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Critics Biodiveristy

Critics Biodiveristy

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Critics Biodiveristy
Critics Biodiveristy

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Published by: Rubén De Gracia Santo on Oct 15, 2013
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337
On a Biodiversity Study Based on Papers of
Taiwan’s Biodiversity

Hsin Chi, Yu-Bin Huang, Yu-Da Dai, Yi-Ying Wu and Jen-Wei Liu
Laboratory of Theoretical Ecology, Department of Entomology,
National Chung Hsing University

Abstract
Since the funding of a long-term ecological study by the National Science Council,
an intensive biological inventory has been carried out in Taiwan over several years.
There were follow up projects on biodiversity targeted by the survey and classification
in some specific areas as well. Biodiversity indices were calculated and the similarities
between different areas were studied. Because some authors did not fully understand
the formulae they were using, there were some errors in interpretation. The description
and interpretation of the data needs to be improved. We reviewed papers on
biodiversity in Taiwan and propose the following suggestions for sampling for
biodiversity, for ecosystem studies of biodiversity, and for permanent sampling plots.
Because of differences in the behavior and temporal and spatial scale of habitats
between species, organisms of different species would be collected with differing
probabilities by the various sampling methods. Therefore, in biodiversity studies, it is
necessary to identify the distribution of habitats of the various organisms. Proper
sampling methods and sample sizes can then be planned and calculated based on the
spatial distribution pattern and the life history of the respective organisms in the habitat.
Only then, can the number of species and their densities be estimated with confidence,
and the diversity index calculated accordingly. In calculating the diversity indices, if
there are still significant differences between the observed number and the actual total
number of species, the calculated diversity indices will change from time to time with
the accumulation of data. This will make the indices meaningless. Because of the
deficiencies in abstract reasoning underlying the definition of most well-known
biodiversity indices, they should be used with caution. From an ecosystem viewpoint ,
knowledge of species number and their abundances are only constitutive elements of the
system. This is not sufficient for a sustainable management of the ecosystem. For an
ecosystem study, not only the composition of a system (the general meaning of
338
“biodiversity”) should be studied, but also the system structure (the relationships
between members in the food-chain or food-web) and the dynamics (the growth,
reproduction, predation, competition, mutualism, or human harvesting, etc.). The
seasonal change of biodiversity of an ecosystem results from the changes of growth rate,
reproduction rate, predation rate and competition of all species and their interactions.
All of these should be studied, based on ecological theories such as life tables, predation,
competition and harvesting. Because of the numbers of components and the complexity
of their interactions, computer simulation is a necessary tool. If the management of
representative permanent sampling plots of an ecosystem is able to minimize human
influence, and if the methods of study and theories are correct, then the theory derived
from abstract reasoning from the data can be confidently used. Robust methods for
conservation and sustainable biodiversity can then be undertaken. Using these methods,
the existence of individual permanent sampling plots can be left to ecological
succession.

Key words: ecosystem, life table, sampling, biodiversity.

