Crop Protection 30 (2011) 1586e1593

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Crop Protection
journal homepage: www.elsevier.com/locate/cropro

A new model for predicting the ight activity of Lobesia botrana

(Lepidoptera: Tortricidae)
Mariano Amo-Salasa, *, Vanesa Ortega-Lpezb, Radoslav Harmanc, Agustn Alonso-Gonzlezd
a
Department of Mathematics, University of Castilla-La Mancha, Camino de Moledores s/n, 13071 Ciudad Real, Spain
b
Department of Civil Engineering, University of Burgos, Calle Villadiego s/n, 09001 Burgos, Spain
c
Department of Applied Mathematics and Statistics, Comenius University, Mlynsk dolina 842 48, Bratislava 4, Slovakia
d
Regulatory Council for the Designation of Origin (D.O.) Ribera del Duero, Calle Hospital, 6 09300 Roa, Burgos, Spain

a r t i c l e i n f o a b s t r a c t

Article history: The grape vine moth, Lobesia botrana (Denis and Schiffermller) (Lepidoptera: Tortricidae), attacks
Received 13 February 2011 vineyards mostly in Southern Europe and Northern Africa. The efciency of most control methods
Received in revised form depends on the treatment of pest populations at their most susceptible stages, therefore the prediction of
2 September 2011
the moths development cycle would help greatly in determining an optimal treatment schedule. One
Accepted 4 September 2011
strategy for protecting against this pest is based on the day of maximum ight of males per generation
(peak ight), when the mating between males and females and the laying of eggs reaches its maximum.
Keywords:
The knowledge of these times would enable us to decide the kind of insecticide and the best time to
Model
Touzeau model
target it. The time of maximum ight can be predicted by the Touzeau model of accumulated average
Ribera del Duero daily temperature exceeding the development threshold. As shown, the t of this model on data from the
Logistic function Ribera del Duero region in Spain is not accurate enough. Therefore, we propose an empirical and bio-
logically plausible model based on logistic transformation of daily temperatures, which proves a better t
and permits simple inclusion of additional variables such as relative humidity.
2011 Elsevier Ltd. All rights reserved.

1. Introduction berries; the third generation (September) affects mature grapes

Wine is an important product in Spain and the Ribera del Duero
region is one of the most important wine growing zones, covering
an area of 20,945 ha. The grape vine moth, Lobesia botrana (Denis
and Schiffermller, 1776), a microlepidoptera member of family the
Tortricidae, is one of the most important pests that attacks vine-
yards in the region. It is a polyvoltine species with three or four
generations per year depending on the latitude. The number of
generations is mainly modulated by the temperature and photo-
period but other factors including relative humidity also have an
important inuence on its development. Temperature and photo-
period operate on the growth rate and the induction of diapause,
respectively. L. botrana passes through eight development stages in
its cycle: egg, ve larval stages, pupa and adult.
In the Ribera del Duero region, the rst generation occurs in
MayeJune and appears during the ower bud stage of its host
(phenological stage H), when it damages the inorescences (Baggiolini,
1952). The second generation (July) is observed during the closing of
the grape clusters (phenological stage L) and it damages the green
* Corresponding author. Tel.: 34 926 295300x6843; fax: 34 926 295361.
E-mail address: mariano.amo@uclm.es (M. Amo-Salas).
(phenological stage N) which can be seriously damaged (Ortega-Lpez
and Alonso-Gonzlez, 2008; Vassiliou, 2009; Armendriz et al., 2010).
The maturity stage of grape berries exerts an inuence on the devel-
opment of L. botrana (Savopoulou-Soultani et al., 1999). The species
remains dormant as a pupa from SeptembereOctober until the
following spring. Fig. 1 shows the life cycle of L. botrana through the
year in the Ribera del Duero region. The inhibition of diapause is
decisively inuenced by the average temperature at the end of winter
and the beginning of spring (Gabel and Roehrich, 1990; Coscoll, 1997).
A rapid development of the larva can ensure that the third generation
does not enter dormancy. The development speed of the rst genera-
tion is the origin of a partial or total fourth generation, which becomes
almost completely lost before winter. This fact could be attributable to
global warming (Martn-Vertedor et al., 2010).
The efciency of the control methods (mating disruption with
pheromones and chemical treatments) depends on the treatment
of pest populations at their most susceptible stages, the prediction
of the moths development cycle would therefore greatly help in
determining an optimal treatment schedule (Moravie et al., 2006).
Pheromone traps provide useful information on male moth activity
in vineyards, but by themselves, do not provide a reliable basis for
timing control methods. However, together with the method of

