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Forest Vegetation of Taiwan, Based on Species Composition

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Forest Vegetation of Taiwan, Based on Species Composition MASARYK UNIVERSITY Faculty of Science Department of Botany and Zoology Diversity of mountain forest vegetation in Taiwan Ph.D. Dissertation Ching-Feng Li Supervisor: prof. RNDr. Milan Chytrý, Ph.D. Brno 2013 1 BIBLIOGRAPHIC ENTRY Author: Ching-Feng Li, M.Sc. Faculty of Science, Masaryk University Department of Botany and Zoology Title of Dissertation: Diversity of mountain forest vegetation in Taiwan Degree Programme: Biology Field of Study: Botany Supervisor: prof. RNDr. Milan Chytrý, Ph.D. Faculty of Science, Masaryk University Department of Botany and Zoology Academic Year: 2013/2014 Number of Pages: 235 Keywords: Altitudinal zones; Azonal vegetation; Classification and regression trees; Cloud forest; Cocktail Determination Key; Eastern Asia; High-montane coniferous forest; Monsoon; Montane forest; Plant community; Seasonality of moisture; Subtropical forest; Supervised classification; Topography; Tropical forest; Zonal vegetation 3 BIBLIOGRAFICKÝ ZÁZNAM Autor: Ing. Ching-Feng Li Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Diverzita horské lesní vegetace na Tchaj-wanu Studijní program: Biologie Studijní obor: Botanika Školitel: prof. RNDr. Milan Chytrý, Ph.D. Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Akademický rok: 2013/2014 Počet stran: 235 Klíčová slova: azonální vegetace; determinační klíč založený na metodě Cocktail; horské lesy; klasifikační a regresní stromy; mlžný les; monzun; rostlinná společenstva; řízená klasifikace; sezónní výkyvy vlhkosti; subtropický les; topografie; tropický les; východní Asie; vysokohorské jehličnaté lesy; výšková zonace; zonální vegetace 5 ABSTRACT In Taiwan, vegetation studies began from the beginning of the 20th century. Due to different approaches applied in vegetation classification, different methods of vegetation data analysis and diverse nomenclature systems used among studies, there is still a lack of a general concept which can be used to understand the ecological patterns of vegetation distribution. In order to have a comprehensive description and definition of plant communities, to provide systematic documentation of plant communities and to understand the relationships of plant communities in Taiwan with those in the neighbouring areas, the use of representative vegetation-plot data to generate a vegetation classification system is necessary. For this purpose, the main aims of this dissertation are: 1) To determine the main forest vegetation types in Taiwan at higher hierarchical levels. 2) To delimit selected vegetation types at the association level. 3) To create formal definitions for all defined forest vegetation types, describe their habitat affinities and their distribution in Taiwan. 4) To construct a hierarchical vegetation classification system. To achieve these aims, vegetation classification was performed following the Braun-Blanquet approach, using the National Vegetation Database of Taiwan, which contains 9822 vegetation plots. Formal definitions of each forest vegetation type were given by unequivocal assignment rules via classification and regression trees or the Cocktail Determination Key. Habitat affinities were explored by ordination analyses. While constructing the hierarchical vegetation classification system, corresponding forest vegetation types in the surrounding area were evaluated by both floristic composition and habitat requirements. The first question was how many main forest vegetation types are there in the whole of Taiwan. The next question focused on the associations and the hierarchical vegetation classification system of forests dominated by coniferous trees, including both the species- poor high-mountain coniferous forests with Abies, Juniperus, Picea and Tsuga, and species-rich subtropical montane cloud forests with Chamaecyparis mixed with evergreen broad-leaved trees. The main conclusions are: 1) Twenty-one main forest vegetation types were distinguished, mainly corresponding to the temperature and moisture differences among habitats, in the subtropical and tropical region in Taiwan. The classification was formalized by the Cocktail Determination Key. 2) In the high-mountain coniferous forests, two alliances and nine associations were recognized based on dominant species. Altitude, topography, snow, wind and soil conditions explain the habitat requirements of these associations. A dichotomous key to determining associations was prepared based on the results of classification and regression trees. 