Revision of the genus Exaesiopus Reichardt, 1926 (Coleoptera, Histeridae, Saprininae)

Abstract The genus Exaesiopus Reichardt, 1926 is revised herein. It now contains seven species; one new combination is proposed: Pachylopus glaucus = Exaesiopus glaucus (Bickhardt, 1914), comb. n., and one species is described as new: Exaesiopus therondi sp. n. from Afghanistan. Subspecies Exaesiopus grossipes berberus Peyerimhoff, 1936 is sunk in synonymy with Exaesiopus grossipes (Marseul, 1855), syn. n. Lectotypes and paralectotypes, respectively, for Saprinus grossipes Marseul, 1855, Exaesiopus grossipes berberus Peyerimhoff, 1936 and a neotype for Pachylopus glaucus Bickhardt, 1914 are designated. Exaesiopus grossipes is re-described; other species are provided with diagnostic descriptions and supplemented by SEM micrographs, colour images, and line drawings of their male genitalia. A key to species is given. Exaesiopus glaucus (Bickhardt, 1914) is newly recorded from the Republic of South Africa; Exaesiopus torvus Reichardt, 1926 is new to Uzbekistan and Russia; Exaesiopus atrovirens Reichardt, 1926 is new to Ukraine and Tajikistan; and Exaesiopus henoni (Schmidt, 1896) is new to Libya and Djibouti.

Differential diagnosis. Members of Exaesiopus are generally morphologically most similar to the Old World species of the genus Hypocaccus, differing from them chiefly by the setose pronotal hypomeron, strongly convex body, thickened metafemora and triangularly dilated and thickened metatibiae. In North America, however, there are at least two species of Hypocaccus (H. propensus (Casey, 1893) and H. servilis Casey, 1893) that are characterized by the presence of hypomeral setae.
Biology. Exaesiopus species are almost exclusively found in sandy soils, beach dunes, river sands, and are also found in sandy areas further inland (e.g. Sahara desert). Morphologically they are well adapted to their fossorial habits. Species are often collected on rotting biological matter, e.g. under faeces, dead fish etc., and are occasionally found under coastal wrack or by shore washing. The middle Asian E. atrovirens and E. torvus are sometimes found burrowing under Tamarix. The biology of E. laevis and E. therondi is unknown, the latter has been found inside the stomach of Kentish plover (Charadrius alexandrinus L. (Aves)).
Distribution. Genus Exaesiopus has a generally circum-Mediterranean-Caspian-Turanian distribution, most westerly occurring on the Canary Islands, reaching Afghanistan in the east. Its members have also been collected in the Sahara desert (Laghouat, Algeria), reaching as far east as northern Somalia (E. laevis) or Djibouti (E. henoni). Exaesiopus glaucus is known only from the Republic of South Africa and Namibia.
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.
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.
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)  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.
Differential diagnosis. Exaesiopus grossipes differs from the three species E. henoni, E. therondi and E. 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 E. atrovirens and E. glaucus it differs chiefly by the absence of a green metallic hue of the dorsum (compare Figs 1 with 73 & 98); from E. glaucus it differs furthermore by thickened and dilated metatibia (compare Figs 13 with 106). On the other hand, some specimens of E. grossipes (especially from N. Africa that formerly belonged to the subspecies berberus) can resemble the specimens of Middle-Asian E. torvus by their densely punctate dorsum. These specimens differ, however, from E. 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 E. grossipes, however (especially those from the northern shore of the Mediterranean Sea and South Europe) have distinctly less punctate dorsum than the specimens of E. 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.
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. Type locality. Aïn Sefra, Algeria.
