A Taxonomic Revision of the Complex (Paeoniaceae) Author(s): Hong De-Yuan and Pan Kai-Yu Source: Annals of the Missouri Botanical Garden, Vol. 91, No. 1 (Apr., 2004), pp. 87-98 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/3298571 . Accessed: 14/02/2015 09:22

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This content downloaded from 128.122.253.212 on Sat, 14 Feb 2015 09:22:03 AM All use subject to JSTOR Terms and Conditions A TAXONOMICREVISION OF Hong De-Yuan2and Pan Kai-YuS THE PAEONIAANOMALA COMPLEX(PAEONIACEAE)

ABSTRACT Basedon field observation,examination of over350 sheets of exsiccatae,and investigationof all the relevanttypes, a taxonomicrevision of the Paeonia anomalacomplex is presented.Typifications are clarifiedfor P. anomala,P. intermedia,P. hybrida,P. sinjiangensis,P. altaica, and P. veitchii.The identityof threewell-known taxa, P. anomala, P. hybrida,and P. intermedia,is also clarified.Two species, P. anomalaand P. intermedia,are recognized,with the circumscriptionof P. anomalaemended to includeP. veitchiias its subspecies.Paeonia anomala includes two sub- species: subspeciesanomala and subspeciesveitchii with the formerdistributed in CentralAsia, Siberia,and the northeasternEuropean part of ,while the latteroccurs in Chinasoutheast of the GobiDesert. Paeonia intermedia is distributedin CentralAsia, but withan isolatedlocality in Georgia.Paeonifl sinjiangensis is treatedas a synonym of P anomala,arld P. hybridaPallas is treatedas a synonymof P. fenuifolia L. The lectotypesof Paeonialaciniata, Paeoniaberesowskii, and Paeoniaintermedia are designatedhere in the presentpaper. Keyuvords: Paeonia, Paeonia anomala complex, Paeonia hybrida, Paeonia intermedia, Paeoniaceae.

The Paeoniaanomala L. compIex (Paeoniac eae) needed a comprehenxivetaxonomic revixion and comprisesa groupof her})aceous peoniesin Central reviewof the nomenclature.For thix purpose,the Asia, Siberia,an(l adjacentnl)rthe,steln Euroy)ean first authorexamined a11 availatele specimens of region,.They are lhara(*teri%ellI)yleavex t)iternate this grouXr)in the herI)ariaat BM, HNWP,K, I,E, with l.eaflets(le(urrent at the t)ase, leaflets finely NWTC,PF. SHI, SHMU,TBI, XJBI, XJNU,and segmente(l,with segmentsof a lowet (the be,st(le- XJU(see A^)penllixl). In aelditio, the ty^es of P velope(l) rangingfrom 7() to 1()()in nullll)erantl anorrl(ll(lXP 1Iybrid(l,I' in.l.erme(li(l,all(3 the othel 4-432mnl in wi(lth.an(l t)y t)ristle,

' The authorsare gratefulto the NationalGeographic Society for financialsupport (Grant 6408-99), whichallowed us to examineherbarium specimens at the KomarovInstitute of Botany,the RussianAcademy of Sciences, and to conductfieldwork in Georgia.The project is also supportedby the NationalNatural Science lSoundationof China(NSFX Grant30130030). We are gratefulto the curatorsof the followingherbaria: BM, HNWP,K, LE, NWTC,PE, SHI, SHMU,TBI, XJBI, XJNU, and XJU.We sincerely thank Zhou Shi-Liang for his assistancewith fieldwork, and Li Qiao- Lingand MaLi-Ming for theirhelp in preparationof lhe manuscript. 2 Laboratoryof Systematicand Evolutionary13Otany, Institute of 13Otany,The ChineseAcademy of Sciences,Xiang- shan, 13eijing100093, [email protected]. ANN.MISSOURI BOT. GARD. 91: 87-98. 2()04.

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laciniata as a synonym of P. anomala. Meyer riety anomalaand varietyintermedia (Pan, 1979). (1830) describedthe fourthspecies, P. interrnedia, Also fromXinjiang, Dai and Ying(1990) described fromthe .Ledebour (1842) citedP. anothernew species, P. altaica, which was stated anornala,P. hybrida,and P. interrnedia,treating P. to have flowerslarger than P. sinjiangensisand one Iaciniataas a synonymof P. anornala.Trautvetter or two underdevelopedflower buds in additionto (1860)recognized only one species, reducingP. hy- the terminalblooming flower. brida as a varietyof P. anornala,and treatingP. Interestingly,Schmitt (1999) failed to recognize interrnediaas a form of variety hybrida.Lynch the distinct differencesbetween Paeonia anornala (1890) cited P. hybridaand P. anornala,and stated and P. interrnediain the root and calyx, and thus that the formerwas native to the Caucasus.Huth treatedthem as a single species, but still recog- (1892) adoptedthe widestspecies conceptof P. an- nizedP. sinjiangensisand P. altaica as distinctspe- omala, which includes four varieties:var. typica, cies. var.hybrida (= P. interrnedia),var. nudicarpa, and Fromthe aboveit is clear thatPaeonia hybrida, var. ernodi(= P. ernodi,confined to the western P. interrnedia,and P. anornalahave been variously Himalayas).A furthertreatment by Krylov(1901) treated.With this paperwe hope to clarifythe bi- divided the complexinto two species, P. anomala ologicalattributes and variationsof the P. anomala and P. hybrida,and reducedP. interrnediaas a va- complex. riety of the latter,P. hybridavar. intermedia (C. A. Meyer)Krylov, in additionto the typicalvariety. In OBSERVATIONSANDDISCUSSION the area of WestTienshan (Kirghizia and Xinjiang, China),Trautvetter (1904) enumeratedonly P. an- To answerhow many species werereally present ornala, treating F! interrnediaas its subspecies. in Xinjiangand whatbiological features they have, Schipezinsky (1921) basically followed Krylov the first authorand coworkers(Hong et al., 1994) (1901), recognizingtwo species, P. anornala(with made an ex)edition to the Tienshan and Altai twovarieties, var. anomala and var.nudicarpa) and Mountains,c onductingfield observationsand pop- P. hybrida(with two varieties var.hybrida and var. ulationsamplings of Paeonia in 1993. Sevenpop- intermedia),and the same treatmentwas adopted ulations in total were observed covering those for the Flora of the USSR (Schipezinsky,1937). characterstates previouslydesc ril)ed for the P. an- These two Paeonia species were groupedby him omalacomplex (Table 1). Wefoun(l there were only as series4, DentataeKom. (Schipezinsky, 1937: 33, twospecies in Xinjiangthat differecl distinctly from "leaflobes incised or with dentatemargin"). Roots each otherin the rootand calyx. One species had of both species weredescribed as tuberous.In con- the rootsbasipetally attenuate (( arrot-shaped)and trast, Stern (1946) recognizedonly one species in mostlycaudate (Fig. 2), y)referringrelatively his monographof Paeonia, P. anomala, in which moisthabitats in woods.This was treatedas P. sin- twovarieties were recognized: variety anomala with jiangensis.The otherhad rootstut)erous or fusiform glabrouscarpels, and varietyintermedia with to- and sepals mostly(at least 2) roundedbut not cau- mentosecarpels. He did not mentionroots of this date at apex, growingon sunny shrubbyor grassy group.Stern (1946: 113) consicleredP. hybridaPal- slopes, or in sparsewoods. This latterwas recog- las as an ambiguousname: "It is doubtfulwhat nized as P. arlomala.We also foundthe presence species Pallas intendedby this name and, as it is or absence of indumentumon the carpelsin this not possible to discoverwhat he meant,this name species groupto be a polymorphism,i.e., individ- of P. hybridahas been omitted."However, in the uals with carpels either glabrousor fromsparsely Flora of ,Gamaulova (1961), following to denselypubescent could be foundwithin a single Krylov(1901) and Schipezinsky(1921, 1937), de- population(Table 1). The indumentumon carpels scribedtwo species, and still recognizedP. hybrida was demonstratedas quitevariable, although it was as a valid name. consideredby the previousauthors as taxonomi- While workingon Paeoniafor the FloraReipub- cally valuable.Therefore, any taxonomicdivision licae PopularisSinicae, Pan (1979) founda speci- based on this character,such as that betweenP. men fromXinjiang whose roots were basipetally at- anomalaand P. anomalavar. nudicarpa (by Huth, tenuate. According to Schipezinsky(1937) and 1892) or P. anomala and P. anomala var. inter- Gamaulova(1961), Paeoniaanomala and P. hybri- media (by Stern, 1946), is artificial.Also not - da havetuberous or fusiformroots, so the plantwas served by the previousauthors was the clear dif- describedby her as a new species, P. sinjiangensis ferentiationin rootand calyx in this group,which K. Y. Pan;the plantswith tuberous or fusiform roots are closely correlatedwith each other(Fig. 2). were treatedas P. anomalawith two varieties:va- Wealso foundthat in the formwith carrot-shaped

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Volume 91, Number 1 Hong & Pan 89 2004 Paeonia anomala Complex

roots,the variedgreatly in length,from 3.5 ;) J to 6.5 cm, and in the Altai Mountainssome indi- : .

