CLASSIFICATION/Conventional Taxonomy (Wild Roses) 111

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::~aritsky G (1997) Background and principles of invitro conservation of plant genetic resources. In:Razdan MK and Cocking EC (eds.) Conservationof Plant Genetic Resources In Vitro, vol 1,

CLASSIFICATION

Contents

Conventional Taxonomy (Wild Roses)Horticultural Classification SchemesCultivar Identification by Image AnalysisChemotaxonomy and Molecular Taxonomy

Conventional Taxonomy(Wild Roses)V Wissemann, Friedrich-Schiller-Universitat Jena,Jena, Germany

1) 2003, Elsevier Ltd. All Rights Reserved.

Introduction

Taxonomy of wild roses is notoriously difficult, for a\'ariety of reasons. On the one hand the enormous

henotypic variability is based on genetic complexi­ity, while on the other hand the close relatedness ofhe genomes and fea tures of the life cycle have begun to

be clarified in recent years. However, traditional tax­onomy based on morphological and anatomical datais in itself subject either to biological phenomena suchas reproductive biology and ecological variability, orro theoretical problems like What is a species? andHow can biological diversity be translated into taxo­nomically and nomenclaturally valid studies? Themost common system in use is the classification by_-\lfred Rehder (see vol. 3 of this Encyclopedia): nev­ertheless, there is an urgent need for an update ofthat system, for both nomenclatural and biologicalreasons. In this article, methods of rose classificationare discussed and the Rehder classification is updated.

Classification of the Genus Rosa

Roses have influenced human cultural history inyarious ways and the enormous interest in roses led

General Aspects, pp. 26-49. Enfield, NH: SciencePublishers.

Towill LE (2002) Cryopreservation of plant germplasm:introduction and some observations. In: Towill LE andBajaj YPS (eds.) Biotechnology in Agriculture andForestry, vol 50: Cryopreservation of Plant GermplasmII, pp. 3-21. Berlin: Springer Verlag.

to several attempts to classify the genus. During theRenaissance, in Baroque times and up to the earlyeighteenth century, rose systems were just dividedinto wild and gentle species, followed by subdivi­sions based on petal colour. The recognition ofhybridization in roses was first remarked by Linnaeus,who wrote in the Species Plantarum of 1753 that hehad the impression that Rosa species are difficult todistinguish, more difficult to determine, and thatnature just for fun mixes species to form new ones.Also Herrmann claimed in his thesis from Strasbourgin 1762 that species are very difficult to determine onthe one hand because of the lack of characters, and onthe other hand because horticulture had merged thespecies so that recognition of 'pure' species was nolonger possible.

The first attempt to create a real system of Rosa wasmade by Linnaeus. He regarded the shape of the hip tobe useful for classification, and this character was ac­cepted nearly exclusively until the beginning of thenineteenth century. Willdenow remarked in 1811that form and presence of prickles as well as hairsand glands can serve as species-specific characters.This remark led to several new systems which werebased more on personal preference rather than on ob­servations in nature. The application of various char­acters to treat the genus systematically was the firstsign of the enormous increase in names up to the be­ginning of the twentieth century (Table 1). The numberof different studies on the genus Rosa at this time isnot a sign of real insight; it is an expression of uncer­tainty facing the bewildering diversity of the genus.

112 CLASSIFICATION/Conventional Taxonomy (Wild Roses)

Table 1 Number of species recognized from the sixteenth tothe nineteenth century

Gessner (1561): 10 speciesDodonaeus (1569, 1616): 10 speciesLobelius (1581): 11 speciesGerard (1597): 16 speciesSpigelius (1633): 16 speciesElsholz (1663) 15 speciesWelsch (1697): 18 speciesSalmon (1710): 12 species and 32 cultivarsLinnaeus (1735): 12 speciesLinnaeus (1772): 10 speciesForsy1h (1794): 28 speciesLaicharding (1794): 31 speciesWilldenow (1811): 34 speciesSmith (1819): 57 speciesTrattinnick (1823): 24 series with > 200 speciesSeringe (1825): 146 speciesLindley (1830): 101 species, about 300 synonymsReichenbach (1832): 77 species, about 200 synonymsDoll (1855): 114 species, about 200 synonymsDeseglise (1877): 15 sections, section Caninae alone

with 329 species

However, not all members of the genus Rosa faced thesame problem. Mainly the dogroses, section Caninae,were subject to intensive species splitting.This culmi­nated in 1886 in the article on 'buissonnomanie' byCrepin, who vehemently accused the rhodologists ofnaming each rosebush with a new name.

