The First Record of the Genus Paralimosina L. Papp (Diptera: Sphaeroceridae) in the Afrotropical Region, with Descriptions of Six New Species

ABSTRACT The Afrotropical species of the genus Paralimosina L. Papp, 1973, are discussed. Six new species, Paralimosina australis sp. n. (Rwanda, Burundi, South Africa), P. congoensis sp. n. (Republic of Congo), P. flavifacies sp. n. (Tanzania), P. heteroneura sp. n. (South Africa), P. paraustralis sp. n. (South Africa), and P. sinelineata sp. n. (South Africa) are described. The morphological variety among the Afrotropical species of Paralimosina seems to be the widest. Therefore, it is expected that many other new species will be described from the region in the future. An identification key to the new taxa is provided.


INTRODUCTION
The genus Paralimosina L. Papp, 1973, includes a number of sphaerocerids of the sub family Limosininae that have robust and strongly sclerotised bodies. Genera in this subfamily have the epandrium of the male genitalia fused below the anal opening. Furthermore, important differentiating characters are located on the cercal part of the epandrial complex as well as the hypandrium, surstylus, postgonite and phallus. On the female postabdomen, the shape, armature and ratios of tergite 8 and sternite 8 can be characteristic of particular species. Likewise, the epiproct (tergite 10), cerci (not fused with tergite 10) and spermathecae may exhibit features that are diagnostic at the spe cific level.
Paralimosina is a diverse genus with 27 described species in the Palaearctic and Oriental regions but hitherto it has not been recorded from the Afrotropical Region (Ri chards 1980; Roháček et al. 2001). Hayashi (2008a) named five other species from Thailand, and Hayashi (2007Hayashi ( , 2008b described four new species from Malaysia.
Although I found six new species during the course of this study, it is highly likely that more new species of Paralimosina will be discovered in the future, since the morpho logical variety among the Afrotropical species of Paralimosina seems to be the widest.

MATERIAL AND METHODS
This paper is based on studies of many double-mounted specimens of Paralimo sina which are housed in the Hungarian Natural History Museum (HNHM; Budapest), Natio nal Museum (BMSA; Bloemfontein, South Africa), and KwaZulu-Natal Museum (NMSA; Pietermaritzburg, South Africa).
Abdomina of up to several (at least one) specimens of each species were removed and treated with 10 % hot sodium hydroxide (NaOH) for several minutes and lactic acid, then rinsed, etc., in the standard way. Those abdomina with genitalia are now kept in glycerol in plastic microvials under the respective pinned specimens.
In the species accounts below, geographical names on specimen labels are given in their present-day spelling, whereas my annotations concerning label data are in square brackets.
I name the sclerotised structures above the phallobase and anterior ventrally to the cer ci (with connections to epandrium and surstyli) as 'subepandrial sclerite' ('medandrium' of Roháček (1998), 'decasternum' auctorum), although its origin may be even more complex. TAXONOMY Genus Paralimosina L. Papp, 1973 Type species: Paralimosina kaszabi L. Papp, 1973: 376, by monotypy. Roháček and Papp (1988) described the genus correctly in respect of diagnostic fea tures. However, below I will nevertheless stress the characteristics of the male and fe male postabdomen and genitalia. Diagnosis: Body rather strongly sclerotised and in some species even granulose, variably with denser microtomentum or more shiny, despite presence of some microtomentum. Body length around 2.5 mm.
Head without inner orbitals, anterior fr-orb always clearly shorter than posterior pair. Frons with or without velvety black (or silvery) M-shaped mark (Roháček 1998: fig. 2). Facial plate normally convex, antennae not in deep hollow, or if in a shallow depression, edges are not sharp. Peristomals weaker, a genal seta present. Several (3-5) pairs of ifr, if one of them is very large, it emerges far from eye margin. First flagellomere not conical but if slightly so, never with rod-like projection. Eye normal or slightly reduced.
