Exaesiopus grossipes (Marseul, 1855)

Lackner, Tomas, 2015, Revision of the genus Exaesiopus Reichardt, 1926 (Coleoptera, Histeridae, Saprininae), ZooKeys 479, pp. 65-108 : 68-75

Taxon classification Animalia Coleoptera Histeridae

Exaesiopus grossipes (Marseul, 1855) Figs 1, 2-9, 10-13, 14-16, 17-25, 26-34

Saprinus grossipes Marseul 1855: 718, t. XX, fig. 153; Schmidt (1885): 315.

Saprinus rugicollis Schmidt 1890: 19 (nomen nudum, given as synonym).

Pachylopus grossipes : Schmidt (1896): 296; G. Müller (1931): 102.

Hypocaccus grossipes : Ganglbauer (1899): 393.

Styphrus grossipes : Jakobson (1911): 651.

Exaesiopus grossipes Reichardt (1926): 16; Reichardt (1941): 329, 330, fig. 117; Peyerimhoff (1936): 227; Kryzhanovskij and Reichardt (1976): 232, Figs 455-458; Vienna (1980): 196, fig. 69; Mazur and Kaszab (1980): 61, Figs 31, 34 D, E, F; Mazur (1984): 101; Mazur (1997): 263; Yélamos (2002): 338, Figs 12E, 161G, 169, 170A; Mazur (2004): 92; Lackner (2010): 112, Figs 19, 54, 89, 118, 339-359; Mazur (2011): 210.

Exaesiopus grossipes berberus Peyerimhoff 1936: 227 - syn. n.

Type locality.

Spain, France: Bayeux, Marseille.

Type material examined.

Saprinus grossipes : Lectotype, present designation, sex undetermined, pinned, right mesotibia, left mesotarus, both hind legs missing, with the following labels: "153 / Saprinus / grossipes / m / Marseille / Barage?" (round illegible label, written); followed by: "MUSEUM PARIS / COLL. / DE MARSEUL 1890" (printed); followed by: “TYPE” (red-printed label); followed by: " Saprinus grossipes / Marseul, 1855 / LECTOTYPE 2014 / des. T. Lackner" (red label, written).

Exaesiopus grossipes berberus: Lectotype, ♀, side-mounted on a triangular point, final two metatarsomeres on right hind leg missing, with the following labels: “Laghouat” (written); followed by: "Coll. Hénon / T Théry” (written); followed by: " Saprinus / grossipes berberus / Peyerimhoff / TYPE" (written); followed by: " Exaesiopus / grossipes berberus / Peyerimhoff, 1936 / LECTOTYPE / des. T. Lackner 2014" (red label, written) (MNHN). Paralectotypes, 2 ♂♂, both mounted on a single pin on triangular mounting points with extracted genitalia, with the following label: “Bône” (written); followed by: "Coll. Hénon / T Théry” (written); followed by: " Saprinus grossipes / berberus Peyerimh / TYPE" (written); followed by: " Exaesiopus / grossipes berberus / Peyerimhoff, 1936 / PARALECTOTYPE / des. T. Lackner 2014" (red label, written) (MNHN).

Note.

Peyerimhoff (1936: 227) distinguished the subspecies Exaesiopus grossipes berberus from the nominotypical one based on elytral punctation that should cover almost the entire elytral disc basally and laterally up to the second dorsal elytral stria. Among the three specimens that he furnished with type labels, however, only the female from Laghouat (Algeria) exhibits these characteristics; the two males from Bône [=Annaba, Algeria] have their apical half (approximately) impunctate and the punctation of their elytral discs terminates in third elytral stria. Therefore, a female from Laghouat has been selected for the Lectotype. Peyerimhoff (1936: 227) himself did not specify which specimen(s) belonged to the type series; he listed several localities with his extremely brief diagnosis of the new subspecies. Both Laghouat and Bône [=Annaba] were among the listed localities.

Additional material examined.

