Diphtheroptila virosae De Prins, Sruoga & Zwick, 2025

Prins, Jurate De, Hartley, Diana, Sruoga, Virginijus, Nicholls, James, Wallace, Jesse & Zwick, Andreas, 2025, Diversity of Australian Ornixolinae (Lepidoptera: Gracillariidae) with taxonomic and nomenclatural acts within the related taxa (Acrocercopinae and Gracillariinae) based on the evidence of museomics, bionomics, and mitogenomics, Zootaxa 5616 (1), pp. 1-340 : 121-143

Diphtheroptila virosae De Prins, Sruoga & Zwick , sp. nov.

( Figs 152, 177–179, 206–208, 228, 229, 250, 251, 261, 271, 278, 287, 638)

Type locality: Australia, Northern Territory, Darwin.

Type specimens: Holotype ♂: [labels verbatim] [1] Australia N.T. [Northern Territory]/ 12.21S 130.52E / Casuarina C.R./Darwin em.[erged]/ 1 Feb 1998 /T. & M. Kumata. [2] Host 5744/ Flueggea / virosa, DNA sample NULT024887, genitalia slide ANIC 6172, ANIC Acc. no 31 085526, in ANIC (Canberra).

Paratypes: 15 specimens (4♂, 9♀, 2 specimens): Paratype 1(♀): Australia, Northern Territory, Darwin, Casuarina C.R., 12.21S 130.52E, 31 January 1998, T. & M. Kumata. Host 5744, Flueggea virosa (Roxb. ex Willd.) Royle ( Phyllanthaceae ). Paratype 2(♂): same data, except the date, 2 January 1998. Paratype 3(♂): same data, except the date, 31 January 1998. Paratype 4(♀): same data, except the date, 1 February 1998. Paratype 5(♀): same data, except the date, 4 February 1998. Paratype 6(♀): same data except the date 2 February 1998. Host 5745, Flueggea virosa (Roxb. ex Willd.) Royle ( Phyllanthaceae ), DNA sample NULT024762, genitalia slide ANIC 6171, ANIC Acc. no 31 085551. Paratype 7(♀): same data except the date 2 February 1998, DNA sample NULT025008, genitalia slide ANIC 6173, ANIC Acc. no 31 085552. Paratype 8: without abdomen; same data. Paratype 9(♀): same data, except the date, 2 January 1998. Host 5744, Flueggea virosa (Roxb. ex Willd.) Royle ( Phyllanthaceae ). Paratype 10(♀): same data, except the date, 2 February 1998. Host 5745, Flueggea virosa (Roxb. ex Willd.) Royle ( Phyllanthaceae ). Paratype 11(♂): same data, except the date, 2 February 1998. Paratype 12(♀): same data, except the date, 31 January 1998, Host 5744, Flueggea virosa (Roxb. ex Willd.) Royle ( Phyllanthaceae ). Paratype 13: without abdomen, same data, except the date, 1 February 1998. Host 5744, Flueggea virosa (Roxb. ex Willd.) Royle ( Phyllanthaceae ). Paratype 14(♀): left forewing, left hindwing absent, same data, except the date, 2 January, 1998, in ANIC (Canberra).

Queensland: Paratype 15(♀): Kuranda, 16.49°S 145.38°E, em.[erged] 7 February 1997, leg. T. & M. Kumata, Host 5708 Glochidion sp. , DNA sample NULT025497, genitalia slide ANIC 6184 , ANIC Acc. no 31 085553, in ANIC (Canberra) .

Type depository: Australian National Insect Collection, Canberra, Australian Capital Territory, Australia.

Diagnosis: The main diagnostic feature of Diphtheroptila virosae sp. nov. is a peculiar pattern of the forewing consisting of an intermix of transverse dirty white-grey irregular fasciae /dorsal stripes, usually coupled or grouped in three with some black spots and patches that are mainly concentrated in the sub-apical costal area. Species belonging to Diphtheroptila cannot be diagnosed by external characters, difficult to diagnose by host plants, since the majority of Diphtheroptila species feed on Glochidion spp. The most reliable, clear and undoubtful diagnostic character set is in genitalia micromorphology. As we see from the type specimens presented above Diphtheroptila virosae sp. nov. feeds on two different host plants belonging to the same host plant family— Phyllanthaceae — Flueggea virosa (Roxb. ex Willd.) Royle and Glochidion sp. Two new species of the genus Diphtheroptila , D. breynella sp. nov. and D. virosae sp. nov., presented in this monograph feed on the same host— Flueggea virosa . Externally D. virosae sp. nov. can be separated from D. breynella sp. nov. due to indistinct, dirty white, hardly traceable oblique fasciae, which is very typical forewing pattern of Diphtheroptila species feeding on Glochidion spp. plants, while in D. breynella sp. nov. dorsal markings of forewing are bright white, short, thick, oblique dorsal strigulae are grouped per three. In the male genitalia of D. virosae sp. nov. the apomorphic character set is found in the shape and structure of valvae. In D. virosae sp. nov. the apical part of valva is more or less triangular, with a sharply pointed cucullus, while in other Diphtheroptila species the apical part of valva is more or less gently rounded. The saccular sclerotised dactyliform appendage, present only in D. virosae sp. nov., is a species-linked apomorphic character and highly diagnostic. In the female genitalia, round ostium bursae with narrow sclerotised marginal ring and two signa, present on corpus bursae (smaller in the posterior part and bigger rhomboid in the anterior part), are diagnostic apomorphies for D. virosae sp. nov. Another diagnostic character set is represented by mitogenomics.