0261-2194/$ e see front matter 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cropro.2011.09.003
 M. Amo-Salas et al. / Crop Protection 30 (2011) 1586e1593
Fig. 1. Diagram of the life cycle of L. botrana during a typical growing season in
Northern Spain (Feytaud, 1924; Baillod and Baggiolini, 1993).
temperature summation, trap samples are important components
of phenological models (Riedl et al., 1976).
It is widely known that models based on daily temperatures are
used to forecast the phenological development of insects, in addition,
predictive methods have been developed to forecast the appearance
of L. botrana in the eld and to monitor its ight activity (Touzeau,
1981; Wagner et al., 1984; Gabel and Mocko, 1986; Baumgrtner
and Baronio, 1988; Caffarelli and Vita, 1988; Logan, 1990; Worner,
1992; Briolini et al., 1997; Milonas et al., 2001; Moravie et al., 2006;
Aylaj and Noussair, 2010), mainly in Central Europe where the moth
has from one to three generations per year (Gabel and Mocko, 1984).
The Touzeau model, for instance, is a prediction model of the
development of L. botrana based on accumulated average daily
temperature exceeding an established threshold. This model is
currently used to predict the stages of the moth in the Ribera del
Duero region. The moths ight activity is also monitored using
pheromone traps and the catch data are used to make comparisons
with the predictions of the Touzeau model.
It is shown in this paper that the t of the Touzeau model in the
Ribera del Duero region is not accurate enough. This could be due to
diverse factors such as the different conditions where the model
was developed and the zone where it is applied. This study aims to
provide a starting point to improve the predictions. A new
approach based on the Touzeau model was developed in order to
improve the t between the actual data available and the predic-
tions of ight activity of male L. botrana.
Firstly, the data available and the monitoring methods are pre-
sented followed by a detailed description of the Touzeau model.
The ts between the predictions of the Touzeau model and actual
data of the moths monitoring are then shown. Finally, a new model
is suggested and new ts displayed.
2. Materials and methods
2.1. Data from pheromone traps
Cardboard delta traps with sticky inserts baited with synthetic
sex pheromone (ECONEX-E,Z-7,9-dodecadienil acetate 1 mg) were
1587
used to monitor the ight activity of male L. botrana. Traps were
hung at a height of 1e1.5 m above ground level. In all vineyards,
traps were hung before the rst ight (MarcheApril) and were
maintained to the end of the season (usually October). Traps were
checked once a week. Sticky inserts were changed as needed and
the pheromone lures were refreshed once every 6 weeks. The data
from trap catches over the years provided information on the
duration of each generation under eld conditions.
Our pheromone trap catch data were collected throughout the
years 2004e2010, from 21 vineyards situated in the Ribera del
Duero region, attending to different parameters (geographical
situation, importance of vineyards in the zone, vineyard age and
their topographic altitudes). Table 1 shows the references, loca-
tions, coordinates, altitudes, grape varieties, ages of plantation,
systems of cultivation and irrigation of these vineyards.
Two traps were placed in each of the vineyards, consequently,
the information obtained corresponded to the catches in each trap,
the average of both traps and the average per trap per day (PTD),
that is, the ratio between the average of both traps and the number
of days between two inspections.
Using this information, ight curves, which express the captures
per trap and day throughout the year, were plotted. This method-
ology enabled the estimation of the actual time of maximum ight
in each generation, once these maximum ights had taken place;
therefore a method to predict these times was necessary. At the
same time, a phenological culture study was carried out, as well as
weather monitoring.
2.2. Weather data
The weather data were collected over the years 2004e2010 and
provided by the Junta de Castilla y Len, Local Government.
Nowadays, the information is collected from 6 weather stations
distributed throughout the area. The stations provide daily
maximum, minimum and average temperatures and readings of
these and relative humidity, solar radiation, precipitation and wind
speed, at 30-min intervals.
Each vineyard which was studied, was assigned to a weather
station following closeness criterion. The largest and smallest
distances between a studied plot and its corresponding weather
station were 25 km and 0.5 km, respectively. Table 2 shows the
locations, coordinates and altitudes of these stations.
In 2004, 2005 there were only three weather stations working.
Moreover, ve vineyards were removed from the study in 2005
because the data obtained from these sites did not provide addi-
tional information (Pelez et al., 2004). The data from 2004 to 2008
were used to compute the parameters of the new model and those
from 2009 to 2010 to check it.
2.3. Touzeau model
In 1981, J. Touzeau (Touzeau, 1981) studied the life cycle of
L. botrana and designed a prediction model. This model is based on
a function which relates the day-to-day development of the moth
to the average daily temperatures. The values that quantify the
degree of development are accumulated only if the average daily
temperature exceeded 10  C. This value establishes the lower
developmental threshold but this model does not establish a higher
one. The Touzeau function is mathematically expressed by:
TTou T maxf0; T  10g (1)
where T is the real variable standing for the average temperature.
The cumulative value of the degree of development on day k of
the year is:
1588 M. Amo-Salas et al. / Crop Protection 30 (2011) 1586e1593

