58
A PhytosociologicalStudy of Serpentine Areas in Shikoku, Japan 'By Tsugiwo Yamanaka I. Introduction It has been well known that the plant life on serpentine and related rocks is strikingly different from that on other rocks. Serpentine is, strictly speaking, essentially a magnesium iron silicate formed by hydro- thermal alteration from ultrabasic rocks, such as peridotite. However, these ultrabasic rocks are grouped together as serpentines by almost all the botanists because of the similarity of the specific e仔ect on the flora and vegetation. Therefore> the 'areas treated in the present paper include the peridotite as well as the real serpentine areas. Ultrabasic rocks occur in many parts of the world. Since Pancic described the flora on serpentine of Serbia in 1859・,the flora and vegetation of serpentine areas have been reported by a large number of authors from Europe・North and South America・New Zealand, New Caledonia, South Africa, Indonesia, and Japan. In Japan, serpentine areas have been well known by botanists as interesting places on account of their very rich and characteristic floras. But no report with particular regard to serpentine appeared until Yoshinaga paid attention to the peculiar flora on serpentine in K6chi Prefecture (Shikoku) in 1914. Thereafter, in !1918, Nishida mentioned the peculiarity of the serpentine flora on Mt. Yupari (Hokkaidd). Up to the year ・1950, Tatewaki's publications on the flora and vegetation on serpentine in Hokkaido were important contributions to the ecological and phytogeographical study. From the year 1950, the plant life on serpentine・ has been noticed by number of botanists. Kitamura and his co-workers (1950-57) have studied serpentine floras in HokkaidS, Honshu, and Shikoku. Toyokuni (1955―58) has described・ the floras of serpentine areas in Hokkaid6. Hattori (1955, 1957) reported the hepatic floras on serpen- tine of Mt Apoi (Hokkaid6) and Mt. Hayachine (N. Honshu). In Shikoku, a little was known about the serpentine flora, though Yoshinaga's report mentioned above was the first one in Japan. In 1935 and 1936, Yoshinaga also noted the serpentine flora in Tokushima Prefecture, and the same district was studied' by Kitamura and Murata in 1952. And then notes on the flora of Nishikiyama in K6chi Prefecture by Nakai appeared in 1942. Apart from the above, the only contributions to the study of the serpentine vegetation in Shikoku are publications made by the present writer (1950 -59). In 1948, at the suggestion of Prof. Yoshiwo Horikawa of the Hiroshima University, the writer, studied the vegetation of Mt. Higashi-akaishi in Ehime Prefecture and of Mt. Shiraga in Kochi Prefecture and made it the subject of his graduation thesis. These two mountains consist of serpentine, and the writer's attention was attracted to the peculiarities of the plant life on serpentine. In 1949, while studying the serpentine

A PhytosociologicalStudy ofSerpentineAreasin Shikoku, Japan · A PhytosociologicalStudy ofSerpentineAreasin Shikoku, Japan ... This district is situated in Yawatciharha city and Mikame

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A PhytosociologicalStudy of Serpentine Areas in Shikoku, Japan

      'By

Tsugiwo Yamanaka

                     I. Introduction

 It has been well known that the plant life on serpentine and related rocks is strikingly

different from that on other rocks.

 Serpentine is, strictly speaking, essentially a magnesium iron silicate formed by hydro-

thermal alteration from ultrabasic rocks, such as peridotite. However, these ultrabasic

rocks are grouped together as serpentines by almost all the botanists because of the

similarity of the specific e仔ect on the flora and vegetation. Therefore> the 'areas treated

in the present paper include the peridotite as well as the real serpentine areas.

 Ultrabasic rocks occur in many parts of the world. Since Pancic described the flora

on serpentine of Serbia in 1859・, the flora and vegetation of serpentine areas have been

reported by a large number of authors from Europe・North and South America・New

Zealand, New Caledonia, South Africa, Indonesia, and Japan.

 In Japan, serpentine areas have been well known by botanists as interesting places on

account of their very rich and characteristic floras. But no report with particular regard

to serpentine appeared until Yoshinaga paid attention to the peculiar flora on serpentine

in K6chi Prefecture (Shikoku) in 1914. Thereafter, in !1918, Nishida mentioned the

peculiarity of the serpentine flora on Mt. Yupari (Hokkaidd). Up to the year ・1950,

Tatewaki's publications on the flora and vegetation on serpentine in Hokkaido were

important contributions to the ecological and phytogeographical study.

 From the year 1950, the plant life on serpentine・ has been noticed by a number of

botanists. Kitamura and his co-workers (1950-57) have studied serpentine floras

in HokkaidS, Honshu, and Shikoku. Toyokuni (1955―58) has described・ the floras of

serpentine areas in Hokkaid6. Hattori (1955, 1957) reported the hepatic floras on serpen-

tine of Mt Apoi (Hokkaid6) and Mt. Hayachine (N. Honshu).    `

                                            メ In Shikoku, a little was known about the serpentine flora, though Yoshinaga's report

mentioned above was the first one in Japan. In 1935 and 1936, Yoshinaga also noted the

serpentine flora in Tokushima Prefecture, and the same district was studied' by Kitamura

and Murata in 1952. And then notes on the flora of Nishikiyama in K6chi Prefecture by

Nakai appeared in 1942. Apart from the above, the only contributions to the study of

the serpentine vegetation in Shikoku are publications made by the present writer (1950

-59).

 In 1948, at the suggestion of Prof. Yoshiwo Horikawa of the Hiroshima University,

the writer, studied the vegetation of Mt. Higashi-akaishi in Ehime Prefecture and of Mt.

Shiraga in Kochi Prefecture and made it the subject of his graduation thesis. These

two mountains consist of serpentine, and the writer's attention was attracted to the

peculiarities of the plant life on serpentine. In 1949, while studying the serpentine

vegetation in the vicinity Qf・ K6chi c‘ity, the writer made collections there with Prof.

Siro Kitamura, who, visiting K6chi^ city for the first time, was much interested in the

flora on serpentine。 Under thぶkind guidance and encouragement of Prof. Horikawa and

Prof. Kitamura・the writer has continued to study the serpentine vegetation in Shikoku

during the last ten years. Consequently・811 the serpentine areas in Shikoku, with the

exception of some smalトoutcrops, have been studied from a phytosociological poit of

view. In the present paper, the writer would outline the results of his study.

 Before going further, the writer wishes to express his sincere thanks to Prof. Yoshiwo

Horikawa, under whose kind direction and guidance this study was carried out. The

writer is also indebted to Prof. Siro Kitamura for his encouragement and invaluable

                 -           Isuggestions. The writer's thanks are due also to Prof. Tokio Suzuki of the Oita Univer-

sity for his criticism in the course of‘this study. Further the writer wishes to thank

Prof. Takeo Sawamura and 'Mr. Chiaki Sakamoto of the K6chi University for their

helpful suggestions in regard to ・petiでological and chemical problems of serpentine. For

the identification of a large niimbfきr of’plants, the writer takes this opportunity of ex-

pressing his thanks to the following taxonomists : Dr. Tisaburo Ohwi of the National

Science Museum, Mr. Yasuichi Momiyama of the Research Institute for Natural

Resources, Mr. Tetuo Koyama of the University of Tokyo, and Mr. Gen Murata of

the Kyoto University. Lastly, particular thanks are due to the various persons who CO-

operated with the writer in the field.

 This study was carried out partly by the Grant in Aid for Developmental Scientific

Research from the Ministry of. Education.

   II. Notes On the flora and vegetation of serpentine aieas in Shikoku

 Shikoku is an island which is situated between lat. 32°45'~34°25' N. and long. 132°~

134°45'E. , extending Over 18773 square kilometers. The highest peak in the island is Mt.

Ishidzuchi rising 1981 mよabove-sea-level. Serpentine and related rocks crop out in the

Nagatoro and Chichibu zones and occur in Ehime, K6chi, and Tokushin!a Prefectures.

In the present paper, adjacent serpentine areas are conveniently grouped together into

districts, such as the Yawatahama", Yusuhara, Kdchi, Tokushima, Mt. Akaishi, and

Mt. Shiraga districts (Fig. 1). 1                       ,

   A. Yawatahama District

 This district is situated in Yawatciharha city and Mikame Town, about 50 km. southwest

of Matsuyama city, containing the largest outcrop in the southwestern part of Ehime

Prefecture.            ’  ヤ

 1. Korodokihana (Yamanaka 1959)

 Serpentine crops out on the borders of Yawatahama city and Mikame Town. The end

                       ・of this outcrop forms a steep slope directly facing the sea, the vegetation being very

sparse in such places (Plate I). However, stable places overlaid by weathered materials

are covered with sとrubs mainly dominated by Quercus phylliraeoidesatta\ning0.5~1m.

                          Fig.1

A. Yawatahamか・District. 1- Korodokihana, 2. Mikame-Futaiwa.

B. Yusuhara District. 3. Shimagawa, 4. Ochimen.

C. K6chi District. 5. Mt. Yokogura, 6. Nishikiyama, 7. Namekawa, 8. Engydji。

  9. Osaka Pass, 10. Okdyama, 11. Aburaishi.

D. Tokushima District. 12. Jinry6, 13. Shirigi Pass, 14. Ryu Pass, IS. Sakashu.

E. Mt. Akaishi District. 16. Mt. Nishi-akaishi, 17. Mt. Higashi-akaishi, 18. Mt。

  Akahoshi, 19. Tomisato.

F. Mt. SI!iraga District. 20. Mt. Shiraga・

a. Matsuyama, b. Yawatahama, C. Mikame, d. Uwajimai e. Yusuhara, f. Ino,

g. K6chi, h. Tosa-Yamada, i. Tokushima, j. Imabari, k. Niihama, 1. Takamatsui

m. Motoyama・n. Oonogahara, o. Mt. Ishidzuchi, p. Mt. Kamegamori, q. Mt. Ishidate,r. Mt. Tsurugi.

in height. As shown in Table 1, the・ floristic composition is not essentially different

from that of Q・uercus piりlliraeoidesscrubson other rocks.

         Table 1. Quercusμびlliraeoides cornmunity of Korodokihana

                (20m., N20W,・5°;2〉く2m.)

Shrub stratum

  Quercus l)りiliraeoides

  Eurya japonica

  Pinus l仙必∂ダgii

  PitiosiJorum tobira

.い(μacciniu附加acteatum

  Rosa wichuraiaれa

  Eurvo emarginaia

  S竹til ax china

  j?ゐ面面endron wりrichii

  Ligustずurn. iaponicum

  Fraxinus sieboldiana

4CM I―I I―I

・●一一●

4321十+1

(十)

 +

421十十十十奉・

41CM.CN*

・r-<i―!・+

●・・一一

 Herb stratum

DicrdnoMeris dichoioma

Miscanihus sinensis

Farfugiurn japonicum

Polygora japonica

ぷglダis spicata

Themedd iaponica

Solidago viダgaurea ssp。

          asiaitca

ChTNsaポhemum ornatumV。

         ashizuriense

Su・ertiaiaponica

Paederia ^cand&n^ v. mairei

23・十・・

22十・、十・

32・・十1

32・ヽ・ss″

22十十・・

十e一

・一一

●●●

一一

・+

4 of the K6chi University Vol. 8, No. 10

 2. Mikame and Futaiwa (Yamanaka 1959)

 This hillock region is well cultivated, and the vegetation in the serpentine area is

also disturbed by human agencies. However, pine forests chiefly composed o? Pinus

densifloraand Pinus thunbereiiate considered to be proper to this area. The forest

dominated by stunted Pinus de7isiflora,which occurs in rather flat places covered with

shallow soils, shows the following composition*:

            Tab\e 2. Pinus(iensiμOTQcommunity of Mikame

                (265m., N75W, 5-10° ; 5×5 m.)

 Shrub stratum

Pinus densiμOTQj?み∂tdodendronweyri凶日E�即畑加nicQSmiiox chinaAmelanc力i∂ダasiaiicaLespedeza homoloba

Rosa onoetF昭ara maれfchuricQRfius frichocarbQLisusitum iaOomcurnDibiomorpha sikokianaメ1di細心t>hne laneφ刄iaViburnu竹1etos≪m

 Herb stratum

Pteridium aquilinum v.           taliusculum

Arunぷnella hirtaMiscawlfius stnensis

言)

混)

づコ

4ヤ)

 呂

1(十)漕)

 1

  Herb stratum

SolidaSO virRaurea ssp.asiatica

ChrNsoれthemum ornaれitnV.

               ashtzuTtenseDiblrnnorpha goni>i

Pairiれia villosa

Paederia scandcnsV. matダ・

Carexspp・

i^iola ovato-oblongaPolygara japonica

Isodon inflcxus

Colomagrostis arundinoceoV.

               brachyiricha

CymbopoRon goerinSiiThemeda japonica

Hziefopappus hisiJidus

/1&μia serf:と2tav. tomentosa

3 other species

 1

 1

 ●

 ●J●

 ●

 ● ●

 ●

 ●

 ●

 1

 1十十

十 1

+++

十十十十十

  Many shrubs and herbs occur in the lower strata, but it is a remarkable fact that there are

never or rarely found many serpentine-characteristic plants belonging to Abelia, Ferりa,

Pourthiaea, Fraエinus, Stephanandra, Ktikianthus, Berberis, Sai£ssurea, Bup加urmn.

Sにb・tosa, etc. On steep slopes where serpentine crops out, forests accompanied by Pinus

蔭山ibergiioccur, and the slirub stratum is predominated by Quercus phylliraeoides. An

example of them is as follows :

 1st tree stratum : Piniisftansiμma 3(十)*, Pinus thunbergii 3(十)

 2nd tree & shrub strata : Quercus phyぶraeoides 4 (1), /?hus javanica 2 (十), Jiosa onoei 1(1),

  Z)iμomofpha sikoゐtana 1 (十),S。tilax china 1 (十),Fagara竹laれtchurica十(十),召erbefis thunbergii

  十, Mallotus加加nicus十, Gloch哨Z卵面部atu四十,Paerteria scandeれs var. 。lairei十,Dioscorca

  graciぷ四α十(十)

 Herb stratum : Miscanlhus sinensis 3, Arundinella hirta 2, Bot力riochloa psr・viflora 2, Chぴsのzl加州g加

  ornatum var. ashizuriense 1, Cymt?idinm virescのIS 1, Pteridiμm aqui山畑m var. latiusculuタフ1+,

  レ^iola ovato-oblonSa十,几面on inflexus十, Patrinia villosa +,Forfugium iapomcum'十,

  Artemisia iai)onica十,Themeda jofiontca十,Liaustれim jotonicum十,Pittosjiaれ4m tobira十

 Such forests also appear on other rocks in coastal districts of Shikoku, and ・ the

characteristic feature of the serpentine area is not found in this district.

 B. Yusuhara Distri≪21t

Yusuhara is a village which is situated about 70 km. west of K6chi city. The serpen

普 Figures in parentheses indicate the dominance in the lower strata

tine outcrops occur in Ochimen and Shimagawa, running from east to west, to th‘e south

of the Oonogahara limestone plateau. Among these, the largest outcrop appears in

Shimagawa, east of Mt. Amadzutsumi. The chromium content of the rock is rather high,

and quarrying was undertaken in Shimagawa・-

 ろShimagawa (Yamanaka 1956, 1959)

 Two outcrops appear about 2 km. apart and are situated between 700 and 1000 m.・

above sea-level. The vegetation is not different from that of Ochimen, and stunted

pine forests are found throughout the outcrops (Table 15 and Plate I). However, the

growth of Pinus densポora "is rather excellent in places overlaid by deep soils. In such

places, the forest represents the following composition :

 1st tree stratum こPinus (iensi/lora S

 2nd tree stratum : Clethra l,αダbinervis 3 (十),£i�eya obtusμoba 2(2), //・ex ^errata2(十), Vibuダs。z

 ヽgずosum 2(十),Magnolia obovaia2, Quercus setiata 1(十), Rhus tnchocarba 1(十), Piびis japonica

  1(十),j?加面庇�加。ダgli。latum十(1), Ilexびenata十(十)

 Shrub stratum : Sssamoifpha purpufascens5(十), Abetia set・■rata 1(2),Cetihaloiaxus haYtingioれio

  十(十)。Lindera sericea十(十),` Srnilax china十(十), Berbがis thunbeなii十,PoMthiaea villosa

  var. zollingがi十,Etも071タmus alatus var.tottt ndatus+

 Herb stratum : Carex spp. 2,Lepiogmmma tofta 1,Athyfium nifionicum十,Trt picfosixfmum

  加加戒cμ加十, Aster ageratoides ssp. amt・Eextfolius十,Cirmamomum iaponicum十, Fagara

  mantchurica+, Acer si必必沁zタ<tum +                 ‥

 Grassy places dominated by Miscanthus shiensis appear as the result of human agen-

cies, and there are found Pteridiuni aq‘uilinumvax. la£iusculum, Arundinella hirta.

Allium thunber'gii. Gen£iana scabravar. buergeri, Svuertia pseudochinensis. As£er

agerato・はes ssp. a‘・'ivbleエザolius.Mete・ropappus hispidus, Saussi£rea pulchella,  etc.

 From the above, it can be concluded that the serpentine vegetation of this district is

similar to that in the K6chi district, though there are some floristic differences between

them. The most remarkable fact is the occurrence of Saussuj?a scaposa which is not

found elsewhere in Shikoku.

 4. Ochimen (Yamanaka 1956)

 Scattered outcrops occurring in this area are characterized by open pine forests (Table

15). It is characteristic that Gynm心なr pyginae略 is abundant in moist places under

p】anted Cりptomeria iaponica.

  C. Kochi District

 This district includes seven main serpentine areas which occur in 10w altitudes in and

near K6chi city. These outcops run from east to west along the southern part of the

Chichibu zone and have a peculiar flora and . conspicuous vegetation.' Therefore, some

endemic and noticeable plants were described from serpentine areas of this district. They

are Coりlopsis spica£a. Hyper icun・1 tosa。ise. He£eropappus hispidus ssp. Iゆ£ocladus

(=H.leptocladus.), Saussurea nipponica ssp・ yoshijiagae(=f S. yoshinagae), and

JLpiniedium tr球固atohinaはm.

 Climatic climaxes in the non-serpentine areas of this district are warm-temperate

forests mainly dominated by Castanopsis ( =Shiia), Cyclohal。lopsis(evergreen Quercus),

Machi臨s, etc., whereas open ・pine forests appear on serpentine.

 5. Mt. Yokogura (YaJnanaka 1955)

 Several small serpentineぺjutcrops occur on Mt. Yokogura and in its・vicinity, about 28

                4km. west of K6chi city. One C)f these outcrops is found at an altitude of about 200 m.,

where the vegetation is principally constituted by Pinus densiflora,Poiびthiaea villosa

var.zollingeri,etc., andトthe following floristic compositions are found:

 A. Pinus densifloraforest

 1st & 2nd tree strata : Pinusdensiμma4(十),Ilex tied鉢肴cutosa2(2), S。nilax china 1(2),Rfius

  trichocarpa十(2); Pourthiaea i・μlosa var. z。Hingeダi十(十),びiburnum erosMirt十(十), Clethra

  barbinervis +             フ●

 Shrub stratum : 乃gダis加加nica 3 (2), /?/7�・dendron wりrichii 3 (1), Abelia ssが■ata 3(1),Rho-

  dodendronkaempfefi7(ミ十),レ'accitヌig加S。nallii Var. glabrum 1(十),£jy・nia elliμica 1 (十), Euりa

  加加nica 1, /?∂Sa onoei 十(1),Ficus eUcta 十(十), Millettia加加㎡Cα十(十),Cりptomefia

  iatonicQ十(十), Ilex crenata十 ノ  ,

 Herb stratum : Mtscanthussi・Iれensis2,Pteridium aquilinμ;w var. latiusculum 1, Os。zz。ldα知加㎡cα

  1,  Afdisiojaponica1, ' Lepisoれ4s thunbergianus十,0。ychium加加タlia四十,Cμ

  g叱y函がi十, Neolitsea sがice皿十・ 。。‘

 BPO・Ur£h.iaea villosavat. z-0111ngeri scrub

 2nd tree & shrub strata : PoUrthiaeaひi尚g vat. zoilinfiぴ■i5(2), Myダica rubra 1 C十),Piwus

  densiμora半(1),FogOTQ manlchuticμ十(十),Rhus trichocarpa十(十), Ligusfれ4精細fionicum

  十(十),Coccui≪s triiobus十・(十),Paedびia scandensvar. maitei十(十),Smilax china十(十),

  RhododeTidronkaempferi十,j?加ゐ加面ダ∂71 gりダichii十

 Herb stratum : Oplis・enus undutatiかlius var. 知加戒as 2, 召rachypodium syliなiticum var. mise畑加

  1,ルfiscanihus sinensis I, 戸lgダidium aqねilinum var. latiusculuタ?Z十, Viola violaceα十, PoWgaTQ

  加加雨Cα十, Solidaao!Dingaureasspレ卯i,atica十,Sθなhum nitidum var. majus十,Cぴμθポeria

  japonica十,Neolifseo seticea十,Rosaonoei十,Rhus trichocarpa十,灸几ぶotus加加nicus十

  〇ther outcrops are found in Kirimigawa, Tokoroyama, etc., but typical serpentine

vegetation is poorly developed there.

 6. Nishikiyama and its vicinity (Nakai 1942 ; Yamanaka 1950, 1951, 1952)

 Large outcrops of serpentine ‘appear in Hidaka Village, about 18 km. west of Kochi

city, and the highest part of thふS・erpentine area reaches about 300 m. above sea-level.

This area is well known for the typical serpentine vegetation markedly different from

the non-serpentine areas, and is characterized by stunted pine forests with Knkianthus

μΓiilattjsvat. japonicur.(Table' 16 and Plate I). Many shrubs and herbs which are

characteristic of serpentir!e ・are ・also fiDund there.

