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 1 23 Your article is protected by copyright and all rights are held exclusively by The Ecological Society of Japan. This e-offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 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 com- position 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 com- monly 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.