!"#$%&'()*+,&-./01+)*&2 3456
School of Forestry and Resource Conservation College of Bioresource and Agriculture
National Taiwan University Master Thesis
7$./89:;<=>?@ABCDEF Leaf Margins and Their Relationships with Annual Temperature and Precipitation in Forests of Taiwan
GH I Chieh Tien
JKLMNOPQ R4 Advisor: Tzung-Su Ding, Ph.D.
STU! 100 > 6 V
June, 2011
誌謝
總不會忘記，那稀疏而華美的景象 當陽光從樹梢間下墜，金暖碎落在肩 我貪婪地用盛夏的翠綠，灌溉心裡的菩提
老實說，我本來想研究台灣的懸鉤子哪一種比較適合園藝化（也就哪一種好 種又好吃），因為手工覆盆子果醬實在太美味了！當然，這個只有衝動無法行動 的提議很快就在各種考量下被駁回（笑）。於是我退而求其次，選擇了跟漂亮的 鋸齒葉有關的研究主題，這個研究說大膽還真有點大膽，因為台灣好像還沒人嘗 試過呢。幸好，幾番小波折後，終究把論文順利完成。
真的，這篇論文能夠完成，要感謝很多很多人。
首先要謝謝我的指導教授丁宗蘇老師，謝謝老師願意接納我這個在他出國進 修前兩星期才貿然跑去請老師收留，而且還不會認鳥的天兵學生；謝謝老師在我 兩年的研究生涯中持續給予各方面的關注，也給了很大的自由，讓我能從不同角 度去發展、解讀自己的研究主題；謝謝老師，在最後要口試那段大家都焦頭爛額 的時期，花心力幫我仔細找出論文很多大大小小的謬誤，還幫我請到了非常專業 的口試委員們，如果不是丁老師，我想論文一定沒辦法這麼順利問世。
在口試本完成以及口試之際，承蒙擔任口試委員的謝長富老師、鹿兒陽老師、
江智民老師，以及林宜靜老師，仔細閱讀我的論文，給予鼓勵之外，也一一提點 了邏輯、圖表、統計，甚至是引用文獻的錯誤，並提出了非常專業的修改及發展 面向的建議。
謝謝401的夥伴們，咪挺時總不吝於給我建議或肯定，特別是阿霞、佩蓉、柯 柯、勁廷、大利、Devil、馥蔓，還肩負了聽我吐槽或吐槽我的艱鉅任務，辛苦你 們了，願還沒畢業的都能高分通過英檢，順利在年限內畢業！已畢業的職場或學
術生涯順遂！也要謝謝助理張美姊，讓我總能迅速完成申請或請領經費，真的，
研究室有這樣專業的助理很幸福耶。
好，我要開始感謝我的親朋好友了，哈哈。謝謝文宜、小隻、章璿、菜菜、
柏因、狗狗、小叮噹、羅小鈞、洪志祐、王宇廷、國王、安琪、毓芩、讓我跟出 差的關老師研究室、咖啡很好喝而且會聽我抱怨的Peloso...還有我可能忘記的一些 朋友們，謝謝你們在這兩年或更長久的時間中給予我研究的各項建議跟幫助，還 有最重要的精神支持及鼓勵，謝謝！
當然，說到幫助，不能不提建融、政道跟建丞，非常非常謝謝你們總是無償 而樂意幫我解決各種資料、統計，甚或是情緒上的困擾，不斷分享自己的經驗安 慰我，跟我一起買醉吃鹹酥雞，下海幫我解決失心瘋買的大堆甜點，讓我不用怕 只胖自己（真是太墮落了），就這樣陪我度過了人生最大的學業考驗。沒有你們，
這篇論文絕對生不出來，能遇見你們真是我上輩子修來的福氣。
說到福氣，一定要謝謝我親愛的家人，我一定是燒過很多好香才有這麼棒的 父母跟弟弟！謝謝我最親愛的爸爸、媽媽，還有天生搞笑咖，帶給我很多歡笑的 田源、田川。很感謝很感謝，感謝上蒼給我如此開朗開明的父母，不管我選哪條 路，只要正正當當就支持到底，讓我能無後顧之憂地完成學業，還總是安慰我說，
人生跟有沒有完成論文沒關係，反正人有很多方法可以養活自己！讓我就算面臨 低潮也知道自己不孤單，撞個幾下牛角尖就馬上潰散，哈哈。謝謝田源，在我抱 怨的時候，還能找出自己更悲慘的遭遇讓我覺得自己其實沒那麼衰（但是這種安 慰方式很淒涼耶）；謝謝田川，努力說笑話安慰我，小時候照顧你真沒有白費！
另外，也要謝謝人在台北，經常給予我生活補給還有打氣的外婆一大家子，謝謝 你們！
然後，謝謝林建融，真的，有你真好。一路走來，你是我最親密的戰友、夥 伴、後援，甚至有時還幫我打前鋒。雖然駁回研究懸鉤子提議的人是你，但是你 也很體貼地和我一起想了另一個我有興趣的研究主題，然後照應我所有需求，不
停地鞭策（？）、鼓勵、陪伴我完成它。謝謝你，因為有你，我才沒有逃避該面 對的事物；謝謝你，在你了解我所有的歡笑、淚水跟醜態後，還願意和我一起面 對。回應你謝誌裡的那句，我願意，也請你在未來的日子跟我攜手同行ɴ很公平 吧，這樣以後如果要塗改，我們兩個人都要進圖書館一本一本拿出來，哈哈哈。
最後，謝謝包容我、成就我的任性的這個世界，希望即將畢業的我也能成為 推動社會的有用齒輪之一。雖然自己的研究不用出差，可是幾次當幫手的經歷會 是我一輩子都無法忘懷的記憶。感謝大地的恩賜，予我足夠的飲食，並讓我有幸 體會到自然的美。
一直記得的是
風輕柔吹撫，枝枒搖曳婆娑 天 那麼藍
而我跟鳥兒一起在樹的頂端
!"#$
!"#$
!"#$
!"#$
!!!""#$%"&#"$'!()%*+&#&'#$,(!")*
+,-./%+)&01#$!!2')-.34(5#'6/0123+4 5266)78#%79:89&844;:<;!=*<)=>%*+?%>+?
@?@>+AB@A@>+,-)CD'45B-B;./%0<#!'!(
)%*+B>+#CD!")1EFG*<EH?2**<EHAB<FEH I#!!!"GJ)GJHIJ(1)G*<EHI)'!()%*+KLMHN
#&'K'!()OP>+LMQNHN&')R&'H1O3)4,P,(
CDGJS:QOQ/(2)RT5EH#!!!"*HI'!()%*+S#&
'/TUG*<EHV)W,*<XUYZ)!V[O\2*)]WCD6&
&'H1O3/AXS*<^72*'!(8A9Y_Z`a):'!(N[
XS*<\QU]^#_`)abbA15;9:*<cXS*<dXUc0.5 hae)CD!"GJf_,dg/(3)eA"$!VMfgh-.&<FEH, -)-!!!"GJih,QijXS*<)L;!!!"<8k=l0,-) mn!""$o)jVk>*<Op#`a/(4)W,45q?r#<'34)
%"stuv)45REH!!!"CD6lHNh,mn-.)789:/(5) w,-Op#xy<n)R!<#'!()%*+*NHN#&')ox@?
z@?pz<#CD6{(HNZ,zy<Bx<)TU!<*<oOpB7 A!V|}B!"GJ)qr#<'Ch~DCs!"/>AE)45
=#%"!'!()%*+B>+LM#'6q12t)8u _v!
#x!!!"&&',/
Abstract
Leaf-margin analysis (LMA) is an univariate regression analysis based on the positive correlation between the ratio of woody dicotyledonous species with entire leaf margins and mean annual temperature, which is often applied to estimate paleo-climate. I used data of 844 plots from!26 native vegetation investigation reports in Taiwan and four climatic variables, mean annual temperature (MAT), annual precipitation, and precipitation of coldest and warmest season, to analyze the ratio of entire leave species (REL) and its correlation with MAT and precipitation in Taiwan and its six biogeoclimatic regions at habitat, mean habitat, and regional scales. The results indicate that: (1) at habitat scale, REL had significantly positive correlation with MAT and some precipitation factors, but the correlations with precipitation factors were much weaker. (2) REL was positively correlated with MAT at all scales. At habitat scale, because the plots were with small area and uneven distribution, REL of plots showed great variation and had weak correlation with MAT. Compiling data from multiple plots, preferably at least 15 plots (about 0.5 ha in total area), could yield consistent results. (3) The LMA regression model from larger multiple plots data was similar with regional scale LMA, indicating that using extreme distribution data could be a reliable data source for LMA. Therefore, using specimen information could compensate the shortcomings caused by insufficient and uneven sampling. (4) LMA regression models in this study all significantly differed from results in other countries. This may be due to the unique ecological environment and plant dispersal history in Taiwan. (5) At habitat scale, MAT was significantly positive correlated with REL in all biogeoclimatic regions.
But the Northeast, Northwest and Central-west regions had significantly lower slopes of regression than Southwest and East regions. Due to uneven and limited
plots in each region, further studies are needed to examine the detail differences among the biogeoclimatic regions. In conclusion, LMA has high potential in Taiwan and results will fill the current information void in East Asia.
Keywords: Regression analysis, temperature gradient, altitude, toothed leaves, Taiwan
%&
%&
%&
%&
!"#$
!"#$
!"#$
!"#$ ... I ABSTRACT ... II
%&
%&
%&
%& ... IV '&
'&
'&
'& ... VII (&
(&
(&
(& ... VIII )
) )
)****+,+,+,+, ... 1
s?!!!"FG ... 1
?!!!")!&!" ... 2
?!!!"&12_u ... 3
1. +!B%*+#,hH ... 3
2. %"./B!#&I ... 4
3. "$o(AB*<XU# ... 4
4. wJ,-#./!V*H ... 5
?!!!"&&'</k ... 5
1. xy!)A!I#'6 ... 5
2. xy!)%KL(B>+#'6 ... 6
?%+&n#,L4!!!"# ... 7
B?45#!!!"12t ... 8
1. 45=#!'!()34>+#'6 ... 9
2. 45=OjT5EH#!'!()34%+#'6 ... 9
3. 45=OjT5EH#!!!"GJ)mn12&EF ... 10
4. 45R./%0<#*<EH!!!" ... 10
--
--****./012./012./012./012 ... 11
s?12<F ... 11
?12z{ ... 12
1. ,-o| ... 12
2. ,-!" ... 13
3 3 3 3****45454545 ... 18
s?*<EHI'!()34LM&'M ... 18
1. >*< ... 18
2. x@< ... 18
3. z@< ... 19
4. pz< ... 19
5. zy< ... 19
6. x<GJ ... 20
?*<EH?2**<EHAB<FEH&!!!" ... 20
1. ! *<EH ... 20
2. !-n 2**<EH ... 21
3. " <FEH&!!!"ICDGJ ... 22
?*<ij4!!!"# ... 22
?45)mn LMA 12&EF ... 23
?45*<EH&%0!<!!!"EF ... 23
66 66****78787878 ... 25
s?45=&*<EH!'!()34>+#'6 ... 25
?45=OjT5EH#!!!"GJBCD6&ChEF ... 25
1. !!!"CD6#k}tCh ... 25
2. REH#!!!"CD6Ch ... 26
3. <FEH#!!!"GJAB<+8} ... 27
?45=OjT5EH#!!!")mn12&EF ... 28
?45R./%0<#*<EH!!!" ... 29
99 99****48484848 ... 31
: : : :****3;"<3;"<3;"<3;"< ... 32
=> => => =>****?@A>?@A>?@A>?@A>(*(BCDEFGH(BCDEFGH(BCDEFGH(BCDEFGHIIII ... 61
'&
'&
'&
'&
~ 1?*<!VAB45./%0!<~ ... 40
~ 2?12~ ... 41
~ 3?%*+,|~………. ... 49
~ 4?%>+,|~……….. ... 42
~ 5?@?@>+……….. ... 49
~ 6?@A@>+……….. ... 42
~ 7?*<EH&>*<!'!()%*+,|~ ... 43
~ 8?2*$*<EH&>*<!'!()%*+,|~ ... 44!
