The genus Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) in Hungary

Winkler, Dániel

doi:10.48311/jibs.12.01.81

The genus Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) in Hungary

Document Type : Checklist

Author

Dániel Winkler

Institute of Wildlife Management and Wildlife Biology, University of Sopron, 9400 Sopron, Hungary

10.48311/jibs.12.01.81

Abstract

A checklist with distribution data of the collembolan genus Pseudosinella of Hungary is presented in this paper. In addition to available relevant references, unpublished data from newly collected material is also included. To date, the occurrence of 23 species of this genus has been confirmed in Hungary, for which a dichotomous key is also provided. Since the publication date of the Hungarian Collembola checklist, a total of nine additional Pseudosinella species have been recorded in the country. In this study, P. edax Gisin, 1967, P. noseki Rusek, 1985, and P. thibaudi Stomp, 1977 are cited for the first time. Up to now, three species of Pseudosinella, namely P. argentea Loksa, 1960 nec Folsom, P. csafordensis Winkler & Mateos, 2018, and P. dungeri Winkler & Mateos, 2018, are recognized as endemic to Hungary.

Graphical Abstract

The genus Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) in Hungary

Keywords

springtails

distribution

endemism

fauna

taxonomy

Subjects

Faunistics

The genus Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) in Hungary

Daniel Winkler

Institute of Wildlife Biology and Management, University of Sopron, 9400 Sopron, Hungary

ABSTRACT. A checklist with distribution data of the collembolan genus Pseudosinella of Hungary is presented in this paper. In addition to available relevant references, unpublished data from newly collected material is also included. To date, the occurrence of 23 species of this genus has been confirmed in Hungary, for which a dichotomous key is also provided. Since the publication date of the Hungarian Collembola checklist, a total of nine additional Pseudosinella species have been recorded in the country. In this study, P. edax Gisin, 1967, P. noseki Rusek, 1985, and P. thibaudi Stomp, 1977 are cited for the first time. Up to now, three species of Pseudosinella, namely P. argentea Loksa, 1960 nec Folsom, P. csafordensis Winkler & Mateos, 2018, and P. dungeri Winkler & Mateos, 2018, are recognized as endemic to Hungary.

KEYWORDS: Distribution, Endemism, Fauna, Springtails, Taxonomy

Citation: Winkler, D. (2026) The genus Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) in Hungary. Journal of Insect Biodiversity and Systematics, 12 (01), 81–95.

INTRODUCTION

Pseudosinella Schäffer, 1897 is one of the largest and most widespread collembolan genera of the family Entomobryidae Tömösváry, 1882, represented by 351 described species worldwide and by around 200 species in Europe (Bellinger et al. 1996–2025). It can be separated from its ancestor genus, Lepidocyrtus, by the number of eyes: 8+8 in Lepidocyrtus and fewer (0–6+0–6) in Pseudosinella (Ponge 2000). The species of this genus inhabit a wide range of habitats from caves (e.g., Gisin & Gama 1969; Arbea 2013; Cipola et al. 2020; Kováč et al. 2023) through diverse forests (e.g., Jordana & Baquero, 2005; Sławska et al. 2017) to open grassland habitats (e.g., Arbea & Jordana 1990; Traser et al. 2006; Winkler & Mateos 2018). Hungary, situated in the Carpathian Basin, is remarkably rich in diverse habitats and, thus, also in fauna diversity, including a high number of endemic species (Varga 1995). Nevertheless, the Collembola checklist of Hungary only lists 14 Pseudosinella species from the country (Dányi & Traser, 2007). This paper aims to present an updated checklist of Pseudosinella species by providing newly discovered data from earlier references and material published since the checklist by Dányi & Traser (2007) was issued. Unpublished data, including new records and species new to the Hungarian Pseudosinella fauna, are also included. A dichotomous key is also provided for the Pseudosinella species of Hungary. Alongside earlier works (e.g., Fiera 2013; Kováč et al. 2023), the present study aims to contribute to the exploration of the apparently rich Pseudosinella fauna of the Central European region.

MATERIAL AND METHODS

For the present work, all available relevant references on the Collembola genus Pseudosinella from Hungary were considered. Unpublished data from newly collected material are also presented here. The newly collected specimens of Pseudosinella were cleared using Nesbitt’s fluid and mounted on slides in Hoyer as a medium (Jordana 2012). The slides were examined under a Leica^® DM2500 LED microscope with conventional bright light and phase contrast. The studied material is preserved at the Institute of Wildlife Biology and Management, University of Sopron, in the author’s collection. Identifications were carried out with the help of the computer-assisted Delta identification key using the combination of chaetotaxic and other characters, originally designed by Christiansen et al. (1990) and regularly updated by Jordana et al. (2024). Figure 1 illustrates the general habitus of Pseudosinella and highlights the key taxonomic characters used in the dichotomous key for the Hungarian species. For each species, the following is reported: a) the scientific name, the name of the author, and the year of description; b. the original name with its bibliographic quotation, and type locality; c. world distribution; d. distribution in Europe; e. localities where the species was recorded in Hungary (Fig. 2); and f. ecological comment based on the collected literature.

Abbreviations. Names of the European countries: Albania—AL; Andorra—AD; Austria—AT; Azores (Portugal)—AZ; Belarus—BY; Belgium—BE; Bosnia and Herzegovina—BA; Great Britain—GB; Bulgaria—BG; Canary Islands (Spain)—IC; Corsica (France)—CO; Crete (Greece)—CT; Croatia—HR; Cyprus—CY; Czech Republic—CZ; Denmark—DK; Estonia—EE; Finland—FI; France—FR; Germany—DE; Greece—GR; Hungary—HU; Iceland—IS; Ireland—IE; Italy—IT; Latvia—LV; Lithuania—LT; Luxembourg—LU; Macedonia—MK; Malta—MT; Moldova—MD; Montenegro—ME; Norway—NO; Poland—PL; Portugal—PT; Romania—RO; Serbia—RS; Slovakia—SK; Slovenia—SL; Spain—ES; Sweden—SE; Switzerland—CH; The Netherlands – NL; Ukraine – UA. Symbols: a.s.l.—above sea level, Th—thoracic tergite, Abd—abdominal tergite, I–VI—segment numbers, pse—pseudoporus.

Figure 1. General habitus of Pseudosinella sp., showing taxonomically important features used in the key to the Hungarian species. A. Labial triangle; B. Detail of foot complex and distal part of tibiotarsus; C. Abd II dorsolateral chaetotaxy; D. Abd IV trichobothrial complex with accessory chaeta s. (Original drawing).

Figure 2. Geographical map of Hungary with localities used in the description of the distribution of Pseudosinella species, with sampling points for species new to the Hungarian fauna. The map was generated using the topographic map by Tschubby (2025) as background.

RESULTS

Class Collembola Lubbock, 1870

Order Entomobryomorpha Börner, 1913

Superfamily Entomobryoidea Womersley, 1934

Family Entomobryidae Tömösváry, 1882

Subfamily Lepidocyrtinae Wahlgren, 1906

Genus Pseudosinella Schäffer, 1897

Type species. Tullbergia immaculata Lie-Pettersen, 1896.

Pseudosinella aelleni da Gama, 1973

Pseudosinella aelleni da Gama, 1973:52. Type locality: Germany.

World distribution. Europe.

