A total of 667 specimens of pseudoscorpions, belonging to 15 species and four families were collected during 2014. Samples were taken from leaf litter at five localities in the Western Carpathians in the Slovak Karst National Park, Slovakia. The most abundant families were Neobisiidae (381 specimens) and Chthoniidae (275 specimens). Furthermore ten chernetid specimens and a single cheliferid deutonymph were recorded. Four of the 15 pseudoscorpion species were recorded in the Slovak Karst for the first time.
The Slovak Karst National Park is one of the most valuable areas in Slovakia in terms of biodiversity (Tomaskinova & Tomaskin 2013). The territory of the National Park consists of the Koniarská, Plešivská, Silická, Zádielská and Jasovská plateaus, the Horný and Dolný hills, and is divided into gorges and valleys. It is one of the largest karst areas in Central Europe. The area of the National Park is about 346 km2. In 1977, the territory of the Slovak Karst was included in the international network of biosphere reserves within the UNESCO Programme “Man and the Biosphere” ( https://en.unesco.org/biosphere/wnbr). The karst phenomenon, together with other abiotic factors, was of great importance in the development of animal communities in this area. Pseudoscorpions from the Slovak Karst have not been systematically studied. The first data from the territory of the Slovak Karst were published by Daday (1888) and later by Krumpál & Kiefer (1981). More recently, additional data by Papáč et al. (2007), Christophoryová (2009, 2010), Christophoryová et al. (2011b) and Jászayová & Christophoryová (2019) have been published. The majority of faunistic data concern the species Neobisium (Blothrus) slovacum (Ducháč 1996, Ducháč & Mlejnek 2000a, 2000b, Christophoryová 2010, Červená et al. 2019). We know very little about the ecology of pseudoscorpions in Slovakia, as they are often only found in random collections. The aim of the present paper was to complement the current faunistic inventory of pseudoscorpions from five localities in the Sites of Community Importance (SCIs) of Slovak Karst in the Western Carpathians, obtained by the method of sifting the forest leaf litter.
Material and methods
Study area
Research on the pseudoscorpions was carried out at monthly intervals (May–September) during the year 2014 at five localities (Fig. 1): Brzotín Rocks (BR), Pod Fabiánkou (F), Drieňovec (D), Hrušovská lesostep forest-steppe (HL) and Jasovské dubiny (JD).
BR: Brzotín Rocks National Nature Reserve is situated at 290–679 m a.s.l. and located on the northwestern slopes of Silická plateau. The area with 433.78 ha is the largest natural reserve in the Slovak Karst. The geological substrate consists mainly of light Wetterstein limestones. Characteristic of this area are the natural forest associations of thermophilic oak-beech (Fago-Quercetum) and linden maples (Tilio-Aceretum). Sampling sites were located at 48.58801°N, 20.49075°E.
F: Pod Fabiánkou Nature Reserve is situated at 490–510 m a.s.l. and is characterized by the occurrence of species characteristic for meadows and pastures, into which thermophilic species from the surrounding xerothermic phytocoenoses penetrate. In the central part of the area forests grow, especially oak-hornbeam (Querco-Carpinetum) and thermophilic oak or beech forests (Fago-Quercetum). Sampling sites were located at 48.55890°N, 20.54735°E.
D: Drieňovec National Nature Reserve is situated at 430–804 m a.s.l., the substrate is built of karst limestones. Forest and rock plant associations on limestone and non-limestone type substrate are present. The vegetation cover consists of beech-oak (Fago-Quercetum) and linden-maple forests (Tilio-Aceretum) and in the southern part of the reserve, there are partially deforested rocky slopes of forest-steppe and xerothermophilic steppe. Sampling sites were located at 48.64085°N, 20.68926°E.
HL: Hrušovská lesostep forest-steppe National Nature Reserve – declared as a protected area in 1984 – is situated at 265–525 m a.s.l. and it has an area of 40.85 ha and represents a typical forest-steppe with xerothermic vegetation of downy oak (Quercus pubescens Willd.) and manna ash (Fraxinus ornus L.). Turkey oak (Quercus cerris L.) also occasionally occurs. Sampling sites are located at 48.59778°N, 20.63278°E.
