Agricultural and Forest Meteorology 97 (1999) 331–349

Improved understanding of dispersal in crop pest and disease

management: current status and future directions
M.J. Jeger ∗
Ecological Phytopathology Group, Wageningen Agricultural University, P.O.B. 8025, 6700 EE Wageningen, The Netherlands

Accepted 21 July 1999

Abstract
Implementation of integrated pest management (IPM) is by definition scale- and location-specific and because these features
govern biophysical processes such as dispersal, information on the latter is relevant for IPM. However, dispersal has not featured
prominently as the defining and critical element of IPM schemes. At the larger regional or inter-continental scales then our
understanding of dispersal has improved with the availability of new tracking technologies (e.g., Doppler weather radar,
see Westbrook and Isard (1999)), but proportionate returns in terms of IPM have not been demonstrated. There is increasing
interest in describing within-field and within-plant dispersal events, but much remains to be done to integrate research findings
into operational IPM programmes. Models are often seen as useful tools for analysing pest and disease outbreaks and for the
evaluation of alternative control options. Dispersal, in full biophysical detail, has rarely featured explicitly in such models, and
indeed the lack of biological balance in complex simulation models places their uses in research, technology development or
implementation somewhat in doubt. Often, however, it is possible to incorporate dispersal or more generally spatial processes
into simplified models that combine applicability to real pest management situations with the insights of theoretical population
ecology. Such insights are relevant to the multiple-species interactions characteristic of biological control and the ways in
which dispersal affects the qualitative attributes of these interactions. Dispersal research has had a demonstrable impact on
pest management and new opportunities will arise in the future. ©1999 Elsevier Science B.V. All rights reserved.
Keywords: Dispersal; Integrated pest management; Biophysical models; Spatial models; Pest and disease dynamics; Multiple-species
interactions; Landscape ecology

1. Introduction behavioural traits are involved. Very crudely, as used

Dispersal as a term has been used with different
nuances of meaning depending often on the organism
in question. In this paper it will be used as a neu-
tral term, irrespective of whether dispersion (scatter-
ing of a group of individuals) takes place or whether
∗ Current address: Department of Agriculture and Horticulture,
Wye College (University of London), Wye, Ashford, Kent, TN25
5AH, UK
E-mail address: m.jeger@wye.ac.uk (M.J. Jeger)
here, dispersal is synonymous with movement and in-
cludes both the movement of individual entities such
as insects or fungal spores, and the growth of inde-
terminate entities such as fungal mycelia. It is not re-
stricted to air dispersal but includes dispersal through
soil and in water. Dispersal by vectors other than hu-
mans is also a legitimate topic for this paper. Disper-
sal operates at different time and spatial scales which
sometimes can and sometimes cannot be separated. A
basic premise is that the importance of dispersal be-
comes apparent at different scales depending upon the

0168-1923/99/$ – see front matter ©1999 Elsevier Science B.V. All rights reserved.
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332 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
system being considered. Dispersal may occur over
tens or hundreds of kilometers between susceptible
crops, over a few meters between individual plants, or
over a few centimetres or less between roots in soil.
The important question is: what is the critical scale
for dispersal, below which a pest or pathogen remains
contained without posing a field problem, and above
which a major pest outbreak or disease epidemic may
occur.
This paper will largely confine itself to arthro-
pod pests of crop plants and plant pathogens (fungi,
viruses, bacteria and nematodes). Dispersal of verte-
brate pests will not be considered and seed or clonal
dispersal of agricultural weed plants will only be con-
sidered in assessing the status of dispersal research
in relation to pest management. Similar comments
apply to the dispersal of arthropod pests and disease
vectors of humans and animals.
During the last 10 years there have been many
authoritative reviews of dispersal in relation to the ma-
jor pest groups (e.g., Fuxa, 1987; Davis, 1987; Fitt et
al., 1989; Aylor, 1990; Madden, 1992; Loxdale et al.,
1993). Similarly, major international conferences of
an interdisciplinary nature have been held during the
last 20 years: including those at NASA Wallops Island,
Virginia in 1978; the Royal Society, London in 1982;
Baton Rouge, Louisiana in 1984; the W.G. Kellogg
Biological Station, Michigan in 1992; and an Ameri-
can Phytopathological Society/Entomological Society
of America symposium at Las Vegas in 1998. Numer-
ous symposia at other national and international con-
ferences have dealt with some of the issues relating
to dispersal and crop pest and disease management.
What must be left out of this paper is far more exten-
sive than what is included.
In an attempt to identify the importance of an im-
proved understanding of dispersal in crop and disease
management the following approach will be adopted.
(i) I consider the role of dispersal research in the de-
velopment and implementation of IPM. In parti-
cular, I ask the question: is dispersal ever the key
defining element in pest or pathogen life history
with respect to management?
(ii) How does the scale at which dispersal operates
affect pest management decision-making?
(iii) How useful for research or management purposes
are models of pest and disease dynamics in which
dispersal is accounted for explicitly or implicity?
(iv) Are there linkages between biophysical ap-
proaches to dispersal with those from theoretical
population ecology (and also genetics) that could
be better exploited?
(v) What priorities should be set for future disper-
sal research: in particular are there new areas
of application, set either from scientific or soci-
etal imperatives, to which dispersal research can
contribute?
2. Dispersal and integrated pest management
The development and deployment of IPM strategies
have been a major preoccupation of crop protectionists
and policy makers for the last 30 years, although ex-
plicit consideration of dispersal within IPM progamme
development and implementation is more recent. In
my view it is debateable whether the pursuit of IPM,
per se,has led either to an increased scientific under-
standing or to an improved management of the pests
and diseases that have afflicted and continue to afflict
the major economic crops worldwide. It is instructive
to consider dispersal in this context. Dispersal can be
seen as a key element in the development of plant
disease epidemics and in the population dynamics of
disease vectors and pests, almost irrespective of other
life history characteristics, provided a relevant tempo-
ral and spatial scale is considered. Dispersal can be
viewed and approached from many different perspec-
tives as befits a topic that truly crosses the boundaries
of many different disciplines. However, is it actually
the case that consideration of dispersal has played a
major role in the development of IPM, or has dispersal
research largely operated in contexts specific to partic-
ular organisms, crops and environments, without real
regard to its importance in management? It is remark-
able that only one review has attempted to specifically
link dispersal with IPM (Vaughn and Wolf, 1979);
where, according to the authors, IPM concepts high-
light the importance of understanding the causes and
effects of dispersal as well as manipulating dispersal
to advantage in management systems. Despite the at-
tractiveness of this mantra, it was proffered only in the
context of insect pests and today remains largely un-
appreciated by practitioners. It still remains the case
that few accounts of the importance of dispersal in
pest management attempt to cover the whole range of
 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
pest organisms (cf. Derron, 1988) or place dispersal
within the broader landscape context that character-
izes agricultural or forest land (Fry, 1995).
