Austrocnephia tonnoiri ( Drummond 1931 ), 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 : 63-72

publication ID	https://doi.org/ 10.11646/zootaxa.4627.1.1
publication LSID	lsid:zoobank.org:pub:A7802D6F-D366-44DE-82D9-F0AAC7468157
persistent identifier	https://treatment.plazi.org/id/038A87CC-FFA5-4442-FF30-FF76AA875496
treatment provided by	Plazi
scientific name	Austrocnephia tonnoiri ( Drummond 1931 )
status

Treatment

Austrocnephia tonnoiri ( Drummond 1931) View in CoL . New combination.

(Figs. 164–195)

Simulium tonnoiri Drummond 1931: 6 View in CoL ; original designation.

Cnephia tonnoiri tonnoiri . Mackerras & Mackerras, 1950: 169; new combination.

Cnephia tonnoiri . Rothfels, 1979: 522.

(‘ Cnephia ’ of authors) tonnoiri . Crosskey, 1987: 443; undetermined genus in Prosimuliini .

(Unplaced species of Prosimuliini ) tonnoiri . Crosskey, 1989: 222.

Cnephia tonnoiri tonnoiri . Trayler et al., 1996: 286.

Paracnephia tonnoiri. Crosskey & Howard, 1997: 18 ; new combination, unplaced to subgenus. Bugledich, 1999: 328. Crosskey & Howard, 2004: 10. Adler & Crosskey, 2008: 26; transferred to Simuliini View in CoL , unplaced to subgenus. Adler, 2019: 33.

Redescription. Adult female (based on a few pinned adults plus pharate adults and literature description). Body: head and thorax brownish orange (Figs. 164, 165); abdomen yellow and black; total length ca. 2.8–2.9 mm. Head (Fig. 166): overall reddish brown; width 0.85–0.97 mm, depth 0.57–0.63 mm; postocciput black, vestiture of dense hairs; frons not markedly narrowed, dark brown, vestiture of moderately dense silvery hairs; frons:head-width ratio 1.0.0:6.0–6.2. Eyes: interocular distance 0.16 mm; dark blackish orange, upper ommatidia lighter; ommatidia diameter 0.023 mm; ca. 37 rows across and down at mid-eye. Clypeus: width 0.18–0.24 mm; dark brown, moderate vestiture of fine silvery hairs. Antenna: total length 0.73–0.78 mm, extended well beyond posterior margin of head; evenly dark brown; scape and pedicel slightly broader than flagellomere I, flagellomere II shorter than first, remainder rectangular, barely tapered to apex. Mouthparts: moderately developed, ca. 0.4× length of head depth; maxillary palp, total length 7.4 mm, palpomeres I & II small, palpomere III darker brown than others, dense vestiture, IV small, V elongated; proportional lengths palpomeres II–V 1.0:0.7:1.5; sensory organ moderately elongated, 0.65× length of palpomere III, opening normal; mandible pointed apically with ca. 27 markedly small inner teeth, outer teeth absent, but with irregular edge, essentially as for A. fuscoflava ; lacinia with 13–15 teeth on inner edge, nine teeth on outer; cibarium ( Fig. 167 View FIGURES 167, 168 ) cornuae broad basally, not markedly flared, tapered and curved apically, medial gap small and angulate. Thorax: length 1.4 mm, width 1.3 mm; evenly orange brown; postpronotal lobe well developed with dense fine hair, scutellum slightly paler than scutum, vestiture of sparse very fine yellowish hairs, long black hairs laterally; postnotum concolourous with scutellum; antepronotal lobe with dense pale hair; proepisternum and fore coxa bare; pleuron light brown, anepisternal membrane bare; katepisternal sulcus shallow and broad. Wing: length 2.8–3.4 mm, width 1.5–1.7 mm; membrane slightly fumose on apex and anal lobe, area between costal and radial sector veins faintly yellowish; distal 2/3 of costa with spines, distal 1/3 Sc bare, Rs not branched; a:b ratio 1.0:2.7; M 1 appearing distinctly doubled; CuA slightly sinuous; markedly distinct small pigmented region at r-m cross veins—comprised of pigmented sclerotized region, pigmented veins and concentration of hairs (as for male, i.e., Fig. 176 View FIGURES 175–179 ). Haltere: stem pale, knob darker. Legs: evenly yellowish brown, joints tending to darker; hind basitarsus with regular row of stout spines, absent distally; calcipala markedly developed with dorsal notch; pedisulcus only moderately expressed; tarsomere II ca. 1.8× as long as distal width; claw talon fine ( Fig. 168 View FIGURES 167, 168 ) and smoothly tapered, basal tooth cone-shaped, 1/3 length of talon, rounded heel moderately expressed. Abdomen: basal scale dark brown, vestiture of long yellowish hairs; anterior segments yellowish, other segments dark brown, vestiture of dense long pale hairs laterally and posteriorly; tergites markedly poorly sclerotized, barely discernable from remainder of dorsum. Genitalia: sternite VIII dark with distinct anterior edges and medial region, vestiture of microtrichia, large strong hairs posterolaterally; hypogynial valves ( Fig. 169 View FIGURES 169–172 ) lightly pigmented, vestiture of triads of microtrichia and strong hairs apically, medial edges of valves slightly concave, lightly strengthened anteromedially, moderately rounded apically, with ill-defined edge; cercus in lateral view elongated and rounded apically, with slight medial depression, anal lobe elongated, subequal in length to cercus ( Fig. 170 View FIGURES 169–172 ); genital fork ( Fig. 171 View FIGURES 169–172 ) finely expressed, anterior stem narrow, rounded apically, when in situ markedly curved dorsally (distorted in image), no indication of membranous lateral areas, lateral arms narrow, apodeme absent, lateral plates bifurcate; spermatheca elongate and ovoid, dark brown, slightly wrinkled, internal fine spines not obvious, clear membranous area at junction with spermathecal duct small with raised edge ( Fig. 172 View FIGURES 169–172 ).

