Hypocaccus (Nessus) curtus ( Rosenhauer, 1847 ) Lackner & Seres, 2018

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 : 432-438

Hypocaccus (Nessus) curtus ( Rosenhauer, 1847) comb. nov.

( Figs 32 , 40–44 , 46–54 )

Saprinus curtus Rosenhauer, 1847: 26 (original description). MARSEUL (1855): 751 (redescription).

Saprinus (Hypocaccus) curtus: GANGLBAUER (1899) : 389 (redescription).

Hypocacculus (Nessus) curtus: REICHARDT (1932) : 49, 122 (keyed, redescription, incl. pl. IV, fig. 9); REICHARDT (1941): 285, 300 (keyed, redescription, incl. fig. 147C).

Saprinus puncticollis Küster, 1849: 30 (original description). MARSEUL (1855): 755 (redescription); BICKHARDT (1916): 96 (synonymy).

Saprinus (Hypocaccus) puncticollis: GANGLBAUER (1899) : 389 (redescription).

Hypocacculus (Nessus) puncticollis: KRYZHANOVSKIJ & REICHARDT (1976) : 204, 213 (keyed, redescription); VIENNA (1980): 179, 181 (keyed, redescription, incl. fig. 64b); MAZUR (1984): 89 (catalogue); MAZUR (1997): 254 (catalogue); YÉLAMOS (2002): 320 (keyed, redescription, incl. fig. 157f); MAZUR (2004): 94 (catalogue); MAZUR (2011): 209 (catalogue); LACKNER et al. (2015): 118 (catalogue).

Saprinus cribellaticollis Jacquelin du Val, 1858:99 (original description). Fauvel in GOZIS (1886): 202 (as synonym of Saprinus puncticollis ). MARSEUL (1862): 509 (redescription).

Saprinus (Hypocaccus) cribellaticollis: SCHMIDT (1885) : 312 (keyed).

Saprinus sicanus Marseul, 1862: 490 (original description, incl. pl. XVII, fig. 47). BAUDI DI SELVE (1864): 233 (as synonym of Saprinus puncticollis ).

Saprinus kuesteri Marseul, 1862: 715 (catalogue; unecessary replacement name for S. puncticollis Küster, 1849 ).

Saprinus revisus Marseul, 1876: 39 (original description). BICKHARDT (1916): 97 (as synonym of Saprinus curtus ).

Type material examined. Saprinus curtus Rosenhauer, 1847 . LECTOTYPE (present designation): ♁ ( Fig. 40 ), originally pinned with pin-hole in its right elytron, mounted on a rectangular mounting card, right antennal funicle and left mesotarsus missing, genitalia extracted and disarticulated, glued to the same mounting card as the specimen, ‘curtus / Rosenh. [written] // Hungaria [written] // herbeus Mars. [written] // Ex Musaeo / Rosenhauer [black-margined, printed label] // pas synonime / d’Herbeus Mars. / Dr. Auzat 1917 [written-printed] // Hongrie / Ex-Musaeo / ROSENHAUER [printed] // Hypocacculus / (Nannolepidius) curtus / ( Rosenhauer, 1847) / Dr. Auzat Dét. 1917 [printed] // Exemplaire provenant de la / collection Vauloger de Beaupré / Marcel (1862-1904) et inclus dans / la collection S. Risser en 2011 [black-margined, printed label] // Saprinus curtus / Rosenhauer, 1847 / LECTOTYPE / Des. T. Lackner 2017 [red label, printed]’ ( ZSM).

Saprinus puncticollis Küster, 1849 . LECTOTYPE (present designation): ♁ ( Fig. 42 ), glued onto a rectangular mounting card, two left and three right mesotarsomeres missing, genitalia extracted, disarticulated and glued to the same mounting card as the specimen, ‘Typ! [written] // Cagliari / Dr. Küster [written] // puncticollis / Küst. [written] // Saprinus / curtus Rosenh. [written] // Saprinus puncticollis / Küster, 1849 / LECTOTYPE / Des. T. Lackner 2017 [red label, printed]’ ( ZSM).

