Anthroherpon Reitter, 1889

Čeplík, Dávid & Hlaváč, Peter, 2022, A contribution to understanding the morphology of the genus Anthroherpon (= Parantrophilon syn. nov.) and new combination for a species from the Balkan Peninsula (Insecta, Coleoptera, Leiodidae, Cholevinae, Leptodirini), Zootaxa 5115 (4), pp. 541-558 : 542-554

Genus Anthroherpon Reitter, 1889

( Figs 1–34 )

Anthroherpon Reitter, 1889: 294 . Type species: Leptoderus cylindricollis Apfelbeck 1889: 61 , subsequent designation.

Anthroherpon Reitter : Newton, 1998: 170; Perreau, 2000: 153; Giachino & Vailati 2005: 150; Perreau & Pavićević 2008: 230; Perreau, 2015: 200; Njunjić, Perreau & Pavićević 2015: 404; Njunjić et al. 2016: 337; Hlaváč, Perreau & Čeplík 2017: 78; Njunjić et al. 2018: 1.

Antroherpon Reitter: Apfelbeck, 1894: 511 (misspelling).

Antroherpon Reitter: Jeannel, 1910: 26; Jeannel, 1911: 546; Jeannel, 1914: 53; Jeannel, 1924: 400; Jeannel, 1930: 127; Pretner, 1968: 38; Guéorguiev, 1976: 31; Pretner, 1977: 142; Guéorguiev, 1990: 242; Giachino & Guéorguiev, 1993: 288.

Antroherpon (Eumecosoma) Müller, 1901: 29. Type species: Antroherpon stenocephalum Apfelbeck 1901: 15 , by monotypy, synonymy in Müller, 1904: 40.

Antrophilon Absolon, 1913: 100. Type species: Antrophilon primitivum Absolon, 1913: 101 , original designation, synonymy in Müller, 1914: 1028.

Euantroherpon Absolon, 1913: 108. Type. species: Leptoderus hoermanni Apfelbeck, 1889: 62 , subsequent designation, synonymy in Jeannel, 1914: 54.

Parantrophilon Noesske, 1914: 17 . Type species: Parantrophilon spelaebatoides Noesske, 1914: 21 , by monotypy. New synonymy.

Protantroherpon Absolon, 1913: 108. Type species: Leptoderus cylindricollis Apfelbeck, 1889: 61 , subsequent designation, synonymy in Jeannel, 1914: 54.

Note: This paper does not represent a revision of the genus Anthroherpon , therefore only generic synonyms and main references (monographs, revisions, phylogenetic studies, catalogues) are listed here. We also provide all cases of misspellings of the generic name. For all species names, references and citations see Perreau (2000: 153–164; 2015: 200–202) and Hlaváč, Perreau & Čeplík (2017: 78–84).

Diagnosis. A member of the subterranean tribe Leptodirini , subtribe Anthroherponina defined by the following combination of characters: leptodiroid body shape, antennae inserted in the posterior quarter of the head, scape longer than pedicel, pronotum slightly to strongly elongate with or without constriction in posterior third, mesoepiventrite slightly or strongly elongate latero–anteriorly, elytra elliptically variable, mesoventral process from short to strongly elongate, sharp or rounded, protarsi with five tarsomeres in males and four in females, claws well developed, aedeagus relatively small, endophallus lacking sclerotized structures, female sternite VIII (spiculum ventrale) with anterior expansion.

