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Entomofauna carpathica, 2024, 36(2): 83-126
83
ANTS (HYMENOPTERA: FORMICIDAE) OF THE EASTERNMOST
PART OF BIELE KARPATY MTS AND ADJACENT AREA
Lukáš JANCÍK
Keblianska 374/38, 020 01 Streženice, Slovak Republic. E-mail: lukas.jancik.857@gmail.com
JANCÍK, L. 2024. Ants (Hymenoptera: Formicidae) of the easternmost part of Biele Karpaty
Mts and adjacent area. Entomofauna carpathica, 36(1): 83-126.
Abstract: The fauna of ants was studied at 73 sites distributed within the easternmost
part of Biele Karpaty Mts and adjacent area (north-west Slovak Republic). All discovered
areas with xerothermophilous grasslands, which were scarce in the investigated area,
were included. The sites also comprised other natural habitats, e.g., wetlands, as well as
ruderal ones disturbed by human activities. In total, 75 ant species were discovered
there, including numerous rarely found ones inhabiting xerothermous forest-steppes,
e.g., Lasius myops Forel, 1894, Lasius reginae Faber, 1967, Proceratium melinum (Roger,
1860), and Strumigenys argiola (Emery, 1869). Some other poorly recorded species
within the Slovak Republic were also discovered, i.e., Lasius sabularum (Bondroit, 1918),
Leptothorax gredleri Mayr, 1855, and Tetramorium immigrans Santschi, 1927.
Noteworthy records are represented also by arboricolous species known to nest in dead
tree parts too, i.e., Lasius bicornis (Förster, 1850), and Lasius citrinus Emery, 1922.
Faunistic data are listed by species, containing all confirmed study sites. A summary of
available past records is provided for some of the discovered species. This study is
enriched by data on the biology of ant species, which were obtained during this study,
e.g., swarming dates, nest construction, and colony demography.
Key words: faunistic, Lasius reginae, Lasius bicornis, ecology, Javorníky Mts, Slovakia,
Považské podolie Val.
INTRODUCTION
The past studies of myrmecofauna within the Biele Karpaty Mts and its adjacent
area were well summarised by BEZDĚČKA & BEZDĚČKOVÁ (2010). The earliest
references to individual findings are documented in ZÁLESKÝ (1939); more
complex studies were carried out by LABUDA (1970), BEZDĚČKA (1992), BEREC
(1997), to a large extent by DEVÁN (2005, 2006a, b, c, 2008b, 2009b, c), and by
DEVÁN (2008). Some additional single records can be found dispersed within
various papers, e.g., ŠILHAVÝ (1935) mentioning three ant species from Kvašov
and Vršatec villages, and SEIFERT (2012) mentioning one finding of Bothriomyrmex
corsicus Santschi, 1923 from Mikušovce village. This study focuses exclusively on
the easternmost part of the Biele Karpaty Mts and adjacent area covering the
Entomofauna carpathica, 2024, 36(2): 83-126
84
Javorníky Mts and Považské podolie Val. (Fig. 1). The study sites in the Biele
Karpaty Mts were in the surroundings of inhabited areas Záriečie (Klecenec),
Zubák, Dohňany, Horná Breznica, Hrabovka (Púchov), Keblie (Púchov), Lednica,
Streženice, and Medné. The localities covered by adjacent part of the Javorníky
Mts were situated near inhabited areas Dešná (Lysá pod Makytou), Lysá pod
Makytou, Záriečie, Zbora, Mostište, Ihrište, Dohňany, and Hrabovka (Púchov).
Within the adjacent part of Považské podolie Val. were included sites in proximity
of village Streženice, and town district Horné Kočkovce (Púchov). Forest-steppe
localities with xerothermophilous grasslands are rare in the investigated area,
span small areas, and most of them are abandoned without any conservation
management, leading to succession process and forest development (Figs 2A, C, D, 3).
MATERIAL AND METHODS
Description of study sites
A list of the study sites (Fig. 1) with their descriptions is presented below. Each
site is described by its unique identification number (1), altitude (Z-coordinate)
[m a.s.l.] (2), coordinates (Y,X) [decimal degrees] (3), geomorphological unit it
Fig.
1
.
Overview of collecting sites: White points (46) indicate sites within the Biele
Karpaty Mts, green points (22) mark sites in the Javorníky Mts, and yellow points
(5) show sites in the Považské Podolie Valley. Red lines outline the boundaries of
the geomorphological units. Picture was created using QGIS software.
Entomofauna carpathica, 2024, 36(2): 83-126
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belongs to (4), administrative affiliation into the cadastral territory it belongs to
(5), and by the biotope (6). The altitude refers to 2D point of the “Y,X”
coordinates. The attribute n. 3 represents approximate value and refer to the
most prominent site’s part. The accuracy of the coordinates is given by the
number of their decimal digits (4). Sites are in descending order based on the “Y”
coordinate, meaning that they are sorted from the northernmost one.
1) 404; 49.2064°,18.1644°; Javorníky; Lysá pod Makytou; Subsoil: unknown. Small
concrete surface serving as a base for a bridge over a stream. Stone-lined stream and
solitary trees in its surroundings.
2) 476; 49.1902°,18.1862°; Javorníky; Lysá pod Makytou; SE facing and very steep
hillside. Subsoil: sandstone. Grassland biotope with occasional stones on the ground surface.
3) 401; 49.1830°,18.2859°; Javorníky; Zbora; Subsoil: unknown. Stands of old Tilia
platyphyllos trees with much dead bark on their trunks, and adjacent area of abandoned
cemetery with dead wood.
4) 402; 49.1800°,18.2591°; Javorníky; Mestečko; Non-overgrown abandoned quarry
with S facing slopes. Subsoil: limestone. Surfaces with bare rocks, gravelly substrate, and
occasional xerothermous grasslands.
5) 401; 49.1720°,18.2360°; Biele Karpaty; Záriečie; S facing and very steep hillside.
Subsoil: limestone. Afforested xerotherm mainly with approximately 70-year-old Pinus
sylvestris stands, but still with small clearings. Rocky soils, subsoil protruding from the
soil in large extent, much of bare stony surfaces. The area directly borders with a small
stream called Klecenecký potok which flows directly under the hillside and is notched to
the subsoil (Fig. 2A).
6) 393; 49.1705°,18.2354°; Biele Karpaty; Záriečie; SE facing slope of a small hill.
Subsoil: limestone. Xerothermophilous grasslands with occasional stones on the ground
surface.
7) 454; 49.1630°,18.3079°; Javorníky; Mostište; S facing slope. Subsoil: limestone.
Sparse Pinus sylvestris forest with undergrowth of grass-herbal vegetation, occasional
stones, and partially burned ground surfaces.
8) 325; 49.1602°,18.2724°; Javorníky; Dohňany; SE facing terrain depression. Subsoil:
sandstone. Grasslands and shrubs with occasional stones on the surface, on an
unmanaged small area.
9) 362; 49.1597°,18.3061°; Javorníky; Ihrište; S facing and very steep hillside. Subsoil:
limestone. Xerotherm with much rock on the ground surface but overgrown with bushes
and trees.
10) 450; 49.1539°,18.3022°; Javorníky; Ihrište; SE facing slope of a hill. Subsoil:
limestone. Small forest clearing with much rock on the surface.
11) 404; 49.1524°,18.3092°; Javorníky; Ihrište; Subsoil: unknown. Country road and
adjacent mowed grass areas.
12) 382; 49.1510°,18.3029°; Javorníky; Ihrište; Subsoil: unknown. Extensive
grasslands with old apple trees and much dead wood.
13) 419; 49.1509°,18.3006°; Javorníky; Ihrište; E facing hillside. Subsoil: limestone.
Small forest clearing with grasslands and many stones on the ground surface.
14) 457; 49.1498°,18.3150°; Javorníky; Ihrište; SE facing and very steep hillside.
Subsoil: sandstone. Forest clearings with much of bare soil surfaces and stones on the ground.
Entomofauna carpathica, 2024, 36(2): 83-126
86
15) 480; 49.1493°,18.2337°; Biele Karpaty; Zubák; SE facing hillside. Subsoil:
limestone. Forest-steppe with xerothermous grasslands and shrubs, also with much rock
on the ground surface.
16) 447; 49.1492°,18.3176°; Javorníky; Ihrište; SW facing slope. Subsoil: sandstone.
Forest clearings with much of bare soil surfaces.
17) 436; 49.1486°,18.3173°; Javorníky; Ihrište; SE facing slope of hill. Subsoil:
limestone. Light Pinus sylvestris forest with rocky soil and grass-herbal undergrowth.
18) 326; 49.1484°,18.2896°; Javorníky; Dohňany; SW facing slope of a hill. Subsoil:
claystone. Surroundings of a branched and little-used country road with mostly bare
ground surfaces and grasslands; in connection with forest ecotone.
19) 399; 49.1475°,18.2989°; Javorníky; Ihrište; Subsoil: limestone. Forest clearing with
xerothermophilous grass-herbal vegetation and much of stones on the ground surface.
20) 354; 49.1451°,18.2993°; Javorníky; Ihrište; E facing and very steep slope of a hill.
Subsoil: limestone. Xerothermophilous grasslands with occasional shrubs and stones
(Fig. 2D).
Fig. 2. Biotopes at study sites. A: Forest clearing on a steep hillside – habitat of
Camponotus piceus and Plagiolepis pygmaea. Photo: September 2019. B: Older
concrete surface with limited vegetation, and gravel-sand of a country road –
habitat of Tetramorium immigrans. Photo: June 2022. C: Forest-steppe in
succession stage – habitat of Tetramorium caespitum, Camponotus aethiops, and
Formica gagates. Photo: July 2022. D: Forest-steppe with xerothermophilous
grasslands on a steep hillside, and adjacent pasture in lowland of a stream. Photo:
September 2023.
Entomofauna carpathica, 2024, 36(2): 83-126
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21) 350; 49.1445°,18.2706°; Javorníky; Dohňany; S oriented slope of a hill. Subsoil:
limestone. Light Pinus sylvestris forest with larger stones surpassing the ground surface.
22) 422; 49.1443°,18.2656°; Javorníky; Dohňany; SW to SE facing and very steep slope
and upper-situated less steep area. Subsoil: limestone. Rocky hill with xerothermophilous
grass-herbal vegetation.
23) 350; 49.1439°,18.3026°; Javorníky; Ihrište; Subsoil: unknown. Tilia stands in
shaded forest area.
24) 300; 49.1394°,18.2895°; Javorníky; Dohňany; E oriented slope of an uphill.
Subsoil: sandstone. Small area in proximity of a little-used country road where occurs
occasional stones on the ground surface.
25) 645; 49.1366°,18.2509°; Biele Karpaty; Dohňany; Upper-situated area. Subsoil:
sandstone. The crest of a hill containing a country road and its surrounding – mixed
forest, and grass areas made up after forest logging. Semi-shaded area.
26) 276; 49.1343°,18.2938°; Biele Karpaty; Vieska-Bezdedov; Subsoil: unknown.
Agricultural and mowed grass areas with single older deciduous trees close to the river.
27) 456; 49.1335°,18.2810°; Biele Karpaty; Vieska-Bezdedov; Upper-situated area.
Subsoil: limestone. Top of a rocky hill with much rock on the surface and occasional
xerothermophilous grass-herbal vegetation.
28) 280; 49.1333°,18.2932°; Biele Karpaty; Vieska-Bezdedov; Subsoil: unknown. Edge
of deciduous forest with many native rocks on the ground surface (a natural part of this
site), and adjacent tree-free area with human-brought gravel and other geological
material, also railway embankment. Partially ruderal area disturbed by human activities.
29) 301; 49.1298°,18.2972°; Biele Karpaty; Vieska-Bezdedov; Subsoil: claystone.
Pasture with lower grass-herbal vegetation cover.
30) 300; 49.1259°,18.2992°; Biele Karpaty; Vieska-Bezdedov; SE facing slope of a hill.
Subsoil: sandstone. Pinus sylvestris forest clearings with grass-herbal vegetation cover.
31) 422; 49.1248°,18.2893°; Biele Karpaty; Púchov; S to SE slope. Subsoil: limestone.
Abandoned overgrown quarry with stones on the surface, and small clearing.
32) 344; 49.1245°,18.3109°; Javorníky; Púchov; S facing slope. Subsoil: limestone.
