AFROTROPICAL . MARK C. WILLIAMS. http://www.lepsocafrica.org/?p=publications&s=atb Updated 11 July 2021

Genus Ernsta Grishin, 2020 Sandmen

In Zhang et al. 2020. Zootaxa 4748 (1): 185 (182-194). Type-species: Pyrgus colotes Druce, 1875 [Proceedings of the Zoological Society of London 1875: 416 (406-417).], by original designation.

The genus Ernsta belongs to the Family Hesperiidae Latreille, 1809; Subfamily Pyrginae Burmeister, 1878; Tribe Carcharodini Verity, 1940; Subtribe Carcharodina Verity, 1940. Other genera of the Subtribe Carcharodina in the Afrotropical Region are , Agylla, Gomalia and Carcharodus.

Ernsta (Sandmen) is an Afrotropical genus of 13 small skippers. One species (zebra) extends extralimitally. Most of the species are found in the eastern Afrotropics and southern Arabian Peninsula but the two forest species are also found in West Africa. They are found in a variety of habitats, from forest to very dry savanna, Karoo and Fynbos but each species has a preferred type of habitat. They are fond of flowers and some species mud-puddle. The males are territorial. The larvae feed on the leaves of plants in the family Malvaceae.

Subgenus Ernsta Grishin, 2020

In Zhang et al. 2020. Zootaxa 4748 (1): 185 (182-194). Type-species: Pyrgus colotes Druce, 1875 [Proceedings of the Zoological Society of London 1875: 416 (406-417).], by original designation.

*Ernsta (Ernsta) colotes (Druce, 1875)# Bushveld Sandman

Males of the Bushveld Sandman (Ernsta colotes) Images courtesy Steve Woodhall

Pyrgus colotes Druce, 1875. Proceedings of the Zoological Society of London 1875: 416 (406-417). Spialia colotes Druce. Swanepoel, 1953a. Spialia colotis (Druce, 1875). Kielland, 1990d. [misspelling of species name]

1 Spialia colotes (Druce, 1875). Pringle et al., 1994: 318. Ernsta (Ernsta) colotes (Druce, 1875). Zhang et al., 2020.

Type locality: Angola. Diagnosis: Median band of hindwing underside composed of separate spots; forewings pointed (Pringle et al., 1994). Distribution: Ethiopia, Somalia, Uganda, Kenya, Tanzania, Zambia, Angola, Mozambique, Zimbabwe, Botswana, Namibia, , Swaziland, Saudi Arabia, Yemen, Oman. Habitat: Dry and moist savanna (bushveld). Also in forest clearings (Cock, 2016). In Tanzania at altitudes from 1 000 to 1 400 m (Kielland, 1990d). Habits: A relatively common species (Larsen, 1991c). Flies low down, often resting on low shrubs or the ground in the shade of trees. Both sexes are fond of flowers. Males do not hilltop, establishing territories anywhere but often in the vicinity of a thorn tree. In Kenya males of subspecies transvaaliae establish territories in forest clearings. They perch on low shrubs or on the ground. Flight period: December to May; commonest from January to April (Pringle et al., 1994). Early stages:

Cock, 2016: 319 [egg, leaf shelters, larva, pupa, parasitoids]

Final instar larva and pupa of Ernsta colotes. Images courtesy Allison Sharp.

Larval food: Hibiscus aethiopicus L. var. ovatus Harv. (Malvaceae) [Otto et al., 2013: 72; South Africa]. Hibiscus fuscus Garke (Malvaceae) [M.J.W. Cock vide Larsen, 1991c: 405; Kenya]. Pavonia burchellii (DC.) R.A. Dyer (Malvaceae) [Otto et al., 2013: 72; South Africa].

Ernsta (Ernsta) colotes colotes (Druce, 1875)

Pyrgus colotes Druce, 1875. Proceedings of the Zoological Society of London 1875: 416 (406-417). Spialia colotis (Druce, 1875). Kielland, 1990d. [misspelling of species name] Ernsta (Ernsta) colotes (Druce, 1875). Zhang et al., 2020.

Type locality: Angola. Distribution: Angola. Specific localities: Angola – Loanga (Plötz, 1884); Bengo Province; Cabinda Province; Luanda Province; Namibe Province (Mendes et al., 2013).

nora Plötz, 1884 (as sp. of Pyrgus). Stettiner Entomologische Zeitung 45: p? (145-150). Angola: “Loanga”.

Ernsta (Ernsta) colotes semiconfluens (de Jong, 1978)

Spialia colotes semiconfluens de Jong, 1978. Tijdschrift voor Entomologie 121: 58 (23-146). Ernsta (Ernsta) colotes semiconfluens (de Jong, 1978). Zhang et al., 2020.

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Type locality: Somalia: “Bihendula”. [near Shikh in northern Somalia (Cock, 2016). Distribution: Ethiopia, Somalia, Uganda (north-east), Kenya (north-east, east & south-east), Saudi Arabia, Yemen, Oman. Specific localities: Somalia – Bihendula (TL). Kenya – Tsavo area (Larsen, 1991c); Besil, on the roadto Namanga (Cock, 2016); Kibwezi (Cock, 2016); Voi (Cock, 2016); Chyulu Hills (Cock, 2016); Teita Hills (Cock, 2016); Shimba Hills (Cock, 2016).

Note: Cock (2016) notes that there appears to be an eco-climatic cline between the subspecies semiconfluens and transvaaliae as many intermediate specimens occur in eastern Kenya.

Ernsta (Ernsta) colotes transvaaliae (Trimen, 1889)# Bushveld Sandman

Pyrgus transvaaliae Trimen, 1889. South-African Butterflies: a monograph of the extra-tropical species 3 Papilionidae and Hesperidae: 286 (438 pp.). London. Spialia colotes transvaaliae (Trimen, 1889). Dickson & Kroon, 1978. Spialia colotes transvaaliae (Trimen and Bowker, 1889). Pringle et al., 1994: 318. Ernsta (Ernsta) colotes transvaaliae (Trimen, 1889). Zhang et al., 2020.

Ernsta colotes transvaaliae. Male (Wingspan 23 mm). Left – upperside; right – underside. Fochville, North West Province, South Africa. 1 February 2008. J. Dobson. Images M.C. Williams ex Dobson Collection.

Ernsta colotes transvaaliae. Female (Wingspan 27 mm). Left – upperside; right – underside. Fochville, North West Province, South Africa. 1 February 2008. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [South Africa]: “Transvaal, Potchefstroom District”. Holotype (male) in the Natural History Museum, London. Original description: “Exp. al., 11 lin. Allied to P. Vindex [= Spialia spio L.], Cram., and Dromus, Plötz. Male. Brownish- black, with very distinct, mostly rounded, white spots. Fore-wing: spots in disposition and relative size most like those of Dromus, but basi-cellular spots very narrow and longitudinally elongate, terminal cellular spot more quadrate, and costal spot of discal row narrower and more distinctly tripartite; spots of strongly-sinuated submarginal series very distinct (in one specimen larger than in the other) and of equal size throughout; no spot 3 beneath basi-cellular one. Hind-wing: a distinct basi-cellular spot; median band resembling that of Diomus, Hopff., but not curved, more oblique, more macular (the crossing nervules forming strongly-marked interruptions); submarginal series of spots more complete than in any of the allied forms, there being a series of three (rather smaller than the rest) somewhat obliquely placed between first subcostal and third median nervules. Under side: hind-wing and costal and apical border of fore-wing glossy yellowish-brown; all spots very distinct; hind-wing with median band not so macular as on upper side, but continuous from costa to submedian nervure, and with a sub-basal curved series of four conspicuous spots. Fore-wing: an elongate small whitish spot immediately below discoidal cell near base (followed in one specimen by a second similar spot); four upper spots of submarginal series much larger and more elongated longitudinally than on upper side. Hind-wing: a white spot at base; first and second spots of sub-basal row rather large and pyriform, third spot very small, fourth spot narrow and elongate, adjoining greyish-white of inner-marginal fold; seven spots in median band (which is irregular in outline on both edges), the two lowest spots almost touching the fourth spot in sub-basal series; spots of irregular submarginal row considerably larger and more elongated than on upper side.” Distribution: Uganda, Kenya, Tanzania, Zambia, Angola, Mozambique, Zimbabwe, Botswana, Namibia, South Africa (Limpopo Province, , North West Province, ), Swaziland. Specific localities: Tanzania – North, central, west, south-east (Kielland, 1990d); Dar es Salaam (Kielland, 1990d). Zambia – A single specimen in the Livingstone Museum bearing an illegible locality on the label (Heath et al., 2002). Angola – Huila Province (Mendes et al., 2013). Zimbabwe – Sabi Valley (Pringle et al., 1994); Mount Selinda (Pringle et al., 1994). Botswana – Mogoditshane (Larsen, 1991l); Tlokweng (Larsen, 1991l); Shashe (Larsen, 1991l); Maun (Larsen, 1991l). Limpopo Province – Warmbaths (Swanepoel, 1953); Potgietersrus (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Tubex (Swanepoel, 1953); Vivo (Swanepoel, 1953); Dendron (Swanepoel, 1953); Blouberg (Swanepoel, 1953); Leydsdorp (Swanepoel, 1953); Percy Fyfe Nature Reserve (Warren, 1990); Letsitele (male illustrated above); Highlands Wilderness (Bode & Bode, unpublished checklist); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015). North West Province – Potchefstroom (TL); De Kroon (Swanepoel, 1953). Gauteng – Tswaing (Williams).

