14th edition (2015).

Genus Swinhoe, 1912

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

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

A genus of small skippers with 20 Afrotropical species, two of which (doris and zebra) extend extralimitally. There are a further eight Palaearctic and Oriental members. The genus was monographed by De Jong in 1978. Most species are found in savanna habitats but Spialia ploetzei is found in forest. All species fly low down and are attracted to flowers, mud, cow 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 larvae have been recorded mainly on herbaceous plants in the dicotyledonous families Sterculiaceae, Malvaceae and Tiliaceae. Spialia doris is exceptional in that its larvae have been found utilising Convolvulaceae.

*Spialia agylla (Trimen, 1889)# Grassveld Sandman

Grassveld Sandman (Spialia agylla bamptoni ). Left – male upperside. Left – male underside. Images courtesy Steve Woodhall.

Pyrgus agylla Trimen, 1889. South-African : a monograph of the extra-tropical species 3 Papilionidae and Hesperidae: 286 (438 pp.). London. Spialia agylla Trimen. Swanepoel, 1953a. Spialia agylla (Trimen, 1889). Dickson & Kroon, 1978. Spialia agylla (Trimen and Bowker, 1889). Pringle et al., 1994: 318.

1 Spialia agylla agylla. Male (Wingspan 21 mm). Left – upperside; right – underside. Noupoort, Eastern Cape Province, . 8 December 2005. J. Dobson. Images M.C. Williams ex Dobson Collection.

Spialia agylla agylla. Female (Wingspan 23 mm). Left – upperside; right – underside. Noupoort, Eastern Cape Province, South Africa. 8 December 2005. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [South Africa]: “Cape Colony; Eastern Districts, between Somerset East and Murraysberg, Murraysberg, Hope Town. Griqualand West; Kimberley, between Riet and Modder Rivers, Vaal River”. Holotype in the Natural History Museum, London. Original description: “Exp. al., 10-11.5 lin. Closely allied to P. Asterodia, Trim., but smaller, with the markings of a purer white, and the discal ones larger; hind-wing with basi-cellular spot distinct and well developed, and submarginal spots larger. Under side: Hind-wing and apex of fore-wing darker in ground-colour, less strongly tinged with ochreous-yellow; nervular interruptions of cilia conspicuously black, without admixture of ochreous yellow. Hind-wing: sub-basal and median white bands much more irregular and denticulate on both edges, and more sharply defined by darker fuscous edging in parts; - the opposite projections of these bands all but touch each other in two places, viz., at origin of subcostal and of median nervules respectively; submarginal white spots enlarged, and near anal angle sometimes partly confluent.” Diagnosis: Distinguished on the underside of the hindwing by the very irregular band, which at vein 2 almost reaches the basal spots. On the underside of the wings the cilia are conspicuously chequered (Pringle et al., 1994). Distribution: Botswana, South Africa, Lesotho. Habitat: The nominate subspecies is found in grassland and, to a lesser extent, in grassy areas in the Karoo and Kalahari. Subspecies bamptoni occurs in grassy areas of the succulent karoo in Little Namaqualand (Pringle et al., 1994). Habits: The flight is fast and low down, with frequent rests on the ground or low bushes (Pringle et al., 1994). Flowers are readily visited. Males do not hilltop but do defend territories from perches on the ground or on stones. Flight period: August to April for the nominate subspecies and from September to December for subspecies bamptoni (Pringle et al., 1994). Early stages:

Clark, in Dickson & Kroon, 1978: 188 [as Spialia agylla bamptoni; Port Nolloth, Northern Cape].

Bampton, in Pringle et al., 1994: 318 [as Spialia agylla bamptoni]. Leaves are partially bound together with silk by the larvae and pupation takes place in the partly open shelter thus formed.

Henning, Henning, Joannou & Woodhall, 1997: 101 (photograph of final instar larva and pupa).

Larval food: comosa Burch. ex DC. (Sterculiaceae) [Kroon, 1999]. Hermannia species (Sterculiaceae) [Dickson & Kroon, 1978: 188; for Spialia agylla agylla]. Hermannia species (Sterculiaceae) [Dickson & Kroon, 1978: 188; for Spialia agylla bamptoni]. burchellii DC. R.A.Dyer (Malvaceae) [Henning, Henning, Joannou & Woodhall, 1997: 100].

2 Spialia agylla agylla (Trimen, 1889)#

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

Spialia agylla agylla. Male (Wingspan 21 mm). Left – upperside; right – underside. Noupoort, Eastern Cape Province, South Africa. 8 December 2005. J. Dobson. Images M.C. Williams ex Dobson Collection.

