Chamaecyparis Montane Cloud Forest in Taiwan: Ecology and Vegetation Classification

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Chamaecyparis montane cloud forest in Taiwan: ecology and vegetation classification

Ching-Feng Li, David Zelený, Milan Chytrý, Ming-Yih Chen, Tze-Ying Chen, Chyi-Rong Chiou, Yue-Joe Hsia, Ho-Yih Liu, Sheng-Zehn Yang, et al.

Ecological Research

ISSN 0912-3814 Volume 30 Number 5

Ecol Res (2015) 30:771-791 DOI 10.1007/s11284-015-1284-0

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1 23 Author's personal copy Ecol Res (2015) 30: 771–791 DOI 10.1007/s11284-015-1284-0

BIODIVERSITY IN ASIA

Ching-Feng Li • David Zeleny´• Milan Chytry´ Ming-Yih Chen • Tze-Ying Chen • Chyi-Rong Chiou Yue-Joe Hsia • Ho-Yih Liu • Sheng-Zehn Yang Ching-Long Yeh • Jenn-Che Wang • Chiou-Feng Yu Yen-Jen Lai • Ke Guo • Chang-Fu Hsieh Chamaecyparis montane cloud forest in Taiwan: ecology and vegetation classiﬁcation

Received: 21 February 2014 / Accepted: 1 June 2015 / Published online: 14 July 2015 Ó The Ecological Society of Japan 2015

Abstract Montane cloud forest is one of the most evergreen broad-leaved forests; it is found at higher endangered ecosystems. However, there are few com- altitudes and is more influenced by the summer mon- prehensive studies on the distribution of subtropical soon than the other alliance. Five associations are de- montane cloud forest (SMCF). Chamaecyparis forest is fined within this alliance. The alliance of Pasanio one type of SMCF in Taiwan, distributed across the kawakamii-Machilion japonicae growing on slopes and whole island. This study describes eleven types of this in valleys contains evergreen broad-leaved forests or forest in Taiwan based on the Braun-Blanquet ap- forests with a mixture of coniferous and evergreen proach. Plots were selected from the National Vegeta- broad-leaved species. Six associations can be determined tion Database of Taiwan. Two alliances were defined, under the alliance of Pasanio kawakamii-Machilion both of which belong to the order Fagetalia hayatae. japonicae. Classification of each syntaxon was formal- Topography and altitude explain the contrasting habitat ized using Cocktail Determination Key. requirements of these two alliances, whereas seasonality of moisture, soil properties and altitude explain differ- Keywords Cocktail Determination Key Æ Seasonality of ences in floristic composition at the association level. moisture Æ Species group Æ Supervised classification Æ The alliance of Chamaecyparidion formosanae on slopes Topography and ridges includes coniferous or mixed coniferous and

Electronic supplementary material The online version of this article (doi:10.1007/s11284-015-1284-0) contains supplementary material, which is available to authorized users. C.-F. Li Æ D. Zeleny´ Æ M. Chytry´ Department of Botany and Zoology, Masaryk University, S.-Z. Yang Æ C.-L. Yeh Kotla´ rˇ ska´ 2, Brno, Czech Republic Department of Forestry, National Pingtung University of Science and Technology, Shue-Fu Rd. 1, Pingtung, Taiwan M.-Y. Chen Department of Life Sciences, National Chung Hsing University, J.-C. Wang Kuo-Kuang Rd. 250, Taichung, Taiwan Department of Life Science, National Taiwan Normal University, Ting-Chou Rd. 88, Taipei, Taiwan T.-Y. Chen Æ C.-F. Yu Department of Nature Resources, National Ilan University, Y.-J. Lai Shen-Lung Rd. 1, Ilan, Taiwan The Experimental Forest, National Taiwan University, Chien-Shan Rd. 12, Nantou, Taiwan C.-R. Chiou School of Forestry and Resource Conservation, National Taiwan K. Guo University, Roosevelt Rd. 1, Taipei, Taiwan Institute of Botany, Chinese Academy of Sciences, Xiangshan Nanxincun 20, Beijing, China Y.-J. Hsia Institute of Nature Resources, National Dong Hwa University, C.-F. Hsieh (&) Da-Hsueh Rd. 1, Hualien, Taiwan Institute of Ecology and Evolutionary Biology, National Taiwan University, Roosevelt Rd. 1, Taipei, Taiwan H.-Y. Liu E-mail: [email protected] Department of Biological Sciences, National Sun Yat-Sen Tel.: +886-02-33662474 University, Lien-Hai Rd. 70, Kaohsiung, Taiwan Author's personal copy 772

cutting of any primary forests in Taiwan. Horng et al. Introduction (2000) estimated that 60 % of the Chamaecyparis forest has been cut since the early 20th century, and about Montane cloud forests are one of the world’s most 48,000 ha remain (according to Chen 2001, however, the endangered ecosystems because of their sensitivity to latter figure might be an optimistic overestimation). changes in unique ecological conditions (Hamilton et al. Across the altitudinal range of nearly 4000 m in 1995; Bruijnzeel et al. 2010). The distribution of mon- Taiwan, Chamaecyparis species mostly grow between tane cloud forests is highly fragmented, and substantial 1500 and 2500 m a.s.l. This altitudinal distribution is isolation of these fragments is assumed to promote associated with two ecological features. First, there is a speciation and endemism. Most studies have focused on prominent fog formation caused by the uplift of air montane cloud forests in tropical regions. There are masses from the sea, which occurs almost daily. A cli- several networks established for the protection and matic station at Yuan-Yang Lake at altitude 1700 m study of tropical montane cloud forest (TMCF), such as (Fig. 1) records an annual average of 342 days with fog the World Conservation Monitoring Centre, Tropical events (Lai et al. 2006). Second, the mid-altitude range Montane Cloud Forest Initiative, and UNESCO Inter- 1500–2500 m is a transition zone for forest physiog- national Hydrological Programme (Bruijnzeel et al. nomy. Above this range, the climate corresponds to a 2010). In contrast, subtropical montane cloud forest cool-temperate or high-montane zone (Su 1984) and (SMCF) has been less well studied, although its bio- forests are dominated by coniferous trees such as Abies logical and conservation value is not less significant. kawakamii, Picea morrisonicola and Tsuga chinensis var. In subtropical eastern Asia, a large proportion of formosana (Lin et al. 2012). The upper limit of this mid- montane cloud forests are dominated by deciduous altitude range is a hard boundary for evergreen broad- broad-leaved or coniferous trees (e.g., Hou 1983;Su leaved species. Below this range is a subtropical or 1984; Da et al. 2009). These can be pure coniferous submontane zone and foothills, with forests dominated forests dominated by Abies spp. or Picea spp., or mixed by evergreen broad-leaved species of mainly Fagaceae, forests of evergreen broad-leaved and coniferous (or Lauraceae, Moraceae, and Theaceae (Su 1984; Li et al. deciduous) trees. The common dominant genera in the 2013). mixed forests include the conifers Chamaecyparis, The aim of this study was to understand which veg- Cryptomeria, Cunninghamia, Picea, Pseudotsuga, Tai- etation types of Chamaecyparis forest grow in Taiwan wania and Tsuga, and the deciduous Fagus. These are and their relationships to important environmental fac- different from the TMCFs, which are dominated only by tors. Here, the Chamaecyparis forest is defined as one evergreen broad-leaved trees (Bruijnzeel et al. 2010). In with Chamaecyparis spp. as diagnostic, dominant or Japan, a typical SMCF is a coniferous and evergreen frequent species. Although several vegetation classifica- broad-leaved mixed forest dominated by Chamaecyparis tion studies in Taiwan focused at least partially on the obtusa, Cryptomeria japonica and Tsuga sieboldii Chamaecyparis forest (see ESM 1 for a literature re- (Miyawaki 1980). In Taiwan, the most representative coniferous and deciduous broad-leaved genera of SMCF are Chamaecyparis and Fagus (Suzuki 1954;Su1984; Chen 2001; Li et al. 2013). Unlike the Fagus forest, which has a very limited distribution in Taiwan (Hsieh 1989; Hukusima et al. 2005), the Chamaecyparis forest grows across the whole island, varying in species composition and habitat conditions. Adult trees of both native Chamaecyparis species, Chamaecyparis formosensis Matsum. and Chamaecyparis obtusa (Sieb. et Zucc.) Endl. var. formosana (Hayata) Rehder, are usually taller than 30 m, with trunks commonly thicker than 1 m at breast height. The Chamae- cyparis forest in Taiwan has received widespread attention from local inhabitants, as evidenced by many legends of the native Taiwanese people. Before the 20th century, these people typically preserved forests with huge Chamaecyparis trees in their ethnic group territory as sacred places, where the spirits of their ancestors rest in peace. No one was allowed to enter these places to collect anything or hunt, which helped preserve the Fig. 1 Geographic locations of Taiwan and two climatic stations pristine nature of the Chamaecyparis forest. However, in with mountain fog observation. Gradient of black and white colors indicates altitudes in m a.s.l. Codes are for the following the 20th century, these forests were extensively logged by ecoregions: NE northeastern, EN north part of eastern, ES south immigrants from mainland China and Japan because of part of eastern, SE southeastern, SSW south part of southwestern, their valuable timber. After 1992, a law prohibited the SW southwestern, CW central-western, and NW northwestern Author's personal copy 773 view), the ecological pattern of that forest across all tation dataset stored in the National Vegetation Data- Taiwan has remained ambiguous because of non-repre- base of Taiwan and used the Cocktail Determination sentative samples, a lack of comparison across the entire Key to define unequivocal assignment rules for each island, and a lack of standardized nomenclature com- vegetation type. parable among studies (Su 2002). A comprehensive study on the vegetation classification of Chamaecyparis forest in Taiwan would assist observation and docu- Materials and methods mentation of its natural distribution. An important requirement of modern vegetation Study area classification is the transparent and formalized proce- dure of assigning individual plots to particular vegeta- Taiwan (21°55¢–25°20¢N, 119°30¢–122°00¢E) is a moun- tion types (Jennings et al. 2009; Chytry´ et al. 2011). tainous island in the subtropical region of eastern Asia There are several statistical techniques that apply (Fig. 1). Backbone mountains roughly run from north to objective classification (Kent 2012; Peet and Roberts south. The climate is mainly controlled by altitude and 2013). However, few of these offer unequivocal assign- monsoon systems. Altitude is strongly correlated with ment rules, especially those with ecological meaning. temperature, which has a lapse rate of 0.55 °C/100 m. In The Braun-Blanquet approach, using plot-based infor- summer, the warm southwestern monsoon and typhoons mation on floristic composition to produce a hierarchi- bring abundant precipitation to the entire island. In cal classification system, is one of the widely used winter, the cool northeastern monsoon brings moderate methods for classifying vegetation (Mueller-Dombois precipitation to only the windward slopes. The area that and Ellenberg 1974; van der Maarel and Franklin 2013). is not influenced by the winter monsoon has a dry period In that approach, presence or absence of several species of 2–6 months. Su (1985) classified Taiwan into seven or species groups is the key argument in the definition of ecoregions, based on climatic data. Here, we separate the a vegetation type representing a particular habitat. Henchun Peninsula in the southernmost region from the Bruelheide (1997) developed the Cocktail method based southwestern region created by Su (1985), defining west- on combination of the presence or absence of specific ern part of the peninsula as the eighth ecoregion (Fig. 1), ecologically meaningful species groups. This method which is characterized by tropical flora and strong wind provides unequivocal rules for assigning plots to vege- in winter. In the lowland of the northeastern region, the tation types (e.g., Bruelheide and Chytry´ 2000; Kocˇ ı´ winter precipitation ratio is the largest (more than 10 %), et al. 2003; Janisˇ ova´ and Du´ brakova´ 2010; Landucci whereas it is the smallest in the lowland of the southwest et al. 2015). Ecologically meaningful means that we (http://www.cwb.gov.tw/V7e/climate/). The smaller the consider species groups that represent a combination of winter precipitation ratio, the longer the winter drought species often occurring together in habitats with similar period at altitudes below 1500 m. There is no dry season environmental features (e.g., stony or acid). The occur- in any part of the island at altitudes above 1500 m, be- rence of such a specific species group indicates that the cause of fog formation at altitudes 1500–2500 m and habitat has a feature related to this species group. frequent precipitation above 2500 m. Moreover, it is reasonable to predict the presence of that The frequency of fog in the montane cloud zone group if habitat conditions are known. One of the main varies seasonally between ecoregions because of the challenges in constructing the Cocktail formulas is the monsoon regime. Ecoregions influenced by the winter search for ecologically meaningful species groups. In monsoon are covered by heavy fog for a longer period well-surveyed regions with a long history of vegetation than other ecoregions. Yuan-Yang Lake weather sta- studies such as Central Europe, definition of the species tion, in an area affected by the winter monsoon (Fig. 1), groups is based on knowledge stored in large and inte- has fog events 40 % of the hours in a year (Lai et al. grated vegetation databases and expert systems (Chytry´ 2006). From October to February, there are fog events 2007). In regions with a lack of extensive vegetation more than 50 % of the hours per month, and over one- databases and expert knowledge of the species groups third of them are heavy fogs. Xi-Tou weather station and their representative ecological meanings, such an (Wey et al. 2011), which is in an area not influenced by approach is more challenging. For this purpose, Li et al. the winter monsoon, has fog events 25 % of annual (2013) proposed the Cocktail Determination Key, which hours. Except for very few fog events in November and is a set of Cocktail formulas defining different vegetation very heavy ones in March, all months have a similar types. Species groups and vegetation units are generated hourly ratio of such events. iteratively while plots and species are sorted in a species- by-site table. In the present study, we asked the following questions: (1) Which environmental factors are the most Vegetation data and environmental variables important determinants of floristic composition in the Chamaecyparis forest of Taiwan? (2) What are ecologi- Vegetation plots were taken from the National Vegeta- cally meaningful Chamaecyparis forest types in Taiwan? tion Database of Taiwan (AS-TW-001), containing 3564 We addressed these questions using an extensive vege- plots mostly of size 20 · 20 m, which were sampled over Author's personal copy 774

