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Phylogenyof Speciesof 7Bfieldiaand from Sequences

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Phylogenyof Speciesof 7Bfieldiaand from Sequences The JapaneseSocietyJapanese Society for Plant Systematics ISSN 1346-7565 Acta Phytotax. Geobot. 60 C3): L31-140 (2010) II. Biosystematic Studies on the Family Tofieldiaceae Phylogeny of Species of 7bfieldia and 7)iantha Inferred from PIastid and Nuclear DNA Sequences MINoRuN.TAMuRAiZ HiResHi JuN YAMAsHITA2, SHizuKA FusE3 ANDAzuMAI,TAKAAKI IsHII4 'DqT)artment Sak.yo-ku, ofBotaay C;raduate Schoot qfScience. K.voto Ultiversio,, 1(itashii'akawa-oiwake-eho, 'Research, lk5,eto 606-8502, JLavan. *[email protected],acv'p (keuthorforcorrespondencel; bistitutefor jMiiseuni Bioresources, Okctyama Univensity, 2-20-1 Chuo, Ktirashiki-shi, Oka.vama 7iO-e046, Jk4Jan; ofIVdture fBotanicalGarctens, andfinman Activities, flyogo, 6 l2ll'oigaoka, Sando-shi, i'lyogo 669-i546, JLipan; Graduate Schoog (ef'Science, Osaka City LJniversity, 2000 kri,saichi, Katano-shi, Osaka 576-OO04, Jtupan Using pLastid trnK, trnL and trnL-F sequcnces and nue]ear ITS sequenccs (4558 bp in total) frem 32 in- dividuals of 13 speeies, we conducted a cembined analysis to clarify phylogenetic rclatienships ot' the spccies of T(ofieldia ancl 1)'iantha and to discoyer new taxonomic eharacters, According to strict consen- sus of 45 most trees, the two are menophyletic with leegi6 bootstrap support. In parsimonious genera 'Tb. 7Zlfieldia, three robust clades (100%) are fbrmed: (1) coccinect, 7b. nudo. Tb. diveF:gens, 7b. yunnanen- sis and 7b. thibetiea, (2) 7b. calyculata, and (3) 7b. okuboi, Tb. gtabra and 7b. pu,s'illa. The species efthe clade 3 arc primarily cltaracterized by a broad and sharply bent leafapex as in some species of 7}'iantha, Jn clade 1, the Chinese species of 7b, dtveFlgens, Tb. yunnanensis and 7b. tkibetiea are groupcd (100%), They share long need]e-like or hair-like proccsses on the leaf margin, ln 7>'iantha, two clades are formed: 'T}'. (1) J'aponica, 7}: occidentalis and 7h giutinosa frDm the USA (76%), and (2) 7>". raeemo,s'a and 7}', glu- tinosa t'rom Canada (91%), This might suggest that 7}t glutinosa is the assetnblage retaining ancestral character states of 7}'iantha and shou]d be divided at least into two species jn the future treatments. '7ofeldia.rlbfieldiaceae, Key words: Alismatales. ITS, molecu]at phylogcny, taxonomic eharacters , T}'i- antha, trnK, trnL, trnL-F [Ibfieldiaceae (Alismatales) are rhizomatous The only molecular phylogeny of Tofieldiace- "Ibfieldia- perennial herbs formerly included in Liliaceae. ae (TUmura et ai. 2e04aj revea!ed that The species grow in rocky alpine places, alpine ceae consist of Tofeldla Huds,, 7}"iantha (Nutt,) meadows, marshes, savannas, rock faces along Baker, Hlriflperocallis McDaniel and Pleea Mi- rivers, etc., and often on calcareous soils. The chx., and resolved the phylogenetic relationships flowers are actinomorphic, homochlamydeous, among the fbur genera. Since that paper aimed at trimerous, hypogynous, and nearly apecarpous, circumscribing 1[bfieldiaceae, many of the spe- The capsules dehisce septicida]ly. Tliese charac- cies were not included. Except fbr the monotypic ter states are believed to be ancestral in the mono- Hdrperocailis and Pleea, enly five species of 7b- cotyledons, and [Ibfieldiaceae are often consid- two