339
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¦³¨Ç¬O´Ó-¹ªÌ¡A¦³¨Ç¬O»G-¹ªÌ¡C¦UºØªº¬Û¹ï¼Æ¶q¤]©¹©¹¨M©w©ó¨ä¦b-¹ª«Ã줤¤§¦a
¦ì¡C¦]¦¹¡A-pºâ´Ï¦a¤¤¯S©w¤@Ãþ¥Íª«ªº diversity index¡A©¿²¤¨ä¤¤-¹ª«Ã줧Ãö«Yªº
°µªk¨ä¥ÍºA·N¸q-ȱoÀË°Q¡C¥H¤U¶È´N±`¥Îªº Shannon-Wiener index »P Simpson’s
index µy¥[°Q½×¡G
1. Shannon-Wiener index
Shannon-Wiener index (Shannon 1948) ¤§¤½¦¡¦p (1)¦¡©Ò¥Ü¡C¦¹¦¡¦b¤Æ¾Ç¼ö¤O¾Ç
¤¤ªº¾ú¥v§ó¤[¡A¨Ò¦p¡G¦pªG§Ú-Ì°Ñ¦Ò McQuarrie (1973)ªº “Statistical Thermo-
dynamics”¤@®Ñ²Ä¤@³¹ªº²ßÃD 1-15 ´N·|¤F¸Ñ Shannon-Wiener index ³ ̤ j- Ȫ º ± ø ¥ ó ¡ A
¤]´N·|¤F¸Ñ¬°¦ó Shannon-Wiener index ªº¨Ï¥Î¤@ª½¦³ª§Ä³¡]¸Ó®Ñªº§@·~ 2-5 »P 2-6
¤]-ȱo°Ñ¦Ò¡^¡C¤½¦¡ 1 ¤¤ s ¬°Á`ºØ¼Æ¡A p
i
¬°²Ä i ºØ¤§-ÓÅé¼Æ¦b©Ò¦³ºØ ¤ § - Ó Å é ¼ Æ Á ` © M
¤¤©Ò¦û¤§¤ñ²v¡C§Q¥Î¦¹¤½¦¡¤§±ø¥ó¬°¡G s ¬°¤wª¾¡A¥B¥ô¦ó¤@ºØªº¨C¤@-ÓÅé³Qµo²{
¤§¾÷²v§¡¬Û¦P¡CµM¦Ó¡A¤j¦h¼Æ§@ªÌ¦b-pºâ Shannon-Wiener index ®É³£µLªk½T©w s
§Y´Ï¦a¤§©Ò¦³ºØ¼Æ¡A¤]µLªk«OÃÒ©Ò-pºâ¤§ p
i
½T¹ê¬°´Ï¦a¤¤¤§¬Û¹ïÂ׫סC-Y¦A¶i¦æ
¤@¦~ªº½Õ¬d¡A¤Sµo²{¤£¤Ö·s±Ä¶°ºØ¡A«h s »P p
i
§¡§ïÅÜ¡A Shannon-Wiener index ¦ÛµM
¤£¦P¡C³oºØ¡u¥Íª«¦h¼Ë©Ê«ü¼Æ³v¦~¼W¥[¡vªº²{¶H¡A¦pªG¤£¬O¦]¬°¡u¥Íª«¦h¼Ë©Ê³v
¦~¼W¥[¡v¡A¦Ó¬O¥Ñ©ó¥ý«e½Õ¬dªº s »P¹ê»Ú¤§¼Æ-Ȥ´¦³¬Û·í®t¶Z¡A«h¬O¤@ºØ¿ù»~¡C
¦]¦¹¡A¦b s ¤§-ÈÁͪñÃ-©w¤§«e¡A¥ç§Y´Ï¦a¤¤¤j¦h¼ÆªººØÃþ³£¤w±Ä¨ì¤§«e¡A¤£©y-p
ºâ Shannon-Wiener index¡C-Y´Ï¦a¤¤¤j¦h¼ÆªººØÃþ³£¤w±Ä¨ì¡A«hÀ³¶i¤@¨B±Ä¥Î¥¿
½Tªº¨ú¼Ë¤èªk¡A¦ô-p¦U-ÓºØÃþªº¬Û¹ïÂ׫סC·íµM¤£¥i¯à¹ï¨C¤@ºØÃþ³£§@ºë½Tªº±Ä
¼Ë¡A¦ý¬O¦Ü¤ÖÀ³°w¹ï¥D-nºØÃþ±Ä¼Ë¡A¦]¨ä¼vÅT³Ì¤j¡C³\¦h§@ªÌ±N Shannon-Wiener
index »~¬° Shannon-Weaver index¡AÁöµM°ê¥~³\¦h±Ð¬ì®Ñ¦-¤w«ü¥X³oºØ¿ù»~¡A¦ý°ê
¤º¤´¦³³\¦h ½×¤å¨Ï¥Î¿ù»~ªº¦WºÙ¡A¨Ò¦p ¡G³¯¤l ¯Eµ¥¡] 1999 ¡^¡B¶ÀÄ£³q µ¥
¡] 2000¡^¡B³¯´Â¦`µ¥¡] 2001¡^¡B¶¾Â׶©µ¥¡] 1998¡^¡CŪªÌ-Y¯à¾\Ū¤Æ¾Ç¼ö¤O¾Çªº
±Ð¬ì®Ñ (¨Ò¡G McQuarrie 1973)»P Shannon (1948)-ì¤å¡]¨ä Appendix 2 ¦³¸Ô²Óªº±À¾É
¹Lµ{¡^¡A¸Ô²Ó¤F¸Ñ¨ä²z½×-I´º¡A¥i¥H½T¹ê¤F¸Ñ¥Î©ó¸ÑÄÀ¥ÍºA¾Ç information ¤§±ø
¥ó¡C



