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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
2
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.
1
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―!・+
3
3
2
ト
●
1
●・・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
●●●
1
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
m
言)
混)
づコ
4ヤ)
呂
皆
皆
1(十)漕)
竹
本
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
十
十
十
●
●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
6
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
+ + +
顕
3(十)示)ブ
憐
川
寸)
-j本
本
-1i
本
t
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●
1
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}
才
竹
工
2こ)
ドミ
2(十)
士
汗
j
(2)
1
1(十)
2 2
2
十+
3
(1)
1(十)
2(5)
1
1(十)
1
1
十
(4)
Flj
親
子
1ダ
(十)
ズ
]E.
才
于
3
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
ト
μ
+ (十) 1
3
りり
刀十十
ぐぐぐ
1
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
1
十十
+
+
2(十)1
十(十)
十(十)●・・sし0・・..
211十十11・十・・・十十
5
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+―
1
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
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
2
+.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
昌
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
1 (十)+
●●
2(十)
+
22十―十十―・・・
凰
{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
(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)
川
2(十)1(十)川
幅)
2(十)
1じ)
1(十)
川
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
2
1
●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.
2
2
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
昌
`;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
?
帽
十
プ
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
2
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
で
/3'
ご
・,.+
・ ● 」
^li:
j5
・(十)
+
5
1
(十)
(十)
1(十)
(十)
+
+
5
十
゛2
十
1 1
、●
ヤ)
仰
1
●+.
● 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
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)
j
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)
3
i
ヤ)
1(十)
ケ)
1
?、 じ)
1(4)
j
詣
3ツ
サ)
+(十)
1.
+.
(?)
+.
j
本
1 臨
●
ヤ)
才
六
+.
●十 ● ●
●ツ)
● ● ● ● ●
1
3
⊃
● ● ● ●
1
2(1)
3
●
4(十)
●+
●
● 1
●
●
●
● (十)
●
● (3)
●
●
●
●
●
●
+
3
2
+
1
●
●
●
●
(十)
1(D
3
5(十.)
1 `
1
1
十
(3)
十
3
2
2
争
2
● 5ツ
●
‰)
7
●
● ソ)
● ソ)
●
●句)
● ● ● ● ●
1
5
ふ
● ● ● ●
T)
九
十(十)ン
1:
十(D六)
ヤ
1
3(十)
1.
阿)
づお
1.
●
十
(十)
●
●
●よ)
●
●
●
●
●
●
1
2
●
●
●
●
●
●
1(D
5(十)
2
●
5(D
●
● (干)
●
● (十)
1(十)
●
●
●
●
●十(1)
●
●
●
●
●
●
5
3
2
●
●
十
●
●
●
疹
2.
句)
他
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
Ⅲ
ⅢIII
IIl
lI
V
V
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
十
十
●
●
●、
●
●
3
2
●
● 1
1
●
●+
●
●
●
●
2
1
2
+ 1
●
●
1
+
●
十
十
十
●
●
1
●+
+
●+
十
十
十
●
●
●
● ●
5
1
十
十I +
? 十
2
4
2
1
1
十
十一
1
十
1
十
十
十
十
十
十
5
十
十
十
2 2
十
十.
十
1
++
十
十十
.●、
5
2
●
●
●
●+
●
●
●十
●
●
2
2
1+
1
十 1
●
●
●
+
+
+
●
●
十
十
●
+
●
●+
●
●
●+
●
● ●
5
1
1
+
.+
3
1
3
1
P
+
1
1
+
+
+
+
十 十
十
十
十
4
1
1
1
●
●
●
●+
+
●
●
●
3
十 2
● 1
●
●
●+
●
+
●
●●
●
●十
●
+
●+
●十
十
●
●
●
●
●
4・
1
11
●’
321
12
十十
1
+
十
十
十 1
+
+
+
十
十
5
2
1
十
.●・ ・
+
5
1
1
+
十
十
十
十
十
十
十・
+
7500
902
203
102
, 52
51
2
2
2
2
1
1
1
3250
1601
1151
378
350
252
228
202
179
150
108
105
105
102
102
57
55
54
7
5
5
5
4
2
1
1
1
1
1
χ
χ
Ⅶ
Ⅳ
Ⅲ
」『
I
I
I
I
I
I
I
χ
Ⅸ
χ
W’
可
Ⅶ
V
M
V
Ⅲ
χ
Ⅶ
Ⅶ
’Ⅳ
Ⅳ
可
M
V
Ⅶ
V
V
V
Ⅳ
I
I
I
I
I
I
* 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
X
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
I
ⅦI
I
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
Ⅳ
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
X
lⅣ
VⅣⅣ
l�
Ⅶ
Ⅲ'
¥ⅣⅣ
Ⅶ
】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
I
llIITI
I
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.
