Saprinus lepidulus Broun, 1881: 665.

Cuticle dark brown to black with faint metallic luster; pronotum almost glabrous, only with faint scattered lateral punctation; elytra punctate and striate; frontal stria weakened, occasionally absent; pronotal hypomeron, prosternum, disc of mesoventrite, lateral disc of metaventrite, metepisternum + fused metepimeron and lateral sides of all abdominal ventrites setose; pronotal depressions present; prosternal foveae absent; prosternal apophysis strongly constricted between procoxae, prosternal process thence strongly expanded; carinal prosternal striae present as vague rudiments on prosternal apophysis; lateral prosternal striae absent; meso- and metafemora thickened, with rows of setigerous punctures. Eighth sternite of the male genitalia fused medially, apices with a row of sparse setae. Eighth tergite densely covered in pores and pseudopores. Spiculum gastrale dilated on both ends. Aedeagus narrow, parameres fused on their basal two-thirds. The densely setose venter in combination with coarsely punctate elytra will readily distinguish this New Zealand endemic from all other Saprininae present in the country.

A psammophilous taxon, found usually in carcasses or under coastal wrack. Several specimens were collected from pitfall traps.

Australopachylopus is endemic to New Zealand and is found on both North and South Islands, but has not been recorded from the Chatham Islands so far (Fig. 752).

Generic epithet of this new genus has been created combining the Latin word for south ‘austral’ and generic name Pachylopus.

Six species are included in the genus Neopachylopus Reichardt, 1926 (Mazur 2011: 211). In the published phylogenetic analysis of Saprininae by the first author (Lackner 2014d) Neopachylopus lepidulus falls separately from the other two members of the genus (N. sulcifrons (Mannerheim, 1843) and N. kochi Thérond, 1963)) which were likewise included in the analysis in order to test the monophyly of the genus. All three taxa (N. lepidulus, N. sulcifrons and N. kochi) were recovered inside a large polytomy of global, mostly psammophilous species united by one ‘strong’ and three ‘weak’ synapomorphies. N. lepidulus, in fact, comes out as sister to the clade uniting another New Zealand endemic Reichardtia Wenzel and Reichardtiolus pavlovskii Kryzhanovskij, 1959; although this purported monophyletic group is not strongly supported (see more in the discussion of Reichardtia). It is interesting to note, however, that the male genitalia of both Reichardtia and Australopachylopus share several similarities, e.g. overall gestalt of the eighth sternite and tergite (including the numerous pores and pseudopores; compare Figs 17–19 and 256–258), a tuft of short setae situated on the apices of eighth sternite, and, very similarly shaped aedeagus (observed from the lateral view; compare Figs 25 and 262). The on-going molecular studies by the senior author will hopefully shed more light on the relationships between New Zealand Saprininae, since all higher taxa have been included. We place N. lepidulus into a new genus based on the setose underside and prosternal apophysis strongly constricted between procoxae, strongly expanded prosternal process; carinal prosternal striae present as vague rudiments on prosternal apophysis; and absent lateral prosternal striae. All of these characters are different from the type species of Neopachylopus (N. sulcifrons).

New Zealand: Wellington: Lyall Bay.

Saprinus lepidulus Broun, 1881: Lectotype, present designation: unsexed specimen, with the following labels: “Type” (round, red-margined printed label); followed by: “New Zealand / Broun Coll. / Brit. Mus. / 1922-482” (printed); followed by: “Lyall Bay / Wellington” (written); followed by: “Pachylopus / lepidulus” (written); followed by: “Saprinus lepidulus / Broun, 1881 / LECTOTYPE 2014 / Des. Lackner & Leschen” (red label, written) (BMNH). Because the number of specimens in the original description is not given this specimen is selected as the lectotype. There are another two specimens in Broun’s collection, both of unknown sex: one bearing the following labels: “1169” (written); followed by “New Zealand / Broun Coll. / Brit. Mus. / 1922-482” (printed). Another one bears identical labels as the preceding specimen, but has one more label: “Invercargill” (printed) (BMNH). These specimens do not belong to the type series of Broun (1881) because Broun (1881) provides the type locality as Lyall Bay, and the second specimen lacks locality data.

NEW ZEALAND: North Island. ND: 4 specs., Ruakaka Beach, north Auckland, 22.ix.1932 (BMNH); 1 spec., Kai Iwi, 28.i.[19]15, Sand in Sun (NZAC); 2 specs., Himatangi, 9.ii.[19]58, R.A. Cumber leg. (NZAC); 1 spec., Kawerua Beach, 5.ii.1975, K. Hoson leg. (LUNZ). WO: 1 spec., Waikato Heads, under Muehlenbeckia, 1.i.1959, B.M. May leg. (NZAC); 1 spec., Kawhia Harbor, 27.i.1958, B.M. May leg., under dead fish on beach (NZAC); 1 ♂, 2 ♀♀, 4 specs., Raglan, 19.ix.1981, C.F. Butcher leg., under dead bird, fish (NZAC). HB: 1 spec., Porangahau, 29.i.1991, C. Duffy leg., pitfall traps (NZAC); SC: 1 spec., Temuka Beach, 22.iii.2008, on dead seagull (CJN). RI: 1 spec., Foxton Beach, 19.v.1996, in dead dry puffer fish on sandy beach (CJN). WN: 3 specs., Paekarariki Beach, 4.xii.1941, G.V. Hudson (BMNH); 1 spec., Wellington (BMNH); 2 specs., Wellington, J.J. Walker (BMNH). South Island. NN: 8 specs., Westport, J.J. Walker (BMNH); 1 spec., Westport, Beach Nelson, 15.x.1971, J.C. Watt leg., under logs (LUNZ); 1 spec., Farewell Spit, Outer Beach, 9.ii.1981, J.W. Early leg., carcass of dolphin (LUNZ); 1 ♂ & 4 specs., Wharariki Beach, 7.ii.1981, J.W. Early leg., under carcass on sand (LUNZ). MC: 2 specs., Lyttelton, xii.1901, J.J. Walker (BMNH). WD: 8 specs., Okarito lagoon, 7.–9.xii.2012, beach (under dead Larus), 43°13.5'S, 170°09.6'E, 1 m, M. Fikáček, J. Hájek & R. Leschen leg. (NMPC; 1 ex. in coll. TLAN); 1 ♀, Mananui Beach, 17.i.1982, J.W. Early leg., sandy beach (LUNZ). DN: 1 ♀, Kuri Bush, nr. Talieri Mouth, 18.iii.2001, on sandy beach (CJN). Unknown localities: 1 spec., New Zealand, 1162, Broun leg. (NZAC); 1 spec., New Zealand, without further data, J.J. Walker (BMNH); 1 spec., New Zealand, without further data, Broun, 5.ix.[18]89 (BMNH); 1 spec., New Zealand, no further data, vi.1904 (BMNH); 1 spec., New Zealand, no further data (BMNH).

Body length: PEL: 4.10–5.50 mm; APW: 1.10–1.75 mm; PPW: 3.15–4.00 mm; EL: 2.65–3.50 mm; EW: 3.50–4.50 mm. Body (Fig. 1) rectangular oval, dorsal surface convex, ventral surface flattened, cuticle black with slight metallic luster, legs, mouthparts and antennae dark brown, antennal club blackish.

Figure 1. 

Australopachylopus lepidulus (Broun, 1881) comb. n., habitus, dorsal view.

Antennal scape (Fig. 2) with carinate margins, with numerous long setae; antennal club (Fig. 3) round, slightly depressed dorso-ventrally; without visible articulation, entire surface with thick short yellow sensilla intermingled with sparse longer erect sensilla, apically with a flat sensory area with short dense sensilla; sensory structures of antennal club (Fig. 16) in a form of a single pear-shaped vesicle situated under sensory area on inner-ventral side of club.

Figures 2–9. 

2 Australopachylopus lepidulus (Broun, 1881) comb. n., head, dorsal view 3 antennal club, dorsal view 4 head, ventral view 5 mentum, ventral view 6 labrum + clypeus, dorsal view 7 propygidium + pygidium 8 prosternum 9 mesoventrite.

Mandibles stout, thickened (Fig. 14) with rounded outer margin strongly curved inwardly, dorsally with sparse punctures, acutely pointed, sub-apical tooth on inner margin of left mandible large, blunt; labrum (Figs 6, 15) punctate, dorsally convex with slight median depression; labral fold small, setae of lateral fringe moderately long; two labral setae present; terminal labial palpomere (Fig. 4) elongated, its width about one-third its length; palpal organ present on both labial and maxillary palpi; mentum (Fig. 5) square-shaped, anterior angles not produced, anterior margin with deep median excavation, surface around it with four longer setae; lateral margins with double row of shorter ramose setae; disc of mentum imbricate, with several short scattered setae; cardo of maxilla with few short setae on lateral margin, stipes triangular, with three short setae; lacinia without lacinial hook (=uncus); terminal maxillary palpomere elongated, its width about one-third its length, approximately twice as long as penultimate.

Figures 10–13. 

10 Australopachylopus lepidulus (Broun, 1881) comb. n., lateral disc of metaventrite + metepisternum 11 protibia, ventral view 12 mesotibia, ventral view 13 metatibia, ventral view.

Clypeus (Fig. 6) sub-quadrate, rounded laterally, slightly concave medially, with scattered microscopic punctures; frontal stria largely interrupted anteriorly (at times completely absent), continuous with weakly carinate supraorbital stria; frontal disc (Fig. 2) with irregular scattered fine punctation; eyes convex, visible from above.

Pronotal sides (Fig. 1) bisinuate, moderately convergent anteriorly, apical angles acute; pronotal depressions well-impressed (occasionally shallow); marginal pronotal stria complete, carinate; disc of pronotum laterally with a band of coarse and dense punctures (occasionally completely smooth), a band of similar punctation present also along pronotal base, pronotal disc otherwise glabrous; pronotal hypomeron with long yellow setae; scutellum small, visible.

Elytral humeri prominent; elytral epipleura smooth; marginal epipleural stria complete; marginal elytral stria straight and carinate, continued as well impressed complete apical elytral stria continuous with sutural elytral stria; along elytral marginal stria a regular row of round punctures present. Humeral elytral stria well impressed on basal fifth (occasionally longer); inner subhumeral stria present medially, rather long, almost attaining marginal elytral stria apically; elytral disc with four deeply impressed dorsal elytral striae 1–4, about the same length, not reaching elytral half apically (occasionally third and fourth dorsal elytral striae intermittent or obliterated by coarse punctation); fourth elytral stria basally not connected with sutural elytral stria; sutural elytral stria well impressed, abbreviated on basal tenth and apical sixth (occasionally continuous with apical elytral stria), interspace between it and elytral suture somewhat elevated, especially on apical third, on basal half a row of fine punctures present; punctation of elytral disc (with exception of elytral humeri and elytral flanks) coarse and dense, punctures separated by less than twice their diameter, on intervals between elytral striae punctation somewhat weakened.

Figures 14–16. 

14 Australopachylopus lepidulus (Broun, 1881) comb. n., mandibles, dorsal view 15 labrum: left half depicting dorsal view and right half depicting epipharynx 16 antennal club depicting sensory structures of the antenna.

Propygidium (Fig. 7) largely covered by elytra; its punctation similar to that of elytral disc; pygidium (Fig. 7) also densely and coarsely punctate, punctures becoming sparser towards extreme apex.

Anterior margin of median portion of prosternum (Fig. 8) slightly projected medially, setose; prosternal foveae absent; marginal prosternal stria vaguely impressed but complete, distanced from anterior margin; prosternal apophysis strongly constricted between procoxae, knife-like, prosternal process thence strongly widening and sloping down anteriorly, surface imbricate, dorso-medially with numerous setigerous punctures; occasionally vestiges of carinal prosternal striae present on prosternal apophysis; lateral prosternal striae absent.

Anterior margin of mesoventrite (Fig. 9) with slight median projection; discal marginal mesoventral stria anteriorly absent, laterally present; disc of mesoventrite convex, with deep and dense setigerous punctures of various sizes; meso-metaventral suture indistinct; meso-metaventral sutural stria absent; intercoxal disc of metaventrite smooth, in male with a longitudinal median depression; laterally with coarse round setigerous punctures; a narrow band of fine punctation present along posterior margin; lateral metaventral stria absent; lateral disc of metaventrite (Fig. 10) slightly excavate, with shallow dense setigerous punctures; metepisternum + fused metepimeron (Fig. 10) with even denser and coarser setigerous punctures, separated by less than half of their diameter; lateral metepisternal stria absent.

Intercoxal disc of first abdominal ventrite with shortened and vaguely impressed lateral stria; disc laterally and anteriorly with round deep punctures of various sizes, median part of disc impunctate, laterally punctures setigerous; all visible abdominal ventrites setose laterally.

Protibia (Fig. 11) slightly widening apically, outer margin with three large widely spaced distal teeth topped by large denticle, diminishing in size in proximal direction, followed by approximately six smaller stout proximal denticles, diminishing in size in proximal direction; setae of outer row thin, sparse; setae of median row approximate to inner protibial margin, but shorter than those of outer row; protarsal groove deep; anterior protibial stria vaguely impressed, shortened on posterior half; protibial spur large, hooked, growing out near tarsal insertion; outer part of posterior surface of protibia smooth, demarcation line between outer and median of posterior surface carinate; posterior protibial stria complete, with dense row of strongly sclerotized long setae growing in size and girth apically; inner margin with double row of short setae.

Mesotibia somewhat thickened, outer margin with a row of approximately 5 denticles growing in size apically, one more denticle present near apical margin; setae of outer row dense and long, strongly sclerotized, growing in size apically; setae of median row rather distanced from outer row, thinner and sparser; posterior mesotibial stria inconspicuous (absent?); anterior surface of mesotibia (Fig. 12) with another dense row of short denticles, becoming sparser and thinner towards inner mesotibial margin; anterior mesotibial stria undulate, with a row of regular short setae growing in size and girth apically; mesotibial spur large; apical margin of mesotibia with a row of about 5 stout denticles; first and second tarsomere ventrally with four long strongly sclerotized setae; third and fourth tarsomere with only two such setae; fifth tarsomere devoid of setae ventrally; dorsally first two tarsomeres with two strongly sclerotized setae; third, fourth and fifth tarsomere with only single seta dorsally; claws of apical tarsomere bent, approximately one-third its length; metatibia (Fig. 13) generally similar to mesotibia, but more slender; denticles of outer margin shorter and sparser; anterior surface of metatibia with three rows of short stout denticles.

Male genitalia. Eighth sternite (Figs 17–18) fused longitudinally; vela short, apices of eighth sternite with a tuft of short setae; eighth sternite laterally with several even shorter setae (Figs 17–19); eighth tergite covered with pores and pseudopores, apical margin inwardly arcuate, basal margin strongly inwardly arcuate; eighth tergite and eighth sternite fused laterally (Fig. 19). Ninth tergite (Figs 20–21) longitudinally fused medially, surface medially with two strongly sclerotized lines; tenth tergite basally inwardly arcuate, apically outwardly arcuate; spiculum gastrale (Figs 22–23) gradually dilated in most of apical half, apex strongly sclerotized; basal end dilated, rounded. Aedeagus (Figs 24–25) slightly thickened, almost parallel-sided; basal piece of aedeagus short, ratio of its length : length of parameres 1 : 5; parameres of aedeagus fused almost along their basal two-thirds; aedeagus only slightly curved from lateral view, apex of aedeagus flattened dorso-ventrally, fringed with a tuft of very short setae (Fig. 25).

Figures 17–25. 

17 Australopachylopus lepidulus (Broun, 1881) comb. n., male terminalia: 8^th sternite + 8^th tergite, ventral view 18 ditto, dorsal view 19 ditto, lateral view 20 male terminalia: 9^th + 10^th tergites, dorsal view 21 ditto, lateral view 22 male terminalia: spiculum gastrale, ventral view 23 ditto, lateral view 24 male terminalia: aedeagus, dorsal view 25 ditto, lateral view.

Diagnosis of this genus is based solely on the species C. aeneovirens that has been recorded from the Australopacific Region. Rather small, metallic ovoid beetle; elytra lighter than pronotum; frons with scattered fine punctures; frontal stria complete; pronotal depressions present, most of pronotal surface punctate, punctures most coarse in pronotal depressions. Dorsal elytral striae well developed, in punctures; prosternum with prosternal foveae, which are linked by marginal prosternal stria. Protibia with 7–8 moderately large teeth topped by denticle.

Figure 26. 

Chalcionellus aeneovirens (Schmidt, 1890) habitus, dorsal view.

The species C. aeneovirens is found in mammal dung and on vertebrate carcasses.

This genus is widespread in the Old World; in the Australopacific Region a single introduced species, Chalcionellus aeneovirens (native to the Afrotropical Region) is found in Western Australia, Victoria and Queensland (Fig. 752).

The sole species of Australian Chalcionellus, C. aeneovirens (Schmidt, 1890) differs from the rest of the Australian Saprininae by the presence of pronotal depressions in combination with well developed and deep prosternal foveae, connected by marginal prosternal stria. Superficially it could be confused with the similarly introduced species Hypocaccus (Nessus) interpunctatus interpunctatus (Schmidt, 1885), but the prosternal foveae of the former species are not connected by the marginal prosternal stria and, furthermore, H. (N.) interpunctatus interpunctatus possesses a characteristic ‘mirror’ (=polished area) on the second elytral interval, absent in Chalcionellus aeneovirens. From the externally similar species of the genus Hypocaccus, the species C. aeneovirens differs by the anteriorly connected prosternal foveae as well as by the absence of frontal rugae, typical for Hypocaccus.

Somalia.

Saprinus aeneovirens Schmidt, 1890: Lectotype, ♂, designated by Gomy & Vienna in 1999, glued on the tip of a triangular mounting card, left mesotibia broken off and glued to the same mounting card as the specimen, genitalia extracted and glued next to the mesotibia on the mounting card, with the following labels: “♂” (written); followed by: “Somaliland / Deyrolle 1.3.85” (written); followed by: “aeneovirens / Schm. Typ” (written); followed by: “coll. J. Schmidt” (printed); followed by: “coll. Schmidt - / Bickhardt” (printed); followed by: “Type” (brick-red label, printed); followed by: aeneovi- / rens * J. Schmidt” (light blue label, written); followed by: “aeneovirens / J. Schmidt / H. Bickhardt det. 1919” (printed-written); followed by: “LECTOTYPUS / Chalcionellus / aeneovirens (Schmidt) / Gomy et Vienna des., 1999” (red label, printed) (ZMHUB). Paralectotype, ♀, designated by Gomy & Vienna in 1999, right mesotarsus and left metatarsus broken off and glued onto the mounting card of the specimen together with the extracted female genitalia, with the following labels: “♀” (written); followed by: “Somali / land” (written); followed by: “Type” (brick-red label, printed); followed by: “coll. Schmidt - / Bickhardt” (printed); followed by: “PARALECTOTYPUS / Chalcionellus / aeneovirens (Schmidt) / Gomy et Vienna des., 1999” (red label, printed) (ZMHUB).

Figures 27–38. 

27 Chalcionellus aeneovirens (Schmidt, 1890) head, dorsal view 28 antennal club, dorsal view 29 mentum, ventral view 30 antennal club, ventral view showing sensory structures of the antenna 31 propygydium + pygidium 32 prosternum 33 mesoventrite 34 lateral disc of metaventrite + metepisternum 35 protibia, dorsal view 36 ditto, ventral view 37 mesotibia, dorsal view 38 metatibia, dorsal view.

