Culex (Culex) dolosus, (Lynch Arribalzaga) (Lynch Arribalzaga, 1891)

Senise, Luana Valente & Sallum, Maria Anice Mureb, 2008, Redescription of Culex (Culex) dolosus (Lynch Arribálzaga) (Diptera: Culicidae), based on specimens from Pico do Itapeva, Serra da Mantiqueira, São Paulo, Brazil, Zootaxa 1683, pp. 51-62 : 52-60

Taxonomic treatment

Heteronycha dolosa of Lynch Arribálzaga 1891: 156.

Culex (Culex) dolosus ( Lynch Arribálzaga, 1891) . Lectotype, adult male, deposited in MACN. Type locality: Buenos Aires Province, Argentina

Culex (Culex) dolosus of Dyar 1922: 3 ( Heteronycha as synonym of Culex ). Dyar 1928: 374, 375 (M, F, L, G). Lane 1953: 362–364 (M, F, P*, L*, G*, in part). Duret & Barreto 1956: 84 (taxonomy). Forattini 1965: 155–157, 198, 204 (M, F, P, L*, G*, in part). Forattini & Rabello 1965: 33–34 (taxonomy). Bram 1967: 59, 60, 61, 117 (L, G*, in part). Casal in Belkin et al. 1968: 15 (lectotype design.).

Culex (Culex) bilineatus Theobald, 1903: 196–198 . Lectotype adult male, deposited in the NHM. Type locality: São Paulo municipality, São Paulo State, Brazil. Peryassú 1908: 190, 191 (M, F). Lane 1951: 334 (syn. design.). Belkin 1968: 13 (lectotype design.).

Culex (Culex) bonariensis Brèthes, 1916: 213 , 214 (M, F, G*). Lectotype adult male, deposited in the MACN. Type locality: San Isidro municipality, Buenos Aires Province, Argentina. Dyar 1928: 374 (syn. design.). Casal in Belkin et al. 1968: 15 (lectotype design.).

