A Review of Afrotropical Acnephalum Macquart, 1838, Including the Reinstatement of Sporadothrix Hermann, 1907 and Descriptions of Two New Genera (Diptera: Asilidae: Stenopogoninae)

ABSTRACT In revising the taxonomy of Acnephalum Macquart, 1838 it was discovered that the Afrotropical species are not congeneric when compared with the Palaearctic type species (A. olivierii Macquart, 1838 from Greece), which is congeneric with species of Pycnopogon Loew, 1847. As a consequence all Palaearctic Pycnopogon species are transferred to Acnephalum which takes precedence over Pycnopogon. In addition it was found that Sporadothrix Hermann, 1907, a genus previously synonymised with Acnephalum, is worthy of full generic reinstatement. The new genus Acnephalomyia is provided for Afrotropical species previously assigned to Acnephalum. The following taxonomic adjustments, innovations and decisions are introduced in this paper: New combinations: Species transferred from Pycnopogon — Acnephalum apicalis (Matsumura, 1916), Acnephalum apiformis (Macquart, 1849), Acnephalum denudatus (Séguy, 1949), Acnephalum fasciculatus (Loew, 1847), Acnephalum hirsutus (Becker, 1913), Acnephalum laniger (Dufour, 1833), Acnephalum leucostomus (Engel, 1939), Acnephalum melanostomus (Loew, 1874), Acnephalum mixtus (Loew, 1847), Acnephalum nikkoensis (Matsumura, 1916), Acnephalum pallidipennis (Brullé, 1936). Species transferred from Acnephalum — Acnephalomyia andrenoides (Wiedemann, 1828); Acnephalomyia dorsalis (Macquart, 1838); Acnephalomyia platygaster (Loew, 1858); Afroholopogon futilis (Wulp, 1899). New synonyms: Dasypogon quadratus Wiedemann, 1828, Acnephalum punctipennis Macquart, 1855, Acnephalum cockerelli Curran, 1934 and Sisyrnodytes sericeus Oldroyd, 1974 = Acnephalomyia andrenoides (Wiedemann, 1828); Acnephalum cylindricum Oldroyd, 1974 = Sporadothrix gracilis Hermann, 1907. New genera: Acnephalomyia (Type species: Dasypogon andrenoides Wiedemann, 1828), Astiptomyia (Type species: Astiptomyia bikos sp. n.). New species: Acnephalomyia eremia (Namibia), Acnephalomyia iota (Namibia, South Africa), Acnephalomyia leukoros (South Africa), Acnephalomyia probolos (South Africa), Ammodaimon platythrix (Namibia), Astiptomyia bikos (Namibia). Lectotype designations: Dasypogon andrenoides Wiedemann, 1828; Acnephalum dorsale Macquart, 1838; Acnephalum futile Wulp, 1899. Keys, tables and illustrations are provided for Afrotropical taxa as identification aids while distribution maps are included where appropriate.

The single Australasian species, A. punctipennis, has, since its description by Macquart in 1855, been largely ignored by taxonomists. Apart from being listed by Kertesz (1909) in a catalogue and listed by Hull (1962) there has been no mention of the species in the literature, including the Australasian Diptera Catalogue (Daniels 1989), since its description. The type, preserved in the Oxford University Museum of Natural History, clearly represents a South African species.

Specimens
Material used in this study is housed in the institutions listed below. The curators that kindly assisted me are named in brackets following the name of the respective institution.

Label data
In recognition of the value of detailed lists of material examined (Dikow et al. 2009), standard formats have been employed when recording label information. When material is considered to have special interest (i.e. type specimens, or when material is not abundant), all label data are reproduced as appearing on labels. For multiple labels each label is demarcated by the use of single inverted commas while each line of data is separated by a spaced slash ( / ). Data that appear on the reverse side of a label are preceded by a '~' symbol. In some instances the colour of a label is provided in square brackets. Square brackets are also used when useful additional information, or comment, not found on labels, is provided. In this regard, coordinates are usually given when these, or a quarter-degree grid reference, do not appear on a label. Coordinates provided are usually for the populated place or geographic feature mentioned immediately before the added note, no attempt being made to estimate coordinates for places recorded as being a certain distance from a populated place. The use of question marks usually indicates unknown or questionable information. When material is abundant, the kind of detail mentioned above is not considered necessary.
Information for these specimens is restricted to locality, date of collection, collector(s) (initials excluded), altitude (when available) and any other potentially useful information available. Should more detail be required, it is available from the relevant institution. All specimens are arranged in geographical order according to latitude and within alphabetically ordered countries to facilitate mapping.

