Forest Ecology and Management 249 (2007) 171–186

www.elsevier.com/locate/foreco

PHENIPS—A comprehensive phenology model of Ips typographus (L.)

(Col., Scolytinae) as a tool for hazard rating of bark beetle infestation
Peter Baier *, Josef Pennerstorfer, Axel Schopf
Institute of Forest Entomology, Forest Pathology and Forest Protection, Department of Forest and Soil Sciences,
University of Natural Resources and Applied Life Sciences, Hasenauerstaße 38, A-1180 Vienna, Austria
Received 1 June 2006; received in revised form 24 January 2007; accepted 5 May 2007

Abstract
We developed the model PHENIPS for spatial and temporal simulation of the seasonal development of Ips typographus at the Kalkalpen
National Park in Austria. The model is based on a digital elevation model used for interpolation of temperature and solar radiation to calculate the
microclimatic conditions (bark temperature) for the beetles’ development. Additionally, the beetles’ phenology at Kalkalpen National Park was
monitored along with air and bark temperature measurements. The onset of host tree infestation in spring was estimated using a lower threshold of
16.5 8C for flight activity and a mean thermal sum of 140 degree-days (dd) from beginning of April 1st onward. Rate of brood development was
calculated from accumulated degree-days of hourly temperature data using upper and lower temperature thresholds of 38.9 and 8.3 8C,
respectively, and a nonlinear function for calculating effective thermal sums. Re-emergence of parental beetles occurred at a time when
49.7% of the thermal sum for total development (557 dd) was reached. The model includes the discontinuance of the beetle’s reproductive activity
at a day length <14.5 h. The rate of successful hibernation of established broods is predicted by assessing the developmental stage of initiated
generations at the beginning of the cold period. For validation we compared the timing of phenological events in the field with predicted events
using both, hourly recorded data at trap trees in the terrain and generated daily topoclimatic data. Using topoclimatic data, the onset of infestation
was predicted with a mean absolute error of 1.3 days. The observed onset of emergence of filial beetles in the field was estimated with a mean error
of 39 dd. Our PHENIPS explicitly considers the strong effects of regional topography and stand conditions on local air and bark temperature and
can be used for precise monitoring of the actual state of bark beetle development at the specific stand/tree level. Using topoclimatic data, PHENIPS
simulates the maximum number of generations which is necessary to assess the potential impact of bark beetle outbreaks at regional scale. Further
applications of PHENIPS for site-specific hazard rating of bark beetle infestation are discussed.
# 2007 Elsevier B.V. All rights reserved.

Keywords: Ips typographus; Scolytinae; Phenology; Hazard rating; Forest protection; Topoclimatic modelling; GIS

1. Introduction
Bark beetle outbreaks are important biotic disturbances that
occur within the natural development and dynamics of forest
ecosystems. Eruptive outbreaks of the European spruce bark
beetle, Ips typographus (L.), result in mass attacks of living
trees and may cause tree mortality at landscape-levels.
Outbreaks of this insect are triggered mainly by heavy storm
events and favourable climatic conditions. Increasing damage
in recent years in Central Europe caused an enormous
economic impact on forestry due to timber losses and high
expenditures for control and sanitation measures (Forster, 1993;
* Corresponding author. Tel.: +43 1 3686352; fax: +43 1 3686352 97.
E-mail address: baier@ento.boku.ac.at (P. Baier).
0378-1127/$ – see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2007.05.020
Schopf and Köhler, 1995; Grodzki, 1999; Turcani and Novotny,
1999; Heurich et al., 2001; Schelhaas et al., 2003). In order to
assess the likelihood of a mass outbreak in time, appropriate
monitoring tools are necessary to accurately predict the number
of generations and the actual developmental process of the bark
beetle population. This is especially important for those areas in
alpine terrain which are difficult to access for pest control, but
have favourable climatic conditions for insect development.
A basic requirement for mapping temperature-dependent
phenology of this bark beetle is the precise spatial interpolation
of point station temperature data across the landscape. Various
approaches for estimating spatial and temporal patterns of
climatic parameters have been tested and applied at different
time scales and spatial resolutions (Russo et al., 1993; Bolstad
et al., 1997; Thornton et al., 1997; Price et al., 2000; Antonić
et al., 2001; Xia et al., 2001; Jarvis et al., 2003). In this study,
172 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186
we established a topoclimatic model based on a digital
elevation model (DEM) and air temperatures recorded from
several weather stations located in the field. The method for
estimating daily air temperature is a common approach based
on the adiabatic lapse rate of temperature using daily multiple
regression analysis (Russo et al., 1993). In complex terrain, the
temperature of the near surface air layer is influenced mainly by
topographic characteristics such as elevation, slope, and aspect
as well as by the adiabatic changes of temperature with
elevation. For the time period critical for bark beetle
development at mid latitudes (April–October), the simple
daily regression analysis of this linear relation between air
temperature and elevation can explain a large part of the spatio-
temporal variability in air temperature. However, to simulate
the temperature condition inside the bark, we had to include the
effect of insolation which accounts for an additional input of
energy by surface heating and essentially contributes to
elevated bark temperatures at sun exposed sites (Bolstad
et al., 1997). Seasonal variability of site-related potential solar
irradiation was defined by topographic sky obstruction,
astronomical, and trigonometric patterns. Site-specific poten-
tial solar radiation of inclined surfaces was estimated by using a
DEM-based geometric radiation model, implemented as a tool
in GIS (Kumar et al., 1997).
Several studies have been conducted in the past to examine
the influence of temperature on the development and
reproduction of I. typographus (Vité, 1952; Annila, 1969;
Zumr, 1982; Anderbrant, 1986). They revealed a develop-
mental pattern of I. typographus that is more complex than for
example the life cycle of the similarly aggressive American
bark beetle species, Dendroctonus ponderosae (Bentz et al.,
1991). In contrast to this univoltine, non-diapausing species, I.
typographus shows a multivoltine life cycle in most parts of
Central Europe with a photoperiodically controlled reproduc-
tive diapause in its adult stage which is also the hibernating
stage (Schopf, 1989; Doležal and Sehnal, 2003). Furthermore,
parental females produce sister broods which considerably
contribute to its population growth (Anderbrant, 1986, 1989;
Kritsch, 2005).
The precise knowledge of the rate and the thermal
thresholds of the beetle’s development (Netherer, 2003;
Wermelinger and Seifert, 1998) are prerequisites to establish
a phenology model. In previous studies, only the linear relation
between developmental rate and ambient temperature above
the lower developmental threshold were used to estimate
degree-days required to complete its development (Coeln
et al., 1996; Führer and Coeln, 1998; Pennerstorfer, 2000;
Netherer and Pennerstorfer, 2001; Netherer, 2003). In this
study, we developed the phenology model PHENIPS which is
based on a nonlinear function for calculating effective thermal
sums as suggested by Wermelinger and Seifert (1998).
Furthermore, we included the prediction of the time of spring
swarming, onset of infestation, re-emergence of parental
beetles, and the number and emergence time of regular and
sister broods which are limited by a photoperiodic threshold
(Doležal and Sehnal, 2003). Model validation was done by
comparing recorded bark temperature, flight activity, and
developmental progress in trap trees at various altitudes and
aspects of sloping ground with predicted phenological events
by the model.
2. Material and methods
2.1. Field studies
Field experiments were conducted at the Kalkalpen
National Park, located in the northern range of the Limestone
Alps in Upper Austria. Time of spring swarming of I.
typographus was recorded during 3 consecutive years (2001–
2003) using flight-barrier traps (Friedrich Theysohn Kunst-
stoff GmbH, Salzgitter, Germany) baited with PHERO-
PRAX1 (Shell Agrar GmbH, Ingelheim, Germany). The
traps were placed at four sites in 2001 (Messerer, Bodingbauer,
Zaglbaueralm and Ebenforstalm) and three sites in 2002–2003
(Krennbauer, Mehlboden and Haslersgatter) at different
elevation (Table 1). We counted the number of captured
beetles at 2–3 day intervals.
In order to observe the onset of tree colonization, brood
development, and presence of parental beetles, we used trap
trees felled at different sites in two series in 2001 and 2002
(April/May and mid to end of July) and only in one series in
2003 (Table 1). Newly bored entrance holes were marked with
coloured tacks at the lower (3/10 of tree height), middle
(crown base), and upper part (mid crown) of felled trap trees. At
weekly intervals, marked breeding systems were inspected by
removing pieces of bark (approx. 20 cm  20 cm). The most
advanced developmental stage of offspring and the presence/
absence of parental beetles were recorded. As soon as the brood
had reached the pupal or teneral adult stage, a log (length:
70 cm) was removed at the crown base and transferred into
photoeclectors located in the garden of the Institute for daily
observation of emerging filial beetles.
The onset of offspring’s emergence in the field was
checked in a weekly interval by inspecting the trap trees for
exit holes on the bark surface. In October, the number of
remaining young adults in the bark of the trap trees was
determined. Beetles emerging from the logs in photoeclectors
were collected daily until emergence ceased in the following
spring. Only logs from trap trees in 2003 were kept in the
laboratory at 23  1 8C and long-day photoperiods of 16 h
light:8 h dark from October 4, 2003 onwards, in order to
induce the emergence of the remaining young beetles from
their brood system. We calculated the ratio of beetles that left
their logs within the same season to the number of beetles that
emerged in total.
Air temperature at the sites of pheromone traps, trap trees,
and photoeclectors was hourly recorded using Gemini
TinytagPlus data loggers TGP-0017 (Gemini Data Loggers
Ltd., Chichester, UK) placed into a radiation shield at 1.5 m
above ground. Likewise, we hourly recorded the bark
temperature at the crown base of each trap tree at four
positions around the trunk (top, bottom, and lateral sides) using
Ni100-sensors connected to a MiniCube VV/VX data logger
(EMS, Brno, Czech).
 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186 173

