Sisyrnodytes major Adams, 1905

Londt, Jason G. H., 2009, A review of Afrotropical Sisyrnodytes Loew, 1856 (Diptera: Asilidae: Stenopogoninae), African Invertebrates 50 (1), pp. 137-137 : 157-160

Sisyrnodytes major Adams, 1905

Figs 3, 4 , 23–26 , 40

Sisyrnodytes major: Adams 1905: 155 ; Oldroyd 1957: 86; 1974: 73 (fig. 66 entire ơ); 1980: 368 (catalogue). Sisyrnodytes niger Bezzi, 1906: 283 .

Sisyrnodytes disjunctus Séguy, 1931: 654 .

Sisyrnodytes erebus Oldroyd, 1957: 85 ; 1980: 368 (catalogue). Syn. n.

As this species displays some geographical variation as well as sexual dimorphism I provide redescriptions of the types of both major (^) and erebus (ơ) as they adequately represent much of the observed diversity.

Redescriptions:

Male (based on holotype of erebus in fair condition; both antennae broken off beyond pedicel, right pro- and mesothoracic legs broken off beyond trochanters and missing, left metathoracic leg lacking terminal four tarsomeres; the specimen is double mounted on a cellulose strip).

Head: Dark red-brown to black with black, white and yellowish setae. Antennae: Dark red-brown (terminal parts missing). Scape and pedicel black setose ventrally, yellowish setose dorsally. Major ventral setae of pedicel missing. Eye to face width ratio 1.9:1. Mystax short, mixed black and yellowish ventrally, white dorsally. Frons and vertex pale yellow setose. Laterally situated frontal setae not extending below antennal sockets. Occipital setae black, postoculars pale yellow except for group of c. 5 black setae behind ocellar tubercle. Palps and proboscis black setose.

Thorax: Dark red-brown to black. Mesonotum entirely pale yellow and orange setose except for narrow paramedial strips, sutural and postsutural spots. Lateral macrosetae pale yellow, other setae mixed white and pale yellowish. Scutellum apruinose with moderately defined transverse groove; disc asetose, apical setae numerous pale yellowish, arranged in about three rows. Pleural setae dark red-brown to black and pale yellowish. Anepisternum with dark red-brown setae anterodorsally and pale yellowish setae posterodorsally. Katepimeron weakly dark red-brown to black setose, katatergals long, shafts more or less smooth, mainly black setose except for some yellowish setae dorsally. Legs: Dark red-brown, mainly black setose (there are some small white and yellowish setae on pro- and mesothoracic legs). Claws longish, dark red-brown; empodia short yellowish (most broken), pulvilli minute to absent. Wings 5.3× 2.6 mm. C white setose basally; membrane pale brownish stained except for narrow hind margin beyond fusion of CuA

2

and A

1

. Haltere red-brown.

Abdomen: Dark red-brown; black setose except for posterolateral parts of T1–6 which are mostly white setose.

Genitalia: Not dissected, but visible structures appear to conform to dissected genitalia of males from Nguruma ( Kenya) ( Figs 23, 24 ) and Zuarungu ( Ghana) ( Figs 25, 26 ). Epandrium greatly reduced, simple in structure, proctiger extending well beyond it. Gonocoxite with external lobe tapering to narrowly-rounded tip, internal lobe with deeply forked tip. Gonostylus shorter than internal lobe of gonocoxite, fairly robust with hooked tip. Hypandrium large, flattish, triangular in ventral view, with broad medial lobe extending almost as far as tip of internal lobe of gonocoxite and obscuring a view of other organs in ventral view. Aedeagus fairly robust, slightly curved in lateral view, with blunt tip.

Female (based on lectolotype of major in good condition, slightly dusty).

Head: Dark red-brown, white, pale yellow and dark red-brown setose. Antennae: Red-brown except for tip of style which is pale yellowish. Scape and pedicel mostly pale yellowish setose (a few small white setae present). Major ventral setae of pedicel project beyond level achieved by postpedicel. Basal element of style short (slightly longer than broad). Eye to face width ratio 2.3:1. Mystax mainly pale yellowish with group of white setae dorsally. Frons and vertex white, pale yellow setose. Laterally situated frontal setae not extending below antennal sockets. Occipital setae white, postoculars mixed white and pale yellow. Palps dark red-brown setose, proboscis white setose.