Table 1
References, locations, coordinates, altitudes, grape varieties, ages of plantation, systems of cultivation and irrigation of the monitored vineyards.

Ref. Municipality Coordinates UTM. Altitude Variety Age of Plantation System Irrigation
Datum: European 1950 (m) (years) of cultivation
P30 Anguix X: 30T 421911 Y:4623037 835 TP 24 Trained Yes
P13 Aranda de Duero X: 30T 443687 Y:4612331 798 TP 15 Trained Yes
P17 Aranda de Duero X: 30T 439406 Y:4611394 834 TP 17 Trained No
P18 Aranda de Duero X: 30T 439035 Y:4611751 820 CS 21 Trained No
P9 Castillejo Robledo X: 30T 458568 Y:4604862 985 TP 17 Trained No
P25 Curiel de Duero X: 30T 409581 Y:4607888 754 TP 16 Trained No
P26 Fuentelcsped X: 30T 447186 Y:4605935 851 TP 38 Bush No
P1 Gumiel de Mercado X: 30T 432320 Y:4616534 808 CS 22 Trained Yes
P2 Gumiel de Mercado X: 30T 432573 Y:4616065 803 TP 12 Trained Yes
P31 La Horra X: 30T 427736 Y:4623251 828 TP 26 Trained and Bush No
P19 Olivares de Duero X: 30T 382148 Y:4609757 751 TP 15 Trained No
P27 Pedrosa de Duero X: 30T 418771 Y:4618325 817 TP 32 Trained No
P34 Peael-Aldeayuso X: 30T 405920 Y:4603198 757 TP 14 Trained No
P11 Pearanda de Duero X: 30T 459750 Y:4611855 878 TP 64 Bush No
P23 Pesquera de Duero X: 30T 401488 Y:4610947 771 TP 19 Bush No
P29 Quintana del Pidio X: 30T 437823 Y:4624067 865 TP 70/10 Bush/Trained No
P15 Roa X: 30T 423462 Y:4617407 784 TP 11 Trained Yes
P32 San Martn de Rubiales X: 30T 418692 Y:4611032 803 TP 11 Trained Yes
P14 Sotillo de la Ribera X: 30T 430867 Y:4625776 875 TP 49 Bush No
P7Q Valbuena de Duero- Quintanilla X: 30T 391615 Y:4609234 735 TP 52 Trained No
P33 Valbuena de Duero X: 30T 390251 Y:4611038 749 TP 13 Trained Yes

TP. Tempranillo CS. Cabernet Sauvignon.

X
k what is considered to be a suitable prediction. This could be dened
Sk s fTou Td (2) as a precise t between predicted days of maximum ight and
d1 actual ones. Therefore, the error is the addition of the absolute
values of the intervals between the predictions of maximum and
where s T1 ; T2 ; .; Tk is the sequence of average daily temper-
the actual days of maximum ight,
atures recorded by the corresponding weather station from 1
January, Td is the average temperature on day d of the year. Fig. 2 X
shows the daily accumulation of temperature with respect to the Err ~y;p;g j
jky;p;g  k (3)
average temperature according to Equation (1). Touzeau estab- y;p;g