~ 9?2*N*<EH&>*<!'!()%*+,|~ ... 45!
~ 10?<FEH&M!0%""$o[%*+,|~ ... 46!
~ 11?'!([*<;oBT5EH&,| ~ ... 47!
~ 12?*<EH&x@<!'!()%*+,|~ ... 48!
~ 13?*<EH&z@<!'!()%*+,|~ ... 49!
~ 14?*<EH&pz<!'!()%*+,|~ ... 50!
~ 15?*<EH&zy<!'!()%*+,|~ ... 51!
~ 16?*<EH&xy<!'!()%*+,|~ ... 52!
~ 17?*<EH&x<!'!()%*+,|~ ... 53!
(&
(&
(&
(&
8 1?9:¡/ ... 54
8 2?>*<&*<'!()34LM&&'!" ... 55
8 3?>*<¢CD&,Ll£|Mo) r2 ... 55
8 4?x@<&*<'!()34LM&&'!" ... 55
8 5?x@<¢CD&,Ll£|Mo) r2 ... 55
8 6?z@<&*<'!()34LM&&'!" ... 56
8 7?z@<¢CD&,Ll£|Mo) r2 ... 56
8 8?pz<&*<'!()34LM&&'!" ... 56
8 9?pz<¢CD&,Ll£|Mo) r2 ... 56
8 10?zy<&*<'!()34LM&&'!" ... 57
8 11?zy<¢CD&,Ll£|Mo) r2 ... 57
8 12?x<&*<'!()34LM&&'!" ... 57
8 13?x<¢CD&,Ll£|Mo) r2 ... 57
8 14?R%*+O&*<o ... 57
8 15?¤+OOj*<ij&¥*XU2*g/l£C ... 58
8 16?Oj*<ijP LMA CD6)XS*< !-t CD6ChQgGJ ... 58
8 17?Oj<F# LMA CD¦o6 ... 59
8 18?45>*<#*<EH?2*$*<EHAB2*N*<EH& LMA CD6{()mn12&ChEF ... 59
8 19?45#*<EH LMA CDR6¡/ ... 60
8 20?45*<EHIR%0!< LMA CD6§¨){(Qg& p-value ... 60
8 21?OjT5EH#!!!"{©ªEF ... 60
) ) )
)* * * *+, +, +, +,
34"%"2'#LM)Lc<"« #¬|)¢®¯
i{°!3+±.,G#</S<&n²/L;s³c%"#´TOµ)¶
!·)"*#34¸/U¹#!&!"dAnalysis of Leaf Morphologye)
"ºV2'#!)^%"!·#QZ?!»?!2?!!²AB¼.¤
¤!O2'?½)1W¾)34#4<'MB&'j²dChaloner and Creber, 1990e)-GJ8<+,-./%"U¿#/%034&"#$<"¬|#
dWolfe, 1993e/;!"N$)s"%0)!&P,(!"À dClimate-Leaf Analysis Multivariate Program, CLAMPe)UXsFÁÂU*c
U+#z{)Ã"!!!"dLeaf Margin Analysis, LMAe/AIÄÅLMA#1 2ÆÇABG45.<#7t/
JJ
JJ****KFLMNOKFLMNOKFLMNOKFLMNO
%"#!!8È!cxy!dtoothed margine)'!dentire margine/
G;)xydteethe#gÉ"!!LÊY|0#Ë)U-HZ,!!
»¨Ì#1/4)L;OZÍη dlobee):ZÏηE#xydAsh et al., 1999;
Royer et al., 2005e/W,xy²!!)'!²!!O:RN-<#34(5) jV)Ð$%"[*Ñs$!!2')m%"ÒdBailey and Sinnott, 1915, 1916eGÓÔ;?IÂ7Ñ)s;AM%"c©`#%7p)!!
"'!²#E¶)}ÕS<<Ö.#%0+AH/;)8A«U¹
#"$'!(h-.34#%+)7U¹#LMA—Q×3+0M!%
"&"$'!(dthe ratio of woody dicotyledonous species with entire leaves, RELe)%*+dmean annual temperature, MATe##&')W¾8AØÙ'!
(,--.34%+#$,(CD¦o/
\Ú#'!()"Gs;%*+QW&¤?M%"Û©`#=²
\)Ð"$Ü^s)GUN"$p'!!2#"$UÛ#E¶/¶´*<
#%*+cÝc17°C)N100$M!0%")-p25$"'!!2)]*<
#'!("25 %/
cÞkNß<"#¬|Biß34#,t)#à.á./%+S/
<"*^#âa/!!!"&UAM!0%"c)"Lc%"
Gã<ØäUpNFP|å,-dWilf, 1997; Wilf et al., 1998e);)AÑ
<%"W¾#LMAGJ)Â}æç%"&#<F&!·|å#'!()CU -.]<F#/%+/Wolfed1979eèéx#=)78%*+)Ñ
<M!0%"'!(&5#ê89GJ/]wJ#%"*0h,+!°
p#%"&)1oë8}O ¡ì@M_+#*</Wing and Greenwood d1993e èéWolfed1979e#GJ)78ís(%*+)!'!(#sîszI CD¦o)6p,(c'!(dREL)i$µ)0≤REL 1≤ e)º,(c%*
+dMAT)$µJºCe)R6IJ
MAT=30.6 REL+1.141×
61-1 P
P P
P****KFLM0KQLMKFLM0KQLMKFLM0KQLMKFLM0KQLM
3+ÑØ%"&W¾!O?½)34ïo5#!"j²)Â8ØÙ|å ,--]/%0/TU)R$34LMl8}%"#<&<`2)¶
+HvN+!8PI4%"!·#>+L(N^Q)!·#+HAB ðaL(¢OP¥>,%"#2ñ)QZ)T34&%"²'?½[L;Sc
Ò#!"4ò/
L;)w_Z#LMAn)QNÒ3+-.!O?½h-]34ïo— Q×U¹#CLAMPdWolfe, 1993e/T)!·8X#£¼ó?!KGô?%
õ2ñ¤[4%"#ðaL(N^):|å,-O|ö(EZ¤÷?½) L;G#$!O?½)34%+#'6V)v"A!!2'?!ø?!ù2ñ
¤¤zÂÓÔ#nO?½c!",L)AÂ<+,/%+#-.dWing and
Greenwood, 1993e/
bTCLAMP4%"2')±.34##$Fc'X):GúDEFLMA
ûüýP/c#$CLAMP)LMAG%+-.#£àHE"þNHNCh)Wilf d1997e¥Xÿ;!Q"#%"&,-CDwJ)ZÏ#;$O=B;
+O=)N^1445;"$/12<F%&'()*H55°)ZÏ;¦b1 2#y!=)G<@N+b>+#,-§./.OÃ0w/]wJ@
1PGJNJ(1)G!O2¼p)%+#2@S<G!!2'EK(2) nC!!An#2¼?½1cdHe3-.%+#f£HK(3)3+$,(!"{
}!"#*0"$o"3+P,(!"{#bb!K(4)Ñ"P,(!"{#w J<*N#$1g[M_+#%0)TU--.4òÃP"h%U¨N& #/
M%"&)G%+An#34LMChFQ#`2I)3+P,(#CLAMP ,--ö/%+8}i|j©/L;Wilf d1997ec)GQPo`ÇI)- .%*+V3+FÁ|#LAME+CLAMPkc'/
R R R
R****KFLMSTUKFLMSTUKFLMSTUKFLMSTU+V+V+V+V
!·)%0#'68}[V5U,|):(-)#VØ)l*
+dCretaceouse, )M%"|åUu#¸)\b)m-jµ.NH
&#£àHdTaylor and Hickey, 1996e)L;LMAG/%+-.E}N!/
#<+/O-12(5[N(}H)+&D0w#(50ZAI!ª/
1. +!B%*+#,hH
CDLMA#12<F)a9Y+&M_+#`Ç)o%*+)sgH
#,|/0w34(5FcMf#*<)"Lc%*+)!!?½&5#&') 4<@#$&./#%0\c1«dWing and Greenwood, 1993e)UA*<
<ë8}2p l"$#<DK*<%*+ACO}YZ)Ã"34j² á.#£àHI>/Dolph and Dilcherd1979eBDolphd1984e!nGm
Carolinas5)Indiana5CD#!!!"12)LLMA6ö)©q#+HC¨cG
±2°Co)U_ZwJ<%&&%*+,|e!nc6°C)3°C78))!!!"j
²#¬C'(ChY9)Âi{PN®É#GJdWilf, 1997e/
2. %"./B!#&I
ALMA-./%+V)W¾CDj²#%"&p:-ö#<F;ac&9
&#%0./<)1@N9#%"!7)þÃ-.i|j©dHinojosa et al., 2010e/Martinetto et al.d2007e3+ÉQ3z@O#%"|åCD!!
!"):-.].Gpã¯< dMessiniane#%+/;wJ¥XÉQ3Alba
=>9<F?#980;%"|å,-)æC#;,-hx?@!AB!
#!O2¼!"j²)TUA6ö#GJOB)@12Òï°W¾j²#%
0./<)%"!#&'C±)0wD@Oqk²t#;j²)f1P]
wJ.#/%+.^gG13.5–15.3°C&5)1EqØÙ#<F)ÑN#
!O2¼!"FG8A-.@!/%"&U¿#%*+/Aizen and Ezcurra d2008eÃAÑHy!+O=#%"&)æ+GOj!!!"j²E)GJ 7Ñ)(A@IJ%"<UW¾#!!!"j²6öy!#+O%"&)%*
+E©q+H3.6–10°C/W;qP):-.&%"&*0)j²"þh&
9./<?&Ni#¬|uv)"!!!"G<+VO8KL#°(/
3. "$o(AB*<XU#
(W¾!!!"#%"*0h<"P*IF_#<F)&*<"$o
_)4i{SM!O2¼[34#NN,|)CUj²#£àHdGreenwood, 2007e/;n)W,"$O(P[ß!Q#)%"5GT±.#!Q 1OLÑ2*#O³HKR*XU)R*"$oSZ)!'!E¶#^7S i|ÑHN#,hdWilf, 1997e/´%"&R)Tñ'#!'!E¶
&¤#Ø8*0*rT)?n"´s;UZXUR#*0)-!'!E¶) -U¿#%"&&&9#%"&l¦8}OjUOqØ8IdGentry, 1969;
Dolph, 1970, 1979; Burnham and Johnson, 1994e/
4. wJ,-#./!V*H
AAdams et al.d2008eG@#12c¶).7Ñ@h%#zOmV
.<)-!'!()%*+#&'I)_78#12N&H#Ch/.