Bulgaria: 1 ♀, Asenovgrad, vi.1963, A. Olexa leg. (TLAN); 1 spec., Nessebar, 30.v.1996, O. Majzlan leg. (TLAN); 1 spec., Kazanlak, vi. 1963, A. Olexa leg. (TLAN); 1 spec., Plovdiv, Rektořík leg. (TLAN); 1 ♂, Newrokop, 19.vi.1938, leg. Hlinikowski (TLAN); 1 spec., SW Bulgaria, 2 km N Gara Pirin, 11.-12.vi. 1983, leg. Hieke (NCB). BOSNIA-HERZEGOVINA: 1 ♀, Mostar, V. Zoufal leg. (ZMHUB). MACEDONIA: 2 specs., 5 km E of Velandovo, 31.v.1992, P. Zahradník leg. (TLAN). Slovakia: 1 ♂, 1 ♀ + 1 spec., Čenkov, 24.vi. 1987, V. Kubáň lgt. (TLAN); 1 spec., ibid, but, 29.v.1993, T. Růžička leg. (TLAN); 2 specs., Kameničná-Balvany 8174c, 13.vii.2000, O. Majzlan leg. (TLAN). Serbia: 1 spec., Veliko Gradište, 2.ix.1955, Stanćić leg. (TLAN); 1 spec., Vranje, 4.vi.1968, collector unknown (TLAN). ITALY: 1 ♀, Piemont, coll. Bickhardt (MNHN); 1 ♂, Torino, coll. Fea (MNHN); 1 ♂ +1 ♀, Lagnola (?), xi. 1910, Sekera leg. (TLAN); 1 spec., PO, Fiume, Piacenza, 2.vi.1963, leg. P. Ratti (NCB); 1 spec., Veneto, Caorle, v. 1999, Clereau leg. (CAS); 2 ♂♂ + 1 ♀, Ponferrada, Paganetti (ZMHUB). FRANCE: 1 ♂, Bouches-du-Rhône, Les Saintes-Maries-de-la-Mer, 18.iv.1978, P. Queney leg. (CYG); 1 ♀, Charente Maritime, île de Ré, viii.1978 (P. Queney leg. (CYG); 1 ♂+1 ♀, Gard, Le Grau-du-Roi, 31.viii.1947, J. Thérond leg. (CYG); 4 ♂♂+1 ♀, Gironde, Soulac, 12.iv.1890, E. Giraud leg. (CYG); 1 ♀, idem, but 6.vii.1975, G. Tempère leg. (CYG); 1 ♀, Manche, Portbail, beach, 22.vi.1955, H. Chevin leg. (CYG); 1 ♂, Pyrénées, (CYG); 1 ♂, Montelimar, 7.v.1912, Laboissere leg. (MNHN); 1 ♀, Plouharnel, 1878, no further data (MNHN); 5 specs., Bretagne, Nicolas, no further data (MNHN); 1 ex., Var, Nice, flooding, v. 1951 (MNHN); 3 specs., Grande Coté, Royau, v. 1918, Chobaut & R. Lebon (MNHN); 1 spec., Aveyron, Millau, 6.iv.1960, Fages lgt. (MNHN); 2 specs., Ile de Ré, coll. Bonnaire, no further data (MNHN); 3 specs., La Rochelle d’Orbigny, no further data (MNHN); 4 specs., Carcassonne, no further data (MNHN); 2 specs., Pluharnel, dept. de Morbihan, no further data (MNHN); 2 specs., St. Jean de Monts, P. Sirguey (MNHN); 1 spec., Agen, 30.v.1908, G. Nicolas (MNHN); 2 specs., Toulouse, Col. D. Grenier (MNHN); 1 spec., Grau du Roi, 11.iv.1955, J. Thérond leg. (MNHN); 2 specs., ibid, but 2.ii.1938, J. Thérond leg. (MNHN); 1 spec., ibid, but 10.ix.1948, J. Thérond leg. (MNHN); 1 spec., ibid, but 19.v.1970, J. Thérond leg. (MNHN); 1 spec., ibid, but 15.v.1951, J. Thérond leg. (MNHN); 1 spec., ibid, but 1.x.1949, J. Thérond leg. (MNHN); 2 specs., ibid, but 9.iii.1940, J. Thérond leg. (MNHN); 2 specs., Pont du Gard, 3.iii.1927, J. Thérond leg. (MNHN); 2 specs., ibid, but 29.v.1928, J. Thérond leg. (MNHN); 2 specs., ibid, but 22.ix.1931, J. Thérond leg. (MNHN); 1 spec., Camargue, St. Maries, 9.x.1928, L. Puel leg. (MNHN); 5 specs., Vendée, St. Jean de Monts, vi.1926, P. Sirguey leg. (NCB); 2 ♂♂ + 1 ♀, ibid, but MNHN; 1 spec., Morgat, Brittany, no further data (BMNH); 2 specs., France, no further data (BMNH); 1 spec., Erqny, Côtes du Nord, H.D. Preston leg. (BMNH); 1 spec., Provence, no further data (BMNH); 3 specs., St. Jean de Monts ( Vendée), P. Sirguey leg., 1926 (BMNH); 1 spec., Manche, Utah Beach, 6.vi.