Figulus nitens Waterhouse 1874
Creators: name: Monte, Cinzia
name: Zilioli, Michele
name: Bartolozzi, Luca
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Coleoptera
Lucanidae
Figulus
Figulus nitens

description: Figulus nitens Waterhouse, 1874
(Figs. 12–15)
Figulus nitens Waterhouse, 1874: 7; Parry 1875: 22; Felsche 1898: 78; Van Roon 1910: 52; Mjöberg 1916: 14; Didier 1937: 30; Nagel 1939: 1; Didier & Séguy 1953: 182; Benesh 1960: 27; Parry 1875: 22; Maes 1992: 39; Moore & Cassis 1992: 10; Krajcik 2001: 12; Krajcik 2003: 81; Maes 2014; Schoolmeesters 2015.
Figulus tambourinensis Mjöberg, 1916: 13; Nagel 1939: 1; Didier & Séguy 1953: 183; Benesh 1960: 29; Maes 1992: 41; Moore & Cassis 1992: 11; Krajcik 2001: 13; Krajcik 2003: 84; Maes 2014; Schoolmeesters 2015. New synonymy. Figulus howei De Lisle, 1967: 542; Lacroix 1980: 126; Bomans 1989: 14; Maes 1992: 38; Moore & Cassis 1992: 9; Krajcik 2001: 10; Krajcik 2003: 77; Fujita 2010: 387; Maes 2014; Schoolmeesters 2015. New synonymy.
Type material. Figulus nitens Waterhouse: Lectotype here designated: Lectotype: “TYPE / SP. [printed]”, “ Figulus / nitens, C. Waterh. / (Type) Ent. M. Mag. 16/4 [handwritten]”, “Ex Musaeo / Parry [printed]”, “MUSÉUM PARIS / 1952 / COLL. R OBERTHÜR [printed]”, [MNHN]; Paralectotypes: 1 paralectotype: “Moreton / Bay [handwritten]”, F. nitens / var. n. Holl / Waterhouse [h]”, Ex Musaeo / Parry [printed]”, MUSÉUM PARIS / 1952 / COLL R OBERTHÜR [printed]” [MNHN]; 1 paralectotype: F. ni t e n s / Water. [handwritten]”, “MUSÉUM PARIS / 1952 / COLL R OBERTHÜR [printed]” [MNHN]. Figulus howei De Lisle (Ƌ): “Holotypus / M.O. de Lisle nom. 1967 [printed]”, “Lord Howe Island / (New South Wales) [handwritten]”, “TYPE [printed]”, “ Figulus / howei m. / M.O. de Lisle det. [printed, handwritten]” [MHNG]. Figulus tambourinensis Mjöberg: Lectotype here designated: Lectotype male: “Mt. Tam / bourine [printed]”, “Queensl. / Mjöberg [printed]”, “8303 [printed]”, “RiksMuseum / Stockholm [printed]”, “ tambourinensis / Mjöb. [handwritten]” [NHRS]; Paralectotype: 1 paralectotype: “Type [printed]”, “Mt. Tam / bourine [printed]”, “Queensl. / Mjöberg [printed]”, “8302 / E91+ [printed]”, “RiksMuseum / Stockholm [printed]”, “ Figulus tambourinensis [handwritten]” [NHRS].
Diagnosis. Figulus nitens differs from the all other described members of the genus in Australia by the shape of the mentum, which has two small tubercles at sides near the base, and by the shape of male genitalia.
Redescription. Measurements: size range (n = 110): TL: 11.0– 16.9 mm; PL: 2.8 3–4.50 mm; EL: 6.21–9.50 mm; PW: 3.58–5.62 mm; EW: 3.66–5.70 mm.
Male (Fig. 12). Body elongate, with subparallel sides, from black or dark brown to reddish brown, glossy.