Pronotal disc (Fig. 35) with ellipsoid to round, rather sparse punctation, punctures separated by their own to several times their diameters, postero-median part of disc always smooth, punctation stopping short of lateral pronotal margin leaving a narrow impunctate band; rest of the pronotum as in E. grossipes. Elytra generally as in E. grossipes; inner subhumeral stria shortly present medially; dorsal elytral striae for short distance surpassing elytral half; elytral punctation variable, in most specimens present only on fourth elytral interval, but can also at times be present on other elytral inter-  vals (a specimen from Libya), or almost completely missing (a specimen from Algeria); along elytral suture can reach almost elytral base, punctures irregular, variously deep, separated often by several times their own diameter, elytral flanks and humeri always smooth. Propygidium (Fig. 38) and pygidium similar to those of E. grossipes; punctation somewhat sparser (compare Figs 5 and 38). Prosternum (Fig. 39) generally similar to that of E. grossipes, but prosternal foveae very weakly impressed, often indiscernible (absent?); prosternal process deeply concave, constricted, prosternal structures and configuration of the two sets of prosternal striae similar to those of E. grossipes. Disc of mesoventrite (Fig. 40) almost smooth, similar to that of E. grossipes, but almost as long as wide; meso-metaventral sutural stria undulate; intercoxal disc of metaventrite with longitudinal depression in both sexes, more prominent in male, smooth, basally with several irregular rows of sparse punctures; lateral metaventral stria (Fig. 41) obliquely arcuate, apically almost reaching metacoxa; lateral disc of metaventrite (Fig. 41) and metepisternum generally similar to those of E. grossipes, but metepisternum with denser and coarser punctation and longer setae; meterpisternal stria unrecognizable beneath setae (absent?). Intercoxal disc of first abdominal sternite as in E. grossipes. Protibia (Figs 42-43) on outer margin with a single massive triangular tooth, followed by another lower tooth; both teeth topped by triangular denticle followed by two-three inconspicuous rounded denticles entombed in outer protibial margin; protibial spur inconspicuous (absent?); outer part of posterior surface of protibia (Fig. 43) smooth, separated from comparatively narrower median part by a definite ridge, posterior protibial stria complete, terminating in two minuscule inner posterior denticles; inner margin of protibia with double row of long dense lamellate setae. Mesotibia (Fig. 44) as in E. grossipes, but denticles on outer margin longer. Metatibia (Fig. 45) even more triangularly dilated and thickened than that of E. grossipes; outer margin with about four strong denticles larger in size apically; dilated anterior margin dorsally with several irregular rows of scattered tiny rounded denticles.
Elytral humeri not particularly enlarged; inner subhumeral stria present only as a row of several punctures; elytral punctation variable, in most specimens reaching elytral base along fourth elytral interval, punctures often present in all elytral intervals, elytral flanks impunctate; punctures regular and deep, separated by about half to several times their own diameter. Propygidium (Fig. 58) and pygidium as in E. grossipes, but covered with denser punctation. Prosternum: prosternal foveae (Fig. 59) weakly impressed; prosternal process otherwise similar to that of E. grossipes. Disc of mesoventrite ( Fig.  60) with scattered shallow punctures; intercoxal disc of metaventrite, lateral disc of metaventrite and metepisternum generally similar to those of E. henoni. Intercoxal disc of first abdominal sternite as in E. grossipes. Protibia (Fig. 61) more dilated than that of E. grossipes; on outer margin with two widely-spaced low teeth, topped by large triangular denticle followed by two low rounded denticles imbedded in outer protibial  margin; protibial spur inconspicuous (absent?) protibia otherwise similar to that of E. grossipes. Mesotibia (Fig. 62) generally similar to that of E. grossipes. Metatibia (Fig. 63) perhaps most triangularly dilated and thickened of all congeners; outer margin with approximately three widely-spaced tiny denticles; inner margin with a dense row of minuscule rounded denticles; no rows of denticles present between the two rows, surface rugulose-lacunose.
Differential diagnosis. Generally the most punctate species of Exaesiopus, which can be confused only with densely punctate specimens of E. grossipes from N Africa. It clearly differs from them by the punctation of pronotum as well as male genitalia (see also Key to species for details).

Figures 80-88. Exaesiopus atrovirens
Diagnostic description. Body length: PEL: 2.50-2.75 mm; APW: 1.00-1.10 mm; PPW: 2.00-2.25 mm; EW: 2.125-2.40 mm; EL: 1.50-1.875. Body shape (Fig. 73) as in its congeners, cuticle with greenish metallic tinge; legs, mouthparts and antennae reddish-brown. Antennae as those of E. grossipes; sensory structures of the antenna not examined. Mouthparts: mandibles somewhat more slender than those of E. grossipes; labrum with large antero-median depression, otherwise similar to that of E. grossipes; mentum and rest of the mouthparts likewise. Clypeus (Fig. 74) rectangular, rugose, anterior margin elevated, depressed medially; frontal, supraorbital and postorbital striae (Fig. 74) as in E. grossipes; frons with several irregularly shaped carinate transverse rugae intermingled with numerous tiny rugae; at times transverse rugae obliterated under numerous tiny rugae; eyes flattened, but visible from above. Pronotum as in E. grossipes. Elytra similar to that of E. grossipes; inner subhumeral stria present medially; elytral punctation, however, mostly confined to apical half of elytra, only rarely punctures present on other than fourth elytral interval. Punctation of propygidium (Fig. 75) and pygidium similar to those of E. grossipes, but punctures on propygidium almost confluent. Prosternum (Fig. 76) most similar to that of E. glaucus, foveae small but deep; prosternal process asetose. Mesoventrite (Fig. 77) occasionally sparsely and finely punctate, otherwise similar to that of E. glaucus; intercoxal disc of metaventrite similar to that of E. glaucus; longitudinal depression in female very faint; lateral metaventral stria, rest of lateral disc of metaventrite, metepisternum + fused metepimeron (Fig. 78) most similar to those of E. glaucus, but the amber setae distinctly longer and denser. Intercoxal disc of first abdominal sternite most similar to that of E. glaucus. Protibia (Fig. 79) similar to that of E. glaucus, differing from it chiefly by lower teeth topped by large triangular denticle. Mesotibia and metatibia similar to those of E. glaucus; metatibia, however, slightly more thickened and dilated.