 7. Naraekawa (Yamanaka 1953, 1り56)

 Namekawa includes Hashiradani, Harihara, and Jori. Serpentine crops out from about

50 t0 300 m. above sea-level. This ' area is also covered with pine forests which are

                 tusually open and stunted (Table 1・6). Though the floristic composition and physiognomy

of the vegetation of this area・resemble those of Nishikiyama and Engy6jV,  Co7-μopsis

spicaは\s not found, and Knkianthus  pcrulatusvar.japonicuf, IS abundant in places.

Grassy places and barrens are also found' in・sevei‘al parts where serpentine・characteristic

plants, such asHetercゆapus hiゆidus ssp・leptocladus,Scabiosaja夕onica,etc., are・

frequently found.                               ,

 8,‘Engyoji and its vicinity (Yamanaka 1950, 1951, 1952)

 The biggest serpentine outcrops in Kdchi city occur in Engydii and its vicinity, in-

eluding Jinzenji, Mitani, Shibamaki, Rendai, and Ori, ranging from about 50 t0 400 m.

above sea-level. The composition of the rock varies considerably, namely, MgO 35~43,

SiO2 30~45, CaO trace: Sakamoto (1955) reported the following ・ values from ・ Engydji :

 MgO 35.27, SiO2 44.00, FeO十Fe2O3十CrzOs 6.91, AI2O3 2.20, NiO 0.25, CaO tr.,

1 g loss 12.00.

 Though grassy places and barrens occur here and there【Tab】e 3 and Plate n), the

Table 3. Afundinaria pygmaeacommunityof Engy6ji

        (80ni., N40W, 30°; 5χ5 m. )

 Shrub stratum

Arundinaria Cygmaea*

Abelia serrataFねxinus sieboidianaCoりlopsissi>tcataPinus densiμora

Pertva glabrescensDiplomorpha siねokianaViburnumぴosumLtgustrum iaponicumRasa OI如etSmilax chinaLespedeza bwずg6ずiDioscorea graciUimaRhododendron locyrichiiRfius trichocafpa         `aglみダabarbiタzsダvis

3 other species

 Herb stratnm

E芦田edium tダifoliatol・ig細田Carex spp.゛“Carey.ciiiaiomar ginaia\^iolaviolaceaChionogrothis japonicaSaussurea nipponica ssp.voshi?tagaeOsmwuda japonicaCarei duvalianaTitりmalus sieboldianus

Gaitu辨知gonanihMmPaedびia scaれdensV. 田αiダeiSa峨口nipponicaAダelicaタlutaタ15Astef scaberMiscanlhus siれensisArundinella hittaBerbetis thunbeずgiiPoりSafa iaponicaStoei'tiafiseudochinensisSerfatula cofnata ssp. insulaダisSanButsorba oJ3^ctnaUs?1∂ダidiumaauilinum v. latiusculu??1

4 other species

  5(1)  3(1)  4(十)  2(1)   (十) 十(十)   (十)   (十)

  1(十) 十(十)  1(十)   (十)

   1   3

   十   +

   十   十

●  ●   +   +   +   十

   十   +   +   十

   +   +   +

3(十)示)ブ

寸)

-j本

-1i

 5(1) 3(D I(十) 3(1) (十) (十) (十) (十) 1(十)

十(十)

 (十)

  2 + +

 十 +

  2  2

 十

 十

 十

5(1)

2(1)

2(十)

2(十)1(十)

十(十)

十(十). 1(十)

 1(十)

十(十)

十(十)  2

  (十)

十(十)

  ● I 十(十)

   3

   2

   2

  +

  +   1

  +

  +

   1

   ●  '   +

  5(1)  ・3(十)

  3(十)

  2(十)

   (十)  十(十)

  十(ナ)

   十一

   (十)

   で  1(十)

  1(十)  十(十)

  十(十)

   3

   2

   +

   +

   + 、   1

   十

   +

   十

   1

   1

・・  十

゛ In the present paper, y1な丿㎡iタ1αダiflpygmaea includes vat. glabra.

昔昔 Cαダ6spp. include C. multifolia and C. subdita.

8Research Reports of the K6c・hiUniversity Vol. 8, N0.10

vegetμi‘on 01 this area is essentially characterized by stunted pine forests accompanied

by a large number of deciduous shrubs, herbs, grasses, and sedges (Table 16 and Plate

n). Among these plantsニthe abundance of Coりlopsis spicata is most characteristic.

Moreover, such serpentinicolous relics and serpentinophytes as HyμΓic征m tosaense.

He£eropappushiゆtdiis ssp. /ゆto cladus,and Saussurea nipponicassp.yoshhiagae are

abundantly found there。

 9. Osaka Pass and its vicinity (Yamanaka 1950, 1951, 1952)

 The Osaka Pass and its vicinity have large serpentine outcrops on and near the bound-

ary between K6chi city and Ok6 Village, and the highest part reaches about 160 m. above

the sea. Though the pine forest is the original vegetation of this serpentine outcrop

(Tab!e 16 and Plate n)j vast areas are now covered with the community dominated by

Ar£もndinaria pymnaea and many other grasses, herbs, and sedges as the result of the

deforestation, fire, grazing, etc. An analysis of the community in such grassy places

is presented in Table 4.

Table 4. Arundinanapygmaea community of Osaka Pass

         (50m., N20E, 5°;1χl m.)

Aれindinariapsgmaea

Miscanthussimれsis

DiplomorphasikoMana

Si)iroeanenJoso

Lespedezahomoloba

y1ダμ卿石タ詔lla hiダ1α

Bupleurμm falcaium

            komarowi

Sanfiuisorbaoが^c'tiialis

Tkalictnemminus

hchaemumぴassiftes

j?osa onoei

532131 1十十十+

532221 1十十十+

532221 十十十十+

532211 1十十十+

522211 11+++

Abelta serrata

Saluiaiaponica

Platタcodon grandifIOfus

Carex subdtta

Cymboftogongo≪ringti

Hypericum tosaense

PolyeoTQjQ知れica

Cyna蹴:hum paniculatuタ?1

Scabtoso ja如れica

Sotidofiovl-TRaureassp。

           asiatica

1・・・・・・・+

・・十十・十十十・

・+・+●一・s一

十十十s・s一e1

1・・・十●・●s

 Floristically, a noteworthy fact is the occurrences of such serpentinicolous relics,

serpentinophytes, and disjunctive e\emeTits as Coryloかsisやicaは, H.ypericum.tosaen'se,

Heteropapus辰砂idusSSpバゆtocladus, Saussurea nipponicasspべyoshindgae.Kenierocallis

thunberg・a, Veratruni maackiivar.reymondianum, etc・

 10. Okoyama (Yamanaka 1950, 1951, 1952)

 Okdyama is a small hill consisting entirely of serpentine, reaching 97.6 m. above the

sea. The composition of the rock is as follows : MgO 37. 91, SiO2 36.30, Fe2O3十AhOs

11.05.

 Pi/tits dejisiflora also dominates in the forest (Table 16 and Plate Ⅲ), but Arundinaria

pygmaeaoccupies large areas as the result of human agencies.

 11. Aburaishi (Yamanaka 1953)

 The most eastern serpentine outcrop within・ this district is found in Aburaishi, about 17

km. northeast of Kdchi city. This serpentine area forms an isolated hill which is situated

between 80an!j150 m. above sea-level. The vegetation of this area is essentially charac-

terized by the stunted pine forest, representing the following floristic composition :

 2nd tree stratum :JPinus

densiμora .5(2). | ,.    . .......

 Shrub stratum : Arundjnorio pygtnaea 5(2),£yonia elliμtea 3(1), R加dodendron uiりrichii 3 0),

  Ilex Pedunculosa 2(1), Rhododendron macrosepalum 2(1), Dioscorcagracitlima 2(1),Poutthtaca

  villosa var. zoUingeri 2(十), Clethra barbinervis 2(十), Myrica rubra 1, Prunus・畑田dsakura 1,

  Liaustrum iaponicum十

 Herb stratum : Dicra,toMerisdichoioma 2, Carex dliaiomarginala2, Siepha,landta tncisa2,Epi-

  刀雲ediumんitamurαnum 1, Miscど7nthus sinensis 1, Akebia tfifoliata 1, Rosa onoei十,Ardtsia

  iaponica十, Cinna。1.。zz。,1加加nicumナ,Lespedeza ho竹1010ba十,Eurya iaponica十

 But there are shrubby or grassy places resulting from human agencies, and floristic

compositions in such places are shown in Tables 5 and 6.             ノ

Table 5. Scrub on serpentine of Aburaishi

    (120m., N55W, 25°; 5X5 m.)

Dicranoi)teris dichotoma

Aru?tdinatia i>y&maea ・

Queずcus sefrata

Cleihfabarbinems

Pinus densiμor a

Les軸心za hornolobo

Rhododendron竹aacroseCaium

Rhododendron tu6タrichii

Rosffl・ono£t

S附・Hax china

Lyonia eUゆtica

Euりa iaponica

Diplomotpha sifeofeiona

532221+11121e

5422221.2++e21

3。53.2+1212+2●-

5432223111211

5322222111221

びoccinium bradeatum

Myrica y幼ずa

Ilex t>edunculosa

Dioscofea gれ

Miscanthus sinensts

Pter・idium aouilinum v。

          latiusculuポ

Ilex crenata

Pteris japonica

Carex mira

G6れtiana scabraV. bueずgがi

Aダundi?lella hiダ1α

Milletia iapo㎡ca

Tab\eらArundinaria pygmaeacommunity of Aburaishi

      (120 m.,・ N55W, 25°; lχ1 m.)

Aれindinaria tvgmaea

Carexciliaiomarginaia

Carexmira

Sanguisorbaoがcinalts

Scabiosaiaponica

召e竹lerocallisthunbぴfiti      ・

DiplomorphasikokianaMiscanthussinensts

Afundin&llahirta

Viola 01なZIθ一助1θタzgαSaluteiaponica

Rosaonofii

Geniia?ia scabrav、fauergeri

ji?hododendron weyrichiiCvmbc

Solifiago tiiygoureossp.asiatica

Srnilaxchina

Dioscoreagracillima

召召をダ∂pappus hispidus ssp. leptocladus

Adenop加忽triphylla v.ialJonico                 t、lncげoHa

Swertia pseudochir認nsis

Ptnusdensiμora

Bupleuれim falcaiumv. komatowi

32十+2十・・・十●・一・―・el一

一一・

531十十・十一・一●一十十●●s一・ 一一●一

52+112+11+

1・十・・・十● 一●一e

3321・―十+3十●一�●一一一一十

521131+13ゆ

●elS

+・・・

SS一番

―・―十

一●魯●

2211

12・―十

十21、・十

十●一一・・+

十・・’・・.

十・‘・・・.

・1↑1十十・

. -<<^ . . . .

413212+22+―+ls’s l●一一 ●・一●

43122十十2・十―●・s・一●一一 一●十・

5十十+1十222・十・・・・・・・・ ・十・・

413222+22sll■

・'―'・・● s’一・+

53322+133

一●●II―f

一一IS

 Floristically, the abundance of Heine-i・ocallist}iu7ibergiiand Ca7~eヱフnira is noticeable

in this serpentine area. Though Coりlopsis spicata, Knki。1£flits perulatusval・i aponiレcus.

etc. are not found, such plants as Myperidぷ111tosaense.Saussi£rea nipponicasspべyosh-i-

imgae.et.c. occur in this area.

10

Research Re

Altitude (m.)ExpositionSteepness

Table .7.‘ySerpen‘tine vegetation of Jinry6

・320N45W

 30

 320N03W 35

Vol. 8, No. 10

 (5〉く5m.)

 320N35W 30

 320N60W 40

 450N40W 10

 1st tree stratum       ,

Pinus densiμora

 2nd tree & shrub strata

Lespedeza buergeタi   ・

Smtiax china

Cαかinus laxifloダ(z         ゛

Q14がcus senaia

Rhododendダ∂S耀びrichii

Fダaximis sieboldiana

Cタclobalanopsis glauca    ・≒.

Eurya iaOonica         -.・

Vibuダnum dμataium

Zabelia integrげotia

Euonfmus alatus v. rotundatus

Sapium japonicum

Clethra borbiれervis

Pertva scandeれs

Perlya giabresccns

Buxus micずol)hvilaV.知夕onica

Diplomofpha sikokiana

Li?tdびa sfittcea

Soずbus gダaciiis

Thujotisis dolabraia      ,

Platvcarvc

Cvcl obalano i)sis salicinav.

               仙附θμび心,

Osmanihus ilictfoUus

Berberis thunber&ii

Tritomodon c6yれuus v. ダubens

spiraea hayaiae     ・,I

 ・ ・ ●  ●               ●Pieris iaponica

Illidum�igiosu肖

£知㎡αぷiptica

Akebia tyげoliaia

Deutzta 'icabra

ぴiburmtmびosum

Cocculus trtlobus

Prunus iamasakura

Anodendron aがine

Rosa onoei

Poufikiaea viUosav. zoUingeri

Loれicera cerasina

 Herb stratum

Carex subdita

Carex cilialo竹larZinata

Dtoscorea gfacillima

Goitum pogonanthum       ご

Brachyelytrum japonicum

Eccoiiopus coiulifer       ’

戸1がidium aQuilinum v. latiusculum

l^iola vioね2cea          '

Heterotropa sp・

Cymbidium i・iダgSCの!S ・

Dii)lot>teりgium glaucum

Miacaれthus siれensis

j4ダg心細ぷαゐiダ1α

ChionograCfiis ioi>onico

Astet aSeratoides ssp. ovaius

Melampyfum laxum

Cvclobalonol'sis acuta

R巳卵outfia japonica     .`

14 other species     `,

    (十)

   2C1)

   1

  、2

   1(十)

  ‘2(十)

   1(十)

' 1(十)

   ヽ3(2)

   2(十)

   1

、- :(十)

   1(1)

 ●  ●  ` 1(十)

    (十)

   十(十)

白 1(十)

   2(十)

 `  ●

   十(十)

  乙 ●   十

.' ●

  ,●

  :、 4

 ..,・5

  .+

    +

ト・  1

   '+

,. +

 ' ●

     2

     2

 ` : 1

     1

 i  ●

 {十}

2こ)

ドミ

2(十)

 (2)

 1

 1(十)

 2 2

 2

十+

3

 (1)

1(十)

2(5)

1

1(十)

 十

(4)

Flj

1ダ

(十)

]E.

11(十)31(十)11(十)

十 1 3(1) 3(十) 2(十)

,2(十) 1(1) (1)

 (十)

 ●.

 1(十)

 1

十(十)÷++

 1(十)

 1 1 1

 +  5 + +  2  2

 十

 十 十

Table 8. Pirws densiμoracommunity 6f Jinryd

  (280-290 m., S50W, 20-40° ; 5〉く5 m.)一一

 2nci tree stratum

Pinus densげlora

 Shrub stratum

Ffaxirms sieboXdia?1a

Zabeliainlegrげ心a

Rhododendfon toeWichii

びiburmtm dilaiaium

Buxus microtitoUoV. jationica

Les1>edeza buergeri

Z:^iplomorpha sikokiana

Abelia ■ierrata

μ外れip6れ4s rigida

Qitercusserrata

Plaiycarンaslrobirocea

Smiiax china

Ilex  Pedunculosa

spiraea hayatae

Cyciobatanotstsがauca

Rhododendron kaetnt)畑i

Euonymus alatus V. roiundatus

Rosa onoei

Pertyd scandens

Deutzia scabra

£yonia elli夕tica

Periya glabresceれs

Ilex integro

Ca。hellia ja知nica

Millettia ia夕onica

Coccuius triEobus

レ'ibiirnumがθsg。I

Sorbus gracilis

BfitchemiaracemosaV.  magna

乃面詰iaea vぷosa V. zoll加即ダi´

Eurya iaponica

 Herb stratum

Pseudofjogonaポh6れ4m quadfinerve

Lirio加州inoダ

Selagi)ぶla tornariscino

Carex ciliatomatk

Mtscanthtts sinensis

Paedびto scajwiensV. mairei

Orchis gramitafolia

Corex chりsolefis

Violaviolacea

HeiBfopappushisがdus

Solidago virgaurea ssp、asiatica

Hugeria iaponica v.  ciliai'is

Saussurea nipbomca ssp.yoshinagae

£μ沁加加加丿ticum V、必eanum

6 other species

2(2)

 り

 I

 Coa

oj.―I

 /'~\/~\

/-->

D十十 十

玖べ玖

μ

+ (十) 1

 3

 りり

刀十十

ぐぐぐ

十(十) 1

 1

 1

221十十十・1・・・・・・

5(1)

り りり りり

十D十十 十十

ぐくぐく ぐぐ

22112+11

十十

2(十)1

十(十)

十(十)●・・sし0・・..

211十十11・十・・・十十

十十

玖~113

2(十)  り りりり り

  十D十十十 十D

  ぐCCぐぐ Cく

21111+211+―

一参

十)

1)

1(1)

一●・・

・・●e

tNI―<r―(f―II―II+―+~●一・・

5(1)

 1

つJ

i-H

+1111

 2

 1

 2

 1

りり り  り

十十D十刀D十

十)

十)

十)

十)

十)

十)

1)

1)

十)1●

2

 1(十)

  (十)

  ● '

2111十・十・1十十・・・

一 一 一 一

5(十)

j   り  りりり りり

、十DD 十DD十十十 十十

くくく くぐぐくぐぐ十くぐ

くくく くぐぐCC

2222+1111

十(・

2(・

 2(十)

十(十)

 (十)

 ヽ(十)

 (十)

1(十)

 (D

+.1+

・・・一

2・―+11十十―・十十1・・・

   D. Tokushima District                               ●

 In Tokushima Prefecture, serpentine outcrops occur mainly in Jinry6・ Fukuhara, and

Sakashu. These serpentine areas are mostly situated in the warm-temperate forest region,

but the vegetation on serpentine is remarkably di琵erent from the neighbouring non-

serpentine areas as first noticed by Yoshinaga (1935, 1936).

 12. Jinryo (Yoshinaga 1935, 1936 ; Yamanaka 1952)

 In Jinry6, about 20 km. southwest of Tokushima city, serpentine crops ont aloug a

12 Research Reports of the Kochi University Vol. 8, No. 10

small stream, ranging from 200 to 300・ m. above sea-level.・The floristic composition

and structure of the vegetation in this serpentine area are di仔erent from place to place

owing to the conditions of occurrences of rocks and soils, and many trees and shrubs

form a closed vegetation. Five examples analysed of j the vegetation are shown in Table

7.        !      ‥

 The abundant occurrence oi Thujopsisdolabratais a characteristic feature (Plate v).

Moreover, it is also a noteworthy fact t]!atZabeliaintesriかlia and BuエMS niicrc巾心1ぬ

var・j呻o・nica.which are abund姐t on some limestone outcrops in Shikoku, are frequently

found in this serpentine area. On・ the other hand, stunted pine forests occur on steep

slopes consisting of firm rocks. Thかfloristic composition is shown in Table 8. The herb

stratum of this forest is comparatively sparse. Ldlium iaponicumvar.abeanum(=£.

abeanum)iS a serpentinophyte described from this serpentine area.

Table 9. Serpentine vegetation of Shingi Pass

     (840m.', N70E, 20°; 5×5 m.)

 2nd tree stratum

Tsuga sieきりIdii          ‘

Pinus d6竹sげI ora

Cりbtomeria iaponica

 Shrub stratum

SasamoreHa tatftwrascens

Pte^is iaponica、

Perlya giabrescetis

Rhus trichocafpa

Rhodo面姐Ton kaempferi

Siiifaea bluポ61

11ex crenata

Abeiia sertaia

Parabenzoin tri昂旨辨     I’ I

Smitax china            I

レ1

\^ibuダnum eダosum         、y

Clethダa bafbineダvis

Triiomndon cefnuuR v. れibens

OneずCMS seftata

7i!hododendron weyj・ichii

£luonymus alatus v. ダotundatus  j

Fyaχinus sieboはiaヽna

Stei>hanandra incisa

£タonia elliptica

Rosa onoei

Ilex Sgすすata            、-

Sorbus gtaciUs

Ma&nolia salicげotia

 Herb stratum      、   ・

Carex conica

Miscanifius sinensis

Ttipiefospermu?

Saussufea nipponica ssp・ yoshinagae

戸1が�turn aQiiilinμm V. latiusculum

StruがtioptがiSタtiponica・

召g祐がis thunbeダgit         、

Viola ovato-oblonga

Osmunda japonica

Geniiana sikokiana

7 other species

 (3)

2

゛り

j(l)

り りりりりり り

十D十十十十十D十

ぐくぐCぐCCくく ~-1●

~1  ・十十+2111

十(2)

2十,

り乙1り/」e

十・・十・・

りりり

n

CN}.―I

ぐくぐ

lcノij

OJ

/-N

/"N

/-^

/―S

 /'-^/■~\ト/'~\/~s

I

/s

I

^-s

。'~s

+11一21+―+     ―

くぐぐCくぐぐぐぐ 十   ぐ

22222211 卜く11121ト

             一―      一

 (十)1

●1・

22十―十・・・十

③出

/-N

/-S

/-N

^/―\

十刀D十DD十D十十D

ぐCCぐくぐCぐCCCls

22221121Q乙

1 (十)+

●●

2(十)

22十―十十―・・・

{十で}

3(1) りり  り jjjjり り

D十十DD十D十十十十十 十

ぐぐぐぐぐくCCぐぐぐぐ●ぐ11

2 31111 1+12CM

 ●

(十)

3十+2+1十十十十

㈲0 口

/・~\/^N/"^S/^S/・'^/~\/'~^/'~^/~\/""S/">.

