Modelling the Effects of Climate Change on the Potential Feeding Activity of

Thaumetopoea pityocampa (Den. & Schiff.) (Lep., Notodontidae) in France

Author(s): Christelle Robinet, Peter Baier, Josef Pennerstorfer, Axel Schopf and Alain
Roques
Source: Global Ecology and Biogeography, Vol. 16, No. 4 (Jul., 2007), pp. 460-471
Published by: Wiley
Stable URL: http://www.jstor.org/stable/30137901
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Global Ecology and Biogeography, (Global Ecol. Biogeogr.) (2007) 16, 460-471

Modelling the effects of climate change

on the potential feeding activity of
Thaumetopoea pit yo ca mpa (Den. &
Schiff.) (Lep., Notodontidae) in France
Christelle Robinet'']", Peter Baier', Iosef Pennerstorfer', Axel Schopf and
Alain Roques'

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 1INRA UR 633 Zoologie Forestiere, Avenue de
la pomme de pin, BP 20619 Ardon, 45166
Olivet, France and 2 BOKU - Department of
Forest
and Soil Sciences, Institute ofFo rest Entomology,
Forest Pathology & Forest Protection,
Hasenauerstr. 38, A -1190 Vienna, Austria
ABSTRACT
Aim We investigated whether climate change has affected the potential feeding
activity of a winter active larva, the pine pro cessionary moth (PPM), Thaumetopoea
pityocampa L., and whether it may explain its range expansion.
Location The study area is France and, at a smaller scale, the Paris Basin.
Methods We used a statistical model derived from Huchon and Demolin [1970
Revue Forestiere Francoise (special issue: La lutte biologique en !oret) , 220-234] to test
whether their model, updated with climate change, could explain the observed range
expansion. Since Battisti and colleagues have recently shown that climate could
affect survival of the PPM through its effect on feeding activity, we also developed a
mechanistic model based on larval feeding requirements (night air temperature
above 0 °C and temperature inside the nest above 9 °C on the preceding day). We
reconstructed the geographical distribution of feeding activity and we compared the
resulting change with the PPM range expansion.

Results The statistical model did not successfully predict the observed expansion
but the mechanistic model showed considerable change in the feeding activity of the
PPM. In the Paris Basin, the PPM border coincided with a zone unfavourable for
feeding activity in the period 1992-96. Feeding conditions became more favourable
in the period 2001-04, and the PPM succeeded in crossing this zone. Over larger
temporal and spatial scales improved feeding conditions in the north-western
part of France were forecast by the mechanistic modeL

Main conclusions (1) The range distribution of the PPM in the Paris Basin is no
longer limited by unfavourable feeding conditions. (2) The pattern of range expan-
sion of the PPM is now governed mainly by its dispersal capabilities and host tree
distribution. (3) At the country scale, this approach gives an approximate prediction
of the potential distribution of the PPM, though the model may not be reliable in
"Correspondence: Christelle Robinet, INRA
mountainous regions.
Zoologie Forestiere, Avenue de la pomme de
pin, BP 20619 Ardon, 45166 Olivet, France.
Keywords
E-mail: robinet@orleans.inra.fr
t Present address: USDA Forest Service, Bioclimatic envelope, climate change, feeding activity, forest defoliator, modelling,
180 Canfield Street, Morgantown, range expansion, spatial dynamics, Thaumetopoea pityocampa.
WV 26505, USA.

in summer (European Environment Agency, 2004). The overall

INTRODUCTION
The world's climate has significantly warmed by 0.6 ± 0.2 °C
during the last century and the mean global temperature is
projected to increase by l.4-5.8 °C by 2100 (IPCC, 2001). Europe
has warmed more than the global average, with an increase of
0.95 °C, and winter temperatures have increased more than those
response of ecosystems remains difficult to predict, but many
recent studies have already highlighted the impact of global cli-
mate change on animal and plant species (Hughes, 2000; Walther
et al., 2002; Parmesan & Yohe, 2003; Root et al., 2003). They show
a great variety of reactions and, in particular, the shift of species
ranges. Thus, European species whose distributions are closely

001: 10.1111/j.1466-8238.2006.00302.x © 2007 The Authors

460 Journal compilation © 2007 Blackwell Publishing Ltd www.blackwellpublishing.com/geb

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 Effects of climate change on the pine processionary moth

