A review of two very rare ground spiders from sandy habitats, new for Slovakia (Gnaphosidae, Araneae)

Abstract

Two rare and interesting spider species (Gnaphosa mongolica Simon, 1895 and Haplodrassus bohemicus Miller & Buchar, 1977) were found during intensive research into the Pannonic sand steppes in Slovakia and represent new records for the country. Numerous specimens of both species were collected between the villages Radvaň nad Dunajom and Marcelová in the years 2017–2020 (145 ind. G. mongolica and 82 ind. H. bohemicus), indicating stable populations. The Slovak records of G. mongolica represent the northernmost and westernmost location of the species in Europe. The characteristic features, photos of the habitus and genitalia, the currently known distribution in Europe and the phenology of both species are provided. A more detailed discussion concerns the morphology of copulatory organs of H. bohemicus, including the first report of a lateral gynandromorphy in this species.

Full text

Arachnologische Mitteilungen / Arachnology Letters 64: 14-24 Karlsruhe, Dezember 2022
Sandy habitats in Slovakia belong to interesting and unique
ecosystems and are inhabited by several rare and endangered
taxa. However, so far only a few studies on spiders were con-
ducted there (Kalivodová et al. 2002, 2008, Prídavka 2002,
Gajdoš & Majzlan 2001, 2008, 2010, Gajdoš et al. 2019). e
Pannonic sand steppes (EUNIS code 6260), one of the most
endangered habitats of the Pannonian region, are character-
ised by semi-natural dry grasslands and scrubland on mobile
or xed sands (EUNIS 2021). In Slovakia, only small frag-
ments of this habitat are found on the Danubian plain and in
the east Slovak lowlands. e major threats are abandonment
of traditional land use and sand extraction, which lead to af-
forestation and spread of invasive species (Šeerová Stanová
et al. 2008). Spiders of the Pannonic steppes localities in
Slovakia were partly studied by Gajdoš & Majzlan (2001).
In recent years, several species new to the Slovak fauna have
been discovered in sandy habitats: Parasyrisca arrabonica Szi-
netár & Eichardt, 2009 (Gajdoš & Majzlan 2010), Erigono-
plus foveatus (Dahl, 1912) (Hollá et al. 2016; although it was
recorded on a tree by accident), Spiracme mongolica (Schenkel,
1963) and Walckenaeria stylifrons (O. Pickard-Cambridge,
1875) (Purgat et al. 2021).
e initial goal of our study was to improve the knowledge
of spider species living in Pannonian sand habitats, which led
to the discovery of two spider species new for the Slovak fau-
na. e ground-dwelling gnaphosids Gnaphosa mongolica Si-
mon, 1895 and Haplodrassus bohemicus Miller & Buchar, 1977
are widespread in the Palearctic (except in Western Europe)
(World Spider Catalog 2022). e rst species, G. mongolica,
can be easily identied using the online key of Nentwig et al.
(2022). However, H. bohemicus can be easily confused with re-
lated species, which causes some uncertainty about its previous
identications and localities. Its diagnosis using known de-
scriptions in the literature is ambiguous due to an inconsistent
terminology (e.g. embolic/terminal apophysis) and poorly de-
ned diagnostic characters that can be misunderstood, which
is due to slightly dierent angles from which the male palp has
been illustrated. erefore, we focused in depth on copulatory
organs of the latter species and discuss misidentied species.
For both species, we also summarize all available data on the
distribution, habitat preferences, phenology and red list status
in Europe. In addition, we present a new case of a bilateral
gynander found in a specimen of H. bohemicus.
Material and methods
Spiders were collected by pitfall trapping using ve traps in
one transect at each study site. ey were emptied monthly
from March 2017 to April 2019 in Domáňovský majer and
from March 2019 to April 2020 at other study sites. e traps
consisted of a plastic owerpot inserted in the ground, into
which a removable plastic cup (diameter 10 cm) with a for-
malin solution was inserted. e here presented phenology
summarizes the new data from Slovakia including published
records and unpublished data, always dated to the last day
the traps were emptied. Habitat summaries were made by
simplifying habitat data into four categories: 1. sand dunes
(habitat with sparse or almost no vegetation on sandy soil), 2.
steppes (grassy habitat with sparse vegetation on sandy soils
or other xerotherm habitats), 3. meadows (dense grassy veg-
etation growing on other type of soil, including records where
A review of two very rare ground spiders from sandy habitats, new for Slovakia
(Araneae: Gnaphosidae)
Anna Šestáková, Ľudmila Černecká, Maria Naumova, Pavol Purgat, Éva Szita & Peter Gajdoš
doi: 10.30963/aramit6402
Abstract. Two rare and interesting spider species (Gnaphosa mongolica Simon, 1895 and Haplodrassus bohemicus Miller & Buchar, 1977)
were found during intensive research into the Pannonic sand steppes in Slovakia and represent new records for the country. Numerous
specimens of both species were collected between the villages Radvaň nad Dunajom and Marcelová in the years 2017–2020 (145 ind.
