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The influence of daylight regime on diurnal locomotor activity patterns of the European hare (Lepus europaeus) during summer

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Knowledge on diurnal locomotor activity pattern in wild nocturnal medium-sized mammals, such as the European hare (Lepus europaeus) is scarce. In this study, we tracked nine European hares during the vegetation period using GPS-transmitters. In particular, we focused on the question how the timing of sunset and sunrise influences the activity peaks in this species. The horal distances between two consecutive hare positions were used as a measure of locomotor activity. European hares showed two distinct peaks in their daily activity. If sunset or sunrise were earlier, the maximum activity peaks of individual European hares occurred after sunset or sunrise, whereas activity peaks were shifted before sunset or sunrise when sunset or sunrise were later. During summer, when the nights are probably too short to allow the hares to cover their energetic requirements, the study animals regularly showed activity peaks in full daylight. In conclusion, our results imply that, although daylight regime normally regulates the diurnal locomotor activity pattern in mammals, other additional factors may play a role in modifying this regulation in European hares during summer.
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Mammalian Biology 77 (2012) 434–440
Contents lists available at SciVerse ScienceDirect
Mammalian Biology
journal homepage: www.elsevier.com/locate/mambio
Original Investigation
The influence of daylight regime on diurnal locomotor activity patterns of the
European hare (Lepus europaeus) during summer
Stéphanie C. Schai-Brauna,, Heiko G. Rödelb, Klaus Hackländera
aInstitute of Wildlife Biology and Game Management, University of Natural Resources and Life Sciences, Vienna, Gregor Mendel Strasse 33, A-1180 Vienna, Austria
bUniversité Paris 13, Sorbonne Paris Cité, Laboratoire d’Ethologie Expérimentale et Comparée (LEEC), F-93430 Villetaneuse, France
article info
Article history:
Received 1 February 2012
Accepted 13 July 2012
Keywords:
Lepus europaeus
GPS
Activity pattern
Locomotion
abstract
Knowledge on diurnal locomotor activity pattern in wild nocturnal medium-sized mammals, such as
the European hare (Lepus europaeus) is scarce. In this study, we tracked nine European hares during
the vegetation period using GPS-transmitters. In particular, we focused on the question how the timing
of sunset and sunrise influences the activity peaks in this species. The horal distances between two
consecutive hare positions were used as a measure of locomotor activity. European hares showed two
distinct peaks in their daily activity. If sunset or sunrise were earlier, the maximum activity peaks of
individual European hares occurred after sunset or sunrise, whereas activity peaks were shifted before
sunset or sunrise when sunset or sunrise were later. During summer, when the nights are probably
too short to allow the hares to cover their energetic requirements, the study animals regularly showed
activity peaks in full daylight. In conclusion, our results imply that, although daylight regime normally
regulates the diurnal locomotor activity pattern in mammals, other additional factors may play a role in
modifying this regulation in European hares during summer.
© 2012 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.
Introduction
In mammals, circadian rhythms are predominantly regulated by
light (Goldman 1999; Cermakian and Sassone-Corsi 2002; van der
Merwe et al. 2011). Numerous studies have shown that the impact
of light as zeitgeber might be affected by various other intrinsic and
extrinsic factors, such as food availability, weather, temperature,
sex, season, reproductive status, and age (Getz 1961; Garshelis and
Pelton 1980; Zielinski et al. 1983; Ferguson et al. 1988; Larivière
et al. 1994; Kolbe and Squires 2007; Wronski et al. 2006; Rödel
et al. 2012). However, sunrise and sunset have been suggested to
trigger the onset and cessation of activity in a wide range of species
(Daan and Aschoff 1975; Benstaali et al. 2001).
Hares (genus Lepus) have been described as mostly nocturnal
mammals (Chapman and Flux 2008), although this seems to be true
only during winter (Homolka 1986; Pépin and Cargnelutti 1994;
Holley 2001). In summer, activity of hares appears to be less con-
sistent and partly diurnal (Mech et al. 1966; Cederlund and Lemnell
1980; Figala et al. 1984). Irrespective of this, also in hares sunset
and sunrise appear to play a major role concerning the onset and
cessation of activity, respectively (Mech et al. 1966; Figala et al.
1984; Pépin and Cargnelutti 1994; Holley 2001).
Corresponding author.
E-mail address: stepbraun@yahoo.com (S.C. Schai-Braun).
In winter, hares start their daily activity shortly after sunset and
end it shortly before sunrise (Cederlund and Lemnell 1980; Pépin
and Cargnelutti 1994). However, studies on different hare species
report contradictory results regarding the influence of sunrise and
sunset as zeitgebers during late spring or summer. Snowshoe hare’s
(Lepus americanus) cessation of activity has been reported to be on
average 1 h before sunrise (Mech et al. 1966; Figala et al. 1984),
however, with notable variation. For some individuals the onset of
activity was 1 h after sunset (Mech et al. 1966), whereas another
one was observed to start its activity more than 2 h before sun-
set (Figala et al. 1984). European hares began to leave their forms
before sunset and to enter them after sunrise as the nights short-
ened in the early part of the year (Holley 2001). That means, in all
studies during late spring or summer sunrise and sunset somehow
trigger onset and cessation of hares’ activity, but the impact of these
zeitgebers is various.
It has been argued that the impact of sunrise and sunset in sum-
mer was altered by the number of daily night hours (Holley 2001).
In this European hare study the duration of the activity period did
not remain constant but decreased from 15 h in January to 12 h at
the end of March, mirroring the number of daily night hours. At this
point, the activity period was not further contracted but the hares
suddenly started to increase their activity period by including day-
light activity. This sudden transition from a totally nocturnal to a
partially diurnal regime was explained by an aversion to daylight
activity. Consequently, we suppose that the number of daily night
1616-5047/$ – see front matter © 2012 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.
http://dx.doi.org/10.1016/j.mambio.2012.07.004
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S.C. Schai-Braun et al. / Mammalian Biology 77 (2012) 434–440 435
hours alter the impact of the predominant zeitgebers sunrise and
sunset. Hence, the hares’ activity pattern should display a sudden
start of daylight activity at the beginning and an abrupt withdrawal
from daylight activity at the end of summer.