Distribution in Europe. AT, DE, HU.

Distribution in Hungary. Moson Plain (Winkler & Traser 2012a), Sopron Mts. (Traser 2008).

Ecological note. Silvicolous, troglophilous (Kampmann 1987; Deharveng & Thibaud 1989).

Pseudosinella aggtelekiensis (Stach, 1929)

Lepidocyrtus (Pseudosinella) aggtelekiensis Stach, 1929:296. Type locality: Hungary.

World distribution. Europe.

Distribution in Europe. HU, SK, RO.

Distribution in Hungary. Aggtelek Karst (Stach 1929; Dudich 1930, 1932; Kovács 1953, 1954; Loksa 1961; Traser 1999).

Ecological note. Caverniculous (Dudich 1932; Kovács 1954).

Pseudosinella alba (Packard, 1873)

Lepidocyrtus albus Packard, 1873:37. Type locality: USA.

World distribution. Cosmopolitan.

Distribution in Europe. AL, AT, BA, BE, BG, CH, CO, CZ, DE, DK, EE, ES, FI, FR, GB, HR, HU, IE, IS, IT, LV, MD, ME, MT, NL, NO, PL, PT, RO, RS, SK, SE, UA.

Distribution in Hungary. Aggtelek Karst (Loksa 1966; Traser 1999), Bakony Mts. (Loksa 1966; Loksa & Rubio 1966; Traser 2010), Balatonfelvidék (Loksa 1960, 1966), Börzsöny Mts. (Loksa 1966; Dunger 1975), Buda Mts. (Loksa 1956b, 1966), Bükk Mts. (Loksa 1956a, 1962, 1966, 1977), Cserhát (Loksa 1978b), Fertő-Hanság (Traser 2002a, 2003; Traser et al. 2006; Winkler & Tóth, 2012), Gerecse Mts. (Loksa 1966; Traser 2006), Gödöllő Hills (Harta et al. 2018, 2021), Keszthely Mts. (Loksa 1966), Kiskunság (Traser & Horváth–Szováti 2004, 2006; Palkó et al. 2020), Kőszeg Mts. (Szent-Ivány 1940), Mezőföld (Traser et al. 2011; Gergócs et al. 2022); Moson Plain (Winkler & Traser 2012a; Szigeti et al. 2022), Mt. Somló (Loksa 1966), Nyírség (Loksa 1991), Őrség (Traser 1995), Pilis Mts. (Loksa 1966), Sopron Mts. (Traser 1981, 2002a), Szigetköz (Traser 1997), Tétény Plateau (Loksa & Loksa 1996a), Vas-Sopron Plain (Traser 1981), Veszprém-Devecser Graben (Winkler 2014; Winkler et al. 2018), Vértes Mts. (Loksa 1966), Visegrád Mts. (Loksa 1966).

Ecological note. Eurytopic, edaphic species, occurring in a wide range of habitats: forest litter and moss, meadows, seashores, cultivated soils, and caves (Dallai & Malatesta 1982; Fjellberg 2007).

Pseudosinella argentea Loksa, 1960 nec Folsom

Pseudosinella argentea Loksa, 1960:262. Type locality: Hungary.

World distribution. Europe.

Distribution in Europe. HU.

Distribution in Hungary. Balatonfelvidék (Loksa 1960).

Ecological note. Caverniculous (Loksa 1960).

Pseudosinella binoculata Kseneman, 1935

Pseudosinella binoculata Kseneman, 1935:6. Type locality: Czech Republic.

World distribution. Europe.

Distribution in Europe. AT, BE, CH, CZ, DE, ES, HU, PL, SK, UA.

Distribution in Hungary. Börzsöny Mts. (Dunger 1975), Dráva Plain (Loksa 1978a).

Ecological note. In caves, forests (Kseneman 1935; Vedovini 1968; Dunger 1975).

Pseudosinella bohemica Rusek, 1979

Pseudosinella bohemica Rusek, 1979: 255. Type locality: Czech Republic.

World distribution. Europe.

Distribution in Europe. AT, BG, CZ, HU.

Distribution in Hungary. Veszprém-Devecser Graben (Winkler et al. 2012; Winkler 2014), Bakony Mts. (Traser 2010).

Ecological note. Silvicolous (Rusek 1979; Traser 2010).

Pseudosinella csafordensis Winkler & Mateos, 2018

Pseudosinella csafordi Winkler & Mateos, 2018:348. Type locality: Hungary.

World distribution. Europe.

Distribution in Europe. HU.

Distribution in Hungary. Vas-Sopron Plain (Winkler & Mateos 2018).

Ecological note. Silvicolous, hygrophilous (Winkler & Mateos 2018).

Pseudosinella decipiens Denis, 1924

Pseudosinella decipiens Denis, 1924 198. Type locality: France.

World distribution. Palearctic.

Distribution in Europe. AT, BA, BE, CH, CT, DE, ES, FI, FR, GB, HU, HR, IE, ME, RO, RS, SE.

Distribution in Hungary. Bakony Mts. (Loksa & Rubio, 1966), Buda Mts. (Loksa 1966), Gerecse Mts. (Loksa 1966), Kiskunság (Traser 2002b), Moson Plain (Traser 2002a).

Ecological note. In caves, and forest soils (Gruia 1970; Fjellberg 2007; Shayanmehr et al. 2024).

Pseudosinella dungeri Winkler & Mateos, 2018

Pseudosinella dungeri Winkler & Mateos, 2018:357. Type locality: Hungary.

World distribution. Europe.

Distribution in Europe. HU.

Distribution in Hungary. Börzsöny Mts. (Winkler & Mateos 2018).

Ecological note. Xerothermophilic, in open habitats (Winkler & Mateos 2018).

Pseudosinella duodecimpunctata Denis, 1931

Pseudosinella duodecimpunctata Denis, 1931:82. Type locality: Italy.

World distribution. Europe, South Asia.

Distribution in Europe. AT, BG, CH, DE, ES, FR, IT, HU, RS, UA.

Distribution in Hungary. Dráva Plain (Loksa 1978a).

Ecological note. In caves, forests, and meadows (Denis 1931; Koledin et al. 1988; Kopeszki & Trockner 1994).

Pseudosinella edax Gisin, 1967 (Fig. 3A)

Pseudosinella edax Gisin, 1967:15. Type locality: Austria.

Material examined. 4 specimens, Zala Hills. Kiskapornak, 214 m a.s.l. 46°50'35"N, 16°58'26"E, from beech forest soil and moss, hand collecting, 01.ix.2018, leg. D. Winkler & Á. Csiszár.

Distribution in Europe. AT, HU (new record, Fig. 2), FR, IT, RO.

World distribution. Europe.

Distribution in Hungary. Zala Hills.

Ecological note. Silvicolous (Gisin 1967).

Pseudosinella cf. fallax (Börner, 1903)

Lepidocyrtus (Pseudosinella) fallax Börner, 1903:153. Type locality: Italy.

World distribution. Europe.

Distribution in Europe. AL, BG, CH, CT, ES, FR, GB, GR, HU, IE, IT, PT, RO, SL, SK, UA.

Distribution in Hungary. Aggtelek Karst (Traser 1999).

Ecological note. In open habitats, cultivated soils, and forests (Börner 1903; Kühnelt 1963; Fratello et al. 1989).