Mesonotum usually with only 1 or 2 dc pairs; when 3 dc present, then R 4+5 slightly re curved (Roháček 1998: fig. 93) or eye strongly reduced; no presutural dc pair. No setulae between apical scutellars; disc of scutellum usually bare. Scutellum never velvety black-marked, with only 2 pairs of long marginal setae, disc always bare. Anepisternum without setae.
Mid trochanter without or with a short seta. Mid tibia always without ventral preapical seta, but usually with one va (sometimes reduced only in males). Long mid ventral se ta on mid tibia present only in species with distinct va on mid tibia (Roháček 1998: fig. 75). Mid tibia with or without mid av. Mid basitarsus without long av and without ventral seta. Hind tibia with a long dorso-preapical seta, at most. Pulvilli and claws weakly de veloped; ventroapical spur of hind tibia mostly reduced.
Wing unicolorous, with cross-veins at most clouded in the Afrotropical species (there are numerous Oriental species with a patterned wing (Hayashi 2008a, b)). Wing without macrotrichia. First costal section basally with 2 shorter (paired) setae, one of them not longer than twice the length of the second, otherwise first costal section with short or medium-length setae only. Vein R 4+5 sinuate or upcurving.
Abdominal tergites not reduced, rather well sclerotised, marginal setae variable. Male postabdomen less downcurved; male cerci and S5 vary (but are usually simple). Male S4 normally without processes; S5 simple, without medio-caudal armature. Synsternite with basic structure characteristic of taxa in the subfamily Limosininae. Right side sclerites of the postabdomen (cf. Papp 2008) usually discernible but seldom with characteristic shape. Hypandrium with ventral single or bifurcate appendage (Roháček 1998: fig. 33). Epandrium with one connection on each side to hypandrium. Surstylus long but mostly not very high, in an anterior and a posterior lobe, the latter without robust ventral spines (Roháček 1998: fig. 108). Phallus in two parts: basiphallus short but usually high, well sclerotised without epiphallus, distiphallus varying, mostly with large ventral lobe(s), always shorter than phallapodeme, which is strong. Ejaculatory apo deme always discernible but its sclerotisation varies; and since it seems to be asymmetrical, its detectable shape is dependent on its positioning.
Female with seven normal preabdominal segments, although the length of the 7 th is variable. Postabdomen not telescoped. T8 consists of 2 lateral sclerites that have a pair of medially directed, narrow ventral processes, and may bear longer setae, mostly on the caudal margin. Features concerning S8 mostly species-specific. Epiproct (= T10, or T9 auctorum) usually flat, with or without some setae that are of diagnostic value. Hypoproct (= S10, or S9 auctorum) larger in several species and with short setulae, which are some times very thick (peg-like). Cerci always short, not fused to T10. Spermathecae 1+2, globular, vesiculate or subspherical, without acute projections. Sclerotised ducts usually short but the length (incl. length of individual ducts when they are paired) is species-specific. Body blackish brown with thick grey microtomentum, head mostly yellow, occiput black, fore femur dirty yellow, mid and hind femora dark brown.
