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

Genus Butler, 1870 Pathfinders

Entomologist’s Monthly Magazine 7: 96 (55-58, 92-99). Type-species: Papilio edipus Stoll, by original designation.

Pardaleodes sator pusiella. Image courtesy Jeremy Dobson.

The genus Pardaleodes belongs to the Family Hesperiidae Latreille, 1809; Subfamily Hesperinae Latreille, 1809; Tribe Ceratrichiini Grishin, 2019. The other genera of the Tribe Ceratrichiini in the Afrotropical Region are Ceratrichia, Ceratricula, Meza, Herila, Argemma and Ankola.

Pardaleodes (Pathfinders) is an Afrotropical genus of five species, occurring in forest.

*Pardaleodes bule Holland, 1896 Pearl-spotted Pathfinder

Pardaleodes bule Holland, 1896. Proceedings of the Zoological Society of London 1896: 76 (2-107).

Pardaleodes bule. Male. Left – upperside; right – underside. Kakamega Forest, . 7 October 1997. Images M.C.Williams ex J. Greyling Collection.

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Pardaleodes bule. Female. Left – upperside; right – underside. Kibale Forest, . 22 October 2014. J. Dobson. Images M.C.Williams ex Dobson Collection.

Type locality: : “Bule country, ”. Distribution: Cameroon, Equatorial (Island of Bioko), , Congo, Central African Republic, Democratic Republic of Congo, Uganda, Kenya (west), (west), Macias Nguema Island (Kielland, 1990d). Specific localities: Cameroon – Bule country (TL). Equatorial Guinea – San Carlos, Bioko (Aurivillius, 1925). Gabon – Ogove River (Vande weghe, 2010). Central African Republic – Dzanga (Noss, 1998). Democratic Republic of Congo – Makala (Bethune-Baker, 1908); Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Mt Mitumba (Ducarme, 2018). Kenya – Kakamega Forest (Cock & Congdon, 2014). Tanzania – Kasoge Forest (Kielland, 1990d); Gombe Stream (Kielland, 1990d). Habitat: Open places in lowland evergreen forest at 800 to 1 000 m (Kielland, 1990d). Habits: Common in the west of its range, rarer in the east (Larsen, 1991c). Occurs in very localized colonies, specimens settling on low vegetation (Kielland, 1990d). Usually rest with closed wings but sometimes sunbathe with the hind wings three quarters open, and the fore wings only slightly open. They feed readily from flowers, for example those of Justicia flava (Cock & Congdon, 2014). Early stages:

Cock & Congdon, 2014: 317. [Kakamega Forest, Kenya]. These authors describe the leaf shelters, penultimate instar larva, final instar larva and pupa. They illustrate leaf shelters and final instar larvae.

Larval food: Setaria megaphylla [Cock & Congdon, 2014: 317; Kakamega, Kenya]. Setaria sulcata (=poiretiana) [Cock & Congdon, 2014: 317; Kakamega, Kenya].

makala Bethune-Baker, 1908 (as sp. of Pandaleodes [sic]). Annals and Magazine of Natural History (8) 2: 480 (469-482). [Democratic Republic of Congo]: “Makala”.

schultzei Aurivillius, 1925 (as ab. female of Pardaleodes reichenowi). Ergebnisse der Zweiten Deutschen Zentral-Afrika-Expedition 1 (18): 1247 (1243-1359). Equatorial Guinea: “Fernando Poo; San Carlos”.

*Pardaleodes edipus (Stoll, [1781]) Common Pathfinder

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Common Pathfinder (Pardaleodes edipus). Left – male; right – female. Male: Boabeng, . Image courtesy Pamela Sai Female: Yangambi, DRC. Image courtesy Bart Wursten.

Papilio edipus Stoll, [1781] in Stoll, [1780-2]. Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en America 4 [part]: 146 (29-252). Amsteldam & Utrecht. Pardaleodes edipus (Stoll, 1781). Evans, 1937.

Pardaleodes edipus. Male. Left – upperside; right –underside. Ipassa, Gabon. 19 June 2016. J. Dobson. Images M.C. Williams ex Dobson collection.