2003–2007. The plot distribution covers all Taiwan and and R v. 3.0.1 (R Core Team 2013) were used for is representative of various ecoregions and altitudes. All manipulating and calculating the environmental factors. vascular plants were recorded, with estimated cover of shrubs, herbs, lianas and epiphytes, plus the cover of trees calculated from their diameters at breast height (Curtis 1959). Both estimated and calculated values were Classification of vegetation expressed as percentages. Information on life forms of all species and leaf types Association is the basic vegetationunitintheBraun- (coniferous, deciduous broad-leaved, or evergreen Blanquet system, whereas alliance is a vegetation unit broad-leaved) of tree species were compiled from the at a higher hierarchical level than association. We de- Flora of Taiwan (Huang and Hsieh 1994–2003). The fined association here as a community unit with proportion of each leaf type in the plots was calculated homogeneous floristic composition and similar habitat to compare the physiognomic structure of different conditions among plots (Mueller-Dombois and Ellen- vegetation types. This proportion was defined as the sum berg 1974). We used two criteria to group associations of cover of one leaf type in a plot divided by the sum of into alliances: (1) associations under the same alliance cover of all trees in that plot. had the same dominant species from several species, Environmental factors were recorded in the field or and (2) associations under the same alliance had at least derived from digital maps in geographic information one defined species group in common. Recognition of systems (GIS; see GIS-derived environmental factors in each association of Chamaecyparis forest and con- ESM 2 for more detail). In the field, altitude, slope struction of their Cocktail Determination Key (Ap- inclination, aspect and geographic coordinates of plots pendix S1 in Li et al. 2013) were achieved by the were measured, and rock cover and rockiness were application of several numerical classification methods estimated in each plot. Topography was recorded on a and field experience. Whether the plots should be 1–6 ordinal scale (1 = ridge, 2 = upper slope, classified as belonging to a particular association was 3 = middle slope, 4 = lower slope, 5 = valley, and determined by expert knowledge, based on both envi- 6 = plain) and was used as a surrogate for soil water ronmental factors and species composition as defined availability and light input (smaller values indicate drier by the Cocktail formula. Details of this process are habitats with greater light input; see Sørensen et al. described in the section Vegetation classification and the 2006). Rock cover was estimated as the percentage of construction of Cocktail Determination Key in ESM 2. area covered by stones larger than 10 cm inside the plot, The nomenclature of associations and alliances follows and rockiness was estimated as the percentage content of Weber et al. (2000). stones with sizes 1–10 cm in the soil. Before data anal- Results of classification are presented in a synoptic ysis, aspect was transformed into southernness by the table that summarizes the similarity or difference of cosine of aspect (Geiger 1966). Southernness can be used floristic composition between vegetation types (Mueller- to reflect the monsoon system affecting a plot, with Dombois and Ellenberg 1974). Diagnostic species were larger values indicating the summer monsoon and lower determined by their fidelity as represented by the phi ones the winter monsoon. Canopy cover and height were coefficient (U) of association, computed with the group estimated in the field, and the former was used as a size equalized to 5 % of the full dataset (Tichy´ and surrogate for light conditions in the understory. Alto- Chytry´ 2006). Fisher’s exact test was used to establish gether, seven environmental variables were used: alti- whether the concentration of species occurrence in a tude, canopy cover, inclination, rock cover, rockiness, particular vegetation type was significant at P < 0.001 southernness, and topography. (Chytry´ et al. 2002). Other variables such as whole-light sky, monthly Nonmetric multidimensional scaling (NMDS; Krus- precipitation, cloud frequency (Sklena´ rˇ et al. 2008), soil kal 1964) based on Bray-Curtis dissimilarities and moisture (Gillies et al. 1997; Sandholt et al. 2002), and square-root transformed species data was used to visu- monthly mean temperature (Lai et al. 2012) were ob- alize relationships among vegetation types and rela- tained from a digital terrain model (DTM; tionships between vegetation types and environmental http://www.csrsr.ncu.edu.tw), global climate data (Hij- factors. Environmental factors were related to the first mans et al. 2005), or the Moderate Resolution Image two axes of NMDS using multiple regression (with the Spectroradiometer (MODIS; modis.gsfc.nasa.gov). relationship quantified by the adjusted coefficient of 2 Whole-light sky is a commonly used environmental determination (Radj) and tested by permutation testing factor in vegetation study in Taiwan. Its values are re- with F-values as the test statistics). lated to topographic shading and the input of solar en- Species data were manipulated and the synoptic table ergy. This term is a synonym of sky-view factor in Lai was prepared using the program JUICE v. 7.0.84 (Tichy´ et al. (2010). All these variables used herein represented 2002; http://www.sci.muni.cz/botany/juice). The ordi- the climate, energy input, light conditions in the under- nation diagram and related analyses were computed story and physical soil conditions, with seasonal differ- using R program v. 3.0.1 (R Core Team 2013) and the ences at each plot. ArcGIS v. 9.0 (http://www.esri.com) library package vegan (Oksanen et al. 2013). Author's personal copy 775

Results

Relationship between ﬂoristic composition and environmental factors

As a complex gradient, altitude was the most important variable, closely related to the variation in floristic composition in the ordination space (Fig. 2a). All 21 environmental factors except the ratio of summer precipitation and southernness were significantly related to the first two axes of NMDS. In the ordination diagram, 2 only factors with the largest Radj value were drawn if several factors were strongly correlated with each other. In that case, among altitude and annual precipitation, the former was chosen. Among the three variables representing soil moisture, annual mean soil moisture was selected, and presented as soil dryness in Fig. 2a because of the negative relationship between the values and soil moisture itself. Among topography and rockiness, the former was chosen. Among monthly mean July temperature, annual mean temperature, monthly mean January temperature, warmth index and annual mean cloud frequency, monthly mean July temperature was selected. The difference between habitats of the Chamaecyparis formosensis dominated forest type and C. obtusa var. formosana dominated forest type is mainly related to topography and related factors such as inclination, rock cover, and whole-light sky. These environmental factors can be represented by the first axis of NMDS. On the ridges (left side of the ordination diagram in Fig. 2a), there is typically well-developed soil and the whole-light sky value is large, indicating that the habitats are less shaded by the surrounding topography than those in Fig. 2 Syntaxa and environmental factors on the ordination diagram of non-metric multidimensional scaling (NMDS). a Two alliances and valleys (right side of the ordination diagram). The 10 environmental factors that were significantly correlated with habitats in the valleys have a large ratio of rock cover floristic composition. 1 = Chamaecyparidion formosanae and 2 = and soil rockiness, and slopes are usually steep. Another Pasanio kawakamii-Machilion japonicae. Soil moisture was replaced habitat feature correlated with the first NMDS axis is by soil dryness because of a negative relationship between values and that summer atmospheric humidity is higher on the left soil moisture itself. b Eleven associations and contours of mean monthly July temperature (°C) in black, and July cloud frequency in side than the right in Fig. 2b. Forests dominated by gray (on a scale of 0–1). Elliptical polygons represent range of Chamaecyparis obtusa var. formosana (the first alliance, the standard deviations calculated from points of each association in left side in Fig. 2a) are more influenced by the summer the ordination space. Association codes are 1.01 = Tsugo formosanae- monsoon than the others because of the higher summer Chamaecyparidetum formosanae,1.02=Vaccinio lasiostemonis- Tsugetum formosanae,1.03=Schefflero taiwanianae-Chamaecy- humidity (see July cloud frequency in Fig. 3 and ESM paridetum formosensis,1.04=Elatostemato trilobulati-Tsugetum 3). In contrast, forests dominated by Chamaecyparis formosanae,1.05= Rhododendro formosani-Chamaecyparidetum for- formosensis are confined to shaded and stony middle mosanae,2.06=Adinandro lasiostylae-Chamaecyparidetum for- slopes and valleys, with less direct influence from the mosensis,2.07=Cyclobalanopsio stenophylloidis-Chamaecy- paridetum formosensis,2.08=Castanopsio carlesii-Chamaecyparide- summer monsoon. tum formosensis,2.09=Arachniodo rhomboideae-Chamaecyparide- Temperature, monsoon systems and related variables tum formosensis,2.10=Symploco wikstroemiifoliae-Machiletum are important in explaining habitat differences at the thunbergii and 2.11 = Pileo brevicornutae-Machiletum japonicae association level (Fig. 2a, b). These factors are strongly correlated to the second axis of NMDS, which is itself ratio of winter precipitation, indicating that the upper positively correlated with temperature and moisture part of the diagram has a stronger influence from the indicators. This axis is positively correlated with the winter monsoon than the lower part. Author's personal copy 776

Fig. 3 Box-plots of the nine most important and representative of each association code is explained in Fig. 2.Completeinformation environmental variables that were strongly correlated with ﬂoristic on all environmental factors is available in ESM 3. Annual mean soil composition in ordination space. Thick horizontal bars are medians; moisture is on a scale of 0–1; a large value indicates low soil moisture. boxes represent inter-quartile ranges; ranges of whiskers include 95 % Meanings of topography codes are: 1 = ridge, 2 = upper slope, of the values. X-axes represent diﬀerent associations and the meaning 3 = middle slope, 4 = lower slope, and 5 = valley

Syntaxonomical synopsis

We described eleven associations and two alliances. All Alliance 2. Pasanio kawakamii-Machilion japonicae belong to the same order: Ching-Feng Li et al. 2015 Order: Fagetalia hayatae Hukusima et al. 2005 Associations: 2.06 Adinandro lasiostylae-Chamaecyparidetum for- Alliance 1. Chamaecyparidion formosanae Tokio Suzuki mosensis Ching-Feng Li et al. 2015 1952 nom. mut. propos. 2.07 Cyclobalanopsio stenophylloidis-Chamaecyparide- Associations: tum formosensis Ching-Yu Liou ex Ching-Feng Li et al. 1.01 Tsugo formosanae-Chamaecyparidetum formosanae 2015 Ching-Yu Liou ex Ching-Feng Li et al. 2015 2.08 Castanopsio carlesii-Chamaecyparidetum formosen- 1.02 Vaccinio lasiostemonis-Tsugetum formosanae Ching- sis Ching-Feng Li et al. 2015 Feng Li et al. 2015 2.09 Arachniodo rhomboideae-Chamaecyparidetum for- 1.03 Scheﬄero taiwanianae-Chamaecyparidetum formosensis Ching-Feng Li et al. 2015 mosensis Ching-Long Yeh et Chien-Chun Liao ex 2.10 Symploco wikstroemiifoliae-Machiletum thunbergii Ching-Feng Li et al. 2015 Ching-Feng Li et al. 2015 1.04 Elatostemato trilobulati-Tsugetum formosanae To- 2.11 Pileo brevicornutae-Machiletum japonicae Ching- kio Suzuki 1952 Feng Li et al. 2015 1.05 Rhododendro formosani-Chamaecyparidetum formosanae Tokio Suzuki 1952 Author's personal copy 777

Table 1 Synoptic table of Chamaecyparis forest in Taiwan

1. 1. 1. 1. 1. 2. 2. 2. 2. 2. 2. Association code 01 02 03 04 05 06 07 08 09 10 11