Head: transverse, about twice as wide as long, slightly depressed. Frontoclypeal region slightly concave at the anterior margin, finely punctate and with two small glossy areas situated before the eyes. Labrum broad with anterior margin slightly convex, moderately protruding, and with long, golden setae. Vertex slightly concave in the middle, inclined anteriorly; with two small, transversal elevations. Two longitudinal carinae, moderately developed, are present along the inner margin of eyes. Eyes large, divided into two parts by the canthi, the upper part smaller than the lower, with distinct facets. Canthus largely protruding, rounded laterally and directed obliquely inwards behind, slightly depressed and smooth upper surface, with a few minute punctures on the lateral margin. Head surface smooth, glossy, finely and sparsely punctate. Mandibles a few shorter than head, strong and asymmetrical, rounded, apices slightly blunted and curved upwards; upper surface at base slightly elevated. Internal margin of the right mandible with a moderately developed basal tooth and a medial strong tooth, with apex blunted directed inwards. Internal margin of the left mandible with a moderately developed basal tooth, a medial tooth and a preapical slight angulation moderately developed. Antenna with 10 antennomeres, with scape slightly shorter than the remaining antennomeres; club of last three antennomeres, which are flatter and broader than the remaining antennomeres of the funicle. Mentum subrectangular (Fig. 13), medially concave, with anterior margin slightly bilobed and with sides almost right or slightly rounded, elevated. Base feebly concave in the middle forming a slightly, elevated process, feebly bilobed anteriorly and reaching the half of the mentum. Two small tubercles, moderately developed, at sides. Surface strongly punctate, rugose in the middle.
Thorax: pronotum subrectangular, with subparallel sides; anterior margin slightly convex, moderately elevated, with a small, blunt medial tubercle and a shallow depression on anterolateral margins; posterior margin slightly bisinuate; anterior angles produced with rounded anterior margin; posterior angles largely rounded. Superficial longitudinal medial groove, deeper in the middle with some large, circular punctures. Pronotal surface glossy, finely and sparsely punctate on the disc, densely punctate around the medial tubercle; sides with dense and large punctures. A patch of large close punctures at midway on the sides. Scutellum subtriangular, surface smooth and glossy. Elytra at base a few narrower than prothorax, and about twice as long as pronotum. Base concave, subparallel sides, apex regularly rounded. Humeral angles acute, with a small tubercle. Elytral surface glossy, with 9 visible subparallel striae, the dorsal striae deeper than the lateral striae. Stria 1 is deep and the only stria reaching apex; formed by a row of large, confluent punctures. Striae 2‒4 formed by a row of large, confluent punctures except at apex; stria 5 as striae 2‒4, but in the posterior stria the punctures are well separated from each other; stria 6 formed by a row of large punctures well separated from each other. Interstriae much larger than striae, slightly convex and finely punctate. Elytral margins and apex strongly punctate, with short, erect, red setae. Wings present. Legs thin and straight. Protibiae with strong, elongate apical fork, and a long, strong spur internally at apex; lateral margin with 4‒5 small teeth, increasing in size distally. Mesotibiae with 4 long, acute, apical teeth, and 2 apical spurs, 1 larger than the other; lateral margin with strong submedial tooth, preceded by 1‒2 smaller teeth. Metatibiae with 4 acute, apical teeth, and 2 long apical spurs, 1 larger than the other; lateral margin with strong submedial tooth, preceded by a smaller tooth. Prosternum subpentagonal in shape behind the coxae, surface with large and deep punctures at anterior edge and with smaller punctures medially and at posterior edge. Metasternum smooth, with a long medial groove, deeper at its posterior edge; sides of metasternum with some horse-shoe shaped, superficial hollows with a short, golden seta in the middle.
Abdomen: first abdominal ventrite with some horse-shoe shaped, superficial hollows with a short, golden seta in the middle. The other ventrites finely punctate, with some larger punctures at sides. Last ventrite with scattered, large punctures, finer at posterior margin.
Male genitalia: penis broad at base, strongly narrowed at apex, with a constriction before the latter (Fig. 14), in dorsal view broader than parameres, slightly sinuose in lateral view, strongly bent at base close to articulation with basal piece. Permanently everted internal sac drawn out into an exceedingly long (several times the tegmen length), broad, colorless or yellow flagellum tapering towards apex. Parameres short (Figs. 14–15) with marginal fringe of short setae entirely enclosed in the cuticle (not represented in figure). Basal piece long, about 2.5 times the length of paramere; ventrally largely membranous, dorsally sclerotized, with a moderately large, elongate sclerotic area along midline. Two long, slender struts run along basal piece, from penis articulation to slightly beyond the base of the lobed expansion of basal piece (Figs. 14–15).
Female. All external morphological characters as in male.