Abandoned quarry with many stones, primarily overgrown with shrubs, but areas with
grass-herbal vegetation cover exist.
33) 277; 49.1218°,18.3035°; Biele Karpaty; Púchov; Subsoil: unknown. Old Acer
stands with much dead bark on their trunks.
34) 302; 49.1203°,18.3028°; Biele Karpaty; Púchov; Subsoil: unknown. Older apple
tree stands in a row, which are extensively managed.
35) 312; 49.1200°,18.2895°; Biele Karpaty; Streženice; Subsoil: unknown. Edge of
mixed forest.
36) 367; 49.1194°,18.2992°; Biele Karpaty; Púchov; S to SW facing slope of a hill.
Subsoil: sandstone. Surrounding approximately 350 meters long little used country road
going approximately in line with the hill’s ridge. There are edges of Pinus sylvestris forest
(including older stands) and southwards-situated originally steppe area affected by
succession process, afforestation, and forest development (Fig. 3A).
37) 324; 49.1193°,18.2940°; Biele Karpaty; Púchov; S facing slope of a hill. Subsoil:
limestone. Extensively managed grass-herbal area with fruit trees and forest ecotone
(Pinus sylvestris).
Entomofauna carpathica, 2024, 36(2): 83-126
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38) 320; 49.1193°,18.2919°; Biele Karpaty; Púchov; S facing slope of a hill. Subsoil:
limestone and sandstone. Mixed forest (also with some older Quercus stands),
surroundings of a little-used country road containing bare soil surfaces, and forest edge.
Area with stones on the ground surface.
39) 306; 49.1190°,18.2915°; Biele Karpaty; Púchov; Subsoil: unknown. Mowed grass area.
40)306; 49.1186°,18.3047°; Biele Karpaty; Púchov; Subsoil: unknown. Managed and
overgrown orchards.
41) 320; 49.1180°,18.3002°; Biele Karpaty; Púchov; S facing slope of a hill. Subsoil:
sandstone. Mixed forest with occasional clearings.
42) 297; 49.1176°,18.2935°; Biele Karpaty; Streženice; Subsoil: unknown. Garden
with young fruit trees and mowed grass area.
43) 280; 49.1162°,18.3012°; Biele Karpaty; Púchov; Subsoil: unknown. Little-used
country road and its surrounding containing mowed grass area, shrub growths, and dead
wood of old trees.
44) 276; 49.1160°,18.3007°; Biele Karpaty; Streženice; Subsoil: unknown.
Surrounding of a small stream where are young tree stands and shrubs.
Fig.
3
.
Biotopes at study sites.
A:
Forest
-
steppe in succession stage with lots of shrubs
and young forest stand. Photo: August 2017. B: Mostly bare surface of rocky soil –
habitat of Tetramorium moravicum. Photo: May 2018. C: Young Pinus sylvestris
forest and grass-herbal undergrowth on rocky soil – habitat of Lasius paralienus,
Ponera testacea, and Temnothorax interruptus. Photo: August 2018. D: Bare rocky
surfaces – habitat of Temnothorax nigriceps. Photo: August 2018.
Entomofauna carpathica, 2024, 36(2): 83-126
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45) 281; 49.1159°,18.3055°; Biele Karpaty; Púchov; S slope and E very steep slope.
Subsoil: limestone. Small forest-steppe area with xerothermophilous grass-herbal
vegetation, also with stones on the surface. Detected floral species are Corylus avellana,
Pinus sylvestris, Robinia pseudoacacia, and Ophrys apifera.
46) 427; 49.1155°,18.2784°; Biele Karpaty; Streženice; N facing slightly steep slope.
Subsoil: sandstone. Approximately 15-year-old mixed forest with much dead wood after
logging but also with remains of older spruce stands. Forest road for logging mechanisms
and tourists (Fig. 3A – the arrow).
47) 290; 49.1154°,18.3304°; Považské podolie; Púchov; Subsoil: unknown. Light Tilia
stands at a cemetery.
48) 479; 49.1139°,18.2613°; Biele Karpaty; Horná Breznica; Subsoil: unknown. Spruce
monoculture within an intensively managed forest.
49) 494; 49.1138°,18.2170°; Biele Karpaty; Lednica; S facing slope of a hill. Subsoil:
limestone. Stony biotope with much larger rocks, xerothermophilous grass-herbal
vegetation, and shrubs.
50) 274; 49.1132°,18.3053°; Biele Karpaty; Streženice; E very steep slope. Subsoil:
limestone. Forest-steppe with xerothermophilous grasslands and occasional stones on
the ground surface.
51) 264; 49.1128°,18.3058°; Považské podolie; Streženice; Subsoil: unknown. Small
area with grasslands and occasional fruit trees in proximity of a road.
52) 334; 49.1128°,18.3033°; Biele Karpaty; Streženice; S to SE slope of a hill. Subsoil:
limestone and sandstone. Overgrowing forest-steppe with many stones on the ground
surface. Small areas with xerothermophilous grasslands exist.
53) 466; 49.1123°,18.2163°; Biele Karpaty; Lednica; Subsoil: limestone. Pasture with
low and sparse xerothermophilous grass-herbal vegetation cover, shrubs, and occasional
stones on the ground surface.
54) 260; 49.1109°,18.3113°; Považské podolie; Streženice; Subsoil: unknown. Tilia
stands on a riverside.
55) 334; 49.1103°,18.3000°; Biele Karpaty; Streženice; SE slope of a hill. Subsoil:
limestone. Forest-steppe with many stones on the ground surface.
56) 302; 49.1099°,18.3027°; Biele Karpaty; Streženice; W slope of a hill. Subsoil:
limestone. Overgrowing forest-steppe with xerothermophilous grasslands partially
covered with rocky grounds. Partially afforested with Pinus nigra (Fig. 2C).
57) 369; 49.1091°,18.2861°; Biele Karpaty; Streženice; Subsoil: unknown. Wetland
with a flowing spring in non-forest.
58) 391; 49.1076°,18.2829°; Biele Karpaty; Streženice; Subsoil: unknown. Wetland
with a flowing spring in non-forest.
59) 358; 49.1071°,18.2854°; Biele Karpaty; Streženice; Subsoil: unknown. Small
stream and its surrounding deciduous stands which border with mowed grass areas.
60) 361; 49.1065°,18.2856°; Biele Karpaty; Streženice; Subsoil: unknown. Wetland in
non-forest.
61) 326; 49.1048°,18.2923°; Biele Karpaty; Streženice; SE slope of a hill. Subsoil:
claystone and limestone. Forest-steppe with stones on the ground surface and partially
with rocky soils. Dominant tree species is Pinus sylvestris. Juniperus communis is
Entomofauna carpathica, 2024, 36(2): 83-126
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abundantly present among the bushes. The site is partially influenced by rock mining in
the past, and a country road crosses it (Fig. 3B, C).
62) 267; 49.1042°,18.3065°; Považské podolie; Streženice; Subsoil: unknown. Older
concrete surface containing gaps with soil and vegetation humus, partially covered with
grass-herbal vegetation. Ruderal biotope disturbed by human activities (Fig. 2B).
63) 267; 49.1041°,18.3063°; Biele Karpaty; Streženice; Subsoil: unknown. Older
concrete surface containing gaps with soil and vegetation humus, partially covered with
grass-herbal vegetation. Also, side of a country road. Ruderal biotope disturbed by
human activities (Fig. 2B).
64) 393; 49.1027°,18.2995°; Biele Karpaty; Streženice; Subsoil: unknown. Extensively
managed pasture with older solitary trees.
65) 382; 49.0996°,18.2845°; Biele Karpaty; Streženice; S facing hillside with a fine
slope. Subsoil: limestone. Light deciduous forest with much rocks on the ground surface.
66) 433; 49.0979°,18.2975°; Biele Karpaty; Streženice; Subsoil: limestone. Abandoned
quarry with shrubs and much rocks on the ground surface.
67) 252; 49.0959°,18.3117°; Považské podolie; Streženice; Subsoil: unknown.
Riverside with non-forest vegetation, and gravel-sandy soils. Ruderal area disturbed by
flooding and human activities consisting of moving the soils.
68) 469; 49.0957°,18.2986°; Biele Karpaty; Streženice; Upper-situated area. Subsoil:
limestone. Top of a rocky hill with much of the rock on the surface and occasional
xerothermophilous grass-herbal vegetation.
69) 446; 49.0915°,18.2932°; Biele Karpaty; Horenice; S facing slope. Subsoil:
limestone. Abandoned quarry with many stones, partially overgrown with shrubs and trees,
and with the presence of xerothermophilous grass-herbal vegetation cover (Fig. 3D).
70) 408; 49.0902°,18.2790°; Biele Karpaty; Medné; SE facing slope of a small hill.
Subsoil: limestone. Stony place where is located a small clearing in surrounding of a
developing forest.
71) 320; 49.0871°,18.2904°; Biele Karpaty; Horenice; S facing steep hillside. Subsoil:
limestone. Small xerothermous area with grass-herbal vegetation cover in the vicinity of a forest.
72) 416; 49.0842°,18.2794°; Biele Karpaty; Medné; SE facing slope of a hill. Subsoil:
limestone. Stony place overgrown with bushes in surrounding of a developing forest.
73) 369; 49.0841°,18.2821°; Biele Karpaty; Medné; Subsoil: limestone. Area of a few
square meters, where are occasional above-ground larger stones, grass-herbal
vegetation cover, and shrubs.
Description of methods and material processing
The altitudinal data come from the map layer of relief model DMR 5.0 accessed
via the web GIS client (https://zbgis.skgeodesy.sk/); they refer to Baltic heights
after adjustment (1957). The positional coordinates (latitude,longitude) refer to
the EPSG:4326 coordinate reference system. Naming, classifying, and boundaries
of the geomorphological units (all kinds of units) follow KOČICKÝ & IVANIČ (2011).
The terms Biele Karpaty Mts, Javorníky Mts, and Považské podolie Val. are
considered as the Biele Karpaty geomorphological unit, the Považské podolie
Entomofauna carpathica, 2024, 36(2): 83-126
91
geomorphological unit, and the Javorníky geomorphological unit, respectively, in
whole paper.
The field study took place from 2014 to 2024. At the study sites, individuals
and nests were searched for by routine field inspection, which consisted of
browsing vegetation cover, vegetation humus, soil surfaces, tree trunks, stony
surfaces, rock crevices, and upper soil layer. To reach some poorly accessible
places, such as cavities in the rock, under bark, nest parts in stony ground, or soil
under deeply embedded stones, a geological hammer, and other equipment
were necessary to deploy (FI – field inspection). Individuals collected directly in
the field were moved into a transport container using a manual or electronic
exhaustor, tweezer, brush, or simply with hands. Litter sifting (LS) and pitfall
traps (PT) were also used to collect material. The sifted material was processed
to extract the individuals – it was browsed manually under light, or a Tullgren
funnel was used. Pitfall traps were exposed at three study sites during the
vegetation seasons – n. 36, n. 45, and n. 52 in year(s) 2018; 2018, 2021, 2022,
2023; and 2018, respectively. These traps were using containers which had input
diameter 3.5 cm, volume 40 ml, and cylindrical shape (years 2018, and 2021), or
containers which had input diameter 7 cm, and volume 300 ml (years 2022, and
2023). Fixing liquid consisted of a solution of 2% formaldehyde. All the study sites
were attempted to be investigated thoroughly, however, not the same effort was
put into all of them. Single findings outside them (the systematic survey was not
conducted at these places) that fall under the overall area of investigation may
also be included in the results. The determinations of specimens followed
primarily keys by SEIFERT (2018). Keys and descriptions of Kutter (1977, 1978),
SEIFERT (1983), CZECHOWSKI et al. (2012), SALATA & BOROWIEC (2013), and WAGNER
et al. (2017) were used too. All the determinations were conducted by the author
(some of them were reviewed). The used nomenclature follows SEIFERT (2018)
and other recent studies (CSŐZS et al. 2023, 2024, KIRSCHNER et al. 2023), which
do not contradict each other. Some of the specimens were integrated into the
author’s collection, and some of them were provided to other myrmecologists,
i.e., Dr. Bernhard Seifert, Petr Werner, and Dr. Adrián Purkart. Camera lens
Olympus M.ZUIKO DIGITAL ED 30mm f/3.5 was used to take Figs 3A, 5A–C, 7B, C,
8, 9B, 11B, C. Figs. 5D, 6B–D, 7D, 9C, D, 10, 11A, D, 12 were taken using a super
macro camera lens Yasuhara Nanoha x5. Figs 5D, 6B–D, 7D, 9C, D, 12 were focus-
stacked using Helicon Focus software (version 8.1.0) from focus-bracketed
photos taken with that lens and Olympus STF-8 macro flash. Multiple pictures
composed into a single figure are ordered chronologically from right to left,
based on the dates of their creation. All photos were taken by the author.
