Acta Zoologica Academiae Scientiarum Hungaricae 50 (2), pp. 77–95, 2004
MOSAIC STRUCTURE OF ANT COMMUNITIES
(HYMENOPTERA: FORMICIDAE) IN EASTERN CARPATHIAN
MARSHES: REGIONAL VERSUS LOCAL SCALES
MARKÓ, B.
1,2
, KISS, K.
2
and GALLÉ, L.
2,3
1
Department of Taxonomy and Ecology, Babes-Bolyai University
3400 Cluj-Napoca, str. Clinicilor 5–7, Romania, E-mail: mbalint@biolog.ubbcluj.ro
2
Department of Ecology, University of Szeged, 6701 Szeged, P.O.Box 51, Hungary
E-mail: galle@bio.u-szeged.hu
3
Tisza Research Group, Hungarian Academy of Sciences, Szeged, Hungary
The ant communities of 12 habitats in four marshlands were studied in the Eastern Carpathians.
Altogether 11 ant species were identified, but none of the species could be considered a spe-
cific relic species of marshlands, though characteristic species could be detected, like Myrmi-
ca rubra, M. ruginodis and M. scabrinodis. The sampling procedure seems to be fairly com-
plete. The between and within region comparisons revealed that there were no regional differences
in the ant communities, but differences could be attributed mostly to differences in habitat at-
tributes like vegetation and related characteristics. The three most abundant Myrmica species
all showed clear habitat preferences, which overlapped just to a lesser extent. The diversity
analysis suggested a highly mosaic-like distribution pattern of ants in different habitats, which
could be attributed to specific constraint factors acting on the ant species inhabiting mountain
marshlands. Thus usually one patch is occupied and dominated by a single species. The exis-
tence of this mosaic-like pattern is also supported by the possibility of competition between M.
rubra and M. ruginodis. The importance of connecting different scales to elucidate mecha-
nisms creating the observed patterns is emphasized.
Key words: Formicidae, communities, mountain marshes, constraint factors, patchiness, com-
petition, scales
INTRODUCTION
The general recognition of the significance of biodiversity emphasizes the
importance of animal community studies revealing the mechanisms, which create
the observed patterns. Considering ecological processes significant in nature con-
servation like extinction, recolonization, recovery, etc. it becomes essential to con-
nect different scales, and to carry out within and between habitat comparisons (SA-
VOLAINEN et al. 1989, PUNTTILA et al. 1994, NIEMELÄ et al. 1996, PUNTTILA et al.
1996, GALLÉ 1999, GALLÉ et al. 2000, BESTELMEYER &WIENS 2001, GALLÉ
2002), in order to elucidate to what extent specific communities are exposed to
threats. The connection of different scales, e.g. regional ⇒ local ⇒ patch, is of
main importance, as it can help elucidate how different processes on different
scales interact and produce a pattern observed on regional level (SAVOLAINEN &
Acta zool. hung. 50, 2004
Hungarian Natural History Museum, Budapest

VEPSÄLÄINEN 1988, BESTELMEYER &WIENS 2001, GALLÉ 2002). These studies
can at least reveal how stable the qualitative-quantitative features of a specific type
of animal community are in similar habitats but in different regions. The accurate
analysis of the characteristics of the same type of animal community in similar
habitats also offers the possibility of assessing the constraints acting on it
(MABELIS 1977, PUNTTILA et al. 1994, NIEMELÄ et al. 1996, PUNTTILA et al.
1996, GALLÉ 1999, GALLÉ et al. 2000, BESTELMEYER &WIENS 2001). The
knowledge of the natural variations of such features, as well as of the constraint
factors acting on the community can help conservationists to elaborate appropriate
protection plans for specific types of communities. This is especially important in
the cases of such extremely vulnerable and unstable habitats as mountain marshes
in Central and Eastern Europe, which are constantly exposed to disturbances either
from the part of forestry or from tourism.
In the Eastern Carpathians there are several marshes, which are known as
glacial refuges for different valuable plant species (POP 1960, KOVÁCS 1984, RUP-
RECHT &SZABÓ 1999, MARGÓCZI et al. 2000). These habitats bear specific condi-
tions, e.g.: high water level and low annual mean temperature potentially acting as
strong constraint factors on the potential set of animal species inhabiting these ar-
eas. The high water level can induce higher mortality, or it can inhibit the larval de-
velopment in the case of insects. The combination of these factors can also act indi-
rectly: e.g. the scarcity of living organisms means narrower prey spectra for preda-
tors. In the case of ants one of the primary constraints is the availability of suitable
nesting sites (MABELIS 1977, BOOMSMA & DE VRIES 1980, WARDLAW &ELMES
1996), as colonies need inundation-free conditions for the normal development of
the larvae and pupae. Another specific limiting factor for ants is the narrow spectra
of organisms serving as prey for them, which can also have a serious impact on
their survivor, or on the size of the colony (MABELIS 1977, ELMES 1991,
WARDLAW &ELMES 1996). Whereas a solitary animal can solely assure the conti-
nuity of its species by finding food sources, and then reproducing, in ants the col-
ony acts as a single reproducing unit, requesting many times more food to maintain
the colony population. Under these conditions ant-communities of marshlands are
expected to bear specific characteristics (GALLÉ et al. 2000). Obviously these
communities can be unique in several ways, e.g. (1) containing specific marshland
species (VEPSÄLÄINEN et al. 2000), (2) bearing specific qualitative and quantita-
tive characteristics (GALLÉ et al. 2000, VEPSÄLÄINEN et al. 2000), (3) having spe-
cifically strong inter- and intraspecific interactions due to the scarcity of resources
(SAVOLAINEN &VEPSÄLÄINEN 1988), (4) developing suitable colony founding
and dispersal strategies e.g. colony fission, and (5) adapting colony systems to
long-time persistence, like polygyny.
78 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004

