植物研究雑誌 J. J. Jpn. Bo t. 甜: 8-22 (1 993)

Taxonomic Status of the East Asian faurieana ()

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 ，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 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. faurieana ， 77 the adjacent islands and Kamchatka resulted in a specimens specimens were available and only 31 for the uniform spread of the Kamchatka specimens sporadically sporadically occurring North American hybrid P. (previous~ treated as P. medi αby， Hulten 1930) αsa r. ぴolia X P. minor. throughout the various subclusters. No grouping by geographical origin of specimens (i. e. ，Honshu ， Results Results Hokkaido ， Sakhalin Is. ， Kurile Is1ands and Morphological Morphological an αlyses The statistical data Kamchatka) was eviden t. summary ， as comp i1 ed in Table 1 and graphically The cluster analysis of the combined data sets portrayed portrayed in Figure 1， shows clearly that P. media for P. faurieana and the North Am erican hybrid is is overall the tallest and most robust of the four P. αsarifolia x P. minor resulted in a relatively taxa taxa compared. It has the tallest scapes ，longest intact separation of the North American hybrid m Oげ同広岡山、】還凶，

Table 1. Surnrnary of statistical data for eleven rnorphological characters in three species and one hybrid of Pyrola.

P.medi α P .f auriean α P.minor P.asarifoU αxP.minor 旬。ロ

N x sm N X S m N X S m N X sm Ehw-

司 d 0000000000002944697。 今 TI'ITi'A'iA 。-ーし『お Scape height (crn) 580 22.15 4.28 0.18 I斗 17 .4 9 3.69 0.4 3 ozJ 16.18 4.17 0.15 32d 16 .4 3 2.89 0.52 『 εJζJ B1 ade length (crn) 583 4.04 0.68 0.03 I 2.93 0.53 0.06 3.03 0.76 0.03 2.84 0.4 3 0.08 Hum520 『 ，、 今 Blade width (crn) 583 3.59 0.66 0.03 I 2.67 0.4 6 0.05 dOO 2.38 0.57 0.02 3 2.62 0.4 3 0.08 『 『 司 Nurnber of flowers 580 9.55 3.03 0.13 IroaIAUd 10 .4 6 3.19 0.36 11.28 3.60 0.13 d 7.03 2.24 0.4 0 回。 今 今 今 Bract Bract length* (rnrn) 247 5.50 0.96 0.06 4.81 0.80 0.10 3 3 4.4 1 0.91 0.05 3 5.74 0.68 0.12 EH 今， 今 今 斗『斗 3du 41d斗・ Bract Bract width * (rnrn) 199 1. 79 0.32 0.02 2.11 0.4 4 0.07 h 1. 42 0.4 9 0.03 2.21 0.54 0.11 岡山『〈。--。∞ 'i'i 司 21tiζus--- I I Sepal Sepal length (rnrn) 449 2.34 0.33 0.02 2.02 0.26 0.03 寸 1. 39 0.24 0.01 2.00 0.16 0.03 IξJ/oro『 『 d 今 Sepal Sepal width (rnrn) 449 1. 68 0.20 0.01 1. 61 0.18 0.02 I 品寸今 1. 56 0.22 0.01 3 1. 90 0.11 0.02

『 今 々 Petal Petal length (rnrn) 214 6.30 0.62 0.04 Ioofo 5.13 0.62 0.08 3ζud3 4.32 0.54 0.03 4 5.55 0.4 9 0.01 Z 『 勾 。・】 Anther length (rnrn) 472 1. 96 0.21 0.01 1. 64 0.15 0.02 10Y 1. 05 0.11 0.004 3 1. 52 0.20 0.04

今 Style Style length (rnrn) 352 4.91 0.79 0.04 3.67 0.63 0.08 斗 1. 49 0.33 0.02 3 3.92 0.67 0.12

Note: Note: Leaf rneasurernents were rnade on largest leaf ，and floral rneasurernents on lowest ， rnost rnature flower; N ， sarnple size; 支， rnean; s， standard deviation; sm' sm' standard deviation of the rnean; *bract rneasurernents are those of the inflorescence bracts.

...... 』司 12 12 植物研究雑誌第68 巻第1 号 平成 5 年 2 月

Scape Scape height (cm) Sepal length (mm) M8 -→ー ME ー+ー FR1 FR1 一一一←一一 FR →ー MI~ -← MI →- AxMI AxMI ーー一一+ーー一四 →- 16 16 17 18 19 20 21 22 1.3 1. 5 1. 7 1. 9 2.1 2.3 lade B lade length (cm) Sepal width (mm) ME ー+ー →ー FR FR 一ー+ー- ー-+一一 MI MI ー←- MI ー+ー AxMI -一←ーー AxMI 一一←司自 2.5 2.5 3.0 3.5 4.0 1. 5 1. 6 1 .7 1. 8 1. 9 Blade Blade width (cm) Petal length (mm) ME -l -← ME 吋ー FR~ ー+ー FR ーー←- MI -l -+- MI →ー AxMI ，ー→ーー AxMI 一ー←ー 2.5 2.5 3.0 3.5 4.0 4.0 5.0 6.0 Number Number of flowers Anther length (mm) MEi -ー ME -I + FR~ 一一←ー一 FR -I -+ MH -← MH ..j. AxMH AxM I1 -一←ー 7 8 9 10 11 12 1. 0 1. 2 1. 4 1. 6 1. 8 2.0 Inflorescence Inflorescence bract length (mm) Style length (mm) 山山内川 一ー→一ー + ー+・ MI MI + AxM1i AxM1i AxM'l →ー 4.4 4.4 4.6 4.8 5.0 5.2 5.4 5.6 5.8 1. 0 2.0 3.0 4.0 5.0

Inflorescence Inflorescence bract width (mm) 」 川内川洲 Mc pppyyyrrrdodaMa mhm 創出 いれ aer ー← 」 lir--t 円2 .no 、 a円 Rd 一一ー十ーー・ H X MIZ n 、』ー J 一→-ー 2s m A AxMI ← Pyrola asarifolia x P. minor 1.3 1.3 1. 5 1. 7 1. 9 2.1 2.3 2.5

