料:a calli nthemum c. sch linensecalli αnthemum (ranunculaceae) from japan and its adjacent...

15
September1989 Journ.Jap. Bot.Vo l. 64 9 KenSATO* & KojiITO 料: A noteonthetaxonomyof Calli α nthemum (Ranunculaceae)fromJapanandits adjacentarea withreferencetoa newsubspeciesof C.s α ch α linense fromHokkaido Japan 佐藤 謙*・伊藤浩司料: 日本近隣キタダケソウ属の分類ノートー 特に北海道産カラフトミヤマイチゲの新亜種と関連して 257 The genus Callianthemum (Ranunculaceae) includesabout14speciesandis disjunctivelydistributedtothesubalpineoralpineregionsofEurasia(Komarov 1937 Turtin eta l. 1964 Wangeta l. 1980). InJapananditsadjacentareas fourspecieshavebeendescribed: C. insigne fromMt.Kanboho in North Korea (Nakai1919 1920 1928) C. miyabeanum fromMt.Apoianditsneighborhoods in Hokkaido(Tatewaki1928) C. hondoense fromMt.KitadakeinHonshu (Nakai & Hara1934) andC. sachalinense fromSouthernSakhalin (Miyabe & Tatewaki 1935). However the taxonomicaltreatments of C. miyabeanum andC. hondoense havebeendiverseamongtaxonomistsinJapan. A question arose as to whether eitherofthemisregardedasaspeciescontrastingwithC. insigne ornot. NakaiandHaratreatedC. miyabea ηum asconspecificwith C. insigne but C. hondoense asadistinctspecies (Nakai1930 Nakai & Hara1934 Hara1935). Onthecontrary OhwiconsideredC. miyabeanum to be adistinctspecies while herstreportedC. hondoense asconspecificwith C. insigne andlaterreduced ittoavarietyof C. insigne i . e.var. hondoense (Ohwi1932 1953). Kitamura & Murata (1962)treatedC. miyabeanum asavariety of C. insigne i. e. var. miyabeanum (Tatew.) KitamuraetMurata butC. hondoense asconspeci :fi cwith C. insigne. Kitamura & Murata'sopinionissimilartoOhwi's one because both ofthemconsideredC. hondoense to bemorerelatedto C. insigne thanC. miya- beanum. ThoughShimizu (1983) recognizedthetwospeciesunderquestionas varietiesof C. insigne thereremainsapairofopinionsastothetaxonomyof * Laboratory of Biology Facultyof GeneralEducation Hokkai-GakuenUniversity Toyohira-ku Sapporo062. 北海学園大学教養部生物学教室. DepartmentofBiosystemManagement Devision of EvironmentalConservation GraduateSchool ofEnvironmentalSciense HokkaidoUniversity Kita-ku Sapporo060. 北海道大学大学院環境科学 研究科環境保全学専攻生態系管理学講座. -1-

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Page 1: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

September 1989 Journ. Jap. Bot. Vol. 64 No~ 9

Ken SATO* & Koji ITO料: A note on the taxonomy of

Calliαnthemum (Ranunculaceae) from Japan and its

adjacent area, with reference to a new subspecies of

C. sαchαlinense from Hokkaido, Japan

佐藤 謙*・伊藤浩司料: 日本近隣キタダケソウ属の分類ノートー

特に北海道産カラフトミヤマイチゲの新亜種と関連して

257

The genus Callianthemum (Ranunculaceae) includes about 14 species and is

disjunctively distributed to the subalpine or alpine regions of Eurasia (Komarov

1937, Turtin et al. 1964, Wang et al. 1980). In Japan and its adjacent areas,

four species have been described: C. insigne from Mt. Kanboho in North Korea

(Nakai 1919, 1920, 1928), C. miyabeanum from Mt. Apoi and its neighborhoods

in Hokkaido (Tatewaki 1928), C. hondoense from Mt. Kitadake in Honshu (Nakai

& Hara 1934) and C. sachalinense from Southern Sakhalin (Miyabe & Tatewaki

1935). However, the taxonomical treatments of C. miyabeanum and C. hondoense

have been diverse among taxonomists in Japan. A question arose as to whether

either of them is regarded as a species contrasting with C. insigne or not.

Nakai and Hara treated C. miyabeaηum as conspecific with C. insigne, but

C. hondoense as a distinct species (Nakai 1930, Nakai & Hara 1934, Hara 1935).

On the contrary, Ohwi considered C. miyabeanum to be a distinct species, while

he五rstreported C. hondoense as conspecific with C. insigne and later reduced

it to a variety of C. insigne, i. e. var. hondoense (Ohwi 1932, 1953). Kitamura

& Murata (1962) treated C. miyabeanum as a variety of C. insigne, i. e. var.

miyabeanum (Tatew.) Kitamura et Murata, but C. hondoense as conspeci:fic with

C. insigne. Kitamura & Murata's opinion is similar to Ohwi's one because both

of them considered C. hondoense to be more related to C. insigne than C. miya-

beanum. Though Shimizu (1983) recognized the two species under question as

varieties of C. insigne, there remains a pair of opinions as to the taxonomy of

* Laboratory of Biology, Faculty of General Education, Hokkai-Gakuen University, Toyohira-ku, Sapporo 062. 北海学園大学教養部生物学教室.

