MEDICAL ENTOMOLOGY STUDIES - Iv.

THE SUBGENERA INDUSIUS AND

EDWARDSAEDES OF THE GENUS AEDES

(DIPTERA: CULICIDAE)!,

John F, Reinert

ABST RACT

The subgenera Indusius Edwards and Edwardsaedes Belkin are redescribed

and compared to other subgenera of the genus Aedes Meigen. Species assigned

to the subgenera are fully illustrated and described. An analysis of the varia-

tion in setal branching in larvae and pupae of imprimens (Walker) is presented.

Aedes suknaensis (Theobald) is returned to synonymy with imprimens.

INTRODUCTION

The monotypic subgenus Jndusius of the genus Aedes Meigen was described

by Edwards in Barraud (1934) and he included in it a single species, pulveru-

lentus Edwards. Two males were doubtfully included by him with females of

pulverulentus on page 269 but later (page 445) assigned by him with little doubt

to this species. He based the subgenus primarily on the marked sexual dimor-

phic features of the male which readily distinguished it from other Aedes ; how-

ever, he could not find any definite features with which to distinguish the fe-

male from other subgenera of the genus. The subgenus has not been treated in

_ the published literature since its original designation except very briefly ina

key by Mattingly (1961) and lists by Horsfall (1955) and Singh (1974). Aedes

pulverulentus also has not been reported since its inclusion in Indusius except

by Aslamkhan (1971) in a list of mosquito species from Pakistan.

Edwardsaedes, also a monotypic subgenus of the genus Aedes, was des-

cribed by Belkin in 1962. The type species, imprimens, however, had been

assigned to several different subgenera of Aedes since its original placement

in the genus Culex Linnaeus by Walker (1860) and these are as follows: Och-

lerotatus Lynch Arribalzaga; Aedimorphus Theobald (= Ecculex Felt); and

Neomelaniconion Newstead (= Banksinella Theobald).

IThis work was supported in part by Research Contract DAMD-17-74-C-4086

from the U.S. Army Medical Research and Development Command, Office of

the Surgeon General and carried out at the Medical Entomology Project,

Smithsonian Institution, Washington, D.C. 20560.

2Major, Medical Service Corps, U. S. Army, Department of Entomology,

Walter Reed Army Institute of Research, Washington, D.C. 20012.

2 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

The present study redescribes the subgenera Indusius and Edwardsaedes

and presents characters which separate each, in all the known stages, from

the other subgenera of Aedes. The single species included in Indusius, pulver-

ulentus, and the single known species of Edwardsaedes, imprimens, are fully

described and illustrated. Tables 1 and 2 list the range and mode of the setal

branching of the pupae and larvae of imprimens.

Many previously unstudied specimens of imprimens were examined during

this review. These specimens expanded the known range of imprimens and

filled in some gaps in the reported distribution of the species. Little material

of pulverulentus was available for study and most specimens were in very poor

condition. All specimens of this species examined were collected from 1911 to

1927 with the exception of 3 females which were taken in 1970 by M. Aslamkhan,

Pakistan Medical Research Center, Lahore, Pakistan.

Abbreviations used in references to literature conform to the BIOSIS List

of Serials, Biosciences Information Service of Biological Abstracts, Philadel-

phia, 1972. Inthe synonymy section, an asterisk following the abbreviations

used (A = adult, 2 = female, “ = male, P = pupa, L = larva, E = egg) indicates

that at least some portion of that sex or stage is figured. In the distribution

section abbreviations used are the same as in the synonymy but with the follow-

ing 2 additions, p = pupal skin and 1 = larval skin. Ten specimens were used

in determining the range and mode of the setal branching in pupal and larval

descriptions and tables. In the pupal description and table the number of

branches on abdominal seta 1-I was measured on the basal third of the seta.

Measurements on the illustrations are in millimeters. Distribution records

are indicated as follows: countries are in capital letters, provinces and pri-

mary administrative divisions are in italics, and place names have the first

letter capitalized. The number of specimens examined from each province

follows the last place name of the province in the distribution section. The

spelling of provincial and locality names was taken from the following Official

Standard Names Gazetteers prepared by the Geographic Names Division, U. S.

Army Topographic Command and the Office of Geography, U.S. Department of

the Interior: India (vols. Iand II, 1952); Indonesia (1968); Japan (1953); Malay-

sia (1970); Pakistan (1962); Philippine Islands (vols. I and II, 1953); South Viet-

nam (1971); and Thailand (1966). Locality names which did not appear in the

eazetteers were spelled according to the collection data sheets and labels on

the specimens.

Information in the bionomics and distribution sections was taken from the

collection data sheets, specimen labels and the published literature.

Nomenclature and chaetotaxy used for the female, male, male genitalia,

pupa and larva follow Knight (1970, 1971), Knight and Laffoon (1970a, 1970b,

1971a, 1971b) and Laffoon and Knight (1973). The terminology of the female

genitalia follows Reinert (1974a).

GENUS AEDES MEIGEN

SUBGENUS INDUSIUS EDWARDS

Type species: Aedes pulverulentus Edwards

Aedes (Indusius) Edwards, in Barraud 1934: 268, 445.

FEMALE. Head. Pedicel of antenna with a few small scales and short

Reinert: Subgenera Jndusius and Edwardsaedes 3

fine hairs mesally; vertex covered with broad decumbent scales, erect and

narrow scales absent; occiput with a few short erect forked scales; eyes wide-

ly separated; interocular space covered with broad scales and interocular

setae well developed. Thovax. Scutum covered with narrow curved scales;

scutellum with broad and narrow scales; acrostichal area with a few setae on

anterior area, setae absent on posterior area; dorsocentral area with a few

setae on anterior area and numerous ones on posterior area; scales present

on antepronotum, postpronotum, propleuron, subspiracular area, postspiracu-

lar area, mesepisternum and mesepimeron but absent on prosternum, para-

tergite, mesomeron and metameron; postpronotum with a number of posterior

setae; propleuron with 6-10 setae; mesepimeron without lower setae. Legs.

Femora I-III short and somewhat thickened; tibiae I-III with numerous erect

stout dark setae, very noticeable on III; posttarsi I and II with ungues equal in

size, each with a tooth, III with ungues equal in size, both simple. Wing.

Alula with a number of very long narrow scales along margin (scales similar

to setae on upper calypter); yemigium without setae. Genitalia, Segment VIII

completely retracted into segment VII; tergum VIII without scales; sternum

VIII with a deep median apical indentation, scales absent or only a very few

present, setae 1-3-S all basad; tergum IX consists of 2 long plates which are

expanded apically, each with several setae; insula tongue-like with a very few

small tuberculi; lower vaginal sclerite absent; upper vaginal sclerite well

developed, heavily pigmented; postgenital lobe with a moderately deep median

indentation; cerci moderately long, scales absent; 3 seminal capsules, one

large and 2 medium sized ones.

MALE. Head, Antenna slightly plumose, noticeably longer than proboscis;

maxillary palpus approximately equal in length to proboscis, 4-segmented,

basal 2 segments short, segment 3 long and approximately one half length of

apical segment which is long and swollen, with numerous short setae; probos-

cis short, apical half swollen and somewhat flattened, numerous long setae on

lateral and ventral areas, length shorter than length of femur I, labellum

large. Legs. PosttarsiI and II each with 2 ungues, empodium large, I with

ungues large, simple and equal in size, II with ungues large, unequal in size,

larger one with a tooth, smaller one simple. Abdomen. Terga with several

short setae, mostly along lateral and posterior margins. Genitalia. Tergum

IX with apex convex, several short setae on each side of midline; gonocoxite

moderately long, broad, somewhat conical, ventral surface with several long

setae on basal portion and a number of short ones scattered over remainder of

area, distal 0.6 covered with a pile of long hair-like spicules; gonostylus short,

broad, with file-like ridges and stout spicules, attached subapically to gono-

coxite; proctiger short with paraproct divided into 2 narrow arms and a broader

basal area, cercuSs membranous without cercal setae; phallosome with param-

eral apodeme moderately long, broad and with a long curved projection on its

caudomesal margin, sternum IX narrow, moderately long, setae absent.

PUPA, LARVA and EGG. Not known.

DISCUSSION. The adults of Indusius are distinguished from the other sub-

genera of Aedes by the following combination of characters: (1) eyes widely

separated and interocular space with broad scales and 3-5 pairs of setae; (2)

vertex of head with only broad decumbent scales; (3) antennal pedicel of female

with only a few small scales and short hairs mesally; (4) anterior and posterior

dorsocentral setae present; (5) a few anterior acrostichal setae present, posteri-

or setae absent; (6) prosternum and paratergite bare; (7) propleuron with 6-10

setae; and (8) wing without remigial setae and alula with several very long thin

scales on margin.

4 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

The very long thin scales of the alula are similar to those of Aedes (Notho-

skusea) chathamicus Dumbleton, Aedes (Halaedes) australis (Erichson), Opifex

fuscus Hutton and species of Culiseta Felt. The feature of widely separated

eyes is similar to Section B of Aedes (Verrallina) Theobald (see Reinert 1974b).

A number of features of the male differ markedly from the other subgenera

of Aedes as follows: antenna only slightly plumose and very long; development

of the maxillary palpus; shape of the proboscis; leg I with femur short and

somewhat thickened, tarsus short with tarsomeres 2-4 very short, posttarsus

with a pair of very large simple ungues and an enlarged empodium; and scutum

with only sparsely scattered narrow curved scales. The enlarged empodium of

the tarsi is similar to that of Deinocerites Theobald while the shortened femur

and tarsus and enlarged ungues resemble those of Opifex fuscus.

The male genitalia display characters of the subgenus that are unusual and

are as follows: gonocoxite broad, somewhat conical, with numerous short and

several long setae and a dense pile of long hair-like spicules; gonostylus short,

expanded, covered with numerous moderately long spicules and apical portion

with file-like ridges; proctiger with paraproct consisting of a narrow caudally

projecting scleritized strip along lateral margin of cerci, an anterior projection

and a broader basal area; shape of tergum IX; and parameral apodeme with a

long curved projection attached to caudal mesal margin. The gonocoxite is

developed somewhat like that of Opifex Hutton while the file-like ridges of the

sonostylus is similar to that of some Paraedes Edwards. The long curved

structure of the parameral apodeme is attached at the same point as the opistho-

phallus of Verrallina, however, it curves ventrally and caudally and does not

form a bridge to the other parameral apodeme.

Indusius possesses several interesting morphological features which are

similar to those of the more primitive members of the tribe Aedini, however,

due to the absence of specimens of immature stages, lack of any biological

information of the immatures and very poor taxonomic condition of the present

adult specimens, its phylogenic relationships and exact taxonomic status re-

main in doubt.

AEDES (INDUSIUS) PULVERULENTUS EDWARDS

(Figs. 1-4)

Aedes pulverulentus Edwards 1922a: 273 (9).

Aedes (subgenus uncertain) pulverulentus of Edwards 1922b: 468.

Aedes (?0chlerotatus) pulverulentus of Barraud 1928b: 374 (@).

Aédes (Indusius) pulverulentus of Barraud 1934: 268 (2,%); Edwards im Barraud

1934: 445 (o'*),

Aedes (Indusius) pulverulentus of Horsfall 1955: 410; Stone et al. 1959: 199;

Aslamkhan 1971: 153; Aslamkhan 1972: 101; Stone and Delfinado 1973:

300.

FEMALE (Fig. 1). Head. Antenna dark brown, 1.03-1.12 length of pro-

boscis, pedicel golden-brown with a few small broad white scales and a few

short fine golden hairs mesally, 13 flagellomeres, flagellomere 1 mostly ©

golden-brown and with several small broad white scales near middle, flagel-

lar whorls with 6 dark brown setae; clypeus dark brown, bare; maxillary pal-

pus dark brown scaled with a few white scales intermixed, 4-segmented, 0.19-

0.25 length of proboscis; proboscis dark brown scaled with a few white scales

intermixed, 1.19-1.27 length of femur I; eyes widely separated; interocular

Reinert: Subgenera Indusius and Edwardsaedes a

space covered with broad white scales, 4-5 pairs of long white interocular

setae; several long brown ocular setae; scales on head all broad and decum-

bent except for a small patch of short erect forked white scales on occiput,

vertex covered with white scales, lateral surface covered with dark brown

scales and with a stripe of white ones extending from area in front of ante-

pronotum to ocular line, postgena white scaled or mostly white scaled with

small patches of dark brown scales. Thorax. Scutal integument blackish-

brown; scutum densely covered with narrow curved white scales except for

narrow curved reddish-brown ones on following areas: a few at scutal angle,

a patch on anterior portion of supra-alar area and a small patch on posterior |

median scutal area, some specimens with a large median area of scutum

covered with dusty-white scales; prescutellar space with only median area

without scales; scutellum with a large patch of broad white scales on median

lobe, lateral lobe with a shaggy patch of long narrow curved white scales;

white to brown setae on following areas: 4-8 median anterior promontory,

.1-7 (usually 1-3) anterior acrostichals, posterior acrostichals absent, 1-6

(usually 1-3) anterior dorsocentrals, numerous posterior dorsocentrals, scu-

tal fossal (5-10 anterior and 5-12 lateral), numerous supra-alar, 4 posterior

medial scutal, 1 postalar callar and scutellar (7-11 long and 6-16 short ones

on lateral lobe, 6-10 long and 4-6 short ones on median lobe); pleural integu-

ment blackish-brown; antepronota widely separated, medium size, covered

with narrow curved scales, upper ones reddish-brown and lower ones white,

16-23 setae, upper ones brown and lower ones white; postpronotum with nar-

row curved reddish-brown scales on upper margin and most of remainder

covered with narrow curved white scales, lower white ones somewhat broader,

0-11 posterior brown setae; propleuron with a large patch of broad white

scales, 6-10 pale brown setae; prosternum, paratergite, mesomeron, meta-

meron and mesopostnotum bare; subspiracular area with a long patch of broad

white scales; postspiracular area with a patch of broad white scales, 3-8 pale

brown setae; mesepisternum with a large upper and a small lower patch of

broad white scales, 2-6 upper and 8-14 posterior brown setae; prealar knob

with several broad white scales, 13-21 pale brown setae; mesepimeron with a

large patch of broad white scales near center, 8-19 golden-white setae dorsad

of scale patch. Legs. Coxae I-III each with several brown setae, I covered

with broad white scales and with a patch of broad brown ones on anterior area,

II with a long patch of broad white scales on anterior area, III with broad white

scales on anterior and dorsal areas; trochanters I-III each with broad white ©

scales; femora I-III white scaled with a few to several brown scales inter-

mixed (brown scales more numerous on I and II); tibia I-III white scaled with

some brown scales intermixed, a number of moderately long erect brown

setae, longer and stouter on III; tarsi I-III brown scaled with white scales inter-

mixed, posteriors of I and II white scaled, III with tarsomeres 1-4 each with a

narrow basal white scaled band, band occasionally indistinct on 4 and some-

times present on 5; posttarsi I-III (Fig. 3) each with 2 large ungues, I and II

with ungues equal in size, each with a tooth, III with ungues equal in size, both

simple. Wing. Dorsal and ventral veins mostly covered with broad white

scales except for the following brown scaled areas: apical 0. 60-0. 65 of an-

terior surface of costa, a few intermixed on basal 0.35-0. 40 of costa, base

and anterior surface of remigium, a median stripe on radius, some on apical

portions of radius 2, 3, 4+ 5, median1+2, 3+ 4, cubitus 1 and 2; alula with

a number of long narrow white scales on margin; upper calypter with a number

of white setae on margin; remigial setae absent. Halter. Pedicel pale; capi-

tellum white scaled. Abdomen. Tergum I white scaled with a few brown

6 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

scales intermixed, laterotergite covered with white scales, II-IV each brown

scaled with a dorsobasal white scaled band which connects with a large irregu-

larly shaped laterobasal white scaled patch which has a small median brown

scaled area, V and VI mainly creamy-white scaled with numerous brown scales

intermixed, each with an indistinct dorsobasal band and laterobasal patch of

white scales; sterna white scaled, III-V each with a small ovoid lateroapical

patch of brown scales; terga and sterna with a number of golden-brown setae,

mostly along posterior margins. Genitalia (Fig. 2). Tergum VIII moderately

pigmented, wide at base and tapering to a moderately broad flat apex, base

nearly straight, numerous short and a few moderately long thin setae scat-

tered over apical 0.94-0.97, basolateral seta absent, covered with minute

Spicules, scales absent, completely retracted into segment VII, VIII-Te index

0.64-0.79, VIIl-Te/IX-Te index 1.32-1.48, length 0.18-0.23 mm, width 0. 24-

0.35 mm; sternum VIII moderately pigmented, moderately wide, base slightly

concave, apex with a deep median indentation, numerous short and a few mod-

erately long and long setae scattered over apical 0.98-0.99, setae on apex short

and thin, setae 1-3-S close together, all basad, 1-S basomesad, 2-S laterad of

1-S, 3-S laterad and slightly distad of 2-S, covered with minute spicules, 0-4

broad scales on basomesal area (usually scales absent), apical intersegmental

fold unpigmented, VIII-S index 1.02-1.10, length 0.28-0.30 mm, width 0. 26-

0.29 mm; tergum IX composed of 2 long narrow moderately pigmented plates

which are connected mesally by a membrane, apical portion of each plate ex-

panded and with 6-14 short thin setae, covered with minute spicules, [X-Te

index 1.31-1.40, length 0.14-0.16mm, width 0.10-0.12 mm; insula moderately

pigmented, tongue-like, covered with minute spicules, 1-2 small tuberculi on

apical area; lower vaginal lip moderately pigmented, narrow, covered with

minute spicules, lower vaginal sclerite absent; upper vaginal lip moderately

to heavily pigmented, narrow, covered with minute spicules; upper vaginal

sclerite large, moderately pigmented with basomesal area produced into a

narrow heavily pigmented structure; postgenital lobe moderately long, mod-

erately wide, apex with a moderately deep median indentation (0. 26-0. 32 deep),

6-10 setae on each side of midline, 12-19 total setae, completely covered with

minute spicules, basal median apodeme well developed and heavily pigmented,

dorsal PGL index 0.85-0.95, ventral PGL index 1.65-1.67, ventral length

0.09-0.10 mm; peri-anal membrane with minute spicules; cercus moderately

long, apex bluntly pointed, mesal margin nearly straight, outer margin bowed,

completely covered with minute spicules, dorsal surface with short and

moderately long setae on apical 0.92-0.95, 2 long setae at apex, scales ab-

sent, ventral surface with a few short and moderately long setae along mesal

and outer margins, index 2.43-2.53, cercus/dorsal PGL index 3.33-4. 14,

length 0.19-0.21 mm; 3 heavily pigmented, spherical seminal capsules, one

large and 2 medium sized ones, each with a few small seminal capsule pores

near orifice, base of accessory gland duct moderately pigmented.

MALE (Fig. 3). Only 2 badly rubbed males known, one of which is mounted

on a microscope slide and has the genitalia missing. Head. Antenna slightly

plumose, dark brown, very long, 1.86-1.88 length of proboscis; 13 flagello-

meres, flagellar whorls with 10-12 setae; clypeus brown, bare; maxillary

palpus dark brown scaled, 4-segmented, basal 2 segments short, segment 3

long and approximately 0.5 length of apical segment which is long and swollen,

1.01-1.03 length of proboscis, with numerous short brown setae; proboscis

short, brown scaled, apical half swollen and somewhat flattened, numerous

long brown setae on lateral and ventral areas, labellum large, 0. 86-0. 91

Reinert: Subgenera Indusius and Edwardsaedes 7

length of femur I; scaling on head as in female; eyes widely separated; inter-

ocular space with broad white scales and 3 pairs of pale setae. Thorax. Scutal

integument mostly golden-brown; scutum with sparsely scattered reddish-

brown narrow curved scales, a narrow stripe of similar scales on acrostichal

area; scutellum completely denuded of scales; brown setae on following areas:

9 median anterior promontory, a few anterior acrostichals, posterior acro-

stichals apparently absent, numerous anterior and posterior dorsocentrals,

scutal fossal (6-8 anterior, 4-6 lateral and 2-4 posterior), numerous supra-

alar, 8 posterior medial scutal, 1 postalar callar and scutellar (10-11 on later-

al lobe and 8-10 on median lobe); pleural integument golden-brown; antepronota

widely separated, medium sized, scales absent, 11-12 brown setae; postpro-

notum without scales, 4 posterior brown setae and 2-3 small hairs; propleuron

with a few broad white scales, 6-8 brown setae; prosternum, subspiracular

area, paratergite, mesomeron, metameron and mesopostnotum bare; post-

spiracular area without scales, 4-6 brown setae; mesepisternum with a medium

size upper and a small lower patch of broad white scales, 3-4 upper and 7

posterior brown setae; prealar knob with a few white scales, 9-11 brown setae;

mesepimeron with a large patch of broad white scales near center, 7 brown

setae dorsad of scale patch. Legs. Coxae I-III each with several golden-brown

setae, I with a patch of broad white scales on anterior area, II and III each with

a small patch of broad white scales; trochanters I-III each with a few broad

white scales; femora I-III brown scaled, II and III each with lower anterior

areas pale scaled, I-III with posteriors brown scaled with large areas of white

scales; tibiae I-III each brown scaled, some pale scales on posterior areas of

I and II; tarsi I-III short, each brown scaled, I with tarsomeres 2-4 very short;

posttarsi I-II (Fig. 3) each with 2 ungues, empodium large, I with ungues large,

simple and equal in size, II with ungues large, unequal in size, larger one with

a tooth, smaller one simple, III missing on both specimens. Wing. Dorsal

and ventral veins rubbed, only brown scales present, anterior surface and base

of costa dark brown scaled; alula with a number of long narrow pale brown

scales on margin; upper calypter with a number of pale brown setae on margin;

remigial setae absent. Halter. Pedicel pale; capitellum pale brown scaled.

Abdomen. Badly rubbed, Tergum I brown scaled, laterotergite covered with

white scales, II-V each brown scaled with a laterobasal patch of white scales,

no apparent dorsobasal white scaled bands or stripes, remainder of abdomen

missing (except badly damaged genitalia of one specimen); sterna brown scaled

with a few pale scales; terga and sterna with several short setae on lateral and

posterior margins, tergum and sternum VIII (Fig. 3) with characteristic shapes.

Genitalia (Figs. 3, 4). Tergum IX heavily pigmented, long, apex convex,

covered with small hair-like spicules, 7-8 short setae on each side of midline;

gonocoxite moderately long, broad, somewhat conical, dorsal surface with a

few long thin setae on basal area, lateral surface with several long thin setae,

ventral surface with several long thin setae on basal area and a number of short

setae scattered over remainder of area, distal 0.6 covered with a pile of long

hair-like spicules, apical portion of mesal membrane with several short setae

and minute spicules (remainder of mesal membrane lost); gonostylus short and

broad, basal area moderately broad, apical area expanded, dorsal surface with

distal area covered with file-like ridges which terminate in a serrated apex, a

few minute setae at base of file-like area, ventral surface with numerous mod-

erately long spicules (many spicules stout) and a few minute setae, attached

subapically to gonocoxite, gonostylar claw absent; basal mesal lobe badly

damaged but a small tergobasal strip and a sternobasal strip each with several

short setae remain, probably consists of a large plate with numerous short

8 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

setae located on basal area of mesal membrane of gonocoxite, connected with

its mate by a heavily pigmented V-shaped structure; proctiger short, paraproct

heavily pigmented, consists of a caudally projecting strip along lateral margin

of cercus, an anterior arm and a broader basal area, cercus membranous with-

out cercal setae; tergum X short, covered with small spicules, base attached

to base of paraproct and apex attached to caudal area of tergum IX; phallosome

missing except for a moderately long and broad parameral apodeme which has

a long curved caudal projection attached to its caudomesal margin; sternum IX

heavily pigmented, narrow and moderately long, small spicules on apical por-

tion, setae absent.

PUPA, LARVA and EGG. Not known.

DISCUSSION. The female of pulverulentus is similar in habitus to Aedes

(Aedimorphus) gouldi Reinert, however, it is distinguished from gouldi by the

following: proboscis and maxillary palpus brown scaled; vertex with only

broad decumbent scales; eyes widely separated; scutal scales white; and re-

migial setae absent. The scutum of pulverulentus is white scaled and resembles

species in subgenera Christophersiomyia Barraud and Finlaya Theobald of

Aedes, but is easily distinguished from them by features listed for the sub-

genus.

Female genitalia of pulverulentus are similar to those of the subgenus

Aedimorphus but differ by the following features: shape of tergum VIII;

sternum VIII with a very deep median apical indentation and placement of

setae 1-3-S; and insula with only 1-2 tuberculi. |

The male shows a strong sexual dimorphism and differs from the female

most noticeably in the following: very long antenna, maxillary palpus, pro-

boscis, tarsus and posttarsus I, and scutal scaling. Other differences exist

between the 2 sexes in the scaling of the pleural areas, legs, wing and abdomen.

The single known male genitalia specimen, dissected and mounted on a

microscope slide, was very badly damaged and consisted of 16 pieces scattered

throughout the mounting medium underneath the cover glass. The specimen

was in such a poor condition it was necessary to remount the fragments so

that the component parts of the genitalia could be determined. A careful

search of the original preparation was conducted prior to remounting the

genitalia and some structures could not be found (e.g. phallosome) and are

presumed to have been lost. The above description and figures 3 and 4 area

result of the reconstructed genetalic fragments and are as complete as could

be determined from the poor specimen.

TYPE-DATA. The holotype female is in the British Museum (Natural

History) (BMNH) and bears the following information on the labels: Aedes

pulverulentus, Edw.; Type H. T.; India, Nowshera, N. W. P., 4. VII. 1911,

Smith [ collector], Capt. P. J. Barraud 1921-166; T74.67 Term. [genitalia

preparation number|. The holotype is in excellent condition and the genitalia

were mounted by me in Canada balsam on a microscope slide. The hand

written label on the holotype gives the collection date as 4. VII. 1911 which

differs from Edwards (1922a: 274) who lists the date as 4. vi. 1911. A paratype

female in the BMNH contains the same label data as the holotype except it

possesses the collection date of 4. VI.1911. This paratype is in good condition.

An additional female in the BMNH is dissected and mounted on a microscope

slide and bears the following information on the label: Aedes pulverulentus &,

Nowshera, N. W. India, per P. J. Barraud. I assume this specimen to be a

paratype.

DISTRIBUTION. 27 specimens examined: 259, 2c.

PAKISTAN. Karachi Federal Territory, Karachi; 162. Northwest Frontier,

Reinert: Subgenera Indusius and Edwardsaedes 9

_ Peshawar, Nowshera; 12 (holotype), 22 (paratypes). Punjab, Multan, Dera

Ghazi Khan, Dera Ismail Khan; 3¢, 20°. Larkana, Sind; 39.

Distribution from literature.

PAKISTAN (Aslamkhan 1971: 153); Karachi, Larkana, Sind, Mari Indus,

Nowshera (Barraud 1928b: 375), Indus Valley, Dera Ismail Khan (Barraud

1934: 269).

BIONOMICS. The 3 females from Dera Ghazi Khan were collected biting

cattle (14 July 1970, M. Aslamkhan). Barraud (1928b: 375) reports specimens

resting under an ice chest at Larkana, Sind.

SUBGENUS EDWARDSAEDES BELKIN

Type species: Culex imprimens Walker

Culex in part of Walker 1860: 144.

Culicada in part of Theobald 1910a: 21.

Ochleroitatus in part of Edwards 1913: 228.

Aedes (Ecculex) in part of Edwards 1922b: 467.

Aedes (Banksinella) in part of Edwards 1924: 371; Knight and Hull 1953: 465.

Aedes (Ochlerotatus ) in part of Edwards 1924: 377.

Aedes (Aedimorphus) in part of Edwards 1929: 5.

Aedes (Neomelaniconion) in part of Stone et al. 1959: 200.

Aedes (Edwardsaedes) Belkin 1962: 408; Huang 1968: 187.

FEMALE. Head. Antenna long, pedicel with a few scattered small broad

scales and short fine hairs mesally; maxillary palpus 5-segmented, segment 5

minute; proboscis noticeably longer than femur I; vertex covered with narrow

curved decumbent scales and numerous erect forked scales; occiput with nu-

merous erect forked scales. Thorax. Scutum covered with narrow curved

scales, similar scales on scutellum; acrostichal and dorsocentral setae numer-

ous; antepronotum with narrow curved scales, numerous setae; postpronotum

covered with narrow curved scales, 6-14 posterior setae; propleuron with

broad scales, 27-42 setae; subspiracular area with a patch of broad white

scales and several to numerous short pale hairs intermixed; postspiracular

area with scales, 8-18 setae; prealar knob with 17-40 setae; mesepimeron

with a large median patch of broad scales, 16-29 setae dorsad of scale patch,

lower setae absent. Legs. Posttarsi I-III with ungues equal in size and each

with a tooth. Wing, Alula with numerous narrow scales on margin; remigial

setae present. Genitalia. Tergum VIII and sternum VIII each with setae on

most of surface, scales usually absent but a very few occasionally present;

sternum VIII large, base and apex each with a deep median indentation, a

small lobe on each side of apical indentation; tergum IX long, apex with a mod-

erately deep median indentation and with several setae on each side of mid-

line; insula long, tongue-like, without setae or tuberculi; lower and upper

vaginal lips narrow; lower vaginal sclerite absent; upper vaginal sclerite

large, heavily pigmented and with a fragmented appearance; postgenital lobe

short, broad, a number of setae on each side of midline; cercus long, narrow,

apex sharply rounded, scales absent; 3 seminal capsules, one large and 2

slightly smaller ones, heavily pigmented, spherical.

MALE. Essentially as in the female. Head. Antenna with numerous long

dorsal-ventrally directed setae in each flagellar whorl; maxillary palpus longer

than proboscis, 5-segmented, segments 1 and 2 short, segments 3-5 long,

10 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

segment 5 and apical portion of segment 4 with numerous long erect lateral

and ventral setae, segment 5 upturned. Legs. Posttarsi I-III with 2 ungues

each bearing a tooth, I and II with ungues unequal in size. Abdomen. Terga

I- VII with numerous long curved setae along lateral margins, VIII witha

number of short stout heavily pigmented setae in a row along apical margin.

Genitalia. Tergum IX with 2 narrow lateral plates each with a caudomesal

lobe bearing several setae, median area between plates membranous; gono-

coxite short, broad, with setae and broad scales, mesal surface membranous,

gonostylus attached at apex; gonostylus moderately long and with setae, gono-

stylar claw absent; basal mesal lobe consists of a broad raised plate with nu-

merous setae, situated on basal area of membranous mesal surface of gono-

coxite; paraproct with a broad basal area and a very heavily pigmented claw-

like tergally curved apical area, cercus membranous except for a pair of small

pigmented rectangular plates on dorsal surface, cercal setae absent; phallo-

some with aedeagus long, narrow, heavily pigmented, apex sagittate, base

divided into 2 slender plates each with a few dorsobasal teeth, paramere mod-

erately long, attached subbasally to parameral apodeme, parameral apodeme

very long, attached near middle to apodeme of gonocoxite; sternum IX broad,

lateral, basal and median areas pigmented, several setae on median caudal

area.

PUPA. Respiratory trumpet. Short, heavily pigmented. Metanotal plate.

Seta 10-CT caudad and laterad of 11-CT. Abdomen. Setae 2, 3-I widely sep-

arated; 9-VII stout, multiple branched, directly cephalad of 6-VII; 9-VIII well

developed, multiple branched. Paddle. Outer margin with minute serrations

on basal area and minute spicules on apical area; 1-P short.

LARVA. Head. Seta 4-C small, multiple branched, mesad and far

cephalad of 5, 6-C; 5-7-C long, barbed, multiple branched; 6-C laterad and

slightly cephalad of 5-C; 7-C laterad and cephalad of 6-C; 12, 13-C approxi-

mated and removed from cephalic border of head; dorsomentum with numer-

ous teeth; ventromedian cervical sclerite well developed (see Fig. 10).

Antenna. Long, slender, with numerous well developed spicules; 1-A long,

multiple branched; 2-A with a subapical constriction. Thovax. Setae 9-12-M,

T located on large heavily pigmented plates bearing a long stout heavily pig-

mented apical spine. Abdomen. Seta 1-VII short, multiple branched; 13-VII

not widely separated from 10-VII; saddle completely rings segment X, acus

absent; 3-X multiple branched, only slightly longer than 2-X; ventral brush

with numerous long multiple branched setae, precratal ones inserted in ven-

tral margin of saddle. Szphon. Moderately long, acus present; pecten with

numerous teeth, a few apical teeth wider spaced than remainder; seta 1-S

small, multiple branched, base attached distad of last pecten tooth.

EGG. See description for imprimens.

DISCUSSION. The subgenus Edwardsaedes is separated from the other

subgenera of Aedes by the following features: in the adults by the combina-

tion of (1) head with vertex covered with narrow curved decumbent and numer-

ous erect forked scales, (2) antennal pedicel with a few scattered small broad

scales and short fine hairs mesally, (3) numerous anterior and posterior dor-

socentral and acrostichal setae, (4) propleuron with numerous (24-42) setae,

(5) prosternum bare, (6) subspiracular area with a patch of broad scales and

short fine hairs intermixed, (7) mesepimeron with 11-29 setae dorsad of scale

patch and without lower setae, (8) posttarsi I-III each with both ungues toothed,

(9) female maxillary palpus 5-segmented, apical segment minute, and (10)

male maxillary palpus long, apical segment upturned, numerous long erect

setae on lateral and ventral surfaces of segment 5 and apical portion of seg-

)

Reinert: Subgenera Indusius and Edwardsaedes 11

ment 4; in the female genitalia by the combination of (1) tergum VIII with setae

on most of its surface, scales usually absent, (2) sternum VIII shape, scales

absent, (3) insula long, tongue-like, setae and tuberculi absent, (4) cercus long,

narrow, scales absent, (5) upper vaginal sclerite large, heavily pigmented,

with a fragmented appearance, and (6) one large and 2 smaller seminal capsules;

in the male genitalia by the combination of (1) tergum IX membranous mesally,

each lateral plate with a caudomesal lobe bearing several short setae, (2)

gonocoxite short and broad, gonostylus attached at apex, (3) gonostylus without

a gonostylar claw, (4) basal mesal lobe a broad raised plate bearing numerous

setae, (5) proctiger with paraproct formed into a heavily pigmented, strongly

curved, claw-like structure, cercal setae absent, and (6) phallosome with

aedeagus long, narrow, apex sagittate, basal area with teeth, parameral apo-

deme very long; in the pupa by the combination of (1) setae 1-3-CT approxi-

mately equally developed, (2) seta 10-CT caudad and laterad of 11-CT, (3)

setae 2, 3-I widely separated, and (4) setae 9-VII, VIII well developed, mul-

tiple branched and with branches thickened and barbed; and in the larva by

the combination of (1) position and development of head setae 4-7-C, (2) an-

tenna long, spiculate, seta 1-A well developed and multiple branched, (3)

thoracic setae 9-12-M, T situated on large heavily pigmented plates bearing

a long heavily pigmented apical spine, and (4) segment X with saddle complete-

ly ringing segment, ventral brush with several setae inserted in ventral margin

of saddle and seta 3-X multiple branched and only slightly longer than 2-X.

Adults of Edwardsaedes are similar in habitus to those of Aedimorphus,

Ochlerotatus and Indusius but can be distinguished from these subgenera by the

features listed above. The male maxillary palpus resembles those of members

of the subgenus Neomelaniconion, however, the male genitalia and other adult

features (see above) differ between these 2 subgenera. The characteristic shape

of the apical portion of sternum VIII allows easy recognition of female Edward-

saedes, even in undissected specimens.

Pupae of Edwardsaedes resemble those of the following subgenera of

Aedes: Neomelaniconion, Aedimorphus, Verrallina and Aedes Meigen. How-

ever, they can be recognized by one or more of the characters listed above.

The pupae of this subgenus also possess setae 6-III-VII multiple branched which

separates them from a number of Aedes subgenera.

Larvae of Edwardsaedes are similar to those of subgenera Aedimorphus ,

Neomelaniconion, Vervallina and Aedes. They can be distinguished from these

subgenera, as well as all subgenera of Aedes, by the above combination of

features. Reinert (1973, 1974b) gives descriptions and illustrations of the

larvae and pupae of Aedimorphus and Verrallina. The subapical constriction

of antennal seta 2-A and the presence of precratal setae of the ventral brush

are shared by members of a number of Aedes subgenera whose larvae inhabit

temporary ground pools (Reinert 1974b). Presence of numerous multiple

branched setae of the ventral brush of segment X, antenna with well developed

spicules and antennal seta 1-A well developed also appear to be shared by lar-

vae of the genus Aedes collected from temporary fresh water ground pools as

compared to species collected from plant container water. The saddle which

rings segment X and with setae of the ventral brush attached along its ventral

margin is similar to those of the genus Psovophora Robineau-Desvoidy. De-

velopment of the ventromedian cervical sclerite is similar to those of Aedes

(Neomelaniconion) lineatopennis (Ludlow) and species of Aedimorphus and

Verrallina.

DISTRIBUTION. Aedes imprimens, the only included species of Edward-

saedes, exhibits a wide geographical range (Fig. 5). The species is found

12 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

throughout Southeast Asia, New Guinea, northern tip of Australia, and east

into the Solomon Islands, the northern range extends throughout Japan and into

South Korea, and the western limit is Bengal, India.

MEDICAL IMPORTANCE. Females of imprimens are aggressive man-

biters, readily bite through clothing and in some areas are troublesome pests.

Very few attempts have been made to isolate pathogens from this species even

though it readily feeds on man. Yamada (1927) in an experimental laboratory

study found zmprimens from Japan not to be a suitable intermediate host for

Wuchereria bancrofti (Cobbold).

BIONOMICS. Immatures usually inhabit temporary unmoving fresh water

in flood pools located in shaded areas in forests. Eggs are laid singly and re-

quire drying for 4-6 weeks prior to hatching. The larvae complete their de-

velopment within 3 days. Females are vicious daytime biters and readily

attack in shaded areas, even feeding through heavy clothing. Adults have

been collected resting in houses, stables, vegetation and in light traps. Speci-

mens have been taken at elevations from sea level to 1,200 m.

AEDES (EDWARDSAEDES) IMPRIMENS (WALKER)

(Figs. 5-11)

Culex imprimens Walker 1860: 144 (2); of Theobald 1901: 422 (2); Blanchard

1905: 306 (2); Brunetti 1907: 348; Brunetti 1912: 470; Theobald 1910b:

350.

Culex imprimiens of Giles 1900: 262 (2, lapsus calami); Giles 1902: 411 (&,

lapsus calami); Theobald 1905: 26 (lapsus calami).

Culex Auratus Leicester 1908: 153 (2); of Knight and Hull 1953: 466.

Culicada suknaensis Theobald 1910a: 21 (2); of Theobald 1910b: 297 (2*);

Edwards 1922b: 467 (synonymy); Barraud 1928a: 663 (questioned

synonymy with imprimens); Barraud 1934: 257 (transferred suknaensis

from imprimens to caecus). |

L. auratus of Brunetti 1912: 467 (lapsus calami).

Culex auratus of Edwards 1913: 228 (questionable synonymy with imprimens;

name preoccupied by Aedes auratus Grabham 1906); Edwards 1932:

170; Stone et al. 1959: 201; Mattingly 1961: 53; Belkin 1962: 409;

Stone and Delfinado 1973: 293.

Ochlerotatus imprimens of Edwards 1913: 228; Brunetti 1920: 136; Senior-

White 1923: 78.

Aedes (Ecculex) imprimens of Edwards 1922b: 467; Brug and Haga 1923: 639.

Aedes (Banksinella) brugi Edwards 1924: 371 (c'); of Edwards in Paine and

Edwards 1929: 314 (suspected synonymy with imprimens); Knight and

Hull 1953: 468 (suggested synonymy with imprimens); Stone et al. 1959: —

200 (questionable synonymy with imprimens); Belkin 1962: 409 (synony-

my).

Aedes (Ochlerotatus) imprimens of Edwards 1924: 377 (2); Edwards 1925: 297.

Aedes (Aedimorphus) imprimens of Edwards 1929: 5; Paine in Paine and Ed-

wards 1929: 306; Brug and Edwards 1931: 258; Edwards 1932: 170;

Taylor 1934: 20; Bonne-Wepster and Brug 1937: 38 (2*); Bohart 1945:

63 (L); Brug and Bonne-Wepster 1947: 185; Bonne-Wepster 1954: 239

(2); Assem and Bonne-Wepster 1964: 98 (2, L*).

Aedes (Aedimorphus) caecus (Theobald) of Edwards 1932: 170 (in part);

Barraud 1934: 257 (in part); Reinert 1973: 28 (transferred suknaensis

from caecus to subgenus Edwardsaedes).

Reinert: Subgenera Indusius and Edwardsaedes 13

Aedes (Banksinella) brugi of Edwards 1932: 170; Taylor 1934: 20; Iyengar 1955:

30.

Aedes imprimens of Edwards in Paine and Edwards 1929: 314; Knight et al.

1944: 35 (key); Stojanovich and Scott 1965: 9, 27 (2*, L*, -key);

Stojanovich and Scott 1966: 45, 117 (2*, L*, key).

Aedes brugi of Knight et al. 1944: 35 (key).

Aedes (Banksinella) imprimens of Hsiao and Bohart 1946: 22; La Casse and

Yamaguti 1950: 102 (2*, o); Waku 1950: 69 (2*, o&*, P*); Knight and

Hull 1953: 465 (2, o*, L*); Horsfall 1955: 527; Iyengar 1955: 30; Hara

1957a: 65 (2*), 1957b: 16 (2*); Macdonald 1957: 21.

Aedes (Neomelaniconion) brugi of Stone et al. 1959: 200; Iyengar 1960: 68.

Aedes (Neomelaniconion) imprimens of Stone et al. 1959: 201; Iyengar 1960:

68: Mattingly 1961: 53 (9*, o’*, P*, L*); Delfinado et al. 1962: 440:

Stone et al. 1966: 50, 57 (key).

Aedes (Edwardsaedes) imprimens of Belkin 1962: 409 (¢, o*, P*, L*); Stone

1963: 130; Kurihara 1963: 196 (A*); Huang 1968: 187 (A*, L*, key);

Stone and Delfinado 1973: 293; Moriya et al. 1973: 54 (E*).

Aedes (Neomelaniconion) impremens of Basio 1971: 24 (o*, lapsus calami).

FEMALE (Fig. 6). Head. Antenna dark brown, 0.91-1.00 length of pro-

boscis, pedicel golden colored with mesal area dark brown and with several

scattered small broad golden and brown scales and short fine brown hairs

mesally, flagellomere 1 with basal 0.50-0.65 pale and with several golden

and few small broad brown scales, flagellar whorls with 6 dark setae; clypeus

brown, bare; maxillary palpus dark brown scaled, 5-segmented, approximate

length of each segment in parentheses, 1 (0.15), 2(0.07), 3 (0.28), 4 (0.47)

and 5 (0.03), total length 0.20-0.23 length of proboscis; proboscis dark brown

scaled, usually with a few pale scales intermixed on ventral and occasionally

on dorsal surfaces, 1.21-1.30 length of femur I; eyes narrowly separated;

interocular setae absent; ocular setae long, brown and numerous; vertex

covered with narrow curved golden and white intermixed decumbent scales,

a few narrow curved reddish-brown scales on anterior portionof coronal

suture, numerous long erect forked golden scales; ocular line with small

narrow curved white or golden-white scales, similar scales on interocular

space; lateral surface with broad decumbent white scales except for a small

patch of dark brown similar scales on dorsoanterior area and a few brown

ones on the postgena; occiput with narrow curved decumbent white scales

and numerous long erect forked scales, median ones golden and several

lateral ones brown. Thorax. Scutal integument brown; scutum covered with

narrow curved bronzy scales except for narrow curved white scales on the

following: median anterior promontory area, anterior scutal fossal area and

extending onto lateral area, basal portion of scutal ridge, supra-alar area,

posterior median scutal area, and along margins of prescutellar space,

patch of narrow curved reddish-black scales on median scutal fossal area,

anterior portion of supra-alar area and occasionally among acrostichal and

dorsocentral setae; scutellum with narrow curved white scales, some with a

brown hue, on each lobe; scutal setae brown, numerous, on following areas:

6-10 median anterior promontory, numerous acrostichal (anterior and posteri-

or), numerous dorsocentral (anterior and posterior), scutal fossal (6-10 an-

terior, 3-9 lateral and 2-5 posterior), numerous supra-alar, 6-8 posterior

medial scutal, 1 postalar callar and scutellar (9-16 long and 4-7 short ones

on lateral lobe, 7-16 long and 4-9 short ones on median lobe); pleural integu-

ment pale brown with postpronotum, sub- and postspiracular areas and prealar

14 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

_knob darker brown; antepronota widely separated, covered with narrow curved

white scales, 25-45 brown setae; postpronotum nearly covered with narrow

curved scales, bronzy ones dorsally and white ones on remainder of area,

some specimens with a few broad white scales ventrally, 6-14 brown posterior

setae, several short pale hairs ventrad of setae; propleuron with a patch of

broad white scales, 27-42 golden to brown setae; prosternum, mesomeron and

metameron bare; subspiracular area with a patch of broad white scales, several

to numerous short pale hairs intermixed with scales; postspiracular area witha

patch of broad white scales, 8-18 golden to brown setae; paratergite moderately

broad, usually bare, however, a few specimens representing the entire range,

possess a few narrow white scales; mesepisternum with a large upper and a

medium size lower patch of broad white scales, 4-8 upper and 12-24 posterior

golden-brown setae, lower ones shorter and golden; prealar knob with 17-40

golden-brown setae; mesepimeron with a large patch of broad white scales on

middle, 16-29 golden setae dorsad of scale patch, lower setae absent (1 speci-

men from Thailand, 00168-111, had a single seta ventrad of scale patch on the

left side, this seta was poorly developed, kinky and considered to be an anomaly).

Legs. Coxae I-III each with several golden-brown setae, I with broad brown

scales, a dorsal and a ventral patch of broad white scales, II with an antero-

lateral patch of broad white scales, some specimens with a few broad brown

scales ventrally, III with a small anteroventral and a small posteroventral

patch of broad white scales; trochanters I-III with broad white scales and short

pale setae; femora I and II each with anterior surface dark brown scaled with

a narrow basal band of white scales, posterior surface dark brown scaled with

a longitudinal white scaled stripe, on median 0. 70-0. 85 of I and on ventral 0. 7-

0.8 of II, stripes broad basally and narrow apically, III white scaled with a nar-

row longitudinal dark brown scaled stripe which forms a broad band on apical

0.25-0.35, a few dorsobasal white scales, I-III each with a few apicolateral

white scales on posterior surface; tibiae I-III each dark brown scaled with a nar-

row basal band of white scales, band very narrow on I, posterior surface of I

and II dark brown scaled with a longitudinal pale scaled stripe from base to

apex, ventral on I and dorsal on IJ; tarsi I-III dark brown scaled, I with white

scaled narrow basal bands on tarsomeres 1-3, some specimens with only a dor-

sobasal white scaled spot on 3, II with white scaled narrow basal bands on tar-

someres 1-4, some specimens with only a dorsobasal white scaled spot on 4,

III with white scaled narrow basal bands on tarsomeres 1-5; posttarsi I-III (Fig.

11) each with 2 ungues, equal in size, each with a tooth. Wing. Dorsal and

ventral veins with dark brown scales except for a small white scaled patch at

base of costa; alula with numerous long narrow brown scales on margin; upper

calypter with a number of long brown setae on margin; 1-3 brown remigial

setae. Halter. Pedicel pale; capitellum with dorsal surface white scaled

apically and brown scaled basally, ventral surface white scaled. Abdomen.

Terga dark brown scaled, I with a large white scaled patch on laterotergite,

II- VII each with a large laterobasal white scaled patch, II-IV also with a dorso-

basal narrow white scaled band which connects with, or nearly with, laterobasal |

white scaled patches, V-VII usually with several dorsobasal white scales, some

specimens with white scales forming a narrow band on V, VI and VII with a few

to numerous golden-brown scales scattered over terga, a few specimens with

much of dorsal surface covered with pale scales; sterna yellowish-white

scaled, II- VI each with a narrow apical band of dark brown scales, VII of some

specimens with a few dark brown scales apically; terga and sterna with numer-

ous short and moderately long golden setae, mostly along posterior margins.

Genitalia (Fig. 7). Tergum VIII lightly pigmented, base strongly concave

Reinert: Subgenera Jndusius and Edwardsaedes 15

mesally, apex slightly concave or flat, lateral margins nearly straight, a>

number of short and moderately long thin setae scattered over entire surface,

occasionally a few short setae on membranous area near lateral and basal

margins, basolateral seta absent, covered with minute spicules, 0-7 (usually

0-3) small moderately broad scales, basal 0.75-1.00 retracted into segment

VII, VIII-Te index 0.93-1.04, VIII-Te/IX-Te index 1. 86-2.07, length 0. 33-

0.35 mm, width 0.32-0.44 mm; sternum VIII lightly pigmented with apical

0.35 moderately pigmented, large, long, base with a deep median indentation,

apex with a deep narrow median indentation and with a small lobe on each side

of indentation, a number of short and a few long setae scattered over entire

surface, setae 1-3-S widely separated, located in a more or less diagonal

line, 1-S basomesad, 2-S approximately 0.48 from 1-S, 3-S approximately

0.52 from 2-S, covered with minute spicules, 0-2 (usually absent) broad

scales, apical intersegmental fold unpigmented, VIII-S index 1.01-1.14,

length 0.39-0.44 mm, width 0.37-0.40 mm; tergum IX moderately pigmented

with midline slightly less pigmented, long, widest point near apical 0.28, apex

with a moderately deep median indentation and with 6-14 moderately long setae

on each side of midline, 15-26 total setae, covered with minute spicules, IX-

Te index 0.91-1.13, length 0.16-0.23 mm, width 0.14-0.21 mm; insula lightly

pigmented, long, tongue-like, without setae or tuberculi, covered with short

hair-like spicules; lower vaginal lip lightly pigmented, narrow, covered with

short hair-like spicules, lower vaginal sclerite absent; upper vaginal lip mod-

erately to heavily pigmented, narrow, covered with short hair-like spicules,

upper vaginal sclerite large, heavily pigmented with unequal pigmentation

toward inner caudal margin giving it a fragmented appearance; postgenital

lobe short, broad, apex with a shallow to small (0.05-0.12 deep) median

indentation, 7-14 setae on each side of midline, 15-26 total setae, covered

with short hair-like spicules, dorsal PGL index 0. 72-0.94, ventral PGL index

1.51-1.95, ventral length 0.13-0.17 mm; peri-anal membrane with short

spicules scattered over entire surface; cercus long, narrow, apex sharply

rounded, 2 long setae at apex, completely covered with minute spicules, dor-

sal surface with numerous short and moderately long setae on apical 0. 84-

0.90, scales absent, ventral surface with a few short setae along outer, apical

and inner margins, cercus index 3. 57-4. 33, cercus/dorsal PGL index 4.17-

0.81, cercus length 0.34-0.37 mm; 3 seminal capsules, one large and 2 slightly

smaller ones, heavily pigmented, spherical, each with a short pigmented neck,

several small seminal capsule pores near orifice, base of accessory gland

duct heavily pigmented.

MALE (Fig. 6). Essentially as in the female but with the following differ-

ences. Head. Antenna 0. 87-0.96 length of proboscis, numerous long brown

setae in each flagellar whorl, setae directed mainly dorsoventrally; maxillary

palpus 1.25-1.34 length of proboscis, brown scaled, 5-segmented, segments

1 and 2, 3 and 4 ankylosed, segment 5 and apical 0.6 of segment 4 with numer-

ous long brown erect setae on lateral and ventral surfaces, segment 5 upturned,

approximate length of each segment in parenthesis, 1 (0.04), 2 (0.02), 3 (0.22),

4 (0.38) and 5 (0. 34); proboscis 1.17-1.30 length of femur I; vertex and lateral

surface of head with dark decumbent scales absent. Thorax. Antepronotum with

several broad white scales in addition to a few narrow curved white ones; post-

pronotum with a few narrow curved golden-white scales and a very few broad

white ones ventrally, 5-9 brown posterior setae; propleuron and postspiracular

area without scales (occasionally a specimen with one or 2 broad white scales);

prealar knob with 13-19 golden-brown setae; mesepimeron with 11-19 golden

setae dorsad of scale patch. Legs. Tarsi I and II with pale scaling reduced;

16° Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

posttarsi I-III (Fig. 11) with 2 ungues each bearing a tooth, I and II with ungues

unequal in size, III with ungues equal in size. Wing. Pale scaled spot at base

of costa very small; 1 remigial seta. Abdomen. Terga brown scaled, II-VII

each with a narrow dorsobasal white scaled band, band somewhat broader me-

sally, laterotergite of I with a large white scaled patch; I- VII with numerous

long curved golden-brown setae along lateral margins, VIII (Fig. 11) witha

broad basal band of pale scales and a number of short stout heavily pigmented

setae in a row along apical margin; sterna II-VII golden scaled with a few

brown scales on apical margins, numerous long golden-brown setae, VIII

golden-white scaled with a narrow apical band of brown scales. Genitalia

(Fig. 8). Tergum IX consists of 2 narrow lateral heavily pigmented plates,

mesal area membranous, each lateral plate with a small narrow lobe on

caudomesal margin bearing 5-9 (usually 7-8) short setae, covered with min-

ute spicules; gonocoxite short, broad, heavily pigmented, covered with min-

ute spicules, dorsal surface with several long and a few short setae along

outer area, lateral surface with a number of long setae and several broad

scales, ventral surface with several long setae on apical area, several moder-

ately long and short setae scattered over basal 0.65 and a few broad scales on

outer area, mesal surface membranous; gonostylus moderately long, approxi-

mately 0.68 length of gonocoxite, heavily pigmented, bifurcated, outer arm

long, narrow and curved, inner arm short, broader and with 4-8 (usually 4-5)

short setae, gonostylar claw absent; basal mesal lobe consists of a broad

apically raised plate situated on basal area of mesal surface of gonocoxite,

narrowly attached to basomesal area of dorsal surface and broadly attached

to basomesal area of ventral surface of gonocoxite, dorsal surface of plate

covered with 38-54 moderately long curved setae, those along inner and apical

margins somewhat stouter, apical setae also on short tuberculi, ventral sur-

face covered with short spicules, basal area connected with its mate by a nar-

row moderately pigmented band which is covered with small Spicules; proctiger

moderately long, paraproct consists of broad heavily pigmented basal area

and a narrow very heavily pigmented claw-like apical area which is strongly

curved tergally, cercus membranous with a pair of small moderately pigmented

rectangular plates on the dorsal surface, each plate attached to distal portion

of broad basal area of paraproct, cercal setae absent; tergum X narrow,

heavily pigmented, curved, base attached to base of paraproct and apex

attached to caudoventral margin of tergum IX at base of lobe; phallosome

with aedeagus long, narrow, heavily pigmented, apex sagittate, base narrow-

ly divided into 2 thin plates each with a few dorsobasal teeth, paramere moder-

ately long, 0. 74-0. 82 length of aedeagus, heavily pigmented, apex attached to

base of aedeagus and attached near base to parameral apodeme, parameral

apodeme very long, moderately broad, heavily pigmented, attached near middle

to apodeme of gonocoxite; sternum IX medium size, lateral, basal and median

areas heavily pigmented, 4-7 short to moderately long setae on median caudal

area.

PUPA (Fig. 9). Chaetotaxy as figured and recorded (Table 1). Moderately

pigmented, some specimens with heavily pigmented areas of cephalothorax,

metanotal plate and abdominal segments I-V; ocular plate without cuticular

facets. Respiratory trumpet. Heavily pigmented, index 2.72-3.41, mean 3.09.

Cephalothorax. Setae 1-3-CT approximately equally developed. Metanotal

plate. Seta 10-CT caudad and laterad of 11-CT. Abdomen. Setae 2, 3-I

widely separated; 9-VII stout, barbed, triple to 5 branched, positioned directly

cephalad of 6-VII; 9-VIII well developed, barbed, 6-10 branched. Paddle.

Ovoid; minute serrations on distal portion of basal 0. 51-0.62 of outer margin;

Reinert: Subgenera Indusius and Edwardsaedes 17

minute spicules on apical 0.38-0.49 of outer and apical 0. 04-0.10 of inner

margins; midrib does not reach apex; seta 1-P short, single or double; index

1.47-1.56, mean 1. 52.

LARVA (Fig. 10). Chaetotaxy as figured and recorded (Table 2). Head.

Covered with granules dorsally and ventrally; seta 4-C small, 6-10 branched;

o-C long, triple to 6 branched; 6-C long, double to 5 branched; 7-C long, 6-14

branched; 12-C small, triple to 6 branched; 13-C moderately long, double or

triple; dorsomentum with 41-43 (usually 41) teeth, heavily pigmented; ventro-

median cervical sclerite heavily pigmented. Antenna. Long, slender, moder-

ately pigmented, numerous well developed spicules; seta 1-A long, barbed,

0-12 branched, base inserted 0.36-0.43 from base of antenna; 2-A long, with

a subapical constriction; 3-A short, 0.28-0.40 length of 2-A; 4-A moderately

long, 0. 43-0. 55 length of 2-A; 5-A flat, apical portion membranous, basal

portion pigmented with a short dorsal spine; 6-A short, 0.28-0.33 length of

2-A. Abdomen. Setae 6-I-VI and 7-I-II located on small heavily pigmented

plates; 8-IV-VII multiple branched; 1-VII short, 5-10 branched; 1, 2-VII on

common small pigmented plate; comb consists of 8-15 (usually 12) scales

arranged in 2 irregular rows, each scale with a long stout median apical spine

and short lateral denticles on laterobasal areas; saddle heavily pigmented,

completely rings segment X, several stout spicules on caudal area, acus ab-

sent; 4 long tapered anal papillae; 1-X short, single; 2-X short, 7-8 branched;

3-X slightly longer than 2-X, double to 5 branched; ventral brush consists of

16-17 setae, 9-10 long multiple branched (5-10 branches) setae on grid, 5-7

shorter multiple branched (4-9 branches) precratal setae which are inserted

in ventral margin of saddle. Szphon. Moderately long, moderately pigmented,

acus well developed, index 3.14-4.13, mean 3.58; pecten on basal 0. 58-0. 63

of siphon, composed of 14-22 (usually 18) teeth, apical 2-4 teeth wider spaced

than remainder of teeth and usually smooth and curved, each tooth long,

slender, pointed and with a long stout ventral denticle, several teeth also

usually with an additional minute denticle; seta 1-S short, double to 7 (usually

3-4) branched, base attached on basal 0. 64-0. 71 of siphon distad of last pecten

tooth.

EGG (Fig. 11). The following account of the egg is taken from Moriya

et al. (1973). The egg chorion has a surface sculpturing consisting of a net-

like pattern of cells when viewed under lower magnification. Under higher

magnification each cell of the pattern had a length of about 20 microns and a

width of about 10 microns. The inner part of each cell possessed a hill-like

process with an uneven surface consisting of small depressions and elevations.

The hill-like process was surrounded by cloud-like structures. Eggs of im-

primens are broadly fusiform in shape, black, with a length of 0.6-0.8 mm

(mean 0.65 mm) and a breadth of 0.18-0.25 mm (mean 0.22 mm). Maximum

width of egg is just before middle. The authors give dorsal and lateral out-

lines (page 49) and a scanning electron microphotograph of the surface sculp-

turing of the egg (page 51).

DISCUSSION. Females of imprimens are similar in habitus to Aedes vex-

ans Meigen and Aedes caecus (Theobald), both in subgenus Aedimorphus, as

demonstrated by a number of specimens found in the U. S. National Museum

(Natural History) (USNM) and the BMNH collections labeled as vexans and

caecus but were actually imprimens and vice versa. Aedes imprimens fe-

males can be distinguished from these species by the development of the max-

illary palpus, presence of hairs mixed with scales on subspiracular area, and

type and color of other pleural scales. Males of imprimens are easily recog-

nized by the development of the maxillary palpus; however, they resemble

18 Contrib. Amer. Ent. Inst., vol. 13, no. 1, [976

those of Aedes lineatopennis. The scutal scaling of the latter species differs

markedly from imprimens. Specimens from Java (Gombong, Padaherang and

Tilatjap) previously reported in the literature as imprimens are vexans vexans.

A female in the BMNH identified as imprimens by Barraud (1928a: 664) from

Rangamati, Chittagong Hill Tracts, is Aedes caecus.

Leicester's (1908: 153) description of auratus fits imprimens very well

and I think there is little doubt the 2 are conspecific. JI also concur with the

synonymy of brugi with imprimens. Theobald (1910a: 21) originally described

suknaensis which was later synonymized by Edwards (1922b: 467) with zmpri-

mens; however, Barraud (1928a: 663) questioned this synonymy and thought

suknaensis was possibly synonymous with caecus. Later Edwards (1932: 170)

also questionably listed suknaensis with caecus and Barraud (1934: 257) un-

equivocably synonymized suknaensis with the latter species. This arrange-

ment was followed by Stone et al. (1959: 191) but Reinert (1973: 28), while

working on the subgenus Aedimorphus, transferred suknaensis to the subgenus

Edwardsaedes and questionably included it with imprimens. After examining

the types and numerous specimens from a wide geographical range I believe

Ssuknaensis and imprimens are conspecific.

The correct date of publication for zmprimens is 1860 and not 1861 as

reported by previous workers.

TYPE-DATA. The holotype female of zmprimens in the BMNH bears the

following information on the labels: imprimens; type; Amb. | Amboyna];

Saunders, 68-4; Culex imprimens Walk. , Identified as the type by E. A.

Waterhouse. Walker (1860: 144) identified the location as Amboyna and the

collector as A. R. Wallace. The holotype male of bDvugi in the BMNH possesses

the following information on the labels: Aedes (Banksinella) brugi Edw.; S.

New Guinea: Merauke, 1922, Dr. S. L. Brug [collector], B. M. 1923-433;

holotype. The holotype female of suknaensis is in the Indian Museum, Calcutta,

India and bears the following information on the labels: Sukna, 500 feet, E.

Himalayas, 2-VII-08, In the dense jungle; cotype F. V. T. A female paratype

in the BMNH contains the same information as the holotype except for the ad-

ditional following data: flies and bites by day, N. A., 1909-272, and T75.7

(genitalia preparation number). Theobald (1910a: 21) records the collection

district as Darjiling and the collector as Annandale for the types of suknaensts.

The female syntype specimens of Leicester's (1908: 153) Culex Auratus are

apparently lost. They could not be found in the BMNH.

DISTRIBUTION. 715 specimens examined: 3832, 91°, 2 p, 41, 142 L, 57

with associated skins (22 p, 35 pl, 11 incomplete).

CAMBODIA. Kompong Sela, Stung Chral; 62, 40, 2L.

INDONESIA. Kalimantan, Mahakkam; 12. Moluccas, Amboina Is.; 1%

(holotype of imprimens), Ceram, Toem,Warasiwa; 72, 1°. Sumatra, Dermajoe,

Moeara Tebo; 42. West Irian, Hollandia, Maffin Bay, Nabire, S. Geelvink Bay,

Sansifor, Vogelkop, Kebar Valley west of Manokwari; 102, 1“ 1, 17, 34 L,

Merauke; 1° (holotype of brugi).

INDIA. Assam or Bengal; 1°. Bengal, Darjeeling Dist., Sukna; 19 (para-

type of suknaensis). Andhra, Musalimadugu Forest Reserve; 3c.

JAPAN. Akigase, Nagano Pref., Norikura-Kagen, Honshu, Omiya; 89.

MALAYSIA. Sabah, Keningau; 12. Sarawak, Kuching; 12. Selangor, Ulu

Gombak, Ulu Langat; 72, 1 lp, 21.

PAPUA-NEW GUINEA. Finschhafen, Huon Peninsula, Gusika, Bougainville

Is., Rauna Plantation, Lae, Milne Bay, Murua near Kerema, S. Vanapa River

3 km, Morobe Dist., Wau, Mt. Missim, Sepik Dist., Wewak, West Highlands,

Ming, Gum Hamlet, Goodenough Is., Watutu Pt., Kiriwina Is. ; 22 pl, 719,

Reinert: Subgenera Indusius and Edwardsaedes 19

30% p, 40c, 38 L.

PHILIPPINE ISLANDS. Mindanao Is., Ft. Pikit; 12. Mindoro Is., San

Jose; 22, 2°, 1 p, 21. Palawan Is., Puerto Princesa; 42. Tawi Tawi Group,

Sulu, Sanga Sanga Is., Lapit-Lapit; 19.

SOLOMON ISLANDS PROTECTORATE. Guadalcanal Is., Belasuma R.,

Flu, Gold Ridge, Honiara, Nini Creek (35 km SE. of Honiara), Roroni (35 km

E. of Honiara), 25 km SSE. of Honiara, Koli Ar., Marovovo, Ruavatu Est.,

Teneru Ar. ; 1632, 50°. New Georgia Group, Rendova Is., 4%. Russell Is. ,

Pavuvu Is.; 12. San Cristobal Is., Arohane; 49.

THAILAND. Chiang Mai, Amphoe Muang, Huey Chang Kien; 12. Chon

Buri, Amphoe Bang Lamung, Khao Mai Keo; 12. Kanchanaburi, Ban La Wa,

Ban Nong Phang Kung, Ban Sai Yok, Huai Mae Nam Noi; 262 pl, 102 p, 798,

4c p, 10°, 56 L. Khon Kaen, Amphoe Chun Phae, Pha Dong Larn, Tham Pho

Ti Yan; 82. Lampang, Huai Mae Phrao; 42 pl. Nakhon Ratchasima, Amphoe

Pak Chong, along Lam Nam Tha Khlong; 192, 20°. Nakhon Si Tammarat,

Amphoe Chawang, Banna Nabon, Ban Tha Phae, Khao Luang; 22. Nan, Ban Ta

Loc; 2L. Songkha, Amphoe Haad Yai, near Ton Nga Chang Waterfalls; 2¢ pl,

42 p, 1o% pl, 9 L. Tak, Khao Salak Phra; 12 p. Ubon Ratchathani, Chongmek;

12. Udon Thani, Amphoe Muang, Ban Kau Noi; 12.

VIETNAM. Binh Dinh, An Khe, Qui Nhan; 192, 1 L.

Distribution from literature.

AUSTRALIA. North of Coen (Marks 1972: 106).

INDIA. E. Himalayas, Darjiling Dist., Sukna (Theobald 1910a: 21); Assam,

Dibrugarh, Dimapur, Golaghat; Bengal (Barraud 1928a: 664).

INDONESIA. Amboina (Theobald 1901: 422, 1910b: 350), Merauke (Edwards

1924: 377); Borneo (Edwards 1922b: 467); Java, Weltevreden; New Guinea (Brug

and Haga 1923: 639); Sumatra, Djambi, Kotta Tjane, Moeara Tebo (Brug and

Edwards 1931: 258); Ceram (Bonne-Wepster and Brug 1937: 38, Bonne-Wepster

1954: 239); New Guinea, Cyclops Mts., Hollandia (Knight and Hull 1953: 468);

Hollandia area (Assem 1961: 24); Kalimantan, Mahakkam; Java, Gombong,

Modjavarno, Padaherang, Tasikmalaja, Tilaljap; Sumatra, Atjeh, Dermajoe

(Mattingly 1961: 54); Moluccas; New Guinea, Kaimana, Manokwari (Assem and

Bonne-Wepster 1964: 99).

JAPAN. Hokkaido; Kyushu (Yamada 1927: 571, Hsiao and Bohart 1946: 22);

Honshu, Sendai area (Waku 1950: 72); Kyushu (LaCasse and Yamaguti 1950:

104); Hokkaido,. Urawa (Hara 1957a: 65); Kanayawa, Ohmura (Kurihara 1963:

196); Saitawa-ken, Kitaurakazu (Moriya et al. 1973: 54).

KOREA. Kwang-Ju Airbase, Osan Airbase (Reisen et al. 1971: Table 26).

MALAYSIA. (Leicester 1908: 153); Selangor, Ulu Gombak (Macdonald 1957:

21), Sarawak, Kuching (Mattingly 1961: 54).

PAPUA-NEW GUINEA. Solomon Is., New Britain (Iyengar 1955: 30),

Bougainville (Belkin 1962: 411, Peters and Christian 1963: 54); New Guinea,

north, east and south coasts, Sepik area (Peters and Christian 1963: 54);

Bismarck Arch. (Steffan 1966: 213). |

PHILIPPINE ISLANDS. Mindanao, Port Banga (Edwards 1929: 5); Min-

danao, Ft. Pikit; Mindoro, San Jose; Palawan, Puerto Princesa (Knight and

Hull 1953: 468).

SOLOMON ISLANDS PROTECTORATE. Solomon Is. (Edwards 1925: 257,

Knight et al. 1944: 52), Guadalcanal (Edwards 1924: 377); Kookoom, Ilu (Paine

in Paine and Edwards 1929: 306); Russell Group, Pavuvu (Knight and Hull 1953:

468); Guadalcanal, New Georgia (Belkin 1962: 411).

THAILAND. Chieng Moeang (Causey 1937: 413).

VIETNAM. Cam Rahn Bay (Parrish 1968: 12, 1969: 554); Phu Cat Airbase

20 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

(Reisen et al. 1971: Table 13).

' BIONOMICS. In Thailand immatures were collected from small and large

flood pools containing clear or turbid, fresh, unmoving, temporary water in

unshaded to moderately shaded areas located in bamboo forests (3 times), teak

groves (twice), rainforests (twice), secondary deciduous forest (once) and a

rubber plantation (once), in valley, plain, hilly and mountain terrain and at

elevations of 76-854 m (most often near 120 m). In New Guinea larvae were ©

collected from water ina shellhole, in South Vietnam from water in a foxhole

and in Indonesia from water in a temporary puddle.

Females in Thailand have been taken biting man between 1400-2100 hours in

bamboo groves and forests with light vegetation. In India females were taken

biting man during the day in the deep jungle and in Japan they have been collected

resting in bamboo groves. Females were collected biting man and animals at

1200-1300 hours in Cambodia and biting man in the jungle in the Philippine Is-

lands and Solomon Islands. In New Guinea adults have been taken at elevations

from sea level to 1200 m.

Immatures have been collected from: a buffalo wallow on a high plateau in

Thailand (Causey 1937: 413); in transient ground water pools in the jungle

(Assem 1961: 24) and temporary pools in half shade at the border of a rain-

forest in Indonesia (Bonne-Wepster 1964: 99); in temporary pools and in flowing

water in flooded areas located in partial to heavily shaded areas of Papua

(Steffan 1966: 213); in ground pools in a forest in Japan (Waku 1950: 72); and

in the Solomon Islands in partly shaded leafy woodland pools (Bohart 1945: 63)

and in temporary pools of a very transient nature usually located in the jungle

(Belkin 1945: 6). Eggs are laid singly, are not cemented to objects (Belkin

1962: 410), require drying for 4-6 weeks (Belkin 1945: 6), hatching taking place

when the shallow jungle depressions in which they lie are flooded by rain and

the resulting larvae complete their development within 3 days and are apparently

capable of withstanding some desiccation (Laird 1956: 39).

Adults have been collected: aggressively biting in deep shade during the

day in the Philippine Islands (Bohart 1945: 63); in biting collections (Macdonald

1957: 21, Macdonald and Traub 1960: 100) and in the jungle, common in some

places but very local (Leicester 1908: 154) in Malaysia; attacking humans during

the day in forest or brush areas in the mountains of Japan (Waku 1950: 72);

readily biting man during the daytime (Steffan 1966: 213) and in New Jersey

light traps in the lowlands of New Guinea (Peters in Peters and Christian 1963:

59, 74); in houses, stables and in the woods (Bonne-Wepster 1954: 239) and

biting man in Indonesia (Assem and Bonne-Wepster 1964: 99); in light traps

during June and July in Korea (Reisen et al. 1971: Table 26); in light traps

during May and June in South Vietnam (Parrish 1968: 12, Reisen et al. 1971:

Table 13); biting in numbers during the day in and near jungle areas, the fe-

males being capable of biting through heavy clothing and attack readily in the

shade and at dusk (Belkin 1945: 6), biting man in the foothill areas (Belkin

1962: 411), feeding on man in large numbers by day, readily biting through the

shirt and stockings, occurring occasionally in marginal parts of plantations,

but were absent at a distance of more than 300-400 yards from forest margin in

the Solomon Islands (Paine in Paine and Edwards 1929: 306); and in the dense

jungle biting during the day in India (Theobald 1910a: 22).

Reinert: Subgenera Indusius and Edwardsaedes 21

ACKNOWLEDGMENTS

Grateful appreciation is expressed to: Bruce F. Eldridge and Ronald A.

Ward, Walter Reed Army Institute of Research for reading and commenting

on the manuscript; to the following for the loan of types and other specimens--

Peter F. Mattingly, British Museum (Natural History), London, Douglas J.

Gould, U. S. Army Component, South East Asia Treaty Organization Medical

Research Laboratory, Bangkok, Thailand, Wallace A. Steffan, Bernice P.

Bishop Museum, Hawaii, J. M. Klein and R. M. Quentin, Office de la

Recherche Scientifique et Technique Outre-Mer, Bondy, France, John N.

Belkin, University of California, Los Angeles, and Kazuo Tanaka and Edward

S. Saugstad, U. S. Army Medical Laboratory (Pacific), Japan; to John E.

Scanlon, University of Texas School of Public Health, Houston, for the loan

of his notes on the type of suknaensis in the Indian Museum; to Young T. Sohn

and Ann Hoskins Dery, Medical Entomology Project (MEP), for preparing the

detailed illustrations; to Susan Hevel, MEP, for laboratory assistance; and to

Janet D. Rupp, MEP, for typing the manuscript for offset reproduction.

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1963. Comparative studies on the pleural structure of the Japanese

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Reinert: Subgenera Indusius and Edwardsaedes 295

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26 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

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Reinert: Subgenera Indusius and Edwardsaedes 27

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28 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

LIST OF FIGURES

1. Aedes (ndusius) pulverulentus--adult female morphology

2. Aedes (Indusius) pulverulentus--female genitalia

3. Aedes (Indusius) pulverulentus--adult male morphology and female posttarsi

4. Aedes (Indusius) pulverulentus--male genitalia

0. Distribution of Aedes (Edwardsaedes) imprimens

6. Aedes (Edwardsaedes) imprimens--adult morphology

7, Aedes (Edwardsaedes) imprimens--female genitalia

8. Aedes (Edwardsaedes) imprimens--male genitalia

9. Aedes (Edwardsaedes) imprimens--pupa

10. Aedes (Edwardsaedes) imprimens--larva

11. Aedes (Edwardsaedes) imprimens--female and male tarsomeres 5 and

posttarsi; male segment VIII; outline of

egg (after Moriya et al. 1973)

LIST OF FIGURE ABBREVIATIONS

Female Genitalia

AGDuB = Accessory gland duct base LVL = Lower vaginal lip

BMA = Basal median apodeme PGL = Postgenital lobe

Ce = Cercus SCa = Seminal capsule

DPGL = Line of attachment of peri- SCaP = Seminal capsule pore

anal membrane to dorsal Tu = Tuberculus

surface of PGL UVL = Upper vaginal lip

H = Hinge UVS = Upper vaginal sclerite

I = Insula VIII-S = Sternum 8

IX-Te = Tergum 9 VIlIl-Te = Tergum 8

Male Genitalia

Ae = Aedeagus IX-Te = Tergum 9

AG = Apodeme of gonocoxite PaA = Parameral apodeme

BML = Basal mesal lobe Par = Paramere

Ce = Cercus Ppr = Paraproct

Gc = Gonocoxite Pr = Proctiger

Gs = Gonostylus VIlI-S = Sternum 8

IX-S = Sternum 9 VIlIl-Te = Tergum 8

X-Te = Tergum 10

Pupa

CL = Cephalothorax MP = Metanotal plate

I-VIII = Abdominal segments 1-8 P = Paddle

7 = Respiratory trumpet

Larva

A = Antenna M = Mesothorax

C = Head MP = Mouthpart

CS = Comb scale Pp = Prothorax

Dm = Dorsomentum PT = Pecten tooth

I- VIII, X= Abdominal segments 1-8, S = Siphon

= Metathorax

10 ei

Aedes (/Indusius) pul/veru/entus °

ventral SCaP

Vill-Te

Aedes (Indusius) pulverulentus

Fig.3

sao g-

Dine hee ‘nem atone "Maye

ove ga oe

soe De 2S Hoc hm ER Se oe

Aedes (/ndusius) pulverulentus

X-Te and Pr

lateral

gals nena 9)

a it

X-Te and Pr

dorsal

0.3

PaA

ventrolateral dorsomesal

Aedes (/ndusius) pul/lveru/entus

Son ales — SU9W/ ICU!

Aedes (Edwardsaedes) imprimens

i |

ventral

Aedes (Edwardsaedes) imprimens

Fig. 8

X-Te and Pr

imprimens

Aedes (Edwardsaedes)

Aedes (Edwardsaedes) /mprimens

Fig. 10

a cervical sclerite

EEE

Aedes (Edwardsaedes) imprimens

Egg (outline, dorsal)

ye,

Id y

NAF,

\ va |

Wh L Vins

A We) }

any

\ i

Uf!)

Age

vl 1 As,

ei Van

rot E

ers ee sa at

RE ony a =

rat $ ng

se IS oe

A ee

ST Se

a -

Vill-Te o&

VIII-S o&%

Tarsomeres 5 and Posttarsi

Aedes (Edwardsaedes) (/mprimens

40 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

APPENDIX: TABLE 1. Record of the branching of the setae on the pupae

of Aedes (Edwardsaedes) imprimens (10 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

Cephalothorax Abdomen I (Cont. ) Abdomen IV

1 2 2 11 1 i 0 1 1

2 2-3 3 Abdomen II 1 3-6 4)

3 2 2 0 1-2 1 2 1 1

4 2-5 S 1 6-13 12 3 6-8 6

5 4-8 & 2 1 1 4 3-8 4

6 2-6 3 3 1-2 1 5 3-5 4

i 3-6 4 q o-11 i‘ 6 2-4 A

8 0-13 7 5) 8-12 8 7 : 2-5 4

9 2-3 2 Abdomen III 8 3-9 4

Metanotal Plate 0 1 1 2 1 1

10 3-6 5 1 5-12 7 10 2-4 3

11 1 1 2 1-2 1 11 1-3 2

12 4-8 5 3 1 1 14 1 1

Abdomen I 4 3-8 ° Abdomen V

1 ZERO 20 5) 9-16 9 0 1 1

2 1-2 1 6 2-4 3 1 3-7 5

3 2-6 3 7 3-7 5) 2 1-2 1

4 6-16 11 8 4-6 5) 3 2-3 3

5 3-6 4 9 1 1 4 4-9 i

6 1-2 1 10 2-4 3 *) 3-4 3

7 2-4 3 11 2-4 2 6 2-3 2

9 1-3 1 14 1 1 7 2-8 6

10 1 1 8 3-9 o

Reinert: Subgenera Indusius and Edwardsaedes

TABLE 1 (Cont. ).

Seta Range Mode Seta Range Mode Seta Range Mode

Abdomen V (Cont. ) Abdomen VI (Cont. ) Abdomen VII (Cont. )

3 1 1 8 3-5 O 7 1-2 1

10 1-3 2 9 1 1 8 4-6 5)

11 2-4 1 10 1-2 ? | Gy BRS 4

14 f 1 11 1-4 1 10 1-2 1

Abdomen VI 14 1 1 11 2-9 os

0 1 1 Abdomen VII 14 1 1

1 4-7 o 0 1 1 Abdomen VIII

2 ‘2 1 1 3-7 5 0 1 1

3 2-3 2 2 1 1 4, 2-4 3

be 4-7 5) 3 3-8 6 9 6-10 8

5) 3-4 3 a 2-3 3 14 1-2 1

6 1-3 2 O 2-5 4 Paddle

7 1-2 | 6 4-8 O 1 1-2 1

42 Contrib. Amer. Ent. Inst. , vol. 13, no. 1, 1976

APPENDIX: TABLE 2. Record of the branching of the setae on the larvae of

Aedes (Edwardsaedes) imprimens (10 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

Antenna Prothorax (Cont. ) Mesothorax (Cont. )

1 o-12 8 3 2-3 3 12 1-2 t

Head 4, 2-3 2 13 7-12 10

0 1 1 5 1-2 1 14 8-11 8

1 1 1 6 1 1 Metathorax

3 1 1 7 2-4 2 1 2-4 3

& 6-10 9 8 2-4 3 2 2-5 5)

5 3-6 3 9 1-3 2 3 8-19 8

6 2-5 3 10 1-2 1 4 0-9 7

7 6-14 8 11 3-5 4 5 1 1

8 2-3 2 12 1 1 6 1-2 1

9 2-3 3 14 2-3 3 7 5-9 7

10 2-4 2 Mesothorax 8 7-11 8

11 6-12 7 1 3-5 4 9 3-5. 4

12 3-6 3 2 2-7 3 10 1 1

13 2-3 2 3 1-3 1 11 1 1

14 1-3 2 4. 2-4 3 12 1 1

15 3-6 6 5 1 1 13 9-14 13

18 1 1 6 4-6 5) Abdomen I

6MP 1 1 7 1 1 2 1-2 1

Prothorax 8 9-8 5 3 4-7 4.

0 8-16 12 9 4-8 7 4 13-24 20

1 1-2 1 10 1 1 9) 3-6 3)

2 2-3 2 11 1 1 6 2-4 3

Reinert: Subgenera Indusius and Edwardsaedes

TABLE 2 (Cont, ).

Seta Range

Mode

Abdomen I (Cont. )

2-3

2-4

1-2

Abdomen II

Seta Range

Abdomen III (Cont. )

Abdomen IV

6-9

1-2

4-5

o-7

1-3

Mode

Seta Range

2-3

3-4

o-9

Mode

Abdomen IV (Cont. )

44 Contrib. Amer. Ent. Inst., vol. 13, no. 1, 1976

TABLE 2 (Cont. ).

"Seta Range Mode Seta Range Mode Seta Range Mode

Abdomen VI (Cont. ) Abdomen VII (Cont. ) Abdomen VIII (Cont. )

2 1 1 2 1-2 1 2 2 2

3 2-9 | 2 3 0-10 7 3 0-10 7

4, 0-10 6 4 2-9 8 4 1-2 1

5) 2-3 2 5) 9-8 5) 3) 3-6 5)

6 2-3 2 6 B18. 10 14 1 1

1 3-6 5) 7 2-3 2 Abdomen X

8 4-7 7 8 0-12 5) 1 1 1

9 1-2 1 9 4-5 O 2 7-8 8

10 1-2 - 10 2-3 3 3 2-5 3

11 3-95 4 11 4-6 4 Siphon

12 2-3 3 12 2-4 3 1 2-7 fy

13 EO-1o) « 12 13 6-12 6 2 1 1

14 1 1 14 1 1 7 i) 1

Abdomen VII Abdomen VIII 8 0-8 D

0 ul 1 0 + 1 ) 1 1

Reinert: Subgenera Indusius and Edwardsaedes 45

INDEX

Valid names are set in romantype. The italicized pages are those

which begin the primary treatment of the taxon. Numbers in parentheses

refer to the figures illustrating the species in question.

Aedes (genus) 1, 2, 3, 4, 8, 9, 10, Halaedes 4

14.12, 1S, b, teeee

Aedes (subgenus) 11 imprimens 1, 2, 9, 10, 11, 12,

Aedimorphus 1, 8, 9, 11, 12, 18 19. 17 ,) 18° 28, 40 42°15, 6,

auratus 12, 18 fairs eve: ee 8 Paes Le 4

australis 4 Indusius 1, 2, 3, 4, 11, 28

bancrofti 12 | lineatopennis 11, 18

Banksinella 1, 9, 13, 18 :

brugi 12, 13, 18 Neomelaniconion 1, 9, 11, 13

Nothoskusea 4

caecus 12, 17, 18

chathamicus 4 Ochlerotatus 1, 4, 9, 11, 12

Christophersiomyia 8 Opifex 4

Cuties. 1,798,432, 136

Culicada 9, 12 Paraedes 4

Culiseta 4 Psorophora 11

pulverulentus 1, 2, 4, 8, 28 (1, 2,

Deinocerites 4 3, 4)

Feculex 1,9; 12 suknaensis 1, 12, 18, 21

Edwardsaedes 1, 2, 9, 10, 11, 12,

: ie las Ory $ =| Verrallina 4, 11

vexans 17

Finlaya 8 vexans vexans 18

fuscus 4

Wuchereria 12

gouldi 8

Contributions

of the

American Entomological Institute

Volume 13, Number 2, 1976

MEDICAL ENTOMOLOGY STUDIES - V.

THE SUBGENUS RHINOSKUSEA OF THE

GENUS AEDES (DIPTERA: CULICIDAE).

John F., Reinert

CONTENTS

PBST ts ee ee es

INTRODUCTION 5 er ae

GENUS AEDES, SUBGENUS RHINOSKUSEA . 2.6 see nt eee ees

KEYS TO SPECIES OF AEDES (RHINOSKUSEA) 2... 1 ee ee eae ss

SPRPCite TART NIe Ne ee ee ee G8

baer Osiy7s (Geeeteh) 8 ok a a es

PUIG?) WIG a a ae ee ee

WavO) MEW GOCCIES: 266 6. i a ee we ee

ACKNOWLEDGMENTS. 20. ee ae es

MITER AT URE CUED 2 ee 8 eh ee ee ee

Dit OF PICU RES se pe ee a a ee Ge ee

LIS] OF FIGURE ABBREVIATIONS. 2.6 6 oka ke 8 oe eek ee

BO es ee ew es we ee ee a ea ee

APPENDICES 2 fue te ee re ee ee eg es

TABLES 1 - 3. Records of the branching of the setae on the

pupae of Aedes (Rhinoskusea) species. ....

TABU 1. VOUMSIVOGIVIS 6 eid oe aa Oe es

TABLE 2... pillait. <i.0. «4 RES EE OE es

POMBE Oe WAVE eee er ee aria Sa as BNE oe oe

TABLES 4 - 6. Records of the branching of the setae on the

larvae of Aedes (Rhinoskusea) species ....

TABLE Ce lQyeIi6 le rac. Sane eee ee es

Ay SOG ah a et cee eae one wea ee ew

Oe eg PU Od esa oe tag Be aah ra aes a gh 9 OR Be ta ae

OS FON Oath. OF OIE OO) OG IOh ROR On, On Oy ek Ore OO) Oh AO Ol Oh Oe Oe Oo Or Ole Ole OR OE lOO HO OO) On hes) ®.

MEDICAL ENTOMOLOGY STUDIES - V.

THE SUBGENUS RHINOSKUSEA OF THE GENUS

AEDES (DIPTERA: CULICIDAE)!.

John F. Reinert2

ABSTRACT

The subgenus Rhinoskusea Edwards is redescribed and compared to other

subgenera of Aedes Meigen. Three species, longirostris (Leicester), pillaii

Mattingly and wardi new species, are included in the subgenus and are described

and illustrated. Tables listing the range and mode of the setal branching of the

pupae and larvae of the 3 species are given.

INTRODUCTION

The subgenus Rhinoskusea of the genus Aedes Meigen was originally

described by Edwards (1929: 342) for the species longirostris (Leicester). The

type species of the subgenus had been included in various generic combinations

prior to its assignment to Rhinoskusea and are as follows: Ficalbia Theobald

by Leicester (1908: 228), Uvanotaenia Lynch Arribalzaga by Taylor (1918: 841,

as hilli) and Skusea Theobald by Edwards (1917: 224).

This primarily Oriental subgenus consists of 3 morphologically very similar

species (longirostris, pillaii Mattingly and wardi new species). The present

study redefines the subgenus and gives characters which separate all stages of

Rhinoskusea from the other subgenera of Aedes. Morphologically similar

features of the subgenus and other subgenera are discussed. All stages of the

3 included species are quite similar except the male which possesses very

distinct characters of the genitalia. Descriptions and illustrations are pre-

sented for the female and male genitalia and immatures of longirostris, pillati

and wardi. The female and male habitus of longirostris are also given.

Tables 1-6 list the range and mode of the setal branching of the pupae and

larvae of the 3 species. :

IThis work was supported in part by Research Contract DAMD-17-74-C-4086

from the U. S. Army Medical Research and Development Command, Office of

the Surgeon General and carried out at the Medical Entomology Project, Smith-

sonian Institution, Washington, D. C. 20560.

2Major, Medical Service Corps, U. S. Army, Department of Entomology,

Walter Reed Army Institute of Research, Washington, D. C. 20012.

2 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

Abbreviations used in references to literature conform to the BIOSIS List

of Serials, Biosciences Information Service of Biological Abstracts, Philadel-

phia, 1972. In synonymy sections, an asterisk following the abbreviations

used (A = adult, 2 = female, “ = male, P = pupa, L =larva) indicates that at

least some portion of that sex or stage is figured. In the distribution sections,

abbreviations used are the same as in the synonymy but with the following 2

additions, p = pupal skin and 1 = larval skin. Seven specimens (6 larvae of

pillaii) were used to determine the range and mode of the setal branching in

pupal and larval descriptions and tables. In the pupal descriptions and tables

the number of branches on abdominal seta l-I was measured on the basal third

of the seta. Measurements on the illustrations are in millimeters. Distribution

records are indicated as follows: countries are in capital letters, provinces and

primary administrative divisions are in italics, and place names have the first

letter capitalized. The number of specimens examined from each province

follows the last place name of the province in the distribution section. The

spelling of provincial and locality names was taken from the following Official

Standard Names Gazetteers prepared by the Geographic Names Division, U. S.

Army Topographic Command and the Office of Geography, U. S. Department of

the Interior: Malayasia (1970); Philippine Islands (vols. I and II, 1953); South

Vietnam (1971); and Thailand (1966). Locality names which did not appear in the

gazetteers were spelled according to the collection data sheets and labels on the

specimens.

Information in the biology and distribution sections was taken from the col-

lection data sheets and specimen labels of the specimens that I examined.

References in the published literature are also given.

Nomenclature and chaetotaxy used for the female, male, male genitalia,

pupa and larva follow Knight (1970, 1971), Knight and Laffoon (1970a, 1970b,

1971a, 1971b) and Laffoon and Knight (1973). The terminology of the female

genitalia follows Reinert (1974).

GENUS AEDES MEIGEN

SUBGENUS RHINOSKUSEA EDWARDS

Type species: Ficalbia longirostris Leicester

Ficalbia of Leicester 1908: 228.

Aedes (Rhinoskusea) Edwards 1929: 342.

Aedes (Rhinoskusea) of Edwards 1932: 160; Barraud 1934: 216; Knight and

Hull 1952: 163; Mattingly 1958: 35; Stone et al. 1959: 178; Mattingly

1961: 10; Huang 1968: 187; Stone and Delfinado 1973: 307.

FEMALE. Small dark mosquitoes with few pleural setae and scales. Head.

Antenna dark, moderately long, 6 setae in flagellar whorls, pedicel with a few

scattered short fine hairs and occasionally a few small dark scales mesally;

clypeus dark, bare; maxillary palpus dark scaled, 4-segmented, short, 0.13-

0.14 length of proboscis; proboscis dark scaled, narrow, long, noticeably

longer than femur I; eyes contiguous; interocular setae absent; ocular setae

numerous; vertex and lateral surface of head with only broad decumbent scales;

occiput with a few erect forked scales. Thorax. Scutum covered with narrow

curved dark scales except for a very small patch of broad dark scales on median

anterior promontory area; prescutellar space with a narrow median longitudinal

Reinert: Aedes (Rhinoskusea) 3

stripe of narrow curved scales; scutellum with a patch of broad dark scales

on each lobe; scutal setae dark and present on following areas: median anteri-

or promontory, dorsocentral (numerous anterior and posterior), acrostichal

(numerous anterior and posterior), scutal fossal (anterior, lateral and posteri-

or), supra-alar (numerous), posterior medial scutal, postalar callar and scu-

tellar (lateral and median lobes); antepronota widely separated, not enlarged,

without scales, 7-14 setae; postpronotum without scales, 2-3 posterior setae;

propleuron without scales, 2-6 setae; prosternum, subspiracular area, meso-

meron and metameron bare; postspiracular area without scales, 1-4 setae;

paratergite narrow, bare; mesepisternum with a small patch of pale scales on

lower posterior area, 1-2 upper and 3-7 long and 3-7 short posterior setae;

prealar knob without scales, 2-5 setae; mesepimeron without scales and lower

setae, 4-10 short setae on posterior dorsal area. Legs. Coxae I-III each

with a patch of scales and setae; femora I-III each long and relatively narrow,

anterior surface dark scaled, III also with a narrow anteroventral pale scaled

stripe on basal area; tibiae long, dark scaled; tarsi I-III each dark scaled, III

with tarsomere 1 long, slightly longer than tibia III; posttarsi I-III each with 2

ungues, each pair of ungues equal in size and simple. Wing. Dorsal and ven-

tral veins dark scaled; alula with a single row of several narrow dark scales

on margin; upper calypter with a number of long dark setae on margin; remigium

usually with 1 short seta (often hidden beneath scales). Halter. Pedicel pale;

capitellum dark scaled. Abdomen. Terga dark scaled, usually with a small

laterobasal pale scaled patch on I-VI; sterna pale scaled; terga and sterna with

numerous short dark setae, mostly along posterior margins. Genitalia. Ter-

cum VIII lightly pigmented, base and apex nearly straight, a number of short

setae scattered over apical 0.45-0.93, those along apex short, basolateral

seta absent, covered with minute spicules, scales usually absent (only 1-2

scales when present), basal 0.65-0.90 retracted into segment VII, VIII-Te

index 0. 68-0. 83, VIII-Te/IX-Te index 1.44-1.91, length 0.16-0.19 mm,

width 0.20-0.26 mm; sternum VIII lightly to moderately pigmented, base

slightly concave mesally, apex with a shallow to moderately deep median inden-

tation (0.02-0.16 deep), numerous short setae scattered over most of area,

setae 1-3-S widely separated, located in a more or less diagonal line, 1-S

basomesad, 3-S apicolaterad, covered with minute spicules, scales usually

absent (only 1-5 scales when present), apical intersegmental fold moderately

pigmented, VIII-S index 0. 72-0.87, length 0.20-0.23 mm, width 0. 25-0. 31

mm; tergum IX moderately pigmented, base without median indentation, apex

with a small median indentation and with 2-6 setae on each side of midline,

0-11 total setae, covered with minute spicules, a short flap on lateral area,

IX-Te index 0. 72-0.87, length 0.09-0.11 mm, width 0.12-0.13 mm; insula

lightly pigmented, short, wide, lip-like, 3-5 long setae, covered with short

hair-like spicules; lower vaginal lip lightly to moderately pigmented, narrow,

covered with short spicules, lower vaginal sclerite absent; upper vaginal lip

moderately to heavily pigmented, moderately broad to broad laterally, covered

with short spicules, upper vaginal sclerite absent; postgenital lobe short, wide,

apex broad with a small median indentation or straight (0.00-0.17 deep), 3-7

setae on each side of midline, 7-13 total setae, covered with short hair-like

spicules, dorsal PGL index 0.38-0.59, ventral PGL index 1.00-1.35, ventral

length 0.06-0.08 mm; perianal membrane with scattered short spicules; cer-

cus moderately long, apex sharply rounded with 2-4 long stout setae, completely

covered with short spicules, dorsal surface with a number of short to moder-

ately long setae on apical 0.71-0.85, scales absent, ventral surface without

setae, cercus index 2.07-3.20, cercus/dorsal PGL index 1.86-2.79, cercus

4 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

length 0.14-0.18 mm; one large seminal capsule, heavily pigmented, spherical,

several small seminal capsule pores near orifice, base of accessory gland duct

moderately pigmented.

