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Biological Con servation 55 1991) 235-254. ~ ' q t~ , . ~ V ~
~
W hat t o P ro tec t Sys t emat i c s and t he Ag ony o f Cho
iceR . I. V a n e - W r i g h t , C . J . H u m p h r i e s P . H .
W i ll ia m s
Biodiversity Programme, Departments of Botany and Entomology,The
Natural History Museum, Cromwell Road, London SW7 5BD, UKReceived 8
September 1990; accepted 29 September 1990)
A B S T R A C TPol i t ic ians an d sc ient is ts a l ike non,
agree that a pr ior i ty l i s t o f g lobal cen tres
fo r prese rvat ion o f b io logical d ivers i ty is required .
Dive rs i ty has general lybeen me asur ed only in terms o f spec
ies richness , or in the fo rm o f indicesc o m b i n i n g r i c h
n e s s w i t h a b u n d a n c e . S u c h m e a s u r e s a r e c
o n s i d e r e dinadequ ate fo r the task in hand. A n ove l index
, based on the in format ioncon ten t o f c lad i s t i c c las s i
f i ca t ions and g iv ing a measure o f taxonomicdis t inc tness ,
i s in troduced. This tax ic d ivers i ty measure , when coupled wi
thde ta i l ed know ledge o f d i s t r ibu t ion , can be used in
mod i f i ed ana ly se s o f thetype prev io us ly deve loped as cr
i t ica l fau na s a nalys is or ne tw ork analys is .Cen t ra l to
a l l such ana ly se s i s the concep t o f com pleme n tar i t y o
f f l o ras or
.faunas . B y em ploy ing comp lemen tar i t y , s t ep -w ise
procedures can iden t i f yop t ima l l y e f f ic i ent , s ing le
- s it e s equences o f p r ior i t y a reas fo r a group , tak
ingex is t in g reserves in to accou nt or not , as required. For
prac t ical p lanning i t i sconcluded that two bas ic rounds o f
analys is are required: f ir s t , recogni t ion o fg loba l pr ior
i ty areas by ta x ic d ivers i ty techniques; secondly , w i th in
any sucharea, ana ly s i s w i thou t tax ic we igh t ing (as be
ing deve loped by M argu le s andh i s co -worker s ) to iden t i f
y a ne twork o f r e se rve s to con ta in a l l loca l taxa a
ndecosys tems . The paper conc ludes w i th a br i e f d i scus
sion o f some imm ed ia teprospec t s . fo r deve lopm en t o f a s
y s t ema t ic approach to g loba l conserva t ionevaluat ion .
I N T R O D U C T I O NBiological d i v e r s i t y is i n c r
is is H u m a n a c t i v it i e s a re d e s t r o y i n g t h e n
a t u r a lworld and i t s b i o t a at an e v e r i n c r e a s i
n g r a te Only a few previous m a s sextinctions, as docum ent ed
by the geological record, could comp are to t h e
235Biol. Conserv.0006-3207/91/503-50 1991 ElsevierScience
Publishers Ltd, England.Printedin Great Britain
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R. I . Vane-Wrigh t , C . J . Humphr ies , P . H . Wi l l i
ams
; n o w o c c u r r i n g . A c c o r d i n g t o M y e r s (1
98 9), w e m a y w e lt b e a b o u t t oh e ss a m a j o r e x t i
n c t i o n s p a s m .ag a in s t th i s r ea l it y , d ep r e ss
in g f o r t h o se w h o see in b io d iv e r s i ty n o t o n lyt
e r ia l b en e f i t ( P r an ce et al . 1 9 8 7 ) b u t a l so sp
i r i t u a l we l lb e in g ( Lo v e lo ck ,9 ) , m o r e a n d m
o r e l o c a l , n a t i o n a l a n d i n t e r n a t i o n a l c
o n c e r n o v e r t h i sv e r s i b l e c h a n g e i s b e i n
g e x p r e s s e d ( T h a t c h e r , 1 9 8 9 ; H R H P r i n c e
o fle s, 1 9 90 ). S o m e c o n c e r n is t ra n s l a t e d i n
t o a c t i o n - - a c t i o n t o s t e m t h e t id e~o llut io
n, an d ac t io n to co n se r v e sp ec i e s o r se t a s id e tr
ac t s o f m an ag ed o r-1 l an d s , in t h e h o p e o f sav in
g a t l e a s t so m e o f t h e Ea r th s b io d iv e r s i ty f o
rse lv es an d f u tu r e g en e r a t io n s ( Ch a lk e r , 1 9 9
0 ) .~ es ou rc es fo r p r o t e c t i n g b i o d i v e r s it y
t h r o u g h t h e c o n t r o l o f l a n d u s e w i lla y s b e
l i m i te d . S e t ti n g a s i d e l a n d f o r c o n s e r v a
t i o n p u r p o s e s is o f t e n i n; c t c o n f l i c t w i t
h e c o n o m i c p r e s s u r e s t h a t p r o d u c e e n v i r
o n m e n t a lt r u c ti o n . C l e a r ly , r e s o u r c e s n
e e d e d f o r c o n s e r v a t i o n a r e b o t h c o n s t r a
i n e da n d d e p e n d e n t u p o n h u m a n e c o n o m y .:
aced wi th th i s p r o b lem we co n s id e r i t h ig h ly d es i
r ab le t o f i n d e f f ec t iv em s o f m e a s u r i n g b i o
d i v e r s i t y , s o a s t o b e a b l e t o q u a n t i f y a n
d t h e r e b yd m i s e th e c o n t r i b u t i o n o f m a n a g
e d a r e as ( b o th n a t u r a l a n d s e m i-a r al : W e s t
m a n , 1 99 0) t o t h e c o n s e r v a t i o n o f t h e E a r t
h s r e m a i n i n go g ica l d iv e r s i ty . Th i s o b jec t
iv e i s i n l in e w i th p r ev io u s ca l l s b y sc i en t i
st sv e n , 19 80 ; R o b e r t s , 1 9 8 8 ) a n d p o l i t ic i
a n s (t h e p r o p o s e d I n t e r n a t i o n a lw e n t i o n
t o C o n s e r v e B i o l o g i c a l D i v e r s i ty : T h a t
c h e r , 1 98 9; I U C N , 1 99 0).d r a w i n g a t t e n t i o n
t o t h e p r o p o s e d C o n v e n t i o n , C h a l k e r ( 1 9
9 0 ) h a s, ific ally s ta t e d t h a t a f u n d a m e n t a l r
e q u i r e m e n t is a g r e e m e n t o n a g l o b a lo f c e n
t r e s o f b i o l o g i c a l d i v e r s i t y f o r p r i o r i
t y c o n s e r v a t i o n . P o l i t i ca l l y ,a r t a t i o n
s m u s t b e t r a n s l a t e d i n t o a c t i o n s . S c i e n
ti fi c al ly , i n o r d e r t ol t if y c r i t i c a l a r ea s
f o r p r io r i t y ac t io n , w e co n s id e r a v i t a l n ee
d to b e th e: t i v e m easu r em en t o f b io d iv e r s i ty i
t s e l f .~ur p u r p o s e h e r e i s t o i n t r o d u c e t h
e i d e a o f a n o v e l i n d e x f o rLsur em en t o f b io lo g
ica l d iv e r s i ty w h ich , we b e l iev e , r e fl ec ts i n f
o r m a t io n9 d ed b y c ladi s t i c ( as d i s t i n c t f r o
m p h y l o g e n e t i c ) re la t ionships a n d t h e n t oly th
i s i n d ex o f t a x i c d i v e r s i ty t o w i ld l if e c o n
s e r v a t i o n e v a l u a t i o n ( U s h e r ,5), a n d s p e
c if ic a ll y t o t h e p r o b l e m o f h o w t o o p t i m i s
e u s e o f r e s o u r c e s f o r~erva t ion o f b iod ivers i ty
(c f . Margu les , 1989) .
I E M E A S U R E M E N T O F B I O D I V E R S I T Y - - A R E
A L L S P EC I E SE Q U A L ?hen you can measure what you are
speaking about, and express it in numbers, youow something about
it; but when you cannot measure it, your knowledge is of a~ager and
unsat isfactory kind: it may be the beginning of knowledge, but you
havercely, in your thoughts, advanced to the stage of science.
'
William Thompson Lord Kelvin), Popu lar Lec tur es Addresses ,
1891-1894
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ystem atics a nd conservation evaluation 37
W h i t t a k e r (1 97 2), w h i l e c o m m e n t i n g t h a
t i f d i v e r s i ty is re c o g n i s e d a s a ne v o l u t i o
n a r y p r o d u c t , it m a y c a u s e n o s u r p ri s e t h a
t n o s i n g le m e a s u r e m e n tse rv es a l l p u rp o ses ,
su g g es t ed th a t t h e r e a r e i n h e re n t d i f fi cu lt
i e s inm e a s u r i n g a s p e c t s s u c h as i n t e ll i g e
n c e , o r g a n i c c o m p l e x i t y a n d phylogene t
icrelationship ( o u r e m p h a s i s a d d e d ) .
H o w i s d i v e r s i t y m e a s u r e d c u r r e n t l y ?
