Hylebainosoma tatranum Verhoeff, 1899

Tajovský, Karel, Mock, Andrej & Papáč, Vladimír, 2014, The genus Hylebainosoma Verhoeff, 1899 (Diplopoda, Chordeumatida, Haaseidae): Redescription of Hylebainosoma tatranum, description of a new troglobiont species and notes to the Hylebainosoma – Romanosoma species group, Zootaxa 3764 (5), pp. 501-523 : 503-516

Hylebainosoma tatranum Verhoeff, 1899

Figs 1–20

Hylebainosoma tatranum Verhoeff, 1899: 136 –137, Figs 3–6

Hylebainosoma tatranum var. Dudichi Verhoeff, 1941: 232 –234, Figs 1–4 Hylebainosoma tatranum jósvaense Loksa, 1962: 158 -159, Figs 9–11

Material examined. Slovakia. Cerová vrchovina Mts. (pseudokarst in volcanic bedrock): Nyáryho Cave (altitude of entrance/length/depth of cave: 570 m a.s.l./ 25 m /0 m), p.t., 8–10 m from the entrance, 9.v.–24.x.2000 (RM), 4 ♂, 2 ♀; Šurický úkryt Cave (525 m a.s.l./ 69 m /- 13 m), 30 m from entrance, p.t., 8.iii.–24.x.2000 (RM), 2 ♂; Čierna hora Mts. (all material sampled by AM): Čierna hora Mt. (1,000–1,020 m a.s.l.), quartzite stony debris on the slope of top, p.t., 13.xi.1997 – 5.v.1998, 1 ♂, 1 ♀; 5.v.–22. vii.1998, 1 ♂, 2 ♀; 13.xi.1998 – 15.x.2002, 8 ♂, 8 ♀; Koľvek (790 m a.s.l.), quartzite stony debris at the peak, p.t., 25.viii.–15. x.1997, 1 ♀; 13.xi.1997 – 5.v.1998, 1 juv.; 13.xi.1998 – 29.iv.1999, 4 ♂, 4 ♀; 16.xi.1999 – 4.iv.2000, 1 ♂, 1 ♀; Malý Ružínok Valley (340 m a.s.l.), alder forest at the stream, p.t., 29.vii.–28. viii.1999, 1 ♀; Vozárska National Natural Reserve (500 m a.s.l.), fir-beech forest, p.t., 5.v.–8. vi.2000, 1 juv.; 14.xii.2000 – 19.i.2001, 2 ♂; 18.iv.–21. v.2001, 1 ♂; Vyšná dolina Valley in the Roháčka Massif (650 m a.s.l.), i.s., 5.v.1998, 1 ♂, 1 ♀; Low Tatra Mts.: Jánska Valley, under the Ludárka Hill (about 1,300 m a.s.l.), mixed forest and meadow, granite bedrock, 29.vi.2006 (AB), 5 ♂ 6 ♀; Krížska Valley (ca 1,200 m a.s.l.), spruce forest and Vaccinium shrub, i.s. (JG), 28.iv.1964, 2 ♀; Salatín Peak (1,634 m a.s.l.), i.s. (JG), 27.ix.1956, 1 ♂, 2 ♀; Slanské vrchy Mts.: Šimonka Peak (1,092 m a.s.l.), i.s. (JG), 30.x.1961, 5 ♂, 11 ♀, 1 juv.; Slovak Karst: Ardovská Cave (314 m a.s.l./ 1,492 m), around the entrance (outside), 8.iii.2001 (AM), 4 ♂, 11 juv.; Brzotín surroundings (600 m a.s.l.), i.s. (JG), v.1946, 1 ♂; Diviačia Chasm (590 m a.s.l./ 468 m /- 123 m), bottom of the entrance shaft, p.t., 28.vi.–29.x.2004 (ĽK), 2 ♂, 3 ♀; deeper part of the bottom of entrance shaft, p.t., 28.vi.