Wolcott 1923: 144, 2013

Kawahara, Akito Y., 2013, <strong> Systematic revision and review of the extant and fossil snout butterflies (Lepidoptera: Nymphalidae: Libytheinae) </ strong>, Zootaxa 3631 (1), pp. 1-74 : 9-10

Key to genera of Libytheinae

1. Rs 2 of forewing with base closer to fork of Rs 3 + Rs 4 than discal cell ( Figs. 53–61 ), band between forewing M 1 and M 3 closer to wing margin than discal cell, male eighth abdominal tergum quadrifid (Figs. 14–21), aedeagus sigmoid ( Figs. 39–46 ), signa present ( Fig. 51 ); Old World....................................................................... Libythea

– Rs 2 of forewing with base approximately midway between apex of discal cell and fork of Rs 3 + Rs 4 ( Figs. 62–68 ), band between forewing M 1 and M 3 closer to discal cell than wing margin, male eighth abdominal tergum trifid (Figs. 22–25), aedeagus curved dorsally ( Figs. 47–50 ), signa absent ( Fig. 52 ); New World.................................... Libytheana

Genus Libythea Fabricius 1807

Libythea Fabricius 1807: 284 [type species Libythea celtis ( Laicharting [1782]) , designated by Latreille (1810)].

Lybithea Rafinesque 1815: 127 .

Hecaërge Ochsenheimer 1816: 32 . Aurivillius 1898: 246 [= Libythea ].

Hecaërgae Hübner [1819], imprint 1816–[1827]: 100 [misspelling of Hecaërge Ochsenheimer ].

Chilea Billberg 1820: 79 . Pagenstecher 1901: 1 [= Libythea ].

Libythaea Godart 1821.

Libythée Godart 1822: 50.

Hypatus Hübner 1822: 3 . Pagenstecher 1901: 1 [= Libythea ].

Libythaeus Boitard 1828: 299 .

Hecaaerge Blanchard 1840: 445.

Lybathea Edwards 1874: pl. [46].

Prolibythea Scudder 1889a: 461 . Shields 1985a: 13 [= Libythea ].

Hecaerge Scudder 1889a: 461 . Hemming 1934: 96 [= Libythea ].

Dichora Scudder 1889a: 470 . Aurivillius 1898: 246 [= Libythea ]. Carcasson 1995: 471 [junior homonym of Dichora Paetel 1875 (Crustacea) ].

Barbarothea Scudder 1892: 21 . Shields 1985a: 13 [= Libythea ].

Hypati Hübner 1894: 350.

Libytahe Heyne [1895]: 771.

Libytheae Kirby 1896: 5.

Libythea (Libythea) : Pagenstecher 1902: 2.

Libythéa: Berge 1910: 56 [unjustified emendation].

Libythia Weeks 1911: xii.

Libethea Wolcott 1923: 144 .

Lybythea Fujioka 1975: 294.

Included Species. Libythea celtis ( Laicharting [1782]) , L. cinyras Trimen 1866 , L. collenettei Riley 1928 , L. geoffroyi Godart 1822 , L. labdaca Westwood 1851 , L. laius Trimen 1879 , L. lepita Moore [1858] , L. myrrha Godart 1819 , L. narina Godart 1819 .

Diagnosis. Species of Libythea share at least five synapomorphies that distinguish it from Libytheana : 1) Rs 2 of forewing with base closer to fork of Rs 3 + Rs 4 than discal cell, 2) band across forewing cells M 1 and M 2 distal, 3) phallus sigmoid, 4) dorsal margin of valve strongly curved ventrad, 5) corpus bursa with signa (Kawahara 2009).

Remarks. Extant Libythea species can be easily separated geographically from Libytheana species because Libythea is found only in the Old World. Pagenstecher (1901) revised this genus and synonymized four generic names, Chilea , Dichora , Hecaërge , Hypatus , and Libythaeus .

Pennington (1994) mistakenly listed Jera Lindsey 1925 as a subgenus of Libythea . Lindsey (1925) unnecessarily proposed Jera as a replacement name for Dichelura Mabille (Hesperiidae) , because Dichelura is a junior homonym of Dichelura Macquart (Diptera) . Pennington (1994) had probably confused Dichelura with Dichora . Carcasson (1995) noted that Dichora Scudder is a junior homonym of Dichora Paetel (Crustacea) .

At least two non- Libythea species appeared in the literature as belonging in Libythea . These are Libythea cuvierii Godart and L. moluccana Felder. Godart (1819: 171) described Libythea cuvierii as a snout butterfly, but Felder (1861) subsequently erected the genus Libythina for cuvierii Godart. In the contents of his publication, Jordan (1898) labeled plate 15, figure 52 of his antennal illustrations as Libythea moluccana . From the captions on plate 15 it is clear that the figure was mislabeled ( Fig. 52 is Acarea moluccana ).

