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Three to four dominant seed-transporting ant species of different size categories (Tetramorium caespitum, Lasius niger, Formica rufibarbis/Formica pratensis) on the plateau of abandoned ore sedimentation basin (tailings containment) were studied as pioneer and subsequent colonisers of this industrial waste deposit, from the viewpoint of their functioning in plant seed dispersal. We examined the role of ants in primary vegetation succession. Experiments of seed removal by ants with plant species found withinclose proximity of tailings were related to the succession. Ant activity generates a considerable shift in the the quality of the colonised surface, as they collectively act as ecosystem engineers.
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1Department of Botany & 2Institute of the Environment, Faculty of Science, Charles
University in Prague, Benátská 2, 128 01 Prague 2, Czech Republic
Received: 15th May 2013, Accepted: 17th July 2013
Three to four dominant seed-transporting ant species of different size categories
(Tetramorium caespitum, Lasius niger, Formica rufibarbis/Formica pratensis) on the
plateau of abandoned ore sedimentation basin (tailings containment) were studied as
pioneer and subsequent colonisers of this industrial waste deposit, from the viewpoint of
their functioning in plant seed dispersal. We examined the role of ants in primary
vegetation succession. Experiments of seed removal by ants with plant species found within
close proximity of tailings were related to the succession. Ant activity generates a
considerable shift in the quality of the colonised surface, as they collectively act as
ecosystem engineers.
Keywords: human-made habitats, industrial-waste deposits, abandoned tailings
containment, primary vegetation succession, plant species diversity, dispersal of plant
seeds, ant-plant interaction, myrmecochory, ecosystem engineers, bioturbation
Ants often play an important role in the dynamics of plant communities acting as seed
dispersal agents (e.g. Hölldobler et Wilson 1990, Jolivet 1996, Kovář et Kovářová 1998).
Foraging workers are likely to take seeds as food into their nests, carrying them a distance
away from the parent plant. Although ants disperse seeds over relatively short distances,
advantages commonly associated with this dispersal strategy include avoidance of parent
competition and of density-dependent predation, as well as dispersal towards microsites
favorable to germination (Wolf et Debussche 1999, Kovář et al. 2001). Activity of
successionally alternating ant species during decades imply significant change of the
substrate by bioturbation during nest-building (Dostál et al. 2005, Frouz et Jílková 2008)
and dynamics in vegetation development (Vlasáková et al. 2009, Vojtíšek 2012). To build
and maintain mounds, ants transport large amounts of substrate from deeper layers (King
1981, Frouz 1996), through a process known as bioturbation (e.g., Meysman et al. 2006).
As a result, ants influence concentrations of chemical elements, soil texture, amount of soil
organic matter, activity and biomass of microfauna, pH as well as the other soil properties
(Czerwinski et al. 1971, Culver et al. 1983, Frouz et al. 1997). This phenomenon, which
reflects physical and chemical effects of the soil biota´s microdisturbations such as anthills
or molehills - having a positive effect on vegetation succession is usually summarised in
the term ecosystem engineering (e.g., Jones 1994, Wright et Jones 2004, Jouquet et al.
2006, Sanders et van Veen 2011). It is apparently functional on spoil substrates enriched in
Journal of Landscape Ecology (2013), Vol: 6 / No. 1
heavy metals in abandoned tailings of our interest, too, where adjacent processes of the
soil improvement are facilitated in this way (Kovář 2004, Kovář et al. 2011, Štefánek et al.
Aim of the study
The main aim of the study is to compare plant cover on the tailings containment and the
influence of the ant activity on dispersal of plants, and facilitation of primary succession of
vegetation after approximately 10 years of spontaneous development. Periods of
comparison: the first one in 1998-2000 (sparse mosaic of lichens, herbs and shrubs;
Jarešová et Kovář 2004) and the second one in 2011-2012 (differentiated mosaic including
tree stands; Vojtíšek 2012).
Study site
An abandoned sedimentation basin with ore deposits is located on the left bank of the
Elbe river, close to the old surface quarry near Chvaletice in the northern part of Železné
hory hills, Czech Republic. Geographical location: 50°02' N, 15°26' E; altitude 200 m. Area
under spontaneous natural restoration: 40 ha. Habitat parameters: Waste sludge of Fe-Mn
ores with high sulphur and phenol contents were hydraulically transferred and deposited in
tailings containment (abandoned gradually in 1975-1980). The non-reclaimed basin has
remained largely treeless or non-vegetated and no manipulations were carried out after
abandonment (across the total surface area in 1984).
Series of vegetation succession (in 1998 and 2011)
Physiognomically distinct vegetational types correlating with successional stages were
selected: (1) in 1998 (a) mosaic of open areas with sporadic vegetation of mosses, lichens
and herbs, almost predominantly with the grass Calamagrostis epigejos), (b) mosaic of herb
stands with seedlings and saplings of aspen (Populus tremula) and birch (Betula pendula),
and (c) relatively tall (max. 3 m) stand of birch and aspen; (2) in 2011 (A) open space
with cryptogams, (B) herb stand (grassland), (C) shrubby stand (seedlings of Betula
pendula, Populus tremula, Salix sp. div., Pinus sylvestris, Tilia platyphyllos, Cerassus
avium, Sarothamnus scoparius), and (D) the most developed community with trees,
mainly birch (Betula pendula) and aspen (Populus tremula), about 5 m tall.
Phytocoenological relevés in plant stands were recorded using seven-grade scale of
abundance (Braun-Blanquet 1928). The results of this analysis in more detailed description
were published in the previous study (Kovář et al. 2011).
Frequency of ant nests in different plant stands
Dominant ant species were studied in the surface plateau of the abandoned sedimentation
basin: Tetramorium caespitum (Linnaeus, 1758); Lasius niger (Linnaeus, 1758), and
Formica rufibarbis (Fabricius, 1793) in 1998-2000; the last one was partly substituted by
Formica pratensis (Retzius, 1783) in 2011. Each of these species represents a different
size category - Tetramorium being the smallest, Lasius being medium-sized, and Formica
the largest. Inside above-mentioned successional formations, 10 x 10 m quadrats were
randomly set, three in each vegetation type. The plots were fixed and marked, closely
prospected and every location of ant nest was recorded.