Description: Wingspan 5.1–5.6 mm; length of the forewing 2.4–2.6 mm ( Fig. 152).

Head ( Figs 177–179): Vertex light fuscous grey consisting of one bunch of tightly supressed piliform scales, gently rounded around base of antenna and on occiput, unicoloured. Frons of the same colour as vertex, thickly covered with piliform supressed scales. Maxillary palpus dirty white, slightly shorter than the first labial palpomere. Labial palpus impressively developed, drooping, with long brown scales, intermixed with white piliform scales, hanging on the second palpomere, second palpomere dilating with broader apical part, dirty white base and brown apex and lateral sides, terminal palpomere snowy white with sharply pointed apex, labial palpus ca. 2, 5× longer than the eye. Antenna beige with fuscous longitudinal lines with intermixed fuscous, consisting of piliform narrow scales forming longitudinal thin lines and lighter apices, ventrally uniformly light ochreous, pedicel short, light grey; scape dorsally light grey, ventrally almost white, 9–12 pecten short, stout, almost white.

Thorax ( Figs 152, 207, 208): lightly beige, intermixed with lighter beige spots, tegula fuscous-grey. Forewing narrowly elongated, equal in width along its entire length, with a gently rounded apex, ground colour beige with fuscous spots, wing markings sometimes could be divided into three sectors: base, median and apical; at base slightly arched dirty white fascia cross the forewing, irregularly bordered with dark fuscous scales, sub-dorsal area of base heavily marked with dark fuscous scales, white sub-costal are is irrorated with light fuscous spots, median part consists of irregularly shaped fascia with shorter stripes or fasciae running parallel to major marking, interrupted by two rows or irregular shorter fuscous stripes (variable), two fuscous spots, variable in shape even on the same specimen left or right forewing, one on sub-dorsal and the other on sub-costal area follow the median fascia; sub-apical area is marked by three oblique dirty white sub-dorsal tripes not reaching the midden of the forewing, followed by a big prolonged dirty white spot situated at tornus of the forewing, apical area and apical spot is marked as irregularly shaped dark fuscous spot, surrounded by two-three short, stripes, three short stripes, the middle is the longest are present in the sub-apical area of forewing. The fringe line is dark fuscous and present only in tornal area; fringe long, at distal part of dorsum, light grey, with light silver shine, and shorter at median part of dorsum of forewing, shading a little bit darker towards the base of forewing. Hindwing narrow, elongate, sharply pointed, ground colour dark greyish fuscous, fringe shorter at costa and long, ca. 6× longer than the width of hindwing at the base, with the longest piliform scales hanging at the base of the dorsum of the hindwing. The fore femur is light beige, the fore tibia is fuscous with white median patch and lighter base, tarsomere I dark fuscous, tarsomeres II–IV fuscous with dirty white apical halves, terminal tarsomere is light grey with darker tip; mid-femur light ochreous-fuscous, mid tibia fuscous with two dirty white prolonged oblique stripes, tibial spurs light beige as long as tarsomere I, tarsomere I fuscous with dirty white sub-basal and apex, tarsomere II–IV fuscous with dirty apices, terminal tarsomere light grey with darker tip; hind femur ochreous beige, hind tibia fuscous with 15–17 stout spines stretching in a row from median to sub-apical part, median spurs very long, almost as long as tibia, hind tarsomeres dark fuscous with dirty white apices, hind tarsomere I with a row of shorter spines, tip is dirty white.

Abdomen ( Figs 206, 261, 271): bronze-beige on tergites I–II, dark fuscous stripe stretches along the median part of the rest of tergites, median part of sternites ochreous-white, lateral sides of abdomen (tergites and sternites) are dirty white with four oblique bronze-ochreous stripes. Margins of abdominal opening on sternum II strongly sclerotised, especially ventral crossing joint which is ca. double as thick as lateral sides, sternal apodemes initiating at the corners of abdominal opening are well developed, clearly observable, straight, thin, ca. 4× shorter than tergal apodemes; a lightly melanised joint connecting lateral sides of abdominal opening on tergum I is present, which is an initiating point of tergal apodemes; tergal apodemes sharply hooked at bases, thin, slightly bent, rather long, reaching sub-posterior part of tergum II; apical part of tergal apodemes blunt.Anterior segment VI in females without sclerotisations; anterior segment VII in males with two narrow, sclerotised semi-rings which are the androconial characters. Anterior margins of segments III–VI in females and III–VII in males narrowly and finely sclerotised.