– 29.x.2004 (AM), 1 ♂, 1 ♀; in the depth 24–38 m, on the wood, 28.vi.2004 (AM), 2 ♂, 2 ♀, 22 juv.; Silická Ice Cave (495 m a.s.l./ 1,100 m /- 110 m), entrance slope, i.s. (JG), 4.vii.1947, 1 juv. ♀; 11.v.1963, 8 ♂, 3 ♀, 23 juv.; 22.vi.1963, 1 ♂, 4 juv., 15.x.1963, 3 ♂, 7 ♀, 6 juv.; 21.v.1963, 1 ♂, 2 ♀, 22 juv.; 18.v.2006 (AM), 3 ♀, 1 juv.; Zvonivá jama Chasm (494 m a.s.l./ 494 m /- 101 m), bottom, on the wood 5.v.2006 (AM), 1 ♂; bottom, wood, 16.xi.2006 (AM), 2 ♂, 2 ♀, 2 juv.; Slovenský raj (Slovak Paradise): Dobšinská Ice Cave (971 m a.s.l.), i.s. (JG), 5.vii.1947, 1 juv. ♀; Grajnár Peak (1,023 m a.s.l.), spruce forest, i.s. (JG), 22.iv.1963, 3 ♂, 6 ♀, 5 juv., 26.v.1963, 1 ♂, 1 ♀, 4 juv., 18.x.1963, 1 ♂, 2 ♀; Tatra Mts.: Danielov dom (1,260 m a.s.l.), habitat opened by cutting of spruce forest after windthrow, p.t. (AM), 19.vi.–20. x.2005, 3 juv.; Hincovo pleso Lake (1,945 m a.s.l.), i.s. (JG), viii.1946, 1 ♀; Hlúpy Peak (2,062 m a.s.l.), i.s. (JG), 28.–31. viii.1955, 12 ♂, 27 ♀, 11 juv.; 27.vii.1957, 1 ♂, 4 ♀; 9.ix.1963, 1 ♂; Jamy (1,075 m a.s.l.), spruce forest destroyed by windthrow, p.t. (AM), 5.x.2011 – 9.v.2012, 1 ♀, 4 juv.; Javorová Valley (1,100–1,300 m a.s.l.), spruce forest, under bark of trees, i.s. (JG), 27.vii.1965, 2 ♂, 3 ♀, 6 juv., 29.vii.1965, 1 ♂, 1 ♀; Plesnivec (1,290 m a.s.l.), spruce forest, i.s. (JG), 29.vii.1965, 1 ♂, 1 juv. ♂, 18 ♀, 3 juv.; Pod Ostrvou (1,520 m a.s.l.), climax spruce forest, p.t. (AM), 5.v.–6. x.2010, 2 ♂; 8.x.2009 – 5.v.2010, 10 ♂, 20 ♀, 9 juv.; Protež (Plesnivec) Chalet (1,450–1,700 m a.s.l.), alpine meadow, i.s. (JG), 7.vii.1953, 1 ♂; Rozpadlé pleso Lake (unclear location), Swiss pine growth, i.s. (JG), 28.ix.1986, 2 ♂, 1 ♀; Smrekovec (1,250 m a.s.l.), spruce forest, p.t. (AM), 1.x.2008 – 28.iv.2009, 1 ♂, 2 ♀; 6.x.2011 – 9.v.2012, 1 juv; Tomanová Valley, under the Poľská Tomanová Peak (1,660 m a.s.l.), subalpine Vaccinium shrubs, granite bedrock, p.t. (KT), 3.vii.–30. ix.1997, 3 ♂, 5 ♀, 30.ix.1997 – 30.vi.1998, 1 ♀, 1 juv, 3.ix.2007 – 30.vi.2008, 1 ♂, 3 ♀; Hvížďalka Valley (about 1,750 m a.s.l.), alpine grassland, limestone bedrock, p.t. (KT), 1.vii.–1. x.1997, 1 ♀; Vihorlat Mts.: Morské oko Lake (618 m a.s.l.), surrounding beech forest, i.s. (JG), viii.1948, 1 ♀; Vtáčnik Mts.: mountain ridge (up to 1,346 m a.s.l.), maple and beech forest growth, i.s. (JG), 27.iii.1957, 2 ♂; Zvolenská kotlina Basin: Ponická Huta (620 m a.s.l.), oak forest, i.s. (JG), 16.x.1955, 6 ♂, 1 ♀, 2 juv.

Czech Republic. Moravskoslezské Beskydy Mts.: Mazácký Grúnik Nature Reserve nr. Ostravice Village (665 m a.s.l.), fragments of fir-beech Carpathian forest, i.s., 6.xi.1991 (KT), 1 ♂, 1 juv., 22.x.1992 (KT), 2 ♂, 9 ♀, 5 juv; Salajka National Nature Reserve nr. Bílá Village (790 m a.s.l.), preserved Carpathian fir-beech virgin forest, p.t. (KT), 19.ix.–31. x.2009, 1 juv. ♂; 31.x.–29. xi.2009, 1 ♀, 1 juv. ♀; 1 juv.; 29.xi.2009 – 18.iv.2010, 8 ♂, 14 ♀.