For each species, faunistic data gathered during the survey are provided
(Data), facultatively are present the additional data (Additional finding(s)) and
the main notes. The Data contain chronological records which are ordered and
Entomofauna carpathica, 2024, 36(2): 83-126
92
structured as follows: date of collection (format “DD.MM.YY”), ID of site,
collecting method, number of captured individuals for each caste (w(w) –
worker(s), g(g) – gyne(s), and m(m) – male(s)), and facultatively a short note. If
the method is individual one (FI), then the collection is from a single ant colony,
if it is not mentioned otherwise. Moreover, the FI collection originates directly
from a single nest or close nest(s)’s surroundings, if it did not contain only sexual
individuals. The short note can be of various character, e.g., a note regarding
determination or a note on the circumstances of the finding. The Additional
findings are structured similarly to the Data, except for sites that do not exist, so
there are listed coordinates and geomorphological units instead.
RESULTS AND DISCUSSION
Dolichoderinae
Bothriomyrmex corsicus Santschi, 1923
Data: 05.06.14, 52, FI, 110 ww, colony nesting in rocky soil; 12.06.18–30.07.18, 45, PT, 1 g.
Dolichoderus quadripunctatus (Linnaeus, 1771)
Data: 17.11.18, 37, FI, 1 g, nested and establishing a new colony (without workers);
05.05.19, 59, FI, 20 ww.
Tapinoma erraticum (Latreille, 1798)
Data: 14.08.17, 38, FI, 100 ww, 1 g; 15.03.18, 36, FI, 10 ww; 07.08.18, 27, FI, 1 w;
11.08.18, 68, FI, 3 ww, unknown whether from a single colony; 19.08.18, 22, FI, 1 w;
30.08.18, 32, FI, 5 ww; 16.09.18, 50, FI, 4 ww, unknown whether from a single colony;
18.04.19, 61, FI, 1 w; 21.04.19, 16, FI, 2 ww, unknown whether from a single colony;
16.07.19, 4, FI, 5 ww; 23.07.19, 29, FI, 1 w; 06.08.19, 20, FI, 3 ww, 5 gg; 17.05.20, 14, FI,
1 w; 15.07.20, 19, FI, 1 w; 28.06.21, 18, FI, 10 ww; 13.08.21, 15, FI, 4 ww, 1 g; 14.06.21–
22.08.21, 45, PT, 15 ww; 02.10.21, 6, FI, 1 w; 25.06.22, 10, FI, 5 ww; 28.06.22, 2, FI, 1 w;
02.07.22, 66, FI, 20 ww; 05.07.23, 21, FI, 1 w; 26.05.24, 42, FI, 1 g, after dispersal.
Tapinoma subboreale Seifert, 2011
Data: 16.07.19, 4, FI, 2 gg, unknown whether from a single colony, after dispersals (?).
The gynes were found during a single visit at the site; they were actively
moving on the ground surface, and both were found up to 5 metres from each
other. Their small abdomens and the late date when they were found compared
with the known swarming period (SEIFERT 2018) suggest that they were not found
immediately after their nuptial flights. Most likely, they were already establishing
new colonies and left their mother chambers due to a lack of nutrition or
Entomofauna carpathica, 2024, 36(2): 83-126
93
disturbances. Such finding suggests the occurrence of their mother
colony/colonies at the site. Targeted attempts to find a nest or workers failed;
only the T. erraticum was found there.
Formicinae
Camponotus aethiops (Latreille, 1798)
Data: 27.07.17, 52, FI, 5 ww, 2 mm; 27.08.18, 61, FI, 1 w; 17.03.19, 52, FI, 5 ww; 13.04.19,
52, FI, 10 ww; 18.04.19, 61, FI, 5 ww; 21.04.20, 16, FI, 2 ww; 06.08.20, 20, FI, 1 w;
18.08.20, 18, FI, 3 ww, 3 gg; 03.03.21, 56, FI, 2 ww; 28.06.21, 56, FI, 30 ww; 26.03.22, 71,
FI, 8 ww; 17.06.22, 46, FI, 3 ww; 25.06.22, 10, FI, 3 ww; 26.06.22, 13, FI, 1 w.
Camponotus fallax (Nylander, 1856)
Data: 25.04.20, 59, FI, 10 ww, 1 g, 1 m; 23.06.23, 37, FI, 20 ww.
Camponotus herculeanus (Linnaeus, 1758)
Data: 24.08.17, 46, FI, 1 w; 04.10.17, 45, FI, 1 w; 03.07.18, 46, FI, 11 gg, unknown whether
from a single colony, after dispersal; 14.08.21, 46, FI, 7 ww, unknown whether from a
single colony.
Camponotus ligniperda (Latreille, 1802)
Data: 22.08.17, 36, FI, 4 ww; 08.07.18, 52, FI, 20 ww; 30.03.19, 14, FI, 1 w; 08.09.19, 38,
FI, 15 ww; 14.09.19, 5, FI, 2 ww; 10.09.21, 61, FI, 4 ww; 25.06.22, 10, FI, 7 ww; 05.05.23,
37, FI, 1 w; 18.05.23, 13, FI, 3 ww.
Camponotus piceus (Leach, 1825)
Data: 12.04.18, 45, FI, 4 ww; 23.05.18, 61, FI, 2 ww, unknown whether from a single
colony; 11.08.18, 68, FI, 7 ww, 1 m; 19.08.18, 22, FI, 2 ww; 14.04.19, 45, FI, 5 ww, 4 mm;
18.04.19, 61, FI, 2 ww, unknown whether from a single colony; 21.04.19, 16, FI, 10 ww;
06.08.19, 20, FI, 1 w; 17.08.19, 20, FI, 5 ww, 3 gg; 14.09.19, 5, FI, 2 ww; 09.05.20, 56, FI,
10 ww; 15.07.20, 19, FI, 1 ww; 18.08.20, 18, FI, 1 w; 28.06.21, 18, FI, 1 w; 26.03.21, 71,
FI, 11 ww; 29.05.21, 10, FI, 1 ww; 26.06.22, 13, FI, 1 ww; 05.07.23, 21, FI, 1 w.
The site n. 5 represents the northernmost known distributional point so far in
the Slovak Republic (ZÁLESKÝ 1939, BELÁKOVÁ 1956, DRDULOVÁ & ZLATOŠOVÁ 1980,
KOŽÍŠEK 1986, KOŽÍŠEK 1989, DRDULOVÁ 1991, BEREC 1997, AMBROS et al. 1998,
DEVÁN 2005, DEVÁN 2006b, WIEZIK 2007, DEVÁN 2008a, b, c, WIEZIK 2008a, c, WIEZIK
et al. 2010, WIEZIK & WIEZIKOVÁ 2012, SUVÁK 2021, JANCÍK & PURKART 2024, MARKO
et al. 2024, JANCÍK, PURKART, unpub. data).
Entomofauna carpathica, 2024, 36(2): 83-126
94
Camponotus vagus (Scopoli, 1763)
Data: 19.08.18, 46, FI, 1 w; 21.04.19, 16, FI, 1 w, 2 gg; 08.06.21, 43, FI, 30 ww; 09.06.21,
43, FI, 7 ww; 14.08.21, 46, FI, 35 ww; 14.08.21, 46, FI, 10 ww.
Four colonies were discovered at the site n. 46. A unique situation occurred
there – a colony of this thermophilous species and a colony of C. herculeanus
were nesting in proximity of tens meters. It is an example of the penetration of
a thermophilous species into the mountains of Biele Karpaty Mts, where it met
with an oligothermic one. Such a situation is characteristic for hills in Biele
Karpaty Mts and was already understood by BEZDĚČKA (1992).
Colobopsis truncata (Spinola, 1808)
Data: 18.11.18, 37, FI, 20 ww; 17.03.19, 36, FI, 10 ww, 1 g; 16.04.20, 59, FI, 10 ww.
Formica cinerea Mayr, 1853
Data: 24.03.19, 67, FI, 45 ww.
Formica cunicularia Latreille, 1798
Data: 06.05.22, 42, FI ,30 ww; 06.07.22, 11, FI, 4 ww.
Formica exsecta Nylander, 1846
Data: 16.07.17, 39, FI, 1 g, after dispersal, fully winged, at 14:07h.
The gyne was transported into the laboratory to potentially start its
laboratory keeping (Fig. 4). Attempts to find colonies of this species were
unsuccessful. Suitable micro-localities with extensive pasture existed a few
hundred meters away. Unfortunately, these were private and inaccessible plots
of land. The gyne’s mother nest probably occurred within either these sites or
within the broader area of the Biele Karpaty Mts. Populations of this species
declined across its distributional range during the past decades due to intensified
land use (SIEFERT 2018). The second known largest polydomous supercolony in
the Europe was found in the Slovak Republic (WIEZIK et al. 2017). Several sites
with confirmed occurrences are known so far within the Biele Karpaty Mts –
48.8949°,17.7991° (Natural Monument Grúň), 49.0399°,18.1557° (Natural
Reserve Drieňová), 48.9698°,17.9486° (Natural Monument Včelíny),
48.8189°,17.4398° (Natural Monument Bučkova Jama), and 49.0588°,18.1369°
(Natural Monument Krivoklátske lúky) (DEVÁN 2009c).
Formica fusca Linnaeus, 1758
Data: 11.04.20, 58, FI, 3 ww; 01.05.21, 36, FI, 3 ww; 05.11.21, 72, FI, 10 ww.
Entomofauna carpathica, 2024, 36(2): 83-126
95
Formica gagates Latreille, 1798
Data: 06.04.19, 36, FI, 40 ww; 21.03.20, 36, FI, 8 ww; 22.03.20, 37, FI, 80 ww; 03.03.21,
56, FI, 2 ww; 06.04.21, 37, FI, 40 ww; 14.06.21–22.08.21, 45, FI, 15 ww; 07.07.22, 45, FI,
4 ww, 4 gg, 6 mm; 17.04.23, 38, FI, 4 ww.
Discovered nests had always above-ground mounds consisting of soil particles
and dry pieces of herbs. The mounds had diameters of its bases from 25 to 40
cm and heights from 10 to 25 cm. Some of the colonies also had their chambers
located under stones. Numerous records are available from the Slovak Republic
(PETRICSKÓ 1892, MOCSÁRY 1897, CHYZER 1902, ZÁLESKÝ 1939, SADIL 1953, BELÁKOVÁ
1956, 1961, LABUDA 1970, DRDULOVÁ 1979, 1991, KOŽÍŠEK 1986, RANDUŠKA 1995,
AMBROS et al. 1998, HOLECOVÁ et al. 2003, DEVÁN 2005, 2006a, b, c, 2007, 2008a,
c, 2009a, b, WIEZIK 2007, 2008a, b, c, WIEZIK & WIEZIKOVÁ 2007, 2012, DEVÁN 2008,
MAJZLAN & DEVÁN 2009, WIEZIK et al. 2010, PURKART 2016, 2017, 2018, 2020,
2023a, ČERVENÁ et al., 2018, ČERVENÁ et al. 2020, PAVLÍKOVÁ 2020, SUVÁK 2021,
MARKO et al. 2024). The northernmost known distributional point of this species
Fig.
4
.
The gyne of
Formica
exsecta
in an interaction with a callow “host” worker of
Formica rufibarbis after they were put together into a keeping facility. The gyne
carries subgenus-specific notched vertex of head, what is in contrast with the
worker which it has roundly shaped. Photo: July 2017.
Entomofauna carpathica, 2024, 36(2): 83-126
96
in the Slovak Republic was near the town Púchov (LABUDA 1970) at 49.13°
(latitude), however, it was not considered by SEIFERT (2018), who mentioned the
northernmost sites at “48.2°N”. This species was not confirmed for the Labuda’s
site during the time of this survey, and most probably has disappeared from
there due to the habitat change.
Formica polyctena Förster, 1850
Data: 15.07.18, 48, FI, 30 ww, polydomous colony with at least 15 nests; 16.04.19, 31, FI,
60 ww.