AUSTRALIA. Western Australia: 1 spec., Dardanup, 1.ii.1979, G. Hall (howden trap) (ANIC); 1 spec., Gingin Brook, 6.x.1977, J. Ridsdill Smith (ANIC); 1 spec., Cataby, 12.vi.1979, G. Hall (howden trap) (ANIC); 4 specs., 15 km S Busselton, 11.i.1983, G.P. Hall (ANIC); 3 specs., ditto, but 12.i.1983 (ANIC); 1 spec., ditto, but 15.i.1983 (ANIC); 1 spec., ditto, but 27.i.1983 (ANIC); 5 specs., 5 km NE Dardanup, 11.i.1983, G.P. Hall (ANIC); 6 specs., ditto, but 13.i.1983 (ANIC; 2 exs. in coll. TLAN); 3 specs., ditto, but 12.i.1983 (ANIC); 10 specs., ditto, but 15.i.1983 (ANIC); 3 specs., ditto, but 27.x.1982 (ANIC); 2 specs., ditto, but 19.x.1982 (ANIC); 1 spec., ditto, but 21.x.1982 (ANIC); 1 spec., ditto, but 4.xi.1982 (ANIC).

Victoria: 2 ♂♂ & 1 ♀, Cheltenham, 2.–3.ii.1918, H. Pottinger (QM).

Queensland: 2 specs., Bangalee Beach, 10 m, 23°04'S, 150°46'E, 16.–19.xii.1999, D. & I. Cook (littoral R/F dung pitfall) (QM).

Figures 39–47. 

39 Chalcionellus aeneovirens (Schmidt, 1890) male terminalia: 8^th sternite + 8^th tergite, ventral view 40 ditto, dorsal view 41 ditto, lateral view 42 male terminalia: aedeagus, lateral view 43 male terminalia: 9^th + 10^th tergites, dorsal view 44 ditto, lateral view 45 male terminalia: spiculum gastrale, ventral view 46 ditto, lateral view 47 male terminalia: aedeagus, dorsal view.

Saprobiont, found often in dung or on carcass.

Described from Somalia, and widespread in East and South Africa; introduced into Australia (Queensland, Western Australia and Victoria; Fig. 753) (Mazur 2011).

The species is diagnosed above, as well as provided with a differential diagnosis that separates it from other Australian taxa. We chose not to fully re-describe it here, leaving its re-description to the revision of the genus Chalcionellus. For the sake of the better species recognition, however, we decided to depict it here, including its male terminalia.

This species-rich genus with 76 described species is distributed across North, Central and South America (Mazur 2011), with one species known also from the Palaearctic and one from Oriental Region, respectively. In the Australopacific Region we report only one introduced species, E. (Neosaprinus) rubriculus (Marseul, 1855) represented in collections by several old specimens presumably collected from New Zealand and Australia.

Diagnosis of this subgenus is based solely on the species E. (N.) rubriculus that has been recorded from the Australopacific Region. Frontal and supraorbital striae absent; eyes visible from above; cuticle dark brown to black; elytral striae 1-4 well developed; carinal prosternal striae divergent on apical half, thence running parallel, united anteriorly by deep sulcus; lateral prosternal striae shortened, attaining carinal prosternal striae at the point where these turn parallel; meso-metaventral stria absent; metepisternal stria complete.

Figure 48. 

Euspilotus (Neosaprinus) rubriculus (Marseul, 1855) habitus, dorsal view/

The biology of the members of the subgenus Neosaprinus is poorly documented, but species of this subgenus are usually found on the carcasses of vertebrates (Arriagada, pers. comm., 2014) or in caves (A.K. Tishechkin, unpublished). Gomy (2005) reports this species from the Island of La Réunion, collected inside a lava tube in the faeces of the Mascarene Swiftlet (Collocalia francica Gmelin, 1789) and concluded that its cavernicolous habitat is nothing exceptional. He (Gomy 2005) hypothesized that this species could have come from Brazil with a shipment of some kind of legumes, probably soybeans or corn. The labels of specimens of E. (N.) rubriculus recorded from Australia and/or New Zealand did not bear any information regarding their biology.

The subgenus Neosaprinus of the genus Euspilotus currently comprises nine described species, most of them occurring in the Neotropical Region (Mazur 2011). One species, E. (N.) scrupularis (J.L. LeConte, 1860) is distributed across North America, from British Columbia (Canada, with doubt), through Texas, Arizona, Utah, California, Washington, Oregon (with doubt) to Florida (USA), as well as in Mexico (Bousquet and Laplante 2006; Mazur 1997; Hatch 1962). One species, E. (N.) loebli Mazur, 1974 has been described from Malaysia and another one, E. (N.) perrisi (Marseul, 1872) occurs in the Palaearctic Region. In the Australopacific Region, we have examined three old specimens of E. (N.) rubriculus: two from Australia and one from New Zealand (Fig. 754).

This taxon is most likely to be confused with the genus Gnathoncus, with which it shares the absence of both frontal and supraorbital striae, but can easily be separated from it by single marginal epipleural stria (double in Gnathoncus) and absence of the hooked appendix between the fourth dorsal elytral and sutural elytral striae, characteristic for Gnathoncus. Furthermore, E. (N.) rubriculus has small prosternal foveae connected by a transverse sulcus whereas species of the genus Gnathoncus lack these structures. The examined specimens from Australopacific Region of this species are: a single female from Sydney (OUMNH), a single male from Brisbane (QM), both Australia and a single female from Tairua, New Zealand (NZAC). This New Zealand specimen was possibly traded or given originally to Thomas Broun who lived in Tairua. We treat this species in our study as a potential introduction because it has also been reported elsewhere outside of its native range (e.g. La Réunion; see Lackner 2013c).

America? (without further data).

Saprinus rubriculus Marseul, 1855: Lectotype, present designation: ♂, with genitalia placed in a small vial under the specimen, both protarsi, left mesotarsus, right metatarsus and left hind leg missing, pinned, with the following labels: tiny, rectangular pink label, followed by: “107 / Saprinus / rubriculus / m. / N.” (round, green label, written); followed by: “Horn / 11 mars 43 ?” (almost illegible label, written); followed by: “6a rubriculus / N. Am” (written); followed by: “MUSEUM PARIS / COLL. / DE MARSEUL 1890” (printed); followed by: “TYPE” (red-printed label); followed by: “Saprinus rubriculus / Marseul, 1855 / LECTOTYPE / des. T. Lackner, 2014” (red label, written) (MNHN). The type locality of this species is “America?” with a question mark. It is unclear why Mazur (1997, 2011) does not mention this doubt with the type locality. This species was designated from unknown number of specimens and the lectotype designation fixes the identity of the species.

Figures 49–57. 

49Euspilotus (Neosaprinus) rubriculus (Marseul, 1855) head, dorsal view 50 antennal club, dorsal view 51 prosternum 52 clypeus + labrum 53 mesoventrite 54 lateral disc of metaventrite + metepisternum 55 protibia, ventral view 56 mesotibia, ventral view 57 metatibia, ventral view.

Saprinus gnathoncoides Bickhardt, 1909: holotype, ♂, side-mounted on a triangular mounting card, with the following labels: “Montevideo / 28.xi.[19]08 / J. Tremoleras” (black-margined printed-written); followed by: “gnathoncoides / Bickh.” (written); followed by: “gnathoncoides / m / det. Bickhardt” (written-printed); followed by: “Type” (red label, printed); “Zool. Mus. / Berlin” (printed); followed by: “10-152” (yellow label, pencil-written, added by the senior author); followed by: “10-166” (yellow label, pencil-written, added by the senior author) (ZMHUB). This species was described from a single specimen (holotype by monotypy) examined here.

URUGUAY: 1 ♀, most likely collected with the type specimen, but without the type label: Montevideo, 28.xi.[19]08, J. Tremoleras (ZMHUB). ARGENTINA: 3 ♂♂, Cordoba prov., Balnearia, 17.–25.ii.2000, collector unknown (TLAN). La REUNION: 1 ♂, L’Eperon, Boyau de lave, Deject salanganes, 28.xi.2003, J. Pousserau leg. (CYG). AUSTRALIA: Queensland: 1 ♂, Brisbane, O.W. Tiegs (QM); 1 ♀ New South Wales: Sydney District, J.J.W, iii.1900 (OUMNH). NEW ZEALAND: North Island: CO: 1 ♀ Broun Tairua, Coll. Wakefield, Arthur Parrott Collection donated June 1983 (NZAC).

Figures 58–60. 

58Euspilotus (Neosaprinus) rubriculus (Marseul, 1855) mandibles, dorsal view 59 labrum: left half depicting dorsal view and right half depicting epipharynx 60 antennal club, ventral view showing sensory structures of the antenna.

Biology of this species is not well documented, but it has been found inside a lava tube on feces of the Mascarene Swiftlet (Lackner 2013c; see also above) where it has been introduced to La Reunion. Arriagada (1987) reports this species from the excrements of common vampire bat (Desmodus rotundus Geoffroy, 1810) in a cave in the Cordoba province of Argentina.

Described from “America?”, recorded from Argentina, Uruguay, Brazil, Venezuela as well as from islands of St. Helena and La Réunion; possibly adventive to New Zealand and Australia (Fig. 754).

Figures 61–67. 

61Euspilotus (Neosaprinus) rubriculus (Marseul, 1855) male terminalia: 8^th sternite + 8^th tergite, ventral view 62 ditto, dorsal view 63 ditto, lateral view 64 9^th + 10^th tergites, dorsal view; spiculum gastrale, ventral view 65 9^th + 10^th tergites + spiculum gastrale, lateral view 66 aedegus, dorsal view 67 ditto, lateral view.

The specimen from “Sydney District” (OUMNH) is slightly larger (PEL: 2.85 mm; APW: 1.05 mm; PPW: 2.00 mm; EL: 1.75 mm; EW: 2.125 mm) in size and has a short median fragment of the inner subhumeral stria, compared to the syntype specimen of S. gnathoncoides Bickhardt, 1909 (ZMHUB) that is slightly smaller (PEL: 2.25 mm; APW: 1.00 mm; PPW: 1.625 mm; EL: 1.25 mm; EW: 1.875 mm) and bears no trace of inner subhumeral stria. It is possible that E. (N.) rubriculus represents a complex of cryptic species though the male terminalia of the lectotype of S. gnathoncoides and those of the specimen from Brisbane are almost identical. The species is diagnosed above, as well as provided with a differential diagnosis that separates it from other Australopacific taxa. We chose not to fully re-describe it here, leaving its re-description to the revision of the genus Euspilotus (Arriagada, in prep.) For the sake of better species recognition, however, we decided to depict it here, including its male terminalia.

Cuticle brown to black, never metallic; frontal, supraorbital striae absent; pronotal hypomeron glabrous; pronotal depressions absent; elytra occasionally imbricate, punctures on apical part of elytra sometimes forming longitudinal rugae; marginal epipleural stria double; fourth dorsal elytral stria never connected with sutural stria; apical elytral stria usually shortened; elytra with characteristic hooked appendix between fourth dorsal and sutural striae at elytral base; anterior ends of fourth dorsal elytral and sutural elytral striae form a small hook. Prosternum without prosternal foveae; median fossa often present; carinal prosternal striae strongly convergent anteriorly, united under sharp angle; lateral prosternal striae shortened, strongly convergent anteriorly; prosternal process flattened, broad; outer-lateral costa reaches prosternal process, its basal margin distinctly elevated; metaventrite of males at times longitudinally concave; ninth tergite of male genitalia divided longitudinally.

Figure 68. 

Gnathoncus communis (Marseul, 1862) habitus, dorsal view.

Gnathoncus is predominantly composed of inquilinous species, present in the nests of birds or mammals; some species are found exclusively inside these nests where they are predators (Kryzhanovskij and Reichardt 1976), presumably preying upon larvae of fleas and other tiny arthropods. Some species, however, are occasionally collected on carrion. Both species of Gnathoncus occurring in the Australopacific Region are typical synanthropes and are often collected in pigsties, dovecotes or chicken coops.

Twenty-four species and subspecies are known to occur worldwide, most in the Holarctic Region (Mazur 2011); several species were possibly distributed over the globe by human activity. In Australopacific Region two introduced species have been collected in New Zealand and Australia (Fig. 754).

This genus can most easily be confused with the species of Tomogenius, endemic to the Australopacific Region, by the combination of absent frontal and supraorbital striae (Fig. 69) and presence of two marginal epipleural striae. Species of the genus Gnathoncus differ from Tomogenius by having a smaller body size, but chiefly by the absence of two large median foveae on the apex of prosternal process (Fig. 702; present in Tomogenius).

Figures 69–80. 

69 Gnathoncus communis (Marseul, 1862) head, dorsal view 70 antennal club, ventral view showing sensory structures of the antenna 71 mentum, ventral view 72 prosternum 73 mesoventrite 74 lateral disc of metaventrite + metepisternum 75 protibia, dorsal view 76 ditto, ventral view 77 mesotibia, dorsal view 78 ditto, ventral view 79 metatibia, dorsal view 80 ditto, ventral view.

USA: New York.

Saprinus communis Marseul, 1862: Lectotype, present designation: most likely a ♀, pinned, left protarsus, right mesotarsus and both metatarsi missing, with the following labels: “Gnathoncus / communis m / Albany / illegible” (green round label, written); followed by: “2937” (written); followed by: “Horn / 19 man / 73” (written); followed by: “1 communis / Et. univ. illegible” (written); followed by: “MUSEUM PARIS / COLL. / DE MARSEUL 1890” (green label, printed); followed by: “TYPE” (red-printed label); followed by: “Saprinus communis / Marseul, 1862 / LECTOTYPE 2014 / des. T. Lackner” (red label, written) (MNHN). This species was described from unknown number of specimens and the lectotype designation fixes the identity of species.

Figures 81–89. 

81 Gnathoncus communis (Marseul, 1862) male terminalia after Ôhara (1994): 8^th sternite, ventral view 82 male terminalia: 8^th sternite + 8^th tergite, dorsal view 83 ditto, lateral view 84 male terminalia: apex of aedeagus, dorsal view 85 male terminalia: aedeagus, dorsal view 86 ditto, lateral view 87 male terminalia: 9^th + 10^th tergites, dorsal view 88 ditto, lateral view 89 male terminalia: spiculum gastrale, ventral view.

NEW ZEALAND. North Island. AK: 2 ♂♂ & 1 ♀, Lynfield, 19.vi.1976, G. Kuschel (hen house) (NZAC); 1 spec., ditto, but 14.xii.1980 (chicken yard) (NZAC); 2 specs., Ranui, Henderson, 29.xi.1955, K.A.J. Wise (ex debris in poultry feed shed) (NZAC); 1 ♂, 1 ♀ & 1 spec., Tropicana Dr., 19.vi.1976, G. Kuschel (hen house) (NZAC); 1 spec., ditto, but 14.viii.1976 (NZAC); 7 ♂♂ & 3 ♀♀, Mahinerang, 26.i.1977, J. Painter (Sturnus vulgaris nest) (NZAC); 3 ♂♂ & 3 ♀♀, Kumeu, 21.vi.1975, J.C.Watt (ex fowl manure) (NZAC). BP: 1 ♂, Tauranga, 13.xi.1976, R.J. Ball (on plant) (NZAC). HB: 2 specs., Longlands, ii.1976, J.A. McKeenan (ex starling nest) (NZAC); 1 spec., Havelock North, 14.xi.1977, collector unknown (on large Pinus sp., ex nest of Sturnus vulgaris) (NZAC); 2 ♂♂ & 3 ♀♀, Graeme Friis’ Farm, Twyford, 2.xi.1976, B.A. Holloway (poultry manure) (NZAC).

In New Zealand found mostly in bird’s nests, chicken coops, less commonly on bird carrion.

Described from New York (USA), where it was probably introduced and widespread across North America. This species is normally spread in Europe, North Africa, Russian Far East, Japan and has been introduced into Australia (Mazur 2011). Mazur (1990:744) reports this species as introduced into Australia (ACT: Canberra). The specimen Mazur examined should be housed in ANIC. However, we were unable to examine it or any other specimens of G. communis from Australia. From New Zealand reported already by Kuschel (1990).

G. communis can be confused with G. rotundatus, but can easily be separated from it by the absence of median fossa of prosternum (present with G. rotundatus), dense and confluent elytral punctation (especially in the apical half of the elytra) and different male genitalia. Most important diagnostic characters to separate the two Gnathoncus species present in the region are outlined in the identification key. We chose not to fully re-describe Gnathoncus communis here, leaving its re-description to the revision of the genus Gnathoncus. For the sake of the better species recognition, as well as for the easier separation from the related G. rotundatus, we decided to depict it here, including its male terminalia.

Poland: Masuria: Ostróda.

None examined for this study. Kugelann’s collection was destroyed during the WWII (T. Huflejt, Warsaw, Poland, pers. comm). It is not known, whether it contained the type material of Hister rotundatus or not.

Figure 90. 

Gnathoncus rotundatus (Kugelann, 1792) habitus, dorsal view.

NEW ZEALAND. North Island. CO: 1 ♂, Earnscleugh, 28.x.1979, G. McLaren (suction trap) (NZAC). HB: 2 ♂♂, Longlands, ii.1976, J.A. McLemmans (starling nest) (NZAC); 1 ♂, Cape Kindnappers, Black Reef, 10.iii.1980, C.F. Butcher & M.F. Tacker (bird’s nest) (NZAC); 1 ♀, Motupiko, 9.ix.1951, G.G. Hole (NZAC); 6 ♂♂ & 7 ♀♀, Twijford, Hawke Bay, 2.xi.1976, B.A. Holloway (poultry manure) (NZAC); 34 ♂♂ & 29 ♀♀, Hastings, Twijford Poultry Farms, 21.x.1976, B.A. Holloway (poultry manure) (NZAC). AK: 1 spec., Auckland, New Zealand, Broun Coll. (BMNH); 1 ♂, Milford, 4.iv.1979, T. Thomas (nest of Passer domesticus) (NZAC); 1 ♂, Pareora Gorge, ii.1951, L.J. Dumbleton (shag nest) (NZAC); 1 ♂, Lynfield, 14.xii.1980, G. Kuschel (hen house) (NZAC); 1 spec., ditto, but 7.iii.1977 (sheep) (NZAC); 1 ♀, ditto, but 27.ii.1978, B.A. Holloway (guinea pig den material) (NZAC); 1 ♀, Kumeu, 21.vi.1975, J.C. Watt (ex fowl manure) (NZAC); 4 ♂♂ & 4 ♀♀, Nelson, 29.i.1971, E.W. Valentine (NZAC). South Island. 1 spec., Addington, Christchurch, 14.ix.1969, M.G. McPherson (inside house) (NZAC); 1 spec., Lincoln College, 18.i.1967, C. Boswell (NZAC); 2 specs., ditto, but 22.xii.1966 (NZAC); 1 spec., ditto, but 19.xi.1966 (NZAC); 2 specs., Prebbleton, 21.ix.1968, R.J. McPherson (ex fowl manure) (NZAC). DN: 6 specs., Otago Peninsula, Tairoa Head, 26.iii.1970, J.R. Jackson (ex Larus dominicanus or Phalacrocorax punctatus nest) (NZAC); 1 spec., Lincoln College, Hilgendorf wing, 8.x.1973, R.D. Welsh (ex window) (NZAC). MC: 2 specs., Banks Penisula, Murrays Mistake, 10.i.1970, J.R. Jackson (ex Phalacrocorax punctatus nest) (LUNZ); 1 spec., Hawkes Bay, Mangaome River, Dartnoor, 28.i.1959, B.M. May (on shingle) (NZAC); 1 spec., Lincoln College, 29.xii.1966, C.C. Boswell (NZAC); 1 ♀, ditto, but 18.vi.1967 (NZAC); 1 ♂, Rangiara, 7.xi.1947, D. Spiller (NZAC); 1 ♀, Maharoa, 9.xii.1941, J. Spiller (ex copra) (NZAC). Unknown localities: 1 ♂, no data (NZAC); 1 spec., Rakaia, 20.ii.1912, Broun Coll. (BMNH).