Spialia agylla agylla. Female (Wingspan 23 mm). Left – upperside; right – underside. Noupoort, Eastern Cape Province, South Africa. 8 December 2005. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: [South Africa]: “Cape Colony; Eastern Districts, between Somerset East and Murraysberg, Murraysberg, Hope Town. Giqualand West; Kimberley, between Riet and Modder Rivers, Vaal River”. Holotype in the Natural History Museum, London. Distribution: Botswana, South Africa (, North West Province, , Free State Province, Eastern Cape Province, Western Cape Province, Northern Cape Province), Lesotho (west). Specific localities: Botswana – Ghanzi (De Jong, 1978). Mpumalanga – Standerton (Swanepoel, 1953). Gauteng – Johannesburg (Swanepoel, 1953); Heidelberg (Swanepoel, 1953). Free State Province – Trompsburg (Swanepoel, 1953); Ladybrand (Swanepoel, 1953); Bloemfontein (Swanepoel, 1953); Boshof (Swanepoel, 1953); Kroonstad (Swanepoel, 1953); Frankfort (Swanepoel, 1953). Eastern Cape Province – Somerset East (Trimen, 1889); Port Elizabeth (Swanepoel, 1953); Cradock (Swanepoel, 1953); Klaarstroom (Swanepoel, 1953); Aberdeen (Swanepoel, 1953); Steynsburg (Swanepoel, 1953); Burgersdorp (Swanepoel, 1953). Western Cape Province – Murraysburg (Trimen, 1889). Northern Cape Province – Hopetown (Trimen, 1889); Kimberley (Trimen, 1889); Noupoort (Swanepoel, 1953); Hantamsberg (Pringle et al., 1994). Lesotho – Maseru (Swanepoel, 1953); Leribe (Swanepoel, 1953).

Spialia agylla bamptoni Vári, 1976#

3 Spialia agylla bamptoni Vári, 1976. Annals of the Transvaal Museum 30: 123 (121-144). Spialia agylla bamptoni Vári, 1976. Dickson & Kroon, 1978. Spialia agylla bamptoni Vári, 1976. Pringle et al., 1994: 318.

Spialia agylla bamptoni. Female. Left – upperside; right – underside. Hondeklip Bay, Northern Cape Province, South Africa. 4 October 1995. J. Greyling. Images M.C. Williams ex Greyling Collection.

Type locality: South Africa: “Hondeklipbaai, Namaqualand Distr.”. Holotype (male) in the Transvaal Museum, Pretoria. Diagnosis: Distinguished from the nominate subspecies by the much more prominent white spotting, particularly on the hindwing underside (Pringle et al., 1994). Distribution: South Africa (Western Cape Province – north, Northern Cape Province – west). Specific localities: Western Cape Province – Brand se Baai (Williams, unpub.). Northern Cape Province – Hondeklip Bay (TL; Bampton); Port Nolloth (Kaplan, vide Dickson & Kroon, 1978).

*Spialia asterodia (Trimen, 1864)# Star Sandman

Male of the Star Sandman (Spialia asterodia). Bulwer Mountain, KwaZulu-Natal Image courtesy Steve Woodhall

Pyrgus asterodia Trimen, 1864. Transactions of the Entomological Society of London (3) 2: 178 (175-180). Pyrgus asterodia Trimen. Trimen, 1866a. Pyrgus asterodia Trimen, 1864. Trimen & Bowker, 1889. Spialia asterodia Trimen. Swanepoel, 1953a. Spialia asterodia (Trimen, 1864). Dickson & Kroon, 1978. Spialia asterodia (Trimen, 1864). Pringle et al., 1994: 318.

4

Spialia asterodia. Male (Wingspan 24 mm). Left – upperside; right – underside. Hillshaven, Gauteng Province, South Africa. 20 September 2008. J. Dobson. Images M.C. Williams ex Dobson Collection.

