Biosystematic Studies on the Family Tofieldiaceae IV. Taxonomy of Tofieldia coccinea in Japan and Korea Including a New Variety

2013·5.9 MB·English

by  Tamura, Minoru N

TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
ISSN1346-7565 ActaPhytotax,Geobot,64(1)2:9-40 (2013)

BiosystematicStudieson the FamilyTofieldiaceaIeVLTaxonomy of

7bLfield cioaccinea in Japan and Korea Including a New Variety
MINoRu N TAMuRAi･', NAM SooK LEE2, TERuo KATsuyAMA3 AND SHIzuKA FusE4
' Dqpartment ofBota aGyracinate School of'Scie Inkc5e,o. tCoh7ivemsi KOiltashirakawa-oiwake-ehe,
Sadyo-ku ,1<),ot 6o06-8502, Jlrpan ."[email protected] (inuthorforcorrespondenceJ;
2 Department qfLij2 iScience E,wha PK]nian Ushivensi4 5J2l Ewhayeoclae-gil ,Seoclaemun-g uS,eoul I20-Z50,
Kbrea; 3 Kbnagawa PrEll?ictu" Mailtseum of?Vbtu Hrisatlot :4z99 Ilyudo ,O(lawara-sh iKa,ncrgawa 250-O031,
Japan; ` MUseum ofAfat uandre Human Activitie sI.lyogo, 6 1?DJoigaoka ,Sando-shi, 11lyog o669-I546, Japan
Te bette runderstand variation in 7ofetdia coccinea in Japan and Korea, we compared morphological
finding wsith molecular phylogenet iancalyses using regions of the plastid strnK, trnL and trnL-F plus
the nuclear ITS region of le species and 19 samples of 7ofeldia, including 10 sampres of T eoccinea.
Based on morphology, we recognized seven varieties of Z coccinea, var. coccinea, var. kondoi ,var.
gracilis ,var. kiusian av,ar, geibiensis ,var, akkana and a new variety, var, diboiry aw,ith a panicle (di-
botryum) .According to the molecular phylogeny T, eoccinea vars. coecinea, kondoi, gracili sand ak-
kana formed aclade with 57-62% bootstra spupport; var, kiusiana and var. dibotrya were excluded from
the clade. The topology supports our recognition Qf 7r eoccinea vars. kiusian aand dibotry aas distinct
from var. gracilis ,which is morphologically most similar, The molecular phylogeny also supports the
inclusion of T coccinea var..tZiuriei within var, kondoi. The name T yoshiiana var. koreana (comb n.ov.)
should be used instead ofT yoshiiana var, kanu,onensis.
Key words: lectotyp emo,lecular phylogeny, new combination, new variety, panicle ,taxonomy,
7Qr7eidi acoccinea, 7tlfietd icoaccinea var. dibotrv a7,byleldiayoshiian avar. koreana, Tbfieldiaceae
Tofeldia Huds, (Alismatales-[IbfieldiSaacve, )aOehw;i, var. kiusiana (Okuyam aOjhwi and
Tamura et aL 2004, Chase et al. 2006) comprises var, kondoi (Miyab e& Kudo) H, Hara. Hulten
12 species (1famu reta al. 2010, 2011) ,among (1943 h)o,wever ,considered T nutans in Asia
which Z coccinea Richardson and T pusill a(Mi- (typ leocalit Eya:st Siberia t)o be conspecific with
chx,) Pers, are widely distribute din circumboreal Z coccinea in NoTth America (ty pleoca]it Cya:n-
and circumarctic regions, while the other 10 spe- adaj, Hara (196 1ag)reed with Hulten ,recogniz-
cies are more or les slecalize dT.he distribution ing fiy evarieties in Japan under T coccinea: var,
range of Z coccinea extends south int otemperate coccinea, var. fcts c(aMiyab &e Kudo) H. Hara,
regions in Japan and Korea, where several variet- var. geihiensi s(M. Kikuchi) H, Hara, var, kiusi-
ie sof T eoccinea are differentia tpreidmarily by ana (Okuyama jH, Hara and var, kondoi (Miyabe
plant size and fiowe r! frui dtensity .In this paper, & Kudo) H. Hara. Hara (196 1pr)oposed the inclu-
we focus on the varieties of Z coccinea in Japan sion ofvar, gracili swithin var. kondbi.
and Korea. Kitamura (1964 )recognized Tofeldia coc-
In Japan ,Toj7eld icoaccinea was firs tidenti- cinea var. akkana (T .Shimizu) T, Shimizu in ad-
fie das T nutans Willd. ex Schult .& Schult ,f ditio nto vars. coceinea, geibiensis k,iusian aand
(Maximowi c18z67), Ohwi (195 3re)cognized five kondoi (= T gracili si)n Japan, without recogniz-
varieties under Z nutans: var. nutans, var. fasca ing T coccinea var. .fusca .The possibili otfy this
(Miya b&e Kudo) Ohwi, var, gracili s(Franc &h. nonrecognition of var. jusc ahad been implie dby
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
30 ActaPhytotax.Geobot, VoL 64