B i o l o g i s t s h a v e t r i e d t o m e a s u r es p ec ie s
r i c h n e s s - - t h e t o t a l n u m b e r o f s p ec ie s o f
a g r o u p r e li a b ly k n o w n t o b ei n d i g e n o u s t o
a p a r t i c u l a r p l a c e . E c o l o g i s t s o f t e n w i
s h t o i n c l u d e , i na d d i t i o n , i n f o r m a t i o n
o n c o m m o n n e s s a n d r a r i t y , b y c a l c u l a t i n
g o n e o rm o r e i n d i c es th a t c o m b i n e m e a s u r e
s o f t h e n u m b e r o f s pe c ie s i n a s a m p l et o g e t
h e r w i t h t h e r e l a t iv e a b u n d a n c e o f t h o s e
s p e c i es ( P e et , 1 97 4; T a y l o r ,1 9 7 8 ) . I n a m o
re e l ab o ra t e sy s t em , r ecen t ly r ev iewed b y Bo n d (1
9 8 9 ) ,W h i t t a k e r ( 1 9 7 2 ) s u g g e s t e d t h a t a
t l e a s t t h r e e m e a s u r e s a r e r e q u i r e d :a l p
h a d i v e r s i t y ( s p e c i e s r i c h n e s s o f s t a n d
a r d s i t e s a m p l e s ) ; b e t a d i v e r s i t y( d if fe
r e n ti a ti o n b e tw e e n c o m m u n i t i e s a l o n g h a
b i t a t g r ad i en t s) ; a n d g a m m ad i v e r s i t y f o r
a g e o g r a p h i c a r e a ( t h e p r o d u c t o f t h e a l p
h a d i v e r s i t y o f i t sc o m m u n i t i e s a n d t h e d
e g r e e o f b e t a d i f f e re n t ia t io n a m o n g s t t h
e m ) .
T h e s e m e a s u r e s a l l d e p e n d o n t h e s iz e o f
a r e a s a m p l e d ( B r o w n , 1 9 88 ).T h e y a l l a l so d
e p e n d o n g o o d t a x o n o m y t o d e t e r m i n e th e n
u m b e r o f s p ec ie si n v ol v e d. A n d t h e y a ll h a v e
a t h i r d f a c t o r i n c o m m o n - - a l l s p ec ie s a r e
t r e a t e da s t a x o n o m i c a l l y e q u i v a l e n t , o
r a s e q u a l u n i t s . Q u i t e a p a r t f r o m m a n yt h
e o r e t i c a l a n d p r a c t i c a l p r o b l e m s t h a t c
o n t i n u e t o a f fe c t t h e s p e c ie s c o n c e p ta n d
i ts a p p l i c a t i o n , is i t a p p r o p r i a t e f o r c o
n s e r v a t i o n p u r p o s e s t o r e g a r d a lls p e ci es
a s e q u a l i n t h is m a n n e r ? T o a c o n s e r v a t i o
n i s t , r e g a r d l e s s o f r e la t i v ea b u n d a n c e ,
is Welwitschia e q u a l t o a s p e c i e s o f Taraxacum? I s t h
e p a n d ae q u i v a l e n t t o o n e s p e ci e s o f r a t? A
t k i n s o n (1 98 9) a n s w e r e d t h i s q u e s t i o n i nt
h e f o l l o w i n g w a y : g i v e n t w o t h r e a t e n e d t
a x a , o n e a s p ec i es n o t c l o s e lyr e l a t e d t o o t
h e r l i v in g s p e c ie s a n d t h e o t h e r a s u b s p e c
i e s o f a n o t h e r w i s ew i d e s p r e a d a n d c o m m o
n s pe ci es , i t s e e m s r e a s o n a b l e t o g iv e p r i o
r it y t o t h et a x o n o m i c a l l y d i s ti n c t fo r m . S
o w e n o w e x p e c t st ill m o r e o f t a x o n o m i s t s -
-n o t o n l y m u s t t h e y b e a b l e to t e ll v a r i o u s
t a x a a p a r t , b u t t h e y m u s t a l s o b ea b l e t o q
u a n t i f y t h e i r d i s t i n c t n e s s .
S y s t e m a t i c s o f f e r s t w o p o s s i b i l i t i e
s t o m e a s u r e t a x o n o m i c d i s t i n c t n e s s :s i
m i l ar i ty m e a s u r e s ( p h e n e ti c o r g e n e t i c d
i st a nc e ) a n d g r o u p m e m b e r s h i p( c la d i s ti c
r e l a ti o n s h i p s , o f t e n i n t e r p r e t e d i n t e
r m s o f a n c e s tr y ) . T w od e c a d e s o f d e b a t e h a
v e l ed t o a n a l m o s t u n i v e rs a l a c c e p t a n c e o
f c la d is t icm e t h o d s i n p r e f e r e n c e to t h o s e
b a s e d o n s i m i l a r it y . A c c o r d i n g l y , w e b a
s e o u ra p p r o a c h e x c l u s i v e l y o n c l a d i s t i
c p r o c e d u r e s .
C l a d i s t i c c l a s s i f i c a t i o n s a r e e x p r e
s s e d a s h i e r a r c h i e s . O n e p o s s i b l ea p p r o
a c h t o m e a s u r i n g t a x o n o m i c d i s t in c t n e ss
is to w e i g h t e q u a l l y g r o u p s a tt h e s a m e t a x
o n o m i c r a n k w i t h r e s p e c t t o t h e a d d i t i v e
w e i g h t s o f t h e i rt e r m i n a l t a x a ( F ig . 1 ). T
h i s is a c h i e v e d b y g i v i n g t h e t e r m i n a l m e
m b e r s o f t h eg r o u p o f l o w e s t r a n k e a c h a w e
i g h t o f o n e, a n d t h e n g i vi n g t h e s i st er g r o u
p t o
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238 R. I . Van e-W right C. J . Hum phries P. H. W il l iams
W P1 6 2 5
1 6 2 5
2 1 2 5
4 2 5
8 50T 16 100
F i g . 1 . E q u a l w e i g h t i n g W ) a p p l i e d t o s
is t e r g r o u p s . C o l u m n P g iv e s t h e p e r c e n t a
g ec o n t r i b u t i o n f o r e a c h t e r m i n a l t a x o n
t o t h e t o t a l d i v e r s it y . S e e t e x t f o r e x p l
a n a t i o n .
t h i s g r o u p a n e q u a l a g g r e g a t e s c o r e , a
n d s o o n u p t h e t r e e . T h i s r e s u l t s i ne v e r y
s i s t e r g r o u p h a v i n g e q u a l a g g r e g a t e w e i
g h t , a n d t h e r e b y g i v e s h i g hr e la t iv e w e i g
h t t o t a x o n o m i c a l l y d is t in c t s p ec ie s s u c h
as t h o s e c o m p r i s i n gm o n o b a s i c g e n e r a o r f
a m i li es ).
H o w e v e r , t h is a p p r o a c h h a s t h e u n d e s i r
a b l e e ff ec t o f o v e r w e i g h t i n g b a s a lt a x a :
w e i g h t a c c o r d e d t o a s t e m s p e ci es w i ll b e e
q u a l l e d o n l y b y s u m m i n gi n d i v i d u a l w e i g
h t s o f al l t h e s u b o r d i n a t e t e r m i n a l t a x a
. A s a r e su l t,t a x o n o m i c r a n k o v e r w h e l m s th
e n u m b e r o f sp ec ie s. F o r e x a m p l e , t h e t w ok n
o w n t u a t ar a s D a u g h t e r y et a l . 1 9 9 0 ) w o u l d
h a v e a s c o r e e q u a l t o t h ec o m b i n e d w e i g h t
o f a ll 6 8 0 0 s n a k e s, l iz a r d s a n d a m p h i s b a e
n i a n s t o w h i c ht h e t u a t a r a s a r e c o n s i d e r
e d t o f o r m t h e s i st e r g r o u p a m o n g s t l i v in g
re p til es ).W h i le w e c e rt a in l y a gr e e t h a t t u a t
a r a s a r e p a r ti c u la r l y i m p o r t a n t a m o n gr e
p ti le s b e c a u s e o f a n c i e n t d i v e r g e n c e o f t
h e i r l in e a g e , a n d s h o u l d s c o r eh i g h l y i n a
n y c o n s e r v a t i o n p l a n , t o a c c o r d t h e m e q u
a l s t a tu s t o 6 8 0 0 o t h e rl iv i ng r e p ti le s i s u n
r e a s o n a b l e i n t e r m s o f s u s t a i n i n g d i v e r
s i t y . W h a t w e n ee dis s o m e l o g i c a ll y d e f e n s
i b le m e a s u r e s e n si ti v e t o b o t h t a x o n o m i c
r a n k a n dn u m b e r o f sp e ci es .