G.mongolica and 82 ind. H. bohemicus), indicating stable populations. The Slovak records of G. mongolica represent the northernmost
and westernmost location of the species in Europe. The characteristic features, photos of the habitus and genitalia, the currently known
distribution in Europe and the phenology of both species are provided. A more detailed discussion concerns the morphology of copula-
tory organs of H.bohemicus, including the rst report of a lateral gynandromorphy in this species.
Keywords: Bulgaria, faunistic, Gnaphosa mongolica, gynandry, Haplodrassus bohemicus, Pannonic sand steppes, species conservation,
sand dunes
Zusammenfassung. Eine Übersicht zu zwei sehr seltenen Plattbauchspinnen aus Sandhabitaten, neu für die Slowakei (Araneae:
Gnaphosidae). Zwei seltene und interessante Spinnenarten (Gnaphosa mongolica Simon, 1895 und Haplodrassus bohemicus Miller &
Buchar, 1977) wurden während intensiven Untersuchungen der pannonischen Sandsteppenhabitate in der Slowakei gefunden und
sind neue Nachweise für das Land. Eine große Anzahl Exemplare beider Arten wurden zwischen den Dörfern Radvaň nad Dunajom und
Marcelová in den Jahren 2017-2020 (145 Individuen von G. mongolica und 82 Individuen von H. bohemicus), was auf stabile Populationen
hindeutet. Die slowakischen Nachweise von G. mongolica stellen die nördlichsten und westlichsten Lokalitäten der Art dar. Die charak-
teristischen Merkmale, Habitus, Genitalien und die aktuell bekannte Verbreitung der Art in Europa sowie die Phänologie beider Arten
werden präsentiert. Eine etwas detailliertere Diskussion befasst sich mit der Morphologie der Kopulationsorgane von H. bohemicus,
einschließlich einer erstmals beobachteten lateralen Gynandromorphie.
Anna ŠESTÁKOVÁ, The Western Slovakian Museum, Múzejné nám. 3, SK-91809
Trnava, Slovakia; E-mail: asestakova@gmail.com
Ľudmila ČERNECKÁ, Institute of Forest Ecology, Slovak Academy of Sciences,
Ľudovíta Štúra 2, SK-96053 Zvolen, Slovakia; E-mail: cernecka@ife.sk
Maria NAUMOVA, Institute of Biodiversity and Ecosystem Research, Bulgarian Acad-
emy of Sciences, Tsar Osvoboditel blvd., 1000 Soa, Bulgaria; E-mail: munny@abv.bg
Pavol PURGAT, Institute of Landcape Ecology, Nitra Branch, Slovak Academy of Sci-
ences, Akademická 2, SK-94901 Nitra, Slovakia; E-mail: pavol.purgat@savba.sk
Éva SZITA, Plant Protection Institute, Centre for Agricultural Research, Herman Ottó
u. 15, H-1022, Budapest, Hungary; E-mail: szita.eva@atk.hu
Peter GAJDOŠ, Institute of Landcape Ecology, Nitra Branch, Slovak Academy of Sci-
ences, Akademická 2, SK-94901 Nitra, Slovakia; E-mail: p.gajdos@savba.sk
Academic Editor: Tobias Bauer
Submitted 29.10.2021, accepted 20.8.2022, online 28.12.2022

Rare ground spiders from sandy habitats in Slovakia 15
no soil or exposure was specied), and 4. other habitats (for-
est, potentially including ecotones, and vineyards). All these
records are summarized in the Appendix Tab. S1. Both maps
were created in QGIS 3.20 software using MapTiler plugin
for basemaps.