* S . F cn vidualsof this formpossessed one or twoadditional X Ut - / cn X e ct o N e underdevelopedflower buds similar to Paeonia = Ct . F . F . F . F 5

- / reitchii. This was again a polymorphism,and not 5 X o o e o o C * consideredby us to be a stable character,and X . thereforeP. altaica was reducedto a synonymof P. d .: ct sinjiangensis(Hong et al., 1994).

. cn 5 o ce ut : N - / After examinationof about 200 sheets of her- 5 5 e ce : . bfi s s . ctF . F . F . F bariumspecimens in the KomarovInstitute of Bot- < C,: 5i:i o o o N N any (LE) in 1999, the first authorfound that two z X E X species couldalso be distinguishedin this complex: . O one with the rootscarrot-shaped and sepals mostly

*: O caudate,while the otherwith the rootstuberous or 5 5 o fusiform,and sepals mostlynon-caudate. These bi- Ct n o c;; 5 cn n: bfi S : ologicalfeatures match those on plantsobserved in X * X ct = .> S C; n ct C X Xinjiang (Hong et al., 1994). Gamaulova(1961, ; ; bfi bfi bfi bfi O t s tab. 2) describedroots of Paeonia anomalaas fu- ;;> * X t .: siform-thickened,and those of P. hybridaas tuber-

C53 Ct ous-thickened.However, roots of P. intermedia(=

Ct C.) F z- z- z- z- ce N c e P. hybrida sensu Schipezinskyand Gamaulova) cn ut ut cn vary in shape from tuberousconsistently to fusi- form,and this variationcould be foundwithin pop- cn ulationsor even on the same individual,whereas Ct g J . oo; ; ; o o o o * _r f : n e ,. in P. anomala neitherfusiform-thickening nor tu- berous-thickeningwere found. k ; k .,, c,, .,. ... Contradictionsexist betweenour naturalobser- X . $ vations(Hong et al. 1994) and earlierdescriptions by Schipczinsky (1937) and Gamaulova(1961). c; X = Stern's(1946) treatmentof this complexas one spe- t . 1 = n * cies withtwo varieties (P. anomalaand P. anomala o e ;:: n cn x, - X var.intermedia) is also at variance.For a clear tax- > = 5 U :> ;> > ct onomicrevision and correctnomenclature, one re-

o e X AD * mainingissue oughtto be resolved:the identityof cn t ) X s X, o X ct i o < Paeonia anomala, P. hybrida,and P. intermedia. '= N The type specimenof P. anomala,with two visible sepals both caudate and relativelythin leaves, is cn = .. perfectly consistent with the form with carrot-

. shaped roots. Accordingto the type specimen at 5 X e ct o LINN[Siberia, no. 692. 3], field observation,and .; n ut examinationof over 350 sheets of exsiccatae, P. = cn; sinjiangensisand P. altaica should be treated as n cn F} ct = X synonymsof P. anomala;the descriptionsof roots of P. anomalaby Schipezinsky(1937) and Gamau- ; ct lova (1961) are erroneous.The type of P. hybrida

ct: F cn . (at BM),on which Pallas's(1789) tab. 86 was ap-

O O t parentlybased, has leaf segments3-4 mmin width, * =: and at least morethan 100 in number,and is thus co within the range of variationin P. tenuifolia L. o C'1 (Hong& Zhou,2003). Therefore,P. hybridaPallas = cn o o t E- X e . shouldbe treatedas a synonymof P. tenuifolia,as ct o , - X . cn b Anderson(1818) suggestednearly 200 years ago.

o X X cn The type specimenof P. intermediaat K possesses o C; cn t. S. tuberousroots, mostly non-caudate sepals, and rel- ct; W. a c; X ativelythick and narrowleaf segments.The

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thathave tuberous or fusiformroots, are distributed Paeonialaciniata Pallas, F1.Ross. 182):93, tab. 85, sub widely fromthe Altai to Tadzhikistanand Uzbeki- F! sibirica,1789. TYPE:the illustration(tab.) cited in the protologue(lect{>types designated here!3. stan, and were namedP. hybridaby Schipezinsky Paeoniasibirica Palla ,, F1. Ross. 1(2):tab 85. 1789. (1921 1937) and Gamaulova(1961) correspondto this type specimenarery well. Pallas's(1789) illustration (tab.) 85 has only one Stern's(1946) statementconcerning the nameof name, P. sibirica, and apparentlyhe used this il- Paeonia hybridais not correct,confusing F! inter- lustrationalso for E laciniata because one sees media with P. anomala.Pan's (1979) treatmentof "Paeon-rcllaciniata Tab.LXXXV" on p. 93. Paeorl- the E anomalacomplex (a new species, P. sinjiarl- ia laciniata was describedin detail by Pallas, but gensis,and P. anomalawith var. intermedia) is also no descriptionof P. sibirica was given by him. unjustifiable.Although Hong and his coworkers Perennials;tap roots > 50 cm long, carrot- (Honget al., 1994) properlydescribed two species shaped, basipetallyattenuate, up to 2 cm diam., and clearly statedtheir morphologicaland ecolog- lateralroots slender, neither tuberous nor fusiform. ical differencesfor this grouT3,the nomenclature Leaves biternate;leaflets finely segmented;lower they used is misapplied. leaves with segments70 to 100 in number,8-32 The Paeonia anomala complex comprisestwo mm in width.Flowers solitary or 2 to 4 on a stem, species: P. anomala and P. intermed7^a.Paeonia often only terminalone fully developedand bloom- ing; sepals 3 to 5, mostlycaudate at apex,rarely 1 emodi,which was treatedas a varietyOr E anomala by Huth (1892), remainsa distinet species, with or very occasionally2 non-caudate;carpels 2 to 5, leaves ternate,without bristles on the upper sur- fromglabrous to densely tomentose. face, leaflets much less segmented,segments no The species prefers relatively moist habitats, morethan 40 in numberand 1.5-4 cm wide, car- growingin forests,on the edges of forests,or rarely pels usually two less frequentlyone, very rarely in bushes. Il is found at altitudesfrom 1100 to threeeand flcowerswhite. Lynch 387() m. was described from Province, China This species is widely distribllte(lfrom central (Lynch 1890), and was said (Stern, 1946; Pan, Chinato the Kola Peninsulaof Russia via Siberia 1979) to differfrom the R anomalagroup in having and CentralAsia. The s;})e(ies is of two allopatric severalflowers instead of a single floweron a stem. subr>l)ecies,with the tyl)icalsubspecies distributed Accordingto our observation,one or two under- northwestof the Cobi DeKsert,while sul)species veit- developedflower buds sometimesalso exist in ad- chii is in Chinasoutheast of the Goti Desert(Fig. dition to one normaland terminalflower in E an- 3). omalaS while E veitchii sometimes has flowers solitary(P. veitchiivar. ztnifora), although it has 3 1 a. Paeonia anomala sut)sp. anolnala or 4 flowersor 1 to 3 underdevelopedflower buds Paeollia sinjiangensisK. Y. Pan, F1. Reilsllbl.Popularis in additionto the terminalflower on a stem. Thus, Sin. 27: 603, fig. 12. 1(379.TYPE: (jllina. Xinjiang: P. veitchiiwas reducedto a subspecies of E an- HalgaheCounty, Lari.X- foreests, ]973, Y.R. Ling1141 omala (EConget al., 2001). (holotype,PE!). Paeorliaaltaica K. M. Dai & l^.H. Ying, lXull.Bot. Res. KEYTO THE PAS(3NIA ANOMAS COMPLEX (Harlgin)10(4): 33, fig 1. 1990. rtYI'Id:China. Xin- la. Roots( arrot-shaped,never tulgerous or fusiform, jiang:Halgahe Countye Wuzlitie 1.5.5() tn, in forests? sepals all or mostlycaudate, less frequentlyone 10 June 1980, E H. Yirlg1007 (holl)type,SHMU; isotype,PE!). or veryrarely two non-caudate 1. E anomala 2a. Flowerssolitary, rarely 1 or 2 underdevel- The typical subspecies is usually foundin de- oped flowerlguds present in additionto ter- minal floweron a stem ------ciduousor coniferforests, in valleys?less frequent- la. P. anomalasulgsp. anomala ly in meadovTsnat altitudesfrom 11()() to 2200 m. 2b. Flowersusually 2 to 4 on a stem or 1 tc)3 Apparentlyit prefersrelatively moist habitats. This underdevelopedflower lguds present in ad- subspecies is of very wide distribution:northeast ditionto terminalflower, rarely flowers sol- itary on a stem ------Kazakhstan,northern Mvngolian from the Altaiand ------11z. P anomalasubsp. veitchii Baikalto the KolaPeninsula in Russia,and in the lb. Roots fusiformor tuberous;inner 2 to 4 sepals Altai and adjacentregions in XinjiangnChina (Fig. roundedbut non-caudateat apex 2. E intermedia 3). TAXONOM1CTREATMENT Additionalspecimens examined. CHINA.Xinjiang: Altay,L. R. Xu s.n. (PE);Mt. Halamaryi,D. K Hong et 1. Paeonia anomala L. Mant. 2: 247. 1771 al. PopulationIVo. 3 (PE),0128 (PE),Xiaoclong Gou val- TYPE: SiberiaSno. 692.3 (holotypenLINNt). ley, T H. Ying 1001 (PE, SHMU).Burqin: Kern, IV. R. Figure1A. Cui86624 (XJNU).Emin: Shiyue Township, BTuerkeheXer,

