Volume 90 Annals Number 3 of the 2003 Missouri Botanical Garden SYSTEMATICS OF EURASIAN Ine´s A´lvarez Ferna´ndez2 AND NORTH AFRICAN DORONICUM (ASTERACEAE: SENECIONEAE)1 ABSTRACT ThegenusDoronicum(Asteraceae:Senecioneae)comprisesperennialherbsdistributedinEurope,NorthAfrica,and Asia.Aworldwiderevisionofthegenusrecognizing26speciesand4subspeciesispresented.Inthepresenttaxonomic treatmentnoinfragenericgroupsarerecognized.Sevennamesarenewlylectotypifiedherein:ArnicadoronicumJacq., Doronicum caucasicum M. Bieb., Doronicum portae Chabert, Doronicum scorpioides Lam., Doronicum souliei Cavill., Doronicum thibetanum Cavill., and Doronicum turkestanicum Cavill. A new chromosome count is provided for D. carpetanumsubsp.diazii. Keywords: Asia,Asteraceae,Doronicum,Europe,NorthAfrica,Senecioneae. The genus Doronicum L. (Asteraceae: Seneci- This genus belongs in the Senecioneae, one of oneae) includes rhizomatous herbs with yellow or the largest and most complex tribes in the Astera- green-tinted radiate capitula. All phyllaries are ceae with 123 genera and around 3200 species similar, generally herbaceous and arranged in two (Cassini, 1819; Bentham & Hooker, 1873b; Hoff- or three rows. Cypselae are cylindric to obovate- mann,1892;Nordenstam,1977;Bremer,1994).Its cylindric with10longitudinalribsandbearapap- 26 species constitute a presumably natural group pus of white-tinted minutely scabrous capillary (Bremer, 1994; involucre without shorter supple- bristles. The pappus can be absent in ray flowers mentary bracts, phyllaries herbaceous arranged in of some heterocarpic species. two or three rows, and cypselae cylindric to ob- 1IamgratefultoGonzaloNietoFelinerforsuggestingandguidingthisworkandforhisconstructivecommentson themanuscript,thecuratorsofAV,B,B-W,BC,BCF,BM,BOLO,BP,BR,BRNM,C,CL,COI-WILLK,E,FI,G,G- BOIS, G-DC, GAZI, GE, GH, GRM, GZU, HVR, IRAN, JACA, JE, K, LAU, LD, LE, LEB, LINN, LY, MA, MACB, MAF,MO,NAP,NY,P-HA,P-LA,RO,S,SANT,SZB,UPS,W,WU,ZA,andtheSa´nchezPedrajapersonalherbarium for the loan of specimens or for the information provided, Jasmin Jakupovik and Jesu´s Sanz for information and commentsonthechemistryofDoronicum,AlbertoHerreroandFe´lixMun˜oz-Garmendiafortheirhelpwithnomencla- turalmatters,aswellasJuanCastilloforthedrawings,Jesu´sMun˜ozformapdesign,andBernardoFerna´ndez,Miguel Jerez, and Raimundo Pradillo for technical assistance. This work has been supported by grant DGES PB96-0849 of theSpanishDireccio´nGeneraldeEnsen˜anzaSuperioreInvestigacio´nCient´ıfica. 2RealJardı´nBota´nico,CSIC,PlazadeMurillo,2,E-28014Madrid,[email protected] ANN. MISSOURI BOT. GARD. 90: 319–389. 2003. 320 Annals of the MissouriBotanicalGarden ovate-cylindric with 10 longitudinal ribs). Its geo- species and subspecies and an index to exsiccatae graphical area extends from Europe and North Af- are presented inAppendices1and2,respectively. rica to Asia, growing in mesic woods and open Observations were made directly or with theaid rockyplaceswithmoistsoil,andnearwatercourses, of binocular lenses. Microcharacters of indumen- from sea level to 5000 m of elevation. tum and cypselae were studied by SEM. Quanti- Althoughthereareseveralregionalstudiesofthe tative characters were recorded using a Brown & genus (Turkey, Edmondson, 1973,1975,1978;Ar- ShapePlusdigitalcaliper(model599-571-3).Mea- menia, Avetisyan, 1980; Iberian peninsula, Cha- surements were made on herbariumspecimens,af- co´n, 1987; Belgium, Duvigneaud, 1992), there is ter flattening and drying. onlyonepreviousworldwiderevisionofDoronicum Distribution maps for each taxon are based on (Cavillier,1907,1911).Fifteennewspecies(Diels, the