The change in understanding of the biological di­versity of European wild roses started in 1873 with thestudy of the Swiss rhodologist Christ, who reduced theinflation of names to about 30 species by applying hissynthetic classification system. This synthetic methodwas based on classification considering combinationsof preferable correlated characters. In contrast to theanalytical methods applied before Christ, his approachcircumscribed taxa which he recognized as being nat­ural groups, rather than artificial entities. However,molecular data clearly indicate (see Classification:Chemotaxonomy and Molecular Taxonomy) thatthese groups have to be revised again. Going backin the history of rhodology it is obvious that the tax­onomic treatment of the genus Rosa is influenced by aEuropean view of the genus. Systems available todayshow that, based on the increase in knowledge of bi­ological diversity, there is a strong need for systematicand taxonomic studies in roses worldwide. Whereasthe knowledge of middle European roses is quite ad­vanced, next to nothing is known about easternEuropean species, and Asian species and even northAmerican species are quite unknown from a system­atic and ecological point of view. Thus morphologicalclassification of the genus results in at least someunnatural groups.

In this article the key for identification of Rehder'smain categories is replaced by a key oriented on thewhole species diversity. Nevertheless, morphologicalclassification is critical in sections like Cinnamomeaeversus Pimpinellifoliae, where unique characters forthe sections are lacking, and where molecular datafrom both ribosomal and chloroplast DNA indicatethat morphological partition is artificial. Another ex­ample is the circumscription of section Carolinae.These roses are generally described as not stoutly built,but they do not show any significant specific charactersfor a sectional classification. Looking at these roses intheir natural habitat, one can instantly see that theirlife in the shadow under trees in the woods forces sucha fragrant growth with single flowers. However, weknow from cultivation experiments in other sectionsthat these characters are strongly dependent on eco­logical features like sun intensity, temperature andhumidity, but there is no knowledge on the plasticityof these phenotypic characters in section Carolinae, sosection and species delimitation may not be natural.

Changes in the Rehder System

From a nomenclature point of view, the subgenusEurosa must be replaced by the subgenus Rosa. Basedon generic type, the subgenus Rosa contains sec­tion Rosa. Section Rosa replaces section Gallicanae,because R. centifolia L. is the generic type of Rosa.Following the rules of the International Code ofBotanical Nomenclature, the first lectotypificationby Britton & Brown in 1913 on R. centifolia as thegeneric type is still a valid choice, but is disputed. Theproposal to replace this choice by R. cinnamomeaewas rejected by the general committee for nomencla­ture at the Tokyo Botanical Congress in 1995. Theproposal was again not accepted in the Saint LouisCode of Botanical Nomenclature in 2000. A revisedproposal will be discussed at the Vienna Congress2005 (Table 2).

Addenda and Corrections to theSystem of Rehder 1940

Subgenus Hulthemia (Dumort.) Focke 1888

Monotypic subgenus: Rosa persica Michx. ex Juss.1789 (=R. simplicifolia Salisb., R. berberifolia Pall.,Hulthemia berberifolia (Pall.) Dumort., Hulthemiapersica Bornmiiller, Lowea berberifolia Lindl.)(2n = 2x = 14).

Subgenus Rosa

Sect. 1. Pimpinellifoliae (DC.) Ser. 1825 About15 species from Asia and Europe. Taxonomic sub­divisions uncertain. Molecular data support the

CLASSIFICATION/Conventional Taxonomy (Wild Roses) 113

-= Ie 2 Key to the sections of Rosa

::

