Lipomelia, Warren, 1893

Sihvonen, Pasi, 2005, Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae), Zoological Journal of the Linnean Society 143 (4), pp. 473-530 : 505-509

THE LIPOMELIA , PROBLEPSIS AND SCOPULA

GROUP OF GENERA

This group of genera is characterized by one synapomorphy ( Figs 133-135 ): the fully developed cerata (79: 2, six occurrences, six reversals). Within the group, the relationships are not resolved. Lipomelia and Problepsis form a compact group, Somatina irregularis and Scopula haemaleata are incertae sedis, and externally heterogeneous Scopula is characterized by a combination of features.

Comments on systematics

See the Dithalama , Isoplenodia , Somatina and Zythos group of genera.

LIPOMELIA WARREN, 1893

( FIGS 6 , 35 , 60 , APPENDIX)

Diagnosis: monophyly supported by ten synapomorphies: forewing costa is wide and grey (11: 2, two occurrences); presence of pale markings on anterior side of hindwing discal spot (23: 1, two occurrences); subterminal shade line and terminal line are touching each other at fore- and hindwings (42: 1, two occurrences, Fig. 6 ); absence of male hindleg pretarsus arolium (68: 0, six reversals); absence of male hindleg pretarsus pulvillus (69: 1, three occurrences); sacculi of valva are asymmetrical (105: 1, seven occurrences); valvuli of valva are asymmetrical (108: 1, seven occurrences); vinculum is asymmetrical (113: 1, six occurrences); anterior margin of juxta is without wing-like processes (116: 0, nine occurrences); presence of parallel folds on vesica (132: 1, seven occurrences).

Imago: wingspan c. 25 mm. Head smooth-scaled; male antennae fasciculate, sensilla in single rows. Pilifer present. Wings brown, base striated, dark area between transverse posterior line and subterminal shade line at forewings; transverse lines cryptic, subterminal shade line and terminal line touching each other at fore- and hindwings ( Fig. 6 ); discal spots white, with iridescent scales, surface smooth; pale markings on anterior side of hindwing discal spots; terminal line discontinuous, equally wide; underside fuscous. Forewings with one areole; R 5 stalked with R 2 –R 4. Outer and inner margin hair pencil of male hindleg tibia well developed; distal spurs of hindleg tibia absent; tarsomeres short, segments 4–5 fused; two claws; arolium and pulvillus absent. Female hindleg tibia with 2 + 2 spurs ( Prout, 1920 -41).

Male abdomen and genitalia: round pouch and anterolateral extensions on male 2nd sternite present. One membranous pouch between male 7th and 8th sternite. Male 8th sternite anterior margin elongated medially; cerata present, coalescent at base ( Fig. 60 ); mappa absent; membranous posterolateral appendices present. Male 8th tergite posterior margin concave. Socii short, setose, separate. Ventral margin of tegumen straight. Valvae asymmetrical, valvula and sacculus separate, ventral margin of left sacculus bifurcate. Transtilla present. Anterior margin of juxta with wing-like processes absent. Aedeagus curved dorsally; plate-shaped sclerotization at proximal end of ductus ejaculatorius; cornutus absent; vesica simple, large lateral diverticulum at base with sclerotized parallel folds, small diverticulum dorsally.

Female genitalia: unknown.

Distribution and species diversity: from India to Taiwan. One species.

Immature stages: unknown.

Biology: unknown.

Comments on systematics: when Warren (1893) described Lipomelia , he did not discuss its systematic position. Prout (1920 -41) considered it a close relative of Scopula on the basis of one areole on the forewing and male genital characters, although he noted that the facies of L. subusta resembles that of Zythos spp. In the present study, Lipomelia is associated with Problepsis on the basis of forewing iridescent scale characters. One of these characters, i.e. the structure of iridescent scales being glossy and smooth (17: 2, Figs 102 , 134 ) is a unique synapomorphy of this clade. I have chosen not to synonymize Lipomelia with Problepsis , however, because the former lacks several characteristic features of the latter, e.g. ocellate discal spots on both pairs of wings (cf. Figs 6, 7 ), fused socii lying ventral to tuba analis (cf. Figs 35, 36 , 117 ), and male 8th sternite anterior margin not being trifurcate and blunt (cf. Figs 60, 61 ). Male genitalia of Lipomelia , including 8th sternite, were found to be structurally similar to those of Scopula .