Exaesiopus laevis Thérond, 1964
Diagnostic description. Body length: PEL: 2.375 mm; APW: 0.825 mm; PPW: 1.75 mm; EL: 1.50 mm; EW: 2.00 mm. Body (Fig. 89) without metallic tinge; legs, mouthparts and antennae light brown; antennal club amber. Antennae as in E. grossipes; sensory structures of the antennal club not examined. Mouthparts: mentum (Fig. 90) glabrous, sub-quadrate, shallowly inwardly arcuate on anterior margin; anterior margin with several rather long setae intermingled with short sparse ramose setae; rest of the mouthparts as in E. grossipes. Clypeus and frons (Fig. 91) similar to those of E. henoni. Pronotum almost smooth, only laterally and behind head with vague patches of shallow sparse punctation; otherwise similar to that of E. henoni. Elytra: inner subhumeral stria absent; elytral disc entirely smooth. Propygidium and pygidium (Fig. 92) similar to other congeners, but only sparsely punctate, punctures separated by several times their own diameter. Prosternum (Fig. 93): prosternal foveae tiny, almost invisible; prosternal process otherwise similar to that of other congeners. Mesoventrite (Fig.  94) glabrous, about as long as wide; metaventrite smooth; lateral disc of metaventrite and metepisternum similar to those of E. henoni. Intercoxal disc of first abdominal sternite similar to that of E. henoni. Protibia (Figs 95-96) similar to that of E. henoni, but outer margin of teeth topped by large triangular denticles, more similar in size Figure 89. Exaesiopus laevis Thérond, 1964 holotype, habitus. than those of E. henoni, furthermore outer part of posterior surface of protibia of E. laevis obscurely variolate, whereas it is glabrous in E. henoni. Mesotibia generally similar to that of E. henoni, but denticles on outer margin shorter. Metatibia (Fig.  97) likewise generally similar to that of E. henoni, but denticles on outer margin more numerous.
Male unavailable. Differential diagnosis. This species is most similar to E. henoni, from which it differs by almost impunctate pronotum (punctate in E. henoni), smooth elytra (punctate in E. henoni) and obscurely variolate posterior surface of protibia (glabrous in E. henoni). From the rest of Exaesiopus species it differs by the characters given in the Key to species (below).
Biology. Unknown, possibly similar to the congeners. Distribution. Known only from north-extreme tip of Somalia: Guardafoui.
Remarks. This species is morphologically rather similar to E. henoni, which is known also from the neighbouring Djibouti. The discovery of a male of E. laevis would help to elucidate the identities of the two respective species.  (Bickhardt, 1914), comb. n.
