Professional Documents
Culture Documents
Michael E. Soulé
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Figure 1. Cancer biology and conservation biology are both synthetic, multidisciplinary sciences. The dashed line indicates the
artificial nature of the borders between disciplines and between “basic” and “applied” research. See text.
al parks should consider the impact of conservation biologists frequently fo holistic is the assumption that multi
the park on indigenous peoples and cus on individual endangered, critical, disciplinary approaches will ultimate
their cultures, on the local economy, or keystone species. ly be the most fruitful. Conservation
and on opportunity costs such as for Conservation biology tends to be biology is certainly holistic in this
feited logging profits. holistic, in two senses of the word. sense. Modern biogeographic analysis
There is much overlap between First, many conservation biologists, is now being integrated into the con
conservation biology and the natural including many wildlife specialists, servation movement (Diamond 1975,
resource fields, especially fisheries bi assume that ecological and evolution Simberloff and Abele 1976, Terborgh
ology, forestry, and wildlife manage ary processes must be studied at their 1974, Wilcox 1980). Population ge
ment. Nevertheless, two characteris own macroscopic levels and that re- netics, too, is now being applied to
tics of these fields often distinguish ductionism alone cannot lead to ex the technology of wildlife manage
them from conservation biology. The planations of community and ecosys ment (Frankei 1974, Frankei and Soulé
first is the dominance in the resource tem processes such as body-size 1981, Schonewald-Cox et al. 1983,
fields of utilitarian, economic objec differences among species in guilds Soulé and Wilcox 1980). Multidis
tives. Even though individual wildlife (Cody and Diamond 1975), pollina- ciplinary research, involving govern
biologists honor Aldo Leopold’s land tor-piant coevolution (Gilbert and ment agencies and wildlife biologists, is
ethic and the intrinsic value of nature, Raven 1975), succession, spéciation, also evident in recent efforts to illumi
most of the financial resources for and species-area relationships. Even nate the question of viable population
management must go to enhancing ecological reductionists, however, size (Salwasser et al. 1984).
commercial and recreational values agree that the proper objective of Another distinguishing characteris
for humans. The emphasis is on our conservation is the protection and tic of conservation biology is its time
natural resources. continuity of entire communities and scale. Generally, its practitioners at
The second distinguishing charac ecosystems. The holistic assumption tach less weight to aesthetics, maxi
teristic is the nature of these re of conservation biology should not be mum yields, and profitability, and
sources. For the most part, they are a confused with romantic notions that more to the long-range viability of
small number of particularly valuable one can grasp the functional intrica whole systems and species, including
target species (e.g., trees, fishes, deer, cies of complex systems without con their evolutionary potential. Long
and waterfowl)—a tiny fraction of ducting scientific and technological term viability of natural communities
the total biota. This distinction is studies of individual components usually implies the persistence of di
beginning to disappear, however, as (Levins and Lewontin 1985, chap. 6). versity, with little or no help from
some natural resource agencies be Holism is not mysticism. humans. But for the foreseeable fu
come more “ecological” and because The second implication of the term ture, such a passive role for managers
December 19 85 729
cal and histo rical ones or to prevail over adaptive, deterministic would preclude genetic differentiation
infrequent catastrophic events, such forces within populations. The sto among the colonies (Soulé 1980).
as inundation, volcanism, and glacia chastic factors in population extinc
tion. In other words, ecological pro tion have been discussed extensively The normative postulates. The nor
cesses belong to an intermediate scale (Shaffer 1981, Soulé 1983, Terborgh mative postulates are value state
of physical size and time (MacArthur 1974) in the context of the minimum ments that make up the basis of an
1972), and these processes begin to conditions for population viability. ethic of appropriate attitudes toward
fail or are overwhelmed near the ex The main implication of this postu other forms of life—an ecosophy
tremities of these ranges. late for conservation is that the prob (Naess 1973). They provide stan
Two major assumptions, or gener ability of survival of a local popula dards by which our actions can be
alizations, underlie this postulate. tion is a positive function of its size. measured. They are shared, I believe,
First, the temporal continuity o f habi One of the corollaries of this postu by most conservationists and many
tats and successional stages depends late is that below a certain population biologists, although ideological purity
on size. The random disappearance of size (between 10 and 30), the proba is not my reason for proposing them.