Male genitalia ( Figs 228, 229): Tegumen short, reduced, teguminal arms are non-sclerotised, anal tube strongly protruding; valvae sharply narrowing from the midden part with narrow, but gently rounded apex; costal margin folded at basal and sub-basal parts till the mid of valvae, basal part of valvae carries paired appendages (on both valvae) that are zigzagging and angulated 2×, ending on saccular part of valva with a prolonged, more or less rectangular shaped, sclerotised tuberculated and enlarged appendage. Valvae symmetrical, rather short, slightly longer than vinculum, heavily setose from mid till apical part, especially ventral area is covered with long, strong, spiculose setae; ventral margin strongly narrowing (ca. 30°) from mid part of valva, ending in a narrow but gently rounded apex; transtilla incomplete; juxta as two round sclerotisations touching each other with tiny narrow appendices; vinculum is short but well developed, vincular internal sclerotisation W-shaped at anterior part, consisting of two fused parts with clearly visible suture, proximately narrow but strongly sclerotised bow reaching the base of tegumen; saccus short but well developed V-shaped, crossed by mid-suture with an arrow-shaped pointed anterior part. Aedeagus is ca. as long as valva, thick, well developed with narrowing and pointed vesica, that carries rather long, oblique strongly sclerotised one cornutus.

Female genitalia ( Figs 250, 251): Papillae anales flattened and fused, covered with short, erect and dense setae; apophyses posteriores short, rather thick with blunt apices; segment VIII is short, weakly sclerotised but carries a broad basal semi-ring of apophyses anteriores; the apical narrow part of apophyses anteriores is ca. as long as of apophyses posteriores with sharp apices terminating in the posterior 1/3 of segment VII. Segment VII is strongly sclerotised and sternum VII is almost completely covered by two-part folded sterigma with V-shaped indentation at the central part of posterior margin of sternum VII; sterigma gently surrounds the ostium bursae, as circular ring opens at anterior margin of sternum VII; margins of ostium bursae are strongly sclerotised, lateral margins of lamella ante-vaginalis are covered with numerous lamellar, flat, nicely arranged brushes of scales that are tightly attached to the borders of sterigmatic sclerotisations. Ductus bursae in corpore is protected by antral, strongly sclerotised, supporting fold; ductus spermathecae short, ca. 1/3 length of segment VII, spiralled into tight six convolutions, bulla small, indistinguishable from the anterior part of ductus spermathecae; ductus seminalis enters the anterior 1/3 of ductus bursae, curved, slightly sinuating, lightly sclerotised, bulla seminalis ca. as long as corpus bursae, sac-shaped, but very weakly melanised; the wall of corpus bursae with visible scales, corpus bursae sac-shaped, without evident distinction between corpus bursae and ductus bursae, except antral sclerotised supporting fold, with two signa of different shapes and size: a smaller signum at posterior 1/3 of corpus bursae, strongly sclerotised small -horn shaped, the bigger signum at anterior 1/3 of corpus bursae, shaped as a sclerotised triangular plate with sharp bigger horn-like process.

Individual variation: There is quite significant variation in the pattern of forewing for interchangeable stripes, dots and small fuscous spots, the ornamental variation can be observed even in the same specimen on right and left forewing. However, the principal pattern of sub-basal arched fascia, median fascia interrupted with fuscous lines/ stripes, oblique dirty white dorsal stripes and fuscous sub-costal patches remain stable.

Bionomics ( Figs 278, 287): The larva is a monophagous leaf miner found feeding on Flueggea virosa (Roxb. ex Willd.) Royle ( Fig. 287) and Glochidion sp. ( Phyllanthaceae ). The peak of mining period is in the second half of January till the first decade of February. The flight period of adults is the end of January up to mid of February.

Mitogenomic data: Despite having been sampled across its geographical range, the mitochondrial genomes of the populations in the Northern Territory and Queensland do not differ significantly, and the monophyly of the species is maximally supported in all analyses. The species is very strongly supported as sister to D. nix sp. nov. + D. crotonella sp. nov. ( Fig. 638).

Distribution: Known from two localities: the type locality in Australia: Northern Territory, Darwin, and Queensland, Kuranda.

Etymology: The specific name is derived from the species name of host plant Flueggea virosa . It is an adjective in feminine gender in genitive case. The adjective virosa in Latin means having unpleasant strong taste or smell.

12. Dysectopa Vári, 1961

“ Dysectopa gen. nov. ”—Vári, L., 1961. Transvaal Museum Memoir 12: xviii (key), 57.

Type species: Dysectopa scalifera Vári, 1961 . Transvaal Museum Memoir 12: 57–58, by original designation. BOLD data: No data.

GenBank data: No data.

Mitogenomic data: No data.

Bionomics: No data.

Distribution: Afrotropical Region: Namibia.