Descriptive passages
Brief generic diagnoses are usually provided. If fuller descriptions are required, those of Hull (1962) or other researchers mentioned in the text may be consulted. Species and size, and so information relating to these characters should not be given undue value.
It should be noted that in order to adequately view and study the normally retracted male genitalia, these need to be excised, softened through maceration in dilute KOH and physically extruded. Extrusion can be achieved by inserting a pin into the opening created after the terminalia have been excised. Genitalia are effectively exposed by carefully dragging the terminalia over the inserted pin. Unfortunately, this method of extrusion may cause some soft, macerated sclerites and associated membranes to buckle slightly. However, most of the organs providing good diagnostic characters (e.g. the hypandrium and gonocoxites) are usually not too adversely affected.
As Afrotropical species currently assigned to Acnephalum are not congeneric it further necessary to provide a new genus for them (see below).
Evidence in support of my actions is embodied in the following redescription and notes relating to the A. olivierii holotype.
Designation of type species of Acnephalum: The designator of A. olivierii as type species has long been accepted to be Engel (1929) who, without explanation, merely states 'Genotype: Olivieri Macq.'. Röder (1882), however, although misspelling the generic name, clearly stated, under the heading 'Ueber Acnecephalum Macq.', that 'Die typische Art der Gattung ist Acnecephalum Olivieri Macq. (Diptères exotiques Tome premier 2. partie pag. 51)'. I believe this statement complies with requirements for a properly constituted type species designation, and that this was probably known to Engel as he makes reference to Röder's work. Diagnosis: Stenopogonine asilids with the following combination of characters. Head: Antennal postpedicel elongate, slender; style composed of 3 elements (2 slender segments and terminal spine-like seta); head clearly wider than high in anterior view (not more-or-less circular); face slightly convex; mystax covering entire face; vertex distinctly excavated; angle of divergence of frons/vertex in anterior view >20°; palpi 2-segmented; proboscis straight. Thorax: Anatergites bare; metepisternal macrosetae present; postmetacoxal area membranous; pulvilli well developed; wing with cells r 1 and m 3 open at margin; costal vein extending around entire wing margin (although weakly around anal cell and alula); 'stump-vein' at base of R 4 absent. Abdomen: Segdensely setose laterally. Acnephalum olivierii Macquart, 1838Figs 6, 12, 13, 19 Macquart's (1838 description is brief and here replicated in full as a matter of convenience. olivierii. In cataloguing the species, Lehr (1988) reported it from Greece (presumably the type material) with queries for the then USSR and North Africa. Remarks: Although the rather badly preserved type specimen strongly resembles Afrotropical species currently assigned to Acnephalum, it demonstrates features that clearly separate it from these species and align it with Pycnopogon. Features that I believe distinguish it (and other specimens of Pycnopogon above) from Afrotropical species covered by this review (i.e. those assigned to the new genus -see below) are shown in Table 1. Although a modern revision of Acnephalum is required before the value of these characters will be known for certain, I believe they serve well to support the taxonomic decisions made here.
As mentioned earlier, with the discovery that the type species of Acnephalum (A. olivierii, a Palaearctic species) is not congeneric with Afrotropical species, it is necessary to describe a new genus for these species. As these species have had a long history of being called Acnephalum it was decided to retain at least part of the name in providing a new one, hence the choice of Acnephalomyia.
During this study it was further decided to re-establish Sporadothrix as a valid genus, and so species assigned to that taxon are handled separately below. In addition, it was discovered that Acnephalum futile is actually a species of Afroholopogon Londt, 1994, and so that species is also handled separately under this name. With these adjustments there were actually only six valid Afrotropical species requiring revision at the commencement of this study (i.e. andrenoides, cockerelli, dorsale, platygaster, quadratus, sericeus). Diagnosis: Stenopogonine asilids with the following combination of characters. Head: Antennal postpedicel elongate, style composed of 3 elements (2 slender segments and terminal spine-like seta); head clearly wider than high in anterior view (not more or less circular); face slightly convex; mystax long, covering entire face; vertex distinctly excavated; angle of divergence of frons/vertex in anterior view < 20°; palpi 2-segmented, well-developed; proboscis straight. Thorax: Dorsocentrals undifferentiated; anatergites bare; metepisternal macrosetae absent; postmetacoxal area membranous; pulvilli present, but poorly developed (c. one-third length of claws); wing with cell m 3 open at margin; costal vein extends around wing margin, terminating at A 1 (i.e. anal cell and alula without bordering vein); stump-vein at base of R 4 commonly present, even if rudimentary. Abdomen Notes on synonymies: The following four species are considered nominal taxa.

Dasypogon quadratus Wiedemann, 1828
Wiedemann's material ( Fig. 1), in good condition, was studied and compared with the andrenoides lectotype. The lectotype is almost identical, and only the slight differences recorded below were noted. There is no doubt that quadratus is a synonym of andrenoides (as suspected by Oldroyd in 1974).
Head: Antenna: Segmental ratios 1.0:0.9:3.1:0.2:1.4. Ocellar setae mostly yellowish. Thorax: Pronotal macrosetae brownish yellow. Mesonotal setae mostly mixed white and brown-yellow except for some uniformly coloured whitish clumps anteriorly. Scutellum with c. 20 yellowish apical macrosetae. Wing: 7.6×3.5 mm. Membrane fairly extensively orange-brown stained. Only central parts of cells bordering wing margin distally unstained. Vein R 4 with short basal stump-vein. Abdomen: Terga fringed laterally with pale yellow setae extending along distal margins of terga for a short distance only. Genitalia dissected and studied (not illustrated as they are similar to those in Figs 24, 25).
Variation: Paralectotypes similar, but one is somewhat smaller than the other specimens, having a wing length of 5.1 mm.