Spialia asterodia. Female (Wingspan 27 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]: “Cape Colony”. Holotype in the Natural History Museum, London. Diagnosis: Median band on underside of hindwing irregularly edged and parallel to the series of basal spots (Pringle et al., 1994). Distribution: Mozambique (south), Zimbabwe, 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: Limpopo Province – Haenertsburg (Swanepoel, 1953); Munnik (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Zoutpansberg (Swanepoel, 1953); Percy Fyfe Nature Reserve (Warren, 1990); Highlands Wilderness (Bode & Bode, unpublished checklist); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015). Mpumalanga – Barberton (Swanepoel, 1953); Ermelo (Swanepoel, 1953); Lydenburg (Swanepoel, 1953); Graskop (Swanepoel, 1953). North West Province – Potchefstroom (Swanepoel, 1953). Gauteng – Johannesburg (Swanepoel, 1953); Pretoria (Swanepoel, 1953). Free State Province – Rouxville (Swanepoel, 1953); Ladybrand (Swanepoel, 1953); Bethlehem (Swanepoel, 1953); Bloemfontein (Swanepoel, 1953); Kroonstad (Swanepoel, 1953). KwaZulu-Natal – Loteni (Swanepoel, 1953); Balgowan (Swanepoel, 1953); Estcourt (Swanepoel, 1953); Ladysmith (Swanepoel, 1953); Vryheid (Swanepoel, 1953). Eastern Cape Province – Port Elizabeth (Swanepoel, 1953); Grahamstown (Swanepoel, 1953); Cathcart (Swanepoel, 1953); Dordrecht (Swanepoel, 1953); Burgersdorp (Swanepoel, 1953); Steynsburg (Swanepoel, 1953). Western Cape Province – Plettenberg Bay (Trimen); Cape Town (Swanepoel, 1953); Swellendam (Swanepoel, 1953); Robinson Pass (Van den Berg); Hermanus (Rossouw); Durbanville (Robertson). Lesotho – Maseru (Swanepoel, 1953); Mokhotlong (Swanepoel, 1953); Giant’s Castle (Swanepoel, 1953). Habitat: Grassland and, to a lesser extent, grassy areas in the fynbos, karoo and bushveld. In the Drakensberg it may be found at altitudes up to 3 000 m (Pringle et al., 1994). Habits: Very fond of flowers but seldom mud-puddles. Males establish territories in grassy areas and perch on the ground. Very occasionally they exhibit hilltopping behaviour. Flight period: August to March (Pringle et al., 1994). Early stages: 5

Clark, in Dickson & Kroon, 1978: 230; plate 13 [as Spialia asterodia; recorded from eggs and larvae from Port Elizabeth, Eastern Cape Province]. “Egg: 6 mm diameter by 0,5 mm high; laid singly among young leaf buds. Pale blue-green with fine white ribbing consisting of 18 to 20 longitudinal ribs, only 8 or 9 of which reach the micropyle. These are crossed by 20 to 23 fine ribs, in the form of reversed diagonal braces in the spaces between the other ribs, with the junctions punctuated by white moles. Eggs hatch after some 11 days. The discarded shell is not eaten. Larva: 1st instar 1,25 to 2,75 mm in 18 days; 2 nd instar 2,75 to 4 mm in 12 days; 3 rd instar 4 to 5,5 mm in 12 days; 4th instar 5,5 to 9 mm in 13 days; 5th instar 9 to 16 or 17 mm in 21 days. The duration of instars is variable. Like those of most Spialia the young larva starts feeding on the inner surface of a very young, unfolding leaf; later, it constructs a purse-like ‘hide-out’ by joining leaves together. With increased growth larger shelters are constructed, the last generally near the ground. Moulting and pupation take place within these shelters. There are about two to three broods, the last having long drawn-out instars, and with hibernation during the pupal period, which sometimes occurs among dead leaves joined together near the roots of the plant. Pupa: 13 mm. Secured by cremastral hooks to silk on the inner side of the leaves forming a shelter. Emergence takes place after 20 days. Parasites: Parasitic attack occurs in the larval state, by species of Tachinidae. The larva turns a pale dirty yellow, and the maggot emerges out of its side. In some cases the latter delays its emergence until its host has pupated. Some 36 hours after emergence the maggot pupates and after a further 12 to 25 days its pupa yields the mature parasite.”

Larval food: Hermannia comosa Burch. ex DC. (Sterculiaceae) [Kroon, 1999]. Hermannia diffusa L.f. (Sterculiaceae) [Dickson & Kroon, 1978: 188; as Hermannia pilosula Harv.]. Hermannia incana Cav. (Sterculiaceae) [Dickson & Kroon, 1978: 188; as Hermannia candicans Ait.]. species (Malvaceae) [Dickson & Kroon, 1978: 188]. Pavonia burchellii (DC.) R.A.Dyer (Malvaceae) [Henning, Henning, Joannou & Woodhall, 1997: 99].

