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Download PDF (881K) The JapaneseSocietyJapanese Society for Plant Systematics ISSN 1346-7565 Acta Phytotax. Geobot. 57 (2): i55-164 (2006) Phylogenetic Analysis ofthe Northern Pinguieula (Lentibulariaceae) Based on Internal Transcribed Spacer (ITS) Sequence KATSUIHIKO KONDOiand HIRO SHIMAI2 iLaboratory qf'Plant Chromosome and Gene Stock Graduate Schoot ofScience, Hiroshima Uhiversity 1-4-3 Klrigamlyama, Higashi-Hiroshima 739-8526 lapan; 27lakamori Orchid Museum, 5i2-73 leuhara, 7bkomori, ?Vdgano 399-3107Jirpan Internal transcribed spacer (ITS) region of 1 8-26S nuclear ribosomal DNA (nrDNA) in 18 species and three subspecies ofPinguicula distributed in the northern hemisphere was sequenced. Based on thc ITS results, species in Section Pinguicula were basically well-agreed with the orthodox classification. Although three subspecies ofR tongCfblia have been described, the present approach showed them to be separate species, The ITS sequences between R vttigaris and R bohemica were identical and suggested that the two species could be conspecific as normally treated. Pingtticula aipina did not show any direct relationship with R namosa and R variagata although they have been taxonomically placed in Section Micranthus. [Ilie present ITS study suggestcd that R vittosa, only the speeies of Section Nana, was close- ly related to R ramosa and R variagata. The northern Pinguicula taxa studied here perfbrmed the same seasonal growth pattern, but it was likely as a result of convergent evolution. Key words: carnivorous plants, DNA, ITS, Lentibulariaceae, phylogeny, Pinguicttla. Pinguicttla L. is distributed throughout all continents Fuchs (1982), Zamudio & Rzedowski (199l), Luhrs except fbr Aritarctica and Australasia and is a mem- (1995a) and Blanca et al. (1999). On the other ber of the Lentibulariaceae together with Utricularia hand, in the last few decades, many species of L, and Genlisea St.-Hil. (Juniper et al. 1989, [faylor Pinguicuifa have been described mostly in Europe 1989). Many species of the genus are endemic to (Tammaro & Pace 1987, Zamora et al, 1996, Casper only a locality or relatively small geographical area, & Steiger 2001) and Mexico (Luhrs 1995a, 1995b, and thus, they are often highly endangered (Zamora Speta & Fuchs 1982, 1989, 1992, Zamudio 1988, et al. 1996, 1997), Some species ofthe genus would 1999, 2001). Thus, approximately 90 species are be relict and their distribution have been shrunken hitherto considered valid in the genus. and eyentually migrated to higher elevations of The species of Pinguicula could bc divided mountains or to the north, in the early pestglacialinto three groups (Huxley 1992) on the basis of period (Pennington 1969). physiology, geographical distribution and horticul- "Nonhern Pinguicula was taxonomically studied by tural points ofview, such as 1) species" or "Temperate Linnaeus (1753), De Candolle (1844), Barnhart sometimes called species;' distributed in (1916) and Ernst (1961). The latest monograph of the mild to colder temperate regions in the northem "Southeastern the genus was completed with three subgenera, 12 hemisphere; 2) North American sections and 46 species by Casper (1966) and some species" mostly distributed in the coastal regions of ` minor modifications of it were proposed by Speta & the southeastern U, S, A,; and 3) ℃ entral American NII-Electronic Library Service The JapaneseSocietyJapanese Society for PlantPlantSystematics Systematics 156 APG Vbl. 57 TABLE l. Tlie taxa ofthe northern Pingnicula group and other Pinguicula group arid two species of thc outgreup in the Lentibulariaceae studied.ima Sour-c.g..