第四章 討論
第四節 促進與競爭
植物演替的機制極為複雜，物種之間存在著促進或競爭的交互關 係，彼此之間因環境或資源壓力不同而產生不同的交互作用，如 Callaway 和 King（1996）在溼地的研究中發現，在低溫環境下，香 蒲（Typha latifolia）的根增加土壤中的氧含量以促進濕地勿忘草
（Myosotis laxa）的生長；但在高溫的環境下卻因微生物及根的呼吸 作用增加，使得土壤中的氧含量減少，進而造成香蒲與濕地勿忘草成 為競爭的關係。Greenlee 和 Callaway（1996）在蒙大拿草原研究發現，
氣候涼爽且濕潤的生長季，叢生草（bunchgrass）與十字花科的植物
（Lesquerella carinata）呈現相互競爭的關係；但在氣候乾熱的生長 季，Lesquerella 因叢生草的遮蔭效果，因而減少缺水的壓力，此時叢 生草和 Lesquerella 之間轉變為正向的關係。本研究與 Chapin（1994）
研究的結果，似乎也呈現相似的現象，在低氮沈降的阿拉斯加地區，
演替早期缺氮的環境，赤楊藉由增加土壤中的氮含量促進雲杉小苗的 生長；但在高氮沈降的臺灣地區，土壤中的氮可能較不缺乏，因此赤 楊與二葉松小苗則為競爭關係。由以上的例子可以看出，物種之間的 關係會因不同的環境而有不同的交互作用，因此未來在植物群落的研 究上，除了瞭解物種組成、物種之間的交互關係及環境狀況外，若能 清楚環境的改變對於生物之間的關係所造成的影響，將對於瞭解植群
27
結構、功能及動態有更深一步的認識。
本研究的赤楊並未促進松樹小苗的生長，然而已生長於此的臺灣 二葉松植群是經由何種演替機制才能成為如今的優勢物種，是個值得 探究的問題。氮沈降的升高是近幾十年的事情，現今生長於此的松樹 植群在古代可能的確是經由赤楊的促進作用而成為現今的優勢物種，
因此在氮沈降量不高的古代，赤楊在植群演替上或許的確是扮演著重 要的角色；但如今卻因過多的氮經由大氣沈降進入此生態系，因而造 成松樹小苗無法藉由赤楊所提供的氮而得到生長上的幫助。但相反的，
也有可能從古至今松樹小苗的生長其實並未受到赤楊的促進，松樹小 苗經由耐陰的方式突破先驅物種的遮蔭，因此最後成為現今的優勢物 種；但如今的高氮沈降環境可能加速松樹小苗的生長，因此縮短松樹 小苗突破遮蔭所需耗費的時間，加速植群演替的過程。然而赤楊未促 進松樹小苗的生長是否對於畢祿溪植群的演替或物種組成造成嚴重 的影響，這些推論都需要更多的歷史資料及後續的調查和研究才能得 到驗證，也將更能明確的指出高氮沈降對於森林演替所造成的影響。
28
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附錄
圖 1、試驗地位置及環境示意圖。（A）試驗地點位置。（B）河階環 境示意圖。
38
圖 2、各階層土壤砏黏土含量比較。誤差線為標準誤（standard error, SE）。各資料點上方有任一相同字母表示 Tukey 事後比較差異不顯 著（P > 0.05）。
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圖 3、各階層土壤氮含量比較。（A）銨根離子濃度比較。 （B）硝 酸根離子濃度比較。誤差線為標準誤（standard error, SE）。各資料點 上方有任一相同字母表示 Tukey 事後比較差異不顯著（P > 0.05）。
40
圖 4、各階層有效物種數量比較。誤差線為標準誤（standard error, SE）。 各資料點上方有任一相同字母表示 Tukey 事後比較差異不顯著（P >
0.05）。
41
圖 5、赤楊與土壤氮含量之關係。（A）赤楊植被覆蓋率與土壤氮離 子含量之關係。（B）赤楊枯落物覆蓋率與土壤氮離子含量之關係。
42
圖 6、赤楊與松樹小苗葉內氮濃度之關係。（A）小苗周圍赤楊株數 與小苗葉內氮濃度之關係。（B）小苗周圍赤楊枯落物覆蓋率與小苗 葉內氮濃度之關係。
43
圖 7、松樹小苗葉內氮濃度與其小苗相對生長高度之關係。
44
圖 8、赤楊與松樹小苗相對生長高度之關係。（A）小苗周圍赤楊株 數與其小苗相對生長高度之關係。（B）小苗周圍赤楊枯落物覆蓋率 與其小苗相對生長高度之關係。
45
圖 9、赤楊與松樹小苗接受光量之關係。（A）小苗周圍赤楊株數與 小苗周圍直射光立地係數之關係。（B）小苗周圍赤楊枯落物覆蓋率 與小苗周圍散射光立地係數之關係。
46
圖 10、松樹小苗接受光量與小苗相對生長高度之關係。（A）小苗周 圍直射光立地係數與小苗相對生長高度之關係。（B）小苗周圍散射 光立地係數與小苗相對生長高度之關係。
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表 1、各樣線調查到的植物名錄 第一條樣線
科名 中文 學名
Aceraceae 槭樹科
臺灣紅榨槭 Acer rubescens Hayata
Asteraceae 菊科
兔兒菜 Ixeris chinensis (Thunb.) Nakai 阿里山薊 Cirsium arisanense Kitam.
