Trachelipus Budde-Lund, 1908

Beron, Petar, 2020, Checklist of Isopoda Oniscidea (Crustacea) of Bulgaria, Ecologica Montenegrina 38, pp. 227-244 : 236-240

Genus Trachelipus Budde-Lund, 1908

Trachelipus myrmicidarum (Verhoeff, 1936) – Zemen, Skakavitsa - Verhoeff (1936: 10); Sandanski – Vandel (1967: 359), Messner (1967: 23); Belassitsa Hut, Kresna, Sandanski – Andreev (2002: 70). Myrmecophilous, with Messor structor rufitarsis , M. barbarus varrialei (Hym., Formicidae ). Bulgarian endemic.

Trachelipus nodulosus (C.L. Koch, 1838) (= balticus Verhoeff, 1907) - Medenik (Vr 18), Dragalevtsi (Vitosha) - Verhoeff (1926: 145); Borovets, 1460 m, Kolibata (Sf 12), Han Maara (Pv 1), General Toshevo - Vandel (1967: 338).

Trachelipus rathkei Brandt, 1833 – Sofia, Vitosha, Kuklen, Bachkovski manastir, Shipka, Kazanlak, Lakatnik, Shtarkelovo gnezdo, Sandanski – Messner (1967: 23); Katsa peshtera (Rz 4) – Vandel (1967: 338); Vitosha, Bachkovska manastir, Malo Konare (Distr. Plovdiv), Tsrancha, Ognyanovo (Distr. Pazardjik), Uzana, Shipchenski prohod (Stara planina), Peshketo (Vr 68) – Andreev (1972: 184-185); Peshterata s dvata vhoda (Bs 13), Kaleto (Bs 16), Tsakalat (Bs 21), Mecha dupka (Bs 24) – Beron (1994: 17 - S. Andreev det.); Deli Burun (Shn 11), Salievata peshtera (Bl 2), Mussinskata peshtera (Tn 12) – Andreev (2002: 69).

Trachelipus squamuliger (Verhoeff, 1907) (= Trachelipus absoloni (Strouhal, 1939)) - Haydushka dupka (Lv 2) – Verhoeff (1926: 145); Sozopol – Frankenberger (1941: 8 – sub “ Tracheoniscus bulgaricus Verh. ”); Strouhal (1939: 201); Razhishka yama (Sf 55) - Guéorguiev & Beron, (1962: 299 – A. Vandel det., sub “ Trachelipus absoloni ”); Haydushkata peshtera (Pn 2) - Vandel (1967: 267 – sub “ Trachelipus absoloni ”); Skoka (Bl 32) – Beron et al. (2011: 594 – S. Andreev det.). Troglophile. According to Schmalfuss (2003), the species T. bulgaricus Verhoeff, 1926 with its two subspecies is synonymous also of T. squamuliger . Below the localities of these possible subspecies are provided.

[ Trachelipus bulgaricus bulgaricus Verhoeff, 1926 ] ( Tracheoniscus bulgaricus ) - Haydushka dupka (Lv 2) – Verhoeff (1926: 145); Saeva dupka (Lv 18) – Verhoeff (1936: 11); Draganchovitsa (Lv 27) - Strouhal (1939: 199); “Cave without name near Karlukovo” – Vandel (1965: 266); Yarkovets (Sf 77), Bankovica (Lv 5), Temnata dupka (Lv 16), Cerovo – Vandel (1967: 339); Haydushka dupka (Pn 2), Devetashkata peshtera (Lv 37) - Beron (1994: 17, Andreev, 2002: 69). Troglophile (or trogloxene?).

[ Trachelipus bulgaricus bureschi Verhoeff,1926 ] - Nirits (Sl 7) – Verhoeff (1926: 145); Dolnata Maaza (Sl 1), “Cave near Krushuna” – Verhoeff (1936: 11 as Tracheoniscus bulgaricus bureschi ); Saeva dupka (Lv 18) – Strouhal (1939: 199); Malkata Humba (Sl 13) - Vandel (1965: 266). Troglophile.

