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Ann. Rev. EntomoL 1983.28:319-35

Copyright © 1983 by Annual Reviews Inc. All rights reserved

DISPERSAL AND MOVEMENT

OF INSECT PESTS

R. E Stinner
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Department of Entomology, North Carolina State University, Raleigh,

North Carolina 27650

c S. Barfield and J. L. Stimac

Department of Entomology and Nematology, University of Florida,
Gainesville, Florida 32611

L. Dohse

Departement de Biologie, Universite Laval, Quebec, Canada GIK 7P4

Perspectives and Overview

Until recently, the study of insect dispersal and movement received little
attention. Even 50 years ago, many ecologists considered studies of move­
ment to be trivial or even worthless exercises (44). Although this view is no
longer held, as late as 1972, Headley (61) wrote that "agricultural pest
control is still handled as though pests were immobile.... "

Since the publication of Southwood's (134) review and Johnson's (68)

monographic work, the study of insect movement has received increasing
attention. Numerous symposia and reviews on various aspects of insect
movement have been published [e.g. evolutionary considerations (6, 35, 37,
38, 48, 59, 60, 112, 134, 135); pest management implications (53, 108, 113,
151)]. Our purpose in this review is not to present an exhaustive list of
references; rather, we wish to compare views and approaches based on our
own, admittedly biased, selection of key references. With few exceptions, we
have limited our review to those works published since Johnson (68).
Our presentation is separated, for organizational clarity only, into sec­
tions dealing with evolutionary studies, experimental considerations, and
modeling efforts. These three areas of interest are o bv iously inseparable
biologically. We shall not attempt here to deal with the numerous papers

319
0066-4170/83/0101-0319$02.00
 320 STINNER, BARFIELD, STIMAC & DOHSE

that discuss the proximal physiological and behavioral aspects of pest move­
ment, since to do so would require an entire book (e.g. 68). The reader is,
however, referred to some of the key literature for specific orders in Table
1.
In this review we do not attempt to distinguish dispersal or migration
from other movements; rather, we consider all insect movements, including
those which could be termed trivial [sensu Johnson (68)], since the behav­
iors involved are not necessarily disjoint (58).
In any real world context, we cannot examine movement as a process
separate from other life processes because the practical and theoretical
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importance of movement is analogous to asking "what is the significance

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of spatially distributed dynamics for the persistence of interacting popula­

tions?" (28). Since we are dealing with problems involving both "escape"
and "colonization" (35), movement must be considered in terms of the
alternatives available to organisms for escape (e.g. diapause or estivation)
and for colonization (e.g. evolution of a wider host range or of more efficient
utilization of present hosts).
Our title implies that there is some reason to examine movement in
"pests," as opposed to "nonpests" as if there were, in fact, differences in
movement between those species that are generally regarded as pests and
those that are not. On an absolute basis, one can safely assume that there
is no difference. However, on a relative basis, strong arguments have been
made (134) that as a group, pests are more apt to be r-strategists. Their
life-history strategies tend to involve greater colonization rates, implying a
greater "importance" to movement. Certainly from the perspective of man­
aging these species in the agroecosystem, a knowledge of their movement
strategies and mechanisms would seem critical (11, 12, 75, 138). Another
obvious difference between those organisms whose major habitat is man­
dominated and those whose habitat is less disturbed is the ability of the
latter to coevolve with their hosts-a survival option that is short-circuited
Table 1 Selected literature on the impact of movement and dispersal on
population dynamics of pest species

Order Reference numbers

Lepidoptera 1,3,7,9,14,16,20,25,34,52,54,62,80,102,113,
133,137,139,151, 152
Coleoptera 23,26,31,86,94,107,120
Diptera 15,18,30,71,101,105,121,125,126,128,156
Hemiptera 36,43,82,98,99
Homoptera 13,42,78,90,93,96,122,155
Orthoptera 5,29,49,53,100
 PEST MOVEMENT 321

in agricultural systems via breeding programs for characteristics that may

or may not involve changes of importance to the fitness of the insects.
However, even with such differences, pests are subject to the same types of
selective laws that "govern" all life, and thus it behooves us to examine pest
movement from an evolutionary perspective.