Although the descriptive comparison of communities cannot elucidate the
underlying structuring processes (VEPSÄLÄINEN &PISARSKI 1982, PISARSKI &
VEPSÄLÄINEN 1989, PUNTTILA et al. 1994, 1996), as the same community struc-
ture can be reached on several routes (VEPSÄLÄINEN &PISARSKI 1982, GALLÉ
1991), but some suggestions can be formulated on the basis of their analysis. Gen-
erally it is agreed that one of the basic structuring processes is competition in ants
(MABELIS 1977, VEPSÄLÄINEN &PISARSKI 1982, SAVOLAINEN &VEPSÄLÄINEN
1988, PISARSKI &VEPSÄLÄINEN 1989, SAVOLAINEN et al. 1989, GALLÉ 1991,
JÁRDÁN et al. 1993, PUNTTILA et al. 1994, GALLÉ 1999), although other spe-
cies-specific factors as dispersal ability, habitat requirements, dependence on other
species (MABELIS 1977, BOOMSMA & DE VRIES 1980, VEPSÄLÄINEN &PISARSKI
1982, PUNTTILA et al. 1994, 1996, JÁRDÁN et al. 1993, BESTELMEYER &WIENS
2001), or even stochasticity (VEPSÄLÄINEN &PISARSKI 1982, PUNTTILA et al.
1996) can also stand at the basis of the presence of a specific ant species in a given
community.
This study proposes to offer answers to some basic questions regarding ant
communities of some Eastern Carpathian marshland areas:
(1) Can the studied marshes be considered as refuges for habitat specialist ant
species?
(2) Are there inter- or intraregional differences among the ant-communities
of different sites?
(3) Does the community structure show the constraints acting on ant-com-
munities e.g. scarcity of nesting sites?
(4) Is competition important in structuring marshland ant communities and to
what extent?
Ants are also potentially ideal bio-monitoring organisms and have proven to
be valuable indicators of environmental conditions and of biodiversity (GALLÉ
1991, JÁRDÁN et al. 1993, GALLÉ 1999, ALONSO 2000, ALONSO &AGOSTI 2000,
KASPARI &MAJER 2000) showing how vulnerable a habitat is. The study of ant
communities should help evaluate the condition of marshlands in the Eastern
Carpathians.
MATERIALS AND METHODS
The following criteria were taken in account when selecting the study sites: to have (1) both
large and relatively small areas, (2) marshes from different regions, (3) and different types of habitats
within one marshland, like sedge meadows, marsh-forests, peat bogs etc. Upon these criteria the ant
communities of four marshlands were analyzed in the Eastern Carpathians, Romania: the marshland
MOSAIC STRUCTURE OF ANT COMMUNITIES IN EASTERN CARPATHIAN MARSHES 79
Acta zool. hung. 50, 2004

