Austrocnephia aurantiaca ( Tonnoir 1925 ), 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 : 8-27

Austrocnephia aurantiaca ( Tonnoir 1925) . New combination.

( Figs. 1 –57)

Simulium aurantiacum Tonnoir 1925: 234 ; original designation.

Simulium (Cnephia) aurantiacum . Edwards, 1931: 147.

Cnephia aurantiacum. Smart, 1945: 498 . Mackerras & Mackerras, 1948: 238. 1949: 383.

Cnephia A. Rothfels, 1979: 522 ; cytological study, perhaps aurantiacum ?

‘ Cnephia of authors’ aurantiacum . Crosskey, 1987: 443; undetermined genus, Prosimuliini .

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

Paracnephia aurantiaca . Crosskey & Howard, 1997: 18; new combination.

Paracnephia aurantiaca . Bugledich, 1999: 327.

‘ Cnephia ’ aurantiacum . Moulton, 2000: 98. Moulton, 2003: 47.

Paracnephia aurantiaca . Crosskey & Howard, 2004: 10; unplaced to subgenus.

Paracnephia aurantiaca . Adler & Crosskey, 2008: 28; transferred to Simulinii, unplaced to subgenus. Hernández-Triana et al. 2017: 350. Adler, 2019: 32.

Redescription. Adult female (based on three paratypes and numerous other specimen). Body ( Figs. 1, 2 ): head dark orange, thorax yellowish orange, abdomen blackish; total length 3.2–5.0 mm. Head ( Fig. 3 ): overall dark in colour; width 0.96 mm; depth 0.70 mm; postocciput densely hirsute with long pale hairs, frons parallel-sided, not markedly narrow, dark brown-black, sparse long black hairs laterally; frons:head-width ratio 1.0:12.0. Eye: interocular distance ca. 0.1 mm; upper ommatidia reddish orange, lower ommatidia darker, ommatidia diameter 0.022 mm; ca. 36 rows across and 44 down at mid-eye. Clypeus: width 0.29 mm; mottled dark brown, vestiture of long hairs laterally. Antenna ( Figs. 3, 4 ): elongate, extended well beyond posterior margin of head; total length 0.95 mm; scape and pedicel pale, similar in size to flagellomere I; flagellomere II–VII similar in size and shape, flagellomeres VIII & IX slightly more elongated. Mouthparts: feebly developed, ca. 0.28× length of head depth; maxillary palp ( Fig. 5 ) total length 0.83 mm, palpomeres I & II small, palpomere III broader and darker than other palpomeres, proportional lengths of palpomeres III–V 1.0:0.8:1.6, respectively; sensory organ moderately elongated, 0.33–0.50× length of palpomere III, opening small, 0.2× vesicle width; mandible ( Fig. 6 ) with ca. 30 weakly developed inner teeth, outer teeth absent; lacinia with 16 and 10 teeth on inner and outer edge respectively; cibarium ( Fig. 7 ) with cornuae sclerotized apically and lightly sculpted, medial gap angulate. Thorax: length 1.2–1.8 mm; width 1.2–1.3 mm; evenly yellowish orange; postpronotal lobe well developed with longer hair than on scutum; scutellar depression with similar vestiture; scutellum slightly paler than scutum, vestiture of sparse very fine yellowish hairs; postnotum concolourous with scutellum, vestiture similar; antepronotal lobe with dense patch of yellow hairs; proepisternum and fore coxa with sparse hairs; anepisternal (aka plural) membrane yellowish brown, bare; katepisternal dark brown, sulcus shallow and broad. Wing ( Figs. 8 , 9 ): length 4.1–5.0 mm; width 1.6–2.2 mm; membrane slightly fumose on apex and anal lobe; veins dark orange; costa with mixture of thin and thickened hairs on distal 2/3rds, the latter not markedly darkened; Rs narrowly divided distally (R 2+3 occasionally expressed as row of a few fine hairs); a:b ratio 1.0:2.5; r-m cross vein darker than adjacent veins, but pigmentation not extended into surrounding membrane; basal medial cell minute or absent; M 1 appearing doubled or tripled; CuA not markedly sinuous; CuP extended nearly to wing margin, as does A 1. Haltere: stem clear, knob yellowish orange. Legs ( Fig. 10 ): forelegs evenly pale; mid and hind legs medium brown; hind basitarsus with ventral row of regular stout spines; calcipala subequal in length and breadth; pedisulcus absent; tarsomere II 2.0–2.4× as long as distal width; claw ( Fig. 11 ) with main talon strongly curved and evenly tapered, basal tooth 0.25–0.50× length of talon, heel substantial and angulate. Abdomen ( Fig. 12 ): basal scale (tergite I) dark brown, vestiture of long dense hairs; anterior few segments pale and yellowish, other segments mottled dark brown; tergites lightly sclerotized, barely discernable from remainder of dorsum, broader in posterior segments; vestiture better expressed on posterior segments. Genitalia: markedly small; sternite VIII with distinct microtrichial array medially, with larger stronger hairs posterolaterally; hypogynial valves ( Fig. 13 ), lightly pigmented, vestiture of triads of microtrichia and strong hairs, medial edges of valves slightly concave, but not touching, broadly rounded apically, valves slightly domed medially; cercus in lateral view ( Fig. 14 ) elongated with slight medial depression, anal lobe also elongate, both with vestiture of long hairs; genital fork ( Fig. 15 ) with anterior stem relatively long and narrowed, slightly expanded apically, no indication of membranous lateral areas, lateral arms narrow, apodeme as membranous triangular region, lateral plates small, trapezoidal- or subtriangularshaped; spermatheca ovoid ( Fig. 16 ), markedly wrinkled, without internal spines, membranous area at junction with spermathecal duct small.

Adult male (based on a paratype and numerous other specimens). Body: overall yellow including head and thorax, abdomen black (Figs. 17, 18); total body length variable, specimens from Pretty Valley, VIC. 3.3–4.5 mm; those originally described by Tonnoir (1925: 234) 2.0– 2.5mm. Head ( Fig. 19 ): width 0.98 mm; depth 0.7 mm. Eyes: upper ommatidia dark orange, not markedly enlarged, diameter 0.04 mm, ca. 23 across and down; lower ommatidia black to blackish orange, markedly smaller, diameter 0.02 mm, ca 24 across, 38 down. Clypeus: black; width 0.19 mm; vestiture of sparse fine black hairs. Antenna ( Fig. 20 ): total length 0.71–0.75 mm; pedicel and scape dark brown; scape short, pedicel wider than broad, flagellomere I narrower than scape and longer than broad, pale basally, other flagellomeres evenly medium brown and tapered slightly to apical flagellomere. Mouthparts: insubstantial; length 0.3× head depth; maxillary palp ( Fig. 21 ) 0.68 mm long, palpomeres I & II small, palpomeres III & IV subequal in length, palpomere V fine and elongated, proportional lengths of palpomeres III–V 1.0:0.8:1.7 respectively, sensory vesicle small, spherical, occupying 0.22× or less palpomere length, opening 0.33× vesicle width; lacinia small, lacking teeth, but with apical hairs; mandible lacking teeth. Thorax: length 1.4 mm; width 1.3 mm; markedly domed, lateral outline semicircular, head angled anteriorly; postpronotal lobe concolourous with scutum, bearing long fine pale hairs; antepronotal lobe with patch of sparse fine pale hairs; proepisternum bare; scutum evenly pale yellow, vestiture of sparse long fine pale hairs, longer in scutellar depression; scutellum lighter than scutum, with markedly long yellow hairs laterally; pleurae yellow, anepisternal membrane bare; katepisternum brown, sulcus distinct, but shallow. Wing: length 3.7–4.3 mm, width 1.7–2.3 mm; membrane slightly fumose at apex, anal lobe very slightly so; anterior veins dark orange; costa with mixture of thin and thickened hairs; Rs narrowly but distinctly branched distally ( Fig. 22 ); a:b ratio 1.0:2.6; basal medial cell present, but minute; M 1 appearing doubled or tripled, CuA not markedly sinuous; CuP extended nearly to wing margin. Haltere: stem pale, knob tan. Legs: overall yellowish, hind leg with darker coloured distal tibia; hind basitarsus, slightly expanded medially with ventral row of stout spines, calcipala and pedisulcus as for female; tarsomere II 2.6× as long as distal width; tarsal claw with short talon, grappling hook of 25–27 teeth ( Fig. 23 ). Abdomen ( Fig. 24 ): black dorsally and posteriorly, pale anteroventrally, vestiture of markedly long golden hairs, more so anteriorly; basal scale (tergite I) black, hairs markedly golden and long, extended to posterior of tergite II; tergite II V-shaped, remainder broad; sternite I present, sternite II absent, sternites III–VIII rounded, distinctly hirsute. Genitalia ( Fig. 25 ): small and lightly sclerotized; cerci well developed; gonocoxa 1.1× longer than its basal width, vestiture of long black sparse hairs and microtrichia; gonostylus 1.6× longer than basal width, markedly shorter than gonocoxite, narrow in ventral view, broad in lateral view ( Fig. 26 ), with two or three apical spines; ventral plate small, 1.6× wider than long in ventral view, apex directed ventrally producing appearance of medial concavity, vestiture absent except for a few hairs medially ( Fig. 27 ), basal arms elongated and narrow; median sclerite in the form of two short, darkly sclerotized, arms arising from anterior edge of ventral plate; parameres plate-like basally, tapered distally; adeagal membrane with 7–15 short stout spines near apex of paramere, membrane otherwise adorned with microtrichia.