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Ants and seed removal experiments
Ant activity supporting dispersal of plants and facilitating primary succession of
vegetation was studied with the help of repeated seed removal experiments (after the period
of approximately 10 years). Seeds were offered in especially constructed two-layer round
dishes, protecting seeds from other predators and from rain and the wind. The base of each
dish had a low edge to avoid problems with access, even by the smallest ant Tetramorium
caespitum. As the subject of this testing was only attractiveness of the seeds, not the
distance of their dispersal, seeds were exposed next to ant nests. In a growing season, seeds
were always offered from June until September when seeds were mature. The length of
exposure was eight hours (10.00 a.m. - 6.00 p.m.). Seeds of 20 plant species (1998-2000)
and more (2011-2012) collected around the ore deposits (just as they ripened) were used for
this experiment. In both periods, three size categories of dominant ants were tested in seed
removal experiments: (1) 1998-2000: Tetramorium caespitum, Lasius niger, Formica
rufibarbis, (2) 2011-2012: Tetramorium caespitum, Lasius niger, Formica pratensis.
Succession of ant species over ten years produced a two-fold increase in their diversity
(when Formica sanguinea and F. pratensis represent semi-parasitic species using the nest
occupation of the pioneers Formica cunicularia or F. rufibarbis.
Table 1: Parameters of particular plant species used in offer experiments (1998-2000)
Length of
Type of
Acetosella vulgaris
Agrostis tenuis
Anthoxanthum odoratum
Carex vulpina
Cerastium vulgatum
Holcus lanatus
Hypericum perforatum
Chrysanthemum leucanthemum
Lotus corniculatus
Luzula multiflora
Lychnis flos-cuculi
Plantago lanceolata
Poa pratensis
Potentilla argentea
Rumex obtusifolius
Trifolium dubium
Tripleurospermum inodorum
Vicia hirsuta
Vicia sativa
Vicia tetrasperma
Life form: H hemicryptophyte, Th therophyte. Length of life: P perennials, A annuals. Distribution:
UNSP non-specified, ANIMe zoochory, presence of elaiosome, ANIMa epi-zoochory (hair, feathers),
ANIMm epi-zooochory (sticky secretions). WIND anemochory. Plant family: Pgo Polygonaceae, Gra
Gramineae, Cyp Cyperaceae, Car Caryophyllaceae, Hyp Hypericaceae, Com Compositae, Leg
Leguminosae, Jce Juncaceae, Pla Plantaginaceae, Ros Rosaceae. According to Grime et al. (1988), and the
last author’s observation.
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Species diversity and effectiveness of myrmecochory
Species-area curves were constructed (2012) for plants by extending the sampling area
and counting the present plant species in each doubled quadrat (all this in four repetitions
on the (1) area with ant nests, and (2) area without ant nests of the abandoned tailings
containment. The potential argument is that differences in plant species diversity, observed
within both areas within the deposit plateau, were not primarily caused by presence/absence
of ant nests. Instead, these differences were attributed to different microhabitat conditions
that evolved the following transplantation control. Five plots 15 x 15 cm were placed
randomly in the area with an absence of ant nests, and the seeds of six relatively common
plant species in many places on the deposit were sown in them: Holcus lanatus, Vicia
hirsuta, Plantago lanceolata, Agrostis tenuis, Trifolium repens, Rumex acetosella. Their
seedlings were monitored during the growing season (2012).
Substrate properties
Six soil samples were taken from each of three layers (0-2 cm, 2-4 cm, and 4-8 cm,
respectively). The following characteristis of the substrate were analysed: incinerable C,
available P, pH and conductivity (measure of salinity) within all 4 vegetation types and/or
habitats (see above). All analytical procedures used were the same, and comparable with
previous studies in the locality (Kovář 2004; in details Vojtíšek 2012).
Ant species and ecological succession
Successional shift from three very weakly-defined clusters of vegetation in 1998
consisted of four relatively well differentiated types of habitats in 2012 (see Methods).
Results of the earlier term within the vegetation succession (Jarešová et Kovář 2004)
showed that lichen and moss species together with the annuals Vicia tetrasperma and V.
hirsuta occupy one semi-open environment. In contrast, woody species and other vascular
plants (Calamagrostis epigejos, Phragmites australis) represent the most advanced type of
plant stand.
It is possible to conclude that differences among vegetation relevés relate to vegetation
coverage. Since all three ant species studied in the period 1998-2000, namely Formica
rufibarbis, indicate their preference to occupy habitat mosaics with open surfaces.
However, the mutual relationship between ants and plants does not show significant
dependencies between an ant species and a particular plant species. Tetramorium caespitum
and Formica rufibarbis are closely associated with the presence of herbs, mostly of Vicia
species, and Calamagrostis epigejos. From nest numbers listed in Table 2, it is apparent
that Lasius niger is the most widespread ant of all the species in this locality. This species
builds its nests mostly in places with low growth of aspen and birch seedlings and saplings.
Tetramorium caespitum and Formica rufibarbis prefer surfaces with sparse vegetation.
The pattern of nests belonging to the aforementioned ant species during the 2011-2012
survey period, confirms the previous knowledge about their dependence on habitat and/or
vegetation types. An increasing proportion of tall woody stands slightly supressed typically
pioneer ant colonisers such as Formica rufibarbis or F. cunicularia. These species probably
became the target of attacks from successionally advanced species (Formica sanguinea,
F.pratensis) known as semi-parasitic invaders to a suitable phase of the ecosystem
development. This was the reason why attention was paid to behaviour of newly established
nests of Formica pratensis, apparently bound with periodic shadow by tree coverage and/or
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relatively dense vegetation. Intraspecific competition among nests of this Formica species
probably caused the success of the one nest (ant hill) with very high numbers of individuals
and wide territorial range of activity (tens of metres). However, it is relatively weak at
competing with the other size categories of ants (Tetramorium caespitum, Lasius niger).
