Japanese-Korean Forest Habitats
High altitude forests, such as those in the so-called Japanese Alps, are largely dominated by endemic conifers such as Abies veitchii, A. mariesii (Pinaceae) and Tsuga diversifolia (Cupressaceae), while other endemics conifers like Chamaecyparis obtuse (family), Larex leptolepis (family), Picea koyamai, P. maximowiczii (Pinaceae) and Thuja standishii (Cupressaceae) are locally common.
In the cool-temperate zone, endemic beech trees, Fagus crenata and F. japonica (Fagaceae), and Quercus crispula dominate the so-called oriental deciduous forests. Fagus japonica is mainly confined to the lower (or warmer) parts of the zone, so the two beeches rarely intermix. The composition of these woodlands also varies on an east - west gradient with those on the Pacific side differing from those on the Sea of Japan side. The endemic Abies homolepis (Pinaceae), for example, only occurs on the Pacific side. Other deciduous trees include endemics such as Acer japonicum (Aceraceae) and Tilia japonicum (Tiliaceae), while endemic shrubs include Camellia rusticana (Theaceae) and Magnolia salicifolia (Magnoliaceae). The undergrowth is largely composed of dwarf-bamboo including the endemic Sasa nipponica (Poaceae). Other rare and endemic ground layer species include Cerastium arvense var. ovata (Caryophyllaceae), Mitchella undulata (Rubiaceae), Hylotelephium tsugaruense (Crassulaceae), Poa ogamontana (Poaceae), Pyrola japonica (Pyrolaceae), Ranzania japonica (Berberidaceae), Shortia uniflora (Diapensiaceae), Silene aomoriensis (Caryophyllaceae) and many orchids such as Calanthe discolor, C. nipponica, Cypripedium yatabeanum, Gymnadenia fujisanensis and Tipularia japonica. Among the fern flora is the endemic Blechnum niponicum (Blechnaceae). An important sanctuary for these species is the Shirakami-sanchi Natural World Heritage site in northern Honshu. Where the soils are very moist or wet, as in certain valley woodland, the species composition may include endemic trees such as Aesculus turbinata (Hippocasttanaceae), Cercidiphyllum japonicum (Cercidiphyllaceae) and Euptelea polyandra (Trochodendraceae). However, in addition to broad-leaved trees, the cool temperate zone includes stands dominated by various endemic conifers such as Chamaecyparis obtuse, Cryptomeria japonica and Thujopsis dolabrata (Cupressaceae).
Broad-leaved evergreen trees with Quercus and Castanopsis
forests of the warm-temperate zone. The principal oak species are Quercus acuta, Q. gilva, Q. glauca, Q. myrsinaefolia, Q. salicina and Q. sessilifolia. These forests are highly stratified with at least two or three tree layers and a shrub layer. Endemic trees include Castanopsis cuspidata (Fagaceae)and Cinnamomum japonicum (Lauraceae)while amongst the endemic shrubs are Camellia japonica (Theaceae)and more locally Abeliophyllum distichum (Oleaceae). In some localities evergreen stands of coniferous trees occur including endemics such as Pinus thunbergii (Pinaceae) and Tsuga siebaldii (Pinaceae). Throughout the temperate zone there are also swamp forests with endemic trees like Alnus japonica (Betulaceae), Fraxinus mandshurica var. japonica (Oleaceae)and Salix sacchalinensis (Salicaceae). The shrub layer of these forests includes Rhamnus crenata and R. japonica while herb layer species are Carex angustinowiczii, C. rhynchophysa, Lysichiton camtschatense, and the ferns Osmunda cinnamomea var. fokiensis and Lastrea thelypteris.
Japanese-Korean Grasslands
Natural climax grasslands only occur in the alpine zones. All of the rest have been established under the influence of human activities including grazing domestic stock, mowing, firing and trampling. They can be broadly classified into: Miscanthus grassland, Pleioblastus grassland, Sasa grassland and Zoysia grassland. In Miscanthus grassland, M. sinensis is usually the dominant species and forms the main type of tall-grass meadow in this BioProvince. Endemics associated with these grasslands include Artemisia japonica (Asteraceae), Cirsium japonicum (Asteraceae) and Eupatorium japonicum (Asteraceae). In contrast, Zoysia grassland is typically short turf pasturelands. The endemic Zoysia japonica (Poaceae) is usually the main species, especially in the cool-temperate regions. Associated endemics include Berberis thunbergii (Berberidaceae) Cirsium tanakae (Asteraceae) Geranium thunbergii (Geraniaceae) Rhododendron japonicum (Ericaceae) and Spiraea japonica (Rosaceae). Pleiobastus grassland is the main dwarf-bamboo pasture or meadow of the warm-temperate zone. The dominant species is usually Pleiobastus distichus var. nezasa, while the associated endemics include Polygala japonica (Polygalaceae). In the cool-temperate zone species of Sasa grassland dominate the main dwarf-bamboo communities. The dominant species are usually the endemic Sasa veitchii or S. nipponica (Poaceae).
