Changing Concepts of The Balance of Nature Egerton 1973

CHANGING CONCEPTS OF THE BALANCE OF NATURE
BY
FRANK
N.
EGERTON
Kenosha,Wisconsin53140
of Wisconsin-Parkside,
University
ABSTRACT
The balance of nature has been a background assumption in natural historysince

antiquity, but even to the present it has seldom been closely studied. The idea of a
balance of nature emerged, but only implicitly,in antiquity. During the 17th century,
with an increased knowledge of natural history,the idea became a functional assumption, but within a theological rather than ecological context. In the 18th century
Linnaeus defined the concept and attempted to make it the foundation of an ecological
science. However, it remained tied to theology and was elaborated without critical
examination. The existence of agricultural pests, the occasional occurrenceof plagues
of animals, and the possibilityof species having become extinct were kinds of evidence
which would have been difficultto reconcile with contemporaryconcepts.
Lamarck was one of the few who perceived some anomaly, and he attempted to save
the old concept by arguing that fossilsrepresentedearly formsof existingspecies rather
than extinctspecies. His ideas were not widely accepted. Wallace found fault with the
Linnaean concept, but it was only a passing thoughtwhich he never published. Darwin
attempted to assimilate the balance-of-natureconcept into his description of natural
selection, but without exploring the inconsistenciesbetween the Linnaean concept and
his theory. As other naturalistsshifted their interpretationsof nature from static to
evolving,few of themappreciated the need to change theirunderstandingof the balance
of nature accordingly.
Some naturalists and ecologists who have thought seriously about the balance of
nature have postulated somewhat mysticalsupraorganismicconcepts. These, like the
general concepts,arose in antiquity,but unlike the general concepts theyhave won only
limitedacceptance.
Balance-of-natureconcepts apparently have receded in importance with the rise of
ecological specialization, probably because ecologists have developed more precise
concepts of productivityand ecosystemwhich can serve about the same explanatory
functions.
FIG. 1. The Peaceable Kingdom, BY EDWARD HICKS
Oil/canvas, 17Y2X 235/8";ca. 1840-1845; reproduced with permissionof The BrooklynMuseum, Dick S.
Ramsay Fund.
Balance of nature concepts have often been influenced by man's hope for harmony and stability.
Edward Hicks (1780-1849), an American Quaker preacher and self-taughtartist, illustrated this hope in
a series of about fiftysimilar paintings entitled "The Peaceable Kingdom." He was inspired by Isaiah 11:6,
which in the 1830's he modified to fithis time and place:
The leopard with harmlesskid laid down,

And not one savage beast was seen to frown,
The wolf did with the lion dwell in peace,
His grim carniv'rousnature there did cease;
The lion with the fatlingone did move,
A little child was leading them in love,
When the great PENN his famous treatymade,
With Indian chiefsbeneath the elm tree's shade.
322
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1973]
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INTRODUCTION
CIENTISTS attemptto explainnature
with generalizationswhich are called

hypotheses,theories, and laws. Scientificgeneralizationsare categorizedaccording to their degrees of comprehensiveness and verification,and a generalization
that has yet to be testedis called a hypothesis.
However, the use of this term implies that it
has been identifiedas one to be tested. Sometimesin the historyof science therehave been
concepts that have remained as background
assumptionsfor long periods of time without
anyone thinking that they needed testing.
These concepts are part of science, yet they
do not easily fitinto the descriptionsof science
by eitherThomas Kuhn or Karl Popper. Whatever disagreementstheremay be between these
philosophers of science, they agree that controversyplays a prominentrole in the development of science. Accordingto Kuhn (1970a, b),
controversyis characteristicof pre-paradigm
and of crisis stages in the development of a
science. According to Popper (1970), controversyshould ideally be a perpetual characteristic of science. Many examples from the historyof biology could be used to illustratetheir
theses. The works of Galen, Harvey, and
Darwin, and the controversiesover epigenesis
and spontaneous generationcome to mind.
Nevertheless,there are also many topics in
the historyof natural historythat seem to have
developed slowly through the ages without
significantcontroversyabout them. In such
cases the emphasiswas upon the accumulation
and organizationof knowledgein the Baconian
spirit, theoretical concepts being assumed
rather than being either tested or defended.
Three biological concepts which originated in
antiquity and had long historiesbefore being
questioned were the concepts of microcosmmacrocosm,the great chain of being, and the
balance of nature.
The microcosm-macrocosmanalogy had
Greek origins,but it flourishedmainly in the
16th century(Conger, 1922; Thorndike, 192358). The great chain of being had a longer
historyas an influentialconcept. Lovejoy (1936)
has shownthat it can be tracedback to implicit
beginnings in the writings of Plato and
Aristotle,and that it remained influentialin
[VOLUME
48
medieval and earlymodernthought. It became

explicit and most influentialin the middle of
the 18th century,but was finally discredited
when Cuvier published his embranchement
theoryin 1812. The longest historyof all is
that of the balance-of-natureconcept, which
had an equally ancient origin and was related
historicallyand logically to the other two. It
has survived into very recent times without
effective
challengeor testing.This is surprising
in view of the fact that the balance of nature
has been a fundamentaltheoreticalconcept in
ecology.
Except forLinnaeus in the 18thcenturyand
possibly A. J. Nicholson and CarringtonWilliams in the 20th century,therehave been few
attemptsto definethe balance of nature. This
deficiencycan be understood as the consequence of the concept having been a background assumptionratherthan a hypothesisor
theory. Any balance-of-natureconcept will assume an approximate stabilityin the populations of species, but since the authors themselves seldom made this assumptionexplicit,it
is not usually possible to state preciselywhat a
particular author's balance-of-natureconcept
encompassed. In tracing the history of this
concept, therefore,one does not progressfrom
one definitionto another,but fromone cluster
of relevantobservationsto another.
Important consequences have followed from
this lack of critical definition. One was poor
articulation of differentparts of the concept,
which led to the retentionof archaic assumptions long after they had become logically inconsistentwith other relevantand betterestablished data. Another was that the limits of
the conceptwere poorlyperceived,whichmeant
that there were opportunitiesfor adding uncritical illustrationsand amplifications.These
accretionsincluded loose analogies between the
balance of nature and other conceptual
units-an
organism, a biotic community,a
and even the universe. Some
political structure,
of these analogies can be classifiedas supraorganismicbalance-of-nature
concepts,but these
special concepts cannot always be easily separated from metaphors that were not meant
literally.
Still another consequence is that not only
were the assimilated elements poorly articu-

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325
lated, but other elements which should have

been assimilated into the concept were not.
A synthetic balance-of-naturetheory would
necessarilyhave included knowledgefromearth
and atmosphericsciencesas well as frombotany
and zoology. Occasionally,naturalistsindicated
an awarenessof this fact,but in practice there
was not, to myknowledge,any detailed attempt
to achieve such a comprehensive synthesis.
Therefore,this surveywill make only passing
referencesto relevantdevelopmentsin the earth
and atmosphericsciencesto indicate when they
could have become assimilated.
that into which destruction,too, happens according to necessity;for they pay penalty and
retributionto each other for their injustice
accordingto the assessmentof Time" (Kirk and
Raven, 1957, p. 117). There is ambiguityin
this statement,yet it foreshadowedthe Democriteanideas of the constancyand conservation
of force and matter. It also conveys a more
abstract impressionof the balance of nature
than is possible in an anthropomorphicmyth.
Practicallyall of the ideas of the pre-Socratic
natural philosophersthat can be related to the
balance of nature are similar to the ideas of
Anaximander in being concerned with forces
and matter,that is, with physical rather than
ANTIQUITY
biological phenomena.
In one formor another,a balance-of-nature It would be difficultto imagine the rise of
concept is a part of most cosmologies. When Greek philosophy and science unaccompanied
nature is viewed anthropomorphically,the by certain basic ideas that were congenial to
balance of nature is thoughtto be within the the developmentof balance-of-nature
concepts.
power of gods, and man assistsin maintaining The pre-Socraticphilosophers did indeed deit by prayer, sacrifice,and ritual. Through bate whetheror not nature was in a state of
these solicitations he hopes to insure good flux, but Greek science was built upon the
weather,bountiful crops, the avoidance of in- belief that nature is constant and harmonious
sect and rodentplagues, or an abundant supply (Vlastos, 1947; Sambursky,1964). The Pythagoof animals forthe hunt. The precariousnessof reans heard musical harmonyin the universe
the food supply in a primitiveeconomytaught (Kirk and Raven, 1957,pp. 229-259), and Greek
man that the constancyof nature could not be medicine taughtthe doctrinesof the balance of
taken for granted. In Greek civilization this humors and the healing powers of nature
stageof anxietyin man's thinkingwas preserved (Jones, 1946; Neuburger, 1943). Accordingly,
in - the myth of Demeter and Persephone ecological harmony and balance would have
(Hamilton, 1953, pp. 49-54; Rose, 1957, p. 27). been a compellingexpectation.
Demeter was the goddess of harvest and her
Herodotus was undoubtedlyindebted to the
daughter Persephone was the goddess of the natural philosophers for some of his own inSpring. When Hades, god of the underworld, terestin natural phenomena,but his answersto
abducted Persephone,her mother'ssorrowleft questions about nature tended to be concrete
the world in perpetual winteruntil the prayers and specificrather than abstract and general.
of men persuaded Zeus to intervene and ar- Furthermore,
in seekingan explanation forthe
range for Persephone'sreturnto earth for half
balance of nature within the living world he
of each year.
confrontedthe question in a differentrealm
With the developmentof pre-Socraticnatural
than they,and mere forces,whetheranimate or
philosophy such mythswaned in importance,
not,
must have seemed to him inadequate to
but the progressiverationalizationof nature by
for all aspects of life. Reports which
account
the Greekswas neverso extensiveas entirelyto
he
fromArabia interestedhim in the
collected
erase from their science a belief in anthropohow each kind of animal can
of
question
morphic forces. Such a belief, however, was
maintain
its
numbers.
Why did not the birds,
not incompatiblewith the idea of natural laws,
the hares? His answer
men
eat
all
and
beasts,
for that idea was derived from the idea of
human laws. This connection is illustratedin was that a superintending Providence had
the cosmologyof one of the firstnatural phi- created the differentspecies with differentrelosophers,Anaximander. He believed that "the productive capabilities. Predatoryspecies usufor existing things is ally had fewer offspringthan did the species
source of *;oming-to-be
326
THE QUARTERLY
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they ate. To illustrate his point he assembled

reports concerning the reproduction of snakes,
hares, and lions. His account is of great interest
because it represents the earliest biological
evidence for the balance-of-nature idea. It
should also be quoted in full, because we shall
refer back to it when discussing the writings of
Aristotle and Sir Thomas Browne.
The Arabians say that the whole world would
swarm with these serpents,if they were not kept
in check in the way in which I know that vipers
are. Of a truthDivine Providence does appear to
be, as indeed one mightexpect beforehand,a wise
contriver. For timid animals which are a prey to
othersare all made to produce young abundantly,
that so the species may not be entirelyeaten up
and lost; while savage and noxious creatures are
made very unfruitful. The hare, for instance,
which is hunted alike by beasts, birds, and men,
breeds so abundantly as even to superfetate,a
thing which is true of no other animal. You find
in a hare's belly, at one and the same time, some
of the young all covered with fur, others quite
naked, others again just fully formed in the
womb, while the hare perhaps has lately conceived afresh.
The lioness, on the other hand, which is one
of the strongest and boldest of brutes, brings
forthyoung but once in her lifetime,and then a
single cub; she cannot possibly conceive again,
since she loses her womb at the same time that
she drops her young. The reason of this is that
as soon as the cub begins to stir inside the dam,
his claws, which are sharper than those of any
other animal, scratch the womb; as the time goes
on, and he grows bigger, he tears it more and
more; so that at last, when the birth comes, there
is not a morsel in the whole womb that is sound.
Now with respect to the vipers and the winged
snakes of Arabia, if they increased as fast as
their nature would allow, impossible were it for
man to maintain himself upon the earth. Accordingly it is found that when the male and
female come together, at the very moment of
impregnation,the female seizes the male by the
neck, and having once fastened, cannot be
brought to let go till she has bit the neck entirely
through. And so the male perishes; but after a
while he is revenged upon the female by means
of the young, which, while still unborn, gnaw a
passage through the womb, and then throughthe
belly of their mother,and so make their entrance
into the world. Contrariwise,other snakes,which
are harmless,lay eggs,and hatch a vast number of
young. Vipers are found in all parts of the world,
but the winged serpents are nowhere seen ex-
[VOLUME
48
cept in Arabia, where they are all congregated