Female. Pale scales varying from white to yellow; dark scales varying from dark yellow to dark brown either with golden or blue sheen; setae mainly dark brown with golden sheen, except when indicated. Head. Integument dark brown; interocular space width 0.06–0.09 mm (mean = 0.08 mm); vertex with narrow, decumbent pale scales, erect forked dark brown scales, and long dark brown setae; occiput with erect forked dark brown scales, medially with erect pale forked scales, and narrow decumbent pale scales; broad appressed white scales above ocular suture extending to postgena; ocular setae dark brown; postgenal setae golden yellow; gena, palpifer and clypeus bare. Antennal length 1.85–2.15 mm (mean = 1.95 mm); pedicel bare, integument dark yellow with brownish areas; flagellum dark brown, flagellomeres 1–13 covered with minute whitish setae, flagellomere whorls with long dark setae, flagellomere 1 1.02–1.15 (mean = 1.10) length of flagellomere 2. Maxillary palpus length 0.40–0.46 mm (mean = 0.42 mm), ratio of length of palpomeres 1–4 to total length of palpus, 1 = 0.12–0.18, 2 = 0.10–0.11, 3 = 0.19–0.23, 4 = 0.49–0.56, palpomere 2 0.44–0.57 (mean = 0.50) length of palpomere 3, palpomere 3 0.33–0.48 (mean = 0.37) length of palpomere 4, maxillary palpus 0.22–0.25 (mean = 0.23) forefemur length, 0.19–0.24 (mean = 0.22) proboscis length, palpomeres dark scaled with few small dark setae, palpomeres 2 and 3 with a pair of long dark brown setae on each side. Proboscis dark scaled, length 1.9–2.2 mm (mean = 2.1mm), labial basal setae dark brown. Thorax. Scutal integument brownish along acrostichal area, dorsocentral area, transversal suture and lateral area of scutal fossa; with a pair of slightly darker stripes between acrostichal and dorsocentral areas and slightly darker spots at middle of scutal fossa, anterolateral area of scutum, prescutelar area, between the antealar/supraalar areas and at posterior dorsocentral area; paratergite, antealar and supraalar areas light yellow to light brown; scutal setae: acrostichal, anterior dorsocentral, anterior scutal fossal, lateral scutal fossal, posterior dorsocentral, posterior medial scutal, posterior scutal fossal, prescutellar, median anterior promontory, antealar and supraalar setae all dark brown with golden sheen; median scutal fossal setae absent; scutal scales: acrostichal, anterior dorsocentral, posterior dorsocentral, posterior scutal fossal, lateral scutal fossal, median scutal fossal and median prescutellar narrow, small and yellowish, with golden sheen; prescutellar, antealar, supraalar, anterior dorsocentral and posterior scutal fossal scales slightly more developed, narrow, pale yellow to whitish; center of prescutellar area bare. Scutellar integument light brown, lighter at midlobe, 5–7 (mode = 6) lateral scutellar setae and 2–5 (mode = 4) median scutellar setae, very long and dark brown, lateral and median scutellar scale whitish, narrow and long. Mesoposnotum bare, integument dark brown medially and light brown laterally. Pleural integument yellowish at middle and lower posterior mesokatepisternum and slightly lighter at anepisternal cleft; whitish at postprocoxal membrane, proepimeron, medial proepisternum, upper anterior of mesokatepisternum, medial and lower posterior of mesanepimeron, mesomeron, metepisternum, metepimeron and metameron; dark yellow to pale brown with medial whitish at subspiracular and hypostigmal areas; darker areas, dark to light brown, at antepronotum, postpronotum, upper and lower proepisternum, upper posterior and lower anterior of mesokatepisternum and upper and lower of mesanepimeron; postpronotal setae dark brown, usually fourth and very long, and following groups of pale setae: antepronotal, postpronotal, proepisternal, upper and lower mesokatepisternal, basalare, several upper mesepimeral and one lower mesepimeral; white and broad spatulate scales: lower prealar, lower and upper mesokatepisternal, upper and anterior mesepimeral; white and narrow spatulate scales at antepronotum; mesokatepimeron, mesomeron, metameron, metepisternum, metepimeron and subspiracular area bare, pre and postspiracular scales and prespiracular setae absent. Halter . Capitellum, scabellum and pedicel integument pale yellow with brownish areas; capitellum apex and outer surface with pale pearly spatulate scales, inner surface bare. Abdomen. Integument pale yellow with some brownish areas at tergum, tergum I dark brown; pale scales pearly and dark scales with blue sheen; length 3.10–3.80 mm (mean = 3.40 mm); tergum and sternum I without scales, only with dark setae; terga II–VII dark scaled with broad basal white bands, smaller on tergum VII, and dark yellow setae; sterna II–VII also with dark setae, scale pattern similar to terga, sternum V–VI with broader basal white bands, sternum VII almost all white scaled; presence of midlongitudinal band of dark scales apically, progressively broader until the apex of sternum VII. Segment VIII. Tergum white scaled with slender apical dark band, sternum white scaled. Legs. Coxal and trochanter integument mostly whitish with pale scales and long and short dark seta; pale to dark brown regions at anterior forecoxa and foretrochanter; femoral bases with pale spots, sometimes as a ring, apex pale; forefemur dorsal, anterior and anteroventral surfaces dark, posterior and posteroventral surfaces pale; midfemur dorsal, upper anterior and upper posterior surfaces dark, ventral, lower anterior and lower posterior surfaces pale; hindfemur similar to midfemur, anteriorly also with dark band distally; fore-, mid- and hindtibia dark scaled, posteriorly pale, apex with pale ring; foretarsomeres 1–5 dark with pale scales, mostly at joints, that might or might not form band at apex and base of tarsomeres, midtarsomeres 1–5 dark, apex of 1, apex and base of 2, 3 and 4 and joints 1–2, 2–3, 3–4 and 4–5 with pale scale bands, hindtarsomeres similar to midtarsomeres, with pale scales bands more apparent on base of hindtarsomere 1. Wing. With dark spatulate scales, dark brown to black, length 3.30–3.85 mm (mean = 3.60 mm); veins C, Sc and CuA covered with broad scales; veins, R 1 and CuA 2 mostly with broad scales and with some narrow scales; veins CuA 1, Rs and M mostly with narrow scales and some broad scales; 1A, R 2, R 3, R 4+5, M 1+2 and M 3+4 with narrow scales.

