Figulus lilliputanus Westwood, 1855

Monte, Cinzia, Zilioli, Michele & Bartolozzi, Luca, 2016, Revision of the Australian species of Figulus MacLeay, 1819 (Coleoptera: Lucanidae), Zootaxa 4189 (3), pp. 447-484: 453-457

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Figulus lilliputanus Westwood, 1855


Figulus lilliputanus Westwood, 1855  

( Figs. 8–11 View FIGURES 8‒11 )

Figulus lilliputanus Westwood, 1855: 219   ; Thomson 1862: 432; Parry 1863: 450; Parry 1864b: 100; Harold 1868: 965; Macleay 1871: 173; Deyrolle 1874: 414; Parry 1875: 22; Felsche 1898: 78; Van Roon 1910: 52; Mjöberg 1916: 13; Arrow 1938: 53, 60, 61; Arrow 1943: 139; Didier & Séguy 1953: 180; Benesh 1950: 53; Benesh 1955: 64, 66; Benesh 1960: 26; Klausnitzer 1982: 15; Bomans 1986: 9 (lectotype designation); Maes 1992: 36, 38; Moore & Cassis 1992: 10; Araya 2001: 13; Krajcik 2001: 11; Krajcik 2003: 79; Klausnitzer & Sprecher-Uebersax 2008: 29; Fujita 2010: 395; Maes 2014. Figulus clivinoides Thomson, 1862: 402   , 432; synonymised by Parry 1863: 450.

Figulus (Figulinus) lilliputanus: Schoolmeesters 2015   .

Type material. Figulus lilliputanus Westwood   , lectotype (designation by Bomans 1986): “S Austral [handwritten]”, “ Figulus   / Liliputanus (sic!) / Tr. Ent. Soc. n.s. III, pl. 12, fig. 5 [handwritten]”, “TYPE / WESTWOOD / Trans. Ent. / Soc. 1855 / P.219 T.12 f.5 / Coll. Hope Oxon. [printed, handwritten]”, “TYPE COL: 333 1 / 2 / Figulus   / lilliputanus   / Westw. / HOPE DEPT. OXFORD [printed, handwritten]”, [OUMNH]; Figulus lilliputanus Westwood   , paralectotype: “S Austral [handwritten]”, “W [printed]”, “TYPE / WESTWOOD / Trans. Ent. Soc./ 1855 P.219. / T.12 fig.5 / Coll. Hope Oxon. [printed, handwritten]”, “TYPE COL: 333 2 / 2 / Figulus   / lilliputanus   / Westw. / HOPE DEPT. OXFORD [printed, handwritten]”, [OUMNH]; Figulus clivinoides Thomson   : Lectotype here designated: Lectotype: “Th. / Type [printed]”, “ Clivinoides   / Type Thoms / Austr [handwritten]”, “Ex Musaeo / James Thomson [printed]”, “MUSÉUM PARIS / 1952 / COLL. R. OBERTHÜR [printed]”, “H.Bomans det., 1973 / = Figulus   / lilliputanus   West. [printed, handwritten]”, [MNHN].

Other material. Figulus lilliputanus Westwood   (Ƌ): “ Lilliputanus   / West. Ent. J. / v. 3 Feb [handwritten]”, “Ex Musaeo / Parry [printed]”, “MUSÉUM PARIS / 1952 / Coll. R. OBERTHÜR [printed]”, [MNHN]; Figulus lilliputanus Westwood   (Ƌ): “Ex Musaeo / Parry [printed]”, “MUSÉUM PARIS / 1952 / Coll. R. OBERTHÜR [printed]”, [MNHN]; Figulus lilliputanus Westwood   (♀): “ Lilliputanus   / West. / Clivinoides   / Thompson [handwritten]”, “Ex Musaeo / Parry [printed]”, “MUSÉUM PARIS / 1952 / Coll. R. OBERTHÜR [printed]”, [MNHN]. Note: this material was indicated as syntypes by S. Boucher (MNHN) in the loan form, but their status as type material is uncertain, as they have no red label or clear indication of being part of the original type series.

Diagnosis. This is the smallest species of Australian Figulus   and differs from all other Australian species by its antenna with only 9 antennomeres. The irregular transversal rows of dense and deep punctures with short erected setae directed downwards on the ventrites 2‒4 are secondary sexual characters, as in other Figulus   species ( Bomans 1989).