In addition, a high ambient temperature might influence the
activity pattern of mammals. For example, in black bears (Ursus
americanus) it was shown that temperatures above 25 C substan-
tially reduced the level of activity (Garshelis and Pelton 1980).
Furthermore, the meadow vole (Microtus pennsylvanicus) was
found to abandon diurnal activity in favour of nocturnal and/or
crepuscular activity when the temperature rose above 20 C(Getz
1961). We hence assumed that the impact of the zeitgebers sun-
rise and sunset might be altered by a high ambient temperature
in which case the hares’ activity would be restricted to the dark
period.
However, detailed quantitative and individual-based data on
daily activity pattern in this genus are still scarce. In this study we
investigated the European hare’s (Lepus europaeus) diurnal loco-
motor activity patterns during summer. In particular, we did not
only focus on the timing of onset and cessation of activity but also
studied subtle changes of activity during 24 h. Our hypothesis was
that sunrise and sunset, the predominant zeitgebers for the Euro-
pean hare’s diurnal locomotor activity pattern, are slightly altered
in their impact by the number of daily night hours (season) and the
temperature. We tested this hypothesis by equipping nine individ-
uals with GPS collars, allowing us to assess their diurnal locomotor
activity patterns.
Material and methods
Study area
The study was conducted in Lower Austria near Zwerndorf
(4820N, 1650E) and the study area consisted of 270 ha arable
land with cereals as the main crop and an average field size of 3.1
(±0.3 SE) ha. Hare density in the study site was estimated in autumn
2009 by spotlight counts (Langbein et al. 1999) and accounted 35
European hares per 100 ha (SSB & KH, unpubl.).
Data collection
Nine adult European hares (4 males, 5 females) were caught
in un-baited box traps from May until September 2009. All ani-
mals were sexed according to secondary sexual characteristics and
equipped with a 70 g GPS collar (Telemetry Solutions, Quantum
4000 Enhanced). The collars were programmed to start working
right after the animal’s release and take 1 fix per hour. For additional
information on the individual hares’ GPS-data see the electronic
appendix. The accuracy of the GPS collars was tested beforehand
(see Harris et al. 1990) and yielded a mean precision of 3.5 m (±1.0
SE). Weather data and the time of sunrise and sunset were provided
by the Austrian Central Institute for Meteorology and Geodynamics.
Temperatures were recorded daily at 7 am and 7pm CET.
Calculation of positional data
The positional data were digitised using the software ArcGIS
9.2 (ESRI). We only included locations with a solution in three-
dimensional mode (based on 4 satellites) (Frair et al. 2010).
The distances (in metres) between two consecutive hare positions
(horal distance) were calculated. Although the horal distance does
not reveal the effective distance the hare covered between these
two fixes, it exposes a minimal distance the hare must have moved
during this hour. In the following, the “horal distance” is used as a
measure of hares’ activity, and the term “activity” is always used in
the sense of the hares’ locomotor activity.
Statistical data analysis
We analysed the data using multivariate (generalized) linear
mixed-effects models, allowing for the use of repeated measure-
ments. Statistical analyses were done with the software R 2.12.0 (R
Development Core Team 2011). Generalized linear mixed-effects
models were fitted using the package lme4 (Bates 2005). P-values
were extracted by likelihood ratio tests (Faraway 2006). When
using linear models, we visually checked normality of the model
residuals by normal probability plots. For all models, the homo-
geneity of variances and goodness of fit were examined by plotting
residuals versus fitted values (Faraway 2006).
We initially included sex in all models tested. However, since
there were never any significant effects of sex in our multivariate
analyses (p> 0.10), this factor was omitted from the models before
re-calculation.
Diurnal activity pattern
We tested the effect of time of the day (covariate, in hours)
on the response variable horal distance by a linear mixed effects
model. In addition, we tested similar models only including data
subsets: one subset included all positional data with a shorter time
interval to sunset than to sunrise, and the other one comprised the
remainder. For all models, the response variable (horal distance)
was log-transformed in order to obtain a normal distribution of the
model residuals.
Since we expected a non-linear time course with at least one
maximum peak, we used polynomials to model the data. For this,
we gradually increased the complexity of the polynomials until the
9th order. All of these models were tested for significance, and,
in addition, we directly compared the support of these models by
using AICc (Burnham and Anderson 1998). The model with the low-
est AICc score can be considered as the best approximating model
of the model set. Note that different models can be considered to
find equally good support by the data when the AICc is smaller
than 2.
All (mixed effects) models included hare identity as a random
factor, in order to allow for the repeated measurements collected
from the different hares, and also an individual-specific code for
the day (“date”) as a second random factor in order to account for
the time series measured for each of the study animals during the
different days of the study. As it could be expected, there was sig-
nificant individual variation among the hares’ locomotor activity
(significant random factor “individual hare”: 2= 199.44, p< 0.001)
with a variance of 0.21 m/h (±0.55 SD).
Influence of different parameters on the activity peaks
A peak can be described mathematically by a parabola, i.e. by
a 2nd-order polynomial. The first derivative is used to find the
vertex of a parabola as the first derivative equals zero at the ver-
tex. At the maximum point of the parabola the hare’s activity is
highest. Therefore, if the horal distance is a function of the time
interval to and since sunrise or sunset, the independent variable
of this function indicates the time interval of maximum locomotor
activity.
Firstly, the time interval with the highest activity was deter-
mined. As the time of sunset and sunrise changes at about 1 min
per day in summer, and as each of the nine hares provided a dif-
ferent number of horal distances, several maximum points for each
animal were calculated. The horal distances of one day did not yield
enough data to calculate a morning and an evening maximum point.