/'~\/"~\/~\/"\

/""^

十十D十十D十十十十十十十 十十十十 十

ぐくぐぐくくくぐぐぐぐ

/Lχぐ Cぐぐぐ C ●●

4211121+11+11+112 +2+’

3十十2・1十・・十

 15. Shingi Pass(Yoshinaga 1935, 1・936 ; Yamahaka ・1952) ・

 The Shingi Pass, about 30 km. southwest of Tokushima city, is situated on the ridge

dividing the basin of the Katuura from that of the Naka. Serpentine outcrops occur from

850 to 1000 m. above sea-leve\. Spiraea blunxeiIS prominent on rocky outcrops, while

Tsugasieboはa and Pinusdensザloraaie rather abundant in stable places (Table 9).

  14. Ryu Pass (Yoshinaga 1935、 1936 ; Kitamura & Murata 1952 ; Yamanaka 19,58)

  About 2 km. southeast of the Shingi Pass、 serpentine crops out between the、altitudes of

500 and 1000 m. on the the northeastern side of the Ryu Pass. Almost the whole、area

consists of barren rocks and is characterized by the abundant occurrence ot spiraea

bluviei with stunted Pinws densiflora(Table 19). The Thujopsis dolabrata iorest is

also found near the ridge、showing the following composition :

 Tree stratum : 7加丿opsis dolαゐrata 5(1)

 Shrub stratum : Sasamoゆha  iJui’teti’ascens4、Si>i≫aea blumei10)、Patabenzotn trilobiもm1(十)、

Table 10. Thuiobsisdolabfaiacommunity of Sakashu

     (540-550 m., N, 40°; 10〉く10m.)

 2nd tree stratum

Thujoteis dolabraia

Cりpiomena jafionica

 Shrub stratum

PaYabenzoin tfilnhumLindera sびiceaV. sXabrataAbelia serrataClethra barbineniisV^tbuynum efosumRhus tnchocarpa

Rhodo海月dron uiびダichiiFtaxtnits s.ieholdianaCallicarpa TOoUis

Zantho%ylum tHixtiれ4mRosa onoeiRubus i>almatusStachyufus praeco%Magnolia salici/oliaHelunngia jatotdca v.imruifoUaHタ(irangea macrophylla ssp. serraiaEuonymus alatus V. rotundatusSmiiax sieboldiStVi'axiatonicaHydrangea paniculataSietManawira iれcisaS竹ailax china

Lonicei/a mochidzukiana£yonia elliptica

7 other species

 Herb stratum

Miscanthus sineタisisj?ubus wダightiiSaussurea nipbonica ssp. VoshinagoeAstがagがatoides ssp. amplex.げotiusCarex blet>hancari)a○がismenus undutatifolius v; japontcus

12 other species

 5(D {:4ミ}

 2

 3(1) 2(1) 2(2) 2(十) 2 1(十) 1(十) 1(十) 1(十)

  (十) 1  (十)

 1(1)

i゛●  (十) 2(十)

 1 1(十)

+ l(十)

  1

4CD C2} (1)

2(十)1(十)2(1)2(十)22(十)十(十)

 1(十) 1(十) (1) (十) (十)・ 1(十)

 (十) l

 (十)

 (十)十(十)

  3  +  2

  +  2

4(2) {31 {十:}

2(十)2(十)2(1)1(D

22(十)2(2)2(十)1(十)1(2)1(十)

 (1)2巾11

 (十)十(十).1

十(十)

 3/ +

 +

  1

  3(D

   {3}

  2{十T}

  2(十)

  3(2)

  1(1)

  2(十)

  1

  1(十)

  2(1)

  1(十)

  1(十)

  2(十)

   (1)

  1 、   (1)

  2(1)

  2

  1(1)

  1

  2

  + ●

   2

   +

  '+ ゛

・ ・ ●, 「

   ●

14 of the Kochi University Vol. 8, No. 10

Table 11. Ptnusdensifioracommunity of Sakashfl

   一(56p.m., S70W, 10°; 5χ5 m.)

 2nd tree stratum

Pinus densiμofa       ミChamaecypatis obtusd  -・  ・ ・Cりpiomeria iaboれica

 Shrub stratum

Viiurnum erosumPettya giobresceMsEnfeicnthus peれhiatusv、iationicusj?みododendダθタlweyダichii   ・.,,Sorbus gracilisAbclia sctratoSrnilox chinaSiJiraeablumeiFraxiれ14ssieboldianaClethダabarbineruisTfiinmodon cernuus v.れxbensUeχt>eduれcutosaBuxus miびo*hsUav.知加雨cα召gダ1?がislみμnbeダgiiRosa onoei         ヽPierisiatonica・S竹lilaxhigoe?tsis      バ  ・Diplomorpha siねokianaRhus trichocarpa£uonymus alatus v. roiundatusDifilomorliha trichototna      'Lordcefa mochidzukiana£yonia elliptica

 Herb stratum

Sasa tenuissirnaCarex spp.゛八■fiscaタithussinensisOsTrtundo iaponica?なダidiumaquilinum v. latiuscutumViola violacea      ''・Ilc% cf&nata

・ jl

 ‘2

 /1(1)

  +..

  `2(十)

  2(1)

  3(1)

  2(1)  1(十)

  .2(1)

,1(十)  1(1)

  2 ‘1(十)

  1(十)

 .1(十)  1(十)

  1(十)

 ''(十)

 j,●

  十(十)

   ,5 ・,.5

 ,‘ 2

  - ●  、+

ヽ・ ●

3(1)

詰)

2(十)1(十)川

幅)

2(十)

1じ)

1(十)

ヤ)

-1?

t

   2   1(1)

   3(1)

   3(十)   3(2)

   3(2)

   2(十)

   3(D  ・1(十)

r,  1(十)

   l(十)

   2(十)

   1(十)

   (十)

   1(十)

   (十)   1(十)

    5    5

    2

    1

    十

    十

131

4(十)3(2)

十(十) 2(十)

十(1) 3(十) 2(十) 1(十) l(十)

 1(1) 2(十) 2(1)

十(十)

 (十) 1(十)

十(十)

++

  5  5  2

  十

十{十万} 2(十)

 2(十) 3(2) 3(1) 1(十) 1(十)

十(十)十(十)十(1)

 1(十)

 2(1)

 1(十) 1(十)

十(十)

十十

 ・5  5  2  +

  +

゛aダ邸spp. include C. biethancarta andC. subdita.

  Euonymus oりμりμss十,Rhus irichocaゆa十, Cleihra barbinervis十

 Herb stratum : C�6x sifeoJ^iana 2,Sottssureo mppo?itca s即・ yoshinagae 1, Hydrangea macrophylla

  ssp. sβがata 1,召ε祐gダis lsC力onos々tana十, Rosa onoei十,Ruims Palmaius十, Ilexびenata十,

  Pertva glabtescB竹s+           .                           ’

 This stand is different fΓom that ・in Sakashfl by the abundant occurrence ofSasmnorpha

加ぽかliirascens 1n the shrub stratum.

 15, Sakashii (Yoshinaga 1935, 1936 ; Kitamura & Murata 1952 ; Yamanaka 1952)

          1           ?●.

 This serpentine area, about 2 k皿southwest of the Shingi Pass and the Ryu Pass,

lies between 400 and 700 m. abo‘ve sea-level. Though barrens are found in several parts,

almost the whole area is well covered with the characteristic vegetation, which contrasts

sharply with non-serpentine areas. Especially, the Tkwiot>sis dolabra£a  iorest(Table

10) and the Pinus densi flora lO rest with Enだianthus peridatusvar・ japonicus(Table

11) are very characteristic. Moreover, it is also of interest that Quercusがりlliraeoides

is frequently found in this serpentine area.

A Phytosociological Study of Serpentine'・Areas・in Shikoku, J [1]`rYAMANAKA)15

   E. Mt. Akaishi District     ……… ≒.・.     ‥≒,/

 Several serpentine areas appear near the eastern boundary of Ehime Prefecture, 50べ・60

km. east of Matsuyama city. The largest outcrop is found on Mt. Higashi-akaishi, and

small ones occur on Mt. Nishi-akaishi and Mt. Akahoshi. Moreover, serpentine is also

found along the valley of the D6zan. The surrounding rocks・ are chiefly crystalline schists

and amphibolite (cf. Horikoshi 1937).                    .

 16. Mt. Nishi-akaishi (Yamanaka 1958)

 This mountain is situated near Mt. Higashi-akaishi, just 2 km. west (higashi = east

and nishi=west). Though the highest part, rising 1626.4 m. above sea-level. consists

of crystalline schists, dunite and serpentinized dunite crop out well in the altitudes

between 1400 and 1500 m.. on the northwest-facing slope (PlateⅢ). This outcrop is

rather small and rocky, so that the coniferous forest is not found there.

 The rocky area is mainly dominated by deciduous shrubs, such as Berberis amurensis

vax. bre^ifolia, Rosa nipponensis, T^ripteりgiu-ni regelii, Kuon:ymus macroptet‘us・

Mag‘noliasieboldii. Spiraea sikokuaり'iina, 1)runus incisa vAT.  feinfeiensis. AbeLia

serratavai. buchvoaldii, Clethra barbinervis, Fra五丿1US lanuginosavar. ser7-どzzど2, and

Rhododendron spp. Noticeable herbs are Gynmocarpium jessoense.Farnassiafoliosa

vaで.れuininularia-, Aruncus dioicus var.tenuiかlius, Filipendulamul£ijuga, Angelica

ubaはkensis, P石tnula  farinosa ssp. modesta,Ruphrasia micropりlla, Leontopodium

japonicuni var. spathulatuni,Saussurea nipponicassp. sikokiana, Festuca O蛍れa,

Melica 7luはns. Careエduvaliana. Ale£risかliata,  etc. The vascular plants occurring

in this serpentine area Eunount t0 100 species, and they occur mostly on Mt.

Higashi-akaishi. On the other hand, the following plants, which are dominant or charac・

teristic on Mt. Higashi・akaishi, are not found, in this atea ・ Thuja standiihii,  Tsuga

diversifolia, Rkodode了idron me£ternichii,. Chamaepericりmenum canadense, Rumohra

■mutica,Sabina sargentii. Vacciniutnvitis-idaeavar.i7xinus, honicera mochidzukiana

val・ filifor mis. Scabiosa japonicavar.alpina, Adenophora triphyllavjr・・puellans.

etc.                                       ,.

 The physiognomy of the above-mentioned vegetation is rather similar to that on the

south-exposed slope of Mt. Higashi-akaishi.

 17. Mt. Higashi-akaishi (Yeimanaka 1950, 1952', 1955, 1958)

 Akaishi, meaning "red stone," is named for the yellowish brown colour of the surface

of weathered serpentine rocks. It has been well known that this, mountain。rising 1706.9

m. above sea-level, has a rich flora and conspicuous vegetation. The rock is dunitic。

          ●consisting essentially of olivine, and is quarried at the Akaishi Mine which is situated at

an altitude of about 1350 m. on the northern side. Chromiferous magnetite and banded

chromite occur as secondary minerals, and the magnesium content of the rock is rather

high, mostly reaching 45%Mg0.This ultrabasic rock area ranges from about 800 m.

upwards on the northern side, but it.appears from about 1000 m. on the southern side.

The main part of this area is situated within the temperate forest region chiefly domi-

・16 ・Research・Reports of the .K6chi University ≒Vり1. 8, NO: 10

nated by broad-lea・ved trees,' whereas the coniferous forest occurs in serpentine and set-

pentinized rock areas as.- a' topograph沁and edaphic climax. I

 The coniferous forest, which is .fairly well developed on the northern side, is dominated

by Chamaecがiaris obtusai Thuja 3tandish.it, Pinus penta].)hyllavar. hiniekomatsu, and

Table 12.‘ Herbaceous community of Mt. Higashi-akaishi

       (1650・m., S20W, 15°; 1〉く1m.)

Scabjosaiatonico v. 4伽声z.、。

Leonioねodium JoftonicuTav. stmihulatum

Carex spp. *

Veratrum maackii

Thalicttum minus

AnRelica ubatakensis

びtola bi flora           ・、一

Pariiassia foiiosaV. numniulaダia

Euf)Jtrasia microtihyila 、

AUttis foliala    ”  一一 ・ヽ

Rhododendron tschonosfeii

Geniiana scahfav. bueずgeri

Hosta sp、       犬

Adenophora tダifJtylla v・ μ・ぶαダis '

spiraea sikokualがu ・.

Abfiliaserrata v. 咄咄waldfi

Bcrberis amutensis vごbret・ifolia

7 other species

4432111十+12十十十・・・

4431十十―十+1十十十十・・・

4331十+1十十―十・・・十・・

242122十十11・+・・+・1

4311十十十―+1十・・・1・・

en CN eg f-H

.―t十+1111+11s十e

433111十十十十・十・・十十+

43321十―十+1十・十十・・・

5331+11+1十十十―・●●・

5331十―――+・十十・十・十・

* Carex spp. include C. blepharicatpa, and C. chrysolejris, andC. alterniflorav.aureobrunnea.

Alt。itude(m.)ExpositionSteepness

Carer, blepharicarpa

Miscanthws oligoslachsus       ,・

Brachypodium sylvaticum V. intsがμ恨

Hyijertcum sihokumonianum-

Geramum shikokianum  ・,     `

Sasamorbha turpurascens

Majanthemum dilatatum v・■nipponicum

Valerianasantbucげoiia v. fauriei.

Astil加話unbergii v. stfeofeuntontanaヽ

F山加ndula multijuga

Patnossia foliosaV.  nummiilafia

Leontopodium jabonicum v. staihulatum

Angelica ubatakensis

Rりnoutrio iatonica

Festuca ovinaI

MiscanffiMssinensis

Aster agei'atoidesssp. amt>!exi/otius,

Lepiosramma totta

Veronicastntm sibiticumV..ausirale `グ

Gentiana scahraV. buergBti   ・.

Sinraea iationica

Rttbus ttalmatus         几

Abelia seffataV.  huchwaldii

HeracUutrtlanaium ssp. moellendor.がit

              v.  tsurμgisaneれSむ

Ad。lotikofa temoiiflota   。・  ∧

戸∂1のitilla dicki刀■sit        j ’

Tithymalus sieboldianus v■ montanus ’一

PedicuiaYis resupinata

Carex fernaldiana

Deutzia crenataV.  nakaiana    ‥

1470

S80E

 20

●1

すle一

l. rttn^ .

.

.+つ」一十●e●

SS・●・1●

2 2

2十

 3

十esel+145十・―・十―・・十一一 ・一●●1●・

十e●eel十35・・2・・―・十十・十 ●●・●●・・

―・・・十+124十・・・十―十十・・十 ・・・●・1●

Table 13. Grassland of Mt.

十奉s一一+345十・1・十―・十十・十

一・・・・

 1480

S80E

 20

―+2+・・5・十――・2十1・十・十十十・+

112十・・5・十――・1・1・十・十十十・+

1131十・5・121・十・十・十・十十十・+

3141十・5・・1+・十・十・十・十十・・+

2141十・5・+1十・十・・・十・十十十・十

・十十・十・・

・・十・十・・

・十・・十・・

・・十・十・・

・十・・十・L

1475

S40E

 15

3+131十・十13212・2・十・1十十・・

2223・・・1123十―・+・+・1十十・・

3212十十・-12111・1・+・―十十・・

CN+1(/`ls・111111els+゜つ`++le

3・21・+・十111・十・1・+・十十十・・

十・・十・・十

・十十十・・・

十十・・・・.ヽ

・+・・・・.

・・十十・・十

Area!・in Shikoku, Japan(T。YAMANAKA)1・7

Tsugαspp. (Table 17 and Plate Ⅲ)。’On・the other hand, the community dominated by

     タ  ●   I    ・        I               I          。    4 1●ゝspiraea sikoku�かina

"is found on the southern' side where the forest is not developed

on account of rocky topography, greater insolation, and higher wind velocities (T゛able

20 and Plate Ⅲ).ln open places covered with soils containing considerable gravels, many

herbs・grasses・and sed呂es occur. An example of analysis in such a place presented

in Table 12 shows that the major constituents of the vegetation are common to those

of scrubs oi Spiraea sikokualpina(cf. Table 20). Secondary grasslands occur in the

most eastern part of this mountain. In this place, serpentine forms a large steep scree

on the northern side (Plate IV), while the southern side covered with rather deep soils

is rich in herbaceous plants as shown in Table 13.

 Thus, in dense forests where the ground is usually covered with the humus layer, the

characteristic flora of serpentine is usually not found. In rocky and gravelly places。how-

ever, are found many relics, endemics, and disjunctive elements. The plants・ described

from this serpentine mountain are Euphrasia  inicroplりlla, Astilbe £hunberがi var.

sikokunioヽn£ana(=A. sikokumontana), Geum japonicuni vax. lyoanuin (=G.りioanu‘m\

spiraea sikokualpina, Adenophoratriがりlla var・ puellaΓis(.―A. puellaris^, honicera

mochidzukiana var.aiiformis‘,and Paris tetraphyll‘a vaT. penduliβora.   '   :■

 18. Mt. Akahoshi「Yamanaka」958)

 Mt. Akahoshi, which is 1453.6 m. above sea-level. is situated about 8 km. northeast

Higashi-akaishi (1〉くlm)

1475SlOE

 10

1480 E

  5

1485S7SE

 20

1550Slow

 30

1475S7SE

 20

 2 4 4 3’3

 2 2 3 1 2

 1 1 1 3 2 2 2 4 2 4

 1 1 1 1 1

・ 1++・

● ● ● ● ●+ 1 1+ 1

 2 3 1 1十

● ● ● ● ● 2 2・ 3 3

・ + 1 3+

 3 2 1十十

● ● ● ● ●++ 11 2

● ● ● ● ● 1 1 1 1+

● + 1 ・ ●十 1 1 1+

+ 1 ・ + 1

● ● 十十・+

● ● ● ● ●

● ● ● ● ●

● ● ● ● ●・ ・ 十十十

 1+ 1十十

 ・ ・ 十十・

 ● ● ● ● ●

 ● ● ● ● ●

 ● ● ● ● ●

  2 3 3 2 3

  5 5 5 5 5

  3 2 2 1 1

  1 1 1 ● +

  ● ● ● ● ● + 1 1++

  2 2 1 2 1

  1 1 2 2 1

  ● ● 十十● 十十・ 1・

  ● ● ● ● ●

  ● ● ● ● ● + ● ● ● ● 十 l ● ● +

  ● ● ● ● ●

  ● ● ● ● ●

’● ● ● ● ●  ● ● ● 十 1

  ● ● ● ● ● 十・十十・

  ・十十十・ ‘1++++

‘+十十十 1

 十十十十1

 + ● ● ● ●

  ● ● ● ● ●,‘ + ● ● ●

  ● ● ● ● ●

  ● ● ● ● ●  ● ● ● ● ●

   2 3 3 3 3

   5 5 5 5 5

   1 1 1-ト 2

  十十十 1・

   4 3 3 3 4

  + ● ● ● +

   1 1 1 1 1

   1 1+ 1 1

  十十十十十

   ● ● ● ● ●

   ● ● ● ● ●

   ● ● ● ● ●

   ● ● ● ● ●  十十・+・

   ● ゛● ● ● ●

   ● ● ●・● ●  ++++・

  + 1+十十

   ● ● ● ● ●

   ● 十 ● ● ●

・  ● ● ● ● ●

  + ● ● ● ●

   ● ● ● ● ●

   1+ 1++

   ● ● ● ● ●

   ● ● ● ● ●  +++++・

   ・++ 1十

   ● ● ● ● ● ゛

   ● ● ● ● ●

   4 5 4+ ●

   3++ ● ●

   1 1 1十 1

   ● ● ● ● ●    1 2 3 3 2

   ● ● ● 3 2

   ● ● ● ● ●    1 1 1 1 2

   ++ 1十十

    1+ ]十 2

   ● ● ● ● ●

   ● ● ● ● ●

   ● ● ● ● ●   十十 1+ 1

   ● ● ● ● ●    5 5 5 5 5

   ● ● + ● ●

   ● ● ● ● ●

   + l ● ● ●

   ● ● ● ● 十

   ● ●’● 十十

   十十十 1・

   ● ● ● ● +

   ● ● ● ● ●

 F  ● ● ● ● 十ヽ

   ● ● ● 1 ●

   ● ● ● ● ●’

   ● ● ● ● ●

   ● ● + 4 3

・  ● ● ●・+ ●

   ++● ● ●

   ● ● ●、● ●

   ● ● ● 十 ●

   ● ● ● ● ●

   ● ● ● ● ●  ‘ 5 5 5 5 5

   ● ● ● ● ●

   ● ● ● ● ●

   ● ● ● ● ’●

   ● ● ● ● ●

   ● ● ● ● ●

   ● ● ● ● ●

   ● ● ● ● ●   2 2 1 3 4

   ● ● ● ● ●

・ ● ●・.● ● ●‘

   ● ● ● ● ●   十十十’1・

   ● ● ● ● ●

   ● ● ● ● ●

   1, ● ● ●・’●

   ● 2● + 1

   ● ● .●・ ● ●

   ● ● ●● ● ●

   千 ● + ●´●

   ● ● ● ● ●

   ● ● ● ● ●

   ● ● ● ● ●

   ●● ● ● 1 2

18 of the K6chi University Vol. 8, No. 10

of Mt. Higashi-akaishi.八small serpentine outcrop appears in altitudes between 1050 an(!

1150 m, 0n the northern side of this, mountain. This ・area is characterized by the com・

munity dominated byspiraea ・bitμnet.The floristic composition of it is presented in

Table 14.          し   I ●・ フ

           Table 1心 spiraea bli。metcommunity of Mt. Akahoshi

                 (5χ5 m.)

