1 s2.0 S0304380007002414 Main

e c o l o g i c a l m o d e l l i n g 2 0 7 ( 2 0 0 7 ) 6184
available at www.sciencedirect.com
journal homepage: www.elsevier.com/locate/ecolmodel
European Corn Borer life stage model: Regional estimates of

pest development and spatial distribution under present
and future climate
a
b , D. Semeradov
M. Trnka a, , F. Muska
a a , M. Dubrovsky c , E. Kocmankov
a a , Z. Zalud
a Institute of Agrosystems and Bioclimatology, Mendel University of Agriculture and Forestry Brno, Zemedelska
1, 613 00 Brno,
Czech Republic
b State Phytosanitary Administration, Brno (SPA), Czech Republic
c Institute of Atmospheric Physics, Academy of Sciences of the Czech Republic, Prague (IAF), Czech Republic
a r t i c l e
i n f o
a b s t r a c t
Article history:
Predicting the potential distribution of agricultural pests, both indigenous and introduced,
Received 4 August 2006
plays a key role in determining the impact of global change on agricultural, horticultural and
Received in revised form
forestry ecosystems. This study investigates changes in the climatic niche of one of the most
2 March 2007
important agricultural pests, the European Corn Borer (Ostrinia nubilalis, Hubner), using the
Accepted 18 April 2007
multi-generational phenology model ECAMON. The model enables us to predict the develop-
Published on line 22 June 2007
ment of the European Corn Borer (ECB), to estimate the risk of its establishing a permanent
population, and to give an indication of climate-related stress factors affecting the species.
Keywords:
The evaluation of ECAMON demonstrated that it provides accurate predictions of the onset
Corn borer
and duration of the key phenological stages over a broad range of sites. It explains over 70%
ECAMON
of the variation in the timing of key developmental stages based only on daily weather data.
GCMs
ECAMON simulations correctly predicted the presence/absence of the ECB over the study
Degree day model
region during the 19611990 reference period. It also helped to explain the sudden increase
Climate change impacts
in the maize infestation over the territory of the Czech Republic during the unusually warm
period of 19912000. The ECAMON results demonstrated that the effect of climate will be
signicant and complex. According to our estimates, the extent of the climate niche will
expand within the next 2030 years to cover almost the entire area suitable for agriculture
by 20402075. The establishment of a bivoltine population is not imminent within the next
decade, but it is likely to take place during the period of 20252050. The timing and extent
of these changes will be affected not only by changes in the means of key meteorological
parameters, but also in their variability. These shifts will be clearly accompanied by an earlier onset of key developmental stages of the pest. The study demonstrated that the level
of uncertainty caused both by emission scenarios and by differences in global circulation
models (GCMs) are of the same order of magnitude. Thus, only the combination of a wide
range of emission scenarios and GCMs can provide insight into the potential effect of climate
change on any particular species. Under future climate conditions, grain maize is expected
to partly replace traditional cereals (e.g. winter wheat, rye, etc.); thus the establishment of
a national or international monitoring scheme is desirable, and an ECAMON-like tool might
serve as the basic modeling platform for such an effort.
2007 Elsevier B.V. All rights reserved.
elsk
Corresponding author at: Institute of Agrosystems and Bioclimatology, Mendel University of Agriculture and Forestry Brno, Zemed
a
1, 613 00 Brno, Czech Republic. Tel.: +420 6 0658 0394; fax: +420 5 4513 3083.
E-mail address: mirek trnka@yahoo.com (M. Trnka).
0304-3800/$ see front matter 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecolmodel.2007.04.014
62
1.
Introduction
Predicting the potential distribution of agricultural pests, both

indigenous and introduced, plays a key role in determining
the impact of global change on agricultural, horticultural,
and forestry ecosystems (e.g. Beaumont et al., 2005; Salinger
et al., 2005; or Gray, 2004). In the early 1990s, Porter et al.
(1991) listed possible impacts of increased global temperature
on the insect pests: (i) extension of geographical range; (ii)
increased over-wintering; (iii) changes in population growth
rates; (iv) increased number of generations; (v) extension of the
development season; (vi) changes in the croppest synchrony;
(vii) changes in interspecies interactions; (viii) increased risk
of invasion by migrant pests; (ix) introduction of alternative
hosts, green bridges and over-wintering hosts. Some of these
assumptions made 15 years ago have been conrmed by subsequent studies (e.g. Olfert and Weiss, 2006) and by observed
shifts in the ranges of crop pests throughout the world (e.g.
Rosenzweig et al., 2001). The speed with which pests are populating potential niches is primarily limited by the abundance
of host plants (e.g. agricultural crops or other host species),
their means of spreading, and by natural predators and/or parasites. Even non-athropogenic wind dispersal allows for the
progress of some insect species by tens of kilometers per year,
even in the rather complex terrain of Central Europe (Sefrov

a
and Lastuvka,
2001). The expansion of the agricultural pests
might be aided by the transfer of infested seeds, forage or other
products, and also by a dense matrix of elds with host plants.
The dynamics of an insect species invasion can be illustrated
by the recent example of the Western corn rootworm (Diabrotica virgifera) or Horse chestnut leaf miner (Cameraria ohridella
Desch & Dem). These cases showed that a vacant climatic
niche might be populated within a few years, providing that
there is a sufcient number of host plants in these regions.
It is quite possible that the direct gains in agriculture
production expected due to higher ambient CO2 concentra
tion (e.g. Parry et al., 2004; Trnka et al., 2004; Zalud
and
2002) could be nullied by the losses caused by
Dubrovsky,
pests (McCarthy et al., 2001; Cannon, 1998). Despite some studies indicating the decreasing importance of some pest species
(e.g. cereal aphids in Western Europe reported by Newman,
2005), many of them will benet by expansion into cooler areas
(e.g. Fuhrer, 2003; Patterson et al., 1999) and by an increased
number of generations (Porter et al., 1991). Even though adopting appropriate agricultural practices and technology might
help in controlling pest populations, such measures increase
the overall cost of production and add yet another pressure on
the environment (Chen and McCarl, 2001). In addition, some
of the techniques of pest control (e.g. deep ploughing) might
conict with efforts focused on soil moisture conservation
through minimum tillage systems. Similarly, the introduction
of genetically modied crops remains problematic in EU countries, and it brings a lot of challenges in its own right (e.g.
Gutierrez and Ponsard, 2006).
The ontogeny of poikilothermic insects is controlled
mainly by temperature (with other weather factors reducing
or enhancing survival rates), and this fact has been utilized
by agrometeorologists and phytopathologists for several generations. A wide variety of modeling techniques exist (e.g.
Guisan and Zimmermann, 2000) and are being applied in ecology studies (e.g. Logan et al., 2006; Beaumont et al., 2005) as
well as by farmers or consultant companies as part of expert
or warning systems (e.g. Aggarwal et al., 2006; Grunwald

et
al., 2000; Hijmans et al., 2000). Models of different complexities have also been used to determine the probability of a
particular pests establishment in a given locale, in case of
unintentional introductions of alien species (e.g. Morrison et
al., 2005; Gray, 2004; Rafoss and Sthre, 2003; Jarvis and Baker,
2001) or in case of a change in the voltinism of the present
populations (e.g. Yamamura and Yokozawa, 2002; Porter et al.,
1991).
Models that are able to estimate the spatial extent of a climatically suitable area for a particular pest allow us not only to
identify the species current potential distribution, but also to
assess which regions will be climatically suitable under future
climate scenarios. Despite some objections regarding the
exclusion of biotic interactions and dispersal scenarios (Davis
et al., 1998), these models have played a key role in assessing
potential distributions of species (Pearson et al., 2002). They
were also among the rst tools used to determine the risks
associated with the behavior of agricultural pests under climatic changes (e.g. Porter et al., 1991; Sutherst et al., 2000). Two
types of approaches have usually been adopted: (i) the inductive approach, which tries to nd links between species presence/absence and key climate variable(s) and (ii) the deductive
approach, which relies on more detailed insight into the
processes associated with the population establishment and
sustainability at a given site/region. The rst approach was
used, for example, in the climate matching mode of CLIMEX
(Sutherst and Maywald, 1985) or BIOCLIM (Houlder et al., 2000).
These tools proved to be quite efcient in cases where detailed
information about the species were not known, since eld surveys can be prohibitive due to the number of species or areas
involved and/or because only the presence-only type of data
are available. When reliable information exists, the deductive
approach (represented, for example, by phenological models)
is assumed to give a more accurate prediction of the probability of establishment than does climate matching (Gray, 2004).
This approach mostly relies on daily observed meteorological data, and thus accounts for weather variability on a ne
temporal scale. This is particularly important when relatively
small and heterogeneous regions are evaluated. In some cases
both approaches are combined, for example, in the CLIMEX,
compare location or compare year modes, to produce an
ecoclimatic index (EI), as a measure of the suitability of a location for establishment (Baker et al., 2000).
The study presented here is focused on one of the most
important corn (Zea mays L.) pests, the European Corn Borer
(Ostrinia nubilalis Hubner). It was chosen because of its nearly
global presence, adaptation potential, and its expected sensitivity to climate change within regions of Central Europe. The
European Corn Borer (ECB) population is presently univoltine
in most of Central Europe. In the southeast USA, it produces up
to four generations per year (Mason et al., 1996), and up to six
generation per year are possible under favorable conditions.1
http://www.cabicompendium.org/cpc/datasheet.asp?CCODE=
PYRUNU(COUNTRY=0), 2007.
According to some studies, the number of generations is connected with the particular pheromone types (so-called Z and
E strains), with the former being closely associated with corn;
whereas the E type is found on other crops (Mason et al.,
1996). The areas with two or more generations per season
suffer much higher economical losses, since the later generations of larvae tunnel through stalks, ear shanks, and
husks. This type of damage causes disruptions of assimilate
translocation throughout the plant, introduction of pathogens
(Keszthelyi and Lengyel, 2003; Gatch and Munkvold, 2002;
Magg et al., 2002), and increased plant lodging (Mason et al.,
1996). Presently, the only Central European region with bivoltine populations is in Hungary (Keszthelyi and Lengyel, 2003)
due to its favorable climatic conditions and host availability
during most of the seasons. This situation could be altered if
the climate conditions would permit the development of the
full second generation in the presently monovoltine areas. In
most of the cases maize is harvested from early September to
mid October, and thus the presence of host plants would not
be a limiting factor for the ECB population development. The
example of the North American population (which probably
originated from Hungary and/or Italy) demonstrated the possibility of rapid colonization of available niches and that the
switch from univoltine to bivoltine populations occur rather
quickly and could have serious economical consequences. The
voltinism of the ECB is not driven solely by the degree-day
accumulation; rather, according to some studies, it is inuenced by the day-length, scotophase (dened here as the part
of a night when the sun is less than 6 below the horizon), or
latitude (e.g. Onstad and Brewer, 1996). In the Czech Republic, the whole larval population enters diapause because the
current climatic conditions do not permit the development
of a complete second generation, even at the warmest sites.
An expected rise in air temperature will likely weaken factors
controlling the monovoltinism of the ECB population, since
a longer and warmer vegetation season might allow for the
completion of the second-generation life cycle. Based on the
historical analogy with the 1930s in the US, such a change
might occur rather suddenly (within two, three seasons) if
climate conditions are optimal.
In order to study the potential risks, a multi-generational
phenology model that accommodates known inuences of climate conditions on the ECB population was developed. The
main aims of this study were (i) to perform an evaluation of
the model using a wide range of eld observations over the
selected case study region (Fig. 1.); (ii) to determine the climatic
suitability of the case study region for the univoltine and bivoltine populations of the ECB under the present and expected
climatic conditions; and (iii) to examine the inuence of the
uncertainties inherent to the climate change scenarios on the
projections of future ECB populations.
2.
Material and methods
The structure of the Environmental Change Assessment