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XE+bt:l; mMS

Figure1. Paeoniaholotype specimens: A. F: anonxala(LINN). B. P. hybridaPallas (BM). C. P. intermediaC. A. Meyer(K): the insertat upperright is the enlargement of the label at lowerright.

cO

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2cm A

2 m B

Figure2. Diagnosticfeatures of Paeoniaanomala (A, B) and P. intermedia(C, D). Tuberousor fusiformroots are correlatedwith non-caudatesepals, while carrot-shapedroots correlated with usually (audate sepals. right bank of Telikete River, XinjiangBranch Acad. Sinica (LE); Mt. Narymsky Range, Ak-tui River valley, D. Ya- 012 (XJBI). Fuhai: Ahebai, glades, s. coll. 3867 (XJBI); korlev 69 (LE); Mt. Saur, Chagan-oboRiver valley, E Rez- Daqiao Forest Farm, T H. Ying1011 (PE, SHMU). Fuyun: nichenko s.n. (LE); Mt. Saurkie, Maly Dgewen River val- C. 1EChang 11356 (XJBI). Habahe: Baihaba, Z. M. Mao ley, B. Keller s.n. (LE). MONGOLIA.Ulan Bator: 10 mi. 10413 (XJBI); Tielieke, Zamanashi Valley, C. t Chang E of Ulan Bator, E Morgan 23 (K). Gloucester: Mongo- et al. 10255 (XJBI); Tiekeliti, 1: H. Ying 1010 (PE, lia-Chinese TurkestanExped. M. P. Price 293 (K). North SHMU); Wuzlity, E H. Ying1022 (PE, SHMU). Jeminay: Mongolia: Mt. Saldzharsky Pass, E. Klements 11a (LE). Xingfu Township, Sanxie Valley, Integ. Exped. 7305 RUSSIA. Altai Region: Altai: between Charga & Sha- (XJBI). Qinghe: Daqinghe, Mudeng Valley, Integr.Exped. bolina, mtn. slopes, E. K. Klements4d (1 E); Chuja road, 11829 (XJBI). Tacheng: Mt. Ba'erleike, X. Y. Li 870147 Saadak-lary, Vereshagin296 (LE). West Altai: Mt. List- (SHI). Toli: Jiayi'er Gold Mine, N. R. Cui 091 (XJNU)- vennitchny, near Nizhne-Uimonskoye, KS.Kolomoitseva Mt. Zayier, N slopes, Integr.Exped. (Biology) 630 (XJBI). 236 (LE). Buryato-Mongolian Autollomous State: Wenquan: A'erxiaoto Gou, E M. Zhang 90-346 (XJNU). Baikal, Tunkam, Turezaninows.n. (LE); Zahaikalje, upper Yumin: Ku'erzhai, Liu & Shen8308 (XJBI). KAZAKHS- Toreika River, 130 km W of Troitskosavsk,R Mihno s.n. TAN. Semipalatinsk Province: Altai, near Altaiskaya (LE). Eubinsk Region: West Sayan, Mt. Alan Range, Go- Station, Mt. Narymsky region, gorge slopes, Ladygin92 lets Taskalyk, I. Krasnoborov6765 (LE); near Buiba Sta- (LE). Southern Altai: Mt. Ivanovsky Range, near Gorno- tion, by river, Krasnoborov& Sannikova s.rl. (LE). Tuva: Ulbinki, I. Rohderis.n. (LE). Ustkamenogorskregion: Mt. Tandinskydistr., East Mt. TannudlaRange, near Shurmak, Altai, Kunduzda River valley, E & A. Reznichenko360 V Hanminchun 515 (LE); Ulug-Hemsky distr., Uiuksky (LE); Mt. Kalbinskie, Saryshbay River valley, E Rezni- Mtn. Range, Bajan-Kol River valley, Orto-Hemtributary, chenko49 (LE); Mt. Kalbinskie, Ulansky, near Asybulak, Lomonosova& Ivanova 89 (LE). Irkutsk Region: Irkutsk by Tainta River, P. Yurchenkos.n. (LE); Mt. Narymsky, Prov.: Lena River valley, upper Gulm River, I. Kuznetsov upper Urkar River, E Reznichenko58 (LE);near Ulbinsky 114 (LE). Balagansky distr.: near Bazheevskoye, N. Mal- gorge, E Iljins.n. (LE); Urunhaika River mouth, S banks tsev 79 (LE). Nizhneudinsky distr.: by Angara River, op- of Marka-KulLake, A. Sedelnikovs.n. (LE). Zaisan region: posite Yandov, S. Ganeshin s.n. (LE); Mt. Hariba-bim Mt. Altai, Ulkunchumek River valley, upper KaldzhirRiv- Range, Angara River valley, Karpovskoye, A. Kororkin er, E Reznichenko106 (LE); MarkakulLake, B. Kellers.n. 290 (LE); by Oka River, A. Krishtofowichs.n. (LE); near