(ZMHB).
Notes: Wiedemann did not designate a holotype so his specimens are considered syntypes. For reasons of taxonomic stability I here recognise and designate a lectotype and two paralectotypes. There are quadratus types in NHMW which are (like the specimens mentioned above under andrenoides exactly the same 'Cap b. sp' label, in ZSMC. As in the case of the andrenoides material mentioned above, I have no means of knowing whether these were available to Wiedemann, and so I prefer to consider them as having no type status. I have also seen the following specimen, assumed to be from Dasypogon / quadratus Wied / CGH. Dr Klug' (OXUM). The specimen carries a note reading 'This could be a / syntype, exchanged / with Mus. Berlin / A. C. Pont det. / 1998'. As confirmation of this suggestion cannot be provided, I also consider this specimen to have no type status.
Remark: Although D. andrenoides and D. quadratus were described in the same pubquadratus with andrenoides as the latter

Acnephalum punctipennis Macquart, 1855
Macquart's holotype, in good condition (antennae broken off beyond pedicels; scutellar macrosetae damaged), was studied and compared with the andrenoides lectotype. Despite being a female it is very similar, and only the differences recorded below were noted. There is no doubt that punctipennis is a synonym of andrenoides.
Head: Antenna: Scape and pedicel uniform red-brown. Thorax: Entirely dark redbrown. Scutellar disc with a few white setae laterally. Legs: Mostly orange-brown. Wing: 8.4×3.6 mm. Membrane mostly transparent, but with some basal staining that extends over costal region. All vein junctions are orange stained giving wings a spotted appearance. Abdomen: Generally strongly white setose, bordering fringe not well-developed (as is normal for females). of two pieces of paper glued together, one upon the other. Probably originally placed below the specimen in a drawer], 'Type Dip.: 155 / Acnephalum / punctipennis / Macquart / Hope Dept. Oxford' (OXUM). Notes: Macquart (1855) gives specimen information as 'De l'Océanie, cap des Aiguilles. M. Bigot.' which somehow established the belief that the species was Australian. There is little doubt that the specimen comes from South Africa as it closely resembles other specimens from the region. While the precise provenance remains a mystery, the locality information appearing on the large label cited above, which is poorly hand this species was entirely and inexplicably, although correctly, overlooked during the preparation of the Diptera catalogue of the Australasian and Oceanian regions.  Curran's holotype, in excellent condition, was studied and compared with the andrenoides lectotype. It is almost identical, and only the slight differences recorded below were noted. There is no doubt that cockerelli is a synonym of andrenoides.
'Acnephalum / cockerelli Note: Although not stated on a label,  says that the holotype was collected by T.D.A. Cockerell.
Sisyrnodytes sericeus Oldroyd, 1974 In reviewing the Afrotropical Sisyrnodytes species, Londt (2009) transferred sericeus to Acnephalum. During the present study it became evident that Oldroyd's species is synonymous with andrenoides. Oldroyd's holotype, in excellent condition, was studied and compared with the andrenoides lectotype. Although smaller, it is almost identical, and only the slight differences recorded below were noted. Variation and sexual dimorphism: A. andrenoides is a highly variable species that also displays some sexual dimorphism. There is, for example, a considerable range in size, and, although all specimens were not measured, males ranged in wing length between 4.3-10.4 mm while females had a similar range of between 4.3-10.3 mm. Both sexes show variation in setal and wing membrane coloration. While most specimens are predominantly white setose, specimens may range in colour to the extent that some are predominantly black setose. Males frequently have well developed abdominal fringes (long laterally situated tergal setae), but there are examples with relatively poorly developed fringes. Although the wings of males are generally more darkly stained than those of females, the extent of staining is variable in both sexes. Male wing membranes may range from being entirely dark brown stained to being pale brown stained proximally and largely unstained distally. Females never have entirely brown-stained wings, but resemble males with predominantly unstained wings. Type locality: The type specimens of A. andrenoides are merely labelled 'Capland Krebs S.' and it is not known exactly where Krebs collected them. Now that the distribution of the species is well understood I designate an area 18 km N Sutherland (32°15'S:20°43'E) as type locality. This area is where the road between Sutherland and Williston crosses the Renoster River. This place is selected for two reasons -specimens similar to the types have been collected there, and the locality is central with respect to the known range. Distribution, phenology (Table 3) and biology: The species is widely distributed (Fig.  65), ranging from southern Namibia, down the west coast of South Africa (Northern and Western Cape Provinces) and eastwards through to the Eastern Cape Province. There are a few records that appear improbable (e.g. Rustenburg in North West Province and Pietermaritzburg in KwaZulu-Natal!) and these specimens may have been mislabelled. In addition there are a few localities from which only female specimens are known.
