Journal of the American Control Association, 19(4):309-322,2ffi3 Copyright O 2003 by the American Mosquito Control Association, Inc.

DESCRIPTION OF BRUCEHARRISONIUS,A NEW SUBGENUS OF OCHLEROTATUS.AND A REDESCRIPTION OF ITS TYPE SPECIES OC. (BRH) GREENII

JOHN F REINERTI

Center for Medical, Agricultural and Veterinary Entomology (CMAVE), United States Department of Agriculture, Agricultural Research Service, 1600/17OO SW 23rd Drive, Gainesville, FL 32608-1067

ABSTRACT, The new subgenus Bruceharrisonizs is described and principal features of all known stages are provided for separating it from other subgenera of Ochlerotatus. The female, male, female and male genitalia, pupa, and 4th-stage larva of the type species, Oc. greenii, are described in detail. Eight species (Oc. alektorovi, Oc. aureostriatus, Oc. doonii, Oc. christophersi, Oc. greenii, Oc. hurlbuti, Oc. okinawanus, and Oc. taiwanus) currently are recognized in the subgenus.

KEY WORDS Brucehanisonius, Ochlerotatus, Oc. greenii, Culicidae, Diptera, new subgenus

INTRODUCTION SUBGENUS BRUCEHARRISONIUS REINERT. NEW SUBGENUS

During studies on the genera and subgenera in Type speciesz Howardina greenii Theobald, tribe , family Culicidae, a new subgenus, 1903 Brucehamisonius, was discovered and is described below. The subgenus is included in genus Ochler- Culex, in part of Doleschall (1857), Giles (1900, otatus Lyncla Aribalzaga (genus recently restored 1902), Theobald (1901). to generic rank by Reinert [2000al) and includes Howardina, in part ofTheobald (1903, 1905, 1907, species formerly placed in subgenus Finlaya Theo- 1910), Blanchard (1905), Bruneui (1907, 1912, bald. Species included in the new subgenus are Oc. 1920). ale kto rovi (Stackelberg), O c. aure o striatas (Doles- (Ochlerotatus) grotp Finlaya, in part of Ed- chall), Oc. doonii (Wattal Bhatia, and Kalra), Oc. wards (1917). christophersi (Edwards), Oc. greenii (Theobald), Ochlerotatus, in part of Senior-White (1923). Oc. hurlbuti (Lien), Oc. okinawanzs (Bohart), and Finlaya, in part of Barraud (L923, 1924), Senior- Oc. taiwanus (Lien). Harrison et al. (1990) elevated White (1926, 1927). Oc. greenii from subspecies to species, synony- Aedes (Finlaya), in part of Edwards (1922a, 1922b, (1926,1931, mized variety kanaranus (Barraud) with it, and 1924,1932, inBanaud 1934), Brug Brug and Edwards (193 l), Barraud clarified the status of Oc. greenii in relation to some 1934), (1934), Taylor (1934), Lee (1944), Bohart of the other species of the subgenus. Tanaka (2OO2) (1946), Bohart and Ingram (1946a, 1946b), King elevated Oc. okinawanas to species status and con- and Hoogstraal (1946), Brug and Bonne-Wepster sidered Oc. taiwanus as a subspecies of it. This (1947), Carter (1950), Knight and Hull (1951), subspecies and the variety doonii of Oc. aureos- Monchadskii (1951), Knight and Marks (1952), triatus are herein provisionally accorded species Bonne-Wepster (1954), Chow et al. (1954), Gut- status based on characters provided by Wattal et al. sevich (1955, 1975), Iyengar (1955, 1960), Hors- (1958), Lien (1968), Tanaka et aI. (1979), and Ta- fall (1955), Srone and Knight (1956), Nakata naka (2002). (1956, 1959), Rao and Rajagopalan (1957), Mac- For adequate examination of specimens, the male donald (1957), Chu (1957, 1958), Wattal et al. genitalia must be dissected to observe the details of (1958), Hara (1959), Bohart (1959), Thurman the claspette and the basomesal structures of the (1959), Stone et al. (1959), Macdonald and Traub gonocoxite. Also, positions of setae 5-7-C and (1960), van der Assem (1961), Stone (1961, measurement of the median length of the dorsal 1970), Kurihara (1963, 1978), Thurman (1963), apotome are best observed in the exuviae of 4th- Scanlon and Esah (1965), Joshi et al. (1965), stage larvae. Terminology used follows Harbach Steffan (1966), Lien (1967, 1968), Huang (1968), (1971), and Knight (1980, 1982), except for terminology Tanaka Stone and Delfinado (1973), (1973), proposed by Reinert (1999, 2000b). Saugstad Hochman and Reinert (1974), Gutsevich et al. (1974), Tanaka et al. (1975), Bhat (1975), Wada et al. (1976), Danilov I Also collaborator, Walter Reed Biosystematics Unit (1977), Knight and Stone (L977), Lien et al. (WRBU), National Museum of Natural History, Smith- (1917), Knight (1978), Tanaka et al. (1979), Gut- sonian Institution, Washington,DC 20560-0165. sevich and Dubitsky (1981), Jayasekera and

309 310 JounNel oF THE AMERIcIN Moseuno CoNrnor- AssocrnrroN VoL. 19, No. 4

Chelliah (1981), Amerasinghe (1982), Lu and Li setae; prealar area with large patch of broad, pale (1982), Lee et al. (1982), Saini and Mishra scales on lower area, several setae on upper area; (1984), Lee et al. (1984), Lei et al. (1984), Toma mesepimeron with large patch of broad, pale scales and Miyagi (1986), Apiwathnasorn (1986), Tsu- covering much of area, several setae on upper, pos- kamoto et al. (1987), Lu and Su (1987), Lee terior area, lower setae absent (except present in (1987), Lee and Zorka (1987), Rattanarithikul Oc. okinawanzs); mesomeron with dorsal margin and Harrison (1988), Amerasinghe and Munasin- well above base of hindcoxa. Legs: Anteprocoxal gha (1988), Lu et al. (1988), Evenhuis and Gon and postprocoxal membranes bare; coxae I-III with (1989), Darsie and Pradhan (1990), Reinert patch of broad white scales, I also with median dark (1990), Townsend et al. (1990), Harrison et al. scales in some species; trochanters with broad, pale (1990), Kaur (1992), Darsie et al. (1991), Ward scales; femora I-III with anterior surface dark- (1992), Stojanovich and Scou (1995, 1996), Lu scaled but normally with pale-scaled areas at base and Ji (1997), Peyton et al. (1999), Reinert and dorsoapically; tibiae I-III often with pale- (2002\. scaled spot ventrobasally; tarsus III dark-scaled Aedes (Finlaya) of Stackelberg (1943), Tanaka with pale-scaled basal bands on at least tarsomeres (2002\. 1-3, some species with pale-scaled basal bands or Aedes, in part of Lee and Lien (1970), Reisen and scales on tarsomeres 4,5 and apical bands or scales Basio (1972), da Costa Pinhao (1974), Sazonova on tarsomeres l-4; posttarsi I,II with 2 ungues, (1981), Mogi et al. (1981), Toma and Miyagi equal in size, both toothed, III with 2 ungues, equal (1981), Lee et al. (1984), Amerasinghe and Mu- in size, both simple. Wing: Dorsal and ventral nasinghe (1984-85), Lee and Egan (1985), Toma veins with dark scales, base of costa with or with- and Miyagi (1992). out white scales; vein R, longer than vein Rr*.; al- Ochlerotatus (Finlaya), in part of Kaur (2003). ula with row of narrow, dark scales on margin; up- per calypter with row of setae on margin; remigium Females. Head: Antennal pedicel with several with 1-3 dark setae. Abdomen: Terga dark-scaled short, fine setae and usually few small, pale or dark with pale-scaled patches basolaterally on I-V[, [- scales mesally; maxillary palpus dark-scaled, short; IV also with or without pale scales dorsobasally proboscis dark-scaled, longer than femur I, narrow, and forming narrow band in some species; sterna more or less straight; clypeus bare; ocular line nar- II-VII pale-scaled proximally and dark-scaled dis- row, with narrow scales; vertex with median patch tally, II usually entirely pale-scaled. pale, of narrow, curved, recumbent scales and sev- Female genitalia. Segment VIII: Laterally com- eral erect forked scales, broad scales laterally; oc- pressed. Tergum VIII: Covered with minute spic- ciput with numerous erect forked scales; eyes con- ules; moderately pigmented; relatively long; base tiguous in front. Thorax: Scutum covered with moderately wide, shorter than length, gently con- narrow, curved scales, background scales dark, pale cave; numerous broad, spatulate scales on distal scales promontory on anterior and antedorsocentral 0.52-0.90; several short setae on distal 0.64-0.87; areas, pale scales forming lines on acrostichal and apical margin gently convex or straight, with 2-4 dorsocentral areas, posterior half of scutum with long and 3-5 short to moderately long setae; ba- pale lines narrow, anterior half of scutum with pale solateral seta present; VII-Te index 0.78-1.06; lines variable from narrow to most of area covered VIII-Te/IX-Te index 2.18-3.06; length 0.234.29 with pale scales, lateral portion of supraalar area mmt width 0.234.35 mrr'. Sternum VIII: Covered pale-scaled, prescutellar space bare except for nar- with minute spicules; moderately pigmented; apex row line of pale scales on each side mesad of pre- gently convex or flat, usually with minute (0.01- scutellar setae; golden to brown setae as follow: 0.02 of dorsal VIII-S length), median emargination, few anterior promontory, acrostichal (few anterior, numerous short and moderately long, relatively 0 to few posterior), few antedorsocentral, dorsocen- straight setae intermixed on apical margin, setae tral (several anterior, several posterior), several pre- tend to increase in length laterally; base narrower scutellar, few scutal fossal, few antealar, several su- than apex, gently concave; numerous broad, spat- praalar (including patch of shorter, blunt-tipped, ulate scales covering distal 0.89-O.96, scales do pale ones in front of wing base), I parascutellar; not, or only partially, overlap presenting somewhat scutellum with narrow, curved scales on all lobes, shaggy appearance; numerous short setae on distal several setae on each lobe; antepronotum with 0.85-0.93; basolateral seta present or absent; setae broad, pale scales, several short and long setae; 1-3-S in more or less diagonal line, l-S basomesad postpronotum with pale scales, narrow, curved ones and some distance from base of sclerite, 3-S api- dorsally and moderately broad to broad ones ven- colaterad; apical, intersegmental membrane nonpig- trally, few posterior setae; postspiracular area with mented; intersegmental membrane between seg- or without scales, several setae; subspiracular area ments VII and VIII moderately long, covered with with elongate patch of broad, pale scales; parater- minute spicules; VIII-S index 0.74-O.95; length gite, mesomeron, and metameron bare; mesokatepi- 0.27-0.36 mm; width 0.33--0.39 rr'll"r.Tergum IX: sternum with upper and lower patches of broad, Covered with minute spicules; comprised of single pale scales, few upper and few to several posterior relatively short (moderately long in Oc. okinawan- Dncerr,rssn2003 BRUCEHARRrsoNrusNsw SuecpNus op OcnrBnortrus 3ll

