Zootaxa 4337 (4): 451–492 ISSN 1175-5326 (print edition) Article ZOOTAXA http://www.mapress.com/j/zt/ Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4337.4.1 http://zoobank.org/urn:lsid:zoobank.org:pub:BA7E7DE5-25C2-41BA-8642-9B429FDC5294 Taxonomic revision of Paraustrosimulium Wygodzinsky & Coscarón: reassignment of Austrosimulium colboi and description of P. obcidens n. sp. from Western Australia DOUGLAS A. CRAIG1, JOHN K. MOULTON2 & DOUGLAS C. CURRIE3 1Department of Biological Sciences, University of Alberta, Edmonton, Canada, T6G 2E9. E-mail: [email protected]. orcID: 0000– 0002–9269–8826 2Department of Entomology and Plant Pathology, 205 Ellington Plant Sciences Building, The University of Tennessee, Knoxville, TN, USA 37996–4560. E-mail: [email protected] 3Department of Natural History, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario, Canada M5S 2C6. E-mail: [email protected] Abstract The hitherto monotypic South American genus Paraustrosimulium Wygodzinsky & Coscarón is revised to accommodate two Australian species: Austrosimulium colboi Davies & Györkös and Paraustrosimulium obcidens n. sp. The generic di- agnosis is updated and the eastern Australian species Paraustrosimulium colboi (Davies & Györkös) n. stat. is re-de- scribed, including the male for the first time. The Western, Australian sister species of P. colbo, namely P. obcidens Craig, Moulton Currie n. sp. is also fully described. The relationship of Paraustrosimulium to other simuliid genera is discussed, as are aspects of historical biogeography. Key words: Australia, South America, Gondwana, Diptera, Simuliidae, Austrosimulium, Paraustrosimulium, Cnesia- mima Introduction The relationships of the southern South American species described as Simulium anthracinum Bigot, 1888, has long intrigued students of Simuliidae. Edwards (1931) was the first specialist to recognize the close relationship between that species and Austrosimulium Tonnoir—a lineage that was, until then, considered endemic to Australia and New Zealand. Although only adult females were known at that time, most specialists accepted the position of anthracinum within Austrosimulium (e.g., Smart 1945, Wydozinsky 1953). Wygodzinsky (loc cit.) described the pupa for the first time and redescribed and figured the female. The larva was first described by Dumbleton (1960) who—although expressing doubts about a close relationship between anthracinum and Austrosimulium, nonetheless retained that taxonomy. Wygodzinsky & Coscarón (1962) described the male for the first time and addressed Dumbleton’s reservations, proposing the monotypic subgenus Paraustrosimulium for A. anthracinum. That segregate was eventually elevated to full genus by Crosskey (1969: 17)—a status generally accepted by most subsequent workers to the present day (e.g., Wygodzinsky & Coscarón 1973, Adler & Crosskey 2017). Recent phylogenetic studies provide contradictory evidence about the relationship of Paraustrosimulium to Austrosimulium. An analysis of morphological characters by Gil-Azevedo & Maia-Herzog (2007) suggests a close (sister group) relationship between the two genera; in contrast, the molecular analysis of Moulton (2003) reveals that P. anthracinum is rather distantly related from all Austrosimulium species analyzed except "A". colboi Davies & Györkös 1988. Indeed, the position of the sister pair P. anthracinum + "A". colboi was so distant from other analyzed Austrosimulium spp. that Moulton (loc. cit.) suggested the latter species should be reassigned to Paraustrosimulium. The objectives of this paper are to (1) to reassess the current assignment of colboi in Austrosimulium, (2) provide an expanded definition of Paraustrosimulium to include colboi, plus a previously Accepted by L. Hernandez-Triana: 5 Sept. 2017; published: 20 Oct. 2017 451 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 