In an attempt to establish the importance attached
to dispersal as a component of IPM, an analysis was
made of relevant literature that purported to link
dispersal with pest, disease and weed management.
Almost 200 publications at least claimed to make this
link over the 25-year period 1974–1998. About a quar-
ter of these publications were considered to be making
exaggerated claims. Others, rather than dealing with
dispersal as a process, dealt with the use of dispersion
statistics in pest management, mostly with respect to
arthropod pests: including their use for determining
optimal sample sizes (Croft et al., 1976; Funderbunk
and Mack, 1989); sampling pattern and design (Zalom
et al., 1984; Li et al., 1992; Clarke et al., 1997), and
methods (Byerly et al., 1978); and sequential sam-
pling plans for action or economic thresholds (Iwao
and Kolodney-Hirsch, 1986; Boivin et al., 1991;
Sanderson and Zhang, 1995). There has been a sim-
ilar trend with regard to plant pathogens and diseases
(Madden, 1989) although in few cases have dispersion
statistics actually been related to dispersal phenomena.
Also omitted from the analysis were a few publica-
tions dealing with dispersal of vertebrate pests (Muria
and Rodriguez, 1989; Cowan and Rhodes, 1993),
arthropod pests or disease vectors of animals/humans
(Toyama and Ikeda, 1974; Legner et al., 1990; Yu et
al., 1995) and invasions of exotic weed species into
native vegetation (Jubinsky and Anderson, 1996).
The papers discussed in the sections below were
chosen as representative of the ways dispersal has been
considered as an important process in attempts to man-
age arthropod pests, plant pathogens and agricultural
weeds. An attempt is made to show that broadly the
same considerations apply irrespective of the pest taxa
considered. An analysis is also made of the impor-
tance of dispersal for successful biological control.
2.1. Dispersal of arthropod pests
There have been many studies of arthropod pest
dispersal in terms of basic ecological and behavioural
information that may influence the management of
these pests, even if not in an IPM context. The general
dispersal behaviour of the scale Fiorinia externa in
333
eastern hemlock forests in Connecticut was observed
(McClure, 1979) and the information used to suggest
better options for control, both chemical and through
predators. Often dispersal from different landscape
elements can cause a major pest problem to arise. The
geometrid Operophtera brumata is an irregular pest in
fruit orchards in Norway, with outbreaks every 12–15
years (Edland, 1981). Larvae are dispersed by wind
from nearby forest slopes and control measures are
then required. A monitoring strategy based on light
traps at different altitudes and distances from forested
areas gave a good forecast of larval dispersal and
when control measures should be taken. Even at a
more local level dispersal between habitats, dependent
upon behavioural and developmental changes, influ-
ences the nature and extent of pest damage as shown
in studies of the hemp moth Grapholitha delineana in
Hungary (Nagy, 1979) and can be related to its control.
Assembly flights in hemp fields generally ended in
edge zones with higher infestations and damage result-
ing in these zones. Information on dispersal within and
between individual plants can also be useful even when
the scale of dispersal is small. For example, a study
of the pine weevil Pissodes strobi in British Columbia
(McIntosh et al., 1996) showed that although consid-
erable diurnal and seasonal dispersal occurred within
white spruce trees, weevils would rarely, on average,
move further than the adjacent tree following mating
and oviposition. Dispersal within a crop may, however,
have significant effects on the ways integrated control
programmes are implemented. Larvae of the tortricid
Lobesia botransa has considerable dispersal capacity
in grapevine (Torres-Vila et al., 1997) and this, in rela-
tion to establishment and survival, must be taken into
account when sampling to determine whether damage
thresholds have been reached.
The relationship between dispersal and pesticide
treatment has often been studied to make better use
of pesticide treatments. In some cases the timing of
treatment is clearly linked to dispersive or mobile
stages so as to obtain the most efficient use of the
pesticide, sometimes outside of the crop (Vercambre,
1982). The soft wax scale Ceroplastes destructor has
an insecticide-sensitive ray stage which settles on
tangelo leaves (Blank et al., 1997); rays later disperse
onto wood forming the wax-protected 3rd instar. The
highest level of control was achieved at about the
time of maximum ray settlement, and insecticide
334 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
application timings were recommended accordingly.
In cases where flight dispersal is linked to temperature
placement of traps and lures can be placed accord-
ingly, for example with the scotylid Xyleborus dispar
(Mani et al., 1992). Similar considerations apply when
mating disruptants (Cordillot, 1987) or other use of
semiochemicals (Paiva et al., 1993) are considered
for use in IPM. Other studies have examined how
application itself is followed by greater dispersal ac-
tivity. The acaricidal activity of pyrethroids is largely
controlled by the extent of repellancy induced by
the chemicals (Penman and Chapman, 1988) which
in turn induces dispersal to other plants. This could
certainly be an important consideration in the case
of non-persistently transmitted plant viruses where
the activity rather than density of virus vectors was
shown to be an important factor in models of the
dynamics of virus transmission (Jeger et al., 1998).
Dispersal can also have a major effect on the evo-
lution of pesticide resistance (Comins, 1979) and its
management (Irwin, 1999), an aspect that has been
little researched and thus remains poorly understood,
except for a few cases such as in the tetranychid mites
(Dunley and Croft, 1992). Immigration of resistant
individuals from a sprayed crop can increase the
frequency of resistance, but equally, immigration of
susceptible individuals from unsprayed habitats can
retard resistance development or lead to its reversion
depending on the level of gene flow.
Somewhat surprising is that the effect of differ-
ent crop management or cultural practices (pruning,
sanitation, defoliation, irrigation) on dispersal has re-
ceived relatively little attention. Increasing the plant
spacing delays or even prevents the spread of the fir
aphid Cinara todocola within plantations (Furuta and
Aloo, 1994). In studies on cotton production prac-
tices in relation to later season crops (Nuessly et al.,
1994) it was possible to define latest irrigation and
defoliation dates for cotton, combined with earliest
planting dates for the other crops, to avoid serious
problems caused by dispersal of the whitefly Bemisia
tabaci into the later crops. Survival and dispersal of
Aphanostigma piri are major factors affecting the in-
cidence and severity of pear phylloxera (Moussion,
1982), and a range of cultural practices that minimize
the pest problem have been developed, involving
manipulation of fertilizer use, ground cover, varietal
susceptibility, and bag protection. At the landscape
level, the spatial pattern of used and abandoned
orchards, together with the surrounding landscape
habitats, can explain a large proportion of the flying
activity of tortricids (Jeanneret, 1997).