Redescription. Measurements: size range (n = 165): TL: 6.0–10.0 mm; PL: 1.6 1–2.49 mm; EL: 3.41–5.43 mm; PW: 1.69–2.97 mm; EW: 1.74–3.0 mm.

Male ( Fig. 8 View FIGURES 8‒11 ). Body elongate, with subparallel sides, dark brown or reddish brown, glossy.

Head: transverse, about twice as wide as long. Frontoclypeal region slightly concave at the anterior margin coarsely punctate with an evident median elevation and two large, glossy areas situated before the eyes. Labrum broad, short, with anterior margin slightly bilobed and with long, golden setae. Vertex slightly concave in the middle, feebly inclined anteriorly, with two small, circular elevations. Two longitudinal, slightly elevated carinae are present along the inner margin of eyes. Eyes small, divided into 2 parts by the canthi, the upper part smaller than the lower with distinct facets. Canthus slightly protruding, rounded laterally and angled behind; upper surface slightly depressed and smooth, with sparse, minute punctures on the lateral margin. Head surface densely punctate with superficial, large punctures smallest on canthi, anterior margin of the frons, longitudinal carinae and transversal elevations of vertex. Mandibles shorter than head, rounded laterally, apices acute; upper surface with external margin slightly raised. Internal margin of mandibles with a medial strong tooth, with apex blunted directed inwards. Antenna with 9 antennomeres, with scape as long as the remaining antennomeres combined; club of last three antennomeres, which are flatter and broader than the remaining antennomeres of the funicle. Mentum subsquare ( Fig. 9 View FIGURES 8‒11 ), slightly concave medially; anterior margin slightly bilobed; base smooth inclined forwards, with sides slightly elevated at mid-length. Surface smooth, with mixed large punctures and wrinkles.

Thorax: pronotum subsquare with subparallel sides; posterior width less than anterior width; anterior margin almost straight, with a small, blunt median tubercle; posterior margin slightly bisinuate; anterior angles produced with anterior margin rounded; posterior angles largely rounded, with slightly crenulate margin. Superficial longitudinal medial groove on pronotal disc, with some large, circular punctures. Pronotal surface glossy with dense, small, superficial punctures, smaller along the anterior margin of prothorax and with patch of large, close punctures at sides. Long, golden setae directed upwards along the lateral pronotal margin. Scutellum long, wedgeshaped; surface smooth and glossy. Elytra 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; each stria formed by a row of large punctures well separated from each other, except for the first stria, deeper than others and where the punctures are confluent. Stria 1 the only stria than reaches apex; stria 2 ending slightly before; stria 3 curved outwards, converging with stria 6; striae 4 and 5 converging at declivity. Interstriae much larger than striae, slightly convex and smooth, with a longitudinal, irregular row of small punctures. Elytral margins and apex strongly punctate, with short, erect, red setae. Wings present. Legs: thin and straight. Protibiae with strong, elongate, apical fork, slightly dilated at the point where it articulates a long, strong spur; lateral margin with 3‒4 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; ventral lateral margin of mesotibiae with 2‒3 small teeth. Metatibiae with 4 acute apical teeth, and 2 long apical spurs, 1 larger than the other; lateral margin with strong submedial tooth. Prosternum subpentagonal in shape behind the coxae; surface with small punctures at anterior edge and with smaller punctures medially and at posterior edge. Metasternal disc smooth, with a long medial groove, deep at posterior edge; sides of metasternum with large and dense, horse-shoe shaped, superficial hollows.

Abdomen: first abdominal ventrite with some large and dense, horse-shoe shaped, superficial hollows with a long, golden seta in the middle. Ventrites 2‒4 with some deep longitudinal depressions along the anterior margin, with short, golden setae emerging from the anterior margin; they are not present in the anteromedial area where there are 1‒3 irregular transversal rows of dense and deep punctures, each with a short seta, erected and directed downwards. Last ventrite with some punctures, which are smaller near the posterior margin.

