taxonID	type	description	language	source
2E392074FF8AFFE9E4E2D0A0FD13F9B9.taxon	description	Zoobank registration: u r n: l s i d: z o o b a n k. org: act: E 83 BF 881 - 52 D 1 - 4204 - A 205 - E 22 AE 305 EEA 9	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF8AFFE9E4E2D0A0FD13F9B9.taxon	type_taxon	Type species: Telesto dominula Plötz, 1884 (hereby designated).	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF8AFFE9E4E2D0A0FD13F9B9.taxon	diagnosis	Diagnosis Atkinsia differs from Dispar Waterhouse & Lyell, 1914, Pasma Waterhouse, 1932 and Anisynta Lower, 1911 by the following characters. The male genitalia of Atkinsia (Fig. 2) have the upper (dorsal) process of the valva rotated vertically downwards at 90 ° and projecting posteriorly well beyond the lower (ventral) process, whereas in Dispar, Pasma and Anisynta the dorsal process is relatively straight or with a slight downward curve in Dispar and Pasma. This unique shape of the processes in Atkinsia gives the valva a ‘ pliers’ - like appearance. The female genitalia of Atkinsia (Fig. 3) possess a unique‘horseshoe-shaped’ sclerotized structure that protrudes from the eighth sterna posterior to the lamella antevaginalis of the sterigma. This structure is absent in the three other genera, and it has not been found in any other species of Trapezitinae (A. F. Atkins, pers. comm., 2020). Although androconia are characters under sexual selection, and therefore not that useful for generic diagnosis, the male sex-brand of Atkinsia (Fig. 4 A) is highly unusual and possibly unique in the Trapezitinae. The sex-brand on the forewing extends as a broad, linear streak from cell CuA 1 to cell 1 A + 2 A, but it consists of two distinct parallel patches of sex-scales, the inner patch black and the outer, broader patch grey. Sex-brands are absent in Pasma and most species of Anisynta and, within the Anisynta group, are present only in Dispar and Anisynta monticolae (Olliff, 1890). In Dispar and Anisynta monticolae, however, the sex-brand consists of single linear streak of dark grey scales and is shorter in length, extending from vein CuA 1 to 1 A + 2 A. Both sexes of Atkinsia are generally larger than adults of the other three genera, and they also differ with regard to the colour pattern elements, particularly the markings on the underside of the hindwing, in which the silvery-white or cream postmedian spots may form a broad convex band that is joined to a prominent tornal spot, an arrangement not found in Anisynta, Dispar or Pasma. In addition, the pupal cap of Atkinsia (Fig. 5) has a long, narrow, sculptured projection that is absent in the other genera and which appears to be unique in the Trapezitinae. In Pasma, the projection of the pupal cap consists of a shorter, laterally spread, raised area (Atkins, 1988) whereas in Anisynta (Atkins, 1975 b) and Dispar (Braby, 2000), the head of the pupa is rounded anteriorly, rugose and lacks a prominent projection (similar to Trapezites); in Anisynta, the cap is divided into three rounded areas.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF8AFFE9E4E2D0A0FD13F9B9.taxon	discussion	Remarks Atkinsia is closely related to Anisynta, Dispar and Pasma. These four genera form a well-supported monophyletic group, with the following relationships according to our molecular results: Dispar + (Atkinsia + (Pasma + Anisynta )), which we refer to as the Anisynta group of trapezitine skippers. Thus, continued recognition of the species dominula in Anisynta renders that genus polyphyletic. The species dominula Plötz, 1884 has had a relatively unstable classification. Originally described in the genus Telesto Boisduval, 1832 by Plötz, Miskin (1891) placed dominula and Telesto drachmophora Meyrick, 1885 (a junior synonym of dominula) under Hesperilla. Meyrick & Lower (1902), however, maintained dominula under Telesto, noting the close similarity between it and drachmophora. However, Lower (1911) later assigned dominula to Hesperilla, noting that Telesto