as) plate, moderately pigmented, but mesal area mented; with 2 heavily pigmented, short, narrow more lightly pigmented; apex with small, median lobes on posterior margin each bearing 3-6 short, emargination; 3-7 short setae distally on each side stout, somewhat flattened setae apically. Gonocox- of midline, 6-12 total setaet base with small to ite: Moderately pigmented; moderately long; distal moderately deep, median emargination; IX-Te in- portion relatively narrow and proximal portion dex 0.64-0.80; length 0.08-0.14 mm; width 0.11- broader; dorsal surface with basal lobe bearing sev- 0.18 mm; dorsal spheres present, dark. Insula: eral moderately long, moderately stout or lanceolate Covered with short spicules; moderately pigment- setae on mesal margin (Oc. alektorovi with these ed; liplike; 2 moderately long, closely set setae on setae short and slender); lateral and ventral surfaces each side laterally, 4 total setae. Lower vaginal lip: with numerous broad scales; ventral surface with Covered with short spicules; moderately pigment- row of moderately long to long, nearly straight se- ed; narrow; hinge narrow; lower vaginal sclerite ab- tae from base to apex, setae decreasing in length sent. Upper vaginal lip: Covered with short spic- on proximal area; mesal margin membranous. Gon- ules; heavily pigmented; narrow with median ostylus: Attached to apex of gonocoxite; lightly to posterior area flattened; upper vaginal sclerite moderately pigmented; long; narrow throughout, small, moderately pigmented. Spermathecal emi- but decreasing in size distally; slightly curved me- nence: Membranous; more or less broadly oval in sad; gonostylar claw short, slendeq slightly curved, outline; few short, simple, spermathecal eminence attached to apex of gonostylus. Claspette: Corn- spicules on basolateral areas. Postgenital lobe: prised of median, ventral, columnar stem with ter- Covered with short spicules; moderately pigment- minal, large, flattened filament, and narrow, lateral ed; moderately long; moderately wide; apex with arm extending from base bearing 2 to several setae, small to moderately deep (0.05-{.22 of dorsal I or more of distal setae flattened.. Proctiger: Par- length), median emargination; 7-13 (1 long, re- aproct narrow, heavily pigmented, but distal portion mainder short) setae distally on each side of mid- more heavily pigmented, apex rounded with short, line, l5-25 total setae; basal mesal apodeme mod- pointed beak; cercus membranous, with 1,2 minute erately pigmented, more or less triangular in setae apicolaterally. Tergun X: Narrow; heavily outline; ventral PGL/Ce index 0.49*0.67; dorsal pigmented; curved; extending from base of para- PGL index 0.65-1.04; ventral PGL index 1.19- proct to base of narrow lobe of tergum IX. Phal- 1.57; ventral length 0.08-O.ll mm. Proctiger: losome: Aedeagus simple, scooplike, ovoid, rela- With scattered small spicules; membranous. Cer- tively short; paramere heavily pigmented, n.urow, cus.' Covered with short spicules; moderately pig- longer than aedeagus; basal piece heavily pigment- mented; moderately long; relatively narrow; apex ed, moderately wide. Sternum 1X.' With 4-9 short sharply rounded; dorsal surface normally without and moderately long setae. scales or when present, usually I or 2, rarely with Pupae. Cephalothorax.' Seta 3-CT very long, 4*6 scales, with number of short setae on distal stout, much longer than l-CT; 5-CT multiple- 0.674.77, 2,3 long and 2-4 moderately long setae branched, noticeably longer than 4-CT; 6-CT short; on approximately distal O.LI4.22; ventral surface 7-CT very long, stout, multiple-branched; 10-CT with 3,4 short setae on apicolateral margin; cercus with several slender branches, shorter than Ll,l2- index 2.O3-2.73; Celdorsal PGL index 2.56-3.60: CT; 1l-CT stout, usually single, occasionally with length 0. 14-0.19 mm; width 0.06-0.O9 rnm. Sper- 2,3 branches, equal to or slightly shorter than 12- mathecal capsules: One large and 2 medium; CT; l2-CT with several branches. Trumpet: Apex heavily pigmented; spherical; with several small, broad; base narrow; tracheoid area basal, weakly spermathecal capsule pores near orifice. Accessory developed. Abdomen: Seta 3-I long, stout, normally gland duct: Basal area moderately pigmented; 2-branched, rarely single or 3-branched; 6-I long, short to moderately long. stout, single, longer than 7-I; l-IJ short, with sev- Males. Similar to females in general habitus. eral to numerous slender branches: 2-II short. lat- Head: Antenna strongly plumose with setae di- erad of l-II, mesad of 3-II; 5-II multiple-branched, rected mainly dorsally and ventrally, shorter than laterad and longer than 4-II; 6-III slender, branched; proboscis and maxillary palpus; maxillary palpus 5-IV-VI long, stout, 5-IV single or 2-branched, 5- dark-scaled, with 5 palpomeres, palpomeres 4 and V,VI single; 2-VI anteromesal to l-VI; 3-VI an- 5 fused, slender throughout, nearly straight but dis- terolateral to l-VI; 7-VI relatively long, single, tal portion slightly upturned, very few short to longer than 6-VI; 6-VII short, with few slender moderately long, dark setae distally on palpomeres branches, anteromesal to 9-VII; 9-VII short, with 3 and 5 and ventrolaterally on palpomere 4, shorter few to several stout, aciculate branches; 9-VIl than proboscis; proboscis longer than femur l. Tho- Iong, with numerous stout, aciculatebranches. Pad- rax.' Scutum sometimes with pale scaling more ex- dle: Relatively narrow without hairlike spicules on tensive on approximately anterior 0.5. Legs.. Post- margins; outer buttress na(Tow, dark, long; midrib tarsi IJI with 2 ungues, unequal in size, both narrow, dark, extending from base to apex; seta l- toothed, III with 2 ungues equal in size, both sim- Pa moderately long, single. ple. Fourth-stage larvae. Head: Seta 1-C relatively Male genitalia. Tergum IX: Moderately pig- long, single, proximal portion moderately stout, 312 JounNel or tHe AMentceN MosQurro CoNrRoL AssocIATIoN Vor. 19,No.4 distal portion slender and attenuated; seta4-C short, tion of 1) eyes are contiguous in front; 2) vertex branched. mesad of 5.6-C. anterior to 5-C, and at has a median patch of pale, narrow, curved, recum- approximately same level as 6-C; 5-C long, ap- bent scales and several erect forked scales; 3) scu- proximately 0.45-0.65 length of dorsal apotome, tum is covered with narrow, curved scales except branched, stout, mesad and posterior to 6-C; 6-C for the bare, median, prescutellar area, the back- exceptionally long, approximately 1.3-1.6 median ground scales are dark, but pale scales form lines length of dorsal apotome, single, stout; bases of 4- on the acrostichal area, dorsocentral area, and along 6-C form small triangle mesad and posterior to lev- the lateral margins of the prescutellar area mesad el of 7-C; 7-C moderately long, branched, posterior of the setae, and patches on the scutal fossal and to antennal base; 12-C branched; l3-C branched, supraalar areas; 4) acrostichal and dorsocentral se- longer than and laterad of l2-C; l9-C present. An- taie ate present; 5) scutellum has narrow scales on tenna: Long, slender; setae l-A normally single all lobes; 6) subspiracular area has an elongate (occasionally 2- or 3-branched). Thorax.' Seta 4-M patch of pale, broad scales; 7) lower prealar area branched;5-M very long, noticeably longer thanT- has a patch of pale, broad scales; and 8) mesepi- M, single; 6-T relatively long, single. Abdomen: meron has the lower setae absent (except present in Setae 6-I,II relatively short; 6-II shorter than 6-I,fII; Oc- okinawanzs); female genitalia by the combi- 7-I longer than 6-I; l2-I present; 7-II short, multi- nation of l) sternum VIII has the apex gently con- ple-branched; 8-II branched; 6-III-VI single (6-III vex or flat and usually with a rninute, rnedian emar- rarely 2-branched); I-VII very long, single; l2-VIl gination and bearing numerous short and branched; 2,4-V[II both branched (very rarely I of moderately long, relatively straight s€tae that tend setae single on one side of