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USE OF MATHEMATICS IN
POPULATION ECOLOGY1
By K. E. F. WATT
Statistical Research Service, Canada Department of Agriculture,
Research Branch, Ottawa

Mathematical models constitute language, not science. Hence the most

meaningful classification of models is in terms of the origin of the state
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ments they describe symbolically rather than their structure. Such a classi
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fication is necessary at the outset of this review to prevent semantic

obfuscation later.
If a polynomial or other simple formula is fitted to data by eye or
regression methods, the result is an equation which is useful for inter
polation but little else. Such empirical formulas are not, strictly speaking,
models.
Models of Kostitzin (62), Lotka (76, 77), Nicholson & Bailey (91),
Thompson (117, 118, 119), and Volterra (131), to mention only a few
representatives of this type, are developed deductively from a priori prem
ises. That is, the model-builder makes certain reasonable assumptions. Then
calculus, algebra, or other mathematical techniques are used to restate the
assumptions in various forms and, thereby, to deduce a number of con
clusions that were not immediately obvious from the assumptions. In con
tradistinction to this approach, which does not make use of precise and ac
curate data, a posteriori deductive models are based on assumptions checked
by measurements or counts.
Finally, complex models may contain a mixture of some terms obtained
by regression methods and other elements of a posteriori deductive origin.
However, it is unusual to encounter models containing both a posteriori
and a priori elements, because, if a thinker is going to reason a priori, he
will reason that way in toto.

THE ROLE OF MODELS IN THE EVOLUTION OF POPULATION E COLOGY

Until very recently, the relation between mathematical models, on the
one hand, and observation and experiment, on the other, has been different
in population ecology from virtually any other science. A detailed study
of the history of physics, for example, reveals that two principle differ
ences exist between the historical roles of theory in physics and popula
tion dynamics:
(a) In physics, theoretical developments have typically been stimulated
by the desire to make sense out of data which were already collected (65).
Newton's gravitational formulae replaced Kepler's empirical formulae which
were based on the observations of Copernicus, Galileo, and so forth. Max
well's equations were intended to explain data of Faraday. Einstein's
1 The survey of the literature pertaining to this review was concluded in February.
1961.
243
 244 WATT
revolutionary (39) paper was clearly stimulated by data from two kinds
of experiments. In population dynamics, on the other hand, all the classical
theories are a priori, not a posteriori, deductive models. The distinction is
subtle but very important. The essence of the modern (Galilean, versus
Aristotelean) scientific tradition is that in the last analysis it is a posteriori.
The theories of modern science are elaborated by deduction, but the basic
assumptions are obtained by observation, not abstract reasoning. Prospec
tive critics of my opinion that classical population models are a priori will
need to arm themselves with actual quantitative data from the early theo
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retical writings of Bailey (4 to 7), Lotka (76, 77), Volterra (131),

Thompson (117, 118, 119), Nicholson & Bailey (91), and Kostitzin (62).
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Such data will not be found.

(b) The historical development of physics has been characterized by a
high degree of integration between theory and experiment. In quantum
mechanics, for example, the closeness of this integration in the last few
decades has been amazing (49). On the other hand, many writers have no
ticed the lack of such integration in population ecology (69, 100, 114, 116,
141). This is an extremely important point, since the single factor most
necessary for rapid evolution of a branch of science is this close integration
of theory and empirical work. It is clear that sophisticated theory by itself
is not adequate since population ecology has had such theories for about
four decades (76, 77, 91, 131). These theories have never had an impact on
the rate of scientific evolution like that of the physical theories which have
often been tested weeks and even hours after they were published (20). It is
also clear that elaborate experimental and observational studies, not di
rected and interpreted by theories, do not greatly advance a science. The
dead files of ecologists abound with vast sets of unused data which were
obtained from studies fundamentally populational in intent. This author
knows of several dozen such sets.
We must conclude that no matter how active a theoretical branch of
science becomes, it will ultimately run into a cul-de-sac if unrelated to the
here-now world of our sense impressions. On the other hand, a field pro
gram will most certainly also be of limited usefulness if unrelated to theory.
One aim of this review is to point out types of activity which can increase
the degree of integration in theory and practice.
I have not attempted to review more than a small fraction of the mathe
matical papers published in this field. It seemed more important to analyze
the logic and implications of various procedures. This end can best be ac
complished in the allotted space by selecting papers representative of cer
tain schools of thought and discussing them in some detail.