Male genitalia: penis tubular ( Figs. 10–11 View FIGURES 8‒11 ), short and narrow, moderately curved in lateral view, strongly bent proximally close to articulation with basal piece, almost completely covered by a dorsal translucid extension of the basal piece. Permanently everted internal sac drawn out into a narrow colorless or yellow flagellum, slightly longer than the tegmen (basal piece + parameres), with two subsequent forkings in the basal half, tapering towards the apex ( Fig. 10 View FIGURES 8‒11 ). Parameres slightly shorter than basal piece, with marginal fringe of setae entirely enclosed in the cuticle (not represented in figure); ventrally the base of the parameres ends in a long spine-shaped apophysis articulated with basal piece ( Figs. 10–11 View FIGURES 8‒11 ). Basal piece ventrally largely membranous, dorsally sclerotized with a moderately large (about one-half of basal piece length), translucent cuticular area along the midline. Two long, slender struts run along basal piece, not reaching its base.

Female. All external morphological characters as in male, except for sternites 2‒4 smooth and glabrous, without irregular, transverse punctation or short setae.

Female genitalia: hemisternites elongate, broader anteriorly, narrower posteriorly; anteriorly broad up to about half of their length; outer edge notched; medial edge convex throughout length. Bursa copulatrix represented by a slight expansion of the wall of spermathecal duct near junction with the median oviduct. Spermathecal duct short, ending with a curve and long expansion, proximally colorless, distally pigmented.

Intraspecific variation. The specimens examined do not show significant morphological variability, except for the body size (see “Measurements” section).