Adult male (lectotypes and other specimens from ANIC). Body ( Figs. 173, 174 View FIGURES 173, 174 ): overall yellowish orange and black; total length 2.3–3.1 mm. Head ( Fig. 175 View FIGURES 175–179 ): overall brown; width 0.90–0.95 mm, depth 0.71–0.79 mm; hairs yellow. Eyes: upper ommatidia bright yellowish orange, large, diameter 0.047 mm, ca. 16 across and down; lower ommatidia blackish orange, markedly smaller, diameter 0.018 mm, ca. 32 across and down. Clypeus: black; width 0.13–0.21 mm; vestiture of sparse stiff black hairs. Antenna: total length 0.57–0.75 mm; darker basally, evenly yellowish brown otherwise; scape and pedicel subequal in length, scape dark brown, flagellomere I slightly narrower than scape, other flagellomeres tapered slightly to apex. Mouthparts: insubstantial; length 0.16–0.20× head depth; maxillary palp 0.63 mm long, palpomeres I & II small, palpomeres III & IV subequal in length, proportional lengths of palpomeres III–V 1.0:0.8:1.8, sensory vesicle spherical in shape, occupying 0.25× palpomere length, opening 0.5× vesicle width; lacinia with hairs apically; mandible not observed. Thorax: length 1.3–1.4 mm, width 0.80–0.96 mm; markedly domed, head angled ventrally; postpronotal lobe with slightly longer hairs than scutum; antepronotal lobes with sparse clump of longish fine pale hairs, scutum evenly medium brown, vestiture of fine golden hairs longer anteriorly, scutellum concolourous with scutum, vestiture of long pale hairs laterally, postnotum darker than scutum; proepisternum bare; pleuron brown, anepisternal membrane bare. Wing: length 2.9 mm, width 1.4 mm; membrane slightly fumose at apex, slightly so on anal lobe; yellowish tint between C and Sc; basal medial cell not apparent; a:b ratio 1.0:3.0; costa with spines, marked pigmented region at r-m cross veins, comprised of concentration of hairs on R 1 and pigmentation of junction ( Fig. 176 View FIGURES 175–179 ); Rs not branched; M 1 appearing distinctly doubled, CuA not markedly sinuous. Haltere: base of stem pale, knob dark tan. Legs: overall yellowish with dark articulations; hind basitarsus with ventral row of stout spines, absent distally; calcipala essentially as for female; tarsal claw as for other species. Abdomen: overall yellow and black, basal scale hairs pale yellow, extended to posterior of segment IV, tergites not markedly sclerotized, 2.0–3.5× as wide as long, markedly hirsute laterally, more so on posterior segments, hairs yellowish, long and dense; pleurites absent; sternites essentially absent, slightly expressed posteriorly. Genitalia: overall small and moderately sclerotized ( Fig. 177 View FIGURES 175–179 ); gonocoxa 2× longer than its basal width, strengthened and scalloped posteromedially, vestiture of sparse hairs ( Fig. 178 View FIGURES 175–179 ); gonostylus in ventral view narrowed and tapered, broad in lateral view, approximately 1.5× longer than basal width, two substantial, short, blunt, apical spines; ventral plate ( Fig. 179 View FIGURES 175–179 ) small, simple, 2.0× wider than long in ventral view, posterior edge thickened apically, slightly concave to convex (depending on view), broadly rounded laterally with straight edge angled anteriorly towards basal arms, slightly convex anteromedially, vestiture essentially absent, but with sparse central hairs, basal arms distinct, albeit not markedly developed, paramere connectors well expressed; median sclerite poorly expressed, as two widely separated arms extended just beyond posterior edge of ventral plate, junction with ventral plate not obvious; parameres plate-like basally, tapered distally with corrugations apically, spines poorly expressed as three or four small blunt structures ( Fig. 178 View FIGURES 175–179 ); adeagal membrane with sparse microtrichia.

