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Ecological Entomology (2016), 41, 276– 283 DOI: 10.1111/een.12297
The importance of diurnal and nocturnal activity
and interspecific interactions for space use by ants
in clear-cuts
MICHAŁ ̇
ZMIHORSKI
1,2and P I O T R ´
SLIPI ´
NSKI
21Department of Ecology, Swedish
University of Agricultural Sciences, Uppsala, Sweden and 2Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw,
Poland
Abstract. 1. Full sunlight conditions in open clear-cuts may limit the activity of ants
as soil surface temperatures reach lethal levels. Therefore, differences may be expected
between the diurnal and nocturnal activity of ants, and in the interactions between ant
species. These predictions, however, have been poorly investigated so far.
2. The circadian activity of ants in clear-cuts in managed forests in Poland was
investigated. Repeated counts of ants were performed during the day and the following
night at the clear-cut edge and in the clear-cut interior. Interspecic interactions and the
effect of plant coverage were also considered.
3. Abundances of Formica fusca Linnaeus and red wood ants were higher during the
day, whereas Myrmica were more common at night. Formica fusca,Lasius and red wood
ants were more common at the clear-cut edge than in the interior. Myrmica showed the
opposite pattern, but at night, its numbers increased at the edge. Plant coverage positively
affected F.fusca and red wood ants.
4. Red wood ants tended to be negatively associated with Lasius, whereas they were
neutral for F. f u s c a . The negative association of red wood ants and Myrmica was stronger
during the day compared to night.
5. The time of day was a strong driver of ant activity in the clear-cuts, whereas the
distribution of red wood ants was of lesser importance. It is concluded that circadian
activity may substantially contribute to niche separation between coexisting species,
therefore, studies performed exclusively during the day cannot reect the real structure
of the community.
Key words. Competition, disturbance, forestry, temperature.
Introduction
Commercial timber harvesting in managed forests severely
affects the ecosystem – removing the previously existing tree
stand creates a totally new habitat. Silvicultural treatments, such
as clearing and thinning, create locally open and semi-open
patches within the tree stand. These habitats are markedly
different from the surrounding forest and have different biotic
and abiotic conditions (Aussenac, 2000). After clear-cutting, the
ecosystem is substantially simplied and the three-dimensional
structure built by the tree stand is almost completely reduced
Correspondence: Michał ̇
Zmihorski, Department of Ecology, Swedish
University of Agricultural Sciences, Uppsala, Sweden, Box 7044, Se
75007 Uppsala, Sweden. E-mail: michal.zmihorski@gmail.com
to two dimensions (i.e. the soil surface). Moreover, the removal
of trees rapidly increases insolation, resulting in an increase of
soil surface temperatures which can exceed 50 ∘C (Aussenac,
2000; ´
Slipi´
nski et al., 2015). As a consequence, the undergrowth
vegetation suddenly exposed to sunlight changes distinctly, and
wind velocity increases (Greenaway, 1981; Retana et al., 1988;
Cros et al., 1997).
These marked changes in microclimatic conditions in
clear-cut areas considerably affect forest animals – some
species increase in number, whereas others withdraw from
such areas (Summerville & Crist, 2002; Niemelä et al., 2007;
Sorvari & Hakkarainen, 2007; ̇
Zmihorski, 2011; Johansson
& Gibb, 2012). Species associated with dense vegetation,
humid microhabitats and trees avoid such open areas and
decrease in number after tree stand removal, whereas those
276 © 2016 The Royal Entomological Society
Ants in clear-cut areas 277
requiring early successional stages, more light or higher tem-
peratures can colonise newly established habitats (Punttila,
1996; Summerville & Crist, 2002; Payer & Harrison, 2003).
One of the organisms of high biocenotic importance in boreal
forests that respond considerably to forest disturbances are
red wood ants. These ants from the Formica rufa group (e.g.
