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ESPINASA, LUIS, SPROUS, PETER, POSSO, KARA, MITCHELL, ANDREW, ESPINASA, MONIKA, LIN, JONATHAN (2023): LUIS ESPINASA, PETER SPROUS, KARA POSSO, ANDREW MITCHELL MONIKA ESPINASA & JONATHAN LIN (2023) Miocene divergence for Texoreddellia? An important component of the cave-adapted fauna of Texas and northern Mexico. Zootaxa, 5256 (3): 267-278. Zootaxa 5296 (4): 600-600, DOI: https://doi.org/10.11646/zootaxa.5296.4.10, URL: http://dx.doi.org/10.11646/zootaxa.5296.4.10

Nososticta chrismulleri Theischinger & Richards, 2016 (Figs 24–29, 63, Map 4, Pl. 3) Nososticta chrismulleri Theischinger & Richards, 2016: 309. Material examined. Holotype ♁, Papua New Guinea, Western Province, Palmer River Catchment, upper Fly River basin, Camp 4 (5°54.477’S, 141°50.773’E; 125 m a.s.l.), 02-viii-2013, S.J. Richards (SAMA 07-001449); 2 ♁♁, same data as holotype (SAMA 07-001592 - 93). Affinities. Based on the present study (Fig. 6), N. chrismulleri is sister group to the remaining taxa of the Western group of species (N. tagula sp. nov., N. stueberi sp. nov. and N. boonei sp. nov.). Differential diagnosis. Nososticta chrismulleri was compared to, and distinguished from, N. salomonis by Theischinger & Richards (2016) as follows: “by the spotted anterior lobe of the pronotum, the markedly longer blue ante-humeral patch, and the lack of a step in the blue mesepimeral patch.” Nososticta chrismulleri is a mainland (New Guinea) species as opposed to the almost exclusively insular species N. africana, N. tagula sp. nov., N. hedigeri sp. nov. and N. salomonis. Whereas the lack of a pale dorsal marking on segment 2 distinguishes the male of N. chrismulleri from N. africana and N. tagula sp. nov. and the lack of a long black basal stalk between the ante-hu meral patches distinguishes it from N. hedigeri sp. nov., an anterodorsally convex metepimeral patch separates the male of N. chrismulleri from N. salomonis (metepimeral patch anterodorsally straight). The male of N. chrismulleri from Western Province is most similar to N. boonei sp. nov. from Gulf Province. It can be distinguished from that species by the following characters: inner and apical margin of the blue ante-humeral patch and the anterior margin of the blue metepimeral patch are more curved/sinuous in male N. chrismulleri than in N. boonei sp. nov.; and blue markings on S7 are lacking in N. chrismulleri, present in N. boonei sp. nov. Nososticta chrismulleri can be separated from N. stueberi sp. nov. by the markedly longer ante-humeral patch and the markedly darker poststernum. Distribution and habitats. Nososticta chrismulleri is currently known only from the lowlands of Western Province, Papua New Guinea (Map 4, Pl. 3), where males perched on low foliage in sunny patches along small, clear-flowing streams in primary lowland forest., Published as part of Theischinger, Günther, Mitchell, Andrew, Richards, Stephen J. & Polhemus, Dan A., 2023, Systematics of the Nososticta salomonis complex (Odonata: Zygoptera: Platycnemididae), pp. 101-146 in Zootaxa 5296 (2) on pages 115-120, DOI: 10.11646/zootaxa.5296.2.1, http://zenodo.org/record/7978135, {"references":["Theischinger, G. & Richards, S. J. (2016) Six new species of Nososticta Hagen, 1860 from Papua New Guinea (Odonata: Platycnemididae). Odonatologica, 45, 291 - 316."]}

Nososticta hedigeri sp. nov. (Figs 34–41, 59, 60, Map 1, Pl. 4) Holotype. ♁, SAMA 07-001675, Papua New Guinea, Bougainville, Pipikei (5.8077° S, 154.8067° E; 240 m a.s.l.), 17–19-x-2019, S. Richards; deposited in the South Australian Museum, Adelaide, Australia. Paratypes. 2 ♁♁, 3 ♀♀, SAMA 07-001673, 07-001676, 07-001674, 07-001677, 07-001678, same data as holotype; deposited in the South Australian Museum, Adelaide, Australia. 2 ♁♁ (No. 313), RMNH.INS.1153719, 1153720, S Bougainville, Buin, 1930, H. Hediger; supposedly in Basel, but sent from RMNH; 1♁, 1♀, RMNH. INS.1153721, 1153722, Bougainville, S. Nasiooi, Agric. Field St., 19-v-1976, HR Wimmer ex USNM; deposited in Naturalis Biodiversity Center Leiden, Netherlands. Etymology. The species is dedicated to its probable discoverer, Heini Hediger (1908–1992), a Swiss biologist noted for work in proxemics in animal behaviour and known as the "father of zoo biology". It is a noun in the genitive case. Description of holotype Head. Largely black; only base of labium pale grey and a transverse frontal bar from eye to eye blue; an ill-defined pale brown mark only barely detectable between antennal base and lateral ocellus. Thorax. Pronotum black, anterior lobe with one blue lateral patch on one side, two on the other, smaller inner patch separate; much larger outer patch connected to large patch of same colour on median lobe; propleura largely blue, some black anteriorly, along part of sutures and at rear of epimeron. Synthorax with pleura largely black; blue restricted to: ante-humeral patch, distinctly longer than ½ the length of mesanepisternum, drop-shaped with inner margin sloping from long wide base to humeral suture, leaving black area between two ante-humeral patches approximately cup-shaped with distinct narrow basal stalk; patch covering dorsal 4/5 length, rarely more, and almost entire width of mesepimeron; partly connected with it a stripe across whole length and anterior 3/4 of metepisternum; patch across most of metepimeron, subbasally to basally shortened by black along metapleural suture extending to poststernum, its anterior margin almost straight; posterolateral tip of meso- and metakatepisternum. Postcoxae blackish brown to black and blue; poststernum pale blue with separate narrow elongate ill-defined patches of blackish brown to black. Legs: Coxae largely blue, anterobasally brown to black, trochanters black and blue, legs otherwise black, with only base of femora, longest in metafemur, pale bluish grey. Wings: Membrane with slight greenish brown tinge, venation black; pterostigma black, slightly more than twice as long as wide; postnodals 16–17/13. Abdomen. S1 dorsally black, laterally largely blue; S2 largely black, blue adjacent to genitalia but blue dorsal mark absent; S3–7 dorsally black, ventrally narrowly margined greyish brown, S3–6 with blue mid-dorsal spot very close to base; S8 largely blue, narrowly lined black basally, more widely and irregularly apically, also a very fine black midline over part of the blue patch; S9 and S10 black. Superior anal appendages largely blue, inferiors brownish black. Dimensions. Hind wing 19.1 mm; abdomen (including anal appendages) 33.2 mm. Description of female Head. Much as in male, but dark structures and areas darker, pale areas largely pale yellow to very pale blue; brownish mark between antennal base and lateral ocellus more distinct. Thorax. Pronotum much as in male, but anterior lobe with larger outer mark only, bluish white and connected to large patch of same colour on median lobe; posterior lobe with vertical dorsal and horizontal ventral lobe; propleura much as in male but pale areas largely bluish white instead of blue. Synthorax with pleura largely black; pale yellow to whitish blue areas include: ante-humeral patch, less than 40% length of mesanepisternum, comma-shaped; mesepimeral patch reduced to tiny remnant, completely fused with stripe across whole length of and anterior 2/3 to 3/4 of metepisternum; a subtriangular to subtrapezoidal patch across most of metepimeron; posterolateral tip of meso- and metakatepisternum. Postcoxae blackish brown to black and pale greyish yellow; poststernum pale greyish yellow with largely narrow elongate patches of dark greyish brown to black. Legs: coxae largely pale to brownish yellow, anterobasally brown to black, trochanters black and pale brownish yellow; legs otherwise dark, with meso- and metafemur partly to largely greyish to pale brownish yellow and tibiae, except for apical portion, brightened to pale and darker yellowish brown. Wings: membrane with very slight greenish brown tinge, venation black; pterostigma black, slightly more than twice as long as wide; postnodals 15–18/12–15. Abdomen. S1 and S2 dorsally largely black, laterally largely pale blue; S3–7 dorsally black, ventrally narrowly margined pale greyish yellow, S3 and S4, sometimes also S5, with very short pale blue dorsal mark very close to base and most commonly divided into two spots by dark midline; S8 black with ventral edge yellow; S9 and S10 black. Anal appendages yellow, inferior anal lamina brown and yellow; ovipositor dull yellow, valves (about 12 teeth) brown to black; supra-anal plate black. Dimensions. Hind wing 17.5–20.5 mm; abdomen 26.2–31.5 mm. Variation in the male paratypes. Inner margin of dorsal region of ante-humeral patch varying between almost straight and sloping and widely or irregularly curved toward humeral suture; black elongate median and lateral patch of poststernum separate or narrowly connected; postnodals 15–17/12–14; hind wing 18.6–20.4 mm; abdomen (including anal appendages) 28.5–34.2 mm. Affinities. Based on the present study (Fig. 6), N. hedigeri sp. nov. is considered sister species to N. salomonis. Differential diagnosis. Nososticta hedigeri sp. nov. can be distinguished from all other species of the N. salomonis complex by having the male ante-humeral patches extending very close to the mid-dorsal carina basally, leaving only a narrow black stalk between them, versus a much wider black stalk, and the very short female ante-humeral patches extending slightly inward basally making them appear comma-shaped, versus more parallel sided and often also markedly longer. Distribution and habitats. This species is currently known only from Bougainville Island at the western edge of the Solomon Islands Archipelago (Map 1, Pl. 4), where it is common along clear-flowing streams in primary and partially degraded lowland and hill forest.

Nososticta tagula sp. nov. ( Figs 50–54, 56, 58, Map 3, Pl. 6) Holotype. ♁, BPBMENT 0000081204, Papua New Guinea, Milne Bay Prov., Louisiade Archipelago, Tagula Is., Kalitau Creek, above estuary, 0–30 m a.s.l., 11°21’52”S, 153°14’26”E; 29-viii-2002, CL 7190, D.A. & J. T. Polhemus (CL 7190); deposited in the Bishop Museum, Honolulu, Hawaii. Paratypes. 6 ♁♁, 1 ♀, BPBMENT 0000081205–81211, same data as holotype; deposited in the Bishop Museum, Honolulu, Hawaii. Etymology. Tagula after the type locality Tagula Island in the Louisiade Archipelago; it is treated as a noun in apposition to the generic name. Description of holotype (all available specimens (type series) apparently turned pale by the way of preservation (ethanol), meaning that in particular the larger now pale greyish blue areas would have been largely bright blue in life). Head. Largely black; base of labium pale brownish yellow, anteclypeus greyish yellow marbled with grey, pleurostoma and transverse frontal bar from eye to eye very pale greyish blue, an ill-defined mark between antennal base and lateral ocellus pale brown as are apex of largely blackish brown scape and pedicel, antennal flagellum dark brown. Thorax. Pronotum black, anterior lobe with two pale greyish yellow to bluish lateral spots on each side, the larger inner more yellowish spot detached and smaller outer more bluish spot only marginally separated from large mark of same colour on median lobe; propleura largely pale greyish blue, black along sutures and at rear of epimeron. Synthorax with pleura largely black except following pale greyish blue areas: ante-humeral patch, very slightly longer than 1/2 length of mesanepisternum, approximately twice as wide basally as dorsally, tip evenly rounded; patch covering about dorsal 4/5 length of mesepimeron, partly connected with stripe across whole length and anterior 3/4 to 4/5 of metepisternum; a subtriangular narrow-tailed patch across posterior half of metepimeron and posterolateral tip of meso- and metakatepisternum. Postcoxae blackish brown to black and pale greyish blue; poststernum pale brownish blue with ill-defined small, narrow patches of greyish brown. Legs: Coxae largely pale greyish blue, meso-and metacoxa anterobasally brown to black, trochanters black and pale greyish blue, legs otherwise black, with only very base of femora and part of basal 1/2 to 2/3 of metafemur pale greyish blue. Wings: membrane almost clear, venation black; pterostigma greyish brown, slightly more than twice as long as wide; postnodals 16/13–14. Abdomen. S1 dorsally largely black, laterally pale greyish blue; S2 largely black, with distinct pale greyish blue apical dorsal mark, and pale greyish blue adjacent to genitalia; S3–7 dorsally black with blue basal mid-dorsal mark, the one on S7 very short, all ventrally margined greyish brown to dull yellow; S8 largely blue, narrowly lined black basally, more widely and irregularly lined black apically; S9 and S10 black. Superior anal appendages largely blue, inferiors brownish grey to brownish black. Dimensions. Hind wing 18.8 mm; abdomen (including anal appendages) 31.3 mm. Description of female Head. Much as in male, but dark structures and areas darker, pale areas largely pale greyish yellow instead of pale greyish blue; no pale mark between antennal base and lateral ocellus detectable. Thorax. Pronotum much as in male, but anterior lobe with ill-defined smaller outer mark only, vaguely connected to large mark of same colour on median lobe, posterior lobe with vertical dorsal and horizontal ventral lobe; propleura much as in male but pale areas largely pale greyish yellow instead of pale greyish blue. Synthorax with pleura largely black; pale greyish yellow areas restricted to: ante-humeral patch, very slightly longer than 1/2 length of mesanepisternum, very narrowly almond-shaped to almost parallel sided; mesepimeral patch reduced to tiny remnant completely fused with stripe across whole length and anterior 3/4 to 4/5 of metepisternum; subtriangular, narrow-tailed patch across posterior half of metepimeron; posterolateral tip of meso- and metakatepisternum. Postcoxae blackish brown to black and pale greyish yellow; poststernum pale greyish yellow with narrow elongate patches of dark greyish brown. Legs: Coxae largely pale greyish yellow, anterobasally brown to black, trochanters black and pale greyish yellow, legs otherwise black, with only very base of femora and part of basal 3/4 of metafemur pale greyish yellow. Wings: membrane rather clear, venation black; pterostigma black; postnodals 16/13–14. Abdomen. S1 and S2 dorsally black, laterally largely pale greyish yellow; S3–7 dorsally black with greyish yellow dorsal mark very close to base, very tiny on S7, ventrally margined greyish brown to dull yellow; S8 black, ventrally margined greyish brown to dull yellow; S9 and S10 black. Anal appendages and inferior anal lamina pale yellow; supra-anal plate and valves brown to black. Dimensions. Hind wing 19.5 mm; abdomen 29.3 mm. Variation in the male paratypes. Small pale spots on anterior lobe of pronotum present or absent. Postnodals 15–16/12–14. Hind wing 18.1–19.4 mm; abdomen (including anal appendages) 30.3–31.5 mm. Affinities. Based on the present study (Fig. 6), N. tagula sp. nov. is closely allied with N. stueberi sp. nov. and N. boonei sp. nov., together these species comprise the sister group of N. chrismulleri. Differential diagnosis. Among members of the N. salomonis complex, only N. africana and N. tagula sp. nov. have abdominal segment 2 dorsally marked with blue. Nososticta africana can be distinguished from N. tagula sp. nov. by having one or two blue antero-dorsal marks and one postero-dorsal mark on segment 2 (vs. only a blue postero-dorsal mark). Nososticta africana is known almost exclusively from the Bismarck Archipelago, while N. tagula sp. nov. is known only from the Louisiade Archipelago. Distribution and habitats. Nososticta tagula is currently known only from Tagula Island in the Louisiade Archipelago (Map 3). It has not been detected on other Milne Bay Islands despite extensive surveys across the region and may be range-restricted Tagula Island endemic., Published as part of Theischinger, Günther, Mitchell, Andrew, Richards, Stephen J. & Polhemus, Dan A., 2023, Systematics of the Nososticta salomonis complex (Odonata: Zygoptera: Platycnemididae), pp. 101-146 in Zootaxa 5296 (2) on pages 128-129, DOI: 10.11646/zootaxa.5296.2.1, http://zenodo.org/record/7978135

Nososticta boonei sp. nov. (Figs 30–33, 64, 65, Map 4, Pls 3, 9) Holotype. ♁, BPBMENT 000008199, Papua New Guinea, Gulf Province, Isaba Creek off Sirebi River, SE of Omo, ~ 40 km N of Kikori, 40 m a.s.l., 7°12’48”S, 144°15’51”E, water temp. 27° C, 2-iii-1995, CL 7005, D.A. Polhemus; deposited in the Bishop Museum, Honolulu, Hawaii. Paratypes. 2 ♁♁, 2 ♀♀, BPBMENT 0000081200–81203, 4 ♁♁, 1♀, BPBMENT 2008035011-35015, same data as holotype; deposited in the Bishop Museum, Honolulu, Hawaii. 2 ♁♁ (SAMA 07-001690 – 91), 1 ♀ (SAMA 07-001692), Papua New Guinea, Gulf Province, Wau Creek, (7.1216°S, 144.3739°E; 20 m a.s.l.), 22–25-iv-2017, S. Richards; deposited in the South Australian Museum, Adelaide, Australia. Etymology. Nososticta boonei sp. nov. is dedicated to the late Jim Boone (1942–2021), Bishop Museum collection manager, who took excellent care of preparing and exporting odonate material including this species for the present study. It is a noun in the genitive case. Description of holotype. Note: Some available specimens, including the holotype, apparently turned paler by the way of preservation (ethanol), meaning that in particular the larger now pale greyish blue areas would have been largely bright blue in life (see Pl 9 for the colour in dried specimens). Head. Largely black; only base of labium pale grey and a transverse frontal bar from eye to eye pale greyish blue. Thorax. Pronotum black with larger whitish blue outer mark on anterior lobe connected to large whitish blue lateral patch on median lobe, a smaller inner mark separate; pleura largely whitish blue, black along suture and at rear of epimeron. Synthorax: pleura largely black with the following pale greyish blue: ante-humeral patch, very slightly longer than 1/2 the length of mesanepisternum, basally approximately twice as wide as dorsally, inner and apical margin almost straight (tip truncate); a patch covering about dorsal 4/5 length of mesepimeron, and partly connected with stripe across whole length and anterior 4/5 of metepisternum, and patch across approximately postero-dorsal 3/4 of metepimeron. Postcoxae largely blackish brown to black; poststernum bluish white with ill-defined patches of blackish brown to black. Legs: Coxae and trochanters bluish grey and black, legs otherwise black, with only extreme base and inner portion of basal 1/2 of metafemur pale bluish to pale and brownish yellow. Wings: Membrane with greenish brown tinge, venation black; pterostigma black, hardly more than twice as long as wide; postnodals 18/14. Abdomen. S1 dorsally black, laterally largely pale bluish; S2 largely black, pale bluish to greyish yellow adjacent to genitalia; S3–7 dorsally black, ventrally margined greyish brown to dull yellow, S3–6 with blue mid-dorsal mark, about as long as wide, very close to base; S7 with pair of small not very well-defined bluish antero-dorsal spots narrowly connected across mid-line, followed by vaguely detectable diffusely bluish area; S8 largely blue, narrowly lined black basally, more widely and irregularly apically; S9 and S10 black. Anal appendages: Superiors largely blue, inferiors brownish black. Dimensions. Hind wing 19.2 mm; abdomen (including anal appendages) 32.7 mm. Description of female Head. Much as in male, but dark structures and areas darker, pale areas largely pale greyish yellow instead of pale greyish blue. Thorax. Pronotum much as in male but anterior lobe with larger outer mark only, which is barely connected to large mark of same colour on median lobe, posterior lobe with vertical dorsal and horizontal ventral lobe; propleura much as in male but pale areas largely pale greyish yellow instead of pale greyish blue. Synthorax: pleura largely black with pale greyish yellow as follows: ante-humeral patch, very slightly longer than 1/2 the length of mesanepisternum, very narrowly almond-shaped to almost parallel sided, 1/5 to 1/4 as wide as long; a tiny remnant of a mesepimeral patch completely fused with stripe across whole length and anterior 2/3 to 3/4 of metepisternum; subtriangular patch across approximately postero-dorsal half of metepimeron continuing narrowly along posteroventral half of it; extreme posterolateral tip of meso- and metakatepisternum. Postcoxae blackish brown to black and pale greyish yellow; poststernum pale greyish yellow with narrow elongate patches of dark greyish brown. Legs: Coxae largely pale greyish yellow, anterobasally brown to black, trochanters black and pale greyish yellow, legs otherwise black, with only very base of femora and part of basal 3/4 of metafemur pale greyish yellow. Wings: Membrane rather clear, venation black; pterostigma black, slightly more than twice as long as wide; postnodals 16/13–14. Abdomen. S1 and S2 dorsally black, laterally largely pale greyish yellow; S3–7 dorsally black, ventrally margined greyish brown to dull yellow, a pale greyish blue dorsal mark very close to base, approximately semicircular on S3– 6, more elongate, between 1/3 to 1/2 the length of segment, on S7; S8 black, with or without 2 tiny greyish yellow spots on each side, ventrally margined dull yellow; S9 and S10 black. Anal appendages pale greyish blue, inferior anal lamina brown; supra-anal plate and valves brown to black; ovipositor greyish brown with tip greyish yellow. Dimensions. Hind wing 19.6–20.0 mm; abdomen 29.5–30.5 mm. Variation in the male paratypes. Postnodals 15–17/13–14. S7 with pair of small ill- to well-defined, separated antero-dorsal spots, posterior to them a diffusely bluish area may or may not be detectable. Hind wing 19.2–19.3 mm; abdomen (including anal appendages) 31.5–33.0 mm. Affinities. Analysis of the two nuclear genes suggest that N. boonei sp. nov. is closely related to, yet distinct from, N. chrismulleri, in close agreement with the morphological evidence. The COI gene, in contrast, places N. boonei sp. nov. as closest to N. stueberi sp. nov. Differential diagnosis. Nososticta boonei sp. nov. is a mainland (New Guinea) species as opposed to the almost exclusively insular species N. africana, N. tagula sp. nov., N. hedigeri sp. nov. and N. salomonis. The lack of a pale dorsal marking on segment 2 distinguishes the male of N. boonei sp. nov. from N. africana and N. tagula sp. nov., and the lack of a long black basal stalk between the ante-humeral patches distinguishes it from N. hedigeri sp. nov. The presence of a blue dorsal pattern element on S7 (vs. absent) separates N. boonei sp. nov. from N. salomonis. The male of N. boonei sp. nov. from Gulf Province is morphologically and genetically most similar to N. chrismulleri from Western Province. It can be distinguished from that species by having the inner and apical margin of the blue ante-humeral patch and the anterior margin of the blue metepimeral patch in males almost straight (vs. curved/sinuous in N. chrismulleri). Nososticta boonei sp. nov. can be separated from N. stueberi sp. nov. by the markedly longer ante-humeral stripe and by the presence of a blue dorsal pattern element on S7 (vs. absent). Distribution and habitats. Nososticta boonei is currently known only from lowland and foothill forests in the Kikori River basin in Gulf Province, southern-central Papua New Guinea (Map 4, Pl. 3) where adults perch on twigs and low foliage in sunny patches along small to wide (> 5 m), clear-flowing rocky and sandy streams. Material from the adjacent Purari River basin also probably belongs to this species., Published as part of Theischinger, Günther, Mitchell, Andrew, Richards, Stephen J. & Polhemus, Dan A., 2023, Systematics of the Nososticta salomonis complex (Odonata: Zygoptera: Platycnemididae), pp. 101-146 in Zootaxa 5296 (2) on pages 120-122, DOI: 10.11646/zootaxa.5296.2.1, http://zenodo.org/record/7978135