2.2. Dispersal of plant pathogens
Published research on pathogen dispersal in rela-
tion to disease management has covered the whole
range of modes of transport, including airborne and
splash-borne dispersal of fungal spores, and soil-borne
dispersal of spores and/or mycelium. However, de-
spite the considerable literature on this topic, many
studies do not specifically consider dispersal in the
light of disease management options — a criterion for
inclusion in this analysis. Many of the different scales
that arise in dispersal of arthropod pests arise again
with pathogens. The patterns of epidemic develop-
ment of tobacco blue mold in the eastern USA and
Canada were consistent with the long-range trans-
port of spores of the pathogen Peronospora tabacina
(Davis and Main, 1984, 1986). Such information can
be used for the evaluation of decisions concerning
disease management (Aylor, 1986) and in operational
disease warning systems, such as the North Carolina
forecasting system for blue mold. Also at the regional
or continental scale the phytosanitary and other op-
tions for managing or limiting spread of Dutch elm
disease and canker stain of plane (Ceratocystis fim-
briata f.sp. platani) within Europe were compared
by Smith (1985); with a widespread distribution of
aggressive strains of the Dutch elm pathogen, but
with human agency largely being responsible for
spread of the latter pathogen. The fungal pathogen
Mycosphaerella graminicola occurs in all wheat
growing areas of northern Europe (Mielke and Ahlf,
1985) with ascospore infection, possibly from distant
sources, giving rise to initial infections followed by
more limited dispersal by asexual spores within-field.
These dispersal characteristics contribute to the ability
to forecast disease development from early infections.
At the individual plant level dispersal of the Karnal
bunt pathogen of wheat (Tilletia indica) from soil to
ears is a key element in predicting disease severity and
developing disease management practices (Nagarajan
et al., 1997). A key factor in understanding the epi-
demiology of fungal plant pathogens is the genetic
 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349 335

composition of the pathogen population, and the role As with phytoplasmas, research on virus dispersal
of dispersal in determining patterns of diversity at a (Thresh, 1985) has tended to emphasize arthropod
range of scales. For example, Limpert (1988) moni- vector dispersal, although of course fungi and ne-
tored airborne populations of barley mildew through- matodes also vector important plant diseases mostly
out cultivated areas in western Europe and found below-ground, where dispersal has been much less
essentially a single epidemiological unit because of studied. A wide range of fungal and bacterial plant
the wind-dispersed nature of the pathogen. On this pathogens are known to be dispersed passively by
basis a system for Europe-wide deployment and use human and animal agents (Warner and French, 1970;
of host resistance and fungicides was proposed. Bashan, 1986).
As with the case for arthropod pests, knowledge of
dispersal can be critical in determining the optimal
2.3. Dispersal and weeds
timing for fungicide sprays. For example, the most
effective treatments with triazole fungicides for con-
Recent research on weed dispersal in relation
trolling Leptosphaeria maculans infections in oilseed
to IPM mostly covers seed dispersal and survival
rape during the 1–6 leaf stage were those applied at
(Regehr and Thomas, 1994; Jordan, 1996; Debaeke,
the beginning of ascospore dispersal (Penaud, 1993).
1997; Wallace, 1997) with some elaborations specific
Some disease warning systems are based on spore
to the nature of the weed problem. The preven-
sampling data to initiate control programmes: for
tion and control of Solanum viarum in pasture and
example, potato early blight in Colorado (Harrison
open forest was achieved by reducing or eliminating
et al., 1965) and hop downy mildew in Germany
seed dispersal by herbivore vectors (Mullahey et al.,
(Kremheller and Diercks, 1983). Some research has
1996). As with insecticides and fungicides, dispersal
also considered the effects of crop management
influences the development of herbicide resistance.
(greenhouse fumigation, nursery practices, mulching,
Herbicide-resistant biotypes of wild oats infest many
plant thinning, sanitation) practices on pathogen dis-
cereal producing regions and a major means of delay-
persal. Dispersal of soil-borne inoculum of Phytoph-
ing or preventing resistance development is to prevent
thora nicotianae var. parasitica to tomato fruits can
dispersal (from immigration or within-field) of oat
be obstructed by polythene mulch, and is as effective
seed into the soil (Thill et al., 1994). When comparing
as a fungicide (Dodan et al., 1994). In glasshouses,
tillage and no-tillage systems for maize it was found
the prevention of recontamination of disinfested soil
that emergence of the most abundant weeds was re-
by airborne spores of Fusarium oxysporum f.sp.
duced by tillage (Mohler and Calloway, 1992) except,
radicis-lycopersici is crucial in controlling crown and
as in the case of Taraxacum officinale, when there was
root rot of tomato (Franceschini et al., 1996). As well
a short time between dispersal and germination. Seed
as for fungi, cultural controls are important for the
dispersal and grazing regime can interact in deter-
prevention of dispersal of nematodes, especially in
mining the optimal management of a weed problem,
nurseries (Esser, 1996), bacteria (Pohronezny et al.,
as found for Hieracium spp. management in New
1990) and phytoplasmas (Purcell et al., 1987). Hand
Zealand pastures (Espie, 1994). Parasitic plants, such
thinning of direct seeded tomato when foliage was dry
as Striga and Orobanche spp., have many features in
in the afternoon reduced disease incidence of bacterial
common with soil-borne plant pathogens possessing
spot (Pohronezny et al., 1990). Use of bacteriocidal
an aerial phase, and their modes of dispersal have
hand washes to avoid human dispersal of the bac-
resulted in an extensive geographical distribution (Fer
terium also proved useful in an integrated programme
and Thalouarn, 1997).
for management of the disease. Cherry decline in
California is caused by a phytoplasma transmitted
by cicadellids (Purcell et al., 1987). The vector over- 2.4. Dispersal of biological control agents
winters on a range of other shrubs and disperses to
cherry in late spring or summer. Management prac- 2.4.1. Arthropod pests
tices now involve the removal of any ornamental food One interesting outcome of the analysis linking dis-
plant of the vector situated close to cherry orchards. persal with pest management is that the dispersal of
336 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
biological control organisms, notably natural enemies
of arthropod pests, has received at least as much atten-
tion as dispersal of the pests themselves. Most of the
research on biological control of arthropods has been
with arthropod natural enemies, with only a few pa-
pers linking dispersal with pathogens of arthropods. In
much of the research on pathogens, emphasis has been
placed on inundative application, where the dispersal
and survival of the pathogen may be counter-indicated,
especially if pathogenicity has been genetically en-
hanced. Many studies have examined dispersal of a
range of natural enemies of a given pest or pest com-
plex. Many of the studies are directed to the ability of
natural enemies to recover after pesticide application
and how the speed of recovery is related to dispersive
ability. Such research has been typical of long-term
projects comparing ‘conventional’ and supervised sys-
tems of pest management such as the Boxworth project
in the UK (Burns, 1988, 1992). Such studies impose
severe methodological constraints on the experimenta-
tion and the interpretations of the results. Investigation
of long-term effects on populations of highly disper-
sive predators may require extensive studies over many
years (Sunderland, 1992).