Evolutionary Considerations
Evolutionary questions have been addressed at various hierarchical levels.
For example, at the global level, Levins (84) dismissed the previously held
dogma that the principal effect of gene flow was to "swamp" a local popula­
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tion and prevent adaptation to local conditions. He argued mathematically

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that gene flow among populations is part of the adaptive system and that
optimum values for flow are functions of the habitat structure. The end
result of such gene flow permits populations to respond through selection
to long-term, regional environmental changes while damping the response
to more local, temporary perturbations. He thus clarified the importance,
in a heterogeneous environment, of movement to species persistence.
In contrast, studies on the applicability of island biogeography, sensu
MacArthur & Wilson (88), to agricultural systems have provided mixed
results in expanding our knowledge of pest movement per se. Census studies
in soybean fields have demonstrated that the colonization process may
promote system instability because of rapid colonization by small her­
bivores, asynchrony in colonization by these herbivores and their natural
enemies, and high extinction rates (110, 129). Thus, extrapolation of island
biogeographic theory as an aid in the interpretation of pest movement will
require considerably more effort. As Rey & McCoy (118) point out, "propa­
gules" will have to be defined more carefully and the effects of habitat
changes (e.g. crop development) on colonization and extinction rates ascer­
tained.
Similar problems of interpretation exist in attempting to examine move­
ment effects on community structure. In a study of arthropods associated
with litter (149), it was shown that migration decreased species richness.
Although possible mechanisms for this effect were discussed, the available
data were not sufficient to elucidate which of the possible mechanisms
hypothesized was involved.
Ecologists examine patterns of occurrence (either observed or theoreti­
cal) and then seek to understand the courses of selection that could have
led to such patterns. For example,based on light trap catches of moths from
many years and locations, Taylor & Taylor (141) offered a conceptual
framework for the regulation of populations by density-related movement.
Fitness is seen as "maximizing the reproductive advantage of a balance
between migratory and congregatory behaviors." However,Hanski (57,58)
 322 STINNER, BARFIELD, STIMAC & DOHSE

interprets the patterns observed by Taylor & Taylor (and his own observa­
tions of two groups of dung-inhabiting beetles) quite differently. He demon­
strates that the rates of immigration and emigration in local habitat patches
(in his case, dung) can be governed by simple stochastic rules (as opposed
to density-related and/or "optimally" determined rates). Hanski (57) con­
cludes that the "difficulties in coping with a stochastic environment, and the
consequences of movements being manifested on many spatial and temporal
levels, favor evolution consisting of a few key behavioral parameters being
singled out and subsequently adjusted by selection" (see also 22, 161).
In search for explanations of the broad patterns and northward expansion
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of many pest species (followed by a characteristic winter "freeze-back"), the

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idea of a "pied piper" effect has been suggested (114). In temperate agricul­
ture, human activities have provided temporarily suitable habitats for many
species that are geographically far from suitable overwintering locations. A
general "random" dispersal of pest species into such regions during periods
of climatic favorability (and "artificial" provision of hosts) would account
for this northward movement in temperate North America. In an interest­
ing article, Walker (154) refutes this "pied piper" effect by arguing that this
model presents an evolutionary dilemma, since selection should operate
against northward moving individuals, unless there is a subsequent autumn,
southward movement. His arguments, however, rest on the unproven as­
sumptions that (a) selection has had enough time to operate since the
development of large-scale agriculture, and (b) this net northward move­
ment is behaviorally different from localized dispersal (or, if it is not, that
a mechanism for determining direction of flight is present in the individual).
Regardless of the correctness of these assumptions, Walker provides a
unique insight into comparative differences in movement behavior between
moths and butterflies and suggests experiments to explain these observed
patterns.
Southwood (135), pointing to the lack of any substantive classification of
ecological strategies, emphasizes the need for the same type of analyses. He
suggests a classification based on the choices available to an organism for
survival and reproduction [temporal ("now or later") and spatial ("here or
elsewhere") dichotomies]. In his most interesting analysis, he points out the
relationship that exists between the ratio of habitat suitability time to gener­
ation time and the necessity for escape (either diapause or movement),
including provision for the stochastic nature of habitat suitability in time
and space. This stochasticity has led some authors to propose (136) that r-,
not k-, selected individuals should be favored as stochasticity increases.
Other scientists disagree (32, 159), primarily based on arguments involving
the use of r and k (as a single vector continuum) to describe a multidimen­
sional space.
 PEST MOVEMENT 323