T h e a p p r o a c h w e s u g g e s t h e r e i s b a s e d o
n a t t e m p t s t o m e a s u r e t h ea m o u n t o f i n f o r
m a t i o n c o n t a in e d o r c o n v e y e d i n h i e r a r ch
i c c l a s s i f i c a t i o n s .T h i s c o m p l e x s u b j e
c t h a s r e c e n t ly b e e n r e v ie w e d b y M i c k e v i c
h a n d P l a t n i c k1989). O u r m e t h o d i n v o l v es a v
a r i a n t o f t w o o f th e i r p r i m a r y i n f o r m a t i
o n
m e a s u r e s , te r m s a n d c o m p o n e n t s . A s d is
c u ss e d b y M i c k e v i c h a n d P l a tn i c k ,a p p l i c
a t io n o f t h es e m e a s u r e s h a s t h e r e s ul t th a t
t h e t o ta l a m o u n t o fi n f o r m a t i o n c o n v e y e d
w i t h in a h i e r a r c h i c c l a ss if ic a t io n v a r i es
w i t ht o p o l o g y , e v e n f o r t h e s a m e n u m b e r o
f t e r m i n a l ta x a .
T o d e m o n s t r a t e t h e m e t h o d , c o n s i d e r a
p e c t i n a te c l a d o g r a m f o r fiv e t ax a ,A - E F ig .
2). F r o m t h is c l a d o g r a m it is p o s si b le to m a k e
f o u r t a x o n o m i c
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ystematics and conservation evaluation 239
Q W P
A 4 3 5 1 10 7
B 4 3 5 1 10 7
C 3 4 67 1 33 14 3
D 2 7 2 21 4
E 1 14 4 42 9
T 14 32 7 9 33 100
F i g . 2 . D e r i v a t i o n o f i n d e x o f ta x o n o m i
c d is t in c t n e s s, o r w e i g h t. T h e e x a m p l e is b
a se d o n af u ll y p e c t i n a t e c l a ss i f ic a t i o n f
o r fi ve te r m i n a l t a x a , A - E . C o l u m n I i n d i c
a te s t h e n u m b e r o fg r o u p s t o w h i c h e a c h t e r
m i n a l t a x o n b e l o n g s w i t h in t h e s y s t e m , t
h e se n u m b e r s b e i n g t h e b a s icm e a s u r e o f ta x
o n o m i c i n f o r m a t i o n . C o l u m n Q g i v e s t h e q
u o t i e n t o f t h e t o ta l i n f o r m a t i o n f o rt h e w
h o l e g r o u p ( i n t h i s e x a m p l e , T I = 1 4) d i v i
d e d b y t h e i n f o r m a t i o n s c o r e f o r e a c h t e r
m i n a l .C o l u m n W g i v e s t h e s t a n d a r d i s e d
weight f o r e a c h t e r m i n a l , t h e Q - v a l u e s f o r
e a c h t e r m i n a lt a x o n h a v i n g b e e n d i v i d e d
b y t h e l o w e s t Q - v a l u e ( in t h i s c a s e , Q m l, =
3 5 ). C o l u m n P g i v e s t h ep e r c e n t a g e c o n t r i
b u t i o n f o r e a c h t e r m i n a l t a x o n t o t h e t o t
a l d i v e r s i ty , i n te r m s o f t h ea g g r e g a t e v a
l u e s f o r Q o r W . T h e t o t a l s r o w ( T ) g i v e s t h
e a g g r e g a t e s c o r e s u n d e r I , Q , W a n d P .S e e
t e x t f o r e x p l a n a t i o n .
s t at e m e n ts a b o u t A it b e lo n g s t o g r o u p s A
B , A B C , A B C D a n d A B C D E ) ,t h e s a m e fo r B , t h r
e e f o r C A B C , A B C D , A B C D E ) , t w o fo r D A B C D ,A
B C D E ) , a n d o n e f o r E A B C D E ) . T h i s g i ve s a to
t a l o f 14 4 + 4 + 3 + 2 + 1)i n f o r m a t i v e s ta t e m e n
t s a b o u t t h i s t a x o n o m i c g r o u p i n g F i g. 2 ,
c o l u m n I).Th e v a lu es 4 , 4 , 3 , 2 , 1) co u ld b e u sed
d i r ec t ly a s we ig h t s . H o w ev e r , t h i s n o to n l y
g iv e s b a s a l t a x a l o w w e i g h ts , a n d v i c e v e r
s a - - t h e o p p o s i t e o f w h a t i sd e s i r e d - - b u
t i t f a i l s a l s o t o m a k e t h e i n d i v i d u a l v a l
u e s d i r e c t l y r e f l e c t t h ep r o p o r t i o n t h a
t e a c h ta x o n c o n t r i b u t e s t o th e to t a l di v e
rs i ty i n f o r m a t i o nc o n t e n t ) o f t h e g r o u p .
B o t h o f th e s e p r o b l e m s a r e o v e r c o m e b y d i
v id i n g e a c ht e r m i n a l s c o r e i n t o t h e t o t a l
- - f o r A , 1 4 / 4 g i v e s 3 . 5 a n d s o o n - - 3 . 5 , 4 .
6 7 , 7a n d 1 4. T h e s e q u o t i e n t s a r e t h e b a s i c
ta x i c w e i g h t s F i g . 2 , c o l u m n Q ). F o r an u m b
e r o f r e a s o n s i n c l u d i n g e a s e o f c a l c u l a t
i o n b y o t h e r m e a n s ) , i t isc o n v e n i e n t t o s t
a n d a r d i s e t h e s e v a l u e s b y d i v i d i n g b y t h
e l o w e s t v a l u et h r o u g h o u t t h u s g i vi n g t h e
lo w e s t r a n k i n g t a x o n a s co r e o f o n e - - F i g .
2,c o l u m n W ) . W e c a n a l s o e x pr e ss t h e c o n t r i
b u t i o n o f e a c h a s a p e r c e n t a g eF ig . 2 , co lu m
n P) .S u c h t e r m i n a l s c o r e s o r w e i g h t s , b a s
e d d i r e c t l y o n a n i n f o r m a t i o n
m e a s u r e o f t h e h i e r a r c h i c c l a ss i fi c a ti
o n , h a v e t h e t y p e o f p r o p e r t i e s w e d e s ir e
.T a x o n o m i c e q u i v a l e n c e e q u a l r a n k ) re s u
lt s i n th e s a m e s c o re . T a x o n o m i c
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24 R. I . Vane-W righ t C . J . Hum phr ies P . H . W i l l
iams
1 1 20 2 1 12 .5 ~ ~ 2 1 14 .31 1 20 2 1 1 2 . 5 | . ~ 2 1 14
.31 1 20 1 2 25 C ~ 2 1 14 .31 1 20 1 2 25 1 2 28.61 1 20 I~ 1 2 25
1 2 28.65 5 7 8 8 72 1 1 6 .7 ~ 2 1 1 6 . 7 ~ 3 1 1 0 .52 1 1 6 .7
~ 2 1 1 6 . 7 ~ 3 1 1 0 .52 1 16.7 1 2 33.3 2 1.5 15.82 1 16 7 2 1
16 7 1 3 31 .61 2 33 .3 2 1 16 .7 F 1 3 31 .69 6 9 6 10 9.5
G
~ 2 1 20 ~ 3 1 1 2 . 5 ~ 3 1 1 3 . 32 1 20 3 1 12 .5 3 1 13 .32
1 20 2 1.5 18.7 3 1 13.32 1 20 2 1.5 18.7 2 1.5 202 1 20 H 1 3 37.5
] 1 3 4010 5 11 8 12 7.5
3 1 1 5 4 3 1 1 4 3 4 1 1 0 7
2 1 5 2 3 1 3 1 1 4 3 3 1 3 1 4 3
2 1 5 2 3 1 3 1 1 4 3 2 2 2 1 4
2 1 6 2 3 1 K 1 3 4 2 8 L 1 4 4 2 91 2 6 5 1 3 7 1 4 9 3
Fig. 3. Th e twelve cladistic topo logie s possible for five
terminal taxa designated A -L ,corre spon ding to M ickevich a nd P
latnick, 1989). Ag ainst each diagram the lef t -hand colum ngives
the informat ion va lue for each termina l t axon and the to ta l
inform at ion for the wholetopolo gy; the central colum n gives the
s tandardised w ight for each terminal taxon and theaggrega te to
ta l ; and the r ight -hand column gives the percentage cont r ibut
ion for eachterminal taxo n to the total diversi ty cf . colum ns I
, W a nd P in Fig. 2). See text for explanat ion.d i s t in c t n e
s s r e s u l ts i n a g r a d e d v a r i a t i o n o f s c o r e
- - n o t a ll sp e c ie s a r e eq u a l .F o r u n e q u a l s is
te r g r o u p p a ir s , t h e g r o u p w i t h t h e h i g h e r
n u m b e r o f t e r m i n a lt a x a w i ll a l w a y s h a v e a
h i g h e r a g g r e g a t e s c o r e t h a n t h e s m a l l e r
g r o u p .A g g r e g a t e d d i v e r s i ty s c o r e s o f m o
n o p h y l e t i c ) g r o u p s o f s p e ci es w il l t h u sd e
p e n d o n b o t h r a n k a n d n u m b e r .