Thorax: Dark red-brown. Mesonotum entirely setose except for narrow paramedial strips, sutural and postsutural spots. Lateral macrosetae light brown; other setae mostly mixed white, pale yellowish and dark red-brown. Scutellum apruinose with poorly defined transverse groove; disc asetose, apical setae numerous, difficult to count (c. 40), red-brown, yellowish and white. Pleural setae mostly white except for some pale yellowish setae dorsally on anepisternum and katatergite. Katepimeron white setose, katatergals white, pale yellowish, long, shafts more or less smooth. Legs: Orange-brown, pro- and mesothoracic femora dark red-brown, mainly white and pale yellow setose, but there are black setae terminally on tibiae and ventrally on tarsi. Claws longish, dark red-brown; empodia short yellowish; pulvilli minute to absent. Wings 6.2× 2.7 mm; C white setose basally; R 4 with obvious remains of supernumerary crossvein (spurvein) basally; membrane pale brownish stained except for distal third of wing. Haltere with dark red-brown knob, orange-brown stalk.

Abdomen: Dark red-brown, blackish and white setose; blackish setae small, confined to central parts of T1–4, white setae longish, erect laterally and on sterna, recumbent along posterior margins of T1–5 (some missing).

Variation and sexual dimorphism: Size – wing length ơ 4.6–7.5 mm (– x= 5.8 mm, n=44), ^4.1 –7.9 mm (– x=6.0 mm, n=48). While on average females are slightly bigger than males the range in size is fairly significant in both sexes. This is also true for a local population (measurable erebus type specimens from Azare) where wing lengths are as follows – ơ 5.8–6.7 mm (x –= 6.1 mm, n=7), ^5.3 –7.3 mm (– x= 6.2 mm, n=14), although ranges were smaller. A fairly uniform species at any one locality, but displays sexual dimorphism in setal coloration (i.e. mystax – ơ black and white (proportions vary), ^pale yellow; pleura – ơ black and white, ^pale yellow; legs – ơ mostly black setose, ^mostly pale yellow setose; and the degree of wing staining, ơ darker stained, ^lighter stained (staining mostly confined to vein margins). Although there is variation within any given population, geographical variation is evidenced in the degree of wing staining, especially in ơ (i.e. West Africa : The Gambia, Ghana – extensively strongly stained except for tip and hind margin ( Fig. 4 ); Benin – basal half of wing strongly stained including entire costal cell. East Africa : Eritrea – basal half of wing strongly stained including entire costal cell; Kenya, Tanzania – strongly stained basally ( Fig. 3 ) including half of costal cell (or c. 2/3 of wing weakly stained). Southern Africa: Mozambique – strongly stained basally up to humeral crossvein (entire costal cell unstained)). Male terminalia show remarkably little variation over this extensive species range .