Saprinus cribellaticollis Jacquelin du Val, 1858 . LECTOTYPE (present designation): ♀ ( Fig.41 ), glued on a rectangular mounting card, both antennal funicles broken off; legs: except for right foreleg and left foretibia, all tibiae broken off; with the following labels:tiny, green rectangular label that is glued onto much larger translucent plastic mounting card (original mounting card of J. du Val) and tiny, red, quadrate label, followed by, ‘ Saprinus cribellaticollis / Jacquelin du Val, 1858 / LECTOTYPE / Des. T. Lackner 2017 [red label, printed]’ ( MNHN; coll. Jacquelin du Val).

Saprinus sicanus Marseul, 1862 . LECTOTYPE (present designation):♁ ( Fig. 43 ), glued onto a rectangular mounting card, right antennal funicle, both protarsi, two segments of right mesotarsus, as well as both metatibiae missing, male genitalia extracted, disarticulated and glued onto the same mounting card as the specimen, with the following labels: small, square-shaped blue label, followed by, ‘ Saprinus / sicanus m. / Schaum ‘59 [round label, written] // 129c / Saprinus / sicanus m. / Sicile / Schm 679 [round label, written] // 47 (129c) Saprin / sicanus m60 / Sicil. [written] // MUSEUM PARIS / Coll. De Marseul / 2842-90 [printed] // TYPE [red-printed label; followed by: “ Saprinus sicanus / Marseul, 1862 / LECTOTYPE / Des. T. Lackner 2017 [red label, printed]’ ( MNHN).

Saprinus revisus Marseul, 1876 . LECTOTYPE (present designation): ♀ ( Fig. 44 ), left antennal funicle, left protarsus, and left metatarsus missing, glued onto a rectangular mounting card, female genitalia extracted, glued to the same card as the specimen, ‘ Saprinus / revisus / rest of label illegible [round, blue label, written] // MUSEUM PARIS / Coll. / De Marseul 1890 [light-green label, printed] // TYPE [red-printed label] // Saprinus revisus / Marseul, 1876 / LECTOTYPE / Des. T. Lackner 2017 [red label, printed]’ ( MNHN).

Additional material examined. ALGERIA: ANNABA: Bône [= Annaba], 1 ♀, coll. Dr. Buysson ( MNHN; coll. Thérond); Bône [= Annaba], 1 ♁, Desbr. ( MFNB). EGYPT: Egypt, no further data, 1 ♀, coll.Ancey, ( MNHN; coll. Thérond). FRANCE: BOUCHES- DU- RHÔNE: Camargue, 2 ♁♁, L. Puel lgt., Auzat coll. ( MNHN; coll. Thérond); Camargue, Vaccares, no date, 1 ♁, 29.v.1937, 1 ♁, J. Thérond lgt. ( MNHN; coll. Thérond); Camargue, La Sauvage, 1.v.1928, 1 ♁, L. Puel lgt. ( MNHN; coll. Thérond); St. Maries de la Mer, 18.vii.1922, 1 ♀, Dr. A. Chobaut lgt., coll.Dr.Auzat ( MNHN; coll.Thérond). ITALY: SARDINIA: Cagliari, Saline di Stato, 10.v.1989, 1 ♁, 3 ♀♀, C. Meloni lgt. (1 ♁ in CTLA, 3 ♀♀ in MSNG); Stagno di Molentargius, 27.iii.1979, 1 ♁, C. Meloni lgt. ( CPVV), 29.v.1988, 1 ♁, 1 ♀, C. Meloni lgt. ( MSNG); Serdiana, 8.vi.2003, 6 ♁♁, 6 ♀♀, Fancello lgt. ( MSNG); Molentargius, 31.i.1979, 1 ♁, C. Meloni lgt. ( MSNG); Cagliari, Campo Santa Gilla, 28.iii.1983, 2 ♀♀, C. Meloni lgt.( MSNG). SICILY: Sicily, no further data, 1♁., 1 spec., Krtz. ( MNHN); Sicilia, no further data, 1 ♀ ( MFNB). LIBYA: TRIPOLI: Tripolis, no further data, 1 ♀ ( MFNB). SPAIN: ANDALUSIA: Andalusia, no further data, 1♀ ( MFNB). TUNISIA: TUNIS: Tunis, 1 spec., collector unknown, Reitter coll. ( ZSM); Tint, i.–ii.1882, 1 ♁, G. & L. Doria lgt. ( ZIN); Carthage, vii. 1914, 1♁, Novak lgt. ( ZIN); Tunis, no further data, iv.[18]83, 1 ♁ ( MFNB); Tunis, no further data, 6 ♁♁, 3 ♀♀ ( MFNB); Tunis, ii.–iii.1882, 1♁, G. & L. Doria lgt. ( MFNB); Radès, iv.1933, 1♁, M. Grossclaude ( MNHN; coll. Thérond). SOUSSE: Sebkha Kelbia lake near Sousse, 8.iv.1962, 1 ♁, Cl. Besuchet lgt. ( MSNG).