=

Leaves entire (1 leaflet), without stipulesLeaves imparipinnate ( > 1 leaflet), with or without stipulesReceptacle smooth or ± gland-tipped acicles or bristlesReceptacle with prickles like a chestnut, sepals pinnate and erectStipules adnate, with divergent, rounded or broadened auricles, leaflet

usually 3 or 5, rarely 7Stipules adnate, with subulate auricles, leaflets usually > 7 (-15)Stipules persistent and half-adnate with auriclesStipules free and ± deciduousReceptacle and younger branchlets tomentose or pubescentNot this combinationSepals erect and persistent after anthesis, receptacle bristlySepals reflexed and deciduous, receptacle smoothSepals conspicuously pinnate (the two sepals outside pinnate on both sides,

the next one only on one side and all of the two inner sepalsSepals entire or only the two outer ones with slight appendicesPrickles glabrous, straight and mixed with gland-tipped bristlesPrickles straight or curved, not mixed with bristlesSepals much longer than petals, younger stems with a purplish bloom,

leaflets with purple pigmentation, prickles straight or curvedNot this combinationSepals short, erect, leaflets glandular and ± hairy, low and compact shrubNot this combinationStyles agglutinated after anthesis like a columnStyles freeSepals after anthesis erect and persistentSepals after anthesis reflexed or spreading, deciduousFlowers mostly solitary, solitary flowers without bractsFlowers solitary or corymbose; if solitary, then with bractsStyles not exserted, covering the orificeStyles exserted, about half the length of the inner stamensSepals entire or only with very few appendices, sepals conspicuously

longer than petals, leaflets not glandular and glabrousOuter sepals significantly pinnate, if entire then leaflets glandular and ± hairyPrickles straight or curved (like a sickle), homoacanthPrickles hooked, sometimes heteracanth (hooked and straight mixed)Leaflets glabrous on the surface, underneath glabrous

or hairy, but not pannose. Veins often prominent like a reliefand glandular. Rhachis with glands. In all, glands more noticeable than hairs.Pinnate sepals deciduous after anthesis, pedicel glandular or with gland-tipped bristles.Leaves multiserrate with glands, fresh leaves not smelling like resin or turpentine

Leaflets conspicuously hairy on both sides. Leaflets underneath with glandshidden in the hairs. Hairs more noticeable than glands. Fresh leaves smellinglike resin or turpentine. Sepals sometimes ± entire

Leaflets sticky, at least on the underside, with numerous glands smelling likeapples or vine. Leaflets glabrous or hairy, glands more noticeable than hairs,hooked prickles sometimes subheteracanth-heteracanth, mixed withgland-tipped bristles

Leaflets not sticky-glandular, leaflets glabrous or hairy, aglandular or withnon-smelling glands

Leaflets hairy, at least always on the glandular midrib, mostly also the rhachis,the veins, the margins of the stipules and the multiserrate leaflet-marginwith glands. ± equal impression of glands and hairs

Leaflets glabrous or hairy, aglandular or sparely glandular on rhachis,veins and margins of the stipules. Leaflets uniserrate-multiserrate.If hairy, then hairs more prominent than glands. Confusion possible betweenglandular forms of R. corymbifera and subsect. Tomentellae

Subgenus Hulthemia24, subgenus Rosa3Subgenus Hesperhodos

Subgenus Platyrhodon75Sect. 10: Bracteatae6Sect. 9: LaevigataeSect. 8: Banksianae8

9Sect. 2: Rosa15 (sect. 3: Caninae)Sect. 3: subsect. Rubrifoliae

10Sect. 3: subsect. Vestitae11Sect. 6: Synstylae121314Sect. 1: PimpinellifoliaeSect. 5: CinnamomeaeSect. 4: CarolinaeSect. 7: IndicaeSubsect. Rubrifoliae

161718Subsect. Trachyphyllae

Subsect. Vestitae

Subsect. Rubigineae

19

Subsect. Tomentellae

Subsect. Caninae

=xistence of three major groups which are mostly-erged together: subsect. Pimpinellifoliae, Luteae::ep. and Sericeae Cn§p. Chromosome numbers

own with 2n = 2x, 4x = 14, 28, balanced. R. ecae

Aitch 1880 (=R. mogoltavica ]uz.), R. foetida J.Herrm. 1762 (=R. lutea Mill.) (2n = 4x = 28),R. hemisphaerica J. Herrm. 1762. R. hugonis Hemsl.1905 (2n = 2x = 14), R. koreana Komar. 1901

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_.............114 CLASSIFICATION/Conventional Taxonomy (Wild Roses)

(2n=2x=14), R. myriacantha DC 1805 (2n=4x = 28), R. omeiensis Rolfe 1912 (2n =2x=14), R. primula Boul. 1936 (2n=2x=14)(=R. ecae Kanitz, non Ait.), R. sericea Lindl. 1820(=R. tetrapetala Royle, R. wallichii Tratt.) (2n =2x = 14). The morphological difference between R.omeiensis and R. sericea is the thickened stalk of R.omeiensis. Usually R. sericea is attributed as havingonly four sepals and petals, but also forms with fiveexist, R. spinosissima L. 1753 (=R. pimpinellifoliaL.): see Rehder no. 2 (2n = 4x = 28). There is still agreat deal of uncertainty about the subdivision of thespecies. Especially under the name R. sp. var. altaica(Willd.) Rehd. or R. altaica Willd., several gene poolsexist which are used for horticultural purposes. R.xanthina Lindl. 1820 (2n=2x=14).