MALE. Essentially as in the female. Head. Antenna moderately long,

lightly plumose, 5-6 setae in flagellar whorls, setae not directed mainly dorsal-

ventrally, apical 2 flagellomeres long, together longer than combined length of

basal 11 flagellomeres; maxillary palpus very short, 0.09-0.13 length of pro-

boscis, 4-segmented, similar to female; proboscis long, narrow, noticeably

longer than femur I. Thorax. Setal groups usually with fewer setae than in

female. Legs. Femora I-III each long and relatively narrow; tarsus III with

tarsomere I long, longer than tibia III; posttarsi I-III each with 2 ungues, I and

II each with one large and one small unguis, larger one simple or toothed, III

with ungues equal in size, both simple. Abdomen. Terga with laterobasal pale

scaled areas reduced in size; lateral setae of terga short. Genitalia. Tergum

IX narrow mesally, broader laterally, setae present or absent; gonocoxite long,

narrow, curved mesally, dorsal surface with several short to moderately long

setae on mesal area and several long stout setae on outer margin, ventral sur-

face with short to moderately long setae on mesal area, long ones on apical por-

tion of outer area, mesal surface membranous; gonostylus moderately long to

long, narrow to broad, with spicules and several short setae, gonostylar claw

short, flared apically, longitudinally striated, heavily pigmented, scoop-like,

attached subapically, gonostylus attached at apex of gonocoxite; basal mesal

lobe large, apical portion free, with setae; proctiger moderately long, para-

proct a narrow heavily pigmented strip from base to apex which terminates in

a small bluntly rounded lobe, cercus membranous with a large lightly to mod-

erately pigmented wrinkled plate covering most of mesal area, cercal setae

present; phallosome simple, aedeagus consists of a short broad trough with

base concave and apex broadly convex, paramere length approximately equal

to length of aedeagus, parameral apodeme short; sternum IX large, extended

cephalad, lateral areas heavily pigmented, with or without setae.

PUPA. Cephalothorax and abdominal segments moderately pigmented,

some segments also with heavily pigmented areas. Cephalothorax. Setae 1,

3-CT approximately equally developed, 2-CT slightly shorter; 4, 5-CT stout,

single to triple, approximately equally developed; 6-CT short, single; 7-CT

long, double to 4-branched; 8-CT double to 5-branched. Metanotal plate. Seta

10-CT double to 5-branched; 11-CT long, stout, double. Respiratory trumpet.

Heavily pigmented; hair-like spicules on inner surface of meatus; basal por-

tion of meatus tracheoid; pinna moderately long; index 4.04-5.93. Abdomen.

Small dark granules on membrane beneath seta 1-I and on median dorsal and

median ventral intersegmental membranes of II-VII; small spicules covering

segments II-VIII and male genital lobe; setae 2, 3-I with bases approximated;

6, 7-I approximately equal in length and development; 3-III removed from

caudal margin; 5-III-VI long, double to 6-branched; 4-VII much longer than

5- VII; 4, 9-VII approximately equally developed; 9-VIII with secondary branch-

ing on apical one half. Paddle. Ovoid with basal area of inner margin de-

pressed; midrib nearly reaches apex; basal area of outer margin with a few

minute serrations; apical portion of outer and inner margins with a few minute

spicules, hair-like fringe absent; seta 1-P moderately long, stout, triple to

5-branched (double on one side of one specimen); index 1. 39-1.90.

LARVA. Head. Moderately pigmented; seta 4-C short, with 4-7 fine

branches, mesad and slightly caudad of 5-C; 5-C long, stout, barbed, with

5-7 branches, slightly mesad and caudad of 6-C; 6-C barbed, double or triple,

longer and stouter than 5-C; 7-C long, stout, barbed, with 9-16 branches,

Reinert: Aedes (Rhinoskusea) 5)

slightly cephalad and widely laterad of 6-C; 11-C moderately long, triple to

11-branched; 12-C small, double or triple; 13-C long, barbed, double to 4-

branched; dorsomentum heavily pigmented, with 19-23 stout teeth; lateral

palatal brush setae filamentous. Antenna. Long, narrow, curved mesad,

moderately pigmented, with numerous spicules; seta 1-A moderately long, with

5-8 branches; 2-A very long; 2, 3-A noticeably subapical in attachment; 3, 6-A

short; 4-A long. Thorax. Setae 1-3-P single, 1-P very long; 5-P very long,

longer than 6, 7-P, single; 9-T barbed, double or triple. Abdomen. Setae

6-III[-VI long, barbed, double; 2-VIII long, single; 1-X long, single, attached

on saddle; 2-X long, with 4-7 branches; 3-X very long, single; ventral brush

of segment X with 10 setae on grid, each with 4-10 branches (caudal 7-8 setae

with 6-10 branches); 4 anal papillae, small, knob-like; saddle small, incom-

pletely rings segment X, with small spicules, acus absent; comb of segment

VIII consists of a large triangular patch of 34-104 scales, each scale with

apical area flared and with a fringe of denticles. Szphon. Moderately pig-

mented; acus well developed, heavily pigmented; index 2.43-3.54; pecten on

basal 0. 44-0. 55 of siphon, composed of 4-15 teeth, distal 1-2 teeth wider

spaced than remainder of teeth; seta 1-S long, barbed, attached far distad

(0. 76-0. 84 from base), distal to last pecten tooth, triple to 6-branched; 2-S

small, single, attached on dorsoapical margin of siphon; 9-S stout, hook-like.

EGG. Not known.

DISCUSSION. The subgenus Rhinoskusea is characterized and separated

from the other subgenera of Aedes by the following: in the adults by the com-

bination of (1) head with vertex with only broad decumbent scales and occiput

with a few erect forked scales, (2) scutum with narrow curved scales except

for a small patch of broad scales on median anterior promontory area, (3)

dorsocentral (anterior and posterior) and acrostichal (anterior and posterior)

setae numerous, (4) scutellum with broad scales on each lobe, (5) thoracic

pleura without scales except for a small lower patch on the mesepisternum

(antepronotum and paratergite also without scales), (6) propleuron with 2-6 setae,

and (7) tarsus III with tarsomere 1 slightly longer than tibia III; in the female

genitalia by the combination of (1) only a single large seminal capsule, (2) in-

sula lip-like with 3-5 long setae, (3) upper vaginal sclerite absent, and (4)

postgenital lobe short and broad; in the male genitalia by the combination of

(1) proctiger with a dorsomedian wrinkled plate and with cercal setae, (2)

aedeagus simple, wide and trough-like, (3) basal mesal lobe with a long apical

portion which is free of gonocoxite and without an apical filament, and (4)

gonostylus with a short, longitudinally striated, scoop-like gonostylar claw; in

the pupae by the combination of (1) small dark granules on membrane beneath

seta 1-I and on intersegmental membranes of II-VIII, (2) seta 11-CT long,

stout and double, (3) seta 5-III-VI long, double to 6-branched, (4) seta 4-VII

much longer than 5-VII, (5) seta 9-VIII multiple branched with secondary

branches on apical one half, and (6) seta 1-P moderately long and triple to

5-branched; and in the larvae by the combination of (1) setae 2, 3-A noticeable

subapical in attachment to antenna, (2) development and position of setae 4-6-C,

(3) comb consists of numerous scales in a triangular patch, (4) seta 1-S long,

multiple branched and attached far distad on siphon, (5) seta 1-X long, single

and attached to saddle, (6) ventral brush with 10 multiple branched setae on

erid, and (7) anal papillae small and knob-like.

Rhinoskusea is a very distinct subgenus whose exact affinities with other

subgenera of Aedes are not evident. It does share characters with several other

Aedes subgenera.

The adults of Rhinoskusea are small unornamented mosquitoes and are most

6 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

similar in habitus to those of the subgenus Levua Stone and Bohart; however,

they also resemble members of subgenera Lorrainea Belkin, Cancraedes

Edwards, Geoskusea Edwards, Veryvallina Theobald and Paraedes Edwards.

Species of these subgenera can be distinguished from those of Rhinoskusea by

one or more of the features listed above.

Males of Rhinoskusea have short maxillary palpi which are similar to those

of the following subgenera of Aedes: Aedes Meigen, Bothaella Reinert, Can-

craedes, Christophersiomyia Barraud, Huaedes Huang, Leptosomatomyia Theo-

bald, Nothoskusea Dumbleton, Paraedes, Verrvallina and most species of Geo-

skusea. Rhinoskusea males are easily separated from these subgenera by the

genitalia and one or more of the above characters.

The male genitalia of the subgenus Rhinoskusea are very distinctive, easily

separated from the other subgenera of Aedes, and show no close affinities to

any of the other subgenera of the genus. Rhinoskusea genitalia are easily

recognized by the development of the basal mesal lobe, proctiger, IX sternum,

and gonocoxite.

Female genitalia of the species of Rhinoskusea possess features which

easily distinguish them from each other and from members of other subgenera

of Aedes. The single large seminal capsule is similar to that found in Can-

craedes, Pavaedes, some African Aedimorphus Theobald and niveus group of

Finlaya Theobald, however, Rhinoskusea genitalia are easily separated from

those of these subgenera by the development of the insula which is lip-like with

long setae, but tongue-like without setae in the former 3 subgenera. From

Finlaya they are easily separated by the development of terga VIII and IX and

sternum VIII. Genitalia of Rhinoskusea are also similar to those of Geoskusea

and Pseudoskusea Theobald but are separated from them by having only a single

seminal capsule.

Pupae of Rhinoskusea species are very similar and are difficult to separate

to species. They do possess features which distinguish them from the species

of the other Aedes subgenera. Pupae of the subgenus Rhinoskusea are similar

to those of subgenera Cancraedes, Geoskusea and Pseudoskusea, however, they

can be distinguished from these 3 subgenera, as well as the other subgenera of

Aedes, by the above combination of features.

Larvae of the 3 species of Rhinoskusea are very similar and are difficult to

separate from each other, however, they possess characters that make them

easily distinguishable from other species of Aedes. Rhinoskusea larvae are

most similar to those of subgenera Cancraedes, Geoskusea and Pseudoskusea.

They can be distinguished from these subgenera, as well as all the subgenera

of Aedes, by the combination of features listed above. The comb resembles

those of Cancraedes, Geoskusea, Halaedes, Levua, Lorvainea, Pseudoskusea

and Skusea of Aedes and Opifex fuscus Hutton. The development of the anal

papillae into small knob-like structures is similar to those of other aedine mos-

quitoes which inhabit small collections of water with a higher saline content

than rain water (e.g., Aedes subgenera Cancraedes, Halaedes, Levua and

Skusea and Opifex Hutton).

DISTRIBUTION. The subgenus Rhinoskusea is found throughout much of

Southeast Asia, Sri Lanka, Papua New Guinea and the northern tip of Australia.

Species of the subgenus are confined to coastal and river areas which contain

brackish water. The distributions of the 3 species of Rhinoskusea are shown

on the map in Figure 1.

BIONOMICS. Immatures of the species of Rhinoskusea usually inhabit

brackish water in crab holes or small ground pools located in coastal mangrove

areas. They have, ona few occasions, been collected from several other types

Reinert: Aedes (Rhinoskusea) t

of small containers in coastal areas such as: beached boat, tin plate, depres-

sion in a log, discrete containers, obstructed earth drain and rock pools.

Adults have been collected in light traps, human- and carabao-baited traps

and resting in crab holes. Females have been taken while feeding on man.

MEDICAL IMPORTANCE. Few investigations have been conducted for

pathogen isolation on members of the subgenus even though the females feed on

man and other animals. One attempt at arbovirus isolation from specimens of

Rhinoskusea from Sarawak was negative (Macdonald et al. 1965).

KEYS TO SPECIES OF AEDES (RHINOSK USEA) *

FEMALE GENITALIA

1. Cercus index 2.07-2.17; tergum VIII with setae on apical 0. 45-0. 58.

pillaii

Cercus index 2. 38-3.20; tergum VIII with setae on apical 0.84-0.93. . 2

2. Cereus Anges 2. BS - 210 ee BOE 8 wardi

Cereus Index 2.613. 20 eS a POs longirostris

MALE GENITALIA

1. Basal mesal lobe not bifid, 12-19 long setae; sternum IX without setae.

pillaii

Basal mesal lobe bifid, 2-5 short apical setae on ventromesal arm;

Ssternuny 1X Wi 3-0 SCG. ee ee a a a a ie

2. Basal mesal lobe with a long stout seta at base; gonostylus narrow at

apex; 0.03-0. 63: leneth of gonccoxile, <6 eee es wardi

Basal mesal lobe without a seta at base; gonostylus broadened at apex,

0703-0. 70 teeth of conccoxite, 2 6. 6 a ee .. longirostris

PUPAE

1. oO Paadie nidex 108-1900, a On ae Ee eS longirostris

Paddle index 1: 39=4 90 SP 6 a EP ew OT Cee 2

2c Beta 1 FP On 1A) Swarr Sine er ea oe es pillai

Seta 11 double at least on one side of IJI-V. ..........26.-. wardai

FOURTH STAGE LARVAE

1. Comb with 66-104 scales. ....... i Ne es longirostris

Comb with 34-04: otalee 636 ee a ee i a ea ea

2. Seta 8-S single; seta 8-M with 6-7 branches. ............ wardi

Seta 8-S usually double; seta 8-M with 8-10 branches. ...... pillaii

*Adult females can not be separated with certainty except by the genitalia.

8 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

SPECIES TREATMENT

AEDES (RHINOSKUSEA) LONGIROSTRIS (LEICESTER)

(Figs. 2, 3, 6, 9, 12, 14)

Ficalbia Longirostris Leicester 1908: 228 (2, “).

Ficalbia longirostris of Brunetti 1912: 494; Brunetti 1920: 172.

Uranotaenia hilli Taylor 1918: 841 (*).

Aedes longirostris of Edwards 1922a: 259 (key); Edwards 1926: 120 (L*);

Stojanovich and Scott 1965: 13, 19 (2*, L*, keys); Stojanovich and Scott

1966: 49, 109 (2*, L*, keys); Parrish 1968a: 4, 5; Parrish 1968b: 2, 4;

Parrish 1969: 554.

Skusea longirostris of Edwards 1917: 224.

Aedes (Skusea) longirostris of Edwards 1922b: 467; Edwards 1924: 391

(?, “*); Barraud 1928: 363 (o*),

Aedes (Rhinoskusea) longirostris of Edwards 1932: 160; Barraud 1934: 216

(2, o*, L); Taylor 1934: 19; Causey 1937: 413; Lee 1944: 79 (L*); Knight

et al. 1944: 25 (A, key); Brug and Bonne-Wepster 1947: 185; Horsfall

1955: 411; Army Mosq. Proj. 1965: 17, 64 (keys); Stone et al. 1966: 51,

59 (keys); Mattingly 1971: Fig. 42e (P*); Grothaus et al. 1971: 18.

Aedes (Rhinoskusea) longirostris in part of Macdonald 1957: 22; Mattingly 1958:

38 (¢*, o*, P*, L*); Stone et al. 1959: 178; Steffan 1966: 211; Basio

1971: 26, 130 (*); Stone and Delfinado 1973: 307.

Aedes (Rhinoskusea) pillaii Mattingly, in part of Mattingly 1958: 21, 39, 40

(2*).

FEMALE (Fig. 2). Head. Antenna dark brown, 0. 85-0. 94 length of pro-

boscis, pedicel dark brown with a few short fine brown hairs mesally; clypeus

dark brown, bare; maxillary palpus dark brown scaled, 4-segmented, approxi-

mate length of each segment in parenthesis, 1 (0.16), 2 (0.15), 3 (0.17) and

4 (0. 52), total length 0.13-0.14 length of proboscis; proboscis long, narrow,

dark brown scaled, 1.09-1.15 length of femur I; ocular setae dark brown,

numerous; vertex and lateral surface covered with broad decumbent blackish-

brown scales, postgena with a small patch of pale brown scales; occiput with a

few erect forked blackish-brown scales. Thorax. Scutal integument dark

brown; scutum covered with narrow curved dark brown scales except for a very

small patch of broad dark brown scales on median anterior promontory area;

prescutellar space with a narrow median longitudinal patch of narrow curved

dark brown scales; scutellum with a patch of broad dark brown scales on each

lobe; scutal setae blackish-brown, numerous, on following areas: 2 long and

3-6 short median anterior promontory, numerous acrostichal (anterior and

posterior), numerous dorsocentral (anterior and posterior), scutal fossal

(2-3 long and 3-6 short anterior, 3-4 long lateral and 2-3 long posterior),

numerous supra-alar, 6 posterior medial scutal, 1 postalar callar and scutel-

lar (3 long and 3 short to 4 long and 4 short ones on lateral lobe, 5 long and 4

short to 7 long and 6 short ones on median lobe); pleural integument brown;

antepronotum without scales, 8-14 dark setae; postpronotum without scales, 1

long and 1-2 short posterior dark setae; propleuron without scales, 4-6 (usually

5-6) dark setae; prosternum, subspiracular area, paratergite, mesomeron and

metameron bare; postspiracular area without scales, 1-4 dark setae; mesepis-

ternum with a small patch of pale brown scales on lower posterior area (scales

often rubbed off), 1-2 upper and 4-7 long and 5-7 short posterior dark setae;

Reinert: Aedes (Rhinoskusea) 9

prealar knob without scales, 2-5 (usually 3-4) dark setae; mesepimeron with-

out scales, 5-10 (usually 6-8) short dark setae on posterior dorsal area. Legs.

Coxae I-III each with several brown setae, I withanterior surface brown scaled witha

small dorsal and a small ventral patch of pale scales, II and III each with a small

patch of pale scales on anterior surface; trochanters I-III each with pale scales

and a few short setae; femora I-III each with anterior surface blackish-brown

scaled, II also with a narrow anteroventral stripe of pale scales on basal area,

I-III each with posterior surface blackish brown scaled with a narrow longitudin-

al pale scaled stripe on basal area, stripe dorsal on I and ventral on II and III;

tibiae I-III and tarsi I-III each blackish-brown scaled, tarsus III with tarsomere

1as long as or slightly longer than tibia III; posttarsi I-III (Fig. 14) each with 2

ungues, each pair equal in size and simple. Wing. Dorsal and ventral veins

with dark brown scales; alula with several narrow dark brown scales on mar-

gin; upper calypter with a number of long brown setae on margin; 1 short remi-

gial seta (often hidden beneath scales). Halter. Pedicel pale; capitellum dark

brown scaled. Abdomen. Terga blackish-brown scaled, I with a small patch

of pale scales on laterotergite, II-VI each with a small laterobasal patch of pale

scales with a brown hue (several specimens from Thailand with pale scaled

patches reduced or absent); sterna II-VI pale scaled, VII with pale scales basal-

ly and brown ones on apical area; terga and sterna with numerous short setae,

mostly along posterior margins. Genitalia (Fig. 3). Tergum VIII lightly pig-

mented, base and apex nearly straight, short setae scattered over apical 0. 84-

0.92, setae along apex short, basolateral seta absent, covered with minute

spicules, scales absent, basal 0.65-0.90 retracted into segment VII, VIII-Te

index 0. 76-0. 83, VIII-Te/IX-Te index 1. 55-1. 82, length 0.16-0.17 mm, width

0.20-0.22 mm; sternum VIII lightly to moderately pigmented, base slightly

concave mesally, apex with a shallow median indentation (0.02-0.05 deep), short

setae scattered over apical 0.93-0.99, seta 1-3-S widely separated, located in

a more or less diagonal line, 1-S basomesad, 2-S approximately 0.32 from 1-S,

3-S apicolaterad and approximately 0.68 from 2-S, covered with minute spicules,

none to 5 scales (usually absent), apical intersegmental fold moderately pig-

mented, VIII-S index 0. 79-0. 86, length 0.20-0.21 mm, width 0.25-0.26 mm;

tergum IX moderately pigmented, base without median indentation, apex with

a small median indentation and with 2-5 (usually 3-4) short setae on each side

of midline, 5-9 total setae, covered with minute spicules, a short flap on

lateral area, [X-Te index 0. 73-0. 80, length 0.09-0.10 mm, width 0.12-0.13

mm; insula lightly pigmented, short, wide, lip-like, 3-5 (usually 4) long —

setae, covered with short hair-like spicules; lower vaginal lip lightly pigmented,

narrow, covered with short spicules, lower vaginal sclerite absent; upper vag-

inal lip moderately pigmented, moderately broad laterally, covered with short

Spicules, upper vaginal sclerite absent; postgenital lobe short, wide, apex broad

with a small median indentation (0. 08-0.17 deep), 4-6 (usually 5) setae on each

side of midline, 8-11 total setae, covered with short hair-like spicules, dorsal

PGL index 0. 52-0.59, ventral PGL index 1.11-1.27, ventral length 0.07 mm;

perianal membrane with scattered short spicules; cercus moderately long,

apex sharply rounded with 2-3 long stout setae, completely covered with

short spicules, dorsal surface with a number of short to moderately long setae

on apical 0. 78-0. 83, scales absent, ventral surface without setae, cercus in-

dex 2.81-3.20, cercus/dorsal PGL index 2.46-2.79, cercus length 0.17-0. 18

mm; one large seminal capsule, heavily pigmented, spherical, several small

seminal capsule pores near orifice, base of accessory gland duct moderately

pigmented.

MALE (Fig. 2). Essentially as in the female but with the following differ-

10 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

ences. Head. Antenna 0. 70-0. 82 length of proboscis, apical flagellomere

long, approximately 0.39 total length, flagellomere 12 approximately 0.21

total length, basal 11 flagellomeres 0.40 total length; maxillary palpus very

short, 0.10-0.13 length of proboscis, 4-segmented, approximate length of

each segment in parenthesis, 1 (0.18), 2 (0.16), 3 (0.23) and 4 (0. 43); pro-

boscis 1.14-1.26 length of femur I. Thovax. Setal differences as follows:

scutal fossal (2 long and 3-4 short anterior, 2-3 long lateral and 1-2 long

posterior); 4 posterior medial scutal; 5-8 on antepronotum; 3-4 on propleuron;

1-2 on postspiracular area; 1 upper and 3-5 long and 4-5 short posterior on

mesepisternum; 2-4 on prealar knob; and 4-7 on posterior dorsal area of

mesepimeron. Legs (Fig. 14). Posttarsi I-III each with 2 ungues, I and II

each with ungues unequal, larger one with a narrow basal tooth, smaller one

simple, III with ungues equal, both simple. Abdomen. Terga with laterobasal

pale scaled patches somewhat reduced in size. Genitalia (Fig. 6). Tergum IX

moderately pigmented, narrow mesally, broadened laterally, setae absent,

covered with small spicules; gonocoxite long, narrow, curved mesally, moder-

ately pigmented, covered with minute spicules, apex of dorsomesal area with

a caudally produced long finger-like projection bearing numerous moderately

long curved setae on mesal and apical surfaces, dorsal surface with a patch of

moderately long setae on apical 0.4, a few long stout setae along outer margin

on apical 0.7, basomesal area with a small narrow lobe bearing 1-3 (usually

1-2) moderately long narrow setae, ventral surface with a few long stout setae

on apical 0.5 and several short to moderately long setae on mesal area from

base to apex, a few setae somewhat stouter on basomesal area, a number of

broad scales on ventral and lateral surfaces, mesal surface membranous;

gonostylus moderately long, 0. 63-0. 70 length of gonocoxite, moderately to

heavily pigmented, basal 0.58 narrow, apical 0.42 moderately broad with

small spicules, 17-25 short setae, mostly along outer margin and apex, some

specimens with a scale basad of setae, gonostylar claw short, flared apically,

longitudinally striated, heavily pigmented, scoop-like; basal mesal lobe large,

bifurcated into a narrow heavily pigmented pointed dorsal arm and a broad

moderately pigmented ventral arm which bears a short dorsoapical flap and an

apical ventrally curved pointed flap, 2-5 (usually 3-4) short fine apical setae,

a few small spicules on basal area of ventral arm, base below furcation broad

with a moderately broad median strip connecting it with its mate and a short

dorsal extension with spicules which is connected to the ventral margin of the

small basomesal lobe of the gonocoxite; proctiger moderately long, paraproct

consists of a narrow heavily pigmented strip along outer margin from base to

apex of proctiger, apex a small heavily pigmented bluntly rounded lobe, cer-

cus membranous with a large moderately pigmented wrinkled plate covering

most of mesal area, 3-6 (usually 4-5) short cercal setae on each side of mid-

line near apex; tergum X narrow, heavily pigmented, curved, base attached to

base of paraproct and apex attached to caudoventral margin of tergum IX; phal-

losome simple, aedeagus consists of a moderately pigmented short broad

trough, base concave, apex broadly convex, paramere with length equal to

length of aedeagus, moderately pigmented, moderately broad near middle,

attached at approximately 0.37 from base to parameral apodeme, parameral

apodeme short, approximately 0.87 length of paramere, narrow, moderately

pigmented; sternum IX large, extended cephalad, lateral areas heavily pig-

mented, a narrow heavily pigmented mesal strip with 3-7 (usually 3-4) short

setae near caudal margin, remainder membranous, covered with minute spicules.

PUPA (Fig. 9). Chaetotaxy as figured and recorded in Table 1. Cephalo-

thorax and abdominal segments moderately pigmented with heavily pigmented

Reinert: Aedes (Rhinoskusea) 11

areas. Respivatory trumpet. Heavily pigmented; index 4.55-5.42, mean 4.71.

Abdomen. Seta 8-VII triple to 5 (usually 4) branched; 9-VIII with 6-10 bran-

ches; 11-III-V single. Paddle. Ovoid with inner basal margin depressed;

very few minute serrations on most of basal 0. 32-0.45 of outer margin, few

minute spicules on apical 0. 55-0. 68 of outer and apical 0. 15-0. 32 of inner

margins; midrib moderately pigmented, nearly reaches apex; seta 1-P moder-

ately long, triple to 5-branched (double on one side of one specimen); index

1.68-1.90, mean 1. 83.

LARVA (Fig. 12). Chaetotaxy as figured and recorded in Table 4. Head.

Seta 4-C with 4-5 branches; 5-C with 5-7 branches; 6-C double, occasionally

triple; 7-C with 10-14 branches; 10-C usually double, occasionally triple; 12,

13-C double or triple; dorsomentum with 20-23 teeth, usually 21. Antenna.

Seta 1-A with 5-8 (usually 6-7) branches, attached 0. 50-0. 53 from base of

antenna; 2-A very long; 3-A short, 0.24-0.29 length of 2-A; 4-A long, 0. 65-

0.73 length of 2-A; 6-A short, 0.32-0.39 length of 2-A. Thovax. Seta 8-M

with 6-9 branches; 2-T double or triple. Abdomen. Seta 13-II with 4-8 (usually

5-6) branches; 1-III, IV single; 2-X with 4 branches, 5-branched in one speci-

men; ventral brush of segment X with 10 setae on grid, each with 4-8 branches;

comb of segment VIII consists of a patch of 66-104 (usually 72-86) scales, each

scale with apical area flared and with a fringe of denticles; saddle small, in-

completely rings segment X, with short rows of small spicules. Siphon. Index

3.07-3.54, mean 3.31; pecten on basal 0. 50-0. 55 of siphon, composed of 10-15

(usually 12-14) teeth, each tooth (except occasionally basal 1-3) long, slender

and with several fine and 2-3 stouter ventral and several fine dorsal denticles

near base; seta 1-S long, triple to 5-branched, attached 0. 77-0. 84 from base

of siphon, distad of last pecten tooth; seta 8-S single.

DISCUSSION. Females of the 3 species of the subgenus are extremely

Similar in habitus features and no consistent characters were found with which

to separate them, however, several features of the genitalia can be used to

distinguish all 3 species (see discussion of pillaii for separating features).

Mattingly's (1958: 21) female genitalia figure (e) is of longirostris and not

pillaii (see discussion under type-data).

Male genitalia of longirostris are similar to those of wardi but can be

separated from them by several features (see discussion section of wardi).

Specimens of longirostris from Thailand have the ventral arm of the basal

mesal lobe narrower than specimens from Malaysia and Singapore. Mattingly

(1958: 37), on his illustration of the male genitalia, mistakenly labeled the

wrinkled mesal plate of the cercus as the phallosome.

Pupae of longirostris are very similar to those of the other 2 species of the

subgenus. Features to separate the 3 species are given in the pupal key.

The larvae of longirostris are very similar to those of both pillaiz and

wardi, however, the 3 species can be distinguished by using the characters

mentioned in the larval key. The siphonal indices can also be used to separate

longirostris and wardi.

The records of longirostris from Nakhon Phanom and Udorn, Thailand by

Parrish (1968b: 10, 16) are questionable and need confirmation. It is doubtful

that longirostris, which breeds in coastal brackish water habitats, would occur

as far inland as the above 2 localities. Unfortunately, none of the specimens

from these localities, on which the records were based, are now in existence.

TYPE-DATA. After examining all the specimens in the British Museum

(Natural History) (BMNH) (kindly loaned to me by Peter F. Mattingly) I believe

the ''series'' of specimens mentioned by Leicester (1908: 229) actually consisted

of several collections of specimens collected in the general vicinity of the mouth

12 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

of the Klang River. The only bionomic and locality data presented by Leicester

in his original description for his new species Ficalbia Longirosiris are as fol-

lows: ''The series were bred from larvae found in stagnant water at Kuala

Klang. Time of capture January.'' His description was also based on both fe-

males and males.

There are 6 males and 5 females in the BMNH that could have been from

Leicester's (1908: 229) ''series"’ of specimens. All these specimens have a

printed label with the following information: Fed. Malay States, Dr. G. F.

Leicester, 1912-350 (BMNH accession number). All the specimens are mounted

on a large octagonal shaped white paper stage (except one male mounted ona

rectangular stage) and have more or less similar information written in ink by

Leicester on the underside. However, each of the specimens bears slightly

different data and none of the specimens was collected in January. The speci-

mens possess the following data on the underside of the mounting stage (each

specimen is here numbered for identification): No. 1, “, "Klang Pt., 29/12/

03, Stagnant pool''; No. 2, “, "Stagnant pool, Klang Pt., behind houses, 29/12/

03'': No. 3, o, ''Stagnant pool behind some houses in Klang Pt., 29/12/03"; No.

4, “, "Larva fr [ from] stagnant pool behind houses, Klang pt., 29/12/03"; No.

5, o, "Larva fr mud hole, Klang Pt. (Aglyph) [ ? spelling], 27/12/03"'; No. 6,

2, "Caught in a crab hole, Klang, Daniells [ sic], 27/11/03"; No. 7, 2, "Caught

in a crab hole, Klang, Daniels, 27/11/03"; No. 8, 9, "Klang, jungle, day, 27/

11/03"; No. 9, 9, 'Klang, jungle, day, 27/11/03"; No. 10, o, ''Crab hole, K.

Klang" [ rectangular stage]; and No. 11, 9, "Port Dickson, 12/4/04". Speci-

mens 8, 9 and 10 each also had "Ficalbia" handwritten on the upper side of the

stage. Specimens 1, 2, 6, 7 and 8 have a small green rectangular card with

the number ''101"' glued to the underside of the stage while specimen No. 11 has

a number ''102" on this green card.

It is my opinion that the above specimens, Nos. 1-7 and 10, constituted

Leicester's "series" from which he described his new species longirostris.

All the specimens, which are mounted similarly, were collected in the same

general area, Klang (3902'N, 101927'E) and Kuala Klang or Klang Pt. (3°00'N,

101924'E) [ taken from Gazetteer No. 11, Malay States, Hydrographic Office,

U. S. Navy Department, H. O. Publication No. 891, October 1944], near the

mouth of the river Klang (Kuala = river or stream), collected as larvae, and

collected just prior to January, the stated time of capture (no collection date

recorded for specimen No. 10). It is possible that the specimens were col-

lected as larvae on 27 November and 27 and 29 December of 1903 and did not

emerge as adults until the following January. Specimens Nos. 8, 9 and 11 are

not considered as belonging to the type-series because of the following: Nos. 8

and 9 were collected in the jungle during the day as adults and not "bred from

larvae''; and No. 11 was collected much later on 13 April 1904 at a different site

(Port Dickson, 2°30'N, 101°48'E), had a different number (102) and no imma-

ture collection information.

The above specimens, Nos. 2-7 and 10, are relabeled as paralectotypes,

each with the following information on the label: Paralectotype, Ficalbia

longirostris Leicester. Specimens Nos. 2, 6 and 7 had previously been marked

as paratypes [ paralectotypes] and specimen No. 1 as hololectotype (sic) | lecto-

type] by P. F. Mattingly (18: [x: 1957). Mattingly's "paratype" [ paralecto-

type], specimen No. 8, and "allotype" | paralectotype], specimen No. 11, are

here considered not to belong to the original type-series for the reasons stated

above. Mattingly (1958: 39, 40) provisionally transferred a female, specimen

No. 6, to his new species, pillaii; however, the females of pillaz and longi-

rostris can not be separated with certainty by habitus features but can be dis-

Reinert: Aedes (Rhinoskusea) 13

tinguished by the genitalia. The genitalia of specimen No. 6 are definitely like

those of longirostris, therefore, this specimen is retained as a paralectotype

of longirostris. Paralectotype male, specimen No. 4, however, is pillaiias

clearly demonstrated by the structures of the genitalia. The lectotype male

contains the following data on the adult labels: [| on underside of octagonal

stage] Klang Pt., 29/12/03, Stagnant pool; 101 [ green label]; Type [ circular

label with red border]; Fed. Malay States, Dr. G. F. Leicester, 1912-350;

and Ficalbia longirostris Leicester, Hololectotype, P. F. Mattingly, 18: IX:

1957. The lectotype is in good condition with all appendages intact except for

right tarsi I-III which are mounted in Canada balsam on a microscope slide.

The scutum is rubbed and somewhat folded. The genitalia are in good condition

and are mounted in a drop of Canada balsam on a small coverslip attached to the

adult pin. —

I have examined one male paratype (with genitalia mounted) of Uvanotaenia

hilli Taylor, deposited in the BMNH, and concur that it is conspecific with

longivostrvis, The paratype possesses the following information on the labels:

G. F. Hill, 3.3.16, Darwin, N.T.; 321; larvae in crab holes mangrove swamps;

and Paratype “, Uvanotaenia hilli Taylor. The genitalia are in good condition

and are mounted in a drop of Canada balsam on a celluloid point attached to the

adult pin. The adult is in poor condition. The holotype male of hilli is deposi-

ted in the School of Tropical Medicine, Sidney, Australia.

DISTRIBUTION. 992 specimens examined: 2709, 271%, 8 pl, 81, 61 L,

and 231 with associated immature skins (137 p, 94 pl).

AUSTRALIA. Northern Territory, Darwin; 1° (paratype of hiliz).

INDONESIA. Java, Batavia; 2 L.

MALAYSIA. Sabah, Kuala Penyu; 59 pl, 1¢% pl, 130°, 1 pl, 4L. Selangor,

Klang; 22 (paralectotypes), 22, Klang Pt. ; 1% (lectotype), 40° (paralectotypes),

K. Klang; 1° (paralectotype), Carey Island, Gunong Swettenham, Jugra, Kg.

Rantau, Kg. Sireh, Port Dickson, Rantau Panjang, Telok Gong; 142 pl, 5c pl,

1° p, 389, 140°, 3 pl, 2L. Trengganu, Kemaman; 29.

PHILIPPINE ISLANDS. Sulu Archipelago, Tawi Tawi Group, Sanga Sanga

Island, Lapit Lapit; 12, 1%.

SINGAPORE. Kuala Simpong, Paya Lebar Airport area, Pulau Ubin, West

coast road at 8,5 miles;232’ pl; 8o ‘pl, 172 pp" 120 p; 809; STs, -P Ip, 8 15°25

THAILAND. Chumphon, Amphoe Muang, Pak Nam Chumphon; 59, 140.

Ranong, Khao Muang Lak; 3792 pl, 10% pl, 192 p, 880° p, 179, 650°, 3 pl, 28 L.

Trat, Tran Mayom Waterfalls, Ko Chang; 32. Samut Prakan, Amphoe Muang,

Bang Po 6; 12. Samut Sakhon, Amphoe Muang, Ta Chalom; 49, 10. South

Thailand; 3c.

Distribution from literature.

AUSTRALIA. Northern Territory, Darwin (Taylor 1918: 841, Edwards

1924: 391, Taylor 1934: 19, Lee 1944: 79, Mattingly 1958: 38). |

PAPUA NEW GUINEA (Australian Trust Territories). North coast; New

Ireland (Peters in Peters and Christian 1963: 54); New Guinea (Steffan 1966:

2%),

INDIA. Andaman Islands, Port Mount (Barraud 1928: 363; 1934: 217,

spelled Port Mouat).

INDONESIA. Java, Batavia (Mattingly 1958: 39).

MALAYSIA. Kuala Klang (Leicester 1908: 229); Selangor, Kampong Sireh,

Rantau Panjang (Macdonald 1957: 22); Port Dickson (Mattingly 1958: 38).

SINGAPORE. Naval Base (Edwards 1926: 120); Blakang Mati Island (Col-

less 1957: 110).

14 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

THAILAND. Haad Raj Samran (Causey 1937: 413); Ban U-Tapao (Parrish

1968b: 7); Chantaburi, Kloong District, Ban Bangcham, Ban Ja Kayang; Chol-

buri, Muang District (Harinasuta et al. 1974: 111-21).

VIETNAM. Phu Cat, Tuy Hoa (Parrish 1968a: 20,23); Phu Cat Air Base,

Tan Son Nhut Air Base, Tuy Hoa Air Base (Parrish 1969: 554); Chu Lai, Da

Nang, Khue Bac, Phu Bai (Grothaus et al. 1971: 18).

BIONOMICS. The normal immature habitat is brackish water in crab :

holes and small ground pools located at or near sea level. Immatures in Malay-

sia were collected from colored, brackish, temporary, unmoving water in

small crab holes, lobster burrows, a small ditch, and small ground pools,

located in unshaded or partially shaded areas of mangrove swamps and coastal

coconut groves at sea level. In Sabah larvae were taken from brackish water

in small ground pools and small buffalo footprints located in a coconut palm

area at sea level. Immatures in Singapore were collected from colored, brack-

ish, temporary, unmoving water in small 1.0-1.5 m deep crab holes, small

ground pools and a small marshy depression with crab holes, in unshaded and

partially shaded areas of a mangrove salt marsh, open swamp, and coastal

coconut palm areas at or near sea level. In Thailand immatures were collected

from clear, brackish, temporary, unmoving water in small crab holes and a

small ground pool located in partially shaded areas of mangroves and at an

elevation near 3 m.

Adults have been taken in Malaysia and Thailand in the jungle during the

day. In Singapore, Malaysia and Thailand adults were collected resting in

crab holes and in the latter 2 countries females were taken feeding on man.

In Malaysia immatures are reported from stagnant water (Leicester 1908:

229), small shady ground pools under nipah palms (Mattingly 1958: 40), and in

ground pools and crab holes in coastal swamps (Macdonald 1957: 22). In Singa-

pore Edwards (1926: 120) reports larvae collected from a mangrove area and

Colless (1957: 108-14) records the immatures collected from mangrove swamp

pools, an obstructed earth drain, discrete containers, and large and small

crab holes. In Thailand immatures were found in rock pools containing brack-

ish water (Causey 1937: 413) and from weedy ponds (Harinasuta et al. 1974:

118).

Adults were collected from crab holes in the Andaman Islands (Barraud

1928: 363; 1934: 217), from crab holes and light traps in Vietnam (Grothaus

et al. 1971: 18), and from CDC light traps in mangrove forests, human bare

leg collection in thick mangrove areas and a net trap using human bait in Thai-

land (Harinasuta et al. 1974: 111-7).

AEDES(RHINOSKUSEA) PILLAI MATTINGLY

(Figs. 4, 7, 10, 14)

Aedes (Rhinoskusea) pillaii Mattingly 1958: 40 (2*, o*, L*).

_ Aedes (Rhinoskusea) pillaii of Stone et al. 1959: 178; Stone and Delfinado 1973:

307. |

FEMALE. Description as for longirostris except for the following minor

differences. Head. Antenna 0. 87-0. 89 length of proboscis; maxillary palpus

0.13 length of proboscis; proboscis 1.03-1.19 length of femur I. Thorax.

Pleural integument somewhat paler brown; pleural setae as follows: 7-10 on

antepronotum; 1-2 long and 1 short on postpronotum; 3-5 on propleuron; 1-3

on postspiracular area; 1-2 upper and 3-4 long and 5-6 short posterior on

Reinert: Aedes (Rhinoskusea) is

mesepisternum; 2-4 on prealar knob; and 4-7 on posterior dorsal area of mese-

pimeron. Genitalia (Fig. 4). Tergum VIII lightly pigmented, base and apex

nearly straight, short setae scattered over apical 0.45-0.58, setae along apex

short, basolateral seta absent, covered with minute spicules, none to 2 scales

(usually absent), basal 0. 65-0. 80 retracted into segment VII, VIII-Te index

0. 68-0. 73, VIII-Te/IX-Te index 1. 44-1.73, length 0.16-0.18 mm, width 0. 24-

0.26 mm; sternum VIII lightly to moderately pigmented, base slightly concave

mesally, apex with a moderately deep median indentation (0. 13-0.16 deep),

numerous short setae scattered over apical 0. 88-0.97, setae 1-3-S widely

separated, located ina more or less diagonal line, 1-S basomesad, 2-S approxi-

mately 0.33 from 1-S, 3-S apicolaterad and approximately 0.67 from 2-S, cov-

ered with minute spicules, scales absent, apical intersegmental fold moder-

ately pigmented, VIII-S index 0.72-0.74, length 0.22-0.23 mm, width 0. 30-

0.31 mm; tergum IX moderately to heavily pigmented, median area wrinkled,

base without median indentation, apex with a median indentation and with 2-4

(usually 3) moderately stout and moderately long setae on each side of midline,

o-7 total setae, covered with minute spicules, a short flap on lateral area,

IX-Te index 0. 79-0. 83, length 0.10-0.11 mm, width 0.13 mm; insula lightly

pigmented, short, wide, lip-like, 4 long setae, covered with short hair-like

spicules; lower vaginal lip moderately pigmented, narrow, covered with short

spicules, lower vaginal sclerite absent; upper vaginal lip moderately to heavily

pigmented, moderately broad to broad laterally, covered with minute spicules,

upper vaginal sclerite absent; postgenital lobe short, wide, apex broad witha

small median indentation (0. 09-0.13 deep), 5-7 (usually 6) setae on each side of

midline, 11-13 total setae, covered with short hair-like spicules, dorsal PGL

index 0. 52-0.99, ventral PGL index 1.02-1.16, ventral length 0. 07-0.08 mm;

perianal membrane with scattered short spicules; cercus moderately long,

apex sharply rounded with 3-4 long stout setae, completely covered with short

spicules, dorsal surface with a number of short to moderately long setae on api-

cal 0.74-0.85, scales absent, ventral surface without setae, cercus index 2. 07-

2.17, cercus/dorsal PGL index 1. 86-2.28, cercus length 0.14-0.16 mm; one

large seminal capsule, heavily pigmented, spherical, several small seminal

capsule pores near orifice, base of accessory gland duct moderately pigmented.

MALE. Essentially as in the female but with the following differences.

Head. Antenna 0.73-0.77 length of proboscis, lightly plumose; maxillary pal-

pus short, 0.09-0.10 length of proboscis; proboscis 1.06-1. 08 length of femur

I. Thorax. Setal differences as follows: scutal fossal (2 long and 3-4 short

anterior, 3 lateral and 1-2 posterior); 4 posterior medial scutal; 5-9 on ante-

pronotum; 1-2 long and 1 short on postpronotum; 5-8 on propleuron; 1-2 on

postspiracular area; 1 upper and 2 long and 4-5 short posterior on mesepistern-

um; 2-3 on prealar knob; and 5-6 on posterior dorsal area of mesepimeron.

Legs (Fig. 14). Posttarsi I-III each with 2 ungues, I and II with ungues unequal

in size, all simple, III with ungues equal in size, both simple. Abdomen. Ter-

ga with laterobasal pale scaled patches somewhat reduced in size. Genitalia

(Fig. 7). Tergum IX heavily pigmented, narrow mesally, broadened laterally,

2-5 (usually 3-4) long stout setae on each side of midline on caudal margin,

covered with minute spicules; gonocoxite long, narrow, curved mesally, heavily

pigmented, covered with minute spicules, dorsal surface with an elongated

patch of short to moderately long setae on basal 0.75 of mesal area, several

long stout setae on median area from base to near apex, ventral surface with

a few long stout setae on apical 0.6 and several short setae along mesal mar-

gin from base to apex, apicomesal area of ventral surface with a very short

projection bearing a few moderately long curved setae, a number of broad

16 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

scales on ventral and lateral surfaces, mesal surface membranous; gonostylus

moderately long, 0.43-0.49 length of gonocoxite, broad, somewhat rectangular

in shape, apex with 7-8 short setae and a mesal apical projection bearing sub-

apically the gonostylar claw which is short, flared apically, longitudinally

striated, heavily pigmented, scoop-like; basal mesal lobe consists of a long

free narrow heavily pigmented lobe bearing 12-19 (usually 13-14) moderately

long setae on mesal margin, outer margin of lobe folded ventrally into a broad

flap, basal area broad, moderately pigmented, covered with short hair-like

spicules, connected with its mate by a moderately broad strip which is covered

with small hair-like spicules; proctiger moderately long, paraproct consists of

a narrow heavily pigmented strip along outer margin from base to apex of

proctiger, apex a small heavily pigmented bluntly rounded lobe, cercus mem-

branous with a large lightly to moderately pigmented wrinkled plate covering

most of mesal area, 3-5 (usually 3-4) short cercal setae on each side of mid-

line near apex; tergum X narrow, heavily pigmented, curved, base attached

to base of paraproct and apex attached to caudoventral margin of tergum IX;

phallosome simple, aedeagus consists of a moderately pigmented short broad

trough, base concave, apex broadly convex with a very small median indenta-

tion, paramere approximately 0.88 length of aedeagus, moderately to heavily

pigmented, moderately broad near middle, attached at approximately 0.44 from

base to parameral apodeme, parameral apodeme approximately 0.87 length of

paramere, narrow, moderately pigmented; sternum IX large, extended cephalad,

lateral areas heavily pigmented, remainder membranous, setae absent, covered

with minute spicules.

PUPA (Fig. 10). Chaetotaxy as figured and recorded in Table 2. Cephalo-

thorax and abdominal segments I-III moderately pigmented with a few small

heavily pigmented areas, remainder of abdomen moderately pigmented. Res-

piratory trumpet. Heavily pigmented; index 4.89-5.93, mean 5.41. Abdomen.

Seta 8-VII double or triple, usually double; 9-VIII with 5-8 branches; 11-III-V

single. Paddle. Ovoid with inner basal margin depressed; a few minute ser-

rations on most of basal 0. 43-0.49 of outer margin, a few minute spicules on

apical 0.51-0.66 of outer and apical 0.13-0.30 of inner margins; midrib moder-

ately pigmented, nearly reaches apex; seta 1-P moderately long, triple to 9-

branched; index 1.41-1.57, mean 1.47.

LARVA (Fig. 14). Chaetotaxy as figured and recorded in Table 5. Head.

Seta 4-C double to 4-branched, usually double or triple; 5-C with 5-7 branches;

6-C double; 7-C with 10-16 (usually 10-14) branches; 10-C double to 4-branched,

usually triple; 12-C double or triple; 13-C double to 4-branched, usually triple;

dorsomentum with 19-22 teeth, usually 21. Antenna. Seta 1-A with 5-7 (usually

5-6) branches, attached 0. 42-0.52 from base of antenna; 2-A very long; 3-A

short, 0.22-0.25 length of 2-A; 4-A long, 0.55-0.66 length of 2-A; 6-A short,

0. 33-0.35 length of 2-A. Thorax. Seta 8-M with 8-10 (usually 8) branches;

2-T single or double, usually single. Abdomen. Seta 13-II with 4-6 (usually

5-6) branches; 1-III, IV single; 2-X with 5 branches, occasionally 4- or 6-

branched; ventral brush of segment X with 10 setae on grid, each with 4-8

branches; comb of segment VIII consists of a patch of 34-54 (usually 39-50)

scales, each scale with apical area flared and with a fringe of denticles; saddle

small, incompletely rings segment X, with spicules, not in rows. Szphon.

Index 2. 46-3.16, mean 2.93; pecten on basal 0. 44-0. 52 of siphon, composed of

4-10 (usually 5-9) teeth, each tooth (except occasionally basal 1-3) long, slender

and with a few fine and 2-3 stouter ventral and occasionally a few fine dorsal

denticles near base; seta 1-S long, with 4-6 (usually 5-6) branches, attached

0. 79-0. 84 from base of siphon, distad of last pecten tooth; seta 8-S double,

Reinert: Aedes (Rhinoskusea) 17

Single in one specimen.

DISCUSSION. Females of longirostris, pillaii and wardi can not be separ-

ated with certainty using habitus features, however, they can be distinguished

by the female genitalia. Female genitalia of pillaii possesses: sternum VIII

with a moderately deep (0. 13-0.16 of total sternum length) median apical inden-

tation, tergum VIII with setae only on apical 0.45-0.58, and cercus broader with

index of 2.07-2.17. Aedes longirostris possesses the following: sternum VIII

with a very shallow (0. 02-0. 05) median apical indentation, tergum VIII with

setae on apical 0.84-0.92, and cercus with index of 2.81-3.20, while wardi

possesses the following: sternum VIII with a very shallow (0. 03-0. 04) median

apical indentation, tergum VIII with setae on apical 0.93-0.98, and cercus

with index of 2. 38-2. 70.

The male genitalia of pillaii are easily separated from those of the other 2

species of the subgenus by the very broad gonostylus, shape of the basal mesal

lobe, and absence of both basal and apical lobes on the dorsal surface of the

gonocoxite, presence of very well developed setae on tergum IX, and absence

of setae on sternum IX.

Males of pillaii can be distinguished from those of longirostris and wardi

by posttarsi I and II which have the ungues simple while the latter 2 species

each have the larger ungue of posttarsi I and II with a narrow basal tooth.

Larvae of pillaii are separated from both longirostris and wardi by setae

8-S and from longirostris by the number of comb scales and pecten teeth.

TYPE-DATA. The holotype male, deposited in the BMNH, possesses the

following data on the adult labels: MALAYA, SELANGOR, Kompong Sireh, 31:

iii: 1953, J. A. Reid; 604 [ collection number]; A. Rhinoskusea sp. indet., Det.

J. A. Reid; Type | circular label with red border |]; and Aédes (Rhinoskusea)

pillati Mattingly, Holotype ©, P. F. Mattingly, 15: X: 57. The holotype adult

is in excellent condition with all appendages intact except for right tarsi I-III

which are mounted in-Canada balsam on a microscope slide. The genitalia are

in good condition and are mounted in a drop of Canada balsam on a celluloid

point attached to the adult pin. Mattingly (1958: 40) also records the following

for the holotype 'from larvae in small shady ground pool under Nipah palm."

The allotype is in fair condition, however, it is rubbed and 3 legs, abdomen

and genitalia are missing. Larval and pupal skins from the allotype are mounted

on a microscope slide. The following information is included on the adult labels:

SELANGOR, Rantau Panjang, 3.7.1952, J. A. Reid; Ex. Coll. I.M.R. Malaya

[on underside of above label]; 463/5 [ collection number]; allotype [ circular

label with red border]; Aédes (Rhinoskusea) pillaii Mattingly, Allotype 2, P. F.

Mattingly, 15: X: 57.

Paratypes are as follows and bear the same collection data as listed for the

allotype except the collection numbers. One paratype male genitalia is mounted

in a drop of Canada balsam on a celluloid point attached to a pin with labels

(adult missing) which possess the collection number 463/2. A microscope slide

with larval and pupal skins contains the same number (463/2). A slide with lar-

val and pupal skins bears the collection number 463/3 and is labeled a paratype.

The adult male from these skins could not be located (Mattingly 1958: 40). I.

have examined all the type material which is deposited in the BMNH.

DISTRIBUTION. 36 specimens examined: 49, 180", 1 pl, 7 with associated

immature skins (2 p, 5 pl).

MALAYSIA. Selangor, Kampong Sireh; 1° (holotype), Rantau Panjan; 12 pl

(allotype), 1° genitalia, 1 p, 1 1p (paratypes), Carey Island, Klang; 110, Klang

Pt.; 1% (paralectotype no. 4 of longirostris).

SINGAPORE. West coast road at 8.5 miles; 22 pl, 1% pl, 12 p, 1c’ p, 2c.

18 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

Distribution from literature.

MALAYSIA. Selangor, Kampong Sirek, Rantau Panjang (Mattingly 1958:

40).

BIONOMICS. Immatures in Singapore were collected from colored, brack-

ish, temporary, unmoving water in small 1mdeepcrabholes, in unshaded and par-

tially shaded areas of a salt marsh and an open swamp at sea level. Adults

were taken in Malaysia resting in crab holes in a mangrove swamp area.

Mattingly (1958: 40) reports the larvae collected in Malaysia from a crab

hole and from a small shady ground pool under a nipah palm.

AEDES (RHINOSK USEA) WARDINEW SPECIES

(Figs, 5, 8, 11, 13;°14)

Aedes longirostris of Senior-White 1927: 69.

Aedes (Rhinoskusea) longirostris of Carter 1950: 89; Chow et al. 1954: 117;

Knight and Hull 1952: 163 (¢, o*, L*); Baisis 1974: 27 (2*, o*, P*, L*).

Aedes (Rhinoskusea) longirostris in part of Macdonald 1957: 22; Stone et al.

1959: 178; Delfinado et al. 1962: 439; Steffan 1966: 211; Mattingly 1958:

38; Basio 1971: 26; Stone and Delfinado 1973: 307.

FEMALE. Description as for longirostris except for the following minor

differences. Head. Antenna 0. 88-0.92 length of proboscis; maxillary palpus

0.13-0.14 length of femur I; postgena with a large patch of pale scales. Thorax.

Pleural setae as follows: 9-11 on antepronotum; 1-2 long and 1 short on post-

pronotum; 2-4 (usually 3-4) on propleuron; 2-4 on postspiracular area; 1-2

upper and 4-6 long and 3-6 short posterior on mesepisternum; 3-5 (usually 4)

on prealar knob; and 5-7 (usually 5) on posterior dorsal area of mesepimeron.

Abdomen. Terga II-VI each with laterobasal pale scaled patches larger in size

and whiter colored. Genitalia (Fig. 5). Tergum VIII lightly pigmented, base

and apex straight, short setae scattered over apical 0. 86-0.93, setae along

apex short, basolateral seta absent, covered with minute spicules, scales ab-

sent, basal 0.75-0.90 retracted into segment VII, VIII-Te index 0. 74-0. 83,

VIlIl- Te/IX-Te index 1.74-1.91, length 0.17-0.19 mm, width 0.22-0.23 mm;

sternum VIII lightly pigmented, base slightly concave mesally, apex with a

shallow median indentation (0.03-0.04 deep), numerous short setae scattered

over apical 0.93-0.98, setae 1-3-S widely separated, located in a more or less

diagonal line, 1-S basomesad, 2-S approximately 0.29 from 1-S, 3-S apico-

laterad and approximately 0.71 from 2-S, covered with minute spicules, scales

absent, apical intersegmental fold moderately pigmented, VIII-S index 0. 75-

0.87, length 0.21-0.22 mm, width 0.27-0.28 mm; tergum IX moderately pig-

mented, base without median indentation, apex with a small median indentation

and with 3-6 short setae on each side of midline, 5-11 total setae, covered

with minute spicules, a short flap on lateral area, IX-Te index 0. 72-0. 87,

length 0.09-0.10 mm, width 0.13 mm; insula lightly pigmented, short, wide,

lip-like, 4 long setae, covered with small hair-like spicules; lower vaginal

lip lightly pigmented, narrow, covered with short spicules, lower vaginal

sclerite absent; upper vaginal lip moderately pigmented, moderately broad

laterally, covered with short spicules, upper vaginal sclerite absent; postgeni-

tal lobe short, wide, apex broad with a very small median indentation or straight

(0.00-0.13 deep), 3-6 setae on each side of midline, 7-11 total setae, covered

with small hair-like spicules, dorsal PGL index 0.38-0.52, ventral PGL index

1.00-1.35, ventral length 0. 06-0. 08 mm; perianal membrane with scattered

Reinert: Aedes (Rhinoskusea) 19

short spicules; cercus moderately long, apex sharply rounded with 3-4 long

stout setae, completely covered with short spicules, dorsal surface with a

number of short to moderately long setae on apical 0.71-0.81, scales absent,

ventral surface without setae, cercus index 2. 38-2.70, cercus/dorsal PGL

index 2.09-2.77, cercus length 0.16-0.17 mm; one large seminal capsule,

heavily pigmented, spherical, several small seminal capsule pores near ori-

fice, base of accessory gland duct moderately pigmented.

MALE. Essential as in the female but with the following differences.

Head. Antenna 0. 68-0. 83 length of proboscis; maxillary palpus very short,

0. 09-0. 12 length of proboscis; proboscis 1.16-1.27 length of femur I. Thorax.

Setal differences as follows: scutal fossal (2 long and 3-6 short anterior, 3

lateral and 2 posterior); 4 posterior medial scutal; 5-9 on antepronotum; 1

long and 1 short on postpronotum; 3-4 on propleuron; 1-3 on postspiracular

area; 1 upper and 4-5 long and 3-6 short posterior on mesepisternum; 2-4 on

prealar knob; and 5-6 on posterior dorsal area of mesepimeron. Legs. Post-

tarsi I-III (Fig. 14) each with 2 ungues, I and II each with ungues unequal in

size, larger one with a narrow basal tooth, smaller one simple, III with un-

sues equal in size, both simple. Abdomen. Terga with laterobasal white

scaled spots somewhat reduced in size. Genitalia (Fig. 8). Tergum IX moder-

ately to heavily pigmented, narrow mesally, broadened laterally, none to 3

setae, covered with small spicules; gonocoxite long, narrow, curved mesally,

moderately to heavily pigmented, covered with minute spicules, apex of dorso-

mesal area with a caudally produced long finger-like projection bearing numer-

ous moderately long curved setae located mostly on mesal and apical surfaces,

dorsal surface with a patch of moderately long setae on apical 0.4, a few long

stout setae along outer margin on apical 0.7, basomesal area with a small

narrow lobe bearing 3-8 (usually 4-6) moderately long narrow setae, ventral

surface with a few long stout setae on apical 0.6 and several short to moder-

ately long setae on mesal area from base to apex, a few setae somewhat

stouter on basomesal area, a number of broad scales on ventral and lateral

surfaces, mesal surface membranous; gonostylus long, 0. 83-0. 88 length of

gonocoxite, moderately to heavily pigmented, narrow with a very slightly

expanded area at approximately 0.63 from base, numerous small spicules,

17-29 short setae, mostly along apical 0.5 of outer margin and apex, some

specimens with 1-2 small scales basad of setae, gonostylar claw short, flared

apically, longitudinally striated, heavily pigmented, scoop-like; basal mesal

lobe large, bifurcated into a long moderately broad dorsal arm with a bluntly

rounded apex, and a very broad heavily pigmented ventral arm which bears a

short dorsoapical flap and a broad short apical ventrally curved flap with a

short point near its base, 3-5 (usually 4-5) short fine apical setae, a single

moderately long stout setae on mesal margin at point of furcation, base below

furcation broad with a moderately broad median strip with short spicules con-

necting it with its mate and a short dorsal extension with spicules which is

connected to the ventral margin of the small basomesal lobe of the gonocoxite;

proctiger moderately long, paraproct consists of a narrow heavily pigmented

strip along outer margin from base to apex of proctiger, apex a small heavily

pigmented bluntly rounded lobe, cercus membranous with a large moderately

pigmented wrinkled plate covering most of mesal area, 4-7 (usually 6-7) short

cercal setae on each side of midline near apex; tergum X narrow, heavily pig-

mented, curved, base attached to base of paraproct and apex attached to caudo-

ventral margin of tergum IX; phallosome simple, aedeagus consists of a mod-

erately pigmented short broad trough, base concave, apex broadly convex with

a very small median indentation, paramere approximately 1.13 length of aedea-

20 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

gus, moderately to heavily pigmented, moderately broad near middle, attached

at approximately 0.37 from base to parameral apodeme, parameral apodeme

approximately equal in length to length of paramere, narrow, moderately pig-

mented; sternum IX large, extended cephalad, lateral areas heavily pigmented,

a narrow heavily pigmented mesal strip with 3-8 (usually 3-4) short setae near

caudal margin, holotype also with a scale mixed with setae, remainder mem-

branous, covered with minute spicules.

PUPA (Fig. 11). Chaetotaxy as figured and recorded in Table 3. Cephalo-

thorax and abdominal segments mostly heavily pigmented. Respiratory trum -

pet. Heavily pigmented; index 4.04-5.13, mean 4.46. Abdomen. Seta 8-VII

triple to 5 (usually 4) branched; 9-VIII with 4-8 (usually 7-8) branches; 11-III-V

usually double, at least double on one side of segments. Paddle. Ovoid with

inner basal margin depressed; a few minute serrations on most of basal 0. 43-

0.45 of outer margin, a few minute spicules on apical 0. 55-0. 57 of outer and

apical 0.15-0.29 of inner margins; midrib moderately pigmented, nearly

reaches apex; seta 1-P moderately long, triple or 4-branched; index 1. 39-1. 59,

mean 1. 51.

LARVA (Fig. 13). Chaetotaxy as figured and recorded in Table 6. Head.

Seta 4-C double or triple, occasionally with 4-6 branches; 5-C with 5-7 bran-

ches; 6-C double; 7-C with 9-13 (usually 10-12) branches; 10-C double or

triple, usually triple; 12-C double; 13-C double or triple, usually double; dor-

somentum with 19-21 teeth, usually 21. Antenna. Seta 1-A with 5-6 branches,

attached 0.49-0.56 from base of antenna; 2-A very long; 3-A short, 0.22-0.27

length of 2-A; 4-A long, 0.73-0.77 length of 2-A; 6-A short, 0.39-0.42 length

of 2-A. Thorax. Seta 8-M with 6 branches, occasionally with 7 branches; 2-T

double. Abdomen. Seta 13-II with 6-9 (usually 8-9) branches; 1-III, IV single

or double, usually double; 2-X with 5-7 branches; ventral brush of segment X

with 10 setae on grid, each with 4-9 branches; comb of segment VIII consists

of a patch of 47-64 (usually 47-62) scales, each scale with apical area flared

and with a fringe of denticles; saddle small, incompletely rings segment X,

with short rows of small spicules. Siphon. Index 2.43-2.57, mean 2. 49;

pecten on basal 0. 50-0. 54 of siphon, composed of 8-12 (usually 10) teeth,

each tooth (except occasionally basal 1-3) long, slender and with a few fine and

2-3 stouter ventral and occasionally a very few fine dorsal denticles near base;

seta 1-S long, with 5-6 branches (one specimen with 4 branches on one side),

attached 0. 76-0. 81 from baseof siphon, distad of last pecten tooth; seta 8-S

single.

DISCUSSION. Females of wardi can be distinguished from the other 2

species of the subgenus only by the genitalia (see discussion of pzllaii for

separating features).

The male genitalia of wavdi resemble those of longirostris but are easily

distinguished from the later species by the longer and narrower gonostylus,

the number of setae (usually 4-6) on the small dorsal basomesal lobe of the

gonocoxite, and the basal mesal lobe which has both dorsal and ventral arms

broader and bears a moderately long stout seta basomesad of furcation.

Larvae of wardi can be distinguished from those of longirostris by the

number of comb scales and the siphonal index and from pillaii by setae 8-M

and 8-S.

Macdonald et al. (1965: 338) record collections of Aedes (Rhinoskusea)

species unidentified which fed on man at Kampong China, Kuching, Sarawak.

The specimens were destroyed by the investigators in virus isolation attempts

and therefore could not be studied here, however, it is probable that they

belonged to the present new species, wardi.

Reinert: Aedes (Rhinoskusea) 21

This new species is dedicated to Ronald A. Ward, Medical Entomology

Project Manager, in appreciation of his support.

TYPE-DATA. The type-series consists of the holotype male, allotype and

4 male and 37female paratypes. The holotype is deposited in the U. S. Nation-

al Museum (Natural History) (USNM) and possesses the following data on the

adult labels: PHILIPPINE ISLANDS, Luzon, Zambales, Subic U. S. Naval

Base, 10 June 1968, LS-3 (Tidal Swamp): S. 360-6 | collection number]; F. E.

Baisas Collection; T75. 5 [| genitalia preparation number]; and HOLOTYPE ¢,

Aedes (Rhinoskusea) wardi, John F. Reinert, April '75. The holotype is in

excellent condition with all appendages intact. The genitalia are mounted in

Canada balsam on a microscope slide and are in excellent condition. Pupal

and larval skins associated with the holotype are mounted in Canada balsam

on a microscope slide. The allotype with its associated pupal and larval skins

possesses the same collection data as the holotype except the collection num-

ber--S. 360-8. Paratypes are as follows: S.360-1(2 pl), S.360-2(c), S. 360-3

(2 pl), S.360-5(c pl), S.360-7(% pl), all with other data as in holotype; S. 331-1

(2? pl), S.331-2(¢ pl), S.331-3(¢), S.331-4(2 1), S.331-5(2 pl), S.331-6(¢ pl),

S.331-7(¢ pl), S.331-8(2 pl), S.331-9(2), S.331-10(2 pl), S.331-11(¢ pl),

S. 331-12(2), S.331-13(¢ pl), S.331-14(2), S.331-15(2 pl), S.331-16(¢ pl),

S. 331-18(2 pl), S.331-19(2), S.331-20(¢), S.221-21(2), all with data as in holo-

type except for the following--20 May 1968, larval site 3, shaded; S.375-1(¢

pl), S.375-2(2 pl), S.375-3(@ pl), S.375-4(2 pl), S.375-5(2 pl), S.375-7(2 pl),

S. 375-8(2 pl), S.375-9(2 pl), S.375-10(@ pl), S.375-11(2 1), S.375-12(2 pl), all

with data as in holotype except for the following--18 June 1968, tidal swamp,

Fairway 14; and S.376-1(¢ pl), S.376-2(2 pl), S.376-3(2 pl), S.376(19, 20°), all

with data as in holotype except for the following--18 June 1968, LS-3-A (larval

site). The type specimens are deposited in the USNM except for 2 female and

one male paratypes (with associated skins) which will be deposited in the BMNH

and 2 female paratypes (with associated skins) which will be deposited in the

Bernice P. Bishop Museum.

DISTRIBUTION. 294 specimens examined: 959, 570", 5 pl, 11, 17 L,

0? with associated larval and pupal skins and 2 with associated larval skins.

INDONESIA. Celebes, Dongkala, Kabaena by Boeton; 1°. Kalimantan,

Tarakan; 32 pl, 169, 24c".

PHILIPPINE ISLANDS. Balabac, Cape Melville; 12, 4 L. Luzon, Wandin.

Olongapo, Subic U. S. Naval Base; 1° pl (holotype), 12 pl (allotype), 27 pl,

20 pl, 221, 82, 2% (paratypes), 182 pl, 142, 160°, 5 pl, 6 L. Mindanao, Zam-

boanga, Zamboanga City; 10°. Mindoro, San Jose; 19, 4%, 7L. Palawan,

Iwahig; 12. Samar, Pintanahon; 3¢ pl, 1% pl, 4c, 11.

SRI LANKA. Trincomalee; 1.

Distribution from literature.

INDONESIA. Kabaena (Brug and Bonne-Wepster 1947: 185, Macdonald

1957: 22), Tarakan (Mattingly 1958: 39).

MALAYSIA. Sarawak, First Division, Kuching, Kompong China (Mac-

donald et al. 1965: 338),

PHILIPPINE ISLANDS. Balabac, Cape Melville; Mindanao, Zamboanga,

Zamboanga City; Mindoro, near Mangarin Bay, San Jose; Palawan, Iwahig,

Iwahig Penal Colony, Tacburos; Samar, Pintanahon near Osmena (Knight and

Hull 1952: 165); Pampanga, Clark Air Base; Laguna, Los Banos (Basio 1971:

26); Luzon, Zambales, Subic U. S. Naval Base; throughout Philippines from

Luzon to Mindanao (Baisas 1974: 28).

SRI LANKA. Trincomalie, Orr's Hill (Senior-White 1927: 69, Mattingly

1958: 39).

22 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

BIONOMICS. Immatures in the Philippine Islands were collected from

shaded and unshaded tidal swamps. Larvae were taken in a nipa palm swamp

at Tarakan, Indonesia.

In the Philippine Islands immatures were collected from brackish water in

crab holes, swamp pools near the beach and a beached boat, from a tin plate in

a grassy area, and a depression in a log (Knight and Hull 1952: 165), and from

pools and ponds affected by salt water tides (Baisas 1974: 28).

Adults in the Philippine Islands were collected resting in crab holes (Knight

and Hull 1952: 165) and females collected from carabao-baited traps (Baisas

1974: 28). Females were taken biting man in Sarawak, Malaysia (Macdonald

et al. 1965: 338).

ACKNOWLEDGMENTS

Grateful appreciation is expressed to: Bruce F. Eldridge and Ronald A.

Ward, Walter Reed Army Institute of Research, for critically reading the manu-

script; to Douglas J. Gould, U. S. Army Medical Component-SEATO, Bangkok,

Thailand, and Shivaji Ramalingam, University of Malaya, Kuala Lumpur, Ma-

laysia, for the collection and preparation of many specimens; to Peter F. Mat-

tingly, British Museum (Natural History), London, and Wallace A. Steffan,

Bernice P. Bishop Museum, Hawaii, for the generous loans of types and muse-

um specimens; to Young T. Sohn, Medical Entomology Project (MEP), for pre-

paring the illustrations; to Virginia M. Ford and Susan Hevel, MEP, for lab-

oratory assistance; to Owilda J. R. Curtis, MEP, for typing the manuscript

drafts; and to Janet D. Rupp, MEP, for typing the manuscript for offset repro-

duction.

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Reinert: Aedes (Rhinoskusea) 23

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Reinert: Aedes (Rhinoskusea) 25

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1958. The culicine mosquitoes of the Indomalayan area. Part III:

Genus Aedes Meigen, subgenera Paraedes Edwards, Rhino-

skusea Edwards and Cancraedes Edwards. Br. Mus. (Nat.

Hist.), London. 61 p.

1961. The culicine mosquitoes of the Indomalayan area Part V:

Genus Aédes Meigen, subgenus Mucidus Theobald, Ochlerota-

tus Lynch Arribalzaga and Neomelaniconion Newstead. Br.

Mus. (Nat. Hist.), London. 62 p.

1971. Contributions to the mosquito fauna of Southeast Asia. XII.

Illustrated keys to the genera of mosquitoes (Diptera, Culici-

dae). Contrib. Am. Entomol. Inst. (Ann Arbor) 7(4): 1-84.

PARRISH, D. W.

1968a. The occurrence and known human-disease relationships of

mosquitoes on USAF installations in the Republic of Vietnam.

U. S. Air Force 5th Epidemiol. Flight, 5EF-TR-68-1. 23 p.

1968b. The occurrence and known human disease relationships of

mosquitoes of USAF installations in Thailand. U. S. Air

Force 5th Epidemiol. Flight, 5EF-TR-68-1. 18 p.

1969. Species composition and human disease relationships of mos-

quitoes on U. S. Air Force bases in the Republic of Vietnam.

Mosq. News 29: 552-6.

PETERS, W. and S. H. CHRISTIAN.

1963. The bionomics, ecology and distribution of some mosquitoes

(Diptera: Culicidae) in the Territory of Papua and New

Guinea. Acta Trop. 20: 35-79.

26 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

REINERT, J. F.

1974, Terminology and preparation techniques of the female geni-

talia of aedine mosquitoes (Diptera: Culicidae). Mosq.

Syst. 6: 46-56.

SENIOR-WHITE, R.

1927. Notes on Ceylon mosquitoes, --II. The larvae of the commer

non-anopheline mosquitoes. Spolia Zeylan. Bull. Natl. Mus.

Ceylon 14: 61-76.

STEFFAN, W. A.

1966. A checklist and review of the mosquitoes of the Papuan sub-

region (Diptera: Culicidae). J. Med. Entomol. 3: 179-237.

STOJANOVICH, C. J. andJ. G. SCOTT.

1965. Illustrated key to Aedes mosquitoes of Vietnam. U. S. Dep.

Health Educ. Welfare, Public Health Serv. 34 p.

1966. Illustrated key to mosquitoes of Vietnam. U. S. Dep. Health

Educ. Welfare, Public Health Serv. 158 p.

STONE, A. and M. D. DELFINADO.

1973. Family Culicidae. p. 266-343. In M. D. Delfinado and

D. E. Hardy, Ed. A catalog of the Diptera of the Oriental

region. Volume I. Suborder Nematocera. Univ. Press

Hawaii, Honolulu. 618 p.

STONE, A., K. L. KNIGHT and H. STARCKE.

1959. A synoptic catalog of the mosquitoes of the world (Diptera,

Culicidae). Thomas Say Found., Entomol. Soc. Am. Vol. VI,

358 p.

STONE, A., J. E. SCANLON, D. L. BAILEY, M. D. DELFINADO and

R. A. BRAM.

1966. Preliminary keys to mosquitoes of Vietnam. Army Mosq.

Project, Smithson. Inst. 92 p.

TAYLOR, F. H.

1918 (1919). Contributions to a knowledge of Australian Culicidae.

No. iv. Proc. Linn. Soc. N. S. W. 43: 826-43.

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Aust. Dep. Health Serv. Publ. (School Public Health

Trop. Med.) No. 1, 24 p.

13.

14.

Reinert: Aedes (Rhinoskusea) 27

LIST OF FIGURES

Distribution of the species of Rhinoskusea

Aedes (Rhinoskusea) longirostris--adult morphology

. Aedes (Rhinoskusea) longirostris--female genitalia

. Aedes (Rhinoskusea) pillaii--female genitalia

Aedes (Rhinoskusea) wardi--female genitalia

. Aedes (Rhinoskusea) longirostris--male genitalia

Aedes (Rhinoskusea) pillati--male genitalia

Aedes (Rhinoskusea) wardi--male genitalia

Aedes (Rhinoskusea) longirostris--pupa

Aedes (Rhinoskusea) pillaii--pupa

Aedes (Rhinoskusea) wardi--pupa

. Aedes (Rhinoskusea) longirostris--larva

Aedes (Rhinoskusea) wardi--larva

Aedes (Rhinoskusea) pillaii--larva; longirostris, pillaii and wardi--female

and male posttarsi; longirostris--male tergu

and sternum VIII |

I- VII

i oll i ee The eat Heese

Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

LIST OF FIGURE ABBREVIATIONS

Female Genitalia

Accessory gland duct base

Cercus

Line of attachment of peri-

anal membrane to dorsal

surface of PGL

Hinge

Insula

Tergum 9

Aedeagus

Apodeme of gonocoxite

Basal mesal lobe

Cercus

Gonocoxite

Gonostylus

Sternum 9

Cephalothorax

Abdominal segments 1-8

Antenna

Head

Comb scale

Dorsomentum

Abdominal segments 1-8,

Ole We Te eae

ee We Ae haat ll

heb ul

Heed es te Nl

Lower vaginal lip

Postgenital lobe

Seminal capsule

Seminal capsule pore

Upper vaginal lip

Upper vaginal sclerite

Sternum 8

Tergum 8

Tergum 9

Parameral apodeme

Paramere

Paraproct

Proctiger

Sternum 8

Tergum 8

Tergum 10

Metanotal plate

Paddle

Respiratory trumpet

Mesothorax

Mouthpart

Prothorax

Pecten tooth

Siphon

Metathorax

Sol Ost Sel Oz sol 06

Cl Sl

11esfid

1psem

$14]80/1Bu0/

° 6

Les

Sal

be be:

pibahaies

is ad

Aedes (Rhinoskusea) /ongirostris

; (Wares ep

Nae a a

eau he,

VINI-S

Vill-Te

Aedes (Rhinoskusea) /ongirostris

JES.

hls

dorsal

os Z

Vill-Te

Aedes (Rhinoskusea) pillaii

Po cuten ne. ile

Rel Seeiv| DSAI A! be

AA US Wat ites et 4)

\ Viphi i i Wd Fl

feral |

fry id

fait

ee ae

eel ie

{ fea il

\ ae se ae! fing

\\ a Fr

‘|

Wey

Ne NE NSN

Se i |

Dh ENG NV re Net kl

ane Ay ver ese A \

Wee fi

A athe ot |

N ee

Q ay o

AL Paice

ay |

ie eh, i rete ee

VIII-S

ventral

| | | hee

\! Ae Accel tle ctelire eqn ea panel | fiche Matsa Tht! (MIBal taka ay }/

Saeed (ylal Tigh A jay et bp rey ploy ty lat (Mal wey ! ees:

al Uae hey jl Py eb pa ghap Geta kt gob tle

a yan by My deree Gaee te peel Iee

| x

dorsal

Vill-Te

Aedes (Rhinoskusea) ward

BML

Thailand

dorsal

Aedes (Rhinoskusea) /ongirostris

Ay ee) Bie ee eae ahr)

Aedes (Rhinoskusea) pillali

aa 9 hos .

ls te apie n, Ue pith, eps ee

aonb ~ ‘ erent aa tied potty nit ctottotetent tad Boy

1 , ae oR “x

a ay rt

¥ /

dorsal

Aedes (Rhinoskusea) wardi

7 \ A \

N\ aS)

Aedes (Rhinoskusea) /ongirostris

Aedes (Rhinoskusea) pillaii

Aedes (Rhinoskusea) wardi

| Fig. 12

Aedes (Rhinoskusea) /ongirostris

Aedes (Rhinoskusea) ward

j i

TLL yt

4‘;

sey

& Vill-Te

0.2

ay) Ng)

bent CRYO)

rant: WN YD

RAEN EST Te Vrain

longirostris & os £11 of VIII-S

pS ee ey nae longirostris

Reinert: Aedes (Rhinoskusea) 45

TABLE 1. Record of the branching of the setae on the

pupae of Aedes (Rhinoskusea) longirostris

(7 specimens).

Seta Range Mode Seta Range Mode _Seta__Range Mode

Cephalothorax Abdomen I (Cont. ) Abdomen III (Cont. )

1 2-3 2 10 1 1 9 Val ih

2 2-3 3 11 1 1 10 2-3 2

3 2-3 3 Abdomen II 11 ‘ 1

4 1-3 2 0 1 1 14 1 1

5 2-3 3 1 13-25 13 Abdomen IV

6 1 1 2 1 1 0 1 1

q 2-3 3 3 2-4 3 1 2-4 3

8 3-4 4 4 2-4 3 2 1-2 i

9 1-2 2 5 2-6 4 3 2-6 5

Metanotal Plate 6 2-4 2 4. 2-4 3

10 3-5 3 7 2-3 2 5 3-4 4

11 2 2 9 1 1 6 1-3 2

12 2-5 3 Abdomen III 7 2-4 3

Abdomen I 0 1 1 8 2-4 3

1 18-36 27 1 3-6 4 9 1 1

2 1 1 2 1-2 1 10 1-3 2

3 2-3 2 3 2 2 11 1-3 1

4 3-6 5 | 4 2-4 3 14 1 1

D 2-6 4 5 2-6 2 Abdomen V

6 1-3 2 6 1-3 2 0 1 1

| 1-3 2 7 3-5 3 1 2-4 3

46 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

TABLE 1 (Cont. )

Seta Range Mode Seta Range Mode _. Seta Range Mode

Abdomen V (Cont. ) Abdomen VI (Cont. ) : Abdomen VII (Cont. )

eee 3 4 a eee

ee: 3 : 2-4 oe 6 2-5 5

po acd 3 .6 2-5 4 7 2 2

6. 14 3 7 122 2 a ee8 4

fo gc4 3 8 2-5 3 9 3-5 3

8 2-4 3 9 1 a 0 2-3 2

9 1 1 4 2 ‘4c 2

1 13 1 iu os eee 14 fos

11 2 1 14 1 1 -- Abdomen VIII

id 1 Abdomen VII Co ST

Abdomen VI 0 1 1 ee

0 1 1 1 i i oe 6-10 7

eae 2 2 1-3 1 24 1

ee 1 3 2-4 > Paddle

ges 2 4 ee 1 B56. 3.

Reinert: Aedes (Rhinoskusea)

TABLE 2. Record of the branching of the setae on

the pupae of Aedes (Rhinoskusea) pillaii

Seta Range Mode

Cephalothorax

1 2-3 2

2 2-3 3

3 2-3 rae

& 2 2

5 2 2

= 1 1

7 2 2

8 2-5 3

9 2 2

Metanotal Plate

2-4 3

2 2

1-3 2

Abdomen I

19-34 20

1 1

Z 2

4-6 4.

3-4 3

1-3 2

2-4 3

1 ‘

(7 specimens)

Seta Range Mode

Abdomen I (Cont. )

10 1 1

11 1 1

Abdomen II

0 1 1

1 9220: 2

2 1 1

3 2-5 3

4 2-4 3

D 3-6 3

6 1-4 2

7 2-3 2

9 1 1

0 1 1

1 3-7 4

2 1 1

3 2-4 3

4 2-4 3

4) 2-9 4

6 1-3 2

7 2-3 2

8 2-3 3

Seta Range

Mode

Abdomen III (Cont. )

Abdomen IV

Abdomen V

1-4

48 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

TABLE 2 (Cont. )

‘Seta. Range Mode Seta Range Mode Seta Range Mode

Abdomen V (Cont. ) Abdomen VI (Cont. ) Abdomen VII (Cont. )

o 2-3 2 4 Ba4 3 5 123 2

4 2-4 3 5 2-4 3 6 3-5 4

5 2-4 3 6 2-6 3 7 1:2 1

6 2-8 3 7 1-2 1 8 258 2

7 1-4 3 8 122 2 9 2-5 3

8 2-8 2 9 1 1 10 2-3 2

9 i 1 10 12 1 11 1 1

10 1-2 1 11 1 1 14 1 1

11 1 1 14 1 1 Abdomen VIII

14 1 1 Abdomen VII 0 1-2 1

Abdomen VI 0 1 il 4 2 2

0 1 1 1 122 2 9 5-8 8

1 2-5 3 2 1 1 14 1 1

2 1 1 3 2-3 2 Paddle

3 1-2 2 4 1-2 2 1 3-5 4

Reinert: Aedes (Rhinoskusea)

TABLE 3. Record of the branching of the setae on

the pupae of Aedes (Rhinoskusea) wardi

Mode

Seta | Range

Cephalothorax

1 2-3 2

2 2-3 3

3 2-3 3

4 1-3 Z

5 2-3 2

6 1 1

7 2-4 3

8 2-5 4

9 2 2

Metanotal Plate

10 2-5 “

11 2 2

12 2-4 2

Abdomen I

1 18-42 26

2 1 1

3 1-2 2

4 3-6 5

5 2-4 4

6 1-2 2

7 1-3 2

9 1 1

(7 spe

cimens)

Seta Range Mode

Abdomen I (Cont. )

1 1

Abdomen II

1 1

14-26 16

1 1

2-3 3

2-4 3

2-5 4

2-4 2

1-3 2

1 1

Abdomen III

1 1

3-6 4

1-2 1

2-3 2

2-4 3

1-3 2

2-9 3

2-4 3

Seta Range

Mode

Abdomen III (Cont. )

1-2

Abdomen IV

2-4

1-2

2-6

2-3

2-4

1-3

2-4

TABLE 3 (Cont. )

Seta

Abdomen V (Cont. )

Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

Range Mode

2-4

2-3

2-4

1-2

Abdomen VI

1-3

1-2

Seta Range Mode

Abdomen VI (Cont. )

1-3

2-4

1-2

Abdomen VII

1-2

2-4

1-2

Seta Range

Mode

Abdomen VII (Cont. )

2-4

3-6

1-2

3-5

3-6

Abdomen VIII

1-2

2-4

4-8

1-2

Paddle

3-4

Reinert: Aedes (Rhinoskusea) o1

TABLE 4. Record of the branching of the setae on the

larvae of Aedes (Rhinoskusea) longirostris

(7 specimens)

‘Seta. Range Mode Seta Range Mode Seta Range Mode

Antenna Prothorax (Cont. ) Mesothorax (Cont. )

1 o-8 6 2 1 1 10 1 1

Head 3 1 1 11 1 1

0 1 1 4 1 1 12 1 1

1 1 1 9) 1 1 13 6-10 8

3 1 1 6 1 1 14 3-6 4

és 4-5 4 i; 2-3 2 Metathorax

9 o-7 6 8 1-2 2 1 1-2 1

6 2-3 2 9 1-2 1 2 2-3 2

7 10-14 13 10 1 1 3 2-4 3

8 2-3 2 11 2-3 3 A 2-3 3

2 2-3 2 12 : 1 5) 1 1

10 a8.14 2 14 2 2 of 1 1

11 o-7 D Mesothorax 7 6-8 7

12 2-3 3 1 1 i 8 2-6 4

13 2-3 2 2 1-2 2 ) 2-3 3

14 1 1 3 1 1 10 1 7

15 3-95 3 é 2 2 11 1 1

18 1 1 9) 1 1 12 1 1

6MP 1 1 6 4-6 5) 13 3-4 3

Prothorax é: 1 1 Abdomen I

0 5-8 7 8 6-9 6 1 1 1

1 1 1 9 o-7 6 2 si 1

Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

TABLE 4 (Cont. )

Seta

Range

Mode

Abdomen I (Cont. )

1-2

2-5

Abdomen II

Seta

Abdomen II (Cont. )

Range Mode

Abdomen III

Abdomen IV

Seta Range

Mode

Abdomen IV (Cont. )

1-2

Abdomen V

Reinert: Aedes (Rhinoskusea) D3

TABLE 4 (Cont. )

“Seta Range Mode Seta. Range Mode | Seta Range Mode

Abdomen V (Cont. ) Abdomen VI (Cont. ) Abdomen VIII (Cont. )

10 1 i 14 1 1 2 1 1

11 1-2 2 Abdomen VII 3 6-9 7

12 1-2 1 0 1 1 4 1 1

13 1 1 1 1-2 1 D 9-13 11

14 1 1 2 i 1 14 1 1

Abdomen VI 3 1-2 2 Abdomen X

0 1 1 “ 1 1 1 1-2 1

1 1 1 5 2 2 2 4-5 4

2 1 1 6 4-5 4 3 1 1

3 1 1 7 1 1 Siphon

4 2 2 8 4-6 4 1 3-5 9)

3) 1-2 1 9 1-4 f 2 1 1

6 2 2 10 2 2 3 1 1

7 1-2 2 11 1 1 4 1 1

8 1-3 2 12 1 1 6 1 1

9 1-2 1 13 1 1 7 1 1

10 1 1 14 i 1 8 1 1

11 2 2 Abdomen VIII 2 1 |

12 1 1 0 1 1 10 1 1

13 5-8 DB 1 3-5 4 11 1 1

12 1 1

54. Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

TABLE 5. Record of the branching of the setae on

the larvae of Aedes (Rhinoskusea) pillaii

(6 specimens)

‘Seta Range Mode Seta Range Mode Seta Range Mode

Antenna Prothorax (Cont. ) Mesothorax (Cont. )

1 5-7 5 2 1 1 10 1 1

Head 3 1 1 11 1 1

0 1 1 4 1 1 12 1 1

1 1 1 9) 1 a 13 0-10 6

3 1 1 6 1 1 14 4-7 6

4 2-4 3 7 2 2 Metathorax

5 5-7 6 8 1-4 2 1 1-2 1

6 2 2 9 1 1 2 1-2 1

7 10-16 12 10 1 1 3 2-3 3

8 2 2 11 1-2 2 4 2-4 3

9 2-3 2 12 1 1 5) 1 1

10 2-4 3 14 2 2 6 1 1

11 o-11 6 Mesothorax t 8-11 8

12 2-3 2 1 1 1 8 4-6 5)

13 2-4 3 2 1-2 1 9 2-3 2

14 1-2 1 3 1 1 10 1 1

15 2-4 2 4 2 2 11 1 1

18 1 1 O 1 1 12 1 1

6MP 1-2 1 6 4-6 9) 13 3-95 4.

Prothorax 7 t 1 Abdomen I

0 6-10 7 8 8-10 8 1 1 1

1 1 1 9 6-8 6 2 1 1

Reinert: Aedes (Rhinoskusea) 55

TABLE 5 (Cont. )

‘Seta Range Mode Seta Range Mode Seta Range Mode

Abdomen I (Cont. ) Abdomen II (Cont. ) Abdomen IV (Cont. )

3 2 2 12 1 1 3 1-2 1

4 4-6 4. 13 4-6 5) 4 1-3 2

D 1-3 3 14 1 1 5) 1 1

6 2-3 3 Abdomen III 6 2 2

7 1 1 0 1 1 i 2-4 2

s) 2-3 2 1 1 1 8 1 1

10 1 1 2 1 1 9 1 1

11 1-2 1 3 1-2 1 10 1 1

12 1 1 4 1-2 2 11 1-2 1

13 1 1 D 1-2 1 12 1 1

Abdomen II 6 2 2 13 1 1

0 1 1 7 2-5 3 14 | 1 1

1 1-2 1 8 1 1 Abdomen V

2 1 1 a 1 1 0 1 1

3 1-2 2 10 1 1 1 1-2 1

~ 3-5 4 11 1 1 2 1 1

5 1-2 1 12 1-2 1 3 1 1

6 3-4 3 13 1 1 4 2-3 3

7 2-3 2 14 1 1 5 1 1

8 1 1 Abdomen IV 6 2 2

9 1 1 0 1 1 7 2-4 3

10 1 1 1 1 1 8 1 1

11 4 1 2 1 1 g 1 1

56 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

TABLE 5 (Cont. )

‘Seta Range Mode Seta Range Mode Seta Range Mode

Abdomen V (Cont. ) Abdomen VI (Cont. ) Abdomen VIII (Cont. )

10 1 ] 14 1 1 2 1 1

11 1-2 1 Abdomen VII 3 6-9 7

12 1 1 0 1 1 4 1 1

13 1 1 1 1 1 D 8-11 9

14 1 1 2 1 1 14 1 1

Abdomen VI 3 2-4 2 Abdomen X

0 1 1 4 1 1 1 1 1

1 1 1 4) 1-2 2 2 4-6 9)

2 1 1 6 4-6 5 3 1 1

3 if 1 7 I 1 Siphon

4 2 2 8 3-5 4 1 4-6 5

5 1 1 9 2-3 2 2 1 1

6 Z 2 10 2 2 3 1 1

7 1 1 11 1 1 4 1 1

8 2-4 2 12 1 1 6 1 1

9 1 1 13 1 1 7 1 1

10 1 1 14 1 1 8 1-2 2

11 1-2 2 Abdomen VIII 9 1 1

12 1 1 0 1 1 10 1 1

13 4-6 6 1 3-4 4 11 1 1

12 1 1

Reinert: Aedes (Rhinoskusea) 7

TABLE 6. Record of the branching of the setae on

the larvae of Aedes (Rhinoskusea) wardi

(7 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

Antenna Prothorax (Cont. ) Mesothorax (Cont. )

1 0-6 5 2 1 1 10 1 1

Head 3 1 1 11 1 1

0 1 1 A 1 1 12 1 1

1 1 1 D 1 1 13 6-11 i

3 1 1 6 1 1 14 o-7 5

4 2-6 3 7 2 2 Metathorax

4) o-7 5 8 1-2 1 1 1-2 1

6 2 2 9 1-2 1 2 2 2

7 9-13 10 10 1 1 3 3-5 3

8 2 2 11 2-4 3 4 2-3 3

9 2-3 2 12 1 1 D 1 1

10 2-3 3 14 2 2 6 1 1

iA 3-6 6 Mesothorax 7 6-8 7

12 2 z 1 i 1 8 5-6 5

13 2-3 2 2 1-2 1 9 2-3 3

14 1 1 3 it 1 10 1 1

15 2-5 3 4 2 2 11 1 1

18 1 1 D 1 1 12 1 1

6MP 1-2 1 6 5-6 5 13 3-5 3

Prothorax 7 | 1 Abdomen I

0 6-10 9 8 6-7 6 1 1 1

1 1 1 9 o-6 6 2 1 1

08 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

TABLE 6 (Cont. )

‘Seta Range Mode Seta Range Mode Seta Range Mode

Abdomen I (Cont. ) Abdomen II (Cont. ) Abdomen IV (Cont. )

3 2 2 12 1-2 2 3 a 2

4 4-6 5 13 6-9 8 4 1-2 2

5 2-4 3 14 1 1 D 1-2 2

6 3 3 Abdomen III 6 2 2

t 1 1 0 1 1 7 3-5 4

9 2-3 3 1 1-2 2 8 1 1

10 1 1 2 1-2 1 9 1 1

11 2-3 2 3 1 1 10 1 1

12 1 1 4 2-3 2 11 2-3 2

13 1-2 2 9) 1-2 1 12 2-3 2

Abdomen II 6 2 2 13 1 1

0 1 1 7 2-4 4 14 1 1

1 1 1 8 1 1 Abdomen V

2 1 1 9 1 1 0 1 1

3 2-3 2 10 1 1 1 1-2 1

4 3-6 3 11 2 2 2 1 1

9) 1-2 2 12 2 2 3 1 1

6 3-4 3 13 1 1 4 3-4 4

7 2-4 3 14 1 1 9) 1-2 2

8 1 L Abdomen IV 6 2 2

9 1 1 0 1 1 7 2-3 3

10 1 1 1 1-2 2 8 1 1

11 1-2 1 2 1 1 9 1 1

Reinert: Aedes (Rhinoskusea) 59

TABLE 6 (Cont. )

Abdomen V (Cont. ) Abdomen VI (Cont. ) Abdomen VIII (Cont. )

10 1 1 14 1 1 2 1 1

11 2-3 2 Abdomen VII 3 7-8 8

12 i 1 0 1 1 4 1 1

13 1 1 1 1-2 2 5 10-13 12

14 1 1 2 1 1 14 1 1

Abdomen VI 3 2 2 Abdomen X

0 1 1 4 1 1 1 1 1

1 1-2 2 D 1-3 1 2 o-7 5)

2 1 1 6 4-6 5 3 1 1

3 1 1 7 1 1 Siphon

4 2 2 8 5-7 5 1 4-6 5

D 1-2 1 9 1-3 1 2 1 1

6 2 2 10 2-3 2 3 1 1

7 1 1 11 1 sf 4 i 1

8 2-4 3 12 1 1 6 1 1

9 1 1 13 1-2 1 7 1 1

10 1 1 14 1 1 8 1 1

11 2-4 2 Abdomen VIII 9 1 1

12 1 1 0 1 1 10 1 1

13 o-7 7 1 4-5 5 11 1 1

12 1 1

60 Contrib. Amer. Ent. Inst., vol. 13, no. 2, 1976

INDEX

Valid names are set in romantype. The italicized pages are those which

begin the primary treatment of the taxon. Numbers in parentheses refer to

the figures illustrating the species in question.

Aedes (genus) 1, 2, 5, 6, 7, 8, 14, longirostris 1, 2, 7, 8, 11, 12, 13,

17, 18, 20; 21, 26 4, 17,18, 20, 26°(4, 2, 3, 6;

Aedes (subgenus) 6 9, 12, 14)

Aedimorphus 6 Lorrainea 6

Bothaella 6 niveus 6

Cancraedes 6 Opifex 6

Christophersiomyia 6 Paraedes 6

Fiealbia &,. 2,.8, 12; 13 pita 1, 2, 7%, 8, t1,.12, 13, 14;

Finlaya 6 17, 20, 26 (1, 4, 7, 10, 14)

fuscus 6 Pseudoskusea 6

Geoskusea 6 Rhinoskusea 1, 2, 5, 6, 7, 14, 17,

Halaedes 6 18, 20, 21, 26

hitit 1, 8, 13s Skusea 1, 6, 8

Huaedes 6 Uranotaenia 1, 8, 13

Leptosomatomyia 6 Verrallina 6

Levua 6 wardid, 7, 11, 17, 18, 20,°21, 26;

@ 5 8 11, 18, 14)

MOSQUITO STUDIES (Diptera, Culicidae)

XXXII. A REVISION OF THE SCAPULARIS GROUP

OF AEDES (OCHLEROTATUS)’

J. Hal Arnell”

CONTENTS

INTRODUCTION ...

MATERIAL AND METHODS

SYSTEMATICS

Taxonomic History .

Taxonomic Characters .

Proposed Classification

Distribution .

Affinities. . .

MEDICAL IMPORTANCE . :

TAXONOMIC TREATMENT .

Scapularis Group. . .

Keys to Subgroups and Species.