Oniscidea incertae sedis

Genus Myrmekiocellio Verhoeff, 1936 (endemic genus for Bulgaria, but according to Schmalfuss, 2003, with doubtful status)

[ Myrmekiocellio squamatus Verhoeff, 1936 – Skakavitsa - Verhoeff (1936: 18, with the ants Tetramorium caespitum ). Bulgarian endemic]. According to Sfenthourakis (in litt.), the description is based on a juvenile of Porcellium recurvatum and is not valid.

Isopoda Oniscidea in Bulgarian caves

A substantial part of the list of Balkan Oniscidea consists of cave dwellers, with many troglobitic species and endemic genera. If we consider only the inhabitants of Bulgaria and the neighbouring Serbia, Rep. N. Macedonia, Greece, Eastern Thrace and Northern Dobrudja, we find the following:

Bulgaria ( Beron, 2015, with corrections) – Oniscidea are known from 185 caves. They belong to 9 families ( Ligiidae , Styloniscidae , mostly Trichoniscidae , Trachelipodidae , Chaetophilosciidae, Platyarthridae , Porcellionidae , Cylisticidae , Armadillidiidae ), 24 genera and 49 species. The 26 troglobitic species belong to Trichoniscidae (24 – Hyloniscus , Trichoniscus , Alpioniscus , Balkanoniscus , Rhodopioniscus , Bureschia , Bulgaronethes , Cyphoniscellus , Vandeloniscellus , Beroniscus ) and Styloniscidae (2 Cordioniscus ). See also Verhoeff (1926, 1929, 1936), Vandel (1965, 1967), Tabacaru (1993), and Andreev (1970, 1972, 1983, 1985, 1986, 1987, 2000, 2002 and others).

Greece ( Alexiou & Sfenthourakis, 2013; Beron, 2016) – the cave Oniscidea of Greece (Dodekanese excluded) belong to 11 families and 25 genera ( Beron 2016). Buddelundiella sporadica belongs to Trichoniscidae and not to a separate family (Schmalfuss, 2003). The 34 troglobitic species belong to Styloniscidae (9 Cordioniscus ), Trichoniscidae (18 - Alistratia , Alpioniscus , Buddelundiella . Graeconiscus , Libanonethes , Trichoniscus ), Scleropactidae ( Kithironiscus ), Armadillidiidae ( Platanosphaera (= Troglarmadillidium ), Schizidium ). Unlike Bulgaria, members of Cordioniscus , Alpioniscus and Schizidium prevail here. See also Andreev (1984, 1985, 1986, 1997, 2001, 2004, 2013), Schmalfuss (1979, 1995 and many others), Sfenthourakis (1996), Vandel (1955, 1958, 1959, 1964, 1968).

Serbia ( Buturović, 1960; Pljakić, 1977; Ćurčić, 2014) - In Serbia (Kosovo included) Oniscidea have been analysed by Pljakić (1970, 1972, 1977) with descriptions of the new taxa Hyloniscus stankovici , H. kossovensis , Microtitanethes licodrensis (n.g., n.sp.), and 10 new Trichoniscus . Buturović (1960) added Hyloniscus kopaonicensis . The list of the cave Oniscidea in Ćurčić, Juberthie & Decu in Ćurčić (2014) contains 13 sp. (all Trichoniscidae , 12 described by Pljakić in 1970 and 1977). One genus ( Microtitanethes ) and all Trichoniscus are endemic to Serbia. Macedoniscus is a synonyme of Alpioniscus (see Schmalfuss, 2003). The cave Oniscidea of Serbia clearly remain understudied.

Rep. N. Macedonia ( Komnenov, 2010; Sket et al., in Ćurčić, 2014) – The list of cave Isopoda terrestria of Komnenov (2010) contains 11 identified to species, including 10 troglobites and Trichoniscus semigranulatus (troglophile, known also from Bulgaria). Alpioniscus vejdovskii is known also from Greece. The other nine (including five Alpioniscus ) are endemic troglobites and include the endemic genera Macedonethes and Mladenoniscus . Vardaroniscus is a synonyme of Cyphoniscellus (see Schmalfuss, 2003). The list of Sket et al. in Ćurčić (2014) contains only seven species (without Mladenoniscus belavodae , Alpioniscus macedonicus , Macedonethes skopjensis , and M. stankoi ) (see also Buturović, 1954, 1955 a, 1955b, I. Karaman, 2003, Karaman & Horvatović, 2008; M. Karaman, 1961, 1966).