Formica pratensis Retzius, 1783
Data: 22.08.17, 36, FI, 6 ww, 2 mm; 13.01.18, 50, FI, 10 ww; 25.01.18, 36, FI, 11 ww.
Formica rufa Linnaeus, 1761
Data: 30.04.21, 40, FI, 90 ww.
Formica rufibarbis Fabricius, 1793
Data: 30.03.18, 42, FI, 2 ww; 24.04.21, 36, FI, 20 ww.
Formica sanguinea Latreille, 1798
Data: 24.08.17, 46, FI, 20 ww; 06.07.18, 36, FI, 10 ww.
Formica truncorum Fabricius, 1804
Data: 23.07.17, 38, FI, 1 g, after dispersal, wingless; 25.05.18, 43, FI, 1 g, after
dispersal, wingless, at 17:45h; 27.05.18, 43, FI, 1 g, after dispersal, wingless;
19.07.20, 3, FI, 6 ww; 14.08.21, 46, FI, 60 ww, 1 m.
The gyne, which was found as the first one at the site n. 43, was put under
laboratory conditions to simulate the natural process of establishing a new
colony of this temporally socially parasitic species. It was assigned a few dozen
pupae of Formica fusca workers from a foreign colony and fed by sugar water
(JANCÍK & DISNEY 2020). Some of the pupae were eaten by the gyne a few days
after installation of the laboratory keeping (JANCÍK & DISNEY 2020). This
observation represented unusual behaviour, compared with a few (4) similar
situations when the colony-establishing gynes did not eat the pupae (JANCÍK,
unpub. data). The keeping revealed that the gyne was infested by a brood of a
parasitic fly species. These flies in the developmental stage of larva were eating
nutrients from the inner of the gyne, possibly explaining the eating of the pupae.
These larvae emerged from the gyne (from an unknown body part), pupated
quickly, and later reached the adult stage. Adults were taken from the keeping
Entomofauna carpathica, 2024, 36(2): 83-126
97
facility and subsequently investigated by R. H. L. Disney, resulting in description
of the species Aenigmatias exreginae Jancík & Disney, 2020 (Diptera, Phoridae)
(JANCÍK & DISNEY 2020). The gyne laid its first egg on June 24, 2018, but another
freshly emerged larva of the parasitic fly appeared on June 27. Therefore, the
ability to lay eggs was not prevented by the presence of the parasite in the gyne’s
body. The gyne survived this infestation, was even able to lay eggs, and
successfully created first generation of workers.
Lasius alienus (Förster, 1850)
Data: 11.03.18, 36, FI, 8 ww; 08.07.18, 52, FI, 5 ww; 14.10.18, 14, FI, 10 ww; 20.10.18,
14, FI, 20 ww; 16.03.19, 52, FI, 8 ww; 21.04.19, 16, FI, 14 ww; 22.04.19, 17, FI, 5 ww;
10.03.20, 32, FI, 10 ww; 15.08.20, 68, FI, 10 ww; 15.09.20, 9, FI, 5 ww, 5 mm; 24.04.21,
36, FI, 30 ww; 30.06.21, 52, FI, 20 ww; 13.08.21, 15, FI, 10 ww, 2 mm; 03.10.21, 36, FI, 30
ww; 07.08.22, 52, FI, 1 g, after dispersal and nested.
Lasius bicornis (Förster, 1850)
Data: 09.05.16, 42, FI, 1 g, after dispersal, fully winged, at 9:26h; 11.05.21, 42, FI, 1 g,
after dispersal, wingless, at 9:02h; 01.05.24, 35, FI, 1 g, after dispersal, fully winged, at
9:21h.
This species is little known (SEIFERT 2018). There are few findings from the
Slovak Republic, most of them refer to catches of alates after their dispersals
(MOCSÁRY 1897, CHYZER 1902, https://www.antweb.org – specimen
CASENT0280461, KOŽÍŠEK 1985a, 1989, DEVÁN 2006b, WIEZIK 2008b, KLESNIAKOVÁ
et al. 2016, PAVLÍKOVÁ 2020, JANCÍK, PURKART, unpub. data).
Species was found three times; each time, they were gynes after dispersals.
Targeted attempts to find nests or workers failed. When a suitable habitat was
found, only different ant species were discovered in proximity to trees – L.
brunneus, L. fuliginosus, or L. umbratus, or even no ant species of that genus.
Findings of single gynes at sites n. 42 (in years 2016, and 2021), and n. 35 (in the
year 2024) do not confirm this species for these sites in the sense of colony
occurrence. The nests or workers were not found at these sites after thorough
searches, therefore, their mother colonies (or single colony) probably existed
outside them. The sites’ surroundings of diameter 1 km included many older
solitary trees (mostly fruit ones) located within private gardens, representing
suitable habitats for this species (SEIFERT 2018). The gynes probably dispersed
from there, or from more distant places within the Biele Karpaty Mts.
Laboratory keeping was used to find out more about the biology of this cryptic
species. The gynes collected in the years 2021 and 2024 were used for this
purpose. Since having gynes of socially parasitic species at the time immediately
after their dispersals, the most complicated activities during the laboratory
keeping are required at its beginning – establishing of a new colony. This involves
Entomofauna carpathica, 2024, 36(2): 83-126
98
establishing a friendly relationship between the gyne and host colony members.
It is unknown what host species is used in the nature, but L. brunneus is the
presumed host because it shares the same habitat (SEIFERT 2018). It is also
unknown whether the parasitic gyne is capable of taking over the queen-right
host colony, or only orphaned colony (GIEHR et al. 2019).
The gyne found in 2021 (Fig. 5A, D) was assigned 15 adult workers originated
from a queen-right colony of L. brunneus. Establishing the relationship was not
successful, and the gyne suffered much harm from non-kin workers. Therefore,
the laboratory keeping was terminated.
Keeping of the gyne found in 2024 firstly required signs of its fertility since it
was found as a fully winged individual, indicating that it may not have been
mated, and if this was true, then the keeping would be irrelevant. It was offered
with sugar water, and the gyne accepted it immediately after it was placed in the
laboratory conditions. To find out the fertility signs, the gyne was put on an open
surface and exposed to strong light. It was not trying to fly off, instead it was
trying to put off its wings using the hind legs. This was the sign that it was mated,
so that the keeping started. Gyne needed the host workers, but to avoid the
mistake of 2021, these workers had to be freshly hatched when they lacked
“loyalty” to their mother colony. Based on the experience of ant-keeping
hobbyists, such workers, combined with pupae of other workers from the same
colony, are considered an ideal mixture to help the gyne smoothly establish its
own colony. This material was not available immediately after the gyne was
incorporated in the laboratory. Therefore, it was supplied with Lasius brunneus
adult (not freshly hatched) workers from a foreign queen-right colony. Only four
individuals were used, fewer than in 2021, and they were added to the gyne one
by one at hourly intervals. The gyne was attacked by these workers, and it lost
part of its funiculus. When one of the four workers was bitten into the gyne for
more than 10 seconds, the gyne simply killed the worker by its mandibles (one
worker was added afterward). Gyne was observed to antennate a worker’s body
surface, and then it rubbed its antennae with forelegs, which subsequently
rubbed with mandibles, and the thorax. On May 2, 2024 – the second day in the
laboratory, the gyne appeared to be stable, located in proximity of the “host”
workers, and all the individuals seemed to be calm. The activity of adding the
four workers aimed to increase the chance that the gyne survive until a freshly
hatched workers and pupae were available. It is likely that gynes of most of the
Chthonolasius species – which do not use the low-temperature strategy for the
colony penetration (SEIFERT 2018) – die within few days if they fail to get workers
which would take care about them since their body reserves are poor. This
behaviour was observed for Lasius umbratus (Nylander, 1846) (JANCÍK, unpub.
data). The gyne was fully winged on May 2, 2024; its attempts to put off wings
did not lead to success. That day, they were put off manually because the gyne
Entomofauna carpathica, 2024, 36(2): 83-126
99
could stick to the surface within the keeping facility, which could potentially
cause its death. Freshly hatched workers and pupae from a foreign colony of L.
brunneus (Poland) were available on May 9, 2024, and were continuously added
to the current keeping from that day till May 14, 2024, when the laboratory
keeping contained the gyne, approximately 100 adult workers, and about 250 pupae.
Gyne was observed to produce one of its first eggs on May 20, 2024. The new
colony contained at least 300 eggs on May 31, 2024. That time, the queen was
physogastric during the period of depositing its eggs, what can be well compared
with the other gyne, at a time after dispersal (Fig. 5A, B). Unexpectedly, the
queen died between June 4, and June 5 as strongly physogastric (Fig. 5C). It was
found motionless in a chamber of the artificial nest, being in proximity of host
workers which were taking care of it. The cause of its death was unknown;
generally, probably physogastric gynes are susceptible to any negative influences
that are causing their deaths. One similar case was observed with queen of
Fig.
5
.
Two gynes of
Lasius
bicornis
–
A, D:
gyne found in the year 2021,
B, C:
gyne
found in the year 2024. A: Wingless gyne with small abdomen, being after
dispersal, is drinking given sugar water. Photo: May 2021. B: Gyne in proximity of
workers of Lasius brunneus in laboratory keeping. Photo: May 2024. C:
Physogastric gyne several hours after its dead. Photo: June 2024. D: Frontal view
of petiolus showing species-specific signs. Scale bar: 0.4 mm. Photo: August 2024.
Entomofauna carpathica, 2024, 36(2): 83-126
100
Anergates atratulus (Schenck, 1852) (PURKART 2023b). The laboratory keeping
was not terminated by this incident, but the queen was extracted and put into
ethanol. Other colony members remained in the keeping facility, aimed to
continue with the development of the first generation of L. bicornis individuals
from the eggs. Colony contained exclusively larvae within all the brood on July 7,
2024. Workers were fed by sugar water, and proteins ad libidum from the May
9, 2024. Temperature conditions varied from 20°C to 30°C and mostly copied
natural ones in place of finding of the gyne, during all the time of laboratory
keeping. Disappearing of larvae and their slow growth was observed from the
July 7 to August 11. No pupae were created till that day. The host workers
continued to take care of this offspring after the queen died – fed it and
protected it when there was any disturbance to the nest, as was observed during
regular daily checks. The reason for either the defect development of the larvae
or less-probably its killing by the workers (not observed) appears unknown. The
laboratory keeping was terminated on August 18, 2024, containing only
approximately 40 middle-sized larvae among the brood.
Lasius brunneus (Latreille, 1798)
Data: 01.08.17, 38, FI, 10 ww; 08.07.18, 52, FI, 5 ww; 26.06.22, 12, FI, 40 ww; 13.05.23,
37, FI, 5 ww; 01.05.24, 35, FI, 10 ww.
Lasius citrinus Emery, 1922
Data: 26.05.19, 42, FI, 1 g, after dispersal, fully winged, at 8:16h
Attempts to find nests or workers at study sites failed. The gyne was dispersed
from its mother nest, which was probably located within the Biele Karpaty Mts,
and probably not at the site where it was found. Known records from the Slovak
Republic were partly summarised by MARKO et al. 2024, additional data were
recorded by ZÁLESKÝ (1939), BEZDEČKA (1996a), DEVÁN (2006a, b, 2007, 2008a, b),
MAJZLAN & DEVÁN (2009), WIEZIK & WIEZIKOVÁ (2010), PAVLÍKOVÁ (2020), and
PURKART (2023a).
Lasius emarginatus (Oliver, 1792)
Data: 17.03.18, 52, FI, 20 ww; 17.03.18, 52, FI, 20 ww; 10.09.20, 36, FI, 5 ww.
Lasius flavus (Fabricius, 1782)
Data: 27.08.18, 61, FI, 5 ww; 14.04.19, 45, FI, 10 ww; 22.04.19, 17, FI, 5 ww; 12.05.20, 36, FI, 4 ww;
12.06.22, 42, FI, 4 ww; 28.06.22, 2, FI, 22 ww.
Lasius fuliginosus (Latreille, 1798)
Data: 26.09.17, 38, FI, 5 ww.
Entomofauna carpathica, 2024, 36(2): 83-126
101
Lasius jensi Seifert, 1982
Data: 03.07.18, 46, FI, 1 g, after dispersal, wingless; 12.06.18–30.07.18, 36, PT, 2 gg;
14.06.21–26.07.21, 45, PT, 3 gg; 06.08.21, 36, FI, 1 g, after dispersal; 14.06.21–23.08.21,
45, PT, 1 g; 10.07.23, 51, FI, 1 g, after dispersal.
