Natural and semi-naturalvegetation in Japan

M. Numata A. Miyawaki and D. Itow

Contents

I. Introduction 436

II. and in Plant life its environment Japan 437 III. Outline of natural and semi-natural vegetation 442

1. Evergreen broad-leaved forest region 442

i.i Natural vegetation 442 Natural forests of coastal i.l.i areas 442

1.1.1.1 Quercus phillyraeoides scrubs 442

1.1.1.2 Forests of Machilus and of sieboldii thunbergii Castanopsis (Shiia) .... 443 Forests 1.1.2 of inland areas 444

1.1.2.1 Evergreen oak forests 444

Forests 1.1.2.2 of Tsuga sieboldii and of Abies firma 445

1.1.3 Volcanic vegetation 445

sand 1.1.4 Coastal vegetation 447

1.1.$ Salt marshes 449

1.1.6 Riverside vegetation 449 lake 1.1.7 Pond and vegetation 451

1.1.8 Ryukyu Islands 451

1.1.9 Ogasawara (Bonin) and Volcano Islands 452

1.2 Semi-natural vegetation 452

1.2.1 Secondary forests 452

C. 1.2.1.1 Coppices of Castanopsis cuspidata and sieboldii 452

1.2.1.2 Pinus densiflora forests 453

1.2.1.3 Mixed forests of Quercus serrata and Q. acutissima 454

1.2.1.4 Bamboo forests 454

1.2.2 Grasslands 454

2. Summergreen broad-leaved forest region 454

2.1 Natural vegetation 455 Beech 2.1.1 forests 455 forests 2.1.2 Pterocarya rhoifolia 457 daviniana-Fraxinus 2.1.3 Ulmus mandshurica forests 459 Volcanic 2.1.4 vegetation 459

2.1.5 Coastal vegetation 461

2.1.5.1 Sand dunes and sand bars 461

2.1.5.2 Salt marshes 461

2.1.6 Moorland vegetation 464

2.2 Semi-natural vegetation 465

2.2.1 Secondary forests 465

2.2.1.1 Pinus densiflora forests 465

2.2.1.2 Quercus mongolica var. grosseserrata-Castanea crenata forests 465

2.2.2 Grasslands 465

Contribution from No. *) JIBP-CT 34 supported by the special research project 'Studies on the dynamic status of the biosphere' sponsored by the Ministery of Education.

2 ) Department of Biology, Chiba University, Yayoi-cho, Chiba, Japan. 3) Faculty of Education, Yokohama Nat. University, Shimizugaoka, Minami-ku, Yokohama, Japan.

4 ) Faculty of Liberal Arts, Nagasaki University, Bunkyo-cho, Nagasaki, Japan.

12 436 BLUMEA VOL. XX, No. 2, 1972

3. Subalpine and subarctic coniferous forests region 467

in 3.1 Abies mariesii-A. veitchii forest Honshu 467

3.2 Mixed forests of Thuja standishii and Tsuga diversifolia 468

Larix 3.3 leptolepis forests 469

3.4 Thickets of Quercus mongolien var. undulatifolia 469 ermanii 3.5 Betula thickets 469

Picea forests in Hokkaido 3.6 jezoensis-Abies sachalinensis 471 Picea 3.7 glehnii forests 471

3.8 Tall herb communities 471

4. Alpine region 472

of Pinus 4.1 Thickets pumila1 472

4.2 Snow-patch vegetation 473 heathlands 4.3 Wind-blown 473

4.4 Alpine desert vegetation 474

IV. Epilogue 474

Literature cited 475

I. Introduction

Studies on the composition of the forests and other vegetation of Japan were initiated by J. Tanaka (1887) in the Meiji Era. According to his introduction his field survey in and resulted into classification of started 1879 a vegetation zones (forest zones), mainly based dominant later of the lowland, on physiognomy and species. In years some other Honda also major proposals of the vegetation zonation were made. (1900) distinguished based his and it with the first altitudinal zones, on survey on dominance, provided vege- tation map. Nakano (1942—1943) stressed the physiognomy combined with species

and succession but did not Kira described composition, sociology, produce a map. (1949) the of the climax and forest zones Japan stressing concept correlating ecology mostly he with with temperature; added a map. Miyawaki (1967) wrote a large book (provided

the in a map) on vegetation of Japan which the climax concept played an important role, did the Suzuki who tried actual as it in study by (1966a) partly to integrate vegetation with climax to the he also coloured types, paying great attention sociological aspect; gave a in of communitiesand map. Miyoshi (1903) introduced Japan the study plant some years later (1907) the study of ecology, and these two disciplines have been pursued by the his Yoshii research schools of pupils, Nakano at Tokyo University, at TohokuUniversity, etc. A large number ofpapers have resulted in these two fields ofvegetation study, mainly published in the Botanical Magazine, Tokyo (Botanical Society of Japan, 1887—), Japanese of Journal ofBotany (Japan Science Council, 1922—), Journal Japanese Botany (Tsumura Acta Geobotanica Laboratory, 1926—), Phytotaxonomica et (Kyoto University, 1932—), Bulletinof the ofPlant Ecological Review (Tohoku University, 1935—), Japanese Society

& Ecol. Ecology (Tokyo University, 1941 —1943), Physiology and Ecology (Physiol. Pub- lish. Ass., Kyoto, 1947—), Hikobia (Hiroshima University, 1950—), Bulletin of the Society Hokuriku of of Plant Ecology (Tohoku University, 1951 —1954), Journal Botany or Jour- nalof of Geobotany (Kanazawa University, 1952—), Japanese Journal Ecology (Ecological

Society of Japan, 1954—), etc. Besides the these of have contributed papers in journals, science reports universities to the development of ecological studies in Japan. A historic review of plant ecology in

Japan and basic concepts, principles, and methodologies of ecology was given by Numata revised edition and brief ofthe literature concerned (1953, 1967) a very bibliography was published in ‘Ecology’ (Numata, 1958).

The ofthis is to describe briefly the purpose paper entirepicture of Japanese vegetation M. Numata et ah: Natural and semi-natural vegetation in Japan 437

with reference the studies. Such overall has been to more important recent aspect strongly and first author requested by many foreign botanists, ecologists, geographers, and the

realized the of concise of when he keenly desirability a summary Japanese vegetation gave brief lecture the of of a at Department Geography, University Kansas (Prof. A. W.

Kiichler) and the Department of Botany, University of Illinois (Prof. L. C. Bliss) during

his the recommendation of Prof. van stay in 1967. Moreover, Dr. C. G. G. J. Steenis the nth Pacific Science in has whom we met at Congress in Tokyo 1966 led to the com- of this The pletion paper. first author, attending an IBP-CT Meeting at Monks Wood,

to discuss the check sheet for reserved submitted England 1965, areas, (with Itow) a paper 'Outline of and environment of this vegetation natural Japan'. From working paper the critical has emanated. present compilation

II. PLANT LIFE AND ITS ENVIRONMENT IN JAPAN

The climate of is in wide it is arctic subarctic in Japan temperate a sense, though or

northern Japan and highlands, particularly in winter, and subtropical in southwestern

The Japan, particularly in summer. four major islands of Japan proper, Hokkaido, Honshu,

0 Shikoku, and Kyushu are located between 30° and 45 NL, the Ryukyu and Bonin

Islands extending further south almost to the Tropic of Cancer. Thus, the Japanese realm chain of islands of latitude. is a long ranging over c. 22° rather cold like the Even in Tokyo at 36° NL it is in winter, andhot in tropicsin summer.

In summer the lowlands in southwestern Japan are hotter than those in Taiwan, and in the Tohoku winter District at about 40° NL is colder than northern Europe at 6o° NL

(Ministry Agr. For., 1961).

The distinction of the four in the seasons is, general, very clear, though periods of

and autumn rather shorter in the northern Weather and climate spring are part. are

variable and differ locally. Japan as a whole is under the influence of a monsoon climate. wind of The southeast monsoon in summer brings a large amount rain on the Pacific

side, and the northwest monsoon wind in winter brings a large amount ofsnow on the

Japan Sea side and in Hokkaido (Nemoto et al., 1959).

There two of thePacific and the because are mainly types climate, type Japan Sea type, of influences of and the backbone mountain the above-mentioned monsoon ranges the four islands. These climatic from the running through major types appear clearly climate diagrams (Fig. 1). The often from the Lowland mountainous topography begins sea coast. plains are few and very valleys are deeply dissected. Pavers are mostly short and swift and even the

river in the Shinano is is longest Japan, River, only 369 km. Of the Japanese land 77 % between and m 18 between and and 0 700 altitude, % is 700 1300 m, 5 % is highlands is over 1300 m. The inclination of slopes rather steep, being over 30° in 24 % of the

0 0 in and less than in The is land, 30°—15 31 %, 15 45 %. geomorphology very compli- cated.

As characteristics of the the principal Japanese climate, annual mean temperature is

from o° C (central Hokkaido) to 18° C (southern Kyushu), the annual precipitation is

600 mm at the minimum(northeastern Hokkaido) and about 4,000 mm at the maximum

(Odaigahara in Kinki District in southern Honshu and Yaku Island south offKyushu).

The annual which a average precipitation in Japan proper is 1,600 mm is high figure with other of the world latitudes and of compared parts at comparable testimony an

oceanic climate.

The frostless in the lowlands is from in northeastern Hokkaido to period 125 days 275 BLUMEA VOL. No. 438 XX, 2, 1972

Sea Fig. 1. Paired climate diagrams of the Japan type (left) and the Pacific type (right), both lowland stations. The is Sea U-shaped curve of the precipitation distribution characteristic to the Japan type and the reversed is that of the Pacific Scales for and U-shaped curve type. temperature precipitation are on The minimum left and right hand respectively. black bar indicates the months with mean temperature

below O° C, and the batched bar indicates the months with the absolute minimum temperature below

Mean O° C. annual temperature and annual precipitation are given in the right-top of the diagram.

southern and southern Kinki and Shikoku. days in Kyushu 300 days in (central Honshu) and A distinct dry season between the summer monsoon rain ('Bai-u', in June-July) the the from the lower reaches typhoon season (in September-October) appears in area of the Yangtze to central Honshu.

Not only the amount of annualprecipitation, but the seasonal distribution of rainfall for life and the in particular is important plant (Numata Mitsudera, 1961). According to the hrs. record maximum rainfall per 24 (Nemoto et al., 1959) was 200 mm in Hokkaido

in there and 500 mm southern Shikoku and Kyushu; are occasionally exceptional figures, such record of hrs. in local rain-storm at as a over 1,100 mm per 24 a Isahaya-shi, Kyushu, on 25 July 1957. several based the conditions The chmate in Japan has been classified into types on mentioned above Such classifications become rather (Fukui, 1933; Sekiguchi, 1959). complicated by the topography which makes application to the various local situations difficult.

climatic Hokkaido Df and others almost Cf the According to Koppen's types, is are in lowland. According to Fukui's classification Japan is divided into three regions, viz. southwestern northern Japan (Hokkaido, except its Oshima Peninsula, having more central than 4 months under 0° C monthly mean temperature), Japan (under 20° C mean and less than months under and temperature 3 o° C in monthly mean temperature), Bonin C in annual all southern Japan (Ryukyu and Is., over 20° mean temperature), figures derived from lowland records. the Regarding phenology, average flowering date of for example Prunus yedoensis (a M. Numata et al.\ Natural and semi-natural vegetation in Japan 439

March in spring-flowering plant) is 31 in southern Kyushu, 10 May the greater part of Hokkaido; the flowering date of (autumn-flowering) Miscanthus sinensis is early in July

in Hokkaido, at the beginning of October in southern parts of Kinki, Shikoku, and

Kyushu. This phenomenon is mainly due to the temperature differences and day-length effect between north and south.

The environment for plant life should be measured regarding its factors working

the Kira conceived kind of accumulated during vegetative season. (1949) a temperature

E — index, the warmth index (W), i.e. W = (t 5) for the lowland. W is an accumulated

temperature of monthly mean temperature as the base temperature is, in general, equal 0 the warmth to 5 C. The climatic classification of Japan by index is shown in Table 1.

Table of after 1. The climatic regions Japan Kira (1949)

Climatic region Warmth index Area Latitude

Subarctic V) 1 Tt" 1 N. E. Hokkaido 43—45° Cool-temperate (45—55)—85 S. W. Hokkaido and Tohoku 37—43°

X W OO 1t— O Kanto-Kyushu 30—37° arm-temperate 1

Subtropical 180 —240 Ryukyu Is. and Bonin Is. 24—30°

Kira also defined a coldness index (C) = £ (5 - t) and mentioned that the Castanea intermediate the and thatof crenata zone is an zone between evergreen Quercus zone the

Fagus crenata zone, covering the zone over 85 in W and over 10—15 in C.

Reversely, Nakamura (1954) maintained that forests of Pinus densiflora, Quercus serrata, of Castanea crenata, Albizzia julibrissin, etc. are established at the sites fire and should not

be used as indicators of classifying forest zones. climatic the stand- Schmithiiscn (1965, private comm.) classified Japanese zones from

point of a European geographer (Table 2).

Table Another classification of Schmithüsen 2. Japanese climatic zones after (1965, private comm.)

Climax Climatic zone

Evergreen needle-leaved forest Cold-temperate

Fagion crenatae Temperate Cool-temperate

sieboldii Tsugion Warm-temperate (Sub-mediterranean)

Sliiion sieboldii Subtropical True-mediterranean

Sclerophyllous forest BLUMEA VOL. No. 440 XX, 2, 1972

As secondary communities, such as pine forests, grasslands, etc. occupy considerable

surfaces, their correspondence to climax and climate should also be considered (Table 3)

Table 3. Grassland vegetation zones in Japan (Numata 1961a, 1969)

Climate Climax Meadow Pasture

Subarctic Evergreen needle-leaved Sasa Poa

forest -Miscanthus pratensis

Cool-temperate Deciduous broad-leaved Miscanthus Zoysia

forest japonica

Warm-temperate Evergreen broad-leaved Miscanthus Pleioblastus forest -Pleioblastus -Zoysia

offorest As stated before, precipitation is sufficient toallow the development throughout the where the low the factor. Japan except in alpine zone temperature is limiting Thus, altitudinal and latitudinal zonation of vegetation in Japan are largely dependent upon conditions. temperature

Climatic climaxes are physiognomically recognized as evergreen broad-leaved forest

in in the the warm-temperate zone, summergreen broad-leaved forest cool-temperate,

needle-leaved forest in the subarctic and and scrub communities in evergreen subalpine,

the alpine zone.

The first three of these climaxes are phytosociologically understood as the Camellietea

japonicae, the Fagetea crenatae and the Vaccinio-Piceetea japonica respectively. The fourth,

which is the Pinus pumila-dominated scrubland, also belongs phytosociologically to the

Vaccinio-Piceetea japonica. the whole climatic These climatic climaxes do not prevail in area of each zone, but

minor natural communities are developed on edaphically, topographically, and micro-

climatically delimited habitats.

to the of each The regional subdivisions of Japan here are made according range

climatic climax, that is, the Camellietea japonicae region where the climatic climaxes also belong to the Camellietea japonicae and minor natural communities are developed

according to habitat variations. In other words, the Camellietearegion is physiognomically characterized the broad-leaved forests and the by evergreen climatically by warm- climate. The the broad-leaved forest temperate Fagetea crenatae region is summergreen region and is cool-temperate; the Vaccinio-Piceeteajaponica region is the evergreen needle-

leaved forest region and is the subarctic and subalpine zone.

