechinger 13641(K,S);Khersonisos Akrotiri (fl.)Gandoger 6.VI.1914
(K. MO); Iraklion (fl.) Wall 27.VII.1930 (S); Kavousi (fl., fr. June); Lemperg 637 (K); Sitia (fl.
May) Rechinger 12673(BM); Rhodes:Kattavia (fl.) Jannone 17.VIII.1934 (FI);ibid. (fl.) Muzzoc-
chi-Alemanni26.IV.1922 (FI); Lindos (fl.June) Rechinger 8440(BM, K).
CYPRUS: Paphos (fl. July) Stavros 42 (BM); Kalavasos (fl. May) Holmboe 621 (C); Kyrenia (fl.
July) Atherton 222 (K); ibid. (fl. July) Atherton 189 (K); ibid. (fl. June) Mapple 86 (K); Nicosia
(fl.) Lindberg 6.VI.1939 (K); ibid. (fl. June) Syngrassides 414 (K); Kythrea (fl.) Sintenis V.1880
(S); AyiaNapa (fl.) Lindberg 10.VII.1939 (S).
TURKEY: Büyükdere (fl.) Berggren s.n. (UPS); Canakkale (fl. Aug.) Sintenis 814 (BM, BR, K,
P, S);between Bergama and Izmir (fl. May) Dudley D34850 (K); Izmir (fl. May) Bornmüller 9764
(P); Aydin (fl. Febr.) Stutz 444 (NY); Marmaris (fl., fr. July) Khan et al. 34(K); Antalya (fl. May)
Tengwall 627(K, S);ibid. (fl.June)D.E.S.T. 79(K);Alanya (fl. Sept.) Baytop 10602(BM); Anamur

(fl.) Peronin 83 (K); Mersin (fl. May) Hennipman et al. 1080 (K, WAG); Adana (fl. May) Balls
1191 (BM, K); Yumurtalik (fl., fr. Sept.) McNeill 806 (K); Osmaniye (fl. May) Davis 42251 (K);
Beien(fl.,fr. July)Gombault 439(P);Iskerendun (fl.Apr.) Davis&HedgeD26941(BM,K);Yayla-
dagi (fl.May) Dinsmore 16987(S);Mardin (fl.,fr. May) Davis 42659(K).
SYRIA: between Aleppo and Antakaya (fl., fr.) Rufish s.n. (BM); Saïda (fl. June) Blanche 692
(P); ibid,(fl., fr.) Gaillardot 5.VII.1855(BM,BR, C, FI, K, P.S,WAG).
LEBANON:Al Majdal (fl., fr.) Lowne 1863-4(BM, K);Bhamdun (fl.) Wall 30.V. 1934(S).
ISRAEL: Lake Hula (fl. Oct.) Jones 224 (BM); Ayyelet Hashahar (fr. July) Eig 10552 (BM); ibid,
(fl., fr. Apr.) Steinberg & Bavazzano 2539 (FI); ibid, (fl., fr. July) Eig 10538 (K); between Lake
Hula and Lake Tiberias (fl., fr. Nov.) Taylor MEG (K); Tiberias (fl.) Osborne 271 (K); Teverya
(fl., fr.) Linder anno 1912(UPS); Har Karmel (fl. Apr.) Steinberg & Bavazzano 3298 (FI); Sha'ar
Ha'amaqin (fl., fr. Oct.) Zohary &Amdursky 456 (AAU, BM, BR, C, FI, K, MO, NY, P, S, UPS,
WAG, Z):Jerusalem (fl. May) Meyers&Dinsmore B2987(S).
JORDAN: Jordan Valley (fr.) Berggren 25.X.1821 (UPS); ibid. (fr.) Michon 5.III.1851 (P); Wadi
At Tayyibah (fl. Apr.) Steinberg & Bavazzano 2992 (FI); Jarash (fl.) Crowfoot 70 (BM); between
Amman and Jarash (fl. Nov.) Carr 15 (NY); Al Mazra'ah (fl. Apr.) Dinsmore 18987 (K, S, Z);
Petra ruins (fr. Apr.) Davis 8771 (K); Wadi Musa (fl. Apr., fr. Oct.) Dinsmore 10987 (K, S); ibid,
(fl.) Marsh4-10.VI.1854(MO);JabalAbu Khushaybah (fr.) Hart XI.1883, II. 1884(BM).
ETHIOPIA: Gheleb (fl.) Rodén 1912-20(S).
OMAN: Salalah (bud) Sammicheli MI.1974 (FI); Sur (fl., fr. Mar.) Radcliffe-Smith 3924 (K);
Ghawrab (fl.Apr.) Lee-Oldfield 36(BM);Wadi Sahtan (fl.,fr. Apr.) Mandaville 6142(BM); Bidbid
(fl. Mar.) Mandaville 3433 (BM); Muscat (fr. Jan.) Bornmüller 501 (BM, BR, K, P, Z); ibid, (fl.,
fr. Jan.) Bornmüller 502(BM, BP, BR, FI, K, P, Z);ibid, (fl.) Leclancher 32 (P);ibid, (fl.) Aucher-
Eloy4925 (BM, FI, FI-W, K, MO, P);Jabal al Akhdar (fl. Mar.) Edmondson 3479 (E);Ar Rustaq
(fr. Oct.)Miller &Whitcombe 2779(E).
UNITED ARAB EMIRATES: Al Fujayrah (fl. Feb.) Edmondson 3133 (E); between Al Khatt and
Daba (fl.May) Western 177(E).
IRAQ: Zakho (fl. July) Rechinger 12141 (AAU); ibid, (fl., fr. June) Yusuf Lazar 815 (MO); 5
kms. S. of Zakho (fl. July) Rawi 23117 (K); Rost (fl., fr. Oct.) Guest 278 (Z); ibid, (fl., fr.) Yusuf
Lazar 21.X.1932 (MO); between Arbil and Rawandiz (fl. June) Bornmüller 1541 (BR, K, P);Arbil
(fr.) Guest 22.III.1930(K); 13 kms.S.W. ofSylaymaniyah (fl.June)Rawi21710(K);between Mosul
and 'Aqrah (fl. Sept.) Lizzi&Omar 35280(K);Abu Ghurab (fl.) Hikmat Abbas 31.VII. 1961 (K).
IRAN: Shahpur (fr.) Bélanger 720 (P); Sheshom (fl. June) Jacobs 6800 (E, GRO, K, L, MIN,
W); between Khorramabad and Andimeshk (fl.Apr.) Wright &Bent425-101 (K);Dezful (fl. May)
Koie 1253 (C); ibid. (fl. June) Loftus 33 (BM); Behbehan (fl.) Hausknecht VI.1868 (BM, K, P);
Kazerun (fr. Feb.) Koie 44 (C); Konar Takhted (fr. Dec.) Bornmüller 513 (K); Shiraz (fr. Dec.)
Bornmüller 3849 (BM, BP, BR, K, Z); ibid. (fr. Nov.) Grant 15091 (MO); ibid (fr. Oct.) Pravitz
356 (S);Kerman (fr. Sept.) Bornmüller 2402(BP, WAG); ibid, (fr.) Gandoger VII.1906(MO); be-
tween Shahdab and Kerman (fr. Sept.) Bornmüller 3850(BM, BP,BR,C, FI, K, P,S,UPS, WAG);
Shahdab (fl.) De Bunge IV.1859 (FI, P); Gudiz (fl. May) Léonard 6248 (BR); between Bam and
Jiroft (fr. Apr.) Léonard 5615 (B); ibid. (fr. Apr.) Léonard 5616 (K); Halil Rud (fl. June) Assadi
et al. 1790(NY);Tarom (fl. Apr.) Rechinger 3282 (S);240 kms. S. of Sa'idabad (fl. May) Léonard
5924 (BR, K); Manujan (fl. Apr.) Léonard 5883 (BR, K); Bandar'Abbas (fl., fr.) Fariab 1.XII.06
(K); Minab (veg.) Collet VIII.1886 (K); Bandar-e-Lengeh (fl., fr. Mar.) Davis & Bokhari D.56161
(K, MO);Nikshahr (fl. Mar.) Faraughi 10665(E).
PAKISTAN: Gwadar (fl. Apr.) Rechinger 27785 (S); between Mand and Suntsar (fl., fr. Feb.) Ali
et al. 1085 (MO); Kalat (fr. Apr.) Rechinger 27379 (AAU, S); between Turbat and Gwadar (fl.,
fr. Apr.) Lamond425(E);Khadeji (fl.,fr. Apr.) Lamond 761 (E);Sibi(fl.July)Andersen &Petersen
479(AAU, C, K);between Nushki and Quetta (fl. May) Sultan ul Abedin 3236(NY); Fort Munro
(fl.) Wilson 30.VIII.1921 (P); Fort Sandeman (fl. May) Duthie 20588 (K); Ara (fl. May) Bis Ram
443 (NY); Rawalpindi (fl. Apr.) Aitchison 464 (K); between Attock and Khushalgarh (fl. May)
Rechinger 30417 (AAU); Shakar Parian (fl.) Manzoor 24.X.77 (MO); Malpur (fl. Aug.) Akram
& Dilawar 32(MO);ibid. (fl.) Manzoor &Javeed 21.X.74 (MO); between Murree and Rawalpindi
(fl.) Bellew 19.VI.1873 (K); Siran Valley (fl. June) Inayat 19938 (K); Chitral (fl., fr.July) Harriss
16364(K);ibid. (veg.Oct.)Toppin 817(K).

AFGHANISTAN:Barikowt (fl., fr. July) Edelberg 1627(C);Chigha Sarai(fl.July)Lamond 2541 (E).
INDIA: Punch (fl., fr. Nov.) Schlagintweit 12606 (BM); between Doda and Ramnagar (fr. July)
Saran et al. 29908 (Z); Pathankot (fl.) Drummond 989 (K); Dharmsala (fl. Sept.) Gammie 18752
(K); Bhadwar (fl. Apr.) Koelz 4286 (MO, NY); Baijnath (fl. May) Koelz 4607 (NY); Mandi (fl.
May) Koelz 8326 (NY); Larji (fl., fr. Nov.) Koelz 3125 (NY); Kotla (fl. May) Stewart 1816 (NY);
Simla(fl. Sept.) Drummond 23559(K,P);ibid, (fl.) Drummond 22622(K);Sirmur (fr. Dec.) Drum-
mond 20625 (K); Kapkot (fl.) Strachey &Winterbottom s.n. (BR); Saharanpur (fr. Aug.) Gamble
25664 (K); Kheri (fl. Apr.) Inayat 22148 (K); Abu (fl. Oct.) Gamble 6662 (K); Someswari R. (fl.
May) Legge63(K).
NEPAL: Silgarhi (fl. Apr.) Bis Ram 275 (NY).
CHINA:Yunnan: Yangpi Pass(fl. Sept.)Ducloux 4432(P);Chi-tsu Shan (fl. Mar.) McLaren 153C
(E).
NOMEN NUDUM
Nerium grandiflorum Desf., Tab.ed.2:92. 1815 = N. oleanderL.
EXCLUDED SPECIES
Not thosepublished in synonymy only.
Nerium antidysentericum Lepech., Reise 1: 270. 1800 = Apocynum venetum

L. = Trachomitum venetum (L.)Woodson (exWoodson Ann. Miss.Bot. Gard.
17: 159.1930).
N. antidysentericum L., Sp. PI. 209. 1753 = Wrightia antidysenterica (L.) R.
Br.
N. caudatum (L.) Lam., Encycl. 3:458. 1789 = Strophanthus caudatus (L.)
Kurz.
N. caudatum Roxb., Hort. Beng.84. 1814,nomen; Fl. Ind. 2:9. 1832 = Stro-
phanthus wallichiiA. DC.
N. chinense Hunter ex Roxb., Hort. Beng. 84. 1814, nomen; Fl. Ind. 2: 9.
1832(as chinensis) = Wrightia religiosa(Teijsm. &Binnend.) Benth. ?( = sup-
posed by Leeuwenberg,pers.comm.).
N. coccineum Roxb., Hort. Beng. 19. 1814, nomen; Fl. Ind. 2: 2. 1832 =
Wrightia coccinea(Roxb.) Sims.
N. coraiaBuch.-Ham. exA. D C , Prod. 8:407, 421. 1844,in syn. and nomen
= Wrightia arborea(Dennst.) Mabberley.
N. coronarium Jacq., Coll. 1: 138. 1787 = Tabernaemontana divaricata (L.)
R. Br.exRoem.&Schuit.
N. divaricatum L., Sp. PI. 209. 1753 = Tabernaemontana divaricata (L.) R.
Br.exRoem. &Schuit.
N. divaricatum Lour., Fl. Cochinch, 1: 142. 1790 = Tabernaemontana divari-
cata(L.)R. Br.exRoem.&Schuit,(asfor thename)and Wrightia antidysenteri-
ca(L.) R. Br.(asfor thedescription and thepre-Linnean synonyms).
N. divaricatum Thunb.,Fl.Jap. 110.1784 = Tabernaemontana divaricata(L.)
R. Br. exRoem. &Schuit, (asfor the name )and Trachelospermum jasminoides
(Lindl.)Lem. (asfor the description).
N. grandiflorum Roxb., Hort. Béng. 19. 1814, nomen; Fl. Ind. 2: 10. 1832
= Cryptostegia grandtflora (Roxb.) R. Br.exLindl. (Asclep.).
N. guineense Brongn. ex Perrot & Vogt in Trav. Mat. Méd. Paris 1912, 9:
61(1913) = Strophanthus gratus (Wall.&Hook.) Baill.
N. obesumForssk., Fl.Aegypt. Arab.205. 1775 = Adenium obesum (Forssk.)
Roem.&Schuit.
N.piscidium Roxb., Hort. Béng. 19.1814,nomen; Fl.Ind. 2:7. 1832 = Melo-
dinusmonogynus Roxb.
N. reticulatum Roxb., Hort. Béng. 19. 1814, nomen; Fl. Ind. 2: 8. 1832 =
Cryptolepis buchananiiRoem. &Schuit.(Asclep.).
N. salicinum Vahl, Symb. Bot. 45. 1791 = Breonadia salicina (Vahl) Hepper
&Wood, Kew Buil. 36:860. 1982.
N. scandens Lour., Fl. Cochinch. 143. 1790 = Strophanthus caudatus (L.)
Kurz.
N. scandensThonn.,inSchum.&Thonn., Beskr.Guin.PI. 148.1827 = Alafia
scandens(Thonn.) De Wild.
N. sibiricum Medic, Beobacht. 15. 1780 = Apocynum venetum L. = Tracho-
mitum venetum (L.) Woodson (ex Woodson Ann. Miss. Bot. Gard. 17: 159.
1930).
N. sinenseHunterexRidley,inJourn.As.Soc.Straits53:81.1909 = Wrightia
antidysenterica (L.) R. Br.(deduced herefrom Ridley'stext).
N. tinctorium Roxb., Hort. Beng. 19. 1814, nomen; Fl. Ind. 2: 4. 1832 =
Wrightia tinctoria R. Br.
N. tomentosum Roxb., Hort. Beng. 84. 1814, nomen; Fl. Ind. 2: 4. 1832 =
Wrightia tomentosa (Roxb.) Roem. &Schult. = W.arborea(Dennst.) Mabber-
ley.
N. undulatum (Lam.) Salisb.,Prod. 148. 1796 = Pittosporum senacia Putted.
N. zeylanicum L., Centur. PI. 2: 12. 1756 ex L. Amoenitates 4: 309. 1759 (as
N. zeylonicum) = Wrightia antidysenterica (L.) R. Br. (ex Ngan Ann. Miss.
Bot. Gard. 52: 166.1965).
POLLINATION OF THE OLEANDER

LOCATING THE STIGMATIC AREA ON THE PISTIL IN NERIUM
OLEANDER L. BY MEANS OF FLUORESCENCE MICROSCOPY
INTRODUCTION:Theflower ofNerium oleanderfeatures apistilwithamorpho-

logicallyhighlydifferentiated pistilhead (Fig. 1).
a b c
FIG. 1. Diagram of the pistil head of Nerium oleander L. a. as visible without enlargement; b. in
transverse section as revealed by microscopy: the specific shape appears to be determined by the
length of the hairs, as suggested by the hatched lines; c. pollen grains will germinate on any part
of thepistil head, but penetration isexclusivelypossibleinthearea indicated by arrows.
The entire pistil head iscovered with bristly hairs, which determine, by their
length, the specific shape of the pistil head. The central cylindrical part of the
pistilhead iscovered with rather short hairs,inserted on a glandular epithelium
which produces an adhesive. At the upper edge of the cylindrical part the hairs
are longer, pointing outward and upward, thus forming a ring or wreath. On
the lower part of the pistil head long hairs, pointing downward, are arranged
ina sinuous line around the cylinder, forming a collar. Below this collar, which
hangs down from the pistil head, isa circular depression marking the transition
tothe style.Thepistil head istopped bytwo short erect appendages.
Thismorphological differentiation isaccompanied byananalogous function-
aldifferentiation. Onlyspecific areasonthepistilhead havea stigmatic function
and arereceptivefor pollen tubes.
An experiment wasexecuted 1)to pinpoint theexact location ofthe receptive
area, and 2) to test compatibility of self-pollinations and of cross-pollinations
between cultivars.
With fluorescence microscopy and a clearing-squash technique the pollen
tubesinthepistilcan bemade clearly visible.
MATERIALS AND METHODS: Living plants of fourteen single-flowered oleander

cultivars (one of each) were available, most of them carrying several inflores-
cences. The plants were provided by the Dutch Oleander Society. The 50 cm
tallplantsweregrowninpotsof 14cmdiameter,placed ontablesinagreenhouse
atatemperature of21°C(glasshouseroomprovided bytheDepartment of Hor-
ticulture oftheWageningen Agricultural University).
About ahundred pollinationswereexecuted byhand, self-pollinations aswell
as cross-pollinations between cultivars. Precautions were taken to avoid un-
wanted natural pollination: the flowers were emasculated and protected with
cheesecloth covers.
Two days after pollination the pistils were removed from the flowers and
placed in Herr's clearing fluid during 24hours to make the tissues transparent.
The clearing fluid is composed of lactic acid (85%), chloral hydrate, phenol,
clove oil, and xylene (2:2:2:2:1, byweight) (HERR 1971).After washing in eth-
anol and water the pistils were macerated during 1 hour in 1 n NaOH at 60°C,
washed in water and placed in a solution of 0.1% aniline w.s. dissolved in 0.1
n K3PO4.The material was kept in the aniline solution for at least 30 minutes.
Then the pistils were placed on a slide in a drop of glycerin and covered with
a glass slip. Squashing was hardly necessary as the weight of the cover slip
spreads thecellsofthepistil sufficiently.
ThepreparationswereexaminedwithaZeissStandard WLmicroscope which
had been made suitable for fluorescence observations by mounting a Zeiss UV
HBO 50Watt high pressure mercury vapour lamp, and the filter combination
BP 365 (Exciter filter)/FT 420 (Dichromatic beam splitter)/LP 425 (Barrier
filter). A Neofluar 16X/0.5 objective and 12X oculars were used. Photographs
weretaken with a built-onZeissM35 camera.
Fluorescence microscopyisbased on secondaryfluorescence: theselectiveup-
take of fluorescent substances by certain parts of the microscopic preparation
(KHO & BAËR 1968).Callose,a substance present inthewallsofthepollen tubes
but absentinthesurrounding tissue,takesupanilineselectively and consequently
fluoresces when illuminated by blue or ultraviolet light. A mercury vapour lamp
supplies short-wave radiation. Filters between light sourceand microscope select
the desired light waves, which illuminate the preparation from above. A filter
between objective and ocularsabsorbstheshort wavesagain,onlythelongwaves
of the fluorescent parts of the tissuebeing transmitted. The pollen tubes are now
visibleinbright yellow-green, contrasting withadark background.
OBSERVATIONS AND CONCLUSIONS: The technique described worked very well

with Nerium oleander.In all preparations the pollen tubes were clearly percept-
ible.
The receptive stigmatic area evidently is located on the circular depression,
right below the collar on the lower part of the pistil head, as shown in Fig.
lc. and in the Photos 11 and 12. Pollen grains will germinate on other parts
of the moist surface of the pistil head as well, but the pollen tubes will grow
along this surface without penetrating it. Penetration is exclusively possible in
theindicated area (Fig. l c ) .
Inmostcasesthepollentubescould befollowed from thepointof penetration
in the stigmatic area through the style until they entered the ovules, as shown
in thePhotos 12-14.
Nerium oleanderappears to be self-compatible. After self-pollination no dis-
turbance ofpollen tube development has been observed. The pollen tubes reach
theovaryandentertheovules.However, innatureself-pollination rarelyoccurs,
asisdescribed inthenext chapter.
All cross-pollinations between cultivars appeared to be compatible. In every
combination ofcultivarstested,thepollentubesreachedtheovuleswithout diffi-
culty.
POLLINATION MECHANISM
The Nerium oleanderflower has an intricate construction (Fig. 2), connected

with arefined pollination mechanism.
Dark, usually carmine red, longitudinal lines in the corolla throat point to
the nectaries around the ovary at the base of the funnel-shaped corolla tube.
However, much ofthepassage isblocked bythecorona lobes and bythe woolly
plug formed by the intertwisted appendages of the anthers. The short-stalked
stamens are inserted at the middle of the corolla tube. The anthers have firm
sagittateplatesoutsideand areintrorse,adheringtothepistilhead,thus forming
a cone in the center of the flower. The pollen is shed inside the cone of anthers
and collects on the top of thepistil head, completely isolated from the stigmatic
areabelowthecollar at thelowerpart of thepistil head.
Between the five triangular anthers are chinks, narrowing to the top of the
cone. The insect's tongue can be inserted in the tube through these chinks or
through the openings between the filaments. In either case, when the tongue
iswithdrawn itgetsjammed betweenadjacent anthers.Apowerful pullisnecess-
arytoreleasethetongueand inthisactionthetonguemovesalongthecylindrical
part of the pistil head, where it gets covered with adhesive, and then through
the pollen chamber, where pollen grains are glued to the tongue. The insect
isprobably disturbed and leavesthe flower. In another oleander flower the pro-
cedure is repeated and when the tongue iswithdrawn it brushes past the collar
hangingdown from thepistilhead.Thus,thepollenisdeposited belowthecollar,
exactly on the receptive stigmatic area. Movingupward thetonguepasses again
along the cylindrical part of the pistil head and through the pollen chamber.
The tongue is covered again with pollen and the insect is ready to pollinate
another oleander flower.
The Nerium oleanderflower isadapted to pollination byLepidoptera (butter-
flies and moths). Only these insects have a tongue long enough to reach the
bottom ofthecorollatubeandhavethestrengthandenduranceneededtorelease
their tongue from thetrap and establish pollination of the oleander flower. The
oleander hawkmoth {Deilephilanerii L., Sphingidae), the larvae of which feed
on oleander leaves, is often mentioned in this respect but it is definitely not
theonlyoleanderpollinator and theinsectalsopollinatesotherflowering plants.
Diptera andHymenoptera often arefound trapped inoleander flowers. Trying
to collect nectar, their mouthparts become lodged between adjacent anthers or
getentangled inthehairyappendages oftheanthers.Theseinsectscannot pollin-
ateoleander flowers.
© frank pagen
FIG. 2. The flower of Nerium oleander L. 1.corolla from above x 3/4; 2. lateral view x 3/4; 3.
lengthwise section of the flower tube, 2 anthers removed x 6; 4. cone of anthers x 7; 5. position
of anther and pistil head x 7; 6. anther, adaxial view x 7; 7. id., abaxial view x 7 (drawn from
livingplants).

APOISONOUS PLANT
Allpartsoftheoleander contain toxicactiveprinciples.Theplantisextremely

poisonous to livestock and other animals, but the shrub is generally left un-
touched by them. However, there are records of animals having been killed by
eating a few oleander leaves. THEOPHRASTUS mentions the poisoning of pack-
animals during Alexander The Great's campaign (334-323 B.C.). DIOSCORIDES
states that oleander leaves and flowers willkill dogs, donkeys, mules and other
four-footed animals. Sheepand goats arekilledevenbydrinkingwaterinwhich
oleander leaves are steeped, according to PLINY (Historia Naturalis 24. 90). In
severalvernacular names theoleander isreferred to asa 'donkey-killer'. In Aus-
tralia six cows out of a dairy herd of seventeen died: they were fed with a load
of grass taken from a lawn near which oleanders were growing. Some leaves
weremixedwiththegrass (Gardeners' Chronicle 1878).The NewYork Tribune
reports that a healthy mare who ate a single tuft of leaves from a branch of
an oleander had partially lost control of her hind limbs the next morning, and
diedbefore assistancecouldbeobtained (Gardener's Chronicle 1881).Aveterin-
arylaboratoryintheDutchEastIndiesreceivedin 1921 thereportthat buffaloes
had died within afew hours after eating some oleander leaves (HEYNE 1927).
PLINY mentioned that honey made by bees of the nectar of oleander flowers
haspoisonousproperties (op. cit. 21. 77). ThesameisstatedbyTHEOBALD(1883
exPERRY 1980)inreference tooleandersin Burma.
Many cases of oleander poisoning of human beings have been reported. The
symptoms are depression, dizziness,fever, nausea, bloody diarrhoea, irregular-
ity of heartbeat, weakened pulse, paralysis, loss of consciousness, and death
through heartfailure. In 1796,someFrench soldiersinCorsicadied immediately
after eating meat skewered on an oleander branch, and during thewar in North
Africa, the majority of a group of soldiers who slept on pallets made from
oleander branches either died or were seriously poisoned, according to Lo-
iseleur-Deslongchamps. The Annals of the Peninsular War states that in 1808
in Spain French soldierscut some oleander branches and, having stripped them
of the bark, used the branches as skewers. Out of twelve who ate of the roast
seven died, and the rest were dangerously ill (Gardeners' Chronicle 1880). In
a supplement to a herbarium specimen from South Africa (Capetown (cult.)
Burtt Davy III. 1847 (K)), a record is given of the death of a four-months-old
baby after drinking milk from acow that had been eating oleander leaves.Chil-
drenhad beenpickingoleander flowers andhad thrown someoftheleavescare-
lessly into the hay. In California a boy died from the poison in an oleander
branch that hehad usedtoskewerhotdogsoveracampfire (Time,Mar. 1.1976).
Oleander rootshavebeenusedincasesofsuicidein India.
Seealso thePhytochemistry chapter.

M E D I C I N A L USES
Nerium oleanderisknown as a medicinal plant since ancient times. DIOSCOR-

IDESstatesthat thepoison oftheoleander, takenwithwineand rue (Rutagraveo-
lens), actsasan antivenin for snake bites.
In pharmacy the fresh or dried leaves,an infusion, decoction, plaster or salve
made from the leaves, the powdered bark or a decoction of the bark, a paste
made from the roots, and the dried flowers or a tallow cataplasm ofthe flowers
are used.
Internally the drug isused with caution as a cardiac stimulant with an action
comparable to Digitalis, to provoke menstruation, as an abortivum, as an anti-
spasmodic in the treatment of angina pectoris, to break the opium-smoking ha-
bit, and to build updeclining strength.
As an external medicine it isused against allkinds of skin diseases:rash, sca-
bies, ringworm, lice, leprosy and boils, to treat skin eruptions or irritations in
herpes, and to destroy maggots in wounds. The powdered leaves are used as
a sneezing-powder.
Nerium oleander has also been used in the treatment of cancer: the flowers,
leaves, leafjuice or latex, bark, and roots have been used against corns, warts,
cancerousulcers,hardapostemes,carcinoma,epithelioma, andindurated, ulcer-
ating orhard tumors (HARTWELL 1967).
Seealso thePhytochemistry chapter.
O T H E R USES
The latex or sap isused as an arrow poison, the bark as a rodenticide. Dried
oleander leaves are laid between papers and documents to keep insects away,
and are also used toexpel fleas. The flowers are among those chosen by Hindus
to offer to the God Siva. In India, Italy and Greece the oleander is associated
with and used at funerals. The oleander was among the flowers used by the
Romans in making garlands. The flowers are used for perfume. It is the floral
emblem of Saint Joseph, and the oleander is popular for making temporary
shelters used in the observance of theJewish Feast of theTabernacles. Adenium
spp.canbegrafted on aNerium oleanderrootstock toproduceprofusely flower-
ingplants.
Above all, Nerium oleander is appreciated as an ornamental plant, as will
bediscussed inPart three:The oleander cultivars.

PART TWO
PHYTOCHEMISTRY OF N E R I U M L.
N. G. BISSET
Pharmacognosy Research Laboratories, Department of Pharmacy,

King's College (KQC), ChelseaCampus, Universityof London,
Manresa Road,London SW3 6LX, U.K.
Chemical studies on the oleander have been reported under the three names
N. oleander L., N. indicum Miller, and N. odorum Sol. Here, only an outline
of the findings can be given, and the references to the literature should be con-
sulted for further details.