料 Departmentof Biosystem Management, Devision of Evironmental Conservation, Graduate School

of Environmental Sciense, Hokkaido University, Kita-ku, Sapporo 060. 北海道大学大学院環境科学

研究科環境保全学専攻生態系管理学講座.

-1-

Page 2: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

258 植物研究雑誌第 64巻第 9号 平成元年 9月

the above two species according to the di旺erenceson the most important diag-

nostic characteristics, which wiII be discussed below. However, the treatments

above mentioned have not been based on the comparisons with C. sachalinense

at all.

The plants of Callianthemum from Mt. Kirigishi in Hokkaido were reported

to be C. miyabeanum, and pointed out to resemble C. sachalinense or C.

hondoense by one of the authors (Ito 1982). Since the taxonomic position of

the plants has not been confirmed, the authors examined many specimens col-

lected from Mt. Kirigishi and the chromosome number of the plants. The

authors have concluded that the plants from Mt. Kirigishi should be a new

subspecies of C. sachαlinense, i. e. subsp. kirigishiense, and that C. miyabeanum

should be treated as a distinct species related to C. sachalinense, contrasting

with both C. hondoense and C. insigne.

Chromosome number As shown in Fig. 1, the chromosome number of this

new subspecies (C. sachαlinense subsp. kirigishiense) was counted to be 2n=

Fig. 1. Drawing of somatic chromosomes

of Callianthemum sachalinense ssp.

kirigishiense (2n=16). The bar indi-

cates 10μ.

16 for the五rsttime, which is the same

as that of C. hondoense (Kurita 1958)

but di百ersfrom those of C. miyabeanum

(Sakai 1935; 2n=32) and C. insigne

(Kurita 1958; 2n=32). However, the

chromosome number of subsp. sachalin-

ense has not yet been found. Fur幽

thermore, the 16 chromosomes have

been counted in C. coriαndrifolium

(Langlet 1932) and C. angustifolium

(Rostovsteva 1981), while the 32 chromo~

somes in C. anemonoides (Langlet 1932)

and C. kernerαnum (Damboldt 1966) at

present.

Petal characteristics The ratios of Iength to width of petals in the popula-

tion level seem to be the most important diagnostic characteristic among the

:above four species in ]apan and its adjacent areas, because its values correspond

to their geographical areas (Fig. 2 & Tab. 1-3). The ratios of C. sachalinense

which is composed of two subspecies and C. miyabeanum are mostly more than

2. The value of subsp. kirigi・shiensewhich distribute between subsp. sachalinense

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Page 3: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

September 1989 Journ. Jap. Bot. Vol. 64 No. 9

S( N=54) I中

K< N=104) I

M (N=56) I =ゴ亡仁コ IH ( N:17)

匹同 I(N=7)

2 3 4 5 6 Fig. 2. Ratios of petal length to petal width of Callianthemum (ranges, standard

deviations and means): [S]: C. sachalinense ssp. sachalinense [KJ: C. sachalin-ense ssp. kirigishiense, [MJ: C. miyabeanum, [HJ: C. hondoense and [I]: C.

znszgne.

259

and C. miyabeanum is intermediate between them. On the other hand, the ratios

of C. hondoense and C. insigne are mostly less than 2. At present most taxo・

nomists do not think that C. sachalinense is related to C. miyabeanum. Based

on the petal characteristic and the finding of a new intermediate subspecies, it

is shown that they are closely related, contrasting with the relation of C.

hondoense to C. insigne.

Proportions of the emarginate petals at the apex in the population level

(PEA in Tab. 1 & 3) seem to be diagnostic as recognized by Nakai & Hara

(1934). Proportions in C. hondoense and C. sαchalz'nense are higher than those

of C. miyabeanum and C. insigne. This characteristic seems to correspond to

the differences in the chromosome number, i. e. 2n=16 and 32. Among about

14 species in the world (Komarov 1937, Tutin et al. 1964, Wang et al. 1980),

the higher proportions of the emarginate petals seem to correspond to the 16

chromosomes such as C. coriandrifolium and C. angustifolium, while the lower

ones to the 32 chromosomes such as C. anemonoides and C. kerneranum.

Therefore, this characteristic is considered to be not originated by species dif-

ferentiation but by parallel evolution in association with diploidy. By this

characteristic, C. hondoense is clearly distinguished from the related species,

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Page 4: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

260 植物研究雑誌第 64巻第 9号 平成元年 9月

but C. miyabeanum does not clearly differ from C. insigne, which has been

pointed out by Nakai & Hara (1934) (Tab. 1 & 3). However, on the basis

of the above consideration, the latter two species should not be conspecific with

each other. As to this characterisitc, the new subspecies also shows inter-

mediate values between subsp. sachalinense and C. miyabeanum.