Nososticta stueberi sp. nov. ( Figs 42–49, 61, 62, Map 4, Pl. 5) Holotype. ♁, SAMA 07-001596, Papua New Guinea, East Sepik Province, Iniok (4°17.296’S, 142°01.236’E; 45 m a.s.l.), 15-vi-2010, S. Richards; deposited in the South Australian Museum, Adelaide, Australia. Paratypes. Papua New Guinea: 1 ♁, 2 ♀♀, SAMA 07-001594, 07-001595, 07-001689, same details as holotype, 14–17-vi-2010, S. Richards; deposited in the South Australian Museum, Adelaide, Australia. 3 ♁♁, 1 ♀, BPBMENT 0000081215–81218, Milne Bay Prov., Bwaona River, E of Mila, upper Sagarai River basin, 90 m a.s.l., 10°30’14”S, 150°18’50”E, water temp. 28° C, 07-iv-2002, CL, 7165, D.A. & J.T. Polhemus; 3♁♁, BPBMENT 0000081223–81225, Milne Bay Prov., Sideia Island, Kwabunamoa River headwaters in south-central interior, 35– 40 m a.s.l., 10°35’59”S, 150°50’54”E, water temp. 26.5° C, 15-i-2004, CL 7288, D.A. & J.T. Polhemus; 2♀♀, BPBMENT 0000081221–81222, Milne Bay Prov., D’Entrecasteaux Is., Fergusson Is., Mebulibuli Creek trib., S. of Basima, 15–75 m a.s.l., 9°30’42”S, 150°52’06”E, water temp. 24° C, 24-viii-2002, CL 7181, D.A. Polhemus; 1 ♁, 1 ♀. BPBMENT 0000081219–81220, Milne Bay Prov., Muguwe Creek, nr Naura, 26 km W of Alotau, 70 m a.s.l., 10°17’20”S, 150°12’10”E, water temp. 25.5° C, 09-iv-2002, CL 7169, D.A. & J.T. Polhemus; all deposited in the Bishop Museum, Honolulu, Hawaii. 2 ♁♁, 3 ♀♀, RMNH. INS. 1153705–1153709, NE New Guinea, Morobe Dist., 6 mi. W of Lae, 17–26-x-1972 ex coll. Donnelly; 1 m, RMNH. INS. 1153639, NE New Guinea, Morobe Dist., creek nr. Gabensis, Wau Rd, 17–26-x-1972 ex coll. Donnelly; all deposited in Naturalis Biodiversity Center, Leiden, Netherlands. 1 ♀, RMNH. INS. 1153640, Camp 1 [Menapi, Cape Vogel Peninsula, 6.5 km. E. of Baniara, 0-20 m, 9°45’37”S, 149°56’00”E, Fourth Archbold Expedition; see Brass (1956)], 24.III.1953; 2 ♀♀, RMNH. INS. 1153641, 1153642, Camp 1 [Menapi, Cape Vogel Peninsula, 6.5 E. of Baniara, 0-20 m, 9°45’37”S, 149°56’00”E, Fourth Archbold Expedition; see Brass (1956)], 2.iv.1953; 2 ♁♁, 3 ♀♀, RMNH. INS. 1153716, 1153643–1153646, Camp 1, 10.iv.1953; 1 ♁, RMNH. INS.1153647, Camp 2, 10.v.1953; 1 ♁, RMNH. INS.1153648, Camp 7 [Peria Creek, off Kwagira River, 50 m., nr. Awani village, vic. 9°39’06”S, 149°23’47”E, Fourth Archbold Expedition; see Brass (1956)], 23.VIII.1953; all deposited in Naturalis Biodiversity Center, Leiden, Netherlands. Indonesia: 1 ♁, 1♀, RMNH. INS.1153649, 1153650, N New Guinea, iii-1931, W. Stüber; 3 ♁♁, RMNH. INS. 1153651–1153653, N. N. Guinea, 11–xii-1930, W. Stüber; 24 ♁♁, RMNH. INS. RMNH. INS. 1153717, 1153718, 1153654–1153675, N. New Guinea, 27.viii–4.ix.1930, W. Stüber; 1 ♁, 1 ♀, RMNH. INS. 1153676, 1153677, cop., N. N. Guinea, 27.viii– 4.ix.1930, W. Stüber; 9 ♀♀, RMNH. INS. 1153678–1153686, N. New Guinea, Humboldt Baai, iii-1931, W. Stüber; 1 ♁, RMNH. INS. 1153687, N. N. Guinea, Humboldt Baai, 19, W. Stüber; or ii-1953, Tamivlabta; 8 ♀♀, RMNH. INS. 1153688–1153695, N. New Guinea, Humboldt Baai, i-1932, W. Stüber; or Kressi, 300 m, 120 km; 1 ♁, 1 ♀, RMNH. INS. 1153696, 1153697cop., N. New Guinea, Humboldt Baai, xii-1932, W. Stüber; or Agaffo Sago; 3 ♁♁, 4 ♀♀, RMNH. INS. 1153698–1153704, N. N. Guinea, Humboldt Baai, 9.iii–11.iv.1936, W. Stüber; or Berap Lakes; 1 ♁, RMNH. INS.1153710, Hollandia, 23-vi-1938, Toxopeus; 5 ♁♁, RMNH. INS. 1153711–1153715, Hollandia, vii-1938, L.J. Toxopeus; all deposited in Naturalis Biodiversity Center, Leiden, Netherlands. Etymology. The species is dedicated to its probable discoverer, Wilhelm Stüber (1877–1942), collector extraordinaire of New Guinean dragonflies, including numerous individuals of this species. The name is a noun in the genitive case. Description of holotype Head. Largely black; only base of labium pale grey and transverse frontal bar from eye to eye blue; a faint ill-defined pale brown mark only barely detectable between antennal base and lateral ocellus. Thorax. Pronotum black with tiny inner spot and small blue mark distally on anterior lobe, very close to large blue lateral patch on median lobe, pleura largely bright blue, black along sutures and at rear of epimeron. Synthorax with pleura largely black except following areas of blue: ante-humeral patch, markedly shorter than 1/2 length of mesanepisternum, basally not more than 1 and 1/2 times as wide as at truncate dorsal end; patch covering at least dorsal 3/4 the length and approximately posterior 1/2/ to 3/4 width of mesepimeron, partly connected with stripe across whole length and anterior 3/4 of metepisternum; patch across posterior half of metepimeron, dorsal half of its anterior margin convex; posterolateral tip of meso- and metakatepisternum. Postcoxae largely blue and black; poststernum pale blue with small and narrow patches of blackish brown to black. Legs: Coxae and trochanters blue and black, legs otherwise black, with only base of pro- and mesofemur and basal inner 1/3 of metafemur partially pale blue. Wings: membrane with very faint greenish brown tinge, venation black; pterostigma black, about twice as long as wide; postnodals 14/12–13. Abdomen. S1 dorsally black, laterally largely pale blue; S2 largely black, pale blue adjacent to genitalia but dorsal blue mark absent; S3–7 dorsally black, ventrally margined bluish white, S3–6 with blue mid-dorsal mark very close to base; S8 largely blue, narrowly lined with black basally, more widely and irregularly lined with black apically, sometimes divided by very fine black mid-line; S9 and S10 black. Superior anal appendages largely blue, inferiors brownish black. Dimensions. Hind wing 19.0 mm; abdomen (including anal appendages) 31.5 mm. Description of female Head. Much as in male, but transverse frontal bar from eye to eye whitish yellow. Thorax. Pronotum much as in male, but anterior lobe with larger outer mark only, bluish white and narrowly separated from large patch of same colour on median lobe, posterior lobe with vertical dorsal and horizontal ventral lobe; propleura much as in male but pale areas largely bluish white instead of blue. Synthorax with pleura largely black. Ante-humeral patch pale yellow to brownish, less than 40% length of mesanepisternum, about 1/4 to 1/3 as wide as long, narrowly oval to almost parallel sided. Bluish white areas include: a remnant mesepimeral patch, variable in length and somewhat in width, completely fused with metepisternal stripe; metepimeral stripe; more extensive posterolateral tip of meso- and metakatepisternum. Postcoxae bluish white and black; poststernum bluish white with largely narrow elongate patches of brown to black. Legs: Coxae largely pale to bluish white, meso-and metacoxa black anterobasally, trochanters black and pale yellow; legs otherwise dark, with base of femora pale to dull yellow. Wings: membrane almost clear, venation black; pterostigma much as in male, greyish brown; postnodals 14–17/12–14. Abdomen. S1 and S2 dorsally largely black, laterally largely bluish white; S3–7 dorsally black, ventrally narrowly margined whitish blue to pale yellow, S3–6, with very short bluish white dorsal mark very close to base, often divided by dark midline; S8 black with ventral edge pale yellow; S9 and S10 black. Anal appendages pale yellow, inferior anal lamina brown and yellow; ovipositor brownish to dull yellow, valves and supra-anal plate black. Dimensions. Hind wing 18.3–20.0 mm; abdomen 28.0– 31.3 mm. Variation in the male paratypes. Postnodals 13–15/12–13; pterostigma ranging in colour from yellowish grey and yellowish brown (due to immaturity and method of preservation) to black and in proportion from slightly less to slightly more than twice as long as wide; hind wing 16.6–19.1 mm; abdomen (including anal appendages) 27.6–32.0 mm. Affinities. Based on the present study (Fig. 6), N. stueberi sp. nov. is closest to N. tagula sp. nov. and N. boonei sp. nov. Differential diagnosis. Nososticta stueberi sp. nov. is the only species of the N. salomonis complex with the blue/pale ante-humeral patch in both sexes markedly shorter than 1/2 the length of the mesanepisternum and close to parallel sided in the female. There appears to be limited morphological variation across this species’ extensive known distribution. Distribution and habitats. Nososticta stueberi has a broad distribution across northern and eastern New Guinea extending from at least the Jayapura region in Papua Province, Indonesia in the west to Alotau in Milne Bay Province, Papua New Guinea, in the east (Map 4, Pl. 5). It occurs in both primary and degraded forest habitats where adults perch on twigs or low foliage along small, shallow streams. It also occurs in forest clearings some distance from water. One of us (DP) has collected this species at multiple sites across the New Guinea region, all of which were creeks or small rivers with cobble and gravel substrates (for examples, see Plate 5, Bwaona River and Muguwe Creek). Similarly, the Peria Creek habitat at Camp 7 of the 4 th Archbold Expedition where this species was taken is described by Brass (1956) as having “...a gravelly bed about 50 meters wide. In May it flowed about knee deep, but in August and early September there remained only deep pools of clear water.” It therefore seems that this species may tend to favour such habitats., Published as part of Theischinger, Günther, Mitchell, Andrew, Richards, Stephen J. & Polhemus, Dan A., 2023, Systematics of the Nososticta salomonis complex (Odonata: Zygoptera: Platycnemididae), pp. 101-146 in Zootaxa 5296 (2) on pages 125-127, DOI: 10.11646/zootaxa.5296.2.1, http://zenodo.org/record/7978135, {"references":["Brass, H. G. (1956) Results of the Archbold Expeditions No. 75. Summary of the Fourth Archbold Expedition to New Guinea (1953). Bulletin of the American Museum of Natural History, 111 (2), 81 - 151."]}

Nososticta salomonis (Selys, 1886) (Figs 7–16, Map 1, Pls 1, 7, 8) Alloneura salomonis Selys, 1886: 188. Material examined. Solomon Islands: Holotype ♁ (photographs). Solomon Is, Mathew (BMNH); the specimen probably comes from Ugi Is near San Cristobal. Other material. As listed below and described in Table 2 from the areas given under Known Distribution. Choiseul Is.: 13 ♁♁, 2 ♀♀, Choiseul Prov., Vangamole River, 3 km N of Mole, 25 m a.s.l., 6°48’40”S, 156°32’42”E, water temp. 28.5° C, 17-iii-2005, CL 7389, D.A. Polhemus; 9 ♁♁, same locality and data as preceding, D.A. Polhemus and R.A. Englund; all in BPBM. 1 ♀, Kitipi River, 80 m a.s.l., 12-iii-1964, P. Shanahan; in RMNH. Guadalcanal: 7 ♁♁, 1 ♀, Guadalcanal Prov., Tenaru River at Tenaru Falls, SE of Honiara, 40 m a.s.l., 9°31’01”S, 160°00’59”E, water temp. 25° C, 21-xi-2004, flying along stream margins, CL 7337, D.A. Polhemus; 8 ♁♁, Guadalcanal Prov., Lungga River gorge, SW of Mt Austen at proposed dam site, 25 m a.s.l., 9°23’44”S, 159°50’48”E, water temp. 25.5° C, 28-vii-2005, CL 7407, D.A. Polhemus; 7 ♁♁, 1 ♀, Guadalcanal Prov., Charebuma River, above Gold Ridge mine, 390–460 m a.s.l., 9°36’00”S, 160°07’23”E, water temp. 23.5° C, 03-viii-2005, CL 7421, D.A. Polhemus; all in BPBM. 1 ♁, Ilu farm, Rystveld, 30-x-1964, R. & G.S. Slooff; 1 f, Komugelea, 1200 ft, Ananastuintje, 23-ix-1965, R. Slooff; 2 m, rivier bij Tenaligi, 25-ix-1965, R. Slooff; 1 ♁, 3 ♀♀, Tenaligi-river, dicht bos, 25-ix-1965, R. Slooff; all in RNMH. Kolombangara Is.: 1 ♁, Western Prov., rocky forest stream 5.5 km N of Ringgi harbour, 120 m a.s.l., 7°12’03”S, 156°54’40”E, water temp. 26° C, 19-iii-2005, CL 7394, D.A. Polhemus; in BPBM. Makira Is. (= San Cristobal): 1 ♁, Makira and Ulawa Prov., Tagataga Creek, 4 km S of Kirakira, 160 m a.s.l., 10°29’34”S, 161°55’58”E, water temp. 26° C, 11-iii-2005, CL 7375, D.A. Polhemus; 1 ♁, Makira and Ulawa Prov., Tawaitara Creek, SE of Tawaitara, 30 m a.s.l., 10°27’35”S, 161°56’37”E; water temp. 26° C, 10-iii-2005, CL 7374, J.T. Polhemus; 3 ♁♁, Makira and Ulawa Prov., Puepue River and trib., 3 km S of Kirakira, 15–50 m, 10°28’40”S, 161°55’33”E, water temp. 26° C, flying along main river channel, 10-iii-2005, CL 7373, D.A. Polhemus; all in BPBM. Malaita Is.: 4 ♁♁, Malaita Prov., Aluta River at bridge on Auki-Atori road, 75 m a.s.l., 8°41’46”S, 160°50’03”E, water temp. 25.5° C, 30-vii-2005, CL 7411, D.A. Polhemus; 4 ♁♁, 4 ♀♀, Malaita Prov., Kwarea River at high bridge on road to Atori, 85 m a.s.l., 8°39’10”S, 160°45’33”E, water temp. 25° C, 29-vii-2005, CL 7408, D.A. Polhemus; 4 ♁♁, 2 ♀♀; Malaita Prov., Kwaisele River at Kwaisele Falls, nr Anomasu, 180 m a.s.l., 8°41’53”S, 160°49’40”E, water temp. 25.5° C, 30-vii-2005, CL 7410, D.A. Polhemus; 2 ♁♁, Malaita Prov., Malaita, Kapolo Creek, 7.5 km N of Auki, 10 m a.s.l., 8°42’09”S, 160°41’59”E, water temp. 26° C, 31-vii-2005, CL 7412, D.A. Polhemus; all in BPBM. 1 ♁ Hojollo, schaduwryke bosbeek, 300 ft a.s.l., 13-x-1967, R. Sloof; 1 ♀, Anoasa Hill, 20 km S. Auki, 400 m a.s.l., 13-ii-1982, G. & T. Racheli; 1 ♁, Tanytalau, 26-ix-1957, J.L. Gressitt; all in RMNH. New Georgia: 2 ♁♁, Western Prov., Dadaloana River, NE of Munda, 20 m a.s.l., 8°08’58”S, 157°21’22”E, water temp. 26° C, 18-xi-2004, CL 7334, D.A. Polhemus; 2 ♁♁, Western Prov., Sakumbare River, near former Sakumbare logging camp, 15 km E of Putagita, 190 m a.s.l., 8°24’30”S, 157°41’34”E, water temp. 26° C, 22-iii-2005, CL 7401, D.A. Polhemus; all in BPBM. Ranongga Is.: 3 ♁♁, 3 ♀♀, Western Prov., Paoroe Creek, above Boroi village, 50–70 m a.s.l., 8°05’00”S, 156°35’51”E, water temp. 26° C, 14-iii-2005, flying along stream margins, CL 7381, D.A. Polhemus; in BPBM. Rendova: 1 ♁, Western Prov., Toropi River headwater trib., 5.5 km S of Kenole, 230–250 m a.s.l., 8°30’30”S, 157°18’45”E, water temp. 26° C, 21-iii-2005, CL 7399, D.A. Polhemus; in BPBM. Santa Isabel: 5 ♁♁, Santa Isabel Prov., Laticie River, above Buala, 2–118 m a.s.l., 8.14212°S, 159.58585°E, 23-vii-2005, R. Englund & D. Boseto; 2 ♁♁, Santa Isabel Prov., Blahitada Stream and large waterfalls above hydropower plant, upstream of Buala, 8.15681°S, 159.59264°E, 280 m a.s.l., 22-vii-2005, R. Englund & D. Boseto; all in BPBM. Shortland Is: 4 ♁♁, in very bad condition, Lofung, 18-x-1936, R.A. Lever; in RMNH. Vella Lavella Is.: 7 ♁♁, 1 ♀, Western Prov., Bainapo Kasi Creek, trib. to Sireri River, 5–15 m a.s.l., 7°45’54”S, 156°36’17”E, water temp. 26° C, 15-iii-2005, hovering along stream margins, CL 7385, D.A. Polhemus; 1 ♀, same locality and data as preceding, R.A. Englund; all in BPBM. Affinities. Although the COI distances of material from San Cristobal, from Guadalcanal and Malaita and from Santa Isabel, Choiseul, Vella Lavella, New Georgia and Rendova are sufficient to consider these three groups as different species, there are large gaps in sampling and the poor quality of some material has prevented adequate morphological comparisons. We have therefore taken a conservative approach and regard all Nososticta from the Solomons archipelago, except the population from Bougainville which is both morphologically and genetically very divergent, as a single geographically variable species, Nososticta salomonis, which is the sister taxon of N. hedigeri sp. nov. from Bougainville. Differential diagnosis. The lack of a pale dorsal marking on segment 2 distinguishes the male of N. salomonis from N. africana and N. tagula sp. nov. and the lack of a narrow black basal stalk between the ante-humeral patches distinguishes it from N. hedigeri sp. nov. A straight anterior margin of the mostly not shortened metepimeral patch separates the male of N. salomonis from N. chrismulleri (anterior margin of the shortened metepimeral patch dorsally convex). The male of N. salomonis can be separated from N. boonei sp. nov. by the lack (vs. presence) of blue pattern elements on S7 and from N. stueberi sp. nov. by the markedly longer antehumeral patch. However, as the ante-humeral patch of N. salomonis ranges from as long as, to markedly longer than, half the length of the mesanepisternum, and can even be absent in the female, and as the poststernum of N. salomonis ranges from yellow over bluish, variously patterned with brown to black patches to completely black in both sexes, it appears helpful to additionally define N. salomonis (in the present concept) also by distribution. Distribution and habitats. Nososticta salomonis as defined in this work is restricted to the Solomon Islands archipelago between at least Choiseul island in the northwest and San Cristobal island in the southeast (Map 1, Pl. 1)., Published as part of Theischinger, Günther, Mitchell, Andrew, Richards, Stephen J. & Polhemus, Dan A., 2023, Systematics of the Nososticta salomonis complex (Odonata: Zygoptera: Platycnemididae), pp. 101-146 in Zootaxa 5296 (2) on pages 110-114, DOI: 10.11646/zootaxa.5296.2.1, http://zenodo.org/record/7978135

Nososticta africana (Schmidt, 1944) (Figs 17–23, 55, 57, Map 2, Pl. 2) Notoneura africana Schmidt, 1944: 43. Material examined. Holotype ♁ (photographs). The labels state: Massawa, Eritrea, 29-xii-1907 (MfN Berlin). The specimen probably comes from Massua on New Britain Island, Papua New Guinea (Gassmann 2015). Papua New Guinea, West New Britain Province: 1 ♁, West New Britain Prov., spring and creek on E. slope of Mt Garbuna, 35 m a.s.l., 5°26’41”S, 150°05’10”E, water temp. 27° C, 15-i-2005, CL 7351, D.A. Polhemus; 2 ♁♁, 1 ♀, West New Britain Prov., trib. to Kopiura River, SW of Ubai, 310 m a.s.l., 5°45’26”S, 150°34’50”E, water temp. 27° C, 12-i-2005, CL 7343, D.A. Polhemus; 1 ♁, West New Britain Prov., freshwater swamp behind beach nr Walindi Plantation, sea level, 5°26’25”S, 150°05’19”E, water temp. 26° C, 11-i-2005, CL 7339, D.A. Polhemus; all in BPBM; following all from New Britain Island: 1 ♀, Lindenhafen, Cocos Grove, 23-iv-1956, J.L. Gressitt (RMNH); 1 ♁, 1 ♀, Metelen R. [Metelen River, trib. to upper Kopiura River], 20 mi SE of Ruango, vii-1970, Jindrich leg. Ex coll. Donnelly (RMNH). East New Britain Province: 1 ♁, 1 ♀, largely pulverised; 3 ♁♁, 6 ♀♀, Gazelle Pen., Kerawat, 60 m a.s.l., Primary bush, ix-1-55, JLG. (RMNH); 1 ♁, 1♀, Gazelle Pen., Warongoi Valley, 24-v-1956; Bishop (RMNH); 2 ♁♁, Vunabakan [Vunavakan Plantation], 16-20-xi-1959, J.L. Gressitt (RMNH). Papua New Guinea, New Ireland Province: 2 ♁♁, 1 ♀, Lihir Island, town area, 3°03’S, 152°36’E, 20–26-x-2001, M. Humphrey & M. Moulds (AM); 3 ♁♁, 1 ♀, Lihir Island, town area, 3°03’S, 152°36’E, 20-26-x-2001, M. Moulds & M. Humphrey (AM); Morobe Province: 1 ♁, NE New Guinea, Morobe Dist. Bulolo, vi-1970, Jindrich; ex coll. Donnelly (RMNH). Affinities. Based on the present study (Fig. 6), N. africana is sister to the remaining taxa of the N. salomonis complex which can be subdivided into a northern and a southern group (Fig. 6). Differential diagnosis. Nososticta africana can be distinguished from all other members of the N. salomonis complex except N. tagula sp. nov. by having abdominal segment 2 marked with blue dorsally. It can be distinguished from N. tagula sp. nov. by having one or two blue antero-dorsal marks and one postero-dorsal mark on segment 2 (vs. only a blue postero-dorsal mark). Nososticta africana is known only from the Bismarck Archipelago, while N. tagula sp. nov. is known only from the Louisiade Archipelago. Distribution. Most records of N. africana are from the islands of New Britain, New Ireland and Lihir in the Bismarck Archipelago, but there is a single record of this species from Bulolo in Morobe Province on mainland New Guinea (Maps 2, 4, Pl. 2). It is an ecologically adaptable species that occurs along small, clear streams in primary forest, degraded forest, and heavily modified landscapes such as gardens and plantations. Gassmann (2015) provides the first record ([his Site 37]) from New Ireland (a male specimen reported from “New Ireland, between Lagagon and Fissoa, 93 km south to Kavieng, at East Coast Highway, 23.vi.1997, stagnant creek in forest (not very clean).”, Published as part of Theischinger, Günther, Mitchell, Andrew, Richards, Stephen J. & Polhemus, Dan A., 2023, Systematics of the Nososticta salomonis complex (Odonata: Zygoptera: Platycnemididae), pp. 101-146 in Zootaxa 5296 (2) on page 115, DOI: 10.11646/zootaxa.5296.2.1, http://zenodo.org/record/7978135, {"references":["Schmidt, E. (1944) Die erste afrikanische Notoneura - Art (Ordn. Odonata). Mitteilungen der Deutschen Entomologischen Gesellschaft, E. V., 12, 43 - 45.","Gassmann, D. (2015) Odonata recorded from northeastern Papua New Guinea including the Bismarck Archipelago in May to July 1997. Faunistic Studies in South-East Asian and Pacific Island Odonata, 10, 1 - 46."]}

