AFROTROPICAL . MARK C. WILLIAMS. http://www.lepsocafrica.org/?p=publications&s=atb Updated 5 February 2021

Genus Hübner, [1818] Wood Whites

In: Hübner, [1808-18]. Zuträge zur Sammlung exotischer Schmettlinge [sic] 1: 13 ([3] + 4-6 + [7] + 8- 32 + [33] - [40] pp.). Augsburg. [7:-:8]. Type-: Leptosia chlorographa Hübner, by subsequent designation (Scudder, 1875. Proceedings of the American Academy of Arts and Sciences 10: 204 (91-293). [extralimital].

= Nychitona Butler, 1870. Cistula Entomologica 1: 34, 41 (33-58). Type-species: Papilio dorothea Fabricius, by original designation.

Synonym based on extralimital type-species: Nina Horsfield.

Opaque Wood White (Leptosia wigginsi wigginsi ). Image courtesy Jeremy Dobson.

The Leptosia belongs to the Family Swainson, 1820; Subfamily Swainson, 1820; Tribe Leptosiaini Braby, 2014. There are no other genera in the Tribe Leptosiaini in the Afrotropical Region.

Leptosia (Wood Whites) is an essentially Afrotropical genus of nine species. Seven species are Afrotropical and two are found extralimitally – L. nina (India to Australia) and L. lignea (Sulawesi). The genus was revised by Bernardi in a series of publications (1952, 1959, 1964, 1966).

* (Stoll, [1782])# African Wood White

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African Wood White (Leptosia alcesta), Burman Bush, Durban. Image courtesy Steve Woodhall.

Papilio alcesta Stoll, [1782] in Stoll, [1780-2]. Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en America 4 [part]: 175 (29-252). Amsteldam & Utrecht. Pontia alcesta Cramer. Trimen, 1862c. Pontia alcesta (Cramer, 1782). Trimen & Bowker, 1889. Leptosia alcesta Stoll. Swanepoel, 1953a. Leptosia alcesta (Stoll, 1782). Dickson & Kroon, 1978. Leptosia alcesta (Stoll, 1780). Pringle et al., 1994: 298. [date of authorship erroneous]

Leptosia alcesta alcesta. Male. Left – upperside; right – underside. Cape 3 Points, western . December 2002. R Vorgas. ABRI-2019-2948. Images M.C. Williams ex ABRI Collection.

Leptosia alcesta alcesta. Female. Left – upperside; right – underside. Putu Range, . December 2010. Safian, Zakar. ABRI-2019-2949. Images M.C. Williams ex ABRI Collection.

Alternative common names: African Spirit; Flip Flop. Type locality: “Côte de Guinée”. Distribution: Senegal, Gambia, -Bissau, Guinea (Bacelar, 1949), Liberia, , Ghana, , Benin (Fermon et al., 2002), , , , , Congo, , , Democratic Republic of Congo, Sudan, , Ethiopia, Uganda, , , Malawi, , Mozambique, , South Africa, Swaziland, . 2 Habitat: Forest and heavy woodland (savanna), especially along river courses. Also in disturbed areas in the forest zone (Larsen, 2005a). In Tanzania ssp. inalcesta occurs from sea-level to 1 850 m (Kielland, 1990d). In Madagascar in forest (Lees et al., 2003). Habits: A very common and ubiquitous forest (Larsen, 2005a). Both sexes keep to the forest understorey where they fly low down, in the shade, with a very weak, bouncing flight. They have been likened, when in flight, to floating pieces of tissue paper. Both sexes feed from small flowers in the forest undergrowth (Pringle et al., 1994). Larsen (1984; 1991; 2005a) recounts that in Nigeria he once saw a specimen settle on water to drink, being supported by surface tension alone, and using its closed wings as a ‘sail’. Males do not normally mud-puddle (Larsen, 1984). They are on the wing from early morning, often at first light (Larsen, 2005a). Flight period: All year but more plentiful in late summer and autumn (Pringle et al., 1994). Early stages:

Clark & Dickson, 1967: 38. [Described from material from Umhlanga Rocks, KwaZulu-Natal]. Egg. Laid singly on young shoots; pure white; 1.5 mm high by 0.4 mm in diameter; 10 longitudinal ribs, 6 of which reach the micropyle; ribs are cross-braced by 28-30 fine ribs; eclosion occurs after 4-6 days. . Exits the egg near the top and consumes the discarded shell. On hatching is 1.5 mm long and transparent water- white, the transparency, until it feeds making it difficult to see. Feeds on the edge of a leaf and grows to 3 mm in 6 days. Second instar larva greenish above and whitish below. Selects a resting place from which it ventures forth to feed. Grows to 6 mm in 3 days. Third instar larva green and shows the same habits as the previous instar. Grows to 8 mm in 3 days. Fourth instar larva green but more setose than previous instar. Grows to 12.5 mm in 3 days. Fifth (final) instar larva slightly darker green with faint whitish sub-spiracular stripe. Grows to 22 mm (male) and 23 mm (female) in 10 days. Pupa. Length 15.5-16 mm and pale green. Secured to a twig by cremastral hooks and supported by a girdle. Pupal period about 10 days. There is a succession of broods throughout the year.

Henning, Henning, Joannou, & Woodhall, 1997: 368 (photograph of final instar larva and pupa).

Final instar larva and pupa of Leptosia alcesta. Images courtesy Allison Sharp.

Larval food: brassii DC. (Capparaceae) [Joannou, in Pringle et al., 1994: 299]. DC. var. zeyheri (Turcz.) Toelken (Capparaceae) [Clark & Dickson, 1967; as C. zeyheri]. Capparis tomentosa Lam. (Capparaceae) [Henning, Henning, Joannou, & Woodhall, 1997: 368]. Maerua juncea Pax (Capparaceae) [Henning, Henning, Joannou, & Woodhall, 1997: 368]. Ritchiea species (Capparaceae) [Larsen, 2005a].

Leptosia alcesta alcesta (Stoll, [1782]) African Wood White

Papilio alcesta Stoll, [1782] in Stoll, [1780-2]. Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en America 4 [part]: 175 (29-252). Amsteldam & Utrecht.

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Leptosia alcesta alcesta. Male. Left – upperside; right – underside. Cape 3 Points, western Ghana. December 2002. R Vorgas. ABRI-2019-2948. Images M.C. Williams ex ABRI Collection.

Leptosia alcesta alcesta. Female. Left – upperside; right – underside. Putu Range, Liberia. December 2010. Safian, Zakar. ABRI-2019-2949. Images M.C. Williams ex ABRI Collection.

Type locality: “Côte de Guinée”. Distribution: Senegal, Gambia, Guinea-Bissau (Bacelar, 1949), Guinea, Liberia, Ivory Coast, Ghana, Togo, Benin (south, central), Nigeria, Cameroon, Equatorial Guinea (Bioko), Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo. Specific localities: Gambia – Fajara, Abuko, Bijilo, Pirang, Brufut, Farasutu, Sanyang, Janjanbureh Island, Tintinto, Basse (Jon Baker, pers. comm, May 2020). Guinea – Ziama (Safian et al., 2020). Liberia – Wonegizi (Safian et al., 2020). Ghana – Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – Noyau Central, Lama Forest (Fermon et al., 2001); Houeyogbe Forest (Coache & Rainon, 2016); see Coache et al., 2017. Nigeria – Oban Hills (Larsen, 2005a); Olokemeji Forest near Ibadan (Larsen, 2005a). Cameroon – Korup (Larsen, 2005a). Gabon – Alembe (Vande weghe, 2010); Lake Evaro (Vande weghe, 2010); Loa-Loa (Vande weghe, 2010); Ipassa (Vande weghe, 2010); Iguela (Vande weghe, 2010); Lope (Vande weghe, 2010).

narica Fabricius, 1793 (as sp. of Papilio). Entomologia Systematica emendata et aucta 3 (1): 187 (488 pp.) “Guinea”.

cepheus Ehrmann, 1894 (as sp. of Pseudopontia). Journal of the New York Entomological Society 2: 77 (77-78). Liberia: “Grand Sess, West Africa”.

Leptosia alcesta inalcesta Bernardi, 1959# African Wood White

Leptosia alcesta inalcesta Bernardi, 1959. Bulletin de la Société Entomologique de France 64: 33 (30-36). 4 Leptosia alcesta inalcesta Bernardi, 1959. Pringle et al., 1994: 298.

Leptosia alcesta inalcesta. Male (Wingspan 35 mm). Left – upperside; right – underside. Lekgalameetse N.R., Limpopo Province, South Africa. 1 September 2002. M. Williams. Images M.C. Williams ex Williams Collection.

