ArticlePDF Available

Differential impact of two dominant Formica ant species (Hymenoptera, Formicidae) on subordinates in temperate Europe

  • Museum and Institute of Zoology (PAS)

Abstract and Figures

Competition is one of the basic mechanisms shaping ant assemblages. Dominant territorial species are known to restrictively influence the traits of subordinates in various ways. However, there could be differences in the effects of dominants due to differences in their colony structure, lifestyle and also behaviour. We tested this hypothesis in natural circumstances in an area where a Formica exsecta Nyl. supercolony neighbours a strong population of the European slave-maker F. sanguinea Latr. For the purpose of our study three different sites were selected: one dominated by F. exsecta, a neighbouring site dominated by F. sanguinea, and a third site where both species co-occurred. We analyzed the structure of subordinate ant communities based on nest counts, and by recording the activity of ants at baits. Based on our findings the structure of ant communities differed significantly among the three sites. The F. exsecta site was characterized by the lowest nest density of subordinates, especially in the case of aggressive species as Lasius platythorax Seifert. At baits even the simple presence of the dominant limited the abundance and occurrence of subordinates. In contrast, no such negative effect could be revealed at the F. sanguinea site. The community of the mixed site showed intermediate features in many respects. Generally, the supercolonial F. exsecta had a bigger influence on the ant community than the slave-making F. sanguinea. On the other hand, our findings revealed a slight protective role of the territorial F. exsecta for potential slave species to F. sanguinea. The current study underlines the importance of differences between dominant ant species in shaping differentially ant communities even within the same restricted area.
Content may be subject to copyright.
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 97
Differential impact of two dominant Formica
ant species (Hymenoptera, Formicidae) on
subordinates in temperate Europe
Gema Trigos Peral1, Bálint Markó2,3, Hanna Babik1, Ioan Tăuşan4, István Maák3,
Zsóa Pál5, Piotr Ślipiński1, Zsolt Czekes2, Wojciech Czechowski1
1 Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza St 64, 00-679 Warsaw, Poland 2Hunga-
rian Department of Biology and Ecology, Babeş-Bolyai University, Clinicilor St 5–7, 400006 Cluj-Napoca,
Romania 3 Department of Ecology, University of Szeged, PO Box 51, H-6701 Szeged, Hungary 4 Department
of Environmental Sciences and Physics, Faculty of Sciences, Lucian Blaga University of Sibiu, Dr. Raţiu St 5–7,
550012 Sibiu, Romania 5 School of Environmental Sciences, Charles Stuart University, PO Box 789, NSW
2640 Albury, Australia
Corresponding author: Gema Trigos Peral (
Academic editor: Jack Ne |Received 29 February 2016|Accepted 17 April 2016| Published 27 June2016
Citation: Trigos Peral G, Markó B, Babik H, Tăuşan I, Maák I, Pál Z, Ślipiński P, Czekes Z, Czechowski W (2016)
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae) on subordinates in temperate
Europe. Journal of Hymenoptera Research 50: 97–116. doi: 10.3897/JHR.50.8301
Competition is one of the basic mechanisms shaping ant assemblages. Dominant territorial species are
known to restrictively inuence the traits of subordinates in various ways. However, there could be die-
rences in the eects of dominants due to dierences in their colony structure, lifestyle and also behaviour.
We tested this hypothesis in natural circumstances in an area where a Formica exsecta Nyl. supercolony
neighbours a strong population of the European slave-maker F. sanguinea Latr. For the purpose of our
study three dierent sites were selected: one dominated by F. exsecta, a neighbouring site dominated by
F. sanguinea, and a third site where both species co-occurred. We analyzed the structure of subordinate
ant communities based on nest counts, and by recording the activity of ants at baits. Based on our nd-
ings the structure of ant communities diered signicantly among the three sites. e F. exsecta site was
characterized by the lowest nest density of subordinates, especially in the case of aggressive species as Lasius
platythorax Seifert. At baits even the simple presence of the dominant limited the abundance and occur-
rence of subordinates. In contrast, no such negative eect could be revealed at the F. sanguinea site. e
community of the mixed site showed intermediate features in many respects. Generally, the supercolonial
JHR 50: 97–116 (2016)
doi: 10.3897/JHR.50.8301
Copyright Gema Trigos Peral et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
F. exsecta had a bigger inuence on the ant community than the slave-making F. sanguinea. On the other
hand, our ndings revealed a slight protective role of the territorial F. exsecta for potential slave species to
F. sanguinea. e current study underlines the importance of dierences between dominant ant species in
shaping dierentially ant communities even within the same restricted area.
Formica exsecta, Formica sanguinea, baits, community structure, competition, interspecic hierarchy, nest
density, slavery, territoriality
Most of the concepts regarding the assembly rules of animal communities point to
competition as a crucial factor in shaping their composition and structure (MacArthur
1972, Wiens 1983, Farris et al. 2015, McFarlane Tranquilla et al. 2015, Sunarto et
al. 2015). is is supported for ants as well by several case studies (e.g. Savolainen
and Vepsäläinen 1988, Pisarski and Vepsäläinen 1989, Cerdá et al. 1998, Parr and
Gibb 2010, Czechowski et al. 2013, Maák et al. 2014, Ślipiński et al. 2014, Dejean
et al. 2015, Spotti et al. 2015) and the work reviewed quite recently by Cerdá et al.
(2013). Specically, ant communities in temperate zones are organized hierarchically
and a crucial role in shaping them can be attributed to ecologically and behaviourally
dominant species. Dominant species can restrictively structure local communities in
various ways, from interference and exploitative competition to direct predation on
lower-ranked species, both quantitatively (decreasing nest densities and colony sizes
of subordinates) and qualitatively (reducing species richness by competitive exclusion
and altering the dominance structure), but also with regards to the spatial patterns
of the nests (e.g. by forced resource partitioning, nest moving, etc.) (Higashi and
Yamauchi 1979, Cherix and Bourne 1980, Pisarski 1982, Pisarski and Vepsäläinen
1989, Savolainen and Vepsäläinen 1988, 1989, Savolainen et al. 1989, Savolainen
1990, 1991, Vepsäläinen and Savolainen 1990, Hölldobler and Wilson 1990, Punttila
et al. 1991, 1996, Sanders and Gordon 2003, Czechowski and Markó 2005, Adler et
al. 2007, Czechowski et al. 2013, Markó et al. 2013, Rzeszowski et al. 2013, Ślipiński
et al. 2014).
Typically, at least in Euro-Asian boreal and temperate zones, the top dominants of
ant communities are territorial species (sensu Pisarski 1982), i.e. those which actively
protect the boundaries of their whole foraging areas against other territorials (Pisarski
1980, Vepsäläinen and Pisarski 1982, Pisarski and Vepsäläinen 1989). In temperate
Europe, such territorial dominants, already well-known for their competitive impact
on subordinate ants, are the wood ant species of the Formica rufa group (subgenus
Formica s. str. L.) (Savolainen and Vepsäläinen 1988, 1989, Savolainen et al. 1989,
Savolainen 1990, 1991, Punttila et al. 1991, 1996, Czechowski and Markó 2005),
species of the subgenus Coptoformica Müll. (genus Formica L.) (Czechowski 1975,
Pisarski 1982, Pisarski and Vepsäläinen 1989), the dendrophilic Lasius (Dendrolasius)
fuliginosus (Latr.) (Czechowski et al. 2013, Markó et al. 2013, Ślipiński et al. 2014),
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 99
and Liometopum microcephalum (Panzer) (Petráková and Schlagamerský 2011). Colo-
nies of territorial species (either allo- or conspecic) do not share the same territory,
but non-territorial species can nest and forage within the dominant’s territory, how-
ever with diverse limitations. eir colonies are the centres of the spatial organization
in ant communities through their dierential eects on subordinate species, which are
‘allowed’ only to nest at certain distances from the nest of the territorial species – the
farther away are the nests of the subordinates, the stronger could be the competition
between the two species (Savolainen and Vepsäläinen 1988, Pisarski and Vepsäläinen
1989). e impact of territorials on subordinates may further be enhanced by specic
social strategies, such as the formation of new colonies through budding, which leads
to the development of a network of related nests over a larger area (so-called supercolo-
nies – see e.g. Helanterä et al. 2009, Robinson 2014 for reviews), a well-known strategy
in many territorial wood ants (Higashi and Yamauchi 1979, Cherix and Bourne 1980,
Rosengren et al. 1985), but also in members of the subgenus Coptoformica (Pisarski
1982, Czechowski 1975, 1977, Markó et al. 2012).
Within subordinates, two competition levels are distinguished: (a) encounter spe-
cies, which are fairly aggressive and usually manage to monopolize (defend) single food
sources, and (b) submissive species, which defend only their nests (Vepsäläinen and
Pisarski 1982). Although they can manage to survive in the territory of dominants,
subordinates are restrictively aected by territorials, and, in general, they fare bet-
ter outside than within territories of dominant species (Savolainen 1990, 1991). On
the other hand, in certain situations, nesting inside a dominant territory can even be
good for certain species. Members of the subgenus Serviformica For. (genus Formica)
are slave species for the facultative slave-maker Formica (Raptiformica) sanguinea Latr.
