CLASSIFICATION/Conventional Taxonomy (Wild Roses) 111-:acey GN, Lynch PT and Benson EE (1999) Plant gene General Aspects, pp. 26-49. Enfield, NH: Science banking: agriculture, biotechnology and conservation. Publishers. Agro Food Industry Hi-Tech 10: 9-14. Towill LE (2002) Cryopreservation of plant germplasm:::~aritsky G (1997) Background and principles of in introduction and some observations. In: Towill LE and vitro conservation of plant genetic resources. In: Bajaj YPS (eds.) Biotechnology in Agriculture and Razdan MK and Cocking EC (eds.) Conservation Forestry, vol 50: Cryopreservation of Plant Germplasm of Plant Genetic Resources In Vitro, vol 1, II, pp. 3-21. Berlin: Springer Verlag.CLASSIFICATION Contents Conventional Taxonomy (Wild Roses) Horticultural Classification Schemes Cultivar Identification by Image Analysis Chemotaxonomy and Molecular Taxonomy Conventional Taxonomy to several attempts to classify the genus. During the Renaissance, in Baroque times and up to the early (Wild Roses) eighteenth century, rose systems were just divided V Wissemann, Friedrich-Schiller-Universitat Jena, into wild and gentle species, followed by subdivi- Jena, Germany sions based on petal colour. The recognition of 1) 2003, Elsevier Ltd. All Rights Reserved. hybridization in roses was first remarked by Linnaeus, who wrote in the Species Plantarum of 1753 that he had the impression that Rosa species are difficult toIntroduction distinguish, more difficult to determine, and that nature just for fun mixes species to form new ones.Taxonomy of wild roses is notoriously difficult, for a Also Herrmann claimed in his thesis from Strasbourgariety of reasons. On the one hand the enormous in 1762 that species are very difficult to determine on henotypic variability is based on genetic complexi- the one hand because of the lack of characters, and on ity, while on the other hand the close relatedness of the other hand because horticulture had merged the he genomes and fea tures of the life cycle have begun to species so that recognition of pure species was nobe clarified in recent years. However, traditional tax- longer possible.onomy based on morphological and anatomical data The first attempt to create a real system of Rosa wasis in itself subject either to biological phenomena such made by Linnaeus. He regarded the shape of the hip toas reproductive biology and ecological variability, or be useful for classification, and this character was ac-ro theoretical problems like What is a species? and cepted nearly exclusively until the beginning of theHow can biological diversity be translated into taxo- nineteenth century. Willdenow remarked in 1811nomically and nomenclaturally valid studies? The that form and presence of prickles as well as hairsmost common system in use is the classification by and glands can serve as species-specific characters._-lfred Rehder (see vol. 3 of this Encyclopedia): nev- This remark led to several new systems which wereertheless, there is an urgent need for an update of based more on personal preference rather than on ob-that system, for both nomenclatural and biological servations in nature. The application of various char-reasons. In this article, methods of rose classification acters to treat the genus systematically was the firstare discussed and the Rehder classification is updated. sign of the enormous increase in names up to the be- ginning of the twentieth century (Table 1). The numberClassification of the Genus Rosa of different studies on the genus Rosa at this time isRoses have influenced human cultural history in not a sign of real insight; it is an expression of uncer-yarious ways and the enormous interest in roses led tainty facing the bewildering diversity of the genus.