Rigoutorum bacchusi (Rigout & Allard, 1997) new combination (Figs 4–6, 10A) Diaphonia bacchusi Rigout & Allard, 1997: 26–27, dorsal image, aedeagus. Holotype male NHML. Type locality: Southern Cross, Western Australia. Diaphonia bacchusi: Calder 2002; Golding 2009: 20 (centre image); Hutchinson & Moeseneder 2013: 2; Hutchinson & Moeseneder 2019: 279; Moeseneder et al. 2019: 545. Specimens examined (57 males, 2 females). Holotype. Male: Southern Cross, W.A., H.W. Brown, Diaphonia bacchusi n. sp. J. Rigout det. 1996 TYPE, Dysdiatheta minima n. sp M.E. Bacchus det, National Museum of Victoria Melbourne, [genitalia on card]; [NHML]. Other specimens examined. 1 male: Yellowdine, WA, 2. Feb. 1970, Ex Carnaby Coll., Diaphonia bacchusi male Rigout & Allard det. T.A. Weir 2016; [WAM]. 1 male: E. Hyden, WA., 26-1-75, Pseudoclithria ruficornis, Bequeathed, H. Demarz Coll.; AFBRC8455-01 [AFBRC]. 9 males: 50 km S. Coolgardie W. Australia, 2. i. 1977, A.M & M.J. Douglas leg. ABFF, Pseudoclithria ruficornis (Westwd.) det. R.M. Young 1989; 88/3007, 88/3008, 88/3009, 88/3010, 88/3011, 88/3012, 88/3013, 88/3014, 88/3015 [WAM]. 4 males: 57 km. E. Hyden W.A. Cruising, 10-2-[19]91, G.P, Golding/Powell Collection donated 12. Feb. 2002; [WAM], AFRBC8459-01, 8459-02 [AFBRC]. 4 males: Yellowdine, W.A., 16-1-1992, male, Donated 2015, Jack Hasenpusch, Aust. Insect Farm; AFBRC8456-01, 8456-02, 8456-03, 8456-04 [AFBRC]. 2 males: E. Sth. Cross, W.A., 22-1-1993, H. Demarz, Pseudoclithria ruficornis male, Donated 2015, Jack Hasenpusch, Aust. Insect Farm; AF- BRC8457-01, 8457-02 [AFBRC]. 1 male: 21 km S. of Yellowdine West. Aust., 28. Jan. 1997, P.H. Pseudoclithria; CET 2164 [PMH]. 1 male: 29 km S. of Lake King W. Aust., 1. Feb. 1999, dead, salt lake, Golding/Powell Collection donated 12. Feb. 2002; [WAM]. 6 males: 104 km E. of Lake King W.A., 3. Feb. 1999, P. Hutchinson, Flying over Euc.[alyptus]; CET 0393 [genitalia on card], 0395, 0397, 0398, 0399 [PMH], AFBRC8460-01 [AFBRC]. 1 male: 9 km SSE Peak Eleanora Upper Lort River W. Aust., 5. Feb. 2000, dead, salt river edge, Golding/Knowles, Golding/ Powell Collection donated 12. feb. 2002; [WAM]. 2 males: L. Hurlstone, W.A., 6.3.2000, dead; AFBRC8458-01, 8458-02 [AFBRC]. 1 male: 53.4 km s. of Southern Cross West. Aust., 30. Jan. 2004, P. Hutchinson, Diaphonia flying over heath; CET 0394 [genitalia on card] [PMH]. 1 male: Grt. E. Hwy. 33 km E. of Yellowdine W. Aust., 5. Feb. 2006, P. Hutchinson; CET 2638 [PMH]. 1 male: Lake King (strandline) West. Aust., 21. Feb. 2006, P. Hutchinson, Diaphonia bacchusi dead lake edge; CET 0391 [genitalia on card] [PMH]. 2 males: 6 km N of Yellowdine, WA, 23 Jan. 2009, TM & LM Hanlon, On Acacia sp leaves; AFBRC8313-01, 8381-01 [AFBRC]. 1 male: Grt. E. Hwy. 35 km E. of Yellowdine W. Aust., 12. Feb. 2009, P. Hutchinson; CET 2639 [PMH]. 2 males: Lake King, W.A., 14 Feb 2011, P. Hutchinson & C. Moeseneder, salt lake, dead on strandline; MIC61002 -005, 61002-006 [AFBRC]. 1 male: 26 km S. of Menzies W.A., 26. Jan. 2012, M. Hanlon & M. Powell on mallee flowers; CET 2164 [PMH]. 2 males: Ghooli, E. of Southern Cross W. Aust., 15. Jan. 2013, P. Hutchinson, Diaphonia bacchusi 0930-1045 hrs flying over plants; CET 0409, 1318 [PMH]. 7 males: Ghooli, E. of Southern Cross W. Aust., 15. Jan. 2013, P. Hutchinson, Diaphonia bacchusi 1045-1110 hrs flying over plants; CET 1199, 1380, 2635, 2636, 2637 [PMH], MIC61102 -001, 61102-002 [AFBRC]. 1 male: 10 km W. Yellowdine Rd. Hs. West. Aust., 26. Jan. 2016, P. Hutchinson, Diaphonia bacchusi dead Male; SCAR135-D4, CET 2203 [PMH], AFBRC8380-01 [AFBRC]. 2 males: Karalee Pumping Station West. Aust., 27. Jan. 2016, P. Hutchinson, Diaphonia bacchusi 0915-0945 [hrs] flying over Acacia; CET 2201, 2202 [PMH]. 1 female: Karalee Pumping Station West. Aust., 27. Jan. 2016, P. Hutchinson, Diaphonia bacchusi 8 ft. high on Acacia 0945 [hrs]; CET 2302 [PMH]. 1 male: Karalee Pumping Station West. Aust., 27. Jan. 2016, P. Hutchinson, Diaphonia bacchusi 1000-1030 [hrs] on Acacia; CET 2200 [genitalia on card] [PMH]. 1 male: Karalee Pumping Station West. Aust., 27. Jan. 2016, P. Hutchinson, Diaphonia bacchusi 1000-1030 on Allocasuarina; CET 2213 [genitalia on card] [PMH]. 1 male: Southern Cross W.A., H.W. Brown, E.F. DuBoulay Collection, Pseudoclithria ruficornis (Westwd.) det. R.M. Young 1989; 88/3016 [WAM]. 1 female: Tammin, W.A., H.W. Brown; AF- BRC8461-01 [AFBRC]. Redescription. Holotype Male (Figs 4, 10A). Condition: right antennal club and one protarsomere 5 missing, one protarsomere 5 attached to metasternum. Length 16.9 mm, width 8.5 mm, widest at humeral umbone. Ovate, depressed. Head. Black. Clypeus quadrate, length:width ratio 1:1, reflexed at apex, clypeolateral ridge present from antennal insertions, high, linear, parallel-sided, angulate to anterolateral angles and continuous across slightly sinuate anterior margin. Lateral declivity deep in apical half, striolate; apical declivity deep and micropunctate, both angulate and visible in dorsal view. Disc flat, abruptly inclined to highly elevated clypeolateral and apical ridge. Surface coarsely punctate bearing dense, very long, yellow pilosity. Clypeofrontal suture absent. Frons flat with coarse punctures bearing dense, very long, yellow pilosity. Ocular canthus narrow, bearing short yellow setae. Antennae with scape black, elongate, along dorsum of length bearing few, short, pale setae and dorsum of apex bearing long, black setae, posterior margin bearing dense setal brush of long, yellow pilosity. Pedicel black bearing sparse, pale microsetae. Antennomere 3 elongate, same length as antennomeres 4 and 5 combined. Antennomeres 3–7 black with apices dark brown and bearing sparse, long, pale setae. Club dark brown, length of antennomeres 3–7, 0.3 x length of head. Antennomere 8 on internal surface bearing setae medially. Antennomere 10 external surface dark brown-black. Thorax. Pronotum black with narrow brown maculae laterally along posterolateral margin. Transverse, with basomedian margin arcuate, basolateral margin linear and angled anteriorly. Basolateral angle narrowly arcuate through 90°. Posterolateral margin linear, slightly attenuate to broadly arcuate midlength then linear attenuate to anterolateral angles.Anterior margin linear. Lateral margins ridged. Disc with evenly dense, moderately coarse punctures becoming rugulose over majority of surface, bearing dense, long, yellow pilosity. Scutellum black, equilateral triangle with coarse punctures except along midline, bearing long, pale pilosity. Elytra yellow-brown, sutural costa black, lateral margins narrowly dark brown. Mesepimeron narrowly exposed, with base of elytra laterally pronounced. Posthumeral arch linear in dorsal view, arcuate in lateral view, weakly exposing metacoxa, obscured by pilosity, then slightly attenuate to broadly arcuate apex. Disc (interstices 1 and 2) coarsely punctate in weakly defined rows becoming coalesced, then rugulose at apical declivity, laterad to costa 2 rugulose; bearing long, pale setae. Humeral and apical umbones raised. Costa 1 broadest, costa 2 narrower, costa 3 barely visible. Sutural costa parallel basally, raised and attenuate in apical half with apices bearing a minute spine. Mesometasternal process black, broad, flat with broadly arcuate apex, declivous at anterior margin of mesocoxa; surface impunctate, glabrous with apical declivity bearing dense, very long, yellow pilosity. Metasternum black, medially with moderately dense punctation, laterally rugulose bearing long, yellow pilosity. Mesepimeron, metepisternum and metepimeron black, rugulose, bearing long, yellow pilosity. Legs. Black. Profemora dorsoventrally flattened, slightly attenuate to apex with ventral surface rugulose bearing yellow pilosity. Protibiae elongate parallel, unidentate with apical denticle elongate acute; dorsal surface with medial and internal row of coarse punctures bearing sparse, long, pale setae and setal brush composed of moderately dense, short, ginger setae; spur stout, reaching apex of tarsomere 1. Protarsomeres 1–5 equal to tibial length; tarsomere 5 long and 2 x length of tarsomere 4. Protarsomeres 1–5 bearing setae less than half tarsomere length. Protarsal claws long, simple and symmetrical. Mesofemora dorsoventrally flattened, parallel with ventral surface rugulose bearing very long, yellow pilosity. Mesotibiae elongate, ventral surface medially with 2 rows of coarse punctures bearing short, yellow setae in basal half. Internal margin and basal half of external margin bearing sparse, short, pale setae. Dorsal surface bearing dense, very long, yellow pilosity. External margin evenly arcuate, non-denticulate, apex bispinose. Internal spine short, acute; external spine long, acute. Spurs long, narrow, attenuate with an acute apex to middle of mesotarsomere 2. Mesotarsomeres 1–5 longer than tibial length. Mesotarsomere 5 longest, 2 x length of tarsomere 4. Mesotarsomeres 1–4 bearing setae equal to tarsomere length. Mesotarsal claws long, simple and symmetrical. Metacoxae black, posterolateral angle 90°, surface rugulose bearing long, yellow pilosity. Metafemora dorsoventrally flattened with anterior margin weakly arcuate and posterior margin linear. Ventral surface medially sparsely punctate, rugulose along margins bearing very long, pale pilosity. Metatibiae elongate with ventral surface rugulose bearing long, yellow pilosity. Internal margin bearing sparse, short, yellow setae. External margin medially with low, very obtuse angle, non-denticulate, glabrous. Dorsal surface bearing dense, very long, yellow pilosity. Metatibial apex bispinose with internal spine short, acute and external spine broad, sinuate. Spurs parallel, apex acute, reaching apex of tarsomere 2. Metatarsomeres 1–5 equal to tibial length; tarsomere 5 longest, 2 x length of tarsomere 4. Metatarsomeres 1–4 bearing setae equal to tarsomere length, with tarsomeres 1–2 bearing setae 1.5–2 x tarsomere length. Claws long, simple and symmetrical. Abdomen. Sternites black. Sternites 2–6 medially with broad, shallow depression, concave in lateral view, with sparse row of punctures bearing short, pale setae and laterally with dense punctures bearing long, yellow pilosity. Sternite 7 medially impunctate and glabrous, laterally rugulose bearing long, yellow pilosity. Propygidium bearing long, yellow pilosity. Pygidium dark yellow with apex broadly black and base narrowly black extending medially to disc. Pygidium length:width ratio 1:1.75, angulate in lateral view, with a narrow, arcuate, slightly obtuse angle post midlength; surface concentrically rugulose with centre past midlength bearing sparse, short, pale setae at apex and basolateral angles. Genitalia (Fig. 10A). Phallobase widest at apex, attenuate to and narrowest at base near parameres, 2 x length of parameres, dorsal surface with a longitudinal, deep groove. Parameres narrow, external margin sinuate concave medially, arcuate to apex. Each paramere narrowing to midlength, divergent to pre-apex then attenuate to apex. Dorsal cleft 45° at base, then broad, ovate in basal two-thirds, apically attenuate to where parameres meet. Paramere in lateral view arcuate through 90° at basal third; surface sparsely punctate. Variation in males (Fig. 5). Length: 13.5–17.5 mm; width: 7.1–9.0 mm. Antennomere 10 external surface with ventral quarter brown. Pronotal basolateral angle rounded. Pronotal lateral maculation absent. Elytra with suture black, not extending to sutural costa. Apical spine absent. Mesotibial external margin angulate pre midlength; internal surface bearing long, yellow pilosity. Metatibial external margin evenly arcuate or serrate. Pygidium with black basal and apical maculation joining medially, enclosing 2 lateral, oblique, elliptical, yellow maculae. Female (Fig. 6). Description based on CET2302 [PMH] (Karalee, 45 km east of Southern Cross). Differs from male by the following characters. Length 16.3 mm, width 8.9 mm. Stouter, broader and deeper. Head. Antennal scape along dorsal length glabrous. Thorax. Pronotum without lateral yellow macula. Basolateral angle arcuate, slightly obtuse. Elytron. Black. Margins narrowly exposing sternites laterally. Legs. Femoral and tibial ventral surfaces bearing black pilosity. Protibiae stout, tridentate, each denticle long and acute with denticle 2 at midlength and closest to basal denticle; setal brush composed of long, black setae. Mesotibial external margin with acute denticle at midlength; apical spines broad, internal spine acute, external spine truncate; spurs reaching apex of mesotarsomere 2. Mesotarsomeres equal to length of tibia; tarsomeres 1–2 bearing setae 2 x length of tarsomere. Metatibial internal and external margins bearing sparse, long pilosity. External margin with short, truncate denticle at midlength. Apical spines broad, truncate. Spurs broad, parallel, internal spur broadest preapically with apex arcuate, reaching midlength of tarsomere 3. Abdomen. Sternites convex. Sternite 6 bearing long, black pilosity across posterior half. Pygidium black. Obtusely arcuate in lateral view. Variation in females. Length 17.2 mm, width 9.0 mm. Femora and tibial surfaces bearing yellow-brown pilosity. Protibial setal brush composed of long, yellow-brown setae. Sternites 5 and 6 bearing long, yellow-brown pilosity across posterior half. Pygidium dark purple-black. Diagnosis. Rigoutorum bacchusi can be recognised by its pronotal sculpture that consists of uniformly dense punctation that bears dense, long pilosity. The colour is dull black with or without a lateral, narrow, yellow maculation (less than one sixth width of pronotum) and restricted to basal half. The lateral maculation is absent in females. Differential diagnosis. Rigoutorum bacchusi is similar to R. jakli but differs in having a duller pronotum with uniformly medium-sized and adjacent punctures. Males are with or without narrow, yellow maculation along the basal half of the lateral margin. Natural history. Rigoutorum bacchusi occur in mallee heath on yellow sandplain with dominant large trees, specifically Eucalyptus (Myrtaceae), Allocasuarina (Casuarinaceae) and Acacia (Fabaceae). Only one specimen was seen on Eucalyptus flowers, even though plants were in bloom nearby, but most were seen flying at the top of taller plants in mallee 1.8–3.6 m high. The beetles circled the top of the tree or bush once or twice and then flew to the next plant, repeating the behaviour. We assume that males were searching for females, but males were never observed flying low to the ground or with a female. Two females of this species are known. One of these was seen at a height of 2.4 m on a 3 m tall, fine-leafed Acacia at 0945 at Karalee Pumping Station. From 0915–0945 approximately 10 males were seen flying over adjacent Eucalyptus, Allocasuarina and Acacia but they were not seen to approach the female. Between 1000–1030 conditions changed, with the weather turning cloudy, and the males stopped flying. Two males were later found on Acacia foliage and one male on Allocasuarina foliage. On 15 January 2013 at 1.5 km south of Ghooli (Fig. 7), many males were observed exhibiting the same behaviour as described above. The beetles were in flight, along a remnant roadside vegetation strip on yellow sand, bordering large areas of farmland. Several plants were examined around their base for signs of larvae, larval faecal droppings or adult remains. Leaf and bark litter beside a mallee (Eucalyptus) was removed, revealing cetoniine larval faecal droppings at no more than 30 cm from the tree trunk and beneath the mulch. One medium-sized and one large larva, assumed to be at L2 and L3 stages, were uncovered along with remains of an adult female beetle, which was retained. Larvae were also retained and transferred to rearing boxes, but these did not develop to adults. Adults were observed from 2 January to 12 February, based on specimen data and PMH trip diaries.

Genus Rigoutorum Hutchinson & Moeseneder, new genus (Figs 4–6, 8–10) http://zoobank.org/NomenclaturalActs/0b3827fe-945e-4a1d-aafb-38e61a5ff30e Type species: Diaphonia bacchusi Rigout & Allard, 1997: 26; here designated. Included species: Diaphonia bacchusi Rigout & Allard, 1997; Rigoutorum jakli new species Description. Length 12.5–18.9 mm. Form elongate ovate. Colour black (females) or brown with yellow elytra and maculation (males). Head. Clypeus quadrate, lateral margins divergent, parallel in apical half, widest apically; clypeolateral ridges present, parallel, anterior margin linear sinuate medially; margins abruptly inclined to high elevations; disc coarsely punctate. Frons coarsely punctate. Antennal club not enlarged in male, same length as antennomeres 2–7. Thorax. Pronotum transverse trapezoidal; basomedian margin weakly concave-linear; lateral margins with broad median angles; disc punctate. Scutellum base broad, elongate, coarsely punctate. Elytron with posthumeral arch linear (in dorsal view), slightly exposing metacoxa, then slightly convergent to broad arcuate apex; mesepimeron slightly exposed; humeral and apical umbones indistinctly raised; sutural costa distinct, each elytron with one complete, indistinct, low costa; disc punctate in irregular lines. Mesometasternal process undeveloped, present as low parallel process, declivous at anterior margin of mesocoxae. Legs. Protibiae unidentate, denticle elongate; protarsomeres elongate and simple. Mesotibiae broadest in basal half, not bidentate or weakly bidentate, concave in apical half. Mesotarsomeres elongate and simple. Metafemora elongate, anterior margin arcuate. Metatibiae elongate, narrow, parallel in apical half, angulate to weakly denticulate at mid-length; spurs elongate parallel, broader in females. Metatarsomeres elongate and simple. Abdomen. Sternites distinctly concave in males, convex in females. Metasternum sparsely punctate medially, rugulose laterally. Pygidium concentrically rugulose. Genitalia (Fig. 10). Phallobase to paramere length ratio 2:1, phallobase widest at apex with narrow, deep medial longitudinal groove on dorsal surface, parameres narrowing from base; sinuate concave widening near apex; dorsal cleft basally acute, narrowly ovate in basal half, then attenuate to apex. Differential diagnosis. The following combination of characters distinguishes Rigoutorum from other genera of Schizorhinini. Size 12.5–18.9 mm; clypeus quadrate with clypeolateral margins abruptly and highly elevated; anterior margin linear; antennal club not enlarged, same length as antennomeres 2–7; head and pronotum punctatesetose; male protibiae unidentate; mesometasternal process undeveloped. Table 2 provides distinguishing characters among Rigoutorum, Diaphonia and Aurum Hutchinson & Moeseneder, 2019. Specimens examined: Rigoutorum bacchusi; Rigoutorum jakli new species; Aurum leonorensis; Diaphonia dispar; Diaphonia xanthopyga; Diaphonia melanopyga Lea, 1914; Diaphonia luteola; Diaphonia vicina; Diaphonia notabilis White, 1846; Diaphonia antoinei Allard, 1995; Diaphonia lateralis Blackburn, 1894. Remarks. In Moeseneder et al. (2019), Rigoutorum keys out at couplet 42 “ Diaphonia bacchusi … Diaphonia Newman (part)”. In Krikken (2018), Rigoutorum keys out at couplet 21 “ Diaphonia and Chondropyga ”. Hutchinson & Moeseneder (2019) give differential characters of Aphanesthes, Aurum, Chondropyga Newman, 1840, Diaphonia, Diaphonia bacchusi and Diaphonia kerleyi Rigout & Allard, 1997. This study is our first step towards addressing the complex relationships among Diaphonia, Aphanesthes, Hemichnoodes, Pseudoclithria and Metallesthes. Mitchell et al. (2020) noted that “While the sampled species in this group are well defined, extensive further work is required to understand the relationships between these species and the status of the five current genera.” Natural history. Behaviourally, Rigoutorum belongs in the non-flower-visiting group of Australian Schizorhinina genera where males are usually seen in flight and females are sedentary and rarely observed. This is also the case in Aurum, Diaphonia, Pseudoclithria, Grandaustralis Hutchinson & Moeseneder, 2013, Navigator Moeseneder & Hutchinson, 2016, and Tapinoschema Thomson, 1880. The flower-visiting group of genera are mostly found congregating on flowers with males and females in similar numbers and both sexes having similar-sized antennae (e.g. Metallesthes, Hemichnoodes, Aphanesthes) (Hutchinson & Moeseneder 2013, 2019; Moeseneder et al. 2014, 2019; Moeseneder & Hutchinson 2016). Aurum and Rigoutorum new genus constitute a subset of the former group in which male antennal clubs are not enlarged. Etymology. The genus is named in honour of the Rigout family. Jacques Rigout contributed to the taxonomy of Batocerini (Coleoptera: Cerambycidae), Cetoniini and Schizorhinini (Cetoniinae), describing numerous new species. The Rigout family travelled to New Guinea researching Schizorhinini. Lydia Rigout operated Hillside Books., Published as part of Hutchinson, Paul M., Moeseneder, Christian H. & Mitchell, Andrew, 2022, Rigoutorum a new genus for Diaphonia bacchusi Rigout and Allard (Coleoptera: Scarabaeidae: Cetoniinae: Schizorhinini), with description of a second species from Western Australia and notes on habitats and behaviours, pp. 239-259 in Zootaxa 5150 (2) on pages 243-244, DOI: 10.11646/zootaxa.5150.2.4, http://zenodo.org/record/6621254, {"references":["Rigout, J. & Allard, V. (1997) Les Coleopteres du Monde. Vol. 25. Schizorhinini 3. Hillside Books, Canterbury, 128 pp.","Hutchinson, P. M. & Moeseneder, C. H. (2019) Aurum, a new monotypic genus of Cetoniinae (Coleoptera: Scarabaeidae) endemic to Western Australia, Australia. Zootaxa, 4683 (2), 277 - 285. https: // doi. org / 10.11646 / zootaxa. 4683.2.6","Lea, A. (1914) Notes on Australian Cetonides, with a list of species and descriptions of some new ones. Transactions of the Royal Society of South Australia, 38, 132 - 218.","Allard, V. (1995) Schizorhinini 1. The Beetles of the World. Volume 23. Sciences Nat, Venette, 152 pp.","Moeseneder, C. H., Weir, T. A., Lemann, C. & Hutchinson, P. M. (2019) Scarabaeidae: Cetoniinae Leach, 1815. In: Slipinski, A. & Lawrence, J. F. (Eds.), Australian Beetles. Vol. 2. Archostemata, Myxophaga, Adephaga, Polyphaga (part). CSIRO Publishing, Clayton South, pp. 531 - 553.","Krikken, J. (2018) Two new genera of Australian flower chafers, with an annotated overview of their relatives (Coleoptera: Scarabaeidae: Cetoniinae). Haroldius, 5, 1 - 34.","Mitchell, A., Moeseneder, C. H. & Hutchinson, P. M. (2020) Hiding in plain sight: DNA barcoding suggests cryptic species in all ' well-known' Australian flower beetles (Scarabaeidae: Cetoniinae). PeerJ, 8: e 9348, 1 - 25. https: // doi. org / 10.7717 / peerj. 9348","Hutchinson, P. M. & Moeseneder, C. H. (2013) Grandaustralis, a new genus and Grandaustralis boomerang, a new species of Cetoniinae (Coleoptera: Scarabaeidae) from Western Australia. Zootaxa, 3669 (1), 17 - 26. https: // doi. org / 10.11646 / zootaxa. 3669.1.2","Moeseneder, C. H., Hutchinson, P. M. & Lambkin, C. L. (2014) Revision of the genus Metallesthes Kraatz and description of Metallesthes anneliesae, a new species of Cetoniinae (Coleoptera: Scarabaeidae) from Queensland and New South Wales, Australia. Zootaxa, 3881 (4), 301 - 327. https: // doi. org / 10.11646 / zootaxa. 3881.4.1","Moeseneder, C. H. & Hutchinson, P. M. (2016) Navigator, a new endemic genus of Cetoniinae (Coleoptera: Scarabaeidae) from Australia with descriptions of two new species and behavioural studies. Zootaxa, 4173 (6), 530 - 556. https: // doi. org / 10.11646 / zootaxa. 4173.6.2"]}