was preoccupied (i. e. a junior homonym of Telesto Lamouroux, 1812, a genus of soft corals in Clavulariidae), and cast doubt on the status of drachmophora. Subsequently, Waterhouse & Lyell (1914) placed dominula and Hesperilla monticolae Olliff, 1890 in the genus Motasingha. Although Lower (1911) introduced the concept of Anisynta a few years earlier, in 1911, dominula (and monticolae) were not transferred from Motasingha to that genus until 21 years later by Waterhouse (1932 b), who also provided a revised diagnosis for Anisynta, and he described the genus Pasma to accommodate the species Hesperilla tasmanica Miskin, 1889 and Hesperilla polysema Lower, 1908 [the concept of Pasma was subsequently revised by Atkins (1973)]. For the past 90 years, dominula has remained in Anisynta. We now place the taxon in a new genus that is monotypic. Waterhouse (1932 a, 1938) recognized and described several infraspecific taxa within dominula, although most of these were not recognized by Braby (2000) owing to the extent of variation. There is negligible variation in the male and female genitalia across the geographical range of the species, from northern New South Wales (Figs 2 A, B, 3 A) to Tasmania (Figs 2 C, D, 3 B). Thus, the nomenclature, new combinations and synonyms recognized for Atkinsia are as follows:	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF8AFFE9E4E2D0A0FD13F9B9.taxon	synonymic_list	Atkinsia dominula (Plötz, 1884) comb. nov. Atkinsia dominula dominula (Plötz, 1884) comb. nov. = Telesto drachmophora Meyrick, 1885 = Anisynta dominula draco Waterhouse, 1938 = Anisynta dominula dyris Waterhouse, 1938 Atkinsia dominula pria (Waterhouse, 1932) comb. nov.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF8AFFE9E4E2D0A0FD13F9B9.taxon	description	Shared characters of the Anisynta group include: (1) the origin of vein CuA 2 of the forewing nearer the end of the discal cell than the base (Fig. 4 A); (2) the origin of vein M 1 of the forewing well separated from R 5, with the intervening discocellular vein (i. e. between M 1 and R 5) inclined toward the termen; (3) the origin of vein CuA 1 of the hindwing closer to M 3 than to CuA 2, with the intervening discocellular vein (i. e. between M 3 and CuA 1) aligned at a similar angle to the middle and lower discocellulars (i. e. between M 1 and M 3) such that the end of the discal cell is broadly truncate (Fig. 4 B); and (4) the hind tibia with two pairs of spurs. Vein 1 A + 2 A of the forewing is slightly distorted in the middle in Dispar and Atkinsia, but it is relatively straight in Pasma and Anisynta. The club of the antenna is evenly curved, with the apiculus short and pointed in Dispar and short and blunt in Atkinsia and Anisynta, whereas in Pasma it is sharply bent before middle, with the apiculus long and blunt. The male genitalia of all four genera are unusual in that the valvae are deeply divided apically into two long processes or ‘ arms’, which are heavily sclerotized, with short spines at their apices (Atkins, 1973, 1994; Sands & Sands, 2017). In all genera, the apical portion of the lower (ventral) process is rotated vertically upward at 90 °, or 135 ° in Dispar, and slightly inwards in Atkinsia. The uncus is similar in profile in Anisynta, Atkinsia and Pasma in that the apex is broadly truncate, whereas in Dispar the uncus is long and narrowly tapered to a point and with a slight downward curve. Members of the Anisynta group occur in temperate grassland, open woodland, woodland, open forest or tall open forest, and all use Poaceae as their larval food plant families. Dispar has occasionally also been recorded on Cyperaceae and Asparagaceae.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF8AFFE9E4E2D0A0FD13F9B9.taxon	etymology	Etymology The new genus is named in honour of Mr Andrew Atkins for his substantial and pioneering research on the taxonomy and biology of Australian butterflies, particularly the Trapezitinae, spanning> 40 years.