F. r u f a Linnaeus, F. polyctena Förster, F. aquilonia Yarrow) are
strong competitors and dominants in relation to other ant species
and also other invertebrates (Reznikova, 1983; Savolainen &
Vepsäläinen, 1988; Savolainen et al., 1989; Domisch et al.,
2005; Hawes et al., 2013). Large, aggressive, and dexterous
red wood ant individuals ght with other territorial ant species
and also reduce the tness of subordinate species living inside
their territories (Savolainen & Vepsäläinen, 1988; Czechowski
& Markó, 2006). Red wood ants also prey on all invertebrates
occurring in their trophic eld and interfere with other predatory
invertebrates inuencing their foraging success and space use
(Reznikova & Dorosheva, 2004; Maˇ
nák et al., 2013). Some red
wood ant species, such as F.rufa, may increase in number after
a local increase in insolation and prefer forest edge or forest
openings for nesting (Punttila, 1996). A recent study indicates
that clear-cuts may not be so poor in nutrition as previously
suggested and ants may harvest as much as 77% honeydew
mass in clear-cuts as they can harvest in mature stands (Gibb
& Johansson, 2010). Because of possible food availability and
territoriality, the red wood ants nesting at a forest/clear-cut
edge are generally present in clear-cuts. However, extreme
microclimatic conditions in clear-cuts may be unfavourable
even for thermophilic wood ants (Sorvari & Hakkarainen, 2007;
Kilpeläinen et al., 2008), which may decrease in number when
the ground surface temperature exceeds 40 ∘C(Cerdàet al.,
1997; ´
Slipi´
nski et al., 2015).
The effect of disturbances in boreal forests on ants and their
interactions with other invertebrates has received a great deal
of attention recently (Punttila et al., 1991; Punttila, 1996; Sor-
vari & Hakkarainen, 2007; Gibb & Johansson, 2010; ̇
Zmihorski,
2010; Sorvari et al., 2014). However, a substantial part of these
studies used pitfall-trapping or daytime snapshot counts as sam-
pling methods, ignoring important circadian variations in the
activity of the studied invertebrates (but see Vepsäläinen &
Savolainen, 1990). Some of the studies showed, however, that
the activity of epigeic ants is strongly temperature dependent
and may be shaped by high temperatures during the day (Cerdà
et al., 1997; Cros et al., 1997; Lescano et al., 2014). One may
expect, therefore, substantial differences between daytime and
nighttime space use by dominant red wood ants and, as a con-
sequence, interactions with subordinate species. These predic-
tions, however, have never been veried so current knowledge
on the effects of forest clearing on species distribution, abun-
dance, and interaction between ants seems to be incomplete.
In this study, we investigated the circadian activity of ants
in clear-cut areas and thus performed repeated daytime and
nighttime ant counts. More specically, we considered the
effects of time of day, the distance to old growth forest, an
abundance of competitors (red wood ants), and plant coverage
as factors possibly inuencing the local abundance of ants in
clear-cuts. We hypothesised that strong insolation (causing high
soil surface temperatures) limits the activity of red wood ants
in clear-cuts. Therefore, we expected the highest abundances of
red wood ants at clear-cut edges (partially shaded) as compared
with the interior and to nd a positive effect of local plant
coverage on the distribution of all studied species. Despite
some abovementioned arguments, we also hypothesised rather
even abundances of ants during day and night. As the last
hypothesis, we expected that subordinate species may benet
from the temporal disappearance of the dominants and increase
in numbers, reecting the predator release effect of Russell et al.
(2009).
Material and methods
Study area
The study was performed in a large forest complex (14 000ha)
located in western Poland (52.80N, 14.30E) in July of 2012
and 2013. The forest is managed for timber production and
the tree composition of the stand is dominated by Scots pine
(Pinus sylvestris), with considerably lower proportions of oaks
(Quercus spp.) and birch (Betula pendula). The forest con-
sists of semi-rich and poor habitats, with an understorey mainly
composed of mosses (Polytrichum spp.) and grasses (Cala-
magrostis spp., Deshampsia exuosa). Plant coverage in the
clear-cut areas was sparse, primarily consisting of Calama-
grostis epigejos,Convolvulus arvensis,Rubus idaeus,Senecio
vulgaris,andDryopteris carthusiana. The stand age in the for-
est ranged from 0 years at fresh clear-cuts to 150 years in the
oldest patches. Clear-cut areas with relatively freshly planted
saplings (1–2 years old) were chosen for the eld work. It was
only possible to identify invertebrates foraging on the soil sur-
face, especially at night, in young forest plantations with good
ground visibility.