undescribed species from Western Australian, (3) redescribe P. colboi, including the male for the first time, (4) describe a new species of Paraustrosimulium from Western Australia, and (5) offer comments about historical biogeography and the relationships of Paraustrosimulium to other austral simuliid genera. Material and methods We follow terms for structures as in Craig et al. (2012), which were based mainly on those of Adler et al. (2004). One character infrequently referred to in the simuliid literature are projections from ventral sclerites of the adult cervix. Such structures were referred to as "cervical laterales" by Davies et al. (1962: 133) and "cervical sclerites" by McAlpine et al. (1981: 24). We adopt the latter term here. For locality data, individual labels are indicated by [ ], with a / indicating a line break. Male and female signs, where given, are as {M} and {F} respectively. Davies & Györkös (1988) stated that type material of "A". colboi was deposited in the Australian National Insect Collection (ANIC); however, Bugledich (1999: 330) noted that it could not be found there. Rediscovery of that material was outlined by Craig (2011: 4). In short, the material was never sent to ANIC, and instead had remained in the D.M. Davies Collection at McMaster University until its rediscovery by DAC (loc. cit.). Only some of the original type material was recovered. Material examined of P. anthracinum was from the D.M. Davies Collection. Other material of colboi was also examined from that collection, plus specimens collected by the authors during three separate collecting trips between 1996 and 2014. Material of P. obcidens n. sp. was collected by JKM in 1996 and by JKM and DCC in 2014. All specimens described in this paper are deposited either in the Australian National Insect Collection, CSIRO, Canberra (ANIC), or the University of Alberta Strickland Museum (UASM), as indicated. The generic affinities of Austrosimulium colboi Davies & Györkös, 1988 In 1968, D.M. Davies, McMaster University, was presented with a collection of Simuliidae by Ron Pilfrey, who had studied the Australian simuliid fauna under the guidance of specialist Ian Mackerras. Twenty years later, Davies & Györkös (1988) described two new and unusual species from that material, even though it was badly bleached. “Cnephia” pilfreyi and “Austrosimulium” colboi, were compared to a number of different genera from Australia, southern Africa and South American, and tentatively assigned to genus pending collection of additional material. The first specialist to seriously question the assignment of colboi to Austrosimulium was Moulton (2003: 53), whose molecular analysis of a broad spectrum of simuliid genera placed "A". colboi as sister to Paraustrosimulium—distantly separated from other species of Austrosimulium in his analysis. Moulton (loc. cit.) further noted possible morphological synapomorphies shared between "A". colboi, P. anthracinum, plus two other species: “Cnephia” pilfreyi and Cnesiamima atroparva (Edwards, 1931). Craig et al. (2012) lent support for this hypothesis by suggesting that "A". colboi could be reassigned to Paraustrosimulium based on features of the male genitalia and pupal gill. Further evidence can be found in a recently published neighbor-joining tree of COI DNA barcodes for species of austral Simuliidae (Hernández-Triana et al. 2017). Although such trees do not represent a phylogeny, "A". colboi and P. anthracinum were relatively closely placed on that tree, but distantly so from seven other species of Austrosimulium, which all clustered together. In view of this growing body of evidence, we formally reassign "Austrosimulium" colboi to Paraustrosimulium. The genus is here re-diagnosed to accommodate that species, plus a previously undescribed one from Western Australia. Paraustrosimulium (Wygodzinsky & Coscarón) Austrosimulium (Paraustrosimulium) Wygodzinsky & Coscarón 1962: 244. New subgenus. Stone 1963: 14. Austrosimulium (Paraustrosimulium). Dumbleton 1963: 333. Dumbleton 1973: 484. Paraustrosimulium. Crosskey 1969: 17. Raised to generic status. Wygodzinsky & Coscarón 1973: 189. Crosskey 1982: 16. Crosskey 1987: 438. Crosskey & Howard 1997: 18. 