Japanese-Korean Alpine Zone
Just above the tree line there are often extensive thickets of coniferous shrubs. These typically include Pinus pumila (Japaneae stone pine) but in the mountains adjacent to the Sea of Japan others include Juniperus communis var. nipponica and Taxus cuspidata var. nana. Broad-leaved deciduous shrubs such as Acer tschonoskii, Alnus maximowiczii and Sorbus matsumurana may also form important stands in this zone, while the ground layer can be important for lichens such as Cetraria crispa var. japonica. Alpine grasslands usually include various species of Calamagrostis including C. longiseta, C. fauriei, C. langsdorffii and C. matsumurana, but one of the most conspicuous plants of this zone is Phyllodoce aleutica. Other flowering plants include Arnia unalaschensis, Cardamine nipponica and the endemic Parnassia alpicola (Parnassiaceae) and Boykinia lycoctonofolia (Saxifragaceae). The clubmoss, Lycopodium sitchense var. nikoense, is also likely to be featured. In the more permanantly wet areas Moliniopsis japonica becomes the chief grass species together with Carex blepharicarpa and Fauris crista-galli. These areas are also characterised by the presence of various tiny endemic primroses such as Primula nipponica and P. cuneifolia (Primulaceae). Other alpine grassland endemics include Minuartia hondoensis, (Caryophyllaceae), various Oxytropis species such as O. japonica (Fabaceae), Plantago hakusanensis (Plantaginaceae), Ranunculus acris var. nipponicus (Ranunculaceae, Trautvetteria japonica (Ranunculaceae), Trollius reiderianus var. japonicus (Ranunculaceae). Finally flourishing in less exposed areas are various alpine heaths. In the mountains of Japan, Empetrum nigrum var. japonicum is one of the most conspicuous elements of these heaths, while other dwarf shrubs are Arcteria nana, Loiseleuria procumbens and Diapensia lapponica var. obovata.
Japanese-Korean Saltmarsh
Saltmarshes are largely composed of halophyte taxa that are distributed throughout the northern hemisphere. More restricted and endemic taxa found here include grasses such as Puccinellia kurilensis, P. nipponica and Zoysia sinica var. nipponica (Poaceae) and the succulent Suaeda japonica (Chenopodiaceae). On the lower shore, seagrass beds include the endemic Zostera caulescens (Zosteraceae).
Japanese-Korean Maritime Cliffs
Sea cliffs in northern zones, such as on the Skimokita Peninsula in Honshu, have a rock crevice community dominated by Chrysanthemum yezoense, Cyrtomium falcatum, Lillium macuratum var. dauricum, Lysimachia mauritiana and Sedum verticillatum, while further inland in the less maritime zones, endemic thickets of Juniperus chinensis var. sargentia with Empetrum nigrum var. japonicum may be present. Other more local endemics of this habitat include Chrysanthemum nipponicum (Asteraceae) and Puccinellia nipponica (Poaceae). In the more southern warm-temperate zones, maritime rocks and scree usually include Carex oahuensis, Miscanthus condensatus and Peucedanum japonicum, while the maritime scrub here includes Elaegnus macrophylla, Euonymus japonica var. radicifer, Eury emarginata, Rhaphiolepis umbellate and Pittospermum tobira. Amongst the shrubs can be found various maritime forbs such as Farfugium japonicum and Boehmeria biloba.
Japanese-Korean Coastal Sand Dunes (General)
The principal mobile dune species are Carex kobomugi, Elymus mollis and Wedelia prostrata. Mobile dune endemics possibly include Linaria japonica (Scrophulariaceae). In the semi-stable zones, Hibiscus hamabo, Rosa wichuraiana and Vitex rotundifolia often form low thickets. Mosses largely dominate the so-called grey dunes but lichens are largely absent. Dune woodland is well represented. The succession of ligneous vegetation on fixed dunes is thought to start with the development of Juniper conferta stands. This gives way the Pinus densiflora dominated vegetation but is eventually succeeded by the endemic Tilia japonica (Tiliaceae) and various oaks. Smaller endemic dune trees occurring in southern areas include Eurya japonica (Theaceae) and Fatsia japonica (Araliaceae). Unfortunately most of the natural dune slacks in this BioProvince have been used for the cultivation of rice.