Species richness: World: 1 species; Australian Region: 0 species.

Type species: Dysectopa scalifera Vári, 1961

“ Dysectopa scalifera spec. nov. ”—Vári, L., 1961. Transvaal Museum Memoir 12: 57–58, pl. 20, fig. 5; pl. 43, fig. 1; pl. 70, fig. 3; pl. 107, fig. 15.

Type locality: South-West Africa [ Namibia], Otjiwarongo District, Abachaus.

Type specimen: Holotype ♀, genitalia slide G7697, in TMSA (Pretoria).

BOLD data: No data.

GenBank data: No data.

Mitogenomic data: No data.

Bionomics: No data.

Distribution: Afrotropical Region: Namibia.

13. Epicephala Meyrick, 1880

“ Epicephala gen. nov. ”—Meyrick, E., 1880. Proceedings of the Linnean Society of New South Wales 5(1): 137 (key), 168.

Type species: Epicephala colymbetella Meyrick, 1880 . Proceedings of the Linnean Society of New South Wales 5: 169, by monotypy.

Synonyms: Iraina Diakonoff, 1955 — Verhandelingen van de Koninklijke Nederlandse Akademie der Wetenschappen (2) 50 (3): 84 (key), 92–93. Type species: Iraina periplecta Diakonoff, 1955 , by original designation, synonymised by Vári (1961: xii, 42).

Leiocephala Kuznetzov et Baryshnikova, 2001 — Trudy Zoologicheskogo Instituta, Rossijskaya Akademija Nauk 291: 32–33. Type species: Epicephala colymbetella Meyrick, 1880 , by original designation. Leiocephala was established to denote a subgenus of Epicephala Meyrick, 1880 , synonymised by De Prins & De Prins (2005: 177). Morphological diagnostic characterisation. Wingspan 6–11 mm; length of the forewing 3–5 mm.

Head: Vertex smoothly covered by long piliform scales projected anteriorly, occiput with two lateral tufts of short piliform scales directed radially, frons more or less triangularly shaped with a loose bunch of piliform scales on labrum, maxillary palpus short with blunt apices, labial palpus slightly longer than the eye, with sharp apices, basal palpomeres covered with short loosely attached scales, a diagnostic character for Epicephala species; antenna slightly longer than forewing, scape carries pecten.

Thorax: Forewing rather broad in comparison with other genera of Gracillariidae , equally wide along its entire length, with a white ornamental pattern on dorsal margin; costal margin from mid of forewing is decorated with very narrow, hardly visible oblique stripes and tiny spots; sub-apical costal margin is marked by three oblique narrow stripes abruptly terminating at sub-apical area; apical spot small, oval, but clearly visible, apical line narrow, distinct, black, continuous; fringe line very short and present only at apex. The wing ornamentation is variable in thickness, edging, and obliqueness of stripes. Hindwings are very narrow, with sharply pointed apices, a character shared by all Epicephala species. Mid legs without pilose covering, which is a supportive character to separate Epicephala species from those of Cuphodes . In Epicephala hind tibia bear a row a sharply pointed, spiculose scales, a characteristic trait for Australian Ornixolinae genera.

Abdomen: Tergal apodemes in segment II with transverse joint, that might be strongly or weakly sclerotised, a diagnostic characteristic for Epicephala . Exception: E. breyniphaga sp. nov. where the anterior margin of sternal plate is strongly sclerotised and plays the function of transverse support.

Male genitalia: With very strongly developed sacculus.

Female genitalia: papillae anales and segment VIII fused into ovipositor, apophyses posteriores and anteriores thick, long, very strong. Fused apophyses posteriores have a piercing function.

BOLD data: https://www.boldsystems.org/index.php/Taxbrowser_Taxonpage?taxon= Epicephala &searchTax=Sear ch+Taxonomy

GenBank data: https://www.ncbi.nlm.nih.gov/nuccore/?term= Epicephala

Mitogenomic data: Epicephala is a species-rich and reasonably well-sampled genus, with relationships between species being robustly to maximally supported ( Fig. 637). Its monophyly is maximally supported by all analyses, while its placement as sister to all other Ornixolinae is recovered by NT, CODON and AA analyses with moderate to strong support, as compared to DEGEN placing the genus without statistical support one node deeper inside Ornixolinae ( Fig. 639).

Bionomics: Mainly fruit piercing and mining. Fabaceae : Bauhinia purpurea L., Bauhinia sp. , B. variegata L. ( E. orientale Stainton, 1856 ), Cajanus cajan (L.) Millsp. ( E. vermiformis Meyrick, 1936 ).