Poland. Bieszczady Mts.: Bieszczady National Park, Smerek Peak, northern slopes (1044 m a.s.l.), dwarf beech forest, soil sampling (KT), 6.x.2006, 1 ♀; Terebowiec Stream Valley nr. Ustrzyki Górne Village: fertile beech forest (789 m a.s.l.), 1 juv. ♀, subalpine bilberry shrubs (1,123 m a.s.l.), 1 juv. ♀, subalpine meadow (1,162 m a.s.l.), 1 ♂; all p.t. (KT) 10.vi.–6.x.2006, 6.x.2006 – 20.vi.2007.

Previously published records. Note: Altitude (in brackets) represents estimated approximate values or their range for published localization within the given territory or concrete locality.

Hungary. Aggtelek Karst: Nagyoldal at Jósvafő (305 m a.s.l.) ( Loksa 1962).

Slovakia. Cerová vrchovina Mts. (525–570 m a.s.l.): pseudokarst caves in basalt, Nyáryho Cave, Šurický úkryt Cave, Stĺpová Cave ( Papáč et al. 2009); Slovak Karst (495–880 m a.s.l.): Diviačia Chasm ( Kováč et al. 2005), Diviačia and Zvonica Caves ( Kováč et al. 2010b); Havrania skala Chasm, Silická ľadnica Cave ( Gulička 1960; Mock & Tajovský 2002), Silická Plateau above Brzotín ( Gulička 1951), Silická ľadnica Cave, Snežná diera Cave ( Gulička 1985); Low Tatra Mts. (800– 2,010 m a.s.l.): Králička, Salatín, Veľká Chochuľa, ( Gulička 1951), Demänovská Valley, Ďumbier, Chabenec, Chopok, Korytnica, Ludrovská Valley – Hučiaky, ( Gulička 1960); altitudinal transect from spruce forest (1,240 m a.s.l.) to alpine Vaccinetum in the Štiavnica Valley (at 1,790 m a.s.l.) ( Tajovský 1997); Muránska planina Mts. (550– 1,400 m a.s.l.): Kľak, Hrdzavá Valley, entrances of the caves and surrounding localities – Bobačka, Brestová, Dlhý vrch, Mokrá Poľana, Studňa, Šingliarka, Veľká lúka, Veľká Stožka, Zlatnica ( Gulička 1985); Tatra Mts. (1,200–2,200 m): “ Kohlbachthale bei Schmecks an der Tatra” = Studený potok Valley near Smokovec = type locality ( Verhoeff 1899), Roháčska Valley, Veľká Studená Valley, Zlomísk Valley, ( Jawlowski 1938), Gáflovka, Hincove plesá Lakes, Kôprovské sedlo, slope of Kôprovský Peak ( Gulička 1951), Bujačí vrch Peak, Furkota, Havran, Hviezdoň, Hlúpy, Jahnence, Javorová dolina Valley, Kamzík, Kolové pleso Lake, Kôprová dolina Valley, Košiare, Malá Studená dolina Valley, Mengusovské sedlo, Mlynická dolina Valley – Skok, Ostrva, Popradské pleso Lake, Skalnaté pleso Lake, Studená dolina Valley – Sesterské plesá Lakes and Zbojnícka chata, Velické pleso Lake, Veľká Svišťovka, Zadné Meďodoly, Zadné Jatky, Ždiarska Vidla ( Gulička 1960), Tomanova Valley – altitudinal transect ( Tajovský 1997), Nefcerka Valley ( Stašiov & Bitušík 2001), Tatra ( Moritz & Fischer 1973 – syntypes!), Tatra Mts. ( Kováč et al. 2010a); Tribeč Mts. (300 m a.s.l.): Velčice (Velsic) ( Verhoeff 1941; Dudich 1958); Veporské vrchy Mts. (950 m a.s.l.): Dobročský prales ( Gulička 1960); Vtáčnik Mts. (1,300 m a.s.l.): Vtáčnik Hill ( Gulička 1960); Zvolenská kotlina Basin (620 m a.s.l.): Ponická Huta ( Gulička 1960).