together. This makes them appear so numerous
(III, 108-109).
Although the report of superfetationin hares

is correct (Hediger, 1947), most of the remainder of this account is not. The winged
serpentscannot even be identified(How and
Wells, 1912, p. 204). Herodotus should have
been able to realize that the situation which
he describedfor lions would not maintain the
population,but would lead to rapid extinction.
One gets the impression that his informants
had gone overboard in attemptingto account
for the balance of nature entirelyby reproduction and predation. Nevertheless,the differential reproductivecapabilities of predatorsand
preyhas remaineda permanentpart of balanceof-natureconcepts, and the idea that species
were created with these capabilities was not
stronglychallenged before Darwin. The correlation that seemed to exist between the
reproductivecapabilityand the habits of different species seemed convincingevidence of the
providential arrangementof the balance of
nature.
This impressionwas strengthened
by Herodotus' reportof a mutuallybeneficialrelationship
betweenNile crocodilesand a speciesof plover:
This bird is of service to the crocodile and lives,
in consequence, in the greatestamity with him;
for when the crocodile comes ashore and lies with
his mouth wide open . . . the bird hops in and
swallows the leeches. The crocodile enjoys this,
and never,in consequence, hurts the bird (II, 68).
These reports left the strong impressionthat

nature is well regulated, but Herodotus did
not formulate an explicit balance-of-nature
conceptbased upon them.
The next contributor,Plato, can be accused
of going to the opposite extreme: his discussions were mainly theoreticaland were based
upon too few factual observations. There are
in Plato's Dialogues two creation myths that
were to be extremelyinfluential sources for
balance-of-natureconcepts. Such an origin is
not unusual. Popper (1962, p. 38) has pointed
out that "historicallyspeaking all-or nearly
all -scientific theories originate from myths,
and that a mythmay contain importantanticipations of a scientifictheory." The one in
Timaeus is the more comprehensive,but also

JUNE
1973]
the less precise of the two. This mythis the

source of two related concepts,the supraorganismicbalance-of-nature
conceptand the microcosm-macrocosmconcept. This creation myth
was related in answerto what mustnow appear
to be a very peculiar question: "In the likeness of what animal did the creatormake the
world?" The answer of Timaeus (30 c-d) was
that God did not make the world like any
one species, but rather as "one visible animal
comprehendingwithin itself all other animals
of a kindrednature."
Such a vague account coming from almost
anyone else might not have had much influence, but Plato's Dialogues have had an incalculable influenceupon Western thought. The
influenceof his mythseems to have been persistent,but the difficulties
involved in articulating the elements of any concept which is
based upon it seems to have prevented the
developmentof detailed explanations in those
concepts. The microcosm-macrocosm
concept
assertsthat the parts of the human body correspond to differentparts of the universe,and
the supraorganismicconcept asserts that the
differentliving beings are actually the organs
of a super-"being." Both are very mystical
concepts,and if it were not for their demonstrable influence,historiansof ecology would
undoubtedly be inclined not to take them
seriously.
For the veryreason that the supraorganismic
concept was vague and never elaborated in
much detail, it was seldom seen to conflictwith
otherecological ideas thatmightsimultaneously
exist. Plato himselfbuilt no explanatorybridge
between it and the creation myth told by
Protagorasin the dialogue of the same name.
At firstsight, the myth of Protagoras seems
more primitivethan that of Timaeus, because
the mythof Timaeus was a cosmologicalspeculation similarin many respectsto those of the
natural philosophers, whereas the myth of
Protagorasinvoked the old mythologicalgods,
Epimetheusand Prometheus.
But if one stripsaway the introductory
matter,Timaeus' idea is seen to be pure speculation,
whereas Protagoras' myth does contain some
observations,however superficial,with important generalizationsbased upon them. This is
the familiarmythabout the formationof the
differentkinds of animals which the foolish
327
god Epimetheusundertook. He bestowedtraits

upon each species which would assist it in
escaping its enemies, protecting it from the
weather, and enabling it to find adequate
nourishment.When he came to man, however,
he had run out of traits, and his brother
Prometheus had to suggest that man be endowed with superiorskills and with fire. The
mythfocused upon man, but of secondaryimportance was its assertionthat each species has
been providentiallyequipped with the means
forits survival.
The influenceof thismythwas much broader
than the one in Timaeus, and its influenceis
easier to follow. In neithercase, however,does
the scientificdevelopmentinspiredby the myth
follow the pattern outlined by Popper (1962,
pp. 50-51, 127-132), who described scientific
mythsas the origin of scientificdebates. The
myth in Protagoras reinforced the implicit
balance-of-nature
concept contained in Herodotus' History, and the effectmight well have
been to stifleratherthan to encourage debate.
Both accounts conveyed the idea that Providence had formed the differentspecies of
animals for a permanentassociationwith each
other, and that each was endowed with traits
that would enable it to continue to exist sucit might
cessfullyin spiteof whateverdifficulties
encounter from other species or from the
environment.
This view of animate nature, which I have
elsewhere called providential ecology (1968a,
p. 179), was teleological. It was thereforelogically in conflictwith the ideas on natural selection of Empedocles and Democritus. The potential debate between proponents of these
views did not, however,materialize. This was
not because either view ceased to attract defenders: Cicero and Plotinus were later to
expound their own ideas on providentialecology, and Lucretius was to provide the most
extensivedefenceof natural selectionto survive
fromantiquity. The debate that was to materialize was between natural selection and
Aristotle's teleological theory of nature.
Aristotlerejected the idea that the traitsof any
species are for the benefitof any other species
than itself (Balme, 1965), and thereforethe
debates over teleologywere generallyconfined
to discussions of the structureof individual
organisms.Aristotlewas not withoutecological

328
THE QUARTERLY
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awareness (Bodenheimer,1954), but his discussions of the balance of nature were brief,
indirect, and scattered among other subjects.
It is probable that the lack of prominence
throughoutsubsequent historywhich balanceof-nature concepts sufferedresulted in part
from his not having come to grips with the
subject more directlythan he did. Aristotle
was a great synthesizerand commentatorupon
Greek scientificthoughtfromThales down to
his own times, and one might have expected
that he would siftthroughthe various ideas on
the balance of nature and then offerhis own
argumentsand pronouncementson the subject.
One can, in fact,findin his De Generatione
et Corruptione the same kind of physical
concept that one findsin the
balance-of-nature
writingsof pre-Socraticphilosophers,a balance
of forcesand elements. The whole process of
and passing-awayhe believed
coming-into-being
was ultimatelycaused by the annual unequal
orbit of the sun around the earth and the
resultingseasons caused by the alternationsin
the amount of heat reaching the earth. But
when one turns to living nature, about which
he wrote so well, one is hard-pressedto findin
his writingseven an implicit balance-of-nature
concept. This silence may reflecta lack of
appreciationfor the importanceof the subject,
but thereis also the strongerpossibilitythat he
felt that his teleological theory,which focused
in itself to
upon the individual, was sufficient
accountforall aspectsof life.
Aristotlethoughtof the developmentof the
individual as being the outcome of its internal
plan of development strivingto organize the
raw materialswhich were to become the organism. Many featuresof a species which others
mightsee as fittingit for its ecological role he
explained as resultingfrom the limitationsof
development. Instead of extending the Epimetheanprincipleof the equitable distribution
of traits,he offeredinstead his own idea of
physiologicallimitation: "No animal that has
horns has also frontteeth in both jaws, those
in the upper jaw being deficient. For nature
by subtractingfromthe teethadds to the horns"
(De PartibusAnimalium,663b36).
Aristotle'sprinciple of physiologicallimitation could also explain the reproductivetraits
of different
species:
[VOLUME
48
Most trees,if theybear too much fruit,wither

away afterthe crop when nutriment
is not reand thisseemsto be what
servedforthemselves,
happens to annuals,as leguminousplants,corn,
and the like. For theyconsumeall theirnutrimentto makeseed,theirkindbeingprolific.
And
somefowlsafterlayingtoo much,so as even to
lay two eggsin a day,have died afterthis. For
boththe birdsand the plantsbecomeexhausted,
and this conditionis an excessof secretionof
residualmatter.A similarconditionis the cause
of the later sterilityof the lioness,for at the
firstbirthshe producesfiveor six, then in the
next year four,and again threecubs, the next
numberdown to one, thennone at all, showing
thattheresidueis beingused up and thegenerativesecretion
is failingalongwiththeadvanceof
years (De Generatione Animalium, 75iOa21-75iObl).
In this passage Aristotleemphasized food and

age as factorsinfluencingthe reproductiveoutput of organisms.In the same work (777b9-16)
he pointed out that the size of a viviparous
species would be the crucial factor for determiningthe lengthof its gestationperiod. Since
he felt that physiologicalcauses were sufficient,
he had no difficultyin rejecting Herodotus'
reporton lion reproduction: "The storyabout
the lioness dischargingher womb in the act of
parturition is a pure fable, and was merely
invented to account for the scarcityof the
animal" (Historia Animalium, 579b3).
However, since Aristotlewas only providing
indirectopposition to an idea that had not yet
even been explicitlyformulated,one should not
expect any militantopposition. He did not go
so far as to deny the existenceof such mutual
relationships in nature as were described in
Herodotus' report of plovers picking leeches
fromthe open mouths of crocodiles. Aristotle
liked that story and mentioned it in three
differentcontexts (Historia A nimaliurn, 612a20;
De Mirabilibus Auscultationibus, 831al 1-14;
Ethica Eudemia, 1236b9), and he also reported
that mutualism existed between the bivalve,
Pinna, and the crustacean, Pinnotheres (Historia Animalium, 547bl6-17). In another pas-
sage he accepted mutualismand implied that it

was compatible with his own physiological
concepts. He wrotethatthe mouthsof dolphins
and sharks were placed on the under surface
"not merely to provide a means of salvation
for other animals, by allowing them opportunityof escape during the time lost in the act

JUNE
1973]
329
tween these categories,and theywere not fully

distinguishedin the minds of naturalistsbefore
the 19thcentury(Egerton,1971).
Aristotleprovided no support,to the best of
concept.
myknowledge,forthe supraorganismic
In one case he suggested that both behavior
There were aspects of his biological thought
and environmental factors prevent overpopulathat could have been adapted to that purpose.
tion in minnows:
In his anatomical writingshe explained that
These fishes shed their eggs little by little, and
simple structureswere organized into organs,
. . . the males swallow the greater part of them,
and the organs were then integratedin strucand some portion of them goes to waste in the
meet the needs of the
water; but such of the eggs as the female deposits ture and function to
on the spawning beds are saved. If all the eggs animal (Historia Animalium, 486a5-14). He
also pointed out that gradation in the comwere preserved,each species would be infinitein
number. The greater number of these eggs so
plexityof species which later became the basis
deposited are not productive,but only those over for the "great-chain-of-being"
concept (Historia
which the male sheds the milt or sperm
Animaliurn, 588b; De Partibus Animralium,
(Historia Animalium,567a31-567b3)
681alO-15, 697bl-4; De Generatione Animalium, 732a25-733bl6). Reasoning fromanalogy,
Yet, Aristotle provided no theoretical context
it would have been easy to argue that these two
for these observations, and consequently their
of gradations are part of a larger whole,
sets
importance went unnoticed in antiquity. His
but he did not. Again, he might have been
most direct support of the balance-of-nature
saved fromdoing so by the focusof his teleology
idea came in his discussion of the reproduction
he observed
upon the individual. Furthermore,
of birds of prey:
that climate varied randomly without regard
The eagle lays three eggs and hatches two of to the needs of organisms(Physica,198bl7-33),
them . . . though occasionally a brood of three which is not what one might expect from a
has been observed. As the young ones grow, the
teleologicalworldsystem.
mother becomes wearied with feeding them and
The balance of nature having received only
extrudes one of the pair from the nest. At the
minimal attention from Aristotle,there were
same time the bird is said to abstain from food,
few other possibilitiesin antiquity for a synto avoid harryingthe young of wild animals...
thesisof knowledgeon the subject. The natural
The phene is said to rear the young one that has
by Pliny and Aelian both focused
histories
been expelled fromthe nest (Historia Animalium,
upon the propertiesof individual species, and
563a16-26).
these authors made little attempt to develop
Discussions such as this show in retrospect a
comprehensive biological theories. Nevertheneed not only for a greater skepticism about
less, these formsof literaturedid provide an
second-hand reports than anyone possessed in
opportunityto add to the knowledge of the
antiquity, but also a need for an ecological
balance of nature. This possibility may be
theory that would define the limits of species
illustrated from Aelian's account of the
interactions. Aristotle's teleological concept
jackdaw:
of turning . . . but also to prevent these fishes

from giving way too much to their gluttonous
ravening after food" (De Partibus Animalium,
696b24-33).
prevented him from exaggerating either the

balance in nature or mutualism, but primarily
by focusing his attention elsewhere. What was
needed was some relative assessment of the
importance of mutualism versus competition in
nature. However, before any answer could be
given, the question itself would have to be
discovered. When discussing the habits of animals, Aristotle did group his examples of competition separately from his examples of predation (Historia Animalium, 6O8b19-609b2O), but
he did not state explicitly the difference be-
The inhabitantsof Thessaly,of Illyria,and of

and have
Lemnosregardjackdawsas benefactors
decreedthat theybe fed at the public expense,
seeingthat jackdawsmake away with the eggs
and destroythe youngof the locustswhichruin
the cropsof the aforesaidpeople. The cloudsof
reducedand the
locustsare in factconsiderably
season's produce of these people remainsundamaged(III, 12).
If carefullycompiled, natural histories could
provide useful evidence for a theory,but there