* Ernsta (Ernsta) confusa (Higgins, 1925)# Confusing Sandman

Hesperia transvaaliae var. confusa Higgins, 1925. Transactions of the Entomological Society of London 1924: 90 (64-120). Spialia confusa (Higgins, 1925). Evans, 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum: 60 (212 pp.). Spialia confusa Higgins. Swanepoel, 1953a. Spialia confusa (Higgins, 1925). Dickson & Kroon, 1978. Spialia confusa (Higgins, 1925). Pringle et al., 1994: 319. Ernsta (Ernsta) confusa (Higgins, 1925). Zhang et al., 2020.

Ernsta confusa confusa. Male (Wingspan 21 mm). Left – upperside; right – underside. Tembe, KwaZulu-Natal, 25 February 2010. M. Williams. Images M.C. Williams ex Williams Collection.

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Ernsta confusa confusa. Female. Left – upperside; right – underside. Ndumu, KwaZulu-Natal, South Africa. 23 February 1997. J. Greyling. Images M.C. Williams ex Greyling Collection.

Type locality: [Malawi]: “Nyasaland”; [Zambia]: “N.E. Rhodesia”; Mozambique. Holotype (female) in the Natural History Museum, London. Distribution: Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, South Africa, Swaziland. Habitat: Moist savanna (dense bush). Populations appear to be localised to discrete areas. In Tanzania subspecies obscura is found in coastal bush and forest at altitudes from sea-level to 500 m (Kielland, 1990d). Habits: A rather scarce (Larsen, 1991c). The flight is fast and low down, but individuals settle frequently, on the ground, grass stems or shrubs. Flowers are often visited. Males establish territories in open places, frequently bare, sandy spots (Pringle et al., 1994). Flight period: All year (Pringle et al., 1994). Early stages: Nothing published. Larval food: species (Malvaceae) [Van Someren, 1974: 325; Sevastopulo, 1974 & 1975; for subspecies obscura; Kenya]. Cock (2016) is dubious about this record and states that it may refer to S. diomus. Triumfetta species (Malvaceae) [Sevastopulo, 1974 & 1975; for subspecies obscura; Kenya]. Cock (2016) is dubious about this record and states that it may refer to S. diomus.

Ernsta (Ernsta) confusa confusa (Higgins, 1925)# Confusing Sandman

Hesperia transvaaliae var. confusa Higgins, 1925. Transactions of the Entomological Society of London 1924: 90 (64-120). Spialia confusa (Higgins, 1925). Evans, 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum: 60 (212 pp.). Spialia confusa confusa (Higgins, 1925). Pringle et al., 1994: 319. [date of authorship erroneous] Ernsta (Ernsta) confusa confusa (Higgins, 1925). Zhang et al., 2020.

Ernsta confusa confusa. Male (Wingspan 21 mm). Left – upperside; right – underside. Tembe, KwaZulu-Natal, 25 February 2010. M. Williams. Images M.C. Williams ex Williams Collection.

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Ernsta confusa confusa. Female. Left – upperside; right – underside. Ndumu, KwaZulu-Natal, South Africa. 23 February 1997. J. Greyling. Images M.C. Williams ex Greyling Collection.

Type locality: [Malawi]: “Nyasaland”; [Zambia]: “N.E. Rhodesia”; Mozambique. Holotype (female) in the Natural History Museum, London. Distribution: Tanzania (central), Malawi, Zambia (north), Mozambique, Zimbabwe, South Africa (Limpopo Province, Mpumalanga, KwaZulu-Natal – north), Swaziland. Specific localities: Tanzania – Mikumi National Park at 500 m (Kielland, 1990d); south-east areas (Kielland, 1990d). Zambia – Petauke (Heath et al., 2002); upper Luangwa Valley (Heath et al., 2002). Mozambique – Xiluvo (Pennington); Inhaminga (Pinhey); Siluve. Zimbabwe – Sanyati Valley (Pinhey). Limpopo Province – Messina (single record). Mpumalanga – near Waterval-Onder (Owen-Johnston); Barberton. KwaZulu-Natal – between Hluhluwe and False Bay, about a kilometre and a half from the shore (Pennington); Makatini Flats (Pringle et al., 1994); north of Mhlosinga (Schoeman and the Pringles); Tembe Nature Reserve (Pringle & Kyle, 2002). Swaziland – Singceni (Pennington; single male).

fasciata Higgins, 1925 (as var. of Hesperia transvaaliae). Transactions of the Entomological Society of London 1924: 90 (64-120). [Malawi]: “Nyasaland, British East Africa”.

Ernsta (Ernsta) confusa obscura (Higgins, 1925)

Hesperia transvaaliae var. obscura Higgins, 1925. Transactions of the Entomological Society of London 1924: 92 (64-120). Spialia confusa obscura (Higgins, 1925). Evans, 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum: 60 (212 pp.). Ernsta (Ernsta) confuse obscura (Higgins, 1925). Zhang et al., 2020.

Type locality: [Kenya]: “British East Africa, Nairobi”. The type locality is suspect because the closest known population of obscura is in the Shimba Hills (Cock, 2016). Distribution: Kenya (east), Tanzania (north-east). Specific localities: Kenya – Nairobi (TL); Arabuko-Sokoke Forest (Larsen, 1991c); Rabai (Larsen, 1991c); Shimba Hills (Larsen, 1991c). Records from the Chyulu Hills by van Someren (1939) are for misidentified specimens of Ernsta colotes (De Jong, 1978). Tanzania – Dar es Salaam (Kielland, 1990d); Pugu Hills (Kielland, 1990d); Dendene Forest at Kisiju (Kielland, 1990d); foot of the Nguru Mountains (Kielland, 1990d).

* Ernsta (Ernsta) wrefordi (Evans, 1951) Wreford’s Grizzled

Spialia wrefordi Evans, 1951. Annals and Magazine of Natural History (12) 4: 1270 (1268-1272). Ernsta (Ernsta) wrefordi (Evans, 1951). Zhang et al., 2020.

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Ernsta wrefordi. Male. Left – upperside; right – underside. Kotido, Uganda. Images courtesy Torben Larsen.

Type locality: Uganda: “Kotido, Karamoja province”. Distribution: Uganda (north-east), Kenya (north). Specific localities: Uganda – Kotido (TL); Kongole (van Someren, vide Larsen, 1991c). Kenya – Samburu (Larsen, 1991c); Mount Kulal (Larsen, 1991c); Marsabit (Larsen, 1991c); Meru National Park (Larsen, 1991c). Habitat: Very dry savanna country (Larsen, 1991c). Habits: Nothing published. Early stages: Nothing published. Larval food: Nothing published.

* Ernsta (Ernsta) paula (Higgins, 1925)# Mite Sandman

Hesperia paula Higgins, 1925. Transactions of the Entomological Society of London 1924: 77 (64-120). Spialia paula Higgins. Swanepoel, 1953a. Spialia paula (Higgins, 1925). Dickson & Kroon, 1978. Spialia paula (Higgins, 1925). Pringle et al., 1994: 320. Ernsta (Ernsta) paula (Higgins, 1925). Zhang et al., 2020.