JD: Jasovské dubiny National Nature Reserve, which is the oldest nature reserve in the Slovak Karst, is situated at 260–340 m a.s.l.The following types of forest are represented there: beech-hornbeam (Fago-Carpinetum), oak-beech (Fago-Quercetum) and oak-beech with a transition to the oak forest with a typical sessile oak (Quercus petraea (Matt.) Liebl.). Sampling sites are located at 48.67851°N, 20.96929°E.
Sampling design
From each locality a volume of about 5 litres of forest leaf litter was sieved. Per 10 m2 five sieving samples were made. Samples were extracted using xereclectors of Moczarski´s type (Besuchet et al. 1987). All pseudoscorpions were studied on temporary slide mounts using lactic acid and then rinsed in the water and returned to 70 % alcohol. The studied material was collected by both authors and identified by the first author. Pseudoscorpion specimens were identified using the key of Christophoryová et al. (2011c). If not otherwise indicated, nomenclature follows the catalogue of Pseudoscorpions of the World (Harvey 2013), Zaragoza (2017) and the updated checklist of pseudoscorpions by Červená et al. (2020). The material is deposited in the collection at the Natural History Department of Šariš Museum in Bardejov, Slovakia.
Results
In total, 667 specimens belonging to 15 species from four families were identified. The most numerous family was Neobisiidae (381 specimens, six taxa) and the second most abundant family was Chthoniidae (275 specimens, six taxa); Chernetidae (10 specimens, three taxa) was less abundant and the family Cheliferidae was represented only by a single deutonymph. The list of taxa collected is given below with date and number of specimens of each sex or developmental stage (tritonymph: Tn, deutonymph: Dn, protonymph: Pn).
Family Chthoniidae Daday, 1888
Chthonius carinthiacus Beier, 1951
Material examined. BR: 5 ♂♂, 5. May 2014; 14 ♂♂, 2 ♀♀, 23. Jun. 2014; 8 ♂♂, 13 ♀♀, 15. Jul. 2014; 8 ♂♂, 2 ♀♀, 5. Aug. 2014; 12 ♂♂, 5 ♀♀, 9. Sep. 2014; F: 7 ♀♀, 26. May 2014; 11 ♂♂, 14 ♀♀, 25. Jun. 2014; 1 ♂, 2 ♀♀, 5. Aug. 2014; 4 ♂♂, 2 ♀♀, 9. Sep. 2014; D: 1 ♂, 1 ♀, 27. May 2014; 5 ♂♂, 1 ♀, 24. Jun. 2014; HL: 3 ♂♂, 6. May 2014; 2 ♂♂, 25. Jun. 2014; JD: 2 ♀♀, 28. May 2014. Distribution. Austria, Czech Republic, Hungary, Italy, Slovakia, Slovenia, Switzerland (Harvey 2013, Červená et al. 2020).
Remarks. Chthonius carinthiacus is an epigean species. It was previously reported from Austria (Beier 1951) and later from several further countries. In Slovakia C. carinthiacus was reported for the first time from Slovenské rudohorie and from the Slovak Karst by Christophoryová et al. (2011b). Additional records from the Slovak Karst were also published recently by Jászayová & Christophoryová (2019).
Chthonius heterodactylus Tömösváry, 1882
Material examined. BR: 2 ♂♂, 5. May 2014; 3 ♀♀, 9. Sep. 2014; F: 2 ♀♀, 25. Jun. 2014; D: 1 ♂, 6. May 2014; 4 ♀♀, 10. Sep. 2014; HL: 2 ♂♂, 16. Jul. 2014; JD: 1 ♂, 6. May 2014; 1 ♂, 11. Sep. 2014.
Distribution. Czech Republic, Hungary, Poland, Romania, Slovakia (Červená et al. 2020), Ukraine (Gardini 2014).
Remarks. This is an epigeic species distributed across the whole Carpathians (Ducháč et al. 2007, Gardini 2014). It has previously been recorded in the Slovak Karst (Mock et al. 2009, Papáč et al. 2009, Jászayová & Christophoryová 2019).
