Austrocnephia fuscoflava (Mackerras & Mackerras), 2019

Craig, Douglas A., Currie, Douglas C., Gil-Azevedo, Leonardo H. & Moulton, John K., 2019, Austrocnephia, new genus, for five species of ‘ Paracnephia’ (Diptera: Simuliidae), with a key to Australian black fly genera, Zootaxa 4627 (1), pp. 1-92 : 41-51

Austrocnephia fuscoflava (Mackerras & Mackerras) . New combination.

( Figs. 99–132 )

Cnephia tonnoiri fuscoflava Mackerras & Mackerras 1948: 236 ; original description.

‘ Cnephia of authors’ fuscoflava . Crosskey, 1987: 443.

fuscoflava . Crosskey, 1989: 222; unplaced species of Prosimulinii.

Paracnephia fuscoflava (Mackerras & Mackerras) . Crosskey & Howard, 1997: 18; Prosimuliini , unplaced to subgenus, new combination. Bugledich, 1999: 328. Adler & Crosskey, 2008: 26, transferred to Simulinii. Adler, 2019: 32.

Redescription. Adult female (numerous reared specimens). A smaller species. Body ( Figs. 99, 100 ): head very dark red, thorax tan, abdomen black; total length 2.2–2.4 mm. Head ( Fig. 101 ): overall dark in colour; width 0.80–0.82 mm; depth 0.63–0.70 mm; postocciput markedly hirsute with long pale hairs, frons margins slightly concave, markedly narrowed, dark brown-black, sparse pale hairs; frons:head-width ratio 1.0:8.5–10.1. Eyes: interocular distance 0.08–0.09 mm; upper ommatidia redder than lower ommatidia, diameter 0.019 mm; lower ommatidia lighter in colour with slightly thicker margins, diameter 0.021 mm, ca. 36 rows across and 44 down at mid-eye. Clypeus: width 0.14–0.19 mm, markedly dark brown, densely hirsute. Antenna ( Fig. 102 ): elongate, extended well beyond posterior margin of head; total length 0.63 mm; scape small and pale, pedicel larger and similar in size to flagellomere I, remainder similar in size, flagellomeres bead-shaped, concolourous, slightly tapered to apex. Mouthparts: feebly developed, ca. 0.23× length of head depth; maxillary palp, total length 0.70 mm, palpomeres I & II small, palpomere III darker brown than remainder with dense vestiture, palpomere IV small, V elongated; proportional lengths of palpomeres III–V, 1.0:0.7:1.6; sensory organ markedly elongated, 0.75× length of palpomere III, opening elongated and 0.6× vesicle width; mandible ( Fig. 103 ) with ca. 26 small inner teeth, arrayed in straight formation (see insert), outer teeth absent; lacinia with 13 teeth on either side; cibarium cornuae flared with narrow terminal extensions, medial gap narrowed and concave, ( Fig. 104 ). Thorax: length 1.3–1.5 mm, width 0.9–1.2 mm; scutum evenly tan; postpronotal lobe well developed with fine hair longer than on scutum, that with overall sparse fine small hairs; scutellar depression similar, concolourous, but with denser long pale hairs; scutellum slightly paler than scutum, vestiture of sparse very fine yellowish hairs centrally, darker laterally; postnotum concolourous with scutellum; antepronotal lobe with distinct patch of yellow hairs; proepisternum and fore coxa bare; pleuron and anepisternal membrane yellowish brown, bare; katepisternum dark brown, sulcus shallow and broad. Wing ( Fig. 105 ): length 2.5–2.8 mm; width 1.2–1.5 mm; very slightly fumose overall, veins yellowish, markedly developed pigmentation at junction of r-m cross vein as well as humeral and other basal cross veins; basal medial cell minute to absent; a:b ratio 1.0:2.4; distal 2/3 of costa with spines, dense apically, Rs not divided apically ( Fig. 106 ); M 1 appears doubled; CuA barely sinuous; CuP extended nearly to wing margin, as does A 1. Haltere: stem clear, knob tan. Legs ( Fig. 107 ): forelegs evenly pale; mid and hind legs blackish brown; hind legs slightly darker; hind basitarsus parallel-sided with ventral row of stout spines; calcipala subequal in length and breadth, as wide as tarsomere; pedisulcus not markedly developed, but obvious; tarsomere II 2.0–2.3× longer than distal width; claws, smoothly tapered, not markedly curved, basal tooth well developed and broad, ca. 0.5× as long as talon, with distinct basal notch, heel small ( Fig. 108 ). Abdomen: mottled dark brown overall, paler anteriorly; basal scale dark brown, vestiture of long dense hair extended to segment IV; tergites markedly poorly sclerotized, barely discernable from remainder of dorsum, broader in posterior segments; vestiture better expressed on posterior segments. Genitalia: markedly small; sternite VIII with no distinct medial region, vestiture of microtrichia, large strong hairs posterolaterally; hypogynial valves ( Fig. 109 ), lightly pigmented, vestiture of triads of microtrichia and strong hairs, medial edges of valves slightly diverged—straight to slightly concave, bluntly triangular apically; cercus in lateral view elongated with faint medial depression, anal lobes smaller, with vestiture of long hairs ( Fig. 110 ); genital fork ( Fig. 111 ) well expressed, anterior arm thickened, flattened laterally, no indication of membranous lateral areas, lateral arms broad, no indication of knee-bend, apodeme absent, lateral plates triangular, small; spermatheca irregularly ovoid to spherical ( Fig. 112 ), externally slightly tuberculate producing mottle effect, internal fine spines present, clear membranous area at junction with spermathecal duct small with series of raised cuticular rings.