Sampling
Ants were sampled in 10 clear-cuts (three clear-cuts in 2012
and seven in 2013). At each location, a pair of two parallel
transects were set up. The rst transect was placed along
the clear-cut edge, close to the tree stand (‘clear-cut edge’
transect). The second transect was placed c. 5 m towards the
clear-cut interior (‘clear-cut interior’ transect). Each transect
was composed of ten 1 ×1 m plots distributed every second
metre in a row along the transect (Fig. 1). The boundary
of each plot was delineated by a removable wooden frame.
Two 1-min counts were conducted within each plot: the rst
count was conducted during the day between 15.00 and 18.00.
After the rst count, the position of a frame was marked with
a small ag to enable detection at the same location. The
second count was conducted the following night between 22.00
and 24.00 hours. During each count, the plot was carefully
monitored and all individuals of the four ant groups present
in the plot or entering it during the count were noted: (i)
Form ica f usc a Linnaeus, (ii) Myrmica sp., (iii) Lasius sp., and
(iv) individuals of the red wood ant group. The recorded number
of individuals of each species was used as the unit of their
activity (e.g. Vepsäläinen & Savolainen, 1990). Additionally,
© 2016 The Royal Entomological Society, Ecological Entomology,41, 276– 283
278 Michał ̇
Zmihorski and Piotr ´
Slipi´
nski
Fig. 1. Schematic distribution of 1×1 m plots (placed every second
metre) in two parallel transects (5 m apart) in a clear-cut area. In total,
10 pairs of transects were set up in the study area.
random samples of individuals from the four groups were taken
to the laboratory to determine species composition according
to the key provided by Czechowski et al. (2012). During the
counts, great care was taken to avoid double counting the
same individuals and at night, two strong torches were used
(hand-held spotlight and head torch) to ensure the effective
inventory of each plot. Additionally, plant coverage (%) was
assessed for each plot and the soil surface temperature was
measured for several randomly selected plots (68 measurements
during the day and 63 measurements at night) using an infrared
hand-held thermometer (DT8750). In total, 387 1-min counts
were performed and used in further analyses (10 clear-cuts ×2
transects ×10 plots per transects ×2 counts per plot equals 400
counts, but 13 counts were omitted due to local disturbances).
Statistical analysis
First, we checked whether the abundances of the four groups
of epigeic ants during the day and at night were correlated.
For this purpose, we used Spearman’s rank correlation between
abundance during the day and at night at a given site for each of
the four studied groups of ants.
Next, we aimed to explain the abundances of the four epigeic
ant groups at all plots. The numbers of ant individuals belong-
ing to the four studied groups were used as dependent variables
in the four separate models. We used zero-inated generalised
linear mixed models with a negative binomial error distribution
and logarithmic link. In each model, we used clear-cut ID and
plot ID as two random factors. The result of a single count on
a1×1m plot was used as a basic record in the analyses (387
in total) and four characteristics, which were the attributes of
a single count were used as the explanatory variables: (i) time
of day (day versus night; xed factor, referred to as ‘T’),
(ii) location of a transect (clear-cut edge versus clear-cut inte-
rior, xed factor; ‘L’); (iii) abundance of red wood ants
(log-transformed; continuous variable, ‘R’ – not used in the
model explaining the abundances of red wood ants); and (iv)
plant coverage as % (continuous variable, ‘P’). Moreover,
we were interested in whether the effects of the three predic-
tors (L, R, and P) differ during the day and at
night. Thus, the following two-way interaction terms were con-
sidered: T ×R, T ×L, and T ×P.
For the purpose of model selection, all possible models were
considered. 𝜔AIC-based model averaging was performed within
the set of models with ΔAIC <6. Averaged parameter estimates
were used for further inferences and parameter estimates with
95% condence intervals not overlapping zero were considered
to be signicant. All the statistical analyses were done in R pro-
gram (R Core Team, 2015) with the help of the ‘glmmADMB’
and ‘MuMIn’ libraries (Barto´
n, 2015; Skaug et al., 2015).
Results
Average soil surface temperatures in clear-cuts during the day
counts were 33.8 ∘C on average and ranged from 24.0 to 53.0 ∘C.
Temperatures at night were signicantly lower (t-test, t=15.1;
d.f. =97.4; P<0.0001) and averaged 18.7 ∘C (range from 12.0
to 23.3 ∘C).
Random samples of ants taken from the clear-cuts allowed
the species to be determined. Individuals belonging to the
Myrmicinae subfamily and further described as the Myrmica
group were actually: M. ruginodis Nylander, M. scabrinodis
Nylander, M. schencki Viereck, and M. lobicornis Nylander.