Regardless of the above "r-k" arguments, the relative importance and

interrelationship of diapause versus movement as escape mechanisms (135)
cannot be ignored. As Dingle [(37) pp. xiiv-xiv] points out, the two are
closely tied both behaviorally (e.g. the promotion of migration by diapause)
and physiologically (a common hormonal system).
Given the existence of varied levels and "kinds" of stochasticity ("patchi­
ness"), it would seem logical that this topic would be of interest in compara­
tive studies. In studies of two species of Tribolium, T. castaneum and T.
confusum, it was shown (160) that the former is a primary colonizer
whereas the latter is a secondary colonist. Since the habitat becomes unsuit­
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able more rapidly for T. castaneum, adults of this species start emigrating
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earlier and are more responsive to density than T. confusum. In addition,

whereas T. confusum adults often exhibit lower emigration in older beetles,
T. costaneum adults continue to emigrate as their age increases.
In another comparative study of movement, Jones (72) found that Plusia
californica larvae, when dislodged, travel rapidly with moderate headwav­
ing on a zigzag course ("radical" search), while Plutella maculipennis
larvae move slowly, headwave frequently, and turn often ("conservative"
search). Since Plutella feed only on cruciferous plants, which tend to occur
in small patches, and Plusia, conversely, is polyphagous, each species
appears to have "adopted" the search behavior most appropriate to its
hosts' distributions. Walker (154) compared five butterfly species with
five moth species for their migratory capabilities and found discernable dif­
ferences in their flight altitudes and times and their responses to physical
environments.
The converse of comparing two or more species in the same habitat is to
compare the same species in different habitats. In such a comparison (74),
Pieris rapae adults were found to exhibit quite different ovipositional search
patterns, again in a manner consistent with host patterns and "expected"
progeny survival.
This concept of "expected" progeny survival (Le. as a measure of
"fitness") has led to a wealth of research concerning "optimal foraging
strategies." While it is beyond the scope of this review to discuss this area
in depth, the reader is referred to several reviews (60, 112) and recent papers
(57, 111, 123). However, Maynard-Smith (92) warns that it is the role of
these approaches not to demonstrate that organisms in fact optimize their
foraging activities (see also 57), but rather to test the adequacy of particular
hypotheses to account for the evolution of'g�ven structures or behavioral
traits.
Interest in foraging strategies has produced a number of studies that deal
with the effect of monocultures as compared to polycultures on the move­
ment of insects (33). In numerous experiments, particularly with crucifer-
 324 STINNER, BARFIELD, STIMAC & DOHSE

ous crops, monocultures have been shown to house organisms with higher
rates of emigration [e.g.Phyllotreta cruciferae (140), Brevicoryne brassicae
(132), Aleyrodes brassicae (132), and "alate aphids" (27 )]. In most cases,
many of the herbivore species studied were oligophagous. In some cases,
inhibition of host recognition has been implicated (117). However, in all
cases small plots were used, leading one to question the generalization of
these results to agricultural systems (i.e. it is implicitly assumed that the
insect pests are mobile enough to reach the crop in regions where it occurs
over large acreages). It is interesting to note in Cromartie's
(27) work, that
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although two Phyllotreta spp. and alate aphid densities were positively
correlated with plot size, densities of Pieris rapae were negatively corre­
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lated. Australian P. rapae also have been shown to exhibit the same discrim­
ination against less acceptable host varieties whether planted together or not
(67).
The "monoculture-polyculture" controversy was intensified by observa­
tions such as those of Farrow (45) that ". . . forest clearance in Borneo, New
Guinea, and Queensland, where monoculture of sorghum has been intro­
duced, has resulted in serious outbreaks of Locusta and other Acridids."
This quotation is taken out of context, and in the article the author gives
an example of reduced locust problems with introduced, small farms. How­
ever, the implication that monoculture is a "cause" of locust outbreaks is
clear. Whether this is due to monoculture per se or to a simple increase in
available host materials has not been demonstrated. At least one matJIemat­
ical study (56) has suggested that "adaptations for achieving dispersal retain
great importance even in uniform and predictable environments." Current
literature thus leads us to conclude that any generalizations concerning
monoculture-polyculture comparisons can only be made in a probabilistic
sense, and a great deal more research is justified.
Up to this point we have discussed movement and movement behaviors
as if they were stable within a species. Certainly, if behavior is adaptive, then
within a system exhibiting spatial and temporal heterogeneity, one could
expect polymorphisms in movement to play an important role. Both Dingle
(38) and Johnson (69) provide detailed treatments of this subject. Although
Dingle (37) has provided insight into the question of polymorphism for
migration as compared to the use of diapause as an escape adaptation, the
potential effect of large, unpredictable weather perturbations on polymor­
phism in agricultural pest species has not been studied. With well-known,
migrant pests (see 2, 64, 103, 113), this would seem to be a particularly
rewarding area of research.