For this reason, we pooled the horal distances of five days for every
hare to calculate one evening and one morning maximum point. If
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436 S.C. Schai-Braun et al. / Mammalian Biology 77 (2012) 434–440
Fig. 1. Locomotor activity (measured as the horal distances moved by the hares)
described by a 6th-order polynomial regression model. Data (squares) are shown as
medians with 25th/75th percentiles. Time of sunrise/sunset during the study period
is indicated by grey shading. See text for details on statistics.
the first derivative at zero described a minimum point or an impos-
sible maximum point, the point was excluded from further analysis.
Out of 171 individual hare days (comprising 3506 hare locations) 36
extrema were calculated for the morning as well as for the evening.
47% of the morning extrema and 75% of the evening extrema were
sensible maximum points.
Secondly, the mean time of sunset and sunrise was calculated for
every five day interval. Each evening maximum point was paired
with its corresponding time of sunset and each morning maximum
point with its corresponding time of sunrise. Subsequently, the cor-
relations were tested by two linear mixed-effects models with hare
identity as random factor (see Fig. 3a and b). The same models were
used to test for correlations between activity peaks and the num-
ber of daily night hours (see Fig. 3c and d) and the temperature,
respectively.
Light conditions and the occurrence of a morning activity peak
Here, we analysed two subsets of the data. One subset included
all fixes taken when the evening activity peak was during the dark
phase, the other one comprised the remainder (see Fig. 4a). To test
the effect of the light conditions on the occurrence of a morning
activity peak, we used a generalized linear mixed effects model for
binomial data with a logit-link function (“logistic regression”). Also
here, hare identity was included as a random factor (see Fig. 4b).
Results
The mean number of satellites used for the location of the fixes
was7(±0.03 SE). The overlapping individual study periods were
on average 11 days (±9.2 SE) long with a minimum of 2 and a max-
imum of 91 days. That is, the number of GPS-fixes taken per animal
ranged from 25 to 2127 with an average of 230 (±219.4 SE). This
resulted in a total of 3528 fixes and thus a total of 3519 horal dis-
tances available for analysis (709 for males, 2810 for females). For
additional information on the individual hares’ GPS-data see the
electronic appendix.
Diurnal activity pattern
The daily time course of the hares’ activity, measured as
the horal distance covered between two fixes, was significantly
explained by a 6th order polynomial (2
6=1078.6, p< 0.001). This
model predicted for the European hares’ diurnal locomotor activity
Table 1
Results of linear mixed-effects models for the effects of different predictor variables
on the timing of locomotor activity peaks, including hare identity as a random fac-
tor. Analyses are based on nevening = 27 and nmorning = 17 five-day activity intervals of
repeated measurements of nine hares.
Response variables Predictor variables Slope 2
1p
Morning activity
peak
Sunrise 5.16 25.06 <0.001
Number of daily night hours 2.31 22.88 <0.001
Morning temperature 0.39 1.57 0.21
Evening activity
peak
Sunset 1.82 10.68 <0.001
Number of daily night hours 1.02 11.06 <0.001
Evening temperature 0.18 1.09 0.30
two distinct activity peaks during night time (see model graph
in Fig. 1). We also verified that the 6th order polynomial was
the best approximating model describing this relationship by a
comparison with higher and lower parameterised models using
AICc values. This model selection procedure revealed that this
model had the lowest AICc score, with AICc > 87 in comparison
to all lower parameterised models, and AICc >1 in comparison to
higher parameterised models.
In relation to sunset and sunrise, the model including a 4th
order polynomial predicted that the hares’ activities were highest
1 h after sunset and again 5h later (2
4=489.75, p< 0.001; Fig. 2a).
Furthermore, a model including a 3rd order polynomial predicted
that hares were most active 5–3 h before sunrise (2
3=445.18,
p< 0.001; Fig. 2b). The animals did not show any notable activities
5–8 h before sunset and 1 h before sunrise until 8 h after sunrise,
when the average distance moved by the hares was less than 10 m
within 1 h. Also here, we verified by model selection with AICc,
that the chosen polynomials were the best approximating mod-
els compared to models including lower-order (AICc > 76 and
AICc > 70, respectively) or higher-order polynomials (AICc > 2
and AICc > 2, respectively).
Influence of different parameters on the activity peaks
Daylight regime
There were significant and negative correlations between both
sunset and the timing of the evening activity peak and between
sunrise and the timing of the morning activity peak (Table 1). If
sunset or sunrise were earlier, the maximum activity peaks of indi-
vidual hares occurred later, whereas activity peaks were shifted
before sunset or sunrise when sunset or sunrise were later (Fig. 3a
and b). Note that the regression slope of the correlation between
sunset and the timing of the evening activity peak was steeper than
between sunrise and the timing of the morning activity peak.
Number of daily night hours
As expected, we found similar effects when testing the variable
number of daily night hours on hare activity peaks (Fig. 3c and d,
Table 1), simply because the number of daily night hours was highly
collinear with sunrise and sunset (correlation of sunrise vs. number
of daily night hours: R2= 0.99, F1,100 = 1.5 ×1031 ,p< 0.001, sunset
vs. number of daily night hours: R2= 0.97, F1,100 = 3628, p< 0.001).
Temperature
There were no statistically significant correlations between the
morning temperature and the timing of the morning activity peaks
and between evening temperature and the timing of the activity
peaks in the evening (Table 1).
Light conditions and the occurrence of a morning activity peak
There was a significant effect of light conditions at the time of
the evening activity peak on the number of observed activity peaks.
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S.C. Schai-Braun et al. / Mammalian Biology 77 (2012) 434–440 437
Fig. 2. Locomotor activity (measured as the horal distances moved by the hares) in relation to the time of (a) sunset described by a 4th-order polynomial regression model
and (b) sunrise described by a 3th-order polynomial regression model. Data (squares) are shown as medians with 25th/75th percentiles. Dark phase is indicated by grey
shading.
The significant models predicted two distinct maxima when the
evening activity peak was during daylight (2
6=230.09, p< 0.001,
Fig. 4a; dashed line), whereas there was only one activity maximum
when the evening activity peak was during the dark phase (2
6=
731.61, p< 0.001, Fig. 4a; solid line). In other words, there was a
higher probability of occurrence of a second activity peak (in 79%
of cases) when the evening activity peaks of individual hares were
in full daylight (2
1=15.50, p< 0.001; Fig. 4b).