Table 1
Location, elevation, aspect, and global site factor (GSF: mean value April–October) for each trap tree with the date of felling, infestation, and observed date of
emergence of filial beetles in the field and from logs in photoeclectors
Location, trap tree series, Elevation Coordinates Aspect GSF (%) Date of felling Onset of Onset of emergence
and year (m a.s.l.) infestation
East North Field Photoeclector
First series 2001
Messerer 580 14.37856 47.80929 NE 10.1 24/04/2001 2/05/2001 31/07/2001 27/07/2001
Bodinggraben 640 14.39343 47.78984 N 42.0 24/04/2001 4/05/2001 31/07/2001 27/07/2001
Zaglbaueralm 940 14.37386 47.79756 S 59.9 24/04/2001 10/05/2001 31/07/2001 27/07/2001
Ebenforstalm 1100 14.42279 47.80468 Level 69.1 18/05/2001 25/05/2001 24/08/2001 30/07/2001
Blumaueralm 820 14.35117 47.78849 S 44.2 18/05/2001 25/05/2001 7/08/2001 7/08/2001
Tiefling 1100 14.38541 47.77244 NE 12.4 18/05/2001 31/05/2001 24/04/2002
Second series 2001
Messerer 580 14.37856 47.80929 NE 29.0 16/07/2001 Not infested
Bodinggraben 640 14.39343 47.78984 N 30.3 16/07/2001 Not infested
Zaglbaueralm 940 14.37386 47.79756 S 66.6 25/07/2001 27/07/2001 2/05/2002
Ebenforstalm 1100 14.42279 47.80468 Level 73.2 16/07/2001 23/07/2001 29/04/2002
Blumaueralm 860 14.34894 47.78935 S 16.5 31/07/2001 2/08/2001 7/05/2002
First series 2002
Krennbauer 726 14.51786 47.69994 NNE 23.8 30/04/2002 6/05/2002 1/08/2002 17/07/2002
Holzgraben I 1050 14.47703 47.71342 S 57.7 6/05/2002 8/05/2002 15/07/2002 11/07/2002
Holzgraben II 1047 14.46958 47.70942 N 32.4 6/05/2002 8/05/2002 19/07/2002 17/07/2002
Mehlboden 860 14.35042 47.75467 Level 10.6 30/04/2002 10/05/2002 15/08/2002 3/08/2002
Haslersgatter 1167 14.37939 47.73847 NE 15.9 30/04/2002 18/05/2002 28/04/2003
Weingartalm 1200 14.41797 47.75683 SW 56.2 15/05/2002 20/05/2002 15/08/2002 28/04/2003
Second series 2002
Krennbauer 647 14.51581 47.70294 N 17.9 15/07/2002 30/07/2002 2/05/2003
Holzgraben I 1050 14.47703 47.71342 S 45.4 15/07/2002 30/07/2002 30/04/2003
Holzgraben II 1047 14.47208 47.70914 N 30.8 15/07/2002 30/07/2002 30/04/2003
a
Mehlboden 860 14.35042 47.75467 SE 23.0 15/07/2002
a
Haslersgatter 1167 14.37939 47.73847 NE 13.2 15/07/2002
Weingartalm 1200 14.41797 47.75683 SW 64.3 15/07/2002 30/07/2002 30/04/2003
2003
b
Krennbauer 647 14.51581 47.70294 N 48.7 8/05/2003 11/07/2003 9/07/2003
b
Holzgraben I 1050 14.47703 47.71342 S 51.4 30/04/2003 11/07/2003 9/07/2003
Haslersgatter 1167 14.37939 47.73847 NE 13.1 9/05/2003 4/06/2003 19/08/2003 1/08/2003
Weingartalm 1200 14.41797 47.75683 SW 44.6 9/05/2003 24/05/2003 24/07/2003 21/07/2003
a
Only sparsely infested.
b
Wind-felled trees.

2.2. Topoclimatic modelling at reference stations (Schoberstein in 2000–2002; Steyregg in

Topographic parameters (slope, aspect, and elevation) were
derived from a digital elevation model (DEM) for the region of
the Kalkalpen National Park with a resolution of 50 m.
Topoclimatic modelling was done for a synoptic grid resolution
of 300 m in which the maximum of potential radiation and the
minimum of elevation of each grid cell were selected as basic
input parameters for topoclimatic modelling. This modelling
was restricted to the major period for bark beetle development
(April–October). Outside this period, the temperatures are
frequently lower than the lower developmental threshold of I.
typographus in the study area. Furthermore, the applied linear
lapse rate model for estimating air temperature is not practical
for use during the winter months due to the frequent occurrence
of temperature inversions.
2003; Table 2). For more than four missing hourly observations
per day, the daily sum was replaced by the monthly mean.
At each trap tree and for the site of solar radiation
measurement in the field (location ‘‘Weingartalm’’), we
analysed hemispherical photos by means of the program
HEMIVIEW 2.0 (Delta-T Devices Ltd., Cambridge, UK). The
calculated monthly mean global site factor (GSF) is the
proportion of global radiation under a tree canopy relative to
that in the open, and includes sky obstruction by topography
and by vegetation (Table 1). Modelling of the bark temperatures
at the sites of trap trees was developed using the observed daily
sum of solar radiation at the reference station reduced by the
monthly GSF at each trap tree (Eq. (1)).
SIredðxl Þi ¼ SIðrefÞi  GSFðxl Þm (1)