Figures 91–98. 

91 Gnathoncus rotundatus (Kugelann, 1792) head, dorsal view 92 antennal club, ventral view 93 mentum, ventral view 94 prosternum 95 mesoventrite + metaventrite 96 lateral disc of metaventrite + metepisternum 97 protibia, ventral view 98 mesotibia, ventral view.

AUSTRALIA. Tasmania: 1 ♂ & 1 ♀, Launceston, 4 & 9.ii.1915, Littler Collection (NZAC). Australian Capital Territory: 2 ♂♂, Lyneham, 5.iv.1969, B.P. Moore, garden compost (ANIC).

A typical synanthrope, found mainly in anthropogenic settings, but also apparent inquiline of bird’s nests.

Holarctic Region, Republic of South Africa, Chile, Saint Paul Island (Mazur 2011). From New Zealand reported already by Kuschel (1990), newly reported here for Australia (Fig. 754).

Figures 99–102. 

99 Gnathoncus rotundatus (Kugelann, 1792) mandibles, dorsal view 100 labrum: left half depicting dorsal view and right half depicting epipharynx 101 anterior margin of mentum, ventral view 102 antennal club, ventral view showing sensory structures of the antenna.

This is a widely distributed species exhibiting some degree of variability regarding the length of elytral striae and density of dorsal punctation. Most important diagnostic characters to separate the two Gnathoncus species present in the region are outlined in the identification key. We chose not to fully re-describe Gnathoncus rotundatus here, the reader is instead referred to Lackner (2010: 118) for the detailed re-description. For the sake of better species recognition, as well as for easier separation from the related G. communis, we decided to depict it here, including its male terminalia.

Figures 103–110. 

103 Gnathoncus rotundatus (Kugelann, 1792) male terminalia after Ôhara (1994): 8^th sternite, ventral view 104 male terminalia: 8^th sternite + 8^th tergite, dorsal view 105 ditto, lateral view 106 male terminalia: 9^th + 10^th tergites, dorsal view 107 ditto, lateral view 108 male terminalia: spiculum gastrale, ventral view 109 male terminalia: aedeagus, dorsal view 110 ditto, lateral view.

Diagnosis of this genus is here based solely on the species H. hyla that has been recorded from the Australopacific Region. A small, ovoid beetle, cuticle dark brown with a faint metallic tinge; frontal stria complete, supraorbital stria lacking, frontal disc punctate. Pronotal depressions absent, almost entire dorsal surface in punctures. Prosternal foveae large; outer margin of protibia with 6–8 low teeth topped by denticle.

Hypocacculus species are normally distributed in open landscapes, often in arid places, where they are usually found on carrion or in mammal excrement. Examined specimens of H. hyla do not carry any biological information on their labels.

The bulk of species of Hypocacculus are distributed in Palaearctic and Afrotropical Regions, with several species occurring also in the Oriental Region (Mazur 2011). Hypocacculus (H.) hyla (Marseul, 1864) was described from New Guinea and is the sole representative of the genus present in the Australopacific Region (Fig. 755).

Species of the genus Hypocacculus are similar in size and general habitus to those of Hypocaccus or Chalcionellus. They differ from members of the genus Hypocaccus by having a punctate frons, which is smooth and adorned with several deep rugae in Hypocaccus, and from Chalcionellus aeneovirens, the sole species of the genus Chalcionellus present in the region, by the absence of pronotal depressions, present in C. aeneovirens.

See above for the diagnosis of Hypocacculus s.l.

See the biology of Hypocacculus s.l.

Members of this subgenus are spread in southern Palaearctic and Afrotropical Regions, with three species occurring also in the Oriental Region. Species H. (H.) metallescens (Erichson, 1834) has been introduced to USA (Florida) (Mazur 2011). In the Australopacific Region, one species, H. (H.) hyla, is present and was described from New Guinea, and occurs also in Indonesia (Java and Sunda), Vietnam, and Myanmar (Mazur 2011). Newly reported from India (see below).

New Guinea.

Saprinus hyla Marseul, 1864: Lectotype, present designation: ♀, side-mounted on a triangular card with right metatibia (metatarsus missing) and left antennal club broken off and glued to the same mounting card, left metatibia missing, last two segments of left mesotarsus missing, both protarsi missing, with the following labels: “Saprinus / hyla / N. Guinée / Wallace illegible text” (yellow, round label, written); followed by: “Saprinus / Hyla / N. Guinea” (written); followed by: “MUSEUM PARIS / Coll. De Marseul / 2842-90” (printed); followed by: “TYPE” (red-printed label); followed by: “Saprinus / hyla Marseul, 1864 / LECTOTYPE / Des. by Lackner & / Leschen, 2011” (red label, written) (MNHN). This species has been described from an unknown number of specimens and the lectotype designation fixes its taxonomic identity.

PAPUA NEW GUINEA: 1 ♂, Huon Golf, Simbang, 1899, Biró leg. (HNMH). INDIA. Orissa: 1 ♂, Koraput Jeypore, 22.x.2006, G. de Rougemont (NCB).

Unknown, probably a saprobiont.

See above (Fig. 755).

The male specimen from India matches the form of the female lectotype, only differing by its color, which is somewhat darker, and the punctation of the elytra, which is less dense apically compared to the New Guinean lectotype.

Body length: PEL: 1.625–1.80 mm; APW: 0.625–0.75 mm; PPW: 1.25–1.625 mm; EL: 1.00–1.25 mm; EW: 1.425–1.75 mm.

Body (Fig. 111) rectangular oval, dorsal surface convex, ventral surface flattened, cuticle light to darker brown with bronze metallic luster, legs, mouthparts and antennae light brown.

Figure 111. 

Hypocacculus (Hypocacculus) hyla (Marseul, 1864) habitus, dorsal view.

Figure 112. 

Hypocacculus (Hypocacculus) hyla (Marseul, 1864) propygidium + pygidium.

Antennal scape (Fig. 113) with lower margin carinate, with two short setae; antennal club (Fig. 113) round, lower half of its surface glabrous, upper half (approximately) with thick short yellow sensilla; sensory structures of antennal club not examined.

Mandibles stout, with rounded outer margin strongly curved inwardly, dorsally with sparse fine punctures, acutely pointed, sub-apical tooth on inner margin of left mandible not examined; labrum punctate, dorsally with median costiform elevation; terminal labial palpomere elongated, its width about one-third its length, about twice as long as penultimate; mentum square-shaped, anterior angles slightly produced, anterior margin with deep median excavation, surface of mentum with sparse setae; stipes triangular, with three short setae; terminal maxillary palpomere elongated, its width about one-third its length, approximately twice as long as penultimate; other mouthparts not examined.

Clypeus (Fig. 113) sub-quadrate, rounded laterally, slightly concave medially, anterior margin slightly carinate, surface mesad to it with dense punctures; frontal stria complete and slightly carinate, slightly outwardly arcuate, continuous with weakly carinate supraorbital stria; frontal disc (Fig. 113) with regular punctation, punctures separated by several times their diameter; eyes slightly flattened, visible from above.

Figures 113–118. 

113Hypocacculus (Hypocacculus) hyla (Marseul, 1864) head, dorsal view 114 prosternum + mesoventrite 115 lateral disc of metaventrite + metepisternum 116 protibia, dorsal view 117 ditto, ventral view 118 metatibia, ventral view.

Pronotal sides (Fig. 111) slightly convergent anteriorly, apical angles obtuse; pronotal depressions absent; marginal pronotal stria complete, carinate; disc of pronotum laterally with shallow, moderately dense punctation, punctures become sparser and finer medially; along pronotal base a denser row of ovoid punctures present; pronotal hypomeron with microscopic sparse amber setae; scutellum small, visible.

Elytral humeri not particularly prominent; elytral epipleura with scattered microscopic punctures; marginal epipleural stria complete; marginal elytral stria straight and carinate; apical elytral stria absent. Humeral elytral stria well impressed on basal third; inner subhumeral stria present as a short median fragment; elytral disc with four deeply impressed dorsal elytral striae 1–4, all striae in punctures which are more prominent apically, striae about the same length, somewhat diminishing in length from first to fourth, reaching approximately two-thirds of elytral length apically; fourth elytral stria basally well connected with sutural elytral stria; sutural elytral stria well impressed, almost complete; basal third of elytral disc and elytral flanks only with scattered microscopic punctation, apical two-thirds with much denser and coarser punctation, punctures separated by about twice their diameter, on apical fifth punctation becomes even denser, punctures separated by their own diameter, extreme apex of elytra with a glabrous band.

Propygidium (112) on anterior half almost impunctate, on posterior half with dense punctures separated by about their diameter; pygidium (112) on basal third with punctures separated by about twice their own diameter, punctures becoming sparser and finer apically; extreme apex of pygidium almost impunctate.

Anterior margin of median portion of prosternum (Fig. 114) rounded; prosternal foveae large and deep; marginal prosternal stria present only laterally; prosternal process densely and coarsely punctate; carinal prosternal striae slightly divergent on prosternal apophysis, running parallel, not united apically; lateral prosternal striae carinate, widely united in front of carinal prosternal striae.

Discal marginal mesoventral stria (Fig. 114) anteriorly slightly inwardly arcuate, complete; disc of mesoventrite with deep and dense punctures, separated by about their own diameter, interspaces imbricate; meso-metaventral suture indistinct; meso-metaventral sutural stria bisinuate and undulate, slightly distanced medially from meso-metaventral suture; intercoxal disc of metaventrite on basal three-fourths with scattered fine punctures, in apical third and especially behind metacoxae much larger and deeper punctures present; lateral metaventral stria carinate, almost complete; lateral disc of metaventrite (Fig. 115) slightly excavate, with deep dense punctures of various sizes, apically becoming sparser; metepisternum + fused metepimeron with even denser and coarser setigerous punctures; lateral metepisternal stria present only on fused metepimeron.

Intercoxal disc of first abdominal ventrite almost completely striate laterally; disc laterally and anteriorly with punctures of various sizes, median part of disc almost impunctate, with alutaceous microsculpture.

Protibia (Fig. 116) slightly widening apically, outer margin with seven obtuse teeth topped by prominent denticle, diminishing in size in proximal direction, followed by two minute proximal denticles; setae of outer row thin, sparse, regular; setae of median row even shorter than those of outer row; protarsal groove deep; anterior protibial stria well impressed, bisinuate, shortened apically; protibial spur small, growing out from anterior protibial margin; outer part of posterior surface of protibia (Fig. 117) almost smooth, demarcation line between outer and median of posterior surface marked by double undulate stria, basally that stria with about seven strongly sclerotized setae; median part of posterior protibial surface almost smooth, imbricate, posterior protibial stria complete, fine, apically ending in two short denticles; inner margin with double row of short setae.

Mesotibia slender, outer margin with a row of several thin denticles growing in size apically; setae of outer row strongly sclerotized, about the size of denticles, growing in size apically; setae of median row present medially, much finer and shorter, posterior mesotibial stria not examined; anterior surface of mesotibia with another dense row of short denticles; anterior mesotibial stria straight and complete, terminates in two short denticles; mesotibial spur short; apical margin of mesotibia with three short denticles; claws of apical tarsomere bent, shorter than its half; metatibia (Fig. 118) similar to mesotibia, but denticles on its outer margin even sparser.

Male genitalia (based on a male from India: Orissa; see above). Eighth sternite (Figs 119–120) gradually narrowing apically, entirely fused medially; apex of eighth tergite (Fig. 120) shallowly emarginate; apex of eighth sternite with vela covered with dense microscopic setae. Eighth sternite and tergite fused medially (Fig. 121). Ninth sternite (Figs 122–123) typical for the subfamily; tenth tergite (Fig. 122) very small, basally inwardly arcuate. Spiculum gastrale (Figs 124–125) dilated on both ends, its body otherwise parallel-sided; basal end spatula-like. Aedeagus (Figs 126–127) gradually narrowing apically, parameres fused on their basal two-thirds (roughly); basal piece of aedeagus rather short, its ratio to fused parameres approximately 1:3.

Figures 119–127. 

119Hypocacculus (Hypocacculus) hyla (Marseul, 1864) male terminalia: 8^th sternite + 8^th tergite, ventral view 120 ditto, dorsal view 121 ditto, lateral view 122 male terminalia: 9^th + 10^th tergites, dorsal view 123 ditto, lateral view 124 male terminalia: spiculum gastrale, ventral view 125 ditto, lateral view 126 male terminalia: aedeagus, dorsal view 127 ditto, lateral view.

Diagnosis of this genus is based on the taxa that occur in the Australopacific Region. Small to moderately sized, often metallic beetles; frontal stria usually well developed, straight; frons coarsely punctate or with several to multiple short to long transverse rugae. Pronotal disc either smooth (subgenus Baeckmanniolus Reichardt, 1926) or adorned with longitudinal rugae or very coarse punctures (subgenera Hypocaccus s.str. or Nessus Reichardt, 1932). Pronotum devoid of pronotal depressions, hypomeron asetose; both sets of prosternal striae present, carinal prosternal striae approximate, usually united apically posterad united lateral prosternal striae; prosternal foveae present, often well developed. Protibia on outer margin with 5–11 teeth topped by denticle; metatibia on outer margin with two (Hypocaccus s.str., Nessus) or three (subgenus Baeckmanniolus) rows of denticles.

Figure 128. 

Hypocaccus (Nessus) interpunctatus interpunctatus (Schmidt, 1885) habitus, dorsal view.

Species of the subgenus Hypocaccus are found on the sandy shores of seas, lakes and rivers (sometimes also on inland sand dunes without the presence of water) where they prey upon dipteran larvae developing in various decomposing organic substances such as excrement, carcasses, seaweed, etc. Species of the subgenus Baeckmanniolus are confined to seashores with similar feeding habits (Lackner 2010).

With four recorded species (one of them introduced), the genus Hypocaccus is poorly represented in the Australopacific Region and has been collected in Australia, New Caledonia and New Guinea (Figs 753, 755, 756, 760). We would expect the widespread species H. (H.) brasiliensis (see Mazur 2011) or H. (H.) sinae to be eventually collected elsewhere within the Region.

Species of the genus Hypocaccus are typified by the presence of smooth to rugose frons, which can be also furnished with transverse rugae and cannot be confused with any other Australian Saprininae.

Key to the subgenera of the genus Hypocaccus of the Australopacific Region

The single introduced Australian species of the subgenus, H. (N.) interpunctatus interpunctatus can easily be differentiated from the rest of the Australopacific Saprininae by its small size (1.70–2.20 mm), presence of well developed prosternal foveae and a characteristic ‘mirror’ (=polished area) found within the second elytral interval. From the other subgenera this species differs by smaller size and absence of several deep rugae on frontal disc.

Figures 129–134. 

129Hypocaccus (Nessus) interpunctatus interpunctatus (Schmidt, 1885) head, dorsal view 130 antennal club, dorsal view 131 prosternum 132 mesoventrite 133 lateral disc of metaventrite + metepisternum, with visible mesotibia 134 metatibia, dorsal view.

Hypocaccus (Nessus) interpunctatus interpunctatus is found in decaying organic matter, most commonly dung or carrion. Often collected using pitfall traps; occasionally found also under stones on beaches.

In the Australopacific Region there is a single introduced species, Hypocaccus (Nessus) interpunctatus interpunctatus recorded from Western Australia (Fig. 753). This species has a scattered distribution: Sicily, Tanzania, Syria, Eritrea, Mozambique, Republic of South Africa, Mongolia and Turkey (Mazur 2011).

Mazur (2011) transferred the subgenus Nessus from Hypocacculus to genus Hypocaccus without comment.

Italy: Sicily.

Saprinus interpunctatus Schmidt, 1885: Lectotype, designated by Vienna & Colla in 2003, ♂, glued to a mounting card, with following labels: “Sicilien / Ragusa” (written); followed by: “coll. J. Schmidt” (printed); followed by: “Zool. Mus / Berlin” (printed); followed by: “interpunctat. / Schm. Typ” (written); followed by: “Type” (brick-red label, written); followed by: “interpuncta / tus Schmidt” (double black-margined label, written); followed by: “♂” (written); followed by: “LECTOTYPUS / Hypocacculus (Nessus) / interpunctatus (Schmidt, 1885) / Vienna & Colla des., 2003” (red label, printed) (ZMHUB). Paralectotypes, designated by Vienna & Colla in 2003, 2 ♀♀, one lacks its left hind leg, the other one lacks its left metatibia, with following labels: “Sicilia / Baudi” (written); followed by: “Type” (brick-red label, written); followed by: “Zool. Mus / Berlin” (printed); followed by: “♀” (written); followed by: “PARALECTOTYPUS / Hypocacculus (Nessus) / interpunctatus (Schmidt, 1885) / Vienna & Colla des., 2003” (ZMHUB). Another paralectotype, designated by Vienna & Colla in 2003, ♂, with genitalia extracted and mounted in Canada balsam on a separate slide under the specimen, with the same “Sicilien/Ragusa” label like the lectotype, followed by: “Zool. Mus / Berlin” (printed); followed by: “Saprinus / interpunctatus Schmidt / Coll. Schmidt-Bickhardt” (printed); followed by: “♂” (written); followed by: “PARALECTOTYPUS / Hypocacculus (Nessus) / interpunctatus (Schmidt, 1885) / Vienna & Colla des., 2003” (ZMHUB).

Figures 135–143. 

135Hypocaccus (Nessus) interpunctatus interpunctatus (Schmidt, 1885) male terminalia: 8^th sternite + 8^th tergite, ventral view 136 ditto, dorsal view 137 ditto, lateral view 138 male terminalia: 9^th + 10^th tergites, dorsal view 139 ditto, lateral view 140 male terminalia: spiculum gastrale, ventral view 141 ditto, lateral view 142 male terminalia: aedeagus, dorsal view 143 ditto, lateral view.

AUSTRALIA. Western Australia: 1 spec., Dardanup, 1.ii.1979, G. Hall (howden trap) (ANIC); 69 specs., 15 km S Busselton, 4.xi.1982, G.P. Hall (ANIC); 30 specs., ditto, but 19.i.1983 (ANIC); 46 specs., ditto, but 9.viii.1983 (ANIC); 1 spec., ditto, but 13.viii.1983 (ANIC); 11 specs., ditto, but 16.vi.1983 (ANIC); 29 specs., ditto, but 12.i.1983 (ANIC); 25 specs., ditto, but 13.i.1983 (ANIC); 26 specs., ditto, but 27.i.1983 (ANIC); 2 specs., ditto, but 27.x.1982 (ANIC); 2 specs., ditto, but 21.x.1982 (ANIC); 4 specs., ditto, but 23.x.1982 (ANIC); 3 specs., ditto, but 11.viii.1983 (ANIC); 3 specs., ditto, but 25.viii.1983 (ANIC); 1 spec., ditto, but 17.viii.1983 (ANIC); 19 specs., ditto, but 15.i.1983 (ANIC); 17 specs., 5 km NE Dardanup, 28.x.1981, G.P. Hall (ANIC); 71 specs., ditto, but 20.x.1981 (ANIC); 20 specs., ditto, but 16.x.1981 (ANIC); 39 specs., ditto, but 4.xi.1982 (ANIC); 9 specs., ditto, but 19.x.1982 (ANIC); 30 specs., ditto, but 27.x.1982 (ANIC); 13 specs., ditto, but 21.x.1982 (ANIC); 5 specs., ditto, but 20.x.1982 (ANIC); 24 specs., ditto, but 23.x.1982 (ANIC); 3 specs., ditto, but 27.x.1982 (ANIC); 8 specs., ditto, but 19.i.1983 (ANIC); 11 specs., ditto, but 27.i.1983 (ANIC); 18 specs., ditto, but 15.i.1983 (ANIC); 6 specs., ditto, but 13.i.1983 (ANIC); 8 specs., ditto, but 12.i.1983 (ANIC); 4 specs., ditto, but 11.i.1983 (ANIC); 16 specs., ditto, but 16.vi.1983 (ANIC); 1 spec., ditto, but 8.vi.1983 (ANIC); 2 specs., ditto, but 11.viii.1983 (ANIC); 6 specs., ditto, but 10.viii.1983 (ANIC); 13 specs., ditto, but 9.viii.1983 (ANIC); 2 specs., ditto, but 14.x.1981 (ANIC); 1 spec., ditto, but 13.x.1981 (ANIC); 1 spec., ditto, but 12.x.1981 (ANIC); 1 spec., ditto, but 1.ii.1979, 33.24S 115.45E, J.D. Majer (ANIC); 4 specs., Rossiter Bay, Cape Le Grand NP, 26.xii.1985, C. Reid (under stones, low heath by beach) (ANIC).