C. hondoense and C. insigne are further characterized by the shorter petals

and the wider petals, respectively (Fig. 3), though the data are still very few.

Other characteristics Whether radical leaves develop before flowering or

after flowering is usually considered to be the most important diagnostic in the

taxonomy of Callianthemum. Petiole length and blade length of radical leaves

at flowering are biggest in C. sachalinense, which shows the development of

radical leaves at丑owering (Fig. 4 & Tab. 1-2). Those of C. hondoense are

developed at flowering, although the length of petiole and blade is relatively

small (Nakai & Hara 1934). On the other hand, the length of them of C.

miyabeanum and C. insigne is small at自oweringand develops after丑owering.

This . characteristic, as well as proportions of the emarginate petals, seems to

correspond to the di百erencesin the chromosome number. Radical leaves in C.

coriandrzjoZz'um and C. angustzjolium which have the 16 chromosomes develop

at宜owering,while those in C. anemonoides and C. kerneranum having the 32

chromosomes develop after自owering. So that, this characteristic is considered

S{N=S4) f 三三二二云 I I r==! r- • I L-ー一寸-

K(N:104) J -土ニ二 I r r==f

M (N=56) I ーーーーー I I r===i

H (N=17) ~ l I =仁コi

1 ( N=7) 田園-i I1

。Fig. 3. Petal length (open) and petal width Csolid) of Callianthemum.

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Page 5: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

September 1989 Journ. Jap. Bot. Vol. 64 No. 9 261

to be derived from the parallel evolution accompanied with diploidy.

Scape length at flowering and flower number on a scape seem to correspond

to the above characteristic (Fig. 5 & Tab. 1-3). C. sαchαlz'nense is distinguish-

able from the related species by the longer scapes, the more numerous flowers

per scape, the longer petioles and blades of radicalleaves, and the longer blades

of cauline leaves, which develop at丑owering(Fig. 2-6 & Tab. 1-2). Among

this species, subsp. kirigishiense shows the shorter or fewer values in the above

characteristics than subsp, sachalinense.

In the ratios of apical petiolule length.to lateral one in radicalleaves (Ohwi

1953), C. miyabeanum has a distinct value, i. e. from 1 to 2 (Fig. 6 & Ta.b.

1-3). This species is further characterized by the petioles of cauline leaves

dilated at the base (Tatewaki 1928), the longer filaments of anthers (Tatewaki

S( N=34)

K(N=39)

暗→M (N=41)

嘩~ H (N=9)

圃国→ 1 (N=10)

10 cm

Fig. 4. Petiole length (open) and blade length Csolid) of radical leaves of Callianthemum at Bowering.

~5-

Page 6: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

262 植物研究雑誌第 64巻第 9号 平成元年 9月

1928; Tab. 1), the round shape of sepals (Ohwi 1953, Kitamura & Murata 1962;

the wider sepals, Tab. 1), and the above-mentioned shorter scapes and leaves

developing after fiowering. . Because these characteristics are mostly specific to

C. miyabeanum, this species should be treated as a distinct species (Ohwi 1953).

S{N=32)

K(N='l.)

M (N=39)

l中 lH (N=7)

IC N=6)

10 20 30 40 Fig. 5. Scape length of Callianthemum at flowering.

S(N=34)

I K(N=33)

t申 M(附

iH H(問

i I I I (N==7)

o 5 10

Fig. 6. Ratios of apical petiolule length to lateral one in radical

leaves of Callianthemum at flowering.

-6ー

50cm

Page 7: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

ωougBσσ円

HU

∞hv

Tab. 1. Comparison of characters among 3 taxa of Callianthemum: [S]: C. sachalinense ssp. sachalinense

from Southern Sakhalin; [K]: C. sachalinense spp. kirigi・shiensefrom Mt. Kirigishi, Hokkaido and [MJ: C.牛 miyabeannmfrom Mt. Apoi, Hokkaido.

M K S Taxa

n m:ts. d. (range) n m:ts. d. (range) n m:ts. d. (range) Characters

』OZ2H・』mH℃・回O門・〈己・白hFZ0

・c

10. 9:t3. 5( 5.0-19.5)39

5. 9:t2.1( 2.9-10.5)41 1.2土0.3(0.7-2.3)41

1. 6:t0. 4( 1. 0-2.0) 8 1.4土0.5(0.9-3.0)26

A scape:

Length a t flowering (cm)

A radical leaf:

Petiole length at flowering (cm)

Blade length a t flowering (cm) RAL* at日owering

RAL * after flowering A cauline leaf:

Petiole length at自owering(cm) Blade length at flowering (cm)

A petal: : Length (mm)

Width(mm)

Ratio of length to width PEA料(%)

Number of petals per flower

A sepal:

Length(mm) 6.3:t1.2( 4.0-8.0)21 6.6:t1.0( 4.0-9.0)59 6.9土1.0(5.0-8.5)27

Width (mm) 3.8土0.7(3.0-5.0)21 3.8:t0.7( 2.0-5.0)59 4.5土1.0(3.0-7.0)27 Number of flowers per scape 2.1:t0.6( 1 -3 )32 1.6:t0.8( 1 -3 )14 1.4:t0.5( 1 -2 )39

Filament: length (mm) ( 2.5-3.5) 6 ( 1. 5-3.5) 11 ( 3.0-5.0) 6

Chromosome number ? 16 [this paperJ 32 [Sakai 1935J

* Ratio of apical petiolule length to lateral one in a radical leaf. ** Proportion of the emarginate petals in the population level. *** The number is not calculated because in most specimens some petals are fallen.

23.8土6.8(10.6-36.2)14

11. 5:t3. 7( 4.5-18.3)39

6. 0:t2. 7( 0.9-11. 3)39

2.6土1.4(1.1-6.5)32 2. 7:t0. 7( 2.1-3.4) 3

29. 8:t6. 4(17.5-47.0)32

14. 2:t3. O( 8.7-22.0)34

6. 4:t1. 7( 3.8-10.2)34

2.7土 1.1(1.7-6.3)34 3.5:t1.9( 1.7-8.0)17

1. 3:t 1.1 ( 0.2-3.2)52

1.0土0.4(0.3-2.5)48

12.4土1.4(9.0-15.0)56

4.7土1.3(2.5-8.0)56

2.0土0.9(1.5-5.6)56

32.1 56

7.7土2.0(5 -12 )36

O. 7 :t 1. 0 ( O. 0-4. 1) 27 2.2:t0.8( 1.0-4.2)27

11.4:t1.1( 9.0-13.5)104

4.8:t1.1( 2.5-7.5)104

2.5士0.6(1.4-4.4)104

47.1 104

7.1:t1. 9( 5 -10 )21

1.4:t2.6( 0.0-13.2)55 2. 5:t 1.1 ( 0.5-4.8)55

12.1:t 1.1 ( 9.5-15.0)54

5.8:t1.0( 4.0-8.0)54

2.1:t0.3( 1. 5-3.0)54

61.1 54 *** . (5 -9 ) 23

民司

N

。ω

Page 8: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

264 植物研究雑誌第 64巻第 9号 平成元年 9月

C. honodoense di百ersfrom the related species in having the leaves intensely

glaucous on both surfaces as recognized by Nakai & Hara (1934). This species

is further characterized by the predominance of the emarginate and shorter

petals, and the shorter scapes and leaves developed at flowering. So that, this

is considered to be a species contrasting with C. insigne.

Habitat As shown in Tab. 4, C. sachalinense and C. miyabeanum, which

distribute nearby each other, are characterized by the occurence in the subalpine

region based on warmth indices (Kira 1948) of about 40ocM. On the other

hand, both C. hondoense and C. insigne occur in the alpine region, of which

warmth indices are below 20ocM. From the geological view point, C. sachalin-

Tab. 2. The t・teston the mean of the characters of Cαllianthemum

shown in Tab. 1.

Taxa S-K K-M S-M

Characters

A scape:

Length at flowering ** (2. 80) **( 6.78) **(14.97)

A radical leaf:

Petiole length at自owering ** (3. 44) ** ( 8.27) **(13.60)

Blade length at flowering (0.77) 料(11.04) **(17.61)

RAL at flowering (0.32) 料(3.51) *( 4.67)

RAL after flowering (1. 31) ( 3.13) **( 4.46)

A cauIine leaf:

Petiole length at flowering ** (1. 75) ** ( 2.44) **( 0.26)

Blade length at百owering (1. 40) ** ( 7.30) **( 9.42)

A petal:

Length ** (3. 79) **( 4.63) **( 1.25)

Width ** (5. 76) **( 0.49) **( 4.98)

Ratio of length to width **(5.59) **( 2.99) ** ( 6.30)

A sepal:

Length (1. 03) ( 1. 29) * ( 1. 85)

Width (0.00) 料(3.29) **( 2.85)

Number of flowers per scape * (2. 09) ( 0.88) **( 5.27)

*: Pく0.05,**: Pく0.01

-8ー

Page 9: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

September 1989 Journ. Jap. Bot. Vol. 64 No. 9 265

Tab. 3. Comparison of characters between Callz'anthemum hondoense from Mt. Kitadake, Honshu [HJ and C. insigne from Mt. Kanboho, North

Korea [IJ.