*Spialia colotes (Druce, 1875)# Bushveld Sandman

Males of the Bushveld Sandman (Spialia 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] Spialia colotes (Druce, 1875). Pringle et al., 1994: 318.

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, South Africa, Swaziland, Saudi Arabia, Yemen, Oman. Habitat: Dry and moist savanna (bushveld). 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

6 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. 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: Nothing published.

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

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

Spialia 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]

Type locality: Angola. Distribution: Angola. Specific localities: Angola – Loanga (Plötz, 1884).

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

Spialia colotes semiconfluens de Jong, 1978

Spialia colotes semiconfluens de Jong, 1978. Tijdschrift voor Entomologie 121: 58 (23-146).

Type locality: Somalia: “Bihendula”. Distribution: Ethiopia, Somalia, Uganda (north-east), Kenya (north-east), Saudi Arabia, Yemen, Oman. Specific localities: Somalia – Bihendula (TL). Kenya – Tsavo area (Larsen, 1991c).

Spialia colotes transvaaliae (Trimen, 1889)#

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.

7

Spialia 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.

Spialia 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 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, Mozambique, Zimbabwe, Botswana, Namibia, South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng), 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). 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 8 (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).

*Spialia confusa (Higgins, 1924)# Confusing Sandman

Hesperia transvaaliae var. confusa Higgins, 1924. Transactions of the Entomological Society of London 1924: 90 (64-120). Spialia confusa (Higgins, 1924). 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, 1924). Dickson & Kroon, 1978. Spialia confusa (Higgins, 1925). Pringle et al., 1994: 319. [date of authorship erroneous]

Spialia 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.

Spialia 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: 9 species (Sterculiaceae) [Van Someren, 1974: 325; Sevastopulo, vide Pringle et al., 1994: 319; for subspecies obscura]. species (Tiliaceae) [Sevastopulo, vide Pringle et al., 1994: 319; for subspecies obscura].

Spialia confusa confusa (Higgins, 1924)#

Hesperia transvaaliae var. confusa Higgins, 1924. Transactions of the Entomological Society of London 1924: 90 (64-120). Spialia confusa (Higgins, 1924). 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]

Spialia 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.

Spialia 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, 1924 (as var. of Hesperia transvaaliae). Transactions of the Entomological Society of

10 London 1924: 90 (64-120). [Malawi]: “Nyasaland, British East Africa”.

Spialia confusa obscura (Higgins, 1924)

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

Type locality: [Kenya]: “British East Africa, Nairobi”. Distribution: Kenya (east and central), 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 Spialia colotes (Larsen, 1991c: 405). Tanzania – Dar es Salaam (Kielland, 1990d); Pugu Hills (Kielland, 1990d); Dendene Forest at Kisiju (Kielland, 1990d); foot of the Nguru Mountains (Kielland, 1990d).

*Spialia delagoae (Trimen, 1898)# Delagoa Sandman

Males of the Delagoa Sandman (Spialia 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.

Spialia 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.

11

Spialia 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, Kenya, 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). Mozambique – Delagoa Bay (TL). Botswana – Lobatse (Pennington; male specimen illustrated above); 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].

*Spialia 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.

Type locality: [Tanzania]: “Msamwia”.

12 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 Spialia 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 (Sterculiaceae) [M.J.W. Cock, vide Larsen, 1991c: 407].

Spialia depauperata depauperata (Strand, 1911)

Hesperia ferax var. depauperata Strand, 1911. Mitteilungen aus dem Zoologischen Museum in Berlin 5: 304 (275-304).

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”.

Spialia depauperata australis de Jong, 1978#

Spialia abscondita Plötz. Swanepoel, 1953a. [Misidentification of Spialia 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.

13 Spialia 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.

Spialia 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).

*Spialia diomus (Hopffer, 1855)# Common Sandman

Males of the Common Sandman, Spialia diomus. Images courtesy Allison Sharp.

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 subspecies ferax] Spialia diomus Hopffer. Swanepoel, 1953a. [referable to subspecies ferax] Spialia diomus (Hopffer, 1855). Pringle et al., 1994: 319.

14

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

Type locality: Mozambique: “Mossambique”. [See note below]. Diagnosis: The light band on the hindwing underside is broad and curved (Pringle et al., 1994). Distribution: Sub-Saharan Africa, including Senegal, Gambia, Guinea, Liberia, Ivory Coast, Ghana, Nigeria, Uganda, Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, Botswana, Namibia, South Africa, Swaziland, Lesotho, Yemen. Habitat: All habitats except forest. Predominantly in savanna. In Tanzania the nominate subspecies is found at altitudes from near sea-level to 2 000 m and subspecies ferax at 2 200 m (Kielland, 1990d). Habits: A common and widespread Afrotropical sandman (Larsen, 1991c). Flies rapidly, close to the ground. Both sexes are readily attracted to flowers (Pringle et al., 1994). 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). 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). 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 [photograph of final instar larva].

