Table Of Content植物研究雑誌
J. Jpn. Bot.
甜:8-22 (1993)
Taxonomic Status of the East Asian Pyrola faurieana (Ericaceae)
Erich HABERaa nd Hideki TA KAHASHlb
aResearch Division，C anadian Museurn of Nature，
P. 0. Box 3443，St ation D，O ttawa，K IP 6P4 CANADA;
bBotanic Garden，Fa culty of Agriculture，H okkaido University，S apporo，0 60 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
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
b
(Received on August 28，1 992)
The east Asian species Pyrola faurieana was investigated to clarify its taxonomic distinction from
the circurnboreal P. minor，t he Eurasian allotetraploid P. media and the North American hybrid P.
asarifolia xP . minor，w ith which it has been confused，a nd to evaluate its possible hybrid origin. In
its overal1 morphology，P. fiαurieana is most simi1ar to the hybrid P.αsαrifo/iαxP. minor，b ut is
distinguished by its rnore floriferous scapes，s horter bracts and higher sepal length/width ratio，its
variable production of viable pollen，s eerningly norrnal seeds and distinctive leaf flavonol rnarkers.
pyrola faurieana has the sarne leaf flavonol pattern as P. media and is sirnilar to it in pollen and seed
rnorphology，bu t differs in being diploid，sr naller in stature and in having distinctively srnaller anthers，
generally rnore flowers，sh orter petals and as rnal1er bract length/width ratio. P. faurieana is recognized
as ad istinct species of hybrid origin with P. minor being one of the parents. The second parent is
suspected of being the Asian vicariad of P. asarifolia，n arnely，s ubsp. incarnata (DC.) Haber et
Takahashi. The first report of chrornosome nurnbers for P. faurieana is docurnented as 2n =4 6.
Introduction the two taxa (Kitamura et al. 1978，Y amazaki
Andres (1912) first described Pyrola faurieana 1981)，b ut others only recognize P. faurieana in
on the basis of a collection by Faurie made at Japan (Ohwi 1965，1 983).
Korsakof，Sa khalin 1s. Itss pecific status has been め'rolafaurieana is also morphologically similar
questioned，ho wever，an d the taxon confused with to the North American hybrid P.αsαrぴoUαXP.
P. minor L. Both P. faurieana and P. minor were minor (Haber 1984) and to the Eurasian P. media
recognized by Hara (19~8， 1948) and distinguished Sw.，a p resumed amphidiploid (2n =92 ) between
ont he basis of style，a nther，a nd sepallengths，a nd the circumboreal P. minor and the Eurasian P.
sepal and stigma shapes. Some authors of recent rotundifolia L. (Hagerup 1941).
Ja panese floras have maintained the distinction of A comparative study of P. faurieana，P. media，
-8ー
February 1993 Journal of Japanese Botany Vol. 68 No. 1 9
P. asarぴoliaxP . minor，a nd P. minor was under- A cluster analysis (average linkage) of morpho-
，
taken to clarify the taxonomic status of P. logical characters for 77 specimens of P. faurieana
fαurieana，e specially with regard to its possible was performed (with missing data for some speci-
hybrid origin. As part of the evaluation of taxo- mens) to determine if any geographic differentia-
nomic status for this taxon，m easurements of a tion was evident between the more northern
number of morphological features were made and Kamchatka specimens (those closest to the range
analyzed statistically. Pollen stainability and ab of the morphologically similar P.α'sarifolia xP .
田
normality were examined and seed set determined minor) and those further south，e specially on
in P. faurieanαwere compared with the other taxa. Hokkaido and adjacent islands. As weU，da ta sets
Chromosome counts were made to determine for P. faurieαnαandP.α'sarぴoliaxP . minor wぽe
ploidy level for P. faurieαna. Distributional ranges combined (108 specimens，to tal) and analyzed to
and habitat preferences of the four taxa were also determine whether any distinctive clustering and
reviewed. The possible relationship of P. asαrifoliα separation of specimens could be found. Statistical
，
Michaux subsp. incαrnatα(DC.) Haber et analyses were performed on ap ersonal computer
Takahashi to P. faurieana was also examined. using commercial software from Systat Inc.
(Evanston，IL ).