Kawano (196 1an)d Hara (1961 S)a.take (1982)de (190 4an)d from Sakhali n(a sT nutans) by Mi-
fbllowe dKitamura's (196 4t)reatment, but con- yabe and Kudo (1914 I)t .was reported from Us-
trary to the then current opinion, Shimizu (1983) suri (a sT nutans) by Kuzeneva (1935 b)u,t not by
distinguish eadgain T coccinea var. gracilis Regel (1861 O)n.ly var. coccinea has been report-
(Fran &c hS.av.) T, Shimizu from T coccinea var. ed from those regions.
kondoi. In this study, we based eur firs ctonclusions
Yamazaki (200 2a)lso recognized Toj7eldia on the taxonomy of Tofeidi acoccinea in Japan
coccinea var. gracili sin addition to var. coccinea and Korea on morphological observations, then
and var. kondoi in Japan, while merging Z coc- tested our finding sby molecular phylogenetic
cinea var. akkana, var. geibiens iasnd var, kiusi- analyses. In the course of the study, we discov-
ana with T coccinea var. gracili sC.ontrary to ered a new variety of T coccinea from Tanzawa,
previous opinions (Matsumu r1a905 ,Miyabe and Kanagawa Pref, Japan, with a panicle (di-
Kudo 1914, Ohwi 1953, Shimizu 1958 ,1983 ,Hara botryum) ,which we describ eas Z coccinea var.
1961, Kitamura 1964, Satake 1982) that var, coc- dibotrya,
(= isdistributed
cinea var. nutans) northward
from Honshu (alpi nzoene), Ylimazaki (2002) Materials and Methods
confined the distributio nto regions northward
from central Hokkaido. Thus, the delimitati oofn Morphologicalobservations
varieties of Z coccinea in Japan has been quite Specimens in the herbari aEWH, HYO, KPM,
confused. KYO, MAK, OSA, SAPS, TI and TNS were ex-
In Korea, Tofeldia .faurie iH. Lev. & Vaniot amined to circumscribe precise ltyhe varieties of
and T taquetii H. Lev. & Vaniot were described Tofeldi acoccinea in Japan and Korea and to de-
from Jejudo Islan d(Levei l19l0e8) and were sub- termine whether plants of Tofeldi afrom [Rinza-
sequently recognized by Nakai (191 1I)n .1914, wa with panicle sand distin cltater ablranches
however ,Nakai reduced T taqueti ite T.fauri eiin were taxonomically distinc tW.e also created a
a report on plants from Jejudo and Wando Is- key to the varieties of TQfield icoaccinea in Japan
lands ,In 1916, Nakai liste dToj7eldi anutans in a and Korea based on herbarium specimens,
report on plant sfrom Rhobong, Pyeonganbuk-
do. Chung (195 7re)cognized T fouri e(i- T ta- Molecularpbylogenet iancalyses
quetii )and Z nutans in Korea. DNA sequences of plasti dtrnK (including
Hara (196 1fu)rthe rreduced Zfauriei (= T ta- matK), trnL and trnL-F regions and the nuclear
quetii )to T coccinea var. konclo ia,nd recognized intern atlranscribed spacer (ITS r)egion of six va-
two varieties under T coccinea in Korea, coc- rieties and seven samples of Ttlfield icaoccinea
cinea (= Z= nutans) and kondbi .W. T, Lee (1996)determine din this study have been deposite din
accepted Hara's taxonomy, but Y. N. Lee (1996)the DNA Data Bank ofJapan (DDBJ )(Tab l1e).
recognized T fauri ieni addition te T coccinea. These new sequence data were subsequently in-
Yamazaki (200 2c)onsidered the differenc ebe- cluded in the matrix of trnK, trnL, trnL-F and
tween Z coceinea and Tfourieito be at the infra- ITS sequences of 10 Tofeldi aspecies (including
specific level and proposed the name T coccinea 2 varieties and 3 samples from T eoceinea) gen-
var,fouriei (H .Lev. & VanioO T. Ytamaz. and also erated by Tamura et aL (201 02,011) (Tab l1e).
recognized anew variety T coccinea var. kanwo- Among the 10 species, T calyculata Wahlb., T
nensis T. Yamaz, Variety kanwonensis, however, glabra Nutt. ,T okztbot Makino and T pusilla
was shown by [I}tmu reat al. (201 1to) belong to T were used as an outgroup because the remaining
yoshiiana Makino. six species (i, Te .eoccinea, T divergen sBureau
Tofeldi acoceinea was also reperted from & Franch,, T farus (eHiiyam aMj. N, Tamura &
Kamchatka by Ledebour (1852 f)r,om Shumshu Fuse, T nudd Maxim,, T thibetica Franch, and Z
Isl i.n the Kuriles (a sZ nutans) by Yabe and Yen- yoshiiandi have always formed a clade with 100%
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnSyts teSmaytsictsematics
June 2013 TAMuRA & AL.-Varieties of 71V7eldi caoccinea in Japan & Korea 31
TABLE l. Sources efrnaterials of 7tvieldia.
Taxon Locality VoucheT Acc, no.(trnK) Acc. no.(trnL no.(ITS)
& trnL-F)Acc.
7bjfeldi acalyculata Wahlb. Italy :Lago di Garda, Vleminckx F. 643 (MO) ABS41029 AB451579 ABS41080
Val di Vesta, 550 m
ZcotcineaRjchardson

var,coccmea USA/Alaska,LimeHills, C. L. Parker di ABS41031 AB451586 ABS41082
SSO-670 m R, Lipkin 88S4 (MO)
Japan/Hokkaido, M. N. 7bmura& AB746433** AB746440** AB746447**
Mt. Yubari, l23S m S. Fuse 15209 (KYO)
var, akkana (T ,Shimizu) Japan]IwatePref.,AkkaX Horii s.n. (KYO) AB746434** AB746441*S AB746448**
T. Shimizu