Taxa

Characters

A scape:

Length at fl.owering(cm)

A radical leaf:

H

m土s.d. (range) n m::ts. d. (range) n

10.8::t2.4( 7.8-14.7) 7 14.6土7.4(6.0-25.0) 6

Petiole length at fl.owering 7.0::t3.1( 3.2-14.6) 9 7.4土2.8(3.5-12.7)10 (cm)

Blade length at fl.owering 3.9::t 0.8 ( 2.7-5.0) 9 2.9土1.4(1.5-5.0)10 (cm)

RAL * a t fl.owering 2. 2土1.0(1.1-4.0) 9 3.7土1.9(1.8-7.5) 7

RAL* after宜owering 2.8::t1.2( 1.6-4.8) 8 2.9::t0.9( 1.4-4.7)22

A cauline leaf:

Petiole length at fl.owering 0.6土0.7(0.0-2.0)15 0.7::t1.1( 0.0-2.7)13 (cm)

Blade length at fl.owering(cm) 1.8::t0.9( 0.7-3.0)15 2.0::t0.9( 1.0-4.1)13

A petal:

Length(mm)

Width(mm)

Ratio of length to width

PEA料(%)

Number of petals per fl.ower***

A sepal:

Length(mm)

Width(mm)

Number of fl.owers per scape

Filament: length(mm) ***

Chromosome number

10.0::!:::0.8( 8.0-11.0)17 12.1::!:::1.0(11.0-13.5) 7

6.2::t1.3( 4.0-7.6)17 7.6::t1.6( 6.0-9.5) 7

1.7::t0.4( 1.3-2.4)17 1.6土0.3(1.2-2.0) 7

88.2 17 28.6 7

(6 -7)3 (6 -9 )4

7.1士0.8(6.0-8.0) 8 6. 7:t0. 7( 6.0-7.4) 8

4.1土0.8(3.0-5.0) 8 3.3::!:::0.4( 3.0-4.0) 5

1.3:t0.5( 1 -2 )12 1.2土0.4(1 -2 )20

( 1.5-2.5) 2 ( 2.0-2.5) 4

16 [Kurita 1958J 32 [Kurita 1958J

* Ratio of apical petiolule length to lateral one in a radical leaf.

** Proportion of the emarginate petals in the population leve1.

*** The number is not calculated, because specimens are few.

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Page 10: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

N

由。Tab. 4. Comparison of habitats among 5 taxa of Callianthemum from ]apan and its adjacent area: [S]:

C. sαchalinense ssp. sachalinense, [K]: C. sachalinense ssp. ki・rigi・shiense,[M]: C. miyabeanum, [H]:

C. hondoense and [1]: C. insigηe.

Plant community Geology Warmth index* CcM)

Latitude (ON) Locality

苗品社出端凶謀関

Altitude (m)

[S] ↑L、:ノ

snud

eqυ

rQd

O咽

i

PTA y

m

ea

n山

wm伊

eu

'HQU

Lfk

chert, slate, sandstone and schalstein with limestone, peridotite, etc (Sasa & Koiwai 1960)

limestone 問問

M'目、ιsL

O

沿

Vd

(l

ae

br

piqu

戸しvvi

'H PA

l

o

-

-

qupqu

tye

aTLvl

e--o

nnf

・1JP川

d

i--

Mpm

U

0

1

scn 綿

C4

alpine windswept herbaceous community (Ohba 1974b), or rarely Pinus pumila scrub (Tatewaki 1928)

alpine windswept herbaceous community (Ohba 1974b) and subalpine tall herbaceous community (Ohba 1974a)

alpine meadow (Nakai 1919)

peridotite

31-49(53009'; Nikolajesk)

25-43(47003'; Cholmsk)

40-42(43015' ; Yamabe)

41-43(43024' ; Furano)

35-46(42010'; Urakawa)

ca 500 Southern Sakhalin, USSR below 700

43017' Mt. Kirigishi, Hokkaido

[K]

950-1000

42006' Mt. Apoi, Hokkaido

[M] o

chert, sandstone and limestone

11-20(35040' ; Kofu)

11-19(35031' ; Iida)

10-14 (39011' ; Wonsan)

13-16(43052' ; Changchun)

400-750

2700-3100

35040' Mt. Kitadake 「tlJH

「L

granite gneiss 41045' Mt. Kanboho, North Korea

[ 1 ]

2300-2500 制如何凶状C

* Warmth indices were estimated on the basis of the climatic data from nearby stations which are shown

in parentheses, and of a lapse rate of 0.550c per 100 m.

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September 1989 Journ. Jap. Bot. Vol. 64 No. 9 267

ense occurs mainly on limestone sites: the present new subspecies (subsp.

kirigishiense) grows on the north-facing (NW-NE), relatively steep (330-520)

screes near the summit of Mt. Kirigishi composed of limestone, and the speci-

mens of subsp. sαchalinense in SAPT were partially collected in “Sekkaiyama"

meaning a limestone mountain. C. miyabeanum occurs on the rocky and gra-

velly slopes consisting of peridotite. However, both C. hondoense and C. insigne

are indi百erentfrom these specific substratum in the alpine region. Therefore,

specific substratum such as limestone or peridotite seems to favor the relic

distributions of C. sachalinense and C. miyabeanum in the subalpine region.