Phyllanthaceae : Breynia fruticosa (L.) Müll.Arg. ( E. lativalvaris Li, Wang & Zhang in Zhang et al. (2012), E. mirivalvata Li, Wang & Zhangin Zhang et al. (2012)) , B. oblongifolia (Müll.Arg.) Müll.Arg. , new record ( E. trigonophora ( Turner, 1900) , E. breyniphaga sp. nov.), B. rostrata Merr. ( E. lativalvaris , E. mirivalvata , B. vitisidaea (Burm.f.) C.E.C. Fisch ( E. sphenitis Meyrick, 1931 , E. vitisidaea Li, Wang & Zhang, 2012 ), Flueggea suffruticosa (Pall.) Baill. ( E. relictella Kuznetzov, 1979 ), Glochidion daltonii (Müll.Arg.) Kurz ( E. daltonii Li, 2016 , E duoplantaria Li, 2016 ), G. ferdinandii (Müll.Arg.) F.M. Bailey ( E. colymbetella ), G. philippicum (Cav.) C.B.Rob. new record ( E. philippa sp. nov.), Glochidion sp. ( E. ancylopa Meyrick, 1918 , E. frenata Meyrick, 1908a ), G. sphaerogynum (Müll.Arg.) Kurz ( E. angustisaccula Li, 2015 , E. camurella Li, 2015 , E. domina Li, 2015 , E. impolliniferens Li, 2015 ), G. zeylanicum (Gaertn.) A.Juss. ( E. bipollenella Li, Wang & Hu in Zhang et al. (2012)), G. zeylanicum var. tomentosum (Dalzell) Trimen ( E. bipollenella ), G. xerocarpus O.Schwarz , new record ( E. xerocarpa sp. nov.), Phyllanthus assamicus Müll.Arg. ( E. assamica Li, 2016 ), P. eriocarpus (Champ. Ex Benth.) Müll.Arg. ( E. eriocarpa Li, Wang & Zhang, 2012 ), P. lanceolarius (Roxb.) Müll. Arg. ( E. lanceolaria Li, Wang & Zhang, 2012 , E. lanceolatella Kawakita & Kato, 2016 , E. perplexa Kawakita & Kato, 2016 ), P. lutescens (Blume) Müll.Arg. ( E. duoplantaria Li, 2016 ), P. meghalayensis Chakrab. & N.P.Balakr. ( E. anthophilia Kawakita & Kato, 2016 ), P. microcarpus (Benth.) Müll.Arg. ( E. laeviclada Li, 2015 , E. macrocarpa Li, 2015 , E. tertiaria Li, 2015 ), P. niruri L. ( E. albifrons ( Stainton, 1859)) , P. sieboldianus T.Kuros ( E. corruptrix Kawakita & Kato, 2016 , E. obovatella Kawakita & Kato, 2016 ), Phyllanthus sp. ( E. albifrons ( Stainton, 1859) , E. scythropis Meyrick, 1930 ), P. subscandens var. subscandens ( E. corruptrix Kawakita & Kato, 2016 , E. jianfenglingina Li, 2016 , E. obovatella Kawakita & Kato, 2016 , P. urinaria subsp. Urinaria ( E. parasitica Kawakita & Kato, 2016 ), P. ussuriensis Rupr. & Maxim ( E. nudilingua Kawakita & Kato, 2016 ), P. wrightii (Benth.) Müll.Arg. ( E. angustisaccula Li, 2015 , E. camurella Li, 2015 ).

Distribution: Afrotropical Region: Mozambique; Namibia; South Africa: Gauteng, KwaZulu-Natal, Limpopo, Mpumalanga; Zimbabwe.

Australian Region: Australia: Queensland, New South Wales; French Polynesia: Marquesas Islands, Papua New Guinea.

Oriental Region: China: Fujian, Guangu , Guangdong, Guangxi, Hainan, Sichuan, Yunnan; India : Bihar, Karnataka, Kerala, Maharashtra, Meghalaya, Tamilnadu, West Bengal; Indonesia : Java; Japan: Amami Oshima , Iriomote Island , Ishigaki Island, Nansei Shoto (Ryūkyū Islands) , Okinawa Island , Tokunishima Island, Yonaguni Island; Moluccas (Buru) ; Malaysia : Selangor, Myanmar ; Sri Lanka; Thailand ; Viet Nam.

Palaearctic Region: China: Fijian, Gansu, Hebei, Heilongjiang, Hong Kong, Yianjin; Japan: Honshū, Kyūshū (Yaku-sima), Korea, Russian Federation: Primorje Region, Taiwan.

Species richness: World: 69 species; Australian Region: 20 species.

Australian species

Based on mitogenomic data and internal morphology of abdominal segment II we divide the Australian Epicephala species into three units:

1) E. albistriatella monophyletic group, consisting of the following species:

E. acrobaphes ( Turner, 1900)

E. albistriatella ( Turner, 1894)

E. dunkensis sp. nov.

E. philippa sp. nov.

E. xerocarpa sp. nov.

2) E. trigonophora monophyletic group, consisting of the following species:

E. breyniphaga , sp. nov.

E. sydnea sp. nov.

E. trigonophora ( Turner, 1900)

3) other species: E. colymbetella Meyrick, 1880 , E. acrocarpa Meyrick, 1927 , E. bathrobaphes Turner, 1947 , E. doddi sp. nov., E. eugonia Turner, 1913 , E. lomatographa Turner, 1913 , E. nephelodes Turner, 1913 , E. periplecta ( Diakonoff, 1955) , E. spinula Clarke, 1986 , E. spumosa Turner, 1947 , E. stephanephora Turner, 1923 and E. zalosticha Turner, 1940 . We have no sufficient data to group the species with unknown placement.