REVIEW OF POPULATION THEORY

Several rather distinct: traditions established themselves at the beginning
of theoretical work in population dynamics and, to a large extent, have re
mained separate ever since.
 MATHEMATICS IN POPULATION ECOLOGY 245
The central tradition in theoretical population ecology has concerned it
self with models of birth, death, competition, and attack (parasitism and
predation). This tradition began with Verhulst's (130) derivation of the
logistic equation, which was not rediscovered until 1920 by Pearl & Reed
(102). Seven of Lotka's 94 papers are on theoretical population dynamics
[most importantly (76, 77)], but his conceptual framework seems to have
come from demography (73) and chemical kinetics (75), Another pioneer
in this work was Thompson (117, 1 18, 119). Apparently it was Thompson's
early writing that stimulated Lotka's (76) paper. This pattern of a mathe
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matician being stimulated to do pioneer work on population dynamics by a

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biologist was repeated several times. Volterra (131) was interested in popu
lation problems by D' Ancona, and Bailey [ (4 to 7) Nicholson & Bailey
(91)] was asked by Nicholson to work in the area in the first half of 1927.
It is clear that the character of early work in theoretical population ecol
ogy is in large part explained by the fact that it was developed by mathe
maticians. A second very important point is that the main features of the
theories were being worked out before a great deal of data had been col
lected. Apart from the occasional pioneering study (27, 101), quantitative
experimental findings on density effects, competition and attack did not ap
pear in numbers until the mid-thirties (80, 96, 99). Large numbers of papers
on attack findings in particular, were not published until 1941 and later
(33, 34, 35, 124, 126, 128) .
The Latka-Volterra M odel.-I have selected one paper for discussion
here (131) because it is one of the most elaborately worked out representa
tives of early a priori work. Since Volterra's original paper is not readily
available, my discussion is based on a translation of a revised version
which was published in Chapman's book (28). (Volterra's original papers
are virtually identical to this version.)
The first noteworthy feature of Volterra's treatment is that he looks pri
marily to factors intrinsic to populations for an explanation of population
fluctuations, In fact, all early mathematical population theories minimized
the importance of climate in population regulation. As noted elsewhere
(83, 120, 143) , for some populations at least, factors such as climate may
never allow a population to become dense enough for density-dependent fac
tors to be brought into play. Unfortunately, there are only a few bodies of
population data available which cover enough years to support this position
adequately (95, 111, 140). However, Volterra's general approach is sound
for situations in which the environment is favorable for the populations in
question and is stable. Several points are made by discussing two of Vol
terra's equations. Let Nl and N2 be the numbers of individuals of a prey
and predator species, respectively. Then, for the rates of change of each
with time, we have (according to Volterra)
dNl
= (el - "(lN2) Nl 1.
dt
 246 WATT

2.

The coefficients <:1 and ez are the coefficients of increase (the instantaneous
birth rate less the death rate), and II and 12 are attack coefficients. Six
points about these two equations are noteworthy:
(a) Birth rates and attack rates are assumed to be density-independent.
Several recent papers summarize evidence (which was not, of course,
known to Volterra) to show that these rates are density-dependent (50,
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51, 52, 127, 139) . Volterra (131) and Lotka (76) both realized inde
pendently that these assumptions were not correct and intended that the
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constants be understood as functions of Nl and N2