Redescription. PEL: 1.60–2.00 mm; APW: 0.75–1.00 mm; PPW: 1.40–1.60 mm; EW: 1.50–1.75 mm; EL: 1.00–1.40 mm. Body ( Fig. 40 ) oblong, oval, rather convex, cuticle dark-brown to black with faint to pronounced greenish hue; legs and antennal funicle light reddish-brown; antennal scape somewhat darker.

Head: mandibles densely punctate dorsally; clypeus densely and coarsely punctate, almost rugose-lacunose, anterior margin slightly elevated; frontal disc with similar, if somewhat weaker punctation; occasionally this punctation is confluent and forms tiny rugae; frontal stria slightly outwardly arcuate, complete to reduced to interrupted medially, supraorbital stria well developed; eyes flattened, but visible from above. Basal third of frontal disc with irregular rounded glabrous area; occipital stria weak, but visible. Antennal scape somewhat darker than reddish antennal funicle, antennae similar to other species of the subgenus, sensory structures of the antennal club studied by DE MARZO & VIENNA (1982).

Pronotum convex, lateral sides slightly narrowing anteriorly; anterior pronotal angles obtuse, marginal pronotal stria complete, its lateral portion observable in some cases from lateral view only. Entire pronotal disc covered with punctures separated by one to several times their diameter, punctation weakens medially. Scutellum very small, triangular.

Elytra: elytral epipleuron impunctate, marginal epipleural stria complete, marginal elytral stria well developed, complete, continued as apical elytral stria for short distance. Humeral elytral stria well developed, present on basal elytral third; internal subhumeral stria present as a median fragment. Dorsal elytral striae 1–4 well developed, first the longest, slightly bisinuate, reaching approximately two-thirds of elytral length apically, occasionally even slightly longer, striae 2–4 shorter, reaching approximately elytral mid-length apically, while second stria may be longer than striae 3–4; fourth stria usually the shortest, formed in most cases of beads of punctures, stopping short of elytral mid-length apically. Fourth dorsal elytral stria usually not connected (connected in specimens that belong to form ‘ cribellaticollis ’) with the basal end of (in)complete sutural elytral stria, which is in punctures and can be basally shortened. Elytral punctation covers approximately apical half of elytral disc, slightly surpassing elytral mid-length basally, slightly and scatteredly entering elytral intervals in some specimens; punctation rather dense, punctures separated by approximately their own diameter. Basal elytral fifth, fourth elytral interval, elytral flanks and extreme elytral apex impunctate, or with scattered microscopic punctation only.

Propygidium and pygidium: propygidium covered with punctation similar to that of elytra; pygidium with much finer and sparser punctation. Prosternum : prosternal process slightly to moderately concave (observed from lateral view); carinal prosternal striae carinate, divergent on prosternal apophysis, running convergent to sub-parallel to almost approximate apically; from mid-length of prosternal process slightly divergent anteriorly, apically united under tiny loop; interspaces between carinal prosternal striae with scattered punctures. Lateral prosternal stria strongly carinate, convergent apically, united in front of united carinal prosternal striae; lateral sides of prosternal process densely punctate; prosternal foveae moderately large, deep.

Mesoventrite: disc of mesoventrite approximately three times as wide as long, with scattered punctures (occasionally almost glabrous); marginal mesoventral stria complete, slightly inwardly arcuate medially; meso-metaventral stria undulate, bisinuate, in punctures, slightly distanced from meso-metaventral suture medially.