Pardaleodes edipus. Female (Wingspan 26 mm). Left – upperside; right – underside. Biakpa Mountain Paradise, Ghana. 24 November 2011. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [West Africa]: “Kaap de Goede Hoop”. [False locality.] Distribution: , Guinea-Bissau (Bivar-de-Sousa & Mendes, 1999), Guinea, , Liberia, , Ghana, , Benin (south, central), , Cameroon, Equatorial Guinea (Island of Bioko), Gabon, , Congo, Central African Republic, Democratic Republic of Congo. Erroneously recorded from South Africa by Trimen, 1866a (as Pamphila edipus Cramer) (MCW). Recorded, probably in error, from Gambia by Evans, 1937 (Larsen, 2005a). Specific localities: Senegal – Basse Casamance (Larsen, 2005a; single record). Guinea – Fouta Djalon (Larsen, 2005a); Ziama (Safian et al., 2020). Ghana – Bobiri Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen

3 et al., 2009). Benin – Lokoli (Tchibozo et al., 2008); Houeyogbe Forest (Coache & Rainon, 2016); see Coache et al., 2017. Gabon – Mondah (Vande weghe, 2010); Nyonie (Vande weghe, 2010); Bitam (Vande weghe, 2010); Ipassa, Ivindo National Park (Vande weghe, 2010); Inkogou (Vande weghe, 2010); Akaka, Loango National Park (Vande weghe, 2010); Rabi (Vande weghe, 2010); Camp PPG, Bateke Plateau National Park (Vande weghe, 2010). Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Mt Mitumba (Ducarme, 2018). Habitat: Mainly in disturbed areas in forest. Habits: This is probably the most common skipper of the West African forest zone (Larsen, 2005a). Mostly flies on forest edges, along forest paths and in forest clearings (Larsen, 2005a). The flight is quite weak and low down, individuals often visiting flowers near the ground (Larsen, 2005a). Males come to bird-droppings (Larsen, 2005a). Early stages: Nothing published. Larval food: Grasses (Poaceae) [Larsen, vide Cock & Congdon, 2014] .

* (Snellen, 1872) Savanna Pathfinder

Pamphila incerta Snellen, 1872. Tidschrift voor Entomologie 15: 29 (1-112). Pardaleodes incerta (Snellen, 1872). Evans, 1937.

Pardaleodes incerta incerta. Male (Wingspan 27 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 13 June 2009. J. Dobson. Images M.C. Willisms ex Dobson Collection.

Pardaleodes incerta incerta. Female. Left – upperside; right – underside. Ziika Forest, Uganda. 28 June 2015. J. Dobson. Images M.C.Williams ex Dobson Collection.

Type locality: “Neder-Guinea”. [False locality] Distribution: Senegal, Guinea-Bissau (Bivar-de-Sousa & Mendes, 1999), Guinea, , Sierra Leone, Ivory Coast, Ghana, Togo, Benin (Tchibozo et al., 2008), Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Angola, Democratic Republic of Congo, , Uganda, Kenya, Tanzania, , Mozambique (Congdon et al., 2010).

4 Habitat: Transition zone between forest and savanna and riverine forest (Larsen, 2005a). In Tanzania, from sea-level to 2 000 m, but seldom above 1 700 m (Kielland, 1990d). Habits: A common species. Usually found on forest paths or in small clearings (Larsen, 1991c). Flies rather slowly for a skipper, except when males are disputing territories (Larsen, 2005a). Both sexes are readily attracted to flowers (Larsen, 2005a). Settles on low vegetation (Kielland, 1990d). Early stages: Nothing published. Larval food: (L.) Raeusch. (Poaceae) [Vuattoux, 1999; Ivory Coast].

Pardaleodes incerta incerta (Snellen, 1872) Savanna Pathfinder

Pamphila incerta Snellen, 1872. Tidschrift voor Entomologie 15: 29 (1-112). Pardaleodes incerta incerta (Snellen, 1872). Evans, 1937.

Pardaleodes incerta incerta. Male (Wingspan 27 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 13 June 2009. J. Dobson. Images M.C. Willisms ex Dobson Collection.

Pardaleodes incerta incerta. Female. Left – upperside; right – underside. Ziika Forest, Uganda. 28 June 2015. J. Dobson. Images M.C.Williams ex Dobson Collection.