at Voslobeni in the Giurgeului Basin, Lacul Dracului marsh in the Harghita Mts, Fagul Rotund, and
Apa Lenta marshes in the Nemira Mts.
(1) The fen area at Voslobeni (46°40’N, 25°37’E) is the remnant of a once existing large
marsh-complex lying in the Giurgeului Basin near the river Mures at cca. 770 m above sea level.
Nowadays this scattered marshland contains wet hayfields, peat bogs, bushes, and large fens, such as
both eutrophic and oligo-mesotrophic habitats are present. This very mosaic area is one of the richest
regions in relic plant species (POP 1960, MARGÓCZI et al. 2000). Three sampling sites were selected:
(a) the core of the marsh (V1) is a peat bog of approximately 60 m diameter with sparse pines (Carici
echinatae-Sphagnetum SOÓ (1934) 1954), where the peat moss (Sphagnum sp.) cover is 70–80%, (b)
V2 is a sedge meadow (Caricetum rostratae RÜBEL, 1912) with sparse trees and shrubs (Betula
pubescens, Salix cinerea, S. pentandra), and the presence of Sphagnum species is quite common in
the moss layer, whereas (c) V3 is a dense marsh forest with willow and alder trees (Alnetum incanae
BROCKMANN 1907) (GALLÉ et al. 2000).
(2) The Lacul Dracului marsh-complex (46°29’N, 25°31’E) is a more isolated area in a small
depression in the Harghita Mts at 1180 m above sea level. It contains both eutrophic and oligotrophic
habitats. Altogether four sample sites were studied. The typical peat bog is represented by LD1
(Eriophoro vaginati-Sphagnetum recurvi HUECK 1925), a relatively open area with sparse Scots
pines and blueberry bushes. The core area of the marsh is an open sedge meadow (Caricetum elatae
KOCH 1926, LD2 and LD3), and the nearby area (LD4) is a dense and wet forested bog with small
spruces (Piceetum sphagnoso-Polytrichetosum SOÓ 1944) (SZABÓ A. and MÁTIS A. pers. comm.).
(3) Fagul Rotund (46°11’N, 26°21’E) is the smallest marsh of all. It lies in the Nemira Mts,
one of the easternmost parts of the Eastern Carpathians, at cca. 1100 m above sea level. The first sam-
ple site (FR1) was an open peat bog (Eriophoro vaginati-Sphagnetum POP et al. 1987), whereas the
second one (FR2) was a sparse Scots pine forest with a dense moss layer and blueberry bushes
(Vaccinio-Pinetum silvestris KLEIST 1929) (KATÓ Z. pers. comm.).
(4) The Apa Lenta marsh complex (46°11’N, 26°17’E) lies also in the Nemira Mts in a valley
next to Fagul Rotund (2.5–3.5 km away) at cca. 1000 m above sea level. It is a mosaic of eutrophic
and oligo-mesotrophic marsh patches. The first sample-site (AL1) was an open sedge meadow
(Caricetum rostratae RÜBEL 1912), the second area (AL2) was a peat bog with sparse birch and
spruce trees (Eriophoro vaginati–Sphagnetum recurvi HUECK 1925), whereas the third site (AL3)
was a dense willow and birch forest (Salici pentandrae-Betuletum pubescentis SOÓ (1934) 1955)
(RUPRECHT &SZABÓ 1999). Sphagnum species were present in the moss layer at every site.
Ants were collected with pitfall-traps using plastic jars of cca. 5.5 cm diameter. Fifteen traps
were set out at each sample site, which were arranged ina3×5grid, each trap being separated by 5 m
from the other traps (SAVOLAINEN et al. 1989, PUNTTILA et al. 1994, 1996, NIEMELÄ et al. 1996,
GALLÉ et al. 2000). Ethylene glycol was used as preserving liquid in the case of Voslobeni, and Lacul
Dracului, whereas in the case of Apa Lenta, and Fagul Rotund marshes brine (saturated NaCl-
solution) replaced the ethylene glycol. The samplings were carried out between 1998 and 2000, as
follows: Fagul Rotund 20–27 July 1998, Apa Lenta 21–28 July 1998, Voslobeni 21–31 July 1999,
Lacul Dracului between 6–13 August 2000.
The collected specimens were identified on the basis of the keys of SEIFERT (1988, 1996) and
RADCHENKO et al. (1997). Two richness estimates were used to see how effective the collections
were in every region. The first-order jackknife estimate is based on the observed species number
(S
obs
) and the number of species occurring in only one sample (L), where n is the total number of sam-
ples (LONGINO 2000):
S=S +L
n–1
n
*
obs