-ö)8}W,]<ªÕ.2FWFcP,)TUµ12G,-¥PEFr T)}G,-Op#<ªAoX{1Poé)UA_#12EFOY>qr
#`2/Adams¤c)(wJ,-#./!VO¶*)i{LÑNØ8
I#%*+,|_`/
W W W
W****KFLMSQXYZ[\KFLMSQXYZ[\KFLMSQXYZ[\KFLMSQXYZ[\
bT<+'!()%*+&'#12UGPN)§b _)6v"EF
k!!ñ'#2S&P34#>/ßdRoyer and Wilf, 2006e)
N·scP¸¡JE#C±/
1.xy!)A!I#'6
Roth et al.d1995ec!#xy"!»Zt01V#[\])o;?½F
Y>A!^$#<N_)LUxy)A!I*g[HG/èé12)!y Oµ[*qN?n3uH#%õB`+adhydathodese)jVQO!#xy"
$[N01b!yvf):bcOHN,Z#]YdedHickey and Wolfe, 1975; Canny, 1990; Roth et al., 1995e)·s?½KN=,%"#^fAB%
KL()gC>+#CDdCanny, 1990e/L;A!^$4>\"+
,2SFP#!!XUd¶xy!e)Umn¸¡#!·KU)8^bL!
·hûkãUiS#}(jk)1AkNN(#>+ßlU\"dCanny, 1990; Wolfe, 1993; Roth et al., 1995; Feild et al., 2005e/
Baker-Brosh and Peetd1997eÃCm)xy8}"+OA!^$G<@#
,ndprecociouse?½)L;.3+Ío14#p|o%Klg!·p#
p|o)¹)AQö18$M!0%"cSn!#>+N(/wJGJ7 Ñ)G14$xy%"$pNq$)<@, #>+HNXp,xy&Î
·ºf¿)Q×!yOµNF,n#>+}t)XnB$xy%"B$'
!!%"èN*#`2/[N<@w)!!XUÛ'!XU#E¶S hSZV)OrN!!#<xIHNF3#Ñò/
2.xy!)%KL(B>+#'6
W,%"D>+Dp¶#p|o»)T%4%"Us)Â NO,>+#^I/L;)qt%õuHF3#!y)}uu%"#T
%L()UN=,%"D>+dBaker-Brosh and Peet, 1997e/Canny d1990eÂc)Cm%õ2*!VG!·E)!y#ðaL(<]@3/L xy#·vi²wP!8|7<xdturbulencee)^_%0&y6óz tdboundary layer resistancee)Uu3!yOµ#%KL(dSchuepp, 1993e/
Gw~k!·xy§%+&'#jV)[¢N4xyLc8Ab
Qö#</?½/TQö%"xy</?I#12)"#$v$34I) xy"ss4%"N3#?½/{")QO!#&'wJc)G+!i|Ug [F/?#%034I)xy%"G<@, #ã!}7~ª)E'!%"
kq<©`dRoyer and Wilf, 2006e/-kc)!!"!·#%K
<F)L;GF/?#<F)Vu#$}Op):+!°p<V)xy PØ8!!G'!pÛ#XUSQ?%KL(F3)CUu>+N ()Sc%"#<©`&sdBailey and Sinnott, 1916; Canny, 1990; Chaloner and Creber, 1990; Wolfe, 1993; Baker-Brosh and Peet, 1997; Royer and Wilf, 2006e/
Royer and Wilfd2006e[uOj!!²&>+N(#wJ/
.QöOj%*+.<&%"!!#>+L()wJ #cEFxy)
xy!"$#@<S<)#2"þF/?<F#%"!·Nxtk#!
!)1èéPwJ78T%C±dGas-Exchange Hypothesise/;C±
W¾GBaker-Brosh and Peetd1997e#o¥Pj²dthe carbon acquisition modele E)^¼GQL$!!²#%KL(drate of gas exchangee"þNCh/
. ¥mPennsylvania5pNorth Carolina5.#M!0%"),j*
#+ 34I)QJã<!#>+L(/wJGJcd1eO½'!&xy)
!!G<@, "!·@q</xI#OµKd2eGj*#+ 34I)x y#!!E'!#!!kqNxIKd3eGj*#+ wJ34I):<.F /?#%"NFqxI#!!/;12#*0ZÍ*BA!I^$)<²N
))L;Royer and Wilfd2006ec]wJGJ)!#*I&<² 1iHN'6)8A+,N@I/??:+!°#p%Idmesice +O=/
] ] ]
]****^_S`a^_S`a^_S`a^_S`abcbcbcbcdKFdKFdKFdKFLMaefLMaefLMaefLMaef
èéµ128y)bT-.#34LM[8}!'!():%*+
)z})'!()!u#&''MdWing and Greenwood, 1993e/TU) cë(LÑ!!?½)±.%*+#&''M)CD!!!"¥*V)v"7 8}%*+&n#,LCh/TU)µQPo12"G *<V0wN .^#@&./#<Fh^bChI)b9·%h)fw{NÒ4>+??
34?m?¨+9¤¤LMH& /
Traiser et al.d2005e!"@!25$!O2¼)34ïo#&''M)4
*<mACG400 mAo)19wx¹#<F)A^b-.8}#3 4L.4!&!"#}B/wJGJc)!'!()%*+N@HN##&
')U>+LM%>+(dMean Annual Precipitatione?@#>+¤
dPrecipitation of Driest Monthe¤))'!([NHN#'6):"&'H
_,%*+/-p)%>+)'!(LHN#&')m%>+cº,o)'!(
)-.!O2¼c,o)8AW¾A!&-.%>+#P^ICDj6 dMultiple Linear Regression, MLRe):j6#6ö£H1O8)oi{6ö
%>+G1000 mmAE#.</
Royer et al.d2009eA!$O=c12<)G%*+&9#12<o)
QJ=B¹?+P8PIp<|ݤLM)xy"$!V&&'/GJ7 Ñ)±.%*+V)?+!8PI3#.<&"$'!(HNF_):=
B¹?$MQZ?$MaQj6¤LM)"$'!(¨NHN&'/
g g g
g****hiaKFLMTUjkhiaKFLMTUjkhiaKFLMTUjkhiaKFLMTUjk
!!!""^W/%0?k<'B34,|#/c/O¾}Aizen
and Ezcurrad2008e#128y)Ay$O%"<W¾#j²h6öy#+
O=%*+)-GJàq)©q+H9):j*#,-æC@&..#j
²)6.+HO}~9qr`Ç/W;8q)W¾Oj%0<?Oj%"&)
\&OjEH#!!!"j²)1eC¥*AB!"#z{)4,!!&!&!
"GR<F5#hjfkNk²}t/
µLMA#12PI,!AB@!CD)b,x<)wWing and
Greenwoodd1993eèéx#%78978LMAj²&n) _ebo
Yang et al.d2007e?Su et al.d2010e¤)"3+pmQ"#%"&)78&
^ãQL&'j²Gpm#<+):45cNÒ¿wzX#12/
45µ¿%"./<6#µ)bTEF&_#LMA12<F)4 5!Q'(#*H,|): ¡#mACwP45#%"&º¢[+H UNHN#,h)jV[N!O³#"$dS/Sud1984be445£!#
;./%0<ABo;<)Oj./%0<#>+@I,|AB@¤#
NUCh/G;ù¥I)45#LMA12O:8+mnD#!"{)JE45
=#!'!()34LM&''M)[8AºV./%0<<F5#ChI EF/nE45#;!=!VGO+?./#<Fd¦§)2008e) Y>_12#p%I=?½)U}¡>*<,-)¿wOj#XS*<i j)hQJv$EH#,-f>À45s*.2,| ¡#34CD!!!
"/45)78¨P#%7129:)}=>k"HF¤©#34,-)4 5#!!!"12GJ);8u _v!#x<&LMA&',/
L;)012ª«45.<#%7899:)¥X*<"$AB«l¤
34,)#$45=#!'!()34>+?34%*+AB*<T5EH
&'6I/AIc012&s!" lJ 1.45=#!'!()34>+#'6
_12))'!(N@&'#34LM"%*+):12[HI)
>+)'!(QHGHN&'/c#234>+}þ¬%*+4'!(k
#T)1JE45"þ)-.12<N#GJ)012[QLG*<EH dhabitat scaleeV)w%*+&n)%>+?@?@>+?@A@>+4'!(
#8}/°45./%0<5#>+ChI);~ªwQJ45#¡K _`n)QLÑ%0!<#GJ)A#$%0<5#8}Ch/
2.45=OjT5EH#!'!()34%+#'6 2a.RT5EH#!!!"CD6
W,,-#T5EHB*<XUQZ'!(#^7GJ)0 12cÞk[T5EHU,h#!!!"_`)>R./%0<#*¼) CD*<EH?2**<EHdZÏ2*$*<EH?2*P*<EHe) AB<FEHdregional scalee#!!!"/-p2**<EH/<FEH"
º4*<®%*+!O)r^7Ðs%*+Oo#2*'!(/'!(/
^72**<EH#'!()"cÞk¯g%*+O#*<ij)'
!(,|&&'/L;012[ß %*+OoOj;o#*<cXS
*<)^7RXS*<&'!()LÑ'!([*<ij#,|_`)BO jXS*<#!!!"GJ/;n)Q4^<FEHI)R%*+!Oo
#"$o)ALÑ45.<Oj!!²#"$o([%*+#qr,|/
2b.REH#!!!"CD6ChQg
0124QgRT5EH#!!!"CD6#{(5B§¨5#C h)AÞkXS*<#&4QZ4!!!"#/
3.45=OjT5EH#!!!"GJ)mn12&EF
W,Oj.<#!!!"GJ*NCh)0124EF45#!!!"C D6&{()mn12#Ch)A#$45!!!"GJ#°?I)1uch G45CD!!!"#&'Ws/
4.45R./%0<#*<EH!!!"