(b) It is obvious that Volterra's (and Lotka's) thinking was analogous
to that previously used in the kinetic theory of gases; Volterra indicates
as much when he considers, for example, that the probable number of en
counters of two species depends on the product of their densities. The ex
tent to which Volterra thinks by analogy with physical systems is made
clearer by a later paper (132).
( c) All assumptions are a priori (no data are mentioned in connection
with these equations except for the remark that the proportion of selacians
increased amongst the Upper Adriatic fish populations during the first
World War).
(d) There is no concern with a comparative a posteriori approach; the
empirical results from different cases are not compared. In view of the
fact that one of the most striking achievements of biology (Darwin's mas
sive documentation of evolution under natural selection) was convincing
largely because of this comparative feature, it is strange that theoretical
population ecology had to wait so long for this type of approach.
(e ) The assumptions made are in fact very specific. For example, the
constant -<:2 indicates that the predator species' instantaneous rate of in
crease decreases linearly with Nz Anyone who has tried to husband preda
tors and prey together in a small, uniform, enclosed universe will have
doubts about the validity of this assumption.
(f) Because Volterra's model is based on an inadequate set of assump
tions, it is of no use in showing how best to control insect pests and, in fact,
is downright misleading in this regard. For example, because the model as
sumes a density-independent birth rate, it does not reveal that spraying can
produce a greater population than would have been without spraying (142).
Also, it does not show that pests can be particularly vulnerable to annihila
tion at very low densities because of the low probability of prospective
mates finding each other (141). Hence, it does not suggest the theoretical
basis for the important novel control methods that use sex-attractants or
manipulation of the genetic content of natural populations. Finally, it does
not show that limited functional and numerical response of parasites and
predators can allow a pest to "escape" their control (87).
In short, all the conclusions which Volterra draws out in a very thor-
 MATHEMATICS IN POPULATION ECOLOGY 247
ough monograph are perfectly valid deductions from the assumptions he
makes. However, the assumptions are not drawn from biological reality.
Also, the fact that events similar to those described by Volterra occur in
nature does not prove that the mechanism which gave rise to the events is
that described by Equations 1 and 2 above. Where experimental results
seem to confirm the findings of Volterra and Lotka, there is typically some
technical explanation. For example, "prey" or "hosts" are not annihilated
because yeast sediments out and escapes ingestion by Paramoecium (44);
prey escape annihilation if they have microniches in which to hide (43);
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the choice of initial parasite and host population densities "forces" cycles
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(34).
Nicholson-Bailey Model.-Similar remarks apply to the Nicholson &
Bailey paper (91). The specific conclusions follow correctly from the sp e
cific assumptions made. However, the conclusions do not relate to
biological reality unless the assumptions relate to biological reality.
The difficulty is not so much that the assumptions made are incorrect, as
that an inadequate number of assumptions is made. The source of the diffi
culty with a priori models is generally deficient imagination rather than
deficient logical power (138). For example, it could not have been ex
pected to occur to anyone, including Nicholson & Bailey, that the efficiency
of parasites per female decreases as the parasite population density increases
(23, 24, 25, 35, 125, 126, 139). As additional empirical work is done, newly
discovered facts will make a priori models seem even further removed from
biological reality (52, 81, 82).
Other related a priori models.-The Lotka-Volterra tradition has not
died, but is flourishing to a greater extent than ever before. This new lease
on life is attributable to the advent of stochastic equations. Complex phe
nomena do not lend themselves to description by stochastic equations be
cause the resultant models are mathematically intractable (85, 90). Hence,
the simple Lotka-Volterra equations are suitable subjects for conversion to
stochastic form, whereas more complex, but biologically realistic, differen
tial equations are not used by stochastic-model biometricians. The result
has been a spate of recent papers on stochastic counterparts of the Lotka
Volterra equations (9,11,12,29,30,41,58,59,70,71,72).
Various other papers have appeared which are related to the Lotka
Volterra tradition. Lotka (78) wrote an interesting paper on theory of
predator-prey encounters. Volterra (133) examined different cases to de
termine the effect of various values for different constants of birth rate and
death rate on population growth and extinction. Hutchinson (54) investi
gated the effect of introducing quadratic terms into Lotka-Volterra com
petition functions. Bodenheimer & Schiffer (19) produced a new model
also based on a priori assumptions. Wangersky & Cunningham (134, 135)
considered the effect of time lags on classical equations of population
growth. Odum & Allee (94) discussed the implications of dome-shaped
fecundity rate versus density curves.
Varley (129) has recently presented an interesting discussion of the
 248 WATT
Nicholson & Bailey (91) and Thompson (119) models. As Varley suggests,
there is truth in both these models. This once again makes the point that the
problem in a priori model building is in not thinking of all the necessary
assumptions.
Mathematical epidemio l ogy.-Mathematical epidemiology is another
branch of population theory with a long history. This field will presumably
be increasingly important to entomologists as pathogens become more widely
used for insect pest control. Farr (40) and Hamer (47) did important early
theoretical work which led to awareness of epidemic waves and the factors
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which caused them. Ross (108) developed and solved three difference equa
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tions to express early theoretical notions about epidemics; these equations