Namely Lasius niger seems to be highly mobile with the construction of new nests in this
industrial environment, and it exhibits a ruderal-type adaptive strategy in diverse
microbahitat mosaics. A total of 12 ant species were identified within the locality: Formica
cunicularia, F. rufibarbis, F. cinerea, F. sanguinea, F. pratensis, Lasius niger, L. flavus, L.
platythorax, Myrmica schencki, M. rubra, Tetramorium caespitum, Temnothorax
crassipinus (underlined names indicate the species recorded in 2012 against the year 1998).
Table 2: Average numbers of ant nests per quadrat (10 x 10 m2) related to vegetation
types: 1 mosaic of open surface with cryptogams and sparse herb vegetation, 2 mosaic
of herb stands, and seedlings or saplings of aspen and birch, 3 woody stands with taller
growth of aspen and birch.
Assessment of variability in plant species richness on the abandoned sedimentation
basins (Vaňková et Kovář 2004) demonstrated high heterogeneity of especially the ore-
washery deposits such as in Chvaletice. Relatively distinct groups of plant taxa associated
with either the ash-slag or ore-washery deposits were distinguished. However, there are
numerous groups of species of wider ecological amplitude lacking a strict connection to the
type of industrial sediments (the bodies of deposits represent early stages of succession
and/or persist in early stages of succession for a long time). Transport of diaspores from the
surrounding landscape, mainly through zoochory and anemochory, is among the factors
governing the development of plant communities in the initial and/or young stages of
succession (Kovář 2004).
A typical S-shape in species-area curves was observed when number of plant species
plotted against increasing plots was demonstrated for the tailings surface (Fig. 1). However,
the steepness of the curve was considerably greater in the area colonised by the ant nests, in
comparison to the area without the ant nests and/or with only their sporadic incidence (the
highest number of nests in the frame of four quadrats 10 x 10 m monitoring was 30 in the
first case, and 9 in the second one). Low numbers of plant species were found in the interior
part of the abandoned sedimentation plateau in 1998 (less than 10), while in 2012 numbers
were approximately four-times higher. A similar relationship is observed for the areas with
extremely rare presence of ant nests in comparison with dense incidence of ant nests (Fig.
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Fig. 1: Species-area relationsips on the abandoned sedimentation basin in Chvaletice,
in the surface plateau area with only rare occurrence of ant nests (left) and in the
surface plateau area with high density of ant nests (right) data from 2011.
Experiments with removal and transportation of seeds by ants
Data from experiments with seed offered to particular ant colonies (1998-2000, Table 1)
were processed with the help of ANOVA. First, the species specific differences in seeds
collecting were tested, including test of significant differences. According to Tukey test,
Lasius niger showed the highest seed dispersal of the three ant species. However, there was
no significant difference in seed dispersal between Formica rufibarbis and Tetramorium
caespitum (Fig. 2). Figure 3 shows a different trend of seed collecting by Lasius niger
during the growing season (1999) compared with the two other ant species. Analogical
comparison at the same abandoned tailings containment 12 years later (Tetramorium
caespitum, Lasius niger,Formica pratensis) shows dominance in the seed dispersal activity
of the largest size category of ant species (Fig. 4). In the first case (Fig. 2) Lasius niger
dominates in gathering of seeds, in the second one (Fig. 4) Formica pratensis is the most
active species in the seed collection process (Table 3). In general, the results showed
significant differences in seed dispersal of several plant species related to collecting
effectiveness of ants.Ant workers of a particular species preferred certain plant species
(Jarešová et Kovář 2004). For example, Chrysanthemum leucanthemum was more preferred
than other plant species. In comparison, Hypericum perforatum and Vicia sativa were
collected in very small amounts.
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±1.96*Std. Err.
±1.00*Std. Err.
a b a
T. caespitum L. niger F. rufibarbis
Table 3: Two groups of plant seeds for experimental offering to ants according to
seasonal period (first group harvested in the second half of June and the first half of July
left, second group harvested in the second half of July and the first half of August), 2011-
First group of seeds for offer
Second group of seeds for offer
Potentila argentea (Po.Arg)
Holcus lanatus (Ho.Lan.)
Rumex acetosella (Ru.Act)
Hieracium laevigatum (Hie.Lae.)
Holcus lanatus (Ho.Lan.)
Plantago lanceolata (Pla.Lanc.)
Calamagrostis epigejos (Cal.Ep)
Trifolium arvense (Tri.Arv.)
Trifolium dubium (Tri.Dub)
Calamagrostis epigejos (Cal.Ep.)
Poa pratensis (Poa)
Melilotus officinalis (Mel.Off.)
Trifolium repens (Tri.Rep.)
Senecio jacobaea (Sen.Jac.)
Vicia hirsuta (Vic.Hirs.)
Lotus corniculatus (Lot.Cor.)
Agrostis tenuis (Agr.Ten.)
Picris hieracioides (Pi.Hie.)
Plantago lanceolata (Pla.Lanc.)
Trifolium repens (Tri.Rep.)
Lychnis flos-cuculi (Lich.F.C.)
Vicia cracca (Vic.Cra.)
Lathyrus tuberosus (Lat.Tub.)
Cirsium arvense (Cir.Arv.)
Centaurium erythraea (Ce.Ery.)
Fig. 2: Comparison of average numbers of effective seed collecting by three ant species
from arranged offers (the graph includes all data from the growing seasons 1998-2000).
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Fig. 3: Box plots show average values of seed collecting by a particular ant species
during 4 months of a season (data from 1999).
Fig. 4: Comparison of average numbers of effective seed collecting by three ant species
(F Formica pratensis, L Lasius niger, T Tetramorium caespitum) from arranged
offers (2011-2012).
Major behavioral similarity is noticeable between the ant species Tetramorium caespitum
and Formica rufibarbis, despite the great difference in their body size: both of them mostly
collected large diaspores of Luzula multiflora and Carex vulpina. Another major collecting
activity was noted for these two species with respect to seeds of Anthoxanthum odoratum
±1.96*Std. Err.
±1.00*Std. Err.
Tetramorium cae spitum
VI VII VIII IX Las ius nig er
Formic a rufibar bis
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and Potentilla argentea. Lasius niger mostly collected Anthoxanthum odoratum and P.
argentea, followed by Trifolium dubium and Vicia hirsuta. For generalisation of these
results, we must to take into account that dispersal of plant diaspores can be, to a certain
extent, affected by the season when the seeds were offered.