cEF45R./%0<#!!!"_`&hj)012wCDR!%
0!<#*<!!!"n)[CD!<!!!"GJ&EF)A#$<5C hUØ8#<'&%0®É/
- - -
-* * * *./012 ./012 ./012 ./012
J J J
J****TUlmTUlmTUlmTUlm
012A450±#%789,-CD!"K450±csU²m
±)cµ¿@*25.3Hb@*20.9H)xz124.6Hbxz119.3H&5)±²
@zcpmm)xXY2-)z³45m´)@Q"&8K@CD%S 450±pyO)[µÉ?¶·ABÁ¸¹º»9)L;º./µh¼) 45jVZÍ$O)+O$%0/450±XUcc36000 km2)-p31 %
cm100 mAI&2.)45 %cm500 mAE#·..2/·.XUp
N70 %#m½1000 m)Um½2000 m#ÃÛN·.XU#26 %)
mG3000 mAE#.2Û·.NXU2.8 %)-pA¶··»@3)cc
3950 m/0±·êPcy@t))±¾2D)-pp¿·»ÀÁ'±)S
c45#x?zOÂ!+Ã/45ì@PQ"?%ÄUNx@@¤Å Æ)c@OOh_+)OÇ(KÈÉrVNs;78#Ê@)' 4lNß>+KÉ@ÃPzy@¤AB$O_%Ë)"'±s%p
(@3#V/@,&r)@¤Ì#N<!dH)Í(-ÃAW yt@#Îhc/>EZ)45XUbOQ):G>m?.2?·»
t?ÏO.2?-?D@¤ABФ¤L.)qN&P,#<
'34)%0!</
012Uw+&./%0<6gl£ÂSud1984a, 1984b, 1985e/
Sud1985e3+@I+H?%(?ì@(?#@P¤%0,-) AX+<!<)445£!cx@9m<?x@o"<?xO<@ª?xO
<yª?z@9m<?z@o"<?pzO9m<?pzOo"<?zy<?
xy<ABѲ<Ò11;<'%0!<&ÆÓ)U4./%0<Æ!cx@
<dNEe?z@<dNWe?xO<dCEe?pz<dCWe?zy<dSWe Bxy<dSEeÒBQ<w+&d~1e/
P P P
P****TU12TU12TU12TU12
W!!!"&'128y):W¾!!?½)%*+#&');aë8}0 w+!(5#Ch)f}G34F#`ÇI)EF'!!)xy!"$43 4%+#</TUuv9:#Ô6)+Õs 1Os)L;012Ò +26 645.<#%7&'9:dÖq81e)wÐ;<'%0!<lN %7*
<d~1e)oë8} Ù45#;!%7²)rï°9:U»&*<,-) o|c"$×Ø)13+*<«lABArcGISR8!"#34ïo/
!!!"!cQOPJ,-o|),-!"/G,-o|~ª¸S*
<34?*<"$,-AB"$?½,-Òs8!"#×ØK,-!"z X)°45#%789*<Fmn12hPZ)c#$*<XUQZ"þ
'!(#LÑ)012w^7*<EH#'!(n)Q^72**<E HAB<FEH&'!()AÞkG*<EH?2**<EH)AB<FEHI)
"þL¥*#T5EHOjUN&h#!!!"GJ/
1P34LMABRT5EH#'!()rRA-'!()34LMC DCD!")A#$IÙs12 lJ
1. 45=&*<EH!'!()34>+#'6) 2. 45=OjT5EH#!'!()34%+#'6) 3. 45=OjT5EH#!!!"GJ)mn12&EF) 4. 45R./%0<#*<EH!!!"/
Ö¤&12d~2eB12ÚIÖZ/
1. ,-o|
(1) o|2669:#%789,-)¡/*<"$&gI,-
W,R9:#%789,-LÑz6Os)L;i{bCDg(,-#
!");n*<89[8}N l#%"$&%7²)LU012G,-¡
/V)º)wM!0%"Û©`#%7dJÛÜ=²e)rW¾!"+
#*<"$,-/;n)W¾*<"$,-V)1O°"$#o()eÝ
Ñ)þ)r4VÞ?s"$#w+ÞÚ)@ß"$obb10$#*<
CD!")¸¡"$ÞÕq»Õ/
(2) Ý"$!!²&gI?½
012®VFlora of TaiwandLarsen, 1995esp#àZ)´"$G]
#!!?½plÃcserrate?biserrate?serrulate?dentate-serrate?crenulate-serrate
¤áâ)4]"$Lcxy!)àZcã'!&xyä[LcqNxy!)(
åã'!äfLc'!!)1æS"$?½4V8AÂ!"/-p)NT12
cη8})!yN#ç}):012®)78&!!!"12)eqN η?½ÒÜ^c'!)ηENxyÒÃÜ^cxy!!Kü!%")j*®
)7812)AZ!#!!?½cÜ^4ò/
(3) *<34,-
Ü+45Q'J,t12pĺé1960–2001%%òè,-U78&%
0~ódLin et al., 2010e)^ã^7S%*+d~3e?%>+dPrecipitation, Prec)
$µmm)~4e?@?@>+dPrecipitation of Coldest Quarter, PrC)$µmm)
~5eAB@A@>+dPrecipitation of Warmest Quarter, PrW)$µmm)~6e) k"Hc 40m é 40m)r3+*<#«l)«ArcGIS´%+B>+~óR
%0ïo)>+,-ZÍ>+B>ê,-)UA>+ÛQPo/
2. ,-!"
(1) '!(^7 a.*<EH'!(
Rs,-8W¾)3+ Microsoft office ,-ëìK Access 2003 í¡)
®VWilfd1997e#z{^7R89*<#"$'!(d^762-1e)R6IJ
!"#$%&'()
!"$%*+
!",'()
62-1 b.2**<EH'!(
°(45PH,./?rI)%789m#*<PIFmnZ)wP
*<5#ChQ)cÞkOj*<ij"þ489#'!(N)012
®1°Cc<³£!R%*+O)^7%*+O#2*'!(/-p)W,LMA
"A'!(-.<Ö.&%*+)1i{-.-%*+&SDdstandard
deviatione)L;012G<!%*+OV)QO4%*+&SD°Go/
£!%*+)[ßGR%*+Oo n;*<cXS*<)Úc!-n
*<)1^7!-n*<#'!(d¶J[ß n;*<)'!"$oca)N
"$cb)Ã]1P#*<'!(ca/be)^ü¥*1000)1P+n;Z*
<cXS*<#'!(2*g)-pnºÀc2?5?10?15?20?25?30?50? 75)XS*<!nA!-2?!-5?!-10?!-15?!-20?!-25?!-30?!-50?!-75Ú&/
%*+O*<oOpnV)OCD]ng#[ߥ*/
AEUP"cR%*+O#2*P*<EH'!()^7{IJ
= !-n
1000
!"#$%
&'(!")*#$%
!
62-2 TU)n¤,1&tdQ×total)8I]%*+O#UN*<eV)1OD
^ü¥* 8#2*'!(g/n=1VbAR%*+O#*<'!(&nr wA*<;o×8)Úc2*$*<EH'!()^7z{IJ
=
!
!"#$%&'(!")*#$%
!"+
62-3
nctdUN*<oeV)ÃA%*+O#'!"$owAN"$o×8)
^7z{IJ
=!"#$%&
'()*+,-!".
)$%&
62-4 Ojij#XS*<&XU¥*@Qg?@ZgB2*g4Ù8ïV)'
!(,|_`)Ã *<oFP#23.5°C?19.5°C?15.5°CAB11.5°C(%*
+OØ8)A ~8Ñ&/
c.<FEH'!(
W,89*<NH)c¬!Q,&5 )012Cm"$,+HîH E"ïÅ!V)î_Z#%*+!O)ÆR"$G%*+!VE#@_)@
3,-)ðª]"$ÝG;%*+'(oKN!V)U1P%*+O#/Ç"
$o/r4R+HOLcs<F)^7]<F&/Ç"$'!()Úc<FEH '!()1W¾]EH'!()%*+#CD/<FEH#%*+O@_+Ü
^8.5°C)Lc453m.<c7°C78ñcò!=O)M!0%"$
Yb/<FEH#'!(^7z{IJ
=!"#$%&'(
)*+,-./#$0
!"1&'(
62-5 (2) #$>+LM)'!(#'6
1P*<EH&'!()0~ª!"4¤2w%*+An)%>+?@
?@>+AB@A@>+4'!(#&'H/L;ís~ª3+^ìKSPSS 12.0)CD'!(?%*+?%>+?@?@>+?@A@>+&5#óô&
'!"dPearson correlatione)Um'!(cº,o)%*+?%>+?@?
@>+B@A@>+sÒ;34LMmc,o)CD¢CD!"dstepwise regressione)U1PR,o&l£|Mo/#$G*<EHI)x@<?z@<?
pz<?zy<ABxO<¤;./<AB45>*<#>+LM)'!(
&'6/xy<L*<NH)34,|1OQ)i{CDNN!"/
(3) 45O!%0<#RT5EH!!!"EF
0~ª lcÞkXS*<)v$ij)!!!"#GJfOrL*
<ij\QUe,)}Ps;dg#!"GJ/L;445=O!%0<#
*<EH?2**<EH)AB<FEH&%*+)'!(#CDCD)1P
61-1&ICD6)r3+ÉIj6dGeneralized Linear Model, GLMe) EF2*N*<EHd!-te&LMACD6))2**<-.EHB<FEH#
LMACD6G{(B§¨#Ch/CD6#{(&§¨ÈssNHNCh)
×Ø8"Oj#CD/
(5) )mn12<&EF
0~ª#!")¡/45>*<#*<EH?2*$*<EHB2*N
*<EH&LMACDGJ))-.8;12<CDEF/-.12.<!n"J xdWolfe, 1979; Wing and Greenwood, 1993e?@dWolfe, 1993e?!dWilf, 1997e?p@!B0dWilf, 1997e?y!dGregory-Wodzicki, 2000e?
