54 the classification hierarchy is delimited by the physiognomy of forests, following the treatment of the US National Vegetation Classification (USNV

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1 8(2) 53-85, Vegetation Classification System of the Machilus- Castanopsis Zone in the Central-West Climatic Region of Taiwan 1 2 3, Ching-Yu Liou 1, Horng-Jye Su 2 and Yen-Hsueh Tseng 3, Endemic Species Research Institute, Jiji, Nantou, Taiwan 2 School of Forestry and Resource Conservation, National Taiwan University, Taipei, Taiwan 3 Department of Forestry, National Chung-Hsing University, Taichung, Taiwan * *Corresponding author 500-1,500m Abstract Sample stands from the Machilus-Castanopsis forest zone in the central-west climatic region of Taiwan were collected for the synthesis of a regional vegetation classification system. The upper level of

2 54 the classification hierarchy is delimited by the physiognomy of forests, following the treatment of the US National Vegetation Classification (USNVC) System. The lower two levels of the hierarchy, alliance and association, were based on floristic composition according to the concept of diagnostic species and floristic-sociological classification of the Braun-Blanquet approach. The original data were subjected to cluster analysis and two-way indicator species analysis (TWINSPAN) for grouping the stands into composite samples and summarization of species abundance. The matrix of the composite samples was re-arranged by the tabular comparison method for developing a hierarchy of the forest vegetation system. The resultant system consisted of 3 formations and 14 alliances, characterized respectively by Ficus septica, Phoebe formosana, Pasania konishii, Castanopsis fargesii, Daphniphyllum glaucescens subsp. oldhamii, Lithocarpus lepidocarpus, Rhododendron formosanum, Trema orientalis, Zelkova serrata, Juglans cathayensis, Carpinus kawakamii, Quercus variabilis, Alnus formosana, and Pinus taiwanensis. A total of 31 associations were proposed under the alliances. Key words: Taiwan, central-west climatic region, Machilus-Castanopsis zone, vegetation classification system Received: October 27, 2005 Accepted: March 20, Humboldt (1931)(1932) (1935) (1938, 1952) (1939) (1980) (floristic region) (physiognomy) (Alpine vegetation) (Abies zone) - (Tsuga-Picea zone) ( Quercus zone) (Machilus-Castanopsis zone) (Ficus-Machilus zone) ( ) ( 1992)

3 8(2) 53-85, (floristics) ( 1995, ) (Su 1984, , 1992, ) (US National Vegetation Classification, USNVC) (Grossman et al. 1998) (UNESCO) (UNESCO 1973) (evergreen broadleaved forest formation group) (evergreen coniferous forest formation group) (deciduous broadleaved forest formation group) 3 (formation group) (laurisilvae) (formation) 1,500-2,500m (diffused light) (submontane zone) 500-1,500m (Su 1984, 1985, , 2005) (character species) (differential species) (constant companion species) (diagnostic species) (diagnostic character) (floristic-sociological approach) ( 2004) (Westhoff and Maarel 1978) (synonym)

4 (Grossman et al. 1998) 2 2 ( ) ( ) (important value index, IVI)( 1983) () (octave scale)0-9 (Gauch 1982) ( 2003) 1( ) (two-way indicator species analysis, TWINSPAN) (Hill 1979 Gauch and Whittaker 1981 Belbin and McDonald 1993) (cluster) ( 1996) 30%

5 8(2) 53-85, (syntaxa) (association) (alliance) (order) (class) (division) (subassociation)(facies) (Westhoff and Maarel 1978) 2 1 ( ) (Braun-Blanquet 1932) (USNVC) (Flora of Taiwan) 2 association 1 alliance ( 1) ( ) (Submontane evergreen broadleaved forest) (subtropical rain forest)

6 58 1. Table 1. Vegetation classification system of Machilus-Castanopsis zone in central-west climatic region of Taiwan

7 8(2) 53-85, ( ) Table 1. Vegetation classification system of Machilus-Castanopsis zone in central-west climatic region of Taiwan (Cont.)

8 60 1. ( ) Table 1. Vegetation classification system of Machilus-Castanopsis zone in central-west climatic region of Taiwan (Cont.)