Type locality: “Neder-Guinea”. [False locality]. Distribution: Angola, Democratic Republic of Congo, Sudan (south), Uganda, Kenya, Tanzania, Zambia (north-west and north-east), Mozambique. Specific localities: Angola – Bengo Province; Bie Province; Kwanza Norte Province; Kwanza Sul Province; Malanje Province; Uige Province (Mendes et al., 2013). Democratic Republic of Congo – 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). Tanzania – Kimboriani Mountains (Kielland, 1990d). Zambia – Ikelenge (Heath et al., 2002); Mwinilunga (Heath et al., 2002); 80 km south of Mwinilunga (Heath et al., 2002); Luongo River (Heath et al., 2002); Lumangwe Falls (Heath et al., 2002);

5 Kalungwishi River (Heath et al., 2002); Lufubu River (Heath et al., 2002); Hillwood Farm (male illustrated above). Mozambique – Mount Mabu (Congdon et al., 2010); Mt Yao [-12.4432 36.5114] (Congdon & Bayliss, 2013).

Note: Populations in eastern Democratic Republic of Congo, southern Sudan, Uganda, most of Kenya and Tanzania are transitional between subspecies murcia in the west and subspecies incerta in the east (Larsen, 2005a).

Pardaleodes incerta murcia (Plötz, 1883) Northern Savanna Pathfinder

Hesperia murcia Plötz, 1883. Stettiner Entomologische Zeitung 44: 229 (26-64, 195-233). Pardaleodes incerta murcia (Plötz, 1883). Evans, 1937. Synonym of Pardaleodes incerta (Snellen, 1872). Ackery et al., 1995. Pardaleodes incerta murcia (Plötz, 1883). Larsen, 2005a: 501.

Pardaleodes incerta murcia. Male. Left – upperside; right – underside. Libreville, Gabon. 12 June 2016. J. Dobson. Images M.C. Williams ex Dobson collection.

Pardaleodes incerta murcia. Left – male upperside; right – female upperside. Male – Ngom, Cameroon. Female – Bassa, Nigeria. Images courtesy Torben Larsen.

Type locality: No locality given. Distribution: Senegal, Guinea-Bissau (Bivar-de-Sousa & Mendes, 1999), Guinea, Burkina Faso, Sierra Leone, Ivory Coast, Ghana, Togo, Benin (south, central), Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo. Specific localities: Senegal – Basse Casamance (Larsen, 2005a). Guinea – Ziama (Safian et al., 2020). Ghana – Kogyae Strict Nature Reserve (Larsen, 2005a); Accra Plains (Larsen, 2005a); Shai Hills (Larsen, 2005a); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – Lokoli (Tchibozo et al., 2008); see Coache et al., 2017. Nigeria – Mkpot 1 (agricultural clearing) (Larsen, 2005a).

6 Cameroon – Korup (fringes) (Larsen, 2005a). Gabon – Mondah (Vande weghe, 2010); Pointe Pongara (Vande weghe, 2010); Alen Nkoma (Vande weghe, 2010); Kinguele (Vande weghe, 2010); Tchimbele (Vande weghe, 2010); Bitam (Vande weghe, 2010); Bakouaka (Vande weghe, 2010); Langoue, Ivindo National Park (Vande weghe, 2010); Waka National Park (Vande weghe, 2010); Rabi (Vande weghe, 2010); Gamba (Vande weghe, 2010); Camp PPG, Bateke Plateau (Vande weghe, 2010).

diluta Robbe, 1892 (as var. of Pardaleodes oedipus). Annales de la Société Entomologique de Belgique 36: 134 (132-134). “Congo”.

*Pardaleodes sator (Westwood, [1852]) Scarce Pathfinder

A male Scarce Pathfinder Skipper (Pardaleodes sator) in a Ugandan forest. Image courtesy Jeremy Dobson.

Cyclopides sator Westwood, [1852] in Doubleday & Westwood, [1846-52]. The genera of diurnal , London: 523 (1: 1-250 pp.; 2: 251-534 pp.). London. Pardaleodes sator (Westwood, 1852). Evans, 1937.

Pardaleodes sator sator. Male. Left – upperside; right – underside. Yekepah, Nimba Mountains, Liberia. 10 January 2014. J. Dobson. Images M.C. Willisms ex Dobson Collection.

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Pardaleodes sator sator. Female. Left – upperside; right – underside. Oyem, Cameroon. October 1994. SCC. ABRI-2019-2428. Images M.C. Willisms ex ABRI Collection.