Metaventrite: disc of metaventrite apart from several rows of tiny punctures situated along basal margin entirely glabrous; lateral metaventral stria almost straight, slightly bisinuate, deeply impressed, in punctures, stopping short of metacoxa; lateral disc of metaventrite depressed, with large oval deep punctures separated by less than their diameter; metepisternum with similar punctation, punctures of smaller sizes than those of lateral disc of metaventrite. First visible abdominal ventrite striate laterally, with scattered fine punctation, occasionally almost impunctate.

Legs: protibia ( Fig. 32 ) on outer margin with 8–11 short to moderately long denticles diminishing in size proximally, protibial groove deep; rest of leg characters similar to preceding species.

Male genitalia: sternite VIII ( Figs 46–47 ) narrowing apically; sternite VIII and tergite VIII fused laterally ( Fig. 48 ). Tergite IX medio-laterally with tiny acute projection ( Figs 49–50 ). Spiculum gastrale ( Figs 51–52 ) similar to other congeners. Aedeagus ( Figs 53–54 ) almost subparallel, bluntly pointed apically.

Distribution. Hungary (?), France, Italy: Sardinia, Sicily, Spain, Portugal, Greece, Malta, Cyprus, Turkey, Tunisia, Algeria, Libya, Egypt.

Biology. According to VIENNA (1980), who repeats THÉROND (1975), H. (N.) curtus is found under detritus in sand near the seacoast, where it was collected from near Suaeda sp. and Statice virgata W. plant roots.

Remarks. The type specimen was part of Rosenhauer’s collection, which later became partly a part of R. Oberthür’s collection (A. Taghavian, pers. comm. 2017), currently housed in MNHN. The senior author has visited MNHN multiple times and failed to locate the type specimen(s) of this species in the collections of MNHN (including R. Oberthür’s collection). Mr. Serge Risser (Pleucadeuc, France) recently purchased the Histeridae collection of the late Marcel René Paul de Vauloger de Beaupré and published its contents in two separate papers ( RISSER 2013a,b). When reading RISSER’ S paper (2013a) we were intrigued by a specimen identified as Hypocacculus ( Nannolepidius !) curtus originating from Hungary and from ‘Musaeo Rosenhauer’. Mr. Risser was kind enough to send this specimen to one of us (T. L.). Having examined it as well as compared it to Rosenhauer’s original description we concluded that this is the long-lost type specimen of Rosenhauer’s species Saprinus curtus . This species was described based on an unspecified number of specimens and therefore we designate a lectotype to fix the species identity.

Saprinus curtus has become a mystery practically since its description, which was, however, rather detailed and served the purpose well. The reason for this was probably the fact that the type specimen(s) were unavailable for comparison and perhaps also because no more specimens matching this species were ever reported from ‘Hungary’. Based on the description alone, BICKHARDT (1916) correctly synonymized the H. (N.) puncticollis ( Küster, 1849) with H. (N.) curtus , which was also followed by REICHARDT (1932). MÜLLER (1937), however, doubted the two species are synonymous since the apical elytral stria in H. (N.) curtus reaches only mid-length of elytral apex, while, according to MÜLLER (1937) it is complete in H. (N.) puncticollis . Furthermore, MÜLLER (1937) advocated using Küster’s H. (N.) puncticollis as the valid (albeit not the earliest) name for this species and suggested, perhaps because of the incomplete description or the absence of the type material, that H. (N.) curtus was a dubious taxon. In the latest treatise on the Histeridae of the USSR (KRYZHANOVSKIJ & REICHARDT 1976), which in fact included almost the entire Palaearctic fauna, Kryzhanovskij upheld MÜLLER’ S (1937) opinion, and the name Hypocaccus (Nessus) puncticollis gained priority. This was followed by MAZUR (1984, 1997, 2011) in all three editions of his world catalogue of the Histeridae as well as by the latest edition of the Palaearctic Catalogue by LACKNER et al. (2015). Having examined both type specimens as well as numerous non-type specimens we can conclude that the two species are synonymous, and the earlier described taxon ( H. (N.) curtus ) has the priority. Regarding external morphological variation of this species, see Remarks section of H. (N.) curtus .