Type specimens examined: ZIMBABWE: ^lectotype 2^paralectotypes ( major ) ‘Salisbury [Harare, 17°50'S: 31°03'E] / S. Africa / F. L. Snow’, ‘ Sept 1900 / 5050ft’, ‘Cotype / Sisyrnodytes / major / Adams’ (SEMC). MOZAMBIQUE: 1^holotype ( disjunctus ) ‘ct Juillet’, ‘Museum Paris / Mozambique / Vallée de Pompoué [Rio Pompué, 16°53'S: 34°40'E] / P. Lesne 1924’, ‘Type’ [red on cream], ‘ Sisyrnodytes / disjunctus / Type ơ / E. Séguy det.1930’ (MNHN). NIGERIA:1 ơ holotype ( erebus ) ‘Type’ [circular with red rim], ‘Shaku. [09°33'N: 06°10'E] / 14.12.10.’, ‘N. Nigeria. / J.J. Simpson. / 1912–460.’, ‘ Holotype / Sisyrnodytes / erebus Oldroyd / det. J.E. Chainey. 1983’ [this rectangular label has a circular red rimmed label glued to its upper right hand corner reading ‘Holo- / type’] (BMNH); 5 ơ 11^paratypes ( erebus ) ‘Nigeria: / Azare. [10°41'N: 10°12'E] / Dr. Ll. Lloyd.’ (BMNH); 1^paratype ( erebus ) ‘Nigeria: / Azare. / 1924 / Dr. Ll. Lloyd.’ (BMNH); 2 ơ 1^1? paratypes ( erebus ), ‘Nigeria: / Azare. / 1928-1929 / Dr. Ll. Lloyd.’ (BMNH). THE GAMBIA: 1^paratype ( erebus ) ‘Salikeni [Salikene, 13°21'N: 14°01'W] / Gambia / 8.iii.1911 / J.J. Simpson’ (BMNH); 1^paratype ( erebus ) ‘Kerewan [13°22'N: 14°13'W] / Gambia / 6.iii.1911 / J.J. Simpson’ (BMNH).

Lectotype designation: Adams (1905) states that he saw three specimens and mentions both the male and female sex. However, all three specimens are female. He did not designate a holotype, and as the designation of a lectotype would provide taxonomic stability for this variable species, I hereby designate the only ‘cotype’ that shows no obvious damage as the lectotype. The other two females, both with visible damage (missing leg or detached wing) I consider to be paralectotypes. The lectotype is the smallest of the three specimens having wing measurements of 6.2× 2.7 mm; the paralectotypes measure 7.1×3.0 and 8.0× 3.6 mm respectively.

Notes on synonymised species:

Sisyrnodytes niger Bezzi, 1906 : I have not seen the type material. Bezzi (1906) provides the following information for his Eritrean type specimen(s) ‘ Anseba, Halibaret, Tellini.’ Although Oldroyd (1957) lists niger as a synonym of major , and states that the holotype is ‘in Udine, Italy’ he may not have studied the material personally as he does not list Bezzi’s locality for major . Although I have not verified the whereabouts of Bezzi’s material, I have seen 3 ơ from Eritrea that carry old identification labels giving the name niger . These conform to my concept of major and so I accept this fairly long-standing synonymy.

Sisyrnodytes disjunctus Séguy, 1931 : The holotype is in excellent condition. Although labelled as a male it is a female as stated by Séguy (1931). I can confirm that disjunctus is indeed a synonym of major .

Sisyrnodytes erebus Oldroyd, 1957 : The synonymy of erebus with major is supported by evidence gained during a study of the variation seen in all the material available to me. In addition male terminalia appear to be remarkably consistent over the extensive range of the species (as can be seen when comparing illustrations of males from Kenya ( Figs 23, 24 ) and Ghana ( Figs 25, 26 )). The fact that there is no male in the major type series does create some doubt concerning this synonymy, but that can only be dispelled when male specimens are available from the type-locality.

Other specimens examined: BENIN: 1 ơ [?] ‘Bas Dahomey 1906 / D’Gallard [? somewhat illegible]’ (MCMI). ERITREA: 3 ơ ‘Sammlung / F. Hermann’, ‘ Sisyrnodytes / niger Bezzi’ [old faded yellowish label], ‘Eritrea / Sisyrnodytes / niger Bezzi’ [pink label found on only one of the specimens], ‘ Sisyrnodytes / niger / Bezzi