specimens studied. Note that geographical ar- 1922; Widder, 1925; Sergievskaja, 1949; Widder eas and countries follow Hollis and Brummitt &Rechinger,1950;Edmondson,1973,1978;Cha- (1992), and major political divisions for countries co´n,1987;Pe´rez&Penas,1990;Pe´rezetal.,1994; wereincludedwhenthatinformationwasavailable. Chen, 1998) and six hybrids (Bornmu¨ller & Koch, 1930; Widder,1934, 1948;Stace,1991)weresub- TAXONOMIC HISTORY sequently described. Two speciesincludedinCav- ThenameDoronicumisapparentlyderivedfrom illier’s monograph have since been transferred to the Arabic word ‘‘darawnay,’’ used for at least two othergenera(i.e.,D.hookeriC.B.ClarkeexHook. different plants (Dozy, 1877). The pre-Linnaean to Nannoglottis (Kitamura, 1980), and D.thibetan- umCavill.toAster(A´lvarezFerna´ndez&NietoFel- botanists(Dioscorides,1554,1557;Dodoens,1574) and other Greek authors referred to speciesofDo- iner, 2000)). Cavillier (1907, 1911) studied the ronicum as Aconitum pardalianches, and the plant morphology of the genus in great detail, especially wasprobablyintroducedinWesternculturebyAv- the indumentum, but his proposed infrageneric icenna (Dodoens, 1574; Dalenchamps, 1587). classification is of rather limited value since these The genus Doronicum was described by Lin- groupsareobscurelydefinedbasedmainlyonnon- naeus (1753) to include four species, only two of exclusive characters. Asa result,classifyingnewly whicharecurrentlyacceptedinthegenus:D.par- described taxa in any infrageneric framework is dalianches and D. plantagineum. The remaining problematic. species correspond to Senecio and Aster, respec- The need to evaluate the newly described spe- tively.Further,onespeciesofArnicadescribedby cies and to assess the infrageneric taxonomy pro- Linnaeus(1753),A.scorpioidesL.,alsobelongsin videsjustificationforthiswork.Theobjectiveswere DoronicumasrecognizedbyLamarck(1786).Sev- to study as many morphological characters (quali- eral pre-Linnaean authors also confused species tative and quantitative) as possible so that(1)only of Senecio, Aster, and Arnica with Doronicum entities that could be consistently diagnosed were (Dalenchamps, 1587; Clusius, 1601; Bauhin, recognizedinthetaxonomictreatment,and(2)spe- 1623; Tournefort, 1700). In particular, the overall cieswereclassifiedasgroupsonthebasisofshared morphological similarity between Arnica and Do- synapomorphies (secondary homologies, De Pinna, ronicum suggested their close affiliation until the 1991). 1970s. However, Nordenstam’s (1977) micromor- phological study of style, anthers, and pollen de- MATERIAL AND METHODS finitively has excluded Arnica from the Seneci- oneae. More than 50 qualitative and quantitative mor- The cypsela dimorphism that occurs in some phological characters were studied in ca. 4300 specieshasbeenarelevantfeatureinthetaxonom- dried specimens from the following herbaria: B, ic history of the genus. Lamarck (1786)referredto BC, BCF, BM, BR, BRNM, COI-WILLK, E,FI,G, the heterocarpic and homocarpic species as ‘‘ar- GAZI,GH,GZU,HVR,IRAN,JACA,JE,K,LAU, niques’’ and ‘‘doronics,’’ respectively. Necker LE, LINN,LY, MA,MACB,MAF,MO,NY,RO,S, (1790)evenproposedthedifferentgenusAronicum SANT, UPS, W, WU, ZA, and theSa´nchez-Pedraja for the homocarpic species, and his classification personal herbarium. From other institutions, only had wide acceptance in the 19th century (de Can- photographs and photocopies of specimens, or ad- dolle, 1838; Koch, 1843; Hausmann, 1851; Rei- ditional information, were available: AV, B-W, chenbach, 1854; Schur, 1866; Ardoino, 1867; Si- BOLO, BP, C, CL, G-BOIS, G-DC, GE, GRM,LD, monkai, 