Pholeuonopsis Apfelbeck, 1901

Čeplík, Dávid, 2021, Restoring an old concept of Pholeuonopsis (= Blattodromus syn. nov.) and new recombination for one species from the Balkan Peninsula (Insecta, Coleoptera, Leiodidae, Cholevinae, Leptodirini), Zootaxa 5016 (4), pp. 559-570 : 560-562

Genus Pholeuonopsis Apfelbeck, 1901 .

Pholeuonopsis Apfelbeck, 1901: 14 . Type species: Pholeuonopsis ganglbaueri Apfelbeck, 1901: 14 , by monotypy.

Pholeuonopsis Apfelbeck : Reitter, 1904: 153; Apfelbeck, 1907a: 402; Apfelbeck, 1907b: 304; Jeannel, 1910: 40; Jeannel, 1911: 461; Breit, 1913a: 357; Breit, 1913b: 309; Breit, 1914: 61; Reitter, 1913: 155; Jeannel, 1914: 41; Jeannel, 1924: 255; Zariquiey, 1927: 157; Knirsch, 1927: 103; Knirsch, 1928: 92; Jeannel, 1930: 228; Pretner, 1968: 17; Guéorguiev, 1976: 101; Perreau, 2000: 238; Ćurčić & Brajković, 2002: 43; S. Ćurčić et al. 2006: 497; S. Ćurčić et al. 2014: 178; Perreau, 2015: 209; S. Ćurčić et al. 2015: 394 ; Hlaváč et al. 2017: 96; S. Ćurčić et al. 2018: 539.

subgenus Silphanillus Reitter, 1903: 210 . Type species: Pholeuonopsis (Silphanillus) leonhardi Reitter, 1903: 211 , by monotypy.

subgenus Scotosites Knirsch, 1928: 88 . Type species: Pholeuonopsis (Scotosites) spaethi Knirsch, 1928: 88 , original designa- tion.

Blattodromus Reitter, 1904: 153 . Type species: Pholeuonopsis (Blattodromus) herculeana Reitter, 1904: 153 , original designation. New synonymy.

Blattodromus Reitter : Breit, 1913b: 308; Jeannel, 1914: 42; Jeannel, 1924: 264; Pretner, 1968: 19; Pretner, 1970: 158; Guéorguiev, 1976: 101; Perreau, 2000: 235; Perreau, 2015: 206; Hlaváč et al. 2017: 92.

Serbopholeuonopsis S. Ćurčić & Boškova, 2002: 11. Type species: Pholeuonopsis cvijici S. Ćurčić & Brajković, 2002: 43 , by monotypy, synonymy in S. Ćurčić et al. 2015: 399.

Diagnosis. A member of the subterranean tribe Leptodirini , subtribe Bathysciina , group Théléomorphes, division II sensu Jeannel 1924, defined by: pholeuonoid body shape, pronotum bell–shaped and very variable, with maximum width approximately in the middle of the length, or maximum width on its base on the hind angles level and variable also in the length and shape of hind angles, protarsi with four tarsomeres in both sexes, protibiae lacking comb (pecten) along lateral external margin, lacking external apical spurs, inner sac (endophallus) of median lobe contains sclerotized structures.

Redescription. BL: 3.50–5.90 mm. Head ( Figs 5, 6 ) longer than wide if head not retracted in pronotum, eyes completely absent, with posterior neck region and genae more or less glabrous. Transverse occipital crest present but weakly–defined. Antennae inserted approximately in the middle of the median third of the head. Clypeus rectangular, clypeofrontal suture distinct, frontoclypeus and labrum densely setose. Mandibles relatively robust, with proximal and subapical incisors and a tooth or teeth between. Terminal maxillary palpomere conical, apically pointed, slender and much shorter than the wide penultimate palpomere. The ventral surface of head (not illustrated) setose but of lower density if compared with dorsal surface. Gula glabrous, gular sutures well defined, genae with some transversal wrinkles and several lateral setae. Submentum separated from gula, separated from mentum by transverse suture. Submentum covered with numerous medium sized and fine setae. Mentum with some longer, medium sized and some thin setae. Cardo with several fine setae, transverse, subtriangular, basistipes and mediostipes subtriangular, basistipes with several fine setae. All antennomeres longer than wide and pubescent. Length/maxi- mum width ratio of antennomeres is species characteristic.Antennal length approximately 0.6–0.8 of the length (L). Antennomere 8 subglobose or cylindrical, from slightly to strongly longitudinal.