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF88FFE7E52AD7BCFE95F902.taxon	type_taxon	Type species: Hesperilla ornata Leach, 1814 (by original designation).	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF88FFE7E52AD7BCFE95F902.taxon	discussion	Remarks Braby (2010) noted that the genera Hesperilla Hewitson, 1868, Motasingha Watson, 1893 and Oreisplanus Waterhouse & Lyell, 1914 are closely related and probably form a monophyletic group. Currently, 14 species are assigned to Hesperilla, whereas Motasingha and Oreisplanus each contain two species: Motasingha dirphia (Hewitson, 1868) and Motasingha trimaculata (Tepper, 1882) in Motasingha (Moulds & Atkins, 1986), and Oreisplanus munionga (Olliff, 1890) and Oreisplanus perornatus (Kirby, 1893) in Oreisplanus (Braby, 2000). Atkins (1984) and Moulds & Atkins (1986) first demonstrated that Motasingha is most closely related to Hesperilla, indicating that it differed chiefly by the form of the apiculus of the antennal club, which is long, stout and blunt rather than long and pointed. However, like Hesperilla, the club is bent before the middle. Other ‘ distinguishing’ morphological characters, such as the wing venation, relative length of the third segment of the labial palpus, number of spurs on the hind tibia, shape of the linear patch of sex-scales (androconia), male and female genitalia, and morphology of the larva (which is semi-translucent) and pupa (which is long and cylindrical, with a strongly produced and heavily sclerotized pupal cap) are otherwise similar to Hesperilla. Indeed, Atkins (1984) noted the strong similarity between Motasingha, which at that time was monotypic, containing the single species Motasingha dirphia, and the Hesperilla idothea (Miskin, 1889) species group of Hesperilla (i. e. Hesperilla donnysa, Hesperilla flavescens, Hesperilla idothea and Hesperilla mastersi) based on comparative illustrations of the wing venation, male androconia and female genitalia. Braby (2000) noted that Oreisplanus is closely allied to Hesperilla but is distinguished by the absence of male androconia (forewing sex-brand) and the form of the club of the antenna, which is evenly curved, with the apiculus short and blunt. Absence of a character does not diagnose a lineage and, moreover, androconia are likely to be under strong sexual selection. Other characters used to diagnose Oreisplanus are shared with Hesperilla, such as the wing venation, labial palpus, hind tibia, and form of the pupal cap, which is strongly produced and heavily sclerotized, in addition to the abdominal segments of the pupa, which, like Motasingha, contain long stiff setae that, in addition to the cremaster, provide support within the silken shelter. Indeed, A. F. Atkins (pers. comm.) considers the Hesperilla ornata (Leach, 1814) species group (i. e. Hesperilla ornata, Hesperilla picta, Hesperilla hopsoni and Hesperilla crypsargyra) to be more closely related to Oreisplanus than to other species of Hesperilla (Braby, 2010), a phylogenetic pattern which is concordant with our findings. This pattern was also partly evident in an earlier phylogenetic study by Warren et al. (2008), who showed that Oreisplanus perornatus was sister to Hesperilla ornata and that this pair was sister to Hesperilla donnysa, rendering Hesperilla paraphyletic. In our phylogenetic analysis, the genus Hesperilla was paraphyletic by inclusion of both Motasingha and Oreisplanus. Motasingha was recovered as sister to a clade that included the Hesperilla idothea species group (Hesperilla donnysa, Hesperilla flavescens, Hesperilla idothea and Hesperilla mastersi) and the Hesperilla malindeva species group (i. e. Hesperilla malindeva, Hesperilla sarnia, Hesperilla sexguttata, Hesperilla furva and Hesperilla crypsigramma) (Atkins, 1978); this clade, in turn, was sister to a clade that included the remaining species of Hesperilla. Oreisplanus was monophyletic with low branch support, nested within a clade comprising the Hesperilla ornata species group (i. e. Hesperilla