AFROTROPICAL . MARK C. WILLIAMS. http://www.lepsocafrica.org/?p=publications&s=atb Updated 30 December 2020

Genus Tetrarhanis Karsch, 1893 On-offs

Berliner Entomologische Zeitschrift 38: 217 (1-266). Type-species: Liptena ilma Hewitson, by monotypy.

= Lectiles Birket-Smith, 1960. Bulletin de l’Institut Français d’Afrique Noire (A) 22: 966 (521-554, 924-983, 1259-1284). Type-species: Lectiles collitorum Birket-Smith, by original designation.

The genus Tetrarhanis belongs to the Family Leach, 1815; Subfamily Doherty, 1886; Tribe Liptenini Röber, 1892. The other genera in the Tribe Liptenini in the Afrotropical Region are Liptena, Obania, Kakumia, Falcuna, Larinopoda, Micropentila, Pseuderesia, Eresina, Eresiomera, Parasiomera, Citrinophila, Argyrocheila, Teriomima, Euthecta, Baliochila, Cnodontes, Congdonia, Eresinopsides, Toxochitona, Mimacraea and Mimeresia.

Tetrarhanis (On-offs) is a purely Afrotropical genus containing 15 species. The distribution of the genus is restricted to the central and west African rainforest zone. The flight is weak, the dark upper surface contrasting markedly with the light underside, giving an ‘on-off’ appearance when they are flying. None of the species appears to be tied to particular ‘ant-trees’, or to be associated with Crematogaster ants, being instead evenly distributed within the forests they inhabit. Females are often seen ovipositing on dead branches on the forest floor (Larsen, 2005a).

*Tetrarhanis baralingam (Larsen, 1998) Baralingam On-off

Thermoniphas baralingam Larsen, 1998. Lambillionea 98: 78 (77-80). [Appears to have inadvertently been described under the incorrect genus as a result of a printer’s error – MCW]. The captions for Tetrarhanis baralingam and Tetrarhanis okwangwo were accidentally transposed in the original description (Larsen, 2005a). Tetrarhanis baralingam (Larsen, 1998). Larsen, 2005a: 152. Tetrarhanis baralingam Larsen, 1998. d’Abrera, 2009: 660.

Type locality: Ghana: “vi. 1993 (T.B. Larsen leg., coll. Natural History Museum, London).” No specific locality given for holotype. Paratypes from various localities in Ghana. Distribution: Guinea, Sierra Leone, Liberia, Ivory Coast (Warren-Gash, pers. comm., 2002), Ghana (south). Specific localities: Guinea – Ziama (Safian et al., 2020). Sierra Leone – Guma Valley, Freetown (ABRI, vide Larsen, 2005a). Liberia – Wologizi (Safian et al., 2020). Ivory Coast – Yapo (Warren-Gash, vide Larsen, 2005a). Ghana – Kakum (Larsen, 2005a); Cape Three Points (Larsen, 2005a); Ankasa (Larsen, 2005a); Atewa Range (Aduse-Poku & Doku-Marfo, 2007). General remarks: The specific name means ‘large penis’ in Sanskrit. Habitat: Primary forest only (Larsen, 2005a). Habits: Appears to be much scarcer than Tetrarhanis symplocus (Larsen, 2005a). Early stages: Nothing published.

1 Larval food: Nothing published.

*Tetrarhanis decempunctata (Schultze, 1923)

Liptena decempunctata Schultze, 1923. In: Schultze & Aurivillius, 1923. Ergebnisse der Zweiten Deutschen Zentral-Afrika Expedition 1(17): 1187 (1113-1242). Liptena decempunctata Schultze & Aurivillius, 1923. d’Abrera, 2009: 652. [Authorship erroneous] Tetrarhanis decempunctata (Schultze, 1923). Libert, 2020. comb. nov.