330
THE QUARTERLY
REVIEW OF BIOLOGY
has never been much chance that a theory

would actually be advanced within such works
because they are organized around descriptive
knowledgeratherthan theory.
The most significanttheoretical discussions
of natural historyfromthe Roman era were in
the philosophical writings of Lucretius and
Cicero. The balance of nature idea received
implicit support fromLucretius' argumentfor
the constancyof nature,but that issue was no
longer in any doubt. Of greaterrelevancewas
Cicero's dialogue, De Natura Deorum, in which
the balance of nature became part of the Stoic
evidence for the wisdom and benevolence of
the Creator (bk. 2). In developing this argument, Cicero utilized the essential points from
the writings of Herodotus and Plato. The
balance of nature was maintained by the
ratesof reproduction,a
existenceof differential
different
place in nature for each species,physical traitsin each species which guaranteed its
survival, and mutual relationships between
certainspecies. Yet, his argumentdoes not add
up to a unifiedtheoryof the balance of nature.
These points remained merely a series of
evidences to illustratethe work of Providence,
as seen in thisexample:
In order to securethe everlastingdurationof
has mademost
divineprovidence
theworld-order,
carefulprovisionto ensurethe perpetuationof
the familiesof animalsand of treesand all the
vegetablespecies. The latterall containwithin
the property
themseed possessing
of multiplying
thespecies;thisseed is enclosedin theinnermost
partof thefruitsthatgrowfromeach plant;and
the same seeds supplymankindwith an abundanceof food,besidesreplenishing
theearthwith
a freshstockof plantsof the same kind (II, 51).
Since Cicero's dialogue presented different
points of view on the nature of God, there
mighthave existed here some potential for the
originof a scientificcontroversy.If the polemic
had developed, a scientifictheorymight have
emerged, as Popper suggested (1962). This
possibilityseems to be supported by the writings of the philosopher Plotinus, who wrote
about two centuriesafterwards.He achieved a
more dynamic idea of the balance of nature
while defending his own theological beliefs.
His goal was to reconcile the existenceof evil
with a belief in an omnipotentand benevolent
creator. Predation was one of the evils requir-
[VOLUME
48
ing an explanation. He explained that predation was essential in order for the greatest
diversityand quantity of life to come into
existence. The positive good of life, he concluded, more than justified the necessity of
death for the sake of feeding other life (Enneads,III, 2:15).
The discussionsby Cicero and Plotinus were
suggestive,but contained neither important
theoreticaladvances nor a definitivesynthesis.
The potentials of the theological avenue to
scientifictheorywere undoubtedlyreduced by
the achievement of a general consensus in
Christianity.
THE
17TH CENTURY
The Middle Ages, in so far as I have ascertained, was not a time in which there was
much discussionof the balance of nature. The
reason might be that a constantlysupervising
God, in which practically everyone believed,
would constantlysupervisethe workingsof nature. Elaborate built-in mechanisms within
nature would seem unnecessary.However,with
the advent of the ProtestantReformationthere
was a new interestin the details of the governance of both societyand nature. A new advancement of the balance-of-natureconcept
began in the 17th century. This advancement
involved the synthesisof old and new knowlconedge within a theological context-a
tinuationof providentialecology.
One of the firstto contributeto the elaboration of the concept in the 17thcenturywas Sir
Thomas Browne, who, good Baconian that he
was, attemptedin 1646 to separatetruescientific
knowledge from old wives' tales. One of the
questions which he investigatedwas whether
or not the whole world, or only a part of it,
had been inhabited before the flood of Noah.
This was only a variation upon an earlier
question concerningthe length of time that it
would have required to populate the world
fromthe time of Adam (Egerton, 1966). Since
the first patriarchs in the book of Genesis
allegedly lived for several centuries,Browne's
attention focused upon the relationship between longevityand rate of reproduction. He
pointed out that species such as man and the
elephant,which reproduce slowly,also tend to
be long-lived(1646, bk.6, ch.6). He thus added

JUNE
19731
the factorof differentiallongevityto the ones

which the ancients had recognized as maintainingthe balance of nature.
However, he did not then go on to explain
how all of the factorswhich had been mentioned mightwork in concert. His organizing
plan was skepticismrather than a theoretical
framework.Consequently,he did not synthesize
his various ideas relating to the balance of
nature.In anotherchapterhe examined Herodotus' report that the female viper kills her
mate aftercopulation and that the young later
kill her by gnawingthroughher womb. Browne
realized that the purpose of the reportwas to
explain why fierce animals do not become
abundant, and harmless ones scarce. He
doubted the truthof the report,however,and
suggestedthat Providence had a betterway of
limiting the numbers of fierceanimals.
If we more nearly considerthe conditionof
Vipers and noxious Animals,we shall discover
an higherprovisionof Nature:how althoughin
theirpaucityshe hath not abridgedtheirmaligeffected
it by their
nity,yethath she notoriously
insecessionor latitancy.For not only offensive
sects, as Hornets,Wasps, and the like; but
sanguineous corticatedAnimals, as Serpents,
Toads and Lizzards,do lie hid and betakethemselvesto covertsin the Winter(bk.3,ch.16).
Hibernation, he believed, served the dual
functionof providing man with some respite
from noxious animals and also restrictingthe
reproductiveseason for such species.
In the year 1662 another Englishman,John
Graunt, added still other factorsto the list of
those which were known to maintain the balance of nature. In his Natural and Political
ObservationsMentioned in a Following Index,
and Made upon the Bills of Mortality he
analyzed for the firsttime birth and death
statistics,and he discoveredthe statisticalregularityof the sex ratio and of most mortality
factors(excludingepidemics)in human populations (Egerton,1972). These discoveriescreated
a sensation amongst the intelligentsiaof Europe. Graunt himselfdid not receive a higher
education, and he was probably not widely
read in the natural historyliterature. He did
not indicate an awareness of the earlier discussions of the balance of nature, nor extensively exploit the implicationsof his own discoveries for this subject. In passing, he did
331
offerhis own suggestionthat the populations

of predatoryanimals,such as foxes and wolves,
might be kept within bounds by a limited
fertilitydue to excessive promiscuity(1662,
ch. 8).
Ideas from both Browne and Graunt were
incorporatedinto a comprehensivetreatiseby
Sir Matthew Hale. Hale's purpose, in The
Primitive Origination of Mankind (1677), was
to defend the Biblical story of creation by
disproving the Aristotelian assertion that the
earth was eternal. Hale believed he could
accomplish this by showing that, in spite of
man's slow rate of reproduction,the human
population had increasedsteadilysince the time
of creation. He even thoughthe could apply
Graunt's estimates of population increase in
England to the whole world and thereby
corroborateorthodoxestimatesof the age of the
earth(Egerton,1967,ch. 2).
For this reason Hale surveyedthe mortality
factorsfor various kinds of animals and for
man. In his opinion, these factorswere less
severe on man than on animals; therefore,he
thought that animal populations remain constant, while man's steadily increases. His was
probably the first comprehensive survey of
mortalityfactorsforanimals, and at the end of
that particular discussion he stated what appears to be the earliest explicit account of the
balance of nature. Before that,naturalistsand
philosophershad discussedonly aspects of it or
else had only alluded to it. Hale's language
was Aristotelian, but unlike Aristotle's De
Generatione et Corruptione,Hale's statements
referredspecificallyto animals, while also taking inanimatenatureinto account:
That the vicissitudesof Generationand Corruptionare by a kindof standingLaw in Nature
fixedin things,and the Notionsand Qualitiesof
Natural thingsare so ordered,to keep always
thatgreatWheel in circulation;and thereinthe
Accessand Recessesof the Sun, the Influxesof
the Heat thereofand of the other Heavenly
Bodies,and the mutualand restlessAgitationof
thosetwogreatEnginsin Nature,Heat and Cold,
of keepingon footthe
are the greatInstruments
and CorrupRotationand Circleof Generations
tions,especiallyof Animalsand Vegetablesof all
sorts.
That yet these Motions of Generationsand
and of the conduciblesthereunto,
Corruptions,
are so wiselyand admirablyorderedand con-

332
THE QUARTERLY
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[VOLUME
48
temperated,and so continuallymanaged and

orderedby the wise Providenceof the Rectorof
all things,that thingsare kept in a certaindue
stayand equability:and thoughthe Motionsof
Generationsand Corruptions,and the Instruments and Engins thereofare in a continual
course,neitherthe excess of Generationsdoes
theWorld,northedefect
oppressand over-charge
dothput a
thereof,
or prevalanceof Corruptions
Period to the Species of things,nor worka total
Dissolutionin Nature(1677,p. 211).
come accepted dogma. Ray remarked (1693,

p. 147) thatif extinctionhad occurred,it would
contradict the wisdom of the ages. He suggested instead that since the world was still
incompletelyexplored, the species known only
as fossilsmightstill live in unexplored regions.
This argumentwas unanswerableat that time,
and it prevailed for more than a century
(Greene, 1959,ch. 4).
The scope of Ray's two natural theology
books was very broad and thereforeprovided
Evidently Hale perceived no conflictbetween a good opportunityfor him to broaden the
the ancient providentialecology and the Aris- scope of the balance-of-nature
concept. To do
totelian systemof nature. By mergingthe two, so was not, however,his goal; as in Browne's
as he did in the above passage, he added an Pseudodoxia, the various elements that could
intermediate cause - heat fluctuations - to
have been united into a theorywere not. For
the purely biological means previouslyrecog- example,Ray discussed,in The Wisdomof God
nized as maintainingthebalance.
Manifested in the Works of the Creation, the
Interestingthough Hale's discussionwas, its hydrologiccycle and its utility (Biswas, 1970,
influenceon natural historyis uncertain. Al- pp. 186-187; Tuan, 1968). This was a kind of
though he presented an original and detailed environmentalbalance that might have been
synthesis,his knowledge was clearly bookish, tied to a discussionof the balance of animate
and naturalistsmight have felt that he had nature. Another of his discussions was in
little to say that was new. By the end of the defense of mountains (see Ray, 1713, p. 34).
17th century a number of naturalists were Some people thought that God had played a
making importantobservationsthat had rele- cruel trickon man by creatingthem. Ray devance for the balance of nature, and with the fended their existence by arguing that they
appearance of their work Hale's must have providedplaces to live forspecies of plants and
soon seemedout of date.
animals thatcould not live in low lands, and he
One of the most prominentof these natural- gave examples. In doing so he may have proistswas an Anglican clergyman,John Ray, who vided the firstreal evidence for the Protagoran
published two works on natural theologythat claim that each species has its own place in
were widelyread. One of the ecological themes nature.
for which he provided a comprehensivesurvey
Among the other ecological topics which
was of the survival mechanisms of different naturalistsinvestigatedlate in the 17thcentury
kinds of animals,forexample,the rapiditywith was parasitism. The development of microwhich young birds develop after hatching scopes had aroused an interestin insects and
(1691, pp. 92-102). This surveymay be seen other small animals. Antoni van Leeuwenhoek
as complementingHale's account of mortality was one of the most diligent of the naturalists
factors,but more likelyRay was respondingto who published importantobservationson such
the arguments of Robert Hooke (1665, pp. small animals. He and others discovered that
109-112; 1705) that petrifiedobjects represent parasiteswere more prevalentthan anyone had
the remains of plants and animals, some of previouslysuspected,and that theywere often
which are probablyextinct. The peculiarityof seriously detrimental or even fatal to their
fossilswas that theyfrequentlydid not resemble hosts (Bodenheimer, 1931). Before those realany known living species. If Hooke were cor- izations it had often been assumed that the
rectabout extinctions,thatfactwould represent relationship between parasite and host was
the firstpotential challenge to the balance-of- mutually beneficial. In fact, that assumption
nature concepts which had been developing persistedin the 18thcenturylong afterevidence
since antiquity. The Protagoran myth had was being accumulated to the contrary(Linimplied that all species have traitsthat prevent naeus, 1759, p. 102; Hoeppli, 1959, pp.
their extinction,and that implication had be- 164-165).
JUNE
19731