Pupa (based on numerous specimens). Body: female length 3.0– 4.2 mm ( Fig 183 View FIGURES 181–183 ), male length 3.2–3.6 mm; cuticle clear brown. Head: frons of female quadratic, ratio of basal width to vertex width and height, 1.0:1.3 and 1.0:1.6 respectively (Fig. 180), that of male ovoid; ratios 1.0:1.9 and 1.0:2.5 respectively ( Fig. 181 View FIGURES 181–183 ), cuticle not tuberculate; in male, frontal and facial setae present, in female frontal setae absent, in both male and female facial setae are well developed and curved—almost spine-like; antennal sheath of female extended beyond margin of ocular shield, that of male markedly not so. Thorax: smooth, dorsocentral setae, stiff, spine-like, tips usually curled. Gill ( Fig. 182 View FIGURES 181–183 ): total length ca. 0.95–1.60 mm; antler-like with 26–36 filaments arising from five to seven markedly short basal trunks, branching at irregular intervals, some branches short, surface with distinct fine pseudoannulations, smooth apically; tips often broken in mature pupae, otherwise as for A. orientalis . Abdomen ( Fig. 184 View FIGURE 184 ): essentially as for A. fuscoflava .

Cocoon ( Fig. 183 View FIGURES 181–183 ). Close fitting, usually covering pupa completely, slightly slipper-shaped; irregular weave, silk fibers fine; considerable extraneous material incorporated.