Uncertain published data (only juveniles or females): Slovakia. Kremnické vrchy Mts. (1,000 m a.s.l.): “bei Körmöcbánya und zwar Skalka” = Skalka pri Kremnici ( Verhoeff 1941); Bukovské vrchy Mts. (490 m a.s.l.): Nová Sedlica ( Ložek & Gulička 1962); Vihorlat Mts. (618 m a.s.l.): Morské oko ( Ložek & Gulička 1962).

Poland. Bieszczady Mts. (790– 1,160 m a.s.l.): Bieszczady National Park, Suche Rzeki, beechwood under Smerek Peak, Ustrzyki Górne, Terebowiec Valley, Szeroki Wierch Peak ( Tajovský & Wytwer 2010).

Description. Body colour yellowish to orange-brown, especially head and dorsal and lateral sides of prozonites, color lightening from anterior to posterior. Paranota with setae are paler. Sexual dimorphism in coloration was not observed. Specimens collected using the formalin or ethylene glycol traps often partially or completely without colour ( Fig 1 ). Material freshly collected in caves sometimes much paler compared to surface dwelling specimens (e.g. in the pseudokarst caves), but sometime fully coloured in cave environment, as shown by the specimens from the bottom of the chasms in Slovak Karst at a depth 20 to 100 m.

Measurements. Length: 8.3 mm (♂) / 8.4 mm (♀); width (15th body segment): 0.8 mm (♂) / 0.8 mm (♀); max. height: 0.7 mm (♂) / 0.8 mm (♀).

Adults with 30 pleurotergites in both sexes (the mature stadium IX). The head width = the width of body segment 3> 2> collum <4 <5 <6, from 6 to 24 more or less equal, caudal body part gently tapering from 24–25 segment toward telson.

Head triangular, with slightly flattened frons and convex cardines and stipetes, antennae medium-sized, total length 1.30–1.65 mm. Antennomeres densely covered with short setae and several elongated setae in basal and apical edges of antenomeres. Number of ocelli: 13/13 (lef/right side of head). Some exceptions like 12/13 or 13/15 were observed, but without dependence on habitat or geography.

Tergites with triplets of macrochaetae; macrochaetal angle up to 110°. Macrochaetae elongated (their length up to 300 µm) oriented partly cranialy and dorsomedially.

Males. Legs of the 1st and 2nd pairs reduced, the 3–7th leg pairs of male markedly robust ( Fig. 1 ) compared to other legs and all legs of female. Beside the gonopods, the 9th and 10th pairs of legs are modified and together with gonopods form a complex for storage and transportation of sperm ( Figs 2, 3 ).

Both gonopods (8th legs) create compact paired unit ( Figs 4–7, 10 ) with unpaired median process (mp) ( Figs 4, 5, 8 , 13 ). Each gonopod is composed from anterior slim branch (cheirite), laterally flattened and only gently laterally and posteriorly curved ( Figs 11, 12 ). Distally it ends with fimbriated tips (t). Small hook (h) below the tip is present on the inner side of anterior gonopod branch, moderately curved dorsally, and toothed on its internal edge. The inner side of the anterior branch below the hooked process bears massive, slightly protruding, brushyfilled (bf) with fine filamentous (distal half) and toothed (proximal half) surface structure. Three xyphoid undulated processes (xp) grow from the posterior base of the gonopod body. On the both lateral sides of massive gonopod base there are one or two long setae. Another two pairs of fields with setae are present on anterior part of the gonopod base: a pair of 3–4 longer setae medially and closer to gonopod branches and pair of fields with up to 10 shorter setae situated more laterally. Pair of rather short, thin, finger-like hyaline processes (hp) is present on the posterior edge of gonopod complex, between the gonopods and near the base of the median process ( Figs 8, 9 , 11, 12 ). Structural fields (sf) covered by small hemispherical protuberances are situated proximally on median sides of both gonopods ( Figs 9 , 12 ). Unpaired median plate or clasp (mp) is slightly shorter than the main gonopod branches, laterally flattened, goose-necked, in the middle extended laterally and fitted with small tubercles. Its tip is expanded, with frayed edges ( Figs 8, 10 , 13 ).

Ninth legs are in the shape of shortened and modified legs with tarsus almost totally, tibia partly reduced ( Figs 14 , 16, 17 ). Apical part of the femur and tibia are covered by long setae. Huge coxae are strongly depressed, forming concave holes at the medial bases by distinct sclerotized medial coxal processes (cp). Processes are broad and laterally flattened with smooth inner surface on their proximal parts ( Figs 16, 17 ), wrinkled between both femurs and as a complex structure they create and protect a channel for transporting spermatophores. Distal tips of coxal processes are orientated dorsally and backwards with two small teeth on apex.