[19]64; 1 spec., Beziers, no further data (BMNH) 1 spec., ibid but ZMHUB; 1 spec., Grau du Roi, 29.iii.1943, J. Thérond leg. (ZIN); 1 spec., Lyon, in Rhône, no data or collector (ZIN). HUNGARY: 1 spec., Hungary, no further data (BMNH); 1 ♂, Jarabszállás, 30.v.1971, P. Polák leg. (TLAN); 1 ♂, Dunakeszi, no further data (ZMHUB). SPAIN: 1 spec., Valencia, no further data (MNHN). IRAQ: 1 spec., Mesopotamia, Millingen, no further data (BMNH); 1 ♀, Mesopotamia, no further data (ZMHUB). Russia: 1 spec., Volgogradskaya obl., Tsimlya, 27.vii.1894, collector unknown (ZIN). Ukraine: 1 spec., Stan. Luganskaya, Lugansk okr., 17.vi.1928, collector unknown (ZIN); 2 ♀♀, Khersonskaya oblast, Alyoshki, Dneprovskij uezd, 26.v.1926, D. Znojko leg.; 2 specs., ibid, but 19.v.1929, N. Kostenko leg.; 1 spec., Khersonskaya oblast, Burkutskie plavni [zapovednik], 17.v.1929, N. Kostenko leg.; 1 spec., ditto, but Kazach village, 7.vi.1928, N. Kostenko leg. (all exs. ZIN). Greece: 1 spec., Peloponese occid., Epitalion, Alfios River, nr. Pyrgos, 13.iv.1995, T. Kopecký leg. (TLAN); 1 ♀, Pirgos, 1.v.1971, leg. Wewalka leg. (TLAN); 1 spec., Peloponesus, Xylokatron, 22.v.1962, H. Pochon leg. (MNHN); 1 spec., Thessalia, no further data (ZMHUB). MOROCCO: 1 ♀, Tauorirt, 10.iii.1993, G. Chavanon leg. (CYG); 1 ♂, Morocco centr., Moyen Atlas, Aguelmame Azegza lake, 22.-26.vi.1998, T. Lackner leg. (TLAN); 1 spec., Ouarzazate prov., Oued Draa River valley, Agdz env., N 30.40.52 W 006.25.08, 29.iii.2011, in human faeces, A. Gusakov leg. (CAS); 1 ♂, Beni Ounif near Figuig, 11.v.1944, Barbier leg. (MNHN). TUNISIA: 1 ♂, Medjez el Bab, v.1935, R. Demoflys leg. (CYG); 1 ♂, Gabès, v. 1944, R. Demoflys leg. (CYG); 1 spec., Zarzia, 5.-11.v.1977, M.A. Hielkema leg. (NCB). 1 ♀, Tunis, i–ii.1882, G. & L. Doria leg. (ZMHUB); 1 spec., Hammamet mer., 25.iii.-4.iv.1992, A. Pütz leg.; 2 ♀♀, 1 ♂ & 1 spec., 6-11.vi.1982, Kairuan, A. Olexa leg. (TLAN). ALGERIA: 1 ♀, Bona [= Bône?], Desbr., no further data (ZMHUB); 2 ♂ + 1 ♀, Oued Sebaou near Tizi-Ouzou, 25.vi.1908, collector unknown (MNHN); 2 ♀♀, Aïn Sefra, v. 1936, collector unknown (MNHN); 1 ♂, ibid, but coll. Bonnaire (MNHN); 1 ♀, Bou-Ktoub, S of Oran, Déchoguat leg. (MNHN); 1 ♀, Biskra, v. 1885, L. Bleuse leg. (MNHN); 1 ♀, Bou-Saada, no further data (MNHN); 1 ♀, south of Oran, no further data (MNHN) 1 ♂ + 1 ♀, Colomb-Béchar, 1912, P. Germain leg. (MNHN).