Ernsta paula. Male. Left – upperside; right – underside. Polokwane, Limpopo, South Africa. January 1980. A. Marais. ABRI-2019-2271. Images M.C. Williams ex ABRI Collection.

7 Ernsta paula. Female. Left – upperside; right – underside. Ysterberg (south of Polokwane), Limpopo Province, South Africa. 30 September 1995. J. Greyling. Images M.C. Williams ex Greyling Collection.

Type locality: [Zimbabwe]: “Buluwayo, S. Rhodesia”; South Africa: “Transvaal”. Holotype (male) in the Natural History Museum, London. Diagnosis: Characterized on the hindwing underside by the central shiny white patch (Pringle et al., 1994). Distribution: Zimbabwe, Botswana (east), South Africa (Limpopo Province, North West Province, Gauteng, Free State Province, Northern Cape Province). Specific localities: Zimbabwe – Bulawayo (TL). Botswana – Lobatse (Dickson & Kroon, 1978). Limpopo Province – Polokwane (Swanepoel, 1953); Munnik (Swanepoel, 1953). North West Province – Swartruggens (Swanepoel, 1953); Rustenburg (Swanepoel, 1953). Gauteng – Haakdoringboom north of Pretoria (Williams). Free State Province – Parys (the Terblanche’s); Viljoenskroon (the Terblanche’s). Northern Cape Province – Windsorton (G. Henning). Habitat: Dry savanna (bushveld). Habits: Flies rapidly about grassy spots, usually on the slopes of hills. Both sexes feed from flowers. Males may show hilltopping behaviour, mostly below the actual summit. Territories are usually established on bare, gravel-covered ground. This is a scarce butterfly in South Africa and is only slightly more common in Zimbabwe. Flight period: August to April but adults are most plentiful from August to October (Pringle et al., 1994). Early stages: Nothing published. Larval food: Nothing published.

* Ernsta (Ernsta) secessus (Trimen, 1891)# Wolkberg Sandman

Pyrgus secessus Trimen, 1891. Proceedings of the Zoological Society of London 1891: 102 (59-107). Spialia secessus Trimen. Swanepoel, 1953a. Spialia secessus (Trimen, 1891). Dickson & Kroon, 1978. Spialia secessus (Trimen, 1891). Pringle et al., 1994: 319. Ernsta (Ernsta) secessus (Trimen, 1891). Zhang et al., 2020.

Ernsta secessus. Male (wet season form) (Wingspan 29 mm). Left – upperside; right – underside. Barberton, Mpumalanga Province, South Africa. 25 November 2006. J. Dobson. Images M.C. Williams ex Dobson Collection.

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Ernsta secessus. Female (dry season form) (Wingspan 31 mm). Left – upperside; right – underside. Munnik, Limpopo Province, South Africa. 12 September 2004. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Angola: “Omrora”. Holotype (male) in the South African Museum, Cape Town. Distribution: Angola, Democratic Republic of Congo (Shaba), Tanzania (west), Malawi, Zambia, Zimbabwe (north), ?Botswana (north-east), Namibia (north), South Africa (Limpopo Province, Mpumalanga, KwaZulu-Natal – north), Swaziland. Specific localities: Angola – Omrora (TL); Luanda Province (Mendes et al., 2013). Tanzania – Ufipa (Kielland, 1990d); Mpanda (Kielland, 1990d); Kigoma (Kielland, 1990d); Ngara District (Kielland, 1990d). Zambia – Ikelenge (Heath et al., 2002); Kafue Flats (Heath et al., 2002); Ndola (Heath et al., 2002); Mufulira (Heath et al., 2002); Mpongwe (Heath et al., 2002); Chalimbana (Heath et al., 2002); Luwingu (Heath et al., 2002); Mporokoso (Heath et al., 2002); Lufubu River (Heath et al., 2002). Zimbabwe – Victoria Falls; Nyanga. Limpopo Province – Haenertsburg (Swanepoel, 1953); Munnik – Buffelsberg (Swanepoel, 1953); Helpmekaar (Swanepoel, 1953); Groenbult (Swanepoel, 1953); Wolkberg (Pringle et al., 1994). Mpumalanga – Barberton (Swanepoel, 1953); Graskop (Swanepoel, 1953). KwaZulu-Natal – Lebombo Mountains (Pringle et al., 1994). Swaziland – Malolotja N. R. (www.sntc.org.sz). Habitat: Grassland, mainly at high altitudes. In Tanzania also in woodland, occurring at altitudes from 1 000 to 2 200 m (Kielland, 1990d). Habits: Flies low down, and not very fast, in short grass, with a zig-zagging flight pattern. Rests frequently on low vegetation or on the ground. Both sexes are readily attracted to flowers. Males hilltop during the midday hours on hot, sunny days and usually perch on the ground. Flight period: July to March in South Africa; throughout the year in the northern parts of its range. There are distinct seasonal forms (Pringle et al., 1994). Early stages: Nothing published. Larval food: Nothing published.

trimeni Aurivillius, 1925 in Seitz, 1908-25 (as f. of Hesperia secessa). Die Gross-Schmetterlinge der Erde, Stuttgart (2) 13 Die Afrikanischen Tagfalter: 564 (614 pp.). “Rhodesia”. Holotype in the Swedish Natural History Museum (no images available at www2.nrm.se/en/lep_nrm/s). [This is the wet-season form of Ernsta secessus.]

* Ernsta (Ernsta) dromus (Plötz, 1884)# Forest Sandman

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Upper- and underside of the Forest Sandman ( Ernsta dromus) Images courtesy Steve Woodhall

Pyrgus dromus Plötz, 1884. Mitteilungen aus dem Naturwissenschaftlichen Vereins von Neu-Vorpommen und Rügen in (Greifswald) Berlin 1884: 6 (1-24). Pyrgus dromus Plötz, 1884. Trimen & Bowker, 1889. Spialia dromus Plötz. Swanepoel, 1953a. Spialia dromus (Plötz, 1884). Dickson & Kroon, 1978. Spialia dromus (Plötz, 1884). Pringle et al., 1994: 319. Ernsta (Ernsta) dromus (Plötz, 1884). Zhang et al., 2020.

Ernsta dromus. Male (Wingspan 26 mm). Left – upperside; right – underside. Lekgalameetse Nature Reserve, Limpopo Province, South Africa. 6 January, 2012. M. Williams. Images M.C. Williams ex Williams Collection.

