Dithalama, Meyrick, 1888

Sihvonen, Pasi, 2005, Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae), Zoological Journal of the Linnean Society 143 (4), pp. 473-530 : 501-505

THE DITHALAMA , ISOPLENODIA , SOMATINA AND ZYTHOS GROUP OF GENERA

This group of four genera is characterized by five synapomorphies ( Figs 133-135 ): presence of two forewing areoles formed by R veins (48: 1, six occurrences and one reversal); forewing vein R 5 runs from the distal areole and is stalked with veins R 2 -R 4 (49: 2, eight occurrences); posterolateral appendices are absent on male 8th sternite (86: 1, five occurrences and six reversals); the medial margin of the sacculus is upturned (110: 1, four occurrences); lamella antevaginalis is flap-like, flexible, formed by one plate (139: 1, 13 occurrences and two reversals). Within the group, the relationships are not resolved. Somatina eurymitra is incerta sedis and recognition of Isoplenodia renders Dithalama paraphyletic. These three taxa form the sister group to Somatina + Zythos .

Comments on systematics

This group of genera, although not found to be supported by unique synapomorphies ( Figs 134 , 135 ), appears to be rather distinctive. Cerata are absent in most cases, and the species have two areoles in the forewings. In the other group of genera (i.e. Scopula , Lipomelia and Problepsis ) the cerata are fully developed and in most species the forewings have only one areole.

DITHALAMA MEYRICK, 1888

( FIGS 3 , 32 , 57 , 82 , 119 , APPENDIX)

Diagnosis: description is based on type species only (see ‘Comments on systematics’, below). Monophyly supported by five synapomorphies: transverse median line of wings is obscure (25: 1, 12 occurrences); posterior margin of male 8th tergite is concave, with two sclerotized lateral spines (88: 2, unique, Fig. 119 ); medial margin of sacculus is not upturned (110: 0, three occurrences); presence of posteriorly directed lobe on top of vinculum with a medial ridge (115: 1, 15 occurrences); apices of wing-like juxta processes are fused to sacculus of valva (116: 2, seven occurrences).

Imago: wingspan 19–24 mm. Male antenna fasciculate, sensilla in single rows; female antenna fasciculate, simple. Pilifer present. Wing patterns variable, transverse lines ranging from almost straight to hardly recognizable; discal spot dark; terminal line discontinuous; wing patterns repeated on underside but indistinct. Forewing with two areoles; R 5 from distal areole, either stalked with R 2 -R 4 or straight from accessory cell. Outer margin hair pencil of male hindleg tibia present, inner either present or absent; distal spurs of hindleg tibia absent; tarsomeres shortened but not fused; claws either two in number or absent. Female hindleg tibia with two pairs of spurs.

Male abdomen and genitalia: pouch on male 2nd sternite round; anterolateral extensions present or absent. One membranous pouch between male 7th and 8th sternite. Male 8th sternite sclerotized; anterior margin elongated medially; laterally turned extensions or two blunt lobes on posterior margin; cerata, mappa and membranous posterolateral appendices absent ( Fig. 57 ). Posterior margin of male 8th tergite concave with two sharp cerate-like projections or with two round lateral lobes ( Fig. 119 ). Socii separate, setose. Ventral margin of tegumen with round lobes, separate or fused. Valvae symmetrical, variably sclerotized, anterior margin of sacculus medially upturned or levelled. Transtilla present. Anterior margin of juxta with wing-like processes, apex fused to sacculus of valva or absent. Vinculum enlarged . Aedeagus straight with one or several teeth, carina, at distal end; vesica simple, curved sac, without sclerotizations .

Female genitalia: lamella antevaginalis sclerotized, flap-like, flexible, medially elongated. Ostium bursae weakly sclerotized; ductus bursae wide, proximal part either membranous or sclerotized; ductus seminalis arises from ductus bursae. Corpus bursae elongated sac; signum either granulate, spines point away from centre or absent; long spines on inner surface of corpus bursae absent.

Distribution and species diversity: Australia and Tasmania. Four species identified tentatively ( McQuillan & Edwards, 1996). See Appendix under Dithalama .

Immature stages: unknown.

Biology: unknown, although D. cosmospila is suspected to live on Eucalyptus ( Prout, 1920 -41).

Comments on systematics: the genus is in need of revision, as was already noted by Prout (1920 -41). He speculated that Dithalama is closely related to Somatina and Autanepsia , and the latter monotypic genus should perhaps be combined with Dithalama . In this analysis Dithalama was recovered as paraphyletic with regard to Isoplenodia . The Dithalama + Isoplenodia connection, although not discussed earlier, is supported by a number of character absences ( Fig. 134 ). In addition to Dithalama + Isoplenodia , the secondary loss of sclerotizations of vesica (125: 1), which is a derived feature in this analysis, has occurred outside of these taxa only in Tasmanian species Dasybela achroa and South African species Zygophyxia relictata , which were found to be unrelated to Dithalama . Absence of male hindleg claws (67: 0), pretarsus arolium (68: 0) and pulvillus (69: 1), on the other hand, are found also in a number of taxa in Zythos , Problepsis and Antitrygodes .