Altitude(m.)ExpositionSteepness

`;60

n00S15W 50

n00S25E 60

n00

S15W

 50

 Shrub stratum

S{>itoeablumei

Quercus senataLespedeza buergeriAbelia Sびrata

Cartiinus corfrinoides   1EuonVmus alatus v. toiuれ/iatus

Rhododendron kaempμriR、decaれdれ4mRosa onoeiZanthoりh4mがperitumRhododendron perdaphタHumF穴

 ?

 十

 t

 ●゛1

・:ノ

 3

 2

・’1

’+

 1

 1

 十

 十

 十

 十

 、+

 Herb stratum

Carex dt4ひalianaC、alterniμoraV. aureobruriKea

            et C. sikokiana

C/Zダysant細川um iタtdicurn

Mtscanthus sinensis

Thalidrum miれ14SCafex ciliaiomareinata

Aster ageratotdes ssp、amOiexi/oUus

Galiu脚力inuta

Brachytodium sjloaticurnv.

                 竹itserumPotentμla dickiタ・ISa

Brochタelvtrum j叫ionicum

Selaeinella tamariscina

2111++

 l

 l++

 ●

 2

 2

 1 1

十 l

 ●

 ●

 ●

 ●+

 In addition, it is a noticeable fact that such calcicolous ferns as Camp£osorus-sibii-tC・us

and Poりstichum eras pedosorun・IOC(沁r‘frequently in serpentine crevices of this area.

 19. Tomisato (Yamanaka 1953)

 As mentioned before・ serpentine outcrops occur in areas along the valley of the D6zan.

The outcrops appear most!y on a rather small scale, being situated in altitudes between

300 and 500m. The areas are,composed of firm rocks often overlaid by weathered materials.

There are several precipices of moderate height along the valley. The rock which is

rich in talc has been quarried in some places.

 The climax vegetation of this region seems to be the forest composed of Tsuga sieboldii

and C:yclobalanopsisspp., which represents the upper part of the warm-temper ate forest

region. Accordingly, Cydobalanopsisがauca, Machilus japonica. Camellia jOpO・・lica,

etc. are not rare on some small serpentine outcrops. However, forests dominated by

Pinus densiμora ate also found i・n this・ region. The forest found at an altitude of 320m.

shows the following floristic ・composition :

 1st & 2nd tree strataご Pinits densiμOTa '4, Tsuga sit・boldii 4 (1), Chamaecyparis obtusa3(1),

  CりMomerio iot>onico・2(1)・

Shrub stratum : Pteris・加加nica 3 (1) ,£yonia elliptica 3 (1) ,Rhododendion decand'Tum3(十),    I   ●●       。・      ●  「  Cyclohalanopsisglauca7(!),μa加dunculosa 1(1),Rhus trichocariya 1(十),aりera j(ゆonica

  1(十)・£“ぴ“加加戒9 1・ C“”1゛ぶ“加加nicα1, Ctethダa barbinervis l, Sorfausiaponica十,VacciれiU竹t

  smallii var. が必ダZ4四十(十), Vaccinium hiftu。十                ‘

 Herb stratum : Hugefiaiatonica var. citiorts 3, Hym㎝(ゆhyllum barbatum 2,Diplopieりgium

         ■       l- ●    |  jμaucum 1, Aダdisiα加加雨c4 !,£・spisoなs thunbeなiα。s十,Miscanthus sinensis十, Castanea

  C旬nata十, Rkododewdton kasmpfeti十,Paederiascandensvar. mairei十,Smilax china十

  This forest is not・l stuntedに10~15 m. high, and is accompanied bv ericaceous plants

in abundance. With a small difference inヽthe floristic composition, similar forests' are

found in other places within this region. For example, a rather open pine forest ・at an

altitude of 350 m. is constituted by such plants ・ as Lespedeza buergeri (abund'ant).

               ・            ・                                r゛Rhododendron decandriLtn (frequent), Cy clobalanopsis elauca, Neoli£sea sericea.

 .ゝ             ●      ・              I.・              ●Cinnamomum japonicum, L)eutzia scabra, Pieris japonica, Fraエinus sieboldiana.

etc・ , and the following species occur in the gravelly places or crevices "■Miscanthus

sineれSIS(abundant), Carにduvaliana(abundant), Po・りstichum craspedosorum (;locally

abundant), Sela^inella tamariscina(locally abundant), Oplismenus undulatifoliusvar.

iaponicus(locally abundant), Coptidipteris ■wilfordii(frequent), and Ardisia japonicd

(frequent).

 Noticeable plants, occurring in such serpentine areas are Coptidipteris ■wilfordii,

Caniptosorus sibiricus-, Polystichum c:raspedosoruTn, Cyr・tomium falcatum. Clematis

tosaensis, Berberis thunbergii, Arabis serratdvar. sikokiana, Buエus microphがlavar.

iaponica, and Carei: duでaliana. Some of these are considered to be calcicolous plants.

Especially, it is very characteristic that C・1や£osorus sibiriciら Poりstichum craspedo-

SOrしini,  and Buエus viicrophyllavar・ Japonic a are frequently found here (Plata Vn). They

are abundantly or frequently found on some limestone outcrops in southern Japan. The

floristic composition of the community characterized by the abundant occurrence of Biにus

microplりllaval・ japonica vs as follows :

 2nd tree stratum : Cvclobalanoteis Etauca3(十), Clethダa bar・binがz心3,  Tortりa nuct/ero 2(十),

  Cephaiotaxus harringtonio 2(十), Actinodaphne lancifolia 2, AcerタフzaタzθvaT. mafmmatum i.

  disscclum 2,Zeikova serrata 1,Prunus iamasakufa 1,Satium japonicum 1, Maackiaμori-

  bunda十, Acer palmatum十,Meliosma myrianiha十

 Shrub stratum : Buxus mic・yo油少(avar.加加nica 3(1),  Abeiia spathulatavat. subtetrasepala 2(十),

  £^euizia scabダα2,£g田田aphyllum miびophyllu四十(十),NeoUlsea sericea十(十),Cinnamomum

  japonicum十(十),  Kertia japonica十, Euonymus alolus var. rotuれ^atus十(十),Paederia scandens

  var. mairei十(十),Smiiax china十(十),CoccmIms ttilobus十

 Herb stratum:Cび6 conica S, Selegineltdiamarisdna 1, Oplismenus undulatφjtiMS var. .iaponicus

  1, Caダex reinii 1, Gonocoダmus minutus十, Lepisotus thunbereiames十,PIびis cretica十> Aster

  agerotoides ssv). omtiexifoiius十,  Dioscorea iokoro十, TrachelosPertnum csiaficum+

   F. Mt. Shiraga District

 Mt. Shiraga, rising 1469. 6m. above sea-level, is situated near the border-line between

K6chi and Ehime Prefectures, about 30 km. north of Kochi city. Serpentine crops out

in altiludes of about 1000 m. and upwards. The surrounding rocks are crystalline schists.

 20. Mt. Shiraga (K6chi Reg. For. Office 1939 ; Yamanaka 1950, 1952, 1954)

 This mountain is known for the eχcellent forest dominated by Chainaecy夕arls ob£lisa.

Climatically, this area belongs to the cool・temperate forest region mainly consisting of

theFagus crenata climax for‘est. However, the Chamaeり夕a石s obはsa forest accom-

panied by Pinus pentaphyllavaT. himekomatsu, Tsuea sieboldii. andTsuga diversiか1祐

is found in the serpentine area as an edaphic climax (Plate v).The floristic・com・

position and structure of this forest are presented in Table 18. The typicai stand of

  .   ●●                  ●  1  1        ・this forest is found on the northern side which is rather flat and moist. On the other

20Research Reports of the・K6chi University Vol. 8, N0.10

hnd

・the south-facing・ steep

。slope.near

' the summit is characterizedby

thedwarf

vege-

tation on account of its topography and dry microclimate(Plate v).lnsuch

a place,

巾efollowing floristic

composition 沁found:

 Shrub stratum ご' ChamaeりParis o!}tusa 4。Tsufia diversげolia 2(十), Clethf・α1。z,・binefvis十(1)

 Herb stratum :・Cay ex aliernifiora

。var.

aurcobrun,nea4,spiraea

sikofeualfHna3,Rhododendずo?1

 ’加?liaphyllum 3, Ilex pedunculosa 2。Fりizi?tus la?iiiRi?tosavar. seffata 2, Vaccinium smaμIf var.

  glabru。z 1, Viola shihokiana 1, Mela。1勿ダμ加ね2XU。2 1, j?ubus palmatus十,Euonyタ92μs隋αびθ-

 , Hefiis十,Rhus trichocarpぶ十,Rhus ambiRua十, Miscanthus sinensis十

 Moreover, exposed rqcky places are characterized by the occurrences of such shrubs

asspiraea sihohu以加na.・Abeliaヽserrata vat. buchzvaldii, etc.An example is as

                    F   II

follows:

 SPifaea sikokualpina 3, Abe111a sefifata var. buchwalぷi 3, Carex alteダniflora var. Qureobrunneo3。μgズ加dunculosa

2, }?hbdodの�ダon pentaP.・'hyllum 2,戸θ砲心iliaぷckinsii 1, Rubus戸ぷmaius十,

 Fダaxinus lanuginosa vat. setfata十,

Melampyダum laxutn十,Saussufea■nipponica ssp,sifeofeiona十

 The forest vegetation of this mountain is similar to that of Mt. Higashi・akaishi,but

the flora is not so characteristic. This・ problem will be discussed later onパ

III. Serpentine vegetation in Shikoku

 In this vegetational study, the quadrat method was adopted, and the belt transect was

also used in case of necessity. Many quadrats varying in size, 10×10m., 5×5m., 2×2m.,

むldいく1 m., were laid dqwn respectively in forests, scrubs, and grasslands. The

object of these quadrats was t9 obtain data with regard to the floristic composition,

dominance, constancy, and fidelity of the component species of plant communities. The

dominance was essentially interpreted as described by 。Braun-Blanqu吋(1951). The

associatiと)nwas determined ・and named from dominant and characteristic species of the

communities. As the total value of every species in one association, the ten class con-

stancy and the coverage value were used。

          ・   ・  ゝ A. Plant communities on serpentine in Shikoku

 Floristic compositions and physiognomies of plant communities in serpentine areas

are usually different from those of neighbouring non-serpentine areas. But in some places,

the characteristc feature is not observed. As already reported in previous papers of the

writer (1957, 1958, 1959), several distinct associations were recognized in serpentine

areas of Shikoku. Besides, there.・are some stands that cannot be included in any of

these .associations. When enough data are not obtained, names ・are distinguished as

provisional.

 1. Pittosporeto-Quercetum phylliraeoidetis, Suzuki et Hatiya (1951)。(Qtierciもs

  j,岫以行aeoides-Pi££osporuin tobira,Association)(Table 1)

 In Japan, the Qtiercus坦りlliraeoidescommunityis found mainly in the coastal district

within the area of the Shiia siebolぶj Alliance, and is also well developed in Shikoku.

 Physiognomically, the Quercusかhylliraeoidescommunity can be divided into two types.

namely, the Maquis-type and .the Garigue-type, and the floristic compositions of these

two types of the community are more or !ess different from each other. Notwithstanding

nJA・reas/inShikoku、Japan (T. Yamanaka)、2j

such・physiognomical・and<floristic

differences; these two types・of the community should

be essentially ・included in one and the same aS:sociation.  ・ ・       ,.  ・ ‥

 This associataon is regarded as an edaphic climax in the warm-temper ate forest・ region.

The physiognomical characters of this association resemble those of sclerophyllous forests

or scrubs which are found in the Mediterranean Basin and California. This association.

however, is quite different from those forests and scrubs, because the climate of districts

where t叫s association occurs is usually characterized by high precipitation and summer

             F   ●がrain (Sato 1946 ; Hatusima 1948 ; Suzuki !953, 1954 ; Yamanaka 1958).

 In serpentine areas of Shikoku, it is of interest that Quercus夕りlliraeoidesoften occurs

in inlands。 Such an example is found in Sakashu and its vicinity, Tokushima Prefecture

(cf. p. 14). However, a typical community on serpent・ine appears only in the coastal

゛9 0fthe Yawatahama district (cf. p. 2), and is often accompanied by Pinus thunbergii

in 'the upper stratum.

 t   lll  t    l    s   ●   ●                               1  ●

 2. Abelieto・Pinetum densiflorae, Yamanaka (1959). (Pinus densiflord・八helia

   serrata Association) (Tables 15 and 16)

 The Pinusdensifloraforest is widely distributed throughout Honshu, Shikoku, Kyushu,

         '  へ   1      1   ゛                      ●  'and adjacent regions, and phytogeographical and sociological studies were made by a

number of authors, such as Hayashi (1952), Yoshioka (1948, 1949, 1958)。Suzuki (1953,

1954), Usui (1953, 1954), Kitagawa (1954), and the writer (1957, 1959). The ・natural

development of this forest is restricted to stands of special edaphic conditions, such as

 f    d   ●                     ¶   ●                ●ridges, rocky outcrops, screes, volcanic ejecta, and alluvial sands (cf. Yoshioka 1949).

0n the other hand, secondary forests dominated by Pinusldensiμoraoccupy extensive

areas ranging from the warm-tempとrate to the cool・temperate forest regions.

 Pi7ias densiがoracan grow on various kinds of rocks and soils because of its particular

                                                       ●characters requiring little mineral nutrient and soil moisture. Accordingly, it occurs

also in serpentine areas where the climatic climax forest is not found.

 Serpentine areas・ in 10w altitudes of Shikoku are mostly characterized by open and

stunted stands oi Pinusdensiがoraaccompanied by a large number of deciduous shrubs,

herbs, and grasses. Floristic compositions of such pine forests are remarkably di仔erent

from those of the pine forests on other rocks. Emphasizing this floristic characteristic.

the pine forest on serpentine is treated as a distinct associat・ion, the Abeleieto-Pineはm

densiβorae.

 In the・serpentine areas of the K6chi district, this association is well observed. In

the Tokushima district, a typical stand of this association occurs in Sakashu (Table 11).

The pine, forest in the Yusuhara district is also essentially not different from that in the

K6chi district. From t・he above,・ it is concluded that serpentine areas in Shikoku, situated

in the warm-temperate forest region, is represented by this association.

 ろ. Rhodoreto-Chamaecyparidetum obtusae, Yamanaka (1957). (Chan!aeりpar・IS

   ohtusa-Hhododendron皿etternichii Association) (Tables 17 and 18)

 C hainae cyparis obはsa .IS distributed in HonshO, Shikoku, and Kyushu, and its natural

forests are well observed in middle Honshu,・■ the Kii Peninsula, and Shikoku.

 As to the distribution and ecology of the Chamaec:yparis‘ohtusa forest, studies were

22 Research of the K6chi Universi Vol. 8, No: 10

Table18; Associationtableof Rhodoreto-Chamaecypandetum

AltitudeCm.)Exposition    ご .Steepness

 ダ     1200       N40W‘ ●     30

1400

 0

 1st tree stratum

Chamaecypris obiusa

Ts昭a dit・eysげolia

Pinus l)e?italitollaV.・himekoniaisuTsuea sieboはii

 2nd tree & shrub strata

j?hododendt・∂が阿gμernichii v.

             hotvloeれS6 .、Sasamorpha purpurasceれs

Suonstnus maampHrus ・

Fれ v. serrata こ

Parabenzoin iril必g阿

Tritomodon cernuus v. rubens ・Acer micfaボhum 。  ’  、I

Clethダa baft?1・ervis

Tfochodendダ071 afalioides  ・ ,

Acer sieboldiartum

Rhododendn削pentaphyllum..Acanthopanas sdadophタItoides・ ,

Sorbus comtnixta

Rhtts ambigua

Tripteぴがum regelii      l

Rhus trichocarpa

内知y四阿gだeolatum   ・,Ilex Pedunculosa   ,

Rubus Palmatus

Iiex maぴ0μ)da        、¨

S other species

 Herb stratum

Carex reinti

Humohta muiica

IlexれxSosaV. stenoがlylla

Skim肖ia japonica v.intermedia

             ・i. reper,!S

Lタcopodium senatumV. thunbが■gii-Strutfiio{)tがisrapomca  、、.

C�ex moi^oiuii"Plaiantkera of^ydioides,

EphippianihMs schmidtii

5 other species

5(十)

 {十万}

・5゛

1

y

)

 :尚

 ・:

 ●゛

/(十)

ご:

jり J

/3'

 ご

・,.+

・ ● 」

^li:

j5

・(十)

 (十)

 (十)

 1(十)

 (十)

  5

 十

゛2

 十

1 1

 、●

 ヤ)

  ●+.

  ● 雨

 1.

  ●  ●  ●  ●  ●  ●  ●  ●

  ● (十)

  5

  ●  2

  j

 +

  ●  ●  ●

5.

十(十)

5.

レ)

 雨

1.

十(十)

 (十)

 j

5(1)

1(1)

1 (1)

1

1

1

十(1)

十(1)

  十  1

  3

  4

  +

  1

 5

 1(2)

 2

  ●

 5(1)、

  ● 2(十)

  ●

  ●

  ●

  ●

  ●

  ●

  ●

  ●

  ●  (1)

  ●

  ●

  ●

  ●

  ●

  ●

  5

  3

  2

  1

  ●

  ●

  ●

  ●

  ●

5

 (2)

2

 ●

5(1)

 ●1(十)

 (十)

 ●

 ●

 ●

 ●

 ● ●

 ●

 ●

 ● (1)

 ●

 ●

 ● (1)

 (十)

 ●

  5

  3

  1

  2

 ● ●

 ●

 ●

 ●

ヅ)

ヅ)

ド)

?)

作七

詰)

2.

5(1)

j(o

 ● ● ● ● ● ● ● ●

 ● ?)

 ● (十)帽

)

 ●

 5 ?

 j

 ● ● ● ●

5ツ

2.

5(1)

1(十)十

゜.

い)

(.1)

汐)

made by Kawada (1930)。Nakano (1942), Hayashi (1951), Maeda (1951), Yamanaka

(1957), etc. Kawada e又p!ainedレits distribution by precipitation throughout the year.

Maeda pointed out・that montlily rainfall in winter indicating about 100 mm. orn!ore was

suitable to the occurrence of 由is forest. At the same time, he also stated that the

podosolization of soil was an index to 出iS forest and was promoted not only by climate

but also by topographic conditions and the character of rocks. The writer concluded that

the Chamaeり1)a元soわぬsaforest .in Shikoku was found as an edaphic climax within                 ・the area from the upper part o卜the warm-temper ate forest region to the cool・temperate

forest region. In Shikqku, ・ this forest occurs usually on a rather small scale on ridges,

rocky outcrops, steep slopes'.,etc・,while it occupies extensive serpentine areas in the Fagus

crenata climax region. Suとh C hamaeりParis-typeforests occurring not only on serpen-

tine but also on various kinds of rocks show floristic compositions fairly different from

obtusaeon Mt. S!liraga1・

         1450

‘     N60E      ,          30

1450-

 0Coveragevalue Constancy

詰)

ヤ)

1(十)

ケ)

?、 じ)

1(4)

 j

3ツ

サ)

+(十)

1.

+.

 (?)

+.

 j

 1 臨

 ●

ヤ)

+.

 ●十 ● ●

 ●ツ)

 ● ● ● ● ●

 1

  3

 ● ● ● ●

 1

 2(1)

 3

  ●

4(十)

  ●+

  ●

  ● 1

  ●

  ●

  ●

  ●  (十)

  ●

  ●  (3)

  ●

  ●

  ●

  ●

  ●

  ●

  +

  3

  2

  +

   1

  ●

  ●

  ●

  ●

 (十)

1(D

3

5(十.)

1 `

 (3)

  3

  2

  2

  争

  2

  ● 5ツ

  ●

 ‰)

 7

  ●

  ● ソ)

  ● ソ)

  ●

  ●句)

  ●  ●  ●  ●  ●

  1

  5

 ふ

  ●  ●  ●  ●

T)

十(十)ン

1:

十(D六)

 ヤ

 1

 3(十)

 1.

 阿)

づお

 1.

  ●

(十)

  ●

  ●

  ●よ)

  ●

  ●

  ●

  ●

  ●

  ●

  1

  2

  ●

  ●

  ●

  ●

  ●

  ●

 1(D

 5(十)

 2

  ●

 5(D

  ●

  ●  (干)

  ●

  ●  (十)

 1(十)

  ●

  ●

  ●

  ●

  ●十(1)

  ●

  ●

  ●

  ●

  ●

  ●

  5

  3

  2

  ●

  ●

  十

  ●

  ●

  ●

2.

句)

 じ)

ヤ)

言)

 1

   4587. 5( 78.5)   2480 ( 652 )   1513 ( 1.5)    25. 5( 51.5)

   6087. 5( 228 )゛   1750 C 113 )    440( 3.5)

    153.5C 4 ) ご    100.5( 1 )    100    88( 2.5),    75( 1.5)    51( 0.5)    50. 5( l.S)    26. 5( 27 ?    26    25( 1 )     3 (1338 )     1.5( 1‘)・     1.5( 0.5)     1(・● 1 )     0.5(100)     0.5( 2 )ヽ     0.5( 1 )

   6064   2563   2412. 5   ・

    653    251.5    27.5、   25

     0.5     0.5

 虻

 環 Ⅶ Ⅲ

 K M一環

 Ⅸ‘

 Ⅲ I・ Ⅳ・` Ⅳ・

 Ⅲ

 I・ Ⅳ・ I I

’IX �・ I I .I ’

 ・l・ I

 士ぺ

 χ

 Ⅶ ’ Ⅳ

 狙 ! I I

one another. However, they can be grouped together into one association, the Riiodore£o-

Chaniaecyparideはmoわはsae.The forests on Mt. Shiraga and on Mt; Higashi-akaishi

a・re typica! stands of this association.

