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

Genus Swinhoe, [1912] Sandmen

In Swinhoe, [1912-3]. Indica 10: 99 (364 pp.). London. Type-: Hesperia galba Fabricius, by original designation [extralimital].

Name based on extralimital type-species: Powellia Tutt. [Invalid; junior homonym of Powellia Maskell.]

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

Spialia (Sandmen) is an Old World genus of small skippers with five Afrotropical species. One species (doris) extends extralimitally. The genus was monographed by De Jong in 1978 and the was revisited by Zhang et al. in 2020. These authors placed most of the species previously in Spialia in the newly erected genera Agyllia and Ernsta. Spialia occur in a variety of habitats but are found predominently in savanna. All species fly with a rapid, buzzing flight, low down. They are attracted to flowers, mud, dung and other foul substances. When feeding the wings are held three-quarters of the way open but at rest are held vertically, closed together. The males are territorial. The larvae have been recorded mainly on herbaceous in the dicotyledonous family . Spialia doris is exceptional in that its larvae utilize Convolvulaceae.

Subgenus Spialia Swinhoe, [1912]

In Swinhoe, [1912-3]. Lepidoptera Indica 10: 99 (364 pp.). London. Type-species: Hesperia galba Fabricius, by original designation [extralimital].

*Spialia (Spialia) mafa (Trimen, 1870)# Mafa Sandman

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Male of the Mafa Sandman (Spialia mafa). Images courtesy Raimund Schutte

Pyrgus mafa Trimen, 1870. Transactions of the Entomological Society of London 1870: 386 (341-390). Pyrgus mafa Trimen, 1870. Trimen & Bowker, 1889. Spialia mafa Trimen. Swanepoel, 1953a. Spialia mafa (Trimen, 1870). Dickson & Kroon, 1978. Spialia mafa (Trimen, 1870). Pringle et al., 1994: 320. Spialia (Spialia) mafa (Trimen, 1870). Zhang, et al., 2020.

Spialia mafa mafa. Male (Wingspan 24 mm). Left – upperside; right – underside. Loding, Mpumalanga Province, South Africa. 16 January 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Spialia mafa mafa. Female (Wingspan 25 mm). Left – upperside; right – underside. Roodepoort, Gauteng, South Africa. 24 August 2003. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [Lesotho]: “Maseru; Koro-Koro”; [South Africa]: “Potchefstroom, Trans-Vaal”. Holotype (male) in the Natural History Museum, London. Diagnosis: Similar to , from which it can be distinguished by the absence of a spot at the base of the costa and by its smaller markings (Pringle et al., 1994). Distribution: Angola, Democratic Republic of Congo, Sudan, Ethiopia, Somalia, Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, Botswana, Namibia, South Africa, Swaziland, Lesotho, Saudi Arabia, Yemen, Oman. Habitat: Grassland on the central plateau of southern Africa, and grassy areas in adjoining biomes (Karoo, fynbos and savanna). In the Western Cape Province it inhabits grassy places on coastal dunes. 2 In Tanzania subspecies higginsi occurs in open montane habitats from 1 200 to 2 400 m (Kielland, 1990d). Habits: A relatively common species (Larsen, 1991c) that occurs in localised colonies (Pringle et al., 1994). It flies close to the ground, in short grass, and frequently visits flowers (Pringle et al., 1994). Males do not hilltop but establish territories within the boundaries of the colony, using low plants, stones or the ground as perches. When defending their territories males are very pugnacious (Larsen, 1991c). Flight period: All year in warmer areas; September to April in high-lying and colder areas (Pringle et al., 1994). Early stages:

Clark, in Dickson & Kroon, 1978: 238; plate 17 [as Spialia mafa mafa; Milnerton, Western Cape]. “Egg: 0.65 mm diameter by 0.6 mm high; laid singly on young leaf-buds. There are 16 longitudinal ribs, composed chiefly of elliptical moles punctuating the junctions with 18 to 20 cross-ribs which form reversed diagonal bracing in the spaces between the main ribbing. The moles and ribs are white, on a pale green shell. Eggs hatch after some 8 days. The discarded shell is not eaten. Larva: 5-instar Group. 1st instar 1.25 growing to 2.5 mm in 8 days; 2nd instar 2.5 growing to 4.5 mm in 9 days; 3 rd instar 4.5 growing to 10 mm in 9 days; 4 th instar 10 growing to 14 mm in 9 days; 5th instar 14 growing to 21 mm in 16 days. 6-instar Group. 1st instar 1.25 growing to 2.5 mm in 9 days; 2nd instar 2.5 growing to 4 mm in 10 days; 3 rd instar 4 growing to 7 mm in 10 days; 4th instar 7 growing to 9 mm in 10 days; 5 th instar 9 growing to 12 mm in 10 days; 6 th instar 12 growing to 21 mm in 19 days. The young larva crawls into a half-open young leaf and feeds on the inner surface. Later, it constructs a ‘purse’ by binding leaves together. The larval and pupal state is spent within leaf-shelters. At first the larva feeds on the surface of a leaf but crawl out to feed on adjoining leaves. As it grows, it has to construct larger shelters and the last one is close to the ground. There is a rotation of broods, with the one in the cold part of the year very protracted. Pupa: 12 mm; at least slightly covered with white powder and secured by cremastral hooks to the inside of its shelter. Emergence takes place after some 18 to 25 days. Parasites: Parasitical attack is similar to that in most Spialia. Recorded from eggs from Milnerton, C.P.”

