國立臺灣大學森林環境暨資源學系碩士論文 Thesis Submitted in partial fulfillment of the requirements for the Degree of Master of Science in the School of Forestry and Resource Conservation National Taiwan University 指導教授:丁宗蘇博士 Adviser:Dr. Ding, Tzung-Su 東亞鳥類生物多樣性熱點之選擇與分析 Selections and Analyses of Avian Biodiversity Hotspots in East Asia 研究生:何麗君 By : Ho, Li-June 中華民國九十四年七月 July, 2005
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國立臺灣大學森林環境暨資源學系碩士論文
Thesis Submitted in partial fulfillment of the requirements for the Degree of Master of Science in the School of
Forestry and Resource Conservation National Taiwan University
指導教授:丁宗蘇博士 Adviser:Dr. Ding, Tzung-Su
東亞鳥類生物多樣性熱點之選擇與分析
Selections and Analyses of Avian Biodiversity Hotspots in East Asia
研究生:何麗君 By : Ho, Li-June
中華民國九十四年七月 July, 2005
I
謝 誌
首先謝謝口試委員李培芬老師以及邱祈榮老師的指正及建議,讓這篇論
文能夠更加完善。
這篇論文能夠完整的呈現,最要感謝的是指導教授丁宗蘇老師。不論課
堂中科學知識的講授傳遞,或是討論時不斷激盪的啟發,老師的教學熱忱及
執著未曾改變。老師教導我的許多道理及態度讓我受用無窮,也成為我鞭策
要更往前進的動力。謝謝袁孝維老師,在我陷入迷惑谷底的時候,聆聽、分
享、關心以及鼓勵我。老師的話讓我從不同的角度看待事情的兩面,
謝謝佳恩和欣怡,常常擔心我、照顧我,有需要的時候立刻能伸出援手、
義不容辭,謝謝你們!謝謝雯菁,在發生意外時,慷慨的立刻答應讓我們借
住他家一個月。謝謝朝婷和怡平,能夠提供我許多珍貴的意見,認識你們就
像打開一扇窗,能夠瞭望世界的天空。也謝謝中芃,微笑樂觀的精神,會一
直陪著我度過風雨。謝謝中衍和元均,難過的時候安慰,高興的時候分享,
總是能告訴我許多的資訊,謝謝你們。也謝謝淑瑋,認真的態度使我獲益良
多。謝謝煥彰,開朗的個性還有不吝嗇的付出幫忙,幫助我度過去年多事的
夏天。還有學弟杜建宗,在每個研究室沈悶的時候總不忘帶給我們許多歡笑。
感謝老天爺讓我擁有一群非常好的朋友,過去曾經一起在 306、308 和 403
的所有伙伴們,很高興我們不論悲傷快樂,都一起相互扶持度過。
研究所三年走來,歷經了許多不同的人生經驗,感觸良多。謝謝招偉總
是在我身旁,包容我的不完美並永遠支持我,謝謝你。
最後感謝我家人,幫我處理部份大量運算的昌陽,讓我節省不少時間。
和我同住、一直照顧我的姊姊,還有雖然很忙,但常常打電話關心我的哥哥;
以及我偉大的父母,沒有他們的體諒和支持就沒有今天的我,謝謝你們所給
我的這一切。
II
摘 要
生物多樣性熱點的選擇與分析對於進行保育區的選擇有相當大的助益。
本研究以 100×100 km 網格劃分東亞地區鳥類分佈圖層,比較(1)所有物種豐
富物、(2)狹佈種豐富度、(3)瀕危種豐富度、以及(4)互補法四種方法所選擇的
生物多樣性熱點網格相關性;更進一步討論,不同分類群其物種多樣性之空
間相關性,以及屬豐富度及科豐富度是否能夠作為物種豐富度之指標物種。
研究結果發現,不同方法所得到的生物多樣性熱點彼此之間相關性不
高,以互補法所選取之熱點在相同之面積下較能包含較多之物種,且屬豐富
度和種豐富度間具有相當高之相關性。因此,不同方法所得到的生物多樣性
熱點難以互相替代。在進行保護區選擇時,可優先考慮狹佈種以及瀕危種為
對象,而且若能應用互補法,則能夠使同樣面積擁有最大物種數,使保育工
作更有效率。若在野外調查種豐富度有其困難時,可考慮以屬豐富度來代替
種豐富度。
III
Abstract
The selections and analyses of biodiversity hotspot are critical for
establishing protected areas. Using a 100×100 km grid system of bird breeding
ranges in East Asia, I compared the spatial correlation of biodiversity hotspots
selected by (1) total species richness, (2) limited-ranged species richness, (3)
endangered species richness, and (4) complementary method. Furthermore, I
examined the distribution of species richness of various avian orders and families
and the correlation among species richness, genus richness, and family richness.