ISOPLENODIA PROUT, 1932

( FIGS 4 , 33 , 58 )

Diagnosis: monophyly supported by eight synapomorphies: absence of pilifer (character and character state 0: 1, unique); ventrolateral sensilla on proximal part of male flagellomere are in multiple rows (8: 1, nine occurrences); absence of anterior ventral lamina of metathoracic metafurca (52: 1, six occurrences); absence of opening on male metathoracic dorsal sclerite (58: 1, character state ‘present’ shown in Fig. 112 , unique); fusion of male hindleg tarsomeres 1–5 (66: 1, two occurrences); cerata are fully developed (79: 2, three occurrences, two reversals); mappa is bare 83: 2 (four occurrences); ventral margin of tegumen is unmodified, straight (100: 0, three reversals).

Imago: wingspan c. 16 mm. Head smooth-scaled; male antenna fasciculate; sensilla arranged in multiple rows, proximal row elongated. Pilifer absent. Wings light brown, transverse lines weakly expressed; discal spot dark; terminal line discontinuous, equally wide; underside light fuscous. Forewing with two areoles; R 5 from distal areole, stalked with R 2 -R 4. Opening on male metathorax dorsal sclerite absent. Outer margin hair pencil of male hindleg tibia short; inner absent; tarsomere segments 1–5 fused, claws absent. Female hind tibia with 2 + 2 spurs.

Male abdomen and genitalia: anterolateral extensions on male 2nd sternite absent. One membranous pouch between male 7th and 8th sternite. Anterior margin of male 8th sternite elongated medially; cerata symmetrical; mappa constricted at base; membranous, posterolateral appendices absent. Posterior margin of male 8th tergite concave. Socii rudimentary, short. Ventral margin of tegumen straight. Valvae symmetrical, soft, setose, valvula and sacculus separate. Anterior margin of sacculus medially upturned, covered with pointing setae. Transtilla present. Juxta weakly sclerotized, narrow at base, but with long wing-like processes. Vinculum enlarged , dorsally narrow, v-shaped ( Fig. 33 ). Aedeagus curved ventrally, with long caecum; without sclerotizations.

Female genitalia: unknown.

Distribution and species diversity: known from Madagascar only. One species.

Immature stages: unknown.

Biology: unknown. Comments on systematics: the genus is retained as valid but only tentatively (see comments on Dithalama ). Prout speculated that Isoplenodia might be of a close relative of the African genera Epicosymbia and Isoplenia , on the basis of two forewing areoles, and long ciliated male antenna. This was not supported in this analysis and the condition of two forewing areoles (48: 1) was shown to be a very homoplastic character. I. arrogans was found to have cerata-like structures laterally on the male 8th sternite, but these are immobile and without apical setae, contrary to other studied material. Examination of female structures could give further clues about the systematic position of this species.

SOMATINA GUENÉE, [1858]

( FIGS 2 , 27 , 31 , 56 , 81 , 120 , APPENDIX)

Diagnosis: monophyly supported by three synapomorphies: sacculi of valva are asymmetrical (105: 1, seven occurrences); valvuli of valva are asymmetrical (108: 1, seven occurrences); anterior margin of juxta is with wing-like processes, apex is fused to sacculus of valva (116: 2, seven occurrences).

Imago: wingspan 23-33 mm. Male antenna fasciculate; proximal row of sensilla sometimes weakly elongated, those sensilla in single rows. Female antenna fasciculate, simple. Pilifer present. Wing patterns variable, colour ranging from white to reddish-yellow, greyish-brown in many species; often irrorated with dark scales; dark band on forewing termen between transverse anterior and median lines in many species; transverse lines dentate, straight or sometimes modified, taking unusual forms; discal spot weak, dark; iridescent scales usually absent, glossy in few species; terminal line usually discontinuous, equally wide; patterns often repeated on underside, fuscous, sometimes absent. Forewing with two areoles; R 5 from distal areole, stalked with R 2 –R 4 or straight from cell. Outer and inner margin hair pencil of male hindleg tibia usually present, outer concealed in a furrow formed by long scales in many species; apical spurs usually absent, sometimes two; tarsomeres normally developed, 5- segmented; two claws. Female hind tibia with 2 + 2 spurs.