Fig. Fig. 1. Graphical comparisons of 11 character means for four Pyrola taxa. Horizontallines represent represent 2x the standard deviation of the mean on either side of the means.

specimens specimens as several interconnected cluster groups. Pyrolafi αuriean αspecimens formed severallarger Five Five out of the 31 hybrid specimens available interconnected cluster subsets. Out of a total of 108 linked linked with adjacent P. faurieana clusters. Of the specimens in the cluster analysis ，a total of 8， or 77 P. faurieana specimens in the analysis ， three about 70/ 0 of the combined sample were “incor- were were included within the hybrid cluster groupings. rectly" aligned with specimens of the other taxon. Two of these three specimens were from Kam- Although the similarity between P. faurieana chatka ，a geographical region adjacent to the and P. asarifolia x P. minor is evident on general western western limit of the range of the North Am erican inspection of specimens and in the lack of signifi- hybrid hybrid in the Aleutian Islands (see Haber 1984). cant differences in the means of six characters ， the February February 1993 Journal of Japanese Botany Vo l. 68 No. 1 13 cluster cluster analysis indicates that the specimens of the this taxon from Japan ， Sakhalin Is. and the Kurile two taxa do exhibit sufficient differences overall Islands have a diameter range of 36-46 (57)μm that that they tend to associate as separate cluster sets. and a mean of 42μm (based on values in Table This This no doubt reflects the significant differences 1 of Takahashi (1986) and additional measure- in in the means for number of flowers ， bract length ， ments by Haber). The occasional collection con- sepal sepal width ，petallength and anther length evident ta¥ns up to 10 0/ 0 giant dyads. In the taxon as a in in Figure 1. whole ，most tetrads tend to be somewhat shrivelled Pollen Pollen tetrad diameters and stainability The and only take on a medium blue stain in lacto- presence presence of some genetic irregularity is evident in phenol and aniline blue. About 50% or more of the the holotype of P. faurieana and in other repre- the tetrads in some collections are highly shrivelled sentative sentative specimens. Tetrads in the holotype 紅 e and presumed abortive. imperfectly imperfectly formed and partly shrivelled with many Two collections at SAPT from M t. Makkari- exhibiting exhibiting a loose tetrahedral arrangement of the nupuri (Nishida 21 Aug 1906 and Konishi 28 Aug four four grains. The large tetrads are about 42μmin 1922) identified as P. fi αurie αna had completely diameter diameter and tend to stain light blue ， unlike the abortive pollen. However ，two other collections deep deep blue stain characteristic of normal plump from the same mountain (13 Aug 1986 ， Takahashi tetrads tetrads in other species. Large empty and small 6662 and 6664 ，SAPT) had only the more typically shrivelled shrivelled tetrads are also presen t. Specimens of incompletely expanded ，somewhat shrivelled

45 l

(目立) nu 門=←

』 4) ω 呂 町喧 十斗士 "0 ，て 3 35 35 。圃.~ 4) E叫 →干芋#

30 + • P. faurieana .A .A P. media ート • P. minor

~ 15 20 25 30 Concurrent Concurrent colpi length (μm)

Fig. Fig. 2. Bivariate plot of sample means and their confidence limits (2 x standard deviation o fo the mean on either side of the means) for concuηent colpi length and pollen tetrad diameter in in three Pyrola taxa. 14 14 植物研究雑誌第68 巻第1 号 平成 5 年 2月 appearance appearance commonly seen in pollen of P. disorder present in the species. This genetic faurieana. faurieana. A somewhat similar set of observations disorder may also be seen in some specimens of was noted for nearby sites in the Shiretoko P. medi αthat not only have malformed and poorly Peninsula ， Hokkaido. A collection from M t. stained tetrads but also have other floral abnor- Okkapake made by Takahashi and Kushibiki 5164 malities. A collection of P. media from the USSR ， (SAPT) ，had highly abortive pollen ， yet a second chromatographed for the present study (Province collection collection by Takahashi & Kushibiki 5157 (CA N)， Orel ，1 Ju l. 1906 ，CA N)， is abnormal in having from Futatsu-like several hundred meters away ， a flowering branch in the lower part of the raceme had mostly well-stained pollen tetrads ， but the in addition to having abnormal tetrads that are grains grains tended to be somewhat collapsed and the poorly stained. tetrads tetrads not always clearly tetrahedral in form. pyrola minor ， in contrast to the three other Pollen Pollen tetrads in P. media range in size from taxa ， has deeply stained ，fully expanded tetrads 37 -4 3μm with a mean of about 41μm. Data within 24 hours. These range in size from 27-39 published published by Takahashi (1986 ， Table 1) for this μm with a mean of 33ρm. Tetrads of P. species species and additional collections examined for the asarifolia x P. minor are highly malformed with present present study by Haber (citations in Appendix) collapsed walls and are stained weakly. The larger indicate indicate that pollen stainability ，and presumably somewhat collapsed tetrads tend to be 36-38μm viab i1i ty ，varies considerably. 恥10st collections of in diameter ， also stain weakly and are considered P. P. media have relatively large tetrads ， but these non-viable. Even after five years in stain ， the grain generally generally tend to be shrivelled to various degrees. contents remain weakly stained ， although some of Estimating Estimating the proportion of grains that might be the larger tetrads have grains with expanded walls ， considered considered to be relatively normal in their a high proportion of malformed tetrads remain. appearance appearance and stainab i1i ty is difficult here ， as in Pollen tetrad diameters are not significantly P. P. faurieana ， because of gradation between tetrads different between P. faurieana ('支 =42μm) andP. that that are somewhat collapsed ， but more or less well media (支 =41μm) ，and both have larger diameters stained ，and those that are progressively more than those of P. minor (支= 33 pm ，Fig. 2). Pyrola collapsed ，emptyor weakly stained. Kn aben (1 944) fα urie αna tetrads are significantly larger than the estimated estimated that pollen viability was reduced to about highly shrivelled and abortive tetrads of the North 23 23 0/ 0. A collection such as that made by H. An dres American hybrid P. asarifolia x P. minor. The in in Germany (Koln ，Aug. 1929 ，CAN) has darkly highly abortive pollen of the Okkapake collection stained stained and mainly fully expanded tetrads ， yet even of P. fauriean ι(Takahashi and Kushibikii 5164) ， here here particularly large ，4-celled tetrads ， about 45 has a mean diameter of about 39μm ， for the more μm in diameter ， are present and in non-tetrahedral fully formed tetrads ， with a broad range in formation. formation. In a British collection (Ambleside ， 24 diameters being present. Al though there is a broad Jun. Jun. 1869 ，CA N) in which the tetrads stain dark overlap in the absolute lengths of concurrent colpi blue ，and are of normal size ，most tetrads are in P. faurieana ('支 =25 pm) ， P. media ('支 =20μm) highly highly shrivelled. Age of pollen samples may affect and P. minor (支= 17μm) ，a small but significant the the ability to rehydrate ， but the collapsed nature statistical difference exists in the means (Fig. 2). of of tetrads ， even after rehydration in stain for over Exine sculpturing in P. faurieana ， P. media and a year ，is more likely a reflection of the genetic P. minor is verrucate (Takahashi 1986). February February 1993 Journal of Japanese Botany Vo l. 68 No. 1 15