lished thresholds which display the generation and the stage of the where ky;p;g are the actual (observed) dates of maximum ight of
development cycle of the moth with respect to the cumulative the year y, on the plot p, of the generation g and ~
ky;p;g are the cor-
value. responding predictions.
Table 3 shows the different stages of the development of The basic building block of the mathematical model considered
L. botrana, their durations and the daily accumulated temperatures in this study was the logistic function given by the formula
necessary to achieve them, expressed in degree-days (DD > 10  C),
according to Touzeau (Touzeau, 1981). Each developmental stage of 1
L. botrana has its own total heat requirement. Development can be f1 Tja; b (4)
1 eabT
estimated by accumulating degree-days between the temperature
thresholds throughout the season. The accumulated degree-days where T is a real argument and a, b are real parameters. In the
from a starting point can help to predict when a developmental function, the value f1 (Tja, b) quanties in intervals of [0,1] the
stage will be reached. Nevertheless, only the maximum ight times development of L. botrana during one day with an average
can be ascertained with our monitoring methods. These times temperature of T. Note that unlike the piecewise-linear function
observed in eld are called actual times of maximum ight and employed by the Touzeau model, the logistic function is more
they can be compared to the ones predicted by the Touzeau model biologically realistic because it is smooth and reects the fact that
to examine the efciency of the model. an increase in temperature beyond some limit does not signi-
cantly foster the speed of development of L. botrana. Daily
average temperatures were accumulated as in Equation (2),
2.4. Logistic function resulting in the new equation the cumulative value on the day k
being dened by,
A new model based on the logistic function to obtain suitable
predictions is proposed in this paper. Firstly, it is necessary to dene X
k
Sk sja; b f1 Td ja; b
Table 2 d1
Locations, coordinates and altitudes of the weather stations.
An estimate of the day of maximum ight of generation g (g is 1,
Municipality UTM Coordinate. Altitude 2 or 3) was determined by the threshold qg applied on the cumu-
Datum: European 1950 (m) lative values Sk. Firstly, we dened
Villanueva de Gumiel X: 30T 448875 Y:4620895 848
    
La Horra X: 30T 427814 Y:4623318 762 kg sa; b; qg min k : Sk sja; b>qg
Sotillo de la Ribera X: 30T 430797 Y:4622423 761
Nava de Roa X: 30T 421072 Y:4608749 706 where kg sja; b; qg is the rst day in which the cumulative value
Aranda de Duero X: 30T 439525 Y:4611525 830
Peael X: 30T 408006 Y:4605558 742
Sk sja; b exceeds the threshold qg. Therefore, the exact time of
maximum ight would be in the interval kg  1; kg , so we
 M. Amo-Salas et al. / Crop Protection 30 (2011) 1586e1593 1589

Daily Accumulated Temp. C and generations that were in data record. Note that another
20 meaningful criterion would be to minimize the sums of squares of
the intervals between the observed and predicted days, but the
error based on the absolute intervals is more robust against
15
outliers.

10 3. Results

Fig. 3 shows the ts between the actual data (black dots) and the
5 predictions of the Touzeau model (breakpoints between the curves
of accumulation of temperatures and horizontal lines) for the years
2004e2008, where horizontal lines are the thresholds established
10 10 20 30
Temperature C by the Touzeau model. The predictions of the rst ight was suit-
able because the actual ights of the rst generation took place
Fig. 2. Daily accumulation of temperature with respect to the average temperature when the accumulation of temperatures were approximately 125
according to Touzeau model.
DD > 10  C. However, for the second generation the ts between
the actual ights and the predictions of the Touzeau model were
interpolated to compute this time. Based on kg sja; b; qg we not accurate enough. Finally, for the third generation (in 2007 there
dened the estimate of the day of maximum ight by were only two generations, and this study included the data up to
the second generation in 2008) the differences between both were
 