Adult male (numerous reared specimens, Stradbroke Island). Body: pinned (Fig. 113), head dark, thorax yellowish brown; in alcohol, overall colour darker, head dark red, thorax greyish brown, abdomen black ( Fig 114 ); total body length 1.8–2.0 mm. Head ( Fig. 115 ): width 0.83–0.84 mm; depth 0.83–0.86 mm; relatively large in relation to thorax. Eyes: upper ommatidia dark reddish orange, enlarged, diameter 0.05 mm, ca. 12–15 across and down; lower ommatidia black to blackish orange, markedly smaller, diameter 0.01 mm, ca. 26 across and down. Clypeus: black; width 0.17–0.18 mm; vestiture of fine black hairs. Antenna ( Fig. 116 ): total length 0.59–0.67 mm; overall evenly light brown; scape short, pedicel as long as broad, flagellomere I longer than wide, flagellomeres II–V quadratic in shape, apical three flagellomeres slightly longer than wide. Mouthparts: insubstantial; length 0.2× head depth; maxillary palp 0.55 mm long, palpomeres I & II small, palpomeres III & IV subequal in length, palpomere IV elongated and as long as III & IV combined, proportional lengths of palpomeres III–V 1.0:0.8:2.0, sensory vesicle small, spherical, occupying 0.2× palpomere length, opening 0.33× vesicle width; lacinia small, lacking teeth, but with apical hairs; mandible not observed. Thorax: markedly domed, lateral outline semicircular; length 1.1 mm; width 0.9 mm; postpronotal lobe with longish fine pale hairs, antepronotal lobe with patch of sparse fine pale hairs, proepisternum bare; scutum evenly greyish yellow (tan), vestiture of evenly sparse longish fine pale hairs, longer anteriorly and in scutellar depression; scutellum yellower than scutum with markedly long yellow hairs laterally, postpronotal lobe concolourous with scutum; pleuron greyish yellow, anepisternal membrane bare, katepisternum dark brown, sulcus distinct, but shallow. Wing: length 2.8 – 2.9 mm, width 1.1–1.3 mm; membrane slightly fumose at apex; a:b ratio 1.0:2.6; veins slightly yellowish, costa with well expressed spines, more so distally, r-m cross vein junction darkly pigmented, as are basal humeral and other cross veins (appear as two spots), Rs shows no sign of bifurcation or thickening distally, M 1 thickened and appears slightly doubled, CuA slightly sinuous, CuP extended nearly to wing margin. Haltere: stem pale, knob tan. Legs: overall bicoloured, pale and black, hind leg with tibia darker coloured distally; hind basitarsus margins parallel; ventral row of stout spines, sparse; calcipala well developed, narrower than basitarsus, with deep dorsal notch; pedisulcus not markedly developed; tarsomere II 2.3× as long as apical width; tarsal claw small and finely expressed, with truncated basal tooth and smaller basal projection, grappling pad of ca. 23 teeth (as for other species). Abdomen: black dorsally and posteriorly, pale anteroventrally, vestiture of markedly long golden hairs, more so anteriorly; basal scale not markedly pigmented, hairs golden, extended to posterior of tergite II; tergites not markedly sclerotized, increased in width posteriorly and more fully pigmented; pleural cuticle concertinaed; sternites moderately developed, sternite II absent. Genitalia ( Fig. 117 ): small, not heavily pigmented; cerci well developed and distinct; gonocoxa, in ventral view, 2.3× longer than basal width, strengthened posteromedially, extended beyond junction with gonostylus, vestiture of long black sparse hairs, microtrichia, posteromedial margin scalloped; gonostylus short, in ventral view curved, in lateral view cone shaped ( Fig. 118 ), approximately 1.5× longer than basal width, strengthened along inner edge, two insubstantial short, blunt, apical spines; ventral plate ( Fig. 119 ) small, 2.0× wider than long, posteromedial apex directed slightly ventrally, appearing as shallow medial concavity, curved laterally, slightly convex anteromedially, vestiture of sparse coarse hairs medially, basal arms finely expressed, paramere connection not markedly expressed; median sclerite poorly expressed—two widely separated short sclerotized arms projected ventrally; parameres plate-like, lightly expressed, slightly crenulated along inner edge, spines absent; aedeagal membrane adorned with microtrichia.