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

Larval food: Hermannia comosa Burch. ex DC. (Sterculiaceae) [Dickson & Kroon, 1978: 189]. 15 Hermannia cuneifolia Jacq. var. cuneifolia (Sterculiaceae) [Dickson & Kroon, 1978: 189; as Hermannia pallius Eckl. and Zeyh.]. Hermannia depressa N.E.Br. (Sterculiaceae) [Woodhall, 2005]. Hermannia diffusa L.f. (Sterculiaceae) [Dickson & Kroon, 1978: 189; as Hermannia pilosula Harv.]. Hermannia incana Cav. (Sterculiaceae) [Dickson & Kroon, 1978: 189; as Hermannia candicans Ait.]. Hibiscus aethiopicus L. (Malvaceae) [Dickson & Kroon, 1978: 189]. Pavonia burchellii (DC.) R.A.Dyer (syn. patens Andr.) (Malvaceae) [Dickson & Kroon, 1978: 189; as Pavonia macrophylla E. Mey. ex Harv.]. species (Malvaceae) [Larsen, 1991c: 404]. Triumfetta brachyceras K.Schum. (Tiliaceae) [Van Someren, 1974: 325; as Triumfetta macrophylla Schum.]. Jacq. (Tiliaceae) [Larsen, 2005a]. species (Sterculiaceae) [Larsen, 1991c: 404].

Note: If the type locality is correct (Mozambique) the nominate subspecies is the one with a southern African distribution and ferax (Wallengren, 1863) (TL [Namibia]) is a synonym. The northern populations should then be Spialia diomus machacoana (Butler, 1899) (TL Kenya) (MCW).

Spialia diomus diomus (Hopffer, 1855)

Pyrgus diomus Hopffer, 1855. Berichte über die zur Bekanntmachung geeigneten Verhandlungen der Königl. Preuss. Akademie der Wissenschaften zu Berlin 1855: 643 (639-643).

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

Type locality: Mozambique: “Mossambique”. Distribution: Senegal, Gambia, Guinea, Liberia, Ivory Coast, Ghana, Nigeria, Uganda, Kenya, Tanzania, Yemen. Recorded, in error for subspecies ferax, from Mozambique by Congdon et al., 2010. Specific localities: Ghana – Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Kenya – Machako’s (Butler, 1899). Tanzania – Central, north and east (Kielland, 1990d); Ngara District (Kielland, 1990d); Katavi National Park (Fitzherbert et al., 2006).

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

Spialia diomus ferax (Wallengren, 1863)#

Syrichtus [sic] ferax Wallengren, 1863. Wiener Entomologische Monatschrift 7: 137 (137-151. Pyrgus diomus Hopffer, 1855. Trimen & Bowker, 1889. [referable to subspecies ferax] Spialia diomus Hopffer. Swanepoel, 1953a. [referable to subspecies ferax] Spialia diomus ferax (Wallengren, 1863). Dickson & Kroon, 1978. 16 Spialia diomus ferax (Wallengren, 1863). Pringle et al., 1994: 319.

Spialia diomus 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.

Spialia diomus 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). Distribution: 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. 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). 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). 17 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).

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”.

*Spialia doris (Walker, 1870) Desert Grizzled

Nisoniades doris Walker, 1870. Entomologist 5: 56 (48-57).

Type locality: [Djibouti]: “Tajora”. Distribution: Mauritania, Sudan, Kenya, Somalia, Djibouti, Saudi Arabia, Yemen, Oman. Extralimitally in Morocco, Egypt, Arabia, Iran and Pakistan. 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 species (Convolvulaceae) [Larsen, 1991c: 405]. Corchorus species (Tiliaceae) [Larsen, 1991c: 405]. Ipomoea species (Convolvulaceae) [Larsen, 2005a].

Spialia doris doris (Walker, 1870)

Nisoniades doris Walker, 1870. Entomologist 5: 56 (48-57).

Type locality: [Djibouti]: “Tajora”. Distribution: Sudan, Kenya (north), Somalia, Djibouti, Saudi Arabia, Yemen, Oman. Extralimital in Arabia, southern Iran and Pakistan. Specific localities: Djibouti – Tajora (TL). Yemen – Aden (Butler, 1885).