Faunistic data are listed in the following order: abbrevia-
tion of the locality name, date of collecting, number and sex
of specimens (( – female, ) – male, sad.)/( – subadult male/
female, j – juvenile), collector. e notes on comparative ma-
terial contain additional information about the locality, name
of the collector and the reference where the record was pub-
lished. Spiders were identied to species level whenever pos-
sible following Nentwig et al. (2022), using the nomenclature
in the World Spider Catalog (2022). e terminology of cop-
ulatory organs for the genus Haplodrassus follows Bosmans et
al. (2018). Digital images were taken using a stereomicroscope
OLYMPUS SZX16, Leica DVM6, and Canon EOS1300D
digital camera attached to Stemi 2000-c stereomicroscope
and were combined in Zerene Stackerv.1.04. Measurements
were made using AxioVisionv.4.6, the values were compiled
from published and original data (see also appendix S1). e
specimens are stored in 70% ethanol and were deposited in
the collection of P. Gajdoš at the Institute of Landscape Ecol-
ogy of Slovak Academy of Sciences in Nitra, Slovakia, the
only exception being the gynandromorph specimen, which is
deposited at the Institute of Biodiversity and Ecosystem Re-
search, Bulgarian Academy of Sciences.
e countries included in the manuscript are divided into
three groups: Central Europe (C-EU): Austria, Czechia,
Hungary, Poland, Slovakia; Eastern Europe (E-EU): Rus-
sia, Ukraine; Southeastern Europe (S-EU): Bulgaria, Greece,
North Macedonia, Romania, Serbia.
Fig. 1: Map and photos of studied sites in Slovakia: 1. Domáňovský majer; 2. Virt; 3. Mašan Nature Reserve; 4. Bašovský Kopec; 5. Marcelovské piesky
Nature Reserve (photo by O. Majzlan)

16 A. Šestáková, Ľ. Černecká, M. Naumova, P. Purgat, É. Szita & P. Gajdoš
Study sites
e study sites (Fig. 1) were situated between the villages
Radvaň nad Dunajom and Marcelová near the Hungarian
border in the Danubian lowland, belonging to the North-
East side of the Pannonian basin (south-western Slovakia).
1. Domáňovský majer – DM (Domány puszta) (47.76290°N,
18.33813°E, 125 m a.s.l.). A farm with a mosaic of dierent
sandy habitats, like the sand steppe overgrown with a small
group of alien Ailanthus altissima (Mill.) trees (location of
study site), and open sandy pastures and abandoned vineyards
used for horse and cattle breeding.
2. Virt – VI (47.76030°N, 18.33380°E, 119 m a.s.l.). A large,
intensively used vineyard on sandy soil alternately with
ploughed and grassy rows, treated with herbicides and culti-
vated by a tractor.
3. Mašan Nature Reserve – MA (47.76976°N, 18.31874°E,
131 m a.s.l.). A small hill, formed by a degraded sand dune
that was used as a military training area in the past. It is cur-
rently covered with psammophytic vegetation. e reserve is
the grassiest of the studied sites. Traps were located near the
top of the hill.
Traps were damaged regularly in July, August and October.
4. Bašovský Kopec – BK (47.77646°N 18.32127°E, 126 m
a.s.l.). An old sand quarry surrounded by natural sandy habi-
tats. Traps were located at the edge of the quarry with very
sparse vegetation.
5. Marcelovské piesky Nature Reserve – MP (47.78660°N,
18.26730°E, 111 m a.s.l.).
Sand dunes near the village cemetery. e reserve is aected
by several human activities, e.g. occasional digging creates
small hills with no or sparse vegetation, illegal dumping of
waste and grazing of sheep.
During July and August, several traps dried up, damaging the
material.
Survey of species
Two new spider species for the Slovak fauna were document-
ed from studied sandy habitats of SW Slovakia. During our
study a total of 145 individuals of G. mongolica (61 )), 13
((, 71 juv.) and 59 individuals of H. bohemicus (33 )), 26 ((,
23juv.) were collected and identied.
Gnaphosa mongolica Simon, 1895 (Figs 2-4)
Material. SLOVAKIA: MA: 17. Apr. – 16. May 2019, 1 ),
1 juv.; 16. May – 5. Jun. 2019, 18 )), 2 ((, 1 sub. (; 05. – 25.
Jun. 2019, 3 )), 1 (, 2 sub. )), 1 sub. (; 19. Aug. – 4. Sept.
Fig. 3: Epigyne of Gnaphosa mongolica from Slovakia
Fig. 2: Dorsal view of Gnaphosa
mongolica from Slovakia. a. fema-
le; b. male

Rare ground spiders from sandy habitats in Slovakia 17
2019, 5 sub. ((. BK: 28. Mar. – 17. Apr. 2019, 1 sub. ), 5 sub.