!dGreenwood et al., 2004e?@!dTraiser et al., 2005e?pmdSu et al. 2010e/
O_ª!"}jVEFCD6#{(§§¨)W,i{1P-.12<
#Ö¤89,-&§¨#l£¬dstandard error of estimate, SEe);OPAR9 :pUõdCD6#{(AB{(#l£¬CDöt-test)AÞkRCD6G{
(OP"þNHNCh/Qgtg&^7z{63-1)6pbcCD6{()SE cj6#l£¬J
1 2
2 2
1 2
t= b -b
SE +SE
63-1 RQgU+&WHddegrees of freedom, dfe63-2)6pSEcj6#
l£¬)ncj6&*0oJ
(
12 22)
24 4
1 2
1 2
SE +SE df=
SE SE
n 2 n 2
" #
$ + %
− −
& '
63-2 z63-1B63-2^7)EF{(Ch#HNH/
(6) #$45.<R%0!<#!!!"Ch
0~ª3+^ìKSPSS 12.0)4*<%*+mcº,o)*<'!(m c,o)CD*<EHI)x@./%0*<?z@./%0*<?pz./
%0*<?zy./%0*<)ABxO./%0*<#!!!"dxy./%
0<L*<#34,|1OQ)i{CD!"e)1AGLMCD%0!<&LMA CD6#{(B§¨&ChEF/
3 3 3
3* * * *45 45 45 45
012Ò^844;*<)-px@<187;?z@<156;?pz<118;?z y<268;?xy<76;?x<39;/*<XU100÷600 m2O¤/M!0%
"Ò^562$)ZÍ80;ø)!'!2^310$?!xy2^252$/
J J J
J****nlnlnlnlopopopopqrFs0tucvSXwqrFs0tucvSXwqrFs0tucvSXwqrFs0tucvSXw 1. >*<
>*<V)'!(;34LMKLHN#&')&'H@3Òc%*
+dr=0.50, p<0.01e)@A@>+&dr=0.15, p<0.01e)%>+ídr=0.11, p<0.01e)@?@>+@ùdr=0.07, p<0.05eK-p%*+)%>+LHN&'dr=
-0.19, p<0.01e))@A@>+LHN#&'dr=0.11, p<0.01e))@?@>+¨N HN&'K%>+)@?@>+dr=0.69eB@A@>+dr=0.14eLHN#&' dp<0.01e)@?@>+)@A@>+LHN&'dr=-0.53, p<0.01ed82e/
>*<#¢CD@P Ù;34,L),L ÙÉÀºc%*
+?%>+?@?@>+AB@A@>+)1P;CD6#Nr2gºc0.245? 0.287?0.302?0.329/HI>*<#'!()wA%*+kn)nC>+,- [}HNmn34LM4'!(,h#kH/;,ol Ù#CD6p)
%*+)%>+&l£|Moc#)@?@>+B@A@>+&l£|Moc
d83e/ 2. x@<
x@<#;34LMp)'!(;34LMKLHN#&')&'H
@3Òc%*+dr=0.44, p<0.01e)%>+&dr=0.38, p<0.01e)@?@>+í
dr=0.33, p<0.01e)@A@>+@ùdr=0.17, p<0.05eK-p%*+)@A@>+L HN&'dr= -0.57, p<0.01e))X;>+LM¨NHN&'K;>+LM 5KLHN#&'dp<0.01e)UA%>+)@?@>+#&'H@3dr=0.94e d84e/
x@<#¢CD@P Ù;34,L)ºc%*+AB@A@>+)
;CD6#Nr2gºc0.196?0.456)%>+?@?@>+Ã0w/;, ol Ù#CD6&,Ll£|MoKc#d85e/
3. z@<
z@<#;34LMp)'!()%*+?%>+B@A@>+LHN#
&'dp<0.01e)&'H@3Òc%*+dr=0.32e)%>+&dr=0.29e)@A
@>+ídr=0.22e)@?@>+)'!(¨NHN'6K-p%*+)%>+
dr= -0.38eAB@A@>+dr= -0.49eLHN&'))@?@>+NHN#&
'dr=0.5eK%>+)@A@>+NHN#&'dr=0.93e))@?@>+¨NHN
&'K@?@>+)@A@>+NHN&'dr= -0.25, p<0.01ed86e/ z@<#¢CD@P Ù;34,L)ºc%*+?%>+AB@
?@>+);CD6#Nr2gºc0.105?0.303?0.327)@A@>+Ã0 w/%*+B%>+&l£|Moc#)@?@>+&l£|Mocd87e/
4. pz<
pz<#;34LMp)'!()%*+B@?@>+LHN#&')&
'H@3Òc%*+dr=0.34, p<0.01e)@?@>+&dr=0.19, p<0.05e)%>
+B@A@>+Ã)'!(¨NHN'6K%*+)R>+LMl¨NHN&
'K%>+)@A@>+NHN#&'dr=0.90, p<0.01eK@?@>+)@A@>
+&5cHN&'dr= -0.19, p<0.05ed88e/
pz<#¢CD@P Ù;,L)º Ù%*+)-c@?@>+)
;CD6#Nr2gºc0.113?0.147)%>+B@A@>+Ã0w)¢C Dp#; Ù,o&l£|Moc#d89e/
5. zy<
zy<#;34LMp)'!()%*+dr=0.57eB@A@>+dr=0.37e NHN#&'dp<0.01e))@?@>+LHN&'dr= -0.45, p<0.01e)%>+
)'!(¨NHN&'K-p%*+)%>+dr=-0.38, p<0.01eB@?@>+
dr=-0.85, p<0.01eLHN&')U)@A@>+LHN#&'dr=0.40, p<0.01eK
%>+)@?@>+dr=0.43e?@A@>+dr=0.63e5KcHN#&'dp<0.01eK
@?@>+)@A@>+5cHN&'dr=-0.28, p<0.01ed810e/
zy<#¢CD@P Ù;,L)ºc%*+?%>+AB@A@>
+);CD6#Nr2gºc0.319?0.370?0.394)@?@>+c Ù/- p%*+B%>+&l£|Moc#)@A@>+&l£|Mocd811e/
6. x<GJ
x<#;34LMp)'!()%*+NHN#&'dr=0.68, p<0.01e))
%>+NHN&'dr=-0.35, p<0.05e))X;>+LM¨NHN&'K%*+
)%>+dr=-0.65, p<0.01e?@A@>+NHN&'dr= -0.60, p<0.01eK%>+
)@A@>+NHN#&'dr= 0.86, p<0.01e))@?@>+¨NHN&'K@?
@>+)@A@>+&5NHN&'dr=-0.32, p<0.05ed812e/¢CDV) e Ù%*+c,L)CD6r2gc0.46)l£|Moc#dp<0.01ed813e/ P
P P
P****nlnlnlnlopopopop****xynlopz{lmopSKFLMxynlopz{lmopSKFLMxynlopz{lmopSKFLMxynlopz{lmopSKFLM
AIRñ¦op#MATØ8%*+d°Ce)REL"'!(d0≤REL 1≤ e)
!8I*<EH)!-ndnJ1, 2, 5, 10, 15, 20, 25, 30, 50, 75, totale8I2**<EH)
"8I<FEH)ncXS*<UZÍ#Z*<;o)p cHNH/
1. !*<EH
*<EH&LMAñ¦ocJ
2
MAT=15.1 REL +8.276 r =0.244 <0.001× ! p
64-1 W~78q)45*<EH#"$'!(,h&Q)ZÍ0B100%)úû
#±.%*+cG5–25°C/:G844;*¼p)'!(30%AINüÊo;) '!(AG0.30–0.90#<5)1!VG%*+8–23°C#'(o/
2. !-n2**<EH
(1) !-12*$*<EH&LMAñ¦o
2
MAT=55.7 REL× !-1 16.629 r =0.896 <0.001p
65-1 2*$*<EH#%*+,|'()*<EH&j):""$'!(!Ve c0.45–0.75)'!(#,|<5E*<EHZd~8e/-'!()%*+&5Q
"HN#&'):r2gEF*<EHQýu){([FQ/
(2) !-n2*P*<EH&LMAñ¦o
(a) MAT=64.3 REL× !-2 20.981 r =0.883 <0.0012 p 65-2 (b) MAT=73.0 REL× !-5 25.109 r =0.908 <0.0012 p 65-3 (c) MAT=74.5 REL× !-10 25.538 r =0.917 <0.0012 p 65-4 (d) MAT=66.5 REL× !-15 20.437 r =0.935 <0.0012 p 65-5 (e) MAT=64.4 REL× !-20 19.060 r =0.934 <0.0012 p 65-6 (f) MAT=62.8 REL× !-25 18.019 r =0.933 <0.0012 p 65-7 (g) MAT=64.3 REL× !-30 18.876 r =0.930 <0.0012 p 65-8 (h) MAT=57.6 REL× !-50 14.740 r =0.907 <0.0012 p 65-9 (i) MAT=51.0 REL× !-75 10.522 r =0.780 <0.0012 p 65-10
2*P*<EH#!!!"&_`sW)%*+)'!(&5KcHN#&
')r2gG0.780÷0.935&5)r2g@_ÒcXS*<!-75/
(3) !-t2*N*<EH&LMAñ¦o
2
MAT=63.7 REL× !-t 18.322 r =0.912 <0.001p
65-11 2*N*<EH#%*+,|'(j2*$*<EH)"$'!(!Vcc
0.45–0.67)'!(#,|<5ÝE2*$EHZd~9e/-'!()%*+&5
Q"HN#&') r2gEF*<EH[Qýu/
(4) R%*+O#*<o,|
%*+24÷25°C&5e4;*<)"*<o@b#%*+O)¡KUs)F 3+ABF_+<#*<oFb)%*+20÷21°C5N128;*<)"*<o@
P#%*+Od814e/
3. "<FEH&!!!"ICDGJ
<FEH&LMAñ¦ocJ
2
"
MAT=64.8 REL× 18.607 r =0.951 <0.001p
66-1
<FEH#CDGJF2*N*<EH/<FEH&"$'!(!Vc 0.43–0.68)!V%*+G8–23°C&5/-'!()%*+&5["HN#&') r2"R$T5EHp@3Ò/
G<FEHI)"$o(G%*+15–20°C#'(od&,45#pm
.<Ve@3)%*+½&_,;'("$olFdH#I>)-p'
!!"$#o(@3¼cG%*+20°C)xy!"$o(#@3¼Ãc%*+
13°C78d~10e/A'!(0.5ØÙ64-3)8yG%*+14°C»9)$!
!2#"$E¶&¤/%*+3,;g)'!"$oFP)oo(C¨N[+H
3UþQ#_`K%*+_,;g)ÃAxy"$oFP/
!!
!!""""#$%&'()*+,-.#$%&'()*+,-.#$%&'()*+,-.#$%&'()*+,-.
XS*<ij&jV)%*+S3)'!( 8gS3KUjs%*+Oo) [¥*&*<ijnQ)'!( 8g#,hHZ?og_dg/A 23.5°C?19.5°C?15.5°C?11.5°C&%*+OæS ~)G%*+23.5°CAB
19.5°C78)XS*<ijW1;mn10;*<V)'!( 8g&2*oN
¢>_#_`)XS*<ijQ,15;Z*<&'!(_,´gg)U<F EHV)'!(¢ÝcEK%*+15.5°COGXS*<EHV#'!( 8g ,|%*+23.5°CB19.5°CO):<FEH#'!( 8gOS>K%*
+11.5°CO#'!( 8g,|ÃO_ZË*Niÿd~11e/Oj*<ij
#XU 8g[*<ijmQU¤E¶mn)¥*#,hHÃ[*<ijmn U^bd815e/
Ojij#XS*<)-LMACD6[Z*<o(Uhd816e/2**<
EH#*<ijG10;9:*<AIV)-CD6#§¨)65-11NHNChK 15;AE#9:*<cXS*<dXUcc0.5 haeV)-CD6O½{(
&§¨l§65-11¨NHNCh)8Lcjs(CDKU65-11)66-1[8 Lcjs(CD/
O½T5EH)45#'!()%*+&5KLHN#&'):64-1&r2 F_d816e/464-1?65-1AB65-11CDEF)Ã!;lNHNCh) 8Lc(Oj#CD/
/ / /
/""""01234012340123401234 LMA 56565656777789898989
O½T5EH)45#'!()%*+&5KLHN#&'))-.12.