Potential slave colonies may prot from the proximity of strong dominants since these
can stop the raiding columns of F. sanguinea or force them to change their route when
protecting their own territory. Such a positive eect of territorials has been reported
for several species of wood ants of the F. rufa group (F. rufa L., F. polyctena Först., F.
aquilonia Yarr., F. lugubris Zett.) for F. fusca L. and F. lemani Bondr., and known slave
species F. sanguinea (Punttila et al. 1996, Czechowski 1999, 2000, Czechowski and
Vepsäläinen 2001, Czechowski and Markó 2006, Väänänen et al. 2010). A similar
mechanism may also work, at least to some extent, in the case of Serviformica species
enslaved by the obligate slave-maker Polyergus rufescens (Latr.) (Czechowski 2006).
e dierences among dominant species regarding their eect on subordinates
may vary depending on their lifestyle (slave-maker vs non-slave-maker), colony struc-
ture (mono- vs polydomous), and on their foraging strategies (narrow vs wide food
spectrum). e dierential eect of dominants on subordinates are best revealed when
dominant species occur syntopically (in the same habitat), consequently, the basic
species pool that could make up the ant community is the same. erefore, if the dif-
ferences between the dominants are negligible, the dierences between the ant com-
munities occurring in their territories/areas should also be minor. In order to test this
hypothesis, an appropriate natural set-up is needed, with the long-term co-occurrence
of territorial allospecic neighbours (see e.g. Czechowski et al. 2013). Recent eld
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
studies performed on the largest European polydomous system of the territorial For-
mica exsecta (Markó et al. 2012) yielded the discovery of a large population of the
slave-maker F. sanguinea, also a dominant species, neighbouring the above-mentioned
supercolony. is unique natural set-up of two dominant neighbours from the same
genus, yet with a dierent colony structure (supercoloniality vs mono- or oligodomy)
and lifestyle (non slave-maker vs slave-maker) oered the chance to perform a com-
parative study. In the framework of the current study, we proposed an analysis of the
structure of ant communities in the area of these two dominants, and formulated the
following questions: (1) are there any dierences between the structure of ant commu-
nities of the two neighbouring dominant Formica species’ territory/area, (2) are there
any dierential eects of these two dominants on the foraging strategies of subordi-
nate species, (3) do the abundance and foraging strategy of members of the subgenus
Serviformica, known as a slave species of F. sanguinea, dier within and outside the F.
sanguinea area?
Materials and methods
Study species and sites
e main subjects of the study are two dominant ant species: Formica (Coptoformica)
exsecta and Formica (Raptiformica) sanguinea.
Formica exsecta is a fairly common North-Palaearctic mound-building territorial
species, a typical forest-ecotone polytope dwelling in forest margins and clearings,
thinned young growth, overgrowing meadows and glades, etc. Its colonies number
several thousand to tens of thousands of workers and occur in two, mono- and po-
lygynous social forms. Polygynous colonies may grow through nest splitting into poly-
domous systems, often with more than 100 nests (so-called supercolonies). Addition-
ally, colonies may be established through the temporary social parasitism of members
of the subgenus Serviformica. e optimal home range of foraging workers from a
single nest does not exceed a few meters. ey are aggressive and predacious ants,
which also feed on honeydew (Pisarski 1982, Seifert 2000, Hughes 2006, Erős et al.
2009, Csata et al. 2012).
Formica sanguinea is a common South-Palaearctic species, a polytope of warm and
dry habitats, both forested and open areas; generally its ecological preferences widely
overlap with those of the former species. It nests in decaying tree stumps, partly cov-
ered with dry plant material and also in the ground, often under stones. In general,
colonies, usually functionally monogynous, number up to several tens of thousands
of workers. It is a facultative socially parasitic species which enslaves workers of the
subgenus Serviformica; the range of its slave raids may reach 100 m (usually < 50 m).
ey are very aggressive and predatory ants; during their raiding period they do not
respect the boundaries of other species’ territories (Czechowski 2000, Hughes 2006).
e territoriality of F. sanguinea is a moot point (see the Discussion).
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 101
As habitat niches of Formica exsecta and F. sanguinea overlap each other to a con-
siderable degree, colonies of these two species often co-occur.
Our study area was a vast semi-moist meadow of the Molinion caeruleae W. Koch
alliance [with, among others, Molinia caerulea (L.) Moench, Deschampsia caespitose
(L.) P. B., Festuca pratensis Huds., Nardus stricta L. and Juncus sp.], densely overgrown
with small birch saplings, located in the southern part of the Giurgeului depression
(46°36'N; 25°36'E; 780 m a.s.l.) in the eastern Carpathians, Transylvania, Romania.
e meadow is fairly intensely grazed by cows for most of the year. is is the location
of the largest known European polydomous system of Formica exsecta. e supercolo-
ny contains 3,347 permanent nests over an area of nearly 22 ha (i.e. the nest density is
ca. 153/ha) (Markó et al. 2012). A population of F. sanguinea was recently detected,
with a small number of F. cunicularia Latr. and F. rubarbis F. slaves, within the same
habitat near the area occupied by the supercolony.
ree sites were selected within the area for the purpose of our study (Fig. 1): (1)
a site clearly dominated by Formica sanguinea right at the border of the F. exsecta poly-
domous system (referred to as the F. sanguinea site; 32 F. sanguinea nests on ca. 3210
m2 with 0.01 nests/m2, and four F. exsecta nests with 0.001 nests/m2), (2) a neighbour-
ing patch within the F. exsecta territory with a single F. sanguinea nests (referred to as
the F. exsecta site; 68 nests on ca. 5780 m2 with 0.012 F. exsecta nests/m2, and one F.
sanguinea nest with 0.0002 nest/m2) and (3) a site with interspersed nests of both F.
sanguinea and F. exsecta (referred to as the mixed site; eight F. sanguinea on ca. 2685
m2 with and 0.003 nests/m2, and 80 F. exsecta nests with 0.03 nests/m2). Within the
whole study area, the meadow was homogeneous in respect of the vegetation, surface
conguration and sun exposure.
e study area was not part of any nature protection area and no specic permits
were required to carry out the eld studies.
Sampling methods
Nest densities of dierent subordinate ant species were obtained using the biocoenom-
etric method, by carefully investigating 15 quadrats of 9 m2 at each of the three study
sites. e quadrats were randomly selected by a blindfolded person in the eld, they
did not border each other, and they were at least 2 m from any Formica exsecta nest
(Fig. 1). A sample of workers was collected for precise identication from each nest
found. Identication of ant species was carried out with the key of Czechowski et al.
(2012). Field work was carried out between 17 and 21 June 2010.
Baiting is commonly used to study the foraging pattern of ants around their nests
and species’ foraging strategies. It is also an appropriate method for determining the
hierarchical positions of particular species within an assemblage (e.g. Czechowski
1979, 1985, Czechowski and Pisarski 1988, Parr and Gibb 2010, Petráková and
Schlaghamerský 2011, Markó and Czechowski 2012). As the food preferences of
ants may change seasonally, we provided a mixture of two dierent kinds of bait:
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
Figure 1. Map of the study area (F. exsecta nests outside the delineated sites are not marked out; grey
area – compact area of the F. exsecta supercolony).
tuna sh akes as a source of animal protein food and a water solution of mixed-
ower honey as a source of carbohydrates. Altogether, 10 observation plots were
distributed along linear transects at each of the three study sites (Fig. 1), and the
baits were put out in the centre of each plot, separated from the next plot by 2 m,
and placed at least 2 m from any Formica exsecta or F. sanguinea nest and any previ-
ously surveyed plot. e bait portions of ca. 3 cm diameter were placed in the centre
of a round green plastic plate of 10 cm diameter. Baiting observations were carried
out on 8 July 2010. Observations were carried out in two periods: between 1000 and
1200 AM, and between 520 and 720 PM. Each period consisted of six observations of
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 103
1 min, separated by 20 minute breaks. During each observation, the number and
species of ants appearing at the baits were recorded, with the exception of Myrmica
spp., where species level identication could not be performed under eld condi-
tions. e baits were put out 10 minutes before each period of observation (morning
and afternoon) and left there until the end of the period when they were retrieved
and the plates cleaned.
Data analyses
e dierences in the abundance of dierent ant species among sites were revealed by
the use of the Kruskal-Wallis test, followed by Tukey and Kramer post-hoc tests. e
diversity of ant communities was calculated with the Shannon-Wiener entropy index
(log2), using data for individual quadrats and baits respectively for nest count and
bait data, while dierences among sites for these parameters were revealed by using
the Kruskal-Wallis test, followed by the Tukey and Kramer post-hoc tests separately
for nest count and bait data. In order to determine the dierences among the ant
assemblages of the three sites, permutational multivariate ANOVA (perMANOVA,
10000 permutations) was applied both for nest count and bait data, while Nonmetric
Multidimensional Scaling (NMDS) was used to visualize the three assemblages, both
in the case of nest and bait data. e SIMPER test was performed to determine the
contribution of each species to dissimilarities between ant communities.
e Generalized Linear Mixed Model approach (GLMM, binomial, maximum
likelihood) was applied to analyze the eect of dominant species on the presence and
absence of subordinate species at baits. e abundance of Formica sanguinea and F.
exsecta workers present at baits were introduced as variables, along with their pres-
ence vs absence data as factors, next to the type of the site (F. sanguinea, F. exsecta,
mixed) as a factor. e period and time of the observation as well as bait ID were
introduced as nested random factors. A similar approach (GLMM, Poisson error,
maximum likelihood) was applied to separately test the eect of dominants on the
abundance of subordinates. e number of all subordinates was pooled together,
since they showed a low abundance separately (see the Results). e same model
structure was used as above.