Latitudinally, the Japanese territory does not extend to the arctic, but altitudinally to the the of the four mentioned alpine zone. Fig. 2 is the map that shows ranges regions of above. A north—south cross section of the Japanese Archipelago is given on the left

Fig. 2. above based the The classification of vegetation regions stated is usually on presence

absence of On the other local characteristics are shown or a special vegetation type. hand,

by the difference of growth and productivity of widely distributed vegetation all over of The the country, for example, the Miscanthus sinensis type grassland. physiological and the zero point of the perennial Miscanthus sinensis is 10° C daily mean temperature M. Numata et at.: Natural and semi-natural vegetation in Japan 441

decided with the end of its growing season is flowering date. The growing period is 60

days in northeastern Hokkaido, 100 days in Tohoku, 160 days in Kanto and Chugoku,

180—200 days in Kinki and Shikoku, and 240 days in southern Kyushu. The effective and accumulative temperature (EAT) over io° C is 1,500°C in Hokkaido Tohoku, and

2 4,000° C in southernKyushu. The effective accumulated insolation (EAI, Kcal/cm /period)

is 22 in Hokkaido and Tohoku, and 100 in southern Kyushu. Besides these, a kind of

aridity index shows a drier climate in Hokkaido and a more humid one in southern

Shikoku and Kyushu. The local productivity of Miscanthus sinensis grasslands correlates and with EAT, EAI, aridity index, etc. (Numata Mitsudera, 1969).

The soils of classified three under the Japan were by Kanno (1953) into main types

concept of Swanson's soil associations; he showed the relationships between each soil N-S annual association, precipitation, quotient, mean temperature, etc. (Table 4).

Table Soil of 4. associations Japan (Kanno, 1953)

Soil association Rain factor N-S quotient Annual mean

temperature °C

-8 Podzol ass. 150—250 < 600—1000 < 5

soil 8 Brown forest ass. 90—170 < 400— 950 < —14.4

Red soil ass. 70—150 < 350— 700 < 14.5—18

A has the map of soil types, scale of 1:200,000, with explanation, been compiled by National of Table Institute Agricultural Sciences (Kamoshita, 1958). 5.

Table Zonal soil of 5. types Japan

Zonal soil types Distribution

I. Podzol northern Hokkaido

2. Rust-coloured forest soil northern sand-dune

(weakly podzolized)

3. Brown forest soil central Honshu

4. Greyish brown forest soil northern Honshu and Hokkaido

5. Red soil southwestern Japan

6. Reddish brown soil southwestern Japan

the soil the numbers and the main and others transition- Among types 1,3, 5 are types are al. Azonal soil types are, in Japan, mostly belonging to mountain soils and colluvial

Intrazonal soils into and in soils. are classified wet soil types (bog soil half-bog soil boggy meadow soil forest and rock soil areas, in paddy areas), wet soil, types (terra rossa, recent VOL. No. 442 BLUMEA XX, 2, 1972

volcanic ash soil, etc.). Muck soil is distributed in Hokkaido, Tohoku, and lowlands of

Kanto. Volcanic ash soil, which is particularly important in relation to the grassland vegetation, is distributed in many areas in Hokkaido, Kanto, Kyushu, etc.

There are some areas severely eroded in forest-lands and grasslands. Such areas are found in Chugoku (western Honshu) and in the mountainous parts of western Shikoku,

sheet erosion. subjected to a predominantly bare or They are caused mainly by the prop- erties of the mother rocks and features. In there is of topographic Japan, a great danger severe accelerated erosion when the plant cover is removed or heavily degraded through rainfall and the heavy steep topography.

28 In Japan there are 23 National Parks and quasi-National Parks. The former occupy of the of the total surface of about 5.3 % area Japan (36,978,000 ha) and the latter about have 1.8 %. Among those, we Special Protection Areas as strictly protected reserves.

Protected National and Besides these, principal vegetation types are reserved as Forests

National Natural Monuments.

III. OUTLINE OF NATURAL AND SEMI-NATURAL VEGETATION

forest I. Evergreen broad-leaved region. Table 6—I. Phot. 9, 10, 12, 14.

forests in this have several of broad-leaved Natural region types evergreen forests.

Major tree species are Machilus thunbergii, Castanopsis sieboldii, and C. cuspidata in coastal and areas, and Quercus glauca, Q. gilva, Q. salicina, Q. myrsinaefolia, Q. acuta (evergreen oaks) in inland areas. In higher elevations, Tsuga sieboldii and Abies firma occur.

The natural vegetation including climax forests and other minor communities in this

the region (Fig. 2) has been largely deformed and destroyed since rice cultivation was introduced to Japan about two thousand years ago. The rice plant, Oryza sativa, is of humid-tropical origin, and its cultivation in Japan restricted the was largely to this warm-temperate region before Meiji Era, about 100 It is certain human time years ago. that population around rice-fields at that was denser than in other regions where rice was not or little planted.

of this denser the still reflected the fact that A consequence population in past is in

twenty-nine out of thirty cities with a population over 300,000 are presently situated in and fields lowland this region. At present, rice- upland prevail there, especially in areas, and secondary forests of Pinus detisiflora, Quercus serrata, and Q. acutissima and grasslands of Miscanthus sinensis, Zoysia japonica, Pleioblastus chino, and P. distichus var. nezasa sub-

stitute the original vegetation on foot-hills and lower mountains (Numata, 1961a).

Natural forests remain fragmentarily in sanctuaries around temples and shrines and on

inaccessible steep slopes of gorges and mountains (Miyawaki and Itow, 1966).

1.1 Natural Vegetation

1.1.1. Natural forests of coastal areas

1.1.1.1. Quercus phillyraeoides scrubs. Phot. 12.

Coastal slopes with shallow soils on the Pacific side support a scrub community that

is dominated and characterized by Quercus phillyraeoides. The scrub is distributed from the Inland southern Kyushu, through Shikoku and Sea (Setonaikai) region, to the Izu Honshu. the Peninsula in central Q. phillyraeoides is a sclerophyllous shrub and community

is physiognomically that of maquis. of the scrub such and shrubs Main components Q. phillyraeoides are evergreen trees as Q. phillyraeoides, Pinus thunbergii, Ligustrum japonicum, Rapanea neriifolia, Camellia japonica, M. Numata et a\.\ Natural and semi-natural vegetation in Japan 443

and Eurya japonica, E. emarginata, Vaccinium bracteatum, herbs as Farfugium japonicum,

Paederia scandens, Cyrtomium fortunei, etc.

Phytosociologically, two associations have beenreported: Pittosporeto-Quercetum philly-

raeoidetis (Suzuki and Hatiya, 1951) and Gleichenio-Quercetum phillyraeoidetis (Imai, 1965). is The former association characterized by the presence of Pittosporum tobira, Pyrrosia and Scutellaria indica the latter lingua, Rhaphiolepis umbellata var. integerrima, var. japonica;

by Gleichenia dichotoma and Rhododendron tosaense.

Quercus phillyraeoides and its scrub also occur on Yaku Island, which is situated south of the mainland. Another of coastal forests from the northeastern Kyushu type is reported

coast of the island. It is composed of Cinnamomum daphnoides, Eurya emarginata, Litsea

japonica, Rhaphiolepis umbellata, Daphniphyllum teijsmannii, Camellia japonica, Euonymus C japonicus, Psychotria serpens, Farfugium japonicum, etc. (Okutomi, 1968). innamomum distributed restricted from the daphnoides, a sclerophyllous shrub, is in a area, ranging of the Danjo Islands at the north, through southernmost coasts Kyushu mainland, Tane-

gashima, Yaku Island, the Tokara Islands and Amani-oshima, to Iwotori-shima of the the south the Ryukyu Islands at (Horikawa, 1961; Toyama et al., 1967). Probably,

Cinnamomum daphnoides scrub is ecologically equivalent to the Quercus phillyraeoides scrub.

and Quercus phillyraeoides Cinnamomum daphnoides are not distributed on coastal slopes of in regions on the Japan Sea side of Kyushu and Honshu. The coastal slope vegetation

those is made of ofthe above-listed these areas up most species except two species, though

details have not yet been studied.

Phot. 1.1.1.2. Forests of Machilus thunbergii and of Castanopsis (Shiia) sieboldii. 9.

Since gentle slopes offoot-hills and alluvial plains are almost completely changed into

farmlands, paddy-fields, and urban areas, relict stands ofnatural forests are the key to the

original communities that were there before man touched them. Based on data from

such relict stands, mesic habitats with deep soils onalluvial plains originally supported the

forests dominated Machilus and by thunbergii (Suzuki, 1952, 1953; Numata Asano, 1965). Although the floristic composition of the M. thunbergii forest shows somewhat regional

variation, common components in Shikoku, for example, are such evergreen broad-

leaved trees as Castanopsis sieboldii, Camellia japonica, Elaeocarpus sylvestris, Ligustrum Neolitsea japonicum, Distylium racemosum, Symplocos lucida, Cinnamomum japonicum, sericea, Ilex Actinodaphne lancifolia, Daphniphyllum teijsmannii, integra, Myrica rubra; evergreen

shrubs as Aucuba japonica, Gardenia jasminoides f. grandiflora, Maesa japonica, Euonymus

deciduous Calli- japonicus, Rapanea neriifolia, Damnacanthus major; shrubs as Ficus erecta, herbs Arisaema carpa japonica var. luxurians; as Alpinia intermedia, ringens, Liriope platyphylla, and lianas Ophiopogon japonicus; ferns as Rumohra aristata, Dryopteris erythrosora; as Trache-

lospermum asiaticum, Piper kadzura, Hedera rhombea, Anodendron affine, Kadzura japonica, four Ficus nipponica, etc. (Yamanaka, 1962). Phytosociologically, associations have been

described: Liriopi-, Rumohro-, Polysticho-, and Polypodio-Machiletum. (cf. Suzuki, 1966b).

Castanopsissieboldii is almost always mixed with Machilus thunbergii, butit is a dominant

species of forests in habitats with rather drier soils on foot-hill slopes and ridges. In rather

inland portions, Castanopsis cuspidata also occurs (Numata and Asano, 1965). Relict

stands are frequently found in forest reserves and national parks, as compared with those

of the sieboldii almost similar of Machilus thunbergii. Component species C. forest are to

those of the M. thunbergii forest, though their quantitative relations are not always alike.

Phytosociologically, seven associations have been described: Bladhio-, Rapaneo- (Suzuki,

1952), Symploco- (Nomoto, 1953), Ardisio- (Hosokawa, 1958), Lasianthero-, Arisaemo-, 444 BLUMEA VOL. XX, No. 2, 1972

and Symploco liukiuensae-Castanopsietum (= Shiietum) (Miyawaki and Ohba, 1963). The

first three associations were reported from Honshu, Shikoku, and the Kyushu main-

land, the fourth from Yaku Island, and the last three from Amani-oshima and Tokuno- shima.

As stated by Hara (1959), Amami-oshima and Tokuno-shima belong to the Ryukyu which number of Floristic Region, in a tropical and subtropical species occur. Examples

are Wendlandiaformosana, Cinnamomumdoederleinii, Symplocos microcalyx, Randia canthioides,

Eurya oshimensis, Illicium anisatum, Helwingia liukiuensis, Ligustrum liukiuensis, Callicarpa

oshimensis, Osmanthus bracteatus, Smilax nervo-marginata, Turpinia ternata, Syzygium buxi- folium, Lasianthus tashiroi, Psychotria rubra, P. serpens, Crataeva religiosa, Schefflera octophylla,

Ardisia quinquegona,and Rhododendron tashiroi. Amami-oshima Island is the northern limit of the first eleven ofthe above-listed of sieboldii species. They are components Castanopsis of island and The of forests the (Miyawaki Ohba, 1963). rest those species are also found

in natural forests on Yaku Island, some of which are distributed further north to the

southernmost part of the Kyushu mainland.

1.1.2. F o r e s t s of inland areas

of the also been Inland areas evergreen broad-leaved forest region have exploited, and and and there presently prevail agricultural areas secondary vegetation such as coppices

grasslands. Based on relict stands of natural forests, however, it is certain that the area of originally supported forests evergreen oaks, Tsuga sieboldii and Abies firma.

oak 1.1.2.1. Evergreen forests. Phot. 10. oaks mentioned here Evergreen are Quercus glauca, Q. gilva, Q. myrsinaefolia, Q. salicina,

and Q. acuta. These oaks are sometimes treated as species belonging to the genus Cyclo- balanopsis from the morphology of their cupules.

Natural forests of Quercus glauca are distributed mostly in the area of the Castanopsis

sieboldii forest, and are infrequently found in inland areas in western Japan (Yamanaka,

The habitat of limestone and andesite and 1966; Okutomi, 1967). is (Suzuki Mori, 1957;

Suzuki et al., 1964; Miyata and Shiomi, 1965; Yamanaka, 1966). The floristic composition forest similar that of the forest. of the Q. glauca is, as a whole, to Castanopsis sieboldii characteristic According to Yamanaka (1966), species are Q. glauca, Xylosma japonica, Eriobotrya japonica, Trachycarpus fortunei, Citrus junos, and Nandina domestica. The Quercus gilva forest is frequently found on deep soils at gentle slopes of foot-hills.

that first the It is thought pre-historic man settled in the habitat of Q. gilva forest (Suzuki, the i960; Miyawaki and Fujiwara, 1969), and therefore relict stands are not abundant.

In the Kanto District of central Honshu, the Quercus myrsinaefolia forest is thought of habitats with as the original forest community that was developed on loamy soils (Yoko-

and Main of yama et al., 1967; Miyawaki, 1967; Miyawaki Fujiwara, 1968a). components

the forest are such trees and shrubs as Quercus myrsinaefolia, Camellia japonica, Eurya

japonica, Aucuba japonica, Ardisia japonica, Fatsia japonica, Neolitsea sericea; herbs as Ophio-

ferns as erythrosora, D. uniformis, etc. pogon japonicus, Liriope platyphylla; Dryopteris (Miya- Natural stands of also inland waki, 1967). Q. myrsinaefolia are found in an limestone area

in western Honshu (Miyata and Shiomi, 1965).

The salicina forest is and with shallow at Quercus developed on steep slopes ridges soils, of some places, with exposed blocks the bedrock. Main components of the forest are

Cleyera japonica, Skimmia japonica, Pieris japonica, Illicium religiosum, Cephalotaxus harring- Camellia Machilus Neolitsea tonia, Quercus acuta, japonica, thunbergii, sericea, N. aciculata, M. in Numata et al.\ Natural and semi-natural vegetation Japan 445

Cinnamomumjaponicwn, Castanopsis cuspidata, Eurya japonica, Aucubajaponica, Trachelosper-

mum asiaticum, etc.