&. voucherspecimen number Northern Pinguieuia Subgenus ll]mnoceras SectionMicranthus RaipinaL, a) Europe to Siberia, and Himalayas (O-4,1OOm), b) Mala Fatra (1,OOOm), Terchova, Slevakia, Kondo & Shimai 5734-LPCGS (HIRO). R ramosa Miyoshi a) E. Japan (1,500-2,300m), b) Koshin-zan (1,700m). Techigi, Japan. Kondo di Shtmai 5735-LPCGS (HIRO). R variegata [[hrcz. a) E. Siberia, Russia, b) Sakhalin Isl., N-49"43'57.S", E-144006'34.9" (240m), Russia. Ik bndo & Shimai 5736-LPCGS (HIRO). Section IVZIna R villosa L. a) N. Eurepe to Siberia and N. America (O-750m), b) Broad Pass (70em), Cantwell, Alaska, U.S.A. Kbndo & Shimai 5737-LPCGS (HIRO). Subg¢ nusPinguicula Section Pinguicttla R halcanica Caspcr a) Balkans (1,800-2,400m), VardoUssia b) (1,gOOm), Fokida, Greece. Kbndo & Shimai 5738-LPCGS (HIRO). R hohemica Krajina a) N. Czech Republic (250rn), b) Cesk6 Lfpa (2SOm), Czeeh Republic, Kbndo & Shimai 5739-LPCGS (HIRO). R conyica Bern. et Gren. ex Gren. et Godr. a) Corsica Isl, (900-2,400m), b) Gorges de la Restonica (1,SOOm), Corsica Isl., France, Kbndo & Shimai 5740-LPCGS (HIRO). R dertosensis (CafiLg.) Mateo et Crespo a) S to SE. Spain, b) Perts de Beceit (500m), Tarragona, Spain. Kbndb & S71imai 574 1-LPCGS (HIRO). RfioriiTbmm.etPace a) Maiella (750-2,500m), Italy, b) VaLle dell'Orfento (1,300rn), Abruzzo, Italy, Kbndo & S],imai 5742-LPCGS (HIRO). R gtundlTlora Lam. subsp. grandijiora Lam. a) W. Europc and Morecco (O-2,500m), b) Lac de Fabreges (1,250m), PyrenEes Atlantiques, France. 1(bndo & suimai 701-Cult.LPCGS. R grandijlora Larn. subsp. rosea (Mutel) Casper a) SE. France (650-I,500m), b) Goncelin (650m), Isere, France. Konclo & Shimai 5743-LPCGS (HIRO). R teptoceras Rehb. a) Alps and Apermines (1,1OO-3,OOOm), b) Col de Tende (2,1OOm), Alpes Maritimes, France. Kondo & Shimai 5744-LPCeS (HIRO), R longij?)tia Ram. ex DC. subsp, longijblia Ram. ex DC. a) Pyrenees (6SO-2,OOOm), Spain, b) Valle de Ordesa (1,500m), Huesca, Spain. Kondo & Shimai 702-Cult,LPCGS, R longijblia Ram. ex DC. subsp, caussensis Casper a) MassifCentral (400-7eOm), France, b) Gorges du Ttm (4SOm), Pweyron, France. Kbndb & Shimai 5745-LPCGS (HIRO), R longijblia Rarn. ex DC. subsp, reichenbachiana (Schindler) Casper a) SE. France and Apennines (500-1 ,400rn), b) Vhllee de la Roya (SOOm), Alpes Maritimes, France. Kondo & S7timai S746-LPCGS (HIRO). maeroceras Link R a) Japan, Sakhalin lsl., Kamchatka to NW/ North America (e-3,OOOm), b) Tbdai-gawa (1,1OOm), Nagano, Japan. Kbndo & Shimai 5747-LPCGS (HIRO). R mundi Blanea, Jamilena, Ruiz-Rej6n et Zamera a) Albacete <1,200m), Spain, b) Nacimiento del Rie Mundo (1,200m), Albacete, Spain. Kbndo & Shimai 5748-LPCGS (HIRO). NII-Electronic Library Service The JapaneseSocietyJapanese Society for PlantPlantSystematics Systematics August 2006 KONDO & SHIMAI: Phylogeny in northern Pinguicula 157 TABLE 1. eontinued. Taxa Seuree & voucher specimen number R nevadensis (Lindbg.) Casper a) Sierra Nevada (2,400-3,OOOm), Spain, b) Sierra Neyada (3,OOOm), Granada, Spain. Konde & Shimai 5749-LPCGS (HIRO). R potdinii Steiger et Casper a) NE. Italy (500m), b) Nhl d'Arzine (SOOm), Friuli-Venezia Giulia, Italy. Kondb & Shimai 702-Cult.LPCGS. R vallisneriijbtia Webb