 4. Thujopsidetum, dolabiatae, nom. prov.  (Thui砂sis dolahraは Association,

  provisional name) (Table 10)

 Thuiopsisdolabraはismostly found 皿middle Honshu and occurs occasionally in

Shikoku and Kyflshu (cf. Hayashi 1952). In Shikoku, the forest dominated by this tree

appears in Tokushima Prefecture. Though the occurrence of this forest is not restricted

to serpentine, typical stands are found in serpentine areas (Yoshinaga 1935, 1936 ;

Miyazaki 1950 ; Yamanaka 1952, 1959).

 As already stated. this forest occupies extensive areas in Sakashu. Thiりopsisdolabrata

24 ■Researcfi of tHe K6chi University' Vol. 8,・NOン110

                       i.

・ Table 19. Association table ’of・IS誠心g,咄細叙umei in Tokushima district . i ≒          1 ● ●』       =

っ迄昌に。)\‥.臨皆言

  14 'I

,卵O、 S20W

  30

I '14680S35E

 35

 14930S05E

 25

 14940S55W

 40

13

850-

 0

Coverage

  value

S、§

l

ロ6

   Shrub stratum

S折はea falumei

Sasamorpha purpurascens

Pertya glabresce7ts,

Pirats densi/ioraAbelia s6がata

Sorbus gracilis

Cleihra barbinervis

S・milax china

Viburmtmぴosu*n

Rhus trichocafta      .・

Quefcus sen at aRhododendron kaempfefi

 ● ・ ●  ●Pierts japonicaHydYanma j)a竹1cM↓ata

Acer夕almatum      .

Rosa OKoei

戸aダabenzoin trilobum

CりAtomびto japonicaμgぶぴenata

j?hododetuiro夕l weyrichii

DRuizia crenataHydYangea macrobhylia

          ssp. sRirataAlnus. hirsuta v. sibirica

Tsuga sieboldii

召erberis thunbergii

Stephanandfa incisa

9 other species

   Herb stratum

Miscanlhus sinenstsSaussurea mppomca ssp・

          SoshtnagaeCoerx sikokiana

Carex cornea

Viola ovato-oblonga

Tripietospermum joijonicum

10 other species

  5(1)

  2(1) .

   ●  ●

   ●  ・

  1(十)

、 ●

  1

  1

  l

 +

   ●  I’

 +

肖  ●

、+

 + j・

   ●  ,

  ’2

   2‘

   1’

   +

   ●I

   7

  5(1)

    ●

.' ●

   (十)

    ●

    ●

    ●

    ●

r  ●

  +

  +

  +

、・  ●

    ●

    ●

    (十)

    (十)

  十

  +

   .●

    ●

   .●

    ●

    ●

    ●

    ●

   ‘1

    +

   ・・1

    ●

  ● ●

  ゛●

5(1)

3

1十十(十)

 1

 1

十(十)+

+十

  2

  1  2

  十  +

5(1)貼)

 ●

 ●21ブ

 1.

 ●

 ● ● ●

'トヤ)

  ●4ヤ)

  ●

  ●

  ●j:(申)

  ●

  2

  1

  ●

  ●  +

5(D

3(十)

lCl)

 (十)2(1)

十(十) (十)

 (十)

 (1)

十(十)

 (十)

十(十)

十(1)  (十う

  (十タ

十(十)

  2

  3

  2

  +

8750(500)

1200(104)

 850(200)

 750 ( 4)

 350(100)

 350

 202 C 2)

 104( 6)

 104 ( 4)

 102( 2)

 102

 102

 100(100)

 100

 100

  6( 6)

  4( 4)

  4( 2)

 `4( 2)

  4( 2)

  4

  2(100)

 ` 2( 2)

  2C 2)

  2( 2)

  2(2)

1500

1302

1300

 350

  6

  4

V’II

ⅢIl

lⅣ

狙ⅢII

IIIⅣ

ⅢIII

IIl

lI

ⅣI

ⅢI

  * cf. Fig. 1.           ’・’

dominates in the tree stratum ・and is often accompanied by such coniferous trees as

Cryp£otneria japonica. Chamaを'りParis obtusa, Sciadopiりs 。erticillata, Pinus

petiteゆhyllavar. himelこOma£SM, an・d Tsuffa sieboldii.Sometimes prostrate branches of

"Thuj opsis dolabrata completely cover the ground.

 The Thujopsis dolabrata forest occurs mainly in the cool-temperate forest region in

middle Honshu, whereas it appears in the warm-temper ate forest region in this district。

               d   l 。・ゝ゜This fact is of great・ interest from phytogeographical and ecological viewpoints. The

comprehensive nature of this forest is not yet established from the study in this district

alone, a provisional name,  iche Thujopsideはm dolahratae, \S presented here.

 5. Associations of Spiraea (Yamanaka 1958) (Tables 14, 19, and 20)

 The Spiraeetum blwinei, Yamanaka (spiraea hluniei Association), which occurs in

serpentine areas of Tokushima Prefecture, is characterized by the dominance oi Spiraea

blumei in the shrub stratum. In this district, Saussureanip夕onica ssp・ :yoshi?taeae\s

Table 20. Association table of SPita-cetum,sikokualtiinaeon・ Mt. Higashi-akaishi・

1 ●   ち    ●   ●g    Altitude (mう

    Exposition‘

    Steepness

1540

S25W

  5

1545

S25W

 25

155S

S30W

  5

F i●- 4●. 1555

S30W

  5

 ・ J 1560

S20W

 20

1565

S20W

 15

1635

Slow

  5

1635

Slow

 15

1665

S20W

 10

1670

S30W

 15

J゛今

`l

ー4・JI

Q

  Shrub stratum

S1>iraea sikokualpina

Abeμどz serrata v.        buchvualdii

Bびberts amurensis v.         1,ダevifolia

Rhododetuiron lagopus

Rosa niPiJonewsis

j?hododendron decandrμ四

Pafabenzoin trilobttm ゛Pinus Peniaphyllav.

       himekomaisu

cutみ?・a barbinervis

Rubus Palmatus   -

Rhododendron tschonoskii

Aceダsi必必jiαがz4四

Rhododendron細丿itatihyllu竹1

  Herb stratum

Care% blepharicarpa

Cαダ6spp.普

Miscanthus olygoslacり≪sScabtosa japonica ^'. atpina

Thalictrum minus

Mtsca?at hits sine竹sisFesluca ovina

Parnassia foliosav.

        琲4mボulaダia

Angelica ubatakensisLeoniopodium japonicum

      v、s1)athulaはずy1

Aster agぴatoides ssp.

       a恨Wexi/oUus

Satissureo rdpponica ssp.

         sikokiaがα

y1面河ophora triμポla V.          iniellaris

Viola biflora

Aruタ1CUSぶ∂iCμS V.         16,面面沁4S

Gsniiana scahraV.          buergeri

Hypericum sihokumontarm竹l£μ

j4Sμtbe thunbergii v.

     sikokumoniana

BrachnPodium sy1むattcum

        V. mtserum

Melica itatafKChionographis iaponica

Viola gりPoceras V. exilis

Reynoutria japonica

Aletris foliata

Potentiμa dickinsii

Hoslo sp・

Primula farinosa ssp.

          modestaド6Γαlyμm maacゐii

 5

 2

 十

 十

 3 2 1

 十 1 1 +

 1

 +

 1

 1

 十 1

 +

 +

 1

・十

 4

 2 2 1

 3 1

十十

 1 2

 1

 1

 1 1

 1

●、

 3

 2

● 1

 1

●+

 2

 1

 2

+ 1

 1

 ●

 ●

 ●

 1

 ●+

 ●+

 ●

 ●

 ●

 ● ●

  5

  1

 十

 十I +

  ? 十

  2

  4

  2

  1

  1

 十

 十一

  1

 十

  1

 十

 十

 十

 十

 十

 十

 5

 2 2

十.

 1

++

十十

.●、

 5

 2

●+

●十

 2

 2

 1+

 1

十 1

●+

 ●

 ●

 ●+

 ●

 ● ●

 5

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 十 十

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 4

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 7500

 902

 203

 102

, 52

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   2

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 3250

 1601

 1151

 378

 350

 252

 228

 202

 179

 150

 108

 105

 105

 102

 102

  57

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* Carex spp. include C. chrysol・s加sand C. altがniflora v.aureobrunnea.

abundant i‘nthe herb stratum. This association is also found on Mt. Akahoshi in Ehime

Prefecture, where the herb stratum is mainly characterized by such sedges as Careぶ

     i

duvaliana,C. alternifl�a var. aureobrunnea ̂C. sikokiana^  and C. ciliatomarginata・・

26 Research R of the K6chi Universi Vol. 8, No. 10

 The Spiraeeはm sikokualpinae,Yamanaka (spiraea sikoku心加na Association) is a

characteristic community which occupies exposed serpentine areas of Mts. Higashi- and

Nishi-akaishi. The physiognomical features of this community resemble those of the

Spiraeeはm blumei.but it can be regarded as an independent association containing a

large number of characteristic species。

 These associations are usually found in dry and sunny serpentine areas where edaphic,

topographic, and microclimatic conditions are not suitable to the establishment of the

forest.

 6. Other plant communities on serpentine in Shikoku      ,

 Besides the above-mentioned associations, there are grassy places in serpentine areas

mainly dominated by gramineous plants, such as Arundinariapyginaea,Miscantkus

sinensis,Arunぷnella hirta.etc..These result mostly from biotic factors, especially

from human agencies.

 Grasslands dominated by Arundinaria pygmaeaexistin low altitudes where pine forests

are cut down. Stable places in rather high altitudes are often dominated by Sasainoゆha

purpurascens. wherever they occur, ざ心・type or Sasa・Cα7?ヱーtype communities are most

common in serpentine areas,Misca?ithussinensisis found throughout the grassy places

in serpentine areas of Shikoku, but the well-developed Mis canthus -typecommunity is not

common. An example can be seen on Mt. Higashi-akaishi (Table 13).

 The vegetation on screes and bluffs in serpentine areas is sparse, and the characteristic

community is usually not found there, though some chasmophytes occur in such places.

 B. Floristic characteristics 01 serpentine vegetation

 The serpentine area supports a vegetation in distinctly physiognomical contrast with

that of surrounding non-serpentine areas and has a unique floristic composition. The ser-

pentine flora is so unusual that it is of great interest from both taxonomical and phyto-

geographical viewpoints. Problems on the serpentine flora have been discussed by alarge

nun!ber of botanists from various viewpoints. In Japan・, particular attention has been

paid to the relics, endemics, and disjunctive elements in the serpentine flora by Tatewaki

(1935, 1938), Kitamura (1950, 1952, 1957), Toyokuni (1955-58), and the writer (1950-

59). In this paper> the floristic composition of serpentine vegetation in Shikoku will be

discussed in relation to phytosociological and ecological problems.

 1. General characters of floristic compositions of serpentine vegetation

 The floristic composition of serpentine vegetation is of course different from one

another, but general characteristics are found throughout the world. According to Rune

(1953), the serpentine flora of North Sweden : (1) is relatively poor in individuals as

well as in species, (2) contains ’several species which are represented by particular

races (ecotypes), (3) includes many plants occurring very disjunctively, (4) contains

basicolous as well as acidicolous plants, (5) has a relatively xeromorphic character,

and (6) is often dominated by certain families or genera. In Shikoku also, all the

characteristics described by Rune, except (1), are observed.

A Phytosociological Study of Serpentine Areas‘・in Shikoku, TeぷanぐT.Yamapヽjaka) 27

          ●               d ●   ●-  ■● I●        ■  I Rune reported that only about 140 spec∃ies of vascular plants were found within the serpen-

tine areas of North Sweden. However, Whittakerイ1954) stated that the major serpentine

areas from the Siskiyou Mountains south into California were floristically rich. Coombe

and Frost (1956) also mentioned that on the Lizard in England plant cover Was usually

nearly closed and the number of species per unit area was certainly greater than

that on the granite soils. In Japan, usually characterized by the warm and humid climate,

serpentine areas have rather rich floras. Therefore, it is very difficult to enumerate all

the plants occurring on serpentine, including serpentine-accidental plants. In Shikoku also.

the flora of serpentine areas is not poor in either species or in individuals. For exan!pie,

                /about 420 species of vascular plants occur on serpentine of Mt. Higashi-akaishi alone.

 In the serpentine areas of Shikoku, families comprising comparatively many species

or dominants are Pinaceae, Cuかressaceae, Faeaceae, Ranunculaceae, a'erberidaceae.

SasiがTaeaceae, Hanianielidaceae, Rosaceae, Leeminosae, Aquifoliacedii Ericaceae,

Caprifoliaceae, Campanulaceae, Composi£ae, Grainineae, C:yperaceae. hiliaceae. etc.

Genera which contain characteristic or dominant species are Tsuga, Pinus・ Thuja・

Thitjofcsis, Quer‘CMS, Thalictru・m, Berberis, Epimedium,  Coりlopsis, Siephanandra,

spiraea, Rosa, Pour£hiaea, Lespedeza, Buエus. Heエ, Euoフリmus, H:ypericum,  Yiola,

£)iがomorpha, Angelica。Rhododendron, Enfeian£fius, Fraエinus, Siuertia, Viburnum,

Lonicera, Abelia, Scabiosa, Adenophora, 1)″り“・He£eropaかpus. Gymnaster, Saussurea,

Arundinaria, Sasa, Sasamorpha, Melica, At・undinella, M.iscanthus,  Carex,  Chiono-

graphis, Yeratrum, hilium, Smiはエ, Dioscore叫etc. It is a general tendency throughout

Japan that the serpentine aeas are characterized or dominated by above-mentioned families

                                                 ゝor genera. Among then!, especially, plants belonging to Ericaceae, Rosaceae, Graniineae>

and Carex are abundant in almost all the serpentine areas. Caryophyllaceous plants

which occur characteristically in serpentine areas in Europe and North America are not

found in Shikoku as characteristic or dominant species.

 The serpentine flora of Shikoku contains a large number of serpentinicolouS relics,

serpentinophytes, and disjunctive elements. It has been recorded from various parts of

the world that the serpentine vegetation has a xeromorphic flora. These examples will

be described and discussed further on.

 2. Characteristics o1 floristic compositions of Serpentine vegetation in

    Shikoku              Λ                  χ  ・   .                        Ft

 As mentioned before, the Quercusやりlliraeoidesscrub on serpentine is found only in

the Yawatahama district. At Korodokihana, twenty・two species were listed for five

quadrats and compared with the floristic composition oi Quei'cus phylliraeoides scrubs

in non・serpentine areas of Shikoku (Table 21).

 From this fact, it may be conc!uded that this scrub on serpentine should be included

in the Pi£tosporeto-Querceはm phylliraeoidetis, andits floristic composition is essentially

not different from that of stands on other rocks.

 The Abelieto・Pinetumdensifloraeshows a very characteristic floristic composition.

The lower strata of this forest are predominated by heliophilous and・xerophilous shrubs

28

Table 21. Quercus phylliraeoides scrub in non-serpentine areas of Shikoku

        (14stands in K6chi, Ehime, and Kagawa Pref.)

 Shrub stratum

Quefcus iitolliraeoidesPiitosPorum tobitaEurva etnargi?lata

Ligustrum j(ゆonteutnPinus thunbefgiiEuna i副!)onica

Paederia scandfiMS v. maireiSmilax chinaFraxinus sieholdiana

16 other species

8135. 6 538.4 125 74.9 72.8 72.1 39.2  4.9  0.7

竃I

Vllv

ⅣI

 Herb stratum

Far/ugium japonicum

MiscanμiUS sinensis2)iびanopterisdichotoma

C/Zぴsanihemum ornatum v.         ashtzunense

12 other species

269.2146.3125.7

 1.4

ⅦI

and herbs as shown in Tables 15 and 16, beeauseP・inusdensifloraon serpentine usually

forms open canopies. The composition of this association varies from place to place・ but

plants having xeromorphic characters are prevalent. Many constituents of this forest occur,

indeed, also on various kinds of rocks. Therefore, such a floristic composition on serpentine

may be comparable to that of special rocks and areas, such as limestone outcrops, granite

barrens, rocky river-banks, or coasts.

 Pinus dejjsiflora Iorestswhich are found in the Seto Inland Sea district of Shikoku are

well known as inferior forests. Comparing the floristic composition of such forests with

that of this association (Table 22), it may be stated that pine forests on serpentine are

  Table 22. Pinus densifloなforest in non-serpentine areas of the Seto Inland Sea District

               (20 stands in Ehime and Kagawa Pref.)

 Tree stratum

Pinus densiflora

Pinus thuタibereii

 Shrub stratum

Arundiれaria pygmaeaRhododendron lλjeyrichiiVaccinium oldhamiEurNa i副i)onica

Juntf'ei'usrtgida£yonia ellipticaRhododendron kaempferij?加ゐ心�yo serpy附加liuタ71Abelia setraiaレ々Zぼiniumsmallii V. glabrumLesf>e<iezaso.Quercus senataレ1Ilex PedunculosaAlnMs pendulaSymplocos prunifolia

64131937.5

17751564

 878.5 828 635.5 614 558 550 290 288 229 142.5 115 114 114 113.5

XV

可XX

1M

ⅨIV

ⅢM

ⅦⅣl

ⅢI

 Shrub stratum

Sポilax china

Viburnurnぴosum ・ ・ ●  ・Pierts japonica

Rosa lλaichuraiana

Pertya scandens

Rfitts tricfiocorSaAmela?ochier asiatica

Wisteriαfloribunda

30 other species

 Herb stratum

Diぴanopteris dichoto竹ta

Diplopteりgium glauc・um

MiscanihMS sine?usis

Cymbidium virescens

Solidago vifgaurea ssp・              astaticα

?1ぞダiぷum aQuilinum v.

            latiusculum

Platanthera mandati tioyurn27 other species

 109

 90 77.5

 53.5

 52.5

 28.5

  3

  2.5

3700. 5

 853.5

 542

 28.5 28

  6.5

  3.5

lⅣ

VⅣⅣ

l�

Ⅲ'

¥ⅣⅣ

】y

remarkab】y different from those on other rocks, though there are many common plants

occurring abundantly or constantly. In short, pine forests on serpentine are characterized

by the poorness of floristic elements ofclimatic climaxes, the occurrence of narrowly

endemic and disjunctive elements, and morphological changes of a number of component

plants。

 The poorness of floristice]ements of c】imaticclimaxes suggests that this association is

A Phytosociological Study of Serpentine Areas ・inShikoku, J (T.Yamanaka) 29・

not a serai stage. This problem will be discussed・further on. ・-

 Endemic and disjunctive elements combined with this association contain distinct charac-

teristic plants, and soine examples are listed below.

 Coりlopsisやicaは(Plate VI) occurs on serpentine in the K6chi district and is abun-

dantly found at Engydji, Nishikiyama, and the Osaka Pass. In Japan, Corylopsiscomprises

several species distributed very disjunctively. 0n serpentine, Corylopsisgotoana(PlateⅥ)

occurs in Jinry6, Tokushima Prefecture, and Coりlopsispauciflora"\sabundant on Mt.

Ooe, western Hon出合. From the above-mentioned fact, Cor-μopsis spicaはISconsidered

as a relic occurring only on serpentine. The cultivation in other soils is very easy, but

it does not grow in general from seeds.

 Ejikianthus peΓ�a以八sa serpentinicolous relic known only from Nishikiyama. Though

this species is rarely found in the above serpentine area, vax.iaponiciもs \s abundantly

found there and occurs also in several serpentine aeras in Shikoku. This variety is also

recorded from Honshu, occurring mostly on serpentine. It is easy to cultivate in other

soils.

 Hypericum.Z。saense is frequently found at Engydji, the Osaka Pass, and Aburaishi,

and may be a serpentinicolous relic. This plant is easily cultivated in other soils.

 Metercゆappushiやidus ssp≒ 1ゆ£ocladus(PlateVI) is frequently or abundantly found on

serpentine in the Kochi district and occurs very rarely in adjacent non-serpentine areas.

This subspecies is a narrow-leaved and glabrous serpentine race.

 Saussurea nipponicasSp・ yoshinagae(Plate VI) is frequently or abundantly found on

serpentine in the K6chi and Tokushima districts. &z心surea nipponica is a relic

including several local subspecies. Subsp・ :yoshinagaehas rather narrow leaves. The

cultivation in other soils is not easy.

 hilium japonicunivar. abeanu。z is frequent in Jinry6 and Sakashu, Tokushima Prefec-

ture. This is an angustifoliate variety di仔erentiated within serpentine areas.

 Clema£ispatens(PlateⅦ)is frequently found on serpentine in the K6chi district. It .is

distributed in Honshfl, Kyushu, and China. In Honshu, this is found on serpentine but

not restricted to it. In Shikoku, there are no specimen collected from non-serpentine

areas.

 Angelicacartilaginomarがnata isixequently found on serpentine in the Kochi district,

but the occurrence is not always restricted to serpentine.

 タaussurea ∫caposa is occasionally found in Kyushu and Shikoku. The occurrence in

Shikoku is restricted to serpentin in Shimagawa, K6chi Prefecture (cf. p. 5).

 Gymnaster pyg・niaetもSχS irequently found in serpentine areas of the K6chi and Yusuhara

districts. Outside serpentine, the occurrence is very rare. In Honshu, this is abundantly

found in some serpentine areas. ゛

 Carex mira \sa rare sedge which was known from several localities in western Honshu

and Korea. In Shikoku, the serpentine area of Aburaishi, K6chi Prefectureバs the only

locality of this species.

 Hemerocallis thunbereiiis frequent or abundant in the serpentine areas of the Osaka

                       1Pass, Ok6yama・ and Aburaishi, occurring also in adjacent non-serpentine areas. However,

the occurrence of this species is more closely connected with serpentine, and it is notice-

ろ0

Research ts of the K6chi Universi Vol. 8, No. 10.

able that this is found disjunctively in such low altitudes. The occurrences oi Scahiosa

idponica and Varatrum maackiivar. reytnojidianu?71, which are usually found in rather

high altitudes, are other examples in the same case.