2004: 10: Adler & Crosskey 2017: 31. Paraustrosimulium. Hernández-Triana et al. 2017: 350. 452 · Zootaxa 4337 (4) © 2017 Magnolia Press CRAIG ET AL. Re-diagnosis. Adult: Antenna with 8 flagellomeres; frons width variable; female mandible expanded subapically (colboi, obcidens), not so (anthracinum), teeth on both sides; lateral cervical sclerites markedly expressed; antepronotal lobe markedly hirsute (anthracinum), or not (colboi, obcidens); anepisternal (aka pleural) membrane bare; metathoracic furcasternum with dorsal projections; wing with small to minute basal cell; costa with spiniform setae; apical end of R with (anthracinum) or without fine spiniform setae; Rs not branched, but may be thickened 1 apically with indication of branching; M appearing thickened apically; Cu curved; female thorax not markedly domed in Australian species, more so for anthracinum, katepisternal sulcus well defined; calcipala well expressed, rounded apically; pedisulcus as series of wrinkles (anthracinum) or definite depression (colboi, obcidens); claw with (anthracinum, obcidens) or without (colboi) definite tooth, when present separated from basal heel by distinct notch; spermatheca externally smooth, internally with sparse acanthae, pigment well extended along duct (obcidens), only slightly (colboi) or not (anthracinum); genital fork with short broad anterior stem, forwardly directed apodemes poorly expressed or absent, lateral arms short, apical lobes large; cercus rounded, anal lobe angulate proximally (colbo, obcidens), tapered (anthracinum); male gonostylus angulate with 2–4 terminal spines; ventral plate angulate laterally, with (colboi, anthracinum) or poorly expressed (obcidens) median keel, median sclerite well developed and M–shaped; parameres plate–like, numerous fine small parameral spines. Pupa: cocoon slipper–shaped, lacking anteroventral collar, well defined anterior edge, no marked medial projection, ventral floor present (colboi) or absent; pupal cuticle brown anteriorly; cephalic and thoracic cuticle with low tubercles; thoracic dorsocentral setae distinct and curled apically; cephalic plate with frontal and facial setae present; gill of simple construction, either annulated tapered tubular construction (colbo, obcidens) or, annulated flattened lamellae (anthracinum), concertinaed in histoblast; abdominal armature not markedly developed, distinct pleurites not present or markedly small; tergites III and IV with 4+4 hooks posteriorly, V to IX with spine combs anteriorly, sternum V with 4+4 hooks posteriorly, sternum VI and VII with 1+1 hooks, or all sternal hooks absent (colboi), small hooks on small pleurites VI and VII; poorly expressed spine comb on tergite IX, terminal spines short and blunt, grapnel hooks exacerbated. Larva: cervical sclerites small, subcircular to elongated, not fused to postocciput or slightly so; postgenal cleft well developed (colbo, obcidens) or shallow (anthracinum); antenna slightly longer than labral fan stem, medial plus basal antennomeres shorter than distal antennomere. Mandibles with apical teeth on narrowed extension, spinous teeth fine and pointed, mandibular serration and sensillum as separate structures; hypostoma with 13 teeth not arranged in distinct groups, underlain by ventral edge of hypostoma; abdominal cuticle bare, except for rectal scales only in anthracinum; anal sclerite X–shaped, with lateral thickening of medial region (aka interarm struts); accessory and semicircular sclerites absent; rectal papillae