Discussion
Locomotor activity pattern in the course of the day
The hares of our study showed two peaks in their daily activity,
having a distinct one in the evening and another less pronounced
one in the morning. Thus, it can be concluded that European hares
have a time of reduced activity in-between an enhanced activity at
the beginning and end of the night. When looking at the activity
of the European hare in relation to sunset, the data indicated an
activity peak around sunset followed by a decrease and a second
increase in activity during the dark phase. During the hour of high-
est activity, hares moved on average 70 m. As the average field size
in the study area was 3.1 ha, we suggest that hares used only a few
different field types per night while being active. Furthermore, the
model predicted a peak before sunrise followed by a long period of
almost no activity. That is, the predicted periods of inactivity did not
last for the whole light phase. As a consequence, there was notable
locomotor activity well before sunset and after sunrise.
Factors influencing the diurnal locomotor activity rhythm
Influence of sunrise and sunset on the diurnal locomotor activity
rhythm
There was a significant negative correlation between both the
timing of the morning activity peak and sunrise as well as between
the timing of the evening activity peak and sunset. As we assumed
that the activity peaks are collinear with onset and cessation of
activity, our results imply that the activity and inactivity of Euro-
pean hares in summer are closely tied to sunset and sunrise. This is
in accordance with other hare studies describing the major role of
sunrise and sunset concerning the onset and cessation of activity
(Mech et al. 1966; Figala et al. 1984; Pépin and Cargnelutti 1994;
Holley 2001). Nevertheless, the negative correlations in summer
imply that the impact of sunrise and sunset differ from winter.
While in winter European hares consistently started their daily
activity shortly after sunset and ended it shortly before sunrise
(Pépin and Cargnelutti 1994; Holley 2001), in summer the hare’s
activity peaks occurred after sunset or sunrise when sunset or sun-
rise were earlier, whereas activity peaks shifted before sunset or
sunrise when sunset or sunrise were later. Our results during sum-
mer confirm the ones of Holley (2001) who observed European
hares’ starting to leave their forms before sunset and enter them
after sunrise in the early part of the year. Thus, we conclude that
the power of the zeitgebers sunrise and sunset is altered in summer
by a seasonal factor.
The steep regression slope between sunrise and the timing of
the morning activity peak resulted in an 8–11 h shift in the timing
of the morning activity peak. In comparison, the shift of the timing
of the evening activity peak was much lower i.e. 5–6 h. This result,
indicating that sunset might be a stronger zeitgeber for the Euro-
pean hare’s diurnal locomotor activity pattern than sunrise, is in
line with another European hare study conducted in winter (Holley
2001). The latter study reported a stronger tie between onset of
activity and sunset than between cessation of activity and sunrise.
Influence of season on the diurnal locomotor activity rhythm
The negative correlation between the timing of the activity
peaks and sunrise or sunset might be explained by the European
hare’s need to fulfil its daily energetic demands, because the length
of the dark phase might limit the hare’s feeding time. Hackländer
et al. (2002) showed that European hares fed with a diet compara-
ble to the composition of stomach contents in free-ranging hares
had a higher food intake yet assimilated less energy than hares fed
with a high fat diet. This might be due to a trade-off between the
energy benefit of increasing food-intake and the additional weight
load in a flight animal (Hackländer et al. 2002). Such a trade-off
might also account for the small resting periods dispersed during
activity (Averianov et al. 2003) which may be used by the European
hares to digest before new food intake can take place. The results
of another study indicated that the duration of European hares’
activity period cannot be contracted further a certain point (Holley
2001). A possible explanation may be the hare’s need to fulfil its
daily energetic demands in combination with required digestive
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438 S.C. Schai-Braun et al. / Mammalian Biology 77 (2012) 434–440
Fig. 3. Correlations between (a) evening activity peaks and sunset, (b) morning activity peaks and sunrise, (c) evening activity peaks and the number of daily night hours
and (d) morning activity peaks and the number of daily night hours with the regression lines (statistically significant). Dark phase is indicated by grey shading. See text for
details on statistics.
breaks. Sunset was late and sunrise early until middle of July and the
hare’s evening and morning activity peaks proceeded in full day-
light. During this time the dark phase seems not to be long enough
for the European hare to accomplish its energetic requirements. The
number of daily night hours was longer in August and September,
and hare’s evening and morning activity peaks took place during
the dark phase. We propose that during the vegetation period the
usual trigger of activity and inactivity, namely sunrise and sunset,
is slightly altered in its impact by the hares’ instinct to accomplish
their daily energetic requirements. Hence, the season in the form
of the number of daily night hours had an influence on the activity
pattern of the European hare and altered the impact of the usual
zeitgebers sunrise and sunset. There was no difference noticeable
in the influence of sunrise and sunset and the numbers of daily
night hours as these parameters were collinear.
Moreover, the season seems to have an impact on the hare’s
inactivity period during daytime. Our results are in line with pre-
viously reported activity data in European hares (Homolka 1986;
Holley 2001) showing that activity during the day is displayed
predominantly in summer. Outside the summer period, on the
contrary, European hare studies reported almost no activity during
daytime (Homolka 1986; Pépin and Cargnelutti 1994; Holley
2001). However, our study animal’s period of inactivity exceeded
the previously reported length of inactivity in European hares
during summer. Homolka (1986) showed that in Moravia (Czech
Republic) during a one all-day visual observation in July 30–50% of
the hares were active with foraging throughout the day except for
about 4 h in the middle of the day. The hares’ distinct length of inac-
tivity during the day might be explained by the different climate
in Southern Moravia and the eastern part of Lower Austria. The
Pannonian climate is responsible for hot and dry summers in our
study area, whereas the continental climate of Southern Moravia
is more moderate. Our study animals may react to the Pannonian
climate in summer with an extended period of inactivity during
the day. We therefore conclude that the length of inactivity during
daytime depends both on season and climatic conditions.