varF.u dsie bvoatr.r ynao vM., N. Tamura & JaTpaann:zKaawnaagawaPref., M(NK.Y7Ol)amNu,ra20324 AB746435** AB746442** AB746449**
var. ,7Ziu r(iHe ,Liev .& Vaniot) Korea: S, Lee NSOO09182 AB746436** AB746443Ss AB746450**
T. Yamaz. Gyeongsangnam-de, (EWH)S.
KoMrt.e aG:ayJae,j u1-40d0o ,m M. Eum Eum030PJ42 AB746437** AB746444,* AB7464Sl**
Mt, Hal[a ,1600 m (EWH)K.
var. gracili s(Franc &h .Sav.) Japan:AkitaPref, Hinyashis.n .(KYe) AB746438** AB746445** AB746452**
T. Shimizu Dakigaeri
var, kiusiana (Okuyama) Japa n/MiyazakiPret, S. Kurogi & AB746439** AB746446** AB746453**
H.Haravar, Mt,Dohdake T Mlr'namitanis.n.(KYO)
kondoi(Miyab e& Kudo) Japan:Hokkaide, M AC rtrmur adi AB541032 AB541061 AB541e83
H. Hara Mt. Apoi, 635 m S. Fuse 15227 (KYO)
Japan: Nagano PreE., M. N. 1bumura, S. Fbus edi AB541033 AB541062 AB54r084
Mt. Amakazari ,19SO in r lshi i14344 CKYO)
T divergens Bureau & Franch. China:Yunnan, M[ N. Tlamura 4704 AB541e34 ABS41063 AB541085
Dali, 2530 m (KYO)M/
Z jurus e(iHiyarn aM). N, Tamura Japan: Tochigi Pref,, rv ,7bmuna & AB561166 ABS61176 AB561186
&FuseZ Ohya, 165 m S. Fuse 20J35 (KYO)
gtabra Nutt. USA:NorthCarolina, ". N. Mbtdenke 113 AB54103S AB4S1590 ABS41086
SofJacksonville (MO)M.
T nuda Maxim Japan/ Mie Pref., iV /Tlrmura, S. Fuse & AB541036 AB541064 AB541087
Washiyama, tlOm 7: Lshii 15i71 (KYO)
T okuhoi Makino Japan:Hokkajdo, M. N. 1lamura& AB541039 AB541067 AB541090
Mt. Yubari. 13S5 m S. Fi!s e15216 (KYO)
Tpusilla(MichxJPers. Austria/Mt.Dachstein, M N. Tlamura & AB541044 ABS41071 ABS4109S
subsp, austriaca H. Kunz 2030mChina/ S. Fuse 160 74 (KYO)
T thibetica Franch. Sichuan, Hongya, C. Il Li 394 (MO) ABS4104S AB451596 AB541096
1150mJapan:KageshimaPref.
ZyoshiianaMakino ff .Okada s.n. {OSCU') ABS61174 ABS61184 AB56I194
var.yoshiiana Yakushima lsl,,
Kosugidani
i OSCU = herbarium, Botanical Gardens, Graduate Schoo lofScience, Osaka City University.
*' DNA sequences that were newly determine idn this study.
bootstra spupport in earlier studies (Tamur eat al. the best model of nucleotide eyolution was esti-
2010, 2011) .Using this new matrix, we conducted mated using Modeltest 3,7 (Posa danad Crandall
parsimony and likeliho odanalyses and con- 1998), The models selected by the hierarchical
structed molecular phylogenetic trees, likelihoo dratio test (hLIr ran)d Akaike informa-
DNA extract{on, pelymerase chain reaction tion criterion (AIC )were TrN+G and GTR ±I+G,
(PCR )amplification and DNA sequencing were respectively. Both models were implemented in
'lamura
perfbrmed as describe dby et al, (2010).PAUP* as describe dby Tamura et aL (201 1M)P.
Maximum parsimeny (MP) analysis using bootstra panalysis (100 0replications) and ML
PAUP' version 4.0 beta 10 (Swoffo 2r0d02) was bootstr aanpalysis (10 0replications) were per-
also perfbrmed as describe dby Tamura et aL formed using PAUP*, as describe dby Tlamur aet
(201 0F)o.r maximum likeliho o(dML) analysis, al, (201 020,11).
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
32 Acta Phytotax. GeoboL Vol. 64
VOucher specimens fbr the molecular phylo- lecte dseveral individua lofs TQfieidi afi:om a pop-
geneti canalyses were deposite din the herbar iofa ulation in Tanzawa, Kanagawa Pref, Japan, and
Kyoto Uniyersity (KYO) ,Ewha Womans Univep transplanted them to the nursery of Osaka City
sity (EWH) ,Museum ofNature and Human Ac- University ,The cultivated plants consistently
tivitie sH,yogo (HYO) or the Botanica lGardens, preduce panicles with distin cltater ablranches.
Graduate School of Science ,Osaka City Univer- 7}'iant h(aNutt B,a)ker, which is sister to Tojleld-
sity. ia, has a reduced panicl ewith three flowers per
node, but lack sbranches .Tofelcfi acoccinea var.
Results and Discussion gracili srarely has two (rare tlhryee) fiower sper
node, but is unbranched. Panicles with distinct
Morphoiogical observations: recognition ofvari-latera lbranches have never been reported in 7b-
eties within Ibfield icoaccinea in lapan and Kb- field iora 7i"ianth a.We therefbr econsider the
rea plant sof Tofelcfi afiro mTanzawa to be distinct
We recegnize seven varieties within Ttlfietdia and undescribed. Since the Iletnza wpalant sdiffer
inJapan Korea: kondoi, littlferom T
coccinea and coceinea, coccinea var. gracilis ,which we
graciiis ,kiusiana ,geibiensis ,akkana and di- consider most similar morphologically to the
botrya (va rn.ov.). The diagnosti ccharacters [lanzawa plants e,xcept for the ramification, we
among the seven varieties include variation in regard the Tanzawa plant sas a variety of Z coc-
leaf lengt hwithin the same individua lle,ngt hofa cinea and describ ei tas T coecinea var, dibotnya,
scape and inflorescence ,infiorescen cteypes,
pedice llength t,epal length ,length ratio of outer Tbfieldi caoccinea Richardso nvar, dibotrya
tepals to jnner ones, anther color, and densit yand M. N. Tamura & Fuse, var. nov. Fig .1
color ofcapsules, The character states ofeach va- Aennis 7ofetdiae coccineae var, gracili ,sed infiorescen-
riety are liste bdelow under `Artificial key te the tia paniculata (dibotr ydiav)ersa.
varieties of 7btfield icoaccinea in Japan and Ko- 7){pu sJ.APAN, Honshu: Kanagawa Pre£, Ashigara-
kami-gun,YlaLmakita-cho,Tanzawa. Cult.inOsaka City
rea7'.
OurtaxonomyagreeswithYamazaki's(2O02),Univ., Japan, 10 Sep. 2010, M N Tlamura 20324 (holo-
KYO; iso-HYO).
the most recent treatment of Tojieicl icoaccinea in
Japan and Korea, in recognizing the distribution Herbs perennia l,glabrou s.Rhizome short.
of var. coccinea to be northward from central Stolons absent, Leaves basal ,distichou sba,sally
Hokkaido(Japan) Pyeonganbuk-defPyeon- linear linear-falca6t-e1,4
and equitant, unifacial, or
gannam-do (Kore aan)d in recognizing T coc- cm long ,1.2-4 mm broad ,variable in lengt hin
einea var. gracili sb,ut differ sfrom Yamazaki the same individua lg,raduall ylong-acuminate
(200 2as) fo11ow sO.ur circumscription of T coc- with apex straight or slightly curved, scabrous
cinea var. kondoi is wider than Yamazaki's with small knobby projection son margin; nerves
(2002 )as, it includes Z coccinea var, j2iuriei4,-11 .Scape with inflorescen c1e5-25 cm long,
which Yamazaki (2002 t)reated as a separate va- with 2 or 3 small sterile leayes ;axis reddish