Pupa (based on numerous specimens). Body: female length 3.3–4.0 mm, male length 3.3–3.8 mm ( Fig. 28 ). Head: frons of female truncated with ratio of basal width to vertex width 1.0:1.5, and basal width to length 1.0:1.7 ( Fig. 29 ), that of male more ovoid ( Fig. 30 ), ratios 1.0:1.9 and 1.0:3.1 respectively; cuticle faintly patterned; frontal and facial setae present, epicranial setae present near antennal sheath; postorbital spine absent; antennal sheath of female with apex extended beyond margin of ocular shield, that of male not extended to margin of ocular shield. Thorax: markedly domed, smooth, with dorsal setae trichoid, short, curved, but not curled at tip ( Fig. 31 ). Gill: total length ca. 1.3–1.5 mm, with 40–50 light brown filaments arising from 5 or 6 short trunks ( Fig. 32 ); filaments paler apically, some branched at irregular distances from base; ventral filaments directed anteriorly, dorsal filaments directed mainly dorsally, but with one or two longer filaments reflexed posteriorly over thorax; filament surface pseudoannulated throughout ( Fig. 33 ). Abdomen ( Fig. 35 ): well sclerotized, especially the anterior tergites, with minute rounded tubercules; tergites III & IV extended more ventrad than other tergites, separated from pleurite by narrow band of striate membrane; tergite V separated from pleurite by band of striate membrane; pleurites absent from segments V & VI; sternites VI & VII divided medially by a semimembranous longitudinal band, sternite VIII a single lightly sclerotised plate with medial marking. Abdominal armature moderately well developed; tergite I with fine hairs; tergites II–IV each with 4+4 dorsal and 3+3 lateral anteriorly directed recurved hooks, the latter smaller than dorsal hooks; tergite V with four hairs and poorly expressed spine comb; tergites VI & VII each with a welldeveloped transverse row of spine comb, plus a double pair of fine hairs anteriorly and posteriorly on either side; tergite VIII with a small spine comb and a pair of fine hairs posteriorly on either side; segment IX without spine comb, tapered posteriorly, with slightly curved terminal spines, other setae simple, long and curved; sternite III with 2+2 anteriorly directed recurved hooks; sternite IV with 5+5 anteriorly directed recurved hooks; sternite V with 6+6 anteriorly directed recurved hooks, sternite VI with 2+2 anteriorly directed recurved hooks, sternite VII with 1+1 anteriorly directed recurved hooks, sternite VIII with a single hair on either side.

Cocoon. A closely fitting shapeless sac covering abdomen and variable proportions of the thorax, head and gills; silk fibers thick ( Fig. 34 ), some markedly so, often with extraneous material incorporated.

Larva (based on numerous last instar larvae, multiple localities). Body ( Figs. 36, 37 ): total length various, mainland populations 8.6–10.1 mm, Tasmanian populations 5.8–7.1 mm; evenly expanded from anterior to posterior abdomen; colour varied, from mottled brown, darker posteriorly, to evenly yellowish brown, or greenish, to pale (particularly for Tasmanian populations). Head ( Fig. 38 ): evenly dark brown to light mottled brown, apotome slightly lighter than genae; head spot pattern weakly developed, tending to negative; length 0.87–1.2 mm, maximum width 0.71–0.78 mm; distance between antennal bases 0.44–0.48 mm; head widest just posterior to stemmata, narrowed irregularly anteriorly, convex posteriorly; cervical sclerites variously developed, fused to postocciput. Antenna ( Fig. 39 ): short, apex not extended to end of labral fan stem; total length 0.33–0.36 mm, basal article unpigmented proximally, darker brown distally, medial article and distal articles evenly brown, basal article markedly shorter than medial article, proportional lengths of basal, medial and apical articles 1.0:1.5:1.2; medial article slightly expanded distally. Labral fan: stem short and broad, not markedly pigmented, unpigmented in early last instar larvae and in some last instar larvae, ca. 50 dark brown substantial rays, 10 posterior rays finer than others, length 1.0 mm, midray width 0.01 mm; no conspicuous microtrichial pattern—most of similar length with longer ones (0.02 mm) at irregular intervals. Mandible ( Figs. 40, 41 ): short and darkly pigmented; brushes well expressed; outer teeth short; apical tooth moderately developed; subapical teeth small and subequal in length; ca. six spinous teeth, the two distal teeth markedly expressed; serration various, not markedly developed; two sensilla present on slightly convex base; blade region short and convex. Maxilla ( Fig. 42 ): heavily pigmented; palp cone-shaped, 1.9× as long as basal width; hair tuft at base of palp markedly developed. Hypostoma ( Figs. 44, 45 , 46, 47 ): darkly pigmented; tooth 0 with apex extended anteriorly to about the same level as that of tooth 4, giving anterior margin of hypostoma a poorly expressed trilobed appearance; tooth 1 closely associated with tooth 0; teeth 2 and 3 shorter and less prominent; tooth 4 longer and flanged; teeth 5–8 decreasing in prominence laterad of tooth 4; lateral serrations typically absent, rarely present in some populations; 4–6 substantial hypostomal setae on each side; ratio of hypostoma: postgenal bridge: postgenal cleft 1.0:1.6:0.7. Postgenal cleft ( Fig. 43 ): shallow, conspicuously wider than deep, with irregular anterior margin; posterior tentorial pits small, rectangular; postgenal bridge lightly pigmented, contrasting with the darker-brown genae. Thorax ( Fig. 48 ): yellowish; gill histoblast with 4–6 primary trunks visible, anterior one thickened with bifurcations showing, all directed ventrally then posteriorly to produce thick L-shaped lobe, filament tips paler and directed anterodorsally. Prothoracic proleg: well developed, hooks of two sizes, lateral plate with vertical portion well-developed and darkly sclerotized, giving plate an overall L-shaped appearance ( Fig. 49 ); lappets typically absent, if present then shorter than apical article of proleg ( Figs. 49 , 50, 51 ). Abdomen: evenly expanded from anterior to posterior segments, not markedly expanded at segments VII & VIII; paler anteriorly, darker posteriorly; colour ranging from banded mottled yellowish brown, to yellow, or grey. Rectal papillae: three simple lobes. Ventral tubercles: absent. Anal sclerite ( Figs. 52, 53, 54 ): anterior arms markedly flared, relatively lightly sclerotized; main body of sclerite with conspicuous hole posteromedially; posterolateral arms usually absent, but slightly expressed in some populations (e.g., Fig. 54 ); posteroventral arms narrower and more heavily sclerotized than anterior arms, of varied lengths; faintly expressed posteriorly directed extensions projecting from proximal base of each arm into the dorsal junction of hooks in circlet; numerous sensilla situated in membrane immediately anterior to arms (e.g., Fig. 53 ); membranous area between posteroventral arms and circlet of hooks with 12–14 campaniform organs (often as short stout trichoid setae). Posterior circlet: markedly developed and directed slightly ventrally, ca. 260 rows of hooks with 30–33 hooks per row (total ca. 8,320 hooks).