were based in part on his experiences with malaria and anophelines. Fur
ther contributions to this theory were made by Lotka (74); Brownlee (21)
fitted Farr's (40) formula for the density-dependence of mortality and an
other equation to human mortality data. The most striking feature of all
these early papers in theoretical epidemiology is that they are affected by
actual data to a much greater extent than early papers on theories of ani
mal population dynamics, presumably because good data on human disease
were available. These early theoretical papers were followed by a number
of contributions to mathematical epidemiology in the form of deterministic
models; then another major step forward was made with the introduction
of stochastic model procedures (2, 9, 10). Bailey (3) gives a comprehen
sive review of literature in mathematical epidemiology, and a recent book
by Muench (88) discusses mathematical theory of epidemics from a rather
different point of view.
Optimum yield theory.-Another facet of theoretical population ecology,
yield theory, is becoming of some interest to entomologists because insects
are being used for research in this area (22, 137). Theoretical work on
the optimum yield began with the early papers of Baranov (8) and Hjort,
Jahn and Ottestad (48). Recent complete reviews of the subject are pre
sented by Beverton & Holt (18), Carlander (26), Ricker (106, 107),
Schaefer (109, 110), and Watt (138, 140). A completely different ap
proach, linear programming, has been applied to forestry problems by Kish
ine (60, 61), The subj ect of optimum yield theory is reviewed in very com
prehensive fashion in a recent book (68). Landahl (66, 67) and Watt (137)
have shown how fairly complex models can be used to study manipulated
populations of flour beetles in the laboratory, These models are systems
analytical in intent in that they attempt to account in detail for the simul
taneous action of a great many factors by using a posteriori models.
Community ecology.-Plant ecology gave rise to an interest in the
organization of populations and species into communities (1, 45). There
is no need to review this area here, because the interested reader can be
keyed to all the relevant literature by using the very comprehensive bibliog
raphies in Hairston's (46) and MacArthur's (79) papers. Another very ac
tive area in which mathematics is being applied is community energetics.
 MATHEMATICS IN POPULATION ECOLOGY 249
The papers by Patten (98), Slobodkin (115), and Odum (93) provide an
excellent introduction to the literature in the various aspects of this field.
Distribution and dispersal.-There have been a great many mathematical
papers on the distribution and dispersal of animals. Papers that are par
ticularly useful in giving insight into the biological meaning of various
distributions are by Pielou (103), Waters (136), and Cole (31). Beall (14)
discusses the relation between various types of distributions, and some in
sight can be obtained from his table which shows the fit of 11 distributions
to a body of corn borer data. Torii (123) gives a great many examples
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of distributions fitted to field entomological data. Cox & Smith (32) show
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how distribution can be analyzed in containers of flour beetles.

Skellam (112, 113) wrote two important mathematical papers on popula
tion dispersal and distribution, and the theory in the first of these has been
applied by Dempster (36). Amongst the most interesting papers on mathe
matical analysis of dispersal are those by Barton & David (13), Jones
(57), Holt (53), and Dobzhansky & Wright (38).
To date there .have been insufficient analyses treating dispersal, distribu
tion, and population dynamics phenomena in the same equation. However,
Beverton & Holt (18) have shown how it is possible to develop such equa
tions. A very recent book by Munroe (89) makes the first serious attempt
to show how mathematics can be used to provide a conceptual link between
popUlation ecology and zoogeography. He points out that the transition
matrices of Markov chains provide a succinct means of expressing not only
the temporal change in numbers at a point but also the temporal change in
numbers at a set of points.