Influence of temperature on the diurnal locomotor activity rhythm
The ambient temperature has been proposed to influence the
activity pattern of some mammal species (Getz 1961; Garshelis
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S.C. Schai-Braun et al. / Mammalian Biology 77 (2012) 434–440 439
Fig. 4. (a) Polynomial regression models describing locomotor activity (measured as the horal distances moved by the hares) with the evening activity peak being in the
light (dashed line) or dark phase (solid line). (b) Probability of the existence of a morning activity peak when the preceding evening activity peak was during the dark or light
phase. See text for details on statistics.
and Pelton 1980). However, this does not seem to be the case for
the European hare, at least not during the vegetation period, as we
could not find significant correlations between the activity peaks
and the temperature. An explanation might be that the European
hare, originally coming from the savannas of Eurasia (Averianov
et al. 2003), is adapted to high ambient temperatures, and, as a
result, the impact of the zeitgebers sunrise and sunset is not altered
by the daily ambient temperature.
Light conditions and the number of activity peaks
Our results revealed a significant influence of light conditions
at the time of the evening activity peak on the number of activ-
ity peaks. If the evening activity peaks of individual hares were in
full daylight, there was a higher probability of occurrence of a sec-
ond activity peak. It seems as if some unknown factor provokes an
increased activity by antedating the evening activity peak into full
daylight and hereinafter by evoking a second activity peak in the
early morning hours. No such findings were ever reported in the lit-
erature regarding hare’s activity patterns. We suggest that future
studies on hares’ activity patterns during the vegetation period
might carefully examine the number of activity peaks during the
night and decipher the factor(s) provoking an increased activity.
Conclusion
In conclusion, our study clearly shows that also in summer,
sunrise and sunset were the zeitgebers for the European hare’s
diurnal locomotor activity pattern. However, the results indicate
that this relationship was altered by the number of daily night
hours (season). We speculate that the hares’ feeding activity during
the short summer nights is not sufficient to cover their daily ener-
getic requirements during the vegetation period. The temperature,
however, had no influence on the locomotor activity pattern of the
European hare.
Acknowledgements
We thank the hunting society of Zwerndorf for cooperation,
especially Walter Metz for his help with hare trapping. The
study was funded by the following foundations or organisations:
Parrotia-Stiftung, Stiftung Dr. Joachim de Giacomi, Basler Stiftung
für biologische Forschung, Messerli Stiftung, Carl Burger Stiftung,
CIC Schweiz, CIC Deutschland, Paul Schiller Stiftung and Karl Mayer
Stiftung. The study complies with the current laws of Austria.
Appendix A. Supplementary data
Supplementary data associated with this article can be
found, in the online version, at http://dx.doi.org/10.1016/j.
mambio.2012.07.004.
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... Our review confirms that the hare is predominantly nocturnal (our prediction 4) and, to a lesser extent, crepuscular, though some diurnal activity is also recorded. Time of day shows a non-linear effect on activity level best supported by a 6th-order polynomial, mirroring the activity pattern reported by Schai-Braun et al. (2012). According to seasonal studies (Pèpin & Cargnelutti 1994, Zaccaroni et al. 2013, Carbone 2019, Viviano et al. 2021, Table 3. ...
... When days become shorter, diurnal herbivores can compensate either by increasing daytime activity (Fattorini et al. 2019) or by starting nocturnal activity (Carnevali et al. 2016). A similar mechanism could thus occur for hares during warm months, when shorter nights may force this nocturnal lagomorph to compensate by increasing its diurnal activity (Schai-Braun et al. 2012, Viviano et al. 2021. Beaudoin et al. (2004) showed that predation risk is fundamental in determining the activity rhythms of a closely related species, the snow-shoe hare Lepus americanus. ...
Article
The European brown hare Lepus europaeus is the most widely distributed hare species of the world, being naturally present throughout Eurasia and introduced as a game species in most continents. Despite the importance of this lagomorph for both management and conservation, a quantitative summary of its spatiotemporal behaviour is still lacking. Taking advantage of 51 selected studies conducted throughout the native range of the hare and spanning the last 40 years, we used meta‐analytic approaches in order to: 1) investigate home range size in relation to ecological factors; 2) test preference across major habitat types; and 3) provide a quantitative synthesis of hare activity patterns. Temporally adjusted home range size of hare populations decreased with increasing cropland cover (cereal and non‐cereal crops), suggesting that home range size gets smaller with increasing food availability, and predicting that hare populations living in absence of crops would double their mean home range size compared to those inhabiting areas covered entirely by cropland. Hare populations where more males were sampled showed larger home ranges, in line with the polygynous mating system of this species. Hares preferred cropland and grassland over other habitats for foraging, thus selecting the two habitats characterised by the majority of trophic resources for this species. Yet, habitat types were used proportionally to their availability overall, except for the general avoidance of human settlements. Hares were mainly nocturnal, with the lowest activity during daytime, when activity was about one third of that at night, and showed moonlight avoidance, probably to limit encounters with nocturnal predators. Our work emphasises the importance of open habitats and especially cropland for this lagomorph, but also suggests some plasticity in the use of space and time by hares. Such plasticity may help this mammal to cope with future environmental changes, providing that landscape heterogeneity is maintained. Home range size of hare populations decreased with increasing cropland cover, suggesting that home range size gets smaller with increasing food availability, and predicting that hare populations living in absence of crops would double their mean home range size compared to those inhabiting areas covered entirely by cropland. Hare populations where more males were sampled showed larger home ranges, in line with the polygynous mating system of this species. As to habitat selection, habitat types were used proportionally to their availability overall, except for the general avoidance of human settlements. Hares were mainly nocturnal, with the lowest activity during daytime, when activity was about one third of that at night, and showed moonlight avoidance, probably to limit encounters with nocturnal predators.
... Additionally, the presence of small carnivores, potential hare predators, has been suggested to alter the spatial behavior of this lagomorph, which shifts its spatial behavior to open areas with short vegetation where detection of potential predators is highest [45,46], or in areas rarely used by predators [47]. The European brown hare is reported to be mainly nocturnal, with activity peaking mostly in the first part of the night or in the crepuscular hours [48][49][50][51]. However, no information is available on its temporal adaptations to limit encounters with its main predators, i.e., mesocarnivores [30,[52][53][54][55]. ...