Long-term management experiments involving a
range of cropping patterns as well as pesticides have
also shown the importance of natural enemy disper-
sal in reducing pest populations, and pinpointing the
problems caused by limited dispersive capacity, for
example of laboratory-bred parasites (Cameron et al.,
1983).
The numbers of studies on dispersal of individual
natural enemy species are too many to document in
full and only representative examples are discussed.
Generally, the research emphasis has been on the
relative dispersive capacity of the natural enemy
compared with the pest, side effects or differential
responses to pesticide application as noted above,
the occurrence of pesticide-resistant natural enemies,
and ways of manipulating the crop and surround-
ing habitats to facilitate the survival and dispersal
of the natural enemy. Particularly prominent in this
respect have been studies on predatory phytoseiid
mites (Burnett, 1979; Hoy et al., 1985; Hoy and Cave,
1988; Courcy-Williams, 1993). For other predators,
the dispersal characteristics of coccinellid beetles (Xu
and Wu, 1989; Tondeur et al., 1993; Dreistadt and
Flint, 1996) and spiders (Thomas, 1988; Thomas et
al., 1990; Jepson and Thacker, 1990) are well repre-
sented in the literature. For parasitoids there has been
a considerable number of studies of the Encarsia for-
mosa/Trialeurodes vaporariorum system in particular.
It is perhaps not coincidental that biological control of
the glasshouse whitefly has been well established in
commercial production in Europe, given the detailed
attention afforded to dispersal of the parasitoid in
glasshouses (van Lenteren et al., 1976, 1992; Hussey
et al., 1976).
Research on the use of insect pathogens has
long been dominated by an inundative approach, al-
though according to Fuxa (1987) this approach is
unlikely to be successful due to the limited num-
ber of quick-acting pathogens; therefore, the greatest
potential for the use of pathogens in IPM lies in
the inoculative augmentation approach. However, it
seems that natural dispersal of entomopathogenic
fungi may be insufficient by itself, especially as
behavioural modifications in infected insects may
restrict conidial dispersal (Carruthers et al., 1985).
However, one possibility is to use other agents,
e.g., honey bees, to vector pathogens from specially
constructed applicators, as reported by Gross et
al. (1994) to deliver Heliothis nuclear polyhedrosis
virus against clover pests. The dispersal and for-
aging behaviour of the honey bee then serves to
disperse the virus to attack phytophagous insects.
The possibilities for improved strain selection and
of genetically engineering pathogens to have greater
and faster-acting effects, however, places the em-
phasis again on an inoculative approach in which
recombinant microbes, once they have done the job
intended, do not survive and disperse in the envi-
ronment. The analogy with pesticide use and the
avoidance of pesticide residues is then complete. Ul-
timately, the risks and ramifications of introducing
genetically-modified microbes for pest control must
be evaluated and quantified (Walter et al., 1990),
and this is a burgeoning area for epidemiological
and ecological insight (Mundt, 1995). Developments
in molecular biology mean that completely new ap-
proaches to ‘pathogen-like’ control of pests are now
possible, such as the cloning of insecticidal genes
and the use of mobile genetic elements as vectors for
the dispersal of the construct through a population
as a consequence of mating (Pfeifer and Grigliatti,
1996).
 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
2.4.2. Weeds
There have been some studies on dispersal in re-
lation to biological control of weeds and most of
these were concerned with arthropods. Sometimes
the very lack of dispersive ability in a potential agent
can lead to dispersal through community intervention
and involvement (Jupp, 1996). More direct successful
introductions of biological control agents based in
part on a knowledge of dispersal characteristics can,
however, be found in the literature. Purple loosestrife
is a wetland perennial introduced to North America in
the 19th century, occurring in large monotypic stands.
A biological weed control programme using several
insect species began in 1986 based on wide-ranging
ecological investigations, including patterns and rates
of insect dispersal (Blossey et al., 1996). The pro-
gramme has resulted in the successful establishment
of some species and the authors claim that biologi-
cal weed management can now be incorporated into
national and international pest management strate-
gies. Dockplants (Rumex spp.) are common plants in
many agricultural and disturbed habitats and in some
situations are considered weeds. The chrysomelid
beetle (Gastrophysa viridula) is a potential biological
control agent for docks but its ability to disperse and
invade changeable habitats must be understood for it
to be successful (Whittaker et al., 1979). Herbicide
application can also affect the dispersive behaviour of
insects used for biological control of weeds. Several
weevils (Neochetina spp.) have been used to con-
trol the aquatic weed water hyacinth. Weevils were
found to consistently disperse from sprayed dying
plants to healthy plants, which could potentially be
exploited in integrated control of the weed (Haag,
1986).
There have been few substantive studies of the
role of dispersal in naturally-occurring pathogens
of weeds, or those applied as mycoherbicides,
where again inundative application and complete
coverage of the target weed is the recommended
practise. The mycoherbicide COLLEGO is a com-
mercial product for the control of the weed northern
jointvetch in rice. Understanding the dispersal of
the pathogen Colletotrichum gloeosporioides by rain
splash (Yang and TeBeest, 1992) and by grasshop-
pers (Yang et al., 1994) was considered to be
an important element in the development of the
mycoherbicide.