andrenoides and platygaster means that some distribution points provided for andrenoides could actually belong to platygaster (and vice versa). The species is active in the adult stage during late winter, spring and summer (July-February) although the majority of collections were made between September and December. Some specimens have been pinned together with prey items. The 20 records available to me are as follows (sex of asilid bracketed): Coleoptera: Hodotermitidae dominate the sample, it is probable that andrenoides, like many other asilids, is a generalist. Similar species: A. andrenoides can be confused with platygaster and, in a few instances, large specimens of dorsalis -especially when female specimens are involved. Although male hypandria are largely withdrawn, it is usually possible to separate the males of these species using the appearance of the distal parts of this organ.
Acnephalomyia dorsalis (Macquart, 1838)  Redescription (Based on lectotype. Condition: Good; dirty and covered with some fungal strands. Anterior pleurites of left side somewhat damaged (probably by a dermestid) and the right mesothoracic leg stuck into position with glue. The specimen is double mounted with tip of abdomen touching card.): Head: Dark red-brown to black, white setose, silver pruinose (evident mainly on frons). Antenna: Scape and pedicel brown-orange, postpedicel and style dark red-brown. Segsubequal in length, white setose; setae of pedicel as long or longer than postpedicel; postpedicel twice as long as scape and pedicel combined; style 2-segmented, tipped with spine, subequal in length to scape and pedicel combined. Face dark red-brown dark red-brown to black, long white setose; angle subtended by eye margins at level of frons/vertex c. 12°. Proboscis orange-brown to dark red-brown, white setose, slightly downturned distally. Palpus 2-segmented, white setose.  (OXUM). Note: The paralectotype is in poor condition: left prothoracic leg missing terminal four tarsomeres; abdomen and posterior part of thorax, including metathoracic legs, entirely missing (apparently consumed by dermestids); wings intact, but tip of left wing slightly damaged. The remaining parts suggest that the specimen was similar to the lectotype. Notes: Macquart indicated that his description was based on a female. Of the two types, one is a male and the other lacks an abdomen. These carry modern syntype labels, and as it is reasonable to believe that Macquart saw both specimens, probably believing them to be female (many mistakes have been made in the intact male as lectotype and the other specimen as paralectotype. Although the specimens do not carry locality data, Macquart records the following for his material 'Du Cap. Collection de M. Serville' and a large drawer label accompanies the material which reads '185 A. Dorsale. / C. B. Speil. [Cap Bone Spei = Cape of Good Hope] / (Coll. Serville) Macq. D. Eu.'. Although Oldroyd (1974) states 'Type in Paris', and so probably never consulted the types, he was able to correctly identify some specimens as dorsale, probably using the rather unreliable pattern of white abdominal setae as a key character. Distribution, phenology (Table 3) and biology: The species is found mainly in the western parts of South Africa (Fig. 66) with records coming from the Western Cape and Northern Cape provinces. Adults are active during spring and summer (August-February) although most collections were made in October and November.
Similar species: Although the male genitalia are similar to those of eremia, leukoros, probolos and platygaster, in that the hypandria are elongate, males can be relatively pale yellowish basal segments of the antennae usually associated with dorsalis is a fairly reliable character.
Figs 38-40, 66 Etymology: From Greek eremia (desert, wilderness); noun in apposition. Refers to the arid conditions under which the species has been found. Description (Condition of holotype fair; antennae broken off beyond pedicels; right prothoracic and left meso-and metathoracic legs missing.): Head: Dark red-brown to black, white setose, mostly apruinose. Antenna (broken off beyond pedicel): Scape and pedicel brown-orange. Segmental ratios 1.0:0.8:?:?:?scape and pedicel subequal in length, pale whitish setose (most macrosetae missing, but single macroseta on dorsal aspect of right pedicel suggests that these would be as long or longer than postpedicel). Face dark red-brown to black, apruinose except for narrow strips adjacent to eye margins, mystax white, covering entire face, but weakly dorsally (some setae appear to have been rubbed off). Frons, vertex and postocular region dark red-brown to black, apruinose except for central part of postocular region, white setose; angle subtended by eye margins at level of frons/vertex c. 17°. Proboscis red-brown, white setose. Palpus dark red-brown, 2-segmented, white setose.
Abdomen: Terga broader than long, mostly dark red-brown to black, but somewhat orange-brown laterally. Terga apruinose, but entirely pitted by setal sockets. Terga with long, recumbent white setae laterally. White setae extend along distal margins of terga gradually becoming shorter medially. Large areas of terga appear asetose, but are covered with tiny reddish-brown setae. Sterna dark red-brown, apruinose, longish white setose. Terminalia largely withdrawn between T6 and S6, distal parts somewhat obscured by setae. Genitalia : Epand moderately well-developed, jutting out to almost same level as achieved by outer lobe of goncx, distally broadly rounded and shallowly incised. Proc somewhat swollen in appearance and jutting out slightly beyond level achieved by epand (lateral view). Exterior lobe of goncx broadly rounded proximally, tapering distally to a broadly rounded, somewhat truncate, distal end; interior lobe rather long, jutting out to about same level as achieved by hypd, slightly downturned distally. Hypd tally projecting medial lobe. Distribution and phenology (Table 3): Known only from the type locality (Fig. 66) where it was collected in August.
dorsalis, leukoros, probolos and platygaster, but shape of gonostylus in lateral view, and especially the distal extremity of outer lobe, is unique as is the shape of the hypandrium.

