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Bulletin of The Natural History Museum Series Botany THE NATURAL HISTORY MUSEUM VOLUME 32 NUMBER 2 28 NOVEMBER 2002 TheBulletin ofTheNaturalHistoryMuseum (formerly: Bulletin oftheBritish Museum (NaturalHistory)),instituted in 1949, is issued in fourscientific series,Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's DepartmentofBotany KeeperofBotany: Prof. R. Bateman EditorofBulletin: Mrs M.J. Stafford Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum,bothbythe scientific staffandby specialistsfromelsewherewhomakeuseoftheMuseum'sresources. Manyofthe papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peerreview before acceptance. SUBSCRIPTIONS BulletinoftheNaturalHistoryMuseum, BotanySeries(ISSN0968-0446)ispublishedtwiceayear(one volume perannum) inJune and November.Volume 32 will appearin 2002.The 2002 subscription price (excludingVAT)ofa volume, whichincludesprintandelectronic access, is 88.00(US $155.00inUSA,CanadaandMexico).Theelectronic-only price available to institutional subscribers is 79.00 (US $140.00 in USA, Canada and Mexico). ORDERS Orders,whichmustbeaccompaniedbypayment,maybesenttoanybookseller,subscriptionagentordirecttothe publisher: Cambridge University Press, The Edinburgh Building, Shaftesbury Road, Cambridge CB2 2RU, UK; or in the USA,CanadaandMexico:CambridgeUniversityPress,JournalsDepartment,40West20thStreet,NewYork,NY 1011-4211, USA. EU subscribers (outside the UK) who are notregistered forVAT shouldaddVATattheircountry's rate.VATregistered members shouldprovidetheirVATregistrationnumber.Japanesepricesforinstitutions(includingASPdelivery) areavailable from Kinokuniya Company Ltd, P.O. 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This consent does not extend to multiple copying for promotional or commercial purposes. Code 0968-0462/2002 $16.00. ISITearSheet Service, 3501 MarketStreet, Philadelphia, PA 19104, USA, is authorisedto supply singlephotocopies of separate articles for private use only. Organizations authorised by the UK Copyright Licensing Agency may also copy material subject to the usual conditions. Forall otheruse, permission shouldbe soughtfrom Cambridge University Press. No part ofthis publication may otherwise be reproduced, stored or distributed by any means without permission in writing from Cambridge University Press, acting forthe copyright holder. Copyright The Natural History Museum, 2002 ELECTRONIC ACCESS This journal is included in the Cambridge Journals Online service which can be found at: http://journals.cambridge.org Forfurtherinformation on otherPress titles access http://uk.cambridge.org orhttp://us.cambridge.org World list abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) ISSN 0968-0446 The Natural History Museum Botany Series Cromwell Road Vol. 32, No. 2, pp. 61-156 London SW7 5BD Issued 28 November 2002 TypesetbyAnnBuchan(Typesetters),Middlesex PrintedinGreatBritainbyHenryLingLtd,attheDorsetPress,Dorchester,Dorset Bull. nut. Hist. Mas. Land. (Bot.)32(2):61-123 Issued28November2002 Studies in the genus Hypericum L. (Guttiferae) 4(2). Section 9. Hypericum sensu lato (part 2): subsection 1. Hypericum series 1. Hypericum NORMAN ROBSON K.B. 62 N.K.B. ROBSON H. ascyron subsp.ascyron [W.SiberiatoJapan,China] Spp.20-42 [SakhalintoBorneo] 2-4.H. undulatumgroup [Macaronesia,Europe Mediterranean,W.Asia] ser. 2.Senanensia Spp. 13-19 [Japan,Korea,C.China] 1. H. maculatums.l. 12.H. scouleri [Euro-Siberia] [W.North America 5.H.perforation toC.Mexico] [Eurasia,Med.] 6.H. altenualum [C.Siberia,.China,Korea] l.H. elegans [C.SiberiatoC.Europe] 8-11. H.yezoensegroup [Japan,Sakhalin,S.Kuriles] Fig. 1 Section9. Relationshipsanddistribution. taxaofsubsect.Erecta(H.yamamotoivar. ripariumY. Kimuraand derivative of a cross between them, especially as H. perforatum H. ovalifolium Koidz.) have abranching woody stembase, likeH. behavescytologicallyasahybrid(seep. 64). However,thenearest ascyron, but the stem internodes are slightly 2-lined or terete, the subspecies ofH. maculatum to H. perforatum morphologically is linesbeingeglandular. Also,theleavesintheH.yamamotoiandH. not the one that occurs in Siberia today (subsp. maculatum) but erectumgroupsofsubsect. Erectaarefrequentlytriangular-lanceo- subsp. immaculatum, which is now confined to the Balkan region. late,likethoseof//, ascyronsubsp.ascyron,whereasacomparable Thehypothesismustthereforebethatsubsp. immaculatumatsome formofleafoccursonlyapomorphicallyinsubsect.Hypericum,i.e. timewaspresentinSiberia,whereithybridizedwithH.attenuatum. only in someofthe moreadvanced species. The chromosomes of the resultant diploid hybrid then doubled, giving rise to an allotetraploid species that is partially fertile (see Distribution andevolution (Fig. 1) p. 88).Subsequentlytheareaof//,maculatumsubsp.immaculatum TnohrethdiasntdriebausttioAnssiaonf,sbuubtsetchtat.o1fsseurbiseesct2. a1nsderiseusbs1ecetx.te2ndasrefrwohmoltlhye bapeocmaomrephgirecattolyitraenddutcheuds;abluitketlhyadteroifvastuibvsep.ofmiat,ceuxltaetnudme,dewahsitcwharids into Siberia. Azores to Mexico. The latter series comprises a western group, based in south-eastern Europe on 1. H. maculatum and extending Sect. 9.Hypericum subsect. 