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

Spialia doris daphne Evans, 1949

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

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

18

*Spialia dromus (Plötz, 1884)# Forest Sandman

Upper- and underside of the Forest Sandman ( Spialia 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.

Spialia 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.

Spialia 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 19 al., 1994). Distribution: Sub-Saharan Africa, including Senegal, Gambia, Sierra Leone (north), Burkina Faso, Guinea (north), Guinea-Bissau (Mendes et al., 2007), Ivory Coast (north), Ghana (north), Nigeria, Gabon (de Jong, 1978); Democratic Republic of Congo, Uganda, Kenya, Tanzania, Zambia, Mozambique, Zimbabwe, Botswana, South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, KwaZulu-Natal, Eastern Cape Province), Swaziland. Specific localities: Ghana – Accra Plains (Larsen, 2005a); Badagry Creek beaches near Lagos (Larsen, 2005a); Boabeng- Fiema Monkey Sanctuary (Larsen et al., 2009). Democratic Republic of Congo – Mpala (Oberthür, 1912). Tanzania – Throughout (Kielland, 1990d); Kitanga (Oberthür, 1912); Zanzibar (Higgins, 1924); 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). 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

20 shelter or among rubbish on the ground, and pupates after some 24 to 36 hours. The parasitic fly emerges from the pupa after 12 to 25 days.”

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

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

Larval food: Melhania species (Sterculiaceae) [Van Someren, 1974: 325]. Triumfetta rhomboidea Jacq. (Tiliaceae) [Dickson & Kroon, 1978: 189; Vuattoux, 1999 (Ivory Coast)]. Triumfetta tomentosa Bojer (Tiliaceae) [Dickson & Kroon, 1978: 189]. Waltheria indica L. (Sterculiaceae) [Van Someren, 1974: 325; as Waltheria americana].

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, 1924 (as var. of Hesperia dromus). Transactions of the Entomological Society of London 1924: 95 (64-120). No locality given.

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

*Spialia kituina (Karsch, 1896) Kitui Grizzled Skipper

Hesperia kituina Karsch, 1896. Entomologische Nachrichten. Berlin 22: 374 (372-378).

21

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

Type locality: [Kenya]: “Sansibargebiete, von Kitui”. Distribution: Kenya (north-west, central, east). Specific localities: Kenya – Kitui (TL); Maunga Inkubwa (Butler, 1898); Lukenia Hills (Larsen, 1991c); Umkambani country (Larsen, 1991c); Kibwezi (Larsen, 1991c); Lokichogio (Larsen, 1991c). Habitat: Dense savanna (Larsen, 1991c). Habits: A widespread but uncommon Kenyan endemic (Larsen, 1991c). Early stages: Nothing published. Larval food: Sida species (Malvaceae) [Van Someren, 1974: 325].

bettoni Butler, 1898 (as sp. of Pyrgus). Proceedings of the Zoological Society of London 1898: 415 (395-444). Kenya: “Maunga Inkubwa”.

*Spialia mafa (Trimen, 1870)# Mafa Sandman

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.

22

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 Spialia spio, 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. 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 23 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: Hermannia comosa Burch. ex DC. (Sterculiaceae) [Kroon, 1999]. Hermannia depressa N.E.Br. (Sterculiaceae) [Henning, G., in Pringle et al., 1994: 320]. Hermannia species (Sterculiaceae) [Dickson & Kroon, 1978: 189]. Hibiscus aethiopicus L. (Malvaceae) [Dickson & Kroon, 1978: 189]. Hibiscus ludwigii Eckl. & Zeyh. (Malvaceae) [Van Someren, 1974: 325; as Hibiscus macrantha]. Pavonia burchellii (DC.) R.A.Dyer (Malvaceae) [Heath et al., 2002: 8]. Pavonia species (Malvaceae) [Larsen, 1991c: 405]. Sida ovata Forssk. (Malvaceae) [Van Someren, 1974: 325; as Sida grewioides]. Sida rhombifolia L. (Malvaceae) [Van Someren, 1974: 325]. 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)].

Spialia mafa mafa (Trimen, 1870)#

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

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. 24 Specific localities: 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). 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 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.).

Type locality: [Kenya]: “Mumias, 4,000 ft”. Distribution: Sudan (south), Ethiopia, Somalia, Kenya, 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 mangana (Rebel, 1899) Arabian Grizzled Skipper

Hesperia (Pyrgus) mangana Rebel, 1899. Anzeiger der Akademie der Wissenschaften. Wien. 36: 360 (359-361).