9 8(2) 53-85, ( ) Table 1. Vegetation classification system of Machilus-Castanopsis zone in central-west climatic region of Taiwan (Cont.)

10 62 1. ( ) Table 1. Vegetation classification system of Machilus-Castanopsis zone in central-west climatic region of Taiwan (Cont.)

11 8(2) 53-85, ( ) Table 1. Vegetation classification system of Machilus-Castanopsis zone in central-west climatic region of Taiwan (Cont.)

12 64 A-1 (Ficus septica alliance) 800m (lowland) (ecotone) (secondary forest) (Machilus japonica var. kusanoi - Ficus septica association) ( Machilus japonica var. kusanoi) (Ficus septica)(machilus zuihoensis) (Ficus nervosa) (Ficus fistulosa) (Ficus virgata) (Ficus variegata var. garciae) (Ficus irisana) (Bischofia javanica) (Dendrocnide meyeniana) (Ficus superba var. japonica)(cinnamomum camphora) - (Machilus kusanoi - Ficus vasculosa association) ( 1980) 800m m 30m cm (Machilus thunbergii) (Beilschmiedia erythrophloia) (Phoebe formosana) (Schefflera octophylla) J 10m 6-8m (Psychotria rubra) (Helicia formosana) (Maesa japonica) (Ardisia cornudentata subsp. cornudentata) (gap) (patch) 800m A-2 (Phoebe formosana alliance) (Turpinia formosana) (Lindera communis) (Lagerstroemia subcostata) (Oreocnide pedunculata) (Ardisia virens) (Pasania hancei var. ternaticupula) (Radermachia sinica) (Neolitsea konishii)(citrus tachibana) (Prunus zippeliana) (Murraya euchrestifolia) (Saurauia tristyla var. oldhamii) (Eurya strigillosa) (Pasania kawakamii)

13 8(2) 53-85, (Perrottetia arisanensis) 2. - (Ardisia virens - Phoebe formosana association) (Mallotus paniculatus) - (Turpinia formosana - Mallotus paniculatus type) ( 1997) - ( 1997) - ( 1997) 700-1,200m (Rhus succedanea) (Trema orientalis) (Cryptocarya chinensis) (Schima superba) (Zelkova serrata) 15-20m (Podocarpus fasciculus) 700-1,100m 3.- (Neolitsea konishii- Phoebe formosana association) - (Phoebe formosana - Turpinia formosana type) ( 1992) 500-1,000m 55% m (Meliosma rhoifolia) J (Eustigma oblongifolium) m 4.- (Saurauja tristyla var. oldhamii-phoebe formosana association) (Aralia bipinnata) (Litsea acuminata) -- (Machilus kusanoi-turpinia formosana-litsea acuminata domin-ance type) ( 1984) - (Schefflera octophylla - Machilus association) ( 1980) 700-1,200m (Prunus phaeosticta)

14 66 900m 5. - (Pasania kawakamii - Phoebe formosana association) (Machilus japonica) - (Pasania kawakamii- Phoebe formosana association) ( 1975) 1,000-1,500m (Lithocarpus lepidocarpus) (Lithocarpus amygdalifolius) (Cyclobalanopsis stenophylloides) (Sapindus mukorossii) (Trochodendron aralioides) (Sloanea formosana) (Mallotus japonicus) J 1,000-1,500m A-3 (Pasania konishii alliance) (Pasania konishii) ( Litsea rotundifolia var. oblongifolia) (Pinus morrisonicola) (Cinnamomum osmophloeum)(styrax suberifolia) (Rhododendron breviperulatum) (Litsea hypophaea) (Gardenia jasminoides) (Diospyros eriantha) (Syzygium formosanum)(ardisia sieboldii) (Wendlandia uvariifolia) (Viburnum formosanum) ( Ormosia formosana) ( Sapium discolor) (Randia cochinchinensis) (Cryptocarya concinna)(nageia nagi) (Schefflera arboricola) (Cyclobalanopsis globosa) (Rhododendron ovatum) (Prunus campanulata) (Fraxinus insularis) (Melicope pteleifolia) (Calocedrus macrolepis var. formosana) (Homalium cochinchinensis) (Litsea cubeba) (Broussonetia kazinoki) (Melanolepis multiglandulosa) (Lindera aggregata) (Champereia manillana) ( Cyclobalanopsis pachyloma) (Camellia japonica) 6. - (Ormosia formosana-pasania konishii association) ( Pinus taiwanensis) (Gordonia axillaris)