The species of E. albistriatella and E. trigonophora monophyletic groups cannot be diagnosed by external characters.

Key to the Australian Epicephala monophyletic groups based on internal morphological characters of abdomen and micromorphology of males sternal plate on sternum II with midline horizontal suture, transverse joint connecting tergal apodemes in sternum II is rudimental or with indentation; valvae in male genitalia strongly narrowed at mid part due to convex ventral margin................................................................................................... E. trigonophora group

- sternal plate on sternum II without midline horizontal suture, strongly sclerotised transverse joint, connecting tergal apodemes is present; valvae in male genitalia more or less equally wide along their entire length............. E. albistriatella group

Key to the Australian species of the E. albistriatella monophyletic group based on internal morphological characters of males

(1) ventral margin of valva with digitiform appendage, sacculus consists of a one-piece plate that acuminates towards the apex, apical part of sacculus triangular-shaped, saccus as a digitiform appendage........................ E. philippa sp. nov.

- ventral margin of valva without appendage, sacculus consists of two parts, apical part of sacculus rectangular or broadly rounded, saccus bulbed or acuminating toward anterior part.................................................... 2

(2) sacculus and valva more or less of equal width, apical part of sacculus broadly rounded, saccus almost round, bulb-shaped........................................................................................... E. acrobaphes

- sacculus almost twice broader than valva, apical part of sacculus rectangular, saccus slender, shaped as a digitiform appendage with acuminating apex................................................................ E. xerocarpa sp. nov.

Note: no data on male genitalia characters of E. albistriatella and E. dunkensis sp. nov.

Key to the Australian species of E. albistriatella monophyletic group based on internal morphological characters of females

(1) segment VII with a pair of leaf-shaped phyllary lamellas, apophyses posteriores and anteriores extend well beyond the anterior margin of corpus bursae, anterior margin of corpus bursae does not extend beyond segment V.............. E. acrobaphes

- segment VII with sclerotisations of other shapes, apophyses posteriores and anteriores do not extend beyond the anterior margin of corpus bursae, anterior margin of corpus bursae extends beyond segment V.................................... 2

(2) lamella post-vaginalis with a pair separate or fused plates that are rectangular posteriorly, signum triangular dentiform, signal area on corpus bursae squamous......................................................................... 3

- lamella post-vaginalis with U-shaped indentation and with narrow, gently rounded lateral sides, corpus bursae without signum or with one tiny star-shaped signum...................................................................... 4

(3) lamella post-vaginalis consists of two separate plates, posterior margin of which is unequal, with numerous indentations or processes, apophyses posteriores do not extend beyond the joint of ductus with corpus bursae, the anterior parts of apophyses anteriores extend ca. ¼ anteriad than the end of apophyses posteriores........................... E. xerocarpa sp. nov.

- lamella post-vaginalis is shaped as a fused plate with an equal rectangular posterior margin; apophyses posteriores and anteriores end almost at the same sector close to the mid of corpus bursae.................................. E. philippa sp. nov.

(4) ovipositor shortened reduced, triangular, ductus bursae ca. 2× shorter than corpus bursae, ends of apophyses posteriores and anteriores reach basal part of corpus bursae, corpus bursae without signum........................... E. albistriatella

- ovipositor long with truncate apex, ductus bursae ca. 2× longer than corpus bursae, ends of apophyses posteriores and anteriores reach the anterior sector of corpus bursae, corpus bursae with one small star-shaped signum................ E. dunkensis

Key to the Australian species of E. trigonophora monophyletic group based on internal morphological characters of males

(1) sacculus oval, almost as long as valva, with broadly rounded apical part, saccus narrow long, ca. as long as valva, aedeagus slender, longer than valva............................................................ E. breyniphaga sp. nov.

- sacculus with sharply pointed apex, ca. 2× shorter than valva, saccus short, ca. 2× shorter than valva, aedeagus thick in girth, ca. as long as valva, with truncate vesica.................................................... E. sydnea sp. nov.

Note: no data on male genitalia characters of E. trigonophora .

Key to the Australian species of E. trigonophora monophyletic group based on internal morphological characters of females

(1) lamella post-vaginalis with convex posterior margin, ductus bursae with a melanised ring, not sclerotised................................................................................................. E. breyniphaga sp. nov.

- lamella post-vaginalis with concave posterior margin, ductus bursae with a digitiform appendage, strongly sclerotised............................................................................................. E. trigonophora

Note: no data on female genitalia characters of E. sydnea sp. nov.