All statistical analyses were carried out using the R 3.1.2 statistical package (R
Core Team 2015). Multiple pairwise comparisons in the case of the Kruskal-Wallis
tests concerning diversity indices were performed using the Tukey and Kramer post-
hoc test after Nemenyi with the PMCMR R-package (Pohlert 2015). Permutational
multivariate ANOVA, NMDS and the SIMPER test were carried out with the vegan
R-package (Oksanen et al. 2013). GLMMs were performed using the glmer function
in the lme4 R-package (Bates et al. 2014). Post-hoc Tukey-tests for sequential com-
parison among factor levels when performing GLMM were carried out with the glht
function in the multcomp R-package (Hothorn et al. 2015).
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
Species composition and abundance
During the quadrat survey, 171 nests were found belonging to 11 subordinate ant
species in addition to the two dominant species (i.e. Formica sanguinea and F. exsecta)
(Table 1). Five subordinate species occurred at every site studied [Tetramorium cf. caes-
pitum (L.), Myrmica scabrinodis Nyl., M. vandeli Bondr., M. schencki Viereck, and
Lasius avus (F.)]. Total nest density of co-occurring ant species was 1.8 times higher
at the F. sanguinea than at the F. exsecta site, whereas it was intermediate at the mixed
site (Table 1). e most abundant subordinate species was the subterranean L. avus
followed by M. scabrinodis on every study site (Table 1).
Only three aggressive encounter species were found: Lasius platythorax, Tetramo-
rium cf. caespitum and Formica rubarbis, and none of them were present in high
densities at the site (territory) of F. exsecta. e three sites diered mainly in the nest
density of two encounter species: L. platythorax and T. cf. caespitum (Table 1). Nest
density of the former was relatively high at the F. sanguinea site, whereas the latter had
a higher density at the mixed site. Nest density of all submissive species [Myrmica spp.,
Leptothorax acervorum (F.) and Serviformica spp.] taken together was the highest at the
F. sanguinea site, owing mainly to the trends noticeable in Myrmica species (Table 1).
e ant community of the Formica sanguinea site was signicantly more diverse
than the F. exsecta site, while the mixed site had an intermediate position (Kruskal-
Wallis χ2 = 7.83, p = 0.02; Fig. 2). e structure of the ant assemblages also diered
signicantly among the three studied sites (perMANOVA F1,40 = 3.51, p = 0.006),
mostly with regards to the ant community of the F. sanguinea site, while seemingly the
communities of the other two sites were quite similar (Fig. 3). According to the results
of the SIMPER analysis, most of the dissimilarities (ca. 70%) among the three sites
could be contributed mostly to three common species: Lasius avus, Myrmica scabrino-
dis and L. platythorax, except for the F. exsecta vs mixed site, where T. cf. caespitum
took the place of L. platythorax in this respect (Table 2).
Foraging strategies and competitive eects
Besides Formica sanguinea and F. exsecta, foragers of other species were also present
at the baits (Table 3). e highest abundance of ants at baits was observed at the F.
exsecta site, exclusively due to the ecient recruitment of F. exsecta workers to the food
sources (Table 3). As expected, F. sanguinea was present at baits in its site, though in a
surprisingly low number, whereas it was totally absent from the F. exsecta site; in turn,
F. exsecta was absent in the F. sanguinea site. Both species exploited baits, though, at
the mixed site (Table 3).
Dierent trends were detected in the abundance of two aggressive (encounter) spe-
cies: Lasius platythorax and Tetramorium cf. caespitum. e former occurred in similar
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 105
Table 1. Nest densities (mean number per 9 m2 ± SD) of subordinate ant species and their proportions
(%) in the three study sites.
F. sanguinea site F. exsecta site Mixed site
mean ± SD
mean ± SD
mean ± SD
Tetramorium cf. caespitum (L.) 0.33 ± 0.49
0.2 ± 0.56
0.67 ± 1.05
Leptothorax acervorum (F.) – 0.07 ± 0.26
Myrmica scabrinodis Nyl. 1.13 ± 1.13
0.53 ± 0.92
0.73 ± 1.1
Myrmica vandeli Bondr. 0.47 ± 0.99
0.47 ± 0.74
0.13 ± 0.35
Myrmica schencki Viereck 0.27 ± 0.8
0.07 ± 0.26
0.13 ± 0.35
Myrmica lobicornis Nyl. 0.13 ± 0.35
0.07 ± 0.26
Myrmica ruginodis Nyl. 0.07 ± 0.26
1.3 – –
Lasius avus (F.) 2.00 ± 1.81
1.33 ± 1.54
1.40 ± 0.99
Lasius platythorax Seifert 0.87 ± 0.74
0.07 ± 0.26
Formica cunicularia Latr. 0.07 ± 0.26
1.3 0.13 ± 0.35
Formica rubarbis F. 0.07 ± 0.26
All subordinate species 5.21 2.87 3.33
All epigean subordinate species 3.21 1.54 1.93
Table 2. Results of the SIMPER analysis: the overall dissimilarity between study sites, the average contri-
bution of subordinated ant species to overall dissimilarity (Dissimilarity contribution), their mean abun-
dances in the compared sites (Mean 1 and Mean 2), and the cumulative percentages of contributions.
Sites (dissimilarity) Species Dissimilarity
contribution (%) Mean 1 Mean 2 Cumulative
contribution (%)
F. sanguinea vs. F.exsecta
L. avus 20.95 2 1.54 29.88
M. scabrinodis 13.49 1.13 0.62 49.13
L. platythorax 12.27 0.87 0.08 66.63
M. vandeli 9.44 0.47 0.54 80.11
T. cf. caespitum 6.13 0.33 0.23 88.85
F. sanguinea vs. Mixed
L. avus 18.83 2 1.5 27.76
M. scabrinodis 13.11 1.13 0.78 47.1
L. platythorax 12.21 0.87 0 65.11
T. cf. caespitum 9.33 0.33 0.71 78.88
M. vandeli 6.18 0.47 0.14 88
F. exsecta vs. Mixed
L. avus 20.35 1.54 1.5 30.92
M. scabrinodis 14.38 0.62 0.79 52.79
T. cf. caespitum 11.16 0.23 0.71 69.75
M. vandeli 8.48 0.54 0.14 82.64
M. schencki 3.46 0.08 0.14 87.91
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
Table 3. Mean number of ant individuals per baits (± SD) for each study site and their relative propor-
tions (%) to other ant species at the same site.
F. sanguinea site F. exsecta site Mixed site
mean ± SD
mean ± SD
mean ± SD
Tetramorium cf. caespitum 31.6 ± 96.78
36.5 10.6 ± 21.49
Myrmica spp. 2.1 ± 3.96
0.2 ± 0.42
4.0 ± 3.83
Lasius platythorax 52.2 ± 83.48
52.2 ± 120.86
Formica cunicularia 0.6 ± 1.9
3.4 ± 4.03
1.8 ± 3.16
Formica sanguinea 0.1 ± 0.32
0.1 1.9 ± 2.56
Formica exsecta 120.4 ± 100.65
14.8 ± 35.67
All subordinate species 86.5 ± 111.43 55.8 ± 121.5 16.5 ± 22.75
99.9 31.6 49.7
All species 86.6 ± 111.35 176.6 ± 102.94 33.2 ± 36.19
100 100 100
Figure 2. Diversity of the ant communities within the study sites based on nest surveys (medians, quartiles,
min–max values and outliers). Survey performed on 9 m2 quadrats (N = 15 in each site). Dierent letters
indicate signicant dierences among groups.
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 107
Figure 3. NMDS graph of the ant communities within the study sites based on nest survey (stress = 0.172).
numbers at baits in the Formica exsecta and F. sanguinea sites, whereas it was absent
from the mixed site (Table 3). In turn, Tetramorium cf. caespitum was absent from the
F. exsecta site, while it was the second most abundant ant in the two other sites (Table
3). e abundance of potential F. sanguinea slave species, such as F. cunicularia, gradu-
ally increased from the F. sanguinea site to the mixed one and then to the F. exsecta
site (Table 3).
e highest diversity was found at baits in the mixed site (Kruskal-Wallis χ2 =
9.11, p = 0.011) followed by the signicantly lower diversities of Formica sanguinea
and F. exsecta sites, which did not dier signicantly from each other based on the
post-hoc test result (Fig. 4). ere were no dierences between the three sites in the
qualitative-quantitative composition of foraging ants (perMANOVA F1,29 = 1.83, p =
0.12), however, baits in the F. exsecta territory seemed to stand apart from all other
sites due to the marked presence of F. exsecta, while the presence of subordinate species
was more characteristic for the F. sanguinea and the mixed sites (Fig. 5).
According to the results of the GLMM analysis, Formica exsecta’s abundance (z =
–3.09, p = 0.002) and presence (z = –2.32, p = 0.02) had a signicant negative eect
on the occurrence of subordinates at baits, while the abundance of F. sanguinea did
not play a major role (z = –1.72, p = 0.08), but its presence did have a positive inu-
ence on the occurrence of subordinates (z = 2.13, p = 0.033). ere were no dierences
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
between the F. sanguinea and the F. exsecta sites with regards to the frequency of the
occurrence of subordinates (z = 1.99, p = 0.11). Also, no dierence was found between
the F. sanguinea and the mixed site in this respect (z = –1.8, p = 0.16), but subordinates
were signicantly less frequent at baits in the mixed site compared to the F. exsecta site
(z = –3.18, p = 0.004).
Generally, a similar pattern with smaller adjustments was found when analyzing
the eect of dominants on the abundance of subordinates at baits. In addition to the
abundance of Formica exsecta (z = –8.33, p < 0.0001), even its simple presence (z =
–2.01, p = 0.044) had a signicant negative eect on the abundance of subordinates.
