植群構造及植物組成在不同微地貌及更新棲位的變化
-以北台灣福山試驗林陡峭林地為例
中文摘要
本研究探討植群構造、組成與微地貌及更新棲位間的相關性。研究地點為台灣北部福山試驗林的一塊2 ha常綠闊葉林，該樣區地勢陡峭，跨越頂坡及溪谷，包含不同微地貌及更新棲位。樣區劃分為7種微地貌(頂坡、上坡、下坡、山腳-溪谷、平緩山溝、陡峭山溝及崩塌地)及3種更新棲位(低孔隙、高孔隙及冠層鬱閉區)，並進一步比較植群構造(以枝葉層垂直結構、平均冠層高、林木胸高斷面積及密度為代表)及孔隙分布在不同微地貌的變化，以及樹種對不同微地貌及更新棲位之適應性。
調查結果指出，不同微地貌之植群構造有顯著差異。頂坡及上坡的有效土壤較深、坡度較為平緩﹔相較於其它微地貌，其林分發育良好，有較高的平均冠層高度，林木胸高斷面積及密度也較大。相反的，孔隙則集中在有效土壤淺薄，容易受到沖蝕之陡峭山溝及山腳-溪谷。崩塌地尚處於林分發育初期之孔隙階段。頂坡及上坡有較高比例的立枯木分布，幹折及根拔林木的分布則無明顯的集中現象。邊坡的穩定性及地表沖蝕所造成的土壤干擾，是影響山區森林結構分化及孔隙分布的重要因子。
樣區內紀錄有99個樹種，多數樹種出現位置與微地貌(92種)或更新棲位(76種)有顯著關聯性；其中稀少種較常見於下坡、山腳-溪谷、陡峭山溝及崩塌地微地貌或孔隙棲位，且多數屬於需光性高之陽性植物；相反的，常見種多生長在干擾度小的頂坡、上坡微地貌，或冠層鬱閉區。與常態對數分布曲線比較，樣區內稀少種之數量明顯偏高，增加了樣區內之樹種多樣性，且上述稀少種多偏好生長在樣區內相對面積較小的易受干擾微地貌，或孔隙棲位。依據不同生活階段的更新棲位適應性，可將17個冠層優勢樹種分為4個類群，大多數冠層優勢種在稚樹階段經常生長在冠層鬱閉區，在幼樹階段則趨向沒有明顯的更新棲位選擇性；部分樹種在不同生活階段對棲位的適應性會有所轉變。
頂坡及山腳-溪谷微地貌組合樣區間的定量物種轉換率為77.7%，低孔隙及冠層鬱閉區間的定量物種轉換率為34.9%；相對而言，微地貌分化對物種β多樣性的貢獻較更新棲位的分化顯著。綜合言之，在陡峭的山區內，微地貌及更新棲位之分化，是造成植群構造及植物組成分化的重要因子，亦是物種多樣性在小面積森林中維持的重要機制。

Variation in Vegetation Structure and Composition in Different
Micro-landforms and Regeneration Niches of a Steep Forest Plot
in the Fushan Experimental Forest, Northern Taiwan
Summary
The variation of vegetation structure and composition among micro-landforms and regeneration niches were examined in a 2-ha plot established on a slope extending from the mountain crest to bottomland in an evergreen broadleaf forest in the Fushan Experimental Forest, northern Taiwan. Seven micro-landforms (crest, upper slope, lower slope, foot slope-bottomland, smooth gully, steep gully, and landslide) and 3 regeneration niches (low-gap, high-gap, closed canopy) were recognized in this plot. Vegetation structure (represented by vegetation height profile, average canopy height, basal area of woody plants, and density of woody plants) and areas of gaps were compared among different micro-landforms. The chi-square test was used for demonstrating the association of certain species with different micro-landforms or regeneration niches.
There was significant difference in vegetation structure among micro-landforms. The vegetation developed better (higher average canopy height, larger basal area and density of woody plants) on the crest and upper slope where the effective soil was deeper and the inclination was flatter. In contrast, there were greater areas of canopy gaps recorded in the steep gully, foot slope-bottomlands and landslide patches where the effective soil was shallower due to soil erosion. Standing dead trees tend to aggregate on the crest and upper slope, while snapped and uprooted trees show no obvious aggregation. In conclusion, the regime of soil disturbance is the predominant factor influencing the vegetation structure and distribution of canopy gaps among different micro-landforms.
Ninety-nine woody species were recorded in 2 ha plot, and most of them significantly adapted to certain micro-landform (92 species) or regeneration niche (76 species). Many rare species tend to appear in the gap and/or micro-landform with frequent soil disturbance, such as lower slope, foot slope-bottomland, steep gully and landslide; and most of them are shade-intolerant. In contrast, common species often appear in closed canopy patch and/or the micro-landform with slight soil disturbance, such as crest and upper slope; and most of them are shade-tolerance species. There are more rare species in 2 ha plot while fitting to the lognormal distribution curve, and many rare species are shade-intolerant. Many rare species occurred in micro-landforms and/or gaps in which the soil is frequently disturbed. As a result, the α diversity in the plot is increased.
According to the regeneration niche selection among life stages, canopy tree (17 species) could be grouped into 4 categories. At sapling stage (dbh< 5 cm), most of canopy species (11 species) tend to exist in area of closed canopy; at young tree stage(5 cm≦dbh≦20 cm), most of canopy species (10 species) are evenly distribute among 3 regeneration niches. The regeneration niche adaptation may be modified at different life stages for some canopy species.
The species turnover rate between crest and foot slope-bottomland is 77.7%, which is much higher than that between low-gap and closed canopy. Therefore, the differentiation of micro-landform contributes more to β diversity than that of regeneration niche. In conclusion, micro-landform and regeneration niche differentiation is important mechanism for maintaining woody species diversity at local scale.