a) S. Spain, b) Sierra de Segura Jaen, Spain. Kondo & Sliimai 5750-LPCGS (800m), (HIRO). R vuigaris L, a) N. North America te Eurepe, W. Russia and Morocco (O-2,50em), b) Velka Fatra (500m), Martin, Slovakia. Kbndo & Shimai 5751-LPCGS (HIRO). Other Pinguicuta Subgenuslsoloba Sectien lsoloba P lusitanica L. a) W, Europe and N, Aftica (O-500m), b) Breekcnhurst (30m), Hampshire, England. Kbndo & Shimai 5752-LPCGS (HIRO). SeetienCZirdiop1tyllam R cr),staUina Sibth. ex Sibth, et Smith a) Tr6odos (1,OeO-1,800m), Cyprus, b> Tr6edos (1,100m), Cyprus. Kbnde & Shimai 5753-LPCGS (HIRO). R hirt(flora Ten. a) Italy, Balkans and Turkey (O-2,OOOm), b) Vietri sul Mare (sea !evel), Carnpania, Italy. Kondo & S7iimai 5754-LPCGS (HIRO). Outgroup Genlisea violacea St.-Hil. a) Brazil, b) Itacambira, Mato Grosso, Brazil. Kbndo & Shimai 703-Cult,LPCGS. UtriculariaminorL, a) N. hemisphere, b) Higashi-Hiroshima, Hiroshima, Japan.Kbndo & Shimai 5755-LPCGS (HIRO). Classificationfo11ewedCasper(1966). a) Generat distribution and altitude range, b) Source of material used in this werk. LPCGS (HIRO)==Vbucher specimen plaeed in the Herbarium ef Hiroshirna University (HIRO). CuLt,LPCGS=Axenic culture strain number ofLaboratory ofPlant Chromoseme and Gene Stock, Graduate Scheol of Science,HiroshimaUniversity. species" distributed in Mexico to Central America, asexual reproduction (Casper 1962a, 1966, 1972), but mestly e'ndemic to Mexico. Some exceptional Thus, they have a clear and common seasonal species of the genus grow in the Caribbean and growth pattern in their habitats, They break dor- South America and thus, were non-categorized to mancy and start vegetative growth when the tem- any of the three groups. perature begins to rise in spring and then set flowers, The group of the northern species in Pingui- and finally form hibernacula for dormancy mostly cuia is not necessarily a particular taxonomical by the end of summer. The completion of this rank but could be defined as; 1) it is distributed in growth pattem takes 2-6 months. The majority ofthe the temperate to arctic or alpine zones in the north- northern species ofthe genus is distributed in Europe ern hemisphere; 2) it forrns hibernacula to resist and several species are distributed in Asia and North low temperature below OOC and dry season; and 3) America (Casper 1962a, 1966, 1972). It is uncertain it produces gemmae at the base ofhibermaculum for whether or not the similarity in the growth pattern in NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics l58 APG Nlo1. 57 TABLE 2, List efG+C contents and lengths ofthe ITS regiQns ofthe members oftbc northern 1'inguicula studied and their relations to chromosome numbers. Basicehromoseme Len hb G+C% number & Species ITS 1 ITS2 ploid levelChromosome number & authors R alpina 259 (73.36) 219 (72,60)8(4 × ) 2n=32 Doulat (1947), Sokolovskaja & Strelkova (196e), Casper (l962a, 1966) Rballeanita 254 × (68.5e) 220(69,55) 8(4 )8(8 2n=32 Casper (1966) Rbcthemica 256 (68.75) 218(69.72) × )8(2 2n=:64 Krahulcova & Jaoliova (1991) R carEi'ca 252 (64.68) 218(69.72) × )8(6 2n=16Contandriopoulos(1957) R clettosensis 254 (66.93)218 (69.72) × )8(4 2n= 48 Zamora et al. (1996) Riibith'RgrTaiidi27ana 254 (66,93) 218 (70.18) × ) 2n=32 [[lammaro & Pace (1987) smbsp. graiidilr7ar:a 254 (65,35) 219 (70.78) 8(4X) 2n= 32 Contaridriopoulos (1 957), Casper (1962a, 1966) RgrTaii{lit7anfi subsp.
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