Experimental Considerations
Conceptual evolutionary arguments have derived from and spawned experi­
mentation on movement in pest insect species. Much of this experimenta-
 PEST MOVEMENT 325

tion has been conducted in controlled environments, and has focused on

responses during flight and reproductive activities as a function of various
factors (e.g. temperature, photoperiod, density of immature stages, adult
density, mating status, etc). Common to most of these studies has been the
need for instrumentation to measure flight activity and for interpretation of
the data acquired (see next section).
Insect movement has been measured with devices ranging from Malaise
traps (153) to airplanes equipped with nets (51) and/or sophisticated radar
equipment (40, 53, 119, 151). Indeed, radar has become an immensely
important tool in the tracking of pests that fiy en masse (53), and the use
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of both ground-stationed and airplane-mounted radar has been investigated

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(113, 151). Although tracking of single organisms can be done, differentia­

tion among species fiying together does not currently appear possible, most
likely owing to little experimental effort in this area (see 151).
Among the tools used by the typical entomologist, various types of light,
pheromone, and/or suction traps have predominated (e.g. 8, 21, 55, 97, 131,
142). The major problems that were perceived early in the history of these
devices still plague entomologists, i.e. the relationship(s) between trap catch
and either ambient or in-field densities of insects present. Even though
researchers still use these types of devices in an "early warning" capacity
for invading pest species, it is clear (11, 12) that trap to actual density
conversions have not been the subject of sufficient experimentation. Since
most crop protection researchers want to use these devices as indicators of
an approaching pest problem, lack of ability to make these conversions
remains a monumental problem.
Much of the laboratory experimentation on insect movement has focused
on distinguishing between the processes of flight and reproduction (both
mating and oviposition). Dingle (36) outlined various approaches that can
be used for this purpose. In studies of this type, a device is usually required
to measure flight (frequency, duration, direction) in relation to reproduction
in the life history of the insect in question. Many of these devices are
modifications of circular flight mills to which the insect is tethered in
various fashions (e.g. 127). Some have been automated and can provide
output amenable to direct data processing (4); others use chart records
attached to signal devices such as photocells, simple counters (e.g. 81),
videoscanners (150), ultrasonic detectors (87), or radar-Doppler type analy­
ses (17). All these are single-insect recorders. However, activity "cages"
have been developed to measure flight in groups of insects (17, 41). Acoustic
actographs (63), self-contained flight chambers (55), visual stimulus detec­
tors (106), and devices with retractable surfaces to stimulate flight (47) have
been developed. P. Wales and C. S. Barfield (unpublished) have developed
a photocell-triggered recorder to measure flight while simultaneously mea­
suring oviposition on a rotating, clock-calibrated resting surface. All these
 326 STINNER, BARFIELD, STIMAC & DOHSE

devices, to some extent, have been tailored to specific research questions and
target insect(s). The most common problems faced in using these tools have
been difficulty in (0) obtaining "representative flight" under laboratory
conditions and (b) quantifying and interpreting flight data.
Many of the insect pest species for which movement is thought to be an
important process in population dynamics and in the establishment of their
pest status are also polyphagous. The impact of variable nutritional requi­
sites, in combination with active or passive modes of transport, has made
these organisms extremely difficult to deal with experimentally (e.g. 11).
The body of information involving specific vectors of environmental (biotic
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and/or abiotic) stimuli associated with flight initiation, direction, duration,