Materials and methods In Pyrola，as in the majority of the Ericaceae，
Specimens from the following herbaria were the four pollen grains derived from the snepollen
創
used for compiling measurement data，di stribu- mother cell are retained as a pollen tetrad at
tional information and as sources of leaf material maturity. Pollen tetrad diameters and concurrent
for flavonol pattern analysis: BG，C ，C AN，D AO， colpi lengths reported in a previous work
GB，G H，K ANA，K YO，M AK，S ，S APT，T I， (Takahashi 1986) for three of the four taxa under
TNS，T US，U PS，H erbarium Ibaraki Univ.， study here，ex cluding P.αsaベfoliaxP. minor，w ere
Kurashiki Museum of Natural History，a nd the evaluated in the context of the present study.
Faculty of Agriculture of Kyushu University. Pollen stainability was determined by the toluidine
Abbreviations follow the latest Index Her- blue-lactophenol method (Radford et al. 1974).
bariorum; for SAPT，se e“News and notes"担 Pollen was also compared with the North
Taxon 32: 703 (1983). American and Asian subspecies of P.α'sarifolia.
Statistical data for the means of 11 morpho- Overall seed lengths and numbers of testa cells
logical characters examined in this study were were determined under a light microscope for
compiled and summarized graphically. A signifi- samples of 20 seeds of P. faurieana (6 samples)，
cant difference was inferred in the character means P. media (3) and P. minor (10). Seeds were
of two taxa being compared when the confidence scattered on glass slides and sealed under ac over
intervals represented by twice the standard devia- glass with paraffin.
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-
Where an overlap was evident，th e significance of alcohol (3: 1) and stored in 700/0 ethanol. Rhizome
the sample variances was judged by comparing the tips were treated apart in aceto-carmine，sq uashed
calculated Fs tatistics with published values at the under ac over slip，a nd heated briefly under a
.05 level of significance. flame，or were softened by heating for one hour
10 植物研究雑誌第68巻第1号 平成5年2月
at 370Ci n 1N H Cl，th en squashed. Young anthers and widest leaves and longest sepals，pe tals，an thers
were dissected from flower buds of about 2m m and style.乃/rolaminor，in contrast，is significantly
diameter and teased apa此inaceto-carmine to free the smallest in blade width，br act length and width，
pollen mother cells. Slides were briefly heated and sepallength，pe tallength，an ther length and style
cells squashed under the cover slip. Chromosomes length.
were counted under oil immersion with the aid of Of the two taxa with which P. faurieanαcan
ac amera lucida drawing tube. Collections yielding be most readily confused，P . media and P.
counts are cited in the Appendix. asarifolia xP . minor，it is most similar to the latter.
The two-dimensional paper chromatographic The statistical summaries of the character means
techniques used in determining the leaf flavonol analyzed (Fig. 1) show no significant differences
patterns of herbarium specimens were those in the sample means of P. fiαurieαna and P.
described by Haber (1983). As in previous studies， asarifolia xP . minor in six of the 11 characters
only the ultra-violet-quenching compounds (scape h勾ht，blade length and width，br act width，
(appearing black under U.Vよwhichserve as con- sepallength and style length). Ont he other hand，
sistent and useful markers，w ere assessed. The all eleven character means are significantly diι
s担lIlarityin flavonol spot patterns was judged on ferent between P. faurieanαand P. media. The
the basis of the average R values of the com- means for only two characters (petal length and
f
pounds as well as their relative positions and con anther length) are significantly distinct for all four
国
centrations. Spot numbering in the patterns is taxa. 1n absolute measurement values，h owever，
complementary to that in earlier publications (e.g.， there is ab road range in overlap of most characters
Haber and Takahashi 1988). Chromatographed so that one must rely on a combination of
specimens of P. faurieana，P. minor，P. media and character measurements as well as other evidence
P.αsa.rぴo，i[αXP. 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 consideration are based on published data P. mediαand P. minor， as judged by the
，
(Haber 1985，H ulten 1930) and on herbarium length/width ratio (l/w) of 3.1 for these taxa as
collections. About 800 and 600 specimens，r es compared with those of 2.3 and 2.6 respectively
四
pectively，w ere measured for P. minor and P. for P. faurieαna and P.αsαrifo，i[αxP. minor.
