Bulletin of the Natural History Museum

100 Pages·2002·8.4 MB·English

by  Natural History Museum (London)

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
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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).