Acnephalomyia iota sp. n.
Figs 41-43, 68 Etymology: From Greek iota (ninth letter of Greek alphabet, anything very small); noun in apposition. The name refers to the small size of the species. Description (Based on holotype. Condition: Excellent): Head: Dark red-brown to black, white setose, partly silver pruinose. Antenna: Uniformly dark red-brown, white setose. Segmental ratios 1.0:1.2:4.2:0.2:1.2 -scape slightly shorter than pedicel; macrosetae of pedicel clearly shorter than postpedicel; postpedicel almost twice as long as scape and pedicel combined; style 2-segmented and tipped with spine, clearly shorter than scape and pedicel combined. Face dark ocular region dark red-brown to black. Frons extensively silver pruinose, white setose (including ocellar tubercle); postocular setae mostly brown (some postocular setae white); angle subtended by eye margins at level of frons/vertex c. 15°. Proboscis short, dark red-brown, brown setose. Palpus 2-segmented, dark red-brown, brown setose. Thorax: Dark red-brown to black, apruinose. Pronotum mostly brown setose (few white). Mesonotum white setose. Lateral macrosetae well developed, pale translucent white (2 npl, 1 spal, 1 pal). Pleura apruinose, largely asetose except for some, long, white katatergals and dorsally situated anepisternals. Katepisternum asetose. Scutellum shiny dark red-brown apruinose with poorly developed transverse, subapical groove. Four white apical macrosetae accompanied by 2 slightly shorter white setae; disc sparsely white setose. Legs: Red-brown, fem slightly darker. Major setae erect, yellowish, minor setae sparse, recumbent, white. Ventral parts of tar and terminal end of tib short, black setose. Claws black, moderately long (shorter than tarsomere 5); empodia slender, yellow, about half length of claws; pulvilli moderately developed (about length of empodium). Haltere pale yellow, base slightly darker. Wing: 2.7×1.2 mm. Veins brown, membrane sparsely microtrichose, transparent, unstained. Vein R 4 with weakly-developed basal stump-vein. Abdomen: Dark red-brown, apruinose, mostly brown setose. Terga broader than long, entirely pitted by setal sockets. T1 with 5 pale translucent white lateral macrosetae, other terga lacking macrosetae, but with a few recumbent white setulae laterally. Sterna sparsely long, white setose. Terminalia withdrawn between T6 and S6. Genitalia : Epand reduced, less than half length of outer lobe of goncx, medially deeply incised to form two lobes. Proc somewhat swollen in appearance and jutting out well beyond level achieved by epand (lateral view). Exterior lobe of goncx broadly rounded proximally, tapering distally to a narrowly rounded tip; interior lobe longish, jutting out to about same level as achieved by hypd. Hypd broadly rounded basally, tapering mystax entirely white. Frons, ocellarium and postocular region entirely white setose. Mesonotum with silver pruinose margins. Scutellum with 6 apical macrosetae, disc asetose. Legs entirely dark red-brown. Wings 3.3×1.3 mm; R 4 lacking stump-vein. Distribution, phenology (Table 3) and biology: Known only from the three type specimens collected in Namibia and South Africa (Fig. 68). The two known localities are fairly widely separated and further collecting in the intervening area is therefore desirable. Collected in September, October and November. Found on gravel plains and sandy river beds. Similar species: A tiny black species not to be confused with any other.
Abdomen: Dark red-brown to black, mostly white setose. Terga apruinose but entirely pitted by setal sockets. Terga with mostly white (a few dark red-brown) setae laterally. White setae extend along distal margins of terga gradually becoming shorter medially; each tergum with small group of white setae anteromedially. Large areas of terga appear asetose, but are covered with tiny dark red-brown setae. Sterna dark red-brown, sparsely long, white setose. Terminalia largely withdrawn between T6 and S6, distal parts somewhat obscured by setae. Genitalia (topotypic paratype male, Figs 44-46): Epand reduced, less than half length of outer lobe of goncx, distally broadly rounded and shallowly incised. Proc somewhat straight, jutting out well beyond epand (lateral view) and to similar level attained by outer lobe of goncx. Exterior lobe of goncx dorsal view), jutting out to similar level as achieved by hypd. Hypd somewhat truncate basally, greatly constricted laterally before midlength, tapering to long, distally clavate medial lobe. Variation: Paratype males are similar to the holotype. Female paratypes range in size with wing lengths of 4.0-6.0 mm (mean 5.0 mm). Females are similar except for the following features: Antennal postpedicel and proximal half of style dark red-brown; dorsal face, entire frons and vertex (not ocellarium) and most of postocular region silver pruinose; mesonotum with clumps of brown setae; mesonotal macrosetae brownyellow; scutellum with 9-14 (mean 11) apical macrosetae ranging in colour (pale white to light brown); wing may be weakly yellow to pale brown stained basally and at crossveins. Distribution, phenology (Table 3) and biology: Known from eight localities in the Western Cape of South Africa (Fig. 67). Adults have been collected in October, November and December and are therefore summer active. Specimens were found resting on sandy ground. Distribution, phenology (Table 3) and biology: Most of the records are from southern Namibia and the Northern Cape Province of South Africa (Fig. 67). Adults have been collected in August through to and including December. Although a specimen has been insect prey and so could confuse some collectors. Similar species: A large species with male genitalia similar to dorsalis, eremia, leukoros, andprobolos. Apart from the distinctive shape of the hypandrium, many males have wings that are darkly stained proximally. Females not associated with males may be confused with those of andrenoides and so a few records may, for that reason, be misplaced.