<#_`sW/464-1?65-1?65-11) _)78#mn12d817eCD EF)Ã45RT5EH#CD)) _U78#12lNHNChd{(C hQgp<0.001ed818e/
: : :
:""""01010101#$;<7=>*$()*+#$;<7=>*$()*+#$;<7=>*$()*+#$;<7=>*$()*+89898989
45R./%0<#*<'!()%*+&5)wxy<An)lNHN#
#&'dp<0.001e):r2gNx<ABzy<F3d819e/x@<*<%*+
G14–22°C#'()'!(,|G0.1–0.9&5)*¼XpG%*+20°C78d~
12e/z@<*<%*+G5–22°C#'()'!(,|G0–0.9&5)*¼QW!
c7)F15–19°C#89<Fd~13e/pz<**<%*+G8–18°C# '()'!(,|G0.3–09&5)G%*+15°C78NFuX#*¼d~14e/ zy<*<%*+G9–24°C#'()'!(,|G0.3–1&5)8¼*¼G+
HB'!(ElNFdH#,|_`d~15eKxy<*<%*+G20–25°C#'
()%*+,|Op)i{"N®É#CDGJdp=0.629ed~16eKx<*<
%*+G11–24°C#'()'!(,|G0.3–1&5)*¼G+HB'!(E#!
Vz@<)QW!c7)%*+19°C78#*¼d~18e/
zEF)R%0!<#CD6{(8A!c7)x@<?z@<?p z<Ò&5#{(¨NHNChdp>0.05e)zy<Bx<&5[¨NHNCh dp>0.05e)-#{(EFKNHNChdp <0.05e)1oAx@-z@-pz<
#{(FZKCD6§¨8A!c7)z@<)pz<&5#§¨¨NHNC hdp>0.05e)zy<)x<5#§¨¨NHNChdp>0.05e)-#EF§¨
KNHNChdp <0.05e/L;>Us)%0!<#!!!"CD68A!c
7)z@<Bpz<#CDGJ8Lcjs(CD)zy<Bx<[8Lcj s(CD)Ux@<#CDGJ)-.!<lOjd820e/
!
!
!
!" " " "#$ #$ #$ #$
?
?
?
?""""01@A7#$;<(B)C2DEFG,HI01@A7#$;<(B)C2DEFG,HI01@A7#$;<(B)C2DEFG,HI01@A7#$;<(B)C2DEFG,HI
012#!"HI)45#>*<&'!()%>+5#&'I)) Traiser et al.d2005e#12GJ&sW/Q×'!()%>+bTG^EN HNI):&'H1O3)U-p#<'®É~TAk)i{é;W¾8#
CD6/45O½!<)þ))'!(#&'@UNsWI#34LM"%*
+)>+LMG!<#8ÑÃTA¼_`/qE)°¾!"R./%0!<
V)¦8}W,!<*<#Ø8IB°¾IvOp)->+LM)'!(5O0 w"CI&')Uqr#&'Ii{W;ày/
¦§d2005ec45$,Holdridged1967e<%O!z&&%H'#
%(dsubhumide)I)pdsemisaturatede&5)1i+#`a/L;012
cOi|G%*+#*^;3#ñÇI)¼&4FZ#>+(,h) '!(,|#dHïG/TU45&>+_`P@¤\dChen et al., 2004e) (}1PT5!VFc*#*0)GH+H,h#_uI)>+@I&<
@>+)'!(#&'[ý"ch8A#$#âa/
J J J
J""""01@AKLMN;<,()*+OPQRST7UV8901@AKLMN;<,()*+OPQRST7UV8901@AKLMN;<,()*+OPQRST7UV8901@AKLMN;<,()*+OPQRST7UV89
>h¼)45#M!0%"G%*+3,14°CVA'!"$FP)S
&)%*+_,14°CAI#±.ÃAxy"$ÛPo);Ñò&Y>Royer and Wilfd2006eu&'!([%*+#6 ,|)Q×xy!"$GO+#_+
<E'!!"$ki|1P<©`):OjT5EH#LMAGJÝNCh/
1.!!!"CD6#k}tCh
45>*<GOjT5EH&LMA#CD6)CD6#r2N&Ch)<
]"*<ij'!(Ø8IUWdGertry, 1969; Dolph, 1970,1979; Burnham and Johnson, 1994e/
*<EH#LMACD6&r21O3)-ö"LEF-.mÌ)012#,-
G&34I*<5#"$,hFQ)b+*<#%*+)'!(,-CD V)r2ÂTAuu/:L"45.2P·)89VTA1PQ'(#*¼*<)%
789+,AP*<{dmultiple plot methodeCD)P*<{"G.2)%"d SFc*¼#.zmZ<)Z<rX>cs8!"#XS*<dGrossman et al., 1998e/45#XS*<QZPG400 m2÷600 m2)*<XU~OpAw'!(N Ø8I)]W-ß\<#.}B!"GJK-.mÌ#LMA*<'(*G1 ha AE)\b"A0.5°×0.5°z*H/Ôcs$µdWolfe, 1979; Wing and Greenwood, 1993; Wolfe, 1993; Wilf, 1997; Gregory-Wodzicki, 2000; Greenwood et al., 2004;
Traiser et al., 2005; Su et al. 2010e)}1PFcdg#'!(,-)>_.#}
B)UuuCD6#k}t/Adams et al.d2008ec*<#*H!
!!"#GJ)0!"&,-hN #I#%789,-)*<i|L7Añ
!V)L;T5*H#`aOiKL/
A!"GJh±)3+2**<EHd!-nEHe#,-hCDLMA)8A e3*<EHd!EHe,-#Op)k>*<!VO*#`a)01L*<
ijZU\<#.)wOj+HO#'!(,-}AFdg#z6LÑ)CD 6#k}tÂ}HNuu/
O2**<EH#*<ij[D)sgQZ)*<#'!(fdg/0 12#GJHI)45'!(#,hH[*<ijnQU^b)r_>_
b´gg)Y>Wilfd1997ec*<XU'!(#±{/;_`ÝLSJ
²!V)<"W,45#xy!"$Fb)L;LÑxy!"$o#Ø8IU D&@Z*<XU)F'!!"$UDkQd23¹?Í45)1989e)]WZ
*<89V|_.!xy"$o)UPF3&'!(K;Ñò6-G'!"$
)xy"$o&CSÎr#<FÔndH)wPCD6Oe§¨N7C){
([8}NUCh/
2.REH#!!!"CD6Ch
O°Ø8I@_#*<EH)e¥!-nEHdn=1?2?5?10?15?20?25? 30?50?75e)2*N*<EHd!-tEHeCDEF)-LMACD6[NHN Ch/A,-?½h¼)'!([*<ij\QUÝLSJ²!V)L;*<ij
ZV8}3.3+<#'!()]W*<Z,!-15V#LMACD6&§¨
)Q*<#GJN7C/012¥ijG!-15AE#!-n*<CD!"V)- '!(Fcdg)LMACD6)2*N*<EH&5iHNCh)8Lcjs(
CD/gPî#")!-15#*<ijQc"0.5 ha)Õcmn)78& LMA 12UÜÏ&@Z*<XUdWilf, 1997e/
-p!-75EH#CD6r2dH7_)Ã-ö"LR%*+O#*<oPG 75;AI)[ß 75;*<V)}ÏÙ!"#%*+Oe;)*¼YbÂw CD6|\<7C);OP&}«8¡!"{&mn,-eC/
3.<FEH#!!!"GJAB<+8}
Shelfordd1913eu&8IgÿdLaw of ToleranceecHLM#@_H dminimume)@3Hdmaximume&5c"$G#8I'()USud1984be
c45%"!VbSM,%+îHE/L;)012Cm"$G+HîHE
#G8I'(×c"$#G!V'()¿wA"$,+H&G!VCD<
FEH#!!!")GJHI<FEH)2*N*<EH#CD6Ò5¨NH NCh)8I45}¶«"$G+HE#!V@3B@_H,-)W¾q8
I#LMACD6/
LMA#12GJz*<+,%"#|å,-E):|ål0[*TAGZX Uo9XqNØ8I#*0)nE|å#ÜX.¼[OqP"];KHxV
#à'µ)-T5k"HFCdGreenwood, 2007e)(4LMA<+,-./%
+)Ü+QEHU1P&CD6Fc>/A45h±)QO!%789#*
<XU):TAqNLMAUD&Ø8I)TU<FEH#!!!"83+Fi
|¥P#"$!V+Õ)[8AÂ+.z#"$§9,-ABqfàµ&l0
,-);²,-&4,*<89,-Oeki|¥P)[Y>|å,-$,"
$!VÕU*<89Õ#?I)O:}mn-./%0UD#,)A0 12#!"GJh¼[">/#{dDilcher, 1973; Christophel and Blackburn, 1978e/Dî#")<FEH#!"{NOP,-"zW8I'(-7Uh)¨
No(#,)L;i{CDg(!")";s!"{#¼/
>Us)3+Ñ<"$!V,)8\<45#*<EH,-)²,- G¥PB¡>EN-rTH):34,EF¸¡),-]¼)sgH) O½g(&gI#^!"K8CD/O45#./34?r)*<EH#, -'!(Ø8IEFOp)(}4Z*<EH#,-¡>7Ø8g)Q×0 12#2**<EH)<"k8#!"EH)-©¼"*<ij8LDU8
¡)U'!(,-§CD6GJlE*<EHNØ8I/b,3+G"$!V ,#<FEH,-),-#¥PFci|)w%7899:n)Q83+"$
!V,-)'!(BCD6[NØ8I):W,OP,-"-½Uh)O|CD
^!")@})g(#!"GJ;&ïVd821e/
!
!
!
!""""01@AKLMN;<,()*+2345678901@AKLMN;<,()*+2345678901@AKLMN;<,()*+2345678901@AKLMN;<,()*+23456789
45*<EHd!EHe?2*$*<EHd!-1EHe)2*N*<EHd!-t EHeU"#(LMACD6)))78#mn12dWolfe, 1979; Wolfe, 1993;
Wing and Greenwood, 1993; Wilf, 1997; Gregory-Wodzicki, 2000; Greenwood et al., 2004; Traiser et al., 2005; Su et al. 2010elNHNCh/-p45*<EH#LMA CD6{(Z,mn12<)8}e"*<Ø8IOpUW):45!-nEH#
LMACD6{(HNQ,mn12<);a´45°?#./?%"î.AB
%"stuv¤zXh#$/
45!-nEH&LMAGJ)mn#{(Chh¼)º"45#%+
Pm3H)+H,|GFZ#¨Ìo8ALÑ)S&-.12<#%*
+Ã"[*HU,|)+H[¨Ì#,|Fp<)LUG#T5EHI)
P%"sa}tCh#)45"$=>#HFQ)&4,j*#%+C h)45#'!(,|HEFZK-)453+<F$,_m#x
¹<)w7EF,UPFP}B)NýP¬Ìp, #=dͳ)2004e)Roth
et al.d1995ec¬Ìp, =#xy"$EFP)453+.<#'!(Â
8}L;7_))_ZiS¡K'!(,|HFZ#`Ç)]W45LMA CD6#{(FQ/
w3+<NFP¬Ìp, #=n)45_+<#=Q)mnNUO j/W,U¿*HO3)45_+<eH,·.)G%*+8°C78)´;!