of three simple lobes. Constituents. Paraustrosimulium anthracinum (Bigot) (South America), Paraustrosimulium colboi (Davies & Györkös) (Victoria, Australia), P. obcidens Craig, Moulton, Currie n. sp. (Western Australia). Paraustrosimulium colboi (Davies & Györkös), 1988 (Figs. 1–42) Austrosimulium colboi Davies & Györkös, 1988: 111. Original designation. Provisional placement to genus. Austrosimulium (Austrosimulium) colboi. Crosskey 1989: 221. Austrosimulium colboi. Crosskey & Howard 1997: 26. Unplaced to subgenus. Austrosimulium (Austrosimulium) colboi. Bugledich 1999: 330. “Austrosimulium colboi”. Moulton 2003: 47. Austrosimulium colboi. Crosskey & Howard 2004: 18. Adler & Crosskey 2009: 19; 2017: 30. Unplaced to subgenus. ?Austrosimulium colboi. Craig et al. 2012: 53. Austrosimulium colboi. Hernández-Triana et al. 2017: 350. Paraustrosimulium colboi. This work, new combination. Redescription. Adult female (based on numbers of reared specimen). Body (Fig. 1): general body colour in alcohol evenly blackish brown; total length 1.9–2.3 mm. Head (Fig. 2): width 0.57–0.60 mm; depth 0.4 mm; postocciput black, hirsute; frons black; frons–head ratio (narrowest width of frons: greatest width of head) 1.0:6.3. Eyes: slightly bicolourous, interocular distance 0.09 mm; ommatidia 0.015 mm in diameter; 35–39 rows up and across at mid–eye. Clypeus: black; 0.17mm wide; vestiture of sparse black hairs. Antenna (Fig. 3): total length 0.41–0.46 mm; pedicel small, scape enlarged, both blackish brown, remainder brown; 8 flagellomeres, basal ones wider than long, distally ones more quadratic, overall tapered, apical flagellomere distinctly so. Mouthparts: substantial, ca. TAXONOMIC REVISION OF PARAUSTROSIMULIUM Zootaxa 4337 (4) © 2017 Magnolia Press · 453 0.7 length of head depth; maxillary palpus (Fig. 4), total length 0.45 mm, articles overall brownish black, 3rd article darker; proportional length of 3rd, 4th and 5th articles 1.0:0.8:1.5; sensory vesicle ovoid, 0.5x width of 3rd article, opening 0.3x width of vesicle; mandible (Fig. 5), broadly triangular apically, sharply pointed with 48–50 inner teeth and 9–11 finely pointed outer teeth; lacinia (Fig. 6) with 13–15 inner teeth and 22–26 outer teeth; cibarial cornuae (Fig. 7) (partly reconstructed) lacking apical fluting or sculpture, sharply terminated, central depression broad. Cervical sclerites markedly developed (Fig. 1). Thorax: moderately domed; length 0.80–0.96 mm; width 0.60–0.63 mm; scutum evenly blackish brown, vestiture of evenly distributed recumbent silver hairs; postpronotal lobe with longer hairs; antepronotal lobe with longer hairs; scutellum and postnotum concolourous with scutum; scutellar depression and scutellum with long substantial black hairs; pleuron and anepisternal membrane blackish brown, bare. Haltere: dark. Wing (Figs. 8, 9): length 2.1–2.4 mm; width 0.9–1.2 mm, veins lightly pigmented, costa not extended to wing apex with spiniform setae distally, absent from other veins; radial veins closely applied to costa; basal cell absent; a:b ratio 1.0:2.8. Metathoracic furcasternum (Fig. 10): dorsal arm with distinct projection. Legs (Fig. 1): overall blackish yellow; hind basitarsus about 5.5x as long as its greatest breadth, ventral row of stout spines absent, calcipala as long as wide 0.75x width of basitarsus, pedisulcus present, but not markedly developed (Fig. 11); tarsal claw (Fig. 12) with moderately developed basal heel and markedly small tooth. Abdomen (Fig. 13): abdominal scale black with dark hairs, not greatly extended; tergite II 5x wider than long, shallowly V–shaped, III– V as wide as long, rounded, VI 2x wider than long; dorsal vestiture of small black hairs increased in density posteriorly. Genitalia: sternite VIII vestiture of sparse coarse black hairs posterolaterally; hypogynial valves (Fig. 14) short, lightly pigmented