1.Hypericum series 1. westtotheAzoresandeasttocentralSiberia,andaneasterngroup Hypericum with two centres, one in eastern Siberia and the other in western sNcoorutlherAi,mewrhiciac.hThhaesNiotsrtmhosAtmeprriicmiatnivceenftroermconisnistthseosnoluytho-fea1s2t.eHr.n Characters andvariation (Figs 2, 3) UnitedStatesandshowstrendsnorthwardtocentralBritishColum- MORPHOLOGY. ThetwomaindivisionsofseriesHypericumcould bia and southward to Michoacan in central Mexico. The basic be separated on whether or not the subsidiary stem lines were species ofthe east Asian centre is 6. H. attenuatum, which has its present,i.e. whetherthesteminternodeswere4-or2-lined, wereit most primitive form in eastern Siberia and a range that extends notfor(a)4.H. triquetrifolium,whichhas 'lost' theminthe4-lined westwardtotheAltairegionandsouthwardtosouth-centralChina. group and (b) 5. H. perforatum, which, as has been explained, Thebasic speciesofthederivativeJapaneseH.yezoensegroup(H. appears to be a hybrid with one parent from each group. Figure 3 tosaense)ismorphologicallyverynearH. attenuatumandhasbeen showsthatthe4-linedstateisprimitiveinthisseries,asindeeditis included in it (Kitamura & Murati, 1962), but in my opinion it is in the whole genus. Intermediate states occur in hybrids of H. specificallydistinct.Hypericumelegans,awesternderivativeofH. perforatum, butnotinthe species. attenuatum, extends the distribution ofthe eastern group as faras Thelaminarleafglandsareabsentormorerarelysparsein l.H. westernGermany. maculatum but more or less dense in the more advanced species. Theremaining species in series 1,5. H.perforatum,does notfit They are all or mostly pale except in 6. H. attenuatum, in which easily into the above scheme. Morphologically it is intermediate blackglandsoften predominate. between 1. H. maculatum and 6. H. attenuatum, and the areas of Thesepalsarebroadwitharoundedapexin 1.H.maculatumand these speciesalmostoverlapinwesternSiberia(Maps 1, 16). Each usuallyin 12.H.scouleri,astheyareintheancestral(?)H.ascyron of them is diploid (2n=16), whereas H. perforatum is tetraploid subsp. ascyron (sect. 7. Roscyna). In all the other species they are (2n=32);soitisconceivablethatH.perforatumisanallotetraploid acute and narrow (Spp. 2, 3 and 5) or acute and broad (6. //. STUDIES INHYPERICUM 63 3a.H. tetrupterum var.anagallidifolium 2b.//. undulalum 11. //.yezoense 3b. H. tetrapterum var. boeticum 12.//. scouleri var.corsicum 16 ~~~~~~~~~~~, 3. H. tetrapterum Za. //. undulatum var. tetraplerum var.undulatum 10. //. momoseanum 16 16,32 I 9.H. iwatelittorale 4.//. triquetrifolium la //. maculatum 16 .ssp. immac\ulatum 6. //. attenuatum -8. //. tosaense (16) 1b. //. maculatum 1c.//. maculatum 1. H. elegans ssp.maculatum ssp. obtusiusculum 5.//.perforatum 32 16 32 ~ 16,32 48 f 5a.//.perforatum ssp.perforatum 5b.//.perforatum 32,48 ssp.songaricum 5d. //.perforatum 5c.//.perforatum ssp.chinense ssp.veronense 32[16] Fig.2 Section9,subsect. 1,series 1.Relationshipsand2nchromosomenumbers. attenuatum) or acuminate and broad (Japanese spp.) except 4. //. CYTOLOGY. Althoughthediploidchromosomenumberinsect.7. triquetrifolium,inwhichthemorereducedformshave(secondarily) Roscynaisusually 18,itrangesfrom22to 16(Robson,2001);and smallroundedsepals.Theapexisentireexceptin2c.//.maculatum 2n=16 appears to be the basic number for the whole of sect. 9. subsp.obtusiusculumand//. xdesetangsii(2.//.maculatumx5.H. Hypericum. Inseries \.Hypericum(Fig. 2), Spp. Ib,2-4,6and 12 perforatum), where itis usuallyeroded-denticulate. are diploid, Sp. la has been shown by indirect evidence from Thepetallaminarglandsareprimitivelypaleandlineartostriiform morphologyandhybridstobediploid,andtheJapanesespecies(8- (la.//.maculatumsubsp.immaculatum, 12.//. scouleri),asinsect. 11)seemlikelytobeso.Tetraploidy(2n=32)hasbeenrecordedin2. 7. Roscyna, and evolve in two directions: (a) the lines become H. undulatum but without reference to morphology, in Ic. H. interrupted, firsttowardtheapex(Spp. Ib, 5,6, 8-10, 12)andthen maculatum subsp. obtusiusculum, which appears to be an throughout the lamina (Spp. 2-4, 7, 11) and sometimes disappear autotetraploidofla.subsp.immaculatum,in7.H. elegans,whichis altogether(Spp.2,3);(b)somepaleglandsbecomeblack,butthese possibly a direct derivative of 6. H. attenuatum, and in 5. H. usually remain in the minority except in 6. H. attenuatum and perforatum,whichisapparentlyanallotetraploidfromthecross la. usually Ib. H. maculatum subsp. maculatum, where all glands H. maculatumsubsp.immaculatumx6.H.attenuatum.Hexaploids become black. (2n=48) have also been recorded in H. perforatum, presumably Thevittaeoftheovaryandcapsuleareprimitivelyallnarrowand resultingfromthefertilizationofanunreduced(apomictic)ovuleby longitudinal, as in sect. 7. Roscyna; but the lateral vittae become anormallyreducedpollengrain(seep.88).Thediploidsreportedin oblique, in4. H. triquetrifoliumonly sporadically, inotherstotally thisspeciesmustbedihaploidsresultingfrompolyhaploidy,i.e.the (Spp.6-11).In5.H.perforatumalonetheobliquelateralvittaefirst secondary occurrence of the diploid number as a result of the becomeenlargedandthenbreakuptoformovaltoglobosevesicles. development ofan unfertilized reduced ovule, which is known to sepalapex acuteto\ rounded 3c acuminate: toobtuse 2a blackpetallaminarglands absent ,- Fig.3 Section9,subsect. 