=ñ,cò!=O)>²#+OA!=dSu, 1984ae/Roth et al.d1995ec A!I[8})'!(N&')Gb²%7#`2I)45#A!^O
_Z#@12<}2S©`=O)8}wPj*#+H,|<5)45§
mn#'!(NÙ)CU!"GJ[ÑCh/
´./?Ih¼)45µ¿,'@%"<êdBoreal floristic kingdome§¢$
O%"<êdPaleotropical floristic kingdome6dTakhtajan, 1986e)1=NOj
#%"stuv)"$hGÃh0-?J?!þ"·<?ABxy¤Pî .<d2Ðz)1972e)LUEF-.78LMA12#.<)qNk?r?ü.
#%"./#$)Ço45.2Fc%&)P@¤[&Q)k"boN
D'(O#<Fd)*)1975K+,-)1976e/>AE<'#$#Oj)[
ý]W45#LMAGJ)mnNHNCn):vDkP&'12A.§hj/
//
//""""01WXY=>$,#$;<()*+01WXY=>$,#$;<()*+01WXY=>$,#$;<()*+01WXY=>$,#$;<()*+
45?r#%"?./AB%0#$w)mnN&Chn)[SMG
/!<#LMAE/45./%0<#LMACD6)8AQW045#x@bz y'!c7/OP&8W@¾D#z)AB@¤ÅÆhk/PU¿
z*H#)45qN$OAB[$O$%0²)nE@¤#1?#¤X iS%0#!|)Sud1984beÂc./%0<&5qNOj#%0(5)U
=Wd2009e´%&!"p<E./%0<5N"$dS#Ch)2Ðz d1972eQc45yO)@O#"$hGOj)$$%0!<G%0²B%
"./#$E#Oj)àq8}wCDGJÑ<FCh/
TU)CD$,(#ICDV)*<!V*H§o(la)sgH)
!"GJfqï°I/:012UÜ+#9:RN-89 #BH)wP*<
z*c7Añ!V)o*<5NF3H#T5&'?!"GJi|Ñ72)
Qg°¾%0<o#,-V)-'!([%*+#,h}þØ8%0<#qr
`2)vN345/ch)[ý8A«¥*FNsWI#(|,-)AB}
p"$!V,#%"ÜXl0)6!<,-Op#r4)ULÑR./%0
<EFNØ8I#!!!"GJ/
%
%
%
%" " " "&$ &$ &$ &$
0127G#$45=#M!0%"&!'!([34LM#,|
)8}:L)Sç¡>)78&%79:)=>%0&GIS,-ë)P/#
!!!"GJ/TU)L9:*<8N#!V)Ô6i{e,)TYSc!"E
#sTHKOj9:89# #Oj)]W*<#mz{?894òABO HÄ(^7#z{[NUCh/OHÄ(i{s¡>)Âi{A"$OHhC Dg(!"K*<XUQZOs)]W'!(OqP>"*<5#¡>&E F)9:89*<Q*L7Añ!V)AW012®./!<QLV)*<Ø8 IB°¾IOp/
L;)èé012GJ)WÑch(:CsCD45#!!!")8r¡
>l0)-.&'&"$!V,)A<FEH#!"mn,-#Ø8I)13 +9ö,-)!"%"<@#34ïo)!!?½#'6)[8Ammê¥
*&Q*<)w9Y!VO*)Ø8I#`a&n)k}A"$©`Hb
.8Ñ'!"$)xy"$G]34I#<}t)U}4%0(5)!!
2'&5#'MNk¤©##$/
`
' ' '
'" " " "()*+ ()*+ ()*+ ()*+
Adams, J., Green, W. and Zhang, Y. (2008) Leaf margins and temperature in the North American flora: Recalibrating the paleoclimatic thermometer. Global and
Planetary Change. 60:523–534.
Aizen, M. A. and Ezcurra, C. (2008) Do leaf margins of the temperate forest flora of southern South America reflect a warmer past? Global Ecology and Biogeography.
17:164–174.
Ash, A. W., Ellis, B., Hickey, L. J., Johnson, K. R., Wilf, P. and Wing, S. L. (1999) Manual of leaf architecture: morphological descriptions and categorization of dicotyledonous and net–veined monocotyledonous angiosperms. Smithsonian Institution, Washington, DC. 67pp.
Bailey, I. W. and Sinnott, E. W. (1915) A botanical index or cretaceous and tertiary climate. Science. 41:831–834.
Bailey, I. W. and Sinnott, E. W. (1916) The climatic distribution of certain types of Angiosperm leaves. Am. J. Bot. 3:24–39.
Baker–Brosh, K. F. and Peet, R. K. (1997) The ecological significance of lobed and toothed leaves in temperate forest trees. Ecology. 78:1250–1255.
Burnham, R. J. and Johnson, S. Y. (1994) Paleoecological and floristic heterogeneity in the plant–fossil record: an analysis based on the Eocene of Washington. United states government printing office. 36pp.
Canny, M. J. (1990) What becomes of the transpiration stream. New Phytologist.
114:341–368.
Chaloner, W. G. and Creber, G. T. (1990) Do fossil plants give a climatic signal?
Journal of the Geological Society. 147:343–350.
Chen, Tsing-Chang, Shih-Yu Wang, Wan-Ru Huang, Ming-Cheng Yen. (2004)
Variation of the East Asian summer monsoon rainfall. Journal of Climate.
17:744–762.
Christophel and Blackburn. (1978) The Tertiary megafossil flora of Maslin Bay, South Australia: a preliminary report. Alcheringa. 2:3ll–319.
Dilcher, D.L. (1973) A Paleoclimatic Interpretation of the Eocene Floras of
Southeastern North America. In: A. Graham, (Editor) Vegetation and Vegetational History oi Northern Latin America. Elsevier, Amsterdam. 39–59.
Dolph, G. E. (1970) A comparison of local and regional leaf characteristics in Indiana.
Proceedings of the Indiana Academy of Science. 80:99–103.
Dolph, G. E. (1979) Variation in leaf margin with respect to climate in Costa Rica.
Bulletin of the Torrey Botanical Club. 106:104–109.
Dolph, G. E. (1984) Leaf form of the woody plants of Indiana as related to environment.
Being alive on land. Tasks for Vegetation Science. 13:51–61.
Dolph, G. E. and Dilcher, D. L. (1979) Foliar physiognomy as an aid in determining paleoclimate. Palaeontographica Abteilung B. 170:151–172.
Feild, T. S., Sage, T. L., Czerniak, C. and Iles, W. J. D. (2005) Hydathodal leaf teeth of Chloranthus japonicus (Chloranthaceae) prevent guttation–induced flooding of
the mesophyll. Plant Cell and Environment. 28:1179–1190.
Gentry, A. H. (1969) A comparison of some leaf characteristics of tropical dry forest and tropical wet forest in Costa Rica. Turrialba. 19:419–28.
Greenwood D. R. (2007) Fossil angiosperm leaves and climate: from Wolfe and Dilcher to Burnham and Wilf. Courier Forschungsinstitut Senckenberg. 258: 95–108.
Greenwood, D. R., Wilf, P., Wing, S. L. and Christophel, D. C. (2004)
Paleotemperature estimation using leaf–margin analysis: is Australia different?
Palaios. 19:129–142.
Gregory-Wodzicki, K.M. (2000) Relationships between leaf morphology and climate, Bolivia: Implications for estimating paleoclimate from fossil floras. Paleobiology.
26: 668–688.
Grossman, D. H., Faber-Langendoen, D., Weakley, A.S., Anderson, M., Bourgeron, P., Crawford, R., Goodin, K., Landaal, S., Met- zler, K., Patterson, K., Pyne, M., Reid, M. and Sneddon, L. (1998) International classification of ecological of
communities: terrestrial vegetation of the United States. Volume I. The National Vegetation Classification System: development, status, and applications. The Nature Conservancy, Arlington, VA, USA. 1–126.
Hickey, L. J. and Wolfe, J. A. (1975) The bases of angiosperm phylogeny: vegetative morphology. Annals of the Missouri Botanical Garden. 62:538–589.
Hinojosa, L. F., Perez, F., Gaxiola, A. and Sandoval, I. (2010) Historical and
phylogenetic constraints on the incidence of entire leaf margins: insights from a new South American model. Global Ecology and Biogeography. 20:380–390 Holdridge, L. R. (1967) Life Zone Ecology. San Jose, Costa Rica: Tropical Science
Center. 54pp.
Larsen, K. (1995) Flora of Taiwan. Nordic Journal of Botany. 15:574–574.
Lin, S. H., Liu, C. M., Huang, W. C., Lin, S. S., Yen, T. H., Wang, H. R., Kuo, J. T. and Lee, Y. C. (2010) Developing a yearly warning index to assess the climatic impact on the water resources of Taiwan, a complex–terrain island. Journal of Hydrology.
390:13–22.
Martinetto, E., Uhl, D. and Tarabra, E. (2007) Leaf physiognomic indications for a moist warm–temperate climate in NW Italy during the Messinian (Late Miocene).
Palaeogeography, Palaeoclimatology, Palaeoecology. 253:41–55.
Roth, A., Mosbrugger, V., Belz, G. and Neugebauer, H. J. (1995) Hydrodynamic modeling study of angiosperm leaf venation types. Botanica Acta. 108:121–126.
Royer, D. L. and Wilf, P. (2006) Why do toothed leaves correlate with cold climates?
Gas exchange at leaf margins provides new insights into a classic
paleotemperature proxy. International Journal of Plant Sciences. 167:11–18.
Royer, D. L., Kooyman, R. M., Little, S. A. and Wilf, P. (2009) Ecology of leaf teeth: a multi–site analysis from an Australian subtropical rainforest. American Journal of Botany. 96:738–750.
Royer, D. L., Wilf, P., Janesko, D. A., Kowalski, E. A. and Dilcher, D. L. (2005) Correlations of climate and plant ecology to leaf size and shape: Potential proxies for the fossil record. American Journal of Botany. 92:1141–1151.
Schuepp, P. (1993) Tansley review No. 59. Leaf boundary layers. New Phytologist.
125:477–507.
Shelford, V. E. (1913) Animal communities in temperate, America. University of Chicago Press, Chicago,Illinois, USA. 375pp.
Su, H. (1984a) Studies on the climate and vegetation types of the natural forests in Taiwan (I): analysis of the variations on climatic factors. Quarterly Journal of Chinese Forestry. 17:1–14.
Su, H. (1984b) Studies on the climate and vegetation types of the natural forests in Taiwan (II): altitudinal vegetation zones in relation to temperature gradient.
Quarterly Journal of Chinese Forestry. 17:57–73.
Su, H. (1985) Studies on the climate and vegetation types of the natural forests in Taiwan (III) A scheme of geographical climatic regions. Quarterly Journal of Chinese Forest. 18:33–44.
