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[]r. Gerardo CeltallosG. t] d s

o What is Conservation
Biolog;y?
A new syntbetic_dis-cipline
addresses
the dynamicsand problems
of perturbeil species,cotnmunities,-and
ecosysteits
MichaelE. Soul6
I

o ns er v at ion bi o l o g y , a n e rv
s t agein t he app l i c a ti o no f s c i Although crisisoriented
\-rf
etrce to conservation probrl e
c rms,
rr)t d
aududr
t c es
s s ssses
conservationbiology
t the
r t r bio
u t o l loogg
y yo
oIf ssppeeccl gi e
s ,s ,
c o mmu nit ies ,and ec os y s te mth
s a t a re
is concernedwith
pe rtu rb ed,eit her dir ectl y o r i n d i re c tly, by hunran activities or orher
the long-term viability
agents.lts goal is to provide princiof whole systems
ples-and tools for preservingbiological diversity. ln this articl. i d.r..it.
c o n se rv at ionbiology , d e fi n e i ts .fu n da me n tal pr opos it ion s , a n d n o te e o n r m e n d a t i o n sa b o u t d e s i g n a n d
fe rv o f i t s c ont r ibut ion s . I a l s o p o i n r ma n a g e me ntbeforehe or she i s comthat ethical norms are a genuine pletely comfortabfe with the theoretic#t
of conservarion biology, ir th.y c a l a n d e rn p i ri calbasesof the anal ysi s
W.t
are in all mission- or crisls-oriented (Ma y 1 9 8 4 , S oul and W i l co.x 1980,
disciplines.
c h a p . l ). T o l erati nguncertai ntyi s often necessary.
Conservation biologists are being
Crisisdisciplines
asked for adviceby governmentagenConservation biology differs from ciesand private organizationson s-uch
most other biological sciencesin one p ro b l e m sa s the ecol ogi caland l real tl r
important way: it is often a crisis c o n s e q u e n c es
of chemi cal pol l uti on,
di sci p l i n e.lt s r : lat ion t o b i o l o g y , p a r- th e i n tro d u c ti on of exoti c speci esanj
t icu l a rl y ec ology ,is ana l o g o u J i o i h a t a rti fi c i a l l y .p r oduced strai ni of exi stot -surgeryto physiology and war to i n g o rg a n i s ms,the si tesand si zes of
po h ti ca l s c ienc e.I n c r is i s d i s c i p l i n e s , n a ti o n a l p a rk s, the defi ni ti onof mi ni on e mu s t ac t bef or e k n o w i n g i l l th . mu m c o n d i t i ons for vi abl e popul af acts;cri s isdis c iplinesa re th u s a m i x - tions of particular target speCieC,
the
t ure o f s c ienc e-and a rr. a n d th e i r tre q u e n c i e sand ki nds of management
p u r s u i t r e q u i r e s i n t u i t i o n a s w e l l a s p ra c ti c e si n e xi sti ngrefuges
and manin fo rma tion. A c ons er v a ti .obni o l o g i s t r-g :d w i l d l a nds, and the ecol ogi cal
may have to make decisions or rec- ffects of development.
Feir poliiicat
reasons,such decisionsmirst often be
Michael E. Soul is an adiunct Drofessor
m a d ei n h a s t e .
in rhe Wildlantl Management Ceiter, Uni-sclrool
F o .re x a mpl e, the rapi di ty and i rreversiry' of Michigan
of Natural
v e rs i b i l i ty o f l oggi ng i nd human reResources. He is coauthor r*.ith Orto H.
s e ttl e m e n t i n W estern N ew Gui nea
Frdnkel of Conseruatlon and Euittntion,
prtrlish:d by Carnbridge Universiry press. (l ri a n J a y a ) prompted the l ndonesi an
Addrc-ss correspondence to 474j Black governntent to establisha system of
national parks. Two of the largest
.}lgqt ti;:- Road, San Diego, CA 92130.
Amc.ican Instiruie of Biological areas recomrnendedhad never been
81985
u_-___
visited by biologists,but it appeared

,.lJer1985
Lr<,...:'..

... -,.--,.:l

l i kel y that these areas harbor ed endemi c bi otas.l R econnai ssance

lat er
confirmed this. The park boundaries
l r - e r ee s t a b l i s h e idn t g g l . a n d s u b s e q.uel l t.devel opment
has al ready pr ecluded all but minor adjustments.
S i mi l ar cri sesare now faci ng m anagers o( endangeredhabitarcrnd speciis
i n the U ni ted S tates-for ex am ple,
grizzly bears in the Yellowstone region, black-footed ferrers in Wyomi1g, old-growth Douglas-fir forists
in the Pacific Northwesl, red-cockaded woodpeckersin the Southeast,and
condors i n C al i forni a.

Other characteristics
of
csnservation biology
As illustrated in Figure l, conservati on bi ol ogy sharescertai ncharact er istics with other crisis-orienteddisciplines. A comparisonwith cancer
bi ol ogy i l l ustratessomeof thesechar acteri sti cs,i ncl udi ngconservationbiol ogy' s syntheri c, ecl ecti c,mult idisciplinary structure. Furthermore,
both fi el ds take many of thei r questions, techniques,and rnethods from
a bloa{ range of fields,not all biofogi cal . Thi s i l l ustrati oni s al so i ntendJd
to show the arti fi ci al i tyof the dichot 9my .between pure' and applied
di sci pl i nes.
Finally, this figure illustrates the
dependenceof the biological sciences
on social sciencedisciplines.Today,
for exampl e, any recommend at ions
about the location and sizeof nationrJaredM. Diarnond,1985,personalcommunicatian. Universiryo{ California,Los Angcles.

