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The importance of diurnal and nocturnal activity and interspecific interactions for space use by ants in clear-cuts


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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 P oland 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. Interspecific interactions and the effect of plant coverage were also considered. 3. Abundances of F ormica fusca L innaeus and red wood ants were higher during the day, whereas M yrmica were more common at night. F ormica fusca , L asius and red wood ants were more common at the clear‐cut edge than in the interior. M yrmica 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 L asius , whereas they were neutral for F. fusca . The negative association of red wood ants and M yrmica 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 reflect the real structure of the community.
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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
1,2and P I O T R ´
21Department of Ecology, Swedish
University of Agricultural Sciences, Uppsala, Sweden and 2Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw,
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. Interspecic 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 reect the real structure
of the community.
Key words. Competition, disturbance, forestry, temperature.
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 simplied 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:
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; ´
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 inuencing 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; ´
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; ̇
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 veried 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 specically, 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 inuencing 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 benet
from the temporal disappearance of the dominants and increase
in numbers, reecting the predator release effect of Russell et al.
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.
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 ´
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-inated 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% condence intervals not overlapping zero were considered
to be signicant. 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).
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 signicantly 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 signicant 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 signicant (Table 1).
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 conrmed 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 conrm some of
the hypotheses presented earlier, as red wood ants appeared to be
signicantly 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; ´
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 ´
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% condence intervals are given for each predictor and ΔAIC of the null model within each set of competing
models is given in rst column. Condence 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 conrmed 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 signicance);
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, conrming the results of former studies
(Savolainen & Vepsäläinen, 1988, 1989; ´
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 modied 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 reect 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 conrmed 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 ´
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.
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 reect the real structure of the community.
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
... Indeed, animal species respond to local-scale environmental change by changing their foraging activity (Gordon 1991). In warmer habitats, species that are physiologically incapable of withstanding the new conditions may shift from foraging in more open microhabitats to foraging in less open microhabitats and from foraging diurnally to foraging nocturnally (Cros et al. 1997;Lázaro-González et al. 2013;Zmihorski and Slipinski 2016). Analyses of foraging activity could reveal the basic mechanisms employed by animal species to cope with the new environmental conditions created by disturbance. ...
... Interestingly, species replacement might have detrimental effects on ecosystem functions (Oliveira et al. 2017;Arnan et al. 2018a) and services, including ant-mediated seed dispersal (Leal et al. 2014) or protection of plants against herbivores Câmara et al. 2018). We also found marked differences in the ant communities present at different times of day, a pattern that has been observed in other regions of the world (Hölldobler and Wilson 1990;Andersen et al. 2013;Houadria et al. 2015;Zmihorski and Slipinski 2016), with ants being mostly diurnal. Temporal differences in foraging activity seem to be related to species physiological limits: nocturnal ants have lower thermal tolerance than do diurnal ants (García-Robledo et al. 2017). ...
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This study examined how chronic anthropogenic disturbance impacts the spatiotemporal dynamics of ant foraging activity and the role played by behavioral traits. Ten plots (0.1 ha) along a gradient of chronic disturbance intensity were sampled in Catimbau National Park (Caatinga vegetation, Brazil). Vegetative structure, ground surface temperature, and ant communities in shaded and sun-exposed microhabitats were characterized during the day and at night. Each ant species’ degree of nocturnality and shaded microhabitat use were determined. Along the disturbance gradient, the frequency of sun-exposed microhabitats increased, as did the daytime ground surface temperatures; also, community composition, but not ant abundance or species richness, changed. Independent of disturbance intensity, community composition differed between day and night, and ant abundance and species richness were higher during the day. Interestingly, most species did not display strictly diurnal habits, nor did they avoid foraging in sun-exposed habitats. However, species common in more disturbed areas were more diurnal and used sun-exposed microhabitats more than species common in less disturbed areas. Many species displayed marked behavioral plasticity that was unrelated to disturbance intensity. Disturbance intensity did influence shaded microhabitat use but not the degree of nocturnality. We conclude that Caatinga ants are already morphologically, behaviorally and physiologically adapted to harsh environmental conditions; that species with different behavioral traits replace each other along the disturbance gradient; and that more plastic species can persist by shifting their microhabitat use.