Pseudosinella hartnerae Winkler & Fiera, 2021

Pseudosinella hartnerae Winkler & Fiera [in Winkler et al.], 2021:123. Type locality: Hungary.

World distribution. Europe.

Distribution in Europe. HU, RO.

Distribution in Hungary. Nyírség (Winkler et al. 2021).

Ecological note. Phytodetriticolous, bryophilous, and hygrophilous (Winkler et al. 2021).

Figure 3. Pseudosinella species habitus in alcohol. A. P. edax Gisin, 1967; B. P. hercynica Rusek, 1979; C. P. noseki Rusek, 1985; D. P. thibaudi Stomp, 1977 (Photos by: D. Winkler).

Pseudosinella hercynica Rusek, 1979 (Fig. 3B)

Pseudosinella hercynica Rusek, 1979:258. Type locality: Czech Republic.

Material examined. 8 specimens, Bakony Mts. Bakonybél, 365 m a.s.l. 47°17'33"N, 17°43'12"E, from soil of beech forest, hand collecting, 16.vii.2016, leg. D. Winkler & Á. Csiszár).

World distribution. Europe.

Distribution in Europe. AT, CZ, HU.

Distribution in Hungary. Bakony Mts. (new record); Sopron Mts. (Winkler 2019).

Ecological note. Silvicolous (Rusek 1979; Winkler 2019).

Pseudosinella horaki Rusek, 1985

Pseudosinella horaki Rusek, 1985:142. Type locality: Czech Republic.

World distribution. Europe.

Distribution in Europe. CZ, HU, MD, PL, RO, SK, UA.

Distribution in Hungary. Aggtelek Karst (Traser 1999), Fertő-Hanság (Winkler & Tóth 2012), Gödöllő Hills (Harta et al. 2021), Kemeneshát (Winkler & Traser 2012b), Moson Plain (Traser 2002a), Sopron Mts. (Traser 2002a).

Ecological note. In forest soils (Rusek 1985; Traser 1999).

Pseudosinella immaculata (Lie-Pettersen, 1896)

Tullbergia immaculata Lie-Pettersen, 1896:16. Type locality: Norway.

World distribution. Palearctic.

Distribution in Europe. AT, CZ, DE, ES, FR, GB, HR, HU, IE, IT, LV, NL, NO, PL, PT, SE, SK, UA.

Distribution in Hungary. Aggtelek Karst (Loksa 1966; Traser 1999), around the Lake Balaton (Farkas, 1995), Bükk Mts. (Loksa 1956a), Fertő-Hanság (Traser 2002a), Mecsek Mts. (Loksa 1966), Mezőföld (Gergócs et al. 2022).

Ecological note. In forests, meadows, and caves (Loksa 1956a; Christian 1997; Fjellberg 2007).

Pseudosinella noseki Rusek, 1985 (Fig. 3C)

Pseudosinella noseki Rusek, 1985:140. Type locality: Czech Republic.

Material examined. 3 specimens, Middle-Tisza Region. Tiszavárkony, 92 m a.s.l. 47°3'18"N, 20°11'39"E, from soil of alluvial poplar forest, hand collecting, 14.vi.2018, leg. D. Winkler & G. Ónodi).

World distribution. Europe.

Distribution in Europe. CZ, HU (new record, Fig. 2), MD.

Distribution in Hungary. Middle-Tisza Region.

Ecological note. Silvicolous, hygromesophilous (Rusek 1985; Buşmachiu 2021).

Pseudosinella octopunctata Börner, 1901

Pseudosinella octopunctata Börner, 1901:705. Type locality: Germany.

World distribution. Cosmopolitan.

Distribution in Europe. AT, AZ, BA, BE, BG, CH, CZ, DE, ES, FI, FR, GB, HR, HU, IC, IE, IT, LV, MD, NO, PL, PT, RO, RS, SK, SL, SE, UA.

Distribution in Hungary. Bakony Mts. (Traser 1980), Balatonfelvidék (Paclt, 1960), Börzsöny Mts. (Dunger 1975), Buda Mts. (Loksa 1956b), Bükk Mts. (Loksa 1977), Dráva Plain (Loksa 1978a), Fertő-Hanság (Traser et al. 2006), Mezőföld (Stach 1929; Traser et al. 2011), Moson Plain (Winkler & Traser 2012a; Szigeti et al. 2022).

Ecological note. Open habitats, grasslands, patchy vegetation in gravel or sand slopes, and cultivated soils (Dunger 1975; Loksa 1978a; Fjellberg 2007; Winkler & Traser 2012a; Mehrafroz Mayvan et al. 2022).

Pseudosinella petterseni Börner, 1901

Pseudosinella Petterseni Börner, 1901:707. Type locality: Germany.

World distribution. Palearctic.

Distribution in Europe. AT, BE, DK, CH, CZ, DE, ES, FR, HU, IE, IT, PT, RS, SK.

Distribution in Hungary. Börzsöny Mts. (Dunger 1975), Bükk Mts. (Loksa 1977), Cserhát (Loksa 1978b), Gödöllő Hills (Harta et al. 2018, 2021), Hárosi-sziget (Loksa & Loksa 1996b), Mecsek Mts. (Loksa 1966), Moson Plain (Winkler & Traser 2012a), Veszprém-Devecser Graben (Winkler et al. 2018), Villány Mts. (Loksa 1966).

Ecological note. In forests, meadows, and cultivated soils (Gisin 1955; Fjellberg 2007; Winkler et al. 2018).

Pseudosinella sexoculata Schött, 1902

Pseudosinella sexoculata Schött, 1902:34. Type locality: Sweden.

World distribution. Cosmopolitan.

Distribution in Europe. AT, BA, BG, CH, CZ, DE, EE, ES, FI, FR, GB, HR, HU, IT, MD, NL, NO, ME, PL, PT, RO, RS, SK, SE, UA.

Distribution in Hungary. Fertő-Hanság (Traser 2002a), Kiskunság (Traser & Horváth-Szováti 2006; Traser 2002b; Flórián et al. 2019; Palkó et al. 2020), Gödöllő Hills (Harta et al. 2021), Mecsek Mts. (Loksa 1966), Mezőföld (Traser et al. 2011; Gergócs et al. 2022), Moson Plain (Winkler & Traser 2012a), Veszprém-Devecser Graben (Winkler et al. 2018), Villány Mts. (Loksa 1966).

Ecological note. Eurytopic species, occurring e.g. in forests, meadows, cultivated soils, organic deposits, and caves (Vedovini 1968; Bogojević 1971; Fjellberg 2007; Winkler & Traser 2012a).

Pseudosinella thibaudi Stomp, 1977 (Fig. 3D)

Pseudosinella thibaudi Stomp, 1977:231. Type locality: Germany.

Material examined. 17 specimens, Mátra Mts., beech forest, 47°53'8.70"N, 19°56'37.53"E; 662 m a.s.l., 27.x.2016, leg. D. Winkler & Á. Erdő, hand collecting.

World distribution. Europe.

Distribution in Europe. CZ, DE, HU (new record, Fig. 2), SK.

Distribution in Hungary. Mátra Mts.

Ecological note. Cavernicolous, eutroglophilous (Stomp 1977; Kováč 2000; Mock et al. 2015).