1886), although some authors only NAP, P-HA, P-LA, LEB, PAL, and SZB. A list of recognized Aronicum as a section of Doronicum Volume 90, Number3 A´lvarez Ferna´ndez 321 2003 Doronicum(Asteraceae) (Ambrosi, 1854; Willkomm & Lange, 1870; Ny- erocarpywasclaimedagain(Gorschkova,1961).In man, 1879; Hoffmann, 1892; Beck, 1893). This the Flora of the U.S.S.R., the classification pro- classification persisted until Bentham and Hooker posed by Willkomm and Lange(1870)wasmerged (1873a) placed Aronicum as a synonym of Doroni- withthatofCavillier(1911)todistinguishtwosec- cum. Tausch (1828) also recognized these two tions in Doronicum (Gorschkova, 1961): section groupsbutreferredtheheterocarpicspeciestoPar- Aronicum (ser. Altaica, ser.Carpatica),andsection dalianches, not Aronicum. Pardalianches (ser. Austriaca, ser. Cardiophylla, Cassini (1817) established the genus Grammar- ser.Macrophylla,ser.Pardalianches,andser.Plan- thron with two species,G.biligulatumandG.scor- taginea). More recently, Edmondson (1978) fol- pioides,now subsumedwithinDoronicum(Jacquin, lowed Cavillier’s classification (1911) instead of 1773; Lamarck, 1786); de Candolle (1836) de- Gorschkova’s (1961), but described the monotypic scribed the monotypic genus Fullartonia (F. ka- subsection Isaurica (D. cacaliifolium Boiss. & maonensis), now D. kamaonense (DC.) A´lv. Fern. Heldr.) within section Doronicum, which corre- (A´lvarez Ferna´ndez, 2001). In 1838, de Candolle sponded to Cavillier’s section Doronicastrum proposed two sections within Doronicum: section (1911). Eudoronicum, including some species of Senecio, The phylogenetic analysis herein confirms what andthemonotypicsectionChromochaeta,withDo- a preliminary morphological study suggested: the ronicum linifolium(Wall.)DC.,nowalsoinSenecio morphologicalcharactersusedaretoolabiletopro- (Maguire, 1943). vide a sound classificationattheinfragenericlevel Webb in Webb and Berthelot (1846) treated the (A´lvarez Ferna´ndez et al., 2001). The molecular genusPericallisD.Don(inSweet,1833–1835:tab. data from nuclear ribosomal and chloroplast DNA 228) including five species as a section of Doron- suggestsomegroupsbutwithoutenoughsupportto icum (D. cruentum, D. echinatum, D. papyraceum, recommend formal taxonomic groupings. One ex- D.tussilaginis,andD.webbii),butitisnowtreated ception is a Mediterranean group of species (D. at its original rank (Nordenstam, 1978). plantagineum group), which receives good support Cavillier (1907, 1911) divided his study of the from both molecular and morphological data. To genus Doronicum, the first devoted to the study of avoidaddingtothealreadycomplextaxonomichis- the homocarpic species (1907), and the second to tory of the genus, a formal infrageneric treatment the heterocarpic ones (1911). However, he later is not proposed here, but is deferred against new concluded(1911)thatthischaracterwasnotuseful evidence. to delimitnaturalgroups.Cavillierproposedanew As already mentioned in the introduction, after classification (1911) that included 3 sections, 7 Cavillier’s revision and until the present work, a subsections,and34specieswithinDoronicum:sec- largenumberoftaxonomicactionsweretakenwith- tion Doronicastrum(subsect.Corsica,subsect.Aus- inDoronicum.Theseincludethedescriptionofnew triaca, subsect. Cardiophylla, subsect. Macrophyl- species (Sergievskaja, 1949; Edmondson, 1973, la, subsect. Pardalianchia, subsect. Plantaginea, 1978; Chen, 1998, among others) and one subsec- and subsect.Grandiflora),sectionSoulieastrum(D. tion (Edmondson, 1978), and a few lectotypifica- stenoglossum Maxim.), and section Hookerastrum