Redescription. Leptodiroid body shape ( Figs 1–5 ), BL: 3.7–8.3 mm. Head hypognathous ( Figs 6 , 8, 9 ), longer than wide (HL1/HW: 1.8–2.2; HL2/HW: 1.4–1.8); head longer than, or approximately of the same length as pronotum (HL1/PL: 1.1–1.6); head from slightly narrower than pronotum to slightly wider than pronotum (HW/PW: 0.9–1.2). Head widest approximately at midlength of HL2 or before level of clypeofrontal suture. Eyes completely absent, with posterior neck region and genae either glabrous or covered with several fine setae. Transverse occipital crest weakly defined. Antennae inserted approximately in the posterior first quarter of HL1. Clypeus subhexagonal, clypeofrontal suture distinct, clypeus and labrum setose. Head densely or sparsely pubescent, punctation dense and relatively deep or sparse and shallow. Mandibles relatively robust, sharply pointed, apical inner margin toothed with several small teeth. Mandibles either with apical incisor or with apical and subapical incisors. Terminal maxillary palpomere conical, apically pointed, slender and shorter than the longer and wider penultimate palpomere ( Figs 7 , 8, 9 ). The ventral surface of the head setose ( Jeannel, 1911: 12: Fig. IV). Gula almost glabrous, gular sutures well defined, genae with transversal wrinkles, glabrous or with several fine setae. Submentum almost confluent with gula, separated from mentum by transverse suture. Submentum covered with several longer, medium sized and fine setae. Mentum with some longer, medium sized and fine setae. Cardo glabrous or with one–two fine setae, basistipes and mediostipes subtriangular, covered with several fine setae. All antennomeres longer than wide and pubescent, the 1st antennomere longer than the 2nd, the 2nd shorter than the 3rd, the 3rd longer than the 4th, the 4th shorter than the 5th, the 5th longer or as long as the 6th, the 6th shorter or longer than the 7th, the 7th shorter or longer than the 8th, the 8th shorter or longer than the 9th, the 9th longer or as long as the 10th, and the 10th shorter or as long as the 11th. Antennal length shorter (several species) or longer (most of species) than the body length.

Pronotum ( Figs 1–5 , 10, 11 ) elongate (PL/PW: 1.2–2.2). Lateral sides convex, considerably variable with or without constriction in posterior third of pronotal length. Lateral rim of pronotum absent or limited approximately to posterior half of pronotal length. Posterior third of pronotum, forms due to a constriction in some species, a prothoracic peduncle, sensu Jeannel. Pronotum widest approximately before or after the anterior third of pronotal length, the narrowest at its base or at the constriction. Dorsal surface sparsely or relatively densely pubescent with shallow and sparse punctation. Anterior and posterior angles weakly defined, obtuse or acute. Prosternum variable according to elongate prothorax, prosternal surface and hypomeron almost glabrous. Procoxal cavities ( Fig. 12 ), median edge weakly defined, more or less visible only if procoxae are free of trochanters.

Venter ( Figs 14, 15–21 ). Mesoventral surface glabrous or slightly pubescent, mesoventral suture absent or weakly defined. Mesoventro–mesoepiventral suture relatively well defined or not and interrupted. Mesoepiventrite latero–anteriorly slightly or strongly elongate ( Figs 15 , 17, 20 ), its latero–anterior part reaches level of posterior part of transverse collar and level of mesonotum. Scutum and scutellum taken together form an escutcheon, sensu Jeannel. Anterior part of elongate mesoepiventrite is more or less visible from dorsal aspect of the habitus ( Figs 2, 3 ) or not ( Figs 1 , 4, 5 ) but more or less visible from latero–ventral aspect and if venter is free of elytra and pronotum ( Figs 15 , 20 ). Scutum and scutellum ( Fig. 13 ) variable, rather “U”shaped. A dorsal sclerite posterior to the scutellum present but weakly defined (= postscutellum, sensu Jeannel). Mesoepimeron slightly or strongly elongate. Mesoepimeral surface almost glabrous. Mesocoxal cavities apparently confluent, separated only by narrow carina, which is visible only when legs are removed, mesoventral process almost absent (intraspecific variability of A. stenocephalum discussed by Njunjić et al. 2016)) or strongly variable in length ( Figs 14 , 22–27 ), from short to long. Metaventral surface pubescent. Metacoxae separated by bifid posterior metaventral process. Sternites pubescent with short or medium sized fine setae.