1,series 1.Limitsofsomecharacters. 64 N.K.B. ROBSON subsect. Erecta Spp.2(M1 4. H. triquetrifqlium subsect. Hypericum 16 2*3. H. undulatum* tetrapterum seriesScnanensia Spp. 13-19 seriesHypericum 3.H. tetrapterum .2.H. undulatum [Svars.J 16~~-:::; ___ [2vars.] 16,32 11.//.. 12.//. scoukri ~~~ 1xa.H. x'laschii 1xb.//. *laschii 16 10. H.^momoseanum nf.laschii nf.froelichii (16) \ (24) 9.//. iwatelittorale la. //. maculatum 6.//. attenuatitm ssp. immaculatum 16 8.//. losaense ^ (16) 1b.//. maculatum ssp.maculatum lc.//. maculalum 5.//.perforatum ,6 ssp. obtusiusculum "^ -"I" [4sspp.] [16]32,48 7.//. elegans 32 ( V 5xb.H. xdesetangsii 5xa. //. xdesetanggssiiii -"~""\. / """"-.>. ! nssp.carinthiacum nssp.desetangsii 5xc.//. xdesetangsii 5 * 7.//.perforatum x elegans 24,40 x^48 --'" nssp. balcanicum -~^ (32) 40 (24,40) 5xba.nf.maculatiforme 5xbb.nf.perforatiforme 5xca. [nf. a] 5xcb. [nf. b] 24 40 (24) (40) Fig.4 Section9,subsect. 1,series 1.Relationships( )andhybrids( ),with2nchromosomenumbers(unverifiednumbersinparenthesis). occurinallopolyploids(seep. 88andRobson, 1998).Thereportof dependingontheploidyoftheotherparentandofthegametesof//. atriploidnumber(n=12)byGagnieu&Wilhelm(1965)islikelyto perforatumitself.Aneuploidnumbersoccurinexperimentalcrosses be an error for n=16. A hypoploid number (2n=8, n=4) has been (e.g. thoseofLihovaet al., 2000), butthey havenotbeen reported reportedfor2.H. undulatum(Guillenetal., 1997),aspecieswhich fromwildpopulations.Noack(1939)foundthat,inhybridsinvolv- is normallydiploid(2n=16)ortetraploid(2n=32). ing diploids, the triploids were intermediate between the parents whereas the pentaploids could not be distinguished from pure H. Hybrids (Fig. 4) perforatum.Martonfietal.(1996/7),however,foundthatapentaploid Natural hybridization is relatively rare in Hypericum (Robson, populationofH. maculatum subsp. maculatum xperforatumfrom 1981: 167, fig. 55)exceptin series 1. Hypericum, where itisquite Slovakia did have some intermediate characters as well as the common largely on accountofthe floral biology ofH. perforatum pentaploidnumber(2n=40). (see p. 88). This species behaves as ahybrid (highpollen sterility, Tetraploid crosses with H. perforatum yield intermediates, at laggingchromosomesatmetaphase,97%apomicticovuledevelop- leastinthefirstgeneration.Thusapossiblehybrid,5.H.perforatum ment with pseudogamy). When crossed with diploids, its reduced x1. H. elegans, isrecordedhereonthebasisofamorphologically pollen grains andovules (n=16) produce triploid (2n=24) hybrids, intermediatespecimenfromtheUkraine;andthecrossH.maculatum while the fertilization of its unreduced ovules by a gamete of a subsp. obtusiusculum xperforatum also yields intermediate prog- diploidspeciesproducespentaploid(2n=40)hybrids.Withtetraploids eny in the Fr The relationship ofthe parents ofthis lattercross is suchasH. maculatumsubsp.obtusiusculum,ofcourse,thehybrids veryclose,however,assubsp.obtusiusculumbehavescytologically are tetraploid (2n=32). Figure 4 (modified and expanded from and morphologically as an autotetraploid ofsubsp. immaculatum, Robson, 1981:fig.55)showsallthehybridsthathavebeenreported not ofsubsp. maculatum, as I first thought (Robson, 1956; 1958a, or inferred from series 1. Hypericum. The status of some of the 1975). This consanguinity facilitates back-crossing and results in hybrids has been inferred from intermediate morphology, while theeventualoccurrenceofacompleteseriesofintermediatesinthe severalchromosomenumbershavebeeninferredfromthoseofthe wild (cf. Crackles, 1990). Counts of2n=48 and40 forthis hybrid parents. from cultivated plants (Robson, 1956; Robson & Adams, 1968) OfthehybridsnotinvolvingH.perforatum,2.H. undulatumx4. presumably result respectively from a cross with an unreduced H. tetrapterum,whichisknownonlyfromplantswithintermediate perforatumgamete(32+ 16)andabackcrossofthishybridtoeither morphology, appears to be rare. As the tetraploid form of H. a 'normal' hybridortoH.perforatumitself(24+ 16). undulatum has been reported only once, the hybrid is likely to be Thespecies3.H. tetrapterumisdiploidonly, soitscrosswith//. diploid. 