On the other hand, F. sanguinea’s abundance (z = 0.04, p = 0.96) and presence (z =
0.96, p = 0.33) had no signicant eect. e abundance of subordinates also showed
signicant dierences among the three study sites (z ≥ 9.92, p < 0.0001).
e main factors which shape biodiversity and structure of animal communities are,
besides habitat heterogeneity (see e.g. MacArthur and MacArthur 1961, Hölldobler
Figure 4. Diversity of the ant communities within the study sites based on bait observations (medians,
quartiles, min–max values and outliers). Dierent letters indicate signicant dierences among groups.
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 109
and Wilson 1990, Tews et al. 2004, Ossola et al. 2015), intra- and interspecic rela-
tionships, such as competition and social parasitism. e importance of the latter is
well known in ants (Savolainen and Vepsäläinen 1988, Hölldobler and Wilson 1990,
Andersen 1992, Ryti and Case 1992, Cerdá et al. 1997, Czechowski and Markó 2006,
Scharf et al. 2011, Ślipiński et al. 2011, 2014). Colonies of dominant species, espe-
cially territorial ones, play a crucial role, as organizing centres of the ant community, by
limiting the establishment of colonies of subordinate species within dominant territo-
ries or by managing their accessibility to various resources (Savolainen and Vepsäläinen
1988, Pisarski and Vepsäläinen 1989, Andersen 1992, Bestelmeyer 2000, Lester et al.
2010, Cerdá et al. 2012, 2013, Czechowski et al. 2013). In our study, we revealed con-
siderable dierences among ant communities living in areas within the same habitat,
but dominated by two dominant species with clearly dierent features.
e species composition of the studied ant communities was quite similar among
all three sites, but dierences were found in the abundance of dierent ant species,
and also in the diversity of ant communities. In almost all respects the Formica san-
guinea site was superior housing the most diverse ant community. us, despite the
low chance of interaction with Lasius avus due to its subterranean lifestyle, the lowest
density of L. avus nests was recorded in the Formica exsecta territory, which could be
Figure 5. NMDS graph of the ant communities within the study sites based on bait observations (stress
= 0.114).
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
the result of the establishment of F. exsecta colonies on the top of their soil mounds,
with the gradual extermination of L. avus colonies (Pisarski 1982). is species was
followed by Myrmica spp. with a low level of aggression compared to L. platythorax
and Tetramorium cf. caespitum, while the lowest abundance was recorded for members
of the subgenus Serviformica, being both a slave species to F. sanguinea and the host of
temporary social parasitism caused by F. exsecta fundatrices (Czechowski et al. 2012).
Generally speaking, there was an increase in the abundance of aggressive and submis-
sive species in the mixed territory where F. exsecta and the slave maker species showed
lower abundance or were absent.
Usually, the exploitation patterns of food resources at the Formica exsecta site was
in agreement with the general rules concerning hierarchy patterns, where the dominant
species, especially a territorial one, monopolizes the area and its resources (Savolainen
and Vepsäläinen 1988, Savolainen et al. 1989). In the F. exsecta site the absence of T. cf.
caespitum at baits, known as an aggressive species (e.g. Brian et al. 1966, 1979) with a
quick reaction towards intruders (Dobrzański and Dobrzańska 1975) and with the abil-
ity to exploit habitat resources eectively (Brian et al. 1966), might have been caused by
its general avoidance of the dominant species during foraging (see Cerdá et al. 2012). On
the other hand, contrary to Formica exsecta, as other authors have already shown before
(e.g. Czechowski 1999, 2000, Czechowski and Markó 2006, Ślipiński et al. 2011), F.
sanguinea behaved as a considerably weaker competitor, exploiting the baits much less
actively than e.g. Lasius platythorax and Tetramorium cf. caespitum, which succeeded in
exploiting the baits within its area quite heavily. e strongest dierence in the exploita-
tion pattern of baits is shown by the results obtained from the mixed territory, where the
presence of almost all species at baits increased, with the exception of Formica exsecta,
which exhibited the highest frequency and abundance in its own exclusive territory.
e low number ofFormica sanguineaforagers at baits, especially in relation to
the numbers of workers of other ant species, requires additional explanations. Recent
studies on the foraging strategy ofF. sanguineahave already shown, that this species
seems to be less competitive towards subordinates to the south (e.g. Romania) than in
the northern parts of the species range in Europe (e.g. Finland) (P. Ślipiński et al., in
prep). One of the major signs of its weaker competitiveness is its reduced presence at
articialbaits, as also conrmed by our present observations. However, it is possible
that at least some of the F. cunicularia workers present at baits were individuals en-
slaved byF. sanguinea,and these de facto acted as foragers of the latter. Since,generally,
slaves fulll intranest tasks in colonies (Kharkiv 1979a,b, see also Czechowski 1996),
the abundance of slaves foraging at baits could anyhow be quite low.
Territorial ants can hold o raiding columns of Formica sanguinea to some extent,
and in this way directly protect possible slave species against their enslaver. is pro-
tective eect would manifest itself when there was a signicantly greater abundance
of slave species within, rather than outside of wood ant territories of the F. rufa group
(Punttila et al. 1996, Czechowski 2000, Czechowski and Vepsäläinen 2001, Czechows-
ki and Markó 2006, Väänänen et al. 2010). However, this is not generally valid for all
territorials, since, e.g., no measurable protective eect of the territorial Lasius fuliginosus
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 111
towards the F. fusca slave species was detected by Ślipiński et al (2014), despite cases of
direct interference of the latter during F. sanguinea raids (Czechowski 1999, 2000). In
our study area, two potential slave ant species (and, at the same time, potential hosts of
the temporary social parasitism of F. exsecta) occurred, both members of the subgenus
Serviformica: F. cunicularia and F. rubarbis. e former is a typical submissive species,
the latter is an aggressive encounter species. Due to this, the relations of each of them
with both local dominant species, i.e. F. exsecta and F. sanguinea, might proceed dier-
ently. On one hand, the submissive, weakly competitive F. cunicularia should be less
restrictively treated by the territorial F. exsecta than the more competitive F. rubarbis.
On the other hand, colonies of the aggressive F. rubarbis (see Mori et al. 2001) should
be much less prone to being destroyed by F. sanguinea, and they should also be more re-
sistant to being taken over by young F. exsecta queens than colonies of the submissive F.
cunicularia. So, one could expect that these dierences in behaviour might be reected
in dierences in the distribution of the two subordinate species. However, both species
were very scarce in the study area. In total, only three nests of F. cunicularia and one
nest of F. rubarbis were found. At the baits, only F. cunicularia foragers occurred, most
abundantly within the F. exsecta site. At the remaining sites (both with F. sanguinea),
they were generally scarce, and what is more, at least some of them may have been F.
sanguinea slaves. is result may suggest some type of protective eect of the territorial
F. exsecta on this slave species against the slave-maker.
While the decisive eect of Formica exsecta as a typically territorial species on the
structure of ant communities is quite evident, this question still remains open regard-
ing F. sanguinea. In the literature, the latter is commonly handled in a similar manner
as the territorial Formica s. str. and Coptoformica species (Vepsäläinen and Pisarski
1982, Savolainen et al. 1989, Punttila et al. 1996, Czechowski 2000, Czechowski and
Markó 2006, Väänänen et al. 2010). However, our ndings seem to contradict the
similarly strong territorial (sensu Pisarski 1982) nature of F. sanguinea. e fact that
some F. sanguinea nests can occur in the vicinity of F. exsecta nests (see the mixed site)
also emphasizes the reduced territoriality of F. exsecta. Other eld observations point
in this direction as well. For example, in the Białowieża Forest (N-E Poland), a very
strong F. sanguinea colony was observed to peacefully coexist with a fairly large F. ex-
secta colony over several years, nesting just ca. 3 m from an edge of the mound of the
latter (W. Czechowski, unpubl.). Also, the above mentioned recent case study on the
competitive strategy of F. sanguinea (P. Ślipiński et al., in prep.) strongly suggests the
non-territoriality of this aggressive and undoubtedly inuential species.
e results of this study underscore the importance of dierences between dominant
ant species in dierentially shaping ant communities, even within the same limited area.
We are indebted for the Apáthy István Society for the housing during the eld works.
e study was carried out with the support of scientic cooperation between the Polish
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
Academy of Sciences and the Romanian Academy. is work was supported by a grant
of the Romanian National Authority for Scientic Research and Innovation, CNCS –
UEFISCDI, project number PN-II-RU-TE-2014-4-1930.
Adler FR, Lebrun EG, Feener DH Jr (2007) Maintaining diversity in an ant community:
mode ling, extending and testing the dominance-discovery trade-o. American Naturalist
169: 323–333. doi: 10.1086/510759
Andersen AN (1992) Regulation of ‘momentary’ diversity by dominant species in exception-
ally rich ant communities of the Australian seasonal tropics. American Naturalist 140:
401–420. doi: 10.1086/285419
Arnan X, Cerdá X, Retana J (2012) Distinctive life traits and distribution along environmental
gradients of dominant and subordinate Mediterranean ant species. Oecologia 170: 489–500.
doi: 10.1007/s00442-012-2315-y
Bates D, Maechler M, Bolker B, Walker S (2014) lme4: Linear mixed-eects models using
Eigen and S4. R package version 1.1-7.
Bestelmeyer BT (2000) e trade-o between thermal tolerance and behavioural dominance in
a subtropical South American ant community. Journal of Animal Ecology 69: 998–1009.
doi: 10.1046/j.1365-2656.2000.00455.x
Brian MV (1979) Habitat dierences in sexual production by two co-existent ants. Journal of
Animal Ecology 48: 943–953. doi: 10.2307/4206
Brian MV, Hibble J, Kelly AF (1966) e dispersion of ant species in a southern English heath.