specimen); comb com- to increase in length laterally, and numerous broad, prised of numerous scales in patch; saddle incom- spatulate scales that nearly cover the sclerite, these plete ventrally; 2-X moderately long, branched with are not, or only partially, overlapping and give dorsal branch shortest; 3-X very long, single; ven- somewhat of a shaggy appeaf,ance; 2) cercus is tral brush with multiple-branched setae on grid moderately long and relatively narrow, the apex is composed of both transverse and lateral bars, 2 or sharply rounded, and scales are normally absent, or 3 shorter, precratal setae, branched- Siphon: Pecten when present consist of usually only 1 or 2, rarely with numerous evenly spaced spines; seta l-S mul- 4-6 scales; 3) tergum IX is comprised of a single, tiple-branched, inserted distal to pecten. normally relatively short, moderately wide plate Eggs. Unknown. that is moderately pigmented, but with the rnesal Material examined (stageVstructures). Oc&- area lighter pigmented, the apex has a small, me- lerotatus alektorovi (fematre, female genitalia, rnale, dian emargination and bear,ing 3-7 short setae d'is- male genitalia, and larva); Oc. aureostriatus (fe- tally on each side of the midline; and 4) insula is male, female genitalia, male, male genitalia, pupa, liplike with 2 closely set, moderately long setae lat- and larva); Oc. christophersl (female, fernale gen- erally on each side of ttrc midline; males by the italia, pupa" and larva); Oc. greenii (female, fernale rnaxillary palpus being dark-scaled, slender genitalia, male, male genitalia, pupa, and larva); throughout, nearly straight but with the distal por- Oc. hurlbuti (female, female genitalia, pupa, and tion slightly uptumed, shorter than the proboscis, larva); Oc. okinawanus (female, female genitalia, and with very few short to rnoderately long setae pupa, and larva); and Oc. taiwanus (fernale and lar- on palpomeres 3-5; male genitatia by the combi- va). The published literature (see list under synon- nation of 1) tergum IX has the posterior margin ymy) also was examined. bearing 2 short, narrow lobes each with 3-5 short, Distribution. Species included in the subgenus stout, somewhat flattened setae; 2) gonocoxite has have been reported from China ( Island and the ventral surface with a row of moderately long Siohuan Province), India, Indonesia, Japan and to long, nearly straight setae extending from the Ryukyu ,{rchipelago, Malaysia, Nepal, Papua New base to the apex, setae decrease in length on the Guinea, Philippine Islands, South Korea, Russia proximal area; 3) gonostylus is long, narrow (Maritime and Ussurii provinces), Sri Lanka, Tai- throughout but is narower distally, and is attached wan. and . to the apex of the gonocoxite, the goilostylar claw Bionomics. The usual habitat of immatures is is short, slender, slightly .ctrved, and is attached water in tree holes and sturnps or internodes of apically; and 4) claspette is comprised of a median, bamboo; however, some species occasionally have ventral, columnar stern with a terminal, large, flat- ,been taken from artificial containers (e.g., earthen tened filament, and a narrow lateral arm extending jars, gravestones, and a wooden box), rock pools, from the base of the stern and bears 2 to several brsmeliads,:and ferns. Adults of some species have setae of which I or more of the distal setae are been collecfed biting humans during the day and in flattened; pupae by the combination of l) seta 3- light traps. CT is very long and rnuch longer than seta 1-CT; Discussion, The following principal features are 2) seta 3-I is long and norrnally 2-branched; 3) pad- most distinctive for Bruceharrisonius, and they can dle is relatively narrow with the rnidrib extending be used to distinguish this subgenus frorn other sub- ftom the base to the apex; and 4) seta l-Pa is single genera of Achlerotatas: females by the combina- and relatively long; and 4th-stage larvae by the DECEMBER2003 BRUCEHARRTSaNIUSNEw SUBGENUsop Ocutaaorarus 313 combination of 1) seta 4-C is short and branched; brown scales mesally, 0.78-0.82 length of probos- 2) seta 5-C is long and branched; 3) seta 6-C is cis; maxillary palpus dark brown-scaled, 0.17-0.19 exceptionally long and single; 4) bases of setae 4- length of proboscis; proboscis dark brown-scaled, 6-C are positioned in a small triangle with seta 4- l.O9-1.21 length of femur I; clypeus dark brown; C mesad of setae 5,6-C, anterior to seta 5-C, and ocular line with narrow, curved, golden scales, se- at approximately the same level as seta 6-C, and tae dark brown; I pair of golden interocular setae; seta 5-C is posterior and mesad of seta 6-C; 5) seta vertex with large patch of narrow, curved, golden, 7-C is branched and at a level anterior to setae 4- recumbent scales and several golden, erect forked 6-C and posterior to the antennal base; 6) seta 19- scales, patch of broad, dark brown scales laterad of C is present; 7) antenna is long, slender, and seta these followed by broad, white scales laterally and l-A is normally single (occasionally 2- or 3- caudally; postgena with broad scales, small patch branched); 8) setae 6-I,II are relatively short and of dark brown ones anteriorly at about level of an- seta 6-II is shorter than seta 6-I,ilI; 9) seta 7-I is tepronotum, white ones below; occiput with nar- longer than seta 6-I; 10) seta I-VII is very long and row, curved, golden, recumbent scales and numer- single; 11) setae z,4-VIll are both branched (very ous moderately long, golden, erect forked scales. rarely is 2- or 4-VIII single on one side); and 12) Thorax: Scutum with integument dark brown, cov- comb is comprised of multiple scales in a patch. ered with niurow, curved scales except for bare Within subgenus Finlaya (genus Ochlerotatus), prescutellar area and small area on each side of adults of some other groups of species possess a anterior promontory between antedorsocentral are- pattern of narrow, pale scales forming lines on the as, background scales dark, reddish-brown, with acrostichal and dorsocentral areas of the scutum numerous golden scales on following: anterior (e.g., Aureostriatus Group, in part, of Knight and promontory, antedorsocentral area, line on acrosti- Marks [952] and the Chrysolineatus Subgroup of chal area from anterior margin to prescutellar area Knight t19681). Adults of Bruceharuisonius species where it forks along lateral margins of prescutellar can be separated from most of these species by one area mesad of setae, anterior portion of line broad or more of the following: the entirely dark-scaled and contiguous with dorsocentral golden line but palpus and proboscis, broader pale-scaled lines of posterior portion of line narrow, line on dorsocen- the scutum, subspiracular area with broad, pale tral area extending from anterior margin but not scales, and the development of the male maxillary reaching prescutellar a.rea, narrow line on mesal palpus. The female and male genitalia, pupae, and margin of supraalar area extending from posterior 4th-stage larvae of these groups differ from those margin of scutal fossal area to posterior margin of of Bruceharrisonius (see above list of principal fea- scutum laterad of prescutellar setae (narrow line of tures). The exceptionally long and single seta 6-C dark, reddish-brown scales along bases of prescu- of larvae of Bruceharrisonius is shared by members tellar setae), scutal fossal area covered and contig- of subgenus Geoskusea Edwards and Oc. (Fin.) Sil- uous with dorsocentral line (except for small, elon- /i (Barraud); however, numerous other differences gate patch of dark, reddish-brown scales on median exist in all life stages from those of Bruceharrison- area), antealar and supraalar areas covered except ias. The most distinctive features of the new sub- for an elongate patch of reddish-brown scales on genus are found in the fumale and male genitalia median area of supraalar area; golden to brown se- and the immature stages. Lien (1967) reported that tae on following areas: 3,4 anterior promontory, 1- in larvae of Oc. hurlbwi seta 6-C was bifid with 1 4 anterior acrostichal, 2-4 antedorsocentral, several branch very long whereas the other branch was dorsocentral (anterior and posterior), 7-10 prescu- more slender and shorteq this condition is atypical tellar. 