media for 11 selected characters (see Table 1). For The cluster analysis of P. faurieana from Ja pan，
the geographically restricted P. faurieana，7 7 the adjacent islands and Kamchatka resulted in a
specimens were available and only 31 for the uniform spread of the Kamchatka specimens
sporadically occurring North American hybrid P. (previous~ treated as P. mediα，by Hulten 1930)
αsa.rぴoliaXP . minor. throughout the various subclusters. Nog rouping
by geographical origin of specimens (i.e.，H onshu，
Results Hokkaido，S akhalin Is.，K urile Is1ands and
Morphological anαlyses The statistical data Kamchatka) was evident.
summary，as compi1ed in Table 1a nd graphically The cluster analysis of the combined data sets
portrayed in Figure 1，sh ows clearly that P. media for P. faurieana and the North American hybrid
is overall the tallest and most robust of the four P.αsarifolia xP . minor resulted in ar elatively
taxa compared. It has the tallest scapes，lo ngest 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.mediα P.faurieanα P.minor P.asarifoUαxP.minor
N x sm N X Sm N X Sm N X sm Ehw-旬。ロ
BSNB1cluaaarddpneeeb elwhreei nidoggtfthh ht f (l((coccrrrwnnne))) r s 555588880330 22493....15055549 4003....26063886 0000....01013833 IIII司『『『IεJζJd斗『 112720....49463976 0033....45613699 0000....40035663 0000000000002。，、dOOozJ 112361....03123888 4003....15767760 0000....01103532 32d3d司今今TI'ITi'A 12267....40863423 2002....44284339 0000....40058802 Hum520。-ーし『お
Bract length* (rnrn) 247 5.50 0.96 0.06 roAU 4.81 0.80 0.10 3今93今 4.41 0.91 0.05 3今'i 5.74 0.68 0.12 回。
BSreapcatl lweindgtthh * ((r rnnrrnn)) 414999 21..7349 00..3332 00..0022 a斗『Id'i斗 22..1012 00..4246 00..0037 I司h今，443du寸今 11..4392 00..4294 00..0013 41d今21A斗・ 22..2010 00..1564 00..1013 EH岡山『〈。
APSSteenytptlaahell e lrlwee inlndgegttnthhgh t ((h(rrr nnn(rrrrnnnn)))r n) 442341759242 4611....99631860 0000....27261902 0000....00001144 IξJ/oro『Ioofo'i『 5311....66161374 0000....16165382 0000....00008822 3ζudI斗『今697 10Yd3品寸今『 4111....04535926 0000....13251324 0000....00001320 4 3334々今勾今tiζus--- 3511....95595220 0000....41621970 0000....00011242 Z--。∞。・】
Note: Leaf rneasurernents were rnade on largest leaf，a nd floral rneasurernents on lowest，rn ost rnature flower; N，sa rnple size;，支rnean; s，s tandard deviation;
sm' standard deviation of the rnean; *bract rneasurernents are those of the inflorescence bracts.
......
』司
12 植物研究雑誌第68巻第1号 平成5年2月
Scape height (cm) Sepal length (mm)
M8 -→ー ME ー+ー
FR1 一一一←一一 FR →ー
MI~ -← MI →-
AxMI ーー一一+ーー一四 →-
16 17 18 19 20 21 22 1.3 1.5 1.7 1.9 2.1 2.3
Bla de length (cm) Sepal width (mm)
ME ー+ー →ー
FR 一ー+ー- ー-+一一
MI ー←- MI ー+ー
AxMI -一←ーー AxMI 一一←司自
2.5 3.0 3.5 4.0 1.5 1.6 1.7 1.8 1.9
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 3.0 3.5 4.0 4.0 5.0 6.0
Number of flowers Anther length (mm)
ー-
MEi ME I- +
FR~ 一一←ー一 FR I- -+
MH -← MH
..j.
AxMH AxMI1 -一←ー
7 8 9 10 11 12 1.0 1.2 1.4 1.6 1.8 2.0
Inflorescence bract length (mm) Style length (mm)
一ー→一ー 山 +
山
ー+・
内
MI 川 +
AxM1i AxM'l →ー
4.4 4.6 4.8 5.0 5.2 5.4 5.6 5.8 .10 2.0 3.0 4.0 5.0
Inflorescence bract width (mm)
川 ←ー Mc」l」pyrdam創いれa
A内川洲 一→-ー 一一ー十ーー・ AxMM円、I2IZir--t←pp Pyy yrr rod olMa a hm asa出nr.noier foa li円HaRd xP. minor X、』 2ーsmJ
1.3 1.5 1.7 1.9 2.1 2.3 2.5
Fig. 1. Graphical comparisons of 11 character means for four Pyrola taxa. Horizontallines
represent 2x t he standard deviation of the mean on either side of the means.