Saprobiont, collected on carrion and dung alike; several Australian specimens were collected under stones or using traps.

Australia: Western Australia (Fig. 753).

This species consists of two subspecies: H. interpunctatus interpunctatus that is known from the above-mentioned countries and H. interpunctatus muelleri Colla, Gomy & Vienna, 2004 occurring in Kenya and southern Ethiopia. The two subspecies differ chiefly in elytral punctation and intervals among striae; furthermore there are differences in the punctation of pygidium and clypeus (Colla, Gomy and Vienna 2014). Hypocaccus (N.) interpunctatus interpunctatus is diagnosed above, as well as provided with a differential diagnosis that separates it from other Australopacific taxa. Most important differences between the Australopacific members of the genus Hypocaccus are outlined in the key to the subgenera above. We chose not to fully re-describe Hypocaccus (N.) interpunctatus interpunctatus here, leaving its re-description to the revision of the genus Hypocaccus. For the sake of the better species recognition, however, we decided to depict it here, including its male terminalia.

From the sole Australian representative of the subgenus Nessus, H. (N.) interpunctatus interpunctatus, members of the nominotypical subgenus differ by larger size and presence of several deep rugae on otherwise almost glabrous frontal disc. From the sole Australopacific member of the subgenus Baeckmanniolus, H. (B.) varians varians, the two species of the nominotypical subgenus are easily separated by the presence of punctation on the pronotum.

Figure 144. 

Hypocaccus (Hypocaccus) brasiliensis (Paykull, 1811) habitus, dorsal view.

Both species of the nominotypical subgenus that occur in Australopacific Region are littoral, occurring under wrack on beach, one specimen of H. (H.) brasiliensis from Australia was also found on a riverbank.

Two very similar littoral psammophilous species, H. (H.) brasiliensis and H. (H.) sinae are present in Australopacific Region (Figs 755, 756, 760). Both species are most likely invaders from north (sensuMatthews and Bouchard 2008: 17), as they occur in the Palaearctic and Indo-Malaysian Regions and the genus Hypocaccus is widely represented there too. The species H. (H.) brasiliensis is tropicopolitan, distributed along beaches in all warmer regions around the globe (Mazur 2011); in Australia it was also found inland on a riverbank of the Colo River (Fig. 756).

Regarding the differences between the nominotypical subgenus and subgenus Baeckmanniolus, Bousquet and Laplante (2006: 80, 196) mention another character, the number of rows of denticles on the outer margin of metatibia (two in Hypocaccus s. str. versus three in the subgenus Baeckmanniolus) and treat the subgenus Baeckmanniolus as a distinct genus. For more discussion see Lackner (2010: 145).

Brazil.

Hister brasiliensis Paykull, 1811: Lectotype, ♀, designated by Dahlgren in 1968, together with extracted genitalia glued to a rectangular mounting card, right foreleg missing, right elytron missing, right metatibia broken off, glued next to the specimen, with the following labels: “24” (pencil-written); followed by: “Uppsala Univ. Zool. Mus. / Gyllenhals saml. TYP nr. / 1153” (red label, printed); followed by: “LECTOTYP / HYPOCACCUS / BRASILIENSIS / PAYK. G. DAHL- / GREN 27.II.1968” (written in black ink) (UUZM).

Figures 145–153. 

145Hypocaccus (Hypocaccus) brasiliensis (Paykull, 1811) head, dorsal view 146 prosternum 147 mesoventrite 148 lateral disc of metaventrite + metepisternum 149 protibia, dorsal view 150 ditto, ventral view 151 propygidium + pygidium 152 metatibia, dorsal view 153 ditto, ventral view.

Saprinus apricarius Erichson, 1834: Lectotype, present designation: most likely a ♀, pinned, left hind leg missing, with the following labels: “Aegypt / xxxxiv Er.(?)” (dark green label, written); followed by: “49233” (printed); followed by: “Hist. -Coll. (Coleoptera) / Nr. 49233 / Saprinus apricarius Er. x / Aegypt., Ehrenberg / Zool. Mus. Berlin” (black-framed label, printed); followed by: “Saprinus / apricarius / Erichson, 1834 / LECTOTYPE 2014 / des. T. Lackner” (red label, written) (ZMHUB). Paralectotypes, present designation: 1 spec., possibly a ♀, pinned, with the following label: “Hist. -Coll. (Coleoptera) / Nr. 49233 / Saprinus apricarius Er. x / Aegypt., Ehrenberg / Zool. Mus. Berlin” (black-framed label, printed); followed by: “Saprinus / apricarius / Erichson, 1834 / PARALECTOTYPE 2014 / des. T. Lackner” (red label, written) (ZMHUB). 1 spec., possibly a ♀, pinned, with the following labels: “49233” (printed); followed by: “Hist. -Coll. (Coleoptera) / Nr. 49233 / Saprinus apricarius Er. x / Sicil, Dahl / Zool. Mus. Berlin” (black-framed label, printed); followed by: “apricarius / Er. / Sicil. Dahl.” (black-framed label, written); followed by: “Saprinus / apricarius / Erichson, 1834 / PARALECTOTYPE 2014 / des. T. Lackner” (red label, written) (ZMHUB). 1 spec., possibly a ♀, pinned, with the following labels: “47”; followed by: “Hist. -Coll. (Coleoptera) / Nr. 49233 / Saprinus apricarius Er. x / Sicil, Dahl / Zool. Mus. Berlin” (black-framed label, printed); followed by: “apricarius / Er. / Sicil. Dahl.” (black-framed label, written); followed by: “Saprinus / apricarius / Erichson, 1834 / PARALECTOTYPE 2014 / des. T. Lackner” (red label, written) (ZMHUB). 1 spec., pinned, with the following labels: “54”; followed by: “Hist. -Coll. (Coleoptera) / Nr. 49233 / Saprinus apricarius Er. x / Sicil, Dahl / Zool. Mus. Berlin” (black-framed label, printed); followed by: “apricarius / Er. / Sicil. Dahl.” (black-framed label, written); followed by: “Saprinus / apricarius / Erichson, 1834 / PARALECTOTYPE 2014 / des. T. Lackner” (red label, written) (ZMHUB). 2 specs., pinned, with the following labels: “Hist. -Coll. (Coleoptera) / Nr. 49233 / Saprinus apricarius Er. x / Sicil, Dahl / Zool. Mus. Berlin” (black-framed label, printed); followed by: “apricarius / Er. / Sicil. Dahl.” (black-framed label, written); followed by: “Saprinus / apricarius / Erichson, 1834 / PARALECTOTYPE 2014 / des. T. Lackner” (red label, written) (ZMHUB). This species has been described from both Sicily and Egypt, among the type series we designate a single female specimen from Egypt as lectotype to fix the taxonomic identity of the species.

Saprinus bistrigifrons Marseul, 1855: Lectotype, present designation: ♂, pinned, right mesotarsus missing, with following labels: tiny pink label, followed by: “161 / Saprinus / bistrigifrons / m. / Mexique / illegible text” (green, round label, written); followed by: “MUSEUM PARIS / COLL. / DE MARSEUL 1890” (green, printed label); followed by: “TYPE” (red-printed label); followed by: “Saprinus / bistrigifrons / Marseul, 1855 / LECTOTYPE 2014 / des. T. Lackner” (red label, written) (MNHN). This species has been described from unknown number of specimens and the lectotype designation fixes the species identity.

Saprinus dentipes Marseul, 1855: Lectotype, present designation: probably a ♂, pinned, left metatarsus and right hind leg missing, with the following labels: “160 / Saprinus / dentipes / illegible text / Mexique / illegible text” (round, green label, written); followed by: “MUSEUM PARIS / COLL. / DE MARSEUL 1890” (green, printed label); followed by: “TYPE” (red-printed label); followed by: “Saprinus dentipes / Marseul, 1855 / LECTOTYPE 2014 / des. T. Lackner” (red label, written) (MNHN). This species has been described from unknown number of specimens and the lectotype designation fixes the species identity.

Saprinus piscarius Blackburn, 1903: Cotype, sex unidentified, with printed label: “Australia / Blackb’s Coll” (printed), followed by “Hypocaccus / vernulus Blackb” (written), followed by “piscarius Bl. Cot. / vernulus placed with Cotype / piscarius in the Bl. Coll” (written), followed by “SAMA Database / No. 25-029592” (printed); 1 spec., “Hypo. piscarius / co-type” (written), followed by “SAMA Database / No. 25-029591 (printed)”; 1 spec., “Saprinus / piscarius / Cotype” (written), followed by “SAMA Database / No. 25-029590” (written) (SAMA). This species was described from ‘near Adelaide’, as well as ‘Also from Victoria (Frankston, Mr. Kershaw)’ (Blackburn, 1903: 108). Because of the ambiguous label data (see above) as well as no locality indication of the cotype we decided not to designate a lectotype of this species.

Figures 154–160. 

154Hypocaccus (Hypocaccus) brasiliensis (Paykull, 1811) male terminalia: 8^th sternite + 8^th tergite, ventral view 155 ditto, dorsal view 156 ditto, lateral view 157 male terminalia: 9^th + 10^th tergites, dorsal view; spiculum gastrale, ventral view 158 male terminalia: 9^th + 10^th tergites; spiculum gastrale, lateral view 159 male terminalia: aedeagus, dorsal view 160 ditto, lateral view.

NEW CALEDONIA. 1 ♂ & 4 specs., Isle Atire, 1.ix.1982, T. Lovegrove (ex dead white-capped noddy) (AMNZ).

New Guinea. New Britain: 12 specs., Ralum, F. Dahl S. (ZMHUB; 2 specs. in coll TLAN).

AUSTRALIA. New South Wales: 2 specs., Tomakin, 35.49S 150.11E, 25.xii.1988, W. Dressler (ex dead shark head) (ANIC); 1 spec., Colo River, Near Colo, 8.xii.1979, D.P. Came (on river sands) (ANIC); 2 specs., Sydney, K.K. Spence (ANIC); 1 spec., Sydney, Lea (SAMA). South Australia: 4 specs., Adelaide, Griffith (SAMA); 1 spec., Marino Rook: Spelhore, 4.–5.i.1978, G. Szelényi leg. (HMNH). Western Australia: 2 specs., Geraldton (SAMA).

Saprinus dentipes has been synonymized with H. (H.) brasiliensis by Mazur (1987). However, upon inspecting its type specimen we conclude that it is not synonymous with H. (H.) brasiliensis, but with the species Hypocaccus (Baeckmanniolus) gaudens (J.L. LeConte, 1851) instead. Thus Saprinus dentipes Marseul, 1855 = Hypocaccus (Baeckmanniolus) gaudens (J.L. LeConte, 1851) syn. n. H. (H.) brasiliensis is a widespread and variable species with variable dorsal punctation. The taxonomic status of both H. (H.) brasiliensis and the following H. (H.) sinae requires further clarification, especially with respect to their rather similar genitalia and minimal differences in dorsal sculpture. Hypocaccus (H.) brasiliensis is above included in the key to the Australopacific species of Hypocaccus that separates it from the related Hypocaccus (H.) sinae. We chose not to fully re-describe Hypocaccus (H.) brasiliensis here, leaving its re-description to the revision of the genus Hypocaccus. For the sake of the better species recognition, however, we decided to depict it here, including its male terminalia.

China: Shanghai.

Figure 161. 

Hypocaccus (Hypocaccus) sinae (Marseul, 1862) habitus, dorsal view.

Saprinus sinae Marseul, 1862: Lectotype, present designation: ♂, mounted on tip of mounting card, with genitalia extracted, dismembered and placed in a separate plastic slide under the specimen, right antennal club, right protarsus and right hind-leg missing, with the following labels: tiny, dark blue rectangular label, followed by: “Saprinus / Sinae m. / Shanghai / Dej. 63” (round, yellow label, written); followed by: “Shanghai” (written); followed by: “52 (158a) Saprinus / Sinae m.60 / Shanghai” (written); followed by: “MUSEUM PARIS / COLL. / DE MARSEUL 1890” (printed); followed by: “TYPE” (red-printed label); followed by: “Saprinus sinae / Marseul, 1862 / LECTOTYPE 2014 / des. T. Lackner” (red label, written) (MNHN). This species was described from unknown number of specimens and the lectotype designation fixes the identity of the species.

Figures 162–173. 

162Hypocaccus (Hypocaccus) sinae (Marseul, 1862) head, dorsal view 163 antennal club, dorsal view 164 ditto, ventral view 165 mentum, ventral view 166 propygidium + pygidium 167 prosternum 168 mesoventrite 169 lateral disc of metaventrite + metepisternum 170 protibia, dorsal view 171 ditto, ventral view 172 mesotibia, ventral view 173 metatibia, ventral view.

Saprinus vernulus Blackburn, 1903: Lectotype, present designantion: ♀, mounted on a card with “♀” (printed), followed by: “7249” (mount card with a red number, printed); followed by: “N.S. Wales” (printed); followed by: “K. 270214” (printed); followed by: “Hypocaccus / vernulus / Cotype” (written); followed by: “Hypocaccus / vernulus Bl.” (written); followed by: “SYNTYPE” (yellow label, written); followed by: “Saprinus vernulus / Blackburn, 1903 / LECTOTYPE des. / T. Lackner 2016” (red label, written) (AMS). This specimen has been described based on unknown number of specimens and the lectotype designation fixes the identity of the species.

Figures 174–182. 

174Hypocaccus (Hypocaccus) sinae (Marseul, 1862) male terminalia: 8^th sternite + 8^th tergite, ventral view 175 ditto, dorsal view 176 ditto, lateral view 177 male terminalia: 9^th + 10^th tergites, dorsal view 178 ditto, lateral view 179 male terminalia: spiculum gastrale, ventral view 180 ditto, lateral view 181 male terminalia: aedeagus, dorsal view 182 ditto, lateral view.

AUSTRALIA. New South Wales: 2 specs., re-mounted on a single pin, with the same labels as the lectotype, (and certainly from the same collection – D. Smith pers. comm.) except the original mount with red number “7248” instead of “7249” and with “K270229” instead of “K. 270214” of the lectotype (AMS). 2 ♂♂, without further data (MAMU); 9 specs., New South Wales, without further data (SAMA); 3 specs., Sydney, Lea (SAMA). South Australia: 3 specs., Adelaide, Griffith (SAMA).

A typical littoral psammophile found on beaches.

Described from Shanghai (China), distributed along the coasts of China, Indonesia, Russian Far East, Thailand, Vietnam, Cambodia, Myanmar, Japan, Korea, Malaysia, Australia (Mazur 2011; present paper). Reported also from Afghanistan by Thérond (1969) together with the preceding species. From the Australopacific Region found only in Australia (New South Wales and South Australia; Fig. 756).

Blackburn (1903: 108) mentioned ‘distinctly green colour in combination with elytra having a very strong external subhumeral stria’ as the main distinguishing characters between H. vernulus and H. sinae. Since the external morphological characters of many Saprininae species are known to be intra-specifically rather variable, we assign the characters used by Blackburn for the separation of H. vernulus from H. sinae as intra-specific variation and synonymize the two species. A single syntype of Saprinus vernulus Blackburn, 1903 (currently Hypocaccus (Hypocaccus) vernulus) housed at ANIC has been inspected and we can herewith conclude that it is conspecific with the species Hypocaccus (Hypocaccus) sinae (Marseul, 1862); thus Hypocaccus (Hypocaccus) vernulus Blackburn, 1903 = Hypocaccus (Hypocaccus) sinae (Marseul, 1862) syn. n. Hypocaccus (H.) sinae is above included in the key to the Australopacific species of Hypocaccus that separates it from the related Hypocaccus (H.) brasiliensis. We chose not to fully re-describe Hypocaccus (H.) sinae here, leaving its re-description to the revision of the genus Hypocaccus. For the sake of the better species recognition, however, we decided to depict it here, including its male terminalia.

The single Australopacific taxon, H. (B.) varians varians can be separated from other congeners by almost smooth pronotum that bears fine punctation in the apical pronotal angles.

Figure 183. 

Hypocaccus (Baeckmanniolus) varians varians (Schmidt, 1890) habitus, dorsal view.

H. (B.) varians varians is a typical littoral psammophilous taxon, found on the beach under dead fish, algae or coastal wrack.

In the Australopacific Region there is only one species present, H. (B.) varians varians, recorded from Western Australia, Northern Territory and Queensland.

Japan.

Saprinus varians Schmidt, 1890: Lectotype, present designation: ♂, glued on the mounting card, with the following labels: “Japan” (written); followed by: “Type” (brick-red label, printed); followed by: “coll. J. Schmidt” (printed); followed by: “varians / m.” (black-yellow margined label, written); followed by: “Saprinus / varians Schmidt / Coll. Schmidt-Bickhardt” (printed); followed by: “Saprinus varians / Schmidt, 1890 / LECTOTYPE 2014 / des. Lackner & Leschen” (red label, written) (ZMHUB). Paralectotypes, present designation: 4 specs., probably ♀♀, carrying the same hand-written labels “Japan” and “Type” and the printed label “Saprinus / varians Schmidt / Coll. Schmidt-Bickhardt”, followed by the paralectotype labels identical to that of the lectotype (ZMHUB). This species has been described from unknown number of specimens and the lectotype designation fixes the taxonomic identity of the species.

Figures 184–192. 

184Hypocaccus (Baeckmanniolus) varians varians (Schmidt, 1890) head, dorsal view 185 antennal club, ventral view 186 mentum, ventral view 187 propygidium + pygidium 188 prosternum 189 mesoventrite 190 lateral disc of metaventrite + metepisternum 191 protibia, dorsal view 192 ditto, ventral view.

AUSTRALIA. Western Australia: 1 spec., Monte Bello Island (ANIC). Northern Territory: 1 ♂ & 1 ♀, Groote Eylandt, N.B. Tindale (SAMA). Queensland: 1 ♀, Cairns (SAMA).

See above.

Figures 193–201. 

193Hypocaccus (Baeckmanniolus) varians varians (Schmidt, 1890) male terminalia: 8^th sternite + 8^th tergite, ventral view 194 ditto, dorsal view 195 ditto, lateral view 196 male terminalia: 9^th + 10^th tergites, dorsal view 197 ditto, lateral view 198 male terminalia: spiculum gastrale, ventral view 199 ditto, lateral view 200 male terminalia: aedeagus, dorsal view 201 ditto, lateral view.

Described from Japan, the nominotypical subspecies is found also on Sakhalin Island, Taiwan, Philippines, Vietnam, Sri Lanka, and Australia. The two other subspecies are H. (B.) varians hatsune Nakane, 1977, described from the Ogasawara Islands (southern Japan) and H. (B.) varians continentalis Reichardt, 1941 described from Russian Far East (Primorsky Kray) and found also in eastern China (Shandong) (Mazur 2011).