Journal of Animal Ecology 35: 281–290. doi: 10.2307/2395
Cerdá X, Angulo E, Caut S, Courchamp F (2012) Ant community structure on a small Pacic
island: only one native species living with the invaders. Biological Invasions 14: 323–39.
doi: 10.1007/s10530-011-0065-0
Cerdá X, Arnan X, Retana J (2013) Is competition a signicant hallmark of ant (Hymenoptera:
Formicidae) ecology? Myrmecological News 18: 131–147.
Cerdá X, Retana J, Cros S (1997) ermal disruption of transitive hierarchies in Mediterranean
ant communities. Journal of Animal Ecology 66: 363–374. doi: 10.2307/5982
Cerdá X, Retana J, Manzaneda A (1998) e role of competition by dominants and tempera-
ture in the foraging of subordinate species in Mediterranean ant communities. Oecologia
117: 404–412. doi: 10.1007/s004420050674
Cherix D, Bourne JD (1980) A eld study on a super-colony of the red wood ant Formica lugu-
bris Zett. in relation to other predatory arthropodes (spiders, harvestmen and ants). Revue
Suisse de Zoologie 87: 955–973. doi: 10.5962/bhl.part.85566
Csata E, Markó B, Erős K, Gál Cs, Szász-Len A-M, Czekes Zs (2012) Outstations as stable
meeting points for workers from dierent nests in a polydomous nest system of Formica
exsecta Nyl. (Hymenoptera: Formicidae). Polish Journal of Ecology 60: 177–186.
Czechowski W (1975) Bionomics of Formica (Coptoformica) pressilabris Nyl. (Hymenoptera,
Formicidae). Annales Zoologici 33: 103–125.
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 113
Czechowski W (1977) Recruitment signals and raids in slave-maker ants. Annales Zoologici
34: 1–26.
Czechowski W (1979) Competition between Lasius niger (L.) and Myrmica rugulosa Nyl. (Hy-
menoptera, Formicidae). Annales Zoologici 34: 437–451.
Czechowski W (1985) Competition between Myrmica laevinodis Nyl. and Lasius niger (L.)
(Hymenoptera, Formicoidea). Annales Zoologici 39: 153–173.
Czechowski W (1996) Colonies of hybrids and mixed colonies; interspecic nest takeover in
wood ants (Hymenoptera, Formicidae). Memorabilia Zoologica 50: 1–116.
Czechowski W (1999) Lasius fuliginosus (Latr.) on a sandy dune – its living conditions and
interference during raids of Formica sanguinea Latr. (Hymenoptera, Formicidae). Annales
Zoologici 49: 117–123.
Czechowski W (2000) Interference of territorial ant species in the course of raids of Formica
sanguinea Latr. (Hymenoptera, Formicidae). Annales Zoologici 50: 35–38.
Czechowski W (2006) e route of Formica polyctena Först. as a factor promoting emancipa-
tion of Formica fusca L. slaves from colonies of Polyergus rufescens (Latr.) (Hymenoptera:
Formicidae). Polish Journal of Ecology 54: 159–162.
Czechowski W, Markó B (2005) Competition between Formica cinerea Mayr (Hymenoptera:
Formicidae) and co-occurring ant species, with special reference to Formica rufa L.: direct
and indirect interferences. Polish Journal of Ecology 53: 467–487.
Czechowski W, Markó B (2006) Uncomfortable protection: Formica polyctena Först. shelters
Formica fusca L. from Formica sanguinea Latr. (Hymenoptera: Formicidae). Annales Zoo-
logici 56: 539–548.
Czechowski W, Markó B, Radchenko A, Ślipiński P (2013) Long-term partitioning of space
between two territorial species of ants (Hymenoptera: Formicidae) and their eect on subor-
dinate species. European Journal of Entomology 110: 327–337. doi: 10.14411/eje.2013.046
Czechowski W, Pisarski B (1988) Inter- and intraspecic competitive relations in Camponotus
ligniperdus (Latr.) (Hymenoptera, Formicidae). Annales Zoologici 41: 355–381.
Czechowski W, Radchenko A, Czechowska W, Vepsäläinen K (2012) e ants of Poland with
reference to the myrmecofauna of Europe. Fauna Poloniae (New Series) Vol. 4. Natura
Optima Dux Foundation, Warszawa, 1–496.
Czechowski W, Vepsäläinen K (2001) Formica rufa protects indirectly F. fusca against raids of
F. sanguinea. Annales Zoologici 51: 267–273.
Dejean A, Ryder S, Bolton B, Compin A, Leponce M, Azémar F, Céreghino R, Orivel J, Cor-
bara B (2015) How territoriality and host-tree taxa determine the structure of ant mosaics.
e Science of Nature 102: 33. doi: 10.1007/s00114-015-1282-7
Dobrzański J, Dobrzańska J (1975) Ethological studies in the ant Tetramorium caespitum. II.
Interspecic relationships. Acta Neurobiologiae Experimentalis 35: 311–317.
Erős K, Markó B, Gál Cs, Czekes Zs, Csata E (2009) Sharing versus monopolizing: distribution
of aphid sources among nests within a Formica exsecta Nyl. (Hymenoptera: Formicidae)
supercolony. Israel Journal of Entomology 39: 105–127.
Farris ZJ, Gerber BD, Karpanty S, Murphy A, Andrianjakarivelo V, Ratelolahy F, Kelly MJ
(2015) When carnivores roam: temporal patterns and overlap among Madagascar’s native
and exotic carnivores. Journal of Zoology 296: 45–57. doi: 10.1111/jzo.12216
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
Helanterä H, Strassmann JE, Carrillo J, Queller DC (2009) Unicolonial ants: where do they
come from, what are they and where they are going? Trends in Ecology & Evolution 24:
341–349. doi: 10.1016/j.tree.2009.01.013
Higashi S, Yamauchi K (1979) Inuence of a supercolonial ant Formica (Formica) yessensis
Forel on the Ishikari coast. Japanese Journal of Ecology 29: 257–264.
Hothorn T, Bretz F, Westfall P, Heiberger RM, Schützenmeister A, Scheibe S (2015) Package
‘multcomp’. Simultaneous inference in general parametric models. R-package version 1.4-
Hölldobler B, Wilson EO (1990) e Ants. Harvard University Press, Cambridge, MA, 732pp.
Hughes J (2006) A review of wood ants (Hymenoptera: Formicidae) in Scotland. Scottish
Natural Heritage Commissioned Report No. 178 (ROAME No. F04AC319).
Kharkiv VA (1997a) Eectiveness of activity and division of labor between slaves and slave-
makers in colonies of Formica sanguinea (Hymenoptera, Formicidae). Zoologicheskij
Zhurnal 76: 438–443.
Kharkiv VA (1997b) Competition as a mechanism of division of labour in colonies of ant
slavemakers Formica sanguinea (Hymenoptera, Formicidae). Zoologicheskij Zhurnal 76:
Lester PJ, Stringer LD, Haywood J (2010) e role of resource dispersion in promoting
the co-occurrence of dominant and subordinate ant species. Oikos 119: 659–668. doi:
Maák S, Markó B, Erős K, Babik H, Ślipiński P, Czechowski W (2014) Cues or meaningless
objects? Dierential responses of the ant Formica cinerea to corpses of competitors and
enslavers. Animal Behaviour 91: 53–59. doi: 10.1016/j.anbehav.2014.02.014
MacArthur RH (1972) Geographical Ecology: Patterns in the Distribution of Species. Harper
and Row, New York, 269 pp.
MacArthur RH, MacArthur JW (1961) On bird species diversity. Ecology 42: 594–598. doi:
Markó B, Czechowski W (2012) Space use, foraging success and competitive relationships in
Formica cinerea (Hymenoptera, Formicidae) on sand dunes in southern Finland. Ethology,
Ecology & Evolution 24: 149–164. doi: 10.1080/03949370.2011.634438
Markó B, Czechowski W, Radchenko A (2013) Combining competition with predation: dras-
tic eect of Lasius fuliginosus (Latr.) on subordinate ant species at the northern limit of its
distribution. Annales Zoologici 63: 107–111. doi: 10.3161/000345413X666156
Markó B, Czekes Zs, Erős K, Csata E, Szász-Len A-M (2012) e largest polydomous system
of Formica ants (Hymenoptera: Formicidae) in Europe discovered thus far in Romania.
North-Western Journal of Zoology 8: 287–291.
McFarlane Tranquilla L, Montevecchi WA, Hedd A, Regular PM, Robertson GJ, Field DA,
Devillers R (2015) Ecological segregation among ick-billed Murres (Uria lomvia) and
Common Murres (Uria aalge) in the Northwest Atlantic persists through the nonbreeding
season. Canadian Journal of Zoology 93: 447–460. doi: 10.1139/cjz-2014-0315
Mori A, Grasso DA, Visicchio R, Le Moli F (2001) Comparison of reproductive strategies and
raiding behaviour in facultative and obligatory slave-making ants: the case of Formica san-
guinea and Polyergus rufescens. Insectes Sociaux 48: 302–314. doi: 10.1007/PL00001782
Dierential impact of two dominant Formica ant species (Hymenoptera, Formicidae)... 115
Oksanen J, Guillaume Blanchet F, Kindt R, Legendre P, Minchin PR, O’Hara RB, Simpson
GL, Solymos P, Stevens HH, Wagner H (2013) vegan: CommunityeEcology package. R
package version 2.0-10.