Sect. 2. Rosa (=sect. Gallicanae (DC) Sec1825) One species in Europe and W. Asia. R. gall­ica L. 1759 (nomen ambiguum R. gallica L. 1753)(2n = 4x = 28) (=R. provincialis Herrm., R. austriacaCrantz, R. pumila Scopoli non ]acq, nom. illegit.).Section Rosa harbours a number of taxa given speciesrank. These taxa are of hybrid origin, long cultivated,the status and knowledge of natural occurrenceare uncertain. All species have (2n = 4x = 28):R. alba L. 1753. R. centi/olia L. 1753 (=R. gallicavar. centifolia (L.) Regel). R. damascena Mill. 1768(=R. gallica var. damascena Voss). R. francofurtanaMiinchhausen 1774 (=R. turbinata Ait.). R.polliniana Spreng. 1813.

Sect. 3. Caninae (DC) Ser. 1825 Stems upright orarching with hooked, slender or straight prickles,outer sepals lobed (three exceptions), sepals persistentand erect, reflexed-deciduous or patent after anthesis.All members characterized by the Caninae meiosiswith unbalanced heterogamous fully sexual reproduc­tion. About 50 species in Europe.

Subsect. Trachyphyllae H. Christ 1873 R. jundzilliiBesser 1815 (=R. marginata Wallr., R. trachyphyllaA. Rau): see Rehder no. 19 (R. marginata Wallr.) (2n= 6x = 42, unbalanced, heterogamous). Of hybrid or­igin, maternal parent from sect. Caninae, paternal par­ent R. gallica L.

Subsect. Rubrifoliae Crip. 1892 R. glauca Pourr.1788 (=R. ferruginea ViII., R. rubrifolia Vill.; nonR. glauca Vill.): see Rehder no. 20 (R. rubrifoliaViII.) (2n = 4x = 28, unbalanced, heterogamous).The smooth or glandular entire sepals sometimeswith lateral appendages. Morphology places thespecies into Cinnamomeae, but meiosis is ofCaninae type.

Subsect. VestitaeH. Christ 1873 R.mollisSm.181'='(=R. mollissima Fr., R. pomifera subsp. mol.'_(Sm.) Schwenschl.): see Rehder no. 13 (2n = 4x, -.~

6x = 28, 35, 42, unbalanced heterogamous) floweLpink-red, white. R. pseudoscabriuscula (R. Kelle­1931: 276) Henker & G. Schulze 1993 (=R. tome';­tosa subsp. pseudoscabriuscula R. Keller) (211 =5x = 35, unbalanced, heterogamous). Shrub up :2 m, prickles slender and curved, homoacanth: leafle­(5}-7, multiserrate and glandular, underneath wi-­glands hidden in the pubescence covering both sid­of the leaf. Rachis pubescent and glandular, sepaspreading but often erect at hip ripening. Flowe::pink. R. sherardii Davies 1813 (=R. omissa Desegl.see Rehder no. 13 (2n = 4x, 5x, 6x = 28, 35, ­unbalanced, heterogamous) flowers dark pink-re­R. tomentosa Sm. 1800 (=R. cinerascens Dumon.see Rehder no. 14 (2n = 5x = 35, unbalanced, here;-­ogamous). Prickles slender and curved, not straigl::..Leaflets mostly uniserrate with conspicuous wide all':short teeth; if biserrate than a slight occurrence ­glands, sepals reflexed and deciduous, flowers sligh .pink-white. R. villosa L. 1753 (=R. pomifera Herrm.see Rehder no. 13 (2n = 4x, 8x = 28,56, unbalance­heterogamous), flowers pink-red, white nail.