Type locality: Cameroon: “Süd-Kamerun: Molundu; Boenga”. Neotype (male) designated by Libert, 2020: Sembe, district de Sounke, Congo, II.1960 (T.H.E. Jackson); Natural History Museum, London. Distribution: Cameroon, Congo. Specific localities: Cameroon – Molundu (TL); Boenga (Schultze, 1923). Congo – Sembe, Souanke district (TL of souanke, which is designated as the Neotype of decempunctata). Habitat: Primary forest. Early stages: Nothing published. Larval food: Nothing published.

souanke Stemppfer, 1962 (as sp. of Liptena). Bulletin de l’Institut Français d’Afrique Noire (A) 24: 1148 (1135-1181). Congo: “Congo ex français, district de Souanké, Sembé”. Syn. nov. for Tetrarhanis decempunctata (Libert, 2020).

*Tetrarhanis diversa (Bethune-Baker, 1904) Diverse On-off

Liptena diversa Bethune-Baker, 1904. Annals and Magazine of Natural History (7) 14: 225 (222-233). Tetrarhanis diversa Bethune-Baker, 1904. d’Abrera, 2009: 659.

Type locality: Sierra Leone: “Sierra Leone”. Distribution: Guinea, Sierra Leone, Liberia, Ivory Coast. Records of this species from central and eastern Africa refer to T. ilala (Larsen, 2005a). Specific localities: Guinea – Fouta Djalon (Larsen, 2005a); Labe (Larsen, 2005a). Ivory Coast – Issia (Larsen, 2005a); Yeale (Larsen, 2005a); Banco (Larsen, 2005a); Tai Forest (Larsen, 2005a). Habitat: Primary forest. Habits: A rare and local species (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

*Tetrarhanis ilala (Riley, 1929)

Liptena diversa ilala Riley, 1929. In Eltringham et al., 1929. Transactions of the Entomological Society of London 77: 493 (475-504). Tetrarhanis ilala Riley, 1929. d’Abrera, 2009: 660.

Type locality: Sudan/Uganda: “Kaia River, Sudan-Uganda frontier”. Distribution: Cameroon, Gabon, Congo, Angola (Mendes et al., 2018), Central African Republic, Democratic Republic of Congo, Sudan, Uganda, Kenya. Habitat: Primary forest. Habits: Nothing published. Early Stages: Nothing published.

2 Larval food: Nothing published.

Tetrarhanis ilala ilala (Riley, 1929)

Liptena diversa ilala Riley, 1929. In Eltringham et al., 1929. Transactions of the Entomological Society of London 77: 493 (475-504). Tetrarhanis ilala ilala Riley, 1929. d’Abrera, 2009: 660.

Type locality: Sudan/Uganda: “Kaia River, Sudan-Uganda frontier”. Distribution: Sudan (south), Uganda, Kenya (west). Specific localities: Sudan – Kaia River (TL). Uganda – Semuliki N.P. (S. Forbes, pers. comm., 2015; misidentified as T. diversa). Kenya – Kakamega Forest (Larsen, 1991c).

Tetrarhanis ilala etoumbi (Stempffer, 1964)

Liptena (Tetrarhanis) etoumbi Stempffer, 1964. Bulletin de l’Institut Français d’Afrique Noire (A) 26: 1236 (1226-1287). Tetrarhanis ilala etoumbi Stempffer, 1964. d’Abrera, 2009: 660.

Type locality: Congo: “Moyen-Congo ex français, Etoumbi”. Distribution: Cameroon, Gabon, Angola (Mendes et al., 2018), Congo, Central African Republic, Democratic Republic of Congo. Specific localities: Gabon – Lope N.P. (zone SEGC) (Vande weghe, 2010); camp Nouna (Vande weghe, 2010). Congo – Etoumbi (TL).

*Tetrarhanis ilma (Hewitson, [1873])

Liptena ilma Hewitson, [1873]. In: Hewitson, [1872-7]. Illustrations of new species of exotic butterflies 5: 86 (127 pp.). London. Tetrarhanis ilma Hewitson, 1873. d’Abrera, 2009: 658.