Pseudosinella wahlgreni (Börner, 1907)

Lepidocyrtus Wahlgreni Börner, 1907:176. Type locality: Egypt.

World distribution. Europe, North Africa.

Distribution in Europe. AT, BA, BE, BG, CH, CZ, DE, ES, FR, HU, IT, LT, MD, PL, RO, SK, UA.

Distribution in Hungary. around the Lake Balaton (Farkas, 1995), Bakony Mts. (Traser 1980, 2010; Loksa & Rubio 1966), Börzsöny Mts. (Loksa 1966; Dunger 1975), Buda Mts. (Loksa 1966), Bükk Mts. (Loksa 1966, 1977), Cserhát (Loksa 1978b), Dráva Plain (Loksa 1978a), Gerecse Mts. (Loksa 1966), Gödöllő Hills (Harta et al. 2021), Keszthely Mts. (Loksa 1966), Kiskunság (Palkó et al. 2020), Mátra Mts. (Loksa 1980), Mecsek Mts. (Loksa 1966), Nyírség (Loksa 1991), Őrség (Traser 1995), Pilis Mts. (Loksa 1966), Somogy Hills (Farkas et al. 1998), Sopron Mts. (Traser 1981), Vas-Sopron Plain (Traser 1981), Veszprém-Devecser Graben (Winkler et al. 2018), Vértes Mts. (Loksa 1966), Villány Mts. (Loksa 1966), Visegrád Mts. (Loksa 1966).

Ecological note. Mostly in forest soils (Dunger 1975; Traser 1981, 1995; Palkó et al. 2020).

Pseudosinella zygophora (Schille, 1908)

Lepidocyrtus zygophorus Schille, 1908:13. Type locality: Poland.

World distribution. Europe.

Distribution in Europe. AT, BG, BY, CZ, HU, PL, SK, UA.

Distribution in Hungary. Aggtelek Karst (Dudich 1932; Traser 1999), Moson Plain (Winkler & Traser 2012a; Szigeti et al. 2022).

Ecological note. In caves and forests (Dudich 1932; Sterzyńska 1995).

Key to the Hungarian species of Pseudosinella

Due to inconsistencies and uncertainties regarding the chaetotaxy, P. wahlgreni was not included in the identification key

1 ........ Head without ocelli. ...................................................................................................................... 2

—................................................................................................................................................................................... Head with ocelli. ................................................................................................................................... 10

2 ........ Unguiculus with large outer tooth. ............................................................................................... 3

— Unguiculus with no large outer tooth. ................................................................................................................................ 4

3 ........ Unguis paired basal teeth unequal, with outer one large and wing-like.......................................
............................................................................................................ P. petterseni Börner, 1901

—....... Unguis paired basal teeth equal. .............................................................. P. argentea Loksa, 1960 nec Folsom

4 ........ Abd IV with 1+3 dorsomedial macrochaetae. ........................ P. aggtelekiensis (Stach, 1929)

— Abd IV with 1+2 dorsomedial macrochaetae. ..................................................................................................................................... 5

5 ........ Th II with 4 macrochaetae. ................................................................. P. decipiens Denis, 1924

—....... Th II with less than 4 macrochaetae. ................................................................................................................................ 6

6 ........ Th II with 1 macrochaeta. .............................................. P. dungeri Winkler & Mateos, 2018

— Th II with 3 macrochaetae. ..................................................................................................... 7

7 ........ Labial chaetae l₁ and l₂ smooth. ..................................... P. immaculata (Lie-Pettersen, 1896)

— Labial chaetae L₁ and L₂ ciliated. ................................................................................................................................... 8

8 ........ Abd IV with microchaeta s. .................................... P. csafordensis Winkler & Mateos, 2018

— Abd IV without microchaeta s. ............................................................................................... 9

9 ........ Unguiculus without outer tooth. ....................................................... P. aelleni da Gama, 1973

—....... Unguiculus with one minute outer tooth. ................................................................................... P. noseki Rusek, 1985

10 ...... Head with 1+1 Ocelli. .............................................................. P. binoculata Kseneman, 1935

—....... Head with more than 1+1 Ocelli. ........................................................................................................................... 11

11 ...... Labial chaetae m₂, e, and l₁ smooth. ......................................................................................... 12

— Labial chaetae M₂, E, and L₁ ciliated. ................................................................................................................................. 17

12 ...... Abd II without chaeta p. ............................................................................................................ 13

—....... Abd II with chaeta p. .............................................................................................................................. 14

13 ...... Head with 2+2 Ocelli. ......................................................................... P. thibaudi Stomp, 1977

—....... .................................................................................................................................................. Head with 3+3 Ocelli. ............................................................................................. P. edax Gisin, 1967

14....... Head with 3+3 Ocelli. ................................................................................................................ 15

—....... Head with more than 3+3 Ocelli. ........................................................................................................................... 16

15 ...... Two ocelli anterior, one posterior in a separate pigment patch, Abd II with chaeta Q₁ as a macrochaeta. ..................................................................................... P. sexoculata Schött, 1902

—....... All three ocelli tightly grouped at the anterior margin of the eye spot, Abd II chaeta q₁ as a microchaeta. ........................................................................................ P. fallax (Börner, 1903)

16 ...... Head with 5+5 Ocelli. .................................................... P. hartnerae Winkler & Fiera, 2021

—....... .................................................................................................................................................. Head with 6+6 Ocelli. ..................................................................... P. duodecimpunctata Denis, 1931

17....... Abd II with chaeta A as a macrochaeta. ................................................................................... 18

—....... Abd II with chaeta a as a microchaeta. ............................................................................................................................. 19

18 ...... Head with 2+2 Ocelli, Abd II with chaeta q₁ as a microchaeta. ....... P. alba (Packard, 1873)

—....... Head with 4+4 Ocelli, Abd II with chaeta Q₁ as a macrochaeta. ................................................................................ P. octopunctata Börner, 1901

19 ...... Head with 2+2 Ocelli. ................................................................................................................ 20

— .................................................................................................................................................. Head with 5+5 Ocelli. ............................................................................................................................... 21

20 ...... Abd IV with 1+2 dorsomedial macrochaetae. ................................. P. bohemica Rusek, 1979

— Abd IV with 0+2 dorsomedial macrochaetae. .......................................................................................... P. hercynica Rusek, 1979

21 ...... Labial chaeta r smooth. .......................................................................... P. horaki Rusek, 1985

—....... ................................................................................................................................................ Labial chaeta R ciliated. ............................................................................ P. zygophora (Schille, 1908)