The Lasius group was comprised of L. niger (Linnaeus) and L.
platythorax Seifert. The red wood ant group was represented by
F. r u f a and F.polyctena, whereas F.fusca from the subfamily
Formicinae was also present in the clear-cuts.
A total of 2662 individuals of the four studied groups of ants
were recorded and the abundance of a single group ranged from
0 to 50 individuals per plot. Red wood ants were the most abun-
dant (1093 individuals in total), followed by Myrmica (708),
F. fusca (508), and Lasius (353). Among the 387 controlled
plots, 47% were occupied (i.e. at least one individual present)
by F. fusca, 51% by Myrmica, 17% by Lasius, and 22% by red
wood ants. So red wood ants occur in high number on relatively
few plots.
The abundance of ants during the day and in the following
night was positively correlated in the case of the four groups.
However, quite a number of plots occupied by F.fusca and red
wood ants during the day were vacated by these species at night,
whereas a reverse pattern was recorded for Myrmica.Inthecase
of Lasius, the association seemed to be less clear (Fig. 2).
The time of day and location of the transect determined the
abundance of all the studied groups of ants. The abundance
of F. fusca and red wood ants was much higher during the
day, whereas Myrmica individuals were more common at night.
Form ica f usc a,Lasius, and red wood ants were more common at
the clear-cut edge than in the interior, whereas Myrmica showed
the opposite pattern (Table 1, Fig. 3). Plant coverage positively
affected the abundance of F.fusca and red wood ants but had
no signicant effect on the remaining groups. The abundance
of F. fusca species in the studied plots showed no association
with the abundance of red wood ants whereas the abundance
of Lasius tended to be negatively linked with the abundance
of red wood ants. Furthermore, the effects of transect location
and abundance of red wood ants on the Myrmica group were
© 2016 The Royal Entomological Society, Ecological Entomology,41, 276– 283
Ants in clear-cut areas 279
Fig. 2. Association between the abundance of ants (log-transformed) during the day and the following night. Slight jittering was added to avoid symbol
overplotting. Blue symbols: plots unoccupied during the day and occupied at night; beige: plots occupied during the day and unoccupied at night. Results
of Spearman’s rank correlation are given for each subplot.
different during the day and at night. During the day, Myrmica
individuals were mainly concentrated in the clear-cut interior,
whereas at night, they spread to the clear-cut edge (Fig. 3).
Furthermore, the abundance of Myrmica was negatively linked
with the abundance of red wood ant workers during the day;
however, at night, this association was less signicant (Table 1).
Discussion
In the study, the strongest effect on epigeic ants was related
to the time of day. We found that F. fusca and red wood ants
were markedly less active at night, whereas Myrmica showed the
opposite tendency. Moreover, time of day shifted the importance
of the other characteristics affecting the abundance of Myrmica
workers. We also conrmed that the abundance of red wood
ants, Lasius,andF. fusca decreased with increasing the distance
from the clear-cut edge, whereas the abundance of Myrmica
increased considerably. We failed, therefore, to conrm some of
the hypotheses presented earlier, as red wood ants appeared to be
signicantly less common at night (we expected no differences)
and had rather a weak effect on the distribution of other ants (we
expected to nd a clear predator release effect).
Two strategies to survive in a clear-cut
At open clear-cut areas with sparse vegetation, sunlight lim-
its the distribution of epigeic ants as the soil surface tempera-
ture reaches lethal levels – at least during the summer period.
For ectotherms (unable to actively control body temperature)
inhabiting sandy habitats, ground temperature is a factor limiting
activity and their survival rate (Lenoir et al., 2009). Every ani-
mal exposed to these conditions for a longer period of time faces
the threat of experiencing heat stroke (Angilletta, 2009). During
the measurements taken by us between 15.00 and 18.00 hours
in the clear-cuts, the temperature of the soil surface exceeded
50 ∘C and was above 40 ∘Cinc. 20% of the measurements. At
midday (12.00–14.00), the temperatures were probably even
higher. It is unlikely, therefore, that any of the studied species
is able to survive in open clear-cuts during full sunlight for a
longer period of time. Usually, only thermal specialists adapted
to such conditions can forage during this part of the day (Cerdà
et al., 1997; Lenoir et al., 2009; Shi et al., 2015; ´
Slipi´
nski et al.,
2015). None of the species observed in this study was a ther-
mal specialist, and the two strategies used by them to avoid
overheating were a shift in temporal activity and a shift in
spatial activity.