2.2.1. Solar irradiation modelling SIred(xl)i is the daily sum of solar radiation reduced with the
For estimating the daily sum of solar radiation in the field, GSF at location xl for day i; SI(ref)i the observed global solar
we used hourly recorded horizontal global irradiation measured radiation at the reference station for day i; GSF(xl)m is the
174 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186

Table 2
Location, recorded parameters (AT: air temperature, SI: solar irradiation), elevation, and number of valid observations of the recording stations at the study area
Recording station Coordinates Recorded parameter Elevation (m a.s.l.) Valid n
East North
Brauneben 14.22149 47.82085 AT 760 846
Ebenforstalm 14.42353 47.80652 AT 1050 792
Feichtaualm 14.32398 47.80247 AT 1370 746
Hengstpass 14.46202 47.70315 AT 990 726
Kogleralm 14.26414 47.77851 AT 1240 856
Mitterweng 14.35668 47.69964 AT 720 691
Rettenbach 14.31615 47.75531 AT 610 824
Rotwagmauer 14.39595 47.79598 AT 750 826
Saubachgut 14.25053 47.76350 AT 875 707
Schoberstein 14.32472 47.90583 AT/SI 1260 507/515
Wurbauer 14.35121 47.72667 AT 850 672
Zaglbaueralm 14.37533 47.79977 AT 990 856
Steyregg 14.35278 48.29028 SI 335 214

‘‘global site factor’’ of the location of trap tree xl during the
month m.
For spatial extrapolation of daily solar irradiation, we
calculated the ratio of observed versus potential solar
irradiation (Kumar et al., 1997) at the reference station in
relation to the potential solar radiation of any location in the
field (Appendix A.1).
2.2.2. Air temperature
Daily mean and maximum air temperatures were estimated
using daily regression analysis (Bolstad et al., 1997; Russo
et al., 1993) that incorporated elevation and relative solar
irradiation (SIrel; Eq. (A.1)) as predictive variables. We
computed for each day a unique set of regression coefficients
(ai, bi, ci) based on aggregated hourly temperature data from 12
stations recorded from April until October in 2000–2003
(Table 2). Estimates of maximum and mean air temperatures
were then calculated using the appropriate daily regression
coefficients, elevation, and estimated solar radiation as input
variables (Eq. (A.2)).
(DTL = 8.3 8C), and a nonlinear function as proposed by
Wermelinger and Seifert (1998).
Effective temperatures for the development from egg stage
to emergence of mature filial beetles were adapted using the
optimum temperature (TO = 30.4 8C), and the lower
(DTL = 8.3 8C) and upper developmental thresholds
(DTU = 38.9 8C) after Wermelinger and Seifert (1998). The
effective temperature (Teff) was estimated using the nonlinear
function of the modified Logan model (Logan et al., 1976;
Lactin et al., 1995) with the assumptions that the relative
developmental rate increases linearly within the range of DTL
and TO and that the relative developmental rate is equal to 1 at
TO. For temperatures outside the lower and upper temperature
thresholds (T  DTL and T  DTU), the Teff was set to zero.
The effective temperature can be calculated with Eq. (2).
T eff ¼ ðT O  DTL Þ  ðeaT  eðaT max ðT max TÞ=bÞ  gÞ (2)
a = 0.02876507; b = 3.5922336; g = 1.24657367;
Tmax = 40.9958913.

2.3. Modelling phenology of I. typographus

2.3.1. Modelling spring swarming and onset of infestation

To predict the onset of spring swarming and infestation we
compared the observed onset of trap catches and trap tree
infestation with the observed thermal conditions for flight
activity. Additionally, the observed onset of spring swarming
and of host tree colonization by I. typographus was compared
with the cumulative thermal sum of maximum air temperature
above the beetles’ lower developmental threshold
(DTL = 8.3 8C; Wermelinger and Seifert, 1998) accumulated
from the first of April onwards using estimated topoclimatic
temperature data.