riety. Our circumscription of Z ceccinea var, brown or green .Infiorescenc ea panicl e(di-
gracili sis narrower than Yamazaki's (2002 a)s, it botryum) ,3-7.5 cm long ,branched ,10-55-fiQw-
excludes Z coccinea vars. kiusiana, geibiensis ered; longest latera lbranch O.8-2. 6cm long;
and akkana. We regard the latte rthree varieties bracts lanceolate ,green. Flowers usually ascend-
as distinc tW.e also describ ea new variety, di- ing ,rarely horizonta lS,eptember ;pedice lI,5-5,5
bots:y af,rom Japan. mm long ,reddish brown or green; calyculus (epi-
calyx) 3-lobed ,Tepal s6, narrowly oblong, 2.5-3
Morphologica lobservations: a new variety qf mm long ,white, sometimes reddish brown abaxi-
1[bfieldiacoccinea ally, Stamens 6, as long as tepals ;filament wshite,
In October 2000, Fuse and Katsuyama col- anthers orange, Pist i2l.8-3. 5mm long ,white or
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
June2013 TAMURA & AL.-Varieties of Tl?fietdi acoccinea in Japan & Korea 33
2
ase
7'IW/
-$kl$gesi.
EEr
tw
FIG,1, 7bj7etdia coccinea Richardson var. dibotFly aM. N. Tamura & Fuse. a, Habir; b, flower in side view; c, flower in back
view; d, inner tepal and stamen in adax ial view; e, outcr tepa land stamen in adaxial view (M. AC 7inmura 20324, KYO). a
panicl weith distinc Ltatcra blranches (M: N 1lrmura & S. Fuse f4899, KYO).
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
34 Acta Phytotax ,Geobot, Vol. 64
slightly creamy; ovary superior; styluli 3. Cap- three plasti dregions were combinable, Further-
sules (exclud sitnylguli) ellipsoid, 3-3.8 mm long, more, the P value ofthe ILD test of the combined
ascending, horizonta lor descending. data set ofthe three plasti dregions and the nucle-
lapanese name. Edauchi-zekishou (nov.). ar ITS data set was O,14 T,herefbre w,e concluded
Distributi oannd habita tK.nown only from that the data set of the three plasti dregions and
the type locality ;wet place sin crevices of cliffs nuclear ITS data set were also combinable.
along streams. The combined MP analysis ofthe trnK+ trnL
Other specimens examined. jAPAN, Honshu: Kana- + trnL-f7 + ITS sequences yielded 2835 equally
gawa Prcf, Ashigarakami-gun, Yamakita-cho, Tbnzawa. most parsimoniou strees of563 steps. The censis-
Cutt, in Osaka City Univ., Japan, M[ IN L7lrmura & S, Fuse tency index (CI )includin ugninformative charac-
14899 (KYO) J,54Z2 (KYO) M, IV /7lrmura 2014S (KYO). ters ,retention index (RI )and rescaled consisten-
cy index (RC )values fbr each tree were O,88 ,O,84
Molecularpbylogenetic analyses: DAL4 seguence and O.75, respectively.
variation veithin Tbfieldi acoecinea The topology of the ML tree using the model
The plasti dtrnK genes ranged from 2552 to selected by hLRT was identic awilth the topology
2614 bp, with the matK gene occupying 1548 bp of the ML tree using the AIC model. No incon-
having three exceptions of 1566 bp in samples of gruence was fbund when they were compared
T coccinea var. coccinea (fro Amlaska and Hok- with the topology of the MP strict consensus tree,
kaido )and Z coccinea var. akkana. The plastid except fbr the clade of Tofeldi aeoccinea var.
trnL gene and trnL-F intergeni cspacer as well as kondoi (Niiga tanadj Z coccinea var. ,fauriei,
the nuclear ITS region invariabl yconsisted of550 which formed in the ML trees but collapsed in the
bp,234 bp 581bp, Values
and respectively, of MP strict consensus tree (Fi g2.),
pairwis esequence divergenc eranged from up to According to the MP strict consensus tree as
O.79% for trnK, O.73% fbr trnL and O.52% for well as the ML trees (Fi g2.), the samples ef 7b-
ITS, DNA sequences of trnL-F were exactly the fiel deioccainea formed a clade that received
same among the 7 varieties and 10 samples ana- 1009i 6bootstra psupport. In this clade, Tofeldia
lyzed. kondoi, fauriei,
coccinea yar, coccinea, var, var,
var. gracili sand var, akkana furthe rfbrmed a
Molecular pbylogenetic analyses: tqpology and subclade, although the bootstra psupport was
inj?)rmatio nofth terees weak at 57-62 9'6 ,Tofeldia coccinea var. di-
In Toj7eld icoaccinea and all ofthe other sarn- botiya as well as T coccinea var, kiusian awere
ple sanalyzed in this study, from the 2750-bp excluded from the subclade.
aligned lengt hof trnK, variable sites cornprised Toj7eldi acoccinea var, dibot?: yias morpho-
299 bp, among which 92 bp were phylogenetical- logical lmoyst similar to Z coccinea var, graciiis,
ly infbrmativ eO.f the 583-bp aligned trnL se- However, only the latte rvariety was included in
quences ,54 bp were variable, and 21 bp were the above subclade; variety dibotry awas exclud-
phylogeneticall yinfbrmativ e.Of the 267-bp ed frem it (Fi g2.) .This evidence supports our
aligned trnL-F sequences, 22 bp were variable morphological observations that Z coecinea var.
and 12 bp were phylogenetica lilnyformative ,Of dibotr) ,ias differe nftrom T coccinea var, graci-
the 593-bp aligned ITS sequences, 89 bp were li sand thus worthy of taxonomic recognition.
variable and 47 bp were phylogeneticall yinfbr- Likewise, our molecular phylogenetic trees agree
'matlve.
with our opinion that Z coccinea var. kiusiana
The incongruence length differenc e(ILD) differ fsrom Z eoccinea var, gracili s(Fi g2.) ,al-
test among the data sets of the three plasti rde- though Yamazaki (200 2co)nsidered var. kiusiana
gions, trnK, trnL and trnL-E returned a P value to be included within var, gracitis,
ef O,25 ,Considerin gthe incongruenc ethreshold In the subclade of Tt2field icoaccinea var. coc-
of O,05 ,we concluded that the data sets ef the cinea, var. kondbi ,var. var. gracili sand
.fattriei,
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
June 2013 TAMIJRA & AL,-Varieties of 7bf7etdi caoccinea in Japan & Korea 35
TL coccinee var. coccinea (U.S. AAl.a,ska)
T coccinea var. coccinea (Hokkai dMto, ,Yubari)'
11 coccinee var. konclo i(Hokkai dMto. ,Apoi) tso:ny--o"oes
TZ coccinea var. konclo i(Niiga tMat., Amakazari)
r coccinea var. fauri ei(Kore aMt,, Gaya)"
Z coccinea var. fauri e(iKore aMt, ,Hal[a)'
7: coccinea var, gracili (sAki tDaak,igaeri>*
71 ceccinea var. akkana (lwat Aek,ka)*
71 coccinea var. kiusian a(Miyaza Mkti, ,Dohdake}*
71 coccinea var. cfibotry evar, nov, {Kanagawa T,anzawa)*
T nudaZ
yoshiiana