Northern Dobrudja ( Romania) ( Giurginca & Curčić, 2003; Tabacaru & Giurginca, 2013) – Among the 41 species of Isopoda terrestrial recorded from Northern (Romanian) Dobrudja there are 17 species recorded from caves and drills, including such interesting taxa like Kithironiscus and Caucasonethes . These and some other amazing troglobites like Trachelipus troglobius , Armadillidium tabacarui and others were described among the unique inhabitants of the “lost world” of the cave of Movile (see also Tabacaru, 1993; Tabacaru & Boghean, 1989).

Eastern Thrace (European Turkey) ( Verhoeff, 1941) - from the cave Barut hane near Yarim Burgaz have been recorded Haplophthalmus stygivagus , Trichoniscus provisorius , and Kosswigius bilselii . Among the Oniscidea of European Turkey the genus Kosswigius Verhoeff, 1941 is notable as it is found only in this area and on Lesvos Island (N. Aegean Sea, Greece).

In the monograph of Guéorguiev (1977) and in some of his earlier papers the origin of the troglobitic terrestrial isopods of the Balkans has been addressed. Many genera have been considered as having a Paleoaegeidian and Nordaegeidean origin, the genera Balkanoniscus , Beroniscus and Bureschia were considered as of Mesoaegeidian and Cordioniscus was ranged among the most ancient Gondwanian relicts in the cave fauna of the Balkans.

Meanwhile many new data have been published. The analysis of Guéorguiev has been updated and completed in the monographs on Bulgaria by Beron (1915) and on Greece by Beron (2016). A remarqable finding was the many new species of Cordioniscus in the caves of Bulgaria and Greece.

The fauna of Oniscidea of the neibouring territories

In Northern (Romanian) Dobrudja 41 sp. of Oniscidea have been recorded ( Giurginca and Ćurčić 2003) in 10 families. The conclusion of these authors is that “..we may emphasize two main features of the Oniscidea from Dobrudja: a high degree of endemicity resulting from the existence of an Euxinic glacial subrefugium and also the preponderance of the Mediterranean, Balkan-Central European, East-European and Holarctic species”. A remarkable finding in Dobrudja was a member of the genus Kithironiscus , which “..marks an isolated northern point in the typically Gondwanian spreading of the Scleropactidae family”. The Oniscidea in Bulgarian Dobrudja are much less well known and some of the interesting genera like Kithironiscus or Caucasonethes could be expected also from Bulgaria.

The Greek fauna of Oniscidea has been analysed by Schmalfuss (1979), Sfenthourakis (2013) and Alexiou and Sfenthourakis (2013), after many papers of Strouhal, Vandel, Schmalfuss, Andreev and others. I myself spent many years collecting Isopoda in Greek caves.

The checklist of Alexiou and Sfenthourakis (2013) contains data on 238 species of 47 genera and 19 families. Again, many species and some peculiar findings ( Kithironiscus ) are from caves. The presence of the mainly South American family Scleropactidae is remarkable. Seven genera are Greek (and Balkanic) endemics: Acteoniscus Vandel , Alistratia Andreev, Paxodillium Schmalfuss , Trichodillidium Schmalfuss , Xeroporcellio Strouhal , Thrakosphaera Schmalfuss , and Rodoniscus Arcangeli , as well as 161 species. Best represented are the families Armadillidiidae (88 sp.), Trichoniscidae (38), Porcellionidae (26), and Trachelipodidae (20). Remarkable are the 13 species of Cordioniscus Graeve ( Styloniscidae ), a genus distributed in the countries of Gondwanian origin, and in Mediterranean ( Vandel, 1968, Guéorguiev, 1977). The cause of this distribution is unclear ( Schmalfuss & Erhard, 1998). All of them are troglobitic and endemic to Greece. Many of them are living in Southern Greece, Crete or the Dodecanese, whose fauna is very different from that of Bulgaria.