2.3.2. Modelling brood development

2.3.2.1. Parameterisation of the bark beetle developmental
model. In order to describe the relation between develop-
mental rate of I. typographus and temperature we used both a
linear function with only a lower developmental threshold
Fig. 1. Relationship between naturally occurring temperatures (T) and effective
temperatures for development of I. typographus (Teff non linear: effective
temperature according to Eq. (2); Teff linear: effective temperature of the linear
model using only the lower developmental threshold).
 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186
The parameters a, b, g, and Tmax of the Eq. (2) were
estimated iteratively by means of a nonlinear regression
analysis for temperatures near the threshold values and within
the range of DTL and TO and their corresponding relative
developmental rates (Fig. 1).
A thermal sum (K) of 334.2 dd is necessary for I.
typographus to complete the preimaginal development (egg
to pupal stage) of the beetle (Wermelinger and Seifert, 1998),
and 222.8 dd (i.e. two third of the thermal sum for preimaginal
development) is required for maturation feeding of the filial
beetles (Wermelinger and Seifert, 1998; Netherer, 2003).
Hence, the thermal sum of 557 dd is required for total
development of I. typographus.
2.4. Modelling bark temperature and effective daily
thermal sums
Modelling of the bark temperature at the upper side of the
bole is based on hourly recorded data measured in the bark of 27
trap trees (Table 1). To estimate the daily mean and maximum
bark temperature we used multiple regression analysis with
estimated solar radiation (SIred) and observed air temperature
as predictive variables.
Since the output of the topoclimatic model provides only daily
meteorological parameters, the calculation of the effective bark
temperature (Eq. (2)) was adapted for daily input data (estimated
daily mean and maximum bark temperature). Within the range of
DTL and TO, the daily effective thermal sum for bark beetle
development was calculated from the difference between the
daily mean bark temperature and the lower developmental
threshold (DTL = 8.3 8C). To avoid overestimating the effective
thermal sum at bark temperatures above the optimum
temperature for beetle’s development (TO = 30.4 8C), we
corrected the estimated daily thermal sums by calculating the
difference between the effective bark temperature sum of the
linear model and the nonlinear model. The correlation between
daily maximum bark temperature and this difference was used
for the correction (see Appendix A).
2.5. Simulation and mapping of the phenology using
PHENIPS
Spatial modelling and mapping of phenology and potential
development of I. typographus at the study area was conducted
for the period 2000–2003. For each grid cell (300 m  300 m;
n = 17,548), the onset of infestation and the numbers of
generations were calculated based on estimated daily solar
radiation, daily mean and maximum air temperature, and daily
thermal sum of effective bark temperature (see Appendix A).
Since we used the maximum incoming radiation for computation
of temperatures without consideration of shading by the canopy,
the maximum number of possible generations is simulated.
2.6. Model validation
To validate the estimated mean and maximum air
temperature, we used independently recorded air temperature
175
data, measured at each trap tree. Validation of time series of
observed and estimated temperatures was done by computing
the mean error (ME), the mean absolute error (MAE), and the
root mean square error (RMSE). For validation of the solar
irradiation model, daily solar irradiation was measured during
the period 3/7/2003 to 30/9/2003 at the location Weingartalm
using a star pyranometer (Schenk, Vienna, Austria). Thermal
sums of estimated bark temperatures were validated using the
relative ratios of predicted and observed sums (relative
percentage mean error).
To determine the accuracy of the developmental model we
compared the effective thermal sum (computed with Eq. (2)
using hourly recorded bark temperatures) at the date of beetle
emergence with the thermal sum required for total development
of one generation (K = 557 dd). For this test we used only trap
trees of the first series that contained complete generation
development within the same season.
For validation of the predicted onset of re-emergence of
parental beetles and emergence of filial beetles calculated from
generated topoclimatic data, we compared the estimated
effective thermal sum at the date of observed re-emergence/
emergence given by the model with the corresponding required
thermal sums (K = 278.5 dd for re-emergence; K = 557 dd for
emergence of filial beetles).
2.7. Statistics, data management, and GIS
Statistical analyses were performed using the statistical
package SPSSTM12.0.1 (SPSS Inc., 1990). Linear relations
were analysed using multiple, linear regression analysis; for
nonlinear relations, the NLR-regression procedure was used.
One-sample Kolmogorov–Smirnov test was applied to verify
normal distribution of test variables. Pairs of means were
compared using the paired samples t-test. Deviations of the
mean of a single variable from a specified test value were tested
by the one-sample t-test procedure.
We used Microsoft SQL Server 2000 as a database program
for topoclimatic modelling and calculation of thermal sums for
predicting phenological events. Mapping was done with the
Esri GIS-Software ArcView 3.2.a and ArcGis 8.3.
3. Results
3.1. Topoclimatic model
3.1.1. Modelling of solar irradiation
For validating estimated solar radiation, we compared the
global solar irradiation recorded at the location Weingartalm
(1200 m a.s.l., south exposed, mature Norway spruce stand) with
the calculated values of the model. The location ‘‘Weingartalm’’
showed a low obstruction by topography and seasonal shading by
the canopy with higher GSF-values during months when the
position of the sun is lower in the horizon. The high accuracy
attained by using the GSF to estimate the daily sum of solar
radiation (Eq. (1)) is indicated by the small deviations from the
recorded solar radiation (mean error = 169.56 Wh m2;
relative mean error = 7.06%; n = 90). Higher deviations
176 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186
Fig. 2. Observed and predicted daily sum of solar radiation at the location
Weingartalm (SIobs: observed solar irradiation; SIred: estimated solar irradia-
tion using GSF (Eq. (1)); SIrel: estimated solar radiation reduced by topo-
graphic obstruction (Eq. (A.1)); SIref: observed radiation at the reference
station).
occurred only when the atmospheric conditions at the reference
station were different from the conditions recorded locally
(Fig. 2).
3.1.2. Modelling of air temperature
Daily mean and maximum air temperature were interpolated
by means of daily regression analysis with elevation and
relative solar radiation (SIrel, Eq. (A.1)) as predictive variables
The estimated mean and maximum air temperatures for the
various recording stations (Table 2) correlated significantly
with the observed values (P < 0.001). Goodness of fit (R2) for
the different years (2000–2003) ranged between 0.957 and
0.987.
We validated the estimated air temperatures by using
temperature data recorded independently at the locations of trap
trees. The estimated daily mean and maximum air temperature
showed a root mean square error of 1.26, and 2.06 8C,
respectively. The grand mean of the ratio between the sum of
estimated versus recorded daily mean and maximum air
temperatures was 99.23 and 99.13%, respectively.
3.1.3. Modelling of bark temperature and daily effective
thermal sum
The observed bark temperatures at the trap trees showed
highly significant relations to ambient temperature and solar
radiation. The daily mean bark temperature can be estimated
with solar irradiation (SIred) and daily mean air temperature as
predictive variables (R2 = 0.961, P < 0.001). The daily max-
imum bark temperature showed a nonlinear relation to the air
temperature maximum with incoming radiation used as an
additional predictive variable (R2 = 0.851, P < 0.001). Based
on these results, the daily mean and maximum bark
temperatures have been estimated using the equations given
in Appendix A (Eqs. (A.3) and (A.4)).
Daily effective thermal sums were calculated with the
procedures described in Appendix A (Eqs. (A.5)–(A.7)). The
daily thermal sum showed a mean absolute error of 0.9 dd
(minimum: 0 dd; maximum: 4.8 dd). The correlation between
estimated and observed daily thermal sum was highly
significant (R2 = 0.9104, P  0.001). Likewise, the estimated
thermal sums accumulated over the entire study period for each
trap tree, showed a highly significant relation (R2 = 0.9671,
P  0.001) with the observed thermal sum (calculated with
Eq. (2) based on hourly recorded bark temperature data).
Cumulative observed and estimated thermal sums did not differ
significantly (paired-samples t-test; mean difference: 6.28 dd;
t = 0.671; d.f. = 26; P = 0.508). The ratio between estimated
and observed thermal sum was on average 101.3%  9.55%.
3.2. Bark beetle phenology model
3.2.1. Spring swarming and onset of infestation
Bark beetles were captured in pheromone traps when the
daily maximum air temperature (ATmax) exceeded 16.5 8C.
Major flight activities (>10% of the total number of beetles
trapped until end of May) occurred on days when ATmax was
above 19.4 8C.
First attacks of I. typographus on trap trees in spring were
observed at a mean ATmax of 21.5  3.33 8C (lowest ATmax:
16.8 8C). Initial infestation of the trap trees occurred in 7 out of
9 cases within 3 days after the first main flight period (Table 3).
In only one case, beetles infested a trap tree prior to beginning
of the first main swarming period (trap tree Mehlboden, 2002).
Besides the lower air temperature thresholds for flight
activity and initial attacks on trees (ATmax > 16.5 and
>16.8 8C, respectively), which may occur occasionally on
warmer days during the winter period, other factors might
prevent hibernating beetles to resume their reproductive
activities too early in the season. Our observations indicate
that flight activity initiated when the thermal sum (ATmax
above >8.3 8C) accumulated from the first of April onwards
was on average 60.54  19.18 dd. First colonization of trap
trees was observed when the thermal sum reached
140.26  23.68 dd.
Using the accumulated thermal sum in combination with the
flight activity threshold, the onset of spring swarming was
estimated with a mean absolute error of 2.4 days, the start of
trap tree colonization in spring can be estimated with a MAE of
1.3 days (Table 3). The difference between the observed and
predicted onset of attacks of 13 trap trees was 1 day earlier at
the minimum and 8 days later at the maximum (Table 3).
3.2.2. Brood development
Generally, trap trees of the first series felled in spring 2001–
2003 showed an intense and rapid colonization each year. Only
two of 16 trap trees located at shaded high elevation sites
(Tiefling, 2001 and Haslersgatter, 2002) were not rapidly
colonized within 1 week. The period from the onset of
infestation until complete colonization of the whole tree lasted
from May 18 to July 31, 2001 at Tiefling, and from May 16 to
June 24, 2002 at Haslersgatter. Trap trees of the second series
felled in mid July, were not colonized at the locations
Bodinggraben and Messerer in the year 2001; those at the
locations Mehlboden and Haslersgatter were only sparsely
 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186 177