CARDIAC GLYCOSIDES
Extensive research hasshown that theplant contains a complex mixture of

cardenolide glycosides, with thequantitatively most important ones being der-
ived from thefollowing four aglycones: oleandrigenin (2a), digitoxigenin( l a ) ,
adynerigenin (3a), a n doleagenin (4a). The sugars attached directly t otheagly-
cones are mostly D-diginose (5),D-digitalose (6),and L-oleandrose (7), a n d
the di-a n d tri-glycosides contain additionally oneor twoD-glucose residues.
See: JÄGER et al. (1959), JANIAK et al. (1963), YAMAUCHI et al. (1975, 1983),
a n d TITTEL a n d W A G N E R (1981).
YAMAUCHI et al. (1976b) found that uzarigenin (lb) glycosides predominated
in theroot bark, thé*main component being odoroside B (lh). In the mixture
of glycosides from thestem bark, on the other hand, digitoxigenin glycosides
were the chief representatives, there being roughly equal amounts of odorosides
A (le), H (le), and G (lg), aswell asofodoroside B(lh).
It is the leaves that have received the most attention, because they arethe
source ofthe glycoside oleandrin, which haslong been inuse clinically (see be-
low). T h emixture of glycosides obtained depends very much o ntheorigin of
the material investigated andon its treatment prior to extraction. Early work
on industrial residues led t othe isolation of four glycosides: the main one olean-
drin (2k), anddeacetyl-oleandrin (2e), adynerin (3c), andneriantin (see: JÄGER
et al., 1959). Subsequent chromatographic studies o n Dalmatian material, by
TURCOVIC (1959c), showed the presence of many more glycosides. The principal
triglycosides, present in approximately equal amounts, were the digitoxigenin
and uzarigenin derivatives odorotrioside G (lg) and K (li), respectively; the
main diglycoside was the gitoxigenin derivative digitalinum verum (2d);a n dthe
major monoglycoside was again the oleandrigenin glycoside oleandrin (2k).
YAMAUCHIetal. (1975)inamore detailed study of air-dried leaves of Japanese
origin obtained the triglycoside corresponding tooleandrin (21,m)and the digly-
coside corresponding t onerigoside (2g) asthe chief glycosides present. I n addi-
tion, the triglycosides corresponding to odoroside A (li),adynerin (3d), a n d
A 1 6 -dehydro-adynerin (3h) were also isolated, aswell asthediglycosides 16-0-
Digitoxigenin,R = H,R t = 5ß-H

Uzarigenin,R = H,R, = 5oc-H
Odoroside A, R = D-Diginose,R, =5ß-H
OdorotriosideA, R = D-Diginose-Gentiobiose, R, = 5ß-H
Odoroside H, R = D-Digitalose,R t = 5ß-H
OdorobiosideG,R = D-Digitalose-D-Glucose,R, = 5ß-H
OdorotriosideG, R = D-Digitalose-Gentiobiose, Rl = 5ß-H
Odoroside B,R = D-Diginose, Rl = 5a H
OdorotriosideK,R = D-Diginose-Gentiobiose,Ki = 5a-H

2a Oleandrigenin, R = H, R^ = CH 3 CO
2b Gitoxigenin, R = R t = H
2c Strospeside, R = D-Digitalose, R t = H
2d Digitalinum verum, R = D-Digitalose-D-Glucose, R, = H
2e Deacetyl-oleandrin, R = L-Oleandrose, R l = H
2f Nerigoside, R = D-Diginose, Kt = CH 3 CO
2g Glucosylnerigoside, R = D-Diginose-D-Glucose, R! = CH 3 CO
2h Neritaloside, R = D-Digitalose, R t = CH 3 CO
2i Glucosyloleandrigenin (= Oleandrigenin ß-D-glucoside),R = D-Glucose,R[ = CH 3 CO
2j Glucosylneritaloside ( = 16-O-Acetyl-digitalinum verum) R = D-Digitalose-D-Glucose,
Ri = CH 3 CO
2k Oleandrin, R = L-Oleandrose, ^ = CH 3 CO
21 Glucosyloleandrin, R = L-Oleandrose-D-Glucose, R t = CH 3 CO
2m Gentiobiosyloleandrin, R = L-Oleandrose-Gentiobiose, Rj = CH 3 CO
3a Adynerigenin, R = H, R, = 5ß-H
3b A16-Adynerigenin, R = H, R, = 5ß-H, 16,17-double bond
3c Adynerin, R = D-Diginose, R, = 5ß-H
3d Gentiobiosyladynerin, R = D-Diginose-Gentiobiose, Rj = 5ß-H
3e 5a-Adynerin,R = D-Diginose, R t = 5a-H
3f A 16 -Dehydro-adynerin, R = D-Diginose, R t = 5ß-H, 16, 17-double bond
3g Glucosyl-A16-dehydro-adynerin, R = D-Diginose-D-Glucose, Rj = 5ß-H, 16,17-double
bond
3h Gentiobiosyl-A16-dehydro-adynerin, R = D-Diginose-Gentiobiose, Rj = 5ß-H, 16,17-dou-
blebond
acetyldigitalinumverum(2j)andA16-dehydro-adynerigeninD-glucosyl-ß-D-di-
gitaloside (3g) and the monoglycosides oleandrigenin ß-D-glucoside (2i) and
odoroside H (le). Strangely enough, while the triglycoside corresponding to
oleandrin is present, the nerigoside congener isabsent; and the reverse is the
4a Oleagenin, R = H
4b OleasideA, R = D-Diginose
4c Oleaside E, R = D-Diginose-Gentiobiose
CH3
H o i — OOM
OH OCH3
6 D-Digitalose 7 L-Oleandrose
5 D-Diginose
case with the corresponding diglycosides. YAMAUCHI et al. (1973) consider the
A 16 -compounds,which in the past have usually been regarded as artefacts aris-
ingduring theextraction process,to begenuineconstituents oftheplant mater-
ial.
YAMAUCHI and EHARA (1972) and YAMAUCHI et al. (1973) observed, on the
other hand, that leaf material oven-dried at a temperature > 80°C yielded a
preponderance of simpler glycosides. Among the more important ones were
oleandrin (2k)(0.13%),odoroside A (lc) (0.03%), adynerin (3c)(0.035%), and
A16-dehydro-adynerin(3f). ABEand YAMAUCHI(1978)isolated 5a-adynerin(3e)
as one of the minor constituents of the mixture. It is the first glycoside with
an aglycone having a trans-A/B ring junction to be obtained from the leaves;
nouzarigenin derivativeshaveyet been detected in them.
ABE and YAMAUCHI (1979) also obtained small amounts of a new series of
new glycosides, among them oleaside A (4b)and E (4c),based on the abnormal
aglycone oleagenin (4a). The fact that material containing a large amount of
oleaside A had very little adynerin, and vice versa, suggests that the two types
of glycoside are related biogenetically, and ABE and YAMAUCHI have been able
to realizethechemicalconversion ofadynerigenin acetate to oleagenin acetate.
On thin-layer chromatographic examination ofextracts from fresh latex and
leaves ofthree Indian varieties of oleander, thosehaving single and double pink
flowers and single white flowers, D E and SHARMA (1980) noted qualitative and
quantitativedifferences intheircardenolidecomposition.Thevarietywithsingle
pink flowers appeared to have the greatest oleandrin content (0.12%) and the
variety with singlewhite flowers theleast (0.06%);theproportions of oleandrin
in the total glycoside mixtures differed less and ranged between ca. 16% and
26%.
TITTEL and WAGNER (1981)have reported on the qualitative and quantitative
high-pressure liquid chromatographic (HPLC) analysis of commercial Nerium
leaf samples of different origins. They found that refluxing the leaf material
Phot. 4
Phot. 5 Phot. 6
PHOT. 3.Sibthorp & Smith. 1819.Flora Graeca. 3t. 248.Nerium oleander. Painted by Ferdinand Lukas
Bauer.
PHOT. 4-6. Oleander cultivars.4.'Magaly'; 5.'Marie Gambetta';6. 'Virginie'.
Phot. 7 Phot.8
Phot.9 Phot. 10
PHOT. 7-10. Oleander cultivars. 7.'Jannoch'; 8.'Soleil Levant'; 9.'Minouche'; 10.'Amboi'na'.

with methanol, with or without further purification, yielded extracts that were
sufficiently pure to be analysed directly. All the samples contained more than
1% glycosides.Carefully dried material contained verymuch higher concentra-
tions of the tri-and di-glycosidescorresponding to oleandrin (21,m)than of the
mono-glycosides such as oleandrin itself (2k) and odoroside A (lc). The olean-
drincontentvaried from 0.013%to0.12%,and innocasewasitthemainglyco-
side present. TITTEL and WAGNER did not detect any oleasides in the materials
they examined - a difference which may be due to local ecological conditions
ortotheexistence ofchemicalraceswithin thespecies.Sampleswith an elevated
monoglycoside content had only traces of higher glycosides; these findings, as
will be understood from the discussion above, must be due to differences in
thedryingand storageconditionstowhichthecommençaimaterial is subjected.
Thus, in addition to providing a quantitative analysis of the sample, the HPLC
analysis furnishes a chromatographic fingerprint that can yield information
about theprevious treatment ofthe sample.
More recently, YAMAUCHI et al. (1983) have quantitatively analysed oven-
and shade-dried leaf samples of 20 horticultural and wild varieties of Nerium
from various parts of the world by a combination of column and thin-layer
chromatography. Again, in most of the oven-dried samples oleandrin (2k) was
themajor glycoside, beingpresent inamounts varying from 0.007% to 0.425%,
but occasionally oleaside A (4b), nerigoside (2f), or A16-adynerin (3f) was the
principal component. In the shade-dried samples, gentiobiosyloleandrin (2m)
dominated, with some samplescontaining over 0.6%.
The oven-dried samples could be divided into two groups, depending on
whether therewasmoreadynerin (3c)than oleasideA(4b)present, or vice versa;
similarly, in the shade-dried samples, either gentiobiosyladynerin (3d) or olea-
side E (4c)was present in greater amount. Most Pakistan, Turkish, Greek, and
Indian samples were found to belong to the adynerin group, but the fact that
in a few samples the oleaside content was greater suggests the presence of an
enzymeabletocatalysetheadynerin-oleaside transformation.
It wasconcluded that overall thereisnocorrelation between the morphology
- flowers: single, semi-double, or double; colour: white deepening through yel-
low and pink to red - and the chemistry of the plants. The horticultural strains
showed considerable variation in their oleandrin content, indicating perhaps
that it might be possible to raise the yield of this glycoside through a selective
breeding programme.
Accordingto TURCOVIC(1959c),examination oftheflowersby paper-chroma-
tography showed that theycontained amixture ofdigitoxigenin and gitoxigenin
glycosides,with odorobioside G (If) asthemain constituent.
The seeds likewise have a complicated mixture of glycosides in them. After
(incomplete) enzymatic hydrolysis of material from Israel, JÄGER et al. (1959)
were able to isolate the digitoxigenin derivatives odoroside A (lc) and H (le)
astheprincipal components;aconsiderable amount of theoleandrigenin glyco-
sidesneritaloside(2h)andnerigoside(2f)wasalsopresent,butonlyalittleolean-
drin (2k).Adynerin (3c)was absent. According to paper-chromatographic stu-
diesonunfermented seedsfrom DalmatiabyTURCOVIC(1959a),the triglycoside
fraction, quantitatively the most important one, consisted almost entirely of
the digitoxigenin glycoside odorotrioside G (lg); the diglycoside fraction was
dominated by digitalinum verum (2d), a derivative of gitoxigenin (2b);and the
monoglycosides included odoroside H (le) and strospeside (2c).Oleandrin was
not detected.
Interestingly, the composition of the glycoside mixture from the pericarp of
the follicles is quite different from that of the seeds. TURCOVIC (1959b) found
that odorotrioside K (li), a derivative of the aglycone uzarigenin (lb), the
5oc-isomerof digitoxigenin (la), was the main glycoside, while odorotrioside G
(lg) was present only in traces and there wasmore odoroside A (le) and B (lh)
than in the seeds. JANIAK et al.(1963) found the pericarp to be a relatively rich
source ofadynerin (3c).
Toxicology
That Nerium is toxic to man and animals was known to the ancients and the
plant was mentioned by THEOPHRASTUS, DIOSCORIDES, PLINY, and GALEN,
among others.An oleander winewas used against thebite of poisonous snakes,
butingeneral theplant wasonlyemployed externally.Thesymptoms of poison-
inginanimals include vomiting, diarrhoea, stupor, trembling, convulsions, and
paralysis. As little as 15-20 g of fresh leaf is the fatal dose for a horse and 1-5
gfor asheep.Inhuman beingsthesymptoms ofpoisoning arenausea, vomiting,
colic,dizziness,staggeringgait,dilatation ofthepupil,bloodydiarrhoea, cardiac
weakness,and death preceded bycoma. Thenectar oftheplant issaid to render
honeypoisonous (WATT and BREYER-BRANDWIJK, 1962).
In 1808, 8 soldiers who ate meat roasted on a skewer made from oleander
wood died, while 4 others became seriously ill. It is said that people who had
eaten hare stuffed with oleander leaves also died as a result (MADAUS, 1938).
The plant has also been used as an abortifacient and to commit suicide. It ap-
pearsinthestatisticsofthePoison Control Centres,butincaseswheretheleaves
or flowers have been ingested, it seems that serious symptoms have not been
registered. No doubt, the intense bitter taste of the cardenolides is an effective
deterrent, preventing excessive consumption of the plant, and the spontaneous
vomiting which often occurs limits the amount of poison absorbed (FROHNE
and PFÄNDER, 1984).
Under the Sanskrit name karavira, the plant ismentioned in the early Indian
medical treatises by Caraka, Susruta, and Vaghbata (SINGH and CHUNEKAR,
1972). The roots and root bark, and also the leaves and flowers, are the parts
used - mostly in external preparations - for treating scorpion and snake bites,
for dealingwith awidevariety of skin complaints, ulcers,haemorrhoids, fevers,
headaches,and for reducingswellings,etc. (NADKARNI, 1954).Asjiazhutao (chia
chut'ao), theleaveshaveaminorroleinChinesetraditional medicine;thereme-
dies are used both internally and externally for heart conditions, bruises and
swellings, ringworm, and resistant fungal infections, as well as an insecticide
(HUNAN, 1978).

Clinicaluse
Interest has centred round the monoglycoside oleandrin (2k),which was ori-
ginallymarketed underthename 'folinerin' (MADAUS, 1938; JANIAKetal.,1963).
Other early preparations such as 'cortenerin', 'foliandrin', and 'cornerin' are
brieflydiscussed byWATTand BREYER-BRANDWIJK(1962).Oleandrinisstill offi-
cial in the Russian pharmacopoeia (REYNOLDS, 1982). Adynerigenin and olea-
genin glycosidesarelargelydevoid of activity.
OTHER CONSTITUENTS
From the root bark and stem bark of Japanese material, YAMAUCHI et al.
(1972b)haveobtained severalpregnenolone glucosides (8a-d),known tobebio-
geneticprecursorsofcardenolides. YAMAUCHIetal. (1974)and ABEand YAMAU-
CHI (1976) have also isolated various 12-hydroxypregnanes containing a
4,6-dien-3-one system, including the ichthyotoxic neridienones A (9a) and B
(9b);similar compounds arepresent in Kibatalia (Paravallaris) microphylla(Pi-
tard) Woods.The occurrence of thesecompounds inNerium issomewhat unex-
pected, for 12-hydroxycardenolides are not otherwise known to occur in the
plant.
Other steroids isolated by YAMAUCHI et al. (1976a) are the intense yellow-
coloured neriumosides (lOa-e), which are present in the root bark. They are
di- and tri-glycosides of 3ß-hydroxy-5ß-carda-8,14,16,20(22)-tetraenolide with
orwithout an additional 21-hydroxyfunction; theextended system of conjugat-
ed double bonds is responsible for the colour of these substances. Another un-
usual steroid derivative, neriaside (11), has been obtained by YAMAUCHI and
ÇH 3
co
ÎXy
/
i IH 1
R2OCH2 0 \ / N i /
A O
«
OR,
8 Pregn-5-en-3ß-ol-20-one ( = Pregnenolone), R = Rj = H
R = H, Ri = D-Glucose
R = D-Glucose, ^ = H
R = Rj = D-Glucose

ABE (1978)asa minor constituent of the leaves;itisthe D-diginoside of a novel
8,14-.s£C0-cardenolide.The corresponding triglycoside is also present and both
substances appear to begenuineconstituents ofthe plant.
The triterpenoid oleanolic (12a) and ursolic (12b) acids have been extracted
from theflowers, leaves,latex, and fruit pericarps (TURCOVIC, 1959b,c;cf. HEG-
NAUER, 1964). The coumarin derivative scopoletin (13a) and its ß-D-glucoside,
scopolin (13b), have been obtained from the stem bark and fruit pericarp. The
leaves have yielded as much as 0.3-0.4% of the branched-chain pentose apiose
(14) (DUFF and KNIGHT, 1963); the leaves and bark also contain the cyclitol
9a Neridienone A 9b Neridienone B
Neriumoside A-l Ri = D-Digitalose-Gentiobiose, R 2 = OH

Neriumsoide A-2 Ri = D-Digitalose-D-Glucose, R 2 = OH
Neriumoside B-l, Ri •• D-Digitalose-Gentiobiose,R 2 = H
Neriumoside B-2, Ri •• D-Digitalose-D-Glucose, R 2 = H
Neriumoside C-l RJ ; D-Diginose-Gentiobiose, R 2 = H
11 Neriaside

J W V \ A J
'COOH
12a Oleanolic acid

12b Ursolic acid
oô
CHO
13a Scopoletin, R = H 14 Apiose I
CHOH
13b Scopolin, R = D-Glucose I
CHOH
HOHjC^CÔH
OH
15 Dambonitol 16a 2-Hydroxy-acetophenone, R = H

16b 2,4-Dihydroxy-acetophenone, R = OH
OvÔ
Me02C
17 Plumericin
dambonitol (15) (see: HEGNAUER, 1964; NISHIBE et al., 1971). From the root
bark and heart-wood, YAMAUCHI et al. (1972a) have separated very small
amounts of4-hydroxy- and 2,4-dihydroxy-acetophenone (16a,b),atypeofcom-
pound also found inApocynum and Trachomitum species(cf. HEGNAUER, 1964).
The antibiotic plumericin (17), which, as its name indicates, was first isolated
from a Plumeria species,occurs in Nerium as such and in glycosidicform (BASU
and CHATTERJEE, 1973).
Rutin (18a) and nicotiflorin (18b) have been identified in the mixture of fla-
vonoid glycosides present in the leaves (1-3%, depending on the time ofcollec-
tion). The phenolic caffeic, ^-coumaric, and chlorogenic acids (19a-c) and syr-
ingic acid (20) have also been detected in the leaves (DURET and PARIS, 1972;
PARIS and DURET, 1974).
The plant contains about 17% fat in its seeds. Other constituents reported
18a Rutin, R = L-Rhamnose-D-Glucose, Rj = OH
18b Nicotiflorin, L-Rhamnose-D-Glucose, Rr = H
CH=CH—COR o
19a Caffeic acid, R = R 2 = R 3 = OH, Rt = H 1— \COOH
19b p-Coumaricacid, R = R 2 = OH, Rj = R 3 = H ^ \
^Si 19c Chlorogenic acid, R = Quinyl ( = H o \ _ _
Au),
R, = R 3 = OH, R 2 = H OH
R2
COOH
N JL 20 Syringicacid
CH,0'^^ÔCH3
OH
to be present are ascorbic acid and traces of rubber and volatile oil (WATT and
BREYER-BRANDWIJK, 1962).Reports indicating thepresenceoftracesof alkaloid
in the plant may be based on false positive alkaloid tests caused by the cardiac
glycosidespresent (BISSET, 1961).
CONCLUSION
Theoutstanding phytochemicalfeature oftheplant is, ofcourse,the presence

of the cardenolide glycosides. It is a character shared by several other genera
of the sub-family Apocynoideae, e.g. Vallaris, Beaumontia, Strophanthus, etc.,
but unique to Nerium isthecombination of aglycones- principally digitoxigen-
in, uzarigenin, gitoxigenin, and oleandrigenin* - and specific sugars - mainly
D-diginose and D-digitalose. The isolation of the novel oleasides and neriumo-
*Very recently, SIDDIQUI et al. (1987) have isolated two new cardiac glycosides, kaneroside and
neriumoside(nottobeconfused withtheneriumosideslOa-eobtainedbyYAMAUCHIetal.(1976a)),
from fresh undried winter leaves from Pakistan, which are formulated as the ß-D-diginosides of
cardenolide aglyconescontaining a2a-hydroxy function.

sides, aswell as of the œco-cardenolide neriaside, raises the question whether
theyreallyarepeculiartoNeriumorwhetherre-examinationofthecardenolide
mixtures present in related genera might bring to light additional substances
belonging to these groups. The occurrence of the neridienones A and B,pre-
gnane derivatives with a somewhat unusual 4,6-dien-3-one system, may point
toaconnection withthegenera oftheApocynoideae that contain steroid alka-
loids,inparticularKibatalia.
REFERENCES
F.Abeand T.Yamauchi(1976),Phytochemistry 15,1745-1748.

F.Abeand T.Yamauchi (1978),Chem. Pharm. Bull. 26,3023-3027.
F.Abeand T.Yamauchi (1979),Chem. Pharm. Bull. 27,1604-1610.
D. Basu and A.Chatterjee (1973),IndianJ. Chem. 11,297.
N. G. Bisset(1961),Annales Bogorienses4,65-144,72.
M. Deand A. K. Sharma(1980),Nucleus (Calcutta), 23,213-225.
R. B.Duff and A.H. Knight (1963),Biochem. J. 88,33P-34P.
S.Duret and R. R. Paris(1972),PI. Méd. Phytothérap. 6,210-215.
D. Frohne and H. J. Pfander (1984),A colouratlas ofpoisonousplants, Wolfe Publishing, London,
pp. 47^*8.
R. Hegnauer (1964), Chemotaxonomie der Pflanzen, Birkhäuser, Basel - Stuttgart, vol. 3,pp. 146,
149,152,153,157.
Hunan Province, Revolutionary Health Committee (1978),A barefoot doctor's manual, Routledge
&Kegan Paul, London - Henley,p.245.
H.Jäger, O.Schindler, and T. Reichstein(1959),Helv. Chim.Acta 42,977-1013.
P. St.Janiak, Ek. Weiss,J.v.Euw,and T. Reichstein(1963),Heb. Chim.Acta 46,374-392.
G. Madaus (1938),Lehrbuch derbiologischenHeilmittel, G. Thieme,Leipzig,pp. 2010-2015.
A.K.Nadkarni(1954),IndianMateria Medica, 3rded.,Popular Book Depot, Bombay,pp.847-849.
S.Nishibe,S.Hisada, and I. Inagaki (1971),Phytochemistry 10,896.
R. R. Parisand S.Duret (1974),PI. Méd. Phytothérap. 8,318-325.
J. E.F. Reynolds (1982),Ed., Martindale. Theextrapharmacopoeia, 28thed.,Pharmaceutical Press,
London, p.544.
S.Siddiqui, F.Hafeez, S.Begum, and B.S.Siddiqui (1987),Phytochemistry 26,237-241.
B. Singh and K. C. Chunekar (1972), Glossary of vegetable drugs in brhattrayl (= Chowkhamba
Sanskrit Studies vol. 87), Varanasi,p.77.
G. Titteland H.Wagner (1981),Planta Medica 43,252-260.
I.Turcovic(1959a),J. Pharm. Belg. [n.s.] 14,263-275.
I.Turcovic(1959b),J. Pharm. Belg. [n.s.]14,376-385.
I.Turcovic(1959c),J. Pharm. Belg. [n.s.] 14,447-455.
J. M. Watt and M. G. Breyer-Brandwijk (1962), The medical andpoisonousplants of southern and
easternAfrica, E.&S.Livingstone, Edinburgh - London, pp. 88-93.
T.Yamauchi and F.Abe (1978), Tetrahedron Lett. no.21,pp. 1825-1828.
T.Yamauchi and Y. Ehara (1972), Yakugaku Zasshi 92,155-157.
T.Yamauchi, M.Hara, and Y. Ehara (1972a),Phytochemistry 11, 1852-1853.
T.Yamauchi, M.Hara, and K. Mihashi (1972b),Phytochemistry 11, 3345-3347.
T.Yamauchi,Y. Mori, and Y.Ogata (1973),Phytochemistry 12,2737-2739.
T.Yamauchi,N.Takata, and T. Mimura (1975),Phytochemistry 14,1379-1382.
T.Yamauchi, F.Abe,and M.Takahashi (1976a), Tetrahedron Lett. no. 14,pp. 1115-1116.
T.Yamauchi, M. Takahashi, and F.Abe (1976b),Phytochemistry 15,1275-1278.

T.Yamauchi, F.Abe,Y.Ogata, and M.Takahashi (1974),Chem. Pharm. Bull. 22,1680-1681.
T.Yamauchi, F.Abe,Y.Tachibana, C.K. Atal, B.M. Sharma, and Z. Imre(1983) Phytochemistry
22,2211-2214.
38 Agric. Univ. WageningenPapers87-2 (1987')

PART THREE
THE OLEANDER CULTIVARS
TENTATIVE CHECKLIST OF O L E A N D E R CULTIVARS
There is no special international registration authority assigned to Nerium

yet. However, international registration would be highly desirable. Oleanders
areverypopular ornamental plantsandcommonlycultivated inalltropical and
subtropical countries as a garden shrub and elsewhere as a pot plant. Many
oleander cultivars are available and new cultivars are added from time to time,
but the naming of oleander cultivars leaves much to be desired. Nomenclature
ofoldercultivarsavailableinthetradetoday isunstable,not uniform, and often
inaccurate. In many cases newly released cultivars are given names that are
contrarytorecommendations orarticlesoftheInternational CodeofNomencla-
ture for Cultivated Plants (ICNCP).
To establish a basis, an inventory of oleander cultivars has been compiled
bythepresentauthor, resultingintheTentativeChecklistofOleander Cultivars.
An important source of information for the inventory has been the unique
andlargecataloguecollection oftheGovernment Institutefor Research on Var-
ietiesofCultivated Plants(R.I.V.R.O.) inWageningen.Thiscollection oforigin-
al nursery and seed catalogues from all over the world includes many recent
catalogues as well as old ones from the nineteenth century, thus representing
an invaluable source ofinformation on cultivar namesand descriptions.
Especially useful was the discovery of a set of catalogues, published in the
second halfofthenineteenth century, ofClaudeSahut'sNurseryin Montpellier,
France,for Sahut hasdeveloped alargenumber oftheoleander cultivars.Addi-
tionalinformation wasfound inotherkindsofliterature,orobtained byperson-
alcommunication. Allinformation has been entered into a data base computer
file.Thechecklist wasgenerated usingthiscomputer file.
A total of 599 different cultivar names, including 157 misspellings and 41
synonyms, were encountered in the inventory, representing 401 distinct culti-
vars.Of these401, about 175areavailable inthetrade today.
Oleander cultivars vary inform, colour and scent oftheflowers, and in habit.
Severalcultivars havevariegated leaves.
FLOWER FORMS:With regard to theflower form threetypesofoleander flowers

can bedistinguished: single,superposed corollas,and double (Fig.3).
I. SINGLE: This flower form consists of a calyx of five sepals, a corolla tube,
five corolla lobes and a corona, asdescribed for the species. Every corolla lobe
is asymmetrical and can be divided in two areas (see Fig. 3: lb). A small strip
(withadarker colour underneath) stretching outward, endinginapointed apex,
and a much broader area stretching out to the right and beyond the pointed
3a
3c
3b
© frank pagen
FIG. 3. Flower forms of oleander cultivars. 1. single; 2. superposed corollas; 3. double, (a. bud;
b.viewfrom above;c.lateralview).

apex.In somecultivars both areas are broad and thecorolla lobeisnearly sym-
metrical.The flower mayvaryconsiderably insize.
ii. SUPERPOSED COROLLAS:The term hose-in-hose can also be used to describe
this form. It features a calyx and two similar corollas with one corolla tube
superimposed insidetheotherand withthecorolla lobesofthetwowhorls alter-
nating. In the outer tube, stamens are absent. Sometimes five slits are visible
in the lower part of the outer tube. A few cultivars feature three superposed
corollas,withstamens present intheinner tube only.
in. DOUBLE: The term "double" in describing a flower form is ambiguous,
but in this classification it is defined by the following characters (not all char-
actersmay bepresent inthesame flower):
- More than ten petalsorpetaloid parts.
- Irregularity.
- Somepetalsfree, some fused.
- Petalsdiffering insize.
- Petalswithclaws.
- Petaloid anthers.
- Broad, swollen buds.
The double oleander flower has an irregular structure. Although the flower
basically is composed of two, three or even four whorls of petals in addition
tothecalyx,thesystemisusuallydistorted and often hard to unravel.
A whorl usually consists of five petals, but whorls of four or six may occur.
The petals of the inner whorl are generally fused to a short tube; some or all
of the anthers may be partly or completely petaloid. The petals in the other
whorls are all provided with a long claw, and may differ widely in size; they
maybefree ormoreorlessunited at their basesingroupsoftwoto four.
Not only the structure of a separate flower isirregular; there isalso consider-
ablevariation in flower structure among theflowers of one plant.
In some cases the flower type with superposed corollas and the double type
may be confused. Several double-flowered cultivars produce flowers basically
with two whorls of petals, in which the petals of the outer whorl are more or
lessfused, and hardly any anther ispetaloid. On the other hand plants of culti-
varswithsuperposed corollasmayproduceoccasionally aflowerwithapetaloid
anther or with another irregularity. However, checking of more flowers on the
sameplant usuallyleadsto aclear identification.
Thecorona,presentinallthreekindsofflowers,alsovariesinform inoleander
cultivars.Twobasictypescanbeclearlyrecognized:atypewithcorona segments
divided in three (sometimes four) lobes, one short and triangular, with a linear
lobeoneitherside(sometimestwoononeside);and atypewithcorona segments
finely divided, usually irregularly, in four to ten long filiform lobes, the two
types originating from the oleander of the Mediterranean, and of the Indian
kind, respectively. However, some intermediates between the two basic corona
typesoccur aswell.