Larval food: comosa Burch. ex DC. (Malvaceae) [Murray, 1959; as camosa auct.; for ssp. mafa]. Hermannia cuneifolia Burch. ex DC. (Malvaceae) [Murray, 1959; as pollens Eckl. and Zehy.; for ssp. mafa]. Hermannia depressa N.E.Br. (Malvaceae) [Henning, G., in Pringle et al., 1994: 320; for ssp. mafa]. Hermannia diffusa L.f. (Malvaceae) [Murray, 1959; as pilosula Harv.; for ssp. mafa]. Hermannia incana Cav. (Malvaceae) [Murray, 1959; as candicamus auct.; for ssp. mafa]. Hermannia spp. (Malvaceae) [Dickson & Kroon, 1978: 189]. aethiopicus L. (Malvaceae) [Murray, 1959]. Hibiscus ludwigii Eckl. & Zeyh. (Malvaceae) [Van Someren, 1974: 325; as Hibiscus macrantha]. Hibiscus macranthus Hochst. Ex A. Rich. (Malvaceae) [Van Someren, 1974; for ssp. higginsi]. Hibiscus species (Malvaceae) [Sevastopulo, 1975; for ssp. higginsi]. arabica Hochst. & Steud. Ex Boiss (Malvaceae) [Otto, 2016: 44; Oman; oviposition only; for ssp. higginsi]. Pavonia burchellii (DC.) R.A.Dyer (Malvaceae) [Murray, 1959; as macrophylla E. Mey. ex Harv.; for ssp. mafa]. Pavonia species (Malvaceae) [Larsen, 1991c: 405]. ovata Forssk. (Malvaceae) [Van Someren, 1974: 325; as Sida grewioides Guill. & Perr.; for ssp. higginsi]. Sida rhombifolia L. (Malvaceae) [Van Someren, 1974: 325; for ssp. higginsi]. Sida schimperiana Hochst. ex A.Rich. and/or Sida tenuicarpa Vollesen (Malvaceae) [Van Someren, 1974: 325; as Sida cuneifolia Roxburgh (which refers to two possible species of Sida)]. Sida sp. (Malvaceae) [Sevastopulo, 1975; for ssp. higginsi]. Sida tenuicarpa Vollesen (Malvaceae) [Van Someren, 1974: 325; as cuneifolia Roxburgh; for ssp. higginsi].

Spialia (Spialia) mafa mafa (Trimen, 1870)# Mafa Sandman

Pyrgus mafa Trimen, 1870. Transactions of the Entomological Society of London 1870: 386 (341-390). 3 Spialia mafa Trimen. Swanepoel, 1953a. Spialia mafa mafa (Trimen, 1870). Dickson & Kroon, 1978. Spialia mafa mafa (Trimen, 1870). Pringle et al., 1994: 320. Spialia (Spialia) mafa mafa (Trimen, 1870). Zhang, et al., 2020.

Spialia mafa mafa. Male (Wingspan 24 mm). Left – upperside; right – underside. Loding, Mpumalanga Province, South Africa. 16 January 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Spialia mafa mafa. Female (Wingspan 25 mm). Left – upperside; right – underside. Roodepoort, Gauteng, South Africa. 24 August 2003. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [Lesotho]: “Maseru; Koro-Koro”; [South Africa]: “Potchefstroom, Trans-Vaal”. Holotype (male) in the Natural History Museum, London. Distribution: Angola, Democratic Republic of Congo (south), Malawi, Zambia (south and south-east), Mozambique, Zimbabwe, Botswana, Namibia (central and north), South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, Free State Province, KwaZulu-Natal, Eastern Cape Province, Western Cape Province, Northern Cape Province), Swaziland, Lesotho. Specific localities: Angola – Huambo Province (Mendes et al., 2013). Malawi – Mount Mulanje (Congdon et al., 2010). Zambia – Livingstone (Heath et al., 2002); Chilanga (Heath et al., 2002); Chalimbana (Heath et al., 2002); Luangwa Valley (Heath et al., 2002); Chipata (Heath et al., 2002). Botswana – Widespread (Larsen, 1991l); Nosop River (Larsen, 1991l); Kgaligadi Transfrontier Park (Larsen, 1991l); Shakawe (Larsen, 1991l); Kazungula (Larsen, 1991l). Namibia – Swakop River, in the Namib Desert (Bampton). Limpopo Province – Polokwane (Swanepoel, 1953); Dendron (Swanepoel, 1953); Highlands Wilderness (Bode & Bode, unpublished checklist); Bateleur Nature Reserve (Williams & Dobson, unpub., 2015). Mpumalanga – Balfour (Swanepoel, 1953); Barberton (Swanepoel, 1953); Crocodile River (Swanepoel, 1953); Percy Fyfe Nature Reserve (Warren, 1990); Argent (male illustrated above). North West Province – Potchefstroom (Trimen, 1870); Utopia Resort (C. Dobson, 2006); Borakalalo Nature Reserve (J. Dobson, unpublished, 2009). Gauteng – Pretoria (Swanepoel, 1953). Free State Province – Ladybrand (Swanepoel, 1953); Bloemfontein (Swanepoel, 1953). KwaZulu-Natal – Howick (Swanepoel, 1953); Impenza (Swanepoel, 1953); Spitzkop (Swanepoel, 1953); Montello (Swanepoel, 1953); Bergville (Swanepoel, 1953); Estcourt (Swanepoel, 1953); Greytown (Swanepoel, 1953); Weenen (Swanepoel, 1953); Ladysmith (Swanepoel, 1953).