Ernsta dromus. Female (Wingspan 31 mm). Left – upperside; right – underside. Umtamvuna River, KwaZulu-Natal, South Africa. 17 December 2008. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [Democratic Republic of Congo]: “Congo”. The type(s) has apparently been lost. Diagnosis: The median band on the underside of the hindwing is characteristically curved (Pringle et al., 1994). Distribution: Sub-Saharan Africa, including Senegal, Gambia, Sierra Leone (north), Burkina Faso, Guinea-Bissau (Mendes et al., 2007), Guinea (north), Ivory Coast (north), Ghana (north), Benin (south, central), Nigeria, Gabon (de Jong, 1978); Democratic Republic of Congo, Uganda, Kenya, Tanzania, Zambia, Angola, Mozambique, Zimbabwe, ?Botswana, South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, KwaZulu-Natal, Eastern Cape Province), Swaziland. 10 Specific localities: Gambia – Tujering, Keneba, Abuko, Brusubi, Tujering, Sifoe (Jon Baker, pers. comm, May 2020). Ghana – Accra Plains (Larsen, 2005a); Badagry Creek beaches near Lagos (Larsen, 2005a); Boabeng- Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – Houeyogbe Forest (Coache & Rainon, 2016); see Coache et al., 2017. Democratic Republic of Congo – Mpala (Oberthür, 1912); Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Mt Mitumba (Ducarme, 2018); Mt Blue (Ducarme, 2018). Tanzania – Throughout (Kielland, 1990d); Kitanga (Oberthür, 1912); Zanzibar (Higgins, 1925); Katavi National Park (Fitzherbert et al., 2006). Zambia – Ikelenge (Heath et al., 2002); Kabompo Gorge (Heath et al., 2002); Mufulira (Heath et al., 2002); Kitwe (Heath et al., 2002); Ndola (Heath et al., 2002); Chalimbana (Heath et al., 2002); Luangwa Valley (Heath et al., 2002); Luongo River (Heath et al., 2002); Kasama (Heath et al., 2002); Mbala (Heath et al., 2002). Angola – Cabinda Province; Huambo Province; Huila Province; Kuando-Kubango Province; Kwanza Norte Province; Kwanza Sul (Mendes et al., 2013). Mozambique – Mount Namuli (Congdon et al., 2010); Mount Mabu (Congdon et al., 2010). Zimbabwe – Burma Road, Vumba (Cookson; male specimen illustrated above). Limpopo Province – Warmbaths (Swanepoel, 1953); Potgietersrus (Swanepoel, 1953); Mica (Swanepoel, 1953); Lekgalameetse Nature Reserve (“Malta Forest”) (Swanepoel, 1953); Tubex (Swanepoel, 1953); Woodbush (Swanepoel, 1953); Munnik (Swanepoel, 1953); Sibasa (Swanepoel, 1953); Louis Trichardt (Swanepoel, 1953); Saltpan (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Chuniespoort (Swanepoel, 1953); Doorndraai Dam Nature Reserve (Warren, 1990); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015); Bateleur Nature Reserve (Williams & Dobson, unpub., 2015). Mpumalanga – Barberton (Swanepoel, 1953); Komatipoort (Swanepoel, 1953); Nelspruit (Swanepoel, 1953); Lydenburg district (Swanepoel, 1953); Graskop (Swanepoel, 1953). Gauteng – Pretoria (Swanepoel, 1953). KwaZulu-Natal – Umkomaas (Swanepoel, 1953); Isipingo (Swanepoel, 1953); Durban (Swanepoel, 1953); Pietermaritzburg (Swanepoel, 1953); Karkloof (Swanepoel, 1953); Greytown (Swanepoel, 1953); Ismont (Swanepoel, 1953); Amahlongwa (Swanepoel, 1953); Eshowe (Swanepoel, 1953); Hluhluwe (Swanepoel, 1953); Kosi Bay Nature Reserve (Pringle & Kyle, 2002); Tembe Nature Reserve (Pringle & Kyle, 2002). Swaziland – Mlawula N. R. (www.sntc.org.sz); Malolotja N. R. (www.sntc.org.sz). Habitat: Mainly forest (clearings and margins) and, to a lesser extent, moist savanna (bushveld). Occasionally in grassland (Gauteng) and dry thorn savanna (Limpopo Province, Mpumalanga and KwaZulu-Natal) (Pringle et al., 1994). In Tanzania at altitudes from near sea-level to 2 000 m (Kielland, 1990d). Habits: A relatively common skipper (Larsen, 1991c). Adults fly close to the ground with a skipping pattern and regularly rest on low vegetation. Flowers and damp places are often visited. Males establish territories, which they defend by chasing off intruding butterflies. Males also occasionally show hilltopping behaviour (Pringle et al., 1994). Flight period: All year (Pringle et al., 1994). Early stages:

Clark, in Dickson & Kroon, 1978: 232; plate 14 [as Spialia dromus; recorded from eggs and larvae from Durban, KwaZulu-Natal]. “Egg: 0,8 mm diameter by 0,7 mm high; laid singly among young shoots. Whitish-green, changing to dirty white, with 20 longitudinal ribs which break up towards the crown and only some seven very disjointed pieces of which reach the micropyle. Cross-bracing is often very indistinct, consisting of about 20 ribs. Eggs hatch after 7 days. The discarded shell is not eaten. Larva: 1st instar 1,5 to 3 mm in 8 days; 2nd instar 3 to 5 mm in 8 days; 3rd instar 5 to 8 mm in 8 days; 4th instar 8 to 12 mm in 8 days; 5th instar 12 to 20 mm in 12 days. The duration of the instars varies, as in the case of other species of the group. The young larva hides in the fold of a young half-opened leaf and feeds on the inner surface. Later it binds leaves together to form a purse-like shelter in which it lives. It feeds on the surface of the leaf or crawls out to feed on adjacent leaves, when it generally eats the edges. Moulting and pupation take place within the ‘purse’. By the time the larva is ready for pupation this shelter is generally situated near the ground. There is a succession of broods. Pupa: 12,5 mm. Secured within its shelter by the cremastral hooks. Emergence takes place after 18 days. Parasites: Larvae are attacked by Diptera. The maggot emerges from the side of the larva or pupa and wriggles to a suitable spot, either within the shelter or among rubbish on the ground, and pupates after some 24 to 36 hours. The parasitic fly emerges from

11 the pupa after 12 to 25 days.”

Cock, 2016: 324 [egg, leaf shelters, larva, pupa]

Congdon et al., 2017 [final instar larva].

Early stages of Ernsta dromus. Left – early instar larva. Right – late instar larve. Images courtesy Steve Woodhall.

Early stages of Ernsta dromus. Pupa. Images courtesy Steve Woodhall.

Larval food: Hermannia spp. (Malvaceae) [Pinhey, 1965; Southern Africa; requires confirmation (Cock, 2016)]. Hibiscus species (Malvaceae) [Pinhey, 1949; Southern Africa]. Melhania species (Malvaceae) [Van Someren, 1974: 325; requires confirmation (Cock, 2016)]. Jacq. (Malvaceae) [Platt, 1921 (South Africa); Vuattoux, 1999 (Ivory Coast); Cock, 2016 (Kenya & Zimbabwe)]. Triumfetta tomentosa Bojer (Malvaceae) [Platt, 1921 (South Africa): 189; Cock, 2016 (Kenya)]. Triumfetta setulosa Mast. (Malvaceae) [Congdon et al., 2017; Mufindi, Tanzania]. Triumfetta sp. (Malvaceae) [Congdon, unpub. 2013 vide Cock, 2016; Tanznia]. indica L. (Malvaceae) [Van Someren, 1974: 325; as Waltheria americana L.; requires confirmation (Cock, 2016)].

zaira Plötz, 1884 (as sp. of Pyrgus). Mitteilungen aus dem Naturwissenschaftlichen Vereins von Neu- Vorpommen und Rügen in (Greifswald) Berlin 1884: 6 (1-24). “Congo”.

melaleuca Oberthür, 1912 (as sp. of Syrichthus). Étude de Lépidoptérologie Comparée 6: 113 (47-120). [Tanzania]: “Kitanga, dans l’Afrique orientale allemande”.

leucomelas Oberthür, 1912 (as sp. of Syrichthus). Étude de Lépidoptérologie Comparée 6: 118 (47- 120). [Democratic Republic of Congo]: “M’pala; Lac Tanganika”.

elongata Higgins, 1925 (as var. of Hesperia dromus). Transactions of the Entomological Society of London 1924: 95 (64-120). Makindu, Kenya.

meridionalis Higgins, 1925 (as var. of Hesperia dromus). Transactions of the Entomological Society of London 1924: 95 (64-120). [Tanzania]: “Zanzibar”.

12

* Ernsta (Ernsta) ploetzi (Aurivillius, 1891) Forest Grizzled Skipper

Hesperia ploetzi Aurivillius, 1891. Entomologisk Tidskrift 12: 227 (193-228). Spialia ploetzi ploetzi (Aurivillius, 1891). De Jong, 1977. Ernsta (Ernsta) ploetzi (Aurivillius, 1891). Zhang et al., 2020.

Ernsta ploetzi ploetzi. Male (Wingspan 28 mm). Left – upperside; right – underside. Kaweri Papyrus, Uganda. 10 January 2010. M. Barnett. Images M.C. Williams ex Dobson Collection.

Ernsta ploetzi ploetzi. Female. Left – upperside; right – underside. Kadera Forest, Uganda. May 2012. JP Leqeux. ABRI-2019-2279. Images M.C. Williams ex ABRI Collection.