PROBLEPSIS LEDERER, 1853

( FIGS 7 , 27 , 36 , 61 , 84 , 102, 105, 106, 108 , 115, 117 , 128 , APPENDIX)

Diagnosis: monophyly supported by five synapomorphies: discal spot of forewing is ocellate and round (20: 3, three occurrences, Fig. 7 ); discal spot of hindwing is ocellate (22: 1, three occurrences); socii are fused at apex, lying ventral of tuba analis (92: 2, two occurrences, Fig. 117 ); ventral margin of tegumen is dentate (100: 1, two occurrences); anterior margin of juxta is with wing-like processes, apex is fused to sacculus of valva (116: 2, five occurrences).

Imago: wingspan 25-50 mm. Male antennae fasciculate; sensilla in single or multiple rows, proximal row elongated in many species. Female antennae fasciculate, simple; sensilla in single rows. Pilifer present. Wings pale to white, greyish in few species; ocellated discal spots ( Fig. 7 ) distinct; smooth, glossy, iridescent scales often present in discal spots or scattered throughout wings ( Figs 102, 105, 106 ); transverse median line often wide, indistinct; subterminal shade line discontinuous, spotted; terminal line continuous or discontinuous, equally wide or wider at vein ends. Patterns not repeated or weakly expressed on underside. Forewings often with one areole, sometimes two; R 5 from proximal or distal areole, stalked with R 2 –R 4. Outer margin hair pencil of male hindleg tibia present or absent; inner margin hair pencil often moderate, sometimes massive or absent; tarsomeres often reduced in length, 5-segmented or segments 4–5 fused; claws either two in number or absent. Female hind tibia with 2 + 2 spurs.

Male abdomen and genitalia: pouch on male 2nd sternite round, posterior margin often invaginated; anterolateral extensions present or absent. One or two membranous pouches between male 7th and 8th sternite. Male 8th sternite elongated; anterior margin often trifurcate, blunt or sometimes bifurcate; cerata absent, rudimentary or fully developed, often fused to mappa; mappa present, bare; membranous posterolateral appendices present or absent. Male 8th tergite posterior margin concave. Socii fused at apex, ventral of tuba analis ( Fig. 117 ), setose, sometimes with two lateral appendices. Ventral margin of tegumen sometimes unmodified, dentate in most species. Valvula and sacculus of valva separate, symmetrical, long, tapering towards apex, acute; medial margin of sacculus rarely upturned. Transtilla variably sclerotized, often invaginated medially. Anterior margin of juxta with wing-like processes, often anterior margin sclerotized only, apex fused to sacculus of valva, sometimes free. Vinculum enlarged , symmetrical, anterior margin often concave. Aedeagus often slightly bent, apex with one or several teeth; vesica large, complex, variably sclerotized, rarely with distinct cornuti; several diverticula, sometimes long and prominent.

Female genitalia: lamella postvaginalis sclerotized in many species; lamella antevaginalis often absent, sometimes weakly developed, rarely flap-like, flexible. Ostium bursae often sclerotized; ductus bursae sclerotized, straight, wide ( Fig. 84 ) in most species. Ductus seminalis arises from ductus bursae or corpus bursae neck. Corpus bursae ovoid, membranous sac. Signum absent sometimes, usually an ovoid patch of separate or fused spines, small compared to size of corpus bursae.

Distribution and species diversity: most species occur from sub-Saharan Africa to Indonesia and Australia. Only a few reach the Palaearctic region. Fifty-one species and several subspecies.

Immature stages: the only larval description is based on a species from south India, whose identity is uncertain. It is possibly P. deliaria Guenée (Bell in Holloway, 1997). The Japanese species illustrated in Sugi (1987) rest at 45 ∞ from the substrate, usually in the mid-rib of a leaf. Based on limited Japanese material ( Nakamura, 1994), the pupal cremaster has one pair of setae only. Pupation takes place in a loose cocoon incorporating particles of earth, usually on the soil surface.

Biology: all host-plant records relate to Oleaceae ( Holloway, 1997) . These include Olea ( Holloway, 1997) , Ligustrum ( Sugi, 1987) and Jasminum ( Singh, 1957) . Many species are from lowland forest, including cultivated areas and dry heath, but Holloway (1997) reports species taken at heights of almost 1800 m in Borneo. In north-east China, P. phoebearia Erschov and P. plagiata (Butler) were taken from a lamp on a warm, south-facing sandy grove (pers. observ.).