Pronotum ( Figs 3, 4 ) bell–shaped and very variable, wider than long. Pronotum widest in mid–length, or mid– width same as width in posterior corners, or widest in posterior corners. Hind corners variable, from obtuse to acute, slightly or strongly protruding backwards and more or less pronounced. Dorsal surface pubescent. Pronotal disc with two basal shallow impressions. Prosternal surface and hypomeron almost glabrous, only slightly pubescent with several fine setae. Procoxal cavities separated by narrow prosternal process (not illustrated). Prosternal process with medial notch forming projection–edge between procoxal cavities, fully visible only if procoxae are free of trochanters, with more or less pronounced median edge.

Elytra ( Figs 1, 2 ) oval, slightly variable, more or less elongate, pubescent, with maximum width approximately at mid–length, apex rounded, longitudinal parasutural striae absent. Elytra wider than pronotum. Marginal furrows distinct, relatively shallow, gradually narrowing towards elytral apex. Punctation more or less dense and deep or shallow and weak, weak punctation and pubescence on apical part of elytra. Scutellum triangular and less pubescent.

Venter ( Figs 7, 8 ). Mesoventral and metaventral surface sparsely pubescent. Mesocoxal cavities separated, mesoventral process well developed. Metacoxae separated by bifid posterior metaventral process. Mesoventral carina well developed, very variable ( Figs 9, 10 ), prolonged posteriorly, not extended over metaventrite, with backwardly oriented setae. Sternites with short and fine setae.

Metatergal apparatus ( Figs 12, 14 ) slightly variable, with relatively narrow and straight metascutum, than slightly curved, alacrista with moderately long basal apophysis. Metendosternite slightly variable, rather of “Y”shape ( Figs 11, 13 ).

Protarsi with four undilated segments in both sexes, first protarsomeres slightly longer than second and third, fourth longest. Mesotarsi and metatarsi pentamerous in both sexes. Claws relatively wide and well developed, empodiums with one bifurcate seta. Protibiae more or less straight or slightly curved, armed with spines, internal subapical side with one shorter and one longer subapical multi–toothed spurs. Mesotibiae and metatibiae relatively narrow and straight, armed with lateral spines, with apical crown of spines (two extrenal spines distinctly longer) of unequal length ( Gnaspini et al. 2020) and two internal apical multi–toothed spurs.

Aedeagus ( Figs 15–17 , 22–24 ) more or less large and relatively robust, elongate, sclerotised. Endophallus lobe elongate, tubular and with developed chitinized structures along its length. Median lobe from dorsal aspect sub–parallel, slightly widened and distinctly narrowed towards rounded apex which is beak–shaped and more or less sharp. Median lobe in lateral aspect elongate, almost straight basally, more or less curved anteriorly, gradually more or less narrowing apically with pointed beak–shaped apex. Parameres ( Figs 15, 18 , 22, 25 ) shorter than median lobe or slightly longer than median lobe, relatively thin, slightly expanded apically, more or less straight in dorsal and lateral aspect or curved inward apically, apex with three or four setae. Endophallus consists of sclerotized structures along its length: reinforcement bands, featherlike structures, connection nodules and basal phanera, with special differences according to species. Median lobe laterally pointed, apical slerotized structures more or less protruding from the median lobe.

Female ventrite VIII pubescent with anterior expansion stout ( Fig. 19 ). Each style with four setae, stylus cylindrical, with one long seta ( Fig. 20 ). Spermatheca ( Figs 21 , 26, 27, 28 ) sclerotized, slightly or strongly curved and more or less narrowing apically, with basal, medial and apical regions.

Distribution: Bosnia and Herzegovina, Serbia and Montenegro (Central and Southern Dinarides; Fig. 30 ).

Remarks: The genus Pholeuonopsis is currentlyclassified in the Leonhardella phyletic line ( Jeannel, 1911; 1924). The genus is distinguished from the morphologically most similar genera Anillocharis , Blattochaeta and Leonhardella as follows:

• from Anillocharis : by having protarsi with four undilated tarsomeres in both sexes against two dilated protarsomeres in males in Anillocharis . Additionaly both genera differ each other by the different general shape of the aedeagus.

• from Blattochaeta : by the pholeuonoid body–shape with bell–shaped and extremely variable pronotal disc against the bathysciod body shape with evenly convex and trapezoidal shape of the pronotal disc in Blattochaeta . Adittionaly Blattochaeta shows some similarities with Pholeuonopsis on the shape and size of the aedeagus, but the structure of the internal sac is slightly different.

• from Leonhardella : by the shape of elytra, more or less inflated against more or less flattened in Pholeuonopsis ; by the shape of metendosternite with a shorter basal part against comparatively longer basal part in Pholeuonopsis and by alacrista (metatergal apparatus) with comparatively shorter basal apophysis against longer basal apophysis in Leonhardella . Additionaly Pholeuonopsis and Leonhardella share some similarities on the shape of the aedeagus but the genus Leonhardella has the apex of the median lobe slightly prolonged.