343
2. Simpson’s index
Legendre and Legendre (1983) ¦b¨ä®Ñ¤¤¼g¹D “The probability that two randomly
chosen specimens belong to the same species is the measure of concentration proposed
by Simpson, i.e., the sum of these combined probabilities for the different species:”
( )
( )
( )
( ) 1
1
1
1
1
1


=




=
=
N N
N N
N N
N N
n
i
i i n
i
i i
(2)
¨ä¤¤ n ¬°ºØ¼Æ¡A N
i
¬°²Ä i ºØ¤§-ÓÅé¼Æ¡A N ¬°©Ò¦³ºØ¤§-ÓÅé¼Æ¤§©M¡C Magurran (1988)
¦b¨ä®Ñ¤¤²Ä 39 -¶¼g¹D “Simpson (1949) gave the probability of any two individuals
drawn at random from a infinitely large community belonging to different species as:

=
2
i
p D (3)
where p
i
= the proportion of individuals in the ith species. In order to calculate the
index the form appropriate to a finite community is used:”
( )
( )

|
|
.
|

\
|


=
1
1
N N
n n
D
i i
(4)
¨Ì¾Ú¾÷²v²z½×¡A¤½¦¡ 2 »P 4 ³£¬O¦Û¦³--¤§¸s»EÀH¾÷¨ú¨ì¨â-Ó-ÓÅé “ÄÝ©ó¦PºØªº¾÷
²v ”¡A¤½¦¡ 3 «h¬O¦ÛµL--¤§¸s»EÀH¾÷¨ú¨ì¨â-Ó-ÓÅé “ÄÝ©ó¦PºØªº¾÷²v ”¡C¦]¦¹¡A
Legendre and Legendre (1983)ªº¸ÑÄÀ¤~¬O¥¿½Tªº¡C Ludwig and Reynolds (1988) ªº¸Ñ
ÄÀ»P Legendre and Legendre (1983)¬Û¦P¡C¦A¨Ì¾Ú¾÷²v²z½×¡A¨ú¨ì¨â-Ó-ÓÅé “ÄÝ©ó¤£
¦PºØªº¾÷²v ”·íµM´N¬O :
( )
( )
D
N N
N N
D
n
i
i i
− =


− =

=
1
1
1
1
1
2
(5)
¤½¦¡ 5 ±`¥Î¨Óªí¥Ü diversity¡CÁöµM¦³¨Ç¤åÄm¡]¨Ò¦p¡G MacArthur 1972¡A Magurran
1988¡^»{¬° 1/D ¤]¥i¥H¥Î¨Óªí¥Ü Simpson’s index¡A¦ý¬O 1/D ¨Ã¨S¦³¥ô¦ó²z½×¨Ì
¾Ú¡C¦Ó¥B¡A D»P D
2
¡]§Y 1 – D¡^§eª½½uÃö«Y¡A¦ý D»P 1/D«h¬°-˼ƨç¼Æ¤§¦±½uÃö
«Y¡F D -Ȭ° 0 ®É¡A 1 – D = 1¡A 1/D «hµL·N¸q¡C©Ò¥H¡A§Ú-Ì»{¬°¤£©y¨Ï¥Î 1/D ªí¥Ü
Simpson’s index¡C°ê¤º¤]¦³¥Î 1/Dªº½×¤å¡A¨Ò¦p¡G¶ÀÄ£³qµ¥¡] 2000¡^¤§µÛ§@¡C
µM¦Ó¡A®Ú¥»°ÝÃD¦b©ó¡G¥Ø«e±`¥Îªº¦UºØ¥Íª«¦h¼Ë©Ê«ü¼Æ»P¬Û¦ü©Ê«ü¼Æ§¡¯Ê¥F
¥ÍºA¾Ç²z½×½×ÃÒ¡A¨Ï¥Î®ÉÀ³²`«ä¼ô¼{¡C¬ì¾Ç®a¾D¹J°ÝÃD®É¡A±`±`§Q¥Î²{¦¨ªº¡u«ü
¼Æ¡v©Î°®¯Ü³Ð¥ß·s¡u«ü¼Æ¡v¸Õ¹Ï¸ÑÄÀ¤@¨Ç²{¶H¡C MacArthur (1972) ¼g¹D “Those
344
who have used diversity to mean a number combining aspects of both the number of
species and the evenness of their abundances have wasted a great deal of time in
polemics about whether

i
i
p
2
1 or


i
i e i
p p log or ( ) ! ! ! !
2 1 n
N N N N ⋅ ⋅ ⋅ or some
other measure is “best.” ”¡C³o¦h¤Ö¤]»¡©ú¤F¨Ï¥Î¡u«ü¼Æ¡vªº§x¹Ò¡C«Ü¦h±Ð¬ì®Ñ»P
ºô¸ô¸ê®Æ³£¹ï¡u¥Íª«¦h¼Ë©Ê«ü¼Æ¡v¦³²`¤Jªº°Q½×¡A-ȱoŪªÌ°Ñ¦Ò¡C¥Ñ©ó diversity
index ¥»¨-°ÝÃD¤´¦h¡A¦ÛµM¤£©y¦A¨Ì¾Ú diversity index ¶i¤@¨B§@¨ä¥L²Î-p¤ÀªR¡C

