Danielsia Theobald, 1904

Reinert, John F., 2009, Descriptions of Danielsia Theobald, a genus recently resurrected from synonymy (Diptera: Culicidae: Aedini), and its type species Dn. albotaeniata Leicester, Zootaxa 1976, pp. 25-38 : 25-29

Genus Danielsia Theobald, 1904

Type species: Danielsia albotaeniata Leicester, 1904 Danielsia in part of Theobald (1907, 1910).

Culex in part of Brunetti (1914).

Aedes (Ochlerotatus) Group Finlaya in part of Edwards (1917).

Ochlerotatus in part of Brunetti (1920), Senior-White (1923, 1927).

Finlaya in part of Barraud (1923a, 1923b, 1924).

Aedes (Finlaya) in part of Edwards (1922a, 1922b, 1928), Brug (1931), Brug & Edwards (1931), Barraud (1934), Brug & Bonne-Wepster (1947), Chow (1949), Carter (1950), Chow & Mattingly (1951), Knight & Hull (1951), Chow et al. (1954), Horsfall (1955), Macdonald (1957), Feng (1958), Stone et al. (1959), Scanlon & Peyton (1965), Gould et al. (1968), Basio (1971), Stone & Delfinado (1973), Baisas (1974), Knight & Stone (1977), Jayasekera & Chellian (1981), Lu & Li (1982), Apiwathnasorn (1986), Tsukamoto et al. (1987), Lu & Su (1987), Lu et al. (1988), Harrison et al. (1990), Townsend et al. (1990), Ward (1992), Darsie et al. (1993).

Aedes (Finlaya) Group F ( albotaeniata -group: Danielsia ) in part of Edwards (1932).

Aedes (Finlaya) Group F, Division I- - albotaeniatus s.str. in part of Knight (1948).

Aedes (Finlaya) Group F ( alboannulatus -group: Danielsia ), Subgroup I, Albotaeniatus in part of Knight & Marks (1952).

Aedes (Finlaya) Group F ( alboannulatus -group) of Macdonald & Traub (1960), Lee et al. (1982).

Aedes (Finlaya) albotaeniatus group in part of Lu & Ji (1997).

Ochlerotatus (Finlaya) in part of Reinert (2000a), Kaur (2003).

‘ Ochlerotatus ’ (‘ Finlaya ’) sensu auctorum in part of Reinert et al. (2004, 2006).

Generic description

FEMALES. Head: Vertex nearly covered with broad, decumbent scales, specimens of some species also with few narrow, curved scales along coronal suture, few semi-erect to erect, forked scales posterior to ocular setae; occiput with several narrow, curved, decumbent scales and numerous erect, forked scales which may extend slightly onto posterior part of vertex; postgena with broad, decumbent scales; ocular line with pale scales and several setae on posterior margin; interocular space with pale scales and few setae; eyes above antennae contiguous to separated by less than 1 ocular facet; antenna dark, pedicel with moderate-sized patch of partially overlapping, moderately broad, pale scales on mesal surface; maxillary palpus dark-scaled, relatively short; proboscis dark-scaled, with pale-scaled area near midlength, longer than forefemur; clypeus without scales. Thorax: Integument dark brown; scutum covered with narrow, curved scales, median, prescutellar area bare, background scales dark with pale-scaled areas; acrostichal setae absent; dorsocentral setae present, 1 or 2 on caudal part of anterior area and several on posterior area; prescutellar area with several setae on lateral margins; scutellum with narrow, curved scales on all lobes (except Dn. harperi (Knight) with broad scales on mid-lobe); parascutellar area, mesopostnotum and paratergite without scales; antepronota widely separated, with broad scales and several setae; postpronotum with scales and few posterior setae; postspiracular area with scales (absent in Dn. harperi ) and few setae; subspiracular area with small patch of scales (absent in Dn. harperi ); proepisternum with broad scales and several setae on upper area, lower area bare; mesokatepisternum with broad, pale scales and several setae; prealar area with patch of broad, pale scales below knob and extending dorsally onto lower part of knob, with few to several setae; mesepimeron with patch of broad, pale scales and several setae on upper area, lower area bare; mesomeron bare, with dorsal margin well above base of hindcoxa; metameron bare. Wing: Dorsal and ventral veins dark-scaled except for pale-scaled patch on anterior surface at base of costa; alula with narrow scales on posterior margin; dorsal tertiary fringe scales present, dark; upper calypter with several dark setae on margin. Legs: Ante- and postprocoxal membranes bare; coxae with broad scales and several setae; femora dark-scaled with some pale-scaled areas; tibiae darkscaled with basal, pale-scaled area; mid- and hindtarsi dark-scaled with basal, pale-scaled bands on at least tarsomeres 1 and 2; fore- and midungues toothed, hindungues simple. Abdomen: Terga dark-scaled, laterotergite of tergum I with broad, pale scales, terga II–VIII each with basolateral, pale-scaled patch and narrow, basal, pale-scaled bands on at least terga IV and V; segments VII and VIII laterally compressed; intersegmental membrane between terga VII and VIII relatively short.