Larva (based on numerous specimens, Kangaroo Gully). Body ( Fig. 185 View FIGURES 185–187 ): total length 6.6–7.6 mm; overall mottled grey; smoothly expanded from the thorax posteriorly. Head ( Fig. 186 View FIGURES 185–187 ): overall markedly bicolourous, yellow and dark brown, head spot pattern positive; length 0.75–0.96 mm, width 0.71–0.83 mm; distance between antennal bases 0.36–0.43 mm; anterior margins of head subparallel, diverging posterior of stemmata; ecdysial lines well visible, slightly divergent until posterior of stemmata, then very broadly curved laterally then medially; cervical sclerites elongated and fused to postocciput; genae brown. Antenna: not extended to end of labral fan stem; total length 0.30–0.35 mm; basal article pale proximally, darker brown distally, medial article and distal articles evenly light brown; medial article slightly narrowed along length, basal article markedly shorter than medial article, proportional lengths of basal, medial and apical articles 1.0:1.2:1.7. Labral fan: stem short, not markedly pigmented, ca. 26–36 fine rays, ten posterior rays finer than others, length 0.74–0.88 mm, mid-ray width 0.014 –0.018 mm; pattern of microtrichia not marked, longer microtrichia subequal in length to ray width. Mandible ( Fig. 187 View FIGURES 185–187 ): darkly pigmented; not noticeably short; outer teeth short and distinct; apical tooth prominent; subapical teeth small and subequal in length; ca. six spinous teeth, distal tooth markedly developed, others short; serration and sensilla well developed, slightly complex; blade region short and straight. Maxilla: palp cone-shaped 2.4–2.6× as long as basal width, very dark; hairs at base of palp not markedly developed, various. Postgenal cleft ( Fig. 188 View FIGURES 188, 189 ): markedly shallow with medial projection—variable; posterior tentoral pits distinct and rounded; postgenal bridge evenly pale contrasting with slightly darker genae; elongated posteroventral muscles spots distinct, but not markedly so; suboesophageal ganglion slightly pigmented and obvious. Hypostoma ( Fig. 189 View FIGURES 188, 189 ): well pigmented; tooth 0 distinct but not markedly prominent, teeth 1 & 2, small, tooth 3 larger (various, often absent from one side), tooth 4 well developed and flanged; teeth 5–7 small and various, tooth 8 small, obvious, and directed slightly laterally; ventral edge of hypostoma partially obscuring teeth 5–7; lateral edges of hypostoma straight, six to nine small lateral serrations, other smaller serrations more laterally; four or five substantial hypostomal setae on each side; ratio of hypostoma to genal bridge and postgenal cleft 1.0:1.3:0.3. Thorax: mottled grey; gill histoblast ( Fig. 190 View FIGURES 190–194 ) broadly L-shaped, with five primary trunks visible, directed ventrally, then posteriorly, then sharply anteroventrally; thinner pale tips of filaments directed anterodorsally. Prothoracic proleg ( Fig. 191 View FIGURES 190–194 ): lateral plates not markedly developed or L-shaped; lappets sometimes present as small clear tubercles. Abdomen: evenly expanded from anterior to posterior segments, not markedly expanded at segments VII & VIII; evenly mottled medium grey, paler anteriorly, darker posteriorly with some yellow. Rectal papillae: three simple lobes. Ventral tubercles: absent. Anal sclerite ( Fig. 192 View FIGURES 190–194 ): anterior arms slightly flared; main body of sclerite not well developed; posteromedial hole open, posterolateral arms absent; posteromedial extensions projecting from base of each posteroventral arm into the dorsal junction of hooks in circlet not markedly developed; posteroventral arms elongated and finely tapered; only five to eight campaniform sensilla between the posteroventral arms and circlet of hooks. Posterior circlet: well developed, ca. 150 rows of hooks with 17–23 hooks per row (total ca. 3,100).

First instar (one specimen). Body. Length 0.79 mm; head ( Fig. 193 View FIGURES 190–194 ). Labral fans: ca. 12 fan rays, length ca. 0.08 mm. Hypostoma ( Fig. 194 View FIGURES 190–194 ): tooth 0 prominent, teeth 1–3 subequal, tooth 4 slightly larger, teeth 5 & 6 present, teeth 7 & 8 barely evident; hypostomal groove distinct. Anal sclerite: barely evident, associated campaniform sensilla distinct. Circlet of hooks: with ca. 45 rows of hooks, with two or three hooks per row (total ca. 112).

Etymology. Named by Drummond (1931: 6) in honour of Australasian simuliidologist André Léon Tonnoir.

Types. Drummond did not designate a holotype, but labeled material as co-types. Subsequent labeling, probably by the Mackerras’, designated a holotype and an allotype. Bugledich (1999: 329) followed suit listing a male holotype, plus one male and four female paratypes. Under present Code, the former should be referred to as the ‘Lecotype’; the others as ‘Paralectotypes’.

Lectotype. Pinned male. Examined by LHG-A in 2007 and photographed ( Fig. 173 View FIGURES 173, 174 ). Type locality Western Australia, Lesmurdie (S31.9900° E116.0500°). 18 Oct. 1930. Coll. F. H. Drummond. (Fig. 195). Exact labeling not recorded. GoogleMaps

Paralectotypes. Two pinned specimens examined. Male, head missing. Label data:- [From pupa/ Lesmurdie/ 18. 10. 30 / Coll. F. H. D.] [Co-type Simulium / tonnoiri] [ Simulium /tonnoiri {M}/ Drum/ Paratype] [Red card] [Aust. Nat./ Ins. Coll.]. Now as microscope slide. Female, same labels, but with {F}.

Additional material. A single specimen is housed in the medical entomology collection, Westmead Hospital, Sydney. Label:- [Species number 01002B1H7] [Lab number (79-1640)] [ Cnephia tonnoiri tonnoiri ] [01/10/29,] [Tillyard] [Lesmurdie Plains, WA]. This predates Drummond’s collection and is assumed here to be the original discovery of the species, that which led Drummond to collect at Lesmurdie a year later. He made no comment. The specimen was not examined.