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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
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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.
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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.
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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.
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・(lnJapanese).
MorishitaヽK. & T. Yamanaka, 1956 :I Distributional and ecological observations of SfiJ’aea
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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-
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Sato, W., 1946 : Studies on the plant-climate in the south-western half of Japan. I-I!. J. Sci.
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Sawamura, T., 1956 : Serpentine resource in the vicinity of K6chi city (lnJapanese).
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Iwate. Japan. 1 ―3. Act. Phytotaχ. Geobot. 17, 85-93, 107-113, 143-154.
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(T・yA・manaka) 45:
39, 145-169.
&M. Kitag耳wa、 1954 : Sur la vegetation et l'ecologie.du pinaies de、PttlUS(ienstμOfa.
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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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Association table of Abelieto-Pinetum densiflorae in'K6chi
district
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⑤ 甫 サ碍年
‥+;+・+++‥+‥‥‥‥‥‥+‥+‥‥‥ヽ・+-・・+-‘ ‥♀♀‥‥‥‥モシ・・べ Wtノ W W WWW 心 WW W 心
OC/〕1呂呂ぢ(りヨ
゜’十十十゜・十十十゜十十’゜゜゛十’’゜’’十゜十十’゜・ tsj・ ・十゜’‥十一’ ‥’十゜万万‥‥万年‥年‥♀♀‥づ年・モjづ年ヅ J。コヴ
+・+・+・・++ト,・十十s・・十十・・十・・+・e、。・・・・ω・・4.ω・・十ト.ト, …‥ト…⑤・皆皆緋・いづ・回≒ ./
W '--^'■'^ W WWWW 心 W 心
小・
‥‥‥‥o‥・+‥ヶ-・‥・+‥・+‥‥‥‥ヽ・・ヽ・・・ヽJヽj ドレ
♀‥‥・矢竹‥万言Eドド・問い回詣 且勺
心づノ 心ハ.八-ハ、ノ yノ y八ノ χ、ノ 心ノ 心いノ
C/つro -0
enOl OOCO
M
‥‥4’‥o‥-ooo‥o‥oo‥・++‥,.・-・一十 ・≒縦‥‥・三回‥ドド・回ド竺か回・竺 Ξ・皐
Wtノ 心/ 心ベノ 心ノ 心ヘノ 心