The species H. (B.) varians has a faint pronotal punctation indicating that it should be transferred into the nominotypical subgenus Hypocaccus. According to Bousquet and Laplante (2006; see above) the outer margin of metatibia of the subgenus Baeckmanniolus has three rows of denticles instead of two rows present in Hypocaccus. The metatibia of Hypocaccus (B.) varians possesses two rows on the metatibia supplemented by another 2 (occasionally 3) denticles between the two rows. The presence of vague and weakened pronotal punctation and by possessing denticles between the two rows on the outer margin of metatibia, this species represents a transitional form between the two subgenera and its exact taxonomic position should be determined by a thorough morphological study of all known species of Hypocaccus s. l. Hypocaccus (B.) varians varians is diagnosed above, as well as provided with a differential diagnosis that separates it from other Australopacific taxa. Most important differences between the members of the genus Hypocaccus are outlined in the key to the subgenera above. We chose not to fully re-describe Hypocaccus (B.) varians varians here, leaving its re-description to the revision of the genus Hypocaccus. For the sake of the better species recognition, however, we decided to depict it here, including its male terminalia.

Iridoprinus myrmecophilus sp. n.

Figure 202. 

Iridoprinus myrmecophilus sp. n., habitus, dorsal view.

Cuticle light brown, elytra with strong blue iridescent metallic luster; frontal stria prolonged far onto clypeus; antennal club large, oval and depressed dorso-ventrally; pronotal disc on apical two-thirds coriarious-punctate, punctures forming elongate wrinkles, confluent; elytra densely imbricate-punctate, punctures with microscopic setae; five dorsal elytral striae present, curved and carinate; inner subhumeral and sutural elytral striae absent; abdominal segments dorsally with microscopic setae; prosternal foveae absent; metepisternum with deep elongate groove for reposing mesotarsus; pro- and mesotibiae slightly dilated.

The paratype has been collected in the nest of Meat Ant Iridomyrmex purpureus (Smith, 1858). Based on the collecting circumstances of the paratype, as well as its morphological characters (dilated tibiae, metepisternum with groove for reposing mesotarsus) we presume that the newly described taxon is a myrmecophile.

Endemic to Australia; known from Northern Territory and New South Wales (Fig. 757).

The generic epithet is created by combining part of the name of the host ant ‘Iridomyrmex’ with part of the word Saprinus (- prinus); the specific epithet of this new taxon relates to its apparently myrmecophilous biology.

By the presence of five strongly curved and carinate dorsal elytral striae and absence of inner subhumeral and sutural striae and presence of peculiar deep longitudinal groove on the metepisternum for the accommodation of mesotarsus, this myrmecophilous Australian genus cannot be confused with any other currently known higher taxon of Saprininae. This is a highly autapomorphic genus (Lackner 2014d), though, due to the paucity of material, critical characters from the antenna and mouthparts were not examined. However, this is the only Saprininae from the Australopacific and Indo-Malayan Regions collected from ant nests, indicating that it may be an obligate myrmecophile (for the outline of ant-inquilinous Saprininae see Lackner 2017). The deep longitudinal groove of the metepisternum to accommodate the mesotarsus while retracted as well as setose body (the microsetae are presumed to exude appeasing liquid substances which are licked by ants), dilated tibiae that probably serve to cover larger space of the venter when retracted, are possible adaptations to life inside ant nests. Its role inside the ant community is unknown, and it does not have obvious trichomes as in other obligate inquilines such as Chlamydopsinae. This taxon was included in the published phylogeny of the subfamily by the senior author (Lackner 2014d) under the name ‘Saprininae gen. n. (Australia)’. In the published cladogram it was recovered as sister to another Australian taxon of unknown biology (Saprinodes falcifer Lewis, 1891); this relationship was supported by one ‘strong’ and two ‘weaker’ synapomorphies. It is interesting to note that another apparent myrmecophile, Euspilotus (Platysaprinus) latimanus (Schmidt, 1890) has been recovered sister to this clade, but this purported monophyly should be regarded as spurious since it might be based on homoplasy rather than homology (the characters supporting this triad are among the ‘weak’ synapomorphies and the resolution of the tree is rather low).

Australia: Coniston Station near Alice Springs.

Holotype, ♂, side-mounted on a triangular card with male genitalia glued to the same card as specimen with the following labels: “Coniston Station, / near Alice Springs / N.W. Mules” (printed), with hand-written text on the reverse side: “Nov 1. Dec.31 1931”; “Good/Saprinus” (hand-written); followed by: “SAMA Database/No.25-029415” (printed); followed by: “10-157” (yellow, pencil-written label added by the senior author); followed by: “Iridoprinus / myrmecophilus / n. gen. & sp. / HOLOTYPE / det. T. Lackner 2010” (red label, written) (SAMA). Paratype, ♀, labelled: “AUSTRALIA / Kudgee / 16.xii.[19]72 / B.P. Moore (printed-written), with written text on the reverse side: “In nest of Iridomyrmex purpureus”; followed by: “10-139” (yellow, pencil-written label added by the senior author); followed by: “Iridoprinus / myrmecophilus / n. gen. & sp. / PARATYPE / det. T. Lackner 2010” (red label, written) (ANIC).

Body length (only 1 specimen, the holotype was measured): PEL: 3.00 mm; APW: 0.85 mm; PPW: 2.35 mm; EL: 2.00 mm; EW: 2.65 mm.

Body (Fig. 202) rectangular oval, moderately convex from above, underside slightly flattened, cuticle light brown, pronotum rufous, elytra with blue iridescent metallic luster; legs and body appendages rufo-castaneous.

Antennal scape (Fig. 203) triangular, thickened, with several setae; club (Fig. 204) rather large, balloon-shaped, on apical two-thirds with short dense sensilla intermingled with sparse longer erect sensilla; sensory structures of antennal club not examined.

Figures 203–211. 

203 Iridoprinus myrmecophilus sp. n. head, dorsal view 204 antennal club, dorsal view 205 propygidium + pygidium 206 prosternum 207 mesoventrite 208 lateral disc of metaventrite + metepisternum 209 protibia, dorsal view 210 ditto, ventral view 211 mesotibia, ventral view.

Mandibles with almost rectangular outer margin, acutely pointed, sub-apical tooth on inner margin of left mandible obtuse; mentum sub-trapezoid, anterior margin almost straight, without notch; other parts of mouth not examined.

Clypeus (Fig. 203) dorsally flattened, rounded laterally, with fine elongated punctures, separated by several times their diameter; supraorbital stria well developed, frontal stria carinate, prolonged onto clypeus; frontal disc (Fig. 203) densely punctuate, punctures deep, separated by about their diameter becoming elongate anteriorly, interspaces imbricate; eyes convex, well visible from above.

Pronotal sides (Fig. 202) feebly convergent anteriorly, apical angles obtuse, marginal pronotal stria present, laterally carinate, absent behind head; disc behind head with almost glabrous triangularly shaped ‘mirror’ (=polished area) with only scattered microscopic punctures, rest of pronotal disc coriarious-punctate, punctures becoming denser and coarser laterally, forming confluent elongate wrinkles; pronotal hypomeron glabrous; scutellum small, well visible.

Elytral epipleuron with microscopic punctures furnished with short setae; marginal epipleural and marginal elytral striae well impressed, complete; marginal elytral stria carinate, continuous along elytral apex as weakened apical elytral stria; humeral elytral stria inconspicuous (absent?); inner subhumeral stria absent; elytral disc with five curved strongly carinate dorsal elytral striae 1–5, striae sub-equal in length, fifth stria the shortest; sutural elytral stria absent. Entire elytral disc coarsely and very densely punctate, punctures rugulose, separated by less than half their diameter, each puncture with microscopic yellow seta well visible from lateral view.

Propygidium (Fig. 205) transverse, about four times as broad as long, partially covered by elytra, its punctation somewhat less dense and coarse than that of the elytra, punctures adorned with setae; pygidium (Fig. 205) with similarly dense and coarse setigerous punctures like those of the elytra, becoming sparser and finer towards apex.

Anterior margin of median portion of prosternum (Fig. 206) almost straight; marginal prosternal stria interrupted; prosternal process flattened, interspace between carinal prosternal striae excavated, smooth, laterally with scattered microscopic punctation, each puncture with minuscule seta; carinal prosternal striae (Fig. 206) divergent on both ends, united anteriorly; lateral prosternal striae carinate, almost straight, attaining apices of united carinal prosternal striae.

Anterior margin of mesoventrite (Fig. 207) almost straight; discal marginal mesoventral stria present only anteriorly, straight; disc flattened, with dense small punctures separated by about their own to twice their diameter, each puncture with microscopic seta, interspaces with alutaceous micro-sculpture; meso-metaventral suture distinct, straight, meso-metaventral sutural stria absent; intercoxal disc of metaventrite completely covered with punctation identical to that of mesoventrite, but interspaces without micro-sculpture. Lateral metaventral stria (Fig. 208) well impressed, shortened, carinate, curved outwardly; lateral disc of metaventrite (Fig. 208) excavated, with confluent shallow elongate punctures fringed with microscopic setae; metepisternum on apical half with deep longitudinal groove (Fig. 208) designed for accommodating mesotarsus, surface of metepisternum glabrous; fused metepimeron with scattered shallow punctures; lateral metepisternal stria present only on fused metepimeron, on metepisternum several large deep elongate punctures present.

Intercoxal disc of first abdominal ventrite completely striate laterally; surface of disc with punctation identical to that of mesoventrite, interspaces with alutaceous microsculpture.

Protibia (Fig. 209) flattened and dilated, outer margin almost without teeth, with 12–14 short thin denticles; setae of outer row short, sparsely paced; median row of setae absent; protarsal groove narrow but deep; anterior protibial stria absent; two thin, very short tarsal denticles present apically; protibial spur minute, growing out from apical protibial margin; outer and median part of posterior surface of protibia (Fig. 210) with several irregular rows of minute setae; demarcation line between outer and median part of posterior surface absent; posterior protibial stria complete, with regularly spaced minuscule setae turning into several thicker setae apically; inner margin with regular row of short setae becoming thicker apically.

Mesotibia (Fig. 211) dilated, outer margin with five widely spaced short denticles growing in size apically; setae of outer row sparse, regular, thin; setae of median row indistinguishable, entire posterior surface of protibia with several irregular rows of minuscule setae; posterior mesotibial stria inconspicuous (absent?); anterior surface of mesotibia with several irregular rows of minuscule setae; mesotibial spur stout, short; apical margin with two tiny denticles; claws of apical tarsomere about half its length; metatibia more slender than mesotibia, outer margin with only two short proximal denticles, in all other aspects similar to mesotibia.

Male genitalia. Eighth sternite (Figs 212–213) apically longitudinally separated medially; vela absent, eighth sternite apically with several setae; eighth tergite and eighth sternite not fused laterally (Fig. 214). Ninth tergite (Fig. 215) longitudinally fused medially; spiculum gastrale (Fig. 215) gradually dilated in most of apical half, basal half moderately dilated, rounded. Aedeagus (Figs 217–218) conspicuously long compared to eighth sternite and tergite, slender; basal piece of aedeagus short, ratio of its length : length of parameres 1 : 6; parameres fused along their basal three-fourths; aedeagus slightly curved from lateral view (Fig. 218).

Figures 212–218. 

212 Iridoprinus myrmecophilus sp. n. male terminalia: 8^th sternite + 8^th tergite, ventral view 213 ditto, dorsal view 214 ditto, lateral view 215 male terminalia: 9^th + 10^th tergites, dorsal view; spiculum gastrale, ventral view 216 male terminalia: 9^th + 10^th tergites; spiculum gastrale, lateral view 217 aedeagus, dorsal view 218 ditto, lateral view.

Body comparatively large; pronotum with bronze luster, elytra with blue metallic luster (old specimens can be dark brown to black without luster); labrum in males with large median projection (Fig. 229); labral pits with a single labral seta; dorsal elytral striae reduced, usually only fourth dorsal elytral stria discernible; supraorbital stria absent; frontal stria prolonged onto clypeus; both sets of prosternal striae absent; metepisternum comparatively widened; sixth abdominal ventrite of male with conspicuous semicircular median elevation; pygidium laterally carinate; metatibia in male more strongly curved than in female; remaining characters as in Saprinus.

Figure 219. 

Notosaprinus irinus (Marseul, 1862) habitus, dorsal view.

This is a typical saprobiont of the open landscapes and forest openings, mostly collected on carrion, where it presumably preys on fly larvae. Some specimens have been collected in pitfall and flight intercept traps.

Notosaprinus is a monotypic genus endemic to Australia: most records are from coastal areas of Queensland and New South Wales with a dubious record from Western Australia, see below (Fig. 757).

Notosaprinus is most similar to species of the genus Saprinus, differing from them chiefly by the absence of both sets of prosternal striae. The median elevation present on the sixth abdominal ventrite in males and sexually dimorphic labrum with single labral seta and absent supraorbital striae could be additional autapomorphies for this genus, however these features must be verified by extensive study of Saprinus species. In the phylogenetic study by Lackner (2014d), Notosaprinus irinus was recovered as sister taxon to Saprinus (Saprinus) semistriatus (Scriba, 1790), the type species of the species-rich genus Saprinus; albeit this relationship received only low support. The clade containing S. (S.) semistriatus and N. irinus was recovered sister to other higher taxa: type species of the genera Xerosaprinus Wenzel, 1962, Hemisaprinus Kryzhanovskij, 1976, Styphrus Motschulsky, 1845, Paraphilothis Vienna, 1994, and Pilisaprinus Kanaar, 1996. Apart from the absence of both sets of prosternal striae, Notosaprinus might be a member Saprinus, for it shares most of other synapomorphies with members of that genus. However, because there was only moderate statistical support for the phylogenetic placement, we maintain these taxa as separate genera.

Australia.

Saprinus irinus Marseul, 1862: Lectotype, present designation: ♀, (genitalia extracted and lost; specimen’s mounting card was pencil-marked with a female sign; probably not the original mounting card of Marseul), side-mounted, both protarsi, right mesotarsus, left metatibia and right metatarsus missing, with the following labels: “22a / Saprinus irinus m. / N. Holl. / T. Dré” (pink, round label, written); followed by one more identical label; followed by: “MUSEUM PARIS / irinus / COLL. / De MARSEUL 1890” (pink label, printed-written); followed by: “TYPE” (red-printed rectangular label); followed by: “Saprinus irinus / Marseul, 1862 / LECTOTYPE / des. T. Lackner 2014” (red label, written) (MNHN). This species has been described from an unknown number of specimens and the lectotype designation fixes the identity of the species.

AUSTRALIA. New South Wales: 3 ♀♀, Richmond River, 1909, collector unknown (BMNH); 2 ♂♂, Richmond River, without further data, Coll Van de Poll (MNHN); 1 ♀, Gerroa, 29.xii.1974, H. & A. Howden (MNHN); 1 ♀, Sydney, without further data (MNHN); 2 specs., N.S. Wales, without further data (MNHN); 1 spec., Chichester S.F., Allyn River, 32.08S 151.27E, T. Weir leg. (ANIC); 3 ♀♀ & 2 ♂♂, Lots, Caparra, 14.iii.1993, S. Watkins leg., under dead cat (ANIC); 1 spec., Mosgeil, without further data (MAMU); 1 spec, Route: Sydney to Mt. Abundance, thence to Maranoa and the Warrego; from there to Nivelle river, back to the Warrego (S., Centr. and W. Queensland, Hely’s Expedition 1852) (MAMU); 1 ♀ & 1 spec., Sydney, without further data, Griffith (SAMA). Queensland: 1 spec., Cairns, 7.ii.1927, no collector (BPBM); 1 spec., Queensland, Challenger exp., no further data (BMNH); 3 ♀♀, Mackay, Queensland, Turner, no further data (BMNH); 3 specs., Ewan Road, 8 miles W of Poluna, 18.i.1970, J.G. Brooks leg., fish lure (ANIC); 1 spec., Atherton, 26.iii.1965, Bornemisza leg. (ANIC); 1 spec., Thorton Range, Daintree, 12.–18.vii.1982, S. & J. Peck leg. (ANIC); 2 specs., Cape Tribulation, 40 km N of Daintree, 10 m, 18.vii.1982, S. & J. Peck leg. (ANIC); 2 specs., Kuranda St. For, 360 m, 27.–31.vii.1982, S. & J. Peck leg. (ANIC); 3 specs., Mount Mee, 7.5 km SW, 27°05'S, 152°42'E, 12.x.1991, Tom Gush leg., under dead snake on the road (ANIC); 4 ♂♂ & 7 ♀♀, Mt. Gavial, 3 km SSW, 18.xii.–14.iii.1999, 23°37'S, 150°28'E, D.J. Cook leg., vine forest 320 m, intercept trap (QM); 1 ♀, Tallebudgera Valley, 420 m, 4.–28.iv.1997, 153°18'E 28°14'S, D. Cook leg., carrion baited pitfall (QM); 2 specs., Brisbane valley, near Ravensbourne Nat. Park, 25.xi.1973, K. McDonald leg. (QM); 3 ♀♀, 2 ♂♂ and 3 specs., Baily Knob summit, 6.xii.1998–6.ii.1999, 17°39'S, 145°30'E, 1100 m, Monteith & Cook leg., intercept (QM); 3 specs., McAfee’s Lookout, 27°26'S, 152°53'E, 300 m, 18.x.1999, G.B. Monteith, under dead goanna (QM); 7 specs., Fraser Island, Yidney Scrub nr. VB52, 1.–2.xii.1975, G. Thompson & A. Slater (QM); 1 spec., Russel R., at Bellenden, Ker Landing, 5 m, 24.x.–9.xi.1981, Baited window trap (QM); 1 spec., Mt. Tamborine, 27.x.1957, S. Breden leg. (QM); 1 spec., Kirrama Range, Douglas Creek Road, 800 m, 10.xii.1986–11.i.1987, Monteith, Thompson & Hamlet leg., flight intercept trap (QM); 1 spec., Kroombit Tops, 65 km SW from Gladstone, 1000–1100 m, 22.–26.ii.1982, open forest, Monteith, Thompson & Yeates leg. (QM); 1 ♀, Mt. Blackwood, 590 m, 21°02'S, 148°57'E, 18.xi.1992–mid iv.1993, D. Cook & G.B. Monteith leg., intercept traps and pitfalls (QM); 1 spec., Maalan Road, 2 km S of Palmerston Highway, 17°36'S, 145°42'E, 7.iii.–15.v.1995, 750 m, Monteith & Hasenpusch leg., intercept trap (QM); 1 spec., Stone Creek, 17°28'S, 146°01'E, 100 m, 1.xi.1995–6.ii.1996, intercept trap, ground level (QM); 11 specs., Mt. Tambourine, xii. 1919, H. Pottinger leg. (QM); 1 spec., Gayndah, Masters (SAMA). Western Australia: 1 spec., K.G. Sound [probably King’s Sound; due to the uncertainty of exact locality not shown on the map] (MAMU).

Unknown localities: 2 ♀♀ & 1 spec., Australia, without further data (BMNH).

A common species found on carrion and often collected by flight intercept traps.

Australia: Coastal regions of Queensland and New South Wales, the single record from Western Australia, is, due to the uncertain locality not shown on the map (Fig. 757).

This species is sexually dimorphic: males have a conspicuous median projection on the labrum, strongly curved metatibia, and sixth abdominal ventrite has a median elevation.

Body length: PEL: 4.25–6.20 mm; APW: 1.50–2.25 mm; PPW: 3.25–4.75 mm; EL: 2.75–3.85 mm; EW: 3.75–5.00 mm.

Body (Fig. 219) rectangular oval, convex; cuticle shining; pronotum dark brown with bronze metallic luster, elytra with blue metallic luster; legs, mouthparts and antennae castaneous brown; antennal club black.