1. H. maculatum x 4. H. tetrapterum (H. x laschii) was perforatum (H. x medium), described as morphologically inter- described as intermediate between la. subsp. maculatum and mediate, is likelytobetriploid; butthecross 1. H. maculatum x 5. tetrapterum and is known from the areaofthat subspecies only. It perforatum (H. x desetangsii) is much more complicated. The tooisthereforelikelytobediploid.Ontheotherhand,thecrosswith hybrid with the tetraploid lc. maculatum subsp. obtusiusculum (the tetraploid) lc. subsp. obtusiusculum, described by Frb'hlich (5xa. //. x desetangsii nothosubsp. desetangsii) is also usually (1911) and named here H. x laschii nothoformafroelichii, should tetraploidandhasbeenconsideredabove.Thatwiththediploid la. provetobetriploid. maculatumsubsp.immaculatum(5xc.//. xdesetangsiinothosubsp. ThehybridsinvolvingH.perforatumcanbetriploidtohexaploid, balcanicum) and Ib. maculatum subsp. maculatum (5xb. H. x STUDIES INHYPERICUM 65 desetangsiinothosubsp.carinthiacum)occursintwoformsdepend- AlthoughaPortuguese specimenofH. undulatum wasobservedto ingonthe sex ofthegametes. WithH. maculatumpollen, triploids have a habit and narrow leaves that approach those of H. (2n=24)areproduced(e.g.5xbanothoformamaculatiforme)whereas triquetrifolium, these characters would seem best interpreted as a with perforation pollen, the progeny are pentaploid (2n=40) (e.g. parallel development within H. undulatum. The first division in 5xbbnothoformaperforatiforme). Thecorrespondinghybridswith group(iv)wouldthenhavebeenbetweenthewesternMediterrean/ 1a.H.maculatumsubsp.immaculatumhavenotyetbeenrecognized Atlantic wet-habitat H. undulatum and the east Mediterranean/W. ornamed. AsiandryhabitatH. triquetrifolium.Thespecies3.H. tetrapterum, with broad stem lines, broaderplane leaves and a more congested Distribution andevolution (Fig. 5) inflorescenceofsmallerflowerssubsequentlyevolved(inthesouth- westernMediterranean?)fromH. undulatumandspreadnorthasfar As has already been explained (p. 61), sect. Hypericum is most as Scotland, Denmark and southern Sweden, north-eastto Poland, closelyrelatedtoH.ascyronsubsp.ascyron(Sect.7.Roscyna),and Latviaandthewestern Ukraine, andeastto Iran. subsect. Hypericum is distributedovermostofthe Paleotemperate Group (ii) divides immediately into an eastern, Old World sub- Region from the Azores to Mexico. The distribution of series group (v) and a western, West American subgroup (vi), both of Hypericum occupies most ofthis area except south-eastern China which start with a single species, respectively the eastern Siberian andsouthJapan.Itisdividedintotwogroups(i)mainlywesternand andChinese6.H. attenuatumandtheBritishColumbiatoMexican (ii) mainlyeastern. 12. H. scouleri. Whilst none of the variation in H. scouleri has Group (i), which extends from the Azores to central Siberia, resulted in speciation, H. attenuatum has speciated eastward and includes the nearest taxon morphologically to series 2, la. H. westward. Tothe west itgave riseto7. H. elegans, which inhabits maculatum subsp. immaculatum, which is restricted to the central steppe regions as far intoEurope as south Germany (Saxony, with Balkan region but differs little from the north Japanese H. isolatedstations furtherwest). Eastwardthere isagroupofspecies kamtschaticum var. pibairense Miyabe & Y. Kimura (series 2), inJapanofwhichthetwosouthern,8.H. tosaense(Shikoku)and9. mainlyinhavingconstantly4-linedgland-bearingsteminternodes. H. momoseanum (C. Honshu)haveatonetimebeen includedinH. Subsp. immaculatum appears to have given rise northward (group attenuatum. They both occur in isolated localities, as does aclose iii) to two subspecies: (a) by vicariance to Ib. subsp. maculatum relativeofH. tosaense,9.H. iwatelittorale,innorthernHonshu.On (Atlantic coast tocentral Siberia) and (b) by autotetraploidy to Ic. the other hand, the northern relative of H. momoseanum, 11. H. subsp. obtusiusculum (=H. dubium Leers) (north-west Europe). yezoense, is widely spread in Hokkaido and northern Honshu and Southwardthereisaderivative(groupiv)thatdividedintoawestern alsooccurs insouthern Sakhalin. (damphabitat)subgroup(Spp.2-3)andaneastern(dryhabitat)one Aswehaveseen,5.H.perforatumhasapparentlyresultedfroma (Sp. 4). In group (iii) the stems are 4-lined, the leaves broad and crossbetweenonetaxonfromgroup(iii)(la.H. maculatumsubsp. planeandthesepalsbroadandrounded(entireornot);ingroup(iv) immaculatum) and one from group (v) (6. H. attenuatum). It was the stem lines become broad (3. H. tetrapterum) orthe subsidiary suggestedthatthecrossoccurredinwesternSiberia,andthepresent pairdisappearaltogether(4. H. triquetrifolium),the leavesbecome distributionandvariationof//,perforatumsupportsthissuggestion. undulate (2a. H. undulatum var. undulatum) and narrow (4. H. Both parents have sessile herbaceous leaves, and the firstdivision triquetrifolium)orremainbroadbutreverttoplane(2b.H.undulatum (vii)isbetween5a.subsp.perforatum, withshortlypetiolateherba- var. boeticum, 3. H. tetrapterum), whilstthesepalsbecome narrow ceous concolorous leaves and 5b. subsp. songaricum, with sessile, and acutebut, inH. triquetrifolium, revert toroundedwhen small. oftensubcoriaceousanddiscolorousleaves. Subsp.perforatumhas 11 Japan(N.Honshu,Hokkaido) Russia(S. Sakhalin,S. Kuriles) 3c EuropeexAcllg.eNr.i,aW.Asia SWEInbgelriaannd,PeSniWn.,WaNleWs,AfSrWicaIreland? W.Canada,12W.USA 10 Lebanon .