Type locality: “Guinea”. Distribution: Sierra Leone, Guinea, Liberia, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo, Sudan, Uganda, Kenya, Tanzania, Zambia (Gardiner, 2010b). Habitat: Forest. In Tanzania at altitudes between 800 and 1 000 m (Kielland, 1990d). Habits: Much scarcer than Pardaleodes edipus or Pardaleodes incerta but the behaviours are similar (Larsen, 1991c; Larsen, 2005a). Usually found along paths in fairly dense forest, flying low down and settling on low vegetation (Kielland, 1990d). Early stages: Nothing published. Larval food: Oplismenus hirtellus (L.) P.Beauv. (Poaceae) [Vuattoux, 1999; Ivory Coast].

Pardaleodes sator sator (Westwood, [1852]) Scarce Pathfinder

Cyclopides sator Westwood, [1852] in Doubleday & Westwood, [1846-52]. The genera of diurnal Lepidoptera, London: 523 (1: 1-250 pp.; 2: 251-534 pp.). London. Pardaleodes sator sator (Westwood, 1852). Evans, 1937.

Pardaleodes sator sator. Male. Left – upperside; right – underside. Yekepah, Nimba Mountains, Liberia. 10 January 2014. J. Dobson. Images M.C. Willisms ex Dobson Collection.

Pardaleodes sator sator. Female. Left – upperside; right – underside. Oyem, Cameroon. October 1994. SCC. ABRI-2019-2428. Images M.C. Willisms ex ABRI Collection.

Type locality: “Guinea”. Distribution: Sierra Leone, Guinea, Liberia, Ivory Coast, Ghana, Togo, Benin (south, central), Nigeria,

8 Cameroon (west). Specific localities: Guinea – Ziama (Safian et al., 2020). Liberia – Wologizi (Safian et al., 2020). 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.

Pardaleodes sator pusiella Mabille, 1877 Southern Scarce Pathfinder

Pardaleodes pusiella Mabille, 1877. Bulletin de la Société Zoologique de France 2: 237 (214-240). Pardaleodes sator pusiella Mabille, 1877. Evans, 1937. Synonym of Pardaleodes sator (Westwood, 1852). Ackery et al., 1995. Pardaleodes sator pusiella Mabille, 1877. Larsen, 2005a: 502.

Pardaleodes sator pusiella. Male (Wingspan 28 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 14 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Pardaleodes sator pusiella. Female (Wingspan 27 mm). Left – upperside; right – underside. Mabira Forest, Uganda. 14 June 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Angola: “Congo, Landana”. Distribution: Cameroon (except west), Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo, Sudan (south), Uganda, Kenya (west), Tanzania (west), Zambia (north-west). Specific localities: Gabon – Mondah (Vande weghe, 2010); Malibe (Vande weghe, 2010); Nyonie (Vande weghe, 2010); Kinguele (Vande weghe, 2010); Lope National Park (Vande weghe, 2010); Waka (Vande weghe, 2010); Evengue, Fernan Vaz (Vande weghe, 2010); Ndougou (Vande weghe, 2010); Iguela (Vande weghe, 2010); Camp PPG, Bateke Plateau National Park (Vande weghe, 2010). Angola – Landana (TL); Cabinda Province; Kwanza Norte Province (Mendes et al., 2013). Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Mt Mitumba (Ducarme, 2018); Mt Blue (Ducarme, 2018). Uganda – Semuliki N.P. (Davenport & Howard, 1996). Kenya – Kakamega Forest (Larsen, 1991c).

9 Tanzania – Kasoge Forest (Kielland, 1990d); Gombe Stream (Kielland, 1990d). Zambia – Ikelenge district (Gardiner, 2010b).

*Pardaleodes tibullus (Fabricius, 1793) Large Pathfinder

Hesperia tibullus Fabricius, 1793. Entomologia Systematica emendata et aucta 3 (1): 326 (488 pp.). Pardaleodes tibullus (Fabricius, 1793). Evans, 1937.

Pardaleodes tibullus tibullus. Male. Left – upperside; right – underside. Tano Ofin, Ghana. Ex ABRI Collectioin. Images courtesy T.B. Larsen.