Incomptus Subgroup :

1. Aedes (O.) connie.

Bogotanus Subgroup ,

2. Aedes (O.) beng mie ;

3. Aedes (O.) deficiens

Trivittatus Subgroup

4. Aedes (O.) pidctanittatuc.

5. Aedes (O.) meprai .

6. Aedes (O.) angustivittatus

7. Aedes (O.) trivittatus .

Tortilis Subgroup . .

8. Aedes (O.) ‘ones

9. Aedes (O.) auratus .

OMONINNDHN A HA BW bP

‘Contribution from project “Mosquitoes of Middle America’ supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478.

* Department of Biology, University of California, Los Angeles, CA 90024.

3 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

Infirmatus Subgroup . . AURORE RRO UN, Oa TE UG ei ae er

10. Aedes (O.) conse. AEG EASES Reape a te IPT nay ON ae or OA RGR

bis wearer oC oe ee Gece ee aS

Be ey ag

PS eee eee a ee ple on ee ee ae

De ee ee ee ei ee

Scapularis Subgroup. . Peay eerie se eae ae, aR MI cae ape Fg eg ge Ce

15. Aedes (O.) nhaconotus Sea eee Wee reat ttle ae gi eho magis is eam

Pee Ey Mes Ae a a Se a ee cue a cy ae

ee a ee a ae a ee ee wk Bae a

Crinifer Subgroup . . AUT aout ee wtee asa Tac TG DAUR va IWE vag Pane au raematta Peage < o

18. Aedes (O.) cnniler SRST SEHD ES Ue 6 SSN Flt clas CU SL a eg ieee

ee ee a a ee ee es

Obturbator Subgroup . . . AGU erate tr ab Nag acer Ga egret lec arc We cats age

20. Aedes (O.) Ce a ae a

Pectinatus Subgroup . . ay Nee AG ARAN Ga pte ae WS hae ere ban re NMR 5

21. Aedes (O.) pectinatus ee oe i eee ue A UR aes re ee

Thelcter Subgroup . . PS ae SEGRE TOLER /ES Teese h ar pe RCO N RIOR CS

22. Aedes (O.) thelcter CAD i ie ag TR aS ES pci ate ols A 2

Subgroup unknown . . Peas pokes AO cay Saar eer ara aga

23. Aedes (O.) sp., Surinam form SENG OG a Sa UN i tah ana ga RN ce)”

De eer te PSO ee Omi PON ok Oar ee ale ee re

REFERENCES CITED .. . PN Gua GE ame ce tae eee ee cee Gee

FIGURES. . SR SSRGT A ER TG er a aN oes ctabe Menge Ey aa Nave na Her Mae Seyeki

TABLE OF DISTRIBUTIONS. ee Ohare eae sey is cary cial. a age Neen

CONSPECTUS OF TAXONOMIC CHANGES aw neath Niston cae bere rah a epahanhtaagy Dep aiateay,

Pe Ee SL A ea Re ee ee ee ae eine ce Eee

INTRODUCTION

The species of the Scapularis Group constitute one of the main evolutionary lines

of the subgenus Ochlerotatus in the New World. Several phyletic lines of Neotropi-

cal ground pool breeding Aedes exhibit strong similarities in one or more stages, and

it is often difficult to define these evolutionary lines when considering less than all

stages together. Asa result, the classification of Neotropical Ochlerotatus is unques-

tionably confused and the present revision is an initial attempt to bring order to this

area of mosquito taxonomy. It is appropriate to begin with the species of the Scap-

ularis Group since scapularis itself is the type of the subgenus Ochlerotatus. It is

hoped that this study might stimulate further revisions of Ochlerotatus, even in the

New World a large and heterogeneous group.

Dyar (1918a:77) recognized the similarities among many of the species of the

Scapularis Group, but since Dyar (1928) and Edwards (1932) no attempt has been

made to define the limits.of the group or to refine its internal classification. As is

true with other Neotropical Ochlerotatus, the species of the Scapularis Group are

very similar in most stages, and often only careful comparison of females, male gen-

italia and larvae provides sufficient characters for adequate identification of species.

Because this has not always been done, the taxonomic literature is replete with mis-

interpretations of species, misidentifications and erroneous distributions. Due to

lack of material in some cases, I have been unable to follow the preferred practice

Arnell: Scapularis Group 3

of comparison of adequate material of all stages and several species interpretations

are provisional. Only collection and study of more material will confirm or refute

these interpretations.

Several species of the Scapularis Group have been implicated in disease transmis-

sion to man or his domestic animals and in view of the widespread distribution and

broad ecological tolerances of these species a thorough knowledge of their taxono-

my should aid in disease control.

I wish to thank John N. Belkin for reading and criticizing the manuscript and for

discussions on several aspects of the study; Thomas J. Zavortink for stimulative dis-

cussion of the study; Peter F. Mattingly for the loan of specimens from the British

Museum (Natural History); Miguel Garcia for the loan of specimens from the Insti-

tuto Nacional de Microbiologia, Buenos Aires; John F. Reinert for the loan of speci-

mens from his personal collection; L. T. Nielsen for the loan of specimens from the

University of Utah; Curtis W. Sabrosky and Ronald A. Ward for kindnesses extend-

ed during my visit to the U.S. National Museum and for arranging the loan and re-

turn of specimens from the USNM; William A. Powder and Sandra J. Heinemann

for the preparation of material; L. Margaret Kowalczyk and Nobuko Kitamura for

the preparation of the preliminary and final drawings; and Kay Metzner for typing

the rough draft and preparing the text copy for lithoprinting.

MATERIAL AND METHODS

The material for this study came primarily from collections made for the project

“Mosquitoes of Middle America” (Belkin, Schick et al., 1965) and is deposited at

the University of California, Los Angeles [UCLA]. Additional specimens were bor-

rowed from or examined at the U.S. National Museum of Natural History [USNM],

including the holotypes of angustivittatus, trivittatus, bracteatus, balteatus, pluto-

craticus, virginensis, condolescens, indolescens, obturbator and thelcter and the lec-

totypes of cuneatus, argentescens, traversus, inconspicuus, habanicus, auratus and

camposanus; the British Museum (Natural History) [BM], including the lectotypes

of tortilis and crinifer; the Instituto Nacional de Microbiologia, Buenos Aires, Ar-

gentina [INM], including the holotype of raymondi; the Department of Biology,

University of Utah [UTAH]; and John F. Reinert, Laurel, MD [JFR]. A total of

12,214 specimens was examined, 1787 males, 6280 females, 1911 pupae and 2236

larvae. Included in this total were 859 individual rearings of which 458 were larval

rearings, 331 were pupal rearings and 70 were incomplete.

The procedures used in this study were primarily those of comparative morpho-

logical taxonomy. The form of presentation and terminology and abbreviations

used in the descriptions follow Belkin (1962) with modifications (Knight, 1970;

Knight and Laffoon, 1970, 1971; Laffoon and Knight, 1971). Descriptions are

composites of all available specimens. In 4 instances only a single specimen was

available. Illustrations were prepared in general from a single specimen but were

corrected to show the modal condition of most available specimens. On the maps,

localities from which I examined specimens are indicated by solid symbols and rec-

ords from the literature by hatched symbols. Political subdivisions and locality

names in the distribution lists conform to the recommended usage in the Official

Standard Names Gazeteers of the United States Board on Geographic Names.

4 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

SYSTEMATICS

TAXONOMIC HISTORY. Culex scapularis was described by Rondani in 1848

and Ochlerotatus confirmatus, conspecific with scapularis, was described by Lynch

Arribalzaga in 1891. By 1910, 21 species now included in the Scapularis Group had

been named, although their close relationship had not been recognized. In the final

edition of his monograph on the Culicidae, Theobald (1910) placed the species

known to him in 3 genera, trivittatus in Pseudohowardina, scapularis in Leucomyia

and crinifer and inconspicuus (conspecific with trivittatus) in Culex. H. G. Dyar

was apparently the first to recognize the similarities among the species. In their

monograph, Howard, Dyar and Knab (1917:770-810) placed the species generally

together, and in a paper on natural mosquito groups based on male genitalic struc-

tures, Dyar (1918a:77) placed scapularis, infirmatus, euplocamus, tortilis, pluto-

craticus, trivittatus, cuneatus, condolescens, balteatus, bracteatus, angustivittatus

and obturbator, the latter 5 being unknown to him in the male, together under one

couplet in Group IV, Ochlerotatus, forms with a well developed claspette (harpago)

and basal and apical sidepiece lobes, in his key. Dyar (1920:103-106) later com-

bined his Groups IV and V (Ochlerotatus and Culicada) under the subgenus Heter-

onycha Lynch Arribalzaga (the name later restricted to Culex), and included all

forms with both basal and apical lobes. Under Heteronycha was the “‘Group scapu-

laris,’’ those forms he had placed together in his previous paper, this being the first

use of the Scapularis Group designation. In subsequent works, Dyar (1922a:51-60;

1924a:117-119; 1928:162-170) refined the subgeneric placement and his concept of

the species included in the group. In the most recent treatment of the Scapularis

Group, as such, Edwards (1932:137, 142-143) broadened the group to include also

dupreei (Coq., 1904) and related species, serratus (Theo., 1901) and related species,

euiris Dyar, 1922 and hortator Dyar & Knab, 1907.

Coquillett (1910:577) designated confirmatus, the second of 2 species included

by Lynch Arribalzaga (1891:146) in his genus Ochlerotatus, as the type of the ge-

nus. This nominal species is now considered to be a subjective synonym of Aedes

scapularis.

TAXONOMIC CHARACTERS. The Scapularis Group, including the subgroups

within it, is defined primarily on the basis of adult characters, especially those of the

female. The immature stages and the male genitalia display a mosaic of characters

which make phylogenetic relationships difficult to determine in most cases. Al-

though the male is much like the female except for sexual differences, the distribu-

tion of scales and patterns of scale colors which constitute the most obvious and

reliable taxonomic characters are more conspicuous and constant in the female

than in the male.

Female characters. The scales covering the scutum except for the usual bare

Spaces are narrow and curved and range in color from dark brown to white. The

color pattern formed by these scales is the most important taxonomic character in

the Scapularis Group not only in identifying species but also in defining the sub-

groups. The presence or absence of white scales on the apices of the femora, most

conspicuous when present on the ventral surface of the midfemur and hindfemur

often are significant in species recognition, and even though all species have baso-

lateral white scale patches on the abdominal terga, white scales often extend across

the dorsal surface at the base of the terga and their presence or absence is signifi-

cant. The Scapularis Subgroup is uniquely marked with white scales in a long stripe

on the anterior of the hindtibia, the white scales often continued to the basal tarsal

Arnell: Scapularis Group 5

segments. The development of palpal segment 4 is of minor importance and must be

observed on slide-mounted specimens.

Male genitalic characters. Considerable confusion has resulted in the past in 1 at-

tempts to separate the species of the Scapularis Group on the basis of the male geni-

talia. Emphasis has been placed on characters which are not constant, while several

reliable characters appear to have been overlooked. The differences in male genitalic

characters between many species are relatively minor, and I have found it impossible

to separate several of the 19 species which are known to me in the male, among

these being several common and widespread species.

Characters which I have found to be of value include sideniees length, the distri-

bution and development of setae on the tergal and sternomesal surfaces of the side-

piece, the development of the setae on the basal tergomesal lobe of the sidepiece,

the claspette and the spiniform of the clasper. Tergite IX, the phallosome and proc-

tiger have proven to be of no value. The apical tergomesal lobe of the sidepiece, a

character on which considerable importance has been placed in many studies of

Ochlerotatus, is undeveloped to well developed in the Scapularis Group and appears

to be of little phylogenetic significance. The taxonomic value of this structure is

further diminished by variation in orientation of the sidepiece in slide preparations.

The subapicotergal setae consist of an irregular, usually oblique, row of 2 to 8,

usually 4 to 6, very well developed setae on the middle of the tergal surface of the

sidepiece at about the level of the base of the apical tergomesal lobe. The setae

proximad of the subapicotergal setae are significantly weaker in development, and

may be separated from the subapicotergal setae by a bare space, and are either uni-

formly short or progressively longer distally. Immediately ventrad of the mesal

membranous surface of the sidepiece is a sclerotized area, often separated from the

sternal surface by a narrow membranous area, extending from the level of the basal

tergomesal lobe to about the base of the clasper, from which arise about 20 to 80

long to very long setae, the number and development of which are relatively con-

stant within a species. The basal tergomesal lobe is well developed, triangular,

though often folded ventrad in slide preparations thus appearing deceptively small.

About 10 to 30 setae arise from the lobe with the peripheral setae of the cluster

usually being slightly longer and with a single basal seta being much longer and

thicker than the remainder. This differentiated seta is thickened basad of the middle

and tapers gradually to a usually recurved apex. The size of this seta appears to be

constant for each species, although accurately assessing the thickness and length is

too difficult for this character to be of practical value. The claspette has a charac-

teristic development in many species and is important in species determination, al-

though to be of value it must be separated from the remainder of the genitalia and

mounted in a lateral view. The ratio of spiniform length to width is significant in a

few species. Measurements were made at a magnification of 860 diameters.

Pupal characters. The pupae of the Scapularis Group are remarkably uniform in

structure and for the most part are difficult or impossible to recognize to species. I

have not found any species or subgroup characters sufficiently reliable or of suffi-

cient magnitude to allow preparation of a key. However, there are several characters

that differ somewhat from species to species and since they are of some value they

are included in the species descriptions.

Larval characters. The thoracic and abdominal integument of all species of the

Scapularis Group is spiculose. There is very little intraspecific variation in spicule

density or length, and, although somewhat inconvenient to use, in several instances

this character is important in species diagnoses. Spicules were measured on slide-

6 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

mounted larvae or larval skins, under a magnification of 860 diameters. The rela-

tively broad spicule base tapers abruptly to a needle-like shaft and the ratio of

basal width to spicule length was measured using an ocular grid on spicules entirely

within the plane of the microscope field.

PROPOSED CLASSIFICATION. The present revision of the Scapularis Group

of Aedes (Ochlerotatus) treats 22 species and 2 unnamed forms. Of the 22 species,

8 are new and | has been resurrected from synonymy. In addition, 3 names, hemi-

surus, camposanus and rhyacophilus, have been reduced to the synonymy of scapu-

laris. Ten subgroups are recognized within the Scapularis Group in the present re-

vision. They are characterized primarily by unique or unusual features of the adult

female but often have distinctive features in other stages. The subgroups are sum-

marized below.

Incomptus Subgroup. Monotypic. Male and female scutum completely dark

brown scaled.

Bogotanus Subgroup. 2 species. Female scutum dark brown scaled with a nar-

row acrostichal line of white scales.

Trivittatus Subgroup. 4 species. Female scutum with narrow to broad dorso-

central lines of light scales extending the entire length of the scutum; pleural integ-

ument dark brown.

Tortilis Subgroup. 2 species. Female scutum with large anterior scutal scale

patch of golden to tan scales and very small subspiracular scale patch.

Infirmatus Subgroup. 5 species. Female scutum with broad anterior stripe of

silvery white scales.

Scapularis Subgroup. 3 species. Female scutum with large anterior hexagonal

patch of yellowish tan to white scales; anterior surface of hindtibia with white

scales.

Crinifer Subgroup. 2 species. Female scutum with narrow acrostichal and dorso-

central lines of tan to white scales or white and dark brown scales in random pat-

tern; distinctive male genitalia; larva with accessory dorsolateral setae on siphon.

Obturbator Subgroup. Monotypic. Scutum with pale tan scales except for dark

brown scales in narrow acrostichal line and on lateral scutal margin; pleural integu-

ments light brown; larval siphon with ventroapical prolongation.

Pectinatus Subgroup. Monotypic. Female scutum completely dark brown scaled;

male scutum with broad patch of white scales on posterior half; larva with 60-70

comb scales; pecten extending well beyond insertion of seta 1-S.

Thelcter Subgroup. Monotypic. Scutum yellowish white scaled with usual broad

— acrostichal line of tan to golden scales; sidepiece of male genitalia short and broad;

larval pecten extending beyond insertion of seta 1-S. .

DISTRIBUTION. The Scapularis Group has an extensive distribution throughout

the Americas from southern Canada to middle Argentina except for the far western

United States and extreme northwestern Mexico and the Pacific slope of the Andes

south of Peru (see figs. 1-4).

AFFINITIES. Due to lack of knowledge of much of the aedine fauna of the Neo-

tropics, the affinities between the Scapularis Group and other aedine phylads is dif-

ficult to determine. It is apparent, also, that before conclusions on relationships

among New World faunas can be made, an extensive knowledge of Old World

Ochlerotatus must be attained. The subgenus Ochlerotatus is often considered to

be primarily a northern group since it has undergone a tremendous species explo-

sion in the Holarctic; however, it is undoubtedly tropical in origin.

Aedes shannoni Vargas & Downs, 1950 from Central Mexico is possibly closely

Arnell: Scapularis Group 7

allied to the Scapularis Group. I have seen males and females of this species. The

male genitalia are similar to species of the Scapularis Group; however, the adult orna-

mentation differs in several respects, most notably in the presence of basal tarsal

bands of white scales, a character which may be of no phylogenetic significance. I

have seen no immatures of shannoni. For the present, I am not including it in the

Scapularis Group.

The morphology and ornamentation of the male and female of calumnior Belkin,

Heinemann & Page, 1970 from Jamaica and possibly the Cayman Islands is remark-

ably similar to obturbator, calumnior being virtually indistinguishable from obtur-

bator, although the larva of calumnior lacks integumental spiculation typical of the

Scapularis Group and the male genitalia are completely unlike any other Neotropical

Ochlerotatus. The remarkable similarity in the adults of the 2 species may be con-

vergence or may indicate hybridization between 2 distinct phyletic lines at some

time in the distant past, possibly during a period of geological instability in the Car-

ibbean area.

On the basis of the male genitalia, the Scapularis Group appears to be closely

allied to the Neotropical euiris Dyar, 1922, milleri Dyar, 1922, scutellalbum Boshell-

Manrique, 1939, and bejaranoi Martinez, Carcavallo & Prosen, 1960, species autoc-

thonous, primarily, to the higher elevations of the Andes, although significant differ-

ences are apparent in the adults and larvae. On the basis of distribution, a case may

be advanced for relationships between these species and the species of the Scapularis

Group, bogotanus, deficiens and pectinatus, apparently endemic to the Colombian

Andes, although it must be kept in mind that the Andes are a young mountain range

and its mosquito fauna is complex and relatively poorly known.

The species of the Serratus Group, which includes the Neotropical serratus

(Theo., 1901), nubilus (Theo., 1903), pertinax Grabham, 1906, eucephalaeus Dyar,

1918, oligopistus Dyar, 1918, hastatus Dyar, 1922 and aenigmaticus Cerqueira &

Costa, 1946, and the Nearctic dupreei (Coq., 1904), atlanticus Dyar & Knab, 1906

and tormentor Dyar & Knab, 1906, resemble the Scapularis Group closely in the

female and a close relationship is often advanced. The 2 groups are distinguishable

primarily by the absence of subspiracular scales in the Serratus Group, although

the individual species can be separated, in most cases, from the Scapularis Group

by other details of ornamentation. On the basis of the male genitalia, the Serratus

Group differs markedly from other New World Ochlerotatus, having an enlarged,

thumb-like apical tergomesal lobe and, more importantly, a basal tergomesal lobe

either partially detached from the sidepiece or attached to the sidepiece by a mem-

brane and more strongly attached to the claspette. Belkin, Heinemann and Page

(1970:160) suggest that this group might be placed in a separate subgenus, the

name Protoculex Felt, 1904 (type species, serratus) being available.

MEDICAL IMPORTANCE

The medical importance of the Scapularis Group is significant, since five species,

scapularis, trivittatus, infirmatus, thelcter and angustivittatus, have been implicated

in the transmission of disease agents. Of the five species, scapularis is by far the

most important, at least 15 viruses having been isolated from this species including

yellow fever and Venezuelan equine encephalitis (VEE) viruses and it also appears to

be a vector of Bancroftian filariasis. It should be pointed out, however, as Sudia,

Newhouse and Chappell (1969:600) state, a mosquito can be considered no more

8 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

than a potential vector of a disease organism unless 4 criteria are met: (1) isolation

of the disease producing agent from wild caught specimens, (2) demonstration of its

ability to become infected by feeding on an infected host, (3) demonstration of its

ability to transmit by bite, and (4) field evidence confirming association of the in-

fected mosquito with the vertebrate population in which the infection is occurring.

DeRodaniche and Galindo (1963:925) report the isolation of Ilheus virus from a

pool of 423 angustivittatus collected near Almirante, Panama in November, 1960,

and an isolation of VEE virus from angustivittatus in Colombia is reported by San-

martin, Mackenzie et al. (1973).

Aedes thelcter appears to be one of the more important potential vectors of VEE

in the localized areas where this species is common. Sudia and Newhouse (1975:3)

report numerous isolates of VEE virus from thelcter collected in the lower Rio

Grande valley of Texas and Mexico during the epidemic wave of VEE through this

area in 1971. During the study of this epidemic, thelcter was found to have the

highest infection rate for VEE virus among all species collected and tested (Sudia,

Newhouse et al., 1975:27).

Although the species does not appear to be particularly important in transmission,

several viruses have been isolated from infirmatus in the southeastern United States.

Lewis, Hammon et al. (1965) and Bond, Hammon et al. (1966) report a California

group virus similar to Trivittatus virus, later confirmed to be Trivittatus virus (Well-

ings, Lewis and Pierce, 1972), recovered frequently and predominantly from infirm-

atus in Florida, although human disease from this source was uncommon. A Trivit-

tatus virus-infirmatus cycle was later confirmed by Taylor, Lewis et al. (1971) in

Florida in a 1966-1967 study in which Keystone virus, another California group

virus, was also isolated from infirmatus. In a summary report on arboviral ecological

Studies in the same area of Florida, Wellings, Lewis and Pierce (1972) report Eastern

equine encephalitis, Keystone, Trivittatus and Tensaw virus isolations from infirma-

tus. One isolation of Trivittatus virus from infirmatus in southern Alabama is re-

ported by Sudia, Chamberlain and Coleman (1968).

Aedes trivittatus is a relatively common mosquito in the midwestern United

States, and virus isolations in several eastern and midwestern states show that several

viruses are not uncommonly found in this species. Trivittatis virus was originally

isolated from trivittatus in North Dakota. Kokernot, Hayes et al. (1969) report

Trivittatus virus from 5 of 24 pools (649 specimens) of trivittatus taken from man

and light trap collections in southern Illinois in 1967. A very high infection rate of

Jamestown Canyon serotype of California encephalitis virus is reported from (trivit-

tatus in Wisconsin by DeFoliart, Ainslow et al. (1969), and in a later paper, De

Foliart, Ainslow et al. (1972) report evidence that Trivittatus virus shows a high

specificity for trivittatus, but since trivittatus feeds on a variety of vertebrates there

is no indication of the relationship between the virus and its vertebrate hosts. Wong,

Dorsey et al. (1970), reporting on surveillance of encephalitis virus in Iowa state

that California group viruses appear as clinical cases in children every year and that

trivittatus is a major cause. This species was the second most abundant mosquito

taken in trapping collections from 1966 to 1968, and one isolate of Trivittatus virus

was made for every 250 pools tested. Wong, Rowley et al. (1973) report 2 isolations

of Western equine encephalitis virus and | isolation of Flanders virus, in addition to

29 isolations of Trivittatus virus from (¢trivittatus in lowa in 1969, 1970 and 1971.

Wills, Pidcoe et al. (1974) report isolates of Trivittatus virus from trivittatus in Penn-

sylvania. Over 26,000 mosquito specimens were collected in the state of Durango,

Mexico in June, 1972, as reported by Sudia, Fernandez et al. (1975) from which 1

Arnell: Scapularis Group 9

isolate of Trivittatus virus and 2 isolates of Bunyamwera virus were made. The mos-

quitoes were identified as angustivittatus, although it is much more likely that the

species was actually trivittatus. A summary of recent research into the activity of

California group arboviruses in the United States and Canada, which contains addi-

tional information on the role of trivittatus, infirmatus and thelcter, is presented by

Sudia, Newhouse et al. (1971).

Aedes scapularis, besides being the most common and widespread species of the

Scapularis Group, is also the most important in the transmission of disease. Rachou,

Lima et al. (1954, 1955), in conducting an epidemiological survey of a village in

southern Brazil for filariasis, concluded that scapularis was a principal vector of Ban-

croftian filariasis outdoors because of its abundance and demonstrated vector capa-

bility. Of 39 individuals captured and dissected, 1 was in infective and 2 in prein-

fective stages.

Aedes scapularis has been shown to be an efficient vector of yellow fever virus

under experimental conditions (Davis and Shannon, 1929:803; Whitman and An-

tunes, 1937:810-811; Shannon, Whitman and Franca, 1938:110). Soper, Penna et

al. (1933) opined that scapularis may well have been a vector of yellow fever during

an outbreak in Espirito Santo, Brazil, and Bugher, Boshell-Maurique et al. (1944) re-

port the possible implication of scapularis in yellow fever transmission in eastern

Colombia. Forattini (1961) has shown that scapularis is apparently becoming do-

mesticated in areas where it has had a long association with man, and since it is an

efficient vector of yellow fever virus, it has a high potential as a vector of urban

yellow fever.

Venezuelan equine encephalitis has been isolated from scapularis in the Amazon

region of Brazil (Causey, Causey et al., 1961:242), Cumana, Venezuela (Sellers,

Bergold et al., 1965), Veracruz, Mexico (Scherer, Dickerman et al., 1971), although

VEE apparently has multiple vectors in that region, and Ecuador (Gutierrez, Monath

et al. as reported by Sudia and Newhouse, 1975:12).

Many other arboviruses whose disease threat to man or his domestic animals is

either unknown or negligible have been isolated from scapularis. These include May-

aro virus from Trinidad (Aitken, 1957:231); Manzanilla virus from Trinidad (Ander-

son, Spence et al., 1960), although the virus was not transmitted by bite by infected

scapularis; Kairi virus from Trinidad (Anderson, Aitken et al., 1960; Causey, Causey

et al., 1961), with antibodies found in humans and several animal species; Caraparu

virus from Trinidad (Causey, Causey et al., 1961) isolated from a mixed pool of

scapularis, Aedes serratus, and Sabethes species; Cache Valley virus from Trinidad

(Downs, Spence et al., 1961); Melao virus, the original isolation of this California

group virus being from scapularis from Trinidad (Spence, Anderson et al., 1961);

a California group virus, TR9375, from Trinidad (Aitken, 1960; Causey, Causey

et al., 1961) and from Brazil (Causey, Causey et al., 1961); the Brazilian strain of

Ilheus virus, TR8349 virus, TR8900 virus, TR10076 virus, and TR20659 virus, all

from Trinidad (Aitken, 1960).

TAXONOMIC TREATMENT

Scapularis Group

FEMALES. Small to medium sized species; integument light brown to dark

brown; dark scales with slight iridescence; light scales exclusive of scutum white to

10 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

gray, on scutum silvery white to yellow or gold; pleural scale patches reduced.

Head: Integument usually dark brown. Eyes moderately separated above anten-

nae, interocular space of 1-3 ommatidial diameters, with narrow, curved scales.

Decumbent scales of vertex and occiput narrow and curved near midline and along

ocular border, broad and flat on dorsolateral, lateral, and ventral surfaces; white but

usually with small dark patch dorsolaterally. Erect scales moderately numerous to

numerous, usually confined to occiput but occasionally extending forward on ver-

tex; mostly white, occasionally dark dorsolaterally and rarely entirely dark. Inter-

ocular setae well developed, 1-3 pairs; ocular setae well developed, about 4-6 pairs

dorsally, 3-4 pairs laterally. Clypeus dark brown, moderately large, bare. Proboscis

long, about 1.25-1.60 of forefemur, entirely dark scaled or noticeably lighter ven-

trally. Palpus short, about 0.16-0.23 of proboscis; 3 or 4 segmented, segment 4

small to minute when present; entirely dark scaled. Antenna slightly shorter than

proboscis; pedicel with yellow integument, darker mesally, without scales or with

few dark scales mesally; flagellar segment 1 longer than segment 2, with few small

scales; flagellar segments 3-13 each slightly longer than preceding segment; segments

2-13 each with basal whorl of 6 moderate setae.

Thorax: Integument light brown to dark brown. Acrostichal setae well develop-

ed in complete row, partially developed and confined to anterior and posterior of

acrostichal line, or restricted to anterior promontory; dorsocentral setae well devel-

oped in complete row, or with anterior setae absent except on promontory; prescu-

tellar and supraalar setae numerous and well developed; humeral setae and posterior

fossal setae well developed or absent. Scutellum with about 6-15 strong setae on

midlobe and lateral lobes. Scutum completely covered with narrow, curved scales

except for inner dorsocentral and median and lateral prescutellar bare spaces; scales

entirely light, entirely dark, dark with large patches of lighter scales, or with con-

trasting longitudinal lines of light and dark scales. Scutellum with numerous narrow,

curved white and/or dark scales on median lobe, scales sparse or absent on lateral

lobes. Postnotum dark brown, bare. Paratergite bare. Apn not enlarged. Pleural

setae on apn, ppn, ppl, psp, stp, pra and upper mep, setae absent on ssp and lower

mep; ppn setae in single row near posterior margin; stp setae in single row from up-

per anterior margin of sclerite to near level of lower margin of meron. Apn with few

narrow, curved, white or dark scales, not closely appressed; ppn nearly covered with

closely appressed, narrow and curved to broad and flat scales, mostly dark but often

white posterioventrally; broad, white, flat, appressed scales in patches on ppl, pst,

ssp, stp below pra, stp caudad of ssp, stp cephalodorsad of midcoxa, mep from up-

per setae to ventral third of sclerite, and usually on psp; scales absent on pcx, hypo-

stigma and metameron.

Legs: Hindcoxa subequal to midcoxa, its base below upper margin of meron.

Legs relatively short, forefemur about 0.85-1.05 (rarely up to 1.25) of distance from

top of thorax to apex of midcoxa. Forecoxa with white or white and dark scales,

midcoxa and hindcoxa with white scales. Forefemur and midfemur generally dark

scaled on dorsal and anterior surfaces and white scaled on ventral and posterior sur-

faces, often white scaling expanded basally and restricted apically; hindfemur most-

ly white scaled with dorsal dark line beginning at about middle and broadening dis-

tally. White posterior knee spots present or absent, when present usually most con-

spicuous on mid and hind femora. Foretibia and midtibia dark scaled, usually with

faint to conspicuous light streak posteriorly. Hindtibia dark scaled, usually with

faint light streaks posteriorly, or, in Scapularis Subgroup, basal fourth to apical

sixth with conspicuous white stripe on anterior surface which occasionally nearly

Arnell: Scapularis Group 11

encircles segment. Tarsi dark scaled, usually with faint to conspicuous light streak

posteriorly on segments 1 to 2 or 3, with additional white streak on anterior surface

of segment 1 in Scapularis Subgroup. Claws of foreleg and midleg with acute subme-

dian tooth, or rarely with posterior claw of foreleg and midleg simple; claws of hind-

leg simple or with acute submedian tooth.

Wing: Scales entirely dark or with small patch of white scales at extreme base of

costa and remigium. Plume scales on R,, Rs, R243, R2, R3, Ra+s, and ventrally

on M,+2, M344, Cu distad of m-cu, and distally on 1A.

Haltere: Knob light or dark scaled.

Abdomen: Tergum I dark scaled dorsally, bare laterally. Terga II-VII with baso-

lateral white scale patches, remainder of terga dark scaled but often with narrow

basal white bands or basomedian white patches. Sterna mostly white scaled.

FEMALE GENITALIA. Segment VIII: Completely or nearly completely retract-

ed into segment VII. Tergum narrowed distally; on midline about 0.5-0.6 of tergum

VII; distal 0.85 with sparse, short setae becoming more dense distally and rarely a

few scales. Sternum about 1.05-1.30 of tergum VIII; broader distally; distal margin

broadly rounded and uniformly sclerotized, or slightly emarginate and more weakly

sclerotized on midline; with sparse, short setae and few longer setae over entire sur-

face, more numerous distally; usually few scales on lateral and distal margins. Ter-

gum IX: Notched distally, maximum length about 0.35-0.50 of tergum VIII; mod-

erately sclerotized; lobes with 3-8 weakly developed setae. Insula: Weakly sclerotiz-

ed; connected to lower vaginal lip; with a pair of setae on each side of midline.

Cercus: Long, about 0.75-1.05 of tergum VIII; apex rounded; setae numerous, the

longest moderately developed; scales absent. Postgenital Plate: Short and broad,

length 0.35-0.55 of tergum VIII; index about 1.35-1.75; apex slightly emarginate in

ventral aspect; 6-8 weakly developed setae and 2 moderately developed seta distally;

basal median longitudinal apodeme weakly developed. Upper Vaginal Lip: Moder-

ately sclerotized; vaginal sclerites moderate in size, weakly sclerotized. Lower Vagi-

nal Lip: Continuous with upper vaginal lip; weakly sclerotized. Basal portion of

spermathecal duct weakly to strongly sclerotized. Spermathecae 3, strongly sclero-

tized, spherical, 1 noticeably larger than others.

MALES. Generally similar to females except for sexual differences. Scutal scale

patterns not as distinct as in females. Head: Proboscis long and slender, length

about 1.35-1.55 of forefemur. Palpus long, about 1.00-1.20 of proboscis; 5 seg-

mented, segments 2 and 3 ankylosed and long, making up about 0.50-0.80 of pal-

pus; segment 4 short, making up about 0.10-0.20 of palpus; segment 5 short, making

up about 0.07-0.20 of palpus; segment 3 slightly elevated and slightly swollen near

apex with segments 4 and 5 slightly downturned, ventral surface of apex of seg-

ment 3 and all of segment 4 and often base of segment 5 densely covered with well

developed setae; or, segment 3 uniformly slender and straight to apex, segments 4

and 5 not downturned and segments 3 and 5 with few short setae near apex; palpus

entirely dark scaled. Antenna shorter than proboscis, about 0.65-0.90 of proboscis;

pedicel much enlarged, without scales; flagellum strongly plumose, segments 1-12

with numerous long setae directed dorsad and ventrad and fewer shorter setae dir-

ected mesad; segment 1 with few scales, segments 12 and 13 elongate, the two com-

bined subequal in length to total length of preceding 11 segments. Thorax: Scutal

scale pattern often not as distinct as in female, with light scales usually more exten-

sive. Legs: Relatively short, forefemur about 0.90-1.20 of distance from top of

thorax to apex of midcoxa. Anterior claw of foreleg moderately to considerably

enlarged, with blunt submedian tooth and with or without acute basal tooth; poster-

12 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

ior claw of foreleg moderately enlarged, with acute basal tooth; anterior claw of

midleg greatly enlarged, with blunt submedian tooth and with or without acute basal

tooth; posterior claw of midleg small or moderately enlarged, with acute subbasal

or basal tooth; claws of hindleg small, with acute median or submedian tooth. Ab-

domen: Distal segments and genitalia not bent ventrad. Terga with basolateral

white scale patches, dark scaled dorsally or with narrow basal white bands. Sterna

usually mostly white scaled.

MALE GENITALIA. Segment IX: Tergum well developed, middorsal portion

usually deeply emarginate on anterior margin with a weakly sclerotized, narrow to

moderately broad caudal bridge connecting the small, prominent tergal lobes; each

lobe with 1-10 moderately to strongly developed setae; sternum large, with 2-4 setae

distally. Sidepiece: Well developed, cylindrical but slightly curved inward, length

about 2.5-4.5 times median width; mesal surface membranous from base to apex;

scales usually restricted to lateral surface; tergal surface with few to numerous short

to moderately long setae, with several thickened and elongate setae at level of base

of apical lobe (subapicotergal setae); lateral and sternal surfaces with few thickened

and elongate setae; sternomesal margin with few to numerous moderately to well

developed setae extending from level of basal lobe to clasper; apical tergomesal lobe

poorly to well developed, with few small setae; basal tergomesal lobe attached to

sidepiece, with 1 specialized basal differentiated seta that is thickened near base,

bent beyond middle and usually hooked at apex, a cluster of about 10-30 undif-

ferentiated setae distally or posteriorly on lobe and rarely 1 enlarged seta on ventral

side of lobe which is stronger than undifferentiated setae but considerably weaker

than differentiated seta; lobe often collapsed or folded ventrad in slide preparations.

Claspette: Stem slender and elongate, curved dorsad, usually with single seta arising

from conspicuous tubercle on ventral surface near base; filament usually subequal

in length to stem, narrow at base, expanded near middle, with or without retrorse

process on outer margin and tapering to acute, often recurved apex. Clasper: Sim-

ple, long, slightly expanded before or near middle, with conspicuous spicules, most-

ly on inner surface, curved inwardly distally; spiniform long and slender. Phallo-

some: Aedeagus small, simple, pyriform to ovate, sclerotized dorsally with slight

apical emargination. Proctiger: Strongly developed; basolateral sclerotization

strongly developed, vertical, produced into lateral tergal lobe ventrad of lobe of ter-

gum IX; paraproct strongly sclerotized, broadly joined to basolateral sclerotization,

with a single, large, heavily sclerotized apical tooth and a short, strongly sclerotized

mesal process; cercal setae short, fine, 1-6.

PUPAE. Cephalothorax: Weakly to moderately brown pigmented, leg cases and

dorsum usually slightly darker. Setae 1-3-C moderately developed, subequal, usually

double (1-4b). Seta 6-C weakly developed, with 2-7 distal forks. Seta 7-C moder-

ately developed, with 2-8 distal forks. Seta 8-C arising anterior to level of base of

trumpet, subequal to and usually with more branches than 9-C, which arises con-

siderably posterior to level of base of trumpet. Setae 10-12-C subequal in length,

10-C 4-8b, 11-C single (1-2b), 12-C usually 3-5b. Trumpet: Uniformly pigmented,

light brown to dark brown, considerably darker than cephalothorax. Broadening

gradually from base to apex; pinna small. Reticulate sculpturing strong, tracheoid

sculpturing absent. Abdomen: Weakly to strongly pigmented, light yellowish

brown to dark brown, lighter posteriorly; usually with area of stronger pigmenta-

Armell: Scapularis Group 13

tion on median anterior portion of most terga. Seta 1-II strong, multiple, 6-20

(4-27) branched, closer to midline than float hair (1-I); seta 1-[V moderately de-

veloped, usually 2-3(1-6) branched; seta 1-V similar in development to 1-IV though

usually with fewer branches; seta 1-VII single to triple (1-5b). Seta 2-II variable in

position, mesad or laterad of seta 3-II; seta 2-III-VII single, variable in position, us-

ually immediately mesad of seta 1 of corresponding segment but often considerably

displaced mesad, or slightly laterad of seta 1 on segments VI and VII. Seta 3-Il

single or double (1-5b), subequal to or weaker than 5-II; seta 3-III moderately de-

veloped, usually single (single or double); seta 3-V single to triple (1-6b); seta 3-

VII usually 1-4 (1-5) branched. Seta 5-IV,V strongly developed, usually nearly as

long as or exceeding length of following tergum, usually double (1-4b); seta 5-VI

strongly developed, usually shorter than length of succeeding tergum, usually double

(1-3b); seta 5-VII weakly developed, shorter than seta 3-VII, usually 2-4 (1-7)

branched. Seta 6-I,II moderately long, fine; seta 6-III-VI moderately developed,

fine, usually single or double, rarely 3 or 4 branched; seta 6-VII 2-10(2-21) branch-

ed. Seta 9-I-VI minute, single; seta 9-VII moderately developed, usually double

(1-4b); seta 9-VIII strong, dendritic, usually with 6-8 main branches. Terminal seg-

ments: Male genital lobe about 0.9-1.3 of tergum VIII. Female cercus projecting

beyond genital lobe, strongly sclerotized externally. Paddle: Very lightly pig-

mented; midrib darker, not extending to apex; external buttress indistinct. Mar-

ginal spicules very poorly developed. Apex usually slightly emarginate. Index about

1.0-1.3. Paddle hair 1-P well developed, long, single.

LARVAE. Head: Broader than long, maximum width about 1.4-1.6 of length,

moderately pigmented, lighter in ocular area and darker posteriorly. Labial plate

narrowed anteriorly. Seta 1-C stout. Setae 4,6-C displaced caudad, not near la-

brum. Seta 4-C weakly developed, usually 2-5 branched, caudad of level of 6-C.

Seta 5-C usually single, occasionally 2-4 branched, more or less in line with 6-C and

laterad of 1-C. Seta 6-C usually single, occasionally 2-3 branched. Seta 7-C usually

4-7 (2-11) branched. Seta 15-C short, multiple, 2-6 branched. Mental plate usually

triangular with 13-16(10-18) teeth on each side of median tooth.

Antenna: Short, slender, uniformly weakly pigmented, with few spicules uni-

formly distributed over entire shaft. Seta 1-A moderately long, usually 2-5(1-9)

branched, inserted near middle of shaft.

Thorax: Epidermis and fat body without conspicuous pigmentation. Integument

with distinct spicules. Tubercles of setae 5-7-P separate. Seta 3-P single or multiple

(1-5b). Seta 7-P usually triple, occasionally double (1-5b). Seta 1-M single or multi-

ple (1-5b). Seta 11-M absent. 3

Abdomen: Seta 1-III moderately developed, usually 3-5(2-11) branched; seta

1-IV,V strongly developed, single or double, rarely 3-8 branched; seta 1-VII strongly

developed, usually double, rarely single or triple. Seta 3-[V,V moderately develop-

ed, usually single or double (1-8b). Seta 6-I single to triple, usually double; seta 6-II

usually double, rarely single; seta 6-III-VI usually single, rarely double. Seta 8-H

weakly developed, usually double (1-3b). Seta 13-II weakly developed, multiple

(5-25b); seta 13-III weakly developed and multiple or strongly developed, usually

single (1-4b); seta 13-IV,V strongly developed, usually single (1-4b).

Segment VIII: Comb scales moderate to small in size; fringed; about 15-85 in

number, in irregular double to triple row or triangular patch.

Siphon: Moderately pigmented, with darker basal band. Short to moderately

long, index about 1.9 to 3.1. Acus distinct, attached, moderate in size. Pecten

teeth 9-25, usually with basal denticles; in straight row, evenly spaced or with

14 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

distal teeth detached, and extending as far as distal 0.2 but usually extending only

slightly beyond middle of siphon. Seta I-S slightly distad of pecten, or within

pecten in thelcter and pectinatus; moderately developed, 6-8(4-13) branched. 2-3

pairs of accessory dorsolateral setal tufts (2a-S) present in crinifer.

Anal Segment: Saddle completely encircling segment; moderately pigmented,

with darker basal band; acus distinct, attached, moderate to small in size. Seta 1-X

moderately developed, usually single (1-4b). Seta 2-X 7-10(5-19) branched. Seta

3-X single. Ventral brush (4-X) strongly developed, with 7 to 9 pairs of setal tufts;

proximal setae usually arising from grid, occasionally arising from within or on mar-

gin of saddle. Anal gills varied in length, usually dorsal and ventral gills subequal.

DISCUSSION. The members of the Scapularis Group can be distinguished from

all other New World Aedes by the following characters: in the females the antenna

is shorter than the proboscis; the pleural scales are white, never silver; the paratergite

is bare; the subspiracular scale patch is present though often reduced; the hypostigial

scale patch is absent; the tarsal segments are without basal or apical light bands; and

the wing scales are all dark except for a few white scales occasionally at the base of

the costa. In the male genitalia, distinguishable from most Neotropical Aedes, the

lobes of tergite IX are present and give rise to several stout setae; the apical tergo-

mesal lobe of the sidepiece, when developed, is not thumblike and gives rise to no

large setae; the basal tergomesal lobe is well developed, with a differentiated seta,

and is completely attached to the sidepiece; the sternomesal margin of the sidepiece

usually has more than 20 long setae between the basal tergomesal lobe and the base

of the clasper; and the claspette stem has a single seta arising near the base. In the

pupae the trumpet is broadened from base to apex; seta 1-II is closer to the midline

than the float hair (1-I); seta 3-III is moderately developed, considerably shorter

than the length of the succeeding tergite; seta 5-II is stronger than or at least sub-

equal in development to seta 3-II; seta 5-IV,V is strong, subequal to or longer than

the length of the succeeding tergum and usually double; seta 5-VII is weaker than

seta 3-VII and usually less than a third the length of the succeeding tergum; the

caudolateral margin of segments III or IV and V lack conspicuous denticles; the

paddle is usually longer than broad, the marginal spicules are very poorly developed

and the apex is usually slightly emarginate. In the larvae the thoracic and abdomi-

nal integument is spiculose; seta 13-VI is short, multiple and subequal to 13-II; the

comb scales number 15 or more in an irregular double row or triangular patch; the

anal saddle is complete; and the anal gills are not tracheate.

It is difficult, if not impossible, to determine relationships among the subgroups

of the Scapularis Group and any attempts to infer relationships beyond the sub-

group level, at this time, would be pure speculation. The arrangement of subgroups

has been with what seem to be the more generalized forms, and, coincidentally the

more common and widespread species, in the middle and the more extremely dif-

ferentiated forms at either end. At a later time when more material, especially

individual and progeny rearings of Scapularis Group species, a more thorough

knowledge of the biogeography of other Neotropical aedine phylads and additional

information from paleogeology and paleobiology of the Caribbean, Central Ameri-

can and northern South American areas is assembled, further conclusions on the

phylogeny of the Scapularis Group may be drawn.

2(1)

DUz).

4(2).

5(4).

6(5).

7(6).

8(6)

Arnell: Scapularis Group 15

KEYS TO SUBGROUPS AND SPECIES

FEMALES

17. comitatus unknown

Scutum with large hexagonal patch or broad stripe of white to yellowish

tan scales on anterior two-thirds, patch or stripe extending well laterad

of dorsocentral line; posterior third of scutum often with narrow stripe

of light scales on acrostichal and dorsocentral lines which are continu-

ous with anterior scale patch .. . a

Scutum with scales unicolorous, or With narrow "te hd ‘pueinidinal

lines of contrasting scales extending entire length of scutum, or with

light and dark scales in pattern suggesting light longitudinal lines on

PUNE COP i he Les ta rn al nin ei Ue vue cb at ee

Hindtibia with conspicuous white stripe on anterior surface, occasional-

ly nearly encircling tibia, and continued on basal tarsal segments; abdo-

men usually with indistinct pale median longitudinal stripe on posterior

segments; proboscis pale ventrally (fig. 35) (Scapularis Subgroup) . .3

Hindtibia entirely dark; abdomen entirely dark dorsally or with narrow

basal segmental white bands; proboscis entirely dark . . . ... .4

Anterior scutal scale patch mostly tan, often yellow to white posteriorly

ee . . . . L5. phaeonotus

Anterior scutal scale batch plac etn leew to tan on lateral margin

CF BG) Oe Bag ome Oe TD RI” ae ca EES

Anterior scutal scale patch golden to tan a 21) (Tortilis Subgroup)

8. tortilis, 9. auratus

Agtenor scutal seale patch entirely white. or - white divided by narrow acro-

stichal line of tan scales (iniirmatus Subgroup) |... oe ee ek wD

Anterior scutal white scale patch divided by narrow acrostichal line of tan

scales (fie, 32) 1. 420; , TON Tee MNOS A 8s Th ros

Anterior scutal scale patch calirely white sy CN ye 5 a A gr eR OE

Knee spots of white scales present posteriorly on apices of femora, most

CONSPICUOUS. Of MEI and NING Femoral eee one Wee Ae A

Knee suits iol White GCales ADeCTE cic ll oa ya tla ta i sia ae

Palpus apparently 3 segmented; distribution Central America and northern

South America. . . . . Il. euplocamus

Palpus 4 segmented, segment 4 relatively large: distribution northern Ar-

senting and southeri Bolivia 6 ek 6k ae Bel, a ee petersani

Abdominal terga with median basal white patches or bands on most seg-

Mmenie 5% «a lianes a oe LO condolescens

Abdominal terga entirely dark dorsally eee naka onion ik ae a

9(1).

10(9).

11¢10).

12(9).

13(12).

14(13).

15(13).

16(15).

17(16).

Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

Scutum uniformly dark brown scaled, or, if slightly lighter scales present

these restricted to posterior dorsocentral line . . . pe ges al)

Scutum with narrow to broad longitudinal lines of contrasting scales, or

with scales unicolorous and light, or scales mixed light and dark in

more or Jés8 fandom pattem i225 se ik ee eee A ace en dd

Knee spots of white scales present on all femora; erect scales of head num-

erous, elongate, extending forward on vertex, mostly dark (Pectinatus

Subgroup) . . . . . . .21, pectinatus

Knee spots of white RCales absent: erect geales of head few, short, restrict-

ed to occiput, mixed light and dark. se a ee eee

Decumbent scales of vertex and occiput entirely white; pleural integu-

ment light brown. . . . . . « « 24. Colombian form

Occipital dark scale patch prominent: pleural integument dark brown (fig.

5) (ncomptus Suberoup) |. .... «2. ns se wee ny LL incomptus

Scutal scales entirely yellowish white except for usual broad acrostichal

line of light brown to golden scales; basal white scale patches of ab-

dominal terga broader on midline, often forming median longitudinal

white stripe (figs. 46,47) rote Seca Roce af aud. Peleter

Scutal scales notasabove .. . pedoes fib aed seeds

Scutum with single dark narrow acrostichal line of white scales, remainder

of scutum dark brown scaled... 4b we ee ee ye IA

Bem nOlde anOve 2 6S ek 0 ee kee wt a

Posterior dorsocentral setae well developed; abdominal sterna with dark

scales in longitudinal median line (fig. 8) (Bogotanus Subgroup) .

ae 2. bogotanus, 3. potions

Dorsocential setae not developed except on anterior promontory; abdom-

inal sterna entirely dark scaled or with dark scales only in distal seg-

mental Dads: g430 6 a ee we a i 28 Sarinam form

Scutum with pale tan scales except for dark brown scales in narrow ac-

rostichal line and usually on lateral scutal margin; pleural ine

tan to light brown (figs. 41,42) (Obturbator Subgroup).. 20. obturbator

Scutal scales not as above; pleural integument dark brown . . . . . 16

Scutum with 2 narrow to broad lines of white to yellow scales in dorso-

central area, these lines continuous and distinct or somewhat irregular

(Trivittatus Subgroup). . . re

Scutum with broad median line of white: or aco scales usually separated

into 3 distinct lines by brown scales laterad of midline, or scales mixed

light and dark in more or less random pattern suggesting 3 light lines on

dark backeround(Crintier Suberoup) ....:. 2.2 ey a fe es 20

Knee spots of white scales present at apices of femora, most conspicuous

Ol Tid aid Rif TemGr. Ce a eas So ES

Wiee BHOLS Of Wiiite Scales absent a a DD

18(17).

19(17).

20(16).

2(1).

3(2).

Arnell: Scapularis Group 17

Dorsocentral line of white or yellow scales complete, unbroken from an-

terior promontory to scutellum and usually expanded laterad in supra-

war ater Clies, a yey. CO 7, SIVIIATUS

Dorsocentral line of white scales ey incomipters: broken at level of

prescutal suture and usually doubled posteriorly, as posterior dorso-

central line and supraalar line (fig. 13). . . . . . . 4. atactavittatus

Dorsocentral line of yellow to white scales uniformly narrow along entire

length, not expanded or divided in supraalar area (fig. 15) . :

: 6. Gres tateas

Dorsocentral line of white séales expanded laterad in antealar and supra-

alar areas, with elongate patch of dark scales laterad of posterior dorso-

Conia IMeCqIe O2). 3. oo oO ee a ee

Scutum with broad median line of white or yellow scales usually separ-

ated into 3 distinct lines by brown scales laterad of midline (figs. 38,

Sy Ye ee LO Crnier

Scutum with white to yellow scales on n background of dark baw scales

in somewhat random pattern but suggesting 3 longitudinal lines (fig.

ee a eee Bch ae

MALE GENITALIA

(3. deficiens, 5. meprai, 13. raymondi unknown)

Sidepiece short and broad, length about 2.5 times median width; basal

tergomesal lobe with about 25-35 setae, differentiated seta only slight-

ly larger than largest of undifferentiated setae (fig.47) . .22. thelcter

Sidepiece length at least 3.5 times median width; basal tergomesal lobe

usually with fewer than 25 setae, differentiated seta much larger than

Upditteren tated sola, es re ee ee 2

Claspette filament considerably longer than stem, straight and uniformly

narrow on basal two-thirds, then slightly broadened with small retrorse

process; spiniform long, moe about 20 times greatest width (fig. 41) .

: ‘ 20. obturbator

Claspette fikanent subequal in ieneth to ‘Stem. wicca dorsad at base,

broadened near or before middle; spiniform shorter, length less than

tS times Srestest With a a

Sternomesal margin of sidepiece with fewer than 30 relatively short setae;

differentiated seta of basal tergomesal lobe straight distad of middle;

apical tergomesal lobe of sidepiece not one (fig. 44) .

; Zi, fecinains

Sternomesal margin ‘of sidepiece with more than 30 setae; differentiated

seta of basal tergomesal lobe curved distad of middle; apical tergomesal

1ODC aC least moderately developed 6.0. he ee

4(3).

5(4).

6(5).

7(5).

8(7.)

9(8).

10(9).

Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

Claspette stem narrow, length about 10 times median width, curved dor-

sad on basal two-thirds and slightly recurved ventrad on distal third;

filament expanded just distad of retrorse process to about 3-4 times

basal width; apical tergomesal lobe very well developed; basal tergo-

mesal lobe with 20-30 setae (figs. 39,41). . 18. crinifer, 19. synchytus

Claspette stem length about 6 times median width, broadly curved dor-

sad; filament expanded just distad of retrorse process to about 1.5-2

times basal width; apical tergomesal lobe moderately to well developed;

basal tergomesal lobe usually with fewer than 25 setae . . ... .5

Claspette filament without retrorse process, or with retrorse process mi-

nute and without spicules basad of process. . . . 6

Claspette filament with large retrorse process and soyly averil elongate

RICiiles Ochad Ol PPOCESR Ci are ee a ee eee ee

Basal tergomesal lobe of sidepiece with conspicuous enlarged ventral seta,

weaker than differentiated seta but stronger than undifferentiated setae

(He S20 a el : . . . 17. comitatus

Basal tergomesal lobe of Sidepicce Wahoo srtoneedd ventral seta (figs. 33,

SO) 8 Oa a oS. pphgeono ms, 1 6. scapularis

Sternomesal margin of sidepiece with about 75 (65-85) well developed se-

tae between basal tergomesal lobe and clasper (fig. 30) . .14. infirmatus

Sternomesal margin of sidepiece with fewer than 60 setae between tergo-

ee Pe BS ae hee a ees iret nc antwriete FOS

Sternomesal margin of sidepiece with about 30 setae between basal tergo-

mesal lobe and clasper (fig. 13) . . . . 4. atactavittatus

Sternomesal margin of sidepiece with 40 to 50 or 60 sciae between basal

fersomesal toue ad Clasper os ene ee eS eee eG Y

Tergal surface of sidepiece with distinct bare space between subapicoter-

gal setae and much shorter, more proximal tergal setae; about 25-30

setae on basal tergomesal lobe (fig. 24) .... . . . 9. auratus

Tergal surface of sidepiece without bare space between subapicotergal se-

tae and more proximal tergal setae; setae of basal tergomesal lobe fewer

fan 25. say rower Mian OO a a, 2 ee 10

Claspette stem expanded into calyciform apex; spiniform short, length 7

to 10 times greatest width; subapicotergal setae 5 or 6 in oblique row

(Tie, US) a : . 6. angustivittatus

Claspette stem not expanded distally: spiniform length at least 10 times

greatest width; subapicotergal setae usually fewer than 5 (figs. 6,9,18,

212A 2ST).

I. incomptus, yy) bogotanus, 7. trivittatus, 8. tortilis, 10. condolescens,

11. euplocamus, 12. patersoni

Arnell: Scapularis Group 19

LARVAE

(5. meprai, 10. condolescens, 13. raymondi, 15. comitatus, 19. synchytus unknown)

4(2).

5(4).

6(5).

7(4).

8(7).

9(8).

10(9).

11(10).

Siphon with 2 or 3 pairs of accessory dorsolateral setae (2a-S) subequal

in development to seta 1-S (fig. 40). . . . ep dk, tO, CHMLLEF

Siphon without accessory dorsolateral setae (2a-S) . Ae NT ayo Oia

Pecten teeth extending well beyond insertion of setal-S ..... .3

Pecten teeth ending before or near insertion ofsetal-S . ..... .4

Comb scales about 60-70, in triangular patch; seta 1-VII single, subequal

to siphon length (fig. 45). . . . . . 21. pectinatus

Comb scales about 16-20, in ‘reoular Baubie: row; eats 1-VII 3 or 4 (2-5)

branched, short, less than 0.5 of siphon length (fig. 48) .... 22. thelcter

Seta 6-I,I] single . . Li uiAce cde iehive tune MAEM dl Be ee

Sarl dante wig ee heres ea eae

Siphon with acute ventroapical prolongation subequal in length to ventro-

lateral valve; ventral brush with 7 pairs of setal tufts; anal saddle with

short, acute spicules on dorsocaudal margin (fig. 43) . .20. obturbator

Siphon without ventroapical prolongation; ventral brush with 8 or 9 pairs

of setal tufts; anal saddle without spicules on dorsocaudal margin . .6

Ventral brush with 9 pairs of setal tufts; seta 13-III well developed, dou-

ble, subequla to 13-IV,V; setae

double or triple (fig. 12) . . . . . J. deficiens

Ventral brush with 8 pairs of setal ‘tufts; seta 13 iI small, yale ole sub-

equal to 13-II, setae 1,13-IV,V well eee ie single (fig. 23)

ARS ARC Ar gece AU Seam Ayia ay AIG 9. auratus

Seta 5-C 2-4 branched, seta 6-C single; ventral brush with 8 or 9 pairs of

setal tufts (fig. 10) . . . . 2. bogotanus

Seta 5,6-C single; ventral brush with: 8 pairs of setal tufts ite ee

Seta 13-III small, multiple, subequal to 13-LII (fig. 22) . . . .8. tortilis

Seta 13-III well developed, single, subean TO AAV oe hes ake he es tet

Seta 6-I usually triple, seta 6-II-VI usually Pe (fig. 14) . ia

4 Wg atactavittatus

Seta art Sonal, double. seta 6-III-VI Taal single RON eran eeeere seer 7.)

Comb scales with well-differentiated median spinule, at least 2.5 to 3

times length of submedian spinules (fig.31) . . . . .14. infirmatus

Spinules of comb scales subequal, or median spinule less than 2 times

TONEUN OF BUDIIOO IAT BITC R i gi ie i aa aaa el a Me i led

Spicules of integument very sparse and short, length less than twice basal

Clam ter (Ui 7 Yk ius’: wlssiisw jhe ted plc’ anlBaws Mle: a MANA. dak a

20 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

Spicules of integument numerous and conspicuous, long, length about 4-

i) ames Dasal digimeter 3 5 2

12(11). Seta 3-P usually at least double (2-5b), very see see a ake

Seta 3-P single, never double . . . . ie eee ele

13(12). Seta 1-III 6-8 branched (fig. 34) . . ...., . . . J5. phaeonotus

Seta I-III usually triple (1-5b) (fig.37) . . . . . . . . 16. scapularis

1412). Seta 1-1V,V usually double; very rarely single. . . ..... .. IS

seta 1-TV,V usually single, rarely double..° . 20 2. ee

15(14). Comb scales without enlarged median spinule, all spinules subequal; spic-

ules of integument long, length about 6-10 times basal diameter (fig.

16) . . .6. angustivittatus

Comb scales with median spinule mnleiued, slightly 1 onger and more prom-

inent than lateral spinules; spicules of integument moderately long,

length about 4-5 times basal diameter (fig. 19) . . . . 7. trivittatus

16(14). Seta 6-III single; spicules of integument moderately long, length about

4-6 times basal diameter (fig. 26). . .... . . . Il. euplocamus

Seta 6-III usually double; spicules of integument ay length about 8-12

times basal Giameter (igs 2S). ek . . . 12. patersoni

INCOMPTUS SUBGROUP

I. Aedes (O.) incomptus Arnell, sp. n.

Figs. 1,5,6,7

TYPES. Holotype female with associated pupal skin (PA 379-110), 6.5 km W of summit of

Cerro Mali, Serrania del Darien (approx. 29 km NE Pucro), Darien, Panama, elev. 1500 m, larva

taken from water hole of collared peccaries in marginal cloud forest, 8 June 1963, A. Quinonez

[USNM]. Allotype male with associated pupal skin and genitalia slide (PA 379-146), same data

as holotype [USNM]. Paratypes: 28 pM (PA 379-101,102,104,109,113-115,120,121,125-127,

130,132,133,141,142,144,145,149,151-153,157), 23 pF (PA 379-103,111,116,117,119,122,124,

128,129,134,135,137,138-140,143,147,148,150,155,156,158,162), 3 p, 36 1, 4 L, same data as

holotype [UCLA, BM, GML, FH].

FEMALE (figs. 5,6). Wing: 3.15 mm. Proboscis: 2.05 mm. Forefemur: 1.60

mm. Abdomen: 2.70 mm. Head: Occipital dorsolateral dark scale patch promi-

nent. Erect scales mostly gray or dark. Palpus 4 segmented. Thorax: Scutum

covered entirely with dark brown scales, or with few light brown scales in narrow

posterior dorsocentral line and surrounding prescutellar bare space. Acrostichal

setae usually absent, anterior dorsocentral and posterior fossal setae present. Scu-

tellum with dark brown scales on lobes. Legs: Forecoxa with extensive area of

dark scales anteriorly. Knee spots of white scales absent posteriorly on apices of

femora. Tibiae and basal tarsal segments of foreleg and midleg with light streak

posteriorly, hindtibia and hindtarsus entirely dark scaled. Claws of hindleg with

submedian tooth. Wing: Scales entirely dark. Abdomen: Terga entirely dark

scaled dorsally.

Arnell: Scapularis Group 21

MALE (figs. 5,6). Wing: 3.15 mm. Proboscis: 2.05 mm. Forefemur: 1.55 mm.

Abdomen: 2.75 mm. Thorax: Scutum mostly dark scaled except for lighter scales

along posterior dorsocentral line and in supraalar area, areas of lighter scales often

indistinct. Legs: Anterior claw of foreleg and midleg considerably enlarged, with

blunt submedian tooth and acute basal tooth.

MALE GENITALIA (fig. 6). Segment IX: Anterior middorsal emargination of

tergum broadly conical and relatively shallow, caudal bridge connecting tergal lobes

broad; tergal lobes with 4-7 setae. Sidepiece: Length about 3.5-4.5 times median

width; sternomesal margin with 50-60 moderate to long setae; apical tergomesal

lobe well developed; basal tergomesal lobe well developed, triangular with anterior

side shorter, lobe often collapsed or folded ventrad in slide preparations, with about

20 setae. Claspette: Stem long, uniformly narrow, broadly curved dorsad, basal

seta relatively small; filament slightly shorter than stem, narrow at base and broad-

ened considerably near middle with large retrorse process and 5-10 elongate spicules

basad of retrorse process extending to about basal 0.1 of filament, and tapered grad-

ually to recurved tip. Clasper: Sharply curved inward distally; spiniform length

about 12-14 times its greatest width. Phallosome: Aedeagus ovate to slightly

pyriform.

PUPA (fig. 6). Abdomen: 2.55 mm. Trumpet: 0.40 mm. Paddle: 0.75 mm.

Cephalothorax: Weakly pigmented, very slightly darker dorsally. Seta 8-C with at

least 9(9-14) branches; seta 9-C 4, rarely 3 or 5 branched. Abdomen: Weakly pig-

mented, very slightly darker anteriorly. Seta 1-IV 5(4-6) branched. Seta 5-IV-V

subequal in length to length of succeeding tergum, 5-VI shorter than length of suc-

ceeding tergum, all double (5-IV rarely triple). Seta 6-III,IV usually 4(2-5) branch-

ed; seta 6-V trjple, occasionally 4 branched. Paddle: Seta 1-P usually forked near

base.

LARVA (fig. 7). Head: 0.80 mm. Siphon: 0.75 mm. Anal Saddle: 0.35 mm.

Thorax: Spicules of integument sparse, inconspicuous, length about 2 times basal

diameter. Seta 3-P single. Abdomen: Seta 6-I,II double; 6-III-VI single. Seta 13-III

strongly developed, single. Segment VIII: Comb scales moderate in size, evenly

fringed, about 22-28 in number, in irregular double to triple row. Siphon: Index

2.3-2.5. Pecten with teeth evenly spaced, extending to middle of siphon. Seta 1-S

inserted slightly distad of distal pecten tooth. Anal Segment: Ventral brush (4-X)

with 8 pairs of setal tufts.

_ SYSTEMATICS. Aedes incomptus can be separated from the other species of the

Scapularis Group by the following characters: in the female (1) the scutum is nearly

completely dark brown scaled with occasional lighter brown scales on the posterior

dorsocentral line, (2) there are no Knee spots of white scales on the apices of the

femora, and (3) the occipital dark scale patch is prominent; in the male genitalia (1)

the sternomesal margin of the sidepiece has 50 to 60 long setae between the basal

tergomesal lobe and the clasper, (2) there is no bare space between the subapicoter-

gal setae and the more proximal shorter setae of the sidepiece, (3) the claspette stem

is broadly curved dorsad and not expanded distally, and (4) the claspette filament is

shorter than the stem, has a large retrorse process and several elongate spicules basad

of the process with the most proximal spicules near the basal 0.1 of the filament; in

the larva (1) the spicules of the thoracic and abdominal integument are sparse and

short, length about 2 times basal diameter, (2) seta 13-III is well developed, single,

subequal to 13-IV,V, (3) the siphon index is less than 2.5, and (4) the comb scales

are evenly fringed.

Aedes incomptus shows no close affinities to any other Scapularis Group species.

22 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

It appears to be, in all stages, one of the more generalized members of the group.

The female shows no unusual characters other than the completely dark scaled

scutum, the male genitalia are indistinguishable from several other species of the

group, and the larva is distinguished mainly by the minute development of the tho-

racic and abdominal integumental spicules. Known only from a single locality in

Panama, incomptus may be the remnants of a developmental line close to the an-

cestral stock that gave rise to the Scapularis Group.

BIONOMICS. Nothing is known of the bionomics of incomptus other than that

the type series was collected in a marginal cloud forest from a small ground pool

used as a water hole by collared peccaries. The pool contained larvae of Psorophora

(J.) sp and Culex (Mel.) sp near jubifer, was in deep shade and contained no vege-

tation.

DISTRIBUTION (fig. 1). Aedes incomptus is known only from a single locality

in the Serrania del Darien, Darien Province, Panama. Material examined: 149 speci-

mens; 29 males, 24 females, 56 pupae, 40 larvae; 53 individual rearings (pupal).

PANAMA. Darien: Cerro Mali, 6.5 km W of summit, type series, see above.

BOGOTANUS SUBGROUP

DISCUSSION. The Bogotanus Subgroup is defined by the female scutal orna-

mentation of dark brown scales divided by a narrow acrostichal line of white scales.

The females of the 2 species in the subgroup, bogotanus and deficiens, are appar-

ently indistinguishable, deficiens is unknown in the male, and the larvae of the 2

species are well marked and easily distinguished.

The subgroup is known from only a few collections in the immediate vicinity of

Bogota, Colombia.

2. Aedes (O.) bogotanus Arnell, sp. n.

Figs. 1,8,9,10

TYPES. Holotype female with associated larval and pupal skins (COB 96-11), Ogamora, nr.

Soacha, Cundinamarca, Colombia, elev. 2400 m, larva from volcanic rockhole, 28 Nov 1965,

Ochoa [USNM]. Allotype male with associated larval and pupal skins and genitalia slide (COB

93-11), “‘Las Acacias,” nr. Usaquen, Cundinamarca, Colombia, elev. 2640 m, larva from small

ground pool in garden, 8 Nov 1965, E. Osorno [USNM]. Paratypes: 3 lpM (COB 93-10,12,13),

1 lpF (COB 93-14), same data as allotype; 2 lpF (COB 96-10,12), 3 L (COB 96), same data as

holotype [UCLA, BM].

FEMALE (figs. 8,9). Wing: 4.55 mm. Proboscis: 2.85 mm. Forefemur: 2.05

mm. Abdomen: 3.30 mm. Proboscis and legs relatively long. Head: Occipital

dorsolateral dark scale patch conspicuous. Proboscis relatively long, about 1.35-

1.45 of forefemur. Antenna about 0.75 of proboscis. Palpus apparently 3 seg-

mented. Thorax: Scutum with narrow acrostichal line of white scales from anterior

promontory to scutellum, surrounding prescutellar bare space; remainder of scutum

with dark brown scales except for few white scales above paratergite and on anter-

ior humeral margin. Legs: Legs relatively long, forefemur about 1.20-1.25 of dis-

tance from top of thorax to apex of midcoxa. Knee spots of white scales absent on

posterior of apices of femora. ‘Tibiae and basal tarsal segments with white streak

posteriorly. Claws of hindleg with submedian tooth. Wing: Scales entirely dark,

Arnell: Scapularis Group 23

or, occasionally with small patch of white scales at extreme base of costa. Abdo-

men: Terga apparently entirely dark scaled dorsally. Sternum with dark scales in

distinct, longitudinal median line.

MALE (figs. 8,9). Wing: 4.80 mm. Proboscis: 3.50 mm. Forefemur: 2.20 mm.

Abdomen: 3.95 mm. Head: Proboscis long, subequal in length to palpus. Thorax:

Scutum with broad acrostichal stripe of white scales, somewhat widened posteriorly;

remainder of scutum with dark scales. Legs: Anterior claw of foreleg and midleg

considerably enlarged, with blunt submedian tooth and acute basal tooth.

MALE GENITALIA (fig. 9). Segment IX: Anterior middorsal emargination of

tergum shallow and relatively narrow, caudal bridge connecting tergal lobes broad;

tergal lobes about as broad as high, with 5-8 setae. Sidepiece: Length about 4.0-

4.5 times median width, tergal surface with setae sparse and uniformly relatively

short and cluster of 5-8 subapicotergal setae; sternomesal margin with about 50-60

long setae, most dense at level of base of apical tergomesal lobe; apical tergomesal

lobe very well developed; basal tergomesal lobe well developed, triangular with an-

terior side shorter and setae arising from posterior side, differentiated seta large,

about 20 undifferentiated setae of 2 distinct sizes, long near periphery of cluster

and much shorter in center of cluster, 4 or 5 long setae usually projecting distad

from posterior side of lobe. Claspette: Stem long, uniformly narrow and broadly

curved dorsad; filament narrow at base, expanded near basal third with large retrorse

process and several elongate spicules basad of retrorse process, and tapered to re-

curved tip. Clasper: Narrow, only slightly broadened near middle; spiniform length

about 12-15 times its greatest width. Phallosome: Aedeagus ovate to slightly

pyriform.

PUPA (fig. 9). Abdomen: 2.85 mm. Trumpet: 0.45 mm. Paddle: 0.95 mm.

Cephalothorax: Weakly pigmented, slightly darker dorsally. Abdomen: Weakly

pigmented, only slightly darker anteriorly, but strongly pigmented at base of seg-

ment III. Seta 1-VII single. Seta 5-IV,V longer than length of succeeding tergum,

both always single; seta 5-VI subequal in length to length of succeeding tergum,

single, very rarely double. Paddle: Apex broadly rounded or slightly pointed,

very rarely emarginate.

LARVA (fig. 10). Head: 1.10mm. Siphon: 1.20 mm. Anal Saddle: 0.45 mm.

Head: Seta 5-C 4(3-4) branched. Thorax: Spicules of integument dense, conspicu-

ous, length about 6-8 times basal diameter. Seta 3-P double (2-3b). Seta 7-P usually

triple (3-5b). Abdomen: Seta 1-III 7-9(5-11) branched; seta 1-[V double (2-3

branched); seta 1-VII triple. Seta 6-I,II double; seta 6-III-VI single. Seta 13-III

strongly developed, double; seta 13-IV double; seta 13-V triple (2-3b). Segment

VIII: Comb scales moderate in size, with slightly differentiated median spinule, 19-

24 in number, in irregular double to triple row. Siphon: Index about 2.6-3.1. Pec-

ten with teeth evenly spaced, extending to middle of siphon. Seta 1-S inserted dis-

tad of pecten, 9(7-12) branched. Anal Segment: Ventral brush (4-X) with 8 or 9

pairs of setal tufts.

SYSTEMATICS. Aedes bogotanus can be separated from the other species of

the Scapularis Group by the following characters: in the female, indistinguishable

from deficiens, the scutum has a narrow acrostichal line of white scales with the re-

mainder of the scutum covered with dark brown scales, and posterior dorsocentral

setae are well developed; in the male genitalia, generally indistinguishable from triv-

ittatus, tortilis, condolescens, euplocamus and patersoni, (1) the sidepiece length is

4 to 4.5 times median width, (2) the claspette filament is subequal in length to the

stem and has a large retrorse process with several elongate spicules basad of the

24 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

process, the spicules seldom extending basad of the basal 0.2 of the filament, (3)

_ the sternomesal margin of the sidepiece has 50 to 60 long setae between the basal

tergomesal lobe and the clasper, (4) the claspette stem is broadly curved dorsad and

not expanded distally, (5) the apical tergomesal lobe of the sidepiece is moderately

to well developed, and (6) the spiniform length is about 12 to 15 times its greatest

width; in the larva (1) seta 5-C is 2-4 branched, (2) seta 6-C is single, (3) seta 6-I,II is

double, and (4) the ventral brush has 8 or 9 pairs of setal tufts.

The apical tergomesal lobe of the sidepiece of the male genitalia is generally larger

in bogotanus than in the species from which I have considered it indistinguishable,

but this character is inconstant in slide preparations and too subjective to be of

value. This species appears to be closely related to deficiens, this relationship being

discussed further under deficiens below.

The name bogotanus is a manuscript name that W. H. W. Komp placed on a num-

ber of specimens of this species that he collected near Bogota in May and June 1942.

BIONOMICS. Aedes bogotanus is known only from the vicinity of Bogota, Col-

ombia. Larvae have been collected on 2 occasions, once from a small, temporary

ground pool in a garden, the pool containing abundant vegetation and larvae of

Aedes (O.) sp near euiris, and once from a volcanic rockhole in a pasture, with

water temporary and without vegetation, and containing larvae of A. (O.) sp near

euiris and Uranotaenia sp. Several collections made by W. H. W. Komp and his

associates in 1942 were adults, mostly males, taken sweeping.

DISTRIBUTION. Aedes bogotanus is known only from the vicinity of Bogota,

Cundinamarca, Colombia. Material examined: 89 specimens; 42 males, 28 females,

8 pupae, 11 larvae; 8 individual rearings (larval).

COLOMBIA. Cundinamarca: Bogota, 28 Feb 1922, F. Miller, 1 M [USNM]. Bogota, 1 Mar

1937 (COR 529), 6 F [UCLA]; same data (COR 530), 4 F [UCLA]. Bogota, 24 May 1942, W.

Komp (KO 200-12, KO 200-15, COK 67), 19 M, 7 F [UCLA]; same data, 2 M, 1 F [USNM].

Bogota, June 1942, W. Komp (KO 200-7), 11 M, 1 F [UCLA]; same data, 1 F [USNM]. Bogo-

ta, May 1944, W. Komp (COK 68), 1 F [UCLA]. Fontibon, 19 May 1942, W. Komp (KO 200-

14, COK 15), 5 M, 2 F [UCLA]. Soacha, type series, see above. Usaquen, type series, see above.

3. Aedes (O.) deficiens Arnell, sp. n.

Figs. 1,11,12

TYPES. Holotype female with associated larval and pupal skins (COB 92-10), Hacienda “‘Con-

jera,” 3 km from Suba, Cundinamarca, Colombia, elev. 2410 m, larva taken from ground pool in

pasture, 7 Nov 1965, Morales, Ochoa and Pardo [USNM]. Paratypes: 2 |pF (COB 92-11,12),

same data as holotype [UCLA].

FEMALE (fig. 11). Wing: 3.55 mm. Proboscis: 2.20 mm. Forefemur: 1.70

mm. Abdomen: 2.75 mm. Apparently indistinguishable from bogotanus.

MALE. Unknown.

PUPA (fig. 11). Abdomen: 3.05 mm. Trumpet: 0.50 mm. Paddle: 0.80 mm.

Cephalothorax: Very weakly pigmented, only slightly darker dorsally. Abdomen:

Weakly pigmented, very slightly darker anteriorly, strongly pigmented at base of seg-

ment III. Seta 1-VII double (rarely single or triple). Seta 5-IV,V subequal in length

to length of succeeding tergum, double on segment IV, single on segment V; seta 5-

Ilf shorter than length of succeeding tergum, single (single or double). Seta 6-IV

triple. Seta 14-VIII relatively long and conspicuous, subequal to seta 11-VII, single

to triple. Paddle: Apex slightly emarginate.

Arnell: Scapularis Group 25

LARVA (fig. 12). Head: 1.00 mm. Siphon: 0.85 mm. Anal Saddle: 0.35 mm.

Head: Seta 5-C double. Seta 6-C single. Thorax: Spicules of integument moder-

ately dense, length about 6-8 times basal diameter. Seta 3-P double (2-4b). Seta

7-P triple. Abdomen: Seta 6-I-VI single. Seta 13-III strongly developed, double

(1-2b); seta 13-IV double; seta 13-V triple. Segment VIII: Comb scales small in

size, evenly fringed, about 42-56, in triangular patch. Siphon: Index about 2.25-

2.40. Pecten with teeth evenly spaced, extending to middle of siphon. Seta 1-S

inserted distad of pecten, 9(7-9) branched. Anal Segment: Ventral brush (4-X)

with 9 pairs of setal tufts.

SYSTEMATICS. Aedes deficiens can be separated from the other species of the

Scapularis Group by the following characters: in the female, indistinguishable from

bogotanus, the scutum has a narrow acrostichal line of white scales and the remain-

der of the scutum covered with dark brown scales; in the pupa seta 14-VIII is rela-

tively well developed, subequal to 11-VII and single to triple; in the larva (1) seta

6,I,II is single, (2) the ventral brush has 9 pairs of setal tufts, and (3) seta 13-III is

well developed and subequal to 13-IV,V.

This species shows no characters in the female to distinguish it from bogotanus,

although the 3 females known of deficiens are depauperate and in extremely poor

condition. Distinguishing characters may be found when better specimens are exam-

ined. The species can be distinguished from bogotanus in the pupa and is very well

marked in the larva. It is probably closely related to bogotanus; however exact

placement must await discovery of the male and examination of additional females.

Although known only from a single collection, deficiens, as well as bogotanus, may

have a more extensive distribution than is now known, since the aedine fauna of the

higher elevations of the Andes is relatively poorly known.

BIONOMICS. Aedes deficiens has been collected only once, from a small, semi-

permanent ground pool in a pasture, the pool containing abundant herbaceous veg-

etation. Larvae of Aedes (O.) milleri were also present.

DISTRIBUTION. Aedes deficiens is known only from a single locality near Bo-

gota, Colombia. Material examined: 9 specimens; 3 females, 3 pupae, 3 larvae; 3 in-

dividual rearings (larval).

COLOMBIA. Cundinamarca: Suba, type series, see above.

TRIVITTATUS SUBGROUP

FEMALES. Head: Occipital dorsolateral dark scale patch conspicuous to nearly

absent. Palpus 3 segmented. Thorax: Scutum with dorsocentral line of white or

yellow scales extending entire length of scutum, this line continuous and distinct or

somewhat diffuse; white or yellow scales mesally on anterior promontory, surround-

ing prescutellar bare space, and often above paratergite; remainder of scutum with

golden brown to dark brown scales. Acrostichal, anterior dorsocentral, and poster-

ior fossal setae absent. Legs: Knee spots of white scales present or absent posterior-

ly on apices of femora. Tibiae and basal tarsal segments with white streak posterior-

ly; hindtibia and hindtarsus dark scaled anteriorly. Claws of hindleg with subme-

dian tooth. Wing: Scales entirely dark. Abdomen: Terga II-VII with large baso-

laetral white scale patches; entirely dark scaled dorsally, with basal white bands, or

basomedian white patches.

MALES. Thorax: Scutal scale pattern generally similar to female although longi-

tudinal lines of light scales usually more extensive laterally; occasionally scutum al-

26 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

most entirely light scaled. Legs: Anterior claw of fore leg considerably enlarged,

with blunt submedian and acute basal tooth, posterior claw of foreleg considerably

enlarged, with acute subbasal tooth; anterior claw of midleg with blunt submedian

and acute basal tooth, posterior claw of midleg small or moderately enlarged, with

acute subbasal tooth. Abdomen: Terga with or without basal bands of white scales.

DISCUSSION. The Trivittatus Subgroup is defined by the female scutal orna-

mentation of 2 narrow to broad dorsocentral lines of light scales extending the en-

tire length of the scutum. The pleural integument is dark brown in color, in con-

trast to the light brown integument of obturbator, which has a somewhat similar

scutal scale pattern.

The subgroup consists of 4 species, atactavittatus, meprai, angustivittatus, and

trivittatus, with an extensive distribution from southern Canada through Central

America and western South America to northern Argentina (see fig. 2). The 2 domi-

nant species of the subgroup are trivittatus in Canada, the United States except for

the Pacific Coast states and Mexico as far south as Oaxaca, and angustivittatus, ex-

tending from central Mexico to Venezuela, Colombia and Ecuador. There is a zone

of hybridization between these 2 species in southern Mexico, discussed below under

angustivittatus. Aedes atactavittatus has a restricted distribution in Campeche, Mex-

ico, and meprai is known from southern Ecuador, southern Bolivia and northern

Argentina and may possibly have a continuous distribution from Ecuador to Argen-

tina. The distribution pattern for the subgroup is similar to that of the Infirmatus

Subgroup, except that the latter is represented in the Antilles. This similarity may

indicate a parallel evolutionary history, especially in Central and South America.

4. Aedes (O.) atactavittatus Arnell, sp. n.

Figs. 2,13,14

TYPES. Holotype female with associated larval and pupal skins (MEX 610-16), E outskirts

Lerma, Campeche, Mexico, elev. 6 m, larva taken from rockhole in back yard of dwelling, 5 Aug

1970, D. Schroeder [USNM]. Allotype male with associated larval and pupal skins and genitalia

slide (MEX 610-13), same data as holotype [USNM]. Paratypes: 3 F (MEX 592), same locality

as holotype, 21 July 1970, K. and D. Schroeder; 1 IpF (MEX 602-11), 5 L (MEX 602), same data

as holotype; 4 F (MEX 607), same data as holotype, 1 L (MEX 608), same data as holotype; 2

IpM (MEX 609-12,15), 2 lpF (MEX 609-13,14), 4 L (MEX 609), same data as holotype; 1 lpM

(MEX 610-10,11,12,15), 1 IlpF (MEX 610-14), 22 L (MEX 610), same data as holotype [UCLA,

BM, ISET].

FEMALE (fig. 13). Wing: 2.90 mm. Proboscis: 1.90 mm. Forefemur: 1.65

mm. Abdomen: 2.40 mm. Thorax: Dorsocentral line of white scales usually in-

complete, broadened laterally and terminating at level of prescutal suture and usual-

ly double posteriorly as posterior dorsocentral line and supraalar line; white scales

present above paratergite. Legs: Knee spots of white scales present posteriorly on

apices of femora. Abdomen: Terga II-VI or VII with complete basal bands of

white scales.

MALE (fig. 13). Wing: 2.10 mm. Proboscis: 1.60 mm. Forefemur: 1.20 mm.

Abdomen: 1.90 mm. Thorax: Scutal scale pattern indistinct; white scales more ex-

tensive than in female, occasionally covering entire scutum. Legs: Posterior claw

of midleg small, with acute subbasal tooth. Abdomen: Terga with basal bands of

white scales.

MALE GENITALIA (fig. 13). Segment IX: Anterior middorsal emargination of

Arnell: Scapularis Group poe

tergum broad and shallow, caudal bridge connecting tergal lobes broad; tergal lobes

about as broad as high, with 3-6 setae. Sidepiece: Length 4 times median width;

tergal surface with few short setae and usually 3 subapicotergal setae; sternomesal

margin with about 30 moderately long setae; apical tergomesal lobe well developed;

basal tergomesal lobe moderately well developed, triangular with posterior side

longer than anterior side, often folded ventrad in slide preparations, with about 20

setae. Claspette: Stem broadly curved dorsad, slightly broadened distally; fila-

ment subequal in length to stem, narrow at base, abruptly expanded near middle

with sharp retrorse process and several elongate spicules basad of retrorse process,

and tapered to recurved tip. Clasper: Spiniform length about 10-14 times its

greatest width. Phallosome: Aedeagus pyriform to ovate. :

PUPA (fig. 13). Abdomen: 2.35 mm. Trumpet: 0.35 mm. Paddle: 0.70 mm.

Cephalothorax: Weakly pigmented, dorsum only slightly darker. Abdomen: Weak-

ly pigmented, only slightly darker anteriorly. Seta 3-V-VII single (3-VI,VII rarely

double). Seta 5-IV-VI slightly longer than length of succeeding tergum, 5-IV usual-

ly single (single or double), 5-V,VI always single. —

LARVA (fig. 14). Head: 0.80 mm. Siphon: 0.70 mm. Anal Saddle: 0.30 mm.

Thorax: Spicules of integument moderately numerous and conspicuous, length

about 6-10 times basal diameter. Seta 3-P single. Seta 7-P triple (2-3b). Abdomen:

Seta 1-IV usually 4(3-4) branched; seta 1-V triple (2-3b). Seta 6-I usually triple (2-

3b); seta 6-II-VI usually double (6-II 2-4b, 6-VI 1-2b). Seta 13-III strongly develop-

ed, triple (3-5b), subequal to seta 13-IV,V which are also triple (2-4b). Segment

VIII: Comb scales moderate in size, usually with moderately differentiated median

spinule, about 23(15-28) in number, in irregular double to triple row. Siphon: In-

dex about 2.25-2.40. Pecten with teeth evenly spaced, extending slightly beyond

middle of siphon. Seta 1-S inserted slightly distad of pecten. Anal Segment: Ven-

tral brush (4-X) with 7 pairs of setal tufts, most proximal tuft often arising from

saddle margin.

SYSTEMATICS. Aedes atactavittatus can be separated from the other species of

the Scapularis Group by the following characters: in the female (1) the dorsocentral

line of light scales on the scutum is broken for a short distance at the level of the

prescutal suture and double posterior to the break and (2) knee spots of white scales

are present on the apices of the femora; in the male genitalia (1) the claspette fila-

ment has a large retrorse process and several elongate spicules basad of the process

and (2) the sternomesal margin of the sidepiece has about 30 setae between the

basal tergomesal lobe and the clasper; in the larva seta 6-I is usually triple and 6-II-

VI usually double.

This species is well marked in the larva, similar to trivittatus but with discrete

scutal markings in the female, and more or less intermediate between angustivit-

tatus and ¢rivittatus in the male genitalia. It appears to be a relict of an early iso-

late of the Trivittatus Subgroup.

BIONOMICS. Aedes atactavittatus was collected at the type locality, an acacia-

thorn scrub forest, in the larval stage from 4 small rockholes, all in the yards of

dwellings. The water in each was temporary and either contained grassy vegetation

or the surface was covered with sticks and leaves. Other species associated with

_ atactavittatus were Psorophora (J.) cyanescens, Culex (C.) coronator, C. (C.) corni-

ger, C. (C.) nigripalpus and C. (Mel.) pilosus. The Hopelchen larval collection was

from a large, semipermanent, roadside ground pool containing grassy vegetation and

floating sticks and bark, and larvae of scapularis, C. pilosus and P. (J.) ferox. Biting

females were captured at the type locality on 2 occasions, once at 1600 hours (MEX

28 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

607) and once at 1700 hours (MEX 592).

DISTRIBUTION (fig. 2). Aedes atactavittatus is known only from two localities

near Campeche, Campeche, Mexico. Material examined: 82 specimens; 9 males, 12

females, 14 pupae, 47 larvae; 14 individual rearings (larval).

MEXICO. Campeche: Hopelchen (20 km W on hwy 180), 80 m, 4 Aug 1970, K. and D.

Schroeder (MEX 600), 2 IpM (10,11), 1 L [UCLA]. Lerma, type series, see above.

5. Aedes (O.) meprai Martinez and Prosen

Figs. 2,32

1953. Aedes (Ochlerotatus) meprai Martinez and Prosen, 1953:1-2. TYPE: Holotype female,

Reserva Nac. ‘Finca El Rey,’ Dep. Anta, Salta, Argentina, Nov 1952 [BA]. Synonymy

with angustivittatus by Ronderos and Garcia (1963b:38).

Aedes (Ochlerotatus) meprai of Stone, Knight and Starcke (1959:148); Forattini (1965:364).

Aedes (Ochlerotatus) angustivittatus of Cerqueira (1943:31); Ronderos and Garcia (1963a:37);

Forattini (1963:348, in part); Stone (1967:207).

Aedes angustivittatus of Del Ponte, Castro and Garcia (1951:236).

FEMALE (fig. 32). Wing: 4.40 mm. Proboscis: 2.55 mm. Forefemur: 2.40

mm. Abdomen: 2.90 mm. Thorax: Dorsocentral line of white scales expanded

laterally in antealar and supraalar areas, with elongate patch of dark scales laterad

of posterior dorsocentral line; white scales present above paratergite. Legs: Knee

spots of white scales absent posteriorly on apices of femora. Abdomen: Terga en-

tirely dark scaled dorsally.

MALE, PUPA, LARVA. Unknown.

SYSTEMATICS. I have seen meprai only in the female, and although similar to

trivittatus and angustivittatus, it can be distinguished from (trivittatus by the absence

of knee spots of white scales on the apices of the femora and from angustivittatus

by the light dorsocentral line of white scales on the scutum which is expanded later-

ally in the antealar and supraalar areas.

This species has been known only from the type locality; however, I have seen

females from 2 localities in southern Bolivia, an unknown locality probably in Peru

and from a single locality in southern Ecuador, all indistinguishable from the para-

type I have seen from Argentina. This indicates a possible continuous distribution

from Ecuador through Peru and Bolivia to northern Argentina, probably at moder-

ate to high elevations. The Ecuador collection is from 2000 meters elevation, and,

in contrast, the closely related angustivittatus has been collected in Ecuador only at

low elevations on the Pacific Coast.

BIONOMICS. The type series of meprai was collected biting man. No more is

known of the bionomics of this species.

DISTRIBUTION (fig. 2). Aedes meprai is known from northern Argentina,

southern Bolivia, southern Ecuador and Peru. Specimens examined: 15 specimens;

15 females.

ARGENTINA. Salta: Reserva [Parque] Nacional Finca el Rey, Nov 1952, Martinez, 1 F

[INM].

BOLIVIA. Chuquisaca: Muyapampa, 3 Apr 1944, Carr (3147), 1 F [USNM]. Santa Cruz:

Camira, 23 Apr 1944, Carr (3148), 3 F [USNM].

ECUADOR. Loja: Vilcabamba (20 km S Loja), 2000 m, 17 May 1974, D. Pletsch (ECU 186),

7 F [UCLA].

PERU. [?]Otapampa, 3 Aug 1927, C. Townsend, 3 F [USNM].

Arnell: Scapularis Group 29

6. Aedes (O.) angustivittatus Dyar and Knab

Figs. 2,15,16

1907. Aedes angustivittatus Dyar and Knab, 1907:9. TYPE: Holotype female, Limon, Limon,

Costa Rica, 28 Sept 1905, F. Knab [USNM, 10140; see Stone and Knight, 1956:214].

Synonymized with trivittatus by Dyar (1922a:52); resurrected from synonymy with

trivittatus by Dyar (1924a:117).

1908. Aedes cuneatus Dyar and Knab, 1908:54-55. TYPE: Lectotype female (422.19) with

associated 2 larval and 1 pupal skins, Cordoba, Veracruz, Mexico, 20 Jan 1908, F. Knab

[USNM, 11964, selection of Stone and Knight, 1956:216]. Considered a subspecies of

trivittatus by Dyar (1922a:54); synonymy with angustivittatus by Dyar (1924a:117).

1908. Aedes argentescens Dyar and Knab, 1908:55. TYPE: Lectotype male (441.4), Cordoba,

Veracruz, Mexico, 20 Jan or 8 Apr 1908, F. Knab [USNM, 11965; selection of Stone

and Knight, 1956:214]. Synonymy with cuneatus by Howard, Dyar and Knab (1917:

770).

19725. We traversus Dyar, 1925b:215-216. TYPE: Lectotype female, sand bar in Zulia Riv-

er, Zulia, Venezuela, L. H. Dunn [USNM, 28480; selection of Stone and Knight, 1956:

226|. Synonymy with angustivittatus by Dyar (1928:163).

Aedes (Ochlerotatus) angustivittatus of Dyar (1925a:144; 1928:163-164); Edwards (1932:142);

Lane (1939:107; 1953:670-671); Arnett (1949:237); Vargas (1956:22); Stone, Knight and

Starcke (1959: 138); Forattini (1965:347-349, in part); Stone (1967:207, in part).

Aedes (Heteronycha) angustivittatus of Dyar (1920:105).

Aedes angustivittatus of Howard, Dyar and Knab (1917:776-778); Dyar (1924a:117-118); Kumm,

Komp and Ruiz (1940:399); Kumm and Zuniga (1942:404).

Aedes (Ochlerotatus) angustivittatus angustivittatus of Martini (1935:4 ).

Aedes (Heteronycha) cuneatus of Dyar (1920:105).

Aedes cuneatus of Howard, Dyar and Knab (1917:770-773); Dyar (1918a:77).

Aedes (Ochlerotatus) angustivittatus cuneatus of Martini (1935:54).

Aedes (Ochlerotatus) trivittatus cuneatus in part of Dyar (1922a:52).

Aedes (Ochlerotatus) trivittatus of Dyar (1918a:77; 1922a:52, in part; 1923:181); Bonne and

Bonne-Wepster (1925 :392-393, in part).

Aedes (Heteronycha) trivittatus in part of Dyar (1922b:51-52).

Aedes trivittatus of Evans (1921:446; 1922:220).

FEMALE (fig. 15). Wing: 3.40 mm. Proboscis: 2.15 mm. Forefemur: 1.65

mm. Abdomen: 3.10 mm. Thorax: Dorsocentral line of light scales complete, un-

broken from anterior promontory to scutellum; line narrow, broadened only slightly

posteriorly, almost never expanded into supraalar area; scales yellow, rarely light

yellow to white. Legs: Knee spots of white scales absent posteriorly on apices of

femora. Abdomen: Terga entirely dark scaled dorsally.

MALE (fig. 15). Wing: 3.40 mm. Proboscis: 2.20 mm. Forefemur: 1.50 mm.

Abdomen: 3.15 mm. Thorax: Scutal scale pattern similar to female; longitudinal

line of light scales rarely broadened laterally. Legs: Posterior claw of midleg moder-

ately enlarged, with acute subbasal tooth. Abdomen: Terga dark scaled dorsally.

MALE GENITALIA (fig. 15). Segment IX: Anterior middorsal emargination of

tergum broad and deep, caudal bridge connecting tergal lobes narrow, less than dis-

tance between lobes; tergal lobes usually higher than broad, with about 4-6(2-10)

setae. Sidepiece: Length about 4.0-4.5 times median width; tergal surface with 5-6

large subapicotergal setae in oblique row; sternomesal margin with 40-50 long setae;

apical tergomesal lobe well developed; basal tergomesal lobe well developed, triangu-

30 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

lar with posterior side shorter than anterior side, lobe often folded ventrad in slide

preparations, with 15-25 undifferentiated setae, those on margin of cluster longer,

with 4 longer setae projecting distad from posterior edge of lobe. Claspette: Stem

broadly curved dorsad, narrow basally and broadened into calyciform apex from

which the filament arises; filament shorter than stem, narrow at base, abruptly ex-

panded near middle with sharp retrorse process and several elongate spicules basad

of retrorse process, and tapered to recurved tip. Clasper: Spiniform length about

7-10 times its greatest width. Phallosome: Aedeagus ovate to pyriform.