Hokkaido Coastal Sand Dunes
As with dunes throughout Japan, the mobile areas are dominate by Carex kobomugi, Elymus mollis, Glechnia littoralis, Ischaemum anthephoroides, Ixeris repens, Lathyrus maritimus and Zoysia macrostachys, but in addition, the northern mobile dunes of Hokkaido are characterized by Carex macrocephala and the near endemic Linaria japanica (Scrophulariaceae). Fixed dunes typically include scrub often dominated by Rosa rugosa, while interspersed among the scrub and occasional grassland are various flowering plants such as Lilium maculatum var. dauricum, Thermopsis lupinoides and the endemic Hemerocallis yezoensis (Hemerocallidaceae). Quercus dentata is usually the main tree species found on fixed dunes but may be associated with a sparse covering of other trees including the endemic Salix bakko (Salicaceae).
Kyushu Coastal Sand Dunes
On the island of Kyushu the following dune zones have been described.
Strandline
These high tide mark ecosystems are well developed supporting both annual and perennial species. The main annual species is Salsola komarovii but Atriplex gmelinii is locally common. Perennials typically include Calystegia soldanella, Glehnia littoralis (Apiaceae), Lathyrus japonicus, Linaria japonica and Xeris repens (Asteraceae).
Embryonic dunes
Early dune formation is often evident in the more stable areas where they typically form a low foredune ridge up to 2 m high. The main sand binding species here seems to be almost exclusively Carex kobomugi.
Foredunes
The main foredune ridge is relatively low with Carex kobomugi again being the main sand binder. However, this species is only capable of fixing a small proportion of wind blow sand and so the resulting fordunes are relatively small. Associated species are the same as those found in the strandline.
Sheltered slopes of foredunes and the zones immediately behind
Here a distinct community can often be found mainly consisting of Carex breviculmis, Ischaemum anthephoroides (a large perennial grass) and Wedelia prostrata.
Fixed dune grassland
These do not appear to develop in this part of the world which may relate to the fact that much of the dune sand is of volcanic origin.
Blowouts and other areas of bare sand
These are not common but where they do exist the characteristic species usually include the sedges Bulbostylis barbata and Fimbristylis sericea, and the mosses Polytrichum spp and Rhacomitrium canescens.
Dune Scrub
In sheltered hind dune areas dense shrubs may occur. The main species are Elaeagnus umbellata and Rosa wichuraiana. Other species typically include the small herb Viola mandshurica.
Japanese Coastal Vegetated Shingle
Vegetated Coastal Shingle Structures in Central and Southern Japan
In central Japan shingle structures can be divided into beaches developed at the base of sea cliffs derived from cliff erosion and beaches deposited by sea currents. The vegetation was divided into the following six communities:
Calystegia soldanella-Lathyrus japonicus Shingle Vegetation
Associated species include Angelica japonica, Arabis stellari var. japonica, Canavalia lineate, Cirsium maritimum, Lysimachia mauritiana, Messerschmidia sibirica, Polygonum senticosum, Rosa wichuraiana and Rumex japonicus.
Polygonum senticosum Shingle Vegetation
Associated species include Ampelopsis brevipedunculata, Atriplex subcordata, Bidens pilosa, Calystegia soldanella, Cnidium japonicum, Commelina communis, Digitaria timorensis, Lathyrus japonicus, Lysimachia mauritiana, Paederia scandens var. mairei and Torillis japonica.
Messerschmidia sibirica Shingle Vegetation
Associated species include Artemisia princes, Atriplex gmelinii, Atriplex subcordata, Calystegia soldanella, Euphorbia jolkinii, Lathyrus japonicus, Polygonum senticosum, Raphanus sativus var. hortensis, Rosa wichuraiana, Rumex japonicus, Setaria viridis var. pachystachys and Tetragonia tetragonoides.