4 Eastern Cape Province – Burgersdorp (Swanepoel, 1953). Western Cape Province – Mamre (Swanepoel, 1953); Tygerberg (Swanepoel, 1953); Malmesbury (Swanepoel, 1953); Durbanville (Swanepoel, 1953); Stellenbosch (Swanepoel, 1953); Milnerton (Dickson & Kroon, 1978). Northern Cape Province – Kimberley (Aurivillius, 1925); Twee Rivieren (Pringle et al., 1994); Tampan (Pringle et al., 1994); Olifantshoek (Pringle et al., 1994); Hotazel (Pringle et al., 1994); Griquatown (Pringle et al., 1994); Wallekraal, near Hondeklip Bay (Pringle et al., 1994). Lesotho – Maseru (Trimen, 1870); Koro Koro (Trimen, 1870).

oberthuri Aurivillius, 1925 in Seitz, 1908-25 (as sp. of Hesperia). Die Gross-Schmetterlinge der Erde, Stuttgart (2) 13 Die Afrikanischen Tagfalter: 565 (614 pp.). South Africa: “Kimberley”.

aurivillii Shepard, 1935 (as replacement name for Hesperia oberthuri Aurivillius). In: Strand, E., Lepidopterorum Catalogus 22 (69): 496 (273-560). ‘s-Gravenhage.

Spialia (Spialia) mafa higginsi Evans, 1937

Spialia mafa higginsi Evans, 1937. A catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum: 62 (212 pp.). Spialia (Spialia) mafa higginsi Evans, 1937. Zhang, et al., 2020.

Spialia mafa higginsi. Male. Left – upperside; right – underside. Nairobi, Kenya. 16 July 2003. Images M.C. Williams ex Gardiner Collection.

Type locality: [Kenya]: “Mumias, 4,000 ft”. Distribution: Sudan (south), Ethiopia, Somalia, Kenya (central, west), Tanzania, Saudi Arabia (south- west), Yemen, Oman. Specific localities: Kenya – Mumias (TL). Tanzania – Northern Highlands (Kielland, 1990d); Njombe (Kielland, 1990d); Ufipa (Kielland, 1990d); Kigoma (Kielland, 1990d); Mpanda (Kielland, 1990d).

*Spialia (Spialia) spio (Linnaeus, 1764)# Mountain Sandman

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Mountain Sandman (Spialia spio). Left – male upperside. Right – male underside. Images courtesy Steve Woodhall

Papilio spio Linnaeus, 1764. Museum Ludovicae Ulricae Reginae: 338 (720 pp.). Holmiae. [According to Ackery et al., 1995]. Papilio spio Linnaeus, 1767. Systema Naturae 1 (2) 12th edn: 533-1328 + [36] pp. Holmiae. [According to de Jong, 1978]. Pyrgus vindex Cramer. Trimen, 1866a. [synonym of Spialia spio] Pyrgus vindex (Cramer, 1782). Trimen & Bowker, 1889. [synonym of Spialia spio] Spialia spio Linnaeus. Swanepoel, 1953a. Spialia spio (Linnaeus, 1767). Dickson & Kroon, 1978. [date of authorship erroneous] Spialia spio (Linnaeus, 1767). Pringle et al., 1994: 319. [date of authorship erroneous] Spialia (Spialia) spio (Linnaeus, 1764). Zhang, et al., 2020.

Spialia spio. Male (Wingspan 26 mm). Left – upperside; right – underside. Tswaing Crater, Gauteng, South Africa. 3 November 2002. M. Williams. Images M.C. Williams ex Williams Collection.