3.4 scutal fossal. 3,4 antealar. numerous su- for the subgenus. praalar (upper ones dark and long, small patch of It is a pleasure to trame the new subgenus Bruce- short, blunt-tipped, pale ones in front of wing base), harcisonius in honor of Dr. Bruce A. Harrison, and I parascutellar; scutellum with narrow, curved, North Carolina Departrnent of Environment and dark, reddish-brown scales on each lobe, also few Natural Resources, Winston-Salem, North Caroli- golden narrow, curved scales on median area of na, in recognition of his outstanding contributions midlobe and few on lateral lobes, midlobe with 3,4 to the biosystematics of the Culicidae, especially to long and 2-6 short setae, lateral lobe with 3-6 long the genus Meigen. The subgenus is mas- and 2,3 short setae; pleural integument dark brown; culine and is abbreviated Bry'r. antepronoturn with broad, dusty-white scales on an- terior, lower lateral, and posterior surfaces, 14-19 setae; postpronotum with narrow, curved, golden Type species description, Ochlerotatus scales dorsally and few dusty-white or golden, (Bnrc eharrisonius, gaeenii (Theobald), 1903 moderately broad scales ventrally, 3-5 posterior se- Fernale (see Fig. 6, Flate III (type form) in Bar- tae; postspiracular area without scales, with 5-8 se- raud 1934). Head: Amtennal pedicel mostly dark tae; subspiracular area with elongate patch of brown but pale anteriorly, with patch of short, fine, broad, white scales; upper proepisternum with dark setae and 2,3 small, broad, dark brown scales patch of broad, white scales, 5-9 pale setae; me- mesally, flagellomere I with small, broad, dark sokatepisternum with upper and lower patches of 314 JourNnL or rur Avrnrcel Moseurro CoNrnor AssocrlrroN Vor. 19,No.4 broad, white scales, 3,4 upper and 4-6 posterior 0.38 mm. Tergum IX; Apex with 3,4 short setae on setae; prealar area with large patch of broad, white each side of midline, 6,7 total setae; IX-Te index scales on lower area, 7-10 setae on upper area; 0.73-O.80; length 0.08-0.10 mm; width 0.11-0.13 mesepimeron with large patch of broad, white mm. Postgenital lobe: Apex with moderately deep scales covering most ofarea, 4-10 upper pale setae, (O.18-O.22 of dorsal length), median emargination; lower setae absent. Legs: Coxa I with broad, white 8-13 setae on each side of midline, 77-25 total scales dorsally and ventrally separated by median setae; ventral PGL/Ce index 0.64-0.67; dorsal PGL patch of broad, brown scales; femora I-III dark index 0.86-0.91; ventral PGL index 1.49-1.57; brown-scaled except for white scales as follow: I ventraf length 0. 10-0.11 mm. Cercus: Dorsal sur- with narrow, white-scaled, basal band, posterior face without scales, with number of short setae on surface with median, longitudinal stripe from base distal 0.68-0.72, 3 long and 3,4 moderately long to apex, stripe broader on basal area and reaching setae on approximately distal 0.22; cercus index posterodorsal margin, II with small, dorsobasal 2.65-2.73; Celdorsal PGL index 2.56-2.73; length patch,2,3 dorsoapical scales and indistinct narrow 0.16-0.18 mm; width 0.06-0.07 mm. Accessory stripe on ventral margin of anterior surface, poste- gland duct: Basal area relatively short. rior surface with posteroventral, longitudinal stripe Male. Similar to female in general habitus. from base to apex, stripe wider on basal area, III Head: Antenna 0.63-0.66 length of proboscis; with approximately anterobasal 0.65 and approxi- maxillary palpus with very few short, dark brown mately posterobasal 0.58 (except dorsal margin setae on palpomeres 3-5, 0.694.73 length of pro- brown-scaled), apex with small, patch on anterior, boscis; proboscis 1.16-1.30 length of femur l. Tho- dorsal, and posterior surfaces; tibiae I-III dark rar.' Scutum with dark-scaled areas reduced and brown-scaled except for small, basal, white-scaled approximately anterior 0.5 primarily golden-scaled. patch on posteroventral area, III and sometimes II Wing: Base of costa dark-scaled. with white-scaled patch slightly larger and extend- Male genitalia (Figs. 2A-2C). Tergum IX: Pair ing onto anteroventral area; tarsi I,II dark brown- of lobes on posterior margin each bearing 3-5 scaled except for white scales on following: II with short, slightly curved, stout, somewhat flattened tarsomere I with narrow, basal band (some speci- setae apically; anterior margin deeply emarginate. mens also with 2,3 dark scales on anterior surface Gonocoxite: Proximal approximately 0.40 of dor- of band), and few at apex, tarsomere 2 with narrow sal surface increasing in width to form more or basal band or with 3,4 dorsobasal scales, tarsomere less triangular lobe with moderately pigmented, 3 of some specimens with 2-4 scales dorsobasally, posteriorly extended, oval area (that normally pro- III with tarsomeres I and 2 with moderate basal jects sternally) bearing 10,1I moderately long, band and nalrow apical bands, tarsomere 3 with stout, lanceolate setae along mesal margin, each narrow basal band and few dorsoapical scales, tar- seta with short, basal sheath, lateral margin of oval somere 4 with few dorsobasal scales. Wing: Dorsal area surrounded by lightly pigmented, narrow, and ventral veins with dark brown scales, base of crescent-shaped strip bearing 9-12 moderately costa with few white scales; remigium with 2,3 rel- long, slender setae; dorsal surface with about 4 atively long, brown setae. Halter: Pedicel pale; very long (extending beyond apex of gonocoxite), capitellum with broad, brown scales except broad, stout setae arising laterally from near base, ap- white scales on anterior and ventral surfaces. Ab- proximately distal 0.20 with several long, slightly domen: Terga dark brown-scaled with small baso- curved, stout setae; lateral and ventral surfaces lateral white-scaled patch, III-VI with narrow, bas- covered with numerous long, spatulate scales; al white-scaled bands, bands incomplete on III and ventral surface covered with moderately long to IV in some specimens, VI with large basolateral long setae scattered over most of area, mesal mar- patch nearly meeting dorsomesally; sternum II gin with irregular row of moderately long and long white-scaled, III-VI dark brown-scaled with basal setae from base to apex, setae increasing in length white-scaled band that becomes broader on lateral distally. Gonostylus: With 1,2 short, slender setae areas; terga and sterna with short setae on lateral near apex. Claspette: Comprised of moderately and posterior margins. long, slender stem (somewhat thickened at about Female genitalia (Fig. 1). Tergum VIII: Basal 0.74 from base), covered with tiny spicules except 0.50-0.60 retracted into segment VII; scales on dis- distal portion, terminating with large, flattened, tal 0.52-{.58; setaeon distal 0.64-0.66; apical mar- leaf-like filament, 3 short, slender setae on thick- gin more or less straight, with 2 long and 4,5 mod- ened area and I short, slender seta at about prox- erately long setae; basolateral seta near basolateral imal 0.34, stem base with long, narrow arm pro- margin; VIII-Te index I .O4-1 .O6; VIII-Te/IX-Te in- jecting laterally and curved dorsally, partially dex 2.68-3.06; length O.25-O.28 mm; width 0.23- connected to sternomesal portion of oval, baso- 0.27 mm. Sternum VIII: Apex gently convex with mesal area of dorsal surface of gonocoxite, nalrow minute (0.01-0.02 of dorsal VIII-S length), median extension bearing several (about ll) setae along emargination; scales covering distal 0.96; setae on margin, distal 5 setae broadly flattened, next 2 or distal 0.85-0.91; basolateral seta absent: VIII-S in- 3 setae proximally lanceolate, and most proximal dex 0.86--0.88; length 0.30{.34 mm; width 0.35- 2 or 3 setae stout, simple, distal 7 or 8 setae each DECEMBER2003 BnucoaenntsoytusNpw SUBcENUSop Ocnrenortrus 315