COLOUR: Flowers of oleander cultivars range in colour from pure white
through cream to yellow, apricot and salmon and from pink (in many shades)
to red,carmine redand purple.
The colour is very much influenced by environmental factors, especially by
light intensity, and can only bejudged properly in flowers that have developed
infull sun.
In onecorolla lobeseveralshadesofonecolour maybepresent. Corolla lobes
may have a different colour beneath. The main colour sometimes is overlaid
with hues of another colour. The corolla lobes may be variegated with stripes
and spots of another (usually paler) colour. The edges of the corolla lobes may
have a different colour. The throat (the inside of the corolla tube) may have
a colour different from thecorolla lobes and often features stripesin a contrast-
ing colour.
The oleander cultivars may be grouped into categories according to flower
form and colour.
Thefollowing classification, usedinthechecklist, isan adaptation of thesys-
temusedby Sahut inhisNerium catalogues:
Superposed
Single corollas Double
Group 1. White 5. White 9. White

2. Pink 6. Pink 10. Pink
3. Red 7. Red 11. Red
4. Yellow 8. Yellow 12. Yellow
White White, cream.

Pink Pink, lilac.
Red Red,carmine,purple.
Yellow Yellow, salmon, apricot, flesh colour, copper, orange
SCENT: Several oleander cultivars have fragrant flowers. The scent may be
faint or powerful and has been described as reminiscent of vanilla, bitter al-
monds,jasmine,magnolia, peruvian heliotrope, or asan agreeablemusky scent.
The oleander of the Indian kind supposedly is the source of fragrance in the
cultivars.
Many cultivarshaveflowers with no scentat all.
HABITUS:Mostcultivarsarelargeandvigorousgrowersand canreacha height

exceeding 2.50 m; medium-sized oleanders are growing 1.25 m to 2.50 m., and
dwarf typesare0.50m to 1.25min height.
VARIEGATED LEAVES: Several oleander cultivars have foliage variegated with

yellow or white. The flowers are usually pink and may be single, double, or
with superposed corollas.
NOTES ON THE CHECKLIST: All cultivar names encountered in the inventory
are listed inalphabetical order in theTentative Checklist of Oleander Cultivars
(seeAppendix).
Ancient cultivar names in Latin are treated as fancy names, and listed as e.g.
'Atropurpureum', 'Carneum Plenum', 'Flore Albo',and 'Foliis Variegatis'.
Wrongly spelled names are listed without a description and with reference
to thecorrect name.
Synonyms are listed with information on thepublication only and with refer-
enceto thepreferable name.
Included inthedescription following acorrectcultivar nameare: encountered
synonyms ofthis name;an indication of flower type,colour, scent,habitus, and
the number of thecategory in which the cultivar isplaced (seediagram on page
42); the name of the originator, if known, and the year of origin (the year in
which the cultivar wasput on the market isconsidered the year of origin here);
the earliest publication of the name encountered by the present author ('First
publication'), and its date. Additional information concerning flower size, in-
florescence, vigour, hardiness,etcetera, isgiven after 'Remarks'.
To somecultivarsplant breeders rightisapplicable and insomecasesatrade-
mark isattached toa cultivar.
HISTORY OF THE CULTIVATED OLEANDER
Nerium oleanderhasbeen incultivation asan ornamental sinceancient times.

In China the cultivation of oleanders was a hobby of literary men, who
adorned their studieswith cut oleander blooms.They especially appreciated the
scent of oleander flowers and the elegant habit of the plant, and chose the
oleander asan emblem ofgraceand beauty. To describe theoleander, the Chin-
ese use three characters, successively meaning 'mingle', 'bamboo', and 'peach
blossom'.
DIOSCORIDES(DeMateria Medica 4. 82)knewtheoleander asaplant growing
wild near theseaand along rivers,but alsoasacultivated plant in gardens.
Oleanders were already planted in Roman gardens in the days of Cicero
(106-43 B.C.). PLINY (Historia Naturalis 17.98) gave directions how to propa-
gate the plant by layering and by seed. In Pompeii the oleander is the plant
most frequently represented in the garden paintings on the walls of peristyle
gardens.Thesepaintings,datingfrom 79A.D.orearlier,andpreservedbyVesu-
vius,weredesigned bythePompeians tomaketheir smallgardens appear much
larger: through the painting the illusion was created that the garden extended
far into the countryside. In these garden paintings the oleander is usually pic-
tured in an informal setting, growing in masses and forming a background for
(painted) fountain-vases and statues. Excavations of villa gardens, e.g. of the
Villa at Torre Annunziata, and analysis of root cavities and remnants of
Ofcrtrityt&WttJ*
FIG. 4.Rembertus Dodonaeus. 1608.Cruydt-boeck, p. 1426:"Oleanderboom". Wood engraving.

^4.8 HORTUS ACADEMICUS LUGDUNO-BATAVUS. 44,9
JJSUVU INDICl/M AyGüSTIFOLWM. ILORIEVS Nniau MDicvH uTiioiwu,n,Qim5tu-
ODORATIS SlUellClBUS. rns ODOMtlS.
FIG. 5.Paul Hermann. 1687.Horti Academici Lugduno-Batavi Catalogus, t.448:"Nerium Indicum

angustifolium Acribus odoratis simplicibus" (left), t. 449: "Nerium Indicum latifolium, floribus
plenisodoratis" (right).Copper engraving.
branchesrevealedthatintheactualgardensoftheVesuvianareaoleanderswere
indeed planted in a similar fashion. Garden paintings featuring oleanders have
also been found elsewhere in Italy: in the garden room (now in the Terme Mu-
seum in Rome) of the Villa at Prima Porta, 14.5 kms from Rome, which was
built by Augustus for his wife Livia, and in the Auditorium of Maecenas in
Rome (JASHEMSKI 1979).
In the twelfth century the oleander was the only flowering shrub, together
with myrtl and rose, used by the Arab gardeners of the Dar-al-Islam (HYAMS
1971).
For a long time thecommon form of the Mediterranean oleander with single
odourless pink to red flowers (see Fig. 4 and Phot. 3)was the only one known
and cultivated in Europe. In 1547a form with single white flowers, discovered
in Crete near Camerachi on Mount Ida (SAVI 1813),was introduced into Italy
by PIER ANTONIO MICHIEL (BLUNT 1979). In 1560 CONRAD GESNER mentions
an oleander in cultivation in Basel. The oleander was introduced into Florida
in 1565 by the earliest Spanish settlers at St. Augustine. In 1596 in England,
JOHN GERARD had ared and awhiteform ofthe oleander.
In 1683the scented Indian oleander (see Figs. 5-7) was introduced into Eur-
ope, the form with single light pink flowers by VAN RHEEDE TOT DRAKESTEIN
from S.W. India; the form with double pink flowers by BEVERNINGK from Cey-
lon (HERMANN 1687, RHEEDE 1689, EDWARDS 1815, SWEET 1830). HERMANN
(1689)hadanIndian oleanderwithvariegated doubleflowersinLeiden; COMME-
LIN (1697) gives an illustration. WEINMANN (1748) shows a variegated-leaved
form of the Mediterranean single pink oleander, and an Indian oleander with
red double flowers. AITON (1789)mentions a double-flowered form both of the
Indian and of the Mediterranean oleander. A yellow single form of the Indian
oleander was named Nerium flavescens by Di Spmo in 1812 (ex ROEMER &
SCHULTES1819).
More and more cultivars appeared in the nineteenth century and the forms
of the Mediterranean and Indian oleander mingled. In 1840 BOSSE mentions
36cultivars;in 1849helists58.
CLAUDE SAHUT in Montpellier, France, produced the greater portion of the
oleander cultivars. In 1898 he could offer 170 cultivars. The flower type with
superposed corollas appeared in 1868inhis nursery.
Oleander cultivars also have been developed elsewhere in the Mediterranean
region,inAsia,and intheUnited States.
THE O L E A N D E R IN CULTIVATION TODAY
Today the oleander is commonly planted in gardens in all tropical and sub-
tropicalcountriesand grown asapot or tub plant inthetemperate zone.
Oleanders areverypopular because oftheir brilliantly coloured flowers, long
flowering season, evergreen foliage, adaptibility and easy cultivation. The
oleander toleratesawidevarietyofsoils,includingpoor and alkaline ones,with-
stands periods of drought, can survive short periods of frost to -12°C (10°F)
and isresistant tourban pollution.
In tropical and subtropical countries the oleander is planted outdoors. The
flowering evergreen shrubs are used in gardens and parks as hedges or screens,
ingroups,asbackground plantings or assolitary specimen plants.The oleander
is also used as a divider plant in the median strips of highways. Larger forms
canbetrained assingle-ormultiple-trunked trees.
Oleanders are planted in pots or tubs, grown as bush plants or trained to
crown standards, for decorating terraces and patios. In the temperate zone the
plantsarewinteredindoors.Intheseareastheoleanderisalsogrown asa green-
house specimen or asa house plant. In thecommercial production of oleanders
as flowering pot plants rooted cuttings are treated with growth retardants to
hasten flowering and toproducecompact plants (KARPER 1979).
Phot. 11
Phot. 12
PHOT. 11. Nerium oleander. Pollen tubes penetrating below the collar on the lower part of the pistil
head, asrevealed by fluorescence microscopy (x 50)(phot. J. van deVooren).
PHOT. 12. Nerium oleander. Pollen grains on the central cylindrical part of the pistil head germinate,
but do not penetrate (top). Penetration of pollen tubes is exclusively possible in the circular depression
below the collar on the lower part of the pistil head (center), as revealed by fluorescence microscopy
(x 50)(phot. J.van deVooren).
Phot. 13
Phot. 14
PHOT. 13. Nerium oleander. Pollen tubes growing down to the ovary through the style, as revealed by
fluorescence microscopy (x 50)(phot. J.van deVooren).
PHOT. 14. Nerium oleander. Pollen tubes have reached the ovary (center) through the style (top left)
and enter theovules,asrevealed byfluorescence microscopy (x 50)(phot. J. van deVooren).
FIG. 6. H. van Rheede tot Drakestein. 1689. Hortus Malabaricus, Vol. 9. t. 1: "Tsjovânna-areli"
Copper engraving.

^755-
der, l&tft>&nl» ïPaütn.

b.ZA/eriizmr&nDicum.2*z£L/oJntm-fb>re rmreo vlenf, 3lu>s,
je Qcrzj£/eßPof$rißify^'ty\lxSa^'&&nnt>tr.
FIG. 7.JohanWilhelmWeinmann. 1748.Duidelijke vertoning,EenigerDuizendinallevier Waerelds

Deelen wassende Bomen, Stammen, Kruiden, Bloemen, Vrugten, en Uitwassen, & c , t. 755: "a.
Nerium Indicum flore carneo,Oleander, Unholdenbaum. b.Nerium Indicum latifolium flore roseo
pleno, Rosagedouble,wohlriechend "gefüllter" Oleander". Copper engraving, hand-coloured.

ACKNOWLEDGEMENTS
The present author is greatly indebted to the Directors and Curators of the
herbaria cited for putting thematerial at hisdisposal:
AAU Aarhus, Denmark: Herbarium Jutlandicum, Botanical Institute, Un-
iversity ofAarhus.
BM London, Great Britain: British Museum (Natural History).
BP Budapest, Hungary: Museum of Natural History, Department of Bo-
tany.
BR Bruxelles,Belgium:Jardin Botanique del'Etat.
C Kobenhavn, Denmark: Botanical Museum and Herbarium.
E Edinburgh, Great Britain: Royal Botanic Garden.
FI Firenze, Italy:Herbarium Universitatis Florentinae, Istituto Botanico.
FI-W Firenze,Italy: Herbarium Webbianum.
G Genève,Switzerland:ConservatoireetJardin Botaniques.
K Kew, Great Britain:The Herbarium and Library.
L Leiden,Netherlands: Rijksherbarium.
MO Saint Louis,Missouri- U.S.A.:Missouri Botanical Garden.
NY New York, NewYork - U.S.A.:The NewYork Botanical Garden.
P Paris, France: Muséum National d'Histoire Naturelle, Laboratoire de
Phanerogamic
S Stockholm, Sweden: Botanical Department, Naturhistoriska Riksmu-
seet.
UPS Uppsala, Sweden: Institute of Systematic Botany, University of Uppsa-
la.
W Wien,Austria: Naturhistorisches Museum.
WAG Wageningen, Netherlands: Department of Plant Taxonomy, Wagen-
ingen Agricultural University.
Z Zürich, Switzerland: Botanischer Garten und Institut für Systematische
Botanik der Universität Zürich.
He is also very grateful for the help he received from Prof. Dr. Ir. L. J. G.
VAN DER MAESEN and from the staff of the Department of Plant Taxonomy
oftheWageningen Agricultural University.
He acknowledges especially Dr. A.J. M. LEEUWENBERG and Ir. W. A. BRAN-
DENBURG for their supervision, Mr. J. VANDEVOORENfor helping with the com-
puterprogram andforphotographs,and Mrs.N.WILDERSforadviceand photo-
graphs.
He is grateful to Prof. Dr. N. G. Bisset of the Department of Pharmacy of
theUniversity ofLondon for hisimportant contribution.
He also wants to express his gratitude to Mr. F. SCHNEIDER for comments
on the cultivar checklist, to Ir. C. J. BARENDRECHT, the Government Institute
for Research on Varieties of Cultivated Plants in Wageningen, Mr. B. VAN DE
LUSTGRAAF, Mr. R. MOELIKER and the Centre for Agricultural Publishing and
Documentation in Wageningen for help and information, to Mr. J. A. SMIT,
Mrs. S. G. SMIT-MIOLÉE and the Dutch Oleander Society for providing living
material and for information, toMr.J.J. KARPER and theDepartment of Horti-
culture of theWageningen Agricultural University for information and for pro-
vidingglasshouseroomfor theoleanders,toDr.A.C. LESLIEoftheRoyal Horti-
cultural Society's Garden in Wisley, Great Britain, to Mr. R. BRAND and Mr.
F. FERREROofI.N.R.A.-G.E.V.E.S., Cavaillon,and Mr.J.M. REYof Pépinières
Jean Rey, Jonquières, in France, to Mr. J. A. BAUML of the County of Los
Angeles Department of Arboreta and Botanic Gardens, Arcadia, California,
Mrs. A. TEASDALE of Monrovia Nursery Company, Azusa, California, Mrs. E.
S. HEADandtheNationalOleander Society,Galveston,Texas,andDr.J. POPEN-
OE of the Fairchild Tropical Garden, Miami, Florida, in the United States of
America, for information and references.
Finally, hewants to express hisgratitude for the grant received from the Wa-
geningen Agricultural University Fund, in order to visit the herbaria of the
Royal Botanic Gardens in Kew (K) and the British Museum (Natural History)
(BM)in London, Great Britain.

REFERENCES
AiTON, W. 1789. Hortus Kewensis, ora catalogue ofthe plants cultivated inthe Royal Botanic
Garden at Kew. London, GeorgeNicol. Ed. 1.1:297.
BAILEY,L.H.&E.Z. BAILEY. 1947.Hortus Second.NewYork: MacMillan Company.496.
BEUCHERT, M. 1983. DieGärten Chinas.Köln, Eugen Diederichs Verlag.51.
BLUNT, W. 1979.The Illustrated Herbal. London, Lincoln.101.
BOSSE, J.F. W. 1854.Vollständiges Handbuch der Blumengärtnerei. Hannover. Ed. 3.2:875-877.
(In German).
BROCE, A.B. &J. IDEKER. 1978. Oleander flowers asinsect traps. Ann. Entomol. Soc. Am. 71.
4:628-629.
COMMELIN,C. 1697. Horti Medici Amstelodamensis rariorum plantarum: descriptio et icônes. Pars
1:TamOrientalis,quamOccidentalisIndiae,aliarumqueperegrinarumplantarum. Amstelodami
(Amsterdam),P.etJ. Blaeu,Abrahamum aSomeren.4 5 ^ 6 , t. 23.(In Latin and Dutch).
CURTIS, W. 1816.Nerium odorum (ß),Double sweet-scented Rosebay, or Oleander. Curtis' Botani-
calMagazine 43: t. 1799.
CURTIS, W. 1818.Nerium odorum (y),Carneum. Curtis' Botanical Magazine 46: t.2032.
DAVIS, P.H.1978. Flora ofTurkey and the East Aegean Islands. Edinburgh, University Press.
6:159.
DIEPEN, H.VAN. 1980. Bacteriekanker bij Nerium oleander (Bacterial gall inNerium oleander).
Vakblad voor deBloemisterij 20:28-29. (In Dutch).
DODONAEUS, R. 1608. Cruydt-boeck, Leyden (Leiden), Francoys van Ravelingen. 1426-1427. (In
Dutch).
DRAPIEZ, P.A. J. 1835.Herbier de l'Amateur de Fleurs, avecun précis d'organographie et de phy-
siquevégétales servant d'introduction àl'ouvrage. Bruxelles.8:t.563,t.564.(In French).
DUMONT DE COURSET, G. L. M. 1811. Le Botaniste Cultivateur. Paris, Weterville & Goujon. Ed.
2.3:268. (In French).
DUTCH OLEANDER SOCIETY. 1981.Tafel der Variëteiten (List ofCultivars). Amsterdam. (In Dutch).
EDWARDS, S. 1815.Neriumodorum ß,Doublesweet-scented RosebayorOleander. Edwards' Botan-
icalRegister l:t.74.
FUCHS, L. 1543.NewKreuterbuch. Basell(Basel),Michael Isingrin.(In German).
GARDEN, THE. 1895.(Great Britain):384.
GARDENERS' CHRONICLE, THE. 1878.1:533.
GRAF, A. B. 1978. Tropica, colorcyclopedia ofexoticplantsand treesfrom thetropics and subtrop-
ics. East Rutherford (N.J.),Roehrs. 1039.
HARTWELL,J. L. 1967.Plantsusedagainstcancer, asurvey.Lloydia 30.4:407.
HERMANN,P. 1687.HortiAcademici LugduniBataviCatalogus.Lugduni Batavorum (Leiden), Cor-
nelium Boutesteyn. 447-450.(In Latin).
HERMANN,P. 1689.Paradisi Batavi Prodromus. Amsterdam. 356.(In Latin).
HERR, J. M. 1971.A newclearing-squash technique for the study of ovuledevelopment inAngiosp-
erms.American Journal ofBotany 58.8:785-790.
HEYNE, K. 1927. Denuttige planten vanNederlandsch Indië. 's Gravenhage (The Hague).2:
1291-1292.(In Dutch).
HYAMS, E. 1971. AHistory ofGardens and Gardening. London,J. M. Dent &Sons.84.
INTERNATIONAL COMMISSION FOR THE NOMENCLATURE OFCULTIVATED PLANTS OFTHEI.U.B.S. 1980.
International CodeofNomenclature for Cultivated Plants (I.C.N.C.P.). Regnum Vegetabile 104.
Utrecht, Bohn, Scheltema &Holtema; TheHague,W.Junk b.v.
JASHEMSKI,W.F. 1979. TheGardensofPompeii,Herculaneum andtheVillasdestroyed byVesuvius.
New Rochelle, Caratzas.
KARPER, J.J. 1979. Teelt Nerium oleander biedt mogelijkheden (The culture ofNerium oleander
hasgood prospects).Vakblad voordeBloemisterij 34.49:40-43. (In Dutch).

KERSTENS,A.H. 1878.Keur van Bloemen en Sierplanten, hare aankweeking, veredeling, vermenig-
vuldiging enbehandeling, 'sHertogenbosch, W.C.vanHeusden. 103-117.(InDutch).
KHO, Y. O. & J. BAËR. 1968.Observing pollen tubes by means of fluorescence. Euphytica 17:
298-302.
KIRTIKAR, K.R.&B.D. BASU. 1933.Indian medicinal plants. Allahabad, Lalit Mohan Basu.Ed.
2.2:1584-1586.
LEEUWENBERG, A.J. M. 1984. Series of revisions ofApocynaceae XIII. Notes on Nerium L.and
Tabernaemontana L.Mededelingen Landbouwhogeschool Wageningen 83.7: 57-60.
LINNAEUS, C. 1737.HortusCliffortianus. Amstelaedami (Amsterdam).76.(InLatin).
LINNAEUS,C. 1753. SpeciesPlantarum. Holmiae (Stockholm).Laurentius Salvius.1:209.(InLatin).
LINNAEUS, C. 1774. Systema Vegetabilium. Gottingae (Göttingen) et Gothae (Gotha), Jo.Christ.
Dietrich. Ed.13.(InLatin).
LODDIGES,C.&SONS. 1822.Loddiges' Botanical Cabinet 7: t.666,t.700.
LUDWIG, F. 1881. Zur Biologieder Apocyneen. BotanischesCentralblatt 8:183-188.(InGerman).
MCCLINTOCK, E.&A. T. LEISER. Received 1985.AnAnnotated Checklist ofWoody Ornamental
Plants ofCalifornia, Oregon and Washington.82.
MILLER,P. 1768.TheGardener's Dictionary. London,J.&F. Rivingtonetal. Ed. 8:2-3.
MON JARDIN & M A MAISON. 1984.12:128.(In French).
NATIONAL OLEANDER SOCIETY. Received 1985.Various promotional publications. Galveston,
Texas.
NICHOLSON,G. 1885.The Illustrated Dictonary ofGardening, apractical andscientific encyclopae-
diaofhorticulture forgardeners and botanists.London, Upcott Gill.447.
NICHOLSON, G.&S. MOTTET. 1895.Dictionnaire pratique d'horticulture etdejardinage. Paris,Li-
braireAgricole.3:465.(InFrench).
OAKMAN, H. 1975. Tropical and subtropical gardening. Queensland, Australia, Jacaranda Press.
206.
PAXTON,J. 1837.Nerium thyrsiflorum. Paxton's Magazine ofBotany 3:73.
PAXTON,J. 1844.Nerium oleander Tangle.Paxton's Magazine ofBotany 11:53.
PERRY, L.M. 1980. Medicinal plants ofEast andSouth-east Asia, attributed properties anduses.
Cambridge,Massachusetts, M.I.T.Press.27.
PLAIZIER, A. C. 1980. A revision of Adenium Roem. & Schult, andof Diplorhynchus Welw.ex
Fie.&Hiern (Apocynaceae). Mededelingen Landbouwhogeschool Wageningen 80.12.
POPENOE,J. 1975.Oleander cultivarsatthe Fairchild Tropical Garden. Florida State Horticultural
Society. 426427.
POTTIER, A. 1879.LesNerium (Lauriers Roses). Revue de l'Horticulture Belge et Etrangère 5:
274-276.(InFrench).
REYNOLDS,J.&J. TAMPION. 1983.Double flowers, ascientific study. London, Pembridge Press.
RHEEDE TOT DRAKESTEIN, H. VAN. 1689. Hortus Malabaricus. Amstelaedami (Amsterdam), J.van
Someren.9:1, t.l, 3,t.2(In Latin).
ROEMER,J.J.&J. A. SCHULTES. 1819.Systema Vegetabilium. Stuttgardtiae (Stuttgart),J.G.Cottae.
4:410.(InLatin).
SANCHEZ MONJE, E. 1980.Diccionario dePlantas Agricolas. Madrid, MinisteriodeAgricultura Ser-
viciodePublicaciones Agrarias. 199.(InSpanish).
SAVI,G. 1818.Flora Italiana. Pisa,presso Niccolo Capurro. 1:23-26,t.9(InItalian).
SCHICK, B. 1980.Untersuchungen über dieBiotechnik derApocynaceenblüte I. Morphologieund
Funktion desNarbenkopfes (Onthefunctional organization oftheApocynaceous flower I. Mor-
phology andfunction ofthe apex of style). Flora: Morphologie, Geobotanik, Oekophysiologie
170:394-432.(InGerman).
SCHUMANN, K. 1895.inEngler, H.G.A. &K.A.E. Prantl (eds.).Die natürlichen Pflanzenfamilien,
nebst ihren Gattungen undwichtigeren Arten: insbesondere denNutzpflanzen. Leipzig, Engel-
mann.4.2:109-122, 176, 180.(InGerman).
SIBTHORP,J.&J. E.SMITH. 1819.Flora Graeca. Londini (London).3:t.248.
SIEBERT, A.&A.Vos. 1896. Vilmorin's Blumengärtnerei, Beschreibung, Kultur und Verwendung
des gesamten Pflanzenmaterials für Deutsche Gärten. Berlin, Paul Parey. Ed.3.1:659-660.(In
German).

SWEET,R. 1830.Hortus Britannicus,oracatalogueofplants,indigenous orcultivated inthe gardens
ofGreat Britain. London, Ridgway. Ed. 2:354.
TOURNEFORT,J. P. DE. 1700.Institutiones ReiHerbariae. Paris.604,t.374(In Latin).
TSIANG, Y. & P. T. Li. 1977. Flora Reipublicae Popularis Sinicae. Peking, Academiae Sinicae.63:
148.
WEINMANN, J. W. 1748.Duidelijke vertoning, eeniger duizend in allevierwaerelds deelen wassende
bomen,stammen,kruiden, bloemen,vrugten,enuitwassen,etc.Amsterdam,Zacharias Romberg.
430-435.(In Dutch).
ZAFRIR, D. 1962.TheNerium oleander inIsrael (In Hebrew and English).
CATALOGUES:
BELGIUM:Verleeuwen. Gent. 1820,1835.

FRANCE:Audibert. Tonelle. 1822,1825;
Batlle.Ille-sur-Têt. 1948,1952-1970;
Buc'hoz. 1799;
Rey.Carpentras. 1973,1984;
Sahut. Montpellier. 1855,1860-1869,1873-1876,1878,1898.
INDIA: Globe Nursery. Calcutta. 1952-1953,1955,1963;
Kohli. Srinagar, Kashmir. Sineanno.
ITALY:Baldacci.Pistoia. 1952,1956-1960,1963-1966,1972,1975-1979;
Battista. Mariano Comense. 1895;
Berti.Milano. 1894,1902;
Bianchi.Pistoia. 1950-1951,1953,1956-1967;
Capecchi. Pistoia. 1959;
Fedi.Pistoia. 1956-1957,1959;
Herb&Wulle.Naples. 1892;
Mati.Pistoia. 1977;
Pacini &Boldi.Pistoia. 1956;
Pagliai. Firenze. 1872;
Perotti.Trieste. 1883-1884;
Pin&Gullino. SanRemo. 1894;
Sciacca.Catania. 1962;
Sqaravatti. Saonara, Padova. 1896-1897, 1899, 1905, 1913, 1930-1938, 1942, 1956, 1959-1966,
1971;
Vanden Borre.Treviso. 1954;
VillaAda. Lago Maggiore. 1882;
Zanoletti.Milano. 1880.
JAPAN:Yokohama Nursery. Yokohama. 1891.
PORTUGAL:De Mattos.Porto. 1910.
SPAIN:Aldrufeu. 1884,1890.
UNITED STATESOFAMERICA:California Nursery. Niles,California. 1926-1927;
Clarke. San Jose,California. 1929-1939,1946-1956,1958,1962-1966;
Glen Saint Mary Nurseries. Glen Saint Mary, Florida. 1925, 1959,1961;
Michel'sGravoisGarden. Saint Louis,Missouri. 1849,1869;
Miller &Sievers.San Francisco, California. 1874-1875;
Monrovia Nursery. Azusa,California. 1952-1953,1957-1967,1970-1971,1983,1985;
Patterson Nursery. Albany, Georgia. 1959;
Payne. LosAngeles,California. 1922,1926-1927,1929;
Peters & Wilson. Millbrae, California. 1954, 1960,1963;
Striblings Nurseries. Merced,California. 1956-1957,1961;
Sunset Seed and Plant. San Francisco,California. 1896;
Thorburn. NewYork, YewYork. 1852.