Female genitalia: hemisternites elongate, separed by 2 small, pigmented patch triangularly shaped; apically with long, golden setae, surrounding the styli; anteriorly broad up to about half of their length and close to the base; outer edge notched; medial edge strongly indented; styli slender, slightly longer than wide, slightly curved outwards. Bursa copulatrix represented by a slight expansion of the wall of spermathecal duct near junction with the median oviduct. Spermathecal duct long ending in a bulbous subcircular expansion proximally colorless then pigmented in the total length; spermathecal gland colorless, small, bulbous, with short duct.
Intraspecific variation. The specimens examined do not show significant morphological variability except for the size of body (see section “Measurements”).
Additional material examined. Australia [state unknown]: 1 specimen, comparé au type (= compared to type), ex Musaeo Mniszech, 6659, H.E.Bomans BMNH (E) 1999-248, 406659 (handwritten) (NHM). AUSTRALIAN CAPITAL TERRITORY: 1 specimen, Mount Ainslie Australian Capital Territory, 28.X.1967, P.B.Carne (ANIC); 1 specimen, Canberra, 31.III.1964, C.G.L.Gooding, C.G.L.Gooding Collection, donated to A.N. I.C. 1979, 8632 (handwritten), H.E.Bomans BMNH (E) 1999-248, 408632 (handwritten) (NHM); 2 specimens, Reid, 19.IX.1960, 705, bred from yellow box log, K.M.L. Key (ANIC). NEW SOUTH WALES: 1 specimen, N.S.Wales, ex Musaeo Möllenkamp 1913, Muséum Paris 1952 Collection R. Oberthür (MNHN); 1 specimen, New South Wales, Muséum Paris 1952, Collection R. Oberthür (MNHN); 1♀, Barrington Tops, 23.I.1927, T.G.Campbell, T 59842, K 293086 (AMS); 1 specimen, Barrington Tops, 13.I. 1947, 500 feet, R.Calven, H.Hughes & A.Musgrave, K 293087 (AMS); 1 specimen, Barrington Tops, Macleay Museum University of Sydney 2006 Australia, 11378 (handwritten), H.E.Bomans BMNH (E) 1999-248, 411378 (handwritten) (NHM); 1 specimen, Barrington Tops State Forest, 31°54'13''S, 151°27'59''E, Gummi Falls, 6–15.I.1999, M.Yee, R.B.Halliday, K.Pullen, & K.Hebbard, at light, open forest (ANIC); 1 specimen, Belanglo State Forest, 34°32'S, 150°14'E, 18.VIII.1990, Tom Gush, under bark cut tree, Tom Gush Collection 2347, Macleay Museum University of Sydney 2006 Australia, 16679 (handwritten), H.E.Bomans BMNH (E) 1999-248, 416679 (handwritten) (NHM); 1Ƌ, Mount Belmore State Forest, N Belmore Road, via Busbys Flat, 4.I. 1999, 300 m, 99:19Ƌ, mv light (ANIC); 1Ƌ, Mount Belmore State Forest, T-Ridge Rd, via Busbys Flat, 4.XII. 1998, 500 m, 98:686 Ƌ, S.G.Watkins, mv light, S.G.Watkins Collection, donated 2001 (ANIC); 1 specimen, Blue Mountains, ex Collection Australian Museum (ANIC); 1 specimen, Blue Mountains, Mount Wilson, 19.XII.1986, D.K.McAlpine, R.deKeyzer, & B.J.Day, K 293094 (AMS); 1Ƌ, 1♀, Broulee, 13.I.1963, in Casuarina, D.Carne (ANIC); 1 specimen, Bungabee State Forest, SE Kyogle, 10.XII.1996, mv light, S.G.Watkins, mv light, S.G.Watkins Collection donated 2001 (ANIC); 1 specimen, Cabbage Tree Creek, Clyde Mountain, 26.X.1982, Doyen & Lawrence, under bark of rotten logs (ANIC); 1 specimen, Cambridge Plateau Richmond Range State Forest, WNW Casino, 15.XI.1996, mv light, 96:567 Ƌ, S.G.Watkins, S.G. Watkins Collection, donated 2001 (ANIC); 1 specimen, Chiswick, 16 km S Armidale, 