Table Table 2. Summary of statistical data for two seed characters of three species of Pyrola.

Sample Sample Seed length (mm) Testa cell number

x sm X sm

P.fi αurie αnα 0.71 0.07 0.03 7.25 0.79 0.35 2 0.69 0.05 0.02 7.35 0.75 0.34 3 0.64 0.05 0.02 7.60 0.75 0.34 4 0.67 0.04 0.02 7.20 0.95 0.4 2 5 0.64 0.05 0.02 7.30 1. 08 0.4 8 6 0.56 0.05 0.02 7.00 0.92 0.4 1 P. P. media 0.73 0.06 0.03 6.85 1. 14 0.51 2 0.68 0.05 0.02 7.30 1. 22 0.55 3 0.69 0.05 0.02 7.15 1. 18 0.53 P. P. minor 0.4 4 0.05 0.02 7.30 1. 30 0.58 2 0.55 0.05 0.02 7.80 0.89 0.4 0 3 0.65 0.05 0.02 9.25 1. 33 0.59 4 0.62 0.04 0.02 8.80 0.95 0.4 2 5 0 .4 8 0.03 0.01 7.60 1. 10 0.4 9 6 0.63 0.05 0.02 8.70 1. 53 0.68 7 0.68 0.04 0.02 8.85 1. 35 0.65 8 0.64 0.06 0.03 8.35 1. 09 0.4 9 9 0.59 0.05 0.02 9.35 1. 09 0.4 9 10 10 0.60 0.05 0.02 8.80 1. 11 0.50

Seed Seed set and size pyrola seeds are minute ， faurieana from 11 metaphase plates in rhizome tips consisting consisting of a small nodule of cells comprising of collections by Takahashi (5249) and Takahashi 由 e undifferentiated embryo ， centered within an and Kushibiki (5157). A haploid number of about elongate ，thin-walled testa. A bivariate graph of 23 chromosomes was counted in 12 plates of a testa testa cell number and seed length of P. media ， P. collection of the same taxon made by Takahashi faurie αna and P. mino 九 based on values in Table and Uematsu 8504 at Hamatonbetsu ， 12 J凹1. 1988. 2 ，is given in Fig. 3. There are no sharp distinc- Perhaps partly because the preserved material was tIons tIons in seed size among the three taxa ， although several years old ，chromosome separation and P. P. media and P. faurieana tend to have somewhat staining was not optima l. Chromosome sizes larger larger seeds than those of P. minor. They also tend ranged between 2.5-5.0μm long. to to have fewer testa cells per length than P. minor. Although chromosome counts were not in- Pyrola Pyrola minor ， in contrast ， has a broader range in dividually conclusive ， they were consistent in seed seed length and testa cell numbers ， with increased magnitude and sufficient in number of counts to seed seed length being correlated with increased testa indicate that the diploid number is probably cell cell numbe r. Capsules of P. asarifolia x P. minor 2n = 46 ， the same as has generally been reported bear bear only aborted ovules ， a reflection of the highly in other diploid species of the genus. This is abortive abortive pollen produced by the hybrid. also supported by the haploid counts. One somatic Chromosome counts Diploid counts of plate was seen that may have had a tetraploid approximately approximately 40 chromosomes were made for P. number. 16 16 植物研究雑誌第68 巻第1 号 平成 5 年 2月

0.8 0.8

0.7 0.7

言 0.6 缶 i 、、 J 7 4 ~ 7 -回圃. 営 0.5 0 F 司句。 ∞ω 0 .4 • P. faurieana ....P. ....P. media • P. minor L → 7 8 9 10 Testa cell number

Fig. Fig. 3. Bivariate plot of sample means and their confidence limits (2 (2 x standard deviation of the mean on either side of the means) for for seed testa cell number and seed length in three Pyrola taxa.

Chrom αtographic patterns of leaf flavonols 持1 and in having 非8.

The 2・dimensional chromatographic patterns of Collections of P. faurieana with pollen that was leaf leaf f1 avonols for P. faurieana and the three other highly abortive (Konishi ，28 Aug. 1922; Nishida ， taxa taxa with which it was compared are given in Fig. 21 Aug 1906; Takahashi and Kushibiki 5164 ，and 4. 4. Based on the previous work with other species Yoshimura and Yokoyama 5 Aug 1938) did not of Pyrola (Haber 1983) ， these UV-quenching differ from the f1 avonol pattern present in collec- flavonols flavonols are all glycosides of quercetin (#1-5 ， 8) tions with more normal levels of pollen viability. and kaempferol (尚). Simila r1 y， the holotype ， with its highly collapsed The f1 avonol pattern of Pyrola fauriean αis pollen ， also did not show any deviation in its identical identical to that of P. media. Pyrola faurieana flavonol pattern from that of other specimens of differs differs from the North American hybrid P. this taxon with higher levels of nearly normal asar ぴ'o Uαx P. minor by having a pair of over- pollen. lapping lapping glycosides (何 and #8) and in lacking Distribution The worldwide distributions for glycoside 再1 of the hybrid. The absence of f1 avonol P. media ， P. minor ， P. faurie αna and P. # 5 in P. asar. ぴolia x P. minor is unusual because asarifoliaxp. minor are given in Fig. 5. Pyrola it it is present in both of its parents. A second minor is a widespread circumboreal species ，found flavonol flavonol (打)， found in P. asarifolia (see Haber primarily in mossy coniferous forests but also in 1983) 1983) is also absen t. This hybrid seemingly lacks mixed woodland sites and deciduous forests in the the full complement of parental f1 avonols. Pyrola northern Sweden and J apan. It is found over a faurieana faurieana differs from P. minor in lacking f1 avonol wide elevational range from near sea level to alpine February February 1993 Journal of Japanese Botany Vo l. 68 No. 1 17