Phytosociologically, Quercus salicina forests are classified into four associations (Suga-

numa, 1965). The first association is Osmantho-Cyclobalanopsidetum (= Osmantho-Quer-

cetum). It is characterized by the occurrence ofOsmanthus ilicifolius and Prunus jamasakura, distributed and is on foot-hills and lower mountains on the Pacific side of central and

western Honshu and in the Inland Sea District. The second is Distylio-Cyclobalanopsidetum that characterized is by Distylium racemosum, Symplocos myrtacea, Ligustrum japonicum, Ilex integra, and Machilus japonica. This association is found in Kyushu and in the southern

half of Shikoku. The third is Carici-Cyclobalanopsidetum, characterized by Carex reinii,

Hydrangea scandens, Lindera sericea var. tenuis, Cephalotaxus harringtonia var. nana, and

Struthiopteris nipponica (Horikawa and Sasaki, 1959a). The fourth is Aucubeto-Cyclo-

balanopsidetum (Sasaki, 1958). These two associations are reported from western Honshu. oak that inlandforests. Quercus acuta is an evergreen occurs frequently in It predominates

in the cloud zone of mountains in Kyushu (Suzuki and Sumata, 1964) and in western Honshu of the (Okutomi, 1967b). Main components Quercus acuta forest, for example, and observed in eastern Kyushu (Suzuki Sumata, 1964), are Q. acuta, Skimmia japonica

Cinnamomum (which are characteristic to the forest), Distylium racemosum, japonicum, Machilus thunbergii, M. japonicus, Quercus salicina, Ilex integra, Neolitsea sericea, N. aciculata,

Actinodaphne lancifolia, A. longifolia, Castanopsis sieboldii, Camellia japonica, Cleyera japonica, Aucuba Ligustrum japonicum, japonica, Trachelospermum asiaticum, Marsdenia tomentosa, etc.

1.1.2.2. Forests of Tsuga sieboldii and of Abies firma. Phot. 4.

The sieboldii forest on and Tsuga is usually developed mountain summits, ridges steep

slopes with shallow soils, at some places, with exposed blocks of the bedrock. Since the habitat of the forest alike that of the salicina of the is to Quercus forest, many components

also are common to both. Phytosociologically, five associations have been described:

Carici-, Rhododendro-, Symploco-, Illici-, and Pieridi-Tsugetum (cf. Suzuki, 1952, 1966b; Yamanaka, 1961). The Abies firma forest, on the other hand, is developed on gentle slopes with deeper with the forest named soils, as compared Tsuga (Yamanaka, 1961). It is phytosociologically Illici-Abietum firmae and is characterized by Illicium religiosum, Schizophragma hydran- Ilex Parabenzoin Ilex Schisandra geoides, crenata, praecox, macropoda, repanda (Suzuki, 1961). and Altitudinally, forests of Tsuga sieboldii Abies firma are situated in the transition

zone between the evergreen and the summergreen broad-leaved forest regions. Therefore, the lower of the and evergreen broad-leaved trees and shrubs occur in portion Tsuga-

Abies-forest zone. Examples are Pierisjaponica, Ilex pedunculosa, Neolitsea aciculata, Quercus acuta, Q. salicina, Ligustrum japonicum, Symplocos myrtacea, Eurya japonica, Illicium religiosum,

Cleyera japonica, etc. On the other hand, deciduous trees and shrubs occur frequently in the forest, especially in higher elevations. Examples are Ilex macropoda, Acanthopanax

scidophylloides, Symplocos coreana, Magnolia salicifolia, Sorbus japonica, Cornus controversa,

Viburnum furcatum, Lindera umbellata, Acer rufinerve, A. sieboldianum, Quercus mongolica

var. qrosseserrata. Carpinus japonica, C. laxifolia, Euonymus oxyphyllum, etc.

shows the distributionof communities the Fig. 3. natural forest in Kanto District in

relation to soil moisture and elevation.

Phot. 1.1.3. Volcanic vegetation. 25, 26. Active and extinct volcanoes abound in the Japanese Archipelago. Plant communities

on beds of lava, ashes, scoria and other ejecta differ from the above-mentioned natural BLUMEA VOL. No. 446 XX, 2, 1972

in Kanto Fig. 3. Two-dimensional representation of the forest community-habitat relationships the region, central Honshu. A. Machilus thunbergii-Polystichum polyblepharum Ass., B. Zelkova serrata--Orixa japonica Ass., C. Zelkova serrata--Acer palmatum Ass., D. Castanopsis cuspidata var. sieboldii-—Bladhia japonica

Ass.,IlliciumE. Quercus myrsinaefolia Ass., F. Quercus salicina-—Osmanthus ilicifolius Ass., G. Abies firma-—

H. Fraxinus I. Ass. religiosum Ass., spaethiana—Dryopteris polylepis Ass., Fagus crenata-—Cornus kousa The last

the broad-leaved forest two associations belong to summergreen region (after Miyawaki, 1967).

Mihara Izu-oshima Island central and communities. Mt. on in Japan Sakurajima in southern of volcanoes situated the Kyushu are representative active in warm-temperate region. They are fields for observing the primary succession on volcanic habitats, since is series of the year of eruptions and lava flows recorded. Fig. 4 shows the the primary of from observations succession plant communities, which was constructed in various habitats on Izu-oahima (Tezuka, 1961). Carex okuboi and Polygonum cuspidatum var. terminale and invade first the weatheredlava flow, and are followed by evergreen decid- shrub and broad-leaved forests dominated uous tree species. Last, evergreen by Casta- nopsis sieboldii and Machilus thunbergii are developed. The structure of the communities

the of the from 'low' and are changed along course succession 'sparse' at early stages to

and 'dense' later Soils also matured the from soils with 'high' at stages. are along course little of and and with thin a amount decomposed organic matters nutrients A horizon, to mature soils with much amount of those and with thick A horizon (Tezuka, 1961).

There also lava flows of different in southern The are ages Sakurajima, Kyushu. primary shown The here succession observed there is in Fig. 5 (Tagawa, 1964). two examples cited alike. and are quite In early stages of the succession, Miscanthus sinensis Polygonum cuspida- M. Numata et ah: Natural and semi-natural vegetation in Japan 447

of in volcanic Fig. 4. Schematic illustration the plant succession habitats on Izu-Oshima, central Japan. Miscanthus 1. Carex okuboi, 2. Polygonum cuspidatum var. terminale, 3. sinensis, 4. Alnus sieboldiana, 5.

6. Prunus 8. Weigela grandifolia, Cornus controversa, 7. lannesiana var. speciosa, Styrax japonica, 9. Fagara Neolitsea ailantoides, 10. Stachyurus praecox var. matsuzakii, 11. Ligustrum pacificum, 12. Eurya japonica, 13. Cinnamomum 16. sericea, 14. japonicum, 15. Camellia japonica, Ilex crenata, 17. Castanopsis cuspidata var. sieboldii, 18. Machilus thunbergii (after Tezuka, 1961).

role turn (var. terminale on Izu-oshima) in accumulating organic matters; then, Alnus

sieboldiana (on Izu-oshima), A.firma (on Sakurajima), and someother deciduousshrubs and

trees follow. Finally, the chmatic climax forest co-dominatedby Machilus thunbergii and

Castanopsis sieboldii is developed.

Coastal sand Table Phot. 1.1.4. vegetation. 6-Ic. 23. There such clear of coastal is not correspondence sand vegetation to the macroclimatic that of northern regions as inland vegetation. However, there are seaside vegetation types, and southern. The coastal zonation in the evergreen broad-leaved forest region is found

as unstable, half-stable, and stable zones from sea-coast to inland. The unstable zone is represented by Zoysia macrostachya, Calystegia soldanella, Carex kobomugi, C. pumila,

Glehnia littoralis, etc. The half-stable zone is represented by Ischaemum anthephroides,

The Fimbristylis sericea, Vitex rotundifolia, etc. stable zone is covered by Imperata cylindrica Machilus var. koenigii, Miscanthus sinensis var. condensatus, Pinus thunbergii, thunbergii, etc. and (Numata, 1949, 1961b; Numata Nobuhara, 1952).

The zonation of coastal vegetation is caused by the fore-shore sea current and wind- 448 BLUMEA VOL. XX, No. 2, 1972

A series of succession assumed from observations Fig. 5. primary of communities on lava flows in

Sakura-jima, southern Kyushu (after Tagawa, 1964, slightly modified).

Fig. 6. Schematic illustration of relationships between salt marsh communities and their habitats in central and western Japan (after Miyawaki and Ohba, 1969). M. Numata et al: Natural and semi-natural vegetation in Japan 449

salt the of saline and the borne (Numata, 1949; Kurauchi, 1964), by invasion water, The sand-movement by wind and sea-waves (Nobuhara et al., 1962, 1964), etc. coastal zonation is expressed by the life-form spectra (Numata and Nobuhara, 1952; Nobuhara,

1965, 1967). is into The coastal vegetation divided two types: open sea type and bay type (Numata, and The former 1949; Mitsudera Numata, 1964). is represented by Calystegia soldanella,

Zoysia macrostachya, Ischaemum anthephroides, etc. and the latter by Atriplex gmelinii, Phot. Calamagrostis epigeios, etc. 23. and The leaves of trees of the coastal forest are classified into two groups: deposite type

The former is leaf in which salt entrance type (Kurauchi, 1956, 1964). a type deposits on their surface into the and the latter is a in which salt scarcely enters tissue, type soon The coastal central enters into the leaves and damages are found. forest in Japan mainly consists of the former such Machilus Camellia Ilex species as thunbergii, japonica, integra,

Cinnamomum japonicum, etc.

Salt h 1.1.5. mars e s. Table 6-Id. Phot. 24.

Salt marsh communitiesare found at estuaries and lagoons where destruction by wave actions is weak. Many of such habitats have been destroyed by successive advances of

salt marshes central and reclamation on lands. Major plant in in western Japan are Zostera Z. Limonium Suaeda S. nana, marina, tetragonum, japonica, maritima, Atriplex gmelinii,

Carex scabrifolia, Zoysia sinica var. nipponica, Artemisia fukudo, Phragmites communis, P. karka, Scirpus iseensis, Aster tripolium, Kochia scoparia f. littorea, Cynodon dactylon, Triglochin maritimum, and some others (Umezu, 1964; Miyawaki and Ohba, 1969). The halophytic characters ofthese marsh and strand dune salt plants plants were studied, especially as to the osmotic pressure, germination, and growth (Tsuda, 1961). Fig. 6 shows the distri- bution of salt marsh communities in relation to habitat conditions (Miyawaki and Ohba,

these the Suaedetum restricted 1969). Of communities, japonicae and the Scirpetum iseense are the of central to northern Kyushu and Korea, and to Ise Bay region Honshu, respectively; the the and the Zosteretum nanae, Zosteretum marinae Phragmites communis community are also common in salt marshes in northern Honshu and Hokkaido; and the rest are dis- tributed only in central and western Honshu, Shikoku, and Kyushu.

is referred section 1.1.8. For the mangrove vegetation to

1.1.6. R i v e r s i d e vegetation

Riverside habitats are different from those of the above mentioned forest lands in soil and water conditions.

Generally speaking, the soils are made up of flood deposits and the ground water table

communis and dominant is high. Phragmites is a very common species on silty and clayey

rivers. deposits with a high water table along lower Alnus japonica is one of the common

trees growing in riverside wet habitats and often in reed swamps. The undisturbed stand of this is and the the species rare data, therefore, are insufficient, because habitat has been exploited for paddy-fields.

Salix gracilistyla and Phragmites japonica are common on gravelly beaches along swift

streams (Okutomi, in Miyawaki, 1967; Miyawaki and Okuda, 1969).

Phytosociological studies on riverside-, levee-, high water channel-, and flood plain for plant communities have been made especially dairy farming (Naohare, 1945, 1948, ofriverside shown from bare 1950, 1951, 1965). An example plant succession is a ground

to Pinus densiflora community through Phragmites communis-, Salix-, Imperata cylindrica-,

Pleioblastus chino- communities, etc. (Kurita, 1943). 450 BLUMEA VOL. XX, No. 2, 1972

modified).

slightly

1967, Miyawaki,

(after Islands Ryukyu

the

in communities

beach

of distribution

the showing Scheme

7.

Fig. M. Numata et al.i Natural and semi-natural vegetation in Japan 451

Pond and 1.1.7. lake vegetation order of and Aquatic plants are arranged in an emergent, floating-leaved, submerged of lakes ones from shallow to deep waters on beaches ponds and (Nakano, 1911—16;

Prominent of the Miki, 1957; Hogetsu, 1945). species emergent plant community are

Phragmites communis, Typha latifolia, T. angustata, Scripus tabernaemontani, S. triangulatus,

Nuphar japonicum, Sparganium stoloniferum; those of the floating-leaved community are

Nymphaea tetragona, Brasenia schreberi, Trapa japonica, Nymphoides indica; and those of the

submerged community are Myriophyllum spicatum, Ceratophyllum demersum, Najas minor,

P. communities also found beaches Potamogeton natans, crispus, etc. Emergent plant are on of the lower coursesof rivers wherethe water flow is slow or almost nil. The most simple

is of of Lemna aquatic community composed one or two such floating plants as paucicostata,

Spirodela polyrhiza, Wolffia arrhiza, Azolla imbricata, and Salvinia natans (Miyawaki, i960).

Islands. Phot. 1.1.8. Ryukyu 13,14.

The Ryukyu Islands are located between Kyushu and Taiwan. The climate is sub- and annual Okinawa tropical. Mean annual temperature precipitation at Naha, Island,

C and the islands of and are 22.0° 2148 mm respectively. Floristically, Okinawa, Ishigaki,

Iriomote, together with Amami-oshima, Tokuno-shima, and the Tokara Islands, belong Floristic to the Ryukyu Region (Hara, 1959).

The natural forest has been considerably disturbed on Okinawa Island, but less so on

Island. Natural broad-leaved forests in Iriomote and Iriomote evergreen (Miyata Odani,

1963) are composed of such trees as Castanopsis sieboldii, Quercus miyagii, Distylium shrubs racemosum, Michelia compressa var. formosana; as Ardisia sieboldii, Psychotria rubra,

Syzygium buxifolium, Lasianthus cyanocarpus, Daphniphyllum teijsmannii, Randia canthioides; ferns Rumohra as Lindsaea chienii, Abacopteris triphylla, aristata, Tectaria phaeocaulis. Dipla- Bolbitis zium virescens, koidzumii, Colysis pothifolia;t; lianas as Freycinetia formosana, Epiprem-

Smilax etc. the forest num mirabilis, Flagellaria indica, china var. kuru, Tree ferns in are

Gymnosphaera podophylla, G. denticulata, Alsophila pustulosa, and Cyathea faurier, epiphytes Details are Neottopteris australasica, Psilotum nudum, and Gastrochilus japonicus. have not

been studied as to the phytosociology and ecology of the natural forest, although the

flora of the Archipelago is well investigated (Hatushima and Amano, 1958). Phot. Mangrove forests at estuaries and lagoons are well developed in Iriomote. 13. There are six species: Bruguiera conjugata, Kandelia candel, Rhizophora mucronata, Sonneratia

alba, Lumnitzera racemosa, and Avicennia officinalis. They are arranged from Avicennia

officinalis and Sonneratia alba at the frontier, through Rhizophora mucronata and Kandelia

candel, to Bruguiera conjugata at the inner portion, and further to the stand community of Pandanus liukiuensis and and Of tectorius var. Barringtonia racemosa (Miyata Odani, 1963). and Kandelia candel these mangrove species, Bruguiera conjugata are also distributed to

Amani-oshima; the latter is further north to the islands of Yakushima and Tanegashima.