Etymology. Not given by Tonnoir, but no doubt in reference to the yellowish- orange adults.

Types. Holotype. Pinned female. NSW, Blue Mountains, Mount Wilson, S38.5000° E150.3700°. 19 Nov. 1921. Coll. Tonnoir. ANIC. Not examined. Exact labeling not recorded. Bugledich (1999: 327) listed the holotype plus 12 female paratypes in the ANIC, with two male paratypes in The British Museum (Natural History).

Paratypes. One was examined and photographed by LHG-A in 2007 ( Fig. 1 ). Only four were found in ANIC by DAC (pers. obs.) in 2011. Two others are known ( Daniels, 1978:416) from the Australian National Museum (Sydney), but were not examined:- Pinned female, label data:- [K.51170– Cnephia aurantiacum – Australia, New South Wales, Blue Mountains, Mount Wilson, (33° 30’ 4” S, 150° 23’ 41” E), 19 Nov. 1921, Paratype]. Pinned male, [K.51171– Cnephia aurantiacum – Australia, Victoria, Sassafras, (41° 17’ S, 146° 30’ E), 22 Oct. 1922, Paratype]. The following in ANIC: adults elbow-pinned (as for MacGillivray, 1903). Pinned male, [Tasm./Cradle Val./ 19 Jan. 1923 / A. Tonnoir] [(blue) PARATYPE / (handwritten – Simulium / aurantiacum)/ A. Tonnoir det.] [(green) Aust. Nat./ Ins. Coll.] [Examined by /D. A Craig/2013] [male symbol]. Condition—antennae absent, right wing with costa broken at half length, left fore- and mid-legs absent, other legs lacking tarsal claws and distal tarsomeres, otherwise in good condition. (S41.6400º E145.9299º). Pinned female, label as for male plus [Examined by /D. A. Craig/2013] {♀}. Condition—head complete, apical half of left wing missing, left hind leg absent, right hind leg missing tarsus, right hind leg absent, overall good condition. Pinned female [Victoria / Sassafras/ 22 Oct. 1922 / A. Tonnoir.], [(blue) PARATYPE / (handwritten—Simulium/ aurantiacum)/ A. Tonnoir det.] [(green) Aust. Nat./ Ins. Coll.] [Examined by /D. A. Craig/2013] {♀}. Condition—poor, all extremities absent. Female, [Victoria / Sassafras/ 22 Oct. 1922 / A. Tonnoir.] [(blue) PARATYPE / (handwritten—Simulium/ aurantiacum)/ A. Tonnoir det.] [(green) Aust. Nat./ Ins. Coll.] [Examined by /D. A. Craig/2013] {♀}. Condition—good, left antenna and middle legs absent.

Additional material examined. Two reared pinned adults with pupal exuviae in microvial—label data:- [ Austrocnephia aurantiaca ] [ AUSTRALIA: Vic/ unnamed stream/ ex. M31 (Hume Fwy/ b/w Boho & Benalla/ 23 IX 1996 / {M}/ Coll. J.K. Moulton] [ ANIC Database No./ 29 026637] and {F} [ ANIC Database No./ 29 026636]. Seven pinned adults (two males, five female) with pupal exuviae in microvials—label data:- [ Austrocnephia aurantiaca ] [ AUSTRALIA: VIC/ Steavenson Falls @/ Marysville/ S 31º 31.4’ / E 145º 46.6’ / 21 September 1996 / Coll. J.K. Moulton] [ ANIC Database No./ 29 026638–026644]. Alcohol material: Considerable material, largely collected by H. & P. Zwick, mainly of larvae and pupae [ ANIC Database No. / 29 026540–026557; 29 026745– 026843], [ UASM #/ 352049, 352050, 352325, 353462, 353471–353475, 354121, 354123]. Slide mounts: All stages [ UASM #/353349–3534621, 370732–370748, 370831].

Bionomics. Tonnoir (1925: 236) noted that A. aurantiaca females had never been recorded as biting humans, even when occurring in considerable numbers. Mackerras & Mackerras (1949: 384) made the same observation. This is in agreement with the markedly small teeth on the mandible, but is at variance with distinct teeth on the lacinia and size of the CO 2 sensory vesicle (Lutz’s Organ) on the maxillary palp ( McIver, 1987; Sutcliffe et al., 1987). A similar suite of character states is discussed by Craig et al. (2012) in regard to non-biting behaviour of Austrosimulium females in New Zealand. Given that A. aurantiaca females possess a distinct basal tooth on the claw ( Fig. 11 ), perhaps they are strictly ornithophilic?

Label data from the extensive Zwick collection indicates that for higher altitude localities in NSW, water temperatures ranged from 8.0°C to 20.0°C, between July to January. For VIC, temperatures were from 6.0°C to 16.0°C, between May to February; in Tasmania, 11.5°C to 17.5°C during December. For localities known at the time, Mackerras & Mackerras (1949: 384) give dates of collection from October to February.

The majority of habitats comprise clear, fast-flowing streams and rivers (Fig. 57). Tonnoir (1925: 236) provides details of habitats visited by him. Mackerras & Mackerras (1949: 384) similarly note that A. aurantiaca is mainly associated with clear mountain streams. Using unpublished notes of Tonnoir’s they reported that he considered it to be an uncommon species, with larvae found in isolation and pupae in clusters, typically on moss and less often on stones. This is in agreement with habitat details included with the Zwick collections of this species. More recent collections, however, show that larvae can be found clustered in large numbers. Velocity determinations were made during the current investigation using the stand-pipe technique, also known as the head-tube technique (Craig, 1987b: 178. Ciborowski, 1991: 110). The water velocities recorded by us for A. aurantiaca immatures ranged from 0.8–1.1 m/s.

Tonnoir (1925: 219) observed that the following species co-occur with Austrocnephia aurantiaca in Tasma- nia: Austrosimulium (A.) cornutum Tonnoir, A. (Novaustrosimulium) furiosum (Skuse) (as simile), A. (N.) victoriae (Roubaud) (as tasmaniense) and A. (N.) torrentium Tonnoir. Blephariceridae have been recorded from A. aurantiaca habitats collected by H. & P. Zwick in 1977, 1981 & 1998. We have also occasionally encountered blepharicerid larvae in A. aurantiaca habitats—an indication of the high-velocity of breeding sites.