¥Íª«¦h¼Ë©Ê½Õ¬d¨ú¼Ë
§Ú-Ì»{¬°¡A¥Íª«¦h¼Ë©Ê½Õ¬d¤u§@À³¥ý¥Ñª«ºØ½Õ¬d¶}©l¡A°£¤F¤@¯ë±`¥Î¤èªk
¥~¡A¤@©w-n¥Î¥ØµøÆ[¹î·j´M¥Íª«¡C¤£-n¤@¶}©l´N¤j¶q±þ¥Í¡C³Ì¦n¦b½Õ¬d¦aÂI±J
Àç¡AÀH®É½Õ¬d¡CµM«á¦A¨Ì¾Úªì¨B¬ã¨s°w¹ï¥D-n¥Íª«©Î¯S©w¥Íª«¶i¦æ¦³¨t²Îªº¨ú¼Ë
½Õ¬d±K«×¡C¥Ñ©ó©Ò¦³ diversity indices §¡°²³]´Ï¦a¤¤¨C-Ó±Ú¸sªº¨C-Ó-ÓÅé³Q±Ä¨ìªº
¾÷²v³£¬Û¦P¡]©Ò¥H¤~±N¦UºØ¥Íª«¤§¼Æ¶q¬Û¥[¦Ó±o N¡^¡A¦]¦¹-Y¨ú¼Ë¤èªk¤£¯à«OÃÒ
¨C-Ó-ÓÅé³£¬OÀH¾÷¨ú±o¡A´N¤£¯àÀ³¥Î³o¨Ç¤½¦¡¡C Magurran (1988)ªº®Ñ¤¤¤]°Q½×¨ì
diversity ¬ã¨s¤¤ sampling ªº-«-n©Ê¡C¡]°ê¤º³\¦h½×¤å³£±N Magurran (1988) ¦C¬°¤Þ
¥Î¤åÄm¡A¦ý¬O¥Ñ½×¤å¤¤ªº¿ù»~«o¥iª¾³\¦h§@ªÌ©|¥¼½T¹ê¾\Ū¸Ó®Ñ¡^¡C¥Ñ©ó¤£¦P¥Í
ª«¤§¦æ¬°»P´Ï©Ò®ÉªÅ¤Ø«×¤£¦P¡A¦UºØ¨ú¼Ë¤èªk±Ä±o¤£¦P¥Íª«¤§¾÷²v¥ç¤£¦P¡C¦]¦¹
°w¹ï¤£¦P¥Íª«»Ý-n¥Î¤£¦Pªº¨ú¼Ë¤èªk»P½d³ò¡A©øÂÎ¥i¯à¤@¥-¤è¤½¤Ø§Y¥i¡A³¾Ãþ¥i
¯à-n¼Æ¦Ê¥-¤è¤½¤Ø¨ì´X¥-¤è¤½¨½¡C¦]¦¹¡A¥Íª«¦h¼Ë©Ê¬ã¨s¤¤¡A-º¥ýÀ³¤F¸Ñ½Õ¬d¦a
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¥¬¡A-q©w¾A·íªº¨ú¼Ë¤èªk¨Ã¦ôºâ©Ò»Ý¼Ë¥»¼Æ¡A¨ä¨ú¼Ëµ²ªG¤~¯à½T¹ê¤Ï¬M¥Ð¶¡¥Íª«
ºØ¼Æ»P±K«×¡A¤]¤~¯à-pºâ¦h¼Ë©Ê«ü¼Æ¡C¦ý¬O¡A-pºâ¦UºØ¥Íª«¦h¼Ë©Ê«ü¼Æ®É¡A¥²¶·
¥ý¤F¸ÑÁ`ºØ¼Æ¡A§_«hµo²{ªººØ¼Æ-Y³v¦~¼W¥[¡A¨C¦~µoªíªº diversity index ¤]³v¦~§ï
ÅÜ¡C¦b¤ñ¸û´Ï¦a¶¡¤§¬Û¦ü©Ê®É¡A¨C¦~µoªíªº similarity index ¤]·|§ïÅÜ¡C¦³Ãö©óºØ
¼Æªº°ÝÃD±N©ó«á¤å°Q½×¡C¦¹¥~¡A¥Íª«¦h¼Ë©Ê¨ú¼Ë½Õ¬d¤u§@¤¤¡A-Y©¿²¤±Ú¸s¥Íªø¹L
µ{¤¤¦~ÄÖ©ÎÄÖ´Áµ²ºc¤§ÅܤơA½Õ¬d©Ò±o¤§±K«×«h¥i¯àµLªk¥¿½T¤Ï¬M¹ê»Ú¥Ð¶¡±K
«×¡A-Y¾Ú¥H-pºâ¤§¦h¼Ë©Ê«ü¼Æ¥ç¤£¥i«H¡C¦]¦¹¡A¤]¥²¶·¨Ì¾Ú¥Íª«ªº¥Í¬¡¥v¡A¿ï¾Ü
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ªº±Ð¬ì®Ñ¬Û·í¦h¡A¨Ò¦p¡G Southwood (1978)¡A Kershaw and Looney (1985)¡A Moore