A stunted stand of this species is found at Kiire in the southernmost Kyushu (Itow, in

Miyawaki, 1967).

Strand communities on raised coral reefs and onsand beaches in the Ryukyus are different

those and from in Kyushu, Shikoku, Honshu. The community distribution on those habitats shown are schematically in Fig. 7.

Pinus luchuensis is endemic to the Ryukyu Islands, ranging from Akuseki at the north, of this through Amami-oshima, Okinawa, to Iriomote. The forest pine is seen on some

disturbed areas. Mowed and/or burnt habitats are dominated by Miscanthus sinensis and

Pleioblastus in linearis, or Imperata cylindrica var. koenigii (Yano, Miyawaki, 1967).

451 452 BLUMEA VOL. XX, No. 2, 1972

1.1.9. Ogasawara (Bonin) and Volcano Islands. Phot. 15. The Islands situated about km south between Ogasawara are 1000 of Tokyo, 26°3o' N in the and of islets. and 27°4o' latitude Pacific, composed eight major islands and many Volcano south km The Islands are situated further between 24°i5' and 24°5o', about 200

southwest of the Ogasawara Is. All the islands are of volcanic origin and the topography climate is generally steep. The is subtropical. Mean annual temperature and total annual and precipitation on Chichi-jima are 22.6° C 1613 mm respectively.

Floristically, the islands belong to the Ogasawara Floristic Region (Hara, 1959), and

in endemism are outstanding high (Hattori, 1908; Nakai, 1930; Hosokawa, 1934; Tuyama, endemic 1953). According to Tuyama (1968), eighty-five out of 128 ligneous species are

to the Ogasawara and the Volcano Islands. Dendrocacalia (Compositae) and Boninia (Ruta- endemic ceae) are genera. The the islands were inhabited since first quarter of the 19th century. The population

about before the World and the either increased to 8000 War II vegetation was destroyed disturbed of the left the islands Honshu then or considerably. Most people to in 1944. Since than and at present, less 200 persons inhabit only Chichi-jima Island. The vegetation has touched this Recent made of been less during period. survey on the Islands Chichi-, Haha-, and Ani-jima (Miyawaki, 1968a) offered an up-to-date information on the vegetation. The disturbed habitats that revegetation in is only superficial. Large parts were formerly

destroyed or disturbed are covered by a dense growth ofLeucaena leucocephala, an exotic shrub introduced from South Feral wild and African snail abundant America. goatruns is

on some islands.

The undisturbed natural forest is restricted inaccessible of the islands. It is to parts com- posed of Cinnamomum pseudopedunclatum *, Ligustrum micranthum'»*. Pittosporum bonitiense*,

Raphiolepis integerrima, Evodia kumagaianaJ*, Distylium lepidotum»*. Boninia glabra»*, Hibiscus

tiliaceus, Syzygium buxifolium, Rapanea maximowiczii<*, Schima mertensiii*. Hibiscus glaberr*.

Ardisia sieboldii, Livistonia boninensiss*, Osmanthus insularis, Pouteria obovata, Calophyllum

Morus inophyllum, Hernandia sonora, kagayamae, Machilus kobu<*, Sideroxylon ferrugineum,

￿ Pandanus boninensis Freycinetia formosana var. boninensis, Trachelospermum foetidumI*, etc. » indicated endemic Floristic (The species by an asterisk are to the Ogasawara Region).

There are two kinds ofnatural communities on windy narrow ridges. One is composed

of Miscanthus boninensisr*. Trachelospermum foetidum>*, Carex hattoriana, Carex bongardii, and others. The Another some community is developed on shallow soils on the ridge. is of lanata and and is almost made up Osteomeles Fimbrystylis diphylla found only on the exposed welded tuff.

Plants on sand beaches are arranged from Ipomoea pes-caprae subsp. brasiliensis at the

foreshore, through Cynodon dactylon and Vitex rotundifolia, toScaevola taccada.The disturb-

ed island slope is covered by Leucaena leucocephala, Miscanthus boninensis, Cymbopogon

angustispica, etc.

1.2. Semi-naturel Vegetation

As stated before, natural forests in the region have been extensively cut over and the various of lands now support types vegetation such as secondary forests, grasslands, weed

etc. the and communities, In present part, secondary forests grasslands are treated.

1.2.1. Secondary forests

1.2.1.1. Coppices of Castanopsis cuspidata and C. sieboldii of have been Coppices Castanopsis are found in forest lands that periodically logged at several decadeintervals data still (Itow, 1968a; Yamanaka, 1968). Though are inadequate, M. Numata et a\.\ Natural and semi-natural vegetation in Japan 453

such stand a appears to be developed in warmer areas of the evergreen broad-leaved

forest region, to which Castanopsis is well adapted. The floristic composition of the forest

almost as of the natural forest of the same of this type is the same that species, except for shade-intolerant Mallotus the occurrence of such trees as japonicus, Albizia julibrissin,

Zanthoxylum ailanthoides, etc. Physiognomically, the sprout-origin coppice is different from the natural low and one in its stature in an even-aged canopy. of edulis Sprout-origin forests Pasania and of Quercus glauca are found, at least, in part

but the details are known. of Kyushu, not yet

1.2.1.2. Pinus densiflora forests. Phot. n.

Natural stands ofPinus densiflora are restrictedto extreme habitats such as narrow ridges,

and lava flows with little soils and of steep slopes, or exposed bedrocks, as peripheries fens The those habitats and and swamps. growth in is usually sparse stunted. On the

other forests of this not the hand, secondary pine are very extensive only in evergreen

broad-leavedforest region (except on islands south of Yakushima), but also in the summer-

green broad-leaved forest region (except on Hokkaido). P. densiflora is now one of the in and its most prominent conifers inhabited regions of Japan, forests are developed on lands that oak originally supported Castanopsis-Machilus forests, evergreen forests, and

Abiesfirma forests. (Pine forests both in the evergreen and the summergreen forest regions

are treated together here.) The has been intervals pine forest long maintained by logging at 15—50 year and by disturbance of the forest floor vegetation.

The floristic composition and the structure differ, more or less, with regions, localities, after disturbance the years the latest logging, and intensity of on forest floor. Based on of the data an analysis in Yoshioka (1958), which were gathered throughout Japan, the species common in forests are Pinus densiflora, Quercus serrata, Castanea crenata, Clethra

Vaccinium and Miscanthus barbinervis, oldhamii, and Ilex crenata, herbs as sinensis, Solidago and Pteridium The above-listed virga-aurea var. asiatica, aquilinum. trees except the pine

are deciduous and shade-intolerant and the last three herbs are of the components grass- land differences vegetation. Regional in the composition are as follows. The forests in the broad-leaved forest of other deciduous summergreen region consist many trees:

Quercus mongolica var. grosseserrata, Sorbus alnifolia, Ilex macropoda, Magnolia obovata, which also beech forests and Fraxinus lanuginosus, occur in which are not or rarely found the forest of the the in P. densiflora evergreen broad-leaved forest region. On other hand,

the following tree species are largely restricted to the forests in the latter region: Eurya

laponica, Juniperus rigida, Vaccinium bracteatum, Quercus glauca, Q. salicina, Ligustrum

laponicum, Ilex chinensis, Castanopsis sieboldii, and Camellia japonica. These species, except for ofthe natural before. Juniperus rigida, are major components evergreen forest, as stated is certain will the It that the pine forest be replaced by summergreen forests in former region and evergreen ones in the latter, ifkept free long enough from human interference.

The floor vegetation of die forest in the latter region is frequently dominated by either central Pleioblastus chino (in Honshu), P. distichus var. nezasa (in western Honshu, Shikoku, and Gleichenia stands the Kyushu), japonica, or Gleichenia dichotoma. At some in Inland Sea region, bare condition of the floor has resulted from repeated gathering of fallen leaves and of the floor associated with condition of the clearing vegetation, dry region. Such a

forest has suffered of erosion and is so deteriorated that the habitat is not potentially the forest capable any more to support Castanopsis that originally was there. of the the The floristic characteristic pine forest is occurrence of Rhododendron species. is in Rh. kaempferi one of the most widely distributed species the pine forests ranging 454 BLUMEA VOL. XX, No. 2, 1972

from northern Honshu to Kyushu. Besides this, Rh. macrosepalum, Rh. weyrichii, and Rh.

the Sea of the kiusianum are characteristic of pine forest of the eastern Inland region, southern half of Kii Peninsula, Shikoku, and Kyushu, and of Mt. Kirishima in Kyushu, respectively (Suzuki, 1966b).

Mixed and 1.2.1.3. forests of Quercus serrata Quercus acutissima (deciduous oaks) of and of the Forests mixed Quercus serrata Q. acutissima are developed on part lands

that originally supported evergreen broad-leaved forests. The forest of the deciduous

oaks has and such been maintained by logging at 15 to 25 year intervals, is composed of

deciduous trees as Prunus jamazakura, Castanea crenata, Clethra barbinervis, Styrax japonica, made of Carpinus tschonoskii, etc. The floor vegetation oftheforest is up Miscanthus sinensis,

western Pleioblastus chino (in central Honshu) or P. distichus var. nezasa (in Honshu), Aster

Potentilla which scaber, Solidago virga-aurea var. asiatica, freyniana, Ixeris dentata, are com- the Miscanthus and ponents ot sinensis grassland (Miyawaki Fujiwara, 1968a; Miyawaki and of broad- et al., 1968; Miyawaki, 1969). In addition, seedlings saplings evergreen the This leaved trees are usually found on forest floor, even though they are sparse. means habitat ofthe deciduousoak forests the forest of that the is potential to support evergreen trees. Those found in the deciduous oak forest in the Machilus-Castanopsis forest region Honshu Aucuba in central are japonica, Machilus thunbergii, Castanopsis cuspidata, C. sieboldii,

Neolitsea sericea, Cinnamomumjaponicum, Dendropanax trifidus, while those in the Quercus

myrsinaefolia (evergreen oak) forest region are Quercus myrsinaefolia, Q. glauca, Eurya If be the japonica. logging would ever ceased, deciduous oak forest will surely be replaced that the climax the by the evergreen broad-leaved forest is climatic of region (Miyawaki, 1967).

Bamboo Phot. 1.2.1.4. forests. 18. There dominated are many bamboo stands, cultivated or semi-natural, mainly by

of and three species Phyllostachys (Ph. bambusoides, Ph. pubescens, Ph. nigra var. henonis).

Phot. 18. The semi-naturalbamboo brakes a serai which the belong to stage converges in

climaxes Shiion sieboldii and a part ofFagion crenatae (Numata, 1955; Ueda and Numata,

1961; Numata and Aoki, 1962; Numata, 1965a). This is judged by the dynamics of

of the east or the ground vegetation Japan type west Japan type. For example, ground flora of the former is composed of Carex lanceolata, Oplismenus undulatifolius, Pleioblastus

Wistaria Viburnum and chino, brachybotrys, erosum, Disporum sessile, Huouttuynia cordata, etc.,

thatof the latter Nandinadomestica, Aucubajaponica, Camellia japonica, Ophiopogon japonicus,

Callicarpa japonica, Dryopteris lacera, Ligustrum japonicum, Torreya nucifera, Eurya japonica,

Asplenium incisum, Polystichopsis pseudoaristata, etc.

1.2.2. Grasslands

Grasslands are described later together with those in the summergreen broad-leaved

forest region.

2. Summergreen broad-leaved forest region. Table 6 —II. Phot. 4, 9.

As shown in the vegetation map (Fig. 2), the summergreen broad-leaved forest region and occupies the montane area in central and southern Japan, and montane to foothill

lowland areas in northern Honshu and Hokkaido. A large number ofnew varieties ofrice

plant adapted to particular areas or to other special conditions have been produced as a result the has of breeding program. Especially, the rice acreage expanded northwards thanks cooler climates result of to new varieties adapted to (Japan FAO Ass., 1958). As a M. Numata et al.: Natural and semi-tiatural vegetation in Japan 455

population incrementin the region, natural habitats have become narrower, while man-

made and man-influenced habitats wider. Nevertheless, the region under consideration least was sparsely populated as a whole and the natural vegetation was less touched, at before World with the broad-leaved forest Since War II, as compared evergreen region.

then, however, the encroachment is advancing further to back countries and cutover

forest lands are reforested uniformly with coniferous trees such as Cryptomeria japonica,

Chamaecyparis obtusa, and Larix leptolepis.