Additional material examined. Australia [state unknown]: 8 specimens, 29.IX.1929, E.Sutton Collection, donated Dec.1964 ( QMB)   ; 1 specimen, ex Musaeo Ed. Brown, Muséum Paris 1952 Collection Oberthür (MNHN)   ; 1 specimen, ex Musaeo W. Möllenkamp, Muséum Paris 1952 Collection R. Oberthür (MNHN)   ; 2 specimens, det. A.M.Lea (MEL); 5 specimens, Nat. Mus. Victoria, COL-85173, COL-85174, COL-85175, COL-85176, and COL- 8517 ( NMVM)   ; 1 specimen (GTC); 1 specimen, ex Musaeo Mniszech, Muséum Paris 1952 Collection Oberthür (MNHN)   ; 1 specimen, Nouvelle Hollande, R.I.Sc. N.B. 21.418, Collection P. de Moffarts, 4764, Comp. au type, H.E.Bomans BMNH (E) 1999-248, 404764 (handwritten) ( NHM)   . AUSTRALIAN CAPITAL TERRITORY: 1 specimen, Black Mountain , 1.II.1968, light trap, M.S.Upton ( ANIC)   . NEW SOUTH WALES: 1 specimen   , New South Wales, Macleay Museum University of Sydney 2006 Australia, 11384, H.E.Bomans BMNH (E) 1999-248, 413884 (handwritten) ( NHM)   ; 1 specimen, New South Wales, Nat. Mus   . Victoria, COL-85784 ( NMVM)   ; 1 specimen, Albury, 16.II.1970, R.Parrot, Collection R.E. Parrot ( MEL); 1Ƌ, Avoca Station off Pomona road 20 km N of Wentworth   , 13.VII.1993, COL-86865 (NMVM); 1 specimen, Ballimore, F.H.Taylor (ANIC); 1♀, 1 specimen, N Belmore Road, 3.2 km SSW of Busbys Flat , 26.I. 1999, 170 m, at mv light, S.Watkins, S.Watkins Collection donated 2001 ( ANIC)   ; 1Ƌ, Bungabbee State Forest , SE Kyogle, 29.I.1997, mv light, S.G.Watkins, 97:199 Ƌ, S.G.Watkins Collection, donated 2001 ( ANIC); 1 specimen, Como, Dr. K.K. Spence, K.K. Spence Collection, K 293062 (AMS)   ; 1 specimen, Coonabarabran , XII.1929, H.J.Carter ( ANIC)   ; 1♀, Fortis Creek National Park, South boundary, N of Grafton , 4.I. 2000, 50 m, at mv light, S.Watkins Collection donated 2001 ( ANIC)   ; 2ƋƋ, Fortis Creek National Park, South boundary, N of Grafton , 17.XII. 1998, 50 m, mv light, 98:837Ƌ and 98:838 Ƌ, S.Watkins Collection donated 2001 ( ANIC)   ; 2 specimens, Maitland , 26.XII.1962, A Neboiss COL-85777 ( NMVM)   ; 1 specimen, Mogriguy, Goonoo State Forest , 13.II.1981, R.H.M., R.H.Mulder Collection, K 293063 (AMS)   ; 1 specimen, Mullaly , XII.1929, H.J.Carter ( ANIC)   ; 2 specimens, Sydney, Hyde Park , XII.1985, K 293074 (AMS)   ; 2 specimens, Wagga, 1937, C.Oke, COL-85913 (NMVM); 2 specimens, Wagga, 1938, C.Oke, Presented by C.G.Oke 1.IX.1948, COL-85883 ( NMVM)   . QUEENSLAND: 1 specimen   , Queensland, Nat. Mus   . Victoria, COL-85916 ( NMVM); 1 specimen   , Queensland, Nouvelle Hollande, Fry Collection 1905-100, BMNH (E) # 607642 ( NHM)   ; 1 specimen, Blackdown Tableland, via Dingo, XII.1978, Maywald & Czechura ( QMB); 1Ƌ, Boggomoss 15 in field, Taroom District , 25°28'S, 150°01'E GoogleMaps   , 12.XI.1996, P.Lawless, 191, QM Reg. No T 34260 (QMB); 1Ƌ, Boggomoss number 8 via Taroom, 25°27'S, 150°02'E, 12.XI.1996, Queensland Mus. survey, day hand collection, 0 39, QM Reg. No T 34259 View Materials ( QMB)   ; 1Ƌ, 2 specimens, Boyne Island, via Gladstone, 23°55'S, 151°20'E, 1995, C.Q. University, 5022 ( QMB) GoogleMaps   ; 1Ƌ, Bribie Island, XII.1925, H.Hacker ( QMB); 1 specimen, Brisbane   , 29.X.1917 and 1 specimen, 4.III.1920, H.Hacker (QMB); 2 specimens, Brisbane (QMB); 2 specimens, Brisbane, Mount Coot-tha , I.1966, C.Oke COL-85890 ( NMVM)   ; 1 specimen, Cape York, 3851 ( QMB)   ; 6 specimens, Clermont , XII.1929, Dr. K.K. Spence, K.K. Spence Collection, K 293070 K 293071, and K 293072 (AMS)   ; 2 specimens, Cooktown , I.1971, J.G.Brooks, J.G.Brooks Bequest 1976 ( ANIC)   ; 1 specimen, Dawson District , Barnard Collection, Muséum Paris 1952 Collection Oberthür (MNHN)   ; 1 specimen, Dawson River , Macleay Museum University of Sydney 2006, 11383 (handwritten), H.E.Bomans 1999-248, 411383 ( NHM)   ; 1Ƌ, 2♀♀, Dunmore State Forest , 27°38'S, 150°58'E, 26.XII.1998, 98:883 Ƌ, mv light, S.Watkins Collection donated 2001 ( ANIC) GoogleMaps   ; 1 specimen, Garradunga Stone Creek , 15.I.1997, J.Hasenpusch, R.deKeyzer Insect Collection, K 293325 (AMS)   ; 1 specimen, 15 km S of Greenvale , 9.I.1911, B.B.Howery ( ANIC); 1Ƌ, Gurgeena Plateau, 25°27'S, 151°23'E GoogleMaps   , 8.VI. 1999, 360 m, open forest, 7741, G.B.Monteith & G.I.Thompson ( QMB); 1 specimen, Hann River , 15°11'S, 143°52'E (GPS) GoogleMaps   , 18.XII.1993 ‒ 14.I.1994, flight intercept trap, P.Zborowski, E.D. Edwards ( ANIC); 1 specimen, 14 km ENE Heathlands , 11°41'S, 142°42'E GoogleMaps   , 8.XII.1992, at light rainforest, P. Zoborowski & W.Dressler (ANIC); 1 specimen, Iron Range, W Claudie Rg, 12°45'S, 143°14'E, 3.XII. 1985, 50 m, rainforest, stick brushing, G.B.Monteith, QM Berlesate No 690 (QMB); 1 specimen, same locality, 3.XII. 1985, 50 m, rainforest, Monteith & Cook ( QMB) GoogleMaps   ; 1 specimen, Iron Range National Park, Gordon Creek ( QMB)   ; 1 specimen, Iron Range , 28.X.1981, J.Hasenpusch, R.deKeyzer Insect Collection, K 293323 (AMS); 1Ƌ, Iron Range, Cape York Peninsula   , VI ‒VII.1977, G.B.Monteith (QMB); 1 specimen, Kuranda , 13.I.1987, R.deKeyzer Insect Collection, K 293322 (AMS)   ; 1 specimen, Maryborough, E.W.Fischer, ex Collection Australian Museum (ANIC); 2 specimens, Moreton Bay , K 15076 View Materials (handwritten), K 293064 (AMS)   ; 1 specimen, Moreton Bay , ex Musaeo Ed. Brown, Muséum Paris 1952 Collection Oberthür (MNHN)   ; 1 specimen, Nipps, COL-85915 (NMVM); 1 specimen, Normanton , II.1982, R.deKeyzer Insect Collection, K 293324 (AMS)   ; 1 specimen, Rockhampton, ex Musaeo Ed. Brown, Muséum Paris 1952 Collection Oberthür (MNHN)   ; 1 specimen, Rocky River, Silver Plains , E Coast   , Cape York Peninsula, 6.I.1960, J.L.Wassel ( ANIC)   ; 2 specimens, Stanthorpe, E.Sutton, F.E.Wilson Collection, COL-85885 (NMVM); 5 specimens, Stanthorpe , 29.VII.‒4.IX.1924, E.Sutton Collection, donated Dec.1964 ( QMB)   ; 2 specimens, Station Creek , N Queensland, 30.XII.1964, J.G.Brooks ( ANIC)   ; 1Ƌ, 2 specimens, Station Creek , 30.XII.1969, J.G.Brooks, J.G.Brooks Bequest 1976 ( ANIC)   ; 1 specimen, Toomba , 19°58'S, 145°34'E, 14‒16.XII.2007, Site 1, 390 m, 14652, mv light, vinescrub edge/paddock, Monteith & Wright, QM Reg. No T147162 ( QMB) GoogleMaps   ; 1 specimen, Townsville, K.K.Spence, K.K.Spence Collection, K 293069 (AMS); 3 specimens, 9 km ENE of Mount Tozer , 12°43'S, 143°17'E, 5‒10.VII.1986, rotten log, T.Weir & A.Calder ( ANIC) GoogleMaps   ; 1Ƌ, Woodstock , 3.VII.1992, B.P.Moore ( LBC)   . SOUTH AUSTRALIA: 2 specimens   , South Australia, ex Macleay Museum University of Sydney 2006 Australia ( GTC, LBC)   ; 1Ƌ, South Australia, R. Mus. Hist. Nat. Belg., I.G. 12.595 ( IRSNB)   ; 1 specimen, Adelaide, BMNH(E) # 607646 (NHM); 2 specimens, Adelaide Hills , V.1930, C.Oke, Presented by C.G.Oke 1.IX.1948, COL- 85892 ( NMVM)   ; 1 specimen, Adelaide, ex Musaeo Ed. Brown, Muséum Paris 1952 Collection Oberthür (MNHN)   ; 1 specimen, Adelaide, Griffith 0 4 (ANIC); 1 specimen, Adelaide, K 28038 (handwritten), K 293073 (AMS); 1 specimen, Barossa, A.H.Elston, A.H.Elston Collection, K 293075 (AMS); 1 specimen, Buckaringa Gorge , Flinders Ra. C. 30 km NNW of Quorn, 19.XII.1985, Reid, Gullan & Lewis, at light trap ( ANIC)   . VICTORIA: 4ƋƋ, Ballarat , VIII.1937, C.Oke, Presented by C.G.Oke 1.IX.1948, COL-85875 and COL-85884 ( NMVM)   ; 1Ƌ, 1♀, Hattah, C.Oke COL-85914 (NMVM); 2 specimens, Lake Hattah , XI.1924, C.Oke, COL-85785 and COL-85786 ( NMVM)   ; 1 specimen, Mount Hope , 27.IX.1947, C.Oke, Presented by C.G.Oke 1.IX.1948, COL-85881 ( NMVM)   ; 2 specimens, Piangil , 20.VII.1919, C.Oke, COL-85912 ( NMVM)   ; 1 specimen, Sea Lake , III.1912, J.C. G., COL- 85894 ( NMVM)   ; 1 specimen, Sea Lake , Goudie, 583, COL-85901 ( NMVM)   . WESTERN AUSTRALIA: 1 specimen, Bakers Hill   , 27.III.1973, light trap, D.Weeldenburg (ANIC); 4 specimens, Blackwood River ( ANIC)   ; 2 specimens, Bridgetown, L.J.Newman, F.E.Wilson Collection, COL-85850 (NMVM); 1 specimen, Champion Bay, ex Collection V-de-P-, Sig. K. vV. Steinwehr Dec./ Nov.1935, Museum Koenig Bonn, 5368, H.E.Bomans BMNH (E) 1999-248, 405368 (handwritten) ( NHM)   ; 3ƋƋ, Gingin, H.J.Carter Collection P. 20.4.22., COL-85891 (NMVM); 1 specimen, Gray River , 6.III.1983, at light, K. & E.Carnaby ( ANIC)   ; 1 specimen, 2 mi. NW Margaret River , 23.XII.1970, mv lamp, G.A.Holloway & H.Hughes, K 293085 (AMS)   ; 1 specimen, Nannup , 26.XII.2004, D.Lord, R.deKeyzer Insect Collection, K 293326 (AMS); 1Ƌ, Pipehead Dam, 15 miles SSE of Armadale   , 26.I.1967, M.S.Upton (ANIC); 3 specimens, Wilga, I.1973 and 1 specimen, 21.IV.1973, K. & Carnaby (ANIC).