specimens as several interconnected cluster groups. Pyrolafiαurieanαspecimens formed severallarger
Five out of the 31 hybrid specimens available interconnected cluster subsets. Out of at otal of 108
linked with adjacent P. faurieana clusters. Of the specimens in the cluster analysis，a t otal of 8，or
77 P. faurieana specimens in the analysis，th ree about 70/0o f the combined sample were“incor-
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 xP . minor is evident on general
western limit of the range of the North American inspection of specimens and in the lack of signifi-
hybrid in the Aleutian Islands (see Haber 1984). cant differences in the means of six characters，th e
February 1993 Journal of Japanese Botany Vol. 68 No. 1 13
cluster analysis indicates that the specimens of the this taxon from Japan，Sa khalin Is. and the Kurile
two taxa do exhibit sufficient differences overall Islands have ad iameter range of 36-46 (57)μm
that they tend to associate as separate cluster sets. and am ean of 42μm (based on values in Table
This no doubt reflects the significant differences 1o f Takahashi (1986) and additional measure-
in the means for number of flowers，br act length， ments by Haber). The occasional collection con-
sepal width，pe tallength and anther length evident ta\ns up to 100/0 giant dyads. In the taxon as a
in Figure 1. whole，mo st tetrads tend to be somewhat shrivelled
Pollen tetrad diameters and stainability The and only take on am edium blue stain in lacto-
presence of some genetic irregularity is evident in phenol and aniline blue. About 50% or more of
the holotype of P. faurieana and in other repre- the tetrads in some collections are highly shrivelled
sentative specimens. Tetrads in the holotype e and presumed abortive.
紅
imperfectly formed and partly shrivelled with many Two collections at SAPT from Mt. Makkari-
exhibiting al oose tetrahedral arrangement of the nupuri (Nishida 21 Aug 1906 and Konishi 28 Aug
four grains. The large tetrads are about 42μmin 1922) identified as P. fiαurieαna had completely
diameter and tend to stain light blue，un like the abortive pollen. However，t wo other collections
deep blue stain characteristic of normal plump from the same mountain (13 Aug 1986，Ta kahashi
tetrads in other species. Large empty and small 6662 and 6664，S APT) had only the more typically
shrivelled tetrads are also present. Specimens of incompletely expanded，s omewhat shrivelled
45
l
門=←
(
目立 nu
)
十斗士
』
4)
ω
呂
町喧
"0
て，3
~ 35
。圃.
E4) → 干 芋 #
叫
+
30 • P. faurieana
ート .A P. media
• P. minor
~
15 20 25 30
Concurrent colpi length (μm)
Fig. 2. Bivariate plot of sample means and their confidence limits (2 xs tandard deviation
oft h e mean on either side of the means) for concuηent colpi length and pollen tetrad diameter
in three Pyrola taxa.
14 植物研究雑誌第68巻第1号 平成5年2月
appearance commonly seen in pollen of P. disorder present in the species. This genetic
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. mediαthat not only have malformed and poorly
Peninsula，H okkaido. A collection from Mt. stained tetrads but also have other floral abnor-
Okkapake made by Takahashi and Kushibiki 5164 malities. Ac ollection of P. media from the USSR，
(SAPT)，h ad highly abortive pollen，ye t as econd chromatographed for the present study (Province
collection by Takahashi & Kushibiki 5157 (CAN)， Orel，1 J ul. 1906，C AN)， is abnormal in having
from Futatsu-like several hundred meters away， af lowering branch in the lower part of the raceme
had mostly well-stained pollen tetrads，b ut the in addition to having abnormal tetrads that are
grains tended to be somewhat collapsed and the poorly stained.
tetrads not always clearly tetrahedral in form. pyrola minor，i n contrast to the three other
Pollen tetrads in P. media range in size from taxa，h as deeply stained，fu lly expanded tetrads
37-43μm with am ean of about 41μm. Data within 24 hours. These range in size from 27-39
published by Takahashi (1986，T able 1) for this μm with a mean of 33ρm. Tetrads of P.