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Vidimka,Vidim River, S. Kucherovskaya282 (LE).Tunkin Tatien-lu(Kangding), 8-10,000 ft., E. H. Wil- distr.:Mt. Sayany,near NylovaPustyn, E Komarovs.n. son (forJames Veitch& Sons)3034 (holotype, (LE);Nizhnaya Tunguska valley, near Nizhnaya Korelina, S. Kokulin254 (LE).Tunkinsky distr.: Kuren Arshan, E K!, photo,PE!). Smirnovs.n. (LE).Verholensky distr.: Biriulka, near Yu- shina, E Alexandrov179 (LE);Harat, G. Dolenko1 08 Paeonia beresowskiiKomarov, Bot. Mater.Gerb. Glavn. (LE);Lena River & Kirengavalley, Toptykan, M. Tomin Bot. SadaRSFSR 2: 5. 15 Jan. 1921. Paeoniaveit- 67 (LE);Tutura, P. Alexandrov219 (LE).Kola Penin- chii var. beresowskii(Komarov) Schipezinsky, Bot. sula: Ponoi River, left bank, 8 km from its mouth,E. Mater.Gerb. Glavn. Bot. Sada RSFSR 2: 46. 26 Mar. Chernov3 (LE);Ponoi River,near its mouth,N. Orlova 1921. TYPE: "ChinaOccid, Szechuan(Sichuan), 240 (LE).Krasnoyarsk Region: Belsk distr.:near A1- Sun-pan-tin(Songpan), Guichua," 9 June 1894, M. shat,I. Kuznetsov4661 (LE);Chadobets, G. Borowikow s.n. Beresowskiis.n. (lectotype,designated here, LE not (LE), Rybnoe,Dranitsyn & Kochubeis. n. (LE). Enisei seen). Prov.,Achinsk distr.: near Ingol Lake,A. Suharevas.n. Paeonia veitchiivar. woodwardii (Stapf ex Cox)Stern, J. (LE),near Maloye Lake, M. Ermolajevas.n. (LE),Bolshe- Roy.Hort. Soc. 68: 130. 1943. Paeoniawoodwardii Uluiskaya,near Bobrovki,I. Kuznetsov53 (LE),Chuksha Stapf ex Cox, P1. Introd.Farrer 43. 1930. TYPE: River Valley,I. Tolmatschew247 (LE),Katanga, G. Bo- China.: Zone (Chuoni),R. Farrer67 (holo- rovikovs.n. (LE);Vyezzhi Log, A. Tugarinovs.n. (LE). type, E not seen). Kanskdistr.: Gutar River valley,by Kamenkatributary, Paeoniaveitchii var. Ieiocarpa W. T. Wang& S. H. Wang, 1E Troitskys.n. (LE),near Kansk,A. Shliahtins.n. (LE), F1. Reipubl.Popularis Sin. 27: 603. 1979. TYPE: RybnayaRiver valley, Perovskoye, I. Kuznetsov193 (LE); China. Sichuan: Jinchuan, Kasa Township,Yin- near Nazimovskoye,Z. Evseeva4406 (LE).Krasnoyarsk changgou,forests by stream,2700 m, 26 Apr.1958, Prov.:Chunia River, right bank opposite mouth of Mutorai X. Li 77248 (holotype,PE!). River,A. Rubins.n. (LE),Hakassia, Shirinsky distr. Bol- Paeoniaveitchii var. uniJYora K. Y. Pan, F1.Reipubl. Po- shayaIiusa Rivervalley, NW slopes, Polozhii& Kanda- pularisSin. 27: 603. 1979. TYPE:China. Sichuan: sova s.n. (LE);near Krasnoyarsk,by MohovayaRiver, E Gaze,Xiongjiling, 3600 m, mtn.summit, bushes, 18 Verhovskaya& M. Mishin104 (LE);Minusinsk, Bolshoye June1974, -XizangExped. Veget. Group 034 KysylkulLake, K. Golubevaet al. s.n. (LE);Nizhnaya Tun- (holotype,PE!). guskaRiver valley, near mouth of HurkakitRiver, Rubin & Maskils.n. (LE);between Salba & Grigorievka,E Kry- This subspecies, thoughdistantly isolated from lov s.n. (LE);Shushensky distr. near Tanzyben, by Black Paeoniaanomala subsp. anomalaby the huge de- TanzybenRiver, Kuminova & Alexeevas.n. (LE);near So- serts of the Gobi still very much resemblesthe rokinoStation, N slopes.Chere/lnirl s.n. (I E). Novosiber- latter. The only remarkabledifference between ia Region: Altai:Elikmanar. upper Karakol River, V Ver- eshagin 345 (I E). KuznetskyAlatau: Kondoma River them is the numberof flowerson a stem. Paeonia valley,Kazany Mine, Mt. Kyon, B. Klo/lotovs.n. (LE);up- anomala subsp. veitchiiusually possesses 2 to 4 per Sary-ChumyshRiver, near Ulus Munai,A. Vyslrins.n. bloomingflowers in additionto up to 2 underde- (LE);Tom River valley, Mt. Tohpan-taiga, B. Klopotovs.rl. velopedflower buds. Veryrarely only the terminal (LE).Mariinsky Prov.: Biisk, Black Anuj,Talitskiebelky flowerblooms in additionto up to 3 flowerbuds. to Kazanda,Pobedimova 697 (IE)- Biisk, Lebed River valley,by ChujaRiver, B. Klopotov180 (LE);Biisk, be- Paeoniaanomala subsp. anomala possesses only a tweenTulatinskaya & Sentelek,N. Kuznetsol1991 (LE); single terminalblooming flower, without or infre- Kainsk,Verhnyaya Nazarova, B. Klopotov113 (LE);Na- quentlywith 1 to 3 underdevelopedflower buds. rym,Ixa River,S. Geninas.n. (LE);Sofronova, E Krylov The subspecies is widely distributedin China: s.n. (LE);Tomsk distr.: Chulym & Chet Rivervalley, P. Sokolov13a (LE). MoshkovskyProv. (Distr.): Uskorniha southeasternand centralGansu, southern , River valley, Kuvajev& Sabitovs.n. (LE).Toguchinsky eastern Qinghai, the Qinling Range of , distr., near Mirny,Lashinsky & Ronginskaya973 (LE). ,western Sichuan, and the easternextreme Omsk Region: TobolskProvince: Berezov distr., Lopsia of Xizang (Tibet)(Fig. 3). Like subspeciesanom- River,tributary of SevernayaSosva, s. coll. s.n. (LE);To- ala7 Paeonia anomalasubsp. veitchiiprefers rela- bolsk distr.,Landinka River valley,E. Visloah73 (LE). : ArcticUral: Lyapin tributary, upper Manja River, tively moisthabitats, growing in forests,grasses on E Sochava100 (LE);Severnaya River valley, Lyapin the edges of forests,bushes, or subalpineand al- tributary,upper Khulga River, B. Gorodkov512 (LE).Mar- pine meadowswith shrubs,at altitudesfrom 1800 iinskydistr., Beket Rivervalley, Kolsonskoye, N. Kuznet- to 3870 m. sov 199 (LE).Tobolsk Province: Berezovsky distr., Bere- zov, Pohle & Rozhdestrenskys.n. (LE); Tarskydistr., Phenology. The subspecies flowers from late Rybinsk,near Pustynnoye, E Varentsovs.n. (LE).Yakutsk April to earlyJune, and fruitsin Augustand Sep- Autonomous State: Yakutia,Suntarsky distr., upper Vil- tember. yuy River, Tuoikhaia,by ChonaRiver, I. Kildushewsky Chromosomes.2n = 10 (Honget al., 1988). 150/22 (LE). Additionalspecimens examined. CHINA.Gansu: Di- lb. Paeonia anomala subsp.veitchii (Lynch)D. ngxi,s. coll. s.n. (CPB).Hezheng: Xinzuang, Gansu Herbs Groups.n. (NWTC).Hezuo: Mt. Dalinke, Q. R. Wang7034 Y. Hong & K. Y. Pan, Novon 11: 317. 2001. (NWTC).Jiangeha: S. Gansu GrasslandExped. 681 Paeonia veitchii Lynch,Gard. Chron. ser. 3, (NWTC).Kangle: Mt. Taizi, G. L. Zhu 80006 (NWTC), 46: 2 tab. 1. 1909. TYPE:China. W. Sichuan: Mt. Lianhua,M. S. Yan1843 (NWTC).Kangxian: s. loco,

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60°

130° fOo

4Oo

120°

30°

60° 70° 80° goo 100° l lOo Figure 3. Distributionmap of the Paeonia anomala species comyalex.BIack circles:Paeoni(z (lraomala subsp. anomala;white circles: E anomalasubsp. veitchii; stars: E intermedia.Each site in the mapis SUp[)OltP(3 by a single or severalvouchers cited in the text. s. coll. s.n. (N%TTC).Lanzhou: lUt. Xinlong,s. coll. 84 e1l. 378 (HI9WP),Baoku, Z. H. Zh(tlg et al. 429] (NGTTC);Mt. Tiane3u,s. coll. s.n. (NWTC).Linxia: near (HNWP).Huangyuan: Mt. Banjie, E C. Ts0e,,7g8842 (PE). Senanpa,by river,7: K Fu 834 .PE). Longdie:Dacaopo, Huangzhong:Sanhei Forest Farm, KS.X. Wang 1065 H. J. Zhou 708 (NWTC).Minxian: Eujing, Zhongehuan- (HNATP);40 km W of Xining, S. X. Zf2erl0351 (PE1. gou, J. Q Wang197 (NWTC);Mt Iauodadoujidela,E P Huzhu:Beishan Forest Farm, Guo & He 9()12 (HNg&P}; VXang15240 (PE); near Mawu,E R ang 4594 (PE); (2iaotou,Gao & Wang6717 (HNWP).Jairlca: near Angla7 betweenLintao and Yuzhong,Hmanghe Exped. 1689 (PE); l,iu & Luo 1040 (HNWP).Ledu: Mayillg 'rownship, Ko- Mt Wutai,Taohe Exped 3243 fPE);Eamathan, K S Hao ngiazui, B. Z. Guo6767 (HNWP).(Jolog: Markog Forest 604 (PE).Taohe: J E Rock12829 (PE) Tsalukuto Shi- Farm,Hongjungou, W. 14 Wang 26818 (HNWP).Menyuan: men, J E Rock13127 (PE) Tianshui:Mt Xinjia, t 14 Semnyi,betw. Lihua & Dalong,Z. 14(vit2g 12/8 (HNWP, Hsia 5707 (PE) Tianzhu:Anyuan, E Q He 4516 (PE), PE);Semnyi, Zhugusi, Xielong7 B. Z. GrllO 7X96(HNWP) Zhucha,E Q He 4915 (PE);JinqiangTownship, Mt Mao- Semnyi, Hankegou,Gansu-Qinghai A,Itl)ed. 2493 (PE,\. mao,Q R Wang1869 (NWTC)Xiahe: Chillgshui, 1: E Minhe: Gushan, Nanxia, L. H. ZZ20ll2528 (EDIMFP) Wang6944 (PE); Tangar'ang,Loqngwagou, Q R War2g Xin'erbeishan,B. Z. Guo7010 (HNW}').Tollgren: llong- 7270 (NWTC)Yongdeng: Liancheng, Tulugou, J L Bai wo, B. Z. Galo10232 (HNWP);Shuarlgferlgxi, B. Z. Cuo 86/3 (NWTC)Yuzhong: Xinlong, rlear Yangdaowan, s 10186 (HNWP).Xining ISining]:Shangwu(huang, K. S. coll. s.n (PE) Ehangxian:Caotan distr, HuangheExped. Hao 779 (PE). Xunhua:Mengda Forest Farm,Glwo & 03183 (PE); Mt Guiqing7Huanghe ExpedS. 4735 (PE), Wang25058 (HNWP),Mt. DaIiJiavG. Z. Zhng 005 HediIing,Mt Kehu, HUangheEXPed5. 4655 (PE) Shi- (HNWP). Shaallxi: Huxian: Taipingyu,Hualingwan, chuan ForestFarme JinhuachiX Lian, ang et AI 79197 PharmacExped. 2019 (PE}.Hwainhsien [Huayin]: Qing- (NWTC) Zhouqu:Taozhou Forest FarmSAAng & J7LB longbei, 1Z E Hsia 4539 (PE). Mt. Taibai:rlear l)adian 00386 (PE) Zone (ChuonI):Kaclie Forest Fann, S(In & temple,1E E Hsia 4571 (PE),Sanchaixia, K. J. Fu 4441 Chen31 (NWTC;);Wancharlg;, Sarlgtanagou, LLAnS & C;hen (PE)-,Dadian temple to Dumugongtemple, Hong &: Zhtk 72 (NWTC)Nillgxia: the LiupaEzMountain Range, Y W PB85065 (PE).Shanxi: Shizipirlg,Xigou, Shanxi Exped. C>i la2sl (PE) Qinghai: Datong:Kuanghuizu, K Al Wang& T1iczn594 (PE).NIt. GTutai: Xitai, Xiachantan vil- LLOU6019 (PE);near Anchiapu,S. M LiOU597W (PE); lage, Xiejiagou,Shanxi Exped. 684 (PE).Sichuan: Baox- Dongxia,Yu, LU, GU & LZ68 (PE},l\At Laoye, LiAng Mt ing: RaozhiSNibagou, Hong & Zhong PB82105 (PE)7