Number of plots 25 49 20 17 37 38 28 35 26 27 33 Average number of all species per plot 43 28 40 59 51 69 63 63 55 69 54 Average number of tree species per plot 17 11 12 18 24 22 19 19 17 21 12 Average number of shrub species per plot 8 6 7 12 10 10 9 9 8 9 5 Average number of herb species per plot 11 7 14 19 11 23 23 21 19 24 25 Average number of liana species per plot 3 2 4 5 3 8 8 8 7 8 8 Average number of epiphyte species per plot 4 3 2 5 4 5 5 5 5 7 5

Common species of Chamaecyparis forest in Taiwan Neolitsea acuminatissima T 96 94 95 100 92 95 79 97 65 30 6 Plagiogyria euphlebia H443330716292 50 86 69 96 48 Araiostegia parvipinnata E 8049508246846883544452 Vittaria flexuosa E 643130825792 50 77 81 63 48 Hydrangea integrifolia L442090 47 32 95 86 51 73 52 55 Hymenophyllum polyanthos E36472594 30 58 32 51 31 70 27 Cyclobalanopsis morii T483165 – 16 11 54 86 58 – 9 Alliance 1 Chamaecyparidion formosanae Chamaecyparis obtusa var. formosana T3649 – 94 51 13 – – – 15 – Vaccinium japonicum var. lasiostemon S1647 5 59 27 – – – – – – Rhododendron formosanum T2445 – 53 705434–– Tsuga chinensis var. formosana T 80 94 80 41 38 50 14 6 – – – Eurya glaberrima T 68 55 60 82 24 32 7 11 19 4 – Schefflera taiwaniana T 60 63 45 41 11 11 11 31 19 – – Viburnum urceolatum S 76 22 – 41 49 26 7 6 8 4 – Damnacanthus angustifolius S 56 16 – 71 65 45 4 – 4 7 – Dendropanax dentiger T 80 55 25 82 86 21 21 46 12 56 3 Plagiogyria formosana H84827594 65 87 57 54 50 19 15 Smilax arisanensis L 76 35 10 82 68 84 11 20 15 4 12 Yushania niitakayamensis S 68 88 90 53 30 32 61 31 4 22 9 Symplocos morrisonicola T 96 45 45 65 51 82 18 46 23 7 12 Ilex tugitakayamensis T4018– 47 57 66 11 6 12 4 – Trochodendron aralioides T 76 33 60 41 51 74 50 34 23 26 9 Alliance 2 Pasanio kawakamii-Machilion japonicae Pasania kawakamii T– – – – 8 79 82 37 50 67 97 Arachniodes rhomboidea H4 6 3029227693 86 88 89 79 Diplazium kawakamii H– – 10185 3457 31 27 48 64 Pilea aquarum subsp. brevicornuta H– – 10– 3 2654 14 27 37 76 Machilus japonica T – – – 18 22 18 86 57 58 37 97 Polystichum parvipinnulum H8 4 55 – 3 68 82 40 38 7 55 Fatsia polycarpa T4 – – – 1466 43 17 23 41 30 Monachosorum henryi H16162035272482 94 77 81 64 Adinandra lasiostyla T122 15– 8 82 4 49 23 – 12 Damnacanthus indicus S 361810357097 86 100 81 81 48 Pellionia radicans H12–– 24145546204667 36 Chamaecyparis formosensis T 16 6 75 6 16 87 64 31 50 22 36 Eurya loquaiana S 64 8 25 29 73 97 68 91 88 78 58 Microsorium buergerianum H 12 4 20 35 22 53 82 69 65 74 70 Prunus phaeosticta T 4 2 5 – 496361465496 55 Stauntonia obovatifoliola L 40 8 20 41 38 97 82 71 46 81 18 Hedera rhombea var. formosana L124 55 – – 63 64 57 42 22 24 Litsea acuminata T 4 2 10 6 65 18 64 94 81 100 91 Lysionotus pauciflorus H5210353524663971425642 Litsea elongata var. mushaensis T 12 – 25 53 22 45 79 17 35 30 30 Eurya leptophylla S 442455123 74 64 43 42 15 21 Ardisia crenata S 40 16 5 65 46 66 50 63 54 33 36 Dryopteris formosana H722765654997 82 63 69 56 36 Elatostema trilobulatum H20205582 3 61 39 69 38 11 18 Acrophorus stipellatus H24162094 38 26 43 43 62 81 36 Association 1.01 Tsugo formosanae-Chamaecyparidetum formosanae Microtropis fokienensis S 88 43 40 29 24 58 21 37 23 15 3 Ainsliaea macroclinidioides H 32 12 5 12 5 5 – 3 4 – – Photinia niitakayamensis T 32 24 20 12 8 – 4 – 4 – – Association 1.02 Vaccinio lasiostemonis-Tsugetum formosanae Lyonia ovalifolia T2433 15 18 16 3 – 3 4 – – Association 1.03 Schefflero taiwanianae-Chamaecyparidetum formosensis Ilex bioritsensis T– 8 30–––––––– Ainsliaea latifolia subsp. henryi H162065 – 11 42 14 11 12 4 – Author's personal copy 778

Table 1 continued

1. 1. 1. 1. 1. 2. 2. 2. 2. 2. 2. Association code 01 02 03 04 05 06 07 08 09 10 11

Rubus formosensis S4 2 40–3–186–119 Pinus armandii var. mastersiana T1210356–13––––– Dryopteris lepidopoda H122 30–331134–– Osmanthus heterophyllus T16183065–43––– Acer morrisonense T166 45– 1613291423119 Lonicera acuminata L8 4 3529– 32112019–– Association 1.04 Elatostemato trilobulati-Tsugetum formosanae Ardisia japonica H–2–53143––4–– Viburnum sympodiale S164– 59143––––– Coptis quinquefolia H128 – 651634–44– Shortia rotundifolia H46–4714–––––– Rubus corchorifolius S861582225259–333 Maclura cochinchinensis L46–471134–4–– Eurya crenatifolia S 4 14 15 76 19 5 25 9 12 37 3 Barthea barthei S248– 76 575 4 171526– Nertera nigricarpa H 4 12 20 4735––4–6 Berberis kawakamii S–22041–37–44– Xiphopteris okuboi E8 33 5 471157–44– Ilex sugerokii var. brevipedunculata T 1218– 35113––––– Cleyera japonica var. taipinensis T322 – 65 51 504–411– Diplopterygium glaucum H42541111846–11– Tripterospermum lanceolatum L128 15353–49–4– Ligustrum liukiuense T42547 14 8 25 14 8 26 – Cleyera japonica T 3210– 59 54– 7 311526– Sarcopyramis napalensis var. delicata H124 15535 18211723433 Polypodium amoenum H281825593 8 144319269 Association 1.05 Rhododendro formosani-Chamaecyparidetum formosanae Schima superba T––––5711–9–11– Plagiogyria dunnii H122 – 2973 3 4 14 4 26 – Symplocos wikstroemiifolia S46–2451–––844 – Michelia compressa T–4––81 50 14 23 46 63 21 Ternstroemia gymnanthera T284 – 4781 50 11 3 42 44 9 Symplocos stellaris T 36 – 10 – 54 504 201973 Association 2.06 Adinandro lasiostylae-Chamaecyparidetum formosensis Rhamnus pilushanensis S–2–––34––––– Calanthe puberula H82–18574 14 9 8 11 – Sycopsis sinensis T82–18366 25 – 15 – – Symplocos migoi T 56 12 20 – 8 92 29 9 23 19 – Viburnum foetidum var. rectangulatum S 121035292284 14 20 8 7 – Ophiopogon japonicus H425––374––7– Trachelospermum formosanum L4–––8507––1915 Zanthoxylum scandens L––––350 11 11 8 – 15 Symplocos heishanensis T 4 6 – 12 30 5879237– Asarum macranthum H––56547 18 3 4 33 – Euonymus spraguei S 241020241161 14 9 15 – – Ligustrum sinense T––56–39 11 9 15 4 3 Ilex goshiensis T 5218–1881 89 14 29 27 37 9 Pyrola alboreticulata H 28 10 – 6 3 34–6––– Lycopodium serratum var. longipetiolatum H – 2 – 12 19 3241144– Rubus kawakamii S16230–3 50 18 23 19 – 24 Cyclobalanopsis stenophylloides T 361030181468 646 271142 Malus doumeri T164 5 – 1134–62373 Camellia tenuifolia S 2065 1814424––37 9 Rubus pectinellus H4225241447 25 9 12 22 9 Goodyera velutina H 24 6 10 12 19 39 11 9 8 15 – Daphniphyllum himalaense subsp. macropodum T 32 8 10 35 24 55 39 – 12 33 27 Athyrium arisanense H841561445 32 34 23 19 – Association 2.07 Cyclobalanopsio stenophylloidis-Chamaecyparidetum formosensis Viburnum arboricolum E–––––55468418 Ligustrum pricei T––––––32––153 Pourthiaea beauverdiana var. notabilis T4–10–5–57 31 19 15 21 Ophiorrhiza japonica H––––321469 121530 Litsea morrisonensis T– 4 35–51646 14 4 7 9 Pyrrosia sheareri E402 256 5 71 71 6 31 – 36 Author's personal copy 779

Table 1 continued

1. 1. 1. 1. 1. 2. 2. 2. 2. 2. 2. Association code 01 02 03 04 05 06 07 08 09 10 11

Schisandra arisanensis L––10–3339681130 Viburnum taitoense S––––51332–4–24 Urtica thunbergiana H––10–––32 3 12 – 27 Ophiopogon intermedius H126 5 291121439––3 Carex brunnea H– – 35––2136 6 12 – 27 Pasania hancei var. ternaticupula T 321435123– 39–4116 Association 2.08 Castanopsio carlesii-Chamaecyparidetum formosensis Gordonia axillaris T82––11–469 12 7 3 Polystichum prionolepis H–––––––34 4 4 – Hydrangea chinensis S4–10––3–57 27 4 3 Lithocarpus amygdalifolius T–––––––37 12 – 6 Castanopsis cuspidata var. carlesii T162––49 45 – 77 19 19 – Callicarpa randaiensis S4––12551860 31 11 9 Lemmaphyllum diversum E–25–1418–43 15 – – Eurya chinensis S4–15––––34 12 – 6 Vaccinium emarginatum E1616––113–54 27 37 12 Eurya strigillosa T – 10 10 24 19 – 4 51 38 7 6 Neolitsea aciculata var. variabillima T 4 – – 1830–1451 – 26 21 Embelia laeta var. papilligera L – 2 – 1214–1146 15 30 15 Plagiogyria stenoptera H4 8 5 241681851 15 30 6 Peperomia reﬂexa H445128–743 31 11 42 Adinandra formosana T 4 18– 2919–– 34 4 7 3 Asplenium normale H4 4 – 2943242157 38 52 18 Association 2.10 Symploco wikstroemiifoliae-Machiletum thunbergii Selaginella doederleinii H–––6531191581 15 Pasania konishii T––––––46–37 – Maesa japonica S––––14814–859 6 Davallia mariesii H––––113113852 6 Osmanthus matsumuranus T––––14–14172763 6 Pasania harlandii T––––11–43441 – Selaginella involvens H12––24811113452 – Pileostegia viburnoides L 4 2 10611– 25203174 36 Asplenium antiquum E––––14311–1959 36 Pyrrosia lingua E–6–6411814231974 21 Itea parviﬂora T––––35111862774 48 Cinnamomum subavenium T––––41–43–44 – Machilus thunbergii T204–670 – 25 80 42 93 9 Pyrenaria shinkoensis T–––128–4––30 – Smilax lanceifolia L 4 105 1235–3274 38 85 45 Cyclobalanopsis longinux T 4 8 5 1243135064278 39 Ficus sarmentosa var. nipponica L4––––2139172352 33 Association 2.11 Pileo brevicornutae-Machiletum japonicae Ficus pumila var. awkeotsang L––––––––4–30 Machilus zuihoensis var. mushaensis T–––––314–12752 Piper kadsura L––––3–18684473 Begonia formosana H–––––––––2239 Oreocnide pedunculata T–––––––341142 Diplazium amamianum H4–––––25 3 15 7 52 Tetrastigma umbellatum L – – 10 – 3 – 25 43 50 19 79 Selaginella delicatula H––––8–73–1942 Lithocarpus lepidocarpus T––––––731 8 – 42 Elatostema parvum H4–5––314315–39 Oplismenus hirtellus H––––––438430 Symplocos modesta S––5–3–42919748 Vandenboschia auriculata E––––352123274145 Hemiboea bicornuta H–––63341781930 Polypodium formosanum H44565111417232636

Values in cells are percentage constancy. Diagnostic species of each syntaxon are sorted by ﬁdelity (U). Underlined numbers indicate U > 0.15, and only those with constancy >30 % are listed. The complete synoptic table is available in ESM 4 and ESM 5. The association 2.09 Arachniodo rhomboideae-Chamaecyparidetum formosensis does not have its own diagnostic species in this table, owing to its low species richness and broad geographic distribution. Letters following the species names are T tree, S shrub, H herb, L liana and E epiphyte Author's personal copy 780