species of 7>iantha ana- .fietdla and only weTe ered to be one of the morphological]y most lyzed. primitjve families among the monocotyledons In this study, we focus on the Ttlfietdia and (Thorne 1992 for Melanthiaceae, Takhtajan 1997, 7}"iantha. Ti"iantha diffbrs firorn Tofieldia in usu- Tamura 1998 fbr Nartheciaceae). ally having three flowers per node, glandular NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics 132 Acta Phytotax, Gcobot. Vol. 60 hairs, and appendaged seeds, Nevertheless, 1)r- antha has often been included in TQfietdia. For Materials and Methods example, 7}'iantha j'cu?onica (Miq,) Bake/r, en- demic to Japan, was treated as nlfieidia in a!1 Piant mate"iaLs' published Japanesc fioras (e.g. Ohwi 1953, Kita- DNA sequences of the plastid regions of the mura 1964, Satake 1982). Tamura et al. (2004aj trnJl gene (including matK), trnL gene and trnL- shosu'ed that the clade of ftiantha is independent Fintergenic spacer and the nuclear region ofITS and sister to the clade containing the species of for nine species, one subspecies and one variety Tofeldia. The results, however, were based on an (20 OTUs) of TQfieldia, three species (10 OTUs) analysis ofonly several species, of 7)"iantha, and Harperocallis flava McDaniel We recognize 11 species of Tofeldia, TQfielcl- and Pleea tenuij?)tia Michx. (as the outgroup) ia coccinea Richardson and lb. pusiUa (Michx.)were determined (Table 1), We selected the out- Pers. are distributed widely in boreal regions. group based on the results of Tamura et al. The other species, including Tb. divergens Bu- (2004a). As fbr 7}"iantha oceidentalis subsp, oc- reau & Franch., 7b. glabra Nutt., 7b. nucla Max- cidentalis and 7k occidentalis subsp. breviso,la im., lb. okziboiMakino and 7b. thibetica Franch., (C. L. Hitchc.) Packer, we failed in determining are localized, In spite of this interesting contrast the DNA sequences of trnK and ITS, and thus we in distribution ranges, the relationships of the obtained data fbr only trnL and trnL-F(Tab]e 1). species of TQfieldia have never been resolved, Alt of the sequence data arc deposited in the 'n'iantha ¢ omprises four species: nt glutinosa DNA Data Bank ofJapan (DDBJ) database. (Michx.) Baker, 7}: occidentalis (S. Watson) R. R. Gates and 7}". racemosa (WaltcD Small in North DAC4 exgraction, PCR amplijication and DIVA se- America (Packer 2002), and 7). ,ioponica in Japan quenclng (Tamura in press), 7>'iantha glutinosa is widely Total genomic DNA was extracted from fresh distributed in the USA and Canada, while the leaves, silica gel dried leaves, or leaves ofherbar- other three species are narrow]y localized. Also ium specimens from thc Missouri Botanical Gar- in n"iantha, the phylogeny ofthe species has nev- den <MO) and Kyoto University (KYO), by the er been analyzed by molccular methods. slightly modified cetyltrimethylammonium bro- The first ai'm o'f this paper is to clarify the rn ide (CTAB) method of Doyle & Doyle (1987). phy!ogenetic relationships of the species ef Tb- The trnK, trnL plus trnL-F; and ITS regions .fieldia and 7>"iantha using amplc materials of the were amplified using the polymerase chain reac- two genera, although we were unable to analyze tion (PCR) with a programmable temperature- the Russian Tofeldia cernua Sm. and the Himala- control system GeneAmp PCR System 9600 or yan Tofetdia himalaica Baker. To achieve our 2700 (Applied