Of the large number of simuliid samples examined by us, only two localities, Godfreys Creek ( NSW, 11 Oct. 1972. Zwick) and Bellell Creek (Highway C153, NSW, Sept. 2014. Craig), yielded each a single parasitized larva. The organism appears as tightly packed spheres (dia. 0.03–0.05 mm) in the abdominal cavity ( Fig. 55 ); similar to that shown by Adler et al. (2004; their Fig. 6.2). Examination at higher magnification reveals that the spheres have a refractive wall and fine grained contents ( Fig. 56 ). It appears to be the Blastocladiales (previously known as a Chytridiales fungus), Coelomycidium simulii Debais 1919 . That species is known too for New Zealand Austrosimulium larvae ( Craig et al., 2012: 33, 294), similarly from cold water, as was the Bellell Creek locality. The organism may, however, possibly be the protist, Tetrahymena sp. ( Adler et al., 2004: 92). As far as is known, this is the first record of such an organism from an Australian simuliid. Litchwardt & Williams (1990) discuss trichomycete gut fungi of Australian simuliid larvae and trichomycetes are now known for Bunyipellum gladiator ( Moulton et al., 2004: 8) and Ectemnoides ( Moulton et al., 2018: 45).

Distribution ( Fig. 196 ). This is a widely distributed species, ranging from Queensland to Tasmania and southwest to the Grampians National Park and the Otway Range, Victoria.

Queensland: Mt. Bartle Frere, Josephine Falls, S17.4420º E145.8600º. 27 Oct. 2002. Adults. Coll. Zwick (ANIC). Mt. Spec , NW Townsville, S18.9500º E146.1800º. 22 Apr. 1955. Adult. Coll. Norris & Common ( ANIC) . Little Nerang Creek , near Gold Coast, S28.1800º E153.2700º. 10 Aug. 1948, 16 Aug. 1950. Adults. Coll. Unknown ( ANIC) . Springbrook , Twin Falls (Little Nerang) Creek, S28.1896º E153.2710º. 23 Jun. 1971. Larvae, pupae, reared adults. Coll. M.H. Colbo ( ROM) . Springbrook , Purling Brook Creek, S28.1898° E153.2709°. 9 Nov. 1973. Larvae. Coll. M.H. Colbo ( ROM) . Carnarvon Gorge National Park , Moss Garden & Hell Hole Gorge, S25.0550º E148.2250º. 13 Feb. 1997. Larva, pupae. Coll. Zwick ( ANIC) . Lamington Plateau , Elabana Falls, S28.1500º E153.0859º. 26 Dec. 1954. Coll. Mackerras ( ANIC) . Wilson’s Peak , Teviot Brook, S28.2500º E152.4800º. 27 Apr. 1971. Larvae. Coll. M.H. Colbo ( ROM) .

New South Wales: Mt. Ebor, Gooch’s Water, S30.4000º E152.3400º. 30Sept.1950.Reared male. 28 Sept. emerged 2 Oct. Reared female. Coll. Unknown (ANIC). New England National Park , S30.5800º E152.4600º. 12 Nov. 1961. Coll. I.F.B. Common & M.S. Upton ( ANIC) . Tubrabucca Falls , S31.8833º E151.4166º. 16 Nov. 1953. Females. Coll. Neboiss ( ANIC) . Barrington Tops, Upper Allyn River, near Whiterock campground, S32.1700º E151.500º. 24 Feb. 2001. Female. Coll. Zwick ( ANIC) . Williams River , S32.4064º E151.7605º. 7 Mar. 1953. Larvae, pupae. Coll. B. McMillan ( UASM) . Newcastle, S32.9300º E151.7800º. Male, female. Coll. Unknown ( UASM) . Crown Creek , S33.1800º E150.1100º. 14 Nov. 1971. Larvae. Coll. Hynes ( ANIC) . Crown Creek, Woods Pl , S33.1818º E150.1179º. 11 Oct. 1972. Larva, pupae. Coll. Zwick ( ANIC & Australian National Museum [ K51170 ]) . Mt. Wilson , S33.5000º E150.3700º. 19 Nov. 1921. Larvae, female. Coll. A.L. Tonnoir ( ANIC) . Govetts Leap Creek, Bridal Veil Falls , S33.6333º E150.3129º. 18 Oct. 2014. Larvae. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) . Wentworth Falls , S33.7274º E150.3742º. 18 Nov. 1921. Male. Coll. A.L. Tonnoir; 22 Nov. 1953. Male. Coll. Neboiss ( ANIC) . Jenolan Caves, S33.8200º E150.0500º. Coll. J. C. Wiburd. Mackerras & Mackerras (1949): Janolan Caves River, below Blue Lake , S33.8202º E150.0269º. 20 Oct. 2014. Larvae. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) . Godfreys Creek , S34.1160º E148.7000º. 2 Apr. 1972, 11/ 12 Oct. 1972. Larva, pupae. Coll. Zwick ( ANIC) . Minnamurra Falls, near Wollongong , S34.6330º E150.7160º. 20 Feb. 2001. Larvae, pupae. Coll. Zwick & Theischinger ( ANIC) . Fitzroy Falls , S34.6470º E150.4825º. 22–27 Nov. 1937. Male. Coll.A.L. Tonnoir ( ANIC) . Pierce Creek State Forest, creek en route to Vanity Crossing , S35.3372º E148.9152º. 11 Oct. 2014. Larvae. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) . Brindabella, Bramina Creek , S35.3835º E148.7410º. 10 Oct. 2011, 4 Oct. 2014. Larvae, pupae, reared adults. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) . Geehi River , S36.2800º E148.0300º. 9 Nov. 1961. Female. Coll. D. H. Colless ( ANIC) . Mt. Kosciuszko, Diggers Creek , S36.2800º E148.5100º. 11 Feb. 1924. Coll. Nicholson ( ANIC) . Mt. Kosciuszko, near Mare Creek, trib. Bogong River , S36.2800º E148.2300º 10 Nov. 2005. Larvae. Coll. Theischinger & Zwick ( ANIC) . Mt. Kosciuszko, Diggers Creek , below falls, S36.3377º E148.4891º. Larvae. Coll. Zwick ( ANIC) . Mt. Kosciuszko, Alpine Way, Bogong Creek , S36.3563 E148.2029. 17 Sept. 2006. Larvae. Coll. Zwick ( ANIC) . Mt. Kosciuszko, Geehi River, Alpine Way , S36.3800º E148.1800º. 4 Nov. 2007. Pupae. Coll. Zwick ( ANIC) . Kosciuszko, Pipers Creek , S36.3839º E148.4347º. 21 Nov. 1972, 4 Jan. 1973. Larvae, pupae. Coll. Zwick ( ANIC) . Pipers Creek , right falls, S36.3839º E148.4347º. 26 June 1972. Larvae. Coll. Zwick ( ANIC) . Mt. Kosciuszko, Snowy River , island, S36.4000º E148.0000º. Aug. 1972. Larvae. Coll. Zwick ( ANIC) . Mt. Kosciuszko, Guthrie’s Creek , S36.4300º E148.3200º. 22 Nov. 1972. Larvae. Coll. Zwick ( ANIC) . Mt. Kosciuszko, Snowy River, Charlottes Pass , S36.4302º E148.3222º. 22 Nov. 1972, 5 Jan. 1973. Larva, pupae. Coll. Zwick ( ANIC) . Mt. Kosciuszko, Threadbo River , S36.5000º E148.3000º. 22/ 23 Oct. 1972. Larva, pupae. Coll. Zwick ( ANIC) . Brown Mountain , February 1939. Coll. Unknown ( ANIC) . Brown Mt., Rutherford Creek , S36.5650º E149.4511º. 10 Mar. 1961. Male. Coll. D.H. Colless ( ANIC) . Brown Mountain, Rutherford Creek , S36.5800º E149.5500º. 10 Mar. 1969. Coll. D.H. Colless ( ANIC) .