Description of associations and alliances

Each syntaxon (association or alliance) is characterized by its name from the literature (ESM 1), ﬂoristic composition (Table 1;ESM4andESM5),vertical structure of vegetation (Table 2), habitat requirements (Fig. 3;Table2; ESM 3), geographic distribution (ESM 6), and information on diﬀerences from similar

EN, NE, NW EN, ES vegetation or habitat types. Fidelity of species to the syntaxa (based on U-value) was used as a criterion for selection of the diagnostic species. The U threshold was set to 0.1 for the deﬁnition of diagnostic species, and the list was sorted in alphabetical order within

CW, EN, ES, NE, NW, SE, SW each life form. Diagnostic species for alliances are listed in the main text, but the U threshold used was

southwestern set to 0.15. ESM 7 lists the diagnostic species of alliances and associations with U 0.1. Figure 4 summa- SW

CW, ES, SE, SW rizes the physiognomic structure of trees in each association. In the nomenclature type releve´ s, species were sorted by their percentage cover within each life form. Header data of each nomenclature type releve´

southeastern, are in Table 3 and species data in Table 4 in Appen- SE NE, NW, SE dix. The Cocktail Determination Key for the 11 evergreen broad-leaved forest. Ecoregion codes for following ecoregions:

3 associations which can be directly imported into the CoDeK program (Appendices S2 and S3 in Li et al. 2013), is presented in ESM 8. For future application of supervised analysis by other researchers, nine ran- northwestern, domly selected plots of each association, together with the type releve´ s, are appended in ESM 9 and ESM 10 NW forest associations in Taiwan (for the ﬂoristic composition and header data, respectively). northeastern, Chamaecyparis NE NE, NW CW, NW CW, ES, NW CW, EN, ES, CW, EN, ES, NE, NW, SE, SW south part of eastern, ES mixed forest of coniferous and evergreen broad-leaved species, and 2 CW, EN, ES, NE, NW, SE, SW north part of eastern, 1800–2600 1800–2700 2150–2650 1600–2300 1300–2400 1800–2300 1400–2500 1800–2300 1500–2500 1100–1700 1200–2100 3–8 3–10 3–6 3–8 3–8 3–8 3–8 3–8 3–8 3–6 – 10–15 – – – 10–15 10–15 – – 10–15 10–15 – 105–145NE, 95–150 NW, SE, SW 80–135 120–145 110–160 100–135 95–160 110–140 105–150 140–170 105–165 coniferous forest, EN 1 month) Æ C ° Physiognomy, geographic distribution and vertical structure of the

central-western, Fig. 4 Ratio of cover composed by coniferous, deciduous broad- leaved, and evergreen broad-leaved trees in each association. small-tree layer (m) altitude (m) index ( height (m) Association codePhysiognomy 1.01Range of 1, 2 1.02 1, 2 1.03 1, 2 1.04 1.05 1 2.06 2 2.07 2 2.08 2.09 2, 3 2.10 2, 3 2, 2.11 3 2, 3 2, 3 Table 2 Range of warmth Physiognomy codes: Height of the Canopy cover (%)Canopy height (m)Sub-canopy 60–80 15–25 60–90 10–20 40–80 10–20 70–90 15–20 70–90 15–25 50–70 25–35 60–80 15–20 60–80 15–25 70–90 15–30 70–80 15–20 60–80 15–20 CW Ecoregions CW, EN, ES, Meanings of association codes are the same as in Fig. 2 Author's personal copy 781

Alliance 1. Chamaecyparidion formosanae Tokio Suzuki 1952 nom. mut. propos. English name: Montane mixed coniferous forest (Su Tien-Chai Chen 1984); Chamaecyparis montane mixed cloud forest (Li et al. 2013). Original name: Chamaecyparidion taiwanensis Tokio Suzuki 1952 (Chamaecyparis taiwanensis = C. obtusa var. formosana). Tze-Ying Chen Nomenclature type association: Rhododendro formosani-Chamaecyparidetum formosanae Tokio Suzuki 1952 (lectotypus hoc loco designatus). Range of altitude and warmth index: 1400–2700 m;

Chang-Fu Hsieh 90–155 °CÆmonth. Diagnostic species: Trees: Chamaecyparis obtusa var. formosana, Cy- clobalanopsis morii, C. sessilifolia, Dendropanax dentiger,

Ho-Yih Liu Eurya glaberrima, Ilex goshiensis, I. tugitakayamensis, Illicium anisatum, Neolitsea acuminatissima, Rhododen- dron formosanum, R. leptosanthum, Scheﬄera taiwaniana, Symplocos morrisonicola, Trochodendron aralioides, Tsuga chinensis var. formosana.

Tien-Chai Chen Shrubs: Barthea barthei, Damnacanthus angustifolius, Microtropis fokienensis, Skimmia arisanensis, Viburnum urceolatum, Yushania niitakayamensis. Herbs: Plagiogyria euphlebia, P. formosana. Lianas: Hydrangea integrifolia, Smilax arisanensis. Chang-Fu Hsieh Epiphytes: Araiostegia parvipinnata, Hymenophyllum polyanthos, Vittaria ﬂexuosa. Chamaecyparidion formosanae is a mixed or coniferous forest. The canopy cover is about 60–80 % and average canopy height 20 m. Three layers are com-

Tze-Ying Chen monly developed. The canopy is mostly composed of valley

5 coniferous trees, where the typically dominant trees are Chamaecyparis formosensis, C. obtusa var. formosana, and Tsuga chinensis var. formosana. Evergreen broad-

Tze-Ying Chen leaved trees such as Cyclobalanopsis morii and Tro- chodendron aralioides are usually mixed with conifers in forest in Taiwan the canopy or sometimes form a distinct sub-canopy

lower slope, and layer. Illicium anisatum, Neolitsea acuminatissima, 4 Rhododendron formosanum and R. leptosanthum are often dominant in the small-tree layer. The shrub layer is Ching-Long Yeh dominated by Myrsine stolonifera and Yushania Chamaecyparis niitakayamensis. Plagiogyria formosana is dominant in sof middle slope, ´ the herb layer. The representative species group of this 3 alliance is composed of Chamaecyparis obtusa var. for-

Tien-Chai Chen mosana, Dendropanax dentiger, Eurya glaberrima, Illi- cium anisatum, Neolitsea acuminatissima, Rhododendron formosanum, Tsuga chinensis var. formosana, Tro-

upper slope, chodendron aralioides, Vaccinium japonicum var. la- 2 siostemon,andYushania niitakayamensis (ESM 8; Chen Appendix S3 in Li et al. 2013). This indicates acid soil on ridge,

1 slopes to ridges, with less topographic shading in the ) 400 400 500 400 400 400 400 400 400 400 400 2 montane cloud zone of Taiwan. The presence of Eri- caceae in both species number and cover is high while 114232422623 )1018402939224029256 ) 121.2873 121.5316 120.7486 121.403 121.3757 121.2888 121.5303 120.8196 120.8471 121.4493 121.3128 ° °

) 24.53854 24.13714 22.61514 24.569 24.54146that 24.53602 of 24.13749 Lauraceae 22.86558 23.47336 is 24.62187 low. 24.02356 The proportion of undecom- ° ) 277 261 306 337 176 45 269 68 47 336 113 Headers of nomenclature type releve °