CURRENT TRENDS AND PROBLEMS IN MODEL-BUILDING

Within the last few years, it has become apparent that several new
trends were showing up in theoretical ecology; these trends indicate likely
direction of work in the future.
The single factor having the greatest impact on the character of modern
theoretical ecology is sampling. Ecologists now realize that the only way
to gain insight into natural popUlations of animals is to collect for many
years precise, accurate data on such populations and the factors that affect
them (86)., Several very large-scale projects are now operating. By far the
most ambitious undertaking by entomologists is that on spruce budworm
population dynamics at Green River, New Brunswick, which has already
produced two papers on models (81, 82) and one monograph (87), in which
a mathematical model for population dynamics of the spruce budworm is
developed and tested.
As the data from more large-scale projects become available, bio
metricians will be faced with the problem of building models very much
more complex than almost anything which has appeared up to now. The
need to build such models will in turn raise a number of quite crucial ques
tions, which are discussed in a recent monograph (144).
 250 WATT

The monograph explains why the effort required to build large and
complex mathematical models of ecological phenomena is justified by the
results of the work, answers criticisms of the model approach, and explains
in detail some of the available techniques for building and using large-scale
mathematical models. A new criticism of models (64) has appeared since
my monograph was written. Labeyrie demonstrates that the "constants" in
models can not really be constant, because, for example, slight differences
in the dispersal of hosts are discovered by each female parasite. This is very
true, but the same instantaneous variation in behavior could be noted for
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particles undergoing radioactive decay or a host of other phenomena for

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which there are satisfactory mathematical descriptions. The important prob

lem is for us to be able to describe average behavior, and we can do this
successfully even when ignorant of between-individual differences (e.g,
81,82).
Many of the important new trends in model-building are illustrated in
an excellent paper by Fujita (42) in which he develops a mathematical
model for the effect of density on fecundity. The first noteworthy feature
.
of this paper is that Fujita has a thorough awareness of experimental find-
ings on the phenomenon for which he develops his modeL He makes imagi
native application of the deductive process to develop his model; his as
sumptions are sound, and he has a reasonably adequate set of assumptions.
A very important feature of this paper is that Fujita recognizes that three
apparently different types of situations are merely different manifestations
of one broad underlying law. This is model-building at its best: a com
parative approach is used to determine what fundamental features are com
mon to a great variety of cases. This procedure leads to fresh insight, since
it can clearly demonstrate why the different cases are different. Finally,
Fujita presents graphs showing how his model behaves for different values
of the constants.
Lanczos (65) distinguishes between comprehensible and incomprehensi
ble models. For example, it is completely impossible to understand some of
the basic equations of quantum mechanics, since we can not conjure up any
mental picture which will make the law expressed by the model seem rea
sonable to us. Yet the laws must be accepted as the best currently available
on the basis of experimental evidence. On the other hand, other mathemati
cal models of physics explain observed phenomena in terms of laws which
seem quite reasonable. Clearly, mathematical models for ecological phenom
ena will be of most use if they are of the latter type. Suppose a particular
model predicts correctly that parasite A is more effective than parasite
B but does not allow us to understand why A is more effective. Such a
model is less useful than one which explains why A is more effective than
B in addition to making the correct prediction, since the latter model tells
us how to look for a parasite that wiII be even better than A.
Another important point about model-building which ecologists are
coming to recognize is that models should concern themselves with the an-
 MATHEMATICS IN POPULATION ECOLOGY 251
alysis of systems and the interactions therein, not with isolated phenomena
lifted out of context. Typically, the effect of any variable in a complex
ecological system is influenced by the simultaneous variate-values of other
variables in the system. For example, whether the defoliation of forest
stands has any effect on starvation by the pests which caused the defoliation
will depend on the population density and genetic composition of the pests
in the year of starvation and following years (87). Also, the effect of any
parasite or predator species depends not only on its density but also on the
density of the vulnerable species (139) and environmental factors (82). A
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recent paper shows that explanation of laboratory population phenomena