APPENDIX
TENTATIVE CHECKLIST OF OLEANDER CULTIVARS

J3 T3
60 O
'E S
ü >
ca -^
o ,2>
jS £
•a = rt
.5
'o. 00
•a .1
43 ü M u
T3 c y g
•S g o §
's gg £
"O .G O t-T ai
J3 -S o as e
S
S -o
o t •,o." rh
CG HG CG
.2 .2 §• oo v a oo
g l l l GÖ& - i l l l i s 11
. ! ? § • § . 3J g § -g.g's g.g. «g .1 g g S. S.
•- -* 'S"3
-j Ä o- - Ä 'g 3
° .2
° O » JG3 •=uSi « ï » °. °u -- nÄ _ i _
«2 5
ai O >•>b û a uabuUP - U «! O«;>-
ob>-Q b Q aau b U b Q
••M
8 *îot
G C Z
•S . og
CA u T-j 25
SS ja -a
g c .2 u
ri 'S
s So
S/ST
&a «
c ~
o 5
O \28
'°. s u s "a »s o
•Sb S « > « e -g
si s £ g ^ i 8' Ses
.3=3 § -s* 1=5 g a a |
£ £ °£ ^ -S G C - K S - G S Sä
-"- C «> G caH 00
8 8 ^ - î f? ^ 5 « r i M .6 « * ï i rôo .5 ,3 S r- .$CT>
m m TftZlbi-lc/3 —U — tN«!diiJ O I O ^ U - e>. l a c- ( * n M > - •
C 5 „ GG „ C Ç
OH O O Û . O O P H O O
3 c 'w "-5 3 c '-3 w 3 •-' *—
O u -n, ca ca O u ' & ca ca O u cd ca
o u
•û XI
V =3 ä w _ - £ 3 < S 3 3 v S ^ ^ •* S'sSg. « S u n - « 3 3
O. O.
& -a •« .£; S 2 « -S P & ?• S > S 3 S . S P 0 - 0 - S > S =S *e u
S 2 5 s 5 ô » c " , ! s i i 8 3 5 Ô » c S.b ea .53 .23 S .s ~
Ö O •§UbU«:0>'bQ -CUbUBi 0-B.Q «EubUtti b û
< < < < <
3
c
e-. E
I I
>^
a 3 | >• I -
« -s .S • fi -S .fi
h "a M w x w „ x h s?
ï < g Ja "a =» S 'S « S >
Ijl
. C •- . . C a
11 |s
g g o .G u
. l | c
s . u .. c : |a-g
o u - u w o> N r" 3 .o u u ÛH 3 £ M u M
•5,-- » •M ' M •= P? w ° ^ M 3S » j J „ ,o 3 -s « j S
OH C
fe a. o o s o , o o g p. o o g s
g. o o
Js 3 \E w § 3 w w S 3 w w 3 'w ^
S o u cd cd « o u ta «s S o o ca ctf a o u «J ça
S £ o, o u B i - D . o u g fc O. O D ,B 4 6 o u
§o^ i i -go£. i i *§.o^ i i a go^. | 3

a n s a ä ä s?össää s ^ s s - s ä ä ss-gsslää
l i l ! i i ä I U I 11^ 11.111U S i | ü l 1 l l |
Sunuaifcû £UhUtfï.Q 5i»Ufcü«iii.Q SKÜUHUKHQ
i SP
u to S ™
I I< 1
1 V-
o 2 «u
•ë E or i_*e
1 "o ÊÖ
H
o £ .5
K < 3 -o
a . - S s s2
h i3 2 "3, -2 o g g
•S B £ § g ft. .2Poc
S <L> 3 i S O n t / S - " ^ öS C
H g g ^ g o o g M^ ft.
^ S E o £ £ 'S, sc N
^ SB. as .* . E < < c
.1 '§ -8 -S
co>; § 3 c g u S 2 ^ g S "a
G
• ' S , .3 £ £ •'B. o 2 .2 .2 B • •£ * 3
C
M £ v ai 3 < Ja17 2 S E E 3 Ü 2 | . i Ä
eSSSg; u u cS.SP&g; u £ 2 S Sä < - 5 3 ^ £ . S S ü ^
cc ce „ cc
o, oo „ o< oo o. oo
3 w w OJD 3 w 'w 3 w w
O u ca ca a o o ca cd - o u es ça
O ft a â ."BÖS äS - .§ -5 ft SS
s
^ s s s ä ä "ta
.2 w s o g tS w
S 3 o "Ö S .= i
^
ta s Be 3s'• ?„ - go-g•=o-g«isaw«ä lo .go . 3&5 '?s .2s s"o sS s.E caa _
. 2 O ft. U oä ft. Cl £ £ £ O ft, U•~aS ft, O £ £ CH £ t «

coU ft. aU O«S' w ft.
& o cQ t* u.
< < < < < < <
58 Agric. Univ. Wageningen Papers 87-2 ( 1987)
S ä 'ë "
= « 7 3 S 2« «
•ä 'I .e
O *- O S fi
a
t; o e-- cd g
a 2 » •§ J= c >,
£ 5 ^ - I 8.^ . g
&S s.s .s >• a ^ s- „j -a| ^ .s a, o
9< O O D H o o a - O O O H
c C
_o O
SE, ss sg, u :§> s s ga §s sa cd eu

O o
X> . o
3 3
=31 g 1 3 &| „. sä 1 g o g.a s ö s s a s. I ^ § s „ 1 O. O.
aj
g 3.2 o «.3 =5 « 3.2 o S S 8.s n 53 .22 <3 .= « J>3 5 ô S S - M *-;
BÜ&.UKH.O s OO,O o<O>• OH Q S ü f c U i f c D ËUEUIBB! E U
5 •< -< <
cd £- cd -JS ^ ö in CL >
c
3 tSSs - 3 g S •
u
~ i> . cd .5G JJ. cc s . -S Ä "S
„ c e ce ec ce
g* oo g, oo „ oo a oo
3 -as ' S - 3 'S 'S x \ 3 'C 3 e -S ÉS
Ë 2 S S EPS ni« u n u • „ o u -S S a
I g Ç Ê 2 2 gOÊ » 2 2 I g 2 2 | ^ ^ »o' 2 3
IEüJll| l i l i i i s H is unguis
5 jC <c «c

* 3
» I
o cd
t/1 3
co S
•a W
r/3
tS . .- -i | -s s s- _o "2
JU
N E "H,
Popenoe. 1
Vigorous.
.g o
u 'S,
<
l î i l ^ . lia« iiîPs^
1975
.S J3
(N en on tn
ei
r- G S 3 3 3 3 ac
o, .2 -2
a o o < a. oo g. e-S -S 3
O a>
"cd
% sa 1 1I s a si sg. ^fll *s
o "55
s
emarks
ate pub
irst pub
CA
O ri tH l , 'S
s O | =2 -e^^ 3 3 °£ »0S22 > S3
M •3 S O u
"3 Q 0 "a
l u i s î t 1Ü1.1S l i l l | - | | l | eu
a. U l l ü B ifeO
o
v..
I 11.| I£ £
S 8 3§1s" | § -S" § I l
* & £ S^S& £ g B" £ f ft.
8 .1.3,35 ^SäSSS* »Êla.öS .I.SSöB

§§ 1& 2.2. g- c.2.2 §• CJJ
1 J
S _ ô l a f M s l ô E â l ë ^ È S i ô l a ë ^ E û îsêôôss-Êa
< < << j< ?•
60 v4gnc.I/niv. WageningenPapers87-2 (1987)

<
1/3
CO 3
8. X•o 'S M wc«

o ccd
i-t
J3 fi 13 $>
o E
*r-ö3 o s eo ts . £ ^L J2
Jxo-cd.LLl • „ c d . r r - c d Ja -3 u r^:
.o _* h « „ ^ . ü -3 (*> JB .e 'S t" £> Q. o o V
•Su — o o r f /2 " oo •- « « oo .5 j - ca oo ,°5 oo o fi « ca \- cd ^
g — O Q ft, « C — M B r - o l r - U - CN <Z) « OO— O — 5 O fc J [ I , Q S
&
•2 B" .2 .2 & 2 .2 S S «
| sarlis gg. -Eli Ia i'E £ u o

ft,
3 3 cd
o
S ££ ° 33 3 „ o^ ëëss o >• B 3 3
C. D, .S
o. g Ö S c o c o , | ? Ï S c o f t f t g > " 3 !a «3 u C
cd
« 1 3 3 o C S .a a & 3 ,2 o 'C 8 .§ * •- 3 .2 ö S il Q 3
O s U ft. U O >Ha, Q Ëfo o - ü O >- (EQ §uEuoi <
| -a s t.
u < eg 83 . >> a o ca
B ft. S 'S ~2 3 "^
3 CA ca
1~ *-,
Large flower
'Double Dar
Largein: flore
Cat.Vill aAd
Cat.Verleeu'
Cat. Cla rke,
Dark purple
o 'S,S
Bright re :d.
DarkRe id.
i - Cl
Double.
Double.
Cat. Pen
'FloreA
Darkpu
'Italia'?
Single.
Single.
'T)
1934
1883
1882
.
11
00
11
B
c c c a a
s §• o .2 ° o.
3 _o o 0- o.
3 .2 -2 P.
C
o o
0
rou
cati
cati
a o u ca ca 6 O tu "ca ta E O o l a ca
pe
S M °- ° 2 3 o o 3 I-i Q^
.0-H
,fc °<
erTy
nyms
•s 3 O >> — =3 V 0>
3 » O >,
varG
crt X I Xi 1«
ez, H 3 3 3 35
g O 2 S
•S. 2 3 S
3
o -
ä O.
«3
a. > t cd ^-
3 2
CL
h.
3
t. 3
3 o . o. Ja
h.
a. C '-B
> ?u O c -g u O. 2 2 * ° S S
O g, 3 .2 0 .3 « o O S?, 3 O O . 3 « S 3 .2 o B .= =a
Ö &o U ft. O ft. Q b ( « O ft. U Pi ft. 0 h. tw U ft, U ft, 0 Ü U ft, U a! ft, Q
< < •< •<

•a
tl .S
-O
cd
5 PH ^ "« a.
0
u
pq ^ > i
O
M __, ai cd
»
_o
I-H
O u .a B
3
£
1> xi tg S 'G
X
t-l
«Z
JS g-s Xi
> Ä .< -B nj a o
c .S 0 H
u< o o u
" t/3 m g o ai 13 3 53 J - |
"&û 6 0 : 3 t o IC (N
HX S g .S 'S * SS g . S S « $ c 5 O ^3 <5
—1 U —i s e-. t/J e*-.<Me~ U — " c^. f . J , | i , Ù Ï Ï « 2 2 (3 2
hjij.ii1 pillais 1imp* üilii^

je a co ca co so
P.
's*
T3
ö
e3 -au
•3 % 1 j-gs
si- o§£
| 8» 3 Ja Ö 8
H£ 8 >. 'Si ? S
8 g § §g « aS c „
"2 > u -r S •- h oo 'S
a°-g *§•«§? I'S S J
„ I onS-5.SS?öS
- £ 2533,38*
•& £? j H . ô 'Sl l3 g
^ "?S^ g . v> <
u =
o. I I a §| a §§ a gg
-als 11 s|& 11| s - H l nigI !s
3| | £H l | gfeS1 äa. .„ggsg-SäS. SI S3 3as. 1
g =j o c; S .h « 5 3 O 0 Ü . S S ^ B O Q - C U S S S 3 0 3 i23 S
S < < •< <
"2 Agric. Univ. WageningenPapers87-2 (1987)

— 3 o-g •o
cj X ) CS
w CO
S
c3 M
e < 'S £>
"°
C
CS
si o
Vi
•o ^
B 'S eu 3
3 o 3
CS
3
eS < J
e*H
Ö o o O
3
c c X
es O
2 o5 3 t-l
5 •S
" o. Ç.a °" ca Du
c
C ca o
o
si *s • 00
_1
•g "H S
G
."8 2 §2 S
c«
o 3
Ü « >> 00 T3 •O -C p -. g
•o
1 £S (U eS M
00
O.C3
U X"~ ° C
00 «"go o -S S
3 is «y 118 S -H*
"Su 3 . ci
o o- c 3 « .g si "S S"
8 8T3
£ S S.ötC
S
e .u°2
i 'Vi S3S . 03 — o S £ 05 • ' Ä C-
CS
Vi <N !» Q n, o5 o. 2
c 0
s c o S
o a.
3 3 3
O Q 2 o o
o O (U 60 ü
%1 k-
U Q. ! O,
•S.O XI 2 o x i j xi
3
x>
S 5
K.2
. °£ - 3 3
o. U 5 3 c o p, a
6 O '5ô S tZ S
o
S S g 3 3
S3 5 ö »
3
a o.
— (U
m <->
e 3 _2 o tl B $ cd t* t - fcn J j cS
« U f c U f c û OS . £ g 3 « pH 1 ü Eu O >• EÛ
03 •S 2 * .2 Ë
9 3 o o S
03
>
Ü
<r OH
e5
OH
PU
C
C c
üu o 3
o
'S.
. c 03
O
e. s cS a
«5 .
2
*-
si
es
(U o (U , 03 t/5
"ôo S G G 00 où S
.S 3 05 c- 03 o — CN Vi ÖH
(H
J U S
es
•
C
c - CÄ c - Vi c- O -H CU Vi ,(JÏ
Tt Vi Vi
c c c
o S
C
c o o o. o O. O
3 G '•
3 3
O (D O e -a , es
O 8. t- CL '60 i o '50
O 'S 'H ' C
S Ü XI X3I X )
O 203 o I 3 2 -5 «X 3
Si s H 3 >* es k- 3 es tn kn
c 'o
s o, o, 3 3-
M
O XI O. C > o 3 o. o, > o 3 3 C O
•a 3
3| 3 ^ O 'C (U .Ü
o o, H o s 5 3 G «i -Ü3
3 es
b in«n
2 * _2 ' S S •s ^ S '3 es
O
U f cO U" O > « t t <
es Vi u E u 05 E Q u oEüO>-EQ
•= « o Euo><Eû
ä Q
Agric. Univ. WageningenPapers87-2(1987') 63

CO
i/3
c
CO
C
3
c
I
33
- «
i- &? ^ o3 §
ill
3 £ éo <"-
I
« e S. h 'S o
ril^ê -e
51 Sr-
| ,11ca e S « <N
c-. C - Û . - J M — y m » ! U > U - • cv- o- r- l/l CN- O — y e i c*i <_>u —
c c c c C C c c
O. OO O. OO O. O O O. O O
-_ 3 ' 3 '-3 3 3 3 ">> 3 C 3 '3 3 3 3
S p cu cd c« O co « co M O co - c . cd cej O co cd ctj
|?H ja-§-§ . | a ^ .si-g E ^ H f °|-§ : 2 | Ï H |-§

2 2 * ° Ë » 2 S 2* > » ES S .5 '-= S o •& >= -g S .S 2 '3 J o - »
? 3^ o ei a • g t s . S o u . c c s "P s .2 o ' C u . 3 ca ? 5 3 £ ö . i ; a
SUfcüoafcu SmUMJuifcO S U fc U O >- euQ S M U H U H D
:cd
60
C
£ c/3 ~
8 3 O "2
§ '5S 05 S « *
I ri > 2 S S
S # .2 O T 3 0 g r-
• s i c . a l z ^ ^ 5
^ cd eu „ ; 13 _ - 3 •O
(L> w ca rn ai - eu r 3 . t. - • C— TTI
</5 rJ > . 2 >< "^ S « UTS g U u ™
C -Ç S <~- u O 0 -3 S 0 0 0 S B C d Eco X!SS"
3
e-Mo- W c - U - S Sr Tf t/3 PH CQ —1 x-i H !> H CQ o - • cs-ts-l>Cac
c e c e c e c
g<
3 c 30 '-3
0 30 - .3 -0 3 0 0 .3 3 30 0 30 . 0
co -r,. cd cd O co cd cd O O) cd cd Ovw
°- i_ • - .u .a .tt O. o .0 t. o. o .0 t o -
| I § 3 S 's ä ä § „ I g s s S. ä s s s s ä S. 1 s s ä g.
52|°:S>S3tss g •= g 2 * ° S S J 2 * .2 S S S S 2 5 0 S S
B 3 . 2 o G e o . c c d e 2 5, 3 ,2 O .3 cd ^ 3 .2 O eo c "3 ^ 3 .2 O 3 cd
u U U u u
64 /4grc'c.f/«iv. WageningenPapers87-2 (1987)
.c |
•5 u c°
"c"~
•s „ c e
S -° -oE
a> u- —•
C J 3 cd
•O ä
•o CJ
<
enu
C3 ;—
U| c3 c -i -sO_s
co a £u " 2
O
O. e jj W —-
k4 £ o
3 1
s
Cat. Rey,
3^ cd c
«JR -s
Hardy.
cj
O 3 60-2 3 m
1973
"M X! i- G XI r~ -S
c M cd oo
>o .g c^-
cd
C« o- 2 2 <N t/5 CO J SctZl — U —
c c C
C C
Q.
S
o.
3 .2 -2 3 G
o '•3
o S
O O.3o. ~_ cd t- l-H
M 3
M
C 'O
s Ci 5
o.
3 > S 3 'S
« \3 _o
s to i î a. •s S '3
e«o en
1 3 o "o
o
i - cd 3 ° S 1 uEuoi O >-EQ
5 Ut, Ua. tin Q U n. ü O >- X
U U a ic
y Mb ui c
w :
Sr i 'S 'J «
s- sV ?e g>
I °S .-I s
s Se MB «
. 3 „3 cd o n
•a U "a ' 3 ^ CM u
•s« la M'.'
S S ES" 3 oo x!
Ä o « r" Q« -a-o
J
x? 3 rtu S =0 -3
- o. - 2; E P3 fi* s aS
MM
cd- ^ -
fp,S
-u T3
tl J | * • •§
S-S
c û2 -X! •
g-»2»
O S c »-
I55 fs
S Ï !
. :§r ° °
I S 2 c $ Js§3 1^ u;^!*
•^ 3 3 3 ^ " 0- T1 3 -S S 3 N > S -
o
c
cc e c u c c ^ „ c c
&
°" .2-2 .2 .2 0H .0 .2 g §• .2 -2
O t u r t c d - o u " c d c d u î d c d S O u e d e d
t a u o o ,fe o. o o o " 2 § A o. <->.2
1 - C T > ^ X 1 X Î C <«(•}>. X ! i 3 ^ - c n X 3 Z 3 j ï c n C T > > ^ = X 1
S^H 3 3 S ËT1H » 3 -o ». g 3-2 fcg ^ H „ 3 .0 -2
I l glaäo. s i l l ö ^ s ä
|2*°S2 S 3 § S S o | | - 2 go
ä | | ä ä § 1 5 f ge s3 32| ä S.
-s = 0 B J o 2
B 3 , 2 3 3 C d O S K a O Q U g t H cd C F . 3 cd S P 3 O o ci 6 .S cd
S U C U U Ó H Q S S w ü f c ü w l ü t t « O S w tt, Q se« UfeUffio-fc. Q
U ü <J U U

CU o
ca s a
> o.g
 5 ra 1
•K .S h -G O IK ca ca •-
o fc.
Hl
ffl ffl „
o 3 S £ „ ._ c c
T3 o T l *-> L* '—' ca ca U
</J ca r. ca t i , ~ ca •a •o
O 3 ca o . c
ca
c
a o CS G c
o o
P o 3 e e
o
S s s
<D -ä C« o t/1
U 3 3 O
U - m
J3 .e
-;-i ra r=9 oo
H ^2 j e j .ü - c-. c/) i>. PH t/1 e*-- U —'
• (/) tr- V3 c- U —
G e
o a s CL
ffl o 3 g '-^ e vo'-S o

<J
60 ca
O CJ t- M '3» 8 o. '5b
00 o o ra <D
O,
-rt ca
00 o
ra
o
s
w o—
I"
« M
3 0 2t - " L •—
O
*^-
Ü >% b 'v- » a
•O H M M ra 'S S 3 ° O £> X>
erat-,
S> » G' so S
o. 2o. « CQ 5 >" » 3 S
ca LH
n n S>H 3 3
5 3 G O O, O,
III
O U IL.
<D . G
oi O >• E O
ca
co
O.
S S
o
öó IS 'S S
g 3 , 2 O S i j u . 3 Ä ra
S < J B H U 0 Ü O > < & . Q
. 2'Cig»uca.bSSraS
2 3 . 2* O
§ U f t i U O > - & i Q
c
o
u O U U
ü
•- 3
W ra
g G 3"°
ca oo O
c 2<
•3<"
'S ff a-g
•So
" c
-o <s ^S «e
<* . 2
'T) w B 'S
O ra oo ça
— T3 o ha •o °
.. C c oo
n " 'S c
Ë ca U •o ' >5 o
C g c
C3 Ö o
o ca -o u ca
u rï3
-8 . c •tu
o - ça
A ca
^ o J3
u G
1 •2 a c o ca ca
u « —• t/5 ca •o
G
Jul
U C >ä ca
o G
ca ca
•a
•dl 2^-g
c •-
§2
u >ra
co o
o ^
•- c
•
'S.
u 'C c
S u o 3
S
CD
T3
a
ca
g
I
O
y
h M > , ^ ü o 60T3 o S 5 J3 3
ra g
o
<L)
. •= .> Ij j i o ^ ca c o * tzi U
^ « J D i e S ^ o ï S y "äb-S, 8 ?? « 2
•g gj ;^, ^- oe g3>§
i j o°g
>5 3
ë
• S A r^ «^ g .2?.H « i - ca oo * oo 8
m c« i-J H
r^ (Zl CQ ( J — J U< oo — U — a
c- o.
3 3 ra
O ÇJ 'oo ca ca
8 g. 8. y r» " i o o ca
s. O S J3 Xi 8,0 H 2 o 5
i= 3 e
cä -tl
3 3 ™ t-i
ca '
3
3
•—' k-
ca
D, o.o. > « 3
" > » 3 S G
D. O, ca
ä J ^ *-;
« tu
Ä 'ÖÖ
S 3S Ô S h 3 O O .tl S S 3 .2 o g Ë
£ o e-, U u,o Ä O fr. U t i u t3 ra o o
u u O >< PH O
u O

o
o -a •o
U. CS
c
v
cy ~ *H
&Û w
S I H » o 'S
1)
3
c a » 5, c 3. •Ö S
JS 3
0> r^ -C 3 eü 3
a M -c
r^ u -2 y .£• a •o 3
^ M c ". UU tu o
es
h
z
M 1>
CS ^
S £g 3 S« 3 S B
_cs x> ca J3.
"o •o u r—
>> c 5^
13) o
e«
.3 E
Cü
s
„ c c cc ce l c cc c c
g.
g
o o
'O 'S
a
3
o o
.2 .2 SE 3o. O.
-s -s
"8 o o
s 'S .2 -2 a
o, o
p u es es Ov es es • - s O u 3
es "cö O U es es "es CS O u 3 3 o. o. g ç > u 3 g a a 3 5| c > Eu >C3i Ö>g o 3 2 5 ft ft
Flow
Syno
Culti
Rem;
Colo
g g3 g.2 ß S S | ° 2 * o E S S co o.
3
= wa5
*
g 2 StS o | - 2 ECfl
c «
S-(2 | o s
feg, 3,2 o &.- s S . S 1 5 Ö o . t s PUb Q6 3 SCL 3 3 S . c M *u 3
C Û o - j i
J y y o
cd
C
-C
o<
•a c«
•3 ~ ^ E
u
ÇS _: ^ ^
g
cet
IJ"?
S aN <s
ji* S * > oo g
es O i - T u 5î • =3 ; ° « * " "*
•a 'S f § R. 5° g S Ï ° Z 2
S 3 £ E | g » 2I § y ^
2 &a § 3 *| S | fiÄ'S ^J
- §. las | =§£ "» c.-s dS-sll Tt
• O ej* 3 > ., es . u -S .. O-^ . C3 ^ 3 u d
U 0 ( Z ) c/î > <-; c/j u ;>,(/] > » 2 u O 2 J3 . S i ' - ,
tCC J !; O " 3 ""> . \o ' S u ^ ; ^ 3 S ^ç ^ "äö Ë ^ 3 ÏT C
S S
=^ £ c 3 £ 1 1" S " l$% S Sri- -. .ÊiSc?^*
5 -tf <*>CuU —< S ° r . p. U U — S^ttziiziBuïI^S —
§ i
S§• -2-2 & 0 .2I §• 2I & 2.2 1 g- §2
32 1 1 1 :Ë 2 1 ; |§ | I s a sI s». BI | | a 1s
l ï t ï 33 JO^ o ë g | Ï O ^ 33 gü^. 3 J ^-OH » 33
! 3 ° O .S a a 3 ° 0 ' C u . = M « 3 ° 0 . 3 « Ë S . 3 ° O S Ö M 3 ° 0 « U " tS
SUöuUüuQ SUBHUOÏHUUQ Sutuuuuû t«Uft.uli.û ' S ü l i ü ^ E6Q7
J O Agric. Univ. WageningenPapers87-2
U (1987) U O
G
•g
'5, '>
•3 60
O
X! cCU
•o
H c tcdH
C3
"o Ü
0 "cdo eu
3 -a IH 'S
D. 3 CU 'B. o
•o EQ 3 o
<U '3 b- 'El
3 O
C o O
o 33 cd
« O cd 'Eb
cd p Bi
3 «J CU
CA _o
Bi
•s roe C
S
tl 5-8 O C/5
73
•o C3 O 75 c«
CS "c o <
S '5.
CD
PH o S Ü OU
e & BH
< o CU
J3 " eu "o
.s •* o ' c
S 0 H O
3 O
o
G eu 'S o S
co o o •o c
eu o X!
O
I-i
3 d
p
o! -SI
« «? CA Q *s <D cd u
OO M cd oo .=? oo *c c e o
.2 -2 3
S PH h j PH CA — C J - H s CU
c o _o '5b
cd
O eu o o o 'e cd 73 Bi
Ü _'5.1 1 S « 'Sb
S çu
2 o 3 2 « O S _ 2 o 3_ x> cd
C- e3 3
Ü >. 3 3
> ¥ 3 —
ta<t; 3 3
G O Q, Q. S 3 c a o. 3 c a O. <
o o
S f .2 oo O \S CU
u
»s »S 3S O
•Ei G CÄ
73
o o
Ï Ü H . U t« eu . 3 cd
O > ti. Q
o CA U PH U CA OM O >- PH Q o >*PH Q
0 CA o
O Û Û
o
•o
CU
cd
CU
60
G
cd
J3
< O
cd
CA cd •s
o
c" cd X!
o CA H
O
G
o 'S
3 G
J3 'S
•U U «5
cü CQ o.
O 3
G O H
O CA bO PH —i o2 CN CA PH
& Q t*> N^ H—^ O
C G c c
O O
S o. o S
o
3 .2 -2 3
cd cd XI o a> 73 M O u
o p cd
O r/i CA rfe Cl. .2 -2 ^- o.
c« 3 x>
i. - ï a 3
CU
•a
3 3
? ^ -° -2
4>
« -s
Cd l-H IH
^ c_ o. >* cd
G >
O S ;
3 cd O. D-
_9 S tS S
3
cd
s i « g EU ». cj i . ^
« > çj 3
S cd 3 3
D. P.
•U 3 . 2 O . 3 Jd G 3 _o Ö 3 oo o .ia s 3 S *H eu
s U S. U CL, Q «CACJPHUP-PHQ u U P- U P- Q a g so Scd P i .G

C»
S ûM
4-1
3
Q Q Û « 3 O Q
eu U PH U
Q
s.lc 3 O
eu
00
E o. eu l-<
43
e o
Ja o*
o
oca"
•0
3 '3
°6*\
3 S3
'M
2oc , o
U H
c "3
ca O
e*-,
O oS B o o
<- M < »
c ca
3
U 5^
e
3
s-i
ja
CA
c
ta. eS
'5b
CU „ - C CU 3. J2 ca
cu taJ 73 00 <
43 o
2 <=° ei - 00
S '-3 <*) CU
oo »O
GO
H4 O ca o
eu § 3 4*! ca G OS < * eu 3
c o — u 43
cta.o
e«: ca
t-
o OB T3
>> JU • *?
"éu
O
o H o
y Ö •o
o
|.Ë cu O
a
S S 3 fa • oo O
1 8 o .5 5- o o\
S * o v Q £ Ö 2
£02 tr-. c--. e>- (J ~ oo [> Û. —
c
o. o. s
a 3 3 o et)
o
O u ca =a O ca « cu CB
o O. o cu o o
a6 tZl C/Î
Üta.>> c Ü £ '43 3
43 43
O >> ' - =5
73 43 43
S ta.H
43 43 (3 o VZH 3
00 3 3 ca U
S 5 „ S3 u u » 3
eu >
e> O. O. >> >
13 eu 3 Q. >3.
o
S3
O £ <D —;
O ca 1 23 * ° OS*_;Scj « ,23 S — c« * -
3 c 73 o OS -2 ô . „ ^ à U
p sö ioü.2i i «i O
O 00
3 ao V Cu U £ Û t-i O Cu O Pu Q
Q û W
c
o
<C
oo
§<
3 00
<
C/2
> Tl-
00
.« en
'Atroplenissim
Cat. Clarke, U
S3
l-l eu
S 43
O o
•o
ca
-, o* Jà" ca
 o o
11
.S -SP »s S
o
TJ- oo NJ U — 2QS
'SD c c c
c
g •o e c S3 S3 g
D. S3. _o \ O S3.
as g- O o eu S .o 3 .2 -2 3 £ 3
o o
*-3 ' I - I
O O eu ca ea cS ta ca O eu ca ca O eu ta ca O eu ca ca
o o u a. o o <Z C i ÇU u J3 S3. o .o
First publi
varG
publi
publi
43 >i w _3
Date publi
erTy
Syno nyms
ri 3 33 3 3
irks
eu © t. ~ 3
a. OS ca u t-l 3
a « ca i- L- 5
ur
a> > u 3 & & > ÇU 3 S3. • > » 3 S3. S3.