with vestiture of sparse short hairs and triads of microtrichia; median edges slightly convex, slightly strengthened along edge, bluntly rounded apically; genital fork (Fig. 15) with anterior stem broad (not easily observed), but well sclerotized and pigmented narrowly medially, flared anteriorly, lateral arms broad, indications of lateral apodeme (as in Gigantodax), more distal apodeme (as in Austrosimulium) present only as ridge, lateral plates large, angular posteromedially, rounded posterolaterally; spermatheca ovoid (Fig. 16), surface un–patterned; sparse internal acanthae in pairs and triads; junction of spermathecal duct pigmented and slightly sculpted; cercus (Fig. 17) as wide as long, broadly rounded in lateral view, anal lobe shallow, angulate proximally. Adult male (reared specimens and others). Body (Fig. 18): general colour evenly black; total length 2.2–2.6 mm. Head (Fig. 19): width 0.89 mm; depth 0.6 mm; wider than thorax. Eyes: upper ommatidia very dark red, 0.04 mm in diameter, ca. 14 across, 20 down; lower ommatidia almost black, 0.018 mm in diameter, ca. 35 across and 23 down. Clypeus: black; vestiture of very sparse black hairs; 0.15 mm wide. Antenna (Fig. 20): total length 0.51– 0.55 mm; pedicel longer and wider than other divisions; first flagellomere longer than wide, others subrectangular; non–tapered, markedly narrow in comparison to that of female; scape and pedicel black, first flagellomere dark brown remainder evenly brown; flagellum markedly not hirsute. Mouthparts: poorly developed; length 0.26x head depth; mandibles insubstantial, finely tapered with apical hairs; laciniae as for mandible; maxillary palpus dark brown, 0.4 mm long, proportional lengths of 3rd, 4th and 5th articles 1.0:1.1:2.0, sensory vesicle small, occupying 0.33x width of article, opening 0.5x width of vesicle. Cervical sclerites markedly developed. Thorax: length 0.9– 1.1 mm; width 0.8 mm; in alcohol, scutum evenly velvety black, vestiture of fine recumbent pale hairs; scutellum and postscutellum concolourous with scutum, coarse long black hairs. Wing: 2.1–2.4 mm in length, 1.0–1.1 mm in width; otherwise as for female. Haltere: tan. Legs: blackish brown; hind basitarsus about 5.5x as long as its greatest breadth, lacking row of stout spines; tarsal claw (Fig. 21) partially covered by grappling pad of ca. 20 hooks, distinct small basal tooth and heel. Abdomen (Fig. 22): black; abdominal scale with long fine hairs, tergites markedly broad, on tergites II–V hirsute laterally, less so on posterior others. Genitalia: ventral view (Fig. 23), ventral plate directed ventrally giving appearance of broadly concave apex, 1.5–2.0x wider than long, median keel well developed, with vestiture of fine hairs, plate roughly sculpted laterally; anteromedial depression broadly U to V–shaped, slight central convexity, basal arms short and pigmented (Fig. 24); median sclerite poorly expressed, broad and slightly divided apically; parameres present, moderately expressed, plate–like, apical rows of small spicules; adeagal membrane with sparse rows of minute microtrichia; gonocoxa 1.7x longer than basal width, markedly coarse black hairs on distal half; gonostylus (Fig. 25), approximately 3.0x longer than basal width, apically with 3 substantial terminal spines—variable and occasionally with one spine markedly displaced to outer apex. Pupa (based on numbers of specimens). (Fig. 26). Body: length, female 2.2–2.8 mm; male 2.2–3.3 mm. Head: frontal cephalic plate lacking dorsal depression; ratio of basal width to vertex width of female 1.0:1.5, for basal width to length 1.0:2.3, rounded apically (Fig. 28), male ratios 1.0:1.6 and 1.0:2.3 respectively (Fig. 29); evenly 454 · Zootaxa 4337 (4) © 2017 Magnolia Press CRAIG ET AL. tuberculate, frontal setae absent, facial setae present, but insubstantial, ocular spine absent. Thorax. Dorsum