THE
18TH CENTURY
During the 18thcenturythe religiouscontext

of natural historyreceded in prominence,but
did not disappear. The belief in a superintending Providence continued to reinforcea
complacent attitude toward the idea of a
balance of nature. There was a greatexpansion
of knowledge about the general descriptions
and geographicalrangesof speciesof plants and
animals, and this knowledge was relevant for
furtherunderstandingof the balance of nature.
However, the contradictionswhich arose from
the expansion of knowledge did not lead to
any significantadjustmentsin the balance-ofnatureconcepts.
William Derham, an English clergymanand
naturalist who was a disciple of John Ray,
published in 1713 a widely read series of
lectures entitled Physico-Theology. His purpose, like Ray's, was to show how nature illustrates the wisdom and goodness of God. He
collected an unprecedentednumber of ecological examples to defend his thesis,and one of
his strongest evidences was the balance of
he claimed:
nature. Confidently
The Balanceof theAnimalWorldis, throughout
all Ages,kept even,and by a curiousHarmony
and just Proportionbetweenthe increaseof all
Animals,and the length of their Lives, the
World is throughall Ages well, but not over-stored
(3rded.,1714,p. 171).
This appears to be the earliestuse of the word
"balance" in this ecological context.
Like Hale, Derham brought into his argument both the vital statisticsof England and
the natural historyof animals. He implied that
forman the statisticalregularitiesfound in the
occurrences of births, marriages, and deaths
was evidence that these were controlledrather
than chance events(1714, pp. 174-175). He was
aware that birth and death statisticsindicated
that the population was increasing, but he
assumed that this was a provision of God for
future disasters. In discussing the regulation
of animal numbershe drew upon the familiar
factorsof differential
reproductivecapacityand
longevity,the diversityof the foods which
animals eat, the importance of predation for
checking the increase of some species, and
different
geographicaldistributionsfordifferent
species. In his account of foods and predation
333
he placed greater emphasis upon ecological

diversitythan in fact exists, with the result
that he overlooked the importanceof competition in nature (Egerton, 1971). On the other
hand, his exploration of ecological diversity
broughthim close to the importantconcept of
food chains (1714,pp. 183-193).
Having emphasizedthe harmonyand stability
of nature, Derham placed himselfin an awkward position fordiscussingplagues of animals.
He could only conclude that noxious species
"serve as Rods and Scorges to chastise us, as
means to excite our Wisdom, Care, and Industry"(1714, p. 55). Since he saw such plagues as
supernatural occurrences,he felt no need to
reconcilethemwith the balance of nature. The
precedentfor his interpretationwas, of course,
the accountof the Egyptianplagues in the book
of Exodus.
Derham's failure to integratethe occurrence
of animal plagues into his balance-of-nature
concept was probably not entirelythe fault of
his theologicalorientation. His contemporaries
who were less theologicallyoriented also failed
to formulate a comprehensive and detailed
balance-of-nature
concept. In studyingthe food
of fishes,Antoni van Leeuwenhoek came closer
than Derham to the concept of a food chain,
for he described several examples, but he did
not take the step from particular cases to a
generalization. He also carried out valuable
researchesinto the reproductivecapabilities of
insects and other animals, but none of these
studies resulted in significantgeneralizations
about the balance of nature (Egerton, 1968b).
Anothercontemporary,
Richard Bradley,was
less thoroughthan Leeuwenhoek,but was more
inclined to draw general conclusions. Bradley's
writingscontained some important ecological
discussions (Egerton, 1969), including interesting remarksabout the balance of nature. The
importance of ecological diversitymust have
been impressedupon him by both his horticultural experience and Derham's book. Bradley's observations indicated not merely that
"every Herb has its peculiar Insect," but also
that "the Insects which nature has designed to
preyupon the Flowerof a Plant will not eat the
Leaves, or any other Part of the same Plant.
The Leaves of Plants have theirInsectsnatural
to them, the Bark and Wood likewise" (1718,
recentincrease
pt. 3, pp. 58-59). Furthermore,

334
THE QUARTERLY
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[VOLUME
48
in the knowledge of parasites (Bodenheimer, even less than Aristotle could he ignore the
1931) caused him to suspect that these various ecological correlationswith reproductivecapacities. EchoingAristotle,Bradleyspeculatedthat
plant parasites
in 1000 yearsthe descendantsfromone fish,if
. . . have theirIcneumonFlies whichfeedupon
all reached maturityand reproduced, would
them,and by the help of fineTubes growing
about them,conveytheirEggsinto thebodiesof occupy a volume the size of the earth. Passing
the Nymphaor Caterpillars,where theyhatch beyond Aristotle, however, he went on to
as soonas theCaterpillars
are laid up in Aurelia, suggestthata proportionaterelationshipexisted
and feed upon them 'till they are themselves between the reproductivecapacity of fish and
and make theirway into the World. theirchances of survival(1721, p. 60). He also
perfected,
... and it may be these Insects which prey upon
discussed the available data on reproductive
others are not withoutsome others of lesser capacitiesof invertebrates,
birds,and mammals,
Rank to feed upon them likewise,and so to although whatever underlying pattern there
Infinity;for that there are Beings subsisting
mightbe eluded him. In spite of this,he conwhich are not commonlyvisiblemay be easily
if thereis any truthin a Micro- cluded that "all Bodies have some Dependance
demonstrated,
upon one another; and that everydistinctPart
scope(1718,pt. 3,pp. 60-61).
of Nature's Works is necessaryfor the Support
When farmersnear London began killing of the rest; and that if any one was wanting,
the birds in their fieldsbecause their turnips all the restmust consequentlybe out of Order"
had been devastated,Bradley convinced them (1721,p. 159).
that the real culpritswere multitudesof caterIn these writingsconsidered so far, Bradley
pillars, and that the birds came to eat these appears to have been developing a consistent
insects,not the turnips (1718, p. 58). In 1723 understandingof the balance of nature. Howone of his readers,S. C., sent him corroborating ever, that consistencywas more apparent than
observations. S. C. counted the number of real, as is to be seen when he discussedagriculcaterpillarswhich a pair of sparrowsbroughtto tural pestsin one of his later works. According
their young within an hour, which was forty. to the statements just quoted, one could
From this figure S. C. calculated that these assume that all species are importantfor the
sparrows brought 3360 caterpillars per week harmoniousworkingof nature. This idea un(Bradley,Aug. 1724, vol. 3, pp. 86-91). These doubtedly appealed to him as long as it
figures seemed to indicate that birds were remained an abstraction, but when he deextremelyimportantfor preventingvast devas- scended to the practicalmatterof raisingcrops,
tationsfrominsects.
he had no qualms about explaining how rats,
Most of Bradley's books were about the wasps,and otherverminshould be exterminated
practicaldetails of agricultureand horticulture, (Aug. 1724, vol. 3, pp. 92-95). He quoted with
but in 1721 he focused upon general natural approval a letterfromanother reader, Charles
history in A Philosophical Account of the Dubois, which reflectedthe farmer'sconstant
Works of Nature. This book, as its subtitle struggleagainst insectsand implied that whatindicated, was organized around the "great- everbalance theremay be is precarious. Dubois
chain-of-being"
concept. Some of its statements was concerned about the ravages of moths,
provideexcellentillustrationsof the connection perhaps codling moths (Carocapsa pomonella),
between that concept and the balance-of-nature amonghis apple trees:
concept. Being familiarwithAristotle'sinverse
If we considerthateveryone of theseMothswill
correlationbetween the size of mammals and
lay about 300 Eggsapiece,whichwill hatchinto
the number of offspringthey can produce,
then the DeCaterpillarsthe Springfollowing,
Bradley speculated that, "was it possible to
structionof an hundred of these Moths is
know the Nature of all Creatures,we might
preventingthe Increaseof 3000 murderingInfind,accordingto theirProportionand different sects,and so likewiseeveryCaterpillaror Insect
that a Bird destroysis preventingat least 300
Habit of Body, a continued Progressionwith
that would otherwisebe troublesometo us the
regard to their Size, Growth, and Length of
Year(Bradley,1724,p. 92).
following
Life" (1721, pp. 96-97). While this statement
expressed the chain of being physiologically,
Thus, Bradley was in the same situation as
JUNE
1973]
Derham. Both reportedobservationsthat could

not easily be reconciledwith theirunderstanding of the balance of nature; yet, since it remained a backgroundconcept, neithershowed
an awarenessof the conflictof ideas.
Throughout the 18th century there was a
steady increase in the knowledge of the life
historiesof plants and animals, and at various
timesthisknowledgewas related to ideas about
the balance of nature. However, it is impractical in this paper to surveyall such discussions. There were literary discussions as
well as scientific.Bradley's discussionof insect
parasites may have inspired Jonathan Swift's
famous rhyme(1733, lines 341-344):
Naturalists
a Flea
observe,
Hath smallerFleas thaton himprey,
AndthesehavesmallerFleas,to bite'em,
Andso proceedad infinitum.
This idea is scarcelycomprehensiveenough to
be much of a contributionto the conceptionof
a balance of nature. However, Bradley's observations,especially those in A Philosophical
Account of the Worksof Nature, may also have
encouraged Alexander Pope's elegant celebration of the great chain of being in his Essay on
Man, which suggests that all species are so
closely interdependentthat the extinction of
one would lead to the destructionof all living
nature (1733, epistle 1, sec. 8, lines 237-246).
The Swedish professor of natural history,
Carl Linnaeus, was one of the importantfounders of the science of ecology,and it was he who
firstgave a name to the balance of nature
concept. Before 1749 it had existed without
name. Linnaeus' contributionsto ecologywere
contained in essays writtenover a number of
years,some of whichwere defendedas thesesby
his students.
In one of his earlieressays,Oratio de Telluris
Habitabilis Incremento (1744), Linnaeus attempted to account for the stocking of the
world with plants and animals since the time
of Noah. He may have been inspiredby some
of the authors who discussed the longevityof
the patriarchs. Hale's PrimitiveOriginationof
Mankind was available in a German translation
(1683), but even if Linnaeus never saw it, there
were numerous other authors who also had
discussedthe subject (Egerton,1967). Linnaeus
reasoned that if God had originally created
335
only one or two individualsof each species,the

numbers of each species must have increased
steadily,just as man's population did. Hale
had not reached thisconclusionbecause he had
argued that the mortalityfactorsoperated more
severelyupon plants and animals than upon
man. Linnaeus supposed that the Garden of
Eden had been a mountain island on the
equator with arctic species living at the top,
tropical species at the bottom, and temperate
species in between. He furthersupposed that
each species increasedin numberseverygeneration, that as theyincreasedtheyalso spread out
over a larger area, and that the size of the
habitable earth increased as the populations
did. Given the last assumption, biological
competition need never become a significant
ecological factor.Among the unresolveddifficulties of this essay,its English translator,Brand,
worried about why the predatoryspecies did
not consume all the prey species before the
lattereverhad a chance to reproduce(Linnaeus,
1781,pp. 119-120).
The Oratio was vulnerable to criticismbecause its assumptionsabout the Garden of Eden
were based upon pure supposition. Nevertheless, it illustratesLinnaeus' propensityto organize his knowledge into comprehensive
explanations. Four years later he was more
successfulwith an essay entitled Oeconomia
Naturae (1749). It was this termwhich became
the firstname for the balance of nature, and
this essay was the firstsketch of a science of
ecology. Linnaeus used his economy-of-nature
concept as an organizingprinciple to unifyan
important, but previously amorphous, part
of natural history. In so doing, he was also
attemptingto transforman important background concept into the central theory of a
new science. Both his concept and his new
science deserveclose attention. (Such attention
is facilitated by von Hofsten, 1957; Stauffer,
1960; Egerton, 1967, ch. 3; and especially
Limoges,1972).
First,the concept. Linnaeus read the Swedish
translationof Derham's Physico-Theology,but
he did not follow Derham in the use of the
term "balance." Staufferhas traced Linnaeus'
term"oeconomia naturae" back to Sir Kenelm
Digby's discourseon the cure of wounds (1658;
see Stauffer,1960), but the true sources of
Linnaeus' ideas are the works from antiquity