Rigoutorum jakli Hutchinson & Moeseneder, new species (Figs 8–9, 10B) http://zoobank.org/NomenclaturalActs/ 698E17EC-7668-4476-81EA-AC7556D797E9 Diaphonia bacchusi: Golding 2009: 20 (left image—female, right image—male). Type series (61 males, 4 females): Holotype male. 66.6 km N. of Bullfinch W.A., 1. Feb. 2000, P. Hutchinson, Diaphonia bacchusi flying over Acacia, [genitalia on card]; CET 0380 [WAM]. Paratypes. 1 male: Marloo Stn Wurarga W.A., 1931–1941, A. Goerling, EB 32, GJB, Diaphonia bacchusi n. sp. J. Rigout det. 1996 PARATYPE, [genitalia on card] [paratype of Diaphonia bacchusi]; [NHML]. 1 female: Marloo Stn. Wurarga W.A., 1931–1941, A. Goerling, C.S.I.R.O; [NHML]. 7 males: 30 km W. Kirkalocka Stn. WA., 16:1:[19]89, M. Powell, Active over Acacia, Golding /Powell Collection donated 12. Feb. 2002; [WAM], AFBRC 8465-01, 8465-02 [AFBRC]. 6 males: 3 km W of [White] Wells Hstd, W.A., 24-1-98, Bequeathed H. Demarz Coll.; AFBRC 8463-01, 8463-02, 8463-03, 8463-04, 8463-05, 8463-06 [AFBRC]. 2 males: Paynes Find Rd W.A., 24-1-1998, H. Demarz; AFBRC 8462-01, 8462-02 [AFBRC]. 7 males: 24 k[m] E. of Pindar W.A., 26-1-[19]92, Golding/Powell Collection donated 12. Feb. 2002; [WAM], AFBRC 8464-01, 8464-02, 8464-03 [AFBRC]. 2 males: 53 km NW. Mt. Magnet W.A., 18:1:[19]93, Golding, Powell, active over sandplain, Golding/Powell Collection donated 12. Feb. 2002; [WAM]. 3 males: Paynes Find Rd N. of Wubin W.A., 24. Jan. 1998, H. DeMarz [sic], Pseudoclithria flying around Acacia CET 0387, 0389, 0392 [PMH]. 2 males: 13.0 km E. of L.[ake] Mongers (Perenjori-Wanarra Rd) W. Aust., 27. Jan. 2000, P. Hutchinson, Diaphonia bacchusi flying over Euc[alyptus]; CET 0384, 0386 [genitalia on card] [PMH]. 1 female: L[ake] Deborah N. Bullfinch West. Aust., 31. Jan. 2000, P. Hutchinson, found dead saltlake, CET 0381 [PMH]; [WAM]. 1 female: 44.8 km N. Bullfinch West. Aust., 31. Jan. 2000, M. Peterson, Diaphonia bacchusi dead edge saltlake; CET 0382 [PMH]. 1 male: 66.6 km N. of Bullfinch W.A., 1. Feb. 2000, P. Hutchinson, Diaphonia bacchusi flying over Acacia; CET 0383 [genitalia on card] [PMH]. 2 males: 86 km NNE of Wubin W.A., 27. Jan. 2004, P. Hutchinson, Diaphonia flying; CET 0388 [PMH], AFBRC 8383-01 [AFBRC]. 15 males: 61 km NNE of Wubin West. Aust., 29. Jan. 2005, P. Hutchinson, Diaphonia bacchusi flying over heath; CET 2621, 2622, 2627, 2628, 2629, 2630, 0400, 0401, 0403, 0404, 0405, 0406 [PMH], MIC61016 -001, 61016-002, AFBRC 8382-01 [AFBRC]. 2 males: Grt. N. Hwy. 61 km N. Wubin West. Aust., 30. Jan. 2005, P. Hutchinson; CET 2623, 2624 [PMH]. 1 female: 18 km E. of Grt. N. Hwy on Kalannie Rd. West. Aust., 3. Feb. 2006, P. Hutchinson, Diaphonia bacchusi on eucalyptus leaves; CET 0385 [PMH]. 8 males: Goodlands Rd., 25km SE Grt. N. Hwy (60 km N. Wubin) West. Aust., 11. Feb. 2009, P. Hutchinson; CET 2619, 2620, 2625, 2626, 2631, 2632, 2633, 2634 [PMH]. 2 males: Lake Baladjie N.R. WA., 20. Dec. 2012 - 19. Jan. 2013, T.M. Hanlon, Blue bucket trap; CET 2162, 2163 [PMH]. Description. Holotype Male (Figs. 8, 10B). Length 15.5 mm, width 8.16 mm, widest at humeral umbone. Ovate, depressed. Head. Black. Clypeus quadrate (length:width ratio 1:1), reflexed at apex, clypeolateral ridge present from antennal insertions high, linear, parallel-sided, angulate to anterolateral angles and continuous across slightly sinuate anterior margin. Lateral declivity deep in apical half, striolate; apical declivity deep and micropunctate, both angulate and visible in dorsal view. Disc flat, abruptly inclined to highly elevated clypeolateral and apical ridge. Surface coarsely punctate, bearing very long, yellow pilosity. Clypeofrontal suture absent. Frons flat, with coarse punctures bearing dense, very long, yellow pilosity. Ocular canthus narrow, bearing short, yellow setae. Antennae with scape black and apex brown, elongate, bearing few, short, pale setae along dorsum of length, dorsum of apex bearing long, black setae, posterior margin bearing dense setal brush of long, yellow pilosity. Pedicel black, apex brown bearing sparse, pale microsetae. Antennomere 3 elongate, same length as antennomeres 4 and 5 combined. Antennomeres 3–7 black with apices dark brown and bearing sparse, long, pale setae. Club dark brown, length of antennomeres 2–7, 0.25 x length of head. Antennomere 8 internal surface bearing setae along dorsal surface in apical half. Antennomere 10 external surface dark brown with ventral third light brown. Thorax. Pronotum black with wide (¼ width) brown maculae laterally along entire margin encircling small black macula at midlength. Transverse with basomedian margin arcuate, basolateral margin linear and angled anteriorly. Basolateral angle narrowly arcuate through 90°. Posterolateral margin linear, slightly attenuate to broadly arcuate midlength then linear attenuate to anterolateral angles. Anterior margin linear. Lateral margins bearing ridges. Disc with sparse micropunctures becoming larger to all margins, rugulose along lateral margins bearing sparse, long, pale pilosity. Scutellum black, equilateral triangle with coarse punctures except along midline, bearing long, pale pilosity. Elytra yellow-brown, sutural costa dark brown-black, lateral margins narrowly dark brown. Mesepimeron narrowly exposed, with base of elytra laterally pronounced. Posthumeral arch linear in dorsal view, arcuate in lateral view, weakly exposing metacoxa, obscured by pilosity, then slightly attenuate to broadly arcuate apex. Disc (interstices 1 and 2) coarsely punctate in weakly-defined rows becoming coalesced, then rugulose at apical declivity, laterad to costa 2 rugulose; bearing long pale setae. Humeral and apical umbones raised. Costa 1 broadest, costa 2 narrower, costa 3 barely recognizable. Sutural costa parallel basally, raised and attenuate in apical half with apices truncate. Mesometasternal process black, broad, flat with broadly arcuate apex, declivous at anterior margin of mesocoxa; surface impunctate, glabrous with apical declivity bearing dense, very long, yellow pilosity. Metasternum black, medially with moderately sparse punctation, laterally rugulose bearing long, yellow pilosity. Mesepimeron and metepisternum black, rugulose bearing long, yellow pilosity, metepimeron black and impunctate. Legs. Black with dark brown suffusions. Profemora dorsoventrally flattened, slightly attenuate to apex with ventral surface rugulose bearing yellow pilosity. Protibiae elongate parallel, unidentate with apical denticle elongate acute; dorsal surface with medial and internal row of coarse punctures bearing sparse, long, pale setae and setal brush composed of moderately dense, short, ginger setae; spur stout, not reaching apex of tarsomere 1. Protarsomeres 1–5 slightly longer than tibial length; tarsomere 5 longest, 2 x length of tarsomere 4. Tarsomeres 1–5 bearing setae less than half tarsomere length. Protarsal claws long, simple and symmetrical. Mesofemora dorsoventrally flattened, parallel with ventral surface rugulose bearing very long, yellow pilosity. Mesotibiae elongate, ventral surface medially with 2 rows of coarse punctures bearing short, yellow setae in basal half. Internal margin basally bearing sparse, short, pale setae. External margin basally evenly arcuate, angulate mid-length, concave apically, non-denticulate, apex bispinose with spines acute and external spine longer. Internal surface bearing rows of very long, yellow pilosity. Spurs narrow, attenuate with acute apex, reaching midlength of mesotarsomere 2. Mesotarsomeres 1–5 longer than tibial length. Tarsomere 5 longest, 2 x length of tarsomere 4. Mesotarsomeres 1–4 bearing setae half tarsomere length. Mesotarsal claws long, simple and symmetrical. Metacoxae black, posterolateral angle 90°, surface rugulose, bearing long, yellow pilosity. Metafemora dorsoventrally flattened with anterior margin arcuate and posterior margin linear. Ventral surface medially sparsely punctate, rugulose along margins bearing very long, pale pilosity. Metatibiae elongate, ventral surface with row of coarse punctures along internal margin and rugulose adjacent to external margin bearing sparse, long, yellow pilosity. Internal margin bearing sparse, short, yellow setae. External margin medially with low, very obtuse angle, non-denticulate, glabrous. Dorsal surface bearing dense, very long, yellow pilosity. Metatibial apex bispinose with internal spine short, truncate and external spine broad, truncate. Spurs parallel, apex acute, internal spur broader and longer reaching apex of tarsomere 2. Metatarsomeres 1–5 equal to tibial length; tarsomere 5 longest, 2 x length of tarsomere 4. Tarsomeres 1–4 bearing setae half of tarsomere length. Claws long, simple and symmetrical. Abdomen. Sternites black. Sternites 2–6 medially with broad shallow depression, concave in lateral view, with row of sparse punctures bearing short, pale setae and laterally with dense punctures bearing long, yellow pilosity. Sternite 7 medially impunctate and glabrous, laterally rugulose bearing long, yellow pilosity. Propygidium bearing long, yellow pilosity. Pygidium dark yellow with apex broadly black and base narrowly black extending medially to disc. Pygidium length:width ratio 1:1.5, angulate in lateral view, with a narrow, arcuate, slightly obtuse angle post midlength; surface concentrically rugulose with centre post midlength bearing sparse, short, pale setae at apex and basolateral angles. Genitalia (Fig. 10B). Phallobase widest at apex, attenuate to and narrowest at base near parameres, dorsal surface with a longitudinal deep groove and 2.15 x length of parameres. Parameres narrow, external margin sinuate concave medially, arcuate to apex. Each paramere narrowing to midlength, divergent to pre-apex then attenuate to apex. Dorsal cleft 45° at base, then broad, ovate in basal two-thirds, apically attenuate to where parameres meet. Paramere in lateral view arcuate through obtuse angle at basal third; surface sparsely punctate. Variation in males. Length: 12.5–17.8 mm, width 6.5–9.4 mm. Antennomeres 3–7 mostly brown. Pronotum with lateral maculation at anterior and posterior ends traversing towards midline or reduced in small specimens to posterolateral margins. Disc with evenly spread macropunctures and lacking micropunctures. Elytral apices microspinose. Sutural maculation expanded at midlength then laterally covering apical declivity. Protibiae external margin bisinuate or bidentate. Mesotibiae external margin unidentate or bidentate. Metatibiae external margin bearing obtuse denticle. Pygidium black maculation not medially extended. Female (Fig. 9). Description based on CET0381 [PMH] (Lake Deborah, 40 km NE of Bullfinch, Western Australia). Differs from male in the following characters. Length: 18.9 mm, width 10.0 mm. Stout, broader and deeper. Head. Anterior margin weakly arcuate convex. Clypeal disc slightly elevated along midline, sparsely microsetose. Frons sparsely microsetose basolaterally. Antennal scape black with antennomere 10 external surface with ventral quarter brown. Dorsum of scape glabrous. Pedicel glabrous. Antennal club slightly shorter than antennomeres 2–7. Thorax. Pronotum uniformly black. Basomedian margin sinuate. Basolateral angle more broadly arcuate through 90°. Lateral margin broadly arcuate at midlength. Pronotum micropunctate medially becoming macropunctate and rugulose laterally. Pronotal and metasternal setae ash-grey to black. Elytra. Black, setae black. Posthumeral arch sinuate in lateral view, distinctly exposing metacoxa. Humeral and apical umbones slightly raised. Costae weaklydefined, barely visible. Metepimeron rugulose bearing yellow pilosity. Legs. Black. Protibiae broader and tridentate. Denticles long, acute, medial denticle at midlength and closest to basal denticle. Dorsal surface internal half with coarse, denser punctures. Mesotibiae ventral surface with punctures restricted to basal half; external margin midlength bearing robust, acute denticle with smaller basal denticle, concave post midlength; apex divergent with spines broad. Mesotarsomeres 1–5 same length as tibia. Metatibiae external margin bearing acute or truncate denticle at midlength; apex divergent. Metatibial spurs parallel with apices arcuate, dorsal spur surpassing apex of metatarsomere 2. Metatarsomeres 1–5 shorter than tibial length. Abdomen dark brown. Sternites convex. Sternite 6 medially with 2 rows of punctures bearing long setae. Sternite 7 medially along posterior half densely micropunctate. Pygidium black; length:width ratio 1:1.8. In lateral view broadly arcuate angle post midlength. Setose across base. Variation in females. Length 15.1–18.9 mm, width 7.6–10.0 mm. Head with anterior margin linear. Antennomeres 1–10 brown. Elytra dark brown. Legs dark brown. Pygidium with 2 lateral, oblique, elliptical yellow maculae. Diagnosis. Rigoutorum jakli can be recognised by its pronotal sculpturing consisting of an impunctate region on the disc, surrounded with micropunctation and radiating outwards, replaced by spaced macropunctures that bear long pilosity. The pronotum in males is shiny black to anterior margins with lateral yellow maculation (rarely complete in minor males), one-quarter width of pronotum, usually extending from basal to anterior margins. Females are uniformly black. Differential diagnosis. Rigoutorum jakli is similar to R. bacchusi but differs in having a shiny pronotum with impunctate disc surrounded with micropunctures, then spaced macropunctures. Discussion. Most of the specimens are consistent morphologically and in colour with a few beetles exhibiting unusual variation (e.g. bidentate protibiae, expanded elytral maculation). The following are noteworthy. Specimens from the northwest of the range of the species (53 km north-west of Mount Magnet, 30 km west of Kirkalocka Station) lack pronotal micropunctures surrounding the impunctate region on the disc; these are replaced by spaced macropunctures. All four specimens examined from Kirkalocka have a bisinuate protibial external margin, and bidentate mesotibial external margin. Natural history. Rigoutorum jakli are found in mallee (Eucalyptus) heath on yellow sandplains with dominant large plants mainly consisting of Eucalyptus, Allocasuarina, Acacia and Melaleuca (Myrtaceae). Adults are localised, and males are encountered in flight. Only three specimens were seen on flowers, namely Eucalyptus, even though plants were flowering nearby. Males regularly flew at the top of taller plants (Eucalyptus, Allocasuarina, Acacia) at a height of 1.8–3.7 m. They frequently flew to a neighbouring plant where they circled the top once or twice and then proceeded to the next tree. The behaviour was then repeated. We assume that males were searching for females. Adult males were never noticed flying low to the ground or with a female. Populations of over 100 individuals were observed. On 27 January 2004, along the Wannara-Perenjori track, 2 km west of the Great Northern Highway, where many males were seen circling the tops of mallee, a single male was observed entering a hole in the ground beside a mallee (Eucalyptus). Only one female was found alive, perched on leaves of a Eucalyptus. No males were observed nearby. Adults have been recorded from 13 January to 11 February (based on data from specimens and PMH trip diaries). Two specimens from Lake Baladjie Nature Reserve, caught in a bucket trap, have label data showing a collection date from 20 December to 19 January. In several instances, specimens were located dead on the edge of salt lakes (“strandlines”). In years when rain fills the salt lakes, individuals may be washed into the lake or land on the lake and drown. In these cases, salt lakes act as natural pitfall traps, and can be used as an indicator of the presence of some local fauna. Strandline specimens deteriorate rapidly due to predation and the detrimental effects of salt and sun. However, if collected within a few days, these effects are minimal, and specimens can be assumed as recently deceased. Etymology. This species is named in honour of Mr. Stanislav (Stan) Jákl (Prague, Czech Republic), an intrepid and prolific researcher of Cetoniinae and Dynastinae. Stan suggested that the specimens that PMH captured north of Wubin, Western Australia may represent an undescribed species., Published as part of Hutchinson, Paul M., Moeseneder, Christian H. & Mitchell, Andrew, 2022, Rigoutorum a new genus for Diaphonia bacchusi Rigout and Allard (Coleoptera: Scarabaeidae: Cetoniinae: Schizorhinini), with description of a second species from Western Australia and notes on habitats and behaviours, pp. 239-259 in Zootaxa 5150 (2) on pages 253-258, DOI: 10.11646/zootaxa.5150.2.4, http://zenodo.org/record/6621254, {"references":["Golding, M. R. (2009) A Pictorial Field Guide to the Beetles of Australia. Part 5: Cetoniidae. Total Digital Solutions, Burswood, Western Australia, 48 pp."]}

Key to species of Rigoutorum 1. Pronotum evenly, densely punctate. Pronotal pilosity dense, long. Males with pronotal lateral margin with maculation narrow and not full-length, or absent. Southeast region of southern Western Australia......... R. bacchusi (Rigout & Allard, 1997) - Pronotum sparsely punctate medially, combination of micro and macro punctures. Pronotal pilosity sparse, shorter. Males with pronotal lateral margin with full-length maculation, 0.25 x width of pronotum (rarely not reaching anterior margin). Northwest region of southern Western Australia....................................................... R. jakli new species, Published as part of Hutchinson, Paul M., Moeseneder, Christian H. & Mitchell, Andrew, 2022, Rigoutorum a new genus for Diaphonia bacchusi Rigout and Allard (Coleoptera: Scarabaeidae: Cetoniinae: Schizorhinini), with description of a second species from Western Australia and notes on habitats and behaviours, pp. 239-259 in Zootaxa 5150 (2) on page 244, DOI: 10.11646/zootaxa.5150.2.4, {"references":["Rigout, J. & Allard, V. (1997) Les Coleopteres du Monde. Vol. 25. Schizorhinini 3. Hillside Books, Canterbury, 128 pp."]}

Hemitelsella Smith, 2016 Hemitelsella Smith, 2016: 72. Type species: Acrotelsella transpectinata Smith, 2015, by original designation., Published as part of Smith, Graeme B., Mitchell, Andrew & Mesaglio, Thomas, 2022, Two New Australian Silverfish (Zygentoma: Lepismatidae: Ctenolepismatinae and Nicoletiidae: Subnicoletiinae), pp. 59-74 in Records of the Australian Museum (Rec. Aust. Mus.) (Rec. Aust. Mus.) 74 (2) on page 61, DOI: 10.3853/j.2201-4349.74.2022.1796, http://zenodo.org/record/7178146, {"references":["Smith, G. B. 2016. On some silverfish taxa from Tasmania (Zygentoma: Lepismatidae and Nicoletiidae). Records of the Australian Museum 68 (2): 45 - 80. https: // doi. org / 10.3853 / j. 2201 - 4349.68.2016.1652"]}

Family Nicoletiidae Lubbock, 1873 Nicoletiadae Lubbock, 1873: 201. Gymnodermata Joseph, 1882: 25. Nicoletiinae Lubbock. — Escherich, 1905: 36. Nicoletiidae Lubbock. — Remington, 1954: 284., Published as part of Smith, Graeme B., Mitchell, Andrew & Mesaglio, Thomas, 2022, Two New Australian Silverfish (Zygentoma: Lepismatidae: Ctenolepismatinae and Nicoletiidae: Subnicoletiinae), pp. 59-74 in Records of the Australian Museum (Rec. Aust. Mus.) (Rec. Aust. Mus.) 74 (2) on page 68, DOI: 10.3853/j.2201-4349.74.2022.1796, http://zenodo.org/record/7178146, {"references":["Lubbock, J. 1873. Monograph of the Collembola and Thysanura. London: The Ray Society, 276 pp. https: // doi. org / 10.5962 / bhl. title. 11583","Joseph, G. 1882. Systematisches Verzeichniss der in den Tropfstein - Grotten von Krain einheimischen Arthropoden nebst Diagnosen der vom Verfasser entdeckten und bisher noch nicht beschriebenen Arten. Berliner Entomologische Zeitschrift 26 (1): 1 - 50.","Escherich, K. 1905. Das System der Lepismatiden. Zoologica (Stuttgart) 43: 1 - 164. https: // doi. org / 10.5962 / bhl. title. 7909","Remington, C. L. 1954. The suprageneric classification of the order Thysanura (Insecta). Annals of the Entomological Society of America 47: 277 - 286. https: // doi. org / 10.1093 / aesa / 47.2.277"]}

Heterolepisma Escherich, 1905 Heterolepisma Escherich, 1905: 63. Isolepisma Escherich, 1905: 61. Notolepisma Tillyard, 1924: 242. Type species: Lepisma pampeana Silvestri, 1902 by subsequent designation, see Paclt, 1967: 25., Published as part of Smith, Graeme B. & Mitchell, Andrew, 2019, Species of Heterolepismatinae (Zygentoma: Lepismatidae) Found on some Remote Eastern Australian Islands, pp. 139-181 in Records of the Australian Museum 71 (4) on page 143, DOI: 10.3853/j.2201-4349.71.2019.1719, http://zenodo.org/record/4653485, {"references":["Escherich, K. 1905. Das System der Lepismatiden. Zoologica (Stuttgart) 43: 1 - 164.","Tillyard, R. J. 1924. Primitive wingless insects. Part I. The silverfish, bristletails and their allies (Order Thysanura). New Zealand Journal of Science and Technology 7: 232 - 242.","Silvestri, F. 1902. Materiali per lo studio dei Tisanuri. I-V. Bollettino della Societa Entomologica Italiana 33: 204 - 249.","Paclt, J. 1967. Thysanura. Fam. Lepidotrichidae, Maindroniidae, Lepismatidae. Genera Insectorum 218 e: 1 - 86."]}

Maritisma dispar (Uchida, 1944) n. comb. Heterolepisma dispar Uchida, 1944: 185. Heterolepisma dispar, a species inhabiting coastal cliffs from the Japanese Island of Honshu, appears, from the original description and illustrations, to share many of the unusual characters such as the macrochaetae along the anterior margin of the frons, the short urotergite X, the long, spaced urosternal combs and the long parameres. The chaetotaxy of urosternite I was not mentioned but the species shares so much of the unusual morphology of H. coralinium sp. nov. that it is likely to also possess a medial comb. Both species lack a glabrous anterior margin to the frons, both have a very broad ultimate article to the labial palp, urotergite X is greatly reduced and they have unusually long urosternal combs with widely spaced macrochaetae. These characters differentiate them from the remaining species with medial combs on urosternite I. However, the illustration of Uchida suggests that the pronotal collar is complete in H. dispar. This Japanese species is here provisionally transferred to the new genus., Published as part of Smith, Graeme B. & Mitchell, Andrew, 2019, Species of Heterolepismatinae (Zygentoma: Lepismatidae) Found on some Remote Eastern Australian Islands, pp. 139-181 in Records of the Australian Museum 71 (4) on page 171, DOI: 10.3853/j.2201-4349.71.2019.1719, http://zenodo.org/record/4653485, {"references":["Uchida, H. 1944. Die Bestatigung der nicht bescriebenen Art, Heterolepisma dispar Silv. (Thysanura). Annotationes Zoologicae Japonenses 22 (4): 185 - 189."]}

Maritisma gen. nov. http://zoobank.org/NomenclaturalActs/ 56115144-6890-4115-88A2-9716C9C5EF95 Type species. Maritisma coralinium sp. nov. Diagnosis. Medium-sized silverfish. Body shape as in Fig. 169. Pigment apparently absent or very limited e.g., around eyes. Macrochaetae smooth with apical bifurcations. Scales multiradiate, rounded with ribs not very close together; lanceolate or triangular scales apparently absent.Antennae possibly with rod-like basiconic sensilla. Chaetotaxy of frons consisting of a row of macrochaetae along anterior margin joining laterally with rows along the sides of the head which run back along the margin and up over the eyes, peri-antennal groups weak. Clypeus with longer macrochaetae laterally and across the face proximally, with small setae medially. Labrum with many macrochaetae distributed across the proximal half and some smaller setae in the anterior half. Eyes of about 12 ommatidia. Apical article of labial palp broad with 3+2 papillae of the aufgelöst type (individual subunits of each papilla not packed closely together), those more distal very long. Pronotum with narrow setal collar which is largely interrupted medially. Thoracic nota with marginal setae and a few submarginal combs each of just a single or two macrochaetae; each lateral margin with two open trichobothrial areas, each anterior trichobothrial area of the pronotum with a macrochaeta laterad of the trichobothrium; posterior margin with 1+1 combs each of a single macrochaeta. Thoracic sternites free, sub-parabolic with submarginal macrochaetae along the more distal lateral regions. Legs typical for the Heterolepismatinae; tarsi with four articles, pretarsi with two claws and a medial empodial claw. Urotergites I–VII with 3+3 small combs, urotergite VIII with 2+2 combs, urotergite IX with 1+1 sublateral setulae. Urotergite X very short, rounded with a few macrochaetae along each lateral margin. Urosternite I with medial comb, urosternites II–VIII with 1+1 long combs, each macrochaeta of all combs distinctly spaced apart from adjacent macrochaetae. Coxites IX of both sexes typical for the Heterolepismatinae, with large, prominent parameres. One pair of styli only (IX). Ovipositor of female simple. Etymology. The genus name derives from the Latin word "maritimus" referring the proximity of known collection sites to the ocean combined with the suffix -isma, used for many silverfish. It is treated as grammatically neuter. Remarks. When initially examined by the first author, the specimen did not immediately appear to belong to the Heterolepismatinae, however with due consideration, most characters were generally in agreement with those associated with the Heterolepismatinae. Molecular data could not be obtained for this species due to the length of time the specimens had been in 70% ethanol. The character analysis places M. coralinium sp. nov. closest to the H. stilivarians group predominantly because both share the presence of macrochaetae rather than setae on the labrum, however the species share little else in common. The stilivarians group is currently under revision and the available molecular data places it quite distant from the remaining Australian species of Heterolepisma for which molecular data is available. The new genus can easily be distinguished from all described Heterolepismatinae (except H. dispar – see below) by the following combination of characters: the presence of chaetotaxy on the anterior margin of the frons, the absence of branched papillae on the ultimate article of the maxillary palp, the presence of 3+3 combs on urotergite I, the presence of a medial comb on urosternite I, the very long 1+1 combs of eight or more spaced macrochaetae on urosternites II–VI, the number of pairs of styli (one pair in both sexes), the wide ultimate article of the labial palp and the much shorter urotergite X., Published as part of Smith, Graeme B. & Mitchell, Andrew, 2019, Species of Heterolepismatinae (Zygentoma: Lepismatidae) Found on some Remote Eastern Australian Islands, pp. 139-181 in Records of the Australian Museum 71 (4) on pages 170-171, DOI: 10.3853/j.2201-4349.71.2019.1719, http://zenodo.org/record/4653485