Type locality: Cameroon: “Camerun”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/p). Diagnosis: Characterized on the hindwing underside by the discal band, which begins at mid costa and runs towards the distal margin at vein 2. In other species of Ernsta the discal band runs towards the hind margin or anal angle (Congdon & Collins, 1998). Distribution: Guinea-Bissau (Larsen, 2005a), Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Gabon, Congo, Angola, Democratic Republic of Congo, Uganda, Rwanda, Burundi, Kenya, Tanzania, Zambia. Habitat: Forest, especially disturbed areas, rather than primary forest (Larsen, 2005a). Forest edges and along paths near streams (Gardiner, 2010b). Habits: A fairly common species (Larsen, 2005a). Flies low down, settling on vegetation along forest paths (Congdon & Collins, 1998) or feeding from flowers (Larsen, 1991c). Larsen (2005a) states that it tends to fly higher up than the savanna species of the genus. Males regularly mud-puddle (Larsen, 2005a). Early stages:

Fontaine, 1988. [as Spialia rebeli; nominate subspecies]

Larsen, 1981. [subspecies occidentalis]

Cock, 2016: 329 [both subspecies; leaf shelters; larva; pupa]

13

Larval food: Triumfetta rhomboidea (Malvaceae) [Vuattoux, 1999 (Ivory Coast)]. Triumfetta species (Malvaceae) [Larsen, 1981 (Lagos, Nigeria); Cock, 2016 (Cameroon & Ivory Coast)].

Ernsta (Ernsta) ploetzi ploetzi (Aurivillius, 1891)

Hesperia ploetzi Aurivillius, 1891. Entomologisk Tidskrift 12: 227 (193-228). Spialia ploetzi ploetzi (Aurivillius, 1891). De Jong, 1977. Ernsta (Ernsta) ploetzi ploetzi (Aurivillius, 1891). Zhang et al., 2020.

Ernsta ploetzi ploetzi. Male (Wingspan 28 mm). Left – upperside; right – underside. Kaweri Papyrus, Uganda. 10 January 2010. M. Barnett. Images M.C. Williams ex Dobson Collection.

Ernsta ploetzi ploetzi. Female. Left – upperside; right – underside. Kadera Forest, Uganda. May 2012. JP Leqeux. ABRI-2019-2279. Images M.C. Williams ex ABRI Collection.

Type locality: Cameroon: “Camerun”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/p). Distribution: Cameroon (except west), Gabon, Congo, Angola (north), Democratic Republic of Congo, Uganda, Rwanda, Burundi, Kenya (west), Tanzania (north-west), Zambia (north-west). Specific localities: Cameroon – Douala (Cock, 2016). Gabon – Tchimbele (Vande weghe, 2010); Ovan (Vande weghe, 2010); Nouna (Vande weghe, 2010). Democratic Republic of Congo – Isiro (Fontaine, 1988); Ituri Forest (Ducarme, 2018); Semuliki Valley (Ducarme, 2018); Central Forest Block (Ducarme, 2018); Mt Mitumba (Ducarme, 2018); Mt Blue (Ducarme, 2018). Kenya – Kakamega (Larsen, 1991c). A record from Mombasa (de Jong, 1978) is almost certainly erroneous (Larsen, 1991c). Tanzania – Kikuru Forest (relatively common) (Congdon & Collins, 1998); Minziro Forest (Congdon & Collins, 1998). Zambia – Ikelenge district (Gardiner, 2010b). Angola – Cabinda Province (Mendes et al., 2013). 14

zebra Rebel, 1914 (as sp. of Hesperia). Annalen des (K.K.) Naturhistorischen Museums. Wien 28: 271 (219-294). [Tanzania]: “Randegebirgen im Nordwesten des Tanganyika-Sees”. [Invalid; junior secondary homonym of Spialia zebra (Butler), 1888 [Hesperiidae].]

rebeli Higgins, 1925 (as [unnecessary] replacement name for Hesperia zebra Rebel). Transactions of the Entomological Society of London 1924: 101 (64-120).

Ernsta (Ernsta) ploetzi occidentalis (de Jong, 1977)

Spialia ploetzi occidentalis de Jong, 1977. Entomologische Berichten 37: 43 (41-44). Ernsta (Ernsta) ploetzi occidentalis (de Jong, 1977). Zhang et al., 2020.

Ernsta ploetzi occidentalis. Male. Left – upperside; right – underside. Mamou, Guinea. September 2001. M. Mane. ABRI-2019-2277. Images M.C. Williams ex ABRI Collection.

Ernsta ploetzi occidentalis. Female. Left – upperside; right – underside. Loloby, Ghana. September 2001. SCC. ABRI-2019-2278. Images M.C. Williams ex ABRI Collection.

Type locality: Ghana. Distribution: Guinea-Bissau (Larsen, 2005a), Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (south, central), Nigeria, Cameroon (west). Specific localities: Guinea – Ziama (Safian et al., 2020). Sierra Leone – Loma Mountains (Larsen, 2005a). Ivory Coast – Yapo Forest (Cock, 2016). Ghana – Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – Houeyogbe Forest (Coache & Rainon, 2016); see Coache et al., 2017. Nigeria – Lagos (Larsen, 1981). Cameroon – Korup (Larsen, 2005a).

15 Subgenus Delaga Grishin, 2020

In Zhang et al. 2020. Zootaxa 4748 (1): 186 (182-194). Type-species: Pyrgus delagoae Trimen, 1898 [Transactions of the Entomological Society of London 1898: 15 (1-16).], by original designation.

* Ernsta (Delaga) mangana (Rebel, 1899) Arabian Grizzled Skipper

Hesperia (Pyrgus) mangana Rebel, 1899. Anzeiger der Akademie der Wissenschaften. Wien. 36: 360 (359-361). Ernsta (Delaga) mangana (Rebel, 1899). Zhang et al., 2020.

Ernsta mangana. Male. Left – upperside; right – underside. Taiz-Turbah Road, Yemen. October 1992. SCC. ABRI-2019-2280. Images M.C. Williams ex ABRI Collection.

Ernsta mangana. Female. Left – upperside; right – underside. Ukambani, Kenya. November 1997. SCC. ABRI-2019-2281. Images M.C. Williams ex ABRI Collection.

Type locality: [Yemen]: “Râs Fártak (Arab. m.)”. Distribution: Yemen, Oman, United Arab Emirates (Feulner & Roobas, 2014), Somalia, Ethiopia, Uganda, Kenya (north). Specific localities: Yemen – Ras Fartak (Larsen, 1991c). Oman – north (Feulner, 2007). United Arab Emirates – Hajar Mountains (Feulner & Roobas, 2014). Uganda – Karamoja area (Larsen, 1991c). Kenya – Suk (Larsen, 1991c); West Pokot (Larsen, 1991c); Kacheliba (Larsen, 1991c); Kibwezi (Larsen, 1991c). Habitat: Rocky ground in very dry savanna (Larsen, 1991c). Habits: This is an uncommon butterfly (Larsen, 1991c). When flying over stony ground their small size makes them difficult to see, except when they are settled on the flowers of Tridax species (Larsen, 1991c).

16 Early stages: Nothing published. Larval food: Melhania muricata Balf. f. (Malvaceae) (suspected) [Feulner & Roobas, 2014; United Arab Emirates].

Relevant literature: Feulner & Roobas, 2014 [Recorded from UAE]. Feulner, 2007 [Recorded from northern Oman].

* Ernsta (Delaga) nanus (Trimen, 1889)# Dwarf Sandman

Dwarf Sandman (Ernsta nanus) male undeside. Image courtesy Steve Woodhall.

Pyrgus nanus Trimen, 1889. South-African Butterflies: a monograph of the extra-tropical species 3 Papilionidae and Hesperidae: 290 (438 pp.) London. Spialia nanus Trimen. Swanepoel, 1953a. Spialia nanus (Trimen, 1889). Dickson & Kroon, 1978. Spialia nanus (Trimen and Bowker, 1889). Pringle et al., 1994: 317. Ernsta (Delaga) nanus (Trimen, 1889). Zhang et al., 2020.

Ernsta nanus. Male (Wingspan 22 mm). Left – upperside; right – underside. Brandvlei Dam, Western Cape Province, South Africa. 12 December 2005. J. Dobson. Images M.C. Williams ex Dobson Collection.

Ernsta nanus. Female (Wingspan 25 mm). Left – upperside; right – underside. Springfontein, Free State Province, South Africa. 5 October 2003. J. Dobson. Images M.C. Williams ex Dobson Collection.