15 8(2) 53-85, (Pinus taiwanensis-ormosia formosana- Gordonia axillaris type) ( 1992) m (Elaeocarpus sylvestris) (Daphniphyllum glaucescens ssp. oldhamii) (Diospyros morrisiana) (Itea parviflora) (Tricalysia dubia) (Callicarpa formosana) (Syzygium buxifolium) m 7. - (Sapium discolor - Pasania konishii association) - (Sapium discolor - Pasania konishii type) ( 1992) m (Alniphyllum pterospermum) (Helicia cochichinensis) (Castanopsis fargesii) (Symplocos stellaris)(engelhardia roxburghiana) m 8. - (Cyclobalanopsis globosa - Pasania konishii association) - (Cyclobalanopsis globosa - Pasania konishii type) ( 1992) m

16 68 (Carpinus kawakamii) ( Neolitsea parvigemma) (Cyclobalanopsis glauca) ( Firmiana simplex) (Adinandra formosana) (Liquidambar formosana) (Mallotus philippensis) (Ternstroemia gymnanthera) (Ligustrum liukiuense) (Maesa perlaria var. formosana) (Vaccinium bracteatum) m 500-1,000m % 25-60% (Pasania synbalanos) J (Malus doumeri) (Taiwania cryptomerioides) m 9.- (Melicope pteleifolia - Pasania konishii association) - (Cinnamomum camphora-daphniphyllum pentandrum type) ( 1993) - (Pasania rhombocarpa-pinus taiwanensis type) ( 1993) - (Mallotus paniculatus- Litsea kostermansii type) ( 1993) - (Trema orientalis- Mallotus paniculatus association) ( 1980) 10.- (Cyclobalanopsis pachyloma - Pasania konishii association) - (Cyclobalanopsis pachyloma - Styrax suberifolia type) ( 1992) 800-1,100m % 60% (Acer serrulatum)

17 8(2) 53-85, ,080m A-4 (Castanopsis fargesii alliance) ( Euonymus laxiflorus) (Meliosma squimulata) (Ilex goshiensis) (Cyclobalanopsis longinux) (Pasania harlandii)(cinnamomum subavenium) ( Neolitsea aciculata var. variabillima) (Ilex formosana) (Symplocos sonoharae) (Pyrenaria shinkoensis) (Ardisia quinquegona) ( Ilex lonicerifolia) (Ficus formosana) (Blastus cochinchinensis) (Osmanthus matsumuranus) (Castanopsis cuspidata var. carlesii f. sessilis) (Castanopsis kawakamii) (Limlia uraiana) (Pasania nantoensis) ( Podocarpus nakaii)(antidesma japonicum var. acutisepalum) ( Camellia furfuracea) (Camellia salicifolia) (Euonymus tashiroi) (Ilex pubescens) (Ilex rotunda) (Helicia rengetiensis) (Castanopsis eyrei) (Embelia laeta var. papilligera)(lasianthus wallichii)(litsea acutivena) (Pachycentria formosana) (Lasianthus curtisii) (Symplocos congesta)(lasianthus fordii) (Ilex ficoidea) (Myrsine seguinii) (Dendropanax dentiger) ( Lasianthus japonicus) ( Microtropis japonica) (Vaccinium randaiense) (Castanopsis cuspidata var. carlesii f. sessilis-castanopsis fargesii association) - - (Engelhardtia roxburghiana-castanopsis borneensis- Castanopsis carlesii var. sessilis dominance type) ( 1984) --- (Castanopsis carlesii var. sessilis- Beilschmiedia chinensis-cyclobalanopsis pachyloma-pasania