ID 02-0609 30-0097 20-0295 02-2077 02-2040 02-0614 30-0096posed 20-0123 material 11-0096 02-2069 in the 29-0166 soil is large. A thick bryophyte ´ layer covers trunks, branches, and the ground. Lightning occasionally strikes the tallest emergent coniferous trees TopographyRockiness (%)Rock cover (%)Sampling year 1Topography 5 codes: 0 2005 1 0 0 2004 2005 5 90 0 2005 0 1 0 2005 35 3 2005 0 1 2004 3 0 2004 30 2 5 2003 10 4 2005 50 5 2007 5 25 0 5 40 60 10 2 Latitude ( Longitude ( Inclination ( Canopy cover (%)Tree height (m)Altitude (m) 50Aspect ( 35 2150 85 20 1985 90 20 2470 70 1950 10 90 1895 15 2130 60 35 1855 75 20 2137 80 2325 15 60 1086 30 1819 80 20 60 15 Author Tze-Ying Table 3 Sampling area (m Association codeReleve 1.01 1.02 1.03 1.04 1.05 2.06 2.07 2.08 2.09 2.10 2.11 Author's personal copy 782 in summer, but the resulting fire affects a small area 1.03 Schefflero taiwanianae-Chamaecyparidetum for- because of the high humidity. Therefore, soil contains a mosensis Ching-Long Yeh et Chien-Chun Liao ex Ching- distinct layer of charcoal in places or patches, with post- Feng Li et al. 2015 ass. nov. hoc loco fire species such as Lyonia ovalifolia, Photinia Nomenclature type releve´ : 20-0295 (holotypus hoc niitakayamensis, Pieris taiwanensis, Pinus taiwanensis, loco designatus). and Rhododendron rubropilosum. This alliance is dis- Coniferous trees such as Chamaecyparis formosensis, tributed across all Taiwan except the tropical region, Pinus armandii var. mastersiana and Tsuga chinensis var. and its habitats are often influenced by the summer formosana mixed with evergreen broad-leaved trees, monsoon. such as Cyclobalanopsis morii, C. stenophylloides and Trochodendron aralioides, are dominant in the canopy. 1.01 Tsugo formosanae-Chamaecyparidetum formosanae Picea morrisonicola is sometimes dominant, whereas Ching-Yu Liou ex Ching-Feng Li et al. 2015 ass. nova hoc deciduous trees such as Acer kawakamii and A. mor- loco risonense are frequent in the canopy. Small trees such as Nomenclature type releve´ : 02-0609 (holotypus hoc Eurya glaberrima, Neolitsea acuminatissima and Sym- loco designatus). plocos morrisonicola are dominant. The shrub layer is Coniferous trees such as Chamaecyparis obtusa var. dominated by Yushania niitakayamensis and Plagiogyria formosana and Tsuga chinensis var. formosana dominate formosana is a dominant herb. This association, the the canopy. Because Chamaecyparis obtusa var. for- coldest and driest among all Chamaecyparis forests in mosana is only found north of the Tropic of Cancer, its Taiwan, occurs at high altitudes on slopes and ridges dominance is replaced by Cyclobalanopsis morii in with large rock outcrops across all Taiwan. Its habitats southern Taiwan. Other conifers, such as Chamaecyparis are similar to those of Rhododendro pseudochrysanthum- formosensis, Pinus armandii var. mastersiana, and Tai- Tsugetum formosanae (Lin et al. 2012), which is an up- wania cryptomerioides are sometimes dominant in the per-montane coniferous forest that grows above the canopy. The sub-canopy is dominated by evergreen montane cloud zone. The species group of montane broad-leaved trees such as Cyclobalanopsis morii, rocky habitats, including Acer morrisonense, Hydro- C. sessilifolia and Trochodendron aralioides, whereas cotyle setulosa, Peranema cyatheoides, Rubia lanceolata Neolitsea acuminatissima, Rhododendron formosanum, and Rubus formosensis, is frequent in this association, R. leptosanthum, Schefflera taiwaniana and Symplocos and absent in the high-montane coniferous forest. morrisonicola are dominant in the small tree layer. Moreover, Rhododendro pseudochrysanthum-Tsugetum Dwarf bamboo, Yushania niitakayamensis, is dominant formosanae contains some subalpine species such as in the shrub layer, and Plagiogyria formosana in the herb Abies kawakamii, Circaea alpina subsp. imaicola, Galium layer. This association occurs on wide ridges or upper echinocarpum, Ribes formosanum, Rubus rolfei and Se- slopes with well-developed soil, island-wide. These wide dum actinocarpum, which are not found in Schefflero ridges have relatively stable landform within the com- taiwanianae-Chamaecyparidetum formosensis. plex mountain terrain. Such habitats were heavily reforested as agricultural land for planting tea or tem- 1.04 Elatostemato trilobulati-Tsugetum formosanae To- perate crops, such as apples, peaches, or cabbage. kio Suzuki 1952 nom. mut. propos. Original name: Pellionieto-Tsugetum sinensis Tokio 1.02 Vaccinio lasiostemonis-Tsugetum formosanae Suzuki 1952 (Pellionia trilobulatum = Elatostema Ching-Feng Li et al. 2015 ass. nov. hoc loco trilobulatum, Tsuga sinensis = Tsuga chinensis var. for- Nomenclature type releve´ : 30-0097 (holotypus hoc mosana). loco designatus). Nomenclature type releve´ : 02-2077 (neotypus hoc Coniferous trees such as Chamaecyparis obtusa var. loco designatus). formosana and Tsuga chinensis var. formosana are Chamaecyparis obtusa var. formosana and Tsuga dominant in the canopy. The dominance of Chamaecy- chinensis var. formosana are dominant in the tree layer. paris obtusa var. formosana is replaced by Cycloba- Illicium anisatum, Neolitsea acuminatissima, Rhododen- lanopsis morii in southern Taiwan. The sub-canopy layer dron formosanum and R. leptosanthum are dominant is absent or composed of few individuals of Elaeocarpus small trees. Barthea barthei, Myrsine stolonifera and japonicus or some species of Fagaceae. Illicium anisatum, Yushania niitakayamensis are dominant shrubs, and Lyonia ovalifolia, Neolitsea acuminatissima and Rhodo- Plagiogyria formosana and P. euphlebia are dominant dendron formosanum are dominant in the small tree herbs. The habitats of this association are on slopes that layer. Yushania niitakayamensis is dominant in the shrub are less shaded by the surrounding topography, with layer, and Plagiogyria formosana in the herb layer. This shallow soils. The forest floor is bog-like, covered by a association occurs on narrow ridges with moderately thick layer of Sphagnum spp. Coptis quinquefolia, Shor- developed soil across all Taiwan. Several species of tia rotundifolia and Viburnum sympodiale indicate this Ericaceae are frequent within this vegetation, indicating cool and very humid habitat. Monachosorum henryi, acid soil. Shrub and herb layers in this association are Plagiogyria euphlebia, P. stenoptera and Sarcopyramis sparse. Vegetation on the narrow ridges is usually spe- napalensis var. bodinieri form a species group repre- cies-poor compared to other Chamaecyparis forests on senting a heavy accumulation of organic material on the wide ridges. Author's personal copy 783 forest floor. All trunks are covered by a thick layer of Neolitsea acuminatissima, Pasania kawakamii, Prunus bryophytes. It is often found that tree seedlings regen- phaeosticta, Symplocos arisanensis, S. migoi. erate on the standing stumps and trunks still alive. There Shrubs: Ardisia crenata, Damnacanthus indicus, Eurya are few Fagaceae species in the canopy or sub-canopy. It leptophylla, E. loquaiana. has been suggested that the shallow groundwater table Herbs: Acrophorus stipellatus, Arachniodes rhom- and humidity prevent Fagaceae growth in this associa- boidea, Arthromeris lehmannii, Diplazium kawakamii, tion (Chou et al. 2000). Geographically, this association Dryopteris formosana, Elatostema trilobulatum, Ly- is confined to northeastern Taiwan, especially around sionotus pauciflorus, Monachosorum henryi, Pellionia Cuei-Fong, Song-Luo and Yuan-Yang lakes. These radicans, Pilea aquarum subsp. brevicornuta, Plagiogyria locations and the substantial heterogeneity of moisture euphlebia, P. formosanum, Polystichum hancockii, conditions within a stand makes this association the P. parvipinnulum. most humid and species-rich community in Chamaecy- Lianas: Hedera rhombea var. formosana, Hydrangea paridion formosanae. integrifolia, Schizophragma integrifolium var. fauriei, Stauntonia obovatifoliola. 1.05 Rhododendro formosani-Chamaecyparidetum for- Epiphytes: Araiostegia parvipinnata, Asplenium wil- mosanae Tokio Suzuki 1952 nom. mut. propos. fordii, Pyrrosia sheareri, Vittaria flexuosa. Original name: Rhodoreto-Chamaecyparidetum tai- Pasanio kawakamii-Machilion japonicae is a mixed or wanensis Tokio Suzuki 1952 (C. taiwanensis = C. obtusa evergreen broad-leaved forest in physiognomy. Canopy var. formosana). cover is about 70–90 % and average canopy height is Nomenclature type releve´ : 02-2040 (neotypus hoc 20 m. Most of the associations of this alliance, except loco designatus). Pileo brevicornutae-Machiletum japonicae, which grows Chamaecyparis obtusa var. formosana, C. formosensis over limestone, have a vertical structure of five layers. and Tsuga chinensis var. formosana are dominant in the The canopy is mostly composed of evergreen broad- canopy. Castanopsis cuspidata var. carlesii, Machilus leaved trees, dominated by Castanopsis cuspidata var. thunbergii and Elaeocarpus japonicus are dominant in the carlesii, Cyclobalanopsis morii, Machilus japonica, sub-canopy. Species of Theaceae such as Pyrenaria M. thunbergii and Pasania kawakamii. Chamaecyparis shinkoensis, Schima superba and Ternstroemia gymnan- formosensis is usually an emergent tree above the canopy. thera are frequent accompanying trees. Neolitsea The sub-canopy is dominated by Litsea acuminata, while acuminatissima, Rhododendron formosanum and R. lep- Neolitsea acuminatissima and Eurya loquaiana are dom- tosanthum are dominant small trees. Myrsine stolonifera inant in the small-tree layer. Yushania niitakayamensis and Yushania niitakayamensis are dominant shrubs, and dominates in the shrub layer and Dryopteris formosana, Dryopteris formosana, Plagiogyria dunnii, P. euphlebia Monachosorum henryi and Plagiogyria formosana in the and P. formosana are dominant herbs. This association herb layer. A species group composed of Castanopsis occupies the south-facing slopes in the western part of cuspidata var. carlesii, Chamaecyparis formosensis, Cy- central and northern Taiwan. In Chamaecypardion for- clobalanopsis morii, Eurya strigillosa, Litsea acuminata, mosanae, the habitat of this association is rocky and Machilus japonica, M. thunbergii and Neolitsea acumi- warm. The proportion of coniferous trees is small natissima is well represented on slopes or in valleys with (Fig. 4). Lauraceae and Fagaceae have substantial cover stony soil in the montane cloud zone of Taiwan (ESM 8; and number of species. Appendix S3 in Li et al. 2013). Seedlings of Chamaecy- Alliance 2. Pasanio kawakamii-Machilion japonicae paris formosensis are usually absent in the understory of Ching-Feng Li et al. 2015 all. nov. hoc loco this alliance; they are mostly found on fresh landslides. English name: Montane evergreen broad-leaved for- Habitats of the associations under this alliance described est (Su 1984); Quercus montane evergreen broad-leaved are assumed to be landslides hundreds to thousands of cloud forest (Li et al. 2013). years ago (Chen 2001). Compared with Chamaecypari- Nomenclature type association: Cyclobalanopsio dion formosanae, this alliance has lower cover of bryo- stenophylloidis-Chamaecyparidetum formosensis Ching- phytes and of Ericaceae, greater cover and species Yu Liou ex Ching-Feng Li et al. 2015 (holotypus hoc number of Lauraceae with greater diversity of herbs, loco designatus). lianas and epiphytes. This alliance is found all over Range of altitude and warmth index: 1000–2300 m; Taiwan except in tropical area defined by Li et al. (2013). 100–165 °CÆmonth. 2.06 Adinandro lasiostylae-Chamaecyparidetum for- Diagnostic species: mosensis Ching-Feng Li et al. 2015 ass. nov. hoc loco Trees: Adinandra lasiostyla, Chamaecyparis formosensis, Cyclobalanopsis morii, C. stenophylloides, Nomenclature type releve´ : 02-0614 (holotypus hoc Fatsia polycarpa, Litsea acuminata, Machilus japonica, loco designatus). Author's personal copy 784

Chamaecyparis formosensis, Pinus armandii var. chinensis and Yushania niitakayamensis are dominant mastersiana and Tsuga chinensis var. formosana domi- shrubs, and Asplenium normale and Monachosorum nate the canopy. In places, Taiwania cryptomerioides is henryi are dominant herbs. A species group composed of an emergent tree above the canopy. Cyclobalanopsis Cyclobalanopsis morii, Eurya strigillosa, Lithocarpus sessilifolia, Sycopsis sinensis and Trochodendron ara- amygdalifolius, L. lepidocarpus, Machilus japonica and lioides are dominant in the sub-canopy. Eurya loquaiana Symplocos sonoharae represents warm habitats without and Neolitsea acuminatissima are dominant small trees. seasonal temperature fluctuation in southern Taiwan. Yushania niitakayamensis is dominant in the shrub layer, Blastus cochinchinensis, Callicarpa randaiensis, Dam- and Arachniodes rhomboidea, Dryopteris formosana, nacanthus indicus, Eurya loquaiana and Hydrangea chi- Pellionia radicans and Plagiogyria formosana are domi- nensis form a representative species group of this nant herbs. Another representative species group of this association in the shrub layer. This association occurs association is composed of lianas, including Ficus sar- primarily on slopes south of the Tropic of Cancer. mentosa var. nipponica, Hedera rhombea var. formosana, Hydrangea integrifolia, Lonicera acuminata, Smilax 2.09 Arachniodo rhomboideae-Chamaecyparidetum for- arisanensis, Schizophragma integrifolium var. fauriei and mosensis Ching-Feng Li et al. 2015 ass. nov. hoc loco Stauntonia obovatifoliola. This liana species group is also Nomenclature type releve´ : 11-0096 (holotypus hoc frequent in the Picea forest which grows on nutrient-rich loco designatus). soil in the montane cloud zone of the island. The soil of Cyclobalanopsis morii, C. longinux, Machilus japonica this habitat is well-developed with some large stones of and M. thunbergii are dominant in the canopy. argillite, which is nutrient-rich after weathering. In Chamaecyparis formosensis and sometimes Taiwania western Taiwan, this association occupies north-facing cryptomerioides are emergent above the canopy. Litsea slopes, while in the eastern part it is on south-facing acuminata and Neolitsea acuminatissima are dominant slopes. Such habitats are less influenced by both summer small trees. Hydrangea chinensis is dominant in the and winter monsoons, indicating less frequent dense fog. shrub layer. Plagiogyria formosana and Strobilanthes flexicaulis are dominant in the herb layer. This associa- 2.07 Cyclobalanopsio stenophylloidis-Chamaecyparide- tion is widespread across Taiwan on steep slopes that are tum formosensis Ching-Yu Liou ex Ching-Feng Li et al. rocky and shaded by surrounding topography. 2015 ass. nov. hoc loco Nomenclature type releve´ : 30-0096 (holotypus hoc 2.10 Symploco wikstroemiifoliae-Machiletum thunbergii loco designatus). Ching-Feng Li et al. 2015 ass. nov. hoc loco Cyclobalanopsis morii, Machilus japonica and Nomenclature type releve´ : 02-2069 (holotypus hoc M. thunbergii are dominant in the canopy and Chamae- loco designatus). cyparis formosensis is an emergent tree above the canopy. Chamaecyparis formosensis, Cyclobalanopsis longi- Litsea acuminata, Prunus phaeosticta and Symplocos nux, Machilus thunbergii and M. japonica are dominant formosana are dominant small trees. Yushania niitakaya- in the canopy, and Cyclobalanopsis gilva and Litsea mensis is dominated in the shrub layer, and Monachoso- acuminata are dominant in the sub-canopy. Prunus rum henryi, Pellionia radicans and Plagiogyria formosana phaeosticta is a dominant small tree and Yushania are dominant herbs. Deciduous broad-leaved trees such as niitakayamensis dominates in the shrub layer. Diplazium Acer palmatum var. pubescens, A. kawakamii, A. morriso- dilatatum, Dryopteris formosana, Monachosorum henryi nensis, Carpinus kawakamii and C. rankanensis are com- and Selaginella doederleinii are dominant herbs. This mon (Fig. 4), indicating relatively dry and rocky habitats. association occurs mostly on north-facing slopes and This association occurs mainly in eastern Taiwan, but it is ridges of northern Taiwan. Such habitats are charac- also found in some locations in the west, near valleys terized by very high atmospheric humidity and a large adjacent to ridges of the backbone mountains. These proportion of winter precipitation. This association oc- habitats are less influenced by the summer monsoon, but curs at lower altitudes than the other Chamaecyparis humidity is high in winter. forest types. There are more lowland species such as Cinnamomum subavenium, Cyclobalanopsis gilva, Itea 2.08 Castanopsio carlesii-Chamaecyparidetum formosen- parviflora, Pasania harlandii and P. konishii in the tree sis Ching-Feng Li et al. 2015 ass. nov. hoc loco layer, and Alpinia japonica and Begonia formosana in the Nomenclature type releve´ : 20-0123 (holotypus hoc herb layer. loco designatus). Castanopsis cuspidata var. carlesii, Cyclobalanopsis 2.11 Pileo brevicornutae-Machiletum japonicae Ching- morii, Lithocarpus amygdalifolius, Machilus japonica and Feng Li et al. 2015 ass. nov. hoc loco M. thunbergii are dominant in the canopy, and Nomenclature type releve´ : 29-0166 (holotypus hoc Chamaecyparis formosensis is emergent above the ca- loco designatus). nopy. Eurya loquaiana, Litsea acuminata and Neolitsea Chamaecyparis formosensis, Litsea acuminata, Ma- acuminatissima are dominant small trees. Hydrangea chilus japonica, M. zuihoensis var. mushaensis and Author's personal copy 785