Ossola M, Nash A, Christie FJ, Hahs AK, Livesley SJ (2015) Urban habitat complexity aects
species richness but not environmental ltering of morphologically-diverse ants. PeerJ 3:
e1356. doi: 10.7717/peerj.1356
Parr CL, Gibb H (2010) Competition and the role of dominant ants. In: Lach L, Parr CR,
Abbott KL (Eds) Ant ecology. Oxford University Press, Oxford, 369–373.
Petráková L, Schlaghamerský J (2011) Interactions between Liometopum microcephalum (For-
micidae) and other dominant ant species of sympatric occurrence. Community Ecology
12: 9–17. doi: 10.1556/ComEc.12.2011.1.2
Pisarski B (1980) Evolution of the competitive behaviour in social insects. Insectes Sociaux 27:
Pisarski B (Ed.) (1982) Structure et organisation des societés de fourmis de l’espèce Formica
(Coptoformica) exsecta Nyl. (Hymenoptera, Formicidae). Memorabilia Zoologica 38: 1–281.
Pisarski B, Vepsäläinen K (1989) Competition hierarchies in ant communities (Hymenoptera,
Formicidae). Annales Zoologici 42: 321–329.
Pohlert T (2015) PMCMR: Calculate pairwise multiple comparisons of mean ranksSums. R
package version 1.1.
Punttila P, Haila Y, Tukia H (1996) Ant communities in taiga clearcuts: habitat eects and
species interactions. Ecography 19: 16–28. doi: 10.1111/j.1600-0587.1996.tb00151.x
Punttila P, Haila Y, Pajunen T, Tukia H (1991) Colonisation of clearcut forests by ants in the
southern Finnish taiga: a quantitative survey. Oikos 61: 250–262. doi: 10.2307/3545343
Reznikova Zh I (1980) Interspecic hierarchy in ants. Zoologicheskij Zhurnal 59: 1168–1176.
[In Russian]
Reznikova ZhI (1983) Interspecic Relations in Ants. Nauka, Novosibirsk, 206 pp. [In Russian]
Reznikova Zh (1999) Ethological mechanisms of population density control in coadaptive
complexes of ants. Russian Journal of Ecology 30: 187–192.
Ryti RT, Case TJ (1992) e role of neighborhood competition in the spacing and diversity of
ant communities. American Naturalist 139: 355–374. doi: 10.1086/285331
Robinsons EJH (2014) Polydomy: the organisation and adaptive function of complex nest sys-
tems in ants. Current Opinion in Insect Science 5: 37–43. doi: 10.1016/j.cois.2014.09.002
Rosengren R (1986) Competition and coexistence in an insular ant community – a manipula-
tion experiment (Hymenoptera: Formicidae). Annales Zoologici Fennici 23: 297–302.
Rosengren R, Cherix D, Pamilo P (1985) Insular ecology of the red wood ant Formica trunco-
rum Fabr. I. Polydomous nesting, population size and foraging. Mitteilungen der Schweiz-
erischen Entomologischen Gesellschaft 58: 147–175.
Rzeszowski K, Babik H, Czechowski W, Markó B (2013) Ants (Hymenoptera: Formicidae) of
Chełmowa Góra in the Świętokrzyski National Park. Fragmenta Faunistica 56: 1–15. doi:
Sanders NJ, Gordon DM (2003) Resource-dependent interactions and the organization of desert
ant communities. Ecology 84: 1024–1031. doi: 10.1890/0012-9658(2003)084[1024:RI-
Gema Trigos Peral et al. / Journal of Hymenoptera Research 50: 97–116 (2016)
Savolainen R (1990) Colony success of the submissive ant Formica fusca within territories of
the dominant Formica polyctena. Ecological Entomology 15: 79–85. doi: 10.1111/j.1365-
Savolainen R (1991) Interference by wood ant inuences size selection and retrieval rate of prey
by Formica fusca. Behavioral Ecology and Sociobiology 28: 1–7. doi: 10.1007/BF00172132
Savolainen R, Vepsäläinen K (1988) A competition hierarchy among boreal ants: impact on re-
source partitioning and community structure. Oikos 51: 135–155. doi: 10.2307/3565636
Savolainen R, Vepsäläinen K (1989) Niche dierentiation of ant species within territories of
the wood ant Formica polyctena. Oikos 56: 3–16. doi: 10.2307/3566082
Savolainen R, Vepsäläinen K, Wuorenrinne H (1989) Ant assemblages in the taiga biome: testing
the role of territorial wood ants. Oecologia 81: 481–486. doi: 10.1007/BF00378955
Scharf I, Fischer-Blass B, Foitzik S (2011) Spatial structure and nest demography reveal the in-
uence of competition, parasitism and habitat quality on slavemaking ants and their hosts.
BMC Ecology 11: 9. doi: 10.1186/1472-6785-11-9
Seifert B (2000) A taxonomic revision of the ant subgenus Coptoformica Mueller, 1923 (Hyme-
noptera: Formicidae). Zoosystema 22: 517–568.
Seifert B (2007) Die Ameisen Mittel- und Nordeuropas. Lutra–Verlags- und Vertriebsgessel-
schaft, Görlitz/Tauer, 368 pp.
Spotti FA, Castracani C, Grasso DA, Mori A (2015) Daily activity patterns and food prefer-
ences in an alpine ant community. Ethology, Ecology & Evolution 27: 306–324. doi:
Sunarto S, Kelly MJ, Parakkasi K, Hutajulu MB (2015) Cat coexistence in central Sumatra:
ecological characteristics, spatial and temporal overlap, and implications for management.
Journal of Zoology 96: 104–115. doi: 10.1111/jzo.12218
Ślipiński P, Markó B, Czechowski W, Włodarczyk T (2011) Space use in Formica sanguinea
my territory my kingdom? Entomologica Romanica 16: 63.
Ślipiński P, Markó B, Rzeszowski K, Babik H, Czechowski W (2014) Lasius fuliginosus (Hy-
menoptera: Formicidae) shapes local ant assemblages. North-Western Journal of Zoology
10: 404–412.
Tews J, Brose U, Grimm V, Tielborger K, Wichmann MC, Schwager M, Jeltsch F (2004) Ani-
mal species diversity driven by habitat heterogeneity/diversity: the importance of keystone
structures. Journal of Biogeography 31: 79–92. doi: 10.1046/j.0305-0270.2003.00994.x
Väänänen S, Vepsäläinen K, Savolainen R (2010) Indirect eects in boreal ant assemblages:
territorial wood ants protect potential slaves against enslaving ants. Annales Zoologici 60:
57–67. doi: 10.3161/000345410X499524
Vepsäläinen K, Pisarski B (1982) Assembly of island ant communities. Annales Zoologici Fennici
19: 327–335.
Vepsäläinen K, Savolainen R (1990) e eect of interference by Formicine ants on the foraging
of Myrmica. Journal of Animal Ecology 59: 643–654. doi: 10.2307/4886
Wiens JA (1983) Avian community ecology: an iconoclastic view. In: Brush AH, Clark GA Jr
(Eds) Perspectives in ornithology. Cambridge University Press, Cambridge, 355–403. doi:
... Territoriality is the most effective strategy to ensure the control of resources in ants (Hölldobler and Wilson 1990;Adams 2016). Territorial ant species usually dominate their foraging grounds, wherein they exert competitive pressure on the cohabiting ant species Vepsäläinen 1988, 1989;Pisarski and Vepsäläinen 1989;Petráková and Schlaghamerský 2011;Czechowski et al. 2013;Adams 2016;Trigos-Peral et al. 2016;Ślipiński et al. 2018). ...
... Such polydomous species efficiently exploit the available food supplies (Debout et al. 2007;Erős et al. 2009;Csata et al. 2012), and because of high worker density, they have a strong negative effect on other ant species (e.g. Czechowski et al. 2013, Trigos-Peral et al. 2016. Under normal circumstances, a lower-ranked species can cohabit with a species of a higher level, but when resources diminish, the competition between them increases to an extent that could end coexistence (Savolainen and Vepsäläinen 1988;Markó and Czechowski 2004;Cerdá et al. 2013;Czechowski et al. 2013;Markó et al. 2013;Ślipiński et al. 2014). ...
... The meadow is fairly intensely grazed by cows for most of the year. The structure of the F. exsecta polydomous systems as well as intraspecific and mutualistic relationships of F. exsecta to other species are well studied (Goropashnaya et al. 2007;Erős et al. 2009;Martin et al. 2009;Csata et al. 2012;Trigos-Peral et al. 2016). ...
Full-text available
Territorial strategy in animals is characterized by the monopoly of resources inside a protected area, the territory. The presence of territorial species considerably alters the behavior of co-occurring submissives, as it is known in several submissive ant species living on the territories of red wood ants in temperate regions. On the other hand, as a rule, territorial species cannot share the same territory and usually exclude each other. However, this ‘rule’ is inferred from the almost complete lack of data on the coexistence of rival territorials, and not from observations regarding the effective exclusion or behavioral inhibition of one territorial species by other. In the frame of this study, we investigated the foraging strategy of the territorial red wood ant Formica pratensis that occurred inside a large polydomous system of another territorial ant species, F. exsecta. Formica pratensis colonies outside the supercolony served as control. Within the F. exsecta supercolony, F. pratensis showed the characteristics of a subordinate species characterized by low discovery and exploitation success of artificial food sources. On the contrary, control colonies outside the polydomous system clearly behaved like typical territorials as they successfully monopolized the majority of the baits. In addition, submissive species were more successful around the F. pratensis nests within the supercolony, than outside of it. As suggested by our results, territorial species could co-occur with other territorials given certain plasticity in their behavior and small colony size. Nevertheless, exclusion also happens as proven by our field observations.
