Formations in the Manchurian BioProvince

Peter Martin Rhind

Manchurian Larch Forest These are either dominated by Larix gmelinii or the endemic Larix olgensis (Pinaceae). The former extends southeastwards from northern Daxingan-ling via Xiaoxingan-ling, Wanda-shan and Zhangguangcai-ling to Laoye-ling. The few other associated trees include Acer mono, Betula platyphylla, Pinus koraiensis and Quercus mongolica. Typical shrub layer species are Ledum palustre, Rhododendron dauricum, Spiraea media and Vaccinium uliginosum. The rich herb layer is often dominated by species of Carex, but may also include Aegopodium alpestre, Dendrantheme zawadskii, Kitagawia terebinthacea, Mitella nuda, Orthilia secunda, Pulsatilla davurica, Pyrola davurica, Sedum aizoon and the two endemic species Geranium maximowiczii (Geraniaceae) and Saussurea amurensis (). Forests dominated by Larix olgensis occur mainly on Changbai-shan, but extend northwards to Zhanguangcai-ling and Laoye-ling. At low altitudes other canopy trees include Ulmus japonica and the endemic or near endemic Fraxinus mandshurica (Oleaceae) and Tilia amurensis (Tiliaceae), while at higher altitudes other trees such as the endemic Betula costata () may be present. The under story comprises a well-developed shrub and herb layers. Typical shrubs include Eleutherococcus senticosus, Deutzia parviflora and the endemic or near endemic Corylus mandshurica (Corylaceae) and Ribes mandshuricum (Grossulariaceae). Vines such as Actinidia kolomikta and Schisandra chinensis may be present, and at ground level Carex species such as C. callitrichus, C. quadriflora and C. ussuriensis are again common. However, on the lower slopes, the herb layer typically becomes dominated by taller species often reaching up to 1 m in height. These include Filipendula angustifolia, Matteuccia struthiopteris, Saussurea grandifolium, Thalictrum tuberiferum and the endemic or near endemic Valeriana amurensis (Valerianceae).

Manchurian Spruce-Fir Forest These are dominated in various amounts Picea jezoensis, P. koriensis and Abies nephrolepis. They are usually associated with wetter area in places such as Changhai- shan, Daxingan-ling and Xiaoxingan-ling. The few other associated trees include the endemic Betula costata (Betulaceae), Phellodendron amurense (Rutaceae) and Tilia amurensis (Tiliaceae) in the better-drained areas at high elevation. The under storey usually consists of a mixed shrub-herb layer. Among the shrubs are various endemic species such as Acer tegmentosum (Aceraceae), Berberis amurensis (), Deutzia glabrata (Saxifragaceae), Euonymus pauciflora (Celastraceae), Lonicera maximowiczii (Caprifoliaceae), Ribes maximowiczianum (Glossulariaceae) and the vine amurensis (). Common herbs are small, shade tolerant species and include many taiga elements like Goodyera repens, Linnaea borealis, Maiathemum bifolium and Trientalis europaea. Among the endemic herbs are Brachybotrys paridiformis (Boraginaceae) and Leptorumohra amurensis (Dyropteridaceae).

Manchurian Pine Forest Forest dominated by pine widely distributed. The most common species is Pinus koraiensis, while less common taxa include Pinus desiflora var. ussuriensis (endemic to a small area around Lake Xingkai), Pinus sylvestris var. sylvestriformis (endemic to a small area on the northern slopes of Changbai-shan), and the near endemic Pinus sylvestris var. mongolica, which is confined to , Mongolia and eastern Siberia. The other associated trees are somewhat similar from forest to forest, but Pinus koriaensis is often associated with Abies nephrolepis, Picea jezoensis and P. koraiensis, while under storey Copyright © 2010 Peter Martin Rhind trees include Populus tremula var. davidiana and the endemic or near endemic Maackia amurensis (Fabaceae). Shrub layer development varies but may include several endemic or near endemic taxa such as Rhamnus diamantiaca (Rhamnaceae), Sambucus recemosa subsp. mandshurica (Caprifoliaceae) and Syringa reticulata var. mandschurica (Oleaceae). Typical herb layer species are Artemisia santolinifolia, Sedum selskianum and Bupleurum longiradiatum, but in some of the dryer sites such as in stands of Pinus syvestris var. mongolica on sand, the herb layer maybe dominated by common steppe species like Agriophyllum squarrosum, Astragalus adsurgens, Filifolium sibiricum, Lilium pensylvanicum, Stipa baicalensis and Thymus dahuricus.