80 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004

The Chao2 richness estimate also incorporates the number of species occurring in two samples
(M) (LONGINO 2000):
S=S +
L
2M
*
obs
2
In both richness estimates S
*
is the assessed number of species.
Regional differences
Principal Coordinate Analysis (distance algorithm: Wishart’s similarity ratio) was used to an-
alyze the structure of the dataset. Two steps were separated. In the first step the origin of the data was
used as grouping criteria. In the second step the type of habitat was considered as possible grouping
factor. It is generally known that differences between ant communities are not following the differ-
ences in the vegetation, but more likely the architectural properties of the vegetation determine the
differences (JÁRDÁN et al. 1993, GALLÉ 1999, GALLÉ et al. 2000). On this basis it was more appropri-
ate to use the following three main habitat types: (1) dense marsh-forests with alder, willow, birch
and spruce, (2) typical peat-bog areas with sparse Scots pines and birches, and (3) sedge meadows.
During the analysis each trap was considered as a separate unit. The computations were carried out
with the SynTax computer software (PODANI 1993). The habitat preferences of the most frequent spe-
cies were analyzed by comparing the number of individuals recorded in traps in different habitat
types using Kruskall-Wallis ANOVA and Mann-Whitney U-test (StatSoft 1995).
Local heterogeneity
In an ideal situation (assumed spatial homogeneity of the habitat) one would expect that ani-
mal species should be distributed evenly in space (LONGINO 2000). Consequently a sample character-
izing this ideal situation should be made up of similar sample units as each species is expected to
occur at the same species-specific probability everywhere (LONGINO 2000). On the basis of this logic
it seems obvious that the more mosaic the species’ distribution is in a given area, the bigger the differ-
ences are between the qualitative-quantitative properties of the sample units and of the sample. The
Shannon-Wiener entropy is an appropriate method for characterizing communities as it implies both
the richness and the evenness of the studied entity, and also emphasizes the presence of rare data
types, e.g. rare species. As such it is a useful tool to assess the structure of a community. According to
the above-presented logic every pitfall trap material was considered to be a small ’representative’
sample unit of the studied site’s community. PUNTTILA et al. (1996) suggested that individual pit-
fall-trap catches (separated by 5 m) should reflect the distribution of the colonies for species with
short foraging ranges as Myrmica, Leptothorax, and foraging preferences of the species with wider
foraging areas like Lasius or Camponotus.VEPSÄLÄINEN and SAVOLAINEN (1990) also suggested that
Myrmica workers’ foraging radius is equal or less than 3 m. Thus by comparing the overall diversity
and evenness values with the values for single traps we should be able to characterize fairly well the
spatial structure of the ant communities.
Interspecific relationships
The analysis of species distribution and abundance data also makes us possible to assume
whether competition could be important in structuring these communities (MABELIS 1977, VEPSÄ-
LÄINEN &PISARSKI 1982, SAVOLAINEN et al. 1989, GALLÉ 1991, PUNTTILA et al. 1996).
MOSAIC STRUCTURE OF ANT COMMUNITIES IN EASTERN CARPATHIAN MARSHES 81
Acta zool. hung. 50, 2004

Mann-Whitney U-test and Spearman rank-correlation coefficient (StatSoft 1995) was utilized
to clarify the existence of interspecific relationships in the case of the most abundant species.
RESULTS
Altogether 11 ant species were identified (Table 1), each of them are widely
distributed in temperate and northern Europe with the exception of Myrmica
vandeli, which has a sporadic distribution (SEIFERT 1988), and it was first identi-
fied in Romania at Voslobeni not long ago (MARKÓ 1999). An additional parasitic
species, Harpagoxenus sublaevis (NYLANDER, 1849), was collected at LD2 in the
nest of Leptothorax acervorum (MARKÓ &CSŐSZ 2001), which species wasn’t
present in the traps.
The identified species are generally characteristic for mountain heaths,
marshes, and forest clearcuts. However, no specific marshland species were de-
tected, like Formica uralensis, F. picea (VEPSÄLÄINEN et al. 2000) or Myrmica
sulcinodis (SEIFERT 1988). Generally the species number was low, it fluctuated
between 3 and 7 (Table 1). Parallel to this the majority of the species showed rather
uneven distribution. Besides M. ruginodis and M. rubra, only M. scabrinodis and
Formica lemani showed higher constancy, and they were also more abundant than
the other species. It is relevant that the most widespread species were also numer-
ous, whereas the other species were characterized not just by sporadic distribution
but also by low abundance (Table 1). The low capture efficiency of the latter spe-
cies (convertible in foraging intensity) suggests that their colonies were small,
probably underdeveloped. The combination of accidental occurrence and low
abundance supports the hypothesis of their reduced preference for mountain
marshes.
The first-order jackknife and the Chao2 richness estimates showed that ap-
proximately one species could be added to the known number of species (Table 2).
In contradiction no additional species are expected to occur in marshlands when
considering every region together. Thus probably one or (at most) two rare species
are expected to occur in every separate region additionally, but the survey of
marshland ant species can be considered fairly complete all things considered.
Ant-communities: region vs. habitat type
The result of the principal coordinate analysis did not support any clustering
tendency of the ant-communities on regional level (Fig. 1). Nevertheless if we ap-
plied the main vegetation type of the given sample site as grouping criteria, there
was a more or less obvious differentiation (Fig. 1). Thus the three main vegetation
82 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004