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and termination is relatively sparse. There are at least four reasons for this.
First, most of these pest species are not boundary layer fliers and often fiy
at night; thus, they are difficult to sample or observe actually in flight (154).
Second, many pest species do not appear to be strong fliers and are thus
thought to be transported passively on weather systems. It is very difficult
to relate relevant weather phenomena to insect movement (see 89, 109),
because of the lack of adequate "in flight" sampling techniques or models
adequately constructed to relate weather to density of fiying insects. Third,
the polyphagous nature of these types of insects suggests that sources of
migrants may be spread over large geographical areas (i.e. many point
sources). The science of assigning "signatures" to source populations has
not been well developed, although much progress has been made in the use
of X-ray dispersive techniques (e.g. 148), allozymes, morphotypes, insecti­
cide resistance, and other natural and artificial markers in estimating popu­
lation source locations (for review see 113). Fourth, colonization and
survival of these species is often heavily dependent upon our determination
of the host plant system structure, both temporally and spatially. These four
problems are not eminent in all species; however, for the majority, their
impact is significant.
Information on movement in pest species appears fragmented and insuffi­
cient to test even conceptual models that exist for nonpest species (see 6,
61). Most of the studies deemed successful for the latter species have been
conducted on boundary layer, day fliers with limited host ranges; thus, the
first of the four problems stated above was avoided. As best we can tell,
researchers interested in pest species have not focused sufficiently on the
intricacies of movement to allow an understanding of this process. At least
some of us argue (11, 85, 114) that this must change if agriculture is to be
successful in forecasting pest outbreaks.
Thus, based on existing information, we can suggest four avenues of
research that are critically needed to understand the role of movement in
elucidating the dynamics of pest species.
 PEST MOVEMENT 327

1. Development of biologically meaningful conceptual models of move­

ment as the basis for structuring and evaluating data on movement in
pest species.
2. Derivation of reliable relationships between (or among) physical phe­
nomena and spatiotemporal appearances of target insects. This includes
knowledge consistent with optimal allocations of sampling tools (e.g.
various traps) to detect and evaluate these relationships.
3. Derivation of a reliable methodology for assigning probabilities to cer­
tain geographical areas as sources of migrants.
4. Identification of physiological and behavioral mechanisms conducive to
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initiation, orientation, maintenance, and termination of flight.

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These mechanisms would appear to involve complex processes set within

a framework of a highly variable environment. Such systems would thus
necessitate the development of complex hypotheses (Le. models).
Movement Modeling
Mathematical models of movement for insect pest species can be descrip­
tive,explanatory,or predictive. The intended use of the model determines
the type of model needed as well as the spatial and temporal resolution
required. Freeman (46) distinguishes between types of models by whether
the mathematical model simply describes what has happened (descriptive),
explains what has happened in terms of parameters of biological significance
(explanatory),or provides a forecast of what may be expected in the future
(predictive). The majority of insect pest models of movement are descrip­
tive, even though evolutionary and management questions about the role
of movement may require complex explanatory and predictive models.
Regardless, it would seem obvious that the variation in behaviors and
spatiotemporal scales among species, as well as different interests of re­
searchers,should lead to a wide variety of models and modeling approaches.
Descriptive models of movement consist of a family of regression equa­
tions relating frequency of catch or insect density to time or distance from
a point source of dispersing organisms (46, 65, 66, 143, 144, 146). Move­
ments of some insect pest species have been described as a function of
weather variables (19,95,115,116,122) or distance of separation between
individuals (147). Without modification, descriptive (regression) models of
insect movement are unlikely to apply to a wide range of situations because
the parameters that a posteriori fit the model to data are influenced by both
abiotic and biotic conditions encountered during the movement process.
Also, such models are not applicable to situations where insects do not
originate from a point source. As pointed out earlier, this is the case for
many insect pests because they are polyphagous and dispersed over large
geographical areas.
 328 STINNER, BARFIELD, STIMAC & DOHSE