W e h a v e a l r e a d y n o t e d t h a t t h e a m o u n t o
f i n f o r m a t i o n i n a c l a s s i f i c a t i o n
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ystematics and conservation evaluation 241d e p e n d s o n i t
s t o p o l o g y . T h e r e a r e 1 2 d i f f e r e n t t o p o l
o g i e s a v a i l a b l e t od esc r ib e a l l p o ss ib l e c l
ad i s t i c r e l a t io n sh ip s f o r f i v e t ax a , i n c lu
d in g th e f u l lyp e c t i n a t e f o r m a l r e a d y d i s c
u s se d , a n d t h e t o t a l l y u n r e s o l v e d t r e e (
o r 'b u s h ') .A l l 1 2 a r e s h o w n i n F ig . 3 , w i t h i
n f o r m a t i o n v a l u e s , w e i g h t s a n d p e r c e n t
a g e sf o r e a c h . N o t e t h a t p o l y c h o t o m i e s d
o n o t o f f e r a n y d i ff ic u lt ie s , a n d t h a t u s eo
f p e r c e n t a g e s e n a b l e s t h e d if f e re n t r e s u
l ts t o b e c o m p a r e d . N o t e a l s o t h a tt h e t o t a
l i n f o r m a t i o n m e a s u r e i s d i f f e r e n t f o r m
a n y , b u t n o t a l l , o f t h eto p o lo g ie s .
T h e f a c t t h a t s o m e i n f o r m a t i o n m e a s u r
e s a r e t h e s a m e i s n o t ju s t ac u r i o s it y . C o m
p a r e F i g . 3 (A ) a n d 3 (G ). A l t h o u g h F i g . 3 ( G
) h a s e x a c t l y t w i c et h e i n f o r m a t i o n c o n t
e n t o f F i g . 3(A ), t h e w e i g h t s a n d p e r c e n t a
g e s a r e a l l th es a m e . W h i l e t h i s m a k e s s e n s
e b e c a u s e a l l t e r m i n a l t a x a h a v e e q u i v a l
e n t r a n k ,c l e a rl y t h e r e is s o m e i n f o r m a t i
o n t h a t w e h a v e f ai le d t o e n c o d e .
M a y ( 19 90 ) h a s p r o p o s e d a m o d i f i c a t io n o
f o u r i n d ex w h i c h t a k e s a c c o u n to f t h e n u m b
e r o f l i n e a g e s a r i s i n g a t e a c h n o d e . A l l n
o d e s a r e s c o r e d b yc o u n t i n g t h e n u m b e r o f
b r a n c h e s a r i s in g f r o m t h e m . E a c h t e r m i n
a l t a x o n ist h e n s c o r e d b y s u m m i n g t h e v a l u
e s fo r a ll s u b t e n d i n g n o d e s . S t a n d a r d i s e
di n d i c e s a r e t h e n d e r i v e d b y d i v i d i n g t h
e s c o r e f o r e a c h t e r m i n a l , i n t u r n , i n t ot
h e m a x i m u m s c o r e . T h i s p r o c e d u r e p r o d u c
e s i d e n t i c a l r e s u l t s t o o u rm e t h o d w h e n a
p p l i e d t o f ul ly r e s o lv e d o r c o m p l e t e ly u n r
e s o l v e d h i e r a rc h i e s ,b u t d o es a f f ec t t h e v
a lu es f o r p a r t i a l l y r e so lv ed t r ee s . Th i s i s
an in t e r e s t in gs u g g e s t i o n , a n d c o u l d b e o f
s p e c i a l v a l u e w h e n d e a l i n g w i t h t h e v e x e
dp r o b l e m o f e v a l u a t i n g s u b s p e c i e s , u t i
li s in g it a s a r e d u c i n g f a c t o r f o rt e r m i n a l
p o l y c h o t o m i e s o f s t r i c t l y a l l o p a t r i c t
a x a .
H o w e v e r , w e c o n s i d e r t h a t a m o r e r a d i c
a l s o l u t i o n i s a l m o s t c e r t a i n l yn e e d e d .
C o n s i d e r F i g . 3( G ) a g a in , i n w h i c h t h e g r o
u p o f f iv e sp e c ie s is j u s td i v i d e d i n t o t w o s
u b g r o u p s . F o r c o n s e r v a t i o n p u r p o s e s , w
e m i g h t w e l l w i s ht o g i v e f ir st p r i o r it y t o c
o n s e r v i n g o n e m e m b e r o f e ac h . W e t h u s c o n
s i d e rt h a t s o m e p r o c e d u r e f o r s a m p l i n g '
a cr o s s ' t h e c l a d o g r a m , t o e n s u r e t h a t a sw
i d e a r a n g e o f c la d e s a s p o s s ib l e is s a m p l e
d i n a d d i t i o n t o i d e n t i f y i n g h i g h l yd i s t
i n c t g r o u p s , i s d e s i r a b le . W e a r e c u r r e n
t l y e v a l u a t i n g o t h e r i n f o r m a t i o nm e a s u
r e s o f ta x ic d i v e r s it y w h i c h m a k e u s e o f th i
s a p p r o a c h ( W i ll ia m s ,H u m p h r i e s V a n e - W r
i g h t , i n p re p .) . O u r m a i n p u r p o s e h e r e is t
o e s t ab l is ht h e id e a t h a t m e a n i n g f u l t a x ic
d i v e r s it y m e a s u r e s c a n b e d e v e l o p e d , a n
d t h a tt h e y c a n b e a p p l i e d t o a s s e s s m e n t o
f c o n s e r v a t i o n p r i o r it i e s .
A R E A L L P L A C E S E Q U A L ?W e n o w c o n s i d e r t h
e s e c o n d s y s t em a t i c a sp e c t o f th e p r o b l e m
o f c o n s e r v i n gb i o d i v e r s i t y - - d i s t r i b u
t i o n a n d b i o g e o g r a p h y o f t a x a. T h e c o n s e
r v a t i o n o fn a t u r e i s i n t i m a t e l y a ff e c te d
b y d i s t r i b u t i o n a n d b i o g e o g r a p h y . I f a t
a x o n i sr e s t r ic t e d t o o n e p l a c e o r r e g i o n ,
it is t h e r e t h a t y o u m u s t p r e s e r v e i t, o r i t
is
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4 R. L Van e-Wrigh t C . J . Hum phr ies P . H . W i l l
iams
f r o m t h e re t h a t y o u m u s t n o r m a l l y o b t a i
n i t if y o u p l a n e x s i t u c o n s e r v a t i o no r i n t
r o d u c t i o n .
M c N a u g h t o n (1 98 9) h a s o b s e r v e d th a t w e h
a v e t o ' d e t e rm i n e w h a t s h o u l db e c o n s e r v e
d a n d h o w it is to b e c o n s e r v e d . A c r i t i c a l p
l a c e s s t r a t e g y . . , c o u l da c c o m p l i s h t h i
s o b j e c t i v e ' . M c N a u g h t o n i s a n e c o l o g i s
t , a n d h i s ' c r i t i c a lp l a c e s ' r e f e r t o r e p
r e s e n t a t i v e e c o s y s t e m s . A s s y s t e m a t i s
t s w e t h i n k i n s t e a do f a r e as o f e n d e m i s m , o
r cr it ic a l f a u n a s a n d f lo r a s f o r p a r t i c u l a
r t a x o n o m i cg r o u p s .
' C r i t i c a l f a u n a s a n a l y s i s ' w a s i n t r o
d u c e d b y A c k e r y a n d V a n e - W r i g h t( 19 84 ) i n
a n a c c o u n t o f m i l k w e e d b u t t er f li e s o f th e
w o r l d . T h e d i s t r i b u t io n so f a ll 15 7 sp e c ie s
t h e n r e c o g n i s e d ( m a n y o f w h i c h a r e p o l y t
y p i c ) w e r ee x a m i n e d o n a g l o b a l b a si s, d i v
i d e d a m o n g s t m o r e t h a n 3 5 0 c o u n t r i e s , r e
g i o n sa n d i s l a n d s . S p e c ie s r i c h n e s s v a r i
e d i n th e s e s a m p l e s f r o m o n e s p e c i e s u p t oa
b o u t 3 5. S o m e s pe c ie s w e r e n a r r o w e n d e m i c
s , o c c u r r i n g i n o n l y o n e s a m p l ea r e a , b u t
m o s t w e r e w i d e s p r e a d . G i v e n t h a t t h e i d e
a o f in d i v i d u a l l yc o n s e r v i n g r e p r e s e n t a
t i v e p o p u l a t i o n s o f al l 1 57 s p e c ie s w a s c o
n s i d e r e du n r e a li st ic , th e f o l l o w i n g q u e s
t i o n w a s a s k e d : w h a t i s t h e m i n i m u m s e t o
fa r e as , o r f a u n a s , w h i c h w o u l d c o n t a i n a t
l e a st o n e p o p u l a t i o n o f e v er ys p e c ie s ? T o c
o v e r a ll 1 5 7 sp e c ie s , d i s t r i b u t e d t h r o u g
h 3 5 0 s a m p l e a r e a s , t h ea n s w e r w a s f o u n d t
o b e 3 1 (F i g . 4 ). P a r s o n s ( 1 98 9 ) h a s s i n ce a d
d e d a 1 5 8 t hs p ec i e s , a n ew a r a n t i c a f r o m N e
w B r i ta i n , o n e o f th e a r e a s i d e n t if ie d .H o w
w a s t h is s h o r t li s t o f 31 f a u n a s d e t e r m i n e
d ? A ll f a u n a s w i t h o n e o rm o r e e n d e m i c w e re
p l a c ed i n s e q u en c e , t h o s e w i th t h e h i g h e s
t n u m b e r o fe n d e m i c s fi rs t, t h o s e w i t h o n l y
o n e e n d e m i c la s t ( th i s p r o c e d u r e i s j u s t i
fi a b l ei n te r m s o f th e r e c o m m e n d a t i o n s p u t
f o r w a r d i n R a v e n (1 98 0); ' gi v e p r i o r i t yt o a
r e a s c o n t a i n i n g t h e ri c h e st a n d m o s t h i g h
l y e n d e m i c b i o t a . . . ' ) . T h eh i g h e st n u m b e
r o f e n d e m i c s ( nin e ) w a s f o u n d o n t h e i sl a n
d o f S u l aw e s i. T h isi s la n d h a s p o p u l a t i o n s
o f 2 4 o t h e r , m o r e w i d e s p r e a d , m i l k w e e d b
u t te r fl ie s .
