Hypocaccus (Nessus) rubripes ( Erichson, 1834 )

Lackner, Tomáš & Seres, Gábor, 2018, Revision of the subgenus Nessus of the genus Hypocaccus from Central Europe, with description of a new species (Coleoptera: Histeridae), Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) 58 (2), pp. 419-439 : 428-432

Hypocaccus (Nessus) rubripes ( Erichson, 1834)

( Figs 25–31 , 33–39 )

Saprinus rubripes Erichson, 1834: 193 (original description).

Hypocaccus (Nessus) rubripes: MAZUR (2011) : 209 (new combination); LACKNER et al. (2015): 118 (catalogue). For complete references see LACKNER (2010): 134 –135.

Saprinus granarius Erichson, 1834: 191 (original description). Fauvel in GOZIS (1886): 202 (synonymy).

Saprinus arenarius Marseul, 1855: 691 (original description). SCHMIDT (1885): 313 (synonymy).

Saprinus corsicus Marseul, 1855: 688 (original description). BICKHARDT (1910): 104 (synonymy).

Saprinus rubripes var. clermonti Auzat, 1920: 4 (original description).

Note. Complete synonymies and literature references of this species are given in LACKNER (2010:134–135) and the reader is referred to them there. For the sake of completeness, and the full list of synonymies examined, however, above we list the references of all original descriptions and the works, in which they were synonymised.

Type material examined. Saprinus rubripes Erichson, 1834 . SYNTYPES: 1 ♁ ( Fig. 25 ), originally pinned with a pin-hole in its right elytron, glued onto a rectangular mounting card, genitalia examined and photographed by the senior author, unfortunately subsequently lost, ‘49207 [printed] // Hist. -Coll. ( Coleoptera ) / Nr. 49207 / Saprinus rubripes Er. x / Sardin. -Lusitan / Zool. Mus. Berlin [black-framed, printed label] // SYNTYPE / Saprinus rubripes / Erichson, 1834 / labelled by MFNB 2016 [red label, printed] // rubripes / Er. / Lusit. Hoffm [black-framed, written label]’ ( MFNB); 1♀, glued onto a rectangular mounting card, originally pinned with a pin-hole in its right elytron, genitalia extracted and glued onto the same mounting card as the specimen, ‘22 [written] // 49207 [written] // Hist. -Coll. ( Coleoptera ) / Nr. 49207 / Saprinus rubripes Er. x / Sardin. -Lusitan / Zool. Mus. Berlin [black-framed, printed label] // SYNTYPE / Saprinus rubripes / Erichson, 1834 / labelled by MFNB 2016 [red label, printed]’ ( MFNB); 1♀, left protibia missing, left mesotarsus missing, left metatibia missing, glued onto a rectangular mounting card with the labels identical to those of the preceding syntype ( MFNB). According to the MFNB staff, the lectotype designation of this species was not allowed, hence no lectotype was designated.

Saprinus granarius Erichson, 1834 . SYNTYPE: 1 ♁ ( Fig. 26 ), originally pinned with a pin-hole in its right elytron, glued onto a rectangular mounting card, left antennal flagellum, left protarsus and right metatibia missing, genitalia extracted, disarticulated and glued to the same mounting card as the specimen, ‘49201 [printed] // Hist. -Coll. ( Coleoptera ) / Nr. 49201 / Saprinus granarius Er. x / Austria, Dahl / Zool. Mus. Berlin [black-framed, printed label] // SYNTYPE / Saprinus granarius / Erichson, 1834 / labelled by MFNB 2016 [red label, printed] // granarius Er. / Austr. Dahl [black-framed, written label]’ ( MFNB).Another♁, without syntype status, identified as ‘ Saprinus granarius’ and labelled as ‘Carthagena [pink label, written] // Saprinus / granarius / Er. [written] // Hist. -Coll. ( Coleoptera ) / Nr. 49201 / Saprinus granarius Er. x / Austria, Dahl / Zool. Mus. Berlin [black-framed, printed label] ( MFNB). According to the MFNB staff, the lectotype designation of this species was not allowed, hence no lectotype was designated.

Saprinus arenarius Marseul, 1855 . LECTOTYPE (present designation): ♁ ( Fig. 27 ), originally pinned, with a pin-hole in its right elytron, glued onto a rectangular mounting card, right metatarsomere missing, genitalia extracted, disarticulated and glued onto the same mounting card as the specimen, ‘132 / Saprinus / arenarius / Dej / Aust. / Dej. [round, written label] // Saprinus 132 / arenarius / Dej. / Aust. [yellow, written label] // TYPE [red-printed label] // MUSEUM PARIS / Coll. / DE MARSEUL 1890 [printed] // Saprinus arenarius / Marseul, 1855 / LECTOTYPE / Des. T. Lackner 2017 [red label, printed]’ ( MNHN). This species was described based on unknown number of specimens and existence of other material cannot be excluded therefore we designate the lectotype to fix its taxonomic identity.