Manchurian Juniper Woods Woods dominated by Juniper rigida occur in southern Changguangeai-ling on dry, nutrient- poor south-facing slopes and ridges. Canopy cover is usually sparse and trees rarely reach more than about 4 m in height although occasionally reaching 10 m. Juniper is usually the single dominant but a few broad-leaved trees such as Acer truncatum, Betula davurica, Prunus sibiricum, Quercus liaotungensis and Q. magnolica may be present. The shrub layer is spare rarely exceeding 40 cm in height and is typically dominated by the shade tolerant endemic Spiraea pubescens (Rosaceae), but also includes Indigofera kirilowii, Lespedeza davurica and Rhododendron davurica. Among herb layer are Asparagus oligoclonos, Patrinia rupestris, Scilla scilloides and Scorzonera austriaca, but their cover is often less than 50%.

Manchurian Oak Forest This BioProvince includes a rich diversity of oak forest types each dominated by a different species of oak. Quercus mongolica forest is widespread with trees reaching 18 m in the more temperate areas. The shrub layer varies from place to place but includes several endemic species such as Rhododendron schlippenbachii (Ericaceae) and Spiraea pubescens (Rosaceae). The herb layer can be up to 80 cm tall and may include Artemisia keiskeana, Dendranthema naktongense, Silene jenisseenis and the endemic or near endemic Diarrhena manshurica (Poaceae). Vines such as the endemic or near endemic Clematis manschurica (Ranunculaceae) may also be present. Quercus dentata forest extends from southeastern south to . Pure stands are confined to steep south facing slopes and rarely reach more than about 4 m tall. The sparse shrub layer typically includes Indigofera kirilowii, Lespedeza davurica, Prunus humilis and Securinege suffruticosa. Likewise the herb layer is pretty sparse and usually dominated by Carex callitrichos, but may also include the endemic Polygonatum stenophyllum (Liliaceae) and Rubia cordifolia (Rubiaceae). Forests dominated by the endemic Quercus liaotungensis (Fagaceae) are confined to central and western Liaoning. They have a well- developed shrub layer that varies from place to place but may include the endemic Exochordia serratifolia (Rosaceae). The herb layer is usually dominated by Carex calitrichos, but also includes Asparagus oligoclones, , Melampyrum roseum and Saposhnikovia divaricata. Forests dominated by Quercus aliena occupy small areas mainly on the Liaodong Peninsula. Associated trees include the endemic or near endemic Juglans mandshurica (Juglandaceae). Alangium platanifolium, Corylus heterphylla, Euonymus alata and Symplocos paniculata are among the dominants of the species-rich shrub layer. Carex again dominates the herb layer mainly C. ciliato-marginata and C. siderosticta, but these are usually joined by other species such as the endemic Geranium koreanum (Geraniaceae). Also found on the Liaodong Peninsula as well as on adjacent islands is forest dominated by Quercus acutissima. The shrub layer of these forests includes Rhus javanica, Weigela praecox and Zanthoxylum schinifolium. Important herbs include Carex calitrichos and C. humida but these may be joined by Cleistogenes polyphylla, Leontopodium leontopodioides and Potentilla discolor. Finally small stands of

Copyright © 2010 Peter Martin Rhind forest dominated by Quercus variabilis can also be found on the Liaodong Peninsula, but confined to southern areas. The shrub layer tends to be dominated by Indigofera kirilowii and the herb layer is again dominated by Carex callitrichos. But many other herbaceous species may be encountered including Codonopsis lanceolata, Cynanchum atratum, Duchesnea indica and Syneilesis aconitifolia.

Manchurian Forest Birch forest in this BioProvince maybe dominated by one of four species - Betula davurica, B. ermanii, B. platyphylla or the endemic B. costata (Betulaceae). Betula platyphylla forest is the most widely spread forest type in northeast covering about 20% of the total forested area, despite being a secondary forest type. Betula davurica forest is far less common being confined to areas south of Daxingan-ling and Xiaoxingan-ling. It occurs in small patches on south-facing slopes between about 300 -700 m. Betula costata forest is another secondary forest type occurring mainly at high elevations and pure stands are rarely encountered. forests are also confined to high elevations mostly near the tree line in the eastern parts of the BioProvince. Other associated trees vary from forest to forest but may include several endemic or near endemic species such as Fraxinus mandshurica (Oleaceae), Juglans mandshurica (Juglandaceae), Phellodendron amurense (Rutaceae) and Tilia mandshurica (Tiliaceae). The shrub layer also varies but endemic species such as Acer pseudosieboldianum (Aceraceae) and Ribes komarovii (Grossulariaceae) may be present. Endemic species possibly encountered in the herb layer include Jeffersonia dubia (Berberidaceae), Ligusticum tenuissimum (Apiaceae) Trollius macropetalus (Ranunculaceae) and Veratrum dahuricum (Liliaceae).