FEMALE GENITALIA. Tergum VIII: Width greater than length; covered with minute spicules; moderately pigmented; base very gently concave to nearly straight; apex broadly rounded, with several moderately long to long setae; setae present on distal 0.47–0.64 (rarely extending onto distal 0.66); basolateral seta very short; numerous broad scales densely covering distal 0.71–0.85; VIII-Te index 0.61–0.74; VIII-Te/IX-Te index 2.22–2.74; length 0.22–0.28 mm; width 0.33–0.46 mm. Sternum VIII: Width greater than length; covered with minute spicules; moderately pigmented; base relatively straight; apex gently sloping from apicolateral corners to midline, with several short, slightly curved setae and few slightly longer, nearly straight setae laterally; setae on distal 0.79–0.91; setae 1–5-S moderately long to long, in more or less angular or diagonal line extending from basomesal area to apicolateral area, seta 1-S inserted moderate distance from basal margin; basolateral seta normally absent; moderate number to numerous broad scales on distal 0.86–0.90; VIII-S index 0.76–0.94 (usually 0.85–0.94); length 0.28–0.38 mm; width 0.33–0.46 mm. Tergum IX: Moderately long and moderately wide; comprised of single, moderately pigmented sclerite, with pair of rounded apical lobes; covered with minute spicules; normally 1–3 (range 0–4, rarely absent on one side) short setae distally on each lobe, 1–7 total setae; IX-Te width/length ratio 0.85–1.17; length 0.09–0.12 mm; width 0.09–0.14 mm. Insula: Liplike; somewhat U-shaped (depressed) transversely; covered with minute spicules; moderately pigmented; with 3–5 (usually 3 or 4) moderately long setae laterally on each side, 6–10 (usually 6–8) total setae. Lower vaginal lip: Covered with minute to short spicules; lightly pigmented; narrow; hinge moderately wide; without lower vaginal sclerite; ventral tuft present, small. Upper vaginal lip: Covered with minute to short spicules; moderately pigmented; narrow laterally and curved outward, caudal part somewhat broader and flattened; upper vaginal sclerite moderately pigmented, small. Spermathecal eminence: Membranous; ill-defined. Postgenital lobe: Covered with short spicules; relatively narrow; apex flat; basal mesal apodeme weakly developed; few to several setae on distal 0.30–0.38 of ventral surface; PGL ventral index 2.50–2.81 (rarely 2.82); PGL ventral width/Ce dorsal width ratio 0.67–0.82. Proctiger: Membranous; with scattered minute spicules. Cercus: Covered with minute to short spicules; moderately long; moderately wide; apex gently oblique, with few moderately long, nearly straight setae and several short, gently curved setae; dorsal surface without scales (rarely with 1 adventitious scale on one cercus in Dn. harperi (Knight)) ; setae on distal 0.68–0.76 of dorsal surface; mesal margin broadly and gently concave; cercus index 2.50–2.95; Ce/dorsal PGL index 2.41–2.83; length 0.16–0.18 mm; width 0.06–0.07 mm. Spermathecal capsules: One large and 2 slightly smaller ones; heavily pigmented; spherical; with few small, spermathecal capsule pores near orifice. Accessory gland duct: Basal area lightly to moderately pigmented, short.