 Besides, several plants which are usually found on the coast appear in serpentine

inlands. A typical example of them is Quercus phylliraeoides.which occurs in the

serpentine area of Sakashu. The same case is observed on serpentine hillocks in the

K6chi district, whereSetafia蛍石disvaT・pachysはchys l. najia, Ischaeinumcrassipes,

Ischaemum antheohoroidesvar. eriostschyum,Asparagus cochinchinejisis,et.c.occur.

 Morphological changes of plants on serpentine is known as "serpentinomorphoses. "

The term "serpentinomorphoses" is sometimes used as a synonym of serpentinophytes

(cf. Rune 1953)よ

 According to Pichi-Sermolli (1948), who studied the serpentinomorphoses of vascular

plants, they contain the following changes : 1. stenophyllism, 2. glabrescence, 3.

plagiotropism, 4. nanism, 5. a great development of root-system, and 6. glaucescence.

Besides, Rune (1953) noticed the tendency to become purplish in colour, and Kitamura

(1956) added lucency and crassism.

 According to the writer's observation in Shikoku, serpentinophytes are characterized

by stenophyllism, nanism, a development of root-system, and glabrescence. Moreover・ the

tendency to become purplish in colour is also found. 0n the contrary, glaucescence and

plagiotropism are not observed in Shikoku.

 Though they are not distinguished as definite taxa, tendencies of these morphological

changes occur in a large number of p!ants in the serpentine area. Some noticeable exam-

pies are described below.

 Stenophyllism is characteristic of Abelia serra£a \nthe Abelieto・Pinetum densザlorae.

and this plant 1S sometimes referable to var. 0bS外法�aは. The writer transplanted it

in other normal soils several years ago, but the small・leaved character is persisting・

Similar cases are also observed in other shrubs and herbs. NaniBm is mostly observed

combined with stenophy!lism. However, this change as well as the greatly developed

root-system is, as already stated by other authors, considered as an adaptive character.

The tendency to become purplish in colour appears in a number of plants. Especially,

it is characteristic of such plants as 7)εΓりa glabrescens, S tephana7idraincisa,Rosa

onoei,Hydrangea macr砂岫lla ssp. serrata, Viola spp., etc. occurring in theAhelieto-

P緬eはm densザlorae. According to the cultivation experiment of Stゆhanandra・incisa-,

this tendency persists in other soils.

 Thus, serpentinomorphoses may have adaptive and non-adaptive changes. Some serpen-

tinomorphoses which seem to be adaptive characters may be xeromorphoses, since similar

morphological changes appear in other special places, such as limestone outcrops, rocky

river-banks, coasts, etc. This fact is confirmed by the writer's observation in Shikoku

(cf. Yatnanaka & Takezaki 1959), but it has been already recognized and discussed by

other authors (cf. Rune 1953).

 The Rhodoreto・Ch。naecypaパぬほm obtiisae χs distinguished from the C hamaecypari s

Obはsa forest in middle Honshil by、such floristic elements asRhododendron ine.ttp.rnir.hii

(et var. hondoense)・neエruBoavar.stenoph:vlla、 lieエs‘uzeroki ssp. loneipedu、iculata、

Viburnu-)n urceolatum. Rhododendron keiskei. etc. In the serpentine areas of Shikoku,

as already stated, the forest dominated by Chamaecy夕aris obtusa Vs iound on Mt. Shiraga

and Mt. Higashi・akaishi.

 The forest on Mt. Shiraga is an excellent stand in Shikoku.Chamaeりparis obはsα;

Pinus pentaphylはvar.himekomatsu..Tsuga diversifolia. andTsuga sieboldii predomi-

nate in the tree stratum, attaining 20m. in height with closed canopies. The shrub stratum

is generally dominated by Rhododendron metternichiivar.  hondoense.whileSasaniorpha

夕a.やurasce刀5 is prominent in some places overlaid by rather deep soils. The herb stratum

is mainly constituted by Carex reinii-, Rumohra mtUica, and such prostrate shrubs as

Hにr叫vat. intermedia l。repens. The

ground beneath the forest is well covered with such mosses as Hyloconiium splendens.

Pleurozium schreheri. Bazzania spp・, etc.

 The forest on Mt. Higashi-akaishi is rather stunted. In the tree stratum,  Thuj a

standishii.which is not found・ on Mt. Shiraga, is abundant. The shrub stratum is

constituted by ]i.hodode7idron me£terniとhii. Her sugerok・i ssp. longipedunc�ata.

Cle£hra barbiner。is, Parabenzoin trilobum, Khodode・ndron 加71£呻JりHum,etc. Care:,『

Mゆharicarpa ls abundant in the herb stratum, but its major constituents are similar to

those on Mt. Shiraga. The H:μocomium-X.ype floor appears also in this forest.

 As already described, the xeromorphic flora usually appears on the south-exposed side

and rocky ridges of the serpentine mountains. On the contrary,。.the characteristic

serpentine flora usually disappears on the moist northern side where・ sciophilous and

hygrophilous plants are found in the forest (cf. p. 19). Therefore, closed and well-

developed Chamaecyparis-\.ype forests lack characteristic plants of serpentine, although

some subalpine and alpine elements occur there. The floristic difference between Mt.

Shiraga and Mt. Higashi・akaishi may partly be explained by the above-mentioned fact.

 In the Chaniae cyparis -type forest, the ground is usually covered with the humus layer.

This has a causal relation to occurrences of acidicolous plants. A typical example is

RujnoKra ・,・nutica.which is frequently found on acid soils in coniferous forests.

 Communities dominated by spiraea are generally found in limited areas, such as rocky

places, ridges, southern sides of mountains, etc. exposed to the intense light. Physiog-

nomical characters of communities dominated by spiraea blumei1nlimestone areas

resemble those on serpentine, but their floristic compositions are more or less di仔erent

from each other as shown in Table 23。

 In spite of these differences, it may be stated that these communities are floristically

characterized by some plants common to serpentine and limestone and the occurrence of

endemic or disjunctive elements. It is observed that several disjunctive elements are found

on limestone as well as serpentine. Moreover, many plants characterized as calcicolous

occur also on serpentine. Some examples in Shikoku, which are not always combined

with one association, are listed below (see also pp. 33-34)。

 Gyninocarpiumjessoense(PlateⅦ)occurs abundantly on Mt. Higashi-akaishi and is

also frequent on Mt. Nishi・akaishi. Outside serpentine, it is found in limestone areas

of Mt. Tsurugi and Mt. Ishidate. In Japan, this fern is distributed very disjunctively.

ろ2・Research Reports・ of the Uniyersity, Vol。8, No. 10.

    Table 23. Spiraea blumei scrub in. limestone areas

(5 stands on Mt. Tsurugii Mt. Ishidate, and Mt. Tengudzuka)

  Shrub stratum

 SWroeo bltttnei

 Salix sieboldiana

 Deutzio firacilis

.Abelia spathulata

”Rhamrois iaponica v.             fieci(>iens*

`Ts昭a <iit)crsi/olia

、CarPinus ca?ゆinoides Zabぷa integrifolia

 P6yりagiobrescens

 Abies homolei>is

 spiraea japonica

 Fずaxinus loMMgitiosa v. sertaia

 Coりlus sieboldiana

 万客ionyボus alatus v. rotundatus 7右・Iwingia知加タ兪a v.・

             parvifolia

 LifLustfum obtusi/ottum

 Abelta senataV. buchuialdii

 Clematis ausifojaponensis**

  Herb stratum

 Thaμdrum minus     ’

 Cafex duvaliana

 Gera?itium shikokianum

 Galium kinuta

" 5350  750

  454

  450

  352

  350

  350

  350

  350  102

  102

  102

  100

  100

  100

  100

    4

    2

  2350

  1900

  750

  750

VIⅣ1

11IIIIIllII

Illl

!VVI

 Herb stratum

Valefiana sambucifoliav..             /auriei

AruMcus dtoicus v. tenuifoliusCarc% blePhaficarPaAdenoPhofa rcmofi/toraAslilbe shikokiaれa゛Brachypodium sylvaticutnv.

            脚iSがg脚Bisiorta majorV. niten?レ゛ぴalritmmaackiiCoptiiii夕1�swilfordiiArabis gernmt fera**Chrysanthemum boreaie*をLigularia stenoce{>fialaFesiuca ovina v.

         chiisanensts**Anapharis margぽitaceav.

          angusti/oliaPedicularis resupi皿ta£芦田edium grandiflorumH郊ぴtcum sikokumonianumTia昭ia tachiroei**Anafk�is si≪ica**Leontopodium iapomcumV.

          spathulatumSaussurea nipponicassp.

            sikokianaSausstがea arnabilis**

550452354202200

200106104102102102102

102

100 6 2 2 2 2

 2

 2 2

Il

ⅢⅢI

!N

ⅢIIII

llIITI

II

    * Not found on sをrpentine in Shikoku. 昔昔 Disjunctive elements occurring on

     limestone and not found on other rocks in Shikoku.

occurring mostly in limestone and serpentine areas (Shimizu 1958). In Europe, closely

related species,  Gyinnocarpium.7・obertianumis regarded as a typical calcicolous plant.

PoりsttchuTiicraspedosor・UTIl\s frequently or’abundantly found on limestone and some-

times occurs on other rocks. As to serpentine, see p. 19.

 Camptosorussihirictts(PlateⅦ) is frequent or abundant on limestone. Sometimes it

occurs on other rocks, such as schists. As to serpentine, see p. 19.

 Coptidipteris ivilfordiiIS 肪nerally found in limestone areas, though it occurs in other

soils. This is frequently found on serpentine of Tomisato, Mt. Higashi・akaishi, etc.

 B・acMりa lanceolata"vsitequently found on limestone at the foot of Mt. Ishidate and

is not rare on serpentine in Sakashu.

 Thalictrum minus\sabundant or frequent on both serpentine and limestone.

 Kpiniediumsrandiflorum isgenerally found throughout limestone areas in Shikoku and

is abundant in serpentine areas of Shimagawa, Nishikiyama, etc.

 Arabis serratavar.sikokiana χS often frequent not only on limestone but also on

serpentine.

 spiraea bl皿^ei is often dominant on limestone as well as serpentine as described above.

 spiraea nervosa is usually abundant on limestone outcrops. It is frequently found on

serpentine of the Osaka Pass.

 Buエus microphyllavar・japonica.See p. 12. and p. 19.

 Hosieaiaponicais a rare plant occurring mostly on limestone. It is occasionally found

on serpentine of Mt. Higashi-akaishi.

 Bupleurumfalca£imivar. komαΓ。由i is sometimes abundant on limestone and often

frequent on serpentine in the K6chi district.

 Zabeliahitegrifolia is a characteristic plant of scrubs on limestone outcrops and is

frequently found on serpentine in Jinryo. See also p. 12.

 honicera■inochidzukianais found in some limestone and serpentine areas.

 Valerianasambucifoliavar.fat£riei is frequent in limestone fields of Mt. Tsurugi

and Mt. Ishidate, and is abundantly found on serpentine soils of Mt. Higashi-akaishi

(cf. Table 13).

 As to the occurrences of the above-mentioned plants, the calcium content of serpentine

may be considered. So far as the composition of serpentine in Shikoku is concerned,

however, the calcium content is as a rule very low, CaO occurring only in trace or not

detected. It mav thus be considered that the occurrences of these plants do not necessarily

result from the calcium content of serpentine soils. Therefore, it can be stated that

other environmental factors, such as topography, physical properties of soils, and

microclimate, have causal relations to them. But 由is problem is so complex that it

must be solved by experimental investigations.

 The^Spiraee£uin sikokualpinaeonMt. Higashi-akaishi represents floristic distinc-

tiveness. Among the constituents of this association, the following serpentinicolous relics,

serpentinophytes, and disjunctive elements are found :

 励行aea sikokuaゆnna (Plate V) is often treated as a variety oi Spiraea blumei(spiraea

bluineivar. pubescens). Many species of this genus occur on serpentine in Japan. Closely

related species, spiraea hluinei and ・spiraeahayatae,are飢so found on serpentine in

Shikoku. Plants of this genus are more or less restricted to special rocks or areas. For

example, spiraea tosaensis,which is closely related to spiraeanippojiicciand Spiraea

ogaiuatoccurring on serpentine in Houshu, is found only on rockv river-banks in Shikoku

(Morishita & Yamanaka 1957). Thus, occurrences of plants of this genus are limited.

but they are very easily cultivated in other soils.'・     F      ふ F・

 l^oniceranioch這zukianavar.filが'or mis isfound only on Mt, Higashi-akaishi. This

should be a variety differentiated froml^onicera mochidzukia・na within the serpentine

area. The leaves of this variety are rather purplish in colour.

 Adcj^ophoratriplリllavaT. pviellaris \san endemic plant which is abundantly found

only on serpentine of Mt. Higashi-akaishi. This is an angustifolia.te variety of Adeno-

かhora triphylla and is closely related to the narrow-leaved form of val・iaponica.

Morphological characteristics persist at cultivation in normal soils。

                                             ● lierherisamtぼensisvat. brevifoliaVs abundantly found in the serpentine areas of Mts.

Higashi- and Nishi-akaishi. This is distributed very disjunctively in Japan, occurring

mostly on serpentine and calcareous rocks (Shimizu 1958 ; Yamanaka 1959).

 V ar. cinium Tjitis-ifin.ROvar.minusoccurs near the summit of Mt. Higashi-akaishi,

and this mountain is the only locality of this alpine plant, in Shikoku.

 Rosa 711pponensis 〇ccurs abundantly in the serpentine areas of Mts. Higashi- and

Nishi-akaishi. Outside serpentine, this appears in the limestone area' of Mt. Tsurugi.

 Viola biflora appears abundantly on serpentine of Mt. Higashi-akaishi. Outside ser・

Dentine, this occurs on limestone of Mt. Ishidate and is also found on other high

M Research ts of the K6chi University Vol. 8, No. 10

mountains.    ,  ・・。

 Angelicauhatakejisis(Plate vn) is abundant on serpentine of Mts. Higashi- and Nishi-

akaishi.         ,

 Primula血百nosa ssp. ■inodesta \sabundant on serpentine of Mts. Higashi- and Nishi・

akaishi. It is occasionally found in several places of high altitudes in Shikoku and occurs

also on limestone of Mt. Ishidate.

 E・ゆhrasia  microphyllais a plant characterized by its very small and slender form.

This is an endemic species in Shikoku occurring abundantly on serpentine of Mts.

Higashi・and Nishi-akaishi. Outside serpentine, it is found on limestone of Mt. Ishidate.

 Scabiosajaponicavar.alpinaoccurs frequently on serpentine of Mt. Higashi-akaishi

and is occasionally found in non-serpentine areas. such as Mt. Ishidzuchi, Mt. Ishidate,

etc.                                。

 C. Physiognomy of serpentine vegetation in 】:elation to vegetation types

 Generally speaking, the vegetation in the serpentine area contrasts strikingly with

that of adjacent non-serpentine areas, and it often serves to delineate the geological

discontinuities of an area. The contrast depends on physiognomical characteristics as

well as floristic compositions of the serpentine vegetation. Therefore, the physiognomical

research is quite necessary in the present work。

 The physiognomical types in the serpentine vegetation are varied, but they have some

common tendencies and characteristics. As already pointed out by Whittaker(1954), the

serpentine vegetation is to be dominated by some one or by some combination of three

growth-forms, namely, coniferous trees, sclerophyllous shrubs, and grass-like plants.

Accordingly, it is also to be emphasized that the dominant species should not be overlooked

from physiognomical viewpoints (cf. Table 24). From the above・mentioned fact, the

Table 24. Comparison between floristiccompositions in terms of major growth-

 forms of climatic climaxes and those of serpentine vegetation. Figures in

 Gothic indicate abundant growth-forms.

Trees Shrubs Herbs

Conifer Evergreen Deciduous Evergreen DeciduousGrasses

  &

SedgesOtherHerbs Ferns

Shiietum '■'

 (Non・serpentine)Pinetum '' (Serpentine)

CyciobalanoOsidefum') (Non-serpentine)Pinetum幻 (Serpentine)Fagetum ^' (Non-serpentine)

C加州aecypari心細附6) (Serpentine)

Chamaecy paridetum?) (Serpentine)

5.3

5.5

4.5

2.6

12.1

 9μ

15.9

 -

13.0

 -

 -

 -

 -

 -

 -

 -

 -

18.4

 -

 -

56.8

 2.6

42.3

 4.5

 5.3

IS.1

13.6

 2.3

36.8

24.1

44.7

44.8

48.5

40.9

 -

14.5

16.4

7.9

9.1

4.6

 6.8

38.2

 1.8

23.9

 7.9

 6.1

18.2

18.2

 2.6

13.0

 6.0

13.2

 9.1

13.6

1) Forest in Usa near K6chi city. 2) K6chi district(Table 16). 3) Forests in Yusu-

 hara and Taish6. 4) Yusuhara district(Table 15). 5) Forests on Mt. Kuishi and

 Nakanokawa Pass.' 6) Mt. Shiraga (Table 18). 7) Mt. Higashi-akaishi (Table 17).

n(TこY八manaka) 55

vegetation types in the sense of Suzuki (1953. 1954) are chiefly adopted in the physiog-

nomical research of the serpentine vegetation.

 According to Suzuki) the vegetation types are similar to Griesebaとh'S "Vegetations-

formen,"which resemble Cajander's forest types, being distinguished among canopy

trees and herbaceous plants respectively. They are picked up from the association domi-

nants.

 In the serpentine vegetation of Shikoku, the following main types can be recognized :

  Canopy types.

    Maquis-type  :    Quercusがりlliraeoides.

    Pinus・tvpe  :  Pinus densiflora,1=>. thiぷnbergii, P. pentaかhylla Vat. hime-

                 lioviatsu.

    TS昭a-type   :   Tsuga diitersifolia, T. sieboldii.

    C haniae cyparis -type : ChamaeりParis obtusa, Tluりa.sはndishii, Thuiopsis dolab-

                 7MZどZ.

Forest-floor types

  Sasa-type

Rゆeris-type

   Rhododendron-type

   Miscanthus-iype :

   Car'ex-type  :

   Dicranopteris・・type

   Uypnum-type :

   Cladot・lia-K.ype :

By combining canopy

  Arundhiaria pygmaea・Sasa tenuissin・la, Sasamorpha pur-

  夕ura∫ceils.

  Heエrueosavar.stenoか1リlla, Skinvm.ia japonicavar. inter-

  ■media i.repens.

 : Rhododendron ■inettemichii.

  Iχdiscan£JlUS sinensis, M. oligos£achyus.

  Car ex spp.

: Dicranopteris dichotoma, L)iplopterygium glaucu,刀l.

  H:ypnuni spp・

  Cはdonia r・1が.μΓリm.

types with forest-floor types, the vegeはtion types of plant

communities in serpentine areas are shown in Table 25.

 As already stated, all the canopy types, except the Maquis-type, are represented by

            Table25. Vegetation types on serpentinein Shikoku

 Canopy types

Maquis-type

Pinus-type

Chamaecyparis-type

  (Tswfia-type)

Forest・floor types

Dicta?loかteris-type

Cladonia-type

Hv{}竹t匹-type

Miscanthtぷs-type

Caダex-type

Sasa-type

Rhododendron-type

ReJ)6ns一旬pe

Hy!ocomium-type

56 Research of th!I K6chi University Vol. 8, No. 10.

coniferous trees in the serpentine vegetation. and Maquis-type scrubs resemble sclerophyl-

lous scrubs. 0n the other hand, such forest・floor types as the Sasa-, Miscanthus・, and

Cαy?ヱーtype are found throughout the serpentine areas. These facts agree well with

Whittaker's description cited before. The climatic climax communities dominated by

broad・leaved trees are represented by such canopy types as the Shiia・, Machilus-,

Cy clobalanops,is-,and Fagus-type. In short, it may be roneluded that the peculialities

of the physiognomy of serpentine vegetation are partly due to such characteristics of the

vegetation types。

 The stunting or impoverishment of growth is one of the most striking features of the

physiognomy of serpentine vegetation. This physiognomical characteristic is well observ-

ed in pine forests, where tree stratum is reduced and the lower strata are increased. Open,

stunted, and discontinuous canopies permit the growth of a large number of heliophilous

and xerophilous plants in the lower strata. In Japan, therefore, serpentine pine forests

are often known as the "garden-like vegetation.”

 When biological spectra of' such pine forests are compared with those of other・forests

in non・serpentine areas, it becomes clear that micro- and nanophanerophytes (especially

nanophanerophytes) are prevalent in serpentine areas. Though Raunkiaer's original

biological spectra seem to be of little value from sociological viewpoints (cf. Suzuki

1954), the above fact shows that the vegetation is stunted and shrubby. In addition.

seasonal aspects are also noticeable in pine forests, because many deciduous shrubs are

found there。

 The Chamaeりparis-type forest forms closed canopies even in the serpentine area, and

the occurrence of the Hylocom・ium-Vype forest-floor, which seems to be closely related

to the subalpine forest, is noteworthy from floristic viewpoints as well as vegetational

ones. The forest on the northern side of Mt. Shiraga is, as already stated, not stunted,

and is an example showing the fact that closed and well-developed forests can be seen in

Table 26. Comparison between physiognomical characters of climatic

  climaxes and those of serpentine vegetation

Climatic zones

Rocks

Forests

Abundant life-

forms

 of flora

 of vegetation

Dominant

canopy-types

Tree stratum

Warm -temperate

Non-serpentine

Evergreenbroad-leavedforests

Phanerophytes

Phanerophytes

Maehiltts

Shit a

(ニlyclobalan砂sis

Closed and

not stunted

Serpentine

Coniferous

forests or scrubs

PhanerophytesHemicryptophytes

PhanerophytesHemicryptophytes

Piれ.us

Open andstunted some-times absent

COOI・temperate

Non-serpentine

Deciduous

broad-leavedforests

Phanerophytes

Phanerophytes

Fagus

Closed andnot stunted

Serpentine

Coniferousforests or scrubs

Phanerophytes

Phanerophytes

Chatnaecyparis

sometimes Tsuga

and Ptnus

Usually closedsometimes stunted

Lower strataSparse~dense

Sparse~dense, )sometimesbarren

densesparse~densesometimes barren

serpentine areas.