... Although ant hierarchy levels were first described in temperate forest species (Vepsäläinen & Pisarski, 1982), these topics have been studied more extensively in ant communities in tropical forest or arid habitats. Although data is relatively scarce, it has been found that ant communities in temperate forest are mainly structured around dominant and subordinate species (Arnan, Cerdá & Retana, 2012;Trigos Peral et al., 2016). Arnan et al. (2018), found that species richness was positively correlated with the presence of dominant native ant species, a pattern that was consistent in temperate and subtropical latitudes, because ant richness is lower in temperate forest than other habitats, dominance is expected to be lower in these communities. ...
... and Temnothorax sp. are cold-climate specialists, which are better adapted for the cooler conditions of the oak forest (Andersen, 1997;Andersen, 2000;Cuautle, Vergara & Badano, 2016). In the case of the Formica genus, different studies point to it being a possible bioindicator in temperate ecosystems (Ellison, 2012) and species of this genus are identified as key species around which temperate communities are structured (Trigos Peral et al., 2016). On the other hand, generalized Myrmicinae, such as M. minimum, are successful competitors that predominate under moderate levels of stress and disturbance but are not as active and aggressive as dominant Dolichoderinae, which are aggressive species most abundant in environments with low levels of stress and disturbance (Andersen, 1997). ...
... Fewer dominant species promotes low competition, which is then reflected in a lower number of registered species and low dominance indices. However, the inclusion of the different species recorded in this study in one of the two categories and subcategories indicated by Trigos Peral et al. (2016), or one of the three categories proposed by Cerdá, Arnan & Retana, (2013), is not so simple. The absence of a dominant element is evident, which also agrees with what is expected for temperate ecosystems. ...
Full-text available
Background The discovery-dominance trade-off is the inverse relationship between the ability of a species to discover resources and the species’ dominance of those resources; a paradigm used to explain species coexistence in ant communities dependent on similar resources. However, factors such as stress (e.g., temperature) or disturbance (e.g., removal of biomass) associated with the change in land use, can modify this trade-off. Here, we aimed to determine the potential effects of land use change on dominance hierarchy, food preferences and on the discovery-dominance trade-off. Methods An experiment with baits was used to investigate the dominance hierarchies of ant communities in a temperate mountain habitat in central Mexico. We evaluated the dominance index (DI), food preferences and discovery-dominance trade-offs of ants inhabiting two types of vegetation: a native oak forest and agricultural land resulting from agricultural land use and grazing. Results The ant communities in both environments were comprised of three species of ants ( Monomorium minimum, Myrmica mexicana , and Camponotus picipes pilosulus ), four morphospecies ( Pheidole sp.1 and Pheidole sp.2, Temnothorax sp. and Lasius sp.) and one genus ( Formica spp.). All Formicidae showed values of intermediate to low DI, and this factor did not seem to be influenced by the change in land use. Ants in the modified vegetation (i.e., agricultural land) were found to be numerically greater. Overall, a higher number of visits were registered to the tuna bait, although the duration of foraging events to the honey baits was longer. However, foraging times were dependent on the species considered: the generalized Myrmicinae, M. minimum , the ant species with highest DI, foraged for longer periods of time in the agricultural land and on the tuna bait. Meanwhile, the cold-climate specialist Formica spp., with a lower DI, foraged for longer periods of time in the oak (although not significant) and on the honey bait. We found little evidence of the discovery-dominance trade-off; instead, we found considerable diversity in the strategies used by the different species to access resources. This range of strategies is well represented by the generalized Myrmicinae M. minimum , the cold-climate specialists Formica spp. and Temnothorax sp., and the rare species, as the cold climate specialist Lasius sp. (insinuators). Conclusions Our evaluation shows that transformation of the original habitat does not appear to affect the hierarchical dominance of the ant communities, but it does affect their food preferences. Species with higher DI values such as the generalized Myrmicinae are more skilled at resource acquisition in modified habitats. Our results suggest that change in land use promotes an increase in the diversity of foraging strategies used by different ant species. This diversity may contribute to resource partitioning which favors coexistence.
... Ants are easy targets for studying the role of competition, since they can be found in high abundance in a wide variety of terrestrial habitats. Competition among ants is studied on many ecological levels, starting from the direct interactions between individuals and colonies (e.g., CzeChowski 1999, 2004, MaRkó & CzeChowski 2004, CzeChowski & MaRkó 2005, CzeChowski & al. 2009, Maák & al. 2014, vepsäläinen & CzeChowski 2014, Ślipiński & Żmihorski 2017 to the role of competition in the formation of multispecies ant assemblages (e.g., savolainen & vepsäläinen 1989, väänänen & al. 2010, tRiGos peRal & al. 2016, Żmihorski & Ślipiński 2016; for a review, see AdAms 2016). ...
... Moreover, in some cases, F. sanguinea does not even exploit all the baits close to its nest. In typical territorial species (e.g., Formica polyctena, F. exsecta, Lasius fuliginosus), the negative impact on the foraging activity of subordinate species, not to mention the use of baits as such, is incomparably more spectacular than that shown here for F. sanguinea (see e.g., savolainen & vepsäläinen 1988, vepsäläinen & savolainen 1990, CzeChowski & MaRkó 2006, tRiGos peRal & al. 2016. ...
Full-text available
There are almost no hard data on the competitive status of Formica sanguinea, and previous studies focus mainly on its spectacular lifestyle as a slave-maker. It is usually described as an aggressive and territorial species, the latter due to its raiding behaviour and morphological resemblance to species of the subgenus Formica s. str. (typical territorial species and top dominants). A series of field observations using baits was carried out across the longitudinal extent of F. sanguinea’s range in Europe, from Finland through Poland to Romania. The main objective of the present study was to determine the competitive abilities of F. sanguinea and its status in the interspecific ant hierarchy. We also investigated the changes in the competitive abilities of F. sanguinea along the meridional axis of the species’ range. Based on our results, the distribution of F. sanguinea foragers around the nest showed a distance dependent pattern, with fewer individuals present in distant arenas and at distant baits. While in the absence of baits, other ant species were seemingly not influenced by the abundance of F. sanguinea, this changed in the presence of baits, showing that F. sanguinea has a negative effect on other species. Still, many baits were exploited by other ant species around the studied nests. There were also clear differences among the study regions in terms of bait utilisation, with Finnish colonies exploiting most of baits, while Romanian colonies mostly neglecting them. In light of the present study and due to the complexity of the competitive strategy of the species, we believe that F. sanguinea does not fall into the conventionally arranged three level hierarchy of interspecific ant competition. In addition, F. sanguinea’s competitive ability may depend on local ecological conditions as revealed by the comparison of different populations across Europe.
... Baiting enable a researcher to record the interactions amongst ant species/colonies and provide the mechanisms the different species employ to access resources such as food [96,97]. Peral et al. [98] used it to investigate if the dominating ant species could be restrictively influencing the traits of the subordinate ones in habitat where they discovered that the presence of dominant ant species limited the abundance and occurrence of subordinates. It can also be used to investigate the interactions amongst individuals of the same ant species from different colonies. ...
Full-text available
Habitat transformation is one of the main drivers of the ecosystem degradation on earth that is ameliorated by restoring some of the degraded ecosystems by regaining their natural ecological functions with all their biotic and abiotic components. The biotic and abiotic components of the ecosystem under restoration can be used to assess the response of the ecosystem to the restoration. Ideal variable to use as the indicator should be able respond positively to the diminishing elements that we causing the degradation and interact positively to some of the biotic and abiotic components expected to prevail when the ecosystem is fully restored. One of such variable is ants. We here provide the information about the eligibility of using ants as indicators of terrestrial ecosystems undergoing restoration and sampling and basic analytical methods to apply when implanting ants at assessing ecosystem undergoing restoration.
... This can highlight the importance of competition for food and nesting sites which has an enhanced effect on seminatural meadows (see also Lenda et al. 2013;Grześ et al. 2018). Competition is usually considered to have a significant effect in shaping ant communities (Savolainen and Vepsäläinen 1988;Braschler and Baur 2003;Trigos-Peral et al. 2016), and unavoidably occurs when the ecological requirements of species overlap (Pianka 1974;Glen and Dickman 2008). The negative effects of competitive interactions can be reduced if the morphological, behavioral, and ecological plasticity of the competing partners allows shifts in their requirements, thus minimizing niche overlap, as observed also in ants (see Cerdá et al. 2013 for a review). ...
Full-text available
Several factors can influence individual and group behavioral variation that can have important fitness consequences. In this study, we tested how two habitat types (seminatural meadows and meadows invaded by Solidago plants) and factors like colony and worker size and nest density influence behavioral (activity, meanderness, exploration, aggression, and nest displacement) variation on different levels of the social organization of Myrmica rubra ants and how these might affect the colony productivity. We assumed that the factors within the two habitat types exert different selective pressures on individual and colony behavioral variation that affects colony productivity. Our results showed individual-/colony-specific expression of both mean and residual behavioral variation of the studied behavioral traits. Although habitat type did not have any direct effect, habitat-dependent factors, like colony size and nest density influenced the individual mean and residual variation of several traits. We also found personality at the individual-level and at the colony level. Exploration positively influenced the total-and worker production in both habitats. Worker aggression influenced all the productivity parameters in seminatural meadows, whereas activity had a positive effect on the worker and total production in invaded meadows. Our results suggest that habitat type, through its environmental characteristics, can affect different behavioral traits both at the individual and colony level and that those with the strongest effect on colony productivity primarily shape the personality of individuals. Our results highlight the need for complex environmental manipulations to fully understand the effects shaping behavior and reproduction in colony-living species.