Model for Ostrinia Nubilalis (ECAMON) is presented in Fig. 2.
ECAMON is a semi-quantitative model that integrates known
relationships between environmental conditions and the
occurrence ECB into a single framework. It consists of phe-
63
Fig. 1 Overview of the case study area. (a) The set of

meteorological stations used for spatial analysis of the ECB
potential niche. The sites marked by circles (45 stations)
were used to develop multiple regression models for the
spatial analysis of ECB potential niche. The stations
marked by triangles (15 stations) were used for the
evaluation of the regression models. The altitudinal map
encaptures the main geographical barriers for ECB
populations dispersal whilst the thing gray lines indicate
district (NUTS4) borders. (b) The geographical locations and
distribution of sites with light or pheromone traps (marked
as black ags) and sites where the ECB eld occurrence was
monitored in correspondence survey (circles). The diameter
of the black dots indicates the length of the record. The gray
shading depicts the spatial distribution of arable land in
the study area.
nological and environmental stress modules and allows us to

map outputs using ArcInfo GIS software.
2.1.
Modeling of the ECB development
The initiation, 25, 50, and 75% development and termination

dates of each stage were estimated using a degree day model
64
Fig. 2 Scheme of the ECAMON model structure.
approach (DDM) that has been found to be as effective as

more sophisticated methods (Got et al., 1996). The base temperature (Tbase ) for the DDM was set at 10 C, as originally
proposed by Apple (1952) and recommended by numerous
other researchers (e.g. Mason et al., 1996; Porter et al., 1991; Got
and Rodolphe, 1989; Jarvis and Brindley, 1965), even though
some studies suggest higher temperature thresholds for at
1962). The degree days calculeast some stages (e.g. Brova,
lation is based on daily values of the minimum (Tmin ) and
maximum (Tmax ) air temperature (both at 2 m above ground)
and a recommended method: e.g. by the method of Arnold
(1960) and Mason et al. (1996):
If Tmax < Tbase then DD = 0
(1)
If Tmin < Tbase and Tmax > Tbase then DD

=
T
max
+ Tbase
2
Tbase
If Tmin > Tbase then DD =
(2)
T
max
+ Tmin
2
Tbase
(3)
The ECAMON model is initiated on the 1st November

in the preceding year; however, the degree day accumulation for a given season starts only after the snow cover has
melted completely and the Tmin does not drop below 0.2 C.
The snow cover presence is simulated according to Running
and Couglans (1988) simple snowpack model as evaluated by
Thornton et al. (2000). Since the rate of the ECB development
decreases when the temperatures reach the upper threshold,
we trimmed the Tmax values used for DD accumulation at
32 C. This value was set in the middle of the range reported

by Got et al. (1994, 1996), Calvin et al. (1991) and Anderson
et al. (1982). It was not possible to determine the rates of
development within the Central European population due to
the scarcity of the experimental data. Therefore, we used the
temperature thresholds dened by Browns2 degree-day model
(Brown, 1982) and Mason et al. (1996). The nal set of values is
listed in the Table 1. Even though the thresholds were based
on the populations from Kentucky, North Dakota, Missouri,
Delaware and Pennsylvania (USA), they agree rather well with
the available observations from Central Europe (e.g. Brova,

1996). The ECB development during
1962; Caga n and Barabas,
a given season is terminated whenever the Tmin drops below
0.2 C for 3 consecutive days. If such an event occurs up to
the stage of ight initiation, the simulation is resumed from
the beginning of the cycle, since there is a chance of some
pupae being still in cold resistant diapause. Our model considers that the later occurrence of low temperatures wipes out
the entire population of the pest unless at least part of the population has reached the 5th instar stage. The ECAMON design
includes the dependency between individual life-stages and,
thus, the proportion of population in a given stage cannot surpass the percentage that successfully nished development in
the previous stage.
The mechanism controlling ECB diapause was addressed in
two ways. The rst methods build on the Caga n (1998) study,
stating two conditions for the partial second generation in
Central Europe: (i) ECB larvae have to undergo development
up to the 4th instar under a daylight duration (sunrise to sun-
http://www.uky.edu/Ag/Entomology/entfacts/dcrops/ef106.
htm, 2007.
65
Table 1 Temperature thresholds of the ECAMON model based on the works of Brown (1982), Mason et al. (1996) and SPA
(1999)
Stage
Pupa
Adult
moths
Flight and
mating
Egg laying
Egg hatch/
1st instar
2nd instar
3rd instar
4th instar 5th instar
1st generation-ADD (accumulated degree days, C)

Initiation
121
216
288
50%
199
316
393
95%
293
404
482
321
454
532
388
516
588
447
599
654
512
677
732
585
754
810
710
827
882
2nd generation-ADD (accumulated degree days, C)

Initiation
782
882
904
50%
882
971
1066
95%
943
1060
1199
949
1154
1293
1050
1232
1366
1109
1299
1421
1174
1371
1493
1247
1443
1560
1372
1610
1927
set) of at least 15.5 h, and (ii) the Tmin in this period may not
decrease below 12 C. This study showed that, in both cases,
when the partial second generation was observed in the eld
conditions in Slovakia, the predened conditions were met.
The proposed length of the photoperiod seems to be rather
high in comparison with the North American ECB population
(e.g. Beck and Hanec, 1960; Beck and Apple, 1961 or Onstad
and Brewer, 1996). These authors reported that part of the
population did not enter diapause if it was reared under daylengths of at least 14.5 h. Therefore, we used two photoperiod,
14.5 and 15.5 h. The second method, proposed by Onstad and
Brewer (1996), allowed us to estimate the percentage of larvae
in diapause (D) based on the conditions from the 1st to the 4th
instar:
D = 384 + 30.8S + 2.33T + 5.11L
(4)
where L stands for latitude in decimal degrees, T for the air

temperature ( C) and S represents the scotophase length (h).
Scotophase is dened as the period when the sun is at least
6 below the horizon, and thus the civil twilight is considered
to be part of the photophase. According Onstad and Brewer
(1996), this method provides satisfactory results for all ECB
populations in North America between the latitudes of 30 and
50 N.
2.2.
Modeling of the environmental stress conditions
for ECB populations
A number of ECB eld studies showed that the uctuations
of the pest populations pertain to the over-wintering larvae
number (Chiang and Hodson, 1959; Chiang, 1961). It was then
argued that the size of the population during a given season
is independent of the egg laying and depends on the available feeding sites of the host. However, climatic conditions
during the season determine whether the available feeding
sites will be lled, how well the population during a given
season becomes established and what the extent of infestation will be. This explanation is supported by the analysis
of catches in light traps during 2000 and 2001 in the Czech
Republic. We noted that the economical losses caused by ECB
in these years were independent of the number of trapped
moths, but the ECB infestation pressure was higher during the
warm and dry episode in the spring of 2000. But weather factors still play a signicant role and in certain regions (e.g. Iowa,
USA), they alone explain 8193% of the population mortality in