Targeted attempts to find a nest or workers (mainly on site n. 36) failed.
Findings of 3 and 4 gynes after dispersals at sites n. 36, and n. 45, respectively,
probably point to existence of their mother nests at these sites. This probability
is increased by the fact that there were collected at least two gynes in a single
pitfall trap at the latter site. The very first finding at the site n. 46 probably refers
to dispersal from a more distant place (e.g., site n. 36) since no typical habitat
existed there. Finding at the site n. 51 can refer to the existence of mother colony
at the neighbour site n. 50, concluding from the habitats. Data of this study
suggest this species to be the only Chthonolasius Ruzsky, 1912 species which
inhabited xerothermophilous grasslands within the investigated area. This
species was reported from a certain site within the Biele Karpaty Mts only once,
excluding this study (BEREC 1997). Also, available records show that it was rarely
found within the Slovak Republic in the past (BEREC 1997, WIEZIK 2008a,
KLESNIAKOVÁ et al. 2016, PURKART 2016, PURKART 2020, 2023a, SUVÁK 2021, MARKO
et al. 2024). However, more findings (and more localities) are known, mostly
based on gynes after dispersals, from recent years (JANCÍK, PURKART, unpub. data).
The site n. 36 represented the northernmost known distributional point of this
species within the Slovak Republic.
Lasius mixtus (Nylander, 1846)
Data: 12.11.20, 40, FI, 7 ww; 02.04.21, 40, FI, 2 ww; 29.04.21, 40, FI, 20; 30.04.21, 40, FI,
20 ww, 19.09.24, 35, FI, 1 g, after dispersal, fully winged, at 10:20 h.
The gyne was found on the second day after a period of cold and rainy
weather had ended, what corresponds with data known for this species (SEIFERT,
2018).
Lasius myops Forel, 1894
Data: 04.06.22, 52, FI, 300 ww; 12.06.22, 52, FI, 200 ww; 18.07.22, 52, FI, 20 ww;
18.07.22, 52, FI, 2 mm, males were pupae; 18.07.22, 52, FI, 3 gg, gynes were pupae;
18.07.22, 52, FI, 125 mm, males were pupae; 18.07.22, 52, FI, 140 mm, males were
pupae; 27.08.22, 52, FI, 50 ww, 18 mm; 09.09.23, 52, FI, 24 ww, 4 gg, 2 mm.
All specimens originated from a single colony; if they were extracted as
pupae, then they were put under laboratory conditions to develop into the adult
stage. This species can be correctly determined by even 2 workers (SEIFERT 1983).
The investigated colony was determined to species using each caste separately
Entomofauna carpathica, 2024, 36(2): 83-126
102
– based on more than 30 workers, 2 gynes, and 30 males (Fig. 6), to maximize
the reliability of the determination. Adult alates were present in the colony at
the typical time following the known data (SEIFERT 2018). Colony members were
extracted directly from the soil where they were nesting near a rock and in the
rock crevices.
This species is rarely found in xerothermic places within the Central Europe
(SEIFERT 2018). Also, it was rather rarely recorded within the Slovak Republic in
the past (LABUDA 1970, KOŽÍŠEK 1986, DEVÁN 2006d (?), 2008, SUVÁK 2021, JANCÍK,
PURKART, unpub. data). The site n. 52 represented the northernmost known
distributional point of this species within the Slovak Republic. The record of
DEVÁN (2006d) referring to a single worker could not be enough for precise
determination (SEIFERT 1983). Therefore, it is considered unreliable, as far the
specimen could be also the sister species – L. flavus. Furthermore, there was a
finding of a single worker at the site n. 52 already on July 08, 2018. This worker
Fig. 6. Colony of Lasius myops. A: Nesting place (Váh river further in the photo).
Photo: June 2022. B: Lateral view of gyne’s head, showing yellowish coloration on
its hind beyond eye. Scale bar: 0.7 mm. Photo: August 2024. C: Mandibles of male
– masticatory borders carry apical and subapical (not highlighted) teeth and
smaller teeth (highlighted). Scale bar: 0.2 mm. Photo: August 2024. D: Lateral
view of a medium-to-big-sized worker. Scale bar: 1.0 mm. Photo: August 2024.
Entomofauna carpathica, 2024, 36(2): 83-126
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had signs of L. myops, but it was insufficient to confirm the species. Based on
that record, subsequent thorough search for additional individuals (and the
whole colony) was successful.
Lasius niger (Linnaeus, 1758)
Data: 16.07.17, 39, FI, 5 ww; 14.03.18, 44, FI, 25 ww; 05.04.21, 60, FI, 3 ww; 16.08.21,
23, FI, 5 ww; 05.11.21, 73, FI, 7 ww; 26.05.24, 62, FI, 10 ww.
Lasius paralienus Seifert, 1992
Data: 02.07.18, 61, FI, 3 ww; 27.18.18, 61, FI, 10 ww, 1 g, 2 mm; 22.09.18, 36, FI, 14 ww;
30.10.18, 61, FI, 60 ww, 1 g; 06.03.19, 55, FI, 10 ww; 07.03.19, 61, FI, 5 ww; 18.04.19, 61,
FI, 10 ww; 17.03.20, 61, FI, 20 ww; 15.08.20, 36, FI, 120 ww; 10.09.20, 36, FI, 5 ww, 55
mm; 11.09.20, 20, FI, 10 ww; 21.04.21, 36, FI, 10 ww; 28.06.21, 18, FI, 8 ww; 28.06.21,
56, FI, 60 ww; 23.07.21, 30, FI, 8 ww; 02.10.21, 6, FI, 20 ww; 29.05.22, 10, FI, 4 ww;
26.06.22, 13, FI, 5 ww; 07.07.22, 53, FI, 7 ww.
Lasius platythorax Seifert, 1991
Data: 24.06.15, 25, FI, 10 ww; 15.07.18, 48, FI, 15 ww; 25.08.19, 10, FI, 1 g, after dispersal
and nested (without workers); 31.03.21, 36, FI, 2 ww; 29.04.21, 56, FI, 90 ww; 14.06.21–
22.08.21, 45, PT, 15 ww; 05.11.21, 72, FI, 20 ww.
Lasius reginae Faber, 1967
Data: 14.08.17, 36, FI, 10 ww, 8 gg; 28.06.18, 36, FI, 10 ww; 28.08.18, 36, FI, 5 ww, 2 gg;
26.09.18, 36, FI, 10 ww; 27.09.18, 36, FI, 11 ww; 05.10.18, 36, FI, 17 ww; 06.10.18, 36, FI,
6 ww; 08.10.18, 36, FI, 12 ww, 3 gg, 1 m; 09.10.18, 36, FI, 3 ww; 17.10.18, 36, FI, 7 ww,
30 gg, 40 mm.
This species was so far reported three times in the Slovak Republic (WERNER
1978, KOŽÍŠEK 1985b, 1987). One colony was found during this survey (Figs. 7, 8)
at site n. 36 which currently represents the northernmost known distributional
point of this species in this country. The nest was discovered on August 14, 2017,
on the margin of the country road. There it was situated near a large root of a
living Pinus sylvestris tree in a forest ecotone, and was surrounded by low and
sparse xerothermophilous grass-herbal vegetation. It had small inconsistent
above-ground mound consisting of soil particles, up to 10 cm of high, and
diameter of up to 30 cm. The workers were taking care about subterranean root-
sap suckers as was observed when the nest was inspected internally. This species
and its sibling one (Lasius carniolicus Mayr, 1861) can be easily distinguished
from the other yellowish Lasius Fabricius, 1804 spp. with no magnification, even
in the field. Gynes have very small body sizes, approximately the same as
workers’ (Fig. 7C). Also, this species is well-distinguishable by its strong citronella
odor released by the colony members when they are endangered. This odor is
Entomofauna carpathica, 2024, 36(2): 83-126
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exceptionally strong and unmistakable with any secretion of other yellowish
Lasius spp. The workers of these two sister species (subgenus Austrolasius Faber,
1967) have typically roundly shaped head sides over mandibles (Fig. 7D). Sexuals
were flying off from stems near the nest during swarming (Fig. 7A); they flew off
in the direction of the sun and were never found near the nest up to 30 meters
after the swarming, even despite their huge number. Flew-offs were observed
on September 26, 2017, at 15:11h–17:01h, and on September 16, 2019, at
15:27h–17:27h. The sexuals were many times observed to be only situated on
the ground surface of the nest, or near the surface, but the swarming did not
occur during that day. Meteorological data were not obtained from the nearest
meteorological station Beluša for these dates, because the station had outages
at those times (SHMÚ pers. comm.). The maximum year-round reproduction
capacity was estimated to be at least 5,000 sexuals – based on their numbers
observed when they were flying off. Freshly mated and dispersed gynes are
Fig. 7. Colony of Lasius reginae. A: Accumulation of gynes on a dry stem near their
nest, before they will fly off to swarming. Photo: September 2017. B: Overall view
of a worker. Photo: July 2022. C: Gyne and worker showing their similar body
sizes. Photo: September 2022. D: Frontal view of worker’s head showing
subgenus-specific and species-specific signs. Scale bar: 0.4 mm. Photo: August
2024.
Entomofauna carpathica, 2024, 36(2): 83-126
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trying to find adoption in Lasius alienus colonies (SEIFERT 2018), as this species is
a temporal social parasite. Colonies of L. alienus and sibling L. paralienus were
nesting in proximity of the nest up to 20 metres.
Lasius sabularum (Bondroit, 1918)
Data: 18.04.21, 65, FI, 1 w; 28.04.21, 41, FI, 50 ww, rev. Seifert, under a stone and close
to the base of an Acer tree; 14.06.21, 41, FI, 9 ww; 29.06.21, 65, FI, 20 ww, under a stone;
14.08.21, 41, FI, 20 ww; 29.06.23, 28, FI, 40 ww, under a stone and close to the base of a tree.
Additional finding: 26.03.22, 49.0846°,18.2151°, Biele Karpaty Mts, FI, 30 ww, in tunnels
in soil of a ground-surpassing mound.
Based on available data, this species was discovered for the first in the Slovak
Republic by Bezděčka (SEIFERT 1988), other published records come from JANCÍK
& PURKART (2024), and MARKO et al. (2024). This survey revealed its permanent
presence (colony existence) directly (colony findings) at all four study sites. Fig. 9.
Fig. 8. Colony of Lasius reginae in nest after removing a nest-covering stone; workers
are taking care about pupae and larvae. Photo: June 2020.
Entomofauna carpathica, 2024, 36(2): 83-126
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Lasius umbratus (Nylander, 1846)
Data: 18.10.18, 37, FI, 10 ww; 23.03.19, 38, FI, 10 ww; 18.04.19, 61, FI, 10 ww; 19.04.19,
45, FI, 9 ww; 09.11.20, 36, FI, 60, morpha Umbrata and morpha Compacta, rev. Seifert;
01.04.21, 36, FI, 123 ww; 10.04.21, 36, FI, 32 ww, morpha Umbrata and morpha
Compacta, rev. Seifert; 11.04.21, 36, FI, 60 ww, morpha Umbrata and morpha Compacta;
27.05.21, 36, FI, 40 ww; morpha Umbrata and morpha Compacta; 12.06.21, 36, FI, 600
ww, morpha Umbrata and morpha Compacta; 15.07.21, 36, FI, 20 ww, 20 gg; 01.11.21,
56, FI, 10 ww; 05.11.21, 73, FI, 20 ww; 17.06.22, 49, FI, 200 ww; 25.06.22, 10, FI, 23 ww,
morpha Umbrata and morpha Compacta; 28.06.22, 36, FI, 16 ww, morpha Umbrata and
morpha Compacta.
Polyergus rufescens (Latreille, 1798)
Data: 23.07.19, 29, FI, 20 ww; 24.07.22, 56, FI, 3 ww, 8 gg, 5 mm, including 2 ergatogynes;
05.09.22, 36, FI, 5 ww; 06.09.22, 56, FI, 7 ww; 30.06.23, 8, FI, 20 ww.
Fig. 9.
Colony of
Lasius
sabularum
from the site n. 28.
A:
Place and habitat of the
colony. Photo: June 2023. B: Few workers accumulated around a root aphid.