Key to species of Libythea

1. Antennal club gradually incrassate ( Fig. 6a of Kawahara [2009])................................................ 2

– Antennal club abruptly incrassate ( Fig. 6b of Kawahara [2009])................................................ 3

2(1). Anterior margin of hindwing lobed near fusion of costa and Sc + R 1 ( Figs. 53 , 69–72 ); Europe and northwestern Africa to western India.............................................................................. Libythea celtis

– Anterior margin of hindwing smooth or weakly lobed near fusion of costa and Sc + R 1 ( Figs. 59 , 119–128 ); from north-central India east to Japan.......................................................................... Libythea lepita

3(1). Forewing apex smooth, wings orange and brown ( Figs. 55 , 73–76 ); Marquesas Islands............... Libythea collenettei

– Forewing apex falcate or weakly falcate; background color of wings blue, orange, or dark brown; Africa, India, Southeast Asia and other islands of Oceania............................................................................. 4

4(3). Forewing dorsum with club-shaped yellow-orange band from cell CuA 1 to wing base ( Figs. 129–140 )..... Libythea myrrha

– Forewing dorsum without club-shaped yellow-orange band from cell CuA 1 to wing base............................. 5

5(4). Forewing dorsum with a single orange, L-shaped apical mark in discal cell ( Fig. 77 ); possibly extinct, Mauritius.................................................................................................. Libythea cinyras

– Forewing dorsum without an apical mark in discal cell, or if present, round, square or rectangular...................... 6

6(5). Hindwing lobe at junction of CuA 2 and wing margin extending more than 1 mm beyond 1A + 2A (e.g., Fig. 58 ), dorsal wing surface brown (e.g., Figs. 105, 109 ); Africa, southern India and Sri Lanka......................................... 7

– Hindwing without lobe at junction of CuA 2 and wing margin, or if lobed, lobe extending less than 1 mm beyond 1A + 2A dorsal wing surface blue (e.g., Fig. 93 ); Mauritius, Continental and Southeast Asia.................................... 8

7(6). Forewing dorsum with discal cell marks connected medially or separated by less than 1 mm, dorsal wing marks pale orange ( Figs. 109–117 ); eastern Africa, Southern India, Sri Lanka, and Madagascar............................ Libythea laius

– Forewing dorsum with discal cell marks separated medially by more than 1 mm, dorsal wing marks white to pale brown ( Figs. 103–108 ); western Africa and Madagascar..................................................... Libythea labdaca

8(6). Hindwing dorsum with a bright median white band from M 1 to CuA 1, dorsal wing surface of male black with white marks ( Figs. 141, 143 )........................................................................... Libythea narina

– Hindwing dorsum without a white median band or very faint white band present from M 1 to CuA 1, dorsal wing surface of male mostly or entirely blue.................................................................... Libythea geoffroyi

Figures

FIGURE 53. Wing venation of Libythea celtis. Fine lines indicate margins of wing markings.

FIGURES 54−62. Wing venation of Libythea and Libytheana. Fine lines indicate margins of wing markings; 54. Libythea cinyras; 55. Libythea collenettei; 56. Libythea geoffroyi; 57. Libythea labdaca; 58. Libythea laius; 59. Libythea lepita; 58. Libythea myrrha; 60. Libythea narina; 61. Libytheana florissanti.

FIGURES 39−50. Lateral illustrations of the aedeagus of Libythea and Libytheana. 39. Libythea celtis; 40. Libythea collenettei; 41. Libythea geoffroyi; 42. Libythea labdaca; 43. Libythea laius; 44. Libythea lepita; 45. Libythea myrrha; 46. Libythea narina; 47. Libytheana carinenta carinenta; 48. Libytheana fulvescens; 49. Libytheana motya; 50. Libytheana terena.

FIGURE 51−52. Lateral view of female genitalia. 51. Libythea celtis. 52. Libytheana carinenta. cb = corpus bursae, db = ductus bursae, gp = genital plate, sgn = signa, st7 = seventh abdominal sternum, st8 = eighth abdominal sternum, t7 = seventh abdominal tergum, t8 = eighth abdominal tergum.

FIGURES 63−68. Wing venation of Libytheana. 63. Libytheana vagabunda; 64. Libytheana carinenta carinenta; 65. Libytheana carinenta bachmanii; 66. Libytheana fulvescens; 67. Libytheana motya; 68. Libytheana terena.

FIGURES 2−13. Lateral view of the male eighth abdominal tergum of Libythea and Libytheana with scales and hairs removed. 2. Libythea celtis; 3. Libythea collenettei; 4. Libythea geoffroyi; 5. Libythea labdaca; 6. Libythea laius; 7. Libythea lepita; 8. Libythea myrrha; 9. Libythea narina; 10. Libytheana carinenta; 11. Libytheana fulvescens; 12. Libytheana motya; 13. Libytheana terena.