Leptosia alcesta inalcesta. Female (Wingspan 40 mm). Left – upperside; right – underside. Lekgalameetse N.R., Limpopo Province, South Africa. 8 April 2006. M. Williams. Images M.C. Williams ex Williams Collection.

Type locality: [Tanzania]: “Tananika, Kilimandjaro, Neu Moschi, 800m”. Paratype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Sudan (south), Uganda, Ethiopia, Democratic Republic of Congo (Kivu, Ituri, Shaba), Kenya, Tanzania, Malawi, Zambia (mainly north), Mozambique, Zimbabwe (east), South Africa (Limpopo Province, Mpumalanga, KwaZulu-Natal), Swaziland. Specific localities: Uganda – Semuliki N.P. (Davenport & Howard, 1996). Democratic Republic of Congo – Semuliki Valley (Ducarme, 2018). Kenya – widespread (Larsen, 1991c). Tanzania – Kilimanjaro (TL); in all parts of the country with suitable (forest) habitat (Kielland, 1990d); Semdoe Forest Reserve (Doggart et al., 2001); Katavi National Park (Fitzherbert et al., 2006). Malawi – Zomba Mountain (Congdon et al., 2010). Zambia – Mwinilunga (Heath et al., 2002); Ikelenge (Heath et al., 2002); mid-Lunga River (Heath et al., 2002); Ndola (Heath et al., 2002); Mufulira (Heath et al., 2002); Victoria Falls (Heath et al., 2002); Lubwe on Lake Bangweulu (Heath et al., 2002); Lake Mweru (Heath et al., 2002). Mozambique – Maputo; Mt Mabu (Congdon et al., 2010); Mt Mecula [-12.0772 37.6297] (Congdon & Bayliss, 2013). Zimbabwe – Dichwe Farm near Chinhoyi (Mullin). Limpopo Province – Lekgalameetse Nature Reserve (“Malta Forest”) (Swanepoel, 1953); Woodbush (Swanepoel, 1953); Mokeetzi (Swanepoel, 1953); Sibasa (Swanepoel, 1953). Mpumalanga – Barberton (Swanepoel, 1953); Graskop (Swanepoel, 1953); Mariepskop (Swanepoel, 1953). KwaZulu-Natal – Umkomaas (Swanepoel, 1953); Durban (Swanepoel, 1953); Verulam (Swanepoel, 1953); Pinetown (Swanepoel, 1953); Eshowe (Swanepoel, 1953); St. Lucia Bay (Swanepoel, 1953); Lebombo Mountains (Swanepoel, 1953); Umhlanga Rocks (Clark & Dickson, 1967); Kosi Bay Nature Reserve (Pringle & Kyle, 2002); Tembe Nature Reserve (Pringle & Kyle, 02); Ndumo Nature Reserve (Pringle & Kyle, 2002). 5 Swaziland – Mlawula N. R. (www.sntc.org.sz); Malolotja N. R. (www.sntc.org.sz).

Leptosia alcesta pseudonuptilla Bernardi, 1959 Ethiopia African Wood White

Leptosia alcesta pseudonuptilla Bernardi, 1959. Bulletin de la Société Entomologique de France 64: 35 (30-36).

Leptosia alcesta pseudonuptilla. Male. Left – upperside; right – underside. Near Mbogi, Ituri, Democratic Republic of Congo. 1 42 N, 30 07 E. 18 March 2017. T. Desloges. Images courtesy T. Desloges.

Type locality: Ethiopia: “Ethiopie, Haute-Orguessa”. Paratype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Democratic Republic of Congo (east Kivu, north-east Ituri), to Ethiopia. Specific localities: Ethiopia – Haute-Orguessa (TL).

Leptosia alcesta sylvicola (Boisduval, 1833) Malagasy African Wood White

Leucophasia sylvicola Boisduval, 1833. Nouvelles Annales du Muséum d’Histoire Naturelle, Paris 2: 168 (149-270).

Leptosia alcesta sylvicola. Male. Left – upperside; right – underside. Zombitse, Madagascar. 16-17 April 2018. M. Williams. Images M.C. Williams ex Dobson Collection.

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Leptosia alcesta sylvicola. Female. Left – upperside; right – underside. Zombitse, Madagascar. 16-17 April 2018. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Madagascar: “dans les bois à Madagascar”. Distribution: Madagascar.

Note: Larsen (2005a: 103) believes that, because of its large size, sylvicola may not be conspecific with L. alcesta.