Photo: June 2023. C: Lateral view of worker. Scale bar: 1.0 mm. Photo: August
2024. D: Frontal view of worker’s head. Scale bar: 0.8 mm. Photo: August 2024.
Entomofauna carpathica, 2024, 36(2): 83-126
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Plagiolepis cf. taurica Santschi, 1920
Data: 25.06.18, 61, FI, 2 ww; 02.07.18, 61, FI, 5 ww; 30.03.19, 14, FI, 10 ww; 30.03.19,
14, FI, 10 ww; 30.03.19, 14, FI, 10 ww;18.04.19, 61, FI, 5 ww; 21.04.19, 16, FI, 10 ww;
18.03.20, 16, FI, 10 ww, 1 g; 15.07.20, 19, FI, 5 ww, 3 gg; 18.08.20, 18, FI, 5 ww, 1 g;
28.06.21, 18, FI, 10 ww, 1 g, 1 m; 13.08.21, 15, FI, 10 ww.
Species Plagiolepis pyrenaica Emery, 1921 (KIRSCHNER et al. 2023) was not
excluded during determinations. Specimens from the investigated area probably
did not belong to that species based on the zoogeography. Also, it has not been
reported from the Slovak Republic yet.
Plagiolepis pygmaea (Latreille, 1798)
Data: 11.03.18, 36, FI, 10 ww; 22.05.18, 61, FI, 3 ww, 1 g; 26.06.18, 38, FI, 10 ww;
02.07.18, 61, FI, 3 ww, 1 g, 2 mm; 07.08.18, 27, FI, 2 ww; 11.08.18, 68, FI, 5 ww; 19.08.18,
22, FI, 3 ww; 30.08.18, 32, FI, 10 ww, 2 gg; 06.03.19, 55, 5 ww; 13.04.19, 52, FI, 10 ww;
14.04.19, 45, FI, 10 ww, 3 gg; 01.06.19, 5, FI, 2 ww, 1 g; 16.07.19, 4, FI, 7 ww, unknown
whether from a single colony; 06.08.19, 20, FI, 1 w; 30.06.20, 56, FI, 5 ww; 15.07.20, 20, FI, 2
ww, 5 mm; 29.04.21, 50, FI, 10 ww; 13.08.21, 15, FI, 10 ww, 5 mm; 14.06.21–22.08.21,
45, PT, 15 ww; 02.10.21, 6, FI, 5 ww; 29.05.22, 10, FI, 3 ww, 5 gg; 05.07.23, 21, FI, 4 ww.
The finding of this tiny xerothermophilous species at the site n. 5 is its new
northernmost known distributional point in the Slovak Republic (CHYZER 1902,
ZÁLESKÝ 1939, SADIL 1953, BELÁKOVÁ 1961, LABUDA 1970, DRDULOVÁ 1979, 1991,
DRDULOVÁ & ZLATOŠOVÁ 1980, AMBROS et al. 1998, DEVÁN 2005, 2006b, 2007,
2008a, c, d, WIEZIK 2007, 2008a, b, c, 2010, WIEZIK & WIEZIKOVÁ 2007, 2012,
KLESNIAKOVÁ & HOLECOVÁ 2015, SUVÁK 2021, PURKART 2023a, JANCÍK & PURKART
2024, MARKO et al. 2024, JANCÍK, PURKART, unpub. data).
Myrmicinae
Aphaenogaster subterranea (Latreille, 1798)
Data: 15.07.17, 52, FI, 4 ww, 1 g; 14.04.18, 52, FI, 1 w; 22.04.18, 36, FI, 5 ww; 31.03.19,
36, FI, 1; 31.08.19, 36, FI, 1 g, after dispersal and nested (without workers); 09.05.20, 56,
FI, 10 ww; 30.06.20, 56, FI, 2; 05.09.20, 36, FI, 3 ww; 29.04.21, 52, FI, 60 ww; 04.05.21,
38, FI, 1 w; 09.0521, 36, FI, 10 ww; 06.08.21, 36, FI, 20 ww, 4 gg, 2 mm; 19.08.22, 26, 10
ww; 25.05.24, 38, FI, 50 ww.
Based on the reported data (ZÁLESKÝ 1939, LABUDA 1970, DRDULOVÁ 1991,
AMBROS et al. 1998, HOLECOVÁ et al. 2003, DEVÁN 2006a, 2008a, 2009a, WIEZIK
2007, 2008a, b, c, WIEZIK & WIEZIKOVÁ 2007, 2012, WIEZIK et al. 2010, PURKART &
HOLECOVÁ 2017, PURKART 2018, 2023a), the site n. 36 is currently the
northernmost known distributional point in the Slovak Republic. About 5
colonies were discovered there; all were nesting near the bases of living Pinus
Entomofauna carpathica, 2024, 36(2): 83-126
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sylvestris trees in the soil, or simply in the soil. Nests were mostly under stones
throughout all the sites except that one (stones were available to a small extent over
there). Northwards distribution in the direction of the Púchov valley (Fig. 1 – lies near
inhabited areas Dohňany, Záričie, and Lysá pod Makytou) seems to be cut off at the
northernmost site n. 36 – near the town Púchov and the Váh river. A suitable habitat
for this species appeared to be at site n. 5 (Fig. 2A). Unfortunately, it was not found
there; maybe it could have been present there in lower abundances, and due to
quick terrain inspections during the only two visits, it was undetected.
Formicoxenus nitidulus (Nylander, 1846)
Data: 16.04.19., 31, FI, 5 ww, 1 g, in nest of Formica polyctena; 18.05.20, 36, FI, 5 ww,
in nest of Formica pratensis.
Leptothorax acervorum (Fabricius, 1793)
Data: 01.09.19, 1, FI, 2 ww, unknown whether from a single colony; 06.04.20, 57, FI, 20
ww; 20.07.20, 64, FI, 10 ww, 10 gg, a colony with at least hundreds of workers, and at
least tens of wingless gynes, under bark of single Pinus sylvestris tree, and few
centimeters above the ground; 22.10.20, 34, FI, 10 ww; 05.7.23, 21, FI, 1 w.
Leptothorax gredleri (Mayr, 1855)
Data: 19.07.20, 3, FI, 5 ww, 2 gg; 19.07.20, 3, FI, 10 ww, 1 g; 17.08.20., 26, FI, 5 ww, 1 g;
21.08.20, 47, FI, 5 ww, 5 gg, 5 mm; 03.09.20, 54, FI, 1 w, 5 gg; 06.07.21, 64, FI, 10 ww, 2
gg; 15.07.21, 33, FI, 2 ww, 2 gg.
Additional findings: 28.03.20, 49.0951°,18.3215°, Považské Podolie Val., FI, 30 ww, 1g,
Salix tree on a cemetery; 14.11.21, 49.1909°,18.2304°, Javorníky Mts, FI, 10 ww, 1 g, Tilia
tree on a cemetery; 05.07.23, 49.1176°,18.2942°, Biele Karpaty Mts, FI, 5 ww, 1 g, Ulmus
tree near a small stream.
This species is considered as endangered in the Red (Ecosozological) List of
Hymenoptera of Slovakia (LUKÁŠ 2001). PURKART (2016) stated that there is very
poor knowledge on its distribution within the Slovak Republic what was and still
is true since the only available records are by KOŽÍŠEK (1985b) and DEVÁN (2007).
Therefore, the multiple newly confirmed sites listed herein substantially extend
the knowledge. This species was observed to nest exclusively under bark on the
trunks of living trees rather close to the ground – in accordance with SEIFERT (2018).
The trees were always older ones with much of the old bark providing enough
micro-spaces for nesting and hunting the prey. During the surveys, there were
no troubles in extracting the majority of colony members, including the queen(s),
directly from their nests as a palette knife was used to peel off the old layer of bark.
Leptothorax muscorum (Nylander, 1846)
Data: 01.09.19, 1, FI, 1 w.
Entomofauna carpathica, 2024, 36(2): 83-126
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Manica rubida (Latreille, 1802)
Data: 24.06.15, 25, FI, 3 ww; 15.10.17, 67, FI, 2 ww.
Myrmecina graminicola (Latreille, 1802)
Data: 12.07.17, 38, FI, 5 ww, 1 g; 14.06.21–22.08.21, 45, PT, 15 ww; 02.09.23, 38, LS, 4 ww.
Myrmica lobicornis Nylander, 1846
Data: 20.07.20, 64, FI, 1 w.
Myrmica rubra (Linnaeus, 1758)
Data: 13.04.20, 60, FI, 10 ww; 30.04.20, 57, FI, 10 ww, 5 gg, macrogynes and microgynes;
29.04.21, 40, FI, 20 ww; 30.04.21, 57, FI, 50 ww; 16.08.21, 23, FI, 10 ww, 5 gg, macrogynes
and microgynes; 04.04.22, 58, FI, 10 ww.
Keeping the data about the microgynes is useful if Myrmica microrubra Seifert,
1993 will be renewed from the current synonymy of M. rubra in the future.
Myrmica ruginodis Nylander, 1846
Data: 08.09.19, 38, FI, 5 ww.
Myrmica rugulosa Nylander, 1849
Data: 21.06.21, 51, FI, 10 ww; 06.05.22, 42, FI, 5 gg, unknown whether from a single
colony, after dispersal; 08.05.22, 42, FI, 12 ww; 06.07.22, 11, FI, 10 ww.
Myrmica sabuleti Meinert, 1861
Data: 22.05.18, 61, FI, 5 ww; 07.08.18, 27, FI, 2 ww; 26.09.18, 36, FI, 10 ww; 12.10.18,
36, FI, 20 ww; 20.10.18, 14, FI, 20 ww; 30.03.19, 14, FI, 5 ww; 31.03.19, 36, FI, 10 ww;
21.04.19, 16, FI, 10 ww; 07.08.20, 7, FI, 30 ww; 18.08.20, 18, FI, 4 ww; 13.08.21, 15, FI,
20 ww; 06.07.22, 11, FI, 10 ww.
Myrmica scabrinodis Nylander, 1846
Data: 05.05.21, 31, FI, 80 ww, 12 gg, 6 mm.
Myrmica schencki Viereck, 1903
Data: 11.05.18, 36, FI, 2 ww; 08.07.18, 52, FI, 1 w; 11.08.18, 68, FI, 6 ww, 1 m; 30.08.18, 32, FI, 1
w; 22.09.18, 36, FI, 1 w; 08.07.20, 70, FI, 1 w; 24.04.21, 36, FI, 4 ww; 10.08.21, 31, FI, 1 w.
Myrmica specioides Bondroit, 1918
Data: 15.07.18, 46, FI, 5 ww; 06.09.18, 36, FI, 10 ww; 15.08.20, 68, FI, 1 ww; 21.05.21,
36, FI, 8 ww.
Entomofauna carpathica, 2024, 36(2): 83-126
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Solenopsis cf. fugax (Latreille, 1798)
Data: 11.03.18, 36, FI, 8 ww; 24.04.21, 26, FI, 20 ww; 14.06.21–22.08.21, 45, PT, 15 ww;
25.08.22, 38, FI, 6 ww; 26.08.22, 52, FI, 10 ww; 28.08.22, 50, FI, 2 ww; 02.09.22, 20, FI, 4 ww.
Specimens were not investigated using the complex morphological analysis
(CSŐSZ et al. 2023), therefore, species S. juliae Csősz et al. (2023) was not
excluded from determinations. The probability of the latter species to occurring
within the investigated area was rather low due to zoogeography. Also, this
species has not been detected in the Slovak Republic yet.
Stenamma debile (Förster, 1850)
Data: 22.09.17, 52, FI, 1 w; 29.04.18, 38, LS, 1 w, 1 g; 01.05.18, 52, LS, 3 ww; 14.09.18,
45, FI, 1 g, after dispersal; 14.10.18, 14, FI, 1 g, after dispersal; 17.09.19, 52, FI, 1 w;
14.06.21–26.07.21, 45, PT, 3 ww, 1 g.
Strumigenys argiola (Emery, 1869)
Data: 30.07.18–30.08.18, 45, PT, 1 m; 14.06.21–22.08.21, 45, PT, 2 gg, 5 mm; 26.08.22–
03.09.22, 45, PT, 1 m.
Findings in the years 2018 and 2021 have already been published (PURKART et
al. 2021); new finding is from the year 2022. Another attempt to record this
species using the pitfall traps was conducted in the year 2023, but this one was
unsuccessful. FI and LS methods were deployed to find a nest or workers (Fig. 10)
at the site, but it did not bring success.