Antennal scape (Fig. 220) slightly thickened, imbricate-punctate, with few short setae; antennal club (Fig. 221) round, depressed dorso-ventrally, without visible articulation, entire surface with dense short sensilla intermingled with sparse longer erect sensilla; sensory structures of antennal club (Fig. 230) in form of four rather large sensory areas on ventral side and one vesicle situated under internal distal margin.

Figures 220–227. 

220 Notosaprinus irinus (Marseul, 1862) head, dorsal view 221 antennal club, dorsal view 222 mentum, ventral view 223 propygidium + pygidium 224 prosternum 225 lateral disc of metaventrite + metepisternum 226 protibia, dorsal view 227 ditto, ventral view.

Mandibles (Fig. 228) punctate, with rounded outer margin, curved inwardly, mandibular apex acutely pointed; sub-apical tooth on inner margin of left mandible small, obtuse; labrum (Fig. 229) convex, densely punctate, depressed medially, in males with large median projection, in females without such projection, with two shallow labral pits, each with single short labral seta; labral fold inconspicuous; terminal labial palpomere elongated, its width about one-third its length; palpal organ minute, but present on both labial and maxillary palpi; mentum (Fig. 222) sub-trapezoid, anterior margin medially with deep notch (Fig. 222) surrounded with two long setae, lateral margins with several shorter setae; disc of mentum imbricate, with several sparse microscopic setae; cardo of maxilla on lateral margin with few short setae; stipes triangular, medially with single long seta, another long seta present laterally; lacinia without lacinial hook (=uncus); terminal maxillary palpomere elongated, its width about one-fourth its length, approximately twice as long as penultimate.

Clypeus (Fig. 220) punctate, narrowed anteriorly, depressed laterally; frontal stria widely interrupted medially (Fig. 220), prolonged onto clypeus, supraorbital stria absent; frontal disc (Fig. 220) with coarse and dense punctures; two vague depressions present on posterior half; eyes large, convex, well visible from above.

Pronotal sides (Fig. 219) moderately narrowing anteriorly, apical angles obtuse, pronotal depressions deep, rather large; marginal pronotal stria complete, carinate; disc of pronotum laterally with a band of coarse and dense punctation, not reaching posterior corners, between it and pronotal margin thin impunctate band present; pronotum medially almost smooth, with scattered microscopic punctation; along pronotal base two to three rows of coarse punctation present; pronotal hypomeron glabrous; scutellum small, well visible.

Elytral humeri slightly prominent, impunctate; elytral epipleura with scattered fine punctures; marginal epipleural stria fine, complete; marginal elytral stria nearly straight, in deep round punctures, continued as complete apical elytral stria. Humeral elytral stria vaguely impressed on basal third; outer subhumeral stria well impressed, thin; inner subhumeral stria erased by coarse elongate wrinkles; dorsal elytral striae 1–3 completely obliterated by coarse longitudinal wrinkles, fourth dorsal elytral stria deeply impressed, straight, shortened apically, present only on basal third of elytral length; sutural elytral stria shortened on basal sixth, well-impressed, apically connected with apical elytral stria. Elytral disc between elytral humerus and fourth dorsal elytral stria along elytral base with very coarse and dense elongate wrinkles reaching apically approximately one-third of elytral length, surface between this coarse band and fourth dorsal elytral stria creating a glabrous band, only with vague row of sparse punctures; surface between fourth dorsal elytral stria and elytral suture on basal third creating a glabrous triangular ‘mirror’ (= polished area); elytral surface otherwise with deep coarse and dense punctures becoming somewhat sparser and finer apically and towards elytral suture; elytral flanks glabrous.

Propygidium (Fig. 223) completely exposed, laterally with depressions, punctate, punctures separated by up to twice their diameter, interspaces finely imbricate; pygidium (Fig. 223) longer than broad along median line; laterally with moderately dense punctures becoming sparser medially, interspaces finely imbricate; carinate laterally.

Anterior margin of median portion of prosternum (Fig. 224) straight; marginal prosternal stria present only laterally; prosternal process flattened, glabrous, antero-laterally large coarse punctures present; both sets of prosternal striae absent.

Anterior margin of mesoventrite shallowly emarginate medially; discal marginal mesoventral stria well impressed, complete; disc of mesoventrite smooth; meso-metaventral sutural stria absent; meso-metaventral suture well impressed, thin; intercoxal disc of metaventrite glabrous, longitudinal suture of metaventrite deeply impressed, area along lateral metaventral stria and posterior margin with fine scattered punctation; along basal margin of metaventrite a band of coarser and denser punctures present; lateral metaventral stria shortened, straight; lateral disc of metaventrite (Fig. 225) slightly concave, with dense large setigerous punctures, interspaces finely substrigulate; metepisternum + fused metepimeron (Fig. 225) comparatively widened, with sparser and finer punctures, punctures not setigerous, interspaces substrigulate; on fused metepimeron punctation somewhat sparser, interspaces glabrous; lateral metepisternal stria complete, deeply impressed.

Intercoxal disc of the first abdominal ventrite almost completely striate laterally; disc with fine scattered punctation; sixth abdominal ventrite in males with distinctive median semicircular projection.

Protibia (Fig. 226) slightly dilated, outer margin with about three large distal teeth topped by stout triangular denticle, followed by four–five much smaller denticles diminishing in size in proximal direction; setae of outer row regular, short; protarsal groove deep; anterior tibial stria shortened apically; setae of median row much shorter and sparser than those of outer row; two apical denticles present near tarsal insertion; protibial spur well developed, bent, growing out from apical margin of protibia; apical margin of protibia ventrally with a single short denticle (occasionally there can be more than one denticle present); outer part of posterior surface (Fig. 227) of protibia smooth, only with several scattered shallow punctures, separated from glabrous median part by vague boundary and an undulate row of short sclerotized setae; posterior protibial stria complete, with moderately dense row of short, well-sclerotized setae; inner margin of protibia with double row of well-sclerotized lamellate setae growing in size and girth apically.

Mesotibia slender, outer margin with two rows of short denticles growing in size apically; setae of outer row regular, dense, about as long as denticles themselves; setae of median row shorter, regular; posterior mesotibial stria shortened apically; anterior surface of mesotibia almost smooth, with scattered shallow punctures; anterior tibial stria complete, terminating in two tiny inner ventral denticles; mesotibial spur rather long and thick; claws of apical tarsomere bent, longer than half its length, each mesotarsomere ventrally with two long, well sclerotized setae; metatibia in males more curved than in females, slenderer and longer than mesotibia, in all aspects similar to it, but denticles on outer margin much sparser.

Male genitalia. Eighth sternite (Figs 231–232) fused longitudinally; vela with single row of sparse short setae; apex of eighth sternite laterally asetose; eighth tergite and eighth sternite not fused laterally (Fig. 233). Tenth tergite (Fig. 234) comparatively large, elongate; ninth tergite (Figs 234–235) longitudinally fused medially; spiculum gastrale (Figs 236–237) abruptly largely expanded on apical third, basally only slightly expanded, triangular. Aedeagus (Figs 238–239) thickened, constricted before apex; basal piece of aedeagus short, ratio of its length : length of parameres 1 : 4; parameres of aedeagus fused almost along their entire length with only a small apical circular opening for median duct; aedeagus only slightly curved from lateral view (Fig. 239).

Figures 228–230. 

228 Notosaprinus irinus (Marseul, 1862) mandibles, dorsal view 229 male labrum: left half depicting dorsal view and right half depicting epipharynx 230 antennal club, ventral view showing sensory structures of the antenna.

Figures 231–239. 

231 Notosaprinus irinus (Marseul, 1862) male terminalia: 8^th sternite + 8^th tergite, ventral view 232 ditto, dorsal view 233 ditto, lateral view 234 male terminalia: 9^th + 10^th tergites, dorsal view 235 ditto, lateral view 236 male terminalia: spiculum gastrale, ventral view 237 ditto, lateral view 238 aedeagus, dorsal view 239 ditto, lateral view.

Cuticle light to dark brown, without metallic luster, entire dorsal surface glabrous; mandibles massive, strongly carinate dorsally; clypeus large, triangular and strongly convex; supraorbital and frontal striae absent; pronotal depressions absent; pronotal hypomeron with long yellow setae; prosternal foveae absent; prosternal apophysis strongly constricted between procoxae, prosternal process thence strongly expanded; both sets of prosternal striae absent, prosternal process setose; mesoventrite constricted between mesocoxae; all femora, meso- and metatibiae strongly swollen; protibia with a dense row of long thin denticles on outer margin; meso- and metatibiae with rows of setigerous punctures.

Figure 240. 

Reichardtia pedator (Sharp, 1876) habitus, dorsal view.

This is a monotypic psammophilous taxon found on the beach under carrion or kelp at depths of 20 cm or more, occasionally also collected walking on sand surface.

Reichardtia is endemic to New Zealand and is found on both North and South Islands, but absent from off-shore islands (Fig. 758).

Based on the characters outlined above, especially the leg morphology, it is impossible to confuse Reichardtia pedator with any other taxon in the region. This New Zealand monotypic endemic is characterized by numerous autapomorphies, including almost impunctate dorsal surface in combination with the absence of supraorbital and frontal striae and a setose pronotal hypomeron. It is a rather derived member of the subfamily (Lackner 2014d), belonging to a global psammophilous clade. In the analysis of the senior author (loc. cit.) Reichardtia was recovered inside a purported monophyletic group Australopachylopus lepidulus (Reichardtia pedator + Reichardtiolus pavlovskii). However, as already noted by Leschen and Ôhara (2017) this relationship will probably not hold true as the characters uniting the triad may be a result of convergence, rather than synapomorphy.

New Zealand: Auckland.

Saprinus pedator Sharp, 1876: Lectotype, present designation: 1 spec., with “Saprinus / pedator / N. Zeal. Type / D.S.” written on the actual mounting card with the specimen, followed by: “Auckland / New Zealand” (printed); followed by: “Sharp Coll. / 1905-313”; followed by: “Type” (red-margined printed round label); followed by: “Saprinus pedator / Sharp, 1876 / LECTOTYPE 2014 / Des. Lackner & Leschen” (red label, written) (BMNH). Paralectotypes, present designation: 1 spec., “Saprinus pedator / D.S. / Otago Hulton 1878” (written on the actual mounting card), followed by: “Sharp Coll. / 1905-313”; followed by: “Saprinus pedator / Sharp, 1876 / PARALECTOTYPE 2014 / Des. Lackner & Leschen” (red label, written) (BMNH). 1 spec., “Saprinus pedator / Sharp / Ind. Typ. / N. Zeald D.S.” (written on the actual mounting card); followed by: “Auckland” (red label, written); followed by: “Auckland / New Zealand” (printed); followed by: “Saprinus pedator / Sharp, 1876 / PARALECTOTYPE 2014 / Des. Lackner & Leschen” (red label, written) (BMNH). 1 spec., “Saprinus pedator / Sharp. / Otago Hulton 1878” (written on the actual mounting card); followed by: “Sharp Coll. / 1905-313”; followed by: “Saprinus pedator / Sharp, 1876 / PARALECTOTYPE 2014 / Des. Lackner & Leschen” (red label, written) (BMNH). 1 spec., with the following labels: “Saprinus / pedator / N. Zeald. Ind. Typ. / D.S.” (written); followed by: “G. Lewis Coll. / B.M. 1926-369” (printed); followed by: “Saprinus pedator / Sharp, 1876 / PARALECTOTYPE 2014 / Des. Lackner & Leschen” (red label, written) (BMNH). Sharp (1876) does not mention the exact number of specimens in his original description and the lectotype designation serves to fix the identity of the species.

Figures 241–252. 

241 Reichardtia pedator (Sharp, 1876) head, dorsal view 242 antennal club, ventral view 243 mentum, ventral view 244 propygidium + pygidium 245 prosternum 246 mesoventrite 247 lateral disc of metaventrite + metepisternum 248 protibia, dorsal view 249 ditto, ventral view 250 mesotibia, dorsal view 251 metatibia, dorsal view 252 ditto, ventral view.

NEW ZEALAND. North Island: ND: 1 spec., Waipapakauri, 7.xi.1991, G. Kuschel leg. (NZAC); 1 ♀, Cape Maria, 20.i.1963, L. Price leg. (NZAC). AK: 1 spec., Ruakaka Beach, N. Auckland, 22.ix.[19]32, C.E. Clarke coll. (BMNH); 1 spec., Muriwri Beach, 12.ix.[19]32, C.E. Clarke coll. (BMNH). TK: 1 spec., Taranaki, New Zealand, Broun Coll. (BMNH). WI: 1 spec., Himatangi Beach, 20.iii.1988, M.O.E. Peters leg. (AMNZ). WN: 1 spec., New Zealand, Parapararaumu, 17.ii.[19]24, O’Connor, G.V. Hudson, in dead fish on beach (BMNH); 1 spec., Wellington, ii.1902, J.J. Walker (BMNH); 1 spec., Manawatu Estuary, 25.iii.1984 (CJN); 1 spec., Waikawa Beach, 2.iv.1995 (CJN); 1 spec., Waikanae Beach, 9.ii.1995 (CJN). South Island. NN: 3 specs., Farewell Spit, Outer beach, 9.ii.1981, J.W. Early leg. (LUNZ); 7 specs., Wharariki Beach, 7.ii.1981, J.W. Early leg. (LUNZ, 3 exs. in coll. TLAN); 1 ♂, Tahunanui, Beach Nelson 21.viii.1976, J.C. Watt leg. (NZAC). SC: 1 spec., Temuka Beach, 22.iii.2008 (CJN). SL: 4 specs., South Island, Curio Bay, 50 km E of Invercargill, sandy beach, 4.xii.2000, Erber (ZMHUB).

DN: 2 specs., Kuri Bush, near Taieri Mouth, 3.xi.2001 (CJN); 2 specs., Sandfly Bay, Otago Peninsula, 18.i.2004 (CJN); 1 spec., Smails beach, Dunedin, 16.iii.1999 (CJN); 1 spec., Boulder Beach, Otago Peninsula, 5.i.2005 (CJN); ditto, but 29.xii.2001 (CJN); 1 spec., Waikouati, 14.i.2002 (CJN). Unknown localities: 1 spec., N. Zealand, no further data, G. Lewis Coll. (BMNH); 1 spec., New Zealand, no further data, Broun Coll. (BMNH); 1 spec; New Zealand, 25.ii.1917, Broun Coll. (BMNH).

Littoral species found on beach under kelp, dead fish, often buried in the sand.

Endemic to New Zealand (Fig. 758).

Body length: PEL: 3.25–4.25 mm; APW: 1.25–1.60 mm; PPW: 2.35–3.00 mm; EL: 2.15–2.90 mm; EW: 2.60–3.35 mm.

Body (Fig. 240) rectangular oval, elytral humeri prominent, strongly convex, cuticle light to dark brown, shining, without metallic luster, entirely glabrous; legs, antennae and maxillary palpi similarly colored.

Antennal scape (Fig. 241) slightly thickened, with numerous long setae; antennal club (Fig. 242) comparatively small, rounded, entirely covered in dense short sensilla, intermingled with sparse longer erect sensilla; sensory structures of antennal club (Fig. 255) in form of a single ventral sensory area with pear-shaped vesicle situated beneath it.

Mandibles (Fig. 253) massively thickened with rounded outer margin, strongly carinate dorsally, bluntly pointed, sub-apical tooth on inner margin of left mandible large, blunt; labrum (Fig. 254) faintly convex dorsally, sparsely microscopically punctate, anterior margin slightly emarginate; labral fold tiny, not conspicuous; setae of lateral fringe conspicuously short; labrum with two labral setae; terminal labial palpomere elongated, its width about one-third its length; palpal organ present on both labial and maxillary palpi; mentum (Fig. 243) sub-trapezoid, anterior angles slightly produced, anterior margin with shallow median excavation and tiny median notch (Fig. 243), surface around it with four long setae, lateral margins with several microscopic setae, surface of mentum imbricate; cardo of maxilla with few short setae on lateral margin; stipes triangular, with four long setae; lacinia without lacinial hook (=uncus); terminal maxillary palpomere elongated, its width about one-fourth its length, about three times as long as penultimate palpomere.

Figures 253–255. 

253 Reichardtia pedator (Sharp, 1876) mandibles, dorsal view 254 labrum: left half depicting dorsal view and right half depicting epipharynx 255 antennal club, ventral view showing sensory structures of the antenna.

Clypeus (Fig. 241) large, rectangular, strongly convex, sloping down anteriorly, rounded laterally, glabrous, surface between clypeus and frontal disc strongly concave; supraorbital and frontal striae absent; frontal disc (Fig. 241) about twice as broad as long, with conspicuous median fovea; eyes slightly flattened, but visible from above.

Pronotal sides (Fig. 240) rounded, apical angles blunt, marginal pronotal stria complete, carinate, somewhat weakened behind head; disc strongly convex, glabrous; pronotal hypomeron with long yellow setae; scutellum small, but visible.

Elytral epipleura punctate; marginal epipleural stria thin, faintly impressed; marginal elytral stria well impressed, continuous along elytral apex as apical elytral stria, connected with sutural elytral stria; humeral elytral stria well impressed, occasionally continuous with inner subhumeral stria forming a stria parallel to first dorsal elytral; dorsal elytral striae deeply impressed, first the longest, reaching approximately half of elytral length apically, striae 2–4 shorter, reaching about one-third of elytral length apically; sutural elytral stria complete, deeply impressed, continuous with apical as well as 4^th dorsal elytral striae. Except for scattered punctures near elytral apex entire elytral disc glabrous.

Propygidium (Fig. 244) transverse, about three times as broad as long, completely exposed, sparsely punctate, punctures separated by several times their diameter, interspaces between punctures with fine alutaceous microsculpture; pygidium (Fig. 244) almost smooth, only on basal third with several scattered shallow almost inconspicuous punctures.

Anterior margin of median portion of prosternum (Fig. 245) bisinuate, setose, anterior angles somewhat produced; marginal prosternal stria absent; prosternal apophysis strongly constricted between procoxae, prosternal process knife-like, strongly widening and sloping down anteriorly, surface imbricate, with scattered punctures fringed with setae.

Anterior margin of mesoventrite (Fig. 246) straight; discal marginal mesoventral stria well impressed, fringed with several short setae; disc convex, only with microscopic punctation, strongly constricted between mesocoxae; meso-metaventral suture deeply impressed, meso-metaventral sutural stria well impressed, overriding meso-metaventral suture; intercoxal disc of metaventrite broad, smooth, in males with concave median longitudinal depression. Lateral metaventral stria well impressed, strongly curved outwardly; lateral disc of metaventrite (Fig. 247) convex, with round shallow large setigerous punctures; metepisternum + fused metepimeron (Fig. 247) with even denser setigerous punctures than those of lateral disc of metaventrite, punctures almost disappear on fused metepimeron; lateral metepisternal stria intermittent, present only on metepisternum.

Intercoxal disc of first abdominal ventrite in males with large depression medially, in females depression much smaller, lateral striae shortened apically; glabrous.

Protibia (Fig. 248) widening apically, outer margin with a row of about 17 densely-set straight long lamellate denticles growing in size apically; setae of outer row moderately dense, long and regular; setae of median row approximate to outer row, but shorter and sparser; protarsal groove deep; anterior protibial stria complete, carinate, next to it another row of much shorter sparse setae present, apically setae growing in length; protibial spur short, hooked, growing out near tarsal insertion; outer part of posterior surface of protibia (Fig. 249) with dense moderately long strongly sclerotized setae, demarcation line between outer, median and inner part of posterior surface of protibia absent; inner margin with double row of sparse sclerotized setae growing in size and becoming denser apically.