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Ants disperse seeds of many plant species adapted to myrmecochory. While advantages of this ant–plant mutualism for myrmecochorous plants (myrmecochores) have been previously studied in temperate region mostly in forests, our study system was a pasture. Moreover, we used a unique combination of observing the effect of ant‐activity suppression on ant dispersal and comparison of the contribution of ant and unassisted dispersal to the distance from mother plant. We established plots without and with ant‐activity suppression (enclosures). We offered diaspores of a myrmecochorous ( Knautia arvensis ), and a non‐myrmecochorous ( Plantago lanceolata ) species in a choice test and followed ants carrying diaspores during days and nights (focus of previous studies was on diurnal dispersal). We measured frequency and distances of ant dispersal and compared them with unassisted dispersal recorded using sticky trap method. The dispersal frequency was lower in enclosures (3.16 times). Ants strongly preferred diaspores of the myrmecochore to non‐myrmecochore with 586 and 42 dispersal events, respectively (out of 6400 diaspores of each species offered). Ant dispersal resulted in more even and on average longer distances (maximum almost tenfold longer, 994 cm) in comparison to unassisted dispersal. Ant dispersal altered the distribution of distances of the myrmecochore from roughly symmetric for unassisted dispersal to positively skewed. Ants dispersed heavier diaspores farther. Ants dropped the majority of diaspores during the dispersal (which reduces clustering of seeds), while several (11%) were carried into anthills. Anthills are disturbed microsites presumably favorable for germination in competitive habitats. Ants provided non‐negligible dispersal services to myrmecochorous K. arvensis but also, to a lesser extent, of non‐myrmecochorous P. lanceolata .
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The mobilization strategies of ants have been studied quite well, but the questions of how far foragers of different species are able to move away from the nest remain unclear. The study of changes in foraging strategies depending on the type of habitat remains relevant. The aim of the work is to study mobilization strategies in 31 ant species. The study was conducted in 2019-2021 on the territory of 2 countries – Ukraine (Kyiv region and Kyiv) and Uzbekistan (Tashkent region, Tashkent). Pairs of baits (one carbohydrate and one with tuna) were laid out at a distance of 3 m from each other, in the form of transects. In total, 16 transects (417 pairs) were laid out in Ukraine in 9 types of habitats, in Uzbekistan – 5 transects (70 pairs of baits) in one type of habitats. The number of ants on each type of bait was recorded every 10 minutes, for 0-90 minutes. The distance to the nest from where the mobilization took place was also determined. It has been established that all ant species can be divided into 4 clusters according to the average distance to the nest from which foragers mobilize on the bait. Cluster 1 included 3 species of dominants, which were able to move away from the nest at a distance of up to 50 m, cluster 2 included 4 species of dominants, whose foragers could move up to a distance of 20 m. Cluster 3 included 23 species that moved away from nest at a distance of 0.2-2.0 m, cluster 4 – 1 species, foragers of it moved to a distance of up to 7 m. Preferences of bait types were noted in 15 ant species. The distance to the nest (F=9.02, p<0.001) had the greatest influence on the number of ants on baits among the considered factors, followed by species of ants (F=6.75, p<0.001) and habitat type (F=4.17, p<0.001). In habitats where an ant species mobilizes a smaller number of foragers, they have to travel, on average, long distances to a food source. Consequently, the abundance of food resources in the habitat of ants is determined by the average distance of mobilization from the nest – the smaller it is, the more resources.