Heterolepisma sclerophylla Smith, 2014 Fig. 7 Heterolepisma sclerophylla Smith, 2014: 9. Type material examined. Holotype. 1♀ (HW 1.28) (AMS K.260990, K.260991 on two slides) NSW: Broulee, 35.8578��S 150.1621��E 15 m asl, 17.xi.2010, Graeme Smith. Paratypes. 1♀ (HW 1.23) (AMS K. 377563 in ethanol) same data as holotype; 1♀ (HW 1.25) (AMS K.261218, K.261219 on two slides) same data as holotype; 1♂ (HW 1.25) (AMS K.260992, K.260993 on two slides) same data as previous; 1♂ (HW 1.23) (AMS K. 377565 in ethanol) same data as holotype; 1♂ (HW 1.13) (AMS K. 377566 in ethanol) same data as holotype; 1♀ (HW 1.15) (AMS K. 377567 in ethanol) same data as holotype; 1 juvenile ♀ (HW 0.84) (AMS K. 377568 in ethanol) same data as holotype; 1♂ (HW 1.15) (AMS K. 377569 in ethanol) same data as holotype; 1♂ (HW 1.10) (AMS K.261074, K.261075 on two slides) same data as holotype; 1♀ (HW 1.08) (AMS K. 377570 in ethanol) same data as holotype; 1♀ (HW 1.20) (AMS K. 377561 in ethanol) same locality and collector as holotype 19.xi.2010; 1♀ (HW 1.40) (AMS K. 377562 in ethanol) same data as previous. Other topotypic material examined. 1♀ (HW 1.20) (AMS K.261216, K.261217 two slides) same locality as holotype, 18.ii.2016, Graeme Smith; 1♀ (HW 1.10) (AMS K.261142 head, thorax and abdominal segments I���IV on one slide); 1♀ (HW 1.10) (AMS K.261143, K.261144 on two slides) same data as previous; 1♂ (HW 1.13) (AMS K.261145, K.261149 on two slides) same data as previous; 1♀ (HW 1.13) (AMS K.261152, K.261153 on two slides) same data as previous; 1♀ (HW 1.18) (gbs004988 in 100% ethanol) same data as previous; 1♀ (HW 1.03) (AMS K.261150, K.261151 on two slides) same data as previous. Material from other localities of the same lineage. 1♂ (HW 1.18) (AMS K.260994, K.260995 two slides) NSW: Guerilla Bay headland, 35.8321��S 150.2313��E 87 m asl, 17.xi.2010, Graeme Smith; 1♀ (HW 1.23) (AMS K.261202, K.261203 on two slides) NSW: Jibbon, 34.0781��S 151.1674��E 18 m asl, 1.i.2016, Graeme Smith; 1♀ (HW 1.00) (AMS K.261184, K.261185 on two slides) same data as previous. Comments. The Broulee group has a comparatively short ovipositor both relative to head width (1.49���1.95 HW) and in the number of divisions (32���37). Most other lineages have ovipositors in mature specimens exceeding twice the head width and with 35���41 divisions. It also seems to have lower levels of pigment overall and the density and strength of the macrochaetae on the head and notal collar is less than for other lineages (Fig. 7). The Nowra lineage has a similarly short ovipositor but appears to have much darker pigment on the labial and maxillary palps and to a lesser extent on the legs. It has denser chaetotaxy along the sides of the head near the eyes (2���4 rows wide versus 2���3 rows wide). The ovipositor length was reported in Smith (2014) as up to 2.09 times head width. This was based on the result from a specimen (originally recorded as K. 377564 in ethanol but now replaced by K. 261219 as slide) and was quite different to other specimens measured (range 1.49���1.79). This specimen was re-measured and the measurement is here corrected to 1.95 times HW. This is still much longer than other specimens from Broulee but the specimen only has 35���37 divisions which is within the normal range for the Broulee specimens (32���37). Each individual division is longer than seen with other specimens of this lineage. The lineage has been collected from three localities along the central and southern coasts of New South Wales. All localities listed have sandy soils, but this may represent a sampling bias rather than be a true indication of habitat. In the type locality it is quite common among dry Eucalypt leaves caught between the fronds of cycads (Macrozamia sp. Zamiaceae). At Guerilla Bay it was collected within abandoned termite galleries on a standing tree and at Jibbon it was collected from dry leaf litter protected from rain beneath a fallen tree or in a large hollow at the base of the tree. It was occasionally found hiding within the bark of certain spongy-barked Eucalyptus or Corymbia trees using pyrethrum sprays, where it was usually collected together with a more abundant (but not yet described) species related to Heterolepisma highlandi Smith. One Broulee specimen (K. 261145) was unusual in that one of its maxillary palps was shorter than usual and had several plumose sensilla along its length rather than the usual three, as found on the other palp where they were more evenly spaced along the article rather than in the distal half. In all specimens examined the plumose sensilla on the palps of the males are much larger (2���3 times larger diameter) and more elaborate than those on the female., Published as part of Smith, Graeme B., Mitchell, Andrew, Lee, Timothy R. C. & Espinasa, Luis, 2019, DNA Barcoding and Integrative Taxonomy of the Heterolepisma sclerophylla species complex (Zygentoma: Lepismatidae: Heterolepismatinae) and the Description of Two New Species, pp. 1-32 in Records of the Australian Museum 71 (1) on page 14, DOI: 10.3853/j.2201-4349.71.2019.1677, http://zenodo.org/record/3837977, {"references":["Smith, G. B. 2014. Two new species of Heterolepisma (Zygentoma: Lepismatidae) from eastern New South Wales. General and Applied Entomology: The Journal of the Entomological Society of New South Wales 42 (2013): 7 - 22 (published 2014)."]}

Heterolepisma coorongooba sp. nov. Figs 6, 8���36 Holotype. ♀ (HW 1.30) (AMS K.261208, K.261209 on two slides) NSW: Glen Davis, above Coorongooba campground, 33.1271��S 150.3232��E 313m asl, 20.vi.2015, Graeme Smith. Paratypes. 2♀♀, 1♂, 1 juvenile, all same data as holotype including 1♀ (HW 1.15) (AMS K.261210, K.261211 on two slides); 1♀ (HW 1.11) (AMS K.261212, K.261213 on two slides); 1♂ (HW 1.08) (AMS K.261204, K.261205 on two slides); 1 juvenile (HW 0 3) (AMS K. 377726 in ethanol). Diagnosis. This species is very similar to Heterolepisma sclerophylla differing in having one fewer pairs of styli in both the male and female (i.e. IX only in ♂, VIII and IX in the ♀). Compared to the lineage from the type locality it also has a longer ovipositor and thicker and perhaps more densely packed chaetotaxy along the pronotal collar and margins of the head. Description Appearance: Medium to large silverfish, scale covering in life uniform or slightly mottled grey with brown antennae, terminal filaments brown with lighter annuli around larger macrochaetae resulting in distinctly banded appearance (Fig. 6). Body length: H+B up to 9.1 mm (♀) 7.0 mm (♂); maximum HW 1.30 mm; thorax: length up to 2.5 mm (or 0.27���0.36 H+B); width up to 2.05 mm, usually slightly widest at the mesonotum; antennae damaged in all specimens, maximum preserved length of antenna 4.4 mm (or 0.56 H+B); terminal filaments damaged in all specimens, maximum preserved length of cercus 2.8 mm (or 0.31 H+B); maximum preserved length of median dorsal appendage 3.4 mm (or 0.42 H+B). Body neither elongate nor broad with thorax slightly wider than abdominal segment I, the following abdominal segments about the same width until the fourth or fifth after which the abdomen tapers posteriorly. Pigmentation: Pigment brown in alcohol preserved specimens. Present around eyes and to a lesser extent behind the peri-antennal group of macrochaetae; pedicel and scape very lightly pigmented distally, rest of flagellum uniformly lightly pigmented; all articles of maxillary palp with pigment especially in the four most distal articles although much less in the ultimate article; labium with pigment on the three most distal articles, being strongest on the penultimate, especially distally. Pigment present in anterior corner and along margins of all nota. Legs not heavily pigmented, with light pigment along outer margin of the coxa and the trochanter, apically on the outer femur and strongest on the tibia especially on the dorsal surface, present on basal article of tarsi, urotergite X and coxites IX with light pigmentation; styli IX pigmented in distal three quarters; other styli with less pigment. Ovipositor with yellowish hue. Terminal filaments with rings of pigment, with the pigment present in all annuli except the annuli bearing the large macrochaetae. Pigmentation is much reduced in juvenile specimens. Macrochaetae: Bifid apically, or simple, light to darker brown in colour. Often quite thick e.g., the stronger macrochaetae of the pronotal collar measured about 50 microns in diameter in the holotype (HW 1.30) compared to 38 microns in the similar-sized holotype of H. sclerophylla (HW 1.28) from Broulee (compare Figs 7, 8). They also appear to be more densely packed. Scales: Unevenly rounded or ovoid, with numerous parallel ribs that do not extend beyond the margin (Fig. 9); in alcohol dorsal scales with dark brown ribs; ventrally mostly hyaline. Lanceolate scales not observed. Scales absent from flagellum of antennae, mouthparts and terminal filaments. Head: Wider than long (Fig. 10) with marginal rows about three macrochaetae wide along the sides of the vertex decreasing to two wide in front of the antennae and six strong curved macrochaetae along the anterior margin, the lateral rows extend back above the eyes, as well as a small 1+1 peri-antennal groups not quite isolated from the marginal rows at the level of each antenna. Clypeus with numerous setae, some long and thin, others more robust arranged in 1+1 lines in the proximal lateral regions but not forming combs. Labrum with thin setae only. Eyes dark, composed of about 12 ommatidia.���Antennal scape with a subdistal rosette of setae, very conspicuous from above (Fig. 11), and numerous setae along the sides and over the ventral face; pedicel short, 0.51 times the length of the scape (range 0.47���0.55), with many setae mostly distally and on the ventral face; repeating intervals of distal end of antennae (Fig. 12) of eight annuli, the most apical annulus of each interval (T-annulus) with a trichobothrium and at least one small inconspicuous rod-like basiconic sensillum (type B of Adel, 1984), the second and forth annuli also each with a sausage-shaped type C sensillum (confirmed also to be present on same annuli in holotype of H. sclerophylla).���Mandibles typical for genus with welldeveloped molar and incisor areas; a group of about nine strong setae distally adjacent to the pectinate molar area and a bush of 50+ setae and macrochaetae externally.���Maxilla (Fig. 13) with three large macrochaetae externally proximal to the palp, the lacinia with three strong teeth, one shorter than the rest, seven lamellate processes and a row of eight simple setae, the galea longer than the lacinia with setulae on the outer face. Palp with rosettes of distinctly stronger setae (some ���carrot-shaped���) subapically on the three basal articles, all articles with numerous fine setae, apical article of maxillary palp 4.6 times longer than wide (range 4.1���4.9) and 1.23 times longer than penultimate article (range 1.16���1.27), the ultimate article in both sexes with three ���branched��� papillae, those in the female less robust than those in the male.���Labium (Fig. 14) short and broad with rows of strong setae on the prementum and submentum; glossae and paraglossae quite broad with short curved setulae; labial palp short, apical article eccentric suboval, 1.1 times as long as wide (range L/W 0.8���1.3) with 2+3 papillae of compact type in a ���cluster formation��� where the slightly larger distal papillae curve around the two smaller proximal papillae and a curved club-like thin-walled basiconic sensillum and at least one rod-like basiconic sensillum, which can also be confirmed as present on the holotype of H. sclerophylla. Thorax: Pronotum (Fig. 15) with strong setal collar of short, apically bifurcated setae and cilia with a row of longer macrochaetae spaced along the back of the collar including 1+1 long thin simple setae about one third in from each side, the density and length of the smaller more marginal macrochaetae of the collar decreases only slightly towards the middle but not the size and density of the larger spaced macrochaetae in the posterior row; lateral margins also with numerous shorter but quite robust, apically bifurcate setae as well as several larger more erect submarginal macrochaetae; trichobothrial areas open and in contact with the lateral margins, the anterior one (Fig. 16) located just anterior to the mid-point along the margin, with or without a large submarginal macrochaeta, when present laterad to the trichobothrium (this macrochaeta missing on the left side of the holotype) and with a cilium and a few setulae; posterior trichobothrial area (Fig. 17) near posterior lateral corner with two submarginal macrochaetae between the trichobothrium and the margin as well as a few setae and setulae; posterior margin slightly concave with 1+1 combs (Fig. 18) each of two macrochaetae, the more postero-mediad lying flatter than the other, each comb associated with a setula and a few cilia.���Mesonotum with lateral chaetotaxy similar to pronotum but less dense (Fig. 19), except the submarginal macrochaetae anterior to the trichobothrial areas are grouped into combs of two, each associated with a cilium and a few setulae, the more posterior of these on the left side of the holotype of only one macrochaeta, both trichobothrial areas (Fig. 20) of similar configuration to those of pronotum except anterior area has a macrochaeta mediad of the trichobothrium. The posterior combs of the mesonotum are unusual in that they both consist of only a single macrochaeta in the illustrated holotype (Fig. 21) whereas in the paratype K. 261204 one consists of two macrochaetae and the other of only one and in the other paratypes (K. 261213 and K. 261210) two macrochaetae are present on both sides.���Metanotum (Figs 22, 23) similar to mesonotum; both posterior combs consist of two equal sized insertion points. The trichobothrium appears to be absent from the right posterior area of the holotype. a Small infralateral setae b More posterior seta insertion smaller Presternum narrow, with transverse row of strong setae and numerous cilia and setulae (Fig. 24).���All thoracic sterna and coxae with hyaline scales. Prothoracic sternum pointed cordiform, almost as long as wide at its base (range L/W 0.91���1.00) and reaching to about two thirds the length of the coxa, rounded apically and with a medial furrow (Fig. 24), most of lateral margins with numerous small marginal setae and cilia, with 6���7 larger submarginal macrochaetae forming weak combs parallel to the edges in the distal third.���Mesosternum (Fig. 25) 1.08 times longer than broad (range 1.01���1.12) with an acutely rounded apex, with setae and cilia along the distal quarter of the margins, with 1+1 distal combs of four apically bifurcate macrochaetae and 1+1 subposterior more pointed macrochaetae.���Metasternum (Fig. 26) 0.79 times longer than wide (range 0.77���0.80) with less pointed, even slightly concave, apex, each comb of four macrochaetae. Legs fairly long (Figs 24, 26), tibia L/W ratio of legs PI 3.0 (range 2.9���3.2), PII 3.4 (range 2.8���4.0), PIII 3.9 (range 3.5���4.2); tarsi L/W ratio PI 7.0 (range 6.3���8.0), PII 7.0 (range 6.3���7.7), PIII 8.5 (range 7.4���9.7). Legs increasingly longer from front to back, mean ratio PI/PIII (tibia 0.62, tarsus 0.72). PI with transverse comb of about six macrochaetae laterally on the precoxa. Coxa of all legs covered with hyaline scales and with strong macrochaetae and numerous cilia and setulae in a row about two macrochaetae wide along the external margin, a strong seta on the inner margin subapically and group of about six curved setae at the apex over the articulation. Trochanter lacking scales, with fine and one stronger seta over the surface. Femur with numerous setae over most of the surface and along the margin; anterior distal end with two strong, quite deeply bifurcate stout macrochaetae and another simple stout macrochaeta more proximal; posterior margin with several strong macrochaetae as illustrated. Tibia with numerous long setae over the ventral surface, with three stout macrochaetae on or near the anterior margin and six or seven stout macrochaetae along the posterior margin; apical spur with several setae. Tibia of PIII with a long thin, laterally projecting trichobothria-like seta inserted dorsal to the proximal stout macrochaeta on the anterior margin, which is about two times as long as the tibia is wide. Tarsus with four articles, all with numerous setae, some on the ventral surface quite long and strong. Pretarsus with long curved lateral claws and a strong curved shorter medial claw. Abdomen: Urotergite I usually with 2+2 combs each of one to three macrochaetae (usually two) located quite close together, urotergites II-VII with 3+3 combs of macrochaetae as in Table 7 (Figs 9, 27���29) noting that the more posteromedial insertion point of each submedial comb was occasionally, quite a bit smaller than the other insertion point, but mostly of about the same size; each comb also associated with up to five marginal setae, five setulae and four cilia. Urotergite VIII with 2+2 combs, lacking the sublateral comb; urotergite IX (Fig. 30) with two infralateral setae on each side as well as a setula and a few cilia. Urotergite X short, parabolic in both sexes (Fig. 31), L/W at base about 0.52 (range 0.51���0.55) with many strong setae along entire margin and obscure 1+1 submarginal macrochaetae in the posterolateral corners. Urosternite I glabrous, urosternites II���VIII (Fig. 32) with 1+1 single macrochaetae (Fig. 33), each associated with 0���2 small marginal setae as well as a few cilia and/or setulae. Coxites of segment VIII in ♀ (Fig. 34) with one or two small macrochaetae, one or two small marginal setae and a cilium mediad of the stylus base and some small setae, setulae and a cilium laterad of the stylus base. Styli in two pairs in the ♀ (VIII���IX); all styli with several noticeably longer and stronger setae apically (Fig. 34) as well as stronger setae along the middle of the ventral face. Styli IX 2.4 times as long as styli VIII (range 2.2���2.8). Coxite IX of ♀ (Fig. 34), the internal process acute apically, about three times longer than the external process (range 2.9���3.2) and 1.5 times as long as broad at its base (range 1.5���1.6), not reaching to half the length of the stylus; external and internal margins of internal process and external margin and apex of outer process with many moderately strong setae directed both up and down.���Ovipositor (Fig. 34), very long and thin (up to 2.34 HW), surpassing the apex of stylus IX by more than the length of the stylus (excluding terminal macrochaetae), composed of 34���39 divisions. Distal divisions of gonapophyses VIII and IX with only short fine setae and setulae. Cerci not well preserved in slide material, with basal divisions shorter than long, gradually becoming longer distally, equally wide as long by about the eighth division after which they become even longer with more annuli each with a rosette of setae and some with trichobothria with the large macrochaetae restricted to the most distal annulus of each division; the most distal surviving divisions with up to four annuli, this annulus without pigment.���Medial filament of similar arrangement. Male: As for female except only one pair of styli (segment IX). Coxites IX (Fig. 35) with acute inner process about 1.3 times longer than wide at its base and about three times longer than the external process, reaching to just under half the length of the stylus. Both process also with several strong setae mostly apically emerging from both the dorsal and ventral surfaces of the processes close to or on the margin. Parameres a little longer than wide, with about thirty fine setae (Fig. 36). Penis typical for genus with numerous glandular setae apically, each set on a protuberance. Subadult stages: the single juvenile specimen available, K.377726 (HW 0.63) had a single pair of styli (IX) and no indication of urosternite VIII dividing into separate coxites nor any nascent genitalia (ovipositor or parameres). The thoracic sternites conformed to those of the adults and the long thin setae on the tibia of PIII was present and very long (about three times the width of the tibia), tergite X was round but shorter than in the adults and the feathered papilla of the maxillary palp could not be seen. Habitat. Heterolepisma coorongooba was collected from leaf litter protected from rain under a fallen but still elevated, log. Etymology. The species name is derived from the proper noun Coorongooba referring the creek that flows through the valley from where it was collected. Comments. The morphology of this species is very close to that of H. sclerophylla, differing from it only in the absence of the most anterior pair of styli in both sexes. It differs from the Broulee lineage in the length of the ovipositor (2.17���2.34 times HW versus 1.49���1.95), but not so much in the number of divisions (34���39 versus 32���37) and the more robust macrochaetae. The Megalong, North Nowra and Glenbrook lineages also have more robust macrochaetae and a longer ovipositor (but more divisions) leaving the fewer styli as the only unambiguous defining character., Published as part of Smith, Graeme B., Mitchell, Andrew, Lee, Timothy R. C. & Espinasa, Luis, 2019, DNA Barcoding and Integrative Taxonomy of the Heterolepisma sclerophylla species complex (Zygentoma: Lepismatidae: Heterolepismatinae) and the Description of Two New Species, pp. 1-32 in Records of the Australian Museum 71 (1) on pages 16-22, DOI: 10.3853/j.2201-4349.71.2019.1677, http://zenodo.org/record/3837977, {"references":["Adel, T. 1984. Sensilleninventar und sensillenmuster auf den Antennen von Thermobia domestica und Lepisma saccharina (Insecta: Zygentoma). Braunschweiger Naturkundliche Schriften 2: 191 - 217."]}

Heterolepisma Escherich, 1905 Heterolepisma Escherich, 1905: 63. Type species: Lepisma pampeana Silvestri, 1902 by subsequent designation (Paclt, 1967: 25). Isolepisma Escherich, 1905: 61. Notolepisma Tillyard, 1924: 241., Published as part of Smith, Graeme B., Mitchell, Andrew, Lee, Timothy R. C. & Espinasa, Luis, 2019, DNA Barcoding and Integrative Taxonomy of the Heterolepisma sclerophylla species complex (Zygentoma: Lepismatidae: Heterolepismatinae) and the Description of Two New Species, pp. 1-32 in Records of the Australian Museum 71 (1) on page 14, DOI: 10.3853/j.2201-4349.71.2019.1677, http://zenodo.org/record/3837977, {"references":["Escherich, K. 1905. Das System der Lepismatiden. Zoologica (Stuttgart) 43 (18): 1 - 164.","Paclt, J. 1967. Thysanura. Fam. Lepidotrichidae, Maindroniidae, Lepismatidae. Genera Insectorum 218 e: 1 - 86.","Tillyard, R. J. 1924. Primitive wingless insects. Part I. The silverfish, bristletails and their allies (Order Thysanura). New Zealand Journal of Science and Technology 7: 232 - 242."]}

Heterolepisma highlandi Smith 2014 Heterolepisma highlandi Smith, 2014: 16. Type material (paratype). 1 juvenile ♀ (HW 0.88) (AMS K. 377604 in ethanol) NSW: Wee Jasper, 35.0591��S 148.6489��E 552 m asl, 21.viii.2010, Graeme Smith and Phil Fleming., Published as part of Smith, Graeme B., Mitchell, Andrew, Lee, Timothy R. C. & Espinasa, Luis, 2019, DNA Barcoding and Integrative Taxonomy of the Heterolepisma sclerophylla species complex (Zygentoma: Lepismatidae: Heterolepismatinae) and the Description of Two New Species, pp. 1-32 in Records of the Australian Museum 71 (1) on page 14, DOI: 10.3853/j.2201-4349.71.2019.1677, http://zenodo.org/record/3837977, {"references":["Smith, G. B. 2014. Two new species of Heterolepisma (Zygentoma: Lepismatidae) from eastern New South Wales. General and Applied Entomology: The Journal of the Entomological Society of New South Wales 42 (2013): 7 - 22 (published 2014)."]}

Heterolepisma buntonorum Smith 2016 Heterolepisma buntonorum Smith 2016a: 58. Material examined. 1♀ (HW 1.43) (AMS K.261244 K.261245 on two slides) TAS: Knocklofty, 42.8752��S 147.2957��E 270 m asl, 13.ii.2016, Stephen Bunton., Published as part of Smith, Graeme B., Mitchell, Andrew, Lee, Timothy R. C. & Espinasa, Luis, 2019, DNA Barcoding and Integrative Taxonomy of the Heterolepisma sclerophylla species complex (Zygentoma: Lepismatidae: Heterolepismatinae) and the Description of Two New Species, pp. 1-32 in Records of the Australian Museum 71 (1) on page 14, DOI: 10.3853/j.2201-4349.71.2019.1677, http://zenodo.org/record/3837977, {"references":["Smith, G. B. 2016 a. On some silverfish taxa from Tasmania (Zygentoma: Lepismatidae and Nicoletiidae). Records of the Australian Museum 68 (2): 45 - 80. https: // doi. org / 10.3853 / j. 2201 - 4349.68.2016.1652"]}

Culicoides subgen. Avaritia Fox, 1955: 218. Type species Ceratopogon obsoletus Meigen, by original designation. Imicola Complex Khamala & Kettle 1971: 8. Nominate species: Culicoides imicola Kieffer 1913: 11. Diagnosis (modified from Meiswinkel 2004). Adults: Females: the only species grouping within C. subg. Avaritia with terga on abdominal segments III–V almost round, not wider than long and with sclerotised plates surrounding gonopore truncate, not partially encircling the gonopore. Males: The only species grouping within C. subg. Avaritia with an asymmetrical apex to the anterior ‘peg’ of the aedeagus and with a single STl on antennal flagellomeres 5– 7 coupled with an absence of STl on flagellomere 8. Description. Adults: Eyes bare. Palpus with five segments, each supporting SCh, segment 3 slightly swollen medially with a round, shallow pit (sensory area reduced to a depression in most specimens of C. nudipalpis). Wing relatively strongly patterned. Mesonotum inconspicuously patterned, legs weakly banded, hind tibial comb with five spines, first from spur longest. Wing with cells r 1 and r 2 subequal in length, cell r 2 tip included in post-stigmatal pale spot to varying degrees, lumens not expansive; pale markings large, distributed in all cells. Female. Eyes contiguous for 1–2 facets. Antennae with basal flagellomeres barrel-shaped, distal flagellomeres elongate; SCo distribution 1, 10– 13, in some species also present on 9; basal whorls of two to four SCh on 2–8 with higher numbers on flagellomeres 2, 4, 6 and 8 than on 3, 5 or 7, SCh always absent from 9–13; one pair of STl and one STc present on flagellomeres 2–8. Mouthparts relatively short, cibarium bare. Pigmentation on third and fourth abdominal tegites not markedly wider than long; sclerotised plate surrounding gonopore truncate; two subequal ovoid or round developed spermathecae with short, tapered necks and one vestigial present, sclerotized ring parallel-sided. Male. Antennae with plume of SCh on flagellomeres 1–10, flagellomeres 2–10 partially fused, 11–13 elongate and subcylindrical; SCo distribution 1, 11– 13; pair of STl on flagellomeres 2–4, single on 5–7; single STc on 2–8; basal whorls of three and two SCh on 11 and 12 respectively. Wings patterned similarly to female but with pale spots generally more expansive. Genitalia with ninth tergite as wide anteriorly as posteriorly with lateral sides concave, apicolateral processes reduced to broad flanges, posterior margin entire and concave. Gonocoxite almost parallel sided for most of its length, ventral and dorsal roots long and narrow, both curving to converge apically; dorsal root arising from gonocoxite at about 75–90 °• Gonostylus slender, distally curving gently to a pointed apex. Ninth sternite with deep caudomedial excavation, membrane bare or spiculate. Aedeagus with low arch; body with almost straight lateral shoulders leading to a tapered distal process with a smoothly rounded apex; anteriorly projected peg not connected to arms of aedeagus by membrane and with an asymmetrically shaped apex. Parameres separate, converging and swelling medially, apical arms tapered, curved and converging to sharp tips. Immatures: Pupa relatively smooth, respiratory horns stout without spines or transverse convolutions, lateral respiratory openings not borne on protuberances; Operculum with long hairlike spines, rarely with reticulate bulges; ad setae long, subequal; d tubercles 1–3 in line and 1 and 2 quite close together; caudal segment with a transverse band of spines across disc; caudal spines short, without spicules. Larva with small eyes made up of a single pigmented spot on each side; head very short and square, broad in front; collar weakly developed. Ventral side of head with longitudinal suture, head setae moderately long; parahypostomal setae located at a distance from subgenal rim; labrum pointed, with two overlying folds; epipharynx with 2–3 combs, dorsal comb with 7–10 teeth. Thorax unpigmented but with diffuse paired post lateral patches of fat body in fourth instar. Larval habitat: Animal dung-pats, moist turf high in organic matter, rotting organic matter. Larvae and pupae are unable to swim and sink in tap water. World distribution. Restricted to the Old world - Afrotropical, Oriental, Palaearctic and Australasian regions.