&C. Mexico Japan(C.Honshu) Levant (v)\ 6 E. Siberia,Mongolia Japan(N. Honshu) E.Med.to Korea,China NW Iran Ib Japan(Shikoku) & NW N. C.Europe Europe SiberiaE.toAngara-Sayan E.toW. Germany,Bohemia W. Siberia Styria&Hungary toW. Europe 5b &NW N.Kazakhstan Xinjiang S.&W.toS.\Ukraine N.&C.EuropeS.toN. Spain,N. Italy,NNEWGreece,NWTurkey SiberiaE.toAngara-Sayan, Mongolia 5c & Macaronesia,Medit. S. Europe 5d ' toE. Kazakhstan&NWIndia China Sudan(JebelMarra),SaudiArabia(Asir) Fig.5 Section9,subsect. 1,series 1.Distribution. 66 N.K.B. ROBSON spreadwestwardthroughoutnorthernandnorth-westernEuropeand persistent. Flowersstellate,homostylous.Sepals5,freeorunitedat eastwardintocentralSiberia.BeyondSiberiaitrecurs,morphologi- base, persistent, erect to recurved in fruit, with margin entire or cally scarcely altered, in Chinato the south-eastofthe Mongolian denticulatetoglandular-ciliate; veins 3-5(-7); laminarglandspale desert and steppe region and is transformed gradually south-east- and/or dark, linear to punctiform; marginal and/or intramarginal ward intoanarrow-leaved form with long branches andcongested glands dark and/or very rarely pale or rarely absent. Petals 5, inflorescences (subsp. chinense). Subsp. songaricum is present in persistent, erectbut nottwisting afterflowering, withoutapiculus; themountainsofnorth-easternKazakhstanandadjacentKyrghizstan margin entire to crenate or eroded-denticulate to ciliate; laminar andXinjiang(China);then,afteragapofover2500km,itrecursin glandslineartopunctiform,paleand/ordark;marginalglandsdark, southernRussia(Astrakhan,Ciscaucasus)andKrym.There,witha immersedto prominent,orabsent.Stamenfascicles'3' (i.e.united smalltransitionalzone,itisreplacedby5c.subsp.veronense,which 2+2+1), persistent, with stamens totalling 20-c. 100; filaments appears to have spread through Turkey westward into southern basallyunited;antherglanddarkorrarelyamber;pollentypesIV?, Europe,theMediterraneanandMacaronesia,southtoSaudiArabia X. Ovarywith 3(4) completely oralmostcompletely axileplacen- and the Sudan Republic (Jebel Marra - if truly native there, see tae, each oo-ovulate; styles 3(4), divergent from discrete bases; Wickens, 1976a, \916b: 98),andeasttothewesternHimalayaand stigma narrowly capitate or narrow. Capsule 3(4)-valved, Central Asia. chartaceous,withvalveslongitudinallyvittateorsometimes later- Distributionsofspecies and species groups in series Hypericum ally vitiate to vesiculate, pale. Seeds cylindric, not carinate or showfewwidedisjunctions,butthosethatdooccurareremarkable. appendiculate; testascalariform-reticulatetofinelyfoveolate. Tvahreiawtiiodnes(ti.ei.spirnimgirtoiuvpe f(ioi)r,m)whoefreH.thaetteeansutaetrnumSiisbenrioawn sceepnatrraeteodf BASICCHROMOSOMENUMBER (x). 8; ploidy2,4(rarely 3, 5, 6). from thatofH. scouleri in Arizona and New Mexico by the north HABITAT. Various;0-3150m. baPAmlaymoscerioHrefuliiiicrc-kelaOecilceyndeniattLsnoijt.ush(nuTa1cphn9tpi5iotso2rhn)dtosistsheahtirarsteiinobpptvulrhotaeielnodrvtnoisinmwnwgiofineuetarlahensdntteclseaeyrse,ntehtemeNhrarnottbroAatssschuuiecpoaAhpumo-sder;irtwsieajtcshunatend.ecirtintHinfoyieNnsprsoeetrrnhaticur-hese DnnnoooIrrrStttThhhR---IwwwBeeeUsssTtttIMOAIofNnrdn.iigcaoal,M(iUaastc,otaaurrKtoohnPr-erewsaaedi,seats,ChhA)isE,niuaarceon((ptemerxeacsle(opepAtxhscyitetahp,ietcsseooruxuetttghhir-oeeenramsnse)t,SneioaTbrsiettbrhei)tat,,,o cCaulmifosrcnoiualenrfiloirsa(aAxmeelmrobde,r19o5f9;thReavArecnto&-TAexretliraordy,e1l9e7m8e)n,twhoefrethiet Qnoirntghh-awiesatndtoEc.enXtirnajliManegx)i,coJ.apRaenp,rweessetnetrendCinanmaadnay,owtehsetrepranrtUs.oSf.At.h,e & worldby H.perforatum asan introduction. oLcacwusrosn)infotrheestYseolflotwhePSiineerr(aPNineuvsadpaon(dMeuronsza&DoKuegclk.,e1x95P9.). ItCi.s 42 species in 2 subsections, one with 2 series (for infraspecific possible, therefore, that H. scouleri has been in western North taxaandhybrids, see subsectionaldescriptions). Americasincethe Miocene. The other notable disjunction has already been mentioned. It Key to subsections and series ofsect. 9.Hypericum would appear that H. maculatum subsp. immaculatum, now con- finedtothecentral Balkans,oncehadadistributionextending into 1 Rootstockherbaceous, stoloniferous; stemspersistently 2^4-lined western(orevencentral)Siberia,whereitcrossedwithH.attenuatum or-winged,withglandsonthelinesandoccasionallyelsewhereor and gave rise to H. perforatum. Its close resemblance to H. kamt- eventuallytereteandeglandular;blacklaminarleafglandsfewor schaticum var. pibairense, of northern Japan, implies that subsp. none (sometimes numerous in Sp. 6); capsule valves sometimes immaculatum(oritsimmediateancestor)migratedsouth-westward withlateralobliquevesicles(subsect. 