A clear shift in temporal activity was observed in Myrmica
workers, which were more abundant at night as compared
to the day. In open clear-cuts, Myrmica were most probably
attracted to forage nocturnally because of the markedly lower
temperatures and higher humidity. As a result of their broad
spectrum of activity, many species of Myrmica can forage
in relatively low temperatures (even around 10 ∘C) and can,
therefore, compensate for periods of low activity when it is
hot (Vepsäläinen & Savolainen, 1990; Punttila, 1996). In the
present study, the shift in the activity of Myrmica was clearly
visible – in several plots, no ants were observed during the day,
but many ants were observed in the following night (see Fig. 2).
As a consequence of this strategy, Myrmica ants occupy both the
clear-cut edge and the interior, being slightly more abundant in
the latter, and are not dependent on plant coverage.
© 2016 The Royal Entomological Society, Ecological Entomology,41, 276– 283
280 Michał ̇
Zmihorski and Piotr ´
Slipi´
nski
Tabl e 1. Averaged parameter estimates of generalised linear mixed models explaining the abundances of ants as a function of habitat characteristics.
Species Predictor B 95% CI
Formica fusca Intercept 0.55 0.13; 0.98
ΔAIC(Null) =150.1 T =night −1.73 −2.16; −1.30
L =interior −0.38 −0.67; −0.09
R −0.13 −0.40; 0.14
P 0.03 0.01; 0.04
T : L −0.28 −0.95; 0.39
T : P 0.01 −0.03; 0.04
T : R −0.61 −1.51; 0.29
Myrmica sp. Intercept −1.29 −2.09; −0.49
ΔAIC(Null) =111.9 T =night 1.47 0.93; 2.01
L =interior 1.02 0.47; 1.57
R −0.34 −0.81; 0.11
P 0.02 −0.01; 0.04
T : L −0.76 −1.29; −0.22
T : P 0.01 −0.02; 0.04
T : R 0.40 0.04; 0.75
Lasius sp. Intercept −1.73 −3.14; −0.32
ΔAIC(Null) =4.9 T =night 0.15 −0.34; 0.64
L =interior −0.94 −1.74; −0.14
R −0.88 −1.81; 0.05
P 0.02 −0.04; 0.07
T : L −0.36 −1.28; 0.55
T : P −0.03 −0.09; 0.03
T : R −0.27 −1.67; 1.12
Red wood ants Intercept −1.27 −2.82; 0.29
ΔAIC(Null) =25.9 T =night −0.29 −0.57; −0.02
L =interior −0.84 −1.25; −0.43
P 0.03 0.00; 0.06
T : L 0.32 −0.12; 0.77
T : P −0.02 −0.06; 0.01
The averaged parameter estimate and its 95% condence intervals are given for each predictor and ΔAIC of the null model within each set of competing
models is given in rst column. Condence intervals not overlapping zero are marked in bold.
Form ica f usc a and red wood ants (and partially also Lasius)
remained active mainly during the day, which is not surprising,
as their diurnal activity was conrmed earlier (Brian, 1956).
However, they shifted space use. These ants were considerably
less active in the clear-cut interior compared with its edge,
and preferred patches overgrown with vegetation. This may be
driven by preferences for shade in the vicinity of trees growing
at the forest edge. Similarly, an association with vegetation
is typical for ants looking for a place to lower their body
temperature (Cros et al., 1997).
Importance of competition
The distribution of ants in clear-cuts is not only driven by
abiotic conditions, but may be shaped by antagonistic interac-
tions between species – the competition theory predicts that red
wood ants, being at the top of the hierarchy, are a territorial and
aggressive species defending their trophic eld, and, therefore,
exclude other aggressive species (Reznikova, 1983; Savolainen
& Vepsäläinen, 1988, 1989; Czechowski et al., 2013). There
are also two levels of subordinate species: encounter and
submissive (Savolainen & Vepsäläinen, 1988). Aggressive
but non-territorial encounter species defend their nest and food
sources, and because of their competitive status, may not coexist
with territorials. Submissive species defend only their nests, and
as a result of their low aggression, may nest within the trophic
eld of aggressive species (Czechowski & Markó, 2006).