18.II.1980, light trap, M.R.Portor (ANIC); 1 specimen, Fortis Creek State Forest, N Grafton, 13.XI.1996, mv light, S.G.Watkins, S.G.Watkins Collection, donated 2001 (ANIC); 1 specimen, Glen Innes, Prison farm, VII.1969 - XII.1970, University of New England Collection donated 1983 (ANIC); 1 specimen, Gordon, Sydney, XI.1989, J.A.Walker, R.deKeyzer Insect Collection, K 293327 (AMS); 1 specimen, Helensburgh, Sydney [error on original label], 16.VIII.1992, R.deKeyzer, R.deKeyzer Insect Collection, K 293328 (AMS); 1♀, Mackellar Range Road, Bungabee State Forest, 15 km SE Kyogle, 25.X.1999, S.G.Watkins, mv light, S.G.Watkins Collection, donated 2001 (ANIC); 1 specimen, Mooney Mooney Creek, near Gosford, 5.XII.1976, D.K.McAlpine, K 293092 (AMS); 1♀, Ogilvie Trail, Banyabba Nature Reserve, SSW Whiporie, 130 m, 27.XI.2000, S.G.Watkins, mv light, S.G.Watkins Collection, donated 2001 (ANIC); 1 specimen, Ourimbah, Sydney [error on original label], 25.II.1925, W.W.Froggatt, W.W.Froggatt Collection (ANIC); 1♀, Pennefathers Road, Cherry Tree State Forest via Casino, 26.XII.1996, mv light, S.G.Watkins, S.G.Watkins Collection, donated 2001 (ANIC); 1Ƌ, 2♀, 5 specimens, Penrose State Forest, 6.V.1972, D.A.Doolan, D.A.Doolan Collection, K 293089, K 293090, K 293091, K 293093, K 293124, K 293125, and K 293126 (AMS); 1 specimen, Platypus Flat, camping area, Nymboi-Binderay National Park, 21.XII.2000, R.deKeyzer, R.deKeyzer Insect Collection, K 293329 (AMS); 1Ƌ, 1♀, Powerful Owl Reserve, Girard State Forest, W of Drake, 7.XII.1998 ‒ 8.I. 1999, 690 m, 98:758Ƌ, S.G.Watkins, mv light, S.G.Watkins Collection, donated 2001 (ANIC); 1 specimen, Sea Acres Reserve, Port Macquarie, 3–4.IX.1970, K.R.Pullen (ANIC); 1 specimen, Track W of Sunnyside Ridge Road, near Bungleboori, picnic area, Newnes State Forest, 1.I.2011, at mv light, R.deKeyzer & A.Scott, K 293330 (AMS); 1 specimen, Tarago, 20.XII.1967, W.J.M.Vestjens (ANIC); 1 specimen, Taralga, Dr.Broom. 99-130, BMNH(E) # 607752 (NHM); 1Ƌ, 1♀, Wadsworth Trail at Potoroo Road, Dingo State Forest, 4.XII.1995 ‒ 24.I.1996, mv light, 95:1097Ƌ, S.G.Watkins Collection, donated 2001 (ANIC). QUEENSLAND: 1 specimen, Mount Glorious State Forest, XI–XII.1986, ex Argyrodendron actinophyllum, Y.Basset Collection (ANIC); 1 specimen, Mount Tamborine, H.J.Carter Collection P. 20.4.22, COL- 85977 (NMVM); 1 specimen, Mount Tamborine, National Museum Victoria, COL-85853 (NMVM). VICTORIA: 1 specimen, Bendoc, I.1938, F.E.Wilson Collection, COL-85166 (NMVM); 1 specimen, Launching Place, I.1919, C.Oke, COL-86014 (NMVM); 1 specimen, Moe, 15.IX.1945, C.G.L.Gooding, C.G.L.Gooding Collection, donated to A.N. I.C. 1979 (ANIC); 4 specimens, Moe, 4.IX.1954, C.G.L.Gooding, C.G.L.Gooding Collection, donated to A.N. I.C. 1979 (ANIC); 1 specimen, Tyers River, IX.1973, T.Brown, C.G.L.Gooding Collection, donated to A.N. I.C. 1979 (ANIC); 1 specimen, Warragul, V.1962, J.M., C.G.L.Gooding Collection, donated to A.N. I.C. 1979 (ANIC). LORD HOWE ISLAND: 1♀, Lord Howe Island, 11–14.V.2003, rotting Howea belmoreana wet forest, slope east of golf course, C.Reid, K 187452 (AMS); 1 specimen, Lord Howe Island, 21–30.I.1965, G.Bornemissza, ex Collection L.Bartolozzi, 11089, H.E.Bomans BMNH (E) 1999-248, 411089 (NHM); 1 specimen, Lord Howe Island, 