Figure 1 Expanding range of the pine processionary moth (PPM) in France. The shaded area is the PPM range distribution observed (a) in
1969-80 (CTGREF- INRA, 1980) and (b) in 2005 (the PPM front was georeferenced with a GPS by INRA Orleans; Robinet, 2006). Dotted lines
indicate that the position of the front is not precisely determined; names in italics refer to mountainous areas.
associated with winter temperatures may be particularly pre-
disposed to undergo changes in the next several years. Birds (Tho-
mas & Lennon, 1999; Visser et al., 2003), amphibians, reptiles
(Araujo et al., 2006), vegetation (Thuiller et al., 2005) and insects
- notably butterflies (Parmesan et al., 1999; Thomas et al., 2001;
Warren et al., 2001; Hill et al., 2002; Konvicka et al., 2003) -
have already been affected by this change in Europe. Some non-
migratory European butterflies have shifted 35 to 240 km
northwards during the last century (Parmesan et al., 1999),
while climatic isotherms have shifted the equivalent of 120 km
northwards (Watson et al., 1998).
Since environmental factors explain a large part of species
distributions, they are commonly used to determine potential
habitat distributions and to predict changes (Guisan & Zimmer-
mann, 2000; Guisan & Thuiller, 2005). Among these factors,
climatic conditions that affect the ability of individuals to survive
and grow define a bioclimatic envelope which is relatively useful
for predicting species distributions and assessing the effect of
climate change (Peterson et al., 2002; Pearson & Dawson, 2003;
Thomas et al., 2004; Thuiller et al., 2006a). The accuracy of this
method has been widely debated due to its apparent lack of real-
ism: it ignores fundamental processes like dispersal and trophic
interactions with other species (Davis et al., 1998a,b). However,
the modelling of a bioclimatic envelope appears to be an efficient
way to specify basic conditions for potential species occurrence.
Phytophagous insects generally provide convenient objects for
exploring this effect due to their physiological dependence on
climate (Karban & Strauss, 2004) and, consequently, their rapid
response to climate change (Bale et al., 2002). Insect species
© 2007 The Authors
Global Ecology and Biogeography, 16,460-471, Journal compilation © 2007 Blackwell Publishing Ltd
whose larvae remain active during winter - such as the pine pro-
cessionary moth (PPM), Thaumetopoea pityocampa (Denis &
Schiffermuller) (Lepidoptera, Notodontidae) - may be particu-
larly affected by global warming (Battisti et al., 2005).
Since latitudinal and altitudinal expansion of PPM outbreaks
have been reported across all of southern Europe (Hodar et al.,
2003; Battisti et al., 2005), we investigated the impact of climate
warming on the range expansion of the PPM in order to identify
areas likely to be invaded in the future. A deeper understanding
of this expansion is of great interest for public health and forestry
because late-instar larvae can release urticating hairs which often
cause severe allergies (Lamy, 1990), and the reduced growth of
pines defoliated by the PPM results in significant economic loss
(Laurent-Hervouet, 1986).
In France, the range of the PPM has expanded considerably
over the last two to three decades (Fig. 1). The northward expan-
sion was surveyed in detail in the southern part of the Paris Basin
(France) (Battisti et al., 2005; Robinet, 2006). The front has
shifted by 27.1 km decade-1 between 1972 and 2004, and has
accelerated during the last 10 years (55.6 km decade") (Battisti et
al.,
2005). To date, the effect of global warming on the southern limit
of-EEM populations in North Africa is unknown. In low latitudes
the moths emerge later (after the warmest temperatures), and
this strategy enables the PPM to avoid extreme heat (Demolin,
1969a).
The study of the impact of climate on the phenology and dis-
tribution of the PPM began long before the question of climate
warming. The first model was based on a correlative approach to
determine the potential range of the PPM (Huchon & Demolin,
461
C. Robi net et a/.
1970). According to this model, the PPM cannot survive when
the mean minimum temperature in January is below -4°C or
when the cumulative annual sunshine is below 1800 hours.
When the temperature is above -4 °C but below 0 °C, 100 hours
of additional sunshine per degree below 0 °C are required to en-
able the PPM to survive. These weather variables were averaged
from 1946 to 1960 and projected on a map to define the theoretical
exclusion area, but the recent expansion invalidates this historical
model.
Battisti et al. (2005) showed that micro climatic conditions
could also constrain the distribution of the PPM by governing
the feeding activity of the larvae during the cold period. Two
conditions have to be satisfied to enable the PPM larvae to feed:
the temperature inside the nest during the day has to reach more
than 9 °C (activation temperature) and the air temperature
during the following night must be above 0 °C (potential feeding
temperature). If one of these conditions is not fulfilled, the larvae
do not feed. The understanding of this ecophysiological process
was based on a combination of laboratory and field experiments
(Battisti et al., 2005).
In this paper, we investigate the effects of climate change on
the distribution of the PPM using two different approaches.
First, we test the historical model derived from Huchon and
Demolin (1970) taking into account climate warming. Since data
for the cumulative annual sunshine are not available, we consider
the solar radiation variable and we use a statistical model to
explain the range of the PPM. Second, we develop a mechanistic
model based on the micro climatic conditions that control feed-
ing activity (Battisti et al., 2005). We then construct a spatially
and temporally localized model for the southern Paris Basin.
Finally, we make a generalization at the country scale and at a
larger temporal scale to determine more clearly the effect of cli-
mate change. This study intends to provide a first approximation
of the potential change of the distribution of the PPM in France.
METHODS
Study system
The PPM is one of the most important defoliators of forests in
southern Europe. The gregarious larvae spin a white silk nest and
develop throughout the winter on pine trees. When conditions
are suitable, they leave nests to feed on needles during the night.
Climate determines the date of the nest construction and the
duration of larval development (Huchon & Demolin, 1970). Larval
survival is affected by the occurrence of extreme temperatures
(the lower lethal temperature of larvae inside the nest being
-16°C; Dernolin, 1969a) and by temperature thresholds
responsible for feeding activity (Battisti et al., 2005). Since global
warming may cause its range expansion, modelling the relation-
ship between climate and distribution of the PPM would enable
forecasts of future PPM population distributions.
As the larval development of the PPM overlaps two consecutive
calendar years, commonly from the autumn of the year n - 1 to
the spring of the year n, we will refer hereafter to the year n
instead of the preceding year (n - 1).
462 Global Ecology and Biogeography, 16,460-471,
Climate scenario
Both statistical and mechanistic models require weather data for
relatively long periods of time to make predictions for the future
over the whole of France. To avoid discontinuity, we used the
data from the climate scenario for previous years (except for the
localized model).
The best approach would consist of testing the effect of differ-
ent climate scenarios in order to determine the resulting range of
predictions (concerning the occurrence and potential feeding
activity of the PPM). As we considered a relatively small spatial
area, we decided to use the regionalized climate scenario of
Meteo- France 'ARPEGE-Climat' (Gibelin & Deque, 2001;
Cloppet, 2002). This climate scenario is based on scenario B2
which assumes a moderate increase of greenhouse gas (the CO2 con-
centration is supposed to only double between 1970 and 2100).
It provides climatic forecasts at 360 locations separated by about
50 km over all of France (211 locations) and boundary areas from
1960 to 2100. Mean temperature is projected to increase by 3.1 °C
over this period.
Including climate warming in the historical model
According to Huchon and Demolin (1970), the distribution of
the PPM is limited by a combination of two climatic variables:
the mean minimum temperature in January (TN], °C) and the
cumulative annual sunshine (hours). Because data on this annual
sunshine variable were not available, we substituted a variable
from the climate scenario 'ARPEGE-Climat: the annual mean solar
radiation [aSRad, watt-hour (Wh) m-2]. The coastline extractor
from the National Geophysical Data Center, NOAA Satellite and
Information Service (http://www.ngdc.noaa.gov/ngdc.html) ,
provided a convenient file for identifying the geographical co-
ordinates of the French border. The range distribution of the
PPM was surveyed across France during 1970-80 (CTGREF-
INRA, 1980) and in 2005 (Robinet, 2006), based on nest moni-
toring (Fig. 1). Each point of the climatic grid was assigned by
the presence or absence of the PPM according to the position of
the front at the two different points in time (1970-80 and 2005).
The historical model used a piecewise linear relationship
to determine areas where the PPM could survive (Huchon &
Demolin, 1970). Since the sunshine variable was not exactly the
same in our study, the relationship might differ. Numerous
methods are currently available for predicting species ranges
from habitat data (Elith et al., 2006). In our case (presence and
absence data, two predictors, and no community data), we
decided to choose a flexible statistical model not restricted to
linear cases: the generalized additive model (GAM routine of the
statistical language R; Hastie & Tibshirani, 1990; Wood, 2006).
This model can generally describe complex relationships with a
high predictive power for such data sets. We considered the PPM
front in 1970-80 and the mean of meteorological variables (TN]
and aSRad) over the same period. To evaluate the model
performance, we calculated the jack-knife estimate of the
correct classification rate (%) and the 95% confidence interval.
Then we applied this model to the meteorological conditions in
© 2007 The Authors
Journal compilation © 2007 Blackwell Publishing Ltd
 Effects of climate change on the pine processionary moth