Reasons supporting the placement of Blattodromus inside Pholeuonopsis :

• The body size, the antennal length, proportions of antennomeres 8, the shape of the mesoventral carina is in general extremely variable even at the species level in numerous genera of Leptodirini e.g., Anthroherpon , Blattochaeta , Drimeotus , Leonhardella , Pholeuon , Sophrochaeta etc... and therefore they can not be considered as diagnostic characters for the validation of Blattodromus as a valid genus.

• The pronotum varies considerably within the whole genus Pholeuonopsis . Blattodromus with its differences on hind angles, which are acute and strongly protruding backwards, more pronounced compared with that of other Pholeuonopsis species can be considered only as a diagnostic character for the definition of species within the genus.

• The aedeagus of Blattodromus is in general of very similar shape and shows similar internal structures as in other species of Pholeuonopsis . The diferenrences like larger size or slightly longer parameres can be considered only as a species level diagnostic characters. Parameres in Blattodromus are bearing three (also mentioned in Pretner, 1970) or four apical setae, it is not unusual that one paramera has three setae while another one has four. Therefore these characters cannot be used as a diagnostic character for the genus.

• The spermatheca in some Blattodromus females is slightly curved ( Fig. 21 ) or curved, usual character state than in Pholeuonopis ( Figs 26–28 ). These differences can be considered as the intra–species morphological variabilities.

Additionaly, no other significant differences between Blattodromus and Pholeuonopsis were observed in the comparison with morphologicaly close genera mentioned above.

Figures

FIGURE 30. Distribution map of the genus Pholeuonopsis.

FIGURES 1–6. Pholeuonopsis (s. str.) herculeana Reitter, 1904, 1: habitus, dorsal aspect, scale bar 1.25 mm, original by P. Krásenský, ex Hlaváč et al. 2017: 92; 3: pronotum, dorsal aspect, scale bar 1.88 mm; 5: head, dorsal aspect, scale bar 0.80 mm. Pholeuonopsis (s. str.) grabowskii grabowskii Apfelbeck, 1907a, 2: habitus, dorsal aspect, scale bar 0.97 mm, original by P. Krá- senský, ex Hlaváč et al. 2017: 96; 4: pronotum, dorsal aspect, scale bar 1.20 mm; 6: head, dorsal aspect, scale bar 0.60 mm.

FIGURES 7–14. Pholeuonopsis (s. str.) herculeana Reitter, 1904, 7: venter, dorsal aspect, scale bar 0.89 mm; 9: mesoventral carina, right lateral aspect, scale bar 0.23 mm; 11: metendosternite, dorsal aspect, scale bar 1.40 mm; 12: metatergal apparatus, dorsal aspect, scale bar 0.94 mm. Pholeuonopsis (s. str.) grabowskii grabowskii Apfelbeck, 1907a, 8: venter, dorsal aspect, scale bar 0.62 mm; 10: mesoventral carina, right lateral aspect, scale bar 0.20 mm; 13: metendosternite, dorsal aspect, scale bar 0.51 mm; 14: metatergal apparatus, left side and medial part, dorsal aspect, scale bar 0.62 mm.

FIGURES 15–21. Pholeuonopsis (s. str.) herculeana Reitter, 1904, 15–16: aedeagus, dorsal and left latero–ventral aspects, scale bar 1.37 mm; 17: median lobe apex, ventral aspect, scale bar 0.44 mm; 18: paramera apex, dorsal aspect, scale bar 0.03 mm; 19: female ventrite VIII, dorsal aspect, scale bar 1.00 mm; 20: style, dorsal aspect, scale bar 0.35 mm; 21: spermatheca, dorsal aspect, scale bar 0.20 mm.

FIGURES 22–29. Pholeuonopsis (s. str.) grabowskii grabowskii Apfelbeck, 1907a, 22–23: aedeagus, dorsal and left latero– ventral aspects, scale bar 0.86 mm; 24: median lobe apex, ventral aspect, scale bar 0.19 mm; 25: paramera apex, dorsal aspect, scale bar 0.02 mm. 26: redrawing of spermatheka of Pholeuonopsis (s. str.) tarensis, from Ćurčić & Pavićević 2018 in S. Ćurčić et al. 2018. 27: redrawing of spermatheka of Pholeuonopsis (s. str.) sljivovicensis, from S. Ćurčić et al. 2015. 28: redrawing of spermatheka of Pholeuonopsis (s. str.) perucensis, from S. Ćurčić et al. 2014. 29: Vilina pećina cave, on the entrance, Lebršnik Mts., Bosnia & Herzegovina.