I found that there were low spatial correlations among biodiversity hotspots
selected by different methods. With same area, the hotspots selected by
complementary method covered more species than other methods. There were
high correlations among species richness, genus richness, and family richness. I
conclude the hotspots of total species, limited-ranged species, and endangered
species were not congruent with each other. In selecting protected areas,
hotspots of limited-ranged species and endangered species should be given
priorities and using complementary method could maximize the species covered
by a given area and increase the efficiency of protected areas. Genus richness
could be a good surrogate of species richness if obtaining species richness is
危種數(e.g. Lombard, 1995; Benayas and de la Montaña, 2003)。大部分研究指
出,當網格所包含之物種豐富度很高時,狹佈種以及瀕危種被包含的數量及
機率也相對提高(Curnutt et al., 1994; Gaston, 1995; Reid et al., 1998; Bonn et
al., 2002; Gjerde et al., 2004)。本研究也呈現類似結果,不同類別之生物多樣
性值之間呈現顯著正相關。然而就不同類別之生物多樣性熱點而言,本研究
中所有物種之生物多樣性熱點多位於亞洲大陸,狹佈種之生物多樣性熱點多
位於華萊士區島嶼,瀕危種之生物多樣性熱點多位於蘇門答臘與婆羅洲,三
23
類熱點網格間的空間相關性並不高。這樣的結果與前人研究大致相符,
(Prendergast et al., 1993; Williams and Humphries, 1994; Lombard, 1995;
Gaston and Blackburn, 1996; Muriuki et al., 1997; Griffin, 1999; Lawton et al.,
1998; Fjeldsa, 2000; Sfenthourakis and Legakis, 2001; Awad et al., 2002; Bonn et
al., 2002; Ojeda et al., 2003; Fox and Beckley, 2005),亦即依照不同方法所選取
的熱點網格其空間相關性並不高,不同類別熱點間難以互相取代。
本研究所探討之三類不同面積定義之狹佈種(E5、E10、E20),其所產
生之生物多樣性值彼此相關程度相當大,彼此熱點的重疊度也相當高。因此,
雖然狹佈種之面積定義不一,但對狹佈種之熱點分布應不會有太大影響。而
就瀕危種之熱點分布,本研究之瀕危種生物多樣性值計算方式包括未加權、
及依物種族群數量及受威脅程度予以加權。結果顯示三者之生物多樣性值彼
此相關程度相當高,且其空間分佈也大致相同,故在未來後續研究中,可不
需考慮對物種族群數量及受威脅程度予以加權。
互補法
生物多樣性熱點若選取數量過多,將難以有效的執行保育工作,尤其在
牽涉到跨國性的範圍時,困難度更高(Mace et al., 2000; Humphries, 2001;
Brummitt and Lughadha, 2003)。互補法可以幫助輕易瞭解在設選擇置保護區
時,最少應該要有多大的面積範圍方能保護最多的物種,以期達到以最小面
積發揮最大功效之能(Williams et al., 1996; Howard et al., 1998; Reyers et al.,
2000; Peterson et al., 2000; Bonn et al., 2002; Lund and Rahbek,2002; Ojeda et
al., 2003; Sauberer et al., 2004),因此互補法的應用可以避免選取數量過多之
熱點。另一方面,若不使用互補法計算熱點網格間的物種,也將不會知道網
格中會有多少物種跟其他熱點網格重複(Margules and Pressey, 2000)。本研
24
究結果顯示,若使用互補法選擇生物多樣性熱點,僅需要 1.2%至 1.5%網格,
便可以保育 95%的所有物種、狹佈種及瀕危種。但若以非互補法方法選取熱
點,則分別需要 64.5%至 66.4%網格,方能保育 95%的物種數。由物種累積
曲線,也可以發現,互補法所累積物種的速度,遠高於傳統的熱點選取方法。
本研究所用之 100×100 km 的網格,對絕大部分鳥類而言,其一萬平方公里
之面積應能足夠支持健康之族群,而達到有效保護該物種之目的。因此,我
認為以互補法選擇保育區不僅能確保每個物種接受到保育,而且也能建立有
效率的保護系統。
不同目、科之間的空間相關性
不同物種其生態需求不同,對環境改變的反應也不同(Lawton et al.,
1998)。在研究區域內的不同目鳥類間,雁形目與其他各目多為負相關。這應
該是因為雁形目鳥類多繁殖於溫帶地區,喜好涼冷的氣候與草原的棲地有
關。而其他各目的鳥類,往往在熱帶地區繁殖種類數量較高,趨勢正好與雁
形目相反。而在燕雀目中,百靈科、文鳥科以及雀科與其他燕雀目內各科的