PUPA (fig. 15). Abdomen: 2.75 mm. Trumpet: 0.45 mm. Paddle: 0.80 mm.

Cephalothorax: Weakly pigmented, dorsum slightly darker. Seta 10-C usually 5-7

(4-9) branched. Abdomen: Weakly pigmented, slightly darker anteriorly, strongly

pigmented at base of segment III. Seta 3-VII double or triple (1-5b). Seta 5-IV,V

longer than length of succeeding tergum, usually double. Seta 6-IV double or triple

(1-4b); seta 6-V double (rarely single).

LARVA (fig. 16). Head: 0.90 mm. Siphon: 0.90mm. Anal Saddle: 0.40 mm.

Thorax: Spicules of integument numerous and conspicuous, length about 6-10

times basal diameter. Seta 3-P single. Seta 7-P triple (2-3b). Abdomen: Seta I-

IV usually double (2-3b); seta 1-V double (1-3b). Seta 6-I double (1-2b); seta 6-II

double (2-3b); seta 6-IIIJ-VI single (1-2b). Seta 13-III strongly developed, single,

subequal to seta 13-IV,V. Segment VIII: Comb scales moderate in size, evenly

fringed, 20-24(16-29) in number, in irregular double to triple row. Siphon: Index

about 2.0-2.5. Pecten with teeth evenly spaced, extending slightly beyond middle

of siphon. Seta 1-S inserted distad of pecten. Anal Segment: Ventral brush (4-X)

with 8 pairs of setal tufts, the most proximal tuft occasionally arising from within

saddle.

SYSTEMATICS. Aedes angustivittatus can be separated from the other species

of the Scapularis Group by the following characters: in the female (1) the narrow

dorsocentral line of usually yellow, occasionally yellowish white, scales on the scu-

tum is broadened only very slightly posteriorly and (2) there are no knee spots of

white scales on the apices of the femora; in the male genitalia (1) the claspette stem

is expanded into a cup-like apex from which the claspette filament arises, (2) the

spiniform is short, only about 7 to 10 times longer than wide, and (3) the subapico-

tergal setae number 5 or 6 in an oblique row; in the larva (1) seta 1-IV,V is usually

double, (2) setae 5,6-C are single, (3) seta 6-I is double and 6-II-VI are single, (4)

the comb scales have all spinules subequal, and (5) the spicules of the thoracic and

abdominal integument are long, length 6-10 times basal diameter.

I have seen 158 specimens consisting of 29 males, 72 females, 17 pupae and 40

larvae (13 individual rearings; 6 larval, 6 pupal, 1 incomplete) from several localities

in Mexico and 1 locality in Guatemala that I consider to be hybrids between angust-

ivittatus and trivittatus, and it is evident that a much more thorough study of these

2 species is required in the area where their ranges overlap. The 2 species are allo-

patric except for an area of Mexico from the states of Jalisco and central Veracruz

on the north to Oaxaca on the south. Since angustivittatus appears to be more

abundant and better established than trivittatus in this area, it may indicate a rela-

tively recent southern invasion by trivittatus. The mixing of the gene pools of nu-

merically dissimilar populations would result in introgressive hybridization, and ex-

tensive backcrossing of the hybrids with the dominant form, angustivittatus, would

prevent the formation of a stable hybrid population. According to Mayr (1973:370,

380), populations meeting in zones of secondary intergradation (a belt of highly var-

iable hybrid populations) may show any degree of difference, which is evident in the

Arnell: Scapularis Group 31

females and male genitalia of the hybrids I have seen.

Females of angustivittatus differ from trivittatus in scutal ornamentation, the dor-

socentral light lines being narrow, usually yellow and not broadened posteriorly in

angustivittatus and wider, usually white, and usually broadened laterally above the

wing in trivittatus; knee spots of white scales on the apices of the femora are absent

in angustivittatus and present in (¢trivittatus; and the abdominal terga are entirely

dark dorsally in angustivittatus and usually have basal bands of white scales in

trivittatus. Hybrids exhibit a usually white scutal stripe broader than angustivittatus

and often as broad as trivittatus and the dark femoral apices and dark dorsal abdo-

men typical of angustivittatus.

The male genitalia of angustivittatus differ from trivittatus primarily in the fol-

lowing characters: (1) the subapicotergal setae number 5 or 6 in an oblique row in

angustivittatus and usually 3 in trivittatus, (2) the claspette stem is expanded into a

cup-like apex in angustivittatus and not expanded in trivittatus, (3) the spiniform

length is 7 to 10 times longer than wide in angustivittatus and 12 to 14 times longer

than wide in trivittatus, and (4) the basal tergomesal lobe is smaller in angustivitta-

tus, has a tendency to fold ventrad in slide preparations and the posterior side is

shorter than the anterior side, while in ¢trivittatus the anterior and posterior sides

are subequal and the lobe is larger and seldom folded. The hybrids exhibit a com-

bination of these characters, most often the sidepiece characteristics of angustivit-

tatus and the claspette and spiniform of trivittatus. A male from near Mixco, Guate-

mala (GUA 115B) is trivittatus-like in all respects although all other specimens from

south of Oaxaca, Mexico are typical angustivittatus.

A large series of specimens was collected by Frederick Knab at Cordoba, Vera-

cruz, Mexico from which Dyar and Knab (1908:54-55) described cuneatus and

argentescens, both later (Dyar, 1924a:117) synonymized with angustivittatus. A\l-

though this series of males and females consists of both angustivittatus and hybrids,

the lectotype female of cuneatus and the lectotype male of argentescens are angusti-

vittatus. In the females of this series, the dorsocentral stripe of the scutum is light

yellow in both forms but the stripe is narrow in angustivittatus and much broader

in the hybrid. The male genitalia show the angustivittatus and hybrid characters

discussed above.

BIONOMICS. Aedes angustivittatus is one of the most common Aedes in Central

America, being found in temporary ground pools formed by rains and stream over-

flow, pools in drying or intermittent streams, animal hoof prints in marshy pastures

and margins of freshwater swamps and ponds. It is found with a variety of species

of Psorophora, Culex and other Aedes, and occasionally with species of Anopheles.

Although collected at elevations of over 2000 meters in Mexico and over 1000 me-

ters in Venezuela and Costa Rica, it is primarily a species of low elevations. Kumm,

Komp and Ruiz (1940:392) found angustivittatus to be the third most common

non-anopheline species taken as adults in traps or by animal bait during a survey in

Costa Rica, totaling 10.5% of 5338 captures and ranking behind only Culex (C.)

quinquefasciatus and Mansonia (M.) titillans.

IIheus virus and Venezuelan equine encephalitis virus have been isolated from

angustivittatus and are discussed above under Medical Importance.

DISTRIBUTION (fig. 2). Aedes angustivittatus is known from the Mexican states

of Jalisco, Distrito Federal and Veracruz south through Central America to Colom-

bia, Venezuela and Ecuador. Material examined: 4152 specimens; 621 males, 2807

females, 256 pupae, 468 larvae, 112 individual rearings (77 larval, 29 pupal, 6 in-

complete).

32 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

BELIZE. Belize: Stann Creek (23 mi W), 1 F [USNM]. Cayo: Central Farm, 70 m, 30 July

1967, D. Bertram (BH 360), 1 F [UCLA]; same locality and collector, 17 Aug 1967 (BH 479),

1 F [UCLA]. Mountain Pine Ridge, 10 Aug 1967, P. Williams (BH 490), 2 F [UCLA]. New

Capital Site, 70 m, 28 July 1967, D. Bertram (BH 350), 1 F [UCLA]. Roaring.Creek, 55 m, 24

July 1967, D. Bertram (BH 324-326), 1 F [UCLA]; same locality and collector, 9 Aug 1967 (BH

429), 1 F [UCLA].

COLOMBIA. Caldas: La Dorada, 25 June 1943, W. Komp (KO 200-10), 18 M, 2 F [UCLA];

same locality and collector, 1944 (KO 202C-7), 20 F [UCLA]. Choco: Atrato River, A. Balfour,

2 M, 2 F [BM]. Cundinamarca: Puerto Lievano [?] and Bogota, 6 Aug 1965, B. Amartegui, 1 F

[USNM]. Magdalena: Cienaga, 11 Jan 1944, A. Barreto (COT 36), 1 F [UCLA]. Meta: Retiro

(nr. Restrepo), 6, 12 Nov 1935, 2M [USNM]. Santander: Lebrija, 10 Apr 1966, C. Marinkelle

(COM 644A), 4 F [UCLA]; same data (COM 64C), 5 F [UCLA]. Locality unknown: (COR 506),

1 F [UCLA] ; (COR 497), 1 F [UCLA].

COSTA RICA. Alajuela: Cascajal, H. Kumm, 1 F [BM]. Ciruelas, 14 Nov 1920, A. Alfaro,

1 F [USNM]. San Ramon, 1000 m, 12 Aug 1971, S. Heinemann (CR 340), 1 IlpF (10), 1 pM

(102), 2 pF (100,104), 1 p [UCLA]; same data (CR 342), 1 IpF (10), 1 pM (100), 11 [UCLA].

Cartago: El Empalme, 16 July 1963, C. Hogue (CR 154), 3 IpM (104,106,109), 5 IpF (101,102,

107,108,110), 14 M, 10 F, 4 p, 15 P, 51,41 L [UCLA]. Turrialba, 9 Aug 1962, F. Truxal, 1 F

[UCLA]. Turrialba, 1 Dec 1962, C. Hogue and W. Powder (CR 57), 2 F [UCLA]; same data

(CR 60), 8 F [UCLA]; same locality and collectors, 2 Dec 1962 (CR 64), 3 F [UCLA] ; same lo-

cality, 8 Dec 1962, C. Hogue,,1 F [UCLA]; same locality, 606 m, June 1954, P. Buxton, 1F

[BM]. Guanacaste: Liberia (2 km W), 27 July 1962, F. Truxal, 2 F [UCLA]. Heredia: Pto.

Viejo de Sarapiqui, 7 Aug 1971, A. Berrios Arias (CR 408), 1 F [UCLA]; same locality, 8 Aug

1971, S. Heinemann (CR 421), 2 F [UCLA]; same locality and date, D. Heinemann (CR 427),

1 F [UCLA]. Limon: Beverly, 10 m, 15 Dec 1971, D. and K. Schroeder (CR 558), 6 L [UCLA].

Boston, 46 m, 9 Oct 1971, D. Schroeder (CR 487), 3 pM (102-104), 1 pF (101), 1 P [UCLA].

Estrada, 46 m, 8 Oct 1971, D. Schroeder (CR 482), 5 IpM (13-15,17,71), 20 IpF (11,12,16,18,19,

60-70,72-75), 1 pM (100), 3 pF (101-103), 1 Ip (10), 22 M, 22 F, 31 p, 86 P, 31,30 L [UCLA].

La Bomba (3.7 km W), 3 Oct 1971, D. and K. Schroeder, 1 F [UCLA]. Limon, 28 Sept 1905, F.

Knab, 1 F (angustivittatus holotype) [USNM]. Pueblo Nuevo, 14 Dec 1971, D. Schroeder

(CR 548), 1 F [UCLA]. Siquirres, H. Kumm, 1 F [BM]. Zent, F. Knab, 1 M, 4 F [USNM].

Zent (0.3 km N), 46 m, 8 Oct 1971, D. Schroeder (CR 485), 2 IpM (12,13), 2 IP (10,11), 6 L

[UCLA]. Zent (4 dist. to Boston), 15 m, 12 Dec 1971, D. and K. Schroeder (CR 540), 2 F,E

[UCLA]. Zent, L. Rozeboom, 1 F [UCLA]. Puntarenas: Boca del Rio Barranca, 7 m 14 Aug

1971, D. Schroeder (CR 373), 1 F [UCLA]; same locality, 20 June 1963, C. Hogue (CR 108),

5 F [UCLA]. Dominical, 28 May 1943, T. Aitken (CRM 19), 5 F [UCLA]; same locality and

collector, 22 June 1943 (CRM 24), 2 F [UCLA]. Esparta, 1938, H. Kumm (KO H14-27), 1 F

[UCLA]. Golfito, 9 July-21 Aug 1957, F. Truxal, A. Menke (CR 172), 1 F, 11,21 L [UCLA].

Rio Aranjuez, F. Knab, 2 F [USNM]. San Vito de Java, 22-24 July 1964, R. Woodruff (CRM 32),

1M, 10 F [UCLA]. San Jose: San Jose, 13 June 1963, C. Hogue (CR 92), 11 F [UCLA] ; same

data (CR 94), 1 F [UCLA]; same locality and collector, 16 June 1963 (CR 95), 1 F [UCLA].

Province unknown: [?] Chase, 1938, H. Kumm (CRK 218), 1 M, 1 F [UCLA]. Locality unspec-

ified: 4 F [BM].

ECUADOR. Guayas: Empalme (3 km S), 6 Feb 1966, J. Belkin et al. (ECU 120), 1 F

[UCLA]. Guayaquil, 15 Feb 1966, E. Gerberg (ECU 172), 7 F [UCLA]. Pichilingue, 16, 17

Mar 1946, E. Hambleton, 6 F [USNM]. Samborondon, 10 Feb 1966, J. Belkin (ECU 138),4 M,

12 F [UCLA]. Los Rios: Valencia, 6 Feb 1966, J. Belkin et al. (ECU 122), 1 F [UCLA].

EL SALVADOR. La Liberdad: Atiocoyo, Oct 1941, 2M [USNM]. San Miguel: San Miguel,

1 F [USNM]. Sonsonate: Sonsonate, F. Knab, 5 P [USNM].

GUATEMALA. Escuintla: Iztapa, 10 July 1964, P. Cowsill (GUA 34), 4 F [USNM]. San

Jose de Guatemala (Puerto de San Jose), 17 July 1943, D. Hall (KO 204A-14), 2 F [UCLA].

Santa Lucia Cotzumalguapa, 3 July 1964, T. Zavortink and P. Cowsill (GUA 31), 3 F [UCLA].

Guatemala: Guatemala, 2 Sept 1964, P. Cowsill and Almengor (GUA 119), 4 IpF (10-13), 1

Ip (14) [UCLA]. Mixco, 1700 m, 1 Sept 1964, P. Cowsill and Almengor (GUA 115B), 1 IpM

Arnell: Scapularis Group 33

(63), 3 IpF (61,62,64), 1 Ip (60), 1 M [UCLA]. Izabal: Bananera, 26 June 1964, P. Cowsill

(GUA 4), 1 F [UCLA]; same locality, 11 Aug 1964, T. Zavortink (GUA 114), 6 F [UCLA].

Mojaca Village, 28 June 1964, J. Zavortink and P. Cowsill (GUA 12), 2 F [UCLA]. Morales, 30

July 1963, T. and J. Zavortink (GUA 73), 1 F [UCLA].

HONDURAS. Atlantida: La Ceiba, 6 m, 7 Feb 1972, D. Schroeder (HON 114), 1 F [UCLA].

Lancetilla, 9 Feb 1972, K. and D. Schroeder (HON 121), 2 F [UCLA]; same data (HON 122), 3

pM (106-108), 2 pF (105,109) [UCLA]; same data (HON 123), 6 pM (100,102-104,107,109),

3 pF (101,106,108), 21 [UCLA]. Rio Nutria (0.8 km E on road from Tela to La Ceiba), 70 m,

8 Feb 1972, D. Schroeder (HON 116), 1 F [UCLA]. Tela (2.5 km NW), 2 m, 9 Feb 1972, D.

Schroeder (HON 118), 6 IpM (14,16,19,20), 11 IpF (11-13,15,18,21,23,25-28), 1 M [UCLA].

Tela, 13 Aug 1964, P. Cowsill (HON 27), 1 F [UCLA]. Cortes: La Lima, 5 Aug 1964, P. Cowsill

(HON 1), 1 F [UCLA]. Puerto Cortes, 12 Aug 1967, A. Adames (HON 67), 1 F [UCLA]. San

Pedro Sula (21 km N), 11 Aug 1967, A. Adames (HON 65), 3 F [UCLA].

MEXICO. Distrito Federal: Mexico City, 1919, A. Herrera, 1 F [USNM]; same locality and

collector, 21 June 1926, 1 F [USNM]. Guerrero: El Marquez, 30 Aug 1964, E. Fisher and D.

Verity (MEX 143), 1 F [UCLA]; same data (MEX 140), 1 F [UCLA]. Jalisco: Puerto Vallarta,

15 m, 31 Aug 1972, J. Belkin (MEX 726), 4M, 3 F [UCLA]. Morelos: Tepotzlan (8.2 km W),

1600 m, 28 June 1970, K. and D. Schroeder (MEX 511), 4 IpM (10-13) [UCLA]. Tabasco: Car-

denas, 30 m, 15 July 1970, K. and D. Schroeder (MEX 564), 1 F [UCLA]. Comalcalco (7.5 km

N), 36 m, 12 July 1970, K. and D. Schroeder (MEX 552), 1 F [UCLA]. Veracruz: Cordoba, 20

Jan or 8 Apr 1908, F. Knab, 6 M, 3 F (440.1); 1 M, 1 F (440.2); 1 M, 1 F, 2 p, 21 (441.1);3 M,

1 F, 3 1 (441.2); 1 M (441.3); 6 M, 6 F (441.4); 1 M, 5 F (441.5); 1 M (425.1); 1 M (425.2); 1 F

(425.3); 1 M (425.4), 1 M (425.5); 1 F (425.6); 1 F (425.7); 1 M, 2 F, 21 (425.8); 1 M,3 F

(425.9); 1 F (425.11); 1 M (441.4, argentescens \ectotype); 1 1,lpF (422.19, cuneatus \ecto-

type) [USNM]. Cordoba, 9 Aug 1965, D. Schroeder and S. Law (MEX 281), 1 F [UCLA] ; same

data (MEX 284), 3 F [UCLA]. Cordoba, 915 m, 11 July 1964, C. Hogue (MEX 376), 3 M, 2 F

[UCLA]. Ixhuatlan-Cordoba (3 km S), 1330 m, 8 July 1970, K. and D. Schroeder (MEX 533),

6M, 11 F [UCLA]. Jesus Carranza (Santa Lucrecia), F. Knab, 14 F [USNM].

NICARAGUA. Comarca de El Cabo: San Carlos (on Rio Segovia) (KO H21-24), 1 F [UCLA].

Leon: Isla Penas Blancas, nr. Puerto Somoza, 16 June 1964 (NI 13), 17 F [UCLA]. Puerto

Somoza, 18 June 1964 (NI 20), 1 F [UCLA]. Zelaya: Bluefields, 60 m, 23 Nov 1971, D. and K.

Schroeder (NIC 84), 2 F [UCLA]; same locality and collectors, 8 m, 24 Nov 1971 (NIC 89), 1 F

[UCLA]. El Recreo, 29 June 1954, K. Neiland, 8 F [UCLA]. Punta Masalla (4 km S Bluefields),

1m, 27 Nov 1971, D. and K. Schroeder (NIC 113), 1 F [UCLA].

PANAMA AND CANAL ZONE. Bocas del Toro: Almirante, Feb 1931 (PAX 22),3 M,1F

[UCLA]. Almirante, July 1934 (PAX 41), 7M [UCLA]. Almirante, 19 Aug 1944, W. Komp

(KO 200-29), 2 F [UCLA] ; same locality and collector, 24 Aug 1944 (KO H21-29), 1 F [UCLA].

Almirante, 27 Apr 1963 (PA 269), 1 F [UCLA]. Almirante, 28 Apr 1963 (PA 270), 252 M, 415

F [UCLA]. Almirante, 30 Apr 1963 (PA 294), 20 F [UCLA]. Almirante, 2 May 1963 (PA 304),

23 M, 247 F [UCLA]. Almirante, 3 May 1963 (PA 305), 1 F [UCLA]. Almirante, 3 May 1963

(PA 308), 36 M, 712 F [UCLA]. Almirante, 3 May 1963 (PA 309), 3 F [UCLA]. Almirante, 5

May 1963 (PA 321), 4M, 467 F [UCLA]. Almirante, 7 May 1963 (PA 331), 2 M, 10 F [UCLA].

Almirante, 8 May 1963 (PA 338), 5 F,321 M [UCLA]. Almirante, 15 Apr 1964 (PA 670), 185 L

[UCLA]. Chiriqui Grande, 17 Apr 1963 (PA 220), 6 F [UCLA] ; same locality, 19 Apr 1963 (PA

243), 2 F [UCLA]. Chiriquisito, 15 Apr 1963 (PA 207), 10 F [UCLA]. Chiriquisito, 16 Apr

1963 (PA 208), 12 F [UCLA]. Chiriquisito, 16 Apr 1963 (PA 217), 2 IpM (102,106), 4 lpF (101,

103,105,107), 1 lp (104), 1 P,67 L [UCLA]. Chiriquisito, 18 Apr 1963 (PA 226), 16 F [UCLA].

Chiriquisito, 19 Apr 1963 (PA 237), 20 F [UCLA]. Chiriquisito, 22 Apr 1963 (PA 249), 1 M,

2 F [UCLA]; same data (PA 250), 18 M [UCLA]. Canal Zone: Albrook Field, W. Komp, 1 F

[UCLA]. Albrook Field (PA 743-C), 1 F [UCLA]. Cocoli, June 1946, C. Pierce, 1 F [UCLA].

Ft. Davis, 7 Nov 1951, S. Carpenter, 16 F [UCLA]. Ft. Davis, 22 Nov 1966, R. Schick and A.

Quinonez (PA 813), 2 F [UCLA]. Ft. Kobbe, 26 Nov 1951, S. Carpenter, 2 F [UCLA]. Ft.

Sherman, 23 Nov 1939, W. Komp (KO 202C-9), 2 F [UCLA]. Gatun, 9 Aug 1923, H. Dyar and

R. Shannon, 3 F [UCLA]. Gatun [?] Tarpon, 12 Dec 1951, S. Carpenter, 5 F [UCLA]. Las

34 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

Cascadas (3 km SE Gamboa), A. Busck, 1 M [BM]. Madden Dam, 27 Oct 1948, 1 F [UCLA].

Madden Dam, 13 Aug 1949, 1 F [UCLA]. Matachin (1 km SW Gamboa), 29 June 1913, J.

Zetek, 2 M [USNM]. Pedro Miguel, 22 Apr 1947, H. Brooks, 2M, 2 F [UCLA]. Tabernilla (12

km W Gamboa), 12 May 1908, A. Jennings, 1 F [UCLA]; same locality and collector, 2 F [BM].

Locality unspecified (CZ 49), 1 M [UCLA]. Chiriqui: David, 6 June 1943, F. Snyder (KO 202A-

29), 2 F [UCLA]. Colon: Colon, L. Dunn, 1 F [UCLA]. Colon, 27 July 1920, W. Chidester, 3 F

[USNM, UCLA]. Colon, 18 June 1929, D. Curry (PAX 3), 2 F [UCLA]; same locality and collec-

tor, 10 Jan 1930 (PAX 13), 1 M [UCLA]. Rio Alejandro (4 km N Puerto Pilon), 14 Feb 1940,

W. Komp, 3 F [UCLA]. Darien: El Real, 25 May 1936, L. Rozeboom (PAR 34), 1 F [UCLA].

Panama: Huile, 23 Nov 1965, A. Quinonez (PA 834), 1 pM (50), 1 pF (101),5 M, 1 1,5L

[UCLA]. La Chorrera, 14 Aug 1944, Wood and Van Doran (ASM 95), 1 M, 3 F [UCLA]. La

Chorrera, 21 Aug 1944 (ASM 124), 2 F [UCLA]. La Chorrera, 26 Sept 1944, P. Adams (ASM

171), 5 F [UCLA]. Tocumen, 10 m, 27 July 1972, H. Arnell (PA 1147), 2 IpM (40,41), 1 L

[UCLA]. Tocumen, 11 Sept 1963 (PA 553), 2 F [UCLA]. Province unknown: [?] Alhajuelo,

17 Apr 1911, A. Busck, 15 F [UCLA]. Locality unknown: 8 Sept 1934, L. Rozeboom (PAR

63), 2 F [UCLA].

VENEZUELA. Aragua: Cagua, 400 m, 12 Aug 1969, J. Valencia and J. Pulido (VZ 344), 1 F

[UCLA]. [La] Trinidad, 30 Jan 1929 (VZR 141), 14M, 3 F [UCLA] ; same data (VZR 162), 16

M, 2 F [UCLA]. Maracay, 20 Aug-25 Oct 1926, M. Nunez Tovar, 5 M, 9 F [UCLA], USNM];

same locality and collector, 10 May 1927 (VZR 4), 2 F [UCLA]; same locality and collector, 3

July 1927, 1 M [USNM]. Maracay (VZR 242), 3 M [UCLA]. Maracay (VZR 247), 32 M, 22 F

[UCLA]. Maracay (VZR 249), 16 M, 13 F [UCLA]. Ocumare de la Costa, 5 July 1927, M.

Nunez Tovar, 1 F [USNM]; same locality, 5 July 1927 (VZR 14), 4 F [UCLA]. Palo Negro, 11

Dec 1967, J. Vera (VZ 112 2), 1M, 1 F [UCLA]. Rancho Grande (1.5 km S), Dec 1967, Hansell

and Rauch (VZ 90), 1 pF (101), 3 F,3 p [UCLA]. San Jacinto Mil. Res. (nr. Maracay), 550m, J.

Valencia and J. Pulido (VZ 242), 1 M [UCLA]; same locality, 30 Oct 1928 (VZR 83), 1 F

[UCLA] ; same locality, 27 Nov 1928 (VZR 98), 7M [UCLA]; same locality, 12 Dec 1928 (VZR

103); 1 M [UCLA]; same locality, 19 Feb 1929 (VZR 150), 2M, 2 F [UCLA]; same locality, 11

Mar 1929 (VZR 153), 1 M, 2 F [UCLA]; same locality, 13 Apr 1929 (VZR 168), 3 M [UCLA];

same locality, 3 July 1929 (VZR 212), 1 M [UCLA]; same locality, 1929, 3 M, 1 F [UCLA].

Turiamo, 11 Sept 1944, W. Komp, 2 M [UCLA]; same locality and collector, 14 Sept 1944 (KO

202A-10), 1 F [UCLA]. Carabobo: Lake Valencia, 29 Nov 1967, Hansell and Rauch (VZ 69), 2

IpF (11,12), 1 lp (10), 1 F, 2 L [UCLA] ; same data (VZ 64), 1 F [UCLA]. Portuguesa: Acarigua

(VZR 230), 3 F [UCLA]. Zulia: Zulia River, L. Dunn, 1 F [USNM]; same data, 1 F (traversus

lectotype) [USNM].

Hybrids

GUATEMALA. Guatemala: Mixco, 1700 m, 1 Sept 1964, P. Cowsill and Almengor (GUA

115B), 1 M [UCLA].

MEXICO. Morelos: Tepotzlan (8 km W), 1600 m, 28 June 1970, K. and D. Schroeder (MEX

508), 1 IpF (10), 4 L [UCLA]; same data (MEX 509), 1 IP (10), 2 L [UCLA]; same data (MEX

511), 1 IpM (15), 1 IpF (14), 2 pM (101,102), 2 pF (100,104), 4 L [UCLA]. Oaxaca: Almoloya,

F. Knab, 18 M, 15 F (309) [USNM]. Asuncion Nochixtlan (1.5 km SE), 24 Aug 1964, D.

Verity (MEX 134), 19 L [UCLA]. Cameron (34 km SE), 21 Aug 1964, E. Fisher (MEX 133), 2 L

[UCLA]. Matias Romero (42 km N, nr. Juchitan), 26 July 1963, E. Fisher (MF 13), 1 F

[UCLA]. San Luis Potosi: Saketepan, 200 m, 20 July 1965, D. Schroeder (MEX 215), 5 F

[UCLA]. Veracruz:. Cordoba, 16 Mar 1908, F. Knab, 1 F [USNM]. Cordoba, 7 Aug 1965, D.

Schroeder and S. Law (MEX 278), 2 IpM (9-10,11), 1 IpF (9-12), 4M, 15 F,3 p,1P,11,3L

[UCLA] ; same locality and collectors, 9 Aug 1965 (MEX 281), 11 F [UCLA]; same data (MEX

282), 6 F [UCLA]; same data (MEX 283), 1 F [UCLA]; same data (MEX 284), 1 IpF (3-12),

1 pM (3-10), 1 pF (3-11), 5 F 21 [UCLA]. Cordoba, 915 m, 11 July 1965, C. Hogue (MEX 376),

3 F [UCLA]; same locality and collector, 12 July 1965 (MEX 378), 2 F [UCLA]. Jesus Carranza

(Santa Lucrecia), F. Knab, 1 F [USNM].

Arnell: Scapularis Group 35

7. Aedes (O.) trivittatus (Coquillett)

Figs. 2,17,18,19

1902. Culex trivittatus Coquillett, 1902:193-194. TYPE: Holotype female, Chester, New Jer-

sey, USA, 10 or 14 Sept, J. B. Smith [USNM, 6702; see Stone and Knight, 1956:226].

1904. Culex inconspicuus Grossbeck 1904:333. TYPE: Lectotype male, Garret Mts., Pater-

son, New Jersey, USA, 5 Oct [USNM; selection of Stone and Knight, 1956:219].

Synonymy with trivittatus by Howard, Dyar and Knab (1917:773).

Aedes (Ochlerotatus) trivittatus of Dyar (1922a:52, in part; 1928:162-163); Root (1924:452);

Bonne and Bonne-Wepster (1925:392-393, in part); Edwards (1932:143); Martini (1935:54);

Matheson (1944:161-162); Ross (1947:75-76); Darsie (1951:21); Carpenter and La Casse

1955:249-251); Horsfall and Craig (1956:372); Vargas (1956:24); Barr (1958:109-110); Stone,

Knight and Starcke (1959:156); Forattini (1965:329-33 1); Trimble (1972: 1535).

Aedes (Heteronycha) trivittatus of Dyar (1920:105; 1922b:51-52).

Aedes trivittatus of Dyar and Knab (1906a:197); Howard, Dyar and Knab (1917:773-776); Dyar

(1918a:77; 1924a:117); Shlaifer and Harding (1946:250); Weathersbee and Arnold (1947:

218); Abdel-Malek (1948:951-954; 1949:19); Breland (1951:362); Judd (1954:107); Price

(1960:558); Breeland, Snow and Pickard (1961:293-294); Steward and McWade (1960:156-

157); Horsfall, Vorhees and Cupp (1970:1711).

Ochlerotatus trivittatus of Coquillett (1906b:21).

Grabhamia trivittatus of Dyar (1905a:48; 1905c:187).

Pseudohowardina trivittata of Theobald (1907:224-227; 1910:227).

Culex trivittatus of Blanchard (1905:339); Theobald (1905:27).

Aedes inconspicuus of Dyar and Knab (1906a:199).

Ochlerotatus inconspicuus of Coquillett (1906b:21).

Culex inconspicuus of Dyar (1905b:108); Theobald (1910:387).

Aedes (Ochlerotatus) trivittatus cuneatus in part of Dyar (1922a:54).

Aedes infirmatus of Rigby (1968:239-240).

FEMALE (figs. 17,18). Wing: 3.75 mm. Proboscis: 2.40 mm. Forefemur:

1.70 mm. Abdomen: 3.25 mm. Thorax: Dorsocentral line of light scales com-

plete, unbroken from anterior promontory to scutellum, usually expanded laterally

in supraalar area; scales usually white, occasionally yellowish white to light yellow;

white scales present above paratergite. Legs: Knee spots of white scales present

posteriorly on apices of femora. Abdomen: Terga entirely dark scaled dorsally or

with narrow basal band or median patch of white scales on terga II-VI.

MALE (figs. 17,18). Wing: 3.50 mm. Proboscis: 2.30 mm. Forefemur: 1.60

mm. Abdomen: 3.55 mm. Thorax: Longitudinal line of light scales usually

broadened laterally, often to scutal margin. Legs: Posterior claw of midleg small,

with acute subbasal tooth. Abdomen: Terga with basal bands of white scales.

MALE GENITALIA (fig. 18). Segment IX: Anterior middorsal emargination of

tergum broad and shallow, caudal bridge connecting tergal lobes broad; tergal lobes

usually broader than high, with 4-8 setae. Sidepiece: Length about 3.0-4.0 times

median width; tergal surface with lateral setae longer than mesal setae and 3 sub-

apicotergal setae; sternomesal margin with about 45-55 long setae; apical tergomesal

lobe moderately developed; basal tergomesal lobe well developed, triangular with

anterior and posterior sides subequal in length, about 20-25 undifferentiated setae,

those on margin of setal cluster longer, with 4 longer setae projecting distad from

posterior edge of lobe. Claspette: Stem broadly curved dorsad, narrowed slightly

distally; filament subequal in length to stem, narrow and slightly angled dorsad near

base, abruptly expanded near middle with sharp retrorse process and usually several

36 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

spicules basad of retrorse process, and tapered to recurved tip. Clasper: Spiniform

length about 12-14 times its greatest width. Phallosome: Aedeagus conical to

slightly pyriform.

PUPA (fig. 18). Abdomen: 2.80 mm. Trumpet: 0.55 mm. Paddle: 0.80 mm.

Cephalothorax: Weakly pigmented, dorsum darker. Seta 10-C usually 6-9(6-10)

branched. Abdomen: Weakly pigmented, darker anteriorly, strongly pigmented at

base of segment III. Seta 3-VII usually double (1-3b). Seta 5-[V-VI subequal to or

slightly longer than length of succeeding tergum; double, very rarely single. Seta 6-

IV,V usually single (rarely double or triple).

LARVA (fig. 19). Head: 0.85 mm. Siphon: 0.80 mm. Anal Saddle: 0.35 mm.

Thorax: Spicules of integument moderately numerous and conspicuous, length

about 4-5 times basal diameter. Seta 3-P single. Seta 7-P triple (3-4b). Abdomen:

Seta 1-[V double (2-3b); seta 1-V double (1-3b). Seta 6-I,] double (rarely single);

seta 6-III-VI single (very rarely double). Seta 13-III strongly developed, single (1-

2b), subequal to seta 13-IV,V. Segment VIII: Comb scales moderate in size, usual-

ly with moderately differentiated median spinule, usually 20-24(16-27) in number,

in irregular double to triple row. Siphon: Index about 2.1-2.7. Pecten with teeth

evenly spaced, extending slightly beyond middle of siphon. Seta 1-S inserted just

distad of pecten. Anal Segment: Ventral brush (4-X) with 8 pairs of setal tufts.

SYSTEMATICS. Aedes trivittatus can be separated from the other species of the

Scapularis Group by the following characters: in the female (1) the dorsocentral

line of white or occasionally yellowish white scales on the scutum is broadened in

the supraalar area and (2) knee spots of white scales are present on the apices of the

femora; in the male genitalia, generally indistinguishable from bogotanus, tortilis,

condolescens, euplocamus and patersoni, (1) the sidepiece length is 3 to 4 times

median width, (2) the claspette filament is subequal in length to the claspette stem

and has a large retrorse process with several elongate spicules basad of the process,

the spicules seldom extending basad of the basal 0.2 of the filament, (3) the sterno-

mesal margin of the sidepiece has 45 to 55 long setae between the basal tergomesal

lobe and the clasper, (4) the claspette stem is broadly curved dorsad and not ex-

panded distally, (5) the apical tergomesal lobe of the sidepiece is moderately to well

developed, and (6) the spiniform length is about 12 to 14 times its greatest width;

in the larva (1) seta 1-IV,V is usually double, (2) setae 5,6-C are single, (3) seta 6-I

is double and 6-II-VI is single, (4) the median spinule of the comb scales is slightly

larger than the lateral spinules, and (5) the spicules of the thoracic and abdominal

integument are only moderately long, length about 4 to 5 times basal diameter.

In the female, this species is inconstant in the amount of white scaling on the dor-

sum of the abdomen, many specimens being completely dark scaled but most having

a narrow basal band or median patch of white scales on the base of terga II to VI,

and the dorsocentral line of light scales on the scutum is usually white but rarely

yellowish white or yellow. There is also some variability in the length of the median

spinule of the larval comb scale, occasionally it being subequal in length to the sub-

apical spinules. Specimens with these variations seem to be more or less randomly

distributed in the trivittatus population.

Aedes trivittatus is one of the more common Aedes species in the upper midwest

and northeast United States as discussed below under bionomics, and is a Nearctic

representative of a primarily Neotropical group (other exceptions are infirmatus and

thelcter in the southern United States, and intrusions of tortilis and scapularis into

the Nearctic in northeastern Mexico and Florida). The present distribution of trivit-

tatus is probably post-Pleistocene and may be northward expansion from Pleisto-

Arnell: Scapularis Group a

cene refugia in northern Mexico and the southwestern United States. If this is in-

deed the case, trivittatus may have been much more common in central and south-

ern Mexico during the Pleistocene, which may account for the apparent introgres-

sive hybridization of trivittatus with angustivittatus in southern Mexico (see discus-

sion under angustivittatus above). The above mentioned variation in female scutal

and abdominal scaling and larval comb scales in some (rivittatus specimens from

throughout its range may be a manifestation of the angustivittatus genotype result-

ing from this mixing of the gene pool.

BIONOMICS. Aedes trivittatus is relatively common throughout the northeastern

and central United States. It occurs in a wide variety of temporary freshwater pool

situations, being commonly found in grassy depressions and bottomland forest pools

filled by rain or stream overflow, although it is also found around the grassy edges

of permanent ponds and marshes. In the more arid western United States and Mex-

ico it occurs primarily along stream courses in overflow pools and blocked stream

beds. In temperate climates trivittatus overwinters in the egg stage, the adults

emerging in late spring. Development is completed in a short time, usually less than

2 weeks. The most common associates of trivittatus are Aedes (Aedm.) vexans and

A. (O.) sticticus. Females of trivittatus are fierce and persistent biters, usually active

in shade in the late afternoon and at dusk, although they will attack at any time of

day or in any light if they are disturbed from their resting place in vegetation. Ab-

del-Malek (1948) discusses the habitat, larval development, feeding behavior and

oviposition, and Abdel-Malek (1949) presents a superficial description of the egg,

4 larval instars and pupa of ¢trivittatus. A more thorough description and figures

of the egg are given by Horsfall and Craig (1956:372).

The considerable extent of arbovirus activity in this species is discussed above

under Medical Importance.

DISTRIBUTION (fig. 2). Aedes trivittatus is known from Nova Scotia, Ontario

and Manitoba in Canada, the United States as far south as Georgia and Texas and

west to Idaho, Utah and Arizona, and Mexico as far south as Oaxaca. Material ex-

amined: 860 specimens; 100 males, 415 females, 137 pupae, 208 larvae; 46 individ-

ual rearings (43 larval, 2 pupal, 1 incomplete).

MEXICO. Coahuila: El Carmen (4 km W; 62 km W of La Rosa on hwy 40), 1500 m, 9 July

1965, R. Schick and D. Schroeder (MEX 184), 1 F [UCLA]. Distrito Federal: Mexico City, 21

June 1926, A. Herrera, 1 F [USNM]. Xochimilco, 4-8 May 1973 (MEX 744), 1 F [UCLA];

Xochimilco, R. Balanzario, 1 F [USNM]. Morelos: Tepotzlan (8 km W), 1600 m, 28 June 1970,

K. and D. Schroeder (MEX 507), 1 IP (10) [UCLA]; same data (MEX 511), 2 pM (103,105)

[UCLA]. Oaxaca: Oaxaca, 30 June 1944, B. Brookman, 1 M [UCLA]. Tule, 3 June 1944, B.

Brookman, 1 F [UCLA]. Zacatecas: Zacatecas (20 km SW on hwy 54), 2000 m, 9 June 1971,

T. Zavortink and L. Nielsen (MEX 684), 2 F [UCLA].

UNITED STATES. Arizona: Duncan, Greenlee Co., 17 Sept 1953, 1 F [UTAH]. Ft. Huachu-

ca, 31 Aug 1966, J. King, 1 M [USNM]. Ft. Huachuca, 7-9 Aug 1973, Fujii, 2 M [Ft. Baker];

same locality and collector, 24 July 1973, 1 M [Ft. Baker] ; same locality and collector, 21-23 Aug

1973, 1 M [Ft. Baker]; same locality, 1 M [UCLA]. Patagonia (4 km SW), 13 Sept 1964, J. Bur-

ger (UCLA 258), 5 M, 4 F, 2 L [UCLA]; same data (UCLA 271), 12 L [UCLA]. Sabino Canyon,

Santa Catalina Mts., 17 Aug 1963, J. Burger (UCLA 399), 2 F [UCLA] ; same locality and collect-

or, 18 Aug 1963 (UCLA 400), 3 F [UCLA]; same locality and collector, 21 Sept 1963 (UCLA

406-1), 1 F [UCLA]; same locality and collector, 24 Sept 1963 (UCLA 407-1), 3 F [UCLA].

Williams, 2100 m, 4,5 Aug 1956, J. Belkin and W. McDonald (UCLA 211,216), 3 F [UCLA].

Colorado: Aurora, 6 Sept 1944, L. Perry, 1 F [UTAH]. Denver, 26 Aug 1948, Anthony, 1 L

[USNM]. Ft. Logan, 10 July 1944, L. Perry, 1 F [UTAH]. Rifle, 12 Aug 1958, F. Harmston,

2 F [UCLA]. Weld Co., 14 June 1950, G. Thompson, 1 M [UCLA]. District of Columbia: Aug

38 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

1928, F. Edwards, 2 M, 1 F [BM]. Idaho: Nampa, 27 June 1945, F. Harmston, 1 F [UTAH].

Illinois: Champaign, June 1972, Busey (UCLA 736), 14 IpM (20-29 40-43), 24 IpF (10-19,30-

39.44.47). 95 M10 F, 35 p, 351,35 L, 5 [UCLA]. Havana, 8 July 1941, Burke and Reigel, 1 F

[UTAH]. Wadsworth, 10 June 1942, Ross and Sanderson, 1 F [UTAH]. Indiana: Spencer, 21

June 1934, 6 F [USNM]. Towa: Mt. Pleasant, 1 June 1923, Yant, 5 F [USNM]. Kansas: Doug:

las Co., Aug 1956, 1 F [USNM]. Lawrence, 25 June 1951, R. Roberts, 3 F [UTAH]. Maine:

Paris, 3 Sept 1910, W. Perham, 5 F [USNM]. Maryland: Baltimore, 5 Aug 1919, 2 F [UCLA].

Massachusetts: S. Amherst, 10 July 1903, G. Dimmock, 1 F [USNM]. Westfield, 30 July 1903,

F. Knab, 1 F [USNM]. Michigan: St. Joseph, 25 June 1942, C. Sabrosky, 1 M, 1 F [USNM]. So.

Haven, 23 July 1938, C. Sabrosky, 1 F [USNM]. Minnesota: Chatfield, 29 Aug 1923, 12 F

[USNM]. Thief River Falls, 29 May 1922, H. Dyar, 2 F [USNM]; same locality and collector,

30-31 May 1922, 28 F [USNM]. Missouri: Camp Crowder, 12 Aug-13 Sept 1942, A. Gumey,

7 F, 2 L [USNM]. St. Louis, 23 June 1952, R. Roberts, 1 F [UTAH]. Montana: Glasgow, 12

July 1921, H. Dyar, 33 F [USNM]. Glendive, 18 June 1922, H. Dyar, 10 F [USNM]. Laurel,

16 July 1917, H. Dyar, 1 IpM (Y3), 4 lpF (Y1,Y2,Y4,Y5), 2 M, 40 F [USNM]. Miles City, 23

July-25 Aug 1915, 6 M, 17 F [UTAH]. Nebraska: Crete, 13 Aug 1944, 1 F [UCLA]. Hall Co.,

21 Aug 1949, G. Thompson, 1 M [UCLA]. Lincoln, 19 Aug 1954, L. Quate, 3 F [UCLA]. Rulo,

Richardson Co., 22 July 1944, 1 F [UCLA]. Valentine, 6 Nov 1942, 1 F [UCLA]. Walthill,

Thurston Co., 29 June 1943, 3 F [UCLA]. Wellfleet, Lincoln Co., 24 June 1961, F. Harmston,

3 F [UTAH]. New Jersey: Chester, 10 or 14 Sept, J. Smith, 1 F (trivittatus holotype) [USNM].

Chester, J. Aldrich, 1 F [USNM]. Culver’s Lake, 13 June 1947, S. Carpenter, 1 M [UTAH] ; same

locality and collector, 11 July 1947, 3 L [UTAH]. Garret Mts., Paterson, 5 Oct,1 M (inconspicu-

us lectotype) [USNM]. New Brunswick, 30 July 1 F [BM]. [?] Gt. Piece Mdw, 6, 15 Sept, J. Al-

drich, 2 F [USNM,BM]. New Mexico: Canjillon, Rio Arriba Co., 12 June 1967, L. Nielsen, 9 L

[UTAH]. San Juan Co., 9 July 1960, L. Jensen, 1 F [UTAH]. Socorro (Escondida Rec. Area),

1400 m, 3 Sept 1966, T. Zavortink (UCLA 324), 2 F [UCLA|. New York: Tuxedo [Park], Aug

1928, F. Edwards, 1 F [BM]. North Dakota: Mandan, 15 June 1923, H. Dyar, 7 F [USNM].

Pennsylvania: Shippensburg, 8 Sept 1933, 5 F [USNM]. Texas: Denison, 22 June 1904, H.

Barber, 1 M, 22 F [USNM]. Utah: Indian Creek St. Park, San Juan Co., 14 June 1970, L. Nielsen

and S. Romney (N-9-70), 18 M, 19 F, 56 p, 36 1,9 L [UTAH]. Canyonlands N. P., San Juan Co.,

15 June 1970, L. Nielsen and S. Romney (N-10-70), 9 M, 15 F [UTAH]. Virginia: Rosslyn, 3

Aug 1916, A. Caudell, 1 F [USNM]. Woodstock, 5 Aug 1904, F. Pratt, 15 F [USNM]. Wiscon-

sin: Darlington, 26 Aug 1933,41 F [USNM]. Univ. Wisconsin Arboretum, Dane Co., 15 June-13

Aug 1960, P. Thompson, 3 M, 22 F [USNM]. Wyoming: Thermopolis, 3 Aug 1944, L. Perry,

1 F [UTAH].

Additional Records from the Literature

CANADA. Manitoba: Pinawa, 10 km NE (Trimble, 1972:1535). Nova Scotia: Pictou (Gib-

son, 1939:109). Ontario: London (Judd, 1954:107).

UNITED STATES. Alabama: Pickwick Res. (Breeland, Snow and Pickard, 1961:293). Geor-

gia: Lee County (Root, 1924:452). South Carolina: Georgetown (Bradley, Fritz and Perry,

1944:109). Texas: Junction (Breland, 1951:369).

TORTILIS SUBGROUP

FEMALES. Head: Occipital dorsolateral dark scale patch conspicuous to nearly

absent. Palpus 3 or 4 segmented. Thorax: Anterior of scutum usually with golden

to tan scales to near level of posterior end of paratergite except for dark brown

scales covering most of fossa; golden to tan scales often extending posteriorly along

acrostichal line and surrounding prescutellar bare space and along dorsocentral line

to scutellum, and occasionally extending laterally into supraalar area; a few whitish

Arnell: Scapularis Group 39

scales above paratergite; remainder of scutal scales dark brown. Scutum rarely uni-

formly dark brown scaled or with anterior scales only slightly lighter in color.

Acrostichal, anterior dorsocentral, and posterior fossal setae absent. Subspiracular

scale patch reduced. Legs: Knee spots of white scales absent posteriorly on apices

of femora. Tibiae and basal tarsal segments with conspicuous white streak posterior-

ly. Claws of hindleg with moderate to small acute submedian tooth, or claws sim-

ple. Wing: Scales entirely dark. Abdomen: Terga II-VII with narrow to broad

basal white bands, usually narrower or absent on more distal segments.

MALES. Thorax: Scutal scale pattern generally similar to female, light scales

usually white. Legs: Anterior claw of foreleg and midleg considerably enlarged,

with blunt submedian tooth and acute basal tooth; posterior claw of foreleg mod-

erately enlarged. Abdomen: Terga with narrow basal bands of white scales.

DISCUSSION. The Tortilis Subgroup is defined by the female ornamentation of

a large anterior scutal scale patch of usually golden to tan scales which contrast

with the dark brown scales of most of the remainder of the scutum, and the very

small subspiracular scale patch, often consisting of fewer than 8 scales.

The subgroup consists of 2 species, tortilis and auratus. Aedes tortilis inhabits

the West Indies south to St. Lucia, with a limited intrusion into southern Florida;

auratus is known only from Kingston, Jamaica.

&. Aedes (O.) tortilis (Theobald)

i Ae Fe 6 a I

1903. Culex tortilis Theobald, 1903b:281-282. TYPE: Lectotype female, Kingston, Jamaica,

20 Aug 1903, M. Grabham [BM; selection of Belkin, 1968:8].

1906. Culex practeatus Coquillett, 1906a:184. TYPE: Holotype female, Havana, Cuba, 1 Nov

1902, J. R. Taylor [USNM, 7753; see Stone and Knight, 1956:215]. Synonymy with

tortilis by Dyar (1922a:55) as subspecies.

1906. Aedes habanicus Dyar and Knab, 1906a:198. TYPE: Lectotype fragments of a larval

skin, Havana, Cuba, 28 Oct 1903, J. R. Taylor [USNM, selection of Stone and Knight,

1956:218]. Synonymy with bracteatus apparently by Howard, Dyar and Knab (1917:

802).

1907. Aedes balteatus Dyar and Knab, 1907:9-10. TYPE: Holotype female, Santo Domingo

(Dominican Republic), Aug 1905, A. Busck [USNM, 10142; see Stone and Knight, 1956:

215]. Synonymy with tortilis by Dyar (1922a:55-56), as subspecies.

1907. Aedes plutocraticus Dyar and Knab, 1907:11. TYPE: Holotype female (no. 10), Nas-

sau, Bahama Islands, 21 June 1903, T. H. Coffin [USNM, 10251; see Stone and Knight,

1956:224]. Synonymy with tortilis by Dyar (1922a:55), as subspecies.

1922. Aedes tortilis virginensis Dyar, 1922a:56. TYPE: Holotype female, St. Thomas, Virgin

Islands, Aug 1905, A. Busck [USNM, 24898].

Aedes (Ochlerotatus) tortilis of Bonne and Bonne-Wepster (1925:394), Dyar (1928:169-170, in

part); Edwards (1932:142, in part); Lane (1939:117, in part; 1953:653-654, in part); Prit-

chard, Seabrook and Mulrennan (1947:11); Carpenter and La Casse (1955:245-247, in part);

Perez Vigueras (1956:296-301); Stone, Knight and Starcke (1959:156, in part); Forattini

(1965:369-370, in part); Montchadsky and Garcia Avila (1966:29,78); Porter (1967:38,39,

40); Stone (1969:5); Belkin, Heinemann and Page (1970:165-168).

Aedes tortilis of Howard, Dyar and Knab (1917:806-809, in part); Dyar (1918a:77); Gerry (1932:

38-39); Staebler and Buren (1946:685); Thurman, Haeger and Mulrennan (1949:171); Torre y

Callejas, Alayo Delmau and Calderon Chapman (1961:67, in part); Garcia Avila and Gutsevich

(1969:6).

40 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

Culex tortilis of Theobald (1905:26); Theobald and Grabham (1905:26); Surcouf and Gonzalez-

Rincones (1911:175).

Aedes (Ochlerotatus) tortilis tortilis (in part) of Dyar (1922a:54-55).

Aedes (Ochlerotatus) tortilis bracteatus, tortilis balteatus and tortilis plutocraticus of Dyar

(1922a:54-56).

Aedes tortilis balteatus of Root (1927:465).

Aedes (Ochlerotatus) bracteatus, balteatus, plutocraticus and virginensis of Bonne and Bonne-

Wepster (1925:393-394).

Aedes (Heteronycha) tortilis, bracteatus and plutocraticus of Dyar (1920:105).

Aedes bracteatus of Pazos (1908:423); Howard, Dyar and Knab (1917:802-804); Dyar (1918a:

71).

Ochlerotatus bracteatus of Coquillett (1906b:20).

Aedes balteatus of Howard, Dyar and Knab (1917:809-810); Dyar (1918a:77).

Aedes plutocraticus of Johnson (1908:70); Howard, Dyar and Knab (1917:804-805); Dyar

(1918a:77).

Aedes (Ochlerotatus) ? condolescens of Root (1922:398-399).

FEMALE (figs. 20,21). Wing: 2.80 mm. Proboscis: 2.10 mm. Forefemur:

1.60 mm. Abdomen: 2.80 mm. Thorax: Anterior of scutum except for fossa

usually with golden to tan scales; scutum rarely uniformly dark brown scaled or

with anterior scales only slightly lighter in color.

MALE (figs. 20,21). Wing: 3.05 mm. Proboscis: 2.20 mm. Forefemur: 1.55

mm. Abdomen: 3.05 mm. Thorax: Anterior of scutum except for fossa with

white to yellowish white scales, white scales often continued posteriorly to prescu-

tellar bare space and above wing; remainder of scutum with dark scales.

MALE GENITALIA (fig. 21). Segment IX: Anterior middorsal emargination of

tergum broad and shallow, caudal bridge connecting tergal lobes broad; tergal lobes

about as broad as high, with 1-8 setae. Sidepiece: Length about 3.5-4.0 times med-

ian width; tergal surface with setae evenly distributed, becoming longer near level of

base of apical tergomesal lobe, and 2-4 subapicotergal setae; sternomesal margin

with about 40-50 long setae; apical tergomesal lobe well developed; basal tergo-

mesal lobe well developed, triangular with anterior side longer and setae arising

from posterior side, with about 10-20 setae. Claspette: Stem uniformly narrow and

broadly curved dorsad; filament slightly longer than stem, narrow at base and ex-

panded near middle with large retrorse process and several elongate spicules basad

of retrorse process, and tapered gradually to recurved tip. Clasper: Spiniform

length about 10-14 times its greatest width. Phallosome: Aedeagus ovate to slight-

ly pyriform.

PUPA (fig. 21). Abdomen: 3.05 mm. Trumpet: 0.40 mm. Paddle: 0.70 mm.

Cephalothorax: Weakly pigmented, slightly darker dorsally. Seta 10-C usually 6-8

(4-10) branched. Abdomen: Weakly pigmented, slightly darker anteriorly and near

base of most segments. Seta 5-IV,V shorter than or subequal in length to length of

succeeding tergum, usually double (1-4b); seta 5-VI about half length of succeeding

tergum, double (1-2b); seta 5-VII about 1/3 length of seta 3-VII and usually 4-6

branched. Seta 6-VI double or triple (1-4b).

LARVA (fig. 22). Head: 0.85 mm. Siphon: 0.75 mm. Anal Saddle: 0.30 mm.

Thorax: Spicules of integument sparse, short, length about 1-2 times basal diame-

ter. Seta 3-P single. Seta 7-P double (1-3b). Abdomen: Seta 1-IV,V single (1-3b).

Seta 6-I,II usually double (1-3b); seta 6-IH-VI single. Seta 13-III weakly developed,

multiple, subequal to seta 13-II; seta 13-IV,V strongly developed, single. Segment

VIII: Comb scales moderate in size, with slightly differentiated median spinule,

Arnell: Scapularis Group 41

about 20 (16-28) in number, in irregular double to triple row. Siphon: Index about

2.1 to 2.8. Pecten with teeth evenly spaced, extending to middle of siphon. Seta

1-S inserted slightly distad of pecten. Anal Segment: Ventral brush (4-X) with 8

pairs (rarely 9 pairs) of setal tufts, the most proximal tuft occasionally arising from

within or on margin of saddle.

SYSTEMATICS. Aedes tortilis can be separated from the other species of the

Scapularis Group by the following characters: in the female, indistinguishable from

auratus, the scutum has a large patch of golden to tan scales anteriorly; in the male

genitalia, generally indistinguishable from bogotanus, trivittatus, condolescens, eu-

plocamus and patersoni, (1) the sidepiece length is about 3.5 to 4 times median

width, (2) the claspette filament is slightly longer than the stem and has a large

retrorse process with several elongate spicules basad of the process, the spicules sel-

dom extending basad of the basal 0.2 of the filament. (3) the sternomesal margin of

the sidepiece has 40 to 50 long setae between the basal tergomesal lobe and the

clasper, (4) the claspette stem is broadly curved dorsad and not expanded distally,

(5) the apical tergomesal lobe of the sidepiece is well developed, and (6) the spini-

form length is about 12 to 14 times its greatest width; in the larva (1) seta 13-III is

weakly developed, multiple and subequal to 13-II, (2) seta 6-I,II is usually double,

and (3) the spicules of the thoracic and abdominal integument are sparse and short,

length only | to 2 times basal diameter.

Howard, Dyar and Knab (1917:802-810) considered as geographically isolated

distinct species plutocraticus from the Bahama Islands, bracteatus from Cuba, tor-

tilis from Jamaica and balteatus from Hispaniola, and Dyar (1922a:54-56) treated

the above populations, including virginensis from the Virgin Islands and Lesser

Antilles as a complex of subspecies, although in his later treatment (1928: 169-170)

he considered these to be a single species. I have seen usually large numbers of

specimens of all stages of tortilis from most of the insular populations and I am in

agreement with Dyar, considering it to be a single species with no more than mod-

erate variation. However, I have seen only females from Florida, the Bahamas, and

Cuba, and these specimens exhibit more variation in the color of the anterior scutal

scales than do specimens from other Antillean islands, the scutal scales occasionally

being concolorous dark brown. Lack of other stages from these areas precludes de-

termination of possible variation coinciding with that of the female.

Aedes tortilis also shows some variability in the number of white scales above the

paratergite and the number of subspiracular scales in the female, and, in the larva,

very slight differences in the density of the thoracic and abdominal spicules; how-

ever, these differences appear to be random and are not correlated with the different

insular populations. The variation in the anterior scutal scale coloration, as men-

tioned above, may be clinal, since it occurs more frequently in the northern pcpula-

tions.

BIONOMICS. Aedes tortilis typically breeds in temporary or semipermanent

freshwater ground pools formed by rain or stream overflow, although it is often

found near the margin of permanent lakes and marshes and in brackish water ground

pools in mangrove swamps. It has been taken from crabholes in Puerto Rico in as-

sociation with Culex (C.) habilitator and Deinocerites magnus, from pools in block-

ed and intermittent streams, from animal tracks and potholes in marshes and from

rockholes. Females bite man regularly and are commonly taken in animal-baited

traps and light traps.

DISTRIBUTION (fig. 1). Aedes tortilis is known from southern Florida, the Ba-

hama Islands, Cuba, Grand Cayman Island, the Greater Antilles and the Lesser Antil-

42 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

les as far south as St. Lucia. Records of tortilis from Mexico and Guatemala by

Martini (1935:54) are undoubtedly in error. Material examined: 1629 specimens;

183 males, 675 females, 324 pupae, 447 larvae; 161 individual rearings (72 larval,

66 pupal, 23 incomplete).

ANTIGUA. St. George Parish: Emerson’s Pond (E of Sea View Farm), 60 m, 15 Sept 1965, T.

Aitken, R. Martinez and A. Guerra (ANT 33), 8 F [UCLA]. Fitches Creek, nr. Piggotts, 7 m, 23

Sept 1965, R. Martinez and A. Guerra (ANT 87), 1 F [UCLA]. St. John Parish: Bendals, nr. St.

John’s, 30 m, 22 Sept 1965, R. Martinez and A. Guerra (ANT 84), 1 pM (100) [UCLA]. Stuart’s

Pasture, nr. Potters, 22 m, 16 Sept 1965, R. Martinez and A. Guerra (ANT 54), 9 F [UCLA];

same data (ANT 56), 5 F [UCLA]; same locality and collectors, 20 Sept 1965 (ANT 77), 14 F

[UCLA]; same data (ANT 78), 7 F [UCLA]; same locality and collectors, 21 Sept 1965 (ANT

81), 22 F [UCLA]. St. Mary Parish: Wallings Reservoir, nr. Sweets, 150 m, 15 Sept 1965, T.

Aitken, R. Martinez and A. Guerra (ANT 46), 1 lpM (30), 2 IpF (31,36), 1 pF (101), 1 lp G3),

11 M, 19 F, 30 p, 3 P, 21, 7 L [UCLA]; same data (ANT 47), 11 M, 9 F [UCLA]; same locality

and collectors, 27 Sept 1965 (ANT 114), 12 L [UCLA] ; same data (ANT 115), 3 L [UCLA]. St.

Paul Parish: Bethesda, 3 m, 13 Sept 1965, R. Martinez and A. Guerra (ANT 15), 1 pM (101), 1

pF (100) [UCLA]; same data (ANT 16), 4 L [UCLA]. Brooks Estate, nr. Liberta, 23 Sept 1965,

R. Martinez and A. Guerra (ANT 92), 3 L [UCLA]. Matthews, nr. All Saints, 60 m, 15 Sept

1965, T. Aitken, R. Martinez and A. Guerra (ANT 44), 2 F [UCLA]; same locality, 29 Sept

1965, R. Martinez and A. Guerra (ANT 120), 2 F [UCLA]; same data (ANT 122), 1 F [UCLA].

All Saints, 60 m, 15 Sept 1965, T. Aitken, R. Martinez and A. Guerra (ANT 38), 1 F [UCLA].

BAHAMA ISLANDS. Andros Island, 21 June 1903, T. Coffin, 2 F [USNM]; same locality

and collector, 26 June 1903, 2 F [USNM]. Eleuthera Island, Powell Point, 1903, T. Coffin, 3 F

[USNM]. Eleuthera Island, Tarpon [Tarpum] Bay, 1903, T. Coffin, 4 F [USNM]. New Provi-

dence Island, Nassau, 21 June 1903, T. Coffin, 1 F (plutocraticus holotype) [USNM] ; same local-

ity and collector, 23 June 1903, 36 F [USNM]. New Providence Island, Winton, Yamacraw

Beach, 5 Aug 1972, T. Rogers (BAH 48-1), 1 F, E [UCLA]. San Salvador Island, 1903, T. Coffin,

6 F [USNM].

CAICOS ISLANDS. [?] Ft. George Cay, 24 July 1930, Bishopp, 1 F [USNM].

CUBA. Havana: Havana, 14 May 1902, J. Taylor, 1 F [USNM]; same locality and collector,

1 Nov 1902, 1 F [USNM]; same data, 1 F (bracteatus holotype) [USNM]; same locality and col-

lector, 28 Oct 1903, 11 (habanicus lectotype) [USNM]; same locality and collector, 10 Nov 1903,

1 F [USNM]. Havana, 1905, P. Serre, 1 M [USNM]. San Antonio de los Banos, J. Pazos, 8 F

[USNM]. Las Villas: Rosario Cay [Cayo del Rosario], 19 Sept 1930, Bishopp, 1 F [USNM].

Oriente: Cayamas, E. Schwarz, 14 F [USNM]. Guantanamo Naval Base, 27 Oct 1953, K. Knight

(378), 1 F [UCLA]; same locality and collector, 27 Oct 1953 (450), 2 F [UCLA]; same data

(452) 1 F [UCLA]. Pinar del Rio: Mariel, J. Pazos, 2 F [USNM]. Locality unspecified: 1914,

J. Pazos, 23 F [USNM].

DOMINICA. Roseau, 15 km N, Clarke Hall Estate, 7 June 1966, G. Steyskal, 1 F [USNM].

Locality unspecified, 1927, A. Carment, 5 M, 3 F [BM].

DOMINICAN REPUBLIC. Dajabon: Dajabon (2 km N), 40m, 11 July 1971, T. Rogers (RDO

114), 1 1lpF (15), 11P (12) [UCLA]. Distrito Nacional: Santo Domingo, Aug 1905, A. Busck, 4 F

[USNM]; same data, 1 F (balteatus holotype) [USNM]. Santo Domingo, Finca Engombe, 20 m,

20 Aug 1971, T. Rogers (RDO 252), 1 IpM (74), 2 IpF (52,53) [UCLA]; same locality and col-

lector, 28 Aug 1971 (RDO 266), 2 F [UCLA]; same data (RDO 267), 1 M, 3 F, 3 L [UCLA];

same data (RDO 268), 1 IpM (49), 1 IpF (40), 1 L [UCLA]; same locality and collector, 29 Aug

1971 (RDO 270), 1 M [UCLA]; same data (RDO 272), 1 pF (111), 5 M, 8 F, 1 L [UCLA]; same

data (RDO 273), § F [UCLA]; same locality and collector, 30 Aug 1971 (RDO 276), 1 F,E

[UCLA] ; same locality and collector, 31 Aug 1971 (RDO 280), 4 F [UCLA]; same locality and

collector, 6 Sept 1971 (RDO 288), 1 M, 1 F [UCLA]; same locality and collector, 13 Sept 1971

(RDO 296), 1 lp (26), 1 L [UCLA]. Duarte: San Francisco de Macoris, 75 m, 17 July 1968, D.

Watson (RDO 7), 2 F [UCLA]. La Vega: La Vega, 75 m, 25 July 1968, D. Watson (RDO 24),

3 F [UCLA]. Peravia: Los Anones, San Jose de Ocoa, 960 m, 23 July 1968, D. Watson (RDO

22), 1 F [UCLA]. Samana: [Santa Barbara de] Samana, 2 m, 27 June 1971, J. Belkin and T.

Arnell: Scapularis Group 43

Rogers (RDO 49), 1 F [UCLA]. Sanchez, 4 Apr 1949, J. Brennan, 13 F [USNM]. Sanchez

Ramirez: Cotui, 55 m, 20 July 1968, D. Watson (RDO 18), 1 F [UCLA]. Valverde: Mao, 400

m, 14 Aug 1971, T. Rogers and J. Gomes (RDO 223), 1 F [UCLA].

FRENCH WEST INDIES. GUADELOUPE. Basse-Terre: Baie Mahault, 21 Oct 1973, G.

Cornely (FWI 955), 2 L [UCLA]; same data (FWI 957), 10 L [UCLA]. Goyave, 17 Feb 1974, G.

Cornely (FWI 1011), 1 P, 3 L [UCLA]. Petit Bourg, 5 Mar 1972, G. Cornely (FWI 889), 1 pM

(14), 1 pF (16) [UCLA]; same data (FWI 890), 8 L [UCLA]. Grande-Terre: Abymes, 20 Oct

1973, G. Cornely (FWI 951), 9 L [UCLA]; same data (FWI 952), 21 L [UCLA] ; same locality

and collector, 27 Jan 1974 (FWI 996), 56 L [UCLA]. Gosier, 50 m, 3 Mar 1974, G. Cornely

(FWI 1016), 27 L [UCLA]; same data (FWI 1017), 27 L [UCLA]. St. Francis, 3 Feb 1974, G.

Cornely (FWI 1000), 8 L [UCLA] ; same data (FWI 1002), 12 P, 1 L [UCLA].

FRENCH WEST INDIES. MARIE GALANTE. St. Louis, 3 m, 15 Oct 1965, P. Fauran (FWI

265), 5 IpM (11-14,17), 5 IpF (10,15,16,18,19),..1 lp, 6 P, 6 1, 20 L [UCLA]; same data (FWI

268), 1 lpM (10), 1 IpF (11) [UCLA].

GRAND CAYMAN ISLAND. 16 Aug 1964, E. Gerberg (CAY 7), 1 F [UCLA]; 13 Aug 1966,

M. Giglioli (CAY 11), 3 F [UCLA]; July 1966, M. Giglioli (CAY 11A), 20 F [UCLA]; 13 Aug

1966, M. Giglioli (CAY 12), 1 F [UCLA] ;9 Sept 1968, J. Belkin and E. Gerberg (CAY 110,111),

15 F [UCLA].

HAITI. Nord: Bayeaux, 12 Sept [1931] (HAR 27), 3 F [UCLA]; same locality, 13 Sept

(HAR 29), 1 F [UCLA]. Roche Platte, 6 Oct 1925, 1 F [UCLA]. Nord-Ouest: Port de Paix (La

Pointe “House of Hope’’), 7 Oct 1970, J. Porter (HAT 114), 1 F [UCLA]; same locality and col-

lector, 8 Oct 1970 (HAT 121), 21 F [UCLA]. Ouest: Godet, 300 m, 6 July 1968, D. Watson

(HAT 12), 1 F [UCLA]. Port-au-Prince, 14 Sept 1930 (HAR 20), 1 F [UCLA]. Port-au-Prince

10 Oct 1970, M. Solis (HAT 133), 1 F [UCLA]. Sud: Trou Zombi, 25 May 1932, S. Cook, 2 F

[USNM]. Locality unspecified: 23-May 1931 (HAC 6), 3 F [UCLA]; 27 May 1931 (HAC 7),

11 F [UCLA]; 4 June 1931 (HAC 8), 1 M, 4 F [UCLA]; 2 July 1931, S. Cook (HAC 9), 12 F

[UCLA]; 9 July 1931 (HAC 10), 10 F [UCLA].

JAMAICA. Clarendon: Milk River Bath, 0 m, 15 Aug 1967, W. Page (JA 730), 2 L [UCLA].

Parnassus, 10 m, 28 July 1967, W. Page (JA 863), 2 pF (38,112), 11, 1 L [UCLA]. Kingston and

St. Andrew: Hill Gardens, July 1924, G. Strathaim, 1 F [USNM]. Kingston, 20 Aug 1903, M.

Grabham, 5 F [BM]; same data, 1 F (tortilis lectotype) [BM]. Liguana, 3 June 1931, Kisliak, 1 F

[USNM]. St. Andrew: Ferry, 6 Sept 1966, D. Watson (JA 610), 5 F [UCLA]. Ferry,0m, 18

Sept 1967, W. Page (JA 796), 2 lpM (19,21), 8 IpF (11,13,15,16,18,23,24,26), 9 pM (90,92,100,

103,104,106,109,110,114), 4 pF (93,94,105,113), 1 lp (10), 3 IP (12,17,22), 1 M, 1 P, 33 L

[UCLA] ; same locality and collector, 17 Sept 1967 (JA 799), 1 F,2 p,11 [UCLA]. Ferry, 15 m,

9 Aug 1967, J. Belkin and W. Page (JA 896), 1 pM (114), 1 pF (100), 19 M, 16 F,4 L [UCLA].

St. Ann: Delight, 200 m, 25 Aug 1967, J. Belkin and W. Page (JA 757), 1 lpF (113) [UCLA].

St. Catherine: Bog Walk, 90 m, 4 Aug 1968, R. Hochman (JA 913), 4 F [UCLA] ; same locality

and collector, 5 Aug 1968 (JA 914), 2 F [UCLA]. St. Elizabeth: Luana, 0 m, 10 Sept 1965, W.

Page (JA 357), 1 F [UCLA]. St. Thomas: Amity Hall, 11 Dec 1962, T. Aitken (JA 104), 1 F

[UCLA]. Cheswick, 22 Oct 1964 (JA 161), 1 lpF (15), 1 P, 1 L [UCLA]. Golden Grove, 1

Feb 1965 (JA 197), 2 L [UCLA]. Holland Bay, 0 m, 11 Aug 1964, H. Tucker and G. Hamilton

(JA 97), 3 lpM (11,12,15), 1 IM (13), 1 pM (100), 2 lp (10,16) [UCLA]. Holland Bay; 5 Oct

1964 (JA 143), 1 pM(5) [UCLA]. Holland Bay, 23 Oct 1964 (JA 159), 1 L [UCLA]. New Pera,

5 Dec 1962, T. Aitken (JA 19), 1 F [UCLA]. Westmoreland: Crab Pond Bay, 14 Sept 1967 (JA

790), 3 F, 1 L [UCLA].

MONTSERRAT. St. Anthony Parish: Plymouth, Elberton Estate, 60 m, 3 Oct 1965, T.

Aitken, R. Martinez and A. Guerra (MNT 1), 1 lpM (10), 1 M, 1 p [UCLA]; same data (MNT 3),

1 IpM (11), 4 IpF (21-24), 1 pF (102), 1 lp (10), 2 1P (25,26), 1 F [UCLA] ; same data (MNT 4),

2 IpM (10,11), 1 pM (100), 1 IP [UCLA]; same data (MNT 5), 2 IpM (10,14), 5 IpF (13,15-18),

SM, 6 F, 11 p, 111, 1 L [UCLA]; same data (MNT 7), 1 F [UCLA]; same locality, 13 Oct 1966,

R. Martinez and A. Guerra (MNT 78), 2 F [UCLA]. Plymouth, Richmond Estate, 60 m, 15 Oct

1966, R. Martinez and A. Guerra (MNT 80), 1 F [UCLA]. St. Peter Parish: Gerald’s Village, 150

m, 17 Oct 1966, R. Martinez and A. Guerra (MNT 109), 2 IpF (10,12), 10 M,6F, 16 p,716L

44 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

[UCLA]. St. John’s Village, 230 m, 17 Oct 1966, R. Martinez and A. Guerra (MNT 101), 1 IP

(10) [UCLA]; same data (MNT 104), 1 IpF (10) [UCLA]; same data (MNT 105), 1 IP (10)

[UCLA]. Sweeney’s Village, 75 m, 17 Oct 1966, R. Martinez and A. Guerra (MNT 112), 2 M,

1 F,4 p, 31 [UCLA]; same data (MNT 113), 1 IpF (11), 7 M, 13 F, 20 p, 121,3 L [UCLA].

PUERTO RICO. Dorado: Levittown, 0 m, 2 Sept 1970, J. Belkin and M. Nelson (PR 160),

1 pF (101) [UCLA]. Playa Dorado-Sardinera, 0 m, 2 Sept 1970, J. Belkin and M. Nelson (PR

151), 1 pM (103), 5 pF (100-102,104,105) [UCLA]; same data (PR 155), 3 lpF (10,12,13), 2

pM (101,106), 9 pF (100,102-105,107-110), 5M, 1 F,4 P, 11, 1 L [UCLA]; same data (PR 156),

2 IpM (10,13), 1 IP (12), 1 M, 1 p, 2 L [UCLA]. Gurabo: Gurabo, June 1943, H. Pratt, 5 L

[UCLA] ; same locality and collector, 18 Oct 1943, 5 L [UCLA]. Mayaguez: Mayaguez, 23

Aug 1935, G. Tulloch, 1 F [UCLA]; same locality and collector, 4 Oct-4 Nov 1935, 43 F

[USNM] ; same locality, 20 m, 25 Aug 1970, J. Belkin (PR 116), 1 M, 1 p [UCLA]. San Juan:

Catano, 5 May 1943, H. Pratt, 1 F [USNM]. Isla Verde, July-Aug 1956, 1 M [UCLA]. San Juan,

Om, 1 Sept 1970, J. Belkin and M. Nelson (PR 149), 2 IpM (10,13), 6 IpF (11,12,14-17), 9 pM

(100,102,103,107-112), 4 pF (104-106,113), 26 M, 27 F, 58 p, 1 1 [UCLA]; same locality, May

1914, through C. Ludlow, 1 F [USNM]. Vega Baja: Camp Tortuguera, 21 Aug 1942, T. Aitken

and H. Pratt (PRA 44), 3 L [UCLA].

ST. LUCIA. Anse La Raye, 12 m, 22 July 1964, R. Martinez and A. Guerra (LU 74), 1 lpM

(10) [UCLA]. Choc Swamp, 0 m, 17 July 1964, R. Martinez and A. Guerra (LU 50), 4 F

[UCLA] ; same locality and collectors, 19 July 1964 (LU 54), 2 F [UCLA]. Grande Anse, 20

July 1964, R. Martinez and A. Guerra (LU 66), 1 IP (11), 15 L [UCLA]. Mahout, 210 m, 28

July 1964, A. Guerra (LU 106), 2 F [UCLA]. Reduit Swamp, 18 July 1964, A. Guerra (LU 53),

3 F [UCLA]; same locality and collector, 19 July 1964 (LU 55), 1 F [UCLA]; same data (LU

56), 3 F [UCLA]. Sapphire Estate, 15 m, 6 Aug 1964, R. Martinez and A. Guerra (LU 162), 5 F

[UCLA]. Sapphire Swamp, 45 m, 24 July 1964, R. Martinez and A. Guerra (LU 90), 1 M

[UCLA] ; same data (LU 92), 1 lpF (20) [UCLA]; same data (LU 93), 3 M, 11 F [UCLA]; same

locality and collectors, 29 July 1964 (LU 122), 1 pF (101) [UCLA]. Vieux Fort, 3 m, 24 July

1964, R. Martinez and A. Guerra (LU 97), 1 F [UCLA]. Volet Swamp, 5 m, 28 July 1964, A.

Guerra (LU 107), 9 F [UCLA]; same data (LU 108), 9 F [UCLA]; same locality and collector,

29 July 1964 (LU 117), 12 F [UCLA].

UNITED STATES. Florida: Key West, 28 Aug 1945, C. Spencer, 1 F [USNM]. [?] Vero

Beach, 14 Mar 1962, Ent. Res. Cent., 1 F [USNM].

VIRGIN ISLANDS. St. Croix: Christiansted, Dec 1936, H. Beatty, 4 M, 2 F [USNM]. St.

Thomas: Nadir, 0 m, 30 Nov 1965, D. Bonnet (VI 2), 1 IlpF (10), 1 1F (11) [UCLA]. Nazareth,

0 m, 30 Nov 1965, D. Bonnet (VI 5), 4 pM (101,103,104,106), 3 pF (100,102,105) [UCLA].

Locality unspecified, Aug 1905, A. Busck, 1 F (tortilis virginensis holotype) [USNM].

Additional Records from the Literature

UNITED STATES. Florida: Fisher Is., Dade Co.; Belle Glade and Jupiter, Palm Beach Co.

(Branch, Logan et al., 1959:159). Clewiston, Hendry Co. (Pritchard, Seabrook and Mulrennan

1947:11).

9. Aedes (O.) auratus Grabham

Figs. 1,23,24

1906. Aedes auratus Grabham, 1906:313-315. TYPE: Lectotype male with genitalia slide

(680827-15), Kingston, Jamaica, 10 Apr 1906, M. Grabham [USNM; selection of Belkin,

Heinemann and Page, 1970:168]. Synonymy with tortilis apparently by Howard, Dyar

and Knab (1917:806); resurrected from synonymy with fortilis by Belkin, Heinemann

and Page (1970:168).

Aedes (Ochlerotatus) auratus of Belkin, Heinemann and Page (1970:168-169).

Aedes auratus of Dyar and Knab (1906b:163); Theobald (1910:598-599).

Arnell: Scapularis Group 45

Aedes (Ochlerotatus) tortilis in part of Dyar (1928:169-170); Edwards (1932:143); Lane (1939:

117; 1953:653-654); Carpenter and La Casse (1955:245-247); Stone, Knight and Starcke

(1959:156); Forattini (1965:369).

Aedes (Ochlerotatus) tortilis tortilis in part of Dyar (1922a:54-55).

Aedes tortilis in part of Howard, Dyar and Knab (1917:806-809); Torre y Callejas, Alayo Delmau

and Calderon Chapman (1961 :67).

FEMALE. Thorax: Anterior of scutum except for fossa with golden to tan

scales.

MALE. Wing: 3.05 mm. Proboscis: 1.95 mm. Forefemur: 1.55 mm. Appar-

ently indistinguishable from fortilis.

MALE GENITALIA (fig. 24). Segment IX: Anterior middorsal emargination of

tergum broad and shallow, caudal bridge connecting tergal lobes broad; tergal lobes

about as broad as high, with 3-6 setae. Sidepiece: Length about 4 times median

width; tergal surface with short setae mesad, longer setae laterad, subapicotergal

setae in row of 5 or 6, separated from more proximal setae of sidepiece by distinct

bare space, and 2 or 3 large setae near base laterally; sternomesal margin with about

40 long setae; apical tergomesal lobe very well developed; basal tergomesal lobe well

developed, large, broadly conical, with about 25-30 setae. Claspette: Stem uni-

formly narrow and broadly curved dorsad; filament slightly longer than stem, nar-

row at base, expanded near middle with large retrorse process and several elongate

spicules basad of retrorse process, and tapered gradually to recurved tip. Clasper:

Spiniform length about 10 times its greatest width. Phallosome: Aedeagus ovate.

PUPA. Unknown.

LARVA (fig. 23). Head: 0.75 mm. Siphon: 0.75 mm. Anal Saddle: 0.35 mm.

Thorax: Spictles of integument sparse, short, length about 1-2 times basal diame-

ter. Seta 3-P single. Seta 7-P double (2-3b). Abdomen: Seta 6-I-VI single. Seta 13-

III weakly developed, multiple, subequal to seta 13-II]. Segment VIII: Comb scales

moderate in size, with slightly differentiated median spinule, about 20 (17-27) in

number, in irregular double or triple row. Siphon: Index about 2.35-2.50. Pecten

with teeth evenly spaced, extending slightly beyond middle of siphon. Seta 1-S in-

serted at level of most distal pecten tooth. Anal Segment: Ventral brush (4-X) with

8 pairs (occasionally 7 pairs) of setal tufts.

SYSTEMATICS. Aedes auratus can be separated from the other species of the

Scapularis Group. by the following characters: in the female, indistinguishable from

tortilis, the scutum has a large patch of golden to tan scales anteriorly; in the male

genitalia (1) the tergal surface of the sidepiece has a distinct bare space between the

subapicotergal setae and the more proximal, short tergal setae, the subapicotergal

setae being in a conspicuous oblique row of 5 or 6 and (2) the setae of the basal

tergomesal lobe number 25 to 30; in the larva (1) seta 6-I,II is single, (2) seta 13-IH

is weakly developed and subequal to 13-II, (3) the ventral brush has 8 pairs of setal

tufts, and (4) the spicules of the thoracic and abdominal integument are sparse and

short, length only 1 to 2 times basal diameter.

The specimens from Grabham’s original collection of auratus constitute all the

known material of this species. The only female, in very poor condition, is impos-

sible to distinguish from tortilis; however, the male genitalia and larva are very well

marked as pointed out above. I am in agreement with Belkin, Heinemann and Page

(1970:167-168) in their recognition of auratus as a full species and their observation

that it is probably a relict population of an early derivative of the tortilis stock.

BIONOMICS. Larvae of auratus were collected by Grabham from temporary

pools formed by seasonal rains in the vicinity of Kingston. No specimens of auratus

46 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

have been taken since the original collection was made in 1906.

_ DISTRIBUTION (fig. 1). Aedes auratus is known only from the type locality,

Kingston, Jamaica. Material examined: 9 specimens; 3 males, 1 female, 5 larvae.

JAMAICA. Kingston and St. Andrew: Kingston, 10 Apr 1906, M. Grabham, 2 M, 1 F,5 L

[UCLA, USNM] ; same data, 1 M (auratus lectotype) [USNM].

INFIRMATUS SUBGROUP

FEMALES. Head: Occipital dorsolateral dark scale patch conspicuous to nearly

absent. Palpus 3 or 4 segmented. Thorax: Scutum with broad anterior stripe of

entirely silvery white scales or silvery white scales divided by narrow acrostichal line

of tan scales; sides of stripe more or less parallel and about midway between dorso-

central line and scutal margin; stripe terminating at about level of posterior end of

paratergite, but often with narrow posterior extensions along dorsocentral line to

scutellum and along midline meeting and encircling prescutellar bare space. Remain-

der of scutum with dark brown scales. Acrostichal, anterior dorsocentral, and pos-

terior fossal setae absent. Legs: Knee spots of white scales present posteriorly on

apices of femora. Tibiae and basal tarsal segments with conspicuous white streak

posteriorly. Hindtibia and hindtarsus dark scaled anteriorly. Claws of hindleg

simple or with submedian tooth. Wing: Scales entirely dark. Abdomen: Terga

II-VII with large basolateral white patches; entirely dark scaled dorsally or with

basal white bands or basomedian white patches.

MALES. Thorax: Anterior scutal scale patch of light scales generally similar to

female. Legs: Anterior claw of foreleg and midleg considerably enlarged, with

blunt submedian tooth and acute basal tooth, posterior claw of foreleg with acute

basal tooth; posterior claw of midleg small, with acute subbasal tooth. Abdomen:

Terga with or without basal bands of white scales.

DISCUSSION. The Infirmatus Subgroup is defined by the female scutal scale pat-

tern, a broad anterior stripe of silvery white scales, rarely divided by a narrow acro-

stichal line of tan scales, and by the anterior surface of the hindtibia being without

light scales.

The subgroup is composed of 5 species, condolescens, euplocamus, patersoni, ray-

mondi and infirmatus, allopatric with one exception, and distributed in the south-

eastern United States, Cuba, the Bahama Islands, Central America, northern South

America, southern Bolivia and northern Argentina (see fig. 3), a distribution pattern

generally comparable to that of the Trivittatus Subgroup.

The distribution of the Infirmatus Subgroup suggests an origin in the Central

American-Greater Antillean region, prior to or concomitant with a probable land

bridge or archipelago connecting these areas.

10. Aedes (O.) condolescens Dyar and Knab

Figs. 3,24

1907. Aedes condolescens Dyar and Knab, 1907:11. TYPE: Holotype female (10), Nassau,

Bahama Islands, 24 June 1903, T. H. Coffin [USNM, 10248, see Stone and Knight, 1956:

216].

Aedes (Ochlerotatus) condolescens of Dyar (1928:165-166); Edwards (1932: 142); Martini (1935:

54-55, in part); Lane (1939:107-108; 1953:656); Perez Vigueras (1956:278-285); Stone,

Arnell: Scapularis Group 47

Knight and Starcke (1959:143); Forattini (1965:357-358); Montshadsky and Garcia Avila

(1966:29); Porter (1967:37,39,40); Belkin, Heinemann and Page (1970:151,164).

Aedes (Heteronycha) condolescens of Dyar (1920:105).

Aedes condolescens of Johnson (1908:71); Pazos (1908:422); Howard, Dyar and Knab (1917:

789-791); Dyar (1918a:77; 1924a:119); Torre y Callejas, Alayo Delmau and Calderon Chap-

man (1961:68).

Aedes (Ochlerotatus) scapularis condolescens of Dyar (1922a:59-60); Bonne and Bonne-Wepster

(1925:388-390).

FEMALE (fig. 24). Wing: 3.00 mm. Proboscis: 1.80 mm. Forefemur: 1.40

mm. Abdomen: 2.30 mm. Head: Palpus 3 segmented. Thorax: Broad anterior

scutal stripe entirely silvery white. Legs: Knee spots of white scales absent pos-

teriorly on apices of femora. Claws of hindleg without submedian tooth. Abdo-

men: Terga [II-VI with broad basal bands of white scales.

MALE (fig. 24). Wing: 3.15 mm. Proboscis: 2.20 mm. Forefemur: 1.16 mm.

Abdomen: 3.05 mm. Abdomen: Terga with conspicuous basal bands of white

scales.

MALE GENITALIA (fig. 24). Segment IX: Anterior middorsal emargination of

tergum broad and shallow, caudal bridge connecting tergal lobes broad; tergal lobes

about as broad as high, with 5-7 setae. Sidepiece: Length about 3.5 times median

width; tergal surface with setae uniformly relatively long and 2-3 subapicotergal

setae; sternomesal margin with 40-50 long setae; apical tergomesal lobe well develop-

ed; basal tergomesal lobe well developed, triangular, with 15-20 undifferentiated

setae, distal setae longer than more proximal ones. Claspette: Stem uniformly nar-

row and broadly curved dorsad; filament subequal in length to stem, narrow at base,

abruptly expanded near middle with large retrorse process and several elongate spic-

ules basad of retrorse process, and tapered to recurved tip. Clasper: Spiniform

length about 12 times its greatest width. Phallosome: Aedeagus ovate.

PUPA, LARVA. Unknown.

SYSTEMATICS. I have seen condolescens only in the adult and it can be separ-

ated from the other species of the Scapularis Group by the following characters: in

the female (1) the scutum has a broad anterior stripe of silvery white scales, (2)

there are no knee spots of white scales on the apices of the femora, and (3) broad

basal bands of white scales are present on abdominal terga II-VI; in the male geni-

talia, generally indistinguishable from bogotanus, trivittatus, tortilis, euplocamus

and patersoni, (1) the sidepiece length is about 3.5 times median width, (2) the

claspette filament is subequal in length to the stem and has a large retrorse process

with several elongate spicules basad of the process, the spicules not extending basad

of the basal 0.2 of the filament, (3) the sternomesal margin of the sidepiece has 40

to 50 long setae between the basal tergomesal lobe and the clasper, (4) the claspette

stem is broadly curved dorsad and not expanded distally, (5) the apical tergomesal

lobe of the sidepiece is well developed, and (6) the spiniform length is about 12

times its greatest width.

This species has been reported from Puerto Rico, the Lesser Antilles, Mexico,

Guatemala, Colombia, Peru, Paraguay and Argentina. Since it is actually known

only from the Bahama Islands, Cuba and Grand Cayman, the erroneous distribu-

tions undoubtedly are either taxonomic misinterpretations or misidentifications of

other Scapularis Group species with similar adult ornamentation, tortilis, euplocam-

us, patersoni or scapularis.

Aedes condolescens appears to be more closely allied to the Central and South

48 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

American members of the subgroup, euplocamus and patersoni, than to the geo-

graphically adjacent infirmatus, which occurs in the southeastern United States,

commonly in southern Florida.

BIONOMICS. Little is known of the bionomics of condolescens. Adults have

been taken biting man in deep to partial shade at 1630-1700 hours on Grand Cay-

man Island.

DISTRIBUTION (fig. 3). Aedes condolescens is known from the Bahama Islands,

Cuba and the Cayman Islands. Material examined: 39 specimens; 2 males, 37

females.

BAHAMAS. Andros Island, 1903, T. Coffin, 4 F [USNM]. Eleuthera Island, Tarpon [Tar-

pum] Bay, 1903, T. Coffin, 1 F [USNM]. Long Island, 1903, T. Coffin, 1 F [USNM]. New

Providence Island, Nassau, 22 June 1903, T. Coffin, 7 F [USNM]; same locality and collector,

24 June 1903, 1 F (condolescens holotype) [USNM]. San Salvador Island, 1903, T. Coffin, 4 F

[USNM].

CUBA. Havana: Havana, 1909, P. Serre, 2 M [USNM]. Havana, 18 Dec 1929, 1 F [|USNM].

San Antonio de los Banos, J. Pazos, 1 F [USNM]. Oriente: Cayamas, 7 June 1909, E. Schwarz,

4 F [USNM]. Guantanamo Bay, U.S. Naval Base, 8 July 1953, K. Knight, 1 F [UCLA]. Locality

unspecified: 1914, J. Pazos, 7 F [USNM].

GRAND CAYMAN ISLAND. 9 Sept 1968, J. Belkin, E. Gerberg (CAY 111), 4 F [UCLA].

LITTLE CAYMAN ISLAND. 13 Sept 1930, H. Peters, 1 F [USNM].

11, Aedes (O.) euplocamus Dyar and Knab

Figs: 3,25;26

1906. Aedes euplocamus Dyar and Knab, 1906a:199. TYPE: Syntypes larvae, Zent, near

Limon, Limon, Costa Rica, F. Knab [NE; see Stone and Knight, 1956:217].

Aedes (Ochlerotatus) euplocamus of Levi-Castillo (1951:383; 1952:262); Vargas (1956:22);

Stone, Knight and Starcke (1959:145); Belkin, Heinemann and Page (1970:164); in part of

Dyar (1925a:144; 1928:167); Edwards (1932:142); Martini (1935:55); Lane (1939:109).

Aedes (Heteronycha) euplocamus of Dyar (1920:105).

Aedes euplocamus of Howard, Dyar and Knab (1917:787-789); Dyar (1918a:77; 1924a:118-119);

Kumm, Komp and Ruiz (1940:400); Kumm and Zuniga (1942:405).

Aedes (Ochlerotatus) scapularis euplocamus of Dyar (1922a:59); Bonne and Bonne-Wepster

(1925:388-390).

Aedes (Ochlerotatus) scapularis of Lane (1951:335); Lane (1953:665-667, in part); Forattini

(1965:331, in part).

FEMALE (fig. 25). Wing: 3.50 mm. Proboscis: 2.20 mm. Forefemur: 1.80

mm. Abdomen: 2.90 mm. Head: Palpus 3 segmented. Thorax: Broad anterior

scutal stripe entirely silvery white. Legs: Knee spots of white scales present pos-

teriorly on apices of femora. Claws of hindleg with submedian tooth. Abdomen:

Terga II-VII entirely dark scaled dorsally or with small basomedian white scale

patches.

MALE (fig. 25). Wing: 3.00 mm. Proboscis: 2.15 mm. Forefemur: 1.55 mm.

Abdomen: 2.95 mm. Abdomen: Terga with basal bands of white scales.

MALE GENITALIA (fig. 25). Segment IX: Anterior middorsal emargination of

tergum broad and shallow, caudal bridge connecting tergal lobes broad; tergal lobes

about as broad as high, with 5-7(3-10) setae. Sidepiece: Length about 4 times med-

ian width; sternomesal margin with about 40-50 long setae; apical tergomesal lobe

moderately developed; basal tergomesal lobe relatively small, triangular with anter-

Arnell: Scapularis Group 49

ior and posterior sides subequal in length although often folded ventrad in slide

preparations, differentiated seta length about 20 times greatest width, with about

20 undifferentiated setae, usually 4 longer setae projecting distad from posterior

edge of lobe. Claspette: Stem broadly curved dorsad and narrowed distally, basal

seta subequal in length to claspette width at origin of seta; filament subequal in

length to stem, narrow at base, abruptly expanded near middle with large retrorse

process and several elongate spicules basad of retrorse process, and tapered to re-

curved tip. Clasper: Spiniform length about 10-14 times its greatest width. Phallo-

some: Aedeagus ovate to slightly pyriform.

PUPA (fig. 25). Abdomen: 2.55 mm. Trumpet: 0.50mm. Paddle: 0.80 mm.

Cephalothorax: Weakly pigmented, dorsum darker. Abdomen: Moderately pig-

mented. Seta 1-VII usually single (1-2b). Seta 3-VII usually triple (1-3b). Seta 5-

IV,V usually considerably longer than length of succeeding tergum; seta 5-VI sub-

equal in length to length of succeeding tergum, single, rarely double. Seta 6-I[II-VI

single, rarely double on segments III, IV, VI.

LARVA (fig. 26). Head: 0.80 mm. Siphon: 0.75 mm. Anal Saddle: 0.35 mm.

Thorax: Spicules of integument moderately numerous and conspicuous, length

about 4-6 times basal diameter. Seta 3-P single. Seta 7-P triple (2-3b). Abdomen:

Seta 1-IV,V usually single (1-3b); seta 1-VII usually double (2-3b). Seta 6-I,II dou-

ble; seta 6-III-VI single. Seta 13-III well developed, single, subequal to seta 13-IV,V.

Segment VIII: Comb scales moderate in size, evenly fringed, 20-24 (13-27) in num-

ber, in irregular double to triple row. Siphon: Index about 2.0 to 2.4. Pecten with

teeth evenly spaced, extending slightly beyond middle of siphon. Seta 1-S inserted

distad of pecten. Anal Segment: Ventral brush (4-X) with 8 pairs of setal tufts.

SYSTEMATICS. Aedes euplocamus can be separated from the other species of

the Scapularis Group by the following characters: in the female (1) the scutum has

a broad anterior stripe of silvery white scales, (2) knee spots of white scales are pres-

ent on the apices of the femora, and (3) the palpus is 3 segmented; in the male geni-

talia, generally indistinguishable from bogotanus, trivittatus, tortilis, condolescens

and patersoni, (1) the sidepiece length is about 4 times median width, (2) the clas-

pette filament is subequal in length to the stem and has a large retrorse process with

several elongate spicules basad of the process, the spicules not extending basad of

the basal 0.2 of the filament, (3) the sternomesal margin of the sidepiece has 40 to

50 long setae between the basal tergomesal lobe and the clasper, (4) the claspette

stem is broadly curved dorsad and not expanded distally, (5) the apical tergomesal

lobe of the sidepiece is moderately developed, and (6) the spiniform length is about

10 to 14 times its greatest width; in the larva (1) seta 1-IV,V is single, (2) seta

13-III is well developed, subequal to 13-IV,V, (3) seta 3-P is single, (4) seta 6-III is

single, and (5) the spicules of the thoracic and abdominal integument are moderate-

ly long, length about 4 to 6 times basal diameter.