Tenth legs ( Figs 15 , 18 ) are short and robust in comparison with following pairs of legs. On the 10th coxae, ladle-like long processes (lp) are present, and orientated like a roof above the median channel between the legs. On their basal inner side there are 2–3 setae, another sole seta is on the tip of the processes. The basal parts of each coxa are broad and equipped with sperm sacs (ss).

Females. All pairs of female legs developped, without any modification or reduction and without any prominent sexual characters, gradually increasing in length from cranial and caudal towards the medial pairs. Female body (in diameter) slightly broader.

Vulvae of females ( Figs 19 , 20 ). Both vulvae are compact and grown together. They have simple oval shape. Bursa (b) has fields of long setae on both lateral sides, growing in the field of dense curved stickers. Operculum (op) is short, flattened and sunk deeper behind the bursa and bears shorter setae in 1-2 irregular rows. Peculiar postvulval structures (ps; postgenital plates sensu Shear 1972) are developed in this species on the posterior side of the vulvae (Figs, 19, 20a). These structures consist of long, flattened, unpaired and apically divided median processes. Their ends are of a float-board shape, concave on anterior side and from external side covered by dense pyramid-like tubercles. The whole structure is movable, it can be deflected from vulvae as demonstrated in Fig. 19 .

Remarks on the previously described subspecies of Hylebainosoma tatranum . Based on the analysed material, no significant differences in gonopod morphology in H. tatranum were found corresponding to the geographic occurrence of particular populations within the known area of distribution, nor to habitat types (caves, surface, woods, alpine habitats). Therefore there seems to be no reason to recognize the previously described subspecies. Both Verhoeff (1941) and Loksa (1962) examined only very low numbers of specimens from isolated localities and in describing their subspecies, they focused only on characters of high variability from the whole distribution of the species.

Isolated populations of this species were recorded in two deep chasms in the Slovak Karst and in pseudokarst caves, but also without any morphological differences. Nevertheless, comparing to aboveground populations, we suppose at least partial adaptation to the cave environment such as permanent occurrence and presence of adults during the whole year (changes in chronobiology).

Hylebainosoma gulickai n. sp. Figs 1 , 21–38

Type material. Holotype: Slovakia, Muránska planina Mts., Tisovec Karst, Kostolík Cave, Sieň pagod and entrance hall, i.s., on wet wood debris, 29.xi.2012, leg.: A. Mock, V. Papáč, Ľ. Kováč, 1 ♂; ( NHMW No. 8210). Paratypes: Slovakia, Muránska planina Mts., Tisovec Karst, Kostolík Cave, Sieň pagod and entrance hall, i.s., on wet wood debris, 29.xi.2012, leg.: A. Mock, V. Papáč, Ľ. Kováč, 1 ♂, 1 ♀ ( NHMW No. 8211); Kostolík Cave, Oddychová sieňka Hall, i.s. on wood, 29.xi.2012, leg. V. Papáč, 1 ♂, 1 ♀; Michňová Cave, same karst region, bottom of the entrance shaft, i.s. on wood, leg. A: Mock, V. Papáč, Ľ. Kováč, 2 ♀ (NHM-SNM No. SZ 7468).