DISCUSSION

To date, a total of 23 species of Pseudosinella have been recorded in Hungary. The very first Pseudosinella species mentioned from the present territory of Hungary is the historic troglobiont springtail P. aggtelekiensis, collected by renowned Hungarian zoologists Elemér Bokor and Endre Dudich from the Baradla cave (Aggtelek Karst) and described by the Polish Collembola specialist Jan W. Stach (Stach 1929). The species was also detected later in the Slovak Karst (Kováč 2000), which forms a geographically contiguous unit with the Aggtelek Karst. Most probably, P. aggtelekiensis can be considered endemic in this region. Its occurrence in Austria (Strouhal & Vornatscher 1975) was already contested by Christian (1987) and also later by Kováč & Rusek (2012). The Romanian data not related to cave habitats (Fiera 2013) also requires validation based on the detailed redescription of P. aggtelekiensis by Kováč & Rusek (2012). Since the publication of the Collembola checklist of Hungary (Dányi & Traser 2007), a total of nine additional Pseudosinella species have been recorded, including three species (P. csafordensis, P. dungeri, and P. hartnerae) new to science (Winkler & Mateos 2018; Winkler et al. 2021). Three species (P. aelleni, P. bohemica, and P. hercynica) new to the fauna of Hungary have previously been reported in various papers (Traser 2008, 2010; Winkler & Traser 2012a; Winkler et al. 2012; Winkler 2014, 2019). An interesting remark should be mentioned regarding the species P. hercynica. In the original description, Rusek (1985) described the species as white in color, while the specimens collected in Hungary are orange-red marbled, varying in intensity from very light to darker (Fig. 3B) tones. Nevertheless, the Hungarian specimens match the original description in all other characters (including the chaetotaxy), and the habitat (beech forest) was also the same as that of the holotype. Three additional species (P. edax, P. noseki, and P. thibaudi), new to the Hungarian fauna, are presented here for the first time. Pseudosinella edax was collected in an old beech forest of Zala Hills, not far from the Austrian location of the holotype and other material in Gisin’s original description (Gisin 1967), also found in beech forest associations. Pseudosinella noseki was identified from soil samples taken in an alluvial poplar forest along the Tisza River. Outside Hungary, this species has only been recorded in the Czech Republic (type locality), Moldova, and Ukraine (Rusek 1985; Buşmachiu 2010; Kaprus & Goblyk 2015). Pseudosinella thibaudi was originally described from a cave in the Franconian Jura in Germany (Stomp 1977), and has been found later in subterranean habitats also in Slovakia (e.g., Kováč 2000; Mock et al. 2015) and in coniferous forests in the Czech Republic (Cuchta & Shrubovych 2015). In Hungary, P. thibaudi was collected from rocky slopes in a beech forest. As has been confirmed for most species belonging to the P. vandeli-group, the distribution of P. thibaudi is also most likely broader (Stomp 1986). To date, three species of Pseudosinella, namely P. argentea, P. csafordensis, and P. dungeri, have been determined as endemic to Hungary. Pseudosinella argentea is a troglobiont species, and it is only known from its type locality (Lóczy-cave) near Lake Balaton. In addition to a detailed redescription, Dányi (2011) also urged the necessity of renaming the species, being the junior primary homonym of P. argentea Folsom, 1902, a cavernicolous species from North America (Christiansen & Bellinger 1998). The other two endemic species (P. csafordensis and P. dungeri) have recently been described; the former from an old remnant alluvial forest, while the latter from a xerophilous pannonic steppic grassland patch (Winkler & Mateos 2018).

AUTHOR′S CONTRIBUTION

The author confirms his contribution to the whole processing steps in the research, examination of the specimens, illustrations, and preparation of the manuscript, and He read and approved the final version of the manuscript.

FUNDING

This research received no specific grant from any funding agencies.

AVAILABILITY OF DATA AND MATERIAL

The specimens listed in this study are deposited in Institute of Wildlife Biology and Management, University of Sopron and are available from the curator upon request.

ETHICS APPROVAL AND CONSENT TO PARTICIPATE

This study only included arthropod material, and all required ethical guidelines for the treatment and use of animals were strictly adhered to in accordance with international, national, and institutional regulations. No human participants were involved in any studies conducted by the authors for this article.

CONSENT FOR PUBLICATION

Not applicable.

CONFLICT OF INTERESTS

The authors declare that there is no conflict of interest regarding the publication of this paper.

ACKNOWLEDGMENTS

The author is greatly indebted to the subject editor and three anonymous reviewers for their valuable comments and suggestions improving the manuscript.

Arbea, J.I. (2013) A new cave species of Pseudosinella (Collembola, Entomobryomorpha, Entomobryidae) from Sima del Campamento (Jaén, Southern Iberian Peninsula). Arquivos Entomolóxicos, 8, 303–308.

Arbea, J.I. & Jordana, R. (1990) New species of Pseudosinella and Lepidocyrtus from Navarra (Northern Iberian Peninsula). Spixiana, 13, 25–31.

Bellinger, P.F., Christiansen, K.A. & Janssens, F. (1996–2025) Checklist of the Collembola of the World. http://www.collembola.org [Accessed October 15, 2025]

Bogojević, J. (1971) Prilog poznavanju faune Collembola Deliblatske peščare. Matica Srpska Journal for Natural Sciences, 40, 153–156. [in Serbian with English summary]

Börner, F. (1903) Über neue Altweltliche Collembolen, nebst Bemerkungen zur Systematik der Isotominen und Entomobryinen. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1903, 129–182. [in German] https://doi.org/10.5962/bhl.part.29866

Buşmachiu, G. (2010) Checklist of springtails (Collembola) from the Republic of Moldova. Travaux Du Muséum National d’Histoire Naturelle “Grigore Antipa”, 53 (1), 149–160. https://doi.org/10.2478/v10191-010-0011-x

Buşmachiu, G. (2021) Fauna Colembolelor (Hexapoda: Collembola) din Republica Moldova. Căpățină Print, Chișinău. 200 pp. [in Romanian]

Christian, E. (1987) Catalogus Faunae Austriae, Teil XIIa.: U.-Kl.: Collembola (Springschwänze). Verlag der Österreichischen Akademie der Wissenschaften, Wien. 83 pp. [in German]

Christian, E. (1997) Die wirbellosen Tiere der Hermannshöhle bei Kirchberg am Wechsel, Niederösterreich. Die Höhle, 50, 205–224. [in German]

Christiansen, K.A. & Bellinger, P.F. (1998) The Collembola of North America north of the Rio Grande; A taxonomic analysis. 2nd Edition. Grinnell College, Grinnell, IA. 1518 pp.

Christiansen, K.A., Bellinger, P. & da Gama, M.M. (1990) Computer Assisted Identification of Specimens of Pseudosinella (Collembola: Entomobryidae). Revue d’Écologie et de Biologie du Sol, 27 (2), 231–246.

Cipola, N.G., Oliveira, J.V.L.C., Bellini, B.C., Ferreira, A.S., Lima, E.C.A., Brito, R.A., Stievano, L.C., Souza, P.G.C. & Zeppelini, D. (2020) Review of eyeless Pseudosinella Schäffer (Collembola, Entomobryidae, and Lepidocyrtinae) from Brazilian caves. Insects, 11 (3), 194. https://doi.org/10.3390/insects11030194

Cuchta, P. & Shrubovych, J. (2015) Soil Collembola communities in montane coniferous forests of the Bohemian Forest. Suva Gabreta, 21 (2), 149–156.

Dallai, R. & Malatesta, E. (1982) Ricerche sui Collemboli. XXVI. Collemboli cavernicoli italiani. Lavori della Società Italiana di Biogeografia, 7, 173–194. [in Italian with English summary]

Dányi, L. (2011) Cave dwelling springtails (Collembola) of Hungary: a review. Soil Organisms, 83 (3), 419–432.

Dányi, L. & Traser, Gy. (2007) An annotated checklist of the springtail fauna of Hungary (Hexapoda: Collembola). Opuscula Zoologica, 38, 3–82.