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Longdongdistr., Zhonggan, Mt. Daping,>S7ichuan-Econom. 1905. TYPE:the Altai Mountains,Ledebour A59-0246 (PE);Ganyanggou, 1: 1: Yu2200 (PE);Dengeh- s.n. (lectotype,designated here. K!). igou, E H. Tu4329 (PE);Lianghekou, Mahuangpo, Zhang & Ren 5572 (PE). Barkam:near Puyajiao,X. Li 71029 Paeonia majkoN. Ketzchoweli,Notul. Syst. Geogr.Inst. (PE); near Kanzhugou,by river, X. Li 70441 (PE); Bot. Tiphlis. fasc. 21: 17. 1959. TYPE: Ge(3rgia. Pu'ermagou,209 LumberingGround, X. Li 70867 (PE3; Kartly,between Igoeti and Lamitskali,16 May1958, Bu'ermagou,X. Li 70903 (PE);near MatanX X. Li 71645 N. Ketschowelis.n. (holotype,TBI!3. (PE);Dalong. iaogou, X. Li 70685 (PE);near Kafeiqiao, X. Paeonia intermediasubsp. pamiroalaica P. N. Ovehinni- Li 71196 (PE);Zonggag, X. Li 70719 (PE),Nazhugou, X. kov, in F1. TadzhikistanskoiSSR. 4: 6-10 & 531. Li 71087 (PE).Emei: Jieyindian, E H. Tu362 (PE).Dege: 1975. TYPE:Tadzhikistan: declivitas australis jugi S. X. Jia 229 (PE).Garze: S bank of YalongjiaRiverX 14 Hissarcic,ad ripamdextram fl. Maichura,5 km ab t Cui 4348 (PE). Heishui:Shidiaolou Township, Sich- ostio, 2500 m, 12.06.1971,A. A. Glebowa6 (holo- ua7z.-Econom.A59-1208 (PE);Sandougou, Sichuan-EcoJI- type, TADnot seen). om. A59-1001 (PE). Jinchuan:Zosijiae E X. Li l Ol37 (PE).Jiulong: Mt. Baitai, between Niupo & Baitaivillage, The species has long been identifiedas Paeonia J. S. Ying 3869 (PE).Kangding: Yulingong, Yingbapo, Hu anomala(Pan, 1979), treatedas a varietyof P. hy- & He 10463 (PE);Yulin Township, Sirnaqiao, Jiang & Xiong 35625 (PE);Yulin Township, Laoyulin, X. C. Jiang brida(Krylov, 1901; Schipezinsky,1921, 1937; Ga- 36014 (PE);Taiyangshan distl:, Hu & He 10373 (PE); maulova, 1961), a variety of E anomala (Fedt- Niuguvillage, t Sichuan Exped. Kuan et al. 243 (PE), schenko, 1905; Stern, 1946; Pan, 1979), a Xirsduqiao,t Sichuan Exped. Kuan et al. 732 (PE); subspecies of R anomala (Trautvetter,1904), or Shade distr., Mi'Along,Nanshui-Beidiao Exped. 02912 even as a formof P anomalavar. hybrida(Traut- (PE);Zhonggu village, Dagaigou, t SichuarlExped. Kuan et al. 352 (PE);letween Kangding& DdwU,t C. Tsoong vetter,1860). These diverseand erroneousassign- 5()04 (PE). Lixian:Longxi, Yiduo village, Z. He 123X9 mentxwere made because: (1) the identity of P. (PE); Miyaluo,Misugou, S. Jiang A-7162 (PE); Shuan- anomala,P. intermedia,and P. hybrida was notpre- jingsi,R X. Li 1()100 (PE);Shuanjingsi, 3 33 kmmilestone? viouslyclear; (2) previousauthors emphasized the Of Cllerlg(lu-Al)ahighway, %h(ltzg & Zholl 2X742 (PE); taxonomi(value of indumentumon e arpels;and (3) Sonlallg,42 knl lllilestolleof Shuanjingisi-Balkalllhigh- way, ZII(ltlA,r& Zlloll 2267T (PE). I uhuo:P C. T.sootlA,rthe characterxof roots an(l ealy(es an(l their ( or- .S()24(1)1<); ;/Jhehot, I)y livel, J. S. Yitl;r4SH2? (Pli). Malxiall: relation were not coonsi(lere(l.Examination of the E T Weltlp,r229X() (1)1<). Maowell: Mt. Maf:arig,Az(hlleltl types (f the three tdxdan(l extenxive()hservati()nx (0t10t7Z. (A-.S9)2.SS4 (I)FJ); Xiallg-Nahn Scltlloggou He 011 hel })ariurll xpecinlens dn(l natural f)()T ulationx d?Slloll 1 X.X@8(1)1<). Mialillilig: Yejill 'laowilshi^), S. F: /11ll interr71esli(l 2()4()/(1'1f,);Hullilig (listr., .lie>;.ingotl, ,. JifltlA,r r76fS (I'fi). ( IaluleI ) xhowthat P. C. A. Meyerix an Muli:Wa-ehill 7' 7' YlleS1X7 (1'19). (t)iallnillg: S of towr in(3epen(1entspee iex, distinctly differillgfrom P Ji(ltl;>rsM./itl ;21 / X (X)FJ);tOI)21WU, Zh(ltz&r & I(ltlt,r .S7 ('1<). sllfl t)y its Iateralloots tut)erous;to long-fuxi- Sertar:I knl fi of W.lilg(la,S. /i(ltlg (9()82(}') %()tS,Iat: folnl, xepalsrll()xtly (at ledst2) roull(le(lat dpeXt)ut HousigotlS. /i(ltz;,rA-727T (PE); Morlt. ()(( i(l. H. ,%nitl not ( au(late . Also differentfrom P anom (ll(lX E in- 2499 (I'F3).'I'iatl(3ua1l: Mt. ,llorig, Xiaolliujirigouealoun(l Xiashan,Hll & He1()()88 ('F). Wellehuan: Wolong l'owll- terme(lia prefersrelatively esunny an(l dry habitats. ship, Si(hll(ltl-E(orzom.A.S9-229() (PE); WolongNatue Carpelsvary from 1 to 5 in numi)erean(l fromgla- Reserve,Mt. 13a1ang, Horlg & Zhll ['BUT()79(PE). Xiao- terousto densely tomentoxe.Ovehinnikov (1975) jin: Xihekou, Niuehallgt,ou, Xtl(lng & Ren 6.X22 (PF); properlytreated the in Tadzhikixtanax E 1JUO6)iFOU Yangpingkou,Zhclrlg & Kerl6()z32 (I'S). Ya- itltermeslia,but his (lescriptionof the new xubspe- jiang: Mt. E:rlllin,Sifhll(lrl Plant.s Clolleftion()&9()5 (PE); vies /)amiroalaica is unjustifiable.Hix cleseription Tanjiaoto Malangeuo, J. S. Ying .X11.S(PE). %arntang: I ukesi Temple,right l)ank, S. Jifltzg893G) (PE); 10 krnW ancl our extensive observationshave not reveale(l of Gangmu(la,S. Jifltlg 8&965(PE). Xizang (Tibet):Jom(la: any significant(lifference. Gamtogdistr., (5:itlghai-YizangExped, Veget(ltiotlGrozlp This species is widely (listribute(lin northern 9893 (PE)- Xinjiang,China (south to the Tienshan),Kazakhs- tan, Kirghizia,Ta(lzhikistan, Uzbekistan, and the 2. Paeonia intermedia C. A. Meyer,in Le(le- Altai of Russia. One small isolatedpopulation was bour,F1. Altaic 2: 277. 1830. Paeonia ano1rn- foun(lin Georgia(Hong & Zhoue2003) (Fig. 3). It ala var. hybridaf. inter1rnedia(C. A. Meyer) grows in grassy and shrubby slopes, meadows, Trautvetter,Enum. P1. Songor. 88. 1860. steppes, or in sparsewoo(ls, at altitudesfrom 900 Paeoniahybrida var. inter1rnedia (C. A. Meyer) to 3250 m. Krylov,F1. Altaya 1: 47. 1901. Paeonia an- Phenology. This species flowersfrom late May o1rnalasubsp. inter1rnedia(C. A. Meyer)Traut- to late June an(l fruitsfrom August to September. vetter,in B. Fe(ltschenko,Tru(ly Imp. S. Pe- Chrornosornes.2n = 10 (Hong et al., unpub- tersburg.Bot. Sa(la 23: 351. 1904. Paeonia lishe(l). ano1rnalavar. inter1rnedia(C. A. Meyer)0. & Additionalspecimens examined. CHINA.Xinjiang: B. Fedtsch., Beih. Bot. Centralbl.18: 216. Altai Mt.:Acad. Sinica XinjiangIntegr. Exped. 10657