18 70 rhombocarpa subtype) ( 1984) --- (Machilus thunbergii - Rhododendron ellipticum -Gordonia axillaris - Elaeocarpus sylvestris subtype) ( 1984) --- (Castanopsis carlesii var. sessilis - Castanopsis hystrix - Helicia formosana - Pasania konishii subtype) ( 1984) 500-1,500m ( Elaeocarpus japonicus) 15-25m (Illicium arborescens) ( Rhododendron ellipticum) (Distyliopsis dunnii)(eurya rengechiensis) 700-1,400m (Castanopsis eyrei - Castanopsis fargesii association) ( Symplocos nokoensis)(rhododendron mariesii) (Ilex micrococca) - (Cinnamomum randaiense - Mallotus paniculatus type) ( 1993) 800-1,600m %45-85% J (Cunninghamia konishii) 800-1,600m (Symplocos congesta - Castanopsis fargesii association) (Diospyros oldhamii) - (Cinnamomum randaiense- Elaeocarpus japonicus type) ( 1992) - (Cinnamomum

19 8(2) 53-85, randaiense- Michelia compressa subtype) ( 1992) - (Elaeocarpus japonicus- Rhododendron formosanum subtype) ( 1992) - (Neolitsea acuminatissima-castanopsis carlesii subtype) ( 1992) - (Diospyros oldhamii- Castanopsis carlesii type) ( 1992) 1,000-1,600m % 60-80% (Castanopsis cuspidata var. carlesii) 12-18m J (climax) 1,000-1,550m (Lasianthus fordii -Castanopsis fargesii association) - (Litsea acuminata - Schefflera octophylla type) ( 1997) - ( 1997) - ( 1997) - ( 1997) 1,000-1,500m ( Meliosma callicarpaefolia) 15-20m J (Michelia compressa) 1,100-1,400m 15.- (Myrsine seguinii - Castanopsis fargesii association) (Skimmia reevesiana) (Symplocos caudata) (Ilex goshiensis) (Rhododendron formosanum) - (Cyclobalanopsis longinux - Rhododendron formosanum type) ( 1997) --- (

20 ) - ( 1997) ( 1997) 1,200-1,800m (Daphniphyllum himalaense subsp. macropodum) ( Damnacanthus angustifolius) J 1,300m 1,834m A-5 (Daphniphyllum glaucescens subsp. oldhamii alliance) (Litsea akoensis) (Tetradium ruticarpum) (Machilus zuihoensis var. mushaensis) 16.- (Cyclobalanopsis stenophylloides - Daphniphyllum glaucescens subsp. oldhamii association) - - (Cyclobalanopsis stenophylloides var. stenophylloides - Daphniphyllum glaucescens var. oldhamii - Litsea acuminata dominance type) ( 1984) 1,100-1,500m (Acer albopurpurascens) (Pittosporum illicioides) 1,200m 17.- (Machilus zuihoensis - Daphniphyllum glaucescens subsp. oldhamii association) (Fraxinus griffithii) (Glochidion rubrum) (Cinnamomum insulari-montanum) (Machilus zuihoensis forest type)

21 8(2) 53-85, ( 1999) ( 1999) - ( 1999) 1,000-1,700m m (Alnus formosana) (Acer kawakamii) L J J 1,165-1,670m (Adinandra formosana-daphniphyllum glaucescens subsp. oldhamii association) (Gordonia axillaris forest type) ( 1999) 1,000-1,700m 1,145-1,175m A-6 (Lithocarpus lepidocarpus alliance) 1,100-1,800m (montane) (Beilschmiedia erythrophloia - Lithocarpus lepidocarpus association) (Cinnamomum kanehirae) (Tetrapanax papyriferus) (Vaccinium dunalianum var. caudatifolium) (Idesia polycarpa) (Beilschmiedia erythrophloia - Machilus japonica - Lithocarpus lepidocarpus - Turpinia formosana dominance type) ( 1984) (Beilschmiedia erythrophloia - Machilus