Male abdominal S5 (Figs 1, 2) not particularly long, medio-caudally with a small thin desclerotised area, longer setae on caudal margin only. S6-8 not fused to each other, S6 large, i.e. comparatively long and broad (Fig. 3), S7 comparatively small with broadened caudal lobe (Fig. 5); S8 very large, particularly long. There is a well scle rotised sclerite located ventrally to S8, which cannot be interpreted as anything but a tergal part. Epandrium with long setae, particularly so ventrally. Cercal lobe (Figs 7,8) large and broad, with no thick setae or sclerotised processes ventrally on epandrial complex. Hypandrium (Figs 4,6) with robust, broad, asymmetrical medial part ('rod'), . Paralimosina flavifacies sp. n., holotype male, postabdomen and genitalia: (1) S5, ventral view; (2) caudal part of S5 at higher magnification; (3) S6, ventral view; (4) hypandrium, ventral view; (5) S7-8, ventral view; (6) hypandrium, lateral view, with ventral process at higher magnification. Scale bars: Fig and with short arms, which are not fused to epandrium. Medial part with an asymmetrical ventral process medially, which is serrate on anterior edge (Fig. 6). Connecting sclerites of hypandrium strong (Fig. 4). It is much shorter than phallapodeme, medial part with a long caudal process. Subepandrial sclerite (Figs 7,8) with long process to surstylus, its body higher than broad. Surstylus (Figs 9, 10) very characteristic: anterior lobe smaller than posterior lobe, subtriangular with short setulae (pegs) only; posterior lobe large, with a short, setose caudal process and a small triangular ventral lobe; longer setae on ventral third of surstylus only. Postgonite (Fig. 13) robust with strong basal part, even apical part of postgonite not thin, with curved blunt apex; anterior edge bears some short setae. Phallus short (Figs 11,12), basiphallus rather small, particularly short. Distiphallus (Figs 11,12) is an intricate structure: it has a large ventral process, which bears a digitiform ventral process; both apical process and ventral edge above the ventral process serrate. Phallapodeme ( Fig. 12) comparatively short and thick. Female T8 in 2 parts as usual (Fig. 14), each with a large, medially directed process having some stronger lateral setae only. S8 very short, although broad, with a pair of wide ly separated setae (Fig. 16). Epiproct extremely large, broad and triangular, with a pair of widely separated setae and longer but fine hairs apically and on a pair of low, slightly curved, dorsal ridges (Fig. 15). Cerci short and thin, with a medium-length se ta and several short pairs of setae. Spectacles-shaped sclerite, rather well-sclerotised, with a stronger, melanised cylindrical structure caudally (Fig. 17). Spermathecae (Fig. 18 Frons with distinct M-shaped mark (silvery on dark brown). Lunular triangle and face shiny dark brown, occiput dark brown. Three or four pairs of almost evenly long ifr plus short anterior pair (holotype with 3 long ifr pairs on the right side, left side with 4 pairs plus a 9 th thinner seta). Height of gena below eye 0.18 mm, longest axis of eye 0.34 mm, vertical diameter 0.28 mm. Upper 0.6 of gena longitudinally hachured. Genal seta emerges 0.09 mm above mouth margin, 0.15 mm long. Anterior fr-orb (0.15 mm) only half as long as posterior one (0.31 mm). Medial seta of scape 0.11 mm long. First flagellomere with cilia 0.025 mm in length. Aristal cilia ca 0.015 mm.
Abdomen shiny black. Longest lateral marginal setae on male abdominal tergite only 0.25 mm. Male S5 (Fig. 23) rather long (but much shorter than that of P. paraustralis), with a very short medio-caudal extension lacking a seta. That medio-caudal part is only slightly more melanised. S6 part of synsternite (Fig. 28) rather large, comparatively long and broad, on the right side distinct sclerotised and melanised sclerites join it ('right side sclerites', see Papp (2008)). Those less formed sclerites are almost per pen dicular to the abdominal axis (and join broadly, although membranously, also to S5). S7 part of synsternite large, with short thin setae on all of its surface. S8 part of synsternite small. Hypandrial arms rather short and slightly curved; medial part strong, with a sagittal ridge and an anteriorly curved asymmetrical process (Fig. 25). Subepandrial sclerite rather narrow (Fig. 24), its