((, 6 juv.; 17. Apr. – 16. May 2019, 4 )), 2 sub. ((, 1 sub. ),
5 juv.; 16. May – 5. Jun. 2019, 19 )), 1 (, 1 juv.; 5. – 25. Jun.
2019, 9 )), 2 ((, 1 sub. ), 4 juv.; 24. Jul. – 19. Aug. 2019,
3((, 2 sub. ((, 3 juv.; 19. Aug. – 4. Sept. 2019, 8 sub. ((,
10juv.; 4. Sept. – 1. Oct. 2019, 1 sub. (; 27. Nov. 2019 – 9.
Jan. 2020, 2 juv. (leg. P. Gajdoš, P. Purgat).
Diagnosis. Gnaphosa mongolica can be confused with G. mus-
corum (L. Koch, 1866), but diers by:
1) absence of the basal spur on the embolus (can be broken o
in G. muscorum);
2) wider and larger epigynal scapus;
3) epigynal ducts extended anteriorly in comparison to closely
spaced ducts in G. muscorum.
Taxonomic notes. Some old records from Europe were
published as G. spinosa Kulczyński, 1897 and G. chaanjoni
Schenkel, 1963. Both names were later synonymised with G.
mongolica by Ovtsharenko et al. (1992).
Measurements. Males. Total length 8.08–10.73 mm. Cara-
pace 3.81–5.14 mm long, 3.14–4.00 mm wide. Opisthosoma
4.27–5.80 mm long, 3.32–3.95 mm wide. Females. Total
length 9.03–13.6 mm. Carapace 4.74–5.18 mm long, 3.63–
3.76 mm wide. Opisthosoma 4.48–8.42 mm long, 3.90–4.68
mm wide.
Distribution (Fig. 5). In Europe it is known from Hungary
(Samu & Szinetár 1999), Serbia (Grbić et al. 2021), Romania
(Weiss & Marcu 1988), Russia (Ponomarev 1981), Ukraine
(Ovtsharenko et al. 1992) and Slovakia (this paper), and also
from Turkey to China (WSC 2022).
Haplodrassus bohemicus Miller & Buchar, 1977 (Figs 6-8,
9a-c, 10)
Material. SLOVAKIA: DM: 17. May – 31. May 2017, 1 );
4. – 17. May 2018, 7 )), 3 (( (leg. P. Gajdoš). VI: 12. Apr. – 4.
May 2018, 1 (; 18. May – 7. Jun. 2018, 1 ). MA: 17. Apr. – 16.
May 2019, 1 ), 1 (, 1 sub. ), 3 sub. ((, 1 juv.; 16. May – 5. Jun.
2019, 1 ), 3 ((, 1 sub. (; 5. – 25. Jun. 2019, 2 ((. MP: 17. Apr.
– 16. May 2019, 7 )), 2 ((, 5 sub. )), 4 sub. ((; 16. May – 5.
Jun. 2019, 10 )), 10 ((, 1 sub. (, 1 juv.; 5. – 25. Jun. 2019, 4 )),
3((; 1. Oct. – 27. Nov. 2019, 1 (, 1 juv. BK: 28. Mar. – 17. Apr.
2019, 2 sub. ((, 17. Apr. – 16. May 2019, 1 ), 1 sub. ), 1 sub.
(; 19. Aug. – 4. Sept. 2019, 1 juv. (leg. P. Gajdoš & P. Purgat).
Comparative material.
H. bohemicus. Czech Republic: Louny, Raná: 50.40655°N,
13.77120°E; 400 m a. s. l., rocky steppe, 4. Oct. – 1. Nov.
1963, 1 ( (paratype, coll. National Museum Prague, Czech
Republic: P6A 5351), leg. J. Buchar (Miller & Buchar 1977);
Bzenec: 48.92657°N, 17.26802°E, 190 m a. s. l., sand dunes,
30. May 1998, 1 ), leg. Bezděčka (Růžička & Bezděčka
2000); Bulgaria: Tsarev Vrah (near peak, above Paril vill.):
41.418°N, 23.664°E, 1100 m a. s. l., 29. Jun. 1937, 1 (, leg.
P. Drensky (Naumova 2009); old border post 1 (above Paril
vill.): 41.413°N, 23.659°E, 1300 m a. s. l., 2. Jul. 1937, 1 (,
leg. Drensky (unpublished); Plovdiv: 42.14883°N, 24.6845°E,
160 m a. s. l., spring 2020, rural habitat on the bank river, 1
gynandromorph, leg. I. Delev (this paper).