Other examined material. Slovakia, Muránska planina Mts., Tisovec Karst: Jaskyňa netopierov Cave (48° 40’ 31” N, 19° 52’ 52” E; altitude of entrance/length/depth of the cave: 601 m a.s.l./ 175 m /-69.5 m), i.s., in depth 35 m, 7.i.2006 (VP), 2 juv.; i.s., in depth 69.5 m, 7.i.2006 (VP), 3 ♀, 2 juv.; Kostolík Cave (48° 40’ 45” N, 19° 54’ 18” E; 480 m a.s.l./ 404 m /- 32 m), corridor Hrádzková-Kotlíková, i.s., on sediment, 8.iii.2006 (VP), 1 ♀; Oddychová sieňka Hall, i.s., on sediment, 8.III.2006 (VP), 1 ♂; Sieň pagod and entrance hall, i.s., on wet wood debris, 29.xi.2012 (AM, VP, LK), 1 ♀, 1 subad. ♂, 1 subad. ♀, 6 juv.; i.s., deeper corridors in the cave, 29.xi.2012 (AM, VP, LK), 1 ♀, 1 subad. ♀, 1 juv.; Michňová Cave (48° 40’ 45” N, 19° 52’ 53” E; 619 m a.s.l./ 336 m /- 72 m), p.t. at the cave bottom, 12.i.-11.iii.1998 (RM), 1 ♂, 2 ♀; on wood debries, i.s., 6.x.2004 (RM), 4 juv.; cave bottom, i.s., 27.x.2004 (AM, LK), 1 ♂, 2 juv. ♀; bottom of the Prvá priepasť Shaft, i.s., on the wood, 27.x.2004 (AM), 1 ♂, 1 ♀; bottom of the entrance shaft, i.s. on the wood, 27.x.2004 (AM), 3 ♀, 1 juv.; i.s., Veľká sieň Hall”, on the wood, 27.x.2004 (AM, LK), 4 juv.; bottom of the cave, i.s., wood, 16.vi.2005 (AM), 1 ♂, 1 juv.; Veľká sieň, i.s., wood, 16.vi.2005 (AM, PL), 1 ♂, 1 ♀, 3 juv.; dtto, (PL), 2 juv.; small side hall in the middle of cave, i.s., wood, 16.vi.2005 (AM), 1 ♀, 2 juv.; bootom of the entrance shaft, i.s., wood, 16.vi.2005 (AM), 1 ♀; Kámenov dóm Hall at the bottom of entrance shaft, i.s., wet wood: 6 ♀, 2 subad. ♂, 1 subad. ♀, 1 juv.; Rysie hniezdo Cave (48° 41’ 16” N, 19° 53’ 43” E; 770 m a.s.l./ 200 m /- 65 m), i.s., bottom of the cave, wood, (VP), 15.x.2007 (VP), 1 ♂; Teplica Cave (48° 40’ 43” N, 19° 54’ 32” E; 462 m a.s.l./ 1,000 m /-51.5 m), Žabia chodba, i.s. (VP), 20.ii.2006, 2 ♂, 3 ♀; i.s. (VP), 18.ii.2009, 1 juv. ♂;, i.s. (VP), 16.VI.2009, 1 subad. ♂.

In total, the whole collected and examined material involves 12 ♂, 5 subadult ♂, 22 ♀, 8 subadult ♀ and 28 juveniles.

All 5 cave localities are connected hydrologically (or paleohydrologically), with sinking waters and wet environment. Almost all specimens were found on very moist wood in the dark zone of the caves. Microclimate in caves: air temperature 5.8–6.8 °C, mean air relative humidity 95 %.

Diagnosis. The new species is strictly troglobiotic, eyeless and without coloration of the body. It differs from congeners by the structure of gonopods (simpler anterior branche, only two xyphoid posterior processes), by the shape of unpaired medial plate and strictly by coxal processes of the 9th legs, which are of broad elk horns shape. Female vulvae posses by large postvulval structures in the shape of broad medial plate and paired lobed lateral processes.

Description. Body completely without pigmentation and eyeless ( Figs 1 , 21 ).

Measurements. Length: 10.3 mm (♂) / 10.4 mm (♀); width (15th body segment): 0.9 mm (♂) / 1.0 mm (♀); max. height: 0.7 mm (♂) / 0.8 mm (♀).

Number of segments in adults: 30 as in preceding species. The head width = body segment 3> 2> collum <4 <5 <6, from 6 to 23 more or less equal, width of caudal body part gently tapering from 24–25 segment toward telson.

Head triangular with slightly convex frons and convex cardines and stipetes. Antennae about 1.63–1.72 mm long. Antennomeres densely covered with elongated fine setae and several long setae in basal and apical edges of antenomeres.

Tergites with triplets of elongated macrochaetae, macrochaetal angle up to 100°. The middle macrochaeta up to 470 µm long (around the 15th body segment), macrochaetae oriented dorsomedially and slightly caudaly.

Males. Legs of the 1st and 2nd pairs reduced, the 3–7th leg pairs of male more robust compared to other legs and all legs of female.

Gonopods (8th legs; Figs 22–30 ) are within the general pattern of congeners. They create a complex unit placed on a broad common basis (modified coxae, Figs 22–24, 27 ).