Pasania kawakamii are dominant in the canopy. There Chamaecyparis spp. are dominant in the Chamaecy- are no obvious sub-canopy or small-tree layers. Sym- paris montane mixed cloud forest or occuring with plocos modesta is dominant in the shrub layer. Ara- scattered individuals in both Quercus montane evergreen chniodes rhomboidea, Diplazium amamianum, broad-leaved cloud and Fagus montane deciduous D. kawakamii, Elatostema lineolatum var. majus and broad-leaved cloud forests. These three main forest types Monachosorum henryi are dominant herbs. Elatostema are treated as three alliances. Two of these, Chamaecy- and Pilea species are common in such rocky habitats. paridion formosanae and Pasanio kawakamii-Machilion The ratio of deciduous broad-leaved trees is relatively japonicae, are described in the present study, and the large, indicating less soil moisture. Araiostegia parvi- third, Fagion hayatae, was described by Hukusima et al. pinnata, Pyrrosia sheareri, Vandenboschia auriculata and (2005). Frequent species of each alliance were listed in Vittaria flexuosa are frequent epiphytes. Arachniodes Table S4 in ESM 11 and Appendix S6 in Li et al. (2013). rhomboidea, Carex brunnea, Coniogramme intermedia, These showed differences of the three alliances, along Diplazium kawakamii, Ellisiophyllum pinnatum, Mi- with their close relationship as demonstrated by large crosorium buergerianum, Polypodium formosanum and numbers of shared species between them. Thelypteris esquirolii make up representative species What is the higher syntaxon for the Chamaecyparis groups for this association. This forest mainly grows in forest in Taiwan? What are other montane cloud habi- eastern Taiwan, often on limestone with shallow soil tats in eastern Asia and what is the relationship between under strong topographic shading. their forests and the Chamaecyparis forest in Taiwan? In Japan, the habitat of C. obtusa forest is affected by frequent fog (Miyawaki 1985). Forests dominated by Discussion C. obtusa in Japan were classified into two alliances (Fujiwara 1996). These are Chamaecyparidion obtusae In Taiwan, the mid-altitude range 1500–2500 m a.s.l. is Yamanaka 1962 included in the order Pinetalia penta- called the temperate or warm temperate zone (Song phyllae Tokio Suzuki 1966, and Tsugion sieboldii Tokio 1999), Quercus zone (Su 1984), or montane cloud zone Suzuki 1953 belonging to the order Saso-Fagetalia cre- (Su 1984; Chen 2001; Li et al. 2013). This zone is char- natae Tokio Suzuki 1966. Another montane cloud acterized by a mosaic of coniferous, deciduous broad- habitat is in Yakushima. Forests there are dominated by leaved, evergreen broad-leaved and mixed forests. Li Cryptomeria japonica (Miyawaki 1980), and their alli- et al. (2013) defined seven main forest types in this mid- ances are Tsugion sieboldii in the order Saso-Fagetalia altitudinal range: (1) Chamaecyparis montane mixed crenatae and Quercion acto-myrsinaefoliae Fujiwara cloud forest; (2) Fagus montane deciduous broad-leaved 1980 in the order Illicio-Quercetalia acutae Fujiwara cloud forest; (3) Quercus montane evergreen broad- 1980 (Fujiwara 1996). Montane cloud forest is also leaved cloud forest; (4) Pasania-Elaeocarpus montane known in mainland China as ‘‘mossy dwarf forest’’ (Xu evergreen broad-leaved cloud forest; (5) Zelkova-Quer- and Wang 2010), but it is mainly evergreen broad-leaved cus rock-outcrop forest; (6) Pinus successional wood- forest and details of its floristic composition remain land; (7) Alnus successional woodland. The above forest unknown. There is no enough published information for types 1, 2 and 3 are defined as SMCF, and type 4 as comparing SMCFs between Taiwan and mainland tropical montane cloud forest. Types 5, 6 and 7 are China. azonal forests, also found outside the montane cloud Comparison of frequent species in Taiwanese SMCFs zone. Another type of evergreen broad-leaved forests, and Japanese Chamaecyparis obtusa and Cryptomeria which is associated with fog formation but does not japonica forests (Japanese montane cloud forests) indi- grow in the mid-altitude range, is Pyrenaria-Machilus cates that Taiwanese SMCFs should be classified into a winter monsoon forest (Li et al. 2013). It is found mostly new order, which does not occur in Japan. Few species in northeastern Taiwan, at altitudes up to 1200 m a.s.l. are shared between Taiwanese and Japanese montane on windward slopes directly exposed to the winter cloud forests (Table S4 in ESM 11). SMCFs in Taiwan monsoon, with cool, foggy and windy winters. Com- have greater cover and species number of ferns in herb pared to forests in the montane cloud zone, Pyrenaria- and epiphyte layers than the Japanese forests as well as Machilus winter monsoon forest is short (usually less much greater dominance and species number of Lau- than 10-m tall), with a dense canopy composed of raceae and Fagaceae in the tree layer. Considering the numerous small individuals with DBH rarely exceeding geographic distribution of species frequent in Taiwanese 20 cm, even for old individuals. SMCFs (Table S5 in ESM 11), 25 % of these are en- Author's personal copy 786 demic, 40 % occur in both Taiwan and mainland China, mental factors or differences in biotic interactions 26 % have ranges across mainland China, Japan and associated with those factors, which vary with ecoregion Taiwan, and only 9 % are found in both Taiwan and and can influence species composition. Japan. Hukusima et al. (2005; 2013) classified the Fagus Empirical observation indicates that the distribution forest in Taiwan into the order Fagetalia hayatae. The of the Chamaecyparis forest in Taiwan is positively present study supports the view that the two alliances of correlated with fog events. Fog events are correlated Chamaecyparis forest in Taiwan, Chamaecyparidion with the microclimate and the soil conditions influenced formosanae and Pasanio kawakamii-Machilion japonicae, by the hydrological process (Bruijnzeel et al. 2010). The should be included in this order. However, further results of the present study suggest that the differentia- comparisons are required to determine whether this or- tion of vegetation types is related to fog seasonality. der should be included in the class, Camellietea japonicae Light conditions in the understory, moisture, and the Miyawaki et Ohba 1963, which represents the laurel degree of decomposition of litter affect establishment of forest in Japan, or Litseo elongatae-Fagetea sp. div. seedlings (Forget et al. 2004). Activity of frugivores and Hukusima et al. 2013, which corresponds with the sub- visibility of seeds determine which species can success- tropical montane Fagus forests in both mainland China fully regenerate (Schupp et al. 2010). Furthermore, fog and Taiwan (unfortunately, the nomenclature is inac- seasonality can alter the aforementioned factors and curately described). benefit species having seeds in specific seasons through The distribution of the associations included in Pasa- enhanced probability of regeneration. Some species have nio kawakamii-Machilion japonicae is often restricted to the ability to absorb nutrients from fog droplets (Lai certain ecoregions (ESM 6). Adinandro lasiostylae- et al. 2007). When increased nutrients input accompa- Chamaecyparidetum formosensis occurs mainly in the nies the peak of dense fog in the appropriate season, e.g., northwestern ecoregion and surrounding areas. Cyclo- the fruiting season of a species, these species can be more balanopsio stenophylloidis-Chamaecyparidetum formosen- successful in occupying a habitat by greater production sis and Pileo brevicornutae-Machiletum japonicae are of seeds than others. Fog-adapted species are sensitive to found in the eastern ecoregion, Castanopsio carlesii- short-duration droughts (Chu et al. 2014; Oliveria et al. Chamaecyparidetum formosensis in the southwestern 2014). Fog seasonality modifies drought duration, ecoregion, and Symploco wikstroemiifoliae-Machiletum causing various fog-adapted species to aggregate and thunbergii in the northeastern ecoregion. One of thereby assemble different vegetation types. There are the possible explanations for this restricted distribution is additional abiotic and biotic interactions correlated with sampling bias introduced by investigators (e.g., preference fog events in SMCFs. Further studies are required to for certain habitats or systematically incorrect determi- properly understand how fog events influence the com- nation of certain species). However, because plots position and structure of subtropical montane cloud belonging to the same association in our study were re- forest communities. peated by different investigators and plots sampled by the same investigator were often classified into different Acknowledgments We appreciate the efforts of the editors and two associations, we suspect that sampling bias is not the anomymous reviewers who greatly helped us to improve this paper. The Forestry Bureau of Taiwan significantly contributed to this reason for such a pattern. Another possible explanation study by supporting the National Vegetation Database of Taiwan. could be differences among local floras, which are caused The work was also supported by the Czech Science Foundation by variations in evolutionary history of individual re- [GAP 505/12/1022 to CFL and MC, and GAP 505/11/0732 to DZ]. gions. If this was the reason, diagnostic species of the aforementioned associations would have locally restricted distributions. This is clearly not the case, because most diagnostic species have wide geographic distributions Appendix throughout the island. Therefore, the most likely explanation of locally restricted vegetation types is environ- See Table 4. Author's personal copy 787

Table 4 Species data of nomenclature type releve´ s (header data is in Table 3)

1. 1. 1. 1. 1. 2. 2. 2. 2. 2. 2. Association code 01 02 03 04 05 06 07 08 09 10 11

Damnacanthus angustifolius S11––––0.5––––– Symplocos heishanensis T6–––0.2–––––– Ilex goshiensis T4–––11––––– Ilex tugitakayamensis T 2 0.3 – 0.2 – 0.5 – – 0.3 – – Lyonia ovalifolia T–14––––––––– Pinus armandii var. mastersiana T–5––––––––– Symplocos modesta S––4–––––––– Hymenophyllum polyanthos E 0.1 0.1 2 1 0.5 0.1 0.2 0.1 – 0.1 – Schefflera taiwaniana T0.5–20.5––––1–– Acrophorus stipellatus H – – 2 1 – 0.5 0.1 – – 0.2 – Photinia niitakayamensis T–––8––––––– Eurya glaberrima T––15––––1–– Prunus matuurai T–––3––––––– Ilex hayataiana T – 0.5 – 2 – – 0.3 1 – – – Arthromeris lehmannii H 0.5 0.1 – 2 – 0.5 – – – – – Diplopterygium chinensis H––––9–––––– Eurya strigillosa T––––5––0.50.5–– Ctenitis kawakamii H––––4–––––– Smilax arisanensis L 0.5 0.1 0.1 – 2 0.1 – – – – – Cyclobalanopsis sessilifolia T 0.5 – – – 0.5 11 – – – – – Sycopsis sinensis T–––––7––––– Symplocos migoi T0.1–1––4––––– Viburnum foetidum var. rectangulatum S––0.2––3––0.5–– Rhododendron rubropilosum S – 0.1 – – – – 17 – – – – Symplocos formosana S – 0.1 – – – – 15 – – 0.5 – Eurya acuminata S––––––12–––– Camellia brevistyla T––––––6–––– Pasania hancei var. ternaticupula T––––––3–––– Litsea morrisonensis T–0.3––––3–––– Vaccinium wrightii T–1––––3–––– Blastus cochinchinensis S–––––––10––– Asplenium normale H––––0.2––7–0.5– Lithocarpus lepidocarpus T––––––0.16––– Adinandra formosana T–––––––6–0.5– Gordonia axillaris T–––––––4––– Polystichum parvipinnulum H – – 1 – 0.5 0.1 0.1 2 – – 1 Sarcopyramis napalensis var. bodinieri H 0.1 – – 0.1 – – 0.1 2 – – – Miscanthus floridulus H––––––––43–– Strobilanthes flexicaulis H––––––––12–– Polygonum chinense H––––––––8–– Pellionia radicans H – – – – – 0.5 0.1 0.5 5 1 – Ellisiophyllum pinnatum H––––––––5–– Microsorium buergerianum H 0.1 – – 0.1 – 0.1 0.1 0.1 4 0.5 – Ligustrum sinense T–––––0.1––3–– Hydrangea angustipetala S––––––––3–– Diplazium dilatatum H–––––––––39– Elatostema lineolatum var. majus H–––––––––33– Fatsia polycarpa T–––––0.5––0.58– Sloanea formosana T–––––––––6– Asplenium antiquum E–––––––––4– Lasianthus fordii S–––––––––3– Pilea melastomoides H––––––––––45 Machilus zuihoensis var. mushaensis T––––––––––24 Pilea angulata H––––––––––15 Cyclobalanopsis stenophylloides T 0.5 – – – – 0.5 0.3 – – – 8 Maesa japonica S––––––––––5 Pourthiaea beauverdiana var. notabilis T––––––––––3 Author's personal copy 788