野茼蒿 Conyza sumatrensis (Retz.) Walker
裂葉蔓黃菀 Senecio scandens Buch.-Ham. ex D. Don. var.
incisus Franch.
黃菀 Senecio nemorensis L. var. dentatus (Kitam.) H.
Koyama
黃鵪菜 Youngia japonica (L.) DC.
鼠麴草 Gnaphalium luteoalbum L. subsp. affine (D.
Don) Koster
臺灣澤蘭 Eupatorium cannabinum L. var. asiaticum Kitam.
歐洲黃菀 Senecio vulgaris L.
Boraginaceae 紫草科
琉璃草 Cynoglossum furcatum Wall.
Caprifoliaceae 忍冬科
臺東莢蒾 Viburnum taitoense Hayata
Onagraceae 柳葉菜科
黑龍江柳葉菜 Epilobium amurense Hausskn. subsp. amurense
Pinaceae 松科
臺灣二葉松 Pinus taiwanensis Hayata 臺灣雲杉 Picea morrisonicola Hayata
Polygonaceae 蓼科
火炭母草 Polygonum chinense L. var. chinense
臺灣何首烏 Polygonum multiflorum Thunb. var. hypoleucum
48
(Ohwi) Liu, Ying & Lai Ranunculaceae 毛茛科
梨山小簑衣藤 Clematis gouriana Roxb. ex DC. subsp.
lishanensis Yang & Huang
第二條樣線
科名 名稱 學名
Aceraceae 槭樹科
青楓 Acer serrulatum Hayata
臺灣紅榨槭 Acer rubescens Hayata
Apocynaceae 夾竹桃科
細梗絡石 Trachelospermum gracilipes Hook. f.
Aquifoliaceae 冬青科
苗栗冬青 Ilex bioritsensis Hayata
Araliaceae 五加科
臺灣常春藤 Hedera rhombea (Miq.) Bean var. formosana (Nakai) Li
Asteraceae 菊科
艾 Artemisia indica
兔兒菜 Ixeris chinensis (Thunb.) Nakai 阿里山薊 Cirsium arisanense Kitam.
野茼蒿 Conyza sumatrensis (Retz.) Walker
裂葉蔓黃菀 Senecio scandens Buch.-Ham. ex D. Don. var.
incisus Franch.
黃鵪菜 Youngia japonica (L.) DC.
鼠麴草 Gnaphalium luteoalbum L. subsp. affine (D.
Don) Koster
臺灣澤蘭 Eupatorium cannabinum L. var. asiaticum Kitam.
蔓黃菀 Senecio crataegifolius Hayata
Betulaceae 樺木科
阿里山千金榆 Carpinus kawakamii Hayata
49
臺灣赤楊 Alnus formosana (Burkill ex Forbes & Hemsl.) Makino
Boraginaceae 紫草科
琉璃草 Cynoglossum furcatum Wall.
Caprifoliaceae 忍冬科
阿里山忍冬 Lonicera acuminata Wall. ex Roxb.
臺東莢蒾 Viburnum taitoense Hayata
Cyperaceae 莎草科
中國宿柱薹 Carex sociata Boott Fagaceae 殼斗科
狹葉櫟 Quercus stenophylloides Hayata
Myrsinaceae 紫金牛科
小葉鐵仔 Myrsine africana L.
Oleaceae 木犀科
小實女貞 Ligustrum sinense Lour.
Onagraceae 柳葉菜科
黑龍江柳葉菜 Epilobium amurense Hausskn. subsp. amurense
Pinaceae 松科
臺灣二葉松 Pinus taiwanensis Hayata 臺灣雲杉 Picea morrisonicola Hayata
Poaceae 禾本科
曲芒髮草 Deschampsia flexuosa (L.) Trin.
Polygonaceae 蓼科
火炭母草 Polygonum chinense L. var. chinense
臺灣何首烏 Polygonum multiflorum Thunb. var. hypoleucum (Ohwi) Liu, Ying & Lai
Rosaceae 薔薇科
Rosaceae 薔薇科