Substrate properties
In general, basic substrate properties of studied deposits in Chvaletice are a long-term
outcome of the process of sulphidic mineralization which was described by Rauch (2004).
More ecotoxicological aspects in the locality is treated in wider publication (Kovář
2004). Sampling of the substrate covered all 4 types of habitats distinguished for the
purposes of studying ant-plant interactions in the abandoned tailings containment during
period of 2011-2012: open space with cryptogams (A), herb stand (B), shrubby stand (C)
and the most developed community with trees (D). Six soil samplings were taken from each
of the three layers (0-2 cm, 2-4 cm, and 4-8 cm, respectively). The following characteristis
of the substrate were analysed: incinerable C, available P, pH and conductivity (measure of
salinity). A very rough illustration of the extent of the above-mentioned variables, which
could result in significant obstacles for the continuity of the ant colonisation of the
substrate, is shown in Table 4.
Table 4: Scheme of pairs of habitat types on the surface of tailing containment in
Chvaletice which significantly differ in a particular soil feature: open space with
cryptogams (A), herb stand (B), shrubby stand (C) and the most developed community with
trees (D).
Depth of soil profile
Substrate parameter
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Density of occurrence of ant nests within habitats and/or vegetation types, on the
abandoned plateau of the ore washery sedimentation basin, shows a relatively high degree
of dependence of ant species on vegetation cover in comparison with a particular plant
species. The most widespread ant species Lasius niger distributes its colony on places
sparsely overgrown by juvenile shrubs and trees. A similar observation was published by
Frouz (1996), who monitored occurrence of this species on dumps formed during brown
coal mining in the Sokolov region (western Bohemia, Czech Republic). Under
environmental conditions comparable with those described in this work, Lasius niger was
also a dominant species. It reached the highest values of abundance and biomass in the
initial stages of vegetation succession on dumps covered with shrub or herb vegetation.
However, on the ore deposit, Lasius niger tended to avoid surfaces overgrown by only herb
vegetation - evidently due to extremely high temperatures on the barren ore deposit surface
(60°C). The moss and lichen layer occurring in the successional stage with tree seedlings
represents a compact but dynamic cover of the soil surface (Pohlová 2004, Hroudová et
Zákravský 2004). The cover which is overheated in sunny days protects the soil below.
The space between the compact substrate and the surface layer offers eligible interface
shelter for ant migration and food transport including plant diaspores. In experiments with
Lasius niger (Brian et al., 1965), this ant survived on very dry soils only if adequate shade
was provided. It is ppossible that aspen and birch growing on ore deposits fulfil this
function. The two remaining ant species (Tetramorium caespitum and Formica rufibarbis)
resisted desiccation more than Lasius niger. This tolerance could explain higher occurrence
of those two ant species in areas with sparse vegetation.
In general, Lasius niger was the most successful ant species in myrmecochorous dispersal
of plants on the surface of the ore deposit. In comparison, in heathlands of southern
England, Tetramorium caespitum exhibited the greatest interest in seeds (Brian 1964).
Lasius niger showed a stronger preference to aphid honeydew than other ant species. The
most efficient seed collecting ant in a semi-natural ecosystem of mountain pasture was also
Tetramorium caespitum, one of 12 ant species present in this ecosystem (Kovář et
Kovářová 1998). However, seedlings of Vicia tetrasperma or Vicia hirsuta were observed
relatively frequently under the installed shelters covering the open substrate of the ore
deposit, which ants inhabited with their colonies - both the Vicia species were included in
seed offer experiments, and consequences of myrmecochory of this type are described in
numerous papers (e.g., Wilson 1971, Hölldobler et Wilson 1990, Jolivet 1996, Agosti et al.
2000). Preference of this ant to a specific plant´s seeds could explain the onset of these
annual Vicia´s following fire in August 1994 (Štefánek 2004).
Brian et al. (1967) also noted higher activity of ants in seed collecting in June and July,
compared to August and September. It is explained by increased dietary requirements due
to nurturing big larvae of sexual individuals in these months. These findings are similar to
our results obtained from the abandoned ore deposit in Chvaletice. The increase was
significant only in July (together with Tetramorium caespitum), but increased seed
dispersal was also exhibited by other species. Major differences between the months might
have been suppressed by low offer of insect feed in this generally poor biotope on ore
deposit, resulting in greater seed collecting activity throughout the year. Another
explanation is that seeds also serve as construction material for ant nests (lack of typical
soil, humus and litter with dead plant material are among the limiting factors). This use of
seeds is mentioned by a number of authors (Gorb et Gorb 1995, Gorb et Gorb 2000, Gorb et
al. 1997). Functioning as a vector of organic litter transport, the ants play a very important
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role in balancing extremity of this environment (overheating of the soil surface, high
concentration of heavy metals in the substrate, salinisation etc.), improving the substrate
conditions and supporting life and/or successional restoration of the habitat (Agosti et al.
Differentiation of the areas on deposit surface plateau into parts (1) poor in presence of
ant nests and (2) rich in density of ants and their nests correlated with plant species
diversity (1 low, 2 high) leads to the question: What is primary? Is is enviroment
unfriendly for establishment of plant seedlings, or does the problem consist of poor
availability and/or dispersal of seeds by myrmecochory? The transplantation experiment
with sowing the seeds of the most common plant species extensively spread throughout the
deposit plateau, rather supported the second hypothesis. Germination success of seeds
(previously tested for germination ability in laboratory conditions) on that part of deposit
with and/or without rare ant nests, confirmed the capacity of local microhabitats to facilitate
the growth of plants (Table 5). It is apparent that very weak success in germination is
exhibited by Plantago lanceolata and Trifolium repens. However, at the end of the growing
season several seedlings of Trifolim repens appeared as well as Rumex acetosella, which
can be presented as an effect of dormancy. Plant species Vicia hirsuta, Rumex acetosella
and Agrostis tenuis, followed by Holcus lanatus were successful. However, Vicia hirsuta
suffered from increased mortality of seedlings (three weeks post-germination) later in the
season. A certain degree of self-thinning after succesful germination of seeds of tested
plants, had a positive effect in increasing survival of the remaining individuals to maturity
and achieving successful reproduction. Similar observations were made by Dauber etal.