336
THE QUARTERLY
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[VOLUME
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Darwinian questions were simply beyond the

scope of Linnaean inquiry. Accepting his
concept, one would not ask why particular
ecological relationshipsdeveloped or how they
might change, because one would assume that
such relationships had been created through
divine wisdom and that they would remain
unchanged. One could legitimatelyask how
such relationshipsworked, and Linnaeus did
so. The extentto whichhe asked and answered
these questions exceeded Derham's, although
his appreciation of the complexitiesseems to
have been about the same.
Organizationally,however, Linnaeus' treatment of the economyof naturewas betterthan
Derham's- better,thatis, if judged by modern
To perpetuatethe establishedcourseof nature
standards
of scientificexplanation. Derham,
in a continuedseries,the divine wisdomhas
the
minister,
used ecological examples to illusthoughtfit,thatall livingcreaturesshouldcontrate
divine
wisdom, while Linnaeus, the
in
stantlybe employed producingindividuals,
that all natural thingsshould contributeand professor,used them as the materials for a
lend a helping hand towardspreservingevery science. From the viewpoint of natural theolspecies,and lastlythatthe deathand destruction ogy, Derham had an elaborate explanatory
to the structure,
of one thingshouldalwaysbe subservient
but only bits of it could serveequally
restitution
ofanother(1759,p. 32).
well fromanotherviewpoint.
The Oeconomia Naturae begins with the
If this statementwere viewed as a justification
for predation, it would seem little different above-quoteddefinitionand then explains how
from Plotinus', but Linnaeus had much more that concept can be used to interpretphenomin inanimate nature and in the plant and
than that in mind. He was tryingto establish ena
animal
kingdoms. For both the plant and
a program which could serve as a guide for
animal kingdomsLinnaeus consideredpropagafutureobservations.
tion, preservation, and destruction as the
The supraorganismic concept might well
phenomena which maintained the economyof
have influencedLinnaeus' thinking,since the
nature. Limoges, by extending the coverage
mythof Protagorasclearlydid. However, Linof Linnaeus' conceptback in timeto the Oratio,
naeus did not give the formerfull recognition
has developed a pyramidalmetaphorto explain
or support. The version of the concept in
the concept.
Plato's Timaeus was probably too vague to
appeal to him. The reappearance of a supraOne can representthis theoryby imagininga
organismic concept in the 19th century was
pyramidin which the geographicaldistribution
of the species representsthe base, with the
based upon a new analogybetweenan organism
phenomena of propagation,preservation,and
and a biotic community,rather than between
the other three sides.
destructionrepresenting
an organismand nature as a whole as it had
The
which
the
are held together
surfaces
apex
by
been in Timaeus. Linnaeus possessed the conis the idea of proportion(1972,p. 10,n. 10).
cept of habitat (under the name of "station"),
but habitat is not a circumscribedunit in the According to Limoges, the Linnaean idea of
way that communityis, and the analogy would "proportion" (or balance) was not merely the
be less appealing when drawn between an effect of the interactions of propagation,
organism and a habitat than between an or- preservation,and destruction,but was actually
ganismand a biotic community.
an additional regulatoryprinciple. Since, howThe organizingvalue of Linnaeus' concept ever, this claim is based upon one brief stateof the economy of nature seems greater than ment in the Oratio (par. 96), I feel that it is
its explanatorypower. A whole series of post- unwarranted to incorporate that idea into
which have been discussed above, along with

natural histories from the 17th and 18th
centuries which belong to the same tradition
as those discussed above. The term "economy
of nature" bore an obvious similarityto the
contemporary term for animal physiology,
"animal economy," which involved studying
how the parts contributedto the functioning
of the whole. Linnaeus may,indeed, have had
in mind an analogy between the organs in an
animal and the species in a habitat,because his
analysisof the interrelationsbetweenthe plants
and animals in nature implied a close and
well-definedinteraction for the good of the
whole.

JUNE
1973]
Linnaeus' theoretical constructsof 1749 and

1760.
Linnaeus' theorizing in the Oeconomia
Naturae and the Politia Naturae (1760) was
mainly confinedto interpretingthe knowledge
which he had assembled and organized. Accordingly,novelty is more apt to be found
in his selection of data than in particular
theoreticalstatements. One discussion in the
Oeconomia Naturae which was evidentlynew
within the balance-of-naturecontext was on
plant succession(1759,p. 65). This phenomenon
mighthave led to some wondermentabout the
stability of habitats, but Linnaeus was able
to envision a full cycle fromlichens to mature
forest to the decay of trees, and perhaps a
repetition of the succession pattern. In the
Politia Naturae Limoges findsa new ecological
337
nature to the status of an explicit scientific

theorywhich was at the foundationof a branch
of science.
Unfortunately,it was not a science that

made rapid progress. Natural historywas not
to be so easily transformed.Most naturalists
continued to organize their observations as
before, into descriptivecatalogues of species.
A contemporary of Linnaeus, the famous
Buffon,effecteda compromise. His Histoire
naturelle included the traditional descriptive
catalogue of species, but Buffon's strong inclination for theoretical discussion found expression in occasional essays which were published with the descriptions. From studies on
both rodents and theirpredatorshe developed
a dynamicalperspectiveon the balance of nature. He reconciled the occurrenceof plagues
theory: ". . . the theory of the articulation of
of rodents with the balance of nature by
the functional groups in rapport with each postulating for a given species an oscillation
other, which is to say, beyond the interde- between extremes of rarity and abundance.
pendence of organismsin the system,the theory If certain forcesin nature permitteda species
of a subordinationof functions,of a hierarchy to become too abundant or too rare, then
of agents" (1972, p. 13, and in press). Linnaeus eventuallyother forceswould come into play
describedthis functionalhierarchyin termsof to restore the balance. The abundance force
an analogybetweenthe roles of different
classes was the reproductivecapacity of the species,
in human societyin comparisonwith the roles and the destructiveforceswere weather,predof various taxonomicgroups in nature. If this ators,and the quantityof available food (Bufidea is consideredto be a theory,it is one that fon,1756,p. 247).
was not well developed. As a metaphorit was
Buffonalso came to suspect that man might
another potential source of a supraorganismic have exterminatedsome of the large animals,
concept, but it was not a potential which such as the mammothsand mastodons,which
Linnaeus developed.
were known only as fossils (Buffon,1753, p.
The Politia Naturae was devoted primarily 173). John Ray's suggestionthat speciesknown
to discussions of the checks on populations only as fossilsmightstill be living in unknown
which preventsome species frombecoming too regions was becoming more and more difficult
numerous and exterminatingothers. Linnaeus to take seriouslyduring the second half of the
noticed the competition between different 18th century. By 1800, most naturalistswere
speciesof herbaceousplants in meadows,and he of the opinion that some species had become
concluded that if it were not for the selective extinct (Greene, 1959, chap. 4). The preservafeedingof insects,some species of plants would tion of all species had remained a definite
probably crowd others out of existence (par. tenet of balance-of-natureconcepts since the
12). Thus, although he had a concern for time of Plato, and since that assumptionwas
insects as agricultural pests (Bryk, 1924), he no longer tenable, the validity of all such
neverthelessemphasized their regulatoryfunc- concepts might legitimatelyhave been questions. He did not develop a critical method- tioned.
ology for determining to what extent his
Buffon'ssuccessorat the Jardin des Plantes,
observationscould serveas a basis forgeneraliza- Jacques-Henri Bernardin de Saint-Pierre,was
tions. Consequently, he, like Derham, often far frombeing the sort of scientistwho might
exaggerated the specialization of species. But have raised such a question. He was by temif Linnaeus achieved no dramatic theoretical perament a sentimental romantic, and the
innovations,he at least raised the balance of harmoniesof nature were so appealing to him
338
THE QUARTERLY
REVIEW OF BIOLOGY
that he eventually published a three-volume

descriptionof them (1815), though that work
was merelya continuationof similar ideas expressed in his ttudes de la nature (1784).
Bernardin de Saint-Pierreseems to have been
stronglyinfluenced by Linnaeus' economy-ofnature concept, but to have exaggerated the
harmoniousnessmuch beyond Linnaeus' thinking. One of his few ideas of value was derived
from his observationson the island of Mauritius. He saw that imported species could
destroythe natural balance of species on such
islands.
[VOLUME
48
Tout se conserve dans l'ordre 6tabli; les

perpetuelsqui
changemens
ct les renouvellemens
dans cet ordresont maintenusdans
s'observent
des bornesqu'ils ne sauroientdepasser;les races
des corps vivanssubsistenttoutesmalgr6leurs
variations;les progresacquisdansle perfectionnementde l'organisation
ne se perdentpoint; tout
anomalie,
cc qui paroit desordre,renversement,
rentresans cessedans l'ordregeneral,et m6mey
concourt;et partout,et toujours,la volontedu
sublimeAuteurde la natureet de tout cc qui
existeest invariablement
executee(1809,vol. 1,
p. 101).
Lamarck,therefore,
believed thatspecieschange
through
time,
but
that in spite of this the
which
it
noxious
insects
is
admitted,
There are,
our corn,nay,our persons. balance of nature remains essentiallythe same.
preyupon our fruits,
and locusts, Fossilsrepresentedold formsof existingspecies,
But if snails,nay bugs,caterpillars,
ravage our plains,it is because we destroythe and the extinctionof species, except for perbirds of our groveswhich live upon them; or, haps a fewof the largest,was merelyan illusion.
counthetreesof foreign
becauseon transporting
Lamarck's theoryof species changes not only
triesintoour own,such as the greatchestnutof
provided
an alternative to the possibilityof
India,the ebony,and others,we have transported
extinction,
but it also minimized the possiwhich
they
insects
with themthe eggs of those
likewise,thebirdsof bility of serious competition in nature. By
nourish,withoutimporting,
the same climate which destroythem. Every exaggerating the potentials of organisms for
countryhas those peculiar to itself,for the change and the inheritance of such changes,
of its plants (1797, vol. 1, ch. 7, Lamarck neglected an important factor for
preservation
p. 244).
any balance-of-natureconcept. A blindness to
the existenceof competitionhad existed among
If this was a new explanation of animal
naturalists since antiquity (Egerton, 1971).
plagues, it was surroundedby so much that was
During the second half of the 18th century
commonplace that it could have made no
several social commentatorsin England had
great impression.Neitherwould it have served
discussed competitionin society,and occasionto explain animal plagues in Europe.
ally they sought support for their arguments
fromreal or hypotheticalexamples fromnature.
THE 19TH CENTURY
However, the implicationsof these discussions
the subject of natural history was only
for
The conflictbetween the fact that species
slowly
appreciated by the contemporarynatuadcan become extinct, which was generally
ralists.
mittedby 1800, and the traditionalview of the
Alexander von Humboldt, writing during
balance of nature seems to have been realized
the
same period as Lamarck, made substantial
by fewnaturalists.One of thosewho did realize
to phytogeographyand other
contributions
the conflictof thoughtwas Lamarck, but his
resolution of it was never popular. It seems ecological subjects,yet even when he observed
probable that the question of extinctionmay conflictand change in nature he still emphahave influencedhis abandonmentof an earlier sized the preservationof harmonyand stability
belief in the stabilityof species. He may not (Egerton, 1970). The firstnaturalist to make
at firsthave realized the theoreticalrelation- significantuse of the conceptof competitionin
de Candolle, who
ship between the extinctionquestion and the naturewas Augustin-Pyramus
,of
the
distributions
plant species
explained
balance-of-nature concept, because in his
between
due
to
the
as
being
partly
competition
Systeme des animaux sans vertebresthe two
in
1816 he
On
to
a
species
(1820).
trip
England
pp.
(1801,
are
discussed
separately
subjects
22-23, 403-411); but in his Philosophie zoologi- had met Malthus (de Candolle, 1862, p. 265),
and there is a possibilitythat his awarenessof
que he explicitlyrelatedthem.
JUNE
1973]
the importanceof competitionin nature might