337
2.4.3. Plant pathogens
Very little research has been concerned with dis-
persal of biological control agents of plant pathogens
within an integrated control strategy. The most suc-
cessful biofungicide for foliar pathogens in commer-
cial use, AQ10TM, is based on the hyperparasitic
fungus Ampelomyces quisqualis. This product is
produced using large-scale fermentation and is for-
mulated as a water-dispersible granule (Daoust and
Hofstein, 1996) and used mainly against powdery
mildew in grapes or under glass against cucumber
powdery mildew. The hypoparasite is partly tolerant
to some fungicides used for powdery mildew control
(Sundheim, 1982b). It is known that passive dispersal
of the hyperparasite by powdery mildew conidia over
large distances occurs (Philipp et al., 1984) and that
the presence of the powdery mildew host promotes
germination (Sundheim, 1982a); however, application
is made solely as a biofungicide without any con-
sideration of the potential for natural dispersal as a
component of integrated control. There is perhaps an
under-appreciation of the potential role of dispersal in
biological control of pathogens; notably the dispersal
of mycoparasites by arthropods above-ground during
pollination (Sutton and Peng, 1993), or below ground
by soil invertebrates.
3. Dispersal as a biophysical phenomenon
As stated in Section 1 this paper takes an inclu-
sive definition of dispersal, dependent only on the
time and spatial scales considered. Above-ground and
below-ground dispersal, and the biophysical factors
involved are both included. In practice however stud-
ies on dispersal below-ground in relation to pest man-
agement are few.
3.1. Dispersal in soil
Wallace (1978) reviewed the topic with respect to
plant pathogens, including nematodes, but there has
been little progress since. Motility of soil beneficial
bacteria is an important aspect of introducing these or-
ganisms into cropping systems (Bashan and Holguin,
1994; Boelens et al., 1994; Parco et al., 1994), and sim-
ilar studies are required for soil-borne pathogens and
338 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
biological control organisms (Parke, 1991). Lateral
flow of subsurface water can be an effective means of
dispersing zoosporic fungi (Kinal et al., 1993) with
potential movement of up to 120 m following intensive
rainfall events. Flooding (Kenerley et al., 1984) and
furrow-irrigation (Grinstein et al., 1983) have been
implicated in the within-field dispersal of soil-borne
pathogens. On a smaller scale, homing responses of
zoosporic fungi are well documented (Deacon and
Donaldson, 1993) and may be critical in determining
the severity of root diseases, as indeed might be the
disturbance of these homing responses by potential an-
tagonists (Dandurand and Menge, 1994). The disper-
sal of fungi through soil by means of mycelial growth
is less understood with few experimental observations
reported (Kenerley and Jeger, 1990; Kenerley et al.,
1998) due to the difficulties in direct visualization. The
rhizomorph forming fungus Armillaria produces ex-
tensive networks extending over hundreds of hectares
but this characteristic again raises the question as to
whether this is growth (of an individual mycelium)
or dispersal of the fungus. There are few studies on
the below-ground dispersal of arthropods in relation
to pest management, except for the case of larval
stages and forecasting the timing of their emergence
in relation to crop phenology (Finch et al., 1996).
The remainder of this section deals with dispersal
above-ground.
3.2. Airborne dispersal
As a biophysical phenomenon, airborne dispersal
integrates environmental physics, meteorology and
models of aerial dispersion; and in the case of insect
dispersal, behavioural responses. As such there have
been major advances in our ability to monitor and
track populations through technological advances in
equipment, information retrieval and data processing.
It is not the intention here to review these develop-
ments, which are covered by other contributors to this
volume (Gage and Isard, 1999; Westbrook and Isard,
1999; Aylor, 1999). Airborne dispersal can be both
long-range and short-range, depending on the time
scale set, and there have been many attempts to explain
the movement of major pests and diseases worldwide
in terms of long-range dispersal between continents.
These attempts range from the anecdotal, involving
inspired calculations (Simmonds, 1994), the tracing of
historical records in major traded crops such as banana
(Burt, 1994), to the development of long-range trans-
port models based on atmospheric conditions (Aylor,
1986; Davis, 1987). In a similar way, Loxdale et al.
(1993) consider the evidence for long-range move-
ment of a particular pest and virus-vector grouping,
aphids, and its significance relative to local move-
ment. They concluded that short-distance movement
has a greater effect on population distribution than
long-distance movement. This conclusion would be
particularly relevant to plant virus epidemiology and
the spread of insecticide-resistant genotypes. Thresh
(1983) specified six conditions that should be satis-
fied in terms of providing evidence for long-distance
‘spread’ of plant viruses, but these have rarely been
rigorously followed. Case studies for a range of other
pests and pathogens, and dispersal types are given
elsewhere (Limpert, 1999; Byrne, 1999; Shields and
Testa, 1999).
In some cases such explicit knowledge on
long-range dispersal can be used directly to inform
strategic and tactical decisions, although generally
this has been the case at regional scales only. From
this perspective aerobiological research can be seen
as a mature science, but does it present the whole pic-
ture and does it really contribute to pest and disease
management at the local level? For spore dispersal
within a crop canopy there has been established a rich
theory combining environmental physics, detailed
observations, epidemiological insight and mathemati-
cal models that was brought together in the late 70s
and early 80s (Aylor, 1978; Legg and Powell, 1979;
Shaw, 1982; Legg, 1983; Raupach and Legg, 1984).
Since then there has been an increased apprecia-
tion of factors such as the role of intermittent wind
(Aylor, 1990) in dispersal and at a finer scale the
micro-conditions governing spore release (Aylor and
Anagnostakis, 1991). Ferrandino (1993) has related
the three-dimensional turbulent dispersal of airborne
spores to the types of contact distributions typically
used in theoretical epidemiological models and shown
how distinctive patterns of disease expansion from
foci can occur, as discussed in the next section. De-
spite this well-developed theory for airborne dispersal
there have been few attempts to incorporate detailed
dispersal information into epidemiological investiga-
tions and disease management schemes at this scale.
 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
Notable exceptions are the work on apple scab man-
agement (Aylor, 1999) and the blue mold warning
system referred to earlier.
It is also the case that surprises can still be found in
studies on airborne dispersal. For example, sorghum
downy mildew is caused by a pathogen that produces
long-lived oospores. Such survival propagules are
normally associated with the soil and the immediate
vicinity of where a susceptible crop is grown. De-
tailed epidemiological studies (Bock et al., 1997) have
shown, however, that oospores, because of the way
they develop within diseased leaf tissue, can be freed
and be dispersed as though they were airborne spores,
with serious implications for attempts to manage this
disease.
3.3. Splash-borne dispersal
The literature on splash dispersal, despite some
pioneering studies in the 50s and 60s, is much more
recent than that of airborne dispersal and is virtually
restricted to fungal and bacterial plant pathogens.