Table 3
Recorded onset of spring swarming, first main flight period, onset of infestation of trap trees, air temperature maximum (ATmax) during the period of first infestation,
estimated onset of swarming and infestation, and minimum absolute deviation of estimated onset from recorded data (MAE: mean absolute error of estimated onset of
swarming and infestation)
Location of Onset of swarming Onset of first main Infestation a ATmax (8C) Estimated onset Minimum abs.
trap/trap tree flight period deviation (days)
Swarming Infestation Swarming Infestation
Messerer 24–26/04/2001 30/04/2001 to 30/04/2001 to 25.83 25/04/2001 02/05/2001 1 0
02/05/2001 02/05/2001
Bodinggraben 24–26/04/2001 02–04/05/2001 02–04/05/2001 28.61 25/04/2001 02/05/2001 1 0
Zaglbaueralm 24–26/04/2001 02–04/05/2001 06–10/05/2001 16.79 29/04/2001 05/05/2001 3 1
b
Ebenforstalm 30/04/2001 to 02–04/05/2001 22–25/05/2001 21.54 30/04/2001 14/05/2001 0
02/05/2001
Krennbauer 27–29/04/02 30/04/2002 to 04–06/05/2002 19.12 26/04/2002 03/05/2002 1 1
2/05/2002
Mehlboden 30/04/2002 to 16–18/05/2002 08–10/05/2002 19.54 01/05/2002 08/05/2002 1 0
02/05/2002
Haslersgatter 06–08/05/2002 16–18/05/2002 16–18/05/2002 20.66 16/05/2002 18/05/2002 8 0
Krennbauer 20/04/2003 25–30/04/2003 30/04/2003 to 24.86 28/04/2003 06/05/2003 8 4
02/05/2003
Mehlboden 21–26/04/2003 29/04/2003 to 26/04/2003 0
02/05/2003
Haslersgatter 29/04/2003 to 04–06/05/2003 07–09/05/2003 18.34 03/05/2003 09/05/2003 1 0
02/05/2003
b
Blumaueralm 18–31/05/2001 23.48
b
Tiefling 18–31/05/2001 19.06
Weingartalm 16–20/05/2002 22.75 09/05/2002 17/05/2002 1
Holzgraben I 07–15/05/2002 18.78 01/05/2002 16/05/2002 1
Holzgraben II 07–15/05/2002 17.61 07/05/2002 17/05/2002 2
Weingartalm 07–09/05/2003 24.29 29/04/2003 06/05/2003 3
Holzgraben I 26–30/04/2003 22.66 25/04/2003 04/05/2003 4
Minimum 1 4
Mean ATmax 21.50 Maximum 8 3
Minimum ATmax 16.79 MAE 2.4 1.31
a
In 2003, the infestation of trees occurred before we felled trap trees. Therefore, the onset of colonization was assessed on infested trees adjacent to the locations of
our trap trees.
b
Trap trees, felled later in the season have not been included.

infested in the year 2002. In 2003, we did not fell a second
series of trap trees.
For validation of the maximum rate of brood development
we compared the observed onset of emergence of the filial
beetles with the thermal sum of hourly measured bark
temperatures at the upper position of the trunk (Eq. (2)).
The thermal sum at the time of emergence was not significant
different from the test value (K = 557 dd) for both emergence in
the field and emergence from logs in photoeclectors (P < 0.05,
Table 4). The mean deviation of the calculated thermal sum
from the required thermal sum (557 dd) was 15.9 dd in the field
and 5.1 dd for logs in photoeclectors, respectively (Table 4).
3.2.3. Status of brood development at the beginning of the
cold period
For successful hibernation, the brood of I. typographus
must complete the preimaginal development (egg–pupae)
(i.e. it requires 60% of the thermal sum for total
development) before the beginning of the cold period.
Based on this ratio, successful hibernation of filial
generations and sister broods can be predicted at the end
of the season. For validation, the observed developmental
stage of the brood in trap trees of the second series was
compared with the predicted stage by the model in October
(Table 5).

Table 4
Mean value, standard deviation (S.D.), test statistics (one-sample t-test with test value K = 557 dd), relative mean error (RPME), and mean difference from K (ME) of
the estimated thermal sum at the date of observed emergence of filial beetles from trap trees in the field and for logs in photoeclectors
Emergence Mean value (dd) S.D. (dd) t-Value d.f. P RPME (%) ME (dd)
In the field (hourly recorded data) 572.9 55.82 1.071 13 0.304 2.85 15.9
From photoeclectors (hourly recorded data) 562.1 43.02 0.428 12 0.676 0.92 5.1
In the field (estimated by topoclimatic data) 595.76 92.09 1.575 13 0.139 6.96 38.8
178 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186

Table 5
Observed developmental stage, relative thermal sum, and predicted developmental stage estimated with observed and calculated bark temperatures in October for trap
trees, felled in July 2001 and 2002
Year Location Date of trap tree Observed Relative thermal Predicted stage (estimated with Predicted stage (estimated
control stage sum (%) hourly observed data) with topoclimatic data)
2001 Zaglbaueralm 9/10 Young adults 83.15 Young adults Young adults
Blumaueralm 9/10 Pupae 61.62 Young adults Young adults
Ebenforstalm 9/10 Young adults 80.18 Young adults Young adults
2002 Krennbauer 8/10 Young adults 67.90 Young adults Young adults
Holzgraben I 8/10 Young adults 70.84 Young adults Young adults
Holzgraben II 8/10 Pupae 55.22 Pupae Pupae
Weingartalm 8/10 Pupae 56.00 Pupae Young adults

Except for one case (trap tree Blumaueralm 2001), the
observed developmental stage in October was accurately
predicted by the model. In the trap tree ‘‘Blumaueralm’’, we
recorded 62% of the thermal sum for total development in
October and predicted the developmental stage as immature
adults while the inspection revealed only the development into
the pupal stage (Table 5). Some of these individuals, however,
were still able to eclose later and successfully hibernated as
adults. As predicted by the relative thermal sum received by the
brood in trap trees of the second series at Holzgraben II and
Weingartalm 2002 (<60%), the larvae developed only to the
pupal stage by October and thus were not able to hibernate
successfully. Only parental beetles re-emerged from these logs
when placed into outdoor photoeclectors in the spring 2003.
3.2.4. Development of sister broods
Earliest time of re-emergence of parental beetles was
observed 29 days after tree colonization of the trap tree
‘‘Messerer 2001’’. Their highest frequency of partial re-
emergence occurred at a time when the offspring generation
had developed into third-instars (Table 6). Parental beetles did
not re-emerge from trap trees felled in mid July, except for one
trap tree (Holzgraben II 2002) where sporadic re-emergence
was observed at the end of August. The relative thermal sum
(expressed as percentage of the thermal sum required for total
development) at the time of first partial re-emergence was on
average 49.3  11.2% (minimum: 31.2%). Thus, for modelling
the re-emergence of parental beetles we used a relative thermal
sum of 50% (i.e. 278.5 dd) (see Eq. (A.10)). Complete re-
Table 6
Relative frequency (%) of first observation of partial and complete re-emer-
gence vs. observed developmental stages of I. typographus
Developmental stage Relative frequency (%)
Parental beetles Parental beetles
partially absent completely absent
First instar 0 0
Second instar 13.33 0 with hourly recorded data. However, the mean value calculated
Third instar 40.00 20.00
Pupae 20.00 6.67
Teneral beetles 26.67 33.33
Dark coloured beetles 0 26.67
Exit holes 0 13.33
emergence of parental beetles was observed at a mean relative
thermal sum of 71.5% (minimum: 39%).
3.2.5. Emergence rate of filial beetles
The rate of emergence of filial beetles showed a high
variability between years and locations. While most of these
beetles, especially at locations of high elevation (Haslersgatter
and Weingartalm) and shady sites (Krennbauer, Holzgraben II)
did not emerge during the summer period in 2002, a high
proportion emerged in 2001 and 2003, even at the shady site
‘‘Haslersgatter’’ (1170 m) (Table 7).
Emergence of filial beetles was markedly influenced by the
time of the year: only mature beetles that had completed their
total development at photoperiods before mid of August
(14.5 h daylight) left their brood system (Fig. 3). Thus, we
used a critical day length of 14.5 h as a threshold for
emergence of beetles to establish a new generation.
3.3. Validation of the phenology model using topoclimatic
data
Using the daily data output of the topoclimatic model, the
timing of the key-phenological events in the bark beetle life
cycle were simulated for the study area with the modelling
procedures of PHENIPS (see Eqs. (A.8)–(A.12)). These include
(1) onset of infestation (start of development), (2) the
cumulative sum of effective bark temperature of parental,
sister, and filial generations, and finally (3) the total number of
generations initiate at day length 14.5 h, which can
successfully hibernate.
The potential development of the bark beetle was calculated
using relative solar radiation (SIrel) for open land. For detailed
point-based analysis of its development in the trap trees, the
global site factor was used for calculation of solar radiation
(SIred, Eq. (1)).
Using topoclimatic data for analysis of the bark beetle
development at the trap trees, the estimated thermal sum at the
time of emergence showed a higher variability than calculated
from the topoclimatic data was not significantly different from
the required thermal sum for total development (test value
K = 557 dd) and deviated by 38.8 dd (Table 4). The application
of estimated bark temperatures for computing the develop-
mental stage at the beginning of the cold period in October
 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186 179