S 2 * ° S S 43 J» 2 * o Ë S 2 eu
•S S o -g u 45 •Ö S O - u
3 3 S o u b ea
g 3 J5 o . s ja 3 3 3 3 O O S ee) 3 3
O U
O ' o e. cl
o ü t t i U C i O O
e
U t i U H i f c D 3 U CL. O CU Q Eu U tu Q
Q Q Q fiO fi
J4
C
"S.
3
3
O
•s 3
_o .2 43
vi CD i« T3
>. CS -o rt
O
E
(3
cO
'S (3
cd J3 C 3 Œ
o CJ — «1 5b
co
^ 3co
43 75 73 CJ
H 60 S s ca •3
4^ c "C 'a 3
e O 'C e
bh CJ ccd oo
'S. s
Free flowerinj
zu .2 CD
Cat. Verleeuw
t-i CO CN
cd
CD Z
c 43 u 75
>
Dark red.
O H •B ó •ó
'M
« in CD tu
£ 73) co
73 "Hb 43
II
C*H' <-* CO c-l
où a> Ö ° co
.S 73
h.
CS
aN CO 0\ G 60
o.«
S ON c
Si
fi
oo
CT* o CJ 0 0
CN ai ft. S <N CO ci
co c —
Û t/3 N
c c c c a e s c
a o o O. _o \ o3 a. o o a o o
3 \2 3 3 "M 3 \3
o ej "cd CO O CJ 7d CO
o CJ
73
CJ 73
O CD CO CO
a o O u O. O O k. D. CJ k. O. CJ CJ
O s-. O 0 E niw
H>. >,
> •
H t/3 43 3 43 H 3 43 3 43
CA 4* 3 3 «1
3 3 M 3 3
co
« 3 3 O. S CO l-l k.
3 o. O. 't»
cO l-l U-i t-i
3
3
O. a cd
> 1-
CD 3 3 O.
2> > °.
(U (D
« 2> g ° E 'S S
Flow
Colo
Culti
S E
First
CJ
- û (Ä -irî
3 J2 o « 5 £- « cd ü ed a 73
V UH O ffi tf U- Q s 73 S. 73
U ft. O
PC Ui û 3 2, o S ü cO 01
CU
a
3 3 uft,o aft.o Ü
C
o S * 'S.
43
43 'S "S
S 4* £ CD
'I
•o
43
3 J
w O 43 S O. cO
I 28
S S
OQ
>»
•" t-.Ss o c
cd cO cO
— •; — CD —— n
ou .s•*
S ja a « • sc O S J . 360
o
t) u - ' CJ
Ö.143 •a S2 ^
u fd CD . BÖ CJ
cd cj
eft
t* CO cd U- '
O =3 43 '
O
» . ' 9
(3 i ?
c
CO
O
W ft. j j
tu u " P
N CJ> ' 3 CD
S I « S
O
cd CD O
 i r - o Qsb -.t. .>- Ccd
S S ^ o j 4 33 t - - ' 3 ! f -
S
oo J3
43 r°5 oo
o 43 -a =?tu£ <^
t>- C/3 c - C/3 c - U —'
i H J 2 J c5)N 2
c o c
s o s
3
c 'So 2o C VJ CD
C '3 o o
T n cd cd 'S ,K o« 60 S g
,60 cj cj 2 O 3 43
2 o3 3 < h f- cd <^- 3 3 feH c» J2 4= 43
3
3
CD 3 C O O. O. ï >2 S
- 5 .S ° - •O. D,
g Ë » S S 2
CO l-i l-i 3
43 2 * S 3 Ë
ij
*-> CD
* s :s> CÔ s s ' 3 O O H CD C CD . S Cd
'S 3 2. O cd CD
co ^ S
3 <"
O O C 1) 3 rj ! cj ft, o w oi O > ft, Q ft. D
t . U O >- 'ft. Q u
70 /Igric. (7/ii'v. WageningenPapers87-2 (1987)
'•g
, ca
!C
°
oS à
c ta -=
<° I
... es
•3 2 o 5!
t/j
-a
<*j S S ^ » S ä .op-g 'S
§2 B-fr-S ^-2>E o
2 g 6ÓT3 R g o CS u ^
Î=fl2| » u« JJ - S fi « s" i»
c c c 2 I £ Bi p: H > u 2 M I a S <S 2 c-. £ e. £ =..u 2
S- o o o. oo o. oo a oo
3 \£3 'Z3 3 'S \C "1-3 \3 w 3 ä •— 'S
s , * * « « S „ 2 * .2 tS ü ' S 2 S O C - U « 2 ^ o 'ü, ü - e
« 3 .2 o .a « «3°Ô.fciM « 3 ° Ô 5 . b « o 3 ° ô ' c S S S
süfcüt, Q s u PHU CL,Q g'ufcUtti&.Q S u ï u o ^ E O
"J 'S u
I S2 -s . a-a'
.a_« .g
J3
«C »S « h e •§ p. CL, S
§§ »-§•§ 2 o£ ^ c £
' S <£ ^ "3 3 ^ S a> . 2 g „• „• Ä M O .g ö
*•* ^:s -si fif | I l i s i tj "£3| -Og

g g.-8l-S H -gî^S ^ | | . i T U .S !
g J5 J : o 5 » > M ' 5 - ï » M» 3 N % W P U'S* * ? 3(N.m
cc cc D- O O
a .2.2 a oo
s 3 3 ü 'T3 \C
O a>
0 >>
o o .« O§ s,>i 'a gs
l-i T 1 •"• •—
T» H »2 ? r h t» S O JD .C
3 3
Cultivai
" Kl l - •£ 3 . * « <« 3 3
mileSali
Flower'
« > CJ 3 3 3 an c/l
S . «. h5 i 3s -a
S 0SO ^D g
. DS,
c J3 c o a a
Rem;
Culti
Habi
lishei
Colo
Flow
Date
First
_o
O o u k u .i; s ^ 3 .2 O H ÜH Q B B BüEu ei o>-Eû
H N
m ça w H
/4gn'c. C//i('v. WageningenPapers87-2 (1987) 71

3 ;w
" Q
3 o
S 09
O
't S § 3 £ § Ö-
6 3 I £ 25
S J 31 -g p, g
3 -- s- ,3 « 'S -
1 O E (Ut- p.
1 M
r c • v ° p
- ^ s >, s si .•ac •=
eu - a -a 53 «
gg § <£ .
P
a M»S ~
S
sa lïs
S ci
g-S
S «J
|
-3^ -S• S
-c
O
•a g û" « „- S CN H 5
§cJ3 g " !s « CA•> _:3H S0- ." 5 'o
t_>rl^ ** ej • . eet o - — "a 55 —<-û«
35 4i •? « w J2 » o- <« 2 S S u . 6 ^ 5
-0Eiag s gsssss E i n s lis* <«SS

c
S CC Ë 'c _ C C C C C C C C
u O O 3 E O. o o 0 0 . . 0 0 o o
C '-3 ' 3 « c 3 '3 '3 \3 'S X 'S '3 '3 '3
S
O e &c o.
d e ö ' S . g O e j c e i e e pJ
8 § e > u 5 c, & c & a ces - cee] o . u a cö 3 ec<3 o. o- cö ra
ir« o «S « u es^ j S So - ^3 ? a o ö- au So - u u a g TL,
o « a u.a ^ o - — a « .a
0
« c«
S ft.3 O43 «<» «« ÄË UTL fH e ü- n
o -O .2 Ë
<« -o&. 43
Q 25 tCn f c 4=Q o.Sc«
e Und
-° -2
ft,
5» I g COH
l I *& S ^ u S g ft,
„ |> 3 3 ft«
Q f, 3 3 £ | ft.
33
S &
O 3
G O.
o, O
T3 s c
M "O Bi) 'S O«
I <§ § 1s
« 4J* -S3 S3 ou 13 <^
Ë C cj ^ e
'3< S g 8 -5 0 2 . 8 (U S
43 "
»JX! -fi
O c3 tuë» ïï 3 2S *3 S S
=3 d
> > •— ce) tx, 1R jr t^ ft,
S
« C . .• 8 ° P.
8 Ë
fc P
^ O' ï ^ S f . I §1 -gI" "l 5 ^-
S ^ s ^ | g | 3 ™ 6 - 3 ^ § ^ 3 „- S o -
4= û « S <= .™ . es .-S . a « g u — es ta „• o -S S
. » H t t . o t t i i f l . S i » - CJ— S TtiziJH-'CL.tt.izi —C J _ H^f MO. Q c o - .
g G C
O O o o P. o
s &3
S C '3 '3 "En eö ed O V cd
' S O U 1
^ A °- v. -S .a .a a Si »- & SP - - Ä — _ -
c u H 1» a o3 3
O CÖ t . t- «<- 3 S
S > u 3 c o o. o. m M
Origi
Flow
Culti
Ixposi
2 g ° c g M a s i g 2 * o « 3
Date
Colo
Year
First
c Ë .2 .S ^ . 2 0 S » ' " . Ü «
s
Ü | , 3 ° 0 . 3 ej
eü
ai 13 3 Ü B . U t>50iO^tt.Q «c«Utt.Utt. Q
U tt. tu p
72 /4gWc.t7n/v.WageningenPapers87-2(1987)
o • T3
C C
cd l>
^ E
Ö E
ä o
c «S
C- k ai
B o U ° -y £
oä
3 =5 •a
5
^ o
<_> . E ,
ä . 'M J2
vi
** ty
o u
O "s _•
et o
S ra
SP 11 3
X
3
a «o
CHH *J
><X
b —
C
o a: o B —
o -«, a I-I
. o 2 •
OH 0- cd
-• 1/3 •O o 3 >>
" 60 IS
'c3 *j
tu O co-a 2 .2 B 13
° E" S « e CC fl) -i-» .•*".
E s.3 •o -' S f o _3 •s S Ä «
C3 5
e> S o "o «- 2 s ~' o
S !T a <c -2
« T S ,-3 <L>
(U (D l—I 3 c/i
M W . O = ts2 C
O
3 -M IJ . . en W
- , ON - , • <N
o ° 'S • ' = ?!u2 •J
2 Q I jj a$ -'O
S 35 2'S ô
eitó(3c2
S 2 _o _o
e e
o o
 X £3 B>>-e2 " XI3 S x3 H3o e
.8 O >> -°
3 -2
8 £ Is o >">
rV C
3 S
OH O. ^ co —en c a o.
_3
^-
"o
O >
» «
> U 3
H M 5
o. o.
y a o, o. ° Sü *H o co u
2 - IH "cd ••= 3 ° o S
œ
o -sa Kg, .g s o w ft, Q
Ö
O S/u ft, u ei E Q
SS, .§ * O trt
ft,
CU Q ft, ft, ft. E tb
oM EO
ft.
3
a
3
O
M XI
o -3 O
S § E
e -o
£ « E cd
3 00
<D 3
«5.2P o o cd O o 3 u
OH o, O OH o. c E
. • -o > cu cd
o 2 J7" ^H
£ 2 CJ *H cj
C/5
2
ö *H ^
?H ft,
3 . cd
< 'S O, 3 =3
t-< ES 5S
o, E{ 3 <o e3
* E- 2
. O 3 I
o _h c t- -c •2 •« Q o •2 m
S3 2 C i^l cd ^ cfl ^t ^^ 4J 'S o^
<C O 1) 00
<lu2 JJ < . 0 . I
0-
O 32 Ï 2 Q E. r3i ?2:TT W^ cd r-^
o. S^ u —
c c
9 c
o o 3 o o o. O O O S
O
s 3 *^ w
cd a cd cd
o u
o u cd cd
S a .sX i CA „ .a .a 3
CL w
'S (X o o
o g xi xi B XI -o X X E X) xi
3
-2 >• 3 2 S 3 S >>2 2
O. O. O, =*> 2 2 OH OH
u p c o. o, < § CA * t ï ° to S ü-3 S3 OH O,
M SS .SS .2 « 0
° 'S s
û o M ft. Q °£ Eû oOX ED
i- cd
ü ÉQ o (^Ê' Û
ft, ft, ft. E
H4gr!'c.Univ. WageningenPapers87-2 (1987) 73
s
c 3
S
3 3
X>
-a T3
c J2 C
cd +2 ca
X s X
Ui vi
'S x>
SO 60 Ë
•3 r-
SO o
B •3 _3
X>
c oo cj c '3
:cd
T3
••m "3 Ü s S
a
3 o C t.
"o c*-i "o r n O
O C 3
> a C > ft. 'E.
O
53
XS
ca
ai
JÜ 'S. ej li Je (u t/3
Ui
O
'S TT
Vi
ej OO
'oo CJ 00
g 'S S « 'G
O
-
^"
CJ 0 0
ft. 3
"5b .SP 60 J3
S oo
C*1
c^. c - <N CU s cd
CN-
c K eu hJ c^. C^. o - sJ - H 3
C
Vi CN
i/5
OQ t/3 c - U
4>
OH
C
O
C
O O.
c C C E c c
o o P. O O
g O. _o _o
Hm'
rrou
Irou
3
cati
cati
1-1
S 'S rt S 3 3 a
pe
O
8.
pe
o
£ U >> M
o
—
x> Ö >>
ü
3
o c
ca
3 3
15 OO
L*
CD
a
O 'C
3)
ca S
o o
X>
E Ui
3 >,
'ist s H
O H O
05
3 3 M S c"- 3
zi 3 33 33
a s CS
> ca I-. i-i
3 O. 3 (H Cu 3
O, > cj 3 CL O. a
_o O o
CÖ
>
ej 3
Ui
s Ô
.3 a O.
Culti
Rem;
Flow
ormo
S
Culti
Flow
Colo
i- o 3cd'Só b tS S
First
Colo
'S > 9 22 "G 'E

S en
(-1 23 ,2^ Os.tl «s "o
o -o 3 Jj o 5 CJ . 3 cd
Û Ë cd tu
u. b-
o
Q U ft, U ft, Q o
ft. ft. 1
£
Oft.üftiO^&.Q
•o T3
ft)
> :ca
£ 60
o c
^« a 2
S .S
•S 3 •a s i J?
|>
cd e«
bO O
Cd r/i
DO O
o ft. 3
•g> _3 •s fe
^
c
•
Is "3
• * Ja
CJ . ej 'S.
60 J *
I OS
, u-i s o j j J3 SJ W Ja & «
60^
so a\ 3 ca .so
"a .S-...
•= 3 «5 .3 .3 .3 'C S S gî
! ( N CÄ CÜ JK-ÏÏ , <N C« ÏÏ _ ) c/3 —. —<
o a « ss
c c c
o E o. .2 -2
3 3 S ca
^3 o o S3 3 a a
:= 3 °* öJ9 3 TS 2 * Og S B
c
82 * o- s
• S U f c ü c< ft. Û § U ft. U a! ft, :q 3 J2 o B .3 ca
•g U ft. U a! ft. Q
o
ft.
l 3 o oi
gUft.Uft,û
a
ta
74 Àgric. Univ. WageningenPapers87-2 (1987)
O
t/3
u.
3 ^
8
o
ca > li. "o .S
« •* ö 8 S . S)
•S £ I a £ .s S
J g a-8. £ 2 8o
<N tz>Q >• o* — « P f c l i h S B i - U - _ * ha J U f , (s 3 DfcJ U 2

.8
». CC CEHSS Ö fi r-"e
Mg- OO g. o O 3 O. OO D. oo
I I & I I >,§£ s#|| fjg: I I l|& II

g3Hj!f l u l l i g -ïSIS .BillllS H H i l B ^
S OfcU I*te,û Ë Ute.Ü £ û< O ><EQ i u E u K E D 2 U E U £ ( 2 E Û
"o 3
o >< O <p
•o a "o S C/3
on »-
O
O 5 O U *
Ö 7f -_C
S -o « 3 c 5 -ö Vi £
B*d -ä & -e § 2 .a -3 2 .c
3 nm 3 « -S _. ~ B J 5 . ' Ä M
M
2 ™ •» .w r"5 « « 2 73 ta O
O 3
03 t*H
C O a C.
Aüllli p i l l s §iilltSBä Hill
• » u E u o ^ E û
a
;a Wî
I M U E U O ^ E Û
OJ
to O O

K 2 ° ai
-'S ^ o
•a ^ z3 a '5T
ge -su
za
° ui _> —
G (fl e -S3 3
^ o - ü c s J
S2?;5 -c > P „
§ — o 'S ca •c « f"«).«
S v. ° 3 '5 ^ . -2 > • ^.Ë >
r
.5 23 0-* .2
- 31S » "S M * f 2s l O
i OB§ S§ «i '3
IL> gü8 3fia°_3Ô
* 5§ .S
Ë «» 2 Ë 3•[° sÔ g. 5 t 5»«S
T2 n * ^ w £ T3 >ä *fi O *S ï i t-i nt O O hl "* H fi Js O . f i ™
S u E ü <* EQ SüEütó EO | « U f c« ücjtft,KUf c»SQ
fci«Q 5 u§ft,UUft,ft.UOft. O
S 5 ?<
o c 2 £ 3
g Ö 5 2 S
.£>
s s r -s 1 .s
T3
C
'S
3 1Ä
c
§1—1 «•?? •§ca .catt - a*R l
M 1
= " . i2
= ufits a-a-ll
ca S- .2 =a 3 g .
-73
S e a g o S ?
CA
SP ^0
'S
Ui
60
c 'S, Ü 6 o«-«
•3 00 Ü
0 •J S M o. ö C 5
•ca
to <N 13
'0
0 rn •a*S « ^ & I 2 S
> •O
«
u.
«J ft]
'M 'C -cf
v-i
.g
c-. '&
O
c^- CC C
00
V5 «•laJliSs «-l^läJfls »Ê «g«s ai§
e Ä fi C c fi a C
P, 0 & 0 0 a 0 0 0 - o o
3 _o \E>
0 a> "ca ca Pö cd«) 0<L> g] td » - O u rtcö
anum'
0 0
er Typ
var Gr
S 33
3
3 0. Cu '3
S.2 * 0 ts S Ö 3 0 0 g ...ca § 3 0 - 3 cag.gta S 3 o o* î 3
S 33 o i ca
ca
2 0 ft, O ft. 0 et U ft. U öS ft« O « O ft, U X O. ft, Q S CJ E O a E O
O Ö ? ? S
'" ^gn'c. t/ra'v. WageningenPapers87-2 (1987)
o
X
o.
.e
'I
•o
2 vi :
t/l
O f>
•CÏ 5 Û OD
O
-J "SÛTÎ 3 C^- .iï1 ^
•* ° c t g " S ê a | 'S^ -o*c
f-s-S f-gS'ils 1 ^ll^S 2 -äf% i|&5.

S5 S „ 5 S _ aa _ ç
g- .2.2 ' f g. .2 .2 g- §2 S- §c§ a §§
3
s3lf.ll l ü J f j l I "sliligis « l l l l ^ l i t i g e
gUfc.üfcQ oufcüait, D |iiüEuslEu f w ü E ü E Q | Ü E U I 2 E Q
O 3
- S
O. es
H «o °-
c > cd u > . ö
'S 5 <S o S V u «
•H * «s-f 8 8 =3 32 2 M-
I£
laissait £ § "s- £ ê £ <

gU | 8 g s a « « S «3-- ,g cj * £
g- e .2I g- g.2 g. gg a g§
i^H §1 sl-^l i^lij-ë'sJJ l i l l o i s SillES

/ïgn'c. Univ. WageningenPapers87-2 (1987) 77
c l-l
'u eu
T3
OJ
m
cd
"B,
T3
T3 t-i
 13 3 •o
C cd ö
o
dj
3
cu "cu S ^
.e cd
CU
OH
CU
>> e
'S
CU
O
H
>
C/S
CU
'S
"o
S
O 3
s ft .2 C*H
c
'•3 £
H
o
'C
o t/5 CU
H
ej Z 8 O 3 *^ CJ
CD
o <L>
t/3 S 3. w
ft S H c o 3
X!
Single.
3 ai C c -
Palepi
<S «J'S
O M 2 <N • r o •3 60 CU 'S)
Ä C XÎ ^ t . r - eu ca
cd
u
60
l-< I'S 00 e cd S
O -4.
(D e
5 . 3 es oo =x oo N ft. N2
2
cd
o- 03 c
2 o- c^> m
't/5
aa Q£
c c e c c o f
ft 8-
3
o
\£
o
\3 ß &. 3
o o
'.£3 ft
3 •2-2
3 •a g 2a
cd
o
cd
o 5" O cu
O. S
o
cd
o
O EL) cd
.8 . o
cd O CU c
*.,or>. 2 o 3- X So 33 0 >> <S O - e
cd <~ -* 3 cd ^ l-i
co
3
3 3 E
s
I*
cd
H LH i-4
3 cd
V3 2? CA Cd hH LH ;
c O O. ft 3 3 ,2 I-H
3 3 o. CJ 3
3 3 g ^ > » 3 I
> y 3 ft O. cd cu O. O.
fc 2 s ,2 'So 23 *^ °o o § O c a> S
S 3 ci a j o <
«&O s
JO CA V) O
CU
B 3 .2 O ' C ej . 3 ed a S 1 3 eu
5 .S cd B 3 .__ O cd . i j cd 3 3 ° O.
«Uft.OO><ft,Q U f c ü ï E Q Ü H Ü
cu
t/5
os: ft. Q a U ft. U X ft. Q
oUfcUB
o
^ £
U CJ
T3
•3
3
^
O.
•a •
cd J3 H M
2>
S P h" e«
S
E
3
&? ü V «
8. •9 S ä£ a X
. ^<
"U
S•§1
c j o t ; _j - • o a ft» 3 33 •s- ?
6 0f- . CN
.S < 3 £ cd 3 cd
J X i t/> , O ïï o Q ? Euïï
o, c C R
o .2 -2
oo p ju cd cd
3
O ca
C '3 ft
3
O CU 'S 'S
C .'Bb
2 r t'j o o
•5 -s 4 -«o>> *h 13 X i ,fe ^ft>
OTl H
en
_ 3 3 3
Cultivai
• * cd e-, u
Datepu
Firstpu
Colour
Flower
il > CU 3 OH & o s B O O. O.

Scent
'B J o s o O '3 S o e O 2 S°
g
c
c
3 O O .ij
U BH U ft. O
* S 3 .2 o
g O ft. U O> E O Ï 3 3 O
.. Ü LL. CJ
78 ^gr;c. [/«i'v.WageningenPapers87-2 (1987)

-à
3 VO
o
o
<
I/)
o
X
ÛH
Vi
«
"9 = s Ç3 «
. 3 2 c E g • " -- -
o s 6 jé S
I î ri 1 .a 8 . S |S i il2 c o^
, u oQ.. o
o .o2.2
-£ ü °uJ o« ä^ Rc .2
.u-s M-a e» u g o *-" c/> i
C F- S<TT
•5 . B 1uS,38 - SMr t>^ .ooà\S ;
•- -S-9 es'S J3&
rH °ô -5 .SLÇ53-S5;
'^ ta "ta oo ta oo
0
9 c
O O OH O
OU 'ca ?3 r/l P u M M O u - S , « « 5 .u „ „• gï g :sO>»
1 s wLS
sg^ gis
3S3,o ja .g 0 ^ t^H' »2 S33
'tu h.
l l E
^ c S 3 f t o- « S ï S ' s U â S ? f e S s ° ° - „>gr5'Ss°5.5.
^3°Ô.ë 5 » 3 0 0 S . S S «-BoôïS.âS | 3 5 ö S ™ 8 s S
-J
üö-Uti, O «unuaifcû Î U f c U O H D SufeUBiO^feû
-J
:W :ca
S g
s e
ta
C 3 ..
ea 3 m3
•c >- —
. o
S u o
-C r-
ûn _r- -C
<9 • =§
1 «so
o » S -
ta s
5M 5o Ê
o
> U ^ Ü
Su U <£ M . U M
2 c u «a a .S
°a^.SÏS£ 'se- ^.gS
c ta R" r7" u ^* ^;
•gll SS _r_ r c«t/3=•—— îl ——
S S
- -- I «** =>- W
M.«. 2ïf^c O^É<r>
u ob
u 2« _; M^> "ob-a
u u SPÖS 'u
u ^« tb •= 3>
u 5u'52a- u
M! Ä£tL,H5u2 ^ K o- w <^.U 2 - a £ £ m e2 - a££« ë2
0
5 _ c c 2 o c c e
9/ 0 0 a o o -g o. 0 0 o. 0 0
<2Ê g|| <2£. s s g g «o>> g 1 op g g

" Î . 2 S O C P - S S s « S O 'ob s " » S'-SSoc-s! u ? '-3 S O Q S U
^ 'T1 !r> îr>
j4gric. Univ. WageningenPapers87-2 (1987) 79

£ g
b 'H.
Ö ni J*;
. PH O £ U- PH
Î R l - O ) O > I- U U i < D <L)
o a> F* u _2 <u F .2= <u g S
S& £ S» B- £ g S- £ £
o BÖ E b -C r - 'S r - * - ^ ö • * <-- 'S r - u c •* S1"- G—

ffl
• S •=• M « C3 oo . 3 00 . * CS t - S 00 fî 00 g •- r -, ° ° ^ ' 2 " ï r'ï ° °
Sf ei'* & 0 -21 & eI

! & §§
-3° ô S £? S .ia RI | 15 Ô 8'C ».S « S i 15 oc »i « S 3^ o i «

.§ut. u Q;O>-BHQ .Suü-uô^fcCi & "on-uoô-û aUB-un.0
-J
B -3
s I
c 3
£? 3
c
'S. 3
,0
1 a «
2 -a c c <5 S g
K 3 . tt.
a»
« ^ -a •5 \o
0) t« <D
2
D . . Ë 2 .C 3 3 « ' S ^ - O ^ =ï 'S
S.SfS£3£ s « .3 -S SR S S S?-3 3 SS
ce cc c c _ eg
O- O O O. O O D . O O D . 0 0
3 C'S ' S 3 e *3 '-B 3 -s 'S 3 C ' 3 'S
§8. :§ss . . 8 g. ^ :&s s s g, g s s g. M:& s .§
££ SS22 g | .o^ ëisï o^r 3 I ££ 2 ë | s
g s s 1 's B.a g> I | « ö s s ' s ä ä rB l b s g. a . I g s s 's a &
illi-|g|s 1| élli^sil U l l i I |U1JJI|
-J J NJ NJ -J HJ
80 /Igrf'c. Univ. WageningenPapers87-2 (1987)

'^ 'O' .J* *1>
-2 H oo JS t!
:ca
"o -^ c b
o Ü > u 2
^ <° O >» e
S M c .• _3
S . S«
•J <D S S
b
Qa p ~
.5 £ 'C •a
a c
c u m . — cS- o a_>
o 3 u u U U S o 3
t4- 1/3
•" o e " CoM J
3
£ u 3 •= b »S i-. g
O
J C/1 £
1 J3 ' I 1 2 3° -3
O
• fra> ^ I |
"C
0. et)
•û Ô 5^ S «o
K -g
3 „
« y e
ä™ «n cd Sa J si ng ' S»?o m SS ^
« (J
O. o •§ «j ° S .§ 3 *3
o O djj u-, < %! 8 -„; g 3 ,2 > -O 3
J o
££,§*»3"3 c l g s »üä *ïfl ' S • *
CQ — E S «s .s ? sa sas «s g
M H K J C u J m c ^ U - S Î"0Q— ï r ;> <J — t t a f c J O c - c--c--bU—
S s
c c 3 C
c e .c2 O
e-2 c e s C C S c e
g. O O O O O . S D . O O 3 O. O O
3 C'S ' 3 \S '-S ' 3 'B S 3 \3 'S c 3 'S '-3
„ p u
* s a
-rîi ca cö
n •- ° a
-^
o o cdcd o
° - « .«u rï o a
td « O u
2 ti o.
cd
o
cd •— O a>
o M i - a ,
cd cd
S u
DO
M^H «OÎ323 s?3O . eC s s33 22 g sis 2?H «2 2 U
5 b sis
CL, (U c O. Q.
llïs s f o ä ä | '3 i l ao gco .*--«3 (U-w a i g. g. « l ö s l ä ä | l s s ä ä
</}
ro
Ï 2 * 5 g §> « » £ « o8 2 3
S" 1/5
S b Q3 g SS | 2 | 5 ES S § H g .2 S S
•a 3 .2 o u C u . s c a •= o S s " « 5 , 3 « S 3 . 2 0 B 3 M S 3 o o "• «
gOb U «i O >
o. T3C- o * Sj - o " .
u ca . a H
3
5 5
3 , ; ' S * ^ S ^ ,2
" « § I § §I «i-c E f c
2
S 3
& ^ a - . » j ^^3 .—I 3C 2« ^ - d O
' 2 «i S «.
C
^ , S ^ , C S „ C C „ o
É
c
0 . 0 0 0 . 0 0 o. 0 0 0 . S O
g -a - a 3 s s 3 -3 -a ^ 3 c 2 's
p uÇ. c .u
d .u
c Ö -„ ^ Ou. u c .0
a d .un J ^1^ o.^ O u .a...a.. c d3sr.baL,o<
a î O o _ .2? .8 'C}.nïc3
1- ^ .. r i r t_ /-. .. . i ^. on o 0
ï > S 3 »• ^ « > S 3 a
? î , S .S S 3 «O.O. j S > U 3 „ S C O C O . 3if
! s o q .= * 3 3 o o i « 2 ^ 3 0 - 3 g ÏÏ3 -S 3 o o g S - C S Ü «
S U b U b Q S U b U b Q 1 / U f c U aibQ g U b U we< 0 > - b Q
H !r> Sri ïr1
^4gr/c. tVm'v.WageningenPapers87-2 (1987) 81