Tetrarhanis ilma ilma. Male. Left – upperside; right – underside. Libreville, Gabon. 29 November 2017. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Angola: “Angola”. Distribution: Gabon, Congo, Democratic Republic of Congo, Angola, Uganda, Kenya, Tanzania. Recorded, erroneously, from West Africa (Larsen, 2005a). Habitat: Primary forest. Habits: Flight slow; frequently settles on bare twigs (Kielland, 1990d). Specimens keep to the shade and roost in small groups, on bare twigs. In flight the dark upperside and white underside gives the

3 a distinctive, pulsating appearance (Larsen, 1991c). Early stages: Nothing published. Larval food: Nothing published.

Tetrarhanis ilma ilma (Hewitson, [1873])

Liptena ilma Hewitson, [1873]. In: Hewitson, [1872-7]. Illustrations of new species of exotic butterflies 5: 86 (127 pp.). London. Tetrarhanis ilma ilma Hewitson, 1873. d’Abrera, 2009: 658.

Tetrarhanis ilma ilma. Male. Left – upperside; right – underside. Libreville, Gabon. 29 November 2017. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Angola: “Angola”. Distribution: Gabon, Congo, Democratic Republic of Congo (Mayumbe, Uele, Ituri, Tshopo, Equateur, Kinshasa, Sankuru, Lualaba, Shaba), Angola. Specific localities: Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Central Forest Block (Ducarme, 2018); Mt Mitumba (Ducarme, 2018).

Tetrarhanis ilma daltoni (Poulton, 1929)

Liptena ilma daltoni Poulton, 1929. In Eltringham et al., 1929. Transactions of the Entomological Society of London 77: 493 (475-504). Tetrarhanis ilma daltoni Poulton, 1929. d’Abrera, 2009: 658.

Type locality: [Tanzania]: “forest near Kakindu Hill, Tang. Terr.”. Diagnosis: Distinguished from other subspecies, in the forewing upperside, by the prominent white discal area in both sexes (Congdon & Collins, 1998). Distribution: Democraic Republic of Congo, Uganda (south-west), Tanzania (north-west). Specific localities: Democratic Republic of Congo – Mt Mitumba (Ducarme, 2018). Tanzania – Kakindu Hill (TL); Minziro Forest (common) (Congdon & Collins, 1998).

Tetrarhanis ilma lathyi (Joicey & Talbot, 1921)

Liptena ilma lathyi Joicey & Talbot, 1921. Bulletin of the Hill Museum, Witley 1: 82 (40-166). Tetrarhanis ilma lathyi Joicey & Talbot, 1921. d’Abrera, 2009: 658.

Type locality: [Democratic Republic of Congo]: “Rutshuru River, N. Kivu”. Distribution: Democratic Republic of Congo (Kivu), Tanzania. Specific localities:

4 Democratic Republic of Congo – Rutshuru River (TL). Tanzania – Kasoge Forest (at the foot of Mount Mahale, Kigoma District) (Kielland, 1990d).

Tetrarhanis ilma ugandae (Stempffer, 1964)

Liptena (Tetrarhanis) ilma ugandae Stempffer, 1964. Bulletin de l’Institut Français d’Afrique Noire (A) 26: 1234 (1226- 1287). Tetrarhanis ilma ugandae Stempffer, 1964. d’Abrera, 2009: 658.

Tetrarhanis ilma ugandae. Male. Left – upperside; right – underside. Radio Hill, Mabira Forest, Uganda. 14 June, 2009. J. & C. Dobson.

Type locality: Uganda: “Masaka, Katera”. Distribution: Uganda, Kenya (west), Tanzania (north-west). Specific localities: Uganda – Masaka, Katera (TL); Semuliki N.P. (Davenport & Howard, 1996). Kenya – Kakamega Forest (Larsen, 1991c). Tanzania – Kikuru Forest (Congdon & Collins, 1998); Munene Forest (Congdon & Collins, 1998).