Australian Capital Territory: Blundell’s, S35.2900º E149.1400º. October. Coll. A.L. Tonnoir (ANIC). Cotter River , bridge, S35.3228º E148.9504º. Larvae. Coll. Zwick ( ANIC) . Coree Creek , S35.3300º E148.8800º. 21 Jan. 1931. Coll. Unknown. Det. A.L. Tonnoir ( ANIC) . Paddy’s River , S35.3600º E148.9500º, 30 Sept. 1962. Coll. Unknown ( ANIC) . Namadgi Nat. Park , Gibraltar Falls, S35.4897º E148.9293º. Larvae. 8 Oct. 2002. Coll. Zwick ( ANIC) . Namadgi Nat. Park , Bushrangers Creek, S35.4156º E148.8313º, 8 Oct. 2002. Pupae. Coll. Zwick ( ANIC) . Gibraltar Creek Falls , S35.4871º E148.9345º. 26 Aug. 1972. Larvae. Coll. Zwick ( ANIC) . Mt. Gingera , Snowy Flats Creek, S35.5000 E148.8166º. Coll. Unknown ( ANIC) .

Victoria: Omeo Hwy, Lightning Creek , S36.6800º E147.4400º. 13 Apr. 1972. Larva, pupae. Coll. Zwick ( ANIC) . Tawonga , S36.7000º E147.1000º. 16 May 1972. Larva, pupae. Coll. Zwick ( ANIC) . Mt. Buffalo, Eurobin Creek , downstream falls on crossing, S36.7171º E146.8421º. 12 April. Pupae. 7 Nov. 2005. Males. Coll. Zwick ( ANIC) . Mt. Buffalo, Eurobin Creek and Falls , S36.7199º E146.8365º. 13 Apr. 1972, 24 Nov 1972, 10 Oct. 2002. Larva, pupae. Coll. Zwick ( ANIC) . Mt . Stirling, headwater stream, N of summit on Circuit Rd , S36.7300º E147.2800º. 23 Nov. 2005. Larva, pupae. Coll. Zwick ( ANIC) . Slippery Rock Creek, Mt. Beauty Falls , S36.7330º E147.1600º. 14 Apr. 1972. Larvae. Coll. Zwick ( ANIC) . Mt. McKay, Pretty Valley, Howmans Gap , stream, S36.8330º E147.2500º. 7 Jan. 1973. Larvae, pupae, adults. Coll. Zwick ( ANIC) . Falls Creek , west bank, S36.8633º E147.2802º. 7 Jan. 1973. Larva, pupae. Coll. Zwick ( ANIC) . Mt. Ben Cairn, Myrtle Creek , S36.9500º E144.4200º. 7/ 9 Oct. 1972, 25 Dec. 1972. Larva, pupae, males. Coll. Zwick ( ANIC) . Mansfeld, Mt. Buller, Chalet Creek , S37.1200º E146.4400º. 6 Nov. 2005. Larvae. Coll. Zwick ( ANIC) . Mansfeld , Mt . Stirling, S37.1200º E146.4400º. 23 Nov. 2005. Larvae, pupae. Coll. Zwick ( ANIC) . Mirimbah, Mt. Buller, Delatite River , S37.1098º E146.397º. 17 Feb. 2001. Larva, pupae. Coll. Zwick ( ANIC) . Mirimbah, Delatite River , S37.1098º E146.3982º. 11 June 1972, Feb. 1987. Larva, pupae. Coll. Zwick ( ANIC) . Mirimbah, Buller Creek , S37.1333º E146.4166º. 1 Sept. 1972, 24 Nov. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Mansfield, Delatite River , S37.1389º E146.1690º. 20 Dec. 1972, Jan. 1987, 06 Nov. 2005. Larvae, pupae. Coll. Zwick ( ANIC) . Mt. Buller , white bridge, S37.1400º E146.4400º. 11 Apr. 1972, 20 Dec. 1972, 8 Jan. 1973, 17 Feb. 2001. Larvae, adults. Coll. Zwick ( ANIC) . Mt. Buller, Chalet Creek , S37.1400º E146.4400º. 17 Feb. 2001. Larvae, pupae, adults. Coll. Zwick ( ANIC) . Mansfield , Mt . Stirling, Bindaree Falls , S37.1490º E146.5562º. 23 Nov. 2005. Larvae, pupae. Coll. Zwick ( ANIC) . Buller Creek , S37.1500º E146.4400º. 2 Oct. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Howgua River, Stockyard Flats , S37.2200º E146.1800º. 15 Apr. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Eildon Lake, Fryers Creek , S37.3456º E146.0703º. 3 Oct. 1972. Larvae. Coll. Zwick ( ANIC) . Big River, Lake Eildon , S37.2300º E145.9700º. 3 Dec. 1972, Feb. 1987. Larvae, pupae. Coll. Zwick ( ANIC) . Taponga River, Eildon Lake , S37.3700º E146.0500º. 11 June 1972, 17 Dec. 1972. Larvae. Coll. Zwick ( ANIC) . The Grampians, Mackenzie River, McKenzie Falls , S37.1109º E142.4087º. 30 Sept. 1954, Larvae. Coll. Neboiss ( ANIC) , 27 Sept. 1996, Larvae, pupae. Coll. J. K. Moulton ( JKM, UASM) . Grampians, Halls Gap, Fyans Creek , nr Mt. Williams Road , S37.2390º E142.5370º. 11 Nov. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Mt. Cole, 20 ml S. Ararat, Ferntree Falls , S37.3100º E143.2400º. 8 Nov. 1972. Larvae. Coll. Zwick ( ANIC) . Jamieson River , S37.2700º E146.1330º 3 Oct. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Baker Creek, Jamieson , S37.3000º E146.1400º. 28 Apr. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Snobbs Creek Falls , S37.3000º E145.8800º. 9 Nov. 1972, 19 Dec. 1972, 28 Dec. 1972, 6 Feb. 1973. Larvae, pupae. Coll. Zwick ( ANIC) . The Grampians, Kalymna Falls , S37.3229º E142.6187º. 9 Nov. 1972. Larvae, pupae, adults. Coll. Zwick ( ANIC) . Royston River , S37.3833º E145.8833º. 28 Dec. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Flourbag Creek, Jamieson , S37.3840º E146.2120º. 28 Apr. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Kinglake West, King Parrot Creek , S37.4283º E145.2496º. 18 Oct. 1953, 13 Dec. 1953. Male. Coll. Neboiss ( ANIC) . Taponga River , S37.4400º E146.0200º. 19 Dec. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Gaffneys Creek, Woods Point Road , S37.4606º E146.1893º. 27 Jan. 1972, 11 Oct. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Kinglake, Masons Falls , S37.4900º E145.2480º. 13 Dec. 1953. Coll. Neboiss ( ANIC) . Marysville, Taggerty River, Lady Talbot Drive , S37.5045º E145.7957º. 21 Nov. 2005. Female. Coll. Zwick ( ANIC) . Narbethong, Acheron River , S37.5284º E145.6884º. Larvae, pupae. Coll. Zwick ( ANIC) . Marysville, upper Big River, Cumberland Rd , S37.56200º E145.9035º. 22 Nov. 2005. Pupae. Coll. Zwick ( ANIC) . Maryville, Steavenson Falls , S37.5317º E145.7727º. 26 Apr. 1972, 11 June 1972, 8 Sept. 1972, 4 Nov. 1972, 29 Dec. 1972. Larvae, pupae. Coll. Zwick ( ANIC) Marysville, Steavenson River, turbine house, Steavenson Falls , S37.53144º E145.7728º. 24 Sept. 2014, 22 Oct. 2014. Larva, pupae. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) . Healesville, Myer’s Creek , S37.5900º E145.5100º. 16 Jul. 1972. Larvae. Coll. Zwick ( ANIC) . Marysville, Keppel Falls , S37.6004º E145.8391º Feb 1987. Larvae, pupae. Coll. Zwick ( ANIC) . Marysville, Cumberland Falls , S37.6300º E145.7700º. 11 June 1972, 28 Dec. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Acheron River, Acheron’s Gap , S37.6800º E145.7500º. 20 Apr. 1972. Larvae. Coll. Zwick ( ANIC) . Warburton, Cement Creek , S37.7239º E145.7542º. 2 Feb. 1972, 4 Feb. 1972, 1 Mar. 1972, 4 July 1972, 7 Oct. 1972, 8 Nov. 1972, 25 Dec. 1972, 20 Nov. 2005. Larvae, pupae, female. Coll. Zwick ( ANIC) . Dandenong Range, Barbers Road, Olinda Creek , S37.8290º E145.3763º. 26 Oct. 2014. Larva, pupae. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) . Noojee, Torongo Fall, S37.8481º E146.0512º. 3 Jan. 1973. Female. Coll. Zwick. ANIC. Baw Baw , East Tanjil River , S37.8590º E146.1873º. 18 Nov. 2005. Larvae, pupae. Coll. Zwick ( ANIC & Australian National Museum [ K51171 ]). Sassafras, S37.8600º E145.3500º. October. A.L. Tonnoir (1925) ( ANIC) . Mt. Dandenong, Sherbrooke Falls, S37.8874º E145.3513º. October. A.L. Tonnoir (1925). 7km W Neerim, Torago River , S37.9500º E145.8900º, 1 March. 1972. Adult. Coll. Neboiss ( MM) . German Creek , S37.9851º E146.1496º. 23 Nov. 1972. Larvae, pupae. Coll. Zwick ( ANIC) . Gippsland, North of Moe, Tyers Riv- er, S38.1600º E146.2400º. 17 Nov. 2005. Larvae, pupae. Coll. Zwick ( ANIC) . Otway Ranges, Carisbrook Creek Falls , S38.6876º E143.8101º. 18 Aug. 1972, 13 Oct. 2002. Larvae, pupae, male. Coll. Zwick ( ANIC) . Great Ocean Road, Carisbrook Creek , bridge, S38.6909º E143.8092º. 20 Nov. 201. Larvae. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) . Great Ocean Road, Smythe Creek , S38.7033º E143.7617º. 21 Nov. 2014. Larva, pupae. Colls. D.A. & R.E.G. Craig ( ANIC, UASM) .