Type species: Epicephala colymbetella Meyrick, 1880

( Figs 288, 289)

“ Epic.[ephala] colymbetella , n. sp. ”—Meyrick, E., 1880. Proceedings of the Linnean Society of New South Wales 5(1): 169.

Epicephala colymbetella — Meyrick 1907: 53, 1928c: 489; Turner 1913: 175, 1940: 56; Fletcher 1929: 80; Viette 1949: 316; Kuznetzov 1979: 835; Clarke 1986: 343; Nielsen & Kumata 1996: 48; De Prins & De Prins 2005: 178; Hu et al. 2011: 447.

Synonym: Epicephala frugicola , n. sp. —Turner, A.J. 1913: 175–176. A junior subjective synonym of Epicephala colymbetella Meyrick, 1880 ; synonymized by A.J. Turner (1940: 56).

Type locality: [ Australia, New South Wales], Sydney.

Type specimens: Syntype 1(♂), in NHMUK (London) ; Syntype 2(♀), in NHMUK (London) .

Specimens examined: Syntype 1(♂): New South Wales, Sydney , 06 January 1978, Epicephala colymbetella , 7/12 Meyr., E. Meyrick det., in Meyrick Coll., BMNH(E) 1406774. Syntype 2(♀): New South Wales, Sydney, 23 December 1978, Epicephala colymbetella , 4/12 Meyr., E. Meyrick det., in Meyrick Coll., BMNH(E) 1406783, NHMUK (London) .

Lectotype designation: Hereby, we designate as the lectotype of the species Epicephala colymbetella Meyrick, 1880 the male specimen ( Fig. 288) with abdomen, belonging to the syntype series and carrying the following labels: [1] ‘Sydney/S.[outh] Australia‘/ 06 January 1878 (handwritten on dark beige paper), [2]’Meyrick Coll./B. M. 1938- 290’ (printed on white paper), [3] ‘Paralectotype’ (printed on white paper with blue ring on the outer margin’ [4] ‘ Epicephala / colymbetella /7/12 Meyr./E. Meyrick det./in Meyrick Coll.’ (three upper lines handwritten with black Indian ink, two lower lines printed on white paper), [6] ‘BMNH(E) 1406774’, in NHMUK (London).

Paralectotype ♀ ( Fig. 289): with abdomen: belonging to the syntype series and carrying the following labels: [1] ‘Sydney/S.[outh] Australia‘/ 23 December 1878 (handwritten on dark beige paper), [2]’Meyrick Coll./B. M. 1938- 290’ (printed on white paper), [3] ‘Paralectotype’ (printed on white paper with blue ring on the outer margin’ [4] ‘ Epicephala / colymbetella /7/12 Meyr./E. Meyrick det./in Meyrick Coll.’ (three upper lines handwritten with black Indian ink, two lower lines printed on white paper), [6] ‘BMNH(E) 1406783’, in NHMUK (London).

The lectotype is designated as part of our taxonomic work to enhance the stability of nomenclature (Declaration 44—Amendment of Article 74.7.3) with the purpose of delineating this species-group taxon Epicephala colymbetella Meyrick, 1880 . This designation will preserve stability in nomenclature ( ICZN Recommendation 74A). We gave the preference to the specimen indicated as ‘7/12 Meyr.’ in the Insect Collection of the Natural History Museum (London) which is with the abdomen, in good shape and curated by the former curator Dr. Jurate De Prins who provided it with the unique QR Code BMNH(E) 1406774 ( ICZN Recommendations 74B, C, D). The locality of the lectotype specimen is verified and included in the Global Taxonomic Database of Gracillariidae https://www. gracillariidae.net/species_by_code/EPICCOLY ( ICZN Recommendation 74E). The syntype specimen with the label data of the type locality and the QR Code BMNH(E) 1406783 is designated as the paralectotype ( ICZN Recommendation 74F).

Note: Four species are found under the same phenotype of Epicephala colymbetella ; however, two of them are phylogenetically distant though both of them belong to the genus Epicephala ( Fig. 637). The assignment of unverified specimens to one of the four species is impossible without detailed internal morphological characters of genitalia and/ mitogenomic data of the type specimens (lectotype ♂, and paralectotype ♀) of Epicephala colymbetella .