2, 3
3 3 3
Xi
•2 =5S c
.s> • s & a
Or-l O >°^ H^ ^
-ïïlli 1 2Iî!ill „Jlïi filial

3 "S ~ 3 .2 -2 S- .2 -2 ^ OO
° 8. 8 8 -„ s2 g. s s sa « <5 o § u s«
£ H "§ 3 2" ^ " ^ Ja? 1 S ^ Ê . 2 S ^ ^ ^ . 2 2
JSSa a -S •§ g s ü "a g. EL 3 « - ^ s g « « £ . «a • £ 3 =
•è% o~ u & §*« s og - S" 3 ^ > g S? S" I .2 g = « °-
3 ° ô . 2 s s 53 oI I s s 13 I •§ si s s I -§ §-S ß2 S
UfcOfc Q | | O E Ô « E Q | O E U * Ê «S | u Ê a * È o
s a «
3 g 
^
S 3 -3 \o
S c c t--
& "5
• B <2 ^ w jo es „- « >: S. a* BU
-p& 11 g -2 I ', I .2 & §| g. §§

|l& I l §.J l | « l î ja 1.1 ig. ^ i l î
|3i3iis ji i i ins mm mma
82 /Igr/c. Univ. Wageningen Papers 87-2 (1987)
3
O. On
•a §
•S 2
3 •=
HÄ U
CQ c
° .2
8c
o
•s
-a ö
'3
a G
00 u
e S- Ë iS ^ S S H& Ë
'C C . — 5
2 oo£ 3
§1 ° -s . ö <=2 • •§ s -g . ^
o i : s c H-T »5 u . m «j . u < J2 * 9 ö -y <«
j?.3»S E'S 52 3 SS
<-•-> t / l -S-S^S .S.SS5S .S| o1 1 £ « £
c c c c _ c c c c
— OO „ O. OO DH 0 0 -.. Q> o 2
3 c -H= - a 3 .. •-= -a -_ 3 _. -H3 -H= .3 3 e^ «
OU 'B, cd C8 M ou cues S o u caçd g o u -g.
o .o .o U ^ H .£> «o Q T H H S -O -S O u h S -2°
Ê l _ , h_ _ô ä _
ä
3 sgssää giss-Sää gg s s-S J ' s
•S3io!ÏS'Co.iïs À3 ^ 'S î S Â3 ó o Sî « •§3 .2ô s 'Efi -~ 3
« U E u M l ï i O r - f c . û « U CLHU fc. û « U H H O B Ü I H D « O UH U tó O >- ö- O
-C
Oi
00
s
"2
-9 §
•" -3
H6
M cd
O
3c3 j .5« C" S 'B
O „ . h oH u • - ' " O
00 >- U u O -3 • oo ir* u Tlcz:U
• K C U U S S ™ . 2 O
o.
00 u a 2g
S'a- £ s s ^ s- £ gfc § £
& I s . -s
« c *r
g - ^ i « ^ m -s
S S -o 5 u «
. .5-a s . -s
Ü 2 <" H S M
M
o u3<-M.m \. 0%j~waùPin.ô ">•3 £?-S 2 vo
> u .rs r- •£ t-- S >.-TÎ3 s b * ^ ^ ^ " C - C J T H X ^ 1* ^~
•g cv- H >c - d r<~
n i es
o <x « oo
- U — aa \ ot?Hi f 'fSl , S
f 1H S
J <»
o r*i
i - »U - Ho vagfe:
- t1 o,2l rS
> i Jn' °0° » i ^ U «
ta c c U c c Z ^ c e
» O , O O M O H O O I B O H O O
Jj 3 c 'S 'H3 «! 3 C *3 'H3 «j 3 C 'S P
s s g. u:agg s s s. ^:§» s.g s u .:§>.§.§
ö°£ »osiss 5°^ »2823 e ï ^ M oëSS
llJlll'ä.lä I U I §1^8.11l i l l l f | . l |
«UO-UOiO^fcHÛ « U H Ü t)5 BÄO >-fc.O £ Uft.U 0< O >- U. Q
Agric. Univ. Wageningen Papers 87-2 (1987) 83

1
*s& î'1
5 > 2 g. §g
^_ S H
| S OU s M
g SS u. Ö Ö H 2 « = C - c ^
»•a a .a a g .a » u s\ ^ >- c .a s
-
Û J . . . . cj ca ** T ^ k CJ ta a j u " ^ * M u ta
& 0 0 . C g c . 3 U'C B . p,o „ ^ Sg e .
ai «^ ~S «* . <->g
~ ft
o *«=„S •§.« D,o S : 2 c a.j S j ) . 2 - ö
O ft % J= 5 P ? ? 3 < N ^ : r n ^ Ô w 2 <" 2 ""> ^j ^> o a u u B û £ ' i 2 < N ^ : ' ^
c c
[ f > i r 9 - C r ï - 1 " S o o r S o o c •- -•= -C i . ca oo =1, oo ,* i f - O « 1 Ç n « » ,1 »
e
-„ „ e c 3 „ c e c e
9 ft O O o ft O O P . 0 0
.5 3 C 'w 'w CL 3 C '-C '-S ".. 3 C *-S '-H
• E P S -n, a s _ p u • a n s J o » -a. ca es
Sg g S „ISoâft gi ö3 ^ S ' S f t f t ggSS 1 loâft

•e 3 ,2 o S s C ü .s a -e 3 .2 o B c 8 b a -a 3 J3 o S c S 2 a
| O t t . U M t t i O > < f c û « U f c U a!0>-li.O • U [i, O ci O >H [ i , O
o g o
S G "O
.2 c fi
ja c
.2 s
0 0 ^
CH c
2 D . "n . 2
*-J M. u
J -O 'S
C ca ca W es ^» •3£
W — .** ;>, c i =3 o S
=b 2 * . - H < >, S * t ".
ffi
S -a =3 g Sä -o ö g .i= S
3 o •& tu i- i= v e ? J= o ; „ o S H
£ & §« 3
a s 1 ic e l i s c§s > o . & . §a l- ia§r a u
^
ca c c . f l o u e s 3 3 M u Ü
P -* 2 ^2? ^ ^ ^ b . e u " • q> o ^ S P s 2 g « 2 ' f i ^ i o
•s «,£11 3 - si s l-s^^s I l g s ä ^ s ;sss8
•5 « h fcoJc-U - 5! s * « J U . « U - vDH&HfcnMucac« —U — "
S A « c g i s ^ a ^ e s c e
1 S& c -S •§ | | S g« e.0 ^ ft o o -s
a S ? ? | 8 | | J! J! 3 ? ^ « U aÇ^ a 2o|| S
| i ^ ö S S o a f t 1 a s g f e s l äft s ? S S J goää g
I I s ^ a j s s g i I 2 * ° g ss 1 3 | ° 1 B »sss g
•o - o 3 . 2 o E i u . c c a -a -a « 3 o o B s=a « 3 .2 o ?4 «3 'CS'-ca -o
« « O P H U O ^ S L - Q « « « Ü fc. U C* B-Q « U f c ü » ) * 0 > H & H Û I
84 v4gn'c. [/n/v. Wageningen Papers87-2 (1987)

ob ~•
5 É S 2 C
•a
pa c o o S cM O
J3
•O O (U O
(-1
O >> e
 2 0
0 a. E0
a •a
s S
cd
2 ni
.; - "I 1 1 1 &§•§«•§ « I J S •a
ai •a
eu
"
U> .^s 3 <N • ro ?• ? f5 o «s °* so • f> J
- u
'« S » S 2
2 s g o o
c e , ofe co o 5 5
•& 9 .2 Ë ç g ^ 0 0 c - 3 - 5
a ê ' w y u es ra g M « s
• ö ^ O n C d i i - t T l -ri Tl Tl >wÔr>l?ï1uirrt
S S
_>,
u
c <^
G .3
»8 2 8 8 e
o ra o g cd cö *-»
sSfc 13 ö &. ö- ~ c
G G . « . </) ÏT ,« . ,, . (U 12
" o g c G " c : ^ c S c o r a
pfl|c|l 1 j^| 1 PI
OD.Ä2'a5?P03<N^:t^ o o « S1" ^ « ' M ^ 00 3 «r> . «o f" f"
£•%'•% £ « r 2 S "S SS^ SS > « 53 ^ SSr ^ SS .3 .5 S3•§ SS * SS 8 8
^ H 3 S h a f c J M - ü - « h o . f c ï i - U H ( N M t J i f l - ü ï - " "
"ca
e
S ~ 5 „ c c _ c o o
00. o .0 __ o, o . o p . o .0 ts
| 2 g, : | g g | 2 E, -als l i g . -al 1 v 1
E? f-? » 0 S 2 3 ( g o ^ „ ë 3 3 3 5 Ç Ê" - s ' S z ü -S o
S > u 3 « S 5 ° ^ °< c > " 3 « c o o . o . S > u 3 ä c o a o , «
Sags ii-as-as 2 Ï Ó - a S - s S s g i l f a s s u •§
• c o ^ o 8 e C u .a * • a o O o Ö c S . a « • S 3 ^ 3 u ' C u . a c 3 -a -o
x
- ' t t , ü M B ! Û > - 1 I . Û 5 U tt. ü Bi O >H u, Q « u BH O Oi O ^ ft, Q « *
Agric. Univ. WageningenPapers87-2(1987) 85

r- TO
c/l <-<
s ^
XI O
•c x:
ffâl
cd t - *•»
ca &
LH
ca CJ
^ • °oT- C
3 5
Ä 'S
c
singL
mme
with
jj a
• 0 8
ta.-S
,3 « Di
§ CO
fO >,
<*-<'-•» 3
CS t-l — (
C u ^ c cd cd
O
2 >
.a «
0
c •d
u °
 0 c C 1-
" -o
.H ö •o 3
bright
Cat. S
'2 " • Si-'S -<*> "oô 00 0 "50 2P 0
1930
g e3 C
<N Ü r^ 03 c/3
î= <N <*> SU J O c- ü ï
'S C a
n
n
•S 3 a -S c-a e ft O 0 Oi 0 0
Ç 3 3
IS ft oo
3 O >cat.^-£c3<-.33 S g^ h J< 3 3 -c S \ ] _ -* 3 3 'C
5 5.i 5 a «.S ° ft ft 'C > » 3 S a a S > Ö3 £ ft ft 3
fi a>
5 5, 3 £ o S -c g.h s £f3 ° o S 2 s £ f 3 ° o u ^ =a rrt 00
| 00Ütt.ü BA O >•fcQ « ü f c ü « £ 0 « U E U (2 E u
s O S
â 51
S, 3°
S ^ b eu
ft u cd .
OS * S 8
£ K U cj ÇJ çd
y §2 £ 1 £ -as
-c .SP U, h " o n- o .S >
= a . M u " B T3 =3 c E?r i
— . 002 c S £2 £ « £2 rh^
«111 £ t I t . o<
«g11 * 11 h 1 iz
. ; So « » « ' S S1 •* 'S 3 "
M - s $ > ; J S "3> J - R o ^ 3 ^ oo S c ^
.9 .S •- c « «s c "S SC; 4 c -5 «<^ 5 « «
& «Il & fl !! g« §§ 1 § §

9 a ft
G ÎT ï-i "•-" *. S .55
* 2
, î S.
M" . :.S>
. SS
* US U c 2
CS8.
f U, : |S S § I l
. £ - 1111 "%
» P e l s ao" P
> . aS
s5i =l 7 «Ofî
Ä O S s§3i oUi
s K 2-s - û . 5.
; ?•ü .•: Ss .JI .5
Ö "ë ™
Jo~ f t _S >. 0 «3 5
qöo5f tSf ot f t fStc ' ft^ & g i f t
- _ _ o S• = « • £ « S > 3 ° 0 - C g . ë « g î o v ï S S ^ 'S I ov. ca
cd
s
86 ^gr/c. C/n/v. WageningenPapers87-2 (1987)
<N a
•* 43
o *
•g •o
« -3
c
'5.
o
43
60
_
™ S o •
o c S
m 43
S" £ 43 o 3 H
o
S *43 1g
- S
* S u g
1 | °
«
c
I" äß
Ö ^ H
s
_i 'c t f c
*1 _
_i ^ 4 ^ F,
»„ 0
S
O . V,
j j >. S
-S . 43
Jj <-> . 2 —.
P
î> S 2 ""> w £> » O S S ' " . Mo .j 2 oo43 j< fc >. js

.s5-assess -a 3 ^ I'S; 8 . a j i Sä . S . Ï . S | ^
cc ce c e _ cc
„ OH O O -,, a oo PH OO CU O O
fa. 3 C '-3 w 33 3 \£3 '-3 3 '3 '3 3 '3 ' 3
.1} O u "C* cö cö *r O o aj rt „ O u cd ai O u cö e3
ï ^ D . 2P o o C t - D - o o \- >- o , o o b O. O O
- ~ > 0 3 C O D , a ts > «J 3 £ O. Q< cd ." > o 3 CUCU > <D 3 „ c a

. £ ' 3 ^ O 'Zn »- +-* U 75 \ 3 ^ O Ö *-* O 'S 3 -~ > o — O " — ' 3 ^ 0 'r^ * J ' O
X > . 3 3 : C . - 5 0 Ü K * W W B 43 £ 45 £3 « w C3 "32 43 5; —3 — -w n ^ ; ^ 4JW3-W
» s ^ o ' C u s « S 3 5 O B S S CQ J 3 ^ o . 3 « 4* 3 .9, o cd.« cd
«UfcUO><ii.Q « U EL, O OSU. Q ^ «| U tt, O fc Q S ü f c U ŒU.Q
1- ^ §
^ u g >i
43 43 S U
S » ^ O
£. 'E O t/3
g?"0- S "3
3
S o 3
cd 4 - .. 3
^ .S O 3
5 S 8 O
X
"3)U. 43 Sc ci
8
^ 3 § | 8 ^ » S ? -S
2Ï& &e- H .^h§2 U
,2 c l 2 i o
-
"§
O S g S - O . ™ -
^ •4
S • 3 32 - r ?2Fo
_ 7 ->
! f l J; 5er scd o
>; »O O
& c
• - -o „ « •'S ,3 - o r l i J c rS
e 43 2 -SP43 <~- 'S <•- c -2 S S <"- c c M g 1 S °° «3 S"
. B3- >
- . J3
43 Î.
E« «» » O
00 J3 OOO -3 c
.5 cd
d cd™
ca co (j,
O ..33. 3
« . ._
- »J
O ra « ov
0 \^ 2 » o
P o\
'
43 ü
CC CC CB2 • „ CC
o- o o - _ D . 0 0 g, o o ™ â & ° 2
3 C 'S '-3 S 3 s 'S „ 3 c '3 S 2 _ 3 'öS
2g, 'SiSS 12g, S3 § 2 g , -agS-ë S 2 " , gS
j O f î » S "S S 3 i Ç f ï - 3 3 !»Ê - s 'S33g J <2 ^. 3 3
5,2 * ° 1 » 3 S 2 s S ^ Ë S S s 2 ^ g a s 3 S i g | s ^ ° a s
Ï 3 5 o iî 'Co.sca S 3 „ o o .3 s E 3 _ o e C «i .s « d t 3 5 o .3 j j
«UfcUai 0>-ü.Q «UtL,üft!fc<Q « U H Ü o a O ^ f c D e . â U f c U f c Û
^4gr('c.t/«î'v. WageningenPapers87-2 (1987) 87
O "m
ja
-..<
H
CO
•S C ^ =' M
"c O. O c "° 2
a £ . 8 8 1S >
3 g .sa .52 £> -S — ö Ä CO <D
'sX-i-^ä'l S^§§*„ . 'I I I I.

§3 g.I'S s | | f •&§£.§ s f jj. 2§§£
U U O J U U ( D 3 3 « i j - ^ o & t t , 3 o._- -o .2.a -3
C
•o-a-o-o-a-occ "^ -a .2 * S J= -CS. — ^ Ç w 2
M
U M
U aÜ j M
U aU i c
U ä ÜO ÜO ,g ^ C O' 13 O <L> >>. 2 «J — O J c<3
S S S S S S S S « 0.j< u «a « i« 5 3 Ëoa 3 « 3 ^ n«
r - r"c- r - r - r - r - r - f -
0 M U i 4 » u 2 « -n * 3CS.OO g .tJ w S X • «N
00 S 3 3 22 MÄ CC G3
£3 o. oo 3 o. o o' o. oo
•3 i-. „ *5 3 C S 'S 60 j 3 \S \ 3 3 \3' 3
chon'
Pout
aguie
guier
«
tins'
's r l 8. .»SS | * S S . S§ 88. SS

:h'
El'
e'
sah £ £ •» o S 2 2 « £ O£ = 3 g O£ B 22
2 « î | 3 Sfflffl
-> p- 0- o_ Û_ o: . .
s
#çu > U 3 s . s ° a ? - s §.£ S 2 £- ^ S.ï S 3 s aD
-
*5c5
Ë S Ë S S S o o o S * .2 B ï S S S « ' à S ^ S S ïï £ * .2 H Su
S2SSSSS2 S U3 f.2c OU 5C< C0 >u - s!U C.SÛ C » C 3 .2 O 3 «
» Ü H . U H D
C 3 .2 O o
»UfcUtó
3 cd
feu
2 S S
.S -fi < K
0\n *
M
os g
o
J3
H
.Ä O S
ccj aj cj ^3
V. >
CH
co , C n ^ M . 3 w f ) 3 .c .. „• .
o I -S-Äl Ö.• 8'S J "« "e -» -i i «g •§sÄ

1-8 s 8' 1 ï. S ^ I = .g " .g gë 1 1 1 - §° ,°
u
.5 -O G ET >s •3 S2" •OT3flHT)-0'0'OT)5t)T3'0-0
ß « 2 <J u s£ G-3 .S?5 3 g-£- ( u u - u ^ u u u u u g ü u n u

• ^ ^ - S s=5 ,£> r ^ O S • • S ' 5 ' C S ^ Ä O S u U U ' < L > < L > < D Ü < D (1 ai 1) Il
_ 3 .S
g- .2 -2 g g- .2.2 « « a 5 a ge
, : 0 )
o u •§, 'S 'S « o u M ' S - < U ^ - Z . i « ' S i 3 2
8
-Z^H? 3= 22'§33 23 3 %V£
2 ~ H « « S3 I g|s 1e'Ss I ^'S"«SJ« gS
3 3 ï 8 "ë*K § fi -
SOS J g o â g . ES s s-3 s.â ^ I ö ö ö o a l l i i - ä 2
„ -a S o lE'ëibS" <;-S&oc «« u 2« , > U O U « « Ü U Ü I >

r~ 'S ^ P~ <~ i~
<N . . —< rs (N rM
w1 ^ O ^ Ä* "^
u ^ \ö u aj u
'3 —T J5 'S 'ü '5
o T3 o o o
t/2 g g t» M 60
2 * | 2 2 2
3 fel3 3 3
3 **H o 's "3 "3
o « « S 5 5
'S K B 'S \3 '3
BC §"9K BC BC
ü
ü T3 r^ 3 ^ u
« «JJ g « g M
C/3 M O '5? 60 C/3 C/3
S3 " o t"* , , C3 CÖ cfl
S 3 u £ "2 'S "°
i; O. rt T ; i- •- _: "E
.2 tM - Ö O . 2 P, 'S P.
ÜH 3 c fc, ti, j E
S u - 0 2 8 2 « u . S u
o 2 - H o . 2 2 ° o .c & o
S
.s.sa.tsp>§•&
PH — n. s
i «l -Q. s
> f lcgägf -cs - §•£
c-. <^. ^. On -H .Sc/3I l> OH*Ï -
.. T|- c/3
S3 .2.2 &§• s
••
2 e3 -2.2 «3 .2.2
r«P- a.S .SAO, ou M <seu u u BC,to. 80
H
ISÊ all --H j 11 *vp i l 1°£ «11
- - * P. gs Ü - » 2 ^ ° B «2 • ' S ^ s s -< 2 g ° s s ^
»i3.2Ou.Ü« »i3P Ou .SU »53P O.3.S «35 Oo,Ü«
Ä U BH Ü S* OH O i ü t - U K ! u. Q i ü i n ü n Q i ü f c U P i f c Q
.0 3 u o s s C * j3 js
3 o c/3 5 - ai 11 r- * 3 < . t ^
" U _. .S3e o, -s 2 •* o \ < * 1 ' *
C
g S §> fe s
.2 ËI 8 (L>
S ög ° - o«5 ° 'S o
; IÊ II >l< 1 si 1
d
^ Sc " â« g '= S^ 'S
"60 ^ O -3 G ° g. 2 'S ° °ft .O o
5 H . - O S « -.2 *< o ç u S
-
•o 'u o a< ij § ' S ' " S ü « f t J »
ë £ g J -â 'I «S 3 2^83
•o
2 . u ^S Ë
a | ^^ Sg<s
^ : . - sSf h
eSS - 24? Ss e s -
t/i u
.S I B | I l | 1 ° c°| a . l §y^ o -o o . s - = .3 » g o
' « 0 0
S .s x
W
0X! C 0 " 0 c e « C C
" 3
S -S -à o .2 .2 a §• .2.2 -s §•
g» S S Ä « ' S g O u « B S O U ranj
^ê I 'I ä , -I I «Saör& „ I l ïo,6 -^
^ U s | 5 S S Oo - u 3 C o 2 g | | «u BCsSog
« 3 5 'Ó i « «>, .Ü i? ^S,3^0Î3 .g « J 3 5 'O « .S S
iUfcUfc O £ « BH Q ^«UfcUa! u, Q | Ü P H Ü ( K &. O
^gn'e. Univ. Wageningen Papers 87-2 (1987) 89

M
•C 'M
M I
c
3 3
g S ca E
c °o
ffl
M.SO . - J Vi -S? •
ï '•& i? -S -S ^ g
S— • Z Z = • U
S O "1 u o Om u
w W
CJ j j «i ^ <D J$ .S >" <L> w
w> « '53 "^ - r* £ "53 ^ OÙ

J I S Ô S S «8 o gS -S> 3 S3 .S | «£
" Û B i J f f l c = S S e- c- CO C -H „ c « > U - c- Cfl c- c/î c- ( J —
c e e c c e c e
o, o o » 0 , 0 0 o. o o g. o ,o
3 •- -s £ 3 S '-S 3 '3 'S g " '" "
OO) n j c d O O U r t c ö O u c d r t o J)
so H »2 S £
S£| 3|
^ H S ^(C
6£ 22
%M ^H
el 2 5 2 2
-?•§:
§ js s a* S. S. g e E sä g. , ösää -« 2 s a g's S. S.
S 3 » Ö u " M C-tS 3 3 . 2 .3 C3 5 3 . 2 O .c « S 3 . 2 O C U .= =$
=ü E u a i E O a« . S u m O ,ï u n. ü n.û SUÖ-UO^PHO
S zz z z z
2 u -S SS ego
w
E C> .2 W u T 3 O
3 n .S C 5 ft. £> NH . " "O C/5 fe
t»
£& e
p& ââ ï . 11 IJS 1.1 'sJU .1.1I
3 S ft O, O c O. CX s > » 3 S O-O. S > U 3
s | s» ^ 2 v, a M •-> > ~ ~ - "> «>•«*«
o 53 .HS «>. •- Ä «
90 /Ign'c. Univ. Wageningen Papers 87-2 (1987)

© S o
— &H F
I 1 S
M a 's
M
"E. 2
ö -a ^
c - I
fa g •?§£ |°!
e
»« s PH
siSa -- :•
§ ESS••o * - sä *^-s
-
^§ at- .:S
Cs o
„111=1 I I I aillai J l l l l ä l J J Î J I
§• I l V .2.2 ^ I l g. §§ i a g§
If I l f a | | gjk U | | £ . f i l 1 | g. 11
il lift ifff iiii.îf Mgilift Ihï-J !
U)
C
(U
6
3
M . 42 .o
^ cu "2 -o
M
o BS =2 *j
? ï r 2 „• S ï K
als" 8 a .S» • £ §,.

•g^êl 3 I? "811"
«fr
> E? 2 £ ~ £ - »of 'S.s aô8 •« .*
m^(i.j^2u2 8 ^. =..<..« g 2 ~ K £ £m g 2
u, u -
! §" c .2.2 .a g- 2 .2 g §• o o a
p
e .:&!! 1 -«lu ! 1 i§& 11 1
îndiîU î nht t sîîBj ï ,lf, il
Îgn'c.t/m'v.WageningenPapers87-2(1987) 91
•O J
»H
r/3 UH
tt.
cd
M2 ô |H
cd 'S
o OÙ
c
<5 Cu cd kl -<u o
u
O o " ÜH
S CS
OH
% JO
E o
_C c h ° 3
oa O 'S
3 3« 3 •—• U «H —»
ai C3
« -2o 0«
< S co
2 cd 3 & S -a
Vigoi
Pope
. Cl
1975
»•o "3 tN
S u £? S
a. éd 3 • .. ed
'S t - S oo 0) » O -3 2 « ^
u
26 c^.
o2
f>- e>- c - tzi o - es- c^- u H :> &o
> J3 - S r-H
P- ^ > cd o o
- U
" '—
c c C c
O.
c
O O.
c c g
3 .2 .2 o.
3
_o
\£ 3 .2 -2 "<L>
t-i .2 -2
O >
TH H •»2 'H
o jo ja SO Z, H
^ C/3
-3 JO
o
T3
O >>
Z. H M 3 -o
<D
I-H
•=
o
= ,_ r- 2 o 2 2
Cd ta. U. <*H
3 9 5
cd - . .in 3 3 -< < iH • ^ 3 3 Ü O ^ « ^ , ^ , 2 ^ 3 3
> l) 3 •Sa o O. 0 . o > ÇU 3 Cd D.
cd i-<
> * " " 1) u 4 , > l > 3 g O O . - H
Colo
ü S * o g SS
53 .S2 ooi >pi Hä( Ls> .sc csd e - s s p ö - a ta e -
'E 1 3s _2
o cO sS s3 cd e c - 3 * » a s s s
Ü H . Ü B l O ' l I . U Q cd
0CS. U Ü- ü OH B- O •3 3 - 2 O 'C oj» . ü cd
0H OH
i O Ü H C J O ^ B - D
a.
o.
o
OH
8
e
cd
t.
tu
—
O
c°
S
H —
CJ
e
Ö 8 .H.
""•
CL. u S 13 cd £
u: e i & £
c
^
•—' 3 Td ÜH _r . «
àt o u ~ » ? 3 •* •*
Ï? .s.s i l . s jj s ^ JU »o
•ar.p: c°1gê §* P g? .3 ^ p gps^
•S S 2 ^ Ä E E , S « ( 3 2 C ! . . <%„• u 2 M B«S<3
c
9 „ co cc cc
O - O O D. O O D. O O 0 . 0 0
3 \s w 3 "irj w 3 c 'S w 3 --J - ^
P « J c d c d O u eded O u - r i c d c d ^ O u c d c d
C
O ,>> ~ •=> O >^ -3-3 0>> S —— M Q >< 3 -
t. H -g -= THH a -°-S i-H So-P-S .S i- H -° -S
. ?s s ä ä . S65 J ä& •i.EBäS'sS.ä fcïBaââ
J S 3 _ 2 o . t ! cd S 3 - 2 o 8 ö . 3 cd I 3 5 ö c 8 .b « 8 3 i5 O 3 «
« ü ft. ü ft, Q ^ O P H U W S . H.Q Ï Ü I H U O ^ I H D ï ü f c U i H U
0. OH 0 . 0 .
92 v4gric. C/«/v. Wageningen Papers 87-2 (1987)

.
I >, XI
o .2 %
o 2
^- 'S
00
**'S60
CU
ca X
O i-T CU
1) S
o cu O
S 3 o
-o
OJ —
';
CU
ccd lH
C M
60 ~ 60
.a <
CZ)
o
d, . o>^ •C& BH
fc- t/3
BH
.g
*C <
c/3 OH O
CU ^-i CU
p, S o o 3 •S 3 D
X3 Oc
3 O.C 2 ca c—
O .g
c/3 w ~ C/3 soi c Z S *
O X "rt 1 "•> c -3 ä « r-
S .2?E ca o o 'S ^ e « S .3. ca X r*-« o\
OH 2 u U U -
c/3 —
u-
c e
a
3 .2 .2
O CU #g cd ca ca ca
t a oo o o o o
O >> o o O
jA- HtH jgi : 33 32 XI Xi
> Sr> 3 S
IH cd «a
o. o. 1- l-i 3 " 2 3
O.
3
O.
o
•3 & O e 0-.5 S 3.SC! CU
;§> * CA W c/3
3 O O u 1 - OJ
U B. U Bi O >-
t - ca
0
^•§-ee
3 O Q aj 53
BH E0
c
'S.
XI
H xl
,60
'C
Xl
o . X
a C/3 *H
ca
ë*
OH
s-g ! BH
O ü
•o c 3 C •a gca^' :3
cd
J-i 3
OH X!
ca C3- C3
a Ë - 3 3
43
«3 o 3
X
ca cd
&o O
X
ca
3 o
6015 ^ 60 60 3
(U c/2
60 3 m Ô "" X I
O cÖ CU 3 (N CO
C/3
•Ü ca Jj X ! r - -= i X r- a> X r- r--
c ca '5 r^ ^ ca oo « oo o ca cd 00 t-i
ca OO ca oo
S BH i—I C/3 ~ O — C/3 c- OO t - BH
U- C/3
CM 'c/5 Q
u
e a G e
o \o£ a3 _o \£3
o o
S 'S
« ca o
I--
fl Sy cao 'S ca
I Ü r>^ o cu ÇU
2 O 23 33 ^CU 0 O o 3 2 ^ Ü t/3 o 'kH