Pupa (based on numerous specimens). Body: female length 2.5–3.0 mm, male length 2.6–2.8 mm; clear yellow cuticle overall. Head: cephalic plates smooth, frons of female bluntly rounded with ratios of basal width to vertex width, and height 1.0:1.3 and 1.0:1.5 respectively ( Fig. 121 ), male ovoid; ratios 1.0:1.7 and 1.0:2.7 respectively ( Fig. 122 ); cuticle lacks pattern; in male, frontal and facial setae present, the former small and fine, the latter substantial and curved, in female facial setae present, frontal setae varied —either present on extreme lateral edge, or absent but on epicranial area; epicranial setae present, closely applied beside antennal sheath; antenna sheath of female extended beyond margin of ocular shield, not so in male. Thorax: notum markedly domed, smooth, but with faint pattern, dorsal setae short, substantial, curled at tip. Gill ( Fig 123 ): total length ca. 0.75 mm; 15–20 brownish grey filaments arising from three or four short basal trunks, paler apically, arranged in antler-like fashion directed anteriorly; filament surface reticulated basally, pseudoannulated distally. Abdomen ( Fig. 124 ): overall, cuticle clear and well sclerotized, with distinct pale yellow sclerites, markedly so for anterior tergites; tergites III & IV connect to the respective sternites with wrinkled cuticle, no pleurites; but pleurites present and well expressed on segments V–VII, pleurite V larger and connected to both tergite and sternite, those of VI & VII smaller and isolated; sternites VI & VII separated medially; abdomen with minute rounded tubercles to absent. Abdominal armature moderately well developed; tergite I with fine hairs; tergite II with 4+4 poorly developed anteriorly directed hooks; tergites III & IV each with 4+4 central anteriorly directed long thin recurved hooks (similar to the ventral hooks), tergite IV with 1+1 lateral hooks, smaller than the more central hooks; tergites V–VIII each with spine comb anteriorly and posterior double pair of fine hairs on each side; segment IX with sharply tapered and slightly curved terminal spines, with other terminal setae simple, long and curved; sternite IV & V with 3+3 (2+2 stronger central and 1+1 poorly developed lateral); sternite VI with 2+2; sternite VII with 1+1 anteriorly directed simple recurved hooks; pleurites VI & VII with 1+1 anteriorly directed, poorly expressed, simple, recurved hooks.

Cocoon. Close fitting, may cover complete pupa, but variable in shape; irregular weave, silk fibers fine, dark brown; considerable extraneous material (algal filaments, sand, etc.) incorporated ( Fig. 120 ).