25

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

Type locality: [Yemen]: “Râs Fártak (Arab. m.)”. Distribution: Yemen, Oman, Somalia, Ethiopia, Uganda, Kenya (north). Specific localities: Yemen – Ras Fartak (Larsen, 1991c). Oman – north (Feulner, 2007). 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). Early stages: Nothing published. Larval food: Nothing published.

Relevant literature: Feulner, 2007 [Recorded from northern Oman].

*Spialia nanus (Trimen, 1889)# Dwarf Sandman

Dwarf Sandman (Spialia 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.

26

Spialia 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.

Spialia 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.

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). 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

27 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].

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

Larval food: Hermannia comosa Burch. ex DC. (Sterculiaceae) [Dickson & Kroon, 1978: 187]. Hermannia cuneifolia Jacq. var. cuneifolia (Sterculiaceae) [Dickson & Kroon, 1978: 187; as Hermannia pollens Eckl. and Zehy.]. Hermannia diffusa L.f. (Sterculiaceae) [Dickson & Kroon, 1978: 187; as Hermannia pilosula Harv.]. Hermannia incana Cav. (Sterculiaceae) [Dickson & Kroon, 1978: 187; as Hermannia candicans Ait.]. Hermannia pulverata Andrews (Sterculiaceae) [Dickson & Kroon, 1978: 187]. Hibiscus aethiopicus L. (Malvaceae) [Dickson & Kroon, 1978: 187].

*Spialia paula (Higgins, 1924)# Mite Sandman

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

28

Spialia 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.

*Spialia ploetzi (Aurivillius, 1891) Forest Grizzled Skipper

Hesperia ploetzi Aurivillius, 1891. Entomologisk Tidskrift 12: 227 (193-228).

Spialia 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.

29 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 Spialia the discal band runs towards the hind margin or anal angle (Congdon & Collins, 1998). Distribution: Guinea-Bissau, 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:

Larsen, 1981.

Larval food: Triumfetta species (Tiliaceae) [Larsen, 1981 (Lagos, Nigeria); Vuattoux, 1999 (Ivory Coast)].

Spialia ploetzi ploetzi (Aurivillius, 1891)

Hesperia ploetzi Aurivillius, 1891. Entomologisk Tidskrift 12: 227 (193-228).

Spialia 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.

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: Gabon – Tchimbele (van de Weghe, 2010); Ovan (van de Weghe, 2010); Nouna (van de Weghe, 2010). 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).

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, 1924 (as [unnecessary] replacement name for Hesperia zebra Rebel). Transactions of the 30 Entomological Society of London 1924: 101 (64-120).

Spialia ploetzi occidentalis de Jong, 1977

Spialia ploetzi occidentalis de Jong, 1977. Entomologische Berichten 37: 43 (41-44).

Spialia ploetzi occidentalis. Male. Left – upperside; right – underside. Ghana. Images courtesy Torben Larsen.

Type locality: Ghana. Distribution: Guinea-Bissau, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon (west). Specific localities: Sierra Leone – Loma Mountains (Larsen, 2005a). Ghana – Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Cameroon – Korup (Larsen, 2005a).

*Spialia 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. Spialia sataspes (Trimen, 1864). Dickson & Kroon, 1978. Spialia sataspes (Trimen, 1864). Pringle et al., 1994: 317.

Spialia 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.

Type locality: [South Africa]: “Cape Colony and British Kaffraria”. Holotype in the Natural History Museum, London. Diagnosis: Closely resembles Spialia delagoae, from which it may be distinguished on the hindwing 31 underside by the absence of submarginal spots (Pringle et al., 1994). Can also be confused with Spialia 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). Specific localities: Eastern Cape Province – Addo (Swanepoel, 1953); Coega (Swanepoel, 1953); Grahamstown district (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 species (Sterculiaceae) [Dickson & Kroon, 1978: 187]. Hibiscus aethiopicus L. (Malvaceae) [Dickson & Kroon, 1978: 187]. Pavonia burchellii (DC.) R.A.Dyer (Malvaceae) [Henning, Henning, Joannou & Woodhall, 1997: 96].

*Spialia 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.

32

Spialia 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.

Spialia 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). 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 (male illustrated above). 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 Spialia secessus.]