Full-text available
Faunistička istraživanja mrava u Mediteranu, posebice u poljoprivrednim sustavima, kod nas su vrlo rijetka. Budući da je mediteransko područje bogato vrstama, a način gospodarenja u poljoprivredi može dodatno utjecati na raznolikost vrsta, istražena je fauna mrava u maslinicima i vinogradima na području Zadarske županije. Uspoređeni su indeksi raznolikosti između područja s različitim načinima upravljanja; u maslinicima i vinogradima s integriranom i ekološkom zaštitom bilja u kojima se koriste različite skupine pesticida, te prirodnom staništu bez gospodarenja, gdje nije bilo uporabe pesticida. Mravi su uzorkovani tijekom vegetacijske sezone u 2018. godini koristeći metodu stresanja, Tullgrenove lijevke i lovne zamke. Vrste su određene morfološki i metodom barkodiranja DNA te je utvrđeno 14 vrsta mrava. Pronađene vrste Aphaenogaster picena i Aphaenogaster balcanica su endemske, te su rijetki nalazi za Hrvatsku. Sve sekvence mravljih vrsta koje su određene u ovom istraživanju bile su prisutne u bazi podataka Nacionalnog centra za biotehnološke informacije (NCBI). Metode lova su utjecale na sastav i brojnost vrsta. Prema prikupljenim podacima, maslinici s integriranom zaštitom bilja u usporedbi s ekološkim maslinicima, imali su manju raznolikost. Stoga bi se utjecaj različitih načina gospodarenja u maslinicima na bioraznolikost mrava mogao detaljnije istražiti proučavanjem zajednica mrava na većem broju ploha.
The hierarchical organization of multi-species ant communities can be determined by the colony size and worker density-dependent differences in behaviour, recruitment efficiency, and aggressiveness between colonies of different species. In this study, we compared the competitive effects of two territorial wood ant species, Formica rufa and F. polyctena, on the ant species subordinated to them (encounterers and submissives) that nest close-by their mounds. The nests of subordinates were mapped in 10 m2 plots around the mounds of each of the two wood ant species, situated at a distance of 10, 20 and 30 m in four directions. The most abundant subordinate species were two Myrmica species: M. rubra and M. ruginodis. Interestingly, the nests of subordinates had an even diversity, density and distribution along with the distance from the wood ant mounds. The wood ant presence had a species-specific negative effect on some of the subordinate species, which can be in relation with the smaller mound sizes in F. rufa compared to F. polyctena. Based on our results it seems that subordinate species, due to their adaptability, might tolerate the circumstances close-by the wood ant mounds by changes in their strategies, foraging behaviour, and switching to the use of alternative food sources, like corpses of wood ants present in high abundance. Although these might not allow reaching normal colony size and reproduction, they may allow the colonies to survive while maintaining relatively large nest densities.
Full-text available
Background Ant-plant mutualistic networks tend to have a nested structure that contributes to their stability, but the ecological factors that give rise to this structure are not fully understood. Here, we evaluate whether ant abundance and dominance hierarchy determine the structure of the ant-plant networks in two types of vegetation: oak and grassland, in two temperate environments of Mexico: Flor del Bosque State Park (FBSP) and La Malinche National Park (MNP). We predicted that dominant and abundant ant species make up the core, and submissives, the periphery of the network. We also expected a higher specialization level in the ant trophic level than in plant trophic level due to competition among the ant species for the plant-derived resources. Methods The ant-plant interaction network was obtained from the frequency of ant-plant interactions. We calculated a dominance hierarchy index for the ants using sampling with baits and evaluated their abundance using pitfall traps. Results In MNP, the Formica spp. species complex formed the core of the network (in both the oak forest and the grassland), while in FBSP, the core species were Prenolepis imparis (oak forest) and Camponotus rubrithorax (grassland). Although these core species were dominant in their respective sites, they were not necessarily the most dominant ant species. Three of the four networks (oak forest and grassland in FBSP, and oak forest in MNP) were nested and had a higher number of plant species than ant species. Although greater specialization was observed in the ant trophic level in the two sites and vegetations, possibly due to competition with the more dominant ant species, this was not statistically significant. In three of these networks (grassland and oak forest of MNP and oak forest of FBSP), we found no correlation between the dominance hierarchy and abundance of the ant species and their position within the network. However, a positive correlation was found between the nestedness contribution value and ant dominance hierarchy in the grassland of the site FBSP, which could be due to the richer ant-plant network and higher dominance index of this community. Conclusions Our evidence suggests that ant abundance and dominance hierarchy have little influence on network structure in temperate ecosystems, probably due to the species-poor ant-plant network and a dominance hierarchy formed only by the presence of dominant and submissive species with no intermediate dominant species between them (absence of gradient in hierarchy) in these ecosystems.
Full-text available
Territoriality of three dominant ant species: Formica rufa L., Lasius fuliginosus (Latreille), and Lasius emarginatus (Olivier), was studied in July and August 2013–2015 in the broadleaf forest areas in Kiev, Ukraine. Thirteen species of ants were found on the soil surface, including 4 subdominants and 5 influents. Subordinate species co-occurred in the plots with L. emarginatus 1.5–2.0 times as frequently as with L. fuliginosus, and at least 3.0 times as frequently as with F. rufa. From 2 (within the foraging territory of the nest complex) to 5 ant species (in the territories of single colonies) were found together with F. rufa. Eight ant species co-occurred with L. fuliginosus and L. emarginatus each, and 7 species, i.e., nearly the same number, were found in the areas without dominants. The dominant ant species can be arranged in ascending order of territoriality as follows: L. emarginatus < L. fuliginosus + F. rufa (single colonies) < F. rufa (nest complex).
Full-text available
First published in 1983 to celebrate the centennial of the American Ornithologists' Union, Perspectives in Ornithology collects together a series of essays and commentaries by leading authorities about especially active areas of research on the biology of birds. Readers will find in this collection a useful overview of many major concepts and controversies in ornithology.
The data on the zonal and biotopical distribution of 70 ant species in the southern Western Siberian Plain and adjacent regions were analyzed. The results showed that, against the background of a considerable diversity of the ant fauna, multispecies communities dominated by Formica pratensis retain the constant species composition and the quantitative ratio of nests over the area from the northern forest steppe to the southern boundary of the steppe zone, and the basic features of their species composition remain unchanged up to the desert zone. In such coadaptive complexes, stable interspecies relationships and characteristic mechanisms of density regulation develop. Field experiments on artificially changing the size of ant families in communities with different dominants demonstrated that ants of dominant species actively regulate the level of dynamic density of subdominant species and exterminate "excess" individuals, estimating their number fairly accurately. Such a form of interspecies relationships is named interspecies social control.
Competition occurs when different species or individuals require the same limiting resources. It can occur within colonies for reproductive rights, between colonies of the same species (intraspecific competition) and between populations of different species (interspecific competition). Evidence for intraspecific competition includes overdispersion of nests, territoriality, and reallocation of castes in response to new neighbours. Evidence for interspecific competition includes spatial ant mosaics, agonistic behaviour, chemical defence and behavioural dominance hierarchies. Experiments in interspecific competition show that it is highly conditional varying with resource quality and quantity, biotic and abiotic conditions. The discovery-;dominance trade-off suggests a possible mechanism for species coexistence. The dominance-impoverishment relationship suggests that species richness is reduced where the abundance of dominant ants is high.
Formica fusca L. slaves from colonies of Polyergus rufescens (Latr.) tend to establish small homospecific satellite nests close to a main nest. P. rufescens maintains the integrity of a mixed colony by means of peculiar integration raids (Czechowski 2005). The present paper describes the history of a group of such satellite nests, temporarily separated from the main nest by a foraging route of Formica polyctena Först, impassable to P. rufescens. The ex-slaves had managed to adopt F. fusca gynes and raised some homospecific worker pupae before the F. polyctena route disappeared and P. rufescens started to raid their nests. During the raids, the mutual attitude of the slave-makers and the ex-slaves quickly evolved from hostility to friendly relations.
The paper gives a description of a Lasius fuliginosus Latr. colony on a sandy dune, the ants' way of life in this beyond-optimum habitat, and their attitude towards other ant species (Formica sanguinea Latr., F. cinerea Mayr and F. fusca L.) revealed during raids of blood-red ants in the neighbourhood. The results are discussed in the context of hierarchical organization of ants species assemblages.
In colonies of ant slavemakers Formica sanguinea Latr. a division of labour between species takes place. Slaves are mainly involved in performing intranest finctions, while the majority of foragers is represented by slavemakers. The expediency of dividing the functions between species was studied. Examining slaves and slavemakers performing different functions has shown that the labour division was not directly related to the productivity of their labour. Slavemakers demonstrated the higher labour efficiency in nest building and brood transport compared with that in slaves (F. cunicularia, F. fusca). Activity of slaves and slavemakers in brood nursing and foraging didn't differ. The data on energy expenditures, available in literature, evidence that the character of dividing the finctions between different species in Formica sanguinea colonies is not quite optimal probably due to using the workers of other species in colonies of ant-slavemakers.
The paper presents four cases of territorial ant colonies interfering in the course of Formica sanguinea Latr. raids against slave species which nest next to or in their territories. In three cases, Formica rufa L., when protecting its own territory, forced out tone case) or held (two cases) a raiding column of ET sanguinea, thus "saving" local Formica fusca L. nests from danger. In one case, Lasius fuliginosus (Latr.) did not let a F. sanguinea column pass its territory; to reach its target (a colony of Formica cinerea Mayr), the column had to by-pass the foreign territory.