T furusei
T thloetica

T divergens

T okuboi
Z glabraT
pusMaT

calyculataoutgreup

FIG. 2. Stric tconsensus of2835 equally most parsirnoniou strees resulting from the combined maximum parsimony (MP)
analysis of the plasti dtrsnK, trnL and trnL-li; and nuclear ITS DNA gequence data from 7hfietdi aT,he topology ofthe
rnaximum likeliheo d(ML) trce using either ofthe models [i. tehe.se selected by hierarchica llikelihoo dratio test (hLRT)
and by Akaike informatio ncriterion (AIC ) ]is identica wlith the topolegy ofthis figure. Arrowhead indicate as branch not
presen itn the MP strict consensus trec. Numbers above branches indicat cMP bootstra pvalues. Numbers below branches
indica tMLe (hLRT )beotstr avaplues ! ML CAIC )boetstra pvalues. Accessions with an asterisk were newly included in
the molecu lar phylogeneti canalysis.
var. akkana, the two samples of Z coccinea var, Yamazaki (200 2t)reated Toj7eldi acoccinea
fauri (eoin efrom Mt, Gaya on the Korean Penin- var. coccinea as being from northern and central
sula and the other from Mt. Halla on Jejud oIs- Hokkaido in Japan and considered plants from
land) were grouped with each other (86-879,6s)ou,thern Hokkaido to be Z coecinea var. kondoi,
Although Yamazaki (200 2c)onsidered T coc- In our molecular phylogenet iacnalysis, the sam-
cinea var. fauri teoi be endemic to Jejudo Island, ple from Mt, Yinbari in central Hokkaido was not
our identificati onof the sample (A iSl, Lee grouped with the sample from Mt. Apoi in south-
A[SOe09182 ,E}VH) from Mt. Gaya, the Korean ern Hokkaido but rather with the sample from
Peninsula, as X coceinea var. fouri seeeims to be Alaska ,U.S.A. (58-659 arlt6ho)ug,h the geograph-
correct, indicatin gthat T coccinea var. fauri iesi ica ldistanc ebetween Mt. Yinbar iand AIaska is
not only on Jejudo Island but also on the Korean far greate trhan that between Mt, Yubari and Mt,
Peninsula, Apoi, Thus, Yamazaki' s(200 2c)ircumscription
In the ML trees ,the two samples of Tojielcita of Z coccinea var. coccinea is supported by our
coccinea var, juuri efirom Korea were further molecular phylogenet icresults as well as our
grouped with T coccinea var. kondbi from Mt, morphological ones,
Amakazari, Niigata, Japan (50-58% )al,though Based on specimens in the KPM herbarium,
this grouping collapsed in the MP strict consen- Ttl17eld icoaccinea var. gracili sis distribut ealdso
sus tree. Based on the morphological similarity in Tanzawa, where T coccinea var. dibotT) ,waas
between 71 eoccinea var. fauri aenid T coceinea found. Unfortunately ,however ,we were unable
var. kondoi ,we prefer Hara' s(196 1i)nclusi oonf to obtain samples of it ,now of Z coccinea var,
var.fourieiwithin T coccinea yar. kondoi ,which geibiensi sfor our molecular phylogenetic analy-
does not conflict with the groupin gin the ML ses, Future molecular phylogeographi cstudies
trees. with more samples covering all the varieties and
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
36 Acta Phytotax. Geobot. Vol.64