Alcohol material: All stages; collected variously by D. Bedo, H. & P. Zwick and JKM. [ ANIC Database No./ 29 026532–026539; 29 026746; 29 026750; 29 026847]; [ UASM #/ 370822–370826]. Slide mounts: All stages [ UASM #/ 370807–370821]

Bionomics. Drummond (1931: 8) and Mackerras & Mackerras (1949: 384) noted that A. tonnoiri breeds in a wide range of stream types. Drummond (loc. cit.) commented that pupae often form clumps with only the posterior tip of the cocoon attached—a behaviour reminiscent of pupae for the aurantiaca species-group. Behaviour of adults is unknown—we assume that the female is non-biting of humans, at least; an assumption supported partly by the markedly small teeth only on the medial side of the female mandible.

Prince (1980) in an ecological study of simuliids in Jane Brook, a small stream in the Darling Range, WA, found that larvae of Austrocnephia tonnoiri were present from March until December, with peak numbers in March. Fast flow and rocky substrate were attributes of the habitats. No adults were ever taken in the field. Austrosimulium furiosum , A. bancrofti and Simulium ornatipes occurred in the same stream reach. Simulium ornatipes was, however, largely separated in time, being more common in the summer and A. furiosum preferred the slower flows.

Distribution ( Fig. 197 View FIGURE 197 ). Western Australia. Near Perth, Bullsbrook, S31.6700° E116.0300°. 16 Aug. 1953. Four adults. Coll. D.L. McIntosh (ANIC). Jane Brook, S31.8808° E116.0862°. ( Prince, 1980). Walyunga National Park. S31.7300° E116.0600°. 26 May 2007. Larvae. Coll. L. Gil-Azevedo (ROM). Perth, S31.9400° E115.8500°. July 1930. Two adults. Coll. Drummond. 15 Nov. 1924. 20 adults. Coll. Nicholson (ANIC). South of Perth, Serpentine Falls, S32.3690° E116.0065°. 26 Oct. 2002. Larvae, pupae. Coll. Zwick (ANIC). Lesmurdie, S.31.9900° E116.0300°. 9 Dec. 1949. Larvae, pupae. Coll. D. Saunders (ANIC). Lesmurdie, S31.9900° E116.0300°. Oct. 1949. Pupae and larvae. Coll. Unknown (ANIC). Lesmurdie Fall, S31.9951° E116.0333°. Sept. 1929. Pupae. Coll. A. Tillyard (ANIC). Lesmurdie Plains, S32.0000 E116.0500. 1 Oct. 1929. Coll. Tillyard (WMH). Jarrah Forest, Brookton Highway, Route 40, Kangaroo Gulley, S32.1181° E116.1527°. 7 Nov. 1996. Larvae, pupae, reared adults. Coll. J.K. Moulton (UASM). Kelmscott, Route 40, Brookton Highway, Stoney Brook, S32.1250° E116.0472°. 14 Nov. 1996. Pupae, reared adults. Coll. J.K. Moulton (UASM). Near Perth, Canning Dam, Kangaroo Gulley, S32.1480° E116.1210°. 11 Jul. 1971. Larvae. Coll. D. Bedo (ROM). Wungong Brook, S32.1921° E116.0081°. Aug. Larva. Coll. Nicholson (ANIC). South of Perth, Serpentine River, S32.3700° E 116.0000°. Mar. 1972. Pupae. Coll. Zwick (ANIC). Pinjarra, east of Waroona. Lane Poole Reserve, Icy Creek, S32.8090° E116.0950°. Oct 2005. Larvae. Coll. Zwick (ANIC). Harvey, S33.0760° E115.9052°. Mar. 1972. Larvae. Coll. Zwick (ANIC). Margaret River, S33.9400° E115.0700°. 10 Nov. 1958. Adult. Coll. E.F. Riek (ANIC). NW Pemberton, Donnely River, S34.1070° E115.9900°. Mar. 1972. Larvae, pupae. Coll. Zwick (ANIC). Stirling Ranges, Moingup Springs, S34.4100° E 118.1500°. 28 Oct. 2005. Larvae, pupae. Coll. Zwick (ANIC). Beedelup National Park, Carey Brook, S34.4100° E115.8000°. Mar. 1972. Pupae. Coll. Zwick (ANIC). Beedelup National Park, Beedelup Falls, S34.4186° E115.8688°. 1972. Larvae. Coll. Zwick (ANIC). Beedelup Falls, S34.4186° E115.8688°. 13 Nov. 1958. Five adults. Coll. E.F. Riek (ANIC).