Saprinus corsicus Marseul, 1855 . SYNTYPE?: ♁ ( Fig. 28 ), glued to a rectangular mounting card, genitalia extracted, disarticulated and glued onto the same mounting card as the specimen, ‘[small, round golden label, which could be an indication that this was indeed Marseul’s type specimen] // Corse [printed] // type / Marseul [written] // Saprinus / granarius [written] // Saprinus / rubripes / v. corsicus, Mars. [written] // Saprinus corsicus / Marseul, 1855 / SYNTYPE? / Des. T. Lackner 2017 [red label, printed]’ ( MNHN; coll. Thérond).

Saprinus rubripes var. clermonti Auzat, 1920 : HOLOTYPE: ♁ ( Fig. 29 ), right antennal funicle, two left metatarsomeres missing, glued onto a rectangular mounting card, genitalia extracted, disarticulated and glued onto the same mounting card as the specimen, ‘Arcachon / Cap Ferret [written] // var./ Clermonti / Type / Dr.Auzat det. 1920 [printed-written] // Coll./ Dr.Auzat [light-green label, written] // TYPE [red label, printed] // Saprinus rubripes var. / clermonti Auzat, 1920 / HOLOTYPE / Des. t. Lackner 2017 [red label, printed]’ ( MNHN). This taxon was described based on a single specimen, which is therefore holotype by monotypy.