26.XI.1966, R.W.Taylor, ex rotten log, Johnston Beach, pasture, <100 ft., R.W.Taylor accession 66.341, 8629 (handwritten), H.E.Bomans BMNH (E) 1999-248, 408629 (handwritten) (NHM); 1 specimen, Lord Howe Island, 21.I.1985, G.F.Bornemissza, Graeme Krake Collection, donated Feb. 2007, COL-86867 (NMVM); 1 specimen, Lord Howe Island, 21.I.1985, G.F.Bornemissza, K 187437 (AMS); 2ƋƋ, Lord Howe Island, 21– 30.I.1985, G.F.Bornemissza (LBC); 1 specimen, Lord Howe Island, 29.XII.1921, R.Baxter, K 45609 (handwritten), K 187441 (AMS); 1♀, Lord Howe Island, 2–3.XII.1966, approximately 200 feet, old settlement, R.W.Taylor, accession 66.452, ex rotten log (ANIC); 1♀, Lord Howe Island, 28031 (handwritten), K 293080 (AMS); 1Ƌ, 1 specimen, Lord Howe Island, Macleay Museum University of Sydney 2006 Australia, 11379-a (handwritten) and 11379-b, H.E.Bomans BMNH (E) 1999-248, 411379·1 and 411379·2 (NHM); 1♀, Lord Howe Island, Macleay Museum University of Sydney 2006 Australia (LBC); 1 specimen, Lord Howe Island, A.M.Lea, K 293082 (AMS); 1Ƌ, Lord Howe Island, XII.1912, A.Lea, 12778 (handwritten), H.E.Bomans BMNH (E) 1999-248, 412778 (handwritten) (NHM); 1Ƌ, 1 specimen, Lord Howe Island, XII.1912, A.Lea (LBC) and (GTC); 1 specimen, Lord Howe Island, A.Lea (QMB); 1Ƌ, Lord Howe Island, 15–17.VIII.1914, R.D.Laurie Collection, British Museum 1954-588, number 34, BMNH (E) # 607416 (NHM); 2 specimens, Lord Howe Island, Australia 1919-46, BMNH (E) # 607414 and # 607415 (NHM); 2 specimens, Lord Howe Island, XI.1955, J.Balderson F.E.Wilson Collection, COL-85169 (NMVM); 2 specimens, Lord Howe Island, 31°30'S, 159°04'E, XII.2002, Mr Ushijima & Mr Chikakura, K 293078 and K 293079 (AMS); 9 specimens, Lord Howe Island, 21.I.1985, G.Bornemissza (ANIC); 2 specimens, Lord Howe Island, 21.I.1985, G.Bornemissza (GBC) and (AKC); 1 specimen, Lord Howe Island, 10–15.II.1992, G.F.Bornemissza (GBC); 1 specimen, Lord Howe Island, public jetty, 24.I.1979, T.Kingston, at light, K 187439 (AMS); 1 specimen, Lord Howe Island, Erskine Valley, 18.XII.1923, G.Whitley, K 48848 (handwritten), K 293081 (AMS); 1♀, Lord Howe Island, Boat Harbour Trail, (CBCR site 27), approximately 40 m, 31°33'41''S, 159°05'43''E, 30.XI.2000, C.Reid, under bark Chionanthus log, K 187446 (AMS); 1 specimen, Lord Howe Island, 800 m along Goat House Track to Intermediate Hill, 31°33'16''S, 159°05'05''E, 8.XII.2000, on Cryptocarya LHI/GC/L14, C.Reid, K 187444 (AMS); Lord Howe Island, Intermediate Hill, 6.XI.1979, G.B.Monteith, rainforest (QMB); 1 specimen, Lord Howe Island, Lower Rd, 31°34'15''S, 159°04'30''E, cliffs just before start of rope traverse, 28.XI. 2000, 75 m, C.Reid, under Cryptocarya bark, K 187445 (AMS); 1Ƌ, Lord Howe Island, 21–30.I.1965, G.Bornemissza (NHM); 1Ƌ, Lord Howe Island, 3.XII.1966, Nichols’ track, in dead wood, E.Britton (ANIC); 1 specimen, Lord Howe Island, halfway between Rocky Point & Muttonbird Point, on path, 4.XII.2003, L.Meades, K 293077 (AMS); 1 specimen, Lord Howe Island, 11–14.V.2003, rotting Howea belmoreana, Valley of Shadows loop track, C.Reid, K 187449 (AMS); 1 specimen, Lord Howe Island, Smoking Tree Ridge, 1.XI.1979, G.B.Monteith, rainforest (QMB); 1 specimen, Lord Howe Island, rotting Olea paniculata, 100 m into forest path at Soldiers Creek, 11–14.V.2003, C.Reid (AMS).