Redescription.

Although this species has been recently re-described by the author ( Lackner 2010: 112-116), I prefer to repeat this re-description here for the reason that the following species are morphologically rather similar to Exaesiopus grossipes . Those species are consequently provided only with diagnostic descriptions outlining their respective differences.

Body length: PEL: 2.10-2.75 mm; APW: 0.825-1.00 mm; PPW: 1.625-2.25 mm; EL: 1.25-2.00 mm; EW: 1.875-2.50 mm.

Body (Fig. 1) oval, convex, cuticle light to dark brown, sometimes with feeble bronze or greenish metallic tinge; legs, mouthparts and antennae rufopiceous.

Antennal scape with few short setae; club (Fig. 2) round, entire surface with thick short yellow sensilla intermingled with sparse slightly longer setae; sensory structures of the antennal club (Fig. 14) in form of stipe-shaped vesicle situated under circular sensory area on internal distal margin of the ventral side of antennal club.

Mouthparts: mandibles (Fig. 15) stout, outer margin slightly curved; mandibular apex bluntly pointed; sub-apical tooth of left mandible large, almost perpendicular; labrum (Fig. 16) sparsely punctate, shallowly depressed medially, two labral pits present, two labral setae arising from each; epipharynx almost completely hidden under labral fold; terminal labial palpomere elongated, its width less than half its length; mentum (Fig. 3) square-shaped, with deep antero-median notch; anterior margin with few long setae, lateral margins with single row of sparse shorter ramose setae; cardo of maxilla with few short setae on lateral margin; stipes triangular, with three much longer setae; terminal maxillary palpomere somewhat thickened, its width less than half its length, about twice as long as penultimate.

Clypeus (Fig. 4) rectangular, almost smooth, can be slightly rugose, anterior margin elevated, clypeus depressed medially; frontal stria well impressed, almost straight (sometimes somewhat curved outwardly), carinate, continued as carinate supraorbital and postorbital striae; frons with two to several irregularly shaped carinate transverse rugae or chevrons; eyes flattened, inconspicuous from above.

Pronotal sides slightly convergent forwards; apical angles blunt; marginal stria complete; pronotal disc convex, with round dense punctation, forming transverse rugae laterally, postero-median part of disc usually smooth, at times entire disc punctate (punctation can also stop short of lateral pronotal margin); pronotal base with a double row of round dense punctures; pronotal hypomeron with amber setae; scutellum small, visible.

Elytral humeri slightly prominent, elytra broad, almost as broad as long at its widest point; elytral epipleura with microscopic punctures, almost smooth; marginal epipleural stria complete; marginal elytral stria deeply impressed, continued as well impressed apical elytral stria; regular row of round punctures present along elytral marginal stria. Humeral elytral stria weakly impressed on basal third, sometimes doubled; inner subhumeral stria present medially, deep and rather long, rarely joining marginal elytral stria; elytra with four dorsal punctate elytral striae 1-4, all striae approximately reaching elytral half apically (occasionally slightly surpassing it), fourth elytral stria basally connected with sutural elytral stria; sutural stria deeply punctured, apically joining apical elytral stria. Elytral punctation variable, often confined to apical half of elytra, along elytral suture reaching almost anterior third of elytral disc, punctures regular and deep, separated by about half to their own diameter, occasionally (often in specimens from North Africa) covering most part of elytral disc (elytral flanks and humeri almost always smooth).

Propygidium (Fig. 5) almost completely exposed, long, covered with coarse and dense regular punctation; punctation of pygidium (Fig. 5) sparser and finer, punctures separated by about 1-3 times their diameter.

Anterior margin of median portion of prosternum (Fig. 6) regularly rounded; prosternal foveae weakly to well impressed, small and often indiscernible under conventional binocular microscope; prosternal process slightly to deeply concave, dorsally impunctate, laterally substrigulate-punctate, few microscopic setae present; carinal prosternal striae divergent between procoxae, subparallel, vaguely united in front, at times obliterated on their anterior third; lateral prosternal striae well impressed, carinate, convergent anteriorly, united in front of apices of carinal prosternal striae.

Mesoventral disc (Fig. 7) somewhat convex, almost smooth, slightly wider than long; meso-metaventral sutural stria well impressed, with several accompanying punc tures; intercoxal disc of metaventrite with longitudinal depression in male, smooth, basally with irregular sparse shallow fine punctures; lateral metaventral stria (Fig. 8) well impressed, carinate, obliquely arcuate, apically almost reaching metacoxa; lateral disc of metaventrite concave, with shallow setiferous punctures of various sizes, separated by approximately their own diameter; metepisternum with even denser and coarser punctation and setae, on apical third + metepimeron punctation much finer and sparser; metepisternal stria deeply impressed, present on metepimeron and approximately apical third of metepisternum.