17

Type locality: [South Africa]: “in the vicinity of [Cape Town], Vogel Vley, Namaqualand District, Robertson”. Holotype in the Natural History Museum, London. Original description: “Exp. al., male 9.5-11.5 lin.; female 11 lin.-1 in. Nearly allied to P. Sataspes, Trim. Male. Brownish- black, with small white spots; spots in number and position almost as in Sataspes, but of a purer white and more sharply defined; hind-wing median band divided as in Sataspes, but not at all oblique. Under side: hind-wing: ground-colour much paler; base with a whitish tinge; both transverse bands whiter, and beginning on costal edge, - the sub-basal one not extending below subcostal nervure, - the median one outwardly dark-edged, biangulated (just above third and just upon first median nervules), and not oblique, its attenuated lower extremity joining white of inner-marginal fold at some distance before anal angle; an indistinct bisinuated submarginal series of white dots. Female. White spots and white of cilia purer, more conspicuous. Under side: all the markings more sharply defined.” Diagnosis: The identifying feature is the unbroken irregular central band on the underside of the hindwing (Pringle et al., 1994). Distribution: Namibia (south), South Africa (North West Province – south, Free State Province, Eastern Cape Province – west, Western Cape Province, Northern Cape Province). Specific localities: Namibia – Aus (Ficq). Free State Province – Bloemfontein (Swanepoel, 1953); Cyferfontein [-30.3736 25.8131] (R. Griesel, unpublished). Eastern Cape Province – Uitenhage (Swanepoel, 1953); Cradock (Swanepoel, 1953); Cookhouse (Swanepoel, 1953); Graaff-Reinet (Swanepoel, 1953); Carlton (Swanepoel, 1953); Somerset East (Swanepoel, 1953); Steynsburg (Swanepoel, 1953); Addo Road (Dickson & Kroon, 1978). Western Cape Province – Cape Town (TL); Robertson (Trimen, 1889); Malmesbury (Swanepoel, 1953); Clanwilliam (Swanepoel, 1953); Klaver (Swanepoel, 1953); Vanrynsdorp (Swanepoel, 1953); Bitterfontein (Swanepoel, 1953); Paarl (Swanepoel, 1953); Worcester (Swanepoel, 1953); Matjesfontein (Swanepoel, 1953); Calitzdorp (Swanepoel, 1953); Beaufort West (Swanepoel, 1953). Northern Cape Province – Kamieskroon (Swanepoel, 1953); Springbok (Swanepoel, 1953); Colesberg (Swanepoel, 1953); De Aar (Swanepoel, 1953); Kimberley (Swanepoel, 1953); Carnarvon (Swanepoel, 1953); Prieska (Swanepoel, 1953); Kuruman (Cottrell). Habitat: Primarily karoo vegetation but extending into the fynbos of the Western Cape Province. Habits: The flight is rapid but close to the ground. Both sexes are fond of flowers. Males establish territories on the flats or slopes of mountains and perch on the ground or on stones (Pringle et al., 1994). Flight period: August to April, with peak emergences in September-October and March-April (Pringle et al., 1994). Early stages:

Clark, in Dickson & Kroon, 1978: 226; plate 11 [as Spialia nanus; recorded from eggs from Addo Road, Eastern Cape]. “Egg: 0,7 mm diameter by 0,5 mm high; laid singly among young buds and cream coloured at first, changing to pale green. There are 17 to 21 heavy, irregular longitudinal ribs, only some 5 of which reach the micropyle, and 16 to 18 very thin cross-ribs. Eggs hatch after some 12 days. The discarded shell is not eaten. Larva: 5-instar Group 1st instar 1,25 to 3 mm in 8 days; 2 nd instar 3 to 4,5 mm in 6 days; 3rd instar 4,5 to 6,5 mm in 10 days; 4th instar 6,5 to 10 mm in 14 days; 5 th instar 10 to 17 mm in 27 days. Larva: 6-instar Group 1st instar 1,25 to 2,5 mm in 8 days; 2nd instar 2,5 to 4 mm in 8 days; 3rd instar 4 to 5,25 mm in 8 days; 4th instar 5,25 to 8,5 mm in 8 days; 5th instar 8,5 to 12,5 mm in 8 days; 6th instar 12,5 to 17 mm in 17 days. The duration of instars in both groups is very variable, depending on climatic conditions. Young larvae start their existence concealed in a partly open, very young leaf, and feed on the inner surface; later, they construct a ‘purse’ by binding leaves together. In this they live, moult and pupate. The final shelter is usually near the ground. There are generally three or four broods, the instars of each being most erratic. Pupa: 11 mm. Secured in its shelter by cremastral hooks; usually covered with white powder. Emergence takes place after 12 days. Parasites: Parasitical attack is similar to that in most Spialia, and the parasites are probably the same. The species is multi-brooded but, as with a number of other members of the group, development of the early stages is retarded in the winter months.”

Henning, Henning, Joannou & Woodhall, 1997: 93 [photograph of final instar larva].

18

Early stages of Ernsta nanus. Egg. Image courtesy Steve Woodhall.

Larval food: Hermannia comosa Burch. ex DC. (Malvaceae) [Murray, 1959]. Hermannia cuneifolia Jacq. var. cuneifolia (Malvaceae) [Murray, 1959; as Hermannia pollens Eckl. and Zehy.]. Hermannia diffusa L.f. (Malvaceae) [Murray, 1959; as Hermannia pilosula Harv.]. Hermannia incana Cav. (Malvaceae) [Murray, 1959; as Hermannia candicans Ait.]. Hermannia pulverata Andrews (Malvaceae) [Dickson & Kroon, 1978: 187]. Hibiscus aethiopicus L. (Malvaceae) [Dickson & Kroon, 1978: 187]. Hermannia sp. (Malvaceae) [Dickson & Kroon, 1978: 187; as Mahernia sp.]. Pavonia burchellii (DC.) R.A.Dyer (Malvaceae) [Murray, 1959; as Pavonia macrophylla E. Mey. ex Harv.].

* Ernsta (Delaga) delagoae (Trimen, 1898)# Delagoa Sandman

Males of the Delagoa Sandman (Ernsta delagoae) Images courtesy Steve Woodhall

Pyrgus delagoae Trimen, 1898. Transactions of the Entomological Society of London 1898: 15 (1-16). Spialia delagoae Trimen. Swanepoel, 1953a. Spialia delagoae (Trimen, 1898). Dickson & Kroon, 1978. Spialia delagoae (Trimen, 1898). Pringle et al., 1994: 317. Ernsta (Delaga) delagoae (Trimen, 1898). Zhang et al., 2020.

19

Ernsta delagoae. Male (Wingspan 22 mm). Left – upperside; right – underside. Thabazimbi, Limpopo Province, South Africa. 23 January 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Ernsta delagoae. Female (Wingspan 26 mm). Left – upperside; right – underside. Stoffberg, Mpumalanga Province, South Africa. 5 February 2006. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: Mozambique: “Delagoa Bay”. Holotype in the South African Museum, Cape Town. Diagnosis: Easily recognized by the unbroken, straight, light band on the underside of the hindwing (Pringle et al., 1994). Distribution: Uganda (north-east), Kenya (north-west), Zambia, Angola, Mozambique, Zimbabwe, Botswana, Namibia (north), South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, KwaZulu-Natal – north, Northern Cape Province), Swaziland. Specific localities: Kenya – Kacheliba (Larsen, 1991c); Kedong (Larsen, 1991c). Zambia – Solwezi (De Jong, 1978). Angola – Luanda Province (Mendes et al., 2013). Mozambique – Delagoa Bay (TL). Botswana – Lobatse (Pennington); Kang (Larsen, 1991l); Middelspits (Larsen, 1991l); Serowe (Larsen, 1991l); Gaborone (Larsen, 1991l); Francistown (Larsen, 1991l); Mpandama-Tenga (Larsen, 1991l); Kasane (Larsen, 1991l); Chobe (Larsen, 1991l). Limpopo Province – Zoutpansberg (Swanepoel, 1953); Vivo (Swanepoel, 1953); Dendron (Swanepoel, 1953); Bandolierkop (Swanepoel, 1953); Rita (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Munnik (Swanepoel, 1953); Chuniespoort (Swanepoel, 1953); Tubex (Swanepoel, 1953); Warmbaths (Swanepoel, 1953); Percy Fyfe Nature Reserve (Warren, 1990); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015). Mpumalanga – Ohrigstad (Swanepoel, 1953); Barberton (Swanepoel, 1953). North West Province – Borakalalo Nature Reserve (J. Dobson, unpublished, 2009). KwaZulu-Natal – Estcourt (Swanepoel, 1953); Weenen (Swanepoel, 1953); Muden (Swanepoel, 1953); Middeldrift (Swanepoel, 1953); Kearsney (Swanepoel, 1953); Hluhluwe (Swanepoel, 1953); Tembe Nature Reserve (Pringle & Kyle, 2002). Swaziland – Mlawula N. R. (www.sntc.org.sz). Habitat: Moist and dry savanna (bushveld). Habits: Appears to be scarce throughout its range (Larsen, 1991c). The flight is quite fast, buzzing and close to the ground. Flowers and damp spots on the edges of streams are visited. Swanepoel (1953) records them drinking from fresh baboon faeces. Males establish territories in areas of short grass on the flats or just below the peaks of hills and ridges. They select perches on the ground or on stones (Pringle et al., 1994). Flight period: All year in the north of its distribution but in South Africa commonest in August- September and February-March (Pringle et al., 1994). Early stages: Nothing published. Larval food: Melhania prostrata DC. (Malvaceae) [Otto et al., 2013: 72].