Euphorbia jolkinii Shingle Vegetation
Assciated species include Agropyron kamoji, Arundo donax, Asparagus lucidus, Bromus japonicus, Calystegia soldanella, Lathyrus japonicus, Lonicera japonica, Miscanthus sinensis, Peucedanum japonicum, Raphanus sativus var. hortensis, Rosa wichuraiana, Rumex japonicus, Torillis japonica, Vicia sepium and Vitex rotundifolia.
Chrysanthemum japonense Shingle Vegetation
Associated species include Calystegia soldanella, Canavalia lineate, Carex fibrillose, Crepidiastrum lanceolatum, Dianthus japonicus, Digitaria violascens, Elaeagnus pungens, Farfugium japonicum, Liriope minor, Lonicera japonica, Lysimachia mauritiana, Miscanthus sinensis, Oxalis corniculata, Paederia scandens var. mairei, Peucedanum japonicum, Rhaphiolepis umbellate, Rosa wichuraiana, Scilla sinensis, Scutellaria indica var. parvifolia, Setaria viridis var. pachystachys and Viola grypoceras.
Rosa wichuraiana Shingle Vegetation
Associated species include Ampelopsis brevipedunculata, Angelica japonica, Asparagus lucidus, Calystegia soldanella, Carex pumila, Cocculus orbiculatus, Dianthus japonicus, Indigofera pseudotinctoria, Lathyrus japonicus, Liriope spicata, Lonicera japonica, Lotus corniculatus var. japonicus, Miscanthus sinensis, Oxalis corniculata, Paederia scandens var. mairei, Peucedanum japonicum and Torillis japonica.
Vegetated Coastal Shingle Structures in South-Western Japan
In south-western Japan vegetated shingle spits are chiefly found in north-western Kyushu. Some of the best preserved examples include Magaribana Spit (near Oshima-mura), Magarizaki Spit, Hinoshima (Island) and Tanoura Spit (Hisakajima, Island).
Calystegia soldanella-Salsola komarovii Shingle Vegetation
This vegetation is typical of the exposed, outer sides of spits with good examples occurring, for example, on the Magarizaki spit. Other species included Canavalia lineata, Commelina benghalensis, Eclipta prostrate, Lysimachia mauritiana, Polygonum senticosum, Portulaca oleracea, Rumex japonicus, Setaria viridis var. pachystachys and Tetragonia tetragonoides.
Messerschmidia sibirica Shingle Vegetation
This vegetation is also typical of the exposed, outer sides of spits particularly along tidal drift lines rich in organic matter. Other species included Calystegia soldanella, Lathyrus japonicus, Setaria viridis var. pachystachys and Tetragonia tetragonoides.
Rosa wichuraiana-Vitex rotundifolia Shingle Vegetation
This dwarf shrub community was found on all shingle spits. Other species included Acalypha australis, Achyranthes fauriei, Agropyron tsukushiense, Allium grayi, Ampelopsis brevipedunculata, Bidens biternata, Bromus japanicus, Calystegia soldanella, Chrysanthemum indicum, Clematis terniflora, Cocculus orbiculatus, Commelina communis, Corchoropsis tomentosa, Corydalis heterocarpa var. japonica, Crepidiastrum lanceolatum, Cyperus rotundus, Dichondra repens, Eurya emarginata, Festuca myuros, Heteropappus hispidus var. arenarius, Ixeris stolonifera, Lespedeza cuneata, Lonicera japonica, Lysimachia mauritiana, Oxalis corniculata, Peucedanum japonicum, Polygonum chinese, Rhaphanus sativus var. hortensis, Rumex japonicus, Sagina maxima, Scilla sinensis, Sedum oryzifolium, Setaria viridis var. pachystachys, Sonchus oleraceus, Tetragonia tetragonoides and Torilis japonica.
Sedum oryzifolium-Lysimachia mauritiana Shingle Vegetation
This vegetation seems to have mainly been associated with man made-habitats comprising heaps of earth and therefore does not represent a natural shingle association. Other species included Asparagus cochinchinensis, Calystegia soldanella, Canavalia lineate, Commelina communis, Corydalis heterocarpa var. japonica, Crepidiastrum lanceolatum, Dianthus japonicus, Heteropappus hispidus var. arenarius, Lathyrus japonicus, Lysimachia mauritiana, Paederia scandens var. mairei, Peucedanum japonicum, Rosa wichuraiana, Sedum oryzifolium, Setaria viridis var. pachystachys and Tetragonia tetragonoides.
Limonium tetragonum Shingle Vegetation
This vegetation was typical of the sheltered, inner sides of spits and was found, for example, on the Magarizaki Spit. It represents a form of shingle saltmarsh. Other species included Artemisia fukudo, Atriplex gmelinii and Suaeda maritima.