Male abdomen and genitalia: pouch on male 2nd sternite round, posterior margin often invaginated; anterolateral extensions usually present. Small lateral pouches between sternites 3 and 4 rarely present. Usually one, sometimes two membranous pouches between male 7th and 8th sternite. Male 8th sternite variable, x-shaped and heavily sclerotized in many species; anterior margin elongated medially or bifurcate; cerata usually absent ( Fig. 56 ), sometimes present as fully developed; mappa absent, setose or bare; posterolateral appendices usually present. Male 8th tergite posterior margin concave, with two round lobes or sharp projections ( Fig. 120 ). Socii separate, covered with long setae. Ventral margin of tegumen unmodified, rarely with small round or elongated projections. Sacculus and valvula of valva separate, with pronounced asymmetry; valvula shape varies from wide, blunt-ended to narrow, long, with sharp apex; medial and lateral margin of sacculus often with upturned projections. Anterior margin of juxta with wing-like processes, apex rarely free, instead fused to sacculus in most species, and sometimes asymmetrical; sometimes juxta processes functionally replacing sacculus of valva. Transtilla narrow weakly sclerotized band, rarely wide. Vinculum enlarged , symmetrical and usually convex. Aedeagus bent, sometimes with small projections near apex; vesica complex, with several large, often twisted diverticula ( Fig. 31 ); variably sclerotized but rarely with distinct cornuti.

Female genitalia: lamella postvaginalis usually absent; lamella antevaginalis large, sclerotized, medially invaginated, flap-like structure in most species. Ostium bursae weakly sclerotized. Ductus bursae usually coiled, weakly sclerotized. Ductus seminalis opens from corpus bursae neck. Corpus bursae ovoid sac; signum elongated, made of separate spines, rather large compared to size of corpus bursae.

Distribution and species diversity: from Africa to the Oriental region and Australia. Forty-four species and several subspecies, although see ‘Comments on systematics’.

Immature stages: I have been unable to find any references. Nakamura (1994) described the pupa of Somatina indicataria (Walker) , but the taxon is considered to belong to Scopula in the present study.

Biology: S. anthophilata has been recorded from Gardenia jasminoides and Randia dumetorum (Rubiaceae) and Rosa (Rosaceae) . Somatina omicraria and S. virginalis have been recorded from the Oleaceae genus Jasminum ( Robinson et al., 2002) .

Comments on systematics: Sterrhinae species with two areoles on the forewings have traditionally been considered to belong to Somatina , in contrast to Problepsis and Scopula , which have one areole. Since this character was shown to be homoplastic in the present study (48: 1), and as both states occur in many lineages, Somatina was considered to be a polyphyletic assemblage of species, as noted by Inoue (1992), with many species formerly ascribed to it shown to belong to either Problepsis or Scopula .

While the remaining species should be revised against the generic descriptions given here and, if necessary, transferred to the appropriate genera (see Appendix for examples), Somatina is, however, likely to be a monophyletic group of species. It has a number of characteristic, although not unique, features. One such feature is the anterior margin of the juxta that is fused to sacculus of the valva (116: 2). The male of S. anthophilata , the type species of the genus, has small lateral pouches between sternites 3 and 4 ( Holloway, 1997), although as these were not seen in other studied taxa, this autapomorphy was not coded. Traditionally, Somatina has been considered a close relative of Scopula or Problepsis (e.g. Prout, 1920 -41, 1934-39), but in the present study species of the redefined genus are associated with Zythos . This connection is supported by the sclerotized posterior part of male 8th sternite (85: 1, Fig. 59 ), which is found elsewhere only in Dithalama desueta , and by the presence of parallel folds on the vesica (132: 1, 129), found also in a few Scopula taxa. Several characters, e.g. the structure of the socii and signum, indicate a close relationship with Scopula .

Somatina irregularis and S. eurymitra were considered incertae sedis, but were retained in their traditional generic combinations, as they did not group together with the recognized genera. Somatina irregularis grouped together with Scopula haemaleata , sharing with it a number of unique features, such as a bifurcated cerata (82: 1) and a juxta with long processes laterally (117: 1). Somatina eurymitra , meanwhile, showed affinities with Somatina and Problepsis ; for example, the male 8th tergite is concave with two sharp lateral projections (88: 3), a feature typical of Somatina , while the cerata fused with the mappa (80: 1) is a common condition in Problepsis . Females of S. eurymitra were not available for study. It remains to be tested whether these taxa actually represent larger monophyletic groups.