Spialia spio. Female (Wingspan 29 mm). Left – upperside; right – underside. Kwamhalanga, Mpumalanga Province, South Africa. 4 December 2010. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [South Africa]: “Cap. b. spei”. The types have apparently been lost. Lectotype designated by Honey & Scoble, 2001: 381. Diagnosis: On the upperside of the wings there is an additional spot between the median and submarginal series along the inner margin. There is also a white spot at the base of the forewing costa (Pringle et al., 1994). See also Spialia mafa for differentiation from this species. Distribution: Sub-Saharan Africa, including Mauritania, Senegal, Gambia, Guinea-Bissau (Aurivillius, 1910), Guinea, Mali, Sierra Leone, Liberia, Ivory Coast, Burkina Faso, Ghana, Togo, Benin (south, north), Nigeria, Niger, Gabon (de Jong, 1978), Angola, Democratic Republic of Congo, Uganda, 6 Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, Botswana, Namibia, South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, Free State Province, KwaZulu- Natal, Eastern Cape Province, Western Cape Province, Northern Cape Province), Swaziland, Lesotho, Saudi Arabia (south-west), Yemen. Specific localities: Gambia – Brufut, Abuko, Pirang, Brusubi, Brufut, Tujering, Kartong, Madiana, Sukuta, Keneba, Sapu, Niani, Tendaba, Basse (Jon Baker, pers. comm, May 2020). Benin – Lokoli (Tchibozo et al., 2008); Houeyogbe Forest (Coache & Rainon, 2016); see Coache et al., 2017. Angola – Bengo Province; Benguela Province; Bie Province; Huambo Province; Huila Province; Kuando-Kubango Province; Kwanza Norte Province; Luanda Province; Malanje Province; Moxico Province 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). Kenya – widespread (Cock, 2016); Chyulu Hills (van Someren, 1939). Tanzania – Throughout (Kielland, 1990d); Mount Longido (Kielland, 1990d). Malawi – Nyika N.P. (J. Timberlake, pers. comm., 2019). Zambia – Mwinilunga (Heath et al., 2002); Kabompo River (Heath et al., 2002); Chingola (Heath et al., 2002); Mufulira (Heath et al., 2002); Ndola (Heath et al., 2002); Mpongwe (Heath et al., 2002); Chalimbana (Heath et al., 2002); Luangwa Valley (Heath et al., 2002); Luongo River (Heath et al., 2002); Mpika (Heath et al., 2002); Mbala (Heath et al., 2002); Nyika (Heath et al., 2002). Mozambique – Mount Namuli (Congdon et al., 2010); Mt Mecula [-12.0772 37.6297] (Congdon & Bayliss, 2013). Botswana – Widespread (Larsen, 1991l); Tshabong (Larsen, 1991l). Limpopo Province – Warmbaths (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Zoutpansberg (Swanepoel, 1953); Haenertsburg (Swanepoel, 1953); Munnik (Swanepoel, 1953); The Downs in Lekgalameetse Nature Reserve (“Malta Forest”) (Swanepoel, 1953); Soetdoring Farm [- 24.561 28.233] (A. Mayer, pers comm. 2015); Bateleur Nature Reserve (Williams & Dobson, unpub., 2015). Mpumalanga – Graskop (Swanepoel, 1953); Lydenburg (Swanepoel, 1953); Barberton (Swanepoel, 1953). North West Province – Potchefstroom (Swanepoel, 1953); Borakalalo Nature Reserve (J. Dobson, unpublished, 2009). Gauteng – Krugersdorp (Swanepoel, 1953); Pretoria (Swanepoel, 1953; male specimen illustrated above). Free State Province – Bloemfontein (Swanepoel, 1953); Ladybrand (Swanepoel, 1953); Kroonstad (Swanepoel, 1953); Cyferfontein [-30.3736 25.8131] (R. Griesel, unpublished). KwaZulu-Natal – Umkomaas (Swanepoel, 1953); Durban (Swanepoel, 1953); Hluhluwe (Swanepoel, 1953); Vryheid (Swanepoel, 1953); Greytown (Swanepoel, 1953); Estcourt (Swanepoel, 1953); Ladysmith (Swanepoel, 1953); Loteni (Swanepoel, 1953); Giant’s Castle (Swanepoel, 1953); Kosi Bay Nature Reserve (Pringle & Kyle, 2002). Eastern Cape Province – Port Elizabeth (Swanepoel, 1953); Cookhouse (Swanepoel, 1953); Grahamstown (Swanepoel, 1953); King William’s Town (Swanepoel, 1953); East London (Swanepoel, 1953); Dordrecht (Swanepoel, 1953); Molteno (Swanepoel, 1953); Steynsburg (Swanepoel, 1953). Western Cape Province – Cape Town (Swanepoel, 1953); Worcester (Swanepoel, 1953); Oudtshoorn (Swanepoel, 1953); Knysna (Swanepoel, 1953); Signal Hill (Dickson & Kroon, 1978); Lion's Head (Dickson & Kroon, 1978). Northern Cape Province – Carnarvon (Swanepoel, 1953); Kimberley (Swanepoel, 1953). Swaziland – Mlawula N. R. (www.sntc.org.sz); Malolotja N. R. (www.sntc.org.sz). Habitat: Occurs in a wide range of habitats, from clearings in forest to savanna woodland, thornveld and grassland. Absent from very dry (desert and semidesert) habitats (Pringle et al., 1994) and from clear-felled wetter forest (Larsen, 2005a). It seems to prefer the lower slopes of hills and valleys (Pringle et al., 1994). In Tanzania at altitudes from near sea-level to 2 600 m (Kielland, 1990d). Habits: A very common and widespread (Larsen, 2005a). Flies with a rapid, buzzing flight, close to the ground. Specimens rest and bask on low vegetation, or on the ground, with the wings held three-quarters open (Larsen, 2005a). Males will often return to a favoured perch (Pringle et al., 1994).

7 Males hilltop during the warmer hours of the day but also establish territories on lower ground. Both flowers and muddy spots attract specimens (Pringle et al., 1994). Occasionally adults feed from herbivore dung, including donkey dung (Cock, 2016). Flight period: All year, but scarce in cold climes in the winter months (Pringle et al., 1994). Early stages:

Clark, in Dickson & Kroon, 1978: 236; plate 16 [as Spialia spio; Signal Hill and Lion's Head, Cape Town, Western Cape]. “Egg: 0.75 mm diameter by 0.7 mm high; laid singly among young shoots. Pale green, with thin white ribbing consisting of some 18 irregular longitudinal ribs and some 16 to 20 fine, reversed diagonal cross-ribs as in the previous species [ ferax], these being punctuated at the intersections by elliptical moles, which represent the main part of the longitudinal ribs. Eggs hatch after 7 days. The discarded shell is not eaten. Larva: 5-instar Group. 1st instar 1.5 growing to 3 mm in 11 days; 2 nd instar 3 growing to 5 mm in 8 to 20 days; 3 rd instar 5 growing to 9 mm in 5 to 20 days; 4 th instar 9 growing to 12 mm in 9 to 20 days; 5 th instar 12 growing to 21 mm in 15 to 20 days. 6-instar Group. 1st instar 1.5 growing to 3 mm in 11 days; 2 nd instar 3 growing to 4.5 mm in 8 to 20 days; 3rd instar 4.5 growing to 7 mm in 5 to 20 days; 4 th instar 7 growing to 10 mm in 9 to 20 days; 5 th instar 10 growing to 17 mm in 15 to 20 days; 6 th instar 17 growing to 21 mm in 15 to 20 days. Six instars are generally passed through in dry seasons. When the foodplant is fresh there are 5 instars of shorter total duration; but the duration of individual instars is variable. Larvae are not gregarious. They feed at first on the surface of a half- opened, young leaf, but later draw leaves together to form a ‘hide-out’. They continue to feed on the surface of the leaf but may crawl out to feed on adjacent leaves. Pupation takes place within the final shelter, generally near the ground. Pupa: 12 mm; covered with a white powder, and secured by its cremastral hooks. Emergence takes place after some 18 days. Parasites: Parasitic attack by species of Tachinidae has been recorded. One of the Diptera, so far identified, has been Thecocarcelia latifrons, ex pupa. This species is multi-brooded. Recorded from eggs and larvae from Signal Hill and Lion’s Head, Cape Town.”

Henning, Henning, Joannou & Woodhall, 1997: 115 [photograph of final instar larva and pupa].

Cock, 2016: 311 [leaf shelters, larva, pupa, parasitoids]

Final instar larva and pupa of Spialia spio. Images courtesy Allison Sharp.

Larval food: Althaea sp. (Malvaceae) [LePelley, 1959; Uganda]. Hermannia coccocarpa (Eckl. & Zeyh.) Kuntze (Malvaceae) [Dickson & Kroon, 1978; South Africa]. Hermannia comosa Burch. ex DC. (Malvaceae) [Murray, 1959; South Africa]. Hermannia cuneifolia Jacq. (Malvaceae) [Murray, 1959; as Hermannia pallius Eckl. and Zeyh.; South Africa]. Hermannia diffusa L.f. (Malvaceae) [Murray, 1959; as Hermannia pilosula Harv.; South Africa]. Hermannia incana Cav. (Malvaceae) [Murray, 1959; as Hermannia candicans Ait.]. Hibiscus aethiopicus L. (Malvaceae) [Gifford, 1965; Malawi]. Hibiscus asper Hook f. (Malvaceae) [Vuattoux, 1999; Ivory Coast]. Hibiscus calyphyllus Cav. (Malvaceae) [Cock, 2016:310; Kenya]. Hibiscus cannabinus L. (Malvaceae) [Le Pelley, 1959; Uganda]. Hibiscus diversifolius Jacq. (Malvaceae) [LePelley, 1959; Uganda]. Hibiscus fuscus Garke (Malvaceae) [Van Someren, 1974; as Hibiscus gossypinus auct.; EastAfrica]. Hibiscus pusillus Thunb. (Malvaceae) [Platt, 1921; as Hibiscus gossypinus sensu Harv. non Thunb. = Hermannia cus L.]. Hibiscus pusillus Thunb. (Malvaceae) [Van Someren, 1974: 325; as Hibiscus gossyphina]. Hibiscus rostellatus Guill. & Perr. (Malvaceae) [Vuattoux, 1999; Ivory Coast]. Hibiscus sp. (Malvaceae) [Le Pelley, 1959; Kenya]. 8 (L.) Webb & Berthel. (Malvaceae) (exotic) [Dickson & Kroon, 1978; as Lavatera arborea L.; South Africa]. Malva species (Malvaceae) [Larsen, 1991c: 404; as Lavatera sp.]. Malvaceae [Fontaine, 1988; DRC]. Pavonia burchellii (DC.) R.A.Dyer (syn. patens Andr.) (Malvaceae) [Platt, 1921; as Pavonia macrophylla E. Mey. ex Harv.; South Africa]. Pavonia columella Cav. (Malvaceae) [Murray, 1959; South Africa]. Pavonia sp. (Malvaceae) [Sevastopulo, 1975; East Africa]. Pavonia urens Cav. var. tomentosa (Hochst. Ex Ulbr.) Brenen (Malvaceae) [Cock, 2016: 310; Kenya; in captivity]. Pavonia urens Cav. var. urens (Malvaceae) [Cock, 2016: 310; Kenya]. Sida cordifolia L. (Malvaceae) [Van Someren, 1974: 325; East Africa]. Sida ovata Forssk. (Malvaceae) [Van Someren, 1974: 325; as Sida grewioides; East Africa]. Sida rhombifolia L. (Malvaceae) [Sevastopulo, unpub.; Uganda]. Sida schimperiana Hochst. ex A.Rich. (Malvaceae) [Van Someren, 1974: 325; East Africa]. Sida schimperiana Hochst. ex A.Rich. and/or Sida tenuicarpa Vollesen (Malvaceae) [Van Someren, 1974: 325; as Sida cuneifolia Roxburgh (which refers to two possible species of Sida)]. Sida species (Malvaceae) [Van Someren, 1939; East Africa]. Sida tenuicarpa Vollesen (Malvaceae) [Van Someren, 1974; as Sida cuneifolia auct.; East Africa]. rhomboidea Jacq. (Malvaceae) [Vuattoux, 1999; Ivory Coast]. Triumfetta species (Malvaceae) [Dickson & Kroon, 1978; South Africa]. Triumfetta tomentosa Bojer (Malvaceae) [Cock, 2016: 310; Kenya]. Urena lobata L. (exotic) (Malvaceae) [Vuattoux, 1999; Ivory Coast]. indica L. (Malvaceae) [Cock, 2016: 311; Benin]. Wissadula amplissima (L.) R. E. Fr. (Malvaceae) [Vuattoux, 1999; as Wissadula rostrata (Schumach.) Hook.f.; Ivory Coast].

vindex Stoll, 1781 in Stoll, [1780-2] (as sp. of Papilio). Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en America 4 [part]: 122 (29-252). Amsteldam & Utrecht. [South Africa]: “Kaap de Goede Hoop”.