tions (Chaco´n, 1987; Pe´rez et al., 1997; Jarvis & (D. hookeri C. B. Clarke ex Hook.). However, this Turland, 1998).Despitealloftheseactionsthege- sectionaltreatmentisnotsatisfactory,sincesection nus was still lacking nomenclaturalstability.Thus, Hookerastrum was described on the basisofaspe- during this study and immediately preceding this cies from another tribe (Nannoglottis hookeri, As- work, several nomenclatural and taxonomic clari- tereae) and the phylogenetic position(A´lvarezFer- fications were done (A´lvarez Ferna´ndez & Nieto na´ndez et al., 2001) of section Soulieastrum’s only Feliner, 1997, 1999, 2000; A´lvarez Ferna´ndez, taxon precludes recognition at the sectional level 2001). The lectotypification of 16 names ofDoron- without artificially splitting the bulk of the genus. icum in current use (A´lvarez Ferna´ndez & Nieto The subsections in Doronicum were defined (Cav- Feliner, 1999) gave the genus nomenclatural sta- illier,1911)mainly fromtheshapeofbasalleaves, bility leading up to this revision. In this work, ad- the size of leaves, and the presence of cypselae ditional lectotypification of seven names belonging dimorphism. Phylogenetic study of the genus (A´l- in Doronicum, although not in current use, is also varez Ferna´ndez et al., 2001) concluded thatthese presented to consolidate and clarify as far as pos- are not synapomorphic characters, and therefore sible the nomenclature of the genus. Despite the Cavillier’s classification does not recognize natural efforts made to locate type material for all the groups. names in current use, 4 out of 30 names (i.e., D. Even after Cavillier’s work the relevance of het- carpaticum,D.clusii,D.corsicum,andD.orientale) 322 Annals of the MissouriBotanicalGarden still required further investigations for lectotype where it can coexist with D. clusii. The alpineand designation.Becauseatpresentthesenamesclear- subalpine habitats of the central and northern half lyrepresentdifferentrecognizedtaxonomicentities, of the Iberian peninsula (except the Pyrenees) are they are cited herein as names in current use, al- occupiedbyD.carpetanum,underwhichfoursub- though their formal identity is not conclusiveuntil species are recognized. lectotypes are designated. Other alpine to subalpine speciesincentraland eastern Europe are Doronicum columnae and D. GEOGRAPHICALDISTRIBUTION carpaticum. The first is widely distributed from It- aly to Romania, and D. carpaticum is restricted to Half of the 26 recognized species of Doronicum the Carpathians. are distributed in Europe and North Africa. Seven TheremainingtwoEuropeanspeciesareendem- of the remainder are from southwesternAsia(Iran, ics, D. cataractarum in the Austrian Alps and D. Iraq,Caucasus,andTurkey),and6speciesaredis- corsicum in Corsica, and both occur in subalpine tributed in central Asia (Turkistan, Altay, Tibet, habitats. Yunnan, and the Himalayas). The genus Doronicum in southwestern Asia is With the exception ofDoronicumorientale,each represented by seven species. Only one, D. oblon- speciesisrestrictedtooneofthethreewell-delim- gifolium (from the Caucasus), is morphologically itedareas:Europe,southwesternAsia,centralAsia. quite different from the others. Three of them are Doronicum orientale is distributed in Europe and widelydistributed:D.macrophyllum(Caucasusand southwestern Asia,abundantlyintheeasternMed- northernTurkey),D.dolichotrichum(Caucasusand iterranean (Greece, western Turkey, southern Italy, south of the Caspian Sea), and D. maximum(east- and Lebanon–Syria), and scattered in central Eu- ern Turkey, and south of the Caspian Sea). Of the rope, where