ornata, Hesperilla picta, Hesperilla crypsargyra and Hesperilla hopsoni). This clade formed a monophyletic group sister to Hesperilla chrysotricha. Given the well-supported phylogenetic relationships among the three genera uncovered in the present study, and the lack of clear synapomorphies by which to distinguish Motasingha and Oreisplanus, we synonymize Motasingha and Oreisplanus under Hesperilla s. l., as foreshadowed by Braby (2010) more than a decade ago. Thus, according to our phylogeny, the nomenclature, new combinations and synonyms recognized for the revised 18 species of Hesperilla s. l. and their constituent subspecies are as follows:	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF88FFE7E52AD7BCFE95F902.taxon	synonymic_list	Hesperilla chrysotricha (Meyrick & Lower, 1902) Hesperilla chrysotricha chrysotricha (Meyrick & Lower, 1902) Hesperilla chrysotricha cyclospila (Meyrick & Lower, 1902) = Hesperilla chrysotricha leucospila Waterhouse, 1927 = Hesperilla chrysotricha plebeia Waterhouse, 1927 = Hesperilla chrysotricha leucosia Waterhouse, 1938 = Hesperilla chrysotricha lunawanna L. E. Couchman, 1949 = Hesperilla chrysotricha naua L. E. Couchman, 1949 Hesperilla ornata (Leach, 1814) Hesperilla ornata ornata (Leach, 1814) Hesperilla ornata monotherma (Lower, 1907) Hesperilla picta (Leach, 1814) Hesperilla perornatus Kirby, 1893 Hesperilla munionga Olliff, 1890 Hesperilla munionga munionga Olliff, 1890 Hesperilla munionga larana (Couchman, 1962) comb. nov. Hesperilla crypsargyra (Meyrick, 1888) Hesperilla crypsargyra crypsargyra (Meyrick, 1888) = Hesperilla crypsargyra lesouefi Tindale, 1953 Hesperilla crypsargyra binna Johnson & Wilson, 2005 Hesperilla hopsoni Waterhouse, 1927 Hesperilla dirphia Hewitson, 1868 Hesperilla trimaculata Tepper, 1882 Hesperilla trimaculata trimaculata Tepper, 1882 = Hesperilla quadrimaculata Tepper, 1882 Hesperilla trimaculata dilata (Waterhouse, 1932) comb. nov. = Motasingha dirphia dea Waterhouse, 1932 Hesperilla trimaculata occidentalis Moulds & Atkins, 1986 comb. nov. Hesperilla donnysa Hewitson, 1868 Hesperilla donnysa donnysa Hewitson, 1868 = Hesperilla donnysa diluta Waterhouse, 1932 = Hesperilla donnysa delos Waterhouse, 1941 = Hesperilla donnysa icaria Waterhouse, 1941 = Hesperilla donnysa patmos Waterhouse, 1941 = Hesperilla donnysa samos Waterhouse, 1941 Hesperilla donnysa aurantia Waterhouse, 1927 Hesperilla donnysa galena Waterhouse, 1927 Hesperilla donnysa albina Waterhouse, 1932 Hesperilla flavescens Waterhouse, 1927 = Hesperilla donnysa flavia Waterhouse, 1941 Hesperilla idothea (Miskin, 1889) Hesperilla idothea idothea (Miskin, 1889) = Telesto dispar Kirby, 1893 Hesperilla idothea clara Waterhouse, 1932 Hesperilla mastersi Waterhouse, 1900 = Hesperilla mastersi marakupa L. E. Couchman, 1965 Hesperilla malindeva Lower, 1911 = Hesperilla malindeva dagoomba Johnson & Valentine, 1994 Hesperilla sarnia Atkins, 1978 Hesperilla sexguttata Herrich-Schäffer, 1869 = Toxidia sexguttata sela Waterhouse, 1932 Hesperilla furva Sands & Kerr, 1973 Hesperilla crypsigramma (Meyrick & Lower, 1902)	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF88FFE7E52AD7BCFE95F902.taxon	discussion	Larvae of Hesperilla specialize on Cyperaceae, especially the genus Gahnia, with one species (H e s p e r i l l a t r i m a c u l a t a) a l s o f e e d i n g o n Haemodoraceae.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF86FFE4E73DD1C7FD3CFC90.taxon	type_taxon	Type species: Toxidia thyrrhus Mabille, 1891 (by monotypy).	