VL

venlrol SCoP

o.2T dor soI I I

-Te Vlll J/.ta"'a,'

Fig. l. Ochlerotatus (Burceharcisonius) greenii, female genitalia. AGDB, accessory gland duct base; BMA, basal mesal apodeme; Ce, cercus; DPGL, line of attachment of proctiger to dorsal surface of postgenital lobe; DS, dorsal sphere; H, hinge; I, insula; IX-Te, tergum IX; LVL, lower vaginal lip; PGL, postgenital lobe; Pr, proctiger; SCa, seminal capsule; SCaP, seminal capsule pore; SE, spermathecal eminence; SES, spermathecal eminence spicule; UVL, upper vaginal lip; UVS, upper vaginal sclerite; VIII-S, sternum VIII; VIII-Te, tergum VIII; VTI ventral tuft; 1-3-S, setae l- 3 of sternum VIIL 316 JouRNnl oF THE AMERICaT Moseuno CoNrnol AssocIATroN Vor-.19. No.4

\--\ \'-)t D

tvI

V-t\

G

Fig. 2. Ochlerotatus (Bruceharrisonius) greenii; male genitalia. (A) claspette, (B) phallosome, (C) tergum IX; pupa, (D) trumpet, (E) abdominal segments VIII and X, genital lobe, and paddle, (F) lateralia; 4th-stage larva, (G) cranium, (H) abdominal segments VII, Vil, and X, and siphon, (I) enlarged comb scale, (J) enlarged pecten tooth. A, antenna; Ae, aedeagus; AeG, aedeagal guide; C, cranium; CII claspette filament; CSt, claspette stem; GL, genital lobe; IX-Te, tergum IX; Lat, lateralia; MCL, median caudal lobe; Pa, paddle; Par, paramere; S, siphon; Sa, saddle; T, trumpet; VII, abdominal segment VII; VIII, abdominal segment VIII; X, abdominal segment X. with a short, basal sheath; bases of claspette stems teriorly, 2 long and 2-4 short to moderately long; connected mesally by wide, spiculate aedeagal narrowly connected laterally to tergum IX. guide. Phallosome: Aedeagus relatively short Pupa (Figs. 2D-2F). Setal branching as in Table with apex narrower than base, narrow lateral areas l. Cephalothorax: Ittegument pale tan with some and approximately basal O.27 heavlly pigmented, darker areas. Trumpet.' Moderately pigmented; remainder lightly pigmented; paramere approxi- apex relatively broad and tapering to narrow base; mately l.l6 the length ofthe aedeagus; basal piece tracheoid area small, basal, and poorly developed; approximately O.74 the length of the aedeagus. index 1.'74-3.25, shortest in male; pinna 0.32-O.38 Sternum 1X.' With heavily pigmented, broad me- the length of the trumpet. Abdomen' Integument dian and narrow lateral areas; 4-6 stout setae pos- pale tan with some darker areas on terga l-IY. Pad- DECEMBER2003 BRUCEHARRTS)NTUSNEw SuscsNus oF OcH LER)TATUS 31'7

d/e.' Relatively narrow; outer buttress on basal o 0.57-0.68 of outer margin; margins of distal area with few small spicules; index 1.58-1.69. Fourth-stage larva (Figs. 2G-2J). Head: Seta l-C relatively long, single, simple; 3-C short, sin- gle, simple; 4-C short, with 9-1 I slender branches; c{ 5-C long, approximately 0.5 the median length of cl n the dorsal apotome, with 3 stout, aciculate branch- N I es, mesad and posterior to 6-C; 6-C exceptionally long, approximately 1.5 the median length of the dorsal apotome, single, stout; 7-C moderately long, with 7-1 1 stout, aciculate branches; 8-C with 3,4 ---- slender branches; 9-C with 5-9 slender branches; _vJ -e.r-99S9-\tio- 10-C with 3-6 slender branches; ll-C with 5-7 o .tttt trl stout, aciculate branches; 12-C short, with 6-10 C'.1-C..la-.16d- slender branches; l3-C moderately long, with 4-6 slender branches; 14-C short, single, near anterior margin of lateralla; 15-C with 3-7 branches; 18,19- 0O C tiny. Antenna: Shaft long, slender, few spicules on proximal area; seta 1-A moderately stout, single, s simple, inserted approximately 0.50-O.55 from base of the shaft; 2-5-A inserted at apex of the shaft. Mouthparfs.' Seta 6-Mx short, single or 2- branched; lateral palatal brush with simple fila- ments; dorsomentum with 25,26 posterior teeth.

o Thorax: Seta 0-P short. with 25-35 slender. den-

b0 dritic branches; l-P with 2,3 branches; 2-P single; a 3-P with 4,5 branches; 2-P>I-P>3-P length; 4-P a a^^^^^ with 3,4 branches: 5-P with 3 stout, aciculate U 3 cn !'1.rt Ol.f.) f branches; 6-P single, stout, simple; 7-P with 2,3 ^ -=-9''iTTJg-T-Jvvvv stout, aciculate branches; 8,11,14-P short, with 2,3 6t + 6 N .q cr.r.n J branches; 9,LO,12-P single; 1-M with 2,3 branches; z,lt-M with 2-4 branches; 3,5,7,10,12-M single, 5-M stout, very long, aciculate; 4-M with 3-6 i branches; 6-M with 3,4 stout, aciculate branches; : 3999S9 3 -9-n09q\9\q-!a- 8-M with 4,5 stout, aciculate branches; 9-M with 4 o rlllllll branches: 13-M short. with 19-23 slender. dendritic .b F$no$n m branches; l-T with 3-8 branches; 2-T with 3-8 o{ branches; 3-T with 5-7 branches; 4-T with 6-9 branches; 5-T single to 4-branched; 6,10,12-T sin- gle;7-T with 6-9 stout, aciculate branches; 8-T g .?) frCn .a *?*ve9*":i-: i-v ) short, with 10-16 slender, dendritic branches; 9-T ^-+ o I with 4,5 stout, aciculate branches; l1-T with 2,3 -l branches; 13-T short, with 7-14 slender, dendritic o N branches. Abdomen: Much of abdominal segments ! III-VI were badly twisted or damaged in the avail- * able specimens. Seta l-I short, with 5-12 slender 6: oo; : branches; 2,10-I single, 2-I short, l0-I moderately long; 3-I with 3-6 branches; 4-l with 12-29 F - fri^Tr-o.6 di i-N branches; 5-I with 5-8 branches; 6-I relatively C.l € short, with 3 stout, aciculate branches; 7-l witlr 2

4 stout, aciculate branches; 9-I with 3-5 branches; 1l-I with 5,6 branches; 12-I with 2-4 branches; 13-

.9x m.i h+6r)++€tif- I single or 2-branched; 1-II with 2-4 branches;2,9- : Jvve !nrl-tt'.nsf9++qoq II single; 3-II with 5,6 branches; 4-II with 13-19 ttttrt-'-.t,1 ri- -- +mN+r,-,i.-€ branches; 5-II with 8,9 branches; 6-II relatively short with 2 stout, aciculate branches; 7-II with 5- {o 8 branches: 8.10-II with 3.4 branches: l1-II with 2 OY O*C.lcOSr)\OF€O\OFcl$ branches; 13-II with 25--29 branches; 6-III-VI sin- ar gle; 1-V[ very long, single, stout; 2-VII single; 3- VII with 3-9 branches; 4-VII with 4,5 branches; 5- 318 JounNal op rns AvEnrcAN MoseulTo CoNrnol AssocrnrroN