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may be impossible unless the structure of interaction effects is understood

as completely as that of single factors (97).
I have discussed in detail elsewhere the characteristics a model should
have in order to be realistic (141, 144). In short, a model of a complex
ecological phenomenon should have a large number of independent vari
ables and should be structured in such a way that a factor (e.g., Tricho
gramma density) is entered correctly in time with respect to the phenologi
cal development of the pest (e.g., only eggs are vulnerable, to Tricho
gramma). The model should describe the complex interactions which exist
in nature and illustrate how some factors can operate in each of several
different causal' pathways. For example, increasing temperature can in
crease mortality of a pest by increasing the rate of searching of its para
sites and predators; on the other hand, it can decrease mortality of the pest
by speeding up its development and hence reducing the time interval in
which its enemies can find it. The model may have to be formulated to in
dicate how restrictions must be placed on control (for technical or eco
nomic reasons). The model must indicate that competition exists at all
levels in the biological world and that biological entities have built-in limita
tions which profoundly affect the way in which they operate. These last
two features make the structure of the terms in ecological models rather
different from the structure of terms in many models of physical phe
nomena.
Another important trend which has appeared recently is that work with
mathematical models is exposing many deficiencies in present ecological
knowledge; subsequent empirical work to fill these gaps has led to more
refined theory. For example, not nearly enough is known about the effect
of fluctuating humidities and temperatures on insect survival to adequately
account for observed correlations (87). Various mechanisms of interaction
of stand factors and climate are not thoroughly understood. The back
ground of information in behavior and physiology is often inadequate to
allow us to see how to build useful models.
The recent models on parasite and predator attack (51, 139), the effect
of density on fecundity (42,141), and the influence of relative density of a
prey species on its contribution to a predator's food (121) all suggest many
avenUes of research on behavior mechanisms which require more attention.
 252 WATT
As a result, there is presently a surge of interest in performing quantitative
studies of behavior (52, 84, 104, 121). Not too much imagination is required
to foresee linking of mathematical work on behavior (78) and modern ex
perimental analyses of behavior (37, 50, 52, 63).
Another new tendency in ecological model-building might be referred to
as the building-block approach. Instead of attempting to build a large model
all at once, the phenomenon is split up into a sequence of consecutively
operating components and, within each of these, a group of concurrently
operating components. Sub-submodels are developed and fitted together to
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form submodels which can then be fitted into a model (87, 144).
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At the time of writing this review, it appears that stochastic, as opposed