Diagnostic description. Body length: PEL: 2.50-2.60 mm; APW: 0.80-1.00 mm; PPW: 1.83-2.00 mm; EW: 2.00-2.18 mm; EL: 1.50-1.60 mm. Body (Fig.  98) similar to the species E. atrovirens, with feeble metallic tinge; legs, mouthparts and antennae light brown. Antennae as in E. grossipes. Mouthparts: as in E. grossipes; labrum with median keel-like elevation, surface anterad of it semi-circularly depressed; mentum (Fig. 99) sub-trapezoid, anterior margin without median notch, fringed with several long setae, lateral margins with single row of sparse shorter ramose setae; stipes with four setae; other mouthparts similar to those of E. atrovirens. Clypeus (Fig. 100) rectangular, obscurely variolate, anterior margin elevated, formed by two transverse tubercles that can occasionally be connected forming thus a ridge-like structure; clypeus and frons otherwise similar to those of E. atrovirens, but without numerous irregular rugae. Pronotum: sides slightly convergent on basal 3/4, strongly convergent on apical ¼; disc with round dense punctation, laterally punctures larger in size and increasingly ellipsoid, occasionally confluent; postero-median part of disc smooth, punctation stops short of lateral pronotal margin leaving a narrow impunctate band; pronotal base with a single row of round punctures; pronotal hypomeron with short amber setae almost Figure 98. Exaesiopus glaucus (Bickhardt, 1914) habitus. invisible from dorsal view; scutellum small, visible. Elytra: humeral elytral stria well impressed on basal fourth; inner subhumeral stria present medially as a short median fragment; elytral punctation confined to apical half of elytra, along elytral suture reaches up to 2/3 of elytral length anteriorly, punctures in most cases do not enter elytral intervals, regular and deep, separated by about their own diameter, punctation does not become denser apically; rest of elytra impunctate. Propygidium and pygidium (Fig.  101) similar to other congeners, with coarse and dense regular punctation. Prosternum (Fig. 102): prosternal foveae well impressed, rather small, but deep; prosternal process slightly concave, otherwise similar to that of other congeners. Mesoventrite (Fig. 103) slightly wider than long, almost smooth; meso-metaventral sutural stria well impressed, undulate; intercoxal disc of metaventrite with longitudinal depression in both sexes, more prominent in male, almost smooth, except for several rows of variously-sized   (Bickhardt, 1914)  deep punctures along base; lateral metaventral stria, lateral disc of metaventrite and metepisternum similar to those of E. henoni. Intercoxal disc of first abdominal sternite as with the rest of congeners. Protibia (Fig. 104) on outer margin with two moderately large triangular teeth, topped by rounded denticle followed by another two lower teeth topped by small round denticle and another tiny denticle entombed in outer protibial margin; setae of outer row regular and short; setae of median row shorter than those of outer row; anterior protibial stria almost complete; protibial groove deep; protibial spur ( Fig. 105) distinct but tiny, growing out from apical margin of protibia; outer part of posterior surface of protibia rugulose-lacunose, clearly separated from comparatively narrower glabrous median part; posterior protibial stria complete, terminating in two tiny inner posterior denticles; inner margin of protibia with single row of short lamellate setae. Mesotibia not particularly dilated or thickened, outer margin similar to that of E. henoni; posterior mesotibial stria fine, shortened apically; mesotibial spur stout, prominent and long; anterior surface of mesotibia smooth; anterior mesotibial stria shortened apically; claws of last tarsomere almost straight, their length approximately half the length of apical-most mesotarsomere. Metatibia (Fig. 106) slightly more dilated and thickened than mesotibia, but always more slender than that of the rest of the congeners; two rows of denticles on outer margin widely separated permitting for placement of another two denticles between the two rows; claws of apical-most metatarsomere shorter than half its length; otherwise metatibia similar to mesotibia. Male genitalia. Eighth sternite (Figs 107-108) entirely fused medially, apically with a setose velum; apex of eighth sternite with short dense setae. Eighth tergite apically weakly inwardly arcuate; eighth sternite and tergite fused laterally (Fig. 109). Ninth tergite (Fig. 110) on apical margin faintly inwardly arcuate; tenth tergite on apical margin regularly rounded, weakly inwardly arcuate basally. Spiculum gastrale  with typical 'head ' and 'tail'; tube-like, sub-parallel, slightly widening apically; parameres fused along their basal three-fourths, apex of aedeagus with pores; basal piece short, ratio of its length : length of parameres approximately 1:5.
Differential diagnosis. E. glaucus is arguably the most distinctive species of the genus differing from all other members by only slightly dilated metatibia (strongly dilated in all other species, compare Fig. 106 with e.g. 97); present and observable protibial spur (very tiny or absent in the rest of species, compare Fig. 105 with e.g. 43). Furthermore, the setae of the pronotal hypomeron are rather short and invisible from dorsal view (in all other species they are protruding from underside of the pronotum and are observable from dorsal view).
Biology. Found on a beach by the technique of shore-washing as well as on a river bank on deposited debris.
Distribution. Described from Namibia; newly recorded from the Republic of South Africa.