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りトーJα)o o ooo t司
十 十十仙十十ト..s礼トー,ω9十ωu 肖、j・・・・・・・・・・・・十・・・・・・・・・・・・・-゜, ,・ ・ ・ ・゜・ヽJ ・ M+ Ca)+ ‥Ξ’♀‥ ゜♀/-\/'~\/'~\/'~\/'~s゜♀ワ♀孚号♀・♀♀♀゜Ξ♀Ξ号 ♀
心/ WW心心心心 心/ 心/心ヘノ 心/ WWχ、/ W 心
呂(/っ呂。
+ ++~+ s八一トo一J-ωωト一十けω・・・・・十十・・・・・・+・十十十・・+・十十十トー+・s・・+・・。ω・トーω,、jω ‥♀常‥゜♀’♀♀ΥΥ ・・ + ・++♀♀♀ ♀常常゜♀♀ΞΞ ‥言
yハノ 心 心心心心/ ‘W WWJWW <:^<-y\^ 心心
S(/)
O OCDgra
’゛’十’゜’十’’゜’レ゜十’゜゜十゛゜十I―' I―'I―<゙ 十・ ・ ・ ・ .・`-J°’’’゜,・.゜ ‥モE°?'・・・・・.十‘万作‥1レモン万作°ミj’十‥早`モj早早{1 to
巾・●
・・・+・・・・・・・・・・・十十十十十・s十十十・・・・l.゜o‥‥・一・ ‥‥‘‥‥‥‥べE・・・・白で学卒.ッ・・学・苦学9合 八でづ
心 WWJW 心 心
・|斟
・・・・・・・・・’‥・十‥’十一‥一十十‥十‥ヽ゛十十‥‥・ヽ・ヽ.’ ‥♀‥‥・・・・・・・・・.’・モF旱べ 巻 十 ドーJ NJto M 十 トーJ トー-Jω ω Q
●・●・●・・●●●●●・・●●●●ト_j●・トーJF一J+トj・+●●}-J・MtQ・・●●・ωK)・ ●・♀♀・・・・・●・・・♀・・●゛Ξ♀・平●・♀・♀゜♀平谷♀ Ξ゜谷
ヽノ-●・ ヽノ ヽ一一ヽノ \^"-^ \^--^ヽノ巾・・
・・・・・・・・・・・・・・.‥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
・+・・・・・・+・・・・+・・・・・+トーJ・トJトーJトJ・トーJ・+・・ドー゛N〉・・・・>-'NJl-'CO ・・♀♀・・・ + ・ ++‘♀‥♀’♀♀’Ξ゛♀♀’♀♀Ξ♀゛尽 ‥Ξ
心ヘノ 心ノ 心八ノ W 心/ Wtノ 心八ノ kノ心 yノ
小
・ヽ・
‥‥oo‥o‥‥-ト・++・++‥++・+・+‥-‥‥一一 皆・♀孚・ト‥プ斗‥仁亜斜勺市・か悶鸚 ・・{1
W心 WW 心/ 心 心 W 心心 心/ W
ット
゜十’゜’十十・・・・・・・十十゜十゜十゜’十1-^1-. ・ I--・ ゜゜・ i^to ・ ・ ・ ・十十,.゜ ‥白と‥勺S゛‥十ブI―'h-f I―'NJ‘十刎学’苦学回答 ‥lj
小
・ヽ・
++ + s.,+ トー ++ 十十ト,+。+e、J ,。+,、,。ω ト一
.・・・・・・・・・・・・.;ト.十十・・・・・+・・+・・・・・・・・+・,。ト.・・ ♀常Ξ+ + + +゜‥常♀ ・T♀゛ΞΥ常常常 ♀゛常゜♀♀ ♀ ‥Ξ
. y八ノ WW 心心 ^.・Sw' 心ヘノ WWJW W 心ノ Wkノ 心/
巾ご
--づ卜 トー ト一トー十ト●HH 十 ド一 to CO -'COCOto Q・・・・・・●・+・・・・・+・・・・・トーJ・・・・・・・+s・・・・・・。・・4+ ・・・・・・Ξ‥♀゛孚♀゜+ + + +Ξ‥♀゜♀♀孚Ξ♀♀゜Ξ ‥Ξ
-----ヽ・- ヽノ--ヽ-ノ-・-
←゛M トーJ M to I―' トーJ NJ 十 十4ムQ ム 十 ム・・・・・・・・・゛゛゜・・・・+・・・・・・・・・・ω,。+・・・・・・・トー・,、34 ・・・・・・・・・孚♀・Ξ゜+ + +♀・・♀‘♀・孚言Ξ♀’Ξ ‘孚
--●・ ヽヽノこヽノ- --- ヽノ ヽノ巾・
+ + 。,..++ ト,s+s.,十 s●+s +ωs。 ω ω・・・・・・+・+・+・・・・++・・・・・・・・・s.・+・・+・・トー・・,、J,、j十. ‥孚♀‥゜ ゜♀♀♀♀・♀・♀♀♀♀・・+ + +++ +Ξ♀゜Ξ ‥苔
WW \-y'-.-y\-y\^ 心 K-y^-^^.^\^ W心ヘノWWW W勺胆
+ sトート.十十ト.js十トー+ 。仙十四肖。ω・・・・・・+・+・s・・.I・十十十・・・・・・・・十・+・・+・・s・・e、j。ト一ω ゜♀♀常‥常常・ ♀♀・♀・+ + + +・・常♀♀常♀+++・Ξ, ♀゛言
ヽノヽノこ ヽノご ヽノヽノ-一一 ヽノ-●・-●ヽノ ヽノ----ヽノヽノヽノ ヽノ
頑§・
‥‥‥+‥・・・・・++-‥‥‥‥‥-・,。・.・+・・・・・・・-・・ 孚・甘‥ド.¨づ‥‥・卦‥i・甘言悶悶岬 しがヽご