1.Hypericum) 2 (without differentiating?) and eastward with minimal differentia- Rootstockwoodyorfibrous,notstoloniferous;stemsalwayseven- tion.Ithassofarnotbeenpossibletoestimatethegeologicalageof tuallyterete,eglandular(sometimesexceptSp. 39);blacklaminar thesedisjunctions. leafglands usually present; capsule valves always longitudinally vittate subsect.2.Erecta 2 Stems persistently 2^4-lined, lines glandiferous, at leastone pair SYSTEMATICTREATMENT1 prominent(sometimesweakandeglandularinSp. 12) series 1.Hypericum Sect. 9. HYPERICUM Stemseventuallyterete(exceptSp. 13),eglandular,linesweak... m ...series2.Senanensia Perennialherbsorveryrarelysuffruticesupto 1.2 tall,withstems erraerecltytroedpdriosshtr)agtle,andgsl;abbrroaunsc,hiwnitghloautteraolr,fwriotmhudpaprekr(obrlmacokstornovdeesr.y Subsect. 1. Hypericum. Type: H.perforatumL. Stems4-2-linedor-wingedwhenyoung,remainingsoorbecoming Stem internodes persistently 2^1-lined or narrowly 4-winged, the terete,eglandularorwithglandsconfinedtothelinesornot.Leaves lines and sometimes elsewhere black- or rarely reddish-gland- opposite or abnormally 3-whorled, decussate, sessile to shortly dotted (very rarely terete and often eglandularin 12. H. scouleri). pseudopetiolate, free, persistent; lamina entire with venation pin- Leaveswithlaminarglandsallpaleorafew(sometimesmorein6. nate, closed, with tertiary reticulation dense to lax and obscure; H. attenuatum)black,punctiform,orabsent.Bractsandbracteoles laminar glands punctiform or rarely shortly striiform, equal or entire.Petalswithlaminarglandspaletoblack,lineartopunctiform, unequal; intramarginalglandsblackoroccasionallyalsopale;ven- orabsent; marginal glands immersed orsessile. Capsule valves (i) tral glands absent. Inflorescence \-c. 70-flowered with branching longitudinally vittate or with (ii) dorsal and lateral vittae or (iii) dichasial-monochasial from 1^ nodes, often with subsidiary vesicles with or without dorsal vittae. Seeds linear-foveolate to branchesfromlowernodes;bractsandbracteolesfoliarorreduced, finely foveolate. Species 1-19. 'Anasterisk(*)beforealocalityorafteraherbariumspecimensymbolindicatesthatthespecimenhasnotbeenseenbyme.Typematerialseenbymeisindicatedbyanexclamation mark(!). STUDIES INHYPERICUM 67 Series 1. Hypericum 7(5) Inflorescence 10-c.70-floweredfrom 1^nodes;stemserect 3a.tetrapterumvar.tetrapterum Hypericumsect.Euhypericumsubsect.HomotaeniumseriesAttenu- ata, series Kamtschatica pro parte quoad H. yezoense, series Inflorescence 1-3-floweredfromterminalnode;stemsprocumbent Acuta, series Quadrangula, series Elegantia, series Crispa toascending 8 Gorschk. in Shishkin & Bobrov, Fl URSS 15: 236-244 (1949), 8(7) Leaveselliptictoorbicular,mostly0.5mmpetiolate,marginplane; nom. invalid, omniasinedescr. lat. inflorescencewithoutfloweringbranches Hypericum sect. Euhypericum subsect. Heterotaenium series Per- 3b.tetrapterumvar.corsicum /noorma.toiGnvoarlsicd,hks.iniendSehsicsrh.kliant.&Bobrov,Fl. URSS15: 247(1949), Lmaeragviensuenlldiupltaitcet;oitnrfilaonrgeuslcaern-coevastoe,mestuibmseesssiwliethtofl0o.w5emrimngpbertaionlcahtees, Hypericum sect. Homotaenium series Quadrangula Y. Kimura in fromupto5nodesbelow Nakai & Honda, Nova Fl. Jap. 10:136 (1951). Type: H. 3c.tetrapterumvar.anagallidifolium Hypqeuraidcruamngsuelctu.mHLo.,monotma.enrieujimciseenrideusmC(r=iHs.pateYt.raKpitmeurruam,Frl.o)c.. cit. 9(1) Lsteeamfmwairdegliynbcrriasnpcehde-dunfdourlmaotes,tloafmiitsnlaeunsgutha,llfyotrrmiiannggublraora-ldapnyceroalmaitde;; (1951). Type: H. crispum L. (=H. triquetrifoliumTurra). sepals 1-3mmlong 4.triquetrifolium Hypericumsect.HomotaeniumseriesBilineata Y. Kimura, loc. cit. (1951). Type: //. attenuatum Fisch. exChoisy. Leafmarginplane,laminaovateoroblongorelliptictolinear;stem narrowly branched rarely below middle, forming at most narrow Stems persistently 2^-lined, lines glandiferous, at least one pair pyramid;sepals(2.5)3mmormorelong 10 prominent(sometimes weakandeglandularin Sp. 12). Spp. 1-12. 10(9) Stem lines (at least) with black glands, principal lines always Key to sect. 9.Hypericum subsect. 1.Hypericum present;petalmarginusuallydistallyblack-glandular-crenate 11 series 1.Hypericum Stem lines eglandular or very rarely with a few reddish glands, 1 Stem internodes completely 4-lined or4-winged; capsule valves dpirsitnaclilpyalbllaicneks-grlaarenldyulaabrs-esnutb(cir.ee.nraatreelyterete);petalmar1g2i.nsecnotuilreertio longitudinally vitiate (rarely interrupted to vesiculate in Sp. 4) 2 11(10) Subsidiary stem lines nearly always developed; sepals often dSitneamlliynvtietrinaotedetsopvaerstiicaulllyat4e-linedor2-lined;capsulevalveslongitu9- epm.geld1ai0na1dt)uelabre-dtewneteincul1.atHe.;moatcheurlacthaurmacatnedrs5.toH.vapreyri5fnxo.graxdtiedogenrsee(etssaeneignksteieyri,- 2(1) eSreopdaelds-duennetqiucaull,ataep;exleraovuensdweidthtovoebntautsieonorsduebnasceulmyinraettei,cuelnattiereantdo