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What Is Conservation Biology?

Michael E. Soul

BioScience, Vol. 35, No. 11, The Biological Diversity Crisis. (Dec., 1985), pp. 727-734.

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Fri Dec 21 11:10:37 2007
What is Conservation Biology?

A n e w synthetic discipline addresses the dynamics and problems

of perturbed species, communities, and ecosystems

Michael E. SoulC

C onservation biology, a new


stage in the application of sci-
ence to conservation prob-
lems, addresses the biology of species,
communities, and ecosysiems that are
Although crisis oriented,

conservation biology

likely that these areas harbored en-


demic bi0tas.l Reconnaissance later
confirmed this. The park boundaries
were established in 1981, and subse-
quent development has already pre-
perturbed, either directly or indirect-
is concerned with
cluded all but minor adjustments.
ly, by human activities o r other the long-term viability
Similar crises are now facing manag-
agents. Its goal is t o provide princi- ers of endangered habitats and species
ples and tools for preserving biologi- of whole systems
in the United States-for example,
cal diversity. In this article I describe grizzly bears in the Yellowstone re-
conservation biology, define its fun- gion, black-footed ferrets in Wyo-
damental propositions, and note a ommendations about design and ming, old-growth Douglas-fir forests
few of its contributions. I also point management before he or she is com- in the Pacific Northwest, red-cockad-
out that ethical norms are a genuine
V

pletely comfortable with the theoreti- ed woodpeckers in the Southeast, and


part of conservation biology, as they cal and empirical bases of the analysis condors in California.
are in all mission- or crisis-oriented (May 1984, Soult and Wilcox 1980,
disciplines. chap. 1).Tolerating uncertainty is of- Other characteristics of
ten necessary.
Conservation biologists are being conservation biology
Crisis disciplines asked for advice by government agen- As illustrated in Figure 1, conserva-
Conservation biology differs from cies and private organizations on such tion biology shares certain character-
most other biological sciences in one problems as the ecological and health istics with other crisis-oriented disci-
important way: it is often a crisis consequences of chemical pollution, plines. A comparison with cancer
discipline. Its relation to biology, par- the introduction of exotic species and biology illustrates some of these char-
ticularly ecology, is analogous t o that artificially produced strains of exist- acteristics, including conservation bi-
of surgery to physiology and war to ing organisms, the sites and sizes of ology's synthetic, eclectic, multidis-
political science. In crisis disciplines, national parks, the definition of mini- ciplinary structure. Furthermore,
one must act before knowing all the mum conditions for viable popula- both fields take many of their ques-
facts; crisis disciplines are thus a mix- tions of particular target species, the tions, techniques, and methods from
ture of science and art, and their frequencies and kinds of management a broad range of fields, not all biolog-
pursuit requires intuition as well as practices in existing refuges and man- ical. This illustration is also intended
information. A conservation biologist aged wildlands, and the ecological to show the artificiality of the dichot-
may have to make decisions or rec- effects of development. For political o m y between p u r e a n d applied
reasons, such decisions must often be disciplines.
Michael E. SoulC is an adjunct professor made in haste. Finally, this figure illustrates the
in the Wildland Management Center, Uni- For example, the rapidity and irre- dependence of the biological sciences
versity of Michigan School of Natural versibility of logging and human re- on social science disciplines. Today,
Resources. He is coauthor with Otto H. settlement in Western New Guinea
Frankel of Conservation and Evolution, for example, any recommendations
published by Cambridge University Press. (Irian Jaya) prompted the Indonesian about the location and size of nation-
Address correspondence to 4747 Black government to establish a system of
Mountain Road, San Diego, CA 92130. national parks. Two of the largest
O 1985 American Institute of Biological areas recommended had never been 'Jared M. Diamond, 1985, personal communi-
Sciences. visited by biologists, but it appeared cation. University of California, Los Angeles.