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CONSERI4ATION

BIOLOGY

BiogeograPhY
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CANCER
BIOLOGY

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Figurc 1. Cancer biology and conservationbiology arc both synthctic, multidisciplinaryscicnces.The dashedline indicates the
artificial nature of the bordcrsbctweendisciplinesand bctween"basic" and "applied" research.Scctcxt.
a l p a rk s s hould c ons i d e rth e i m p a c t o f
4qthe park on indigenous peoples and
cultures, on the local economy,
Etheir
and on opportunity costssuch as forfeited logging profits.
There is much overlap between
conservation biology and the natural
resourcefields, especiallyfisheriesbiology, forestry, and wildlife management. Nevertheless,two characteristics of these fields often distinguish
them from conservationbiology. The
6rst is the dominance in the resource
fields of utilitarian, economic objecti ve s.E v en t hough in d i v i d u a l rv i l d l i fe
b i o l o gis t shonor A ldo L e o p o l d ' sl a n d
e th i c and t he int r ins icv a l u e o f n a tu re ,
most of the financial resources for
ma n a gem ent m us t g o to e n h a n c i n g
co mmer c ial and r ecre a ti o n a lv a l u e s
for humans. The emphasisis on orlr
naturaf resources.
The second distinguishing characteristic is the nature of these resources.For the most part, they are a
sma l l num ber of par ti c u l a rl yv a l u a b l e
target species(e.g.,trees, fishes,deer,
and waterfowl)-a tiny fraction of
the total biota. This distinction is
beginning to.disappear, howe.ver,as
some natural resource agencies be"ecotogical" and because
come rnore

728

c o n s e rv a ti onbi ol ogi stsfrequentl yfocus on individual endangered,critical,

or keystone species.
Conservation biology tends to be
holistic, in two sensesof the word.
First, many conservation biologists,
including many wildlife specialists,
a s s u m eth a t ecol ogi caland evol uti onary processesmust be studied at their
own macroscopic levels and that reductionism alone cannot lead to exp l a n a ti o n sof communi ty and ecosyst e m p r o c e s s e ss u c h a s b o d y - s i z e
differencesamong species in guilds
(C o d y a n d D i amond 1975), pol l i nator-plant coevolution (Gilbert and
R a v e n 1 9 75),,succes5i on,speci ati on,
a n d s p e c ies-arearel ati onshi ps.E ven
ecological reductionists, however,
agree tha.t ,h: proper obiective of
conseft'ation is the protection and
c o n ti n u i ty of enti re communi ti esand
e c o s y s te m s.
The hol i sti c assumpti on
of conservationbiology should not be
confusedwith romantic notions that
o n e c a n g r asp the functi onal i ntri cacies of complex systemswithout conducting scientific and technological
studies of individual components
(Levinsand Lewontin 1985, chap. 5).
Holism is not mysticism.
The second implication of the term

ltolisticis the assumptionthat multiwill ultimateapproaches

disciplinary
lv be the most fruidul. Conservation
biology is certiinly holistic in this
analysis
Modernbiogeographic
sense.
is now beinginttgratedinto the conservationmovement(Diamond 1975,
Simberloffand Abele 1976,Terborgh
1974,t0filcox1980). Populationgenetics,too, is now being aPPliedto
the technologyof wildlife management(Frankefi 974,Frankeland Soul6
et al. 1983'
1981, Schonewald-Cox
Soul6 and Wilcox 1980). Multidisciplinary resgarch,!ryg.t"ile. Bovernand-wildlifebi;lctgsts,is
mentagenoes
also.uld.nt in recenteffortsto illumi"
nrt. tt. questionJ uit*. population
'n.i'
et al. 19841..
size(Salwasser
,
ract{l'$:
ch.a
ishing
Anotherdistingu
biology.ls tts ttme
tic of conservation
practltlonersatits
Generally,
scate.
maxi'
aesthetics.'
to
ir.i,'r.*';;Glil
tnq
,nJProfitab.ilitY'
mum vields,
o'
vlablllty,.
long-range
the
io
more
lncludlng
and species'
wholesystems
their evolutionarypotentlal' Lgtlgtermviabilityof narural.commumtls
oj ol'
usuallvimpliesthe persiste.nc.e
help.trom
no"
versity,with little or
But for ,h* fottteeable fui;;t;;.
a'passiverole for manag*s
tu.tt
tu*,
BioscienceVoL 35