112 CLASSIFICATION/Conventional Taxonomy (Wild Roses)Table 1 Number of species recognized from the sixteenth to In this article the key for identification of Rehdersthe nineteenth century main categories is replaced by a key oriented on theGessner (1561): 10 species whole species diversity. Nevertheless, morphologicalDodonaeus (1569, 1616): 10 species classification is critical in sections like CinnamomeaeLobelius (1581): 11 species versus Pimpinellifoliae, where unique characters forGerard (1597): 16 species the sections are lacking, and where molecular dataSpigelius (1633): 16 species from both ribosomal and chloroplast DNA indicateElsholz (1663) 15 speciesWelsch (1697): 18 species that morphological partition is artificial. Another ex-Salmon (1710): 12 species and 32 cultivars ample is the circumscription of section Carolinae.Linnaeus (1735): 12 species These roses are generally described as not stoutly built,Linnaeus (1772): 10 species but they do not show any significant specific charactersForsy1h (1794): 28 species for a sectional classification. Looking at these roses inLaicharding (1794): 31 speciesWilldenow (1811): 34 species their natural habitat, one can instantly see that theirSmith (1819): 57 species life in the shadow under trees in the woods forces suchTrattinnick (1823): 24 series with > 200 species a fragrant growth with single flowers. However, weSeringe (1825): 146 species know from cultivation experiments in other sectionsLindley (1830): 101 species, about 300 synonyms that these characters are strongly dependent on eco-Reichenbach (1832): 77 species, about 200 synonymsDoll (1855): 114 species, about 200 synonyms logical features like sun intensity, temperature andDeseglise (1877): 15 sections, section Caninae alone humidity, but there is no knowledge on the plasticity with 329 species of these phenotypic characters in section Carolinae, so section and species delimitation may not be natural. Changes in the Rehder SystemHowever, not all members of the genus Rosa faced thesame problem. Mainly the dogroses, section Caninae, From a nomenclature point of view, the subgenuswere subject to intensive species splitting.This culmi- Eurosa must be replaced by the subgenus Rosa. Basednated in 1886 in the article on buissonnomanie by on generic type, the subgenus Rosa contains sec-Crepin, who vehemently accused the rhodologists of tion Rosa. Section Rosa replaces section Gallicanae,naming each rosebush with a new name. because R. centifolia L. is the generic type of Rosa. The change in understanding of the biological di- Following the rules of the International Code ofversity of European wild roses started in 1873 with the Botanical Nomenclature, the first lectotypificationstudy of the Swiss rhodologist Christ, who reduced the by Britton & Brown in 1913 on R. centifolia as theinflation of names to about 30 species by applying his generic type is still a valid choice, but is disputed. Thesynthetic classification system. This synthetic method proposal to replace this choice by R. cinnamomeaewas based on classification considering combinations was rejected by the general committee for nomencla-of preferable correlated characters. In contrast to the ture at the Tokyo Botanical Congress in 1995. Theanalytical methods applied before Christ, his approach proposal was again not accepted in the Saint Louiscircumscribed taxa which he recognized as being nat- Code of Botanical Nomenclature in 2000. A revisedural groups, rather than artificial entities. However, proposal will be discussed at the Vienna Congressmolecular data clearly indicate (see Classification: 2005 (Table 2).Chemotaxonomy and Molecular Taxonomy) thatthese groups have to be revised again. Going back Addenda and Corrections to thein the history of rhodology it is obvious that the tax- System of Rehder 1940onomic treatment of the genus Rosa is influenced by a Subgenus Hulthemia (Dumort.) Focke 1888European view of the genus. Systems available todayshow