Pardaleodes tibullus torensis. Female. Left – upperside; right – underside. Kakamega Forest, Kenya. 5 October 1997. Images M.C.Williams ex J. Greyling Collection.

Type locality: [Africa]: “in Indiis”. [False locality.] Distribution: Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo, Sudan, Uganda, Kenya. Habitat: Open places in forest in reasonable condition (Larsen, 2005a). Habits: Relatively common, with a flight that is much more powerful than its congeners (Larsen, 2005a). Individuals are fond of flowers, particularly those of the Acanthaceae and Lamiaceae. Early stages:

Sourakov & Emmel, 1997. [Ghana; ssp. tibullus]. Briefly describe and illustrate the final instar larva.

Cock & Congdon, 2014: 320. [Kakamega, Kenya; ssp. torensis]. Describe the cast final instar cuticle and head capsule and pupa. Illustrate the pupa.

Larval food: Cenchrus species (Poaceae) [Larsen, 2005a; Kakum, Ghana; as Pennisetum]. Setaria megaphylla (Poaceae) [Cock & Congdon, 2014: 320; Kakamega, Kenya].

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Pardaleodes tibullus tibullus (Fabricius, 1793) Large Pathfinder

Hesperia tibullus Fabricius, 1793. Entomologia Systematica emendata et aucta 3 (1): 326 (488 pp.). Pardaleodes tibullus tibullus (Fabricius, 1793). Evans, 1937.

Pardaleodes tibullus tibullus. Male. Left – upperside; right – underside. Tano Ofin, Ghana. Ex ABRI Collectioin. Images courtesy T.B. Larsen.

Type locality: [Africa]: “in Indiis”. [False locality.] Distribution: Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo, Sudan (south). Specific localities: Guinea – Ziama (Safian et al., 2020). Liberia – Wologizi (Safian et al., 2020). Ghana – Aburi (Plötz, 1879); Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Cameroon – Bonjongo (Plötz, 1879; Eningo (Plötz, 1879); Bangwe (Reuss, 1920); Korup (Larsen, 2005a). Gabon – Alen Nkomo (Vande weghe, 2010); Ipassa, Ivindo National Park (Vande weghe, 2010); Waka National Park (Vande weghe, 2010); Lake Oguemoue (Vande weghe, 2010); Kangwe (Vande weghe, 2010). Angola – Uige Province (Mendes et al., 2013).

reichenowi Plötz, 1879 (as sp. of Plastingia). Stettiner Entomologische Zeitung 40: 357 (353-364). Ghana: “Aburi”; Cameroon: “Bonjongo”; Ivory Coast: “Eningo”. Synonymized with tibullus by Evans (1951).

festus Mabille, 1890 (as sp. of Pardaleodes). Annales de la Société Entomologique de France (6) 10: 33 (17-51). “Congo”; Gabon.

aurivillii Reuss, 1920 (as sp. of Pardaleodes). Entomologische Rundschau 37: 25 (25). Cameroon: “N. Kamerun, Bangwe”.

Pardaleodes tibullus torensis Bethune-Baker, 1906 Eastern Large Pathfinder

Pardaleodes torensis Bethune-Baker, 1906. Annals and Magazine of Natural History (7) 18: 342 (339-346). Pardaleodes tibullus torensis Bethune-Baker, 1906. Evans, 1937. Pardaleodes tibullus toroensis Bethune-Baker, 1906. Larsen, 1991c. [Misspelling of subspecies name]. Synonym of Pardaleodes tibullus (Fabricius, 1793). Ackery et al., 1995. Pardaleodes tibullus torensis Bethune-Baker, 1906. Larsen, 2005a: 502.

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Pardaleodes tibullus torensis. Male. Left – upperside; right – underside. Kakamega Forest, Kenya. Ex ABRI Collectioin. Images courtesy T.B. Larsen.

Pardaleodes tibullus torensis. Female. Left – upperside; right – underside. Kakamega Forest, Kenya. 5 October 1997. Images M.C.Williams ex J. Greyling Collection.

Type locality: [Uganda]: “Toro, E. Africa”. Distribution: Uganda, Kenya (west). Specific localities: Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Mt Mitumba (Ducarme, 2018). Uganda – Toro (TL); Semuliki N.P. (S. Forbes, pers. comm., 2017). Kenya – Kakamega Forest (Larsen, 1991c); Kabras (Larsen, 1991c).

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