T h u s a p r o g r a m m e t o c o n s e r v e t h e e n t i r
e m i l k w e e d b u t t e r f l y f a u n a o fS u l a w e s i c
o u l d p r e s e rv e r e p r e s e n t a t i v e s o f 3 3 /1 5
8, o r 2 1 o f t h e w o r l ds p ec ie s . T h e n e x t h i g h e
s t n u m b e r o f e n d e m i c s w a s f o u n d o n B i a k , a
n i s l a n di n I ri a n B a y , N e w G u i n e a . I t h a s a t
o t a l o f 1 8 sp e ci es , f o u r o f w h i c h a r e n o tf o u
n d a n y w h e r e e ls e. O f t h e o t h e r 1 4, f iv e o c c u
r o n S u l aw e s i , b u t n i n e d on o t . S o c o n s e r v i
n g S u l a w e s i + B i a k c o u l d p r e s e r v e a t o t a l
o f 3 3 +4 + 9 ( = 46 ) s e p a r a t e m i l k w e e d b u t t e
rf l y s p e c i e s - - 2 9 o f t h e w o r l d f a u n a .F o r a
l l n a r r o w e n d e m i c s , 2 4 r e g i o n s a r e r e c o g
n i s e d a s e s s e n t i a l - - b u t t h e s e2 4 r e g i o n
s d o n o t i n c l u d e r e p r e s e n t a t i v e s o f al l 1
58 sp e c ie s . A m i n i m u m o fs e v e n f u r t h e r r e g i
o n s is n e e d e d t o c o m p l e t e t h e l i s t - - t h e 31
c r it ic a l f a u n a sf o r m i l k w e e d b u t t e r f l y c
o n s e r v a t i o n ( F i g . 4 ) .
C o l l i n s a n d M o r r i s (1 985 ), i n t h e i r I U C N
R e d D a t a B o o k , a p p l i e d cr it ic a lf a u n a s a n a
l y s i s t o a l l 57 3 w o r l d s p e c i e s o f s w a l l o w
t a i l b u t t e r f li e s t h e yr e c o g n i s e d . U n l i k
e m i l k w e e d b u t t er f li e s, s w a l l o w t a i ls a r e
s t r o n g l y r e p r e s e n -
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Systematics and conservation evaluation 243
. . . . . - u ~ ~ \ j r
/ .
. . . . . . . . . . . . . ~ . . . . . ' > ~ ~ : . . . . . . .
t . . . . . . . . ~ - - - - ~ B - U ~ - - - ' ~ - + - . . . . .. .
. . . . . . . : . . . . .. . . . . ~ . . . .. . . .. 1 . . . .. .
.. . ~ - - - t - - ~ i - r ~ ' ~ . . . . . . . .. .
Fig. 4. The m inimum set of faunal regions, or critical faun as,
that includes at leas t onerepresentative population of all know n
spe cies of m ilkw eed butterflies (Lepidoptera:Danainae)i Cou
ntries or islands listed in italics include species only found in
those particularregions; the list is not in priority order. 1, Com
oro Islands; 2 Seyche lles; 3 M auritius; 4, Zaire;5 Sri Lanka; 6
southern India;7, Ne pal; 8, Burm a; 9, Sumatra; 10, Java; 11
Borneo; 12, Luzon;13, Negros; 14, Mindanao; 15, Sulawesi; 16,
Sumbawa; 17, Sumba; 18, Flores; 19, Timor; 20,Seram; 21, Biak; 22,
New Guinea; 23, New Ireland; 24, New Britain; 25, Guadalcanal; 26,
SanCristobal; 27, N ew Caledonia; 28, Hispaniola; 29, Cuba; 30, Co
sta Rica; 31, Bolivia. It shouldbe noted that the New W orld has a
danaine fauna o f 14 species, the Afrotropical Region has21, and
the Indo-Pacific has 126 (world total 158 species; original data
from Acke ry and Vane -W right (1984) and Parsons (1989)). Over 350
separate islands, regions or countries w eresurveyed; the s i te
conservat ion efficiency (Pres sey Nic holls, 1989a), for
minimumrepresentation, is better than 0.91.t e d in P a l a e a r c
t ic , N e a r c t i c a n d N e o t r o p i c a l r e g i o n s a
s w e l l a s O l d W o r l dt ro p i c s. A l t h o u g h n o t b
a s e d o n s t r ic t ly c o m p a r a b l e g e o g r a p h i c a
l u n i ts , th et o p O l d W o r l d t r o p i c a l e l e m e n
t s i n th e i r li st o f 5 1 c r i ti c a l f a u n a s f o rs w
a l l o w t a i l s a r e c l o s e ly m a t c h e d b y t h e f ir
st p a r t o f t h e m i l k w e e d l is t( T a b l e I ) - - - s
tr o n g l y s u g g e s t i v e th a t i t m a y i n d e e d b e p
o s s i b l e to r e c o g n i s ea r e a s o f e n d e m i s m c o
m m o n t o a n u m b e r o f d if fe re n t g r o u p s ( s o -c a
ll ed ' h o ts p o t s ' ) .
A c l o s e l y s i m i la r s t e p -w i s e a p p r o a c h t
o th e c ri ti c a l fa u n a s p r o c e d u r e h a sb e e n d e
v e l o p e d i n d e p e n d e n t l y b y M a r g u l e s a n d h
is c o - w o r k e r s ( M a r g u l e s e tal . 1 9 88 ; P r e s s
e y N i c h o l l s , 1 9 8 9 a, b ), b a s e d o n a n o r ig i n
a l id e a o fK i r k p a t r i c k ( 1 9 83 ) a n d K i r k p a t
r i c k a n d H a r w o o d ( 19 83 ). M a r g u l e s a n d h i sc
o l l e a g u e s a r e c o n c e r n e d w i t h e v a l u a t i o
n p r o c e d u r e s w h i c h w il l l e ad t om a x i m u m e f
fi ci en c y in e s t a b l i s h m e n t o f n a t u r e r e s e
rv e ' n e t w o r k s ' t oc o n s e r v e a l l o r m o s t s p e
c i e s o r l a n d s y s t e m s w i t h i n a g i v e n a r e a
.
I n b o t h t h e n e t w o r k a p p r o a c h a n d t h e c r
it ic a l f a u n a s p r o c e d u r e , a lls p e c ie s ar e t r
e a t e d a s t a x o n o m i c a l l y e q u a l . A m i n i m a l
l is t is o b t a i n e d b yr e c o g n i s i n g t h a t a ll n a
r r o w e n d e m i c s ( s pe c i e s r e s t r ic t e d t o j u s
t o n e o f th es a m p l e a r e a s ) h a v e t o b e i n c l u d
e d i f a ll t h e s p e c i e s a r e t o b e t a k e n i n t
o
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244 R. L Vane-W right, C. J. Hum phries, P. H. Williams
T A B L ESome Crit ical Faunasa for Milkweed and Swallowtail
ButterfliesThe table lists the first four regions in each of the
original critical faunasanalyses of Ackery an d Vane-W right (1984)
for the Danainae, an d o f Collins andM orris (1985) for the
Papilionidae. No te th at the m ilkweed butterflies are m
ainlydistributed in the Ind o-A ustr alian tropics, whereas the
swallowtails are far moreevenly distributed on a global b asis. N
ote also th at Sulawesi, Biak and N ewG uine a are all effectively
included in the political division Indonesia, and tha tMindanao is
the main southern island of the Philippines (see also text).
M ilkweeds Dana inae) Swa llowtails Papilionidae)1. Sulaw esi (9
endem ics) 1. Indon esia (53 endem ics)2. Bia k (4) 2. Ph ilippin
es (21)3. M ind an ao (3) 3. Ch ina (15)4. New G uin ea (3) 4.