A B 2・D 2・D

0.5 0.5 s 0.5 ;.1 1 4 ~21

1・D 0.5 1・D ・ 0.5

Pyro la as 官官， blla x Pyr ola mlnor Pyro la mlnor

C 2・D 、D 2・D

8 8 s.2 s.2 0.5 0.5 6'. 6'. 6，・ ‘•s 2 1岨 D 0.5 1・D 0.5

Pyro la medla Pyr ola faurfeana

Fig. Fig. 4. Two-dimensional chromatographic patterns of UV-quenching leaf flavonols in four pyrola pyrola taxa. meadows. In the more southerly extensions of its range. It occurs most commonly in J apan range range in the Rocky Mountains of Colorado ，it (northern Honshu and Hokkaido). However ，it is reaches reaches elevations of 3600 m and in SE Tibet 4α 旧 sporadic and considered rare. Northward it is m. pyrola minor is rare in northern Japan. found on Sakhalin Is. ， where the type was col- pyrola pyrola media is Eurasian with a relatively lected ，on the Kuriles ， southern Kamchatka and scattered scattered distribution within its overall broad Bering Is. It is a of higher elevations ， range. range. Like P. minor ，its preferred habitats are occurring in the Pinus pumila (p all.) Regel scrub mossy coniferous woodlands. It occurs near sea zone and alpine meadows in northern Japan ， level level but is more commonly found in mountainous associated with such species as Empetrum nigrum terrain terrain reaching the upper subalpine meadows. 1n L. and Tilingia ajanensis Rege l. Occasionally it is the the Caucasus Mountains ， it reaches alpine found with P. minor and P. al1 フ的 αH. Andres.

meadows at about 2800 m. pyrola medi αextends， Specific site localities for the North Am erican eastwards eastwards in Siberia to about Lake Baika l. hybridP.α sar ぴ'o[i， αxP. minor ， as plotted in Figure pyrola pyrola faurieana has a relatively restricted total 5， have been updated from those first published 18 18 植物研究雑誌第68 巻第1 号 平成 5 年 2 月

Fig. Fig. 5. Worldwide distributions of the four Pyrola taxa studied. by Haber (1984). Three new records are included other species of Pyrola ， including those in this for for Al berta ， Colorado and Utah. One record for study ， are short ，ぽ ect or spreading cylindricallobes Colorado Colorado plotted by Haber (1984) but not cited in projecting beyond the reflexed stylar tissue that that that publication is also cited in the Appendix. Thi s forms a collar below the lobes (see Haber 1984). hybrid hybrid has been regularly collected with one or The short ，1 mm long ， anther sacs of P. minor ， both both of its parental species. with their truncated ends terminating in large pores The distributional ranges of P. faurieana and the width of the sacs and the relatively slender P. αsarifoli αX P. minor are virtually contiguous ， filaments are also distinctive features of the species. meeting meeting at the western end of the Al eutian Is1ands pyrola minor is a diploid species (2n = 46) with (Attu (Attu Is.) and the Commander Islands off the east highly fertile ，well-stained and developed pollen coast coast of Kamchatka. tetrads. At the other extreme is P. media ， a tetraploid Discussion Discussion and taxonomic conclusions (2n = 92) ，and in most features ， the largest of the Of the four taxa compared in the present study ， four taxa. 1ts white ，occasionally pink-tinged ， P. P. minor has the most distinct morphology. It globular flowers have a slightly exsert ，straight to tends tends to be the smallest of the four ， both somewhat declinate style. The comparatively large vegetatively vegetatively and in its floral features. It is readily anther sacs ， mainly about 2 mm long and with differentiated differentiated by its small ， white to pink-tinged ， relatively large pores ， serve as the most drooping ， actinomorphic flowers that possess a characteristic morphological feature ，together with short ，straight insert style. It s stigma consists of the generally robust size ，distinguishing this species five five radially arched crests that become everted from from the others under consideration. It s pollen within within the stylar tip as the outer tissue of the tip tetrads are substantially larger than those of P. reflexes reflexes on maturation. The stigmatic crests in minor and vary considerably in their stainab i1i ty February February 1993 Journal of Japanese Botany Vo l. 68 No. 1 19 and degree of collapse of grains. incarnata (DC.) Haber et Takahash i. This

pyrola pyrola faurieana is similar in size and mor 田 subspecies of P. asarifolia is found in northern phology phology to the North American hybrid P. asari- Japan ， the Kuriles ，central Kamchatka and eastern folil αx P. minor. The two are alike in general habit ， continental Asia (see Fig. 1，Haber and Takahashi their their pink-tinged globular flowers and possession 1988) and overlaps the ranges of both P. minor of of attributes that are associated with hybrids in and P. fi αurie α na. However ， subsp. incarn α ta which which P. minor is one of the parents. These presently tends to occur in northern Japan only at attributes attributes include ，most specifically ，styles that are lower elevations in broad-leaved deciduous forests slightly slightly exsert and straight ， or nearly so ，and and open Larix forests. Both P. as αr， ぴ'o lia subsp. anthers anthers that have large pores the full width of the incarnata and P. minor may be found in closer truncated truncated or only slightly elongated ends of the saω proximity in the Kuriles and Kamchatka. Sub- (see (see illustrations in Haber 1984 ， 1987). Hybrid species α'sarifolia and subsp. incarnata differ specimens specimens with such styles and anthers invariably primarily in the narrower sepals of subsp. in- have have high levels of pollen irregularity and poor carnata and in the leaf flavonol pattern of this staining. staining. 1n sterile hybrids ，common abnormalities subspecies that resembles more closely that of the such such as split styles or bifid sepal apices or floral Eurasian P. rotundifolia (Haber and Takahashi branches branches in the raceme are also found. 1988). Pyrola fi αurieana ，like P. αsarifolia subsp. The possibility of a hybrid origin for P. inc αrn αtα ， has a sepal width mean that is fauriean αhas never been suggested ， although significantly narrower than that of P. Takahashi Takahashi (1986) had indicated the presence of asarifolia x P. minor. high high levels of incomplete expansion of tetrads in Pyrola faurieana is also similar to P. media in most most collections and in some cases up to 10% giant general morphology and flavonol pattern. dyad dyad formation. Additional examination of pollen However ，this is interpreted to be a reflection of for for the present study has confirmed this ，and as its past origin as yet another hybrid with P. minor well ，several collections with completely abortive as one of its parents. The similar size of pollen pollen pollen were found. tetrads in these two argues taxa against their being