MOSAIC STRUCTURE OF ANT COMMUNITIES IN EASTERN CARPATHIAN MARSHES 83
Acta zool. hung. 50, 2004
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types, as (1) peat-bogs with sparse Scots pines and birches (61% of the peat bog
samples included in the group), (2) sedge meadows (66% of the meadow samples
included in the group), and (3) marsh-forests (94% of the forest samples included
in the group) formed loosely defined clusters. The existence of these groups
proved, nevertheless, only tendencies, for none of the three clusters was separated
clearly by the others. Besides, the low value of the cumulative percentage of
eigenvalues of the first two axes (35.33% and 46% for the first three axes respec-
tively) also revealed the heterogeneity of the dataset.
The three most common ant species – Myrmica rubra, M. ruginodis and M.
scabrinodis – were separately analyzed for their habitat preferences. These three
84 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004
Fig. 1. PCoA–plot of the ant–communities, each dot representing a single trap: filled dots – open
peat–bog habitats with sparse trees; empty dots – sedge meadows; crosses – marshland forests; A –
marshland forests; B – peat–bog habitats; C – sedge meadows
Table 2. Different types of richness estimates of the number of ant species occurring in the four
studied marshlands
Voslobeni Lacul Dracului Fagul Rotund Apa Lenta Altogether
No. of species 6 7 7 7 11
1st order jackknife 6.97 7.97 7.96 8.94 11
Chao2 6.5 7.25 7.5 –* 11
*M = 0, invalid formula

species have almost the biggest niche-overlap among other Myrmica species (ELMES
1991). Two of these species, namely M. rubra and M. ruginodis were present in
each of the three major habitat types, whereas M. scabrinodis occurred only in peat
bogs and sedge meadows. The presence of M. scabrinodis did not show any biases
regarding the two habitat types (Table 3). On the contrary M. rubra and M. rugi-
nodis showed differences in their preferences: they both preferred peat bogs and
marshland forests to sedge meadows (Table 3). There were no significant differ-
ences, however, in their preferences regarding marsh forests or peat bogs (Mann-
Whitney U-test, z = –0.16, p = n.s. for M. rubra, and z = –1.7, p = n.s. for M. rugi-
nodis), though it seemed that M. rubra was generally more abundant in marsh for-
ests, whereas M. ruginodis in peat bogs (Table 3). These results along with the
PCoA-analysis showed clearly, that while in peat bogs all the three Myrmica spe-
cies could be found in great abundance, sedge meadows were generally character-
ized by the presence of M. scabrinodis, and marsh-forests by the presence of M.
rubra and to a somewhat lesser extent by the occurrence of M. ruginodis.VEPSÄ-
LÄINEN and SAVOLAINEN (1990) also found that M. scabrinodis and M. ruginodis
preferred different types of habitat (open terrains vs. forests), and they did not oc-
cur together usually.
Local heterogeneity
The data from pitfall-traps could also be used to make a fair assessment on
the number of species present in the traps’ range of catch. The mean species num-
ber per traps was very low which showed that generally patches were inspected by
foragers of one, or in a few cases, of two species (Table 4). Thus the mean diversity
values of patches were strikingly lower than the overall site diversity values in al-
most every case (Table 4). On the basis of the comparison of the patches’ average
evenness values with the evenness of the overall site diversity values (Fig. 2, Table 4)
MOSAIC STRUCTURE OF ANT COMMUNITIES IN EASTERN CARPATHIAN MARSHES 85
Acta zool. hung. 50, 2004
Table 3. The presence of the three most frequent ant species in different types of marshland habitats
Species Mean no. of specimens (SD) Comparison
peat–bog sedge meadow marsh forest
Myrmica scabrinodis 1.6 (0.84)
n = 10
4.56 (1.04)
n = 23
– Mann–Whitney U–test,
z = –1.72
Myrmica ruginodis 4.02 (5.68)
n = 43
1.2 (0.41)
n = 15
1.57 (0.95)
n = 7
Kruskall–Wallis ANOVA
(df = 2) chi
2
= 9.16**
Myrmica rubra 7.7 (7.05)
n = 30
5 (9.13)
n = 18
10.3 (14.55)
n = 26
Kruskall–Wallis ANOVA
(df = 2) chi
2
= 6.1*
*p < 0.05, **p < 0.01

it could be concluded that the ant fauna of the patches were clearly dominated by a
few (generally one) species, while the ant community of the given site was more
even in qualitative-quantitative aspects. It becomes obvious, that the structure of
ant communities was very mosaic-like: foragers of mainly one species dominated
the patches.
Interspecific relationships
As it was stated in the previous sections, the three most abundant Myrmica
species showed separate habitat preferences. However, only the morphologically
and behaviourally very similar M. rubra and M. ruginodis were present in every
habitat type, thus it seemed obvious to investigate the probability of interspecific
exclusion on finer scale only in the case of this species pair.
Only those sites were taken in consideration where both species occurred, ex-
cluding site LD1, where only Myrmica rubra was present, because there was no
evidence whether M. ruginodis was missing due to exclusion by M. rubra,oritwas
absent because of other factors. We also did not consider data from traps where
both species were absent, for the above-presented reasons. The Spearman rank-cor-
relation coefficient (r = –0.47, t = –5.36, p < 0.000, n = 103) revealed a significant
negative relationship between the two species that could imply the existence of
86 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004
Fig. 2. The overall evenness vs. the evenness of the mean patch–diversity values for each sample site
(V – Voslobeni, LD – Lacul Dracului, FR – Fagul Rotund, AL – Apa Lenta)