December 1985 727


1 -" CANCER - \
BIOLOGY

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.

a1 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 kevstone s~ecies. ly be the most fruitful. Conservation
and on opportunity costs such as for- ~ohservatiAnbiology 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 over la^ 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 macrosco~iclevels 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 (Frankel 1974, Frankel and Soul6
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 Soul6 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-plant coevolution (Gilbert and ment agencies and wildlife biologists, is
ethic and the intrinsic value of nature, Raven 1975), succession, speciation, 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- ecosvstems. The holistic a s s u m ~ t i o n tach less weight to aesthetics, maxi-
teristic is the nature of these re- of cdnservation biology should n'ot 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 o r 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

728 BioScience Vol. 35 No. 11


is unrealistic, and virtually all conser- biota. For example, the responses of tant as breeding or feeding sites for
vation programs will need to be but- prey to a predator's appearance or of animals mav initiate seauences of
tressed artificially. For example, even a phytophagous insect to potential causally linked events that ultimately
the largest nature reserves and nation- host plants are continually "tuned" lead to further extinctions (Frankel
al parks are affected by anthropogen- by natural selection. and Soul6 1981, Gilbert 1980, Ter-
ic factors in the surrounding area This postulate merely asserts that borgh and Winter 1980).
(Janzen 1983, Kushlan 1979), and the structure, function, and stability Introductions of generalists may re-
such refuges are usually too small to of coevolved, natural communities duce diversity. The introduction of
contain viable populations of large differ significantly from those of un- exotic plant and animal species may
carnivores (Frankel and SoulC, 1981, natural or svnthetic communities. It reduce diversity, especially if they are
Shaffer and Samson 1985). In addi- does not necessarily rely on determin- large or generalist species (Diamond
tion, poaching, habitat fragmenta- istic factors like density-dependent 1984, Elton 1958). Apparently, the
tion, and the influx of feral animals population dynamics or the molding larger the land mass, the less the
and exotic plants require extraordi- by competition of morphological re- impact of exotics (e.g., Simberloff
nary practices such as culling, eradi- lationships in communities over both 1980).
cation, wildlife immunization, habitat ecological and evolutionary time. In The evolutionary postulate and its
protection, and artificial transfers. addition. this ~ o s t u l a t eis neutral on corollaries formalize the evidence that
Until benign neglect is again a possi- the issue of holistic versus reduction- natural communities comprise species
bility, conservation biology can com- istic analysis of community structure. whose genetic makeups have been
plement natural resource fields in (In practice, a reductionistic method- mutually affected by their coexistence
providing some of the theoretical and ology, including autecological re- (Futuyma and Slatkin 1983, Gilbert
empirical foundations for coping with search, may be the best way to estab- and Raven 1975). An alternative the-
such management conundrums. lish the holistic structure of ory, the null hypothesis that commu-
communities.) nities are randomly assembled, is usu-
There are many "corollaries" of ally restricted t o "horizontal"
Postulates of conservation this postulate. Strictly speaking, most subcommunities such as guilds, spe-
biology of them are empirically based general- cific taxa, or trophic levels (e.g.,
Conservation biology, like many of izations. The following all assume the James and Boecklen 1984). In gener-
its parent sciences, is very young. existence of community processes as al, this latter thesis lacks empirical
Therefore, it is not surprising that its well as a coevolutionary component support, except that competitive
assumptions about the structure and in community structure. structuring within guilds or trophic
function of natural systems, and Species are interdependent. Not levels is often absent or difficult to
about the role of humans in nature, only have species in communities demonstrate (Strong et al. 1984), and
have not been systematized. What are evolved unique ways of avoiding that harsh environments or the vaga-
these postulates? I propose two sets: a predators, locating food, and captur- ries of dispersal may often be more
functional, or mechanistic, set and an ing and handling prey, but mutualis- important than biological interac-
ethical, or normative, set. tic relationships are frequent (Janzen
1975, Seifert and Seifert 1979). This
-

tions in determining local communitv


composition (e.g., Underwood and
The functional postulates. These are is not to say that every species is Denley 1984).
working propositions based partly on essential for community function, but The second functional postulate
evidence, partly on theory, and partly that there is always uncertainty about concerns the scale of ecological
" L
Dro-
on intuition. In essence, they are a set the interactions of species and about cesses: Many, if not all, ecological
of fundamental axioms, derived from the biological consequences of an ex- processes have thresholds below and
ecology, biogeography, and popula- tinction. Partly for this reason, Aldo above which thev become discontinu-
tion genetics, about the maintenance Leopold (1953) admonished conser- ous, chaotic, or suspended. This pos-
of both the form and function of vationists to save all of the parts tulate states that many ecological pro-
natural biological systems. They sug- (species) of a community. cesses a n d patterns (including
gest the rules for action. A necessary Many species are highly special- succession, nutrient cycling, and den-
goal of conservation biology is the ized. Perhaps the majority of animal sity-dependent phenomena) are inter-
elaboration and refinement of such species, including phytophagous in- rupted or fail altogether where the
principles. sects, parasites, and parasitoids, de- svstem is too small. Smallness and
The first, the evolutionary postu- pend on a particular host (Price randomness are inseparable.
late states: Many of the species that 1980). This means that the coattails Nonecological processes may also
constitute natural communities are of endangered host species can be dominate at the other end of the
the products of coevolutionary pro- very long, taking with them dozens spatial and temporal scale, in very
cesses. In most communities, species (Raven 1976) or hundreds (Erwin large or very old systems. In very
are a significant part of one another's 1983) of small consumer species large systems, such as continents, cli-
environment. Therefore, their geneti- when they go. matic and physiographic phenomena
cally based physiological and behav- Extinctions of keystone species can often determine the major patterns of
ioral repertoires have been naturally have long-range consequences. The the landscape, including species dis-
selected to accommodate the exis- extinction of major predators, large tribution. In very old systems, eco-
tence and reactions of a particular herbivores, or plants that are impor- logical processes give way to geologi-