Figure 2 Location of the study area in the

southern Paris Basin.
1995-2005 and we calculated the error made by comparison
with the front in 2005. A projection of PPM occurrence was then
established for 2020-30.
Modelling nest temperature and feeding ability
Microclimatic conditions in the nest are crucial factors for the
survival of winter-active PPM larvae. Winter sun can warm the
nests to several degrees above ambient temperature. Solar energy
plays a central role in the metabolism and development of PPM
larvae, especially when the ambient maximum temperature is
too low for feeding activation and digestion of previously
ingested food (Demolin, 1969b; Breuer, 1997; Netherer et al.,
2005).
The relationship between surrounding air temperature, solar
radiation and nest temperature was investigated under semi-field
conditions in the garden at the Institute of Forest Entomology in
Vienna, Austria. Air temperature and nest temperatures at the
front and back sides of six PPM nests (origin: Venosta Valley,
northern Italy) were recorded hourly using Tinytalk data loggers
from 22 January until 30 March 2004 (n = 70 days). Solar radia-
tion was measured using a pyranometer (Starpyranometer 8101,
Schenk, Vienna, Austria). For data analysis, the hourly mean
values of nest temperatures measured at the back and front
sides of the six observed nests were calculated. The relationships
between minimum nest temperature and air temperature, and
also between maximum nest temperature, air temperature and
solar radiation were analysed by linear regression (see Appendix S1
in Supplementary Material) using the statistical software package
sr-ss 12.0.1 (SPSS Inc., 1990).
To calculate the duration of feeding activity during the night,
continuous hourly recordings of temperatures were needed to
check whether the temperature reached 9 °C in the nest during
the day and was higher than 0 °C at night outside the nest
(Battisti et al., 2005). Since only daily minimum and maximum
temperatures are commonly provided by meteorological services,
we considered that PPM larvae could feed effectively if the daily
maximum temperature in the nest was above 9 °C and the daily
minimum air temperature of the following day was above 0 "C.
© 2007 The Authors
Global Ecology and Biogeography, 16, 460-471, Journal compilation © 2007 Blackwell Publishing Ltd
We reconstructed the temperature inside the nest using the
previous regression model and we calculated the resulting potential
feeding activity of larvae at various locations in the field (see
below for the precise locations) according to the mechanism
described by Battisti et al. (2005). For each day, we checked
whether both the daily maximum nest temperature and minimum
air temperature requirements were realized. If so, we concluded
that PPM larvae were able to feed. For each winter period (October-
March) and each location, the feeding activity of the larvae was
estimated by the number of feeding days (NFD, or more exactly
the number of feeding nights) and the longest period of no feed-
ing (or starvation, LPS). Feeding ability is particularly important
during the 'cold period' defined by Battisti et al. (2005) (i.e. the
period during which the weekly mean of the daily minimum air
temperature is below 0 °C). Since this period could differ between
locations and years, we defined the 'cold period' in this study as
the time from October to March to obtain comparable results.
This period coincides with larval development of the PPM in
most areas of France. Mean NFD and LPS were calculated over
several years (see below for the precise years) for each location (R
Development Core Team, 2004) and then it was spatially
interpolated using kriging (Surfer version 7.00, copyright
©1993-99, Golden Software, Inc.).
Localized model of feeding ability
In order to analyse the role of feeding ability in PPM range
expansion, we considered the area in central France where the
insect is currently expanding northwards up to the Paris Basin
(Fig. 2). This study area includes the core area, which was histor-
ically colonized (hereafter our reference is Tours; 47°23' N,
00°4-1' E); the expansion area, which has been mostly colonized
after 1992 (our reference is Orleans; 47°55' N, 01 °54' E); and far
beyond the current front up to the Paris Basin (our reference for
this uncolonized area is Melun; 48°32' N, 02°40' E).
Two discontinuous periods representative of two distinct
phases in the range expansion of the PPM were chosen: 1992-96
(first detection of PPM expansion) and 2001-04 (uninterrupted
shift of the front carefully observed) in order to detect eventual
463
C. Robinet et a/.
changes in feeding ability and to compare the results with the
observed range expansion (front position reported in 1992, 1996
and 2004 by Battisti et al., 2005).
Meteorological stations (n = 11) were selected over the core
area, the expansion area, and outside the PPM distribution area.
Meteorological conditions associated with the feeding capacity of
the PPM were used in this model: daily minimum and maximum
temperatures (0C) and daily solar radiation (Wh m-2) (data
source: Meteo- France; European Climate Assessment, Tank et
al.,
2002). Missing values of solar radiation were estimated via Meteo-
Sat images (see Appendix S2 in Supplementary Material).
In addition to these data, field conditions were recorded at
eight sites where the experiment to investigate the feeding mech-
anism was conducted (Battisti et al., 2005). These sites were
located on two parallel axes going from the core area to beyond