We conclude that P. faurieana is of hybrid different ploidal variants of the s副 ne taxon. Pyrola origin origin with P. minor being one of the parents. fauriean αis maintained here as a distinct species Unlike the completely sterile hybrid P. of hybrid origin with P. minor as one of its asarifolia asarifolia x P. minor ， P. faurieana has reduced and parents. It is morphologically most sim i1 ar to variable variable pollen viability but produces seemingly pyrola asarifolia x P. minor ， produces seemingly normal normal seeds. As past attempts to propagate normal seeds ，and has reduced pollen viability. from Pyrola seeds have not been successful Diagnostic key and synoptic comparison of the (Christoph (Christoph 1921 ，LihnellI942) ， the viability of P. four taxa studied Measurement ranges are not faurieana faurieana seeds is difficult to establish. absolute minimumlmaximum values but represent 1n 1n view of the morphological similarity of P. values for about 70070 of specimens as determined faurie αna to P. asarifolia x P. minor and the by the means + 1- the standard deviation. contiguous contiguous nature of their ranges ， the second 1 Style insert ，straight ， about 1.5 mm long ， stigma parent parent of P. faurieana is suspected of being the peltate with 5 radially spread ridges; anthers Asian Asian vicariad of P. asarifolia ，namely ， subsp. mainly about 1 mm long and plump ， ends of 20 20 植物研究雑誌第68 巻第1 号 平成 5 年 2 月

sacs sacs without tubes ， truncate and open at fo 口ning basal rose 社 e representing l-several years dehiscence; dehiscence; pollen highly fertile ，tetrad diameter growth; petioles commonly shorter or subequal to about about 33μm; 2n = 46 ...... P. minor blades ，occasionally longer; blades ovate to elliptic ， l' l' Style exsert ，straight to somewhat declinate ， or orbicular ，occasionally obovate ， 1-4.5 cm averaging averaging 3-6 mm long ， stigma of 5 erect to long ， 1.5-4.5 cm wide; apex rounded or divergent divergent cylindrical lobes projecting from a mucronulate; base rounded or shallowly cordate reflexed reflexed collar ， anthers mainly over 1. 5 mm and commonly abruptly decurrent; margins entire long long and elongate ，sacs constricted at ends into to crenulate. Scapes 10-26 cm tall ， with several short short tubes with conspicuous pores; pollen ovate bud scales at base and l-several scales

with with variable viability ， tetrad diameter over below raceme. Raceme symmetric ， 4-18 聞 36μm ...... 2 flowered; bracts ovate ，3-7 mm long ， 1-3 mm 2 An thers mainly 1.8-2.2 mm long; petals wide ，commonly longer than pedicels. Flowers ， 5.5-7.0 5.5-7.0 mm long; inflorescence bracts globular ，radially symmetric ，drooping ， white to 1.5-2 1.5-2 mm wide; pollen only about 30070 pink ，6-8 mm across. Sepals elliptic to broadly normal ，tetrad diameter averaging 41μm; ovate ，1 .5 -2 .5 mm long ， 1.3-2.0 mm wide; apex plants plants mainly 18-26 cm tall; 2n = 92 . blunt to subacute; bases generally overlapping; P.medi α margins entire to erose. Petals 4-7 mm long ， 2' 2' An thers mainly 1. 3- 1. 8 mm long; petals 3.3-4.7 mm wide. Stamens included ， uniformly 4.5-6.0 4.5-6.0 mm long; inflorescence bracts positioned around pistil; filaments narrow ， 1.7-2.7 1.7-2.7 mm wide; pollen viab i1i ty variable ， 2.5-5.0 mm long. Anthers 1. 4-2.0 mm long ， tetrad tetrad diameter variable; 2n = 46 ...... 3 creamy white to tan ，oblong ， without clearly de- 3 Sepals mainly 1.8-2.0 mm wide; pollen fined tubes; pores relatively large ， at ends of the sterile sterile with tetrads highly malformed ， narrowed sacs. Pollen tetrads 35 --4 6μm diameter ，

largest largest tetrads about 36-38μm in 支=42ρm. Style straight to somewhat declinate ， diameter; diameter; 2n = 46(?) .. barely exserted ， 2.0-4.8 mm long; stigma with . P. as αr ぴ'ou， αxP. minor conspicuous reflexed collar and 5 divergent ，cylind- 3' 3' Sepals mainly 1. 4- 1. 8 mm wide; pollen ric lobes 0.5 mm long. Capsule pendent ，5-10cular ， generally generally with perhaps 50070 nearly 5-6 mm diameter; dehisced valves connected by normal normal tetrads ，diameter variable ，36 --4 6 fibers. Chromosome number 2n = 46. Mainly (56)μm ， average about 42μm; 2n=46 alpine meadows and sometimes edges of open . P. faurieana subalpine deciduous broad-leaved forests of N The following is an updated description of P. Honshu ，Hokkaido ，Kuriles ，Sakhalin ，Kamchatka fα urie α na. and Bering Is.; 10-2000 m elevation. Pyrola Pyrola faurieana H. Andres ，Verhand l. Bo t. Ve r. Prov. Brandenburg 54: 218-227 (1 912) - The first author would to like thank research Pyrola Pyrola yezoensis (in sched.) - P. minor L. sensu assistant Laurie Consaul and museum volunteer some Jap. authors. - TYPE: Sakhalin Is. ， Robert Couch for their assistance ，partic 叫ar1 y with Korsakof ， Faurie 611 (holotype: E!) data analysis and mapping. The second author is Perennial ，scapose ，evergreen herb ，clonal from indebted to curators at BG ，C ，GB ，S and UPS for spreading rhizomes. rhizomes. spreading Leaves several ，approximate ， permission to measure herbarium specimens. The February February 1993 Journal of Japanese Botany Vo l. 68 No. 1 21