(2008), Lenoir (2009), Veen et Olff (2011) and Chen et Li (2012).
Table 5: Degree of germination and survival success of chosen plant seeds sown into
the deposit surface area characterised by absence and/or rarity of ant nests, and low
plant species diversity.
Number of germinated and survived seeds of 60 in total
Number of plot/plant
Biodiversity: The results show that species diversity of ants increased moderately (rather,
dominant species has changed) and diversity of plants increased several times during one
• Plant-ant relationship: there is a strong dependence of plant species diversity on presence
of numerous ant nests.
• Role of myrmecochory: manipulative experiments with plant seed removal (offer of seeds
near ant nests) demonstrated plant dispersal.
Journal of Landscape Ecology (2013), Vol: 6 / No. 1
• Survival of seedlings established after the ant transport of seeds into ant nests confirmed
the role of ants as eco-engineers in restoration.
• Selected parameters of the substrate in abandoned tailings containment show differences
among four habitat types (successional stages).
• Extreme soil conditions are balanced in areas inhabited by ants which facilitate the
acceleration of vegetation succession.
We thank to P. Pech (Dept. Biology, Faculty of Science, University of Hradec Králové)
and J. Frouz (Inst. Environment, Faculty of Science, University of Prague) for
determinantion of ant species. The work has been supported by the grant no. 206/93/2256
of the Grant Agency of the Czech Republic “Biotic interactions during vegetation
succession on toxic substrates”, grant no. 200/1997/B/BIO of the Grant Agency of Charles
University and by the research project MSM 113100004, chapter “Ecological plasticity and
taxonomical variability of expansive plant species“ of the Czech Ministry of Education.
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... On the plant-ant relationship, the focus has been mainly addressed to the role of ants in seed dispersal (Crist & Wiens 1994;MacMahon et al. 2000;Ness et al. 2004;Guarino et al. 2005) and in affecting plant growth through defoliation and herbivory (Dibner et al. 2015;De Almeida et al. 2020). More rarely researchers have investigated the role of vegetation as habitat for ant species (Danin & Yom-Tov 1990) and the effect of ant species on plant communities (Dean et al. 1997;Kovář et al. 2013;Nemec 2014). In fact, the presence of anthills and mounds in grasslands may disturb and modify the plant assemblage structure with change in species richness, diversity, patchiness and evenness (Dean et al. 1997;Nemec 2014). ...
... Therefore, further investigations could develop these aspects (soil chemistry, species-specific differences between anthills). However, this study could be considered as a first step that clarifies some structural differences between plant assemblages of anthills when compared to those not disturbed typical of the urban Mediterranean grasslands dominated by Dasypyrum villosum, therefore corroborating the role of ants as ecological engineers (Kovář et al. 2013;Ghobadi et al. 2016). ...
We explored the structure of plant assemblages that settles around the anthills of a guild of Hymenoptera Formicidae (Messor wasmanni Krausse 1910, Tapinoma nigerrimum Nylander 1856 and Aphaenogaster spinosa Emery 1878), observable in urban grasslands dominated by Dasypyrum villosum (Rome, Central Italy). Since it is known that ants act as a disturbing factor on plant assemblages of grasslands, our hypothesis was that vegetation structures suffer from some stressors that affect, in this plant association, the dominant plant structure. We compared the plant assemblages observed in the plots of the anthills with the control grassland assemblages using a diversity/dominance diagram. We recorded 63 plant taxa. The average number of plant species was found to be significantly lower in anthill plots than in control grassland plots. In anthill plots, dominant plant species (Polygonum romanum, Poa trivialis Vulpia myuros, Aira elegantissima and Vulpia ligustica) have been found to be different from control grassland plots (Convolvolus arvensis, Dasypyrum villosum, Poa trivialis and Sheradia arvensis). Anthill plant assemblages were found to be significantly different (One-way PERMANOVA) and poorer in terms of species richness, less diversified, and with a reduced species turnover than control grassland plots. Whittaker plot analysis seems to show that the plant assemblages of the control, with greater evenness, emphasize a stressed condition in anthill plant assemblages. Non-metric multidimensional scaling shows a set of species strictly linked to anthills. Our data seem to confirm the role of ants as a stressor in the plant assemblage structures of Mediterranean urban grasslands. However, our results also show that the peculiar ant nest conditions may favor plant species less represented in the Dasypyretum grasslands (i.e. Polygonum romanum) or even absent in this plant assemblage (i.e. Aira elegantissima, Cynodon dactylon and Poa annua), thus increasing the overall local plant diversity.
... There is great inconsistency in nutrient composition among studies (Farji-Brener and Werenkraut 2017) and anthills are sometimes richer in three fundamental nutrients i.e. phosphorus, nitrogen and carbon (e.g. for P-Lasius niger; Wu et al. 2010 for N, P-Formica sanguinea, Lasius flavus, C-Formica sanguinea; Ehrle et al. 2017 for P-Lasius flavus). This is presumably due to the organic material deposition (Beattie and Culver 1983;Rudolphi 2009;Kovář et al. 2013), however opposite results were also found (e.g. Dean et al. 1997-for N, P-Lasius favus, Lasius alienus and Formica rufibarbis; Frouz 2006, Ehrle et al. 2017 for C, N-Lasius flavus). ...
... Dispersal of seeds of myrmecochorous plants (myrmecochores), which provide nutrient rich elaiosomes as reward for ants, is well known (Sernander 1906). Nevertheless, ants were also demonstrated to disperse some seeds without elaiosomes (Kovář et al. 2013). Thus, the total number of seeds was reported to be higher in nests (Schütz et al. 2008). ...