88 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004
Fig. 3. Number of Myrmica rubra specimens in the absence (1), and in the presence (2) of M. rugi-
nodis (Mann–Whitney U–test, z = –5.15, p < 0.000, n
1
= 47, n
2
= 56)
Fig. 4.Number of Myrmica ruginodis specimens in the absence (1), and in the presence (2) of M. rub-
ra (Mann–Whitney U–test, z = –5.14, p < 0.000, n
1
= 29, n
2
= 74)

DISCUSSION
From regional to local level
None of the species can be considered to be specific relic species of the Car-
pathian marshes, though the two most abundant Myrmica species, M. rubra and M.
ruginodis, and even M. scabrinodis can be regarded as core species of marshland ant
communities. VEPSÄLÄINEN et al. (2000) report M. scabrinodis to be a typical peat
bog species in Finland, whereas in Romania this species is generally quite frequent
in mountain open areas (GALLÉ et al. 2000). It is interesting that GALLÉ et al.
(2000) did not find Myrmica rubra and M. ruginodis to be common in the
hayfield-marshland complex at Voslobeni. Nevertheless the high constancy of
these two species in the studied marshes indicates that these species can be handled
as characteristic species of Eastern Carpathian mountain marshes. The low fre-
quency and low abundance of other species emphasizes the restrictive character of
these habitats for ants. Although territorial Formica s. str., Raptiformica and
Coptoformica species occur in the surroundings of marshes (GALLÉ et al. 2000,
GALLÉ et al. unpubl.), none of these species was detected in the studied areas. Ac-
cording to VEPSÄLÄINEN et al. (2000) virgin and early succession ditched bogs are
unsuitable habitats for wood ants due to the microclimate, but the lack of large
aphid colonies can also lead to the absence of these obligate aphid-tending species.
Nevertheless the low abundance of their host Serviformica species (or their lack in
the case of Voslobeni) can solely diminish the possibility of colonization. The dis-
tance to the nearest inhabited patches, or the degree of isolation from these areas
can also account for the colonization probability in the case of Formica s. str. spe-
cies (MABELIS 1994, MABELIS &KORCZYŃSKA 2001). The lack of these territo-
rial species causes changes in the competitive hierarchy: species considered to be
submissive (BRIAN et al. 1966 in SAVOLAINEN &VEPSÄLÄINEN 1988, VEPSÄ-
LÄINEN &PISARSKI 1982, PISARSKI &VEPSÄLÄINEN 1989, PUNTTILA et al. 1996)
become top species of these communities, overruling in abundance the lower
ranked, but aggressive species, like Lasius niger,orCamponotus herculeanus,
which occur sporadically and in low abundance.
The lack of regional differences clearly supports the idea of strong con-
straints acting on the set of ant species, which could inhabit these areas, and as the
species pool is small and constant there is no possibility to develop specific com-
munities. There is though some evidence that the habitat type can slightly differen-
tiate among ant communities (MABELIS 1977, JÁRDÁN et al. 1993, NIEMELÄ et al.
1996, GALLÉ et al. 2000, VEPSÄLÄINEN et al. 2000) on local level. GALLÉ et al.
(2000) emphasizes the importance of the vegetation’s architecture in structuring
MOSAIC STRUCTURE OF ANT COMMUNITIES IN EASTERN CARPATHIAN MARSHES 89
Acta zool. hung. 50, 2004