H. pseudosignifer. Russia: Altai Republic (Katun' River val-
ley): 50.133°N, 86.083°E, 895 m a. s. l., 22. Jun. – 26. Jul. 1983,
1 ) (paratype, ISEA (Institute of Systematics and Ecology of
Animals, Siberian Branch of the Russian Academy of Sci-
ences, Novosibirsk, Russia) 000.122, ex-BI-1405), leg. H.
Hippa (Marusik et al. 1996); Novosibirsk Area (Karasuk Dis-
trict): 53.72272°N, 77.72887°E, 101 m a. s. l., under birch, 01.
Jul. 2007, 9 )), 1 ( (ISEA 001.1872), leg. I.I. Lubechankii;
53.73167°N, 77.86467°E, 102 m a. s. l., 13. May 2001, 1 )
(ISEA 001.8928), leg. G.N. Azarkina.
Diagnosis. In Slovakia and neighbouring countries, H. bo-
hemicus is most similar to H.signifer (C. L. Koch, 1839), but
males of H. bohemicus dier by:
1) a shorter embolic apophysis and embolus (Fig. 9);
2) the base of the embolic division reaching 1/2 of bulbus
length in comparison to 2/3 in H. signifer;
Fig. 4: Male palp of Gnaphosa
mongolica from Slovakia. a.prola-
teral view; b. ventral view; c. retro -
lateral view

18 A. Šestáková, Ľ. Černecká, M. Naumova, P. Purgat, É. Szita & P. Gajdoš
3) the retrolateral tibial apophysis has a step-like dorsal mar-
gin and is apically oblique, not rounded as in H. signifer;
4) the female epigyne of H.bohemicus has narrower sub-parallel
lateral pockets in comparison to a wide and reniform one;
5) hood is narrower than areola, not wider as in the latter spe-
cies.
Gynandromorphy. One recently collected specimen from
Bulgaria was found to have a lateral gynandromorphy, type
1 sensu Roberts & Parker (1973). Male characters are on
the left (bulbus, darker coxa 1, darker and slimer half of the
opisthosoma) and female on the right (half of the epigyne
with lateral developed pocket, larger opisthosoma). e left
and right eyes are dierent as well (by shape and size) es-
pecially the posterior median ones (Fig. 6d, 7d, 8b, d-e). It
represents the rst record of gynandry for the species and rst
gynandromorph spider found in Bulgaria as well.
Taxonomic notes. While studying the two fundamental
works on the genus Haplodrassus (Kovblyuk et al. 2012, Bos-
mans et al. 2018), we noted the absence of some important di-
agnostic features. For example, it is necessary for determina-
tion to extract the cymbium because from the commonly used
ventral view a certain sclerite behind the embolus is usually
not visible (Fig. 10, sclerite “X”). is sclerite is only rarely
mentioned in the genus Haplodrassus (e.g. Grimm 1985: Dor-
salbereich des Embolus [“a dorsal side of embolus”]; Chatzaki
2021: dorsal apophysis). According to its position, it could
be a terminal apophysis and appears to be important for the
delimitation of related species (e.g. Grimm 1985, Marusik et
al. 1996, Kamura 2007). Unfortunately, in H. bohemicus, this
sclerite was drawn only from an apical view by Kovblyuk et
al. (2012). In addition, slightly dierent angles of the bulbus
may cause a signicantly dierent appearance of sclerites, e.g.
the presence, width or shape of the apical embolic lamella;
the shape and edges of the embolic apophysis; the presence
of a step-like dorsal margin in the retrolateral tibial apophy-
sis. aler (1984) also observed a further diagnostic feature
on the tip of the embolus in the genus Haplodrassus, but its
detailed structure is only apparent by SEM or probably on
microscopic slides.
e Slovak specimens match the original description by
Miller & Buchar (1977), and to the paratype female. Un-
fortunately, the female holotype and male allotype were not
found, probably because they were accidentally moved to an-
other storage box (Petr Dolejš pers. comm.). Compared to
other published drawings or photographs of this species, with
the exception of those by Bosmans et al. (2018), all visible
structures match the drawings by Miller. In our opinion, spec-
imens from Ukraine and Russia described by Kovblyuk et al.