 Summarizing the above physiognomical characteristics of the serpentine vegetation com-

pared with thsoe of climatic climaxes on other rocks, evident contrasts are presented

in Table 25.

D. Succession of Serpentine vegetation

 As mentioned before, the serpentine areas of Shikoku are situated from the warm-

temperate to the cool-temperate forest regions. In the warm-temperate forest region,

evergreen broad-leaved forests consisting mainly of C卵なnopsis, Cyclobalanop了is, and

Machilus,ate iound,while the cool・temperate forest region is mostly predominated

by deciduous broad-leaved forests, such as 凡7g心。?7iata forests. But typical stands of

these climatic climaxes are of course not found in the serpentine areas.

 It may be considered that forests in serpentine areas have characters of preclimaxes

which occur on dried and infertile soils. It may therefore be assumed that such plant

communities are serai or successional and will converge to essential identity with climatic

climax communities with the development of soil maturity.

 In fact, there are some places overlaid by rather deep and weathered soils in serpentine

areas, where fragmental stands of the climatic climax forest occur. Judging from edaphic

factors, however, it should not be assumed that serpentine areas are to be occu-

pied by climatic climaxes. Whittaker (1954) also recognized that the Siskiyou serpentine

vegetation is not a serai stage but is a distinctive climax. The writer cannot support the

monoclimax theory. Consequently, it is considered that the Maquis-, Pi7ius-, and

Chaniaecyparis-typescrubs or forests are to be re・cognized as climaxes.

 The Maquis-type scrub and the Chaniaeりparts-type forestare of course edaphic or

topographic climaxes which occur on various kinds of rocks as well as serpentine. Though

the Pinnx-type forestis generally characterized by apparent initial stages from its

floristic and physiognomical features, it can also be regarded as an edaphic climax. .

 As mentioned above, edaphic or topographic climaxes are to be essentially recognized

in serpentine areas of Shikoku, and occurrences of grasslands result from other biotic

factors, especially from human disturbances. In the latter case, the vegetation often

continues an initial stage.

 The development of vegetation on serpentine is rather slow as compared with that on

other rocks, and this tendency is marked in denuded or barren areas. Consequently,

the secondary vegetation occupies extensive serpentine areas where the establishment

of the climax communities cannot be expected in the near future.

   IV. Comparison with serpentine vegetation in other parts of Japan

 From the results of the vegetational study described above, the following facts can be

summarized :

 1. The Maquis-type scrub dominated by Quercusphylliraeoidesappears in the coastal

district.・                   ●        ぐ心,

 2. The Pi・nits-typeforest dominated by 7)iivus densifloraoccurs in the warm-temperate

ろ8 ts of the K6chi University Vol. 8, No. 10

forest region.

 3. The C hamaeりParis-type forest with the j)inus- andTsuga-type is mostly found in

the cool-temperate forest region.

 4. Spir.αea is observed on serpentine in some places.

 5. The Sαjα-Car?ヱーtype grassland is commonly found as the secondary community.

 When the serpentine vegetation in Shikoku is compared with that which has been

reported from various parts of Japan, it is obvious that the above-mentioned tendencies

are common throughout serpentine areas in Japan.

 Maquis-type scrubs often accompanied by Pinus thunberzii ate found in the Kinki

district, western Honshu. The Pi nits thunbersii-Quercus phylliraeoides community on

Sugashima, Mie Prefecture (Kitamura & Momotani 1952), and the Qt£ercus pりlliraeoides

community on Mt. Asama, Mie Prefecture (Taniguchi 1958), are examples.

Such communities dominated by Querctts phがliraeoides are similar to those in the

Yawatahama district and should be included in one and the same association.

 Pinus-type forests in coastal districts of western Honshu are often dominated by Piniis

thui・tbergit.For example, the 7)inus thunbergii一Btにus microかhvll lavar・]dponicacom-

munity on Sugashima (Kitamura & Momotani, 1. c.) and the PlJlWS thunberg・ii・Pour£hiaea

villosa vaて, laevis community on Mt. Asama (Taniguchi, 1. c. ) occur in the serpentine

areas. However,Finns d。isiflora  iorests are commonly found in lowlands of western

Honshu (Kitamura, Hiroe, & Nakai 1950 ; Kitamura, Murata, & Torii 1953 ; Ogawa

1950 ; Taniguchi, 1. c). Physiognomies and major constituents of these pine forests

are similar to those oi the Abelieto-Pinetum densiflorae in Shikoku. Therefore, the

flora and vegetation on serpentine in lowlands of westsrn Honshu are well comparable

to those of Shikoku. Occurrences of the following characteristic and noticeable plants

show that the serpentine vegetation in lowlands of western Honshu, which is as a rule

sharacterized by open pine forests, has general floristic features in common with

the serpentine vegetation in lowlands of Shikoku ■。Bucfele・りa lanceolata. Clematis patens.

Be・rberis thunbergit, Coりlopsis go£oana, S£ephan。・idra incisa. Btにus microph^ぶ.a var.

japonica・ L)iplomort)ha sifeofetana, E-nkianthus perulatusvai・ iaponicus, Abelia serra£a,

Perりa glabresce・ls・ Dioscorca graciUi・ma. Tithymalus pekinensis^ Bupleuruni falcatu・,・,l

vat. komaroivi. Scabiosajaponica, Heteropappus hispid・ws, Gymnaster j):ygmaeus.

A£・racりlodes japonica. He・mero c� Its thunbergii.etc.

 Ch,ainaecyparis-type forests on serpentine in the mountainous region of middle and

northern Honshu are tobe represented by the vegetation on Mt. Shibutsu and on Mt.

Hayachine. The TJ・luia standishii forest on Mt. Shibutsu, Gumn!a Prefecture, was treated

as the Thuj etum Sはndishii, nom. prov.by Suzuki(1954). Besides, the flora and vegetation

of this mountain are, as already pointed outby Kitamura (1952), closely related to those

of Mt. Higashi・akaishi. The following plants are common to both mountains : Gymヽuo’

calやi・Em・1 jcssoense,  Ti・luja staTidisl・iti, Tsuga diversifolia, Sabina sargentii,Berbe石s

a切.arensisvar. brex瞳folia, Mag・1011a salic・げolia, Saエ迂raga for£‘unei, Sorbus commixta.

Rosaiiipponensis. Tit

Iりmalus siebold・ianusvar. -montanus,  Viola bifiora,  Chaniae-berv-

climenum canadense, Menztesia pen£andra,  Vaccinium vitis-idaea, Pri・niulafarinosa

ssp. modesta.  Viburnum. furcatu!・n, Aletris foliata. etc. Related plants which include

the vicarious are as follows ■・ Pinus pentaphylla(on Mt. Shibutsu)→7)・pentaphyllavar.

himekomatsu(on Mt. Higashi・akaishi), spiraea nipponica→S. sikoktialpina. Ilex

n(Tr.YAMANAKA)ろ乳

sugeroki ssp. bre・vipec

degronianuni→尺.niettemichii. Rhododendron tschonoskii var. Zパフlervど→R. tschono一

sfeii, Tritoinodon. campanulaは5 var・palibini→T,cernuus,Euphrasia insignis→£.

mic・ropりHa, Leontol>od・turnjaponicum. -* L. iaponicuin var. spathulatum, Solidago

vireaureassp. leiocarpa→5. vir・gαzz7?a ssp. asiatica, Festuca ovinavax. s・upi・na→F.

nvina var. ovina, Vera£r副n maackii var.reyinondianum et var・ par折夕ora・,n→y.

inaackii, etc. (cf. also Hara & Mizushima 1954).

 The Thujopsis dolabratavaiンho・ndai Iorest on Mt. Hayachine (Kitamura 1952) has also

floristic and physiognomical resembrances to the Rhodoreto-Chan・はeりparideはm obtusae

and the T Kuj opsidetumdolahratae in Shikoku.

 Spiraeaoccurring on Sざrpentine contains several major species in Hokkaidd and Honshu

(cf. Kitamura 1952). spiraea betidifolia and its related species occur in Hokkaid6 and

northern Honshfl. In middle and northern Honsh危, spiraea nipponica \s often abundant.

and its related species, 励iΓaea ogamai, is abundantly found in some serpentine areas of

the Kii Peninsula, western Honshu. spiraea 711ervosaoccurs frequently near Sekinomiya

in Hydgo Prefecture, western Honshu.

 It is to be added that the Picea-Vype forest and the Pi・nus -type scrub are not found in

Shikoku, whereas the P・icea glehni iorest occurs in Hokkaidd (cf. Tatewaki 1944 ;

Tatewaki & Morimoto 1933) and the Pinus pamila scrub is found in Hokkaid6 and

Honshu. These forest and scrub are found in subfrigid and frigid zones.

       V. Environmental factors affecting vegetation on serpentine

 The serpentine vegetation varies greatly with the different p芦rts of the world. This

vegetational difference is of course due to different geographical and macroclimatic con-

ditions. On the other hand, concerning remarkable characters of the serpentine

vegetation contrasting with surrounding vegetation on other rocks, it w川be em-

phasized that edaphic factors have causal relations to the peculialities of serpentine

vegetation. Therefore, particular attention has been paid to the nature of rocks and soils

of serpentine. However, it is a matter of course that the peculiar plant life on serpen-

tine may ecologically be understood on the basis of considering the total environments.

 A. Topography and microcliraate

 According to differences in the climatic zones, various types of serpentine vegetation

appear in different parts of the world. In Japan also, the types of serpentine vegetation

are diverse as already mentioned (cf. pp. 37-39). On the other hand, local distinctiveness

in vegetation within a serpentine area is mostly due to topography and microclimatic

elements.

 IIt has been already stressed by all authors that the serpentine vegetation occurs to be

characterized by xeromorphism. This character may be closely connected with topography

and dry microclimate in serpentine areas. In the serpentine vegetation of Shikoku, the

Pinus-X.ypeforest has the most xerophilous and heliophilous characters, while the Chamae-

り'戸αΓむ・type forest・is more hygrophilous and sciophilous. Therefore, the characteristic

features of serpentine vegetation appear mostly in open stands where the plants are ex・

、40 Re、search of the K6c‘hiUniversity Vol. 8, No. 10

posed to the intense light, higher temperature, and air movement. Such microclimatic ele-

merits vary distinctly with eχpositions. For example, conditions of the northern side of

the mountain differ greatly from those of the southern side, and differences of the flora

and vegetation in such case were already ・mentioned. In short, it may be stated that the

microclimatic elements, together with topography and other edaphic factors, are more or

less responsible for floristic compositionsand physiognomical characters of serpentine

vegetation.

 B. Physical and chemical properties of serpentine soils

 Since Novkk (1928) explained the influence of serpentine on the vegetation, many

authors discussed the problems of rocks and soils of serpentine in relation to the plant life,

and it has become a well-known fact that the serpentine soils are infertile for the growth

of most plants. Recently, these problems were reviewed in detail by Krause (1958)。

 In most areas, serpentine rocks weather slowly. Soils are usually shallow, and parent

rocks crop out on steep slopes and in places with sparse vegetation. Though mechanical

compositions of serpentine soils differ from place to place, they generally contain con-

siderable unweathered rock particles. This tendency is more or less peculiar to serpen-

tine soils in 10w altitudes. The gravel content exceeding 60%iS established of the soil

in the pine forest of Nishikiyama. Such soils result in reduced moisture and nutrient

levels. Physical characteristics of soils, such as gravelly texture and low clay contents,

are unfavourable for the growth of plants. Besides, as already discussed, the depth of

soils is often closely correlated to the vegetation. Thus, the shallowness of soils is also

a cause of the occurrence of characteristic serpentine vegetation. On the contrary, ser-

pentine soils often form heavy clay in rather damp places. Some plants which are not

found in other parts of serpentine areas occur in such places.

 Chemical compositions of rocks and soilsof serpentine vary considerably with situation,

but high contents of magnesium and very 10w contents of calcium are striking. Therefore,

Novdk(1928) firstpaid attention to the ratio MgO/CaO in serpentine soils. Afterwards,

Lammermayr (1928) criticized Novdk's theory, attaching importance to physical

properties of serpentine soils. More recently, Walker (1954, 1955) and Kruckeberg (1951,

1954) stated that low calcium level was the basic cause of the peculiarity of serpentine

soils. Rune (1953), supporting the assumption of Robinson and others, concluded that

the rather high contents of chromium and nicke! were general and their toxity was a

dominant cause of infertility. On the other hand, Pichi・Sermolli (1948), recognizing the

effect of high concentration of chromium, emphasized that such factors as microclimate.

mechanical composition of soils, and others might be more responsible for the unique

character of serpentine vegetation。

 Serpentine rocks in the vicinity of Kochi city contain more than 35%Mgo in general,

while the calcium contents are only trace or are not detected at all. Chromium and

nickel also occur and are detected in rather high concentration from the subterranean

water in serpentine areas. The relations between the chemical composition of serpentine

soils and the plant life cannot be solved without experimantal investigations. Therefore,

full discussion cannot be done here becau‘seno eχperimental result is available in Japan.

  In scrubs and open forests; the ground is hardly covered with the humus layer. The

pH values are usually between 6.2 and 6.9 in serpentine soils in the vicinity of K6chi

                  4city and Tokushima Prefecture. On the contrary, in the ChamaeりParis-typeforest,

・A-horizon is strikingly developed, the soil being quite acidCcf. Miyazaki 1942). It will be

realized from such facts that, as alreaby mentioned by Rune, plants found in communities

retaining the initial stage are more or less basicolous, while those in the climax'forest are

usually acidicolous.

  Itmay be said in conclusion that the infertility of serpentine soils is caused by chemical

properties and is also connected with physical properties and many other environmental

factors.                      '・

  C. Biotic factors

  There is no biotic factor which has causal connection only with serpentine. However,

it may be stated that such human agencies as the fire, grazing, etc. are noticeable from

floristic, structural, and successional viewpoints of serpentine vegetation.

  Finally, in such special habitats as serpentine, competition is not severe. It can be

recognized through the study of serpentine vegetation that this low competition must be

stressed in connection with the problems of distribution and ecology of plants.

                      VI. Summary

 Recently, the remarkable plant life on serpentine in Japan has been noticed by some

botanists, though the peculiar serpentine florawas already described by Yoshinaga in 1914.

During the last ten years・the writer has studied the vegetation in serpentine areas of

Shikoku, and summarized his study in the present paper.

 In Shikoku, serpentine occurs in Ehime, K6chi, and Tokushima Prefectures, and each

area studied is outlined on pp. 2-20.

 As the result of vegetational analyses, the following main plant communities were

recognized in serpentine areas of Shikoku :

 (1)Pittosporeto-Quercetum phylliraeoide£is.This association is a Maquis-type com-

munity dominated by Quercus pりlliraeoides.whichoccurs mainly in coastal districts of

southwestern Japan. In serpentine areas of Shikoku, this is found in the Yawatahama

district. The physiognomical characters of this association resemble those of sclerophyllous

scrubs. The floristic composition of this community in the serpentine area is not essentially

different from that of stands on other rocks.

 (2)Abelieto-Finetum densiAorae. Thisassociation is an open and stunted pine forest.

Serpentine areas in low altitudes of Shikoku are mostly characterized by this association.

Floristic and physiognomical characteristics of this association contrast strikingly with

those of the vegetation in surrounding non-serpentine areas. Similar pine forests occur

also on serpentine within the area of the warm-temperate forest region in HonshO.

 (3)Rhodoreto-Chaniaecypari.detum ohtusae.ThIS Chainaeりparts・■typeforest occurs

mainly within the area of the cool-temperate forest region. Though ChamaeりParisohtusa

                iforests which are included in one and the same association are found on various kinds of

rocks, typicil stands of this association appear in serpentine areas of Mt. Shiraga and

42

Mt. Higashi-akaishi. Similar forests, sud! as the Tfiwia sほ?idishii iorest on Mt. Shibutsu

and the Thuiopsis dolabratavar. hondai forest on Mt. Hayachine, occur on serpentine

in Honshu.

 (4)Tfiuj opsidetiandolabratae.nom. prov. The forest dominated by "Thujopsisdola-

braはoccurs on serpentine in Tokushima Prefecture. Though the occurrence of this forest

is not restricted to serpentine, it is a remarkable fact that such a community appears in

the warm・temperate forest region.

 ・(5) Associations oi Spiraea. Communities dominated by S piraea M四面and S piraea

sikohualpinaoccur on serpentine in Tokushima and Ehime Prefectures. These com-

munities are generally found in rocky places, on southern sides of mountains, etc. ex-

posed to the intense light.

 (6) There are grassy places mainly dominated by gramineous plants, such as

ArU・ndinaria pygmaea, Miscanthus sinensis,Arundinella hirta, etc. Occurrences of

such communities are mostly due to biotic factors, especially to human agencies.

 Characteristics of the floristic composition of serpentine vegetation in Shikoku are

similar to the general characters of serpentine flora mentioned by Rune, except for the

rather rich flora. The serpentine flora of Shikoku is not so poor in either species or in indivi-

duals, being rich in relics, narrowly endemics, and disjunctive elements. Families and

genera comprising characteristic and dominant species are listed on p. 27. Considering

endemic, disjunctive, and other noteworthy plants, an account of floristic characteristics

of the serpentine vegetation is given・ on pp. 26―34。

 Serpentinomorphoses are recognizable in a number of plants. Especially, stenophyllism.

glabrescence, nanism, a greater development of root-system. and the tendency to become

purplish in colour are characteristic in the serpentine flora of Shikoku. Some of these

serpentinomorphoses, such as nanism and the greatly developed root・system, which are

adaptive characters seem to be xeromorphoses, since the same morphological changes

of plants appear in dry places, such as limestone outcrops, rocky river・banks, etc.

 The physiognomy of serpentine vegetation is researched with reference to the vege-

tation types. Canopy types of serpentine vegetation are represented by the Maquis-,

Pintis-, and Chamaeりparts-type. In open pine forests, the forest floor is characterized

by the Sasa-, Miseanthtふs・, Careエー, Hy pniati・, or Cladonia-Vype. However, in the

Cflan・laecyparis-lype forest with closed canopies, the Khododcndron-, Repens-, 〇r Hy-

locoiniuiルtype are characteristic in the forest-floor.

 The Maquis-, Pinus。and Ch。laeりpar・is-typecommunities on serpentine are con-

sidered as edaphic climaxes, though the Pinu.s-typeforest appears to be an initial stage

in its physiognomy and floristic composition. An interpretation as regards to the climax

and succession is given on p. 37.

 It is generally noticed in Shikoku that climax communities in serpentine areas appear

to be characterized by coniferous forests or Maquis-type scrubs, and the similarity to the

vegetation on serpentine in other parts of Japan is presented and discussed on pp. 37-39.

 The serpentine vegetation varies with the different parts of the world, the variation

being due to geographical and climatic conditions. However,・it・will be stated that edaphic

factors have causal relations to the remarkable plant life on serpentine. As alreaby dis-

                  「cussed by many authors, chemical properties of serpentine soils are responsible for the

日ora and vegetation on serpentine. On the other hand, it should be emphasized that

other environmental factors, such as , microclimate, topography, and physical properties

of rocks and soils, also tend tて)・produce the distinctiveness of the flora and vegetation on

serpentine. Finally, it may be stated that the low competition in the serpentine ecosystem

is to be stressed in connection with the problems of distribution and ecology, of plants.

References

Braun-Blanquet, J., 19S1 : Pflanzensoziologie.

Coombe, D. E. & L. C. Frost, 1956 : The heaths of the Cornish serpentine. J. Ecoi. 44, 229-

 256.

Hara, H., 1934-1938 : Preliminary report on the flora of southern Hidaka, Hokkぷdo(Yezo).

 Bot. Mag. Tokyo 48-52.                  `

& M. Mizushima, 1954 : Vegetation of the Ozegahara Moor and its surrounding districts,

 central Japan. Sci. Res. Ozegahara Moor 401 -427.

1954 : List of vascular plants of the Ozegahara Moor and its surrounding dis

 tricts. Ibid. 428-479.

Hattori, S. , 1955 : Hepaticae of Hokkaido. I. On the Hepaticae・of southern Hidaka with special

 reference to the Hepaticae occurring on serpentine on Mt. Apoi. J. Hattori Bot. Lab. No. 15,

 75-92.

1955 : Hepaticae occurring on serpenti‘neon Mt. Apoi (Hokkaid6). Bot. Mag. Tokyo 68,

1957 : Hepatic戸eof Hayachine Mountain with special reference to Hepaticae ocとurring

1952:SPifaeaof Japan and its neighbourhood. Ibid. 14, 149-159 (In Japanese).

1952 : Serpentine flora of Mt. Shibutsu, Prov. Kodzuke, Japan. Ibid∠14, 174-176 (In

19S2 : Serpentine flora of Mt. Hayachine, Prov. Rikuchu, Japan. Ibid. 14, 177-180 (In

1956 : Serpentine flora of Mt. Apoi, Prov. Hidaka, Hokkaido.・Ibid. 16, 143-148.

1957 :Serpentine flora of Nupromapporo Valley, a branch ot Teshio River, Teshio。

& Y. Moraotani, 1952 : Serpentine flora of Island Sugashima, Prov. Shima, Japan. Ibid.