International audience; The monophyly of the Imicola complex, a natural species complex within subgenus C. subgen. Avaritia Fox of the biting midge genus Culicoides Latreille, is supported using morphological and molecular analyses. A diagnosis for the group along with comparative redescriptions of the male and female of the species represented in Australasia, C. brevitarsis Kieffer and C. nudipalpis Delfinado and a description of C. asiatica Bellis sp. nov., are presented together with keys for their specific determination and molecular support for their status.

Culicoides brevitarsis Kieffer (Figs 2, 6, 7, 10, 11, 13, 16, 19, 22, 24, 26, 29, 32, 37, 38, 39, 44) Culicoides brevitarsis Kieffer 1917: 187; Lee & Reye 1962: 362 (C. robertsi synonymy); Kettle & Elson 1976: 313 (description of larva and pupa); Dyce 1979: 52 (C. radicitus synonymy); Dyce & Wirth 1983: 224 (C. superfulvus synonymy); Wirth & Hubert 1989: 258 (description of male and female, SE Asian records; designation of neotype); Lien et al., 1998: 27 (misident. Taiwan record, description of male and female). Yu et al. 2005: 911 (redescription of male and female, China record); Dyce et al., 2007: 19 (female wing illustrated). Culicoides robertsi Lee & Reye 1953: 386; Tokunaga, 1960: 74 (C. orientalis male of Tokunaga 1959 misdet.); Tokunaga, 1962: 485 (male and female redescribed; New Guinea and Solomon Islands); Culicoides radicitus Delfinado 1961: 657 (in part). Culicoides superfulvus Das Gupta 1962: 253 Culicoides orientalis Macfie (misident.); Tokunaga, 1959: 254 (Indonesian Papua record). Type material examined. C. radicitus: Philippines Tala, Rizal, 21.v. 1958, M Delfinado, (neotype male for C. brevitarsis, designation by Wirth & Hubert 1989, USNM). C. superfulvus: India Baguiati (near Calcutta) by sticky trap, August, ��� 60, Das Gupta (cotype female, designation by Dyce & Wirth 1983, USNM). C. brevitarsis: Australia, NSW. (neoallotype female, Lismore, 13.Nov. 2007, H. McKenzie, ANIC). Non-type material examined. Australia. NSW: Paterson, 6.Feb. 2008, Lt Tp. (1 female, NTQIC); NSW, Grafton, 14.Dec. 2007, H. McKenzie (1 female, NTQIC); Qld: Oxley Ck, Corinda, 18.March. 2009, M. Shivas, (1 female, NTQIC); Bamaga, 27.Jan. 2009, J Sailor, (1 female, NTQIC). Timor Leste, Surucraic, 9.05583 ��S; 125.5444 ��E, 15.Nov. 2001, Lt Tp, E. Watkins, (1 female, NTQIC); 4 km. N. Same, Lt. Tp. 24 th Aug 1969, D. Nicholls (3 females, ANIC); Cape Tefara, lt. tp. 5 th Aug 1969, DG. Nicholls (1 female, ANIC); Cape Tefara, 11km. S.E. Suai, Lt. Tp. 6 th August 1969, D. Nicholls (2 females, ANIC); Cape Tefara, lt. tp. 5 th Aug 1969, DG. Nicholls (1 female, ANIC); Los Palos, lt. tp. 6���7.ix. 69, DG. Nicholls (1 female, ANIC); Desa Pollo, 13 Sept 1984, Light trap, NT. Hunt (3 females, ANIC); Desa Pollo, 12���13 Sept 1984, NT. Hunt & Phillip, light trap by penned cattle (1 female, ANIC); Oecusse, Samora, 22.Feb. 2005, G. Bellis (1 female, NTQIC); Cova Lima, Suai, 9.31 ��S, 125.26 ��E, 3.Dec. 2003, E. Watkins (1 female, NTQIC);. Solomon Islands, Malaita, Kwailatutu, 8 �� 56 ��� 24 ������S, 160 �� 46 ��� 43 ���E, 18.May. 2010, L. Halling (1 female, NTQIC); Western Province, Tuiai via Nila,. 7.09491 ��S, 155.86068 ��E, light trap on edge of mangroves with 1 penned pig and 3 cattle, 0m, 19.Feb. 2012, G. Bellis (3 females, NTQIC). Papua New Guinea, West New Britain, Numondo, 5 ��31.0'S, 150 ��5.0'E, 28.Feb. 2007 LT, P. Boland, (1 female, NTQIC); Markham Farm, 6 �� 32.684 'S; 146 �� 38.724 'E, 11���17.June. 2008, Lt Tp, J. Schmidt & N. Harris, (1 male, NTQIC). Indonesia, Bali, Denpasar, 9.Feb. 1989, L.T. over cattle, HA. Standfast (2 females, 1 male, ANIC); Pedang bay, Denpasar, 35km NE, 17.Oct. 1969, DG. Nicholls (1 female, ANIC). Kalimantan, 80 -0818- 29, VLi- 95, V. Lee (1 female, ANIC). Flores, 80 -0812- 5, VLi- 30, V.Lee (2 females, ANIC). Sumbawa, 22.Oct. 1969, DG Nicholls (3 females, ANIC). West Timor, Kupang, Quarantine Station, L.T. over cattle, 7.Feb. 1989, HA. Standfast (3 females, ANIC). Papua, Merauke, L.T. 15.Feb. 89, HA Standfast, ov. cattle (7 females, ANIC). Thailand, Chiengmai, Apr- May 1958, V. Notananda, light trap (1 female, 1 male, ANIC). China, Hainan, Mengnya, 20.08 ��S, 110.37 ��E, Lt Tp, 28.Nov. 2010, G. Bellis (2 males, NTQIC); Sanlian Can, 20.08 ��S, 110.37 ��E, Lt Tp, 28.Nov. 2010, G. Bellis (1 male, NTQIC). Diagnosis. We have been unable to reliably distinguish the morphology of C. brevitarsis from C. bolitinos Meiswinkel so the following diagnosis applies to both of these species and separates these two species from the remaining species in the Imicola complex. Female: The only species in the Imicola complex with the combination of wing with posterior margin of apical pale marking in cell m 1 straight and never attaining vein M 2 before margin of the wing, proximal dark marking on costa not longer than stigmatic dark spot and apical fifth of cell r 2 included in post-stigmatic pale spot. Male: The only species in the Imicola complex with the combination of wing with posterior margin of apical pale marking in cell m 1 straight and never attaining vein M 2 before margin and proximal dark marking on costa no more than 1.5 times as long as the stigmatic dark spot. Description. Adult. In addition to characters listed in the diagnosis, palpus (Fig. 13) with 3 rd segment bearing shallow pit of diameter about �� width of segment and containing few, elongate, capitate sensilla; wing (Fig. 6, 7) relatively strongly patterned with about 1 / 5 of cell r 2 included in the post-stigmatic pale marking; legs (Fig. 19) pale brown, hind femora unbanded, mid and fore femora with subapical pale band, all tibiae with pale subbasal band, mid & hind tibia additionally with weak pale apical bands; haltere pale. Male hypopygium (Fig. 37) with ninth sternite with ventral membrane bare. Aedeagus (Fig. 39) with peg strongly sclerotised and quite dark. Immatures. Fourth instar larvae and pupae were adequately described by Kettle & Elson (1976). Distribution. (Fig. 44) Widespread in the Oriental and Australasian regions. Australia: NSW, Qld, NT, WA; PNG, Solomon Is, Fiji, New Caledonia, Vanuatu, Tonga, Timor Leste, Indonesia, Philippines, Thailand, China and India. Records from within the Oriental Region require confirmation as they may refer to C. asiatica. Biology. Closely associated with bovids. Immatures live in discrete bovid dung pats (Canon & Reye 1966, Campbell & Kettle 1976) while adults feed on sheep, cattle, marsupial, buffalo, human and horse (Lee et al. 1962; Kay 1973; Muller & Murray 1977; Muller et al. 1981; Kay & Lennon 1982). Adults are crepuscular (Campbell & Kettle 1979 b; Bellis et al. 2004), roost in ground herbage during daylight hours (Bishop et al. 1995) and mate in swarms at sunset (Campbell & Kettle 1979 a). This species has been implicated in the transmission of 19 different viruses, including several of economic importance (Doherty et al. 1972, St George et al. 1977, 1978, 1979, 1983; Muller et al. 1982; Zakrewski & Cybinski 1984; Parsonson & Snowdon 1985; Standfast et al. 1985; Cybinski & Muller 1990). Remarks. Lee & Reye (1962) suggested the holotype specimen of C. brevitarsis may have been destroyed in the fire at the National Museum of Hungary in Budapest in 1956. Debenham (1979) was unable to locate the holotype specimen of this species confirming that it was lost in 1956 and prompting Wirth & Hubert (1989) to designate a neotype using the neotype male of C. radicitus. The designation of an allotype female from the type locality in Australia is however, desirable due both to the marked sexual dimorphism in wing pattern of this species and the confusion surrounding the identity of this species in Asia. Kieffer (1917) gave only ��� Australie ��� as the type locality but morphological and genetic analyses of C. brevitarsis from various localities in Australia (this study, Gopurenko et al. in preparation) has revealed there to be only a single species present indicating that any locality within Australia is suitable for an allotype specimen. Within the Imicola group, C. brevitarsis is morphologically most similar to the African C. bolitinos and the Asian C. asiatica and this grouping is strongly supported by the molecular data. The relative lengths of the first and second dark marking along the costa enables reliable separation of C. brevitarsis and C. bolitinos from C. asiatica, but does not enable separation between C. brevitarsis and C. bolitinos. Meiswinkel (1995: 31) offered the length:width ratio of flagellomeres 4���7 and the absolute length of all flagellomeres of the female antenna as a means of distinguishing C. brevitarsis from C. bolitinos. Data for these characters for C. asiatica, C. brevitarsis and C. bolitinos (Table 6) illustrate that although differences are apparent between the mean values of these characters, the overlap evident between species makes identification of individual specimens unreliable. These three taxa are however, reliably separated using either the COI and CAD genes (Fig. 46���48). Despite the closer morphological similarity of C. brevitarsis to C. bolitinos than to C. asiatica, both CAD and COI analyses indicate that C. asiatica is the sister species of C. brevitarsis (Fig. 46���48). Although COI analysis suggests that the Chinese specimens of C. brevitarsis may represent a further cryptic species (Fig. 47), this was not supported by the CAD analysis, which nested these specimens among other genotypes of C. brevitarsis (Fig. 48) suggesting that the COI division between Chinese and Australasian specimens reflects maternal phylogeographic structuring within the species rather than indicating presence of separate cryptic species., Published as part of Bellis, Glenn, Dyce, Alan, Gopurenko, David, Yanase, Tohru, Garros, Claire, Labuschagne, Karien & Mitchell, Andrew, 2014, Revision of the Culicoides (Avaritia) Imicola complex Khamala & Kettle (Diptera: Ceratopogonidae) from the Australasian region, pp. 401-427 in Zootaxa 3768 (4) on pages 405-407, DOI: 10.11646/zootaxa.3768.4.1, http://zenodo.org/record/218566, {"references":["Kieffer, J. J. (1917) Chironomides d'Australie conserves au Musee National Hongrois de Budapest. Annales Historico- Naturales Musei Nationalis Hungarici, 15, 75 - 228.","Kettle, D. S. & Elson, M. M. (1976) The immature stages of some Australian Culicoides Latreille (Diptera: Ceratopogonidae). Journal of the Australian Entomological Society, 15, 303 - 332. http: // dx. doi. org / 10.1111 / j. 1440 - 6055.1976. tb 01711. x","Dyce, A. L (1979) Culicoides radicitus Delfinado: A synonym of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae). Journal of the Australian Entomological Society, 18, 52. http: // dx. doi. org / 10.1111 / j. 1440 - 6055.1979. tb 00811. x","Dyce, A. L. & Wirth, W. W. (1983) Reappraisal of some Indian C ulicoides species in the subgenus A varitia (Diptera: Ceratopogonidae). International Journal of Entomology 25 (2 - 3), 221 - 225.","Wirth, W. W. & Hubert, A. A. (1989) Culicoides of Southeast Asia. Memoirs of the American Entomological Institute No. 44, Florida, USA, 508 pp.","Lien, J. - C., Lin, C. - C. & Weng, M. - H. (1998) A revision of the genus Culicoides in Taiwan. Part II. Subgenus Avaritia (Diptera, Ceratopogonidae). Journal of Taiwan Museum, 51, 21 - 48.","Yu, Y. - X., Liu, J. - H., Liu, G. - P., Liu, Z. - J., Hao, B. - S., Yan, G. & Zhao, T. - S. (2005) Ceratopogonidae of China, Insecta, Diptera, Vols. 1 - 2. Military Medical Science Press, Beijing, 1699 pp. [in Chinese]","Dyce, A. L, Bellis, G. A. & Muller, M. J. (2007) Pictorial atlas of Australasian Culicoides wings (Diptera: Ceratopogonidae). Australian Biological Resources Study, Canberra, 88 pp.","Lee, D. J. & Reye, E. J. (1953) Australasian Ceratopogonidae (Diptera, Nematocera). Part VI. Australian species of Culicoides. Proceedings of the Linnean Society of New South Wales, 77, 369 - 394.","Tokunaga, M. (1960) Notes on biting midges I. Akitu, 9, 72 - 76.","Tokunaga, M. (1959) New Guinea biting midges (Diptera: Ceratopogonidae). Pacific Insects, 1, 177 - 314.","Tokunaga, M. (1962) Biting midges of the genus Culicoides from New Guinea (Diptera: Ceratopogonidae). Pacific Insects, 4, 457 - 516.","Delfinado, M. D. (1961) Philippine Zoological Expedition 1946 - 1947. The Philippine biting midges of the genus Culicoides (Diptera: Ceratopogonidae). Fieldiana Zoology, 33, 627 - 675.","Das Gupta, S. K. (1962) Culicoides (Dipt., Ceratopogonidae) from suburbs of Calcutta. Entomologist's Monthly Magazine, 98, 253 - 254.","Cannon, L. R. G. & Reye, E. J. (1966) A larval habitat of the biting midge Culicoides brevitarsis Keiffer (Diptera: Ceratopogonidae). Journal of the Entomological Society of Queensland, 5, 7 - 9. http: // dx. doi. org / 10.1111 / j. 1440 - 6055.1966. tb 00670. x","Campbell, M. M. & Kettle, D. S. (1976) Number of adult Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) emerging from bovine dung exposed under different conditions in the field. Australian Journal of Zoology, 24, 75 - 85. http: // dx. doi. org / 10.1071 / zo 9760075","Kay, B. H. (1973) Seasonal studies on a population of Culicoides marmoratus (Skuse) (Diptera: Ceratopogonidae) at Deception Bay, Queensland. Journal of the Australian Entomological Society, 12, 42 - 58. http: // dx. doi. org / 10.1111 / j. 1440 - 6055.1973. tb 02153. x","Muller, M. J. & Murray, M. D. (1977) Blood-sucking flies feeding on sheep in south-eastern Australia. Australian Journal of Zoology, 25, 75 - 85. http: // dx. doi. org / 10.1071 / zo 9770075","Muller, M. J., Murray, M. D. & Edwards, J. A. (1981) Blood-sucking midges and mosquitoes feeding on mammals at Beatrice Hill. N. T. Australian Journal of Zoology, 29, 573 - 588. http: // dx. doi. org / 10.1071 / zo 9810573","Campbell, M. M. & Kettle, D. S. (1979 b) Abundance and temporal and spatial distribution of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) on cattle in South-East Queensland. Australian Journal of Zoology, 27, 251 - 260. http: // dx. doi. org / 10.1071 / zo 9790251","Bellis, G. A., Melville, L. F., Hunt, N. T. & Hearnden, M. N. (2004) Temporal activity of biting midges (Diptera: Ceratopogonidae) on cattle near Darwin, Northern Territory Australia. Veterinaria Italiana, 40 (3), 324 - 328.","Bishop, A. L., McKenzie, H. J., Barchia, I. M. & Harris, A. M. (1995) The effects of habitat on the distribution of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) during its resting phase. Australian Journal of Zoology, 43, 531 - 539. http: // dx. doi. org / 10.1071 / zo 9950531","Campbell, M. M. & Kettle, D. S. (1979 a) Swarming of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) with reference to markers, swarm size, proximity of cattle, and weather. Australian Journal of Zoology, 27, 17 - 30. http: // dx. doi. org / 10.1071 / zo 9790017","Doherty, R. L., Carley, J. G., Standfast, H. A., Dyce, A. L. & Snowdon, W. A. (1972) Virus strains isolated from arthropods during an epizootic of bovine ephemeral fever in Queensland. Australian Veterinary Journal, 48, 81 - 86. http: // dx. doi. org / 10.1111 / j. 1751 - 0813.1972. tb 02220. x","St. George, T. D., Standfast, H. A., Christie, D. G., Knott, S. G. & Morgan, I. R. (1977) The epizootiology of bovine ephemeral fever in Australia and Papua-New Guinea. Australian Veterinary Journal, 53, 17 - 28. http: // dx. doi. org / 10.1111 / j. 1751 - 0813.1977. tb 15812. x","St. George, T. D., Standfast, H. A. & Cybinski, D. H. (1978) Isolations of Akabane virus from sentinel cattle and Culicoides brevitarsis. Australian Veterinary Journal, 54, 558 - 561. http: // dx. doi. org / 10.1111 / j. 1751 - 0813.1978. tb 02412. x","St. George, T. D., Cybinski, D. H., Filippich, C., & Carley, J. G. (1979) The isolation of three Simbu group viruses new to Australia. Australian Journal of Experimental Biology and Medical Science, 57, 581 - 582. http: // dx. doi. org / 10.1038 / icb. 1979.60","St. George, T. D., Cybinski, D. H., Standfast, H. A., Gard, G. P. & Della-Porta, A. J. (1983) The isolation of five different viruses of the epizootic haemorrhagic disease of deer serogroup. Australian Veterinary Journal, 60, 216 - 217. http: // dx. doi. org / 10.1111 / j. 1751 - 0813.1983. tb 09587. x","Muller, M. J., Standfast, H. A., St. George, T. D. & Cybinski, D. H. (1982) Culicoides brevitarsis (Diptera: Ceratopogonidae) as a vector of arboviruses in Australia. Arbovirus Research in Australia, 3, 43 - 49.","Zakrewski, H. & Cybinski, D. H. (1984) Isolation of Kimberley virus, a rhabdovirus, from Culicoides brevitarsis. Australian Journal of Experimental Biology and Medical Science, 62, 779 - 780.","Parsonson, I. M. & Snowdon, W. A. (1985) Bluetongue, epizootic haemorrhagic disease of deer and related viruses: Current situation in Australia. In: Barber, T. L. & Jochim, M. M., (Eds.), Bluetongue and Related Orbiviruses. A. R. Liss, New York, pp. 27 - 36.","Standfast, H. A., Dyce, A. L. & Muller, M. J. (1985) Vectors of bluetongue virus in Australia. Progress in Clinical Biological Research, 178, 177 - 186.","Cybinski, D. H. & Muller, M. J. (1990) Isolation of arboviruses from cattle and insects at two sentinel sites in Queensland, Australia, 1979 - 1985. Australian Journal of Zoology, 38, 25 - 32. http: // dx. doi. org / 10.1071 / zo 9900025","Debenham, M. L. (1979) An annotated checklist and bibliography of Australasian Region Ceratopogonidae (Diptera, Nematocera). School of Public Health and Tropical Medicine, University of Sydney and Commonwealth Department of Health Monograph Series. Entomology Monograph, 1, xiv + 671 pp.","Meiswinkel, R. (1995) Afrotropical Culicoides: biosystematics of the Imicola group, subgenus Avaritia (Diptera: Ceratopogonidae), with special reference to the epidemiology of African horse sickness. MSc (Agric.) dissertation. University of Pretoria, South Africa, iv + 359 pp."]}

Key to females of Australasian species of the Imicola complex 1. Apical half of cell r 2 included in post-stigmatic, apical pale marking in cell m 1 reaching or crossing vein M 2 subapically then narrowing apically and not touching vein M 2 at wing margin (Fig. 8)..................................... nudipalpis - apex of cell r 2 with apical 1 / 5 to �� included in post stigmatic pale spot, apical pale marking in cell m 1 not narrowed apically and either touching or approaching vein M 2 at wing margin (Fig. 4, 6).......................................... 2 2. Wing with proximal dark marking on costa of similar length to stigmatic dark spot, apex of cell r 2 with apical 1 / 5 included in post stigmatic pale spot (Fig. 6).................................................................. brevitarsis - Wing with proximal dark marking on costa distinctly longer than stigmatic dark spot, apex of cell r 2 with apical 1 / 5 to �� included in post stigmatic pale spot (Fig. 4).......................................................... asiatica, Published as part of Bellis, Glenn, Dyce, Alan, Gopurenko, David, Yanase, Tohru, Garros, Claire, Labuschagne, Karien & Mitchell, Andrew, 2014, Revision of the Culicoides (Avaritia) Imicola complex Khamala & Kettle (Diptera: Ceratopogonidae) from the Australasian region, pp. 401-427 in Zootaxa 3768 (4) on page 405, DOI: 10.11646/zootaxa.3768.4.1, http://zenodo.org/record/218566