Moreover, C. miyabeanum with many specific morphological characteristics is

considered to represent“ultrabasicosaxomorphosis" of C. sachalinense because of

the closely related petal characteristics and diploidy of the latter.

In plant community, the slopes where the present new subspecies grows are

covered with an ordinary tall herbaceous community and partially with the

mixed forest. The constituents of the former are Saussurea γiederi var.

yezoensis, Aconitum yuparense, etc, while those of the latter are Tilia ja戸onica,

Betula ermanii, Abi・es sachalinensis, etc. Whereas, the habitats of subsp.

sachalinense are“under珂oor of the valley forest" (Sugawara 1939). These

habitats of two subspecies of C. sachalinense are characterized by more heavily

snow-covered, humid, limestone-rich sites in the subalpine region.

The habitats of C. miyabeanum are chie丑ycovered with an alpine windswept

herbaceous community (Ohba 1974b) and rarely Pinus pumila scrub (Tatewaki

1928), which appear to be windswept, less snow-covered dry in winter, although

they do not attain the alpine region. The. habitats of C. hondoense are mainly

windswept slopes covered with an alpine windswept herbaceous community

(Ohba 1974b) and partially screes covered with a subalpine tall herbaceous

community (Ohba 1974a), and those of C. insigne were reported as“an alpine

meadow" (Nakai 1919). These habitats are clearly distinguished from those of

C. sachalinense‘

Key to the taxa treated

1. Ratios of length to width of petals mostly more than 2................. .2.

1. Ratios of length to width of petals mostly less than 2 ..................4.

2. Blades of radical leaves 1-2 cm long at fiowering; Ratios of apical petiolule

length to lateral one of radical leaves usually 1-2. .......... C. miyαbeanum

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268 植物研究雑誌第 64巻第 9号 平成元年 9月

2. Blades of radical leaves 3-9 cm long at flowering; Ratios of apical petiolule

length to lateral one of radical leaves usually 1-4 ........ (C. sachalinense)

3. Scapes mostly 17-31 cm long at flowering; Number of flowers per scape

mostly 1-2 ............................................ subsp. kz.rz.gishiense

3. Scapes mostly 23-36 cm long at flowering; Number of flowers per scape

mostly 2-3............................................. .subsp. sachalinense

4. Petals mostly emarginate at the apex, 8-12 mm long and 5-8 mm wide; Leaves

intensely glaucous on both surfaces............................C. hondoense

4. Petals mostly round or obtuse at the apex, 12-15 mm long and 6-10 mm wide;

Leaves not glaucous on both surfaces . . .. .. . . .. . . .. . . . . . . . . . . . . . . C. insigne

Callianthemum sachalinense Miyabe et Tatewaki subsp~ kirigishiense K.

Sato et Ko. Ito, subsp. nov.

Callianthemum miyabeanum sensu Ko. Ito in Journ. Jap. Bot., 57: 376 (1982),

non Tatewaki (1928).

A白nisad subsp. sachalinensi, sed di百erta quo scapis brevioribus paucira-

mosis,日oribuspaucis et foliis radicalibus petiolatis brevioribus.

Plantae rhizomatosae perennes glaberrimae, radicibus numerosis五liformibus

in sicco ca. 1 mm crassis. Folia radicalia 2-6 petiolata, petiolis plerumque 8-15

cm longis basi vaginatis albidis raro purpurascentibus; lamina supra viridescens

subtus pruinosa ternata lata ovata 1-11 cm longa; segmentis terminalibus ambitu

late ovatis ternatis, petiolulis-l0-53 mm longis 1-4-plo longioribus quam latera-

libus; segmentis lateralibus late ovatis trisectis vel ternatis; lobis trisectis vel

tripartitis breviter stipitatis vel sessilibus, lobulis tri自diset trilobatis vel flavel-

latim incissis. Scapi plerumque 17-31 cm alti folia radicalia superantes teretes

plerumque 1-2 ramosi uniflori. Folia caulina 1-3 superne fere sessilia inferne

petiolata usque ad 4 cm longa, petiolis basi vix dilatatis raro semiamplexicaulis;

laminis late ovatis tripartitis vel trisectis, lobis flabellatim incisis vel simplicibus.

Flores 19-27 mm in diametro, albidi praecoces; sepalis 5 plerumque ellipticis in-

aequalibus 4-9 mm longis 2-5 mm latis pallide viridulis vel albidis; petalis 5-10

anguste obovatis vel obovatis 9-13.5 mm longis 2.5-7.5 mm latis apice emarginatis

vel rotundatis basi rubro・aurantiacisnectariferis. Stamina numerosa, filamentis

linearibus 1.5-3.5 mm longis, antheris oblongis ca. 1 mm longis. Carpella 5-13, stylis brevibus recurvis. Achenia ovatoglobosa 3.5-5.5 mm longa 1.5-3.5 mm lata

supra leviter nervosa.

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September 1989 Journ. Jap. Bot. Vol. 64 No. 9

Chromosomatum numerus 2n=16.