the leading edge of the current PPM distribution. We calculated
the feeding activity (NFD and LPS) on these sites and we
included the results in the spatial interpolation for the second
period (2001-04) to obtain a higher spatial resolution of the
feeding ability. Missing data were estimated by a regression based
on the nearest meteorological stations.
Indicator of feeding ability
To study the intrinsic effect of climate change, we had to consider
a longer time period than just the two previous periods (1992-96
and 2001-04) because natural oscillation of the climate during
relatively short periods could affect the results. Since we had no
data on observed solar radiation over such a long period, we
searched for a general climatic indicator explaining the annual
number of feeding days (NFD) and the longest period of no feed-
ing (LPS) in order to reconstruct the feeding ability in the past
and to investigate the existence of a possible trend. Correlation
analysis, based on the data from 11 meteorological stations
during the nine cold periods (winters from 1991-92 to 1995-96
and from 2000-01 to 2003-04), was conducted to find the best
explanatory variable ('simple indicator') among those used in
the literature or those likely to explain the feeding ability during
the cold period. We considered the following variables: the abso-
lute minimum air temperature; the mean minimum air temper-
ature in January; the annual cumulative sunshine expressed in
hours (variables used by Demolin, 1969a and Huchon & Demolin,
1970); and the mean of minimum and maximum air temperatures
from October to March (period corresponding to PPM larval
development and the feeding period in the Paris Basin). A 'two
sample r-test' was also applied to these variables to compare
them before and after 1988 (middle of the period) along the axis
Tours (historically colonized )-Orleans (colonized in 1992)-
Melun (not yet colonized). We investigated a possible trend in
the simple indicator from 1972 to 2004 using a linear regression.
In order to reconstruct the feeding ability at larger temporal
and spatial scales (using the climate scenario), we conducted
multiple regressions using two predictors among the previous
variables and solar radiation, and we tested all possible comb ina -
tions. The 'global indicator' was defined as the combination for
which the regression showed the best fit.
464 Global Ecology and Biogeography, 16, 460-471,
Extended model of feeding ability
In order to get an overview of the potential change of the PPM's
range distribution at a larger scale, we utilized the 'global indicator'
of feeding ability to simplify and extend the feeding model to
France for the next decades. We calculated a projection of this
feeding ability for the whole of France using the climate scenario
'ARPEGE-Climat'. In this analysis, we considered 30-year periods
(1971-2000, 2001-30 and 2031-60) in order to eliminate the
natural oscillation of climate and to keep representative condi-
tions of the warming trend. The projection produced for the first
period (from 1971 to 2000) was used as a reference for compari-
son with future predictions.
RESULTS
Including climate warming in the historical model
The generalized additive model applied to the PPM distribution
in 1970-80 generated errors around 11%: it incorrectly predicted
the presence of the PPM in 6.2% of the cases (mostly in central
France at the border of the range distribution in the Massif
Central) and it incorrectly predicted the absence for 5.2% of the
points (mostly in the northern range distribution) (Fig. 3a). The
mean of the jack-knife estimate of the correct classification rate
was 83.4%, with a 95% confidence interval of73.9-92.9%. When
we applied this model to the climatic conditions in 1995-2005
and compared the results with the PPM front in 2005, it incor-
rectly predicted PPM presence for l.9% and absence for 19.9%
with a total number of errors of around 22% (Fig. 3b). When we
considered the projected climatic conditions during 2020-30,
the model still incorrectly predicted PPM absence for 13.7% of
the points compared with the front in 2005 (mainly in the
middle of the northern range distribution, Fig. 3c).
Localized model of feeding ability
The mean number of feeding days (Fig. 4a-c) and the longest
period of no feeding (Fig. 4d-f) showed a similar pattern.
Feeding conditions became more favourable across the whole
area. A relatively unfavourable region appeared in the first
period, splitting the potential feeding area into two parts: the first
one was located in the core area around Tours (where the PPM
was historically present) and another one in the Paris Basin,
where the PPM was absent. In contrast, climatic conditions for
feeding were far better nearly everywhere in the whole region
during the second period (except for a small area in the southern
part of the Paris Basin). The pattern of the front in 2004 was
particularly interesting, because the front apparently shifted more
quickly where the unfavourable area was thinner but the front
moved less rapidly near the most unfavourable areas (especially
north-east of Orleans). Also, minimum temperatures recorded
in the field were on average 1.46 °C lower than temperatures
recorded by the meteorological stations of Meteo- France
(for comparison, we considered the two nearest Meteo- France
stations for each site). Therefore, the estimated feeding ability of
© 2007 The Authors
Journal compilation © 2007 Blackwell Publishing Ltd
 Effects of climate change on the pine processionary moth