the eld conditions (Showers et al., 1978) and they are consid 1959).
ered important in the Central Europe as well (e.g. Brova,
Beck and Hanec (1960) demonstrated that the over-wintering
5th instar requires rather high relative air humidity in order
to reach the pupal stage. The sensitivity to air moisture is
also characteristic of the latter developmental stages. Low air
humidity, for example, decreases the longevity of the moth
by 60% (Balachowsky and Mesnil, 1935), which directly inuences the occurrence of oviposition and the number of eggs
deposited per female. Showers et al. (1978) showed that the
ECB population can be reduced by the occurrence of drought
stress, especially when it is combined with high temperatures.
Similarly, Anderson et al. (1982) presented evidence that a low
moisture diet markedly slows down the ECB development. Boring larvae (i.e. later 3rd5th instar) tend to be less sensitive to
ambient moisture conditions, since they are protected by the
tissues of the host crop (e.g. Ellsworth et al., 1992). In order
to account for these factors, the ECAMON model includes the
ratio of the actual (ETa ) and crop reference (ETc ) evapotranspiration as an indicator of the water stress imposed on the
hosts and on the ECB population itself. The PenmanMonteith
model (Allen et al., 1998) is considered to be a reliable method
for estimating the reference evapotranspiration (ETr ), which
is dened as the evapotranspiration from well-watered grass.
The ETc value describes the evapotranspiration of a wellwatered canopy (e.g. grain maize) and could be estimated from
ETr using the dual crop coefcient approach (e.g. Allen et al.,
1998). The ETa brings together ETc and the available soil moisture on a given day. We considered days with the ETa to ETc
ratio below 0.4 to be dry, and those below 0.2 to be extremely
dry. When the ratio equaled 1.0, the days were marked as
wet.
The developmental stages from the moth ight up to the
2nd instar of the ECB are sensitive not only to drought, but
also to heavy precipitation. During this period, eggs or larvae
feeding on leaf surfaces can be washed onto the soil, which
sharply increases mortality. ECAMON incorporated this effect
by counting the number of days with precipitation over 20 mm
during sensitive stages. The number of the droughts and wet
and rain stress days are summed for each stage to allow for
comparisons between sites or seasons.
Special attention was paid to the air humidity during the
period from oviposition to egg hatching; Godfrey and Holtzer
(1991) demonstrated that the effect of temperature and mois-
66
ture stress accumulates during stressful periods, and remains

even after the stress is relieved (e.g. during the night). They
also showed that survival rates could be as low as 1.9% under
a combination of high air temperature and low relative air
humidity. The overall decrease of egg hatchability was determined using Godfrey and Holtzers (1991) formula:
y = a(ebT )zc
(5)
where y is the percentage of eggs that hatch, T the mean air

temperature ( C), z the vapor pressure decit (mb), and a, b and
c are constants. The ECAMON environmental stress module
serves as a semi-quantitative indication of the climate niche
characteristics. We are aware of the potential drawbacks of the
methodology, including the lack of experimental data quantifying the effect of the stress day number on the population
density of ECB. One can also question the suitability of the
chosen stress threshold levels for the eld populations in Central Europe. Nevertheless, it is, in our view, useful to make a
comparison between the levels of known stress factors during
the critical developmental stages, not only between sites, but
also between different climate change scenarios. This part of
the study also points out areas where more research on the
ECB life cycle and its interaction with weather parameters is
needed.
2.3.
Spatial analysis of the ECB characteristics
In order to carry out a spatial analysis of the ECB climatic niche

for the case study region, a generalization of ECAMON outputs was required. Since the density of the weather stations
with long-term and complete datasets was low, we turned to
the approach used in similar studies, e.g. by Gray (2004) or
` and Sharov (1999). The phenological model was run

Regni
ere
for individual weather stations, and selected model outputs
were interpolated. The observed weather series of 19612000
were extended by ve randomly chosen years that were used
for model initialization at each site. The primary assumption
preceding every run was that the ECB might eventually survive at any of the stations examined. We then evaluated the
outputs of the ECAMON model for all years and stations and
discarded those where the given developmental stage of the
pest was not found. Multiple linear regression models (MLR)
were consequently tted to each dataset using altitude, northing, and easting (as well as their rst order interactions
and squares) as covariates of individual ECAMON outputs (e.g.
mean date of the ight initiation). The MLRs were developed
using the statistical package UNISTAT v. 5.1.11. The performance of each MLR was evaluated by a comparison of the
original and interpolated values at 45 weather stations by
means of a coefcient of determination (R2 ), mean bias error
(MBE), and root mean square error (RMSE). Fifteen additional
weather stations (Fig. 1a) were used to independently verify
interpolated values with those calculated by ECAMON with
the station data. In the last step, the appropriate MLR was coupled with the digital landscape model (DLM) with 1 km 1 km
grids, and nal maps or table outputs were produced. The
DLM also included, in addition to coordinates and altitude,
information on the crop suitability, soil conditions, and land
use; thus it enabled us to exclude all but the grids of arable
soil suitable for maize production. The MLR method produced

results similar to those of other interpolation techniques, e.g.
co-krigging or thin spline, despite a relatively low density of
weather stations (i.e. one station per approximately 1800 km2 ).
Even though applying method proposed by Jarvis et al. (2003)
would seem to be more consistent in methodological terms,
our results suggest that the MLR method does not decrease
precision and that it is not computationally intensive. In addition, it prevents occurrence of non-homogeneities between
meteorological parameters that would likely appear due to the
simultaneous use of multiple meteorological variables and low
station density.
2.4.
Weather input data
ECAMON requires daily values of Tmax and Tmin , daily sums

of global radiation and precipitation, as well as the mean
water vapor pressure and wind speed. The complete series
were available from 45 stations of the Czech Hydrometeorological Institute (CHMI) and included 40 years of complete
data, from 1961 through 2000 (Fig. 1a). This set of stations is
likely to be applied for the operational version of the model
in the near future since they belong mostly to the rural sites
with good data quality. An additional 15 sites used for verication of the MLR performance (Fig. 1a) had, in some cases,
up to 3 years of missing data that were reconstructed by CHMI
experts based on observations from at least 10 surrounding
stations. In all cases, the station location and measurement
procedures followed the WMO standards. The spatial distribution of the basic set of 45 stations (Fig. 1a) indicates that only
two stations are located higher than 800 m a.s.l., which is an
upper boundary for any sensible agriculture production, both
in present and future climates. The lowest station is located at
171 m above the sea level (with a mean annual temperature of
9.3 C and mean annual sum of precipitation 481 mm during
the 19612000 period). The station with the highest altitude
is at 1324 m (with a mean annual temperature of 2.6 C and
annual precipitation 1391 mm). The mean altitude of all of the
stations is 448 m, which is close to the country mean altitude
(430 m). The mean annual temperatures and annual sum of
precipitation of the selected weather stations are almost identical to the mean climatological values of the Czech Republic,
i.e. 7.5 C and 674 mm, respectively. For evaluation of the ECAMON at the light trap sites, data from more than 100 weather
stations across the region were available.
2.5.
Experimental data for model testing
The authors carried out an overview of the relevant regional

literature and gathered all available information on the ECB
occurrence in the Czech Republic. The rst data source
included dates and the number of caught moths at the light
and pheromone traps (Fig. 1b) during the period from 1977 to
2005. These data enabled us to validate the initiation, 50%,
and the end of the ECB ight scenarios simulated by ECAMON. They were complemented with the published data from
the light traps and data on oviposition from Slovakia (Fig. 1b).
The second data source included reports on the ECB presence/absence in grain corn elds across the Czech Republic
that were published by the State Phytosanitary Adminis-
tration (SPA) between 1961 and 2003. We also conducted a

correspondence survey. The survey was conducted among
the farmers, breeders, and researchers, and the combined
database presently includes over 4400 reliable reports of the
pest presence/absence on corn from over 230 sites (Fig. 1b). To
overcome the insufcient data on bivoltine populations and
to allow for ECAMON validation, we used experimental data
published by Jarvis and Brindley (1965). This dataset was combined with daily weather data for the Boone weather station
in Iowa, USA (42.06N, 93.88W and 353 m a.s.l.) between years
1950 and 1961.
The agreement of the ECAMON outputs, relative to the
observed data, was assessed by several statistical methods.
These included a comparison of the basic descriptive statistics, e.g. mean, median, extreme values and coefcient of
variance, as well as further examination by the mean bias
error (MBE), root mean squared error (RMSE), Pearson correlation coefcient (r), modeling efciency index (MEI) according
to Wilmot (1982) and Theils inequality coefcient (U) (Theil et
al., 1970). The main reason for reporting our results in terms
of several statistical values was to allow comparison with
the results of other authors and at the same time to enable
other researchers to make the same comparison. Since some
of these methods cannot be compared directly (e.g. MEI and
U), we prefer to provide values for each of them. MEI is a very
useful indicator that is specically derived in order to assess
the performance of models compared with the experimental
data; it is formulated as follows:
n
(J
i=1 est
n
MEI = 1
|J
i=1 est
Jobs )
Jestmean ||Jest Jobsmean |

(6)
where Jobs and Jest are, e.g. the observed and estimated dates
of month ight initiation. The values have to be totaled for
each possible pair of estimated and observed values. The index
results in a number between 0 and 1. The closer the number
is to 1, the better the t between the model and eld observations. MEI refers to the accuracy of prediction, where accuracy
is regarded as the degree to which model predictions approach
the magnitude of their observed counterparts.
Theils inequality coefcient (U) penalizes large errors more
than small ones and it also assesses a models ability to
duplicate a turning point or rapid changes in the data. It is
formulated as follows:

U=
1/n
1/n
(Jest Jobs )
2 +
Jest
1/n
2
Jobs
2.6.
Determination of the ECB climatic niche using
ECAMON
The outputs of ECAMON include key dates of phenological
development and the outputs of the stress modules, both for
individual seasons and for the statistics for selected number of
years. These outputs could be evaluated in a number of ways
including the effect of the interannual climate variability on
the population or the population stability. Furthermore, multiple indices (e.g. Jarvis and Baker, 2001) could be developed
according the requirements of the user. Since we intend to
explore a number of future climate scenarios, we decided to
use the climate suitability index (CS) that gives rather comprehensive picture about the ECB year-on-year stability. The index
for univoltine population (CSI ) is derived from the proportion of seasons during which the ECB successfully completes
one-generation life cycle. The establishment of a viable ECB
population requires the stability of environmental conditions.
Therefore, the calculation of CSI was considered only at sites
that allowed, in all years, at least 5% of the ECB population
to develop. A particular year was marked as suitable when it
enabled the development of 95% of the population, and the
value of CSI was calculated as:
CSI =
(7)
The value of U ranges from 0, indicating perfect t, to 1,

indicating a total absence of any t between the simulated
and observed values. One of the main advantages of the coefcient is the possibility of calculating proportions of estimated
bias (UB ), variance (US ) and covariance (UC ). Whilst UB indicates systematic error, US measures the ability of the model
to replicate the degree of variability in the data. Any remaining
error after accounting for the bias and variance effects is called
covariance proportion (UC ). The ideal distribution of inequality over these three sources is for the bias and variance effects
to equal 0 and the covariance to equal 1 (Theil et al., 1970).
Xsuitable
n
(8)
where Xsuitable represents the number of suitable seasons

based on the criteria dened above, and n is the total number of years. CSI therefore represents the proportion of years
during which the entire univoltine population successfully
completes its development. Those sites that allowed for at
least 5% of the second generation to complete its development were considered climatically suitable for the bivoltine
ECB population (CSII ). A lower threshold for CSII was chosen
to make ECAMON more sensitive to the onset of the bivoltine
ECB population. Sites with CSI or CSII values higher than 0.70
(i.e. requirements fullled in 7 seasons out of 10) are referred
to as primary niche areas that are characterized by the highest infestation pressure of ECB. The regions with CSI < 0.55 are
thought to be unsuitable for the long-term establishment of
a viable univoltine ECB population. An occasional occurrence
of the ECB in these otherwise marginal regions might still be
possible, especially during warm years, and at patches close
to the areas with an existing ECB population.
2.7.
67
GCM-based climate change scenarios
The climate change scenarios applied in this paper are based

on the transient simulations made by four global circulation
models (GCMs), which were performed for the IPCCs Third
Assessment Report (Houghton et al., 2001). These models,
whose outputs were downloaded from the IPCC Data Distribution Center,3 included CSIRO-Mk2, ECHAM4, HadCM3 and
NCAR-PCM. The simulations were run at the SRES-A2 emission scenario. All GCMs included in the analysis are models
coupled with ocean circulation. The horizontal resolution of
the atmospheric part of the model ranges from 2.8 to 7.5 in
the zonal direction, and from 2.25 to 5.6 in the meridional
http://cera-www.dkrz.de/IPCC DDC/SRES/index.html, 2007.
68
Fig. 3 The standardized GCM-based climate change

scenarios for the Czech Republic: (a) daily mean
temperature and (b) precipitation. The GCMs are:
CSIRO-Mk2, ECHAM4/OPYC3, HadCM3, and NCAR-PCM. Y is
related to the changes in the annual means. The changes
are with respect to thebaseline period (19611990).
direction. The details of the models may be found on the

IPCCs web page (http://www.mad.zmaw.de/IPCC DDC/html/
SRES TAR/index.html). The localized climate change scenarios were constructed by the pattern scaling technique of
Santer et al. (1990): the scenario is dened by a product of the
standardized scenario (Fig. 3) and the change of the global
mean temperature. The standardized scenarios, which relate
responses of climatic characteristics to a 1 K rise in the global
mean temperature (TG ), were determined from a 20102099
period of the above-mentioned GCM runs. Changes in TG for
time periods (2010, 2015, 2020, 2025, 2030, 2040, 2050 and 2075)
were calculated by a simple climate model MAGICC (Harvey

et al., 1997; Hulme et al., 2000), assuming combinations of
an emission scenario and climate sensitivity (equilibrium
change in global mean surface temperature following a
doubling of the atmospheric equivalent CO2 concentration,
TG,2CO2 ) given in Table 2. Finally, the standardized scenario
from each GCM was scaled by the values of TG (Table 2),
which represent low, middle, and high estimates of the
global mean temperature rise. The low estimate is based
on low climate sensitivity (TG,2CO2 = 1.5 K) and the SRES-B1
emission scenario. It is a rather optimistic scenario, assuming
a convergent world that puts an emphasis on global solutions to economic, social and environmental sustainability.
The upper estimate is based on a high climate sensitivity
(TG,2CO2 = 4.5 K) and an SRES-A2 emission scenario, which
assumes a very heterogeneous world and a primarily regionally oriented economic development. The middle estimate of
TG is based on a combination of the SRES-A1T emission scenario, which represents the conservative estimate of future
emission developments, with a medium climate sensitivity
(TG,2CO2 = 2.5 K). More details on the SRES (The Special Report
on Emission Scenarios) scenarios may be found in Houghton
et al. (2001), and the details (including the model validation)
on constructing GCM-based climate change scenarios for
the Czech Republic are in Dubrovsky et al. (2005). Since the
GCM-based outputs are only in the monthly time step and
have a very low spatial resolution (one grid box usually covers
the entire Czech Republic), the further statistical downscaling
for individual sites was necessary. The stochastic weather
1997) was initially trained
generator Met&Roll (Dubrovsky,
on the 40 year series of observed data from each weather
station. The parameters of the generator (e.g. monthly mean
temperature) were then changed according to the appropriate
GCM-based climate change scenario and the daily series of
synthetic weather data were generated for each site. More
details about this downscaling technique can be found in
Dubrovsky and Zalud

(2002) and Dubrovsky et al. (2004).
The climate change scenarios used in this study do not
consider changes in the variability of climatic characteristics because the applied outputs of the GCM models included
only monthly series that did not allow us to reliably estimate
changes in the daily weather variability. This might be a serious limitation, considering the concerns raised that changes
in climatic variability might have a greater impact than do
changes in means of climatic variables (e.g. Mearns et al.,
1997). Therefore, a sensitivity analysis was performed as a part
of the study to demonstrate the potential effect of temperature
variability on an ECB climate niche. The analysis was carried
Table 2 Changes in the global mean temperature ( C) for three combinations of emission scenarios and climate
sensitivity used in the study
Scenario
2010
2015
2020
2025
2030
2040
2050
2075
A2-HIGH
A1T-MED
B1-LOW
+0.63
+0.47
+0.32
+0.77
+0.59
+0.37
+0.93
+0.71
+0.43
+1.10
+0.85
+0.49
+1.29
+0.98
+0.55
+1.67
+1.24
+0.65
+2.08
+1.47
+0.76
+3.13
+1.92
+1.02
The changes are with respect to the baseline period (19611990) and were calculated by MAGICC with only the effect of greenhouse gasses
considered (no effect of atmospheric aerosols is taken into account).
out for a combination of a SRES-A2 scenario, high climate sensitivity, and an ECHAM GCM. The mean values of individual
weather elements were modied according to the appropriate
scenario in each time period. The standard deviation param 1997),
eters of the weather generator, Met&Roll (Dubrovsky,
were modied in such a way as to imply a reduction/increase
of the temperature variability by 25, 50 and 100% (compared to
the present climatic conditions). One of the major drawbacks
of the weather generator use is its tendency to underestimate the number and intensity of extreme events, such as
extreme temperatures or the duration of heat waves (Kysely
2005). But according to our experience, the
and Dubrovsky,
stochastic weather series rarely exert a signicant inuence
over general tendencies in the results of complex models,
e.g. crop or hydrological (Dubrovsky et al., 2005). In this case,
the additional tests using both observed and generated data
showed small differences between key ECAMON outputs. In
fact, the mean difference between the ight initiation was
less than 0.5 day; whereas the values of CSI obtained by these
two methods for the 19611990 period at 45 weather stations
were virtually identical (P = 0.993). Finally, it should be noted
that the uncertainty introduced by various SRES scenarios and
GCM models is more than one order of magnitude higher than
uncertainties caused by the used downscaling method.
3.
Results
3.1.
Evaluation of the ECAMON
69
The ECAMON explains over 73% of the variability in moth

ight initiation dates (Fig. 4a) with negligible bias (Fig. 4a), but
it tends to estimate higher developmental rates in the ECAMON population for latter stages of ECB (Fig. 4b and Table 3).
The model correctly depicts interseasonal variability of ECB
development (Fig. 4a and c and Table 3), as well as differences between sites, with values of r, MEI and Uc very close
to 1 (indicating good t of the model to observed data) and
relatively small RMSE (Fig. 4b and Table 3). Also, oviposition
initiation is simulated rather well (Fig. 4b and Table 3), but
in this case only ve data points were available. The ECAMON estimated the ight and oviposition dates of the rst
and second generation for Boone County in Iowa, US (Jarvis
and Brindley, 1965) with the similar MBE, MEI and Uc values
as with the Czech data (Table 3 and Fig. 4b). The diapause
modules were tested in addition on datasets published for two
locations by Caga n (1998), Caga n et al. (2000) and Tancik and
Caga n (2004) in Slovakia and Poland, where ndings of empty
pupae substantiated the presence of the partial second gen-
Fig. 4 results of the ECAMON model evaluation: (a) observed vs. estimated ight initiation at Central European locations
(n = 61); (b) values of MBE and RMSE of the estimated ight and oviposition dates across sites in Central Europe and U.S.; (c)
comparison of the observed and simulated development dynamic during extremely warm (2003) and normal years (2005) at
the experimental site in Kunovice.
70
eration in 1994. At both sites, the ECAMON results were in