Culicoides nudipalpis Delfinado (Figs 3, 8, 9, 14, 17, 20, 23, 27, 30, 33, 40, 41, 42, 45) Culicoides nudipalpis Delfinado 1961: 655; Wirth & Hubert 1989: 281 (description of male and female); Meiswinkel & Baylis, 1998 (separation from C. imicola); Lien et al., 1998: 35 (Taiwan record, description of male and female); Yu et al. 2005: 1018 (China record, description of male and female); Dyce et al., 2007: 19 (female wing illustrated). Type material examined. Philippines: Tala, Rizal, 21.May. 1958, M. Delfinado (paratype, female, USNM) Non-type material examined. Timor Leste: Hera, 30.Mar. 2004, Lt Tp., E. Watkins, (2 females, NTQIC); Cape Tefara, lt. tp. 5.Aug. 1969, DG. Nicholls (5 females, 1 male, ANIC); Suai, Cafe Cooperativa Timor, 9 �� 23.046 'S; 125 �� 10.675 'E, Lt Tp, 13���14.May. 2010, B. Cookson (1 female, 2 males NTQIC); Ainaro, Surucraic, 15.Nov. 2001, E. Watkins (1 female, NTQIC), Bobonaro, Maliana, Rai Fun, 7 -May- 2002, E. Watkins (1 female, NTQIC); Oecusse, Samora, 24 -Feb- 2005, G. Bellis (1 female, NTQIC); Cova Lima, Suai, 9.31 ��S, 125.26 ��E, 5.May. 2002, G. Bellis (1 female, NTQIC); Oecusse, Pante Makassar, 7 -Dec- 2002, G. Bellis (1 female, NTQIC); Ermera, Letefoho Eraulo, 22.Jul. 2004, E. Watkins (1 female, NTQIC); Viqueque, 19.Jul. 2000, A. Moss (1 female, NTQIC); Dili, Cristo Rei, Hera, 19.Jul. 2000, A. Moss (1 female, NTQIC). Indonesia, Irian Jaya, Merauke, Kumbe, VLi378, 5.Aug. 1980 (1 male, ANIC). Diagnosis. Female: The only species in the Imicola complex with the combination of wing with proximal dark marking on costa of similar length to stigmatic dark spot, apical half of cell r 2 included in post-stigmatic pale spot, posterior margin of apical pale marking in cell m 1 curving posteriorly to reach vein M 2 well before the wing margin, often crossing vein M 2 and joining the pale spot in cell m 2 and with PH ratio less than 0.75. Male: there are currently no reliable characters for separating males of this species from those of C. imicola. These two species are the only species in the Imicola complex with the posterior margin of apical pale marking in cell m 1 curving posteriorly to reach vein M 2 well before the wing margin, often crossing vein M 2 and joining the pale spot in cell m 2 Description. Adult. In addition to characters listed in the diagnosis, palpus (Fig. 14) pale brown with 3 rd segment slightly swollen, broadest about mid way along, sensory area a superficial rounded patch of aggregated sensoria in a shallow depression spanning nearly the full width of segment; legs (Fig. 20) pale brown, fore & mid femora with pale apical band, all tibiae with pale basal band and weak, broad apical pale band; haltere pale. Male hypopygium (Fig. 40) with ventral membrane of ninth sternite bearing a sparse covering of spicules. Aedeagus (Fig. 42) with peg broad at base and noticeably paler than distal process. Immatures. Unknown. Distribution. (Fig. 45) Philippines (Delfinado 1961; Wirth & Hubert 1989), Indonesia: Bali, West Timor, Papua, Flores, Sumbawa, Lombok and Maluku (Wirth & Hubert 1989; Sukarsih et al. 1993; Dyce et al. 2007), Timor Leste (Wirth & Hubert 1989), Taiwan (Lien et al. 1998) and China, Hainan (Yu et al. 2005). Biology. Largely unknown. Bellis et al. (2005) investigated some of the aspects of biology pertinent to vector capacity and found this species to feed on mammals and have a relatively high survival rate. Sukarish et al. (1993) reported collecting this species in association with cattle. Remarks. Dyce (1979) reported some confusion in the type series of this species. He could not locate the holotype female and the specimen illustrated as C. nudipalpis by Delfinado (1961) was apparently labeled as an allotype of C. radicitus. Wirth & Hubert (1989) designated a neotype which is now lodged in the USNM. Meiswinkel & Baylis (1998) correctly noted that C. nudipalpis is morphologically most similar to C. imicola and reported on characters used to distinguish these two species. They were unable to observe differences between males of these two species and although differences are apparent in the ratio of the length between the interocular seta and the base of the cibarium to the length between the base of the cibarium and the apex of the labium (Bellis, Garros & Labuschagne unpublished data), these differences require further study to confirm their validity. Results from the present study confirm the observations of Meiswinkel & Baylis (1998) that the PH ratio of female C. nudipalpis is less than that reported for C. imicola and this remains the most reliable means of separating these species. Genetic analyses reported herein support the validity of C. nudipalpis as a taxon and analyses of CAD and concatenated CAD + COI data place it as sister to C. imicola as morphological similarity might suggest although COI analysis did not agree with this close relationship. Wirth & Hubert (1989) reported that these two species do not occur sympatrically although Yu et al. (2005) have subsequently reported both species occurring on Hainan in China and Lien et al. (1998) reported C. nudipalpis in nearby Taiwan so these species are not as geographically separate as previously thought and workers in this region are encouraged to check material to ensure accurate identification. Antennal flagellomere Antennal flagellomere Gene: COI codon 1 st 2 nd 3 rd all Notes: # informative = number of parsimony informative sites; CI = consistency index; CI-ex = consistency index excluding invariant sites; RI = retention index; RC = rescaled consistency index (CI * RI)., Published as part of Bellis, Glenn, Dyce, Alan, Gopurenko, David, Yanase, Tohru, Garros, Claire, Labuschagne, Karien & Mitchell, Andrew, 2014, Revision of the Culicoides (Avaritia) Imicola complex Khamala & Kettle (Diptera: Ceratopogonidae) from the Australasian region, pp. 401-427 in Zootaxa 3768 (4) on pages 413-422, DOI: 10.11646/zootaxa.3768.4.1, http://zenodo.org/record/218566, {"references":["Delfinado, M. D. (1961) Philippine Zoological Expedition 1946 - 1947. The Philippine biting midges of the genus Culicoides (Diptera: Ceratopogonidae). Fieldiana Zoology, 33, 627 - 675.","Wirth, W. W. & Hubert, A. A. (1989) Culicoides of Southeast Asia. Memoirs of the American Entomological Institute No. 44, Florida, USA, 508 pp.","Meiswinkel, R. & Baylis, M. (1998) Morphological confirmation of the separate species status of Culicoides (Avaritia) nudipalpis Delfinado, 1961 and C. (A.) imicola Kieffer, 1913 (Diptera: Ceratopogonidae). Onderstepoort Journal of Veterinary Research, 65, 9 - 16.","Lien, J. - C., Lin, C. - C. & Weng, M. - H. (1998) A revision of the genus Culicoides in Taiwan. Part II. Subgenus Avaritia (Diptera, Ceratopogonidae). Journal of Taiwan Museum, 51, 21 - 48.","Yu, Y. - X., Liu, J. - H., Liu, G. - P., Liu, Z. - J., Hao, B. - S., Yan, G. & Zhao, T. - S. (2005) Ceratopogonidae of China, Insecta, Diptera, Vols. 1 - 2. Military Medical Science Press, Beijing, 1699 pp. [in Chinese]","Dyce, A. L, Bellis, G. A. & Muller, M. J. (2007) Pictorial atlas of Australasian Culicoides wings (Diptera: Ceratopogonidae). Australian Biological Resources Study, Canberra, 88 pp.","Bellis, G. A., Melville, L. & Walsh, S. (2005) Preliminary assessment of the vector capacity of Culicoides nudipalpis and C. orientalis for bluetongue virus. Arbovirus Research in Australia, 9, 28 - 32.","Dyce, A. L (1979) Culicoides radicitus Delfinado: A synonym of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae). Journal of the Australian Entomological Society, 18, 52. http: // dx. doi. org / 10.1111 / j. 1440 - 6055.1979. tb 00811. x"]}

Culicoides asiatica Bellis sp. nov. (Figs 1, 4, 5, 12, 15, 18, 21, 25, 28, 31, 34 ���36, 43) Culicoides brevitarsis Kieffer (misident.) Kitaoka 1984 (in key); Henna et al. 1991, Wada et al. 1996 (Japan records); Lien et al. 1998 (Taiwan record); Matsumoto et al. 2009 (COI and COII sequence); Yanase et al. 2011 (Kyushu record), Yanase et al. 2013 (larval COI). Type material examined. Holotype: Japan: Kagoshima, Minamikyushu, 31 �� 19 ���N; 130 �� 23 ���E, 11.Sept. 2009, T. Yanase, (female, NMNS), Paratypes: same data as holotype (1 female NTQIC); Okinawa, Yonaguni, Feb. 2005, T. Yanase, (3 females, NTQIC), Okinawa, Nakijin, 26 �� 41 'N; 127 �� 56 'E, 9.Dec. 2010, T. Yanase (2 males, NTQIC, 1 male NMNS, 2 males KRSC), Kagoshima, Kanoya, 31 �� 27 'N; 130 �� 49 'E, 16.Oct. 2009, T. Yanase, (1 female NMNS, 2 females NTQIC, 2 females KRSC); Okinawa, Yonaguni Is, 16.Nov. 1974 Lt.tp. S. Kitaoka (4 females, ANIC); Timor Leste, Surucraic, 9.05583 ��S; 125.5444 ��E, 15.Nov. 2001, Lt Tp, E. Watkins, (1 female, NTQIC); 4km.N. Same, Lt Tp 24.Aug. 1969, D. Nicholls (1 female ANIC). Indonesia: Lombok, Tabane, East Lombok, 20.Oct. 1969, DG. Nicholls (1 female, ANIC), West Timor, Kupang, Quarantine Station, Ov. Cattle L.T. 7.Feb. 1989, HA. Standfast (1 female, ANIC). Malaysia: Ipoh, Light Trap, 31.May. 1978, CS. Shanta, (8 females, ANIC), Ipoh, 7.Dec. 1977, S. Kitaoka (2 males, ANIC); Selangor, Kuala Lumpur, Aug 1958, R. Traub, light (2 females, ANIC), Selangor, Kuala Lumpur, March 1958, R. Traub, light (1 female, ANIC); Laos: Vientiane, Hatsayfong, Lt Tp. Aug. 2010, P. Soysouvanh (1 female, NTQIC). Thailand: Phangnga, Pulau Panjang, 17.Nov. 1954, cowshed - night, DH. Colless (1 female, ANIC); TaLee Dist. 8��� 9.June. 1959, manop, col. light (1 male, ANIC). Diagnosis. Female: The only species in the Imicola complex with the combination of wing with apical pale marking in cell m 1 not narrowed apically and either touching or approaching vein M 2 at wing margin, proximal dark marking on costa distinctly longer than stigmatic dark spot and apical third to half of cell r 2 included in poststigmatic pale spot. Male: The only species in the Imicola complex with the combination of wing with apical pale marking in cell m 1 not narrowed apically and touching vein M 2 at wing margin and proximal dark marking on costa distinctly greater than twice as long as the stigmatic dark spot. Description. Adults: In addition to characters listed in the diagnosis, eyes bare in both sexes; palpus (Fig. 12) pale brown with 3 rd segment slightly swollen medially with a round shallow pit with a diameter of about half the width of 3 rd segment bearing emergent, capitate sensilla; legs (Fig. 18) pale brown, fore & mid femora with weak apical pale bands, hind femora dark to apex, all tibiae with pale basal bands and unbanded apex; haltere pale. Male hypopygium (Fig. 34) with ventral membrane of ninth sternite bare. Immatures. Unknown. Distribution. (Fig. 43) Japan, Taiwan, Laos, Thailand, Malaysia, Indonesia (Lombok, West Timor) and Timor Leste. Biology. Takayoshi et al. (1994), Goto et al. (2004) and Yanase et al. (2010) detected bluetongue virus from field-collected females of this species (reported as C. brevitarsis) in Japan. Yanase et al. (2011) reported collecting large numbers of this species (as C. brevitarsis) in light traps set at cowsheds near pasture. Label data of specimens examined herein and molecular analysis of larvae (Yanase et al. 2013) indicates that C. asiatica breeds in discrete cattle dung pats. Remarks. The presence of a species morphologically similar to C. brevitarsis in Asia prompted the need to reassess the status of the junior synonyms of C. brevitarsis in Asia, namely C. radicitus and C. superfulvus. Unfortunately, the holotype specimens of both of these species have been lost (Dyce 1979, Dyce & Wirth 1983) but the neotype male of C. radicitus designated by Wirth & Hubert (1989) and the cotype female of C. superfulvus designated by Dyce & Wirth (1983) were kindly loaned by the USNM. Based on the relative lengths of the first two dark markings on the costa of these two specimens (Fig. 10 and 11), they are both referable to the current definition of C. brevitarsis and thus remain as junior synonyms of this species. The extensive geographical sympatry between C. brevitarsis and C. asiatica does not allow reliable interpretation of many previously published studies on C. brevitarsis senso lato in Asia. The record of Lien et al. (1998) of C. brevitarsis from Taiwan is supported by a wing photograph which clearly shows that the first costal dark spot is longer than the stigmatic dark spot and this specimen is consequently referable to C. asiatica. Similarly, illustration of the wing of C. brevitarsis from China provided by Yu et al. (2005) is indeed referable to that species although it is not clear if the specimen illustrated is from An Hui, Hainan or Taiwan as all three localities are listed by Yu et al. (2005). Specimens examined herein indicate that C. brevitarsis senso stricto is present in Hainan but the presence of this species in Taiwan and An Hui requires confirmation. All specimens examined herein from Japan, including representatives from the populations studied by Matsumoto et al. (2009) and Yanase et al. (2010; 2011; 2013), are referable to C. asiatica so it is reasonable to assume that all reports from Japan previously attributed to C. brevitarsis should be referred to C. asiatica. Workers in this region however need to be vigilant for the presence of C. brevitarsis in future studies. All specimens examined from Australia, PNG and the Pacific are referable to C. brevitarsis and any work reported from this region remain referable to that species. The presence of C. asiatica in nearby Indonesia and Timor Leste however, warns that care should be taken when identifying specimens from this region., Published as part of Bellis, Glenn, Dyce, Alan, Gopurenko, David, Yanase, Tohru, Garros, Claire, Labuschagne, Karien & Mitchell, Andrew, 2014, Revision of the Culicoides (Avaritia) Imicola complex Khamala & Kettle (Diptera: Ceratopogonidae) from the Australasian region, pp. 401-427 in Zootaxa 3768 (4) on pages 407-413, DOI: 10.11646/zootaxa.3768.4.1, http://zenodo.org/record/218566, {"references":["Kitaoka, S. (1984) Japanese Culicoides (Diptera: Ceratopogonidae) and keys for the species. Bulletin of the National Insitue of Animal Health, 87, 73 - 108. [in Japanese]","Henna, M., Shinjo, K. & Kokuba, T. (1991) Identification of biting midges (Culicoides) and their distribution in Okinawa prefecture. Bulletin of the Okinawa Prefecture Institute of Animal Health, 27, 43 - 49. [in Japanese]","Wada, Y. Miyagi, L. Takagi, M. & Tsuda, Y. (1996) An annotated list of Culicoides biting midges in Yaeyama islands in the Southern Part of Japan (Diptera: Ceratopogonidae). Tropical Medicine, 38, 61 - 68.","Lien, J. - C., Lin, C. - C. & Weng, M. - H. (1998) A revision of the genus Culicoides in Taiwan. Part II. Subgenus Avaritia (Diptera, Ceratopogonidae). Journal of Taiwan Museum, 51, 21 - 48.","Matsumoto, Y., Yanase, T., Tsuda, T. & Noda, H. (2009) Species-specific mitochondrial gene rearrangements in biting midges and vector species identification. Medical and Veterinary Entomology, 23, 47 - 55. http: // dx. doi. org / 10.1111 / j. 1365 - 2915.2008.00789. x","Yanase, T., Hirata, M., Matsumori, Y, Matsumura, M., Kato, T., Shirafuji, H., Yamakawa, M., Hayama, Y. & Tsutsui, T. (2011) Detection of Culicoides brevitarsis Activity in Kyushu. Journal of Veterinary and Medical Science, 73 (12), 1649 - 1652. http: // dx. doi. org / 10.1292 / jvms. 11 - 0231","Yanase, T., Matsumoto, Y., Matsumori, Y., Aizawa, M., Hirata, M., Kato, T., Shirafuji, H., Yamakawa, M., Tsuda, T. & Noda, H. (2013) Molecular Identification of Field-Collected Culicoides Larvae in the Southern Part of Japan. Journal of Medical Entomology, 50 (5), 1105 - 1110. http: // dx. doi. org / 10.1603 / me 11235","Takayoshi, K., Kokuba, T., Kubo, M. & Miura, Y. (1994) Isolation of bluetongue virus from a calf and antibody survey among cattle in Okinawa. Journal of the Japanese Veterinary and Medical Association, 17, 651 - 654. [in Japanese]","Goto, Y., Yamaguchi, O. & Kubo, M. (2004) Epidemiological observation on bluetongue in sheep and cattle in Japan. Veterinaria Italiana, 40, 78 - 82.","Yanase, T., Kato, T., Katagiri, Y., Aizawa, M., Nakamura, K., Kokuba, T., Araki, M., Shirafuji, H., Yamakawa, M. & Tsuda, T. (2010) Isolation and characterization of Bluetongue virus from Culicoides brevitarsis (Diptera: Ceratopogonidae) in Okinawa. Medical Entomology and Zoology, 61 (1), 85 - 91. http: // dx. doi. org / 10.7601 / mez. 61.85","Dyce, A. L (1979) Culicoides radicitus Delfinado: A synonym of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae). Journal of the Australian Entomological Society, 18, 52. http: // dx. doi. org / 10.1111 / j. 1440 - 6055.1979. tb 00811. x","Dyce, A. L. & Wirth, W. W. (1983) Reappraisal of some Indian C ulicoides species in the subgenus A varitia (Diptera: Ceratopogonidae). International Journal of Entomology 25 (2 - 3), 221 - 225.","Wirth, W. W. & Hubert, A. A. (1989) Culicoides of Southeast Asia. Memoirs of the American Entomological Institute No. 44, Florida, USA, 508 pp.","Yu, Y. - X., Liu, J. - H., Liu, G. - P., Liu, Z. - J., Hao, B. - S., Yan, G. & Zhao, T. - S. (2005) Ceratopogonidae of China, Insecta, Diptera, Vols. 1 - 2. Military Medical Science Press, Beijing, 1699 pp. [in Chinese]"]}