Natural 2.1. Vegetation

2.1.i. Beech forests. Phot. 2.

of the The beech forest is the climatic climax that occupies large parts region. Main

the forest are such deciduous trees components common in in Japan as Fagus crenata, Tilia Kalopanax septemlobus, japonica, Quercus mongolica var. grosseserrata, Magnolia obovata,

Acanthopanax sciadophylloides, Fraxinus sieboldiana, Sorbus alnifolia, Acer mono, A. palmatum shrubs Rhus vines var. matsumurae;; deciduous as Viburnum furcatum, trichocarpa; and as

Schizophragma hydrangeoides, Rhus ambigua, Hydrangea petiolaris, etc. stated the differs Pacific the As has been before, climate with regions on the side and on side. Beech forests subdivided into which these Japan Sea are also two major types occupy

Such difference in the beech forest was out two regions, respectively. a firstly pointed

by Suzuki (1952). Later, Sasaki (1964) confirmed it and studied forests in detail throughout

Japan. kurilensis It TheFagus crenata-Sasa forest is developed in regions on the Japan Sea side. is characterized by Sasa kurilensis, Acer japonicum, Hugeria japonica, Oxalis acetocella var. japonica, Mitchella undulata, Rumohra mutica, Plagyogyria matsumureana, Aucuba japonica

var. borealis, Daphniphyllum macropodum var humile, Cephalotaxus harringtonia var. nana,

Skimmia japonica var. repens. Those in species are absent, or very rare, the Fagus crenata-Sasamorpha purpurascens Pacific side. Sasa kurilensis and the last four ofthe forest that is developed in regions on the shrubs. have in the above-listed species are evergreen They an ecological advantage

after melts crenata- photosynthetic activity soon the accumulated snow away. Fagus forests characterized Sasamorpha purpurascens (phot. 2) are by Sasamorpha purpurascens, Betula Lindera Acer sieboldiana, A. shirasawanum, A. micranthum, grossa, Symplocos coreana,

umbellata, Stewartia pseudo-camellia, S. monadelpha, Fagus japonica, Parabenzoin trilobum,

Sasa nipponica, Abies homolepis, Tsuga sieboldii, and Skimmia japonica (Sasaki, 1964). beech forest the Phytosociologically, the two types of the were first regarded as asso- ciations: Saseto- and Sasamorpheto-Fagetum crenatae (Suzuki, 1952, 1966b; Sasaki, 1964). each But recent detailed studies revealed several associations which are distinctive from

the other in theirfloristic composition and ecological relation to habitat(Miyawaki et ah,

1964, 1968, 1969). The Corno-Fagetum crenatae is found on mountain slopes with rather shallow soils and the Miricacalio-Fagetum crenatae is developed in the cloud zone with in high air humidity in Mts. Tanzawa, Fuji, and Hakone, which are situated regions on

the Pacific side of central Honshu (Miyawaki et ah, 1964, 1968, 1969). In the Okutadami central the other Aucubo- and Hamamelo- region on the Japan Sea side, Honshu, on hand, Fagetum crenatae have developed in low and high elevations of the beech forest zone the respectively (Miyawaki et ah, 1968). Based on these detailed studies, major two types and of the beech forest in Japan, which were regarded as associations, viz. Saseto- Sasa- be alliances: Saseto- and morpheto-Fagetum crenatae, might treated as Sasamorpheto-Fagion

which the of the Sea side and of the Pacific side crenatae, occupy montane regions Japan respectively, and to which several associations belong (Miyawaki et ah, 1964, 1967, 1968,

1969). 456 BLUMEA VOL. XX, No. 2, 1972

8. the the Fig. Map of Japan showing range of (I) Fagus crenata-Sasa kurilensis forest and (II) the Fagus forest. Detailed crenata-Sasamorpha purpurascens explanation in text (after Sasaki in Miyawaki, 1967).

Geographical variation in the floristic composition of beech forest is shown in Fig. 8

in The I in 8 is the of the crenata-Sasa (Sasaki, Miyawaki, 1967). Region Fig. range Fagus

Beech la characterized kurilensis forest ( Saseto-Fagion crenatae). forests in are by Sasa cernua, those Hamamelis Hydrangea macrophylla var. megacarpa and Pachysandra terminalis; in lb by japonica var. obtusata, Rhododendron albrechtii, and Carex morrowii var. temnolepis. Beech forests in Ic show a transitional composition between the Regions I and II, and are charac- Lindera and terized by Cryptomeria japonica var. radians, umbellata, Symplocos coreana, The II in is of F. Rubus pectinelleus. Region Fig. 8 the range the crenata-Sasamorpha pur- the Pacific side. Beech purascens forest ( Sasamorpheto-Fagion crenatae) on forests in Ila are the lib typical type of the beech forest on thePacific side; those in are characterized by Abies homolepis; and those in lie by Parabenzoin trilobum, Rhododendron metternichii, and

Enkianthus cernuus (. rubens (Sasaki, in Miyawaki, 1967).

of is in Fagus japonica is a minor counterpart F. crenata. It distributed montane regions Pacific side. found in limited scale only on the Forests ofF. japonica are infrequently a (cf. Tohyama, 1965). M. Numata et al.: Natural and semi-natural vegetation in Japan 457

2.1.2. P t e r o c a r y a rhoifolia forests

Moist habitats in bottoms and alluvial fans in the forest valley on summergreen region forests. Such habitats surrounded beech forests unless the support Pterocarya rhoifolia are by

area and soils well drained and made of and sands is disturbed, are up boulders, gravels,

with decomposed organic matter (Saito, 1967). Common plants of the Pterocarya rhoifolia forest are Ulmus laciniata, Cercidiphyllum japonicum, Phellodendron amurense, Pachysandra

Stellaria are terminalis, diversiflorum, Athyrium pycnosorus, Matteuccia struthiopteris (which Sorhus characteristic to the P. rhoifolia forest), Acer mono, alnifolia, Ligustrum tschonoskii, Rhus ambigua, Schizophragma hydrangeoides, Hydrangea petiolaris, Asperula odorata, Galium

japonicum, Laportea bulbifera, Dryopteris crassirhizoma, etc. (which are common, more or of the less, to some summergreen forests in the region).

Phytosociologically, P. rhoifolia forests are subdivided into three associations. One is

the Polysticheto-Pterocaryetum (Suzuki et al., 1956; Miyawaki et al., 1968). It is mostly Aesculus distributed in regions on the Japan Sea side and characterized by turbinata, Laportea macrostachya, Viola vaginata, Polystichum retroso-paleaceum, P. tripteron, Dryopteris

monticola. The second association is Dryopteridi-Fraxinetum commemoralis (Suzuki, 1952),

which with et is synonymous Chrysosplenio-Fraxinetum spaethianae (Miyawaki al., 1964).

This is distributed in regions on thePacific side and is characterized by Fraxinus spaethiana,

Acer carpinifolium, A. diabolicium, A. argutum, Scutellaria shikokiana, Chrysoplenium macro-

stemon, Cardamine anemonoides, Asarum caulescens and Dryopteris polylepis. The third is

from western Honshu Polysticheto-Aesculetum turbinatae, which was reported (Horikawa

and Sasaki, 1959a). The species composition is similar to the first association.

Schematic of distribution in relation latitude and soil moisture Fig. 9. summary plant community to in the broad-leaved forest of Forests the side gradient summergreen region Japan. on Japan Sea are given and onleft half; the Pacific side, right half; Hokkaido, above the broken line; and Honshu, Shikoku, Kyushu,

below. A. Pinus densijlora forest, B. Abies firma-Tsuga sieboldii forest, C. Tsuga diversifolia forest, D. Quercus

E. crenata-Sasa F. mongolica var. grosseserrata forest, Fagus kurilensis forest, Fagus crenata-Sasamorpha purpu-

rascens forest, G. Pterocarya rhoifolia-Polystichum tripteron forest, H.Fraxinus spaethiana-Pterocarya rhoifolia-

K. I. Dryopteris polylepis forest, Quercus mongolica var. undulatifolia forest, Salix-Alnus hirsuta forest, J.

Phragmites communis moorland (fen), L. Picea glehnii forest, M. Quercus tnongolica var. grosseserrata-Quercus dentata forest, N. Fraxinus mandshurica-Ulmus davidiana forest, O. Sphagnum bog (after Miyawaki, 1967). VOL. No. 458 BLUMEA XX, 2, 1972

Successive in and Fig. 10. changes coverage (abcissa) height class (ordinate) of the important taxa com- posing the plant community on the central part of the Akaikawa pumice flow. A. (trees): I. Betula platy- Larix phylla var. japonica, 2. Populus maximowicziana, 3. leptolepis, 4. Pinus densiflora, 5. Quercus mongolica

B. 6. C. 8. var. grosseserrata ; (subtrees): Salix bakko, 7. Salix sachalinensis; (shrubs): Salix integra, 9. Hydrangea D. paniculata; (herbs): 10. Polygonum sachalinensis, 11. Dryopteris rhizophragma; E. (mosses and lichens): 12.

Racomitrium vesvianum Polystichum commune, 13. canescens, 14. Stereocaulon (after Yoshioka, 1966). M. Numata el a\.\ Natural and semi-natural vegetation in Japan 459

Ulmus d d a r a x n u s mandshurica forests 2.1.3- a v i i n a-F i

Habitats with a high ground water tablein lowlands ofnorthern Honshu and Hokkaido forests that different from the above-mentioned support deciduous are ones. One is the Alnus forest. and japonica It is developed on the periphery of fens and reed swamps on

flood deposits along the lower river courses. The ground water table of those habitats is

and than the level. If the water table as high as in some places even higher ground is lower,

Alnus japonica is mixed with Fraxinus mandshurica and futher replaced by Ulmus davidiana.

Based on our observations of relict stands, alluvial plains where cities and agricultural

lands are now extensive originally supported the mixed forest of Ulmus davidiana and

mandshurica. Main of and Fraxinus components the forest are such deciduous trees shrubs

as Ulmus laciniata, Pterocarya rhoifolia, Acer mono, Prunus maximowiczii, Syringa reticulata, Trillium Euonymus oxyphyllus, E. macropterus, Sambucus sieboldiana var. miquelii; herbs as kamtschaticum, Fritillaria camtschatcensis, Meehania urticifolia, Scopolia japonica, Anemone

flaccida, Urtica platyphylla, Impatiens noli-tangere, Cardamine leucantha, Filipendula kam-

tschatica, Cacalia hastata var. orientalis, Veratrum grandiflorum, Maianthemum dilatatum, Carex

dissitiflora, C. parciflora, etc. (Miyawaki, 1967).

2.1.4. Volcanic vegetation. Phot. 4, 25, 26. Several and volcanoesin the broad-leaved forest extinct active summergreen region are

good fields for observing the primary succession on volcanic habitats. An example is Mt. southermost Hokkaido. took Komagatake, which is situated in An eruption place on it

in 1929 and the vegetation on the slopes was damaged, and, on some slopes, was com-

and ash. After the the pletely covered by pumice eruption, development and recovery flow Yoshii Yoshioka of vegetation on a new pumice were studied by and in 1933, 1935,

1938,1942,and 1948, and by Yoshioka in i960and 1965. The details reported by Yoshioka

(1966) are given in Fig. 10.

Vegetational development on lava can be observed on the andesitic lava bed dating

the northeast of Mt. central Honshu. Phot. The from 1783 on slope Asama, 26. plant the bed still and low. growth on (1350 m in altitude) is quite sparse Species observed there Salix S. are Hydrangea paniculata, vulpina, bakko, Betula platyphylla var. japonica, Polygonum cuspidatum, Alnus firma, Larix leptolepis, Pinus densiflora, Tripetaleia paniculata, Loiseleuria Vaccinium uliginosum, procumbens, Empetrum nigrum var. japonicum, Cassiope The of the lycopodioides, Phyllodoce nipponica, P. aleutica, etc. first five above-listed species above-cited flow The last are common to the pumice on Mt. Komagatake. six species

that in elevations in this Their are alpine plants usually grow high exceeding 2300 m area.

to occurrence in the cool-temperate zone is extraordinary and seems result from the

severe conditions on the lava flow that resembles conditions in the alpine zone. habitats with loose and On the scoria, sand, ash are developed herbaceous communities.

Such habitats in the montane, subalpine, and alpine areas on Mt. Fuji support three asso-

ciations; viz. the Carex stenantha-Stellaria nipponica Ass., Cirsium purpuratum- Campanula and the punctata var. hondoensis Ass., Angelica hakonensis-Miscanthus oligostachyus Ass. The

first and second associations are developed on rather unstable substrata, while the third stable habitats with dense of found these occurs on growth plants. Species commonly in Arabis associations are serrata var. serrata, Artemisia pedunculosa, Astragalus adsurgens.

Polygonum cuspidata, Salix reinii, Carex doenitzii, Hedysarum vicioides, Larix leptolepis, and The of these the Polygonum weyrichii var. alpinum. last species grows alone on most

unstable habitats in high elevations (Miyawaki et al., 1967). those Plant communitiesaround active solfataras differ from just mentioned. Phot. 25.

The habitat is characterized by ofC0 S0 H S, etc. and the soil is acidic. Obser- gases 2 , 2 , 2 460 BLUMEA VOL. XX, No. 2, 1972

topographic are Linaria the and on species Lathyrus, dominant Calystegia,

the 1962). indicate Ischaemum, Ishizuka, names trends. (after plant

The successional bar. the Ischaetnum-group sand of

the types the

as on the here system indicate called C

and B, collectively topography-vegetation A, capitals

the The

of symbols. developmentby

The denoted

II. units Fig. M. Numata et al.: Natural and semi-natural vegetation in Japan 461

made on Hakkoda and northern Honshu vations Mts. Osoreyama, (Yoahioka 1951; al. indicatethat surface soils solfataras Yoshioka et 1965) near are extremely acidic ranging from in which found. 1.5 to 3.8 pH value, on no plants are Soils supporting plant growth

also are from to value. Plants such habitats acido- acidic, 2.0 4.5 in pH growing on are

philic or acid-tolerant. Examples observed on Mt. Osoreyama are Haplozia crenulata var. and gracillima, Drepanocladus fluitans, Carex angustisquama. These three species are appar- the observed ently acidophilic. In addition, following species are there: Deschampsia Miscanthus flexuosa, Juncus brachyspathus, Moliniopsis japonica, sinensis, Eubotryoides grayana

Ilex crenata var. grayana, and var. paludosa. Scrub communities in a distance from the

solfatara are composed of Ledum palustre var. diversipilosum, Ilex crenata var. paludosa,

Vaccinium smallii, Rhododendron fauriei, Hydrangea paniculata, Ilex sugerokii var. brevipe- Sorbus dunculata, Sasa paniculata, S. kurilensis, Eubotryoides grayana var. grayana, commixta,

Viburnum furcatum, etc. Among the species listed above, Deschampsia flexuosa, Ledum and palustre var. diversipilosum, Rhododendron fauriei, Ilex sugerokii var. brevipedunculata

are those whose distribution is usually restricted to the subalpine and alpine zones. Their of occurrence around the solfataras results apparently from the severe conditions the

habitat. The same is true of the solfatara area on Mt. Hakkoda (Yoshioka, 1951).

2.1.5. Coastal vegetation. Phot. 17. Sand dunes and sand bars. Phot. 2.1.5.1. 23. Plant communities coastal zonation of on sand dunes vary with habitats. Typical the dune understood komarovii communities is as ranging from (1) Salsola on nitrophytic with habitats decomposed tidal drifts, through (2) sparse communities on unstable slopes

and with loose sand that consist of Carex crests kobomugi, Elymus mollis, Ixeris repens,

Glehnia littoralis, Calystegia soldanella, Linaria japonica, and Carex macrocephala, to (3) the shrub community of Rosa rugosa and Malus baccata var. mandshurica, and further to (4)

Quercus dentata forests on fixed dunes (Miyawaki, 1967).

On sand outward the bars or spits extending into open sea, various types of maritime

communities and submaritime plant are found (Ishizuka, 1961, 1962). The interspecific the of there well with associations or disassociations major species are correlated similarity those of or dissimilarity among topographic-distributional patterns species. Three groups can be recognized as positive associations, such as 1) Suaeda maritima, 2)

Elymus mollis and Carex pumila, and 3) Ixeris repens, Carex kobomugi, Carex macrocephala, Glehnia littoralis, Ischaemum anthephoroides, Calystegia soldanella, Lathyrus maritimus, and

Linaria japonica.

The interrelation between the vegetation and the dune topography on sand bars is

a of mutual and regarded as system interactions development (Fig. 11).