Ephippiochthonius boldorii (Beier, 1934)
Material examined. BR: 1 ♀, 5. Aug. 2014; D: 3 ♀♀, 5. Aug. 2014; HL: 5 ♀♀, 16. Jul. 2014; 4 ♂♂, 3 ♀♀, 6. Aug. 2014; JD: 2 ♂♂, 4 ♀♀, 6. Aug. 2014 (rev. J. Christophoryová).
Distribution. Austria, Croatia, Germany, Italy, Slovakia, Slovenia and Switzerland (Muster et al. 2004, Zaragoza 2017, Červená et al. 2020).
Remarks. In Slovakia, this species was recorded in oak-hornbeam forest at lower altitudes from the Malé Karpaty Mts. and in the Trnavská pahorkatina Highland. These localities have habitats and conditions suitable for the occurrence of stable and relatively numerous populations of this species (Christophoryová & Krumpál 2005, 2007, Christophoryová 2013). Ephippiochtonius boldorii is an epigeic species occurring in oak-hornbeam forests (Muster et al. 2004, Christophoryová & Krumpál 2007).The highest localities in the Alps exceed 1500 m a.s.l., but on the other hand, this species was also recorded at sea-level in Venice. The species occurs predominantly in leaf litter in mesophilic habitats (Muster et al. 2004). One female was recorded from compost heaps (Christophoryová et al. 2017b). We found E. boldorii in all studied localities except Pod Fabiánkou.
Ephippiochthonius fuscimanus (Simon, 1900)
Material examined. BR: 1 ♂, 1 ♀, 5. May 2014; HL: 3 ♂♂, 2 ♀♀, 28. May 2014; 2 ♂♂, 6. Aug. 2014; 5 ♂♂, 1 ♀, 10. Sep. 2014. Distribution. Austria, Czech Republic, Georgia, Germany, Hungary, Italy, Poland, Slovakia and Turkey (Harvey 2013, Červená et al. 2020).
Remarks. Ephippiochthonius fuscimanus was recorded in Slovakia from the oak-hornbeam forest of the Malé Karpaty Mts (Christophoryová & Krumpál 2007, Christophoryová et al. 2012, Christophoryová 2013), from the Slovak Karst in front of cave entrances (Jászayová & Christophoryová 2019) and from bird nests in different parts of Slovakia (Christophoryová et al. 2011a). One deutonymph and one female of the species were recorded in tree hollows and dead wood in western Slovakia (Christophoryová et al. 2017a).
Ephippiochthonius tetrachelatus (Preyssler, 1790)
Material examined. D: 1 ♀, 10. Sep. 2014; HL: 14 ♂♂, 5 ♀♀, 25. Jun. 2014; JD: 2 ♀♀, 6. Aug. 2014.
Distribution. Ephippiochthonius tetrachelatus is widely distributed, known from the Palaearctic region, Argentina, Cuba, Seychelles and the USA (Harvey 2013); in Europe it occurs in Austria, Czech Republic, Germany, Hungary, Poland, Slovakia, Slovenia and Switzerland (Červená et al. 2020).
Remarks. In Slovakia, Ephippiochthonius tetrachelatus is one of the most common species, inhabiting different types of habitat and preferring dry or slightly moist conditions (Christophoryová 2013, Christophoryová et al. 2017b). This species was previously confirmed from a cave entrance in the Slovak Karst National Park, from Ardovská cave (Jászayová & Christophoryová 2019).
Mundochthonius carpaticus Rafalski, 1948
Material examined. BR: 2 ♀♀, 5. Aug. 2014; 2 ♀♀, 9. Sep. 2014; F: 5 ♀♀, 5. Aug. 2014; 1 ♂, 5 ♀♀, 9. Sep. 2014; D: 1 ♀, 6. May 2014; 2 ♀♀, 5. Aug. 2014; JD: 1 ♂, 6. May 2014.
Distribution. Czech Republic, Hungary, Poland, Slovakia, Ukraine (Harvey 2013, Červená et al. 2020).