Comments on systematics: traditionally, Problepsis has been delimited to include species that have one forewing areole, large ocellated discal spots with iridescent scales and no spurs on male hind legs. Its closest relatives were assumed to be Antitrygodes , Scopula and Somatina ( Sterneck, 1941; Prout, 1934- 39; Holloway, 1997). In the present study, the clade comprising Lipomelia + Problepsis was found to be the closest relative of Scopula , in addition to two taxa incertae sedis ( Fig. 133 ). Ocellated discal spots, whether round or semiround (20: 3 or 20: 4), were found to be a unique synapomorphy of Problepsis . Number of forewing areoles was shown to be a homoplastic character (Ch. 48), and although the majority of Problepsis species have one, two are also found, e.g. in P. centrophora (Prout) . Absence of male hindleg spurs was found to be a plesiomorphic feature in Scopulini (62: 0). Fused socii of male genitalia (92: 2), which lie ventral to tuba analis, was found to be a diagnostic character delimiting the genus; it is present in all studied Problepsis taxa, and found outside of it in Antitrygodes cuneilinea (Walker) only. It is possible that a few Problepsis species are misplaced in Somatina , see Appendix. See also Discussion under Lipomelia .

SCOPULA SCHRANK, 1802

( FIGS 8–26, 29 , 37–54 , 62–79 , 85–100, 104, 107 , 110–113, 116 , 118, 122, 123, 126, 127, 130 , APPENDIX)

Diagnosis: monophyly supported by one synapomorphy: absence of posterolateral appendices on male 8th sternite (86: 1, five occurrences).

Imago: wingspan 11–70 mm, usually 25-35 mm. Male antennae fasciculate; sensilla in single or multiple rows, proximal row sometimes elongated markedly; rarely bipectinate. Female antennae usually fasciculate, simple, occasionally sensilla arranged in single or even multiple rows. Pilifer present. Wings very variable, no single character can be used to define this group; almost all colours are found, but usually straw or grey-brown are dominant; often irrorated, sometimes unicolorous; iridescent scales absent; three wavy transverse lines running from forewing to hindwing are common, dentate in many species, reduced to series of spots in other or absent; sometimes two dark marks on forewings medial to transverse posterior line and another dark mark at forewing termen; few species with reticulate pattern; few with wide dark areas on outer third of wings; terminal line at forewings usually discontinuous, equally wide; smoothly curved around forewing apex occasionally or absent. Discal spot usually dark, small; cryptic in some, rarely absent; fringe usually unicolorous, occasionally chequered. Patterns usually repeated on underside, weak, sometimes absent. Forewings usually with one areole, sometimes two; R 5 usually from proximal areole, either stalked with R 2 –R 4 or free, sometimes from distal areole, stalked with R 2 –R 4 or free or even from M 1. Hindwing Sc + R 1 and Rs usually stalked for short distance, then divergent or sometimes connected by R 1. Outer and inner margin hair pencil of male hindleg tibia either present or absent; sometimes pronounced; if absent, replaced with two spurs at distal end in many species, occasionally one or absent; tarsomeres usually normally developed, sometimes reduced in length, 5-segmented; rarely segments 1–5 or 2–5 fused; two claws, absent in few groups. Female hindleg tibia with 2 + 2 spurs, sometimes only two distally.