and Chapman (1985)¡A Ludwig and Reynolds (1988)¡A Krebs (1999)¡C¦³¨Ç®Ñ¤¤¦³¿ù
»~¡AŪªÌ¥²¶·¥J²Ó§PÂ_¡C³Ì¦n¤ñ¸û¤£¦P±Ð¬ì®Ñ¡A§ä¥X¥¿½Tªº¤èªk¡C°Ñ¦Ò½Ķªº®Ñ
Äy®É¡AÀ³¥[-¿ÂÔ·V¡A¦]¬°Â½Ä¶¿ù»~ªº±¡§Î¬Û·í´¶¹M¡C¦³Ãö¨ú¼Ë²z½×¤è-±¡A«hÀ³¸Ó
°Ñ¦Ò¥H²Î-p¾Ç¬°°ò¦¤§®ÑÄy¡A¨Ò¦p¡G Cochran¡] 1977¡^»P Thompson¡] 1992¡^´N¬O
¨â¥»-ȱo¾\Ūªº®Ñ¡C¡]¥»¤å¤¤¦U¤½¦¡¤§¸Ô²Ó»¡©ú§¡½Ð°Ñ¾\-ìµÛ©Î±Ð¬ì®Ñ¡A¥H«K²`
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»P¦ôºâ©Ò»Ý¼Ë¥»¼Æ¡C¦bªÅ¶¡¤À¥¬¤è-±¡A³Ì±`¨£ªº¬OÀH¾÷«¬¤À¥¬»P»E¶°«¬¤À¥¬¡CÀH
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»E¶°«¬¤À¥¬¦b©øÂάã¨s¤¤±`¥Îªº¬O negative binomial distribution¡]-t¤G¶µ¤À¥¬¡^¡A
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346
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distribution »P negative binomial distribution ¤£¦ý©M¨ú¼Ë¦³Ãö¡A¤]©M Simpson’s index
¦³Ãö¡C Simpson (1949)ªº½×¤å¤¤´N¥Î¨ì¤F³o¨âºØ¤À¥¬¡C
¨M©w½Õ¬d¤§´Ï¦a½d³ò«á¡A¨Ò¦p¦b¯ó¦a½Õ¬d¬YºØ¼u§À¥Ø©øÂΡ]«UºÙ¸õÂΡ^¡A¥i
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¥ÍºA¨t¤§¨ú¼Ë®e³\¸û¤j»~®t¡^¡F t ¬° Student’s t ¤À¥¬¤§-È¡A t
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347
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n ¬°§Q¥Î¦³--±Ú¸s®Õ¥¿¤§¼Ë¥»¼Æ¡A N ¬°¼Ë¥»ªÅ¶¡¥À±Ú¸sÁ`¼Æ¡]¦¹³B«Y«ü¨ú¼Ë
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¡^¡C·í n/N > 0.05 ®É¡A«KÀ³±Ä¥Î
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¸q¡A³o¬O¤j®a³£»{¦PªºÆ[©À¡C Legendre and Legendre (1983) ¤]¼g¹D “As a final
caution, the reader should be warned that a meaningful interpretation of a data matrix
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