Splash dispersal is intimately linked to rainfall events
(Fitt et al., 1989; Madden, 1992) and with the forma-
tion and subsequent fate of water drops and films on
plant surfaces (Butler, 1990). Detailed photographic
analysis has revealed the sequence of physical events
by which fungal spores contained in water drops
or films on leaf surfaces are dispersed within the
secondary droplets caused by the impaction of rain
on the original droplet (Walklate, 1989; Yang et al.,
1991). A range of innovative experimental studies
simulating rainfall of different qualitative and quan-
titative characteristics has revealed the patterns and
extent of splash dispersal possible during rainfall
(Reynolds et al., 1987, 1989). Simple monitoring de-
vices (a splash meter) have been proposed as a means
to forecast when significant disease increase follow-
ing splash dispersal can be expected (Shaw, 1987).
Physical models of splash dispersal have also been
developed. Yang et al. (1991) applied a modified dif-
fusion equation to parameterize the effects of rainfall
intensity and surface topography on splash dispersal
of strawberry pathogens, and considered the model to
hold considerable promise for predicting the effects
of different ground covers. Subsequently, Pielaat and
van den Bosch (1997) and Pielaat et al. (1998) have
339
developed a mechanistic model based solely on phys-
ical considerations and shown that under limiting
circumstances the solution approaches that of the dif-
fusion equation. This is an excellent example of how
a relatively parsimonious model based on biophysical
principles can be developed and related to a more
general model that has found many areas of applica-
tion in population ecology. This is a theme that will
be returned to later in this paper.
There have been few studies on the effects of rain-
fall on dispersal of arthropod pests or vectors, or of
dispersal within splash droplets in any way analogous
to splash-borne plant pathogens; although the effect of
rain on disease vectors at the mesoscale has been well
described (Smith, 1983) and rainfall generally inhibits
active movement of most arthropod pests (Gage and
Isard, 1999).
4. Dispersal and pest and disease development
There have been many experimental and observa-
tional studies on the influence of dispersal on pest
and disease dynamics. It is not possible to cover all
the relevant details in this section, but relevant ex-
amples include: the timing of initial establishment of
the aphid vectors of beet yellowing viruses (van der
Werf et al., 1992), and the spread of maize chlorotic
dwarf virus in relation to introduction of the leafhop-
per vector (Madden et al., 1990) as examples of vec-
tored dispersal; the spread of bean rust in relation to
local dispersal of the pathogen (Aylor and Ferrandino,
1989, 1990) and the relationship between aerial spore
concentrations and disease development (Aylor and
Kiyomoto, 1993) for fungal pathogens. It is sometimes
possible to use characterizations of disease gradients
as an indicator of dispersal (gradients) especially when
the disease source is known and molecular markers are
available to identify genotypes (Zwankhuizen et al.,
1998). This study in the Netherlands was able to un-
equivocally identify major sources of inoculum for
the regional epidemic, including organic farms, and
as a consequence make control recommendations in a
locality where conventional and organic farms are in-
termingled. Dispersal research can then lead to broader
cultural and political implications.
Sometimes, especially with virus diseases, there is
an apparent discrepancy between the known biology
340 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
and behaviour of the vector and the apparent rate
of spread of the disease in the field. An example
of this is cocoa swollen shoot disease caused by a
mealybug-transmitted virus that occurs in all the co-
coa growing areas of west Africa. Despite the very
limited mobility of the vector, which only spreads
by branch-to-branch contact and occasional wind-
borne dispersal of first instar nymphs, the disease had
spread apparently rapidly and showed no signs of
diminishing despite an eradication campaign that has
been one of the largest ever mounted against a plant
disease (Thresh et al., 1988). The resolution of this
discrepancy comes essentially from the ‘landscape’
features of the environment in which cocoa is grown,
in close proximity to wild hosts and with little sep-
aration between the individual (small) holdings. In
the case of cocoa swollen shoot disease, modelling
was found useful in attempts to understand the im-
plications of the dispersal process and to evaluate the
effectiveness of control options such as the introduc-
tion of a cordon-sanitaire around new plantings (Jeger
and Thresh, 1992) and the removal of diseased trees
(Chan and Jeger, 1994).
Dispersal can be incorporated conceptually and
mathematically into models of disease epidemics and
pest outbreaks occurring at a more local scale. Broadly
two approaches have been followed, which treat dis-
persal either explicitly or implicitly in the models.
4.1. Dispersal as an explicit process
In the first approach detailed knowledge of disper-
sal within-field or between fields, where it exists, is
combined with information on colonization/infection
and reproduction/sporulation for pests/diseases, and
an attempt is made using complex simulation models
to integrate these processes spatially and temporally
to obtain an overall prognosis on a desired manage-
ment intervention. There have been many examples
of the development of complex simulation models in
the entomology, plant pathology and weed science lit-
erature, but only one example will be referred to in
this section, mainly because it has the rare distinction
of covering insects, plant pathogens and a weed plant
within the same paper. The simulation modelling ap-
proach was illustrated by van der Werf et al. (1989)
for the biological control of an understory cherry in-
vading forest habitats, the motivational and preda-
tory behaviour of a carabid beetle, and the dispersal
of green peach aphid causing beet yellowing disease.
Undoubtedly in these examples there was consider-
able insight obtained into the processes underlying
spatial dynamics and some clues obtained as to pos-
sible control interventions; however, it is a personal
view that this approach has rarely led to either an im-
proved insight scientifically or more effective man-
agement. However, individual-based models in which
the characteristic dispersal or movement of individual
insects is integrated into an aggregated description of
population dispersal (as with the carabid beetle exam-
ple cited above), represent a relatively new approach
that can lead to new insights into the significance of
dispersal for pest management.
One of the major problems associated with a com-
plex simulation approach is that it is very difficult to
ensure sufficient balance in the weights attached to the
individual components that comprise the model. The
temptation is always to incorporate as much biologi-
cal and empirical detail as is available irrespective of
the paucity of information in other components (Jeger,
1986). Thus, with dispersal there may be limited in-
formation available but yet a linkage is attempted with
what might be considerable information on other com-
ponents of the pest life cycle or disease epidemiology.
An extreme example, not specifically related to pest
biology or management, may be found in a recent
paper describing a landscape simulation model which
aims to investigate the effect of habitat heterogeneity
on species diversity patterns (Ziv, 1998). This is a com-
plex but general simulation model which requires con-
siderable information according to the object-oriented
design, and yet dispersal (movement of individuals be-
tween patches in the landscape) is described by a sin-
gle idealized distribution, the generality of which is
simply not known.