Remarks. The placement of this species in Exaesiopus must be regarded as tentative, as it differs from the rest of the members chiefly by only slightly instead of strongly dilated metatibiae. Hypocaccus from the Old World, however, does not contain any species with ciliate pronotal hypomera, and keeping E. glaucus in Hypocaccus would make it heterogeneous. Note that it was already Reichardt (1926) who remarked that this species should be, based on its ciliate pronotal hypomeron, moved into the genus Exaesiopus. Thérond Type material examined. Holotype, ♂, side-mounted on a triangular point, right hind leg missing, genitalia glued to the same mounting point as the specimen, with the following labels: "N AFGHANISTAN: / Hamud-i-Sabari / 26.iii.1949 Danish / Central Asian Expedn." (written in black ink); followed by: "Pachylopus / sp. not in BM / J. Balfour-Browne det. / v. 1964" (written-printed); followed by: "St. No. / 7" (printed-written); followed by: "Brit. Mus. / 1964-302" (printed-written); followed by: "Ex stomach of / Charadinus a. / alexandrinus L." (written in black ink); followed by: "Ex-   Diagnostic description. Body length: PEL: 2.125 mm; APW: 0.875 mm; PPW: 1.825 mm; EW: 2.05 mm; EL: 1.55 mm. This species (Fig. 114) is externally very similar to E. henoni, differing from it chiefly by its densely punctate pronotum, which is furnished with two round glabrous patches amongst the punctation laterally. The structure of frons (Fig. 115) is also different; whereas E. henoni always possesses only two well-defined chevrons on a completely glabrous surface, E. therondi has its chevrons beset on all sides with irregular rugae. The punctation of propygidium and pygidium (Fig. 116) is similar to that of E. henoni (Fig. 38). The prosternal process (Fig.  117) of E. therondi is more setose than that of E. henoni; prosternal foveae are absent. Anterior face of profemora (Fig. 117) is covered with dense amber setae in E. therondi, whereas only several sparse short setae are present in E. henoni. Anterior face of protibia ( Fig. 117) is rugulose-lacunose in E. therondi while it is glabrous in E. henoni. Further differences are found on male genitalia: Eighth sternite (Figs 118-119) is more slender, setae on apex are shorter; eighth sternite and tergite apically more slender (seen from lateral view; compare Figs 48 and 122). The rest of the male genitalia is markedly similar between the two species.
Differential diagnosis. E. therondi most resembles the Saharan species E. henoni, differing from it by rugulose-lacunose anterior face of protibia (glabrous in E. henoni), and the different structure of the frons (E. henoni has its frons glabrous with two chevrons whereas E. therondi has the chevrons surrounded by tiny rugae).
Remarks. Although this newly described species does strongly resemble the Saharan species E. henoni, and it has furthermore been found in a stomach of a bird, it is unlikely that they are conspecific, given the vast geographic stretch between African Sahara and Afghanistan. If it had been consumed by a Kentish plover in Africa and discovered in its stomach in Afghanistan it would have probably passed through the digestive tract of the bird by the time the bird migrated from the Sahara Desert to Afghanistan and would be beneath recognition at best. Instead, given the perfect shape of the insect, I consider it highly probable that the bird consumed it in Afghanistan and thus this species is an element of the Afghan fauna.

Discussion
Exaesiopus is a taxon that is morphologically well adapted to the psammophilous and fossorial way of life by the thickened metafemora as well as dilated pro-and especially metatibiae. A setose underside of the body is common to most obligate psammophiles in Histeridae and serves as further adaptation to life in sand; setae possibly prevent tiny particles of sand entering the body cavities. Although morphologically united by at least one weak synapomorphy (ciliate pronotal hypomeron), which is possibly a parallelism shared by some Hypocaccus spp. from North America, the monophyly of the genus Exaesiopus is likely questionable. The taxonomical uncertainties between (mostly) littoral taxa Hypocaccus, Exaesiopus, Pachylopus, Neopachylopus, Eopachylopus, etc. lie chiefly in the morphological similarities resulting from ecological pressures causing multiple parallelisms and convergences of characters. A future phylogenetic analysis of all littoral Hypocaccus-like taxa should focus on characters in systems putatively independent of the environmental selection pressures; otherwise characters that are prone to homoplasies (e.g. setae, denticles, rugae, trichomes etc.) could continue to obscure true phylogenetic relationships. In the recently published phylogeny of the subfamily by the author (Lackner 2014d), which included mostly the type species of the Saprininae genera, the type species of Exaesiopus was recovered among the members of a large clade of mostly psammophilous taxa whose interrelationships are unresolved.
Members of Exaesiopus are found in sandy soils or in sand over a vast geographic area rivalling perhaps only the distribution of Xenonychus Wollaston, 1864 (see also Lackner 2012). The distribution of Exaesiopus covers the area from the Canary Islands, circum-Mediterranean, South Europe, Caucasus, Iraq, Somalia, Djibouti, as far east as Afghanistan. Identity of the Somali species E. laevis Thérond, 1964 is uncertain; the species is known from a single female only. Other related genera (sensu Lackner, 2014) e.g. the species-rich and widespread Hypocaccus or Hypocacculus, or monotypic and localized Eopachylopus, Reichardtia etc., are distributed along most of the world beaches, as well as inland sand-systems; their inter-relationships shall be the focus of future phylogenetic studies.