that, based on the increase in knowledge of bi- Monotypic subgenus: Rosa persica Michx. ex Juss.ological diversity, there is a strong need for systematic 1789 (=R. simplicifolia Salisb., R. berberifolia Pall.,and taxonomic studies in roses worldwide. Whereas Hulthemia berberifolia (Pall.) Dumort., Hulthemiathe knowledge of middle European roses is quite ad- persica Bornmiiller, Lowea berberifolia Lindl.)vanced, next to nothing is known about eastern (2n = 2x = 14).European species, and Asian species and even north Subgenus RosaAmerican species are quite unknown from a system-atic and ecological point of view. Thus morphological Sect. 1. Pimpinellifoliae (DC.) Ser. 1825 Aboutclassification of the genus results in at least some 15 species from Asia and Europe. Taxonomic sub-unnatural groups. divisions uncertain. Molecular data support the
CLASSIFICATION/Conventional Taxonomy (Wild Roses) 113-= Ie 2 Key to the sections of Rosa Leaves entire (1 leaflet), without stipules Subgenus Hulthemia Leaves imparipinnate ( > 1 leaflet), with or without stipules 2 Receptacle smooth or ± gland-tipped acicles or bristles 4, subgenus Rosa Receptacle with prickles like a chestnut, sepals pinnate and erect 3 Stipules adnate, with divergent, rounded or broadened auricles, leaflet Subgenus Hesperhodos usually 3 or 5, rarely 7 Stipules adnate, with subulate auricles, leaflets usually > 7 (-15) Subgenus Platyrhodon Stipules persistent and half-adnate with auricles 7 Stipules free and ± deciduous 5 Receptacle and younger branchlets tomentose or pubescent Sect. 10: Bracteatae Not this combination 6 Sepals erect and persistent after anthesis, receptacle bristly Sect. 9: Laevigatae Sepals reflexed and deciduous, receptacle smooth Sect. 8: Banksianae Sepals conspicuously pinnate (the two sepals outside pinnate on both sides, 8 the next one only on one side and all of the two inner sepals Sepals entire or only the two outer ones with slight appendices 9:: Prickles glabrous, straight and mixed with gland-tipped bristles Sect. 2: Rosa Prickles straight or curved, not mixed with bristles 15 (sect. 3: Caninae) Sepals much longer than petals, younger stems with a purplish bloom, Sect. 3: subsect. Rubrifoliae leaflets with purple pigmentation, prickles straight or curved= Not this combination 10 Sepals short, erect, leaflets glandular and ± hairy, low and compact shrub Sect. 3: subsect. Vestitae Not this combination 11 Styles agglutinated after anthesis like a column Sect. 6: Synstylae Styles free 12 Sepals after anthesis erect and persistent 13 Sepals after anthesis reflexed or spreading, deciduous 14 Flowers mostly solitary, solitary flowers without bracts Sect. 1: Pimpinellifoliae Flowers solitary or corymbose; if solitary, then with bracts Sect. 5: Cinnamomeae Styles not exserted, covering the orifice Sect. 4: Carolinae Styles exserted, about half the length of the inner stamens Sect. 7: Indicae Sepals entire or only with very few appendices, sepals conspicuously Subsect. Rubrifoliae longer than petals, leaflets not glandular and glabrous Outer sepals significantly pinnate, if entire then leaflets glandular and ± hairy 16 Prickles straight or curved (like a sickle), homoacanth 17 Prickles hooked, sometimes heteracanth (hooked and straight mixed) 18 Leaflets glabrous on the surface, underneath glabrous Subsect. Trachyphyllae or hairy, but not pannose. Veins often prominent like a relief and 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 glands Subsect. Vestitae hidden in the hairs. Hairs more noticeable than glands. Fresh leaves smelling like resin or turpentine. Sepals sometimes ± entire Leaflets sticky, at least on the underside, with numerous glands smelling like Subsect. Rubigineae apples or vine. Leaflets glabrous or hairy, glands more noticeable than hairs, hooked prickles sometimes subheteracanth-heteracanth, mixed with