*Leptosia bastini Hecq, 1997 Congo Wood White

Leptosia bastini Hecq, 1997. Lambillionea 97(4) (Tome I): 482 (482-484).

Leptosia bastini. Male. Left – upperside; right – underside. Mbaiki, Lobaye, Central African Republic. October 1994. SCC. ABRI-2019-2955. Images M.C. Williams ex ABRI Collection.

Leptosia bastini. Female. Left – upperside; right – underside. Mbaiki, Lobaye, Central African Republic. December 1994. SCC. ABRI-2019-2956. Images M.C. Williams ex ABRI Collection.

Type locality: Central African Republic: “R.C.A. Ft de N’Gotto, 20.II.1996. (Didier Bastin leg.).”

7 Holotype in MRAC, Tervuren. Distribution: Cameroon (south-east), Central African Republic. Specific localities: Central African Republic – Ft de N’Gotto (TL); Pama Quelle-Baiki (Gaede, 1916). Habitat: Forest. Early stages: Nothing published. Larval food: Nothing published.

Note: Larsen (2005a: 104) says that bastini might be a subspecies of L. marginea (Mabiile, 1890).

fuscolimbata Gaede, 1916 (as var. of ). Internationale Entomologische Zeitschrift 9: 105 (105-106, 109-112, 125-126). “Pama Quelle-Baiki” [Central African Republic].

* Bernardi, 1952 PlainWood White

Leptosia hybrida Bernardi, 1952. Bulletin de la Société Entomologique de Mulhouse 1952: 12 (12).

Leptosia hybrida hybrida. Male. Left – upperside; right – underside. Cape 3 Points, western Ghana. December 2002. R Vorgas. ABRI-2019-2951. Images M.C. Williams ex ABRI Collection.

Leptosia hybrida hybrida. Male. Left – upperside; right – underside. Ipassa, Ivindo N.P., Gabon. 14 June 2016. J. Dobson. Images M.C. Williams ex Dobson collection.

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Leptosia hybrida hybrida. Female. Left – upperside; right – underside. Kongu, Ivindo N.P., Gabon. 23 November 2017. J. Dobson. Images M.C. Williams ex Dobson Collection.

Alternative common name: Hybrid Spirit. Type locality: Guinea: “Mt. Nimba, Guinée française”. Distribution: Guinea, Guinea-Bissau (Bivar de Sousa et al., 2007), , Liberia, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo, Uganda, Kenya, Tanzania, Zambia. Habitat: Primary and riparian forest, including riverine vegetation that penetrates moist savanna. Also in somewhat degraded forest (Larsen, 2005a). In Tanzania ssp. somereni occurs at altitudes from 800 to 1 300 m (Kielland, 1990d). Habits: A common forest species. The flight is weak but stronger than in other members of the genus. It spends most of its time flying around in the shady forest understorey but is often found visiting flowers along forest roads (Larsen, 1991c). Early stages: Nothing published. Larval food: Capparis species (Capparaceae) [Van Someren, 1974: 316].

Leptosia hybrida hybrida Bernardi, 1952 PlainWood White

Leptosia hybrida Bernardi, 1952. Bulletin de la Société Entomologique de Mulhouse 1952: 12 (12).

Leptosia hybrida hybrida. Male. Left – upperside; right – underside. Cape 3 Points, western Ghana. December 2002. R Vorgas. ABRI-2019-2951. Images M.C. Williams ex ABRI Collection.

Leptosia hybrida hybrida. Male. Left – upperside; right – underside. Ipassa, Ivindo N.P., Gabon. 14 June 2016. J. Dobson. Images M.C. Williams ex Dobson collection.

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Leptosia hybrida hybrida. Female. Left – upperside; right – underside. Kongu, Ivindo N.P., Gabon. 23 November 2017. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Guinea: “Mt. Nimba, Guinée française”. Distribution: Guinea, Guinea-Bissau (Bivar de Sousa et al., 2007), Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (south), Nigeria, Cameroon, Equatorial Guinea (Bioko), Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo (except east). Specific localities: Guinea – Nimba Mountains (TL). Ivory Coast – Man (Dufrane, 1947). Ghana – Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – see Coache et al., 2017. Cameroon – Korup (Larsen, 2005a). Equatorial Guinea – Sampaka, Bioko (Martin, 2015). Gabon – Ipassa (Vande weghe, 2010); Kongou (Vande weghe, 2010); Waka (Vande weghe, 2010); Kinguele (Vande weghe, 2010); Bakouaka (Vande weghe, 2010). Democratic Republic of Congo – Binga (Dufrane, 1947).