Male. As in female, except for the following sexual differences. Head. Palpomere 3 entirely dark brown, with broad apex and long dark setae, from middle to apex, at ventral and outer surfaces, joint of palpomeres 3–4 with tuft of white scales, more evident at inner surface, palpomere 4 covered with dark long setae, dorsal, ventral and outer surfaces dark, inner surface dark with pale longitudinal stripe, broader near base, joint 4–5 with tuft of white scales, palpomere 5 dorsal, ventral and outer surfaces dark, inner surface dark with pale spot near base. Antennal length 1.43–1.65 mm (mean = 1.55 mm), male pedicel diameter approximately 1.3 female pedicel diameter, filament integument pale, flagellar whorl dark brown, with tufts of very long dark setae, shorter on apex, flagellomere 14 longer than 12 and 13 and covered with minute white setae, antenna apex with pair of long dark setae. Proboscis with a ventromedial pale band, usually forming a ring. Abdomen. Terga II–VII with dark setae mostly at apex and margin, scale arrangement similar to female terga; sterna integument lighter and greenish, sternum I with few long setae, II and III bare, IV–VII white scaled with midlongitudinal dark stripe and long dark setae along surface. Legs. Foretibia apex with pale spot dorsally, anteriorly and posteriorly; hindtarsomeres 1–5 dark with pale ring at joints, more evident in 5, claws simple. Genitalia ( Figs. 1A–D ). Gonocoxite ( Fig. 1A ). Covering of short spicules and long setae, two setae near gonostylus base; subapical lobe prominent and undivided with set of small setae on base and three longer setae above that; setae a–c, f and h as a rod, a–c stout, a slightly rounded at apex, b and c with hooked apex, f and h slender, f hooked at apex, h pointed at apex and bending medially, g leaf form, with lanceolate shape. Gonostylus with two small and slender setae on dorsal surface, gonostylar claw small, with sides folding toward medial surface, acquiring the shape of a tent. Gonocoxite length (dorsally, from acetabulum to gonostylus base) 0.28–0.33 mm (mean = 0.30 mm), gonostylus length (dorsally, from base to base of gonostylar claw) 0.15–0.17 mm (mean = 0.16 mm), gonostylar claw length 0.011 –0.015 mm (mean = 0.013 mm). Segment IX ( Fig. 1B ). Ninth tergal lobe narrow, rounded laterally, with spiculed integument and 5–9 (mean = 5) long setae arranged in a single straight row, that might be unevenly-spaced when in bigger number, medially with lowered hyaline region, similar to a membrane; rectangular sternal lobe, with spicules arranged in horizontal lines. Proctiger ( Fig. 1C ). Paraproct with basal lateral arm long and curved and prominent acetabulum as small lobe, both strongly sclerotized; paraproct crown with row of 4–6 (5) stout and sharply pointed spines, moderately sclerotized, remaining spines slender, longer, heavily sclerotized and sharply pointed; cercal sclerite elongate; presence of 3–4 (mean = 3) cercal setae; X–T slightly rectangular. Phallosome ( Fig. 1D ). Lateral plate with 3–6 (4) large and 1–2 (1) smaller, strongly sclerotized teeth, and a flat lateral lobe, base of lobe continuous with base of dorsal process, lateral lobe similar in shape to lateral teeth, less sclerotized, directed laterally; dorsal process a strong hook, projected dorsomedially, broad at base, tapering to apex and ending in a sharp point, strongly sclerotized; ventral arm bend dorsomesad forming a somewhat right angle on dorsal surface, connected with a ventral projection, forming a somewhat triangular, moderately sclerotized ventrolateral lobe; dorsal arm flattened, large, tapered distally, moderately sclerotized, extending beyond ventral arm; aedeagal sclerite and paramere strongly sclerotized, basal piece moderately sclerotized.