body only 0.07-0.08 mm high, ventral processes comparatively short and thick. Cercal lobes of epandrium very small. They can be regarded as reduced, with sagittal connection of the epandrial complex very low (Fig. 24). Surstylus with very large anterior lobe being an intricate structure: anterior and dorsal parts less strongly sclerotised (Fig. 29). Posterior lobe of surstylus (Fig. 29) much smaller; strongly scle ro tised and melanised, with numerous medium-length setae. Phallapodeme strong (Fig. 26), ca 0.3 mm long and almost straight in lateral view. Basiphallus (Figs 26, 27) short and high, ventral apex blunt and slightly curved. Distiphallus (Fig. 26) short, only 0.17 mm, without conspicuous processes. Postgonite rather long (0.22-0.23 mm from apex to base), basal part not strongly broadened, clearly curved with blunt apex (Fig.  30). Ejaculatory apodeme (Fig. 27) discernible, but only 0.055 mm long. Female abdomen 1.62 mm long and ca 1.14 mm wide, without long lateral marginal setae. Desclerotised 'window' of T1+2 with a pair of less sclerotised, round plates. T5 only slightly broader than half the width of abdomen. S2 as broad as ⅔ of width of S3 (i.e. 0.35 mm). S6 is 0.46 mm long and 0.165 mm wide. T8 (Fig. 20) subquadratic in lateral view, without medial process, with several short setae laterally. S8 (Fig. 19) rather long and rounded, with a pair of long setae laterally and having 7 or 8 short setae; anterior margin free of small hairs. Epiproct with a pair of long, more closely set setae (Fig. 21); lateral anterior edges free of small hairs. Cerci (Fig. 21) with a pair of very long (0.21 mm) apical setae and a somewhat shorter pair of medial subapical setae; a third pair of long lateral setae also present. Spermathecae (Fig. 22)  Body dark brown; frons with M-shaped pattern. As regards body features, the male of the new species is very similar to P. australis sp. n. but differs in the following: mid tibia without a row of thick black setae ventrally, but apical 0.4 has long dense hairs; no va on male mid tibia; fore coxa with some longer hairs only; costal index 1.0. S5 large and very long (Fig. 31), even longer medially; that medial extension with a very short medio-caudal edge, which does not bear a seta. The medio-caudal part is only slightly more melanised. S5 medially with a pair of long subcaudal setae, short setae present between them and also laterally. S6 part of synsternite (Fig. 32) rather large, comparatively long and broad, 'right side sclerites' less distinct than in P. australis. S7 part of synsternite with an exceptionally large dorsal part. S8 part small. Hypandrial arms (Fig. 37) rather short and broad, not curved, medial part strong, with a lower sa gittal ridge and an anteriorly curved, asymmetrical process (Fig. 37). Subepandrial scle rite (Fig. 34) rather similar to that of P. australis. Cercal lobes of epandrium can be regarded as reduced but sagittal bridge definitely broader (i.e. higher). Surstylus with extremely large, intricately structured anterior lobe: anterior and dorsal parts less strongly sclerotised (Fig. 35). Posterior lobe of surstylus (Fig. 35) much smaller, with a double-curved long black bristle (Figs 34, 35); posterior lobe strongly sclerotised and melanised and bears several setae of medium length. Phallapodeme (Fig. 33) less strong, ca 0.23 mm long and with anterior part ventrally curved in lateral view. Basiphallus (Figs 26, 27) short and high, ventral apex blunt and slightly curved. Distiphallus short (Fig. 33), in contrast to that of P. australis, with several apical processes and a ventral process. Postgonite (Fig. 36) shorter than in P. australis (less than 0.2 mm from apex to base), thinner and less curved than that of P. australis, basal part not strongly broadened, with blunt apex. Remark: The discovery of two such closely related species of the Afrotropical Paralimosina must make us aware of the possible existence of similar sister species in the region, i.e. of the need for genitalia preparations from specimens in the future.

Figs 38-41
Etymology. The species is named after Congo. Description:

Female.
Measurements in mm: body length 1.94, wing length 1.54, wing width 0.70. Body dark brown, including frons (the specimen was kept in alcohol for about 3 decades but colours have not faded much); tarsi ochre.