December 1985 729


cal a n d historical ones o r t o brevail over adabtive. deterministic would preclude genetic differentiation
infrequent catastrophic events, such forces within pobulations. The sto- among the colonies (Soul6 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 198 1, Soule 1983, Terborgh mative ~ostulates are value state-
of physical size and time (MacArthur 1974) in the context of the minimum ments th'at 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 ~ositivefunction of its size. measured. They are shared, I believe,
First, the temporal continuity of 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 1 0 and 30), the proba- is not my reason for proposing them.
resources or habitats will occur fre- bilitv of extinction from random de- Diversity of 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 conceD-
larval settlement, or catastrophic ing organisms will suffer a chronic tual mind may accept or reject the
events. as well as the dvnamics 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 Soule 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 their habitats 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- thrng else, but it may be as close to a
dominant s~eciesrise above sustain- geous in outbreeding species (Beard- universal norm as we can come. This
able levels, ;hey can destroy local prey more 1983. Soule 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 s~ecies. 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 t'he 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 inherentlv a wide arrav of habitats and re-
ture reserves can also increase the rate diseqzrilibrial 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 a c A r t h u r a n d Wilson 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- ciallv maintained for manv 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, speciation, 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 dis~ersalare 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 (Soul6 serves because reserves are nearly al- is offset by speciation. 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- organ.isms
" 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 t o they are to persist. Such gene flow mond 1984, Frankel and Soul6 1981),