ZYTHOS FLETCHER, 1979

( FIGS 5 , 34 , 59 , 83 , 101, 103 , 114 , 121, 124, 125, 129, 131, 132 , APPENDIX)

Diagnosis: monophyly supported by 20 synapomorphies: iridescent scales of wings are restricted to a few rows, excluding discal spots (16: 3, unique); iridescent scales of wings are glossy and longitudinally grooved (17: 1, unique, Fig. 101 ); discal spot of forewing is not unicolorous and dark (19: 1, six occurrences);discal spot of forewing is lunular (20: 2, two occurrences); presence of a spot at vein endings at forewing margin (40: 1, two occurrences); absence of hair pencil on inner margin of male hind tibia (61: 1, seven occurrences); male hindleg is with one claw only (67: 1, unique, Fig. 114 ); absence of pretarsus arolium on male hindleg (68: 0, six reversals); two pouches are present between male 7th and 8th sternites (76: 1, four occurrences, Fig. 59 ); absence of posterolateral appendices on male 8th sternite (86: 1, five occurrences, five reversals); posterior margin of male 8th tergite is convex (88: 4, unique, Fig. 121 ), posterolateral margin of male 8th tergite has two membranous appendices (90: 1, Fig. 121 ); absence of setae on socii (96: 0, unique); anterior margins of tegumen are dorsally fused (101: 1, unique, Fig. 125 ); absence of transtilla (102: 2, unique); valva have symmetrical sacculi (105: 0, two occurrences); valvula of valva lies ventral of sacculus (107: 1, unique, Fig. 124 ); valvuli of valvae are symmetrical (108: 0, two occurrences); anterior margin of juxta is without wing-like processes (116: 0, nine occurrences); signum has a bare zone laterally on both sides of median ridge (135: 1, unique, Fig. 131 ).

Imago: wingspan 32-40 mm. Male antennae fasciculate, sensilla in single rows; female antennae fasciculate, simple. Pilifer present. Sensilla styloconicae forming serrate lining at distal end of proboscis. Chaetosemata prominent. Wing pattern variable, often with fine striations on extensive areas of wings; costa and middle parts often olive grey; often with longitudinally grooved iridescent scales on outer half of forewing, excluding discal spots ( Fig. 101 ); subterminal shaded line and terminal line touching each other at fore- and hindwings in many species; discal spots dark, often lunular on forewings; pale markings on anterior side of hindwing discal spots in many species; terminal line discontinuous, equally wide, dark spot at vein end at forewing margin; underside often unicolorous, ranging from light brown to light orange. Forewings with two areoles; R 5 from distal areole, either stalked with R 2 –R 4 or straight from accessory cell. Hair pencil of outer margin of male hindleg tibia present, point of origin prominent; inner margin hair pencil absent; tarsomeres normally developed; one claw ( Fig. 114 ); arolium absent; pulvillus present. Female hindleg tibia with 2 + 2 spurs.

Male abdomen and genitalia: pouch on male 2nd sternite round, large; anterolateral extensions present. Two membranous pouches between male 7th and 8th sternite. Male 8th sternite heavily sclerotized, anterior margin bifurcate; posterior margin with blunt extensions or medial projections; cerata, mappa and membranous posterolateral appendices absent ( Fig. 59 ). Male 8th tergite posterior margin convex, with two membranous appendices on posterolateral margin ( Fig. 121 ). Genitalia capsule oval. Socii wide, separate, without setae. Ventral margin of tegumen slightly curved inwards. Valvae symmetrical, valvula and sacculus separate; anterior margin of sacculus medially upturned or levelled; valvula moved ventrad of sacculus ( Fig. 124 ), often curved dorsally. Transtilla absent; instead, anterior margins of tegumen are fused dorsally ( Fig. 125 ). Anterior margin of juxta without wing-like processes. Vinculum enlarged . Aedeagus almost straight, apex with small blunt extension; vesica prominent, long, often with long lateral diverticulum and sclerotized parallel folds.

Female genitalia: lamella antevaginalis flap-like, flexible, medially either concave or bilobed. Ostium bursae heavily sclerotized. Ductus bursae membranous, coiled; ductus seminalis arises from ductus bursae. Corpus bursae an elongate sac; signum often with narrow medial, sclerotized strip followed by bare zone ( Fig. 131 ) and granulate field of separate spines on both sides laterally.

Immature stages: unknown.

Distribution and species diversity: from India to Papua New Guinea. Morphologically, a compact genus with 11 species.

Biology: adults occur predominantly in the understorey of lowland rain forests and have been collected from carrion-baited pitfall traps ( Holloway, 1997). The serrated distal end of the proboscis indicates an adaptation to piercing. Frequently taken from secondary forests as well ( Chey, 1994).

Comments on systematics: Prout (1920 -41), followed by Holloway (1997), considered Zythos to be a close relative of Ignobilia on the basis of striated facies and pale underside of wings. In the present study Zythos is associated with Somatina (see Discussion under that genus).

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 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.

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 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.