MALES. Differ from females primarily as follows. Head: Antennal flagellomeres with numerous long setae directed dorsally and ventrally, apical 2 flagellomeres with few short setae, both elongate but penultimate flagellomere noticeably longer; maxillary palpus dark brown-scaled, slightly shorter to slightly longer than proboscis, entire length slender, palpomere 1 very short, palpomeres 2 and 3 long, palpomeres 4 and 5 short but 4 longer than 5, distal part of palpomere 3 and entire length of palpomere 4 with numerous moderately long to long setae, palpomere 5 with several short setae ventrolaterally and at apex ( Dn. harperi with fewer setae on distal part of palpomere 3 and on palpomeres 4 and 5); proboscis dark-scaled with pale-scaled area near midlength or entirely dark brown-scaled. Thorax: Scutellum with narrow, curved, white scales on midlobe ( Dn. harperi with broad, white scales on midlobe). Wing: Dorsal tertiary fringe scales absent. Legs: Fore- and midtarsi each with 2 ungues, unequal in size, larger unguis with 2 teeth, smaller unguis with 1 tooth, hindtarsus with 2 ungues, equal in size, both simple.

MALE GENITALIA. Tergum IX: Covered with minute spicules; lightly to moderately pigmented; narrow mesal area separating moderately broad, lateral areas with outer parts projecting caudally presenting broad Vshaped appearance, each lateral part with very small, somewhat flattened, sclerotized, submedian lobe bearing 1 or 2 short, slender setae. Gonocoxite: Covered with minute spicules; moderately pigmented; moderately long; relatively narrow; broad, spatulate scales on outer dorsal, lateral and ventral surfaces; dorsal surface with basomesal area bearing compact patch of several to numerous relatively slender setae, some setae longer; lateral surface with long, stout setae; ventral surface with number of short to relatively long setae, mesal area near midlength bearing patch of long, moderately broad to broad, lanceolate scales, some lanceolate scales narrower and some shorter, some scales may be fusiform; mesal surface membranous. Gonostylus: Attached to apex of gonocoxite; relatively short; relatively narrow with distal part somewhat narrower; single, long, narrow, gonostylar claw attached at apex. Claspette: Comprised of single, moderately long, relatively narrow, columnar stem with few short setae, with several minute spicules on most of surface; claspette filament attached at apex, moderately long, flattened, expanded postbasally and tapering to narrow, bluntly rounded apex. Proctiger: Paraproct narrow, apex heavily pigmented, curved; cercus membranous with few minute cercal setae. Phallosome: Aedeagus moderately pigmented, simple, somewhat scoop-like, approximately proximal 0.75 broadly ovoid, distal part noticeably narrower; paramere heavily pigmented, narrow, shorter than aedeagus length. Sternum IX: Covered with minute spicules; moderately long; few short to moderately long setae on posteromesal area.

PUPAE. Cephalothorax: Seta 1-CT shorter than 3-CT; 2-CT shorter than 1,3-CT; 4-CT longer than 5-CT; 6-CT short; 7-CT long; 11-CT long, stout, single. Trumpet: Moderately long to long; tracheoid area at base of trumpet, poorly developed. Abdomen: Seta 1-I with several brush-tipped branches; 1-II with 2 slender branches; 2-II shorter than and inserted lateral to 1-II; 2-VI inserted mesal to 1-VI; 3-I very long, stout, single, longer than 6-I; 3-II single, longer than 6-II, inserted mesal to 2-II; 3-III stout, single, longer than 5-III; 3-VI inserted lateral to 1-VI; 5-II shorter than 3-II, inserted lateral to 4-II; 5-V very long, longer than mid-dorsal length of tergum VI; 6-III single; 6-VII shorter than and inserted posterior to 9-VII; 9-VIII with 6–12 stout, aciculate branches. Paddle: With apical margin rounded; without fringe of hairlike spicules; midrib extending to apex of paddle; seta 1-Pa 0.30–0.42 length of paddle; 2-Pa absent.