resources or habitats will occur fre bility of extinction from random de Diversity o f organisms is good.
quently in small sites but rarely, if mographic events increases steeply Such a statement cannot be tested or
ever, in large ones. The reasons in (Shaffer 1981). proven. The mechanisms by which
clude the inherent randomness of The next three corollaries are ge such value judgments arise in con
such processes as patch dynamics, netic. First, populations of outbreed sciousness are unknown. The concep
larval settlement, or catastrophic ing organisms will suffer a chronic tual mind may accept or reject the
events, as well as the dynamics of loss of fitness from inbreeding depres idea as somehow valid or appropri
contagious phenomena such as dis sion at effective population sizes of ate. If accepted, the idea becomes part
ease, windstorm destruction, and fire. less than 50 to 100 (Franklin 1980, of an individual’s philosophy.
The larger an area, the less likely that Soulé 1980). Second, genetic drift in We could speculate about the sub
all patches of a particular habitat will small populations (less than a few conscious roots of the norm, “diversi
disappear simultaneously. Species hundred individuals) will cause a pro ty is good.” In general, humans enjoy
will disappear if th eir h a b ita ts gressive loss of genetic variation; in variety. We can never know with cer
disappear. turn, such genetic erosion will reduce tainty whether this is based on avoid
Second, outbursts reduce diversity. immediate fitness because multilocus ing tedium and boredom or some
If population densities of ecologically heterozygosity is generally advanta thing else, but it may be as close to a
dominant species rise above sustain geous in outbreeding species (Beard- universal norm as we can come. This
able levels, they can destroy local prey more 1983, Soulé 1980, and refer is probably one of the reasons for the
populations and other species sharing ences cited below). (The genetic bases great popularity of zoos and national
a resource with such species. Out of these two corollaries may be the parks, which in recent years have had,
bursts are most probable in small sites same: homozygosity for deleterious, respectively, over 100 million and
that lack a full array of population recessive alleles.) Finally, natural se 200 million visitors annually in the
buffering mechanisms, including hab lection will be less effective in small United States. Perhaps there is a ge
itat sinks for dispersing individuals, populations because of genetic drift netic basis in humans for the appeal
sufficient predators, and alternative and the loss of potentially adaptive of biotic diversity (Orians 1980, Wil
feeding grounds during inclement genetic variation (Franklin 1980). son 1984). After all, humans have
weather. The unusually high popula The fourth functional postulate is been hunter-gatherers, depending on
tion densities that often occur in na that nature reserves are inherently a wide array of habitats and re
ture reserves can also increase the rate disequilibrial for large, rare orga sources, for virtually all of the past
of disease transmission, frequently nisms. There are two reasons for this. several million years.
leading to epidemics that may affect First, extinctions are inevitable in A corollary of this postulate is that
every individual. habitat islands the size of nature re the untimely extinction of popula
Taken together, the corollaries of serves (M acA rthur and W ilson tions and species is bad. Conservation
this postulate lead to the conclusion 1967); species diversity must be artifi biology does not abhor extinction per
that survival rates of species in re cially maintained for many taxa be se. Natural extinction is thought to be
serves are proportional to reserve cause natural colonization (reestab either value free or good because it is
size. Even though there is now a lishment) from outside sources is part of the process of replacing less
consensus that several small sites can highly unlikely. Second, spéciation, well-adapted gene pools with better
contain as many species as one large the only other nonartificial means of adapted ones. Ultimately, natural ex
site (when barriers to dispersal are replacing species, will not operate for tinction, unless it is catastrophic, does
absent), the species extinction rate is rare or large organisms in nature re not reduce biological diversity, for it
generally higher in small sites (Soulé serves because reserves are nearly al is offset by spéciation. Natural extinc
and Simberloff, in press). ways too small to keep large or rare tions, however, are rare events on a
The third functional postulate con organisms isolated within them for human time scale. Of the hundreds of
cerns the scale of population phenom long periods, and populations isolat vertebrate extinctions that have oc
ena: Genetic and demographic pro ed in different reserves will have to be curred during the last few centuries,
cesses have thresholds below which maintained by artificial gene flow if few, if any, have been natural (Dia
nonadaptive, random forces begin to they are to persist. Such gene flow mond 1984, Frankei and Soulé 1981),