(2012) could represent a variability, or the dierences may be
due to slightly dierent angles shown. e apical view shows
the beak-shaped tip of sclerite X (Kovblyuk et al. 2012: gs 3,
6), as in our Slovak specimens. A comparison of our material
with photographs of this species from North Ossetia (Pono-
marev et al. 2021) and Rostov (unpublished, photographed
by Ponomarev) conrm the presence of such a sclerite at least
in other specimens of H. bohemicus collected from the eastern
part of the distribution. However, it must be said that modern
genetic methods would certainly help to solve the problems of
species identity within this group.
e male specimens in Bosmans et al. (2018), tentatively
designated as H. bohemicus, correspond to the description of
H. pseudosignifer Marusik, Hippa & Koponen, 1996 accord-
ing to:
1) apically rounded retrolateral tibial apophysis without a
step-like dorsal margin;
2) shorter and more massive median apophysis;
3) tip of embolus directed upwards;
4) embolic apophysis with distinct ridge.
Other important characters of the male palp are not easily
comparable with published descriptions and gures. Also, a
comparison with H.pseudosignifer from Russia (including a
paratype from Altai) does not solve the problem, because we
cannot see further details that are only visible after the ex-
traction of the endaparatus or cymbium. e female epigyne
depicted in Bosmans et al. (2018) has a broad sclerotised are-
ola compared to a very narrow one in H. bohemicus. It resem-
bles H. concertor (Simon, 1878), but it diers by a wider hood.
In addition, it is very similar to the drawings of the epigyne
Fig. 5: Distribution of Gnaphosa mongolica (blue triangle) and Haplodrassus bohemicus (yellow circle, red stroke = oldest records; CZ: Miller & Buchar
(1977), BG: Naumova (2009), RU: Ponomarev & Tsvetkov (2006)) in Europe (not all published records from European Russia included)

Rare ground spiders from sandy habitats in Slovakia 19
of H. pseudosignifer from Crimea in Kovblyuk et al. (2012).
How ever, in both cases (Bosmans et al. 2018: g 204; Kov-
blyuk et al. 2012: g. 75) there are also dierences (possible
variability) that distinguish them from the original descrip-
tion (Marusik et al. 1996: g. 69):
1) epigyne has a narrower hood than areola;
2) fovea is narrower with constricted lateral sclerotised edges;
3) areola is rounded at its outer edge.
Unfortunately, without re-examination of the specimens by
Bosmans et al. (2018) we cannot make a nal decision on spe-
cies status of the specimens collected by us. However, it should
be noted that the occurrence of H. bohemicus in Greece has
not been questioned by this assessment. e identication of
the rst country record (Van Keer et al. 2010) was recently
discussed with Johan Van Keer (pers. comm.), and he con-
rmed that two females are identical to the original species
description.
Measurements. Males. Body length 5.21–6.65 mm. Cara-
pace 2.17–2.80 mm long, 1.75–2.17 mm wide. Opisthosoma
3.04–3.72 mm long, 1.71–2.01 mm wide. Females. Body
length 5.63–7.32 mm. Carapace 2.26–2.65 mm long, 1.66–
2.12 mm wide. Opisthosoma 3.37–4.67 mm long, 2.10–2.81
mm wide. Gynandromorph. Body length 7.50 mm. Carapace
3.36 mm long, 2.71 mm wide. Opisthosoma 4.28 mm long,
3.07 mm wide, strongly asymmetric.
Distribution (Fig. 5). Austria (Milasowszky et al. 2008),
Bulgaria (Naumova 2009), Czech Republic (Miller & Buchar
1977), Greece (Van Keer et al. 2010), North Macedonia (Ste-
fanovska et al. 2008), Russia (Esyunin & Tuneva 2020), Ser-
bia (Grbić et al. 2021), Ukraine (Polchaninova & Prokopenko
2019), Hungary (Keresztes et al. 2012, Szinetár pers. comm.)
and now known from Slovakia (this paper).
Comments. e species was described from the Raná Na-
tional Nature Reserve (Czech Republic) in the 1960s, but
during a repeated survey in 1988, Buchar did not nd it there
(Růžička & Bezděčka 2000). e latest ndings in the Czech
Republic come from South Moravian sandy sites (Hula et
al. 2004, Hula 2014). e species were recorded in 1937 in
Bulgaria (Naumova 2009; revision of Drensky’s collection in
2021 by M. Naumova & C. Deltshev) and in 1975 in Russia
(Kalmykia) (Ponomarev & Tsvetkov 2006). It is interesting
that two of the earliest ndings in Europe were collected far
away from the type locality, which could indicate a relict-like
character of the species’ localities and a scattered distribution
in the Western Palearctic.