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF86FFE4E73DD1C7FD3CFC90.taxon	discussion	Remarks Braby (2010: p. 26) remarked ‘ The genus Toxidia comprises a divergent assemblage of taxa [currently nine species] that have probably been grouped together more for convenience than on phylogenetic grounds. ’ Notably, he pointed out that the genus Neohesperilla on available evidence is poorly defined; the known morphological characters do not distinguish it from Toxidia (see Braby, 2000). Moreover, the morphology of the larva and pupa, general biology and larval food plants of Neohesperilla (Johnson et al., 1994) are similar to those of Toxidia (Crosby, 1973; Atkins, 1988; Wood & Muller, 2003; Jenkinson, 2009). For example, in both taxa the head of the final instar larva is black and finely pitted or rugose, and the head of the pupa is sclerotized and rounded, lacking a prominent projection. Currently, four species are referred to Neohesperilla, but phylogenetic analysis of this genus has, until now, been lacking to establish its systematic relationship and determine whether Neohesperilla should be maintained or synonymized with Toxidia. In the phylogeny generated in the present study, Toxidia was paraphyletic and polyphyletic with regard to Neohesperilla and Signeta, respectively. Although Neohesperilla comprised a monophyletic lineage, it was embedded within a larger clade that consisted of six other species of Toxidia, while the two species of Signeta formed a monophyletic group with two other species of Toxidia (Toxidia peron and Toxidia melania) that was sister to the remaining species of Toxidia plus Neohesperilla. Given the strong phylogenetic relationships uncovered in the present work, previous morphological studies, and the lack of clear synapomorphies by which to distinguish Neohesperilla from Toxidia, we synonymize Neohesperilla under Toxidia. The issue of polyphyly with regard to Signeta is dealt with below, in which we remove two species from Toxidia. Thus, the nomenclature, new combinations and synonyms recognized for the revised 11 species of Toxidia and their constituent subspecies are as follows:	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF86FFE4E73DD1C7FD3CFC90.taxon	synonymic_list	Toxidia arfakensis (Joicey & Talbot, 1917) Toxidia inornata (Butler, 1883) Toxidia inornata inornata (Butler, 1883) = Hesperia maykora Plötz, 1885 = Telesto uniformis Swinhoe, 1905 Toxidia inornata sekara (Plötz, 1885) = Toxidia damora Fruhstorfer, 1911 Toxidia inornata anga Evans, 1949 Toxidia andersoni (Kirby, 1893) Toxidia rietmanni (Semper, [1879]) Toxidia rietmanni rietmanni (Semper, [1879]) = Telesto extranea Plötz, 1884 = Telesto leucostigma Meyrick & Lower, 1902 Toxidia rietmanni parasema (Lower, 1908) Toxidia doubledayi (C. Felder, 1862) = Telesto leachii C. Felder, 1862 = Carystus vallio Mabille, 1883 Toxidia thyrrhus Mabille, 1891 = Telesto bathrophora Meyrick & Lower, 1902 Toxidia parvula (Plötz, 1884) = Hesperilla humilis Miskin, 1889 = Hesperilla ismene Anderson & Spry, 1894 Toxidia xanthomera (Meyrick & Lower, 1902) comb. nov. Toxidia senta (Miskin, 1891) comb. nov. Toxidia xiphiphora (Lower, 1911) comb. nov. Toxidia crocea (Miskin, 1889) comb. nov. = Hesperilla satulla Mabille, 1891 The larvae of Toxidia feed exclusively on Poaceae.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF85FFE4E497D2C2FBA5FAC5.taxon	type_taxon	Type species: Timoconia thiele Strand, 1909 (by original designation).	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF85FFE4E497D2C2FBA5FAC5.taxon	discussion	Remarks Signeta currently contains two species: Signeta flammeata (Butler, 1882) and Signeta tymbophora (Meyrick & Lower, 1902). Signeta is distinguished from Toxidia, to which it is closely allied, by differences in the shape of vein 1 A + 2 A in the male forewing, which is sharply distorted (i. e. bowed toward the dorsum) near the base, and the presence of a large ovoid central black patch on the upperside of the forewing in the male (Waterhouse & Lyell, 1914; Atkins et al., 1991; Braby, 2000). However, evidence from the morphology of the immature stages (Jenkinson, 2018), male genitalia and adult stage morphology (A. F. Atkins, pers. comm.) indicates that Toxidia peron (Latreille, 1824) is more closely allied to Signeta than to other members of Toxidia. Moreover, in a molecular phylogenetic analysis that included a limited subset of trapezitine taxa (Warren et al., 2008, 2009), Toxidia peron emerged