Male abdomen and genitalia: pouch on male 2nd sternite round, posterior margin occasionally invaginated or rarely elongated, surface smooth or with fine decorations, one- or occasionally two-chambered; occasionally laterally elongated; with or without pointing setae, sometimes absent; anterolateral extensions present or absent. Sometimes tufts of setae on male 6th and 7th segments laterally. Usually one membranous pouch between male 7th and 8th sternite, rarely two or absent. Male 8th sternite anterior margin often medially elongated, sometimes cup-like or bifurcate, rarely trifurcate; cerata usually present, asymmetrical, free ( Fig. 62 ), with apical setae, or rarely fused to mappa, sometimes seen as bifurcate, rudimentary or absent; mappa usually present, bare ( Fig. 76 ), sometimes setose or posterior margin separate, rarely absent; posterolateral appendices usually absent, pronounced occasionally; posterior margin of male 8th sternite not sclerotized. Male 8th tergite posterior margin usually concave with two round lobes ( Figs 118, 122 ) or rarely with sharp projections; laterally constricted in few species. Socii size and shape variable, typically separate at apex, setose, well developed, slightly upturned, occasionally reduced to tuft of setae, rarely massive, or in few instances fused at apex, dorsal of tuba analis. Tegumen ventral margin usually unmodified, rarely with lateral nondentate projections. Transtilla present, usually U-shaped, occasionally W-shaped; weakly sclerotized. Sacculus and valvula of valva separate, partially fused in few instances; size and shape very variable; symmetrical in most cases but pronounced asymmetry not unusual; valvula usually soft, blunt-ended (e.g. Fig. 48 ), bent ventrally, setose, blunt-ended, often with apical spines; sacculus usually more sclerotized, tapering towards apex, sometimes acute or dark; bifurcate in some species; occasionally rudimentary; various projections may arise from lateral and ventral margins. Anterior margin of juxta typically with wing-like projections ( Fig. 52 ), sometimes absent; apex usually free, but both or one extensions fused to sacculus in few species; often asymmetrical, size and shape variable; juxta often tube-shaped, well sclerotized with cup-like extension ventrally. Vinculum enlarged , margin typically convex, rarely concave; symmetrical, sometimes asymmetrical; cup-shaped and turned ventrally in few species. Aedeagus size and shape variable, in many instances narrow and straight or wide and slightly bent; caecum usually a continuation of aedeagus, enlarged occasionally; sharp projections at apex (carina) sometimes; apex sharp or blunt; vesica size and shape variable; often simple sac, with small diverticula and plate-shaped sclerotization at proximal end of ductus ejaculatorius; or large, complex, with elongated, twisted diverticula, variably sclerotized; distinct cornuti usual but more often sclerotizations are less definitive; pronounced plate-shaped sclerotizations at distal end of vesica occasionally; all instances between extremities occur.

Female genitalia: papillae anales often round, ventral margin upturned occasionally. Lamella postvaginalis usually weakly developed, irregular plate or made of parallel folds or absent. Lamella antevaginalis usually sclerotized, flap-like, semiround ( Fig. 99 ), posterior margin invaginated in few species; horseshoe-shaped in others, laterally wider; asymmetry usual; normally bare but occasionally heavily setose. Ostium bursae usually sclerotized, with lateral extensions, reaching proximal part of ductus bursae, dorsally membranous. Ductus bursae variable in length, width, shape and amount of sclerotization; membranous and coiled in many cases, or short and wide, or indistinct. Ductus seminalis arises from ductus bursae, corpus bursae neck or corpus bursae. Corpus bursae elongated sac, usually smooth but heavily wrinkled in others. Signum usually present as an elongated patch of separate spines ( Fig. 130 ), sometimes two patches, which may be fused medially along axis, sometimes spines fused forming long and narrow band. Usually corpus bursae with spines on inner surface, length and density variable.

Distribution and species diversity: virtually ubiquitous. The genus has successfully colonized even the remotest islands in Polynesia and New Zealand ( Holloway, 1997; Dugdale, 1988). About 800 described species and numerous subspecies. See ‘Comments on systematics’ under Somatina .

Immature stages: material is scarce in museums and pertinent literature is scattered, often dealing with single species. Too little is known to make comprehensive conclusions. The most detailed descriptions can be found in McGuffin (1967: 12), who dealt with Canadian species. Their application to the worldfauna may be questionable, but they provide a good starting point: ‘Egg round, somewhat longer than wide, usually larger at one end than at other (figs 77, 84c) with longitudinal ridges and cross striae; white, cream coloured, or light green; turning pink, scarlet, brown, or black; attached to leaves of food plant or laid loosely on ground near food plant.’ The illustrations of eggs of Palaearctic Scopula species fit the description well ( Sannino & Balbiani, 1984; Sannino & Espinosa, 1999). The caterpillar is usually long, slender, green or brown in colour, with longitudinal patterns. The resting posture is either stick-like, at an angle of 45 ∞, or ventrally bent to form a complete loop ( Sugi, 1987; Murase, 1998; Ohbayashi, 2000; Tominaga, 2000; Ebert, 2001). According to McGuffin (1967: 12): ‘20–30 annulets on each anterior abdominal segment; cuticle smooth at 50¥; body setae short, with blunt tips; setae on legs relatively long, pointed; prolegs of mature larva with 6–10 crochets in two incompletely separated groups (fig. 82g).’ If disturbed, the larvae of many Palaearctic species drop down and make several powerful side-to-side movements (pers. observ.).