Larva ( Figs. 2A–C ). Position and development of setae as figured; range and modal number of branches in table 1. Head ( Fig. 2A ). Wider than long, length and width not measured, moderately pigmented, yellowish, darker on median labral plate and posterior area of lateralia, dark yellow; collar heavily tanned; dorsomentum with large median tooth and 5–10 (mean = 6) smaller teeth on either side; seta 4–C extending beyond insertion of 7–C, setae 5,6,7–C with aciculate branches. Antenna weakly pigmented, pale yellow, darker mesally, above 1–A insertion and on antennal base; length 0.60–0.77 mm (mean = 0.71 mm), width 0.06–0.08 mm (mean = 0.07 mm), 8.70–12.40 (mean = 10.40) longer than wide; 1–A inserted 0.44–0.54 mm (mean = 0.50 mm) from base with 20–30 strongly aciculate branches, ratio of distance 1–A to base/ antennal length 0.63–0.73 (mean = 0.70). Thorax ( Fig.2B ). Integument hyaline with tiny spicules, more evident dorsally and laterally; 0–P symmetric and long, 1–8,10-P with aciculate branches, 5–10,12-M with aciculate branches, 4-M long; 1-T symmetric and long, 4-T forked and long, 7,9,10,12-T with aciculate branches. Abdomen ( Figs. 2B–C ). Integument hyaline, spicules smaller than those on thorax, more distinct on segment VIII; setae 1-I and 1-II long. Segment VIII ( Fig. 2C ). Comb with 32–57 short scales, fringed on sides and apex, apical fringe more distinct. Siphon ( Fig. 2C ). Index 5.40–7.90 (mean = 6.10), moderately pigmented, brownish, strongly darker on acus, base, spiracular apparatus, spiracular apodeme and apex; spiracular apodeme ratio approximately 0.25; pecten on base 0.25–0.40 (mean = 0.33) of siphon, with 10–15 pair of spines, usually with two teeth; seta 1-S laterally inserted near most distal pecten spine, length of setae exceeding the width of siphon at its insertion. Segment X ( Fig. 2C ). Saddle complete, spiculate, moderately tanned, slightly darker on base and apex; length 0.36–0.52 mm (mean = 0.43 mm), siphon/ saddle index 3.30–5.30 (mean = 4.0).

Pupa ( Figs. 3A–B ). Position and development of setae as figured; range and modal number of branches in table 2. Cephalothorax ( Fig. 3A ). Wing cases, antennal cases, head shield and middorsal ridge weakly pigmented, pale yellow, mouthparts cases and metanotum slightly darker than the former, posterolateral region, above wing cases, with yellowish spot; trumpet laticorn, weakly pigmented medially and darker, at tracheoid area and apex, apex somewhat chalice-shaped. Trumpet index 7.0–10.0 (mean = 8.60), pinna 0.15–0.22 (mean = 0.18) length of meatus, tracheoid length 0.27–0.37 mm (mean = 0.31 mm); 1-CT very long, 12-CT longer than 10, 11-CT, 10-CT usually simple, rarely with aciculate branches. Abdomen ( Fig. 3B ). weakly pigmented, pale, yellow, segment I, between setae 1-I insertion darker; anteromedial portion of tergum and sternum strongly sclerotized and strongly pigmented, brownish, particularly sternum II–V; abdomen length 2.5–3.30 mm (mean = 2.90 mm), ratio of length of segment III/II 1.05–1.35 (mean = 1.17), IV/III 1.05–1.25 (mean = 1.10), V/IV 1.01–1.20 (mean = 1.10), VI/V 1.00–1.15 (mean = 1.10), VII/VI 1.00–1.15 (mean = 1.10), VIII/ VII 0.70–0.80 (mean = 0.75),VIII/VI 0.70–0.85 (mean = 0.80), VIII/II 1.00–1.30 (mean = 1.20); seta 1-I dendritic and well developed, 5-IV-VI very long, exceeding the inferior margin of segment, 5-VII shorter than the other, 6-I,II very long, 6-VII usually simple but occasionally with aciculate branches, 9-II very small, inserted close to 6-II, 9-VIII long, well developed and moderately pigmented, with aciculate branches. Genital lobe ( Fig. 3B ). Lightly tanned in female, slightly darker in male, male genital lobe approximately 1.60 length of female genital lobe. Paddle ( Fig. 3B ). Almost hyaline, except for external buttress, midrib and base, lightly tanned; length 0.70–0.90 mm (mean = 0.80mm), width 0.50–0.65 mm (mean = 0.60 mm), paddle index 0.60– 0.75 (mean = 0.70).