Fig. 10. Worker of Strumigenys argiola in laboratory keeping, showing its elongated
mandibles. Specimen originated from S Slovakia. Photo: February 2024.
Entomofauna carpathica, 2024, 36(2): 83-126
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Temnothorax affinis (Mayr, 1855)
Data: 14.03.18, 44, FI, 20 ww, 1 g; 17.03.18, 52, FI, 20 ww, 1 g; 16.04.18, 52, FI, 3 ww; 29.06.18, 36,
FI, 10 ww; 14.08.18, 65, FI, 5 ww; 04.09.18, 52, FI, 20 ww, 1 g; 06.03.19, 55, FI, 5 ww, 1 g; 17.03.19,
36, FI, 10 ww, 1 g; 23.03.19, 36, FI, 20 ww, 1 g; 23.07.21, 30, FI, 3 ww; 26.06.22, 12, FI, 5 ww.
Temnothorax albipennis (Curtis, 1854)
Data: 08.07.20, 70, FI, 40 ww, 1 g, nest under moss and on stone; 05.11.21, 73, FI, 54 ww, 1
g, nest in rock crevices, rev. Seifert; 17.06.22, 46, FI, 24 ww, 11 gg, nest under stone and on
rocky soil; 07.07.22, 53, FI, 25 ww, 3 gg, 2 mm, nest in grass tuft and rocky soil; 29.06.23, 24,
FI, 4 ww, nest in rock crevices; 05.07.23, 21, FI, 10 ww, 1 g, nest in rock crevice and in soil;
24.07.23, 56, FI, 1 w; 29.07.23, 56, FI, 75 ww, 1 g, nest in rocky soil.
Available records from the Slovak Republic were well summarised by JANCÍK &
PURKART (2024). Additional data were reported by PURKART et al. (2024) and
Fig. 11.
Temnothorax
albipennis
.
A:
Worker with colony
-
average pigmentation. Scale
bar: 0.9 mm. Photo: September 2021. Individuals in photos B–D were part of the
single colony collected at the site n. 24. B: Alated gyne left mother colony and is
headed to swarming, locating on a limestone rock. Photo: July 2023. C: Colony
members including brood in laboratory keeping. Photo: December 2023.
D: Workers processing a prey – Pholcus phalangioides (Füssli, 1775) individual.
Photo: January 2024.
Entomofauna carpathica, 2024, 36(2): 83-126
112
MARKO et al. (2024). Thorough searches were conducted to discover this species,
which resulted in detection of 7 colonies. Findings from the Biele Karpaty Mts
come from even 5 sites, what contrasts with no records from that area until now.
This can be explained by the lower abundances of the species in the field, as
confirmed by this survey, where only one nest was discovered at each site. Also,
it can be explained by possible misidentifications with any of the sibling species
(mainly with T. tuberum (Fabricius, 1775)) sharing similar body features in the
past. One such similarity was observed during this study when some colonies had
abnormally darkly pigmented average workers (Fig. 11A) – resembling T.
tuberum pattern. Such a colony came from site, e.g., n. 73, which included a
worker depicted in the Fig. 11A. This phenomenon can be confusing during
determinations, what requires consideration of morphological signs, too.
Temnothorax corticalis (Schenck, 1852)
Data: 21.09.18, 36, FI, 9 ww, under the bark of living Pinus sylvestris tree; 20.07.20, 64,
FI, 4 ww, unknown whether from a single colony, on the bark of a living coniferous tree;
20.07.20, 64, FI, 5 ww, 1 g.
Temnothorax crassispinus (Karavajev, 1926)
Data: 15.03.18, 36, FI, 10 ww, 2 gg; 21.08.20, 47, FI, 10 ww; 03.09.20, 54, FI, 10 ww;
22.10.20, 34, FI, 20 ww; 06.04.21, 37, FI, 20 ww, 1 g; 15.06.21, 45, LS, 10 ww; 15.07.21,
33, FI, 20 ww; 01.05.24, 35, FI, 15 ww.
Temnothorax interruptus (Schenck, 1852)
Data: 28.07.18, 66, FI, 20 ww, 1 g, macrogyne; 02.08.18, 61, FI, 20 ww, 1 g, macrogyne;
02.08.18, 61, FI, 10 ww, 1 g, macrogyne; 02.08.18, 61, FI, 10 ww, 1 g, macrogyne;
02.08.18, 61, FI, 10 ww, 1 g, macrogyne; 15.09.18, 61, FI, 20 ww; 07.08.19, 20, FI, 1 w;
17.08.19, 20, FI, 4 ww, unknow whether from a single colony; 17.03.20, 61, FI, 10 ww, 1
g, macrogyne; 15.07.20, 19, FI, 1 w; 11.09.20, 20, FI, 1 w; 10.10.20, 61, FI, 30 ww, 1 g,
macrogyne; 02.10.21, 6, FI, 10 ww, 1 g, macrogyne; 17.06.22, 49, FI, 10 ww, 1 g,
macrogyne; 17.06.22, 53, FI, 2 ww; 25.06.22, 36, FI, 1 g, macrogyne, after dispersal and
in soil without workers – probably establish a new colony; 08.05.23, 56, FI, 1 w; 30.06.23,
8, FI, 38 ww, 1 g, 11 mm, macrogyne; 05.07.23, 21, FI, 1 w; 24.07.23, 36, FI, 10 ww;
29.07.23, 56, FI, 80 ww, 1 g, macrogyne; 29.07.23, 56, FI, 8 ww, 2 gg, 9 mm, macrogynes.
BEZDĚČKA (2000, 2018) summarised findings of ant colonies which built their
nests entirely or partly within shells of snails or clams. My two last findings at site
n. 56 in the year 2023 refer to colonies completely nesting in shells of the snail
Xerolenta obvia (Menke, 1828). This ant species extends the Bezděčka’s list for
the species of snail. The shells were not damaged; they were located on the
ground surface, slightly in vegetation humus, and surrounded by grass-herbal
vegetation cover. The shells’ opening parts were silted by the ants with soil
Entomofauna carpathica, 2024, 36(2): 83-126
113
particles, causing only a small entrance hole with a diameter of 1–2 mm to be
kept for each shell. This species also forms microgynes, but no such individual
was part of the collected material.
Temnothorax nigriceps (Mayr, 1855)
Data: 30.03.18, 66, FI, 20 ww, 1 g; 28.07.18, 66, FI, 10 ww, 1 g; 07.08.18, 27, FI, 5 ww,
unknown whether from a single colony; 11.08.18, 68, FI, 3 ww, 1 g; 11.08.18, 68, FI, 10
ww, 1 g; 16.08.18, 69, FI, 20 ww , 1 g; 19.08.18, 22, FI, 2 ww, unknown whether from a
single colony; 30.08.18, 32, FI, 2 ww, unknown whether from a single colony; 15.04.19,
31, FI, 10 ww, 1 g; 01.09.19, 1, FI, 2 ww, unknown whether from a single colony; 08.07.20,
70, FI, 5 ww, unknown whether from a single colony; 17.06.22, 49, FI, 10 ww; 05.07.23,
21, FI, 10 ww; 26.05.24, 62, FI, 40 ww, 1 g, under moss on the concrete surface.
Temnothorax parvulus (Schenck, 1852)
Data: 22.04.18, 36, FI, 10 ww; 21.04.20, 65, FI, 1 w; 30.06.20, 53, FI, 1 w; 07.07.20, 61, FI,
1 w; 05.09.20, 45, LS, 6 ww; 05.09.20, 36, LS, 6 ww; 15.06.21, 45, LS, 10 ww; 31.10.21,
56, FI, 100 ww, 2 gg, wingless gynes; 17.09.22, 45, FI, 10 ww, 1 g.
Temnothorax saxonicus (Seifert, 1995)
Data: 07.08.18, 27, FI, 2 ww, unknown whether from a single colony; 19.08.18, 22, FI, 1
w; 30.08.18, 32, FI, 10 ww, 1 g, nest in rock crevice; 30.08.18, 32, FI, 1 w.
This species was considered as vulnerable in the Red (Ecosozological) List of
Hymenoptera of Slovakia (LUKÁŠ 2001). The finding of the nest was done only at
the site n. 32, all other findings contained workers collected outside their nests.
The site n. 22 was the northernmost known distributional point in the Slovak
Republic (KOŽÍŠEK 1986, DEVÁN 2006a, 2009b, WIEZIK & WIEZKOVÁ 2012, SUVÁK
2021, PURKART 2023a, MARKO et al. 2024).
Temnothorax tuberum (Fabricius, 1775)
Data: 21.04.19, 16, FI, 8 ww, 1 g, nest in rock crevice; 15.06.20, 14, FI, 14 ww, 1 g, nest in
moss.
Temnothorax unifasciatus (Latreille, 1798)
Data: 30.03.18, 66, FI, 20 ww, 1 g; 30.03.18, 66, FI, 20 ww, 1 g; 03.04.18, 52, FI, 10 ww, 1 g;
04.04.18, 66, FI, 20 ww, 1 g; 11.05.18, 36, FI, 5 ww, 1 g; 02.08.18, 61, FI, 10 ww, 1 g; 16.08.18,
69, FI, 15 ww, 1 g; 30.08.18, 32, FI, 10 ww, 1 g; 07.09.18, 31, FI, 10 ww, 1 g; 22.09.18, 36, FI,
10 ww; 14.10.18, 14, FI, 5 ww, 1 g; 14.10.18, 14, FI, 10 ww, 1 g; 04.03.19, 55, FI, 20 ww, 1 g;
06.03.19, 55, FI, 20 ww, 1 g; 07.03.19, 61, FI, 10 ww, 1 g; 21.04.19, 17, FI, 10 ww, 1 g; 22.04.19,
17, FI, 10 ww, 1 g; 14.09.19, 5, FI, 20 ww; 18.03.20, 16, FI, 10 ww, 1 g; 30.06.20, 56, FI, 10 ww,
1 g; 19.07.20, 3, FI, 1 w; 07.08.20, 7, FI, 20 ww, 1 g; 29.05.22, 10, FI, 20 ww; 17.06.22, 49, FI,
16 ww, 1 g; 05.05.23, 45, FI, 20 ww, 1 g; 05.05.23, 38, FI, 15 ww, 1 g; 05.05.23, 37, FI, 10 ww;
07.05.23, 27, FI, 1 w; 09.05.23, 28, FI, 5 ww; 13.05.23, 22, FI, 2 ww; 13.05.23, 21, FI, 1 w;
Entomofauna carpathica, 2024, 36(2): 83-126
114
16.05.23, 6, FI, 1 w; 16.05.23, 4, FI, 25 ww; 16.05.23, 18, FI, 2 ww; 16.05.23, 20, FI, 2 ww,
unknown whether from a single colony; 18.05.23, 13, FI, 13 ww; 18.05.23, 19, FI, 3 ww; 21.05.23,
68, FI, 4 ww; 01.06.23, 65, FI, 3 ww; 03.06.23, 70, FI, 41 ww; 06.06.23, 71, FI, 2 w; 08.06.23, 15, FI, 2
ww; 10.06.23, 9, FI, 12 ww, 1 g.
Tetramorium caespitum (Linnaeus, 1758)
Data: 25.06.22, 10, FI, 10 ww, 1 g, 1 m; 08.05.23, 36, FI, 30 ww, 7 gg, 2 mm; 08.05.23,
56, FI, 20 ww, 2 gg, 8 mm.
Colony samples of Tetramorium caespitum species complex also containing
males were determined to the species level based on the male specimens, and
to a small extent, based on the workers too. These determinations always had
clear results for T. caespitum, or T. immigrans. Colony samples that were not
determined to the species level did not contain males. Nevertheless, species T.
immigrans and T. hungaricum were ruled out from them based on the worker
caste. Worker specimens were not investigated with complex morphological
analysis.
Tetramorium immigrans (Santschi, 1927)
Data: 21.06.22, 62, FI, 84 ww, 1 g, 1 m, sexuals were pupae; 28.06.22, 63, FI, 80 ww, 14
gg, 13 mm, part of sexuals were pupae; 02.07.22, 63, FI, 30 mm; 17.07.22, 63, FI, 40 mm;
25.06.23, 28, FI, 20 ww, 6 gg, 5 mm, sexuals were pupae.