Subgenus Platygnathia Picard, 1948

Ref? Type-species: ? phlomidis Herrich-Schäffer [extralimital].

*Spialia (Platygnathia) doris (Walker, 1870) Desert Grizzled Skipper

Nisoniades doris Walker, 1870. Entomologist 5: 56 (48-57). Spialia doris (Walker, 1870). Ackery et al., 1995. Spialia (Platygnathia) doris (Walker, 1870). Zhang, et al., 2020.

Type locality: [Djibouti]: “Tajora [Tadjoura]”. Distribution: Mauritania, Sudan, Kenya, Somalia, Djibouti, Saudi Arabia, Yemen, Oman. Extralimitally in Morocco (ssp. daphne Evans), Egypt (ssp. amenophis (Reverdin)), southern Iran to Gujurat, India (ssp. evanida (Butler). Habitat: Dry water courses and rocky ground in semi-desert (eremic habitats) (Larsen, 1991c; Larsen, 2005a). Habits: Specimens are attracted to flowers (Larsen, 1991c). Early stages: Nothing published. Larval food: Convolvulus hystrix Vahl (Convolvulaceae) [Pittaway, 1985; western Saudi Arabia; ssp. doris]. Convolvulus lanatus Vahl (Convolvulaceae) [Graves, 1925 (Transactions of the Entomological Society

9 of London, 1925: 17-125); Egypt; ssp. amenophis]. Convolvulus species (Convolvulaceae) [Larsen, 1991c: 405]. Corchorus depressus (L.) Stocks (Malvaceae) [Pittaway, 1980 & 1985; Quatar; ssp, doris]. Corchorus species (Malvaceae) [Larsen, 1991c: 405]. Ipomoea imperati (Vahl) Griseb. (Convolvulaceae) [Benyamini, 1984; as stolonifera (Cirillo) J.F. Gmel.; Sinai Peninsula]. Ipomoea species (Convolvulaceae) [Larsen, 2005a].

Spialia (Platygnathia) doris doris (Walker, 1870)

Nisoniades doris Walker, 1870. Entomologist 5: 56 (48-57). Spialia (Platygnathia) doris doris (Walker, 1870). Zhang, et al., 2020.

Type locality: [Djibouti]: “Tajora [Tadjoura]”. Distribution: Sudan, Kenya (north), Somalia, Djibouti, Saudi Arabia, Yemen, Oman. Extralimital in Arabia, southern Iran and Pakistan. Specific localities: Djibouti – Tajora [Tadjoura] (TL). Yemen – Aden (Butler, 1885). Kenya – Tana River between Garissa and Bura (De Jong, 1978).

adenensis Butler, 1885 (as var. of Pyrgus evenidus). Proceedings of the Zoological Society of London 1884: 493 (478-503). Yemen: “Aden”.

Spialia (Platygnathia) doris daphne Evans, 1949

?Spialia doris daphne Evans, 1949. ?Annals and Magazine of Natural History (12) 2: 54-56. Spialia (Platygnathia) doris daphne Evans, 1949. Zhang, et al., 2020.

Type locality: ? Distribution: Mauritania. Extralimitally in south-western Morocco. Specific localities: Mauritania – Akjoujt, 250 km N of Nouakchott (H. Warren-Gash, vide Larsen, 2005a).

*Spialia (Platygnathia) diomus (Hopffer, 1855) Common Sandman

Pyrgus diomus Hopffer, 1855. Berichte über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1855: 643 (639-643). Pyrgus diomus Hopffer, 1855. Trimen & Bowker, 1889. [referable to ferax] Spialia diomus Hopffer, 1855. Higgins, 1925. Spialia diomus diomus Hopffer, 1855). Evans, 1937. Spialia diomus Hopffer. Swanepoel, 1953a. [referable to ferax] Spialia diomus ferax (Hopffer, 1855). Pringle et al., 1994: 319. Spialia diomus (Hopffer, 1855). Cock, 2016. Spialia (Platygnathia) diomus (Hoppfer, 1855). Zhang, et al., 2020.

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Spialia diomus. Male. Left – upperside; right – underside. Kenya. Images courtesy Torben Larsen.

Spialia diomus. Female. Left – upperside; right – underside. Arabuko-Sokoke Forest, Kenya. June 1997. SCC. ABRI-2019-2276. Images M.C. Williams ex ABRI Collection.