Flinders Island (Bass Strait): Mt. Strezlecki, S40.2400º E148.0800º. Jan. 1953. Coll. R. Mykytowycz. Larvae (1 slide), pupae (ANIC).

Tasmania: Cradle Valley, S37.8600º E145.3500º. 19 Jan. 1923. A.L. Tonnoir (ANIC); Jan, February. ( Mackerras & Mackerras 1949). Duck River, 6km SW Roger River, S41.0058° E145.0672°, 29 Nov 1974. Adult. Coll. Neboiss (MM). Roger River, Buffs Road, Roger River Road, S41.0432º E145.0172º, DUCK28. 16 Mar. 1995. Larvae. Coll. M. Nelson, DPIPWE (TMAG). Burnie, S41.0500º E145.8900º. Coll. Tonnoir (1925). Penguin, Penguin Creek, Hayward Street, S41.1139º E146.0690º, BLYT04. 5 Apr. 2006 Larvae. Coll. D. Horner, DPIPWE (TMAG). Meander River, Falls Road, S41.2307º E147.6098º, MEAN13. 14 Nov. 2002. Larvae. Coll. D. Horner, DPIPWE (TMAG). Hogarth Rivulet, end of Falls Road, S41.2307º E147.6098º, GFOR31. 19 Nov. 2002, 20 Mar. 2003, 7 Oct. 2003, 5 Oct. 2004, 19 Apr. 2005, 4 Nov. 2005, 1 Apr. 2009. Larvae. Colls. T. Krasnicki, D. Horner, S. Meyer, Latton, DPIPWE (TMAG). Mackenzie Valley Road, Mackenzie Rivulet, S41.2312º E147.5494º, GFOR35. 10 May 2005. Larvae. Coll. D. Horner, DPIPWE (TMAG). Wilmot River, Alma Reserve, S41.2606º E146.2323º, FORT06. 17 Mar. 2004. Larvae. Coll. A. Foley, DPIPWE (TMAG). Arthur River, Pykes Bridge, S41.2636º E145.5154º, ARTH01. 27 Oct. 2004. Larvae. Coll. A. Harvey, DPIPWE (TMAG). Great Forester River, off East Diddleum Road, S41.2670º E147.5070º, GFOR17. 6 Oct. 2003, 4 Oct. 2004, 8 Nov. 2005. Larvae. Colls. M. Read, S. Meyer, D. Horner, T. Krasnicki, DPIPWE (TMAG). Hellyer River, S41.2700º E145.6100º. 2 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). St. Patricks River, Corkerys Road, S41.2936º E147.4160º, NESK22. 29 Mar. 1995. Larvae. Coll. D. Oldmeadow, DPIPWE (TMAG). South George River, St. Columba Falls Road, S41.2981º E147.9572º, GEOR07. 30 Mar. 1995. Larvae. Coll. D. Oldmeadow, DPIPWE (TMAG). St. Patrick River, S41.3600º E147.3100º. Coll. Tonnoir (1925). Launceston Gorge, S41.4400º E147.1200º. 8 Dec. 1972. Larvae. Coll. Zwick (ANIC). Whyte River, Waratah Road, S41.4667º E145.3906º, PIEM02. 27 Oct. 2004. Larvae. Coll. J. Latton, DPIPWE (TMAG). Cradle Mt. road waterfall, near Celthana Dam, S41.4700º E146.1600º. 9 Dec. 1972. Larvae. Coll. Zwick (ANIC). Mersey River, upstrm of Union Bridge, S41.5100º E147.3300º, MERS06. 14 Nov. 2002. Larvae. Coll. D. Horn- er, DPIPWE (TMAG). Dove Lake, S41.6660º E145.9666º. 9 Dec. 1972. Pupae. Coll. Zwick (ANIC). Meander River, dwnstr dam site, S41.5200º E147.0300 º, MEAN10. 22 Oct. 2003. Larvae. Coll. T. Krasnicki, DPIPWE (TMAG). Mersey River, Dogs Head Hill, S41.5210º E145.2870 º, MERS07. 11 Nov. 2002. Larvae. Colls. D. Horner, C. Amos, DPIPWE (TMAG). Blythe River, South Riana Road, S41.5688º E147.2391º, BLYT01. 29 Oct. 2003. Larvae. Coll. A. Foley, DPIPWE (TMAG). Jackeys Creek, dwnstrm of Jackeys Marsh, S41.6716º E146.6467º, MEAN31. 19 Oct. 2006. Larvae. Coll. D. Spiers, DPIPWE (TMAG). Liffey River, upstm of Liffey, S41.6809º E146.9260º, MEAN16. 23 Oct. 2003, 18 Nov. 2005, 11 Oct. 2006. Larvae. Colls. A. Foley, T. Krasnicki, J. Beard, DPIPWE (TMAG). Forth River, waterfall, S41.6900º E146.0750º. 29 Jun. 1967. Adult. Coll. E. F. Riek. 10 Dec. 1972. Larvae. Coll. Zwick (ANIC). Mt. Farrell, S41.7500º E145.6300º. Coll. Tonnoir (1925). Fish River, Mersey Forest Road, S41.7714º E146.2251º, MERS34. 15 Nov. 2005. Larvae. Coll. T. Krasnicki, DPIPWE (TMAG). Stitt River, upstream of Roseberry, S41.7919º E145.5409º, PIEM22. 11 Apr. 1995. Larvae. Coll. M. Nelson, DPIP- WE (TMAG). Mackintosh River, S41.8833º E145.8833º. 3 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). Ewart Creek, Zeehan Highway, S41.9576º E145.4631º, HENT13. 29 Apr. 2003, 16 Mar. 2004, 26 Oct. 2004, 28 Mar. 2006. Larvae. Colls. J. Latton, A. Harvey, M. Read, D. Warfe, DPIPWE (TMAG). Queenstown, Ewarts Creek Road, S41.9941º E145.4716º. 3 Dec. 1972, 6 Dec. 1972. Larvae. Coll. Zwick (ANIC). Allens Rivulet, Kaoota Road, S42.0199º E147.1652º, HOBA15. 5 Oct. 2005. Larvae. Coll. J. Latton, DPIPWE (TMAG). Strahan, Botanical Creek, Hogarth Falls, S42.1498º E145.3479º. 4 Nov. 2014. Larvae. Colls. D.A. & R.E.G. Craig (UASM). Collingwood River, S42.1600º E145.9200º. 4 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). Henry River, S42.2200º E146.0210º. 3 Dec. 1972. Pupae. Coll. Zwick (ANIC). Florentine River, Florentine Road, S42.4433º E146.5052º, UDER008. 16 Mar. 2004, 19 Oct. 2005, 16 Oct. 2006. Larvae. Colls. D. Horner, K. Hoyle, D. Spiers, DPIPWE (TMAG). Russell Falls, S42.6700º E146.7100º. Dec. 1922. Larvae, pupae. Coll. Tonnoir (UASM) Mt. Field Nat. Park, Lady Barron Falls, S42.6838º E146.7172º. 6 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). Mt. Field Nat. Park, Russell Falls, S42.6841º E146.7199º. 5 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). Mt. Field National Park, Lady Barron Creek, S42.6881º E146.6989º. 7 Nov. 2014. Larvae. Colls. D.A. & R.E.G. Craig (UASM). Mt. Field Nat. Park, Tyenna River, S42.7200º E146.6700º. 6 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). Ferntree, nr Hobart, S42.9200º E147.2600º. 7 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). North West Bay River, Betts Road, S42.9544º E147.2066º, HOBA02. 20 Apr. 2006. Larvae. Coll. D. Horner, DPIPWE (TMAG). North West Bay River, upstream of Allens Rivulet, S42.9818º E147.1996º, HOBA06. 21 Oct. 2005. Larvae. Coll. D. Horner, DPIPWE (TMAG). North West Bay River, Margate weir, S42.9893º E147.2214º, HOBA05. 28 Apr. 2006. Larvae. Coll. D. Horner, DPIPWE (TMAG). Hartz Mts., Huon River, S43.1600º E146.6900º. 6 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). Kermandie River, Oigles Road, S43.1681º E146.8787º, HUON013. 24 Oct. 2003, 28 Sept. 2005. Larvae. Colls. D. Warfe, D. Spiers, DPIPWE (TMAG). Hartz Mts., Arve River, S43.2600º E146.8000º. 7 Dec. 1972. Larvae, pupae. Coll. Zwick (ANIC). Esperance River, Esperance River Road, S43.2774º E146.8765º, HUON046. 8 Apr. 2004, 12 Oct. 2006. Larvae. Colls. A. Harvey, D. Spiers, DPIPWE (TMAG). Bruny Island, S43.4000º E147. 2600º. Coll. A.L. Tonnoir (1925).