is u.nrealistic,
and virturirllyall conser- b i ota. For exanrpl e,
rhe responscsof ranr as brecdi ng
y at lon pr ogr a rn sw i l l n c e d ro b e b u rt> r fe eding sir cs f or
p rey to a predator' sappearr,.,..< l r of a n i m a l s
rnay initiate ,"qi.,r..,
r r es s edar t if ic i a l l y .F o r:e x a m p l e e
, vcp
l h l t o R h a g o u s i n s c c i r o p o r e n t i a l c a u s a l l yl i n k e,de v e n r st h a t ' u t r i m a or cf l y
r
r he lar ges naru
rere s e rl i s ,n i n a ri o n - h3 o s t p l a n r s a r c
c o n t i n u a l l y. . t u n e d " l e a d r o f u r t h e r e x r i n c t i o n J ' i i r " * t . t
al parks are affe.ctedby anrhropogen_ b y n a r u r a l
sclcction
a n d S o u l e 1 9 8l , G i l b c r r l 9 g { J , T e r _
ic f ac r or s in th e s u i ro u n d rn s a re a
Thi s posrul aremercl y asscrrsrhat b o r g h a r r dW i n r e r 1 9 8 0 ) .
.
{ J a n z e n -1 9 8 3 , K u s h l a n l g 7 g i , a n d rh e srrucrure,funcri on,
and srabi l i ty
lntroductions of generalists ntitl r_
such rcfugesare usually roo sriiall ro o f c o e v o l v c d ,
n a t u r a l c o m m u n i r i e s duce di t,ersi ty.T' h; i rr r r o. lucr ion
of
c onr ain v iabl e p o p u l a i i o n s o f l a rg e d i ff er
si gni fi canrl yfronr those of un- exori c p.l antand ani mal
speciesnr ay
c ar n- iv or es( F ra n k e la n d So u l i , l 9 g j ,
n a r ural or syntheri ccommuni ti es. l t reduccdi vcrsi ry,espec iaill, 'r f
t hey ar e
S haf f er and S a m s o n 1 9 8 5 ). In a d d i _ d o e snot necessari l y
rcl y on derermi n- targe or general rsrspe cies( Diam ond
r ion, poac hi n g , h a b i ra t fra g men ra _ i s ri c facrors l i ke
-a n i m a l s
dcnsi tv-depe
ndenr 1 9 8 4 , E l t o n 1 9 5 8 ) . A p p a r e n r l y ,r h e
t ion, and t he i n fl u x o f fe ra l
p o pul ari on dynami csor i he mol di ng l arger the l and mass, ''r he
leis r he
and exotic planrs require extraordi- by comperirion
of morphological rel
i Tpl ., of exori cs (e . g. , Sim ber lof f
nar.y pracricessuch as culling, eradj_ l a ri o nshi psi n
communi i i esoi er both 1 e 8 0 ) .
carion, wildlife immunizarionlhabitar e c o l ogi cal
The evol uri onaryposr ulat eand it s
.and evol uti onary ti me. In
prorection, and aruficial rransfers. a d d i ri on, rhi
s posrul arei s neutral on corollariesforrnaliz. the evidence
rhar
Until benign neglecr.is again a possi_ th e i ssueof hol i sri c
versusreducti on- natural conrmfunities
comprise species
Dr t lt y ,c ons er v a n o nb i o l o g y c a n c o m - i s ti c anal ysi sof
cornmuni rysrrucrure. w h o s e . . g e n e t i m
c a k e u p s 'h a v e ' b e e n
ple menr narural r.rour.r. fields in ([n pracri ce,
a reducri oni si i cmethod- murually affecredby thiir coexisrence
pr ov iding s omeo f th e rh e o re ri c aal n d o l o g y , i n c l u d i n g
a u r e c o l o g i c a l r e - ( F u t u y m aa n d S l a r k i n 1 9 8 3 , G i l b e r r
ernpirical foundarionsfor coping with search,may be
the besr\4,ay-toestab- a n d R a v e n l 9 7 S ) . A n a l r e r n a r i v er h e such managementconilndrunrs]
l i s h t h e h o l i s t i c s r r u c r u r c o f ory, the nul l hypothesist har
com m ucommuniries.)
ni ti esare randoml y assem bled,is usuT h ere are many " corol l ari es' , of a l l y r e s t r i c t e d t o " h o r i z o n t a l "
Postulatesof conservation
th i s .p osrul are.S tri ctl yspeaki ng,mosr subcommuni ti essuch
as guilds,
biology
- (spe. . g. ,
of rhem a19eTpiricaliy 6ased
feneral- ci fi c taxa, or .trophi c levels
i
z
a
tions.
The fol l ow rngal l assi me rhe Jamesand B oeckl en 1984) . I n
Conservation biology,
'is like many of
sener existenceof communlty processesas al , thi s l arter thesi s
l a cks em f ir ical
lt11 parenr sciences, very young.
Therefore, ir is nor surprising it ar iis w e l l as a coevol uti onary' component s u p p o r t , e x c e p r r h a t c o n r p e t i r i v e
-trophic
assumptionsabour the srruciure and in community structure.
srrucru.ringwithin guilds or
Spec.ies
are
t u n c t i o n o f n a t u r a l , s y s r e m s ,a n d
interdependent. Not levels is ofren absenr or difficuli to
about the role of humans in narure, o n l y have speci es i n cornmuni ti es demonstrare(S rronget a l. 1984) ,and
have not beensysremar{zed.
Whar are e v o l v e d u n i q u e w a y s o f a v o i d i n g that harsh envi ronmenrsor t he vagapredators,locating food, and caprur_ ries of dispersalmay
these
pro$se
I
rwo
sers: a
often be nlore
.postulares?
tunctronal, or mechani$ic, set and an i n g a nd handl i ng pr.y, but nrurual i s- imporrant than biological interacti c re l ari onshi psai e frequent fl anzen tions in determining local
erhical, or normative, $r.community
7 9 7 5 , S ei ferrand S ei feri tl ZfJ. tt i s composirion (e.g., Underwood
and
The_ functional postulafes. These are is nol
19 say that every species is Denfey 1984).
working proposirions
partly
on e s s e n ti alfor communi ryfuncti on, bur
$sed
The second functi on al post ut at e
evidence,parrly on rhefry, .nd pirrly
th a r th erei s al w ays uncerrai nty .6our concernsthe scale
of ecological proon rnfuttron. In essence,arhey
are a ser th e i n reracti onsof speci csanj
abour cesses:MIry, if not all, ecological
of fundamental axioms$derived fr"m
th e b iol ogi calconseq,,.n... of an ex- processeshaue
thresholds belotiand
ecology, biogeogr"ph-f,i and popula_ rrn c rr on.
P artl y for rhi s reason, A l do aboueuhicb they beconrc discotttiuution genetics,aboui rhd mainienance L e o p o l d ( 1 9 j 3 )
a d m o n i s h e dc o n s c r _ ous, cbaotic, or suspenderl.This posof both the form and function of v a n o nl sts
ro save al l of the parts tul atesraresrhar many ecologicr r l
pr obiologicalsysteirs. They sug- (s p e ci es)
of a communi ry.
i1,:lllrhe
cesses and parrrns (including
gest
rules for action. A necessarv
. illofy -species are .ltigy ly .special- succe.ssi on,
nutri ent cycl ing, and dengoal of conservation bioloey is rhl
i z e d ..Perhapsrhc rrr.r;onty
of ani nral si ty-depcndcnr
phenon-r cn";ar e int er elaborarion and refinegrent-of such s p e c t es,i ncl udi ng
phyrophasous In_ rupted or fail altogether r,r,herethe
principles.
i
parasircs,and parisirdids, de_ system
i s too srnal l . S m allnessand
l.,.,tl
. The firsr, the evoludonary posru- p : l d o n a p a r r i c u l a r h o s t ( p r i c e randomnessare inseparable.
Iate srares: Mauy
,of th:e spegiis t:hat, 1 9 8 0 ) .,Thi s means rhat .n.-* ri rr" i l ,
Nonecological processesmay also
codstitute natural communities ale of endangered host
,
specics can be domtnare
at the other end of t he
the products o{ coeuoSttiortdry pro.
very long,. ra-king rvith rhcm tlozens spati al and
tenrporal sc ale, in ver y
cesses.ln mosr commuhiries, speciesl (R a v e n 1976.r or hundreds (E rw i n
l arge or very ol d sysre m s. I n ver y
are a significanrpart of one anolherts , 1 9 8 3 ) o f s m a l l c o n s u n r e r s r r e c i e s
largg systemsrsuch as conrinents,cli'
environment.Therefori their geneti- when rhey go.
matic and physiographic phenomena
cally basedphysiological
and 6ehav- . Extinrciio"is of heystone speciescatl otten dererminethe maior parternsof
ioral re.perroires
have beennarurally hau.e lgng-r1nge
.consequences. The th.e la.ndscape,including speciesdisselected to accommodatethe exls- extinction of maior
predarors, large trtbutton. ln very old systems, ecotenc and reactionsof a particular h e rb rvores,or pl ants that are i mpor- logical processes
give way'ro geologi-