Manchurian Aspen-Poplar Forest These forests can be be broady divided into three mainly secondary forest types - Populus koreana forest, P. maximowiczii forest, P. suaveolens forest and P. tremula var. davidiana forest. The most widespread of these is P. tremula forest which occurs mainly on leeward slopes between 800 – 1000 m. Populus koreana and P. maximowiczii forests are far less common being mainly confined to eastern areas, while P. suaveolens forest occurs primarily on Daxingan-ling. Their canopies vary in height from about 23 m in P. tremula forest to about 30 m in P. maximowiczii forest. Associated tree vary from forest to forest but include many endemic or near endemic species such as Fraxinus mandshurica (Oleaceae), Juglans mandshurica (Juglandaceae) and Tilia amurensis (Tilaceae). The shrub and herb layers also vary but several endemic species may be encountered such as the shrubs Atraphaxis manshurica (Polygonaceae) and Lonicera maximowiczii (Caprifoliaceae), and the herbs Fritillaria maximowiczii (Liliaceae) and Plectranthus excisus (Lamiaceae).

Manchurian Elm Forest One or two of three elm species dominate these forests - Ulmus japonica, U. macrocarpa and U. pumila - depending on location. Natural forests of U. macrocarpa and U. pumila are confined to the temperate steppe region of Northeast China and in places both species occur together. Canopy height varies from 7-10 m. Typical shrub layers include Caragana microphylla, Sambucus williamsii and Spiraea fritschiana, while the varied herb layer often comprises the endemic or near endemic Erysimum amurense (Brassicaceae). Ulmus japonica forest is usually confined to alluvial soils and rivers banks in eastern areas where it often forms monodominate canopies. Typical shrub layers include Salix gracilistyla and Sorbaria sorbifolia, while the well-developed herb layer comprises species such as Aconitum kusnezoffii, Angelia dahurica, Equisetum sylvaticum, Galium maximowiczii and Urtica angustifolium.

Copyright © 2010 Peter Martin Rhind Manchurian Lime Forest Forest dominated by the endemic or near endemic Tilia amurensis (Tiliaceae) occur widely in eastern areas as well as in western Liaoning Province. The canopy can be very dense with a cover of up to 90% formed by a diverse mixture of broad-leaved trees. Other characteristic species include Acer mono, Aralia elata, Carpinus cordata, Fraxinus rhynchophylla and several endemic taxa. The shrub layer is often high, up to 2 m and very rich in species including Myripnois dioica, Ostryopsis davidiana, Philadelphus pekinensis and a great variety of endemic taxa. Vines are also prolific with species like the endemic or near endemic Aristolochia manchuriensis (Aristolochiaceae) and Vitis amurensis (Vitaceae). Endemic or near endemic taxa are also well represented in the rich herb layer with species such as Asarum heterotropoides var. mandshuricum (Aristolochiaceae) and several spring-ephemerals like Adonis amurensis (Ranunculaceae) and Gagea hiensis (Liliaceae).

Manchurian Ash Forest These forests are either dominated by Frazinus mandshurica or F. rhynchophylla. The former occurs in eastern areas mainly along rivers or on gentle slopes, while the latter is confined to slopes in Liaoning Province. They share a similar assemblage of associated trees including several endemic or near endemic species like Maackia amurensis (Fabaceae) and Syringa reticulata var. mandschurica (Oleaceae). The shrub and herb layers also have several species in common including the endemic shrub Rhamnus diamantiaca (Rhamnaceae) and the endemic herb Plectranthus excisa (Lamiaceae).