Remarks. This species is a Carpathian faunal element known from the whole of Slovakia. It occurs in leaf litter of mixed forests, in humus, under stones, pieces of wood (Christophoryová et al. 2017a, Christophoryová & Krajčovičová 2019), and in the Slovak Karst it was recorded in front of cave entrances (Mock et al. 2009, Papáč et al. 2009, Jászayová & Christophoryová 2019).
Undetermined material from the family Chthoniidae Daday, 1888
BR: 5 Tn, 9. Sep. 2014; 12 Tn, 5. Aug. 2014; F: 2 Dn, 25. June 2014; 1 Dn, 3 Tn, 9. Sep. 2014; 9 Tn, 5. Aug. 2014; D: 1 Pn, 24. June 2014; 2 Tn, 5. Aug. 2014; 1 Dn, 10. Sep. 2014; HL: 1 Pn, 1 Tn, 28. May 2014; 3 Dn, 5 Tn, 6. Aug. 2014; 2 Tn, 10. Sep. 2014; JD: 1 Tn, 6. May 2014; 4 Tn, 6. Aug. 2014.
Family Neobisiidae Chamberlin, 1930
Neobisium (Neobisium) brevidigitatum (Beier, 1928)
Material examined. BR: 2 ♂♂, 6 ♀♀, 23. Jun. 2014; 1 ♀, 15. Jul. 2014; 1 ♀, 5. Aug. 2014; 4 ♂♂, 7 ♀♀, 9. Sep. 2014; F: 5 ♀♀, 15. Jul. 2014; 2 ♂♂, 3 ♀♀, 5. Aug. 2014; D: 1 ♂, 2 ♀♀, 24. Jun. 2014; 2 ♀, 15. Jul. 2014; 2 ♂♂, 16 ♀♀, 10. Sep. 2014; HL: 1 ♀, Sep. 2014.
Distribution. Georgia, Poland, Romania, Slovakia (Harvey 2013), Hungary (Novák 2015), Ukraine (Novák 2017).
Remarks. This species was first described from Romania (Beier 1928). The presence of Neobisium (N.) brevidigitatum was confirmed in the High Tatra Mts, Great Fatra Mts and National Park of Pieniny (Krumpál 1980, Šťáhlavský & Dolejš 2019).
Neobisium (Neobisium) carcinoides (Hermann, 1804)
Material examined. BR: 1 ♀, 23. Jun. 2014; F: 1 ♀, 9. Sep. 2014.
Distribution. This species is distributed in West, Central, North (except Lithuania),East (except Belarus and Moldovia) and South (except Albania) Europe, in Africa (Algeria, Morocco, Tunisia and Kenya) and Asia (India and Kazakhstan) (Harvey 2013, Červená et al. 2020).
Remarks. Neobisium (N.) carcinoides is a eurytopic, mainly epigeic species and is by far the most frequently recorded Neobisium species in Central Europe (Drogla & Lippold 2004, Christophoryová 2013). In our study, we found only two females. The occurrence of this species in the Slovak Karst was also reported by Jászayová & Christophoryová (2019).
Neobisium (Neobisium) carpaticum Beier, 1935
Material examined. BR: 1 ♀, 15. Jul. 2014; D: 1 ♀, 5. Aug. 2014.
Distribution. Poland, Slovakia (Červená et al. 2020).
Remarks. The species is regarded as a Carpathian element (Beier 1963, Christophoryová et al. 2012). The typical habitat of this species is oak-hornbeam forest (Krumpál & Krumpálová 2003, Christophoryová & Krumpál 2007) and cave entrances (Krumpál 2000). From the Slovak Karst, N. (N.) carpaticum was recorded for the first time in front of the entrance to Silická ľadnica cave (Jászayová & Christophoryová 2019).