Many species are polyphagous, and recorded hostplants range from woody to herbaceous taxa. They include Scrophulariaceae (Striga) , Sterculiaceae (Theobroma) , Asteraceae (Lactuca) , Polygonaceae (Rumex) , Lamiaceae(Thymus) , Poaceae (Zea) , Moraceae (Ficus) Rosaceae ( Filipendula , Sorbus ), Liliaceae (Litanthus) and Ericaceae (Vaccinium) for Scopula ; Rubiaceae ( Adina , Breonia , Mitragyna , Hymenodictyon and Mussaenda ) and Marantaceae (Maranta) for the Antitrygodes -group; Rubiaceae ( Oxyanthus , Randia ) and Sapindaceae (Blighia) for the Aletis -group ( Staude, 1999; Tanahara, 1999; Robinson et al., 2002); Lamiaceae (Thymus) , Caryophyllaceae ( Dianthus , Gypsophila ) and Brassicaceae (Alyssum) for the Glossotrophia -group ( King, 2000; Hausmann & Dötterl, 2003) and Rubiaceae (Pygmaeothamnus) for the Epicosymbia -group ( Staude, 1999). A few species are known to be of minor pest status, attacking tobacco ( Sannino & Balbiani, 1984; Sannino & Espinosa, 1999) and groundnut ( Satpathi, 1995).

Pupal cremaster has usually four pairs of setae, terminal pair usually much enlarged ( Khotko, 1977; Patocka, 1994), reduced to two in S. indicataria ( Nakamura, 1994) . In the Glossotrophia -group the proboscis case extends beyond the cremaster, curving inwards ( King, 2000; Hausmann, 2001).

Biology: Scopula includes species of both forested and open habitats as shown in the host-plant list. A number of species have been recorded feeding as adults on the tears, sweat and blood seeping from wounds of large mammals in South-east Asia ( Bänziger & Fletcher, 1985). Many species are day-active, and the species of the African Aletis and Cartaletis groups are exclusively diurnal.

Parasitoids: Homolobus truncator (Say) ( Braconidae : Homolobinae ) has been reported to parasitize the larva of S. annae (Mentzer) of the Glossotrophia group ( King, 2000).

Distinct species groups can be defined within Scopula . Their description is, however, beyond the scope of this paper. They include the Antitrygodes , Epicosymbia , Glossotrophia and Aletis groups.

Comments on systematics: Scopula is the largest and externally most heterogeneous genus of the tribe. Although only one character was found to support the monophyly of this genus, overall support is nevertheless not as weak as it may appear from the cladogram. Having studied a large number of species across different biogeographical regions, I have seen that the moths of this genus have many characteristic features such as separate sacculus and valvula of the valva, urceolate shaped juxta and spined signum, in addition to diagnostic structures of the male 8th sternite and tergite. All were found to be homoplastic, and accordingly, many are present in the majority of species, although one or a few may be absent in any given species. For example, the absence of ceras in many apical species such as Scopula sentinaria , Stigma kuldschaensis and Dasybela achroa was found to be due to secondary losses (79: 0, CI = 0.09, RI = 0.51). It is thus difficult to define this genus in terms of unique features; instead, Scopula can be characterized by a combination of characters, as already noted by Covell (1970). Many distinctive species groups within Scopula were found, but their relationships remain unresolved, probably due to the high degree of homoplasy that the characters display ( Fig. 134 ).

All the synonymized genera were found to have diagnostic structural characters of Scopula , although some of the taxa were found to have very divergent external appearance from the type species of the genus, S. ornata (Appendix) .

Figures

Figure 133. Strict consensus of 20 equally parsimonious cladograms. based on 141 characters of adult morphology and ecology (L = 876, CI = 0.21; RI = 0.58). Species are mentioned in their generic combinations prior to analysis. Names on the right-hand margin indicate genera as recognized here.

Figure 134. Cladogram 19 of 20 based on 141 characters from adult morphology and ecology (L = 853, CI = 0.22; RI = 0.59). Numbers above hash marks indicate character numbers and below character states. Black bars indicate unique and white bars non-unique synapomorphies. Species are mentioned in their generic combinations prior to analysis. Names on the right-hand margin indicate genera as recognized here.