*Reference front: 1970-1980 *Reference front: 2005 *Reference front: 2005

(
,A
 • PPM presence observed* and predicted o PPM presence observed' but not predicted PPM presence not observed" but predicted

Figure 3 Maps of PPM range distribution over three periods of time: (a) 1970-80, (b) 1995-2005 and (c) 2020-30, resulting from the
generalized additive model (using the points defined by the climate scenario).

140
138
136
134
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~-0 132
130
0> 128
c
:0 126
Q)
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a.
c 16
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~ 15

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(i) 14

(i)
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0>
c 12
~ 0

11

10
1992-1996 2001-2004 2001-2004, Including field observations

e Meteorological stations used to build the map

£ Field observations used to build the map

:~ PPM front in 2004
o Other meteorological stations used for temporal analysis
~ ••• ••••••• •••• ••••• PPM front in 2004 (disconnected observations)

Figure 4 Spatial distribution of the mean number of feeding days, NFD (a.b,c), and the mean of the longest period of no feeding, LPS (d.e.f),
of PPM larvae in the Paris Basin region during 1992-96 (a.d) and during 2001-04 (b,e) according to the feeding model. For the second period,
two maps have been plotted. The second ones additionally incorporate temperatures recorded in the field (c.f), The position of the PPM front
for different years is also indicated.

© 2007 The Authors

Global Ecology and Biogeography, 16,460-471, Journal compilation © 2007 Blackwell Publishing Ltd 465

PPM front in1972

- - - - - PPM front in1992
• - • - • - PPM front in 1996
C. Robinet et al.
Table 1 Coefficient of determination (R2) of various temperature The longest period of no feeding was badly estimated by
parameters as predictors of the number of feeding days (NFD) and multiple regression (R2 < 0.33). Therefore, we chose to evaluate
the longest period of no feeding (IPS)
the feeding ability only by the number of feeding days during the
cold period.
Predictors NFD IPS

r.: Oct-March (Oe) R2 = 0.77 (P < 0.001) R2 = 0.29 (P < 0.001) Extended model of feeding ability
T max Oct-March (0C) R2 = 0.55 (P < 0.001) R2 = 0.23 (P < 0.001)
The maps based upon the regression on Tmin and SRad and using
Tmin Jan (0C) R2 = 0.06 (P = 0.011) R2 = 0.09 (P = 0.003)
the climate scenario (Fig. 5) showed evidence for a future geo-
graphical change in PPM feeding capacity. In 1971-2000, Orleans
PPM was lower when field observations were included in the was close to the isoline of 130 feeding days. This isoline moves
analysis (Fig. 4c,f). further east very quickly in the 2001-30 period and more slowly
after 2030. Eastern France and mountainous regions are likely to
Climatic indicators of the feeding ability remain relatively unfavourable areas.