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Ants can shape vegetation as seed dispersers and ecosystem engineers. When anthills are long-lasting, they are known to change soil and vegetation characteristics. However, it is unclear whether plant species traits and species composition vary between ant guilds and between parts of individual anthills. We compared different aspects of soil and vegetation (composition, seedling abundance, and functional traits) between anthills and the surrounding mesophilous pasture in Czechia. This pasture hosts eight ant species, which belong to both seed dispersers and non-dispersers. Where feasible, we divided anthills into centres and margins for the analyses. Anthills (area 90.5–4051.7 cm²; 5–19 plant species) displayed different and more heterogeneous (less similar within anthill plot type) vegetation composition when compared to the surrounding area, with more seedlings and several species restricted to anthills. Further, anthills were more functionally diverse and exhibited several differences in traits, both at the community and intraspecific level. Anthill centres had higher surface temperatures in hot sunny days, higher levels of phosphorus and pH than margins, while margins had higher moisture and carbon content than surrounding vegetation. Further, anthill vegetation differed between ant guilds with more myrmecochorous species found at nests of seed dispersers. Overall heterogeneity in this mesophilous pasture was enhanced by the presence of anthills. Further, the anthills themselves are heterogeneous due to variable sizes, persistence, and differences between their centres and margins on long-lasting anthills. Anthills can thus enhance plant diversity by maintaining disturbed microsites and enhancing the growth of seedlings and less competitive plants.
... Ants were found as pioneers and subsequent colonizers of this industrial waste deposit from the viewpoint of their functioning in plant seed dispersal. So ants play the role in primary vegetation succession (Kovar et al, (56) 2013) . ...
Agricultural systems are areas of land where humans manipulate physical, chemical, and biological processes using a cluster of practices to obtain a benecial product for their use. To understand the level of organization in a particular region we must consider biotic and abiotic factors and the interaction between them to employ best conservation practices to restore biodiversity in this region. Ants perform a different role in agro ecosystems like pollination, soil turbation, bio indication and the regulation of crop-damaging insects. Agro ecology management of ants in tree cropping systems required some information about positive and negative impacts on crops and pest species. This review article provides an overview of knowledge of the roles performed by ants in agricultural habitats.
... Ants (family: Formicidae) occur in almost all terrestrial ecosystems, while tropical regions harbor peaks of their diversity [1]. Some ants are key predators [2,3], ecosystem engineers [4][5][6], seed dispersers [7,8], and biocontrol agents [9,10]. However, some ant species are notorious pests of households, agriculture, and forests. ...
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The information available on the diversity of ant species and their distribution and interaction with forest health in Nepal remains limited. As part of a nationwide project on forest health, we conducted inventories to assess the diversity and distribution of forest ants and their role in forest management in Nepal. Ants were collected from 187 plots of 10 m × 10 m size along the north–south belt transects in eastern, central, and western Nepal. We used vegetation beating, sweeping, and hand collection methods in selected forest types. In each transect, we designed six plots in each major forest type (Sal, Schima–Castanopsis, and broadleaf mixed forests) and three plots each in deodar, Alnus, riverine, and Cryptomeria forests. We recorded 70 ant species from 36 genera and six subfamilies. This includes five genera and nine species new for the country, as well as eight tramp species, four of which are major ecological, agricultural, and/or household pests. Our study indicates that forest ant species richness is high in western Nepal and the Siwaliks, and it decreases as elevation increases. The high diversity of ant species in the forests of Nepal needs to be assessed with further exploration using multiple sampling methods covering all seasons and forest types. Ants can be useful indicators for ecosystem management and human impacts on forests. Reports of invasive ants in Nepalese forests indicate the relevance of urgent interventions through sustainable forest management initiatives to prevent future incursions.
... The situation is different if such transport was preferentialonly plant macro remains -or the transport of pollen was of too limited magnitude to induce a recognizable homogenization of the pollen archive. Preferential downward transport of seeds is for example known from ants (Majer et al., 2008;Robins and Robins, 2011;Kovář et al., 2013). Evidently, such faunal activity will not extend below the mean lowest groundwater level, but short dry spells may have sufficed for some bioturbation to have occurred. ...
Distal tephra from the major Somma-Vesuvius Avellino (AV) eruption is widespread in the coastal basins of Southern Lazio (Central Italy). Dated to 1995 ± 10 cal yr BC in 2011, later on doubts arose about the reliability of this frequently cited age. This led to a major effort to date AV tephra holding sections, based on a thorough methodological approach. Various aspects were studied to identify sections yielding reliable ¹⁴C ages, including bioturbation, inbuilt age, and variable sediment accumulation rate. Lowered rates upon deposition of tephra, particularly in anoxic marshy environments and attributed to toxic F contents, showed up as sharp increases in pollen density. The ‘sampling error’ was quantified for specific sedimentary environments and derived from coring data and published data on accumulation rates for similar Central Mediterranean sites. Next, two Bayesian analyses were performed, a traditional using the full set of samples and a novel, based on samples that were deemed as suitable (no bioturbation, inbuilt age, etc.) and of which the age was corrected for the sampling error. The age obtained by the novel analysis had the smallest range (1909–1868 cal yr BC), differs about a century, and is virtually identical to the ages published by Passariello et al. (2009) and Alessandri (2019). The earlier found age (2011) is ascribed to a statistical coincidence. The results solve a long debate on the age of the AV eruption, which is the youngest of the three major eruptions in the Central Mediterranean Bronze Age. Ages of the other two, the Agnano Mt Spina (Phlegrean) and FL eruption (Etna), are still uncertain and disputed. This study illustrates the need for a thorough approach in ¹⁴C dating tephra holding sediment archives in the Central Mediterranean, and employed a methodology that can be applied in such approach. Attention is called for potentially toxic fluorine concentrations in Campanian tephra, which may have had a serious impact on the contemporary environment and induced chronological hiatuses, but hitherto were not reported for the early tephra.
... A strongly cemented horizon with red ferric oxides and gypsum developed at lower depths of soil profiles (Mrázek, 2004;Kovář et al., 2011) and caused retrogradation in successional progress through periodical extinction od the tree seedlings (Kovář et Herben, 2004). Repeated restoration of initial colonization with biota includes the microhabitat dynamics at small-scale level (creation of biological soil crust) and processes of the organic matter decomposition (litter fall, dead biomass supply), bioturbation activity of the soil fauna, establishment of plant seeds and their seedlings or the insect/plant interactions, seed dispersal by zoochory and anemochory improve and use the environmental properties in the mosaic of microsites and shift the ecosystem far towards life-supporting state of spontaneous balanced succession (Kovář, 2004;Neustupa et al., 2009;Štefánek et al., 2012;Dostál et Kovář, 2013;Kovář et al., 2013). Hence, above mentioned contradictions imply irregular shifts in vegetation succession on toxic postindustrial wastes and differentiated qualitative composition of plant stands inside and outside the deposited substrate body. ...