Nom. Jap. Kirigishi~So (nov.)

Hab. on limestone substrates, Mt. Kirigishi (ca. 950-1000 m alt.).

Distr. Endemic and calciphilous.

269

Typus. in SAPT (Herbarium of Fac. Agr. and Environ. Sci., Hokkaido

Univ.)“K. Sato, 23 June 1986, Mt. Kirigishi, Prov. Ishikari".

The present new subspecies is very close to the typical subspecies but is

distinguished from it by shorter, few branched scapes with fewer丑owersand

shorter petioles of radical leaves. It is an endemic to Mt. Kirigishi, which

stands in Central Hokkaido and a calciphilous plant.

The authors are grateful to Dr. K. Samejima of Sapporo Gakuin University

and Dr. T. Nishikawa of Asahikawa College, Hokkaido University of Education

for their valuable suggestions. Our thanks are extended to Dr. H. Kanai and

Dr. Y. Kadota of National Science Museum, Tokyo for the kind permission to

use the collections of Callianthemum in TNS.

References

Damboldt, J. 1966. Chromosomezahlen einiger sudalpiner Endemiten. Ber. Deutsh.

Bot. Ges. 78: 374-375. Hara, H. 1935. Preliminary report on the fiora of

Southern Hidaka, Hokkaido (Yezo) IV. Bot. Mag. Tokyo 49: 9-10. Ito, K.

1982. New localities of Hokkaido plants (1). Journ. Jap. Bot. 57: 376. Kira,

T. 1948. On the altitudinal arrangement of climatic zones in Japan. Kanti-

Nogaku 2: 143-173. Kitamura, S. & G. Murata 1962. New names and new

conceptions adopted in our Coloured Illustrations of Herbaceous Plants of ]apan

II (Choripetalae). Acta. Phytotax. Geobot. 20: 204. Komarov, V. L. 1937.

Flora URSS 7: 54-57. Kurita, M. 1958. Chromosome studies in Ranunculaceae

X. Karyotypes and chromosome numbers of some genera. Rep. Biol. Ehime

Univ. No. 6: 12. Langlet, O. F. 1932. Uber Chromosomenverhaltnisse und

Systematik der Ranunculaceae. Sv. Bot. Tidskr. 26: 386. Miyabe, K. & M.

Tatewaki 1935. Contributions to the fiora of North Japan VI. Trans. Sapporo

Nat. Hist. Soc. 14: 73. Nakai, T. 1919. Notulae ad Plantas Japoniae et

Coreae XX. Bot. Mag. Tokyo 33: 49. Nakai, T. 1920. Anemone insigne

Nakai. In Matsumura, J.: Icon. Plant. Koisikav. 4: 85. Nakai, T. 1928.

Callianthemum insigne Nakai. Scientific Knowledge 8: 41. Nakai, T. 1930.

Vegetation of Mt. Apoi in the province of Hidaka, Hokkaido. 49. Nakai, T.

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Page 14: 料:A Calli nthemum C. sch linenseCalli αnthemum (Ranunculaceae) from Japan and its adjacent area, with reference to a new subspecies of C. sαchαlinense from Hokkaido, Japan

270 植物研究雑誌第 64巻第 9号 平成元年 9月

& H. Hara 1934. Callianthemum Novum Japonicum. Journ. Jap. Bot. 10: 49-

50. Ohba, T. 1974a. Syntaxonomische Studien ueber die Stauden丑urengeseI-

Ischaften der subalpinen Stufe Japans 1. Bull. Kanagawa Pref. Mus. Yokohama

7: 23-56. Ohba, T. 1974b. Vergleichende Studien ueber die alpinen Vegeta-

tion Japans 1. Carici rupestris-Kobresietea bellardii. Phytocoenologia 1: 339-

401. Ohwi,]. 1932. Symbolae ad Floram .Asiae Orientalis 6. Acta. Phytotax.

Geobot. 1: 301. Ohwi, ]. 1953. Flora of Japan. 527-528. Shibundo. Tokyo.

Rostovsteva, T. S. 1981. Chromosome number of some species of the family

Ranunculaceae 2. Bot. Zhurn. SSSR 66: 1751-1755. Sakai, K. 1935. Studies

on the chromosome number in alpine-plants 11. Jap. Jour. Genet. 11: 68-73.

Sasa, Y. & T. Koiwai 1960. Geological map of Sakhalin. Shimizu, T. 1983.

The new alpine flora of Japan in color 2: 48-49. Hoikusha. Osaka. Sugawara,

S. 1939. Illustrated flora of Saghalien with descriptions and自guresof Phanero-

gams and higher Cryptogams indigenous to Saghalien 2: 942-943. Karafuto

Shokubutsu Zushi Kankokai. Tokyo. Tatewaki, M. 1928. A new species of

Calliαnthemum from Japan. Trans. Sapporo Nat. Hist. Soc. 10: 79-80. Tutin,

T. G. et aI. 1964. Flora Europaea, 1: 208-209. Cambridge Univ. Press. Wang,

W. T. et al. 1980. Flora Reipublicae Popularis Sinicae 28: 242-246. Science

Press‘ Beijing.