2.1.5.2. Salt marshes. Phot. 24.

Salt natural condition the eastern coasts ofHokkaido marshes in a remain abundantly on

(Ito, 1963). Plants and communities of the salt marshes are distributed in relation to the

of and and the of actions. The of the salinity water soils, to intensity wave arrangement

communities observed in eastern Hokkaido are schematically shown in Fig. 12, in which

the communities are named according to the phytosociological tradition (Miyawaki and The Ohba, 1965). plant distribution along the salinity gradient given on the top of communities row Fig. 12 is shown in Fig. 13. in which the sea weed are shown together. 462 BLUMEA VOL. XX, No. 2, 1972

Miyawaki

(after Hokkaido

of

coast eastern

the

on actions wave modified). and salinity slightly

to 1965,

relation Ohba,

in and communities

marsh

salt

of illustration Schematic

12.

Fig. M. Numata et al.: Naturel and semi-natural vegetation in Japan 463

in Fig. 13. Schematic illustration of plant-habitatrelationships salt marshes at the easterncoast of Hok-

Zostera Zostera Salicornia kaido. Only dominant species are shown here. 1. marina, 2. nana, 3. brachystachya, maritima 4. Puccinellia kurilensis, 5. Spergularia marina, 6. Glaux var. obtusifolia, 7. Potentilla egedei var. grandis,

8. Inula Juncus gracillimus, 9. Eleocharis kamtschatica, 10. Calamagrostis epigeios, II. japonica, 12. Phragmites communis. A. Zostera marina Ass., B. Zostera nana Ass., C. Salicornietum, D. Puccinellietum (1. typical subass., Glaux 2. subass. of maritima var. obtusifolia), E. Juncetum (1. typical subass., 2. subass. of Calamagrostisepigeios)

(after Miyawaki and Ohba, 1965, slightly modified). BLUMEA VOL. No. 464 XX, 2, 1972

1 d Phot. 2.1.6. M o o r a n vegetation. 22.

Moorlands under cool and cold climate are found abundantly in montane regions of northern Honshu and central and in Hokkaido, but infrequently at high elevations in

southern Japan (Miyawaki 1968b). Of the moorlands in Japan, those in the Ozegahara

region and its vicinity (Mts. Tashiro, Aizukoma, and Hiragatake) have been studied in

detail (Horikawa and H. Suzuki, 1954; T. Suzuki, 1954; Yoshioka, 1954; Miyawaki et al., and The deals with the 1967, 1968; Miyawaki Fujiwara, 1968b). present paper mainly these moorland community in areas.

Twenty-one species of Sphagnum have been reported from the Ozegahara region and The (Horikawa H. Suzuki, 1954). high or raised moor in the areas is made up of a

mosaic ofhummocks and hollows. Hollow plants differ with the depth of water. Hollows

with water are characterized the of limosa 5 —30 cm deep by sparse growth Carex (the

community is named Caricetum limosae, Miyawaki et al., 1967), while plants in shallow

hollows with 0—3 cm deep waterare represented by Scheuchzeriapalustris, Rhynchospora alba,

Drosera anglica (which in Honshu is found only in the Ozegahara Basin but common in

moorlands in Hokkaido), and Lycopodium inundatum (Scheuchzerio-Rhynchosporetum albae, This Miyawaki et al., 1967). community comprises constantly some other species such as and the Carex middendorffii Drosera rotundifolia; Sphagnum pulchrum frequently grows in

and the waterless and water Carex omiana in hollow (Miyawaki et al., 1967; Miyawaki

Fujiwata, 1968b). The several of hummock communities are characterized by species Sphagnum. Sph.

papillosum is characteristic to the 'young' hummock which fringes the hollow( Sphagnetum

pappilosi, Miyawaki et al., 1967, 1968; Miyawaki and Fujiwara, 1968b). This community

is distributed from low-elevation moorlands near the seacoast in Hokkaido (Miyawaki,

1968) through montane regions of Honshu to high elevations in Yakushima Island,

and southern Kyushu (Aragane, 1963). Sphagnum magellanicum Sph. compactum are repre-

sentatives of the intermediate hummock (Sphagnetum magellanici and compacti, Miyawaki

and Fujiwara, 1968); Sph. fuscum is characteristic to the mature hummock ( Sphagnetum fusci, Miyawaki et al., 1967). These four hummock communities comprise other common

bog plants such as Oxycoccus quadripetalus, Andromeda polifolia, Rhynchospora alba, Carex

middendorffii, C. omiana, Moliniopsis japonica, Drosera rotundifolia, Tofieldia japonica, Eriopho-

rum vaginatum, etc. Sphagnum ambryphyllum, Sph. rubellum, and Sph. pulchrum are infre- The and hummock make quently found in these communities. hollow communities up of the complex mosaic the high moor vegetation. The of Moliniopsis japonica-Carex middendorffii community is representative the tran-

sition moor of Japan. It is named Carici-Moliniopsietum japonicae (Miyawaki et al., 1967,

1968). Physiognomically, this community is different from the above-stated high moor because of the dense of communities growth such tall prominent herbs as Hemerocallis

middendorffii, Solidago virga-aurea var. asiatica, Sanguisorba officinalis, Eriophorum vaginatum,

Carex Myrica gale var. tomentosa, Moliniopsis japonica, and middenddorffii. Heloniopsis orien-

talis Drosera and also found rotundifolia, Oxycoccus quadripetalus, Pogonia japonica are com- the of the moorland of this dominated monly in transition moor. Some parts type are by Osmunda one of following species: Sphagnum capillaceum, Sph. ambryphyllum, Sasa oseana,

asiatica, Tofieldia japonica, Lysichiton camtschatcense, and Hosta albo-marginata (Miyawaki et

al., 1967; Miyawaki and Fujiwara, 1968b). the moorland Bog pools in support an aquatic community. It is composed of Comarum

palustre, Menyanthes trifolia, Caltha palustris var. membranacea, Carex limosa, Lobelia sessili- folia, Equisetum limosum f. limosum, and E. limosum (. verticillata. The community is named Comareto-Menyanthetum (Suzuki, 1954). M. Numata et a\.\ Natural and semi-natural vegetation in Japan 465

The high and transition moorland communities mentioned above are expected to be

foundin Hokkaido (Ito and Tohyama, 1968). Ledum palustre var. diversipilosum is restricted

only to the latter (Miyawaki, 1968b). studies of and Ecological peat bogs (Yoshii Yoshioka, 1940; Iwata, 1940, 1941; Suzuki,

1941; Jimbo, 1941, 1942, 1948; Nakamura, 1942, 1949; Ishizuka, 1949) clarified aquatic and of lake flora vegetation, zonation and succession, pollen analysis deposits and peat, microflora, and environmental conditions.

2.2. Semi-natural vegetation

S d forests. Table 2.2.1. e c o n a r y 7

2.2.1.1. Pinus densiflora forests. Phot. n.

of before Secondary forests Pinus densiflora are treated under the evergreen broad- leaved forest region.

2.2.1.2. Quercus mangolica var. grosseserrata-Castanea crenata forests

Secondary forests of mixed Quercus serrata and Q. acutissima are not developed in the while the found summergreen broad-leaved forest region, Quercus serrata forests are in the lower montane areas in region (Ishizuka, 1968). So far as we know, the latter is different from the former in lacking, at least, Q. acutissima and seedlings and saplings of and shrub evergreen tree species. However, many species are common to both. The forest Quercus mongolica var. grosseserrata-Castanea crenata is representative of in secondary forests the region underconsideration. They are, more or less, varied with localities habitats. different and Components common to such forests in localities are Rhus Quercus mongolica var. grosseserrata, Castanea crenata, trichocarpa, Acer palmatum var.

matsumurae, A. mono, Ilex macropoda, Clethra barbinervis, Styrax japonica, Viburnumfurcatum,

Sorbus alnifolia, S. japonica, Cornus kousa, Carpinus tschonoskii, C. japonica, Acanthopanax

sciadophylloides, etc.

2.2.2. Grasslands. Phot. 19—21.

Grasslands of the warm- and cool-temperate and subarctic regions are treated here

together. They are divided into two categories in terms of their origin. One is the

natural grassland; another is the secondary one.

The natural grassland is developed in the alpine zone and on windy ridges on which the

growth of trees is prevented and on volcanic habitats. Natural grasslands in northern

Honshu and Hokkaido dominated Sasa S. S. are by kurilensis, paniculata, nipponica,, or nikkoensis Those and S. (Oshima, 1961). in central western Japan are characteristic in of allied restricted accompanying species Rhododendron or genera, and are to high windy ridges. Ericaceous species are Rhododendron tsusiophyllum on Mts. Tanzawa and Hakone

(extinct volcanoes) in central Honshu (Miyawaki et ah, 1964; 1969), Gaultheria adenothrix

in western Honshu (Horikawa and Sasaki, 1959), Rhododendron tschonoskii in Shikoku Rh. and (Yamanaka, 1964), kiusianum on Mts. Kuzyu, Aso, Unzen (active volcanoes) in

Kyushu (Suzuki and Abe, 1959), and Rh. metternichii var. yakushimanum on Yaku-shima Island.

The is and disturbed maintained secondary grassland developed on deforested areas, and is

by mowing, grazing, or burning. Dominance-types categorized by dominant (and sub-

dominant) species are: Sasa-, Miscanthus sinensis-, M. sinensis-Pleioblastus-, Zoysia japonica-,

Z. japonica-Pleioblastus-, Pleioblastus-, Pteridium aquilinum-, and Poa pratensis-types.

The Sflsa-type secondary grassland is mainly distributed in northern Honshu and Dominant Sasa S. S. Hokkaido. species are kurilensis, paniculata, or nipponica. As is well 466 BLUMEA VOL. XX, No. 2, 1972

abounds of the known, the Japanese Archipelago in species genus Sasa. Most of these are

associated withforest communities as their undergrowth in the cool-temperate, subalpine,

and subarctic region. When timbers of the forest are logged down, there remains a dense

almost ofthe growth of Sasa species which cover completely the logged forest land. Most and Sasa-type grasslands except on windy ridges on volcanic habitats result from such disturbance. The and function of structure Sasa communities are recently analysed from

the standpoint of production ecology (Oshima, 1961, 1962).

The Miscanthus sinensis is grassland (phot. 19) maintained by yearly or every two-year distributed the subarctic mowing with frequent or infrequent burning. It is mainly in and

and warm- and cool-temperate regions from Hokkaido, through Honshu Shikoku, to

Kyushu (and further south to the Ryukyu Islands). The grassland is not natural but a serai If free from human community, as stated before. kept long interference, it is replaced

by shrublands and successively by forests. Main components of the Miscanthus sinensis

such Arundinella shrubs grassland are grasses as M. sinensis, hirta, Spodiopogon sibiricus; as

Lespedeza cyrtobotrya, L. bicolor, L. cuneata, Smilax china, Salix vulpina; tall herbs as Aster

scaber, Solidago virga-aurea var. asiatica, Lysimachia clethroides, Cirsium japonicum, Patrinia

scabiosaefolia, Eupatorium lindleyanum, Platycodon grandiflorum, Pteridium aquilinum, Adetto- and small phora triphylla var. japonica; herbs as Carex lanceolata, Potentilla freyniana, Ixeris

Viola etc. dentata, Disporum smilacinum, grypoceras, (Yoshioka, 1955; Itow, 1963; Numata,

1966; Miyawaki and Fujiwara, 1968a; Miyawaki et al., 1968). The another that distributed Zoysia japonica grassland (phot. 20) is type is widely in Japan. It is developed only in the continuously grazed habitat, and is distributed from southwestern Hokkaido the north the south. is sod at to Kyushu at Zoysia japonica a grass and forms extensive turfs by vegetative reproduction of rhizomes and stolons. Herbaceous Miscanthus plants found in the sinensis grassland also occur frequently or infrequently in

the Zoysia japonica grassland in a reduced, stunted, and injured condition. In addition, the common species of grassland are such grazing- and trampling-tolerant perennials as

Haloragis micrantha, Polygala japonica, Carex nervata, Lotus corniculatus, Luzula capitata,

Geranium nepalense var. thunbergii, Agrostis clavata, Hydrocotyle ramiflora, Plantago asiatica, and annual Trifolium repens; plants as Poa annua, Cerastium holosteoides var. hallaisanense,

Mazus miquelii, Kummerowia striata, etc. (Yoshioka, 1955; Itow, 1963; Suganuma, 1966).

sites in the weed of Heavily trampled grassland support a community consisting Plantago

asiatica, Poa annua, Trifolium repens, and some other weeds (Iizumi, 1962; Itow, 1963).

is The community similar to that on trampled trails (Numata, 1961a; Miyawaki, 1964). the Phytosociologically, Zoysia japonica grasslands were classified into four associations:

Violo-, Geranio-, Erigero-, and Arundinello-Zoysietum (Suganuma, 1966). Our data from

in association is lowland grasslands Kyushu suggest a new which developed on coastal grazed lands.

The Pleioblastus-type and its allied types (Miscanthus sinensis-Pleioblastus- and Zoysia

in central Honshu chino japonica-Pleioblastus-types) are distributed (where Pleioblastus Honshu occurs), and in western and Kyuhsu (where P. distichus var. nezasa occurs). They

are prevailing on volcanic ash soils in the Mts. Kuzyu, Aso, and Kirishima areas of

central Kyuhsu (Itow, 1968b) and in the Mt. Sanbe area of western Honshu (Suganuma, central 1966); and on non-volcanic soils in western and Honshu (Okuda and Miyawaki,

of these 1966; Miyawaki and Fujiwara, 1968b). Grasslands types are mostly, but not

exclusively, distributed in the warm-temperate region. The floristic composition is

similar to the Miscanthus sinensis grassland whenit is yearly mowed, and to the Zoysia

japonica grassland when grazed. of Miscanthus sinensis while it Pteridium aquilinum occurs as a component grasslands, M. Numata et ah: Natural and semi-natural vegetation in Japan 467

frequently grows very densely within or contiguous to the M. sinensis and Zoysia japonica grasslands. The physiognomy is quite different from these, but the floristic composition of the Miscanthus is almost the same as that sinensis grassland when the bracken community is developed within or contiguous to it (Itow, 1963), and the same as that of the Zoysia

when it is within japonica grassland or contiguous to the latter (Suganuma, 1966).

the sown and fertilized careful The Poa pratensis grassland is pasture. It is maintainedby management, mostly in northern Japan (Numata, 1961a). Poa pratensis was introduced from in the 20th and is established semi-natural Europe early century now as a good under of Hokkaido. The pasture proper intensity grazing, especially in pasture is phyto- sociologically named Phleum pratense-Poa pratensis association (Miyawaki, 1962).