Larva (based on numerous last instar larvae). Body ( Fig. 125 ): total length 6.5–7.3 mm, evenly light grey and brown, markedly expanded posteroventrally. Head ( Fig. 126 ): mottled medium brown, head spot pattern poorly expressed, but tending to positive; length 0.74–0.93 mm, maximum width 0.61–0.74 mm; distance between antennal bases 0.36–0.38 mm; apotome broad, ecdysial lines distinct and divergent until posterior of stemmata, cervical sclerites distinct, separate from postocciput and situated medially. Antenna: apex not extended to end of labral fan stem; total length 0.27–0.32 mm, overall brown; basal article short and clear proximally, darker brown distally, medial article and distal articles darker brown than remainder, basal article markedly shorter than medial article, proportional lengths of basal, medial, and apical articles 1.0:1.4:2.0, apical article relatively longer than others of the genus. Labral fan: stem short and broad, pigmentation ranging from pale to light brown, ca. 36–38 dark brown fine rays, ten posterior rays finer than others, length 0.8 mm, mid-ray width 0.01 mm; no microtrichial pattern as such—all of similar length (0.02 mm) and markedly longer than ray width. Mandible ( Fig. 127 ): well pigmented; brushes typical; outer teeth markedly short; apical tooth moderately developed; subapical teeth small and subequal in length, blunt; ca. 7 spinous teeth, first subequal in length to apical tooth, others slightly decreased in length proximally; serrations and sensillum well expressed; blade region slightly convex. Maxilla: heavily pigmented; palp cone-shaped, 2.6× as long as basal width; hair tuft at base of palp not markedly developed. Postgenal cleft ( Fig. 128 ): shallowly V-shaped, with irregular anterior edge; postgenal bridge concolourous with genae. Hypostoma ( Fig. 129 ): darkly pigmented; tooth 0 not prominent, but distinct, subequal to tooth 3, teeth 1 & 2 shorter and less prominent, tooth 4 longer and flanged medially, teeth 5–7 markedly small and not obvious, tooth 8 more basal, small and directed laterally; all teeth as a unit with parallel sides; serrations lightly expressed on convex lateral edge of hypostoma; four to six hypostomal setae on each side, anterior ones more substantial; ratio of hypostoma: genal bridge: postgenal cleft 1.0:1.5:0.7. Thorax ( Fig. 130 ): gill histoblast broadly rounded with four or five substantial dark trunks visible basally, all directed ventrally then broadly posteriorly, then curled anteroventrally, with pale filament endings then turned dorsally along anterior edge of histoblast. Prothoracic proleg: well developed and elongated with sharply V-shaped lateral plates; lappets absent. Abdomen: evenly expanded from anterior to posterior segments, but markedly expanded posteroventrally at segments VII & VIII. Rectal papillae: three simple lobes, well developed. Ventral tubercles: absent. Anal sclerite ( Fig. 131 ): anterior arms flared, strengthened laterally; main body of sclerite not markedly developed, posteromedial clear area open, posterolateral arms absent, posteroventral arms finely expressed and tapered, numerous campaniform sensillae anterolateral of posteroventral arm; seven or eight campaniform sensilla between the posteroventral arms and circlet of hooks; posteromedial extensions into circlet of hooks junction absent. Posterior circlet: well developed and directed posteriorly, ca. 130 row of hooks with 13–15 hooks per row (total ca. 1,820).

Eggs. Laid in mats on trailing grass. Typical triangular shape.

Etymology. Not expressly given by Mackerras & Mackerras (1948: 236), but no doubt in reference to the tan colour (dark blonde) of the adult scutum.

Types. Holotype. The female holotype ( ANIC) was examined and photographed by LHG-A in 2007 ( Fig. 99 ). Exact labeling not recorded. The type locality ( Mackerras & Mackerras, 1948: 236) is merely given as “Dunwich, Stradbroke Island”; the original collection was by E. N. Marks, 28 Nov. 1947. Stradbroke Island is also now known as ‘ Minjerribah’.

Paratypes. Bugledich (1999: 328) listed 10 male and nine female paratypes in the ANIC with two male and two female specimens in the Natural History Museum, London. There were 16 ANIC specimens available for our examination, including six paratypes:- three males [Dunwich/ em. 12 Oct. 47] [ Cnephia / tonnoiri fuscoflava / M&M/ PARATYPE. {male symbol}] [Aust. Nat./ Ins. Coll.] [ANIC Database No./ 29 031821 –823]. The first numbered is now mounted on a microscope slide. Three females with data as for males, but with female symbol and [ANIC Database No./ 29 031824 –03826]. The first is now mounted on a slide.