* * * *

日本近隣のキタダケソウ属植物について,かつて“ヒダカソウ"として報告した北海

道蛙山産植物(伊藤 1982)を含んで, 分類学的検討を行なった。蛭山産植物は南サハ

リンのカラフトミヤマイチゲの新亜種,キリギシソウ(新和名)とすべきであるとの結

論に達した。キリギシソウはカラフトミヤマイチゲとは,花茎長,花茎当り着花数,根

生葉葉柄長などがより小さな値を示す点などから区別された。最も重要な形質として花

弁の長さ・幅比に注目し, 2亜種を含む種としてのカラフトミヤマイチゲと北海道アポ

イ岳のヒダカソウは本州北岳のキタダケソウや朝鮮半島のウメザキサバノオと大別され

ることを示した。次いで,花弁における凹頭花弁の割合と根生葉の展開時期は,世界的

にみても染色体数の違いに対応していることを示した。 2h=32の植物(ヒダカソウ,

ウメザキサバノオなど)は凹頭花弁の割合が低く,根生葉が花後に展開するのに対して,

2n=16の植物(キリギシソウ:初報告,キタダケソウなど)はその割合が高く, 花期

には展開している違いがあった。前者の特徴は後者から染色体数の倍化に伴って平行的

に生じたものであると考察した。根生葉における第 1次の頂小葉小葉柄と側小葉小葉柄

の長さの比(大井 1953)は, ヒダカソウが特徴的な値を示した。 ヒダカソウはすでに

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September 1989 Journ. Jap. Bot. Vol. 64 No. 9 271

知られているさらに多くの特徴を持つので明らかな種と考え,さらにカラフトミヤマイ

チゲから染色体数の倍化によって生じたものであると推定した。生育地の比較は,最も

重要とした花弁の長さ・幅比による分類と対応していた。新亜種を含む種としてのカラ

フトミヤマイチゲとヒダカソウは亜高山帯の,それぞれの石灰岩とかんらん岩の特殊岩

に結びついて生じ,一方,キタダケソウとウメザキサバノオは特殊岩と無関係に高山帯

に生じることを示した。

0高等植物分布資料 (127) Materials for the distribution of vascular plants in

Japan (127)

0トクガワザザ Sasa Tokugawana Makinoトグガワザサはササ属アマギザサ節に含

まれ,主な分布地は神奈川県箱根山ならびに伊豆半島天城山系風速峠周辺に限られてい

る。分布の上でも,形態の上でもミヤマクマザサ Sasa Hayatae Makino に似るが,

それよりも分布域が狭く,また稗鞘に聞出する長毛を密生する点で区別される。

筆者らは1986年12月18日に八溝山系の地質ならびにササ類の分布調査を行い,その途

上で栃木県那須郡馬頭町脇郷においてトクガワザサの群落を発見した。トクガワザサの

分類形質である稗鞘の開出長毛は冬期には脱落しやすく判別が困難なことがあるので,

正確を期するため1987年 5月29日に再度同地を訪れ,新しく伸びた植物体の稗鞘の毛を

確認し 同定した。群落の生育状況は極めて良好で, 手早の高さ 220cm,直径 9mm,

葉の長さ 31cm,幅 7cmに達するものがある。稗鞘は節聞の長さの半分以下で稗の上

方のものは所々で繊維状に細裂する。節は膨出し,稗の上方で、盛んに分枝する。葉裏に

軟毛を密生し,古い葉には細くくまどりがみられる。葉鞘は無毛,葉舌は低い山形,肩

毛は放射状に発達する。証拠となる標本は宇都宮大学教養部植物標本庫に保管されてい

る。生育地は那珂川の支流・大内川より女体山(標高 409.3m)の南東面へ入り込んだ

沢に面した痩せた尾根の北西斜面で,中生代ジュラ紀の砂岩・泥岩互層に貫入した馬頭

岩体(ハンレイ岩一花崩閃緑岩複合体)からなり,標高 150m から 210mに位置して

いる。周囲はほとんどスギ, ヒノキの人工林で占められているが,わずかに取り残され

たように急傾斜地上に成立するシデ類,カスミザ、クラやタカノツメなどからなる落葉広

葉樹林の林床である。斜面の下には小川が流れ,水田によって自動車道より隔てられ,

湿潤な環境が保持されている。周辺地域をくまなく調査した結果,他に女体山の山腹の

北斜面の一部,標高 150mから 200mにかけての落葉広葉樹林林床にもトグガワザサ

の群落を発見した。これらは, トクガワザサの分布地の北限である。

一方, ミヤマクマザサの分布地の北限は福島県西白河郡表郷村犬神であるが,その生

育地は鈴木貞雄博士によって写真撮影された後, 1970年頃のダム建設によって水没L,

-15ー