Figure 135. A simplified cladogram of Figure 133 transformed to show generic relationships within the Scopulini. Taxa of uncertain generic association (incertae sedis) are shown in grey.

Figures 1–25. Type species of genera included in the phylogenetic analysis. The numbers on the figures refer to the numbers of the character states. 1. Idaea aversata. 2. Somatina anthophilata. 3. Dithalama cosmospila. 4. Isoplenodia arrogans. 5. Zythos turbata. 6. Lipomelia subusta. 7. Problepsis ocellata. 8. Scopula ornata. 9. Ignobilia urnaria. 10. Antitrygodes divisaria. 11. Epicosymbia dentisignata. 12. Glossotrophia confinaria. 13. Isoplenia trisinuata. 14. Cartaletis libyssa. 15. Aletis helcita. 16. Leucoxena lactea. 17. Stigma kuldschaensis. 18. Dasybela achroa. 19. Pseudocinglis eurata. 20. Cinglis humifusaria. 21. Autanepsia poliodesma. 22. Antilycauges pinguis. 23. Oar pratana. 24. Zygophyxia tornisecta. 25. Scopuloides fucata. Scale bars = 5.0 mm.

Figures 30–54. Male genitalia and aedeagus of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 30. Idaea aversata (slide number) PS704. 31. Somatina anthophilata (slide number) BMNH GEO20369. 32. Dithalama cosmospila PS707. 33. Isoplenodia arrogans BMNH GEO20527. 34. Zythos turbata BMNH GEO20371. 35. Lipomelia subusta BMNH GEO20530. 36. Problepsis ocellata BMNH GEO20370. 37. Scopula ornata PS692. 38. Ignobilia urnaria BMNH GEO20531. 39. Antitrygodes divisaria BMNH GEO20375. 40. Epicosymbia dentisignata BMNH GEO20365. 41. Glossotrophia confinaria BMNH GEO20382. 42. Isoplenia trisinuata BMNH GEO20363. 43. Cartaletis libyssa BMNH GEO20372. 44. Aletis helcita BMNH GEO20360. 45. Leucoxena lactea BMNH GEO20376. 46. Stigma kuldschaensis PS859. 47. Dasybela achroa PS897. 48. Pseudocinglis eurata PS894. 49. Cinglis humifusaria PS648. 50. Autanepsia poliodesma BMNH GEO20526. 51. Antilycauges pinguis BMNH GEO20381. 52. Oar pratana PS714. 53. Zygophyxia tornisecta BMNH GEO20380. 54. Scopuloides origalis PS58 (male of S. fucata type species of the genus was not available for study). Figures are not to scale.

Figures 55–79. Male 8th sternite of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 55. Idaea aversata (slide number) PS704. 56. Somatina anthophilata (slide number) BMNH GEO20369. 57. Dithalama cosmospila PS707. 58. Isoplenodia arrogans BMNH GEO20527. 59. Zythos turbata BMNH GEO20371. 60. Lipomelia subusta BMNH GEO20530. 61. Problepsis ocellata BMNH GEO20370. 62. Scopula ornata PS692. 63. Ignobilia urnaria BMNH GEO20531. 64. Antitrygodes divisaria BMNH GEO20375. 65. Epicosymbia dentisignata BMNH GEO20365. 66. Glossotrophia confinaria BMNH GEO20382. 67. Isoplenia trisinuata BMNH GEO20363. 68. Cartaletis libyssa BMNH GEO20372. 69. Aletis helcita BMNH GEO20360. 70. Leucoxena lactea BMNH GEO20376. 71. Stigma kuldschaensis PS859. 72. Dasybela achroa PS897. 73. Pseudocinglis eurata PS894. 74. Cinglis humifusaria PS648. 75. Autanepsia poliodesma BMNH GEO20526. 76. Antilycauges pinguis BMNH GEO20381. 77. Oar pratana PS714. 78. Zygophyxia tornisecta BMNH GEO20380. 79. Scopuloides origalis PS58 (the male of S. fucata. the type species of the genus. was not available for study). Figures are not to scale.