the ant communities of the hayfield-marshland complex around Voslobeni. Alike
the vegetation (MARGÓCZI et al. 2000), the water content of the soil can be one of
the major differentiating factors in ants (MABELIS 1977, JÁRDÁN et al. 1993,
NIEMELÄ et al. 1996). The segregation of habitat preferences learned in the case of
the three most abundant Myrmica species also support the possible effect of vege-
tation and factors related to it.
The key factor of being successful in marsh habitats generally characterized
by shortage in nesting sites can be the efficiency of the applied dispersal strategy.
The success of Myrmica species over Formicines can be attributed to the smaller
size of their colonies, which allows them to inhabit smaller suitable patches, than
Formicines (BRIAN et al. 1976 in VEPSÄLÄINEN &SAVOLAINEN 1990). It is clear
that ant species, which use colony fission can more successfully colonize the avail-
able nesting sites, and parallel to this the presence of other species, which disperse
exclusively by queens, can become more accidental. BANSBACH and HERBERS
(1999) report that Myrmica punctiventris can expand its colony’s territory via poly-
domy, thus it seems appropriate to suggest, that these Myrmica species, which are
also well-known for the existence of polygynous colonies (ELMES 1991, WARD-
LAW &ELMES 1996, WALIN &SEPPÄ 2001), use colony fission as dispersal strat-
egy (BRIAN &BRIAN 1955 in PUNTTILA et al. 1994, ELMES 1991, WARDLAW &
ELMES 1996, WALIN &SEPPÄ 2001). It is also known in M. punctiventris that col-
onies frequently move (BANSBACH &HERBERS 1999), which was also observed in
M. ruginodis in forests (MARKÓ and KISS pers. obs.) and in M. gallieni (PETAL
pers. comm.). This suggests that if a nesting place becomes unsuitable the whole
colony could move onto another site (ELMES 1991) even in marshes.
The resistance to occasional risings of water level can also lead to the domi-
nance of both Myrmica species, and even to the noticeable presence of the M. scab-
rinodis. As stated by BOOMSMA and ISAAKS (1982) M. rubra and M. scabrinodis
tolerate inundations in sandy coastal plains surviving for several weeks. The ants
usually prefer well-aerated Festuca mounds (BOOMSMA &ISAAKS 1982), which
elevate 1–2 m from the sea level. MABELIS (1977) also reports that usually M.
rubra and M. ruginodis are the two ant species present in forests with high water
level. In accordance Myrmica colonies are generally found in Sphagnum mounds in
marshes, and M. rubra was found to survive the inundations in Voslobeni and in
other regions, too (GALLÉ unpubl.). Contrary to M. rubra, Lasius flavus is consid-
erably vulnerable to rising water (BOOMSMA &ISAAKS 1982), which, combined
with its endogeic life style of tending root aphids (not too frequent in marshes),
could also explain why this species, otherwise abundant on mountain heaths, has a
low occurrence in marshes.
90 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004

The low annual mean temperature of the marshlands can also interplay in the
success of some species. VEPSÄLÄINEN and SAVOLAINEN (1990) suggest that
Myrmica scabrinodis and M. ruginodis are predominantly low-temperature spe-
cies. Other factors, as differences in larval development among species can cause
biases in community structure additionally. It is known that in M. ruginodis the lar-
val development is the fastest among all other European Myrmica species (ELMES
&WARDLAW 1983), which makes possible the development of larger colonies in
shorter period, than needed by the other species. This advantage over the other spe-
cies is even more emphasized by the relatively short warm period in mountain areas.
From local to patch level
The low availability of nesting sites combined with other restricting factors
like narrow prey spectra causes a mosaic-like structure of the ant communities:
suitable patches are monopolized by mostly one species, and these patches are
rarely visited by other species. This is clearly supported by the low evenness values
of the patches. As a result the foraging territories of different species are well sepa-
rated, the probability of overlapping is reduced. VEPSÄLÄINEN et al. (2000) also
reports patchy distribution of three Myrmica species on ditched bogs in Finland,
which leads us to the presumption that this is a general feature of the marshland ant
communities.
On the level of patches this mosaic-like structure can be supported by inter-
specific competition, which is generally considered to be of main importance in
structuring ant communities (MABELIS 1977, VEPSÄLÄINEN &PISARSKI 1982,
SAVOLAINEN &VEPSÄLÄINEN 1988, PISARSKI &VEPSÄLÄINEN 1989, SAVOLAI-
NEN et al. 1989, GALLÉ 1991, BANSBACH &HERBERS 1999, VEPSÄLÄINEN et al.
2000). The different types of shortages specific for marshes can enhance even
more the probability of inter- and intraspecific conflicts. The two most abundant
species of marshes, namely Myrmica rubra and M. ruginodis are not just morpho-
logically very close to each other, but their ecological requirements also overlap
(MABELIS 1977, SEIFERT 1988, RADCHENKO et al. 1997, VEPSÄLÄINEN et al.
2000, CZECHOWSKI et al. 2002). Both species live in forests at low altitudes; M.
rubra generally prefers oak-forests in plains, lowlands, while M. ruginodis can be
found in forests of the hilly regions (SEIFERT 1988, RADCHENKO et al. 1997,
ALVARADO &GALLÉ 2000, CZECHOWSKI et al. 2002). In mountains M. rubra
generally inhabits wet pastures, and marshes. Its sister species occurs on pastures,
marshes, and in spruce forests. Though they seem to replace each other, they can
also be found together. The colonial properties of both species are also similar
(SEIFERT 1988, ELMES 1991, WARDLAW &ELMES 1996), nevertheless M. rubra
MOSAIC STRUCTURE OF ANT COMMUNITIES IN EASTERN CARPATHIAN MARSHES 91
Acta zool. hung. 50, 2004