Biosystems). The amplification re- goals, w ¢ sequenced both plastid and nuclear action mixture was prepared using TaKaRa Ex DNA regions, which were 4558 bp in total after Taq DNA polymerase fo11owing its instructioll the aljgnment. manual (Takara Shuzo). We used sixteen, two, of and six Perhaps because of the simple morphology PCR primers (Table2) forthe amplifica- [[bfieldiaceae, very few morphoLogical characters tion of trnK, trnL + trnL-b] and ITS regions, re- have been used for the taxonomy of npteldia and spectively, and the tbllowing PCR profile: a 30- n"iantha. The second aim of this paper is to dis- 37 cycle reaction with denaturation at 940C fbr aover new taxonomic characters fbr Tofegdia and O.5-1 min, annealing at 50-620C for O.5-1 min, 7}'iantha by analyz{ng the detailed phylogenetic and extension at 720C for O.5-3 min, in addition relationships of species of the two genera based to an initial denaturation at 940C fbr 2,5-5 min on the molecular analysis, and a final extension at 720C for 5-7 min. We pu- rified the PCR products with MicroSpin S-300 HR Columns (Pharmacia Biotech) or QIAquick NII-Electronic Library Service The JapaneseSocietyJapanese Society for PlantPlantSystematics Systematics rt(arch 20]O TAMuRA &AL.-1'hylogeny of 7kij7etdia and 7}'iantha 133 TARLE L. Sources gfmateriats of Tofietdiaeeae Acc. no.(trnK)Acc.]i[].vrn[&rrsiL-EiAcc.no.{TTS) '1'axon Locality Veucher 'TLIfieldia ccrlycugata Wahlb. Gcrmuny:Frankrurt,Palmcngarten C. Bayer ,s.n. (KYO) AB541027 ABS410S9 AB541078 Germany:Garmisch-Partenkirch- A. Cirb'ger 20.0ZV.1 CKYO) AB54102g ABS410GO AB541079 en, Schachen, l70e m lvTesta, Jta[y: Lago di Gardu, Val di eveminekx E 643 CMO) AB5410Z9 AB451S79 ABs41ogo 550mItaly : Prov, de Trento, Tinemalzo, J'. Maquei S2yPO (MO) AB541030 AB4S15SQ AB541081 1800m 7bjieidiacoccineaRichardson vttr. coccinea USA/ Alaska, Lime Hills, 550 -67U rn C. L. Parker di R. Lipkin SS84 ABS41031 AB451586 AB5410g2 (MO),Lt var. kondoi {Miyabe & Kudo) Japan/ Hokkaido, Mt. Apoi, 63S m N. Jlrmura di S. Fuse 1.i227 AB541032 AB541061 AB541eS3 H.Hara CKYO),lt Japan:Nagano1iretl,Mt. ILI 7}imaura, S, Fuse & 7/ AB541e33 AB541062 AB5410S4 Amakazari, 19SO ]n ls･hii 14344 (KYO) 7}zfieidia divergens Bureuu & China: Yunnan, Dali. 2530 m ,･td. N. lhmura 4704 (KYO) AB541034 AB541063 AB5410g5 Franeh,nVieJdi[rglabraNutt. USA; NorLh Care[ina, S of H. Al ,ldi]ldenke ILI CMO) AB541035 AB451S90 ABS41086 Jacksonville Tbjieidia nuda Maxim, japan/MiePvef,,Washiyama,11OmM. N, 7kemztt'a. S. Ri,y'e & Z ABS41036 ABS41064 ABS41087 Ishii 1517i (KYO) Japan: Hyogo Pret, KamegaLsubo, ,V. tN', 7timura & S, Fttse lj532 ABS41037 ABS410fiS AB541088 43a 1-Japan:Ilokkaido, (KYO)jdi ]QfietdiaekuboiMakinv Mt, Yubari, 1480m N llimuna & S. Fleise I5220 ABS41038 AB541066 AB541089 {KYO)M, Japan: Hokkaido. Mt. YubarL 1]55 m iV. IlimTtra & S, ")/se IS2I6 AB541e39 AB5.41e67 AB54iD9D (KYO),W. Japan/ Hokkaide, Mt. Yuhari. 134S m iV. 7lrmltra di S. )'use 1521)' ABS4104U AB541068 AB541091 (KYO),l{ Jupan/ lshikawa Preft, Ml. N. 7inmura [E S. Fute r)',519 AB54f041 ABS41U69 ABS41092 Halcusan, 2580 rn (KYO),if Japan/ Tshikawtt Pre CZ, Mt. N. 71rmura & S. Fuse i.i521 AB54t042 ABS41070 ABS41e93 Hakusan,2500m {KYO) 7}ijietdiapttsiUa (Michx.) PeFs. subsp.pusiila Denmark: Grecnland. SW of B. Fredtskild di C. Bay 3I48 AB54LQ43 tLB451593 ABS41094 Nonv'uk,5m {MO),4I subsp.
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