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(PE). Altay: Qiao'ati, G. L. Zhu et al. 6325 (PE)- to Hal- GissarskyMountain Range: Buhara,Gissar, Mtn. Zar- amaryi, Acad. Sinica XijiangExped. 10227 (PE); Klasu kamarPass, E Lipsky1697 (LE).Darvazsky Mtn. Range, Gou, R. C. Ching2433 (PE); Haxiong Gou, T E Chou N slopes, left bank of ZarakoRiver, oppositeHavdak, 652100 (XJBI); Mt. Halamaryi, D. E Hong et al. Popu- Shinova& Kinzikaeva1550 (LE).Imam-Askari, Mt. W of lation No. 2 (PE); Xiaodong Gou, D. E Honget al. Pop- Darai-Imam,Bochantsev & Egorova1014 (LE). Peter I ulationNo. 4 (PE). Barkol: Nanshan, s. coll. 780 (XJBI). Range,N slopes,Nazarak Valley, upper reaches, 15 kmS Fuhai: Fuhai Forest Farm, G.L. Zhu 5665 (PE); N slopes, of Tadzhikabad,Ladygina, Ikonnikov & Fridman1337 T E Chouet al. 652150 (PE, XJBI). Fukang: Tienci, Shi- (LE). VahshskyMtn. Range, Mt. Hozreshi-Sho,above men, D. E Hong et al. 0190 (PE). Fuyun: Mica No. 3 Sary-Hosor,Sidorenko 185 (LE).Varzob Valley, right bank Mine, Integr.Exped. s.n. (XJBI);Mica No. 4 Mine, Integr. of MaihurGorge, NE slope, Stepanenko& Kudriashova Exped. s.n. (XJBI). Habahe: Tielieke, Kelimu 10171 2514 (LE);Mountain pass betweenrivers Havag & Kets- (XJBI);Wuzliti, Kelimu10614 (XJBI). Hoboksar:Songshu gug, N. Gontscharow2051 (LE);upper Horanchon,Za- Gous. coll. 7385 (XJBI); Mt. Qiaganebo, Integr.Exped. priagaev351 (LE).UZBEKISTAN. Tashkent Province: 10563 (PE, XJBI). Huocheng: Aksu Commune, E R. Ling BolshoyChimgan, Z. von Minkwitz1190 (LE);Bolshoy 74848 (PE); Daxigou, Acad. Sinica Exped.10402 (PE); Chimgan,N stonyslopes, E Bochantsev547 (LE). Xin'ertai, Yeguolin, C. 1E Chang et al. 3316 (XJBI); Xiaoxigou, A. R. Li & J. N. Zhu 10414 (PE, XJBI). Mori: Nangou, S. L. Chen0040 (PE). Qapqal: 58 km from Yining LiteratureCited to Zhaosu, Inst. Biologyand Pedology66 (XJBI). Qitai: Anderson,G. 1818. A monographof the genusPaeonia. Biliuhe, J. C. Zhao82-4288 (PE, XJU); Kuangou,Biology Trans.Linn. Soc. London.12(1): 248-283. 81-8381574 (XJBI); Tangfangmen Forest Farm, Integr. Dai, K. M & T. H. Ying.1990. A newspecies of the genus Inst. Bio. Res. 5788 (XJBI); Yuejin Commune, S. Q. Lin Paeoniafrom China. Bull. Bot. Res. (Harbin)10(4): 33- et al. 34 (XJBI). Tacheng: Mt. Baerleike, N slope, Integ. 41. Exped.1200 (XJBI). Toli: Mt. Albakzin, mountain slope, De Candolle,A. P. 1818. Regni VegetabilisSystema Na- K. C. Kuan 2611 (PE, XJBI). Xinyuan: Nanshan, T E turale1: 386-394. Paris. Chouet al. 650392 (PE). Yining: Yining Forest Farm, Jir- . 1824. ProdromusSystematis Naturalis Regni Ve- toulong valley, D. E Honget al. Population1 & 092 (PE). getabilis1: 56-66. Paris. Yumin:The Wild Bada PrunusNature Reserve, J. C.Zhao Fedtschenko,B. A. 1904. FloraZapa(lnage Tyan-shanya 85-293 (PE, XJU). GEORGIA. Kartly: between Lamis- 1: 103-105. Acta Hort.Petrop. 23(2): 351-353. tskali and Igoeti, Hong& ZhouH99033 (PE). KAZAKHS- Gamaulova,A. P. 1961. Paeonia.Pp. 12-13 in N. V.Pav- TAN. Ili-Pribalkhash: NW ridges of Mt. Kirgizsky lov (editor),Flora of Kazakhstan,4. rl'ab.2. Alma-Ata. Range, upper Sugata River, E Golos-Kokovs.n. (LE); Is- Hong,D. Y. & S. L. Zhou.2003. Paeoni(l(Paeoniaceae) syk-Kul Provin(e, Santash, Dongulsky garden, Aidarova in the Caucasus.Bot. J. Linn.Soc. 143: 135-150. & Kashenkos.n. (LE), Semirechensk Province, N slope of , Z. X. Zhang& X. Y. Zhu. I988. Studieson the Alatau, between Verny to Kastek mountain pass, V Lad- genusPaeonia (1)-Report of karyotypesof somewild ygin s.n. (LE); SW SongorianAlatau, Mt. Emelsky Range, species in China.Acta Phytotax.Sin. 26(1):33-34. Mt. Matai, E Goloskokovs.n. (LE), Taldy-KurganProvince, , K. Y. Pan & X. Y. Li. 1994. Paeoniain Xinjiang, Mt. Altyn-Emel Range, Mt. Engi-Altyn-Emel pass to road China.Acta Phytotax.Sin. 32(4):349-355. Sary-Ozek-Dzharkent,Grubov & Luboarsky229 (LE);Zai- , K. Y. Pan & N. J. Turlancl.2001. Paeoniaan- liisky Alatau, Usun-Kargali River Valley, W Sukonnaya omalasubs p. veitchii(Paeoni ace ae) , a newcombination . Fabrika, E Goloskokov(LE). E. Kazakhstan: Mt. Kalbin- Novon11: 315-318. sky Range, Mt. Kok-tau, near Sabinskie Lake, A. Yunatov Huth, E. 1892. Monographieder GattungPaeonia. En- 24 (LE); Semipalatinsk Province, Zaisan distr., Kusto gler'sBot. Jahrb.14: 258-276. Gorge, B. Shishkins.n. (LE); Zaisan distr., near Kender- Krylov,P. 1901. Floraof the Altai l: 47. rlaomsk. lyk, B. Shishkin.s.n. (LE); near Zaisan, Mt. Kichken-tau, Ledebour,C. F. 1842. FloraRossica l: 74. Stuttgart. Tatarowa& Borsowa52 (LE). Mountain Kazakhstan: Linnaeus,C. 1771. MantissaPlantarurn. 247. Holmiae. Semirechensk Prov., Songorian Alatau, S of Lepsinsk, R. Lynch, R. I. 1890. A new classificationof the genus Kozhevits55 (LE); W ridges of Songorian Alatau, Koksu Paeonia.J. Roy.Hort. Soc. 12: 428445. Gorge, near Koksuisky, E Goloskokov(LE); Tarbagatai, Meyer,C. A. 1830. Paeonia.Pp. 276-279 in C. F. Led- Chenarak, steppes, Grigoriev& Buhaeva298 (LE); Mt. ebour(editor), Flora Altaica. Berolini. TsarytsynskyPass, deep gorge from the pass, Schipezinsky Ovehinnikov,R. 1975. FloraTadzhikistanskoi SSR 4: 6- 125 (LE). S. Kazakhstan: Mt. Chatkalsky Range, Chat- 10 & 531. kal Valley, Mt. Piazak, O. Knorring60 (LE); Prichuisky Pallas,P. S. 1789. FloraRossica 1(2): 92-95, tab.84-87. Mujun(Kum), Sulunger Gorge, E Zapriagaev66 (LE). St. Petersburg. KIRGHIZIA. Dzhalalabad Province: Bazar-Kurgan Pan, K. Y. 1979. Paeonia.In: W. '1'.Wang (editor), Flora distr., Mt. Fergansky Range, S slopes, near AK-Terek,Le- Reipublicae Popularis Sinicae 27: 37-59. Science bedeva& Faleevas.n. (LE); Kara-Alma,Toguz (Bulak), N Press,Beijing. slopes, Poniatovskaya,s.n. (LE); Tianshan, Mt. Ketmen Schipezinsky,N. V. 1921. Synopsisof the genusPaeonia. Range, Mt. Elchin-Ustriuk, Kizilbulak Gorge, s. coll. s.n. Not. Syst. Herb.Hort. Bot. Petrop.2(11-12): 4147. (LE). Fergana Province: Kirgizsky Alatau, Makbal . 1937. Paeonia.In: V. L. Komarov(editor), Flora Gorge, Ty-KarynRiver, G. Igolkin97 (LE); Oshsky distr., URSS7: 24-35. EditioAkademiae Scientiarum URSS, Ak-Bura valley, Mt. Kanida Pass, O. Knorring25 (LE). Leningrad. Semirechensk Province: Pishpeksky distr., Uzungyr- Schmitt,E. 1999. Les Pivoines.Etude systematiquedu skaya Volost, Mt. Aleksandrovsky Range, Mt. Shekule N genrePaeonia L. Plantesde Montagne191: 574-583. slope, A. Savenkova29 (LE). RUSSIA. Altai: AlexShrenk Stern,F. C. 1946. A Studyof the GenusPaeonia. Royal s.n. (LE); Altai, Barnaul Province, between Kalmytskie HorticulturalSociety, London. Mysy & Mahanova,R Krylovs.n. (LE). TADZHIKISTAN. Trautvetter,E. R. 1860. Enumeratioplantarum songori-