-ヽヽノ-● ヽ.・ - ヽヽノ ヽ.・-ヽノ-ヽノ -・-司レ
+ 肖ト。十十十sμ』-J to >―'^ to ω ω・・・・・・++・・・・・・。+・+・9・・+・・・+・+・・s+・・・・,。e、,。e、J ・・♀・・・♀・・♀・♀ ・孚・ Ξ・♀♀・♀孚ヽ‥♀♀号孚・言 Ξ・7
W W W W 心 WW 心ヘノ WW W J巾・
‥‥‥‥+‥‥++++申‥+‥‥・-‥‥‥‥‥-.+-. ・づ・モヤ‥‥岬・緋‥‥牡‥・託づ‥か竺 √(゛
心 心/ W WW 心J WW 心 心 心/巾・
++ +。 ωトー+ ,、j,。 ,、34Qω ω Q・・・・・・・・+・・・・・+・++・・+・・・・・・・+・・・・・,。・。・トー・トーω ・・・・・・・・・・常 ・♀♀‥Ξネ♀・・♀♀‥♀Ξ常常・言 ‥
心/ 心/心ノ W心/ 心心/ yノ k八ノ
巾・ ++ + + + + 。++s s,り りり ω ω り
・・・・・++++・・・・・+十十十・・・++・+・s・・・・+ト.・4・十十ωむ。十 ♀♀竿‥ ‥ワ’♀♀‥’言T♀?♀♀‥♀言‘Ξ゜号 ‥言
㎜ WWW 心 W心 WWJ K^'-.-y 心
さ稽・
ドー-り-Jトーj“ H-j 1―' to (o toω ω
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COtOtOCOtOCOtOCOlOft!>.Ji..4^ai<DCDC0ぱ)ぁ(Jlc⊃ocoNj>-oototDCncn.Ji.CnドーJ isj J^ Jik Ji. Kj C刀cJ】むlk、JONJtOtOCOib'COi―'I―'J^ムドーJりJ^ J^ ,& OONJCJ M4トJ
OOドーJトーり一一Jドーjωω400ドー-■ Csj COωふ4(乃○トーJQドー-■(ocjcooaioi―'ooドーJト■'oooooeoot―』 り叫^J CJ O O I-' lO OドーJK)4トーJω○○肖肖り払○トーjOωto o t―' oドー-^ o o o ooo
●● ● ● ●●● ●● ●●● ● ● ● ● ● ● ●●● ● ● ● ● ● ● ● ●● ●●● ● ● ● ●● ● ● ● ●●●●●●●・● ●● ● 畢 ●● ●●● ●●● ●
ムQ)りり(乃○りto CD tti. J^ふり り*>■ CO CO C乃J^ MOD O) Ji. lO J^ 03 C乃(コl心仏ム 00 ifc. *■ CT> QC刀心(71 CO Ol*. to 4^ ifc. cnc刀ife. ji. a>りCO ≫K CO CDムり心41Q IQ4ふ ` 4
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● ←J
トJ H H ドーJム Ok、】 Q
Q CO Q り 4Q ω(乃^ to 1―'[sj りぁり00 t!)MトーJ(χ) 00 Ol
MNJ4^4^tsjOtOtO Qω(刀44μj回心ドーJQ(刀CO .tk Ji. i{^ CD toぱ3 6O Ji> to O tsj り N)
●●●● ●●● ● ●●●● ● ●●●●● ● ● ●● ●● ● ●
00 00 CXI CO J^ CT) CT) J^ i^ 00 CTl 03 C乃り4^ Cfl 4i>. ^ 4(刀 4(乃44 ふ 4 ヽノ-●・---~一之ヽノ ヽノ-●ヽノこ-ヽ-●--ヽノ-ヽノ--ヽ------ヽノ-・ヽ-- - ヽ●
W
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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・・・・・十・・・・・+・・,、j+・・ ・・・・・・・・・・・・・孚‥Ξ♀‘♀♀♀ ♀‥ΞΞ
- ヽ-● --- -
…………→-・・・・・。 ‥忖レ仁・悦仁‥悩聯 E・半農
WJW 心 WW 心 心JW 心
W i-,回色S
SO
・ ・++・ + ・++・+ + !-・+・ ・ ・ ゛・トト・万言≒諮‥・とに回旱 孚・惣芭
心ヘノ W W WWJWJ W
‥‥‥‥++・・++・・・・+ ‥‥‥‥ヤ‘バ‾⑤シ.゛・ヨデモFモシモFy ♀・回心
W W心 vy v>v._^S_V̂^ W + ++ + +++ + 卜.+トー十。 ωωω
・+・・・・++++s+トーJ・・・・ト,j ・・+ ++゜♀‥ 采♀♀’ ’孚 孚 孚 Ξ゜♀Ξ孚
心 ・^-^・v-^ W 心/WJ W 心 W W J