Table 7
Emergence ratio of filial beetles for trap trees in the field and for logs in photoeclectors
Year Trap tree (first series) Emergence behaviour, classified in the field Emergence from logs in photoeclectors
Ratio of emergence (%) Classification
2001 Messerer Mainly emerging 98.13 Emerging
Bodinggraben Emerging 79.38 Emerging
Zaglbaueralm Emerging 98.12 Emerging
Ebenforstalm Emerging 97.30 Emerging
Blumaueralm Emerging 70.16 Mainly emerging
Tiefling Not emerging 0.00 Not emerging
2002 Krennbauer Mainly emerging 34.17 Partially emerging
Holzgraben I Emerging 86.41 Emerging
Holzgraben II Mainly emerging 13.54 Not emerging
Mehlboden Partially emerging 2.46 Not emerging
Haslersgatter Not emerging 0.00 Not emerging
Weingartalm Partially emerging 0.00 Not emerging
2003 Krennbauer Emerging 93.38 Emerging
Holzgraben I Emerging 74.52 Mainly emerging
Haslersgatter Partially emerging 69.95 Mainly emerging
Weingartalm Mainly emerging 75.72 Emerging

Status of emergence Criteria for field Ratio of emergence within the

behavioura observations same season of infestation (%)
Emerging Many exit holes, only few young adults remaining >75
Mainly emerging Frequent exit holes; many young beetles in the bark 51–75
Partially emerging Few exit holes; many young beetles in the bark 26–50
Not emerging Few or none exit holes; young beetles almost all in the bark 0–25
a
Criteria for classification of emergence behaviour of filial beetles within the same season of infestation.

revealed that only in one case (location ‘‘Weingartalm’’) the
developmental stage was overestimated (Table 5). Using bark
temperature data calculated by means of topoclimatic data
resulted in a relative thermal sum of 52.17  13.89% at the
time of re-emergence. This mean value was not significantly
different from the relative thermal sum for re-emergence
calculated with hourly observed bark temperatures (t = 0.605;
d.f. = 14; P = 0.555).
The accuracy of our model in estimating the bark beetle’s
phenology is demonstrated by the comparison of the recorded
Fig. 3. Influence of photoperiod and thermal conditions after maturity of the
young beetles (i.e. thermal sum cumulated for 21 days after completion of brood
development) on the emergence behaviour of filial beetles in the field and for
logs in photoeclectors.
with the predicted and potential developmental conditions
(effective thermal sum) and the observed, predicted and
potential developmental stage of the beetles in trap trees at two
selected sites ‘‘Holzgraben I and II’’ in 2002 (Fig. 4; Table 8).
Both sites are located at similar altitudes (1050 m), but clearly
differ in their aspect and GSF (57.7% and 32.4%; Table 1). Due
to the higher sky obstruction by the canopy and the northern
exposition, the low GSF value at ‘‘Holzgraben II’’ results in
lower bark temperatures of the trap tree with a markedly slower
development and a lower number of generations compared with
the conditions in the trap tree at the southern-exposed site.
The point-based comparison revealed a high conformity
between the phenological events predicted by recorded
temperature data at the local trap tree and those predicted by
the topoclimatic model. This indicates the possibility to
compute precisely local temperature conditions inside the bark
by the model. The higher deviation between observed and
predicted events might be due to uncertainties by the weekly
control intervals and to the variability in beetle’s life processes
which are not temperature-dependent. Especially the re-
emergence of parental beetles showed a high variability and
may be strongly affected by intra-specific competition or by the
nutritive quality of the host tree. However, the application of
PHENIPS using detailed data of site-specific conditions can
detect relevant differences concerning the phenology of I.
typographus. The estimation of the potential development can
be used as an indicator of the maximum number of possible
generations for hazard rating.
180 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186

Fig. 4. Time series of observed and estimated daily mean bark temperature, relative thermal sums of the potential development and relative thermal sum of the
predicted development of I. typographus using observed and generated temperature data at two selected locations in 2002. Potential development of a new generation
starts at predicted onset of infestation. Predicted development starts at the observed date of infestation of the first and second trap tree. The observed onset of re-
emergence is indicated by a white asterisk; the black asterisk indicates the date of observed emergence of filial beetles (F1). The dotted horizontal line marks the
threshold for successful hibernation (relative thermal sum 0.6); the dotted vertical line indicates the threshold for induction of diapause (day length = 14.5 h); boxes
illustrate the potential generations.

Table 8
Observed, predicted, and potential phenological events at two selected locations at Kalkalpen National Park in 2002
Phenological event Location Date of event/brood status for hibernation
Observed Predicted by recorded data Predicted by topoclimatic data Potential development
Infestation Holzgraben I 8/5 – 16/5 16/5
Holzgraben II 8/5 – 17/5 17/5
Re-emergence Holzgraben I 24/6 15/6 16/6 20/6
Holzgraben II 24/6 21/6 22/6 24/6
Emergence F1 Holzgraben I 15/7 14/7 13/7 20/7
Holzgraben II 19/7 27/7 1/8 1/8
Hibernation F2 Holzgraben I Adults Adults Adults Adults
Holzgraben II Pupae Pupae Pupae Pupae