& G. Murata, 19S2 : Serpentine flora of Prov. Awa, Shikoku, Japan. Ibid. 14, 120-122

320-323.

 on serpentine rocks. J. Hattori Bot. Lab. No. 18, 106-118.

Hatusima, S., 1948 : On the distribution o{ QueYcus仙郷iraeoidcsA. Gray in Japan. Ec�. Rev?

 11, 101-106 (In Japanese).

Hayashi, Y., 1951-1954 : The natural distribution・ of important trees, indigenous to Japan.

 Conifers report 1-3. Bull. Gov.For. Exp. St. No. 48, 1-240 ;.No. 55, 1-251 ; No. 75, 1-173.

Honda, M. & Y. Takenaka, 1930 : Natural monuments of Iwate Prefecture. Tennen-kinembutsu

 Ch6sa H6koku (Shokubutsu) No. 10, 1-10 (In Japanese).・        / :

Horikoshi匹Y・, 1937 : Geology and petrology of the vicinity of Bessi,' Ehime Prefecture (Prelimi-

 nary report). J. Geol. Soc Jap. 44, 121-140.                ,・・

Kitamura, S., 1950:Adaptation arid isolation on the serpentine areas. Act. Phytotax. Geobot.

 12, 178-185 (In Japanese).

Japanese)

Japanese).

Hokkaido. Ibid. 17, 41-45.

14, 118-119.

, M. Hiroe, & G. Nakai, 1950 : Serpentine flora near Sekinomiya, Prov. Tazima. Ibid.

 12, 185-187 (In‘Japanese).

, G. Murata, & K. Torii, 1953 : Serpentine flora of Mikawa Province. Ibid. 15, 1-3

 (lnJapanese).                      ●..

of the' K6chi University Vol. 8, No. 10

Tokushima

K6chi Regional Forestry Office, 1939 : Report on the forest vegetation in the district under the

  jurisdictionof K6chi Regional Forestry Office(lnJapanese).

Krause, W., 1958 : Andere Bodenspezialisten. Handb. der Pfl. -physiol. Bd. 4, 755-806.

& W. Liidwig, 1957 : Zur Kenntnis der Flora und Vegetation auf Serpentinstandorten

1954 : The ecology of serpentine soils. III. Plant species in relation to serpentine soils.

1927 : -

1954 :

II. Das Problem der Serpentinpflanzen, eine Kritische 6ko】ogische

(II) Bull. Fac. Educ. Kobe Univ. 9, 7-31.

 des Balkans. 2. Pflanzer】gesel】schaftenund Standorte in Gostavic-Gebiet (Bosnien). F】ora 145,

 78-131.

Kruckeberg,A. R., 1951 : Intraspecific variabilityin the response of certain native plant species

 to serpentine soil. Amer. J. Bot 38, 408-419.

 Ecol. 35, 267-274.

Lammermayr, L。 1926 : Ma terialien zur Systematik und Okologie der Serpentinflora. I. Neue

 Beitrage zur Kenntnis der steirischen Serpentinflora. Sitz. Akad. Wiss. Math. -Nat. Kl. Abt

 1, 135, 369-407.                            ‘

Studie. Ibid. 136, 2S-69.

1928 : Weitere Beitrage zur Flora der Magnesit- und Serpentinboden. Ibid. 137, 55-59.

1928 : Vierter Beitrag zur Okologie der Flora auf Serpentin- und Magnesitboden. Ibid.

 137,・825-859:

Maeda, T., 1951 : Sociological study of Chamaecyparisobtusa forest and its Japan-sea elements.

 Misc. Inform. Tokyo Univ. Forests N0. 8, 21-44.

Miyazaki, S.,1937 : Sylvicultural consideration on the 7加加戸sis dolat?rata forest in

 Prefecture. K6chi Rin-yu N0.206, 1-23 (lnJapanese).

1942 : On the relation between the forest vegetation and the soil morphology of Shikoku

1942 : Outline of the flora of Nishikiyama. Ibid. No. 19, 37-42 (In Japanese).

1954 : Notes on 哨e flora of the Gaspe Peninsula. Sv. Bot. Tidskr. 48, 117-136.

1954 : L'Alliance du Shiion Sieboldi. Vegetatio S-6, 361-372.

& K. Hatiya, 1951 : The forest vegetation of Izu Peninsula. Bull. Tokyo Univ. Forests

 ・(lnJapanese).

MorishitaヽK. & T. Yamanaka, 1956 :I Distributional and ecological observations of SfiJ’aea

 tosoensisYatabe. Jap. J. Ecol- 6, 50-53.

Nakai, T., 1930 : Vegetation of Mt. Apoi in the Province of Hidaka, Hokkaido. Tennen-kinem-

 butsu Ch6sa Hbkoku (Shokubutsu) No. 12, 1-80 (In Japanese).

Nakano, H. , 1942 : The deciduous forest vegetation in Japan. Ann. Phytoecol. 2, 57-72 (In

 Japanese).

Nishida, S., 1918-1919 : On the distribution of plants in the Yupari mountain range. Trans.

・ Sapporo Nat. Hist. Soc. 7, 71-92, 136-177 (In Japanese).

Novak, F. A., 1928 : Quelques re marques relatives au probleme de la vegetation sur les

 terrains serpentiniques. Preslia 4, 42-71..

Ogawa, Y., 1950 : Serpentine flora of Mt. Kurosawa, Prov. Kii. Act. Phytotaχ. Geobot. 12, 187

 -188 (In Japanese).

Ohwi, J., 1933 : Flora of Japan (ln Japanese).

Pichi-Sermolli, R., 1948 : Flore e vegetazione delle serpentine e delle altre ofiliti dell'alta valle

 del Tevere (Toscana). Webbia 6,1-380.

Raunkiaer, C., 1934 : The life forms of plants and statistical plant geography. `・

Rune, O., 1953 : Plant life on serpentines and related rocks in the North of Sweden. Act. Phyto-

 geogr. Suecica 31,1-139.

Sakamoto, C., 1955 : Studies on utilization of serpentine. I. On the treating of calcined ser-

 pentine by sulphuric acid. Res. Rep. K6chi Univ. 4(48), 1-8.

Sato, W., 1946 : Studies on the plant-climate in the south-western half of Japan. I-I!. J. Sci.

 Kanazawa Normal Coll. 1, 1-59, 61-72 (In Japanese).

Sawamura, T., 1956 : Serpentine resource in the vicinity of K6chi city (lnJapanese).

Shimizu, T., 1953 : On the noteworthy plants from the limestone range in Shimohei-gun, Pref.

 Iwate. Japan. 1 ―3. Act. Phytotaχ. Geobot. 17, 85-93, 107-113, 143-154.

Suzuki, T., 1953 : The forest climaχes in East Asia. Jap. J. Bot. 14, 1一12.

一一19S4 : Forest and bog vegetation within the Ozegahara Basin. Sci. Res. Ozegahara Moor

 205-268.

(T・yA・manaka) 45:

39, 145-169.

&M. Kitag耳wa、 1954 : Sur la vegetation et l'ecologie.du pinaies de、PttlUS(ienstμOfa.

Akamatsu・ ni kansuru Kenkyu Rombun-shu 163こ-168.

& H. Usui, 1953 : The secondary forest vegetation of North Kant6 District. ]. Jap.

1928 : The vegetation of Mt. Apoi, Prov. Hidaka, Hokkaido。Japan. Ibid. 5, 49-134.

1932 : A list of plants collected in the Teshio University‘Experimental Forest. II. Ibid.

1935 : The distribution of ?i。gs戸umilain northern Japan. Ecol. Rev. 1, 23-36 fin

1938 : Rhytogeographical study on the alpine plants of Yezo. (I). Ibid. 4, 101-112.

1944 : Phytosocio】ogical study of Picea glehni forest. Res. Bull. Coll. Exp. For. Hokkaido

& T. Morimoto, 1933 : Report on the ecological survey of the Teshio Experimental Forest,

1954 IV. The vegetational response to serpentine soils. Ibid. 35, 275-288.

1950 : Studies on the serpentine vegetation. I-II.、Seibutsugakkaishi of Hiroshima

1951 : Sociological studidS on the serpention vegetation. 1. Bull. Fac. Educ. K6chi

1952 II. Ibid. No. 2, 59-74

1952 : Studies in the vegetation and the flora on serpentine. Res. Rep. K6chi Univ. 1

1952 : Flora and vegetation of Mts. Akaishi, Shikoku. Act. Phytotax. Geobot. 14,

1952 : Some problems on the f】oraand the vegetation of、theserpentine region. Ibid. 14、

1953 : Serpentine vegetation of・Aburaishi in Okusuue, Prov. Tosa, Shikoku, Japan.

1953 : Serpentine vegetation of Tomisato, Prov. Iyo, Shikoku, Japan. Ibid. IS, 91-94

1953 : Serpentine vegetation of Hashiradani and Harihara in K6chi city, Shikoku, Japan.

1954 : Sociological studies on the serpentine vegetation. Ⅲ. The vegetation on Mt.

19S5 : Studies on the limestone vegetation in Shikoku, Japan. Res. Rep. K6chi Univ

1955 : Sociological studies on the serpentine vegetation. IV. The vegetation on Mt

1955 : Serpentine vegetation of Mt. Yokogura. Act. Phytotax. Geobot. 16, .43 (In

 For. Soc. 3S, 9-13.                                    ,.

Tadros, T. M., 1957 : Evidence of the presence of an edapho-biotic factor in the problem of ser-

 pentine tolerance. Ecol. 38, 14-22.                       ,

Taniguchi, M., 1958 : Phytosociological study of serpentine area of Mt. Asama, Mie Pref. Act.

 Phytotaχ. Geobot. 17, 122-!27.

Tatewaki, M., 1928 : A list of plants collected in the Teshio University Experimental Forest.

 I. Res. Bull. Coll. Eχp. For. Hokkaido Univ. S, 1-48 (In Japanese).

7, 181-208 (In Japanese).

Japanese).

Univ. 13 (2), 1-181 (In Japanese).

 Hokkaido. II. On the synecological survey of the forest of Picea glehni. Ibid. 8, 1-287.

Toyokuni, H., 1955-1958 : On the ultrabasicosaχicolous flora of Hokkaido, Japan. (l)-(7).

 Hokuriku J. Rot. 4, 97-101; 5,12-15, 81-84, 115-116; 6,17-20,・63-67; 7,37-38.

Usui, H. , 1954 : The plant communities and their successional relationship on the・vegetation

 of the Utsunomiya University Forest. Bull. Coll. Agr. Utsunomiya Univ. 2, 169-176. ・

Walker, R. B., 1954 : The ecology of serpentine soils. II. Factors affecting plant growth on

 serpentine soils. Ecol. 35, 259-265.                               。

     ,H. M. Walker, & P. R. Ashworth, 1955 : Calcium-magnesium nutrition with special

 reference to serpentine soils. Pi. Physiol. 30, 214-221.

Whittaker, R. H., 1954 : The ecology of serpentine soils. I. Introduction. Ecol. 35, 258-259.

Yamanaka, T., 1950 : The serpentine vegetation of Nishikiyama. Act, Phytotax. Geobot. 12,

 188-192 (In Japanese).

Univ. 2 (2), 23-24, 25-26.

Univ. No. 1, 95-117.

(9), 1-8.

129-132 (In Japanese).

195-197 (In Japanese).

Ibid. 15, 61-63 (In Japanese).

(ln Japanese).

Ibid. IS, 95-96 (In Japanese).

Shiraga, K6chi Pref. Bull. Fac. Educ. K6chi Univ. No. 5, 47-53.

4 (2), 1-12.

Higashi・akaishi, Ehime Pref. Bull. Fac. Educ. K6chi Univ. No. 8, 49-57.

砧   ’

 Japanese).

Research of the k6chi University Vol. 8、No、10

1956 : Sociological studies on the serpentine vegetation. V. Namekawa and Yusuhara

1957 : On the Ghamaecs{>orisObtusa forest in Shikoku, southern Japan. Jap. J. Ecol. 6,

1957 : The forest vegetation of the Inland Sea District of Shikoku, Japan. Res. Rep.

1958 : Serpentine flora of Mt. Nishi-akaishi, Shikoku. Act. Phytotax. Geobot. 17, 94-96

1958 : Serpentine flora of the foot of Mt. D6go, Hiroshima Pref. Journ. Jap. Bot. 33,

1953 : Sociological studies on the serpentine vegetation. VI. On the communities of

1959:The Pirusd6竹sげlofaforest on serpentine. Jap. J. Ecol. 9, 54-58.

1959 : Sociological studies on the serpentine vegetation. VII. General considerations on

1959 : Serpentine vegetation of Yawatahama and Mikame in Ehime Prefecture, Shikoku.

& K. Takezaki, 1959 : Distribution and ecology of j?hododendron ri加〃ssMakino, with

1935 : The occurrence of Thujol)stsdolabraia andother peculiar plants in the basin of

193S : Further notes on the occurrence of nzφ・Psis加labrata. Ibid. No. 195, 7-13 (In

1940 : Serpentine plants. Ibid. Nc . 233, 1-4 (In Japanese).

1949 : Sociological studies on the pine forests in Japan, especially with regard to their

1953 : Ecoloc!ical studies of Japanese pine forests (ln Japanese).

in K6chi Prefecture. Bull. Fac. Educ. K6chi Univ. No. 9, 35-39.

149-152.

K6chi Univ. 6 (6), 1-16.

(lnJapanese).

120-121.

 SPlfaeain Tokushima and Ehime Pref. Bull. Fac. Educ. Kbchi Univ. No. 10, 71-76.

一一― 1958 : The Quefcus仙郷ii'aeoidcscommunity in Shikoku, Japan. Res. Rep. K6chi Univ

 7 (9), 1-6.

the ssrpentine vegetation in Shikoku. Bull. Fac. Educ K6chi Univ. No. 11, 87-104

Act. Phytotax. Geobot. 18, 29-32 (In Japanese).

 reference to the vegetation and flora on rocky river・banks. Journ. ]ap. Bot. 34, 215-224.

Yamawaki, T., 1957 : Studies on the distribution of the Genus Hemerocaliisand its natural

 variation. Kusunoki N6h6 11 (8),・12-26.

Yoshimura, I., 1958 : Ecological observations on Cladonia, a genus of the Lichen, in Shikoku.

 Gakugei No. 1, 44-51 (In Japanese).

Yoshinaga, T., 1914 : The relation between serpentine and peculiar plants in Prov. Tosa. Bot.

 Mag. Tokyo 28, 40-42 (lnJapanese).

the River Naka, Tokushima Prefecture. K6chi Rin・yu N0.184, 1-10 (In Japanese).

Japanese).

Yoshioka, K., 1948 : On the types and development of Japanese pine forests. Ecol. Rev. 11,

 204-216 (In Japanese).

structure and development. Sci. Rep. Tohoku Univ. Ser. 4, 18, 229-242.

                   E xplanation of Plates

                        Plate l

1. The serpentine Outcrop of Korodokihana directly facing the sea. The Quercus仙郷ii'aeoifies

 scrub (Pitiosponto-QuBtceium ^hylHraeoirieiis)occur in rather stable places. (Oct., 1958)

2. An open and stunted pine forest G4かぷd∂・Pinetum心nsiflびae) at an altitude of 900 m. in

 Shimagawa一一                                    (Oct., 1958)

3. The serpentine barren in Shimagawa. The vegetation of this area is essentially characterized

 by the pine forest.                               (May, 1956)

4.A view of the stunted iJine forest (AbelieioヽPitieturadensiμorae)atthe 280 m. altitude in

 the vicinity of Nishikiyama.                          (Nov., 1956)

                      ‘ Plate II

5.A stand of the pine forest (じAhelieto-Pinetum fiensiルrce)at an altitude of 370 m. in the

 vicinity of Rngy6ji. Barrens occur within the pine forest.          (Nov., 1958)

6. The serpentine area at an altitude of 380 m. in the vicinity of Engybji. Pinus deiisi/1びa

 is abundantly found.                               (Nov., 195S)

7. The open pine forest (Abelieto-Pinetu回心s切印・ae) in altitudes

 the Osaka Pass.

between 40 and 80 m. 0f

     (March, 1951)

8. The serpentine area of the Osaka Pass covered with Afundinaria夕ygmaea. Scatteredstun tec!

trees are戸畑μsder (March, 1951)

                         Plate 曹

 9. The pine forest (.Abeμgl∂一月netu。zjのisiflθrae) at an altitude of 60 m. of Okbyama. The

 shrub stratum is dominated by Afundinaria pyg竹^aea.              (Oct., 1950)

10. A view of Mt. Nishi-akaishi (background) from the western place of Mt. Higashi-akaishi

 (foreground). The south-exposed serpentine area of Mt. Higashi・akaishi is characterized by

 the stunted coniferous forest. The serpentine outcrop of Mt. Nishi・akaishi is shown by an

 arrow。                                                (Oct., 1957)

11. The southern side near the summit of Mt. Higashi-akaishi. The coniferous forest is very

 stunted, and the scrub of S1)iraeasikokuaゆina(SPi^aeeiumsikokuaゆ・inae) occurs here。

                                              (Aug.. 1955)

12. An inside view of the coniferous forest (Rhodoreto-Chamaecybaridetumobtusae)at an altitude

 of 1200 m. 0n the northern side of Mt. Higashi・akaishi.         (Aug., 1950)

                         Plate Ⅳ

13. The northern side of Mt. Higashi・akaishi covered with the coniferous forest. In the eastern

 part of this mountain, serpentine forms a large scree.            (Aug., 1956)

14. A view of the east ridge of Mt. Higashi・akaishi from the summit.   (Aug., 1956)

15. The summit of Mt. Higashi-akaishi seen from the east ridge.     (Oct., 1957)

16・ The summit of Mt. Shiraga. Coniferous trees, such as Chamaecipafis obtttso.Piims peniapkylla

 var.himekomaisu, and TsMgafiiuersげolia,aでe stunted.          (June, 1958)

                         Plate V

17. The southern side near the summit of Mt. Shiraga. The vegetation is characterized by the

 stunted coniferous forest with many withered trees.              (Jnne, 1958)

18. A typical stand of the coniferous forest (j?加doreto-Chamaeりか・ridetutn obtusae) at an altitude

 of 1200 m. on the northern side of Mt. Shiraga.             (March, 1955)

19. Spiずaeasikokuaゆi丿la Koidz. with flowers in the rocky place near the summit of Mt. Shiraga。

                                              (June, 1958)

20. ShrubbyTftuiopsis dolabrataSieb. et Zucc. in the serpentine area of Jinry6. (July, 1951)

                         Plate M

21.

22.

23.

24.

Blooming C∂がopsis spicata Sieb. et Zucc. in the vicinity of Nishikiyama. (Apr., 1957)

Coり!oftsisspicaiaSieb. et Zucc. with young fruits and leaves at the Osaka Pass.

   ゛゛                                   (Apr., 1953)

CoりloOsis gotoa皿Makino with leaves at Jinryb.

Hete・y砂appushtspidusssp. 1ゆiodadusKitam. with flowers at the Osaka

25. Saitssぼea nipponicassp.タoshinagae■K\tam. with buds at the Ryu Pass

                         Plate Ⅶ

25.

27.

28.

23.

30.

  (July, 1951)

Pass.

  (Oct., 1950)

  (July, 1958)

£iliw.り1加;り㎝iaりzvar. al・gasりIKitam. from the serpantine area of Sakashu. (May, 1S57)

Gymnoccぴ岫4ヽmjessoenseKoidz. in the serpentine crevice of Mt. Higashi -akaishi.

Clematis戸atens M、)rr.et Decaisn. with flowers at the Osaka Pass.

Ange!ica ubatake・〃jsKitagawa at the summit of Mt. Higashi-akaishi.

Cα・mtitosofussibricitsRupr. in the serpentine crevice of Tomisato.

(July,

(Apr.,

(July,

(Apr.,

1953)

1957)

1953)

1953)

(Received August 28, 1959)

Table 15.

Association table of

Abelieto-Pinetum densiflorae in Yusuhara district

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                                                 WWW WWW W   W  WWW 心ヘノ      心鴫L

                                                                   +     +     s ,、Jト., ト..十十。十十。む。。     。

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                                -9-9uu-u-u4nS乱丿高ぶ哀咽賢 SJ                                、、●。。、● 、。,、、.,。。。。。。。。。。。。。。。。。、●。。。。。。。。。   。。。

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・●・一一≒≒4≒=<1=(M≒!ilrrぶ4≒ !^^^.=s;一一・μ一泡4面一・4≒SIr491     M≒≒≒ Mess-・=1砂=4sョ-4一・i一≒四面g-・|り4・isS==を・・1両両岸MiM面細   細細M Constancy

*cf.  Fig.

 1.

叫a-

S‘

Association table of  Abelieto-Pinetum densiflorae in'K6chi

district

4タや

⑤   甫   サ碍年

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・・・・・・・・・・・・・・.‥十十‥゜十‥‥十‥十‥’‥‥’-一一゜    ♀‥♀‥‥゛‥‘‥早‥‥・早・五万‥‥早゜早早ヅ ≫                                                      十 士卜叫J    ドj NJ K) り トーJトーJ りωωω 4     り

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頑§・

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                                                                 ●                              ←J

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吻011

--●≒≒≒=0==4『同一i一一≒細細同一味=;g-・i≒凹地1≒『-i同≒遡一一・4細―μヒiョ1≒同s≒gべMiig     細田il!1ぶ自‐『同一ぶ-qiμぶ細≒μS4向哨べ4ぶぶぶ同べ同哨べ4M   『≒M Constancy

*cf.

 Fig.

 1.

Table 17

yS0ciation table of Rhodoreto-Chamaecyparidetum obtusae on Mt.

  Higashi-akaishi

a       -●・・●a ●                 a    、        □ 、           乙;●

           ■≫           a

爪甲

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Plate VI

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Plate Ⅶ