gland-tipped bristles Leaflets not sticky-glandular, leaflets glabrous or hairy, aglandular or with 19 non-smelling glands Leaflets hairy, at least always on the glandular midrib, mostly also the rhachis, Subsect. Tomentellae the veins, the margins of the stipules and the multiserrate leaflet-margin with glands. ± equal impression of glands and hairs Leaflets glabrous or hairy, aglandular or sparely glandular on rhachis, Subsect. Caninae veins and margins of the stipules. Leaflets uniserrate-multiserrate. If hairy, then hairs more prominent than glands. Confusion possible between glandular forms of R. corymbifera and subsect. Tomentellae=xistence of three major groups which are mostly Aitch 1880 (=R. mogoltavica ]uz.), R. foetida J.-erged together: subsect. Pimpinellifoliae, Luteae Herrm. 1762 (=R. lutea Mill.) (2n = 4x = 28),::ep. and Sericeae Cn§p. Chromosome numbers R. hemisphaerica J. Herrm. 1762. R. hugonis Hemsl. own with 2n = 2x, 4x = 14, 28, balanced. R. ecae 1905 (2n = 2x = 14), R. koreana Komar. 1901
" , . . . -: ,=~~~ I _ - - ~ ............. _ 114 CLASSIFICATION/Conventional Taxonomy (Wild Roses) (2n=2x=14), R. myriacantha DC 1805 (2n= Subsect. VestitaeH. Christ 1873 R.mollisSm.181= 4x = 28), R. omeiensis Rolfe 1912 (2n = (=R. mollissima Fr., R. pomifera subsp. mol._ 2x=14), R. primula Boul. 1936 (2n=2x=14) (Sm.) Schwenschl.): see Rehder no. 13 (2n = 4x, -.~ (=R. ecae Kanitz, non Ait.), R. sericea Lindl. 1820 6x = 28, 35, 42, unbalanced heterogamous) floweL (=R. tetrapetala Royle, R. wallichii Tratt.) (2n = pink-red, white. R. pseudoscabriuscula (R. Kelle- 2x = 14). The morphological difference between R. 1931: 276) Henker & G. Schulze 1993 (=R. tome;- omeiensis and R. sericea is the thickened stalk of R. tosa subsp. pseudoscabriuscula R. Keller) (211 = omeiensis. Usually R. sericea is attributed as having 5x = 35, unbalanced, heterogamous). Shrub up : only four sepals and petals, but also forms with five 2 m, prickles slender and curved, homoacanth: leafle- exist, R. spinosissima L. 1753 (=R. pimpinellifolia (5}-7, multiserrate and glandular, underneath wi-- L.): see Rehder no. 2 (2n = 4x = 28). There is still a glands hidden in the pubescence covering both sid- great deal of uncertainty about the subdivision of the of the leaf. Rachis pubescent and glandular, sepa species. Especially under the name R. sp. var. altaica spreading but often erect at hip ripening. Flowe:: (Willd.) Rehd. or R. altaica Willd., several gene pools pink. R. sherardii Davies 1813 (=R. omissa Desegl. exist which are used for horticultural purposes. R. see Rehder no. 13 (2n = 4x, 5x, 6x = 28, 35, - xanthina Lindl. 1820 (2n=2x=14). unbalanced, heterogamous) flowers dark pink-re- R. tomentosa Sm. 1800 (=R. cinerascens Dumon. Sect. 2. Rosa (=sect. Gallicanae (DC) Sec see Rehder no. 14 (2n = 5x = 35, unbalanced, here;-- 1825) One species in Europe and W. Asia. R. gall- ogamous). Prickles slender and curved, not straigl::.. ica L. 1759 (nomen ambiguum R. gallica L. 1753) Leaflets mostly uniserrate with conspicuous wide all: (2n = 4x = 28) (=R. provincialis Herrm., R. austriaca short teeth; if biserrate than a slight occurrence - Crantz, R. pumila Scopoli non ]acq, nom. illegit.). glands, sepals reflexed and deciduous, flowers sligh . Section Rosa harbours a number of taxa given species pink-white. R. villosa L. 1753 (=R. pomifera Herrm. rank. These taxa are of hybrid origin, long cultivated, see Rehder no. 13 (2n = 4x, 8x = 28,56, unbalance- the status and knowledge of natural occurrence heterogamous), flowers pink-red, white nail. are uncertain. All species have (2n = 4x = 28): R. alba L. 1753. R. centi/olia L. 1753 (=R. gallica var. centifolia (L.) Regel). R. damascena Mill. 1768 Subsect. Rubigineae H. Christ 1873 R. agres (=R. gallica var. damascena Voss). R. francofurtana Savi 1798 (=R. sepium Thuill.): see Rehder n Miinchhausen 1774 (=R. turbinata Ait.). R. 16 (2n = 5x, 6x = 35, 42, unbalanced heterogamous polliniana Spreng. 