VII with 6-9 branches; 6-VII with 2l-30 branches; ACKNOWLEDGMENTS 7-VII with 3 branches; 8-VII with 17-31 branches; 9-VII with 3-5 branches; lO-V[ with3,4 branches; Appreciation is expressedto Donald R. Barnard ll-V[ single; I2-VII with 4-6 branches; I3-VII (CMAVE) for providing facilities; to Ralph E. Har- with 9-11 branches; I-VIII with 4-7 slender bach (NHM, London, United Kingdom), Richard C. (WRBU), branches; 2-VIII with 3-5 slender branches; 3-VIII Wilkerson and Thomas J. Zavortink (Bo- with 2-4 stout, aciculate branches; 4-VIII with 2,3 hart Museum, University of California, Davis, CA) for reviewing slender branches; 5-VIII with 4-6 slender branch- the manuscript; to Thomas V. Gaffi- gan (WRBU) and Theresa M. Howard (NHM) es; segment VIII with comb of 47-68 scales in for the loan of specimens;and to Taina R. Litwak (Dar- patch, scales moderately long, apex rounded, with nestown,MD) for preparingFig. 1. slender spicules on apical and lateral margins; sad- dle heavily pigmented, incomplete ventrally, few short, stout spicules along upper, posterior margin, REFERENCES CITED short rows of 3-5 minute spicules scattered over Amerasinghe FP 1982. Observations on the mosquitoes entire surface, saddle/siphon index 3.23-3.6O; seta (Diptera: Culicidae) of Udawattakele Forest, Sri Lanka. l-X short, with 2 (rarely 3) branches, inserted on J Natl Sci Counc Sri Innka lO:81-97. saddle near posterior margin; 2-X with 3 branches, Amerasinghe FP, Munasinghe NB. 1984-85. Further ob- dorsal branch noticeably shortest; 3-X very long, servations on the mosquito fauna of Udawattakele For- single, noticeably longer than 2-X; ventral brush est, Sri Lanka. Ceylon J Sci (Bio Sci) 17 and 18:l-21. with 8 (occasionally 9) setae attached to grid, setae Amerasinghe FQ Munasingha NB. 1988. A predevelop- ment mosquito survey in the Mahaweli Development fanlike with 6-8 branches. and 2 short. 5- or 6- Project area, Sri Lanka: adults. "/ Med Entomol 25:276- branched, precratal setae, grid with both lateral and 285. transverse bars; 4 anal papillae, dorsal pair longer Apiwathnasorn C. 1986. A list of mosquito species in than ventral pair. Siphon: Index 3.18-3.39 (dorsal Southeast Asia Bangkok, Thailand: Museum and Ref- length to width at 0.5 length); acus small, at base erence Centre, SEAMEO-TROPMED Nature Centre of of pecten; pecten on proximal approximately 0.39 Thailand, Mahidol University. Barraud PJ. 1923. A revision of siphon, comprised of 25-30 evenly spaced of the culicine mosquitoes of India. Part VII. The larvae of some species of Srego- spines, basal 2 or 3 spines reduced in size, remain- myia (Theo.), the larvae of some species of Finlaya der of spines heavily pigmented with 2,3 small (Theo.), the larva of Christophersiomyia thomsoni spicules near mid ventral margin; seta 1-S with 4- (Theo.), the larva of Mimomyia chamberlaini (Ludl.), 7 aciculate branches, borne distal to pecten; 2,6-5 the larva of Aedomyia fsic'! catasticta (Knab.). Indian single; 8-S with 3-9 branches. J Med Res ll:495-505 + pl. XXXIV-XL. Type data. The holotype female is mounted on Barraud PJ. 1924. A revision of the culicine mosquitoes a minuten pin attached to a circular paper stage of India. Part VIII. Further descriptions of Indian spe- cies of FinlayaTheo. Indian J Med Res I l:845-865 + attached to an pin and has 5 paper labels with pl. XLVIII-LVI. the following data: /93, Ceylon, E. E. Green/ /Per- Barraud Pl. 1934. The fauna of British India, including adeniya, Ceylon, II.l900/ /133/ IHolotype/ and /Ae- Ceylon and Burma. Diptera Volume Y Family Culici- des greenii Type FVT/. The holotype is deposited dae. Tribes Megarhinini and Culicini London, United in The Natural History Museum (NHM), London, Kingdom: Taylor and Francis. United I(ingdom. Bhat HR. 1975. A survey of haematophagous Discussion. The above description of Oc. green- in western Heimalayas, Sikkim and Hill Districts of West Bengal: records of mosquitoes collected from Hi- ii is based on the holotype female (see above) and malayan region of Uttar Pradesh with ecological notes. numerous topotypic males and females with asso- Indian J Med Res 63:1583-1608. ciated pupal and 4th-stage larval exuviae and 4th- Blanchard R. 1905. Les moustiques histoire naturelle et stage larvae (9 : female; d : male; p: pupal meelicale Paris, France: E R. de Rudeval, Imprimeur- exuviae; I : Iarval exuviae; L : Iarva) with the Editeur. [In French.] following collection data: Sri Lanka, Central Prov- Bohart RN. 1946. New species of mosquitoes from the (Diptera, ince, Kandy District, Peradeniya, Royal Botanic Marianas and Okinawa Culicidae). Proc Biol Soc Wash 59:39-46. Garden, E. L. Peyton and Y.-M. Huang collectors, Bohart RN. 1959. A survey of the mosquitoes of the immatures collected fiom small bamboo stumps, 30 southern Ryukyus. Mosq News 19:194-197. June 1975, collection number 142 (3?pl, 89p, Bohart RN, Ingram RL. 1946a. Four new species ofmos- 7dp), collection number 145 (59p1, l9p, 2dp), quitoes from Okinawa (Diptera: Culicidae). J Wash collection number 162 (2?pl,2L), collection num- Acad Sci 36:46-52. ber 167,1 July 1975 (19p, l9), collection number Bohart RN, Ingram RL. 1946b. Mosquitoes of Okinawa 180, I July 1975, immatures collected from leaf and isletnds in the Central Pacffc NAVMED 1055. Washington, DC: Bureau of Medicine and Surgery axils of ground bromeliad (l9pl, adpl, 26p, lpl, Navy Department. 3L). The topotypic specimens are deposited in the Bonne-Wepster l. 1954. Synopsis of a hundred common National Museum of Natural History, Washington, non-anopheline mosquitoes of the Greater and Lesser DC. Sundas, the Moluccas and New GuineaRoyal Tiopical DECEMBER2003 BRUCEHARRISoNTUs New SuscpNus op Ocatznorerus 319

Institute of Amsterdam Special Publication CXI. Am- lasian and Oceanian regions Honolulu, HI: Bishop Mu- sterdam, The Netherlands: Elsevier Publishing Co. seum Press. p l9l-218. Brug SL. 1926. The geographical distribution of mosqui- Giles GM. 1900. A handbook of the gnats or mosquitoes toes in the Malayan Archipelago. Meded Volksgezon- giving the anatomy and lift history oJ the Culicidae dheid Ned-Ind 4:47 l-482. London, United Kingdom: John Bale, Sons and Dan- Brug SL. 1931. Culiciden der Deutschen Limnologischen ielsson, Ltd. Sunda-Expedition. Trop Binnengewass 2:1-42. [n Giles GM. 1902. A handbook of the gnats or mosquitoes Dutch.l giving the anatomy and life history of the Culicidae 2nd Brug SL. 1934. Notes on Dutch East Indian . ed. London, United Kingdom: John Bale, Sons and Bull Entomol Res 25:501-519. Danielsson, Ltd. Brug SL, Bonne-Wepster J. 1947. The geographical dis- Gutsevich AV. 1955. New and little known forms of mos- tribution of the mosquitoes of the Malay Archipelago. quitoes (Diptera, Culicidae). Akad Nauk Zool Inst (Tru- Chron Nat lO3:179-197. dy) 18:32O-324. [In Russian.] Brug SL, Edwards FW. 1931. Fauna Sumatrensis (Bijdr- Gutsevich AV. 1975. The determination of mosquito fe- age nr. 68), Culicidae (Diptera). O v er g edrukt Tij ds c hr ift males (Culicidae) by microscopic preparations of the Entomol 74:257*261. head IV. A key to species of the genus Aedes. Mosq Brunetti E. 1907. XXV.-Annotated catalogue of Oriental Syst 7:164-173. [Originally published in Russian in Culicidae. Rec Indian Mus (Calcutta) l:297-371 . P arazito lo giya 8:329 -335, 197 4.1 Brunetti E. 1912. X. Annotated catalogue of Oriental Cu- Gutsevich AV, Dubitsky AM. 1981. New species of mos- licidae-supplement. Rec Indian Mus (Calcutta) 4'.403- quitoes in USSR fauna Leningrad, USSR: USSR Acad- 517. emy of Science, Zoological Institute. Brunetti E. 1920. I. Catalogue of Oriental and south Asi- Gutsevich AV, Monchadskii AS, Shtakel'berg AA. 7974. (Calcutta) at'ic Nemocera. Rec Indian Mus 17:l-300. Fauna of the U.S.S.R. Diptera. Mosquitoes family Cu- Carter HF 1950. Ceylon mosquitoes: lists of species and licidae Jerusalem, Israel: Israel Program Scientific names of mosquitoes recorded from Ceylon. Ceylon J Translations. [Translation from Russian.] Sci (B) 24:85-115. Hara J. 1959. Two new mosquito records from Japan (Dip- Chow CY Thevasagayam ES, Tharumarajah K. 1954. In- tera: Culicidae). Jpn J Sanit Zool 10:225-229. sects of public health importance in Ceylon. Rev Ecuat Harbach RE, Knight KL. 1980. Taxonomists' glossary of Entomol Parasitol 2: I 05-150. mosquito anatomy Marlton, NJ: Plexus Publishing, Inc. Chu E 1957. Collection of megarhine and culicine mos- Hiubach RE, Knight KL. 1982. Correcrions and additions quitoes from Hainan Island, south China, with descrip- to Tcsonomist' glossary of mosquito anatomy. (1981) tion of a new species. Acta Zootax Sin 9:145-163 + pl. Mosq Syst l3:.2ol-217. I-III. [In Chinese.] Hartison BA, Rattanarithikul R, Peyton EL, Mongkolpan- Chu F-I. 1958. Advances in the study of culicine mos- ya K. 1990. Thxonomic changes, revised occurrence re- quitoes of Hainan, south China. Indian J Malariol 12: cords and notes on the Culicidae of Thailand and neigh- 109-1 12. boring countries. Mosq Syst 22:196-227. da Costa Pinhao R. 1974. Sobre a situacao zoogeografica Hochman RH, Reinert JF. 1974. Undescribed setae in lar- de Timor. An Inst Hig Med Trop 7;467-47O. vae of Culicidae (Diptera). Mosq Syst 6:1-10. Danilov VN. 1977. On the synonymy of species names of Horsfall WR. 1955. Mosquitoes their bionomics and re- Aedes [sic] mosquitoes (subgenera Finlaya [sic] and lation to disease New York, NY: Ronald Press Co. Neomelaniconion [sic]) in the Far East fauna. Parazi- Huang Y-M. 1968. A new subgenus of Aedes (Diptera, tologiya I I :181-184. [In Russian.] Culicidae) with illustrated key to the subgenera of the Darsie RF Jr, Pradhan SP | 990. The mosquitoes of Nepal: Papuan Subregion (Diptera: Culicidae). J Med Entomol their identification, distribution and biology. Mosq Syst 5:169-188. 22:69-13O. Iyengar MOT 1955. Distribution of mosquitoes in the Darsie RF Jr, Pradhan Sf; Vaidya RG. 1991. Notes on the South Pacifc Region Technical Paper 86. Noumera, mosquitoes of Nepal I. New country records and re- New Caledonia: South Pacific Commission. (Diptera, vised Aedes keys Culicidae). Mosq Syst 23: Iyengar MOT. 1960. A review of the mosquito fauna of 39-49. the South Pacffic (Diptera: Culicidae). Technical Paper Doleschall CL. 1857. Tweede bijdrage tot de kennis der 130. Noumea, New Caledonia: South Pacific Commrs- dipterologische fauna van Nederlandsche Indie. Na- s10n. tuurk Tijdschr Ned-Ind 14:377-418. [In Dutch.] Jayasekera N, Chelliah RV. 1981. An annotated checklist Edwards FW. 1917. Notes on Culicidae, with descriptions of mosquitoes oJ Sri Innka (Diptera: Culicidae). Pub- of new species. Bull Entomol Res 7:2Ol--229. licaton 8. MAB-UNESCO-Man and the Biosphere Edwards FW. 1922a. A synopsis of adult Oriental culicine Nat Committee of Sri Lanka (including megarhinine and sabethine) mosquitoes. Part Joshi G, Pradhan S, Darsie RF Jr. 1965. Culicine, sabeth- l. Indian J Med Res lO:249-293. ine and toxorhynchitine mosquitoes of Nepal including Edwards FW. 1922b. A synopsis of adult Oriental culicine new country records (Diptera: Culicidae). Proc Ento- (including megarhinine and sabethine) mosquitoes. Part mol Soc Wash 67:137-146. ll. Indian J Med Res lO:43O-475. Kaur R. 1992. Mosquito types present in the collections Edwards FW. 1924. A synopsis of the adult mosquitos of of National Institute of Communicable Diseases, Delhi the Australasian Region. Bull Entomol Res l4:35I-4O1. (Diptera: Culicidae). Orient 26:367-382. Edwards FW. 1932. Genera insectorum. Diptera, fam. Cu- Kaur R. 2003. An update on the distribution of mosquitoes licidae Fascicle 194. Bruxelles, Belgium: Desmet-Ver- of the tribe Aedini in India (Diptera: C]ulicidae). Orient teneuil, Imprimeur-Editeur. Insect 37:439-455. Evenhuis NL, Gon SM III. 1989. Family Culicidae. In: King WV, Hoogstraal H. 1946. Species of Aedes (Finlaya) Evenhuis NL, ed- Catalog of the Diptera of the Austra- [sic] of the papuensis group in the Australasian Region 320 Joumnl or rHs AMERTcIN Mosqulro CoNTRoL AssocrlrloN 19,No. 4