Notes on ecology and conservation
Phenology
Both species have adult stages that are active from May to
June (Fig. 11), which means that damaged traps in later pe-
riods (see “Material and methods”) did not aect the data
on the phenology of adult specimens. Adults of G. mongolica
were recorded in Slovakia from April to August with the
highest abundance in June. In comparison with other records
from Europe, our results correspond to the central European
ndings (Fig. 11a). Although only a few records from south-
ern Europe have been published, we assume a shift in the oc-
currence of adults to August. In Hungary, G. mongolica over-
winters as juveniles or subadults (Szita et al. 2006). e same
was conrmed in Slovakia by our ndings of immature stages
from September to April.
Adults of H. bohemicus were recorded in Slovakia mainly
from May to June, with the highest abundance in June. Only
one female was sampled late in November, suggesting that
some adults may be overwintering. In comparison with other
records from Europe, our results correspond to ndings from
Central Europe, where H. bohemicus was mostly collected in
May and June (Fig. 11b). On the other hand, in southern Eu-
rope more than 80% of all adults were recorded in June. In
eastern Europe, the highest activity was found in June and July.
A comparison of the phenology of H. bohemicus and H.
signifer collected on our study sites in Slovakia suggests that
H. bohemicus is mostly active in May and June, while H. signi-
fer is signicantly active only in May (Fig. 12).
Fig. 6: Habitus of female and male of Haplodrassus bohemicus from Slovakia and gynandromorph from Bulgaria. a. female, dorsal view; b. eyes asymmetry
of gynandromorph; c. male, dorsal view; d. idem., ventral view; e. gynandromorph, ventral view

20 A. Šestáková, Ľ. Černecká, M. Naumova, P. Purgat, É. Szita & P. Gajdoš
Fig. 7: Epigyne of two females of Haplodrassus bohemicus from Slovakia and one gynandromorph from Bulgaria. a. ventral view; b. extracted and cleared
(hood slightly deformed), ventral view; c. Idem., dorsal view; d. gynadromorph, ventral view
Fig. 8: Left male palp, Haplodrassus bohemicus. a–e. male (Slovakia); d–e. gynandromorph (Bulgaria); a, d. ventral view; b, e. retrolateral view; c. retrolate-
ral tibial apophysis (without scale). Arrow pointed step-like process
Fig. 9: Bulbus with extracted cymbium.
a-c. H. bohemicus (Slovakia); d-f. H. signi-
er (Slovakia); g-i. H. pseudosignifer (Rus-
sia, Novosibirsk); a, d, g. dorsal view; b, e.
ventral view; c, f, i. ventro-prolateral view;
h.retrolateral view

Rare ground spiders from sandy habitats in Slovakia 21
Habitat
Both species inhabit xerothermic steppe habitats. Gnaphosa
mongolica was recorded in Europe mainly from sandy mead-
ows, which also ts to our recent ndings from Slovakia (Fig.
13). In Hungary (Szinetár et al. 2005, Szita et al. 2006) it
seems to be the dominant species on sandy grasslands and
on clearings of Juniperus downs on sand. Several specimens
were found on sand dunes with sparse vegetation in Romania
(Weiss & Marcu 1988) and Russia (Ponomarev et al. 2011,
2017, Ponomarev & Abdurakhmanov 2014). One study site
in Slovakia (Bašovský kopec) was located very close to the
sand pit where this species was most abundantly collected
(Fig. 14). Its retreat was found built under stones, which is
characteristic for ground spiders (Seyyar et al. 2008, Grbić et
al. 2021).
Haplodrassus bohemicus occurs in meadows, steppes and
sand dunes (Kovblyuk et al. 2012). In central Europe, the
majority of ndings derive from sandy grasslands. In Ser-
bia (Southern Europe), the species has been recorded from a
sandy meadow that was dominated by xerophilic steppe veg-
etation on brown sand (Grbić et al. 2021). e ndings from
Slovakia presented herein are also mostly from a sandy grass
steppe, with only two specimens that were recorded from an
intensively used vineyard (see above). However, specimens
from south-eastern and eastern Europe were frequently col-
lected on mesophilic meadows and meadows with a xerother-
mic character (low precipitation, typical plant composition),
which can be explained by the dierent climate compared to
central Europe (Fig. 13). O. Machač observed this species un-
der stones and fallen trunks (ČAS 2021).