/ ơ / det. E. O. Engel’ (ZSMC); 1 ơ ‘Eritrea / Aswara-Keren [Keren, 15°47'N: 38°28'E] / Rd. 2.12.45 / K. Guichard’ (BMNH); 1^‘ Eritrea / Mt Rumio [?] / 7/2/54’ (BMNH). THE GAMBIA: 3 ơ 7^‘ Keneba [13°20'N: 16°01'W], Gambia / 19.xi.74 [1 ơ 1^] 25.xi.74 [1 ơ] 26.xi.74 [1^] 11.xii.74 [1^] 27.xii.74 [1 ơ 3^] 27.ii.75 [1^] / Grassland in burnt bush’, ‘W.F. Snow collection / pres. W.F. Snow, 1996’ (OXUM). GHANA: 1 ơ 2^‘ Gold Coast / Zuarungu [10°47'N: 00°48'W] / 15.xi.1952 / J. Bowden’ (NMSA); 1 ơ ‘Gold Coast / Zuarungu / 22.xi.1952 / J. Bowden’ (NMSA). KENYA: 1 ơ ‘Lake Rudolf. [L. Turcana, 03°30'N: 36°00'E] / Sept. 1944 / Mrs J. Adamson’ (BMNH); 1 ơ 1^‘ Samburu Dist. [01°15'N: 37°00'E] / June, 1944, / E. Opiko.’ (BMNH); 1^‘ Kabarnet Dist / Baringo [00°28'N: 35°58'E], 1–44 [i.1944] / Museum Staff.’ (BMNH); 1^‘ O.U.E.C.Exp. / Mt. Kenya. / B.M. 1949-562’, ‘E. Africa: / Kijabe [Research Station, 00°55'S: 36°35'E] Plateau. / Southern extreme. / 6,800 ft. 17.vii.1949. / J.A. Riley’ (BMNH); 2 ơ 2^‘ Kenya #61 / Nguruma, Kajiado dist / 01°50'S: 36°56'E 700m / vi.1990, Rift valley / Coll: I.M.I. Abu-Zinid / Alluvial plains’ (NMSA); 1 ơ 3^‘ Brit. E.Africa. / Near Wangi [Wange, 02°00'S: 40°55'E], / coast of mainland. / 21– 22 Feb. 1912. / S.A. Neave’ (BMNH); 1 ơ ‘Kenya / near Kitui [? Kituti, 02°08'S: 38°07'E] / 9.ix.74 / D.J. Greathead (BMNH); 4 ơ 1^‘ Brit. E.Africa. / Witu [02°23'S: 40°26'E] / 28.ii.12. / S.A. Neave’ (BMNH); 2 ơ ‘Kenya / Mtito / Andei [02°41'S: 38°10'E] / 22.v.1963 / D.J. Greathead’ (BMNH); 1 ơ 1^‘ Kenya / Taveta [03°24'S: 37°41'E] / 9.vi.1962 / D.J. Greathead’ (BMNH); 1^‘ Kenya / Masai [? Masai Manyatta, 03°43'S: 37°51'E] near / Orgisaille / 23.vi.74 / D.J. Greathead/ (BMNH). MAURITANIA: 1 ơ ‘N. Africa. / Mauritania; betw. Kiffa [16°37'N: 11°24'W] & Tidjidja / 1931 / Mrs Mary Steele’ (BMNH). NIGER: 1 ơ ‘Niger: / Nr. Zinder. [13°48'N: 08°59'E] / 2.xii.1960. / G. Popov. / B.M. 1966-359’ (BMNH). SOMALIA: 1 ơ 1^‘ Somaliland: / Gemhisem [somewhat illegible,? Gemadin, 10°18'N: 47°09'E] / 25/ii/53 / Desert Locust Survey’ (BMNH). SOUTH AFRICA: 1^‘ N. E. Zoutp.[Soutpansberg, 22°45'S: 30°00'E] / dist.7&8.16 [vii & viii.1916] / H.G. Breyer’ (NMSA); 1 ơ ‘Maputa [26°59'S: 32°45'E] / June ’ [19]14 / H.G. Breyer’ (NMSA). SUDAN: 1 ơ 1^‘ Sudan / Darfur, Teiga [15°38'N: 25°40'E] / 27.x.1958 / G.B. Popov’ (BMNH). TANZANIA: 2 ơ ‘Tanganyika / Old Shinyanga. [03°33'S: 33°24'E] / 13.vii.55 / E. Burtt’ (BMNH); 1 ơ 1^‘ Tanganyika / Shinyanga Dist. / Old Shinyanga / Dr E. Burtt. / on / burnt ground / 26.vii.56’ (BMNH); 3 ơ 3^‘ Tanganyika / Shinyanga Dist. / Old Shinyanga / Dr E. Burtt. / 27.vii.56’ (BMNH); 1 ơ ‘Tanganyika / Old Shinyanga / on unburnt / ground / E. Burtt. / 31.vii.56’ (BMNH); 1 ơ 1^‘ Tanganyika / Arusha Chini [03°35'S: 37°20'E] / 7.vi.1962 / D.J. Greathead’ (BMNH); 1 ơ ‘Tanzania: / Same, [04°04'S: 37°44'E] Rt. B1 / 8–16.ix.1992 / A. Friedberg’ (NMSA); 2 ơ ‘T. Nash 6/9/1928 / Tanganika Territory / 489 [1 ơ] 490 [1 ơ] / Kikorie [? Kikorue, 07°18'S: 35°33'E]’ (OXUM); 1 ơ 1^‘ Njombe [09°20'S: 34°46'E] / 6000–6500 ft. / Tanganyika / 8.8.1951 W.P.’ (BMNH).UNKNOWN [? Sudan]: 1^‘ Anglo-Egyp.Sud./ H.H. King’ (BMNH). Distribution, phenology and biology: Oldroyd (1957) says of major : ‘One of the most widely distributed species, occurring from Eritrea: Asmara-Keren Road (Guichard), through Kenya: Witu and Wangi Dt. (Neave), Tanganyika: Shinyanga District (Burtt) to Southern Rhodesia: Salisbury (type of major ) and Moçambique: Pompué Valley (same latitude as Salisbury) (type of disjunctus )’. For erebus he recorded ‘ Holotype ơ Northern Nigeria: Shaku (Simpson). Paratypes: Azare (Lloyd), 5 ơ 15^. Gambia: Kerewan, 1^; Sankeni, 1^(Simpson)’. With the synonymy of erebus the distribution of major is greatly extended into West Africa ( Fig. 40 ). According to Oldroyd (1957) major is active in the adult phase during ‘December–February in north; July in south.’ Specimens listed in this paper were collected in every month of the year except April (Table 1). The species appears to be mainly winter-active, bearing in mind that it is found both north and south of the equator (Table 1). Although there is very little label data that provides information about habitat preferences, Oldroyd (1957) reports that ‘Dr. Burtt took it on burnt ground, where its charred, black-and-white appearance would give good concealment’. This association with burnt vegetation is supported by label data on some Gambian specimens (see above).