Manchurian Alder Forest These forests, dominated by Alnus hirsuta, are mainly confined to mountainous areas in the east and north of the BioProvince. The canopy can reach 10-12 m and may include Malus buccata, Prunus padus and Pyrus ussuriensis. Shrub layer species can reach 2 m in height and typically include Ribes procumbens, Rosa davurica and Rubus matsumuranus. The herb layer is usually well developed with species such as Comarum palustre, Geranium eriostemon, Maiathemum dahuricum and Pedicularis resupinata.

Manchurian Chosenia arbutifolia Forest These forests usually form narrow strips along rivers and streams on well-drained soils in northern and eastern areas. They are strongly dominated by Chosenia arbutifolia, but may also include Populus maximowiczii and the endemic or near endemic Fraxinus mandshurica (Oleaceae). The shrub layer usually includes Cornus alba, Salix udense, Sambucus buergeriana and the endemic or near endemic Syringa reticulata var. mandschurica (Oleaceae). With herb layer cover of up to 70 %, it includes a large variety of species such as Anemone dichotoma, Girardinia cuspidata, Lysimachia davurica, Parasenecio hastata, Picris davurica, Saussurea parviflora and Veronicastrum sibiricum.

Manchurian Kalopanax septemlobus Forest These forests are confined to the middle and lower slopes of Changbaishan. Associated trees include several endemic species such as Abies holophylla (Pinaceae). Shrub layer species comprise Rubus crataegifolius, Viburnum burejaeticum and stunted specimens of Acer ukurunduense. Typical herb layer species are Lilium distichum, Syneilesis aconitifolia and the endemic or near endemic Adonis amurensis (Ranunculaceae).

Manchurian Celtis bungeana Forest These forests are confined to the western parts of Liaoning Province where they usually occupy dry, south-facing slopes between altitudes of 500 to 600 m. Other trees may include Acer truncatum, Tilia mongolica and Ziziphus spinosa. Major shrub layer taxa

Copyright © 2010 Peter Martin Rhind include Ampelopsis humulifolia, Rhamnus arguta and the endemic Spiraea pubescens (Rosaceae). Dominant in the herb layer are Carex humida, C. lanceolata, Cleistogenes nakai and C. caespitosa.

Manchurian Carpinus turczaninovii Forest These forests are confined to south-facing slopes on the Liaodong Peninsula and in western Liaoning Province. Carpinus turczaninovii strongly dominates the canopy but there may also be Fraxinus rhychophylla and Quercus mongolica. Shrub layer species include Celastrus flagellaris, Clerodendron trichotomum and Symplocos paniculata. The herb layer is quite sparse with a cover no more than about 10%. The main species are Calamagrostis arundinacea var. brachytricha, Carex lanceolata, Polygonatum odoratum and Spodiopogon sibiricus.

Manchurian Koelreuteria paniculata scrub These scrublands are found on south-facing slopes along the coast of the southeastern part of the Liaodong Peninsula. They reach a height of up to 2.5 m. The few other shrubs include Celtis bungeana. Herb layer species typically include Allium macrostemon, Artemisia argyi var. gracilis, while less common are Dentranthema lavandulifolium, Pinellia ternata and the endemic or near endemic Viola mandshurica (Violaceae).

Manchurian Pinus pumila scrub Pinus pumila forms mono-dominant scrub throughout northeastern and is the most cold tolerant of the pine species. It mainly occurs at high altitudes greater than 1200 m and in this BioProvince can be found on Changbai-shan and Daxingan-ling where it normally forms a low growing scrub never reaching more than about 2 m in height. The few associated canopy species include Betula ermani, B. nana subsp. exilis and Juniperus davurica. Common understorey species include Aquilegia flabellata var. pumila, Empetrum nigrum var. japonicum, Hedysarum alpinum, Hierchloë alpina, Pedicularis labradorica, Polygonum ajanense, Saxifraga bronchialis and Tilingia ajanensis.

Further information required.

References

Kolbek, J., Srutek, M. & Box, E. O. 2003. Forest Vegetation in Northeast Asia. Kluwer Academic Publishers.

Richardson, S. D. 1977. Forestry in communist China. The John Hopkins Press.

Wang, Chi-Wu. 1961. The forests of China with a survey of grassland and desert vegetation. Maria Moors Cabot Foundation. Publication No. 5.

Yang, X. & Xu, M. 2003. Biodiversity conservation in Changbai Mountain Biosphere Reserve, northeastern China: status, problems, and strategy. Biodiversity and Conservation, 12: 883-903.

Copyright © 2010 Peter Martin Rhind