The best predictors of the number of feeding days were: the mean
minimum air temperature from October to March; the mean
maximum air temperature from October to March (though it
was highly correlated with minimum temperature; R 2 = 0.69,
P« 0.001); and the mean minimum temperature
(Table 1). The first predictor explained a particularly large part of
the establishment of the PPM in the Paris Basin, whereas the
absolute minimum temperature, the mean minimum tempera-
ture in January, and the duration of sunshine did not differ
greatly between the locations (Table 2). The longest period of no
feeding was also explained by these variables but the correlations
were not as strong as for the number of feeding days (Table 1).
Therefore, the mean minimum air temperature from October to
March (T min' °C) could be used as a 'simple indicator' for the
estimation of both the number of feeding days and the longest
period of no feeding, and for evaluating the long-term fluctuation
of feeding ability (see Appendix S3 in Supplementary Material).
Multiple regression analysis showed that the mean minimum
temperature from October to March (T min) and annual mean of
solar radiation (SRad) accounted for 81 % of the variance of the
number of feeding days (NFD). The coefficients of this multiple
regression were highly significant (P < 0.001):
NFD = 47.25 + 14.43 Tmin + 0.016 SRad.
This combination defined the 'global indicator'.
DISCUSSION
Improvement in the PPM distribution model
in January The historical model (Huchon & Demolin, 1970) is consistent
with the stable PPM range distribution in the past but it does not
successfully predict the current expansion. The main reason for
this failure seems to be the correlative approach of simply com-
paring meteorological variables with the observed PPM range
distribution. In contrast, it is valuable to describe the complex
relationships between PPM survival, climatic conditions and nest
temperature. Our results confirm that maximum nest temperature
is determined both by the maximum air temperature and the
daily amount of solar radiation as reported by Breuer et al. (1989)
and Breuer and Devkota (1990). Our approach takes into account
physiologically based processes (such as feeding ability of the
PPM during the cold period) for modelling and mapping of the
range and the expansion of the PPM under unstable climatic
conditions. This approach substantially improves our insight into
the relevant mechanisms of recent and future PPM range expansion.
Localized model and the effect of microclimates
Modelling of the feeding ability in the Paris Basin revealed that
the climatic conditions are now favourable for winter survival of

Table 2 Mean and standard deviation (SD, in brackets) of four climatic parameters: the mean of minimum air temperature from October to
March (0C), the absolute minimum air temperature (0C), the mean of minimum daily air temperature in January (0C) and the annual duration
of sunshine (hours). Their value is given for three locations along a gradient of PPM presence over two periods of time. For a given variable, if
two mean values are followed by the same letter, this means that they are not significantly different (two-sample t-test, P < 0.05). The shaded
area shows that temperature in the expansion area (Orleans and Melun) during the second period was not significantly different from
temperature in
the core area (Tours) during the first period

Tmin Oct-Mar (Oe) Abs. T min (0C) Tmin Jan (Oe) Sunshine (h)

Location 1972-87 1989-2004 1972-87 1989-2004 1972-87 1989-2004 1972-87 1989-2004

Tours -'7.lO a (4.44) -7.13 a (2.52) 1.29 ab (2.58) 2.32 a (1.49) 1838 a (153) 1857 a (179)
Orleans -9.31 a (3.82) -8.53 a (2.54) 0.17 b (2.71) 1.69ab (1.50) 1791 a (150) 1790 a (210)
Melun 2.44 c (0.85) -9.36 a (4.01) -8.28 a (2.68) 0.19 b (2.92) 1.45 ab (1.50) 1667 b (161) 1795 a (182)

© 2007 The Authors

466 Global Ecology and Biogeography, 16,460-471, Journal compilation © 2007 Blackwell Publishing Ltd
 Effects of climate change on the pine processionary moth

Figure 5 Feeding activity generalized to France using the climate scenario of Meteo-France over three periods of time: (a) 1971-2000, (b) 2001-
30 and (c) 2031-60.
the PPM over the entire area. An isolated PPM colony recently
discovered (November 2003, J.-P. Raimbault & c. Robinet, pers.
obs.) in eastern Paris - probably introduced accidentally-
confirms that the PPM is able to survive far beyond its present
area of colonization.
Figure 4(c) and (f) indicate a clear difference between data
recorded by meteorological stations and data directly recorded in
the field. Forest temperatures were consistently lower than those
temperatures recorded by meteorological stations. This phenom-
enon highlights the importance of microclimate in forests, but
it does not affect the qualitative result of this study. Even if the
feeding activity calculated from meteorological station data is
overestimated, its temporal and spatial patterns give useful infor-
mation about the relative trend.
Air temperature in Paris is much higher than temperatures
anywhere in the south of the Paris Basin (see Appendix S3) and
this area is particularly favourable for PPM feeding activity (Fig.
4). This mild climate could be associated with the urban heating
and concentrated anthropogenic activity around Paris (Menut et
al.,
2000). However, one may wonder whether PPM populations
could have expanded so quickly without this effect. In this case,
extensive urbanization seems to enhance the PPM expansion
dynamics.
Even though this model reconstructed the feeding abilities
during the past few years, showing considerable changes between
1992-1996 and 2001-2004, we cannot conclusively evaluate the
impact of a warming trend from such short periods.
Changes in the indicator of feeding ability
The mean minimum temperature during the cold period is the
best predictor of feeding ability and its long-term observation
enables us to detect the effect of global warming on feeding activity
of the PPM. The sudden increase of this indicator in Orleans
took place just a few years before colonization by the PPM in
1992 (see Appendix S3). During the last few years, the indicator
in Melun has reached approximately the level recorded in
© 2007 The Authors
Global Ecology and Biogeography, 16,460-471, Journal compilation © 2007 Blackwell Publishing Ltd
Orleans when it had been colonized. Consequently, Melun can
now be considered potentially favourable to the establishment of
the PPM.
This sudden warming trend during the last 20 years is a global
event (National Assessment Synthesis Team of the US Global
Change Research Program, 2000). Half of the last century's tem-
perature rise has occurred since the late 1970s and 17 of the 18
warmest years in the 20th century occurred since 1980. Further-
more, the 1990s were the warmest decade in the observational
record (European Environment Agency, 2004).
The balance between climatic factors
Feeding failures are mostly due to a lack of heat during the
day (i.e. the maximum temperature inside the nest does not
reach the required threshold) according to field experiments
in the Paris Basin (Battisti et al., 2005). This lack could be
attributed to a low maximum air temperature or low solar
radiation (or both).
According to Huchon and Demolin (1970), the northward
distribution of the PPM in France would be limited mainly by
the sunshine barrier (threshold defined as a minimum of 1800
hours of sunshine). The sunshine records (Table 2) and the
climate scenario 'ARPEGE-Climat' give the evidence that solar
radiation has not changed and will not change considerably in
the future. In contrast, maximum temperature during the cold
period has increased by 0.04 °C year! in Orleans over the period
1972-2004, which is consistent with the general increase of
annual temperature observed in Europe between 1976 and
2000 (Watson et al., 2002). Thus, the sunshine barrier might
be of minor importance in northern France if the increase of
maximum temperature compensates for the low amount of
sunshine.
Changes in precipitation totals may also indirectly affect the
expansion. While southern Europe is likely to experience a severe
decrease in precipitation, northern Europe may face an increase
(European Environment Agency, 2004).
467
C. Robi net et al.