December 1985

t1nt*o1|.,I J - ,*oL"nl rzs

'cal

and historical ones or to

i n f r e q u e n tc a t a s t r o p h i ce v e n t s ,s u c h
a s i n u n d a t i o n ,v o l c a n i s m a, n d g l a c i a t ion. ln ot her v t' o rd s ,e c o l o g i c rlp rc rbelongto a n i n te rl n e d i a tes c a l e
c es s es
o f p h y s i c a sl i z ee n d t i m e ( N l a c A r t l r u r
1972) , and t lre s e p ro c c s s e sb e g i n to
f ail or ar e o1' e r!r' h e l rn ende a r th e e x t r enr it iesof t he s era n g e s .
T wo m ajor a s s u mp ti o n so, r g e n e raliz at ions , un d e rl i e th i s p o s tu l a te .
First, the tentporal continuity of babitats and successional stagesdepends
on s iz e.T he r a n d o m d i s a p p e a ra n coef
r es our c esor h a b i ta ts w i l l o c c u r fre quent ly in s nra l l s i te s b u t ra re l y , i f
ever, in large ones. Jlre reasonsinc lude t he inh e re n t ra n d o mn e s s o f
s uc h pr oc es s e sa s p a tc h d y n a mi c s ,
larval settlement, or catastrophic
ev ent s , as r v e l l a s th e d y n a m i c s o f
c ont agious ph e n o me n a s u c h a s d i s eas e,r v inds t or md e s tru c ti o n ,a n d fi re .
T he lar geran a re a ,th e l e s sl i k e l y th a t
all pat c hesof a p a rti c u l a rh a b i ta tu ' i l l
disappear simultaneously. Species
will disappear if their habitats
disappear.
Second, outbursts reduce diuersity.
I f populat ion d e n s i ti e so f e c o l o g i c a l l y
dom inant s pec i e sri s e a b o v e s u s ta i n able levels,they can destroylocal prey
populations and other speciessharing
a resource with such species.Outbur s t sar e m os t p ro b a b l ei n s rn a l ls i te s
t hat lac k a f u l l a rra y o f p o p u l a ti o n
buffering mechanisms,including habit at s ink s f or d i s p e rs i n gi n d i v i d u a l s ,
sufficient predators, and alternative
f eeding gr oun d s d u ri rrg i n c l e m e n t
weat her .T he u n u s u a l l y h i g h p o p u l a t ion dens it iest h a t o fte n o c c u r i n n a t ur e r es er v es
c a n a l s o i n c re a s eth e ra te
of dis eas e t r a n s mi s s i o n , fre q u e n tl y
leading t o epi d e mi c sth a t m a y a ffe c t
ev er y indiv idu a l .
T ak en t ogeth e r, th e c o ro l l a ri e so f
t his pos t ulat el e a d to th e c o n c l u s i o n
t hat s ur v iv al r a te s o f s p e c i e si n re serv'es are proportional to reserve
size. Even though there is now a
consensusthat s'everalsmall sites.can
c ont ain as m an y s p e c i e sa s o n e l a rg e
site (when barriers to dispersal are
absent), the speciesextinction rate is
gener allyhigh e r i n s m a l l s i te s(S o u l i
and S im ber lof f,i n p re s s ).
T he t hir d f un c ti o n a l p o s tu l a tec o n cerns the scaleof population phenomena: Genetic and dentographic processes haue thresholds below which
nottadaptiue, random forces begin to
i