species and additional collections examined for the asarifolia xP . minor are highly malformed with
present study by Haber (citations in Appendix) collapsed walls and are stained weakly. The larger
indicate that pollen stainability，a nd presumably somewhat collapsed tetrads tend to be 36-38μm
viabi1ity，va ries considerably.恥10stcollections of in diameter，al so stain weakly and are considered
P. media have relatively large tetrads，b ut these non-viable. Even after five years in stain，th e grain
generally tend to be shrivelled to various degrees. contents remain weakly stained，al though some of
Estimating the proportion of grains that might be the larger tetrads have grains with expanded walls，
considered to be relatively normal in their ah igh proportion of malformed tetrads remain.
appearance and stainabi1ity is difficult here，as in Pollen tetrad diameters are not significantly
P. faurieana，b ecause of gradation between tetrads different between P. faurieana 支'(=42μm)andP.
that are somewhat collapsed，bu t more or less well media 支(=41μm)，and both have larger diameters
stained，a nd those that are progressively more than those of P. minor (支=33 pm，Fi g. 2). Pyrola
collapsed，e mptyor weakly stained. Knaben (1944) fαurieαna tetrads are significantly larger than the
estimated that pollen viability was reduced to about highly shrivelled and abortive tetrads of the North
23 0/0. Ac ollection such as that made by H. Andres American hybrid P. asarifolia xP . minor. The
in Germany (Koln，A ug. 1929，C AN) has darkly highly abortive pollen of the Okkapake collection
stained and mainly fully expanded tetrads，ye t even of P. faurieanι(Takahashi and Kushibikii 5164)，
here particularly large，4- celled tetrads，a bout 45 has am ean diameter of about 39μm，fo r the more
μmi n diameter，ar e present and in non-tetrahedral fully formed tetrads，w ith a broad range in
formation. In aB ritish collection (Ambleside，24 diameters being present. Although there is ab road
Jun. 1869，C AN) in which the tetrads stain dark overlap in the absolute lengths of concurrent colpi
blue，a nd are of normal size，m ost tetrads are in P. faurieana 支'(=25pm)，P. media 支'(=20μm)
highly shrivelled. Age of pollen samples may affect and P. minor (支=17μm)，a s mall but significant
the ability to rehydrate，bu t the collapsed nature statistical difference exists in the means (Fig. 2).
of tetrads，ev en after rehydration in stain for over Exine sculpturing in P. faurieana，P . media and
ay ear，is more likely ar eflection of the genetic P. minor is verrucate (Takahashi 1986).
February 1993 Journal of Japanese Botany Vol. 68 No. 1 15
Table 2. Summary of statistical data for two seed characters of three species of Pyrola.
Sample Seed length (mm) Testa cell number
x sm X sm
P.fiαurieαnα 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 0.42
5 0.64 0.05 0.02 7.30 1.08 0.48
6 0.56 0.05 0.02 7.00 0.92 0.41
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. minor 0.44 0.05 0.02 7.30 1.30 0.58
2 0.55 0.05 0.02 7.80 0.89 0.40
3 0.65 0.05 0.02 9.25 1.33 0.59
4 0.62 0.04 0.02 8.80 0.95 0.42
5 0.48 0.03 0.01 7.60 1.10 0.49
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 0.49
9 0.59 0.05 0.02 9.35 1.09 0.49
10 0.60 0.05 0.02 8.80 1.11 0.50
Seed set and size pyrola seeds are minute， faurieana from 11 metaphase plates in rhizome tips
consisting of as mall nodule of cells comprising of collections by Takahashi (5249) and Takahashi
由eundifferentiated embryo，c entered within an and Kushibiki (5157). A haploid number of about
elongate，th in-walled testa. A bivariate graph of 23 chromosomes was counted in 12 plates of a
testa cell number and seed length of P. media，P. collection of the same taxon made by Takahashi
faurieαna and P. mino九basedon 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 in seed size among the three taxa，al though several years old，c hromosome separation and
P. media and P. faurieana tend to have somewhat staining was not optimal. Chromosome sizes
larger seeds than those of P. minor. They also tend ranged between 2.5-5.0μm long.
to have fewer testa cells per length than P. minor. Although chromosome counts were not in-
Pyrola minor，in contrast，ha s ab roader range in dividually conclusive，t hey were consistent in
seed length and testa cell numbers，wi th increased magnitude and sufficient in number of counts to
seed length being correlated with increased testa indicate that the diploid number is probably
cell number. Capsules of P. asarifolia xP . minor 2n =4 6，th e same as has generally been reported
bear only aborted ovules，a r eflection of the highly in other diploid species of the genus. This is
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 at etraploid
approximately 40 chromosomes were made for P. number.