22 74 japonica-turpinia formosana-litsea nantoensis subtype) ( 1984) (Cinnamomum osmophloia-machilus kusanoi - Litsea akoensis-turpinia formosana subtype) ( 1984) - (Beilschmiedia erythrophloia-turpinia formosana forest type) ( ) - (Lithocarpus lepidocarpus-litsea akoensis subtype) ( 1991) - (Lithocarpus lepidocarpus - Beilschmiedia erythrophloia forest type) ( 1994) - (Lithocarpus lepidocarpus - Beilschmiedia erythrophloia subtype) ( 1994) - (Aphananthe aspera - Litsea akoensis subtype) ( 1994) - (Lithocarpus lepidocarpus - Turpinia formosana type) ( 1993) - (Machilus zuihoensis - Litsea acuminata vegetation type) ( 1994) 1,000-2,000m % 40-80% 1m (Symplocos modesta) J 70 ( Embelia lenticellata)(helwingia japonica subsp. formosana) (Sinopanax formosana) A-7 (Rhododendron formosanum alliance) 1 - (Vaccinium wrightii)(lyonia ovalifolia) (Vaccinium wrightii-rhododendron formosanum association) (Pieris taiwanensis) - (Rhododendron formosanum - Lyonia ovalifolia var. lanceolata type) ( 1992) - (Cinnamomum randaiense - Elaeocarpus japonicus type) ( 1992)

23 8(2) 53-85, (Elaeocarpus japonicus - Rhododendron formosanum subtype) ( 1992) 1,300-2,050m 80% 5-10m (Rhododendron oldhamii) ( ) (Submontane deciduous broadleaved forest formation) A-8 (Trema orientalis alliance) 800m (Macaranga tanarius) (Aphananthe aspera) (Paulownia taiwaniana) (Adina racemosa) (Glochidion zeylanicum) (Glochidion acuminatum) (Broussonetia papyrifera) (Macaranga tanarius - Trema orientalis association) (Celtis sinensis) - (Macaranga tanarius - Trema orientalis forest type) ( 2002) m %75% 1,500 /ha 7m 5-20 cm10-35 cm (Vitex quinata) 675m 810m 22.- (Ficus irisana - Trema orientalis association) - (Trema orientalis-ficus irisana type) ( 1993)

24 76 500m % 7-8m cm J J 500m A-9 (Zelkova serrata alliance) (Pistacia chinensis) (Ulmus parvifolia) (Koelreuteria henryi) (Rhamnus parvifolia)(symplocos konishii) (Acacia confusa) (Ligustrum pricei) (Camellia nokoensis) (Glycosmis citrifolia) (Melia azedarach)(rhamnus nakaharae) ( Osmanthus lanceolatus) (Bridelia balansae) (Scolopia oldhamii) (Pourthiaea lucida) 23.- (Pistacia chinensis - Zelkova serrata association) -- (Cyclobalanopsis glauca - Pistacia chinensis - Zelkova serrata forest type) ( 2002) 500-1,200m %60-90% 1,000 /ha 10-15m 5-20 cm (Murraya paniculata) (Celtis biondii) 725-1,195m (Symplocos konishii - Zelkova serrata association) (Bridelia tomentosa) (Morus australis) (Acacia confusa forest type) (

25 8(2) 53-85, ) 500-1,000m %50-70% 1,280 /ha 750m 7m10-30 cm (Celtis formosana) 750m 818m (Acer albopurpurascens - Zelkova serrata association) --- (Lagerstroemia subcostata - Acacia confusa - Acer albopurpurascens - Zelkova serrata var. serrata dominance type) ( 1984) - (Zelkova serrata- Lagerstroemia subcostata association) ( 1980) m 6-10m (Rhus javanica var. roxburghiana) m 26.- (Bridelia balansae-zelkova serrata association) (Zelkova serrata type) ( 1992) m m A-10 (Juglans cathayensis alliance) (Juglans cathayensis association) ( Juglans

26 78 cathayensis) ( Lindera megaphylla) ( Hydrangea aspera) (Ulmus uyematsui) (Trichodesma calycosum) (Ehretia acuminata) (Ficus erecta var. beecheyana) (Fatsia polycarpa) (Juglans cathayensis forest type) ( 1991) 1,500m % 800-1,500m 1,300-1,500m A-11 (Carpinus kawakamii alliance) 28. (Carpinus kawakamii association) (Platycarya strobilacea) 900-1,100m % 60-90% 15-20m (Deutzia pulchra) (Pluchea indica) (Alangium chinense) ( Litsea coreana) ,090m A-12 (Quercus variabilis alliance) 29. (Quercus variabilis association) ( Quercus variabilis) ( Elaeagnus thunbergii) (Coriaria japonica subsp. intermedia) (Pouzolzia elegans) (Zanthoxylum ailanthoides) (Stachyurus himalaicus) (Tetradium glabrifolium) - (Cyclobalanopsis glauca