See comment on T. caespitum. T. immigrans was reported only once from the
Slovak Republic WAGNER et al. (2017). The site n. 28 represented the
northernmost distributional point of this species in the Slovak Republic. This
species was first discovered at the site n. 63 where it nested in spaces between
older concrete where the substrate was gravel-sand and vegetation humus. A
single colony was found at the site n. 28 where it nested in gravel sand near the
country road and the railway embankment. If the sexuals were collected as
pupae, then they were kept alive and put under laboratory conditions to develop
into adult stage, and subsequently the determinations to be possible based on
the male caste. This species is synanthropic in the Central Europe (SEIFERT 2018),
and the ruderal habitats where it was found during this survey are well-known
for it (SEIFERT 2018). Average worker is the largest one among all the species in
the T. caespitum species complex discovered in the Slovak Republic so far. Its
occurrence in the Slovak Republic is strongly under-recorded (JANCÍK, unpub.
data), what is probably because its similarity to other species of the complex and
due to its synanthropic occurrence, usually at grubby places, as are, e.g.,
roadsides or pavements – places usually not interesting for myrmecological
research at the first sight.
Entomofauna carpathica, 2024, 36(2): 83-126
115
Tetramorium moravicum Kratochvíl, 1941
Data: 23.05.18, 61, FI, 12 ww; 27.08.18, 61, FI, 20 ww; 18.04.19, 61, FI, 60 ww; 10.09.21,
61, FI, 10 ww; 19.06.22, 61, FI, 40 ww.
The site n. 61 represented the northernmost known distributional point
within the Slovak Republic (DRDULOVÁ & ZLATOŠOVÁ 1980, AMBROS et al. 1998,
WIEZIK 2007, 2008a, b, c, WIEZIK & WIEZIKOVÁ 2007, 2012, WIEZIK et al. 2010,
PURKART et al. 2019, PURKART 2020, SUVÁK 2021, MARKO et al. 2024, JANCÍK,
PURKART, unpub. data). Only one colony was discovered on a small rocky slope
(Fig. 3B).
Tetramorium sp(p). (caespitum species complex)
Data: 18.08.18, 68, FI, 30 ww; 27.08.18, 61, FI, 20 ww; 30.08.18, 32, FI, 20 ww; 04.09.18,
52, FI, 30 ww; 14.04.19, 45, FI, 20 ww; 20.04.19, 67, FI, 20 ww; 21.04.19, 16, FI, 20 ww
See comment on T. caespitum.
Ponerinae
Ponera coarctata (Latreille, 1802)
Data: 06.07.18, 36, FI, 1 w; 21.04.19, 16, FI, 1 w; 30.06.20, 56, FI, 1 w; 06.07.20, 42, FI, 1 w.
Ponera testacea Emery, 1895
Data: 22.08.17, 36, FI, 1 w; 30.09.17, 36, FI, 3 ww; 25.06.18, 61, FI, 1 w; 26.06.18, 38, FI,
2 ww; 08.07.18, 52, FI, 1 w; 30.08.18, 32, FI, 1 ww; 19.04.19, 45, FI, 1 w; 22.04.19, 17, FI,
1 w; 12.5.20, 36, FI, 1 w; 15.09.20, 45, FI, 2 ww; 29.04.21, 50, FI, 2 ww, 3 gg; 25.06.21, 45,
FI, 1 w; 06.08.21, 36, 8 ww, 3 gg; 25.06.22, 36, FI, 1 w; 28.06.22, 36, FI, 1 w; 23.06.23, 37,
FI, 1 w; 30.06.23, 8, FI, 1 w.
The specimens collected at sites n. 17 and n. 61 were found at semi-shaded
places in light Pinus sylvestris forest with grassy undergrowth. Although, these
sites were gradually becoming more and more shaded due to succession process
and forest development during past decades, permanent occurrence of this
species was confirmed here, and, therefore, these observations extend its
ecological data (SEIFERT 2018).
Proceratiinae
Proceratium melinum (Roger, 1860)
Data: 26.09.17, 36, FI, 1 g, after dispersal, fully winged, at 15:50h.
Entomofauna carpathica, 2024, 36(2): 83-126
116
The gyne (Fig. 12) was found during wind-free weather at the site containing
suitable, though degrading, habitat. It is likely that its mother colony was located
at that site at that time. However, targeted attempt to find nest or workers
failed, anyway, cryptic subterranean habits make this species mostly
unreachable. The gyne was caught when located on the ground surface on a
Pinus sylvestris root protruding from the country road in the ecotone. It was
immediately transported into laboratory to study its behaviour, where it was
placed into a small tube with diameter of seven mm with a sufficient humidity
level. It was trying to put off its wings for even 4 days from the day when it was
found until when all its wings were put off on the September 29, 2017. This long
time could be influenced by the environment in the laboratory keeping. Wings
putting off behaviour was carefully observed. The gyne was pushing its abdomen
upwards, moving the wings aside, and downwards, and trying to wrest the wings
by hind leg. Wings at one side were wedged between the femur located in front
of the wings, and the tibia located behind them. After the wings were wedged,
the gyne was trying to pull out them by moving the leg downwards. The gyne
was also observed to move slightly backward from time to time, what was
causing movement of its body, but wings were mostly not moving since they
were moved aside and downwards, touching the surface.
Fig. 12. Dealate gyne of Proceratium melinum. Specimen originating from the site n.
36. Scale bar: 0.6 mm. Photo: August 2024.
Entomofauna carpathica, 2024, 36(2): 83-126
117
This species was rarely reported from the Slovak Republic (WERNER & WIEZIK
2007, KLESNIAKOVÁ et al. 2016, PURKART & REPTA 2022). The newly presented site
was the northernmost known distributional point of this species within the
Slovak Republic, making it more precious as it is the only known site with natural
habitat so far. Other records with known habitat come from synanthropic areas
(KLESNIAKOVÁ et al. 2016, PURKART & REPTA 2022, JANCÍK, unpub. data). Species was
reported from neighbouring Moravian region at more (two) sites with the natural
habitats (BEZDĚČKA 1992, 1996b, 2007b, 2008a, BEZDĚČKA & BEZDĚČKOVÁ 2013).
DISCUSSION
A total of 75 ant species were detected across the study sites, and for each
species its permanent presence (meaning colony occurrence) was confirmed
(directly or indirectly) at least for a specific bounding geomorphological unit, if
not for a certain site. It represents more than 62% of the total number of 120
free-living ant species reported from the Slovak Republic (KOŽÍŠEK 1984, 1987,
SEIFERT 1988, WERNER & WIEZIK 2007, DEVÁN 2008a, WAGNER et al. 2011, SEIFERT
2012, WIEZIK & WIEZIKOVÁ 2013, BEZDĚČKA & TĚŤÁL 2013, SEIFERT & GALKOWSKI 2016,
WAGNER et al. 2017, SUVÁK 2021, PURKART & REPTA 2022). Base for this species
enumeration is the last specimen-based checklist by WERNER & WIEZIK (2007)
which confirmed 106 species. Each one of most of the later studies (7) document
one species. The other later studies of WAGNER et al. 2017 and SUVÁK 2021 bring
per 2 species. KOŽÍŠEK (1987) reported the species Formica picea Nylander, 1846,
which specimen(s) was not available for the last revision (WERNER & WIEZIK 2007).
The authors of the checklist omitted and did not refer to KOŽÍŠEK (1984) and
SEIFERT (1988). Reported species from these studies are Lasius carniolicus Mayr,
1861, and Lasius sabularum (Bondroit, 1918), respectively. BEZDĚČKA & TĚŤÁL
(2013) reported the species Cardiocondyla elegans Emery, 1869, but revision of
SEIFERT (2023) marked this record as Cardiocondyla dalmatica Soudek, 1925.
This survey was primarily focused on the easternmost part of the Biele
Karpaty Mts where the ant fauna was studied on 63% of all the sites. Permanent
presence of 73 ant species was confirmed for this geomorphological unit,
representing portion of more than 60% of the total Slovak myrmecofauna (as
referred above).
Summarising ant species that have been reported from the Biele Karpaty Mts
(see Material and Methods) so far brings territory incompatibility with an area of
“Biele Karpaty” referred by BEZDĚČKA & BEZDĚČKOVÁ (2010) in their checklist.
Therefore, species from the checklist cannot be included within the Biele Karpaty
Mts, otherwise, some of them could potentially drop from the summary. The
summarisation can be done only for an area of Biele Karpaty Mts Merged
Entomofauna carpathica, 2024, 36(2): 83-126
118
comprising both the mentioned territories. This is because records for some
species are available only in the checklist where they absent any concrete
collecting data, making these records unable to assign or not to be assigned to
the area of the Biele Karpaty Mts. This way, unfortunately, the area of the Biele
Karpaty Mts Merged is undefined since the area of “Biele Karpaty” was
undefined by BEZDĚČKA & BEZDĚČKOVÁ (2010).
The summary of ant species with confirmed colony occurrence for the Biele
Karpaty Mts Merged consists of species records reported in the past, and these
ones are presented herein. The old records comprise 71 ant species – DEVÁN
(2006a) (1), BEZDĚČKA & BEZDĚČKOVÁ (2010) (69), and PURKART et al. (2021) (1). The
checklist lacks species Anergates atratulus, which was reported by DEVÁN (2006a)
and referred to by PURKART (2023b). The authors of the checklist operated
probably only with material of specimens which was available to them. This study
adds 10 species – P. melinum, P. testacea, T. albipennis, T. corticalis, T.
immigrans, L. paralienus, L. myops, L. sabularum, L. bicornis, and L. reginae.
Therefore, the summarised list contains 81 species representing more than 68%
of the total Slovak myrmecofauna (as referred above). This large list increase was
only partly predicted by the authors of the checklist – they predicted the possible
occurrence of 8 species that could be additionally found in the future, but only
two of them are being confirmed herein – P. testacea, and L. paralienus. The
author of this study supposes that Myrmica gallienii Bondroit, 1920, Myrmica
karavajevi (Arnol'di, 1930), Tetramorium staerckei Kratochvíl, 1944,
Myrmoxenus ravouxi (André, 1896), Strongylognathus testaceus (Schenck,
1852), and Plagiolepis xene Stärcke, 1936 can be found within the area in the
future, ordering them by the highest probability.
ACKNOWLEDGEMENTS
I would like to thank landowners for permitting the field survey. I would like to
thank Dr. Bernhard Seifert for his determinations. I would like to thank Dr. Adrián
Purkart for provided part of used chemistry deployed in the field. I would like to
thank Petr Werner for specimens coming from his personal collection. Lastly, I
am grateful to Hubert Kozaczuk for supplying live ant material.
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SÚHRN
Práca prináša nové faunistické údaje mravcov prevažne z oblasti
najvýchodnejších častí Bielych Karpát, čo je oblasť rozprestierajúca sa najmä pri
meste Púchov a obci Streženice. Toto hornaté územie je zároveň v severnej časti
daného geomorfologického celku a tiež má relatívne severnú pozíciu v rámci
krajiny. Aj napriek tomu tu boli zistené vzácne teplomilné druhy mravcov. Nie je
preto prekvapením, že niektoré z týchto druhov tu dosahujú svoje známe
severné hranice rozšírenia v rámci Slovenskej Republiky; sú nimi napríklad
Aphaenogaster subterranea, Lasius myops, Lasius reginae, či Proceratium
melinum. Práca prináša aj nové poznatky o biológii vzácneho arborikolného
druhu Lasius bicornis, kde sa v chovných podmienkach podarilo simulovať jav
založenia kolónie v prírode. Niekoľko skúmaných lokalít sa nachádza aj v okolí
Bielych Karpát – v rámci geomorfologických celkov Javorníky a Považské podolie.
Dôkladný, 10 ročný prieskum spoločenstiev mravcov mal za cieľ odhaliť mnohé
zriedkavo nachádzané druhy, čo sa vo veľkej miere aj podarilo. Za takéto druhy
možno považovať najmä Lasius myops, Lasius reginae, Lasius sabularum,
Leptothorax gredleri, Proceratium melinum, Strumigenys argiola, Temnothorax
albipennis, či Tetramorium immigrans. Celkovo bolo zistených 75 druhov
mravcov zo všetkých piatich čeľadí, ktorých zástupcovia sú známi zo Slovenska.
U vybraných druhov mravcov sú uvedené všetky dostupné historické údaje o ich
výskyte, čo okrem iného dokumentuje vzácnosť ich nálezov.