Deharveng, L. & Thibaud, J.M. (1989) Acquisitions recentes sur les Insectes Collemboles cavernicoles d’Europe. Mémoires de Biospéologie, 16, 145–151. [in French with English summary]

Denis, J.R. (1931) Sur la faune Italienne des Collemboles, IV. Memorie della Societa Entomologica Italiana, 10, 80–85. [in French]

Dudich, E. (1930) Az Aggteleki barlang állatvilágának élelemforrásai. Állattani Közlemények, 27, 62–85. [in Hungarian with German summary]

Dudich, E. (1932) Biologie der Aggteleker Tropfsteinhöhle „Baradla” in Ungarn. Verlag Speläologisches Institut, Wien, 13, 1–246. [in German]

Dunger, W. (1975) Collembolen aus dem Börzsöny–Gebirge. Folia Historico-Naturalia Musei Matraensis, 3, 11–33. [in German with English summary]

Farkas, J. (1995) Collembola fauna from the shore of Lake Balaton, Hungary. Opuscula Zoologica, 27–28, 45–47.

Farkas, J., Szathmáry, K. & Strenner, A. (1998) Ecofaunistical investigations of Collembola, Araneae and Coleoptera in mosaic-like habitats in the Cinege Valley, Hungary. Opuscula Zoologica, 31, 49–61.

Fiera, C. (2013) Species of the genus Pseudosinella (Collembola: Entomobryidae) in the Romanian fauna. Acta Oecologica Carpatica, 6, 53–66.

Fjellberg, A. (2007) The Collembola of Fennoscandia and Denmark, Part II: Entomobryomorpha and Symphypleona. Fauna Entomologica Scandinavica, 42, 1–264.

Flórián, N., Ladányi, M., Ittzés, A., Kröel-Dulay, G., Ónodi, G., Mucsi, M., Szili-Kovács, T., Gergócs, V., Dányi, L. & Dombos, M. (2019) Effects of single and repeated drought on soil microarthropods in a semi-arid ecosystem depend more on timing and duration than drought severity. PLoS ONE, 14 (7), e0219975. https://doi.org/10.1371/journal.pone.0219975

Fratello, B., Sabatini, M.A., Mola, L., Uscidda, C. & Gessa, C. (1989) Effects of agricultural practices on soil arthropoda: Organic and mineral fertilizers in alfalfa fields. Agriculture, Ecosystems & Environment, 27 (1–4), 227–239. https://doi.org/10.1016/0167-8809(89)90088-1

Gergócs, V., Flórián, N., Tóth, Z., Szili-Kovács, T., Mucsi, M. & Dombos, M. (2022) Crop species and year affect soil-dwelling Collembola and Acari more strongly than fertilisation regime in an arable field. Applied Soil Ecology, 173, 104390. https://doi.org/10.1016/j.apsoil.2022.104390

Gisin, H. (1955) Recherches sur la relation entre la faune endogée de Collemboles et les qualités agri-coliques de sols viticoles. Revue suisse de Zoologie, 62, 601–648. [in French]

Gisin, H. (1967) Espèces nouvelles et lignées évolutives de Pseudosinella endogés (Collembola). Memórias e Estudos do Museu Zoologico da Universidade de Coimbra, 301, 1–25. [in French]

Gisin, H. & Gama, M.M. da (1969) Espèces nouvelles de Pseudosinella cavernicoles (Insecta: Collembola). Revue Suisse de Zoologie, 76, 143–181. [in French] https://doi.org/10.5962/bhl.part.146030

Gruia, M. (1970) Donnés sur certains Collemboles lapidicoles et lithoclasicoles des Carpates. Travaux de l’Institut “Emile Racovitza”, 9, 191–195. [in French]

Harta, I., Winkler, D. & Füleki, G. (2018) Impact of former long-term fertilization on springtail communities in a reforested experimental area. Columella – Journal of Agricultural and Environmental Sciences, 5 (1), 13–25. https://doi.org/10.18380/szie.colum.2018.5.1.13

Harta, I., Simon, B., Vinogradov, S. & Winkler, D. (2021) Collembola communities and soil conditions in forest plantations established in an intensively managed agricultural area. Journal of Forestry Research, 32, 1819–1832. https://doi.org/10.1007/s11676-020-01238-z

Jordana, R. (2012) Capbryinae and Entomobryini. In: Dunger, W. & Burkhardt, U. (eds) Synopses on Palaearctic Collembola. Soil Organisms, 84 (1), 1–391.

Jordana, R. & Baquero, E. (2005) New species of Pseudosinella (Collembola, Entomobryidae) from Tesla Mountains, Burgos, Spain. Serie Zoologica, 29 (1), 1–7.

Jordana, R., Baquero, E. & Ariño, A.H. (2024) Continuously updated: Collembola DELTA database. University of Navarra. https://www.unav.es/unzyec/collembola/Pseudosinella/ [Accessed July 14, 2024]

Kampmann, T. (1987) Untersuchungen an Collembolenzönosen in Wäldern der Bundesrepublik Deutschland. Ph.D. Dissertation, University of Bonn. 179 pp.

Kaprus, I.J. & Goblyk, K. (2015) Ecological and sozological evaluation of soils of the Transcarpathian Lowland with using Collembola communities. Proceedings of the State Natural History Museum, 31, 45–58.

Kopeszki, H. & Trockner, V. (1994) Effects of skiing on the collembolan fauna of an alpine meadow in Grodental (South Tyrol). Zoologischer Anzeiger, 233, 221–239.

Koledin, D., Matic, R., Tancic, N. & Stamenkovic, S. (1988) Collembola community dynamics in beech forest habitats on Jastrebac mountain (Yugoslavia). Arhiv bioloskih Nauka, 40 (1–4), 53–68.

Kovács, I.E. (1953) Biológiai terepszemle az aggteleki Békebarlangban. In: Jakucs, L. (ed.) A Békebalang Felfedezése. Művelt Nép, Budapest, pp. 86–91.

Kovács, I.E. (1954) Első vizsgálatok az aggteleki Békebarlang faunáján. Állattani Közlemények, 44 (1–2), 95–98. [in Hungarian with German summary]

Kováč, Ľ. (2000) A review of the distribution of cave Collembola (Hexapoda) in the Western Carpathians. Mémoires de Biospéologie, 27, 71–76.

Kováč, L. & Rusek, J. (2012) Redescription of two troglobiotic species of the genus Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) from the Western Carpathians. Zootaxa, 3341, 32–45. https://doi.org/10.11646/zootaxa.3341.1.2

Kováč, Ĺ., Žurovcová, M., Raschmanová, N., Jureková, N., Papáč, V. & Parimuchová, A. (2023) Troglomorphic adaptations on the northern European frontier: the phylogeny of the cave Pseudosinella (Hexapoda, Collembola) in the Western Carpathians. Frontiers in Ecology and Evolution, 11, 1169911. https://doi.org/10.3389/fevo.2023.1169911

Kseneman, M. (1935) Apterygota z území rybníku Lednickych II. Zvláštní otisk z Časopisu Národního Musea, 1931, 1–27. [in Czech with French summary]

Kühnelt, W. (1963) Über den Einfluβ des Mycels von Clitocybe infundibuliformis auf Streufauna. In: Doeksen, J. & van der Drift, J. (eds) Soil Organisms. North Holland Pub. Co., Amsterdam, pp. 281–287.