its proximity to inhabited places sug- three remaining species a limited number of spec- gestspossiblyhavingescapedfromgardens.Delim- imens are known, and this results in a scattered iting the natural areas of distribution is also distribution. difficult in the case oftwootherEuropeanspecies: AllofthecentralAsianspecies(D.altaicum,D. D. plantagineum and D. pardalianches. Both were briquetii, D. falconeri, D. gansuense, D. kama- used as ornamental plants in previous centuries onense, D. stenoglossum) overlap at least in one (Pena, 1571;Miller,1787)andnowareconsidered point of their distributions. alienplantsintheUnitedKingdom(Harron,1986; Clement & Foster, 1994). MORPHOLOGY The species discussed above (D. orientale, D. RHIZOMES plantagineum,and D.pardalianches)occurinsim- ilar mesic habitats from sea level up to subalpine All representatives of Doronicum are perennial regions, but not in high mountain habitats(theup- rhizomatous herbs. The shape and structure of the per tree-line). Doronicum hungaricum, occupying rhizome are constant within each species, but are similar habitats in Eastern Europe, can be consid- notexclusivetoanyone.Thesecharactersareuse- ered vicariant withD.plantagineuminthisregion. ful, sometimes indispensable, to discriminate be- Doronicum austriacum is widely distributedinEu- tween species.Therearefleshyorwoody(orsome- rope, most abundantly in the Austrian Alps, Mac- whatwoody)rhizomesinDoronicum.Thischaracter edonia, and Ukraine, always in subalpine regions, was described by Cavillier (1911: 199), who pro- andintheIberianpeninsulaitoccursonlyinafew vided histological diagrams, as ‘‘tube´reux’’ and localities in the eastern Pyrenees. ‘‘non tube´reux,’’ respectively. To distinguish be- The strictly alpine species of the genus in Eu- tween these, observations on fresh material arere- rope are represented by Doronicum grandiflorum, quired, although when pressed, fleshy rhizomes D.clusii,andD.glaciale.Thefirstofthesespecies flatteneasilywhilethewoodyonesretaintheirorig- is the most widely distributed of them. It is abun- inal more or less terete shape. When fresh, fleshy dant in the Alps, the Pyrenees, and in the Canta- rhizomes are succulent and brittle, while woody brian range(northernSpain).Inaddition,thereare ones are fibrous and tough. Fleshy rhizomes are two specimens from Corsica dated1878and1917, easily recognized in some European species (e.g., suggesting its extinction on this Mediterranean is- D. plantagineum, D. pardalianches, D. hungari- land, which has well known floristic affinities with cum), while woody rhizomes are well represented the Alps (Briquet, 1901). Doronicum clusii is pre- in Asian species (e.g., D. macrophyllum, D. maxi- sentintheAlpsandCarpathians,whileD.glaciale mum, D. stenoglossum). In a few cases, rhizomes isrestrictedtotheAlps(mainlytheAustrianAlps), are fleshy to somewhat woody and cannot be as- Volume 90, Number3 A´lvarez Ferna´ndez 323 2003 Doronicum(Asteraceae) signed to either type (e.g., D. grandiflorum, D. ca- fresh,andpaleyellowtobrownwhendry.Generally taractarum). stems are straight, but zigzag stems occur in some Within a species, rhizome internodes may have species (e.g., D. austriacum). The stems are often roughly constant length and width (e.g., D. altai- simple and end in a single capitulum. Whenbear- cum, Fig. 1G), or may vary in length and width, ing several capitula, the stem is branched only in resulting in stolon-like structures. Most species the upper part. Exceptions are seen in D. stenog- have the former condition, and only D. orientale lossum and D. kamaonense, which sometimeshave (Fig. 