has usually larger colonies (ELMES 1991). ELMES (1991) suggests that displace-
ment between M. rubra and M. ruginodis is high. PUNTTILA et al. (1996) detected
negative relationship between coexisting M. ruginodis and M. sulcinodis in Finn-
ish taiga clearcuts, whereas MABELIS (1977) found the same relationship in the
case of M. rubra and M. ruginodis species pair. Thus the mere possibility of coex-
istence of the two abundant Myrmica species having similar ecological require-
ments raises questions on the existence and nature of interspecific relationship, and
suggests that they should form a locally improbable species pair (VEPSÄLÄINEN &
PISARSKI 1982, SAVOLAINEN &VEPSÄLÄINEN 1988). Although the study of com-
petition on the basis of distribution patterns has been criticized many times, be-
cause many processes or combination of processes may lead to an observed pattern
(VEPSÄLÄINEN &PISARSKI 1982, SAVOLAINEN &VEPSÄLÄINEN 1988) in the
case of these two Myrmica species we believe that the statistical analysis of the dis-
tribution data could indirectly detect the mere possibility of competition due to the
above-presented similarities of the species. The results show that interspecific
competition can indeed well occur between the two most abundant Myrmica spe-
cies, which is also valid for the relationship between these species and the other
species to a lesser extent. Potential coexisting competitors should exhibit resource
partitioning, which could be reflected especially by habitat, time and food type
partitioning (SAVOLAINEN &VEPSÄLÄINEN 1988). In this case the results support
only the existence of habitat partitioning, but other types of partitioning cannot be
excluded. However, these datasets refer only indirectly to the spatial pattern of the
distribution of the ants, and since direct evidences for negative interactions are not
available, these results should be handled carefully (SAVOLAINEN &VEPSÄLÄI-
NEN 1988, PUNTTILA et al. 1996). In addition there is no available data on the role
of intraspecific competition in this case, though it is known to exist and to be of rel-
evant importance in Myrmica species (ELMES 1991, MARKÓ unpubl.).
Connecting different scales
The absence of regional differences proves the uniformity of regional species
pools in great lines. Nevertheless there can be specific species, as Myrmica van-
deli, which appear only sporadically, but their presence in marshes is generally
negligible. The presence or absence, and the dominance of specific ant species
mostly depend on their dispersal abilities, as well as on their colony foundation
strategies. The differences among ant communities of major marsh habitat types
are probably caused by differences in the vegetation as well as other factors con-
nected to it, like shadiness and water level. The restrictive nature of these factors,
one of which is most probably the nesting site shortage, causes a mosaic structure
92 MARKÓ, B., KISS, K. & GALLÉ, L.
Acta zool. hung. 50, 2004

of ant communities, supported then by the competition on the level of patches.
Consequently it would be a mistake to consider that competition is the only factor
structuring these communities. As presented previously the combination of several
species-specific factors, as well as stochastic processes (VEPSÄLÄINEN &PISARSKI
1982, SAVOLAINEN &VEPSÄLÄINEN 1988, PUNTTILA et al. 1994, PUNTTILA et al.
1996, VEPSÄLÄINEN et al. 2000) and certainly competition, too, can shape the
ant-communities of marshes. These different processes, and factors act on differ-
ent scales, but all are important on their turn contributing to the community forma-
tion process.
*
Acknowledgements – We are grateful for the considerable help of ESZTER RUPRECHT and
ANNA SZABÓ in identifying the plant associations of the sample sites. We owe thanks for the valuable
comments of the anonymous referee, and for the articles kindly provided to B. M. by Dr. DONAT
AGOSTI and Dr. BRAM MABELIS. We are also indebted to all of those who assisted us in the fieldwork,
especially to DÁVID FÜLÖP,RÓBERT GALLÉ, Dr. KATALIN MARGÓCZI,KUNIGUNDA MACALIK,
ATTILA MÁTIS,ISTVÁN MIKÓ,EMŐKE NAGY, Dr. ENDRE SÁRKÁNY-KISS and TAMÁS SIKE.
The fieldwork at Apa Lenta, Fagul Rotund, and Lacul Dracului marshes made part of a re-
search programme of the Eco Studia Society (Cluj-Napoca, Romania) supported by the Illyés Fund
(Budapest, Hungary). The fieldwork at Voslobeni was carried out in the frame of a research project of
the Department of Ecology, University of Szeged (Szeged, Hungary), supported by grant no
1–0379/98 of the Hungarian Nature Conservation Office (Ministry of Environment), and coordinated
by L.G.
B. M.’s work was supported by a research grant of the A.N.S.T.I. Romanian Government
(grant no. 6203/B27/2000), and of the Hungarian Ministry of Education – Department for Hungari-
ans Abroad (spring 2002), as well as by the Arany János Fund for Science (Budapest, Hungary) (au-
tumn 2002) during the preparation of the MS. K.K. was also supported by a research grant from the
Hungarian Ministry of Education – Department for Hungarians Abroad (Budapest, Hungary) during
this period.
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Revised version received August 30, 2004, accepted September 6, 2004, published November 24, 2004
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