Zoysia sinica var. nipponica Shingle Vegetation
This also represents a form of shingle saltmarsh typical of the sheltered, inner sides of spits and was found, for example, on the Magarizaki and Tanoura spits. Other species included Limonium tetragonum and Suaeda maritima.
Polygonum polyneuron-Atriplex gmelinii Shingle Vegetation
Restricted to the sheltered, inner side of spits the vegetation occurs in both saltmarsh and tidal drift lines. No other associates were recorded.
Euphorbia jolkinii Shingle Vegetation
The tidal drift line vegetation was confined to shelter, inner side of spits and found, for example, on the Magaribana Spit. The only other associate was Messerschmidia sibirica.
Raphanus sativus var. hortensis Shingle Vegetation
This vegetation type was only recorded in the Oshima-mura area and here it was heavily influenced by grazing. Other species included Dicondra repens, Messerschmidia sibirica, Oxalis corniculata, Sagina maxima, Sonchus oleraceus and Torilis japonica.
Vitex rotundifolia-Crinum asiaticum var. japonicum Shingle Vegetation
This vegetation was found on all shingle spits except in Oshima-mura and typical of the high-tide line. Other species included Ampelopsis brevipedunculata, Artemisia princeps, Canavalia lineate, Chrysanthemum indicum, Clematis terniflora, Crepidiastrum lanceolatum, Crinum asiaticum var. japonicum, Lonicera japonica, Miscanthus sinensis, Nethrolepis auriculata, Oxalis corniculata, Rosa wichursaiana, Scilla sinensis together with the endemic Trachelospermum asiaticum (Apocynaceae) and Wisteria floribunda (Fabaceae).
Hibiscus hamabo Shingle Vegetation
Found on the sheltered, inner side of spits this vegetation represents a form of semi-mangrove, and was found, for example, on the Magarizaki and Tanoura spits. Other species included Atriplex gmelinii, Crinum asiaticum var. japonicum, Euonymus japonicus and Rosa wichursaiana.
Myoporum bontioides Shingle Vegetation
Comprising monspecific stands Myoporum bontioides, this vegetation also represented a form of semi-mangrove confined to the sheltered, inner side of spits, and was found, for example, on the Magarizaki spit.
References
Doing, H. 1981. A comparative scheme of dry coastal sand dune habitat, with examples from the eastern United States and some other temperate regions. Veroff. Geobot. Inst. Rubel, 77: 41-72.
Hara, H. 1972. Corresponding taxa in North America, Japan, and the Himalayas. In: Taxonomy, Phytogeography and Evolution. Ed. D. H. Valentine. Academic Press.
Hosokawa, T., Tagawa, H. & Chapman, V. J. 1977. Mangals of Micronesia, Taiwan, Japan, the Philippines and Oceania. In: Ecosystems of the World 1 - Wet Coastal Ecosystems. Ed. V. J. Chapman. Elsevier Scientific Publishing Company.
Kanai, H. 1963. Phytogeographical observations on the Japano-Himalayan Elements. Journal of the Faculty of Science, University of Tokyo, Sect 3. Bot. 8: 305-339.
Kolbek, J., Srutek, M. & Box, E. O. 2003. Forest Vegetation in Northeast Asia. Kluwer Academic Publishers.
Nakanishi, H. 1982. Coastal vegetation of the shingle spits of southwestern Japan. Phytocoenologia, 10: 57-71.
Nakanishi, H. 1984. Phytosociological studies on shingle beach vegetation in central and southern Japan. Hikobia, 9: 137-145.
Numata, M. (ed). 1974. The flora and vegetation of Japan. Elsevier Scientific Publishing Company.
Numata, M., Miyawaki, A. & Itow, D. 1972. Natural and semi-natural vegetation in Japan. Blumea, 20: 435-481.
Ohwi, J. 1965. Flora of Japan. Smithsonian Institution. Washington D.C.
Satoo, T. 1983. Temperate broad-leaved evergreen forests of Japan. In: Ecosystems of the World 10 - Temperate Broad-Leaved Evergreen Forests. Elsevier.
Sun, B. & Stuessy, T. F. 1998. Preliminary observations on the evolution of the endemic angiosperms of Ullung Island, Korea. In: Evolution and Speciation of Island Plants. Eds. T. S. Stuessy and M. Ono. Cambridge University Press.