... In general, hares were confirmed to be mostly nocturnal, in line with previous studies [44,[48][49][50][51]. Mammalian predators and competitors may however play a major role in structuring the spatiotemporal behavior of this important prey species. ...
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Analysis of spatiotemporal partitioning is pivotal to shed light on interspecific coexistence. Most research effort has involved large-sized carnivores and their prey, whereas little attention has been devoted to lagomorphs. We assessed spatiotemporal overlap among the European brown hare Lepus europaeus and its potential competitors and predators through camera-trapping in an area in Central Italy. We estimated the interspecific patterns of the spatiotemporal activity rhythms of brown hares, its potential predators (the red fox Vulpes vulpes, the pine marten Martes martes, the domestic cat Felis catus, and the domestic dog Canis familiaris), and a competitor, the roe deer Capreolus capreolus. Brown hare activity was studied in natural conditions as well as in a fenced area that excluded terrestrial predators and competitors. Free-ranging hares developed a more nocturnal behavior to avoid diurnal predators (i.e., domestic carnivores and martens). Although high temporal overlap was observed between free-ranging brown hares and both red foxes (82%) and roe deer (81%), hares avoided fox by being more active on darkest nights, as well as avoided roe deer through spatial partitioning. We suggest that hares may adapt their spatiotemporal behavior to avoid potential predators and competitors.
... 4) 'Flushed and shot fired': two observers approached a hare using the UHF beacon, homing in on the hare and fired a shotgun shot in the air when the hare escaped, simulating a hunting situation. All disturbance experiments were conducted during the late morning, between 09:30 and 13:00 h, when hares were typically inactive (Schai-Braun et al. 2012, Mayer et al. 2018) and people are typically active, and the exact time of the disturbance was recorded (or arbitrarily assigned during the late morning for controls). The duration of each disturbance ranged between 5 and 10 min, depending how quick we detected the hare (independent of the disturbance type). ...
Article
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Capture and handling of wildlife is an important component of wildlife studies, and hunting can be a central tool for wildlife management. However, human-caused disturbance of animals can cause various negative effects on individuals. Thus, an increased understanding of different disturbances on animals will allow improved mitigation of human stressors for wildlife, and provides the basis for data-censoring when using information obtained from captured individuals. Here, we investigated the effects of capture and handling, as well as experimental disturbance, on the movement behavior of GPS-collared European hares Lepus europaeus. Of 28 hares captured in box traps, three died during handling to fit GPS collars, likely due to acute stress. Apart from an 11% decrease in activity in both sexes the first four days after capture compared to later, capture events had no significant effects on subsequent movement behavior. Hares that were disturbed experimentally, i.e. flushed with or without a shotgun shot fired, moved on average (± SD) 422 ± 206 m directly subsequent to the disturbance, leading to a spatial displacement of their short-term home range and an increased daily home range size on the disturbance day. Home range sizes returned to their before disturbance size on the following days, but hares remained further from field edges and spent more time in short vegetation in the days after simulated hunting, though this effect was comparatively small. Overall, our findings indicate that hares only marginally changed their movement behavior in response to short-term disturbances. Therefore, capture and hunting disturbance should not have severe negative effects on the movement behavior of individuals, but future studies should aim to reduce acute capture-related stress to avoid mortalities. We recommend that researchers should censor the first four days after capture from their analyses to avoid using potentially biased data.
... Pépin and Cargnelutti (1994) also found that the average daily activity of the hares in winter started near sunset and ended near sunrise. During summer, European hares had two distinct peaks in their daily activity before or after sunset and sunrise, but peaks were also recorded in full daylight (Schai-Braun et al. 2012). However, it is known that any species within the Lepus genus is not territorial (Mori et al. 2020). ...
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The Spišská Magura mountain range, located in the Middle Spiš, is one of the regions in Slovakia most contaminated by heavy metals resulting from mining and smelting activities. Heavy metals and other potentially toxic elements have accumulated in mountain areas via atmospheric transport. The influence of the daily range size of the European hare on its contamination by heavy metals was investigated in three habitat types (forest, woodland edge, meadow) in the Spišská Magura mountain range in the West Carpathians. Individual hares (n = 21) were traced and located by GPS following snowfall. Pair samples of their faeces (n = 64) and food (n = 64) were collected from feeding sites. The maps created were used for determination of the size of the daily range as being small or large. All hares that have a small daily range avoid meadows and open spaces due to the higher predation risk. However, individuals with a large daily range feed in all habitats, including meadows. Hares with a small daily range in a forest habitat ingested higher amounts of bio-elements Ca, Cr, S, and Mn as well as higher amounts of heavy metals Ba and Pb than hares with a large daily range. Moreover, dominant hares with a small daily range, with access to abundant food sources in a forest habitat, may gradually take on higher levels of bio-elements including heavy metals that are present in their food source. In contrast, in the woodland edge, hares with a small daily range had a smaller concentration of Ca, Cr, Mn, S, Ba, and Pb compared to hares with a large daily range. Caecotrophy plays a very significant role as far as the intake of nutrients and other elements is concerned. We found significant dependence between concentrations of the elements Cr, S, Ba, Pb, and Cd in the food of European hares and in their faeces.
... (Arias-Del Razo et al. 2011). This bimodal pattern has been also recorded for other genera of lagomorphs (Southern 1940;Schai-Braun et al. 2012;Oberosler et al. 2017), which may have evolved a trade-off between mostly nocturnal behaviour and the need to avoid their main predators (Carbone 2019). For instance, the wild rabbit shapes its spatiotemporal behaviour to limit encounters with its predators, e.g. the red fox and the Iberian lynx Lynx pardinus (Southern 1940;Penteriani et al. 2013). ...