The anterior branch (cheirite) of gonopod is wider than posterior branches, slightly laterally flattened, before the end gently extended and topped with fibriform tail (t). The inner side of the anterior branch is fitted with brushy structure without any other processes or hooks, partly covered by brushy structure ( Figs 28, 29 ). Fibriform distal tail of anterior part (t) is moveable in a range of 90°, often either inclined apically (e.g. Fig. 26 ) or erected forward ( Figs 28, 29 ). The lower half of the anterior branch is provided on medial edge with prominent wide hook-like process (wp) oriented backwards ( Fig. 29 ). Posterior part of the gonopod is formed by two sword-like branches or processes (xp) with moderately curved tips ( Figs 28, 29 ). Gonopod processes as well as basal inner surfaces are smooth without any specific surface structures. Broad common basis of the gonopod complex is armed on the anterior (frontal) and lateral sides with setae arranged in three paired groups: dense alignment of 8–10 setae in more lateral position, group of 2–3 setae arranged more anteriorly and medially near the basis of median plate (mp), and one or two setae in the middle of both lateral margins of the basis. Small finger-like hyaline processes (hp) are present on the posterior edge of the gonopod complex. Slim unpaired median plate (mp), situated between gonopods, is almost as long as gonopod branches ( Figs 22, 23 ). This median plate (mp) is of keel-like triangular shape with the shortest side basal, laterally flattened, with rounded, narrow, unextended apex slightly armed by fine tubercles and scratches ( Figs 22–30 ).

Ninth legs are shortened and modified, their distal parts are strongly reduced, with femur and reduced tibia covered by long setae, as usually in the family Haaseidae . Coxae are robust with deep holes on medial sides covered by unusually short and broad coxal processes (cp) ( Figs 31 , 33, 34 ). The shape of coxal processes resemble elk horns or coxcombs, with denticles on anterior edges. Processes and basal part of coxae together enclose a narrow space, the route for spermatophores is on the outer side of processes between the distal parts of coxae ( Figs 33, 34 ).

Tenth legs (Figs, 32, 35) are of normal shape but slightly robust compared to following pairs of legs. The 10th coxae bear ladle-like elongated processes (lp) oriented medially and sperm sacs (ss) on the base. Beside the several setae in the basal part of coxa, another separate seta is on the top of each ladle-like process.

Females. Female legs as in preceding species. Body slightly longer and broader in width and height ( Fig. 21 ).

Vulvae of females ( Figs 36–38 ). Vulvae compact and grown together in one complex, both rounded or slightly broader than long. Operculum (op) is narrow, flattened and armed by short setae, both lateral sides covered by fine and short spines. Surfaces of both lateral and caudal edges of the bursa (b) are covered by dense fields of smaller or slightly elongated spines; long setae only on both lateral sites of the bursa. Postvulval structures developed on the posterior side of the vulvae. Beside the medial broad rounded postvulval plate (psm) there are present another two postvulval processes in the shape of paired elongated lobes (psl) erected laterally behind the vulvae complex ( Figs 36–38 ). These flattened and apparently less sclerotized processes (not stable in shape) are connected together on the base deeper under the vulvae ( Figs 37 , 38 b). Fine honeycomb structure is more developed on apical part of medial plate, on other parts of postvulval structures it is only weakly visible.

Etymology. The species is named after Ján Gulička (1925–2009), the Slovak myriapodologist and biospeleologist, one of the most important zoologists and promoters of ecological research in Slovakia in the past century.

Figures

FIGURE 1. General habitus of males of the genus Hylebainosoma: above— H. tatranum (Slovakia, Low Tatras Mts., Jánska Valley), in the middle— H. tatranum (Slovakia, Cerová vrchovina Mts., Nyáryho Cave), below— H. gulickai n. sp. (Slovakia, Muránska planina Mts., Michňová Cave). Material fixed in ethanol. Scale bar: 1 mm.

FIGURES 2 – 10. Hylebainosoma tatranum, male: Figs. 2 – 3: Slovakia, Cerová vrchovina Mts., Šurický úkryt, pseudokarstic cave; Figs. 4 – 10: Slovakia, Tatra Mts., Tomanova Valley. SEM photos. Fig. 2: Lateral view on the body section within the zone of the 5 th – 10 th leg pairs, with gonopods and the 9 th pair of legs, Fig. 3: Detail of gonopods in situ, ventral view, Fig. 4: Complex of gonopods, lateral view from left, Fig. 5: Gonopods from latero-anterior view, Fig. 6: | Gonopods, anterior view, Fig. 7: Gonopods, posterior view, Fig. 8: Right gonopod with median plate, median (inner) view, Fig. 9: Left gonopod, median view without median plate, Fig. 10: Complex of gonopods, ventral view. Scale bar: 100 Μm. Abbreviations: cp: Coxal processes, hp: Hyaline process, lp: Ladle-like processes of 10 th legs, mp: Median plate, sf: Structural field.