Regarding the Oniscidea of northern Greece, immediately bordering Bulgaria (the territory East of Struma, or Strymon River and the islands Thassos and Samotraki), according to the checklist of Alexiou and Sfenthourakis (2013) from this area are known at least 40 species. Some (mainly cave species) are endemic: Cordioniscus graevei (cave of Alistrati), C. vandeli (caves near Zygos), Alistratia beroni (cave of Alistrati, endemic genus), Alpioniscus thracicus (cave of Maronia), Porcellium collicola (Greek Makedonia, Thrace), Thracosphaera schawalleri (Thrace, endemic genus), Armadillidium jerrentrupi (Nestos) , A. pangaionum (Pangaion) , A. peraccae (Thassos island), A. petralonense (from Thessaloniki to Petralona), A. phalacronum (Phalacron, Rhodopes) , A. tuberculatum (Kavala) . Particularly interesting is the endemic genus Thrakosphaera ( Tendosphaeridae ) and the species of Alpioniscus , which could be expected also from Bulgaria. Other species are widespread, littoral or halophilic, or riparian ( Ligia , Ligidium , Tylos , Stenoniscus ).

Some species are also present in Bulgaria ( Monocyphoniscus bulgaricus , Trachelipus squamuliger , Trichoniscus rhodopiensis and some widespread or myrmecophilous species).

On the map in the paper of Sfenthourakis & Hornung (2018) we see that 132 species are known from continental Greece, 24 from Rep. N. Macedonia, 67 from Bulgaria, 49 from European Turkey, 25 from Albania, 45 from Montenegro, 44 from Serbia ( Kosovo included), 65 from Bosnia and Herzegovina, 140 from continental Croatia, and 67 from Slovenia.

The fauna of Rep. North Macedonia contains endemic genera like Macedonethes Buturović. It has been studied mostly by Buturović (1954, 1955 a, 1955b), who described Vardaroniscus (= Cyphoniscellus ) tetraceratus, Macedoniscus (= Alpioniscus ) vardarensis, Alpioniscus slatinensis , A. karamani , and others. Karaman (1961) recorded other Oniscidea from North Macedonia. He recorded ( Karaman, 1966) also “ Tendosphaera verrucosa ” near Skopje, but Schmalfuss rations that this is certainly a species of Tendosphaera , but can neither be the verrucosa nor graeca ”[these are the only known species of Tendosphaera ]. The family is not yet known from Bulgaria, but most probably occurs there. I. Karaman & M. Horvatović (2008) described the new (endemic) genus and species Mladenoniscus belavodae from the cave Bela Voda near Demir Kapija.

Oniscidea of Serbia have been explored by Buturović (1960), who described in 1970 – 1977 several more species (including some near the border with Bulgaria) – Hyloniscus stankovici , H. kossovensis , Macedoniscus (now Alpioniscus ) metohicus, Microtitanethes licodrensis , Trichoniscus bogovinae , T. serbicus , T. buturovici , T. naissensis , T. serboorientalis , T. bononiensis timocensis , T. bononiensis sotirovi , T. pancici , T. licodrensis . Pljakić (1970) also reported some Oniscidea ( Ligiidae , Mesoniscidae , Trichoniscidae ).

Regarding Eastern (Turkish) Thrace Verhoeff (1941) published several species from the cave Barut hane near Yarim Burgaz and some others from other Thracian places .

Isopoda Oniscidea expected from Bulgaria

On the Balkan Peninsula (the Alpine part of Slovenia and the Dodecanese excepted) 20 families of Isopoda Oniscidea are known, in ca.80 genera. The Slovenian genera like Moserius and Calconiscellus are rather part of the Italian or Apine fauna. Almost half of the genera belong to the family Trichoniscidae , 32 genera are endemic for the Peninsula (including the islands). Most of the endemics are troglobites.

The following three families, known from the Balkans, but not yet found in Bulgaria.

Mesoniscidae – Mesoniscus graniger (Friv.) is known from Serbia and Romania.

Scleropactidae – the (relict?) genus Kithironiscus Schmallfuss, 1995 of this mostly Neotropical family is known from Kithira Island (S. Greece) and from Northern Dobrudja.

Tendosphaeridae – the genus Thrakosphaera Schmallfuss, 1998 is endemic for Thrace (the Greek province Thraki, not far from the Bulgarian border).

Several genera of other families ( Leptotrichus Budde-Lund and others) could also be found in Bulgaria, as well as new species, mostly from caves. Several more species of Armadillidium Brandt and Alpioniscus Racovitza are also to be expected from Bulgaria.