As stated before, secondary grasslands are developed on deforested and disturbed areas and shrubs and maintained by mowing, grazing or burning. They are invaded by trees, if kept long free from human interference. Succession of grasslands was first studied by Oseko and later several authors and (1937) by (Yoshida, 1951; Sugawara Iizumi, 1954, Iizumi 1964; et al., 1957, 1961; Numata, 1961a; Itow, 1963). These studies are based on regional or local data. Stands of the grassland differ, more or less, from each other not

only in the floristic composition but also in the successional status. The problem to be

solved is the objective judgement ofthe dynamic status ofeach stand and ofeach grassland

in terms of succession. For there number ofstands ofthe type vegetational example, are a

Miscanthus sinensis grassland, ranging from those in floristic composition close to the those close the shrub the Zoysia japonica grassland to to community. To represent ecolog-

ical distance among grassland types and among stands under study, an index was pro-

posed (Numata, 1961a). It is called the 'Degree of Succession' (hereafter abbreviated as

DS), that is,

DS = [i7(l x d) / n] x v,

where 1 is the life of the d is the dominance values span component species, as expressed

by Summed Dominance Ratio (SDR, the summation percentage of the ratios of the

is in the coverage, height, frequency, density, etc.), n the number of species occurring minimal and the total The life of area, v is ground cover in percentages. span (1) plant for species is assumed as 1 for Therophytes, 10 Geophytes, Hemicryptophytes, and

Chamaephytes, 50 for Nanophanerophytes, and 100 for Micro-, Meso-, and Mega- theDS for each phanerophytes. Values of grassland type of Japan are distributed as shown

in Fig. 14 (Numata, 1969a). The index floristic DS is applicable to grassland communities, beyond climatic zones,

provinces, and continents, since it is based on the life-form of plants, not on species. This of and was proved by applications to grasslands Nepal Himalaya (Numata, 1965b)

southern Korea (Park, 1965).

forest 8. 3. Subalpine and subarctic coniferous region. Table 6—III. Phot. 7,

Subarctic, subalpine, and alpine regions of Japan are less affected by man as compared with the other treated before. of this have been regions Large parts region designated as

national in order the natural parks to preserve landscape.

Abies veitchii forests Honshu 3.1. mariesii-A. in

The subalpine region of Honshu and Shikoku supports coniferous forests dominated by Abies mariesii and/or A. veitchii. The coniferous forest is distributed from Mt. Hakkoda

in northern Honshu, where A. veitchii is lacking, through the Japan Alps in central

Honshu where A. mariesii and A. veitchii are usually mixed, to Mt. Ishizuchi where only

A. veitchii var. shikokiana is found.Altitudinal lower limit of coniferous forests is gradually

467 VOL. No. 468 BLUMEA XX, 2, 1972

Fig. 14. Curves showing the frequency of each grassland type on the DS-index gradient. 1. Erigeron is in canadensis community (which a pioneer community onabandoned fields, not treated the present paper),

distichus 2. Zoysia japonica-, 3. Pteridium aquilinum- 4. Miscanthus sinensis-, 5. Pleioblastus var. nezasa- and

P. chino-, and 6. Sasa-type grasslands (after Numata, 1969).

higher southwards, from 700 m above sea level in northern Honshu, through x 500 m at

in central Honshu, to 1900 m Shikoku (cf. vegetation map).

A stand of the coniferous forest on Mt. Tashiro (Miyawaki et al., 1967), for example,

of such and shrubs is composed trees as Abies mariesii, A. veitchii, Picea jezoensis var. hondoensis, Acer ukuruttduense, Sorbus commixta, Betula ermanii, B. coryfolia, Acer tschonoskii,

Cornus canadensis; and herbs as Coptis trifolia, Plagiogyria stenoptera, Carex dolichostachya,

Olopanax japonicus, Oxalis acetosella, Lycopodium serratum var. thunbergii, Pteridophyllum racemosum Papaveraceae an endemic genus), Tripterospermum japonicum, Smilacina yesoensis, ,

Cacalia adenostyloides, Dryopteris austriaca, Trillium smallii, Smilacina hondoensis, Clintonia The is almost the in udensis, Pyrola alpina, Diphylleia grayi, etc. same true of forests other regions in central Honshu (cf. Masamune, 1961; Suzuki, 1964).

Mixed forests of standishii and diversi- 3.2. Thuja Tsuga folia

The mixed forest of Thuja standishii and Tsuga diversifolia is another type of the conif- the with shallow erous forests in subalpine region of Honshu. It is developed on ridges soils on which blocks of the bedrock are infrequently exposed on the ground surface.

The forest is different from the Abies mariesii-A. veitchii forest in its floristic composition.

For example, the Thuja standishii-Tsuga diversifolia forest which is contiguous to the above-cited stand is lacking the last thirteen of the above-listed species and comprises such additional species as Pinus parvifolia, Ilex sugerokii var. brevipedunculata, Ilex rugosa,

Rumohra and Viburnum mutica, Rhododendron metternichii var. pentamerum, urceolatum var.

procumbens..The forest is also developed onnarrow ridges in the beech forest area (Miyawaki et al., 1967, 1968). M. Numata et al.: Natural and semi-natural vegetation in Japan 469

The above-mentioned the Abies- andthe forests found arrangement in Thuja- is usually in

the subalpine region ofHonshu (Suzuki, 1964; Suzuki et al., 1963; Miyawaki et al., 1968).

for ests. Phot. 3.3. Larix leptolepis 5. The natural distribution ofLarix leptolepis is restricted mostly to montaneand subalpine regions in centralHonshu, although the reforestation ofthis conifer has been made widely

in the summergreen broad-leaved forest region. The natural forests are fragmentarily or

extensively developed on talus and volcanic habitats whose soils are less capable in holding

water. L. leptolepis is a pioneer tree of these habitats. The floor vegetation of the forest is

varied in types and in floristic composition from site to site. According to Tatewaki et ah, the floor and (1965), types categorized by constant dominant species are Sasa-, Fern-,

Lonicera-, Malus sieboldii-, Azalea-, Rhododendron-, Grass-, Heath-, Moss-, Sapling of and Prevalent these and Azalea- conifers-, Herb-types. types among are Sasa- types in the forests below and Rhododendron- and Herb- elevations. 1900m, types in higher Major Larix Mt. Asama components of the leptolepis-Sasa paniculata forest on (1720 m), for

example, are Quercus mongolica var. grosseserrata, Rubus palmatus var. coptophyllus, Celastrus Astilbe orbiculatus, Festuca parvigluma, Artemisia montana, thunbergii var. congesta, Aquilegia those of stand buergeriana, Sanguisorba officinalis; a high-elevation in the Nikko area Maianthemum (2320 m) are Prunus ermanii, dilatatum, Polygonum cuspidatum, Solidago

and Larix decurrens, Trietttalis europea var. europea;, those of the leptolepis-Vaccinium vitis-

idaea forest are Salix reinii, Betula maximowiczii, and Pyrola incarnata.

in moist Larix leptolepis also grows naturally and wet habitats like stream-side deposits,

and marshes. The herb of such for is fens, layer a stand at Nikko, example, made up of moorland such plants as Oxycoccus quadripetalus, Andromeda polifolia, Moliniopsis japonica, Osmunda Malus asiatica, Eriophorum vaginatum, Sphagnum sp. and sieboldii (Tatewaki et ah 1966).

Thickets of 3.4. Quercus mongolica var. undulatifolia (Nanoquercetum) half of Mountains in the western northernHonshu are outstanding in lacking conif- forests the thickets dominated undulati- erous in subalpine zone; by Quercus mongolica var.

folia are developed. This is apparently because the mountains are subject to a severe and northwesterly winter monsoon to heavy snowfall which attains usually several

metres. Mts. Chokai, Zawo, Iide, Aizu-Komagatake, and Shibutsu and the Okutadami

such in their habitat. The area support a thicket, more or less, subalpine community is

very dense and about 2 m tall. It is contiguous to beech forests at its lower limit and to communities the in and the alpine at upper limit very snowy areas; community and the

coniferous forest make mosaic in mountains where wind and snowfall up a pattern are for not so severe. Main components of the Quercus mongolica var. undulatifolia thicket, in example, the Okutadami area, are Hamamelis japonica var. obtusata, Sorbus commixta, Viburnum furcatum, Menziesia multiflora, Sasa kurilensis, Maianthemum dilatatum, Vaccinium

smallii, Streptopus streptopoides var. japonicus, Plagiogyria stenoptera, Paris tetraphylla, the Clintonia udensis, Tilingia holopetala, etc. (Miyawaki et al., 1968). Fig. 15 shows ecolog- communities ical relation of the thicket with other observed in the Okutadami area,

central Honshu.

3.5. Betula ermani thickets. Phot. 7.

Betula ermani and Alnus maximowiczii are deciduous shrubs found in the subalpine and

Mixed stands of these habitats with alpine regions. or pure species are developed on 470 BLUMEA VOL. XX, No. 2, 1972

region, Okutadami

the

of zones

alpine

and subalpine, 1968). al., montane, et

the Miyawaki

in (after relationships Honshu vegetation-habitat central

of illustration Schematic

15.

Fig. M. Numata et al.: Natural and semi-natural vegetation in Japan 471

boulders and shallow soils along snow avalanche tracks and ravines (Masamune, 1961;

and volcanic Itow et ah, 1964; Suzuki, 1964) on subalpine habitats (Tohyama, 1966; the Miyawaki et ah, 1967). In many examples observed in transition zone between the the subalpine and alpine zones, the A. maximowiczi thicket is found in mesic bottom of

ravines and snow avalanche tracks; the B. ermanii thicket on the slope; and Pinus pumila, shrub the the a representative of alpine region, is on the ridge. Species common to A. and thickets Sorbus maximowiczi the B. ermanii are such shrubs as matsumurana,S. tschonoskii, Honshu and Acer ukurunduense, Rubus vernus, Weigela middendorfiana (only in northern

Hokkaido); and herbs as Glyceria alnasteretum, Streptopus amplexifolius var. papillatus,

Trautvetteria japonica, Polystichum microchlamys, Athyrium melanolepis, Dryopteris austriaca,

etc. (Ohba, in Miyawaki, 1967).

3.6. Picea jezoensis-Abies sachalinensis forests in Hok-

kaido. Phot. 1.

Prevailing coniferous forests in Hokkaido are dominated by Picea jezoensis and Abies

or Based on the dominance of the sachalinensis, accompanied by Picea glehnii not. type floor the several Prevalent vegetation, forests can be divided into types. types of the for ofcentral Hokkaido floor, example, in part (Tatewaki et ah, 1955), are Sasa paniculata-, Carex sachalinensis-, Dryopteris amurensis-types, etc. Floristic composition of these forest is relations of consid- types, however, quite alike, unless quantitative the components are

ered. Plants commonly found in the Picea jezoensis-Abies sachalinensis forest are such

trees as Betula ermanii, Sorbus commixta, shrubs as Ribes sachalinense, Euonymus macropterus, Ribes Vaccinium smallii, V. praestans, Acer ukurunduense, horridum, and herbs as Cornus

canadensis, Oxalis acetosella, Tiarella polyphylla, Maianthemum bifolium, Streptopus strepto- poides, Circaea alpina, Trillium tschonoskii, Dryopteris austriaca, Polypodium vulgare, Rumohra

mutica, Lycopodium serratum var. thunbergii, Gymnocarpium robertianum, etc.

3.7. Picea glehnii forests

Picea glehnii is distributed in Hokkaido and in southernmost Sakhalin and southern forests both habitats shallow Kuriles. Its are developed on dry with soils like volcanic

ejecta, rocky or gravelly ridges and slopes, and serpentine areas, and in wet habitats like and marshes fens, sedge swamps, (Tatewaki, 1944).

of forests on habitats are and Early stages P. glehnii serpentine sparse in physiognomy with associated Pinus pumila, Juniperus communis var. montana, Berberis amurensis, Crepis last of these characteristic the gymnopus (the two species are to area), and later stages are associated with dense undergrowth of Sasa kurilensis. The forest on rocky and gravelly

with Picea Abies The floor slopes are usually mixed yezoensis and sachalinensis. vegetation either is dominated by Rhododendron fauriei, Menziesia pentandra, Carex sachalinensis, or

mosses. Wetland forests of their Picea glehnii are various in composition. Based onthe dominants of the the forests classified floor, are into six types: Carex middendorffii, Moliniopsis japonica,

Phragmites communis, Ledum palustre var. diversipilosum, Menziesia pentandra, Lysichiton Osmunda and camtschatcense, asiatica, Sphagnum spp. (Tatewaki 1944).

Tall 3.8. herb communities. Phot. 16.

Along snow avalanche tracks and in similar habitats, a community of tall herbs is

developed in the montane, subalpine, and alpine regions. The community is distinctive

from others in its physiognomy and floristic composition. Several examples are given below. Phot. 8. A community observed on Mt. Hakkoda (Yoshioka, 1948) is composed of 472 BLUMEA VOL. XX, No. 2, 1972

16. Schematic Pinus Fig. illustration of community distribution on analpine ridge. I. pumila community;

2 and 3. wind-blown heathland, 2. Arcterio-Loiseleurietum, 3. Kobresia bellardii community; 4. snow-patch

in community. Broken line indicates the snow accumulation winter (after Ohba in Miyawaki, 1967, slightly modified).

Angelica pubescens var. matsumurae, Aconitum gigas var. hondoense, Hydrangea macrophylla var. macrophylla, Rodgersia podophylla, Filipendula kamtschatica, Aralia cordata, Trautvetteria

Plectranthus japonica, Petasitesjaponicus var. giganteus, trichocarpus, Athyrium pycnosorum, etc.

Dominant species of the community on Hokkaido (Tatewaki 1942) are Angelica ursina,

Artemisia montana, Cirsium kamtschaticum, Cacalia hastata var. orientalis, Senecio cannabifolius, Petasites japonicus var. giganteus. Major components of the similar community in the and of subalpine alpine regions centralHonshu (Suzuki and Nakano, 1965) are Aconitum senanense, Cirsium babanum var. otayae, Rumex arifolius, Polystichum microchlamys, Veratrum stamineum, Saussurea nikkoensis var. sessiliflora, Artemisia monophylla, Angelica edulis, Galium kamtschaticum Carex multifolia, var. acutifolium, etc.

Based on these examples and other observations, it is summerized that the characteristic

of the components tall herb community are species belonging to Angelica, Saussurea, and Cirsium, Aconitum (Ohba, in Miyawaki, 1967; Miyawaki et al., 1968). Phot. 8. Ecolog- ical characteristics of altherbosa are analysed tracing precisely their developmental process, dry matter production, and absorption and accumulation of nutrient elements (Midorikawa, 1959).

4. Alpine region. Phot. 5. Thickets ofPinus Phot. 4.1. pumila. 3. Pinus is pumila a prominent shrub in the alpine region of Japan.It is distributed from the alpine region of central Honshuat the south, to Hokkaido, and further northto theKuriles, and Siberia. thickets elevations Sakhalin, Kamchatka, P. pumila in Japan occur in high exceeding the timber line and consist of Pinus pumila, Vaccinium vitis-idaea, V. axillare,

Cornus Rhododendron Empetrum nigrum var. japonicum, Gaultheria miqueliana, canadensis, Trientalis aureum, Rubus pedatus, Coptis trifolia, europea, Shortia soldanelloides, Sorbus sambucifolia, Tripetaleia bracteata, etc. There are some floristic variations from habitat to habitat and from locality to locality.

The thickets are exclusively restricted to habitats that are sheltered by accumulated

in and that snow winter are exposed early to sunlight in late spring or early summer.