FIGURES 69−84. Right dorsal and ventral wing surfaces of Libythea. The second image of each specimen shows the ventral surface of the wings. 69, 70. Libythea celtis (Laicharting): ♂, Russia, Kirgizia, Kirgizaskii Ridge, Ala−Archa, Ak−Sai (TFC); 71, 72. Libythea celtis (Laicharting): ♀, France, St. Maxime (BMNH); 73, 74. Libythea collenettei Riley: ♂, Marquesas Islands, Ua Pou, Poohekaei Summit (BPBM); 75, 76. Libythea collenettei Riley: ♀, Marquesas Islands, Nuku Hiva, Toovi Plateau (BPBM); 77, 78. Libythea cinyras Trimen: Holotype, Mauritius, Moka District (BMNH); 79, 80. Libythea geoffroyi alompra Moore: Holotype ♂, Myanmar, Ataran Valley (BMNH); 81, 82. Libythea geoffroyi ceramensis Wallace: ♂, Indonesia, Seram (BMNH); 83, 84. Libythea geoffroyi ceramensis Wallace: ♀, Indonesia, Seram, Mansuela (BMNH).

FIGURES 117−132. Right dorsal and ventral wing surfaces of Libythea. The second image of each specimen shows the ventral surface of the wings. 117, 118. Libythea laius lepitoides Moore: ♂, India, Nilgiris, Conoor (BMNH); 119, 120. Libythea lepita formosana Fruhstorfer: Holotype ♂, Taiwan (BMNH); 121, 122. Libythea lepita amamiana Shirôzu: ♂, Japan, Okinawa, Nago (MTC); 123, 124. Libythea lepita amamiana Shirôzu: ♀, Japan, Okinawa, Shimajiri, Kitadaito−mura (TFC); 125, 126. Libythea lepita lepita Moore: ♂, Japan, Yamanashi, Hinoharu (AYK); 127, 128. Libythea lepita lepita Moore: ♀, Japan (BMNH); 129, 130. Libythea myrrha myrrha Godart: ♂, Indonesia, Java (BMNH); 131, 132. Libythea myrrha myrrha Godart: ♀, Indonesia, West Java (BMNH).

FIGURES 101−116. Right dorsal and ventral wing surfaces of Libythea. The second image of each specimen shows the ventral surface of the wings. 101, 102. Libythea geoffroyi howarthi Petersen: Paratype ♂, Solomon Islands, Rennell (BMNH); 103, 104. Libythea labdaca ancoata Grose−Smith: Holotype ♂, Madagascar, Irast (BMNH); 105, 106. Libythea labdaca labdaca Westwood: ♂, Uganda, Mulange (CMNH); 107, 108. Libythea labdaca labdaca Westwood: ♀, Cameroon, Bafia (CMNH); 109, 110. Libythea laius laius Trimen: ♂, Malawi, Cholo (CMNH); 111, 112. Libythea laius laius Trimen: ♀, Malawi, Cholo (CMNH); 113, 114. Libythea laius tsiandava Grose−Smith: Holotype ♂, Madagascar (BMNH); 115, 116. Libythea laius tsiandava Grose−Smith: ♀, Madagascar, Mahabo (BMNH).

FIGURES 85−100. Right dorsal and ventral wing surfaces of Libythea. The second image of each specimen shows the ventral surface of the wings. 85−86. Libythea geoffroyi genia Waterhouse: ♂, Australia, 14 km SE of Kalumburu (ANIC); 87, 88. Libythea geoffroyi genia Waterhouse: ♀, Australia, 14 km SE of Kalumburu (ANIC); 89, 90. Libythea geoffroyi geoffroyi Godart: ♂, Indonesia, Timor, Suai, Portug (BMNH); 90, 91. Libythea geoffroyi geoffroyi Godart: ♀, Indonesia, Timor, Suai, Portug (BMNH); 92, 93. Libythea geoffroyi philippina Staudinger: ♂, Philippines, Palawan (BMNH); 94, 95. Libythea geoffroyi philippina Staudinger: ♀, Philippines, Palawan (BMNH); 96, 97. Libythea geoffroyi pulchra Butler: ♂, Papua New Guinea, New Ireland (BMNH); 98, 100. Libythea geoffroyi pulcrha Butler: ♀, Papua New Guinea, New Ireland (BMNH).

FIGURES 133−148. Right dorsal and ventral wing surfaces of Libythea and Libytheana. The second image of each specimen shows the ventral surface of the wings. 133, 134. Libythea myrrha hecura Fruhstorfer: ♂, Borneo (BMNH); 135, 136. Libythea myrrha rama Moore: ♂, Holotype, Sri Lanka, Colombo (BMNH); 137, 138. Libythea myrrha sanguinalis Fruhstorfer: ♂, India, Assam (USNM); 139, 140. Libythea myrrha sanguinalis Fruhstorfer: ♀, India, Assam, Naga Hills (BMNH); 141, 142. Libythea narina Godart: ♂, Myanmar, Tenasserim, Ataran Valley (BMNH); 143, 144. Libythea narina Godart: ♀, Philippines, Palawan (BMNH); 145, 146. Libytheana fulvescens (Lathy): ♂, Dominica, 1−2 km north of Colihaut (MCLB); 147, 148. Libytheana fulvescens (Lathy): ♀, Dominica, Copt Hall (USNM).