目 錄
附表目次 ………………………………………………………….....Ⅰ
附圖目次 ………………………………………………………….....Ⅱ
中文摘要 ………………………………………………………….....Ⅲ
英文摘要 ………………………………………………………….....Ⅴ
壹、緒言 ……………………………………………………………...1
貳、前人研究…………………………………………………………..3
參、材料與方法………………………………………………………..6
一、試驗地概述…………………………….………………….......6
二、調查方法………………………………….………………….....8
(一) 樣區測量及微地貌分類 ……………..……………….……...8
(二) 植物調查 ………………………………………………….....10
(三) 環境因子及林相因子之調查、評估 …………………….....11
(四) 植群高度剖面結構及更新棲位的標定方法 …………….....12
三、樣區資料建檔及分析方法 …………………………………....13
(一) 植物調查原始資料檔建立 ……………………………….....13
(二) 不同密度等級之物種豐富度 …………………………….....13
(三) 樹種α及β多樣性計算方式……….………………….......14
(四) 不同微地貌區塊的環境及林相因子比較……….…….......15
(五) 枯死木數量及孔隙在不同微地貌區塊的分布 ….……......16
(六) 樹種對微地貌及更新棲位偏好之分析方法 …..…..……...16
肆、結果……………………………………………………………...17
一、 微地貌的劃分…………………………….……………….....17
二、 植群高度剖面結構…………………………………..………..17
(一) 全區植群高度剖面結構 ………………..…………….......17
(二) 不同微地貌區塊之植群高度剖面結構……….…….........19
三、孔隙分布 ………………………………………………........20
四、枯死木分布 ……………………………………………........23
五、不同微地貌區塊的環境及林相因子………………….........24
(一) 環境因子 …………………………………………….........24
(二) 林相因子 …………………………………………….........25
(三) 環境及林相因子間的相關性 ……………………...........26
六、樣區植物組成及不同密度等級物種之豐富度 ………………..27
七、微地貌及更新棲位之物種多樣性 …………………………....31
（一） 不同微地貌及更新棲位的物種α多樣性 ………………...31
（二） 不同微地貌及更新棲位之物種β多樣性 ………………...33
八、 物種對不同微地貌及更新棲位之適應性 …………………….35
九、 優勢樹種在不同生活階段的更新棲位適應性 ……………….49
伍、討論 …………………………………………………..........51
一、植群構造的變化 ……………………………..……………....51
二、孔隙與枯死木的分布 ………………………….………….....54
三、樹種的微生育地適應性 ………………………………………..56
四、優勢種不同生活階段的更新棲位適應性……………………….58
五、微生育地分化對物種多樣性之影響 …………..………………60
陸、結論 …………………………………….……………….......65
柒、引用文獻 ………………………………………………........67
附錄 1. 樣區植物名錄 …………………………………………....74
附錄 2. 調查樣區內不同更新棲位組合樣區之物種組成 ………..79
附錄 3. 調查樣區內不同微地貌組合樣區之物種組成 …………..82
附表目次
表 1. 三種更新棲位對不同微地貌適應性之卡方檢驗結果 ……….22
表 2. 枯死木(胸徑≧20 cm)對不同微地貌適應性之卡方檢驗結果.24
表 3. 不同微地貌區塊之環境及林相因子之變異數分析及鄧肯多
變域分析比較 ………………………………………………...25
表 4. 樣區環境及林相因子間之相關性比較 ……………………….26
表 5. 樣區紀錄物種之相對密度、相對優勢度及重要值 ………….28
表 6. 不同微地貌區塊內紀錄之植物種數及Shannon多樣性指數 ..32
表 7. 不同更新棲位出現之植物種數及Shannon多樣性指數 ……..32
表 8. 不同微地貌區塊間之物種組成相異性(物種轉換率) ……….35
表 9. 不同更新棲位間之物種組成相異性(物種轉換率) ………….35
表 10. 樣區木本植物對不同微地貌適應性之卡方檢驗結果 ……..38
表 11. 樣區木本植物對不同更新棲位適應性之卡方檢驗結果 …..42
表 12. 優勢種在不同生活階段對更新棲位適應性的卡方檢驗結果.50
表 13. 台灣東北內陸區（northeast inland region）楠櫧林帶之
植群調查結果比較 ……………………………………………64
附圖目次
圖 1. 永久樣區於福山試驗林之設置地點 …………………………8
圖 2. 坡面微地貌之分類方式(仿Hara et al. 1996) ………… 10
圖 3. 微地貌在調查樣區之分布 …………………………………..18
圖 4. 全樣區之植群剖面結構 ………………………………………19
圖 5. 七個微地貌區塊之植群剖面結構 ……………………………20
圖 6. 三種更新棲位在調查樣區之分布 ……………………………21
圖 7. 樣區木本植物在不同對數密度等級之分布頻度 ……………31
圖 8. 七種不同微地貌區塊組合樣區在DCA第1及第2軸之分布....33
圖 9. 不同密度樹種在不同更新棲位之實測值與期望值差異之卡
方檢驗………………………………………………………….46
圖 10. 不同密度樹種在不同微地貌之實測值與期望值差異之卡
方檢驗...………………………………………………….…47
圖 11. 樣區喬木(成熟植株胸徑>5cm)種類在不同對數密度等級
之分布頻度 ………………………………………………….62

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