^g sa; b; qg  k sa; b; qg   1
k even higher.
g
The lack of t showed in Fig. 3 could be expressed numerically
qg  Skg sja;b;qg 1 sja; b
using Equation (3). The predictions of maximum ight of the Tou-
Sk sja;b;qg sja; b  Sk sja;b;qg 1 sja; b zeau model were made using Equations (1) and (2) and the
g g
thresholds of Table 3. The cumulative error committed with the
where kg  1 is the day before kg , ^
kg is the interpolation of integer predictions of the Touzeau model was,
days kg  1 and kg . This value is a real value and therefore the X
function used for the estimation of the parameters is continuous. ErrTou jky;p;g  Touy;p;g j 4228:12 days
Under the condition of continuity the procedure of estimation is y;p;g
more precise. Skg is the cumulative value from 1 January to kg and
Note that this error is cumulative in the sense that it depends on
Skg 1 is the cumulative value from 1 January to kg  1.
^g sja; b; qg is the day (not neces- the total amount of data considered (year, plot and generation). In
Graphically, the estimate k
this study, the total number of times of maximum ight was 247, as
sarily an integer) on which the piecewise-linear interpolation
a result the relative error was 17.12 days per year, plot and
of the points Tk ; Sk sja; b, k 1, 2,., crosses the threshold qg .
generation.
The software used for the parameter estimation was Mathe-
The predictions were based on the temperature records of the
matica 7.0.
weather stations, which means that the predictions for the plots
Clearly, the estimates of the dates of maximum ight strongly
assigned to the same weather station were necessarily the same.
depend on the values of parameters a, b, q1, q2, q3. Hence, we
However, the actual days of maximum ight differed even for plots
needed to calibrate the model, which required nding the values of
with the same weather station, which implies that there is a lower
the parameters to minimize prediction error. In our model we used
limit on the achievable error. To compute this error limit, the
the error criterion of the form
median of the set of days of maximum ight for each year, weather
X    station and generation was considered as the optimum prediction.
Erra; b; q1 ; q2 ; q3 ^  
jky;p;g  k y;p;g sp;y a; b; qg (5) Thus, the minimum error was:
y;p;g
X
where ky;p;g are real (observed) days of the maximum ight of the
minErr jky;p;g  kw j 1015 days
^y;p;g sp;y ja; b; qg are y;p;g
year y, on the plot p, of the generation g, and k
the corresponding predictions yielded by the model with param- where kwy;p;g is the median of the set of days of maximum ight for
eters a; b; qg under the temperature record sp;y on the plot p during each year, weather station and generation. The median was
year y. The sum ranged through all the combinations of plots, years considered because it is robust against the outliers. In this case the

Table 3
Steps of development and accumulated temperatures sum necessary according to Touzeau model.

Step Temperatures Accumulated temperature Average duration of different

sum ( C) sum ( C) steps (days)
DD > 10  C DD > 10  C
Maximum ight 1st G 125 125 The adult lives 10e15 days
Egg eclosion 1st G 75 200 7e10 days after the laying
Larval development 170 370 5 larval stages: 21e30 days
Pupal formation and 130 500 Pupa: 12e14 days before the adults emerge
adult appearance of 2nd G (adult life: 10 days)
Egg eclosion 2nd G 65 565 7e10 days after the laying
Larval development 255 820 5 larval stages: 19e25 days
Chrysalis formation and 130 950 Pupa: 12e14 days before the adults emerge
adult appearance of 3rd G (adult life: 10 days)
Egg eclosion 3rd G 75 1025 7e10 days after the laying / hibernated larvae
1590 M. Amo-Salas et al. / Crop Protection 30 (2011) 1586e1593

Sum of Accumulated Temp. C 2004 Sum of Accumulated Temp. C 2005

1400 1500
1200

1000
1000
800

600

400 500

200

Days Days
50 100 150 200 250 300 350 50 100 150 200 250 300 350

Sum of Accumulated Temp. C 2006 Sum of Accumulated Temp. C 2007

1200
1500
1000

1000 800

600

500 400

200

Days Days
50 100 150 200 250 300 350 50 100 150 200 250 300 350

Sum of Accumulated Temp. C 2008

1200

1000

800

600

400

200

Days
50 100 150 200 250 300 350

Fig. 3. Plot of the accumulation of temperatures using the Touzeau model along the years 2004e2008. Black dots stand for the actual days of maximum ight and horizontal lines
stand for the thresholds of maximum ight according to the Touzeau model. Different lines for each year stand for the different weather stations.

relative error was 4.11 days per year, plot and generation. Notice the
large interval between the minimum error and the error of the
Touzeau model.
In order to compute the new model, the function of the error
dened by Equation (4) was minimized for the values of the
parameters
The weather stations provided other variables besides the
average temperature. An important factor related to the develop-
ment of the moth is the relative humidity. Another advantage of
this model is simplicity of extension to the case with more
exploratory variables. In this case, the function was dened by,

a 5:43; b 0:387; q1 24:32; q2 73:01; q3 113:8 Daily Development

1.0
The achieved value of the error was 1704.6 days, which
amounted to 6.9 days per year, plot and generation. This is an 0.8
empirical model based on the data in order to obtain precise ts
with the actual data, but the range of years considered in this study
0.6
was large enough to assume that the model obtained will provide
suitable predictions. Fig. 4 shows a graph of the function f1 (see
equation (4)) with the values of a and b given above and shows 0.4