1813. sepals reflex and deciduous after anthesis. Back - sepals without glands. R. elliptica Tausch 1 1- Sect. 3. Caninae (DC) Ser. 1825 Stems upright or (=R. graveolens Gren. et Godr.): see Rehder no. 1: arching with hooked, slender or straight prickles, (synonym of R. inodora) (2n = 5x, 6x = 35, outer sepals lobed (three exceptions), sepals persistent unbalanced heterogamous), sepals erect, persiste-- and erect, reflexed-deciduous or patent after anthesis. after anthesis. Flowers pinkish. R. inodora Fr. 181- All members characterized by the Caninae meiosis (=R. elliptica subsp. inodora Fries), see Rehd-- with unbalanced heterogamous fully sexual reproduc- no. 15 (2n = 5x, 6x = 35, 42, unbalanced hetero- tion. About 50 species in Europe. gamous), somewhat of an intermediate species be- tween R. agrestis and R. elliptica, sepals patent ar::. Subsect. Trachyphyllae H. Christ 1873 R. jundzillii deciduous. R. micrantha Borrer ex. Sm. 1812: Besser 1815 (=R. marginata Wallr., R. trachyphylla Rehder no. 16 (2n = 4x, 5x, 6x = 28, 35, A. Rau): see Rehder no. 19 (R. marginata Wallr.) (2n unbalanced, heterogamous), usually small hips yj-- = 6x = 42, unbalanced, heterogamous). Of hybrid or- reflexed and deciduous sepals early after anthe is igin, maternal parent from sect. Caninae, paternal par- R.rubiginosa L. 1771 (=R. eglanteria L., R. umbel! ~ ent R. gallica L. Leers): see Rehder 15 (synonym of R. eglanteri- (2n = 5x = 35, unbalanced, heterogamous), aroma :. Subsect. Rubrifoliae Crip. 1892 R. glauca Pourr. smelling leaf (apple), sepals persistent and ere 1788 (=R. ferruginea ViII., R. rubrifolia Vill.; non R. rubiginosa subsp. columniftra Schwertschl. 191 R. glauca Vill.): see Rehder no. 20 (R. rubrifolia (=R. columnifera (Schwertschlager) Henker ViII.) (2n = 4x = 28, unbalanced, heterogamous). G. Schulze 2000 (homonym illeg., non Fries 181- The smooth or glandular entire sepals sometimes (2n = 5x = 35 also 6x?, unbalanced, heterogamou: with lateral appendages. Morphology places the intermediate species combining characters species into Cinnamomeae, but meiosis is of R. rubiginosa and R. micrantha. Sepals patent a Caninae type. partly deciduous, flowers pinkish-white.
,,,;,".;~..~ ;. _ • • --- ~_:::- ~ ,- r .~.. _~~._"" ...: , ,~ - .".- . - .- CLASSIFICATION/Conventional Taxonomy (Wild Roses) 115 ;sect. Tomentellae H. Christ 1873 R. tomentella diate species between R. canina and R. dumalis, sepals n in Cassini 1818 (R. obtusifolia auct. (mult.) patent and deciduous. Flowers pale pink. R. sub- -::. Desv.) (2n = 5x = 35, unbalanced, heteroga- collina (H. Christ) R. Keller 1891 (=R. coriifolia f. - . ). Shrub up to 2 m, homoacanth, prickles curved. subcollina H. Christ; R. caesia subsp. subcollina ~-jets 5-7, mostly pubescent on both sides, veins (H. Christ) S06) (2n = 5x = 35, unbalanced, heterog- :~rneath the leaflets densely covered with red amous). Shrub up to 3 m, intermediate species be- --ds which do not smell, increase of glands towards tween R. corymbifera and R. caesia, sepals patent ~ margin of the leaflets. Rhachis pubescent and with and deciduous. Flowers pink. -- s. Sepals reflexed and deciduous after anthesis. ~::: tacle usually smooth as well as the pedicel, Sect. 4. Carolinae Crep. 1891 5(?) species from ~ers white. R. abietina Gren. ex Christ 1873 North America. R. carolina L. 1753 (2n=4x=28) =.K obtusifolia subsp. abietina (Gren. ex Christ) (=R. humilis Marsh., R. lyonii Pursh, R. serrulata - -:-ferrmann) (2n = ?, unbalanced, heterogamous). Rafinesque). R.foliolosa Nutt. ex. Torr. & A. Gray --:.Ib about 1.5 m high, prickles slender or hooked. 1840 (2n=2x=14). R. nitida Willd. 1809. R . ~O ets (5)-7, both sides or only underneath pu- . palustris Marsh 1785 (2n = 2x = 14) (=R. carolina -:-ent, always with glands. Pedicels stipitate-glandu- Gray non L.) R. virginiana Herrm. 