Additional material examined. ALGERIA: BÉCHAR REGION: Bèni Abbès, Sahara, 20.x.1980, 1 ♁, A. Olexa lgt. ( CTLA). ORAN REGION: Oran,no further data,1♁ ( MMBC). AZERBAIJAN: LANKARAN REGION: Mamusta env., 12.v.2001, 5 spec., T. Lackner lgt. ( CTLA). BULGARIA: BLAGOEVGRAD REGION: Struma River valley, Sandanski, 21.–22.iv.1987, 1 spec., J. Mertlik lgt. ( CTLA). BURGAS REGION: Burgas, on the beach, 5.viii.1981, 1 ♀, collector unknown ( NHMW); Nessebar, 4.viii.1994, 2 spec., 22.viii.1996, 4 spec., T. Lackner lgt. ( CTLA); Arkutino, 14.–16. ix.1988, 1 spec., J. Růžička lgt. ( CTLA); Sozopol env., 1.iv.2014, 1 spec., P.Kylies lgt. ( CTLA); Primorsko, vii.1980, 1♀, J. Pokorný lgt.( MMBC). VARNA REGION: Škorpilovici, vii.1983, 1 ♀, J. Pokorný lgt. ( MMBC). FRANCE: Gallia, no further data, 1 ♀ ( MMBC). BOUCHES- DU- RHÔNE: St.Maries, Camargue, 9.x.1928, 1♀, L.Puel lgt. ( MNHN; coll. Thérond); Grau du Roi, 10.ii.1939, 1♁, 1♀, 19.vi.1933, 1♁, J.Thérond lgt.( MNHN; coll. Thérond). CORSE: Oletta, 13.viii.1981, 1 ♀, Wewalka lgt. ( NHMW); Bonifacio, Révélière, 1 ♀, Col. A. Grouvelle ( MNHN, coll. Thérond); Corse, 1 ♁, 2 ♀♀, coll. Croissandeau ( MNHN; coll. Thérond); Corse, 1 ♁, coll. Bonnaire ( MNHN; coll. Thérond). HÉRAULT: Sète, no further data, 1 ♁ ( MNHN; coll. Thérond). GREECE: CORFU: Acharawi west, 24.vi.2017, 1 spec., O. Majzlan lgt. ( CTLA). KAVAVALA: Thassos Island, SW Potos env., saline, 10.vii.2004, 1 spec., P. Bulirsch lgt., ( CTLA). THESSALY: Leptokaria, 30.vii.–15.viii.1993, 2 ♁♁, J. Háva lgt. ( CTLA). HUNGARY: BÁCS- KISKUN MEGYE: Kalocsa, no further data, 1 ♁, Speiser ( NMPC);Bócsa,in sand-hills, 17.vi.1956, 3 spec., Kaszab & Székessy lgt. ( HNHM); Kéleshalom, vi.1955, 7 spec., Dr. Lenci lgt. ( HNHM); Bócsa, 17.vi.1956, 10 spec., Dr. Lenci lgt. ( HNHM); Kiskunsági National Park, Fülöpháza sand-hills, 22.vi.1978, 9 spec. (pitfall trap baited with cheese), Ádám & Hámori lgt. ( HNHM); Kalocsa, 4 spec., Speiser lgt., coll. Speiser ( HNHM);Soltvadkert, 100 m,sandy pasture, 31.iii.1975, 1 spec. (from cow dung), O. Merkl lgt. ( HNHM). CSONGRÁD MEGYE: Nagyszéksós, vii.1922, 2 spec., Szabó-Patay ( HNHM). FEJÉR MEGYE: Budapest env., Martonvásár, no date, 1 spec., H.Diener coll.( HNHM). PEST MEGYE: Budapest, no further data, 1 ♁, Gammel lgt. ( MMBC); Budapest, 12.xii.1934, 1 spec., Kaszab lgt. ( HNHM); Budapest, no further data, 1 spec. ( HNHM); Budapest env., Újpest-Alag, no date, 2 spec., coll. H. Diener ( HNHM); Budapest, Pest, no further data, 3 spec., Gimmel lgt. ( HNHM); Budapest, no further data, 1 spec., Kuthy lgt. ( HNHM); Pest m., Pusztavacs, Strázsa hill, 100 m, Festucetum vaginatae danubiale, from plant debris, 29.vi.1990, 30 spec., L. Ádám lgt. ( HNHM); Táborfalva, 5.vi.2015, 47º3′39″N, 19º27′36″E, 1 spec. (car-netting on the driving course) ( HNHM); Nagykőrös, Csókás forest, 4.iv–9.v.2010, 1 spec. (on sand in the forest), 10.–27.viii.2010, 1 spec. (on sand in the forest), Tallósi lgt. ( HNHM);Táborfalva, 30.vi.2012, 1 spec. (car-netting on the driving course), O. Merkl lgt. ( HNHM); Budapest-Rákos, no date, 1 spec., H. Diener coll. ( HNHM); Óbuda, iv.1903, 1 spec., coll. H. Diener ( HNHM); Budapest, no further data, 1 spec., coll. Pillich ( HNHM). SOMOGY MEGYE: Zamárdi, Balatonszéplak shore, 17.vii.–10.viii.1951, 1 spec. (on sandy meadow), Kaszab lgt. ( HNHM); Siófok, no further data, 3 spec., Lichtneckert lgt. ( HNHM); Balatonlelle, no further data, 6 spec., coll. Peregi ( HNHM); Öszöd, no date, 1 spec., viii.1903, 1 spec., vii.1905, 1 spec., vii.1906, 2 spec., viii.1906, 2 spec., Ehmnann lgt., coll. Dr. R. Streda ( HNHM). INDIA: ANDRA PRADESH: 35 km SE of Rajahmundry, Kottipale, Godavari River bank, 23.–24.ii.1994, 4 spec., Z. Kejval lgt. ( CTLA). KERALA: 10 km E of Punalur,bank of Kallada River, 8º59′N, 77º01′E, 20.–21.i.1994, 1 spec., Z. Kejval lgt. ( CTLA); Shoranur, Ponnani River, 10º46′N, 76º16′E, 31.i.1994, 3 spec., Z. Kejval lgt.( CTLA). ORISSA: 30 km NE of Jaleswar, riverbank of the Balasor River, 13.ii.1999, 1 ♁, Z. Kejval lgt. ( CTLA); Kalasandhapur, N of Berhampur, on a river bank, 20.–21.ii.1994, 1 spec., Z. Kejval lgt. ( CTLA). ISRAEL: Arvat Sedom, 8.–29.iv.2014, 1 spec.,I. Renan lgt.,spring ( CTLA). ITALY: GORIZIA: Grado, 1 ♀, J. Matcha lgt. ( NMPC). SARDINIA: no further data, 1 spec., Gené ( MFNB). TUSCANY: Pisa-Calambrone, 9.iv.2014, 2 spec., P. Kylies ( CTLA). KAZAKHSTAN: AKTYUBINSK REGION: Khobda River, 25.v.2000, 3 spec.,collector unknown ( CTLA). MONTENEGRO: BUDVA: Budva, no further data,1♁ ( NMPC). MOROCCO: FAS- MEKNAS REGION: MoyenAtlas,AguelmameAzegza Lake,lake shore, 22.–26.vi.1998, 1 spec., T. Lackner lgt. ( CTLA). SPAIN: CATALONIA: Girona, Sant Pere Pescador, 42°10.628′N, 3°06.608′E, 24.iv.2010, 1 ♁, J. Krátký lgt. ( CTLA). TUNISIA: JENDOUBA REGION: Chemtou env., Mejerda River, 30.iv.–1.v.1997, 2 spec., J. Mertlik lgt. ( CTLA). TURKEY: SAMSUN REGION: Samsun, 17 km N of Çarsamba beach, 19.v.1989, 4 spec., P. Kanaar lgt. ( CTLA). UKRAINE: CHERKASSY REGION: Dniper River, 10.–25.vii.2000, 1 ♀, Vasko lgt. ( NHMW). CRIMEA: Zurzut, 28.v.1999, 2 ♁♁, Putchkov lgt. ( NHMW). KHERSON REGION: Golopristansky district, Bolshevik, 1.vi.2000, 1♀, Putchkov lgt. ( NHMW). UNITED ARAB EMIRATES: ABU DHABI: Abu Dhabi,Channel Street,Al Raha Beach, 24°26′07.62″N, 54°33′42.26″E, 22.–31.iii.2015, 1 ♁, A. Pütz lgt. ( CTLA).