Note: Congdon & Collins (1998: 70) aver that subspecies ugandae and daltoni, because of their preference for ecologically different forests, may be distinct species.

*Tetrarhanis laminifer Clench, 1965

Tetrarhanis laminifer Clench, 1965. In Fox et al., 1965. Memoirs of the American Entomological Society No. 19: 300 (438 pp.).

Type locality: Cameroon: “Batanga”. Distribution: Cameroon, Equatorial Guinea (Mbini), Gabon. A male labelled ‘Liberia’ (Good) in the Carnegie Museum is almost certainly mislabelled (Larsen, 2005a). Specific localities: Cameroon – Batanga (TL). Habitat: Primary forest. Habits: This appears to be an extremely rare species (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

Note: d’Abrera, 2009: 658 appears to consider that laminifer Clench, 1965 is possibly a synonym of nubifera (Druce, 1910) but does not make any formal taxonomic changes.

5 *Tetrarhanis nubifera (Druce, 1910) White On-off

Liptena nubifera Druce, 1910. Proceedings of the Zoological Society of London 1910: 362 (356-378). Tetrarhanis nubifera Druce, 1910. d’Abrera, 2009: 658.

Type locality: Cameroon: “Bitje, Ja River, Cameroons, 2000 ft”. Distribution: ?Nigeria (south-east), Cameroon, Gabon, Congo. Specific localities: Cameroon – Bitje, Ja River (TL). Gabon – Lope N.P. (Vande weghe, 2010); Waka (Vande weghe, 2010); Bitam (Vande weghe, 2010); Langoue (Vande weghe, 2010). Habitat: Primary forest. Early stages: Nothing published. Larval food: Nothing published.

*Tetrarhanis ogojae (Stempffer, 1961) Ogoja On-off

Liptena (Tetrarhanis) ogojae Stempffer, 1961. Annales Musée Royal de l’Afrique Centrale (8) 94: 44 (73 pp.). Tetrarhanis ogojae Stempffer, 1961. d’Abrera, 2009: 659.

Type locality: Nigeria: “province d’Ogoja, Ikom”. Distribution: Nigeria (south and Cross River loop), Cameroon (west), Gabon. Specific localities: Nigeria – Ikom, Ogoja Province (TL); Gashaka-Gumpti (Larsen, 2005a). Cameroon – Korup (Larsen, 2005a). Gabon – Nouna (Vande weghe, 2010). Habitat: Primary forest. Habits: A very scarce species, only one specimen seen per fifty or so of Tetrarhanis simplex (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

*Tetrarhanis okwangwo Larsen, 1998 Okwangwo On-off

Tetrarhanis okwangwo Larsen, 1998. Lambillionea 98: 78 (77-80). Tetrarhanis okwangwo Larsen, 1998. d’Abrera, 2009: 660.

Tetrarhanis okwangwo. Female, Nigeria, underside. Image courtesy Torben Larsen.

Type locality: Nigeria: “Nigeria, Okwangwo Division of Cross River National Park, Butatong area, xi.

6 1995 (T.B. Larsen leg.).” Distribution: Nigeria (east and Cross River loop), Cameroon (west), Congo. Specific localities: Nigeria – Okwangwo (TL); Oban Hills (Larsen, 2005a). Cameroon – Korup (Larsen, 2005a); Ebogo (ABRI, vide Larsen, 2005a ). Congo – Ouesso (ABRI, vide Larsen, 2005a). Habitat: Primary forest. Habits: This appears to be a very rare butterfly (Larsen, 2005a). Flight period: Recorded in November and January. Early stages: Nothing published. Larval food: Nothing published.