16 植物研究雑誌第68巻第1号 平成5年2月
0.8
0.7
言
0.6
、、缶 i 4
J 7
7
~
-回圃.
営 0.5
0
F司
句。
ω
∞
0.4
• P. faurieana
....P. media
• P. minor
L→
7 8 9 10
Testa cell number
Fig. 3. Bivariate plot of sample means and their confidence limits
(2 xs tandard deviation of the mean on either side of the means)
for seed testa cell number and seed length in three Pyrola taxa.
Chromαtographic patterns of leaf flavonols 持1and in having非8.
The 2dimensionalchromatographic patterns of Collections of P. faurieana with pollen that was
・
leaf f1avonols for P. faurieana and the three other highly abortive (Konishi，28 Aug. 1922; Nishida，
taxa with which it was compared are given in Fig. 21 Aug 1906; Takahashi and Kushibiki 5164，a nd
4. Based on the previous work with other species Yoshimura and Yokoyama 5A ug 1938) did not
of Pyrola (Haber 1983)，t hese UV-quenching differ from the f1avonol pattern present in collec-
flavonols are all glycosides of quercetin (#1-5，8) tions with more normal levels of pollen viability.
and kaempferol (尚). Similar1y，th e holotype，wi th its highly collapsed
The f1avonol pattern of Pyrola faurieanαis pollen，a lso did not show any deviation in its
identical to that of P. media. Pyrola faurieana flavonol pattern from that of other specimens of
differs from the North American hybrid P. this taxon with higher levels of nearly normal
asarぴ'oUαxP. minor by having ap air of over- pollen.
lapping glycosides (何 and#8) and in lacking Distribution The worldwide distributions for
glycoside再1of the hybrid. The absence of f1avonol P. media， P. minor， P. faurieαna and P.
#5 i n P. asar.ぴoliaxP . minor is unusual because asarifoliaxp. minor are given in Fig. 5. Pyrola
it is present in both of its parents. A second minor is aw idespread circumboreal species，fo und
flavonol (打)， found in P. asarifolia (see Haber primarily in mossy coniferous forests but also in
1983) is also absent. This hybrid seemingly lacks mixed woodland sites and deciduous forests in
the full complement of parental f1avonols. Pyrola northern Sweden and Jap an. It is found over a
faurieana differs from P. minor in lacking f1avonol wide elevational range from near sea level to alpine
February 1993 Journal of Japanese Botany Vol. 68 No. 1 17
A B
2・D 2・D
;.1
0.5 s 0.5
4 ~21
1 ・
1・D 0.5 1・D 0.5
Pyrola as官，官bllaxP yrola mlnor Pyrola mlnor
C 、D
2・D 2・D
・，
‘
s.2
8 8
2
0.5 6 0.5
•
6'.
s
1D 0.5 1・D 0.5
岨
Pyrola medla Pyrola faurfeana
Fig. 4. Two-dimensional chromatographic patterns of UV-quenching leaf flavonols in four
pyrola taxa.
meadows. In the more southerly extensions of its range. It occurs most commonly in Jap an
range in the Rocky Mountains of Colorado，it (northern Honshu and Hokkaido). However，it is
reaches elevations of 3600 m and in SE Tibet α4旧 sporadic and considered rare. Northward it is
m. pyrola minor is rare in northern Japan. found on Sakhalin Is.，w here the type was col-
pyrola media is Eurasian with a relatively lected，o n the Kuriles，s outhern Kamchatka and
scattered distribution within its overall broad Bering Is. It is a plant of higher elevations，
range. Like P. minor，it s preferred habitats are occurring in the Pinus pumila (pall.) Regel scrub
mossy coniferous woodlands. It occurs near sea zone and alpine meadows in northern Jap an，
level but is more commonly found in mountainous associated with such species as Empetrum nigrum
terrain reaching the upper subalpine meadows. 1n L. and Tilingia ajanensis Regel. Occasionally it is
the Caucasus Mountains， it reaches alpine found with P. minor and P. al1フ的αH. Andres.
meadows at about 2800 m. pyrola mediαextends Specific site localities for the North American
，
eastwards in Siberia to about Lake Baikal. hybridP.αsarぴ'o[i，αxP.minor，as plotted in Figure
pyrola faurieana has ar elatively restricted total 5，h ave been updated from those first published