Material examined. Twenty four specimens of Cx. dolosus were examined, as follow. BRAZIL, State of São Paulo, Pindamonhangaba, Pico do Itapeva (22º 45.50´S 45º 30. 87´W), Sallum and Wilkerson coll. 20- XI-2001, Sallum det. 2002: E-12375 M LePe G, E-12376 M Pe G, E-12377 F LePe, E-12378 F LePe, E-12381 M Le G, E-12382 F LePe, E-12383 M LePe G, E-12385 M LePe G, E-12390 M LePe G, E-13391 M LePe G, E-12394 F LePe, E-12395 M LePe G, E-12396 F, E-12397 M LePe G, E-12398 M LePe G, E-12405 F Pe, E- 12408 F Pe, E-12414 Pe; Sallum et al. coll. 17-II-2006, Senise det. 2006: E-12924 Le, E-12925 Le, E-12926 Le, E-12927 LePe; Lane coll. XII-1936, Lane det., accession n o 4779 G, n o 4780 G. Other material examined: Cx. (Cux.) eduardoi , BRAZIL, State of São Paulo, São Paulo municipality, Bairro do Limão , Susan coll.; VI- 1973, Sallum det. 2007: E-5021 LePe G, E-5022 LePe G; Pariquera-Açú municipality, Sítio Galiléia, Gomes and Kakitani coll. 1985, Sallum det. 2007: E-7651 LePe G, E-7653 G; Iguape municipality, Sítio Palmeiras, Gomes coll. 22-II-1989, Lab. Ent. FSP-USP det.: E-8183 PeLe G, E-8080 LePe; State of Paraná, Cerro Azul municipality, Natal coll. 16-VI-1991, Natal det. 1991: E-9050 L; 17-VI-1991, Sallum det. 1992: E-9060 L.

Distribution. The geographical distribution of Cx. dolosus , i.e. Bolivia, Brazil, Chile, Ecuador, Paraguay, Uruguay and Argentina ( Bram 1967; Knight & Stone 1977), is probably an overestimate because of misidentifications.

Bionomics. Immature stages of Cx. dolosus have been found in natural and artificial habitats, such as cans, permanent or temporary ponds, with clear or turbid, stagnant water, and either associated or not associated with aquatic vegetation. In urban areas in Argentina, immatures were taken from non-polluted water ( Almirón & Brewer 1996; Maciá 1997; Fischer & Schweigmann 2004).

Pico do Itapeva, Pindamonhangaba municipality is situated in high altitude and upland forest vegetation. In a field collection from 2006, immatures of Cx. dolosus were found in a car track, temporary ground pool. The water was fresh, stagnant, unpolluted, with emergent grass and green algae, in the shade or in full sun. Immatures of Cx. dolosus were associated with those of Cx. (Cux.) coronator Dyar & Knab, Cx. (Melanoconion) pilosus (Dyar & Knab) , and Ae. (Ochlerotatus) crinifer (Theobald) .

Lourenço-de-Oliveira et al. (2004) collected adult females of Cx. dolosus in peridomiciliary areas. The medical importance and feeding behavior of the species is unknown.

Discussion. Specimens from Pico do Itapeva, Pindamonhangaba can be identified as either Cx. dolosus or Cx. eduardoi based on the following characteristics of the male genitalia: ventral arm of the lateral plate distinct from the teeth of the lateral arm, with pointed apex and bending laterally in a right angle; the dorsal process is strong at base, tapering to apex, ending in a pointed apex, distinctly bent dorsoposteriorly, forming a strong pointed hook-like structure. However, adult females can be identified as Cx. dolosus by having wing scales entirely dark, postspiracular scales absent, scutum with antealar spot of yellowish scales, basal white bands on abdominal terga, and pale scales on the joints of fore-, mid- and hindtarsomeres 1–5, which are more apparent in the hindtarsomeres. Darsie (1985) and Forattini (2002) indicated that pale scales are either absent or present in Cx. dolosus but, when present, are restricted to narrow bands or patches at tarsomere joints. It is plausible to hypothesize that the polymorphism noted by these authors indicates that more than one species has been misidentified as Cx. dolosus .

Considering the fourth instar larval characteristics, Dyar (1928) postulated that Cx. dolosus could be distinguished from other species of the Culex (Culex) by a siphon that is approximately 6 times as long as the basal width, and the presence of 4 pairs of multibranched seta 1-S. Darsie (1985) and Forattini (2002) used the siphon index (ratio of the length at dorsal surface / width at base)> 5.0 and the presence of 4 pairs of seta 1-S as characteristics to distinguish Cx. dolosus from other Culex (Culex) species. Larvae of Cx. dolosus from Pico do Itapeva, Pindamonhangaba have a siphonal index of approximately 6.10, and 4 to 5 pairs of seta 1-S. In addition, the length of the most basal 1-S seta exceeds the width of the siphon at the seta insertion.

The pupae of Cx. dolosus seem to be similar to those of other Culex species. However, it is noteworthy that setae 1-CT, 5-IV-VI and 6-I, II are very long, the trumpet is darkly pigmented at the apex and tracheoid area, and weakly pigmented medially, with the apex somewhat chalice-shaped.