*Tetrarhanis onitshae (Stempffer, 1962) Onitsha On-off

Liptena (Tetrarhanis) onitshae Stempffer, 1962. Bulletin de l’Institut Français d’Afrique Noire (A) 24: 1146 (1135-1181). Tetrarhanis onitshae Stempffer, 1962. d’Abrera, 2009: 659.

Type locality: Nigeria: “Onitsha, Awka, Mamu Forest”. Distribution: Nigeria (east). Known only from two males and a female collected by Jackson in Awka Forest, Onitsha in May, June and October 1959 (Larsen, 2005a). Specific localities: Nigeria – Mamu Forest, Awka, Onitsha (TL). Habitat: Primary forest. Habits: Nothing published. Early stages: Nothing published. Larval food: Nothing published.

*Tetrarhanis rougeoti (Stempffer, 1954)

Liptena (Tetrarhanis) rougeoti Stempffer, 1954. Bulletin de la Société Entomologique de France 59: 92 (88-93). Tetrarhanis rougeoti Stempffer, 1954. d’Abrera, 2009: 659.

Type locality: Gabon: “Douano”. Distribution: Gabon, Congo, Democratic Republic of Congo (Equateur). Specific localities: Gabon – Douano (TL). Habitat: Primary forest. Early stages: Nothing published. Larval food: Nothing published.

*Tetrarhanis schoutedeni (Berger, 1954)

Liptena (Tetrarhanis) schoutedeni Berger, 1954. Annales du Musée Royal du Congo Belge. C-Zoologie (4) 1: 308 (306-309). Tetrarhanis schoutedeni Berger, 1954. d’Abrera, 2009: 659.

Type locality: [Democratic Republic of Congo]: “Sankuru, Katako-Kombe”. Diagnosis: Can only be distinguished from Tetrarhanis stempfferi by examining the genitalia (Larsen, 2005a). Distribution: Cameroon, Democratic Republic of Congo (Sankuru). Specific localities: Cameroon – Makak Station (Birket-Smith, 1960). Democratic Republic of Congo – Katako-Kombe, Sankuru (TL). 7 Habitat: Primary forest. Early stages: Nothing published. Larval food: Nothing published.

Note: d’Abrera, 2009: 658 appears to consider that schoutedeni (Berger, 1954) is possibly a synonym of stempfferi (Berger, 1954) but does not make any formal taxonomic changes.

collitorum Birket-Smith, 1960 (as sp. of Lectiles). Bulletin de l’Institut Français d’Afrique Noire (A) 22: 968 (521-554, 924-983, 1259-1284). Cameroon: “A small clearing in dense forest near the river Nyong just south of the Station of Makak (Loc. 146), French Cameroon”.

*Tetrarhanis simplex (Aurivillius, 1895) Simple On-off

Perched Simple On-off. Cameroon, May 2019. Image courtesy Jeremy Dobson.

Liptena ilma var. simplex Aurivillius, 1895. Entomologisk Tidskrift 16: 201 (195-220, 255-268). Tetrarhanis simplex Aurivillius, 1895. d’Abrera, 2009: 658.

Tetrarhanis simplex. Male. Left – upperside; right – underside. Petit Savanna, Cameroon. 4 May 2019. J. Dobson. Images M.C. Williams ex Dobson collection.

Type locality: Cameroon: “Ikoti, Kitta, Ekundu”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/i). Distribution: Nigeria (east and Cross River loop), Cameroon, Congo, Central African Republic. Recorded, in error, from west of Nigeria (Larsen, 2005a). Specific localities: Cameroon – Ikoti (TL); Kitta (Aurivillius, 1895); Ekundu (Aurivillius, 1895). Central African Republic – Dzanga (Noss, 1998). Habitat: Primary forest, with some tolerance for forest degradation (Larsen, 2005a). Habits: A common species, but rarely numerous (Larsen, 2005a). Clusters of three to five are often found on the tendrils of vines or on dry twigs (Larsen, 2005a). Larsen (2005a) observed a female

8 ovipositing on mossy sticks lying on a path in Korup (Cameroon). Early stages: Nothing published. Larval food: Nothing published.