Phenological event Location Model parameter Relative thermal sum at observed date of event
Recorded data Topoclimatic data Potential development
Re-emergence Holzgraben I 0.5 0.69 0.70 0.59
Holzgraben II 0.56 0.54 0.50
Emergence F1 Holzgraben I 1 1.02 1.05 0.95
Holzgraben II 0.91 0.85 0.82
Hibernation F2 Holzgraben I 0.6 0.71 0.81 1.09
Holzgraben II 0.55 0.52 0.56
 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186
4. Discussion
Monitoring the development of concealed living insects like
phloem feeding bark beetles is a challenging problem due to the
complex modelling of the specific microclimatic conditions
inside the bark tissue. Especially the effect of solar irradiation
may significantly alter the bark temperature of exposed sites
compared to the ambient air temperature. With PHENIPS we
developed a model that combines the estimation of the
microclimatic conditions inside the bark with the eco-
physiological characteristics of the aggressive spruce bark
beetle, I. typographus. PHENIPS differs from other bark beetle
phenology models (Logan and Bentz, 1999; Ungerer et al.,
1999) in that it is developed for estimation of the phenology and
development of I. typographus by explicit consideration of the
strong effects of regional topography and stand conditions on
local air and bark temperature.
On one hand, PHENIPS can be applied for assessing the
likelihood of outbreaks by modelling the regional potential
development of the bark beetle for simulation the maximum
number of generations (‘‘worst case’’). On the other hand, the
utilization of detailed data of incoming solar irradiation for
modelling bark temperatures provides a precise monitoring of
the actual state of bark beetle development at the specific stand/
tree level.
Due to the limited number of weather stations that record
solar irradiation, the availability of these data is limited. Since
the incoming irradiation shows a relatively high spatial
autocorrelation it is possible to estimate solar irradiation on
a local scale by combining daily solar irradiation recorded at a
reference station located nearby with the potential solar
irradiation derived from a DEM. The estimated relative solar
irradiation is used in an insolation-modified lapse rate model as
a predictive variable for estimating daily air temperature.
Furthermore, the estimated insolation is an important input
parameter for the prediction of daily mean as well as maximum
bark temperature. Actual daily solar irradiation is determined
by daily varying atmospheric conditions (transmissivity). The
forest canopy additionally reduces the incoming solar irradia-
tion. Sky obstruction by the canopy can be estimated in detail
using hemispherical photographs. In practice, forest inventory
data about stand density and canopy closure may be used for
roughly estimating the insolation below the canopy, whereby
the solar irradiation exponentially decreases with increasing
canopy closure (Pennerstorfer, 2000). Within closed stands,
irradiation has a negligible effect on bark temperatures, which
is then equivalent to the air temperature. As it is shown in the
comparison of two local sites with different sky obstruction and
exposition (Section 3.3), the development of the bark beetle is
significantly accelerated due to higher insolation at the exposed
site. However, for estimating the maximum (potential)
developmental rate, the maximum amount of daily solar
irradiation received on open land is relevant, such as the case
after wind throw or at south-exposed stand edges.
Our nonlinear function for simulating bark beetle develop-
ment is based on quantitative experimental data of develop-
mental thresholds and thermal sums (Wermelinger and Seifert,
181
1998; Netherer, 2003). This allowed accurate simulation of
both, preimaginal development and maturation of young adults
in the field (mean relative error: 2.9%); additionally the
overestimation of the brood development in trees exposed to
high insulation was eliminated. Furthermore, completion of
preimaginal development was predicted accurately by the
model at the end of the growing season. Since only the adult
stage of I. typographus can successfully survive cold winter
temperatures (Faccoli, 2002; Netherer, 2003), determination of
the status of brood development, i.e. uncompleted generations
at the end of the season, allows us to estimate the loss of local
populations due to hibernation mortality.
As a comprehensive model for simulating the phenology of
I. typographus, PHENIPS can be used to estimate brood
development, to predict the onset of spring swarming and tree
infestation, and to estimate the development of sister broods
and the number of generations that can be completed prior to
hibernation. The observed onset of spring swarming and flight
activity of I. typographus in the study area confirmed the
temperature threshold of 16.5 8C for flight activity (Lobinger,
1994). After chilling during hibernation, diapause is terminated
and followed by a quiescence stage until the temperature rises
and ovarian development can proceed (Doležal and Sehnal,
2003). However, short-day photoperiods (prior to mid of April)
apparently suppress untimely emergence from hibernation sites
(Dobart, 2006) like it is reported for Ips accuminatus by
inhibiting follicle formation in ovarioles (Gehrken, 1985).
Using both, the lower threshold for flight activity and a specific
thermal sum of daily maximum air temperature above the lower
threshold for beetle activity, the onset of swarming and
infestation was predicted accurately.
Usually, the majority of parental beetles of I. typographus
re-emerge 2–3 weeks after initial attack and establish at least
one sister generation (Martinek, 1956, 1957; Thalenhorst,
1958; Annila, 1969; Anderbrant, 1989; Kritsch, 2005). Besides
density-dependent intra-specific competition (Anderbrant
et al., 1985), the residence time of parental beetles is mainly
temperature-dependent (Annila, 1969; Anderbrant, 1986,
1989). Therefore, the mean re-emergence time is estimated
using thermal sums, accumulated since the brood has been
established (278.5 dd, i.e. 50% of the thermal sum for total
development). In comparison, for modelling mean re-emer-
gence time Annila (1969) suggested that I. typographus
requires 150–200 dd above 5 8C, and Anderbrant (1986)
suggested 158–262 dd with a threshold of 7.5 8C. The
application of the thermal sum for re-emergence additionally
can be used for interpretation of flight activities monitored by
pheromone-trapping to differentiate between swarming of re-
emerged parental beetles and dispersion of filial beetles.
Under favourable thermal conditions and at lower elevations
in Central Europe I. typographus is usually able to establish
more than one filial generation within the same season.
However, also these multivoltine populations are restricted in
their reproductive activities by short-day photoperiods (Schopf,
1985, 1989; Doležal and Sehnal, 2003; Kritsch, 2005).
Apparently, this arrest in gonad maturation, which represents
a facultative adult diapause of I. typographus, is a develop-
182 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186

mental adaptation to seasonal changes and to unfavourable
climatic conditions. It helps to avoid population losses due to
high winter mortality of preimaginal stages. Thus, the
multivoltine life cycle of I. typographus with re-emerging
parental beetles and photoperiodic thresholds influencing flight
and reproduction activity is much more complex and more
challenging to model than simpler life cycles of insects, like
Dendroctonus ponderosae, that depend only on thermal
conditions (Bentz and Mullins, 1999; Bentz et al., 1991).
The modelling procedures of PHENIPS can be used to
conduct a comprehensive analysis of the potential development
and phenology of I. typographus in complex mountainous
terrain. Our model can be used for landscape-scale forecasts.
Using interpolated topoclimatic data, the spatial and temporal

Fig. 5. Onset of infestation (A) and number of generations (B) for the years 2000–2003 at the Kalkalpen National Park (black line: boundary line of the National
Park).
 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186
Fig. 6. Potential succession of generations and theoretically possible number of
offspring originating from one parental female per year in relation to the relative
annual thermal sum (F1–F3: first, second, and third filial generation; SB: sister
generation [prefixed Fx indicates additional filial generations initiated by
offspring of sister brood; arrows indicate parental origin of sister brood]).
The theoretical number of offspring per generation is indicated as the product of
number of females and mean number of offspring per female for each initiated
generation. The total number of offspring is summarised below.
simulation of the potential number of generations per year, like
it is shown for the region of the Kalkalpen National Park
(Fig. 5), is a basic requirement for reliable assessment of the
likelihood of bark beetle establishment and occurrence of
outbreaks. The output of PHENIPS identifies a key-factor and
provides essential data for site-and stand-related hazard rating
of bark beetle infestation (Netherer and Nopp-Mayr, 2005).
Especially for spruce-dominated National Parks, hazard rating
is a crucial tool for bark beetle management. Hazard rating can
be used to establish buffer zones where control measures have
to be performed in order to protect adjacent managed spruce
forests.
Based on the theoretical population growth associated with
the number of potential generations per years (Fig. 6), relative
scores were assigned to the potential number of generations
(Fig. 7). These relative scores can be used for further rating of
site-related hazards of bark beetle infestation (Netherer and
Fig. 7. Scoring of the potential number of generations per year as an indicator
of site-specific predisposition to I. typographus. (1G: one generation; 1G + SB:
one generation and one sister brood; 2G: two generations; >2G: more than two
generations).
183
Nopp-Mayr, 2005). When combined with knowledge-based
hazard rating systems that indicate the key site- and stand-
related factors influencing the susceptibility of hosts to bark
beetle attack, PHENIPS contributes to the identification of
vulnerable areas of forest stands and sites both in space and
time.
Using inputs of air temperature and solar irradiation data
from a nearby weather station, the model can predict the actual
state of maximum bark beetle development on local scale. By
adding local irradiation conditions, PHENIPS further allows
precise monitoring of the brood development at selected
locations in the terrain. This again is important to compare
developmental conditions for the bark beetles at stands which
are already indicated to be highly susceptible to bark beetle
infestation.
PHENIPS may also help to gain deeper insight into the
population dynamics of I. typographus, by prospective and
retrospective analyses of its development. Furthermore, the
model can be used to create scenarios about the likelihood of
bark beetle outbreaks under changing environmental condi-
tions. The interpretation of trap catches and the accuracy of
assessing population densities using pheromone traps (Faccoli
and Stergulc, 2005) could be substantially improved by
combining monitoring programs with models such as
PHENIPS.
Our model can be used with point-based as well as
interpolated meteorological data. The increasing capability to
manage huge data sets and the availability of new tools for
spatio-temporal analysis within GIS will enable the land-
scape-wide application of PHENIPS with high spatial
resolution. The parameterization of PHENIPS was done for
application of the model in mountainous regions of Central
Europe, but it can be adapted by minor changes of the model
parameters (especially concerning photoperiodic thresholds
which depend on latitude) for application at a broader
geographical range.
Acknowledgements
The financial support of this study by the National park O.ö
Kalkalpen GmbH and by the Federal Ministry of Agriculture,
Forestry, Environment and Water Management is gratefully
acknowledged. We thank the ÖBf AG (National Park
management unit Kalkalpen) for technical assistance in field
work.
Appendix A
A.1. Topoclimatic model
To estimate the daily sum of solar irradiation (Wh m2) for
any location in the field (eq. grid cell), we used the potential
solar radiation (Eq. (A.1)), which depends on topographic
factors (elevation, slope, and aspect), latitude, daytime, and
season. Monthly means of the potential solar radiation for each
grid cell were calculated within GIS using DEM-data. The
estimated solar radiation (SIrel) represents the maximum of
184 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186
daily radiation without canopy shading.
 