Intercoxal disc of first abdominal sternite almost completely striate laterally; disc almost smooth, with sparse punctures along apical margin; lateral portion of disc of all visible abdominal sternites with short setae.

Protibia (Fig. 9) on outer margin with two to three low teeth, topped with triangular to rounded (blunt, if worn) denticle followed by two inconspicuous rounded denticles; setae of outer row sparse, moderately long; setae of median row shorter than those of outer row, sparse; anterior protibial stria shortened apically; protibial groove shallow; protibial spur (Fig. 10) minuscule, growing out from apical margin of protibia; outer part of posterior surface of protibia (Fig. 10) obscurely variolate, vaguely separated from comparatively narrower median part, posterior protibial stria complete, terminating in two minute inner posterior denticles; inner margin of protibia with double row of short dense ramose setae.

Mesotibia (Fig. 11) moderately dilated and thickened, outer margin with two rows of sparse short denticles; setae of outer row well sclerotized, comparatively short; setae of median row shorter and sparser, covering most of posterior surface; posterior mesotibial stria vaguely impressed, shortened apically; mesotibial spur stout, prominent and long; anterior face of mesotibia (Fig. 12) smooth; anterior mesotibial stria shortened apically; claws of last tarsomere bent, shortened, shorter than half its length.

Metatibia (Fig. 13) triangularly dilated and thickened apically; outer margin with four widely-spaced short rounded denticles, a single row of tiny sparse rounded denticles present dorsally on thickened anterior face of metatibia; setae of intermedian row shorter and denser, cover almost the entire posterior face of metatibia; otherwise metatibia similar to mesotibia.

Male genitalia. Eighth sternite (Figs 17-18) on apical half longitudinally separated medially, with tiny asetose vela, eighth sternite and tergite fused laterally (Fig. 19). Ninth tergite (Figs 20-21) apically with faint emargination; basally deeply emarginated; tenth tergite (Fig. 20) apically outwardly arcuate, basally faintly inwardly arcuate. Spiculum gastrale (Figs 24-25) typical for the subfamily representing the most common type with ‘head’ and ‘tail’ (sensu Caterino & Tishechkin, 2013); ‘tail’ cordate, ‘head’ with two narrow, curved arms. Aedeagus (Figs 22-23) tube-like, slender, basal piece of aedeagus rather short, ratio of its length: length of parameres 1:3.5; parameres fused along their basal two-thirds, aedeagus slightly curved ventrad (Fig. 23).

Differential diagnosis.

Exaesiopus grossipes differs from the three species Exaesiopus henoni , Exaesiopus therondi and Exaesiopus laevis chiefly by the shape of its protibia, which is on its outer margin furnished with three low teeth topped by triangular or rounded denticles (Figs 9, 10), whereas the three other mentioned species have their protibia furnished with two large teeth topped by triangular denticles on outer margin (Figs 61, 79 & 114). From Exaesiopus atrovirens and Exaesiopus glaucus it differs chiefly by the absence of a green metallic hue of the dorsum (compare Figs 1 with 73 & 98); from Exaesiopus glaucus it differs furthermore by thickened and dilated metatibia (compare Figs 13 with 106). On the other hand, some specimens of Exaesiopus grossipes (especially from N. Africa that formerly belonged to the subspecies berberus) can resemble the specimens of Middle-Asian Exaesiopus torvus by their densely punctate dorsum. These specimens differ, however, from Exaesiopus torvus by their respective male genitalia (compare Figs 17-34 with 64-72) and the less punctate pronotal disc (see also Key to the species for details, below). Most specimens of Exaesiopus grossipes , however (especially those from the northern shore of the Mediterranean Sea and South Europe) have distinctly less punctate dorsum than the specimens of Exaesiopus torvus .

Biology.

This species is found on the beach under coastal wrack as well as further away from the waterfront, almost exclusively on sandy soil. Beetles can be found under rotting fish, excrements or buried under vegetation.

Distribution.

Known from the Canary Islands, Morocco, Algeria, Tunisia, Libya, Spain, France, Italy, Greece, Bosnia and Herzegovina, Macedonia, Bulgaria, Russia, Serbia, Slovenia, Ukraine, Slovakia, Hungary, Austria, Iraq.

Remarks.

A variable species, covering vast area from the Canary Islands in the west to Iraq in the east. Its external morphology as well as male genitalia exhibit a certain degree of variation (compare Figs 17-25 and 26-34), but I find it difficult to discern discrete states among the variation and prefer to lump all examined specimens under the same species.