Another problem with deterministic simulation
models is that processes are averaged out, whereas it
is often extreme values that are important, as shown
by Mayer et al. (1993) for dispersal of the screw-
worm, an exotic pest in Australia (Fig. 1(a)). This
study compares the outcome of individual-based sim-
ulations using a stochastic unbounded distribution to
represent dispersal distances from an initial invasion
point, compared with a deterministic version. By the
fifth generation of the screwworm fly the ‘far movers’
 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349 341

Fig. 1. (a) Spatial dispersal pattern for the screwworm fly from an invasion point at the sea–land interface (Mayer et al., 1993): LEFT,
dispersal is modelled on an individual female basis using a Cauchy distribution for distance and a uniform distribution for direction;
RIGHT, dispersal is modelled on a grid cell basis with populations moving a random distance in a random direction; in both cases the
median distance dispersed was 2 km and each character represents a 20 × 20 km cell with 䊏, +, and − representing >50, 5–50 and <5
screwworm flies, respectively. (b) Two-dimensional simulated epidemic based on diffusion and spatial probability models for disease spread,
with infections represented by dots (Minogue, 1989): UPPER, dispersal is modelled using an exponential distribution; LOWER, dispersal
is modelled using a Pareto (non-exponential tail) distribution; in both cases the mean distance of dispersal events was set at 3% of the long
axis. (c) Colonization of an array of demes by grasshopper races (Nichols and Hewitt, 1994): UPPER, colonization is modelled analagously
to the Fisherian advance of an advantageous gene; LOWER, colonization is modelled by weighing the sum of two normal distributions
allowing for long-distance dispersal; in both cases the variances of the distributions were bounded and equal, but the tails differed.
342 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349

in the stochastic model had clearly established low physically-based model of focus expansion which
density populations beyond the advancing point of described how disease would develop in time and
the deterministic model (Fig. 1(a)). space. An interesting feature of this model was that it
predicted non-wavelike expansion of the focus front.
4.2. Dispersal as an implicit process Previously, based on epidemiological models which
treat dispersal implicitly through a contact distribution,
most models proposed for plant pathogens predicted
An alternative to the complex simulation approach
an asymptotically constant expansion rate, in which
is to consider dispersal (providing the spatial ele-
the front moved at a constant distance per unit time
ment) implicitly within basically temporal models.
(Zadoks and van den Bosch, 1994; van den Bosch et
This again has limitations in terms of providing di-
al., 1999). By contrast the dispersive fronts predicted
rect management recommendations, but does provide
by Ferrandino (1993) accelerate as the front expands.
valuable information on sampling protocols and the
In fact some theoretical models also predict
derivation of pest/disease and damage relationships.
non-wave-like expansion and the critical aspect is not
It has also proved possible to indicate the relative
so much the introduction of stochastic elements in
importance of dispersal compared to other epidemi-
the model, but rather the form of the contact distri-
ological or population dynamic processes, and the
bution chosen. In fact if the dispersal function is not
extent to which a measure of dispersal can be taken
bounded with an exponential tail then characteristic
as an indicator of the future severity of an epidemic
non-wave-like or dispersive fronts emerge (Minogue,
or pest outbreak. An example of this approach is also
1989; Shaw, 1995) (Fig. 1(b)). The two authors used
given by van der Werf (1995) in which a threshold
slightly different distribution functions to represent
criterion is determined for an aphid vector outbreak
non-exponential tails but in both cases the mean and
in terms of life history parameters relating to the
median dispersal distances were equated to that of
aphid and also to a coccinellid predator. Immigration
an exponential distribution and the patterns of focus
and emigration (relative) rates appear in the model
expansion simulated. As in the case of invasion of the
specification and also in the derived criterion. Thus,
screwworm fly, the pattern that emerges in Minogue’s
it is possible to evaluate the influence of dispersal
simulations (Fig. 1(b)) clearly shows the emergence
treated implicitly as immigration/emigration against
of satellite foci in the non-exponential-tail version
other parameters such as birth and death and feeding
ahead of the advancing front of the exponential-tail
rates. A similar approach to a more complex situation
version. Shaw’s results using the Cauchy distribution
was also taken by Jeger et al. (1998) who developed
for dispersal distances (also used by Mayer et al.
a general model of virus-vector dynamics and derived
(1993) for the screwworm fly) gave similar patterns,
a composite basic reproductive rate, which effectively
although the analysis was taken further by looking at
gives a criterion as to whether a virus epidemic will
the features of dispersal on different scales and the
occur or not. In one version of this model the ef-
fractal properties observed.
fect of immigration and emigration was included. It
An important innovation by Zadoks and co-workers
is again a personal view that this approach is more
was to generalize the process of focus expansion and
suited than complex simulation to study dispersal in
to designate zero-order, first-order and second-order
more complicated contexts involving multiple-species
epidemics, depending largely on the spatial and time
interactions, as found in biological control systems
scales considered. Typically foci expand at rates of
and vectored plant diseases.
cm/day, km/day and km/year for these classes, re-
spectively (Zadoks and van den Bosch, 1994). These
4.3. Spatiotemporal models authors also claim the general applicability of their
methods to virus diseases, insect pests, parasitoids
The final example of how dispersal may be con- and vertebrates. Recently, van den Bosch et al. (1999)
sidered implicitly within basically temporal models showed that it is possible to scale up from small-scale
concerns focus expansion in disease epidemics. As disease expansion to large-scale pandemics in which
mentioned earlier, Ferrandino (1993) developed a diseases literally sweep across whole continents and
 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
moreover to account for discontinuities in cropping
seasons — a feature which was previously supposed
to present intractable analytical problems.
An interesting parallel to the work on focus
expansion is given by the effect of perhaps rare
long-distance dispersal on gene frequencies in pop-
ulations during periods of range expansion. In cir-
cumstances where such rare events are occurring, the
advance of a population can explain geographical
patterns of allelic distributions that were previously
not understood (Nichols and Hewitt, 1994), in this
case for grasshoppers (Fig. 1(c)). These authors mod-
elled colonization distances using the weighted sum
of two normal distributions (which have exponential
tails), such that by varying the value of one parame-
ter the (bounded) variances could be equated but the
tails would differ substantially. The parameter can
be varied to represent the conventional steady Fish-
erian wave of advance (of an advantageous gene) at
one extreme to irregular progress by a sequence of
long-distance colonizations at the other as shown in
Fig. 1(c). The importance of this study is that it shows
that although a non-exponential tail may be sufficient
to generate non-wave-like expansion it appears not to
be necessary.