Additional material examined. Three pinned females [Dunwich, Q./ 11. 8. 51 / {female symbol}] [Aust. Nat./ Ins. Coll.]. A further eighteen ( ANIC) slide mounts of all stages were examined, plus topotype material (Queensland Museum) and considerable newer material collected in 1996 by JKM, and that recently in 2015, collected by DCC and JKM. Alcohol material: All stages [ ANIC Database No./ 29 026741–026744]; [ UASM #/370832–370834]. Slide mounts: All stages [ UASM #/ 370835–370846].

Bionomics. In the original description, Mackerras & Mackerras (1948: 236) noted that eggs were unknown. We have examined, however, a sample collected in 1951: the eggs are laid as a mat on grass leaves and are of normal triangular shape. Mackerras & Mackerras (1948: 239) further noted that larvae of all ages were abundant in clear, moderately fast, somewhat brown water flowing in a narrow man-made channel, plus in a small, sunlit, natural stream just before it discharged onto a beach, both streams with sand substrate. Larvae were attached to vegetation, submerged sticks and logs, and especially to barely-submerged long grass blades. Pupae were in similar situations as the larvae and covered by irregular cocoons with extraneous material incorporated (e.g., Fig. 120 ). This is a notable feature for pupae of this species. No adults were originally collected from the type locality and there were no records of any biting. Given that Stradbroke Island is a tourist location and biting, as far as known, not been reported, the species may be autogenous, or highly ornithophilic as are most Cnephia -grade taxa.

More recent collections generally corroborate the original data. Of interest is that much of the newer material was collected from fast water ( Fig. 132 ). Given collections from Stradbroke Island contain a full array of stages over an extended period of months, the probability that A. fuscoflava is multivoltine is high.

Distribution ( Fig. 197 ). Austrocnephia fuscoflava was originally known only from Stradbroke Island (aka Minjerribah) near Brisbane ( Mackerras & Mackerras, 1948: 236). No other simuliid species has been reported from the island. Bugledich (1999: 328) listed distribution of A. fuscoflava as widespread, from the NE coastal, SE coastal, NSW, QLD, TAS, VIC, but no such material is in the ANIC (DAC pers. obs., 2014). We assume that this given distribution is in error. Still, a single pupa collected by H. Zwick from Cement Creek, Victoria, was considered possibly that of A. fuscoflava . We have re-examined the specimen, but could not make a firm identification. Mackerras & Mackerras (1949: 384) gave Little Nerang River as a locality, but later ( Mackerras & Mackerras, 1950: 170) retracted that. Moulton, however, collected A. fuscoflava material in 2002 from the mainland north of Brisbane, at Cooloola Creek, Rainbow Beach Road (S26.0050º E153.0590º), so the species is likely to occur elsewhere, but not as widespread as indicated by Bugledich (loc. cit.).

The exact position of the type locality is unknown. Moulton’s 1996 Stradbroke material and that recently in 2015, collected by DCC and JKM was from Capembah Creek, Myora Springs Environmental Park (S27.4687º E153.4257º) ( Fig. 132 ). The stream is generally with sandy substrate and of low gradient, but near the road there was a fast stretch and small waterfall that yielded specimens.

Remarks. Number of hooks (ca. 1,800) in the posterior circlet of the Stradbroke Island material is unusually low for the genus and in keeping with the lower velocity inferred for habitats of the original collections—streams with sand substrates. As noted previously, this agrees with the correlation detailed by Palmer & Craig (2000) and others, regarding number of hooks and velocity of flow. That, as also noted elsewhere, is at variance with the abdominal shape—adapted for fast water.

An interesting point regarding slides prepared by the Mackerras’s of the October 1947 material, is that some slides, originally labeled as Prosimulium sp., has that crossed out and relabeled Cnephia tonnoiri fuscoflava .

Figures

FIGURE 105. Austrocnephia fuscoflava female. (105) Wing. Paratype. Scale bar = 0.5 mm.