Neobisium (Neobisium) crassifemoratum (Beier, 1928)
Material examined. BR: 1 ♀, 5. May 2014; 12 ♀♀, 23. Jun. 2014; 2 ♂♂, 15. Jul. 2014; 3 ♀♀, 5. Aug. 2014; 9 ♂♂, 7 ♀♀, 9. Sep. 2014; F: 3 ♂♂, 3 ♀♀, 25. Jun. 2014; 1 ♀, 15. Jul. 2014; 1 ♂, 4 ♀♀, 5. Aug. 2014; 10 ♂♂, 20 ♀♀, 9. Sep. 2014; D: 1 ♂, 6. May 2014; 2 ♀♀, 24. Jun. 2014; 8 ♂♂, 5 ♀♀, 10. Sep. 2014; HL: 1 ♂, 6. May 2014; 2 ♂♂, 3 ♀♀, 16. Jul. 2014; 1 ♂, 10. Sep. 2014; JD: 2 ♀♀, 28. May 2014; 1 ♂, 1 ♀, 25. Jun. 2014; 1 ♀, 6. Aug. 2014.
Distribution. Central and Eastern Europe, also Turkey, Georgia, Azerbaijan (Harvey 2013), Lithuania (Krajčovičová et al. 2018).
Remarks. This species occurs mostly in the leaf litter (Krumpál & Krumpálová 2003). In Slovakia N. (N.) crassifemoratum was recorded in the Great Fatra Mts (Krumpál 1980) and National Park of Poloniny (Krumpál & Krumpálová 2003). A few specimens were also recorded from bird nests (Fenďa et al. 1998, Christophoryová et al. 2011a). From the Slovak Karst, N. (N.) crassifemoratum was recently recorded just outside the entrance to a cave (Jászayová & Christophoryová 2019).
Neobisium (Neobisium) erythrodactylum (L. Koch, 1873)
Material examined. BR: 2 ♀♀, 23. Jun. 2014; F: 4 ♀♀, 5. Aug. 2014; D: 1 ♂, 10. Sep. 2014; JD: 2 ♂♂, 1 ♀, 16. Jul. 2014; 1 ♂, 11. Sep. 2014.
Distribution. This species is widespread in Central Europe. It occurs also in Armenia, Azerbaijan, Bosnia and Herzegovina, Croatia, Georgia, Greece, Iran, Italy, Romania, Serbia, Turkey and Ukraine (Harvey 2013, Červená et al. 2020).
Remarks. Neobisium (N.) erythrodactylum is a typical epigeic species living in forest leaf litter and in the upper part of the soil (Krajčovičová & Christophoryová 2014). The species was reported from numerous faunistic surveys in Slovakia (Christophoryová et al. 2012).
Neobisium (Neobisium) sylvaticum (C. L. Koch, 1835)
Material examined. F: 1 ♀, 5. Aug. 2014; HL: 1 ♀, 6. May 2014.
Distribution. This species is widespread throughout Europe except for the United Kingdom, North Europe and European Russia. It occurs also in Armenia, Georgia and Turkey (Harvey 2013).
Remarks. Neobisium (N.) sylvaticum occurs in forest habitats in the leaf litter and moss, but is also known from the entrances of caves and was found under trees or in bird nests (Krumpál & Krumpálová 2003, Drogla & Lippold 2004, Christophoryová 2010, Christophoryová et al. 2017a). Some data from the Slovak Karst were already published by Papáč et al. (2009) and Jászayová & Christophoryová (2019).
Undetermined material from family Neobisiidae Chamberlin, 1930
BR: 3 Tn, 5. May 2014; 12 Pn, 10 Dn, 6 Tn, 23. Jun. 2014; 2 Pn, 2 Dn, 5 Tn, 15. Jul. 2014; 1 Pn, 3 Dn, 8 Tn, 5. Aug. 2014; F: 4 Tn, 26. May 2014; 2 Dn, 6 Tn, 25. Jun. 2014; 3 Pn, 3 Tn, 15. Jul. 2014; 1 Pn, 16 Dn, 6 Tn, 5. Aug. 2014; 1 Pn, 24 Dn, 3 Tn, 9.Sep. 2014; D: 1 Tn, 6. May 2014; 3 Pn, 5 Tn, 24. Jun. 2014; 1 Dn, 2 Tn, 15. Jul. 2014; 1 Pn, 6 Dn, 9 Tn, 5. Aug. 2014; 1 Pn, 4 Dn, 3 Tn, 10. Sep. 2014; HL: 3 Tn, 6.May 2014; 1 Tn, 6. Aug. 2014; JD: 3 Tn, 28. May 2014; 1 Tn, 16. Jul. 2014; 2 Dn, 2 Tn, 6. Aug. 2014; 3 Tn, 11.Sep. 2014.