Saprinus puncticollis was described from a specimen found in Cagliari by Küster himself, as well as from specimen(s) brought by Mr. Handschuh from Cartagena ( Spain) ( KÜSTER 1849). The depository of the Spanish specimens is unknown and hence we designate the male specimen from Cagliari (Sardinia) as the lectotype to fix the identity of this taxon for purpose of synonymy.

Saprinus cribellaticollis was described based on unknown number of specimens.A single specimen was located in the original collection of Jacquelin du Val, deposited in MNHN, under the label ‘ Saprinus cribellaticollis ’. Jacquelin du Val did not provide his specimens with any labels, but, according to the curator of Coleoptera in MNHN, A. Taghavian, he kept his types in his private collection. Therefore we presume that this specimen, which completely matches J. du Val’s description, is a syntype. The species was described based on an unknown number of specimens and therefore we designate the lectotype to fix the taxon identity for purpose of synonymy.

Saprinus sicanus was described from Sicily ( Italy) based on an unspecified number of specimens, therefore we designate the lectotype to fix the taxon identity for purpose of synonymy.

Saprinus revisus was described from Algiers ( Algeria) based on an unknown number of specimens, therefore we designate the lectotype to fix the taxon identity for purpose of synonymy.

The type of S. curtus was found in mid-19 th century ‘Hungary’. This vague locality could refer to anywhere in the former Hungarian monarchy, which stretched south to the Adriatic Sea. It is possible that this species will be discovered in countries of the former Yugoslavia. It is a rather rare and seldom-collected species apparently spread around the Mediterranean Sea. Its rarity and slight morphological differences regarding dorsal punctation or course of carinal prosternal striae probably account for its numerous synonymies.

Hypocaccus (Nessus) controversus (G. Müller, 1937) ( Figs 45 , 55–63 )

Hypocacculus controversus G. Müller, 1937: 115 (original description).

Hypocacculus (Nessus) controversus: KRYZHANOVSKIJ & REICHARDT (1976) : 204, 212 (keyed, redescription); MAZUR (1984): 89 (catalogue); MAZUR (1997): 252 (catalogue); MAZUR (2004):93 (catalogue).

Hypocaccus (Nessus) controversus: MAZUR (2011) : 208 (catalogue); LACKNER et al. (2015): 117 (catalogue).

Type material examined. Hypocacculus controversus . LECTOTYPE (present designation): ♀ ( Fig. 45 ), mounted on a triangular mounting card, right metatarsus missing, ‘ ♀ [written] // Banat 1909 / Herkulesbad / leg. M. Hilf / Coll. O. Leonhardt [printed] // sbsp. / controversus [written] // TYPUS [light-ochre label, printed] // scat. / Hist. 6 [yellow label, written] // Hypocacculus / ( Nessus ) / controversus / G. Müller, 1937 / LECTOTYPE / des. T. Lackner 2017 [red label, written]’ ( CST) . PARALECTOTYPES: 1 ♀, side-mounted on a triangular mounting point, left meso- and metatarsus missing, ‘Athen / Phaleron [written] // Da Scat. / 6 [yellow label, written] // Hypocacculus (Nessus) / controversus Müll. / Det. T. Lackner 2017 [printed-written] // Hypocacculus / ( Nessus ) / controversus / G. Müller, 1937 / PARALECTOTYPE / des. T. Lackner 2017 [red label, written]’ ( CST). 1 ♀, ‘Saloniki / Schatzmayr [written] // Da Scat. / 6 [yellow label, written] // Hypocacculus (Nessus) / controversus Müll. / Det. T. Lackner 2017 [printed-written] // Hypocacculus / ( Nessus ) / controversus / G. Müller, 1937 / PARALECTOTYPE / des. T. Lackner 2017 [red label, written]’ ( CST).