The existence of a synthetic program of research on what was then termed the “nocturnal problem” and that we might now call “nighttime ecology” was declared more than 70 years ago. In reality, this failed to materialize, arguably as a consequence of practical challenges in studying organisms at night and instead concentrating on the existence of circadian rhythms, the mechanisms that give rise to them, and their consequences. This legacy is evident to this day, with consideration of the ecology of the nighttime markedly underrepre-sented in ecological research and literature. However, several factors suggest that it would be timely to revive the vision of a comprehensive research program in nighttime ecology. These include (i) that the study of the ecology of the night is being revolutionized by new and improved technologies; (ii) suggestions that, far from being a minor component of biodiversity, a high proportion of animal species are active at night; (iii) that fundamental questions about differences and connections between the ecology of the daytime and the nighttime remain largely unanswered; and (iv) that the nighttime environment is coming under severe anthropogenic pressure. In this article, I seek to reestablish nighttime ecology as a synthetic program of research, highlighting key focal topics and questions and providing an overview of the current state of understanding and developments.
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The abiotic conditions of the desert habitat fluctuate in a circadian rhythm of hot days and cold nights. Species living in desert habitats evolved many adaptations to increase their chances of survival. However, abiotic conditions in xerothermic habitats of a temperate climate are much different. Diurnal fluctuations are not as strong, but animals have to cope with seasonal changes and hibernate during the winter, which may potentially influence their adaptations to critical temperature conditions. We attempted to assess heat resistance adaptations using the example of a widely distributed xerothermic ant Formica cinerea. Using Real-Time PCR, we measured the expression of three heat shock protein genes (Hsp60, Hsp75, Hsp90) and assessed the adaptations of F. cinerea to enable foraging in risk prone conditions. The analysis of gene expression using the Generalized Linear Model surprisingly indicated that there was no significant effect of temperature when comparing workers from the control (23ºC) with workers foraging on the surface of hot sand (47-54ºC). As a next step we tried to estimate the threshold of a thermal resistance with the use of thermal chambers. Expression of all Hsps genes increase compare to the control group, expression of Hsp60 and Hsp90 continued up to 45ºC.
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In forest ecosystems in the temperate and boreal zones in Europe, red wood ants (RWA, Formica rufa group) have a significant affect as predators and competitors in communities of ground-dwelling arthropods. Therefore, the spatiotemporal distribution and abundance of RWA affect the distribution of many other species. The hypothesis that a reduction in the abundance of RWA in clear-cut areas enables other arthropods to increase in abundance was tested. The study was conducted in NW Poland in 2007 and 2008. A total of 276 1×1 m plots were sampled and 1,696 individuals recorded. The probability of the occurrence of RWA decreased significantly towards the center of clear-cut areas and increased with increasing plant cover. The frequency of Lasius platythorax, Formica fusca and spiders in the plots significantly increased towards the edge of a clear-cut area. Moreover, the occurrence of L. platythorax was negatively associated with the presence of RWA, while that of the Myrmica species was positively associated. The effect of the distance to the edge of a clear-cut area seems to be much more pronounced than the effect of RWA. This suggests that the arthropods studied prefer habitats close to the edge that are utilized by RWA than RWA-free sites located in the centre of clear-cut areas.
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The question if dominant ant species affect habitat use of other ant species was studied around two mounds of the territorial Formica polyctena. We sampled foraging ants at 10, 30 and 60 m from the mounds in four vertical layers on rocky outcrop interspersed with small vegetation patches and in neighbouring forest. We tested expected interactions among ant species on the basis of a linear competition hierarchy concept consisting of three levels: territorial (top competitors), encounter (aggressive but nonterritorial) and submissive (nonaggressive) species. We focussed on resource partitioning by space (not by food) and shifts in use of vertical layers of the habitat in presence of the territorial species. F. polyctena was present everywhere except in the litter. Its numbers decreased with distance from its mound, although its activity was substantially patchy within each distance zone. The encounter species occurred occasionally in places where F. polyctena was scarce. The submissive F. fusca, morphologically similar to the top dominant, did not respond by layer shifts; but its numbers decreased toward the mound of F. polyctena. The submissive Myrmica shifted from surface of ground to litter and shrub layers at high densities of the dominant. Small colonies and short foraging distances of the submissive species allow coexistence within the territory in low-density patches of F. polyctena. Our ant community consists of three functional guilds corresponding to taxonomic and morphological guilds: the larger above-ground Formicinae, the smaller and compacter Formicinae, and the small litter-inhabiting Myrmicinae. Interference competition is stronger and more effective among the Formicinae than among the Myrmicinae or between the subfamilies, but the top dominant affects all ant species of the community. Coexistence between the submissives and the top dominant is facilitated by niche differentiation and behavioural responses in the presence of the top dominant.