have been deepened by their conversation.
The meritsof de Candolle's essaywere fully
appreciatedby Charles Lyell, who utilized it in
his own even more comprehensivediscussion
of the distributionof species. As a geologist
Lyell was interested in both temporal and
spatial distribution,and the scope of his Principles of Geology (3 vols., 1830-33) enabled
him to bring togetherthe various aspectsof the
balance of nature concept which have been
discussed thus far. In rejecting Lamarck's
theoryof evolution,he also rejected Lamarck's
uncritical belief in the great capacity of individuals forchange. As an opponent of evolution Lyell found it necessary to explore as
thoroughlyas possible the causes of extinction.
Cuvier's idea of periodic catastropheshe found
no more convincingthan Lamarck's ideas, and
instead he placed his emphasis upon competition as a leading cause of extinction(Egerton,
1968c,p. 235).
While developing his ideas on the distribution and extinction of species, Lyell became
very much impressedwith the ecological writings of Linnaeus. Having assimilatedLinnaeus'
ideas, Lyell then made his own contributions
to the understandingof factorswhich tend to
preservethe balance of nature. Most notably,
he describedwhat can be called the buffering
effect. Its operation is conceptually simple.
It is a biological example of the law of supply
and demand, though Lyell did not draw this
analogy. His understandingof it is conveyed
in the following hypothetical examples.
339
peds, as rabbitsand fieldmice,

but also birds,
frogs,lizards,and insects,the profusionof any
one of these last may cause all such general
feedersto subsistmore exclusivelyupon the
speciesthusin excess,and the balancemay thus
be restored
(1832,pp. 138-139).
Lyell's grasp of ecological principleswas thus
based upon both an awarenessof the relevant
literature and some observationsof his own.
Although he possessed a fine critical mind, in
one instancehe uncriticallyfollowedLinnaeus'
example of describingthe extremedevastations
from insect plagues and then at the next
moment emphasizing the role of insects as
regulatorsof the populations of plants (1832,
p. 134). Before the extinction of species was
admitted,one mighthave reconciled this paradox in the styleof Buffon,but in the time of
Lyell the constantpreservationof the balance
of nature could no longer be so securely assumed. One of Lyell's readers who realized
this was AlfredRussel Wallace, who remarked:
Some species exclude all others in particular
tracts.Whereis the balance? When the locust
devastatesvast regionsand causes the death of
animalsand man,whatis the meaningof saying
the balance is preserved?[Are the devastations
of] the Sugar Antsin the West Indies and the
locustwhichMr. Lyell sayshave destroyed
800,000 men an instanceof the balance of species?
To humanapprehension
thereis no balancebut
a strugglein whichone oftenexterminates
another(McKinney,
1966,pp. 345-346).
Wallace wrote these remarks in his "species

notebook" around 1855. Here, at last, was
the kind of questioning which Popper insists
Althoughit may usuallybe remarkedthat the should be a
constant characteristicof science.
increaseof some one species is
extraordinary
followedand checkedby the multi- However, Wallace's doubts remained unpubimmediately
plicationof another,yet this is not alwaysthe lished until 1966. In his famousessay "On the
case,partlybecausemanyspeciesfeedin common Tendency of Varieties to Depart Indefinitely
on the same kinds of food,and partlybecause from the Original Type," which he sent to
manykindsof foodare oftenconsumedindiffer- Darwin in 1858, Wallace also raised the quesentlyby one and the same species. In the for- tion of the balance of nature, but from a
animals differentperspective. Under the influenceof
mer case, where a varietyof different
have preciselythe same taste,as for example, Malthus, he assumed that all species must
birds and reptilesde- produce more offspring
when manyinsectivorous
than could survive,and
vour alike some particularfly or beetle, the
he was primarilyinterestedin what the checks
usual numbersof the lattermay only cause a
of to increase were and what were the conseaugmentation
slightand almostimperceptible
each of thesespeciesof bird and reptile.In the quences of their operation. The balance-ofotherinstance,whereone animalpreyson others natureconcepthad receded fromits momentary
of almosteveryclass,as forexample,whereour prominence in his thinkingto its more usual
Englishbuzzardsdevournot onlysmall quadru- statusas a backgroundassumption.
340
THE QUARTERLY
REVIEW OF BIOLOGY
[VOLUME
48
Another British naturalist who had read

Lyell's Principles of Geology and who attemptedto reconcileextinctionwiththe balance
of nature was Edward Blyth. He was a careful
observer, but he seldom was able to bring
his observationsto bear upon his theoretical
ideas, and he thereforemade no permanent
theoreticalcontributions. He postulated that
a "physiological system" of nature exists in
which
truly new concept of balance in nature, and

Chambersincorporatedthis knowledgeinto his
interpretationof the fossilrecord. He argued
that land plants probablyhad to alter the composition of the air somewhatbeyond what the
aquatic plants had achieved before animals
could exist on land (p. 89); that since reptiles
require less oxygen than mammals,they were
able to flourishearlier; but with the alteration
of the atmosphere by terrestrialplants, the
mammals became dominant and the reptiles
. . . the existenceof one speciesis necessaryto receded. "In short,we see everywherethroughthatof another,and whichbindseach race to its
out the geological history,strong traces of a
locality;where the presenceof each is alike
necessaryto preservethe equilibriumof organic parallel advance of the physicalconditionsand
have the organic forms"(p. 150). As the conditions
being around; and when circumstances
changed,and the necessityfor its agencyno changed, species either became extinct or
longer remains,a whole race perishes (1836; evolved into new formssuitable for the new
1959,p. 124).
reprinted
conditions.
Chambers' general picture of an evolving
The brevityand vagueness of these thoughts world was a fine achievementfor an amateur
must have prevented them from being taken naturalist,yet he was vulnerable on many deveryseriouslyby anyone else, and Blyth never tails. One of the notable scientistswho leaped
expanded them in his other writings.
to the attack was the Cambridge professorof
Similar facile commentsabout the balance geology,Adam Sedgwick. He published a long
of nature could be quoted fromother natural- anonymousrefutationof Vestigesin which he
ists of the period, but without adding much took the unknown author to task for lacking
that would be new and useful. It seems un- scientificrigor. Nevertheless,Sedgwick's own
likely that verymuch that was new could have arguments were often something less than
been said on the subject until after the pre- scientific;one of his claims even bringsto mind
vailing ideas on the stability of species had the supraorganismicmythfrom Timaeus.
been successfullychallenged. The man who
is a miracleas incompreEach organicstructure
attemptedto create a comprehensivenew synhensibleas the creationof a planetarysystem;
thesis of knowledge on species, a synthesisinrelatedto all
is a microcosm
and each structure
volving the balance of nature, was Robert
otherworldswithinthekenofsense;yetgoverned
naturalist.
amateur
and
a
publisher
Chambers,
by laws and revolvingcycleswithinitself,and
His Vestigesof the Natural Historyof Creation
implied in the veryconditionsof its existence
(1844) was so unorthodox,fromboth scientific
(1845;p. 62).
and religiouspoints of view, that he dared not
sign his name to it. Prominentnaturalistspub- In spite of greatstridesin knowledge,orthodox
lished hostile reviewsof it, but to little avail. theory(and whetherthis is science,philosophy,
It went throughten editions by 1853. What or religion seems unclear) seems to have proChambers had to say about the balance of gressedin some respectsremarkablylittle.
nature was widely read, if not universally
The situation was ripe for Darwin's clean
respected.
sweep of obsolete, but persistent,theory,and
His evolutionary theory embraced all of one mightexpect his sweep to brush aside not
only obsolete species theory,but obsolete econature, thus being ecological in scope. He
achieved an explanation of a balanced nature logical ideas as well. This expectation is enin a changingworld. Since Priestleyhad pub- hanced by the knowledge that Darwin had
lished his firstpaper on plants and gases in read the ecologicalwritingsof Linnaeus, Buffon,
1772, many naturalistshad become aware of Humboldt, De Candolle, and Lyell, and by the
the reciprocalgas exchangesamong plants and fact that he was somethingof a skeptic.
During his voyage on the Beagle Darwin's
animals and the intermediateair. This was a
JUNE
1973]
341
geological outlook was shaped by Lyell's Prin- theory and some assumptions related to the
ciples, but in other respectshe was profoundly economy-of-nature
concept. In 1852 Darwin
influencedby Humboldt's Personal Narrative asked himselfwhy populations which become
(Egerton,1970). Darwin's Journalof Researches separateddivergeinto different
species.Limoges
(1839) and his "Notebooks on Transmutation" (1971) has suggestedthat the answer came to
(1837-39)indicate that he was at the time more him afterhe had read Milne-Edwards'lectures
stronglyinclined toward Humboldt's perspec- on the division of physiologicalwork among
tive of harmonyin nature than toward Lyell's the organs of an organism. EvidentlyDarwin
emphasis upon interspecificcompetition. He
drew an analogy in his mind between the relachanged his mind in October, 1838, afterread- tionshipof the organs to an organismand that
ing Malthus-recommended,
ironically, in between the organismsand their habitat. He
of all
Humboldt's writingss.Darwin then went on to then realized "that the modifiedoffspring
extend the idea of biological competitionto the dominantand increasingformstend to become
intraspecific
level (Egerton,1973).
adapted to many and highlydiversifiedplaces
Darwin's theoryof natural selection,with its in the economy of nature" (Darwin, 1959, p.
emphasis upon competitiondue to population 121). That a supraorganismicmental image was
pressure,implicitlycorrectedthe exaggeration part of his thinking on this subject is subof niche specialization which is found in the stantiated by a statement in The Origin of
writings of Plato, Derham, Linnaeus, and Species (1859, pp. 115-116): "The advantage of
others. Furthermore, Darwin's theory also diversificationin the inhabitants of the same
offereda new means of explaining the differ- region is, in fact, the same as that of the
ential reproductive capacities of species- in physiologicaldivision of labour in the organs
termsof theirsurvivalrequirements,which are of the same individual body-a
subject so
both physiological and ecological. This ex- well elucidated by Milne-Edwards." This was
planation thereforerepresentsa synthesisof a common type of supraorganismicstatement
Herodotus' environmental explanation and in being neitherprecise nor lengthy. It was a
Aristotle'sphysiologicalexplanation.
suggestionratherthan a proof,and would not
Limoges (1970, p. 151) has commentedthat have converteda skepticto a belief in a supraDarwin's theoryof natural selection replaced organismic theory. It might have reassured
Linnaeus' theory of the economy of nature. someone who already believed such a theory,
There can be little doubt about a significant but there is no clear reason to assume that
incompatibilitybetween importantelementsin Darwin was one of them.
the two theories,but the logical incompatibility
Undoubtedly Darwin's most importantconseems to have been overlookedby Darwin and tributionto the balance-of-natureconcept was
his contemporaries.Linnaeus' concepthad been his description of the high reproductive poso readily accepted as a truismthat even Dar- tential of species which is kept in check by a
win's skepticismdid not encompassit. In fair- varietyof mortalityfactors(1859, pp. 65-69).
ness to him, the economy of nature was a
comExcept forhis emphasisupon intraspecific
subsidiarypart of his argument and did not petition,much of what he said on the subject
receive the deep considerationfrom him that
was common knowledge. However, since this
might have exposed the logical contradictions.
discussionwas part of his theoryof evolution,
As a result,Darwin assimilatedwhat he should
therewas a new, albeit indirect,emphasisupon
either have rejected or else severelyqualified.
the instabilityof the situation. Yet, this genOne of the corrollariesof Darwin's theoryof
did
natural selection is the competitive-exclusion eral theoreticalemphasis upon instability
some
from
also
Darwin
not
lending
prevent
principle,which suggeststhat close competitors
do not share the same niche (Hardin, 1960). specific support to the balance-of-natureconOddly enough, it seems that this late addition cept.
His most conspicuous support of the conto his theorycame to him througha supraorcame in a descriptiveaccount, based in
cept
ganismic mental image, an image that might
not have occurredto him if he had previously part upon a paper by Newman (1850) and in
realized the incompatibilities between his part upon his own observations.
342
THE QUARTERLY
REVIEW OF BIOLOGY
[VOLUME
48
slight modifications,his account remained essentiallythe same throughall six editionsof the
Origin (Peckham, 1959, p. 157). A check on
the validity of Darwin's account was finally
undertakenin this centuryby W. L. McAtee.
He found that of the two species of forage
mentioned, Viola tricolor is commonly selfpollinated and Trifoliumpratenseis commonly
pollinated by honey-beesas well as by humblebees (McAtee, 1947). Furthermore,the importance of field mice in destroyingthe nests
of humble-beeshad been exaggeratedby Newman. In short,Darwin's account might have
occasional validity,but his investigationsinto
this subject had not been extensiveenough to
enable him to make reliable generalizations.
The major significanceof Darwin's cats-andclover account for the present discussion is
that it was an old-fashionedexplanation surrounded,in The Origin of Species,by radically
of natural history.By assonew interpretations
ciation, the account became part of the new
outlook; yet,since it really representedan old
and familiar idea, it was subjected to rather
less skepticismthan much else of what he wrote.
A popular story is often spun out into a
Darwin's scientificrevolutionhad considerably
progressively
more elaborate one the more it is less effectupon balance-of-nature
conceptsthan
repeated. Such was the fate of Darwin's ac- it should have had.
count. In later versions the cats were said to
In January,1860, HerbertSpencer published
belong to spinsterswho kept them for com- his essay, "The Social Organism," which depany; the women were spinsters because so scribedvarious parallels betweenorganismsand
many Englishmen were in the navy; and the societiesof differing
complexities.It was written
sailors ate dried beef that came from cattle too soon after publication of The Origin of
which grazed in the clover fieldspollinated by Species for Darwin's thoughts to have influthe humble-bees(Milne and Milne, 1960,chap. enced Spencer, who built upon his own more
1). In short, the elaborated version of this cosmictheoryof evolution. However,the strikbalance-of-naturestory implied that if the ing parallels which Spencer found could lead
spinstershad decided to confine their cats to to no definiteconclusionsbecause he was untheir houses, the British Empire would have able actually to tie social and biological evolucollapsed long before it did, because as the tion together with a definite law. The immice increased, the bees, clover, cattle, and plications of these parallels seemed substantial,
sailorswould all have decreasedin numbers.
however, in suggesting a new evolutionary
Few if any naturalists or ecologists would microcosm-macrocosm
concept. Yet, for all his
have believed the most extremeconclusionsin interestin biology,Spencer was a philosopher
the above story,but I have mentionedthem to and not a scientist,and by 1860 it was already
show how easily a good storycan get of hand. impractical to build scientifictheories upon
If it is appealing, the common urge is to speculation rather than first-handresearch.
embellishit ratherthan check its validity. The
Nevertheless,the writingsof a popular philosofirstskepticalreactionto Darwin's accountcame pher mightwell have influencedthe thinking
in Robert Owen's anonymoushostile reviewof of scientistswho read his works.
The Origin of Species (Owen, 1860, p. 494).
Whether or not Stephen A. Forbes was one
Although Darwin subsequently made some of these,he at least found the old idea of the
From experiments which I have tried, I have
found that the visitsof bees, if not indispensable,
are at least highly beneficial to the fertilisation
of our clovers, but humble-bees alone visit the
common red clover (Trifolium pratense), as other
bees cannot reach the nectar. Hence I have very
little doubt, that if the whole genus of humblebees became extinct or very rare in England, the
heartsease and red clover would become very
rare, or wholly disappear. The number of
humble bees in any district depends in a great
degree on the number of field-mice,which destroy
their combs and nests; and Mr. H. Newman, who
has long attended to the habits of humble-bees,
believes that "more than two-thirdsof them are
thus destroyed all over England." Now the
number of mice is largely dependent, as every
one knows,on the number of cats; and Mr. Newman says, "Near villages and small towns I have
found the nests of humble-bees more numerous
than elsewhere,which I attribute to the number
of cats that destroythe mice." Hence it is quite
credible that the presence of a feline animal in
large numbers in a district might determine,
through the interventionfirstof mice and then
of bees, the frequencyof certain flowersin that
district!(1859,pp. 73-74)