Figures 100–108. Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 100. Extensions on the anterior part of dorsal sclerite between antennae. Somatina microphylla (slide number) PS711. 101. Structure of iridescent scales. Zythos turbata. 102. Structure of iridescent scales. Problepsis conjunctiva. 103. Chaetosemata. Zythos avellanea. 104. Scales among chaetosemata. Cartaletis variabilis. 105. Dark patch at forewing transverse median line that is separated by vein CuA1 (veins drawn on top). Problepsis conjunctiva. 106. Dark patch at forewing M2 (veins drawn on top). Problepsis centrophora. 107. Anterior and posterior ventral lamina of metathorax metafurca. Scopula imitaria PS779. 108. Pouch on posterior margin of male metathorax coxa. Problepsis ocellata (slide number) BMNH GEO20421.

Figures 109–117. Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 109. Base of male metafurca (posterior view). Idaea aversata (slide number) PS766. 110. Base of male metafurca (posterior view). Cartaletis variabilis (slide number) BMNH GEO20575. 111. Membranous sac at posterior part of male metathoracic coxa (dorsal view). Scopula silonaria BMNH GEO20433. 112. Opening on dorsal sclerite of male metathorax (dorsal view). Scopula ternata PS718. 113. Hair pencils of tibia on outer and inner margin of male hindleg. Ignobilia urnaria BMNH GEO20579. 114. Pretarsus of male hindleg. Zythos turbata BMNH GEO20443. 115. Pretarsus of male hindleg. Problepsis achlyobathra BMNH GEO20724. 116. Pouch on male 2nd sternite with lateral invaginations. Antitrygodes parvimacula PS883. 117. Lateral appendices of socii. Problepsis plagiata PS709.

Figures 26–29. Interpretation of wing pattern homologies. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 26. Glossotrophia confinaria. 27. Problepsis aegretta. 28. Somatina chalyboeta. 29. Antitrygodes divisaria. Abbreviations: t.a., transverse anterior line. t.m., transverse median line. t.p.l., transverse posterior line. s.t.s.l., subterminal shade line. a.t.s.l., adterminal shade line. t.l., terminal line. Figures are not to scale.

Figures 80–99. Female genitalia of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 80. Idaea aversata (slide number) PS828. 81. Somatina anthophilata (slide number) BMNH GEO20472. 82. Dithalama cosmospila PS832. 83. Zythos turbata BMNH GEO20483. 84. Problepsis ocellata BMNH GEO20485. 85. Scopula ornata PS835. 86. Ignobilia urnaria (insert: signum) BMNH GEO20532. 87. Antitrygodes divisaria BMNH GEO20480. 88. Epicosymbia dentisignata BMNH GEO20462. 89. Glossotrophia confinaria BMNH GEO20476. 90. Isoplenia trisinuata BMNH GEO20464. 91. Cartaletis libyssa BMNH GEO20468. 92. Aletis helcita BMNH GEO20469. 93. Leucoxena lactea BMNH GEO20478. 94. Pseudocinglis eurata PS895. 95. Cinglis humifusaria PS644. 96. Antilycauges pinguis BMNH GEO20479 (insert: corpus bursae spines). 97. Oar pratana PS804. 98. Zygophyxia tornisecta BMNH GEO20474. 99. Scopuloides fucata PS59. Figures are not to scale.

Figures 118–132. Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 118. Posterior margin of male 8th tergite. Scopula submutata (slide number) PS693. 119. Posterior margin of male 8th tergite. Dithalama cosmospila PS707. 120. Posterior margin of male 8th tergite. Somatina eurymitra PS888. 121. Posterior margin of male 8th tergite. Zythos avellanea PS879. 122. Posterior margin of male 8th tergite. Antitrygodes divisaria (slide number) BMNH GEO20375. 123. Apex of coremata. Scopula rhodocraspeda BMNH GEO20700. 124. Position of sacculus. Zythos obliterata BMNH GEO20591. 125. Dorsally fused anterior margins of tegumen (dorsal view). Zythos turbata BMNH GEO20371. 126. Plate-shaped sclerotization at proximal end of ductus ejaculatorius. Antilycauges pinguis BMNH GEO20381. 127. Sclerotization of vesica. Scopula rivularia BMNH GEO20684. 128. Dorsal diverticulum of vesica with sharp sclerotization at apex. Problepsis plagiata PS709. 129. Parallel folds of vesica. Zythos turbata BMNH GEO20371. 130. Signum. Somatina indicataria PS834. 131. Bare zone of signum. Zythos obliterata BMNH GEO20592. 132. Flexible lamella antevaginalis divided into two parts. Zythos avellanea PS880.