Article
Assessing wildlife activity rhythms is crucial to design effective management programmes for invasive alien species. Among the most widespread alien mammals in Italy, the Eastern cottontail Sylvilagus floridanus has been introduced from North America for hunting purposes and it is now very common in the Northern and Central regions. In this study, we conducted a camera-trapping survey to determine seasonal patterns of activity rhythms of this small lagomorph in Northern Italy, as well as the overlap of temporal rhythms with its main mammal predators. Eastern cottontails were mostly crepuscular and nocturnal throughout the year. Inter-seasonal overlaps of activity patterns were always lower than 75%, as activity of the Eastern cottontail was linked to photoperiod. The bimodal activity, with significant peaks at dawn and dusk, may represent an adaptation of this species to limit encounters with nocturnal predators, i.e. red foxes Vulpes vulpes and stray cats Felis catus in our study area. Although activity increased during the first hours of daylight during the mating season, the temperature seems to be not correlated to cottontail activity. Cottontails responded to the increased predation risk on bright moonlight nights, when red foxes and stray cats were most active, by significantly reducing their activity in the full moon throughout the year and in every single season. Conversely, spatial overlap between cottontails and predators was high. We suggest that individual counts and direct removal methods (i.e. firearms or trapping) for this alien lagomorph should be carried out at dawn or at dusk, when the probability to spot active cottontails is the highest.
... The 31 habitat types used to classify the study area's land use, a classification into seven categories and their area covered in percent before, during and after harvest in Lower Austria in the year 2018 up hares near the transects for faecal pellets. Due to the increased locomotor activity of European hares during night and dawn(Schai-Braun et al. 2012), we started early in the morning (04:45 am) collecting fresh faecal pellets and stopped when identification of freshness was no longer possible (10:30 am) to avoid further bacterial degradation of fGCMs ...
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Anthropogenic disturbances, such as habitat modifications and machines, are associated with increased levels of faecal gluco-corticoid metabolites (fGCMs) in mammals, an indicator of a stress response. One human-caused process provoking incisive habitat alterations is harvesting arable crops. We investigated the effect of cereal harvest on fGCM concentrations in European hares (Lepus europaeus) in arable landscapes in lower Austria during the year 2018 by collecting 591 faecal samples before, during and after cereal harvest. fGCMs were analysed using an enzyme immunoassay, and data were analysed using linear mixed-effects models. We found that neither cereal harvest nor farming practice (organic vs. conventional) caused an overall increase in the hares' stress level. Lower vegetation density and higher proportions of bare ground were negatively correlated with fGCM concentrations, whereas the proportion of stubble fields was significantly positively correlated with fGCM concentrations in European hares. A change to more open landscapes might decrease time spent avoiding predation, and fallen grains may provide a beneficial additional food source for the hares. This indicates that European hares are well adapted to an opening up of the landscape and short-term disturbances such as cereal harvesting. In conclusion, cereal harvest had no large impact on European hares' adrenocortical activity in an arable landscape with small average field size and enough available non-farmed areas.
... We found that most species were either nocturnal (small mammals) or facultatively nocturnal (pumas, foxes, hares, rabbits), meaning they can shift diel activity according to environmental/biological cues. Guanacos were the only diurnal species, as reported previously [95] and hares showed some selection for crepuscular periods, which has been described elsewhere [96,97]. ...
Article
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Coexistence between species with similar ecological niches implies species must segregate along one or more niche axes to survive. Space, time, and trophic resources are regarded as the principal axes upon which species segregate. We examined segregation along these niche axes to determine mechanisms underlying coexistence between the two main predators, puma (Puma concolor) and culpeo foxes (Lycalopex culpaeus) in the Andes of Central Chile. We used occupancy modeling to examine space use and overlap, Kernel Density Estimation to determine temporal activity patterns and overlap, and analysis of prey remains in feces to assess diet breadth and similarity. We found high spatial overlap and positive associations between detection of the carnivores lending little support for spatial segregation. Similarly, we found high nocturnal, temporal overlap between pumas and foxes that matched peaks in activity of prey. In contrast, we found relatively low dietary overlap indicating niche segregation likely occurs along the dietary axis. The Puma diet was dominated by introduced, exotic hares and foxes appeared to shift away from hares to rabbits, small mammals, and seeds. Given that lagomorphs are the main dietary resource for pumas in particular, management decisions regarding the control or eradication of such exotic species could negatively affected puma survival.
Article
Selecting appropriate candidates for genetic rescue mostly relies on previous genetic research and monitoring, while ecological and behavioural traits of the remnant and source populations are rarely considered for such conservation measures. Because of their slow recovery, Eurasian lynx Lynx lynx populations in Central and Western Europe have been a repeated target of genetic reinforcements and reintroductions in the past 50 years. Once inhabiting much of south-eastern Europe, the Balkan lynx L. l. balcanicus is now critically endangered and confined to a small population. Long-term isolation has caused loss of genetic diversity and has possibly led to inbreeding depression. Immediate actions need to consider genetic reinforcement to increase the genetic diversity and secure population viability. Here, we compared the Balkan lynx with two neighbouring populations: Dinaric population originating from the Carpathian subspecies (L.l. carpathicus) and Anatolian population of Caucasian subspecies (L.l. dinniki) to determine is better suited source from an ecological standpoint. Main findings suggest that the L.l. carpathicus is ecologically more similar to the L.l. balcanicus and therefore likely better suited for the environment of south-western Balkans on the basis of prey preference (roe deer being the main prey), local prey availability (lower lagomorph and higher ungulate availability) and habitat use (predominant use of the mixed and broadleaved forests). We discuss the contrasting results of genetic and ecological analyses from both the evolutionary and conservation perspective and provide potential solutions that would take into account both aspects to pave the road towards potential genetic rescue of the Balkan lynx.