JUNE 1973]
microcosm-macrocosm serviceable in some form,

for he entitled a well-known address in 1887
"The Lake as a Microcosm." Forbes was a very
capable field naturalist who conducted ecological studies on the lakes of Illinois and
summarized his findings in the address. He
explained:
The animals of such a body of water are, as a
whole, remarkably isolated -closely
related
among themselvesin all their interests,but so far
independent of the land about them that it every
terrestrialanimal were suddenly annihilated it
would doubtless be long before the general multitude of the inhabitants of the lake would feel
the effectsof this event in any important way.
It is an islet of older, lower life in the midst of
the higher, more recent life of the surrounding
region. It forms a little world within itself-a
microcosmwithin which all the elemental forces
are at work and the play of life goes on in full,
but on so small a scale as to bring it easily within
the mental grasp.
Nowhere can one see more clearly illustrated
what may be called the sensibility of such an
organic complex, expressed by the fact that whatever affectsany species belonging to it, must have
its influence of some sort upon the whole assemblage (p. 537).
It may be that this statement reflects the influence, not only of Darwinism and earlier balance-of-nature concepts, but also of the bioticcommunity concept. This latter concept had
been developing for several decades before finding explicit expression in Karl Mobius' treatise
on oysters (1877; Allee, Emerson, Park, Park,
and Schmidt, 1949, pp. 34-35).
Be that as it may, ecology was still an infant
science when Forbes made these remarks, and
the techniques for describing quantitatively the
interrelations which he discovered were as yet
undeveloped. He had to confine himself to the
less complex descriptions of the physical conditions of the lakes, the kinds of life found in
them, the food chains, and the effects of
changing seasons. These descriptions were undoubtedly useful, but were not exhaustive
enough for one to imagine that he had actually
proven his conclusions.
Perhaps no phenomenon of life in such a
situation is more remarkable than the steady
balance of organic nature, which holds each
species within the limits of a uniform average
343
number, year after year, although each one is

always doing its best to break across boundaries
on every side. The reproductive rate is usually
enormous and the struggle for existence is
correspondingly severe. Every animal within
these bounds has its enemies, and Nature seems
to have taxed her skill and ingenuity to the
utmost to furnishthese enemies with contrivances
for the destructionof their prey in myriads.For
every defensivedevice with which she has armed
an animal, she has invented a still more effective
apparatus of destruction and bestowed it upon
some foe, thus strivingwith unending pertinacity
to outwit herself; and yet life does not perish in
the lake, nor even oscillate to any considerable
degree, but on the contrarythe little community
secluded here is as prosperous as if its state were
one of profound and perpetual peace. Although
every species has to fight its way inch by inch
from the egg to maturity, yet no species is
exterminated,but each is maintained at a regular
average number which we shall find good reason
to believe is the greatestfor which there is, year
after year, a sufficientsupply of food" (p. 549).
It is doubtful that Forbes intended to imply
that no species is ever exterminated; yet, as it
stands, this conclusion sounds remarkably like
an echo of Herodotus and Plato. However, the
cause which Forbes invoked, unlike Herodotus,
was not divine providence but rather "the
beneficent power of natural selection which
compels such adjustments of the rates of destruction and of multiplication of the various
species as shall best promote this common
interest" (p. 550). It seems that Forbes was
placing an extremely strong emphasis upon the
stabilizing effects of natural selection. The
causes of the balance of nature might change
with different scientific theories, but the existence of that balance was not questioned. The
time dimension which was so important for
Darwin's theory of evolution was not carried
over into the discussions of the balance of
nature.
THE
20TH
CENTURY
During the 20th century ecology became a

coherent science, or perhaps two related coherent sciencesplant ecology and animal
ecology. There was no science built, as Linnaeus would have had it, around the balanceof-nature concept. Instead, a number of descriptive approaches were developed, to some

344
THE QUARTERLY
REVIEW OF BIOLOGY
of which the balance-of-nature concept could

be tied, but apparently without being essential. These include studies of biotic communities, populations, productivity, and ecosystems. Because of the volume of modern
ecological literature, the discussion of this
century will be even more limited than the
foregoing discussions of the concept during the
previous two centuries. However, since this
discussion does include some of the more important publications, it is probably sufficiently
representative.
After ecologists accepted the community concept, one of them soon formulated a supraorganismic-community concept. The plant ecologist, Frederic Clements, defended this concept
in several publications, particularly in his
Plant Succession (1916). There he drew both a
morphological and a developmental analogy
between organisms and communities.
The unit of vegetation,the climax formation,is
an organic entity. As an organism, the formation arises, grows,matures,and dies. Its response
to the habitat is shown in processes or functions
and in structureswhich are the record as well
as the result of these functions. Furthermore,
each climax formationis able to reproduce itself,
repeating with essential fidelitythe stages of its
development.The life-historyof a formationis a
complex but definite process, comparable in its
chief features with the life-historyof an individual plant. The climax formationis the adult
organism, the fully developed community, of
which all initial and medial stages are but stages
of development. Succession is the process of the
reproduction of a formation,and this reproductive process can no more fail to terminatein the
adult form of vegetation than it can in the case
of the individual plant (pp. 124-125).
At least one ecologist, Phillips (1934-35),
strongly supported these developmental analogies, but William Cooper bluntly rejected them
(1926, pp. 399-400), and A. G. Tansley found
them unsatisfactory (1935, pp. 289-299). Popperian controversy had finally begun. Charles
Elton stated in more general terms:
"The balance of nature" does not exist, and perhaps never has existed. The numbers of wild
animals are constantlyvaryingto a greateror less
extent,and the variations are usually irregularin
period and always irregular in amplitude. Each
variation in the numbers of one species causes
[VOLUME
48
on the numbers
directand indirectrepercussions
of the others,and since manyof the latterare
varyingin numbers,
themselvesindependently
(1930,p. 17).
is remarkable
confusion
theresultant
EvidentlyClementstook seriouslythe criticisms
fromCooper and Tansley. In a paper written
in 1936 on "Nature and the Structureof the
less explicit
Climax," Clementswas significantly
in drawing his analogy between the development of individualsand biotic communities(p.
257), and when he collaborated with Victor E.
Shelford on Bio-Ecology, his developmental
analogies were omitted. Instead, the co-authors
quoted the above passage fromElton (Clements
and Shelford,1939,p. 173).
Clements' apparent abandonment of his
earlier claim mightseem to indicate the death
of the supraorganismicconcept, and perhaps
even that of the whole balance-of-natureconcept. Life, however, lingered in both. The
possibilityof such an easy resolution of the
question was prevented by the fact that the
concept had never been adebalance-of-nature
quately defined or defended. Therefore it remained elusive.
New opportunitiesfor defendingbalance-ofnature concepts would constantlyarise. For
example, the homeostasismechanismsin warmblooded vertebratesprovided the substancefor
a new analogy,which became part of the most
authoritative modern defense of a supraorganismicbalance of natureconcept.This occurs
in that great compendium by the Chicago
professorsAllee, Emerson, Park, Park, and
Schmidt,Principles of Animal Ecology (1949).
Apparentlyunder the influenceof their own
studies on termites,ants, and other social
animals,theyconcluded that:
maintainsa certainbalance,
. . .the community
a bioticborder,and has a certainunity
establishes
parallelingthe dynamicequilibriumand organizationof otherlivingsystems.Naturalselection
resystem,
operatesupon the whole interspecies
sultingin a slowevolutionofadaptiveintegration
and
and balance. Divisionof labor,integration,
characterize
the organismand suprahomeostasis
organismicintraspeciespopulation. The interspeciessystemhas also evolvedthese characteristicsof the organismand maythusbe called an
(1949,p. 728).
ecologicalsupraorganism
Henry (1955) and Bodenheimer(1958, pp. 190201) have criticized the above concept, but