Unverified specimens examined: Queensland: Specimen 1: without abdomen, Millaa , 17.5117°S 145.6136°E, North , 24-09-1934, leg. nihil, ANIC Acc. no 31 075703. Specimen 2: without abdomen, Burpengary, 27.1615°S 152.9581°E, North, No date, leg. nihil. Specimen 3: idem locality and collecting data. Specimen 4: idem locality and collecting data. Specimen 5: without abdomen, Brisbane, 27.4705°S 153.0260°E, 20-04-1919, leg. nihil, ANIC Acc. no 31 075701. Specimen 8: without abdomen, Dunk Island , 17.9338°S 146.1437°E, North Queensland, DNA Voucher specimen, Sample ID: 11 ANIC-16209 , BOLD Proc. ID: ANICY209-11, 07-06-1928, leg. nihil, ANIC Acc. no 31 053780. Specimen 9: without abdomen, Tamborine Mountain , 27.9284°S 153.1845°E, 24-11-1940, leg. nihil, ANIC Acc. no 31 075702. Specimen 10: without abdomen, Dunk Island , 17.9338°S 146.1437°E, 06-06- 1928, leg. nihil. Specimen 11: Brisbane, 27.4705°S 153.0260°E, 15-04-1919, leg. nihil. Specimen 12: Dunk Island , 17.9338°S 146.1437°E, 06-06-1928, leg. nihil., ANIC (Canberra) Specimen 16: Brisbane, 27.4705°S 153.0260°E, as E. frugicola , no date, Goldfinch G.M. Specimen 19: idem collecting and locality data, February, Specimen 20: idem collecting and locality data, December. Specimen 21: idem collecting and locality data, Australian Museum (Sydney) .

New South Wales: Specimen 6: without abdomen, Gosford, 33.4267°S 151.3417°E, 21-11-1924, leg. nihil. Specimen 7: without abdomen, idem locality and collecting data. Specimen 13: without abdomen, Sydney , 33.8688°S 151.2093°E, Barcode of Life , DNA voucher specimen, Sample ID: 11 ANIC-16208 , BOLD Proc.ID: ANICY208- 11, 04-01-1933, leg. Goldfinch G.M., ANIC Acc. no 31 053779. Specimen 14: idem collecting and locality data. Specimen 15: without abdomen, idem locality and collecting data except the date 29-10-1932. Specimen 17: Sydney, 33.8688°S 151.2093°E, 25-10-2014, leg. Goldfinch G.M. Specimen 18(♀): idem collecting and locality data except the date 16-11-2014, Australian Museum (Sydney) .

Morphological diagnostic characterisation: Wingspan 8.2–8.6 mm; length of the forewing 3.7–3.9 mm ( Figs 288, 289). The main diagnostic set of characters for this species are found in micromorphology and in mitogenomics. This species is monophagous on Glochidion ferdinandii (Müll.Arg.) F.M. Bail ( Phyllanthaceae ).

Head: vertex shining white, smoothly covered by long piliform scales projected anteriorly, occiput with two lateral tufts of shining white, short piliform scales directed radially, frons more or less triangularly shaped with a loose bunch of piliform scales on labrum, maxillary palpus short with blunt apices, labial palpus slightly longer than the eye, with sharp apices; antenna slightly longer than forewing, unicolourous light ochreous grey with silver lustre, scape ca. as large as two flagellomeres, more or less equally broad at base and at apex carrying a tuft of long radially oriented scales.

Thorax ( Figs 288, 289): unicolourous white, tegula light ochreous grey, concolourous with the ground colour of forewing; forewing rather broad in comparison with other genera of Gracillariidae , equally wide along its entire length, with a white ornamental pattern on dorsal margin; costal margin from mid of forewing is decorated with very narrow, hardly visible oblique stripes and tiny spots, while the base of dorsal margin is snowy white and mid part of dorsal margin bears three thick white, oblique, not edged strigulae; sub-apical costal margin is marked by three oblique narrow stripes abruptly terminating at sub-apical area, narrow shining, edged with black scales fascia; apical spot small, oval, but clearly visible, present on light ochreous patch which is bordered by a small white spot at apex and a big triangular spot on tornus; apical line narrow, distinct, black, continuous; fringe line very short and present only at apex. The wing ornamentation is variable in thickness, edging, and obliqueness of stripes. The general pattern is that the dorsal margin is more decorated than the costal margin and usually carries three oblique clearly detectable strigulae. In E. colymbetella hind tibia bears a row of erected, sharply pointed, spiculose scales, a characteristic trait for Australian Ornixolinae genera.

Abdomen ( Figs 288, 289): tergites light ochreous, matte, without shine, last genital segments white, sternites dirty white with three dark brown oblique stripes.

Female and male genitalia: No data.

BOLD data: https://www.boldsystems.org/index.php/Taxbrowser_Taxonpage?taxon =colymbetella &searchTax=Se arch+Taxonomy

GenBank data: No data.

Mitogenomic data: No data.

Bionomics: Phyllanthaceae : Glochidion ferdinandii (Müll.Arg.) F.M. Bailey ( Turner 1913: 176) . This species is monophagous. One species of Australian Gracillariinae , Caloptilia xanthopharella Meyrick, 1880 , and two species of Australian Ornixolinae , Epicephala colymbetella and Diphtheroptila ochridorsellum , feed on the same host plant Glochidion ferdinandii . The diagnosis between them can be easily made by external morphological wing pattern characters.

Distribution: Australian Region: Australia: Queensland ( Meyrick 1907: 105), New South Wales ( Meyrick 1880: 169); French Polynesia: Marquesas Islands ( Meyrick 1928c: 504).