Aedes euplocamus can be separated from the closely related patersoni only with

some difficulty in the female, male genitalia and larva. The presence of the rela-

tively large fourth palpal segment in the female of patersoni and its absence in eu-

plocamus is constant in the specimens I have examined, although this character is

variable in other species of the Scapularis Group. Although the differentiated seta

of the basal tergomesal lobe is somewhat longer and thicker in patersoni than in

euplocamus, the male genitalia are separated primarily by the longer basal seta of

the claspette stem of patersoni. The separation of the larva is tenuous, seta 6-III

being usually double though often single in patersoni and always single in euplo-

camus, and the spicules of the integument are longer in patersoni than in euplo-

camus.,

50 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

There is no evidence to indicate how long euplocamus and patersoni have been

geographically isolated; however, evolutionary pressures on neither species have

been sufficient to produce significant and morphological changes between the two.

BIONOMICS. Larvae and pupae of euplocamus have been collected from volcan-

ic streamside rockholes in Costa Rica, in partial shade, the water temporary, the

rockholes without vegetation, and associated with several species of Culex and Uran-

Otaenia. In Venezuela, larvae of euplocamus have been collected from a tire with

larvae of Aedes (S.) aegypti and pupae from a small, temporary ground pool.

Kumm, Komp and Ruiz (1942:405) collected larvae of euplocamus in El Salvador

only once, in a seepage area at about 900 meters elevation containing abundant

algae, although adults were taken often in houses, horse traps, caves and in the

forest during the day.

DISTRIBUTION (fig. 3). Aedes euplocamus is distributed from Veracruz, Mex-

ico through Central America to the Panama Canal Zone, and is also known from

Aragua, Venezuela and Meta, Colombia. There are no reliable distribution records

in the literature other than the localities included in the following list of specimens

examined. Material examined: 186 specimens; 39 males, 73 females, 55 pupae, 19

larvae; 35 individual rearings (7 larval, 26 pupal, 2 incomplete).

COLOMBIA. Meta: Locality uncertain, probably near Villavicencio, 8 Nov 1936 (COR 136),

4 F [UCLA].

COSTA RICA. Alajuela: Ciruelas, 24 Nov 1920, A. Alfaro, 2M, 2 F [USNM]. Ciruelas, 26

July 1921,4 M [USNM]. Rio Alajuela, nr. Atenas, 500 m, 31 July 1971, S. Heinemann and A.

Berrios Arias (CR 308), 1 IpF (10), 1 pF (105), 1 IP (11), 11, 2 L [UCLA]; same data (CR 309),

1 pM (102), 3 pF (100,101,103), 1 IP (10), 1 F, 1 p, 1 L [UCLA]. Guanacaste: El Coco, 19

July 1962, F. Truxal, 1 F [UCLA]. Heredia: Heredia, 1 M [USNM]. Limon: Matina, 21 May

1925, L. Rozeboom (CRR 6), 1 M [UCLA].

EL SALVADOR. San Salvador: San Salvador, F. Figueroa, 1 F [USNM]. Sonsonate: Son-

sonate, F. Knab, 1 M, 3 F [UCLA]. Usulutan: Berlin, Sept 1941, 4 M, 2 F [USNM]. Locality

unspecified: 1 M.

MEXICO. Oaxaca: Almoloya, F. Knab, 6 F [USNM]. Matios Romero (42 km N), 26 July

1963, E. Fisher (MF 13), 1 F [UCLA]. Veracruz: Cordoba, 8 Apr 1908, F. Knab, 1 F [USNM].

PANAMA AND CANAL ZONE. Canal Zone: Ancon, 18 July 1908, A. Jennings, 1 M

[USNM]. Corozal, 24 May 1943, H. Crowell (KO 202C-3), 3 M, 2 F [UCLA]. Corozal, J.

Shropshire, 1 F [USNM]. La Pita, 16 Jan 1922, J. Shropshire, 1 F [USNM]. Pedro Miguel

1916, L. Dunn, 1 M, 2 F [USNM]. Locality unspecified (CZ 49), 1 lpF (14), 3 pM (10,101,

105), 4 pF (104,106,107,109), 1 M, 1 F, 1 P [UCLA]. Panama: La Chorrera, 1 Feb 1945,

R. Arnett and Frick (ASM 400), 4 F [UCLA]. Paitilla Pt., 31 July 1923, H. Dyar and R.

Shannon, 5 M, 5 F [BM, USNM].

VENEZUELA. Aragua: San Jacinto Mil. Res. (nr. Maracay), 550 m, 17 July 1969, J. Valencia

and J. Pulido (VZ 237), 3 M, 12 F, 15 p, 51, 1 L [UCLA]. Turmero, 540 m, 29 Aug 1966, E.

Vasquez (VZ 26), 4 pM (100,104,107,110), 8 pF (101,105,108,109,111-114), 3 p [UCLA]:

same locality and collector, 14 Sept 1966 (VZ 42), 3 lpM (90, 96 98), 3 IpF (93,94), 2 pF (92,

95 [UCLA]. State unknown: Trinidad, 30 Jan 1939 (VZR 141), 1 F [UCLA].

12. Aedes (O.) patersoni Shannon and Del Ponte

Piss: 3527.28

1928. Aedes (Ochlerotatus) patersoni Shannon and Del Ponte, 1928:73-74. TYPE: Lectotype

female, San Pedro de Jujuy, Jujuy, Argentina, 27 Apr 1926, Paterson, Shannon and

Shannon [INM; selection of O. H. Casal in Belkin, Schick and Heinemann, 1968:14].

Synonymy with crinifer apparently by Edwards (1932:142); resurrected from synonymy

with crinifer by Del Ponte, Castro and Garcia (1951:229).

Arnell: Scapularis Group 51

Aedes (Ochlerotatus) patersoni of Del Ponte, Castro and Garcia (1951:229); Ronderos and Garcia

(1963a:35-36); Garcia and Ronderos (1963:29); Stone (1967: 207).

Aedes (Ochlerotatus) crinifer in part of Edwards (1932:142); Lane (1939: 108; 1953:668-669);

Stone, Knight and Starcke (1959:143); Forattini (1965:358).

FEMALE (fig. 27). Wing: 3.75 mm. Proboscis: 2.45 mm. Forefemur: 1.90

mm. Abdomen: 3.25 mm. Head: Palpus 4 segmented, segment 4 relatively large.

Thorax: Broad anterior scutal stripe entirely silvery white. Legs: Knee spots of

white scales present posteriorly on apices of femora. Claws of hindleg with sub-

median tooth. Abdomen: Terga II-VII entirely dark scaled dorsally or with small

basomedian white scale patches.

MALE (fig. 27). Wing: 3.60 mm. Proboscis: 2.40 mm. Forefemur: 1.70 mm.

Abdomen: 3.40 mm. Abdomen: Terga with bands of white scales.

MALE GENITALIA (fig. 27). Essentially asin euplocamus with following excep-

tions: differentiated seta of basal tergomesal lobe of sidepiece somewhat longer and

thicker, length about 12 times greatest width; basal seta of claspette stem usually

about double in length to claspette width at origin of seta.

PUPA (fig. 27). Abdomen: 3.55 mm. Trumpet: 0.60 mm. Paddle: 0.80 mm.

Cephalothorax: Weakly pigmented, dorsum considerably darker. Seta 9-C usually

single (1-2b). Abdomen: Moderately pigmented. Seta 1-VII usually single (1-2b).

Seta 3-VII usually single (1-2b). Seta 5-IV,V usually longer than length of succeed-

ing tergum. Seta 6-III usually double (1-2b); seta 6-IV-VI usually single (1-2b).

LARVA (fig. 28). Head: 0.85 mm. Siphon: 0.80 mm. Anal Saddle: 0.35 mm.

Thorax: Spicules of integument numerous and conspicuous, length about 8-12

times basal diameter. Seta 3-P single. Seta 7-P triple. Abdomen: Seta 1-IV,V single

(1-2b). Seta 6-I,II double; seta 6-III double (1-2b); seta 6-IV single (1-2b); seta 6-V,

VI single. Seta 13-III well developed, single, subequal to seta 13-IV,V which are

single. Segment VIII: Comb scales moderate in size, evenly fringed, about 20(18-

23) in number, in irregular double to triple row. Siphon: Index about 2.1-2.3.

Pecten with teeth evenly spaced, extending slightly beyond middle of siphon. Seta

1-S inserted distad of pecten. Anal Segment: Ventral brush (4-X) with 8 pairs of

setal tufts.

SYSTEMATICS. Aedes patersoni can be separated from the other species of the

Scapularis Group by the following characters: in the female (1) the scutum has a

broad anterior stripe of silvery white scales, (2) knee spots of white scales are pres-

ent on the apices of the femora, and (3) the palpus is 4-segmented with segment 4

relatively large; in the male genitalia, generally indistinguishable from bogotanus,

trivittatus, tortilis, condolescens and euplocamus, (1) the sidepiece length is about

4 times median width, (2) the claspette filament is subequal in length to the stem

and has a large retrorse process with several elongate spicules basad of the process,

the spicules not extending basad of the basal 0.2 of the filament, (3) the sterno-

mesal margin of the sidepiece has 40 to 50 long setae between the basal tergomesal

lobe and the clasper, (4) the claspette stem is broadly curved dorsad and not ex-

panded distally, (5) the apical tergomesal lobe of the sidepiece is moderately de-

veloped, and (6) the spiniform length is about 10 to 14 times its greatest width;

in the larva (1) seta 1-IV,V is single, (2) seta 13-III is well developed, subequal to.

13-IV,V, (3) seta 3-P is single, (4) seta 6-Iil is usually double, and (5) the spicules of

the thoracic and abdominal integument are long, length 8 to 12 times basal diameter.

This species is morphologically very similar in all stages to euplocamus, to which

it is undoubtedly closely related. This is discussed above under euplocamus.

52 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

BIONOMICS. Paterson, Shannon and Shannon reared the type series of patersoni

from larvae collected from a heavily vegetated temporary rain pool which also con-

tained larvae of Psorophora (J.) ferox and Culex (Lutzia) sp. Data from records of

2 UCLA collections of patersoni from Argentina indicate that larvae were obtained

from a semipermanent pond associated with scapularis and from a temporary road-

side pool containing no vegetation and associated with P. ferox.

DISTRIBUTION (fig. 3).- Aedes patersoni is known from extreme northern Ar-

gentina and southern Bolivia. Material examined: 433 specimens; 64 males, 22

females, 233 pupae, 114 larvae; 49 individual rearings (33 larval, 15 pupal, 1 in-

complete).

ARGENTINA. Jujuy: San Pedro de Jujuy, 27 Apr 1926, Shannon and Shannon, 1 M, 2 F

[USNM]. Salta: Gen. Enrique Mosconi (Campamento Vespucio), 29 May 1969, Fernandez,

Garcia and Casal (ARG 742), 2 IpM (11,14), 3 pM (106,108,111), 1 pF (100), 1 F, 30 L [UCLA];

same data (743), 1 F (18) [INM]. Tablillas (nr. Vespucio), 3 June 1969, Fernandez (ARG 755),

2 IpM (25,29), 7 IpF (20-24,26,28), 3 pM (101,104,112), 8 pF (100,102,103,105,106,108,109,

113), 1 lp (27), 29 M, 22 F, 178 p, 6 P, 50 L [UCLA].

BOLIVIA. Chuquisaca: Muyupampa, 3 June 1944, Carr (3147), 1 F [USNM]. Locality un-

known: Carr (2911), 1 F [USNM]; Carr (2921), 2 M [UCLA].

13. Aedes (O.) raymondi Del Ponte, Castro and Garcia

Figs. 3,32

1951. Aedes (Ochlerotatus) raymondi Del Ponte, Castro and Garcia, 1951:235-236. TYPE:

Holotype female (587), San Pedro, Jujuy, Argentina, 27 Apr 1926, Shannon and Shan-

non [INM].

Aedes (Ochlerotatus) raymondi of Stone, Knight and Starcke (1959: 152); Forattini (1965:368).

FEMALE (fig. 32). Wing: 3.25 mm. Proboscis: 2.05 mm. Forefemur: 1.70

mm. Abdomen: 2.90 mm. Thorax: Broad anterior scutal stripe divided by narrow

acrostichal line of tan scales. Legs: Knee spots of white scales present on apices of

femora. Claws of hindleg with submedian tooth. Abdomen: Terga II-VII with

basomedian white scale patches.

MALE, PUPA, LARVA. Unknown.

SYSTEMATICS. Aedes raymondi, very closely related to euplocamus and pater-

soni, shares the same habitat with the latter species, having been described from

specimens in the type series of patersoni. Known only in the female, raymondi is

separated from patersoni and the other species of the Scapularis Group by a single

character, a narrow acrostichal line of tan scales dividing the characteristic broad

anterior scutal stripe of silvery white scales of the Infirmatus Subgroup. In the ab-

sence of contrary evidence from other stages and the presence of this unusual scutal

marking, I am recognizing raymondi as a distinct species.

BIONOMICS. Aedes raymondi was described from 2 females from the type

series of patersoni. The series was reared from larvae collected from a heavily vege-

tated temporary rain pool with associated larvae of Psorophora (J.) ferox and Culex

(Lutzia) sp.

DISTRIBUTION (fig. 3). Aedes raymondi is known only from the type locality,

Jujuy, Argentina. Material examined: 2 specimens, 2 females.

ARGENTINA. Jujuy: San Pedro de Jujuy, 27 April 1926, R. and E. Shannon, 1 F (holotype)

[INM]; same data, 1 F [INM].

Arnell: Scapularis Group 53

14. Aedes (O.) infirmatus Dyar and Knab

Figs. 3,29,30,31

1906. Aedes infirmatus Dyar and Knab, 1906a:190,197. TYPE: Syntypes larvae, Baton

Rouge, Louisiana, USA, J. W. Dupree [NE; see Stone and Knight, 1956:219].

Aedes (Ochlerotatus) infirmatus of Dyar (1928:166); Edwards (1932:142); Matheson (1944:158-

159); Coher (1948:102); Carpenter and LaCasse (1955:199-200); Stone, Knight and Starcke

(1959:147); Craig and Horsfall (1960:17); Forattini (1965:323-324); Reinert (1970: 148).

Aedes (Heteronycha) infirmatus of Dyar (1920:105; 1922b:50-51).

Aedes infirmatus of Dyar and Knab (1906b:162); Howard, Dyar and Knab (1917:781-783);

Dyar (1918a:77; 1924a:119); Shlaifer and Harding (1946:250); Weathersbee and Arnold

(1947:216); Breland (1951:362-371); Breeland, Snow and Pickard (1961:288).

Aedes (Ochlerotatus) scapularis infirmatus of Dyar (1922a:59); Root (1924:452).

FEMALE (figs. 29,30). Wing: 3.75 mm. Proboscis: 2:40 mm. Forefemur:

1.90 mm. Abdomen: 3.30 mm. Head: Palpus 3 segmented. Thorax: Broad anter-

ior scutal stripe entirely silvery white. Legs: Knee spots of white scales absent pos-

teriorly on apices of femora. Claws of hindleg with submedian tooth. Abdomen:

Terga II-VII entirely dark scaled dorsally.

MALE (figs. 29,30). Wing: 3.60 mm. Proboscis: 2.55 mm. Forefemur: 1.70

mm. Abdomen: 3.55 mm. Abdomen: Terga without basal bands of white scales

although basolateral white scale patches large and prominent on distal segments.

MALE GENITALIA (fig. 30). Segment IX: Anterior middorsal emargination of

tergum relatively broad and deep, caudal bridge connecting tergal lobes narrow; ter-

gal lobes usually broader than high with 5-7(2-7) setae. Sidepiece: Length about

3.5 times median width, tergal surface with short setae mesad, longer setae laterad

and about 3 subapicotergal setae; sternomesal margin with numerous, about 70-80,

long setae; apical tergomesal lobe well developed; basal tergomesal lobe well devel-

oped, triangular with anterior and posterior sides subequal, with 20-25 undifferent-

iated setae of 2 sizes, long dorsally and ventrally and short in center of cluster, usu-

ally with 3 or 4 long setae projecting distad from posterior edge of lobe. Claspette:

Stem uniformly narrow and broadly curved dorsad; filament subequal in length to

stem, narrow at base, abruptly expanded near middle with large retrorse process

and several elongate spicules basad of retrorse process, and tapered to acute, re-

curved tip. Clasper: Spiniform long, length 12 times its greatest width. Phallo-

some: Aedeagus pyriform.

PUPA (fig. 30). Abdomen: 3.35 mm. Trumpet: 0.55 mm. Paddle: 0.95 mm.

Cephalothorax: Moderately pigmented; leg cases, wing case and dorsum darker.

Seta 8-C with 6-8(4-9) branches. Trumpet: Strongly pigmented, very dark brown.

Abdomen: Moderately pigmented, dark brown at base of segment III. Genital lobe

considerably darker than segment VIII. Seta 1-IV 4-5(4-7) oranched, 1-V 2-4(2-6)

branched; seta 1-VII triple (2-5b). Seta 2-VI usually mesad of seta 1-VI. Seta 3-V

triple (244b). Seta 5-[V,V usually subequal to or slightly shorter than length of suc-

ceeding tergum; seta 5-VII 4-5(2-6) branched. Seta 6-III usually triple (3-4b); seta

6-IV 4(2-4) branched; seta 6-V triple (1-3b); seta 6-VI double (2-3b).

3 LARVA (fig. 31). Head: 0.90 mm. Siphon: 0.90 mm. Anal Saddle: 0.45 mm.

Thorax: Spicules of integument sparse and inconspicuous, short, length about 2-3

times basal diameter. Seta 3-P single. Seta 7-P triple (2-3b). Abdomen: Seta 6-I, II

double, very rarely triple; seta 6-III-VI single. Seta 13-III strongly developed, single,

subequal to seta 13-IV,V which are also single. Segment VIII: Comb scales moder-

54 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

ate in size, with well-differentiated median spinule, at least 2% to 3 times length of

-submedian spinules, 20-25 in number, in irregular double row. Siphon: Index

about 2.6-3.0. Pecten with teeth evenly spaced, extending to middle of siphon.

Seta 1-S inserted distad of pecten. Anal Segment: Ventral brush (4-X) with 8 pairs

of setal tufts, the most proximal tuft occasionally arising from within or on margin

of saddle.

SYSTEMATICS. Aedes infirmatus can be separated from the other species of the

Scapularis Group by the following characters: in the female (1) the scutum has a

broad anterior stripe of silvery white scales, (2) there are no knee spots of white

scales on the apices of the femora, and (3) the abdominal terga are entirely dark

scaled dorsally; in the male genitalia the sternomesal margin of the sidepiece has at

least 65 and as many as 85 well developed setae between the basal tergomesal lobe

and the clasper; in the larva (1) the spicules of the thoracic and abdominal integu-

ment are short and sparse, length about 2 to 3 times basal diameter, (2) seta 13-III

is well developed, single and subequal to 13-IV,V, (3) the siphon index is greater

than 2.5, and (4) the median spinule of the comb scales is long, about 3 times the

length of the submedian spinules.

Primarily on the basis of distinctive male genitalic characters, infirmatus appears

to be the most advanced species of the Infirmatus Subgroup. It is also the only

Nearctic species in the subgroup, probably invading continental North America from

the West Indies.

In most areas where species of the Infirmatus Subgroup are sympatric with other

species of the Scapularis Group the former seem to be the less common, an excep-

tion being the southeastern United States where infirmatus and trivittatus of the

Trivittatus Subgroup occur together. This may indicate that infirmatus became es-

tablished in this area prior to the probable post-Pleistocene invasion of North Amer-

ica proper by trivittatus and has been the more successful competitor.

BIONOMICS. Aedes infirmatus is found throughout most of the southeastern

United States and is abundant in localized areas. It breeds in temporary woodland

pools formed by rain or stream overflow and along the grassy edges of permanent

marshes and ponds. Most breeding sites contain an abundance of vegetation. There

seems to be no preference for either shade or sun. This species overwinters in the

egg stage, the larvae hatching in the late spring and adults appearing 2-3 weeks later.

Larvae are produced whenever conditioned eggs are covered by rains or floodwaters

so a variable number of broods are produced each year. Adults of infirmatus will

attack whenever their habitat is invaded but are typically more active at dusk.

The egg of infirmatus is described and figured by Craig and Horsfall (1960:17)

and all larval instars are described by Breland (1951).

The role of infirmatus in the transmission of arboviruses is discussed above under

Medical Importance.

DISTRIBUTION (fig. 3). Aedes infirmatus is found throughout the southeastern

United States north to Delaware and Missouri and west to Texas. The record of

infirmatus from Arizona by Rigby (1968:239) refers to trivittatus. Material examin-

ed: 371 specimens; 60 males, 159 females, 113 pupae, 39 larvae; 37 individual

rearings (4 larval, 23 pupal, 10 incomplete).

UNITED STATES. Arkansas: Scott, 31 Aug 1908, J. Thibault, 8 F [USNM]. Delaware:

Thompsonville, 21 Aug 1961, R. Lake, 1 L [USNM]. Florida: Archbold Bio. Sta., Highlands

Co., 13-19 Apr 1970, W. Wirth, 1 F [USNM]. Archer, 31 July 1969, J. Reinert, 1 M, 1 F [JFR];

same locality and collector, 9 Aug 1969, 1 M [JFR]. Astor Park, Lake Co., 10 Nov 1933, 2M

[USNM]. Avon Park, Aug 1943, R. Arnett, 7 F [UCLA]; same locality, 21 May 1944, 1 L

Arnell: Scapularis Group 55

[USNM] ; same locality, 2 L [UCLA]. Chantilly Acres, Alachua Co., 25 Apr 1970, W. Wirth,

2 F [USNM]. Everglades N.P., Long Pine Key, 8 July 1971, T. Rogers, S. Chew (FLA 414A),

2 F [UCLA]. Gainesville, 8 Apr-6 Aug 1969, J. Reinert, 24 F [JFR]; same locality and collect-

or, 17 Jan 1970 (69.84), 5 pM (3,12,20,71,73), 8 pF (2,5-7,16,18,22,42), 1 pM, 6 lp, 3 M,

5 F, 21 p, 3 P, 51, 1 L [JFR]; same locality and collector, 28 Jan 1970 (70.2), 1 pF, 1 M [JFR];

same locality and collector, 21 Feb 1970 (70.13), 3 p, 2 L [JFR]. Miami, 10 Dec 1921, G.

Moznette, 1 F [UCLA]. Micanopy, Alachua Co., 20 Feb 1970, J. Reinert (70.11), 4 L [JFR];

same locality and collector (70.19), 4 pM (2,3,5,26), 4 pF (1,22,23,25), 19M, 15 F, 12 P, 7 L

[JFR]; same locality and collector, 9 Apr 1970 (70.37), 6 M, 5 F, 13 p, 1 P [JFR]; same data

(70.38), 1 lp (15), 8 M, 10 F, 16 p [JFR]. Seminole Co., 4 Oct 1937, 2 Ip, 2 1 [USNM].

St. Joseph’s St. Pk., Gulf Co., 1-3 May 1970, W. Wirth, 4 F [USNM]. Tampa, Dyar and Caudell,

1 F [USNM]. Yeehaw Jct., 53 km W_ Vero Beach, 17 July 1972, T. Rogers (FLA 62), 1 F

[UCLA]. Georgia: Brooklet, Bullock Co., 20 Sept 1933, 1 F [UCLA]. Hinesville, 29 Sept

1933, 12 F [USNM]. Homerville, 1 Sept 1933, 3 F [USNM]. Savannah, 28 Apr 1951, G. Heid,

1 M, 1 F [UCLA]. St. George, Charlton Co., 30 Aug 1933, 1 M [USNM]. Summerville, 15 May

1934, 2 F [USNM]. Louisiana: Alexandria, 31 Mar 1943, W. Wirth, 3 L [USNM]. Baton

Rouge, 7 June 1941, W. Wirth, 1 F [USNM]; same locality and collector, 10 May 1942, 2 F

[USNM]. Harihan, 30 May 1943, J. Belkin (SE 479), 1 F [UCLA]. New Orleans, 15 Sept 1914,

W. King, 2 IpM (218.8,231.1), 2 IpF (218.11,218.12), 1 lp (226.1) [USNM]; same locality,

3 Apr 1927, R. Turner, 2 M, 4 F [USNM, BM]. Norco, 15 Oct 1944, F. Young (ASM 250-7),

15 F [UCLA]. Rapides Co., 1 Apr 1943, W. Wirth, 2 L [USNM]; same locality and collector,

7 Apr 1943, 2 M, 2 F [USNM]; same locality and collector, 4 July 1943, 1 M [USNM]. Mississip-

pi: Gulfport, 30 Apr 1929, T. Griffiths, 1 F [USNM]. Missouri: Jefferson City, 21 Oct 1926, F.

Bingham, 1 F [USNM]. North Carolina: [?] Camp Davis, 18 June 1942, 1 L [USNM]. Char-

lotte, 1920, H. Barret, 1 F [USNM]. Wilmington, 28 Aug 1931, Bishopp, 1 F [USNM]. South

Carolina: Jasper Co., 20 Apr 1951, 2 F [UCLA]. Kingstree, 26 July 1934, 1 F [USNM]. [?]

Stark Gen. Hospital, 23 Aug 1943, Middlekauff, 1 L [USNM]. Texas: El Paso, 26 May 1922,

R. Tarbett, 3 F [USNM]. Virginia: Assateague [Is.], 9, 11 July, 16 Oct 1962, K. Ludlam, 3 F

[USNM].

Additional Records from the Literature

UNITED STATES. Illinois: Massac County (Hayes, 1965:165). Texas: Brownsville (Dyar,

1922b:52; McGregor and Eads, 1943:938). Palmetto St. Pk., nr. Luling (Breland (1951:369).

SCAPULARIS SUBGROUP

FEMALES. Head: Occipital dorsolateral dark scale patch reduced or absent.

Erect scales usually unicolorous, white. Proboscis usually distinctly paler ventrally.

Palpus 4 segmented. Thorax: Scutum with large rounded hexagonal patch of light

scales from anterior promontory to scutal angle and posteriorly to near level of pos-

terior end of paratergite, often with posterior extensions along dorsocentral line to

scutellum and surrounding prescutellar bare space; scales silvery white to yellowish

tan. Acrostichal, anterior dorsocentral, and posterior fossal setae absent. Legs:

Foreleg and midleg with conspicuous white streak posteriorly on femora and basal

tarsal segments. Hindleg with conspicuous white stripe on anterior surface of tibia,

occasionally nearly encircling tibia, and continued on basal 1 or 2 tarsal segments.

_ Claws with acute submedian tooth. Wing: Small patch of white scales present at

extreme base of costa and often remigium. Abdomen: Terga II-VII with large baso-

lateral white patches and basomedian patches of pale scales, more obvious poster-

iorly and often forming an indistinct longitudinal line.

56 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

MALES. Thorax: Anterior scutal scale patch of light scales similar to female

though often somewhat reduced in extent laterally. Legs: White stripe on anterior

surface of hindtibia and basal tarsal segments often inconspicuous. Posterior claw

of foreleg moderately enlarged, with acute subbasal tooth. Abdomen: Terga dark

scaled dorsally. | |

DISCUSSION. The Scapularis Subgroup is characterized in the female by a large

usually hexagonal patch of yellowish tan to white scales on the anterior of the scu-

tum, a stripe of white scales on the anterior surface of the hindtibia and usually an

indistinct longitudinal stripe of light scales on the abdomen. In the male genitalia,

the claspette filament lacks or has a very small retrorse process on the convex mar-

gin and the apical tergomesal lobe is moderately to well developed.

The Scapularis Subgroup consists of 3 species, the abundant and widely distrib-

uted scapularis, phaeonotus from the island of Grenada in the Lesser Antilles and

comitatus from the vicinity of Villavicencio, Colombia. Aedes phaeonotus is known

in all stages and differs from scapularis primarily in the color of the anterior scutal

scales of the adult. Known from only 2 males, comitatus has distinctive male gen-

italia as pointed out in the species diagnoses.

15. Aedes (O.) phaeonotus Arnell, sp. n.

Figs. 4,33,34

TYPES. Holotype female with associated larval and pupal skins (GR 104-102), Chantimelle

Village, near Sauteurs, St. Patrick Parish, Grenada, elev. 200 m, larva taken from temporary stream

pool, 30 Oct 1963, R. Martinez [USNM]. Allotype male with genitalia slide (730813-2)

(GRR 82), Grenada, locality unspecified, 1929 [USNM]. Paratypes: 1 F (GRR 21), Belair

School, St. Andrew Parish, Grenada, 8 July 1929; Grenada, locality unspecified, 1 M, 3 F(GRR

25), 8 July 1929; 9 M, 10 F (GRR 30), 10 July 1929; 1 F (GRR 54), 16 July 1929;1 M(GRR

75a), 23 July 1929; 4 M, 16 F (GRR 82), 1929; 7 M, 1 F (GRR 139), 21 Aug 1929, F. Root

[UCLA, BM, FH].

FEMALE (fig. 33). Wing: 3.00 mm. Proboscis: 2.20 mm. Forefemur: 1.55

mm. Abdomen: 2.65 mm. Thorax: Hexagonal patch of light scales on anterior of

scutum yellow to yellowish tan.

MALE (fig. 33). Wing: 3.10 mm. Proboscis: 2.40 mm. Forefemur: 1.55 mm.

Abdomen: 3.15 mm. Generally similar to female except for sexual differences.

MALE GENITALIA (fig. 33). Segment IX: Anterior middorsal emargination of

tergum broad and relatively deep, caudal bridge connecting tergal lobes narrower

than distance between lobes; tergal lobes about as broad as high, with 4 or 5 setae.

Sidepiece: Length about 4 times median width; tergal surface with setae relatively

long, and 3 or 4 subapicotergal setae; sternomesal margin with 30-40 setae; apical

tergomesal lobe well developed; basal tergomesal lobe well developed, broadly coni-

cal, with about 15 undifferentiated setae, most on ventral surface of lobe. Clas-

pette: Stem uniformly narrow and broadly curved dorsad; filament slightly longer

than stem, narrow at base, expanded near middle, and tapered to acute tip. Clasper:

Spiniform length about 10 times its greatest width. Phallosome: Aedeagus ovate

to slightly pyriform.

PUPA (fig. 33). Abdomen: 2.75 mm. Trumpet: 0.40 mm. Paddle: 0.65 mm.

Cephalothorax: Weakly pigmented. Seta 10-C 9-10 branched. Trumpet: Weakly

pigmented, light yellowish brown in color. Abdomen: Weakly pigmented; base of

segments HI and IV darker. Seta 3-V single; seta 3-VII double. Seta 6-III-VI single.

LARVA (fig. 34). Head: 0.80 mm. Siphon: 0.70 mm. Anal Saddle: 0.30 mm.

Arnell: Scapularis Group 57

Thorax: Spicules of integument numerous and conspicuous, length about 6-10

times basal diameter. Seta 3-P single or double. Seta 7-P triple. Abdomen: Seta

1-III 6,8 branched; seta 1-IV,V strongly developed, single. Seta 6-I,II double; seta

6-III-VI single. Seta 8-II single. Seta 13-III strongly developed, single, subequal to

seta 13-IV,V which are single. Segment VIII: Comb'scales moderate in size, evenly

fringed, about 22 in number, in irregular double to triple row. Siphon: Index about

2.25. Pecten with teeth evenly spaced, extending to middle of siphon. Seta 1-S in-

serted distad of pecten. Anal Segment: Ventral brush (4-X) with 9 pairs of setal

tufts.

SYSTEMATICS. Aedes phaeonotus can be separated from the other species of

the Scapularis Group by the following characters: in the female (1) the anterior sur-

face of the hindtibia and basal hindtarsal segments has a conspicuous stripe of white

scales and (2) the large light scale patch on the anterior scutum is yellow to yellow-

ish tan; in the male genitalia, indistinguishable from scapularis, (1) the claspette fil-

ament is subequal in length to the claspette stem and without or with a very minute

retrorse process, (2) the apical tergomesal lobe of the sidepiece is moderately devel-

oped, and (3) there is no enlarged undifferentiated seta ventrally on the basal tergo-

mesal lobe of the sidepiece; in the larva (1) seta 3-P is at least triple, (2) seta 5-C is

single, (3) the siphon has no ventroapical prolongation, and (4) seta 1-III is multiple

(6-8 branched).

Although common and very widely distributed, scapularis is apparently absent

from Puerto Rico, the Virgin Islands and the Lesser Antilles. The Scapularis Sub-

group is, however, represented on Grenada, one of the southernmost of the Lesser

Antilles, by phaeonotus, distinguishable from scapularis in the female by the yellow

to yellowish tan anterior scutal scale patch, and, tenuously, in the larva by the more

branched seta I-III. I have seen only one larva of phaeonotus and this character

may not hold when more specimens are examined.

Aedes scapularis is common in Trinidad, Tobago and adjacent northern South

America. The presence of distinct but closely related species on Trinidad and Gre-

nada is found in other aedines, for example busckii and ioliota of Aedes (Howard-

ina) and splendens and celeste of Haemagogus (H.).

BIONOMICS. The holotype larva of phaeonotus was taken in seasonal deciduous

forest from a temporary stream pool covered with surface scum. Other species as-

sociated with phaeonotus were Anopheles (N.) argyritarsis, Culex (Mel.) jocasta and

Psorophora (G.) sp near cingulata. The allotype and paratype adult collections, lo-

cality unknown, were made during a 1929 malarial survey of Grenada (Root and

Andrews, 1938).

DISTRIBUTION (fig. 4). Aedes phaeonotus is known only from the island of

Grenada in the Lesser Antilles. Material examined: 58 specimens; 23 males, 33 fe-

males, 1 pupa, 1 larva; 1 individual rearing (larval).

GRENADA. Type series, see above.

16. Aedes (O.) scapularis (Rondani)

Figs. 4,35,36,37

1848. Culex scapularis Rondani, 1848:109. TYPE: Female(s), Brazil, locality unspecified;

TYPE LOCALITY here corrected to vicinity of Belem (Para) [possibly NAPLES]. See

discussion below under systematics.

1891. Ochlerotatus confirmatus Lynch Arribalzaga, 1891:146. TYPE: Lectotype female, Na-

58 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

varro, Buenos Aires, Argentina, May 1887, F. Lynch Arribalzaga [BA; selection of O.

H. Casal in Belkin, Schick and Heinemann, 1968:14]. Synonymy with scapularis by

Blanchard (1905:335).

1906. Aedes hemisurus Dyar and Knab, 1906a:199. TYPE: Larva, Kingston, Jamaica; based

on figure in Grabham (1905:405) [NE; see Stone and Knight, 1956a:218]. Synonymy

with scapularis by Howard, Dyar and Knab (1917:787). Resurrected from synonymy

with scapularis by Belkin, Heinemann and Page (1970:161). NEW SYNONYMY.

1907. Aedes indolescens Dyar and Knab, 1907:11-12. TYPE: Holotype female, Cayamas,

Cuba, May, E. A. Schwarz [USNM, 10249; see Stone and Knight, 1956:219]. Synon-

my with hemisurus and scapularis apparently by Howard, Dyar and Knab (1917:787).

1918. Aedes (Ochlerotatus) camposanus Dyar, 1918b:128-129. TYPE: Lectotype male, Guay-

aquil, Guayas, Ecuador, date not specified, F. Campos Ribadeneira [USNM, 21916;

selection of Stone and Knight 1956:216]. Synonymy with euplocamus apparently by

Dyar (1925a:144). Resurrected from synonymy with euplocamus by Levi-Castillo

(1951:383). NEW SYNONYMY.

1933. Aedes (Ochlerotatus) rhyacophilus Lima, 1933:404-407. TYPE: Syntypes males, te-

males with larval exuviae and pupae, Valle do Chanaan, Esperito Santo, Brazil, 1932,

J. Serafim (TYPE LOCALITY restricted to Sao Joao de Petropolis by Belkin, Schick

and Heinemann, 1971:19) [LU]. NEWSYNONYMY.

Aedes (Ochlerotatus) scapularis of Bonne-Wepster and Bonne (1923:126); Dyar (1923:181-182;

1928:167-168, in part); Shannon and Del Ponte (1928:73,121-122); Shannon (1931a:7;

1931b:149); Kumm (1931:56-57); Edwards (1932:142); Soper, Penna et al. (1933:576); Lane

(1939:112-114; 1953:665-667, in part); Cerqueira (1943:32; 1961:145); Matheson (1944:

159-160); Coher (1948: 101-102); Levi-Castillo (1952:262); Carpenter and La Casse (1955:224-

226); Duret and Barreto (1956:89); Perez Vigueras (1956:286-295); Vargas (1956:23); Rachou,

Lima et al. (1958:417); Stone, Knight and Starcke (1959:153); Fauran (1961:26); Garcia and

Ronderos (1963:29); Prosen, Carcavallo and Martinez (1964:101-102); Forattini (1965:331-

341, in part); Montchadsky and Garcia Avila (1966:29,75); Porter (1967:37-40); Xavier and

Mattos (1970:445-446); Belkin, Heinemann and Page (1970:164).

Aedes (Heteronycha) scapularis of Dyar (1920:105).

Aedes scapularis of Howard, Dyar and Knab (1917:783-787); Dyar (1918a:77; 1921:150); Evans

(1921:445-446); Martini (1931a:118; 1931b:204); Prado (1931:201); Komp (1936:62); Del

Ponte, Castro and Garcia (1951:236); Komp (1956:38); Torre y Callejas, Alayo Delmau and

Calderon Chapman (1961:68-69); Mattos and Xavier (1965:278); Garcia Avila and Gutsevich

(1969:6); Morales-Ayala (1971:142).

Culex scapularis of Hunter (1900:281); Blanchard (1905:335); Lutz, de Sousa Araujo and da

Fonseca Filho (1919:86).

Leucomyia scapularis of Theobald (1910:315); Surcouf and Gonzalez-Rincones (1911:160); Lutz

and Nunez Tovar (1928:37).

Aedes (Ochlerotatus) scapularis scapularis of Dyar (1922a:57-59); Bonne and Bonne-Wepster

(1925:388-390).

Aedes scapularis scapularis of Dyar (1924a:118).

Ochlerotatus confirmatus of Hunter (1900:280); Coquillett (1906b:19,21).

Culex confirmatus of Theobald (1901:42-44; 1903a:191; 1905:26); Giles (1902:443-444); Lutz

(1904a:7; 1904b:4); Theobald and Grabham (1905:25); Aiken (1907:68).

Aedes scapularis hemisurus of Dyar (1924a:118); Root (1927:465).

Aedes (Ochlerotatus) hemisurus of Belkin, Heinemann and Page (1970:161-165).

Aedes hemisurus in part of Dyar and Knab (1906b: 162).

Aedes (Ochlerotatus) camposanus of Levi-Castillo (1951:383; 1952:262); Stone, Knight and

Starcke (1959:140); Forattini (1965:337); Belkin, Heinemann and Page (1970:164).

Aedes (Ochlerotatus) rhyacophilus of Soper, Penna et al. (1933:575);; Lane (1953:671-673);

Stone, Knight and Starcke (1959:152); Forattini (1965:368); Garcia and Casal (1968:108).

Aedes (Ochlerotatus) euplocamus of Dyar (1925a:144, in part; 1928:167, in part); Edwards

Arnell: Scapularis Group 59

(1932:142, in part); Martini (1935:55, in part); Lane (1939:109, in part); Pritchard, Seabrook

and Mulrennan (1947:11).

Aedes euplocamus of Dyar and Knab (1906b:162, in part); Del Ponte, Castro and Garcia (1951:

236).

Aedes (Ochlerotatus) condolescens in part of Martini (1935:54-55).

Aedes condolescens of Martini (193 1b:204); Morales-Ayala (1971:141).

Aedes angustivittatus of Komp (1936:62; 1956:38).

FEMALE (figs. 35,36). Wing: 3.10 mm. Proboscis: 2.10 mm. Forefemur:

1.50 mm. Abdomen: 2.55 mm. Thorax: Hexagonal patch of light scales on anter-

ior of scutum silvery white, often yellow to tan on lateral border.

MALE (figs. 35,36). Wing: 2.70 mm. Proboscis: 2.20 mm. Forefemur: 1.40

mm. Abdomen: 2.80 mm. Generally similar to female except for sexual differ-

ences.

MALE GENITALIA (fig. 36). Segment IX: Anterior middorsal emargination of

tergum broad and usually deep, though occasionally relatively shallow; caudal bridge

connecting tergal lobes usually narrower than distance between lobes; tergal lobes

about as broad as high, with 3-6(2-8) setae. Sidepiece: Length about 3.5-4.0 times

median width; tergal surface with short to relatively long, uniformly distributed

setae, and 2 or 3 subapicotergal setae; sternomesal margin with 30-40 setae; apical

tergomesal lobe moderately developed; basal tergomesal lobe moderately developed,

triangular with anterior and posterior sides subequal, often folded ventrad in slide

preparations, with about 12-20 undifferentiated setae, those in center of cluster

shorter, and 3-4 longer setae projecting distad from posterior edge of lobe. Clas-

pette: Stem uniformly narrow and broadly curved dorsad; filament subequal in

length to stem, narrow at base, expanded near middle into broadly obtuse outer

angle, often with small retrorse process, and tapered to acute, recurved tip. Clasper:

Spiniform length about 10 times its greatest width. Phallosome: Aedeagus ovate

to pyriform.

PUPA (fig. 36). Abdomen: 2.60 mm. Trumpet: 0.45 mm. Paddle: 0.75 mm.

Cephalothorax: Weakly pigmented. Seta 10-C usually 3-4(2-7) branched. Abdo-

men: Weakly pigmented; base of segments HI and IV slightly darker. Seta 3-VII

usually single (1-2b). Seta 6-III-VI single, very rarely double.

LARVA (fig. 37). Head: 0.80 mm. Siphon: 0.80 mm. Anal Saddle: 0.35 mm.

Thorax: Spicules of integument numerous and conspicuous, length about 6-10

times basal diameter. Seta 3-P usually double (2-5b). Seta 7-P triple (2-4b). Ab-

domen: Seta I-III usually triple (3-5b); seta 1-IV,V strongly developed, single.

Seta 6-I,II double (1-3b); seta 6-III-VI single (rarely double). Seta 8-II single (rarely

double). Seta 13-III strongly developed, single, subequal to 13-IV,V which are

single. Segment VIII: Comb scales moderate in size, evenly fringed, 20-24 (12-32)

in number, in irregular double to triple row. Siphon: Index about 2.0-2.6. Pecten

with teeth evenly spaced, extending slightly beyond middle of siphon. Seta 1-S in-

serted slightly distad of pecten. Anal Segment: Ventral brush (4-X) with 8 pairs

of setal tufts.

SYSTEMATICS. Aedes scapularis can be separated from the other species of the

Scapularis Group by the following characters: in the female (1) the anterior surface

of the hindtibia and basal hindtarsal segments has a conspicuous stripe of white

scales and (2) the large light scale patch on the anterior scutum is white; in the male

genitalia, indistinguishable from phaeonotus, (1) the claspette filament is subequal

in length to the claspette stem and without or with a very minute retrorse process,

60 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

(2) the apical tergomesal lobe of the sidepiece is moderately developed, and (3)

there is no enlarged undifferentiated seta ventrally on the basal tergomesal lobe of

the sidepiece; in the larva (1) seta 3-P is at least double, (2) seta 5-C is single, (3) the

siphon has no ventroapical prolongation, and (4) seta 1-III is usually triple.

I consider scapularis to be a single species extending from the southwestern

United States through Central and South America to Argentina with an intrusion

into the Greater Antilles. JI-am reducing hemisurus from Jamaica and the Greater

Antilles, camposanus from western South America and rhyacophilus from south-

eastern Brazil to the synonymy of scapularis. These species have been considered

to be distinct from scapularis on the basis of variation in the scutal ornamentation

of the female and details of the male genitalia.

The light scale patch on the anterior of the scutum of scapularis varies from being

straight from the anterior promontory to the scutal angle with few dark brown

scales between the light scale patch and the anterolateral scutal margin to being

concave to a greater or lesser degree with a proportionately larger number of dark

brown scales in the concavity. I find that these differences are a matter of individ-

ual variation with the entire range of variation appearing in most populations, al-

though the frequency may be clinal, the populations from Central America and the

West Indies having fewer individuals with a reduction in white scutal scaling and

those from South America having more. The extent of white scaling on the hind-

tibia shows slight individual variation as does the presence or absence of a small

retrorse process on the outer angle of the claspette filament of the male genitalia.

These variations appear to be strictly characteristic of individuals rather than pop-

ulations.

Aedes scapularis is the most common and widely distributed species of the Scapu-

laris Group and one of the most widely distributed mosquitoes in the New World.

It is probably one of the more advanced species in the Scapularis Group, as evi-

denced not only by its distribution and abundance, but also, in comparison with

other members of the group, modern morphological characters such as light scales

on the hindtibia and reduction of the retrorse process on the claspette filament of

the male genitalia. |

Belkin, Schick and Heinemann (1971:20,41) restricted the type locality of scapu-

laris to the vicinity of Rio de Janeiro, Brazil; however, Papavero (1971-1973:341-

343) evidences that the entomological collections from which Camillo Rondani de-

scribed scapularis were made during a 3 month period in the summer of 1846 in the

immediate vicinity of Belem by Vittore Ghiliani. I am therefore correcting the type

locality to the vicinity of Belem, Para, Brazil. Belkin (1968:7,53,56) states that the

location of the type series of scapularis is unknown, though possibly in the Instituto

di Zoologia, Bologna. Nelson Papavero (1975, in litt.) indicates that part of the

Rondani collection, possibly including the types of scapularis, may be either in the

Instituto di Zoologia, Naples or in the possession of the Rondani family.

BIONOMICS. Aedes scapularis is found at low to moderate elevations through-

out most of tropical and subtropical America. It breeds in a wide variety of tempo-

rary or semipermanent freshwater situations, primarily temporary rain-filled or

stream overflow pools but including pond and swamp margins, rockholes and crab-

holes, in either sun or partial shade. Associated with larvae of scapularis are a wide

variety of species of Anopheles, Uranotaenia, Culex, Psorophora and Aedes. In tem-

porary, grassy, rain-filled pools near Rio de Janeiro, Brazil, the most common mos-

quito associates were Psorophora (J.) sp, P. (P.) ciliata and Culex (C.) sp. In Ja-

maica, scapularis is most often found with fortilis.

Arnell: Scapularis Group 61

Females of scapularis attack man readily, and though primarily crepuscular, will

bite anytime they are disturbed. Forattini (1961) presents evidence that in areas of

prolonged association with man, scapularis is adapting to human habitations and is

becoming more concentrated around dwellings in rural and semirural areas, even en-

tering buildings to bite. Causey and Kumm (1948) recaptured marked females of

scapularis 4.1 km from the point of release in a forested area of Brazil, indicating a

relatively large dispersal range for this species.

Aedes scapularis is by far the most important species of the Scapularis Group in

the transmission of disease organisms, and this aspect of its biology is discussed

above under Medical Importance.

DISTRIBUTION (fig. 4). Aedes scapularis extends from the Rio Grande Valley

of Texas through the Atlantic lowlands of Mexico to Honduras, the Pacific lowlands

of Guatemala to northern Costa Rica; the Bahamas and Greater Antilles except for

Puerto Rico; and throughout South America at low and moderate elevations with

the exception of Chile and southern Argentina. Material examined: 3601 speci-

mens; 531 males; 1635 females, 668 pupae, 767 larvae; 305 individual rearings (183

larval, 106 pupal, 16 incomplete).

ARGENTINA. Buenos Aires: Otamendi (nr. Campana), 22 Mar 1969, O. Casal (GA 697), 4

pM (101-103,105), 1 pF (106) [INM]. Chaco: Quitilipi, May 1952, 1 M,3 F [USNM]. Resist-

encia, 21-26 Feb 1927, R. Shannon, 8 F [USNM]. Resistencia, H. Parker, 6 F [USNM]|. Formo-

sa: [Puesto] El Coati, May 1950, 1 M, 1 F [USNM]. Jujuy: Calilegua, 5 May 1926, R. Shannon,

3 F [USNM]. Ledesma, 19 Mar 1926, N. Davis and R. Shannon, 1 F [USNM]. San Pedro de

Jujuy, 28 Apr 1926, R. Shannon, 48 L [USNM|. Mendoza: Mendoza, 5 Jan 1927, F. and M.

Edwards, 1 F [BM]. Misiones: Iguazu, 5 Oct 1927, R. and E. Shannon, 1 M, 2 F [USNM].

Posadas, 13-14 Jan 1927, F. and M. Edwards, 1 F [BM]. Salta: Balbuena (nr. Tartagal), 4 June

1969, Garcia, Casal and Fernandez (GA 764), 3 pM (101,102,106), 1 pF (103) [INM]. Campo

del Cuervo (nr. Tartagal), 4 June 1969, Garcia, Casal and Fernandez (GA 763), 1 pM (101), 2 pF

(104,105) [INM]. Embarcacion, 4 May 1926, R. Shannon, 4 F [USNM]. Gen. Enrique Mosconi

(Campamento Vespucio), 29 May 1969, Fernandez, Garcia and Casal (ARG 742), 5 IpM (12,18,

19,27,28), 5 IpF (13,17,22,24,29), 6 pM (10,21,102,104,107,113), 6 pF (15,20,101,109,110,

112), 1 lp (23), 2 F, 1 P, 6 1, 50 L [UCLA]; same data (GA 743), 2 lpM (14,15), 1 IpF (12)

[INM]; same data (GA 744), 7 pM (104,105,108,110,111,113,114), 5 pF (103,106,107,109,

112), 1 M [INM]; same locality, 5 June 1969, Garcia (GA 765), 2 pF (100,101) [INM]; same

locality, Mosconi Airport, 29 May 1969, Garcia and Fernandez (GA 746), 1 pF (113) [INM].

Santa Victoria, May 1960, 1 F [USNM]. Tablillas (nr. Vespucio), 5 June 1969, Casal, Garcia

and Fernandez (GA 759), 2 IpF (11,14), 2 pM (12,101) [INM]. Taragal R., Tartagal, 31 May

1969, Garcia and Casal (GA 747), 2 IpF (13,14), 5 pM (102-104,110,112), 4 pF (100,106,109,

114) [INM]; same data (GA 753), 1 M [INM]. Tucuman: Monteros, 12 Mar 1926, R. Shannon,

1 M [USNM]. Vipos, 4 Feb 1927, R. Shannon and E. Del Ponte, 5 F [USNM]. Vipos, 3 Apr

1927, E. Del Ponte, 1 L [USNM]. ,

BAHAMA ISLANDS. Andros Island, 26 June 1903, T. Coffin, 1 F [USNM].

BELIZE. Belize: Belize, 0m, 11 Aug 1967, D. Bertram (BH 443), 1 F [UCLA]. Belize (mile

25 west), 30 m, 10 July 1967, D. Bertram (BHA 269), 1 F [UCLA]. Freetown, 6 m, 16 Mar

1967, D. Bertram (BHA 57), 4 F [UCLA]. Ladyville (nr. Stanley Airport), 0 m, 31 July 1967, D.

Bertram (BH 362), 2 F [UCLA]. Maskalls, 180m, 30 July 1970, K. and D. Schroeder (BH 506),

1 F [UCLA]. Cayo: Central Farm, 70 m, 30 July 1967, D. Bertram (BH 360), 1 F [UCLA];

same locality and collector, 17 Aug 1967 (BH 479), 1 F [UCLA]. Roaring Creek, 70 m, 28 July

1967, D. Bertram (BH 350), 1 F [UCLA]; same locality and collector, 8 Aug 1967 (BH 396), 1 F

[UCLA] ; same locality and collector, 9 Aug 1967 (BH 435), 1 F [UCLA]; same data (BH 436),

1 F [UCLA]. Stann Creek (mi 49 on Hummingbird Hwy), 180 m, 9 Aug 1967, D. Bertram

(BH 426), 1 F [UCLA].

BOLIVIA. Beni: Baures, W. Komp, 2 F [USNM]. L. Rogagua, Nov 1921, W. Mann, 2 F

[USNM]. Rio Matos, 25 Sept 1943, Carr (3059), 12 F [USNM]. San Borja, 6 Oct 1943, Carr

62 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

(3061), 4 F [USNM]. San Ignacio, 18 Sept 1943, W. Komp (KO 202A-18), 28 F [UCLA]. San

Joaquin, 6 June 1964, P. Woke, 5 F [USNM]. Santa Clara, 27 Oct 1943, 3 F [USNM]. Santa

Cruz: Cercado, Jan 1939, 1 F [USNM]. Rincon de Palometas, Sara, 22 Jan 1944, Carr (3101),

3 F [USNM]. Santa Cruz, 21 Feb 1953, J. Munro, 4 F [USNM] ; same locality, 26 Jan 1944,

3 F [USNM]. Locality unspecified: Carr, 1 F (2765B), 1 F (2773B), 1 F (2778), 1 F (2783 A),

1 F (2783B), 3 F (2784), 2 F (2785A), 1 F (2785B), 2 F (2785D), 3 F (2786A), 1 F (2793 A),

1 F (2797), 1 F (2840), 3 F (2844), 2 F (2845), 1 F (2847), 3 F (2848), 1 F (2849), 1 F (2852),

2 F (2855), 3 F (2863), 1 F (2867), 3 F (2868A), 7 F (2870), 1 F (2871), 1 F (2873), 1 F (2875),

1 F (2881), 5 F (2884), 2 F (2898), 1 F (SFA 2971), 1 F (SFA 2972), 1 F (SFA 2974), 2 F

(SFA 2975), 1 F (SFA 2976), 2 F (SFA 2977), 1 F (SFA 3027A), 2 F (SFA 3027C),: 218

(SFA 3929C), 1 F (SFA 3030A), 1 F (SFA 3038A), 5 F (SFA 3038C), 1 F (SFA 30394),

3 F (SFA 3042B) [UCLA, USNM].

BRAZIL. Bahia: Aratu, P. MacCreary, 1 F [USNM]. Caravelas, May 1944, MacCreary and

Bricker, 1 M [USNM]. Piraja, 11 Apr 1929, R. Shannon, 1 lp, 3 M, 24 F [UCLA, USNM] ; same

locality, Feb-Mar 1929, N. Davis and R. Shannon, 9 M, 2 F [USNM]; same locality, 1930, R.

Shannon, 1 M [USNM]; same locality, 2 Nov, R. Shannon, 1 M [USNM]. Salvador, 1972, J.

Fowler (BRS 164), 3 L [UCLA]; same data (BRS 183), 2 L [UCLA]; same data (BRS 274),

3 L [UCLA]; same data (BRS 281), 1 L [UCLA]. Salvador, Dec 1928, R. Shannon, 1 M

[USNM]. Salvador, H. Kumm, 3 M [BM]. Espiritu Santo: Vale do Canaan, May 1932, 2 F

[USNM]. Mato Grosso: Chapada, 600 m, Ponce and Lane, 1 M, 3 F [UCLA]. Maracaju, July

1938, 6 M, 9 F [UCLA]. Salobra, July 1939, 1 F [UCLA]. Para: Belem, APEG Forest, 24

Sept 1970, T. Aitken et al., 11 M [UCLA]; same locality and collectors, 29-30 Sept 1970, 1 F

[UCLA]; same locality and collectors, 8 Oct 1970, 1 M [UCLA]; same locality and collectors,

22 Oct 1970, 1 M [UCLA]. Belem, 30 m, 29 Sept 1970, T. Aitken (BRA 57), 1 M [UCLA].

Belem, 11 Apr 1941, W. Komp (KO 202A-35), 1 F [UCLA]. [?] Belem, Durham, 2 F [BM].

Rio de Janeiro: Iguassu, Aug 1938, R. Shannon, 1 F [USNM]. Jacarepagua, 0 m, 29 Dec 1975,

J. Belkin, M. Faran, G. Bryce (BRA 237), 2 IpM (10,11), 3 IpF (12-14), 3 pM (100,101,103),

5 pF (102,104-107), 1 IP (15) [UCLA]; same data (BRA 238), 3 lpM (10,11,14), 16 IpF (12,

13,15-19,90-98), 4 pM (102-105), 1 pF (100), 22 M, 13 F, 123 P, 1 p, 123 L [UCLA]; same

data (BRA 239), 4 1pM (11,12,15,16), 4 lpF (10,13,14,17), 1 pM (101), 1 pF (100), 1 L [UCLA].

Niteroi, 0 M, 6 Jan 1976, J. Belkin et al. (BRA 266), 1 lpM (10), 1 IpF (11) [UCLA]. Petropolis,

May 1938, R. Shannon, 1 M [USNM]. Rio de Janeiro, 9 Dec 1899, C. Moreira, 4 F [BM]; same

locality, Sept 1938, 3 M [USNM]. Rio de Janeiro, Ilha do Fundao, 0 M, 28 Dec 1975, J. Belkin,

M. Faran, G. Bryce (BRA 235), 7 IpM (20-26,70), 4 IpF (20,27-29), 2 L [UCLA]; same data

(BRA 236), 6 IpM (20-22,27-29), 11 IpF (11,23,24,26,41-45,49,51), 6 pM (100-105), 2 1P (25,

50), 84 M, 94 F, 1 P, 196 p, 53 L, 1321 [UCLA]; same locality and collectors, 29 Dec 1975

(BRA 240), 14 M, 28 F [UCLA]. Santa Cruz, L. Fajardo, 1 F [BM]. Rio Grande do Norte:

Natal, 5 Aug 1943, MacCreary and Bricker, 1 M, 3 F, 1 L [USNM]. Santa Catarina: Nova Teu-

tonia, 10 July 1938, F. Plaumann, 1 F [BM]. Sao Paulo: Itaquaquecetuba, 23 Jan, C. Town-

Sh ce ee Sao Paulo, A. Lutz, 1 F [USNM]. Locality unspecified (BRAP 411), 1 M, 2

COLOMBIA. Antioquia: Medellin, 1923, L. Dunn, 1 F [USNM]. Turbo, 27 May 1943,1F

[USNM]. Cordoba: Carillo (nr. San Pelayo), 20 m, 15 July 1965, Incora (COB 80), 2 lpM (12,

13), 3 IpF (10,11,14), 1 F, 1 L [UCLA]. Monteria (13 km N), 4 Oct 1969, W. Page (COL 419),

39 F [UCLA]; same data (COL 420), 15 F [UCLA]; same locality and collector, 5-6 Oct 1969

(COL 422), 2 F [UCLA]. San Carlos, 80 m, 16 July 1969, Incora (COB 82), 4 IpM (10,11,13,

14), 2 IpF (12,15) [UCLA]. San Pelayo, 20 m, 15 July 1965, Incora (COB 81), 4 lpM (10,21,31,

34), 5 IpF (11,12,20,30,32), 2 F, 4 L [UCLA]. Magdalena: El Banco, 1 Dec 1943, L. Pinzon

(COT 50), 1 F [UCLA]. Santander: Lebrija, 10 Apr 1966, C. Marinkelle (COM 64A), 1 F

[UCLA] ; same locality and collector, 11 Apr 1966 (COM 64B), 3 F [UCLA]. Valle del Cauca:

Buenaventura, Feb 1940, W. Komp, 3 M, 7 F [USNM]. Locality unspecified: (COR 175), 1 F

[UCLA] ; (COR 484), 1 F [UCLA] ; (COR 485), 1 F [UCLA]; (COR 497), 1 F [UCLA].

COSTA RICA. Guanacaste: Palo Verde OTS Sta., 10 m, 18 Aug 1971, S. and D. Heinemann

(CR 448), 1 F [UCLA] ; same locality, 20 June 1975, J. Hayes (CR 602),6 F [UCLA].

Arnell: Scapularis Group 63

CUBA. Havana: Havana, Dec 1903, J. Taylor, 1 M, 2 F [USNM, BM]. Havana, 1909, P.

Serre, 1 F [USNM]. Marianao, Apr 1936, H. Carr, 1 M, 6 L [USNM]. San Antonio de los Banos,

J. Pazos, 3 F [USNM]. Oriente: Cayamas, E. Schwarz, 5 F [USNM]. Cayamas, 8 May 1906, E.

Schwarz, 1 F (indolescens holotype) [USNM]. Levisa (nr. Nicaro), 9 July 1953, K. Knight (379),

1 F [UCLA].

DOMINICAN REPUBLIC. Dajabon: Dajabon, 38 m, 10 July 1971, T. Rogers (RDO 110), 1 F

[UCLA] ; same locality, 40 m, 11 July 1971, T. Rogers (RDO 111), 1 L [UCLA]; same data

(RDO 114), 1 lpM (18), 2 IpF (16,17), 2 IP (10,11), 1 M [UCLA]. Distrito Nacional: San Felipe,

39 m, 29 June 1971, J. Belkin and T. Rogers (RDO 57), 1 F [UCLA]. Santo Domingo, A. Busck,

6 F [USNM]. Santo Domingo, Calle Caonaba, 27 m, 25 July 1971, T. Rogers and E. Drake

(RDO 157, 1 F [UCLA]. Santo Domingo, La Venta, 50 m, 6 Aug 1971, T. Rogers (RDO 207),

22 F [UCLA]. Santo Domingo, Finca Engombe, 2 m, 22 June 1971, J. Belkin, E. Drake and T.

Rogers (RDO 29), 1 IpM (21), 1 IpF (20), 1 pF (100), 2 P, 10 L [UCLA]; same locality, 2 Aug

1971. E. Drake and T. Rogers (RDO 196), 1 M [UCLA]; same locality, 27 Aug 1971, T. Rogers

(RDO 265), 3 IpM (10,11,13), 1 pM (102), 1 Ip (12), 1 P, 21,1 L, E [UCLA]; same locality and

collector, 28 Aug 1971,(RDO 266), 3 F [UCLA]; same data (RDO 267), 8 IpM (20,22,25,28,31,

33 34,36), 7 lpF (23,24,29,32,35,37,39), 1 lp (30), 1 IP (21), 5 M, 16 F, 13 L [UCLA]; same data

(RDO 268), 1 IpF (48), 2 M,2 F,8P, 2p, 1 L [UCLA] ; same data (RDO 269), 1 IpM (10), 1 IpF

(11) [UCLA]; same locality and collector, 29 Aug 1971 (RDO 271), 1 IpF (21), 1 Ip (20)

[UCLA]; same data (RDO 272), 1 pF (108), 3 M, 5 F [UCLA]; same data (RDO 273), 6 F

[UCLA]; same locality and collector, 30 Aug 1971 (RDO 278), 1 lpM (10), 1 lpF (11), 1 pF

(100), 6 F, 3 L, E [UCLA]. Duarte: San Francisco de Macoris, 185 m, 22 July 1968, D. Watson

(RDO 20), 4 F [UCLA]. El Seibo: Sabana de la Mar, 8 m, 30 June 1971, J. Belkin and T. Rogers

(RDO 59), 3 F [UCLA]. La Vega, 75 m, 25 July 1968, D. Watson (RDO 24), 3 F [UCLA].

Peravia: San Jose de Ocoa, Los Anones, 960 m, 23 July 1968, D. Watson (RDO 22), 1 F [UCLA].

Samana: Santa Barbara de Samana, 2 m, 27 June 1971, J. Belkin and T. Rogers (RDO 49), 1 M,

6 F, 7 p [UCLA]. San Cristobal: San Cristobal, La Toma, 120 m, 23 July 1968, D. Watson

(RDO 23), 2 lpM (20,21), 1 lp (22), 1 L [UCLA]. Valverde: Mao, 400 m, 14 Aug 1971, T.

Rogers and J. Gomes (RDO 223), 5 F [UCLA].

ECUADOR. Guayas: Guayaquil, 5 Feb 1966, J. Belkin et al. (ECU 100), 7 lpM (11-14,20,23,

25), 8 IpF (15-19,21,22,24), 1 pM (102), 1 p, 2 1, 8 L [UCLA]; same data (ECU 101), 20 F

[UCLA]; same data (ECU 102), 1 IpM (20), 3 pF (100,101,103), 24 L [UCLA]. Guayaquil, 9

Feb 1966, J. Belkin and E. Gerberg (ECU 130), 92 F [UCLA]. Guayaquil, 22 May 1974, D.

Pletsch (ECU 187), 3 F [UCLA] ; same data (ECU 188), 2 F [UCLA] ; same data (ECU 188A), 3 F

[UCLA] ; same data (ECU 188B), 1 F [UCLA]. Guayaquil, F. Campos R., 1 F (camposanus lecto-

type), 9 M, 5 F [USNM, BM]. Guayaquil, Mar 1940 (KO 202A-25), 12 F [UCLA]. Guayaquil

(km 12 on rt 3), 11 Feb 1966, W. Hjort and D. Schroeder (ECU 144), 16 F [UCLA]. Guayaquil

(km 10 on rt 3), 12 Feb 1966, E. Gerberg (ECU 162), 1 F [UCLA]; same locality, 15 Feb 1966,

J. Belkin and E. Gerberg (ECU 168), 1 lpM (22), 1 IpF (21), 2 pM (103,106), 2 pF (104,105), 1 Ip

(20), 1 1 [UCLA]; same data (ECU 169), 2 pM (85,113), 1 pF (81), 2M, 1 F,3 p,3 L [UCLA];

same data (ECU 170), 3 F [UCLA]. Pascuales, 5 Feb 1966, W. Hjort and E. Gerberg (ECU 108),

9 F [UCLA]. Samborondon, 10 Feb 1966, J. Belkin (ECU 138), 69 M, 89 F [UCLA]. Los Rios:

Babahoyo, Apr 1938, F. Campos (KO H-11-7), 8 F [UCLA]. Manabi: Manta, 14 Mar 1972 (ECU

184), 2 F [UCLA]. Locality unspecified: 1938, H. Hanson (ECUK 67), 5 F [UCLA].

EL SALVADOR. La Union: Canton El Cipres, 24 June 1966, J. Austin (SAL 45), 6 F

[USNM]. San Miguel: Laguna el Jocotal, 20 km E Usulutan, 21 Sept 1965, J. Austin (SAL 11),

1 F [USLA]; same locality and collector, 5 May 1966 (SAL 41), 4 F [UCLA]; same locality and

collector, 24 May 1966 (SAL 43), 1 F [UCLA]. San Salvador: San Salvador, 14 June 1958, L.

Bottimer, 2 F [USNM]. Locality unspecified: 2 F [USNM].

FRENCH GUIANA. Guyane: Cabassou, 26 Nov 1968, J. Clastrier (FGC 3875), 1 F [UCLA];

same locality, 31 Jan 1965, T. Aitken, R. Martinez and A. Guerra (FG 15), 1 M [UCLA]; same lo-

cality and collectors, 3 Feb 1965 (FG 43), 2 F [UCLA] ; same data (FG 45), 1 F [UCLA]. Mont-

joly (9 km E Cayenne), 8 Feb 1965, T. Aitken and R. Martinez (FG 69), 1 M [UCLA].

GUATEMALA. Escuintla: Puente El Carrizo (nr. Iztapa), 10 July 1964, P. Cowsill (GUA 34),

64 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

28 F [UCLA]. Izabal: Bananera, 26 June 1964, P. Cowsill (GUA 4), 2 F [UCLA]. Bananera, 11

Aug 1964, T. Zavortink (GUA 114), 1 F [UCLA]. Bananera (13 km W), 4 Aug 1964, T. and J.

Zavortink (GUA 95), 3 L [UCLA]. Mojaca Village (nr. Morales), 28 June 1964, J. Zavortink and

-P. Cowsill (GUA 12), 1 F [UCLA]. Navajo (betw. Morales and Puerto Barrios), 27 June 1964, T.

and J. Zavortink and P. Cowsill (GUA 7A), 1 F [UCLA]. Peten: La Liberdad, Apr 1942,

(KO 50-8), 2 F [UCLA]. San Juan Acul, Apr 1942, 2 F [USNM]. :

GUYANA. Berbice: Berbice, Jan 1908, J. Aiken, 2 F [USNM]. Corentyne R., Village 116, 12

Oct 1962, T. Aitken [UCLA]. New Amsterdam, Oct 1966, M. Giglioli (GUY 11), 5 M, 5 F

[UCLA] ; same locality and collector, May, June 1967 (GUY 16), 49 F [UCLA] ; same locality, E.

Rowland, 2 F [BM]. Ptn. Blairmont, Nov 1920, G. Bodkin, 1 F [BM]. Demerara: Georgetown,

2 May 1944, C. Giglieli, 3 M, 3 F [USNM]. Hyde Park, 1 Aug 1941, L. Rozeboom (BGR 5), 1 F

[UCLA]. Locality unknown: Dr. Low, 1 F [BM].

HAITI. Nord: Plaine du Nord, 16 Jan 1930 (HAR 16), 1 F [UCLA]. Nord-Ouest: Port de

Paix, 8 Oct 1970, J. Porter (HAT 117), 1 F [UCLA]. Ouest: Bord de Mer Royale, 3 July 1968,

D. Watson (HAT 10), 1 F [UCLA]. Petit Goave, 18 Oct 1924 (HAR 1), 4M [UCLA]. Port-au-

Prince, Oct 1929, Cook (HAC 2), 1 F [UCLA]. Port-au-Prince (Cite Simone Duvalier), 9-10 Oct

1970, M. Solis (HAT 133), 2 F [UCLA]. Sud: Les Cayes, 600 m, 8 July 1968, D. Watson (HAT

14), 2 F [UCLA]. Locality unspecified: 2 July 1931, Cook (HAC 9), 2 F [UCLA] ; 29 Oct 1929

(HAR 8), 2 M, 1 F [UCLA]; (HAR 43), 1 F [UCLA]; (HAC 22), 6 F [UCLA]; (HAC 23), 2 F

[UCLA].

HONDURAS. Atlantida: Lancetilla, 9 Feb 1972, K. and D. Schroeder (HON 121), 1 F

[UCLA]. Tela, 30 Oct 1940, W. Komp, 1 M, 1 F [USNM]. Tela (2.5 km NW), 1.5 m, 9 Feb

1972, D. Schroeder (HON 118), 1 IpM (10) [UCLA]. Colon: Corocito, 3 Apr 1924, 1 F

[UCLA]. Puerto Castillo, 24 Nov-21 Dec 1942, J. Duncan, 10 M, 9 F [USNM, UCLA] ; same lo-

cality and collector, 24 Dec 1942 (KO 204A-9), 10 F [UCLA]; same locality, K. Maxwell, 1 F

[USNM]. La Lima, 5 Aug 1964, P. Cowsill (HON 1), 1 F [UCLA]. San Pedro Sula (21 km N),

11 Aug 1967, A. Adames (HON 65), 10 F [UCLA].

JAMAICA. Clarendon: Goshen, 10 Sept 1965, J. Belkin (JA 353), 1 F [UCLA]. Parnassus,

10 m, 28 July 1967, W. Page (JA 863), 5 lpM (21-24,26), 4 IpF (20,25,27,28), 1 pM (37), 3 M,

1 P, 11,11 L [UCLA]. Yorks Pen, 50 m, 6 Sept 1966, D. Watson (JA 604), 1 lpF (10) [UCLA].

Kingston and St. Andrew: Kingston, from M. Grabham, 1 F [USNM]. Rio Cubre Dam, M.

Grabham, 1 F [USNM]. St. Andrew: Ferry, 15 m, 9 Aug 1967, J. Belkin, W. Page (JA 896), 4

IpF (13,23,24,28), 3 pM (102,104,111), 5 pF (96,101,103,109), 1 lp (22), 33 M, 51 F, 1p, 10L

[UCLA]; same locality, 17 Sept 1967, W. Page (JA 799), 1 F, 1 P [UCLA]; same locality, 18

Sept 1967, W. Page (JA 796), 1 IpF (14), 2 pM (91,102), 1 pF (112), 9 L [UCLA] ; same locality

(JA 768), 2 L [UCLA]; same locality (JA 890), 1 F [UCLA]. St. Ann: Delight, 200 m, 25 Aug

1967, J. Belkin, W. Page (JA 757), 1 IpF (45), 1 pM (112) [UCLA]. St. Catherine: Bog Walk, 90

m, 4 Aug 1968, R. Hochman (JA 913), 1 F, E [UCLA]. Caymanas, 26 Nov 1965, W. Page (JA

404), 5 F [UCLA]. Central Village, 20 m, 29 Aug 1967, W. Page (JA 759), 1 lpF (20) [UCLA].

Spanish Town, 6 Feb 1964, H. Tucker (JA 37), 1 lpF (14) [UCLA]. St. Thomas: Golden Grove

(nr. Morant Bay), 1 Feb 1965 (JA 197), 1 L [UCLA]. Grants Pen, 150 m, 30 Oct 1966, G. Berlin

and D. ee (JA 649), 4 F [UCLA]. Locality unspecified: F.Cundall and M. Grabham, 1 M,

4 F [BM].

MEXICO. Campeche: Hopelchen (12 km W on hwy 180), 80 m, 4 Aug 1970, K. and D.

Schroeder (MEX 600), 1 pF (100) [UCLA]. Lerma, 6 m, 21 July 1970, K. and D. Schroeder

(MEX 592), 1 F [UCLA]; same locality, 5 Aug 1970, D. Schroeder (MEX 605), 1 F [UCLA];

same data (MEX 607), 9 F [UCLA]. Nuevo Leon: Monterey, 500 m, 11 July 1965, R. Schick

and D. Schroeder.(MEX 188), 1 F [UCLA]. Tabasco: Cardenas, 30 m, 15 July 1970, K. and D.

Schroeder (MEX 564), 5 F [UCLA]. Comalcalco (12 km N), 37 m, 12 July 1970, K. and D.

Schroeder (MEX 544), 1 F [UCLA]. Comalcalco (7.5 km N), 38 m, 12 July 1970, K. and D.

Schroeder (MEX 552), 10 F [UCLA]. Frontera, 18 Feb, C. Townsend, 1 F [USNM]. Tenosique,

17 July 1970, K. and D. Schroeder (MEX 569), 1 F [UCLA]. Tamaulipas: Ciudad Mante, 120

m, 14 July 1965, R. Schick and D. Schroeder (MEX 198), 33 F [UCLA]. Veracruz: Cerro

Guzman ( ? Guzmantla), 9 July 1970, K. and D. Schroeder (MEX 535), 2 F [UCLA]. Cordoba,

Arnell: Scapularis Group 65

F. Knab, 2 F [USNM]. Cordoba, 1000 m, 6 July 1970, D. and K. Schroeder (MEX 527), 1 F

[UCLA]. Cuitlahuac (11 km E), 410 m, 15 July 1965, E. Fisher and D. Verity (MEX 35), 6 F

[UCLA]. Santa Lucrecia (Jesus Carranza), F. Knab, 5 F [USNM]. Veracruz, Crawford, 2 F

[USNM].

NICARAGUA. Locality unspecified: 6 Apr 1943, T. Aitken, 2 F [UCLA].

PARAGUAY. Boqueron: Gen. Eugenio A. Garay Colony (Fortin Coronel Eugenio A. Garay),

June 1964, W. Murdock, 4 F [UTAH]. La Cordillera: San Bernardino, Fiebrig, 2 M, 2 F

[USNM]. Locality unknown: [?] Eng. Mission, Choco, 2 June 1927, L. Rozeboom, | F

[UCLA].

PERU. Amazonas: Bagua, 10-15 Jan 1954, W. Ebeling, 19 F [UCLA]. Ica: Pisco, Aug 1925,

T. Cockerell, 1 M [USNM]. La Liberdad: San Idelfonso, 50 m, 25 Feb 1973, F. Morales-Ayala

(PER 109), 23 F [UCLA]; same data (PER 111), 8 F [UCLA]. Lima: Canete, Feb, Mar 1941, E.

Hambleton, 2 F [USNM]. Lima, spring 1943, E. Hopkins, 2 M [USNM]. Piura: Piura, 22 Dec

1968, M. Nelson (PER 10), 1 pF (10),[UCLA] ; same locality and collector, 3 Jan 1969 (PER 23),

5 IpM (10,11,16-18), 4 lpF (12-15), 2 pM (100,101), 1 IP (19), 4 P, 1 p,3 1, 1 L [UCLA]. Sul-

lana, 3 Jan 1969, M. Nelson (PER 17), 8 F [UCLA]. Talara, Rio Parinas, 12 Dec 1943, 1 F

[USNM]. Talara, Mar 1942, J. Ruddy (KO 105-20), 1 F [UCLA]. Tumbes: Tumbes, 30 Dec

1968, M. Nelson (PER 14), 10 F [UCLA] ; same data (PER 16), 1 F [UCLA].

SURINAM. Suriname: Lelydorp, 17-21 Apr 1964 (SUR 78), 1 F [UCLA]; same locality, 21-

23 Apr 1964 (SUR 79), 4 F [UCLA]; same locality, 26-29 May 1964 (SUR 81), 4 F [UCLA];

same locality, 29 May-2 June 1964 (SUR 113), 3 M, 7 F [UCLA]. Zanderij, 26-30 June 1964

(SUR 82), 2 F [UCLA]; same locality, 1-5 July 1964 (SUR 83), 3 F [UCLA] ; same locality, 6-10

July 1964 (SUR 84), 3 F [UCLA] ; same locality, 14-17 July 1964 (SUR 85), 7 F [UCLA] ; same

locality, 21-24 July 1964 (SUR 87), 2 F [UCLA]; same locality, 21-26 June 1964 (SUR 88), 5 F

[UCLA] ; same locality, 27-30 July 1964 (SUR 90), 2 F [UCLA]; same locality, 31 July-3 Aug

1964 (SUR 91), 2 F [UCLA] ; same locality, 27-30 Aug 1964 (SUR 99), 1 F [UCLA] ; same local-

ity, 11-14 June 1964 (SUR 114), 3 F [UCLA]; same locality, 14-17 June 1964 (SUR 115), 1 F

[UCLA] ; same locality, 17-22 June 1964, D. Geijskes (SUR 116), 8 F [UCLA]. Locality unspeci-

fied: 12 Feb 1908, J. Aiken, 2 p [USNM].

TRINIDAD AND TOBAGO. TOBAGO. St. Andrew: Bacolet, 3 Aug 1963, T. Aitken, 1 F

[UCLA]. Scarborough, 60 m, 15 Nov 1965, T. Aitken, R. Martinez, A. Guerra (TOB 2), 1 lpM

(20) [UCLA]; same locality and collectors, 75 m (TOB 4), 1 IpF (50) [UCLA] ; same data (TOB

6), 1 IP (60) [UCLA]; same data (TOB 7), 1 L [UCLA] ; same data (TOB 11), 1 F [UCLA]; same

locality, 20 Nov 1965, R. Martinez and A. Guerra (TOB 65), 1 F [UCLA]; same data (TOB 66),

1 F [UCLA]; same data (TOB 67), 4 F [UCLA]. St. Patrick: Buccoo, 12 Jan 1966, R. Martinez

and A. Guerra (TOB 158), 1 F [UCLA]. Canaan, 19 Nov 1965, T, Aitken, R. Martinez, A. Guerra

(TOB 57), 1 L [UCLA]; same data (TOB 59), 4 F [UCLA]. Golden Grove Road, 30 Nov 1965,

R. Martinez, A. Guerra (TOB 145), 1 IP (10) [UCLA]. St. Paul: Roxborough, 16 Nov 1965, T.

Aitken, R. Martinez, A. Guerra (TOB 28), 1 F [UCLA]; same locality and collectors, 17 Nov 1965

(TOB 50), 1 M, 4 F [UCLA] ; same locality, 30 Dec 1965, R. Martinez, A. Guerra (TOB 153), 2 F

[UCLA]. Locality unspecified: (TOB 161), 1 F [UCLA]. TRINIDAD. Caroni: Todd’s Road,

17 June 1941, Hopkins (TRR 8), 1 M, 1 F [UCLA]. St. Andrew: Cumuto, 11 June 1941, Hop-

kins (TRR 5), 4 F [UCLA]. Cumuto, 11 July 1941, Hopkins (TRR 21), 3 F [UCLA]. Cumuto,

21 July 1941, Hopkins (TRR 28), 4 F [UCLA]. Cumuto, 5 Aug 1941, Hopkins (TRR 40), 5 F

[UCLA]. Cumuto, 14 Aug 1941, Hopkins (TRR 47B), 1 F [UCLA]. Cumuto, 5 Aug 1941

(TRR 62), 1 F [UCLA]. St. Andrew: Mt. Harris, 75 m, 16 July 1964, F. Powdher (TR 573), 5 F

[UCLA]. Mt. Harris, 23-31 July 1924, C. Withycombe, 5 F [BM]. Nestor Village, 14 June 1964,

A. Guerra (TR 484), 1 F [UCLA]. St. Georges: Monos I., 27 July 1931, E. de Verteuil, 2 M

[BM]. Port of Spain, 2 Sept 1929, W. Hoffman (LAR 54), 2 M [UCLA] ; same data (LAR 55), 9

M, 9 F [UCLA]. St. Patrick: Cedros, 23 Aug-19 Sept 1920, J. Pawan, 4 F [BM]. Cedros, C.

Hewlett, 1 M, 1 F [BM]. Fyzabad, 12 Jan 1924, S. Neave [BM]. Victoria: San Fernando, 1-2

Feb 1912, G. Marshall, 2 F [BM]. Locality unspecified: 3 Mar 1931, M. Beattie, 1 M, 1 F [BM];

A. Busck, 3 F [USNM]; F. Urich, 2 F [USNM]. :

UNITED STATES. Texas: Brownsville, 28 Aug 1916, M. High, 1 F [USNM]. Brownsville,

66 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

Oct 1932, A. Niven, 2 F [USNM]. Brownsville, 5 Jan 1940, 2 F [USNM]. Brownsville, R.

Turner, 1 F [USNM]. Cameron Co., 19 Oct 1943, Pfeifer, 1 F [USNM]. Mission, 4 May 1924, R.

Turner, 13 F [USNM]. Mission, 20 Sept 1944, C. Joyce, 2 L [USNM]. Rio Hondo, 11 June

1942, W. Reeves, B. Brookman and R. Eads, 1 M [USNM]. San Benito, 17 June 1942, W. Reeves,

B. Brookman, R. Eads, 1 M [USNM]. Santa Maria, 30 Apr 1942, B. Brookman and W. Reeves, 1

M, 1 P,3 L [UCLA, USNM].

VENEZUELA. Aragua: Cagua, 400 m, 12 Aug 1969, Valencia and Pulido (VZ 344), 15 F

[UCLA]. Maracay, Dec 1967, Hansell and Vera (VZ 91), 1 IpM (10), 1 L [UCLA]. Maracay, 24

Aug 1926, M. Nunez Tovar, 5 F [USNM]. Maracay, 10 May 1927 (VZR 4), 2 F [UCLA]. Mara-

cay, 25 July 1927 (VZR 18), 10 M, 4 F [UCLA]. Maracay, Aug 1928 (VZR 57), 1 M,1F

[UCLA]. Maracay (VZR 247), 2M [UCLA]. Maracay (VZR 249), 2 F [UCLA]. Ocumare de

la Costa, 100 m, 12 July 1969, Pulido and Valencia (VZ 185), 2 F [UCLA]; same locality, 0 m,

15 Aug 1969, Pulido (VZ 352), 2 lpM (10,11), 3 pM (100-102), 2 P, 1 1, 2 L [UCLA] ; same local-

ity, 1927, M. Nunez Tovar, 1 F [USNM]; same locality, 5 July 1927 (VZR 14), 16 F [UCLA].

San Jacinto Mil. Res. (nr. Maracay), 27 Nov 1928 (VZR 98), 3 M [UCLA]; same locality, 19 Feb

1929 (VZR 150), 1 M, 1 F [UCLA]; same locality, 11 Mar 1929 (VZR 153), 2M, 4 F [UCLA];

same locality, 13 Apr 1929 (VZR 168), 3 M [UCLA]; same locality, 3 July 1929 (VZR 212), 2

M [UCLA]. Turiamo, 14 Sept 1944, W. Komp (KO 202A-24), 7 F [UCLA]. Turmero, 400 m,

12 Aug 1969, Valencia and Pulido (VZ 336), 1 pM (100) [UCLA]; same data (VZ 345), 4 F

[UCLA]. Carabobo: Lake Valencia, 29 Nov 1967, R. Hansell (VZ 62), 1 F [UCLA] ; same local-

ity and date, P. Rausch (VZ 66), 1 F [UCLA]. Puerto Cabello, 30 Nov 1967, P. Rausch, R.

Hansell and Vera (VZ 75), 2 pM (101,103), 1 pF (100) [UCLA]. Distrito Federal: Juan Diaz,

Dec 1927 (VZR 31), 4 F [UCLA] ; same data (VZR 32), 1 F [UCLA]; same data (VZR 33), 2 F

[UCLA]. Miranda: La Trinidad, Mar 1929 (VZR 162), 2M [UCLA]. Locality unspecified: 10

Oct 1926, M. Nunez Tovar, 1 M [USNM].

Additional Records from the Literature

ARGENTINA. La Pampa: Planicie de la Loma Negra (Dyar, 1921:150).

BOLIVIA. Beni: Riberalta (Cerqueira, 1943:32).

BRAZIL. Amapa: Amapa; Macapa. Amazonas: Codajas; Manaus. Goias: Pedro Afonso;

Porto Nacional. Mato Grosso: Chapada do Guimaraes; Cuiaba (Cerqueira, 1961:145).

COLOMBIA. Meta: Restrepo (Komp, 1936:62).

17. Aedes (O.) comitatus Arnell, sp. n.

Figs. 4,32

TYPES. Holotype male with genitalia slide (740523-64), Bosque Ocoa, E of Villavicencio,

Meta, Colombia, 23 May 1944, through W.H.W. Komp (KO 200-17) [USNM]. Paratype: 1M

with genitalia slide (740523-63), same locality as holotype, 16 July 1943, through W.H.W. Komp

(KO 200-2) [UCLA].

FEMALE. Unknown.

MALE. Wing: 2.80 mm. Proboscis: 2.05 mm. Forefemur: 1.55 mm. Abdo-

men: about 2.45 mm. Generally similar to scapularis. Thoracic and abdominal

scaling indistinct, specimens badly rubbed but with scattered white scales on an-

terior of scutum. Head: Proboscis short, about 1.35 of forefemur. Palpus about

1.10 of proboscis; segments 2 and 3 making up 0.66 of palpus, segments 4 and 5

relatively short, their combined length about 0.28 of palpus. Antenna about 0.90

of proboscis. Legs: Forefemur 1.20 of distance from top of thorax to apex of

midcoxa. Anterior claw of foreleg and midleg considerably enlarged, with blunt

submedian tooth and apparently acute basal tooth; posterior claw of midleg mod-

Arnell: Scapularis Group 67

erately enlarged, apparently with acute basal tooth. Wing: Scales apparently all

dark. Abdomen: Terga apparently entirely dark scaled dorsally and with basal

white scale patches laterally. Sterna mostly white scaled.

MALE GENITALIA (fig. 32). Segment IX: Anterior middorsal emargination of

tergum broad and shallow, caudal bridge connecting tergal lobes broader than dis-

tance between lobes; tergal lobes broader than high, with 4 or 5 setae. Sidepiece:

Length about 3.5-4.0 times median width; tergal surface with setae relatively short

and sparse, and cluster of 4-6 subapicotergal setae; sternomesal margin with 30-40

well developed setae concentrated at distal third of sidepiece; apical tergomesal lobe

moderately developed; basal tergomesal lobe moderately developed, apparently con-

ical, differentiated seta large, with about 10 undifferentiated setae and a single seta

on ventral margin of setal cluster which is considerably larger than remainder of un-

differentiated setae. Claspette: Stem uniformly narrow and broadly curved dorsad;

filament slightly longer than stem, narrow at base, expanded near middle and taper-

ed to very acute tip. Clasper: Slightly curved inward distally; spiniform length

about 10 times the greatest width. Phallosome: Aedeagus ovate.

PUPA, LARVA. Unknown.

SYSTEMATICS. Aedes comitatus is known only in the male, and can be separ-

ated from all other species in the Scapularis Group by the following characters of

the male genitalia: (1) the claspette filament has no retrorse process and (2) the bas-

al tergomesal lobe of the sidepiece has a ventral seta which is smaller than the differ-

entiated seta but considerably larger than the undifferentiated seta.

Although both specimens are rubbed, there are sufficient numbers of thoracic

scales present to indicate the Scapularis Subgroup scutal scale pattern and slight in-

dications of hindtibial white scales, although the hindtibial white scale streak is rare-

ly prominent in males of the Scapularis Subgroup.

W. H. W. Komp (1936:62; 1956:38) mentions taking, in yellow fever surveys in

the vicinity of Villavicencio, Colombia, adult specimens of scapularis, as well as lar-

vae of angustivittatus that produced adults that “chad the thorax suffused over with

white...the...species thus much resembl(ing) scapularis.”” Scapularis Group speci-

mens that I have been able to locate that may be attributable to Komp’s collections

in this part of Colombia include possibly 4 females of euplocamus collected in 1935,

2 males of angustivittatus, with typical angustivittatus scutal ornamentation, collect-

ed in 1935 and the 2 males of comitatus, collected in 1943 and 1944. It is impossi-

ble to know what species Komp had before him in his reference to “‘scapularis-like

angustivittatus,’’ euplocamus, scapularis, which, in view of its abundance in Colom-

bia probably inhabits the Villavicencio area although I have seen no specimens from

there, or possibly comitatus, which may agree with his cryptic description in the

female.

BIONOMICS. Nothing is known of the bionomics of comitatus. The holotype

was a “hand catch.”

DISTRIBUTION (fig. 4). Aedes comitatus is known from a single locality near

Villavicencio, Meta, Colombia. Material examined: 2 specimens; 2 males.

COLOMBIA. Meta: Type series, see above.

CRINIFER SUBGROUP

FEMALES. Head: Occipital dorsolateral dark scale patch present or absent. Pal-

pus 3 or 4 segmented. Thorax: Scutum with 3 narrow lines of light scales extend-

68 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

ing entire length of scutum, often joined into single, broad line; or, light scales on

dark background in somewhat random pattern but suggesting 3 narrow longitudinal

lines, 1 acrostichal and 1 dorsocentral; white scales along humeral margin, above

paratergite, along supraalar margin, and surrounding prescutellar bare space. Acro-

stichal, anterior dorsocentral, and posterior fossal setae well developed. Legs: Knee

spots of white scales present posteriorly on apices of femora. Tibiae and basal tarsal

segments of foreleg and midleg with light streak posteriorly; hindtibia and hindtar-

sus entirely dark. Claws of hindleg with submedian tooth. Wing: Scales entirely

dark. Abdomen: Terga entirely dark scaled dorsally.

MALES. Thorax: Scutal scale pattern generally similar to female although light

scales more extensive. Legs: Anterior claw of foreleg with submedian tooth and

acute basal tooth. Abdomen: Sterna usually mostly dark scaled.

DISCUSSION. The Crinifer Subgroup is defined by the female scutal scale pat-

tern of narrow acrostichal and dorsocentral lines of light scales extending the length

of the scutum or light and dark scales in a more or less random pattern that suggests

acrostichal and dorsocentral light lines, and by the male genitalia (see species ac-

counts below).

The subgroup consists of 2 species, crinifer and synchytus, with a distribution

throughout southeastern Brazil, Uruguay, northern Argentina and probably eastern

Paraguay (see fig. 1). Aedes crinifer appears to be relatively common in this area;

however, synchytus is known only from the type locality.

18. Aedes (O.) crinifer (Theobald)

Figs. 2,38,39,40

1903. Culex crinifer Theobald, 1903a:209-210. TYPE: Lectotype female, Sao Paulo, Sao

Paulo, Brazil, A. Lutz. [BM; selection of Belkin, 1968:4].

1910. Culex lynchii Brethes, 1910:470-471. TYPE: Lectotype female, Buenos Aires, Argen-

tina, 19 Jan 1903, J. Brethes [BA, selection of O. H. Casal in Belkin, Schick and Hein-

emann, 1968:13]. Synonymy with crinifer by Edwards (1932:142).

1917. Culex tapinops Brethes, 1917:227-229. TYPE: Lectotype male, San Isidro, Buenos

Aires, Argentina, 2 Feb 1917, J. Brethes [BA, selection of O. H. Casal in Belkin, Schick

and Heinemann, 1968:13-14]. Synonymy with lynchii by Dyar (1922a:56).

1928. Aedes (Ochlerotatus) iguazu Shannon and Del Ponte; in Dyar, 1928:168-169. TYPES:

Syntypes, male, female, Misiones, Argentina [LU]. Synonymy with crinifer by Edwards

(1932:142).

Aedes (Ochlerotatus) crinifer of Dyar (1922a:57; 1928:164-165); Bonne and Bonne- Wepster

(1925:391); Shannon and Del Ponte (1928:123); Shannon (1931b:149); Edwards (1932:142)

in part); Antunes and Lane (1934:35-37); Lane (1939:108, in part; 1951:335; 1953:668-669,

in part); Cerqueira (1943:31; 1961:143-144); Duret and Barreto (1956:90); Rachou, Lima et

al. (1958:417); Stone, Knight and Starcke (1959:143, in part); Garcia and Ronderos (1963:

29); Prosen, Carcavallo and Martinez (1964:99); Forattini (1965:358, in part); Belkin, Heine-

mann and Page (1970:164); Xavier and Mattos (1970:445).

Aedes (Heteronycha) crinifer of Dyar (1920: 105).

Aedes crinifer of Prado (1931:201); Davis (1944:225); Del Ponte, Castro and Garcia (1951:236);

Mattos and Xavier (1965:277).

Culex crinifer of Lutz (1904a:8; 1904b:4); Theobald (1905:29; 1907:pl.13,14; 1910:361);

Surcouf and Gonzalez-Rincones (1911:181); Lutz, de Sousa Araujo and da Fonseca Filho

(1919:87).

Aedes (Ochlerotatus) lynchii of Dyar (1922a:56-57; 1928:165); Shannon and Del Ponte (1928:

122).

Arnell: Scapularis Group 69

Aedes (Ochlerotatus) iguazu of Lane (1939:110).

Aedes iguazu of Del Ponte, Castro and Garcia (1951:236).

Aedes (Heteronycha) dolosa of Lynch (1891:156; in part); Dyar (1920:105).

Culicelsa confirmatus of Dyar (1905c:186).

FEMALE (figs. 38,39). Wing: 4.00 mm. Proboscis: 2.40 mm. Forefemur:

1.90 mm. Abdomen: 2.80 mm. Head: Occipital dorsolateral dark scale patch us-

ually present. Palpus 4 segmented. Thorax: Scutum with tan to white scales in

narrow acrostichal line and narrow dorsocentral line extending entire length of scu-

tum, these lines separated by brown scales which are often reduced in extent, re-

sulting in a broad median stripe of light scales; brown scales laterad of dorsocen-

tral line.

MALE (figs. 38,39). Wing: 4.25 mm. Proboscis: 3.00 mm. Forefemur: 1.85

mm. Abdomen: 4.30 mm. Thorax: Scutum with broad median stripe of yellow

to tan scales extending entire length of scutum between dorsocentral lines.

MALE GENITALIA (fig. 39). Segment IX: Anterior middorsal emargination of

tergum broad and shallow, caudal bridge connecting tergal lobes broad; tergal lobes

usually broader than high, with 3-8 setae. Sidepiece: Length about 4.0-4.5 times

median width; sternomesal margin with 50-60 long setae, most dense at level of

base of apical tergomesal lobe; apical tergomesal lobe very well developed; basal

tergomesal lobe well developed, triangular with anterior and posterior sides subequal

in length, setae arising from near posterior side, with about 20-30 undifferentiated

setae, a row of usually 6-8 projecting distad from posterior side of lobe. Claspette:

Stem long, length about 10 times median width, uniformly narrow, broadly curved

dorsad on basal two-thirds, distal third recurved ventrad; filament uniformly narrow

on basal half, angled dorsad near basal fourth, abruptly expanded near middle with

strong retrorse process and usually several elongate spicules basad of retrorse process

and tapered abruptly to recurved tip. Clasper: Narrow, only slightly broadened be-

fore middle; spiniform length about 10-13 times its greatest width. Phallosome:

Aedeagus ovate.

PUPA (fig. 39). Abdomen: 3.00 mm. Trumpet: 0.45 mm. Paddle: 0.80 mm.

Cephalothorax: Weakly pigmented, very slightly darker dorsally. Abdomen: Weak-

ly pigmented, only slightly darker anteriorly. Seta 1-II usually 4-6(4-8) branched.