Distribution. Australia: Australian Capital Territory, New South Wales, Queensland, Victoria, South Australia, Western Australia, and Lord Howe Island.
Natural history data. According to label data, most adults were collected at mercury vapor lights, mainly from November to January, but there are single records in February, May, August, and September. They were most commonly found in rotten logs, in dead wood, under the bark of rotten logs, or recently cut trees; less frequently in urbanized areas at light, camping areas (Nimboy-Binderay National Park), and picnic areas at mv light (near Bungleboori, I.2011). The elevation ranges from 40 to 800 m. Some data on host plants were obtained from labels, namely Argyrodendron actinophyllum (Malvaceae), in Queensland. Other records from Lord Howe Island are Casuarina (Casuarinaceae); Chionanthus (Oleaceae), under bark log; Cryptocarya (Lauraceae); rotting Howea belmoreana (Arecaceae); rotting Olea paniculata (Oleaceae).
Remarks. Concerning this species, Bomans (1989: 14) wrote that " M.O. de Lisle (1967: 542) a décrit cet insecte d’après un exemplaire unique provenant de l’Ile Lord Howe (N.S.W., Australia). J’ai examiné le type conservé au Muséum d’Histoire Naturelle de Genève, et j’ai pu constater qu’effectivement la cuvette oculaire de l’insecte est présente mais entièrement chitinisée. Ce Figulus est donc bien aveugle comme le déclare de Lisle" [M.O. de Lisle (1967: 542) described this insect upon a single specimen from Lord Howe Island (N.S.W., Australia). I examined the type, housed in the Muséum d’Histoire Naturelle in Geneva, and I could verify that eye basin of the insect is indeed present, yet wholly chitinized. Therefore this Figulus is really blind as de Lisle states]. Bomans adds that "le C.S.I.R.O. à Canbérra m’a transmis pour identification une série de Lucanides parmi lesquels j’ai trouvé 3 exemplaires de Figulus howei provenant, comme l’holotype, de l’île Lord Howe. J’ai constaté qu’un seul exemplaire est aveugle, les deux autres sont normaux " [C.S.I.R.O. (Canberra) sent me for identification a series of Lucanidae, among which I found three Figulus howei specimens, coming from Lord Howe Island as the holotype. I verified that only one specimen is blind, the other two are normal].
We examined the holotype from Lord Howe Island and several specimens from the same locality and verified that neither the holotype nor the other specimens seem to be really blind. The cuticle covering the eye may be normal or in various degrees thickened, as in other species of the genus, yet ommatides are always visible at the stereomicroscope. We dissected the male specimen labeled " Type " of F. h ow e i, and compared the genitalia with those of F. nitens. We see no significant differences between F. how ei and F. nitens, both in external and in genitalia characters, except for the variability falling within the individual range. Therefore we consider F. howei as a junior synonym of F. nitens (new synonymy).
The same situation resulted from examination of external morphology and genitalia structures of the male of F. tambourinensis. The characters given by Mjöberg (1916) to distinguish F. tambourinensis from F. nitens as “black head, sculpture of prothorax, and striae of the elytra” are within the range of the individual variability we have seen in F. n i t e ns. Therefore F. tambourinensis is also here considered as a junior synonym of F. nitens (new synonymy).
description: Published as part of Monte, Cinzia, Zilioli, Michele & Bartolozzi, Luca, 2016, Revision of the Australian species of Figulus MacLeay, 1819 (Coleoptera: Lucanidae), pp. 447-484 in Zootaxa 4189 (3) on pages 457-461, DOI: 10.11646/zootaxa.4189.3.2, http://zenodo.org/record/166170