The studies by Mayer et al. (1993), Nichols and
Hewitt (1994), and Shaw (1995) are important in that
they show not only that dispersal can be modelled
in similar ways for pathogens and insect pests, but
that the same general patterns and conclusions regard-
ing wave-like and non-wave-like spread emerge in the
different areas covered of population dynamics and
genetics. Moreover, it is clear that further theoretical
insight can be gained by continued research in this
area.
5. Dispersal and multiple-species interactions
A further perspective to be considered in relating
dispersal to IPM is that of theoretical population eco-
logy. There is a voluminous literature in theoretical
population ecology and some of this considers the role
of dispersal, usually represented in implicit terms, on
multiple-species interactions. Dispersal in spatially
complex environments leads to equally complex pat-
terns of abundance (Kareiva, 1990) that may change
or be a consequence of the ways in which species
343
interact. Predator–prey and host–pathogen interac-
tions are affected by heterogeneity, in which again
dispersal may play a part. Theoretical models (Auger
and Faivre, 1993) have shown that where species are
either competing, or where there is selective pre-
dation, the phenomenon of extinction waves arises,
presumably subject to the same kind of conditions as
found for epidemic expansion or the movement of an
advantageous gene.
In particular, questions concerning the stability
properties of species interactions have long dominated
the theoretical population ecology literature. Whether
in the specific context of metapopulations or spatially
distributed populations more generally, the question
of how dispersal affects stability properties has been
raised. It seems that dispersal would rarely stabilise a
species interaction such as occurs in biological con-
trol, but can often serve to destabilize depending on
the differential dispersal characteristics of the species
involved (Rohani et al., 1996). It is doubtful whether
this sort of generalization gives sufficient justice to
the myriad patterns of dispersal known across all pest
biota, but nevertheless the mathematical analysis has
considerable merit and relevance given the impor-
tance attached to the dispersal of biological control
agents noted earlier in this paper.
In several sections of this overview the biophysi-
cal and theoretical population ecological approaches
have been contrasted. One of the most significant chal-
lenges for future dispersal research is to find a means
of integrating these two approaches, each with differ-
ent histories and scientific traditions, into models that
can deal with the complexites of multiple-species in-
teractions in agricultural, forest and natural habitats
in a parsimonious way. Although parsimonious, the
models should still incorporate sufficient biophysical
realism to turn generality into a basis for pest man-
agement action.
6. Dispersal and new opportunities for research
The above sections have attempted to show how
an improved understanding of dispersal could prove
important in developing and implementing IPM pro-
grammes. The realization that dispersal may be criti-
cal to the success of such programmes may have come
late, but there is no excuse now for not deepening and
344 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
broadening the scope of dispersal research. Studies
on dispersal must broaden in scope from single pests
or pathogens to interacting species, as exemplified for
example by vectored plant diseases and biological con-
trol, and an appropriate theoretical framework both for
modelling and the design of appropriate experiments
will be required. Additionally, it can be argued that re-
search on dispersal has over-concentrated in the past
on pests/diseases of agricultural or forest crops. The
current debate, scientifically and politically, on the im-
portance and value of biodiversity raises issues that
necessarily involve consideration of the role of disper-
sal, for example in the conservation of natural vege-
tation. This may simply mean applying the tools of
dispersal research to invasive species or exotic pests
but with the emphasis on natural systems. Examples
of such research already exist: with weeds (Bergelson
et al., 1993); fungi as evidenced by the widescale
destruction of the west Australian jarrah vegetation
by Phytophthora cinnamomi; and with insects, cer-
tainly in an agricultural context (van Lenteren et al.,
1987).
Within a broader biodiversity framework there is
much current debate on the merits of in situ conserva-
tion of the progenitors, wild relatives and land races
of the major economic crops. Arguably such conser-
vation is desirable because it provides an arena for
the continuing evolution of these plant populations
with their pathogens and pests. Thus, through these
dynamic processes valuable resistance traits are both
conserved and generated de novo. It follows that if we
seriously wish to promote in situ conservation of plant
genetic resources then this necessarily implies in situ
conservation of pathogens and pests. If sites are to be
identified and managed for conservation, what are the
implications in terms of dispersal: either as immigra-
tion from near or far; or as emigration, and the threats
posed, to surrounding agricultural habitats? This again
is an argument for broadening the horizon of dispersal
research within a landscape ecological perspective.
7. Conclusions and key questions
This paper has attempted to provide an overview of
the importance of dispersal in crop pest and disease
management, to evaluate options for incorporating im-
provements in understanding of dispersal into IPM,
and to outline new directions for research over the next
decade. No overview is complete without a set of con-
clusions. However in this case, given the broad scope
of the paper, covering all the major crop pest taxa, it
is necessary to counterbalance each conclusion with a
key question.
IPM, by definition, can only make sense in terms
of implementation. Implementation, however, is bet-
ter secured if the specific details of site, scale, and the
host–pest complex are incorporated within an under-
lying conceptual model which recognizes the impor-
tance of dispersal.
A key question (1) is: for what circumstances can
dispersal be considered as the defining or crucial ele-
ment in IPM?
Dispersal research has matured considerably over
the period that IPM has been promoted and imple-
mented. Information on long-distance dispersal of
some pests is relevant for IPM and methodologies for
monitoring and predicting long-range dispersal events
have become increasingly sophisticated.
A key question (2) is: if short-range dispersal is
of more significance for IPM are there diminishing
returns in investing in further technological advances
relevant for tracking long-distance dispersal?
Models of pest outbreaks and disease epidemics
have proved useful tools in the development of IPM
programmes. These, however, rarely give an explicit
account of dispersal as a population-dynamic or epi-
demiological process.
A key question (3) is: does this matter, or is it suf-
ficient to model dispersal implicitly in exploring the
likely benefits of pest management actions?
Theoretical population ecological models predict
the effects of dispersal on the stability properties of
multiple species, especially biological control, inter-
actions and provide some clues as to general qualita-
tive outcomes.
A key question (4) is: are these predictions actually
testable, and can the theoretical models be comple-
mented/combined with biophysical models to enable
their use in specific pest management systems?
Dispersal research has matured as a biophysical sci-
ence, and is increasingly seen as crucial for the de-
velopment and implementation of IPM in agricultural,
horticultural and forest crops.
A key question (5) is: can the same approaches
and skills be used in applying dispersal research to
 M.J. Jeger / Agricultural and Forest Meteorology 97 (1999) 331–349
issues relating to biodiversity conservation and land-
scape ecology?
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