the wholedistribution area of Z coccinea areneeded to confirm our taxonomy,
Artij7ci kaaly to thevarieties ofTbfieldiaco cicn .ilinpann ea andKorea
la. Infioreseenc ea panicte ,with distinc t]atera lbranche s-..H...,....,..,,.,.....,....,, .v.a.r.. ,d.i.b.o.t.t,l.y.a..,,,,..,,....,..
lb. tnflorescenc ea simple raceme, very rarely a reduced panicle without any branches ...........,,,... .2.........,.....,.
2a .Pedicel Os,5-2 mm long ;capsules dense ,dark brown or blackis bhrown .,,....,.....,.. .v.a,r.. .co.c.ci.ne,a.,,.,,,,
2b. Pedicels 2-10 mm long; capsules relatively loosel sypaced, brown ....,,....,,....,.,..-,.H.,.H.. .3.-..H-...-...-.......
3a. Scape with infiorescenc e5-13. 5cm tong ;pedicel 2s-S(-5,5) mm long; leaves subequa] in length on same
individu a-l,--."".."...H....,....,,....,,...H...-,,-,,,H..H..-....-....,...... .v.ar... k.o.n.d.o.i............,...H..H-.H-...,,,,,H,,."...
3b .Scape with infioresce n8c-e30 em long; pedicel 3s-10 mm long ;leave vsariable in length on same individ-
ual.....,,"",,H..."...".."...-...H....,....,,,,,,,.,......"..."..H....-..-...,....,,,,,,,.....,....,.....,.........."...".......H....H...H....H......-4
4a. Pedicels 3-6 rnm long ..,,.,,,,,,...,....H..H".H..H...H....,.,.,",,.,....,,....,.....,. ,5.,,,..."..."....-.H...H...H.....-.-H.,"M,..."..
4b.Pedicels6-10mmlong.,,....,,,,,-,,.H...---.--..m..m.............,,....,....H...H,.,,",,."....,....,...H..H"..H...H-.･-･･,,-･.･.6
5a. Scape with infiorescen 8c-e20 cm long ;tepals 2.5-3, 5mm long ;anthers purplish .......... .va.r.. .g.x.ac.i.t.is
5b. Scape with infloresce n1c5e-30 cm long; tepals 3.5-4 mm long ;anthers pale purplish ..... .v.a.r.. .kittsiana
6a .Outer tepals nearly as Iong as inner tepals .,,-....,....,,....,,....,,....,....".. .v-ar... g.e.i-b,i.e-n.s.is.H..--..H-..-...
6b. Outer tepals 1/2-213 times as long as inner tepals ..,,.....,........."....H...-....-.H v.a.r., Hak-k,a"na",,,,,...,....H....
7lrxonomic tTeatment Kudo) H. Hara £ rishiriensis (Miyab e& Kudo) Sug-
Tofieldi caoccinea Richardson in Franklin, Narr. irn. ,Keys Herb, Pl. Jap. 2: 566 (1973 )=I.Ype: Japan,
JourneyPolarSea:736 (1823-)[.[tsrpeC:anada, Prov .Kitami (Hokkaido I)s,l .Rishiri ,Mt. Rishir ia,t
Nunavut, at th e Ar ctic Sea near Bathu rst Inlet, tShAeP Ssu!m,mi t,hic Adueg.s i1g8n9a9t ,e7s 1)KA.anowtahkearm is ysn.tny.p e:(l eJcatpaont,ype
anon. s.n. (holo-BM d,igita ]image BM!). Prov .Kitami (Hokkaido I)s,l .Rishiri ,Mt. Rish{r{ ,at
the surnmit, 3 Aug. I896, M Hirose s,n. (SAPS!).
var. coccinea (Fi g3,) Jinpanese name. Chishima-zekishou.
7 t zf7i e(l2d in) uat:1a5ns73 (W1il8l3d0 . ie)xn oSbcsehruvl.t &.su bS cZh ulcet r£n.u,a .Sy s-t 4.Vpeeg:, KDiosrterainb unatmieo. na nSduk -heauhni-tkak to.Rtusjsainag -(peo.astward
Russia, eastern Siberi aN,, PaUas s.n, (holo- Bcl,igital from E Siberia),Mongolia, NE China,Korea
imageB!).
Tt lfieid iafasMciyaabe & Kudo inTrans.SapporeNat. (northw afrrdom Pyeonganbuk-do f Pyeongan-