Fig. 16. M. Numata et el.: Natural and semi-natural vegetation in Japan 473

Phytosociologically, four associations have been reported from the alpine region of

Honshu and Hokkaido. They are Vaccinio- (from central Honshu) (Suzuki, 1964; Suzuki and Umezu, 1965), Sorbeto-, Rubeto-, and Ledeto—Pinetum(from Hokkaido) (Kobayashi,

1967).

The main associations of the alpine thickets in Hokkaido are as follows according to

Pinus Tatewaki (1963): pumila-Rhododendron aureum-Empetrum nigrum var. japonica ass.,

P. pumila-Arctous alpina var. japonica-Vaccinium uliginosum ass., P. pumila-Ledum palustre P. P. ass., P. pumila-Sorbus matsumurana ass., pumila-Sasa kurilensis ass., and pumila-non plant cover ass.

4.2. Snow-patch vegetation. Phot. 6.

The snowfall blown by north-westerly winds in winter is accumulated on leeward slopes of ridges and in depressions. Accumulated snow is usually more than 5 m thick. the The snow melts away, or remains unmelted as snow-patches insummer. Slopes below snow-patch are mesic to wet, and those above the patch are dry. The plant community of the upper slopes, which is named Anaphalio-Phyllodocetum aleuticae (Ohba, in Miyawaki, sitchense Shortia soldanel- 1967), is made up of Phyllodoce aleutica, Lycopodium var. nikoense, loides f. alpina, Geum pentapetalum, Arnica unalascensis var. tschonoskyi, Anaphalis alpicola,

Deschampsia flexuosa, etc. (Ishizuka, 1950; Masamune, 1961; Itow et al., 1964; Ohba, in Miyawaki, 1967). Phot. 6. and flat below the On the other hand, mesic gentle slopes places snow-patch support different a community composed of Moliniopsis japonica, Fauria crista-galli, Carex blephari-

Shortia carpa, soldanelloides f. alpina, Tojieldia japonica, Coptis trifolia, Tilingia ajanensis, etc. and Plantago hakusanensis Primula cuneifolia var. hakusanensis (in central Honshu) or P.

is nipponica (in northern Honshu) are characteristic to the community that developed on habitats below the the the further wet patch in alpine region on Japan Sea side (Yamazaki and Nagai, 1961; Ohba, in Miyawaki, 1967).

Flat ridges and gentle slopes in the subalpine and alpine regions of the Japan Sea side are outstanding in lacking forest communities, or in limited development of these, as

before. The stated herbaceous community developed on such areas is also dominated by

Fauria Carex Other Moliniopsis japonica, crista-galli and/or blepharicarpa. components are almost similar to the above-listed species, though with somelocality-to-locality variations

(Miyawaki et al., 1967, 1968; Shimizu, 1967).

4.3. Wind-blown heathlands

wind-blown heathlands found with little accumulation Alpine are on ridges snow whichare subject to severe winds in winter. The compositional characteristics ofheathland communities the of such dwarf scrubs are prevalence as Arctous alpinus var. japonicus,

Arcterica nana, Loiseleuria procumbens, Vaccinium uliginosum, V. vitis-idaea, Empetrum nigrum

and var. asiatica, Diapet lapponica var. obovata, etc. (Ohba, 1964; Suzuki, 1964; Suzuki Herbaceous Umezu, 1965; Ohba, in Miyawaki, 1967). species are Carex stenantha, Gentiana algida, Geum calthaefolium, and Deschampsia flexuosa; common lichens are Cetraria islandica var. orientalis. Thamnolia vermicularis, Cladonia rangiferina, C. alpestris, Cetraria chrysantha, etc. The first three species of the above-listedlichens andLoiseleuriaprocumbens are common to the European alpine flora (Suzuki, 1964). Phytosociologically, two associations of heathland communities have been reported: Arcterio-Loiseleurietum (Ohba, 1964; Suzuki,

1964; Suzuki and Umezu, 1965) and Leontopodio-Arcterietum (Shimizu, 1967). The of the heath Hokkaido follows main associations alpine in are as according to Tatewaki (1963): Empetrum nigrum var. japonicum-Rhododendron aureum ass., Vaccinium VOL. No. 474 BLUMEA XX, 2, 1972

uliginosa-Arctoüs alpinus var. japonicus ass., Diapensia lapponica var. obovata-Arcterica nana- Loiseleuria procumbens and aleutica ass., Phyllodoce caerulea-Phyllodoce ass.

desert 4.4. Alpine vegetation The desert loose substrata alpine vegetation is developed on with, more or less, movable sands and boulders is on slopes, and made up of sparse growthof plants. It is phytosociolo- classified into and gically 18 associations 8 communities corresponding to associations (Ohba, 1969).

IV. EPILOGUE

The of the purpose present paper was to outline the natural and semi-natural vegetation of in relation to environmental conditions Japan (Tables 6—7). Here we would like to refer the trend of to changing man-vegetation relationships and to the ecological and the phytosociological approaches to problems. like other industrial In Japan, in densely populated countries, the original or natural

vegetation is restricted to limited portions of the territory (Miyawaki and Itow, 1966). The broad-leavedforest evergreen region whichoccupies the lower region of the southern half of the of Japanese Archipelago is course most disturbed of all and at present only fragments of this climax forest This are found. destruction still goes on. As stated before,

secondary forests and grasslands have long been maintainedby human interference such

as there in logging, mowing, burning, or grazing. Apparently, was the past an equilibrium between and In man vegetation. recent years, however, the population increment follow- the industrialization ing is outstandingly rapid in such big cities as Tokyo, Chiba, Yokohama, and and the Nagoya, Osaka, Fukuoka, consequence results, in part, in the

thoughtless development ofresidential areas and traffic facilities in and around the cities. The balanced man-vegetation relationship is now upset by intrusion of big-machine- the technology and pesticides in suburban areas of secondary forests, grasslands, and farmlands; the environment of human life is deteriorated and uniform becoming in a

depauperate state.

In forest natural the summergreen broad-leaved region, forests have been logged down

over surfaces. large The logged forest lands are mostly reforested uniformly by coniferous

trees such as Cryptomeria japonica, Chamaecyparis obtusa, or Larix leptolepis (Miyawaki and Itow, 1966).

The deforestation is also the policy encroaching on parts of primeval subalpine conifer- and ous forest region; the deformation destruction of natural habitat by tourism develop-

ment invade further the alpine region, where the natural communitiesof Pinus pumila, dwarf and herbs shrubs, are developed under severe conditions. The restoration of these disturbed 'fragile' communities, once or destroyed, by their own revegetation potential

can hardly be expected.

Under such of circumstances, some Japanese ecologists and phytosociologists have shown the scientific basis for the wise land and for the use revegetation planning of dete-

riorated or denuded areas in terms of their disciplines. Examples are those in the plannings of new town development and plant cover restoration made in the following areas in the of Tsukuba vicinity Tokyo: in (Yokoyama et al., 1967), Kohoku New Town area (Miya- waki et al., 1968), Minami-Tama New Town area (Miyawaki, 1969), Fujisawa (Miya- waki and and Chiba Fujiwara, 1968a), New Town area (Numata, 1969b). A revege- also offered toward tation program is the development plan of tourist facilities and roads the north on south and slopes ofMt. Fuji (Miyawaki et al., 1967,1969). In the Oze region, M. Numata et al: Natural and semi-natural vegetation in Japan 475

the and rehabilitation studies are making steady progress regarding ecological judgement of the degraded moorland communities that result from trampling by hikers (Miya-

waki and Fujiwara, 1968b). and Fundamental researches for the management and establishment of national parks

have a and national parks and nature reserves been made in number of existing proposed reserves. In some of those researches, a vegetation map is compiled, for example, hi the

from north to central Hokkaido et 1955), following areas (arranged south): (Tatewaki al.,

the Mts. Tashiro and Aizu-koma region (Miyawaki et al., 1967), Okutadami region

(Miyawaki et al., 1968), Mt. Tanzawa (Miyawaki et al., 1964), Mt. Fuji (Miyawaki et al.,

1967), Mt. Hakusan (Masamune, 1961), Ise-Shima region (Miyawaki and Fujiwara, 1969), its Minoo (Miyawaki, 1966c), Mt. Dogo (Horikawa and Sasaki, 1959b), Sandankyo and Is. vicinity (Horikawa and Sasaki, 1959a), Okunoshima Is. (Horikawa et al., 1962), Danjo

(Toyama et al., 1968), etc. (cf. Miyawaki, 1966a, b). made The mapping ofactual vegetation in Japan in 1/200,000 is now being on the basis of prefectural unit under the sponsorship of the Ministry ofEducation. The comparison

the of natural shows the of map existing with that ofpotential vegetation not only vege- also offer the basis for the land the tation-habitat relationships, but objective use, preser- of deteriorated habitats vation of valuable endangered natural areas, and the restoration such and vegetation (Miyawaki, I968d, e). A most important premiss in vegetation studies

its relation to is accurate observation, a wide view of Japanese vegetation as a whole and and integration with that of other parts of the world (Miyawaki, 1967).

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in the Shiretoko Phot. 1. Climax forest ofAbies sachalinensis with a goodregenerative structure Peninsula, northeastern Hokkaido. (Photo M. Numata)

Phot. 2. Sasamorpha purpurascens, a representative of the undergrowth species in the beech forest on the Pacific side of Honshu. Hakonc, central Honshu. (Photo A. Miyawaki) VOL. No. 484 BLUMEA XX, 2, 1972

A is the ermatii Phot. 3. subalpine to alpine transition landscape. Pinus pumila (left) on ridge; Betula on the slope ( Alnus maximowiczii in the bottom of ravine, invisible). Elevation about 2500 m, Mt. Norikura, central Honshu. (Photo A. Miyawaki)

A view forest of Picea obtusa Phot. 4. bird’s-eye of mixed pioneer polita, Tsuga sieboldii, Chamaecyparis lava bed that in north of etc. growing on the flowed 865. Elevation about 900 m, Aokigahara at the foot

Mt. Fuji, central Honshu. (Photo S. Itow) M. Numata et at.: Natural and semi-natural vegetation in Japan 485

Larix the line Mt. alt.Phot. 5. Shrubby growth(Photoof leptolepis, Tsuga diversifolia,M.etc. atNumata)tree on Fuji, 2500 m

Phot. 6. An alpine community of Phyllodoce aleutica and Anemone narcissiflora. Elevation about 2800 m, Mt. Tateyama, central Honshu. (Photo S. Itow) BLUMEA VOL. No. 486 XX, 2, 1972

and maximowiczii Phot. 7. A subalpine landscape. Betula ermatii grows on upper slope Alnus on the middle slope (foreground) separated by grassland. Kurobe, central Honshu. (Photo S. Itow)

Phot. 8. Tall herb community of Angelica pubescens var. matsumurae and Aconitum senanense. Elevation

A. about 2500 m, Mt. Norikura, central Honshu. (Photo Miyawaki) M. Numata el al.: Natural and semi-natural vegetation in Japan 487

Phot. 9. Interior of the Castanopsis sieboldii forest in coastal area. The floor communityis composed of

shrubs such Aucuba Fatsia etc. central evergreen as japonica, japonica, Ligustrum japonicum, Kamakura, Honshu. (Photo A. Miyawaki)

Phot. 10. A climax forest of Cinnamomum camphora at Fukuoka, Kyushu. (Photo M. Numata) BLUMEA VOL. No. 488 XX, 2, 1972

Phot. A Pinus 11. stand of densiflora, a representative of secondary forests. Tsukuba, central Honshu

(Photo S. Itow)

Phot. 12. Coastal vegetation composed of Quercus phillyraeoides, Pittosporum tobira, Pinus thunbergii, etc. (Pittosporeto-Quercetum phillyraeoidetis). Kii Peninsula, central Honshu. (Photo A. Miyawaki) and in M. Numata et al.\ Natural semi-natural vegetation Japan 489

The Island. Lumnitzera

and Iriomote

in mucronata, River Rhizophora

Nakamagwa del, can the of Kandelia estuary Miyawaki) the conjugate, A. at landscapeBruguiera (Photo of Ryukyu. mangrove consists

A forest Iriomote, 13. Phot. mangrove racemosa. BLUMEA VOL. No. 490 XX, 2, 1072

Phot. 14. A stand of Livistona chinensis var. subglobosa which is distributed from southernmost Kyushu and Shikoku southward. Aoshima Island, Kyushu. (Photo A. Miyawaki)

Natural Livistona Ardisia Phot. 15. forest composed of boninensis, Schima mertensii, sieboldii, Hibiscus glabra, Raphiolepis integerrima, etc. Mt. Yoake-yama, Chichi-jima, Bonin Is. (Photo A. Miyawaki) semi-natural in M. Numata et at: Natural and vegetation Japan 491

Iris Phot. 16. A moist grassland on peat covered by Spodiopogon sibiricus, setosa, Filipendula kamtschatica,

The is M. Numata) etc. et Kerochi, Hokkaido. upper part a sown pasture. (Photo

Lake beach the Sea of Okhotsk near the Phot. 17. A sandy grassland dominated by Elymus mollis, facing

Notoro, Hokkaido. (Photo M. Numata) BLUMEA VOL. No. 492 XX, 2, 1972

Phot. 18. A natural bamboo brake of Phyllostachys bambusoides on a steep slope near Kyoto. (Photo M. Numata)

Phot. General view ofa meadow dominated Miscanthus sinensis distributed 19. high-grass type by widely in Japan, at Kawatabi, northern Honshu. (Photo M. Numata) in 493 M. Numata et al.: Natural and semi-natural vegetation Japan

dominated distributed in Phot. 20. General view of a short-grass type pasture by Zoysia japonicamostly N. M. the cool-temperate region. Hakkoda, Honshu. (Photo Numata)

Mt. Phot. A dominated Pleioblastus distichus var. nezasa on Kuju, 21. dwarf-bamboo type pasture by Kyushu. (Photo M. Numata) 13LUMEA VOL. 494 XX, No. 2, 1972

Phot. Moor in the 22. community complex Ozegahara Basin. Major species are Moliniopsisjaponica,

Vacciniun A. Drosera rotundifolia, oxycoccus, Andromeda polifolia, Sphagnum papillosum. (Photo Miyawaki)

Phot. A Ischaetnum 23. coastal sand-dune covered by anthephroides, Carex kobomugi, Calystegia soldanella, etc. at Kashimanada, central Japan. (Photo M. Numata) M. Numata et at.: Natural and semi-natural vegetation in Japan 495

Numata)

M. (Photo Hokkaido.

eastern Atsukeshi,

lagoon,

a of shore

the along herbacea Salicornia

of vegetation halophytic

A

24.

Phot. 496 BLUMEA VOL. XX, No. 2, 1972

Phot. A solfatara of Miscanthus 25. vegetation Pinus pumila, Ledum palustre var. yesoense, sinensis, etc. at Iozan, Hokkaido. (Photo M. Numata)

Phot. 26. Sparse vegetation ofPinus thunbergii, Pittosporum tobira, etc. on the lava ofSakurajima, southern Kyushu. (Photo M. Numata)