Loksa, I. (1956a) Zoologische Untersuchungen von Kollembolen im Bükkgebirge. Acta Zoologica Academiae Scientiarum Hungaricae, 2 (4), 379–419. [in German]

Loksa, I. (1956b) Die zönologische Untersuchung von Kollembolen in einer ungarischen Querceto- Potentilletum Albae-Assoziation. Acta Zoologica Academiae Scientiarum Hungaricae, 2, 199–243. [in German]

Loksa, I. (1960) Faunistisch–systematische und ökologische Untersuchungen in der Lóczy-Höhle bei Balatonfüred (Biospeologica Hungarica, XI.). Annales Universitas Scientiarum Budapestinensis, Sectio Biologica, 3, 253–266. [in German]

Loksa, I. (1961) Ökologisch-faunistischetische Untersuchungen in der Freiheitshöhle bei Égerszög. (Biospeologica Hungarica, XIII.). Acta Zoologica Academiae Scientiarum Hungaricae, 7, 219–230. [in German]

Loksa, I. (1962) Über die Landarthropoden der István-, Forrás- und Szeleta-Höhle bei Lillafüred. Karszt– és Barlangkutatás, 3, 59–81. [in German]

Loksa, I. (1966) Die bodenzoozönologischen Verhältnisse der Flaumeichen–Buschwälder Südostmittel-europas. Akadémiai Kiadó, Budapest. 437 pp. [in German]

Loksa, I. (1977) Quantitative Analysen des Collembolenbesatzes im Jahre 1975. im Untersuchungsgebiet bei Síkfőkút. Annales Universitatis Scientiarum Budapestinensis de Rolando Eötvös, Nominate, Sectio Biologica, 18–19, 205–218. [in German]

Loksa, I. (1978a) Die Collembolen-Fauna der Urwacholder aus der Umgebung von Barcs. Dunántúli Dolgozatok, Természettudományi Sorozat, 1, 51–64. [in German]

Loksa, I. (1978b) Mikrohabitate und ihre Bedeutung für die Verteilung der Collenbolengemeinschaften in einem Hainbuchen-Eichenbestand. Opuscula Zoologica, 15 (1–2), 93–117. [in German with English summary]

Loksa, I. (1980) A siroki Nyírjes-tó ugróvillás rovarai (Collembola). Folia Historico–Naturalia Musei Matraensis, 6, 107–114. [in Hungarian]

Loksa, I. (1991) The Collembola fauna of the Bátorliget Nature Reserves (NE Hungary). In: Mahunka, S. (ed.) The Bátorliget Nature Reserves – after forty years. Studia Naturalia, 1 (1), 269–277.

Loksa, I. & Loksa, I. (1996a) Adatok a Tétényi-fennsík ízeltlábú faunájához. In: Simon, T. (ed.) Természeti kincsek Dél-Budán; A Tétényi-fennsík és a Háros-sziget növény- és állatvilága, természetvédelme. Cserépfalvi Kiadó és Zöld Jövő Környezetvédelmi Egyesület, Budapest, pp. 40–51. [in Hungarian]

Loksa, I. & Loksa, I. (1996b) Adatok a Háros-sziget ízeltlábú faunájához. In: Simon, T. (ed.) Természeti kincsek Dél-Budán; A Tétényi-fennsík és a Háros-sziget növény- és állatvilága, természetvédelme. Cserépfalvi Kiadó és Zöld Jövő Környezetvédelmi Egyesület, Budapest, pp. 70–79. [in Hungarian]

Loksa, I. & Rubio, I. (1966) Anagben zu den Kenntnissen über die Collembolenfauna des Bakony- Gebirges. Opuscula Zoologica, 6 (1), 139–156. [in German]

Mehrafroz Mayvan, M., Sadeghi-Namaghi, H., Shayanmehr, M. & Greenslade, P. (2022) Contribution to the knowledge of Entomobryomorpha (Hexapoda: Collembola) from Northeastern Iran with new records and a key to the species. Journal of Insect Biodiversity and Systematics, 8 (2), 151–174. https://doi.org/10.52547/jibs.8.2.151

Mock, A., Šašková, T., Raschmanová, N., Jászay, T., Ľuptáčik, P., Rendoš, M., Tajovský, K. & Jászayová, A. (2015) An introductory study of subterranean communities of invertebrates in forested talus habitats in southern Slovakia. Acta Societatis Zoologicae Bohemicae, 79, 237–250.

Paclt, J. (1960) Über eine kleine Apterygoten-Ausbeute aus Ungarn. Opuscula Zoologica, 3 (3–4), 155–158. [in German]

Palkó, Á., Ónodi, G., Rédei, T. & Winkler, D. (2020) Talajfaunisztikai- és ökológiai vizsgálatok alföldi reliktum homoki tölgyesekben és a helyükön létesített idegenhonos faállományokban. Erdészettudományi Közlemények, 10 (1–2), 123–137. [in Hungarian with English summary] http://doi.org/10.17164/EK.2020.010

Ponge, J.P. (2000) Acidophilic Collembola: living fossils? Contributions from the Biological Laboratory, Kyoto University, 29, 65–74.

Rusek, J. (1979) Three new Pseudosinella species from Czechoslovakia (Collembola, Entomobryidae). Acta Entomologica Bohemoslovaca, 76, 255–265.

Rusek, J. (1985) New Palearctic Lepidocyrtus and Pseudosinella species (Collembola: Entomobryidae). Vestnik Ceskoslovenske Spolecnosti Zoologicke, 49, 132–146.

Shayanmehr, M., Yoosefi Lafooraki, E., Tully, T., D’Haese, C. & Ghajar Sepanlou, M. (2024) New records of springtails (Hexapoda: Collembola) for Iran from the Bula Hyrcanian forest. Journal of Insect Biodiversity and Systematics, 10 (1), 31–42. https://doi.org/10.61186/jibs.10.1.31

Sławska, M., Bruckner, A. & Sławski, M. (2017) Edaphic Collembola assemblages of European temperate primeval forests gradually change along a forest-type gradient. European Journal of Soil Biology, 80, 92–101. https://doi.org/10.1016/j.ejsobi.2017.05.003

Stach, J. (1929) Verzeichnis der Apterygogenea Ungarns. Annales Historico-Naturales Musei Nationalis Hungarici, 26, 269–312. [in German]

Sterzyńska, M. (1995) Collembola of secondary succession of the pine forests of Puszcza Białowieska. Fragmenta Faunistica, 37, 354–369.

Stomp, N. (1977) Nouvelle contribution a l’etude des Pseudosinella endoges d’Europe (Collemboles, Entomobryidae). Revue d’Écologie et de Biologie du Sol, 14 (1), 231–239.