visit visit to Scandinavian herbaria (1 989-1990) was pyrolafaurieana (citations are provided for all records seen of of this taxon because of the common confusion with P. made possible through a grant from the exchange minor; + indicates specimen was chromatographed; *pollen program between the Japanese Society for the was examined; ! chromosome counts were made). JAPAN. HOKKAIDO. Abashiri Province: Mt. Io-zan Promotion of Science and the Royal Swedish (=Iwo) ， 18 Ju l. 1951 ，Misumi et al. 41172 (SAPT); Mt. Academy of Sciences. Shiretoko ，900-1050 m ， 19 Ju l. 1952 ，Samejima et al. (SAPT). (SAPT). Shari ， 15 Aug. 1965 ， Shiota and Yoshitake (Herb. Ibaraki Ibaraki Univ.) ，M t. Kikin ， 21 Ju l. 1974 ， Matsuki (MAK); Appendix of Specimen Citations M t. Shari ， 1500 m ， 12 Aug. 1983 ， Takita 1669 (KYO); + *! Pyrola Pyrola asarifolia x P. minor (+ indicates specimen was Shiretoko Pen. ，Futatsu-ike ， 1300-1400 m ， 12 Aug. 1984 ， chromatographed; chromatographed; *pollen was examined). CANAD A. Takahashi and Kushibiki 5157 (SAPT ，CA N); + *Shiretoko Alberta: Alberta: *Poboktan Creek ， 26 Ju l. 1908 ，Brown 1351 ， Pen. ，M t. Okkapake ， 12 Aug. 1984 ， Takahashi and (C AN); Moraine Lake ， Banff National Park ， 11 Aug. 1964 ， Kushibiki 5164 (SAPT). Hidaka Province: M t. Poroshiri Scoggan Scoggan 16314 (CAN). British Columbia: + *Sustut L.， 23 (= Porojir i)， 10 Ju l. 1929 ，Tokunaga and Sakamoto Ju l. 1977 ，Gillett and Boudreau 17401b (CA N); + Peak L. ， (SAPT); +M 1. Tottabetsu ，Kar ，9 Sep. 1933 ，Hoshino