In this study, the negative effect of the dominant species (red
wood ants) on the presence of subordinate species was not
strong (see also Neuvonen et al., 2012 for similar results). Only
Lasius avoided red wood ants (on the verge of signicance);
this is most probably driven by the competitive status of both
L.niger and L.platythorax, which are aggressive encounter
species (Punttila et al., 2004). In contrast, red wood ants coex-
isted with F.fusca, conrming the results of former studies
(Savolainen & Vepsäläinen, 1988, 1989; ´
Slipi´
nski et al., 2014).
In the case of Myrmica ants, the situation was more compli-
cated. The results suggest that the distribution of these ants is
substantially modied by red wood ants – the abundance of
Myrmica increases in the area (interior) and at the time (night) of
the absence of red wood ants. The observed preference of Myr-
mica to the clear-cut interior cannot be attributed to the abiotic
conditions, which seem to be better (lower insolation and higher
food availability) at the edge. Thus, it seems that such a distribu-
tion may be driven by the avoidance of red wood ants. Myrmica
were much more active at night, which may also reect an
© 2016 The Royal Entomological Society, Ecological Entomology,41, 276– 283
Ants in clear-cut areas 281
Fig. 3. Abundance of the four groups of ants (y-axis, log-transformed) showed with kernel density estimation (irregular polygons, kernel density values
on x-axes are not shown) and mean (horizontal thick line) separately for day and night (beige and blue, respectively), as well as for clear-cut edge and
clear-cut interior (left and right columns for each subplot).
avoidance of red wood ants (Vepsäläinen & Savolainen, 1990).
Myrmica may shift to night activity at sites with a high presence
of F. polyctena (Savolainen & Vepsäläinen, 1989), and this, in
turn, may drive its lower activity on the surface during the day
as observed by us. This presumption is partly conrmed by the
strong interaction between time of day and transect location
(T : L, see Table 1), showing that at night (when red
wood ants are less active), Myrmica workers expand their space
use towards the clear-cut edge. Furthermore, Myrmica seem to
be less affected by red wood ants at night. This may be driven by
the fact that red wood ants use visual cues for enemy detection
(Dorosheva et al., 2011) and we cannot exclude, therefore,
© 2016 The Royal Entomological Society, Ecological Entomology,41, 276– 283
282 Michał ̇
Zmihorski and Piotr ´
Slipi´
nski
that they detect the enemy less effectively when it is dark.
Alternatively, the red wood ants may not be able to aggressively
interact with Myrmica because of lower temperatures at night.
Conclusions
In clear-cut areas, environmental conditions seem to be more
important for the distribution of subordinate ants than the abun-
dance of red wood ants (see ̇
Zmihorski, 2011; Neuvonen et al.,
2012 for similar results). The distance to the clear-cut edge
affected all of the studied species, and the time of day shaped
the abundances of three of them. The presence of red wood ants
seemed to be of lower importance for space use by the remaining
species of ants. What is most important, our investigation docu-
mented clear differences between diurnal and nocturnal patterns
of the distribution of epigeic ants in clear-cuts and in the interac-
tion between species. For instance, the share of Myrmica among
the four studied groups denoted 14% and 40% during the day
and at night, respectively. In contrast, Form ica f usc a constituted
31% of ants observed during the day, whereas only 7% of those
observed at night. Temporal changes in the activity of epigeic
ants in clear-cut areas constitute an important element of their
strategy, and nocturnal foraging is important for some of the
species. Moreover, differences in circadian activity may substan-
tially contribute to niche separation between coexisting species,
leading to an overall increase in diversity. We conclude, there-
fore, that investigations of epigeic ants performed exclusively
during the day can be seriously biased towards diurnal species
and will not reect the real structure of the community.
Acknowledgements
This research was funded by a grant from Iuventus Plus (IP2011
064771) funded by the Ministry of Science and Higher Edu-
cation in Poland. We are grateful to Violetta Hawro for helpful
comments on the manuscript. Barbara Przybylska kindly
improved the English. M. ̇
Z.: project design, data collection,
data analysis and manuscript writing; P. ´
S.: project design, data
collection, and manuscript writing.
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Accepted 19 November 2015
First published online 16 February 2016
Associate Editor: Adam Hart
© 2016 The Royal Entomological Society, Ecological Entomology,41, 276– 283