Acnephalomyia probolos sp. n.
Figs 50, 51, 68 Etymology: From Greek probolos (any protruding or jutting object or prominence); noun in apposition. The name refers to the long, projecting hypandrium. Description (Based on holotype. Condition: Good; double mounted, right prothoracic leg missing terminal four tarsomeres, left mesothoracic leg broken off and glued to mounting strip.): Head: Dark red-brown to black, white setose, weakly silver pruinose. Antenna: Dark redbrown except for tip of style which is pale yellow. Segmental ratios 1.0:0.9:3.0:0.2:1.2, scape and pedicel subequal in length, pale whitish setose, macroseta on ventral aspect of pedicel longer than postpedicel. Face dark red-brown to black, apruinose except for narrow strips adjacent to eye margins, mystax white, covering entire face. Frons, vertex and postocular region dark red-brown to black, apruinose except for central part of postocular region, mainly white setose (some dorsal setae yellowish); angle subtended by eye margins at level of frons/vertex c. 17°. Proboscis red-brown, white setose. Palpus dark red-brown, 2-segmented, pale yellowish setose.
Abdomen: Terga broader than long, dark red-brown to black. Terga apruinose, but entirely pitted by setal sockets. Terga with longish white setae laterally. White setae extend for a short distance along distal margins of terga, becoming shorter medially. Large areas of terga appear asetose, but are covered with tiny reddish brown setae. Sterna dark red-brown, apruinose, longish white setose (wavy anteriorly). Terminalia largely withdrawn between T6 and S6, distal parts somewhat obscured by setae. Genitalia (Figs 50, 51): Epand highly reduced. Proc in lateral view longer than epand. Exterior lobe of goncx and jutting out to about same level as achieved by aedeagal tip. Gonst small. Hypd in projecting medial lobe. Hypd basodorsally with a disc-like projection that lies between gonocoxites. Variation: The female paratype agrees well with the holotype, but is slightly bigger (wing length 4.7 mm) and stump-veins are moderately well developed.
Distribution and phenology (Table 3): Known only from the type locality (Fig. 68) and collected only in September. Similar species: Male genitalia are similar to those of dorsalis, eremia, leukoros, and platygaster of females of these species is sometimes problematic as they so closely resemble each other.

Key to species of Acnephalomyia
The following key is useful only for male specimens, and, because it makes use of male genital characters, it may be necessary to excise and macerate terminalia. If this hypandrium with illustrations provided in this paper. On receipt of the type material it was immediately apparent that futilis should not have been described in Acnephalum as the specimens fail to display the diagnostic characteristics of the genus. Indeed, it is doubtful that any researcher has consulted the types since their original description. The species is fully congeneric with Afroholopogon and is therefore unhesitatingly transferred to that genus. Redescription (Format follows that of Londt (2005). Based on lectotype (Fig. 2). Condition: Excellent; right antenna broken off beyond pedicel.): Head 1.0:1.1:3.3:0.3:1.3. Face black, entirely silver pruinose, mystax white, extending to antennal sockets, with vertical asetose strip medially. Face width:head width ratio 1.0:4.4. Frons and vertex black, frons apruinose except for silver lateral areas and a central spot.
Abdomen: Terga dark red-brown, shiny apruinose except for silver areas posterolaterally (that extend medially to almost centre of tergum, but not along posterior margin), white setose. Sterna dark red-brown. silver pruinose, short white setose. Genitalia (Figs 52-54): Hypopygium unrotated; epand (in dorsal view) quite short (about as long as proc), shallowly incised to form short distolateral lobes; goncx externally distally bilobed and projecting (in lateral view) to a level beyond that reached by epand; hypd (laterally) of moderate length (longer than external lobe of goncx) and (ventrally)  Lectotype designation: Wulp described the species on 'a pair in coitu, from Aden'. As he did not designate a holotype, the specimens are considered syntypes. For reasons of taxonomic stability I hereby designate the male as lectotype and the female as paralectotype. Distribution, phenology and biology: The species is only known from the type locality. Specimens were collected in February. Nothing is known of its biology. Similar species: Wulp (1899) stated that futile was 'closely allied to Dasypogon (Acnephalum) andrenoides', a comment echoed by Engel (1929). The similarity is, however, minimal and it is surprising that Engel did not suspect a misallocation.
Afroholopogon was fully revised by Londt (2005), who gave an historical account together with redescriptions and descriptions of 17 species, most of which are southern African. Using Londt's key to species reveals that futilis keys out at the end of the key together with three other speciesdasys Londt, 2005, pardosoros Londt, 2005and uranopia Londt, 2005. Two of these are known only from South Africa (pardosoros from the Tierberg Nature Reserve, and uranopia from the Graaff-Reinet area) while dasys is known from three specimens from Eritrea, Oman, and the Yemenese island of Abd al Kuri. Besides the characters provided in the key below there are obvious differences to be observed in male genital form.
A. futilis can easily be incorporated into Londt's (2005)  Sporadothrix : Hermann : 8 [1908. Type species: Sporadothrix gracilis Hermann, 1907, by original Londt (1994: 76), after studying the holotype of S. gracilis, and noting its obvious similarity to Acnephalum cylindricum Oldroyd, 1974, synonymised Sporadothrix with Acnephalum. Now that a revision of Acnephalum (as Acnephalomyia) has been completed, I have come to a different conclusion. Not only is gracilis similar to cylindricum, cylindricum must fall as a synonym of gracilis. In addition, this species shows a number of marked differences when compared to all other species previously placed in Acnephalum and so I reverse my 1994 decision and here reinstate Sporadothrix, which, as a result of the synonymy of cylindricum, remains monotypic.