FIGURES 11 – 15. Hylebainosoma tatranum, male (Slovakia, Tatra Mts., Tomanova Valley). Fig. 11: Right gonopod, lateral view, Fig. 12: Right gonopod, medial view, Fig. 13: unpaired medial plate in (a) lateral and (b) anterior view, Fig. 14: Anterior view of the 9 th pair of legs, Fig. 15: Anterior view of the 10 th pair of legs. Scale bar: 100 Μm. Abbreviations: bf: – Brushy field, cp: Coxal process, h: Hooked process, hp: Hyaline process, lp: Ladle-like processes, sf: Structural field, ss: Sperm sac, t: Fimbriated tip, xp: Xyphoid processes.

FIGURES 16 – 19. Hylebainosoma tatranum: Figs. 16 – 18: Male (Slovakia, Tatra Mts., Tomanova Valley), Fig. 19: Female (Slovakia, Slovak karst, Diviačia Chasm). SEM photos. Fig. 16: 9 th pair of legs with coxal process curved posteriorly, anterior view, Fig. 17: Detail of coxal process of the 9 th pair of legs, ventro-lateral view, Fig. 18: 10 th pair of legs, anterior view, Fig. 19: Vulvae in ventral view. Scale bar: 100 Μm. Abbreviations: b: Bursa, cp: Coxal process, lp: Ladle-like process, op: Operculum, ps: Postvulval structures, ss: Sperm sacs.

FIGURE 20. Hylebainosoma tatranum, female (Slovakia, Tatra Mts., Tomanova Valley), undivided vulvae. Fig. 20 a: Ventral view, Fig. 20 b: Ventro-anterior view. Scale bar: 100 Μm. Abbreviations: b: Bursa, op: Operculum, ps: Postvulval structures.

FIGURE 21. Hylebainosoma gulickai n. sp., adult female, Slovakia, Kostolik Cave. Photo: Ľ. Kováč, A. Mock. Scale bar: 1 mm.

FIGURES 22 – 27. Hylebainosoma gulickai n. sp. (Slovakia, Michňová Cave), male. SEM photos. Fig. 22: Complex of gonopods, latero-anterior view, Fig. 23: Gonopods, anterior view, Fig. 24: Gonopods, lateral view from right, Fig. 25: Gonopods, posterior view, Fig. 26: Left gonopod, median view without median plate, Fig. 27: Complex of gonopods, ventral view. Scale bar: 100 Μm. Abbreviation: mp: Median plate.

FIGURES 28 – 32. Hylebainosoma gulickai n. sp. (Slovakia, Michňová Cave), male. Fig. 28: Right gonopod, lateral view, Fig. 29: Right gonopod, medial view, Fig. 30: Unpaired medial plate, lateral view, Fig. 31: Anterior view of the 9 th pair of legs, Fig. 32: Anterior view of the 10 th leg pair. Scale bar: 100 Μm. Abbreviations: cp: Coxal process, hp: Hyaline process, lp: Ladle-like processes, ss: Sperm sac, t: Fimbriated tip, wp: Wide hook-like process, xp: Xyphoid processes.

FIGURES 33 – 35. Hylebainosoma gulickai n. sp. (Slovakia, Michňová Cave), male. SEM photos. Fig. 33: 9 th leg pair with broad toothed coxal process, anterior view, Fig. 34: Detail of coxal processes of the 9 th leg pair, ventro-lateral view, Fig. 35: Left leg of the 10 th pair, anterior view. Scale bar: 100 Μm. Abbreviations: cp: Coxal process, lp: Ladle-like process, ss: Sperm sacs.

FIGURES 36 – 37. Hylebainosoma gulickai n. sp. (Slovakia, Michňová Cave), female. SEM photos. Fig. 36: Vulvae in ventral view, Fig. 37: Left vulva, posterior view. Scale bar: 100 Μm. Abbreviations: b: Bursa, op: Operculum, psl: Postvulval lateral process, psm: Medial postvulval process.

FIGURE 38. Hylebainosoma gulickai n. sp. (Slovakia, Michňová Cave), female. Vulvae in (a) ventral and (b) posterior view. Scale bar: 100 Μm. Abbreviations: b: Bursa, op: Operculum, psl: Postvulval lateral process, psm: Medial postvulval process.