Subsect. Rubigineae H. Christ 1873 R. agresSavi 1798 (=R. sepium Thuill.): see Rehder n16 (2n = 5x, 6x = 35, 42, unbalanced heterogamoussepals reflex and deciduous after anthesis. Back ­sepals without glands. R. elliptica Tausch 1 1­(=R. graveolens Gren. et Godr.): see Rehder no. 1:(synonym of R. inodora) (2n = 5x, 6x = 35,unbalanced heterogamous), sepals erect, persiste-­after anthesis. Flowers pinkish. R. inodora Fr. 181­(=R. elliptica subsp. inodora Fries), see Rehd-­no. 15 (2n = 5x, 6x = 35, 42, unbalanced hetero­gamous), somewhat of an intermediate species be­tween R. agrestis and R. elliptica, sepals patent ar::.deciduous. R. micrantha Borrer ex. Sm. 1812:Rehder no. 16 (2n = 4x, 5x, 6x = 28, 35,unbalanced, heterogamous), usually small hips \yj-­

reflexed and deciduous sepals early after anthe isR.rubiginosa L. 1771 (=R. eglanteria L., R. umbel! ~Leers): see Rehder 15 (synonym of R. eglanteri­(2n = 5x = 35, unbalanced, heterogamous), aroma :.smelling leaf (apple), sepals persistent and ereR. rubiginosa subsp. columniftra Schwertschl. 191(=R. columnifera (Schwertschlager) HenkerG. Schulze 2000 (homonym illeg., non Fries 181­(2n = 5x = 35 also 6x?, unbalanced, heterogamou:intermediate species combining charactersR. rubiginosa and R. micrantha. Sepals patent apartly deciduous, flowers pinkish-white.

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CLASSIFICATION/Conventional Taxonomy (Wild Roses) 115

;'sect. Tomentellae H. Christ 1873 R. tomentellan in Cassini 1818 (R. obtusifolia auct. (mult.)

-::. Desv.) (2n = 5x = 35, unbalanced, heteroga-- . ). Shrub up to 2 m, homoacanth, prickles curved.~-jets 5-7, mostly pubescent on both sides, veins':'~rneath the leaflets densely covered with red--ds which do not smell, increase of glands towards~ margin of the leaflets. Rhachis pubescent and with-- s. Sepals reflexed and deciduous after anthesis.~::: tacle usually smooth as well as the pedicel,~'ers white. R. abietina Gren. ex Christ 1873

=.K obtusifolia subsp. abietina (Gren. ex Christ)- -:-ferrmann) (2n = ?, unbalanced, heterogamous).

--:.Ib about 1.5 m high, prickles slender or hooked.~O' ets (5)-7, both sides or only underneath pu­-:-ent, always with glands. Pedicels stipitate-glandu--. ack of the sepals covered with glands. Sepals

=::;:x and deciduous. Flowers reddish-pink.

:-sect. Caninae R. caesia Sm. 1812 (=R. coriifolia-:: .: see Rehder no. 18 (synonym R. coriifolia)- . = 5x, 6x = 35, 42, unbalanced heterogamous).

: Is first patent after anthesis, than erect and- -,::l'tent. Flowers intensely pink. R. canina L.

- ~ oJ (=R. nitidula Besser, R. afzeliana Fr., R.-~iana Leman, R. squarrosa (A. Rau) Boreau, R.

,--Z'r ta Crep., R. canina agg. Ehrendorfer): see~o er no. 19 (2n = 5x = 35, unbalanced, hetero­'~ous). Leaflets and rhachis not pubescent. Sepals

- ::;:x and deciduous after anthesis. Flowers pinkish­'::"::e. R. corymbiftra Borkh. (R. dumetorum Thuil!.,

btusifolia Desv., R. deseglisei Boreau, R. brilon­..::s G.H. Loos): see Rehder no. 18 (2n=5x=35,. lanced, heterogamous). Leaflets pubescent on

. ::l ides or only underneath or even only the rhachis..:.als reflex and deciduous after anthesis. Flowers.:ite, sometimes pinkish. R. dumalis Bechst. 1819

=R. vosagiaca Deseg!., R. glauca Vii!. ex Loisel, R.-__.~eri (Godet) Reuter): see Rehder no. 19 (2n = 5x,-~=35, 42, unbalanced heterogamous). Sepals first-:-:::.: fit after anthesis, than erect and persistent.- montana Chaix in Villars 1786: see Rehder no.