Hist. Soc. 5: 75 (191 4-)7b.j7etd inatctans Wirld. ex nam-do), Japan (northw afrrdom C Hokkaido),
Schult .& Schult .f var. fasc a(Miyab e& Kudo) Alaska, Canada and Greenland .Alpine slopes
Koidz .in Bot. Mag, (Tokyo 3)1 :138 (19]7 -)7,Llfieldiaand rocky places.
ny6ua8tab n(es 1& 9 WK5iul d3-lo)7d)1,. 1eOxfh wiSiecl hid cnui olBaccu ti&ln.ela . SNca htR.ui clShtc afi,r .dvMsaeurn.s, f Tyiaornk.sy j ec(u aM3s3ic:-a Mi yaAbreot & eO. nKeu doof fthreo ms yJntayppaesn, H oonfs Thou,fe Plrdoiv af. aSshic-a
(Miyab c& Kudo) H. Hara in J ,Jap. Bet, 36: 392 nano (Nagan oPref) ,Mt. Shirouma, 12 Aug.
(1961 -)7,tlfield cioaccinea Richardson fi fins c(aMi- 1904, S. Kbmatsu s.n. (SAPS! i)s T coccinea var.
yabe & Kudo) Q S. Sun in P. Y. Fu, Clavis Pl. Chinae koncfoi.
Boreal.-Orie nedt.. 2: 764 (1995 )-T.lype :Japan ,Prov.
NIisshhiikcalr aiCsH,no. kk(aliedco Mt)to,. t YySupAbaPerSi!, , 7-h9i cA udge, si1g9n1a3t, eSs,; var, kondoi (Miyab e& Kudo) H. Hara in J.
isolectoty pSeAPS!) .Other syntypes: Japan, Prov. Jap .Bot. 36: 392 (1961 )(.Fi g4). -TQfieldia kondoi
Ishika r(iHokkai dMot). ,Yubari ,on grasslands in Miyabe & Kudo in Trans .Sapporo Nat. Hist. Soc,
northern part ofthe meuntain, 6 Aug. 1912, H/ Minag- 5: 74 (191 4-)To.jieldiafasca Miyabe & Kudo var,
isawa & A. Hbmana s,n. (SAPS! )P;rov .Ishikari kondoi(Miyabe& Kudo) Tatew. inRes.Bull,
(NiHsohikdka a&i Hd . Mo}t)Z. ,iAnsahgiiuspae wtsa.nn.u p(uSr Ai3,P-5S A!u)g,. 1913, S. Exp. Fo r. Hokkai do Univ. 5 : 12 (1 92 8 -)T,Qf ieldia
71zfieldiajus eMaiyabe & Kudo f .rishiriensis Miyabe & nutans Willd. ex Schult .& Schult .f var. kondbi
K-7ubdyo7 eiln dT ircaaocncsin.e aSap pRoircoh aNradts, onH ivsart.. Sjoucs. c5 :( a7M6 i(y1a9 b1&e4).(52M:i55y9a(b &1e 9K3ud8-o)T)}. iHp.e :HJaarpaa ni,n PBroovt.. MHiadga.k a(T(bHkoyko-)

NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
June 2013 T･"dL'R,x &K[ . --Variet iofes Tl?fieidi acoecinea in Jupan & Korea 37
F]c] .3. Dense raceine of 71)fieid icoa('('inea Richardson vur. FEce 4.. Sli.ght ]lyoc}s reaceme of 1?tfield cioaccinea R{chard-
coeei"ea frem Mt. Yubari, Hokka{do. Japan. son Nar. kondoi (Milr'a be& Kudo) H. Hara from Mt.
Apol, Hekkuido. Japan.
kaido) S,amani, Mt. Apoi. 17 Aug. 1912, K. Kon- 193S. B. Sonokis,n, (holo-SAPS! i:so-SAPS!).
do s.n, (holo-SAPS!). T ? lnJda'pi d. Bicooact'c. '3in6e:a 3 9R2i c(hIa9r6d1s )o-n.T fy p.Jc): aJlalepsacne .Hn osHn.s Hhua.r a Piron Jv..
7tlfieidia.17nt HJ.' iLe6iv. & Vaniot in Repert. Spcc, NoN, Rikuchu (I"al ePret). Mt. Iwate. 13 Aug. 1929. }[
Regni Veg. 5: 283 (1908) ,-Tl?fie/ dctoeaeinea Rieh- NaJ'ii c,rs. n(,holo-Tlb.
ardson var..ftutriei cH. LeN･ &' ,VanioO T. Yamaz, in J,
Jap. Bot. 77i 302 ･{2002 )-,b'pe: Kereu. Jeju-do ,Mt, .lttpanes enaine. Apoj-zekishou.
Ilalla ,Aug. 1907, U. FZittr i2e107(holo-P. dlg'tt ai[m- Kt"'ean name, Halla-kkotjang-po.
age P!; iso-KYO!. Tl!). Distri ihtt.icn andhabitat. Japan (S Hokkaido
7 ? tfjRicue.l-ud nditi aaVoMqctt,g.t .e Ht5iai: l H2l.8a 3. L 1e(7x01' 0.9& m0 .V8 a)'Onct.itco.; t q1i9in 0tZRe -e tpT7ie}1lr'itqp, te St:K epo4ter0ec4 a. (.N loJevec.-- aKnedre Nan & PCen iHnosnuslhau )a nadnd MKto reHaa l(lMa to n.G aJyeaj uond otIhes-
totype P .digjt ailmage P!. hic designates: isolecto- land) .Alpine and subalpine slopes and rocky
t},pe KYO!). .ALnoth esyrntype: Korea. Jeju-do .i.h-OO places,
m. Oet. 1906, U. Fat,ri e264 CKYO!. P. Tl!. di.gi tiam]-
age P!).
ft?fiel dsui:aoensis 'Miyabe & Kudo in Trans ,Sapporo N'ar. gracili s(Franc h&, Sav, )T. Shimizu,
Nal. Hist .Soe. 5: 73 {19]4 )-.T.ype :Japan. Proi. [buri Ne"･ Alp. Fl. Jap. in Color 2: 358 (1983 ()F.ig ,5)
CHokkaido ).Mt. Yeni"'a ,Aug. I895. Z K(ni'akafni -7??fieldi gai'ac'i lFirsanch .& Sav. ,Enum. Pl .Jap,
s,n, (ho]o-SAPSI), 2:89(1877), 531(1879)-.71?fieldia
n ? fiTeaJtdei wa.. ii'ne :Toreann ss.iM isSyaapbpeo ro& NKautd. oH ivsart.. Sooktci.s hii]'6en: si1s]5 Wi 1l d. ex Sc hnuoml.t &. e tS chu lt f.i var. gracili s( Fntrttaansnch.
(1940) .-fype: Japan. Prov. Shiribeshi (Hokkaido). & Sav. )Oh"'i in Bull. Nat, Sci ,Mus. Tokyo 33: 68
Isl .Okushiri. the upper Kamoishi River. ]O ,ALu.o.(I953 )-T.ype: Japan. in northern mountain, Sa-
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
38 ActaPhytotax.Geobot, Vet.64