Seta 5-IV-VI subequal in length to length of succeeding tergum, double on all seg-

ments.

LARVA (fig. 40). Head: 0.75 mm. Siphon: 0.80 mm. Anal Saddle: 0.35 mm.

Head: Seta 5-C triple (3-5b). Seta 6-C triple. Seta 7-C 8-10 (8-11) branched.

Thorax: Spicules of integument numerous and long, length about 5-8 times basal

diameter. Seta 3-P triple (3-5b). Seta 1-M 4(3-5) branched. Abdomen: Seta 13-III

strongly developed, single; seta 13-IV,V usually double. Segment VIII: Comb

scales moderate in size, with well developed median spinule, about 16-20(9-25) in

number, in irregular double to triple row. Siphon: Index about 2.45-2.60. Pecten

with teeth evenly spaced, extending slightly beyond middle of siphon. Seta 1-S in-

serted distad of pecten. /2 or 3 pairs of accessory dorsolateral setal tufts (2a-S)

present, each usually 4(3-5) branched, and subequal to 1-S. Anal Segment: Ven-

tral brush (4-X) with 8 pairs of setal tufts.

SYSTEMATICS. Aedes crinifer can be separated from the other species of the

Scapularis Group by the following characters: in the female the scutum has tan to

white scales in a narrow acrostichal and dorsocentral line extending the entire length

of the scutum, these lines usually separated by dark brown scales, but occasionally

70 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

the light lines merge to form a broad median light stripe bordered laterally by dark

brown scales; in the male genitalia, indistinguishable from synchytus, (1) the clas-

pette stem is narrow and curved dorsad on the basal two-thirds and recurved ventrad

on the distal third, (2) the claspette filament is angled dorsad near the basal fourth

and is very broad just distad of the retrorse process, and (3) the apical tergomesal

lobe is very well developed; in the larva the siphon has 2 or 3 pairs of accessory

dorsolateral setae (2a-S) subequal in development to seta 1-S.

The presence of accessory dorsolateral setae on the siphon of crinifer is unique

in the Scapularis Group and rare except in the Culicini and Sabethini. In New

World Aedini these setae are known in only 2 other species, well developed in Aedes

(O.) trichurus (Dyar), which is widespread in southern Canada and the northern

United States, and poorly developed in Aedes (O.) bicristatus Thurman and Winkler,

a species known only from northern California. Although the accessory dorsolateral

setal tufts of crinifer are subequal in development to the siphon tuft (1-S), I consid-

er them to be a multiplication on seta 2-S rather than of 1-S because of their loca-

tion. / :

Brethes (1910:470-471) described lynchii from females previously described by

Lynch Arribalzaga (1891:156) as Heteronycha dolosa, Lynch apparently describing

his species from male Culex and female Aedes. Brethes (1916:212) later described

the supposed male of lynchii, in reality a Culex (brethesi Dyar, 1919), and then did

not recognize the true male of lynchii and described it as tapinops (1917:227-229).

Dyar (1922a:56) included tapinops in the synonymy of lynchii, and Edwards

(1932:142) synonymized lynchii under crinifer.

Apparently from a large series of adults collected at Iguazu Falls, Argentina by R.

C. and E. M. Shannon during the week of 4-10 October 1927, Shannon and Del

Ponte described iguazu (Dyar, 1928:168-169), and stated that types were deposited

in the U.S. National Museum. Belkin, Schick and Heinemann (1968:14) reported

that 3 female syntypes of iguazu, without type labels but with the above collection

data, were present in the USNM collection. I was unable to locate those specimens

or any other material that could be identified as part of the syntype series of iguazu.

However, 2 males and 15 females of crinifer, with which the Shannon and DelPonte

description of iguazu agrees in every respect, with the above mentioned Shannon

and Shannon collection data, were in the USNM collection. I am therefore consider-

ing iguazu as a synonym of crinifer.

BIONOMICS. Immature stages of crinifer have been collected in temporary

ground pools in all documented cases except | report by Davis (1944) who collected

crinifer larvae in cut bamboo at Teresopolis, Brazil, at a time when all ground pools

were dry. This species was collected near Itapetininga, Brazil from a small, tempo-

rary, grassy ground pool associated with larvae of Anopheles (N.) evansae, Culex

(Mel.) pilosus, Culex (C.) sp near coronator and Aedes (O.) dupreei. Causey and

Kumm (1948) recaptured marked adults of crinifer 2.3 km from the release site

in a forested area of Brazil.

DISTRIBUTION. Aedes crinifer is found throughout southeastern Brazil, Uru-

guay, northeastern Argentina and undoubtedly Paraguay. Although crinifer has

been reported from Panama, Colombia (Komp, 1936:62), Venezuela, French Gui-

ana (Fauran, 1961:26), Ecuador, Peru, Bolivia (Cerqueira, 1943:31; Prosen, Carca-

vallo et al. (1964:99) and northern Brazil (Cerqueira, 1961:143-144), these records

are undoubtedly misidentifications or misinterpretations of this species. A. crinifer

appears to be restricted to the Parana River system and adjacent coastal drainage of

southeastern Brazil and Uruguay. Material examined: 134 specimens; 26 males, 84

Arnell: Scapularis Group 71

females, 13 pupae, 11 larvae; 12 individual rearings (4 larval, 4 pupal, 4 incomplete).

ARGENTINA. Buenos Aires: Buenos Aires, J. Brethes, 4 F [USNM]. Buenos Aires, 1920, J.

Petrocchi, 7 F, 1 L, E [USNM]. Campana, 24 Mar 1966, Garcia and Casal (240), 2 M [INM];

same locality and collectors, 24 July 1968 (648), 2 pM (104,106), 2 pF (103,107), 1 p [INM].

Rio Santiago, 18 Nov 1915, J. Petrocchi, 1 F [USNM]. Corrientes: Paso de los Libres, 23, 25

July 1970, Bejarano (GA 1100,1106,1107,1112), 6 M, 2 F [INM]; same locality and collector,

28 July 1971 (GA 1132,1133,1135), 2M, 1F [INM]. Entre Rios: Concepcion del Uruguay, 3 F

[USNM]. Misiones: Iguazu, 4-10 Oct 1927, R. and E. Shannon, 2 M, 15 F [USNM]. San Ig-

nacio, 1911, E. Wagner, 3 F [USNM]. Province unknown: Miguel Garcia, 10 Dec 1915, J. Petroc-

chi, 1 F [USNM]. Locality unspecified: J. Petrocchi, 1 M [USNM].

BRAZIL. Mato Grosso: Maracaju, June 1937, 19 F [USNM]. Parana: Cambara, Oct 1936,

1 M [USNM]. Curitiba, Dec 1938, 1 M [UCLA]. Rio de Janeiro: Mangaratiba, Nov 1938, 5 F

[USNM, UCLA]. Teresopolis, May 1938, R. Shannon, 4 F [USNM]. Santa Catarina: Nova Teu-

tonia, 27 July-26 Sept 1938, F. Plaumann, 2 M, 2 F [BM]. Sao Paulo: Guaratuba River, 30

Sept 1972, H. Xavier, 1 F [UCLA]. Itapetininga, 30 Nov 1973, H. Xavier, 1 F [UCLA]. Itap-

etininga (28 km W), 5 Feb 1975, T. Rogers and K. Linthicum (BRA 167), 2 IpM (21,24), 2 IpF

(23,70), 3 IP (20,22,25), 1 P, 1 1, 1 L [UCLA]. Juquia, Nov 1938, J. Lane [USNM]. Sao Paulo,

A. Lutz, 1 F (crinifer lectotype) [BM]. Sao Paulo, 6 Aug 1903, 1 M [USNM]. Sao Paulo, J.

Lane and P. Antunes, 1 M, 2 F [BM, UCLA]. Locality unspecified: A. Lutz, 1 F [BM].

URUGUAY. Rivera: Rivera, 3 Sept 1944, 1 M [USNM]. [?] Rocha, San Carlos, 11 Mar

1945, Parker and Hackett, 2 F [USNM]. Treinta y Tres: Treinta y Tres, 7 Mar 1945, Parker and

Hackett, 1 F [USNM].

LOCALITY UNSPECIFIED. 1M, 1 F [BM].

Additional Records from the Literature

BRAZIL. Goias: Anapolis (Mattos and Xavier, 1965:277). Mato Grosso: Cuiaba (Cerqueira,

1961:144). Dourados (Xavier and Mattos, 1970:445). Minas Gerais: Passos (Causey and Kumm,

1948:469). :

19. Aedes (O.) synchytus Arnell, sp. n.

Figs. 4,41

TYPES. Holotype female, Cataratas del Iguazu, Misiones, Argentina, 4-10 Oct 1927, R. C.

and E. M. Shannon [USNM]. Allotype male with genitalia slide (2354), same data as holotype

[USNM]. Paratypes: 41 females, same data as holotype [UCLA, USNM, BM, ISNM, FH].

FEMALE (fig. 41). Wing: 4.00 mm. Proboscis: 2.60 mm. Forefemur: 2.05

mm. Abdomen: 2.40 mm. Head: Occipital dorsolateral dark scale patch reduced

or absent. Palpus 3 segmented. Thorax: Scutum with white to yellow scales on

background of dark brown scales in somewhat random pattern but usually suggest-

ing 3 narrow longitudinal lines, 1 acrostichal and 1 dorsocentral; white scales pres-

ent along humeral, antealar, and supraalar margins and surrounding prescutellar bare

space; light scales often concentrated along prescutal suture.

MALE (fig. 41). Wing: 3.40 mm. Proboscis: 2.45 mm. Thorax: Scutum with

brown and white scales mixed in apparent random pattern.

MALE GENITALIA (fig. 41). Apparently indistinguishable from crinifer.

PUPA, LARVA. Unknown.

SYSTEMATICS. Aedes synchytus is known only in the adult and can be separat-

ed from the other species of the Scapularis Group by the following characters: in

the female the scutal scales are of 2 contrasting colors, white to yellow and dark

brown, and arranged in a somewhat random pattern that often suggests 3 light long

itudinal lines, acrostichal and dorsocentral, on a dark background; in the male geni-

72 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

talia, indistinguishable from crinifer, (1) the claspette stem is narrow and curved

dorsad on the basal two-thirds and recurved ventrad on the distal third, (2) the

claspette filament is angled dorsad near the basal fourth and is very broad just

distad of the retrorse process, and (3) the apical tergomesal lobe is very well de-

veloped.

The type series of synchytus was found in the undetermined excess collection at

the U.S. National Museum, and is part of a large series of adults collected by Shan-

non and Shannon at Iguazu Falls, Argentina in October 1927. It is closely allied to

crinifer, as demonstrated by the male genitalia and the general facies of the adults;

however, the scutal scale pattern is unlike any other species of the Scapularis Group.

BIONOMICS. Nothing is known of the bionomics of synchytus. The type series

consists entirely of adults collected during a 1 week period. Present in the same col-

lection were specimens of crinifer and scapularis.

DISTRIBUTION (fig. 2). Aedes synchytus is known only from Iguazu Falls,

Misiones, Argentina. Material examined: 43 specimens; 1 male, 42 females.

ARGENTINA. Misiones: Cataratas del Iguazu, type series, see above.

OBTURBATOR SUBGROUP

20. Aedes (O.) obturbator Dyar and Knab

Figs. 2,41,42,43

1907. Aedes obturbator Dyar and Knab, 1907:9. TYPE: Holotype female, Tarpon Bay (Tar-

pum Bay), Eleuthera Island, Bahama Islands, 7 July 1903, T. H. Coffin [USNM, 10141].

Aedes (Ochlerotatus) obturbator of Dyar (1922a:52; 1928:163); Edwards (1932:142); Lane

(1939:112; 1953:653); Maldonado-Capriles, Pippin and Kuns (1958:67-68); Stone, Knight

and Starcke (1959:150); Forattini (1965:365); Porter (1967:37,39,40); Belkin, Heinemann

and Page (1970:151-152).

Aedes (Heteronycha) obturbator of Dyar (1920:105).

Aedes obturbator of Johnson (1908:70); Howard, Dyar and Knab (1917:778-779); Dyar (191 8a:

77; 1924a:117).

FEMALE (figs. 41,42). Wing: 3.25 mm. Proboscis: 2.30 mm. Forefemur:

2.05 mm. Abdomen: 2.75 mm. Pleural integument tan to brown, much lighter

in color than in remainder of Scapularis Group. Head: Occipital dorsolateral dark

scale patch much reduced. Pedicel uniformly tan or yellow in color. Palpus 3 seg-

mented. Thorax: Scutum with pale tan scales except for dark brown scales in very

narrow acrostichal line and narrow line or patch near scutal angle and in supraalar

area, and white to yellow scales surrounding prescutellar bare space. Acrostichal,

anterior dorsocentral and posterior fossal setae well developed. Legs: Relatively

long, forefemur about 1.20 of distance from top of thorax to apex of midcoxa.

Forecoxa without dark scales. Light scaling on femora, tibiae, and tarsae much re-

duced; femora with white scales only on posterioventral surface and in small poster-

ior knee spot; tibiae and tarsi entirely dark scaled. Claws of foreleg and midleg

with submedian tooth, or posterior claw of foreleg and midleg with tooth reduced

or absent; claws of hindleg with submedian tooth. Wing: Scales entirely dark. Ab-

domen: Terga II-VI with narrow basal white bands.

MALE (figs. 41,42). Wing: 3.00 mm. Probosics: 2.55 mm. Forefemur: 1.75

mm. Abdomen: 2.95 mm. Head: Palpus relatively short, subequal in length to

proboscis; segments 2 and 3 long, making up about 0.80 of palpus, segments 4 and

Arnell: Scapularis Group 73

5 short, making up 0.10 and 0.07 of palpus, respectively; segment 3 straight and

uniformly slender to apex, segments 4 and 5 slender and straight, segments 3 and

5 with few short setae near apex. Thorax: Scutal ornamentation similar to female,

but pale tan scales more extensive, extending laterally nearly to scutal angle and to

scutal margin in supraalar area. Legs: Relatively long, forefemur about 1.20 of dis-

tance from top of thorax to apex of midcoxa. Anterior claw of foreleg and midleg

with blunt submedian tooth and without acute basal tooth; claws of hindleg with

acute median tooth. Abdomen: Terga with broad basal bands of white scales.

MALE GENITALIA (fig. 41). Segment IX: Anterior middorsal emargination of

tergum broadly conical, caudal bridge connecting tergal lobes relatively broad; tergal

lobes about as broad as high, with 3-6 setae. Sidepiece: Length about 3.5-4.0 times

median width; tergal surface with sparse, short setae from basal fourth to near base

of apical lobe, and 3-4 extremely long, thick subapicotergal setae, laterotergal sur-

face with 1 or 2 extremely long setae arising near basal fourth; sternomesal margin

with 40-50 well developed setae; apical tergomesal lobe poorly developed; basal

tergomesal lobe conical, differentiated seta tapered gradually from base to apex,

apex usually hooked, with about 12-20 undifferentiated setae. Claspette: Stem

uniformly narrow, curved dorsad, more abruptly near distal third; filament long,

with basal two-thirds uniformly narrow and straight, then expanded, with small,

sharply retrorse process, and tapered to acute, recurved tip. Clasper: Broadly ex-

panded near middle, sharply curved inward distally; spiniform length about 20

times its greatest width. Phallosome: Aedeagus ovate.

PUPA. Unknown.

LARVA (fig. 43). Head: 0.60 mm. Siphon: 0.75 mm. Anal Saddle: 0.28 mm.

Thorax: Spicules of integument sparse, inconspicuous, length about 3 times basal

diameter. Seta 3-P single. Seta 7-P double. Abdomen: Seta 6-I-VI single. Seta

13-III strongly developed, double, subequal to 13-IV,V, which are double or triple

Segment VIII: Comb scales moderate in size, evenly fringed, about 20 in number.

Siphon: With acute ventroapical prolongation, subequal in length to ventrolateral

valve. Index about 3.0. Pecten with teeth evenly spaced, extending beyond middle

of siphon. Seta 1-S inserted distad of pecten. Anal Segment: Saddle with relatively

large spicules on dorsocaudal margin. Seta 1-X double. Ventral brush (4-X) with 7

pairs of setal tufts.

SYSTEMATICS. Aedes obturbator can be separated from the other species of

the Scapularis Group by the following characters: in the female (1) the scutum is

covered with pale tan scales except for dark brown scales in a very narrow acrosti-

chal line and on the scutal margin and (2) the pleural integument is very light

brown; in the male genitalia (1) the claspette filament is very long, narrow and

nearly straight, with a small retrorse process, (2) the spiniform is long, about 20

times its greatest width, and (3) 1 or 2 very long, thick setae arise from the lateral

surface of the base of the sidepiece; in the larva (1) the siphon has a conspicuous

ventroapical prolongation about equal in length to the ventrolateral valve and (2)

seta 6-I, II is single.

Only females of this species are known from the type locality in the Bahama Is-

lands. I am considering the specimens from Mona Island, Puerto Rico of females,

males and larvae (Maldonado-Capriles, Pippin and Kuns, 1958:67-68) to be conspe-

cific with obturbator, the females from the two localities being indistinguishable.

The relationships between obturbator and the other members of the Scapularis

Group are obscure. The scutal scaling of obturbator resembles that of (trivittatus,

although the lines of pale tan scales corresponding to the dorsocentral lines of white

to yellow scales in trivittatus are much broader in obturbator, and correspondingly

74 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

the dark brown scales of the acrostichal line and especially the lateral scutal margin

are much reduced in extent. The male genitalia and immature stages of obturbator

give no indication of close affinities to other Scapularis Group species.

BIONOMICS. Maldonado-Capriles, Pippin and Kuns (1958:67) report collect-

ing obturbator larvae from semipermanent rockpools and cement catchment basins

and taking adults feeding day and night on Mona Island, Puerto Rico. This species

apparently has not been taken in the Bahama Islands since Coffin collected the type

series in 1903.

DISTRIBUTION (fig. 2). Aedes obturbator is known only from Eleuthera Island

in the Bahama Islands and Mona Island between Puerto Rico and Hispaniola. Mater-

ial examined: 36 specimens; 4 males, 30 females, 2 larvae; 2 individual rearings (2

incomplete).

BAHAMA ISLANDS. Eleuthera Island, Tarpon [Tarpum] Bay, 1903, T. Coffin, 19 F

[USNM, BM]; same locality and collector, 7 July 1903, 1 F (obturbator holotype) [USNM].

PUERTO RICO. Mona Island, Feb-Nov 1955, W. Pippin, 2 1F, 4 M, 8 F [USNM].

PECTINATUS SUBGROUP

21. Aedes (O.) pectinatus Arnell, sp. n.

Figs. 1,44,45

TYPES. Holotype female (COB 1), Rio Cuja, nr. Fusagasuga, Cundinamarca, Colombia, elev.

1400 m, larva from rock hole in stream margin, 29 Oct 1964, E. Osorno et al. [USNM]. Allotype

male (COB 1) with genitalia slide (690210-19), same data as holotype [USNM]. Paratypes: 1 F

(COB 1), 1 M (COB 1) with genitalia slide (741023-1), 4 Ip (COB 1-10,11,12,13), same data as

holotype [UCLA, USNM].

FEMALE (fig. 44). Wing: 4.15 mm. Proboscis: 2.70 mm. Forefemur: 1.95

mm. Head: Occipital dorsolateral dark scale patch large and conspicuous; erect

scales numerous, elongate, extending far forward on occiput, mostly dark. Pro-

boscis long, about 1.60 of forefemur. Thorax: Scutal scales entirely dark brown

except for few white scales above paratergite. Acrostichal and dorsocentral setae

absent anteriorly. Scutellum with dark scales on lobes. Legs: Conspicuous knee

spots of white scales present on apices of femora. Tibiae and tarsae entirely dark

scaled. Claws of hindleg with submedian tooth. Wing: Scales entirely dark. Abdo-

men: Terga entirely dark scaled dorsally.

MALE (fig. 44). Wing: 4.35 mm. Proboscis: 3.05 mm. Forefemur: 2.20 mm.

Abdomen: 4.05 mm. Thorax: Scutum with white scales in broad patch on poster-

ior half between dorsocentral lines, patch with narrow anterior projection along

acrostichal line which does not reach anterior promontory; remainder of scutum

with dark scales. Legs: Anterior claw of foreleg and midleg considerably enlarged,

with blunt submedian and acute basal tooth.

MALE GENITALIA (fig. 44). Segment IX: Anterior middorsal emargination of

tergum deep, rectangular to pandurate in shape; caudal bridge connecting tergal

lobes very narrow; tergal lobes broader than high, with 5-7 setae. Sidepiece: Length

about 4 times median width; sternomesal margin with about 20-30 setae in irregu-

lar double row; apical tergomesal lobe not developed; basal tergomesal lobe conical,

differentiated seta expanded and slightly curved before middle and straight beyond

middle, with about 20 undifferentiated setae. Claspette: Stem long, uniformly

narrow and broadly curved dorsad; filament slightly longer than stem, narrow at

base, expanded at middle into broadly obtuse outer angle or small, sharply retrorse

Arnell: Scapularis Group 75

process, and tapered to recurved tip. Clasper: Broadly curved inward distally;

spiniform length about 10 times its greatest width. Phallosome: Aedeagus ovate.

PUPA (fig. 44). Abdomen: 4.05 mm. Trumpet: 0.70mm. Paddle: 1.00 mm.

Cephalothorax: Weakly pigmented, slightly darker dorsally. Abdomen: Weakly

pigmented, dark at base of segment III. Seta 1-II 22-27 branched. Seta 5-IV,V

long, about 1.5 times length of succeeding tergum, usually double on segment IV

(1-2b), always single on segment V; seta 5-VI subequal in length to length of suc-

ceeding tergum, single. Paddle: Seta 1-P long, single, about 0.33 of paddle length.

LARVA (fig. 45). Head: 1.00 mm. Siphon: 1.05 mm. Anal Saddle: 0.40 mm.

Head: Seta 5-C triple (2-3b). Seta 6-C 2-3 branched. Mental plate with lateral 4 or

5 teeth much reduced, median teeth normal or slightly enlarged. Thorax: Spicules

of integument moderately dense, thickened basally, length about 6 times basal diam-

eter. Seta 3-P 4(3-5) branched. Abdomen: Seta 1-VII very strongly developed,

single, subequal to siphon length. Seta 13-III strongly developed, single. Segment

VIII: Comb scales small in size, spatulate and evenly fringed, 60-70(55-85) in

number, in large triangular patch. Siphon: Index about 2.8-2.9. Pecten teeth 20-

24, distal 4-6 teeth without basal denticles and slightly detached; pecten extending

to distal 0.75-0.80 of siphon. Seta 1-S 4(3-5) branched, inserted within pecten, at

about distal 0.70 of siphon. Anal Segment: Ventral brush (4-X) with 8 pairs of

setal tufts, all arising from grid. Anal gills long, about 6 times dorsal saddle length.

SYSTEMATICS. Aedes pectinatus can be separated from the other species of the

Scapularis Group by the following characters: in the female (1) the scutum is entire-

ly dark brown scaled and (2) knee spots of white scales are present on the apices of

the femora; in the male genitalia (1) the sternomesal margin of the sidepiece has

only 20 to 30 relatively short setae, (2) the apical tergomesal lobe of the sidepiece

is not developed, (3) the differentiated seta of the basal tergomesal lobe is usually

not curved distad of the middle, and (4) the claspette filament is without or has a

small retrorse process; in the larva (1) the pecten teeth extend beyond the insertion

of seta 1-S and (2) the comb scales number 60-70.

One of the most aberrant species of the Scapularis Group, pectinatus exhibits a

number of characters found nowhere else in the group. Although the males of most

species of the group are less distinct in scutal and abdominal scale patterns than the

females, pectinatus is the only species to exhibit true sexual dimorphism, the female

having a completely dark scaled scutum and the male having a broad patch of white

scales on the posterior half of the scutum. The numerous, completely dark erect

occipital scales and the very long proboscis in the female, the absence of the apical

tergomesal lobe of the sidepiece and the reduction in setae on the sternomesal mar-

gin of the sidepiece of the male genitalia, and the large number of comb scales and

pecten teeth, the extreme development of seta 1-VII and the reduction in the lateral

teeth of the mental plate in the larva all point to an early divergence from the main

line of Scapularis Group evolution.

BIONOMICS. The only information on the bionomics of pectinatus is from the

collection records for the type series. The larvae were collected from a rockhole at

the margin of the Rio Cuja in an area of coffee plantations and second growth for-

est. The water in the rockhole was semipermanent and no vegetation was present.

Associated with pectinatus were larvae of Aedes (O.) scutellalbum, A. (O.) fluvia-

tilis, Culex (C.) sp near articularis and Culex (Cu.) sp near coronator.

DISTRIBUTION. Aedes pectinatus is known only from 2 localities near Bogota,

Cundinamarca, Colombia. Material examined: 13 specimens; 3 males, 2 females, 4

pupae, 4 larvae; 4 individual rearings (incomplete).

COLOMBIA. Cundinamarca: El Colegio, 1750 m, 8 Nov 1940, Bates a Osorno, 1 M

[USNM]. Rio Cuja, nr. Fusagasuga, type series, see above.

76 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

~THELCTER SUBGROUP

22. Aedes (O.) thelcter Dyar

Figs. 1,46,47,48

1918. Aedes (Taeniorhynchus ? ) thelcter Dyar, 1918b:129. TYPE: Holotype female, Browns-

ville, Texas, U.S.A., 29 Aug 1916, M. M. High [USNM, 21718].

1947. Aedes (Ochlerotatus) keyensis Buren, 1947:228-229. TYPE: Holotype female, Key

West, Florida, U.S.A., 15 Oct 1946, E. Fernandez [A]. Synonymy with thelcter by

Thurman, Haeger and Mulrennan (1949:171).

Aedes (Ochlerotatus) thelcter of Dyar (1922a:60; 1928:169); Dyar and Ludlow (1922:62-63);

Edwards (1932:143); Martini (1935:51-52); Matheson (1944:160); Thurman, Haeger and Mul-

rennan (1949:171); Carpenter and La Casse (1955:240-241); Vargas (1956:24); Stone, Knight

and Starcke (1959:155).

Aedes (Heteronycha) thelcter of Dyar (1922b:52).

Aedes thelcter of Dyar (1924a:118; 1924b:132).

FEMALE (figs. 46,47). Wing: 3.55 mm. Proboscis: 2.40 mm. Forefemur:

1.70 mm. Abdomen: 3.25 mm. Head: Eyes rather broadly separated, interocular

space about 3 ommatidial diameters. Occipital dorsolateral dark scale patch often

reduced or absent. Palpus 4 segmented. Thorax: Scutal scales entirely yellowish

white except for light brown to golden scales in broad acrostichal line extending to

posterior third of scutum or occasionally to prescutellar bare space; scales in acro-

stichal area occasionally only slightly darker than surrounding scales making acro-

stichal line indistinct. Acrostichal setae confined to anterior promontory, anterior

dorsocentral setae not developed. Scutellum with white scales, broader than scutal

scales, in large patch on midlobe and often small patch on lateral lobes. Lobes with

10 to 15 strong setae. Ppn covered with narrow, curved golden scales except for

patch of broad white scales posterioventrally. Scale patch on ssp usually extensive,

with a few scales extending dorsad toward ppn. Legs: Short, forefemur about 0.85-

0.90 of distance from top of thorax to apex of midcoxa. White scales on femora,

tibiae and basal tarsal segments usually more extensive but in same pattern as in re-

mainder of group. Claws of hindleg with acute submedian tooth. Wing: Scales

entirely dark. Abdomen: Terga II-VI or VII with basomedian white scale patches,

broader on midline, often extending to distal margin of terga forming distinct me-

dian longitudinal white line on entire length of abdomen; basolateral white scale

patches large, often extending to distal margin of tergum. Sternum entirely white

scaled.

MALE (figs. 46,47). Wing: 3.00 mm. Proboscis: 2.15 mm. Forefemur: 1.45

mm. Abdomen: 3.15 mm. Generally similar to female except for sexual differ-

ences. Head: Proboscis long, about 1.55 of forefemur. Palpus about 1.10 of

proboscis; segments 2 and 3 making up about 0.52 of palpus, segments 4 and 5

each making up about 0.21 of palpus; segment 3 slightly elevated and swollen

near apex, segment 5 slightly downturned at base; apex of segment 3, segment 4

and base of segment 5 densely covered with well developed setae ventrally. Antenna

about 0.75 of proboscis. Thorax: Scutal scales entirely yellowish white except

for broader white scales surrounding prescutellar bare space. Legs: Anterior claw

of foreleg moderately enlarged, with blunt submedian tooth. Abdomen: Terga

with narrow basal bands of white scales.

MALE GENITALIA (fig. 47). Segment IX: Anterior middorsal emargination of

Arnell: Scapularis Group a

tergum broad and shallow, caudal bridge connecting tergal lobes very broad; tergal

lobes broad, with 3-8 setae. Sidepiece: Short and broad, length about 2.5 times

median width; tergal surface with few short to moderate setae and 1-3 long, thick

subapicotergal setae; sternomesal margin with about 20 moderately long setae; api-

cal tergomesal lobe moderately developed, with row of 3-4 short setae on mesal

margin; basal tergomesal lobe prominent, triangular, differentiated seta relatively

small, often only slightly longer than largest of about 25-35 undifferentiated setae.

Claspette: Stem short, narrowed and slightly angled dorsad distally; filament nar-

row at base, expanded near basal fourth into obtuse outer angle; often with small

retrorse barb, and gradually tapered distally to recurved tip. Clasper: Sharply

curved inward distally; spiniform relatively short, length about 10 times its greatest

width. Phallosome: Aedeagus ovate.

PUPA (fig. 47). Abdomen: 2.90 mm. Trumpet: 0.45 mm. Paddle: 0.75 mm.

Cephalothorax: Weakly pigmented, considerably darker dorsally. Abdomen:

Weakly pigmented, darker anteriorly and at base of most segments. Seta 5-IV-VI

subequal in length to length of succeeding tergum, double, rarely single on seg-

ments IV and VI. Seta 6-VII short, dendritic, with about 12-18 main branches.

Paddle: Marginal spicules extremely poorly developed.

LARVA (fig. 48). Head: 0.80 mm. Siphon: 0.80 mm. Anal Saddle: 0.40 mm.

Head: Broad, width about 1.6 of length. Thorax: Spicules of integument sparse

and short, length about 2 times basal diameter. Seta 3-P triple (1-3b). Seta 1-M tri-

ple (2-4b). Abdomen: Seta 1-IV 4-6(4-8) branched; seta 1-V 3(3-5) branched; seta

1-VII 3-4(2-5) branched. Seta 3-IV 5(4-8) branched; seta 3-V 4(3-5) branched. Seta

13-HI weakly developed, 8(6-10) branched. Segment VIII: Comb scales moderate

in size, with well differentiated median spinule and 1-3 smaller lateral spinules,

about 16-21 in number, in irregular single to triple row. Siphon: Index about 1.9-

2.05. Pecten with distal 2 or 3 teeth detached, extending to about distal 0.70-0.75

of siphon. Seta 1-S inserted slightly basad of middle of siphon, 9-13 branched.

Anal Segment: Seta 1-X double (1-4b). Seta 2-X 15(14-19) branched. Ventral

brush (4-X) with 9 pairs of setal tufts.

SYSTEMATICS. Aedes thelcter can be separated from the other species of the

Scapularis Group by the following characters: in the female (1) the scutal scales

are entirely yellowish white except for a usual broad acrostichal line of light brown

to golden scales and (2) the abdominal terga have basal bands of white scales which

are usually expanded on the midline and often form a median longitudinal white

line the entire length of the abdomen; in the male genitalia (1) the sidepiece is short

and broad, length about 2.5 times median width and (2) the basal tergomesal lobe of

the sidepiece has 25-35 setae; in the larva (1) the pecten teeth extend beyond the

insertion of seta 1-S and (2) the comb scales number about 16-20.

Aedes thelcter shows no close affinities to any other Scapularis Group species.

The structure of the male genitalia with the compact sidepiece, densely hirsute basal

tergomesal lobe and short claspette, and the relatively short and broad cercus of the

female genitalia set this species apart from the remainder of the group. Probably

less significant but obvious characters are the unusual scutal and abdominal scaling

of the adult and the extensive row of larval pecten teeth.

BIONOMICS. Aedes thelcter is found commonly in irrigation overflow pools and

temporary rain pools in the lower Rio Grande valley. Miller, Doll and Wheeler

(1964:459) report collecting thelcter on the Pecos River near Artesia, New Mexico

following flash floods which inundated large areas of Tamarix marshes that had not

held water for several years. In this collection of thelcter were also larvae of Psor-

78 | Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

ophora (G.) discolor, P. (G.) confinnis, P. (G.) signipennis, P. (J.) cyanescens, Aedes

(Aedm.) vexans, A. (O.) dorsalis and A. (O.) sollicitans. Aedes thelcter was collect-

ed in considerable numbers in light traps in the lower Rio Grande valley during VEE

surveillance activities in 1971 (Sudia and Newhouse, 1975). Thurman, Haeger and

Mulrennan (1949) describe in detail the breeding habitat of thelcter at Long Key

and Rock Harbor in the Florida Keys, in rain-filled, shallow pools in limestone,

with several species of Psorophora and Aedes and Deinocerites cancer being assoc-

iated with thelcter. Numerous collections of thelcter have been taken in light traps

in the Florida Keys.

Aedes thelcter appears to be a significant vector of VEE in Texas. This is dis-

cussed under Medical Importance above.

DISTRIBUTION. Aedes thelcter is known from the Florida Keys and from the

Rio Grande Valley of Texas and southern New Mexico northeastward to central

Oklahoma. It undoubtedly occurs in northeastern Mexico; however, the records of

thelcter from Jalisco by Vargas (1956:23) and Los Mochis, Sinaloa by Martini

(1935:51) are probably in error. Material examined: 136 specimens; 16 males, 101

females, 4 pupae, 15 larvae; 4 individual rearings (3 larval, 1 incomplete).

UNITED STATES. Florida: Key West, 14-21 Oct 1946, [E. Fernandez], 7 F [USNM]. Long

Key, 5 Dec 1947, J. Haeger, 1 lpM (32) [USNM]; same locality and collector, 9 Dec 1947, 1 F

[USNM]; same locality and collector, 29 Dec 1947, 2 IpF (1,42), 1 lp (56) [USNM]. Texas:

Brownsville, 29 Aug 1916, M. High, 33 F [USNM, BM]; same data, 1 F (thelcter holotype)

[USNM]. Brownsville, 12-15 Feb 1924, R. Turner, 9 M, 13 F [USNM]; same locality and col-

lector, 20 Mar 1924, 1 M, 1 F [USNM]. Brownsville, Oct 1932, A. Niven, 1 F [USNM] ; same lo-

cality and collector, 5-8 July 1933, 1 F [USNM]. Brownsville, 15 Apr 1942, W. Reeves, B. Brook-

man and R. Eads, 1 M [USNM]. Brownsville, 10 Aug 1944, C. Joyce, 4 L [USNM]. Brownsville,

Apr 1940, 1 F [BM]. Brownsville, Brooklyn Mus. Coll., 1 M [USNM]. Corpus Christi, 29 June-8

July 1924, R. Turner, 5 F [USNM]. Glasscock Co., 23 Aug 1971, 6 F [JFR]. Howard Co., 9

Aug 1971, 9 F [JFR]. Mission, 23 Feb-8 Apr 1924, R. Turner, 2 M, 3 F,3 L [USNM, BM].

Pecos Co., 20 Aug 1971, 1 F [JFR]. Rio Grande Valley, Oct 1923, R. Turner, 2 F [USNM].

San Antonio, 16 Sept 1942, E. Ross, 1 M, 4 F [USNM]. Sonora, 12 June 1934, E, Cushing,

1 F [USNM]. Valverde Co., 11,26 Aug 1971, 8 F [JFR]. Webb Co., 2 June 1943, 1 Bosh

[USNM].

Additional Records from the Literature

UNITED STATES. New Mexico: Artesia (Miller, Doll and Wheeler, 1964:459). Oklahoma:

Anadarko; Clinton; Guthrie; Hollis; Walters; Watonga (Griffith, 1952:10).

SUBGROUP UNKNOWN

23. Aedes (Ochlerotatus) sp., Surinam form

Fig. 1

FEMALE. Medium sized species. Wing: 2.75 mm. Forefemur: 1.20 mm. Ab-

domen: 2.20 mm. Head: Decumbent scales of vertex and occiput white mesally

with large dark scale patch dorsolaterally. Proboscis entirely dark scaled. Thorax:

Integument dark brown. Acrostichal and dorsocentral setae confined to anterior

promontory; prescutellar and supraalar setae well developed. Scutellum with about

6 strong setae on lobes. Scutum with narrow acrostichal line of silvery white scales

from anterior promontory to scutellum; remainder of scutum with dark brown

Arnell: Scapularis Group 79

scales. Scutellum with large patch of silvery white scales on median lobe and small

patch of dark brown scales on lateral lobes. Apn scales entirely dark. Legs: Fore-

coxa with large patch of dark scales. Knee spots of white scales at apices of femora

absent. Hindtibia entirely dark scaled. Wing: Entirely dark scaled. Abdomen:

Terga II-VI with basolateral white scale patches; entirely dark scaled dorsally.

Sternum either entirely dark scaled or white scaled with dark scales distally on

segments.

MALE, PUPA, LARVA. Unknown.

SYSTEMATICS. This form can be separated from the other species of the Scapu-

laris Group by the narrow acrostichal line of silvery white scales with the remainder

of the scutum being entirely dark brown scaled and the absence of dorsocentral se-

tae except on the anterior promontory. It appears to belong to the Scapularis

Group but its placement within the group is uncertain, although it most closely re-

sembles the Bogotanus Subgroup. The 2 specimens of the form are indistinguish-

able from each other except for the scales of the sternum, the Colombian specimen

being entirely dark scaled and the Surinam specimen white scaled with dark scales

distally on each segment.

BIONOMICS. The Colombian specimen was apparently reared from a larva taken

from clear water at the grassy margin of a large pond in a pasture and associated

with several species of Anopheles, Uranotaenia and Culex. Nothing is known of the

bionomics of the Surinam specimen.

DISTRIBUTION (fig. 1). This form is known from Surinam and low elevation

Colombia. Material examined: 2 specimens; 2 females.

COLOMBIA. Meta: Laguna de la Palmita, nr. Villavicencio, 12 May 1947, L. Rozeboom

(CV 377), 1 F [UCLA].

SURINAM. Suriname: Zanderij, 17-22 June 1964 (SUR 116), 1 F [USNM].

24. Aedes (Ochlerotatus) sp., Colombian form

FEMALE. Medium sized species. Wing: 3.05 mm. Proboscis: 1.95 mm. Fore-

femur: 1.55 mm. Abdomen: 2.20 mm. Head: Decumbent scales of vertex and

occiput entirely white. Proboscis relatively short, length about 1.25 of forefemur,

entirely dark scaled. Thorax: Integument moderately light brown. Acrostichal se-

tae confined to anterior promontory; anterior dorsocentral setae absent except on

promontory; prescutellar and supraalar setae well developed. Scutellum with 4-6

setae on lobes. Scutal scales entirely dark brown. Scutellum with white scales on

midlobe. Apn with few dark scales; psp and ssp scales few. Legs: Forecoxa with

scales entirely white. Knee spots of white scales at apices of femora absent. Hind-

tibia entirely dark scaled. Wing: Entirely dark scaled. Abdomen: Terga II-VI with

basolateral white scale patches; entirely dark scaled dorsally. Sternum white scaled

with dark scales distally on segments.

MALE, PUPA, LARVA. Unknown.

SYSTEMATICS. This form can be separated from the other species of the Scapu-

laris Group by the entirely dark scaled scutum, the absence of knee spots of white

scales on the apices of the femora and the absence of dark scales on the occiput. I

have declined to name this species because of the absence of collection data and un-

certain placement due to lack of stages other than the female. It resembles incomp-

tus in most respects and may be related to that eastern Panamanian species.

DISTRIBUTION. This form is known from an unspecified locality in Colombia.

Material examined: 6 specimens; 6 females.

COLOMBIA. Locality unknown: (COR 494), 6 F [UCLA, USNM].

80 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

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1972. Agents encountered during arboviral ecological studies: Tampa Bay area,

Florida, 1963 to 1970. Am. J. Trop. Med. Hyg. 21:201-213.

Whitman, Loring and P. C. A. Antunes

1937. Studies on the capacity of various Brazilian mosquitoes, representing the

genera Psorophora, Aedes, Mansonia and Culex, to transmit yellow fever. Am.

J. Trop. Med. 17:803-823.

Willis, William, V. Pidcoe, D. F. Carroll and J. E. Satz

1974. Isolation of California group arboviruses from Pennsylvania mosquitoes—

1971, 1972. Mosq. News 34:376-381.

Wong, Y. W., D. C. Dorsey, M. J. Humphreys and W. J. Hausler, Jr.

1970. Arbovirus encephalitis surveillance in Iowa. Health Lab. Sci. 1970:117-

ies

Wong, Y. W., W. A. Rowley, J. A. Rowe, D. C. Dorsey, M. J. Humphreys and W. J.

Hausler, Jr.

1973. California encephalitis studies in lowa during 1969, 1970 and 1971. Health

Lab. Sci. 10:88-95.

Xavier, Sebastiao H. and S. S. Mattos

1970. Distribucao geografica dos culicineos no Brasil (Diptera, Culicidae). II.—

Estado de Mato Grosso. Rev. Bras. Malariol. Doencas Trop. 22:441-460.

FIGURES

1. Distribution of Incomptus, Bogotanus, Tortilis, Pectinatus and Thelcter Sub-

groups and Surinam form

2. Distribution of Trivittatus, Crinifer and Obturbator Subgroups

3. Distribution of Infirmatus Subgroup

4. Distribution of Scapularis Subgroup

5. Aedes (O.) incomptus; male and female head and claws; male foretarsus 5; fe-

male thorax, wing, legs and genitalia

6. Aedes (O.) incomptus; male and female scutum; male genitalia, pupa

7. Aedes (O.) incomptus; larva

8. Aedes (O.) bogotanus; male and female head and claws; male foretarsus 5; fe-

male thorax, wing, legs and genitalia

9. Aedes (O.) bogotanus; male and female scutum; male genitalia, pupa

10. Aedes (O.) bogotanus; larva

11. Aedes (O.) deficiens; female scutum, pupa

12. Aedes (O.) deficiens; larva

13. Aedes (O.) atactavittatus; male and female scutum; male genitalia, pupa

14. Aedes (O.) atactavittatus; larva

15. Aedes (O.) angustivittatus; male and female scutum; male genitalia, pupa

16. Aedes (O.) angustivittatus; larva

17. Aedes (O.) trivittatus; male and female head and claws; male foretarsus 5; fe-

male thorax, wing, legs and genitalia

18. Aedes (O.) trivittatus; male and female scutum; male genitalia, pupa

19. Aedes (O.) trivittatus; larva

20. Aedes (O.) tortilis; male and female head and claws; male foretarsus 5; female

thorax, wing, legs and genitalia

21. Aedes (O.) tortilis; male and female scutum; male genitalia, pupa

Arnell: Scapularis Group 91

. Aedes (O.) tortilis; larva

. Aedes (O.) auratus; larva

. Aedes (O.) auratus; male genitalia; Aedes (O.) condolescens; male and female

scutum; male genitalia

. Aedes (O.) euplocamus; male and female scutum; male genitalia, pupa

. Aedes (O.) euplocamus; larva

. Aedes (O.) patersoni; male and female scutum; male genitalia, pupa

. Aedes (O.) patersoni; larva

. Aedes (O.) infirmatus; male and female head and claws; male foretarsus 5; fe-

male thorax, wing, legs and genitalia

Aedes (O.) infirmatus; male and female scutum; male genitalia, pupa

. Aedes (O.) infirmatus; larva

. Aedes (O.) meprai and Aedes (O.) raymondi; female scutum; Aedes (O.) comi-

tatus; male genitalia

. Aedes (O.) phaeonotus; male and female scutum; male genitalia, pupa

. Aedes (O.) phaeonotus; larva

. Aedes (O.) scapularis; male and female head and claws; male foretarsus 5; fe-

male thorax, wing, legs and genitalia

. Aedes (O.) scapularis; male and female scutum; male genitalia, pupa

. Aedes (O.) scapularis; larva

. Aedes (O.) crinifer; male and female head and claws; male foretarsus 5; female

thorax, wing, legs and genitalia

. Aedes (O.) crinifer; male and female scutum; male genitalia, pupa

. Aedes (O.) crinifer; larva

. Aedes (O.) synchytus and Aedes (O.) obturbator; male and female scutum;

male genitalia

. Aedes (O.) obturbator; male and female head and claws; male foretarsus 5; fe-

male thorax, wing, legs and genitalia _

. Aedes (O.) obturbator; larva

. Aedes (O.) pectinatus; male and female scutum; male genitalia, pupa

. Aedes (O.) pectinatus; larva

. Aedes (O.) thelcter; male and female head and claws; male foretarsus 5; female

thorax, wing, legs and genitalia

. Aedes (O.) thelcter; male and female scutum; male genitalia, pupa

. Aedes (O.) thelcter; larva

SATIN ALALVLS

00s

0001

SYA LAWOTA

voruainy TIGA |

WO} WeULING »

o ie

4sn}Buljoed ¥

= ge

F 7 A

Le Z

4 dnoi3qng snyBuyyoeq hi be ala ees

3 pg eee aie

suatoljop=

_ dnoisqns snuej0s0g

= Ne

ie Ew am |

Oy snj}dwooule

oS. dnossqng snjduoouy]

it a

aS * |

dnoi3qng SIJIZIOL

S1]110}

snjeine-

o1sqns

J9}9]94}

J9}9[9Y f :

9,2

Cok el

OCHLEROTATUS

eee SG

V3 : : i

toa “a fe ene

Trivittatus Subgroup, | a St Pe

Re a cey

Ciaran i of can

etrivittatus. ae Ay oF Obturbator Subgroup _""”

fay)!

“Ss

otriv-angust hybrid Kat or

aatactavittatus

sangustivittatus

Equator

oe emenral..

synchytus4

j crinifere

Crinifer Subgroup

NORTH & SOUTH AMERICA

KILOMETERS

x 1500 2000 2500

1000 1500

STATUTE MILES

SATIW ALALVLS

00s

ooo! 00

SYA LAWOTIA

VOIWANV WIdd

snweoojdne=

———

e i

4 0

IpuowAvl=*. .

+ i

dnoigqne snjewstu

noszqng snypuyU]

SALVLOWATHOO

i ‘Sy a

RPE NT ae

OCHLEROTATUS

ee SM Flo

Scapularis' Subgroup

f GGG

NORTH & SOUTH AMERICA

KILOMETERS

STATUTE MILES

MOSQUITOES OF MIDDLE AMERICA UCLA - ZOOLOGY - ENT - 1971

tie

Aya

iyi

\ :

m\\\

ra y,

v ® y Yj

i \ by

ma Yy by; S

Yr wx. SS

Mm ee

iff Vy We ~S a,

,\ ~ —

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en ‘

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mK OC Mii Q

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PT RR ’ \ SS

a :

i 2c i

LANA NN % ~~ <

VW \

alls

aH R

HM RS

Blk

4 ia

‘i, x

—s

= =? :

fae i

Hh |) AN

if Y i

Ai Mf He

y i : y ih ji i i i

Hh 4 y fh y ae

bith | y an Mi

mt rah ya 2

Wh Hid in Hf CES

mt Hh) Bi SZ

Di.) hia 7, hh eZ

iin Hin Ae

Mi Hh Huy rh i

TH yi AR

he Hs Ha

Hh 1) Wil} ‘ |

ae i) i iy

Wf Hy Hi Li’ Wii

phil HI HH

q ! Hh , Whi! hii )

oy bhi \ Vi'\

i, t \ if, y}) r\ By

a Ws) WA,

ie bi | it

ae ie Hh

a of Ai

Ai We Wy 1 Wht

mh) We i i Hl

fh y THIN YY

HE Hi ip

ri Hi fk

in ae ih

MN yi Gis

Hi bi yf

NN Bi \ ah

Ahh Up Y fi) \

IN Y MM if \\

AN HAN Hi il

}\s {AAA \N aisle

ah ik\ Zan

Mm nt Sinn

a ht anal

{ N hh NN Ah

HN AS 1)

i, VN wt AW

AN a HN

ae) A

im (iN

AN)

i Wh AX iN

iN " LN cay

TN AN i XN

Li * HIS TAN

Ul h TN \ Ii N

Yy i] \ Hy i\h jis

AN ii Hin wit’

whi ji HIN

i in

Mj if \\f i \ A

if WA \ ANN Hal N

Wh \ Hit: Hj

iyi a y\ iW HAN

yi i A

Y An HW N mh Q

ft ®\ AWE Ji

i iN a \ PN

EM \N IWS iin

YP si. Hy) fee Hii

AN PS A

Fi \ TINS UVES

Ye TN AN

Yi AN we

Uf PHN HNN

Ae a ie

Lies A aN

De ai

4A wi HEN fat ae

E77) pi,\\ MN Fan

yy / iN

am we \ y, :

m HN yi)\ Wy 9

j \ Y oft . Uy 2 y

Di Wy] Y ‘ 9 6 com

Uy, TN f Uu S

Wii

Fei NN

BA -

Ai’ bys

Zh anama

Fig. 6

OCHLEROTATUS

4 i A %

incomptus

PA 379

Darien

Panama

Hig? |

OCHLEROTATUS

incomptus

PA 379

Darien

Panama |

VII

SG, A

a 6

1.0

719 12 1A:

i 1 7 pr h Ke

pia a ee

MID A

4 ‘

i } i, Z

i 4: i ii my OM ee A ee eee eS See

rh Z|

| Wn Wi WY; Wy .

| 1 Dr

MN tr Y) phi

iM ye Oh

A ii , )) y, Ih Tap

‘a ‘ Wows

Nit ! N\\ " . on Hf} | MOA it Wy ? \ \N

ma wi \ v ih | il I

TN Wis i\

Hine aN " bn i

a i i iN v Ri y

i i \ Ay} iN y

UNL AN ANY

Hijhstin iN

OCHL R sh Nr iin nh 7

a a 5 EROTATUS

\ ‘

ANT a i

AF ANA

; iy Ait WY ST

KARR epee rece RATT TY W FN

aeaeeRee

U, Z Vi

ply

WY yy

KO 200-15

Zag}

COR 530

HIND

bogotanus

Cundinamarca

Colombia

Fig.9

OCHLEROTATUS

6 x

SSS

Li een

TM \

bogotanus

DO are oe Ag SH

C4 ita (us: A g Cundinamarca

Cee i ‘|

ee i | Colombia

i’

| A’ i

bogotanus

COB 96

Cundinamarca

Colombia

a eid

2 As Ais

/ j Ala

Ce il

Ys a grO\\OX

YAY AS rf Y// D

INS IN FAX

NY 2 ‘

WW \ A ee

My pov) \

2 : A 24 Ca

Fig. Il

OCHLEROTATUS

deficiens

COB 92

Cundinamarca

Colombia

Figiz J \.

{ OCHLEROTATUS , , oe

deficiens

COB 92

Cundinamarca

Colombia

i ke

Cs 7 A ‘

XK t AMA

a4, Bape

Vill

ne \ by

top

Fig.13

‘ OCHLEROTATUS

MEX 609

MEX 610

atactavittatus

Campeche

Mexico

MEX 610

Fig. 14

OCHLEROTATUS

atactavittatus

MEX 609

Campeche

Mexico

VII

-sp

yk 2

7, Af}

Vill

POD

LA:

¥ An eee

: Fig. 15

OCHLEROTATUS

S ij}

74)

Af,

}+-—__——— 1.0 ————_—__ CR 64

Cartago

Costa Rica

Cartago

Costa Rica

angustivittatus

CR 482

Limon

Costa Rica

Fig. 16 ( |

OCHLEROTATUS

angustivittatus

CR 482

Limon

Costa Rica

hi \,

HAA Mh

iN A ‘

bi HNN

TTT

ih v ‘ "

YUN ian

‘ -\ OCHLEROTATUS

OWN “ \ :

; A

ANTS TASCA

WN ANNAN AN)

“zy

foe

Ze,

~ s

—

= =S SS

ANY Yi

A, pa Uf

Wee Vi fii

oo 2 fe

Ms Hi ie ONIN

“NN es Apel y Hy, yy

t uN i, sun IAN NS

AVN Y Ht Hira \

NR A HT dal

i a iN

N. HNy Ny

ws UR AM

us Ks ‘ y, f

4 aN pi)\ :

s\ IR \) i N

at ys Hi

Ny iy } nN Mx

4 q i N Ht

A hg yin

a AA

| an aa

Bill ISS HANS

Uh \ i i ie

AN ANN Hie

py t TN Is q

TAN WN. AiLAN)

yh a

“HN iN ARN

oo a

yi uN A

Bee any 7} a Yf t.

ii \ Yh N Hin

y)) WAN i ie

Ail) 0 /\ Nae

ie ~~ Uh

Jy gt WN HN

if aN he oot

| om iN trivittat

MY iis] Ri

AR y UCLA 736

bi Lip Illinois

A \\

WEN)

Vis \

LAN

JAR

— Viper {NY

Gy iy

Ait

4 \

Lie

__Z ue

Bat

Sypi

Spygay

EY.

_ Zu

“ul

— < RA

‘ Re, mee AON

eS PA

Nansen eon

—

SSS

9.18

o awe

c QCHLEROTATUS i)

ivittatus

UCLA 736

Illinois

USA

Fig. 19

OCHLEROTATUS

trivittatus le

UCLA 736

Illinois

Fig.20 *

SSIS

| Li yy

—

WH hin) ee < ERAN

Ii 177 Rapa: i aenn

Fe Sree REAR SERS Tl

Dai Hat haldaldttat KCN

SKN

. \ \ WA NN

‘ NANA XY

CX ARH

QS SSN Gs uN

= Se 2 ~ : Y NS ANY SY

“%

\ Wig

=—S

ZZ,

Nj fk RDO 272

Ly Dist. National

if Dominican Republic

Dist. National

‘Dominican Republic

Why Gin

Wi Ms

—s

—

—S Ss

SE a Re

SS SSS SSS ~~

WO SS SSS SS

SS a SSS

= SSS SSSA : =

SSS SSS SSS SS ™

——— =: WS >

SS SSS “

PUN

i Aig

. vik \

a Hi} Va

\ ‘ Hy bi \

‘ fi, hi

ia My iN

at ANS

ih \ a

AN Hae

H) \N ih Ws

| aes

Be i ('\\ \

i WN

ih A aN

i ‘ \ IN ZA St. Catherine

fi \ YUAN 4 Jamaica

ry WK \ \ JA

BRS HD

ai KN \

fats Ah VN

Lhi\ HIN

Li aN dN

HS At N

a \\ i »

ARS TN.

__Ie® ERS

yi HN

Y; yay Ni

TAs HAN

ae ih

wi \

¥ ‘ \

Samana

Dominican Republic

e Fig. 21

OCHLEROTATUS

! 2)

Bagcatas. “> ‘

eat

Be IN

nm N

— Ae

— : Zip 1 10 \

Z///\\e -

(UY

py UU vy

St. Catherine

Jamaica

tortilis

JA 796

St. Catherine

Jamaica

JA 796

St. Catherine

Jamaica

tortilis

jo}

—

OCHLEROTATUS |

Fig. 23

OCHLEROTATUS

ye TS

Voy a Jo F

SNe

auratus

St. Andrew

Jamaica

auratus

Kingston

Jamaica

\ AY

Xy x XN

N «J ®

ae “t

NA , < 5)

OCHLEROTATUS

S —_ 7 :

D2 BA >

New Providence

Bahama Islands

condolescens

( Fig, 25

| OCHLEROTATUS cc

me a |

CZ 49

Canal Zone

Panama

Guanacaste

Costa Rica

et)

euplocamus

CR 309

Alajuela

Costa Rica

Fig. 26

OCHLEROTATUS

euplocamus

CR 309

Alajuela ;

Costa Rica fe

Fig. 27

OCHLEROTATUS

ARG 742

Salta é

Argentina

SN 1

SSN

ARG 755

ARG 755 e : i

Salta

Argentina paterson!

Fig. 28

OCHLEROTATUS

patersoni

ARG 742

Salta

Argentina

Pr, my

an|

S ~

sally

VE ince

Lisa aint

. eR \\

Ratt ANNA

—— SSS :

ea ae

N : SS

SASs

S SS Hf

Louisiana

USA

South Carolina

USA

FLA 41A

Florida

USA

Fig. 30

OCHLEROTATUS

Florida

USA

-_—_————_-1.0 —__—__"4

Louisiana

USA

Fig. 31

OCHLEROTATUS

1.0

11 9

13 10 a

infirmatus

Florida

USA

=, t Heil 1g |

Plas =

1 50 &

Fig. 32

OCHLEROTATUS

aiskanlan ea emaamceaaigel. “Ce meme I ae as

raymondi

Salta Jujuy

Argentina Argentina

comitatus

KO 200-2

Meta

Colombia

| Fig. 33 \\

OCHLEROTATUS

GRR 82

Grenada

SZ Ma

ae d [See ee Ws

6. =~ "M@il)) >

De {l) a |

it Zoe

GR 104

GRR 82 eT / Eon AS ‘ es aa Vee Grenada

Grenada SE ssa it

Fig. 34

OCHLEROTATUS

phaeonotus

GR 104

Grenada

SSS

SSSssxsS&.a.

SSN a

= SSS S

EE Ee =

<==

ae me

—-:

SSS

BEA

ess

SSS

SS SSS

SSS Ss

> SS

Piast a rcarh

——

}

Hiss,

HA: «

SSS SS

fe scapularis

’) ie: m

Uh |,

f | ll i iW Dy ivan ’ “ ~~. i

ih mM Aan \

OCHLEROTATUS Li }

SS NN

‘ AN PN \\\S

hy i i WAYNE « ~ — >» A NC

N \,

)

nt SS ash

Hii Wi

iy Ny

‘a ay

pe Wf y] Wi y)

Mato Grosso

Brazil

YO)

Ui a

SSS

La y : Mato Grosso

Li Brazil

FORE MID HIND

Bahia

Brazil 0.3

. Mato Grosso

Brazil

Fig. 36

OCHLEROTATUS

ARG 742

Salta

Argentina

he >

pee

Mato Crosso ~“S

Brazil

BRA 238

Rio de Janeiro

BRA 237

Rio de Janeiro

Brazil

Mato Grosso

Brazil

Fig. 37

OCHLEROTATUS

scapularis

ARG 742

Salta

( i \

N\ss\

MY yyw

y y — Re. AN AS

an i i i | m AT iy i‘ K eas wT AN \\ \\ nN

wi TUT etre pr CT

We ' \OCHLEROTATUS SN iiiiANNNNNeA aN Rae URS

vf NY

i IN h

Hah’

ith NN \\ = >

“ Ss

a SS

ni SSS

ma SS

Hi SS

i ANN SSs—-x

HUN 4 “

a SS

ANN

Corrientes

Argentina

Hips i

ip Hy i

YA ai 4)

ie |

Misiones

Argentina

yis')N

PANS

| Fig. 39

Ee OCHLEROTATUS

BRA 167

Sao Paulo

Brazil

Mato Grosso

Brazil

crinifer

Sao Paulo

Brazil

Fig. 40

OCHLEROTATUS

crinifer

BRA 167

Sao Paulo

Brazil

VIII

3 *T® lan

synchytus

Misiones

Argentina

Eleuthera Island

Bahama Islands

Mona Island

Puerto Rico Mona Island

Puerto Rico

Ly < obturbator

me AD in

RNS

a : eS

— SEI \ AN AAS

os iN :

VA Pa ANE PARA

mi Nh KAMAN

SSS pee

\\"

with

‘\ \ > . ‘ Nh \\

\ ee ONY

in i eS ¥ ok

Vgc rae

Wat \ ran

i - 4 \

ih me R

iM SSS ; .

Mi G

Lh Ny

AWN

LSS

WSS

SSS

ESS

Mis \

Ws:

Eleuthera Island

Bahama Islands

Ss PEt

tuite Z

SWSSsssSss

SSS

obturbator

Mona Island

Puerto Rico

Fig.43 3 \:

OCHLEROTATUS

obturbator

Mona Island

Puerto Rico

VIll

{) Fig. 44

OCHLEROTATUS |

pectinatus

COB 1

Cundinamarca

Colombia

— Fig. 45

OCHLEROTATUS

Vill

7 ule

p ay 3

| LA

Y), 1 CD

ft \ p y

AA \\\)

fl . AS {| a ie) i

= y, | \ Yi

vA i

Nes W)

OB 1

Cundinamarca

Colombi

ig. 46 =e

) | \y Hh ee ® Se oe a ® RN

: Hi " ey Mes, | ~~ Leen

OCHLEROTATUS

ANNA TTT TRRRRERR Strrmm AWN AN

(TT eR

SQW

7 Capa aS

——es SS

SSS, >

SS >

SS SS SSS

\N BAIN

iN mi k

‘) a

AN ‘ iN \ \ :

Tye Lyin \\Y AN

Zi Y) Vs RN \ ‘ \

H\\ hi A®.\ WN

aiK\ Hin

Bh \N LAN

aan theicter

vii N N\,

m (fF

eA Sy

| Texas

USA

thelcter

Texas

USA

Fig. 48

OCHLEROTATUS

| thelcter

a Texas

USA

jon

USA

BAHAMAS

GREATER ANTILLES

LESSER ANTILLES

MEXICO

GUATEMALA :

ECUADOR

PERU

VENEZUELA

TRINIDAD

GUIANAS

BRAZIL

BOLIVIA

ARGENTINA

PARAGUAY

URUGUAY

Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

TABLE OF DISTRIBUTIONS

: . 2

= 3 ai & = =

= cS co os > ed) ”n

=| = os oS a 2 > = -

S) o mS O Sy = a= Ss cS

O Se o D fi 3 - =

& te ao e & a 5 = =

= NN oa) st aN Ne} ~ ore) ON

CANADA LE Bee aS

North :

Southeast

Southwest

North

South

Ria as

Pan 2a

oe.

Rech

Pao) ae

lesa ae

ot. it lel |

Low elev.

Amazon

Southeast

141

Arnell: Scapularis Group

TABLE OF DISTRIBUTIONS

19}9[9Y} *7Z

snjeurjood *{7

10}e€gGINIGO “OC

Sn}iAYOUAS "6 ]

IOJIULIO “QT

SN}eWIWOD */ |

suie[ndeos 9]

snjouoseyd *c]

SNPEUWITUT “Pp |

IPUOWABI ‘¢]

Tuosiojed °7 |

snuleoojdns *[ |

SUdOS9TOPUOD ‘O]

142

Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

CONSPECTUS OF TAXONOMIC CHANGES

NEW TAXA

BeACEAVILESTU oa a a a 2G

bocetanus 40k ee ee ee 2

CONMEAGIE Ue ay en OG

deficiens = Fok a oe eS et

WCOMIPIS ae a ee

Pecttianies 30 ee ee eet

phaconotis owe ee eer DO

SCR VES oe ee Al

CHANGES IN TAXONOMIC STATUS

CHIMPOSAKUS, BYNONVIIY 2 Re a OT

Nernuisurus, SVHOAVINY 6 ce ee OT

Meprat to specilic- rank yk. OL ee oe es 28

VERVGCOPNIIIS, SYNONVYY 6 oe ne ae

Arnell: Scapularis Group 143

INDEX TO SCIENTIFIC NAMES

Aedes, 2, 6, 14,31, 36, 60, 70, 78

Aedini, 70

aegypti (Aedes), 50

aenigmaticus (Aedes), 7

angustivittatus, 3, 4, 7, 8, 9, 17k, 18k, 20k,

26, 27, 28, 29-34, 37, 67; figs: 15, 16

angustivittatus of authors, 28, 59

Anopheles, 31, 60, 79

argentescens, 3, 29,31, 33

argyritarsis (Anopheles), 57

articularis, sp near (Culex), 75

atactavittatus, 17k, 18k, 19k, 26-28; figs. 13,

14; 142

atlanticus (Aedes), 7

auratus, 3, 15k, 18k, 19k, 39, 41, 4446;

figs. 23, 24

balteatus, 3, 4, 39,40, 41, 42

bejaranoi (Aedes), 7

bicristatus (Aedes), 70

bogotanus, 7, 16k, 18k, 19k, 22-24, 25, 36,

41, 47, 49, 51; figs. 8-10; 142

Bogotanus Subgroup, 6, 16k, 22, 79; fig. 1

bracteatus, 3, 4, 39, 40, 41, 42

brethesi (Culex), 70

busckii (Aedes), 57

calumnior (Aedes), 7

camposanus, 3, 6, 58, 60, 63, 142

cancer (Deinocerites), 78

celeste (Haemagogus), 57

ciliata (Psorophora), 60

cingulata, sp near (Psorophora), 57

Colombian form, 16k, 79

comitatus, 15k, 18k, 19k, 56, 66-67; fig. 32;

142

condolescens, 3, 4, 15k, 18k, 19k, 23, 36,

41, 46-48, 49, 515 fig. 24

condolescens of authors, 40, 59

confinnis (Psorophora), 78

confirmatus, 4, 57, 58

confirmatus of authors, 69

corniger (Culex), 27

coronator (Culex), 27

coronator, sp near (Culex), 70, 75

crinifer, 3, 4, 14, 17k, 18k, 19k, 68-71, 72;

figs. 38-40

crinifer of authors, 51

Crinifer Subgroup, 6, 16k, 67-68; fig. 2

Culex, 4,31, 50, 60, 70, 79

Culex species, 52, 60

Culicada, 4

Culicidae, 4

Culicini, 70

cuneatus, 3,4, 29, 31, 33

cuneatus of authors, 35

cyanescens (Psorophora), 27, 78

deficiens, 7, 16k, 17k, 19k, 22, 23, 24-25;

figs. 11, 12; 142

discolor (Psorophora), 78

dolosa of authors, 69, 70

dorsalis (Aedes), 78

dupreei (Aedes), 4, 7, 70

eucephalaeus (Aedes), 7

euiris (Aedes), 4, 7

euiris, sp near (Aedes), 24

euplocamus, 4, 15k, 18k, 20k, 23, 36, 41, 46,

47, 48-50, 51, 52, 58, 67; figs. 25, 26

euplocamus of authors, 58, 59

evansae (Anopheles), 70

ferox (Psorophora), 27, 52

fluviatilis (Aedes), 75

habanicus, 3, 39, 40, 42

habilitator (Culex), 41

hastatus (Aedes), 7

hemisurus, 6, 58, 60, 142

Heteronycha, 4

hortator (Aedes), 4

iguazu, 68, 69, 70

incomptus, 16k, 18k, 19k, 20-22, 79; figs.

5-7; 142

Incomptus Subgroup, 6, 16k, 20; fig. J

inconspicuus, 3, 4, 35, 38

indolescens, 3, 58, 63

infirmatus, 4, 7, 8, 9, 15k, 18k, 19k, 36, 46,

48, 53-55; figs. 29-31

infirmatus of authors, 35.

Infirmatus Subgroup, 6, 15k, 46, 543 fig. 3

ioliota (Aedes), 57

jocasta (Culex), 57

jubifer, sp near (Culex), 22

keyensis, 76

Leucomyia, 4

lynchii, 68, 70

magnus (Deinocerites), 41

meprai, 17k, 19k, 26, 28; fig. 32; 142

144 Contrib. Amer. Ent. Inst., vol. 13, no. 3, 1976

milleri (Aedes), 7, 25

nigripalpus (Culex), 27

nubilus (Aedes), 7

-obturbator, 3, 4; 7, 16k, 17k, 19k, 26, 72-74;

figs. 41-43

Obturbator Subgroup, 6, 16k, 723 fig. 2

Ochlerotatus, 2, 4, 5, 6, 7

oligopistus (Aedes), 7

patersoni, 15k, 18k, 20k, 23, 36, 41, 46, 47,

48, 49, 50-52; figs. 27, 28

pectinatus, 7, 14, 16k, 17k, 19k, 74-75; figs.

44, 45; 142

Pectinatus Subgroup, 6, 16k, 74, fig. 1

pertinax (Aedes), 7

phaeonotus, 15k, 18k, 20k, 56-57, 59; figs.

33, 34; 142

pilosus (Culex), 27, 70

plutocraticus, 3, 4, 39, 40, 41, 42

Protoculex, 7

Pseudohowardina, 4

Psorophora, 31, 60, 78

Psorophora species, 22, 60

quinquefasciatus (Culex), 31

raymondi, 3, 15k, 17k, 19k, 46, 52; fig. 32

rhyacophilus, 6, 58, 60, 142

Sabethes, 9

Sabethini, 70

scapularis, 2, 4,6, 7,9, 15k, 18k, 20k, 27, 36,

47, 52, 56, 57-66, 67, 72; figs. 35-37

scapularis of authors, 48

Scapularis Group, 2, 3, 4, 5, 6, 7, 9-14, 21,

23, 25, 21; 30,36, 45, 47, 49.51, 54; 57,

59.60, 61567,09, 70; 71, 72, 73, 14075.

Weee,

Scapularis Subgroup, 6, 15k, 55-56, 57, 67;

fe. 4

scutellalbum (Aedes), 7, 75

serratus (Aedes), 4, 7, 9

Serratus Group (Aedes), 7

shannoni (Aedes), 6, 7

signipennis (Psorophora), 78

sollicitans (Aedes), 78

splendens (Haemagogus), 57

sticticus (Aedes), 37

Surinam form, 16k, 78-79; fig. J

synchytus, 17k, 18k, 19k, 68, 70, 71-72; fig. ~

41; 142

Tamarix, 77

tapinops, 68, 70

thelcter, 3, 7, 8, 9, 14, 16k, 17k, 19k, 36,

76-78; figs. 46-48

Thelcter Subgroup, 6, 16k, 76; fig. 1

titillans (Mansonia), 31

tormentor (Aedes), 7

tortilis, 3, 4, 15k, 18k, 19k, 23, 36, 39-44,

45,47, 49, 51, 60; figs. 20-22

tortilis of authors, 45

Tortilis Subgroup, 6, 15k, 38-39; fig. J

traversus, 3, 29, 34

trichurus (Aedes), 70

trivittata of authors, 35

trivittatus, 3, 4, 7, 8, 9, 17k, 18k, 20k, 23,

26, 27, 28, 30, 31, 35-38, 41, 47, 49, 51,

54, 73; figs. 17-19

trivittatus of authors, 29

Trivittatus Subgroup, 6, 16k, 25-26, 27, 46,

D4; fig, 2

Uranotaenia, 50, 60, 79

Uranotaenia species, 24

vexans (Aedes), 37, 78

virginensis, 3, 39, 40, 41, 44

Contributions

of the

American Entomological Institute

Volume 13, Number 4, 1977

MEDICAL ENTOMOLOGY STUDIES - VI.

A REVISION OF THE SUBGENUS LOPHOCERA OMYIA

OF THE GENUS CULEX IN THE ORIENTAL

REGION (DIPTERA: CULICIDAE).

Sunthorn Sirivanakarn

CONTENTS

a aS nak a Gee ticca ant tees a a laa oe) Sorte ge PE Geo Ay es ge 1

BA a HON ns a, ee SR aw Rai ee dae a noe Blige ee eee ee a

EEE ee de IEG A PRE 06 Se, eau ks Roos ges a oe ew wee 2

ee ke Ee Py hp ce yh ca ek | Behe en de Wee ce 4

Serene POPOL ENA 2 ee oka gs lve ee ee ee 4

Keys to Groups of Oriental Loprocevaomyia. . 6. 6 ee ee ee ee 12

eee i Rr COE ee eg ete a is ek ae cas bce Rk Ww wae BO we 13

Keys to Subgroups, Complexes and Species...........246. 16

VETS S175 WO VOUS S CIPS GS. bow BLES Segre BRE aL

Te) RUSS tries GeO). se So we oe Sou ee ke Be 22

Be UL OP CR Ee sata ki tak Ree te te Raa Ok, BR ae. Ae vas

ic HMC Ae ee 4 gag Sab ee: Be ew ees 26

A BU VOM a oe eh os AED ere telieeseat ee sin eee Bs 29

Ae? GORIOF) BAPLONE ciienia-decienes 0s ay 4 Me iy ern Se 31

Den CHIChELIUS POWAVOS 4) 6 8c hil sowie Dae eae ler els Ae eae aa 5

ee RO a) gs egret al ed Seige Mafdis. a doe we eve, oul 36

Le TUDE OVOCTS (LIEICESLEY )ii5 iis bck Secce ca eae bo, Hide eee wk Sal

Be eV ETE OL Oi 2 gue eh gale a eS Gece we ee we 40

Oe he Ue AC a Ma SU ame an ata elie hile aye ee 41

DO IDLE MOP Oe ye Yue aloe yale a et a? os) oe eet oboe 8 43

Pies teers TaWwarde) sg a. bk eG Sle cea wie bk ee > 44

Do re Sy el gg eee ge be ee SO Se eutel kl es ee wey 48

Bee eres MAC gs pike ha Pad Cet eee BR Sa glial well 49

We LCG EEU He Wire Be es kan, a aarti ae pclae tee tae caine? gt QOL, Choe ad ag ore 20

EDs) MCCO? We OLB BS «ea bk> area we 2 NAA aa FS: 4 0 MMe ol Yah eS YS ol

Rs Or ESIC GMb R ig.) oocbe. MOR Kee Rw ee 03

LW jOSC PUTAS BAISaS Se a We SP a GR oie eset Desh 06

BO O71 1a tis COMES fees sii elia lg en ad eran ah, Pate eo he 2 oO”

TOUR eC Se BO ee oN got Le ety 6 a be wae 58

Pe IO GT OHO UO OG) ate is. eG ale, wencetes ee tee eS, eos eve 59

Bs HMO Oe OU CSS ec ais Gn i oi sek a ee ek te 61

Eee Gd hE Atari alle rica he eB ce sie gt sol Se terla GR. OR 63

RUNS, BR SERRE NS aT aca a ame a a 65

Vy ee CR yoy ne cal gw wots wee te wl oe ee ee 67

Keys to Subgroups, Complexes and Species. .........0.6-6 69

WATE? GOOYOUD facts Ge Sn a we GS ee oe eee ee 78

VE GSO Ma TOO eh ea ey ae ek a ee ale ek ee 80

DES TERR Fn) 11) BSS E) Ge hs eae Re ce) pe ag an cet ES ea at, a a a nen ee a 82

Ge. MAEFO ret Si TATOO AIA) he Boma lank AS WOE eet ale ce oe 84

Zier: EO Od On er ls pe eae WO we a ee 86

28 Mee Fer A PIGOGLET) oe rera$ seers a eet aye eR we 87

Ce re te a earl pg Cerne ote agi gag) eb. ag) A oS 90

or MCR TEGUS OC OLLOR GR Ugh Eee ma el GONNA RG BeBe ee os 91

Ot. eee MOLL GMs toe eta See RTE eR BESS giana es ew 93

32... Spiculosus Bram,and Rattanarithikul .. 2. 2 2 2 2 ee 96

Boost a ie CORI cu ca ate e ee lara Rk Ge eo Re 8 98

4. COV HS | CORIO) Wy aos ee aha oak we Sec eee Oe a 102

BO ROCCHI STS BeV a Se ge ee a) ee ay 104

ye SURI Yh oe 2 eos 21S) is SRM alae SRD we girth) es tN” UE Pee er ae 105

CONTENTS

37, panmst Kine and Necestraal oui sia Se ao oe ee

33. CYOSSICOINUS COMERS Git EE IO OR oe

39. incomptus Bram and Rattanarithikul..........

40°). benerpensie Paria ty fos anak eas ote en ea

41. peytoni Bram and Ratiandrithiol Spe. ee.

42. eukrines Bramand Rattanarithill.:.... 20. 84 2

43. pholetey Bram and Rattanarithikul., 2. . 06.40. 2

$4. JIACLCOPNIS- Berra Fa: esis, eo ee lc ng

4. (Seema Hy Ste gk en ean ae ge nena

Brevihaipus Gao evomes cae iy seas Oe ee as FE oe Se ALS

40... GIQHTIS Tawars ee Soe Ge eee ara ee ee ede

Bis \COCTIHOACOIS FWA, he can Dee ae a a eas ee

40, Rew Cia ese ree Sees ees ee Sas

49. -jensoni (de Meera ei eget a) ah ew whee ete ea

D0: - Derepepd ius TP heobeldy). wih hae ee es ate ee ee

obs deer is COESS hen i wR aieks Babs Oe eae a

52, eminent tieicesteryo no Jarek. .techins! See

Dos CHE POLINA ACO Be. aad rer deca Ssh ee Bowes

De ONE Peer Umass cry te sd) Seer aOR a a

Ds =| SHORT GNSS ABTS} is a5. et? Gh aa. Gee ke oa “ea

WILFRED) GROUP see Awe Ps th Song otis ale eS 5 2 OR i ee eee,

REVS 10: SpO0TR8 i ees Has, 6 20S Se I ee

Gy Tea pee CORR BSS eee Wy a MS es Ba ee

nl, pilifemoralis. Wang and: Feng. 2. Goa Gow.

BOs avid Lads Ua Bg ae oe tea eae Bek

BANOO ste Gh BU tho ge a tia Ae ee BP ae SE

BICERAL Re Cte ecdson see ie oe eine eae a ee

bie? OF PIGURES, 2a 2, ose eo aig Tae al, ee ee

PCRS gels ee Aa es a es ae ee

APPENDIX A. TABLE OF DISTRIBUTION AND BREEDING

ERPASE EP a I a BE ge ee ig PS ge PU de

ili

MEDICAL ENTOMOLOGY STUDIES - VI.

A REVISION OF THE SUBGENUS LOPHOCERAOMYIA

OF THE GENUS CULEX IN THE ORIENTAL REGION

(DIPTERA: CULICIDAE)!,

Sunthorn Sirivanakarn2

ABST RACT

This revision is based on the study of some 8, 000 specimens (3, 789

males, 2,521 females, 77 pupae, 1,616 larvae). Included among these are

3,115 adults with correlated pupal and/or larval skins (1,314 p, 1, 801 lp)

associated through individual field rearings.

Altogether 58 species are recognized and of these, 9 are new; 49 are re-

validated and redescribed. The new species include: @lorensis from Alor,

Indonesia; pavaculeatus from Sabah, Malaysia and the Philippines; aestivus

from Sarawak, Malaysia; gvacicornis and pairoji from Peninsular Malaysia

(Malaya) and Thailand; impostor from Sabah and Sarawak, Malaysia; wardi

and lasiopalpis from Sri Lanka and hirtipalpis from Thailand. Keys to all

available stages and illustrations of every species are provided. The taxonom-

ic treatment includes a revised morphological characterization of the sub-

genus, followed by a review of distribution, taxonomic discussion, scheme of

internal classification, bionomics and medical importance. The groups, sub-

groups and complexes, as adopted and currently developed here, are defined and

characterized. The descriptions of 49 species are based on all stages (female,

male, pupa and fourth instar larva) and those of 9 other species are based

only on the males and presumptive female of one species.

New synonyms proposed include: pachecoi Baisas (= quadripalpis (Edwards));

fuscosiphonis Bram and Rattanarithikul (= demissus Colless); hui Lien (= spi-

culosus Bram and Rattanarithikul): plantaginis Barraud (= minor (Leicester))

and uniformis mercedesae Baisas (= kuhnsi King and Hoogstraal). Culex bi-

cornutus(Theobald)is resurrected from the synonymy with minor (Leicester);

barkerii(Theobald) bernardi(Borel) and mindanaoensis Baisas are reduced to

nomina dubia.

I This work was supported by Research Contract No. DAMD-17-74-C-4086

from the U. S. Army Medical Research and Development Command, Office

of the Surgeon General, Washington, D. C.

2Medical Entomology Project, Smithsonian Institution, Washington, D. C.

20560.

2 Contrib. Amer. Ent. Inst., vol, 15, no. 4, 1977

INTRODUCTION

The genus Culex, subgenus Lophoceraomyia is an Old World taxon with

characteristic distribution confined to the Oriental, eastern Palearctic and

Australasian regions and the islands of the western and south Pacific Ocean.

Of all the subgenera of Culex in the Oriental region, Lophoceraomyia is ap-

parently the richest in number of species. The majority of these occur in

Southeast Asia and neighboring areas where they form one of the common

elements of the culicids of tropical rain forests. After the recognition of the

group by Theobald (1905a, 1905b), the subgenus has been the subject of several

taxonomic studies, but most of these are strictly local, or limited to the des-

criptions of species and additional descriptions or records of previously known

species. A number of regional studies prior to 1960, in which the group was

treated as a unit include Leicester (1908), Borel (1926, 1930), Barraud (1934),

Baisas (1935), Bohart and Ingram (1946) and LaCasse and Yamaguti (1950).

Since the discovery of arbovirus infections in wild-caught adults by the U. S.

Army Medical Research Unit in Peninsular Malaysia (Anonymous 1960), fur-

ther significant intensive studies of local species have been made. These in-

clude an excellent review by Colless (1965) in Malaya, followed by the studies

of Delfinado (1966) and Baisas (1974) in the Philippines, Bram and Rattanarithi-

kul (1967) and Bram (1967a) in Thailand. Other works dealing only with des-

criptions of new species during this later period are Wang and Feng (1964) in

Yunnan, southern China; Rahman, Choudhury and Kalra 1968 (1969) in India

and Lien (1968) in Taiwan.

During the past 10 years since the Southeast Asia Mosquito Project (SEAMP,

1964-1974, currently replaced by the Medical Entomology Project (MEP)) was

initiated, a considerable amount of Lophoceraomyia material from various

parts of the Oriental region has accumulated and more information on distribu-

tion, bionomics and medical importance of many species has been obtained.

This material, which represents the largest collection from the area, consists

of numerous reared specimens of nearly all previously known species and a

number of undescribed species. A long term study of this material together

with my subsequent study of the various types and topotypic specimens of spe-

cies from India, Sri Lanka and other countries at the British Museum (Natural

History) and the U. S. National Museum now make it possible to attempt a

broad regional revision of the subgenus.

The present revision includes almost all species previously described

from the Oriental region. A total of 58 species are treated. Of these, 49 of

the previously recognized species are redescribed and 9 are considered as

new. Approximately 8,000 specimens | 3,789 males, 2,521 females, 77 pupae,

1,616 larvae; 1,314 p, 1,801 lp] have been examined. Forty-nine species are

known and described in all stages and 9 other species are described only from

the males, including one also with presumptive female.

MATERIAL AND METHODS

The largest portion of the material for this study came from the extensive

collections made since 1964 by the Department of Medical Entomology, U. S.

Army Medical Component--SEATO, Bangkok, Thailand; the Malaysia Mosquito

Project, Department of Parasitology, University of Malaya and by several

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 2

individuals from U. S. Army Medical units, local governmental agencies and

private institutions in the Philippines, Malaysia, Vietnam, Indonesia and the

Ryukyus, Japan. Numerous additional fresh adult specimens, both field-

caught and reared have been obtained from the recent collections in Sri Lanka

by the Smithsonian Institution's 'Biosystematic Studies of the Insects of

Ceylon" project, in Indonesia by S. Ramalingam, University of Malaya, T.

Kurihara, Teikyo University and Yoshito Wada, Nagasaki University, Japan,

in the Andaman Islands and Madras, India by N. L. Kalra and B. N. Mohan.

The remaining specimens examined were largely field-caught adults, repre-

senting the type or parts of the original type-series and topotypic specimens,

from loans made to the project by the British Museum (Natural History); Divi-

sion of Entomology, CSIRO, Australia; Bishop Museum, Honolulu; Institut

Voor Tropische Hygiene, Amsterdam, the Netherlands; Field Museum of

Natural History; California Academy of Sciences; Academy of Natural Sciences

(Philadelphia) and the collection of the U. S. National Museum.

Vital to this study were the collections of the British Museum (Natural

History) and U. S. National Museum. The collection of the British Museum

was most important because of its broad representation of species and accumu-

lation of the types (holotypes, lectotypes) and other specimens in the original

type-series. The collection of the U. S. National Museum has become increas-

ingly important in recent years. This collection contains all holotypes and

most of the specimens in the type-series of the species described by Bohart

(1946), Stone and Bohart (1944), Delfinado (1966) and Bram and Rattanarithikul

(1967) and several paratypes deposited by Colless (1965) and Lien (1968). In-

cluded also are a few types and some topotypic specimens of the species des-

cribed by Ludlow (1909), Dyar (1920) and Baisas (1935) from the Philippines.

Among the recent collections which significantly contributed to a better

knowledge (biological, taxonomic, distributional) of several previously known

species, 4 deserve special mention here. The excellent collection of the

Malaysia Mosquito Project directed by Shivaji Ramalingam is well balanced,

containing many reared specimens with adequate associated pupal and larval

skins of most Malaysian species and the material collected and prepared is of

high research quality. The collections throughout Thailand of the SEATO

Laboratory directed by John E. Scanlon (1960-64) and Douglas J. Gould (1964-

76) and the earlier collections by D. C. and E. B. Thurman (1952-57) are the

largest of all, containing numerous field-caught (by light traps or nets) and

reared specimens of most local species. The private collection of the late

F. E. Baisas in the Philippines, which was donated to the U. S. National

Museum in 1968, has a good series of reared specimens of locally common

species but most of these were from Subic Naval Base in Luzon. Finally, the

most recent collection by E. L. Peyton and Y.-M. Huang in Sri Lanka of the

Smithsonian Biosystematic Studies of the Insects of Ceylon, is perhaps the

most comprehensive collection ever made in that country. This collection has

numerous reared specimens of nearly all known local species, additional new

records of the species known elsewhere and some undescribed species.

The format of presentation and descriptions of species in general follows

the preceding Culex papers by Bram (1967a) and Sirivanakarn (1972) and sub-

sequent papers by the latter author. The terminology of important taxonomic

characters used throughout were those of Sirivanakarn (1968), which were

modified from Belkin (1962) and Colless (1965). The only changes which have

been made here are certain features of the male genitalia. To simplify the

description of the specialized setae and leaflets of the subapical lobe, the

alphabetical designations such as a, b, c, etc., as used by Belkin (1962) are

4 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

adopted. In describing the lateral aedeagal plate of the phallosome of the

species in the Mammilifer Group, I have introduced the term ''external pro-

cess" to substitute the "dorsal process" used by Colless (1965), simply to dis-

tinguish it from the other process, the internal process of the same author.

TAXONOMIC TREATMENT

SUBGENUS LOPHOCERAOMYIA THEOBALD

Lophoceraomyia Theobald 1905a: 245. Type-species: L, uniformis Theobald

1905a, Ceylon (Sri Lanka); monobasic. Considered as a lapsus for

Lophoceratomyia by some authors.

Lophoceratomyia Theobald 1905b: 93. Type-species: L. fraudatrix Theobald

1905b, New Guinea; the first of 2 included species, selection of Brunetti

(1914: 64). Considered as a lapsus for Lophoceraomyia by some authors,

as a valid emendation by others.

Philodendromyia Theobald 1907: 623. Type-species: P. barkerii Theobald

1907, Sarawak (Malaysia); monobasic. :

Cyathomyia de Meijere 1910: 921. Type-species: C. jenseni de Meijere 1910,

Java (Indonesia); monobasic.

Culex (Lophoceraomyia) Theobald, Bohart and Ingram (1946: 1-110); King and

Hoogstraal (1955: 1-11); Bohart (1956 (1957): 66-70); Dobrotworsky (1957

(1958): 317-21); Stone, Knight and Starcke (1959: 232-5); Stone (1961: 45);

Belkin (1962: 248-72); Stone (1963: 133-4): Assem and Bonne-Wepster

(1964: 111-3); Colless (1965: 261-307); Delfinado (1966: 101-15); Stone

(1967: 213-5); Bram (1967a: 42-113); Sirivanakarn (1968: 75-186); Baisas

(1974: 111-22).

Culex (Lophoceratomyia) Theobald, Leicester (1908: 18-261); Edwards (1924:

396-7); Borel (1930: 352-64); Edwards (1932: 196-8); Barraud (1934: 359-

76); Baisas (1935: 167-79); Feng (1938: 296-9); Carter and Wijesundara

(1948: 144); LaCasse and Yamaguti (1950: 192-201); Bonne-Wepster

(1954: 111-3); Wang and Feng (1964: 37-41).

FEMALE. Small to medium sized (wing length 2.0-3.5 mm), usually

blackish, sometimes yellowish, reddish or tan-brown; palpus, proboscis and

legs without any pale markings. Head, Decumbent scales on dorsum of ver-

tex varying from narrow, linear to broad clavate or ovate, color varying from

pale whitish or yellowish, forming a distinct ocular line to predominantly dark,

‘brownish or blackish; erect scales numerous, usually entirely dark brown or

black, sometimes largely pale, yellow or golden; lateral patch of broad scales

at side of eye usually pale beige or bluish white, sometimes entirely dark;

frontal tuft of scales poorly developed or absent; frontal and dorsolateral ocu-

lar bristles very strong, usually dark, sometimes golden brown; subocular

bristles weak, dark or slightly pale. Palpus entirely dark scaled, 4-segmented,

0.20-0.25 of proboscis length; segment 4 longest, as long as or longer than

combined length of segments 1-3. Proboscis entirely dark scaled, slightly

longer than forefemur; labial basal setae weak, hairlike, usually 2-4 in num-

ber, sometimes 6. Antenna as long as proboscis; pedicel with or without

minute patch of tiny setae or scales on inner dorsal surface; flagellum com-

posed of 13 subequal flagellomeres; flagellomere 1 with several short setae in

middle; flagellomeres 2-13 with numerous short setae and each with a basal

whorl of 5-8 bristles, the latter 1.5-2.0 times as long as length of individual

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 9)

flagellomere. Cibarial Armature: Well developed; cibarial bar with 15-60

distinct teeth in concave row; length and thickness of teeth variable; cibarial

dome oval or hemispherical, with numerous minute tubercles or denticles.

Thorax. Mesonotal integument usually dark brown, sometimes yellowish,

reddish brown or almost black; mesonotal scales narrow, curved, sparse

or moderately dense, entirely dark brown or black; acrostichal bristles usually

absent except on extreme anterior promontory, sometimes present, weakly

developed, extending to anterior 0.5 of disc; dorsocentral, supraalar and

scutellar bristles well developed, very strong, dark brown or black. Pronotal

integument same color as mesonotum; anterior pronotum (apn) with short setae and

bristles only; posterior pronotum (ppv) entirely bare or with scattered short and

weak setae and narrow scales on anterior surface cephalad of 4-6 posterior

bristles. Pleuron slightly paler or concolorous with mesonotum, usually

without distinct scale patches on propleuron (pl), sternopleuron (St) and

mesepimeron (mep), sometimes with a few to many pale scales restricted to

upper corner and posterior border of stp and upper mep; 1 lower mep bristle

usually present, sometimes absent. Legs. Forecoxa with dark scale patch

and several strong, curved setae on anterior surface; midcoxa with a vertical

row of 4-6 dark bristles and a small pale scale patch on antero-lateral sur-

faces; hindcoxa with lateral pale scale patch and a posterior row of setae; an-

terior surface of forefemur and midfemur dark brown or black, ventral sur-

face pale; anterior surface of hindfemur usually with longitudinal white stripe

extending from base to near apex, sometimes absent or entirely dark; tibiae

and tarsi of all legs dark brown or bluish black; all claws small, equal and

simple; pulvilli well developed. Wing. Scales on all veins dark brown or

black; scales on C, Sc, Rand Ry dense and broad; plume scales on Ro, Re

moderately dense, narrow, linear or broad clavate; fringe scales on wing

margin dusky; cell Ro 1.5-2.0 times as long as its stem (R9,3); alula with

a few short marginal setae; upper and lower calypter fringed with several

long, hairlike setae. Halter, Peduncle pale and bare; capitellum covered

with dark scales. Abdomen. All terga usually dark brown or black, some-

times with basolateral pale spots or complete basal pale bands; sterna usually

pale beige yellowish, sometimes largely dark. Genitalia, Very similar in

all species. Segment VIII partially retracted into segment VII; sternum VIII

with several bristles in middle of posterior caudal margin; median caudal

margin lightly or moderately emarginate. Segment IX largely retracted into

segment VIII; tergum IX narrow, ribbonlike, with a lateral row of 3-4 weak

setae on each side. Cercus thumblike, short or moderately long, with several

setae and numerous spicules. Postgenital plate broad, oblong, not projecting

beyond apex of cercus; caudal margin broadly rounded or lightly emarginate at

middle; apical 0.5 with 4,5 setae in 2 parallel rows, rest of plate largely

spiculose. Posterior cowl very narrow, entirely spiculose. Sigma largely

membranous and densely spiculose. Insula with a tuft of 8 strong setae.

Vaginal sclerite in form of an inverted U, weakly sclerotized, lightly to mod-

erately pigmented.

MALE. In general similar to female, differing in smaller size and scantier

scales on the wing veins. Palpus. Long, slender, 5-segmented, usually ex-

ceeding proboscis by 0.5-1.0 of the length of segment 5, sometimes reduced to

0.5 of proboscis length; segment 1 with or without 2 pairs of small, short finger-

like processes on lateral and ventral surfaces; segment 3 with 1,2 ventral rows

of short, tiny setae, sometimes with numerous fine setae forming dense rows,

apical 0.25 with a weak ventrolateral tuft of dark bristles, number of bristles

varied; segments 4 and 5 weakly to strongly plumose, usually subequal, seg-

6 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

ment 5 sometimes considerably reduced in length. Proboscis. Slender or

moderately thick; false joint present at 0.25 from base or absent; dorsal surface

of labium with or without a double row of upright, sinuous setae extending from

0.25 of length to almost apex; lateral and ventral surfaces usually with a few

rows of short setae, widely spaced along whole length, sometimes with a few

long hairs or a dense group of several setae in middle; labial basal setae short,

hairlike or spinelike, 10-12 in number, in transverse row. Antenna, Pedicel

smooth, rounded, with or without distinct nipplelike spiculose prominence (or

mammiliform process) on inner dorsal surface; flagellum moderately to densely

long plumose; flagellomeres 1-11 subequal, 12,13 elongate; flagellomeres 5-10

or 5-9 with modified tufts of scales and setae, sometimes modified tufts re-

stricted to flagellomeres 7 and 8, rarely absent entirely; minor whorl of short

setae distad of normal whorls absent or not developed on flagellomeres 2-11.

Thorax, Setae on ppl usually more numerous than infemale. Legs, Claws

of fore- and midlegs enlarged, with basal denticles; claws of hindleg small and

Simple, as in females.

MALE GENITALIA. Segment Lx. Tergum narrow, lateral tergal lobe small,

poorly developed, with 3-5 weak or moderately strong setae; sternum broad,

more or less rectangular, usually with one or more transverse rows of strong

setae and scales towards caudal margin, sometimes absent. Basimere. Small

slender, roughly conical, with convex lateral tergal margin and concave tergo-

mesal margin, sometimes modified; scales absent; strong bristles confined to

lateral tergal surface; inner tergal surface usually with a few to several strong-

ly differentiated submarginal setae, in row or patch from near base to level of

subapical lobe, sometimes absent or with short and weak setae only; tergo-

mesal margin with one or a few rows of marginal setae. Subapical Lobe.

Prominent, usually not clearly divided into proximal and distal divisions,

setae on lobe densely packed, usually well differentiated; proximal part usually

with 3 stout, subequal rods @-c); distal part usually with a mesal group of 1

club-shaped leaflet (g5) and 4-6 flattened bladelike, narrow rodlike or fine

hairlike setae (d-f) and a lateral group of 1 leaflet (€;) and 1 strong basal seta

(h). Distimere, Slender, curved in middle, distal portion usually tapered into

a blunt recurved apex, latter sometimes enlarged or modified; subapical dorsal

surface usually without a crest of spicules; dorsal and ventral tiny setae

present, usually placed subapically, sometimes near middle of curvature; sub-

apical claw (or spiniform) slender, short, small and simple. Phallosome,

Aedeagus relatively simple, composed of 2 lateral plates which are connected

by a broad basal and a narrow submedian tergal bridge; apical portion of lateral

plate [ or dorsal lobe (DL)] simple, beaklike, with apex projecting tergad or

composed of a stout, denticulate or tuberculate external process (EP) anda

slender simple spinelike internal process (IP), sometimes represented by a

broad, heavily denticulate external process only; basal 0.5 of lateral plate

represented by a simple, rounded ventral (or sternal) lobe (VL) which is more

or less prominent in lateral aspect, sometimes reduced or indistinct; paramere

and parameral apodeme well developed, very similar in all species. Pyroc-

tiger. Apical crown of spicules small to large; number and texture of spicules

varied, paraproct narrow, well sclerotized, its apex usually with distinct

lateral and sternal lobes, placed adjacent to apical crown; basal sternal pro-

cess absent; basolateral sclerotization poorly or well developed, apparently

attached to base of paraproct; cercal sclerite poorly sclerotized; cercal setae

2-4 in number.