BioScience Vol. 35 No. 11


whereas the rate of anthropogenic phic population containing unique al- camellias, bougainvilleas, daffodils,
extinctions appears to be growing leles or genetic combinations has eucalyptus, and begonias are every-
exponentially. greater d u e , for example, than a where similar.
It may seem logical to extend the small, genetically depauperate popu- This combination of local variety
aversion of anthropogenic extinction lation of the same species. Also, the and geographic homogeneity pro-
of populations to the suffering and fewer the populations that remain, duces several pleasant benefits for hu-
untimely deaths of individuals be- the greater the probability of the si- mans. Not only are the exotic species
cause populations are composed of multaneous extinction (random or more spectacular, but the world trav-
individuals. I do not believe this step not) of all populations, and thus of eler can always feel botanically at
is necessary or desirable for conserva- the s~ecies.Hence. how ~ r e c i o u sa home. In addition. manv cities now
tion biology. Although disease and population is is a function of how have a greater diversity df plant fam-
suffering in animals are unpleasant many such populations exist. ilies and tree species than did the
and, perhaps, regrettable, biologists Ecological complexity is good. This original habitat destroyed to make
recognize that conservation is en- postulate parallels the first one, but way for the city. But these aesthetic
gaged in the protection of the integri- assumes the value of habitat diversity benefits are costly. The price is low
ty and continuity of natural process- and complex ecological processes. Ar- geographic diversity and ecological
es, not the welfare of individuals. At riving at this judgment may require complexity. Botanical gardens, zoos,
the population level, the important considerable sophistication, training, urban parks, and aquaria satisfy, to a
processes are ultimately genetic and and thought. Someone familiar with degree, my desire to be with other
evolutionary because these maintain descriptive plant and animal biogeog- species, but not my need to see wild
the potential for continued existence. raphy, trophic levels, nutrient cycling, and free creatures or my craving for
Evolution, as it occurs in nature, edaphic heterogeneity, and other as- solitude or for a variety of landscapes
could not proceed without the suffer- pects of ecological classification is in and vistas.
ing inseparable from hunger, disease, a better position to fully appreciate Evolution is good. Implicit in the
and predation. the complexity in a tidepool or forest. third and fourth functional postulates
For this reason, biologists often Like the first one, this postulate is the assumption that the continuity
overcome their emotional identifica- expresses a preference for nature over of evolutionary potential is good. As-
tion with individual victims. For ex- artifice, for wilderness over gardens suming that life itself is good, how
ample, the biologist sees the aban- (cf. Dubos 1980). When pressed, can one maintain an ethical neutrality
doned fledgling or the wounded however, ecologists cannot prove that about evolution? Life itself owes its
rabbit as part of the process of natu- their preference for natural diversity existence and present diversity to the
ral selection and is not deceived that should be the standard for managing evolutionary process. Evolution is the
"
rescuing" sick, a b a n d o n e d , o r habitats. For example, even if it could machine, and life is its product. One
maimed individuals is serving the spe- be shown that a decrease in s ~ e c i e s possible corollary of this axiom is an
cies or the cause of conservation. (Sal- diversity led to desertification, iutro- ethical imperative to provide for the
vaging a debilitated individual from a phication, or the piling up of organic continuation of evolutionary process-
very small population would be an material. it is still not a logical
" conclu- es in as manv undisturbed natural
exception, assuming it might eventu- sion that such consequences are bad. habitats as po&ible.
ally contribute to the gene pool.) For example, such events in the past Biotic diversity has intrinsic value,
Therefore, the ethical imperative to created fossil fuels (although " not ev- irresoective of its instrumental or
conserve species diversity is distinct eryone would argue that this was utilitarian value. This normative pos-
from any societal norms about the good). tulate is the most fundamental. In
value or the welfare of individual Ecological diversity can be en- emphasizing the inherent value of
animals or plants. This does not in hanced artificiallv. , , but the increase in nonhuman life, it distinguishes the
any way detract from ethical systems diversity can be more apparent than dualistic, exploitive world view from
that provide behavioral guidance for real (especially if cryptic taxa and a more unitary perspective: Species
humans on appropriate relationships associations are considered. such as have value in themselves. a value
with individuals from other species, soil biotas and microbial communi- neither conferred nor revocable,
especially when the callous behavior ties). In addition, humans tend to but springing from a species' long
of humans causes animals to suffer sacrifice ecological and geographic evolutionary heritage and potential or
unnecessarily. Conservation and ani- heterogeneity for an artificially main- even from the mere fact of its exis-
mal welfare, however, are conceptu- tained, energy-intensive, local species t e n ~ eA. ~large literature exists on this
ally distinct, and they should remain diversity. Take, for example, the large subject (Devall and Sessions 1985;
politically separate. numbers of plant taxa maintained in Ehrenfeld 198 1; Passmore 1974; Rol-
Returning t o the population issue, the warm-temperate and subtropical ston 1985, p. 718 this issue; Tobias
we might ask if all populations of a cities of the world. Most of these
given species have equal value. I think species are horticultural varieties that ZHunters, loggers, and developers often express
not. The value of a population, I do well in landscaped gardens and the same love for nature as do professional
believe, depends o n its genetic parks. One sees a great variety of such conservationists, but for many reasons, includ-
ing economic ones, honorable people may be
uniqueness, its ecological position, plants in Sydney, Buenos Aires, Cape unable to behave according to their most cher-
and the number of extant popula- Town, Athens, Mexico City, Miami, ished values, or they honestly disagree on what
tions. A large, genetically polymor- and San Diego. But the roses, citrus, constitutes ethical behav~or.

December 1985
species. Ryder and Wedemeyer (1982)
pioneered retrospective genetic analy-
Retention of 90% of Original Genetic sis of c a ~ t i v estocks with the obiective
Variation for 200 Years of equalizing founder representation.
At the National Zoo in Washington,
DC. Ralls and Ballou (1983) have
pro;ided incontrovertibie evidence
for the universality of inbreeding de-
pression in mammals [see November
1984 BioScience 34: 606-610, 6121.
Many authors have appealed for
larger founder sizes in groups of cap-
Founder size N, = 10
tivelv bred animals to minimize in-
breeding problems and the loss of
genetic variability (Senner 1980, Tem-
leto on and Read 1983). but s~ecific
'guidelines have been lacking. ~ e c e n t
A analyses have clarified the interrela-
tionships between founder size and
several other variables, including gen-
Founder s ~ z eN, = 20 eration length, maximum captive
group size (carrying capacity), and
group growth rate (Figure 2).
Conservation biology has also con-
tributed to the design and manage-
ment of wildland areas. An example
is the new field of ~ o ~ u l a t i oviabilitv
L L