O 33 33
3 ca V-. 3 3
Ü ca <*H
c o o. o, O" > S
C O o. O.
j ca ^H
« > g <cd^' rt
c O
CH
O. O.
c/3 o So Ui CU tH CU
2 xï 'S cd
C 
Vi
O
<
3 pi o
an
< „' <
s
J
cd
zo" CJ C/2
cd
L-i 'S P i-
cd" BH
"
.M £ (D
0) o cj 3 CU
pa (U -—
e B. I/î JO oa •3
'S. 2 G ^H C 3
. 3 • u •o O >» -O S CN
c - 3 M ta cd
. s3 •=
e « - ? OS
1 P .3 "S
• 03
O Os
u <S t f l &H J U -• « u2 E 2 Q£ U 2
o,
3 cd çd cd cd
O CJ * y cj „ <~
sO ^ ^ ' C -.3 3! >, O 3 O £• «O - O
2 O JO -O. O « •°3 'S e *- H
u 3 s3
p. 3
S Un co « ' S 3 3
« S ^ ^ •* c o o.* - O. > g 3 cd o. .!§.§.
co *- cj oa >-i cd 1H
G > CJ~ •-- co
Sao,
•— cd u. i-i t- i 2 S .2 Ë ° «S -IB .M _ Ô J' S ° O O3 -^« CJ
- 1 > g 3 cd 3 , 2 O co 3 cd 3 c j j t »c ü BH O B. Q
a 2u *E 5u e!
£ O >< E ! CJ U, CJ Oi BH Q c« E Q £ ft, 3
Ö 3 .2 O u
cd
o
u
<D O
fi'is
h- 60
J3 ai >> O, C T3
3
60 •S 2"§
c 6 O
fi • S ? c oa
cd 'C
'S I-I
ü cd (D 3 IH . 3 ^^
cd J3 cO «•al
CJ 00 t->o
c
•a
O CD 'I 's-g °-
cj cd s
cd 3
T3 09 • 60 O JS
"O E ! -O a "°
ai 3 eu -fi _
vi 5 3 O O )M ^ "
1-
cd 'S C : c 'IS T3
O « i-i C/3 •8 -S * JS
i- J-l cd *cd <ù • O cd
O <£ o t-H CU 0> •s s
o cd £ CU
«
•o * co •o
'1 .S O 3
S si
o S -o
& S MCU
eu G cd
i-i
cd
3
O 3
O co
•o -
cj
^ O
BH O
ai 1 1 J5
cd o ,
2§
Go
o. s I-H 2 .3
cd
t/5
.é_i cd •SUS
co tvi
<D ^ 'B.V «
3 Ow
O J 3 •M OO i- • OO W H ^ (U y O (U C 3^
S C S
PJ 00 C cd o 'S o^ i* >-> "i cd oo . n , oo
VO H io S & 1 H
cd o o
J l f l -
r- H '-J IZI f^ ( Ä oa BH J on — U — OH
.02
c c
S a
O. o S
o o.
o .o 3 c
3 3 'S 'S 3 o
O CJ ;so | g '60
b O- op ö ! •—.
o h- o. i-. *n
eu 'Ü -a Ü
s Ü £ Vi o co O o 3 3 c ». o o
ta ' ^ S «a 3 g o A I, u
cd I-I k,
' P, o . 3 O c o. ei CJ a o
O > CJ 3 cd Ü 3 „ .„ c
' I S as ^- +j u S
ulti
low
oio
v 2, c g
CU
s '»H eu . 3 _çd
3 3,
, S U i H Ü « i a ! 0 >eu' f. 3c cd
P
Cl » « CJ t 3 OO
» U H U
K § v, eu
00 0i O >-
U &, U C4 O >- B- Q CJ
0.
£ ft,
E
94 ^4grî'c. [/«i'v. WageningenPapers87-2 (1987)
•o
tl)
•o
• >
'5.
•5
•$ 3
a
•* c
ia
•o -o
-c 'I s
T3 O »
U —. pi
.2*2 «Sog § . - • 5
Sc14- o ï g f c o S e » " l
o r t u i ) o -»-• o >-« <D 5 33 2
u 3 o -o ° ° 3 e E ° 5 Ê « 3 c
s ooo g u b U "u g 'T m Q •§ T 3 l 3 = s
p.f 5 !^£ 5 I S S <S it *

h ^ O t i «-. * u •£ l/J iE ifc n" J) SÄ « m
ô
<- „ c c - - c e fi c e
« g- .g .2 S §• .2 .o « &, o .2
 s 3 5 § 'S £ =&S^-1Hfl's55
^ U OHU O >• UnQ | U B . u a ! 0 > < t H Û 2 2 2 S B Ü E O O ^ D
3 T3
c 0 2
o
60
T3 . «
? a
o
SU KB! £3
JS yaeg
•< ^2 ^ 3 -* X
"E. U w
P H CJ
S « £ -SP •
* §su §^11« §• i
J « ! M Ç O N tn V r« 3 . 3 c
3 g, u ^ -d c-; ;g .c
-C U
w
. - QM J -H ^ . c- c- Q 2 !S ^ « y £ ,Sfflc - r.Q M » f . U -
. i „ c c c e c e oc
$a .2-2 o. 2 - 2 2 - 2 * °2
!.§£. 11 s n 1 1 -e 11 s.I & _ l ' l l
E 2 S O - U § 2 S o S - s g o - s i2*5'a5SS
g 3.2 o,~ « e 3.2 otf .c M S1S. .a « s l ^ o C o b «
g U PHO a, Q SUfcUïfc O §w fc Q g O h ü O >- e- Q
55 AH OH b
^4gn'c. {ƒ/!!>. WageningenPapers87-2 (1987) 95

cj ^
T3 ca
CJ O
00 t-
•O - 3
o H
S ,j -9
'o. S g
o ~ »o
O ^ H 3 3§
•o 5 73 -a 2£ •- ft.
o g, 3 3
o . o „
oa 6« OD !-.
S T! ni -S
. a >bO5u Î-H
g1s.
l a og OH c a- ft. o
o. •"c S
« 2 Sg 3 8 ° 3 3
-c
J3
(L) (U on *s is c« ^ Ç- -M C/3
C8
ta M M S O J3 o 3 i n . vo
P. & ,2 . 73 > oo - ^ . t - ca oo .=1 oo
»-; t—i Ö « tu J3 r - -S t--
• 3 S .•" ca oo , rt . oo
CI t/5 95 ft. 1/3 —' U - .
c
O.
S •S o. 3 S
3 O O .2 -2 2 3
o e •*: '-H M 3
S p
'5b | | o. ;aj§
O O
&> 2 o x> Xi
'ÓO
=3=1 s5 u's 3 3
^3 a3 fiÖ
A
s-.
S-
M S O 2 - Û
3 3
o> •ï Ta «-• 3 3 ca Ct b w c o
•a S
<2 3 » 73
„ S e O R » .« m * -2 . 5 cj »
*
u i s gs i ca «
a wo.
SUH.U s e 's? es » " » U f c U « ! 0 > ' H P 2üB-ÜBiO>-B-U
Cu
OH
11
1-3
O.
c.
3
ca s
M
il.
3 'S J><
J3 -° X)
CD 73
S 3
£•-3 O » o ÎS
3 -t? — ca —I •O
s C
" S ca
5 S o c -75 ca o»
3 C
O | 5 -S o
• o * tS
.1-Si.. f i l 00 ^3 >
! C 73
u =u °ca un
l O
e s .
! 73 73 -3 ., 3 u o ^
3 i o . >s 3 " tu ^~ •-H ca w O 3 Ü
Q — <D ta .. 3 ; U 73
•a
s
ca
« -3 h ft,
ca » i • P
3 oo c a.73 "' : s? 3
a
c o
u o S8g S .9
O. 5ca 2 . g. u
s 3
! - 3 ID =- -ri 3 LH S, 3
3 00 73 3 r-1 - m
> O. o
|- C - a r - -g t - » u
S P.-3 S-C 3 73 J3 .«! S
ca o ca oo - ^ oo O p.
^ l ó j ö ^
c
S 3 S
O O
a .2 .2
S ' 3 '-g
00 ü o -ï
o «u
ca ca s ä O
2 o -= -° » 2 o £ £
# ta c - 3 ?
3 >> «'s s 3 ca t.
s o o. a
s >
o 'S _ S •S I •a
>. 3 o 75 3 » ' U 14 U . S »
M O fc U O > n, û U ft. ( (*• O >• ft. Q
96 Agric. Univ. Wageningen Papers 87-2 (1987)

1
3
c 1 ,£, o
o
J3
•<
3
a-
C
<* .2w
in
0\ M
.. e
c. s
s
2. o
IA .• o e c o
<L> 0) . S -*•* -*- - ^
LH CU
=: o g
'c g g 'S o o o M O JHM
O . g g 1-ro c c c B S O ewr, J j T3
t- in M M M
c 3 O OS S S E °Z - u •>: . CU
O
O IH
a— L,
3
t.
3
t-
3
S~
3 3
L-l
3
U.
o y 57 ö i-i >^ m C 2 Meu
eu
CO
eu
V3
eu
CO
eu
CO
eu eu
CO CO 2
•0
CU <%T3
V3 V3 CO CO CO CO
l -22 u tu
O O
C CU
c/i cd O O O O O .23 >> - s M 3 C ° ri >,
u eu O. C/î
VI
«S «S Cu OH OH CU CU CL.
O M O B aj -J"
a:
3
3
in
o. c g M 0 C 3
£ 0 2 o OO
01 a i 01 n ! H eu fa Êü2 00 H on t u QC u2 0
Û E z
o e e c c c B s C
-0 .2 .2 o. o P. .2 .2 s 0. 0 0
3 C seu 3 \£
G M M G ~co O eu
,3 .0.Sa - - B O O C t- Q 'Eb M M O
ÏH
eu
0.
M
eu
M
eu
•3 v. S o ü O *- 'C .S -S E
§ a |= u g S -° -S o o 3 3 s ÏMH H>
in
> 3 3
3
a M in M M- 3 S S 3 CL
IH IH ^ M M M 3 M 3 a 3 C O
eu 3
s « Ü O2 E E E O
e
2 S O 0 . ^3
co 0. a
0 3 s
_o V ) eu
3 .2 3 ~ _ ~ S 3 ^
'5b ^ 0 LH
M
<*- SL . 3 «*
2 c« ft. Q
%—• %—• U—i U-i
0 0 0 0
U—i U—i
0 0
«H 3
2 (J tu
°
3* £ Q BU Q
s
s U ft, U E Q
0. i-i ; * I H I H I H
L*
O . CL. O H O H CL. CL.
eu eu
S "2
'B.-3 Ë SM
_o
•Sa
M j3 „ 's Q o
o Q £•
J3 M M eu M
co J-
cïï °
S.a (*H "IT1
ÇH Q
ö l Q <U t 3 CU T3
tu
tu -M
tu cd
0 *J 0 tu co on üco
C C 1, vi < UI
s- ^ 3 T3 _3
O
-C o a •S
a
H3 eu
0
eu 3 t-T
O çu
CU
O
0 J3 S B
o 0 C cd
B g.-S
ra eu c/î
G
O
00 O.
cd
s- M
0 eu
3 JS
ui eu
3 J2
M eu
O. s *
J3 'M Û H Û H
E eu G ft, '>
S*Jft,
3 O 1-1
eu .S "> =* S ob O 3 3 G 3
cU
G
T3 C 0 O
eu M M (U J l V2 r-^ c-^
W5
s 3 3
O CO ^
M 00 ou
eu eu
LH
3 (N
onr^ -s £ . ô 0 en 5 O in
eu M 4S r - r- \o 0 bb 00 eu
& ja t« i-,
M O CÖ 00 M 00 G 0 M
er. ÇU . G 00
ói 00
J C on m on Ci U z PH
z ,g 00
V3
ft,
"eu
G G G G "G G G T3 c G
O O "n P. O O M _o O G _o O
O)
c 'Jï 3 .G M
M M O au a eu M M M
'5b O eu O,
M
eu _o 3 O eu ^
3 eu eS
eu
ce 0!-
0
a. - t O H CA
i-u
O 0 HO.0 h- ^3 3 Q
CO
3 3 Q S 3
in
3
e
wer'
;tpu
^S 3 3 L. 3 l i
>, 3a 3
> 3a 3
C« U-t l-H
M
>
our
C 0 O. O O G
_> a Cfl G
Ä S o B g tl
cd
eu
eu on O eu t/!
1Z!
O eu
g.3,2
- S 3 eu
0
tu 'C tu M «sa3U ft,
^ 3 s M c2
0
G LH M .U G
0 e/3
M
M U ft. U U 0 >- Q U ft, 0 Q E Q
on
E 00
eu
& OH

g §g
3 003
S (gffl 5
fe
%M ï
1 ^ il -a i
T3 'S r 3 ~0 . . o ° -
1- r _^ r _ fis. ,_ ||.|| U» e »
l i i SSrsi
«
•sg i ^ i'«i§-u:§:' •§o«' -so 1 1(ôi^J2
» M™
§ S<Ä 2pç
U ol si a Z " g-a .S
3 s
..g|>^ a i aa |>^ a |HSJS ^ S |
«JSSSÜ S888 ^ J IS 8 „m** „1213*35
«• § I ï I I S; I l s.& „I I
i& ï ï I I B s». I l 1 1 ! ,:!.!!
10EÔ*È û SPffSfS SÄE O S -güEoEEQ &ÔE ô i2<5 £ E û
os ai aï & &, oi eé as «
••ca
M
S
e
CÖ
8 8 8
ca
c T3 e
o . t-ï b
TO )-« i-U ^ ^ T
J
^ 3 « S| 3
"cd 5" ^ ^ "rt
"äo 3 ^ rn O M Jf^
Ü .S 'S * SS > 'S ca «
"» <z>es-c/> CN- O -H \oHffl"JU
cc „ c
& a-S-S 3 -S-
s& .:&ss sa s.
3 gs ^ o Mas s 1 2 * » e ts
«9,3°Ô'Cu.3i5 ~ 3 ° o S . 3
•§ <» <_> b u o ^ &. o a u u. u pi PH I
98 ^grc'c. C/m'v.WageningenPapers87-2 (1987)

C cd
C O. CÄ
8 O « « s .-S
I ä S s. 2 | |
W J
C ^ TO 5 w5 3
TO - r„ u - S C C^ ç
^ M O U _T . +- O e î0 H - - J . s t 1
aj -4-T
*>-*_Ö *J C « - - ^ g -g g o - w) g . ^-Î £ <N *§ 'C 13 *j ^
3
t__ o c Q c e c e e c
«S- ° ° H °" OO Q. O O D . o o
» 3 ta -B «3 .S „ 3 c '-3 *3 3 'S 'S 3 •= 'S
* "2 ^ 2823 | . 12Ê »2S23 S?£ 32 »£• » § S

• § 5 § s | o ä ä | 'f | gbs-ga 's ä ä •§*s s l a S. -e I s s'Saä
J 3 3 . 2 o C u . 3 . S O a) S 3 . 2 O B E U . H C S 5 3 .2 O C5. - es « 3 .2 O S 3 es
iUfcUO>"li.Q .2 o gUfcOûSO^B-Q oUfcU SCO SuEuaiËQ
o: ai (* « oi »
••ei
e»
c
3
3
ai .e 3
- ^ Xt
-g ajS S o r-cca
^
j»s 73
.: S ° 3 -• 73 S S u cd S ^ .So n
.I<§s 1 ! .s^! |^ i i .ïia>g

j j a ü « i» S £ £ „g g
„ S S , »ftft | S15 Ö S S. o. ä S & 0 * ''S "H * > s 3 - S a 5. ca
u S
2 * 5 g « SS ï s ° 2 M s S c g s s S « S3 2 g ° c ^ -
O)
Ufc(J<*0>"ü-Q Swüft-üfcu Ö <zT ü<Q .2 .H .a U li. U « B n, u
? Q^ 05 Bi X

U Oß
•S <
O
os
rt Ë 73
t.
0 e0 3
t u
0 3
«S S
C 5O S
ö u aiOuS s
u. c g
0 OH O
C
O
•a zÔ
u
a
0 y 0 CÖ
U
C os
.c
CA 0 "3?
SES -c 2 S'£• ^ 3 32
cd «U.«>
C— Ui O 0
C G
"^ O E
.S | oi S 5 3 O 3 u o s
O Vi . .S 73
S 0c
Palep
Single
Cat. S
O
1873
60 g 3 U
0
> G -O O.
cd
a <N
ce „ _ _, C_ a c c
Q. O O O O "y D. O ,0 O 0 a, _ G.
3 .2 .2
O u 73 75 ï 75 cd 2f O u 75 75 Oü 75 75 O
t-
u 75 ai
0. 0 0
» ,b o. o S -a u " .S , t o. ou <- D. ou O >,
». H
— -3
-0 «o
§S £u 0H s ËL-e-e
& "2 uÈ?s D.D.
-e-e I Iï sT:ss-11 s o.»-2-2ï ?u
. s
1:s
H So-a -Ss-Sg0 5 s, &
« 3 ° o .= tf S 5 a ni » 3 o 0 5 = c a u 3 J2 o u 3 « ä 3 .2 o . 3 =s
Ó U U Ü n n o « tu Q SUUHÜOS II, Û g U I I Ü BS PHO S U f c U f c P
ce o; o: « as
o
c
43
H
< s »
60
s2 -J o —
vi
on -g H '£öH &.g g-g e
u «2 § ..S C O <B " C .3 £ « c 5 n "i e

. 0 . 2 ûj « a « .g 3 S « 25 « u u « (~) oS j?
«C
M
c c c C
5 _ _ O " S H , _ c e 5
O. o O D. -O O D. o O * D. 0 0 7 :
3 -X3 '-C 3 ^2 'Z3 3 "Z3 '-3 5 3 C '-*- '+- u
O <D c3 «î O u cöcd O u cd aj .S O u -rv cfl <S O
t. o, 0 o u. o. u o % ^ a u u a ,b a _ .SP o o -a
ÇÊ 3 2 -M 2° s-? » 3 2 3£H «g 3 £££ » ° SS3 8
as c« -a t« i-, 1-, -n g 5fta 5 « t. » î s ft g . d»N s « ^ 5 go
« 5 ^ ° t ÄD .S O. Up, W!? >2 SS3 oS g - u S O > us 3SSo g s° - u > "US3^ S oS g C O»Oa- sD ü
S . «c
S 3 . 2 0 . 3 . 5 g « 3 2^ 0^ uË. 3S = S SO 3 s ° Ô ou g. cs c«u ' 2 S 0 ug
3 . 2 o 5 . 3 =« 3 S 3 ^ 0 U 3 CS " -S §3 3. 2° o0 u Cc3.3«=S
„S u i L , U f t . „Q _o _SCuJi LLL, .U
C0J aSi lLLL ,. Q
D i u[ n
S ) I -L U[ o
) aS! f i c
l, . QQ S SO üI Lf cI )U QSöS O>~IL,Q0H
OS OS OS
100 ^[gn'e. t/n/v. Wageningen Papers87-2 (1987)

üc
3 O
e o
o • . c
c ~* h .S
^2 « So.
•Eft- . g- 8 c 8 .23
.S « 8 .E S ^ S H rj
S3g £ S S ^ 2 ^ _, I cg
> ï Un 3 JO " -C «-• u C O—
..|£ | | I .-s| S jj-gl ,. J S | 1

^ K < .3 (5 c.. U Î S t ^ Ä Q u S t ^ - Q Q u 2 S ä l ü S ^ d i
c c c c c c c c
&
o- o o a ° .2 ' ° .2 ^ .2 .2
o o -c^rtsa O u a « S o u « s o u « es.
^ H <« o o 3 .£> Ë Ï H .o J= # E il H -= -° * il H -° -S
ft
> 5 3 3 « C O Q P , s C > 1) 3 ft D.
• ~ 3 0 ^ u E u 3 C 3 _o S 5 3 J2 13 .3 M fe 5, 3 . 2 O .3 M £• 3 . 2 O .3 «
§ u p- u ou O >» fc Q 3 •= « u n . u u . Q S M U H U H Û S Ü C U H O
tt ai et et et
M
O u
at y 9
o -
s e S -u
c e u g
" H S © T3
O g PH .
g a S u
o S c S
9* - J& «2 5
ai
o
00 >, JU >, O. c
rS • CS WH P3 ^ S" ^ rS
° B - S Ê - ^ U O ir0- O C WH
w QJ^ £ 3 Ä ft 3 « S 3 <" 3 3
° • ft — o g a 5 C o g 3 oo a
oâ1 "x £ SP ÄJ « 5 2 Pt à ê •$ O-1 o -5
U^ ( U G O H UÔ- U . <L> <, U o . u C/3 P Q- t/3
?« S3<N 53 E Ë Ü 2 ? < S ü ï „ <N »5£ .3 U 2 e- 2 Q £ j u 2 ?- $ U 2
§ I
e c g c c f e c c e c „ c c
a 0 0 - 2 5 o ,2 o. o o „ a o o « oo
3 \p 'S OH '3 S S 3 \3 ' 3 S 3 3 s u \S '-3
sg. nn a s s •§ 2 & ss | 2 g. ss §• ss
C b r J2 1H 3 ÖH g a :J U b ^ J2 3 3 E S. 3 -*S 3 3 o S. 3 3
3 2 2 " * o c s 2 = i » - 3 2 ^ ° E S 2 I g s ^ E S S 3 2 SS
S S 3 0 3 g 2S « c g -g 3 3 o ô S .s a 9 S 3 ° Ô 8 . 3 =« S S, i i
St« u E u a i E û SM E n S u E u o i C Q S ^ U E Ü O Ä M H Q o ^ u, û
? ?! * et ai
Agric. Univ. Wageningen Papers87-2 (1987) 101

•o < ^
8 | 8' g g 8
£ %. g. £ ^ •> c £ S- £
J 1 II |1 I I Si
•S« 3 ÜC3 . C i 8 V> . „ ea
S, 2 <*> r5 3 o » . »2 . ,ï ,ï Ja -s M
.5 -3 3 £ g 5 «5 .a -S S3 ,* £ g g .s -S rts ss
Tf- C/3 C/3 ( J ~ " C-- Q c - U — ( S Ï I P H J U - " S C-- (/] e-- t/> e - O «
_ c e c e c e c e
D . O O C X O O O H O O OH O O
3 TS 'S 3 -S '-S 3 ' 3 'S 3 C 'S 'S
o t u c a c a o o ca ca o o caca o o -c. cace
IL H £ X c ~ t_. .o . o J £ ^ 1 H »J /i %, " Ï 7 L H 2 O x> .o
C > o 3 O. O. C Eli > u 3 Q. O. tr > u 3 3 O. O. .S PL, fc>i>3C°0.0.
O 2 * ° S Ï § ï 2 s os s ^1s =*.2 s -s s ^ 3 es s o'a j= Ï «
c a o 0 i. a c « 3 o o 3 =e •—' 3 o o o ü ca ~ 2 3 3 ^ o ' C o c = a
IUÜHUP-O 3 äüU-UfcQ 5üö.üa!ii.u g ^ «SjUU-UO^'fcQ
cc t/i cc cc an c/5 cc
e
3
«
8 -e
s a
iü M b^» QDf.
O« r^ «a g£3
8 5b -g-S -S^ e
a .S ^| §"§ ge £
3 c ü ,j 5 . o u ä M *
o o EL, S JS ä £ 'S J2 3 «
^ c « o. ^ û c ^ o
0
Te .a 8 3$ =»§ 8ï s»
ps 1
3 o
«' j . -H M
. ce
pli 3 ^ . 3 ^
« -ff 2 -S
aâ .
o o -a 1-,
. . .
--il i^
J2 O. o S
. . 3 CL- t^
u w
LU
S o -S r- S c"- 1) S « 8 C J3 O •— vo c -S « 'G n " ""

» p 5w = » "3 5 > ,3 • - > £ «S o> .Scata.s Oooo
c e _ c e c e c e
B* 2 .2 *- ^ 2 .2 ^ 2-2 ^ 2 - 2
0 0 • r i . ' c a t a t . o o ' c a a «• o o "cata 0 0 -r,. "ce te
,to. ,_ .SP o o M L- o . o o .S t o . 0 0 t o . ^ » 0 o
» ilH 2 0-0-° e Tl H -° ^ —Z.H -0-° , I H cn°OJ3 .O
a S ^ S ' ï S g , S S^. ^ 3 3 « S ^ v . 3 3 3 s( S u u T < 3 < « 3 3
J 2 > 0 3 C O O - C L eo > o 3 P. O. > >. o 3 .r! O, O, C > " 3 c a e O O . o,
12So'a^-S le 3 * .2 S S i S * 5 2 S S a S ^ g ' ï S S B
. E 3 . 2 0 E o . 3 c a « 3 o o.S ta « 3 o o ce ~ ce — 3 . 2 o S 'C o 3 ca
•i Vtt.U O>- EL,Q SU6.ÜH D SO&.OKfc.Q gO LL,U <*O >» &, O
S/3 i/ 3 S7> y
ft
3
Vi Vi
O
0 S
s S S
u CH 0 0
Cu cu M
CU eu ~o>
, •
fc-
0
>* >-,
O 3 -C JS
'ufi(« _e E
E S e C
'? G •|.s
Ö 'ra
w
'w 'r0a.
j =
.•3
c«
0
'3
0.
" 'ra ^ra
a Vi tu Vi •c •a a
tu t «
"S Cu
en e S a £j en
? 3' Ç •0
tu 3"
s
'vi 3'
M
gçu
ra 3"
00
ra _-
j/5 'C tu J ^
<2 CD
'tM 3
3
-0
0> ra- 30 CU
cy} C
3
0 T3
tM > 3
ra• 0
'S
>
OJ
ra T3
W5
„ fc-
cu' 'ra ai E tu' > ^H
< 3 'S
V)
< .0 Xi "0 tu < s Xi ra
Cat.
c ra00 00 cu
0 0 .2
'C «0 O a CU
a. 0 0 ra Sra OO 0 0 C
ra 0 0 ]w 0
O C
Q U 3 s Q QJ U û tu U Q
„ VI •c Vi 'S
"S
0 0
v05 0
cX
0
s
3 E C «
SI
.ï
c e e
0 0
sCU 0. O
c0 ctu sV C
0 0 U

i- ra ra
0 0 t-< CU ra 0«
0 t-
0 O. ra ra
0 0 y t-
D. ra
u ra 0
VI U
c ra0 ra
.2
O Hft
u
o ;3 ;3 O _o ;3 is u? •5
>, 3 « 0 O i>
-> -M 33 x>
O H>-, 0 Hs->
E 8 ,0 -O 0
23 X3 O
Wî t/2
3 .ft
3 E u. ra 3 3
E ra 3 3 u.x ra u ^4 b « 2=> ^3
m 0. 3 0 . O. E
m ra tu 3 •S
'S e .23 > ra a ft a a S ft ft
CU «1 0> 3 V)
ft «
'w > . > Remi Kt
>
Flow
Culti
Colo
tu
c o ra C c 2 % ° V, B C e 2 * .2 B » " s W2
ra
'55
2 * 0 gs a
u G tu
s 3 S. 0 ts ra c
eu
>>
s C/3 Q e
CU
a s 3 J2 0 G 3 ra S E û 3 3. 0 S 3 ra
U CM O ö. Q U CM U <* CM Û s
P c/3 { « 'Vi 35 CÄ C» u i M ü a t t D
o
E
<
Vi
CM
p
^< I l < S « ft
'«S t E g 3 1
•- P fi- £ ^ •-3 S -3
I - c - „ -3 öig 2 S3 Ä
"S S | g g .a sêM •%
Ù ê1*
a
•Si» . JB -a c« c Ja S «; o oO c 3 . u <
Qôo .5 ra « 00 Û .S « « do .5 i a «S \7, o 5 -S ra « îS
" " 3 3 C C 3 3 3 0 3 3
-ti O O ft O O ft O O ft O O ft O O
T3 ' 3 '-C 3 ö \^ '3 " 3 'S * 3 3 '^ '-3 3 *"" ' ^
* I.§ l ï t 3 S § | 2 ï S§ SR SS 88. S3
U
î'33 t_T 0 S 3 3 I s 0 ^ 3 3 .O ^ „ 2 3 D
|O>> „ 3 3 0 , 0
g C f t f t 0 > j ) 3 c O f t f t ! / 3 C > ! U 3 f t - v S > U 3 S f t f t « i C ^ t U B r a -
•o g s a ? 5 s o'a s s B g ° 2 ^ ° a a " < 3 ^ E S S » ° 2 * ° g SS
'5 S i K ~3.2oEtD.sra £ S . 3 2 o ü " ä S 3 ° o f i s r a § 5 . 3 ° 0 < B ü = !
gcnCi-D 5 U C M U O ^ C M Û ^ G O U C M U C M Û ^ U C U (Ü Ö. û g ^ U B, Ü « 'tt. O
y4gr/c. f/«i'v. WageningenPapers87-2 (1987) 103

•5<<-: -Sr-: 2<
u-i c ^ ^ g ^ 5 c
O o
U 5 C
"2 ° «u 2 •?
g§ - * <* ï Ss
si s gaii si
eÖ~,s s ^ - S CÖA5
- 0 3 3 5
o - ; " a u * - ° •? 13 D
0 0
'3 .» o ' i T i M S a .S o J |
•a & g - 'g s g s g §.g S .s
111 H - 11? lil** 113S

