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Page 1 植物研究雜誌 I. Jpn. Bot. 68: 8-22 (1993) Taxonomic Status Of 植物研究雑誌 J. J. Jpn. Bo t. 甜: 8-22 (1 993) Taxonomic Status of the East Asian Pyrola faurieana (Ericaceae) Erich HABER a and Hideki T AKAHASHl b aResearch aResearch Division ， Canadian Museurn of Nature ， P .0. Box 3443 ，Station D ，Ottawa ，KIP 6P4 CANADA; bBotanic bBotanic Garden ， Faculty of Agriculture ，Hokkaido University ，Sapporo ， 060 JAPAN 東アジア産カラフトイチヤクソウの分類 E. へイバーヘ高橋英樹 b a カナダ オタワ カナダ自然、博物館研究部 b 北海道大学農学部附属植物園 060 札幌市中央区北 3 条西 8 丁目 (Received (Received on August 28 ， 1992) The east Asian species Pyrola faurieana was investigated to clarify its taxonomic distinction from the the circurnboreal P. minor ， the Eurasian allotetraploid P. media and the North American hybrid P. asarifolia asarifolia x P. minor ， with which it has been confused ， and to evaluate its possible hybrid origin. In its its overa l1 morphology ， P. fi αurieana is most sim i1 ar to the hybrid P. αsα rifo /i αxP. minor ， but is distinguished distinguished by its rnore floriferous scapes ，shorter bracts and higher sepal length/width ratio ，its variable variable production of viable pollen ，seerningly norrnal seeds and distinctive leaf flavonol rnarkers. pyrola pyrola faurieana has the sarne leaf flavonol pattern as P. media and is sirnilar to it in pollen and seed rnorphology ， but differs in being diploid ，srnaller in stature and in having distinctively srnaller anthers ， generally generally rnore flowers ，shorter petals and a srna l1 er bract length/width ratio. P. faurieana is recognized as as a distinct species of hybrid origin with P. minor being one of the parents. The second parent is suspected suspected of being the Asian vicariad of P. asarifolia ，narnely ， subsp. incarnata (DC.) Haber et Takahash i. The first report of chrornosome nurnbers for P. faurieana is docurnented as 2n = 46. Introduction Introduction the two taxa (Kitamura et al. 1978 ，Yamazaki Andres (1 912) first described Pyrola faurieana 1981) ，but others only recognize P. faurieana in on the basis of a collection by Faurie made at Japan (Ohwi 1965 ， 1983). Korsakof ，Sakhalin 1s. It s specific status has been め'rola faurieana is also morphologically similar questioned ，however ，and the taxon confused with to the North Am erican hybrid P. αsαr ぴoU αXP. P. minor L. Both P. faurieana and P. minor were minor (Haber 1984) and to the Eurasian P. media recognized recognized by Hara (19~8 ， 1948) and distinguished Sw. ，a presumed amphidiploid (2n = 92) between on the basis of style ，anther ，and sepallengths ，and the circumboreal P. minor and the Eurasian P. sepal sepal and stigma shapes. Some authors of recent rotundifolia L. (Hagerup 1941). J apanese floras have maintained the distinction of A comparative study of P. faurieana ， P. media ， -8 ー February February 1993 Journal of Japanese Botany Vo l. 68 No. 1 9 P. P. asar ぴ， olia x P. minor ，and P. minor was under- A cluster analysis (average linkage) of morpho- taken taken to clarify the taxonomic status of P. logical characters for 77 specimens of P. faurieana fα urieana ，especially with regard to its possible was performed (with missing data for some speci- hybrid hybrid origin. As part of the evaluation of taxo- mens) to determine if any geographic differentia- nomic status for this taxon ， measurements of a tion was evident between the more northern number of morphological features were made and Kamchatka specimens (those closest to the range analyzed analyzed statistically. Pollen stainability and ab 田 of the morphologically similar P. α'sarifolia x P. normality normality were examined and seed set determined minor) and those further south ，especially on in in P. fauriean αwere compared with the other taxa. Hokkaido and adjacent islands. As weU ， data sets Chromosome counts were made to determine for P. faurie αnαandP.α 'sar ぴolia x P. minor w ぽ e ploidy ploidy level for P. faurie α na. Distributional ranges combined (108 specimens ，total) and analyzed to and habitat preferences of the four taxa were also determine whether any distinctive clustering and reviewed. reviewed. The possible relationship of P. as α， rifoli α separation of specimens could be found. Statistical Michaux subsp. inc αrnat α(DC.) Haber et analyses were performed on a personal computer Takahashi Takahashi to P. faurieana was also examined. using commercial software from Systat Inc. (Evanston ， IL). Materials Materials and methods In Pyrola ， as in the majority of the Ericaceae ， Specimens Specimens from the following herbaria were the four pollen grains derived from the s創 ne pollen used used for compiling measurement data ，distribu- mother cell are retained as a pollen tetrad at tional tional information and as sources of leaf material maturity. Pollen tetrad diameters and concurrent for for flavonol pattern analysis: BG ，C ，CAN ，DAO ， colpi lengths reported in a previous