with very small tubercles, no pattern (Fig. 30); dorsocentral setae spine–like and curled apically. Gill (Fig. 26, 27): basically a single expanded tapered tube directed anteroventrally, 1.7–2.1 mm, full length 0.3 mm at greatest width with ca. 19 annulations on the longer anterior portion with 2 or 3 on a stub–like posterior lobe (Fig. 27), fenestra normal (Fig. 30), fine filaments absent. Abdominal armature: sternal hooks absent, but sternite VIII with sparse multi–pointed scales; tergites I & II with 4–6 moderately expressed setae; tergite III with four hooks posteriorly per side, 3 or 4 other fine setae, no lateral hook; tergite IV as for III, but with single small hook laterally; tergites V– VIII with markedly poorly–expressed spine comb anteriorly, laterally morphing into low multi–pointed scales, four to five fine setae posteriorly; tergite IX with poorly–expressed broad row of low spines and scales anterad, terminal spines short, not markedly sharp, grapnel hooks well expressed (Fig. 31). Cocoon (Fig. 26). Surface smooth, fabric coarsely woven, silk filaments obvious, medium brown; distinctly slipper–shaped fully covering pupa with well defined anterior edge, slight median projection, complete ventral floor, close fitted to pupa. Larva (based on numbers of last instar larvae). Body (Fig. 32): overall grayish brown, stubby (head large in relation the body), thorax and anterior abdomen subequal in diameter, expanded smoothly posteriorly; total length 4.0–4.9 mm. Head (Figs. 33): distinctly bicolourous, background pale translucent yellowish brown, head spots marked, but not strongly pigmented, medial and posterior spots form distinct inverted T, mediolateral spots distinct; most spots with negative centre; probable males have a lighter head pattern than females; head length 0.69–0.70 mm, width 0.50–0.60 mm; distance between antennal bases 0.30–0.38 mm; lateral margins of head smoothly convex, more so posteriorly; cervical sclerites well developed and pigmented, rounded posteriorly but flared anteriorly, not fused to postocciput; anterolateral edges of apotome sharply pigmented; genae with darker 'eye brow' above stemmata. Antenna (Fig. 34): overall clear pale brown; total length 0.4 mm; well extended beyond labral fan stem; distal article slightly longer than other two articles combined; proportions of basal, median and distal article 1.0: 0.6: 1.8. Labral fan: stem markedly translucent; 62–64 fine rays, 0.56 mm in length, 0.01 mm in width; microtrichia as long as ray width, distinct pattern with ca. 5 microtrichia decreased rapidly in length. Mandible (Fig. 35): apical brush well developed with distinct pigmented base: apical teeth not markedly developed; subapical teeth small, 8–9 spinous teeth; 2 distinct serrations widely separated, sensillum distinct and finely pointed; blade region long, smooth and slightly concave. Maxilla (Fig. 36): lobe markedly cone–shaped, asymmetrical, palp longer than lobe, closely applied, 3x as long as basal width. Postgenal cleft (Fig. 37): small, but U–shaped with irregular anterior apex, sclerotized posterior tentorial pit cuticle extensive, pits ovoid; ratio of hypostoma, bridge and cleft 1.0:1.9:0.7; suboesophageal ganglion not pigmented. Hypostoma (Fig. 38): ventral edge as raised dome; 13 teeth, median tooth barely protruded beyond edge, two sublateral teeth smaller and subequal in length, other larger and protruded beyond edge, lateral teeth larger and well protruded, paralateral teeth smaller and sharp, variable expression, no others evident; no hypostomal serrations; four hypostomal setae per side. Postgenal bridge: pale and concolourous with genae. Thorax: (Fig. 39) anterior prothorax dark brown, remainder paler; pharate pupal gill as black, paddle–shaped horn; annulations of gill concertinaed. Abdomen: evenly medium brown, darkened posteriorly; abdominal