limestone,
wet, often rocks.
va: kiusiana (Okuyam aH), Hara in J .Jap.
Bot.36:393(l961-)TQ.fieldiakittsianOakuyama

inJ,Jap,Bot.26:294 (1951-)T.ofeldia
nutans
Willd, ex Schult . & Schuit . f var. kiztsiana
(OkuyamaOjhwi inBull,Nat.Sci.Mus. Tbkyo

33:68(1953-)[[,Ype:Japan,Kyushu,Prov,Hyuga

(MiyazakPireb, Mt. Horagatake (Mt.Dohdake),

23 Aug. 1915,Z. 7Zishiros.n(.holo-TNS!).

lapanese Nagae-chabo-zekishou.
name.
Disxributi oannd habita tE,ndemic to .lapan:
Kyushu (Miyaza Plr<eiE). At the southern border
ofthe distributio rnange of Tofeidia coccinea, On
limestone
rocks.
var, geibiens i(sM. Kikuchi) H. Hara in J. Jap.
Bot.36:393 (1961-)T.ofelcliakiusianaOkuyama

var. geibiensi sM. Kikuchi in Ann. Rep. Gakugei
Fac.IwateUniv.11(2)6:7, 2(1957-)IY,pe: Ja-
£
pan, Honshu, Prov. Rikuchu (Iwa t?ere£) ,Hi-
gashi-iwai-gun (lchinoseki -Gseihbii)-,ke i,15
Aug. 1947,AdiKikttchis.(nh.olo-iUiMs;o-TNS!).

Geibi-zekishou.
.llypanese name,
Distrihutio nand habitat .Endemic to Japan:
Honshu (lwat ePref). On limeston ecliffs along
rlvers.
var, akkana <T .Shimizu) T, Shimizu in J.
Fac, tcxt, Sci. tcchnol. Shinshu Univ. 36, ser. A,
F[G. 5. Extreinel yloese raceme or 7}?fietd icoaccinea Rich- Biol ,12: 74 (1963 -)Tl.lf7eldt aakkana T. Shimizu
ardson var. gracili s(Franc h&. Sav. )T. Shjmizu fvom inActaPhytotax.Geobot,17:153,f,13(1958-).
Dakigaeri ,Akita Pre£, japan. [f ype:J apan,Hons hu,Iwa teP ret, Sh im ehei-g un,

Iwaizumi-cho, Akka, 15 Oct. 1957,T Shimizu

2364 (holo-KYO!),

3749 (holo-dPig,italimage P!). lapanese Akl<a-zekishou.
vatien nan2e.
7b y f1e1t d:4i 3csoi7r d(id1o86 M7 a-)xT.iLiVnf. eilnd Bi nuazlttla. nsAe aWdi.l lScd i,e .xS tS.c-bPeutlet r&.sb. H o nDsih.us 't(rIibwuat i tPoanrnedef )h. abLijtmaets .tEonndee micrcev itcoes J apanadn:
Schult .f var. sordidti (Maxim .T), Shimizu jn Acta gravelly slopes.
Phylotax, 6eobot. 17: 153 (1958 )-1.lype :Japan, cult.
Yedo (fokyo )fr,om the neighboring mountain (n.v.). var. dibotrya M. N. Tamura & Fuse (Fi g6.)
11zlleldia Franch.& Sav.,Enum, Pl.Jap,2:530
stenantha (see
(187 9=f)yp.e: Japan, in Hakone region, Savatie r1235 above)
(holo -diPg,ita timag¢ P!), Correct name and assignment of rlbfieldi acoc-
kanwonensis
.fLipanes ename. Chabo-zekishou. cinea van
Distributi ocxnnd habita tE.ndemic to Japan: Yamazaki (200 2d)escribe dTbfield iacoc-
Honshu, Shikoku and Kyushu (Ohi tParef) ,On cinea var. kanwonensis based on a specimen
NII-Electronic Library Service