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This pilot case study compares genome sizes of two groups of species (conspecific plants) which spontaneously colonize interior space within abandoned industrial area and/or deposits, and those ones occurred in adjacent vicinity. Testing of the hypothesis “There is functional significance of small versus large genomes of plant species by comparing their occurrence in unreclaimed toxic deposits as an example of stressed environment and in their populations from neighbouring habitats” confirmed this idea.
... On the other hand, the presence of ants, which act as an ecosystem engineers, can promote small-to-large scale changes, such as soil water infiltration rates, erosion reduction and organic material increment (Cammeraat and Risch, 2008;Cerda and Jurgensen, 2008;Cerda et al., 2009), which enable positive interactions with the flora. The role of ants in establishing the plant community can take many forms, from facilitating mycorrhizal colonization with nitrogen and phosphorus enhancement (Dauber et al., 2008) to seed dispersal during the restoration process (Kovář et al., 2013). ...
The full text is avaliable in a personalized URL providing 50 days' free access to the article. Anyone clicking on this link before April 01, 2020 will be taken directly to the final version of article on ScienceDirect, which they are welcome to read or download. No sign up, registration or fees are required: The aim of this study was to evaluate the influence of succession stages of regeneration of the Brazilian savanna (Cerrado) on the richness, diversity, trophic guilds, and community composition of ants looking for taxonomic and functional changes in the community. The areas included an active pasture that had been created within the past six months, areas with several years of abandonment with different degrees of natural Brazilian savanna regeneration (from 2 to 15 years), and two large remnants of native forest that had been preserved for at least 40 years. Collections were carried out in 56 plots using sardine baits and an active search for a period of one hour in each plot. In addition to calculating the regeneration age, we evaluated successional stages by measuring vegetal characteristics in each plot. A total of 60 species was obtained, distributed in 28 genera and 8 subfamilies, and these demonstrated a direct association with the regeneration age of the areas, as well as their richness and diversity of ants. Areas with initial regeneration showed a greater proportion of generalist species and, as plant succession increased, generalist species decreased. A temporal threshold of 5–6 years for regeneration age was observed, when the community structure tended to stabilize. We observed that the maintenance of a minimum quantity of vegetation in areas destined for livestock can efficiently preserve ant diversity.
A brief and comprehensive introduction to restoration ecology is given. After a short sketch of the history of this subdiscipline of ecology, concepts, general objectives, reference systems, and a definition of ecosystem restoration are presented. Since ecosystem restoration is based on ecological key concepts such as e.g., species pool, functional groups, succession, stress, disturbance, and ecosystem functions, it might be an “acid test” for ecology. General objectives could be the restoration of natural ecosystems, traditional and diverse land-use systems, and “novel” ecosystems without a reference in the natural or traditional cultural landscape. By giving an overview on measures, the practice of ecosystem restoration is addressed. An emphasis is put on the transdisciplinary peculiarities of restoration ecology and ecosystem restoration, bridging the natural and the social sciences.KeywordsEcological key conceptsEcosystem restorationNovel ecosystemsRestoration ecologyRestoration measuresTransdisciplinarity
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Post-industrial sites form a unique phenomenon in the landscape. They enable us to study the human-altered succession of communities. Regarding this, we studied an ant community in three types of habitats-reclamation and spontaneous succession in an ore basin together with unaltered surroundings in the Czech Republic. More than 30 years after being abandoned, the site with spontaneous succession was more species rich than the reclaimed one. Moreover, spontaneous succession created a habitat that was more similar regarding ant diversity to the unaltered surrounding environment than that after traditional reclamation. Ants dependent on tree vegetation were rather rare in both the reclaimed and spontaneous succession parts of the ore basin compared to the surrounding landscape. The relative abundance of socially parasitic ants increases in a gradient from the reclaimed basin, through the basin with spontaneous succession to the unaltered surroundings. Our study highlighted the fact that the formation of ant communities at post-industrial sites is clearly more complicated than for other arthropods, including related aculeate hymenopterans. The potential of both reclaimed and spontaneous succession basins for harbouring endangered species appeared to be lower for ants than for other taxa indicated by recent studies.
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In Shapotou, northeast of the Tengger Desert in China, Formica cunicularia Lat. proliferated after mobile sand dune stabilization by artificial revegetation. In the present paper, the anthill properties of F. cunicularia after sand stabilization were systematically investigated along a chronosequence. The soil properties and soil seed banks in the anthill and adjacent soils without anthills were also examined. The results showed that the revegetation age, landform, and their interactions significantly affected the anthill density, anthill coverage, and soil turnover capacity of F. cunicularia. The longer the dunes were stabilized, the more ant nests proliferated in the dunes. More nests were located in the hollow and windward slopes than in the dune top and leeward slopes. Soil organic matter, total and available N, P, and K, electrical conductivity, as well as soil water content were all significantly higher in the nests than in the adjacent soils without anthills. The soil seed bank in the anthill was much greater than in the adjacent soil for both moss- and algal-crusted soils. The current research suggested that the occurrence of F. cunicularia after mobile sand dune stabilization greatly influenced both soil properties and soil seed bank.
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Of the three main seed-transporting ant species on the plateau of an abandoned ore sedimentation basin, the most efficient was Lasius niger. Seeds of 20 plant species, both myrmecochorous and non-myrmecochorous, were collected in dif-ferent amounts by different ant species. Chrysanthemum leucanthemum, Vicia sativa, Carex vulpina, Anthoxanthum odoratum and Potentilla argentea were preferred over the course of a season. The three dominant ant species studied show also different habitat preferences. Lasius niger builds its nests mostly clo-se to aspen and birch seedlings growing from a cover of lichens and mosses, whi-le Tetramorium caespitum and Formica rufibarbis inhabit more open surfaces with sparse vegetation.