work GB ，GH ，KANA ，KYO ，MAK ，S ，SAPT ，TI ， (Takahashi 1986) for three of the four taxa under TNS ，TUS ，UPS ，Herbarium Ibaraki Univ. ， study here ，excluding P. αsa ベfolia x P. minor ， were Kurashiki Kurashiki Museum of Natural History ，and the evaluated in the context of the present study. Faculty Faculty of Agriculture of Kyushu University. Pollen stainability was determined by the toluidine Abbreviations Abbreviations follow the latest Index Her- blue-lactophenol method (Radford et al. 1974). bariorum; bariorum; for SAPT ，see “News and notes" 担 Pollen was also compared with the North Taxon 32: 703 (1983). American and Asian subspecies of P. α'sarifolia. Statistical Statistical data for the means of 11 morpho- Overall seed lengths and numbers of testa cells logical logical characters examined in this study were were determined under a light microscope for compiled compiled and summarized graphically. A signifi- samples of 20 seeds of P. faurieana (6 samples) ， cant cant difference was inferred in the character means P. media (3) and P. minor (10). Seeds were of of two taxa being compared when the confidence scattered on glass slides and sealed under a cover intervals intervals represented by twice the standard devia- glass with paraffin. tion tion of the mean on either side of the character Chromosome counts were made using rhizome means did not overlap (Hubbs and Hubbs 1953). tips and flower buds of P. faurieana fixed in acetic- Wh ere an overlap was evident ， the significance of alcohol (3: 1) and stored in 70 0/ 0 ethano l. Rh izome the the sample variances was judged by comparing the tips were treated apart in aceto-carmine ， squashed calculated calculated F statistics with published values at the under a cover slip ，and heated briefly under a .05 .05 level of significance. flame ， or were softened by heating for one hour 10 10 植物研究雑誌第68 巻第1号 平成 5 年 2 月 at at 37 0 C in 1N HCl ， then squashed. Young anthers and widest leaves and longest sepals ，petals ，anthers were were dissected from flower buds of about 2 mm and style. 乃/rola minor ， in contrast ，is significantly diameter diameter and teased apa 此 in aceto-carmine to free the smallest in blade width ，bract length and width ， pollen pollen mother cells. Slides were briefly heated and sepallength ，petallength ， anther length and style cells cells squashed under the cover slip. Chromosomes length. were were counted under oil immersion with the aid of Of the two taxa with which P. fauriean αcan a camera lucida drawing tube. Collections yielding be most readily confused ， P. media and P. counts counts are cited in the Appendix. asarifolia x P. minor ， it is most similar to the latter. The two-dimensional paper chromatographic The statistical summaries of the character means techniques techniques used in determining the leaf flavonol analyzed (Fig. 1) show no significant differences patterns patterns of herbarium specimens were those in the sample means of P. fi αurie αna and P. described described by Haber (1 983). As in previous studies ， asarifolia x P. minor in six of the 11 characters only only the ultra-violet-quenching compounds (scape h勾ht ， blade length and width ，bract width ， (appearing (appearing black under U.V よwhich serve as con- sepallength and style length). On the other hand ， sistent sistent and useful markers ， were assessed. Th e all eleven character means are significantly diι s担lI larity in flavonol spot patterns was judged on ferent between P. fauriean αand P. media. The the the basis of the average R f values of the com- means for only two characters (p etal length and pounds as well as their relative positions and con 国 anther length) are significantly distinct for all four centrations. centrations. Spot numbering in the patterns is taxa. 1n absolute measurement values ，however ， complementary complementary to that in earlier publications (e.g. ， there is a broad range in overlap of most characters Haber and Takahashi 1988). Chromatographed so that one must rely on a combination of specimens specimens of P. faurieana ， P. minor ， P. media and character measurements as well as other evidence P. αsa r. ぴo[i， αX P. minor are cited in the Appendix. to delimit the taxa. Some qualitative differen 切 S The distributional ranges for the four taxa in shapes are evident ， as in the narrower bracts of under under consideration are based on published data P. medi αand， P. minor ， as judged by the (Haber (Haber 1985 ， Hulten 1930) and on herbarium length/width ratio (l /w) of 3.1 for these taxa as collections. collections. About 800 and 600 specimens ，res 四 compared with those of 2.3 and 2.6 respectively pectively ， were measured for P. minor and P. for P. faurie αna and P. αsα rifo [i， αxP. minor. media for 11 selected characters (see Table 1). For The cluster analysis of P. faurieana from Japan ， the the geographically restricted P.
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