1A), D. plantagineum, D. pardalianches, and branches on the lower part of the stem. sometimesD.hungaricumhaveclearlyirregularin- Themainstemterminatesinacapitulum,which ternodes. matures first. Further capitula, if any, are on ter- Sometimes rhizome nodes have brown-tinted minal lateral branches, which for the most part scales remaining from the sheath of basal leaves overtop the main head. Each speciesgenerallyhas from previous years (e.g., D. carpetanum, Fig.1F). a characteristic number of capitula, e.g., one inD. The persistent remains of basal fibers from old falconeri, up to 5 in D. pardalianches, and more petioles occur, for example, in Doronicum oblon- than 5 (up to 20) in D. corsicum. gifolium, but they are frequently absent (e.g., D. austriacum, Fig. 1C). LEAVES Hyaline, shiny, and smooth trichomes aresome- timespresentontheyoungernodesofrhizomesand Leavesaresimpleandalternate.Leafcharacters also in the axils of basal leaves. Sometimes these havebeentraditionallyusedinthetaxonomyofthe trichomesarelong,abundant,entangled,andwhite genus(Cavillier,1911),buttheirusefulnessislim- to yellow, and they can cover a large part of the ited to the specific level. The shape and size of rhizome(e.g.,D.orientale).Suchrhizomeswerere- leaves are variable even within a single specimen ferred to as ‘‘e´riopode’’ by Cavillier (1911: 199) in for some species. For this reason, basal leaves contrast to ‘‘gymnopode’’ rhizomes,whichlackthis (those inserted on the rhizome nodes) and cauline indumentum (e.g., D. columnae). In many cases it leavesarenecessaryfordescriptivepurposes.Sim- is difficult to see trichomes on rhizomes, because ilarly, cauline leaves are distinguished by position they are short and scarce and can be coveredwith as lower, middle, and upper, i.e., inserted in the leafremains(e.g.,D.carpetanum,D.clusii,D.gla- basal,middle,andupperthirdofthestem,respec- ciale, and D. grandiflorum). tively. In some species, basal and lower cauline Buds are evident on some fleshy rhizomes (e.g., leaves are usually absent at flowering time. D. hungaricum, Fig. 1B). These stem buds can be Basalleavesarepetiolate,thepetiolebeingshort seen in plants two years or older, but these must and wide (e.g., D. briquetii), or much longer than be collected carefully. Sometimes the scales that the leaf blade (e.g., D. columnae). In species with cover young buds can also be observed. large basal leaves, sheaths are conspicuous (e.g., Two species each have unique rhizomes.Doron- D. macrophyllum, D. maximum, and D. dolichotri- icum cacaliifolium has moniliform rhizomes with chum). Acropetally, along the stem, the petiole uniform, spherical, swollen internodes, alternating gradually shortens, often leading to fiddle-shaped with nodal constrictions, sometimes covered by a leaves. The upper cauline leaves arereduced,ses- fibrous net. In the second type, seen in most D. sile, and ovate to bract-like. This leaf transitionis stenoglossum,themainstemisinsertedonaconvex marked in D. austriacum, D. carpetanum, D. ma- swollen woody surface. Sometimes, pieces of a woody organ perpendicular to the stem were also crophyllum, and D. pardalianches. collected. Although the whole structure has not Leaves may beorbicular,ovate,elliptic,andob- been seen, it is presumed to be a kind of distinct ovate, as well as fiddle-shaped or bract-like. The woody rhizome, but further study of the subterra- baseofleavescanbecordate(Fig.2A),subcordate, nean organ is needed. truncate, or attenuate (Fig. 3A, C, F, I). Leaf mar- Adventitious roots are present in Doronicum gins are generally entire to subentire, sometimes stenoglossum