Mesotibia (Fig. 250) strongly thickened, outer margin with a dense row of thick denticles growing in size apically; setae of outer row strongly sclerotized, dense, growing in size but becoming thinner apically; setae of median row rather distanced from outer row, thinner and sparser; between the two rows several additional setae present; posterior mesotibial stria inconspicuous (absent?); anterior surface of mesotibia convex, with several dense rows of short denticles, becoming sparser and thinner towards inner mesotibial margin; anterior mesotibial stria inconspicuous (absent?); mesotibial spur inconspicuous; first and second tarsomeres ventrally with four long strongly sclerotized setae; third and fourth tarsomeres with only two such setae; fifth tarsomere devoid of setae ventrally; first two tarsomeres dorsally with two strongly sclerotized setae; third, fourth and fifth tarsomeres dorsally with only single seta; claws of apical tarsomere bent, shorter than half its length; metatibia (Figs 251–252) basically similar to mesotibia, but even more thickened, denticles of outer margin and anterior surface much denser and stouter than those of mesotibia.

Male genitalia. Eighth sternite (Figs 256–257) fused longitudinally; vela absent; apex of eighth sternite laterally with several short setae; eighth tergite and eighth sternite not fused laterally (Fig. 258). Ninth tergite (Figs 259–260) longitudinally fused medially; spiculum gastrale (Fig. 259) gradually dilated on apical half, basal end strongly dilated, resembling flukes of a whale. Aedeagus (Figs 261–262) slightly thickened, constricted before apex; basal piece of aedeagus very short, ratio of its length : length of parameres 1: 7; parameres of aedeagus fused almost along their basal two-thirds; aedeagus only slightly curved from lateral view, apex of aedeagus flattened dorso-ventrally.

Figures 256–262. 

256 Reichardtia pedator (Sharp, 1876) male terminalia: 8^th sternite + 8^th tergite, ventral view 257 ditto, dorsal view 258 ditto, lateral view 259 male terminalia: 9^th + 10^th tergites, dorsal view; spiculum gastrale, ventral view 260 male terminalia: 9^th + 10^th tergites; spiculum gastrale, lateral view 261 male terminalia: aedeagus, dorsal view 262 ditto, lateral view.

Cuticle light to dark brown with faint bronze metallic tinge, entire dorsal surface (with the exception of vaguely delimited ‘mirrors’ on pronotal disc and elytra) rugulose-lacunose; labral pits and setae absent; pronotal depressions absent; prosternal foveae absent; bases of lateral prosternal striae with distinctive projection; protibia without teeth or denticles, very slender and elongate; protarsal groove very deep; protibial spur large; spiculum gastrale not expanded basally.

Figure 263. 

Saprinodes distinctus Dégallier, 1993, habitus, dorsal view.

Unknown, most of the specimens have been collected using pitfall traps.

Endemic to Australia: Queensland and New South Wales (Fig. 759).

Saprinodes is the only Australopacific saprinine with the protibia devoid of teeth or denticles on its outer margin. Furthermore, its protibia is apically narrowed, terminating in a large protibial tooth. The peculiarly-shaped protibia of Saprinodes is not found in any other currently known Saprininae genus. The absence of labral pits and setae and the distinctive prosternal projection near the bases of lateral prosternal striae are also autapomorphies. In the morphology-based performed phylogenetic analysis published by the senior author (Lackner 2014d), the type species of the genus, S. falcifer Lewis, 1891 was recovered sister to Iridoprinus myrmecophilus with one ‘strong’ and two ‘weaker’ synapomorphies supporting their relationship (see also remarks section of Iridoprinus).

Australia: Queensland: Danbulla S.F.

Saprinodes distinctus Dégallier, 1993: holotype, ♂, genitalia glued to the same mounting card as the specimen, with the following labels: “AUSTRALIA: n. Qld. / Danbulla S.F., 13 km / NE of Yungaburra / 28.VII - 3.IX. 1987 / Storey & De Faveri” (printed-written); followed by: “MDPI Intercept / Trap Site No. 27” (printed-written); followed by: “Saprinodes / distinctus / HOLOTYPE / N. DEGALLIER / T. 13244” (red label, written) (QM). Allotype, ♀, ditto, but 20.XII.[19]86 - 13.I.[19]87, and Trap Site No. 21 (QM). Paratype, ♀, with genitalia glued to the same mounting card as the specimen, with the following labels: “The Crater, near / Herberton, N. Qld. / 16 dec. 1961 / McAlpine + Lossin” (written); followed by: “Saprinodes / distinctus / DEGALLIER / PARATYPE” (red label, written); followed by: “K 204388” (printed); followed by: “10-114” (yellow label, pencil-written, added by the senior author) (AMS).

AUSTRALIA. Queensland: 1 spec., Wongabel S.F., 5 km S Atherton, 800 m, 5.–14.xii.1988, Monteith & Thompson (FIT) (QM); 1 ♂ & 2 specs., Batavia Downs, 12.41S 142.41E, 22.vi.–23.viii.1992, P. Zborowski & J. Cardale (FIT) (ANIC).

Unknown.

Endemic to Australia: Queensland (Fig. 759).

S. distinctus is rather similar to S. falcifer. Characters that best separate the two species are mentioned in the above key. The males of the two species differ only in the structure of eighth abdominal ventrite (compare Figs 270–271 with 292–293). Since we provide the type species of the genus, S. falcifer, with a full re-description we give for S. distinctus only a brief diagnostic description.

Body length: PEL: 2.50–2.80 mm; APW: 0.75–0.85 mm; PPW: 1.75–2.00 mm; EL: 1.50–1.80 mm; EW: 2.00–2.25 mm. Body (Fig. 263) generally similar to S. falcifer, but elytral ‘mirror’ (=polished area) slightly larger, punctures slightly larger than in S. falcifer (compare Figs 263 and 279); legs, antennae and mouthparts similarly coloured between the two species. Antennal scape (Fig. 264) as well as antennal club (Fig. 265) almost identical to S. falcifer; sensory structures of antennal club not examined. Mandibles with rounded outer margin, acutely pointed, sub-apical tooth on inner margin of left mandible moderately large, rounded; labrum faintly convex dorsally, knobby, otherwise completely agrees with that of S. falcifer; mentum (Fig. 266) very similar to that of S. falcifer, but anterior margin slightly bisinuate and median emargination less deep than in following species (compare Figs 266 and 282); other mouthparts not examined.

Figures 264–269. 

264 Saprinodes distinctus Dégallier, 1993 head, dorsal view 265 antennal club, ventral view 266 mentum, ventral view 267 prosternum 268 mesoventrite 269 protibia, dorsal view.

Clypeus (Fig. 264) also similar between two species, but more constricted laterally (compare Figs 264 and 280), and more coarsely punctuate, margined laterally by prolonged divided frontal striae; supra-orbital striae well developed, carinate; frontal stria (Fig. 264) widely separated medially, continuous even further on clypeus than in S. falcifer; frontal disc (Fig. 264) more coarsely punctuate than that of S. falcifer, otherwise very similar to it; eyes convex, well visible from above.

Pronotal sides (Fig. 263) strongly convergent apically, stronger so than in following species, rest of pronotum extremely similar to S. falcifer with exception of small angulate projection of frontal stria medially behind head which is absent in S. distinctus and present in S. falcifer; scutellum very small. Elytral striae striae 1–4 faintly recognizable. Entire elytral disc very coarsely and densely punctuate, punctation rugulose-lacunose, even coarser than that of S. falcifer, punctures on apical half confluent, aciculate and striolate (Fig. 263) on apical third punctures disappear, forming deep elongate wrinkles; basally between fourth dorsal elytral and sutural stria a small (however, slightly larger than that of S. falcifer) round ‘mirror’ (=polished area) present, its surface glabrous. Propygidium and pygidium very similar between the two species. Prosterna of the two species almost identical, curious projection near basal ends of lateral prosternal striae of S. distinctus (Fig. 267) even more prominent than in following species (Fig. 284). Mesoventrites (Fig. 268) as well as metaventrites of the two closely related species very similar; metepisterna also very similar. Intercoxal disc of first abdominal ventrite very similar in both species. Legs of both species (compare e.g. Figs 269 and 287) almost identical. Male genitalia (Figs 270–278) of S. distinctus differ basically only in the structure of eighth abdominal ventrite that is in S. distinctus fused medially (separated in S. falcifer) and vela that is in the case of S. distinctus almost asetose; apices of eighth sternite are with few setae in S. distinctus whereas they are without setae in S. falcifer (compare Figs 270–271 with Figs 292–293). Eighth abdominal tergite is medially deeply inwardly arcuate in S. distinctus whereas it is less so in S. falcifer.

Figures 270–278. 

270 Saprinodes distinctus Dégallier, 1993 male terminalia: 8^th sternite + 8^th tergite, ventral view 271 ditto, dorsal view 272 ditto, lateral view 273 male terminalia: 9^th + 10^th tergites, dorsal view 274 ditto, lateral view 275 male terminalia: spiculum gastrale, ventral view 276 ditto, lateral view 277 male terminalia: aedeagus, dorsal view 278 ditto, lateral view.

Australia: Queensland: Rockhampton.

Figure 279. 

Saprinodes falcifer Lewis, 1891 habitus, dorsal view.

Saprinodes falcifer Lewis, 1891: lectotype, designated by Dégallier in 1993, ♂, with genitalia glued to the same mounting card as the specimen, labelled: “Rockhampton/Queensland” (hand-written); followed by: “Saprinodes/falcifer/Type Lewis” (hand-written); followed by: “G. Lewis Coll./B.M. 1926-369.” (printed); followed by: “Type” (round label with red margins, printed); and a consecutive red label: “Saprinodes/falcifer/LECTOTYPE” (hand-written); followed by a yellow label: “10-120” (pencil-written, added by the senior author) (BMNH).

AUSTRALIA. Queensland: 11 specs., 25°27'S, 150°08'E, Taroom District, Nathan Gorge, Riverine Forest, 12.ix.–13.xi.1996, P. Lawless leg., pitfall (QM; 2 exs. in coll. TLAN). New South Wales: 2 ♂♂, Bogan River, F.H. Taylor leg. (ANIC); 7 ♂♂ & 3 ♀♀, ditto, but J. Armstrong (all exs. ANIC; 1 ♂ in coll. TLAN); 2 ♂♂ and 7 specs., ditto, but J. Armstrong leg. (MAMU).

Unknown.

Endemic to Australia: New South Wales and Queensland (Fig. 759).

In this sexually dimorphic species the male has a large depression on the metaventrite and the first abdominal ventrite has coarser and larger punctures than that of the female. Females are also substantially larger than males. The minute setae associated with dorsal punctures observed by Dégallier (1993: 48) were not observed in our specimens.

Body length: PEL: 2.50–3.75 mm; APW: 0.85–1.05 mm; PPW: 1.80–2.55 mm; EL: 1.55–2.20 mm; EW: 2.10–3.00 mm.

Body (Fig. 279) ovoid, flattened dorso-ventrally, cuticle light brown to castaneous, with slight metallic tinge; ventral side darker than dorsal; legs and body appendages similarly colored.

Antennal scape (Fig. 280) slightly thickened, punctuate dorsally, with few short setae; antennal club (Fig. 281) rather large, oval, dorsally without any visible structures, ventrally with two visible oval sensory areas, entirely covered in dense short sensilla, intermingled with sparse longer erect sensilla; sensory structures of antennal club (Fig. 291) in form of two ventral sensory areas, apical sensory area larger than basal, with pear-shaped vesicle situated beneath it.

Figures 280–288. 

280 Saprinodes falcifer Lewis, 1891 head, dorsal view 281 antennal club, ventral view 282 mentum, ventral view 283 pygidium 284 prosternum 285 mesoventrite 286 lateral disc of metaventrite + metepisternum 287 protibia, dorsal view 288 ditto, ventral view.

Mandibles (Fig. 289) with rounded outer margin, acutely pointed, sub-apical tooth on inner margin of left mandible moderately large, rounded; labrum (Fig. 290) faintly convex dorsally, sparsely punctuate, anterior margin slightly emarginate; labral fold not particularly developed; setae of lateral fringe moderately long; labrum without pits or setae; terminal labial palpomere elongated, its width about half its length; palpal organ present on both labial and maxillary palpi; mentum (Fig. 282) sub-trapezoid, anterior angles produced, anterior margin with deep median excavation, surface around it with several short setae, lateral margins with a single row of much shorter sparse ramose setae, disc on apical half with few scattered setae, basal half asetose; cardo of maxilla with few short setae on lateral margin; stipes triangular, with two short setae; lacinia without lacinial hook (=uncus); terminal maxillary palpomere elongated, its width about half its length, about twice as long as penultimate palpomere.

Clypeus (Fig. 280) large, rectangular, flattened, rounded laterally, with sparse fine punctures, separated by several times their diameter; supra-orbital striae well developed, carinate; frontal stria (Fig. 280) prolonged onto clypeus; frontal disc (Fig. 280) anteriorly somewhat depressed, punctuate, punctures separated by about their diameter, becoming coarser and denser near frontal stria, disappearing medially; eyes convex, well visible from above.

Pronotal sides (Fig. 279) on posterior half feebly convergent anteriorly, on anterior half strongly convergent, apical angles produced, marginal pronotal stria complete, carinate, behind head medially with tiny angulate projection; disc antero-laterally with deep, rugulose-lacunose oval punctation, medially punctures becoming much sparser and finer, separated by several times their diameter, forming vague impunctate areas (in some specimens these can be better delimited forming ‘mirrors’), interspaces between punctures on entire pronotal disc with sparse microscopic punctation; pronotal hypomeron glabrous; scutellum very small.

Elytral epipleura punctuate; marginal epipleural stria thin; marginal elytral stria well impressed, thin, continuous along elytral apex as apical elytral stria, connected with sutural elytral stria; humeral elytral stria erased under elytral punctation; inner subhumeral stria present medially, almost unrecognizable beneath elytral punctation; dorsal elytral striae almost unrecognizable from dorsal view, from oblique view striae 1–4 faintly recognizable, first the longest, originating near elytral base running to 3/4 of elytral length apically, striae 2–4 abbreviated basally and apically; sutural elytral stria complete, deeply impressed, surface between it and elytral margin with scattered microscopic punctation. Entire elytral disc very coarsely and densely punctate, punctation rugulose-lacunose, punctures separated by less than half of their diameter, on apical third punctures disappear, forming deep elongate wrinkles; basally between fourth dorsal and sutural striae a small round ‘mirror’ (= polished area) present, its surface with scattered microscopic punctation.

Propygidium transverse, about five times as broad as long, partially covered by elytra, its punctation much finer and sparser than those of the elytra, punctures on apical half separated by about their diameter, on basal half propygidium almost impunctate, interspaces between punctures with fine alutaceous microsculpture; punctation of pygidium (Fig. 283) much coarser and denser, resembling that of elytra; pygidium carinate laterally.

Anterior margin of median portion of prosternum (Fig. 284) straight; marginal prosternal stria almost complete, interrupted laterally; prosternal process deeply concave on anterior half, laterally with sparse large oval punctures, interspaces imbricate; dorsally prosternal process with sparse shallow punctures; carinal prosternal striae (Fig. 284) carinate, slightly divergent anteriorly and united in front under a rounded loop; lateral prosternal striae carinate, widely convergent and ‘open’ anteriorly, near their origin a curious tiny projection present, unseen in any other members of the subfamily (Fig. 284). Lateral costa of antennal groove not reaching prosternal process.

Anterior margin of mesoventrite (Fig. 285) deeply emarginate medially; discal marginal mesoventral stria anteriorly well impressed, absent laterally; disc with antero-lateral depressions, with sparse large round punctures separated by several times their diameter, each puncture with a microscopic seta, interspaces between punctures with scattered microscopic punctation; meso-metaventral suture indistinct, meso-metaventral sutural stria absent; intercoxal disc of metaventrite in males medially with large depression; metaventrite of females with only slight median longitudinal depression. Disc of metaventrite in both sexes almost smooth, punctation appears only along lateral and posterior margins. Lateral metaventral stria (Fig. 286) well impressed, almost straight, not reaching metacoxa; lateral disc of metaventrite (Fig. 286) depressed, with round shallow large punctures; metepisternum evenly punctate with similar punctation, punctures on fused metepimeron much sparser than those of metepisternum; lateral metepisternal stria (Fig. 286) present, deeply impressed, complete.

Figures 289–291. 

289 Saprinodes falcifer Lewis, 1891 mandibles, dorsal view 290 labrum: left half depicting dorsal view and right half depicting epipharynx 291 antennal club, ventral view showing sensory structures of the antenna.

Intercoxal disc of first abdominal ventrite in males with large depression, in females not depressed, completely striate laterally; surface of disc in males with scattered oblong punctation, punctures becoming sparser and finer medially, in females punctation much sparser and finer.

Protibia (Fig. 287) slender, only very slightly dilated apically, terminating in massive protibial spur; setae of outer row very short, sparsely spaced; setae of median row even shorter than those of outer row, even; protarsal groove deep, designed to accommodate entire protarsus in repose; anterior protibial stria complete, costate; outer part of posterior surface of protibia (Fig. 288) with a row of sparsely spaced minuscule denticles; median part of posterior surface with a single row of regular minuscule setae; posterior protibial stria complete, very thin, with scattered minuscule setae; inner margin with single row of sparse microscopic setae.

Mesotibia slender, outer margin with a single row of around five denticles growing in size apically; setae of outer row sparse, minuscule; setae of median row similar to those of outer row, between the two rows an additional complete stria present; posterior mesotibial stria shortened apically; anterior surface of mesotibia convex, with dense row of well-sclerotized short setae; anterior mesotibial stria complete; mesotibial spur inconspicuous; claws of apical tarsomere about half its length; metatibia basically similar to mesotibia, but denticles of outer margin much sparser than those of mesotibia.

Male genitalia. Eighth sternite (Figs 292–293) separated longitudinally; vela with sparse short setae; eighth tergite and eighth sternite not fused laterally (Fig. 294). Ninth tergite (Figs 295–296) longitudinally fused medially; spiculum gastrale (Figs 297–298) gradually dilated on apical half, basal end slender, not dilated. Aedeagus (Figs 299–300) slender, slightly widening apically, slightly constricted before apex; basal piece of aedeagus very short, ratio of its length: length of parameres 1 : 10; parameres of aedeagus fused almost along their entire length, with small circular aperture for median duct; aedeagus only slightly curved from lateral view, apex of aedeagus flattened dorso-ventrally.

Figures 292–300. 

292 Saprinodes falcifer Lewis, 1891 male terminalia: 8^th sternite + 8^th tergite, ventral view 293 ditto, dorsal view 294 ditto, lateral view 295 male terminalia: 9^th + 10^th tergites, dorsal view 296 ditto, lateral view 297 male terminalia: spiculum gastrale, ventral view 298 ditto, lateral view 299 male terminalia: aedeagus, dorsal view 300 ditto, lateral view.

Some members of Saprinus are the largest of the Australian saprinines with specimens of S. (S.) viridanus measuring up to 6.70 mm (PEL). The genus is also the most species-rich; it contains 18 species, including 4 newly described and 2 introduced species. All Australopacific species (with the exception of S. detritus, S. pseudodetritus, and S. chathamensis from New Zealand/Chatham Islands, and S. nitiduloides Fairmaire, 1883 from New Britain/Solomon Islands and S. artensis Marseul, 1862 from New Caledonia that are dark brown) are characterized by strong metallic luster on the dorsum of their bodies, especially on elytra, and absence of red, orange or yellow elytral patches (often present in Palaearctic, Oriental and African species). Likewise, all have complete frontal stria (with the exception of S. grandiclava from New Guinea and two newly described species from the Chatham Islands). The frontal stria is usually widely interrupted in the Palaearctic or African species and is used to diagnose the Palaearctic members of Saprinus. One species, S. viridipennis (Australia), lacks carinal prosternal striae and two species, S. grandiclava (New Guinea) and S. viridanus (Australia), share unusually large, circular antennal clubs.

Figure 301. 