daily development of L. botrana with respect to the temperature,

now the daily development of the moth is expressed in the interval 0.2
[0, 1]. The curve is more realistic from a biological point of view.
Note that based only on the empirical data, the model reveals the Temperature C
10 20 30
fact that under 10  C the development of the moth is negligible, and 10

an increase of the temperatures beyond the limit of approximately Fig. 4. Daily development of the moth with respect to the temperature for the new
22  C does not have a further positive effect. model.
 M. Amo-Salas et al. / Crop Protection 30 (2011) 1586e1593
1 Touzeau model was 17.38 days per year, plot and generation, for the
f2 T; Hja; b; g
1 eabTgH
where H is the relative humidity. The procedure of calibration was
the same, the resulting optimum values of the parameters and the
thresholds were
a 3:32; b 0:3; g 0:015; q1 26:62;
q2 70:96; q3 114:81
and the error was 1626.1 days, i.e., 6.58 days per year, plot and
generation. The inclusion of a new parameter reduced the error,
nevertheless the value of error for the model with and without
relative humidity was close. Fig. 5 exhibits a graph of the function f2
(see equation (6)) with the values of a, b and g given above and
shows the daily development of L. botrana with respect to
temperature and relative humidity.
Thus, using this procedure the Touzeau model was improved.
The function dened by the Touzeau model was assumed but new
thresholds were computed in order to minimize the error. These
new computed thresholds were
q1 144:25; q2 666:48; q3 1216
and the error was 1816.5 days, equal to 7.35 days per year, plot and
generation. These thresholds are measured in degrees-days
DD > 10  C and are larger than those of the Touzeau model.
Hence, the calibration of the thresholds improved the predictions
of the Touzeau model. Notice that the model based on the logistic
function (with both temperature and humidity as variables)
improved the precision of the prediction by almost one day
compared even to the optimal thresholds used with the standard
Touzeau model. This can be explained by the fact that the logistic
function has a more natural shape biologically than the unbounded
and non smooth function of the Touzeau model, as well as by the
inclusion of humidity as another relevant factor.
Finally the data from 2009 to 2010 were used to validate the
model computed. Fig. 6 shows the ts between the predictions with
the different models (the Touzeau model, the new model with the
variable temperature, the new model with the variables tempera-
ture and relative humidity and the Touzeau model with the new
thresholds) and the actual days of maximum ight for the years
2009 and 2010 (in 2010 data were included up to the second
generation). It shows a clear difference between the ts for the
Touzeau model and the ts for the new model computed. These ts
could be compared numerically using the denition of error of
Equation (3). Thus, for the data of 2009 and 2010 the error of the
100
Relative humidity
50
0
1.0
0.5 Daily Development
0.0
30
20
10
0 Temperature C
10
Fig. 5. Daily development of the moth with respect to the temperature and relative
humidity for the new model.
1591
(6)
new model with the variable temperature the error was 8.48 days
per year, plot and generation and for the new model with the
variables temperature and relative humidity, the error was 8.24
days per year, plot and generation. Finally, for the Touzeau model
with the new thresholds the error was 9.84 days per year, plot and
generation.
4. Discussion
Pheromone trap observations and daily temperature forecasts
could be helpful in identifying the peak ight of L. botrana,
achieving better timing of treatments. Considering the increasing
interest for biorational insecticides where precise timing of treat-
ments is important, daily temperature models could be a useful tool
in improving their efciency.
At present, L. botrana is being monitored with an aim to knowing
its population and the dates of maximum ight in the Ribera del
Duero region. The actual data exhibited signicant discrepancy with
the predictions of the widely used Touzeau model. These differences
have also been observed in nearby vineyard regions (Armendriz
et al., 2010). This lack of t might be the result of the differences
between the region where the Touzeau model was developed and
where it is now applied. The Touzeau model was developed in
France where the temperatures are milder while the Ribera del
Duero region has a continental climate where the daily difference
between the maximum and the minimum temperature is high, even
in summer. The climate conditions (between 1 April and the 5
October) for a region close to the place where the Touzeau model
was developed, showed that 11% of days are not favorable for the
activity of L. botrana, according to the climate-graph of Stelwaag
(Bovey, 1966), while in the Ribera del Duero region this percentage is
around 26% (Armendriz et al., 2010). Moreover, in the Touzeau
model, the development of the moth increases linearly with the
temperature while, for instance, in a day with an average temper-
ature of 40  C the development of the moth is not similar to the
development of three days with 20  C. In fact, extremely high
temperatures do not have a positive effect on the moth, but can even
have a negative one (Tzanakakis et al., 1988). Hence, it is necessary
that better models be developed. The model developed in this paper,
which predicts the times of maximum ight, could be used as
a complementary tool to the Touzeau model, which details the
different stages of the development of L. botrana.
Our empirically-based model is merely intended to describe the
timing of maximum moth ight and not the number of moths
ying. The model proposed is based on the logistic function. This
function led to a model that provides better estimates. It is also
biologically more realistic and suitable for the incorporation of
additional quantitative variables. The Touzeau cumulative proce-
dure was used with the logistic function and the error was dened
as the absolute value of the distances between the actual data and
predictions.
A new model with only one variable, average temperature, was
developed. The parameters of the logistic function and the
thresholds for each generation were estimated in order to minimize
errors. This model reduced the value of the error of the predictions
by more than 10 days per year, plot and generation with respect to
the Touzeau model. The values of the thresholds showed that the
second and third generations need approximately the same sum of
average temperatures to reach the adult stage, while the rst needs
less. This variability could be caused by many factors such as vari-
ation in development time or in oviposition duration and decreased
efciency of monitoring tools over time.
With the new model with two variables, average temperature
and relative humidity, the error was further reduced. Note these
1592 M. Amo-Salas et al. / Crop Protection 30 (2011) 1586e1593