Table 4 continued

1. 1. 1. 1. 1. 2. 2. 2. 2. 2. 2. Association code 01 02 03 04 05 06 07 08 09 10 11

Arachniodes festina H––––––––––3 Pteris wallichiana H––––––––––3 Pilea aquarum subsp. brevicornuta H––––––0.1–––3 Diplazium amamianum H––––––––1–3 Tetrastigma umbellatum L––––––0.1–0.5–3 Urtica thunbergiana H––––––––0.1–3 Michelia compressa T––––––––––3 Acer serrulatum T––––––––––3 Cyrtomium hookerianum H–––––––––0.13 Selaginella mollendorﬃi H–––––––––0.13 Strobilanthes rankanensis H – – – – – 0.1 – – – 0.1 3 Alnus formosana T––––––––––3 Rhododendron leptosanthum T10–––17––1––– Trochodendron aralioides T3–0.2––2––––– Cleyera japonica var. taipinensis T2––3–0.1––––– Cyclobalanopsis morii T2––––––0.513–– Adinandra lasiostyla T2––––3––––– Rhododendron formosanum T – 14 – – 3 – 0.3 – – – – Myrsine stolonifera S–3––2–––––– Microtropis fokienensis S 1 2 0.1 – – 0.5 – 0.5 – 2 – Hydrangea chinensis S––21––––14––– Eurya leptophylla S––6––8––0.5–– Hydrangea integrifolia L 0.1 – 2 3 0.5 1 0.1 – 0.1 0.5 – Eurya crenatifolia S–––3––2––0.5– Litsea elongata var. mushaensis T – – – 2 0.2 0.5 – – 1 – 10 Barthea barthei S–––27––––0.5– Plagiogyria dunnii H––––27––6––– Diplazium kawakamii H––––3–––––10 Taiwania cryptomerioides T––––2–––31–– Acer morrisonense T–––––3––0.1–3 Cyclobalanopsis longinux T – 0.5 – – 0.2 – 2 – – 1 5 Castanopsis cuspidata var. carlesii T–––––––242–– Machilus thunbergii T–––––––5–15– Chamaecyparis obtusa var. formosana T37––356–––––– Symplocos wikstroemiifolia S–8––3––––3– Dendropanax dentiger T13–20.5––2––– Illicium anisatum T––383–––––– Symplocos arisanensis T–––3–4–3––– Monachosorum henryi H – – 0.5 – – 0.5 11 5 0.1 0.5 5 Damnacanthus indicus S – – – 0.2 1 1 11 5 – 0.1 3 Pasania kawakamii T–––––0.52––24 Prunus phaeosticta T – – – – 0.2 0.3 – 3 – 3 3 Yushania niitakayamensis S892–66––36–––– Symplocos morrisonicola T3–30.544––––– Plagiogyria euphlebia H 3 – – 4 8 1 0.1 2 0.1 0.2 – Eurya loquaiana S2–––14––14–18 Elaeocarpus japonicus T16––15–2––15 Litsea acuminata T––2–0.5––4–821 Machilus japonica T––––2–1–20726 Arachniodes rhomboidea H – – – 0.2 2 2 0.1 0.1 6 0.5 3 Tsuga chinensis var. formosana T 8 36 11 4 – 23 6 – – – – Dryopteris formosana H – 0.1 2 2 17 4 0.1 8 2 – – Plagiogyria formosana H 63 0.1 79 7 3 67 11 3 – – – Neolitsea acuminatissima T4383260.590.5–– Chamaecyparis formosensis T – – 33 – 16 19 25 23 11 36 45 Pyrrosia sheareri E – – – – – 0.1 0.1 – 0.5 – 0.1 Ligustrum liukiuense T – – – 0.2 – – 1 0.5 0.1 – – Author's personal copy 789

Table 4 continued

1. 1. 1. 1. 1. 2. 2. 2. 2. 2. 2. Association code 01 02 03 04 05 06 07 08 09 10 11

Asarum macranthum H – – – 0.3 – 0.1 0.1 – – 0.1 – Dryopteris sparsa H – – 0.1 – – 0.1 – 0.1 0.1 – – Hedera rhombea var. formosana L – – 0.5 – – 0.1 0.1 – 0.5 – – Skimmia reevesiana S 0.1 – – 0.2 – 0.5 – – – 0.5 – Elatostema trilobulatum H – – 0.3 0.5 – 0.1 0.1 0.1 – – – Pileostegia viburnoides L – 0.1 – 0.2 – – 0.1 – 0.1 – 0.1 Lysionotus pauciﬂorus H 0.1 – – – – 0.1 0.1 0.1 0.1 0.1 – Ardisia crenata S 0.1 – – 0.5 – 0.1 – 0.5 1 – 1 Araiostegia parvipinnata E – – 1 1 0.3 0.1 0.1 0.1 0.5 – 0.1 Stauntonia obovatifoliola L – – – 0.5 0.1 0.5 0.1 0.1 0.1 0.1 – Vittaria ﬂexuosa E 0.1 – 0.2 0.1 0.5 1 0.1 0.1 0.5 0.1 0.1

Association codes are 1.01, Tsugo formosanae-Chamaecyparidetum formosanae; 1.02, Vaccinio lasiostemonis-Tsugetum formosanae; 1.03, Schefflero taiwanianae-Chamaecyparidetum formosensis; 1.04, Elatostemato trilobulati-Tsugetum formosanae; 1.05, Rhododendro formosani- Chamaecyparidetum formosanae; 2.06, Adinandro lasiostylae-Chamaecyparidetum formosensis; 2.07, Cyclobalanopsio stenophylloidis- Chamaecyparidetum formosensis; 2.08, Castanopsio carlesii-Chamaecyparidetum formosensis; 2.09, Arachniodo rhomboideae-Chamaecy- paridetum formosensis; 2.10, Symploco wikstroemiifoliae-Machiletum thunbergii; 2.11, Pileo brevicornutae-Machiletum japonicae. Letters following the species names are T tree, S shrub, H herb, L liana, E epiphyte. Numbers in the table are percentage covers Other species whose cover is always less than or equal to 2 % and are present in less than four plots: Trees: Acer kawakamii 09:0.2, 10:0.5; A. palmatum var. pubescens 06:0.5; Carpinus rankanensis 07:0.5; Cephalotaxus wilsoniana 11:1; Cinnamomum subavenium 05:0.2, 10:2; Cleyera japonica 05:2; Daphniphyllum himalaense subsp. macropodum 06:2; Eurya gnaphalocarpa 07:0.5; Ficus erecta var. beecheyana 10:0.5; Ilex ficoidea 10:0.5; I. sugerokii var. brevipedunculata 01:0.1, 04:2; I. suzukii 04:1; Illicium arborescens 08:2; Itea parviflora 07:0.3, 10:2; Lithocarpus amygdalifolius 08:2; Morus australis 11:1; Neolitsea aciculata var. variabillima 04:2; Oreocnide pedunculata 09:0.1; Osmanthus heterophyllus 03:0.1; O. matsumuranus 07:0.5; Perrottetia arisanensis 09:1; Phoebe formosana 10:0.5; Photinia serratifolia 07:0.5; Pourthiaea villosa var. parvifolia 07:0.3; Prunus transarisanensis 02:1; Pyrenaria shinkoensis 10:2; Quercus tatakaensis 02:1, 07:0.5; Schima superba 05:2; Symplocos caudata 08:0.5; S. sonoharae 08:0.5; S. stellaris 01:0.1, 06:0.5; Ter- nstroemia gymnanthera 01:2, 04:0.2; Tetradium glabrifolium 09:0.5; Vaccinium kengii 05:2 Shrubs: Berberis kawakamii 03, 07:0.1, 04:0.2; B. mingetsuensis 06:0.5; Bredia oldhamii 08:2; Callicarpa formosana 09:2; C. randaiensis 04, 05, 08: 0.1; Daphne arisanensis 09:0.5; Euonymus spraguei 03, 06, 07: 0.1; Eurya chinensis 08:0.5; Lasianthus appressihirtus 10:0.2; Mahonia japonica 06:0.1; M. oiwakensis 09:0.5; Melastoma candidum 04:0.1; Pieris taiwanensis 03:0.1; Rhamnus crenata 04:0.5; R. pilushanensis 06:0.5; Rhododendron pseudochrysanthum 03:0.1; Rubus corchorifolius 04:0.5; R. croceacanthus 09:0.1; R. formosensis 07:0.1; R. kawakamii 06:0.1, 09:1; Sageretia thea 07:0.1; Salix fulvopubescens 09:0.2; Sambucus chinensis 09:0.1; Vaccinium japonicum var. lasiostemon 02, 04:0.1; V. merrillianum 10:0.2; V. randaiense 08:0.5; Viburnum erosum 04:0.5; V. integrifolium 08:2; V. luzonicum 04:0.5; V. propinquum 07:1; V. sympodiale 04:0.5; V. urceolatum 01, 06:0.1, 05:0.5 Herbs: Alpinia intermedia 10:0.1; Anoectochilus formosanus 09:0.1; Ardisia japonica 04:0.1; Arisaema formosanum var. brevipedunculatum 03:0.1; Asarum caudigerum 10:0.1; Athyrium arisanense 04:0.3; A. erythropodum 05:0.5, 06:0.1; A. subrigescens 06:0.1; Begonia formosana 10:0.3; Calanthe arcuata 06:0.1; C. densiflora 10:0.5; C. puberula 04, 06:0.1; Carex brunnea 08:0.5; Collabium chinense 10:0.5; Coniogramme intermedia 07:0.1, 11:1; Coptis quinquefolia 04:0.2, 10:0.1; Crypsinus engleri 04:0.1; C. quasidivaricatus 02:0.1, 03:0.5; Ctenitis apiciflora 03:1, 09:0.5; C. transmorrisonensis 03:1, 06:0.1; Cyclosorus subpubescens 04:0.1; Dennstaedtia scabra 06:0.1; Dictyocline griffithii 10:0.5; Diplazium wichurae 11:1; Diplopterygium glaucum 04:0.1; Disporopsis fuscopicota var. arisanensis 06:0.1; Dryopteris atrata 06:0.1; D. austriaca 05:0.2; Elatostema parvum 11:1; Eutrema japonica 09:0.1; Goodyera foliosa 06:0.1; G. kwangtungensis 06:0.1; G. velutina 06:0.1; Hemiboea bicornuta 10:0.2, 11:1; Hydrocotyle nepalensis 09:2; Lecanthus peduncularis 08:0.5; Lepisorus monilisorus 03, 10:0.1; 09:0.5; L. morrisonensis 02, 07:0.1; L. obscurevenulosus 06:0.1; L. thunbergianus 04, 08:0.1, 05:0.5; L. tosaensis 04, 08:0.1, 05:0.5; Leptorumohra quadripinnata 01:0.1; Loxogramme salicifolia 04, 06:0.1; Luzula effusa 07:0.1; Lycopodium serratum var. longipetiolatum 04, 08:0.1; Lysimachia ardisioides 07:0.1; Microlepia tenera 05:1; Myriactis humilis 04:0.1; Myrmechis drymoglossifolia 02:0.1; Nertera nigricarpa 03, 04: 0.1; Ophiopogon intermedius 04, 07:0.1; O. japonicus 06:0.5; O. japonica 06, 09:0.1; Oxalis acetosella subsp. griffithii var. formosana 03, 06:0.1, 04:0.5; Parathelypteris glanduligera 10:0.5; Peperomia reflexa 09:0.1; Peranema cyatheoides 03:0.2; Pilea plataniflora 09:0.1; Pla- giogyria stenoptera 10:0.1; Polygonum pilushanense 03:0.1; Polypodium amoenum 03:0.3, 04:0.1, 10:0.2; P. mengtzeense 06:0.1; P. formosanum 11:0.1; Polystichum hancockii 06, 07:0.1, 11:1; P. piceopaleaceum 08:2; P. stenophyllum 07:0.1; P. wilsonii 06:0.1; Pteris scabristipes 09:0.1; P. setulosocostulata 11:1; P. alboreticulata 01:0.5; Pyrola morrisonensis 04:0.1; Rubia lanceolata 03, 06:0.1; Rubus pectinellus 04, 06, 10:0.1; Sarcopyramis napalensis var. delicata 03:1; Selaginella delicatula 07:0.1, 10:2, 11:1; S. doederleinii 09:0.5; S. involvens 04:0.1; S. remotifolia 07:0.1; S. stauntoniana 07:0.1; Shortia rotundifolia 04:0.1; Viola adenothrix 05:0.1; V. formosana 07:0.1 Lianas: Actinidia chinensis var. setosa 09:0.1; Aristolochia heterophylla 07:0.1; Cayratia japonica 10:0.5; Celastrus kusanoi 06:0.1; Clematis grata 03:0.1; Embelia laeta var. papilligera 08:0.1, 10:0.5; Ficus pumila 07:0.1; F. sarmentosa var. nipponica 07, 11:0.1; Heterosmilax japonica 07:0.1; Lonicera acuminata 06, 07, 09:0.1; Maclura cochinchinensis 04:2, 05:0.2; Piper kadsura 07, 11:0.1, 10:0.2; Rhus ambigua 04:1; Rubus liuii 04:0.5; R. mesogaeus 10:0.1; R. swinhoei 07:0.1; R. trianthus 05:0.2; Schizophragma integrifolium var. fauriei 09:0.1; Smilax bracteata 09:0.1; S. china 09:1; S. discotis 04:0.5; S. riparia 03:0.1; S. lanceifolia 07:0.1, 08, 10:2; S. sieboldii 08:0.1; Stauntonia obovata 03:0.1; Thladiantha nudiflora 09:0.1; Trachelospermum jasminoides 07:0.1, 11:1; Tripterospermum taiwanense 06:0.1 Epiphytes: Asplenium wilfordii 06, 07:0.1; Bulbophyllum setaceum 03:0.1; Crepidomanes bilabiatum 10:2; Crypsinus echinosporus 06:0.1; Dendrobium furcatopedicellatum 01:0.1; Drymotaenium miyoshianum 01:0.1; Elaphoglossum yoshinagae 01, 06:0.1; Gastrochilus formosanus 01:0.1; Lemmaphyllum diversum 09:1; L. microphyllum 07, 08:0.1; Lepisorus pseudoussuriensis 07:0.1; Prosaptia contigua 10:0.1; Pyrrosia linearifolia 01:0.1; P. lingua 02, 07, 11:0.1; Rhododendron kawakamii 01:0.1; Vaccinium emarginatum 02, 08:0.1; Vandenboschia auriculata 07:0.1, 10:02; Xiphopteris okuboi 03:0.1 Author's personal copy 790