科豐富度相關性較為不一致,原因也是這些科的鳥種大多偏好氣候乾冷的溫
帶地區環境,以致於熱點分佈地區和其他燕雀目各科大為不同。由本研究對
東亞地區鳥綱動物(Aves)之分析,可以看出隨著階層越高,不同分類群間
物種豐富度的空間相關性也越高。這應該是由於分類階層越高其所包含之物
種也就越高,大幅降低物種間在棲地需求與分佈範圍上的差異,而呈現出一
個巨觀的趨勢。若分類階層越低,則越有可能受到物種獨特的棲地需求影響,
使得整體趨勢越為紛歧。
25
屬豐富度與科豐富度
使用較高的分類階層取代物種,能使物種的辨識工作進行更為簡單快速
而節省人力物力,同時所取得資料也有良好的可靠性(Cleary, 2004)。但研
究指出分類階層越高,其豐富度和物種豐富度間的相關性會越低(Balmford et
al., 2000; la Ferla et al., 2002)。然而在本研究中,鳥類屬豐富度和種豐富度呈
現相當高的相關性;因此在缺乏完整的種類資料時,或是物種辨識有其困難
時,可以考慮以屬豐富度來代替種豐富度(Harcourt, 2000)。而科豐富度在
本研究中雖與種豐富度也呈現顯著的正相關,但其相關度較低,因此以屬豐
富度來替代種豐富度會較為適合(Balmford et al., 2000)。
尺度
在生態學中,尺度是很重要的議題(Shmida and Wilson, 1985; Wines, 1989;
Levin, 1992)。成功的熱點分析取決於尺度的選擇及考量(Curnutt et al., 1994;
Bobson et al., 1997)。在空間尺度上,以不同大小面積的地區相互比較生物多
樣性將會有誤導性的結果,小面積的相對多樣性將會被高估(Brummitt and
Lughadha, 2003);因此本研究中選擇同樣尺度標準化做比較將會有較為客觀
的結果(Stockwell and Peterson, 2003)。
雖然最適切的空間尺度沒有一定依據(Howard et al., 1998),但大部分分
析結果的呈現和解釋都會受到尺度選擇的影響(Prendergast et al., 1993; Allen
et al., 1997; Howard et al., 1998; Lawton et al., 1998; Bonn et al., 2002)。以不同
物種類別而言,在國家尺度下為狹佈種,在全球尺度下可能就不是狹佈種;
同樣地,國家尺度下為瀕危種,在全球尺度下,可能族群數量龐大(Bonn et
al., 2002)。狹佈種和瀕危種的資料無法由全球尺度直接轉換為國家尺度
(Gärdenfors, 2001);在轉換的過程中,窄尺度和廣尺度不同定義的資料無法
26
進行類比式的彼此轉換。
此外,窄尺度保育計畫有其不可取代的價值;部分較破碎、面積狹小的
重要棲地,會在廣尺度的分析中被忽略而無法呈現。所以在棲地異質性高的
地方,需要良好解析能力的尺度(窄尺度)(Allen et al., 2001; van Rensburg et
al., 2002)。但相對而言,廣尺度能夠掌握不同地景的特性,這是比較小的窄
尺度沒辦法作到的(Fuller et al., 1998)。本研究中,選取的面積範圍越大,
不同類別熱點間重複也越高;另一方面而言,熱點選取範圍越大,保護這些
熱點所需投注的成本也會越高。因此在社會、政策及經濟條件影響下,設計
規劃保護區的大小也是值得慎重考慮的課題(Kunin, 1997; Shafer, 2001;
Olson et al., 2002; Parks and Harcourt, 2002; Kati et al., 2004)。
保育-誰為優先
如何選擇保護區並讓其發揮功效,是對經營管理者的一大考驗。設計一
個能夠長久維持生物多樣性的保護區時,會牽涉到許多複雜的因子在內
(Cabeza and Moilanen, 2001)。僅以不同選擇對象並就實務操作的角度而
言,在選擇生物多樣性熱點時,狹佈種及瀕危種較容易吸引社會大眾的注意,
而有能更多的基金挹注生物多樣性的保護工作(Willams et al., 2000)。但如
同本研究之結果,由瀕危種以及狹佈種所挑選的熱點網格,並不能夠代表所
有物種分佈的情形( Bonn et al., 2002)。在有特定之保育物種目標時,可依
個別之目標進行該物種之保護區劃設(Virolainen et al., 2000)。但若沒有特定
之保育目標時,擁有較多瀕危種以及具有重要生態特色之狹佈種地區,應納
入生物多樣性熱點之優先保育目標(Ceballos et al.,1998)。瀕危種由於受威脅
程度較大,因此應當被列入優先考慮。此外,分佈範圍狹小物種也較易受到
滅絕的威脅(Lawton, 1993; Dobson et al., 1997),因此擁有重要狹佈種之地區
27
也應被列為優先保育的地區。最後,在劃設保護區時,若能利用互補法來選
擇熱點,應能使保護區之保育效果更加具有效率。
28
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表 1. 不同生物多樣性值間之相關程度(T:所有物種;E5:繁殖範圍為五萬
平方公里以下狹佈種種數;E10:繁殖範圍在十萬平方公里以下狹佈種種數;
E20:繁殖範圍在二十萬平方公里以下狹佈種種數;R:未加權之瀕危種豐富
度;RC:瀕危種依受威脅程度加權總值;RP:瀕危種依族群大小加權總值)
r2 T E5 E10 E20 R RC RP
RP 0.422* 0.021* 0.032* 0.037* 0.988* 0.997* -
RC 0.447* 0.022* 0.034* 0.038* 0.978* -
R 0.378* 0.008* 0.028* 0.032* -
E20 0.006* 0.651* 0.716* -
E10 0.009* 0.950* -
E5 0.005* -
T -
* Correlation is significant at the 0.01 level (2-tailed)