Type locality: Mozambique: “Mossambique”. Distribution: Senegal, Gambia, Guinea, Liberia, Ivory Coast, Ghana, Benin (south, north-west), Nigeria, Democratic Republic of Congo, Uganda, Kenya, Tanzania, Zambia (Cock, 2016), Mozambique (north) (Cock, 2016), Yemen. Recorded, in error for ferax, from Mozambique by Congdon et al., 2010. Specific localities: Gambia – Pirang, Abuko, Brufut, Marakissa, Tujering, Dumbutu, Tampoto, Keneba, Tintinto, Basse (Jon Baker, pers. comm, May 2020). Ghana – Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – see Coache et al., 2017. Democratic Republic of Congo – Ituri Forest (Ducarme, 2018); Mt Blue (Ducarme, 2018). Kenya – Machako’s (Butler, 1899). Tanzania – Central, north and east (Kielland, 1990d); Ngara District (Kielland, 1990d); Katavi National Park (Fitzherbert et al., 2006). Habitat: All habitats except forest. Predominantly in savanna. In Tanzania it found at altitudes from near sea-level to 2 000 m (Kielland, 1990d). Habits: A common and widespread Afrotropical sandman (Larsen, 1991c). Males are attracted to a variety of foul matter, including fresh cow pats, skins being cured in the sun, carcasses, and leopard scat (Larsen, 2005a). Cock (2026) records it feeding from fresh elephant dung in Kenya. They are also known to mud-puddle (Cock, 2016). Early stages:

Cock, 2016: 314 [egg, leaf shelters, larva, pupa; parasitoids]

Larval food: Hermannia spp. (Malvaceae) [Sevastopulo, 1975; East Africa]. Hibiscus species (Malvaceae) [Sevastopulo, 1975; EastAfrica]. Pavonia species (Malvaceae) [Larsen, 1991; East Africa]. Pavonia burchellii (DC.) R.A.Dyer (Malvaceae) [Kielland, 1990; East Africa; as Pavonia macrophylla E. Mey. ex Harv.].

11 Sida ovata Forssk. (Malvaceae) [Cock, 2016; Kenya]. Sida rhombifolia L. (Malvaceae) [Cock, 2016; Kenya; in captivity.] Sida species (Malvaceae) [Le Pelley, 1959; East Africa]. Triumfetta brachyceras K.Schum. (Malvaceae) [Van Someren, 1974: 325; as Triumfetta macrophylla E. Mey. ex Harv.]. Triumfetta rhomboidea Jacq. (Malvaceae) [Larsen, 2005a]. Triumfetta species (Malvaceae) [Van Someren, 1974]. Waltheria indica L. (Malvaceae) [Cock, 2016; Kenya, Rwanda, Benin]. Waltheria species (Malvaceae) [Larsen, 1991c: 404; Kenya].

machacoana Butler, 1899 (as sp. of Pyrgus). Proceedings of the Zoological Society of London 1899: 426 (417-427). Kenya: “Machako’s”.

*Spialia (platygnathia) ferax (Wallengren, 1863)# Striped Sandman

Males of Spialia ferax. Images courtesy Allison Sharp.

Syrichtus [sic] ferax Wallengren, 1863. Wiener Entomologische Monatschrift 7: 137 (137-151. Pyrgus diomus Hopffer, 1855. Trimen & Bowker, 1889. [referable toferax] Spialia ferax Wallengren, 1863). Higgins, 1925. Spialia diomus ferax (Wallengren, 1863). Evans, 1937. Spialia diomus Hopffer. Swanepoel, 1953a. [referable to ferax] Spialia diomus ferax (Wallengren, 1863). Dickson & Kroon, 1978. Spialia diomus ferax (Wallengren, 1863). Pringle et al., 1994: 319. Spialia ferax (Wallengren, 1863). Cock, 2016. Stat. rev. Spialia (Platygnathia) ferax (Wallengren, 1863). Zhang, et al., 2020.

Spialia ferax. Male (Wingspan 29 mm). Left – upperside; right – underside. Kwamahlanga, Mpumalanga Province, South Africa. 5 December 2010. M. Williams. Images M.C. Williams ex Williams Collection.