Similar species: This species is similar to apicalis , aterrimus and vestitus .

Figures

Figs 1–5. Sisyrnodytes species: (1) S. brevis (Macquart, 1838), dorsal view of a Gonnemanskraal ơ; (2) S. curtus (Wiedemann, 1819), lateral view of Knersvlakte ^; (3, 4) S. major Adams, 1905, variation in wing staining: (3) Nguruma ơ; (4) Azare paratype ơ; (5) S. aethes sp. n., wing venation of Meiringspoort paratype ^.

Figs 19–24. Sisyrnodytes species, ơ terminalia: (19, 20) S. dasykylon sp. n., 23 km SE of Middelpos paratype, lateral and ventral; (21, 22) S. irwini Oldroyd, 1974, holotype, lateral (shape of robust aedeagus shown with a broken line) and ventral; (23, 24) S. major Adams, 1905, Nguruma, Kenya, lateral and ventral. Scale lines = 1 mm.

Figs 25–30. Sisyrnodytes species, ơ terminalia in lateral and ventral aspects: (25, 26) S. major Adams, 1905, Zuarungu, Ghana; (27, 28) S. nilicola (Rondani, 1850), Tel Aviv, Israel; (29, 30) S. niveipilosus Ricardo, 1925, Bulawayo, Zimbabwe. Arrowed is a broken structure. Scale lines = 1 mm.

Fig. 40. Distribution of Sisyrnodytes aterrimus Engel, 1929 (•), major Adams, 1905 (·) and nilicola (Rondani, 1850) (9).