*Tetrarhanis stempfferi (Berger, 1954) Stempffer’s On-off

Liptena (Tetrarhanis) stempfferi Berger, 1954. Annales du Musée Royal du Congo Belge. C-Zoologie (4) 1: 307 (306-309). Tetrarhanis stempfferi Berger, 1954. d’Abrera, 2009: 659.

Type locality: [Democratic Republic of Congo]: “Lualaba, Kapanga”. Diagnosis: Can only be distinguished from Tetrarhanis schoutedeni by examining the genitalia (Larsen, 2005a). Distribution: Guinea, Liberia, Ghana, Nigeria, Cameroon, Gabon, Congo, Central African Republic, Democratic Republic of Congo, Uganda, Kenya. Habitat: Primary forest. Habits: A scarce butterfly in West Africa (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

Tetrarhanis stempfferi stempfferi (Berger, 1954)

Liptena (Tetrarhanis) stempfferi Berger, 1954. Annales du Musée Royal du Congo Belge. C-Zoologie (4) 1: 307 (306-309). Tetrarhanis stempfferi stempfferi Berger, 1954. d’Abrera, 2009: 659.

Type locality: [Democratic Republic of Congo]: “Lualaba, Kapanga”. Distribution: Guinea, Liberia, Ghana, Nigeria (south), Cameroon, Gabon, Congo, Central African Republic, Democratic Republic of Congo (Kivu, Uele, Tshopo, Sankuru, Lualaba). Specific localities: Guinea – Ziama (Safian et al., 2020). Liberia – Wologizi (Safian et al., 2020). Ghana – Atewa Range (Larsen, 2005a). Gabon – Tchimbele (Vande weghe, 2010); Mboumie (Vande weghe, 2010); Waka (Vande weghe, 2010); Lake Evaro (Vande weghe, 2010); Ipassa (Vande weghe, 2010). Democratic Republic of Congo – Kapanga, Lualaba (TL); Ituri Forest (Ducarme, 2018); Central Forest Block (Ducarme, 2018); Mt Mitumba (Ducarme, 2018).

Tetrarhanis stempfferi kigezi (Stempffer, 1956)

Liptena (Tetrarhanis) stempfferi kigezi Stempffer, 1956. Annales du Musée Royal du Congo Belge (8) (Sciences zoologique) 49: 8 (54 pp.). Tetrarhanis stempfferi kigezi Stempffer, 1956. d’Abrera, 2009: 659.

Type locality: Uganda: “Kayonza, Kigezi”. Distribution: Uganda (south-west – Kigezi district), Kenya (west). Specific localities: Uganda – Kigezi, Kayonza (TL).

*Tetrarhanis symplocus Clench, 1965 Clench’s On-off

Tetrarhanis simplex symplocus Clench, 1965. In Fox et al., 1965. Memoirs of the American Entomological Society No. 19: 9 302 (438 pp.). Tetrarhanis symplocus Clench, 1965. d’Abrera, 2009: 659.

Type locality: Liberia: “Harbel, Marshall Territory”. Distribution: Guinea, Liberia, Ivory Coast, Ghana, Togo, Benin (south), Nigeria (west). Specific localities: Guinea – Ziama (Safian et al., 2020). Liberia – Harbel (TL); Wologizi (Safian et al., 2020); Wonegizi (Safian et al., 2020). Ghana – Bobiri Butterfly Sanctuary (Larsen et al., 2007). Benin – see Coache et al., 2017. Nigeria – Okomu (R. Warren, vide Larsen, 2005a). Habitat: Primary forest. Habits: A common forest butterfly, widely distributed within a particular forest, possibly because they are not closely tied to ant-trees (Larsen, 2005a). They have been noted perching, with closed wings, on twigs just above the forest floor (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

10