Brazil (11)To tal: 69 species Total: 296 species(over 40 world
total) (over 50 world total)
a All 31 critical faun as for the D ana ina e are listed,
unordered, in the legend toFig. 4; the 51 critical faun as list for
the Papilionidae, a s given by Collins an dM orris (1985), is com
pleted thus: 5, M adagas car; 6, India; 7, M exico; 8, Taiw an;9,
Malaysia; 10, Papua New Guinea; 11, USA; 12, Cuba; 13, Ecuador;
14,Co lom bia; 15, Australia; 16, A nd am an s an d Nicobars; 17,
Jamaica; 18, Zaire; 19,Cam eroon ; 20, USSR; 21, Tanz ania; 22,
French Gu iana; 23, Japan ; 24, Can ada;25, Sri La nk a; 26, Haiti;
27, New Caledonia; 28, Co m oro Islands; 29, Peru; 30,Bolivia; 31,
Venezuela; 32, Ga bo n; 33, Bhutan; 34, G han a; 35, Solomon
Islands;36, Afghanistan; 37, South Africa; 38, lran; 39, Vanuatu;
40, Mauritius; 41,Rr un ion ; 42, Fiji; 43, W estern Sam oa; 44,
Burma; 45, Laos; 46, Hon du ras ; 47,Argentina; 48, Uganda; 49,
Ethiopia; 50, Mozambique; 51, I ta ly. Siteconservation efficiency
(Nicholls & Pressey, 1989a) for minimal representationo f
Papilionidae: 0.7 (abo ut 170 countries analysed by Collins an d M
orris (1985)).
a c c o u n t . T h e n e t w o r k o r c r it ic a l f a u n a
s l i s t -s e q u e n ce i s h i g h l y d e p e n d e n t ,t h e
r e fo r e , o n a c c o r d i n g , a p r i o r i , h i g h v a lu
e s to e n d e m i c s . G i v e n t h a t w e m a yb e a b l e t o
s a v e o n l y a p o r t i o n o f t h e E a r t h 's b i o t a ,
a n d t h a t m a n y e x i s t i n gr e s e rv e s a r e n o t i n
o p t i m a l a r e a s ( a n d o f t e n l a r g e l y d u p l i c
a t e e a c h o t h e r , i nt e r m s o f s p e c i e s r e p r e
s e n t a t i o n ; c f. P r e s s e y & N i c h o l l s , 1 9
8 9 a, b ), t h e c r i t ic a lf a u n a s o r n e t w o r k s t e
p - w i s e p r o c e d u r e s d o n o t o f f e r a s u f f ic i
e n tl y f le x ib l es t r a t e g y a t a g l o b a l l ev el . N
o n e t h e l e s s , b o t h p r o c e d u r e s a r e b a s e d o
n a b a s icg u i d i n g p r i n c i p l e - - w h i c h w e ca ll
c o m p l e m e n t a r i t y .
I n c a r r y i n g o u t a s i m p l e c r i ti c a l f a u n a
s a n a l y s i s , o n c e t h e f i rs t c h o i c e h a sb e e n
m a d e , a ll f u r t h e r c o n s i d e r a t i o n s o f sp e c
i es i n c l u d e d w i t h i n t h a t r e g i o na r e e l im i
n a t e d . T h e s e c o n d a r e a is t h e n d r a w n f r o m
t h e t a x o n o m i cc o m p l e m e n t o f t h e f i r s t - -
t h e r e m a i n i n g f a u n a w i t h th e h ig h e s t n u m b
e r o fe n d e m i c s , a n d a ll a d d i t i o n a l n o n - e n
d e m i c s t h a t t h e a r e a h a p p e n s t o c o n t a i n
.
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246 R. L Van e-Wright C. J. Hum phries P. H. W illiams
W R 1 R 2 R 31
B
C 1 3
D 2
E 4
T 9 3 3 3 4 3 7 3P 1 3 5 4 6 7 8P2 2 2 1 1
Fig . 5 . Theore t ica l p r io r i ty a rea ana lys is , based
on top o lo gy and tax ic we igh t ing g iven inF ig . 2 . Three o
f the f ive te rmina l taxa o ccu r in each o f th ree a reas , R l
-R 3 , acco rd ing to theth ree -co lum n m at r ix a t the r igh t
. Co lum n W g ives the tax ic we igh ts (c f . F ig . 2) . R ow T
g ivesthe to ta l (aggrega te ) sco res fo r a ll f ive taxa , and
fo r each o f the th ree reg ions ; row P1 g ives thepercen tag e d
ive rs i ty sco res fo r each o f the th ree reg ions a t the f ir
st s tep , ind ica t ing tha t R3 i st h e t o p - p r io r i t y r
e g io n ; r o w P 2 g iv e s t h e p e r c e n t a g e d iv e r s
it y s c o r e s f o r t h e r e m a in in g tworeg ions wi th
respec t to the taxa complementary t o t h o s e o c c u r r i n g
i n R3 , a n d i n d i c at e s t h a tR I i s the second p r io r
i ty ; f ina lly, the fac t tha t the h ighes t va lues o f the f
ir st tw o s teps sum to100 indicates tha t the analysis is com
plet e (Pressey & Nic ho lls ' s i te eff ic iency value: 0
.33) .
See tex t fo r exp lana t ion .a d d i t i o n a l t a x ic d i
v e rs i ty , w h e r e a s R e g i o n 2 of fe rs o n l y o n e u
n i t ( 1 1 ) .T h u s , d e s p i t e b e i n g l e a s t d i v e
rs e i n t e r m s o f w e i g h t e d s c o re s , R e g i o n 1
is t h es e c o n d p r io r it y . A n d n o t e n o w t h a t R e
g i o n s 3 + 1 g iv e a c o m b i n e d o rc o m p l e m e n t a r
y t a x i c d i v e r s it y s c o r e o f ( 7 8 + 2 2 ) = 1 0 0 .
T h i s in d i c a t e s t h a tt h e w e i g h t e d s t e p - w i
s e a n a l y s is i s c o m p l e t e .
B y s e c u r i n g R e g i o n s 1 a n d 3 , a l l f iv e t a x
a a r e r e p r e s e n t e d . B u t t h isis n o t t o s a y t h
a t R e g i o n 2 is o f n o v a l u e, o r is u n i m p o r t a n
t i n o t h e rr e s p e c t s - - m e r e l y t h a t i n t h i s
i n i t i a l a n a l y s i s i t i s n o t r e q u i r e d t o a c
h i e v e1 0 0 r e p r e s e n t a t i o n o f t h e g r o u p in q
u e s ti o n .
F i g u r e 6 s h o w s t h e t o p o l o g y o f a c u r r e n
t c l a s s if i c at i o n o f th e b u m b l e b e e so f t h e
Bo m bus s ibiricus group ( Wi l l i am s , i n p r e ss ) . Tax ic
we ig h t f o r eacht e r m i n a l t a x o n c a n b e r e a d , f
r o m t h e b o t t o m l i n e , a g a i n s t t h e n o d es u b
t e n d i n g t h e t e r m i n a l t a x o n i n q u e s t i o n .
W i l l i a m s h a s m o d i f i e d h i sW O R L D M A P c o m p
u t e r p ro g r a m t o ca r ry o u t p ri or it y a re a a n aly
s is o n th eg r o u p . R e p r e s e n t a t i v e s o f t h e 43
s p e c ie s o c c u r i n 1 2 0 o f t h e 2 5 0 e q u a l - a r e
ag r i d - s q u a r e s e m p l o y e d i n t h e m a p p r o j e
c t i o n ( W i l l ia m s , in p r e ss ; F i g . 7).
A l t h o u g h t h e g r e a t e s t n u m b e r o f sp e ci es
i n o n e a r e a o c c u r s i n t h e g ri d -s q u a r e c e n t
r e d o n E c u a d o r , t h e 10 s p ec ie s f o u n d i n th a t
s q u a r e c o n t r i b u t e
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S y s t e m a t i c s a n d c o n s e r v a t i o n e v a l u a
t i o n 247
L .
5 5 3 7 2 8 2 2 1 8 1 6 1 4 1 2 1 1 1Fig. 6. Cladogram topology
for the B o m b u s s i b i r i c u s g r o u p based on Williams,
in pressand unpublished)which forms the basis for the analyses show
n in F igs 7 and 8; the row o fstandardised taxic weights bottom)
applies to the term inal taxa arising at the co rrespondingnodes.
See also text for explanation.le ss t h a n 15 o f t h e t o t a l
w e i g h t e d d i v e r s it y s c o re , w h e r e a s th e G a
n s u a r e ao f C h i n a , w i t h n i n e ( m o r e h i g h - r
a n k i n g ) s p e c ie s i n t h e g r id , c o n t r i b u t e
sa l m o s t 2 3 . T h e h i g h e s t c o m p l e m e n t t o G a
n s u is t h e B i g H o r n r e g i o n o fN o r t h A m e r i ca
, w h i ch , a l t h o u g h i t h a s o n l y f o u r s p ec ie s,
c a n a d d m o r e t h a n15 w e i g h t e d d i v er s it y t o t
h e G a n s u t o ta l . T h e t h ir d c o m p l e m e n t is t h
e na d d e d b y t h e E c u a d o r s q u a r e , w i t h 1 4
(i.e. t h is m e a n s t h a t n o n e o f t h e l 0s p e c ie s i
n E c u a d o r i s f o u n d i n B ig H o r n o r G a n s u ) . T
h u s , t h e f ir st th r e es q u a r e s o u t o f t h e to t a
l o f 12 0 s c o r e 5 2 o f t h e w e i g h t e d d i v e r s it y
i n d e x fo rt h e g r o u p ( a n d a l s o h a p p e n t o i n c
lu d e m o r e t h a n h a l f o f t h e w o r l d s p ec ie s) .B
y t h i s m e t h o d , a l l s p e c i e s ( 1 0 0 d i v e r s i t
y s c o r e ) a r e r e p r e s e n t e d w i t h i n am i n i m a
l s e t o f 1 3 g r i d a r e a s , a r r a n g e d i n p r i o r i
t y s e q u e n c e ( F i g . 7 ) .