SIpotðxk Þm
SIrelðxk Þi ¼  SIðrefÞi (A.1)
SIpotðrefÞm
 
SIpotðxk Þm
SFtopoðxk Þm ¼  100 (A.1.1)
SIpotðrefÞm
SIrel(xk)i is the estimated solar radiation at the location xk for
day i; SIpot(xk)m the potential solar radiation at the location xk
for month m; SIpot(ref)m the potential solar radiation at the
reference station for month m; SI(ref)i the observed global solar
radiation at the reference station for day i; SFtopo(xk)m is the
site-related, topographic factor.
Daily mean and maximum air temperatures for any grid cell
were calculated using the daily regression coefficients,
elevation and estimated daily sum of solar radiation (SIrel)
as input variables (Eq. (A.2)).
ATðmean=maxÞðxk Þi ¼ ai þ bi  elevðxk Þ þ ci  SIrelðxk Þi
(A.2)
AT(mean/max)(xk)i is the air temperature (mean/maximum) at
the location xk for day i; elev(xk) the elevation of the location xk;
SIrel(xk)i the estimated solar radiation at the location xk for day i
(Eq. (A.1)); ai, bi, ci are the daily coefficients from regression
analysis for ATmean and ATmax, respectively.
The calculation of the bark temperature is based on
regression analysis with air temperature and solar radiation
as predictive variables (Eqs. (A.3) and (A.4)). The calculation
of the effective bark temperature using the nonlinear function
(Eq. (2)) was adopted for usage with daily temperature data,
provided by the topoclimatic model. For daily maximum bark
temperatures below the beetles’ developmental optimum
(TO  30.4 8C), the daily sum of effective bark temperature
was calculated using the simple linear function (Eq. (A.6)). In
order to account for the daily variability in bark temperature
with temporary daily maxima above TO, the difference in the
effective thermal sum calculated with hourly data according to
Eq. (2) compared with the linear computed thermal sum
(Eq. (A.6)) was estimated for daily bark temperature maxima
above TO. This difference (Diff_BTsum) showed a highly
significant correlation with the daily bark temperature
maximum (R2 = 0.844, P  0.001) and was estimated with
Eq. (A.5). For BTmax > TO and Diff_BTsum > 0 dd the
effective thermal sum was calculated with Eq. (A.7).
BTmeanðxk Þi
¼ 0:173 þ 0:0008518  SIrelðxk Þi þ 1:054
 ATmeanðxk Þi (A.3)
BTmaxðxk Þi ¼ 1:656 þ 0:002955  SIrelðxk Þi þ 0:534
 ATmaxðxk Þi þ 0:01884  ðATmaxðxk Þi Þ2
(A.4)
310:667 þ 9:603  BTMaxðxl Þi
Diff BTsumðxl Þi ¼ (A.5)
24
If BTmaxðxk Þ ¼ To then BTsum effðxk Þi
¼ ðBTmeanðxk Þi  8:3Þ
If BTmaxðxk Þ > To and Diff BTsumðxk Þi > 0
then BTsum effðxk Þi ¼ BTsumðxk Þi  Diff BTsumðxk Þi
(A.6)
where
BTsum effðxk Þi
¼ 0 dd if BTsumðxk Þi  Diff BTsumðxk Þi < 0 dd (A.7)
BTmean(xk)i is the mean bark temperature at the location xk for
day i; AT(xk)i the mean air temperature at location xk for day i;
BTmax(xk)i the estimated daily maximum of bark temperature
for the location xk for day i; SIrel(xk)i the daily sum of solar
radiation reduced with the GSF at location xk for day i;
ATmax(xk)i the maximum air temperature at location xk for
day i; Diff_BTsum(xk)i the estimated difference between linear
and nonlinear modelled thermal sum at the location xk for day i;
BTsum(xk)i is the daily effective thermal sum of bark tempera-
ture at location xk for day i.
Estimated daily air temperature maximum (ATmax) and
daily effective thermal sum of bark temperature (BTsum_eff)
were used as climatic input parameters for calculating the
seasonal phenology of I. typographus with PHENIPS.
A.2. Phenology model of I. typographus
PHENIPS comprises following stepwise computations for
any location (grid cell):
 Calculation of the year day of onset of infestation
(YDinfestation) based on the threshold for flight activity and
on cumulative daily maximum air temperatures (ATmax)
above DTL from the April 1st (YD = 92) onwards
X
YDinfestation if ATmax > 16:5 C and ðATmax
 8:3Þ  140 dd (A.8)
 Calculation of the cumulative sum of effective bark
temperature (BTsum_eff) relative to the necessary thermal
sum for total development (K = 557 dd) of parental genera-
tion
if YD  YDinfestation
P 
BTsum eff (A.9)
TsumF1 ¼
K
 Calculation of the onset of sister broods and their cumulative
sum of effective bark temperature (BTsum_eff) relative to the
necessary thermal sum for total development (K = 557 dd) of
sister brood
YDsister brood if TsumF1 > 0:5 and ATmax > 16:5 C and
day length  14:5 h
if YD  YDsister brood
P 
BTsum eff
Tsumsister brood ¼
K (A.10)
 P. Baier et al. / Forest Ecology and Management 249 (2007) 171–186
 Calculation of the onset of second and third filial generation
and their cumulative sum of effective bark temperature
(BTsum_eff) relative to the necessary thermal sum for total
development (K = 557 dd)
YDF2 if Tsum > 1 and ATmax > 16:5 C and day
length  14:5 h
if YD  YDF2
P  Sturmschadensgebieten. Schweizerische Zeitung f. Forstwesen, pp. 767–776.
BTsum eff
TsumF1 ¼
K (A.11)
YDF3 if Tsum > 2 and ATmax > 16:5 C and day
length  14:5 h
if YD  YDF3
P  Working Party 7.03.10, Sion-Chateauneuf, Switzerland, April 20–23.
BTsum eff
TsumF2 ¼
K (A.12)
Finally, the relative thermal sum of each initiated generation
at the end of October of each year and of each grid cell is
evaluated for its probability of survival during the following
cold period. At relative thermal sums higher than 60% of the
necessary thermal sum for total development (TsumFx 0.6),
the initiated generation has completed its preimaginal
development and can successfully hibernate as young adults.
Therefore, initiated generations with relative thermal sums
<0.6 are ignored for calculating the total number of potential
generations per year.
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