agreement with observations and indicated the presence of
a partial second generation (i.e. some non-diapauzing larvae)
in 1994, but not in other years during 19931996 experimental
period.
Modeling efciency index (Wilmot, 1982) indicates the goodness-of-t of the predicted values. The value of the coefcient lies between 0 and 1, with 1 indicating perfect t.
Theils inequality coefcient tests the goodness-of-t of the predicted values. The value of the coefcient lies between 0 and 1, with 0 indicating perfect t.
3.2.
Mean standard deviation, minimum and maximum dates (expressed as Julian days) are given for each stage. The relationship between observed and estimated series is assessed by means of the
Pearson correlation coefcient (r), modeling efciency index (MEI) and Theils inequality coefcients (U) including individual components (see Eq. (7) for details).
0.71
0.77
0.80
0.78
0.11
0.21
0.15
0.03
0.18
0.02
0.05
0.09
0.62
0.53
0.61
0.59
165
168
217
220
162
167
215
226
140
143
196
199
141
153
200
206
7
7
7
8
158
161
208
212
5
4
5
7
12
11
11
11
Data from boone, Iowa, USA
1st generation ight: 5% of population
1st generation oviposition: 5% of population
2nd generation ight: 5% of population
2nd generation oviposition: 5% of population
155
160
207
213
0.99
0.99
0.92
0.63
0.86
0.85
0.68
0.71
201
206
227
166
210
205
237
170
146
158
170
183
137
163
172
189
11
12
13
7
168
182
192
177
11
10
13
10
61
54
53
5
Data from Central Europe
1st generation oviposition: 5% of population
165
186
212
181
Estimated
Observed
Estimated
Observed
Estimated
Observed
0.98
0.98
0.99
0.99
0.02
0.02
0.01
0.01
0.78
0.63
0.78
0.47
0.01
0.01
0.01
0.12
0.21
0.36
0.21
0.41
0.02
0.02
0.05
0.09
UC
US
UB
U-components
Ub
MEIa
r
Max (JD)
Min (JD)
Mean S.D. (JD)
n
Developmental stage
Table 3 Comparison of the observed data and estimates based on the ECAMON for sites in the Central Europe (Czech Republic and Slovakia) and in Boone station
(Iowa, USA)
ECB climatic niche during the period 19612000
The mapping of the CSI for the period from 1961 to 1990
showed that all the locations where the occurrence of ECB on
maize was reported were limited to the areas with CSI > 0.55
(Fig. 5a). In total, 77 sites with long-term ECB observations
comprising of 411 ECB seasonal damage reports were available. Among them, 60 sites (i.e. 383 ECB reports) were located
in the area with CSI > 0.70 and the remainder in the adjacent
region with CSI > 0.55. Unfortunately, these site-based observations allow only for a qualitative assessment of the ECAMON
performance, since they did not cover the whole territory of
the country evenly.
To carry out a partial quantitative assessment of the ECAMON results. We determined those districts (Fig. 1a) where
CSI > 0.70 at least in some areas, and these were then considered to be the districts with the ECB presence. This estimated
ECB presence/absence in each district was then compared
with the SPA reports that were based on various numbers
of observational points at which key agricultural pests were
monitored during the 19611990 period. Since the location of
these points was changed annually and were not mapped,
they could not be used for more detailed model evaluation.
The comparison of observed and estimated values showed
agreement in 71 out of 77 districts, with only one case of
false negative ECAMON prediction. The unweighted coefcient kappa value (Cohen, 1960) equaled 0.74, indicating
relatively a good t of the model. As shown at Fig. 5a, the
bulk of the ECB population during 19611990 was concentrated
in the southeast of the country. Two sites in the northwest
reported an ECB infestation of grain maize only during the
unusually warm season of 1988. The economically signicant
losses and areas treated by ECB-specic insecticides were limited exclusively to the southeastern region (Fig. 5b). However,
the northwest prime niche has shown signs of ECB pres
ence since the 19th century (e.g. Zima and Zavorka,
1938),
but only as a pest of other crop species (especially Hop
vine, Humulus lupulus), with no signicant damage to grain
maize.
The quality and extent of the ECB niche has been signicantly altered during 1990s (Fig. 5b and d). During this period,
the mean annual temperature at a number of representative
Czech weather stations increased by 0.20.9 C as compared
n,
2001). At the same time,
to the 19611990 baseline (Kveto
92 out of 100 of the highest monthly mean temperatures were
recorded during the 19912000 period, as compared to 8 during
the 19611990 baseline. The modeled shift of climatically suitable areas as indicated by the model has been conrmed by an
enormous increase in the number of reports of ECB occurrence
on maize in the northwest of the country that was associated
with unprecedented level of crop damage (Fig. 5d). For the rst
time, ECB was observed in the northeast of the country as
well. Out of 169 sites where ECB presence was reported (i.e.
370 seasonal presence reports), 141 sites (i.e. 338 reports) were
located in the area with CSI > 0.70 and 19 sites (i.e. 23 reports)
in adjacent region with CSI between 0.55 and 0.70. The fact
that the remaining nine sites are situated in the area with CSI
between 0.40 and 0.54 could be explained by the short period
of time for which CSI was determined and/or as an indication
of continuing expansion of the current niche. The unweighted
71
coefcient kappa value equaled to 0.64, which still indicates a

rather good t between the modeled and observed data especially when we consider the non-stationarity of the population
during 1990s.
There is also a close relationship between the mapped
CSI values and the number of reported infestations, which
Fig. 5 (a and b) Value of the CSI and CSII during 19611990 (a) and 19912000 (b), respectively. The dots represent sites
where the ECB occurrence in maize was observed under the eld conditions. (c and d) Areas where ECB causes economic
losses expressed as a proportion of years when the ECB treatment in the region was reported during the periods of 1961 (c)
and 19912000 (d). Notes: For better visualization both (a) and (b) and Fig. 7af include the whole territory of the country
(excluding areas above 800 m a.s.l.) rather than the arable land only. The resolution of (c) and (d) depends on the size of the
reporting units, i.e. districts with mean area of 1000 km2 .
72
Fig. 5 (Continued ).
document the reliability of ECAMON and its development

in time. The positive effect of higher temperatures during
19912000 on the ECB climatic niche was partially offset by
an increased number of drought stress days from oviposition
until the second larval instar, despite an earlier initiation of
these stages. The egg-hatching module indicated, on average,
a 612% decrease in hatchability during the 19912000 period,
as compared to the reference period that could result in a
higher mortality.
3.3.
niche
Impact of the climate change on the ECB climatic
The climate change will lead to an earlier beginning of the

growing season and will accelerate pest development (Fig. 6a).
By 2025, the ight initiation could take place, on average, 410
days earlier than during the reference period, and the ECB life
cycle will be completed 915 days earlier. Differences exist
between the scenarios, with some ECAMON runs predicting
73
Fig. 6 (a) Example of the life cycle of ECB under present climatic conditions and changed climatic conditions in 2025 and
2050 (A2 emission scenario, high climate sensitivity and ECHAM model) at the Lednice station (171 m a.s.l.; 16.83E; 48.78N).
Each bar represents mean dates when individual stages are initiated and 95% of the population nished the given stage.
The vertical lines delimit the period of the year with a day-length (sunrisesunset) of 14.515.5 h, respectively. (b) Example
sowing timing and the key developmental stages of grain maize at the Lednice station under present and changed climatic
conditions (as in a) estimated by the CERES-Maize model. Legend: (1) From sowing to emergence, (2) from emergence to the
end of juvenile phase, (3) from the end of juvenile phase to the oral initiation, (4) from the oral initiation to the end of leaf
growth, (5) from the end of leaf growth to the beginning of grain lling, and (6) the grain lling phase.
the completion of the rst generation more than 1 month earlier by 2050, as compared to the reference period. Since it has
been already shown for the example of the 19912000 period
(Fig. 5), the increase in air temperatures leads to a larger climate niche for ECB (Fig. 7af). This will be accompanied by
a spread of the pest into these regions. As the differences
between the values for different emission scenarios in Table 4
illustrate, the uncertainty in the suitable area is substantial
and increases with time. A combination of B1 emission scenario, low climate sensitivity, and NCARPCM model assumes
only a small temperature increase and an enlargement of the
ECB prime niche area in 2010 by only a 4.0% compared to
19611990 period. According this scenario, the prime niche
area would increase to 31.0% of all arable land by 2025, and
up to 43.0% by 2050. Such development is not probable, since
the rate of green house gas emissions assumed by the SRESB1 scenario is very low. The mid-range estimate (based on
SRES-A1T emission scenario, medium climate sensitivity, and
the HadCM model) would result in the expansion of ECB to
62% of arable land by 2025. Almost all agricultural land would
allow for the existence of an univoltine ECB population by 2050
(Table 4). If we consider the possibility of a rapidly progressing
climatic change (i.e. SRES-A2 emission scenario, high climate
sensitivity, and the ECHAM model), then the entire arable land
area would be potentially suitable for ECB between 2025 and
2030 (Fig. 7e and Table 4).
The probability of an occurrence of the partial second
generation inside two main maize production areas (i.e. the
southeast and northwest of the country) increases significantly after 2025 (Fig. 6a). The permanent presence of a
74
partial second generation might cause higher economical

losses since some of the larvae could damage stems and husks
during grain lling. ECB is known to feed on a number of other
plant species (e.g. Nault et al., 2001) that eventually might
have a higher nutritional value or water content than maize.
In such case higher rates of development should be expected
1984 or Anderson et al., 1982) and, in warm years,
(e.g. Brova,
such change of diet could enable development of the fth ECB
instar before the rst frosts and the bivoltine life cycle could be
established. If we assume no change of the host preferences by