Family Chernetidae Menge, 1855
Chernes cimicoides (Fabricius, 1793)
Material examined. BR: 1 ♂, 23. Jun. 2014; 2 ♀♀, 9. Sep. 2014. Distribution. Central Europe (Červená et al. 2020), Eurasia (Harvey 2013).
Remarks. In Slovakia, this species is common in tree hollows and under tree bark (Krajčovičová & Christophoryová 2014) and can also be found in anthills (Drogla & Lippold 2004) or bird nests (Krištofík et al. 2002). We found three specimens of C. cimicoides at the Brzotín Rocks.
Chernes similis (Beier, 1932)
Material examined. D: 4 ♂♂, 10. Sep. 2014.
Distribution. Austria, Bulgaria, Czech Republic, Hungary, Macedonia, Montenegro, Poland, Romania, Slovakia, Turkey (Novák 2012, Harvey 2013, Christophoryová 2013, Červená et al. 2020), Lithuania (Krajčovičová et al. 2020).
Remarks. A few records of this species are known from Slovakia (Krumpálová & Krumpál 1993). Chernes similis was found by Šťáhlavský & Dolejš (2019) in Domica in the Slovak Karst. In our study, we only found four male specimens at the Drieňovec site.
Pselaphochernes scorpioides (Hermann, 1804)
Material examined. F: 1 ♀, 5. Aug. 2014; HL: 1 ♂, 10. Sep. 2014.
Distribution. This species is widespread across Europe (Červená et al. 2020) and in some countries in Asia and Africa (Harvey 2013).
Remarks. Pselaphochernes scorpioides occurs in Slovakia mainly in leaf litter, decomposing material, tree hollows, bird nests, decaying wood and under tree bark (Beier 1963, Krumpál & Cyprich 1988, Krištofík et al. 2009, Christophoryová 2013, Krajčovičová & Christophoryová 2014, Kaňuchová et al. 2015, Christophoryová et al. 2016, Christophoryová et al. 2017b) or in ant nests (Jászayová et al. 2015).
Undetermined material from family Chernetidae Menge, 1855
F: 1 Tn, 25. June 2014.
Undetermined material from family Cheliferidae Risso, 1827
F: 1 Dn, 9. Sep. 2014.
Discussion
The present paper reports faunistic data on the occurrence of 15 species of pseudoscorpions from the Slovak Karst National Park. We present four new species for the study area. A total of 20 pseudoscorpion species are presently known from the Slovak Karst, which is almost 39 % of the fauna of Slovakia (Tab. 1). The area shows a high species richness in the leaf litter – 15 species at six localities in our study as compared to 12 species at 26 localities in southwestern Slovakia (Christophoryová 2013). The results were obtained by similar methods in similar habitats, mainly in oak and oak-hornbeam. The published results originate from intensive research of cave habitats, the area around cave entrances and warm oak-beech forests. Further research should focus on different habitats and microhabitats prevalent in the territory of the Slovak Karst, such as decayed wood, tree hollows, compost, ant, bird and mammalian nests, or different wetlands, including peat bogs and fens. Generally, the Slovak Karst represents one of the most valuable areas in terms of biodiversity for the whole of Slovakia. For example, in beetle research several new species for the fauna of Slovakia were found in this area (Jászay 1998, Jászay & Hlaváč 2016, Jászay & Jászayová 2018).
Tab. 1:
List of known pseudoscorpions from the Slovak Karst (original name of locality in square brackets)
Acknowledgements
The authors express their gratitude to Jana Christophoryová (Bratislava) for revising the species difficult to identify,especially Ephippiochthoniusboldorii.We are also grateful Katherine R.Day (Tallahassee) for the correction of English language and Patricia Antlová (Vienna) for the German correction and two anonymous reviewers for their constructive comments.