Remarks. Austrocnephia aurantiaca is a highly variable species, as noted by Tonnoir (1925: 236) in his original description. Based on our examination of a large series of material from many localities, ranging from near Cairns in Queensland, south along the Great Divide Mountain Range to Tasmania and west to the Otway Range and the Grampians in Victoria, we conclude that several species likely occur under this name. In fact, Rothfels (1979: 522), in a cytological study of what might have been A. aurantiaca (as Cnephia A), reported the presence of two sibling species. Of particular note are the morphological difference between larvae of mainland A. aurantiaca ( Fig. 36 ) and those from Tasmania ( Fig. 37 ). The latter are distinct in colour, shape and size of the pupal gill histoblast, and size of the posterior proleg. The male paratypes designated by Tonnoir (1925) from Tasmania also exhibit differences from mainland material. It is highly likely that the Tasmania population of A. aurantiaca is specifically distinct from those on the mainland. Another character state, the presence of small but distinct prothoracic proleg lappets in certain populations ( Figs. 49–51 ) is perhaps also indicative of specific difference. This feature is markedly expressed in A. strenua , but is entirely absent from most populations of A. aurantiaca . The intermediate stage as expressed in certain populations of ‘ A. aurantiaca ’ requires further scrutiny. Finally, the apically branched Rs, is distinct in some populations and less so in others (i.e., is represented by just a few hairs in a row). Further morphological cytogenetic and molecular studies are needed to confidently circumscribe entities in this complex.

Males of A. aurantiaca and A. strenua are distinct, although the latter exhibits a great deal of colour variation ( Tonnoir, 1925: 236). Overall the body in A. aurantiaca males is bright yellow ( Fig. 18 ) whereas that of A. strenua is darker brown (Fig. 73). Further, the eyes of A. strenua males have fewer rows of larger upper ommatidia and are dark brown.

The ventral plate of A. aurantiaca is rather complex and must be interpreted with caution. Few of the males examined so far have the depressed apex of the ventral plate as illustrated by Mackerras & Mackerras (1950: their Fig 3 ), where it is described as being ‘distinctly concave’. When examined in three dimensions, the tip is directed ventrally, giving the plate a concave appearance in ventral view ( Fig. 25 ), thence the original description. If the ventral plate is flattened out on a slide mount ( Fig. 27 ), it appears to be rounded apically. All males examined have parameres plate-like and broadly attached to both the gonocoxa and basal arms, with the paramere connection apparently absent. The parameres are tapered posteriorly with various development of small spines distally on the aedeagal membrane. Apical spines of the gonostylus are poorly developed in A. strenua (Mackerras & Mackerras, loc. cit.). Female A. aurantiaca possess a rugose-wrinkled spermatheca as originally described ( Fig. 16 ). Pupa possess only a small pleurite on abdominal segment V ( Fig. 35 ) and this agrees largely with Mackerras & Mackerras (1950: 169) who state that pleurites are absent.

Tonnoir (1925: 226. His Fig. 10B ) illustrated the anal sclerite of A. aurantiaca larvae as having a distinct ‘hole’ at the posteromedial base of the structure and the posterolateral arms single, albeit substantial. With the widespread collections now available, the first character still is of value for identification, but the second not. The posteroventral arms are varied and while can be single, may be substantial and paralleled by a posterolateral arm, as seen in larvae of A. strenua . Similarly, as for A. strenua larvae, there are a large number of campaniform sensilla between the posterolateral arm, as there are number of hooks in the posterior circlet—indeed, these two character states are diagnostic for the aurantiaca species-group.

Mackerras & Mackerras (1955:105) noted that the Flinders Island (Bass Strait) population of Austrocnephia aurantiaca was subtly different from those of the mainland and Tasmania, but did not give any details. We illustrate ( Fig. 46 ) the larval hypostoma, but with no recent material available we have not pursued their observation. However, with the gross differences in larvae (cf. Figs. 36, 37 ) of A. aurantiaca of the Australian mainland and Tasmania, as noted previously, we are fully of the opinion that they will be shown to be related but separate taxa.

Collection data reveals that A. aurantiaca was present in The Grampians National Park and environs from at least the mid 1950s through the mid 1990s; however, more recent efforts (2011, 2014) to collect this species from those same localities have proven unsuccessful. While high-velocity habitats remain within the Park, such as the Mackenzie Falls, the region has recently suffered from drought and intensive forest fires. Consequently, many of the watercourses where A. aurantiaca immatures were previously collected are now dry or characterized by low flow. Could an extended period of drought account for the apparent lack of A. aurantiaca from The Grampians? Other local species of simuliid, such as Nothogreniera fergusoni (Tonnoir) and Simulium ornatipes Skuse , are adapted to breed in intermittent streams that flow during the winter and early spring. Another commonly encountered species, Austrosimulium furiosum , is a generalist species that can breed in a wide variety of running water habitats. Whether Austrocnephia aurantiaca has truly been extirpated from The Grampians warrants further investigation.