the ECB, the simulations based on the SRES-B1 emission scenario, low climate sensitivity, and the NCARPCM model show
extremely low probability of a bivoltine population establishment, even prior to 2075 (Table 4). A combination of an SRES-A2
emission scenario and high climate sensitivity with HadCM,
ECHAM and NCARPCM GCMs would on the contrary enable
the existence of a stable bivoltine population in the southeast
between 2025 and 2050 (Table 5b). According to these scenar-
Fig. 7 Development of the climatically suitable areas for the 1st and 2nd generations expressed in terms of CSI and CSII
during 2025 and 2050. The estimates are based on the ECHAM GCM model with a combination of the B1 SRES scenario and
the low sensitivity of a climate system (ECHAM LOW), A1T SRES scenario and medium climate sensitivity (ECHAM MED) and
A2 SRES scenario and high climate sensitivity (ECHAM HIGH).
ios, by 2050, two generations of the pest will be common in

the northwest of the country, with the southeast population
area reaching CSII > 0.90 (Fig. 7e and f; Table 5b).
In order to assess possible changes in synchronization
between the pest and host (i.e. grain maize) development, we
estimated the expected duration of phenological stages using
crop growth model CERES-Maize (Jones and Kiniry, 1986). This
model was extensively calibrated by Zalud

and Dubrovsky
(2002) and results indicate relatively small shifts in mutual
timing of ECB and maize stages (Fig. 6b). Even though the
crop model predicts shortening of some maize developmental
stages (especially from owering to maturity) under climate
75
change, we have to take into account the possibility of earlier

sowing dates, which will mitigate the effect of higher overall
temperatures on the shortening of maize development. The
results show that a large part of the second ECB generation
will manage to complete its life cycle before the harvest (Fig. 6)
in most years.
3.4.
Effects of a changed climate on the number of
stress events
The changing climate conditions and consequent increase in
the availability of suitable areas will affect other environmen-
Fig. 7 (Continued )
76
Fig. 7 (Continued ).
tal variables that inuence ECB. An increase in temperature

will lead to earlier ight periods as well as to the earlier
completion of the development (Figs. 6a and 8a). An example
of the experimental site in Lednice shows that, by 2025, the
mean ight period will be 410 days earlier and, by 2050, this
shift would be as large as 18 days on average (Fig. 8a). Under
present conditions, the individuals complete their rst generation developmental cycle in early August (Fig. 5a), whereas by
2025 it is likely to be completed during the last 10 days of July.
According to some scenarios, by 2050, the development of the
rst generation could be completed by the 10th of July. The

number of drought and wet stress days for the ECB population
is not expected to change signicantly for the rst generation
as we expected that the maize will be sown earlier. However,
the second generation will be faced with a much higher water
decit and number of dry days than the present univoltine
population. Model results also indicate signicant changes
in the egg hatchability (Fig. 8b) due to higher water vapor
pressure decit. In some years, hatchability might decrease
to below 50%, thus reducing the infestation pressure. The
77
Table 4 Changes of the relative area of arable land suitable for one (CSI ) and two (CSII ) generations of ECB under present
and changed climates
SRES-scenario GCM
2010
2015
2020
2025
2030
2040
2050
2075
Proportion of arable land with CSI > 0.7

B1-LOW NCARPCM
0.18
A1T-MED HadCM3
0.39
A2-HIGH ECHAM
0.51
0.22
0.45
0.70
0.26
0.56
0.82
0.31
0.62
0.95
0.35
0.77
0.98
0.37
0.89
1.00
0.43
0.98
1.00
0.56
0.99
1.00
Proportion of arable land with CSII > 0.7

B1-LOW NCARPCM
A1T-MED HadCM3
A2-HIGH ECHAM
0.02
0.04
0.24
0.03
0.38
0.24
0.99
A2-HIGH stands for SRES-A2 scenario and high sensitivity of the climatic system; A1T-MED stands for SRES-A1T scenario with medium climate
sensitivity and B1-LOW stands for SRES-B1 scenario with a low climate sensitivity to the increase of green house gases. The individual emission
scenarios were used as boundary conditions for the several GCM model.
number of drought stress days will increase, especially during

the summer months, and will inuence the egg hatchability
and mating capability of moths, especially those from the
second generation. Since the drought stress will also affect
the host plants, the effect of increased environmental stress
on ECB population will not likely benet the farmers in
the end.
3.5.
Role of climatic variability under expected climate
conditions
The effect of the climate variability on the ECB proved to be
complex and highly dependent on the degree of change of the
mean values of key weather variables. As is apparent from
the results for the Lednice station (Fig. 8c and d), the relative
Fig. 8 Analysis of the ECB climatic niche characteristics under baseline conditions and expected climates. (a) The date of
the rst generation ight initiation; (b) The date of 1st generation development completion; (c) Percentage of hatched eggs.
For construction of (a) and (b), the same set of scenarios as in Fig. 5 were used. The (c) and (d) are based on the combination
of A2 emission scenario, high climate sensitivity, and the ECHAM model. Note: the black ends of each bar at (ac) represent
minimum and maximum values, the white area delimits 1 standard deviation from the mean value, which is marked by
the vertical black line.
78
impact of the increased climate variability decreases with the

increase of the mean values. This explains why the effect of
variability is lower in 2050 compared to 2025 (Fig. 8c). A lower
variability results in a more uniform timing of development
(Fig. 8c) and more stable populations (Fig. 8d). The doubling of
the weather variability leads to a high population instability,
even at presently suitable sites (Fig. 8d). The change of variability inuenced the egg hatchability and number of stress
days, with some years showing a decrease in egg hatchability below 30%. The results of the Wilcoxon test showed that
alterations of weather variability in the range of 12.5% will
not cause a statistically signicant alteration of the ECB population for any of the climate change scenarios. A change of
temperature variability by 25% or more would lead to a signicant change in the distributions of the selected parameters
(i.e. ight initiation, egg hatchability, and number of stress
days).
4.
Discussion
The rst study considering the potential impact of climate

change on the ECB climate niche in the Central Europe
was performed by Porter et al. (1991). The authors applied
a very simple approach based on the accumulated DD per
one generation and climate change scenarios available at
that time with rather coarse resolution. The methodology
used was not veried with experimental data from Central
Europe and did not take into account the effect of interannual variability or other meteorological elements besides the
temperature. The ECAMON not only allows detailed insight
into the ECB life stage timing, but it also introduces a simple
and efcient DD accumulation initiation procedure, using a
combination of Tmin temperature threshold and snow cover
presence, which ensures more reliable results, at least in
the Central Europe compared to those used by Mason et
al. (1996) or Jarvis and Baker (2001). As is frequently the
case for similar studies (e.g. Logan et al., 2006 or Gray,
2004), available eld data on ECB mortality are scarce; thus
we had to deal with the fact that only a few phenological
stages are well monitored in Central European ECB populations compared to other regions (e.g. Got and Rodolphe,
1989 or Phoofolo et al., 2001). The lack of reliable experimental data was substituted by using a semi-independent
module that provides the user with at least a notion of the
changes in abiotic stresses relative to the present state. This
approach is in line with the ndings of Roy et al. (2001),
who used an extensive U.K. buttery database to develop and
test quantitative models for prediction of buttery populations; they found such models to be reliable as qualitative
but not quantitative tools. The experience of the ECAMON
development shows that the key threshold value could be
transferred across different populations, which is in accordance with the ndings of the US-based study of Calvin et al.
(1991), who found only small differences between geographically isolated populations in terms of their developmental
rates. Despite overall satisfaction with the ECAMON performance, there are several points that need to be discussed
further.
4.1.
Methods of degree day accumulation and spatial
analysis
During evaluation of the ECAMON at individual sites, we noted
a bias in the key phenological dates at some sites (e.g. Fig. 4).
These could be explained by (i) differences in trap sensitivity
associated with their location in the terrain or (ii) by the distance to the nearest weather station. The latter explanation
is supported by distinct patterns in the model residuals versus trap to weather station distance assessed by pattern index
(Donatelli et al., 2004). Yamamura and Yokozawa (2002) suggest that the ambient air temperature at 2 m over a standard
grass surface is not a good proxy for estimating temperature
of some poikilothermic species since global radiation can lead
to an increase in body temperature well above the temperature of the environment. Our results indicate that this effect
does not have to be considered in the case of ECB. This is partly
due to the biology of the pest, since its life cycle takes place
on the bottom side of leaves, in the whorl, deep inside the
stem or, in the case of adult moth, during the night; thus, the
role of solar radiation is relatively small. Moreover, the ECAMON showed a tendency to predict the onset and especially
the termination of the developmental stages earlier than was
indicated by the light traps (Fig. 4; Table 3), which in a way
contradicts the signicance of global radiation to ECB body
temperature. These faster than observed developmental rates
could be caused by three factors, individually, or in combination: (i) either by the fact that at least part of the observed
catches belong to populations feeding on other hosts, which
might lead to lower rates of development (Anderson et al.,
1982); (ii) by the difference between air temperature and crop
temperature or (iii) because of the presence of two ECB strains.
It is possible that ECAMON performance could be improved
by the use of an estimated canopy temperature for later ECB
stages, rather than the one of ambient air. Such algorithms are
available, e.g. as a part of complex crop growth models (e.g.
Brisson et al., 2003) and will be the subject of further research,
since they require detail parameterization of the crop model at
a given site. Probably the most likely explanation of the slower
development of the ECB population than predicted by ECAMON is the presence of both ECB strains in the area. Mason et
al. (1996) reported higher developmental rates for the E strain
than for the Z strain. According to Hrdy et al. (1986), both Z and
E strains are present in the southeast region, where most of
the validation data were collected and it is very likely that both
strains were caught in the light traps. This then explains the
higher variability in the observed developmental rates of Central European populations compared to ECAMON estimates,
which were based mostly on E strain specic data.
The evaluation of the MLR performance using an independent set of 15 stations (Fig. 1a) showed that the CSI values
for 19611990 and 19912000 were estimated with MBE lower
than 0.02 and RMSE between 0.08 and 0.11. At the same time
the values of r 0.92 and 0.97 indicated a very good t between
the interpolated data and those estimated at the station level.
Similar results were achieved with the interpolated values
of mean ight initiation date (MBE < 0.4 days and RMSE < 3.2
`
days), which is similar to values reported by Regni
ere
and
Sharov (1999). The highest magnitude of error was found in the
northeastern lowlands, which are characterized by a colder
and wetter climate compared to the rest of the country. The