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This article examines the consequences of a fire (August 1994) which burned the vegetated surface of abandoned tailings containment in mosaic pattern, in Chvaletice (Eastern River Elbe basin, Czech Republic). Vegetation succession with adjacent processes (such as leaf litter decomposition) was then monitored in comparisons on burned and unburned plots. The influence of the introduced plant organic matter by mulching on the biological soil crust was also investigated. Apart from the naturally burned plots, some plots were also experimentally burned. The soil moisture and surface temperatures in different temporal distances from the fire experiment were measured. There were no significant differences in decomposition rate between the plots that were affected and/or unaffected by fire. The daily temperatures of the burned soil surface increased after the fire, while its night temperatures decreased. The moisture content of the soil surface was reduced after the fire. Addition of dry local aboveground plant biomass on the substrate surface suggests that substrate roughness provides retention to seeds transported by wind, and protects the rhizosphere against extreme heat, drought and salt incrustation. Furthermore, it positively modifies the hydrological regime of microsites and stimulates the creation of a humus soil layer and enrichment of the substrate by nutrients. Finally, it facilitates colonisation by plant seedlings from seeds transported by anemochorous or zoochorous mechanisms.
The mounds built by Lasius flavus F. (Hymenoptera: Formicidae) reflect the history and age of the grasslands in which they occur. An index based on ant-hill size was sucessfully used to estimate the dates at which thirteen chalk grasslands in Wiltshire, England, were last ploughed. The mean volumes of the five largest and-hills out of a thousand increased linearly with the actual age of the grassland, in fields last ploughed 54-164 years ago. Other evidence which supports the hypothesis that large ant-hills only occur in old grasslands is discussed.
Attractiveness of five deciduous forest myrmecochorous and diplochorous plants for Formica polyctena ants were studied in field experiments. Seeds of different plant species were removed at different rates (RRS) by ants. Experiments with demonstrations of seeds of one or a pair of species and with partly cut elaiosomata or seed bodies showed that seed attractiveness increased with seed size increase in Viola matutina, Chelidonium majus, V. mirabilis, V. hirta and Asarum europaeum, and was weakly correlated with elaiosome size. The exponential character of RRS from groups was especially pronounced for pure myrmecochors. The RRS obtained for F. polyctena near (5 m) were about ten times shorter than those obtained far (15 m) from the nests. Ants taking seeds of A. europaeum, V. hirta and V. mirabilis were significantly larger than ants taking Ch. majus. It is suggested, that in habitats with colonies of F. polyctena with a prevalence of small individuals, the small-seed myrmecochors receive some advantages in seed dispersal, and in habitats with colonies with a prevalence of large size classes - the large-seed myrmecochors have an advantage.
The vegetation and soil chemistry of 15 abandoned mounds of Formica canadensis were compared to control quadrats in a meadow at 2900 m near Gothic, Colorado, USA. Principal Components Analysis indicated the mound vegetation was relatively homogeneous and distinct from nonmound vegetation. Discriminant Analysis indicated that mounds were characterized by Bromus polyanthus and Achillea millefolium, and nonmounds by Poa interior. A parallel analysis of 15 soil chemicals indicated some statistically significant chemical differences between mounds and nonmounds. Several micronutrients (Fe, Zn, Mn), were lower in mounds but were unlikely to be deficient. Ant-dispersed plants (myrmecochores) were almost entirely restricted to nonmound quadrats.
Seed dispersal by ants was studied in three Mediterranean old-fields (6, 15 and 43 years after abandonment) and a Quercus pubescens forest that represented the last successional stage. In each plant community, we investigated 1) the occurrence of both seed dispersal strategies involving ants, namely myrmecochory and dyszoochory, relatively to plant species frequency and cover, 2) the abundance of disperser species likely to perform one dispersal type or the other, 3) the relative effectiveness of myrmecochory and dyszoochory, by experimentally offering elaiosome-bearing seeds of Viola alba and Euphorbia nicaeensis, and seeds of the same species deprived of their elaiosome. The proportion of dyszoochorous plants and the abundance of granivorous ants followed the same trend along the successional gradient. The youngest stage had the highest proportion of dyszoochorous plants and the highest abundance of granivorous ants, including the harvester ant Messor structor, whereas the oak forest showed the lowest proportion of dyszoochorous plants and held no granivorous ants. In contrast, there was no coincidence during the succession between the number of myrmecochorous plants and the number and abundance of ant species collecting elaiosomes. More generally, myrmecochorous plants were rare while ant species collecting elaiosomes were present from old-field to forest. Seed removal experiments showed an overall significant increase in the removal rate of elaiosome-bearing seeds over that of seeds without elaiosome, but in the youngest old-field the difference was not significant for V. alba and was attenuated for E. nicaeensis seeds, due to high harvester ant activity. We suggest that dyszoochory by ants could be a major mechanism influencing plant dynamics in open and grassy vegetation in the Mediterranean region. We also suggest that, though apparently efficient, myrmecochory is underrepresented because of biogeographical features (rarity of the medio-European flora) and lack of strong selective pressure. We emphasize that dispersal strategies involving animals, be it ants, birds or mammals, all share the same dichotomous pattern that shows the evolutionary strength of two mechanisms, one directly derived from predation (dyszoochory), the other generally more deeply shaped by mutualistic processes (e.g. myrmecochory).
1. The worker populations of a number of marked colonies of the ant Tetramorium caespitum have been estimated by Petersen's mark-recapture method in two consecutive years. 2. At the same time the area of territory used, the wet weight and the head width of workers and the numbers and weights of sexual forms have been obtained. In one year seed production per territory was also estimated. 3. Analysis shows that worker population does not correlate significantly with any of these variates. 4. Territory area, worker size, and male and female sexual production are all positively and significantly correlated. This is taken to indicate that variation in resources influences the production of sexuals (and their precursive large workers) rather than the worker population size. Colony maturity is thus a condition that may be reached at a wide range of colony sizes. 5. Most colonies were mature and the total sexual weights were 43% (1963) and 53% (1964) of the worker biomass. 6. With a food supply estimated at about 240 g/year at least, this ant needs less than 10 g new material a year to continue to survive and emit sexuals.