730

i * - i

preuail o{'er adaptiue, detenninistic w oul d precl udegeneti cdif ir r ent iat r on
forces witbin prspulations. l-he sto- a m o n gt h e c o l o n i e s( S o u l 61 9 8 C ) .
c h a s t i cf a c t o r si n p o p u l a t i o n e x t i n c t i o n h a v e h e e n d i s c u s s e de . r t e n s i v e l v The normati ve postulat es.The nor -l-erhorgh
( S h a f f e r1 9 8 1 ,S o u l e 1 9 8 - 1 ,
mative postulatesare value state1 9 7 4 ) i n t h e c o n t e x to f t h e m i r r i m u m m e n t s t h a t n r a k e u p t h e b a s i s o f a n
c o n t l i t i o n s f o r p o p u l a t i o n v i a h i l i t v . ethi c o[ appropri ateatt it udest or var d
T h e m a i n i n r p l i c a t i o no f t h i s p o s t u - other forms of l i fe-an ecosophy
l a t e f r l r c o n s e r v a t i o ni s t h a t t h e p r o b - (N aess 1973). Thev pr ovide st ana b i l i t y o f s u r v i v a lo f a l o c a l p o p u l a - dards by w hi ch our a ct ions can be
ti o n i s a posi ti ve functi on of i ts si ze. measured.They are sh ar ed,I bclieve,
O n e of the corol l ari esof thi s postu- by most conservati on ist sand m any
l a te i s that bel ow a certai npopul ati on bi ol ogi sts,al though i deologicatpur it y
s i z e (betw een10 and 30), the proba- i s not my reas(,nfor proposing t hem .
Diuersity o{ organisms is gaod.
b i l i ty of exti ncti on from random dem o g raphi c events i ncreases steepl y S uch a statementcannot be t est edor
( S h a f f e r1 9 8 1 ) .
proven. The mechanism s by which
The next three corollaries are ge- such val ue i udgments ar ise in conxre.unknown. The concepn e tic. Fi rst, popul ati onsof outbreed- sciousness
i n g organi sms w i l l suffer a chrcni c tual mi nd may accept or r eject t he
loss of fitnessfrom inbreedingdepres- idea as somehow valid or approprision at effective population sizesof ate. If accepted,the idea becomespart
l e s st h a n 5 0 t o 1 0 0 ( F r a n k l i n1 9 8 0 , o f a n i n d i v i d u a l ' sp h i l o s o p h y .
W e coul d specul atea bout t he subg e n e t i cd r i f t i n
S o u l 1 9 8 0 ) .S e c o n d o
"diver si(l
ess
few
a
consci
ousroots of the n or m ,
than
popul
ati
ons
s mal l
h u n dred i ndi vi dual s)w i l l causea pro- ty i s good." l n general,hum ans enioy
g re ssi vel oss of geneti c vari ati on; i n variety. We can never know with certurn, such geneticerosion will reduce tainty whether this is based on avoidimmediate fitnessbecausemultilocus i ne tedi um and boredom or som eh e te rozygosi tyi s general l y advanta- thi ng el se,but i t may be as close t o a
g e o us i n outbreedi ngspeci es(B eard- universalnorm as we can come. This
m o r e 1983, S oul 6 1980, and refer- is probably one of the reasonsfor the
encescited below). (The geneticbases great populariry of zoos and national
o f t hese tw o corol l ari es may be tl re parks, which in recentyearshave had,
same: homozygosity for deleterious, respectively,over 100 million and
re c essi veal l el es.)Fi nal l y, natural se- 200 mi l l i on vi si tors annually in t lr e
lection will be less effective in small United States. Perhaps there is a gep o p ul ati ons becauseof B eneti cdri ft neti c basi s i n humans f or t he appeal
a n d the l oss of potenti al l y adapti ve of biotic diversiry (Orians 1980, Wilg e n e t i cv a r i a t i o n( F r a n k l i n 1 9 8 0 ) .
son 1984). A fter al l , hum ans have
T he fourth functi onal postul atei s been hunter-Batherers,depending on
that nature reserues are inherently a w i de array of hab it at s and r edisequilibrial fo, large, rare orga- sources,for vi rtual l y all of t he past
n i s m s.There are tw o reasonsfor thi s. severalmi l l i on years.
A corol l ary o[ thi s p ost ulat eis t hat
F i rst, exti ncti ons are i nevi tabl e i n
h a b i tat i sl andsthe si ze of nature re- the unti mel y exti nction of populas e r v e s ( l v l a c A r t h u r a n d \ V i l s o n ti ons and speci esi s bad . Conser vat ion
1 9 6 7); speci esdi versi tymust bc arti fi - bi ol ogy does not abhor ext inct ionp. er
c i a l l y mai ntai ned for many tara be- se.N i i ural exti ncti onis t hought t o be
c a u s e n a t u r a l c o l o n i z a t i o n { r e e s t a b - either value free or good becauseit is
l i s h nrei rt) from outsi de sources i s part of tl re processof r eplacingless
h i g hl y unl i kel y. S econd, speci ati on, wetl-adapted gene pools with better
th e onl y other nonarti fi ci al nreansof adaptedones. U l ti mate ly, nat ur al exreplacing species,will not operatefor tinciion, unlessit is cetastrophic,does
ra re or l arge organi smsi n nature re- not reduce biological diversity, for it
s e rvesb.cause ,i r.ru.t are nearl yal ' i s offsetby speci ati on.Nat ur ai ext incrvays too small to keep.L.tgq or rare tions, however, are rare events on a
o rs a ni sms i sol ated w fthi n them for human ti me scal e.Of th e hundr edsof
loie periods, and populations isolat- vertebrate extinctions that have oc;i fi iik.t.", t.t.i".i will haveto be curred during the last few cenruries,
ut artificialseneflorvif few, if any, have been natural (Dia;;l;;l;;J
i-[;t ;t. to persist.Suc[ geneflow arond 1984, Frankelan d Soul6 1981) ,