Jennings MD, Faber-Langendoen D, Loucks OL, Peet RK, Ro- References berts D (2009) Standards for associations and alliances of the U.S. National Vegetation Classification. Ecol Monogr Bruelheide H (1997) Using formal logic to classify vegetation. Folia 79:173–199 Geobot 32:41–46 Kent M (2012) Vegetation description and data analysis, 2nd edn. Bruelheide H, Chytry´ M (2000) Towards unification of national Wiley-Blackwell, Hoboken vegetation classifications: a comparison of two methods for Kocˇ ı´ M, Chytry´ M, Tichy´ L (2003) Formalized reproduction of an analysis of large data sets. J Veg Sci 11:295–306 expert-based phytosociological classification: a case study of Bruijnzeel LA, Scatena FN, Hamilton LS (2010) Tropical montane subalpine tall-forb vegetation. J Veg Sci 14:601–610 cloud forests: science for conservation and management. Kruskal JB (1964) Nonmetric multidimensional scaling: a numer- Cambridge University Press, Cambridge ical method. Psychometrika 29:115–129 Chen YF (2001) Vegetation of Taiwan—Chamaecyparis cloud Lai IL, Chang SC, Lin PH, Chou CH, Wu JT (2006) Climatic forest zone. Avanguardbook, Taipei (in Chinese) characteristics of the subtropical mountainous cloud forest at Chou CH, Chen TY, Liao CC, Peng CI (2000) Long-term eco- Yuanyang Lake Long-Term ecological research site, Taiwan. logical research in Yuanyang Lake forest ecosystem. I. Vegeta- Taiwania 51:317–329 tion composition and analysis. Bot Bull Acad Sinica 41:61–72 Lai IL, Schroeder WH, Wu JT, Kuo-Huang LL, Mohl C, Chou Chu HS, Chang SC, Klemm O, Lai CW, Lin YZ, Wu CC, Lin JY, CH (2007) Can fog contribute to nutrition of Chamaecyparis Jiang JY, Chen J, Gottgens JF, Hsia YJ (2014) Does canopy obtusa var. formosana? Uptake of a fog solute tracer into foliage wetness matter? Evapotranspiration from a subtropical mon- and transport to root. Tree Physiol 27:1001–1009 tane cloud forest in Taiwan. Hydrol Process 28:1190–1214 Lai YJ, Chou MD, Lin PH (2010) Parameterization of topographic Chytry´ M (ed) (2007) Vegetation of the Czech Republic 1. Grass- effect on surface solar radiation. J Geophys Res 115:D01104 land and heathland vegetation. Academia, Prague (in Czech) Lai YJ, Li CF, Lin PH, Wey TH, Chang CS (2012) Comparison of Chytry´ M, Tichy´ L, Holt J, Botta-Duka´ t Z (2002) Determination MODIS land surface temperature and ground-based observed of diagnostic species with statistical fidelity measures. J Veg Sci air temperature on complex topography. Int J Remote Sens 13:79–90 33:7685–7702 Chytry´ M, Schamine´ e JHJ, Schwabe A (2011) Vegetation survey: a Landucci F, Tichy´ L, Sˇ umberova´ K, Chytry´ M (2015) Formalized new focus for Applied Vegetation Science. Appl Veg Sci classification of species-poor vegetation: a proposal of a con- 14:435–439 sistent protocol for aquatic vegetation. J Veg Sci 26:791–803 Curtis JT (1959) The vegetation of Wisconsin: An ordination of Li CF, Chytry´ M, Zeleny´ D, Chen MY, Chen TY, Chiou CR, Hsia plant communities. University Wisconsin Press, Madison YJ, Liu HY, Yang SZ, Yeh CL, Wang JC, Yu CF, Lai YJ, Da LJ, Kang MM, Song K, Shang KK, Yang YC, Xia AM, Qi YF Chao WC, Hsieh CF (2013) Classification of Taiwan forest (2009) Altitudinal zonation of human-disturbed vegetation on vegetation. Appl Veg Sci 16:698–719 Mt. Tianmu, eastern China. Ecol Res 24:1287–1299 Lin CT, Li CF, Zeleny´ D, Chytry´ M, Nakamura Y, Chen MY, Forget PM, Lambert JE, Hulme PE, Wall SBV (2004) Seed fate: Chen TY, Hsia YJ, Hsieh CF, Liu HY, Wang JC, Yang SZ, predation, dispersal and seed establishment. CABI Pub, Cam- Yeh CL, Chiou CR (2012) Classification of high-mountain bridge coniferous forests in Taiwan. Folia Geobot 47:373–401 Fujiwara K (1996) Classification of plant communities in the Miyawaki A (1980) Vegetation of Japan 1. Yakushima. Shibundo Vegetation of Japan 1980–1989. Bull Inst Environ Sci Technol Publishing, Tokyo (in Japanese) Yokohama Natl Univ 22:23–79 Miyawaki A (1985) Vegetation of Japan 6. Chubu. Shibundo Geiger R (1966) The climate near the ground. Harvard University Publishing, Tokyo (in Japanese) Press, Cambridge Mueller-Dombois D, Ellenberg H (1974) Aims and methods of Gillies RR, Cui TNC, Kustas WP, Humes KS (1997) A verification vegetation ecology. John Wiley & Sons Inc, New York of the ‘triangle’ method for obtaining surface soil water content Oksanen J, Blanchet FG, Kindt R, Legendre P, Minchin PR, and energy fluxes from remote measurements of the Normalized O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Wagner Difference Vegetation Index (NDVI) and surface radiant tem- H (2013) vegan: community ecology package. R package ver- perature. Int J Remote Sens 18:3145–3166 sion 2.0-6. http://CRAN.R-project.org/package=vegan. Ac- Hamilton LS, Juvik JO, Scatena FN (1995) The Puerto Rico tropical cessed 30 April 2013 cloud forest symposium: introduction and workshop synthesis. Oliveria RS, Eller CB, Bittencourt PR, Mulligan M (2014) The In: Hamilton LS, Juvik JO, Scatena FN (eds) Tropical montane hydroclimatic and ecophysiological basis of cloud forest dis- cloud forests. Springer-Verlag, New York, pp 1–18 tributions under current and projected climates. Ann Bot Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A (2005) 113:909–920 Very high resolution interpolated climate surfaces for global Peet R, Roberts D (2013) Classification of natural and semi-natural land areas. Int J Climatol 25:1965–1978 vegetation. In: van der Maarel E, Franklin J (eds) Vegetation Horng FW, Ma FC, Yu HM, Hsui YR, Chang HM (2000) An ecology, 2nd edn. Wiley-Blackwell, Oxford, pp 28–70 estimation of original Chamaecyparis forest area in Taiwan and R Core Team (2013) R: a language and environment for statistical its implication for conservation. Q J Chin For 17:143–153 computing. R Foundation for Statistical Computing, Vienna, Hou XY (1983) Vegetation of China with references to its geo- Austria. http://www.R-project.org. Accessed 30 April 2013 graphical distribution. Ann Mo Bot Gard 70:509–549 Sandholt I, Rasmussen K, Andersen J (2002) A simple interpre- Hsieh CF (1989) Structure and floristic composition of the beech tation of the surface temperature/vegetation index space for forest in Taiwan. Taiwania 34:28–44 assessment of surface moisture status. Remote Sens Environ Huang TC, Hsieh CF (1994–2003) Flora of Taiwan, vol. I–VI, 2nd 79:213–224 edn., National Taiwan University, Taipei Schupp EW, Jordano P, Go´ mez JM (2010) Seed dispersal effec- Hukusima T, Lu SY, Matsui T, Nishio T, Liu CL, Pignatti S (2005) tiveness revised: a conceptual review. New Phytol 188:333–353 Phytosociology of Fagus hayatae forests in Taiwan. Rend Fis Sklena´ rˇ P, Bendix J, Balslev H (2008) Cloud frequency correlates to Acc Lincei 16:171–189 plant species composition in the high Andes of Ecuador. Basic Hukusima T, Matsui T, Nishio T, Pignatti S, Yang L, Lu SY, Kim Appl Ecol 9:504–513 MH, Yoshikawa M, Honma H, Wang Y (2013) Phytosociology Song YC (1999) Perspective of the vegetation zonation of forest of the beech (Fagus) forests in East Asia. Springer, Heidelberg region in eastern China. Acta Bot Sinica 41:541–552 (in Chi- Janisˇ ova´ M, Du´ bravkova´ D (2010) Formalized classification of nese) rocky Pannonian grasslands and dealpine Sesleria dominated Sørensen R, Zinko U, Seibert J (2006) On the calculation of the grasslands in Slovakia using a hierarchical expert system. topographic wetness index: evaluation of different method Phytocoenologia 40:267–291 based on field observations. Hydrol Earth Syst Sci 10:101–112 Author's personal copy 791

Su HJ (1984) Studies on the climate and vegetation types of the van der Maarel E, Franklin J (2013) Vegetation ecology: historical natural forests in Taiwan (II)—Altitudinal vegetation zones in notes and outline. In: van der Maarel E, Franklin J (eds) relation to temperature gradient. Q J Chin For 17:57–73 Vegetation ecology, 2nd edn. Wiley-Blackwell, Oxford, Su HJ (1985) Studies on the climate and vegetation types of the pp 45–59 natural forests in Taiwan (III)—A scheme of geographical cli- Weber HE, Moravec J, Theurillat JP (2000) International code of matic regions. Q J Chin For 18:33–44 phytosociological nomenclature, 3rd edition. J Veg Sci Su HJ (2002) The diversification and synthesis of vegetation clas- 11:739–768 sification. J NTU Exp For 16:269–282 (in Chinese) Wey TH, Lai YJ, Chang CS, Shen CW, Hong CY, Wang YN, Suzuki T (1952) The forest vegetation of East Asia. Kokin-Shoin, Chen MC (2011) Preliminary studies on fog characteristics at Tokyo (in Japanese) Xitou region of central Taiwan. J NTU Exp Forest 25:149–160 Suzuki T (1954) The forest vegetation of the north-Formosa (in Chinese) mountains. Jpn J Ecol 4:7–13 (in Japanese) Xu J, Wang XH (2010) Distribution and characteristics of the Tichy´ L (2002) JUICE, software for vegetation classification. J Veg mossy dwarf forest in China. J East China Nor Univ (Nature Sci 13:451–453 Science) 4:44–57 (in Chinese) Tichy´ L, Chytry´ M (2006) Statistical determination of diagnostic species for site groups of unequal size. J Veg Sci 17:809–818