Sinutbesrimdeidairaytestemlinesabsent;sepalsentire;othercharactersnot1s2o laminarglandsabsentorscattered,relativelylarge (1.maculatum) 3 12(11) Capsulevalveswithlateralvittaeorvesiclesandoftendorsalvittae (5.perforatum) 13 Sepals subequal toequal, apex acute to acuminate, entire; leaves with venation laxly reticulate; laminar glands always present, Capsulevalveswithlongitudinalvittaeonly 16 varyinginsize 5 13(12) Leaves(atleastonmainstem)sessile;petallaminarglandsusually 3(2) Stem internodes always with complete subsidiary lines; inflores- allpale 14 caepnecxeenbtriarne;chpeestamlsakenitnirge,nawrirthowlaamnignlaer(cg.la3n0d)swbiltahcksatenmd;dsoetpsatlosswhiotrht Leavesallpetiolate;petallaminarglandspaleand/orblack 15 streaksorlinearandpale 4 14(13) Capsulevalveswithlateralvittaelinearorslightlyswollenoverall, Stem internodes with complete or incomplete subsidiary lines; notinterrupted; sepalsfinelyacuminate; leavesoblongtooblong- inflorescence branches making a broad angle (c. 50) with stem; ovate,base shallowlycordate-amplexicaul sepals with apex usually finely eroded-denticulate; petals usually 5b.perforatumsubsp.songaricum distallyunilaterallycrenate,withlaminarglandsblack,mainlylines Capsulevalveswithlateralvittaeswollenatbaseand/orinterrupted tostreaks Ic.maculatumsubsp.obtusiusculum toshortandirregular(vesicular); sepalsacute; leavesusuallynar- 4(3) Sepalsandpetalswithonlyplaal.emgalacnudls;atpeutmalsgulbasnpd.silmimneasctuolsatrteuakms 5ro-w1l0ymtmrialnognugl)a,r-blaasnecerooluantedetdotloinceuanreaotr,eifbroader, then short (c. 5c.perforatumsubsp.veronense Sepalsandpetalswith mostly blackglands; petalglandsdotsand shortstreaks Ib.maculatumsubsp.maculatum 15(13) Inflorescence and/orpartial inflorescences notusuallycongested, branchesrelativelyshort,straight;leavesusuallyoblongtoovateor 5(2) Steminternodes equallynarrowly4-winged;leafmmarmginusually elliptic;atleastsomepetallaminarglandspale crisped-undulate; inflorescence lax; flowers 12-17 in diam.; 5a.perforatumsubsp.perforatum sepalsnearlyalwayswith3-28laminarblackglands;petalsbright yellow,nearlyalwaysred-tingeddorsally(2.undulatum) 6 Inflorescence and/or partial inflorescences congested, branches relativelylong,curved-ascending;petallaminarglandsallblackor Steminternodes broadly4-winged,theprincipallineswider;leaf absent 5d.perforatumsubsp.chinense marginnearlyalwaysplane;wholeinflorescenceorpartialinflores- cencesdense;flowers 10-15mmindiam.;sepalswithoutorrarely 16(12) Sepalsblack-glandular-ciliateorwithsessileblackmarginalglands with1-2laminarglands;petalsratherpaleyellow,notred-tinged(3. 7.elegans tetrapterum) 7 Sepals entire or with apical black gland only, without or with 6(5) Petalstingedreddorsally;leaveselliptictoovate,margincrisped- immersedblackmarginalglands 17 undulate;planterect 2a.undulatumvar.undulatum 17(16) Sepals andpetals with pale and black laminarglands; stem often Petals not tinged red; leaves broadly elliptic to orbicular, margin withscatteredblackorreddishglands;sepalsacutetosubacuminate entire;planterecttodecumbent ....2b.undulatumvar.boeticum ....6.attenuatum 68 N.K.B. ROBSON mm Sepals and petals with pale laminarglands only; stem with black elliptic, entire. Flowers (15-)20-25(-35) in diam., stellate to glandsconfinedtoraisedlinesornearthem;sepalsacutetoabruptly reflexed; buds broadly ellipsoid, rounded to obtuse. Sepals 5, acuminate 18 unequal, 3.5^.5(-5) x 2-2.7(-3) mm, broadly elliptic to broadly 18(17) Leaveswithoutblacklaminarglands;innerpetal margincrenulate ovate or broadly oblong, rounded to obtuse or subacuminate (unknown in9. iwatelittorale); stems usually erect, ratherwoody (varying in same flower), entire to eroded-denticulate, erect to 19 recurved in bud, recurved in fruit; veins 5(-7), not or slightly Leavesusuallywithafewblacklaminarlands;innerpetalmargin branched;laminarglandspaleandsometimesalsoblack,punctiform plane;stemsusually ascending,herbaceous 20 and sometimes striiform; intramarginal glandspaleoroccasionally blackoroftenabsent.Petals5,goldenyellow,nottingedredinbud, 19(18) Leaves all sessile, mosmtmly elliptic or oblong-ellmipmtic to narrowly (8-)10-15 x4-7.5 mm,c. 3 x sepals,broadlyelliptictoobovateor oblong:flowers15-18 indiam.;sepals2.5-3 long,gradually more rarely symmetrically oblanceolate, distally crenate orentire; toabruptlyacuminate,intramarginalglandsalwayspres8e.ntt,ossapeancseed laminarglandspaleand/orblack,punctiformandsometimesproxi- mally striiformto linear(in subsp. immaculatumalmostall linear); Lfsuleboaawvceeursmsion9na-t1ela0.teirmnatlmrabmraianrngcidhnieaaslm.gp;leatinsodelspualflaetswe,3o-rel4albipsmteinmct9t.oliownoagbt,lealnaiccteutotolreaatlete;o gi1nl0ta0r)n,admba'3lrag'c-ikfn.aaslcOivgclalaerndy,ds3l-oflenowgc,eulsdatirs,7t-a2l1-0oArxmums1u,.a5l-cl.2y.05a.bm6s5em-n,t0..9Sbtxraompaeedtnlasyls5o;0va-oni8td0h(et-ro ovoid-ellipsoid; styles 3, free, 3-4 mm, l-1.5(-2) x ovary, rather 20(18) Stemssingle.0.25-0.65m,erectorbasallyascending;inflorescence narrowlyspreading;stigmasnarrowlycapitate. Capsule6-10x4-6 7be-l1o1w-floweredfrom2-3nodes,usuallywith1-4f10l.owmeorimnogsberaanncuhems mlomn,gict.u2di-n2a.l5vxitsteapea,llsi,neoavroitdo-setlrliiifposromi.dSteoedbsrdoaardklbyroovwoni,d;0.v8a-lv1e.s2mwimt,h Stemsusuallycaespitose,0.1-0.35m,erecttoprocumbent;inflores- cylindric,notcarinateorappendiculate;testafinelylinear-foveolate. cence 3-7(-9)-flowered from 1-2 nodes, with 0-1 flowering 2n=16,32(seeundersubspecies). branchesbelow 11.yezoense In usually rather damp habitats - woods, scrub, grassland, moor- 1. Hypericum maculatumCrantz, Stirp. austr. 