F i g u r e 8 p l o t s t h e s t e p -w i s e a c c u m u l a t
i o n o f w e i g h t e d d iv e r s it y , t o g e t h e rw i t h
t h e o r e t i c a l b o u n d s t h a t w o u l d d e s c r i b e
to t a l s y m p a t r y a t o n e p l a c e (i.e.
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248 R. L Vane-Wright C. J. Humphries P. 11. Williams
Fig . 7 . Sequ ence of c ri ti ca l f auna s for the bum ble
bees of the Bom bus sibiricus-group givingthe highes t success ive
scores for taxica l ly weighted complementary divers i ty. The
areas arelabelled b y reference to m ajo r geo-poli tica l fea
tures near the cen tre of the par t icular gr id-squares , as fol
lows: 1, Gansu (China) ; 2, Big Horn (USA); 3, Ecuador ; 4, Turkey;
5, Baikal(US SR ); 6, H ind u K ush (Afghanis tan) ; 7, M icho acan
(Mexico) ; 8, La P az (Bol ivia); 9,Sam arkan d (USS R) ; 10 , Ka
shm ir ( India) ; 11 , Nep a l ; 12 , Aru nacha l P radesh ( India)
; 13 ,Nor the rn Ca l i fornia (USA) .The m ap (Wi ll iams , in pre
s s) i s based on a cylindri ca l, equa l -a rea projec t ion , or
tho m orph ica t 45 N /S (where bu m ble bee r ecords a re pa r t i
cula r ly num erous ). Th e gr id i s ca lcula ted f romintervals
of 10 Lo ngi tud e to provid e equa l a reas tha t a pp ea r a s
squa res on th is projec t ion .The po r t i on o f t he g ri d s
how n c ove r s the k now n na t ive d i s t ri bu t i on o f a ll
bum bl e be es .
o n e s q u a r e w i t h a l l 4 3 s i b i r i c u s - g r o u
p s p e c i e s c o n t a i n e d w i t h i n i t) , o rc o m p l e
t e a l l o p a t r y w i t h a ll s p e c i e s r e g a r d e d a
s t a x o n o m i c a l l y e q u a l ( t h is l in es t a r ts a t
2 3 % , o n e s p e c i e s o u t o f 4 3 , a n d r e a c h e s 1 0
0 % a t a r e a 4 3 ). T h es h a p e o f t h e c u m u l a t i v e
d i v e r s i ty c u r v e f o r t h e p r io r i t y s e q u e n c
e ( w h i c h i nt h is a n a ly s i s , a s a l r e a d y n o t e
d , r e a c h e s 1 0 0 % a t a r e a 1 3) g i v e s a m e a s u r
e o ft h e g e o g r a p h i c d i s p e r s i o n o f t a x i c d
i v e r s i t y f o r t h e g r o u p , w o r l d w i d e . W eb e
l i ev e th a t t h is m e a s u r e , o n c e p r o p e r l y q u
a n t i f i e d , w i ll b e o f u s e in e c o l o g ya n d c o n
s e r v a t i o n p l a n n in g . T h e 1 0 0 % p o i n t i n F i
g . 8 c o r r e s p o n d s t o t h e( si te ) e f f ic i e n c y m
e a s u r e o f P r e s s e y a n d N i c h o l l s ( 1 9 8 9 a ) (
E , e f fi c ie n c y , i sd e f in e d b y th e f o r m u l a E =
1 - ( X / T ) , w h e r e X is t h e m i n i m u m n u m b e r o re
x t e n t o f s i t es n e e d e d t o c o n t a i n a ll a t t r i
b u t e s r e q u i r e d [ t a x a ] , a n d T i s t h et o t a l
n u m b e r o r a r e a o f s i t e s s a m p l e d ; P r e s s e y
& N i c h o l l s , 1 9 8 9a ). D i f fe r e n tp a t t e r n s
o f d i s p e r s i o n a n d a c c u m u l a t i o n w i ll c a ll
f o r q u i t e d i f f e re n t a c t io n s i fc o n s e r v a t
i o n p l a n s a r e to b e ef fe c ti ve .
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ystematics and conservation evaluation 2491
9
87
65o
-~ 3 oE~ 2
lO1 2 3 4 5 6 7 8 9 1 11 12
n o o f a r e a sFig. 8. W eighted taxic diversi ty accu mu lat
ion histogram for the 43 bumb le bee species ofthe Bom bussibiricus
group. Th e abscissa represents the num ber o f sample areas ( the
sibiricusgro up occu rs in 120 of the 250 grid-squares show n in
Fig. 7). Th e ord inate scales percentag ediversi ty (based on the
agg regate of the 43 weights for the terminal tax a given in Fig.
6). Th e13 po ints co rres po nd to the sequence o f grid-squares
listed in the legend to Fig. 7, the 13prior i ty areas. T hus the f
irs t poin t is area 1, Ga nsu, con tr ibut ing 22.95 % o f the
total w eighteddiversi ty; area 2 is Big H orn , contr ibut ing an
ad di t ional 5.25 % of com plem entary weighteddiversi ty, and so
on up to area 13, N orth ern Cal i fornia, which co ntr ibutes the
last 2-19% o fweighted diversity, and br ings the accum ulat ion to
100% (the cum ulat ive nu m ber of speciestak en in to acc ou nt at
eac h successive step, up to the to tal o f 43 at the 13th, is 9,
13, 23, 28, 30,32, 35, 38, 39, 40, 41,42, 43). T he (incomplete) ve
rtical straight l ine indicates how the h istogr amwo uld loo k i f
there was at least on e gr id-square con taining al l 43 species;
the s loping st raightl ine is a bou nd indicat ing how the histog
ram wo uld lo ok i f a ll 43 species occurred in
differentgrid-squares an d al l were t reated as taxon om ical ly
equal. See also text for explanat ion.
P L A N N I N G N A T U R E R E S E R V E N E T W O R K S W I T
H I NC R I T I C A L A R E A SI d e n t i f i c a t i o n o f w h o
l e c o u n t r i e s , g e o g r a p h i c r e g io n s , i s la n
d s o r l a r g e g r i ds q u a r e s a s c r i t ic a l f a u n a
s o r f l o r a s is a b a s i c s te p , b u t s u c h u n i t s r
a r e l yr e p r e s e n t p r a c t ic a l ly c o n s e r v a b l
e a r e a s i n t e r m s o f c u r r e n t e c o n o m i cr e s o
u r c e s . F u r t h e r m o r e , w i t h i n c r i t i c a l a r
e a s , a ll s p e c i es o f i n t e r e s t r a r e l yc o e x i
s t a t s in g l e l o c a li t ie s . I t is t h e r e f o r e n e
c e s s a r y t o u n d e r t a k e f u r t h e rr o u n d s o f a
n a l y si s , w i t h i n e a c h p r i o r i t y r e g i o n , to
d e v e l o p e f f ec t i v e a c t i o np l a n s f o r c o n s e
r v a t i o n .F u n d a m e n t a l t o s e c o n d a r y a n a l
y se s is t h e v i ew t h a t a ll a n i m a l a n d p l a n ts p
e ci e s w i t h i n a c r i t i c a l a r e a , r e g a r d l e s
s o f g e n e r a l a b u n d a n c e o r l o c a le n d e m i c i
t y , s h o u l d b e ( i n s o f a r a s i t i s p o s s ib l e )
e q u a l l y a n d a d e q u a t e l y
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25 R. I . Vane -Wrigh t C . J . Hum phr ies P . H . W i l l
iams
p r o t e c te d . T h i s is b ec a u s e r a re o r e n d e m
i c ta x a , t o g e t h e r w i th c o m m o n o rw i d e s p r e
a d s p ec ie s, d o n o t e x is t in i s o l a t i o n - - t h e
y e x i st w i t h i n m o r e o r le ssw e l l -d e f in e d e c o
s y s t e m s o r c o m m u n i t i e s , o n w h i c h t h e y a r
e d e p e n d e n t , a n dt o w h i c h t h e y m a k e t h e i r
o w n u n i q u e c o n t r i b u t i o n s . T h u s , o n c e i d
en t if ie d ,t h e w h o l e b i o t a o f a p r io r i t y r e se
r v e a r e a n e e d s a d e q u a t e p r o t e c t i o n a s af
u n c t i o n a l e c o l o g i c a l s y s t e m , o r s e t o f s
u c h s y s t e m s .