PUPA. Pigmentation of cephalothorax and abdomen varying from uniformly

pale, creamy or yellowish white to tan brown. Trumpet. Yellowish to dark

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 7

brown; usually very slender, long, more or less cylindrical (length 0. 5-0.8

mm; index 10-16), sometimes short, stubby, bell-shaped or funnel-shaped

(length 0.2-0.3 mm; index 3-4); pinna same diameter as meatus or slightly

flared towards apex, with or without slit extending to meatus. All setae

developed, relative position of setae essentially similar to other Culex sub-

genera. Metanotum. Seta 10-C usually double, sometimes single, rarely

triple or more branched; 11-C subequal to 10-C, single or double. Abdomen,

Seta 1-II usually brushlike or strongly dendritic, with several distal branches,

sometimes forked into 3-5 branches; setae 5-IV-VI strong, 1-2 times as long

as segment following; 9-IV 1-7 branched; 5-V 1-5 branched; 5-VI single or

double rarely triple; 9-VIII well removed from caudolateral angle. Paddle,

Usually oval shaped; sometimes hemispherical; usually pale whitish to almost

transparent; margin of outer part distinct, smooth or lightly spiculated in

basal 0. 50-0. 75, apical 0.25-0.50 rather indistinct; setae 1, 2-P usually

present, minute. :

LARVA. Generally similar to other Culex subgenera. Head. Usually

broader than long; ocular bulge usually prominent; collar dark: labrum well

developed; seta 1-C usually dark, spiniform, with or without accessory lateral

spicules or spines, sometimes pale, blunt-tipped or tapered into a fine point;

2-C absent; 14-C usually double, sometimes single or more branched; 16,

17-C usually present, sometimes absent. Antenna shorter than or nearly as

long as head; basal 0.5 of shaft usually with numerous strong spicules, some-

times absent or poorly developed; seta 1-A usually large, fan-shaped, multiple,

sometimes strongly reduced, 3,4 branched; 2, 3-A usually long, bristlelike,

situated subapically, sometimes very weak, situated apically. Mouth brush

filamentous, sometimes more or less flattened. Mental plate with 6-8 strong

lateral teeth on each size of median tooth. Thorax. Spiculation usually lightly

to moderately developed or completely absent, sometimes strongly developed

and extremely dense, setae 1,2-P strong, long, subequal and single; 3-P

reduced to 0.25-0.50 of length of 1,2-P, usually single or double, sometimes

more branched; 4-P strong, subequal to or slightly shorter than 1,2-P, 1-4

branched; 7-P strong, 1-3 branched; 8-P usually subequal to 7-P, sometimes

greatly reduced, 1-3 branched; 14-P single or double. Seta 1-M shorter or

as long as 3-M, single. Abdomen. Spiculation as in thorax; seta 7-I usually

double, sometimes single or 3-5 branched. Comb scales usually numerous

forming a broad oval or triangular patch. Saddle complete, concolorous with

siphon, caudolateral margin lightly to strongly spiculate, sometimes practi-

cally smooth or with fringe of numerous spines; setae 4-X (ventral brush)

usually with 6 pairs of branched setae (total 12), sometimes 5, all inserted

within grid; anal gills as long as or longer than saddle. Siphon. Usually very

long, slender or very thin, distally tapering, 4-6 times as long as saddle,

sometimes reduced to 0.5-2.0 of saddle length; pigmentation varied from pale

yellow to dark brown; pecten well developed; siphonal tufts weak, widely

spaced or strong, closely spaced, 3-8 pairs (total 6-16), all of which are

situated subventrally distad of pecten; median caudal filament of spiracular

apparatus present or absent.

SYSTEMATICS. The subgenus Lophoceraomyia is characterized in the

male by the presence of modified scales and setae forming conspicuous tufts

at the bases of the flagellar whorls of the antenna. These peculiar tufts are

developed on at least flagellomeres 7-8 or in the vast majority of species on

flagellomeres 5-10 or 5-9. Currently, 2 members of the subgenus are known

including: impostor n. sp from Sabah, Malaysia and sumatranus Brug 1931

from Sumatra, Indonesia, in which these characters are absent. Another

8 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

unusual feature of Lophoceraomyia is the presence of acrostichal bristles on

the mesonotal disc in certain species: paivojin. sp. from Thailand and

kusaiensis Bohart | 1956 (1957)] from Micronesia. Accordingly, I have given

above a more detailed description of the subgenus than in the past by including

the multitude of characters of the adults and the immatures.

All stages of Lophocevaomyia can be readily recognized and distinguished

from other subgenera of Culex in the Oriental region (including Acalleomyia,

Culiciomyia, Eumelanomyia, Culex, Lutzia and Barraudius) by the following

combination of characters: adults of both sexes by (1) relatively small size

(wing length not exceeding 3.5 mm); (2) general coloration usually blackish or

dark brownish, without pale markings on palpus, proboscis and legs; (3)

scales on mesonotum entirely dark, moderately dense or sparse, producing

a rough appearance; (4) acrostichal bristles absent on mesonotal disc (except

pairoji); (5) pleural scale patch largely absent (except sumatranus and curti-

palpis); (6) 1 lower mep bristle usually present; (7) wing scales moderately

dense or sparse and (8) abdominal terga usually completely dark, or some-

times with basolateral pale spots or basal transverse pale bands; male by (1)

antennal pedicel sometimes with spiculose prominence on dorsal inner surface;

(2) antennal flagellomeres 5-10, 5-9 or 7-8 with modified tufts of scales and

setae (except impostor and sumatranus); (3) minor whorls of short setae distad

of normal large whorls of long setae on flagellomeres 2-11 absent; (4) palpus

very slender, usually as long as or longer than proboscis, sometimes reduced

to 0.5 of proboscis; (5) palpal segment 1 sometimes with small, fingerlike

processes and (6) proboscis sometimes with false joint at 0.25 from base;

male genitalia by (1) basimere without any scales; (2) distimere usually with-

out distinct crest of spicules on dorsal subapical surface; (3) lateral plate of

phallosome relatively simple, sometimes with small distinct denticles or

tubercles and (4) basal sternal process of proctiger absent; pupa by (1) trumpet

usually long, slender and cylindrical (except in certain pitcher plant species);

(2) seta 10-C single or double, or not exceeding 3 branches and (3) seta 9-VIII

well removed from caudolateral angle; larva by (1) setae 16, 17-C usually

present; (2) seta 14-C usually bifid; (3) seta 3-P short, 0.25-0.5 of 1, 2-P;

(4) seta 4-P strong, slightly shorter or subequal to 1,2-P; (5) 7-P double or

triple; (6) 8-P usually subequal to 7-P, sometimes strongly reduced, single or

double; (7) 14-P single or double; (8) siphon very slender and long (except for

certain pitcher plant species); (9) all siphonal tufts inserted subventrally and

(10) ventral brush of saddle (or 4-X) with 6 pairs of branched setae.

Practically all Lophoceraomyia species have been recognized on the basis

of the differences in the male antenna, palpus, proboscis and genitalia. The

females of most species are difficult to identify except by correlation with the

males through associated larval and/or pupal skins from individual field rear-

ings. The female cibarial armature may be significant as a specific or group

character but the female genitalia are of little or no taxonomic value. The

larvae and pupae which have been studied so far also provide several important

taxonomic characters for separating as well as grouping species at various

levels. They are, in several cases, of equal significance to the male charac-

ters.

Prior to this study, approximately 55 Lophoceraomyia species were con-

sidered as valid taxa for the Oriental region (Stone, Knight and Starcke 1959;

Stone 1967; 1970). In this study, nearly all nominal species previously des-

cribed by Borel (1930), Barraud (1934), Baisas (1935; 1974), Colless (1965,

1967), Delfinado (1966), Bram and Rattanarithikul (1967), Bram (1967a) and

other authors are treated. The only nominal forms not included here are:

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 9

bernardi (Borel 1926)from Vietnam, mindanaoensis Baisas 1935 from the Philip-

pines, szemaonensis Wang and Feng 1964 and harpagophallus Wang and Feng

1964 from Yunnan, China and vaghavanii Rahman, Choudhury and Kalra 1969

from India. The types of bernardi and mindanaoensis are apparently lost and

none of the topotypic material has been found to agree with the published des-

criptions. It is possible that they were erroneously described and both are

perhaps better reduced to nomina dubia for the present. The types and topo-

typic specimens of szemaonensis,harpagophallus and vaghavanii were not avail-

able for this study, but from the descriptions and the figures of the male anten-

nae and genitalia by their authors, it appears probable that szemaonensis may

be conspecific with ganapathi Colless 1965, harpagophallus with minor (Leicester

1908) or bicornutus (Theobald 1910) and vaghavanii with flavicornis Barraud 1924.

In this study, 58 species of Lophoceraomyia are recognized from the

Oriental region. Of these, 49 are revalidated and 9 are considered as new.

In treating the previous nominal species, the following taxonomic changes have

been made: (1) synonymizing pachecoi Baisas 1935 with quadripalpis (Edwards

1914a); (2) resurrecting bicornutus (Theobald 1910) from the synonymy with minor

Leicester 1908 proposed by Bram (1967a); (3) synonymizing hui Lien 1968 with

spiculosus Bram and Rattanarithikul 1967; (4) synonymizing fuscosiphonis Bram

and Rattanarithikul 1967 with demissus Colless 1965; (5) synonymizing plan-

taginis Barraud 1924 with minor (Leicester 1908); and (6) relegating barkerii

(Theobald 1907), bernardi (Borel 1926)and mindanaoensis Baisas 1935 to nomina

dubia. The new species recognized are: alorensis from Alor, Indonesia;

paraculeatus from Sabah, Malaysia and the Philippines; aestivus from Sabah,

Malaysia; gvacicornis from Peninsular Malaysia (Malaya); pairojifrom Thai-

land; impostor from North Borneo, Malaysia; wardi and lasiopalpis from Sri

Lanka and hirtipalpis from northern Thailand.

CLASSIFICATION. The classification of the Oriental Lophoceraomyia

adopted here is modified from Edwards (1932; in Barraud 1934) and Colless

(1965). The 58 species treated here apparently fall into 3 natural groups:

Fraudatrix, Mammilifer and Wilfredi on the basis of the male phallosome,

antennal pedicel, palpus and proboscis as indicated in the key to groups. Of

these, 23 species belong to the Fraudatrix Group, 32 to the Mammilifer Group

and 3 to Wilfredi Group. The Fraudatrix Group is subdivided into 2 subgroups:

Minutissimus and Fraudatrix. The latter is further split into 7 complexes:

seniori, cinctellus, rubithoracis, inculus, quadripalpis, variatus and alphus,

The Mammilifer Group is divided into 2 subgroups: Mammilifer and Brevipalpus

(or subgroups B, and Bo of Colless 1965). The mammilifer Subgroup includes

all species previously treated by Colless (loc. cit.) except wilfredi, It is split

into 8 complexes: impostor, traubi, mammilifer, ganapathi, minor, peytoni,

pholeter and flavicornis, The Brevipalpus Subgroup is divided into 5 com-

plexes: navalis, hewitti, jenseni, brevipalpus and eurtipalpis, For more de-

tailed discussions on the characterization and the alignment of species accord-

ing to this scheme, see the treatment of the groups and subgroups.

AFFINITIES. Among the subgenera of Culex, Lophoceraomyia apparently

exhibits the strongest affinity with Eumelanomyia as pointed out by Sirivana-

karn (1971, 1972). This relationship is evident in the following features: (1)

adults - scales on mesonotum dark, narrow, sparse or moderately dense;

pleuron without distinct scale patches; female cibarial armature with numer-

ous teeth; male proboscis without median tufts of long hairs on ventral surface;

(2) male genitalia - development of setae and leaflets of the subapical lobe;

distimere without crest of strong spicules on subapical dorsal surface; shape

and relatively simple structure of phallosome; absence of basal sternal process

10 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

of the proctiger; (3) pupae - long, slender trumpet; seta 10-C single or

double, rarely more branched; (4) larvae - stout, spiniform seta 1-C, pres-

ence of setae 16,17-C; seta 3-P weaker and shorter than 1, 2-P; seta 14-P

‘single or double; siphon slender and relatively long (except for certain pitcher

plant species); ventral brush (4-X) of saddle with 6 pairs of branched setae.

Based on the similarity in the adults and male genitalic characters, Lopho-

ceraomyia is apparently closest to the rubinotus-vima group of Eumelanomyta

and it seems plausible, as suggested earlier (Sirivanakarn 1971, 1972) that it

was probably a derivative of that group in the latter subgenus. Ona broad

comparative basis, Lophoceraomyia also exhibits some similarity to Culicio-

myia in the simple male phallosome and general external features of the adults.

but differs from all known members of the latter subgenus very strikingly in

several other features of the male, male genitalia and immature stages. The

affinity between Lophoceraomyia and Culiciomyia as suggested by Colless

(1965) does not seem to be tenable on the basis of present knowledge.

EVOLUTION. Only a broad generalization on the evolution of Lophocerao-

myia is discussed here. Although this topic has been discussed in general by

Colless (1965: 265), my knowledge of the group in the Papuan subregion (Siri-

vanakarn 1968) together with the current study of the Oriental fauna and others

has warranted further speculation on the phylogeny and other aspects of the

evolution in the subgenus. We still lack knowledge of the subgenus in some

critical areas, especially among the islands of Indonesia, including Borneo

and other islands to the west of New Guinea and it is quite conceivable that

further study of these areas would fill numerous gaps in providing more con-

crete evidence concerning the place of origin and the phylogeny of some major

group taxa.

On the basis of morphology, breeding habitats and zoogeography, it appears

most probable as suggested by Colless (1965) that the most primitive forms of

Lophoceraomyia are representatives of the Mammilifer Group (or Group B)

rather than the more widespread Fraudatrix Group (or Group A). These primi-

tive forms probably originated in the Indomalayan area or Southeast Asia where

the subgenus Eumelanomyia is predominant. The males of these forms may

have been primitively Eumelanomyia-like in the absence of special modifications

of the antennal pedicel and flagellum, as exhibited by some Lophoceraomyia

species, such as impostor from Sabah, North Borneo, sumatvanus Brug from

Sumatra; bolii Sirivanakarn and digoelensis Brug from New Guinea (Sirivana-

karn 1968: 100-5). These presumptive Eumelanomyia-like forms were probably

ground pool breeders and were presumably widely scattered among the areas

between Borneo and the Moluccas and subsequently gave rise to various phyletic

lines of the Mammilifer Group, which were further differentiated into several

forms through specialization in breeding habitats and geographic isolation.

Concurrently or subsequently some of these primitive Mammilifer forms gave

rise to the members of the Fraudatrix group which were widely dispersed

northwest into Asia and east and southeast into New Guinea, northern Australia,

islands of the Central and Western Pacific (Micronesia) and the South Pacific.

As opposed to the view of Colless that New Guinea may be the ancestral home

of the Fraudatrix Group, I however believe that the several islands of Indo-

nesia to the west of Wallace's and Weber's lines are the more likely places

where the ancestors of this group originated and subsequently radiated in all

directions. The distribution patterns as shown by the contemporaneous faunas

appear to correspond to Dispersal 5 of Belkin (1962) and the origin and the

course of evolution of the group probably took place during the Tertiary period

(75 million years ago) according to the view tentatively advanced by Colless

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 11

(1965).

As can be discerned, the most important factors in determining the course

of speciation of Lophoceraomyia are primarily the types of breeding habitat

and geographic isolation. In addition, other factors such as population size,

flight range and adult behavior are also important in stabilizing population

structure as well as in preventing newly adapted forms from cross breeding.

These intrinsic factors together with the types of breeding habitats apparently

play a major role in the proliferation of the numerous species in the Mammili-

fey Group. Because of the restriction in breeding sites, such as bamboos,

tree holes, pitcher plants and other natural containers associated with the

types of tropical forest formation, the majority of the species in this group

are largely endemic or indigeneous and are not susceptible to introduction into

a new zone through geographic isolation and dispersal agencies. This may

explain why the group is almost exclusively Indomalayan. The occurrence of

several closely related forms within this area strongly indicates that the evolu-

tion in the Mammilifer Group largely took place through sympatric speciation.

The same generality can be accounted for the Fraudatrix Group. However, as

it is composed largely of breeders in ground pools, there is obviously a

tendency for these forms to be capable of invading a new zone and become

widely established through geographic isolation and various dispersal mechan-

isms.

BIONOMICS. The majority of Lophoceraomyia species are restricted to

humid tropical forests and only a few members occur in open cultivated lands

such as rice fields and plantations. In the Oriental region, they are found

breeding in a variety of habitats, ranging from general ground pools (most

species in the Fraudatrix Group) to rock holes, tree holes, bamboo stumps,

leaf axils of pandanus and pitcher plants (most species in the Mammilifer

Group). Elsewhere, in the South Pacific (Belkin 1962), some members of the

Fraudatrix Group were also reported from leaf axils of aroids and pandanus

and artificial containers. In the Papuan subregion (Sirivanakarn 1968),they

have also been found breeding in palm bracts, sago stumps and sago leaf axils.

The adults of most species have been obtained from rearing the larvae or pupae. By

sweeping with nets, several were also caught while they were flying or resting on

leaves of plant seedlings or shrubs. They were also caught in Malaise traps and oc-

casionally in lighttraps. In Malaya, Reid (1941) reported that anunknown Lopho-

ceraomyia species was observed to feed on the tree snake(Passerita prasina) and

Colless (1965) noted that a number of species are known to attack man under forest

cover but are rarely taken with mammalian baits in the open. Ina study using

the precipitin test (Colless 1959), the females were normally found to obtain

blood meals from wild birds. Lophocevaomyia were reported to feed on frogs

and occasionally on man in Australia (Marks 1960). In Japan, females of C.,

infjantulus were reported to feed readily on frogs, snakes, lizards, turtles,

chicks and mice in the laboratory (Miyagi 1972, 1973). In New Guinea, several

wild-caught females were found to be engorged with a reddish or blackish sub-

stance, which suggested that they might feed on wild birds or other vertebrates

in nature (Sirivanakarn 1968).

MEDICAL IMPORTANCE. Little is known about the medical importance

of Lophoceraomyia. In Malaysia, adults of some unidentified species have

been reported to be naturally infected with 4 different arboviruses: Bakau

(MM-2325), Ketapang (MM-2549) and Bebaru (MM-2354) and Lahore (a strain

of Bakau No. 114)(Anonymous 1960, Berge 1970). It is probable that certain

Lophoceraomyia species play an important role as a natural reservoir of

arboviruses.

12 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

DISTRIBUTION. Lophoceraomyia, as far as known, is confined almost

exclusively to the tropics and subtropics of the Oriental and Australasian

regions, the islands of the western and central Pacific Ocean (Micronesia)

and the South Pacific with a small extension into Japan, Korea and northern

China in the Palearctic. Its westernmost limit appears to be in India and

Pakistan and it has never been reported further to the west in the Middle East

and the Ethiopian region. Of the 3 major groups, the Fvaudatrix Group is the

most widespread and has been recorded from all areas within the reported

range of the subgenus. The Mammilifer Group is almost exclusively Oriental

with only a few members occuring to the north in southern China, Taiwan and

the Ryukyus and to the east as far as Papua New Guinea. The Wilfredi Group

is exclusively Oriental, where it has been recorded only from Peninsular

Malaysia, Thailand and southern China.

KEYS TO GROUPS OF ORIENTAL LOPHOCERA OMYIA

ADULTS

L, Female: Decumbent scales on dorsum of vertex largely broad;

scales on veins Roy and Rg of wing usually broad, clavate; abdo-

minal terga ee with B onlatenal pale spots or sometimés

basal pale bands; Male: Basal segment 1 of palpus with 1 or 2

pairs of spiculose fingerlike processes; modified tufts of scales

and setae present on flagellomeres 7-8 or 5-10.

FRA UDATRIX GROUP

Female: Decumbent scales on dorsum of vertex largely narrow,

linear; scales on veins Ro and Rg of wing narrow, linear; abdominal

terga completely dark or without pale markings as above; Male:

Palpus without basal fingerlike processes; modified tufts of

scales and setae present on flagellomeres 5-9 or 5-10 or com-

pletely absent. ..eo ee Oe ee 2

2(1). Male: Antennal pedicel with spiculose prominence on inner dorsal

surface; modified tufts of scales and setae usually present on

flagellomeres 9-8 or 5-9; anterior surface of forefemur without

a dense tuft of setae in apical half. .... MAMMILIFER GROUP

Male: Antennal pedicel without spiculose prominence on inner

dorsal surface; modified tufts of scales and setae present on

flagellomeres 5-10; anterior surface of forefemur with a dense

tuft of several strong setae in apical half... WILFREDI GROUP

MALE GENITALIA

Lb. Dorsal lobe (DL) of lateral plate of phallosome (aedeagus) repre-

sented by a slender, simple, apical beaklike process, projecting

tergally; ventral lobe (VL) prominent and very distinct in lateral

aspects; proctiger crown relatively small. . FRAUDATRIX GROUP

Dorsal lobe (DL) of lateral plate of phallosome represented by a

large denticulate or tuberculate external process (EP) and with

or without a simple, spinelike internal process (IP); ventral lobe

(VL) reduced and not as distinct as above; proctiger crown large. 2

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 13

2(1). Lateral plate of phallosome bowl-shaped, with a partially denticu-

late or tuberculate external process (EP) and a simple, spine-

like internal process (PP) eee ee MAMMILIFER GROUP

Lateral plate of phallosome compact, barrel-shaped, with a

broad, heavily denticulate external process (EP) only.

WILFREDI GROUP

PUPAE

dy Trumpet short, bell-shaped or funnel-shaped, 0.2-0.4 mm in

length; occurring in pitcher plants.

MAMMILIFER GROUP (in part)

Trumpet long, cylindrical, 0.5-0.8 mm in length; occurring in

ground pools, rock pools, tree holes, bamboos and leaf axils. . 2

2(1). Pinna of trumpet with slit extending into meatus........... 3

Pinna of trumpet without slit extending into meatus.

MAMMILIFER GROUP (in part)

3(2). Seta 5-IV double (except mammilifer, wardi and bengalensis).

MAMMILIFER GROUP (in part)

Seta. S41Vi @-'Ppnanthed., eee, hoe A Se eee ae a 4

4(3)s5 «Seta 8<C single ordouble,... 1. eee. FRA UDA TRIX GROUP

Sela 6+C 4,0 pravcheds, 620 ee ee .. « -WILPREDI GROUP

LARVAE

fe Setae 2,3-A placed apically; setae 16,17-C absent; breed in pitcher

planta ese ee ee MAMMILIFER GROUP (in part)

Setae 2,3-A placed subapically; setae 16, 17-C present; breed in

ground pools, rock pools, tree holes, bamboos and leaf axils. .. 2

2(1). Seta 7-P triple (except veidi); seta 1-M 0. 25-0. 50 of the length

of seta 3-M; breed in general ground pools. . FRAUDATRIX GROUP

Seta 7-P double; seta 1-M usually as long as seta 3-M; breed in

rock pools, tree holes, bamboos, leaf axils or sometimes

Sroune HOOISEE tre. Pie ae ea, Se 3

3(2). Seta 14-P double; usually breed in rock pools, tree holes, bamboos

or sometimes leaf axils (except mammilifer, bengalensis and

pholeteyvy 20h 3 ee ee A eS MAMMILIFER GROUP

Seta 14-P single, breed in ground pools only... WILFREDI GROUP

FRAUDATRIX GROUP

FEMALE. As described for the subgenus; coloration varying from yellow,

orange to dark brown, sometimes darker to almost black. Head, Decumbent

scales of vertex relatively broad, clavate or ovate, occupying an extensive

area in center, color of scales predominantly dark or sometimes with pale

14 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

ones forming a narrow ocular line; narrow decumbent scales rather few,

largely restricted to dorsal midline of vertex and occiput; erect scales numer-

ous, entirely dark brown or black; lateral patch of broad appressed scales

partially pale to completely dark. Proboscis usually with 2 hairlike labial

basal setae, sometimes 4-6, as long as or slightly longer than palpus. Cibari-

al Armature, Cibarial bar with 40-60 teeth in concave row, all teeth subequal

in length and size and apically blunt. Thorax, Integument of mesonotum

usually dark brown, sometimes pale yellow or orange; all mesonotal scales

dark brown, sometimes few pale scales present, restricted to marginal areas

on anterior promontory, humerus and scutal angle; pleural scale patches

practically absent, sometimes with a few scattered scales on upper corner

and posterior border of stp; ppl with 2,3 dark bristles and a few other pale

weak setae. Wing. Scales on veins Ro, R. usually broad, clavate, sometimes

narrow, linear. Abdomen. All terga usually entirely dark, sometimes with

basal pale bands or basolateral pale spots.

MALE. Generally similar to females except for scantier scales on wing

veins and presence of more numerous ppl bristles. Palpus, Longer than

proboscis by 0.5-1.0 of the length of segment 5; segment 1 usually with 1 or

2 pairs of conspicuous fingerlike processes; segment 3 with 1,2 rows of short,

tiny setae on ventral surface; segments 4 and 5 upturned, moderately to

strongly plumose. Proboscis. Labial basal setae stout, spiniform, 8-12, in

irregular transverse row on ventral surface; false joint present, usually marked

by flexion at 0.25 from base, sometimes absent; apical 0.5 or more usually

with a double row of sinuous setae on dorsal surface, sometimes absent.

Antenna. Pedicel smooth, rounded, without nipplelike spiculose prominence

on inner dorsal surface; flagellar whorls densely long plumose; modified tufts

of scales and setae usually present on flagellomeres 5-10, sometimes 6-10 or

restricted to flagellomeres 7-8; size and shape of modified tufts varied.

MALE GENITALIA. Segment IX. Tergal lobe with 3-4 weak setae; sternum

usually entirely bare, sometimes with a few setae toward caudal margin. Basi-

mere. Small, conical; bristles and setae on outer lateral surface rather

sparse; inner tergal surface usually with differentiated submarginal setae, in a

single row parallel to tergomesal margin, number and length varied. Subapical

Lobe. Not clearly divided into proximal and distal divisions; proximal part

with 3 stout rods @-c) and usually 1 hairlike seta ventrad of seta a; setae of

distal parts varied, usually with a mesal group of 4-6 bladelike setae (a-f) and

1 club-shaped leaflet (>) and a lateral group of 1 broad leaflet (21) and 1 strong

basal seta (1); sometimes leaflets (£1, g) not developed or absent. Distimere.

Normal; subapical portion very weakly serrated, without distinct crest of fine

spicules extended to middle of curvature on dorsal surface; subapical claw,

short, slender, distally dilate. Phallosome. Dorsal lobe of lateral plate in

form of a beaklike process projecting tergad, which is simple or entirely bare

(except for infantulus which is strongly imbricate with reticules); ventral lobe

broad, prominent, with upper sternal margin at about same level of apical

margin of dorsal beaklike process. Proctiger. Crown small, with relatively

small number of spicules; apical and sternal lobes of paraproct poorly or well

developed; cercal setae 2-3 in number.

PUPA. Cephalothorax and abdomen generally whitish or creamy white with

indefinite darkened areas. Trumpet. Slender, relatively long and more or less

cylindrical, length varies from 0.5-0.8 mm, index 10-18; pinna usually with

slit extending to meatus. Cephalothorax, Setae 8,9-C usually double, some-

times single or triple. Metanotum, Seta 11-C double. Abdomen. Seta 5-IV

usually 4-6 branched (3-8); 5-V 2-5 branched; 5-VI usually double (1-3); 5-V, VI

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 15

shorter or as long as individual segment following; 6-III-VI 2-6 branched;

9-VII subequal in length to 9-VIII. Paddle, Usually entirely pale; midrib

usually weak, pale, sometimes dark.

LARVA. Head, Integument yellowish white; seta 1-C stout, dark, simple

spiniform; 4-C usually single or forked, short, usually 0.5 of distance between

bases of the pair, sometimes shorter or longer; 5,6-C strong, subequal; 9-C

usually double, sometimes single or triple; 6-C always double. Antenna nearly

as long as head; with numerous strong spicules on outer dorsal surface in basal

0.50-0. 75 or more; seta 1-A large, fan-shaped, multibranched, placed at 0.75

of length from base; 2,3-A bristlelike and dark, situated subapically. Thorax,

Integument with or without conspicuous patch of spicules; seta 3-P usually

Single or double, sometimes multibranched; 4-P double or sometimes single;

7-P triple; 8-P subequal to 7-P, double; 14-P usually single, sometimes

double; 1-M usually very short, inconspicuous, much shorter than 3-M, rarely

as long as the latter. Abdomen. Spicules absent; live or whole mount speci-

mens usually with light and dark greenish bands on segments I-VI; segments

VII-VIII pale; setae 6-I, II pale, usually triple; 7-I pale, usually double, rarely

Single; 6-III-VI subequally long, 3-5 branched. Comb scales numerous, in

broad oval patch, all scales small, subequal, apically rounded, with even

fringe of fine spicules; seta 2-VIII usually double, sometimes single. Saddle

complete; posterior caudal margin lightly spiculate; anal gills slender, usually

longer than saddle. Szphon. Very slender, long, distally gradually tapered;

1.0-2.0 mm; index 8-12; siphon/saddle ratio usually 4,5; pecten 8-14; subven-

tral tufts 4 pairs (or total 8), all relatively weak, short, subequal, as long as

or slightly longer than siphonal width at points of attachment; median caudal fila-

ment of spiracular apparatus well developed; seta 2-S weak, pale and short;

ventral and dorsal valves of spiracle small.

DISCUSSION. The Fraudatrix Group in the broad sense occupies the entire

reported range of the subgenus and is particularly common in the Oriental

region (Southeast Asia), the Papuan part of the Australasian region and the

South Pacific. Elsewhere, it has been reported from Australia [ Edwards 1924;

Dobrotworsky 1957 (1958)]; the western and central Pacific in Micronesia

[| Bohart 1956 (1957)] and from Japan in the eastern Palearctic (LaCasse and

Yamaguti 1950).

Edwards (1932: 196-8) in his original scheme restricted the Fraudatrix

Group to his group B (Lophoceratomyia), but later (in Barraud 1934: 360-72)

he expanded it to also include group A (Minutissimus), The latter scheme was

subsequently adopted by Colless (1965: 264) and is also followed here. The

only modifications which I have made are to recognize 2 subgroups within this

group: Minutissimus and Fraudatrix and to subdivide the second subgroup into

7 complexes. The 23 Oriental species are segregated according to the present

scheme as follows: (1) Minutissimus Subgroup includes: minutissimus Theo-

bald, alorensis n. sp. and infantulus Edwards; (2) Fraudatrix Subgroup repre-

sented by: (a) seniori complex with seniori Barraud, (b) cinctellus complex

with cinctellus Edwards and fulleri (Ludlow), (c) rubithoracis complex with

rubithoracis (Leicester), niger (Leicester) and gibbulus Delfinado, (d) inculus

complex with inculus Colless, (3) quadripalpis complex with quadripalpis

(Edwards), aculeatus Colless, paraculeatus n. sp., aestivusn. sp. and reidi

Colless, (f) variatus complex with variatus (Leicester), josephinae Baisas,

cubitatus Colless, gracicornis n. sp., whartoni Colless, macdonaldi Colless

and pairojin. sp. and (g) alphus complex with alphus Colless. For the accounts

on the relationships among the various complexes and species, see the discus-

sion of the subgroups.

2(1).

3(1).

4(3).

5(4).

6(5).

7(6).

8(3).

9(8).

10(9).

11(10).

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

KEYS TO SUBGROUPS, COMPLEXES AND SPECIES

FEMALES!

Abdominal terga with basal transverse pale bands. ....... 2

Abdominal terga entirely dark or with small basolateral pale

Spots (praudcive, Suberoup, in part). 2. sec. a 3

Proboscis with 4-6 labial basal setae; plume scales on wing

veins Ro, Re narrow, linear (Minutissimus Subgroup).

(minutissimus, alorensis and infantulus inseparable, use male genitalia)

Proboscis with 2 labial basal setae; plume scales on wing veins

Rog, Rg broad clavate or ovate (Fraudatrix Subgroup, in part).

cinctellus

Small species. wine leneth 2.4-2. 7mm. 2 6 io. 4

Medium or large sized species, wing length 2.8-4.0 mm. .... 8

Integument of thorax yellowish, orange or golden; lower mep

Bristle Ansel, sie ie ea ee rubithoracis

Integument of thorax brownish to blackish; lower mep bristle

DECGENE G3 ee a a ee 5)

Lower posterior mep with a minute patch of tiny semierect scale-

WRC SELAG Che a SS ae niger

Lower posterior mep without patch of setae as above. ..... 6

Labial basal setae of proboscis strong, 6 in number. .... alphus

Labial basal setae weak, .2 in-number... 20. <0s 0.0. 8 eee 7

Abdominal terga with basolateral pale spots. ........ inculus

Abdominal terga without basolateral pale spots. ........ reidi

Plume scales on wing veins Rg, Rg relatively narrow, linear or

clavate (quadripalpis COMmplom). 6 teh. Vee a ee 9

Plume scales on wing veins Ro, Rg relatively broad, ovate. ... 10

Wing length usually 3.4-3.7 mm (3. 0-4.0 mm); cibarial armature

AVA OU EECUNs oie. ds oe Ae es Rae ee I ae quadripalpis

Wing length usually less than 3.0 mm (2.7-3.0 mm); cibarial

armature with 35-40: teeth: £5 2.20. ee aculeatus

ACrostichal bristles present... fo a ee ee patroji

merostiehal bristles absent... of Ses Was a 11

Inseparable, see key to males and male genitalia.

fulleri, variatus, josephinae, cubitatus, whartoni and macdonaldi

seniori, gibbulus, pavaculeatus, aestivus and gracicornis are unknown.

4(1).

0(4).

6(5).

7(6).

8(7).

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 17

MALES AND MALE GENITALIA

Antennal flagellomeres 7 and 8 with inconspicuous mesal tufts of

modified, stout spinelike or bristlelike setae; subapical lobe of

basimere of genitalia without any leaflet (Minutissimus

Sub@rOup)sr.: 6 ce 5 swine ce Ge cee Je en ee 2

Antennal flagellomeres 5-10 or sometimes 6-10 with distinct

lateral, ventral and mesal tufts of modified scales and setae;

subapical lobe of basimere of genitalia with at least 1 or

usually 2 leaflets (Fraudatrix Subgroup). ........... ft

Dorsal beaklike process of phallosome short, stout, and simple. . 3

Dorsal beaklike process of phallosome long, slender and strongly

imbricate with reticules. -4 ee we 26.453 46 3 ee tnfantulus

Dorsal beaklike process of phallosome with a posterior recurved

spine sternad; proctiger crown with 14-17 dark spinelike ©

SPICULES J ieas -2Ci Nee a e S alorensis

Dorsal beaklike process of phallosome without posterior

recurved spine; proctiger crown with 4-5 pale toothlike

SPICHIOSs 45 Gs a eee a ee ee minutis StmMuUus

Modified tufts of scales and setae poorly developed and restricted

to antennal flagellomeres 6-10 only; apical half of proboscis

without dorsal sinuous setae (seniori complex). ..... seniovi

Modified tufts of scales and setae well developed on antennal

flagellomeres 5-10; apical half of proboscis usually with

dorsal: simuous Selaess: <3". 46. 2 ee ee ee 9)

Antennal flagellomere 5 with a small tuft of narrow, hairlike

setae or broad scales which are shorter or slightly longer

than combined length of next 3 flagellomeres.......... 6

Antennal flagellomere 5 with a large fan-shaped tuft of several

broad, blunt tipped scales which are as long as or longer

than combined length of next 4 flagellomeres. ........ 17

Basimere of genitalia stout, swollen or convex on tergomesal

margin and with a dense group of several strong submarginal

Setae in PALCRe 6 eae a ae 7

Basimere of genitalia slender, with concave tergomesal margin

and relatively few strong submarginal setae in a single regular

Dorsal process of lateral plate of phallosome anvil-shaped,

with numerous toothlike ridges (inculus complex). .... inculus

Dorsal process of lateral plate of phallosome beaklike and

simple (ciictelins Complex). oe ole gee ae 8

Submarginal setae of basimere relatively weak, subequal and more

or less widespread; leaflets g1, go of subapical lobe moderately

DrOAG: 54 Ge eek re a ee a: -einetellus

9(6).

10(9).

11(10).

1219):

13(12).

14(13).

15(14).

16(5).

17(16).

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Submarginal setae of basimere strong and densely packed, 4-9

basal ones very strong, remaining weaker and subequal;

leaflets 21, go of subapical lobe very broad. ....... fulleri

Modified tuft of antennal flagellomere 5 rather inconspicuous,

consisting of narrow, flattened, fine tipped setae only

(PUORNOTE CIS COMDIEN) « Gch ah RT ee A aS 10

Modified tuft of antennal flagellomere 5 very conspicuous,

consisting of some broad, dark scales and fine tipped,

flattened setae (quadripalpis complex). ...........-. 12

Proboscis without dorsal sinuous setae; paraproct of proctiger

with: @ lone, apical thumblike lope: ioe. ej ee eS niger

Proboscis with dorsal sinuous setae; paraproct of proctiger with

a short, pointed or rounded apical lobes: 5. So ae 11

Distal division of subapical lobe elongate, leaflet g, absent;

basimere with 1 submarginal seta. ........... gibbulus

Distal division of subapical lobe not elongate, ieane: £1 present;

basimere with a row of 3 submarginal setae... . rubithoracts

Basimere with a prominent row of 7-8 long submarginal

SET ESRI OP Crem PR ir 7 ace Ene ear ra ae eee

Basimere with a weak row of 2-4 short submarginal setae.

Submarginal setae strong, flattened, usually 4 in number; one

of rodlike a-c of subapical lobe with characteristically

expaiiced apex (hier hS)) Var Sells oh See. rveidi

Submarginal setae weak, fine, hairlike, 1-3 in number; none of

rodlike a-c of subapical lobe with modified apex as above. .. 14

Basimere with 3 strong submarginal setae. ........... 15

Basimere with 1 weak submarginal setae. ......... aestivus

Dorsal beaklike process of lateral plate of phallosome slender,

thin and remarkably long; antennal flagellomere 7 with well

developed comb-shaped tuft of modified setae. .... aculeatus

Dorsal beaklike process of lateral plate of phallosome stout and

relatively short; antennal flagellomere 7 with poorly developed

comb-shaped tuft of modified setae. ........ paraculeatus

Small, wing length at most 2.6 mm; modified tuft of antennal

flagellomere 5 with scales in ventral group strongly expanded or

swollen toward apices (alphus complex). ......... alphus

Medium or large, wing length more than 2.8 mm or usually 3 mm;

modified tuft of antennal flagellomere 5 without scales in ven-

tral group modified as above (variatus complex). ...... 17

Modified tuft of fused setae of antennal flagellomere 8 with charac-

teristic kink toward apical J-hook; humerus and anterior margin

of fossa of mesonotum with a dense patch of numerous setae. . 18

Modified tuft of fused setae of antennal flagellomere 8 in form of a

typical, smooth J-hook; humerus and anterior margin of fossa

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 19

without a distinet patch Of Setae. ck kw 3 ale ei ais 19

18(17). Dorsal beaklike process of lateral plate of phallosome slender,

long, sustaining about 90 degrees with main stem. ... vwariatus

Dorsal beaklike process of lateral plate of phallosome stout,

short, and strongly bent, forming an acute angle with

MSN Stem. 65. av eeeiwie re db oh eke pO eee ee ae jos ephinae

19(17). Basimere with a prominent row of 6,7 strong submarginal

SOCAG, 6/0 AOR AGN tale ae ee a a ees pairoji

Basimere with a weak row of 3-4 weaker submarginal setae. ... 20

20(19). Modified tuft of antennal flagellomere 5 largely composed of very

broad dark, scales onbiso0540 2 tiliwel whe Sie ees whartoni

Modified tuft of antennal flagellomere 5 composed of broad dark

scales dorsally and narrow, pale scales or flattened setae

1atOTALLY One: VOR TaRy Sc Wis ol ee ee ge are: Ge ais 21

21(20). Seta a of subapical lobe characteristically elbowed in apical

PUA hs 6d oc a3 tak ateanver a ae Oe wea ks ge Ea CP aR: cubitatus

Seta a of subapical lobe straight or uniformly smooth, without

SHOvO DOG ICAlL en ciohs extas Rie sete ere tO DAt Lee. ele 22

22(21). Submarginal setae of basimere usually 4 in number; basal

processes of palpal segment 1 long. ......... macdonaldi

Submarginal setae of basimeres 3 in number; basal processes

of palpal segment 1 relatively short. ......... gvacicornis

PUPAE!

- Setae 6-III-IV 5, 6 branched (Minutissimus Subgroup).

minutissimus, tnfantulus

Setae 6-III-IV usually 1-4 branched (Fraudatrix Subgroup). .... 2

2(1) a Alri a a tl be l= a ee ult pi gl ee ah ee ALES GP WE ge AN alphus

Seta LOE ddouble:or more: branchednsiaalealiviek TY -< ema ay . 3

3(2) Seta 9-C usually 3, 4 branched; seta 10-C 3,4 branched. . cinctellus

Seta 9-C single or double; 10-C double. .........4..2..2.. 4

4(3) Seta 5-V usually 5, 6 branched; 6-IV-VI usually 4-6 branched. .. 5

Seta 5-V usually double or at most 4 branched; 6-IV-VI usually

2%: DBE TSO vig cies prentaciibcle ek eat alto uaa ang = Bice tan Veneers Scat 6

5(4). Seta 7-II usually triple; 9-VII usually 4,5 branched. . . quadripalpis

Seta 7-II usually double, 9-VII usually triple. ...... aculeatus

EF SUES

alorensis not included; sentori, fulleri, gibbulus, paraculeatus, aestivus

and gvacicornis unknown.

6(4).

7(6).

8(7).

9(8).

10(7).

11(10).

12(11)-

13(12).

20:

3(2).

4(3).

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Seta -6=C: Single. 6 wr e4 oe GOP Oe whartoni

SebarG=C MOuble. 6 Laos a ee ee ee

Seta 1-C double; 6-IV, V usually double... 2 2 ee es 8

Seta 1-C triple or 4 branched; 6-IV,V 3,4 branched. ...... 10

Seta 5-1] 5 branched; 5-1] 4,5 branched..........5..%. cubitatus

Seta 5-II 3,4 branched; 5-IJI 3,4 branched. ............ 9

Integument dark brown; seta 5-IV 4,5 branched. ...... inculus

Integument pale yellowish; seta 5-IV double or triple. .... reiat

Integument dark brown or with striking pattern of dark and light

areas> seta 5-IV usually 7 branched... se a 11

Integument more or less uniformly pale yellowish or without

striking pattern of dark and light areas; seta 5-IV usually 6

branched. G5 oe ie Pe ee Ee a ee 12

Cephalothorax and abdomen entirely dark brown; paddle with dark

ald. Very StYOne MIdTID. 4 bo ke ie ee es niger

Cephalothorax and abdomen with pattern of dark brown and pale

areas; paddle with pale and weak midrib. ...... vubithovacits

Margin of mid-dorsal ridge of cephalothorax strongly folded or

very irregular; seta 4-VIII usually double. . macdonaldi, pairoji

Margin of mid-dorsal ridge of cephalothorax weakly folded or

more or less regular; seta 4-VIII usually triple. ....... 13

meta 02 V 5,4 branched: oo. er a a ee ee variatus

Seta 5-V dOUDIG. Gi ee jos ephinae

LARVAE!

Seta 2-VIII single; 14-P double (Vinutissimus Subgroup).

| minutissimus, tnfantulus

Seta 2-VIII double; 14-P usually single (Fvaudatrix Subgroup)... 2

Fraudatrix Subgroup

Seta S-Pisingle, oa a a re 5)

Sela 22 P doubters ee RS 6

Seta 5-C single (imculus complex). ........2.6.. oe aneulus

Seta 5-C double (quadripalpis complex). ......4.2.2+4-+4.-.

Seta 7-P usually double; 7-I single. .........2.+-e6.--. reid

Seta 7-P triple; f=) double,“ 0b a ee a 5)

La lorensis, not included; seniori, fulleri, gibbulus, pavraculeatus, aestivus

and gvacicornis unknown.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 21

5(4). Seta 3-P usually double; each siphonal tuft usually double.

aculeatus

Seta 3-P usually single; each siphonal tuft 3, 4 branched.

quadripalpis

6(2); Seta.5-C triple (einctellus complex) sinus i eek) Bw we cinctellus

Seta 5+€ double or, sinelewian siete 8 a ee ie Sea bee ae

7(6). Pecten teeth simple or with inconspicuous fringe of numerous fine

spines; comb scales strongly differentiated into an anterior

group of short ones and a posterior row of remarkably long

ones (a lbhus complex) aval oh aie li ee oe alphus

Pecten teeth with distinct fringe of strong denticles; all comb

scales similar in jeneth and, Size. 4:65 idee bk See ap 8

8(7). Seta. o-@ singlerig hs) aia mews creche: el er pies Ae ee niger

petacdsC dow le sca et sees siaie! Diocese Parti ay pe es Sr g

9(8). Seta 3-P multiple or at least 4 branched; 7-I single. . . vubithoracis

Seta 3-P single or double or at most 4 branched; 7-I double

(WEVICTUS COMDIER =: 4c ay chly wectas bay eg Pages eee Cae Es 10

10(9). Seta 14-P usually double; 2-X of saddle double. ...... cubitatus

Seta 14-P single; 2-X of saddle 3,4 branched. .......... 11

11(10). Seta 3-P 3,4 branched; 4-C longer than distance between bases

OE EINE HOA Mee Gai tices ee eee Gaia aap ap ung rus ie rales Sg a ok whartoni

Seta 3-P single or double; 4-C at most 0.5 as long as distance

betweenbases Of the pair. cs eo ye eee. Sea a 12

12(11). Seta 9-C larger than 8-C and placed at a considerable distance

eephaladsof the latter: ii). gd ee Oe a ee Wal macdonaldi

Seta 9-C subequal to 8-C and placed slightly cephalad of the

(he on eae err ES COUER en Te Ucsat gear Term nM Sr RT rar eect ene oe 13

13(12). -Setae 4 and-10-Vil-usually triples: 66k see ie ad oo patrvoq

setae’ 4 and 10=Vil double. erg wks So ee ee as 14

14013) Seta 3eP wusuallvidouble ss Vireo ei aera es variatus

peta. o> P usually singles 000 shane eee ee a eee josephinae

Minutissimus Subgroup

The Minutissimus Subgroup is characterized chiefly by the following char-

acters: in the adults of both sexes by (1) presence of basal transverse pale

bands on abdominal terga II-VII and (2) plume scales of veins Ro, Rg of wing

rather narrow, linear; in the female by proboscis with 4-6 labial basal setae;

in the male by (1) palpal segment 1 with rudimentary basal spiculose processes;

(2) palpal segments 4, 5 weakly plumose or with rather few lateral and mesal

bristles; (3) proboscis without dorsal upright sinuous setae and (4) modified

tufts of setae present only on mesal surface of antennal flagellomeres 7 and 8;

in the male genitalia by (1) distal division of subapical lobe without leaflets;

22 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

(2) submarginal setae of basimere poorly differentiated and widely spaced; (3)

phallosome with single beaklike process on dorsal lobe of lateral plate and (4)

proctiger with a small crown of few to several spinelike spicules; in the pupa by

(1) long, slender trumpet; (2) pinna of trumpet with slit extending to meatus; (3)

setae 10, 11-C double; (4) seta 5-IV 4-8 branched and (5) setae 6-III-VI 5, 6

branched; in the larva by (1) seta 4-C 1. 0-1.5 times as long as distance be-

tween bases of the pair; (2) seta 5, 6-C double; (3) seta 3-P usually single; (4)

seta 14-P double; (5) seta 7-I double; and (6) seta 2-VIII single.

DISCUSSION. The Minutissimus Subgroup is a complex of 3 species:

minutissimus, alorensis and infantulus, The first 2 of these are apparently

restricted in distribution; minutissimus is known only from Sri Lanka, India

and Thailand and alorensis from Alor, Indonesia while infantulus is widespread

with the range extending north and northeast into Japan and possibly also China

and Korea in the Palearctic.

The relationships between the Minutissimus and Fraudatrix Subgroups are

not clear, but both apparently fall into the same phyletic line (Fraudatrix

Group) on the basis of the phallosome of the male genitalia and the multitude

of characters of the immature stages. Colless (1967: 519-20), in discussing

the affinity of infantulus, pointed out that it does not fit either to the Fraudatrix

(group A) or the Mammilifer (group B) and suggested its placement in a third

group. This view is not justified since the other 2 members, minutissimus and

alorensis,do show the fraudatrix-type of male phallosome. Infact, the male

phallosome of infantulus is only slightly modified, especially in being strongly

reticulate, but its shape essentially resembles the other 2 related species and

all known members of the Fraudatrix Group. On the other hand, the presence

and the development of modified setal tufts on male antennal flagellomeres 7 and 8

in the Minutissimus Subgroup are, somewhat surprisingly, identical to those

of the Digoelensis Subgroup of the Mammilifer Group in New Guinea (Sirivana-

karn 1968: 101-6). The Minutissimus larvae resemble most members of the

Mammilifer Group by having seta 14-P double and seta 2-VIII single, as noted

by Colless (1967) for infantulus, but generally conform to the Fraudatrix Group

in most features of the chaetotaxy.

1. CULEX (LOPHOCERA OMYIA ) MINUTISSIMUS (THEOBALD)

(Fig. 1) 3

Culiciomyia minutissima Theobald 1907: 235 (9).

Culiciomyia nigerrima Theobald 1910: 233 (2); Edwards 1913: 235 (synonymy).

Melanoconion juxtapallidiceps Theobald 1910: 456 (¢); Edwards 1913: 235.

(synonymy).

Culex (Lophoceratomyia) minutissimus (Theobald), Edwards 1922a: 280

(*, key); Barraud 1924: 39 (o*, 2, L); Senior-White 1927: 71 (L*);

Edwards 1932: 197 (taxonomy); Barraud 1934: 363 (o‘*, 9, L); Feng

1938: 299 (distribution); Brug and Bonne-Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) minutissimus (Theobald), Stone, Knight and Starcke

1959: 234 (catalog); Harrison et al. 1974: 156 (distribution).

FEMALE. Wing: 2.8mm. Forefemur: 1.3 mm. Proboscis: 1.6 mm.

Small species; essentially conforms to the description of the Fvaudatrix Group

with the following diagnostic features. Head, Decumbent scales on dorsum

of vertex largely broad and entirely dark, not forming pale ocular line anteri-

orly; erect scales slender, entirely dark; lateral patch of broad scales dingy

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 23

white or dark bluish. Palpus dark scaled; 0.15-0.20 of proboscis length.

Proboscis dark scaled; labial basal setae 4-6, 2 lateral ones longest, as long

as palpus. Cibarial Armature, Not studied. Thorax. Mesonotal integument

dark; scales narrow, dark except for a few pale ones on extreme anterior

promontory, humerus and scutal angle. Pleural integument without pattern of

dark and pale bands; scales practically absent; ppl with 2 dark, strong bristles

and 3-4 pale, weak setae; 1 lower mep bristle present. Legs. Anterior sur-

face of hindfemur entirely dark. Wing. Plume scales on veins Ro, R3, R4i5

and branches of M and Cu narrow, linear and fine. Abdomen. Terga II-VI

with narrow, complete basal pale bands; sterna entirely dark or with banding

pattern as terga. :

MALE (Fig. 1). In general similar to female except for the following

sexual characters. Palpus. Slender, exceeding proboscis length by 0.5-1.0

of segment 5; segment 1 with a mesal pair of basal processes which are rudi-

mentary and rather inconspicuous; segments 4, 5 upturned, very weakly plu-

mose or with only a few bristles; apex of segment 5 bears 1,2 slender, dark

spines. Proboscis. False joint submedian, not clearly marked; apical 0.5 of

labium without upright setae on dorsal surface; labial basal setae 8-10, all

dark, weak, long, hairlike, as infemale. Antenna. Pedicel rounded, without

prominence; flagellum densely long plumose, with small modified tuft of setae

only on flagellomeres (F) 7 and 8, both mesal in position, readily seen from

above; F-7 with a minute tuft of 2,3 short, dark stout setae; F-8 with a strong

tuft of 3-4 long, dark bristlelike setae, as long as combined length of next 3

flagellomeres; F-12 about 1.5 times as long as F-13.

MALE GENITALIA (Fig. 1). Segment LIX, Tergum narrow, ribbonlike;

tergal lobe very small, bearing 2,3 weak setae; sternum broad, without

setae or scales. Basimere. Small, slender, conical, 0.20-0.25 mm. in

length; submarginal setae 2-6, widely spaced, in row parallel to tergomesal

margin. Subapical Lobe. Proximal rodlike setae a-c stout, straight, apically

hooked, 1 of which is thicker and broader than the other 2; weak seta present or

absent, ventrad of seta a; distal part not clearly separated from proximal part;

setae d-f4,5, all subequal, flattened, bladelike, 0.75 of length of setae a-c;

leaflets 21, 8) absent; seta h and 1,2 other weak setae present, laterad of

setae d-f. Distimere. Short and slender, about 0.5 of the length of basimere;

basal 0.95 straight and uniformly broad, apical 0.5 slightly curved downward

and tapered into a blunt apex; subapical claw slender and moderately long;

dorsal and ventral tiny setae present beyond middle of curvature, dorsal distad of

ventral. Phallosome. Dorsalprocess simple, beaklike, more or less resembling

duck head; ventral lobe large and rounded on apical sternal margin. Proctiger.

Apical crown very poorly developed, consisting of 4-6 toothlike spicules, in single

row; paraproct broad and strongly sclerotized, apical portion inform of a broad

plate; cercal sclerite broad and well sclerotized; cercal setae 4.

PUPA. Abdomen: 2.5mm. Paddle: 0.67 mm. Trumpet: 0.65 mm;

index 10. As figured for infantulus (Fig. 2). In general as described for the

group and subgroup with the following diagnostic chaetotaxy. Cephalothorax,

Seta 1-C 4,5 branched; 3-C usually triple (2-4); 5-C usually 5 branched (4-6);

8,9-C double. Metanotum, Seta 10-C usually triple (2-3); 11-C double. Abdbdo-

men. Seta 5-IV 4-8 branched; 5-V usually double, sometimes triple; 6-III-VI

0, 6 branched; 9-VII usually triple (2-4); 4-VIII double; 9-VIII usually 6,7

branched (4-7). Paddle. Broad, pale to almost transparent; midrib variable,

from very weak, pale to strong and dark pigmented.

LARVA. Head: 0.65 mm. Siphon: 1.6 mm; index 8-10. Saddle 0.3 mm;

siphon/saddle ratio 5-6. As figured for infantulus (Fig. 3). Head, Seta 1-C

24 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

dark, stout, spiniform; 4-C double or single, long, 1.0-1.5 times as long as

distance between bases of the pair; 5,6-C strong, as long as antenna, double;

7-C usually 7 branched (5-7); 10-C double or triple; 13-C 5,6 branched; 14-C

usually double (2-3). Antenna. Nearly as long as head; pigmentation largely

pale, cream-colored proximally, dark at base and beyond the insertion of seta

1-A; spicules numerous and distinct. Thorax. Spiculation absent; seta 3-P

usually double, sometimes single; 4-P double; 7-P triple; 8-P double, strong;

14-P double. Abdomen. Live specimens with dark greenish bands on seg-

ments I, II, III, V and VI, largely pale on segments IV and VIII; setae 6-I, II

triple; 7-I double; 6-III-VI 4 branched; 1-IJI-VI usually 3, 4 branched (3-5);

seta 1-VII usually 6 branched (4-8); 4 and 7-VII single; 13-VII 4 branched.

Comb scales about 40, in broad oval patch; all scales similar, subequal in

length and size and with rounded apical fringe of evenly fine spicules; seta 1-

VIII usually 5 branched (4-6); 2-VIII single; 3-VIII 7, 8 branched; 5-VIII 4, 5

branched. Saddle same color as pale portion of siphon; spiculation very weak;

seta 1-X usually triple (2-4); 2-X 3,4 branched; anal gills slender, pale,

fusiform, as long as or slightly longer than dorsal length of saddle. Siphon.

Slender, straight, long, distally tapering and with median dark ring; pecten

teeth 12-14, most distal tooth with 6,7 graded denticles; subventral tufts weak

and short, 4 pairs (or total 8), double or triple each, as long as siphonal width

at points of attachment.

TYPE-DATA. (1) Culiciomyia minutissima Theobald; Holotype °*,

CEYLON | SRI LANKA], Peradeniya, date not specified, collected by Green,

British Museum (Natural History) (BMNH). (2) Culiciomyia nigerrima Theobald;

Holotype 2*, CEYLON [SRI LANKA], Trincomalee, October 1907, collected

by Green (BMNH). (3) Melanoconion juxtapallidiceps Theobald; Holotype ¢*,

CEYLON [ SRI LANKA], Trincomalee, October 1907, collected by Green

(BMNH).

DISTRIBUTION. Known only from India, Sri Lanka and Thailand. 31

specimens examined: 10, 142, 1 P, 6 L; 8 with associated immature skins

(Pp, i lp),

INDIA. Bombay: Trombay; Karanja Jungle; Coimbatore; 5c", 22, 1 P,

5L(P. J. Barraud, 1921).

SRI LANKA. Colombo; Padaviya; Peradeniya; Trincomalee; 2°, 62 (in-

cluding 32 in the type-data), 1 L, 1 lp. :

THAILAND. Phrae: Padaeng Khawi; 30, 69, 7p.

Additional records from the literature: INDIA, Bombay: Deccan; Hubli;

Madras: Nilgiri Hills; Central Province: Nowgong; Orissa; Punjab (Barraud

1924: 41; 1934: 365); CHINA, Kwangtung (Feng 1938: 299).

TAXONOMIC DISCUSSION. Culex minutissimus is identical to alorensis

and infantulus in the male antenna and in the general external characters of

the adults. It can be definitely separated from the latter 2 species by the

shape of the male phallosome and by the details of the proctiger as indicated

in the key and as described and figured here. In the immature stages, the

minutissimus pupa differs slightly from that of infantulus in having most setae

relatively stronger and longer; seta 10-C usually triple (usually double in

infantulus) and seta 5-V usually double (usually 3,4 branched in infantulus),

The larva differs from infantulus in having seta 6-III 4 branched (triple in

infantulus) and in the presence of a median dark ring on the siphon (usually

absent in infantulus),

Except for the localities given above in the distribution data, most of the

previous records of minutissimus in the literature are doubtful and require

confirmation. The record from Vietnam by Borel (1930: 352-6) is undoubtedly

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 29

erroneous judging from the figures and the description of the male phallosome

and proctiger by this author. It appears most likely that Borel's specimens

are in fact those of infantulus which is more common and more widespread in

the Oriental region.

BIONOMICS. Culex minutissimus is a typical ground pool breeder. In

India it was reported from rock springs, pools in ravine and river-beds,

coconut shells, shallow wells, stagnant water in shaded culverts, etc. (Barraud

1934: 364-5). In Sri Lanka, the immatures were collected from eround pools

without further particulars. In Thailand, the pupae were collected from flooded

pools in mountain areas. The adults came from rearing the pupae or larvae

and were also caught by sweep nets near the water sources. The adult biology

is unknown.

2. CULEX (LOPHOCERAOMYIA) ALORENSIS NEW SPECIES

(Fig. 1)

Culex (Lophoceratomyia) minutissimus of Brug and Bonne-Wepster 1947: 186

(misidentification).

FEMALE. Small, light brown species; wing length about 2.7mm. Es-

sentially similar to minutissimus and infantulus in the presence of basal pale

bands on abdominal terga II-VII, differing slightly in the following features.

Head. Decumbent scales on dorsum of vertex narrower and predominantly

pale, forming distinct ocular line; lateral patch of broad scales whitish, ap-

parently broader and more distinct. Thorax, Integument of mesonotum pale

brown or yellowish brown but not blackish or reddish; mesonotal scales

same color as underlying integument.

MALE (Fig. 1). Essentially similar to minutissimus and infantulus in

palpus, proboscis and antennal characters, differing as described for female

and in the following additional features. Palpus. Variable in length, from

slightly shorter to longer than proboscis by length of segment 5. Antenna.

Flagellar whorls weakly to moderately plumose. Abdomen. Basal pale bands

usually well developed and distinct on terga IV-VII or V-VII, poorly developed

and indistinct on terga II-III or IIl-IV, latter sometimes completely dark

scaled or with only a few pale scales forming basolateral pale spot

MALE GENITALIA (Fig. 1). As figured; extremely similar to minutissi-

mus, differing from it particularly in the following features. Basimere.

Inner tergal surface with 1 strong seta in line with 2,3 other weaker setae.

Subapical Lobe. Seta h of distal part stronger and longer; setae d-f consist

of 3 subequal, sharply pointed blades and 2 shorter and weaker hairlike setae.

Distimere. Subapical claw longer; its apex strongly inflated or expanded.

Phallosome. Dorsal beaklike process more slender, shorter and with a re-

curved spine on posterior margin. Proctiger. Apical crown with a dense tuft

of several dark spinelike spicules; paraproct narrow with small rounded apical

lobe; cercal setae 3 in number.

PUPA. Abdomen: 2.2 mm. Paddle: 0.56mm. Trumpet: 0.55 mm;

index 10-11. Essentially similar to minutissimus and infantulus in general

and in detailed chaetotaxy, differing in the following combination of charac-

ters. Cephalothorax. Seta 8-C usually triple (2-3); 9-C single or double.

Abdomen. Seta 1-III usually 7 branched (6-9); 1-IV usually 7 branched (5-8);

1-V usually 5 branched (3-6); 1-VI, VII usually triple (3-4); 5-IV usually

triple (3-4); 5-V usually double (1- 2): 6-III usually triple (2-4); 6-IV-VI

26 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

usually 3,4 branched (3-5).

LARVA. Head: 0.65 mm. Siphon: 1.6 mm; index 8-9. Saddle: 0.30

mm; siphon/saddle ratio 6. Essentially similar to minutissimus and infantu-

lus and as figured for the latter (Fig. 3), differing from both particularly in

the following. Thorax. Seta 7-P usually double (1-2); 8, 9-M usually 4

branched (4-5). Abdomen. Setae 1-III-VI usually triple (2-3); 6-IV-VI triple;

1-VII usually 4 branched (3-4). Saddle seta 2-X double. Siphon. Yellowish

white, median dark ring absent. |

TYPE-DATA. Holotype & (80-65) with slide of antenna and genitalia (69/

297), INDONESIA, Alor, date not specified, collected by Rodenwaldt, U. S.

National Museum (USNM). Paratypes: 1% (80-63) with slide of antenna and

genitalia (69/299) and 10° (80-62) with attached genitalia mount on pin, Pantar,

June 1925, collected by Von Beck (USNM); 2, (TM-0047), 2 lp? (TM-004702,

14) (USNM), 2% (TM 0047), 2 lp? (TM 0047-1, 2) and 22 (TM 0047) (J.C. Lien

coll., Taiwan), Timor, Halibot, larvae in stream pool, 6-7 October 1973,

J. C. Lien and Soeroto, collectors.

DISTRIBUTION. Known only from Alor, Pantar and Timor, Indonesia.

13 specimens examined: 7°, 69; 4 with associated larval and pupal skins.

INDONESIA. Alor (type-locality); 1% (holotype), as indicated in the type-

data; Pantar; 20° (paratypes), as indicated in the type-data; Timor: Halibot,

Malak Timur, Belu Reg. 4c, 69, 4 lp; (all designated as paratypes); reared

from larvae collected in stream pool, 6-7 October 1973, J. C. Lien and

Soeroto.

TAXONOMIC DISCUSSION. Culex alorensis was previously recorded by

Brug and Bonne-Wepster (1947: 186) as minutissimus based on the adult

males from Alor and Pantar. Preliminary study of this material and sub-

sequent study of the specimens recently reported by Lien et al. [| 1975: 335 as

Culex (Lophoceraomyia) sp. | from Timor has shown that it is distinct from

minutissimus and infantulus in the shape of the phallosome and in the develop-

ment of the proctiger crown of the male genitalia. It is also quite possible

that the record of minutissimus by Brug and Bonne-Wepster (loc. cit.) from

Sulawesi (Celebes) is in fact that of alorensis,

Culex alorensis is closely related to minutissimus and infantulus and with

the latter 2 species, falls into the Minutissimus Subgroup as recognized here.

The male phallosome and proctiger of alorensis are apparently more similar

to members of the Fvaudatrix Subgroup than to that of minutissimus and injfan-

tulus.

BIONOMICS. The specimens from Alor and Pantar presumably came from

a general field catch but no definite field data were given. In Timor, immatures

were collected in a stream pool. Nothing is known about the adult biology of

alorensis,

3. CULEX (LOPHOCERAOMYIA) INFANTULUS EDWARDS

(Figs. 2; 3)

Culex (Lophoceratomyia) infantulus Edwards 1922a: 287 (“*, 9°, key);

Edwards 1932: 197 (taxonomy); Baisas 1935: 174 (c"*); Causey 1937: 416

(distribution); Feng 1938: 298 (distribution); LaCasse and Yamaguti 1950:

197 (o*, 9*, P*, L*); Brug and Bonne-Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) parainfantulus Menon 1944: 389 (%, ¢); Mattingly

1949: 224 (synonymy).

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region ar

Culex (Lophoceraomyia) infantulus Edwards, Bohart 1945: 75 (o°, L); Bohart

and Ingram 1946: 73 (o*, 9, L*, P*); Iyengar and Menon 1955: 10 (L*);

Peters and Dewar 1956: 46 (L); Stone, Knight and Starcke 1959: 233

(catalog); Delfinado 1966: 106 (o“*); Colless 1967: 519 (taxonomy); Bram

1967a: 58 (o°*, 2, L*); Baisas 1974: 111 (“, °, P, L); Harrison etal.

1974: 156 (distribution).

Culex (Lophoceratomyia) minutissimus of Borel 1930: 353 (misidentification).

FEMALE. Wing: 2.3-2.8 mm (average 2.6 mm). Forefemur: 1.4 mm.

Proboscis: 1.7mm. Abdomen: 1.6-2.3 mm. Small, blackish species; as

described and figured by LaCasse and Yamaguti (1950: 197-9)and Bram (1967a:

09-61). In general extremely similar to minutissimus from which it can not

be separated with certainty. Cibarial Armature (Fig. 2). Cibarial bar with

about 60 narrow, long, apically blunt teeth in an even concave row. Legs.

Anterior surface of hindfemur usually with pale whitish stripe, sometimes

entirely dark. Abdomen. Basal transverse pale bands of abdominal terga

variable, usually broad, sometimes narrow or incomplete.

MALE. Essentially similar to minutissimus and alorensis in palpal,

labial and antennal characters and cannot be separated from the latter 2

species with certainty.

MALE GENITALIA (Fig. 2). As figured, differing strikingly from

minutissimus and alorensis in the phallosome and slightly in other features

as in the following. Basimere. Submarginal setae 5, 6 in number, 3 of which

are strong, subequal in length, 2 or 3 other setae weaker and shorter. Sub-

apical Lobe. Setae a-c subequal in length and thickness. Distimere. Subapi-

cal claw slender, long, as in alorensis. Phallosome. Dorsal beaklike pro-

cess of lateral plate slender, long, largely reticulose or strongly imbricate

with numerous reticules. Proctiger. Apical crown larger, consisting of

numerous dark spinelike spicules; paraproct narrow, weakly sclerotized, with-

out distinct apical lobe; cercal setae 3-5.

PUPA (Fig. 2). Abdomen: 2.4-2.8 mm (average 2.6 mm). Paddle:

0.52-0.69 mm (average 0.65 mm). Trumpet: 0.56-0.78 mm (average 0.65

mm); index 9-12. As figured; extremely similar to minutissimus, differing

in having most setae relatively weaker and particularly in the following.

Metanotum. Seta 10-C usually double (2-3). Abdomen. Seta 5-V usually 3, 4

branched (2-4); setae 6-III-VI usually 5,6 branched, sometimes 3, 4 branched.

LARVA (Fig. 3). Head 0.65-0.78 mm (average 0.72 mm). Siphon 1.5-

2.0 mm (average 1.8 mm); index 8-10. Saddle 0.27-0.34 mm (average 0.3

mm); siphon/saddle ratio 5-6. As figured, differing slightly from minutissi-

mus particularly in the following. Thorax, Seta 3-P single or double; 4-P

usually triple, sometimes 4 branched. Abdomen. Seta 6-III usually triple

(3-4). Siphon. Median dark ring usually absent or sometimes median portion

slightly darkened, but not forming a distinct ring. }

TYPE-DATA. (1) Culex (L.) infantulus Edwards 1922; Holotype “* with

attached genitalia mount; HONG KONG, 1914, collected by Dr. H. Macfarlane

(BMNH). (2) Culex (L.) pavainfantulus Menon; Type (not specified); INDIA,

Trivandrum, Tvavancore (Madras states), location of the type unknown.

DISTRIBUTION. Widely distributed throughout the Oriental region. 548

specimens examined: 259", 1932, 7 P, 89 L; 248 with associated immature

skins (138 p, 110 lp).

INDIA. Bombay: Kawar, N. Kanara; 1c, 19.

NEPAL. Hetaura; 4c.

MALDIVE ISLANDS: Fiori; 1.

28 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

BURMA. Rangoon; 5c.

SRI LANKA. North Central Province: Anuradhapura, Padaviya; 1°.

THAILAND. Mae Hong Son: Doi Chang; Mae Sariang; Chiang Mai: Muang;

Chom Tong; Huey Kaeo; Chang Phuak; Ban Chang Kien; San Kampang; Ban

Rong Rua Taeng; Ban San Kha Yom; Nan: Na Noi; Ban Pong Mon; Lampang:

Ngao; Ban Pha Khoi; Ban Pha Daeng; Ban Rong; Ban Rong Pako; Ban Rong Na;

‘Ban Pha Khoi; Nakhon Ratchasima; Koraj, Pachong; Huey Sai Noi; Prachin

Buri: Ban Thap Lan; Ban Bu Phram; Chanthaburi: Tha Mai; Ban Plu; Nontha-

buri; Ayutthaya: Bang Pa-In; Chon Buri: Bang La Mung; Khao Mai Kaeo; Khao

Mai Hawa; Prachuab Khiri Khan: Khlong Hin Chong; Krabi: Ban Mai Kien Tai;

Ban Pru Taci; Surat Thani: Koh Samui; Pathum Thani; Satun; 123%, 1072, 42 L,

142 p, 40 lp. :

VIETNAM. Cam Ranh; 2c, 3 L.

MALAYSIA. Peninsular Malaysia (Malaya): Serdang; Perak: Pusing;

Kuala Kangsar; Kedah: Changlum; Perlis: Kg. Sanglang; 200, 15%, 8 L, 19 lp.

' PHILIPPINES. Luzon: Agoo La Union; Subic Naval Base; Malawin Creek,

Mt. Makiling; Mindoro: San Jose; Samar: Osmena; Leyte: Palo; Lago Lago,

Baybay; Mindanao: Parang; Zambales; 340, 352, 13 L, 49 lp.

HONG KONG. 4c, 49.

JAPAN. Ryukyu Islands - Okinawa: Chizuka; Kana village; Nakasoni;

Nago; Iriomote: 1.5 mi. N. Toyokara village; Yabu; 400, 13°, 10 L, 3 lp;

Nagasaki; Honshu; Kyoto; 5“, 392, 4L.

Additional records from the literature: INDONESIA, Java (Brug and

Bonne-Wepster 1947: 186); CHINA, Anhwei, Chekiang, Kiangsi, Kwangtung

(Feng 1938: 298).

TAXONOMIC DISCUSSION. Culex infantulus is one of the most widespread

forms of Lophoceraomyia in the Oriental region. It occurs sympatrically

with minutissimus in northern Thailand, southern India, Sri Lanka and possibly

with alorensis among the Lesser Sundas of Indonesia. It is possible that some

of the previous records of minutissimus from several localities in India by

Barraud (1924, 1934) are actually infantulus, Of all stages, the male genitalia

of infantulus is most distinctive, particularly in the reticulate dorsal process

of the phallosome.

Culex infantulus exhibits variations in all stages, but there is no indication

that it is locally differentiated into any recognizable geographic forms among

the material I have examined. The adults from Japan are larger and paler than

those from Southeast Asia and the basal abdominal bands are variable from

broad, narrow, complete to incomplete or sometimes absent, especially

among the material from Peninsular Malaysia but all are remarkably constant

in the details of the male genitalia. The pupa and larva of infantulus exhibit

much overlap with minutissimus in most features of the chaetotaxy but can be

separated from the latter by certain diagnostic features as indicated in the

above description.

BIONOMICS. The pupa and larva of infantulus are most frequently found

in small ground pools, including ditches, ponds, wells, footprints, tracks

and occasionally in rock pools, natural or artificial containers, such as palm

bracts, axils and earthenware jars and the margins of slow moving streams

and creeks. The records from tree holes and bamboos in Thailand by Bram

(1967a: 61) appear to be incorrect. They were probably based on erroneous

field notes. The breeding sites are at a broad range of elevation and are usual-

ly under shade of tropical forests. The adults have been collected while resting

on damp vegetation and stream banks and occasionally also from light traps.

In a successful colonization of infantulus by Miyagi (1972) in Nagasaki, Japan,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 29

the females were reported to readily feed on frogs, snakes, turtles as well as

on chicks and mice. The development from larvae to emergence of imagos

at room temperatures of 24-26°C lasts 9-11 days and the adults have a life

span of 15 to 18 days (Miyagi 1973).

Fraudatrix Subgroup

The Fraudatrix Subgroup is characterized by the following characters:

in the adults of both sexes by (1) abdominal terga usually entirely dark, some-

times with basolateral pale spots or with basal pale bands and (2) plume scales

of wing veins Ro, Re relatively broad clavate or ovate; in the female by pro-

boscis usually with 2 labial basal setae, sometimes 4-6; in the male by (1)

palpal segment 1 with 2 distinct pairs of spiculose basal fingerlike processes;

(2) proboscis usually with 2 prominent rows of upright sinuous setae on dorsal

surface, sometimes reduced or absent; (3) labial false joint present, usually

marked by flexion at 0.25 from extreme base and (4) modified tufts of setae

and scales present on antennal flagellomeres 5-10 or 6-10; in the male geni-

talia by (1) subapical lobe of basimere usually with 2 leaflets (£1, 8), some-

times 1; (2) submarginal setae of basimere usually strongly differentiated,

forming a distinct row on inner tergal surface, sometimes in a broad patch;

(3) lateral plate of phallosome with a simple apical beaklike process (or dor-

sal process) on dorsal lobe (DL) and (4) proctiger crown small, with few to

several spicules; in the pupa by (1) long, slender trumpet; (2) pinna of trumpet

with slit extended to meatus; (3) setae 10, 11-C usually double; (4) seta 5-IV

3-7 branched and (5) setae 6-III-VI usually 2-4 branched; in the larva by (1)

seta 4-C 0.25-1.00 times as long as distance between bases of the pair; (2)

seta 3-P usually single or double, sometimes multiple; (3) seta 14-P usually

single; occasionally double; (4) seta 7-I usually double, sometimes single and

(5) seta 2-VIII double.

DISCUSSION. The Fraudatrix Subgroup as recognized here includes

practically all species previously placed in the Fvaudatrix Group in a strict

or broad sense of Edwards (1932), Belkin (1962), Colless (1965) and Sirivana-

karn (1968). The Fraudatrix Subgroup is the largest in the subgenus, compris-

ing approximately 68 species: 20 in the Oriental region, 33 in the Australasian

(Papua New Guinea and Australia); 12 in the South Pacific and 3 in Micronesia.

All representatives of the Fraudatrix Subgroup in each respective area are

basically similar in the comparatively simple male phallosome, but are slight-

ly or strikingly differentiated from one another in the details of the male pal-

pus, proboscis, modified tufts of antennal flagellomeres 5-8 and 9, submar-

ginal setae of the basimere, setae of the subapical lobe, development of the

apical dorsal beaklike process of the phallosome and the apical lobes of the

paraproct of the proctiger. The relationships among the members of com-

plexes of the Fvaudatrix Subgroup are very close. However, there is apparent-

ly little or no overlap in species composition between the Oriental fauna and

those from the Australiasian regions and the Pacific islands. There is also

a strong indication of endemicity and independent development of the subgroup

in each faunal area, which seems to further suggest that the speciation of the

group probably occurred at the same time subsequent to the separation of the

land masses. Most of the Oriental forms appear to be more primitive than

those from the Australasian regions and the Pacific islands, but this can not

be accurately determined until the entire fauna is more thoroughly known and

compared in details with respect to their character states.

30 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

The Fraudatrix Subgroup is represented in the Oriental region by 7 com-

plexes of 20 species. The characterization of the 7 complexes given below is

limited to the most clearly marked and constant correlated features to indicate

the diversity within the subgroup.

(1) seniorvi complex. The seniori complex includes only seniovi, It is

chiefly characterized in the male, the only known stage by: (1) the rudimen-

tary basal fingerlike processes of the palpus; (2) the absence of dorsal upright

setae on the proboscis; (3) modified tufts of the antenna poorly developed and

are restricted only to flagellomeres 6-10; (4) leaflet g] and seta / of the sub-

apical lobe absent or not developed; (5) submarginal of the basimere poorly dif-

ferentiated; (6) apical dorsal beaklike process of the phallosome strong and

moderately long; (7) paraproct with a single, short, rounded apical lobe adja-

cent to crown and (8) abdominal terga with basal pale bands.

(2) cinctellus complex. This complex includes the widespread cinctellus

and the Philippine fulleri. It is characterized in the male by (1) well developed

basal fingerlike processes of the palpus; (2) presence of dorsal upright setae on

the proboscis; (3) modified tufts of the antenna well developed on flagellomeres

5-10; (4) F-5 with a small tuft of 2, 3narrow or broad lanceolate scales, as long

as the next 2 flagellomeres, followed by 2,3 narrow, finely pointed scales of

increasing length; (5) leaflet g; and seta h of the subapical lobe present; (6)

submarginal setae of basimere numerous, in a broad patch; (7) apical dorsal

beaklike process of the phallosome strong and moderately long; (8) apical lobe

of paraproct poorly developed and (9) abdominal terga with basal pale bands;

in the larva by (1) seta 5-C 3,4 branched; (2) seta 4-P double and (3) seta 7-I

double.

(3) vubithoracis complex. The vubithoracis complex is represented by

rubithoracis which is common and very widespread throughout the Oriental

region, the Malaysian niger and the Philippine gzbbulus, This complex is dif-

ferentiated from the cinctellus complex and the next 4 complexes below particu-

larly in the following: the male by (1) dorsal upright setae of proboscis pre-

sent or absent; (2) relatively short basal fingerlike processes of palpus; (3)

F-5 with a small, inconspicuous tuft of 4,5 short, narrow, linear, finely

pointed scales or setae; (4) basimere with a row of 1-3 relatively weak sub-

marginal setae; (5) all specialized setae and leaflets of subapical lobe well

developed; (6) apical dorsal beaklike process of phallosome slender and

moderately long; (7) paraproct of proctiger with well developed apical lobe or

lobes and (8) abdominal terga without basal pale bands; the larva by (1) seta

5-C double or single; (2) seta 4-P double and (3) seta 7-I single.

(4) inculus complex. The inculus complex is represented by imculus which

is a rare Southeast Asian form. It differs from rubithoracis and the next 3

complexes particularly in the following: the male by (1) dorsal upright setae

of proboscis present; (2) basal fingerlike processes of palpus relatively long

and distinct; (3) F-5 with a small, but distinct tuft of 8-10 dark, narrow, finely

pointed scales; (4) basimere strongly swollen, with 1,2 prominent rows of 10-.

12 strong submarginal setae; (5) leaflet g, and gq of subapical lobe very broad;

(6) apical dorsal beaklike process of Si inceine anvil-shaped, with several

fine and sharp spicules on ventral surface and (7) paraproct without differenti-

ated apical lobe; the larva by (1) seta 5-C single; (2) seta 4-P single and (3)

seta 7-I single.

(5) quadripalpis complex. The quadripalpis complex comprises 6 species:

quadripalpis, aculeatus, pavaculeatus, aestivus and reidi. It is characterized

by the following combination of characters: in the male by (1) dorsal upright

setae of proboscis present; (2) basal fingerlike process of palpus relatively long

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 31

and very distinct; (3) F-5 with a distinct medium-sized tuft of 3-8 broad,

blunt tipped or abruptly pointed scales and narrow fine-tipped scales as long

as the next 3-4 flagellomeres; (4) basimere normal, usually with 3-7 weakly

to strongly developed submarginal setae in a row, sometimes 1; (5) setae of

subapical lobe well developed, leaflets g; and go varied; (6) apical dorsal

process of phallosome typically beaklike, length and thickness varied and (7)

paraproct usually with well developed lateral and sternal apical lobes, shapes

varied; in the larva by (1) seta 5-C double; (2) seta 4-P single and (3) seta 7-I

double.

(6) variatus complex. The variatus complex is composed of 7 species:

variatus, josephinae, cubitatus, gracicornis, whartoni, macdonaldi and pairoji.

It is strongly characterized in the male by the large fan-shaped tuft of several

broad scales on antennal flagellomere 5 and by the following additional features:

(1) dorsal upright setae of proboscis present, well developed; (2) basal finger-

like processes of palpus varied, well developed; (3) basimere with a row of

3-7 strong submarginal setae; (4) all specialized setae of subapical lobe well

developed, varied; (5) apical dorsal beaklike process of phallosome stout,

length varied and (6) apical lobes of paraproct poorly or well developed; in the

larva by (1) seta 5-C double or triple; (2) seta 4-P double and (3) seta 7-I

double.

(7) albhus complex. The alphus complex is restricted to alphus which

strongly resembles the members of the variatus complex in the modified tuft

of male antennal flagellomere 5. It is chiefly characterized by the following:

in the male by (1) basal fingerlike processes of palpus relatively long, reach-

ing beyond apex of palpal segment 1; (2) modified tuft of antennal F-5 large,

fan-shaped, composed of several broad scales; 3-6 scales in lateral ventral

group with characteristic swelling in distal subapical portion; (3) apical dorsal

beaklike process of phallosome very short and slender and (4) paraproct of

proctiger with a small apical sternal lobe only; in the larva by (1) seta 5-C

double, reduced in length, about 0.5-0.75 of 6-C; (2) seta 4-P double; (3) seta

7-I single; (4) comb scales strongly differentiated in length and size, those in

the posterior row elongate, 2,3 times as long as the anterior ones and (5)

pecten teeth simple or with a weak fringe of numerous minute denticles.

seniovi complex

4, CULEX (LOPHOCERAOMYIA) SENIORI BARRAUD

(Fig. 6)

Culex (Lophoceratomyia) seniori Barraud 1934: 365 (c'*),

Culex (Lophoceraomyia) seniori Barraud; Stone, Knight and Starcke 1959: 235

(catalog).

FEMALE. Unknown.

MALE (Fig. 6). Based on the holotype (wing length 2.8 mm). Palpus.

Slender, long, exceeding proboscis by 0.5 of the length of segment 5; segment

1 with 2 pairs of very short, tiny, subequal basal fingerlike processes which

are about 0.5 of distal portion of segment 1; apical 0.1 of segment 3 with ven-

trolateral tuft of 4,5 bristles; segments 4 and 5 moderately plumose. Pro-

boscis. Dorsal upright setae on labium absent or not developed; labial basal

setae 10, all of which are short, subequal, stout, spinelike, in a transverse

row on ventral surface; false joint not clearly marked. Antenna. Pedicel

P Contrib. Amer. Ent; Inst., vol. 13, no, 4, 1977

simple, rounded; flagellar whorls densely long plumose; modified tufts of

scales and setae present only on F-6 to F-10; F-6 with a small mesal tuft of

4-5 dark, thickened, more or less fused setae, as long as the length of 1 flagel-

lomere; F-7 with a rudimentary ventral tuft of 3 dark, short, flattened, api-

cally blunt setae and a small mesal tuft of 3,4 longer hairlike setae; F-8 with

a large, prominent mesal tuft of 5 dark, fused bristlelike setae, as long as the

next 3 flagellomeres; F-9 with a mesal tuft of 2 dark, long bristlelike and 2, 3

hairlike setae, as long as the next 5, 6 flagellomeres; F-10 with similar tuft

of setae as on F-9. Thorax. Integument of mesonotum and pleuron light

brown; mesonotal scales dark, narrow, moderately dense; pleuron without

distinct scale patch, a few scales present on upper corner of stp; ppl bristles

4: 1 lower mep bristle present. Legs. Anterior surface of fore- and midfe-

mora dark; anterior surface of hindfemur with longitudinal pale stripe from base

to near apex; tibiae and tarsi of all legs dark. Wing. Scales on all wing veins

dark and rather scanty; scales on veins Ro, Rg and R4,5 small, broad clavate.

Abdomen. Tergal, II entirely dark; terga III- VII with basal transverse pale

bands; sterna entirely pale beige or yellowish.

MALE GENITALIA (Fig. 6). As figured and as in the following description.

Basimere. Conical; tergomesal margin lightly convex; inner tergal surface

with several strong setae, which are widely scattered, not forming a row along

tergomesal margin. Subapical Lobe. Rodlike setae a-c of proximal part sub-

equal and straight, setae a and c apically hooked, seta } tapered into a point;

distal part with 5 bladelike setae in group d-f, most distal seta (= leaflet £9)

longest, lanceolate, the rest shorter and narrower; leaflets gj and seta h ab-

sent. Distimere. Normal, without subapical crest of spicules; subapical claw

moderately long; 1 tiny ventral seta present distad of middle curvature, dorsal

seta absent. Phallosome. Apical dorsal lobe of lateral plate represented by

a stout, short, beaklike process, sustaining about 90 degrees with the main stem;

ventral lobe well developed and rounded on sternal margin. Pyroctiger. Apical

crown relatively small, composed of dark, short, spinelike spicules only;

paraproct well sclerotized, broad, with a distinct apical lobe; cercal sclerite

well sclerotized; cercal setae 2.

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype “* with slide of genitalia, palpus, proboscis and

antenna; Garden Reach, Calcutta, INDIA, October 1931, R. Senior-White

(BMNH).

DISTRIBUTION. Known only from the type-locality. 1 specimen examined:

o (marked as type), as indicated in the type-data.

TAXONOMIC DISCUSSION. Culex seniori is known only from the single

type male originally described and figured by Barraud (1934: 365). It is

essentially similar to all members of the Fraudatrix Subgroup in the develop-

ment of the basal processes of the palpus, the spinelike labial basal setae of

the proboscis and in the type of phallosome of the genitalia but can be readily

recognized by (1) presence of basal pale bands on the abdominal terga; (2)

absence of dorsal upright setae on the proboscis; (3) characteristic modified

tufts of setae on antennal flagellomeres 6-10 and (4) details of setae of the sub-

apical lobe of the genitalia as indicated in the key and as described above. Be-

cause of several distinctive features of the male, seniovi is here placed in its

own complex separated from the other complexes of the Fraudatrix Subgroup.

BIONOMICS. The male of seniovi apparently came from a general field

catch. No further data are available.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 33

cinctellus complex

9. CULEX (LOPHOCERAOMYIA) CINCTELLUS EDWARDS

(Figs. 4, 5)

Culex (Lophoceratomyia) cinctellus Edwards 1922a: 287 (new name for Lopho-

ceratomyia taeniata Leicester 1908, nec Wiedemann; o*, key); Barraud

1924: 42 (*); Edwards 1932: 197 (taxonomy); Barraud 1934: 366 (c'*, &,

L); Brug and Bonne-Wepster 1947: 186 (distribution); Chu 1958: 110

(distribution).

Culex (Lophoceraomyia) cinctellus Edwards, Stone, Knight and Starcke 1959:

232 (catalog); Colless 1965: 277 (“*, 2, L*); Delfinado 1966: 103 (o"*);

Bram: 1967ay 55 (0%). 2s 1*)s

FEMALE. Wing: 3.0mm. Forefemur: 1.44 mm. Proboscis: 1.85 mm.

Abdomen: 2.2 mm. Medium-sized species, distinguished from all members

of the Fraudatrix Subgroup except seniori by the presence of basal pale bands

on the abdominal terga and by the following features. Head. Decumbent scales

of vertex largely broad, pale anteriorly, forming a distinct ocular line, darker

posteriorly; narrow decumbent scales restricted to occiput and dorsal midline

(coronal suture) of vertex; lateral patch of broad scaleswhitish, distinct. Pal-

pus varying from 0.15-0.20 of proboscis length. Proboscis with 2 labial

basal setae which are dark, hairlike, about 0.5 of palpal length. Czbarial

Armature (Fig. 4). Cibarial bar concave, with 60 or more teeth in close-set

row, all of which are narrow, fine, more or less pointed, subequal in size

and length. Thorax. Mesonotal integument light to dark brown; scales nar-

row, dark brown. Ppn with some scattered dark scales and tiny setae ce-

phalad of 3,4 posterior bristles. Pleuron same color as mesonotum or slightly

paler; upper corner of stp with or without a few pale scales; Ppl bristles 4-6;

upper mep setae 7; 1 lower mep bristle present. Legs. As in seniorvi, with-

out any marked coloration; anterior surface of hindfemur with longitudinal pale

stripe in basal 0.5, apical 0.5 dark scaled. Wing. Plume scales on veins Ro,

R3 and R4,5 moderately broad, clavate. Abdomen. Tergum II with narrow

basal pale bands, not completely extended laterad; terga III-VII with complete

basal pale bands; sterna entirely pale. :

MALE (Fig. 4). In general conforms to the Fraudatrix Subgroup, with the

following distinctive characters. Palpus. Exceeding proboscis by a little more

than the full length of segment 5; basal fingerlike processes short, less than

0.5 of the length of distal portion of segment 1; segment 3 with 4-10 ventro-

lateral bristles in apical 0.1-0.2, ventral surface with 1,2 fine, short, hair-

like setae; segments 4, 5 upturned and strongly plumose. Proboscis. Apical

0.5 or more of labium with a double row of fine, upright, sinuous setae on

dorsal surface, longest setae about 3 times as long as labial width; labial basal

setae short, stout, spinelike, 12 in number; false joint present at 0.20-0. 25 of

the length from base. Antenna. Flagellar whorls densely long plumose,

modified tufts cf scales and setae well developed on lateral basal tubercles of

long normal setae of flagellomeres 5 to 10; F-5 with a small dorsolateral tuft

of 2,3 broad, lanceolate or abruptly pointed scales which are as long as the

next 2 flagellomeres, followed by 2,3 narrower, fine-tipped scales of increas-

ing length; F-6 with a dark crumpled tuft of several heavily curled setae on

lateral ventral surface; F-7 with a lateral tuft of about 10 curled, apically

twisted setae in row resembling the shape of a comb and a ventral mesal tuft

34 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

of 3,4 stout, proximally fused, curled setae; F-8 with a ventral slender tuft of

6-8 fused setae in form of a J-hook; F-9 with a ventral tuft of 4,5 long, distally

broad, bladelike scales and several short, hairlike setae; F-10 with a ventral

tuft of 2,3 dark, long, distally curved bristlelike setae.

MALE GENITALIA (Fig. 4). Segment IX. Tergal lobe very small, bearing

a row of 2 or 3 short setae; sternum broad, usually entirely bare or sometimes

with 1 strong seta on median caudal margin. Basimere. Very stout, conical,

about 0.2 mm in length; basal 0.5 lightly to strongly swollen; inner tergal sur-

face with 15 or more strong submarginal and marginal setae aggregating into a

patch along tergomesal margin, 4,5 lateral setae are longest and strongest,

the rest shorter and weaker; setae on outer tergal area laterad of submarginal

setae and subapical lobe sparse. Subapical Lobe. Prominent; rodlike setae a-c

of proximal part very stout and gently curved in middle; distal part with setae

in group d-f strong, bladelike, 5,6 in number, seta 5 strongly differentiated

from setae d-f as a broad, lanceolate leaflet; leaflet g; very broad, with acu-

minate apex, setah strong. Distimere. Thick, more or less straight from

base to 0.75 of total length; apical 0.25 weakly curved, lightly to strongly

swollen on ventral surface; subapical claw well developed; ventral subapical

seta very strong, 2,3 times as long as dorsal subapical seta. Phallosome.

Apical dorsal beaklike process of lateral plate strong and moderately long,

apical margin smooth. Proctiger. Apical crown medium-sized, with several

dark spinelike spicules; paraproct strongly sclerotized, moderately to strongly

pigmented, apical lobe not developed; cercal sclerite moderately to strongly

pigmented; cercal setae usually 2, sometimes 3.

PUPA (Fig. 4). Abdomen: 2.5mm. Paddle: 0.7mm. Trumpet: 0. 65-

0.70 mm; index 10. Chaetotaxy as figured, distinctive in the following.

Cephalothorax, Seta 1-C double; 8-C single; 9-C 3,4 branched. Metanotum.

Seta 10-C 3,4 branched. Abdomen. Seta 6-III-VI single or double; 5-IV 4, 5

branched; 5-V 2-4 branched; 5-VI usually double (1-3); 9- VII double or triple;

4-VIII usually double (2-3); 9-VIII usually 5 branched (4-6). Paddle. Midrib

moderately to strongly pigmented.

LARVA (Fig. 5). Head: 0.7mm. Siphon: 1.8 mm; index 9. Saddle: 0.35

mm; siphon/saddle ratio 4. General features and chaetotaxy as figured, the

following are diagnostic. Head. Seta 4-C distally forked into 3, 4 branches,

its length about 0.5 of the distance between bases of the pairs; 5-C 3, 4 branched;

6-C double. Thorax. Spiculation not developed; seta 3-P usually double, some-

times single or triple; 4-P double. Abdomen. Unspiculated; seta 7-I double;

1-III-VI usually 4 branched (3-5); 6-III-VI usually 4 branched (4-5). Saddle

lightly spiculated on posterior caudal margin; seta 2-X usually 4 branched;

anal gills as long as or longer than saddle. Siphon. Very slender, same

color as head capsule, without median dark ring; pecten teeth 10-11, most

distal tooth with 12, 13 fine denticles; subventral tufts very weak and rather

inconspicuous, 4 pairs, all of which are double, subequal, shorter or as long

as siphonal width at points of attachment; 2-S very weak and short; median

caudal filament of spiracular apparatus well developed.

TYPE-DATA. Lectotype “* (originally marked as a cotype of Lopho-

cevatomyia taeniata Leicester 1908) with attached genitalia mount, marshy

ground, Circular Road, Kuala Lumpur, (Selangor), Malaya | MALAYSIA],

December 10, 1903, G. F. Leicester (BMNH; selection of Colless 1965: 278).

DISTRIBUTION. Widespread throughout Southeast Asia, also reported

from India. 140 specimens examined: 1040", 259, 11 L, 9 with associated

immature skins (4 p, 5 lp).

INDIA. Assam: Rupsi; Dibrugarh; 2.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 30

THAILAND. Khon Kaen: Chumphae; Pa Dong Lan; Chon Buri; Siracha,

Bang Phra; Bang La Mung; Khao Mai Kaeo; Nakhon Si Thammarat; Choung

Khao; Navathiwat: Ban Kado; 120, 1L, 2p, 1 lp.

VIETNAM. Danang; Ankhe; Lai Khe; 22, 1 L.

MALAYSIA. Peninsular Malaysia - Selangor: Kuala Lumpur, Circular

Road; Pacific Tin; Kuang; Pahang: Cameron Highlands, Mt. Brinchang;

Bentong Road; Perak: Strait Settlements; Kedah; Pusing; Kuala Kangsar;

Johore: Kota Tinggi; Malaysia - Sarawak, Kuching; Sabah: Kota Kinabalu

Tawau; 330°, 212, 10 L; 2p, 4 lp.