2 Aug. 1977 ，Gillett and Boudreau 17711b (CA N). North ‘ (SAPT); M 1. Poroshiri ， 1600-1700 m ， 26 Ju l. 1971 ， west west Territories: *Brintnell L.， Red 恥1tn. ，7 Aug. 1939 ， Koyama et al. 3906 (TNS); 14 Aug. 1979 ，Umezawa (TI) ， Raup and Soper 9737 (CAN). Yukon: + *恥 11. Sheldon ， 13 18 ∞m ，1 Aug. 1983 ， Takahashi 4588 (SAP T); M t. Chiroro ， Aug. 1944 ， Porsild and Breitung 11092 (CAN). USA. 1710-1848 m ，7 Aug. 1985 ， Takahashi et al. 5865 (SAPT). Alaska: Alaska: Nome ，4-8 Aug. 1945 ，Scamman 3897 (GH); Iburi Province: + *Mt. Yotei ・zan =( Makkarinupuri) ， 21 Attu Attu Is. ， Massacre Bay ，1 Sep. 1945 ， Schaack 973 (GH). Aug. 1906 ， Nishida (SAPT) ，+ *28 Aug. 1922 ， Konishi Colorado: Colorado: Marshall Pass ， 27 Ju l. 1896 ， Shear 5146 (NY); (SAPT) ， 1610 m ， 13 Aug. 1986 ， Takahashi 6662 and 6664 Cottonwood Pass ， 14 Aug. 1971 ， Childers and Pinkava (SAPT). Ishikari Province: + *M t. Yubari (= Yuparo ， 8952 8952 (ASU). Utah: 24 km E of Cedar City ， 12 Ju l. 1940 ， Yupari) ，8 Aug. 1912 ，Yanagisawa (SAPT) ，7-9 Aug. Maguire Maguire 19464 (CAN). 1913 ， Nishida (SAPT); M t. Tokachi ，Aug. 1915 ， Koidzumi (TI); (TI); M t. Yubari ，Aug. 1916 ，Koidzumi (TI) ， 22 Ju l. 1933 ， Pyrola Pyrola media (+ specimens used for chromatographic Hara (TI) ，Aug. 1938 ，Honda (TI); + *M t. Ishikari ，8 Aug. analyses; analyses; *pollen evaluation) FINLAND. + Jaaski ，9 Ju l. 1952 ，Samejima and Misumi (SAPT); M 1. Yubari ， 1380 m ， 1927 ， Valle s.n. (DAO); + Nylandia ， 26 Ju l. 1952 ， 29 J叫.1987 ， Takahashi 7576 (SAPT) ， 1400 m ，7 Aug. 1987 ， Marklund s.n. (DAO); + Etela-Savo ，6 Ju l. 1983 ， Ulvinen Yamazaki (TI) ，Uno 16820 and 17231 (Herb. Kurashiki s.n. s.n. (CA N). GER 恥1A NY. + *Koln ，Aug. 1929 ， Andres s.n. Mus.). Kushiro Province: M t. Meakan ，Aug. 1893 ， (CAN). GREAT BRITAIN. *Ambleside ，Lus(?) ， 24 Jun. Fujimura (SAPT) ，Aug. 1897 ，Kawakami (SAPT) ，2 Aug. 1869 1869 (CA N). SWEDEN. + Sjobottom ，9 Ju l. 1964 ，Porsild 1937 ，Yokoyama 4273 (SAPT); M t. Mashu ，4 Sep. 1954 ， 23296 23296 (CAN). USSR. + *Province Orel ，District Brjansk ， Okamoto 1166 (KYO); Teshikaga ，Pekere ， 13 Ju l. 1959 ， 1 Ju 1. 1906 ，Protopopov ，Rudnev and Chitrovo s.n. (CA N). Kimura (TNS); M t. Oakan ， 1250 m ，7 Aug. 1981 ， Takita 678 678 (KYO). Soya Province: Rebun Is. ，5 Ju l. 1920 ， pyrola pyrola minor (all specimens used for chromatographic Tatewaki 19699 (SAPT) ， 200 m ， 28 Jun. 1983 ， Takahashi analyses) analyses) CANAD A. Alberta: Cypress Hills ， 16 Aug. 1947 ， 4320 (SAPT) ，400-490 m ， 18 Aug. 1984 ， Takahashi 5241 Breitung Breitung 5658 (DAO). British Columbia: 40 m i. W of (SAPT); Wakkanai ，Noshappu-misaki ， 22 Ju l. 1956 ，Hara Anahim Lake ， 11 Jun. 1956 ，Calder ， Parmelee and Taylor (TI); Rishiri Is. ， 800 m ， 19 Ju l. 1982 ， Takahashi 2802 17321 17321 (DAO); 10 m i. N. of Bull R. ， 13 Ju l. 1958 ， Taylor (SAP T) ，+! 930 m ， 20 Aug. 1984 ， Takahashi 5249 (SAP T); and and Ferguson 2730 (DAO). Manitoba: Reindeer L.， 10 Aug. !Hamatonbetsu ，Beniya-genseikaen ， 12 Jun. 1988 ， 1951 ， Baldwin 2397 (CA N). Newfoundland: Gander ，7 Ju l. Takahashi and Uematsu 8504 (SAPT). Tokachi Province: 1949 ，Bassett 434 (DAO). Ontario: Jackfish ， 19 Ju l. 1937 ， M t. Saoro (= Sahoro) ， 13 Aug. 1913 ， Nishida (SAPT); M t. Hosie ， Losee and Bannan 2397 (CAN). Quebec: Corner of Satsunaitake ， 24 Ju l. 1917 ， Nishida (SAPT). HONSHU. the the Beach ， 15 Ju l. 1946 ，Terrill4466 (CAN). Saskatchewan: Akita Prefecture: M t. Eboshi ， 16 Ju l. 1958 ， Kikuchi (TNS). Cypress Cypress Hills Park ， 22 Ju l. 1947 ， Breitung 4962 (DAO). Aomori Prefecture: M t. Sukayu ， 27 Aug. 1897 ， Hiratsuka Yukon: mile 95 ，Canol Rd. ， 11 Ju l. 1944 ， Porsild and (SAPT). Iwate Prefecture: M t. Iwate-san ，Aug. 1905 ， Breitung Breitung 10398 (CAN). FINLAND. Lake Oulunjarvi ， 15 Sakurai (TNS). Miyagi Prefecture: M t. Katta ， 14 Ju l. 1936 ， Ju l. 1969 ，Alho and Laine s.n. (DAO). FRANCE. Hara (TI); Daikokuten ， 1500 m ，4 Ju l. 1950 ，Kimura and Clermont-Ferand ， 15 Ju l. 1960 ，Lugagne 3923 (DAO). Sugaya (TUS); M t. Sugigamine ， 1720 m ， 24 Ju l. 1977 ，Ueno GERMANY. Wissental ， 30 Jun. 1933 ，Schumacher s.n. and Suzuki 20182 (TUS). Yamagata Prefecture: M t. (DAO). GREENLAND. Nigerdleq ，8 Ju l. 1966 ， Jorgensen Azuma-san ，Aug. 1927 ，Yuhki (KYO). USSR. KURILE 66-168 (CAN). JAPAN. Rishiri Is. ，20 Aug. 1984 ， ISLANDS: Shumushu Is. ，Bettobi ，2 Aug. 1943 ，Akiyama Takahashi Takahashi 5248 (CAN). SCOTLAND. Bl ack Loch ， Jun. (KANA); Paramushir Is. ，Urafutoyama ，8 Aug. 1920 ，Kudo 1959 ，Lamond 3062 (DAO). SWEDEN. Angermanland ， 28 (TUS) ， Ziguro (?)， Kozima (KYO); Matsuwa Is. ， Jun. Jun. 1921 ， Samuelsson 1265 (CAN); Jukkasjarvi ， 22 Aug. Yamamotowan-Ainuwan ，7 Aug. 1928 ， Tatewaki and 1949 ，Asplund s.n. (DAO). Tokunaga 12160 (SAP T); Shimshir Is. =( Simushir) ， Broton 22 22 植物研究雑誌第68 巻第1 号 平成 5 年 2 月