20 * Ernsta (Delaga) zebra (Butler, 1888) Zebra Grizzled Skipper

Pyrgus zebra Butler, 1888. Annals and Magazine of Natural History (6) 1: 207 (132-151, 196-209). Ernsta (Delaga) zebra (Butler, 1888). Zhang et al., 2020.

Ernsta zebra bifida. Male. Left – upperside; right – underside. Ngong, Kenya. June 1979. SCC. ABRI-2019-2282. Images M.C. Williams ex ABRI Collection.

Ernsta zebra bifida. Female. Left – upperside; right – underside. Nandi, Kenya. October 1971. SCC. ABRI-2019-2283. Images M.C. Williams ex ABRI Collection.

Type locality: [India] Pakistan: “Campbellpore [Attock]; Futch Khan’s bungalow, Kooteer, Chittar Pahar Range [Chittar Pahar], 2000 to 3000 feet”. Distribution: Yemen, Oman, Ethiopia, Somalia, Sudan, Democratic Republic of Congo, Uganda, Kenya, Tanzania (subspecies bifida). The nominate subspecies is from Pakistan (Punjab). Habitat: Open country (Larsen, 1991c). Savanna, including dry bush country, at altitudes from sea- level to 1 800 m in Tanzania (Kielland, 1990d). Habits: Subspecies bifida is commoner in Kenya than elsewhere in its range (Larsen, 1991c). Flies very close to the ground, thus being very difficult to follow with the eye (Larsen, 1991c). It often flies along roads and paths (Kielland, 1990d). Early stages:

Cock, 2016: 332 [egg; leaf shelters, larva, pupa; parasitoids]

Larval food: Dombeya burgessiae Gerrard ex Harv. (Malvaceae) [Cock, 2016; Kakamega, Kenya]. Dombeya species (Malvaceae) [Larsen, 1991c: 407]. Hermannia exappendiculata (Mast.) K. Schum. (Malvaceae) [Cock, 2016; Jadini Forest, Diani Beach, Kenya]. Melhania ovata (Cv.) Spreng. (Malvaceae) [Van Someren, 1974: 325; ssp. biifida]. Melhania rotundata Hochst. Ex Mast. (Malvaceae) [Cock, 2016; as Melhania taylori Baker f.; Kenya]. Melhania velutina Forssk. (Malvaceae) [Van Someren, 1974: 325’ ssp. bifida].

21

Ernsta (Delaga) zebra bifida (Higgins, 1925)

Hesperia bifida Higgins, 1925. Transactions of the Entomological Society of London 1924: 111 (64-120). Spialia zebra bifida (Higgins, 1925). Ackery et al., 1995. Ernsta (Delaga) bifida (Higgins, 1925). Zhang et al., 2020. Stat rev.

Ernsta bifida. Male. Left – upperside; right – underside. Ngong, Kenya. June 1979. SCC. ABRI-2019-2282. Images M.C. Williams ex ABRI Collection.

Ernsta bifida. Female. Left – upperside; right – underside. Nandi, Kenya. October 1971. SCC. ABRI-2019-2283. Images M.C. Williams ex ABRI Collection.

Type locality: [Kenya]: “Nairobi, Mombasa, British East Africa”; Uganda. Distribution: Yemen, Oman, Ethiopia, Somalia, Sudan (south), Democratic Republic of Congo, Uganda (east), Kenya, Tanzania (north). Specific localities: Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Mt Blue (Ducarme, 2018). Kenya – Nairobi (TL); Widespread (Larsen, 1991c); Mombasa (Higgins, 1925). Tanzania – Dar es Salaam (Kielland, 1990d); Singida (Kielland, 1990d); Ruaha National Park (Kielland, 1990d); Mount Meru (Kielland, 1990d); Oldeani (Kielland, 1990d); Mount Lolkisale (Kielland, 1990d); Himo River (Kielland, 1990d); Same, South Pare Mountains (Kielland, 1990d); Geita (Kielland, 1990d); Mweze Mountain in Mpanda (Kielland, 1990d); Moshi (Cordeiro, 1995).

* Ernsta (Delaga) sataspes (Trimen, 1864)# Boland Sandman

Pyrgus sataspes Trimen, 1864. Transactions of the Entomological Society of London (3) 2: 178 (175-180). Pyrgus sataspes Trimen. Trimen, 1866a. Pyrgus sataspes Trimen, 1864. Trimen & Bowker, 1889. Spialia sataspes Trimen. Swanepoel, 1953a.

22 Spialia sataspes (Trimen, 1864). Dickson & Kroon, 1978. Spialia sataspes (Trimen, 1864). Pringle et al., 1994: 317. Ernsta (Delaga) sataspes (Trimen, 1864). Zhang et al., 2020.

Ernsta sataspes. Male (Wingspan 24 mm). Left – upperside; right – underside. Baviaanskloof Mountains, Eastern Cape Province, South Africa. 15 December 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Ernsta sataspes. Female. Left – upperside; right – underside. Llandudno, Western Cape Province, South Africa. February 1965. C. Dickson ABRI-2019-2284. Images M.C. Williams ex ABRI Collection.

Alternative common name: Boland Sandmannetjie (Afrikaans). Type locality: [South Africa]: “Cape Colony and British Kaffraria”. Holotype in the Natural History Museum, London. Diagnosis: Closely resembles Ernsta delagoae, from which it may be distinguished on the hindwing underside by the absence of submarginal spots (Pringle et al., 1994). Can also be confused with Ernsta depauperata but on the underside of the hindwing the shape of the median band differs in the two species (Pringle et al., 1994).

Distribution: South Africa – Eastern Cape Province, Western Cape Province, Northern Cape Province.