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ContextRoads are ubiquitous in human inhabited landscapes, and can impact animal movement and population dynamics, due to barrier effects, road mortality, but also by providing resources at road verges. Thus, we need a better understanding of how roads, in interaction with seasonal changes in habitat structure, affect space use and habitat selection of the animals that persist in these landscapes.Objectives Here, we used the European hare ( Lepus europaeus ) as model species to investigate how human-induced changes in landscape composition – measured as road density, land cover type, and field size – affect home range location, seasonal habitat selection and road crossings, which are likely to correlate with wildlife-vehicle collision risk.Methods We collected >240,000 GPS positions of 90 hares from three populations (one in Denmark and two in Germany) that differed regarding agricultural intensification and road density. Using this data, we analyzed home range location and habitat selection (using step-selection functions) in relation to roads, habitat composition, and seasonality, and quantified how these factors affected road crossings by hares.ResultsIn comparatively more heterogeneous landscapes, hares established home ranges in areas with lower road densities compared to the surrounding area, but not in more simple landscapes. Moreover, hares generally avoided main roads and selected for minor roads during the vegetation growth seasons, especially in areas with comparatively less heterogeneous habitat structure. After accounting for main road density, the number of main road crossings was comparatively higher in simpler landscapes, and the number of daily main road crossing changed seasonally.Conclusions Our findings emphasize that it is important to distinguish between road types, as different roads can have different impacts on animals (e.g. small roads providing foraging opportunities via roadside vegetation and large roads being avoided). Moreover, animals in comparatively more heterogeneous landscapes are better able to adjust their habitat selection to avoid main roads than animals inhabiting simpler landscapes. More generally, homogenous landscapes increase the space use requirements of animals, leading to increased probability of road crossings, which in turn might affect population dynamics via increased road mortality risk.
Article
Agricultural land‐use practices have intensified over the last decades, leading to population declines of various farmland species, including the European hare (Lepus europaeus). In many European countries, arable fields dominate agricultural landscapes. Compared to pastures, arable land is highly variable, resulting in a large spatial variation of food and cover for wildlife over the course of the year, which potentially affects habitat selection by hares. Here, we investigated within‐home‐range habitat selection by hares in arable areas in Denmark and Germany to identify habitat requirements for their conservation. We hypothesized that hare habitat selection would depend on local habitat structure, that is, vegetation height, but also on agricultural field size, vegetation type, and proximity to field edges. Active hares generally selected for short vegetation (1–25 cm) and avoided higher vegetation and bare ground, especially when fields were comparatively larger. Vegetation >50 cm potentially restricts hares from entering parts of their home range and does not provide good forage, the latter also being the case on bare ground. The vegetation type was important for habitat selection by inactive hares, with fabaceae, fallow, and maize being selected for, potentially providing both cover and forage. Our results indicate that patches of shorter vegetation could improve the forage quality and habitat accessibility for hares, especially in areas with large monocultures. Thus, policymakers should aim to increase areas with short vegetation throughout the year. Further, permanent set‐asides, like fallow and wildflower areas, would provide year‐round cover for inactive hares. Finally, the reduction in field sizes would increase the density of field margins, and farming different crop types within small areas could improve the habitat for hares and other farmland species.
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In agrocoenoses in S Moravia, Czechoslovakia, in winter and spring the foraging activity of the hare begins between 1600 and 1700 hrs and ends round 0700 hrs. The period of daily rest lasts 9-10 hr. In summer the boundaries between the periods of activity and rest are indistinct, a considerable part of individuals being active throughout the day, except for c4 hr in the middle of the light phase of the day. During the light phase, hares spend 18% of their time foraging in spring, 29% in summer; in autumn and early winter (before the winter solstice) they forage only at night. After the winter solstice, foraging takes up 8.5% of the time in the light phase of the day and 61.5% at night. -from Author
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Daily movements during winter of 2 adult male (Nos 1 and 3) and 2 adult female (Nos 2 and 4) radiotracking hares Lepus europaeus Pallas, 1778 were estimated from simultaneous bearings using a 50 x 50 m grid.
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Activity patterns of 15 female black bears (Ursus americanus) were studied using motion-sensitive radiotransmitters during late summer and autumn of 1990 and 1991 in La Mauricie National Park, Québec. Female black bears primarily were diurnal during this period, and no difference in activity patterns was detected between solitary females and females with young. Onset of activity followed sunrise by an average of 30 min, and cessation occurred on average 141 min after sunset. Activity and resting periods averaged 245 and 57 min, respectively. The proportion of active bears was highest during the ripening time of berries (Rubus idaeus, Vaccinium myrtilloides, and Prunus virginiand) in late August and of beechnuts (Fagus grandifolid) in early October. Bears denned earlier in 1991 than in 1990, probably because of the poor beechnut crop.
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The rimes of 241 entales into and 573 exits by brown hares from their forros on a 65 ha area of the Somerset Levels, South West England, were recorded over a ten year period to reveal the variation during the year of the duration of the daily activity period. In December when nightlength was l6 h, activity was exclusively nocturnal; intervals between entry and sunrise were longer and less consistent than those between sunset and exit. In June with nightlength down to 7.4 h, activity was part diurnal, post-sunrise and pre-sunset, for a total of 6 h. From a peak duration of 14.5 h in December, the activity period declined to 12 h in the third week of March, mirroring night time duration. It then increased to 13.5 h in late midsummer before again reducing in the autumn. Ir is suggested that the proximate cause of this circannual cyclicity is an aversion to daylight activity.
Article
A comparison was made between the responses to live-traps of marked and unmarked individuals of Microtus pennsylvanicus and Peromyscus leucopus. Previously captured individuals of M. pennsylvanicus were more readily captured than were the unmarked ones, even after an interval of a month. There was no difference in the rates of capture of marked and unmarked P. leucopus. Comparisons of a given live-trapping technique utilized in two different habitats (marsh and old field) indicated no significant difference in its effectiveness after the second day of trapping. M. pennsylvanicus, in an old field, was found to abandon diurnal activity in favor of nocturnal and/or crepuscular activity when the temperature rose above 20⚬ C. During the winter, when the temperature dropped below 0⚬ C, activity was much less at all times. Except for less activity during the days the temperature rose above 20⚬ C, such responses were not observed in the marsh. Blarina brevicauda was found to be more active on cloudy days than on those that were sunny or rainy. No correlations between activity and temperature variations was observed for this species.