Subantennal swelling of facial plate rather large. Frons with silvery-striped M-shaped pattern. Three pairs of ifr, anterior one minute, 2 nd and 3 rd pairs of medium length (0.08 mm), the posterior pair is anteriad of anterior fr-orb. Anterior fr-orb 0.09 mm, posterior 0.15 mm long. Vibrissa 0.25 mm long, genal seta not far from mouth margin, strongly upcurving, 0.11 mm long. Antenna 0.23 mm in length. Scape with a thin but very long (0.105 mm) seta. Pedicel with similarly long setae. First flagellomere with dorsal apical conus and with cilia 0.04 mm long. Arista short (ca 0.50 mm), aristal cilia 0.04 mm.
Wing membrane light brownish, veins ochre, longest sub-basal seta of costa 0.12 mm. Setae on first costal section 0.06-0.07 mm, on second section half as long. Costa terminates just at conjunction with R 4+5 . Second costal section 0.42 mm, third costal sec- form an open ring of 0.26 mm in diameter. T8 sub-pentagonal, with several moderately long, caudally directed setae (Fig. 39). S8 bipartite (Fig. 40), proximal part with seta(e) of 0.08 mm and a belt covered by dense, short, sharp setulae; distal part quadratic but the more melanised part is T-shaped.
Postabdomen slightly protruding. Epiproct rather broad, with a pair of long setae (Fig.  38). Cerci small and only 0.05 mm long, with a pair of dorsal medial setae 0.13 mm in length and with some short setae. Spermathecae (Fig. 41) are not flattened, and are vesiculate like some of the Palaearctic species (see Roháček & Papp 1988), ducts of medium length and not bulbous. Ten mature yellowish eggs of 0.56×0.10 mm and several immature eggs were found in the abdomen of the holotype when dissected.
Body dark blackish brown, legs black, tarsi brown. Frons with distinct M-shaped pattern (silvery on blackish brown). Face moderately shiny. Three pairs of long ifr plus a minute anterior pair (holotype), paratype female with a 4 th posterior ifr on the left side. The posterior ifr pair is posterior to anterior frorb. Genal seta 0.11 mm long and emerges only 0.03 mm above mouth margin. Scape and pedicel dirty reddish, first flagellomere dark grey. Medial seta of scape light and thin; and 0.10 mm in length. Longest cilia on first flagellomere 0.05 mm, aristal cilia slightly more than 0.01 mm.
Male S5 (Fig. 42) with a quadrate medio-caudal extension, which is bare. At least 1 pair of very long, submedial, subcaudal setae and several other long setae on S5. A broad, medial, caudal, less melanised area detectable, which bears short setulae. S6-7 parts of synsternite broadly and strongly fused (Fig. 43), S8 component medium-large and not fused to S7 section. Ventral portion of S6 thin, and no sclerotised 'right side sclerites' were found. Hypandrium (Figs 44, 45) comparatively short but robust. Hypandrial arms not fused to either the epandrium or to the medial part ('rod'), arms short with broadened caudal ends (to epandrium); medial part with a large asymmetrical ventral process. No cercal lobes on epandrial complex (Figs 47,48). Ventral part of epandrial complex with some medium-length setae and dense thin hairs. Subepandrial sclerite about as broad as high, ventral processes short (Fig. 48). Surstylus with a bilobed, less sclerotised and melanised anterior part; both lobes are rounded (Fig. 50). Part connecting to posterior section comparatively thin (narrow) but well-sclerotised. Posterior region of surstylus with a long, blunt ventral lobe and several long setae. Phallapodeme 0.24 mm, curved in lateral view (Fig. 46). Phallus (Fig. 46) short, ca 0.20 mm. Basiphallus short and not very high, distiphallus with a long, thin, ventrally and caudally curved pair of processes and having a ventrally directed digitiform medial process. Postgonite rather broad-based (Fig. 49), basal 0.8 almost straight, only apical 0.2 narrowed and anteriorly curved. Apex of postgonite narrowly rounded.