Like descriptive models, explanatory models describe what has hap­

pened; however, in the latter, emphasis is placed on explaining why the
resulting movement occurred. Parameters of explanatory models have bio­
logical significance in terms of our knowledge of the species' ecology. Many
attempts at developing explanatory models have been made within a frame­
work of continuous time and homogeneous environment, and using diffu­
sion equations (e.g. 83, 104). While these models have provided some
interesting mathematics, we feel that such models have limited applicability
because of the complexity in using nonisotropic diffusion equations (39).
Although experimental systems have been set up to validate some simple
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diffusion models (77), they utilized a highly artificial design to reduce the
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natural complexity.
Another approach has been to divide an entire region into cells (124) and
to calculate the probability of moving into adjacent cells. Such a modeling
method was used to simulate movement of a cereal leaf beetle between
cereal fields in Michigan (124), the interleaf movement of aphids (16), and
the ebb and flow of the western tent caterpillar (157, 158). Here, each cell
is likely to be homogeneous and the herbivore tends to move only to an
adjacent cell. This approach is limited to regions where the habitat is large
relative to the total geographic area. Where the habitat area is small (10%),
such as often occurs in field crops, this grid method tends to be inefficient
if the insect under consideration has some degree of flight mobility.
Both the diffusion equation and grid modeling approaches assume that
each individual of a population [or phenotype, as in (104)] has an equal
chance of moving from one unit to an adjacent one (i.e. random walk),
which does not account for a possible "migration-prone" phase in the
insect's life cycle.
Jones (72) elucidated the relationships between movement patterns of
three caterpillar species and the spatial "patterning" of their environment.
This analysis of movement patterns included rules for initiation, duration,
and termination of movement. Behavioral changes in individuals resulting
from size, condition, or distribution of host plants or patches can alter
values of model parameters in equations that describe the distance or direc­
tion of movement. In this research, Jones (72) used detailed simulation
models incorporating probabilistic rules for movement to generate extended
movement sequences and to extrapolate broader consequences of movement
(73).
Similar approaches have been used by Siniff & Jesson (130) and Kaiser
(76). Emphasis is placed on how one individual responds to given environ­
mental conditions, and actual population sizes are not considered. Kitching
(79) extended the model of Siniff & Jesson to encompass a larger number
of individuals, simulating movement of Tribolium confusum from a fixed
 PEST MOVEMENT 329

starting point over an area containing several resource patches. This same
basic framework has also been used to model among-field movement in
Epi/achna varivestis (39).
Clark et al (24) used simulation to analyze interactions of patchiness,
movement, and population dynamics for the spruce budworm, Choris­
toneura fumiferana, in an attempt to explain observed spatiotemporal
changes in budworm patterns. The authors found that highly directional,
often aggregative, behavior was observed in a wide array of natural systems
and that random diffusion (i.e. Brownian movement) does not adequately
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explain movement of insect pest species. Five rules for movement are identi­
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fied as being more realistic than that of random diffusion (24).

1. The proportion of animals leaving a location is a function of per capita

habitat quality.
2. The exodus population determined by rule 1 moves as a patch, not as
independent individuals.
3. The population patch will go off in a single direction and settle together.
4. Migrating populations will not settle within a sizable refractory distance
of their original location.
5. Settlement of different populations will be contagious in space because
of biological and physical factors.

Although these rules may be necessary for derivation of explanatory models

of some insect pest species, Taylor (145) demonstrated the adequacy of a
simpler simulation approach based on the movement of individual desert
locusts, Schistocerca gregaria.
Descriptive and explanatory models are valuable aids in gaining a
biological/ecological understanding of insect movement. However, for in­
sect pest species, the desired application of movement models is to predict
pest influxes (e.g. 70; see 12). Stimac & Barfield (138) present a conceptual
model to identify various levels of movement processes that determine how
mobile pests are partitioned into crop and noncrop host plant communities.
Many of the biological events that determine how insect pest inoculum is
partitioned among different spatial units are poorly understood (10).
As stated earlier, a number of difficult experimental problems must be
overcome bifore data necessary to elucidate sources and patterns of move­
ment for insect pest species can be acquired. Until such time, hypotheses
on the mechanisms of movement of these species cannot be tested. Conse­
quently, the possibilities for developing predictive models of insect pest
movements are bleak.
Clark et al (24) admit the immense difficulty in developing truly predic­
tive models of habitat-movement and pest population dynamics. However,
 330 STINNER, BARFIELD, STIMAC & DOHSE

they also point out that movement studies have progressed from an earlier
preoccupation with statistical problems of pattern description (91) to the
present treatment of movement as an ecological process on the same order
as predation or reproduction. In these latter processes, a shift in perspective
from treatment as static parameters to component-structured ecological
processes yielded substantial improvements in both theory and experimen­
tation (50). Hopefully, a similar shift in perspective for the study of move­
ment of insect pests is underway.

Literature Cited
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