Bay ， 13 Aug. 1928 ， Tatewaki and Tokunaga 11613 (SAP T)， Hara H. 1938. Pyrola faurieana H. Andres. J. Jpn. Bo t. Yamagoshizaki ， 16 Aug. 1928 ， Tatewaki and Tokunaga 14: 427-428 (in Japanese). 11797 11797 (SAPT); Etorofu 1s. ，Shibetoro ， 30 Ju l. 1884 ，恥1i yabe 一一一一 1948. Enumeratio Spermatophytarum Japoni- (SAPT) ， 19 Ju l. 1906 ，Miura (SAPT) ， 21 Ju l. 1906 ，Miura carum. 1. 1wanami Shoten ， Tokyo. (SAPT) ，Moyoro-Shibetoro ，5 Aug. 1938 ，Yoshimura and Hubbs C. L. and Hubbs C. 1953. An improved graphical Yokoyama (SAPT); Kunashiri 1s. ， M t. Chacha ， 26 Ju l. analysis and comparison of series of samples. Sys t. 1929 ， Nagai and Shimamura (SAPT) ，1 Aug. 1929 ，Okada Zoo 1. 2: 49-56 ， 92. (TNS) ， M t. Tomari ， 20 Aug. 1936 ， Tatewaki 25520 (SAPT). Hulten E. 1930. Flora of Kamtchatka and the adjacent SAKHALIN: Kaibato Is. ， M t. Dainan ， 28 Ju l. 1931 ， islands. vo l. 4. Kong 1. Svenska Vetenskapsakad. Hand 1. Kimoto et al. (SAPT); Precise locality unknown ，4 Aug. se r. 3. 8(2). Almqvist & Wiksells ， Stockholm. 1910 1910 (Tl); Shikka (= Sikka-gun): M t. Asase-yama ，9 Aug. Kitamura S. ，Murata G. and Hori M. 1978. Coloured 1935 ，Sugawara (SAPT); Chirikoro ・daiichisiryu ，2 Aug. illustration of herbaceous plants of Japan 1. 1935 ，Sugawara (SAPT); M t. Kawashima ，9 Aug. 1936 ， (Sympetalae) ，revised ed. ，Hoikusha Pub l. Co. Lt d. ， Yoshimura (SAPT); Horonoberenzan ，Mimizuku ， Osaka (in Japanese). Nagamatsu (KYO); Hoshi (?)， Sugawara (KYO). W. Coas t: Knaben G. 1944. Studier over norske Pyrola-arte r. Arb. Anbetsu ， 16 Ju l. 1927 ， 1shiyama (SAPT). Motodomari: M t. Univ. Bergen Ma t. Naturvitensk. Se r. 1943(6): 1-18. Kashipo ，7 Aug. 1928 ， Hiratsuka (SAPT); M t. Tossozan ， Li hnell D. 1942. Keimungsversuche mit Pyrolasamen. Kume 2597 ，Uno 21226 (Herb Kurashiki Mus.). Ootomari Symb. Bo t. Upsa l. 6(3): 1-37. (= Ohdomari ，Korsakof f): Sakaehama ， 22 Ju l. 1937 ， Ohwi J. 1965. Flora of Japan. (F. G. Meyer and E. H. Yoshimura and Hara (SAPT) ，Otogiri (KYO); Ohdomari ， Walker ，eds ふ Smithsonian 1ns t.， Washington ，D.C. 28 28 Ju l. 1936 ，Hara (S); Takadai ， 22 Ju l. 1937 ，Yoshimura 一一一一一 1983. New Flora of Japan. (revised ed. ，by M. and Hara 132 (TNS); 16 Ju l. 1907 ，Miyake (SAPT) ， 16 Ju l. Kitagawa) ， Shibundo Pub l. Co. Lt d. ，Tokyo (in 1907 ，Miyake (TNS) ， 12 Ju l. 1906 ，Miyabe and Miyagi Japanese). (SAPT) ，8 Aug. 1923 ，Sawada (TI) ，Aug. 1924 ， Numajiri Radford A. E.， Dickison W. C. ，Massey J. R. and Bell R. (TNS) ， 31 Ju l. 1933 ，Akiyama (KANA) ， Faurie (KYO) ， 17 1974. Systematics. 891 pp. Harper & May 1907 ，Miyake (TNS) ， 17 May 1907 ，Miyake (SAPT) ， Row ，New York. 28 28 Ju l. 1936 ，Hara (S). S. KAMCHATKA: between Takahashi H. 1986. Pollen morphology of Pyrola and its Petropavlovsk Petropavlovsk and Avatcha volcano ，6 Aug. 1920 (Hult 釦 taxonomic significance. Bo t. Mag. Tokyo 99: 771 771 (S) ， 22 Aug. 1920 ， Hulten 1025 (S); Avatcha volcano ， 137-154. 5 Aug. 1920 ，Hult 白1 747a (S); pass between T(?)avan and Yamazaki T. 198 1. Pyrolaceae. In Satake ， Y. et al. (eds.) ， T(?)hadulka T(?)hadulka rivers ，1 Aug. 1921 ， Hulten 2575a (S); Savoiko ， Wild flowers of Japan ， Herbaceous plants II 1. Heibon- 28 28 Aug. 1928 ，Eyerdam (S). sha Pub l. Co. Ltd. ，Tokyo (in Japanese).

References References 要旨 Andres Andres H. 1912. Zwei neue Pirolaceae aus der Subsection Erxlebenia Erxlebenia (Opiz). H. Andres nebst einigen Bemerkungen zur Systematik der heimischen Arten. 東アジアに分布する Pyrola fl αune αnαカラフ Verh. Verh. Bo t. Ve r. Brand. 54: 18-227. トイチヤクソウの分類学的取扱には，これまで問 Christoph Christoph H. 192 1. Untersuchungen uber die mykotrophen Verhaltnisse Verhaltnisse der “" und die Keimung von 題があった.特に，周北極要素の P. minor エゾ Pirolaceen. Pirolaceen. Beih. Bo t. Centralb 1. 38(17): 115-157. イチヤクソウ，ユーラシアに広がる異質 4 倍体の Haber E. 1983. Morphological variability and flavonol P. medi α，北アメリカに生育する自然雑種 P. chemistry chemistry of the pyrola αsα rifolia complex (Ericaceae) in in North America. Syst. Bot. 8: 277-298. αsarifoli α〉く minor との区別が難しかった.外部 一一一一 1984. A comparative study of pyrola minor x P. 形態，花粉と種子の微細形態・稔性，染色体数， asarifolia asarifolia (Ericaceae) and its parental species in North America. America. Can. J. Bo t. 62: 1054-106 1. 葉フラボノール成分，地理分布の比較から ， P. 一一一一一 1985. The hybrid identity and taxonomic status of fα une αnαは2 倍体の独立種であることを明らか Pyrola Pyrola sikkimensis (Ericaceae) from eastern Himalaya. にした.これら 4 種類を区別するための検索表を Can. Can. J. Bo t. 63: 133-137. 一一一一 1987. Hybridization of Pyrola chlorant 加 つくり，さらに P. fl αune αnαの最新の記載と充 (Ericaceae) (Ericaceae) in North America. Can. J. Bo t. 66: 分な標本引用をおこなった.これらの形質から総 1993-2000. 1993-2000. Haber E. and Takahashi H. 1988. A comparative study 合判断して， P. fl αune αnα は片親に P. minor を of the North American pyrol αα sarifolia and its Asian もっ雑種起源の種と推定された.もう一方の片親 vicariad ， P. inc αrnat α(Ericaceae). Bo t. Mag. Tokyo 101: 101: 483-495. は，北アメリカに分布する P. αsarifoli αのアジ Hagerup O. 194 1. Zytookologische Bicornes ・.Studien. ア産姉妹種 P. αsarifolia subsp. inc αrn αtα ベニ Planta Planta 32: 6-14. バナイチヤクソウである疑いがある.