2: 64 (1763), 2nd land, meadows and ditches; (W. Scotland) - 2650 m (SE ed. 2: 98 (1769); Allioni, Fl. pedem. 2: 45 (1785), 3: t. 83 f. 1 Switzerland). (1785)proparte,excl.syn.H.androsaemifoliumVill.;Schinz& Europe from Scotland (Ross-shire), northern Scandinavia (to KKaeillseerr,l.FlA.kSacdh.weiWizs,s.3,rdMeadt.,h.1-:N3a5t8ur(w1i9s0s9).;KAl..F1r2o0h(l1.)m:Si5t3z8un(g1s9b1e1r)., HweisntneorynaS(ibLeorfioat,ensoIust.)h,t6o8n2o2r'thNe)r,n nSopratihne,rnnoErtuhreorpneaItnalRy,usnsoirathaenrdn in Oesterr. hot. Z. 63: 18(1913); Thellung inAllg. Bot. Z. Syst. Greece and the central Ukraine, and east to south-western Siberia 18: 19(1912); Hayek,Prodr. Fl. Pen. bale. 1: 534(1925); Hegi, (R. Ob) with eastern outlier near Krasnoyarsk). Introduced into ///. Fl. Mitt.-Eur. 5(1): 517 pro parte, excl. f. 2004 (1925); N. Canada(BritishColumbia). Robson inBot. Soc. Brit. IslesProc. 2: 237 (1957), inop. cit. 3: 99(1958), inDavis,Fl. Turkey2: 400(1967)inadnot., inTutin Hypericummaculatum isthe mostprimitivespecies inthe(mainly & etal..Fl. Europ.2:268(1968);Zelenyetal.inFutak Bertova, European) series Hypericum, its nearest relative in the (eastern) Fl. Slovenska3: 302, t. 37 f. 1 (1982); A. Ramos in Trab. Dept. series Senanensia (Spp. 13-19) being 13. H. kamtschaticum var. Bot. Univ. Complut. Madrid12: 49,t. 3 f. 1 (1983), inActaBot. pibairense from Hokkaido (N. Japan). It comprises two diploid Malac. 11: 167, f. 6e(1986), inCastroviejoetal., Fl. Iberica3: subspecies(subspp. maculatumandimmaculatum)andatetraploid 168 (1989); Hagemann in Flora 173: 112-115, tt. 12-14, 39 subspecies (subsp. obtusiusculum) that, from chromosome mor- (1983);N.Robson&StridinStrid,MmFl.Greece1:607(1986); phologyandsynthesisbycolchicinedoublingofsubsp.maculatum N. Robson in Cullen et al., Europ. Gdn Fl. 4: 60 (1995); in (Robson, 1958a), would appeartobe autotetraploid. The latterhas Wisskirchen & Haeupler, Standardliste Farn-u. Bliitenpfl. sometimesbeen treated as aspecies (H. dubium Leers); but (i) the Deutschl.:269(1998).Type:Austria, 'alpen\ 1760(fl),Breynin autotetraploidyand(ii)theoccasionaldifficultyinassigningaplant Crantz 828 (BPI-lectotype, N. Robson, selectedhere). tooneorothertaxonindicatethatsubspeciesisthemostappropriate Fig. 6, Maps 1, 2. rank,paceMartonfietal. (1999) (seep. 75). Prioenforlteoinrnnegisabcalesnhece,er,wbilta0th.e2sr-tb0er.ma6snmcnhuetmadlelra,obeourevscettofoofrreewa.,s2ci/en3intdoiiaflntlghyefuinrrbolrmenacgntrcheh.eepSditnbegemlsaon4wd- actoinTofhniseneoddvieptrollotiahdpepsisunobgustphiencBiadeilssktaronifsbuHta.inodnm.alacOcunklseabtl(usamucbkscpgo.lmapnirdmismsaeocntutlwhaoetppueomtp)aullsi-,s lined, the subsidiary ones (decurrent between leaves) often less thepaleonesbeinglinearandtheleavesusuallypale-gland-dotted. plsbieernsnoesemsia;liteni;hen,ltnaetcmrhoniaronrdaetevas1ce5en1o^0pu-0as5r;tx0iaam1pl0mle-y,x2a0srbhsomoeurmnntt,de2er,bdtur,hsoauamnadalltrlyogyetiwoxnicnteaphelradbrniloenaw,gclkybleagealslvleaienpsrtd.ioscLu,oenpnadavetledhe;ser coIwtabisltdlcueyshsirpiutrosemciasoudlsvuaoemnmr.deyTsnlhsiuiegmmhiobtltleahyrreorhvtaeposrolpsnauuoplbtsastypiie.ntoanmbreae(acesuuntbhlscaapott.uounfmmta,secudbu,lslepbas.usttuismmmo)om,ratpcowhuholsiluacobthgsuipim-.s venation: 3(2) pairs ofmain laterals from lowerquarterto base of inazonefromBosniatosouthern Bulgaria, haspetalswithpuncti- midrib, tertiary reticulation dense; laminar glands absent or rarely formtoshortlystriiformblackglands,thepale(punctiform)glands pale,scattered,sometimesalsoblack,few,punctiform;intramarginal being few, andthe leaves usually lackpaleglanddots. glands black, close, irregular in size. Inflorescence (2-)3-c. 15- flowered, from 1-3 nodes, with flowering branches narrowly or 1a. Hypericummaculatumsubsp.immaculatum(Murb.)A.Frohl. curvedascendingfromupto5nodes,thewholecylindrictobroadly mSitzungsber.Kaiserl.Akad. Wiss.,Math.-Naturwiss.Kl.120(1): pyramidal or subcorymbiform; pedicels 1.5-4 mm; bracts and 547, f. 8 (1911), in Oesterr. Bot. Z. 63: 18 (1913), in Mitt. mm bracteoles upto 5 long, narrowly triangular-ovate to narrowly Naturwiss. VereinesSteiermark51: 223 (1915); Hayek, Prodr. :Partialorcompleteabsenceofsubsidiarylinesmaypossiblyindicatehybridi/.ationwithH.perforation(cf.H. xdesetangsii,p.75).

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