3) In the subtropical montane cloud forests dominated by Chamaecyparis spp., two alliances and eleven associations were recognized and their diagnostic species groups defined. Altitude, soil conditions and the seasonality of moisture in different ecoregions explain the habitat requirements of these associations. The classification was formalized by the Cocktail Determination Key. 4) Altitude and topography explain the differences of floristic composition at the level of alliance in the high-mountain coniferous forests and the subtropical montane cloud forests, respectively. 5) Forest vegetation types in the subalpine, upper-montane and montane cloud zones of Taiwan, and those in subtropical mainland China, should be classified to the same phytosociological classes, which are different from those occurring in the temperate and boreal zones of eastern Asia because of the dominance of evergreen broad-leaved species. 7 ABSTRAKT První vegetační studie na Tchaj-wanu začaly na počátku 20. století. Dodnes však chybí jednotný koncept vegetační klasifikace, který by umožnil pochopení ekologických zákonitostí v rozšíření vegetačních typů. Je to především proto, že autoři jednotlivých studií používali různé přístupy ke klasifikaci vegetace, různé metody analýzy vegetačních dat a různou nomenklaturu. Pro získání detailního popisu, definice a systematické dokumentace rostlinných společenstev a pro pochopení vztahu vegetace na Taiwanu k vegetaci v okolních zemích je zapotřebí vybudovat jednotný klasifikační systém, založený na reprezentativní databázi vegetačních zápisů. K naplnění tohoto cíle má pomoci i tato disertační práce, jejíž hlavní cíle jsou: 1) popsat hlavní typy lesní vegetace Tchaj-wanu na vyšších hierarchických úrovních, 2) pro některé z nich popsat vegetační typy na úrovni asociací, 3) vypracovat formální definice pro všechny popsané typy lesní vegetace a popsat jejich stanovištní nároky a rozšíření na Tchaj-wanu, 4) sestavit hierarchický systém vegetační klasifikace. Pro vybudování hierarchického klasifikačního systému, založeného na Braun-Blanquetově metodě, byla použita Národní vegetační databáze Tchaj-wanu, která aktuálně obsahuje 9822 vegetačních zápisů. Formální definice jednotlivých vegetačních typů byly založeny na jednoznačných přiřazovacích pravidlech, které byly získány jednak metodou klasifikačních a regresních stromů, jednak pomocí determinačního klíče založeného na metodě Cocktail. Stanovištní nároky byly popsány pomocí ordinačních metod. Při sestavování hierarchického klasifikačního systému lesní vegetace Tchaj-wanu bylo přihlíženo k existenci analogických vegetačních typů v okolních zemích, k jejich floristickému složení a stanovištním nárokům. Nejdříve bylo třeba zjistit, kolik hlavních typů lesní vegetace je možné na Tchaj-wanu vlastně rozlišit. V dalším kroku byl připraven popis asociací a sestavena hierarchická klasifikace lesní vegetace s dominancí jehličnatých dřevin, zahrnující jak druhové chudé vysokohorské jehličnaté lesy s druhy rodů Abies, Juniperus, Picea a Tsuga, tak i druhově bohaté subtropické horské mlžné lesy s charakteristickým výskytem rodu Chamaecyparis a stálezelených listnatých dřevin. Hlavní závěry této práce jsou následující: 1) Bylo rozlišeno 21 hlavních typů lesní vegetace Tchaj-wanu, které se odlišují hlavně nároky na teplotu a vlhkostními poměry stanoviště. K formalizaci této klasifikace byl použit determinační klíč založený na metodě Cocktail. 2) V rámci vegetace vysokohorských jehličnatých lesů byly rozlišeny dva svazy a celkem devět asociací, a to především na základě dominance některých druhů. Stanovištní nároky těchto asociací se liší faktory prostředí jako je nadmořská výška, topografie, výskyt sněhové pokrývky, síla větru a vlastnostmi půdy. Metodou klasifikačních a regresních stromů byl získán dichotomický klíč k určování jednotlivých asociací. 3) V rámci vegetace horských cypřiškových mlžných lesů byly rozlišeny dva svazy a celkem 11 asociací a definovány skupiny diagnostických druhů. Rozdíly ve stanovištních nárocích jednotlivých asociací jsou dány především rozdíly v nadmořské výšce, vlastnostech půdy a sezónnosti srážek mezi jednotlivými ekoregiony. Pro formalizaci klasifikace byl použit determinační klíč založený na metodě Cocktail. 4) Svazy rozlišené v rámci vysokohorských jehličnatých lesů lze vysvětlit především rozdíly v nadmořské výšce, zatímco rozdíly mezi svazy cypřiškových mlžných lesů jsou způsobeny především rozdíly v topografii. 5) Vegetační typy lesní vegetace v subalpinské, vysokohorské a mlžné vegetační zóně na Taiwanu a v subtropické části kontinentální Číny by měly být zahrnuty do stejných tříd, které jsou zároveň odlišné od tříd vyskytujících se v temperátní a boreální zóně východní Asie. Důvodem je především výskyt a někdy i dominance stálezelených listnatých dřevin. 9 中文摘要 台灣的植群研究開始於 20 世紀初期；在植群分類的研究上，不同的研究採用不同的分類概念， 不同的資料分析方法，和不同的命名系統，使得台灣的植群分類系統一直缺乏很明確的植相定 義，尤其是統一的而且可以完整陳述台灣生態系統的植相定義。為了對台灣的植群做完整的定 義和描述，並對台灣的植群提供系統性的建檔記錄，以及瞭解台灣的植群和鄰近地區的植物社 會彼此間的關聯，本論文的主要目的如下： 1) 瞭解台灣的森林植群在較高的分類階層中有幾種主要的類型。 2) 判識所選定的森林植群中各有多少種群叢
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