Biascience
Vol.35No. 1I

1
{

t'

l ,

* nr r . r , t he r a te o f a n th ro p o g e n i c p h i c p o p u l a r i o nc o n t a i n i n gu n i q u ea l - camel l i as, bougai nvi l l e as, daf f odils,

e x r inc t ions app e a rs to b e g ro rv i n g l e l e s o r g e n e r i c c o n r b i r r a r i o n sh a s eucal yptus,and begonrasar e ever ye x p o n en t i a l l y .
B r e a r e rv a l u e , f c l r c x a r n p l e ,t h a n a u' heresi mi l ar.
I t m ay s eem l o g i c a l to e x tc n d rh e s m a l l , g c n e r i c a l l yd c p . r u p e r a rpeo p u J ' h i s c o n r b i n a t i o no f l o c a l v a r i e t y
av er s ionof ant h ro p o g e n i ce x ri n c ri o n l a t i o n o f t h c s a m e s p e c i c sA
. l s o , r h e and geographi c horno gcneit y pr oo f populat ions ro rh e s u ffe ri n g a n d f e w c r t h e p o p u l a t i o n s t h a t r c t n a l l " t , ducesseveral pl easantbenef it s f or huu n t i m e l y d e a t h s o f i n d i v i d u a l s b c - t h e g r e a t e rt h c p n r b a b i l i r yo f t h c s i - m a n s .N o t o n l y a r e t h e e x o t i c s p e c i e s
c aus e popular i o n sa rc c o mp t-rs c do f m u l t a n e o u s ex t i n c t i o n ( r a n d o n r o r morc spcr-tacul ar,
bur the wor ld t r avi n d i v i d u a l s I. d o n o t b e l i e v et h i s s r c p n o t) r .rfai l pti pul ari ons,and thus of el cr can al w ays feel bor anically at
y d e s i ra b l efo r c o n s c rv a - the species.Hcncc, horv preci<;usa home. In addi ri on, nl any cit ies now
i s nc c es s ar or
tion biology . A l th o u g h d i s e a s ca n d p o p u l ati on i s i s a frrncti on of how havea grcaterdi versi tyo f plant f am s uf f er ing in an i m a l s a re u n p l e a s a n t m a n y s u c h p o p u l a t i o n se x i s r .
i l i cs and tree speci esthan did t he
a nd, per haps ,re g rc tta b l e ,b i o l o g i s ts
Ecologicalcomplex ity isgood. This ori gi nal [,abi tat destroyed r o m ake
recognize that conservarion is en- p o s rul ateparal l cl s the fi rsr one, but w ay for the ci ty. B ut th ese aest her ic
gaged in the prorecrionof rhe inregri- a s s u rnes
the val ue of habi tat di versi ry benefitsarc costly. The price is low
ty and c ont inu i ty o f n a tu ra l p ro c e s s - and complex ecologicalprocesses.
Ar- geographi cdi versi ty and ecological
es , not t he welf a re o f i n d i v i d u a l s .Ar ri v i n g at thi s j udgment may requi re compl exi ry.B ofani cal ga r dens,zoos,
the populat ion l e v e l , rh e i n rp o rta n r c o n si derabl csophi sri cati on,rrai ni ng, urban parks, and aquari a sat isf y,t o a
pr oc es s esar e u l ti ma te l y g e n e ri ca n d a n d thought. S omeonefami l i ar w i th degree,rny desire to be with orher
ev olur ionar y be c a u s erh e s ema i n ta i n d e s cri pti vepl ant and ani mal bi ogeog- speci es,but not my need t o see wild
the pot ent ialf o r c o n ri n u e de x i s te n c e . r a p h y ,t r o p h i cl e v e l sn, u t r i e n tc y c l i n g , and free creaturesor my cr aving f or
E v olut ion, as i t o c c u rs i n n a ru re , e d a p hi c hcterogenci tyo
and other as- sol i tudeor for a vari ety o f landscapes
could not pr oc e e dw i rh o u r rh e s u ffe r- p c c ts of ecol ogi calcl assi l i cati oni s i n and vi sras.
ing ins epar abl efro m h u n g e r,d i s e a s e , a b e tter posi ti on to ful l y appreci are
Euohfttott is good. Implicit in the
a nd pr edar ion.
th e c ompl exi tyi n a ti depoolor forest. thi rd and fourth functi on alpost ulat es
For this reason, biologisrs often
L i ke the fi rst one, thi s postul ate i s rhe assumpri onrhat th e cont inuit y
overcome their emotional idenrifica- expressesa preferencefor nature over of evolutionaryporential is good. Astion wit h indiv i d u a l v i c ti m s . F o r e x - a rti 6 ce, for w i l dernessover gardens sumi ng that l i fe i tsel f i s good, how
a m ple, t he bio l o g i s r s e e s rh e a b a n - ( c f . D u b o s 1 9 8 0 ) . W h e n p r e s s e d , can one mai ntai nan ethical neut r alit y
d o n e d f l e d g l i n g o r t h e w o u n d c d h o w ever,ecol ogi stscannot prove that about evol uti on? Li fe i tself owes it s
rabbit as part of the processof natu- their preferencefor natural diversity existenceand present diversity to the
ral selectionand is not deceivedthat should be the standard for managing evolutionaryprocess.Evolution is the
"rescuing"
s i c k , a b a n d o n e d , o r l ra b i tats.For exampl e,eveni f i t coul d machine,and life is its product. One
m aim ed indiv id u a l si s s e rv i n gth e s p e - be shown that a decreasein species possiblecorollary of this axiom is an
cies or the causeof conservation.(Sal- diversity led to desertification,eutro- erhical imperative to provide for the
vaging a debilitaredindividual from a p h i c a ti on,or the pi l i ng up of organi c continuationof evolutipnary processv er y s m all pop u l a ri o n w o u l d b e a n m a re ri al ,i t i s sti l l not a l ogi calconcl u- es in as many undisturbed natural
exception, assumingit mighr evenru- sion rhat such consequencesare bad. habitatsas possible.
ally contribute ro rhe gene pool.) F o r e xampl e,such eventsi n the past
Biotic diuersity has intrinsic ualue,
Therefore, the erhical imperative ro c re a tedfossi l fuel s (al rhoughnot ev- irrespective of its instrumental or
conserve speciesdiversiry is distinct e ry o ne w oul d argue that thi s w as utilitarian value. This normative posfrom any socieral norms about rhe g o o d ).
tulate is the most fundamental. In
v alue or t he w e l fa re o f i n d i v i d u a l
Ec o l ogi cal di versi ty can be en- emphasizing the inherent value of
anim als or plan rs . T h i s d o e s n o r i n hancedartificially,
but the increasein nonhuman l i fe, i r di sti n guishest he
-more
any way detract from ethical sysrems diversity can be
apparent rhan dualistic,exploitive world view from
that provide behavioral guidance for real (especially if cryptic taxa and a more unitary perspective: Species
hum ans on app ro p ri a tere l a ti < l n s h i p s associationsare considered, such as have val ue i n themselves,a value
wit h indiv idual s fro m o rh e r s p e c i e s , s o i l bi otas and mi crobi al communi - n ei t h e r c o n f er r ed n o r r e v o c a b l e ,
es pec iallywhen th e c a l l o u s b c h a v i o r ti e s ). In addi ti on, humans tend to bur springing fronr a species' long
oi humans causes animals to suffer sacrifice ecological and geographic evolutionaryheritageand potenrial or
unnec es s ar ilyC. o n s e rv a ti o na n d a n i - heterogeneityfor an artificially main- even from the mere fact of its exismal welfare, however, are conceptu- tained, energy-intensive,local species tence.2A large literature exists on this
ally dis t inc t ,an d th e y s h o u l d re m a i n diversity.Takeofor example,the large subj ect {D eval l and S essions1985;
numbers of plant taxa maintained in E hrenfel d1981; P assmo r e1974; Rolpnlitically separate.
the warm-ternperateand subtropical ston 1985, p. 718 thi s issue; Tobias
i
s
s
u
e
,
Ret ur ning r o th e p o p u l a ti o n
we m ight as k if a l l p o p u l a ri o n so f a ciries of the world. Most of these
g iv en s pec iesha v ee q u a l v a l u e .I rh i n k s p e c iesare horti cul tural vari eti esthat rHunters, loggers, and developers often express
n ot . T he v alue o f a p o p u l a ti o n , I do well in landscaped gardens and the samc love for naturc as dc professioual
but for rnany reasons, includb e l i e v e , d e p e n d s o n i t s g e n e t i c parks. One seesa great variety of such conservationists,
ing econornic ones, honorable people may be
uniqueness, irs ecological position, plants in Sydney,Buenos Aires, Cape unablc ro behave according ro rheir mosr chcrand the number of extanr popula- Town, Athens, Mexico City, Miami,
ishcd values, or they honestly disagree on what
tions. A large, generically polymor- and San Diego. But the roses,citrus, constirutcs edrical behavior.