1762 -. ack of the sepals covered with glands. Sepals (2n=4x=28) (=R. virginiana Mill. 1768, in most =::;:x and deciduous. Flowers reddish-pink. cases the citation of the author for this species is wrong; the name by Miller is a younger homonym). :-sect. Caninae R. caesia Sm. 1812 (=R. coriifolia -:: .: see Rehder no. 18 (synonym R. coriifolia) Sect. 5. Cinnamomeae (DC) Ser. 1825 Uncertain - . = 5x, 6x = 35, 42, unbalanced heterogamous). number of species (about 80) in Asia, North America, : Is first patent after anthesis, than erect and Europe. R. acicularis Lindl. 1820 (2n = 4x, 8x = 28, - -,::ltent. Flowers intensely pink. R. canina L. 56) (=R. alpina Pall. non L., R. granulosa Keller, - ~ oJ (=R. nitidula Besser, R. afzeliana Fr., R. R. gmelinii Bunge). R. amblyotis CA. Mey. 1847 -~iana Leman, R. squarrosa (A. Rau) Boreau, R. (2n = 2x = 14) (=R. davurica Hulten non PalL). ,--Zr ta Crep., R. canina agg. Ehrendorfer): see R. arkansana Porter ex LM. Coult. 1874 (2n = ~o er no. 19 (2n = 5x = 35, unbalanced, hetero- 4x = 28) (=R. rydbergii Greene). R.banksiopsis Baker ~ous). Leaflets and rhachis not pubescent. Sepals 1914 (2n=2x= 14). R. beggeriana Schrenk 1841 - ::;:x and deciduous after anthesis. Flowers pinkish- (2n = 2x = 14). R. bella Rehd. & E.H. Wilson 1915 ::"::e. R. corymbiftra Borkh. (R. dumetorum Thuil!., (2n = 4x = 28). R. blanda Ait. 1789 (2n = 2x = 14). btusifolia Desv., R. deseglisei Boreau, R. brilon- R. davurica Pal!. 178 8 (2n = 2x = 14) (=R . ..::s G.H. Loos): see Rehder no. 18 (2n=5x=35, cinnamomea Ldb. non L., R. wildenowii Sprengel). . lanced, heterogamous). Leaflets pubescent on R. californica Cham. et Schltdl. 1827 (2n = . ::l ides or only underneath or even only the rhachis. 4x = 28). R. caudata Baker 1914 (2n = 2x, 4x = 14, .:.als reflex and deciduous after anthesis. Flowers 28). R. corymbulosa Rolfe 1914 (2n=2x=14) .. .:ite, sometimes pinkish. R. dumalis Bechst. 1819 R. davidii Crep. 1874 (2n = 4x = 28). R. ftdts- =R. vosagiaca Deseg!., R. glauca Vii!. ex Loisel, R. chenkoana Regel 1878 (2n = 4x = 28). (=R. -__.~eri (Godet) Reuter): see Rehder no. 19 (2n = 5x, caraganifolia Sumn., R. coeruleifolia Sumn., R. -~=35, 42, unbalanced heterogamous). Sepals first epipslia Sumn., R. laurenkoi Sumn., R. lipschitzii -:-:::.: fit after anthesis, than erect and persistent. Sumn., R. minusculifolia Sumn., R. oligosperma - montana Chaix in Villars 1786: see Rehder no. Sumn.). R. forrestiana Boulenger 1936. (2n = 2 (2n = 5x = 35, unbalanced, heterogamous). 4x = 28), R. gymnocarpa Nutt. ex Torr. et A. Gray -:: stylosa Desvaux 1809: see Rehder no. 17 1840 (2n=2x=14). R. laxa Retz 1803 (2n= ~. = 5x, 6x = 35, 42, unbalanced, heterogamous). 2x = 14) (=R. soongarica Bge., R. alpina Ldb. non ~3J1ets 5-7, the single one often formed like a flame L.). R. majalis Herrm. 1762 (wild: 2n=2x=14 ; ~ dandle. Pedicels glandular and very much longer hart.: 4x, 8x = 28, 56) (=R. cinnamomea L. 1759 =..L1 the eglandular receptacle. Sepals on the back non 1753, R. spinosissima Rydb.). R. marretii H. _ out glands, reflexed and deciduous after anthesis. Lev. 1910 (2n = 2x = 14). R. moyesii Hemsl. & E.H. ::. "oers pinkish-white. Styles agglutinated but not Wilson 1906 (2n = 4x = 28). R. multibracteata ;.: :mate as in section 6: Synstylae. Disc formed like Hemsl. et E.H. Wilson 1906 (2n = 4x = 28) (=R. _ :::one. R. subcanina (H. Christ) R. Keller 1891 latibracteata Bou!., R. orbicularis Baker, R. reducta =R. reuteri f. subcanina H. Christ, R. dumalis Baker, R. rotundibracteata Cardot). R. nanothamnus :::.::-sp. subcanina (H. Christ) S06) (2n = 5x = 35, un- Boulenger 1935 (2n = 4x = 28) (R. beschnauensis -~anced, heterogamous). Shrub up to 2.5 m, interme- Sumn., R. botryoides Sumn., R. vitaminifera. .-~~ =~--~~== - ------ - - --- - - - . I _~;.1