•§^S)<o •aëSsSS" -ägScK^B^ ^ o s : >
o S 5 s « S -S J3 .S 1 « S o S - n ' ^ » 2 „ ° » •; I '^ S * n ^
s
£0> c e c e . 2 c e c
T a O O O. O O I - D . o o g . o
O 3 •— -^ 3 •— ' ^ CM 3 'S '-3 3 '^
"^ O u rt cd O (D rt cd W O (D cd Çd ~ -- -
J= -S
5 J 3 . 2 O B . 3 C « M 3 5 O O . 3 « î | 3 ° O o --JS M 3 . 2 O 1> . 3 «
Büfcüainn çUfcUK ü- Q ,S U fc U o! ft. Q ,Ç U fc ü »! tt. D
sea
M
c
•O J3 J3
4= * *
a ca ca
ja Ja Ja
•n t^- %-
C O O
on .. '3 •j ' M Ö PH
O. •3 VO -a o. "53 °~ 3 3
e/î c r- S v S
ta
v- O t/3
:cd oo
• 3~
Cfl (N
ÓD 3 Dû 3 S «i
i- u
U Ä
"o ro .ab «a3 u c£
b ei -
i»
S
(U
*2 "(*)Ö0)
sis'?
ro i-i cd
> -a > -o > TS c
e.
J^
fi
S
3
3 ^C
«1
CJ < ,
00 . 'S-
ai
c«
W
'S3
3
3
u <
oo a
3
3 ca » j OO
Lar
C O C en
Cat
o
188
aj oo u ca C »
«2û o- O - o- C
O o 00 s O
Q
0)
"o
D B C
CU (U
DC
n
s c u s ci
n
C C
O.
3 .2 .2 O.
3
o .o o.
3 .2 o •< O.
3
o O
\£
scd
X>
O a> rt Cd O U
« aï s to
CD cd '^
ni O u
'S , t-
t O. o .a O
O- o CJ s
Ü
. O. CJ _o Û O^. P. o rt
o 3CJ CL»
OH
H>> H>. 3 H>,

M
»o s
S Z. H t. 3 33 3 h.
«5 ^
3
C > 
Cd t.
3 D. VI > 0> 3 xn
CD 'vi "Vi
1s * ° ess 3
U
O O .3
a
'S 1 23 * ° B ts S
o o o S o
s
eu
B
2 * 5 E S S
3 O O » 3 1
C
Ç
es
CD
d?u n. u a: n. a ft. c U ft. Q ttî fc Q g
!» K U ft. U Pi ft. D c55 'Ä
104 ^4gric. t/n;'v. Wageningen Papers 87-2 (1987)

c
.3
33 »
o J3
ëm
60
•S? e c
> o
•c
00 -
o
•g 6
g*
o -o OH •-
UH - OH
u Ö <D
R £ SS = O eS
5 -o
s c W «5
o »
00 : <= B. C
M
•a es
oa u
<S •J2
a .Ss ° »» »5 S^ ï "5
ou ou j ^ 5 ~ ' <_; >; ;«
* £ « S o
vis -S > r £ =s £ V "3
O S « !» •S 'C es « 00 * oo •S >S Ur i —§ H .S 5 es
(Nt/5fflJyi-U- • c« t MOJ u -
a
o. 9 o.
3 .2 -2 3 3
O 4J
o o CS es
J-. r v »| | Ü O i, t.
'S io £ 2 o3 3
=S <a 3 -G.
-g SE o 3 3
o, s e>* es O H
an 3 „ es - c3 s--
e~ 3 3 S 3 M ftft
LS p o O \3
eu 2
3 »g O
o- « -
o o. o. S O g - «J
< 73 eju
ri
-6-0' eS J» *J •â i I-, ^
' a « «D
U ,2 O u 3 es
c«uE U C« O >- P- Q § U 3Ü O
. "U O ; O > -1>P. 3H eS " S O U ,u" EQ "S M U B H o a i un Q
u bo
•£<: es CJ
.2 s
^ E
S | S 00
<^ o -£? C3 o e <c
es o
ja "
o •H S
" c
e> 2
o U ^
O es
," o
't- O
eS D
a
3
0
25.1
3 S
— -O
»O
3
H ^ 'S O
° ( , .S a o 3
§"^33
<-> H * eu C/5 <
B5 1) >—.
J3
3
a
t - X5 --H
eS \ ~ ^
t - *r" n
00
c
B 3 S
'SÄ.S
1% C«
-• 2
.
00
i-i g C B. •o
u •s s <
J3 SP c
<C o
o£ " -
u u c ?! g o
. . 2 >> o JS
S S
oca u a\ .2 «j >r>
„aï S i B)
ss
Bs •3 > r»H .. es -=S rrs
-= O 3 *H
la« o 'S ^
> c ö£2 - t/3 > U . —i r-Su2 -* S3 >- S3 8
c c
a3 a S o S
o
3 .2 .2
E o u rt es o tu "es es
.S -S J O S .a -H
-°
3 -S
es es t- t-
a3 o.3 a a h b 'S S & >. g g
[fi > » 3
u 'S 5 O H - CJ
? > g 3 M
»5 * og v3 +- 5 5 |
o. o.
< S °" a
n V] 4-1 1H M S
"Bk 33 .2
^2 O CJ ^b 3 ^ o S t- es 3b 3 _2 o « > , . 3 es
t- es .S O BH O ÖS U I H U B ! BH Q
O BH O Pi fe Q Cu O S M [I. D
VI ai

•S-o ^
ja
O w
•O H «
cd
J« -o
CS 3
- t u
s • ; ij S 3 _ .2 a cd O j 5 o 5 «
? i | 2 .£ £f £ £ 3 o^g§£
>
-o g 8 o 2 3 a ^ 6 S S a S -o 'S « 3 g
llllii | &s
Jtini l HUIi C/3 - w ^ - ^ C - j - j C / ï
rt
3 ,# -S 2 « O M OO r"? OO tj .5 * S S 1 « CO , ^ OO £ 3 * ••? i5 rS* °° n °°
•*• .2 Ce
2 ., .S 2 2 SS „ .S -52 S I S », .S ' -
££ £ » 3'gSS S? ^ » 2 2 2 JS ï ° £ -2ë22
S 2 ^ o B « S S ï S2 | « E » Sî! S S § 2 g2 E .»f S s £
> 32 O 5 C v ^ c s >• 3 2 O fljCcubccJ > > 3 ,2 O uCoj.ücö
t» CC Vi W
> c
TJ
<D 3
o C cd o
•> ^ . & -s
ja g 8 -a
- es 2c >>
•o .^ « o
u
D « « ï -ï ^ r - C/J
"•» « 2 O S C -3T3t.
2 > 'S _ " " & £ S— 2 3
•s 2 o g S g ^ B ~ " o s o
c
*u cd
M
jj
u>5
•S 3 2 ^ »e ü » s 3 5 3 ' oo
</1
S 3 "S
•=f U o
Pi
c c
a
3
o o
\2
00 a a 3
o cd
i~ u «
o. o o
0t . H>» JO
3
s l a a f t Sc^Öëao, ijsjs feS'oäa ' « - ^ ^ I l ^
. .. ^ Ä « S g 2 2 * ° S S S 2 ^ 2 e a l î S S S 2 c î
fi 3 o O . ü « " > , 3 2 o . ä ! 5 S = 2 o u 'E o .S3 « >
• 3 O P - U Ü . Q gdoCJBHUU-Q §CJti.CJ D ^ O > - B H Q g
c« ce ce ce
106 ^gn'e. t/n;'v. WageningenPapers87-2 (1987)

S S
.3 5£<
*_ "u =a a.
•s« =i 6 >>sz
u
'S u <-> « O n
•n ca « es » g S o
»ft. ftft- X T3 ^ I
os
«h S " ta
-C _ö
J2 « ft.
T? <->
- 1-
w C ~
S-
Cr.
« . <L> ^
,_• S-•s £~ a H
>
oa «.«!X-G .
0
o 5° 3^ ,/ £ ^ -
. . S3 132 o3
S £ ja 3 ?" . -2 "g = £ >;
ü ü -s -c <» 3 2 ^ -s 0* » *ï 3 s »i « u >> u Z u oS
~ > J5 cd « oo N i d p a .«. » g .S 3 i h s co , ^ oo « ci .3 ca o nj «j ca~
S a
e c ^ e c u e c cj c e
OH O O . C O . o o ** o. o O _ Q. o o
3 C '.5 '-£ î> 3 c w '•£ '-C 3 C 'w w S 3 C '-3 '-3
Sg, »gg s2& 'ag g £ 2 â « g § a gg, -a s s
° £ »o'ëgg «O£. „ S ' g i s s „ O£ M o'g33 J i ° ( - ? » So33
gga-Si-sta ï I g a - s i - s f a ; g g s S - g l - s l a ï I I aa-S f o t â
3 .2 o 5 E u . - ca £ 3 ^ o S C u . 3 c a 6 3 5 o « » C ô . i : s •; >, 3 .2 o 53 ïcï.b u
ütt.U«!0><ft,Q § Uft.UaiO >• u- Q gOft.U BCai O >- u- Q gc«u ft. U ai O >-ft.O
c ca "O s ca
u SÏ S S
C r/î —- en t/1
ca w g J4 vi
C CL) ^ "3 —-
g §-S S
13 ~ r P. « u
'S S S3•= g o.
Ó; eg-S s s
u-i . O 0.3 g. «3
1 "§«i§ > S e
SS Q .? H ,S8 « g§ 1
h b ^3 £3 s ;; 5 c ^ *-• " - a * -
£< £ g 55 g | g o fc « 5 fc
* CL, ^ «j o ^ a S Ë S u L - ' u D . 2 L . " U
il«?
* .2 fi Üj £
F/5 IS N
-öÄ
^ S Ip M
' c ëI^S i ca .hls i • u ca a „ ca
« 8 3 . Ë,« g 3fe1^äS« S ' <? u ^ S j ä
3 |
c c ^ û cc cc
3 -S -S = .a 3 ° ° ra & -2.2
?^ ja 1 1 -s I Ï H ja^'§ll 3ïÊ 52'oss

3 ° o S ,= s > I 3 oô » S'C S| a » 3 0 3 gÏ 5 2 3
Uft.ua: ft. D g guft.u ^ o ^ n û gut.u aio >£ a
</) t/3 W
^gri'c. t/ni'v. WageningenPapers87-2 (1987) 107

e
3
S
.ë'S-s
J3 'C
"c« c«
C
aj
^
J3
>VI
• "'
; "o-ou
-O
8
3
C e
3 '1'öi>e
3
•o C
aj
3
a *J
JJ
3, 'ja
c/3 "ÖJ
aj b. •s -s
o <
M 00 m ^ca "aaä> «
s
C3 -3
CJ S SO
o O ~x c
OJ
QJ
c
CJ
-o •3 ;
U
Vi
C
O
c
S
3 . aSj V) OJ e
QJ E c S.: 5
a (H
3 U 3 3 C 3
3, g I oS ™


CJ
M J2 g to . m
Y
to o > B
CJ in "et) CJ 3 - u cj m
in •o S m g -a *J S
rt 5 °c ca 00
Su o oo 'S -2 o £ •-
3 •*« S
° S'5 S
Onu aj
pU oo O u B OO
CJ
. 3 CJ
o So Q ai U
S c es
'cd
s c C CJ
C C •g C C >
fi c o. o _o
~cj
3 _o O t/5 P. O O Vi > a o
'o '3 O3 CJ 3
o
o
cj
o
u
o<
S 'S
o u
3
O o
Vi
'S
_o o
"cd fa.O
OJ
O b-
aj
a
S
O
sÇJ 'M
*o
?=
"5
o a>
*-< O-
«
o
U O >> Q O >> U.
3 3 VI B 3 3 VI
3 3 V) £ 20 S «
3 3 3 3 3 3
S«
c c s
eu
B cd
H 3 O. a.
C
c ex CJ >H k-
3
o. C
QJ
« oC" sT2> 5<D^ o3Clesr-„ O-
*- en_
CJ
•o • Ö S O aj • o •a o aj
•a aj O
S QJ •o J-I
C S 3 .2*o b « c c c S B 3 5 ö i « c S 5 3 5 o S .Ü ca
3. "S. U PHU UN û 3, 3
Q "3. U fr. Ufr, Û 3,
c/3 vi t » c«. w &n
.S ^
• 'S E
>4 S » i M
3 W C ^ ~
o- u « u "a-2"*
S e g ë jj.§«
•a ra Qj . 3 j
t- /Ä
o a -o „- o«
!s O 5 X .SP e
g f e # "g dB-s | -g
50 £ 3 a l s « (
d ,
a ^
? . ? a * t^ s 1- cj c ja - " r-~ 3. O Q J U 3 O \ S — O o S S S ^
1 00 W fr, Ù0 — U — -^ c-!*>c^(/JC-"U— Ï J — Ö [In < , ' a rt y - H K l i - l U - ,
c
s
Ä
aj
3
O CJ •S 's '5 C '3 3
Culti varGr
60 O CJ
ouven ir d'uni
Flow er Typ
2h ' O
J JD
^ ^X3
w a" P 2 o 2 * -v ET.F S * -„ets- JS-S-S
3<- 33
Colo ur
O O Q. C •3 S > o j 3 c o a o , 2 e > u 3i °- « o » 3 U S ft»

g 52 M u S s S « 2 2 ^ » S "SSHS^cg-sS
Origi
Date
Year
c
First
QJ > ü s O o - c S ä « •Se,3°o.£ S g g , 3 ° o 8 " - S «

CJ
g |,Ufr.UO>fr.û E c / T U f r - U t , Q -g,« ufr.Ut« aifr,Q
Vi
108 ^gr/c. [/«/v. WageningenPapers87-2 (1987)

E S
_3 3
« S
J3 ja J3
3 -_ * 3
•q B .-3 T3
K S 'S È u
O -t-*
c
(D
._• !D . fT O <L) "
b ÖD . U ^ P . Û Û . -' ^
. IlS g g £ !'f S gt
•a 8=§ °° 2 s ë §°° 1"
ä r-J2 .2SSo«^ fe u
I .2
I s»*S«o n Ié .S »
C 3 .G > _: 3 . • 2 . Ë O > _ J S a
c ( D
.ß . CQ o-^^wr« ^ ?- - « ! t -T f 33Û3
S ^ ^ ï ï ö 3 . * S ' ü •* J Ï JJ 3^??S5'ä3 ü — ^J ^
3
~- U CLH U —« r -H P ÛH CQ c - C-C--C--U— ï"r-PiXi_lfl3 c-H u T O —•
I-I
S °
a
« o- 2 .2 °" 2 -2 "e 2 -2 3 ° .2
H H H
ï £ s 3 •§ -s 33 j l s .a-s 3 •§&•§•§
• a o - s S o - " » -a ' 3 S o •" « -a o \ B is o e -5: , 3 . 2 o .3 s S 3 i o .S « 5 >, 3 .2 o «3 . 3 « g 5 .s a
U ü f c U f c O •H.ertO&HUfc, O •B.UU.UfcQ -B, t « O fc O Oi fc 0 • a « 6 , Q
sc co co co M
e e
3 . 3
co a po
t. 2 t,
•o ^ -o
2 =§
« g
-au Ü S a b . ä e ö> .
ga _- o ^ bo « .5 '-3 s=
H
r
e * s" -_ e S 8 a • * ° ^ c a = " . o m
Jä'S«§1j I Jä'S"S| Ja-S S « ^ S U U § JäHÜ^S

s,S-ï33^ ÏSSÎ53S silÎSi» 8 s a=:lâ«SJ5 Ss
9- o o a o . o o a a O O 2 Ö
f 2| SS Br|& S§ SS8. S S 'S S1 | S& S S
î Ç ^ » 3 3 « £ £ « S S S=Ö>- „ g g O ei « S O £ . g g
llöslää i^öslaä §2Ö3^1 g. a g g i & 2 f e 3 _ ä a
§ 3 ° ô S . ë S S 3 ° Ô O . b « g 3 ° ô | g .g S g g g C j l o » S S
71 Vi VI uï à? 03
/4gric. C/n/'v.WageningenPapers87-2 (1987) 109

3
S
S
XI •
3
C
i/i
£ 5"
a
âg
(U eu
> i - 0 tu
£ 'S,
•§) M>
S
M
J= 3 JÜ
fc
S . -a •?..•§ . . • " J3
ta o
C
6 0 c E
o ^ „ S « h S^& 2-3 -gin

'M Ü 2 .c _; ^ -S "» M'S 5 h Z •jr> "3) 3 -en oD'O-o « i «
'G .«
ft
c c > ,"
O O ^ i ^ D .
ce OO ft
c=
OO
_ ft
e gO O
3 'S \E 'C t« 3 \3 'Z3 3 3 'J3 w 3 \3 'w
sa ss o s s a S3 sa '§>gs ga s§
-_° £ » 3 3 fï hO>. „ 2 3 « ° £? 2 S33 ££ » 33
l l s s l S. ä | I l s s i ä ä S t a s i ' s as. ' B l g s l - S ä S .
o 3 ° ô S £ a o ° - 3 ° o S . s es = 3 ° O - C U . ! = M * 3 5 o » » .S «
lüüUü! tuQ 2 »Ufcüa! tu Q gunUOûÛ SUü-UKoiPuQ
3 ^
„• . S ta
o "o -o o
OH
O
l-H
C/2 SX)
e J-i
M
ai
(U
00
3
s O o o S c •3 VC
u ta c oo
vi co CO "c D,
•3 G C
O tu fi ri
-o
3 C
•3 T3
o "S O
c c: 3
•a S . ai
cj c
OH
C J= > •6
o. O. "H- ai C ta
S W
S" (75
00
—û
• § 1 'S
O Si 'S-
Oß JU r 1•" oo
• VC 3
ai
fi
</3
'cj ^"
. S "ta •9 ~
u U U c
u
fi
d)
CO
(U
en o 'c/5 .S3S ta
U OH
ca oo
<N 1/3 0 . c« —
S
C" c^- o -
O
03 c —
u oo
O
C
1
ca
(U c C C C c c C 3
O. a, O .o o _o
s S 3 .2.2
3
3 ^c .2 -2 3
4J 3 O 
a> <
>-ta• H 2ta § 33 3
S Ü H 3 3
e "a
e S3
H
3 3
'C ca
3 3
a 5 a 2
M-;
B s Ja ta> S-u
uO
S '3 > o
UH
3 D. >
1-1
3 O. a > U 3 co o. 3 o. o.
'S _2 3 -J ' S S O Su g o% H eu a. 'S S (U
C
.2 .S o
3 ° O c 3 S, O fa ed 3 .2 o C u .s ta
•fed o
tu
3° O .fa ta
'C tu Û s U UH O ÜH û s U tu U O >• tuQ O tu û
3 U tu U
(Z)
55 05 H H H
a
_3
«
ci JD
C.
•g
W5
'S
T3 ~3 •a
S u
.S -ft u cd o
3S J3 PL,
ai i-
Sahut, Montpellier.
CJ
Cat. Verleeuwen, B
Rey, Carpentras, F
Hardy. About 10pi
Large flower. Buds

Pink, spottedwith'
O
s f~- o G
Cd
c l-H
_w CN cd
Carmine red.
k,
PH
"o rn U,
>-d >, 3
Double.
Apricot.
CD
Single.
oi ai on
o 00 • ô
S t--
oo CN g
Ü 00 u 22
2
es. c^- c^. e^> m
c 't«
c
iblication
0 C C
O P. D.
rGroup
b 3 C
o 3 c .2 -2
O (U 'S S u O (U %S
Type
cd
O C D. u '5b
=iVft •- •-
3 g. Ü >> t-i 'n 0 0 a t- u .o
o 3 3 « 2 o
a u. n g 3 U cd k- 3 3 s cd H 33 33
> 
t-<
3 > u 3 ÛH
> u 3 O. O. o p. 3 O
S ? ° SS •a ä § .2 e :SP «• (A Q 2 * 5 Ë 3 S a- <->ai l
*c«
S 3 S O .3 3 ,2 O != «
3 J^ O u k. u
t. 'S M 3 ,2 o - E Q O Bc U «S O >< CL, Q
C O
O <D
•o I -
(U
J5
(Ä
(U
JD
£> t/5
3
O, 3
(^ S
u
3 S s
J3
^T3 c-
"crt
o cd
a
3
su ni ed
 1 en
3 •X
r/2 u
in
Os cd o
O 3 : | » 3 S CL, cd
r--
(N
"î3 "o cd'
5 =3 ^ 60
>> CL, _CJ
'C — • ^
o
u "o f i 'p,
S .0
•J
•3 C
> dcd < o
cd s (— ^ T3 ed a, t-,
-o
ereiE
dgeo
• P\ cd cd
osse.
t/5 c 3 ^ > O 1—»
H
"ob.* 00
354
CS 00 3 Cl
0 0 ^ CJ Z oo
.S -S u c
J3
Cd 'S r-
.S -S 3, o o 3 3 cd (U
U cd
(N !» S OH eu m s —
C/5 c
CN. u oo
CN 't» E • J U =
9° a E QJ
cd
Ü
Os
C C c C 3 3
.: ex
3 .2 .2 3 o O o, p. O .O
0
( O eu
rt "5 S '^ 3
O v « rt jr 3
cd cd
i A D- o .o k. '2P o O o
! C5 >> Ü O O r>> en 00
H>>
stpub
marks
ultivar i
o XI
te pub
lower T
J3 *
S *
3 3 3 3 Ë 3 X
3
olour
( H
C o O. C w £ >i cd P, O.
cd k. h
c (U 3 O, P .
3
e« o \s>£ o s
c3
C a 2 5 _o ]S) cd u
O CJ 3 cd
S 'C CJ
Vi
i-
CU ï£ _u
'•is I I ra »SS
•^ S _- ° -e •~ ^
tJW ai aiS » » c ° Ä » 2 ^
812 .§•£*§ -slllg s Îlgs

^. c-..c^. M e — — c/3 !> U — -(oSfflO.« S (NC/>:>U —
g g a g§ a g § "Ig. JJ
S o u rt ed Ou eö ça O u ça « !Z 2 X ?1o
« 3 0 - 3 " M s 3 .2 "o § « Ä 3 .2 ^3 .3 « •- •- a -2 o.3 a

S ü E ü i ï Q rfOB-UP-Q fUH-UOn Q £ J Ü H U H . Q
> P ? ? s?
=cd cd
00 C•
—
M
i
g :a o
C
C 58
w- d
W •»
-- - a2su, «-"
à 'S. sw s t/i __a
_ 3 ^ 2
•ar, « i> ja o< & a « üsi s « _
4( « -g
" H 8 ' S 3 -0 ^ S ^ P s P g>3|

o. o .0 o. .2 .2 §• .2 -2 §• .2
o« ' i 1 o» ; |1 S Su 2.1 28."
»2 15 « s s 13 j » B g ' a a s s 'S2SOB ts s S« s ° s .a
•S 3 o Ô .3 « 'Eb •&3 © ô « S -C S .gS • a 3 5 o . .ë s « 3 o"o.S«
t S ü f c ü f c û .3 i ü t i , ü Ä ( i i O > - C k O i u E u « £0 Su EU£O
112 Agric. Univ. Wageningen Papers87-2 (1987)

<s <s
Ze Ze
es s«
^ <0
D
on >>
o cd .
_g c
O T2
& 3< Soo Ü
<-> 33 <
o>
fr*
c<3
<S 5 ä «
ö.S .2 „, «Zag
i »1 ( j
. -S S â c - g " JS e e 3 g~ > ° •-
u S g - a o g £ S 3 u '5. 3 -o o g o 2 •-
o. oo o. oo
O o -rï. ca ca . , O y 35 3 e o o. a „ > Ö3 3 S c o o, o. u
2 * ° 2 e » S5 2 'S H ^ .2x g '& Sa a 'S
a i os» CS .s s 2 3 .2 o « u C u i c a S
U t i - U X o ; O >- U- Q H U U - U X B ! O >" ft. 0, H
3 0 <• g
•3 M • ca
u fi ZC
•go g là
•O cl g -S
Il ?Ô
O O . S R O
<! ^ - 6 5 .
e
•8£ , fi SS ö . g
«s s « ^ D «zo.g.2
" c § 2 s | .ÏÏT)>5„- 2
2 I a-o »-S • d â 8 2 « | § «
I "O
*9 S > 3 •= .«I * ä .5 ,5 •£ g >, C ^ ri » 3
c>. o. P . 2 ï ? K • * C«0- Ü —• H » I Q 2 H J l J j 2 r . Ü H <
Ö C C Ö q q
D H O O P H O O Û , OO
3 \3 •;: 3 \3 'S 3
.- o u 3 ca o T H •» £ O X>X> ,5
^ > « 3 o- o, „ > u 3 o. o. > o 33 3 c o o. Q. u
5 2 s os s s» s05« 2 s0 3 c 'a s -s s •a
5 3 » o . î | .2 3 ° ô .£ « s oô a § 'C S s 3 g
r ü f c ü f c Û % U CL. Uft. Q U H Ü Ï A 0>-fcÛH
? >"

Oleanders Nerium L and The Oleander Cultivars Ser-Wageningen University and Research 290321 PDF

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Oleanders Nerium L and The Oleander Cultivars Ser-Wageningen University and Research 290321 PDF

Uploaded by

Copyright:

Available Formats

SERIES OF REVISIONS OF APOCYNACEAE

00000307 7829 \^n 2.02,^3

© Agricultural University Wageningen, TheNetherlands, 1988.

Printed intheNetherlands by Drukkery Veenman B.V., Wageningen

PART ONE:NERIUM OLEANDER 5

PART TWO: PHYTOCHEMISTRY OFNERIUM L. (N. G. BISSET) 27

PART THREE: THE OLEANDER CULTIVARS 39

Agric. Univ. WageningenPapers87-2 (1987)

Agric. Univ.WageningenPapers87-2 (1987)

The genus was established by TOURNEFORT in 1700 as 'Nerion' and was

The binomial Nerium oleanderwas given by LINNAEUS in his Species Plantar-

Inaddition tothespeciesmentioned above,severalotherNerium specieshave

Agric. Univ. WageningenPapers87-2 (1987)

LEEUWENBERG'S description of Nerium oleander L.,including synonyms and

PHOT. 1. The lectotype specimen of Nerium PHOT.2.Labelofthelectotypespecimen of Ner-

Agric. Univ. WageningenPapers87-2 (1987)

Agric. Univ. WageningenPapers87-2 (1987)

The distribution map of Nerium oleander(Map 1)is prepared by the present

Distribution: Libya, Niger, Tunisia, Algeria, Morocco, Portugal, Spain,

Nerium oleanderis generally found in a wet habitat on sites well exposed to

10 Agric. Univ. WageningenPapers87-2 (1987)

Agric. Univ. WageningenPapers87-2 (1987) 11

Nerium oleander,awide-ranging species,showsavariability inflowering time

1. Southern France and northern Italy: June-Sept.

English: Oleander, Rose Bay (USA; another plant called Rose

12 Agric. Univ. WageningenPapers87-2 (1987)

Aselection ofthespecimensseenby LEEUWENBERGand/or PAGEN

LIBYA:Derna (fl.,fr.) Gregory 23.VII.1908 (BM,MO);ibid.(fl. May)Pampanini&Pichi-Sermolli

14 Agric. Univ. WageningenPapers87-2 (1987)

Agric. Univ. WageningenPapers87-2 (1987') 15

16 Agric. Univ. WageningenPapers87-2 (1987)

Agric. Univ. WageningenPapers87-2 (1987) 17

Nerium grandiflorum Desf., Tab.ed.2:92. 1815 = N. oleanderL.

Not thosepublished in synonymy only.

Nerium antidysentericum Lepech., Reise 1: 270. 1800 = Apocynum venetum

POLLINATION OF THE OLEANDER

INTRODUCTION:Theflower ofNerium oleanderfeatures apistilwithamorpho-

MATERIALS AND METHODS: Living plants of fourteen single-flowered oleander

OBSERVATIONS AND CONCLUSIONS: The technique described worked very well

The Nerium oleanderflower has an intricate construction (Fig. 2), connected

Agric. Univ. WageningenPapers87-2 (1987) 23

Allpartsoftheoleander contain toxicactiveprinciples.Theplantisextremely

24 Agric. Univ. WageningenPapers87-2 (1987)

Nerium oleanderisknown as a medicinal plant since ancient times. DIOSCOR-

Agric. Univ. WageningenPapers87-2 (1987) 25

Pharmacognosy Research Laboratories, Department of Pharmacy,

Agric. Univ. WageningenPapers87-2 (1987) 27

Extensive research hasshown that theplant contains a complex mixture of

Digitoxigenin,R = H,R t = 5ß-H

28 Agric. Univ. WageningenPapers87-2 (1987)

PHOT. 7-10. Oleander cultivars. 7.'Jannoch'; 8.'Soleil Levant'; 9.'Minouche'; 10.'Amboi'na'.

32 Agric. Univ. WageningenPapers87-2 (1987)

Agric. Univ. WageningenPapers87-2 (1987) 33

Neriumoside A-l Ri = D-Digitalose-Gentiobiose, R 2 = OH

34 Agric. Univ. WageningenPapers87-2 (1987)

12a Oleanolic acid

15 Dambonitol 16a 2-Hydroxy-acetophenone, R = H

Theoutstanding phytochemicalfeature oftheplant is, ofcourse,the presence

36 Agric. Univ. WageningenPapers87-2 (1987)

F.Abeand T.Yamauchi(1976),Phytochemistry 15,1745-1748.

Agric. Univ. WageningenPapers87-2 (1987) 37

38 Agric. Univ. WageningenPapers87-2 (1987')

TENTATIVE CHECKLIST OF O L E A N D E R CULTIVARS

• ^:s -si fif | I l i s i tj "£3| -Og