2 (2n = 5x = 35, unbalanced, heterogamous).-:: stylosa Desvaux 1809: see Rehder no. 17~. = 5x, 6x = 35, 42, unbalanced, heterogamous).~3J1ets 5-7, the single one often formed like a flame~ dandle. Pedicels glandular and very much longer

=..L'1 the eglandular receptacle. Sepals on the back_' out glands, reflexed and deciduous after anthesis.

::. "oers pinkish-white. Styles agglutinated but not;.: :mate as in section 6: Synstylae. Disc formed like_ :::one. R. subcanina (H. Christ) R. Keller 1891=R. reuteri f. subcanina H. Christ, R. dumalis:::.::-sp. subcanina (H. Christ) S06) (2n = 5x = 35, un­-~anced, heterogamous). Shrub up to 2.5 m, interme-

diate species between R. canina and R. dumalis, sepalspatent and deciduous. Flowers pale pink. R. sub­collina (H. Christ) R. Keller 1891 (=R. coriifolia f.subcollina H. Christ; R. caesia subsp. subcollina(H. Christ) S06) (2n = 5x = 35, unbalanced, heterog­amous). Shrub up to 3 m, intermediate species be­tween R. corymbifera and R. caesia, sepals patentand deciduous. Flowers pink.

Sect. 4. Carolinae Crep. 1891 5(?) species fromNorth America. R. carolina L. 1753 (2n=4x=28)(=R. humilis Marsh., R. lyonii Pursh, R. serrulataRafinesque). R.foliolosa Nutt. ex. Torr. & A. Gray1840 (2n=2x=14). R. nitida Willd. 1809. R.

. palustris Marsh 1785 (2n = 2x = 14) (=R. carolinaGray non L.) R. virginiana Herrm. 1762(2n=4x=28) (=R. virginiana Mill. 1768, in mostcases the citation of the author for this species iswrong; the name by Miller is a younger homonym).

Sect. 5. Cinnamomeae (DC) Ser. 1825 Uncertainnumber of species (about 80) in Asia, North America,Europe. R. acicularis Lindl. 1820 (2n = 4x, 8x = 28,56) (=R. alpina Pall. non L., R. granulosa Keller,R. gmelinii Bunge). R. amblyotis CA. Mey. 1847(2n = 2x = 14) (=R. davurica Hulten non PalL).R. arkansana Porter ex LM. Coult. 1874 (2n =4x = 28) (=R. rydbergii Greene). R.banksiopsis Baker1914 (2n=2x= 14). R. beggeriana Schrenk 1841(2n = 2x = 14). R. bella Rehd. & E.H. Wilson 1915(2n = 4x = 28). R. blanda Ait. 1789 (2n = 2x = 14).R. davurica Pal!. 1788 (2n = 2x = 14) (=R.cinnamomea Ldb. non L., R. wildenowii Sprengel).R. californica Cham. et Schltdl. 1827 (2n =4x = 28). R. caudata Baker 1914 (2n = 2x, 4x = 14,28). R. corymbulosa Rolfe 1914 (2n=2x=14) ..R. davidii Crep. 1874 (2n = 4x = 28). R. ftdts­chenkoana Regel 1878 (2n = 4x = 28). (=R.caraganifolia Sumn., R. coeruleifolia Sumn., R.epipslia Sumn., R. laurenkoi Sumn., R. lipschitziiSumn., R. minusculifolia Sumn., R. oligospermaSumn.). R. forrestiana Boulenger 1936. (2n =4x = 28), R. gymnocarpa Nutt. ex Torr. et A. Gray1840 (2n=2x=14). R. laxa Retz 1803 (2n=2x = 14) (=R. soongarica Bge., R. alpina Ldb. nonL.). R. majalis Herrm. 1762 (wild: 2n=2x=14 ;hart.: 4x, 8x = 28, 56) (=R. cinnamomea L. 1759non 1753, R. spinosissima Rydb.). R. marretii H.Lev. 1910 (2n = 2x = 14). R. moyesii Hemsl. & E.H.Wilson 1906 (2n = 4x = 28). R. multibracteataHemsl. et E.H. Wilson 1906 (2n = 4x = 28) (=R.latibracteata Bou!., R. orbicularis Baker, R. reductaBaker, R. rotundibracteata Cardot). R. nanothamnusBoulenger 1935 (2n = 4x = 28) (R. beschnauensisSumn., R. botryoides Sumn., R. vitaminifera

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