JUNE
1973]
nevertheless both Clements' developmental

analogy and the homeostaticanalogy of Allee
et al. have survived in some form. For some
ecologists these analogies seem to be merely
heuristicdevices,but in othercases (e.g., Odum,
1969) it is not altogether clear whether the
usage is onlymetaphorical.
Since the time of Darwin the dominant idea
about the controlof populations has been that
natural selection has determined how many
offspring
a species produces,and the numberis
always somewhatin excess of what can survive
to maturity,the excess dying in the struggle
for existence. Although Darwin noted cases
where the strugglewas against the environment
rather than against other organisms(1859, p.
62), his emphasis was upon intraspecificand
interspecificcompetition. In the 20th century
there has been much discussion of the extent
to which populations are controlled by competitionas againstotherfactors.
The debate seems to have begun with Kropotkin (1902), who argued that weather is
more important than competition for controlling populations. His observations were
largelyconfinedto Siberia and were not necessarilyrepresentativeof other climates. During
the followingyears the distinctioncame to be
made between density-dependentand densityindependent factors,but the debate over the
relative importance of the various factorsaffectingpopulations has not yet yielded a consensus among ecologists(Allee et al., 1949, p.
331; Andrewarthaand Birch, 1954, pp. 16-25;
Lack, 1966, pp. 281-312; McLaren, 1971).
While that debate is directlyrelevant to the
presentdiscussion,it is too involved to be summarized adequately here. Some notable examples, however,should not be omitted.
The papers of A. J. Nicholson have attracted
much discussion, and two early ones focus
specificallyupon the balance of populations
(Nicholson, 1933; Nicholson and Bailey, 1935).
Responding to Elton's statement,Nicholson
remarked,"It appears to be usual at presentto
deny the existence of the 'balance of nature,'
about which there has been so much vague
talk. Therefore, before consideringthe question of the balance of animal populations,
we mustexamine the argumentsforand against
the existence of such balance" (1933, p. 132).
There were two argumentswhich had been
345
used to reject the balance of nature: populations do not remain at a constant level; and
natural selection cannot maintain the balance.
However, neither of these claims seemed to
Nicholson to invalidate the existence of a
balance, and in support of the balance of
nature he cited several laboratory studies on
insectpopulations.
He also offeredin defenseof the balance an
analogy that seems as elusive as the balance of
nature itself:
The balance of animal populationsis similar
to thatof a balloonactedupon by the changing
of nightand day. Such a balloon
temperatures
risesand fallsin relationto the changein temperature,for this varies the volume of the
air.
balloon and the densityof the surrounding
in a stateof tending
The balloon is continually
towardsa position of stationarybalance, but
continuesto riseand fallbecausethe positionof
balanceis changingall the time(1933,
stationary
p. 133).
This analogy has drawn firefromother ecologists, such as H. G. Andrewarthaand L. C.
Birch, who reject the idea that density-dependent factorscontrolpopulation. They complained thatin Nicholson'sdiscussion,"It is not
easy to understandwhat preciselyis meant by
theword 'balance'" (1954,p. 20).
Other notable examples are concerned with
mechanismswithin species for regulatingtheir
numbers. Wynne-Edwards(1962) has argued
that there are behavioral mechanismsfor this,
but his interpretationshave not won widespread acceptance (Lack, 1966, pp. 311-312; G.
Williams, 1966, pp. 243-246). MacArthurand
Wilson (1967) have found that for island
species the regulatorymechanism is natural
selection. They concluded that species having
high rates of reproductionare more likely to
surviveduring the early colonizationof islands
when populations are low, but that selection
pressurewould later favor species with lower
rates of reproduction. Krebs, Gaines, Keller,
Meyers,and Tamarin (1973) have mostrecently
argued that natural selection might regulate
populations not only by changes in rates of
reproduction,but also by changes in migratory
habits.
What, then, is the status of the balance-ofnature concept today? For many ecologists it
may have been superseded by more precise

346
THE QUARTERLY
REVIEW OF BIOLOGY
ecosystem concepts of productivity, succession,

and energy flow (e.g., Margalef, 1963). If this is
true, it may mean that these ecologists only
think in terms of the balance of nature when
they need a poetic or propagandistic phrase for
a discussion of conservation or pollution. If
this is the case, such a usage could still influence their own thinking as well as that of the
general public.
A few ecologists have actually cogitated regarding the concept and have attempted to
give it a modern meaning. Carrington Williams
wrote a book entitled Patterns in the Balance
of Nature (1964), in which he described mathematically not merely the population fluctuations of one or two species, as had commonly
been done in the recent past, but rather "the
pattern in which all the species in an association are balanced" (p. v). To Williams, however, the balance of nature represented an approach to a problem more than it did a
hypothesis.
[VOLUME
48
steady state of balance might theoretically be

achieved. Any sudden change, such as the introduction of a new species, produces a state of
"imbalance" and the various forcesof population
controlwill set about returningto a new balance,
although this may be only a new variation of
the older pattern. Such cases are well known in
the accidental introductionof insects into a new
country, where they have rapidly increased in
numbers to become a major pest. Man then
makes effortsto restore the previous balance by
the introduction of enemies of the new pest
(p. 295).
Two other biologists, Lorus J. Milne and

Margery Milne, have recently written a book
entitled The Balance of Nature (1960), but it
provides no evidence for the current viability
of a balance-of-nature theory. The book is
written for the general public, and although it
provides some documentation for the cases
described, it is far short of being a scientific
treatise. The provision of rigorous evidence of
causal relations is beyond the book's scope, but
Each species establishes temporarily an uneasy even if it were not, no general theory emerges.
As in times gone by, the balance of nature is
balance of numbers among all the others. The
pattern of the relative abundance of all the spe- simply taken for granted, and a series of
cies in a mixed community (and all wild com- specific illustrations described. It is very much
munities are mixed) is thus a synthesisof all the in the spirit of Herodotus, whose story about
competition and co-operation, and all the diffi- the
relationship between the Nile crocodile and
culties and facilities, that have surrounded all
plover is actually mentioned (p. 66).
the species of the communityin the recent past;
Not all recent ecologists have shared the
and every minute the position of any one spebelief of Williams and the Milnes that the
cies is changingin relation to the others.
balance of nature can remain any longer a
The pattern of relative abundance is thus an
expression of the momentarybalance which has
viable concept. Erhlich and Birch, reacting to
been set up among all the species of the associa- a passing reference to the balance of nature
tion, and it is important to find out whether,as
by Slobodkin, responded as follows (1967, pp.
time passes, the fundamentalpattern changes, or
97-98).
if the species move in their relative abundance
within a more or less stable pattern. This is the
The existence of a supposed balance of nature
approach to the problem of the "balance of
is usually argued somewhat as follows. Species X
nature" from the point of view of quantitative
has been in existence for thousands or perhaps
synecology(p. 2).
millions of generations,and yet its numbers have
never increased to infinityor decreased to zero.
Although Williams' statistical descriptions may
The same is true of the millions of other species
be very useful, they led to a series of limited
still extant. During the next 100 years,the numconclusions rather than to a broad, well-defined
bers of all these species will fluctuate; yet none
will increase indefinitely,and only a few will
theory of the balance of nature. One of these
become extinct. Furthermore,most species have
conclusions which does fall within the bounds
at least some populations living in areas where
of traditional balance-of-nature thought turns
they are well able to cope with the climate, yet
out to be his most speculative.
even these populations never increase indefinitely.
Such "observations" are made the basis for the
If the population remains for a long period
without interferencefrom outside, and if the
statementthat population size is "controlled" or
physical environment also remains constant, a
"regulated," and that drastic changes in size are
JUNE
1973]
the results of upsetting the "balance of nature."

Sometimes this is put in other ways, such as "on
the average, the species just replaces its numbers
in successivegenerations"or "on the average, the
numbersof individuals over a long period of time
are constant." An extreme version can be found
in Slobodkin (1961, p. 46). "Despite this enormous variation in reproductive patterns, each
female adult animal alive now -in everyspecies,
in almost every location -will
be replaced by
preciselyone female alive a generationfromnow.
If this were not the case, the size of animal populations would be changing permanently and
strikinglyat a much greater rate than any existing evidence indicates."
To the critique which followed these words,
Slobodkin, Smith, and Hairston (1967) replied
that in their opinion it was unlikely that any
contemporary biologists actually held the concept which Ehrlich and Birch refuted. This
controversy has seemed significant enough for
the papers to have been reprinted in a collection of articles on population and community
ecology (Hazen, 1970).
In spite of all these negative comments, it
seems likely that to sound a death knell for
the balance-of-nature concept would at present
be precipitous. A better approach might be
for ecologists to define the limits of validity of
the balance-of-nature concept within the context of modern ecology. The balance of nature
347
is analogous to the stabilityof species. There

are forces which tend to preserve the demographic stability of species just as there are
forces which tend to preserve their genetic
stability. In the short run, both biotic communities and species appear to be stable, but
in the long run there are dramatic permanent
changes in both. Mutations and natural selection gradually change species. Extinctions,
species evolution, changes in species composition, and environmental alterations change
natural communities. Some species, and some
communities,are much more stable than others.
When one studies evolution and genetics it is
important to recognize both the forces of
stabilityand of change and to formulatescientifichypotheses,theories, and laws accordingly. The same is true in ecology. The forces
of stability and change must be carefully
studied and then applied to the subject of the
balance of nature. Then criticalscientificgeneralizations can be formulatedwhich do not
have theirfoundationin hidden traditionsthat
extend all the way back to Greek mythology.
ACKNOWLEDGMENTS
I wish to express my thanks to the many colleagues with whom I have discussed this subject.
I have benefitted in numerous ways from their
comments.
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CHANGING CONCEPTS OF THE BALANCE OF NATURE

The balance of nature has been a background assumption in natural historysince

The leopard with harmlesskid laid down,

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THE BALANCE OF NATURE

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CIENTISTS attemptto explainnature

with generalizationswhich are called

medieval and earlymodernthought. It became

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THE BALANCE OF NATURE

lated, but other elements which should have

they ate. To illustrate his point he assembled

cept in Arabia, where they are all congregated

Although the report of superfetationin hares

These reports left the strong impressionthat

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THE BALANCE OF NATURE

the less precise of the two. This mythis the

god Epimetheusundertook. He bestowedtraits

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Most trees,if theybear too much fruit,wither

In this passage Aristotleemphasized food and

that mutualism existed between the bivalve,

sage he accepted mutualismand implied that it

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THE BALANCE OF NATURE

tween these categories,and theywere not fully

of turning . . . but also to prevent these fishes

prevented him from exaggerating either the

The inhabitantsof Thessaly,of Illyria,and of

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has never been much chance that a theory

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THE BALANCE OF NATURE

the factorof differentiallongevityto the ones

offerhis own suggestionthat the populations

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temperated,and so continuallymanaged and

come accepted dogma. Ray remarked (1693,

THE BALANCE OF NATURE

During the 18thcenturythe religiouscontext

he placed greater emphasis upon ecological

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THE BALANCE OF NATURE

Derham. Both reportedobservationsthat could

only one or two individualsof each species,the

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Darwinian questions were simply beyond the

which have been discussed above, along with

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THE BALANCE OF NATURE

Linnaeus' theoretical constructsof 1749 and

nature to the status of an explicit scientific

Unfortunately,it was not a science that

that he eventually published a three-volume

Tout se conserve dans l'ordre 6tabli; les

THE BALANCE OF NATURE

the importanceof competitionin nature might

peds, as rabbitsand fieldmice,

Wallace wrote these remarks in his "species

Another British naturalist who had read