Oenochroma barcodificata Hausmann & Young, sp. nov. Type material. Holotype: Ƥ, ‘TAS[MANIA], Chimney Pot Hill, nr Ridgeway, 20 OCT. 1996, eucalypt forest, leg. P.B. McQuillan’, coll. TMAG, genit.slide ‘ ZSM G 13863 ’, specimen ID ‘ BC ZSM Lep 0 2948 ’ [sequence ID GWORB 1068 -07]. Paratypes: coll. BIO: 63 Australia, Tasmania, Hobart, Kingston Beach, lat. - 42.986 x long. 147.317, alt. 110 m., 0 2., 11. and 18. XII.2005, 10. X.2006, 27. XII.2006, 22. I. 2007, leg. R. D. Ward, sequence IDs: LOTSA 134 -06/06-TASA-00134, LOTSA 125 -06/06-TASA-00125, LOTSA 122 -06/06-TASA- 0 0 122, LOTSC 391 -07/06-TASB-01390, LOTSC 814 -07/06-TASB-01813, LOTSD 507 -08/07-TASB-0507; coll. M. Sommerer: 1 Ƥ, Tasmania sept., vic. Mt. Roland, Silver Ridge Lodge, 41 ° 27 ’ S / 146 ° 15 ’ E, 320 m, 25-28.I. 2006, leg. M. Sommerer, BC ZSM Lep 0 8033. coll. ANIC (153, 16Ƥ): 13 ‘Mt Wellington 870m [Tas.] 6 Dec. 1980 L. Hill’; 13 ‘ 43.03 S 146.17 E Huon Camping Area Tas. 25 Jan 1983 J.C. Cardale’; 13 ‘Lake Fenton 1060m Tas. 14 Jan 1978 P.B. McQuillan’ ‘ ANIC Genitalia slide 18601 ’; 13 ‘Mt Read nr Rosebery Tas. 4–5 Dec 1990 P.B. McQuillan’; 13 ‘ 41.51 S 145.33 E Mt Read Tas. 500m 9 Jan 1989 P.B. McQuillan E.S. Nielsen’; 13 ‘ 43.25 S 146.09 E Melaleuca Tas. 15 Jan 1991 E.S. Nielsen E.D. Edwards’; 23 ‘ 43.25 S 146.09 E Melaleuca Tas. 28 Nov 1991 E.S. Nielsen M. Horak’; 13 ‘ 43.25 S 146.09 E Melaleuca Tas. 25 Nov 1991 E.S. Nielsen M. Horak’; 13 ‘ 43.25 S 146.09 E Melaleuca Tas. 14 Jan 1991 E.S. Nielsen E.D. Edwards’; 13 ‘ 43.25 S 146.09 E Melaleuca Tas. 17 Jan 1991 E.S. Nielsen E.D. Edwards’; 13 ‘ 43.23 S 146.08 E Claytons Bathurst Harb. Tas. 16 Jan 1991 E.S. Nielsen E.D. Edwards’; 1 Ƥ ‘Mt Wellington 1000m [Tas.] 8 Dec 1980 L. Hill’; 1 Ƥ “Mt Wellington 870m [Tas.] 6 Dec 1980 L. Hill’; 1 Ƥ “ 42.02 S 146.33 E 12km NNE Bronte Park Tas. 20 Jan 1983 J.C. Cardale’; 1 Ƥ ‘ 42.02 S 146.33 E 12km NNE Bronte Park Tas. 2 Feb 1983 J.C. Cardale’; 2 Ƥ “Pensford Tas. 920m [Tas.] 22 Dec 1981 L. Hill’; 1 Ƥ ‘ 40.58 S 148.01 E 1km SSE Gladstone Tas. 29 Jan 1983 J.C. Cardale’; 1 Ƥ ‘Mt Nelson Tas. 160m 3 Nov 1979 P.B. McQuillan’; 1 Ƥ ‘ 41.51 S 145.33 E Mt Read 500m Tas. 9 Jan 1989 P.B. McQuillan E.S. Nielsen’; 1 Ƥ ‘ 43.25 S 146.09 E Melaleuca Tas. 14 Jan 1991 E.S. Nielsen E.D. Edwards’; 1 Ƥ ‘ 43.23 S 146.08 E Claytons Bathurst Harb. Tas. 16 Jan 1991 E.S. Nielsen E.D. Edwards’; coll. TMAG: 13 ‘Tas. Pt. Sorell. Tas. U. V. L. 14.x. 1988 L. Hill’ ‘Registration No. F 4702 ’ ‘ TMAG genitalia slide F 4703 ’; 13 ‘ 41 ° 52 ’S 146 ° 30 ’E TAS. Central Plateau L. Augusta to L. Ada 1150 m 3 FEB. 1994. P. B. McQuillan uvl’ ‘Registration No. F 4704 ’ ‘ TMAG genitalia slide No. F 4705 ’; 13 ‘ 41 ° 52 ’S 146 ° 30 ’E TAS. Central Plateau L. Augusta to L. Ada 1150 m 3 FEB. 1994. P. B. McQuillan uvl’ ‘Registration No. F 4706 ’ ‘‘ TMAG wing slide No. F 4707 ’; 13 ‘S.W. TAS. 12 Trees Hill (Strathgordon) U.V.L. 17.xi. 1989. T. Semmens’ ‘Registration No. F 4708 ’ ‘ TMAG genitalia slide F 4709 ’; 13 ‘ 40 ° 6 ’ 8.53 ”, 148 °0’ 15.07 ” Bluff Rd. Whitemark Flinders Is. Tas. 31 Oct. 2008 C. J. Young U.V.L’. ‘Registration No. F 4710 ’; 1 Ƥ ‘ 41 ° 11 ’ 5 ”, 146 ° 19 ’ 5 ” Stony Rise Centre, Devonport. Tas. 23–29 Sept. 2004 L. Hill’; coll. ZSM: 1 Ƥ S.W. TAS. 12 Trees Hill (Strathgordon) U.V.L. 17.xi. 1989. T. Semmens’ ‘Accession No. 104062 ’; 1 Ƥ ‘Mt Wellington 870m [Tas.] 12 V UV 3 Nov. 1981 L. Hill’; 23 ‘ 40 ° 6 ’ 8.53 ”, 148 °0’ 15.07 ” Bluff Rd. Whitemark Flinders Is. Tas. 31 Oct. 2008 C. J. Young U.V.L’. ‘Accession Nos 104058, 104060’. Other material examined: New South Wales, coll. ANIC (all excluded from the type series): 1 Ƥ ‘Newnes S[tate].F[orest]. [NSW] 20 Nov 1993 J.C. Keast’; 1 Ƥ ‘ 36.28 S 148.27 E Rangers Stn 6km NE by E Thredbo NSW 1260m 6 Jan 2002 E.D. Edwards’; 1 Ƥ ‘ 36.23 S 148.25 E 1.5km NNW Smiggin Holes NSW 1700m 9 Jan 2002 E.D. Edwards’; 1 Ƥ ‘ 36.23 S 148.25 E Saddle 2km NW Smiggin Holes Kosciusko Nat. Pk 1680m 23 Jan 1987 E.D. Edwards’; 1 Ƥ ‘ 1.3 km E of Island Bend NSW 1158m 30 Nov 1970 I.F.B. Common J.S. Dugdale’; 23 ‘ 36.23 S 148.22 E Guthega Village Kosciusko N.P. 1700m [NSW] 24 Jan 1987 E.D. Edwards’, one with ‘ ANIC genitalia slide 18603 ’; 13 ‘ 36.23 S 148.25 E Saddle 2 km NW Smiggin Holes Kosciusko N.P. 1680m [NSW] 23 Jan 1987 E.D. Edwards’. Description. Wingspan 3 42–47 mm. Habitus and external characters (Figs. 2–3): Forewing apex tapered, termen convex. Hindwing termen straight, angled at apex and tornus. Ground colour variable, usually purplish with dark suffusion and with blackish dotting of forewing costa. Fringe crimson. Antemedial line of forewing vague, at costa usually well marked by a costal spot. Postmedial line of forewing almost straight but slightly undulating, ochre with dark grey dots basally, obliquely leading into forewing apex. Border of postmedial line on the hindwing more violet and forked towards costa. Forewing apex with blackish fringe. Cell spots slightly elongate, black, with small hyaline filling. Underside of forewing with a large, round dark violet spot close to the inner termen at 2 / 3, i.e. at the position of the dotted postmedial line. Underside of hindwing with large white and grey speckled spot close to the costa at 2 / 3, i.e. at the position of the dotted postmedial line. Palpi, frons, and vertex concolorous with ground colour. Frons flat. Palpi at upperside of tip with dark grey scales, last segment narrow, length of palpi ca 1.5 times diameter of eye in both sexes. Proboscis well developed. Antennal flagellum thick in both sexes, male antennae unipectinate to 2 / 3 of length, female antennae filiform. Frenulum strong in 3, absent from Ƥ. Wing venation, forewing: R separate from Rs, anastomosing with Sc for a short distance in the forewing, R 2–5 stalked. Hindwing: M 2 located at mid-point between M 1 and M 3. Hindleg with four very short spurs in both sexes. Ansa narrow at the base, widening mesally, tapering apically. Last abdominal segment with extremely large interior coremata. 3 genitalia (Fig. 8): Uncus long, narrow. Gnathos slender; medial process broad, sub-acute posteriorly, posterior surface covered with rows of short, broad spicules. Juxta large, well-sclerotised, divided. Saccus broad rectangular, with central invagination. Costa of valva sclerotised, setose. Valva asymmetrically adorned with subapical processes on ventral margin: left valva with single, large, sclerotised subapical, flattened, acute process, right valva with two short, broad processes. Aedeagus comparatively broad; vesica well sclerotised posteriorly, with longitudinal ridges; long narrow sclerotised process attached anteriorly, no discrete cornuti; caecum long, slender, tapered. Ƥ genitalia (Fig. 10): Apophyses anteriores, posteriores long, slender. Lamella postvaginalis membranous. Lamella antevaginalis poorly developed. Sternum A 7 slightly sclerotised. Ductus bursae short, close to corpus bursae strongly sclerotised, towards antrum dilating. Corpus bursae with strong anterior and posterior dilatations, constricted between; posterior half sclerotised, strongly ridged; anterior half membranous, plicate. Signum absent. Morphological diagnosis (most congeners illustrated in BOLD (2008)): Oenochroma vinaria in habitus and external characters (Figs. 4, 6) very close to Oenochroma barcodificata and externally only distinguished, usually, by the straighter postmedian fascia of the forewing, often with a continuous dark proximal border. Dark suffusion of ground colour and blackish dotting of forewing costa usually reduced. Oenochroma pallida Warren, 1898 differs from both Oenochroma barcodificata and Oenochroma vinaria at once by the fore tibiae having an anterior apical hook (cf. Prout 1910, Turner 1932), and by the ochreous brown fringe of wing termen, the larger forewing cell spot on a paler ground colour, inner termen of hindwing underside with a narrow, dark spot; Oenochroma orthodesma Lower, 1894 has pale ochreous grey ground colour and ochreous fringe, an ochreous postmedian line edged anteriorly with pale yellow, the (ochreous) discal dots mostly wanting, hindwing at apex with pink suffusion and no spot on underside; Oenochroma decolorata Warren, 1896, has grey forewings with fine brown irroration and purplish fringe, and a pale ferruginous postmedial fascia on both wings; Oenochroma phyllomorpha Lower, 1899 is of light brown ground colour, the forewing postmedial line is sinuate, fringe fuscous, and the cell spot lacking; Oenochroma cycnoptera Lower, 1894 has anterior tibiae with a strong apical hook (cf. Turner 1932), a very faintly ochreous postmedian line not reaching apex of forewing, fringe pale brownish, hindwings pale, without pattern and with whitish fringe; the New Caledonian Oenochroma unifasciata Holloway, 1979, is broad-winged, without discal dots to the forewing. Molecular diagnosis (see Fig. 1 and Table 2): DNA barcode analysis revealed a 3.34 % K 2 P divergence between Oenochroma vinaria and O. barcodificata. Each species displays a very low mean intraspecific variation, with 0.05 % (SE= 0.03 %, maximum distance of 0.33 %) and 0.03 % (SE= 0.03 %, maximum distance of 0.15 %) for O. vinaria and O. barcodificata respectively and are thus unambiguously characterized by their DNA barcodes. Distribution. Recorded in south-eastern and northern parts of Tasmania at altitudes from 0 to 870 m above sea-level, probably distributed all over the island. Further material from the central and southern tablelands of New South Wales dissected (T Edwards) but excluded from the type series to maintain geographical homogeneity in the type material. O. barcodificata 0.03 % 0.03 % 0.15 % n= 9 O. pallida 0.00% n= 3 O. decolorata 0.15 % n= 2 continued. B mean distance standard error minimal distance O. vinaria vs O. barcodificata 3.34 % 0.65 % 3.1 % O. barcodificata vs O. pallida 5.35 % 1.17 % 5.2 % O. barcodificata vs O. decolorata 6.31 % 1.02 % 6.2 % Biology and morphology of immature stages. Egg: Marked on all surfaces by round to hexagonal, concave cells with broad walls (Figs 13, 15– 18), micropylar cell walls narrow (Fig. 14). Aeropyles, slightly elevated, openings very small (Fig. 16), present on all surfaces apart from top of wider lateral side (Fig. 17). Micropylar cells slightly recessed. Micropyles offcentred (Fig. 13). Chorion undulating, irregularly pitted (Figs 16–18). Colour: Off-white (Fig. 12), becoming irregularly blotched rust-red, then transparent grey on maturity. Size (mm): (n= 4) L = 1.01 ± 0.01 (SE), W = 0.82 ± 0.01 (SE), T = 0.69 ± 0.01 (SE). Width/length: 0.81. Aeropylar opening size (micrometers) (n= 10): L = 1.44 ± 0.07, W = 1.08 ± 0.04; relative aeropylar opening size [aeropyle length (micrometers)/egg length (mm)]: 1.4. Micropyles (Figs 13, 14): distinct; no. of openings: 6; no. of cells in rosette: 9; no. of rows of cells in micropylar area: 5. Shape: Broad, bluntly ovoid, dorso-ventrally flattened, anterior pole angled to horizontal axis (Figs 12, 13, 15). Oviposition: 2 Ƥ code nos.: G 87, G 307. Oviposition period: 26 Sept.– 31 Oct. Batch size range: 5–60 (n = 2). Realised fecundity (range): 4–60. Incubation time (days): 9. Batch configuration and attachment: Attached, singly. Larval description. First instar (newly emerged): Head capsule width: 0.53 ± 0.03 mm (SE) (n= 4). Head ground colour burnt yellow with chevroned dark-brown blotches, mouthparts lighter in colour, stemmata black, frons convex. Thorax and abdomen ground colour cream, except A 6 -A 10 yellow, with numerous longitudinal wavy, dark brown stripes; posterior third of each segment, except A 6 -A 10 and thoracic segments white; thoracic stripes less dense; venter dark chocolate brown. Thorax and abdomen becoming a uniform chocolate brown broken by numerous thin, wavy cream stripes after feeding. Setae very short, white, blunt on small dark brown pinacula on small raised black tubercles. Thoracic legs white with sparse dark brown spots. Prolegs on A 5, A 6, A 10, rudimentary on A 5. Spiracles very small, brown. Anal shield, no stripes, yellow with sparse orange streaks. Crochet arrangement, incompletely interrupted mesoseries. Resting position about 30 ° from substrate. Second instar: Head capsule width: 0.91 ± 0.01 mm (SE) (n= 7). Head ground colour light yellow with dark-brown blotches, dorso-lateral orange stripes, mouthparts pale brown, stemmata black, frons convex. Thorax and abdomen ground colour dark brown, broken by numerous longitudinal, narrow wavy, white stripes, venter smoky brown. A 1 with dark brown spot on dorso-lateral line adjacent to anterior margin, directly adjacent and anterior to burnt orange blotch directly ventral to mid-dorsal line; A 3 with two small orange blotches directly adjacent to mid-dorsal line, about one third segment length from posterior margin in some specimens, venter with black speckles. Setae very short, black, blunt on small dark brown pinacula on small raised black tubercles. Thoracic legs white with small brown blotches. Prolegs on A 5, A 6, A 10, rudimentary on A 5, same ground colour as body. Spiracles small, cream, peritremes black. Anal shield, same ground colour as body. Third instar: Head capsule width: 1.48 ± 0.03 mm (SE) (n= 5). Head ground colour cream with chevroned black blotches dorsally just lateral of mid-dorsal line, becoming two large circular, black blotches with orange centres, just posterior of frontoclypeus, dorso-lateral orange stripes, frontoclypeus pale brown streaked with black and brown, other mouthparts pale brown; stemmata black, frons convex. Thorax and abdomen ground colour cream, broad, dense-grey mid-dorsal band, centred by narrow white stripe connecting mid-segmental diamond shapes, thin wavy, grey stripes on rest of dorsum becoming yellowish-brown laterally. A 1 with medial horn-like protuberances just lateral of mid-dorsal line, apex with black pinacula and setae, colour burnt orange with diagonal yellow stripe extending from mid-dorsal line to apex, black velvety blotch just anterior of protuberance at dorso-lateral position; similar small protuberances on A 3. A 8 D 1 pinacula large, raised, bright burnt orange, venter paler from A 3 to A 10. Setae short, black, blunt, longer, transparent on venter A 6- 10. Thoracic legs cream with small black blotches. Prolegs on A 5, A 6, A 10, rudimentary on A 5, same ground colour as body. Spiracles pale brown, peritremes black. Anal shield, same ground colour as body. Fourth instar: Head capsule width: 2.31 ± 0.04 mm (SE) (n= 4). Head ground colour light orange-brown mottled heavily with dark orange-brown, lighter stripes extend from mid-dorsal line to frons, becoming lighter and wider anteriorly, mouthparts pale brown, stemmata black, antennae brown; frons convex. Thorax and abdomen ground colour light brown, scattered sparsely with small white spots; dorsal band white speckled with black, on thoracic segments band is defined loosely by lines of black spots, on abdominal segments band is constricted at segment margins and bulges at segment centres; large fleshy horn-like dorsolateral protuberances on A 1 just posterior of middle of segment, protuberances on A 2 similar to A 1 but smaller, apices of protuberances mottled orange-brown, large orange blotch mottled yellow, just anterior of protuberances, not extending laterally ventral of protuberances; venter pale brown speckled black; D 1 pinacula on A 8 on bright orange protuberances. Thoracic legs pale brown with sparse small black blotches. Prolegs on A 5, A 6, A 10, rudimentary on A 5, same ground colour as body. Spiracles orange, peritremes black, on white blotched with grey large spot. Fifth instar (Fig. 19): Head capsule width: 3.78 ± 0.03 mm (SE) (n= 4). Length: 45–50 mm (n= 3). Similar to 4 th instar, except for the following. Head ground colour pinkish brown with faint orange marbling, mouthparts pale brown, stemmata black, antennae reddish brown; frons convex. Thorax and abdomen ground colour light brown, scattered sparsely with small white spots; dorsal stripe light brown speckled with small black spots, two lines of small white spots circled in black define dorsal stripe on abdominal segments. Two large fleshy dorso-lateral protuberances on A 1, apices black anteriorly, yellow posteriorly, horns preceded by large mid-dorsal yellow blotch narrowing anteriorly to A 1 anterior margin; relatively smaller protuberance on A 3 similar to protuberance on A 1. Venter pale brown blotched dark brown, white between A 6 –A 10. D 1 pinaculi on A 8 on bright orange protuberances. Black spots surround pinaculi of sub-dorsal setae. Setae short, pale brown, blunt. Thoracic legs pale brown with sparse small black blotches. Prolegs on A 5, A 6, A 10, rudimentary on A 5, same ground colour as body. Spiracles orange, peritremes black. Chaetotaxy: trisetose SV setae on A 1; SV 1, SV 3 and V 1 setae on A 1 unaligned; three lateral setae on A 6 proleg. Pupa. Length: ɗ 23, 24 mm; Ψ, 24, 25 mm. Width: ɗ 7, 7 mm; Ψ, 6, 8 mm. Colour: reddish-brown on maturity; wings duller than shiny abdomen. Silken cocoon constructed from soil, debris and body fluids. Pupa large, stout. Labrum sub-trapezoidal, slightly convex, well-defined. “Mandibles” flat, rugose, margin between labrum and “mandibles” broadly ridged. Labial palpus large, pentagonal, rugose. Border between genae, maxilla less steep than that between oculus and pro-leg; border between pro-tibia and antenna about same length as border between pro-tibia and oculus. Pro-tibia same length as mid-tibia. Pro-femora not visible, Protibia meets maxilla at 2 / 3 length of maxilla, mid-tibia meets maxilla at 5 / 6 length of maxilla, antenna broad, almost reaches apex of maxilla, hind-leg barely visible. Hindwing becoming concealed at A 4. Meta-notum short. Wing-bud callosity pronounced. Thoracic spiracles not visible, spiracles visible on A 2 -A 8, poorly developed on A 8; elliptical, elevated, pre-spiracular slit visible. Punctation on A 1 -A 8, punctures small, shallow, numerous on A 2 -a 7, sparse on A 1, A 8; randomly arranged, uniformly sparse on venter. Setae very short. Anal area large, anal slit bordered by longitudinal furrows. Male genitalia area, simple longitudinal slit. Cremaster, long, slender, roughly trigonal, posterior third, very rugose; one pair of terminal, robust, hamate, long, parallel setae, directed ventrally; pronounced dorsal and lateral grooves, six pubescent teeth on anterior margin of A 10. Larval hostplants. Reared on Grevillea sp. (Proteaceae). Etymology. The species name refers to t

Genus Oenochroma Guen��e, 1858 All 20 species of the genus Oenochroma Guen��e, 1858 as listed in Scoble (1999) are distributed in mainland Australia except one species which is restricted to New Caledonia (Oenochroma unifasciata Holloway, 1979). From dissections, it is most likely that Oenochroma hieroglyphica (Warren, 1906), which was described from Papua New Guinea, Mt. Kaindi, is a junior synonym of Oenochroma turneri (Lucas, 1892), described from Toowoomba near Brisbane. Two species are known to occur both in mainland Australia and in Tasmania: Oenochroma vinaria and Oenochroma vetustaria (Walker, 1860) (McQuillan 2004). The type species of the genus, Oenochroma vinaria, boasts the violaceous red "wine" colour ("rose-lie de vin" in the original description) that may have suggested to Guen��e the chosen denominations of the species and of the genus. Only Oenochroma vinaria and the new species described here are intended to fall in the ��� vinaria speciescomplex.��� It is possible that they may group together, phylogenetically, with Oenochroma pallida and Oenochroma decolorata. However, both the definitions of infrageneric species-groups such as of the whole genus Oenochroma await an urgently needed major revision. Collection material (ANIC, ZSM, TASAG) suggests that the genus includes several other undescribed species., Published as part of Hausmann, Axel, Hebert, Paul D. N., Mitchell, Andrew, Rougerie, Rodolphe, Sommerer, Manfred, Edwards, Ted & Young, Catherine J., 2009, Revision of the Australian Oenochroma vinaria Guen��e, 1858 species-complex (Lepidoptera: Geometridae, Oenochrominae): DNA barcoding reveals cryptic diversity and assesses status of type specimen without dissection, pp. 1-21 in Zootaxa 2239 on page 7, DOI: 10.5281/zenodo.190505, {"references":["Guenee, A. (1858) Uranides et Phalenites. In: Boisduval & Guenee: Histoire naturelle des Insectes. Species general des Lepidopteres I, Vol. 9.","Scoble, M. J. (ed.) (1999) Geometrid moths of the world: a catalogue. Apollo Books, Stenstrup, 116 pp.","McQuillan, P. B. (2004) An overview of the Tasmanian geometrid moth fauna and its conservation status. Journal of Insect Conservation, 8, 209 - 220."]}

Oenochroma vinaria Guen��e, 1858, redefined Oenochroma vinaria Guen��e, 1858: in Boisduval & Guen��e, Hist. nat. Insectes (Spec. g��n. L��pid.) 9: 185, pl. 7, fig. 2. Syntypes 231 �� (MNHN), lectotype �� designated by Viette (1950) using the term ���type��� (cf art. 74.5, Code ICZN 1999), Nouvelle Hollande [Australia], Barcode no. BC-MNHN 0 0 0 7 (Figs 4, 5). One further paralectotype, mentioned in Viette (1950) as ���paratype���, could not be traced in MNHN and may be lost or destroyed. Monoctenia decora Walker, 1869: Charact. Undescr. Lepid. Heterocera: 76 (Australia, not exactly specified, but very likely ���Victoria��� (TE)). Syntype (s) �� (MVMA). At MVMA only one syntype could be traced, which is designated herewith as lectotype (cf. Figs 6, 7), in accordance with Article 74.7. 3 of the Code (ICZN), in order to define the herewith presented synonymy. Synonymization with true vinaria (cf. McQuillan & Edwards 1996, Scoble 1999) is supported by the straight postmedial line of the forewing, and structure of genitalia (examination by CY). Redescription. Wingspan 40���50 mm. Habitus and external characters (Figs. 4, 6) very close to Oenochroma barcodificata. Ground colour variable, usually wine-red, sometimes brown or sand colour. Postmedial fascia of the forewing straight, often with a continuous dark proximal border. Dark suffusion of ground colour and blackish dotting of forewing costa usually reduced. Tasmanian populations of Oenochroma vinaria are characterised by a darker ground colour than in the nominotypical continental populations. It is worth noting that the figure in Boisduval & Guen��e is too drab and not enough rosy as Guen��e himself asserts in the original description. 3 genitalia (Fig. 9): Uncus slightly longer and narrower than in Oenochroma barcodificata. Valvae broader. Process of left valva slightly larger. Right valva without the two spines arising from the edge subapically but with vestigial spinules on the inner surface of valva. Tip of aedeagus narrower than in Oenochroma barcodificata. Sclerite of vesica much smaller. �� genitalia (Fig. 11): Differing from those of Oenochroma barcodificata in the strong sclerotisation of corpus bursae at the junction with the ductus bursae. Anterior and posterior dilatations of corpus bursae less pronounced. Distribution. Tasmania and mainland Australia, widely distributed from Murchison River in West Australia through South Australia, Victoria and New South Wales to the Atherton Tableland in northern Queensland. Biology. Larvae of ��� O. vinaria ��� are recorded as feeding on various species of the Proteaceae genera Hakea, Grevillea and Banksia (McFarland 1988. Herbison-Evans & Crossley 2006; T. Edwards pers. obs.), for details see McFarland (1988). All these data refer to populations outside the known range of the sister species O. barcodificata, and for material from the host-plant genera Hakea, and Grevillea species identity as O. vinaria could be verified definitely (T. Edwards). Remarks. Both McFarland (1988) and Herbison-Evans & Crossley (2006) highlight the large variability of larvae in colouration and even in shape e.g., length of fleshy dorsal appendages on segment A 3. Further research is needed to examine if this variation refers to the presence of different taxa also in South Australia and Victoria. The following illustrations in the literature refer to true O. vinaria after re-examination: McFarland (1988: 157), Common (1990: pl. 10, fig. 12; pl. 26, fig. 10), Coupar & Coupar (1992: 48), Zborowski & Edwards (2007: 143), Daley (2007: 194), Richardson (2008: 19), Willan (2008). Unipectinate antennae, although rare in the Geometridae, are characteristic of Proteaceae-feeding oenochromines (Scoble & Edwards 1990). Oenochroma vinaria, O. barcodificata and other Oenochrominae s. str. possess two wing venational characters usually present in Geometrinae: R separate from Rs and anastomosing with Sc for a short distance in the forewing and R 2���5 stalked in the forewing (Young 2006 b). Also, the ansa is similar to the geometrine type i.e., narrow at the base, widening mesally and again tapering apically, and is not the more typical tapering ansal morphology characteristic of Oenochrominae s. l. (Cook & Scoble 1992). As in the Geometrinae, the caecum of the aedeagus is long, slender and tapered and cornuti are reduced, tending not to be discrete rods and spines (Young 2006 b). Similar aedeagus characteristics are found in the Australian oenochromine s. str. Monoctenia falernaria Guen��e, 1858 (Young 2006 b). This apparently close relationship between the Oenochrominae s. str. and the Geometrinae is also supported by molecular data (Young 2006 b, Yamamoto & Sota 2007). The mature larva of O. barcodificata was exceptional among other geometrids examined in a study of Australian Geometridae by Young (2006 b) by possessing trisetose SV setae on A 1. The bifurcate cremastral spines, punctation, dorsal and lateral grooves and mesothoracic spiracles described in O. barcodificata here were also noted in the oenochromines Arhodia lasiocamparia, Guen��e, 1858, Monoctenia falernaria Guen��e, 1858, M. smerintharia Felder & Rogenhofer, 1875, Dinophalus drakei (Prout, 1910), Hypographa Guen��e, 1858, Parepisparis lutosaria (Felder & Rogenhofer, 1875) and Phallaria ophiusaria Guen��e, 1858 (McFarland 1988)., Published as part of Hausmann, Axel, Hebert, Paul D. N., Mitchell, Andrew, Rougerie, Rodolphe, Sommerer, Manfred, Edwards, Ted & Young, Catherine J., 2009, Revision of the Australian Oenochroma vinaria Guen��e, 1858 species-complex (Lepidoptera: Geometridae, Oenochrominae): DNA barcoding reveals cryptic diversity and assesses status of type specimen without dissection, pp. 1-21 in Zootaxa 2239 on pages 18-19, DOI: 10.5281/zenodo.190505, {"references":["Guenee, A. (1858) Uranides et Phalenites. In: Boisduval & Guenee: Histoire naturelle des Insectes. Species general des Lepidopteres I, Vol. 9.","Viette, P. (1950) Sur quelques especes de geometrides decrites par Guenee (1857). Bulletin mensuel Societe linneenne Lyon, 19, 201 - 206.","ICZN, International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature, Fourth Edition, adopted by the International Union of Biological Sciences. The International Trust for Zoological Nomenclature, c / o The Natural History Museum, London, 306 pp.","McQuillan, P. B. & Edwards, E. D. (1996) Geometridae. In: Nielsen, E. S., Edwards, E. D., & Rangsi, V. (eds): Checklist of the Lepidoptera of Australia. CSIRO, Melbourne, pp. 220 - 228, 355 - 357.","Scoble, M. J. (ed.) (1999) Geometrid moths of the world: a catalogue. Apollo Books, Stenstrup, 116 pp.","McFarland, N. (1988): Portraits of South Australian Geometrid Moths, Allen Press, Lawrence, 399 pp.","Herbison-Evans D. & Crossley, S. (2006) Oenochroma vinaria Guenee, 1857. In: Herbison-Evans D. & Crossley, S. (eds): Geometridae of Australia. Available from: http: // www. usyd. edu. au / museums / larvae / geometridae / geometridae. html (accessed 1 December 2008).","Common, I. F. B. (1990) Moths of Australia, Melbourne University Press, Melbourne, EJ Brill, Leiden, 535 pp.","Coupar, P. & Coupar, M. (1992) Flying Colours. Common caterpillars, butterflies and moths of south-eastern Australia. New South Wales University Press, Kensington.","Zborowski, P. & Edwards, E. D. (2007) A field guide to Australian moths. CSIRO Publishing. Collingwood.","Daley, E. (2007) Wings. An introduction to Tasmania's winged insects. Riffles, Buckland. P.","Richardson, B. (2008) Mothology: Discover the magic. Moths and moth art from the World Heritage listed wet tropics of Queensland Australia. LeapFrogOz, Kuranda.","Willan, L. (2008) Oenochroma vinaria Available from: www. ento. csiro. au / gallery / moths / Oenochromavinaria (accessed 11 December 2008).","Scoble, M. J. & Edwards, E. D. (1990) Parepisparis Bethune-Baker and the composition of the Oenochrominae (Lepidoptera: Geometridae). Entomologica Scandinavica, 20, 371 - 399.","Young, C. J. (2006 b) Molecular relationships of the Australian Ennominae (Lepidoptera: Geometridae) and implications for the phylogeny of the Geometridae from molecular and morphological data. Zootaxa, 1264, 1 - 147.","Cook, M. A. & Scoble M. J. (1992) Tympanal organs of geometrid moths: a review of their morphology, function, and systematic importance. Systematic Entomology, 17, 219 - 232.","Yamamoto, S. & Sota, T. (2007) Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 44, 711 - 723.","Prout, L. B. (1910) Geometridae, Oenochrominae. In: Wytsman, P. (ed.): Genera Insectorum. Lepidoptera Heterocera. Fasc. 104, 120 pp. (p. 40 - 42)"]}