to deterministic mathematical model-building will have only a limited im
pact on the mainstream of ecological thinking. Apparently, because of the
great difficulty of working with complex stochastic models, this type of
equation will be used largely to gain insight into relatively simple phe
nomena (144). It is noteworthy that Neyman & Scott (90), in one instance,
had to simplify their assumptions in order to build a stochastic model they
could manipUlate.
Many of the earlier ecological models were designed for steady-state,
or equilibrium, conditions in which large-scale changes of the physicochemi
cal environment did not occur (18, 76, 91, 131). However, as more and
more data become available which show that very large-order fluctuations
are influenced by weather, steady-state models may decline in popularity
(87, 95, 111, 140).
There seems to be a definite tendency away from use of multiple re
gression analysis and polynomials and toward models based on reasonably
realistic differential equations. This presumably reflects the fact that ecolo
gists have an increasing awareness of the following undesirable features
of the use of multiple regression analysis (144): The meaning to be at
tributed to a high or low correlation coefficient may be in question. The
fitted equation may yield no insight into the mechanics of a system. The
phenomenon may involve non-Iinearities of a type that can not be handled
by regression equations (e.g., several asymptotic effects). Finally, multiple
regression analysis may have an insidious effect on the development of
theoretical work in a subject matter field if men try to force information
into an artificial conceptual mold. The main advantage of multiple regres
sion analysis is that its procedures are mathematically straightforward
rather than that the fitted equations bear a close correspondence to events
in biological reality.
Another new tendency in papers about models concerns comparisons of
various models. In the older literature, with few exceptions, a new model
was presented without any attempt to show that it gave a better fit to data
than previous models. One notable exception was the paper by Janisch (55)
in which he compared the fit of six formulae to temperature-development
time data. However, nowadays so many models are already in the litera-
 MATHEMATICS IN POPULATION ECOLOGY 253
ture that it behooves authors of new models to demonstrate that their
models really represent an advance rather than a further cluttering of an
already well-cluttered literature. Holling (51), Miller (82), and Watt (139,
141) have compared the agreement between values observed, and cal
culated, from two or more models.
One very new trend is the attempt to explain phenomena at a macro
scopic level in terms of processes at a microscopic level (56, 145); in this
case, responses of insects and other organisms to temperature are analyzed
in terms of molecular kinetics. An important consequence of this activity
is that a more solid link is being established between theoretical work in
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physics and chemical kinetics, on the one hand, and physiology, behavior,
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and ecology, on the other. All fields should benefit from the resulting cross
fertilizations of ideas. Also, this type of theoretical work helps clarify the
relationship between the effect of a factor and the mechanisms which de
termine the magnitude of the effect.
Within the next decade, the need for work on models that link mathe
matical population dynamics and population genetics will be conclusively
demonstrated by new types of data (146).
There wiII be an enormous increase in the use made of computers in
mathematical population ecology in the next few years. One of the most im
portant reasons for this activity is that population theoreticians are becom
ing increasingly concerned with fitting quite complex equations to data. It
requires a prodigious amount of effort even to fit the relatively simple
fecundity model of Watt (141) unless a computer is used to apply the
method of differential correction (92). As more work of this type is done,
we can expect to see experimental work on computers to determine which
of the several available fitting routines (105) is most efficient for a given
type of equation. Another reason for the increased interest in computers
is that many ecologists have become interested in simulation (93). That is,
we construct a mathematical model of a population, preferably mimicking
the simultaneous effect of weather, density-dependent factors, and treatment
by man (122, 142, 147). Then the computer calculates how the population
will behave over a sequence of generations with no treatment and with dif
ferent types of treatment. Such research will be very revealing; for example,
one recent study showed that spraying pests could produce, under certain
conditions, more pests at a later time than there would have been if the
pests had not been sprayed (142).
A major new development is the application to ecology of branches of
mathematics developed recently to cope with the complexities of managing
complex manufacturing, transportation, and military, scientific, and engi
neering systems. One of these branches of mathematics is dynamic pro
gramming, which shows how to select the best sequence of decisions to be
made over a period of time (15, 17). Clearly, since insect pest control has
almost never turned out to be a "one-shot" campaign, mathematical meth
ods designed for multistage decision processes must be relevant for the
 254 WATT
economic entomologist. More recently, Bellman (16) has published a book
on adaptive process-control theory. This material should be of concern to .
entomologists, since insect pest control is an adaptive process control; that
is, we attempt to control a pest by using procedure A, then analyze what
went wrong, and use this information to devise a superior process for con
trol, B. This process of improving control procedure on the basis of in
formation about defects in previous control procedures continues until
control is perfect. Ideally, a computer should be integrated into the cycle;
test control procedure, and the defects in control procedure should be an
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alyzed, refined, and new procedures tested. It should be noted that Bell
Annu. Rev. Entomol. 1962.7:243-260. Downloaded from www.annualreviews.org

man's point of view with respect to the role of mathematical models is quite
different from that of most biologists. Instead of assuming a description of
a problem structure and finding out the best solutions to the problem in this
particular structure, Bellman uses mathematics to determine the best topo
logical structure for the problem; that is, mathematics is used to determine
an optimum policy, not the best parameter values for a given policy. It
follows from Bellman's approach that the best use of mathematics may
often be to show what not to do rather than what to do. In essence, mathe
matical programming puts bounds on the range of policies available to us
by eliminating those that are infeasible for some reason.
Since mathema ti cal programming as developed by Bellman and others
is designed to cope with problems of managing large-scale systems, it al
lows us to consider problems in a natural and sensible manner. For ex
ample, we can consider the whole economic problem of a. forest all at once,
not just the smaller problem of pest control lifted out of context. The rea
son for considering the whole problem at once is that the best control policy
from the standpoint of minimizing pest survival may not be the optimum
policy from the standpoint of maximizing profit from a given tract of tim
ber.
 MATHEMATICS IN POPULATION ECOLOGY
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