sister to S. flammeata rather than to Toxidia doubledayi (C. Felder, 1862) (the other exemplar of the genus studied). Atkins (2012, 2017) hypothesized further on the putative close relationship between Toxidia peron and Signeta, and he placed Toxidia peron informally under Signeta, although he did not comment on the status of other members of Toxidia, such as Toxidia melania (Waterhouse, 1903), for which the biology and life history remain poorly known. As noted above, the genus Toxidia was polyphyletic in our analysis, with two species (Toxidia peron and Toxidia melania) showing a closer relationship to Signeta than to the other species of Toxidia. The logical course of action would be to change the composition of these genera by transferring Toxidia peron and Toxidia melania across to Signeta to maintain monophyly; otherwise, Signeta would have to be synonymized under Toxidia. However, since Timoconia thielei Strand, 1909 (the type species of Timoconia) is a junior synonym of Hesperia peron Latreille, 1824, Timoconia Strand, 1909 is a valid and available name for this clade of four taxa and therefore becomes the senior synonym of Signeta Waterhouse & Lyell, 1914 because it was published five years earlier. Thus, the nomenclature and new combinations recognized for the four revised species of Timoconia are as follows:	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF85FFE4E497D2C2FBA5FAC5.taxon	synonymic_list	Timoconia peron (Latreille, 1824) comb. nov. = Telesto kochii Felder, C., 1862 = Hesperilla doclea Hewitson, 1868 = Telesto arsenia Plötz, 1884 = Timoconia thielei Strand, 1909 Timoconia melania (Waterhouse, 1903) comb. nov. Timoconia flammeata (Butler, 1882) comb. nov. = Telesto eclipsis Butler, 1882 = Hesperilla atromacula Miskin, 1889 Timoconia tymbophora (Meyrick & Lower, 1902) comb. nov.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF85FFE4E497D2C2FBA5FAC5.taxon	discussion	Larvae of Timoconia exploit a wide range of monocots, including Asparagaceae, Cyperaceae, Flagellariaceae, Phormiaceae and Poaceae. Flagellariaceae were recently recorded as food plants for T. melania (E. A. Petrie, unpublished data).	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
2E392074FF85FFE5E7EDD449FDA9FD83.taxon	discussion	The genus Prada contains three species endemic to mainland New Guinea (Parsons, 1998). Evans (1949) considered Prada to be closely related to the monotypic Tiacellia Evans, 1949 (also from New Guinea), and he classified this pair within the Plastingia group of the subfamily Hesperiinae, although he noted a possible link with the Trapezitinae. Parsons (1998) pointed out that the male genitalia and hindwing venation of Prada rothschildi (Evans, 1928) was similar to that of Rachelia extrusa (C. & R. Felder, 1867). Subsequently, Atkins (2017) transferred Prada and Tiacellia to the Trapezitinae and suggested that Prada and Tiacellia might be closely related to Trapezites and Rachelia based on study of male genitalia and wing venation. Cong et al. (2019) reached a similar conclusion regarding Prada based on molecular data of a limited set of Trapezitinae (10 species representing 10 genera), in which Prada rothschildi was sister to Trapezites symmomus Hübner, 1823 in most of their analyses, rather than to other members of the Hesperiinae in which Prada has traditionally been placed. In our comprehensive molecular phylogeny of Trapezitinae, Prada was placed as sister to Trapezites, thus supporting previous views that this genus belongs in the Trapezitinae and not in Hesperiinae. Our analysis also supports, in part, the hypothesis of Atkins (2017) of a close relationship between Prada and Trapezites. Tiacellia, however, was not recovered within the Trapezitinae, falling well outside this subfamily (Fig. 1). Larval food plants remain unknown.	en	Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A., Kawahara, Akito Y. (2022): Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421, DOI: 10.1093/zoolinnean/zlab086