Consequences of extremely hot temperatures
Temperature increases are likely to improve PPM survival, espe-
cially the survival rate of founder populations in the expansion
areas (Battisti et al., 2005), and to enhance flying activity and
expansion (Battisti et al., 2004; Stastny et al., 2005). But increased
temperatures can also produce negative feedbacks. The excep-
tional heat that occurred during the summer of 2003 killed a
large proportion of PPM eggs and young larvae in the Paris Basin
(DSF, 2004; DSF Nord-Ouest, 2004). In this area, the PPM's pheno-
logical cycle does not enable the population to avoid extremely
hot periods during the summer as in southern (Mediterranean)
regions (Demolin, 1969a). But, due to the phenological plasticity
of the PPM, its life cycle can be expected to adapt to ongoing
climate warming (Benigni & Battisti, 1999). The role of a prolonged
diapause (Demolin, 1969a) in the persistence of the PPM is a
possibility that has not been investigated.
Extended model: its outcome and its restriction
The feeding duration is extended in the north-western quarter of
France where an eastward climatic gradient is prevalent. Some
caution in interpreting these results in mountainous areas is
needed because the model was initially built for the flat region
near the Paris Basin. The movement of the PPM front from
1970-80 to 2005 (Fig. 1) clearly shows a northward expansion
and also an expansion into higher elevations in the Massif Central
(in the south central part of France) and in the French Alps.
Topography strongly affects climatic conditions, and conse-
quently can also be expected to affect the presence of the PPM.
Hence, a specific topoclimatic model is necessary in order to
determine favourable areas for the PPM in mountainous terrain.
Trophic interactions involved in range expansion
The expansion of the PPM's range seems to be associated with
the increase of temperature since the mid-1980s (see Appendix S3),
but the analysis and prediction of responses to climate change
may be misleading if trophic interactions are ignored.
The results of modelling the potential feeding activity in
France at both local and regional scales, as well as the reconstruc-
tion of the feeding ability, indicate that the survival of the PPM
could have been possible in the past in some areas outside its
former range. The high number of potential feeding days, the
short periods of starvation and the low frequency of extreme
frost events suggest that the past PPM range in France was not
completely in equilibrium with the climatic conditions. Notably,
the effects of host plant availability and physical and behavioural
barriers to the dispersal of the PPM may indicate that the past
distribution of the PPM in France was not equivalent to its
potential climatic range.
French forests have greatly increased in area since the middle
of the 19th century. There were approximately 8.9 to 9.5 million
ha of forest in 1830. There are now 14.9 million hectares (source
French National Forest Inventory, http://www.ifn.frl). The range
of the main host plants of the PPM (Pinus spp.) covers a large
468 Global Ecology and Biogeography, 16, 460-471,
part of France, especially in the PPM's core area, but it also
extends beyond the present PPM distribution. A great trophic
plasticity enables the PPM to sometimes attack exotic conifer
hosts (Roques et al., 2002). The French forest area is still very
fragmented, especially in north-western and central France.
Since female dispersal capabilities are relatively low (maximum
of 2 km; Dernolin, 1969a), PPM expansion strongly depends on
the presence of relay trees (planted along the roads or in private
gardens) (Robinet, 2006).
Land use directly affects host-tree distribution and indirectly
affects dispersal and distribution of the PPM. Thuiller et al.
(2004) showed that a land-cover variable which was not explained
by climate could improve the prediction of species distribution,
but generally this was not the case in Europe. Due to the great
importance of isolated host pine trees and their widespread
occurrence over France and Europe, a land-cover variable may
not improve static models of PPM distribution. In contrast, the
inclusion of host-tree distribution and dispersal capabilities in a
dynamic model, such as the diffusion model (Robinet, 2006),
could improve the prediction of past and future PPM expansions.
Furthermore, climate change will undeniably affect host-tree
distributions. Some recent studies have predicted a significant
change in the distributions of European tree species in future
decades (Bakkenes et al., 2002; Thuiller, 2003; Thuiller et al., 2006b;
CARBOFOR project http://www.nancy.inra.fr/extranet/com/
carbofor / carbofor -D 1-resume.htm). Long-term modelling of the
distribution of the PPM should include this fundamental change.
Climate change will probably affect whole trophic interac-
tions. The consequences of climate change coupled with the
PPM's range shift on the natural enemies of the PPM are still