Additional material examined. CYPRUS: Cyprus, no further data, 1 spec. (probably a male, genitalia lost), Baudi, ( MFNB). GREECE: Greece, 1 ♁ (genitalia lost, sexed by the protarsi), 1 ♀, Emge lgt., C. & O. Vogt coll. (1 ♀ in CTLA, 1 ♁ in MSNG); Greece, 1 ♁, ( MFNB). ATTICA: Attica, no further data, 2 ♀♀ ( MFNB). CRETE: Lerapetra E, 0–20 m, 17.–23.iv.2000, 1 ♀, A. Kopetz ltg. ( MSNG). IONIAN ISLANDS: Zante [=Zakynthos],Kalamaki,1909, 1♁, M.Hilf lgt., Coll. O. Leonhard ( MNFB). JORDAN: IRBID: 5 km NE of El Karama, 31.iii.1994, 31.58°N, 35.36°E, 200 m, 1 ♀, S. Bečvář jun. & sen. lgt. (dubious identification) ( MSNG); Toten Meer [= Dead Sea], 10.v.1963, 1 ♀, J. Klapperich lgt. (dubious identification) ( MSNG). ROMANIA: BANAT: Banat, Orșova, 1909, 1 ♀, M. Hilf lgt., coll. O. Leonhard ( MFNB). TUNISIA: DJERBA: Rass Taguernes, 10.–20.ii.1997, 1 ♀, Egger Manfred lgt. (dubious identification) ( MSNG). TURKEY: IZMIR: Smyrna? [=Izmir], no further data, 1 ♁ ( MFNB).

Diagnostic description. This species is externally rather similar to the preceding species and therefore here we provide only the diagnostic description outlining the differences between the two taxa. Body ( Fig. 45 ) somewhat more round and more flattened, light to dark brown, with light bronze hue (never with greenish hue). PEL: 2.00–2.30 mm; APW: 1.00–1.10 mm; PPW: 1.50–1.70 mm; EW: 1.65–1.90 mm; EL: 1.25–1.50 mm. Frontal disc more finely punctate than the one of H. (N.) curtus ; pronotum medially almost impunctate. The first dorsal elytral stria is only slightly longer than the second (apically both striae 1–2 surpass slightly elytral half), never reaching ¾ of the elytral length apically (in H. curtus the first dorsal elytral stria is substantially longer, occasionally surpassing ¾ of elytral length apically). Sutural elytral stria always connected basally with fourth dorsal elytral stria (in H. curtus these two striae are joined only in specimens that belong to the ‘ cribellaticollis ’ form), can occasionally be shortened apically. Carinal prosternal striae strongly convergent apically, their apices very approximate, stopping posterad of united lateral prosternal striae; their united apices not forming a ‘loop’ as in H. curtus . MÜLLER (1937) mentioned another character: the mesoventral punctation is supposed to be denser and coarser in ‘ controversus ’ than in ‘ puncticollis ’ (= H. curtus ). According to our observations, this is a valid, but not entirely stable character, since even among the few ‘ controversus ’ specimens we were able to examine we saw a specimen with only weak mesoventral punctation; the majority of specimens had their mesoventrite densely punctate. Male genitalia ( Figs 55–63 ) are generally similar to the preceding species, the aedeagi differ most markedly: the one of H. (N.) curtus is sub-parallel and blunted apically, while the one of H. (N.) controversus is shorter, stouter, slightly dilated in apical third with acutely pointed apex (compare Figs 53 and 62 ).

Note. The two female specimens from Jordan as well as the female from Tunisia are generally somewhat narrower, and their frons is adorned with coarse elongate rugae in place of dense punctures that are present at the type specimens. Therefore we identified these specimens with doubts as H. (N.) curtus .

Distribution. Romania, Greece (including Zakynthos Island and Crete), Montenegro, Spain, Morocco, Jordan (?), Turkey, and Saudi Arabia ( LACKNER et al. 2015). Newly reported from Cyprus and from Tunisia (with doubt).

Biology. According to KRYZHANOVSKIJ & REICHARDT (1976) this species is found on sandy banks of rivers and seas. The examined specimens did not bear any ecological data.