Discussion on the red list status
e abundance (in each site and in total) of H. bohemicus
(82 ind.) and G. mongolica (145 ind.) appears to be, at least
Fig. 10: Bulbus of H. bohemicus. a. dorsal view; b. prolateral view; c. ventro-prolateral view; d. ventral view. E = embolus, Ea = embolic apophysis, Ma =
median apophysis, Sc = spermatic canal, X = sclerite “X”
Fig. 5: Comparison of phenology of adults G. mongolica (a) and H. bohemicus (b) for subregions of Europe based on published and unpublished data. C-EU
= Central Europe, E-EU = East Europe, S-EU = Southeast Europe

22 A. Šestáková, Ľ. Černecká, M. Naumova, P. Purgat, É. Szita & P. Gajdoš
partly, inuenced by anthropogenic disturbances of the habi-
tat. Horváth et al. (2015) found that increasing isolation of
grassland fragments decreases the abundance of G. mongolica,
a habitat specialist, which also seems unable to settle down
permanently in adjacent lands altered by human activities.
Grbić et al. (2021) suggest that G. mongolica should be classi-
ed as an endangered species due to the limited geographical
distribution in Europe and its apparently narrow ecological
niche that is under increasing economic pressure. In Slovakia,
G. mongolica inhabits only natural sandy habitats and seems
to avoid sites with anthropogenic inuences (pastures and
vineyard), judging by the fact that all sites are very close to
Fig. 13: Habitat preferences in
each subregion of Europe ex-
pressed as relative abundance of
G. mongolica and H. bohemicus
adults based on data in this work
and other published and unpu-
blished data (see appendix Tab.
S1). C-EU = Central Europe, E-EU
= East Europe; S-EU = South and
Southeast Europe, Dots = step-
pes (xerothermic grasslands with
sparse vegetation; sandy mea-
dows, steppes, sand pits etc.),
black = sandy dunes (very sparse
or no vegetation), vertical lines =
meadows (grasslands with dense
vegetation, incl. grassy habitats
without closer specications of
soil and exposition), grey = other
habitats (vineyards and forests)
Fig. 12: Occurrence over the year of adults of H. bohemicus (black) and H. signifer (orange) in Slovakia, collected with pitfall traps
Fig. 14: Comparison of specimens
numbers of G. mongolica (blue)
and H. bohemicus (yellow) collect-
ed on each study site in Slovakia
from March to November in 2019
(when traps were active on all
localities). Abbreviations: BK =
Bašovský kopec, MA = Mašan, MP
= Marcelovské piesky

Rare ground spiders from sandy habitats in Slovakia 23
each other without signicant migration barriers. In contrast,
H. bohemicus, whose occurrence was most likely overlooked
in Slovakia, seems to cope better with anthropogenic inu-
ence in its habitat, as shown by the high numbers collected at
Marcelovské piesky (Fig. 14) and the presence of the species
at each of the ve Slovak locations that were investigated. A
quick revision of a spider collection from Gönyű (Hungary)
showed that H. bohemicus has been confused with the similar
H. signifer in some cases (Szinetár, pers. comm.). e latter
situation could also have happened in Slovakia, but it should
be noted that these habitats have not been studied intensively
in the past.
Acknowledgements
We are grateful to Peter Dolejš and the National Museum in Prague
(Czech Republic), who provided comparative photographs of the
paratype female of H. bohemicus. Our thanks also go to Robert Bos-
mans, Johan Van Keer and Yuri Marusik for the valuable discussion,
Christo Deltshev, Csaba Szinetár and Norbert Milasowszky, who
provided us important details on their published and unpublished
records and Galina Azarkina for comments and photographs of H.
pseudosignifer. anks to Ivelin Mollov and Ivan Delev (Bulgaria,
University of Plovdiv “Paisii Hilendarski”, Faculty of Biology), who
provided the gynandromorph specimen. Finally, we thank the editor
Tobias Bauer and the reviewers Milan Řezáč and Mykola Kovblyuk.
is research was funded by the Operational Program of Integrated
Infrastructure: DNA barcoding of Slovakia (SK-BOL), as a part of
the international initiative International Barcode of Life (iBOL)
(ITMS2014+313021W683) and by the Slovak Grant Agency of the
Ministry of Education, Science, Research and Sport of the Slovak
Republic, as part of project VEGA No. 2/0135/22 (Research of specic
landscape elements of bio-cultural landscape in Slovakia).
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Electronic supplement
Tab. S1: compilation (.xls) of additional data and records of Gnaphosa
mongolica and Haplodrassus bohemicus in the literature