(Diptera, Culicidae). Proc Entomol Soc Wash 48:135- Aedes and seven new species of Culex. Trop Med lO: 157. 2r7-262. Knight KL. 1968. Contributions to the mosquito fauna of Lien JC, Kawengian BA, Partono E Lami B, Cross JH. Southeast Asia.-IV. Species of the subgroup Chryso- 1977. A brief survey of the mosquitoes of south Sula- lineatus of group D, genus Aedes, subgenus Finlaya wesi, Indonesia, with special reference to the identity Theobald. Contrib Am Entomol Inst (Ann Arbor) 2(5\: of Anopheles barbirostris (Diptera: Culicidae) from the 1-45. Margolembo area. J Med Entomol 13:719-72'7. Knight KL. 1978. Supplement to a catalog of the mos- Lu B, Chen H, Xu R, Ji S. 1988. A checklist of Chinese quitoes of the world Washington, DC: Thomas Say mosquitoes (Diptera: Culicidae) Guiyang, China: Foundation, Entomological Society of America. Guizhou People's Publishing House. [In Chinese.] Knight KL, Hull WB. 1951. The Aedes mosquitoes of the Lu B, Ji S. 1997. Subgenus Finlaya Theobald, 1903. In: Philippine Islands L Keys to species. Subgenera Macr- Lu B, Li B, Ji S, Chen H, Meng Q, Su L, Qu f;, Gong dus, Ochlerotatus, and Finlaya (Diptera, Culicidae). Z, Zhang Z, eds. Diptera: Culicidae I. Fauna Sinica, Pac Sci 5:211-251. Insecta Yofume 8. Beijing, People's Republic of China: Knight KL, Marks EN. 1952. An annotated checklist of Science Press. p 99-173, Figs. l8-60. [In Chinese.] the mosquitoes ofthe subgenus Finlaya [sic], genus Ae- Lu B, Li P. 1982. Chapter 1. Identification of Chinese des [sic]. Proc US Natl Mus 10l:513-574. Mosquitoes. In: Lu B, ed. Ilandbook for the identifi- Knight KL, Stone A. 1977. A catalog of the mosquitoes cation of Chinese important medical zoology Beijing. of the world (Diptera: Culicidae) College Park, MD: China: People's Hygiene Publishing House. p l-159, Thomas Say Foundation, Entomological Society of 941-944. [In Chinese.] America. Lu B, Su L. 1987. A handbook for the identification of Kurihara T. 1963. Comparative studies on the pleural Chinese aedine mosquitoes Beijing, People's Republic stnrcture of the Japanese mosquitoes. Jpn J Sanit Zool of China: Science Press. [In Chinese.] l4:.l9l-2o7. [In Japanesewith English summary.] Macdonald WW. 1957. Malaysian parasites, XVL An in- Kurihara T, 1978. Collection records of mosquitoes' in terim review of the non-anopheline mosquitoes of Ma- Indonesia. Teixz\o J Med l:333-338. [In Japanese with laya. In: Andy, JR, ed. Malaysian parasites XVI- English abstract.l XXXN. Study no.28 Kuala Lumpur, Malaysia: Institute Lee DJ. 1944. An atlas of the mosquito larvae of the Aus- for Medical Research of the Federation of Malaya. p tralasian Region tribes-Megarhinini and Culicini. t-34. Australia: North Melbourne Victorian Railways Print- Macdonald WW, Traub R. 1960. Malaysian parasites ing Works, by authority of HQ Australian Military XXXVII. An introduction to the ecology of the mos- Forces. quitoes of the lowland dipterocarp forest of Selangor, Lee DJ, Hicks MM, Griffiths M, Russell RC, Marks EN. Malaya- In: Macdonald WW, ed. Malaysian Parasites 1982. The Culicidae of the Australasian region Volume XXXV-XLIX. Study no. 29 Kuala Lumpur, Malaysia: In- ll Nomenclature, synonymy, lite rature, distribution, bi- stitute for Medical Research of the Federation of Ma- ology and relation to disease. Genera ABDEOMYIA, laya. p 79-109. genzs AEDES (subgenera [Aedes], Aedimorphus, Mogi M, Miyagi I, Suzuki H. 1981. New locality records Chaetocruiomyia, Christophersiomyia, Edwardsaedes of five mosquito species in Japan. Jpn J Sanit Zool 32: and Finlaya). Monograph Series. Entomology Mono- 124-126. graph 2. Canberra, Australia: School of Public Health Monchadskii AS. 1951. The larvae of bloodsucking mos- and Tropical Medicine, Australian Government Publish- quitoes of the USSR and adjoining countries (subfam. ing Service. ). Tabl Anal Faune URSS Moscow Zool Inst Lee KW. 1987. Checklist of mosquitoes (Culicidae) in Ko- Akad Nauk SSSR 37:l-290. rea. Korean J Parasitol 25:2O7-2O9. [In Korean with Nakata G. 1956. A new species of mosquito, Aedes (Fin- English summary.l laya) kabayashii n. sp.: from Kyoto, lapan. Jpn J Sanit Lee KW, Egan PJ. 1985. Illustrated taxonomic keys to Zool 7:135. [In Japanese.] genera and species offemale mosquitoes of Korea Part Nakata G. 1959. Redescription of Aedes (Finlaya) kobay' I. San Francisco, CA: Department of the Army, 5th Pre- csftii Nakata, 1956, Jpn J Sanit Zool 1O:16-2O + 2 pL' ventative Medicine Unit, 18th Medical Coinmand. Peytori EL, Pecor JE, Gaffigan TV. Trpis M, Rueda LM, Lee KW Lien JC. l9'7O. Pictorial keys to the mosquitos Wilkerson RC. 1999. The Johns Hopkins University of Korea WHO/VBC 70.196. Geneva, Switzerland: School of Hygiene and Public Health. Lloyd E. Roze- World Health Organization. boom mosquito collection. J Arn Mosq Control Assoc Lee KW, ZorkaT. 1987. Illustrated taxonomic keys to gen- l5:526-551. era and species of mosquito larvae of Korea. Part II Rao TR, Rajagopalan PK. 1957. Observations on mos- San Francisco, CA: Department of the Army, 5th Pre- quitoes of Poona District, India, with special reference ventative Medicine Unit, l Sth Medical Command. to their distribution, seasonal prevalence and the biol- Lee VH, Nalim S, Olson JG, Gubler DJ, Ksiazek TG, Aep ogy of adults. Indian J Malariol ll:l-54. S. 1984. A survey of adult mosquitoes of Lombak Is- Rattanarithikul R, Harrison BA. 1988. Aedes (Finlaya) re- land, Republic of Indonesia. Mosq News 44:784-191. inerti, a new species from northern Thailand felated to Lei X, Yang C, Lu B. 1984. Notes on Aedes (E) chris- Aedes (Finlaya) formosensis Yamada (Diptera: Culici' tophersi [sic] new to China (Diptera: Culicidae). Sdcy't- dae). Mosq Syst 20:77-96. uan J 7,ool 9:8-9. [In Chinese with English summary.] Reinert JE 1990. Medical entomology studies-XVll. Lien JC. 1967. New species of mosquitoes from Taiwan Biosystematics of Kenknightia, a new subgenus of the (Diptera: Culicidae) Part I. Two new species of Aedes mosquito genus Aedes Meigen from the Oriental Re- (Finlaya) lsicl. Trop Med 9:177-185. gion (Diptera: Cuticidae). Contrib Arn Entomol Inst Lien iC. 1968. New species of mosquitoes fiom Taiwan ( Gaine sville ) 26(2):1-l 19. (Diptera: Culicidae) Part V. Three new subspecies of Reinert JE 1999. Descriptions of Zavortinkius, a new sub- Bn u c rn tnnts ot ttu.s New SuscpNusop Oc n tp portru s 321