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carum,a Dr. Alex Schrenkannis 1840-1843 collectar- (2); Iljin,V. s.n. (LE)(la); Integr.Exped. 630 (XJBI)(la), um Bull. Soc. Nat. Mosc.33: 87-88. 1200 (XJBI)(2), 7305 (XJBI)(la), 10563 (PE, XJBI)(2), . 1904. Paeonia. In: B. A. Fedtschenko(editor), 11829 (XJBI)(la), s.n. (XJBI)(2); Integr.Inst. Hydrol. Flora of the West Tianshan(Acta Hort. Petrop.)23: Pedol & Biol. Res. 5788 (XJBI)(2). 351-353. Jia, S. X. 229 (PE) (lb); Jiang& Jin 00386 (PE)(lb), 2113 (PE) (lb); Jiang& Xiong 34242 (PE) (lb), 34319 APPENDIX1 (PE)(lb), 35625 (PE)(lb); Jiang,S. 5768 (PE)(lb), 8874 TO EXStCCATAEEXAMINED (PE)(lb), 8930 (PE)(lb), 8965 (PE)(lb), 9082 (PE)(lb), A-7162 (PE) (lb), A-7275 (PE) (lb); Jiang,X. C. 36014 Thefigures in parenthesesrepresent: la = Paeoniaan- (PE)(lb), 36433 (PE)(lb). omala subsp.anomala; lb = P. anomalasubsp. veitchii; Kelimu 10171 (XJBI)(2), 10369 (XJBI)(2), 10614 2 = R intermedia. (XJBI)(2); Keller,B. s.n. (LE)(la); Ketzchoweli,N. s.n. Acad. Sinica Bot. Inst. Exped. Xinjiang 1803 (PE, (TBI)(2, type of Paeonia majkoN. Ketzchoweli);Kildu- XJBI)(2); Acad. Sinica Exped. 10402 (PE)(2); Acad. Sin- shewsky,I. 150122(LE) (la); Klements,E. 4d (LE)(la), ica XinjiangExped. 2496 (PE, XJBI)(la), 10227 (PE) lla (LE)(la); Klopotov,B. 113 (LE)(la), 180 (LE)(la); (2); Acad.Sinica Xinjiang Integr. Exped. 10657 (PE)(2); Knorring,0. 25 (LE) (2), 66 (LE) (2); Kokulin,S. 254 Agr. College 1051 (PE) (2); Aidarova& Kashenkos.n. (LE)(la); Kolomoitseva,S. 236 (LE)(la); Komarov,V. s.n. (LE)(2); Alexandrov, P. 179 (LE)(la), 219 (LE)(la); Alex (LE) (la); Korovkin,A. 290 (LE) (la); Krasnoborov,I. Shrenks.n. (LE)(2). 6765 (LE)(la); Krishtofowich,A. s.n. (LE)(la); Krylov, Bai, J. L. 8613 (NWTC)(lb); Beresowskii,M. s.n. (LE) P. s.n. (LE)(2); Kuan,K. C. 2611 (PE, XJBI)(2), 77309 (lb, typeof Paeoniaberesowskii Komarov); Biology Exped. (PE)(lb); Kuanet al. 243 (PE)(lb), 352 (PE)(lb); Kuch- 79250198(XJBI) (2), 81-8330478(XJBI) (2); Bochantsev, erovskaya,S. 282 (LE) (la); Kuminova& Alexeevas.n. V. 547 (LE) (2); Bochantsev& Egorova1014 (LE) (2); (LE)(la); Kuvajev& Sabitovs.n. (LE)(la); Kuznetsov,I. Borovikov,G. s.n. (LE)(la); Borovikovet al. s.n. (LE)(la). 53 (LE)(la), 114 (LE)(la), 193 (LE)(la), 4661 (LE)(la); Chang,C. W. 11356 (XJBI)(la); Chang,C. W. et al. Kuznetsov,N. 199 (LE)(la), 1991 (LE)(la). 3316 (XJBI)(2), 4037 (XJBI)(2), 4357 (XJBI)(la), 10255 Ladygin92 (LE)(la), s.n. (LE)(2); Ladygina et al. 1337 (XJBI)(la); Chen & Ju s.n. (NWTC)(lb); Chen, S. L. (LE)(2); Lashinsky& Ronginskaya973 (LE)(la); Lebed- 0040 (PE)(2); Cherepnin s.n. (LE)(la); Ching,R. C. 1803 eva & Faleevas.n. (LE)(2), Ledebour,s.n. (K)(2, type of (PE) (la), 2433 (PE) (2), 2496 (PE) (la); Chou, T. Y. Paeoniaintermedia C. A. Meyer);Li, A. R. & Zhu,J. N. 652100 (XJBI)(2); Chou,T. Y. et al. 650392 (PE) (2), 10414 (PE, XJBI)(2); Li, P. X. 10100 (PE) (lb), 10137 650868 (PE, XJBI)(2), 652150 (PE, XJBI)(2); Cui, N. (PE)(lb), 10143 (PE)(lb); Li, X. 70247 (PE)(lb), 70441 R. 091 (XJNU)(la), 86624 (XJNU)(la); Cui,Y. W.4348 (PE)(lb), 70510 (PE)(lb), 70685 (PE)(lb), 70719 (PE) (PE)(lb), 10281 (PE)(lb). (lb), 70867 (PE)(lb), 70903 (PE)(lb), 71029 (PE)(lb), Dolenko,G. 108 (LE)(la); Dranitsyn& Kochubeis.n. 71087 (PE)(lb), 71196 (PE)(lb), 71645 (PE)(lb), 74842 (LE)(la). (PE) (lb), 74936 (PE) (lb), 77248 (PE) (lbF type of Ermolajeva,M. s.n. (LE) (la); Evseeva,Z. 4406 (LE) Paeoniaveitchii var. Ieiocarpa W. T. Wang& S. H. Wang); (la). Li, X. Y. 870147 (SHI)(la), 870169 (SHI)(2); Lian & Fang,W. P. 4213 (LE)(la), 6037 (LE)(la); Farrer,R. Chen31 (NWTC)(lb), 72 (NWTC)(lb); Lian et al. 378 67 (E) (lb, type of Paeonia woodwardiiStapf ex Cox); (HNWP)(lb), 79179 (NWTC)(lb); Licent,P. 2022 (PE) Fedorov& Iljina72 (LE)(2), 128 (LE)(2); Fu, K. J. 4441 (lb); Lin, S. Q. et al. 34 (XJBI)(2); Ling, Y. R. 1141 (PE) (PE)(lb); Fu, T K. 834 (PE)(lb). (la, type of Paeoniasinjiangensis K. Y. Pan),74243 (PE) Ganeshin,S. s.n. (LE) (la); GansuHerbs Groups.n. (2), 74848 (PE) (2), 74849 (PE) (2); Liou, K. M. 5659 (NWTC)(lb); Genina,S. s.n. (LE) (la); Gansu-Qinghai (PE)(lb), 5970 (PE)(lb), 6019 (PE)(lb), 6178 (PE)(lb); Exped.2493 (PE)(lb); Goloskokov,V. s.n. (LE)(2); Gon- Lipsky,V. 1697 (LE)(2); Liu & Luo 1040 (HNWP)(lb); tscharow,N. 2051 (LE)(2); Gorodkov,B. 512 (LE)(la); Liu & Shen 8308 (XJBI)(la); Liu, S. W. 2115 (HNWP) Grigoriev& Buhaeva298 (LE)(2); Grubov& Luboarsky (lb); Lomonosova& Ivanova89 (LE)(la); Long,X. F. 85 229 (LE) (2); Guo, B. Z. 6767 (HNWP)(lb), 7010 (NWTC)(lb). (HNWP)(lb), 7396 (HNWP)(lb), 10186 (HNWP)(lb), Maltsev,N. 79 (LE)(la); Mao,Z. M. 10413 (XJBI)(la); 10232 (HNWP)(lb); Guo& He 9012 (HNWP)(lb); Guo Mihno,P. s.n. (LE)(la); MinkwitzZ. von 1190 (LE)(2); & Wang6008 (HNWP)(lb), 6717 (HNWP)(lb), 25058 Morgan,P. 23 (K)(la). (HNWP)(lb); Guo& Yang9608 (HNWP)(lb). Nanshui-BeidiaoExped. 02912 (PE)(lb). Hanminchun,V. 515 (LE) (la); Hao, K. S. 779 (PE) Pharmac.Exped. 2019 (PE)(lb); Pobedimova697 (LE) (lb); He, Y. Q. 4516 (PE) (lb), 4915 (PE) (Ib); He, Z. (l a); Pohle & Rozhdestvenskys.n. (LE)(l a); Polozhii& 12339 (PE) (lb); He & Zhou 13338 (PE) (lb); Hong D. Kandasovas.n. (LE) (la); Poniatovskayas.n. (LE) (2); Y. et al. Population2 (PE) (2), Population3 (PE) (la), Price,M. P. 293 (LE)(la). Population4 (PE) (2), Population5 (PE)(la), Population Qing,Z. Y. 1218 (HNWP,PE) (lb); Qinghai-XizangEx- 1 (PE)(2), Population0190 (PE)(2); Hong D. Y. & Zhong ped. Veget.Group 034 (PE)(lb, type of Paeoniaveitchii PB82105 (PE) (lb); Hong D. Y. & Zhu PB85019 (PE) var.uniflCora K. Y. Pan),9893 (PE) (lb); Qu, Z. X. 2792 (lb), PB85040(PE) (lb), PB85065(PE) (lb); Hong,Luo (PE)(lb). & He H95034 (A, CAS, K, M0, PE, US) (lb); Hong& Reznichenko,V. 49 (LE)(la), 58 (LE)(la); Reznichen- Zhou H99033 (PE) (2); Hsia, W. Y. 4539 (PE) (lb) ko, V. & A. 106 (LE)(la), 360 (LE)(la), s.n. (LE)(la); 4571(PE)(lb), 5707 (PE) (lb); Hu, W. G. 13050 (PEj Rock,J. F. 12260 (PE)(lb), 12829 (PE)(lb), 13127 (PE) (lb); HuangheExped. 1689 (PE) (lb), 3183 (PE) (lb), (lb); Rohderi,I. s.n. (LE)(la); Rozhevits,R. 55 (LE)(2); 4655 (PE) (lb), 4735 (PE) (lb); Hu & He 10088 (PE) Rubin,A. s.n. (LE)(la); Rubin& Maskil,s.n. (LE)(la). (lb), 10373 (PE) (lb), 10463 (PE)(lb), 11173 (PE)(lb); Sanadzeet al. s.n. (TBI)(2); Savenkova, A. 29 (LE)(2); Huang,Luo & Jiang730 (PE)(lb). Schipezinsky125 (LE) (2); s. coll. Siberia,(LINN) (la, Igolkin,G. 97 (LE)(2); Ikonnikov& Fridman892 (LE) type of Paeonia anomalaLinnaeus); Sedelnikov, A. s.n.