Metanotum ( Fig. 13 ) slightly variable, reduced to a transverse narrow arc (= etroite bandelette transversale, sensu Jeannel). Metendosternite slightly variable, rather “Y”shaped ( Fig. 28 ).

Elytra elongate, elliptically variable ( Figs 1 , 2–5 ), EL/EW: 1.5–2. Punctation from shallow to deep, from sparse to dense. Pubescence from short to long. Punctation and pubescence is species characteristic. Elytral surface with or without microreticulation between punctures. Elytra widest approximately before or after mid–length, apex rounded, longitudinal parasutural striae absent. Elytra wider than pronotum.

Legs very long, from slender to thicker and pubescent. Protarsi with five undilated segments in males and four undilated segments in females. Mesotarsi and metatarsi pentamerous in both sexes. Tarsal empodium with two setae. Claws well developed, more or less robust and curved. Protibiae straight or slightly curved inwards. Mesotibiae and metatibiae relatively narrow and straight or only slightly curved. Internal subapical side of tibiae with one or two spurs (visible from ventral aspect).

Aedeagus relatively small ( Figs 29–30 ; Jeannel, 1924: 410, 412: Figs 465–474, Figs 476, 477, 479–481; Njunjić, Perreau & Pavićević 2015: 407: Figs 6–11 ). Aedeagus approximately 0.4–0.8 mm long. Median lobe elongate, from thiner to thicker shape, with rounded or pointed apex. Endophallus of median lobe lacking sclerotized structures. Parameres variable, shorter than median lobe or slightly longer than median lobe, relatively thin or slightly expanded, straight or slightly to strongly curved from dorsal and lateral aspects, straight or slightly to strongly inward apically, apex with three setae.

Female sternite VIII pubescent, anterior expansion developed ( Fig. 33 ). Female sternite IX ( Fig. 34 ) with reduced sclerotized parts. Spermatheca ( Figs 31–32 ) curved, weakly sclerotized, rather of “C”shape.

Sexual dimorphism. Protarsi with five undilated segments in males and four undilated segments in females.

Distribution. Croatia (single species from Sniježnica Mts.), Bosnia and Herzegovina, Montenegro and Albania (several species from Prokletije Mts.), (Central and Southern Dinarides), ( Fig. 38 ).

Remarks. The genus Anthroherpon is distinguished from the morphologically similar genera Leptomeson and Graciliella (subtribe Anthroherponina ) as follows:

• from Leptomeson : by lacking sclerotized structures in median lobe of the aedeagus versus having sclerotized structures in Leptomeson . Additionally, by less elongate anterior part of the mesoepiventrite (= either more or less visible from dorsal aspect of the habitus or more or less visible only from lateral aspect if venter is free of elytra and pronotum) versus extremely elongate in Leptomeson (= always visible from dorsal aspect of the habitus).

• from Graciliella : by less elongate anterior part of the mesoepiventrite (= either more or less visible from dorsal aspect of the habitus or more or less visible only from lateral aspect if venter is free of elytra and pronotum) versus more elongate in Graciliella (= always visible from dorsal aspect of the habitus) and by variable shapes versus relatively constant shape of the pronotum with constriction in the posterior third in Graciliella .

Additionaly, the genus Anthroherpon is distinguished from the genus Spelaeobates (subtribe Spelaeobatina ) as follows:

• by having protarsi with five tarsomeres in males against four tarsomeres in males in Spelaeobates ; by the first antennomere longer than the second against the first antennomere approximately of the same length as the second in Spelaeobates ( Fig. 36 ). Spelaeobates also differs by its construction of the pronotum ( Fig. 35 ) and slightly different maxillae ( Fig. 37 ).

Note: Some morphological convergences–parallelisms (e.g., in general slightly similar habitus ...) among phyletically different taxa, belonging to different subtribes but inhabiting the same or very similar subterranean niches can be found here as examples of the convergent evolution of taxa.