genus of Aedes, and the eleven included species from Washington, DC: Thornas Say Foundation, Entomolog- the Afrotropical Region (Diptera: Culicidae). Contrib ical Society of America. Am Entomol Inst (Gainesville) 31(2):l-1O5. Thnaka K. 1971. Mosquitoes of the Ryukyu Islands. Jpn Reinert JF 20O0a. New classification for the composite J Sanit Zool 2l:4. gents Aedes (Diptera: Culicidae: Aedini), elevation of Thnaka K. 2002. Studies on the pupal mosquitoes of Japan subgenus Ochlerotatus to generic rank, reclassification (6) Aedes (Finlaya) (Diptera, Culicidae). Jpn J Syst En- of the other subgenera, and notes on certain subgenera tomol 8:137-17'7. and species. J Am Mosq Control Assoc 16:175-188. Thnaka K, Mizusawa K, Saugstad ES. 1979. A revision Reinert JF 2000b. Comparative anatomy of the female of the adult and larval mosquitoes of Japan (including genitalia of genera and subgenera in tribe Aedini (Dip- the Ryukyu Archipelago and the Ogasawara Islands) tera: Culicidae). Part I. Introduction, preparation tech- and Korea (Diptera: Cr.tlicidae). Contrib Am Entomol niques, and anatomical terminology. Contrib Am En- Inst (Ann Arbor) 16:1-98'7. tomol Inst ( Gainesville ) 32(2): l-18. Tanaka K, Saugstad ES, Mizusawa K. 1975. Mosquitoes Reinert JF.2OO2. Comparative anatomy of the female gen- of the Ryukyu Archipelago (Diptera: Culicidae). Mosq italia of genera and subgenera in tribe Aedini (Diptera: Syst 7:2O7-233. Culicidae). Part XIII. Genus Ochlerotdtus Lynch Arri- Taylor FH. 1934. A check list of the Culicidae of the Aus- balzaga, Contrib Am Entomol Inst (Gainesville) 33(l): tralian region Service Publication (School of Public 1-lI1 Health and Tiopical Medicine) 1. Glebe, Australia: Reisen WK, Basio RG. 1972. Oviposition trap surveys Commonwealth of Australia, Department of Health, conducted on four USAF installations in the western Australasian Medical Publishing Co. Pacific. Mosq News 32:107-108. Theobald FV. 1901. A tnonograph of the Culicidae or Saini RS, Mishra AK. 1984. Mosquito fauna of Rewa. mosquitoes. Mainly compiled from the collections re- Geobios New Rep 3:57-58. ceived at the British Museum from various parts of the Saugstad ES. 1973. Two mosquito species new to the world in connection with the investigation into the Ryukyu Islands. Jpn J Sanit Zool 23:284. cause of malaria conducted by the Colonial Office and Sazonova ON. 1981. Mosquitoes of the Soviet Far East. the Royal Society London, United Kingdom: British Cytogenetics and genetics of vectors Proceedings of the Museum (Natural History). Symposium of the XVI International Congress of En- Theobald FV. 1903. A monograph of the Culicidae or tomology. New York, NY Elsevier Biomedical Press. mosquitoes. Mainly compiled from the collections re- p 65-74. ceived at the British Museum from various parts of the Scanlon JE, Esah S. 1965. Distribution in altitude of mos- world in connection with the investigation into the quitoes in northern Thailand. Mosq News 25:13'7-144. cause of malaria conducted by the Colonial Office and Senior-White R. 1923. Catalogue of Indian insects part the Royal Sociery, London, United Kingdom: British 2-Culicidae Calcutta, India: Superintendent of Govern- Museum (Natural History). ment Printing. Theobald FV. 1905. Genera insectorum. Diptera, fam. Cu- Senior-White R. 1926. Physical factors in mosquito ecol- licidae Fascicle 26. Bruxelles, Belgium: M. P Wytsman. ogy. Bull Entomol Res 16:187-248, Theobald FV. 1907. A monograph of the Culicidae or Senior-White R. 1927. Notes on Ceylon mosquitoes,-Il. mosquitoes. Mainly compiled from collections received The larvae of the commoner non-anopheline mosqui- at the British Museum London, United Kingdom: Brit- toes. Spolia Zeylan Bull Nat Mus Ceylon 14:61-76 ish Museum (Natural History). Stackelberg AA. 1943. A new species of Finlaya, Tlteo. Theobald FV. 1910. A monograph of the Culicidae or (Diptera, Culicidae) from USSURI Land. Bull Entomol mosquitoes- Mainly compiled from collections received Res 34:311. at the British Museum London, United Kingdom: Brit- Steffan WA. 1966. A checklist and review of the mos- ish Museum (Natural History). quitoes of the Papuan Subregion (Diptera: Culicidae). Thurman EB. 1959. A contribution to a revision of the J Med Entomol 3:179-237. Culicidae of northe rn Thailand Br:Jletin A- 100. College Stojanovich CJ, Scott HG. 1995. Mosquitoes of Asiattc Park, MD: University of Maryland Agricultural Exper- Rzzssia Metairie, LAr Published by Authors. iment Station. Stojanovich CJ, Scott HG. 1996. Mosquitoes of Korea Thurman EB. 1963. The mosquito fauna of Thailand (Dip- Metairie, LA: Published by Authors. tera: Culicidae). Pac Sci Congr (Entomol) 9:47-57. Stone A. 1961. A synoptic catalog of the mosquitoes of Toma T Miyagi I. 1981. Notes on the mosquitoes col- the world supplement I (Diptera: Culicidae). Proc En- lected at forest areas in the northern part of Okinawa- tomol Soc Wash 63:29-52. jima, Ryukyu Islands, Japan. Jpn J Sanit Zool 32:271- Stone A. 1970. A synoptic catalog of the mosquitoes of 279. Un Japanese with English summary.l the world, supplement IV (Diptera: Culicidae). Proc Toma T, Miyagi I. 1986. The mosquito fauna of the Ryu- Entomol Soc Wash 72:131-171 . kyu Archipelago with identification keys, pupal descrip- Stone A, Delfinado MD. 1973. Family Culicidae. In: Del- tions and notes on biology, medical importance and dis- finado D, Hardy DE, eds. A catalog of the Diptera of tribution. Mosq Syst 18:l-109. the Oriental Region Yoltrme I Suborder Nematocera Toma T Miyagi I. 1992. A survey of larval mosquitoes Honolulu, HI: The University Press of Hawaii. p 266- of Kume Island, Ryukyu Archipelago, Iapan. J Am -14-f _ Mosq Control Assoc 8:423-426. Stone A, Knight KL. 1956. Type specimens of mosquitoes Townsend FC, Chainey JE, Crosskey RW, Pont AC, Lane in the United States National Museum: II, The genus RR Boorman JPI Lowry CA. 1990. A catalogue of the Aedes [sic] (Diptera, Culicidae). J Wash Acad Sci 46: types of bloodsucking,fles Occasional Papers on Sys- 2t3,228. tematic Entomology 7. London, United Kingdom: Nat- Stone A, Knight KL, Starcke H. 1959. A synoptic catalog ural History Museum. of the mosquitoes of the world (Diptera, Culiciauel Tsukamoto M, Miyagi I, Toma t Sucharit S, Tumrasvin -)zz JounNal oF THE AMERICAN MoseulTo CoNrnol Assocr.qrroN 19,No. 4

W, Khamboonruang C, Choochote W, Phanthumachinda ma Island and the overwintering of Japanese encepha- B, Phanurai P. 1987. The mosquito fauna of Thailand litis virus. Trop Med 17:187-199. "A (Diptera: Culicidae): an annotated checklist. Jpn J Trop Ward RA. 1992. Thtrd supplement to catalog of the Hyg 15:291-326. mosquitoes of the wnrld" (Diptera: Cu,licidae). Mosq van den Assem J. 1961. Mosquitoes collected in the Hol- Syst24'.177-23O. landia area, Netherlands New Guinea, with notes on the Wattal BL. Bhatia ML. Kalra NL. 1958. Some new re- ecology of larvae. Tiidschr Entomol 1O4:17-30. cords of culicines of Dehra Dun (Uttar Pradesh) with a Wada X Mogi M, Oda T, Mori A, Suzuki H, Hayashi K, description of a new variety. Indian J Malariol 12:217- Miyagi I. 1976. Notes on mosquitoes of Amami-Oshi- 230.