FOURTH-INSTAR LARVAE. Head: Seta 4-C branched, moderately long, inserted anterior and mesal to 6-C; 5-C long, single (rarely with 2–4 branches in Dn. albotaeniata ), inserted posterior and mesal to 6-C; 6-C moderately long, with 2–4 branches, inserted mesal and anterior to 7-C; 7-C moderately long, with 3–6 branches, inserted lateral and anterior to 5-C; 12-C branched, inserted mesal and posterior to 13-C; 13-C single; 14-C with 4–11 short, stellate branches; ventromedian cervical sclerite present. Antenna: Relatively short; without spicules; seta 1-A short, single to 3-branched. Thorax: Setae 1–3-P inserted on pigmented, common, setal support plate, 1,3-P branched, 2-P single; 4-P equal to or longer than 3-P; 5-P branched, longer than 6-P; 1-M 3.81–4.00 length of 2-M; 4-M single; 5-M longer than 7-M; 1,4-T stellate; 2,6-T single; 7-T long, stout, single, slightly longer than 6-T. Abdomen: Setae 1,2,5-I multiple-branched, stellate; 6-II with 2–4 branches, longer than 6-III; 6-III with 2 or 3 long branches; 7-II long, stout, single or 2-branched; 8-II branched; 1-VII relatively long; 2-VII branched, inserted relatively near 1-VII; 3-VII single to 3-branched, inserted mesal and anterior to 1-VII; 10-VII stellate; 12-VII single, inserted posterior to 13-VII; 1-VIII with 5–12 branches; 2,4- VIII single; comb comprised of 10–30 scales in patch ( Dn. albotaeniata ) or 6–8 scales in curved row ( Dn. harperi ); segment X with saddle incomplete ventrally, with well-developed spicules on posterior margin, acus absent; 1-X long, stout, single (rarely 2-branched), inserted on saddle; 2-X with 2 branches, 0.68–0.70 length of 3-X; 3-X single; ventral brush (seta 4-X) with setae borne on boss ( Dn. albotaeniata ) or grid with transverse bars ( Dn. harperi ), 2 shorter preboss/precratal setae. Siphon: Moderately long to relatively long; acus attached; pecten with distal 1–3 spines more widely spaced; seta 1-S with 2–6 branches, inserted distal to pecten; 2-S single; 9-S short, relatively straight.

EGGS. Unknown.

Included species. Danielsia albotaeniata , Dn. harperi and Dn. lepchana (Barraud) .

Material examined. Danielsia albotaeniata : female and genitalia, male and genitalia, pupa and fourthinstar larva; Dn. harperi : female and genitalia, male and genitalia, pupa and fourth-instar larva; and Dn. lepchana : male and genitalia. Material from the type series of each species was examined.

Distribution. India, Indonesia, Malaysia, Nepal, People’s Republic of China, Philippine Islands, Sri Lanka and Thailand.

Bionomics. Danielsia albotaeniata adults were reported by Leicester (1907) as bred from larvae collected in cut bamboos. Barraud (1934), Chow (1949), Macdonald (1957) and Gould et al. (1968) reported the occurrence of immature stages of this species in bamboos. Macdonald & Traub (1960) collected the immature stages of Dn. albotaeniata in Malaysia from treeholes and bamboos, i.e., bamboo pots, stumps, fallen, split, and large, moderate and small holes. Darsie et al. (1993) collected immature stages of this species from a reservoir with clear, fresh water at an elevation of 2,200 meters in Nepal. Macdonald (1957) reported that females of Dn. albotaeniata bite humans during the day in forested areas. Knight (1948) indicated that adults of Dn. harperi were reared from larvae collected in cut bamboo and bamboo stumps, Basio (1971) collected larvae from a bamboo stump, and Baisas (1974) reported the presence of immature stages in bamboos and the females as apparently zoophagous. Barraud (1923a) indicated that males of the type series of Dn. lepchana were reared from larvae taken from bamboo stumps.

Discussion. Theobald (1904) described the genus Danielsia and included a single species, Danielsia albotaeniata Leicester. The genus was later synonymized with other generic-level taxa as noted in the previous usage section above. Reinert et al. (2008) restored Danielsia to generic rank and included three species, i.e., albotaeniata , harperi and lepchana . Reinert (2008c) provided a generic description of the female genitalia and described and illustrated the female genitalia of the type species.

Danielsia albotaeniata was described by Leicester (in Theobald, 1904) from females and males, and later redescribed the species in 1907. Theobald (1907) provided a description of the male and female and included a partial illustration of the latter. Barraud (1934) described and illustrated the male and part of the genitalia and provided a brief description of the larva of this species. Chow & Mattingly (1951) illustrated the male genitalia and partially illustrated the pupa. Edwards (1922a) described mikiranus as a new species and provided information on only two adult characters. Edwards (1922b) and Barraud (1923) considered it a valid species and the latter author described and illustrated the female, but Barraud (1934) reduced it to a variety of albotaeniata . Subsequent authors considered it a variety of albotaeniata , including Chow & Mattingly (1951) who described the fourth-instar larva and pupa and partially illustrated these life stages. Lu & Su (1987) and Lu & Ji (1997) illustrated the male genitalia and fourth-instar larva of mikiranus (as a subspecies of albotaeniata ). Harrison et al. (1990) considered mikiranus as nothing more than a clinal variation of albotaeniata and formally synonymized it with albotaeniata .