Diagnostic description. This taxon was redescribed and figured in detail by LACKNER (2010) and the reader is referred for the detailed redescription there. For the sake of better recognition we provide here only a short diagnostic description supplemented by a habitus image ( Fig. 25 ) and genitalia drawings ( Figs 33–39 ). Clypeus with elevated anterior margin, somewhat margined laterally, rugulose- lacunose; frontal stria well impressed, straight, carinate; continued as a well-impressed carinate supraorbital stria; frontal disc normally with irregular longitudinal rugae intermingled with sparse microscopic punctation; eyes flat, inconspicuous from above. Pronotal disc laterally with coarse punctation, between it and pronotal margin present a smooth longitudinal band; medially punctation much finer and sparser. First dorsal elytral stria the longest, usually reaching approximately three-fourths of elytral length apically; second, third and fourth dorsal elytral striae about the same length, reaching approximately elytral half apically; fourth dorsal elytral stria basally connected with sutural elytral stria; sutural stria well impressed, in shallow punctures, shortened on its apical tenth. Elytral disc on apical half (except for elytral flanks) with coarse and dense punctation, punctures separated by about their own diameter, anteriorly reaching about half of elytral length; basal half with only fine microscopic punctation; extreme apex of elytra with an impunctate band. Protibia ( Fig. 31 ) flattened and somewhat dilated, outer margin with four low teeth topped with short denticle followed by three minuscule denticles. Male genitalia. Sternite VIII ( Figs 33–34 ) longitudinally separated medially, apically with tiny inflatable membrane (velum); fringed with single short seta; tergite VIII and sternite VIII not fused laterally ( Fig. 35 ). Morphology of tergite IX ( Figs 38–39 ) typical for the subfamily; spiculum gastrale ( Fig. 37 ) expanded on both ends. Basal piece of aedeagus ( Figs 36–37 ) rather short, ratio of its length to length of parameres equals to 1: 3; parameres fused along their basal two-thirds; aedeagus curved ventrad ( Fig. 37 ).

Distribution. Hypocaccus (Nessus) rubripes , as presently understood, covers a large area from Portugal in the west to the Russian Far East in the east. It is spread in the entire Mediterranean subregion, the Netherlands, Central, Eastern and Western Europe, Turkey, Georgia, Armenia and Azerbaijan, Iran, Middle Asia, Mongolia, India as well as entire tropical Africa ( MAZUR 2011).

Biology. This species is most often found in sandy soils, often on riverbanks and seashores where it can be encountered on dung, carcass or under decaying vegetation.

Remarks. As noted already by REICHARDT (1932) and LACKNER (2010) this species exhibits a large degree of variation regarding its colouration of the cuticle and other external morphological characters, and might represent a complex of cryptic species. In this study, we try to sum up and depict ( Figs 25–29 ) the most common variations (‘forms’ – most of them originally described as species) of H. (N.) rubripes as elaborated already by REICHARDT (1932).

Hypocaccus (N.) arenarius Marseul, 1855 is a darker form ( Fig. 27 ) without metallic hue, whose colour can be attributed to the specimens’ age and has no taxonomic meaning.