Sum of Accumulated Temp . C 2009 2010

Sum of Accumulated Temp . C

1400
1500
1200

1000
1000
800

600
500
400

200

Days Days
50 100 150 200 250 300 350 50 100 150 200 250 300 350

2009
Development Accumulated Development Accumulated 2010

150 140

120

100
100
80

60
50
40

20

Days Days
50 100 150 200 250 300 350 50 100 150 200 250 300 350

Development Accumulated 2009 2010

Development Accumulated

150 140
120
100
100
80
60
50 40
20
Days Days
50 100 150 200 250 300 350 50 100 150 200 250 300 350

2009 2010
Sum of Accumulated Temp . C Sum of Accumulated Temp . C

1500 1400
1200
1000
1000
800
600
500
400
200
Days Days
50 100 150 200 250 300 350 50 100 150 200 250 300 350

Fig. 6. Plot of the accumulation of temperatures using the Touzeau model (rst row), developments accumulated using the new model with variable temperature (second row),
developments accumulated using variables temperature and relative humidity (third row) and accumulation of temperatures using the Touzeau model with the new thresholds
(fourth row) along the years 2009 and 2010. Black dots stand for the actual days of maximum ight and horizontal lines stand for the thresholds of maximum ight according to the
Touzeau model, the new model with variable temperature, the new model with the variables temperature and relative humidity and the Touzeau model with the new thresholds,
respectively. Different lines for each year stand for the different weather stations.

models are empirical models based on the data collected from the following way: using meteorological variables such as temperature
Ribera del Duero region. or humidity, the advisor could update (on a daily basis) a cumula-
While the thresholds do not have a biological interpretation, tive sum quantifying the estimated stage of the development of the
they could guide decisions of a crop protection advisor in the moth. As the sum approaches the threshold, the advisor would
 M. Amo-Salas et al. / Crop Protection 30 (2011) 1586e1593
know that the day of maximum ight is close. Of course, the day of
maximum ight does not need to exactly correspond to the day on
which the cumulative sum reaches the threshold; such a perfect
prediction is impossible in practice. However, we have tuned the
parameters of the model so that the expected error of the predic-
tion has been minimized.
Finally, the thresholds of the Touzeau model were recomputed
to minimize the error. With these new thresholds the value of the
error was reduced although it was still higher than the error of the
new model. Moreover, the model was validated with the data of
2009 and 2010. Fig. 6 shows the improvement of the predictions of
the new model with respect to the Touzeau model.
Acknowledgments
We want to thank the editor Jerry Cross and the anonymous
referees for helpful comments. We would also thank Regulatory
Council for the Designation of Origin (D.O.) Ribera del Duero and
Junta de Castilla y Len. This work was sponsored by Slovak VEGA
Grant No. 1/0077/09 and Junta de Comunidades de Castilla-La
Mancha PEII10-0291-1850.
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