Knight (1948) described Dn. harperi (as Aedes harperi ) based on the male and its genitalia, the female, and the fourth-instar larva. Also, he included illustrations of the head and terminal segments of the larva, and male genitalia. Illustrations of the female and descriptions of the female, larva and pupa were given by Baisas (1974).

Danielsia lepchana was briefly described originally from males by Barraud (1923a) but he later (1934) listed this species as a synonym of albotaeniata . Knight (1948) listed it as a valid species, but noted it was possibly a synonym of albotaeniata . Knight & Marks (1952) and Knight & Stone (1977) listed both albotaeniata and lepchana as valid species. Reinert (2008b) redescribed the holotype male and genitalia of Dn. lepchana in detail.

The following combination of morphological features is most distinctive for Danielsia : females with occiput bearing numerous erect, forked scales, eyes contiguous to separated by less than diameter of one ocular facet, scutum with narrow, curved scales, white-scaled patch on anterior portion of acrostichal area or expanded and covering acrostichal and dorsocentral areas, acrostichal setae absent, paratergite without scales, mesepimeron without lower setae, costa with white-scaled patch at base of anterior surface, mid- and hindtarsi dark-scaled with basal, pale-scaled bands on at least tarsomeres 1 and 2, and abdominal segments VII and VIII laterally compressed; female genitalia with insula liplike, somewhat U-shaped (depressed) transversally and with 3–5 moderately long setae laterally on each side, lower vaginal lip lightly pigmented which sharply contrasts with moderately pigmented insula, cercus moderately long, moderately wide, apex gently oblique, sternum VIII with moderate number to numerous scales on most of surface, apex gently sloping from apicolateral corners to midline, and upper vaginal sclerite small; males with maxillary palpus dark-scaled, slightly shorter to slightly longer than proboscis, entire length slender, palpomeres 4 and 5 short, fore- and midtarsi each with two ungues, unequal in size, larger unguis with two teeth, smaller unguis with one tooth, hindtarsus with two simple, equal ungues; male genitalia with tergum IX comprised of narrow, median area separating moderately broad, lateral areas with outer parts projecting caudally and presenting V-shaped appearance, one or two short, slender setae inserted on very small, somewhat flattened, sclerotized, submedian lobe on posterior margin of each side, gonocoxite relatively narrow, dorsal surface with basomesal area bearing compact patch of relatively slender setae, some setae noticeably longer, ventral surface with mesal area near midlength bearing patch of long, moderately broad to broad, lanceolate scales, claspette comprised of single, moderately long, columnar stem and bearing apically one moderately long, flattened filament that is expanded postbasally and tapered to narrow apex, cercal setae present, and aedeagus simple, somewhat scooplike with approximately proximal 0.75 broadly ovoid, distal part noticeably narrower; pupae with seta 1-CT shorter than seta 3- CT, seta 4-CT longer than seta 5-CT, seta 6-CT noticeably shorter than seta 7-CT, seta 1-II with two slender branches, inserted mesal to seta 3-II, seta 3-I very long, stout, single and longer than seta 6-I, seta 3-VI inserted lateral to seta 1-VI, seta 9-VIII with 6–12 stout, aciculate branches, and paddle with apical margin rounded, without fringe of hairlike spicules, midrib extending to apex; fourth-instar larvae with seta 4-C branched, moderately long, inserted anterior and mesal to seta 6-C, seta 5-C long, normally single, inserted posterior and mesal to seta 6-C, seta 6-C with 2–4 moderately long branches, inserted mesal and anterior to seta 7-C, seta 13-C single, antenna relatively short, without spicules, setae 1–3-P inserted on common, setal support plate, setae 2,6-T single, numerous stellate setae on thorax and abdomen, seta 6-II, III branched, seta 7-II long, stout, single or 2-branched, setae 12-VII, 2,4-VIII single, segment X with saddle incomplete ventrally, with well-developed spicules on posterior margin, acus absent, seta 1-X long, stout, single (rarely 2- branched), seta 2-X moderately long, 2-branched, and siphon with attached acus, pecten with distal 1–3 spines more widely spaced, seta 1-S with 2–6 branches, inserted distal to pecten. The above data is based on adults and immatures of Dn. albotaeniata and Dn. harperi , but only the male of Dn. lepchana .