A s a l r e a d y n o t e d , e c o l o g i s t s a n d c o n s
e r v a t i o n i s t s w o r k i n g f o r n a t i o n a la n d s
t a t e w i l d li fe s er v ic e s i n A u s t r a l i a a r e d e
v e l o p i n g s te p - w i s e a n a l y s e si n t e n d e d t o
se r v e t h i s p a r t i c u l a r f u n c t i o n ( M a r g u l
e s , 1 9 86 ; M a r g u l e sN i c h o l l s , 1 98 7; M a r g u l
e s e t a l . 1 98 8; M arg u l e s S t e i n , 1 98 9; M cK en z i
e e tal . 1 98 9; P r e s s e y N i c h o l l s , 1 98 9a ,b ). T h
e i r a n a l y s e s c a n t a k e i n t o a c c o u n tt h e d e
s i r e to p r o t e c t a ll s p e ci e s a n d r e p r e s e n t
a t i v e e c o s y s t e m s w i t h i n ar e g io n . B y p l a n
n i n g m u l t i p l e r e p r e s e n t a t i o n s w h e r e v e
r p o s s ib l e , t h e y a l s om a k e a l l o w a n c e f o r p
o p u l a t i o n e f f ec ts , s u c h a s p o s s i b i l it ie s
o f l o c a le x ti nc t i o n. T h e i r a p p r o a c h a p p e a
r s c a p a b le o f a c c o m m o d a t i n g a nyt h e o r e t i
c a l o r p r a c t i c a l d e v e l o p m e n t s a f f e c ti n
g t h e siz e, s h a p e a n d d e s i g n o fn a t u r e r e s er
v e s. F o r o p t i m i s i n g e ff ic ie n c y, h o w e v e r ,
t h e y w i l l a l s o n e e d t oa d d r e s s t h e a n a l y t
i c a l p r o b l e m s a s s o c i a t e d w i t h t h e p o t e n
t i a l a d v a n t a g e sa n d d i s a d v a n t a g e s o f s e
t- w i se v e r s u s st e p -w i s e m e t h o d s .
W e b e li ev e , n o n e t h e l e s s , t h a t t h e s e a n
a l y ti c a l m e t h o d s d e s i g n e d t o i d e n t i f ye f
fi c ie n t n a t u r e r e s e r v e n e t w o r k s w i t h i n s
p e c i fi e d o r l o c a l r e g i o n s a r e w e ll -s u i te d
t o t h e n e e d s o f s e c o n d a r y a n a l y se s . H o w e
v e r , t h e p r i m a r y a n a l y s e ss h o u l d i n v o l v
e t h e i d e n t i f i c a t i o n o f p r i o r i t y a r e a s o
n a g l o b a l s c a le , o rm e a s u r e m e n t o f t h e g lo
b a l c o n t r i b u t io n s m a d e b y e x is ti n g n a t i o
n a l p a r k s ,t h r o u g h u s e o f p r i o r i t y a r e a a
n a l y s e s, a s a lr e a d y o u t l in e d . O n l y b y a na p
p r o a c h o f th is t y p e c a n r e q u i r e m e n t s f o r t
h e p r o p o s e d I n t e r n a t i o n a lC o n v e n t i o n (
C h a l k e r , 1 99 0) b e m e t w i t h a n e f fi c ie n t s y s
t e m a t i c p e r sp e c t i v e .
G L O B A L P R O S P E C T SM a n y b i o l o g i s t s , e s p
e c i a l l y n o n - s y s t e m a t i s t s , m a y t a k e e x c
e p t i o n t oi n t r o d u c t i o n o f t a x o n o m i c w e i
g h t in g in t o c o n s e r v a t i o n p r o c e d u re s . S u
c h ar e a c t i o n i s l i k e l y b e c a u s e t o d o s o c o
u l d b e s e e n t o a d d a n e w l e v e l o fu n c e r t a i n
t y . I t h a s o f t e n b e e n a r g u e d t h a t h i g h e r t
a x a a r e a r ti fi ci al , w h e r e a ss p e c ie s a r e n a t
u r a l o r ' s t a b l e ' u n i ts . O n a w o r l d s c a le t h
is is c e r t a i n l y n o t t h ec a s e : h i g h e r t a x a a
r e n o m o r e d i f fi c u lt t o r e c o g n i s e o r d e f i n
e t h a n s p e c i e sl ev e l t a x a . I n d e e d , i n m a n y
c a s e s (e .g . M a m m a l i a v e r s u s m a n y s p e c i es
o f M u so r R u b u s v e r s u s m a n y o f it s 's p ec ie s')
, h i g h e r t a x a a r e m u c h m o r e s t a b le a n de a s i
ly d e l i m i t e d .
I f c o n s e r v a t i o n e v a l u a t i o n i s t o b e b a
s e d o n c o n c e p t s o f d iv e r si ty , t h e n w eb e li ev
e t h a t i m p e r f e c t i o n s o f o u r k n o w l e d g e o f
t h e d i v e rg e n c e o f t a x a
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ystematics and conservation evaluation 251t h r o u g h ti m e c
l a ss i fi c a ti o n ) a n d d i s t r i b u t i o n in s p a c e
b i o g e o g r a p h y ) h a v et o b e a c c e p t e d , a n d c
o n t i n u a l a d j u s t m e n t s m a d e a s o u r u n d e r s
t a n d i n g o fb o t h c h a n g e s a n d i m p r o v es . T h e
c u r r e n t e s ti m a t e o f th e w o r l d n u m b e r o fb i
o l o gi c a l s p ec ie s is u n c e r t a in b y a n o r d e r o
f m a g n i t u d e o r m o r e W i ls o n ,1 9 8 9 ; May , i n p r
e ss ) . Of th o se 1 .5 - 1 .8 m i l l i o n p l an t an d an im a
l sp ec i e sn a m e d t o d a t e , o n l y a s m a l l fr a c t i
o n c a n b e c o n s i d e r e d w e l l - k n o w n in t e r m so
f sp ec i e s - l im i t s , h ig h e r c l a s s i f i ca t io n ,
d i s t r i b u t io n , an d b as i c eco lo g ica lr e q u i r e
m e n t s . T h u s t h e g l o b a l a n a l y s e s c a n b e b a
s e d , f o r t h e i m m e d i a t ef u t u r e a t l e a s t , o
n l y o n t h o s e p l a n t a n d a n i m a l g r o u p s w h i c
h a r e a l r e a d yw e l l - k n o w n , o r f o r w h i c h s u
i t a b l e s t u d y m a t e r i a l a l r e a d y e x i s t s
.
O u r n e x t c o n c e r n , a p a r t f r o m r e fi n in g t
a x ic i n f o r m a t i o n m e a s u r e s a n dt h e i r a p p l
i c a t i o n t o p r i o r i t y a r e a p r o c e d u r e s , i s
t o a s s e m b l e a f e d e r a t i o n o fs y s te m a t i s ts
w i l li n g t o p o o l d a t a o n p a r t i c u l a r a n d a p
p r o p r i a t e g r o u p s .I n i t i a l ly a t le a s t w e i
n t e n d t o c o n f i n e o u r a t t e n t i o n t o t e r r e s
t r ia l o r g a n i s m s .W i t h i n o u r o w n i n s t i t u t
i o n w e a r e c u r r e n t l y a b l e t o c o m m i t w o r k o
nd a n a i n e , i t h o m i i n e a n d p a p i l i o n i d b u t
t e r f l i e s , s p h i n g i d m o t h s , a n dC o n i f e ra l
e s. T h r o u g h c o l l a b o r a t i o n w i t h t h e R o y a
l B o t a n i c G a r d e n s , K e w ,w e a n t i c i p a t e m a
j o r a n a l y s e s o f t h e t r e e f a m i l i e s C h r y s o
b a l a n a c e a e a n dL e c y t h i d a c e a e . A s t h e l e
a d i n g s p e c i a l i s t o n t h e s e g r o u p s i s a l r e
a d y d e e p l yi n v o l v e d i n c o n s i d e r a t i o n s o
f p r i o r i t y a r e a s f o r c o n s e r v a t i o n P r a n c
e , 1 99 0),t h is p a r t i c u l a r i n it ia t iv e w i ll p r
o v i d e u n i q u e o p p o r t u n i t i e s t o d e v e l o p a
n dt e s t o u r m e t h o d s .
W i t h i n t h e n e a r f u t u r e w e h o p e t o s e c u r
e p o t e n t i a l c o l l a b o r a t i o n w i t hw o r k e r s
o n m a n y m o r e g r o u p s . S c i e n ti fi c a ll y , a f u
r t h e r t a s k w i l l t h e n e m e r g ea s t o h o w b e s t
t o e v a l u a t e i n f o r m a t i o n g e n e r a t e d f r o m
s u c h d i s p a r a t eg r o u p s f o r th e m a j o r t a s k
in h a n d - - r e c o g n i t i o n o f p r i o ri t y ar e a s, a
n d t h ea s s e s sm e n t o f t h e c o n t r i b u t i o n t h a
t e x i s ti n g n a t i o n a l p a r k s a n d o t h e rr e se r
v es m ak e . Po l i t ic a l ly , i f we a r e t o b e e f fec tiv
e , we w i l l a l so n ee d to a l lyo u r f e d e r a t i o n a n
d s y s t e m t o o n e o r m o r e o f t h e m a j o r i n t e r n
a t i o n a la g e n c ie s c o n c e r n e d w i t h t h e c o n s
e r v a t i o n o f w i ld li fe .
A C K N O W L E D G E M E N T SW e w o u l d l ik e t o t h a n
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l C h a l m e r s ,I a n G a u l d , L a u r e n c e M o u n d , J
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