Figures

FIGURE 1. Anthroherpon spelaebatoides Noesske, 1914, scale bar 3 mm, original by D. Beňo, ex Hlaváč, Perreau & Čeplík 2017.

FIGURES 2–5. 2: Anthroherpon cylindricollis cylindricollis Apfelbeck, 1889, scale bar 4.3 mm. 3: Anthroherpon zariquieyi Jeannel, 1930, scale bar 3.8 mm. 4: Anthroherpon primitivum primitivum Absolon, 1913, scale bar 4.5 mm. 5: Anthroherpon piesbergeni Zariquiey, 1927, scale bar 4.6 mm. All originals by D. Beňo, ex Hlaváč, Perreau & Čeplík 2017.

FIGURES 6–7. Anthroherpon spelaebatoides Noesske, 1914. 6: head, dorsal aspect, scale bar 0.64 mm; 7: right maxilla, dorsal aspect, scale bar 0.32 mm.

FIGURES 8–11. 8: Anthroherpon primitivum primitivum Absolon, 1913, head, dorsal aspect, scale bar 0.38 mm; 9: Anthroherpon piesbergeni Zariquiey, 1927, head, dorsal aspect, scale bar 0.43 mm; 10: Anthroherpon spelaebatoides Noesske, 1914, pronotum, right lateral aspect, scale bar 0.60 mm; 11: Anthroherpon piesbergeni Zariquiey, 1927, pronotum, right lateral aspect, scale bar 0.90 mm.

FIGURES 12–15. Anthroherpon spelaebatoides Noesske, 1914. 12: pronotum, ventral aspect, scale bar 0.70 mm; 13: mesono- tum—scutum, scutellum, metanotum, scale bar 0.44 mm; 14: venter, scale bar 0.70 mm; 15: mesothorax, right latero–ventral aspect, without scale bar.

FIGURES 16–21. 16: Anthroherpon cylindricollis cylindricollis Apfelbeck, 1889, venter, without scale bar, 17: mesothorax, right latero–ventral aspect, without scale bar; 18: Anthroherpon zariquieyi Jeannel, 1930, venter, without scale bar; 19: Anthroherpon primitivum primitivum Absolon, 1913, venter, without scale bar, 20: mesothorax, right latero–ventral aspect, without scale bar; 21: Anthroherpon piesbergeni Zariquiey, 1927, venter, without scale bar.

FIGURES 22–27. Mesoventral process. 22: Anthroherpon cylindricollis cylindricollis Apfelbeck, 1889; 23: Anthroherpon hossei Winkler, 1925; 24: Anthroherpon piesbergeni Zariquiey, 1927; 25: Anthroherpon primitivum primitivum Absolon, 1913; 26: Anthroherpon stenocephalum stenocephalum Apfelbeck, 1901; 27: Anthroherpon zariquieyi Jeannel, 1930.

FIGURES 28–32. Anthroherpon spelaebatoides Noesske, 1914: 28: metendosternite, dorsal aspect, scale bar 0.95 mm, 29: aedeagus, dorsal aspect, scale bar 0.46 mm, 30: median lobe, right lateral aspect, scale bar 0.34 mm, 31: spermatheca, scale bar 0.08 mm; 32: Anthroherpon cylindricollis cylindricollis Apfelbeck, 1889, spermatheca, scale bar 0.19 mm.

FIGURES 33–37. Anthroherpon spelaebatoides Noesske, 1914: 33: female sternite VIII, dorsal aspect, scale bar 0.56 mm; 34: female sternite IX, dorsal aspect, scale bar 0.23 mm. Spelaeobates novaki Müller, 1901: 35: pronotum, right lateral aspect, without scale bar; 36: scape and pedicel, without scale bar; 37: left maxilla, dorsal aspect, without scale bar.

FIGURES 38–40. 38: Black dot on the map shows location of Grebci, Bosnia and Herzegovina; 39: Grab(r)ovica pećina cave, topographical map, ex Malez, 1970; 40: The entrance of Grab(r)ovica.