27 8(2) 53-85, Quercus variabilis vegetation type) ( 1994) 1,000-1,700m % 10-20m (Dodoneae viscosa) 1,300m 1,430m 10-20m 5-15 cm 1,190m 1,135m 1,780m ( ) (Submontane evergreen coniferous forest formation) A-14 (Pinus taiwanensis alliance) A-13 (Alnus formosana alliance) 30. (Alnus formosana association) (Debregeasia orientalis) (Boehmeria densiflora) (Alnus japonica forest type) ( 2002) 1,100-1,800m %40-80% 3,000 /ha 31. (Pinus taiwanensis associaton) - (Pinus taiwanensis- Alnus japonica forest type) ( 2002) 800-1,200m % 60% 1,175m

28 80 (sociological progression)(braun-blanquet 1964) (affinity) () ( ) 500-1,500m 500m 1,500m ( 2003) ( ) ( ) ( 2003)

29 8(2) 53-85, ( 1) 1,500m m ,200m 1. Fig. 1. Succession of associations of Machilus-Castanopsis zone in central-west climatic region of Taiwan.

30 ( Damnacanthus indicus) (Zanthoxylum nitidum) ( Rhamnus formosana) 1,200m - (1,000m ) (1,000m ) (Strobilanthes flexicaulis) - ( 1) 2 ()( )(Su 1985) ( 2003) () ( ) 2 -

31 8(2) 53-85, (Cyclobalanopsis salicina)- - - ( 2003) ( 2003) A B A B A B A B (narrow endemic species)( 2003) (I) ( ) 111:

32 (1): ( ) 18: (I) (4): (3): ( ) 9(2): (IV) 39: (1): (4):

33 8(2) 53-85, : Belbin, L. and C. McDonald Comparing three classification strategies for use in ecology. Journal of Vegetation Science 4: Braun-Blanquet, J Plant Sociology. (Transl. by G. D. Fuller and H. S. Conad). New York. xviii 439 pp. Reprint Braun-Blanquet, J Pflanzensoziologie, Grundzuge der Vegetationskunde. 3rd ed. Springer, Wien-New York. 865 pp. Gauch, H. G. and R. H. Whittaker Hierarchical classification of community data. Journal of Ecology 69: Gauch, G. H Multivariate analysis in community ecology. Cambridge University. Press, Cambridge. 298 pp. Grossman, D. H., D. Faber-Langendoen, A. S. Weakley, M. Anderson, P. Bourgeron, R. Crawford, K, Goodin, S. Landaal, K. Metzler, K. Patterson, M. Pyne, M. Reid and L. Sneddon Terrestrial vegetation of the United States, vol. I. The natural vegetation classification system: Development, status and application. The Nature Conservancy, Virginia. 119 pp. Hill, M. O TWINSPAN A FORTRAN program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. Ecology and Systematics, Cornell University. Ithaca, New York. 90 pp. Su, H. J Studies on the climate and vegetation types of the natural forests in Taiwan. (II). Altitudinal vegetation zones in relation to temperature gradient. Quarterly Journal of Chinese Forestry 17(4): Su, H. J Studies on the climate and vegetation types of the natural forests in Taiwan. (III). A scheme of geographical climatic regions. Quarterly Journal of Chinese Forestry 18(3): Su, H. J Species diversity of forest plants in Taiwan. Institute of Botany, Academia Sinica Monogrraph Series No.14: UNESCO International classification and mapping of vegetation. Series 6. Ecology and conservation. United Nations Educational, Scientific and Cultural Organization, Paris, France. Westhoff, V. and E. van der Maarel The Braun-Blanquet Approach. In: Whittaker, R. H. (eds.). Classification of plant communities. Dr. W. Junk by Pub. The Hague, Boston. pp

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