Tools for performing model selection and model averaging. Automated model selection through subsetting the maximum model, with optional constraints for model inclusion. Model parameter and prediction averaging based on model weights derived from information criteria (AICc and alike) or custom model weighting schemes. [Please do not request the full text - it is an R package. The up-to-date manual is available from CRAN].
Temperature pervasively impacts the phenotypes and distributions of organisms. These thermal effects generate strong selective pressures on behaviour, physiology, and life history when environmental temperatures vary over space and time. Despite this fact, progress toward a quantitative theory of thermal adaptation has lagged behind empirical descriptions of patterns and processes. This book draws on current evolutionary paradigms (optimization, quantitative genetics, and genetic algorithms) to establish a theory of thermal adaptation. It initially focuses on simple models that describe the evolution of thermosensitivity, thermoregulation, or acclimation. Later chapters focus on more complex models describing the coadaptation of traits or the coevolution of species. Throughout the book, various lines of evidence are used to question the major assumptions of these models. Furthermore, the predictions of these models are confronted with experimental and comparative data. Empirical examples represent a wide range of taxa, including bacteria, plants, fungi, and animals. The result is a synthesis of theoretical and empirical studies of thermal biology that offers insights about evolutionary processes.
Keeping cool Silver ants inhabit one of the hottest and driest environments on Earth, the Saharan sands, where most insects shrivel and die moments after contact. Shi et al. show that the triangular shape of the silver hairs that cover their bodies enables this existence. The hairs both increase the reflection of near-infrared rays and dissipate heat from the ants' bodies, even under full sun conditions. Evolution's simple solution to intense heat management in this species could lead to better designs for passive cooling of human-produced objects. Science , this issue p. 298
NOTE! THE AUTHORS HAVE NO PRIVATE FILE OF THE BOOK TO SHARE This book on ants (Hymenoptera: Formicidae) is an updated, corrected and expanded version of the previous monograph of the Polish myrmecofauna (Czechowski et al. 2002). It contains 103 ant species of 25 genera reported in Poland until 2010, the occurrence of which in the country are unquestionable or at least considered by the authors probable. The book consists of four main chapters: (1) a systematic checklist of the ant taxa (subfamilies, genera, species) of Europe, (2) a faunistic catalogue of the ants of Poland, (3) characteristics of the myrmecofauna of Poland, and (4) keys for identification. The checklist is the newest updated list of the European ants, and it displays recent alterations in the ant taxonomy. It contains nine subfamilies, 57 genera and 613 valid species; the list is complemented by 10 recognised but not yet formally described species. The faunistic catalogue provides a taxonomic survey of the Polish ant species together with information about their general distribution in the Palaearctic and the distribution in Poland, with notes on the biology of each species. For some taxa (species, genera), notes on their taxonomic history are provided. The chapter compiles all faunistic data published until the end of the year 2010; published reports are supplemented by confirmed unpublished data available to the authors. For every species maps of their ranges in the Palaearctic and distributions in the geographical regions in Poland are enclosed. The catalogue is closed by a list of the species ever reported in Poland and later owing to a specified reason excluded from the Polish fauna. The next chapter includes zoogeographical and ecological characteristics of the Polish myrmecofauna, with reference to the European myrmecofauna as a whole. The last chapter includes keys for identification of the ant taxa (subfamilies, genera and species), individually for workers, queens and males (when distinguishable). The keys for subfamilies and genera involve all European taxa of these ranks. The keys for species include all ant species known in Poland, and those not reported from Poland but present in adjacent regions of Central Europe, and recog nised as possible to be found in the country. The keys are illustrated with SEM photographs. The main chapters are preceded by the foreword presenting general trends in myrmecological research in the world throughout history, and the introduction, in which the past and the present of Polish myrmecology is outlined.