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Spialia ferax. Female (Wingspan 31 mm). Left – upperside; right – underside. Hennops River, Gauteng Province, South Africa. 7 August 2002. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [Namibia]: “in territorio fluminis Kuisip”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/f). Diagnosis: The light band on the hindwing underside is broad and curved (Pringle et al., 1994). Distribution: Tanzania, Malawi, Zambia, Angola, Mozambique, Zimbabwe, Botswana, Namibia, South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, Free State Province, KwaZulu-Natal, Eastern Cape Province, Western Cape Province, Northern Cape Province), Swaziland, Lesotho. Specific localities: Tanzania – Bisi Mountain, Ufipa, 2 200 m (Kielland, 1990d; single female). Malawi – Mount Mulanje (Congdon et al., 2010). Zambia – Solwezi (Heath et al., 2002); Ndola (Heath et al., 2002); Mansa (Heath et al., 2002); Shiwa Ngandu (Heath et al., 2002); Mbala (Heath et al., 2002). Angola – Huila Province (Mendes et al., 2013). Mozambique – Njesi Plateau (Congdon et al., 2010); Mount Inago (Congdon et al., 2010); Mt Mecula [-12.0772 37.6297] (Congdon & Bayliss, 2013). Botswana – Widespread (Larsen, 1991l); Kgalagadi Transfrontier Park (Larsen, 1991l); Motokwe (Larsen, 1991l); Molopo Valley (Larsen, 1991l). Namibia – Kuisip River (TL). Limpopo Province – Throughout (Swanepoel, 1953); Zoutpansberg (Oberthür, 1912); Warmbaths (Swanepoel, 1953); Potgietersrus (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Munnik (Swanepoel, 1953); Letaba (Swanepoel, 1953); Percy Fyfe Nature Reserve (Warren, 1990); Lekgalameetse Nature Reserve (“Malta Forest”); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015); Bateleur Nature Reserve (Williams & Dobson, unpub., 2015). Mpumalanga – Throughout (Swanepoel, 1953); Lydenburg (Swanepoel, 1953); Buffelspoort Nature Reserve (Williams). North West Province – Throughout (Swanepoel, 1953); Utopia Resort (C. Dobson, 2006); Borakalalo Nature Reserve (J. Dobson, unpublished, 2009). Gauteng – Throughout (Swanepoel, 1953); Witwatersrand Botanical Gardens (J. Dobson, unpublished checklist, 2001). Free State Province – Throughout (Swanepoel, 1953); Platberg, Harrismith (Dobson, Williams & Schutte, unpublished, 2010); Cyferfontein [-30.3736 25.8131] (R. Griesel, unpublished). KwaZulu-Natal – Throughout (Swanepoel, 1953); Estcourt (Swanepoel, 1953); Ladysmith (Swanepoel, 1953); Muden (Swanepoel, 1953; male specimen illustrated above); Tugela Estates (Swanepoel, 1953); Tembe Nature Reserve (Pringle & Kyle, 2002). Eastern Cape Province – Throughout (Swanepoel, 1953); Port Elizabeth (Dickson & Kroon, 1978). Western Cape Province – Throughout (Swanepoel, 1953). Northern Cape Province – Throughout (Swanepoel, 1953). Swaziland – Mlawula N. R. (www.sntc.org.sz). Habitat: All habitats except forest. Predominantly in savanna. In Tanzania ferax is found at altitudes around 2 200 m (Kielland, 1990d). Habits: Flies rapidly, close to the ground. Both sexes are readily attracted to flowers (Pringle et al., 1994). Males hilltop, selecting perches on bare ground, and vigorously defending their territories against intruders. They may defend such territories for most of the warmer hours of the day. Territories may also be established on the slopes and flats (Pringle et al., 1994).

13 Flight period: All year in warm areas; August to April in colder climates (Pringle et al., 1994). Early stages:

Clark, in Dickson & Kroon, 1978: 234; plate 15 [as Spialia diomus ferax; recorded from eggs and larvae from near Port Elizabeth, Eastern Cape Province]. “Egg: 0,75 mm diameter by 0,7 mm high; laid singly on young shoots. It is pale blue-green, with some 16 longitudinal ribs, cross-braced by about 18 very fine ribs. The junctions of the ribs are capped by elliptical moles. Eggs hatch after some 10 days. The discarded shell is not eaten. Larva: 1st instar 1,75 to 3,25 mm in 8 days; 2nd instar 3,25 to 4,5 mm in 5 to 7 days; 3 rd instar 4,5 to 8,25 mm in 8 to 9 days; 4 th instar 8,25 to 13 mm in 13 days; 5th instar 13 to 22 mm in 20 days. Duration of instars variable. Young larvae at first feed on the surface of a partially opened, young leaf. They form ‘hide-outs’ by drawing the leaf edges together with silk. Later, leaves are spun together, and feeding may occur on the edges of adjacent leaves. The whole larval state is spent within the protection of these shelters. Pupation also takes place in them. Pupa: 13,5 mm secured by cremastral hooks. Emergence occurs after 15 to 33 days. Hibernation may tide the pupa over winter, especially in cold areas. Parasites: Larvae are chiefly attacked by species of Tachinidae. The maggot emerges either from the larva or pupa and it pupates, itself, generally among debris on the ground. Parasites so far identified are the Diptera, Thecocarcelia incedens and T. latifrons, both ex pupa. The species is multi-brooded.”

Henning, Henning, Joannou & Woodhall, 1997: 113 [as Spialia diomus ferax; photograph of final instar larva].

Fianl instar larva and pupa of Spialia ferax. Images courtesy Allison Sharp.

Larval food: Hermannia comosa Burch. ex DC. (Malvaceae) [Murray, 1959; South Africa; as camosa]. Hermannia cuneifolia Jacq. var. cuneifolia (Malvaceae) [Murray, 1959; South Africa; as Hermannia pallius Eckl. and Zeyh.]. Hermannia depressa N.E.Br. (Malvaceae) [Woodhall, 2005; South Africa]. 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: 189]. Pavonia burchellii (DC.) R.A.Dyer (syn. patens Andr.) (Malvaceae) [Murray, 1959; South Africa; as Pavonia macrophylla E. Mey. ex Harv.]. Sida cordifolia L. (Malvaceae) [Staude in CRG database, 2016; Magaliesburg, Gauteng]. Sida dregei Burtt Davy (Malvaceae) [Staude in CRG database, 2016; Magaliesburg, Gauteng].

abscondita Plötz, 1884 (as sp. of Pyrgus). Mitteilungen aus dem Naturwissenschaftlichen Vereins von Neu-Vorpommen und Rügen in (Greifswald) Berlin 1884: 21 (1-24). “Africa”.

lacreuzei Oberthür, 1912 (as sp. of Syrichthus). Étude de Lépidoptérologie Comparée 6: 117 (47-120). South Africa: “Zoutpansberg, Nord-Transvaal”.

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