and sometimes in D. kamaonense a markedly dentate (e.g., D. cacaliifolium,D.colum- few centimeters above the subterranean woody or- nae, D. corsicum, and D. grandiflorum). gan, suggesting that the lowest vertical part of the Number and arrangement of cauline leaves de- stem was buried. termine to a large extent the architecture of the plant.Insomespeciesthenumberofcaulineleaves STEMS is low (2 to 4, e.g., D. orientale) and they are con- The stems in Doronicum are always erect, fistu- fined to the basal third of the stem. In other leafy lose, cylindric, and slightly ribbed, green when species (D. austriacum, D. corsicum, or D. altai- 324 Annals of the MissouriBotanicalGarden Figure1. RhizomesinDoronicum.—A.Doronicumorientale(drawnfromWilling3515,B).—B.Doronicumhun- garicum (drawn from Grundl s.n., G, as D. longifolium). —C. Doronicum austriacum (drawn fromStridetal.18585, B).—D.Doronicumcataractarum(drawnfromHo¨pflingers.n.,BM).—E.Doronicumcolumnae(drawnfromSladen9/ 4/452,BM).—F.Doronicumcarpetanumsubsp.carpetanum(drawnfromLucen˜o&Vargas208,MA).—G.Doronicum altaicum(drawnfromKrasnoborovetal.959,K). Volume 90, Number3 A´lvarez Ferna´ndez 325 2003 Doronicum(Asteraceae) Figure2. A,B.Doronicumpardalianches(drawnfromRivas-Godays.n.,MA).—A.Basalleaf.—B.Indumentum ofbasalleaf.C,D.Doronicumdolichotrichum(drawnfromDavis&Hedge29493,K).—C.Uppercaulineleaf.—D. Indumentum of upper cauline leaf. E, F. Doronicum briquetii (drawn from Rock 22380, E). —E. Upper caulineleaf. —F.Indumentumofuppercaulineleaf.G,H.Doronicumaltaicum(drawnfromKrasnoborovetal.959,K).—G.Basal leaf.—H.Marginofbasalleaf. 326 Annals of the MissouriBotanicalGarden Figure 3. A, B. Doronicum grandiflorum (drawn from Maquet 83/56, MA). —A. Basal leaf.—B.Indumentumof basal leaf. C–E. Doronicum glaciale (drawn from Steininger s.n., B). —C. Basal leaf. —D, E. Indumentum of basal leaf.F–H.Doronicumclusii(drawnfromCastroviejoetal.11615,MA).—F.Basalleaf.—G,H.Indumentumofbasal leaf.I,J.Doronicumoblongifolium(drawnfromAlburyetal.3176,K).—I.Basalleaf.—J.Marginofbasalleaf. Volume 90, Number3 A´lvarez Ferna´ndez 327 2003 Doronicum(Asteraceae) cum) leaves arearranged along thestem.Thelarg- CAPITULA est leaves are usually seen at the middle or basal AllDoronicumspecieshaveradiate,hemispheric parts of stems. to widely campanulate, homochromous capitula Leafvenationisagoodtaxonomiccharacter,eas- (Fig. 5A, E) with yellow or green-yellow corollas. ily observed in dry specimens and preferablyfrom Capitulum diameter ranges from 8 to 15 mm (e.g., basal leaves. For its description andcategorization D.cacaliifoliumandD.kamaonense)and7to8cm thetermsproposedbytheLeafArchitectureWork- (e.g.,D.falconeriandD.cataractarum).Therecep- ing Group (1999) are used. Most species have an tacle is convex to hemispheric, glabrous or pubes- actinodromousvenationforfirstveincategory(e.g., cent. In fruit, the base of the capitulum is some- D.grandiflorum, D.carpetanum,D.reticulatum)in times widely turbinate. which all secondary and tertiary veins are more or Rayflowersarefemalewithstrap-likeornarrow- less equally evident. Pinnate venation for the first lyelliptictoslightlyobovaterays,generallyending vein category occurs in central Asian species. In in three or two teeth, sometimesentire(e.g.,D.al- this type, the tertiary veins are not well marked, taicum). Disk flowers are hermaphrodite, actino- and both the secondary veins and the main vein morphic, and narrowly funnel-shaped. are equally prominentandthick(e.g.,D.altaicum, Phyllaries are herbaceous to slightly papery at D. gansuense, D. kamaonense, D. stenoglossum). the base in some species (e.g., D. austriacum)and The acrodromous type of venation for the firstvein arrangedin2or3rows,theouterbeingwiderthan category is restricted to a European group of spe- theinner.Inmostspeciesthephyllariesareclearly cies (D. hungaricum, D. orientale, D. plantagi- shorterthantherayflowers,buttheycanbealmost neum, and D. columnae). Intermediate cases be- equal or even longer than them (e.g., D. stenoglos- tween the latter and the actinodromous type occur sum, D. pardalianches). Phyllaries are ovate-trian- inD.columnae,D.carpaticum,andD.pardalianch- gular, ovate-elliptic, or ovate-lanceolate to linear. es, and between pinnate and actinodromous vena- The phyllary apex is usually acute, except in D. tion in D. clusii and D. glaciale. gansuense where it bears a sessile gland (Fig.4A– C). Phyllary margins are entire,exceptinD.haus- HABIT sknechtiiwheretheyareslightlyfimbriate.Agroup of species (D. orientale, D. plantagineum, D. hun- Four main habit classes can be distinguished: garicum, D. carpaticum, and D. columnae) have (1) An ‘‘orientale’’ type: solitary capitulum with a phyllaries with ciliate margins, bearing thin, stiff, scapose stem,sometimesbearingbract-likeleaves; acute, and equidistant trichomes (0.2–1.5 mm) a few cauline leaves (2 to 4) inserted in the basal (Fig. 5E–G). thirdofthestem.ItisdisplayedbysomeEuropean species (e.g., D. orientale, D. columnae, D. plan- FRUITS tagineum). (2) An ‘‘altaicum’’ type: generally a single capitu- Some species of the genus have dimorphic cyp- lum; a mostly leafy stem and a variable number of selae (heterocarpy), evident primarily by the ab- uniform leaves (41) along the stem length, or at sence of a pappus in ray flowers. Cypselaewithout leastin itslowerhalf.Thisispresentinsomecen- pappuses are also generally larger than those with tral Asian as well as European species (e.g., D. a pappus in the same specimen (Fig. 6A, B). Het- altaicum, D. falconeri, D. grandiflorum). erocarpyoccursinothergeneraofAsteraceaesuch (3) A ‘‘macrophyllum’’ type: several capitula;stem as Senecio, Crepis, Erigeron, Leontodon, and Het- branched in the upper third; large cauline leaves erotheca, as well as in other families, Caryophyl- (3 to 5) mainly in the lower half of the stem, and laceae, Apiaceae, Poaceae (Zohary, 1950). These bract-like leaves on the upper stem. This is re- morphologic differences serve different functions. stricted to the southwestern Asian species. In Heterotheca latifolia, in which the type of di- (4) A ‘‘corsicum’’ type: several capitula; a variable morphism is similar to Doronicum, Venable and number of6 uniform leaves(51)6evenlyinsert- Levin (1985) suggested that the pappose cypselae ed along the stem. This is characteristic of D. cor- are dispersed away by wind, while epappose cyp- sicum and D. austriacum. selae fall near the mother plant. This double dis- In some species the habit does not correspond persalstrategyimpliesthepotentialtocolonizedif- to these patterns (e.g.,D. kamaonenseandD.sten- ferent habitats (Venable & Levin, 1985; Tanowitz oglossum, which are sometimes branched from the et al., 1987; Imbert et al., 1996). Although plants base), and sometimes intermediate patterns occur with incompletely developed pappuses on ray flo- (e.g., D. pardalianches, D. cacaliifolium). retsarefoundinsomespecies(e.g.,D.carpaticum, 328 Annals of the MissouriBotanicalGarden Figure 4. A–D. Doronicum gansuense (drawn from Rock 12192, E, as D. thibetanum). —A. Capitulum. —B. Phyllary.—C.Apexofphyllary.—D.Rayflower.E–H.Doronicumstenoglossum(drawnfromRock12941,GH).—E. Capitulum.—F.Phyllary.—G.Indumentumofthebaseofphyllary.—H.Rayflower.