overall performance of the MLR technique is satisfactory and
it seems that, even with a relatively low station density, rather
good nation-wide estimates could be derived, which is crucial
for the ECAMONs application as a monitoring tool.
4.2.
Historical and future patterns of ECB infestation
pressure
The use of ECAMON enabled to elucidate the reasons behind
the sharp increase of pest incidence and crop damage during the 1990s. The ECB prominence in recent years has been
mostly attributed to the general increase in the grain maize
acreage, an overall decrease of crop protection, the widespread
use of the minimum tillage technologies, and a general decline
of the quality of farming practices (e.g. Daems et al., 2006).
These changes correlated with social and economical reforms,
started in 1989, which led to a decade-long crisis in the agriculture sector. However, we are convinced that these factors
alone cannot explain the spread of ECB that caused damage
in 40 new districts, its establishment in completely new areas
and even at altitudes over 600 m above the sea level (Fig. 6b).
Even though we lack experimental data that would allow the
analysis similar to Yamamura et al. (2006), the ECAMON results
clearly show that the underlying cause of the increase in the
ECB infestation was the major increase in the size and quality of the prime niche area (Fig. 6b) compared to 19601990
(Fig. 6a). In 19611990, only 8.1% of arable was found suitable by ECAMON for ECB population (CSI > 0.7); whereas in
19912000, it was 17.3%. This doubling of the climatically suitable area was found to be directly proportionate to the sharp
increase in the number of spots where ECB-caused crop damage was observed (Fig. 6b and d) during the 1990s. Against the
conclusions of Daems et al. (2006), also stands the fact that
the overall area of the grain maize acreage oscillated between
29 and 40,000 ha during the 1990s, with maxima in 1990 and
2000. During the same period the total maize growing area
decreased by 40% (from 426 to 267,000 ha) in the same time due
to decreasing demand for silage maize. Furthermore, the ECB
infestation intensity is still growing, even after the consolidation of the agriculture sector in late 1990s, despite much higher
attention was being paid to plant protection and tillage practices. During 20012006, the ECB occurrence was documented
from sites above 700 m a.s.l. during 2006. Analysis of this particular season by ECAMON showed that in 2006 the unusually
warm weather in July enabled rapid and homogenous development of the ECB population, and, that at least part of the
ECB population could survive there in 2 preceding years.
ECAMON results also showed the presence of the partial
second generation occurrence (although only at the warmest
sites) during 1983, 1993 and 2000 by at least one of two available diapause models. In all 3 years the development of the
larval instars was initiated about 2 weeks earlier than in an
average year. In 1993 and 2000, the ECB developmental season started 20 days earlier due to an unusually warm spring
whilst in case of 1983 the high June temperatures speeded up
the development of the latter ECB stages. The existence of the
partial second generation has been proposed by Czech experts
since the 1980s (e.g. Dusek, 1983). Yet these claims were based
only on indirect signs, which included secondary peaks during
79
the moth ight, partially bivoltine populations in the adjacent

1998; Barabas
et al.,1985) and the
regions of Slovakia (Caga n,
experiment by Brova (1984), who showed that larvae kept in
near eld conditions on energy rich diets will not enter diapause, but will establish a partial second generation in Central
European conditions. The reason why it has been so difcult
to prove partial second generation in the Czech Republic is
easily explained by a very low ratio of non-diapauzing ver 1998)
sus diapauzing larvae, which is less than 1:2000 (Caga n,
and probably by the sheer low frequency of occurrence, as
ECAMON results suggest.
The evaluation of ECAMON uncovered a certain degree
of uncertainty in our understanding to the diapause control
factors. There are certain signs that in Central Europe the
diapause is not primarily controlled by the length of photoperiod as Caga n (1998) proposed. It is clear that in the
southeastern region even the requirement for at least a 15.5 h
day-length during the development of 1st to 4th instar is fullled in number of the years (Fig. 6a). Thus, the onset of
diapause in the Caga n (1998) algorithm is in fact controlled
by the proposed threshold of Tmin = 12 C under which all larvae will enter diapause. It is a well-established fact that many
insect species adjust their life cycles to the seasonal phenology by latitudinally changing the photoperiodic response of
diapause (Danilevsky et al., 1970; Taylor and Spalding, 1986)
and that the photoperiodic response is rather easily changed
by the process of local evolution. Studies from North American ECB populations indicate that shorter day requirements
(e.g. 14.5 or 14.75 h) are associated with higher air temperatures and lower latitude (Beck and Apple, 1961); whereas
others point out that at least part of civil twilight should be
included into the photoperiod (e.g. Onstad and Brewer, 1996).
Thus, the length of the critical photoperiod for the Central
European population might be longer than that proposed by
Caga n or the temperature has larger effect on diapause than
day-length. This was the main reason why we applied the
method of Onstad and Brewer (1996), originally developed for
North American population, because it includes the effect of
day-length (including civil twilight), temperature and latitude.
However, there are obvious limitations in the transferability of
this method to Central Europe: (i) it is highly sensitive to small
changes of latitude above 50 N and (ii) there is a known disparity between the latitudinal temperature gradient between
geographically corresponding locations in Central Europe and
North America. Thus, studies similar to Beck and Apple (1961)
and Onstad and Brewer (1996) with the ECB population from
Central Europe should be performed and a model specic for
the region should be developed. However, neither photoperiod nor temperature seems to be limiting bivoltine population
establishment in the southeast of the Czech Republic under
the changed climate, as can be seen from Fig. 6a or Fig. 7c, e
and f. We believe that the results of ECAMON are sufciently
accurate and in accordance with the experimental data.
Despite caveats, the ECAMON helps us to gain insight into
the present status and the possible changes of the ECB prime
niche under future climatic conditions. The use of a large
number of scenarios (compared to similar studies, e.g. Gray,
2004 or Beaumont et al., 2005) also allowed us to demonstrate that the uncertainty in the phenological model could
be marginal compared to other sources of uncertainties. The
80
Table 5a Changes to the relative area of arable land suitable for one (CSI ) and two (CSII ) generations of ECB according to
different emission scenarios
Scenario
B1-LOW
A1T-MED
A2-HIGH
2010
2025
2050
2010
2025
2050
0.36
0.41
0.51
0.44
0.60
0.95
0.57
0.99
1.00
0.02
0.02
0.38
A2-HIGH stands for SRES-A2 scenario and high sensitivity of the climatic system; A1T-MED stands for SRES-A1T scenario with medium climate
sensitivity and B1-LOW stands for SRES-B1 scenario with a low climate sensitivity to the increase of green house gases. The individual emission
scenarios were used as boundary conditions for the ECHAM GCM model.
Table 5b As (a), but using a single emission scenario SRES-A2, high climate sensitivity, and four different GCM models
GCM
ECHAM
NCARPCM
HadCM
CSIRO
2010
2025
2050
2010
2025
2050
0.51
0.37
0.53
0.56
0.95
0.61
0.96
0.82
1.00
0.98
1.00
0.99
0.02
0.38
0.05
0.36
0.42
overall uncertainty of the estimate for ECB prime niche area

is high and the main factor affecting the outcome of the analysis is which emission scenario is combined with which GCM
and climate sensitivity assumption. The rate of expansion
of the prime niche would be the slowest according to the
SRES-B1-based projections, which is half that of the SRESA2-based estimates (Table 5a). Differences between various
GCMS (Table 5b) are the second major contributing factor to
the uncertainty. The results indicate that estimates based on
the NCARPCM model yield a signicantly slower expansion of
the climate niche area than do the other three GCMs tested
(Table 5b). The uncertainty caused by GCMs (at least in case
of SRES-A2 emission scenario) is more pronounced in the near
future (i.e. time horizons of 2010 and 2025). In the long-term
(e.g. 2050 or 2075), the uncertainty is attributable mostly to
emission scenarios (Table 4). We expect that by 20302040, at
least part of the present maize growing areas will be endangered by the second ECB generation, with the rst outbreaks
occurring about a decade earlier. In the southeast of the Czech
Republic the probability of the bivoltine populations establishment is higher, and might take place 510 years sooner than
in the northwest of the country. In order for a ECB population
to complete two generations, the ECAMON model indicates
that the second generation ight period would have to begin
during mid-August at the latest (Fig. 6a). Therefore, ndings
of pupal remains in the beginning of August or any notable
peaks in trap catches or ndings of newly laid eggs during the
rst two decades of August should serve as a warning of an
impending 2nd generation. Similarly, seasons with the ight
initiation of the rst generation prior the 5th of June might be
potentially suitable for the establishment of the second generation. An early warning system based on a combination of
eld observations and models is capable of providing farmers
with an early warning that might help in limiting economical
loses. The potential risk of the second ECB generation can be
partially mitigated by the introduction of genetically modied
maize cultivars, particularly Bt-varieties (Daems et al., 2006).