houzeaui Dufrane, 1947 (as f. of Leptosia medusa). Bulletin et Annales de la Société Royale Entomologique de Belgique 83: 46 (46-73). Ivory Coast: “Man, Côte d’Ivoire”.

reducta Dufrane, 1947 (as ab. of Leptosia medusa). Bulletin et Annales de la Société Royale Entomologique de Belgique 83: 46 (46-73). Ivory Coast: “Man, Côte d’Ivoire”.

submarginata Dufrane, 1947 (as ab. of Leptosia medusa). Bulletin et Annales de la Société Royale Entomologique de Belgique 83: 46 (46-73). Democratic Republic of Congo: “Binga, Congo”.

Leptosia hybrida somereni Bernardi, 1959 Bufambo PlainWood White

Leptosia hybrida somereni Bernardi, 1959. Revue Française d’Entomologie 26: 227 (227-231).

Leptosia hybrida somereni. Male (Wingspan: 43mm). Left – upperside; right – underside. Near Mbogi, Ituri, Democratic Republic of Congo. 1 42 N, 30 07 E. 10 June 2017. T. Desloges. 10 Images courtesy T. Desloges.

Leptosia hybrida somereni. Female. Left – upperside; right – underside. Wingspan: 45mm. Zambezi Bridge, Zambia. 24 March, 1981. A.J. & M.W. Gardiner. (Gardiner Collection).

Type locality: Uganda: “Bufambo Forest, W. Elgon, Ouganda”. Diagnosis: Differs from the nominate subspecies in the following respects: smaller; forewing upperside apical patch narrower and subapical black spot absent (Kielland, 1990d). Distribution: Democratic Republic of Congo (north-east), Uganda, Kenya (west), Tanzania, Zambia. Specific localities: Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Central Forest Block (Ducarme, 2018); Mt Blue (Ducarme, 2018). Uganda – Bufambo Forest (TL). Tanzania – Rovuhu River in Ngara District (Kielland, 1990d); Kemfu Forest (single female) (Kielland, 1990d). Zambia – Ikelenge (Heath et al., 2002); Zambezi Bridge, Ikelenge (male and female illustrated above).

*Leptosia marginea (Mabille, 1890) Black-edged Wood White

Pontia marginea Mabille, 1890. Annales de la Société Entomologique de France (6) 10: 27 (17-51).

Leptosia marginea. Male. Left – upperside; right – underside. Near Mbogi, Ituri, Democratic Republic of Congo. 1 42 N, 30 07 E. 3 December 2016. T. Desloges. Images courtesy T. Desloges.

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Leptosia marginea. Female. Left – upperside; right – underside. Maan, southern Cameroon. November 2008. PA. ABRI-2019-2950. Images M.C. Williams ex ABRI Collection.

Alternative common name: Black-edged Spirit. Type locality: [Ivory Coast]: “Assinie”. Distribution: Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (south), Nigeria (south), Cameroon, Equatorial Guinea (Bioko), Gabon, Congo, Central African Republic, Democratic Republic of Congo, Uganda (Davenport, 1996), ?Tanzania. Specific localities: Ivory Coast – Assinie (TL). Cameroon – Kitta (Aurivillius, 1895); Ekundu (Aurivillius, 1895); N’Dian (Aurivillius, 1895); Bibundi (Aurivillius, 1895); Korup (Larsen, 2005a). Equatorial Guinea (Bioko) – Caldera de Luba, Bioko (Martin, 2015); Río Riaco (Martin & Cobos, 2016). Gabon – Malibe (Vande weghe, 2010); Alen Nkoma (Vande weghe, 2010); Kinguele (Vande weghe, 2010); Waka (Vande weghe, 2010); Ipassa (Vande weghe, 2010); Kongou (Vande weghe, 2010); Langoue (Vande weghe, 2010); Nouna (Vande weghe, 2010); Bakouaka (Vande weghe, 2010). Central African Republic – Dzanga (Noss, 1998). Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Mt Mitumba (Ducarme, 2018). Habitat: Primary forest. Habits: A local and relatively scarce species of undisturbed forest (Larsen, 2005a). Early stages: Nothing published. Larval food: Capparis species (Capparaceae) [Van Someren, 1974: 316].

immaculata Aurivillius, 1895 (as var. of Nychitona medusa). Entomologisk Tidskrift 16: 257 (195-220, 255-268). Cameroon: “Kitta, Ekundu, N’Dian, Bibundi”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/m).