unknown, but they could in turn affect the control of the PPM.
CONCLUSIONS
Modelling of the feeding activity of the PPM revealed fundamen-
tal changes between past, present and future conditions that
could not be detected with a statistical model. This model pre-
dicted that the PPM would no longer be limited by the feeding
requirements in the Paris Basin and a large part of north -western
France may become favourable for the PPM in the future decades.
The distribution of the PPM in the Paris Basin could not have
been totally in equilibrium with past climatic conditions.
Climate change and the decrease in fragmentation of pine forests
contributed in concert to the range expansion. The effective shift
of the PPM's range will essentially depend on female dispersal
capacity and environmental heterogeneity. Future research
should focus on developing a comprehensive understanding of
how entire trophic chains will respond to global change.
ACKNOWLEDGEMENTS
We acknowledge financial support from the EU projects Promoth
QLK-CT-2002-00852 and ALARM GOCE-CT-2003-S0667S
(FP6 Integrated Project, Assessing large-scale environmental risks
with tested methods), and also from the Region Centre 'programme
de recherche inter-organismes et interdisciplinaires 2003 de la
© 2007 The Authors
Journal compilation © 2007 Blackwell Publishing Ltd
Effects of climate change on the pine processionary moth
Region Centre'. The climate scenario was kindly provided by
Meteo-France (Alain Braun and Emmanuel Cloppet). We thank
Andrea Battisti, Jacques Garcia, Francis Goussard, Andrew Lieb-
hold, Patrick Pineau, Jean-Paul Raimbault, Jerome Rousselet,
Daniel Sauvard and Regis Young for assistance and discussion.
We are grateful to Wilfried Thuiller and an anonymous referee
for their useful comments.
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Thuiller, w., Lavorel, S., Araujo, M.B., Sykes, M.T. & Prentice, I.e. bioclimatic modelling and risk assessment.
(2005) Climate change threats to plant diversity in Europe.
Josef Pennerstorfer holds a Master's degree in
Proceedings of the National Academy of Sciences of the USA, 102,
Geographic Information Systems from the Manchester
8245-8250.
Metropolitan University (UK) and is currently doing his
Thuiller, W., Broennimann, 0., Hughes, G.O., Alkemade, J.R.M.,
PhD in Land and Water Management and Engineering at
Midgley, G.F. & Corsi, E (2006a) Vulnerability of African
BOKU (Vienna, Austria).
mammals to anthropogenic climate change under conservative
land transformation assumptions. Global Change Biology, 12, Axel Schopf is head of the Institute of Forest
424-440. Entomology, Forest Pathology and Forest Protection
Thuiller, W., Lavorel, S., Sykes, M.T. & Araujo, M.B. (2006b) (BOKU, Vienna, Austria) with a research focus on
Using niche-based modelling to assess the impact of climate physiological ecology and host-parasitoid interactions.
change on tree functional diversity in Europe. Diversity and
Alain Roques is the head of the Forest Zoology Unit
Distributions, 12,49-60.
(INRA, Orleans, France). He studies the biology and
Visser, M.E., Adriaensen, E, Van Balen, J.H., Blondel, J., Dhondt,
population dynamics of forest insect pests and
AA., Van Dongen, S., Du Feu, c. Ivankina, E.V., Kerimov,
A.B., De investigates the impact of insect invasion and expansion
Laet, J., Matthysen, E., McCleery, R, Orell, M. & Thomson, D.1. related to human activity.
(2003) Variable responses to large-scale climate change in
European Parus populations. Proceedings of the Royal Society of
Editor: Jose Alexandre F. Diniz-
London Series B, 270, 367-372. Filho
Walther, G.-R, Post, E., Convey, P., Menzel, A., Parmesan, c.,
Beebee, T.J.C., Frometin, J.-M., Hoegh-Guldberg, o. &
Barlein, F. (2002) Ecological responses to recent climate SUPPLEMENTARY MATERIAL
change. Nature, 416, 389-395.
The following supplementary material is available for this article:
Warren, M.S., Hill, J.K., Thomas, J.A, Asher, J., Fox, R., Huntley,
B., Roy, D.B., Telfer, M.G., Ieffcoate, S., Harding, P., Jeffcoate, Appendix S1 Modelling nest temperature
G., Willis, S.G., Greatorex-Davies, J.N., Moss, D. & Thomas,
Appendix S2 Estimation of solar radiation
C.D. (2001) Rapid responses of British butterflies to
opposing forces of climate and habitat change. Nature, 414, 65- Appendix S3 Long-term fluctuation in the indicator of feeding
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Watson, R.T. (ed.) (2002) Climate change 2001: synthesis report.
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A contribution of Working Groups I, II and III to the Third
http://www.blackwell-synergy.com/doi/absIl0.llll1j.1466-
Assessment Report of the Intergovernmental Panel on Climate
8238.2006.00302.x
Change. Cambridge University Press, Cambridge.
Watson, RT., Zinyowera, M. & Moss, RH. (1998) The (This link will take you to the article abstract).
regional impacts of climate change, an assessment of Please note: Blackwell Publishing is not responsible for the
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