n
ULTIMATE EFFECTIVE POPULATION SIZE (K) analysis, whose goal is to estimate the
(effective) number of individuals
Figure 2. Combinations of effective population sizes and generation lengths (in years) in needed to maintain a species' long-
managed populations required to maintain at least 90% of the genetic variation that term genetic fitness a n d ensure
existed in the source population; the program lasts 200 years. The calculations on against extinction from other, nonge-
which the curves are based assume an intrinsic population growth rate of 1.0% per netic causes. Several relatively inde-
year. For curve C, the founder size is equal to the ultimate size of the managed pendent pathways of research in pop-
population. Minimum founder sizes for most species are in the range of 15 to 30 (from ulation biology, community ecology,
Soul6 et al., in press). and biogeography are being joined in
this effort. which I believe will con-
tribute significantly to theoretical
1985; and the journal Environmental birds will have to be maintained arti- population biology. One approach is
Ethics). ficially if they are to avoid premature to integrate demographic stochasti-
Endless scholarly debate will prob- extinction (Mvers 1984. Soul6 et al.. citv (random variation in birth and
ably take place about the religious, in press). EveAtual advances in tech: deAth rates and sex ratio) and envi-
ethical, and scientific sources of this nology may enable some, if not most, ronmental variation to predict the
postulate and about its implications such species to be kept in a suspend- probability of survival (Leigh 1981,
for policy and management. For ex- ed, miniaturized state, such as frozen Shaffer and Samson 1985). This ap-
ample, does intrinsic value imply sperm, ova, and embryos. Mean- proach is leading to very large esti-
egalitarianism a n d equal rights while, however, traditional ways to mates for long-term viability.3
among species? A more profitable dis- maintain most of the planet's mega- Genetics is also i m ~ o r t a nin
t viabil-
cussion would be about the rules to fauna must be improved. ity analysis. At leas; in outbreeding
be used when two or more species In recent years, the breeding of species, it appears that relatively het-
have conflicting interests (Naess endangered species has undergone erozygous individuals are frequently
1985). profound changes as physiologists more fit than relatively homozygous
and geneticists have become involved. ones. Many fitness criteria have been
Contributions of conservation Active research is sponsored by many studied, including growth rates, over-
biology zoos. At the San Diego Zoo, new winter survival, longevity, develop-
techniques were developed for the mental stability, metabolic efficiency,
Recently, rapid progress has been determination of sex in sexuallv and scope for growth (for reviews see
made by zoos and similar institutions monomorphic bird species (Bercovitz Beardmore 1983, Frankel and Soul6
in the technology and theory of cap- et al. 1978). Other workers (e.g., Ben-
tive breeding of endangered species. It irschke 1983) have found cvtogenetic> " 'Gary Belovsky and Daniel Goodman, 1985,
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2000 species of large mammals and tive performance of several mammal gan and Montana State University.

732 BioScience Vol. 35 No. 11


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What Is Conservation Biology?
Michael E. Soul
BioScience, Vol. 35, No. 11, The Biological Diversity Crisis. (Dec., 1985), pp. 727-734.
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References cited

Associations Among Protein Heterozygosity, Growth Rate, and Developmental Homeostasis


J. B. Mitton; M. C. Grant
Annual Review of Ecology and Systematics, Vol. 15. (1984), pp. 479-499.
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Duties to Endangered Species


Holmes Rolston, III
BioScience, Vol. 35, No. 11, The Biological Diversity Crisis. (Dec., 1985), pp. 718-726.
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A Heliconia Insect Community in a Venezuelan Cloud Forest


Richard P. Seifert; Florence Hammett Seifert
Ecology, Vol. 60, No. 3. (Jun., 1979), pp. 462-467.
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Minimum Population Sizes for Species Conservation


Mark L. Shaffer
BioScience, Vol. 31, No. 2. (Feb., 1981), pp. 131-134.
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Population Size and Extinction: A Note on Determining Critical Population Sizes


Mark L. Shaffer; Fred B. Samson
The American Naturalist, Vol. 125, No. 1. (Jan., 1985), pp. 144-152.
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Island Biogeography Theory and Conservation Practice


Daniel S. Simberloff; Lawrence G. Abele
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Preservation of Natural Diversity: The Problem of Extinction Prone Species


John Terborgh
BioScience, Vol. 24, No. 12. (Dec., 1974), pp. 715-722.
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