The record ( Fig. 196 ) of a single A. aurantiaca adult from Mt. Spec, Queensland, (clustered with A. strenua ), should be regarded with caution. It is anomalously north of other known localities.

Figures

FIGURE 9. Austrocnephia aurantiaca female. (9) Wing, showing veins and basal medial cell (bm) and basal radial cell (br). (Brindabella). Scale bar = 1.0 mm.

FIGURE 28. Austrocnephia aurantiaca pupa. (28) Habitus, male. (Carisbrook). Scale bar = 1.0 mm.

FIGURE 35. Austrocnephia aurantiaca pupa. (35) Abdominal armature. (Brindabella).Arrow indicates pleurite (plt). Scale bar = 0.5 mm.

FIGURE 196. Distribution of Austrocnephia aurantiaca species-group (A. aurantiaca, A. strenua).

FIGURES 1–6. Austrocnephia aurantiaca female. (1) Dorsal view of paratype. (Mount Wilson). Image by LHG-A. Scale bar = 0.5 mm. (2) Habitus. (Steavenson Falls). Scale bar = 1.0 mm. (3) Frontal view of head. (Steavenson Falls). Scale bar = 0.2 mm. (4) Antenna, showing flagellum, pedisulcus (pd) and scape (sc). (Brindabella). Scale bar = 0.1 mm. (5) Maxillary palp, showing palpomeres III–V, sensory vesicle (s v) and lacinia (lc) (Brindabella). Scale bar = 0.1 mm. (6) Mandible (mnd) and lacinia (lc). (Steavenson Falls). Scale bar = 0.05 mm.

FIGURES 7, 8. Austrocnephia aurantiaca female. (7) Cibarium showing cornuae (cn). (Steavenson Falls). Scale bar = 0.05 mm. (8) Wing veins showing branched Rs. (Brindabella). Scale bar = 0.1.mm.

FIGURES 10, 11. Austrocnephia aurantiaca female. (10) Calcipala (cpl) and tarsomere II. (Steavenson Falls). Scale bar = 0.1 mm. (11) Tarsal claws, heel (h) and basal tooth (t). (Brindabella). Scale bar = 0.02 mm.

FIGURES 12–16. Austrocnephia aurantiaca female. (12) Abdominal tergites. (Brindabella). Scale bar = 1.0 mm. (13) Hypogynial valves (hyp v). (Brindabella). Scale bar = 0.1 mm. (14) Anal lobe (a l) and cercus (c). (Brindabella). Scale bar = 0.05 mm. (15) Genital fork, anterior stem (a st), apodeme (ap) and lateral plate (l p). (Brindabella). Scale bar = 0.05 mm. (16) Spermatheca. (Steavenson Falls). Scale bar = 0.1 mm. FIGURE 17. Austrocnephia. aurantiaca male. (17) Holotype. (Mount Wilson). Image by LHG-A. Scale bar = 1.0 mm.

FIGURES 18, 19. Austrocnephia aurantiaca male. (18) Habitus. (Taggerty River). Scale bar = 1.0 mm. (19) Frontal view of head. (Brindabella). Scale bar = 0.2 mm.

FIGURES 20–23. Austrocnephia aurantiaca male. (20) Antenna. (Brindabella). Scale bar = 0.1 mm. (21). Maxillary palp and lacinia. Paratype. (Mount Farrell, Tasmania). Scale bar = 0.1 mm. (22) Wing veins. Arrow indicates R2+3. (Brindabella). Scale bar = 0.1 mm. (23) Claws and grappling hooks (g h). (Brindabella). Scale bar = 0.02 mm.

FIGURES 24–27. Austrocnephia aurantiaca male. (24) Abdominal tergites (t) and sternites (st). Paratype. (Mount Farrell, Tasmania). Scale bar = 0.5 mm. (25) Genitalia (ventral view). Note downturned apex of ventral plate. (Carisbrook). Scale bar = 0.5 mm. (26) Gonocoxa (gc) and gonostylus (gs). Paratype. (Mount Farrell, Tasmania). Scale bar = 0.05 mm. (27) Slide mounted genitalia showing adeagal membrane (a m), basal arms (b a), parameres (p), spines (s) and ventral plate (v p). (Brindabella). Scale bar = 0.05 mm.

FIGURES 29, 30. Austrocnephia aurantiaca pupa. (29) Female cephalic capsule. (Brindabella). Arrows indicate setae. Scale bar = 0.2 mm. (30) Male cephalic capsule. (Brindabella). Scale bar = 0.2 mm.

FIGURES 31–34. Austrocnephia aurantiaca pupa. (31) Female thoracic cuticle. (Steavenson Falls). Scale bar = 0.2 mm. (32) Gill. (Carisbrook). Scale bar = 0.5 mm. (33) Gill filament apex surface. (Carisbrook). Scale bar = 0.05 mm. (34) Cocoon silk. (Brindabella). Scale bar = 0.1 mm.

FIGURES 36, 37. Austrocnephia aurantiaca larva. (36) Habitus, last instar. (Steavenson Falls). Scale bar = 1.0 mm. (37) Habi- tus, last instar. (Hogarth Rivulet, Tasmania). Scale bar = 1.0 mm.

FIGURES 38, 39. Austrocnephia aurantiaca larva. (38) Dorsal view of head, last instar. (Steavenson Falls). Scale bar = 0.25 mm. (39) Antenna. (Steavenson Falls). Scale bar = 0.05 mm.

FIGURES 40–45. Austrocnephia aurantiaca larva. (40) Mandible. (Steavenson Falls). Scale bar = 0.05 mm. (41) Mandible apex showing apical tooth (ap t) and spinous teeth (sp t). (Pretty Valley). Scale bar = 0.05 mm. (42) Maxilla and palp (mx p). (Carisbrook). Scale bar = 0.1 mm. (43) Ventral head cuticle showing postgenal cleft (pg c). (Steavenson Falls). Scale bar = 0.1 mm. (44) Hypostoma. Numbers indicate teeth. (Steavenson Falls). Scale bar = 0.05 mm. (45) Hypostoma. (Carisbrook). Scale bar = 0.05 mm.

FIGURES 46–49. Austrocnephia aurantiaca larva. (46) Hypostoma. (Flinders Island). Scale bar = 0.1 mm. (47) Hypostoma. (Huon River, Kerranda, Tasmania). Scale bar = 0.1 mm. (48) Pupal gill histoblast. (Steavenson Falls). Scale bar = 0.25 mm. (49) Prothoracic proleg. Arrow indicates rudimentary lappet. (Steavenson Falls). Scale bar = 0.1 mm.

FIGURES 50, 51. Austrocnephia aurantiaca larva. (50) Prothoracic proleg.Arrow as for Fig. 49. (Steavenson Falls). Scale bar = 0.25 mm. (51) Prothoracic proleg, Arrow as for Fig. 49. (Steavenson Falls). Scale bar = 0.2 mm.

FIGURES 52–56. Austrocnephia aurantiaca larva. (52) Larval anal sclerite and circlet of hooks showing anterior arm (a a), posteroventral arms (pv a), campaniform sensilla (c s), circlet of hooks (c h). (Carisbrook). Scale bar = 0.2 mm. (53) Anal sclerite and circlet of hooks. (Flinders Island). Scale bar = 0.2 mm. (54) Anal sclerite and circlet of hooks, showing posterolateral arms (pl a). (Pretty Valley). Scale bar = 0.2 mm. (55) Parasitized larva. (Godfreys Stream, NSW). Scale bar = 0.5 mm. (56) Coelomycidium (?)simulii. (Godfreys Stream, NSW). Scale bar = 0.1 mm. FIGURE 57. Austrocnephia aurantiaca typical habitat. (57) Steavenson River, NSW. 24 Sept. 2014.