Acnephalomyia Sporadothrix
Antenna: Length of style (all elements) relative to lengths of scape and pedicel combined.
Style clearly shorter than scape and pedicel combined.
Style clearly longer than scape and pedicel combined. Head: Extent of pruinescence.
Thorax: Shape of mesonotum in dorsal view (maximum length to maximum breadth ratio).
Mesonotal length equal to or shorter than breadth (i.e. almost circular in appearance).
Mesonotum clearly longer than wide (c. 1.3 times as long as broad).
Katepisternal setae absent to poorly developed (few and short).
Katepisternal setae well developed (many and long). Legs: Development of pulvilli.
Present, but poorly developed (c. one-third the length of claws).
Absent (or so minute as to be considered absent).
Abdomen length and breadth of terga).
Abdomen by number of clearly visible terga and sterna).
Segments 1-6 well-developed and clearly visible. Segments 7-8 and terminalia much reduced, withdrawn and largely hidden from view.
Segments 1-8 welldeveloped and clearly seen. Terminalia not greatly reduced and hidden from view.
Epandrium well developed, almost completely incised medially, forming a pair of well-developed lobes.

Genitalia exposed (not withdrawn between terminal terga and sterna) (Figs 55-57):
Epand extent as external parts of goncx. Proc juts out to about same level achieved by hypd (lateral view). External lobe of goncx suboval in lateral view, interior lobe slender, slightly down-curved distally. Hypd broadly rounded basally, tapering distally to narrowly rounded apex which has a small medial lobe.
Note: As the antennae and wings of the holotype are damaged, the following measurements are those of the cylindricum holotype, which is larger than the gracilis type (distance between humeral crossvein and tip of R 1 being 7.4 mm). Antennal ratios: 1:1.2:3.5:1.1:2.2. Wing: 8.6×3.4 mm. Illustrations of an entire specimen (Fig. 3), antenna (Fig. 16), wing (Fig. 9) and tar 5 (Fig. 21) are those of other specimens. Variation: Previously known only from the two type specimens listed below, I can now list further 31 specimens. This is a remarkably consistent species showing no variation being slightly smaller than females (wing length 6.3-10.9 mm).  (Table 3) and biology: The species appears to be fairly widely distributed being recorded from four places in Namibia and eight in the Northern Cape Province of South Africa (four being within the Witsand Nature Reserve) (Fig. 68), as well as from one unknown locality in Botswana. Adults are active in summer and have been collected in December -April. Specimens have been taken from grass in dry, sandy, Acacia savannah habitats. Remarks: Oldroyd (1974: 83), who had not seen the holotype, records, for gracilis, 'Type in ? Munich. Type-locality: Kalahari Desert. Distribution. I have seen one male that I assign to this species, from S.W. AFRICA: Noachabeb, 43 km N.N.E. Grunan [Grünau], 10-12.i.1972(B. M. S Afr. Exped., 1972.' Firstly, the type is not in Munich, but in the ZMHB collection. Secondly, I have studied the Noachabeb male gracilis by Oldroyd (1974) and commented upon by Londt (1994)) and, as suspected, it has proved to be a new species of Ammodaimon Londt, 1985, which is described below. Oldroyd (1974 characterised cylindricum in a key, providing a description of fewer than forty words and no illustrations. His statement regarding the material used Gobabis, January (W. D. Haacke).' The unique holotype, is actually a female, now in the NMSA, and is labelled 80 miles S of Gobabis, in present day Namibia.
Abdomen: Cylindrical, uniformly dark red-brown to black, mostly white setose (some pale yellowish on distal terga). Discal setae long, erect, normal, translucent white; minor setae scale-like, white, recumbent. Eight segments discernable, genitalia rotated through 90°, not withdrawn between subterminal terga and sterna. Terga apruinose, virtually all setae short, white, scale-like and recumbent. Genitalia (Figs 55-57) slightly clockwise rotated (<90°): Epand lobes fused proximally; lobes falling short of distance achieved by external parts of goncx. Proc juts out to about same level achieved by hypd (lateral view). External lobe of goncx in lateral view rounded basally, interior lobe slender, projecting beyond levels reached by either proc or hypd; interior lobe fairly broad in lateral view with slightly downturned (lateral view), bifurcate (dorsal & ventral views) tip. Gonst small, normally hidden from view except in ventral view, slightly curved with two relatively well-developed ventral setae distally.  (1974). Although he correctly stated in his text that it was a male, the caption for his useful illustration gives the sex as female. Distribution (Fig. 68), phenology (Table 3) and biology: The species is known only from the holotype collected in January (A. acares being collected in September). The species probably has a similar behaviour to A. acares which rests on the ground in open, fairly arid situations (Londt 1985 Head: Dark red-brown to black, mainly white setose, some silver pruinescence on face and frons. Antenna (Fig. 18): Orange-brown except for dark brown proximal part of style. Segmental ratios 1.0:1.2:3.3:0.3:1.8 -scape and pedicel subequal in length, macrosetae of pedicel pale translucent yellowish, longer than postpedicel; postpedicel almost twice as long as scape and pedicel combined; style 2-segmented, tipped with spine, subequal in length to scape and pedicel combined. Face dark red-brown to black, mystax ventrally dense white, dorsally sparse yellowish (narrow asetose strip below antennae). Frons, vertex and postocular region dark red-brown to black, short white setose; angle subtended by eye margins at level of frons/vertex c. 12°. Proboscis orange-brown white setose. Palpus 2-segmented, brown-orange, white setose.
Abdomen: Terga broader than long, T1 mostly dark red-brown to black, T2-T6 orangebrown. Terga apruinose but entirely pitted by setal sockets. Terga with recumbent white setae laterally, these extending for a short distance along distal margins of terga. Large areas of terga appear asetose, but are covered with tiny blackish setae. Sterna brownbetween T6 and S6. Genitalia (Figs 61-63): Epand moderately well-developed, slightly longer than half length of goncx, distally somewhat truncate and hardly if at all incised medially. Proc fairly short (about half length of epand), jutting out to about level achieved by goncx (lateral view). Exterior lobe of goncx broadly rounded proximally, same level as external lobe). Hypd rather short, cup shaped, broadly rounded basally, tapering rapidly to short, pointed, upwardly directed distal end. Variation: Paratypes agree well with holotype but are slightly larger (mean wing length abdomen generally more orange. Most of the paratypes have more apical scutellar farm, 860m / 15-II-1974, ME. Irwin / vegetated, moving dunes' (NMSA). Distribution, phenology (Table 3) and biology: Known only from the type locality (Fig.  68) and collected only in February. Nothing is known of the biology of this species.