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{LE) (la); S. Gansu Grassland Exped. 681 (NW TC)(lb), Y. et al. 27347 (HNWP) (lb); Wang & Zbou 19 (HNWP) Shanxi Exped. 684 (PE) (lb); Shanxi Exped. Wang & Tian {lb), 198 A (HNWP) (lb). 594 (PE) (lb); Shinoara& Kinzikaeva 1550 (LE) (2); Xinjiang Branch Acad. Sinica 012 (XJBI (la); Xu, L. Shishkin, B. s.n. (LE) (2); Shliahtin, A. s.n. (LE} (la); R. s.n. (PE) (la). Shukin, S. s.n. (LE) (la); Sichuan-Econom. A59-0246 Yakovlev, D. 69 (LE) (la); Yan, M. S. 1843 (NWTC) {PE) (lb), A59-1001 (PE) (lb), A59-1208 (PE) (lb), A59- {lb); Ying, J. S. 3113 (PE) (lb), 3869 (PE) (lb), 4582 2554 (PE) (lb), Sichuan-Econom. & Ya 838 1PE) (lb); (PE)(lb)7 4591 (PE)(lb); Ying, T. H. 995 (PE, SHMU) Sichuan Plants Collection 0905 (PE) (Ib); Sidorenko 185 (I a), 1001 (PE, SHMU) (la), 1006 (PE, SHMU) (la), 1007 (LE) (2); Smirnov, V. s.n. (LE) (la); Smith, H. 2499 (PE) (SHNIU,PE) (la, type of Paeoniaaltaic(l K. M. Dai & T. (lb); Sochava, V. 100 (PE)(lb); Sokolov, P. 13a (LE) (la); H. Ying), 1008 (PE) (la), 1009 (PE, SH1WU)(la), 1010 Song, Z. P. 38544 (PE) (lb), 39107 (PE) (lb); Stepanenko (PE, SHMU) (la), 1011 (PE, SHMU) (la), 1014 (PE, & Kudriashova2514 (LE) (2); Suhareva, A. s.n. (LE) (la). SHMU) (La), 1022 (PE, SHMU) (la); Yu et al. 68 (PE) Tang, T. 938 (PE) (lb); Taohe Exped. 3243 (PE) (lb); (lb); Yu, T. T. 2200 (PE) (lb), 6137 (PE) (lb); Yunatov, Tatarova& Borsowa52 (LE) (2); Tolmatschew,I. 247 (LE) A. 24 (LE) (2); Yurchenko, P. s.n. (LE) (la). (la); Tomin, M. 67 (LE) (la); Tripolitova,T. s.n. (LE) (2); Zapriagaev,F. 66 (LE) (2), 351 (LE) (2); Zhang, F. M. Troitsky,W. s.n. (LE) (la); Tsoong, P. C. 5004 (PE) (lb), 90-346 (XFNU) (la); Zhang, G. Z. 005 (HNWP) (lb); 5024 (PE) (lb), 8842 (PE) (lb); Tu, T. H. 362 (PE) (lb) Zhang & Lang 57 (PE) (lb); Zhang & h 0502 (HNWP) 4329 (PE) (lb); Tugarinov,A. s.n. (LE) (la); Turezaninow, (lb); Zhang & Ren 5572 (PE) (lb), 6082 (PE) (lb), 6322 s.n. (LE) (la)- (PE)(Ib); Zhang, Z. H. et al. 0097 (HNWP) (lb), 0414 Varentsov, V. s.n. (LE) (la), Vereshagin, V. 296 (LE) (HNWP) (lb), 4291 (HNWP) (lb), 4317 (HNWP) (lb), (la), 345 (LE) (la); Verhovskaya& Mishin 104 (LE) (la); 4320 (HNWP) (lb); Zhang & Zhou 22092 (PE) (lb), Vislouh, E. 73 (LE) (la); Vydrin, A. s.n. (LE) (la). 22155 (PE) (lb), 22675 (PE) (lb), 23742 (PE) (lb); Zhao, Wang, F. T. 2293() (PE) (lb); Wang, J. Q. 197 (NWTC) J. C. 82-293 (PE, XJU) (2), 82-4288 (PE, XJU) (2), s.n. (lb); Wang7Q. R. 1869 (NWTC)(lb), 7034 (NWTC)(lb), (PE, XJU) (2); Zhen, S. X. 0351 (PE)(lb); Zhou, H. J. 7270 (NWTC) (lb); Wang, S. X. 1065 (HNWP) (la); 708 (NWTC) (la); Zhou, L. H. 2528 (HNWP) (]b); Zhu7 Wang, T. P. 197 (NWTC)(la), 15240 (NWTC) (la), 4594 G. L. 5665 (PE) (2), 5755 (PE) (la), 6345 (PE) (la), (NWTC)(la), 6944 (PE) (lb); Wang,W. Y. 26771 (HNWP) 80006 (NWTC)(lb); Zhu, G. L. et al. 6325 (PE) (2), 6386 (lb), 26818 (HNWI') (lb), 27060 (HNWP) (lb); Wang,W. (PE) (2); Zhu, S. F. 20491 (PE) (lb).

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