Stomp, N. (1986) Le groupe de Pseudosinella vandeli (Insectes, Collemboles), dans les Alpes Orientales. In: Dallai, R. (ed) Second International Seminar on Apterygota. University of Siena, Siena, pp. 85–91. [in French]

Szent-Ivány, J. (1940) Adatok a Kőszeg-vidéki ugróvillás rovarok (Collembola) ismeretéhez. Dunántúli Szemle, 7 (7–8), 425–439. [in Hungarian]

Strouhal, H. & Vornatscher, J. (1975) Katalog der rezenten Höhlentiere Österreichs. Annales Naturhistorisches Museum Wien, 79, 401–542. [in German]

Szigeti, N., Berki, I., Vityi, A. & Winkler, D. (2022) Soil mesofauna and herbaceous vegetation patterns in an agroforestry landscape. Agroforestry Systems, 96 (4), 773–786. https://doi.org/10.1007/s10457-022-00739-6

Tschubby (2025) Reliefkarte Ungarn. https://w.wiki/GHie [Accessed October 15, 2025]

Traser, Gy. (1980) Adatok a farkasgyepűi bükkösök avarszintjének Collembola (ugróvillás) faunájához. Erdészeti és Faipari Tudományos Közlemények, 2, 19–23. [in Hungarian with English summary]

Traser, Gy. (1981) Talajinszekticidek hatása a talajfaunára. Erdészeti és Faipari Tudományos Közlemények, 3 (2), 129–138. [in Hungarian with English summary]

Traser, Gy. (1995) The Collembola fauna of Őrség (Insecta: Collembola). In: Víg, K. (ed.) Natural History of Őrség Landscape conservation Area I. Savaria Múzeum, Szombathely, pp. 43–47.

Traser, Gy. (1997) Egy szigetközi nemesnyáras biodiverzitása: a talajfauna. In: Pájer, J. & Fabich, E. (eds) V. Erdészeti Szakmai Konferencia kiadványa a WoodTech keretében. Sopron, pp. 57–64. [in Hungarian]

Traser, Gy. (1999) Springtails of the Aggtelek National Park (Hexapoda: Collembola). In: Mahunka, S. (ed.) The fauna of the Aggtelek National Park. Akadémiai Kiadó, Budapest, pp. 49–59.

Traser, Gy. (2002a) The Collembola of the Fertő–Hanság National Park. In: Mahunka, S. (ed.) The fauna of the Fertő-Hanság National Park. MTM, Budapest, pp. 259–270.

Traser, Gy. (2002b) Borókás–nyáras erdők ugróvillás (Insecta: Collembola) faunája. In: Barna, T. (ed.) Alföldi Erdőkért Egyesület Kutatói nap 2001-2002., Alföldi Erdőkért Egyesület, Gyula, pp. 46–52.

Traser Gy. (2003) Hansági nemesnyár és éger erdők ugróvillás (Insecta: Collembola) faunája. III. Kárpát-medencei Biológiai Szimpózium kiadványa. Magyar Biológiai Társaság, Budapest, pp. 153–157. [in Hungarian]

Traser, Gy. (2006) A gyarmatpusztai vadgesztenyés ugróvillás (Insecta: Collembola) faunájának diverzitása. Folia Musei Historico-Naturalis Bakonyiensis, 23, 33–39. [in Hungarian with English summary]

Traser, Gy. (2008) Rovartani érdekességek (Insecta: Collembola) egy patak menti égererdőből. In: Barna, T. (ed.) Alföldi Erdőkért Egyesület Kutatói Nap 2008. Alföldi Erdőkért Egyesület, Szeged, pp. 120–125. [in Hungarian with English summary]

Traser, Gy. (2010) A Collembola fauna (Hexapoda: Entognatha) Porva körzetében a harmadik Biodiverzitás Nap gyűjtései alapján. A Bakonyi Természettudományi Múzeum Közleményei, 27, 49–54.

Traser, Gy. & Horváth-Szováti, E. (2004) Tájhasznosítás hatása a talajfaunára: ugróvillás (Parainsecta: Collembola) közösségek diverzitása Csévharaszton. In: Barna, T. (ed.) Alföldi Erdőkért Egyesület Kutatói Nap 2004. Alföldi Erdőkért Egyesület, Kecskemét, pp. 51–62. [in Hungarian with English summary]

Traser, Gy. & Horváth-Szováti, E. (2006) Land–use intensity and diversity parameters: Collembola (Insecta) communities in Csévharaszt (Hungary). In: Csóka, Gy., Hirka, A. & Koltay, A. (eds) Biotic Damage in Forests. Proceedings of the IUFRO (WP7.03.10) Symposium held in Mátrafüred, Hungary, pp. 329–339.

Traser, Gy., Szűcs, P. & Winkler, D. (2006) Collembola diversity of moss habitats in the Sopron Region, NW-Hungary. Acta Silvatica et Lignaria Hungarica, 2, 69–80.

Traser, Gy., Winkler, D. & Molnár, M. (2011) A mezőföldi Szent László-víz völgyének ugróvillás (Collembola) faunája. A Bakonyi Természettudományi Múzeum Közleményei, 28, 71–79. [in Hungarian with English summary]

Varga, Z. (1995) Geographical patterns of biological diversity in the Palearctic region and the Carpathian Basin. Acta Zoologica Academiae Scientiarum Hungaricae, 41 (2), 71–92.

Vedovini, A. (1968) Contribution a l'inventaire faunistique des cavites souterraines des environs de Marseille. Annales de Speleologie, 23, 229–242. [in French]

Winkler, D. (2014) Collembolan response to red mud pollution in Western Hungary. Applied Soil Ecology, 83, 219–229. https://doi.org/10.1016/j.apsoil.2013.07.006

Winkler, D. (2019) Ugróvillások (Collembola). In: Kárpáti, L. (ed.) Soproni Tájvédelmi Körzet - Monografikus tanulmányok a Soproni-hegység természeti és kulturális értékeiről. Szaktudás Kiadó Ház Zrt., Budapest, pp. 133–135. [in Hungarian]

Winkler, D. & Mateos, E. (2018) New species of Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) from Hungary. Zootaxa, 4382 (2), 347–366. https://doi.org/10.11646/zootaxa.4382.2.7

Winkler, D. & Tóth, V. (2012) Effects of afforestation with pines on Collembola diversity in the limestone hills of Szárhalom (West Hungary). Acta Silvatica et Lignaria Hungarica, 8, 9–20. https://doi.org/10.2478/v10303-012-0001-8

Winkler, D. & Traser, Gy. (2012a) Collembola Diversity in Agricultural Environments (Lajta Project, Western Hungary). In: Proceedings of the International Scientific Conference on Sustainable Development and Ecological Footprint. 26–27 March 2012, Sopron, Hungary, pp. 1–5.

Winkler, D. & Traser, Gy. (2012b) Eco-faunistic study on the Collembola fauna in the Vasvár-Nagymákfa area. Natura Somogyiensis, 22, 39–52. https://doi.org/10.24394/NatSom.2012.22.39

Winkler, D., Korda, M. & Traser, Gy. (2012) Rare and little known Collembola species from Hungary. Opuscula Zoologica, 43 (1), 89–95.

Winkler, D., Bidló, A., Bolodár-Varga, B., Erdő, Á. & Horváth, A. (2018) Long-term ecological effects of the red mud disaster in Hungary: Regeneration of red mud flooded areas in a contaminated industrial region. Science of the Total Environment, 644, 1292–1303. https://doi.org/10.1016/j.scitotenv.2018.07.059

Winkler, D., Németh, M.T. & Fiera, C. (2021) A new species of Pseudosinella Schäffer, 1897 (Collembola, Entomobryidae) from Hungary and Romania, with redescription of the related species Pseudosinella huetheri Stomp, 1971. ZooKeys, 1063, 121–137. https://doi.org/10.3897/zookeys.1063.73094