Taxonomy
While the taxonomic position of Afroholopogon appears fairly stable, the situation with respect to the other genera reviewed in this paper, and Sisyrnodytes, is somewhat controversial. As alluded to in my review of Sisyrnodytes (Londt 2009), Dikow (2009a) placed Sisyrnodytes and Acnephalum, as well as Trichoura Londt, 1994, together with two Nearctic genera (Willistonina Back, 1908and Ablautus Loew, 1866 in a new subfamily named Willistonininae. Although Dikow (2009b) has provided further evidence, derived from a DNA sequencing analysis, I remain unconvinced and so refrain from adopting Dikow's arrangement until he is able to extend his analysis to include a greater number of the species covered in this and my previous work on Sisyrnodytes. As previously stated (Londt 2009), Dikow used cylindricum to represent Acnephalum. This study shows this species to be distinctive and meritorious of its own generic position (i.e. Sporadothrix), and I would venture to suggest that Sporadothrix has more in common with Ammodaimon than it does with Acnephalum, Acnephalomyia or Sisyrnodytes. At present, I believe Trichoura to be a rather distinctive genus, easily separated from Acnephalomyia and Sisyrnodytes, as well as the other genera reviewed in this paper. So, while I have the greatest admiration for Dikow's work, I prefer to await further developments before adopting his subfamily Willistonininae.
In retaining the Stenopogoninae (sensu inherent in my key to the genera (Londt 1999). After separating out a group of some seven taxa possessing setose anatergites (to this number Ontomyia Dikow & Londt, 2000 must now be added -as was done in a revised key to these genera published by Dikow & Londt in 2000), the 1999 key then divides the remaining genera into two groups on the basis of costal vein characteristics -those with a complete costa, and those having a costa that fails to border the anal cell and alula. The latter group, with reduced costal veins, was then split into two groups using the extent of development of the pulvilli -those with normally developed pulvilli and those with reduced or no pulvilli. It was into the group with reduced pulvilli that Acnephalum, Ammodaimon and Sisyrnodytes fell. It is now necessary to update the 1999 key through the incorporation of Sporadothrix and Astiptomyia. The following key picks up from couplet 9 of the Key to genera of Stenopogoninae without, or with poorly developed, pulvilli 1 Abdomen broad and dorsoventrally compressed (width : length ratio of T2 >2)..

Distribution
All the genera covered in this review, except for Afroholopogon, are southern African endemics that are largely restricted to rather arid biomes (i.e. Desert, Fynbos, Nama-Karoo and Succulent Karoo) as is shown for Acnephalomyia species (Fig. 64). This means that populations are found mainly in the south-western parts of southern Africa -including parts of southern Namibia, the western parts of the Northern Cape and Western Cape Provinces, and the western parts of the Eastern Cape Province of South Africa. Species occur both in the winter rainfall and summer rainfall areas of the subcontinent.
Biology Table 3 indicates that adult Acnephalomyia most species being found resting on the ground during October and November. Sporadothrix, Ammodaimon and Astiptomyia peaks during the summer months of January and February. Sporadothrix, however, may also be found during the autumn month of April. While little is known of the habits of Astiptomyia, and Ammodaimon, like Acnephalomyia, species appears to frequent the ground, while Sporadothrix has been found both on the ground and perching in vegetation. The few prey records that are available suggest that Acnephalomyia is a generalist, feeding on anything of an appropriate size that it gains access to.  3 Phenology of species of Acnephalomyia, Sporadothrix, Ammodaimon and Astiptomyia reviewed in this paper.
Months start with July so as to centre the data for these mainly summer-active species.