‥‥・+‥+‥++‥-‥ ‥.几‥‥へ・モ卜口・ ・ ・ ・ 'P 'P 首.c^)゛右
゜゜十゜゜゛十゜’゛゛゜十十゜・ t- ・ ・ ゛十゜’’’゜十‥‥宍‥‥十゛且♀ピ モF.c^)`)竺
十・・・十・+・十十十十。・・・・・ω ‥‥・♀ト‥・孚‥‥・ヤ万万 ・ぐ昆
心/yノ tノ 心いノ 心 W
C/つを-JS旨色
w o
十十・●十・十・十十十十・十・・・・ ・・・・.太太太h―' . .太.....只゛‘j i_ii__iCO .⌒、欠坦坦
古士士 士 士 士 士 ’じ古など
++ ト_, + +,。十e、J ωωω
・+・・+・+・・トー十十・+・・・。 ゜゜゜’゜♀♀‥‥常‥・+ ++゜常常 ツΞ言言
心/心ノ 心 W WW 心心 W
++ + +十十十 s ト. +H-.+ト.,ω ,。ωω‥‥十‥十゛‥十十‥・.゜・ 平常‥ ♀平♀‥ワ゜♀’Ξ♀Ξ♀Ξ ワ ΞΞ
ヽノ之 --こ ごヽノ ヽノ - ヽノ
+ 十十十 十s 肖。u uω・・・・・・・+・・・十・・・+・. ゛Ξ’‥Ξ♀゜♀‥・♀’♀゛Ξ♀Ξ孚Ξ ツ 白白
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2にj回色目
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+十 十十 十ω トー4・・・・・・・・十・・・・・・十・・J ・・・‥♀♀・・・・・・♀ワ‥゜♀ツ ツΞ谷゛
tノW W 心ヘノ 心ヘノ ● ’ 十 トJ 十 り WNJCOトJ
・゜゜゜゛・・・・゛゜・゛・゛トー+s .・・・・・・・・・・・・・T‥ワ゜Ξワ ワΞ言 - - - -
十十 ト一so、J 四十・・・・・・・・・・・・・・・●.・. ・・・・・・♀’’゜’♀ワ‥’♀゜Ξ゜♀ ゜ΞΞ言
ヽノ -- ヽ-● -
………・‥・・・・・・ヽ ……4デ……モtデ・・べ 十 十 H トーJ (刀 Qトー吋、j
・・・●・・・・・・・・・・・+・・. ・・・・・・孚・・・・・・孚‘゜゜T’♀゜孚 ’Ξ ΞΞ
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w o
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- - -
F I Sゝ H H Kに、)ドj・・・・・・・+・・・十・・・・・・. ‥♀‥‥♀‥‥ワ’ ・Ξ’Ξワ ゜言言言言
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+ +s 、+ Q ,。 4,。ω
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t\j t>j en en トJトーJ SJ CO 010)0 ト-< CO en CO
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ドJトーJトjトーJトーJトーJ CO CO NJ h一一JCsJ CDりト-' O NJ Oトーj トーJトー』トーJトーjトーJトーJωcot-'トJトーJトjeQωocsjotOt&^Njroo oo o o o
●● ● ●・ ● ●● ● ● ● ●● ●●● ●●● ● ●● ●●●● ●●●● ● ● ● ● ●
QQN〉WOIO) 00 CO C乃ム4 to Ji" CO Ji. O) to N3に■, C7) en en lo a> to tsj M (刀心(JI み Q)ム ム CX)
⌒⌒⌒⌒⌒⌒⌒⌒⌒ ⌒⌒⌒⌒⌒⌒⌒⌒⌒⌒⌒ ⌒⌒⌒⌒⌒ トーJ トー゛トーJ en toωぱ)
4 Q 4M K3 fO CD Q O (7りQCχ0 NJ O
OCs、)トーり一一JMQωo o to CO en o tvj o enχjcnj^ i~j Qeりトー-JOトーJ
・● ● ● ● ●●● ●● ●●● ●● ● ● ● ● ●● 号 C04i.tsJcn (DtOCOOO CDrf^Olttv^J^Cχ)ム 4M *.aiCDlD こ--・ノヽノヽノ-ヽノヽノ -----ヽノ--ヽノ-- -----
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国加==回国l=(l=l自明回向国回詞呻≒=41=4詞匝=!爾 Ml=i!磯回自畑=4-・=ia≒s4≒呻哨.か・一回『-・i固ぶ~ぶべ 哨ぶMべχ Constancy
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