Prince (1980; her Fig. 3 View FIGURES 1–6 ) mapped some 70 localities for A. tonnoiri , from north of Perth to Stirling Range National Park, southern Western Australia, but detailed localities were not given.

Remarks. Mackerras & Mackerras (1949: 380, 384) originally considered material from Canberra and Coree Ck., ACT, as A. tonnoiri , but subsequently ( Mackerras & Mackerras, 1950: 170) restricted distribution to Western Australia and considered that the then Cnephia tonnoiri consisted of three subspecies, tonnoiri , orientalis and fuscoflava .

Rothfels (1979) in a cytological examination of Australian Austrocnephia (as Cnephia at the time), stated that A. tonnoiri occurred not only in Western Australia, but also Tasmania. That distribution has been cited since (e.g., Crosskey & Howard, 1997: 18). Of note was that A. tonnoiri consisted of two chromosomal siblings, but it is not clear if they were separate in Western Australia and Tasmania. This latter point is of biogeographical interest, since no material of A. tonnoiri is currently known from Tasmania, given the broadly-based collections (see Distribution(s) given) that have been made there. Austrocnephia orientalis is, however, widely distributed. Is it possible that A. orientalis of Tasmania is more closely related to the Western Australian A. tonnoiri than to A. orientalis on the neighbouring mainland? Certainly, larvae of Tasmanian A. orientalis are distinct from populations on the mainland, as noted previously.

Further, in none of the Mackerras’ work is C. tonnoiri recorded as occurring in Tasmania. Indeed, they comment that Cnephia material from the peninsula south of Adelaide, South Australia, was not as might be expected of C. tonnoiri tonnoiri from Western Australia, but was of the eastern C. tonnoiri orientalis . Prince (1980: 71) also does not report A. tonnoiri (as C. tonnoiri tonnoiri ) occurring in Tasmania. Thence, we are of the opinion that the records of A. tonnoiri in Tasmania are, for the present, in error. Bugledich (1999: 329) too has A. tonnoiri present in eastern Australia—again in error?

Colbo (1974) described and illustrated the head of a first instar larva of Cnephia tonnoiri orientalis —now Austrocnephia orientalis . That, similar to first instar A. tonnoiri , dealt with already, possesses labral fans, clearly indicating that, as he noted, these species are not Prosimuliini . The hypostoma is similar in first instar larvae of both species ( Figs. 161 View FIGURES 161, 162 , 194 View FIGURES 190–194 ), showing a developmentally plesiomorphic arrangement for those teeth. Similarly, Craig (1975: 466, 1997: 890) showed that highly apomorphic hypostomal teeth in Tahitian larval simuliids, have, in their first instar larvae, a very similar basic arrangement to that shown here—indicative perhaps that this is a plesiomorphic arrangement for Simuliini larval hypostomas? Be that as it may, Colbo (1974: 123) illustrated hypostomas of Austrosimulium bancrofti and Aust. pestilens first instar larvae, expressed both as in their respective mature larvae and not in the suggested plesiomorphic state as above.

Drummond (1931: 4) in discussing predators of Australian simuliids, commented that for A. tonnoiri , where the pupa are clumped and accumulate material, chironomid larvae were often present in and around the cocoons. He suggested that perhaps the chironomid larvae were preying on the simuliids, but in a footnote, comments that Tonnoir suggested the chironomids were merely commensals. We too have noted presence of chironomid larvae in the pupal mats and tend to agree with Tonnoir. Drummond (loc. cit.) further commented that cocoons of A. tonnoiri are very rough with a wide array of foreign material incorporated ( Fig. 183 View FIGURES 181–183 ). Also, that when the cocoons are clumped, often in groups of 10 or more, only the posterior of the cocoon is attached to the substrate (we too have noted this arrangement for aurantiaca and strenua ). Cocoons are usually found on stones, rarely on grass, in both fast and slow water (Fig. 195). Also noted was the dichromatic condition of the female eye (Fig. 166) and the marked pigmentation at the r-m veins junction ( Fig. 176 View FIGURES 175–179 ). That latter character, consists of campaniform sensillae located at the base of Rs, plus a concentration of pigmented hairs on R 1 and slight pigmentation of the wing membrane; producing the appearance of three closely adjacent pigment spots.