*Spialia spio (Linnaeus, 1764)# Mountain Sandman

33

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 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, Guinea,

34 Mali, Sierra Leone, Liberia, Ivory Coast, Burkina Faso, Ghana, Togo, Benin, Nigeria, Niger, Gabon (de Jong, 1978), Uganda, Kenya, Tanzania, 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: Benin – Lokoli (Tchibozo et al., 2008). Tanzania – Throughout (Kielland, 1990d); Mount Longido (Kielland, 1990d). 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). 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 skipper (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). 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). 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

35 the previous species [Spialia diomus 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].

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

Larval food: Althaea species (Malvaceae) [Kroon, 1999]. Hermannia coccocarpa (Eckl. & Zeyh.) Kuntze (Sterculiaceae) [Pringle et al., 1994: 319]. Hermannia comosa Burch. ex DC. (Sterculiaceae) [Henning, Henning, Joannou & Woodhall, 1997: 115]. Hermannia cuneifolia Jacq. (Sterculiaceae) [Dickson & Kroon, 1978: 189; as Hermannia pallius Eckl. and Zeyh.]. Hermannia diffusa L.f. (Sterculiaceae) [Dickson & Kroon, 1978: 189; as Hermannia pilosula Harv.]. Hermannia incana Cav. (Sterculiaceae) [Dickson & Kroon, 1978: 189; as Hermannia candicans Ait.]. Hibiscus aethiopicus L. (Malvaceae) [Dickson & Kroon, 1978: 189]. Hibiscus fuscus Garke (Malvaceae) [Heath et al., 2002: 8]. Hibiscus pusillus Thunb. (Malvaceae) [Dickson & Kroon, 1978: 189; as Hibiscus gossypinus sensu Harv. non Thunb. = Hermannia cus L.]. Hibiscus pusillus Thunb. (Malvaceae) [Van Someren, 1974: 325; as Hibiscus gossyphina]. Lavatera arborea L. (Malvaceae) (exotic) [Henning, Henning, Joannou & Woodhall, 1997: 115]. Lavatera species (Malvaceae) [Larsen, 1991c: 404]. Pavonia burchellii (DC.) R.A.Dyer (syn. patens Andr.) (Malvaceae) [Dickson & Kroon, 1978: 189; as Pavonia macrophylla E. Mey. ex Harv.]. Pavonia columella Cav. (Malvaceae) [Kielland, 1990d: 236]. Sida cordifolia L. (Malvaceae) [Van Someren, 1974: 325]. Sida ovata Forssk. (Malvaceae) [Van Someren, 1974: 325; as Sida grewioides]. Sida schimperiana Hochst. ex A.Rich. (Malvaceae) [Van Someren, 1974: 325]. 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)]. Triumfetta species (Tiliaceae) [Larsen, 1991c: 404]. Urena lobata L. (Malvaceae) [Larsen, 2005a]. Wissadula rostrata (Schumach.) Hook.f. (Malvaceae) [Larsen, 2005a; as Wissadula amplissima].

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

36 Africa]: “Kaap de Goede Hoop”.

*Spialia wrefordi Evans, 1951 Wreford’s Grizzled Skipper

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

Spialia wrefordi. Male. Left – upperside; right – underside. Ghana. 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.

*Spialia zebra (Butler, 1888) Zebra Grizzled Skipper

Pyrgus zebra Butler, 1888. Annals and Magazine of Natural History (6) 1: 207 (132-151, 196-209).

Type locality: [India] Pakistan: “Campbellpore; Futch Khan’s bungalow, Kooteer, Chittar Pahar, 2000 to 3000 feet”. Distribution: Yemen, Oman, Ethiopia, Somalia, Sudan, Uganda, Kenya, Tanzania. 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: Nothing published. Larval food: Dombeya species (Sterculiaceae) [Larsen, 1991c: 407]. Melhania ovata (Cv.) Spreng. (Sterculiaceae) [Van Someren, 1974: 325]. Melhania velutina Forssk. (Sterculiaceae) [Van Someren, 1974: 325].

Spialia zebra bifida (Higgins, 1924)

37 Hesperia bifida Higgins, 1924. Transactions of the Entomological Society of London 1924: 111 (64-120).

Type locality: [Kenya]: “Nairobi, Mombasa, British East Africa”; Uganda. Distribution: Yemen, Oman, Ethiopia, Somalia, Sudan (south), Uganda (east), Kenya, Tanzania (north). Specific localities: Kenya – Nairobi (TL); Widespread (Larsen, 1991c); Mombasa (Higgins, 1924). 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).

38