segments expanded smoothly; posteroventral tubercles not markedly developed. Anal papillae: three simple lobes. Posterior proleg (Fig. 40): rectal scales absent; anal sclerite X– shaped with median region poorly expressed, anterior arms slightly flared, shorter than posterior arms, interarm struts distinct, posterior arms short; accessory sclerite absent, pigmented semicircular sclerite absent, but clear cuticle evident in that position. Posterior circlet: ca. 76–80 rows of 11–13 hooks (total ca. 930). Etymology. Named by Davies & Györkös (1988) after Murray H. Colbo. Types. The original type material as designated by Davies & Györkös (1988: 111) was stated to be a holotype pharate female with parts mounted on slides. Paratypes of 5 pupae, a pupal exuviae in alcohol with filaments on slides and 2 mature larvae mounted on slides. Label data as "Halls' Gap (37º 07´S/ 142º 07´E) in a slow flowing drain in a grassy forest, 26.viii.1958. I. M. Mackerras". As partially explained by Craig (2011), the dissected materials were never permanently mounted on slides and, when recovered, were still in depression slides, in glycerine, of which most had crept out of the depression. Alcohol material was dry—not surprising after some 20 years, or so. Much of the described material by Davies & Györkös (loc. cit.) was not recovered. That which was, was badly bleached and essentially useless. Some, however, was stained in Chlorozoal Black and examined to confirm it was of P. colboi, then placed in microvials, pinned, labeled as below and deposited in the Australian National Insect Collection (ANIC), CSIRO, Canberra. TAXONOMIC REVISION OF PARAUSTROSIMULIUM Zootaxa 4337 (4) © 2017 Magnolia Press · 455 FIGURES 1–6. Paraustrosimulium colboi female. (1) Habitus. Reared. ETOH. Arrow indicates lateral cervicales. Scale bar = 0.5 mm. (2) Head. Frontal view. ETOH. Scale bar = 0.2 mm. (3) Antenna. Scale bar = 0.05 mm. (4) Maxilla. Scale bar = 0.05 mm. (5) Mandible. Note teeth on both sides. Scale bar = 0.02 mm. (6) Lacinia. Scale bar = 0.02 mm. 456 · Zootaxa 4337 (4) © 2017 Magnolia Press CRAIG ET AL. FIGURES 7–11. Paraustrosimulium colboi female. (7) Cibarium. (Partly reconstructed). Scale bar = 0.05 mm. (8) Wing veins. Scale bar = 0.2 mm. (9) Wing. Scale bar = 0.5 mm. (10) Metathoracic furcasterum. Scale bar = 0.05 mm. (11) Basitarsus, calcipala & pedisulcus. Scale bar = 0.02 mm. TAXONOMIC REVISION OF PARAUSTROSIMULIUM Zootaxa 4337 (4) © 2017 Magnolia Press · 457 FIGURES 12–17. Paraustrosimulium colboi female. (12) Claw. Scale bar = 0.02 mm. (13) Abdominal tergites. Scale bar = 0.2 mm. (14) Hypogynial valves. Scale bar = 0.1 mm. (15) Genital fork. Scale bar = 0.05 mm. (16) Spermatheca (flattened) Scale bar = 0.05 mm. (17) Cercus & anal lobe. Scale bar = 0.05 mm. 458 · Zootaxa 4337 (4) © 2017 Magnolia Press CRAIG ET AL. FIGURES 18–23. Paraustrosimulium colboi male. (18) Habitus. Reared. ETOH. Scale bar = 0.5 mm. (19) Head, frontal view. ETOH. Scale bar = 0.2 mm. (20) Antenna. Scale bar = 0.05 mm. (21) Claw. Arrow indicates grappling hooks. Scale bar = 0.02 mm. (22) Abdominal tergites. Scale bar = 0.2 mm. (23) Genitalia, ventral view, cleared, in glycerine. Scale bar = 0.1 mm. TAXONOMIC REVISION OF PARAUSTROSIMULIUM Zootaxa 4337 (4) © 2017 Magnolia Press · 459 FIGURES 24–25. Paraustrosimulium colboi male. FIGURES 26–29. Paraustrosimulium colboi pupa. (24) Ventral plate & parameres (slide mounted). Abbreviations: b a—basal arm, m s—median sclerite, p—paramere, s—spines, v p—ventral plate. Scale bar = 0.02 mm. (25) Gonocoxa—gc, gonostylus—gs. Lateral view. Scale bar = 0.05 mm. (26) Pupa and cocoon. Scale bar = 0.1 mm. (27) Pharate gill, slightly expanded. Scale bar = 0.2 mm. (28) Female cephalic capsule. Scale bar = 0.2 mm. (29) Male cephalic capsule. Scale bar = 0.2 mm. 460 · Zootaxa 4337 (4) © 2017 Magnolia Press CRAIG ET AL.