*Leptosia medusa (Cramer, 1777) Dainty Wood White

Papilio medusa Cramer, 1777. Die Uitlandsche Kapellen voorkomende in de drie waereld-deelen Asia, Africa en America 2: 86 (151 pp.). Amsteldam & Utrecht.

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Leptosia medusa. Female. Left – upperside; right – underside. Cape 3 Points, western Ghana. December 2002. R Vorgas. ABRI-2019-2952. Images M.C. Williams ex ABRI Collection.

Alternative common name: Dainty Spirit. Type locality: [West Africa]: “Côte de Bengale”. [False locality.] Larsen (2005a) states that the type locality is probably Sierra Leone. Distribution: Guinea-Bissau (Larsen, 2005a), Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (south, central), Nigeria (west), Equatorial Guinea (Bioko) (Martin, 2015). Recorded, in error, from Uganda by Davenport, 1996. Specific localities: Guinea – Ziama (Safian et al., 2020). Liberia – Wologizi (Safian et al., 2020); Wonegizi (Safian et al., 2020). Ivory Coast – Yapo (Larsen, 2005a). Ghana – Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – Houeyogbe Forest (Coache & Rainon, 2016); see Coache et al., 2017. Nigeria – Lagos district (Larsen, 2005a); Okomu (Wojtusiak, vide Larsen, 2005a). Habitat: Forest. Habits: Widespread and common in West Africa, in localities with a complete vegetation canopy (Larsen, 2005a). Larsen (2005a) recounts that he once noted a string of ten males chasing each other for several minutes, with not more than ten centimetres between each individual. Early stages: Nothing published. Larval food: Nothing published.

dorothea Fabricius, 1793 (as sp. of Papilio). Entomologia Systematica emendata et aucta 3 (1): 194 (488 pp.) “India”. [False locality.]

empeda Godart, 1819 in Latreille & Godart, [1819], [1824] (as sp. of Pieris). Encyclopédie Méthodique. Histoire Naturelle [Zoologie] 9 Entomologie: 139 (1-328 [1819], 329-828 [1824]). Paris. [West Africa]: “Bengale”. [False locality.]

* (Butler, 1873) Immaculate Wood White

Nychitona nupta Butler, 1873. Cistula Entomologica 1: 175 (150-177). Leptosia nupta (Butler, 1873). Pringle et al., 1994: 299.

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Leptosia nupta nupta. Male. Left – upperside; right – underside. Ivindo N.P., Gabon. 26 November 2017. J. Dobson. Images M.C. Williams ex Dobson Collection.

Leptosia nupta viettei. Female. Left – upperside; right – underside. La Mandraka, Madagascar. December 2002. ABRI Coll. ABRI-2019-2954. Images M.C. Williams ex ABRI Collection.

Alternative common names: Petite Wood White; Immaculate Spirit. Type locality: Angola: “Bembe Mines”. Identification: Similar to Leptosia alcesta but smaller; wings more translucent; lacks discal spot on forewing upperside (Pringle et al., 1994). Distribution: Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Angola, Democratic Republic of Congo, Uganda, , Kenya, Tanzania, Zambia, Zimbabwe, Madagascar. Recorded, in error, from Ghana by Berger (1981) (Larsen, 2005a). Believed to be extinct in Guinea-Bissau (Bivar-de-Sousa et al., 2016). Habitat: Wet, primary forest. In Madagascar in forest and forest margins (Lees et al., 2003). Habits: More localized than other members of the genus but where found it may be abundant (Larsen, 2005a). Flight feeble and keeps to the shade (Kielland, 1990d). In Tanzania from 800 to 1 600 m altitude (Kielland, 1990d). Early stages: Nothing published. Larval food: Capparis species (Capparaceae) [Kielland, 1990d: 71]. Rorippa madagascariensis (DC.) Hara (Brassicaceae) [Heath et al., 2002: 35].

Leptosia nupta nupta (Butler, 1873) Immaculate Wood White

Nychitona nupta Butler, 1873. Cistula Entomologica 1: 175 (150-177).