Culex dolosus is morphologically similar to Cx. eduardoi . The adult male of Cx. dolosus can be distinguished from that of Cx. eduardoi by the presence of a pale ring on proboscis, which is absent in Cx. eduardoi ; the adult female by the presence of basal white bands on abdominal terga, and pleural integument with dark areas. Additionally, the presence of antealar spots of yellowish scales is usually used to separate Cx. dolosus from other species of the subgenus Culex , including Cx. eduardoi . According to the original description of Cx. eduardoi (Casal & García 1968) , the scutal scales are mostly brown with a reddish sheen and there is no mention to the presence of yellowish scales in the antealar area of scutum. Casal & García (1968) also reported the presence or absence of postspiracular scales in the male of Cx. dolosus , which are always absent in Cx. eduardoi . In the specimens of Cx. dolosus from Pico do Itapeva, Pindamonhangaba, the postspiracular scales are always absent. Culex eduardoi and Cx. dolosus are indistinguishable by male genitalic characteristics. The fourth instar larva of Cx. dolosus can be distinguished from those of Cx. eduardoi by the following characters: setae 1-T and 1-I longer, dorsomentum narrower with more developed teeth, thoracic and abdominal integument spiculate (vs. glabrous or sparsely spiculate in Cx. eduardoi ), and 4, 5 pairs of seta 1-S ( Cx. eduardoi has 6 pairs). Knight & Laffoon (1971) postulated that some Culex species possess setae 16-C and 17-C, a pair of small seta inserted close to the postocciput. Casal & García (1968) noted the presence of setae 16-C and 17-C in fourth instar larvae of Cx. dolosus , and thus used this character to separate Cx. dolosus from Cx. eduardoi . However, no specimens of Cx. dolosus from Pico do Itapeva, Pindamonhangaba possess these setae. Casal & García (1968) also noted the absence of spicules on the thoracic integument of fourth instar larvae of Cx. eduardoi , whereas this structure is spiculate in Cx. dolosus . In contrast to Casal & García, we recorded minute, sparse spicules on the thoracic integument of fourth instar larva of Cx. eduardoi . Additionally, setae 0-P, 4-M, 1-T, 4-T and 1-II are longer in Cx. dolosus from Pico do Itapeva, Pindamonhangaba than those of Cx. eduardoi .

Culex bilineatus was described by Theobald (1903) based on specimens collected in São Paulo municipality, Brazil, and subsequently synonymized with Cx. dolosus by Lane (1951). In comparing adults of Cx. eduardoi and the description of adult Cx. bilineatus , we noticed that both have the proboscis entirely dark scaled, whereas those of Cx. dolosus have a pale ring approximately at the middle of the proboscis. By examining specimens collected in São Paulo municipality that were identified as Cx. dolosus , we noticed they belong to Cx. eduardoi . Because both Cx. bilineatus and Cx. eduardoi possess a proboscis that is entirely dark, we hypothesize that Cx. bilineatus is conspecific with Cx. eduardoi rather than Cx. dolosus . However, it is premature to propose any taxonomic change without examining additional specimens of Cx. eduardoi from the type localities in Argentina and to compare them with the type specimen of Cx. bilineatus .

Figures

FIGURE 1. Cx. dolosus, male genitalia. (A) Gonocoxite (Gc), gonostylus (Gs), gonostylar claw (GC), setae a–c and f–h. (B) Sternum IX (IX–S), tergum IX (IX–T). (C) Paraproct (Ppr), paraproct crown (PpC), cercal seta (CSe), cercal sclerite (CSc), basal lateral arm (BLA), tergum X (X–Te), acetabulum (a). (D) Phallosome: aedeagal sclerite (AeS), dorsal arm (DOA), ventral arm (VA), lateral lobe (LL), ventrolateral lobe (VLL), dorsal process (DP), basal piece (BP). Scales are in mm.

FIGURE 2. Cx. dolosus, fourth-instar larva. (A) Head, left side dorsal, right side ventral. (B) Pro- (P), meso- (M) and metathorax (T), and abdominal segments I–VI, left side dorsal, right side ventral. (C) Abdominal segments VII–X side view, spiracular apparatus (S), comb spines (CS), pecten spines (PS). Scales are in mm.

FIGURE 3. Cx. dolosus, pupa. (A) Cephalothorax (CT). (B) Metathorax, abdominal segments I–IX, left side dorsal, right side ventral, paddle (P), genital lobe (GL). Scales are in mm.