However, their introduction poses a dilemma for the growers, not only due to higher cost of the seeds, but also due to
an existing opposition by consumers. Other mitigation strategies that focus on targeting various ECB stages (e.g. removal of
the plant residues followed by deep ploughing or an increase
of pesticide treatments) would increase production costs. An
abandonment of minimum tillage strategies might limit the
possibility of soil moisture conservation and lead to higher
soil compaction. The occurrence of a bivoltine ECB population might also lead to a much higher infestation of the forage
maize. Under present conditions, the occurrence of the ECB
larvae is not economically important, since the forage maize
is usually harvested earlier, i.e. before the ECB larvae could
cause damage to the husks or stems. We expect that the shifts
in the ECB developmental rates and more abundant ECB population will result in a higher exposure of forage maize to the
pest. This would inevitably inuence both forage production
and quality and could lead to signicant economical losses,
since forage maize acreage is ten times higher than that of
grain maize.
5.
Conclusions
The ECAMON model is a multi-generational phenology model

that predicts the development of the ECB, estimates the risk of
establishment, and gives an indication of the climate-related
stress factors. It requires no user input regarding initiation
dates and uses no arbitrary cut-off date. The predictions
of CS and phenological events are a function of the temperature regime during the given generation, and depend on the
completion of a life cycle during the preceding generation.
For input, the model requires records of daily maximum and
minimum temperatures, global radiation, precipitation, wind
speed, and air humidity. The evaluation of the model showed
81
that it produces accurate estimates of the onset and duration

of the key phenological stages over a broad range of sites.
At the same time, we demonstrated that the climate suitability determined with the help of ECAMON corresponds to
the observed ECB occurrence. ECAMON simulations helped
to explain the sudden increase of the ECB caused damage
over the territory of the Czech Republic during the unusually warm period of 19912000. This makes ECAMON an ideal
tool for testing the effect of future climate change on the
ECB population within a region of interest. We demonstrated
that the effect of climate will be signicant and complex.
According to our estimates, the niche suitable for univoltine
populations will increase rapidly within next 2030 years and
will cover almost the entire area suitable for agriculture by
20402075 (depending on the scenario). The prospect of the
bivoltine population becoming established is not imminent
within the next decade, but it is likely to take place during the
20252050 period. The timing and extent of these changes will
be affected not only by changes in the means of key meteorological parameters, but also in their variability. These shifts
will be accompanied by an earlier onset of key developmental
stages of the pest. The change in developmental patterns of
maize (including earlier sowing dates) might lead to a different
interaction between the ECB and host plants. Despite a higher
frequency of unfavorable conditions in some years, the ECAMON does not indicate a signicant increase in environmental
stress for the rst generation because of the earlier onset of the
development. The effect of environmental stress (especially
droughts) on the population of the second ECB generation
remains to be seen since there are no experimental data available in the region. We expect that the lower egg hatchability
and a lack of moisture will partly limit the second generation
development, especially when the increased weather variability is taken into account. The uncertainty in the emission
scenario, climate sensitivity to increases of greenhouse gases,
and differences between individual GCMs are of the same
order of magnitude and represent the main sources of uncertainty in estimating the future ECB potential niche. Thus, only
a combination of wide range of emission scenarios and GCMs
can provide insight into the potential effect of climate changes
on any particular species. Under future climate a larger area
will allow for successful grain maize production and the area
of host plants will likely increase. Grain maize is also expected
to partly replace traditional cereals (e.g. winter wheat, rye,
etc.), especially in the most productive areas. It seems that the
introduction of a national (or preferably international) monitoring scheme is desirable, and ECAMON could serve as the
basic modeling platform for such effort.
Acknowledgements
The development of the ECAMON model was funded through
the National Agency for the Agricultural Research (project
QG60051) as a part of the vulnerability assessment of the main
agrosystems; whereas the methodology of the spatial assessment of climate change impact on pest species was developed
and tested with support of project funded by the Grant Agency
of the Czech Republic (Project No. 522/05/0125). The support
of Dr. Trnka and Dr. Dubrovsky by 6th FP EU project CECILIA
(project no. GOCE 037005) is appreciated. We are grateful to Dr.

Jrgensen and to three anonymous reviewers for their inspiring comments and suggestions.
Appendix A. List of symbols and abbreviations

The following list includes all symbols, constants and coefcients that are used in the paper. In case of adopted parts of
the ECAMON the nomenclature of the paper tries to follow the
original symbols used by the authors.
Symbol
a
b
c
CSI
CSII
D
DD
ETa
ETc
Jest
Jobs
L
MBE
MEI
r
R2
RMSE
S
T
Tbase
Tmax
Tmin
U
UB
UC
US
y
z
Equationa
Description
Empirical coefcient-Godfrey
and Holtzer (1991)
and Holtzer (1991)
and Holtzer (1991)
Climate suitability index for
univoltine population
Climate suitability index for
bivoltine populations
Percentage of larvae in
diapause (%)
Degree day ( C)
Actual evapotranspiration
(mm)
Crop reference
evapotranspiration (mm)
Estimated values
Observed values
Latitude (decimal degrees)
Mean bias error
Modeling efciency index
(Wilmot, 1982)
Pearson correlation coefcient
Coefcient of determination
Root mean square error
Scotophase (day of the night
without civil twilight) (h)
Mean air temperature ( C)
The base temperature of
ECB > 10 C ( C)
Maximum daily air
temperature ( C)
Minimum daily air
temperature ( C)
Theils inequality coefcient
Theil coefcient proportion of
estimated bias
estimated covariance
estimated variance
Egg hatchability (%)
Vapor pressure decit (mb)
(5)
(5)
(5)
(8)
(8)
(4)
(6) and (7)

(6) and (7)
(6)
(4)
(1)(3)
(1)(3)
(2) and (3)
(7)
(5)
(5)
82
Global circulation models and SRES scenarios-related

abbreviations
CSIRO
GCM
ECHAM
HadCM
NCAR
SRES
SRES-A1T
SRES-A2
SRES-B1
GCM made by Australian

Commonwealth Scientic and
Research Organization (version
Mk 2 was used in the study)
Global circulation model
GCM made by Max Planck
Institute for Meteorology and
the German Climate
Computing Centre (version 4.
was used in the study)
GCM made by the UK Met
Ofce Hadley Centre (version
Mk 3 was used in the study)
GCM made by National Centre
for Climate Research USA (PCM
version was used in the study)
Special report on emission
scenarios (refers to the source
of the emission scenarios that
used to drive GCMs), http://
www.grida.no/climate/ipcc/emission/
Emission scenario A1T as used
in the Houghton et al. (2001)
Emission scenario A2 as used
Emission scenario B1 as used
a Equation
number in this column is given only if the parameter is used in the specic equation.
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e c o l o g i c a l m o d e l l i n g 2 0 7 ( 2 0 0 7 ) 6184

journal homepage: www.elsevier.com/locate/ecolmodel

European Corn Borer life stage model: Regional estimates of

Received 4 August 2006

Received in revised form

Accepted 18 April 2007

Published on line 22 June 2007

Degree day model

Climate change impacts

Predicting the potential distribution of agricultural pests, both

even in the rather complex terrain of Central Europe (Sefrov

or warning systems (e.g. Aggarwal et al., 2006; Grunwald

Material and methods

The structure of the Environmental Change Assessment

Fig. 1 Overview of the case study area. (a) The set of

nological and environmental stress modules and allows us to

Modeling of the ECB development

The initiation, 25, 50, and 75% development and termination

Fig. 2 Scheme of the ECAMON model structure.

approach (DDM) that has been found to be as effective as

If Tmin < Tbase and Tmax > Tbase then DD

If Tmin > Tbase then DD =

The ECAMON model is initiated on the 1st November

32 C. This value was set in the middle of the range reported

the available observations from Central Europe (e.g. Brova,

4th instar 5th instar

1st generation-ADD (accumulated degree days, C)

2nd generation-ADD (accumulated degree days, C)

where L stands for latitude in decimal degrees, T for the air

USA), they alone explain 8193% of the population mortality in

ture stress accumulates during stressful periods, and remains

where y is the percentage of eggs that hatch, T the mean air

Spatial analysis of the ECB characteristics

In order to carry out a spatial analysis of the ECB climatic niche

` and Sharov (1999). The phenological model was run

soil suitable for maize production. The MLR method produced

Weather input data

ECAMON requires daily values of Tmax and Tmin , daily sums

Experimental data for model testing

The authors carried out an overview of the relevant regional

tration (SPA) between 1961 and 2003. We also conducted a

Jestmean ||Jest Jobsmean |

The value of U ranges from 0, indicating perfect t, to 1,

where Xsuitable represents the number of suitable seasons

GCM-based climate change scenarios

The climate change scenarios applied in this paper are based

http://cera-www.dkrz.de/IPCC DDC/SRES/index.html, 2007.

Fig. 3 The standardized GCM-based climate change

direction. The details of the models may be found on the

were calculated by a simple climate model MAGICC (Harvey

Dubrovsky and Zalud

Evaluation of the ECAMON

The ECAMON explains over 73% of the variability in moth

eration in 1994. At both sites, the ECAMON results were in

ECB climatic niche during the period 19612000

coefcient kappa value equaled to 0.64, which still indicates a

document the reliability of ECAMON and its development

Impact of the climate change on the ECB climatic

The climate change will lead to an earlier beginning of the

partial second generation might cause higher economical

established. If we assume no change of the host preferences by

ios, by 2050, two generations of the pest will be common in

model was extensively calibrated by Zalud