December 1985

731

R e t e n t i o no f 9 0 % o f O r i g i n a lG e n e t i c
Variation for 200 \'ears

',
.l',

F
t:i
-1

-10

z
o

Foundrr tirc N. = l0

t-

fr20
zl&l

Foundrr rire N. = fg

B
C

Foundcr rir.e N. = (

100

xn

300

400

Joo

ULTIMATEEFFECTIVE
POPULATIONSIZE(K)
Figure2. Combinations of effectivepopulation sizesand generationlengths(in years)in
managed populations required to maintain at least 90"/" o( the geneticvariation that
existed in the source population; the program lasts 200 years. The calculationson
which the curves are based assumean intrinsic population growth rate of 1.0Y" per
year. For curve C, the founder size is equal to the ultimate size of the managed
population. Minimum founder sizesfor most speciesare in the range of l5 to 30 {from
Soul et al., in press).

1 9 85; and t he jou rn a l E n u i ro n me n ta l

Ethics).
Endlessscholarly debatewill proba b ly t ak e plac e a b o u t th e re l i g i o u s ,
e th ic al, and s c ie n ti fi cs o u rc e so f th i s
p o s t ulat e and ab o u t i ts i mp l i c a ti o n s
for policy and management.For exa m ple, does int ri n s i c v a l u e i m p l y
e g a l i t a r i a n i s ma n d e q u a l r i g h t s
a mong s pec ies A
? mo re p ro fi ta b l ed i s cussion would be about the rules to
he used when rlvo or more species
have conflicting interests(Naess

b i rd s wi l l have to be mai ntai nedarti fi c i a l l y i f they are to avoi d premature

e x t i n c t i o n( M y e r s 1 9 8 4 , S o u l 6e t a l . ,
i n p re ss).E ventual advancesi n techn o l o g y may enabl esome,i f not ntost,
such"speciisto be kept in a susperrde d , m i ni aturi zedstate,such as frozen
s p e rm , ova, and emhryos. Meanw h i l e , how ever, tradi ti onal w avs to
ma i n ta i n most of the pl anet' s megaf a u n am u s t b e i m p r o v e d .
l n recent y" tri , the breedi ng of
e n d a n gered speci es has undergone
p ro fo und chang,esas physi ol ogi sts
l98s).
i nvol ved.
have hecontea n d g eneti ci sts
try many
is
sponsored
research
Active
Contributions of conservation
new
Zoo,
l)iego
Sarr
At
the
zoos.
biology
techniques were developed for the
Rec ent ly , r apid p ro g re s s h a s b e e n d e t e r m i n a t i o n o f s e x i n s e x u a l l Y
m a d e b y z o o sa n d s i m i l a r i n s t i t u t i o n s m o n o m orphi c hi rd speci es(B ercovi tz
in the technology and theory of cap- e t a l . 1 978).Other w orkers (e.g.,B eni rs c h ke 1933) have found cytogeneti c
tive breedingof endangeredspecies.It
i s bec om ing ap p a re n t th a t n e a rl y explanations for the poor reproduc2000 speciesof large mammals and tive perforrnanceof several mammal