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Leptosia nupta nupta. Male. Left – upperside; right – underside. Ivindo N.P., Gabon. 26 November 2017. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Angola: “Bembe Mines”. Distribution: Nigeria (east), Cameroon, Equatorial Guinea (Bioko), Gabon, Congo, Angola, Zambia. Nominate ssp. recorded, in error, from Uganda by Davenport, 1996. Believed to be extinct in Guinea-Bissau (Bivar-de-Sousa et al., 2016). Specific localities: Nigeria – Omo Forest (Larsen, 2005a); Abo Abo in Okwangwo (Larsen, 2005a). Equatorial Guinea (Bioko) – Caldera de Luba, Bioko (Martin, 2015); Campamento UPM-Campamento Hormiga (Martin & Cobos, 2016). Cameroon – Nyong R., just south of Makak station (Birket-Smith, 1960); Korup (Larsen, 2005a). Gabon – Tchimbele (Vande weghe, 2010); Franceville (Vande weghe, 2010). Angola – Bembe Mines (TL). Zambia – Ikelenge (Heath et al., 2002).

nuptilla Aurivillius, 1910 in Seitz, 1908-25 (as f. of Leptosia alcesta). Die Gross-Schmetterlinge der Erde, Stuttgart (2) 13 Die Afrikanischen Tagfalter: 31 (614 pp.). “Ruwenzori Mts”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/a).

pygmaea Birket-Smith, 1960 (as f. of Leptosia medusa medusa). Bulletin de l’Institut Français d’Afrique Noire (A) 22: 541 (521-554, 924-983, 1259-1284). Cameroon: “near the river Nyong just south of Makak station, French ”.

Leptosia nupta pseudonupta Bernardi, 1959 Central Immaculate Wood White

Leptosia nupta pseudonupta Bernardi, 1959. Bulletin de la Société Entomologique de France 64: 35 (30-36). Leptosia nupta pseudonupta Bernardi, 1959. Pringle et al., 1994: 299.

Leptosia nupta pseudonupta. Male (Wingspan 34 mm). Left – upperside; right – underside. Mpanga Forest, Uganda. 13 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [Democratic Republic of Congo]: “Congo belge, Stanleyville”.

15 Distribution: Democratic Republic of Congo, Uganda, Rwanda, Kenya (west), Tanzania (west), Zambia, Zimbabwe (Vumba and Mutare – only two specimens). Specific localities: Democratic Republic of Congo – Stanleyville (TL); Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Central Forest Block (Ducarme, 2018); Mt Mitumba (Ducarme, 2018); Mt Blue (Ducarme, 2018). Uganda – Semuliki N.P. (Davenport & Howard, 1996). Kenya – Kakamega Forest (Larsen, 1991c; male illustrated above). Tanzania – Ntakatta Forest (Kielland, 1990d); Kigoma District and adjoining areas of Mpanda to Ngara District and Munziru Forest north of Bukoba (Kielland, 1990d). Zimbabwe – Vumba (E. W. Lannin, March 1928; single record); Mutare (Carcasson, August 1946; single record). These two records may indicate that these specimens were strays from further north. Paré averred that they might be extreme dry season forms of Leptosia alcesta (Pringle et al., 1994).

Leptosia nupta viettei Bernardi, 1959 Malagasy Immaculate Wood White

Leptosia nupta viettei Bernardi, 1959. Bulletin de la Société Entomologique de France 64: 31 (30-36).

Leptosia nupta viettei. Male. Left – upperside; right – underside. Ranamofana, Madagascar. 23 March 2016. J. Dobson. Images M.C.Williams ex Dobson Collection.

Leptosia nupta viettei. Female. Left – upperside; right – underside. La Mandraka, Madagascar. December 2002. ABRI Coll. ABRI-2019-2954. Images M.C. Williams ex ABRI Collection.

Type locality: Madagascar: “Madagascar centre, Massif de l’Ankaratra, forêt d’Ambahona, Manjakatompo”. Distribution: Madagascar. Specific localities: Madagascar – Ankaratra (TL); Ambahona Forest (Bernardi, 1959); Manjakatompo (Bernardi, 1959).

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*Leptosia wigginsi (Dixey, 1915) Opaque Wood White

Opaque Wood White (Leptosia wigginsi wigginsi ). Image courtesy Jeremy Dobson.

Nychitona medusa wigginsi Dixey, 1915. Transactions of the Entomological Society of London 1915: 7 (1-15).