Distribution: South Africa (Eastern Cape Province, Western Cape Province, Northern Cape Province). Specific localities: Eastern Cape Province – Addo (Swanepoel, 1953); Coega (Swanepoel, 1953); Grahamstown district

23 (Swanepoel, 1953); Bathurst (Swanepoel, 1953); Port Alfred (Swanepoel, 1953); Port Elizabeth (Pringle et al., 1994); Adelaide district (Pringle et al., 1994); Bedford district (Pringle et al., 1994). Western Cape Province – Cape Town, Muizenberg (Swanepoel, 1953); Llandudno (Swanepoel, 1953); Lion’s Head (Swanepoel, 1953); Ladismith (Swanepoel, 1953); Knysna (Swanepoel, 1953); Plettenberg Bay (Swanepoel, 1953); Signal Hill (Dickson & Kroon, 1978); Cape Peninsula (Pringle et al., 1994); . Habitat: Grassy areas in fynbos vegetation. Mainly on the lower slopes of mountains but occasionally in flat country. In the Eastern Cape Province it penetrates karroid country (Pringle et al., 1994). Habits: Both sexes are very fond of flowers. Males do not show hilltopping behaviour. They establish and defend territories on the slopes and use grass stems or low shrubs as perches. They are alert and restless (Pringle et al., 1994). Flight period: September to March (Pringle et al., 1994). Early stages:

Clark, in Dickson & Kroon, 1978: 228 plate 12 [as Spialia sataspes; recorded from eggs and larvae from Signal Hill, Cape Town, Western Cape Province]. “Egg: 0,75 mm diameter by 0,55 mm high; laid singly and concealed in young shoots. Whitish, with 20 heavy, irregular, longitudinal ribs, only some 6 to 7 of which reach the micropyle. There are some 25 very thin cross-ribs. Eggs change to very pale dull yellow and hatch after 11 days. The discarded shell is not eaten. Larva: 1st instar 1,25 to 2,5 mm in 14 days; 2 nd instar 2,5 to 3,5 mm in 9 days; 3 rd instar 3,5 to 6,5 mm in 9 days; 4 th instar 6,5 to 11 mm in 9 days; 5th instar 11 to 18 mm in 12 days. The duration of instars is very variable, depending upon climatic conditions. Young larvae feed on the surface of very young, half-opened leaves. Later, they bind more mature leaves together to form a ‘purse’, and feed on the surface of the leaf within this shelter, but they may crawl out and feed on adjacent leaves. Moulting and pupation take place within these shelters. Pupa: 11 mm. Secured by cremastral hooks to the inside of a shelter made generally of dried leaves near the ground. Emergence takes place after 16 to 20 days. Parasites: Attacked in the larval state by Tachinds, the maggot emerging from the side of the larva, which has turned a pale yellow; but there may be delayed emergence of the maggot, in which case it will find its way out of the side of the pupa. This species is multi-brooded but may hibernate in the pupal state in the colder part of the year.”

Larval food: Hermannia comosa Burch. ex DC. (Malvaceae) [Murray, 1959]. Hermannia cuneifolia Jacq. (Malvaceae) [Murray, 1959; as Hermannia pollens Eckl. and Zehy.]. Hermannia diffusa L. f. (Malvaceae) [Murray, 1959; as Hermannia pilosula Harv.]. Hermannia incana Cav. (Malvaceae) [Murray, 1959; as Hermannia candicans Ait.]. Hibiscus aethiopicus L. (Malvaceae) [Dickson & Kroon, 1978: 187]. Pavonia burchellii (DC.) R.A.Dyer (Malvaceae) [Murray, 1959; as Pavonia macrophylla E. Mey. ex Harv.].

* Ernsta (Delaga) depauperata (Strand, 1911)# Wandering Sandman

Hesperia ferax var. depauperata Strand, 1911. Mitteilungen aus dem Zoologischen Museum in Berlin 5: 304 (275-304). Spialia depauperata (Strand, 1911). Pringle et al., 1994: 317. Ernsta (Delaga) depauperata (Strand, 1911). Zhang et al., 2020.

24 Ernsta depauperata depauperata. Male. Left – upperside; right – underside. Kibale, Uganda. June 1993. IB, PW. ABRI-2019-2273. Images M.C. Williams ex ABRI Collection.

Ernsta depauperata depauperata. Female. Left – upperside; right – underside. Cholo Mountain, Malawi. September 1969. JCO Chitty. ABRI-2019-2274. Images M.C. Williams ex ABRI Collection.

Type locality: [Tanzania]: “Msamwia [Ufipa Plateau, south-western Tanzania]”. Diagnosis: Very similar to Spialia diomus but can be distinguished on the upperside of both wings by the absence of basal cell spots; on the forewing upperside by the presence of only one outer median spot (as opposed to two) in area 1B; on the underside of the hindwing by the series of submarginal spots starting at the end of vein 7 (as opposed to the end of vein 8) and by the costal edge of the median band, which is extended further outwards (Pringle et al., 1994). Also close to Ernsta zebra but in depauperata there is usually a discal spot in area 6 on the upperside of the hindwing (Larsen, 1991c). Distribution: Uganda, Kenya, Tanzania, Democratic Republic of Congo, Malawi, Zambia, Mozambique, Zimbabwe, Botswana, Namibia (Swart, 2004), South Africa, Swaziland. Habitat: Dry savanna (bushveld), especially Acacia dominated vegetation (Pringle et al., 1994). In Tanzania from 1 000 to 2 000 m (Kielland, 1990d). Habits: Flies low down in areas with short grass. Flowers and damp spots are both visited. Males do not hilltop and establish their territories in grassy areas usually perching on the ground (Pringle et al., 1994). Flight period: Mainly flies from August to April. In favourable seasons it may be found in any month of the year (Pringle et al., 1994). Early stages: Nothing published. Larval food: Melhania species (Malvaceae) [M.J.W. Cock, vide Larsen, 1991c: 407; erroneous (record was for Ernsta zebra (Cock, 2016) – the host plant(s) for S. depauperata is therefore unknown].

Ernsta (Delaga) depauperata depauperata (Strand, 1911)

Hesperia ferax var. depauperata Strand, 1911. Mitteilungen aus dem Zoologischen Museum in Berlin 5: 304 (275-304). Ernsta (Delaga) depauperata depauperata (Strand, 1911). Zhang et al., 2020.

Ernsta depauperata depauperata. Male. Left – upperside; right – underside.

25 Kibale, Uganda. June 1993. IB, PW. ABRI-2019-2273. Images M.C. Williams ex ABRI Collection.

Ernsta depauperata depauperata. Female. Left – upperside; right – underside. Cholo Mountain, Malawi. September 1969. JCO Chitty. ABRI-2019-2274. Images M.C. Williams ex ABRI Collection.

Type locality: [Tanzania]: “Msamwia”. Distribution: Uganda, Kenya, Tanzania, Democratic Republic of Congo, Malawi, Zambia, Mozambique. Specific localities: Kenya – Kibwezi (Larsen, 1991c); Mount Elgon area (Larsen, 1991c); Menfort (Larsen, 1991c). Tanzania – Msamwia (TL); Mpala (Oberthür, 1912); Ufipa (Kielland, 1990d); Mpanda (Kielland, 1990d); Kigoma (Kielland, 1990d); Kitesa Forest, west of Songea (Kielland, 1990d); Nyumbenito Mountain (Kielland, 1990d); Mufindi in the Uzungwa Range (Kielland, 1990d). Democratic Republic of Congo – Mpala region (Oberthür, 1912). Zambia – Solwezi (Heath et al., 2002); Mpongwe (Heath et al., 2002); Kamapanda (Heath et al., 2002); Petauke (Heath et al., 2002). Mozambique – Mount Mabu (Congdon et al., 2010).

rehfousi Oberthür, 1912 (as sp. of Syrichthus). Étude de Lépidoptérologie Comparée 6: 117 (47-120). [Democratic Republic of Congo]: “Région de M’pala, près du lac Tanganika”.

Ernsta (Delaga) depauperata australis (de Jong, 1978)# Wandering Sandman

Spialia abscondita Plötz. Swanepoel, 1953a. [Misidentification of Ernsta depauperata australis] Spialia depauperata australis de Jong, 1978. Tijdschrift voor Entomologie 121: 105 (23-146). Spialia depauperata australis De Jong, 1978. Pringle et al., 1994: 318. Ernsta (Delaga) depauperata australis (de Jong, 1978). Zhang et al., 2020.

Ernsta depauperata australis. Male (Wingspan 26 mm). Left – upperside; right – underside. Rustenburg Nature Reserve, North West Province, South Africa. M. Williams. Images M.C. Williams ex Williams Collection.

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Ernsta depauperata australis. Female (Wingspan 32 mm). Left – upperside; right – underside. Munnik, Limpopo Province, South Africa. 12 September 2004. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: South Africa: “Muden (Natal)”. Holotype in the Natural History Museum, London. Distribution: Mozambique, Zimbabwe, Botswana (north and east), Namibia (Swart, 2004), South Africa (Limpopo Province, Mpumalanga, North West, KwaZulu-Natal), Swaziland. Specific localities: Botswana – Lobatse (R. Plowes vide Larsen, 1991l). Namibia – Otavi (Swart, 2004). Limpopo Province – Munnik (Swanepoel); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015). North West Province – Rustenburg Nature Reserve (male specimen illustrated above). KwaZulu-Natal – Muden (TL).

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