732

s p e c i e sR. y d er a n d W e d e m e y e{r1 9 8 2 )
pi oneeredretrospecti ve
ge net icanalr 's i so f c a p t i v es t o c k sw i t h t h e o h j e c t i v e
o f e q u a l i z i n gf o u n d e rr e p r e s e n t a r i o n .
A t r h e N a t i o n a l Z o o i n \ t h s h i r r-g} .troanv,e
DC. Ralls and Ballou {l9SJ)
provi ded i rrcontroverti b le evicler r ce
f o r t h e u n i v e r s a l i t lo' f i n b r e e d i n gd e p r e s s i o ni n m a m m a l s[ s e eN o r . e m b e r
1 9 8 4B i o S c i e n c3e4 : 6 0 5 - 6 1 0 . 6 l 2 J .
l rl any authors have appealed f or
l argerfoundersi zesi n gro ups of capt i v e l y b r e d a n i m a l st o m i n i m i z e i n breedi ng probl ems and t he loss of
geneti cvari abi l i ty(S enne r1980, Tem pl eton and R ead 1983), but specif ic
gui del i neshave been l acking. Recent
anal yseshave cl ari fi ed the int er r elati onshi psbetw e{n founder size and
severalother vari abl es,i ncf udinggeneration length, maximum captive
group si ze (carryi ng cap acit y) , and
group grorvth rate (Fi gur e 2) .
C onservati onbi ol ogy ha s also contri buted to the desi gn and m anagement of w i l dl and areas.An exam ple
i s the new fi el d of popul ation viabilit y
w hosegoal i s to e st im at et he
anal ysi s,
( e f f e c t i v e )n u m b e r o f i n d i v i d u a l s
neededto mai ntai n a sp ecies'longterm genetic fitness and ensure
againstextinction from other, nongenEtic causes.Several relatively independentpathwaysof researchin popul ati on bi ol ogy, commun it y ecology,
and bi ogeography
-*lrich are bei ng ioined in
I believJ will conthis effo-rt,
tri bute si gni fi cantl y to t heor et ical
population biology. One approach is
i o i ntegrate demograph ic st ochast ici ty (random vari ati on i n bir t h and
death rates and sex rati o) and environmental variation to predict the
p r o b a b i l i t yo f s u r v i v a l ( L e i g h 1 9 8 1 ,
S hafferand S arnson1985) . This approach is leading to very large estimatesfor l ong-termvi abilit v. s
Geneti csi s afsoi mportant in viabili t1' anal ysi s.A t l east i n out br eeding
speci es,
i t appearsthat re lat ivelvhet erozygousi ndi vi dual s are f r equent ly
more fit than relatively homozygous
ones.Many fi tnesscri teria have been
studi ed,i ncl udi nggrow th r at es!over w i nter survi val , l ongevit y, developmentalstabi l i ty, metabolic ef f iciency,
and scopefor growth (for revieu'ssee
B e a r d m o r e1 9 8 3 , F r a n k e l a n d S o u l 6
sGary Belovsky and Daniel Goodman, 1985,
personalcom m un ications. Un iversity of tvlichigan and Montana State UniversitY.

BioScienceVol. JJ No. I I

cited
i '9 trL and M it t on a n d G ra n t 1 9 8 4 ). g e re ds p eci es,A t thi s poi nr i n hi srory, References
Barrow'and
a
m
a
j
o
r
t
h
r
e
a
t
[-ande
t
o
s
o
c
i
e
t
y
i
s
a
n
d
n
a
t
u
r
c
Rur..fl
.George
B e a r d m o r e .J . A . 1 9 8L E x t i n c t i o r r ,s u r v i v a l .
ar e pr oposi n g th a t p o p u l a - te c h n o l ogl ' ,so i t i s appropri
ate that
a n d g e n e t i c v a r i a t i o n . F a g e s I 2 5 - | - 5| i n
' to
'. i i n u gha
C . N { . S c h n n e ' * ' a l d - C o xS, . i r { . C h : r m h e r s ,B .
sci encearrd
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a
n d { - o n s t r y + 7 t i o nB. e n j ; r m i r r . ( . u m m i n g s
f
o
r
q
u
a
n
t
i
t
a
t
i
v
e
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a n d l e g i s l a t i v ree s p o n s c s .
f,",,.ri.
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u
'
i
l
l
num
b
e
rs
b
e
n
e
e
d
e
d
Lar
ger
O
u
r
e n v i r o n m c n t a la n d e t h i c a l B e n i r s c h k eK, . l 9 f l . T h e i r n p ; r cot f r e c e a r c ho n
Qra i ts.
varf those
t h e p r o p a g a t i o no f e n d ; n q e r e d s p e c i e si n
f o r q u a l i t a t i v et r a i t s ,i n c l u d i n gg e n e t - p r o b l e m s , h o n ' e v e r , d r-t-he,
z o o s . P a g e s4 0 2 4 1 - 1 i n C . N l . S c h o n e r v a l d currrnt
i c p o l . " - r nor phis mTsh. e U S F o re s tSe r- fa c e d b y our ancesrors.
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