Leptosia wigginsi wigginsi. Male (Wingspan 43 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 15 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Leptosia wigginsi wigginsi. Female. Left – upperside; right – underside. Mapimbi, North Kivu, Democratic Republic of Congo. August 2009. ABRI Leg. ABRI-2019-2953. Images M.C. Williams ex ABRI Collection.

Alternative common name: Opaque Spirit. Type locality: Uganda: “Toro country on the eastern slopes of Ruwenzori; Entebbe”; Kenya: “Ugaia country, south of Kavirondo Gulf”; “Victoria Nyanza”. Distribution: Senegal, Guinea-Bissau (Larsen, 2005a), Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (Fermon et al., 2002), Nigeria, Cameroon, Equatorial Guinea, Gabon, Angola (Mendes et al., 2018), Central African Republic, Democratic Republic of Congo, Uganda, Kenya, Tanzania. Habitat: Primary lowland forest. In West Africa it occurs in both ‘wet’ and ‘dry’ forests (Larsen, 2005a). In Tanzania the nominate subspecies is found at altitudes from 800 to 1 200 m (Kielland, 17 1990d). Habits: In West Africa it is the rarest species of its genus (Larsen, 2005a). Keeps to shady places in the forest (Kielland, 1990d). Habits similar to those of L. hybrida (Larsen, 1991c). Early stages: Nothing published. Larval food: Capparis species (Capparaceae) [Van Someren, 1974: 316].

Leptosia wigginsi wigginsi (Dixey, 1915) Opaque Wood White

Nychitona medusa wigginsi Dixey, 1915. Transactions of the Entomological society of London 1915: 7 (1-15).

Leptosia wigginsi wigginsi. Male (Wingspan 43 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 15 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Leptosia wigginsi wigginsi. Male (Wingspan 43 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 15 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Leptosia wigginsi wigginsi. Female. Left – upperside; right – underside. Mapimbi, North Kivu, Democratic Republic of Congo. August 2009. ABRI Leg. ABRI-2019-2953. Images M.C. Williams ex ABRI Collection.

Type locality: Uganda: “Toro country on the eastern slopes of Ruwenzori; Entebbe”; Kenya: “Ugaia 18 country, south of Kavirondo Gulf”; “Victoria Nyanza”. Distribution: Democratic Republic of Congo (east), Uganda, Kenya (south-west), Tanzania (west). Specific localities: Democratic Republic of Congo – Ituri Forest (Ducarme, 2018). Uganda – Ruwenzori Mtns (TL); Entebbe (Dixey, 1915); Semuliki N.P. (Davenport & Howard, 1996). Kenya – Ugaia (Dixey, 1915). Tanzania – From Luegele River to Malagarazi River in Kigoma (Kielland, 1990d); adjoining areas in Mpanda (Kielland, 1990d).

uganda Neustetter, 1927 (as sp. of Leptosia). Internationale Entomologische Zeitschrift 21: 7 (7-8, 14- 16, 23-24, 32). Uganda.

Leptosia wigginsi pseudalcesta Bernardi, 1965 Western Opaque Wood White

Leptosia wigginsi pseudalcesta Bernardi, 1965. Bulletin de la Société Entomologique de France 69: 260 (258-260).

Type locality: Senegal: “Sénégal, M’Boa”. Paratype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Senegal, Guinea-Bissau (Larsen, 2005a), Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (south), Nigeria, Cameroon, Equatorial Guinea (Bioko), Gabon, Angola, Central African Republic, Democratic Republic of Congo (most of). Specific localities: Senegal – M’Boa (TL); the ‘niyaes’ near Dakar (Larsen, 2005a); Basse Casamance (Larsen, 2005a). Ghana – Kissi Forest (Larsen, 2005a); Shai Hills (Larsen, 2005a); Bia National Park (Larsen, 2005a); Krokosua Hills (Larsen, 2005a); Bobiri Butterfly Sanctuary (Larsen et al., 2007). Benin – Noyau Central, Lama Forest (Fermon et al., 2001); Houeyogbe Forest (Coache & Rainon, 2016); see Coache et al., 2017. Nigeria – Isheri near Lagos (Larsen, 2005a); Okwangwo (Larsen, 2005a); Oban Hills (Larsen, 2005a). Equatorial Guinea – Biapa (Conception), Bioko (Martin, 2015). Gabon – Masouna 2000 (Vande weghe, 2010); Lope (Vande weghe, 2010).

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