Distribution. Australia: Australian Capital Territory, New South Wales, Queensland, Western Australia, Victoria, South Australia. The species is widely distributed in Australia, with the exception of the Northern Territory.

Natural history data. According to label data, adults were collected thoughout the year, mostly from December to April. Most of them were collected at light trap with mercury vapor light; a single specimen was captured in a flight intercept trap (Hann River, XII.1993 – I.1994). Few other specimens were found in rotten logs, in vine scrub. The elevation ranges from 50 to 390 m. The larvae and host plant(s) are unknown.

Remarks. Concerning Figulus lilliputanus Westwood   and Figulus clivinoides Thomson, Bomans (1986: 9)   wrote that: “- Il n’est pas absolument certain que ces deux insects soient synonymes. L’examen des types permet de constater que le premier est nettament plus petit que le second, 6,2 et 7,5 mm pour 8 mm. De plus lilliputanus   a été découvert en Australie méridionale et occidentale, et clivinoides   ou du moins l’espèce la plus grande, en Nouvelle Galle du Sud et au Queensland ” [It is not sure that the two species are synonyms. Examination of types shows that the first is clearly smaller than the second, 6.2 and 7.5 mm instead of 8 mm. Moreover F. lilliputanus   has been discovered in southern and western Australia, and F. clivinoides   , or at least the largest species, in New South Wales and Queensland]. We dissected the male labeled "Type" of F. clivinoides   , and we compared the genitalia with those of F. lilliputanus   . We do not see any significant difference between F. clivinoides   and F. lilliputanus   , both in external morphology and in genitalia characters (e.g., in the flagellum shape). Therefore we consider F. clivinoides   as a junior synonym of F. lilliputanus   (new synonymy).

We found in the collection of NHM a specimen labeled “Tasmania” (A.Simson leg., 8116, 4765, H.E.Bomans), but this locality is most probably erroneous, since we have never seen any other specimens of this species from Tasmania.


Queensland Museum, Brisbane


University of Nottingham


Australian National Insect Collection


Museo Entomologico de Leon


University of the Philippines at Los Ba�os


Gifu Type Culture Collection


Institut Royal des Sciences Naturelles de Belgique














Figulus lilliputanus Westwood, 1855

Monte, Cinzia, Zilioli, Michele & Bartolozzi, Luca 2016

Figulus lilliputanus

Fujita 2010: 395
Klausnitzer 2008: 29
Krajcik 2003: 79
Araya 2001: 13
Krajcik 2001: 11
Maes 1992: 36
Moore 1992: 10
Bomans 1986: 9
Klausnitzer 1982: 15
Benesh 1960: 26
Benesh 1955: 64
Didier 1953: 180
Benesh 1950: 53
Arrow 1943: 139
Mjoberg 1916: 13
Van 1910: 52
Felsche 1898: 78
Parry 1875: 22
Deyrolle 1874: 414
Macleay 1871: 173
Harold 1868: 965
Parry 1864: 100
Parry 1863: 450
Parry 1863: 450
Thomson 1862: 432
Thomson 1862: 402
Westwood 1855: 219