Female T8 without medially directed process and with only ca 4 medium-length setae (Fig. 53). S8 trapezoid (Fig. 54), with a pair of sublateral setae 0.08 mm long, as well as 4 other short setae; anterior part free of short hairs. Hypoproct, as usual, broad and narrow, with a pair of short widely separated setae. Epiproct broad and semicrescentic (Fig. 51), with a pair of short (ca 0.03 mm) setae (on the only female paratype, a third asymmetrically placed seta also present). Cerci short, with a long (0.10 mm), al most straight pair of setae; other cercal setae short. Spermathecae (Fig. 52)  Frons without M-shaped mark, finely microtomentose brown, anterior part ochre. Four ifr pairs, anteriormost one prominent but not more than 0.05 mm. Antennae well separated. Medial seta on scape 0.06 mm long, genal seta 0.06 mm. First flagellomere with cilia 0.025 mm long (apical aristomeres broken off, so aristal cilia not measurable).
Anterior katepisternal seta only 0.04 mm long, posterior one 0.14 mm. Wing comparatively short, rounded, uniformly light brown, veins slightly darker, only costal vein darker brown. Basal seta on costa 0.09 mm. Male S5 (Fig. 55) broad and rather long with a broad, medial caudal extension that continued into a narrow membranous margin, which bears short hairs; there are mediumlength setae caudally, medially and also on lateral edges. S6-7 parts of synsternite strongly and broadly fused, medial part of S6 not extended much to the right, dorsal part of S7 small. S8 component of synsternite not fused to S7 part, broad on the left side dorsally, narrowed to the right side. No cercal lobes on epandrial complex but the connection of the two halves is broad sagittally (Fig. 57). Subepandrial sclerite al most as high as broad, ventral processes rather short. Hypandrium robust, with a large asymmetrical ventral process (Fig. 59). Surstylus (Fig. 58) with rounded, less scle rotised and melanised anterior part, with comparatively broad connecting part and rather large posterior section, which bears numerous long and medium-length setae. Phal lapodeme ca 0.22 mm long, slightly curved in lateral view. Phallus (Fig. 60) short (<0.2 mm), basiphallus short but comparatively high, distiphallus with a dorsal, short process medially and a pair of subapical processes, having a serrated apex (Fig. 60). Postgonite with broad base, apical ⅓ narrowed, apex sharp (Fig. 61). Anterior edge with 2 medium-length (a little more than 0.02 mm) setae.
Female abdomen 0.75 as broad as long (postabdomen not at all telescopic), without any long setae, longest marginal setae of tergites only 0.08 mm. S2 trapezoid, medial ly very weakly sclerotised and melanised; as a result of NaOH preparation, it looks bipar tite. S3-5 are ⅓ to / as broad as tergites. S7 in its natural position (arched) 0.10 mm long and 0.25 mm broad. S8 shield-shaped, with broadly rounded caudal margin, 0.12 mm wide, 0.11 mm long with setae ca 0.05-0.06 mm in length on its apical 0.6. Syntergite 0.4 section as long as wide, T1 part weakly sclerotised and light medially, T2 component less darkly melanised medially but not less sclerotised than other parts. T3-6 are 0.25, 0.25, 0.23, 0.2 as long as width of abdomen at their position. T7 is 0.3× as broad as syntergite. T8 in 2 completely laterally placed, comparatively large sclerites (ca 0.11 mm long and 0.15 mm broad), and with a ca 0.04 mm long and nearly 0.20 mm wide membranous central part. Epiproct 0.175 mm broad but only 0.050 mm long, no setal pair on epiproct. Hypoproct 0.18 mm in width, broader than a semicircle, with numerous mar ginal setulae 0.020-0.025 mm long. Cerci strongly oblique to the body axis (ca 45°), with a seta 0.11 mm in length and several shorter pairs of setae. Spermathecal ducts not long, so sper mathecae are situated in the 7 th segment. Spermathecae pear-shaped,