Hypocaccus (N.) clermonti Auzat, 1920 is another form ( Fig. 29 ), which represents specimens worn-out by age; its frontal disc is almost glabrous (missing the numerous elongate rugae of the typical form) and the fore tibiae are devoid of teeth and denticles (worn off by age). This form has likewise no taxonomic meaning.

Hypocaccus (N.) corsicus Marseul, 1855 occurring in Corsica, Sardinia and Sicily is a viable candidate for a subspecies ( Fig. 28 ) and in fact has been treated as such (even as a bona fide species by several authors (e.g. SAINT- CLAIRE- DEVILLE 1907)). In this form, the sutural elytral stria is missing, the elytral punctation is much finer and its aedeagus is somewhat shorter and less dilated. In Corsica, this form occurs together with the typical form.

Hypocaccus (N.) granarius Erichson, 1834 occurs chiefly in southern Russia, in the Caucasian republics ( Armenia, Azerbaijan, Georgia) and is characterized by anteriorly effaced sutural elytral stria, occasionally not connected with the fourth dorsal elytral stria ( Fig. 26 ). REICHARDT (1932: 129, figs 16: A, B), when elaborating on different ‘forms’ and variation of this species put the most emphasis on the different pronotal shapes between (geographically) different populations. The two extremes of the pronotum are represented by acute ( REICHARDT 1932: 129, fig. 16:A) vs. obtuse ( REICHARDT 1932: 129, fig. 16:B) anterior pronotal angles. In case of acute anterior pronotal angles the marginal pronotal stria is well marked and easily discernible, whereas in case of obtuse anterior pronotal angles the marginal pronotal stria is rather difficult to distinguish. REICHARDT (1932) noted that the form with ‘acute anterior pronotal angles’ is much more frequent and almost all specimens from southern Russia, Crimea and Caucasian republics belong to it, as well as most specimens identified as the ‘ granarius ’ form. On the other hand, most specimens from North Africa belong to the form with obtuse anterior pronotal angles. Although there is no clear-cut difference between these two forms, specimens of both extremes are very different. A thorough morphological as well as molecular study of the possible ‘superspecies’ H. (N.) rubripes could possibly resolve this conundrum. In fact, the small handwritten label ( Fig. 30 ) found in Thérond’s collection (housed in MNHN) summarizes the situation with different ‘forms’ of this species up nicely. It is believed that Jean Thérond wrote up this label (Y. Gomy pers. comm. 2017).

Saprinus corsicus Marseul, 1855 . The examined specimen does not bear the labels typical for Marseul’s types, but it bears the tiny round golden label, which could indicate that it actually is the original Marseul’s type specimen. The description of MARSEUL (1855: 688) generally agrees with this specimen, because of the uncertainty of the exact status of the specimen we refrain from designating it as the lectotype. In the general collection of the Histeridae housed at MNHN (which contains Marseul’s collection) the type specimen(s) of Saprinus corsicus is missing. It is possible that this specimen was part of Auzat’s collection, which was later purchased by Thérond (N. Dégallier, pers. comm., 2017).

After the publication of LACKNER (2010) this species (together with the rest of the taxa included in the subgenus Nessus Reichardt, 1932 ) was transferred from the genus Hypocacculus Bickhardt, 1914 into genus Hypocaccus C. Thomson, 1867 by MAZUR (2011) without explanation.

Figures

Figs 25–28. Hypocaccus (Nessus) rubripes, dorsal habitus. 25 – Saprinus rubripes Erichson, 1834, syntype. 26 – Saprinus granarius Erichson, 1834, syntype. 27 – Saprinus arenarius Marseul, 1855, lectotype. 28 – Saprinus corsicus Marseul, 1855,?syntype.

Figs 29–30. Hypocaccus (Nessus) rubripes (Erichson, 1834). 29 – Saprinus rubripes var. clermonti Auzat, 1920, holotype, habitus, dorsal view. 30 – varieties of Hypocaccus (Nessus) rubripes as summed up by Jean Thérond.

Figs 31–32. Protibiae, dorsal view. 31 – Hypocaccus (Nessus) rubripes (Erichson, 1834); 32 – H. (N.) curtus (Rosenhauer, 1847) (both re-drawn from VIENNA 1980).

Figs 33–39. Male genitalia of Hypocaccus (Nessus) rubripes (Erichson, 1834): 32 – VIII sternite and tergite, ventral view; 33 – ditto, dorsal view; 34 – ditto, lateral view; 35 – aedeagus, dorsal view; 36 – ditto, lateral view; 37 – IX & X tergites, dorsal view + IX sternite (spiculum gastrale), ventral view; 38 – IX & X tergites + IX sternite (spiculum gastrale), lateral view.