Remarks. This species was described from the following localities, but the number of specimens from each locality was not specified: Romania, Banat: Băile Herculeane; Greece: Thessaloniki; Phaleron near Athens; Parnass (= Mount Parnassus?), and Zakynthos Island: Kalamaki. We were able to examine the specimens from Romania, Thessaloniki, Zakynthos Island: Kalamaki and Phaleron near Athens. We chose the best-preserved female specimen from Romania as the lectotype, since it was the only specimen bearing a “type” label and the remaining three specimens as the paralectotypes, respectively. The remaining specimen(s) from Parnass (= Mount Parnassus?) should qualify as paralectotype (s), but their depository is unknown to us. The specimen from Zakynthos Island: Kalamaki was labelled by the MFNB staff as ‘ Hypocacculus rufipes Payk. ’, since it was placed among other specimens of H. (N.) rufipes originating from the collections of Schmidt and Bickhardt. Although MÜLLER (1937) mentioned that he examined specimen(s) from Zakynthos Island, we cannot be sure that this very specimen can be attributed a paralectotype for the following reasons: REICHARDT (1932: 124) already mentioned a series of five specimens with the same label data ( Greece, Zante [=Zakynthos] Island, Kalamaki, 1909, Hilf lgt., coll. Leonhard); at least one of these specimens was examined also by Müller. According to REICHARDT (1932), two of these were identified by Müller as H. (N.) puncticollis (= H. curtus ); two were deposited in Schmidt’s collection and identified as H. (N.) curtus var. aenescens Schmidt in litt.; and a single specimen was deposited in Schmidt’s collection and identified as H. (N.) rufipes Payk. The five specimens were supposedly divided between MFNB and Deutsches Entomologisches Institut, Müncheberg, Germany. According to Reichardt, who examined the whole lot, the five specimens doubtlessly belonged to the same species, albeit Reichardt was not sure to which, and placed them as ‘near to rufipes or transitional forms between rufipes and curtus , or even hybrids of the two species’. The specimen from Zakynthos we were able to examine is most likely the one that was identified as ‘ rufipes ’ in Schmidt’s collection, currently housed in MFNB and therefore probably not examined by Müller, when he described H. (N.) controversus . Albeit the specimen cannot be ascribed a paralectotype status this is the only male specimen of H. (N.) controversus we have seen and we therefore depict its genitalia here.

REICHARDT (1932) expressed his frustration with a couple of specimens of Hypocaccus (Nessus) curtus from North Africa, which look externally as Saprinus revisus but the aedeagus is different and similar to another species, Hypocaccus (Nessus) emendatus (Peyerimhoff, 1917) occurring in Algeria, Libya, Tunisia, and Egypt (LACKNER et. al. 2015). Without examination of the type of H. (N.) emendatus , we are unable to solve this riddle and opt for keeping the status quo.

Figures

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 40–43. Hypocaccus (Nessus) curtus (Rosenhauer, 1847), dorsal habitus.40 – Saprinus curtus Rosenhauer, 1847, lectotype.41 – Saprinus puncticollis Küster, 1849, lectotype. 42 – Saprinus cribellaticollis Jacquelin Du Val, 1858, lectotype. 43 – Saprinus sicanus Marseul, 1862, lectotype.

Figs 44–45.Hypocaccus (Nessus) curtus (Rosenhauer, 1847), dorsal habitus. 44 – Saprinus revisus Marseul, 1876, lectotype. 45 – Hypocacculus (Nessus) controversus G. Müller, 1937, lectotype.

Figs 46–54. Male genitalia of Hypocaccus (Nessus) curtus (Rosenhauer, 1847). 45 – VIII sternite and tergite, ventral view; 46 – ditto, dorsal view; 47 – ditto, lateral view; 48 – IX and X tergites, dorsal view; 49 – ditto, lateral view; 50 – IX sternite (spiculum gastrale), ventral view; 51 – ditto, lateral view; 52 – aedeagus, dorsal view; 53 – ditto, lateral view.

Figs 55–63. Male genitalia of Hypocaccus (Nessus) controversus (Müller, 1937). 55 – VIII sternite and tergite, ventral view; 56 – ditto, dorsal view; 57 – ditto, lateral view; 58 – IX and X tergites, dorsal view; 59 – ditto, lateral view; 60 – IX sternite (spiculum gastrale), ventral view; 61 – ditto, lateral view; 62 – aedeagus, dorsal view; 63 – ditto, lateral view.