Saprinus (Saprinus) amethystinus Lewis, 1900 habitus, dorsal view.

An aedeagal peculiarity among Australopacific Saprinus is the presence of well-separated parameres (see also below; e.g. Fig. 475) in the Australian endemic species S. amethystinus Lewis, 1900, S. australis (Boisduval, 1835), S. laetus Erichson, 1834, and S. tyrrhenus Blackburn, 1903. The three New Zealand endemics, as well as S. grandiclava Kanaar, 1989 (endemic to New Guinea), have their parameres separated in the apical half and are more approximate than in the Australian species (Fig. 456).

Species of Saprinus are typical volant predators with a preference for open grasslands and xerothermic habitats where they prey upon fly larvae found in decomposing organic matter, such as mammalian dung or carcasses (Lackner 2010). Their large and well-developed sensory patches on the ventral side of their antennal clubs probably enable them to locate the carrion from a long distance (but these are lacking in S. grandiclava). The natural history of several rare species (S. grandiclava, S. amethystinus and S. tyrrhenus) is completely unknown. S. rarus sp. n. was collected in the nest of the arboreal Tree termite (Nasutitermes walkeri (Hill, 1942)) and is presumed to be a specialised termitophile.

There are eighteen species of Australopacific Saprinus: nine native and two introduced (S. chalcites and S. cupreus) are known from Australia; three species of Saprinus are present in New Zealand/Chatham Islands; New Caledonia has one endemic and one non-endemic species. New Guinea has a single endemic species, with two other, non-endemic congeners. We describe one endemic species from the Kiribati atoll (Figs 753, 755, 758–765).

Australopacific species of the genus Saprinus differ from the similar-looking species Notosaprinus irinus (Marseul, 1862) by the presence of lateral prosternal striae (usually both carinal and lateral prosternal striae are present). See discussion of Notosaprinus for more details. The genus Saprinus is the most species-rich genus of the subfamily (see e.g. Lackner 2010 or Mazur 2011 for details) with the bulk of its representatives occurring in the Palaearctic and Afrotropical Regions. Although moderately diverse also in the Oriental (Indo-Malayan) and Nearctic Regions, Saprinus of the Australopacific Region with 16 native species are rather species-poor when compared to their counterparts from the other regions. Only the Neotropical Region, with only 7 known species has fewer than Australopacific Region.

Based on the structure of the aedeagus there are two main groups of Australopacific Saprinus: in each group there are eight species with widely separated parameres and with fused parameres. Regarding the group of species with fused parameres, the following species are morphologically very similar: S. cyaneus cyaneus, S. artensis, S. nitiduloides and S. pacificus (compare Figs 335, 420, 493 and 510). Based on their overall similarities, including the structure of male genitalia we believe that they likely share a recent common ancestor that probably reached the region from the north, since a subspecies of S. cyaneus, S. cyaneus auricollis, is distributed also in the Indo-Malayan Region up to Southern Japan. These species can possibly be regarded among the so-called northern invaders, sensu Mitchell and Bouchard (2008). Another species with fused parameres, Saprinus splendens, is widely distributed in the tropical regions of Africa/Arabian peninsula and the Oriental (Indo-Malayan) Region as far east and north as Japan, and its occurrence in the Australopacific Region (Australia and Marianna Islands) is not surprising. The three remaining species with fused parameres (S. rarus, S. viridanus, and S. viridipennis) are Australian endemics. Saprinus rarus has a uniquely triangulate dilated and thickened antennal scape (Fig. 529), strongly dilated tibiae (Fig. 536) and elongate body shape (Fig. 528), features that are probably reflecting its termitophilous lifestyle, and may protect it against termites (thickened antennal club and dilated tibiae protect larger space of body venter when in repose). S. viridipennis lacks the carinal prosternal striae entirely (Fig. 609; an autapomorphic feature within Saprinus), while the antennal club of S. viridanus is unusually large and circular, resembling the form present in the New Guinean endemic S. grandiclava (compare Figs 460 and 586).

The remaining eight species with (widely) separate parameres (Australian S. amethystinus, S. australis, S. tyrrhenus, S. laetus; New Guinean S. grandiclava and New Zealand S. detritus, S. pseudodetritus and S. chathamensis) are unique among the 157 described species of Saprinus (sensuMazur 2011): no other species of Saprinus have this character. In the published phylogenetic analysis of the senior author (Lackner 2014d) only the type species of the genus, S. (S.) semistriatus, a taxon with fused parameres, was analysed. As noted above (see remarks section of Notosaprinus), S. (S.) semistriatus was recovered sister to Notosaprinus irinus; this clade itself was recovered sister to a larger clade containing North American Xerosaprinus, African Pilisaprinus and Paraphilothis, Middle-Asian Styphrus, Palaearctic Phaonius (itself subgenus of Saprinus). None of these relatives of Saprinus possess separated parameres. On the other hand, two members of the genus Microsaprinus Kryzhanovskij, 1976 that was recovered in the aforementioned analysis closer to the root, possess separated parameres of the aedeagus (albeit not widely; see Secq and Secq 1995 for figures). Looking at the character value nodes of Saprinus near and even more distant relatives the basal aedeagal condition does not appear therefore to be the separated-parameres character state, with the single exception of the two Microsaprinus species.

Australia: Queensland: Taylor Range.

Saprinus amethystinus Lewis, 1900: Lectotype, present designation: ♂, side-mounted, terminalia and pygidium glued to the same card as the specimen, right antennal funicle missing, right mid-leg missing, right metatarsus missing, left protarsal claw missing, with the following labels: “Taylor Range / Queens Land / (Janson)” (written); followed by: “Saprinus / amethystinus / Type Lewis” (written); followed by: “G. Lewis Coll. / B.M. 1926-369.” (printed); followed by: “Type” (round, red-margined label); followed by: “09-088” (yellow, pencil-written label, added by the senior author); followed by: “Saprinus / amethystinus / LEWIS, 1900 / LECTOTYPE / des. T. Lackner ‘011” (red label, written) (BMNH). This species was described from an unknown number of specimens and the lectotype designation fixes the species identity.

AUSTRALIA. New South Wales: 1 ♂, Quirindi, G.E. Bryant, 2.xi.[19]08, G. Bryant Coll., 1919–147, Dahlgren det. (BMNH); 1 spec., Caragabal, 13.ix.1966, Z. Liepa (ANIC); 10 specs., Euglo ex., Humbug Creek, 12.xi.1972, D.A. Doolan leg. (AMS; 3 specs. in coll. TLAN). Queensland: 1 ♀, Urangan, Manski, collector unknown (ANIC).

Unknown.

Australia: New South Wales and Queensland (Fig. 759).

This is a rare species of Australian Saprinus known from only a handful of specimens; the last ones were collected in 1972.

Body length: PEL: 3.25–3.85 mm; EL: 2.00–2.40 mm; APW: 1.25–1.60 mm; PPW: 2.25–2.75 mm; EW: 2.50–3.00 mm.

Body (Fig. 301) rectangular oval, dorsally convex, ventrally rather flattened; cuticle dark brown, shining, pronotum darker, piceous black, with bronze metallic luster; elytra on basal third light brown, rest of elytra darker, elytra with dark blue metallic luster; legs, mouthparts and antennal scape castaneous brown; antennal club light brown.

Antennal scape (Fig. 302) thickened and triangularly dilated, punctuate, with several setae; antennal club (Fig. 303) covered with dense short sensilla intermingled with sparse longer erect setae; lower third of club asetose; sensory structures of antennal club not examined.

Figures 302–310. 

302Saprinus (Saprinus) amethystinus Lewis, 1900 head, dorsal view 303 antennal club, dorsal view 304 pygidium 305 prosternum 306 mesoventrite 307 lateral disc of metaventrite + metepisternum 308 protibia, dorsal view 309 ditto, ventral view 310 metatibia, dorsal view.

Mandibles carinate laterally, dorso-laterally densely and coarsely punctuate, rounded, mandibular apex acute; mentum square-shaped, anterior margin with deep conspicuous median notch; labrum finely and sparsely punctuate, convex, with shallow median depression; labral pits present, each with a single labral seta; other mouthparts not examined.

Clypeus (Fig. 302) evenly densely punctuate; frontal stria weakened, but complete and outwardly arcuate medially, supra-orbital stria carinate; frontal disc (Fig. 302) coarsely and densely punctuate; eyes convex, well visible from above.

Pronotal sides (Fig. 301) moderately narrowing anteriorly, apical angles acute, pronotal depressions absent, anterior incision for head shallow, semicircular; marginal pronotal stria complete, carinate, visible along its entire length from dorsal view; pronotal disc laterally with a band of deep dense elongate punctures, becoming finer and sparser behind head, median part of pronotal disc almost smooth, with only scattered microscopic punctation; double row of fine ovoid punctures present along pronotal base; pronotal hypomeron with short yellow setae; scutellum very small, but visible.

Elytral epipleura with sparse fine punctures; marginal epipleural stria complete; marginal elytral stria well impressed and slightly carinate, continued as complete apical elytral stria. Humeral elytral stria well impressed on basal third, in most cases connected to inner subhumeral stria; variously long fragment of inner subhumeral stria present also laterad of humeral stria (in two of the three studied specimens); three dorsal elytral striae 1–3 well impressed, impunctate, carinate, apically slightly surpassing elytral half, fourth dorsal elytral stria strongly abbreviated, present as short basal fragment, basally not connected with sutural elytral stria; sutural elytral stria well-impressed, in punctures, abbreviated on basal fifth, apically connected with apical elytral stria; elytral disc on apical four-fifths punctuate, punctures becoming denser apically, forming almost elongate strioles near elytral apex.

Propygidium with dense round deep punctures separated by less than their diameter intermingled with much finer and sparser punctures; pygidium (Fig. 304) with similar, if somewhat sparser punctation; punctures much larger than those of propygidium.

Anterior margin of median portion of prosternum (Fig. 305) almost straight, rounded laterally; marginal prosternal stria present laterally and also as a rather long medial fragment; prosternal process flattened, surface between carinal prosternal striae depressed, with coarse punctures, apico-laterally substrigulate-punctate; carinal prosternal striae well-impressed, carinate, sub-parallel, slightly divergent anteriorly, forming a rounded loop, united in front (Fig. 305); lateral prosternal striae carinate, rather short, apically attaining carinal prosternal striae at about four-fifths of their length, surface around them with microscopic setae.

Anterior margin of mesoventrite (Fig. 306) almost straight, slightly inwardly arcuate; discal marginal mesoventral stria well impressed, complete; disc with coarse and large punctation; meso-metaventral sutural stria largely absent, present only as two short lateral fragments; intercoxal disc of metaventrite flattened; disc of metaventrite for the most part almost smooth, only with scattered microscopic punctation, along posterior margin (especially in area behind hind coxae) denser and coarser punctation appears; lateral metaventral stria (Fig. 307) well impressed, carinate, almost straight, shortened; lateral disc of metaventrite (Fig. 307) slightly concave, with dense shallow large setigerous punctures; metepisternum (Fig. 307) with similar setigerous punctures, on basal two-thirds and on fused metepimeron punctures becoming much sparser, not setigerous; metepisternal stria present as short intermittent fragments.

Intercoxal disc of first abdominal ventrite completely striate laterally; disc along basal and lateral margins with deep round punctures becoming finer and sparser medio-apically.

Protibia (Fig. 308) slightly dilated, outer margin with three triangular teeth topped by large triangular denticle, second and third teeth conspicuously larger than first, followed by two tiny low teeth topped by denticle; setae of outer row regular, moderately spaced; protarsal groove shallow; anterior protibial stria present on basal two-thirds, next obliterated; setae of median row shorter and sparser than those of outer row; two tarsal denticles present near tarsal insertion; protibial spur (Fig. 309) massive, bent, growing out from apical margin of protibia; outer part of posterior surface obscurely variolate, separated from glabrous median part of posterior surface by definite carinate boundary; posterior protibial stria complete, almost along entire length with dense row of long well sclerotized setae; inner row of setae single, setae sparse, shorter than those of posterior protibial stria.

Mesotibia moderately dilated, outer margin with a row of sparse long denticles growing in size apically, another row of much shorter sparser denticles situated on anterior surface of mesotibia; setae of outer row regular, thick, almost as long as denticles themselves; setae of median row irregular, shorter and finer; posterior mesotibial stria not examined; anterior surface of mesotibia sparsely punctuate; anterior mesotibial stria almost complete; mesotibial spur stout, moderately long; apical margin of mesotibia anteriorly with two short denticles; inner margin of mesotibia with sparse row of long setae; claws of apical tarsomere slightly bent, shorter than half its length; metatibia (Fig. 310) slenderer and longer than mesotibia, in all aspects similar to it, but denticles on outer margin much shorter and sparser.

Male genitalia. Eighth sternite (Figs 311–312) strongly sclerotized, separated medially approximately on its apical half, apex with several microscopic setae, velum absent; eighth tergite and eighth sternite fused laterally (Fig. 313). Ninth tergite (Figs 314–315) typical for the subfamily; tenth tergite (Fig. 314) outwardly arcuate, rounded; spiculum gastrale (Figs 316–317) gradually dilated on most of its apical half; basal end slightly dilated. Aedeagus (Figs 318––319) with parameres separated almost on their entire length; basal piece of aedeagus short, ratio of its length : length of parameres 1 : 5; aedeagus slightly curved from lateral view (Fig. 319).

Figures 311–319. 

311Saprinus (Saprinus) amethystinus Lewis, 1900 male terminalia: 8^th sternite + 8^th tergite, ventral view 312 ditto, dorsal view 313 ditto, lateral view 314 male terminalia: 9^th + 10^th tergites, dorsal view 315 ditto, lateral view 316 male terminalia: spiculum gastrale, ventral view 317 ditto, lateral view 318 male terminalia: aedeagus, dorsal view 319 ditto, lateral view.

New Caledonia.

Figure 320. 

Saprinus (Saprinus) artensis Marseul, 1862 habitus, dorsal view.

Saprinus artensis Marseul, 1862: Lectotype, present designation: ♂, side mounted on a triangular card, with terminalia glued on the same triangular card as specimen, right protarsus and left mesotarsus missing, both metatarsal claws broken off, with the following labels: “13 / Saprinus / artensis m. / illegible text” (round pink label, written); followed by: “artensis / type” (pencil-written label added probably by G. Dahlgren); followed by: “TYPE” (red-typed label); followed by: “MUSEUM PARIS / artensis / COLL / DE MARSEUL 1890” (pink label, typed-written); followed by: “09-052” (yellow, pencil-written label, added by the senior author); followed by: “Saprinus / artensis / Marseul, 1862 / LECTOTYPE / Des. T. Lackner ’11” (red label, written) (MNHN). This species was described from an unknown number of specimens and the lectotype designation fixes the identity of species.

1 ♂, New Caledonia (MNHN). Apparently this specimen is also from Marseul’s collection, but does not originate from the same sampling as the lectotype. There is no “Type” label on it and therefore it is not designated as the paralectotype; 5 ♂♂, “New Caledonia”, all from Schmidt’s collection, one male has a small label: “Type” – it is not clear whether it forms a part of the syntype series or not (ZMHUB); 1 ♀, New Caledonia, A. Deyr[olle] (NCB). 1 ♂ & 1 ♀, N. Caledonia, Fauvel (BMNH).

Unknown, presumably similar to congeners.

Endemic to New Caledonia, rarely collected (Fig. 760).

Body length: PEL: 3.35 mm; EL: 2.00 mm; APW: 1.25 mm; PPW: 2.50 mm; EW: 2.80 mm (only the type specimen was measured).

Body (Fig. 320), convex, cuticle dark brown, almost black, shining, without metallic luster; legs, mouthparts and antennal scape dark brown; antennal club black.

Antennal scape (Fig. 321) black, slightly thickened, finely punctuate, with two setae; antennal club (Fig. 322) covered with dense short sensilla intermingled with sparse longer erect setae; sensory structures of antennal club not examined.

Figures 321–329. 

321Saprinus (Saprinus) artensis Marseul, 1862 head, dorsal view 322 antennal club, ventral view 323 mentum, ventral view 324 propygidium + pygidium 325 prosternum 326 mesoventrite 327 lateral disc of metaventrite + metepisternum 328 protibia, ventral view 329 mesotibia, ventral view.

Mandibles dorso-laterally densely punctuate, rounded; labrum finely and sparsely punctuate, convex, with shallow median depression; labral pits present, each with a single labral seta; mentum (Fig. 323) almost quadrate, furnished with sporadic seate, anterior margin with deep triangular notch furnished with several longer setae; other mouthparts not examined.

Clypeus (Fig. 321) broad, flattened, sloping down laterally, its punctation denser and coarser than that of labrum; frontal and supra-orbital striae complete, slightly carinate; frontal disc (Fig. 321) finely punctuate, punctures separated by about several times their diameter; eyes convex, well visible from above.

Pronotal sides (Fig. 320) moderately narrowing anteriorly, apical angles obtuse, pronotal depressions shallow but present, anterior incision for head shallow; marginal pronotal stria complete, slightly carinate, visible along its entire length from dorsal view, weakened anteriorly; pronotal disc laterally with a band of deep dense elongate punctures originating approximately in pronotal depressions, but not reaching basal angles of pronotum, between it and pronotal margin a narrow smooth band present; rest of the pronotal disc almost smooth, with only scattered microscopic punctation; double row of fine ovoid punctures present along pronotal base not reaching ante-scutellar area; pronotal hypomeron glabrous; scutellum small, visible.

Elytral epipleura with a double row of fine punctures; marginal epipleural stria complete; marginal elytral stria well impressed and slightly carinate, continued as complete apical elytral stria. Humeral elytral stria well impressed on basal fourth, faintly connected to rather long inner subhumeral stria; four dorsal elytral striae 1–4 well impressed, all about the same length (third elytral stria slightly shorter than others), not reaching elytral half apically; first and second dorsal elytral striae in fine punctures, first elytral interval with elongate strioles; third and fourth striae in large sparsely set punctures, surface of second and third elytral intervals almost smooth, only with microscopic punctation; fourth dorsal elytral stria curved towards sutural elytral stria but not connected with it; sutural elytral stria well-impressed, in fine punctures, abbreviated on basal fourth, apically connected with apical elytral stria; elytral disc on apical 4/7 in dense large punctures; punctures separated by about their own diameter, laterally and especially apically punctures aciculate; elytral flanks almost impunctate.

Propygidium (Fig. 324) very densely punctuate, punctures separated by less than their diameter; pygidium (Fig. 324) with similar, if somewhat sparser punctation, interspaces in both cases imbricate.

Anterior margin of median portion of prosternum (Fig. 325) almost straight, rounded laterally; marginal prosternal stria present laterally and also as apical fragment; prosternal process between carinal prosternal striae flat, punctuate; carinal prosternal striae carinate, parallel on basal three-fourths, on apical fourth slightly divergent, united in front (Fig. 325); lateral prosternal striae carinate, rather short, apically attaining carinal prosternal striae at about three-fourths of their length.

Anterior margin of mesoventrite (Fig. 326) distinctly inwardly arcuate; discal marginal mesoventral stria well impressed, carinate; disc with sparse fine punctation, punctures becoming larger near meso-metaventral suture; meso-metaventral sutural stria indicated by a row of large punctures; intercoxal disc of metaventrite flattened, with slight longitudinal median depression; disc of metaventrite for the most part almost smooth, along posterior margin several rows of punctation appears; lateral metaventral stria (Fig. 327) well impressed, carinate, almost straight, shortened; lateral disc of metaventrite (Fig. 327) slightly concave, with dense shallow large setigerous punctures; metepisternum (Fig. 327) similar, but with deeper and larger punctures without setae, on fused metepimeron punctures becoming much sparser; metepisternal stria present along fused metepimeron, along metepisternum present as short intermittent fragments.