FIGURE 124. Austrocnephia fuscoflava pupa. (124) Abdominal armature. Arrow indicates pleurite (plt). Scale bar = 0.5 mm.

FIGURES 132. Austrocnephia fuscoflava larva. (132) Typical habitat of A. fuscoflava immatures. Capembah Stream, Strad- broke Island. Image JKM. FIGURES 133–135. Austrocnephia orientalis female. (133) Holotype, frontal view. Image by LHG- A. Scale bar = 0.25 mm. (134) Head. (Hume Highway). Scale bar = 0.25 mm. (135) Mandible (mnd) and lacinia (lc). (Hume Highway). Scale bar = 0.05 mm.

FIGURE 197. Distribution of Austrocnephia tonnoiri species-group (A. fuscoflava, A. orientalis, A. tonnoiri).

FIGURES 99–102. Austrocnephia fuscoflava female. (99) Holotype, frontal view. (Stradbroke Island). Image by LHG-A. Scale bar = 0.5 mm. (100) Habitus. (Stradbroke Island). Arrow indicates wing spot (w s). Scale bar = 0.5 mm. (101) Frontal view of head. Paratype. Cleared. (Stradbroke Island). Scale bar = 0.2 mm. (102) Antenna. Paratype. Scale bar = 0.2 mm.

FIGURES 103, 104. Austrocnephia fuscoflava female. (103) Maxillary palp, lacinia and mandible. Insert—mandible apex. Paratype. Scale bar = 0.1 mm. (104) Cibarium. Scale bar = 0.1 mm.

FIGURES 106, 107. Austrocnephia fuscoflava female. (106) Wing veins showing non-branched Rs. Scale bar = 0.1 mm. (107) Calcipala and pedisulcus. Paratype. Scale bar = 0.1 mm.

FIGURES 108–112. Austrocnephia fuscoflava female paratype. (108) Tarsal claws. Scale bar = 0.02 mm. (109) Hypogynial valves. Scale bar = 0.05 mm. (110) Anal lobe (a l) and cercus (c). Scale bar = 0.05 mm. (111) Genital fork. Scale bar = 0.05 mm. (112) Spermatheca. Scale bar = 0.05 mm. FIGURE 113. Austrocnephia fuscoflava male paratype. (113) Dorsal view. Image by LHG-A. Scale bar = 0.5 mm.

FIGURES 114, 115. Austrocnephia fuscoflava male. (114) Habitus. Scale bar = 1.0 mm. (115) Frontal view of head. Scale bar = 0.2 mm.

FIGURES 116–119. Austrocnephia fuscoflava male. (116) Antenna. Scale bar = 0.1 mm. (117) Genitalia, ventral view. Scale bar = 0.05 mm. (118) Gonocoxa (gc) and gonostylus (gs). Scale bar = 0.05 mm. (119) Genitalia, slide mounted, showing aedea- gal membrane (a m), paramere connector (p c), paramere (p) and ventral plate (v p). Scale bar = 0.05 mm.

FIGURES 120–123. Austrocnephia fuscoflava pupa. (120) Habitus. Scale bar = 1.0 mm. (121) Female cephalic capsule. Scale bar = 0.2 mm. (122) Male cephalic capsule. Scale bar = 0.2 mm. (123) Gill. Tips of some filaments broken. Scale bar = 0.2 mm.

FIGURES 125–127. Austrocnephia fuscoflava larva. (125) Habitus, last instar. Scale bar = 1.0 mm. (126) Head, dorsal view, last instar. Scale bar = 0.5 mm. (127) Mandible apex showing subapical (sb t) and spinous (sp t) teeth. Scale bar = 0.05 mm.

FIGURES 128, 129. Austrocnephia fuscoflava larva. (128) Ventral head cuticle. Scale bar = 0.1 mm. (129) Hypostoma, note lateral serrations (l s). Scale bar = 0.1 mm.

FIGURES 130, 131. Austrocnephia fuscoflava larva. (130) Pupal gill histoblast. Scale bar = 0.2 mm. (131) Anal sclerite showing anterior arms (a a), campaniform sensilla (c s), posteroventral arms (pv a) and circlet of hooks. Scale bar = 0.2 mm.