Su, T., Xing, Y. W., Liu, Y. S., Jacques, F. M. B., Chen, W. Y., Huang, Y. J. and Zhou, Z. K. (2010) Leaf margin analysis: a new equation from humid to mesic forests in China. Palaios. 25:234–238.
Takhtajan, A. (1986) Floristic regions of the world. Engl. edn. Translated by T. S.
Crovello, ed. by Arthur Cronquist, Univ. Calif. Press Berkeley. 522 pp.
Taylor, D.W. and Hickey L.J. (1996) Flowering Plant Origin, Evolution, and Phylogeny.
Chapman and Hall, New York. 402 pp.
Traiser, C., Klotz, S., Uhl, D. and Mosbrugger, V. (2005) Environmental signals from leaves-a physiognomic analysis of European vegetation. New Phytologist.
166:465–484.
Wilf, P. (1997) When are leaves good thermometers? A new case for leaf margin analysis. Paleobiology. 23:373–390.
Wilf, P., Wing, S. L., Greenwood, D. R. and Greenwood, C. L. (1998) Using fossil leaves as paleoprecipitation indicators: An Eocene example. Geology.
26:203–206.
Wing, S. L. and Greenwood, D. R. (1993) Fossils and fossil climate – the case for equable continental interiors in the Eocene. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 341:243–252.
Wolfe, J. (1979) Temperature parameters of humid to mesic forests of eastern Asia and relation to forests of other regions of the northern hemisphere and Australasia. US Geological Survey Professional Paper. 1106: 1–37.
Wolfe, J. (1993) A method of obtaining climatic parameters from leaf assemblages. US Geological Survey bulletin (USA). 71 pp.
Yang, J., Wang, Y. F., Spicer, R. A., Mosbrugger, V., Li, C. S. and Sun, Q. G. (2007) Climatic reconstruction at the Miocene Shanwang basin, China, using leaf margin analysis, clamp, coexistence approach, and overlapping distribution analysis.
American Journal of Botany. 94:599–608.
Ò:p?<Å; (2005) z`+ëX+<%7!"&12)m¾<5Q<",G
=>?qJ=12)19(1):1–16/@
p: (2000) <5@OoAF_m$O;!=&'¬Ì)93B/@
CÉ (2006) DÓAF%7²)34LM&&'I12)m¾p·Q<"
øêEF½9)65B/
GHI (2005) ê·JAK<¹"^±L34%7Mö12)D?>ÐÔÕ
=ÏÔN12^OêÙ)94–03–8–012)78B/
G:?@PQ (2001) RQ.<%7<'89dí%e)D?>ÐÔÕ=
ÏßÖ12êÙ)90–032)144B/@
ͳ (2004) 45zyOSTAIU<=)Ö=&%"dS,ÀÙ%
²)m¾45Q<",G=>?qJ=12)18(2):115–127/ ͳ (2006) <5zyOVWXApEUX+<*;!=%7<'&12)m
¾p·Q<"øê12UYF½9)96B/@
|Z[ (2005) Y\]=wUqJ=&%712)m¾p·Q<"øê12U
EF½9)60B/@
=^_ (1988) y;!^ß`<%7<'&12)m¾45Q=12UE
F½9)118B/@
=W ( 2009) 45%7P*IdSB!V&#$)m¾45Q=12UE
F½9)127B/
=am (2009) 'bA=<'ê}B.%7kã&12)m¾pQ<%ø
ê12UEF½9)93B/@
=cd (2004) qeAEU.<;!=&%712)m¾p·Q<"øêEF
½9)68B/@
¦§?GHI?=YQ?f1g@(2005) <5.<Holdridge<%O&hæ)m ÌR×)15(1):61–78/@
¦§?=4?f1g?ij?@PQ?GHI?i!Ö@(2008) <5G%
72&!z&)=Ð12@k)30(4):89–112/@
)* (1975) 45lmn=7A<'&12)45>)92:134–178/
op (2007) p2@A=%"50mîH&"$P*I12)m¾ØÑQ
T,Gê12UEF½9)109B/@
d (2007) qÈI±Ùr·%7<'&12)m¾sxøFQ=ê12U
EF½9)71B/@
0M?tuO?vwx?Úy (2002) ØÑx@<&·.%7P,o!")m
¾<5Q<",G=>?qJ=12)16(3):201–214B/@
0p?zH[?{|}?G: (2009) ,]XA@¶·%7&12)pp=
@k)42:1–22/@
0~ (2004) <5x@O@`AEU&%7!")m¾45Q=12UE
F½9)92B/@
ÉS (2009) Û+ë%7)¡>I./,MöêW¾&12)m¾pQ
<%øê12UEF½9)104B/@
(2009) Ã.<&%7)m¾pQ<%øê12UEF½9)72 B/@
+Ü (2006) Y mÌR×AF%",Gù¥89)oOWY
mÌR×Y/¿)58B/
+,- (1976) lm45n=²&=PU=^$kã)=!!V&^1
2)45=Ð)2(7):6–18/
! (2003) ]·%7<'89dí%e)D?>ÐÔÕ=ÏßÖ1
2êÙ)92–142)96B/@
2 (2004) <5mV·»@ª=%"7A&12)m¾xpQT,GY
/12UEF½9)71B/
23¹?Í45 (1989) =%"<')<54)462B/
2Ðz (1972) 450%")4pJpQ>?ÔÕ)887B/
@PQ?GHI (2003) êmÌR×%7<'89–vå.<)oOW$ê
mÌR×Y/¿ÔN12)80B/@
@PQ?=4 (2004) êmÌR×%7<'89–QZ.<)oOW$
êmÌR×Y/¿ÔN12)116B/@
{|} (2009) %·.<%7<'12)m¾µÉQ>?=Ð=T,
-12UEF½9)148B/
mù (2005) QAA%7<'12)m¾µÉQ>?=Ð=T,-1
2UEF½9)130B/
Í (2001) <@9_m;!=&12)m¾45Q%"12UEF½
9)124B/
~1?*<!VAB45./%0!<~d8I*<µ)NEcx@<)NW cz@<)CEcx<)CWcpz<)SEcxy<)SWczy<e
~2?12~
EFOj'!(^7{#®É@
EFOjT5EH#$,(LMAGJ
!"#$%&&&&&&&&&&'()*&
<FEH ,-o|J
1. *<34 2. *<"$,- 3. "$?½,-
2**<EH
*<EH A*<Ý&
%*+W¾@
*<#'!(@
¡>"$!V,-)
^7%*+O&@
"$'!(@
CDCD!"#GJ±d)EF
1.45#*<EH!'!()34>+&'6 2.45=$OjT5EH#!!!"
3.45=OjT5EH#!!!")mn12&EF 4.45R./%0!<#!!!"Ch
£!%*+O)@
^7%*Oo&@
2**<'!(@
A'!(§34LMCDJ 1.)%*+"$,(CD 2.)ïo"P,(CD
A'!(,-§%*+CDJ 1.'!()%*+"$,(CD 2.!""$o[%*+&,|
圖3、年均溫（MAT，單位°C ）變化圖 圖4、年降水（Prec，單位mm）變化圖
圖5、最冷季降水（PrC，單位mm） 圖6、最暖季降水（PrW，單位mm）
~7?*<EH&>*<!'!()%*+,|~/x¾"*<'!(dREL!e) y¾"A°C8I#%*+dMATe/x?yg81P&ICD¦oIJ
MAT = 15.1× REL!8.276)r2 = 0.244)df=843)p<0.001
64-1
~8?2*$*<EH&>*<!'!()%*+,|~/x¾"%*+O#2
*$*<'!(dREL!-1e)y¾"A°C8I#%*+dMATe/x?yg81P
&ICD¦oIJ
MAT = 55.7×REL!-1 - 16.629)r2 = 0.896)df=16)p<0.001
65-1
~9?2*N*<EH&>*<!'!()%*+,|~/x¾"%*+O#2
*N*<EH'!(dREL!-te)y¾"A°C8I#%*+dMATe/x?yg8 1P&ICD¦oJ
MAT = 63.7×REL!-t - 18.322 )r2 = 0.912)df=16)p<0.001
65-11
~10?<FEH&M!0%""$o[%*+,|~/x«l¾"A°Cc$
µ#%*+dMATe)7Zy«l¾"]%*+O#"$o,|/uðd----e c'!"$o) ÅðdZZZZ[[[[\\\\[[[[\\\\ecxy"$o/8Zy¾"]%*+O
&xy"$o^µ'!"$o&g)G~pAÎq8I/
~11?'!([*<;oBT5EH&,| ~dbox-plote)"G23.5°C? 19.5°C?15.5°C?11.5°C(%*+Oo)[ß 2;?5;?10;?15;?20
;?25;?30;?50;?75;*<cXS*<1^ü¥*1000)B 1;
*<§UN*<V#'!(2*gKx¾E#18I s;*<#2*$*<E H)#8I2*N*<dQ×!-teEH)"8I<FEH#GJ)-#oá"[ß
¥*&*<oKy¾8I'!(RELdi$µ)0≤REL 1≤ e/zc 8'!(
&,h)zIzÎcís!µo)£z&Èqcpµo)qÄc 2*o)zEzÎcí!µo)TÄcê g&7Cg)R%*+
OGOj*<o1P& 8'!(#2*oA2¡ïl/
~12?*<EH&x@<!'!()%*+,|~/x¾"*<'!(dREL!e) y¾"A°C8I#%*+dMATe/x?yg81P&ICD¦oJ
MAT = 5.2×REL!+ 15.994)r2 = 0.192)df=186)p<0.001
68-1
~13?*<EH&z@<!'!()%*+,|~/x¾"*<'!(dREL!e) y¾"A°C8I#%*+dMATe/x?yg81P&ICD¦oJ
MAT = 8.7×REL!+ 9.706)r2 = 0.099)df=155)p<0.001
68-2
~14?*<EH&pz<!'!()%*+,|~/x¾"*<'!(dREL!e) y¾"A°C8I#%*+dMATe/x?yg81P&ICD¦oJ
MAT = 5.5×REL! + 11.984)r2 = 0.105)df=117)p<0.001
68-3
~15?*<EH&zy<!'!()%*+,|~/x¾"*<'!(dREL!e) y¾"A°C8I#%*+dMATe/x?yg81P&ICD¦oJ
MAT = 20.9×REL! + 4.754)r2 = 0.316)df=267)p<0.001
68-4
~16?*<EH&xy<!'!()%*+,|~/x¾"*<'!(dREL!e) y¾"A°C8I#%*+dMATe/x?yg81P&ICD¦oJ
MAT = -0.8×REL! + 22.597)r2 = 0)df=75)p=0.629
68-5
~17?*<EH&x<!'!()%*+,|~/x¾"*<'!(dREL!e)y
¾"A°C8I#%*+dMATe/x?yg81P&ICD¦oJ MAT = 20.1×REL! + 6.112)r2 = 0.448)df=38)p<0.001
68-6