SINGAPORE. 3°, 29.

INDONESIA. Java: Djakarta; Kebon Ros; Belakang Ponrok; Surabaja,

Bengkulu; 50,

PHILIPPINES. Luzon; Samar; San Antonio; Leyte: Palo; Dulag; Tacloban,

Diit River, Burugwin; 49.

Additional records from the literature. CHINA: Hainan Island (Chu 1958);

INDONESIA: Sumatra (Brug and Bonne-Wepster 1947); INDIA: Malabar Coast,

Pudupadi (Barraud 1934): JAPAN: Ryukyus, Yaeyama (Bohart 1959).

TAXONOMIC DISCUSSION. The adults of cinctellus can be readily recog-

nized by the presence of basal pale bands on the abdominal terga and by the

features of the palpus, proboscis and the modified tufts of the antenna of the

male as indicated in the keys and in the above description. The cinctellus fe-

male may be confused with those of infantulus, minutissimus, alorensis and

seniorvi which exhibit similar abdominal tergal pale bands. It can, however,

be separated from these species by the relatively larger size, the broader

plume scales on wing veins Ro, Rg and R45; the presence of only 2 labial basal

setae of the proboscis and by having pale decumbent scales of vertex of the

head forming a very distinct ocular line. The male genitalia are most diagnostic

in the shape of the basimere, the aggregation of several strong submarginal

setae into a broad patch on the inner tergal surface of the basimere and in the

development of the setae of the subapical lobe. The pupa can be readily recog-

nized by the single seta 8-C; the larva by the 3,4 branched seta 5-C and by

several combinations of characters as given above.

Culex cinctellus is closely related to fullevz Ludlow from Mindanao, the

Philippines, on the basis of the male genitalia and with the latter apparently

falls into a distinct complex of the Fraudatrix Subgroup. Both species resemble

seniovi more than any other in the shape of the basimere and apical dorsal

process of the phallosome of the male genitalia.

BIONOMICS. The immatures of cinctellus have frequently been collected in

large bodies of water on the ground such as marshy depressions, swamps,

ditches and stream margins under partial or heavy shade of forests. On occa-

sion, they have also been collected from small ground pools. The elevation

ranges from above sea level to 1,000 m. The adults were collected by light

traps, animal bait traps and sweep nets while resting or biting. In Malaysia,

they were commonly taken while resting among vegetation in forests or more

open areas but were rarely taken with mammalian bait (Colless 1965: 278).

The females are probably largely bird feeders (Colless 1959) but they were

also reported to bite and feed on man in the forest canopy (Macdonald and

Traub 1960).

36 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

6. CULEX (LOPHOCERAOMYIA) FULLERI (LUDLOW)

(Fig. 6)

Oculiomyia fulleri Ludlow 1909: 97 ().

Culex (Lophoceratomyia) fulleri (Ludlow), Edwards 1922a: 287 (c'*, key);

Edwards 1932: 197 (taxonomy).

Culex (Lophoceraomyia) fulleri (Ludlow), Bohart 1945: 74 (o"*); Stone and

Knight 1957: 50 (lectotype designation): Stone, Knight and Starcke 1959:

233 (catalog); Delfinado 1966: 104 (c", 9).

FEMALE. Wing: 3.4mm. Forefemur: 1.6 mm. Proboscis: 2.0 mm.

As described for cinctellus, differing from it in the absence of basal pale

band on abdominal terga and in the following characters. Head. Broad decum-

bent scales of vertex largely pale. Czibarial Armature, Cibarial teeth appar-

ently finer and more numerous, about 80 in number. Thorax. Ppl bristles

6,7. Abdomen. Terga entirely dark dorsally, terga III-VII with small baso-

lateral pale spots.

MALE (Fig. 6). Differing from cinctellus as described for the female

and in the following additional features. Palpus. Fine hairlike setae on ven-

tral surface of palpal segment 3 more numerous and rather denser. Antenna,

Modified tuft of F-5 with 4,5 narrower, fine tipped scales; F-9 with 5, 6 darker

and broader bladelike scales; F-10 with 3,4 stronger, more flattened bristle-

like setae.

MALE GENITALIA (Fig. 6). As figured; exceedingly similar to cinctellus,

differing constantly in the following details. Basimere. Larger, about 0.25

mm in length; basal portion strongly swollen; submarginal setae stronger,

densely packed, 4 or 5 of which are longest, 5 times as long as shorter sub-

marginals. Subapical Lobe. Leaflets 1 and £9 broadly foliate in apical por-

tion; setae d-f shorter and narrower. Distimere. Thicker and longer, strong-

ly swollen beyond middle before tapering to a blunt point apically; subapical

portion strongly annulated; subapical claw shorter. Pyroctiger. Paraproct

with well developed apical sternal lobe sternad of crown of spicules; cercal

setae 4 in number.

PUPA and LARVA. Unknown.

TYPE-DATA. Lectotype: ¢*, Parang (Cotabato), Mindanao, PHILIPPINES,

October 25, 1908, Major Fuller (USNM; selection of Stone and Knight 1957: 50).

DISTRIBUTION. Known only from Mindanao and Jolo Jolo islands in the

Philippines. 137 specimens examined: 77°", 609.

PHILIPPINES. Mindanao: Parang, Ludlow Barracks; Kabakan; Agusan S.

Francisco; 15¢, 602; Jolo Jolo'ls. ; 2¢.

TAXONOMIC DISCUSSION. The Philippines records of fulleri from Leyte

and Luzon by Bohart (1945) and Delfinado (1966) are doubtful and need to be

confirmed since I have not seen any specimens from these areas. It is pos-

sible that these records were based on the misidentified specimens of cinctel-

lus which is widespread in those islands.

Culex fulleri exhibits constant differences from cinctellus in several de-

tails of the male genitalia and in the absence of the basal pale bands on the

abdominal terga but is basically similar to the latter in the characters of the

male antenna, palpus, proboscis and in the shape of the basimere and the

phallosome. It also appears to be geographically isolated from cinctfellus,

suggesting that it may be only subspecifically distinct.

BIONOMICS. The breeding site of fullervi is not known but is most probably

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 37

the ground pool habitat as in cinctellus, All of the adults apparently came from

general field catches by net and light traps. Nothing is known about the adult

feeding behavior and its medical importance.

vubithoracis complex

7. CULEX (LOPHOCERAOMYIA) RUBITHORACIS (LEICESTER)

(Figs. 7, 8)

Lophoceratomyia rubithoracis Leicester 1908: 119 (%, 9).

Culex (Lophoceratomyia) rubithoracis (Leicester), Edwards 1932: 197

(taxonomy); Barraud 1934: 367 (*, 2); Hsiao and Bohart 1946: 29

(distribution); Bohart 1946: 16 (key); LaCasse and Yamaguti 1950: 192

(o*, O*,--P*#e L*)> Hara 1957%:-57(9 eenttalia*).

Culex (Lophoceraomyia) rubithoracis (Leicester), Mattingly 1949: 224 (L*,

key); Stone, Knight and Starcke 1959: 235 (catalog); Colless 1959: 114

(L); Lien 1962: 632 (distribution, biology); Colless 1965: 281 (o*, @

L*); Delfinado 1966: 113 (<); Bram 1967a: 69 (o'*, 9, L*).

FEMALE. Wing: 2.1-2.7 mm (average 2.2 mm). Forefemur: 1.0 mm.

Proboscis: 1.4mm. Small, yellowish brown species; in general as described

for the Fraudatrix Group and Subgroup with the following diagnostic features.

Head, Decumbent scales of vertex broad, pale along upper eye margin, form-

ing distinct ocular line, dark posteriorly; narrow, linear decumbent scales

relatively few in number, restricted to coronal suture and occiput; erect scales

short, dark and rather sparse; lateral patch of broad scales pale beige or yel-

lowish white. Palpus very slender, 0.2 of proboscis length. Proboscis with

2 labial basal setae, nearly as long as palpal length. Cibarial Armature (Fig.

7). As figured, cibarial bar relatively short, with about 30 teeth in concave

row; median 4-6 weak, short, lateral teeth stronger, flattened and apically

blunt. Thorax. Mesonotal integument with characteristic hue of bright yellow,

orange or sometimes reddish brown, but not dark brown or chestnut brown;

scales on mesonotal disc narrow, linear, dark brown, sparse or moderately

dense. Ppn with some scattered, semi-erect narrow scales cephalad of 2-4

strong posterior bristles. Pleural integument same color as mesonotum;

scales on upper corner of stp practically absent; ppl bristles few, 1 or 2 of

which are strong and dark; lower mepbristle absent. Legs. Without any

marked coloration. Wing. Scales on all veins rather scanty; plume scales

On veins R Rg and Ry,5 narrow, linear in basal portion, broader in apical

portion. Asdamans Terga entirely dark, basolateral pale spots absent; terga

V-VII usually with narrow lateral pale stripe: sterna whitish or yellowish.

MALE (Fig. 7). In general similar to female, with the following diagnos-

tic features. Palpus. Very slender and thin, longer than proboscis by 0. 5-

1.0 of segment 5; basal fingerlike process of segment 1 minute, short, rather

inconspicuous; basal 0.5 of segment 2 with a row of dark, short, spinelike

setae; ventral surface of segment 3 with a row of fine, minute setae which are

dense in basal 0.5, sparse in apical 0.5, its apical 0.25 with a ventrolateral

tuft of 5,6 dark bristles; segments 4 and 5 moderately to strongly plumose.

Proboscis, Slender; ventral surface slightly paler than dorsal surface;

distal 0.5 of labium with distinct double rows of dorsal sinuous setae

which are 2 or 3 times as long as labial width; labial basal setae dark,

stout, spiniform, 10 in number. Antenna. Flagellar whorls densely long plu-

38 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

mose; modified tufts well developed on F-5 to F-10; F-9 with a small tuft of

2-5 very narrow, finely pointed scales which are pale yellowish to blackish,

subequally long, as long as the combined length of the next 3 flagellomeres;

-F-6, 7 with dark crumpled tufts of heavily curled setae as described for the

eroup; J-hook tuft of F-8 very slender, composed of 6, 7 fused setae; F-9 with

5 dark, very broad, bladelike scales; F-10 with 2,3 dark long bristlelike setae.

MALE GENITALIA (Fig. 7). As described and figured by Colless (1965:

281-2). Segment IX, Tergal lobe very small, bearing 2,3 weak and short

setae. Basimere. Normal, small, slender, 0.17 mm in length; inner tergal

surface with 3 strong submarginal setae in a row parallel to tergomesal mar-

gin; marginal setae 6,7 widely spaced, 2,3 basal ones strongest; strong bris-

tles largely confined to lateral tergal areas. Subapical Lobe. Rodlike setae

a-c of proximal part stout, subequal, lightly curved in distal portion; leaflets

1, gg and seta # of distal part well developed; leaflet g1 club-shaped; setae

d-f 5,6, all flattened, bladelike, 2,3 of which are strongly bent in middle.

Distimere. Slender, lightly or strongly curved distad of midpoint; subapical

claw small, short, apically dilated and rounded; 1 ventral and 1 dorsal tiny

seta present near apex. Phallosome. Apical dorsal beaklike process of

lateral plate short and relatively slender. Proctiger. Apical crown very

small, or poorly developed, consisting of only a few pale or dark spinelike

spicules; apex of paraproct with a characteristic budlike apical lobe; lateral

paraproct and cercal sclerite well sclerotized; cercal setae 2.

PUPA (Fig. 7). Abdomen: 2.0mm. Paddle: 0.52 mm. Trumpet: 0.65

mm; index 11-12. Cephalothorax yellowish, with or without brownish areas

on apex of head shield, posterior mid-dorsal ridge, leg, wing and labial cases.

Trumpet darker than underlying integument. Chaetotaxy as figured, the fol-

lowing combination of seta branching is diagnostic. Cephalothorax, Seta 1-C

usually triple (3-4); 3-C usually double (2-3); 8,9-C double. Metanotum,. Set-

ae 10, 11-C double. Abdomen. Seta 6-III, IV usually triple (2-4); 6-V, VI usu-

ally 4 branched (3-4); 5-IV 6-8 branched; 5-V usually 4 branched (3-6); 5-VI

double or triple, rarely single; 9- VII double or triple; 4-VIII double or triple;

9-VIII 4-6 branched. Paddle. Basal external margin of outer part usually

with characteristic infuscated area; midrib weak and pale with characteristic

dark spot towards apex; setae 1,2-P present, minute.

LARVA (Fig. 8). Head: 0.6mm. Siphon: 1.0-1.4 mm; index 9. Saddle:

0. 28-0.30 mm; siphon/saddle ratio 4. As characterized for the group and

complex; chaetotaxy as figured; the following are diagnostic. Head. Seta 13-C

3,4 branched; 16,17-C very well developed and distinct. Thorax. Spiculation

light to moderate, usually visible under 10X objective; seta 3-P usually 4 or

more branched (2-11); 4-P double. Abdomen. Unspiculated; seta 7-I always

single; 1-III-VI 4, 5 branched; 6-III-VI usually 4,5 branched (4-6). Comb

scales numerous (40-50), all similar in size and with even apical fringe of fine

spicules. Saddle same color as siphon; seta 2-X with 3 short and 1 long

branched; ventral brush (4-X) with 5 pairs of setae; anal gills 1.5 times as

long as saddle. Siphon. Slender, thin, moderately long and uniformly yellow-

ish distally; pecten teeth 10, each barbed with 10-12 fine denticles; subventral

tufts 4 pairs (total 8), all subequal, very weak, short, double or triple each;

slightly longer than siphonal width at points of attachment; 2-S very short and

weak; median caudal filament of spiracular apparatus very well developed.

TYPE-DATA. Lectotype “* with attached genitalia mount, Circular Road,

Kuala Lumpur, [ Selangor], Malaya | MALAYSIA], marsh, November 6, 1903,

G. F. Leicester (BMNH; selection of Colless 1965: 281).

DISTRIBUTION. Widespread throughout the Oriental region; also reported

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 39

from Japan. 413 specimens examined: 360, 42°, 14 L; 28 with associated

immature skins (11 pupal, 17 larval).

INDIA. Assam: Dibrugarh; Bengal: Lalmanirhat; 2°.

BURMA. Rangoon; 2°, 39.

SRI LANKA. Western Province: Colombo District, Waga; 29.

THAILAND. Chiang Mai: Muang; San Pha Thang, Ban San Khayom; Mae

Rim; Ban Chang Kien; Udon Thani: Muang, Ban Nong Bua; Khon Kaen; Nontha-

buri: Pak Kret; Bangkok; Phaket: Ban Borae; Ban Pha Thong; 3890, 49, 5 L;

9 py od Ip:

CAMBODIA. Phnom Penh; 4%, 1% genitalia.

VIETNAM. Con Son; Phu Lai; Vung Tau; Cam Ranh; 4c, 9, 3 L.

MALAYSIA. Peninsular Malaysia - Johore: Kg. Kahang; Tvengganu:

Kuala Brang; Pahang: Pdg. Tungku; Pevak: Pusing; Chior; Tg. Rambutan;

1807.1. 259,)-3-19 2 8A

SINGAPORE. 4c, 39.

INDONESIA. Java: Djakarta; 90°; Sumatra: Dermajo; 1“; Celebes; 2°.

PHILIPPINES. Luzon: Subic Naval Base; Mindoro: San Jose; Leyte:

Tacloban; Palo; Sulu Archipelago (Mindanao): Jolo Is,; 25%, 4°, 9 lp.

HONG KONG. 1¢.

JAPAN. Honshu: Kyoto; 1°, 1L.

Additional records from the literature. MALAYSIA and INDONESIA:

Borneo (Brug and Bonne-Wepster 1947: 186); CHINA: Chekiang; Kwangtung

(Feng 1938: 299); TAIWAN (Lien 1962: 632).

TAXONOMIC DISCUSSION. The adults of rubithoracis can be readily

recognized by the small size, the orange or bright yellow integument of the

thorax and by the absence of the lower mesepimeral bristle. Males can be

recognized by the small modified tuft of 2-5 narrow, finely pointed scales of

antennal flagellomere 5 and the dark, subapically bladelike scales of antennal

flagellomere 9. The male genitalia are very diagnostic, particularly in the

presence of a row of 3 submarginal setae on the inner tergal surface of the

basimere; the small proctiger crown and in the characteristic budlike apical

lobe of the paraproct. In the immature stages, the pupa is readily recognized

by the dark spot on the apex of the paddle midrib and the larva by the multi-

branched seta 3-P, the single branched seta 7-I and by having only 9 pairs of

setae in the ventral brush (4-X) of the saddle.

Culex rubithoracis is one of the most common species in the subgenus and

is apparently as widespread as infantulus, All stages of rubithoracis are

variable, but there is no indication of geographic differentiation among the

material I have examined. It is evidently closely related to niger and gibbulus

on the basis of the male characters as well as the immature stages. Together

with these 2 species it falls into a distinct complex of the Fraudatrix Subgroup.

The members of the rubithoracis complex also appear to be related to the New

Guinea pseudorubithoracis complex of Sirivanakarn (1968: 178-80); however,

their true affinity can be definitely determined only when the immature stages

of the New Guinea forms are known.

BIONOMICS. The immatures of vubithoracis are most frequently found in

open, sunlit habitats such as ricefields and marshy depressions, quite differ-

ent from most forms of the Fvaudatrix Group which usually breed in heavily

or partially shaded ground pool habitats in forests. The breeding sites of rubi-

thoracis are in most cases, small ground pools, puddles and ditches with

emergent or submerged aquatic vegetation such as grasses, duckweed and Lily

40 Contrib. Amer. Ent. Inst., vol. 138, no. 4, 1977

pads in the flat swampy ground just above sea level. The numerous adults

collected in Thailand came from light traps. Nothing is known about adult

feeding preferences and the medical importance.

8. CULEX (LOPHOCERAOMYIA) NIGER (LEICESTER)

(Figs. 9, 10)

Lophoceratomyia niger Leicester 1908: 123 (“, 9).

Culex (Lophoceratomyia) atratulus Edwards 1922a: 287 (new name for niger

Leicester 1908, in error).

Culex (Lophoceratomyia) niger (Leicester), Edwards 1932: 197 (taxonomy).

Culex (Lophoceraomyia) niger (Leicester), Stone, Knight and Starcke 1959:

235 (catalog); Colless 1965: 282 (“*, 9, L*).

FEMALE. Wing: 2.7mm. Forefemur: 1.3 mm. Proboscis: 1.9 mm.

Small, blackish species with the following distinctive features. Head, Broad

decumbent scales on upper eye margin and posterolateral area of vertex black-

ish; narrow decumbent scales pale, shiny brown, numerous, occupying a broad

area in center and on occiput; lateral patch of broad scales predominently dark

except for a few bluish white ones along eye margin. Palpus rather short,

about 0.17 of proboscis length. Proboscis with 2 labial basal setae which are

moderately long, about 0.5 of palpal length. Ciberial Armature: Not studied.

Thorax. Integument of mesonotum dark brown to almost black; mesonotal

scales blackish; dorsocentral and scutellar bristles very strong and dark.

Ppn with a few dark scalelike setae cephalad of 4 posterior bristles. Pleuron

as dark as mesonotum in upper 0.5, slightly paler in lower 0.5; Ppl with 4

bristles and 3,4 other short and weak setae; 1 lower mep bristle present; lower

posterior portion of mep with a minute patch of about 5 semi-erect scalelike

setae. Legs. Anterior surface of hindfemur with a silvery white longitudinal

stripe extending from base to near apex. Wing. Scales in apical portion of

veins Ro and Rg moderately broad clavate. Abdomen. Terga entirely dark or

black; terga III-VII with basolateral pale spots; sterna pale yellowish.

MALE (Fig. 9). As described for the female except for the absence of

basolateral pale spots on the abdominal terga, differing from other members of

yubithovacis complex in the following characters. Palpus, Longer than pro-

boscis by the full length of segment 5; basal fingerlike process of segment 1

distinct, moderately long, about 0.5 of the distal part of the segment; segment

3 with 1 ventrolateral bristle subapically; segments 4 and 9 weakly plumose,

with several weak and short bristles. Pvoboscis, Dorsal upright setae ab-

sent. Antenna. Flagellar whorls strongly plumose; F-5 with a small tuft of

4-6 dark, narrow, finely pointed scales which are subequally long, as long as

the combined length of the next 4 flagellomeres; F-6 to F-8 with similar

modified tufts of setae as in rubithoracis; F-9 with 2,3 bladelike scales; F-10

with 2 long bristlelike setae. 3

MALE GENITALIA (Fig. 9). Exceedingly similar to rubithoracis, differing

from it particularly in the following features. Segment Ix, Sternum with 1

strong seta towards caudal margin. Basimere. Inner tergal surface with 1 or

sometimes 2 other weak submarginal setae in a row; median tergal area laterad

of subapical lobe without any setae. Subapical Lobe. Bases of rodlike seta

a-c of proximal part with a group of 4, 5 apically hooked fine setae on mesal

surface; setae in group d-f 5, all of which are flattened, bladelike; leaflets 91,

go broad foliate and apically acuminate, leaflet gy apparently shorter and

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 41

smaller than leaflet go. Phallosome. Apical dorsal beaklike process of later-

al plate very slender, long, with apex terminated into a sharp, curved point.

Proctigery. Crown small, consisting of about 10 spinelike spicules aggregating

into a tuft on mesal surface; apex of paraproct very characteristic, with a long,

slender thumblike external lobe and a broad, apically rounded sternal lobe;

cercal setae 2,3.

PUPA (Fig. 9). Based on 1 associated skin. Abdomen: 2.2 mm. Paddle:

0.65 mm. Trumpet: 0.65 mm; index 10. Essentially similar to vubithoracis

in the detailed chaetotaxy, differing from it particularly in the following.

Cephalothorax and Metanotum, Entirely brownish. Trumpet. Slender, long,

as dark as underlying integument. Abdomen. Segments I-V brownish or same

color as cephalothorax; segments VI-VII slightly paler. Paddle. Midrib

strong and dark pigmented; areas surrounding midrib strongly infuscate; basal

external margin of outer part pale or not infuscate.

LARVA (Fig. 10). Based on 1 associated skin. Head: 0.65 mm. _ Siphon:

1.3 mm; index 7. As figured, differing from rubithoracis in the following

characters. Head, Seta 5-C single; 7-C very dark, with 10 flattened, strong-

ly plumose branches. Thorax. Spiculation absent or not developed; seta 3-P

double. Abdomen. Seta 7-I double. Siphon. Subventral tufts stronger and

longer, 9 pairs, each 3,4 branched, 1.5-2.0 times as long as siphonal width

at points of attachment.

TYPE-DATA. Lectotype “* with slide of antenna, palpus, proboscis and

genitalia, Circular Road, Kuala Lumpur, | Selangor|, Malaya | MALAYSIA],

October 7, 1903, G. F. Leicester (BMNH: selection of Colless 1965: 282).

DISTRIBUTION. Known only from Selangor, Malaysia. 6 specimens

examined: 30, 3°, 1 with associated lp skins.

MALAYSIA. Peninsular Malaysia. Selangor: Kuala Lumpur; 1“ (holo-

type), 12 (cotype); Puchong Rd. ; 20, 29, 1 lp.

TAXONOMIC DISCUSSION. Culex niger is a rare Malaysian species. It

exhibits a stronger affinity with rubithoracis than with any other species on the

basis of the morphology of all stages. It can be readily distinguished from

rubithovacis and other members of the Fraudatrix Subgroup in the adults of both

sexes by the presence of semi-erect scalelike setae on the posterior lower por-

tion of the mesepimeron (mep); the male by the small tuft of narrow, pointed

scales of the antennal flagellomere 5 and the absence of dorsal upright setae

on the proboscis; the male genitalia by the long thumblike apical lobe of the

paraproct of the proctiger; the pupa by the dark brown cephalothorax and abdo-

men and in the larva by the single seta 5-C, the dark, flattened and plumose

branches of seta 7-C; the double seta 3-P and the double seta 7-I.

BIONOMICS. According to Colless (1965: 283), the larvae of niger have

been found on one occasion amongst lotus leaves at the edge of a partly shaded

Swamp in association with the vubithoracis larvae. Nothing is known about the

adult biology.

9. CULEX (LOPHOCERAOMYIA) GIBBULUS DELFINADO

(Fie, 1%)

Culex (Lophoceraomyia) gibbulus Delfinado 1966: 106 ('*).

FEMALE. Unknown.

MALE (Fig. 11). Wing: 2.7mm. As described by Delfinado (1966: 105-

6), with the following additional description and corrections. Head. Vertex

42 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

with broad pale scales restricted to eye margin, forming distinct ocular line;

narrow decumbent scales numerous, occupying an extensive area in center;

lateral patch of broad scales distinct. Palpus, As long as proboscis; basal

fingerlike process of segment 1 about 0.5 of the distal part of the segment;

segment 3 with 1,2 dark, short, ventrolateral bristles subapically; segments

4 and 5 very weakly plumose, with a few dark, short bristles on lateral and me-

sal surfaces. Proboscis. Labium without dorsal upright setae; labial basal

setae relatively long. Antenna. Flagellar whorls moderately plumose; F-5

with a small tuft of 4,5 narrow, dark, hairlike setae, slightly longer than the

combined length of the next 3 flagellomeres; modified tufts of F-6, 7 as des-

cribed for the group, comb tuft of F-7 with about 14 curled setae, 4 dorsal-

most of which are strong and long, the remainder weaker and shorter; F-8

with normal J-hooked tuft; F-9 with 4 yellowish bladelike scales; F-10 with

2,3 bristlelike setae. Thorax. Integument of mesonotum brownish, mesonotal

scales narrow, same color as integument. Pleuron same color as mesonotum;

ppl bristles 3; 1 lower mep bristle present. Legs. As in rubithoracis, Wing.

Plume scales of veins Ro, Rg small, clavate. Abdomen. Terga entirely dark,

sterna pale yellowish.

MALE GENITALIA (Fig. 11). Segment IX. Sternum broad, with a trans-

‘verse row of 10 strong setae towards caudal margin; tergal lobe small, bearing

a few weak setae. Basimere. Small, normal, conical; inner tergal surface

with 1 strong submarginal seta. Subapical Lobe, Elongate; rodlike setae a-c

relatively long; setae in group d-f 4,5, placed on a hairy, elongate tubercle or

stem, leaflet g5 bladelike, club-shaped; leaflet g, (or external leaf) absent or

not developed; seta / strong. Distimere. Normal, relatively long, nearly as

long as basimere. Phallosome. Apical dorsal beaklike process of lateral

plate moderate in length and thickness as in vubithoracis. Proctiger. Crown

relatively larger, with 5,6 coarse spicules laterally and several fine pointed

spicules internally; paraproct narrow, without distinct apical lobe; cercal

setae 2.

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype o* with slide of antenna and genitalia, San Jose,

Mindoro, PHILIPPINES, March 9, 1945, E. S. Ross (USNM). 3

DISTRIBUTION. Known only from Mindoro (type-locality) and Palawan,

the Philippines. 2 specimens examined: 2%,

PHILIPPINES. Mindoro: San Jose; 1“ (holotype); Palawan: Brook's Point,

Uring-Uring; 1 slide of “ antenna and genitalia, rest of specimen lost.

TAXONOMIC DISCUSSION. Culex gibbulus is readily separated from the

other 2 members of the rubithoracis complex, rubithoracis and niger in the

male (the only known stage), by the peculiar development of the comb-shaped

tuft of antennal flagellomere 7, the absence of leaflet g; and the elongation of

the distal part of the subapical lobe of the genitalia. The male of gibbulus also

strikingly differs from rubithoracis in the complete absence of the dorsal up-

right setae of the proboscis, the presence of a lower mesepimeral bristle and

the dark brown coloration of the thorax; from ”iger in the smaller tuft of anten-

nal flagellomere 5, paler coloration of the thorax and the absence of the minute

patch of the semi-erect scalelike setae on the lower posterior portion of mese-

pimeron. In theoverall male morphology, gibbulus apparently exhibits stronger

affinity with the 2 members of the vubithovacis complex than with the other

members of the Fvaudatrix Subgroup. It is tentatively assigned to this complex,

pending a thorough knowledge of the immature stages.

BIONOMICS. Nothing is known about the biology of gzbbulus,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 43

inculus complex

10. CULEX (LOPHOCERA OMYIA) INCULUS COLLESS

(Figs): 12,013)

Culex (Lophoceraomyia) inculus Colless 1965: 280 (“*, 9, L*).

FEMALE. Wing: 2.5mm. Forefemur: 1.1 mm. Proboscis: 1.6 mm.

Small, blackish species. Head, Decumbent scales on dorsum of vertex large-

ly broad and dark, not forming a distinct ocular line; lateral patch of broad

scales whitish, very distinct. Labial basal setae of proboscis strong, nearly

as long as palpus. Cibarial Armature. Not studied. Thorax. Mesonotal

integument dark brown to black, scales narrow and black. One lower mep

bristle present. Legs and Wing. Without any marked coloration. Abdomen.

Terga entirely dark; basolateral pale spots present on terga IJ-VII; sterna sil-

very white scaled.

MALE (Fig. 12). As described by Colless (1965: 280-1); distinctive in the

following features. Palpus. Longer than proboscis by a little more than the

length of segment 5; basal fingerlike processes distinct, relatively long, as

long as the distal part of segment 1; segment 3 with 6,7 ventrolateral bristles

subapically; segments 4 and 5 moderately plumose. Proboscis. Dorsal up-

right setae present in apical 0.5 of labium. Antenna. Flagellar whorls strongly

plumose; F-5 with a small dark tuft of 8-10 narrow, acute scales of variable

length, longest ones as long as the next 2 flagellomeres; F-8 with a relatively

thick J-hooked tuft of 9 proximally fused setae; F-9 with 3, 4 dark bladelike

scales; F-10 with 3 strong bristlelike setae.

MALE GENITALIA (Fig. 12). Very distinctive. Segment IX. Tergal lobe

with 4,5 very strong setae; sternum without any scales or setae. Basimere.,

Modified conical shaped; basal half strongly swollen, with a pronounced convex

tergomesal margin, apical half narrowed; submarginal setae very strong, 10-

13 in number, in 1,2 prominent rows on convex tergomesal margin; marginal

setae numerous and dense; bristles and setae on lateral tergal surface relative-

ly stronger than most forms. Subapical Lobe. All specialized setae well de-

veloped; leaflets 8} and 8 very prominent; 85 club-shaped, distally broad and

bent upwards; g, broad, acuminate. Distimere, Stout, more or less straight

or not strongly curved in middle. Phallosome. Apical dorsal process anvil-

shaped, lightly or strongly imbricate with irregular ridges or folds, ventral

surface of elongate tergal beaklike process provided with several tiny spines.

Proctiger. Crown well developed, medium sized, with several fine pointed

spicules; apex of paraproct without differentiated apical lobes; cercal sclerite

broad, well sclerotized; cercal setae 1, 2.

PUPA (Fig. 13). Abdomen: 2.3mm. Paddle: 0.60 mm. Trumpet: 0.52

mm; index 10. In general as described forthe group. Cephalothorax and abdo-

men pale yellow or cream-colored. Trumpet. Brownish, cylindrical or slight-

ly widened towards apex. Cephalothorax. Setae 1,3-C double; 5-C triple; 8,

9-C double. Metanotum. Setae 10,11-C double. Abdomen. Seta 5-IV 4,5

branched; 5-V, VI double; 6-III, IV double; 6-V, VI usually double (2-3); 4-VIII

single or double. Paddle. Very pale to almost transparent, midrib weak and

_ lightly pigmented.

LARVA (Fig. 13). Head: 0.65 mm. Siphon: 1.0 mm; index 6. Saddle:

0.26 mm; siphon/saddle ratio 4. Generally conforms to the group, with the

following diagnostic features. Head, Seta 4-C minute, 0.25 of the distance

44 Contrib. Amer. Ent. Inst., vol. 13, no. 4; 1977

between bases of the pair; 5-C single; 14-C double. Thorax, Unspiculated;

seta 3-P single; 4-P single. Abdomen, Seta 7-I single; 6-III-VI usually

triple (3-4). Comb scales 40, normal. Saddle seta 2-X with 1 short and 1

long branch; 4-X with 6 pairs of setae. Szphon. Uniformly yellowish, without

‘median dark band and moderately long; pecten teeth 14-16, distal teeth with

1,2 strong basal denticles and 6,7 distal denticles; subventral tuft strong,

prominent, 4 pairs, all more or less subequal, 5-7 branched each, about 1.5

times as long as siphonal width at points of attachment.

TYPE-DATA. Holotype “* (604/20) with associated larval and pupal skins,

slides of antenna (CH 132) and genitalia (CT 427), Kampong Sireh, Selangor,

Malaya | MALAYSIA], March 31, 1953, J. A. Reid, Australian National Insect

Collection, Canberra (ANIC).

DISTRIBUTION. Malaysia and Cambodia; 3 specimens examined: 2° (in-

cluding holotype), 12; 2 with associated larval and pupal skins.

MALAYSIA. Peninsular Malaysia - Selangor; 1, 192, 2 lp (type-series).

CAMBODIA. Sihanoukville; 1° (only antenna and genitalia slide, collector:

J. M. Klein, 15 Jan. 67).

TAXONOMIC DISCUSSION. Culex inculus is strongly differentiated from

the rest of the Fraudatrix Subgroup by the peculiar anvil-shape of the male

phallosome and by several other features of the male genitalia as described

above. The male is somewhat intermediate between the vubithoracis and

quadripalpis complexes in the characters of the palpus and the antennal tuft of

flagellomere 9 and in the chaetotaxy of the larva. The female resembles vreidi

in the quadripalpis complex more than any others and cannot be distinguished

from that species with certainty. This species can be well assigned to the

quadripalpis complex but because of the very distinctive male genitalia, it seems

to represent a distinct line or separate complex.

BIONOMICS. Culex inculus is a rare species as it has been collected only

twice, once from the type-locality in Malaysia and once from Cambodia. In

Selangor, Malaysia, the immatures of inculus were collected from pools ina

nipa palm swamp in association with specimens of reidi (Colless 1965: 281).

In Cambodia, the single male was incidentally collected by sweeping with a net

in a forest along the sea coast. Nothing is known about the adult biology and its

medical importance.

quadripalpis complex

11. CULEX (LOPHOCERAOMYIA) QUADRIPALPIS (EDWARDS)

(Figs. 14, 15)

Lophoceratomyia sylvestris Leicester 1908: 125 (“, 2; non Theobald 1901).

Lophoceratomyia quadripalpis Edwards 1914a: 80 (&).

Culex (Lophoceratomyia) quadripalpis Edwards 1922a: 280 (key); Edwards

1922b: 473 (synopsis); Edwards 1932: 197 (taxonomy).

Culex (Lophoceraomyia) quadripalpis (Edwards), Mattingly 1949: 227 (L*);

Stone, Knight and Starcke 1959: 235 (catalog); Colless 1965: 285 (o'*, 9,

Et); Bram 2967a:.65 (¢*,- 2. 1b).

Culex (Lophoceratomyia) roubaudi Borel 1926: 112 (o*, 2, L*); Borel 1930:

362 (o*, 9, L*); Edwards 1932: 197 (synonymy).

Culex (Lophoceratomyia) pacheécoi Baisas 1935: 171 (o*, 2); Bohart 1945: 76

(“*, key); Stone, Knight and Starke 1959: 235 (catalog); Delfinado 1966:

112 (**, P, L). NEW SYNONYMY.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 45

Culex (Lophoceratomyia) barkerii (Theobald 1907), in part of Edwards 1913:

236 (taxonomy); Edwards 1917: 227 (taxonomy).

FEMALE. Wing: 3.0-4.0 mm (average 3.5mm). Forefemur: 1.7 mm.

Proboscis: 2.0 mm. Medium or large sized species, color dark brownish to

blackish; generally conforms to the Fraudatrix Subgroup. Head, Vertex with

a narrow band of broad pale scales forming distinct ocular line along upper eye

margin; narrow decumbent scales numerous, predominantly dark, occupying

an extensive area in center; erect scales coarse and blackish; lateral patch of

broad scales whitish, very distinct. Palpus about 0.2 of proboscis length.

Proboscis with 4 labial basal setae, 2 lateral ones strong, 0.5 of palpal length,

2 median ones shorter. Cibarial Armature (Fig. 14). Cibarial teeth numerous,

60 or more, all of which are narrow, subequal, apically blunt or pointed, ina

close-set row. Thorax. Mesonotal integument dark brown; mesonotal scales

narrow, dense and entirely blackish. Ppn without scales or setae cephalad of

posterior bristles. Pleuron same color as mesonotum, scales practically

absent; Ppl with 3 bristles and 8-10 weak setae; 1 lower mep bristle present,

sometimes 2. Legs. Dark, without marked coloration; anterior surface of

hindfemur with a broad pale stripe extending from base to apex. Wing. Scales

dark, narrow and relatively dense on all veins; plume scales on veins Ro, Rg

rather narrow, linear. Abdomen. Terga entirely dark; basolateral pale spots

present or sometimes absent; sterna yellowish scaled.

MALE (Fig. 14). In general as described for the female. Head, Ocular

line of vertex wider, narrow decumbent scales paler. Palpus. Length varying

from as long as to longer than proboscis by the full length of segment 9; basal

fingerlike process rather thick and relatively long, reaching slightly beyond

apex of segment 1; segment 3 with 3,4 ventrolateral bristles subapically; seg-

ments 3 and 4 weakly to moderately plumose. Proboscis. Dorsal upright

setae well developed, numerous, forming a dense double row in distal 0.75

of total length; most distal setae longest, 2,3 times as long as labial width at

point of attachment; labial basal setae 12. Antenna. Flagellar whorls densely

long plumose, modified tuft of F-5 variable, usually very distinct, with some

broad dark scales and lanceolate setae; 3,4 dorsalmost scales long, blunt

tipped or abruptly pointed, as long as the combined length of the next 4 flagel-

lomeres, followed laterally by 3 acute lanceolate scales of decreasing length

and 1 dark, longer bristlelike seta; F-6, 7 with usual crumpled tufts of setae

as described for the subgroup; F-8 with relatively thick J-hooked tufts of 6-8

fused setae; F-9 with 4, 5 yellowish bladelike scales; F-10 with 3,4 long bristle-

like setae.

MALE GENITALIA (Fig. 14). Segment IX. Tergal lobe small, with 3,4

short, weak setae; sternum usually bare, sometimes with a few setae towards

caudal margin. Basimere, Normal, conical; with a prominent row of 6-8 strong

submarginal setae on inner tergal surface; marginal setae 6,7, all weak, short,

widely spaced on tergomesal margin. Subapical Lobe, Rodlike setae a-c of

proximal part stout, subequal, smoothly curved, with hooked or truncate api-

ces; several fine hairlike setae present at bases of rods on mesal surface; dis-

tal part with 1 hairlike and 4 bladelike setae in group d-/; leaflet £5 distally

broad, spatulate; leaflet g; well developed, broad, apex blunt or acuminate;

seta 4 strong and long. Distimere. Normal, rather thick, gently or strongly

curved in middle; subapical claw short, distally dilated; 1 ventral and 1 dorsal

tiny seta present at 0.75 from base; dorsal subapical surface very lightly

serrated. Phallosome. Apical dorsal beaklike process of labial plate moder-

ately thick and long, posterior margin with or without minute spicules. Pvoc-

46 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

tiger, Crown small, with a distinct tuft of few flat and several spinelike

spicules; apex of paraproct with a short, broadly rounded, apical sternal lobe

and a poorly developed external lobe; cercal setae usually 3,4, sometimes 2.

PUPA (Fig. 14). Abdomen: 2.5mm. Paddle: 0.70mm. Trumpet: 0.78

mm; index about 12. Cream-colored or yellowish white. Trumpet. Relatively

long, uniformly cylindrical. Cephalothorax, Seta 1-C usually 4 branched (3-4);

8,9-C double. Metanotum. Setae 10,11-C double. Abdomen. Seta 6-III-VI

usually 4,5 branched (3-6); 5-IV usually 7 branched (7-8); 5-V usually 5

branched (4-7); 5-VI double; 9-VII usually 4 branched (4-5); 4-VIII usually

double (2-3). Paddle. Entirely pale, without infuscate areas; midrib weak,

moderately pigmented or same color as outer and inner parts of paddle.

LARVA (Fig. 15). Head: 0.76 mm. Siphon: 1.8-2.0 mm; index 9-10.

Saddle: 0.30 mm; siphon/saddle ratio 5-6. In general as described for the

group with the following diagnostic features. Head, Yellowish white; seta

4-C 0.5 or more of the distance between bases of the pair, single or bifid; 5,

6-C double; 7-C usually 6, 7 branched (5-8), all branches narrow and weakly

pectinate; 14-C double. Thorax, Spiculation absent; seta 3-P usually single,

rarely double; 4-P usually single (1-2); 14-P single or double. Abdomen.

Seta 6-I, II usually triple (3-4); 7-I double; 1-III-VI usually 4, 5 branched (3-7);

6-IV-VI usually 4 branched (3-6). Comb scales 30-40, all subequal, with api-

cal fringe of evenly fine spicules. Saddle yellowish white, seta 2-X with 1 short

and 1 long branch; ventral brush (4-X) with 6 pairs of setae; anal gills as long

as saddle. Siphon. Slender, long, usually with median dark ring, pecten teeth

11-15, distal tooth with 2,3 strong basal denticles and 4-6 fine, graded denticles

distally; subventral tufts 4 pairs (total 8), 2-4 branched each, as long as or

slightly longer than siphonal width at points of attachment; 2-5 short and weak;

median caudal filament of spiracular apparatus very well developed.

TYPE-DATA. (1) Lophoceratomyia sylvestris Leicester; Lectotype “*

with slide of genitalia, Kuala Lumpur, (Selangor), Malaya | MALAYSIA], pool

in hillside stream, Pahang Rd. jungle November 23, 1903, G. F. Leicester

(BMNH; selection of Colless 1965: 285). (2) Lophoceratomyia quadripalpis

Edwards; Holotype o* with slide of antenna and genitalia, Sarawak, N. Borneo

[ MALAYSIA], 1908, J. Hewitt (BMNH). (3) Culex (Lophoceratomyia) rou-

baudi Borel; Type non-existent, Terres Rouges, Cochin-China, [ VIETNAM].

(4) Culex (Lophoceratomyia) pachecoi Baisas 1935; Lectotype o* (F-150),

marked as paratype, with slide of antenna and genitalia, College of Agriculture,

Los Banos, Laguna, Luzon, PHILIPPINES, larva breeds in semi-stagnant

edges of forest creeks, February 2, 1935, F. E. Baisas (USNM; present

selection).

DISTRIBUTION. Sri Lanka, Thailand, Malaysia and the Philippines; also

reported from Vietnam, Singapore, Indonesia (Sulawesi). 160 specimens

examined: 71c', 229, 67 L; 54 with associated immature skins (19 p, 35 lp).

SRI LANKA. Western Province: Colombo District, Labugama; 1c, 3.

THAILAND. Chiang Mai: Doi Sutep; Khon Kaen: Phuweang; Nakhon Rat-

chasima: Khao Yai; Ban Khanong, Phra Nua; Khlong Pai; Kabinburi Pak Thong

Chai; Khao Saladi; Nakhon Nayok: Khao Yai; Phangnga: Nam Tai; Songkhla;

Satun; Narathiwat: Khau Lau; 23, 169, 65 L, 19 p, 15 lp.

MALAYSIA. Peninsular Malaysia - Selangor: Ulu Gombak; Pahang:

Gunong Benom; Sungai Temau; Perak: Chior F. R.; Kedah: Sintok F. R. ; 8c",

62, 1 L, 2p, 11p. Malaysia (Borneo) - Sarawak: Kuching; Sabah: Tawau, N.

Kalabakan; Sandakan Bay, Sepilak F. R.; 17°.

PHILIPPINES. Luzon: Laguna, Los Banos, College of Agriculture;

Malawin Creek, Mt. Makiling; Palawan: Puerto Princesa; Iwahig; Cabayuyan;

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 47

Ludlow Barracks; 220.79; 1°, °O0"Ip.

Additional records from the literature. VIETNAM (as roubaudi, Borel

1930: 362); SINGAPORE (Colless 1965: 286); INDONESIA: Sulawesi (Celebes)

(Brug and Bonne-Wepster 1947: 186).

TAXONOMIC DISCUSSION. Iam following Edwards (1932) and subsequent

authors (Stone, Knight and Starcke 1959, Colless 1965, Bram 1967a) in using

the name "quadripalpis" for this species in spite of the fact that there are

some dubious points about its validity as pointed out by Colless (1965: 286).

Although the earlier name "sylvestris" Leicester was apparently the one to be

adopted after it had been removed from the synonymy with Culex sylvestris

Theobald which is an Aedes, there was, however, no indication that "sylvestris"

was ever resurrected by Edwards (1922a, 1922b, 1932). In the earlier paper,

Edwards (1917: 237) had considered both sylvestris and quadripalpis to be syn-

onymous with barkervii Theobald, but later in his world catalog (Edwards 1932:

197), he revalidated quadripalpis and listed barkerii and sylvestris as its syn-

onyms. It seems that Edwards' final action was based purely on the grounds

that the status of quadripalpis was more stable and had been more frequently

used than the other 2 names. Edwards' treatment appears to be justified ex-

cept for the synonymy of barkerii Theobald which was clearly unwarranted as

indicated in the discussion by Colless (1965).

Culex quadripalpis is apparently one of the widespread Southeast Asian

species. It exhibits a great range of variation in the number and size of the

modified tufts of male antennal flagellomere 5, overlapping with the members

of the rubithovacis complex and the other members of the quadripalpis complex.

It can, however, be readily recognized in the male genitalia by the prominent

row of 6-8 strong submarginal setae of the basimere, the stout, moderately

long apical dorsal beaklike process of the phallosome and by the broadly rounded

apical sternal lobe of the paraproct of the proctiger. The female and pupa do

not appear to show any striking difference from other members of its own

complex, but can be distinguished from most members in the other complexes

of the Fraudatrix Subgroup by the numerous, fine cibarial teeth of the females

and by the combination of characters of the pupa as described above. The

larva is very distinct from other complexes, particularly in having seta 4-P

single, 14-P single or double and the relatively long siphon.

The newly proposed synonymy of the Philippine pachecoi Baisas with

quadripalpis above is only tentative, based partly on the similarity in several

details of the male genitalia and the immature stages. In comparing the males

of both forms, I found that pachecoi differs slightly from the typical quadvi-

palpis in usually having fewer, narrower, finely pointed scales in the modified

tuft of antennal flagellomere 5, more or less resembling some members of

rubithovacis complex, and in the weaker modified tufts of antennal flagello-

meres 6-8. However, there is also some overlap in all of these features. It

is quite probable that the patchecoi form is only subspecifically distinct and

that it probably represents a local or geographic segregate of a single wide-

spread species.

BIONOMICS. The breeding site of quadripalpis ranges from small ground

pools, rock pools along stream margins to large bodies of water such as

flooded grounds, seepage ponds and swamps. These habitats are usually under

heavy shade of tropical forest, containing numerous leaves, fallen trunks or

branches of trees. The adults were collected while resting on leaves or bushes

near the breeding sites and do not appear to be attracted to light. In Malaysia,

Colless (1965: 286) noted that adults occasionally attack man.

48 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

12. CULEX (LOPHOCERA OMYIA) ACULEATUS COLLESS

(Fig: 16)

Culex (Lophoceraomyia) aculeatus Colless 1965: 286 (*, 2, L*); Bram 1967a:

a0 1G, 2; L*).

FEMALE. Wing: 2.7mm. Forefemur: 1.2 mm. Proboscis: 1.55 mm.

Extremely similar to quadripalpis, differing in relatively smaller size and in

the following. Head. Erect scales of vertex finer. Cibarial Armature. As

figured for quadripalpis, cibarial bar narrower, with a close-set row of 35-40

teeth which are stronger and apically pointed. Thorax, Ppl with 1,2 bristles

and 4,5 weak setae. Wing. Plume scales on veins Ro, Rg finer; scales on

most veins rather sparse. Abdomen. Terga entirely dark, basolateral spots

completely absent.

MALE (Fig. 16). As described for female, differing from quadripalpis

particularly in the following features. Palpus. More slender; basal fingerlike

process of segment 1 apparently shorter, not exceeding apex of the segment;

segments 4 and 5 weakly plumose, with fewer bristles on lateral and mesal

surface. Pvoboscis. Dorsal upright setae in distal portion of labium fewer

and shorter; labial basal setae weaker, more or less hairlike, but not spine-

like. Antenna. Flagellar whorls moderately plumose; F-5 always with

prominent tuft of 6-8 dark scales, dorsalmost 3,4 scales broad, apically blunt,

usually slightly exceeding the combined length of the next 3 flagellomeres,

sometimes as long as the next 4 flagellomeres; followed laterally by 3, 4

lanceolate or apically pointed scales of decreasing length; 2,3 setae among

normal long hairs of inner whorl of F-5 dark, flattened, very strong and distinct;

F-8 with a strong J-hooked tuft of 6,8 fused setae; F-9 with 3,4 long, yellowish,

bladelike scales; F-10 with 2,3 bristlelike setae.

MALE GENITALIA (Fig. 16). Rather slender and small, differing from

quadripalpis strikingly in having only 3 submarginal setae on the inner tergal

surface of basimere and in the following details. Segment IX. Tergal lobe

very poorly developed, bearing 2 tiny setae; sternum with or without a row of

strong setae near caudal margin. Basimere. More slender; inner tergal sur-

face with 3 moderately strong submarginal setae in a row on its inner tergal

surface; sometimes with 1 other weaker seta in addition. Subapical Lobe,

Leaflets 21, £9 of distal part broad, foliate, with acute apices; g9 with apex

drawn out into a filament. Distimere. Slender and gently curved in apical

0.5; subapical claw short. Phallosome. Very distinctive; apical dorsal beak-

like process of lateral plate very long, thin, distally tapering into a sharp

point. Pyroctiger, Lateral spicules of crown coarser; apex of paraproct with

a distinct lateral lobe which is well sclerotized, elongate, somewhat tongue-

like; cercal setae 3.

PUPA. Abdomen: 2.3mm. Paddle: 0.62 mm. Trumpet: 0.66 mm;

index 10. Essentially as figured and described for quadripalpis, differing

slightly in the following. Cephalothorax, Seta 1-C 4,5 branched; 5-C 4-6

branched. Abdomen. Seta 5-IV 5-6 branched; 9-VII usually triple (3-4).

LARVA. Head: 0.70 mm. Siphon: 1.4-1.6 mm; index 8-10. Saddle:

0.3 mm; siphon/saddle 5. As figured and described for quadripalpis, differ-

ing from it particularly in the following. Head, Seta 14-C usually triple or

4,5 branched. Thorax. Seta 3-P usually double, rarely single; 14-P always

single. Abdomen. Seta 6-I always triple. Saddle seta 2-X with 2,3 short

branches and 1 long branch. Siphon. Relatively shorter and with lower index

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 49

and siphon/saddle ratio; pigmentation uniformly yellowish in proximal and dis-

tal portion, median dark ring usually absent; subventral tufts weaker and

shorter, double or triple, as long as siphonal width at point of attachment.

TYPE-DATA. Holotype “ with associated pupal and larval skins and slide

of antenna and genitalia, Kg. Gajah F. R. Perak, Malaya | MALAYSIA], Nov-

ember 13, 1958, D. H. Colless (ANIC).

DISTRIBUTION. Thailand and Malaysia, not known elsewhere. 116 speci-

mens examined: 530°, 2792, 36 L; 33 with associated immature skins (19 P,

14 lp).

THAILAND. Chiang Mai: DoiSutep; Lampang: Ngao; Tak: Doi Sam Sao;

Khao Salak Phra; Huai Lan Lang; Nakhon Ratchasima: Khao Yai; Prachin Buri:

Ban Tung Fack; Chanthaburi: Ban Bo Phu Pit; Ban Chak Yai; Khao Sai Dao;

Ranong: Khao Hin Chong; Trang: Muang; Phangnga: Khao Soong; Phuket:

Khao Prathin; Bang Khian; Prachaup Khiri Khan: Klong Hin Chong; Songkhla:

Ratta Poom; Yala; Yala Bong; Stun; Narathiwat; Khau Lau; 36, 119, 36 L,

18 p, 8 lp.

MALAYSIA. Peninsular Malaysia - Selangor: Ulu Gombak; Ampang F. R.;

Pahang: Gunong Benom; Perak: Chior F. R.; 13o, 162, 1p, 6 lp. Malaysia -

Sabah: Tawau; 4.

TAXONOMIC DISCUSSION. Culex aculeatus is strongly differentiated from

the rest of the quadripalpis complex especially by the long, very slender apical

dorsal beaklike process of the male phallosome. The other male features which

are also very diagnostic of aculeatus are the presence of 2,3 dark flattened

setae on the inner whorls of antennal flagellomere 5 (fine, thin, hairlike in all

others of the complex) and the elongate, tongue-like apical lobe of the para-

proct of the proctiger. The female cannot be separated from quadripalpis with

certainty on the external characters, but is obviously different from it in hav-

ing fewer cibarial teeth as described and figured here. Both the larva and pupa

of aculeatus exhibit considerable overlap with those of quadripalpis from which

it can be clearly separated by using the combinations of the usual characters

of the setae and other general features as indicated in the above description.

There is apparently little variation in the diagnostic characters of aculeatus

and all of the specimens which have been identified as this species essentially

conform to a single type. Although aculeatus is known only from Thailand and

Malaysia, there is a possibility that it may be widespread, extending into other

areas of Southeast Asia.

BIONOMICS. The immatures of aculeatus have been frequently collected

from pools at the edges of streams and occasionally from swamps, marshy

depressions and rock pools. These breeding sites which are under heavy

shade usually contain aquatic vegetation, decayed leaves and fallen branches

of trees and are at a high elevation. At Khao Yai, Nakhon Ratchasima, Thai-

land, specimens were collected at an elevation of 1,200 m. A number of the

adults were collected resting on vegetation and from sweeping with a net among

bushes near the breeding sites. The females are not known to bite man.

13. CULEX (LOPHOCERAOMYIA) PARACULEATUS NEW SPECIES

(Fig. 16) :

Culex (Lophoceratomyia) fraudatrix of Baisas 1935: 173 (c'*, 9).

FEMALE. Unknown.

MALE (Fig. 16). Exceedingly similar to aculeatus in the details of antenna,

50 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

palpus, proboscis and general facies: differing slightly in the following. fPal-

pus. Basal fingerlike processes thicker and longer, nearly reaching the apex

of segment 1. Pvroboscis. Distal 0.75 of labium with more numerous dorsal

upright setae. Antenna. Setae among normal long hairs of inner whorl of F-5

all fine, hairlike; modified tuft with 4-6 dark scales, 2,3 dorsalmost scales

longest and apically blunt, slightly longer than the combined length of the next

3 flagellomeres, 2-3 lateral ones shorter, lanceolate; F-7 with weak, markedly

reduced tuft of curled setae; F-8 with more slender J-hooked tuft, consisting

of only 4 dark, fused setae.

MALE GENITALIA (Fig. 16). As figured, exceedingly similar to aculeatus ;

differing chiefly in the following. Subapical Lobe. Leaflet g9 narrow in basal

0.5, broad spatulate in apical 0.5, apex blunt. Phallosome. Apical dorsal

beaklike process much shorter, more or less resembling that of quadripalpis.

Proctiger, Apical sternal and lateral lobes of paraproct short and rounded;

crown with weaker and fewer spicules.

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype “ (B60/22) with slide of antenna and genitalia

(SEAMP 69/106), Tawau, (Sabah), North Borneo, [| MALAYSIA], February 1960,

D. H. Colless (BMNH). Paratype 1° (B60/22) with slide of head, palpus,

proboscis and genitalia (SEAMP 69/107), same data as holotype (USNM).

DISTRIBUTION. Malaysia (Sabah) and the Philippines (Leyte, Mindanao).

specimens examined: 4°.

MALAYSIA. Sabah; 2, as indicated in the type-data.

PHILIPPINES. Leyte: Diit River, Tacloban, 1c; Mindanao: Parang, 1°.

TAXONOMIC DISCUSSION. Culex paraculeatus is recognized as distinct

from aculeatus on the basis of the difference in the considerably shorter and

stronger apical beaklike process of the male phallosome and in the markedly

reduced tufts of antennal flagellomere 7. The 2 males from the Philippines un-

doubtedly belong to this species since they agree well with the males in the

type-series in the genitalic characters. At the type-locality in Tawau, Sabah,

paraculeatus was found to be sympatric with aculeatus without intergradation in

their diagnostic features. The Philippine specimens described by Baisas (1935:

173) as fraudatrix apparently belong to this species judging by his figures of the

male antenna and genitalia.

BIONOMICS. The males of paraculeatus came from collecting with a

sweep net among leaves of plants in a tropical rain forest.

14. CULEX (LOPHOCERA OMYIA) AESTIVUS NEW SPECIES

(Fig. 17)

FEMALE. Unknown.

MALE (Fig. 17). Exceedingly similar to aculeatus and paraculeatus in most

features of the antenna, palpus, proboscis and general facies; differing from

both particularly in the following. Antenna. Modified tuft of F-5 with all

broad, dark scales abruptly ended into a fine point; inner whorl of normal long

hairs of F-9 as in paraculeatus; F-8 with a strong J-hooked tuft, consisting

of 8 dark, fused setae.

MALE GENITALIA (Fig. 17). As figured, differing from aculeatus and

paraculeatus particularly in the following. Basimere, Inner tergal surface

with only 1 submarginal seta. Subapical Lobe. Leaflet g, of distal part absent

or not developed, go broad, foliate, apically drawn out into a fine point, as

in aculeatus, Phallosome, Apical dorsal beaklike process short and stout,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region ol

as in pavaculeatus but apparently heavier. Proctiger. Crown smaller, with

fewer and finer spinelike spicules; paraproct with distinct lateral apical lobe

which is longer than Pavaculeatus but shorter than aculeatus.

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype “ with slide of antenna and genitalia (SEAMP 69/

105), Long Tebangan, Savawak, MALAYSIA, June-July, 1953, D. H. Colless

(BMNH).

DISTRIBUTION. Known only from the type-locality. 10° examined, as

indicated in the type-data.

TAXONOMIC DISCUSSION. Culex aestivus is somewhat intermediate be-

tween aculeatus and paraculeatus but is distinct from both in the presence of

1 submarginal seta on the basimere and the absence of leaflet g; in the distal

part of the subapical lobe of the male genitalia. Although aestivus is known

from a single male, it appears to be distinct. There is no indication that its

diagnostic features specified above fall into the range of variation of aculeatus

or paraculeatus.

BIONOMICS. The single male of aestivus was collected in the field while

resting among shrubs.

15. CULEX (LOPHOCERAOMYIA) REIDI COLLESS

(Fig. 18)

Culex (Lophoceraomyia) reidi Colless 1965: 279 (o*, 9, L*); Delfinado 1966:

113 (o*)}) Bram 1967a: 66 (o*, 9, L*).

Culex (Lophocevatomyia) quadripalpis of Edwards and Given 1928: 352 (L,

misidentification).

FEMALE. Wing: 2.7mm. Forefemur: 1.3 mm. Proboscis: 1.7 mm.

In general as described for the complex and subgroup, not readily distinguished

from other species. Head, Vertex largely covered with dark, broad decum-

bent scales; not forming a distinct ocular line; narrow decumbent scales re-

stricted to occiput; erect scales dark or blackish, lateral patch of broad scales

predominantly dark, not distinct. Palpus relatively short, about 0.15 of pro-

boscis length. Proboscis with 2 labial basal setae, both of which are dark,

about 0.5 of palpal length. Cibarial Armature. Not studied. Thorax, Meso-

notal integument and scales dark brown. Pleuron slightly paler than mesonotum;

1 lower mep bristle present. Legs. Without marked coloration; anterior sur-

face of hindfemur with a narrow longitudinal pale stripe from base to near apex.

Wing. Scales on all wing veins moderately dense, plume scales on veins Ro,

Rg narrow, clavate. Abdomen. Terga entirely dark; basolateral pale spots

usually absent, sometimes present on terga II-VII; sterna usually pale yellow-

ish scaled, sometimes dusky.

MALE (Fig. 18). In general similar to female; essentially as described by

Colless (1965: 279) and Bram (1967a: 66). Palpus. Exceeding proboscis by

slightly more than the full length of segment 5; basal fingerlike processes

strong and relatively long, as long as the distal part of segment 1; segment 3

with 1 strong and 2 weak bristles in apical 0.1; segments 4, 5 weakly to moder-

ately plumose. Proboscis. Apical 0.5 of labium with distinct double rows of

dorsal upright setae, distal setae longest, 2 times as long as labial width at

point of attachment. Antenna, Flagellar whorls strongly long plumose; modified

tuft of F-5 distinct, consists of 4-6 narrow or broad scales, all of which are

tapered into a fine point, 2,3 dorsalmost scales longest, as long as the combined

D2 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

length of the next 3 flagellomeres, followed laterally by 2,3 shorter scales

and 1 narrow scale of increasing length; modified tufts of F-6 to F-10 normal,

as described for the subgroup and essentially similar to guadripalpis and acu-

leatus.

MALE GENITALIA (Fig. 18). As figured, as described by Colless (1965:

279) and as in the following description. Segment IX. Tergal lobe well de-

veloped, bearing 5-7 relatively strong setae. Basimere. Submarginal setae

usually 4, sometimes 3 or 5, all apparently flattened, strong, moderately

long and heavily bent distad of middle. Subapical Lobe, One of the 3 rods in

group a@-c broadly expanded and strongly angulate in apical portion, other 2

rods slender, tapered into hooked apices; distal part with 5 narrow blades and

1 hairlike seta in group d-f/; leaflet g. club-shaped; 8; broad, acuminate; basal

seta (h) weak. Distimere. Dorsal subapical surface with a distinct crest of

fine spicules, extending from apex to middle of curvature; subapical claw short.

_Phallosome. Apical dorsal beaklike process of labial plate slender and short.

Proctigery, Crown small, composed largely of fine, pointed spicules; apex of

paraproct adjacent to crown broadly rounded, apparently without distinct exter-

nal (or lateral) and sternal lobes; cercal setae 3, 4 in number.

PUPA. Abdomen: 2.3mm. Paddle: 0.60 mm. Trumpet: 0.52 mm;

index 8. As figured for quadripalpis (Fig. 14); pigmentation of cephalothorax

and abdomen light, yellowish brown, without definite pattern of dark and light

areas, trumpet darker; the following setae are diagnostic. Cephalothorax.

Seta 1,3-C double; 8,9-C double. Metanotum., 10, 11 and 12-C double. Abdo-

men, Seta 1-III, IV 6 branched; 1-V, VI 3,4 branched (2-4); 1-VIII 1-3 branched;

o-IV 2-4 branched; 5-V-VI double; 6-III-VI usually double, sometimes single.

Paddle, Midrib strong and dark, inner and outer parts pale, without darkened

areas.

LARVA (Fig. 18). Head: 0.70 mm. Siphon: 1.2 mm; index 9. Saddle:

0.26 mm; siphon/saddle ratio 4.5. Essentially conforms to the quadripalpis

complex in the single seta 4-P, distinctive in the following. Head. Seta 4-C

very short, about 1.5 of the distance between bases of the pair, placed very

close to seta 6-C; usually distally forked into 2,3 branches; 5, 6-C double,

subequal; 7-C 6,7 branched, weak; 14-C double. Thorax. Seta 3-P single;

4-P single; 7-P usually double (2-3); 8-P double; 14-P single. Abdomen,

Seta 6-I, II triple; 7-I single; 6-III-VI usually triple (3-4); 1-IIJ-VI double or

triple. Comb scales about 40, all of which are subequal and with apical fringes

of fine spicules. Saddle seta 2-X with 2 short branches and 1 long branch;

ventral brush with 6 pairs of setae; anal gills as long as or shorter than saddle.

Siphon. Moderately long, uniformly yellowish white or sometimes with median

dark band; pecten teeth 11-12, distal teeth with 2 strong basal and 6-10 fine

apical denticles; subventral tufts 4 pairs, 3-6 branched each, 3 proximal pairs

subequal, slightly longer than siphonal width at points of attachment, most

distal pairs shorter; median caudal filament of spiracular apparatus well de-

veloped.

TYPE-DATA. Holotype ©“ with associated larval and pupal skins and slide

of antenna and genitalia (CT544), SINGAPORE, reared from a single egg-raft,

December 18, 1956, D. H. Colless (ANIC).

DISTRIBUTION. Thailand, Malaysia, Singapore and the Philippines; 22

specimens examined: 60, 2°, 14 L; 5 with associated immature skins (1 p,

5 Ip).

THAILAND. Bangkok: Pathumwan; Bang Khun Non; Bang Mod; Thonburi:

Bangkok Noi; Chanthaburi: Ban Wak; 3c, 12, 6 L, 1p, 2 lp.

MALAYSIA. Peninsular Malaysia - Selangor: Rantau Panjang; Perak:

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region o3

Trong; 1%, 12, 6 L, 2 lp; Malaysia - Sabah; 1~.

SINGAPORE. 2 L.

PHILIPPINES. Palawan: Uring Uring, Brooke's Point; 1“ (genitalia only).

TAXONOMIC DISCUSSION. Culex reidi is differentiated from the other

members of the quadripalpis complex in the adults of both sexes by the pre-

dominantly broad and dark decumbent scales of the vertex of head; in the male

by (1) the relatively long basal fingerlike processes of palpus and (2) the

pointed or acute scales of the modified tuft of antennal flagellomere 5; in the

male genitalia by (1) the presence of a row of 4 markedly flattened submarginal

setae; (2) one of the 3 proximal rods of the subapical lobe broadly expanded and

angulate in apical portion; (3) the short and slender apical beaklike process of

the phallosome and (4) apical lobe of paraproct of the proctiger broadly rounded;

in the pupa by (1) the dark and strong midrib of the paddle and (2) setae 6-III- VI

usually double; in the larva by (1) seta 7-P usually double and (2) seta 7-I

single.

The affinity of veidi appears to be intermediate between the other members

of the guadripalpis complex and inculus. It is very similar to imculus in the

general facies of the adults, pupal and larval chaetotaxy but is similar to most

members of the quadripalpis complex in several details of the male genitalia.

BIONOMICS. The breeding sites of veidi are fresh water ground pools in

a nipa palm swamp near to or in the vicinity of the seacoast. It appears to be

rather rare. Nothing is known about adult biology.

variatus complex

16. CULEX (LOPHOCERAOMYIA) VARIATUS (LEICESTER)

(Figs. 19, 20)

Lophoceratomyia variata Leicester 1908: 121 (“, ).

Lophoceratomyia fraudatrix Theobald 1905b, Edwards 1913: 235 (as senior

homonym of L.variata Leicester).

Culex (Lophoceratomyia) fraudatrix Theobald of Barraud 1924: 41 (o*);

Edwards and Given 1928: 351 (L); Edwards 1932: 197 (taxonomy);

Barraud 1934: 368 (o*, 9°, L); Brug and Bonne-Wepster 1947: 186

(distribution, in part); Chu 1957: 151 (o*, distribution).

Culex (Lophoceraomyia) fraudatrix Theobald, Stone, Knight and Starcke 1959:

233 (catalog).

Culex (Lophoceraomyia) variatus (Leicester), Colless 1965: 270 (o*, 2, L*);

Bram 1967a: 72 (o*, 2, L*); Stone 1967: 214 (catalog).

FEMALE. Wing: 2.8-3.3 mm (average 3.0 mm). Forefemur: 1.2-1.5

mm (average 1.4 mm). Proboscis: 1.5-1.9 mm (average 1.75 mm). Medi-

um to large sized in the subgenus; generally conforms to the description of

the Fvaudatrix Group. Head, Decumbent scales of vertex largely broad and

dark, sometimes partially pale along upper eye margin, forming a distinct

ocular line; narrow decumbent scales restricted to dorsal midline, extending

anteriorly to frons and posteriorly to occiput; lateral patch of broad scales

pale brown or dingy white. Palpus 0.2 of proboscis length. Proboscis with

2 labial basal setae which are 0.5 of palpal length. Cibarial Armature (Fig.

19). Cibarial bar with a concave row of 40-50 narrow fine teeth which are

equally long and apically pointed or blunt. Thorax. Mesonotal integument

pale brown to dark chestnut brown; scales very narrow, moderately dense and

04 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

dark except for a few pale ones on extreme anterior promontory. Ppn with

some narrow fine scales or setae cephalad of 4,5 posterior bristles. Pleuron

same color as mesonotum; ppl with 3,4 bristles and a few weak setae; 1 lower

mep bristle present. Legs. Anterior surface of hindfemur with a longitudinal

pale stripe extending from base to 0.5-1.0 of whole length. Wing. Scales

moderately dense; plume scales on R,, R, relatively broad clavate. Abdomen.

Terga entirely dark; basolateral pale spots present or absent; sterna pale

yellowish.

MALE (Fig. 19). Palpus, proboscis and antenna as described and figured

by Colless (1965: 270); sexual dimorphism well developed, particularly in the

presence of numerous fine setae, forming a dense patch on anterior part of

fossa of the mesonotum. Palpus. Relatively long, exceeding proboscis by

nearly the combined length of segments 4 and 5; basal fingerlike processes

distinct, about 0.5 of the distal part of segment 1; segment 3 with a ventral

row of short tiny setae which are dense in basal 0.5, sparse in apical 0.5;

ventral lateral tuft of setae not developed or absent; apical 0.2 with 5-10 dark

bristles; segments 4 and 5 strongly plumose. Pyvoboscis. Dorsal upright

setae present, extending from basal 0.25 to apex of labium, their length 3

times as long as labial width at point of insertion. Antenna, F-5 with a large

prominent fan-shaped tuft of 7-10 dark broad apically blunt scales which are

as long as the combined length of the next 4 flagellomeres, followed laterally

and ventrally by 10-12 distally pale flattened setae of subequal length; F-6, 7

with dark crumpled tufts of curled setae as described for cinctellus and other

preceding species; mesal tuft of F-7 rather thick, with 6,7 subapically twisted

setae; J-hooked tuft of F-8 with characteristic twist or kink subapically; F-9

with 5 yellowish bladelike scales; F-10 with 2,3 strong bristlelike setae.

MALE GENITALIA (Fig. 19). Essentially as described and figured by

Colless (1965: 271) and Bram (1967a: 74). Segment IX, Tergal lobe small,

bearing 4-6 weak setae; sternum without any scales or setae towards caudal

margin. Basimere. Normal, with a row of 3 moderately long submarginal

setae on slight convex inner tergal surface; 4,5 marginal setae strong, 0. 50-

0.75 of submarginals. Subapical Lobe. Rodlike setae a-c stout, subequal,

smoothly curved; setae d-f 5 in number; leaflet g5 broad club-shaped, apex

pointed or blunt; leaflet g, rather narrow, short, with acute apex; seta “ mod-

erately long. Distimere. Normal; subapical dorsal surface lightly annulated;

subapical claw short, more or less dilated apically; ventral tiny seta longer

than dorsal. Phallosome. Apical beaklike process slender, long, sustaining

about 90 degrees with main stem and distally tapered into a sharp point. Proc-

tiger. Crown small, consisting of 3,4 coarse spicules laterally and several

spinelike spicules internally; paraproct with a small apical lateral lobe and a

broad round apical sternal lobe; cercal setae 3.

PUPA (Fig. 19). Abdomen: 2.5mm. Paddle: 0.65mm. Trumpet: 0. 72-

0.75; index 11-12. Pigmentation variable, usually yellowish white or some-

times with indefinite brownish areas on cephalothorax and metanotum. Trum-

pet. Relatively long, darker than underlying integument, meatus slightly

swollen in middle, narrow towards basal and apical portions, pinna narrow,

with slit extended to meatus. Cephalothorax., Seta 1-C usually triple (2-4);

3-C triple; 5-C usually 4 branched; 8-C single or double; 9-C usually double

(2-3). Metanotum., Setae 10,11-C double. Abdomen. Seta 5-IV 4-7 branched;

5- V usually 4 branched (3-5); 5-VI double; 6-III-VI usually 4 branched (3-4);

9-VII usually triple (2-4); 4-VIII usually triple (2-3); 9-VIII as long as 9-VII,

usually 7 branched (5-8). Paddle, Broad, inner part lightly darkened, outer

part pale except for darkened area along midrib, midrib strong and usually

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region ayy)

very dark.

LARVA (Fig. 20). Head: 0.72 mm. Siphon: 1.6-1.9 mm; index 7-10.

Saddle: 0.36 mm; siphon/saddle ratio 5. Essentially conforms to the group,

with the following diagnostic features. Head, Seta 4-C distally forked into

2-4 branches, its length about 0.5 of the distance between bases of the pair;

9, 6-C double; 10-C 3, 4 branched; 14-C double. Thorax. Lightly to moderately

spiculated, spicules distinct under 10X objective, seta 3-P usually double (1-2);

4-P double; 7-P triple; 8-P double; 14-P single. Abdomen. Unspiculated; seta

7-I double; 6-III 3,4 branched; 6-IV-VI usually 4 branched (4-5). Comb scales

normal, 40-50. Saddle lightly spiculated on posterior caudal margin; seta 2-X

3,4 branched; anal gills as long as saddle length. Siphon. Slender, long; me-

dian dark band present or absent; pecten teeth 9-14, distal teeth with about

10 graded denticles; subventral tufts weak, rather inconspicuous, 4 pairs,

each tuft double or triple, as long as siphonal width at point of attachment; 2-S

fine, hairlike; median caudal filament of spiracular apparatus well developed.

TYPE-DATA. Lectotype “*, Kuala Lumpur, [ Selangor], Malaya | MALAY-

SIA], marshy ground, Circular Road, October 19, 1903, G. F. Leicester (BM

NH; selection of Colless 1965: 270).

DISTRIBUTION. Thailand, Vietnam and Malaysia; also reported from

India and Hainan (China), but not known from the Philippines; 151 specimens

examined: 880, 30°, 33 L; 48 with associated immature skins (17 p, 31 lp).

THAILAND. Bangkok; Thon Buri: Bangkok Noi; Trat: Koh Chang; Ban

Salak Phet; Chanthaburi; Ban Si Phraya; Ban Wak; Ban Sung Bon; Ranong:

Kraburi; Songkhla: Boripat Waterfall; Satun: Muang; Narathiwat: Ba Chau Vil-

lage; Ban Tone; 180, 162, 28 L, 11 p, 20 lp.

VIETNAM. Vung Tau; 1 L.

MALAYSIA. Peninsular Malaysia - Selangor: Rantau Penjang; Kuala

Lumpur; Pahang: Bentong Rd.; Tungku K. Lipis; Bt. Betong K. Lipis; Mela

K. Lipis; Perak: Kg. Kuala Dipang; Kuala Kangsar; Johore: Kota Tinggi; Per-

lis: Kg. Gunong; Kedah; Sintok, F. R.; Trengganu: Marang; 21“, 52, 4L, 3p,

4pl. Malaysia - Sabah: Lipasar Papar; N. Sandakan; Papar; Beaufort; Sanda-

' kan Bay; Kota Kinabalu; Tawau; 490", 92, 3p, 7 lp.

Additional records from the literature: INDIA: Assam; Andaman Is. (Bar-

raud 1934: 370, as fraudatrix); CHINA: Hainan Island (Chu 1957: 163, as

fraudatrix); INDONESIA: Sumatra; Java, Borneo (Brug and Bonne-Wepster

1947: 186, as fraudator, in part); MALAYSIA: Sarawak; SINGAPORE: Johore

(Colless 1965: 273).

TAXONOMIC DISCUSSION. The adults of variatus are variable in colora-

tion. In Malaysia, the adults are locally differentiated into the pale inland

form and the dark coastal form, as pointed out by Colless (1965: 273). The

pupa and larva are also variable in the color of the integument, more or less

correlated with their adults. However, as there are no significant correlated

differences in the male characters, including the genitalia and the diagnostic

pupal and larval chaetotaxy, it is probable that only a single species is involved

and that these variations are environmentally induced.

The female of varviatus is difficult to separate from most members of the

Fraudatrix Subgroup but the male, pupa and larva are distinctive and can be

readily recognized by the characters as given in the keys and as described

above. One of the most diagnostic features of the vaviatus male not described

before, is the presence of several fine hairlike setae laterad of the dorso-

central bristles on the anterior surface of the fossa. This character provides

a clear cut separation of the variatus males from all sympatric forms in the

variatus complex.

06 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Until it was resurrected by Colless (1965), vaviatus had been considered

by several authors as synonymous with fraudatrix Theobald from New Guinea.

The relationships between vaviatus and the typical fraudatrix as well as among

the members of these 2 complexes are extremely close and all may well be

assigned to the same complex as suggested by Sirivanakarn (1968: 155-6).

BIONOMICS. Culex variatus utilizes various types of ground pools as

breeding habitats, including: puddles, ditches, ponds, marshy depressions,

swamps under partial or heavy shade of forest or nipa palms. In Malaysia,

the larvae were reported to be most common in stagnant, usually fresh water,

partly shaded ground pools containing much organic matter in nipa palm swamp.

They were sometimes collected from palm axils, tins, cavities in fallen logs

or bamboos and crab holes (Colless 1965). The adults have been collected from

rearing the larvae, but several were also collected by light traps and by sweep-

ing with nets. The dark adult form readily attacks man by day under forst

cover in certain areas but its normal hosts are probably birds or forest mam-

mals (Colless 1965).

17. CULEX (LOPHOCERAOMYIA) JOSEPHINAE BAISAS

(Fig. 21)

Culex (Lophoceratomyia) josephinae Baisas 1935: 172 (o*, &).

Culex (Lophoceraomyia) fraudatrix Theobald, Bohart 1945: 74 (taxonomy);

Stone, Knight and Starcke 1959: 233 (catalog).

Culex (Lophoceraomyia) josephinae Baisas, Delfinado 1966: 106 (c‘*).

FEMALE. Exceedingly similar to the pale form of variatus and is essen-

tially as described for the latter, differing slightly in having predominantly

pale broad decumbent scales and more numerous yellowish brown narrow de-

cumbent scales on the vertex of head. Cibarial Armature (Fig. 21). Essen-

tially similar to variatus from which it is indistinguishable.

MALE. Extremely similar to variatus in palpal, proboscis and antennal

characters, differing slightly in having less numerous fine hairlike setae on

anterior surface of fossa. |

MALE GENITALIA (Fig. 21). As figured and described by Baisas (1935:

172-3) and Delfinado (1966: 106-7); exceedingly similar to variatus in the de-

tails of basimere and distimere, differing from it particularly in the following.

Subapical Lobe, Leaflet 21 of distal part markedly reduced to a narrow, acute,

bladelike seta. Phallosome, Apical dorsal beaklike process very stout, short,

strongly bent downwards, sustaining an acute angle with the main stem, its

posterior apical margin provided with a number of minute spicules. Proctiger,

Paraproct with poorly developed apical lateral lobe; apical sternal lobe broad

and rounded; cercal setae usually 2 in number.

PUPA. As described and figured for variatus (Fig. 19) from which it is

indistinguishable.

LARVA. As described and figured for variatus (Fig. 20) from which it

differs slightly in the following features. Head, Antennal shaft with weaker

and finer spicules in basal 0.5. Thorax. Spiculation practically absent; seta

3-P always single. Siphon. Uniformly yellowish white, median dark ring ab-

sent.

TYPE-DATA. Types “, ¢ (lost), Del Carmen, Pampanga Province, Luzon,

PHILIPPINES, larva breeds in a clear vegetated river slew, March 6, 1930,

F. E. Baisas (Bureau of Health, Manila; the types were probably destroyed

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 57

during World War II).

DISTRIBUTION. Known only from the Philippines; 92 specimens examined:

~ T10", 202, 1 L; 2 with associated immature skins (2 lp).

PHILIPPINES. Mindoro: San Jose; Samar: Osmena; Leyte: San Rosa;

Tacloban; Diit, Lago Lago, Bay Bay; Dagamit, Mt. Lobi; Mindanao: Parang,

Ludlow Barracks; Kabakan; Zamboanga; Paranonco; Agusan, S. Francisco;

Surigao; Su-francisco: 71 2027 4415 2-1p, |

TAXONOMIC DISCUSSION. Culex josephinae is restricted to the Philippines

where it is one of the most common forms of Lophoceraomyia. It is probably

widespread throughout the area, including several small and large islands.

The phallosome and a few other characters of the male genitalia of josephinae

as given above are remarkably constant, providing a clear-cut separation from

those of variatus. It is possible that josephinae is only subspecifically distinct

from variatus, but, as there is no overlap in the diagnostic features of the

male genitalia between the 2 forms, it appears most practical to provisionally

recognize both as full species.

BIONOMICS. Culex josephinae is a typical ground pool breeder, as in

variatus, The recorded breeding sites include marshes and swamps associated

with stands of nipa palm and forests in coastal areas. Several adults were col-

lected while resting on vegetation by sweeping with a net as well as from mass

rearings in the field. Nothing is known about the adult biology and its medical

importance.

18. CULEX (LOPHOCERAOMYIA) CUBITATUS COLLESS

(Fig.-21)

Culex (Lophoceraomyia) cubitatus Colless 1965: 273 (o'*, 2, L*); Sirivanakarn

1973: 215 (distribution).

FEMALE. Wing: 3.0mm. Forefemur: 1.5mm. Proboscis: 1.9 mm.

Medium-sized, brownish species, essentially as described for variatus, dif-

fering from it slightly in the following characters. Head, Broad decumbent:

scales of vertex predominantly pale, forming a distinct ocular line; narrow

decumbent scales in the center more numerous. Czbarial Armature, As in

Fig. 21. Abdomen. Terga II-VII with lateral pale scaling, more or less con-

trast with dark scaled areas on dorsal surface.

MALE (Fig. 21). As described by Colless (1965: 273-4), differing from

variatus particularly in the following features. Pvroboscis, Dorsal upright

setae stronger and more numerous; lateral and ventral surface of labium with

several distinct setae (largely absent in variatus). Antenna, Modified tuft of

F-95 larger, composed of 14-18 broad, dark, blunt tipped scales in dorsal

eroup, followed laterally and ventrally by about 10 pale, hairlike setae which

are gradually increased in length; F-8 with smoothly curved J-hooked tuft;

F-9 with 5 darker and broader bladelike scales. Thorax, Anterior surface of

fossa without a dense patch of fine hairlike setae laterad of dorsocentral bristles.

MALE GENITALIA (Fig. 21). Extremely similar to variatus, differing from

it particularly in the following. Basimere. Inner tergal surface usually with a

row of 4 submarginal setae, sometimes 3 or 5. Subapical Lobe. One of the 3

proximal rodlike setae in group a-c strongly angulate, or with a characteristic

elbow beyond middle. Phallosome. Apical dorsal beaklike process stouter,

shorter, distally tapered into a blunt apex. Pyvroctigey. Crown larger, with

more numerous spicules; paraproct with short, broadly rounded apical lateral

08 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

and sternal lobes.

PUPA. Abdomen: 2.4mm. Paddle: 0.65mm. Trumpet: 0.63 mm;

index 10-12. As figured and as described for variatus, differing from it in the

following characters. Tvumpet, Rather thicker, shorter, darker and strongly

curved in middle. Cephalothorax, Seta 5-C 5 branched; 8-C double, much

stronger and longer than 9-C. Abdomen, Seta 5-IV 4,5 branched; 5-V, VI

double; 6-III-V usually double (1-3); 6-VI triple. Paddle, Midrib moderately

to strongly infuscated, inner and outer parts of paddle pale.

LARVA. Head: 0.70 mm. Siphon: 1.7-2.0 mm; index 9-11. Saddle:

0.35 mm; siphon/saddle ratio 4. As figured for variatus from which it differs

chiefly in the following features. Head, Seta 4-C shorter; 10-C single or

double. Antenna shaft darker, with stronger spicules. Thorax. Spiculation

absent or not developed; seta 14-P usually double, rarely single. Abdomen.

Seta 6-III usually triple. Comb scales apparently fewer, about 23-40 (average

30). Saddle seta 2-X double. Siphon. Rather longer, thinner, uniformly

yellowish white or brown; median dark ring absent.

TYPE-DATA. Holotype “ (57/48/1) with associated pupal and larval skins,

slides of antenna (CH 118) and genitalia (CT 474), Mandai Road, SINGAPORE,

December 29, 1957, D. H. Colless (ANIC).

DISTRIBUTION. Singapore, Malaysia, the Philippines (Mindanao), Ceram,

Indonesia and Papua New Guinea; 40 specimens examined: 170°, 59, 18 L; 6

with associated immature skins (2 p, 4 lp).

SINGAPORE. Mandai Road (type-locality); Mc Ritchie Reservoir; 3%

(including holotype), 12, 13 L, 3 lp.

MALAYSIA. Peninsular Malaysia - Pahang: Cameron Highlands; 4 L.

Sabah: ‘Tawau; Papar: Beaufort; 7¢, 2¥,.1'L; 2p; 1 lp.

PHILIPPINES. Palawan: Princesca; Mindanao: Agusan, S. Francisco;

selene e

INDONESIA. Ceram; 1%.

PAPUA NEW GUINEA. Sepik: Vanimo; 1c.

TAXONOMIC DISCUSSION. Culex cubitatus is differentiated from variatus

and other members of the variatus complex by the combination of the male

characters, larval and pupal chaetotaxy as described above. This species has

been seldom encountered but apparently exhibits a very broad range of distri-

bution, extending into the Philippines (Mindanao),Ceram and New Guinea (Siri-

vanakarn 1973: 215). In Singapore, Colless (1965: 274) noted that cubztatus is

sympatric with the dark form of variatus, but occupies a distinct ecological

niche, restricted to the inland areas, similar to the pale form of variatus in

Malaysia. Those facts are well supported by the recent collections in Malaysia.

BIONOMICS. In Singapore, cubitatus was found breeding in jungle ground

pools away from the tidal zone or coastal areas (Colless 1965). In Peninsular

Malaysia, the cubitatus larvae were collected from a pool at a high elevation

in the Cameron Highlands. The adults from Sabah, Malaysia were collected

by a sweep net and by rearing the pupae and larvae. The adults recorded else-

where all apparently came from general field catches by sweeping with nets.

Nothing is known about the biology or medical importance of the adults.

19. CULEX (LOPHOCERAOMYIA) GRACICORNIS NEW SPECIES

(Fig): 22)

Culex (Lophoceraomyia) sp. near cubitatus Colless 1965: 274 (o*, $).

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region o9

FEMALE. Unknown.

MALE (Fig. 22). As described and figured by Colless (1965: 274-5); es-

sentially similar to cubitatus in palpal, labial and antennal characters, differing

from it chiefly in the following. Palpus. Segments 4 and 5 more strongly plu-

mose. Thorax, Dorsocentral bristles relatively weaker; 2,3 rows of several

short, fine setae laterad of anterior dorsocentrals present, extending to middle

of fossa (absent or only a few present in cubitatus),

MALE GENITALIA (Fig. 22). As figured; exceedingly similar to cubitatus,

differing from it particularly in the following. Basimere. Inner tergal surface

with only 3 submarginal setae ina row. Subapical Lobe. Rodlike setae a-c

smoothly curved, seta c (or external rod of Colless) not angulate or without a

distinct elbow beyond the middle. Phallosome, Apical dorsal beaklike process

slender, relatively longer and distally tapered into a sharp point. Pvroctiger,

Paraproct with an elongate budlike apical lateral lobe which is apically blunt

or pointed, apical sternal lobe broadly rounded.

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype “ with slides of antenna (CH 122) and genitalia

(CT 524), Ayer Mati, Perak? MALAYSIA (ANIC). Paratype 1° with slides of

antenna (CH 123) and genitalia (CT 523), same data as holotype (ANIC).

DISTRIBUTION. Known only from Thailand and Peninsular Malaysia.

Specimens examined: 40.

THAILAND. Chiang Mai: Chang Puak; Kanchanaburi: Ban Sai Yok; 2.

MALAYSIA. Peninsular Malaysia - Perak?: Ayer Mati; 2<.

TAXONOMIC DISCUSSION. Culex gracicornis is morphologically intermedi-

ated between cubitatus and variatus. The male, the only known stage of

gracicornis, is very similar to cubitatus in the details of the modified tufts of

antennal flagellomeres 5-10, but its genitalia are similar to vaviatus in the

phallosome and in the rodlike setae of the subapical lobe. This species was

previously described by Colless (1965: 274) as sp. near cubitatus based on 2

males from Malaya. In this study, 2 other males were also discovered among

the Thailand material, which on the basis of close comparison, perfectly agree

with the Malayan specimens. Based on these records, I am convinced that

gvacicornis is distinct and should be properly recognized as a separate species

within the variatus complex.

BIONOMICS. The type-males of gvacicornis apparently came from a

general field collection. In Thailand, the male from Kanchanaburi was obtained

from rearing an immature collected from a pool on the flooded ground in an un-

shaded area near a native plantation and the other from Chiang Mai presumably

came from a light trap.

20. CULEX (LOPHOCERAOMYIA) WHARTONI COLLESS

(Fig. 22)

Culex (Lophoceraomyia) whartoni Colless 1965: 275 (o"*, 2, L*).

FEMALE. Wing: 3.0 mm. Forefemur: 1.4 mm. Proboscis: 1.7 mm.

In general exceedingly similar to the dark form of variatus, slightly differing

from it in the following. Head. Broad decumbent scales of vertex entirely

dark; narrow decumbent scales sparse, yellowish brown, restricted to dorsal

midline in center; lateral patch of broad scales dark. Cibarial Armature (Fig.

22). Cibarial bar with about 40 teeth in a concave row which is more or less

produced in middle. Thorax, Mesonotal integument deep chestnut brown;

60 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

mesonotal scales entirely blackish. Ppn with several narrow, short, dark

scales or setae cephalad of posterior bristles. Wing, Plume scales on Ro,

Rg relatively broader. Abdomen. Terga II-VII usually with basolateral pale

spots; sterna slightly paler or as dark as terga.

MALE (Fig. 22). As described and figured by Colless (1965: 275-6); gen-

erally dark or blackish as in female; differing from variatus and other related

forms particularly in the following. Palpus, Basal fingerlike processes rela-

tively long, reaching apex of segment 1. Proboscis, Labium with fewer and

weaker dorsal upright setae; lateral setae practically absent. Antenna. Modi-

fied tuft of F-5 very prominent, distinct in shape, composed of 16-22 broad

scales only, 10-16 scales in dorsal group longest, black, apically blunt or

abruptly pointed, followed by lateral group of 5,6 distally pale or entirely

dark curved, sharply pointed scales of decreasing length; F-8 with a typical

or normal J-hooked tuft of fused setae, as cubitatus. Thorax, Anterior sur-

face of fossa without a dense group of fine hairs laterad of dorsocentral bris-

tles.

MALE GENITALIA (Fig. 22). Exceedingly similar to cubitatus, variatus

and gracicornis, differing slightly in the following features. Basimere, Sub-

marginal setae usually 4 innumber. Subapical Lobe, All rodlike setae a-c

smoothly curved in distal portion; leaflet 2, relatively broad, as long as or

longer than leaflet g,. Phallosome, Paraproct with a distinct apical lateral

lobe which is apically rounded.

PUPA. Abdomen: 2.5mm. Paddle: 0.65 mm. Trumpet: 0.65 mm;

index 10. As figured for variatus (Fig. 19) and essentially similar to the

latter in coloration of cephalothorax, abdomen and paddle and in the shape of

trumpet, differing in the following. Trumpet. Shorter, with lower index.

Cephalothorax. Seta 8-C usually single (1-2) and rather weak. Abdomen.

Seta 1-II weakly dendritic or with smaller number of branches; 5-IV 3, 4 bran-

ched; 5-V, VI double; 6-III-VI usually double (1-2); 4-VIII usually double (2-3).

LARVA. Head: 0.76 mm. Siphon: 1.5 mm; index 8. Saddle: 0.3 mm;

siphon/saddle 5. As figured for variatus (Fig. 20), very similar to the latter

in most chaetotaxy, differing particularly in the following. Head, Seta 4-C

longer, as long as or longer than distance between bases of the pair; 16,17-C

rather inconspicuous or sometimes absent. Thovax. Spiculation lighter; seta

3-P 3,4 branched, sometimes more; seta 1-M very strong, as long as or

longer than 3-M; 1-T as long as 2-T. Abdomen. Seta 6-III 4,5 branched.

Comb scales 28-36. Seta 2-X of saddle usually triple (3-4). Siphon. Rather

shorter, thicker; median dark ring usually present; subventral tufts stronger,

more conspicuous, usually 3,4 branched (2-4) each.

TYPE-DATA. Holotype “ with associated pupal and larval skins, slides of

antenna and genitalia, SINGAPORE, reared from a one ege raft (Ss ELS),

June 22, '56, D. H. Colless (ANIC).

DISTRIBUT ION. Thailand, Malaysia, Singapore and Indonesia; 125 speci-

mens examined: 56, 422, 27 L; 27 with associated immature skins (2 p, 25

lp).

THAILAND. Narathiwat: Koh Kien; 1%.

MALAYSIA. Peninsular Malaysia - Selangor: Banting; Bt. Ulu Bakau;

9%, 102, 6 lp. Malaysia (N. Borneo) - Sabah: Tawau; Beaufort; 36%, 279,

27 oe 2 py ai dp,

SINGAPORE. McRitchie Reservoir; Mandai Road; Nee Soon Rifle Range;

8S Oe 2 Ip.

INDONESIA. Sumatra: Benkoelen, Dermajoe; Achin, Kutaraja; 2°.

TAXONOMIC DISCUSSION. The adults of whartoni are apparently darker

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 61

than variatus and cubitatus and can generally be separated from the latter 2

species in the female by the dark broad scales on dorsal and lateral surfaces

of the vertex and the deep brown or black mesonotum and pleuron. The female

cibarial armature is also distinctive in having the concave row of teeth pro-

duced in the middle, quite different from those as described for vaviatus and

other species in the Fvaudatrix Subgroup. The male can be readily separated

from the other members of the vaviatus complex by the modified tuft of anten-

nal flagellomere 5; the fewer and shorter dorsal upright setae of the proboscis

and by the relatively longer basal fingerlike processes of the palpus. The

male genitalia resembles cubitatus more than any other species, but can be

separated from it by the smoothly curved rodlike setae a-c and broader leaflet

81 of the subapical lobe. Both pupa and larva of whartoni are very distinct in

the diagnostic chaetotaxy given above and can be readily separated from vavriatus

and cubitatus as indicated in the keys. All specimens of whartoni essentially

conform to a single type and there is no indication of local differentiation in the

material I have examined. Evidence from comparative male morphology indi-

cates that it is closer to cubitatus than to variatus.

BIONOMICS. Culex whartoni is less common than variatus but has been

more frequently encountered than cubitatus in the collections from Malaysia and

Singapore. The reported breeding sites are ground pools, ditches and swamps

along the seacoast under deep shade of jungle or in the open near forests. In

Selangor, it was reported to prefer acid peaty soils as a breeding site (Colless

1965). Most of the adults were obtained from individual or mass rearings from

the larvae and pupae but, some were also taken while resting near its breeding

sites.

21. CULEX (LOPHOCERAOMYIA) MACDONALDI COLLESS

(Figs. 23, 24)

Culex (Lophoceraomyia) macdonaldi Colless 1965: 276 (“*, @, L*); Delfinado

1966; 108 (2*,. P*; L*): Bram 1967a: 61:(¢*, L*):

FEMALE. Wing: 2.3-2.8 mm (average 2.6 mm). Forefemur: 1.25 mm.

Proboscis: 1.6 mm. In general resembling other members of the variatus

complex, differing in smaller size and in the following characters. Head,

Broad decumbent scales of vertex entirely dark; erect scales slender; lateral

patch of broad scales bluish white, distinct. Palpus and proboscis slender.

Cibarial Armature (Fig. 23). Cibarial bar with a concave row of about 30

teeth, median 4-6 teeth shorter and thinner than lateral teeth. Thorax. Meso-

notal integument reddish brown (field caught specimens) to dark, blackish

(reared specimens); mesonotal scales narrow, blackish. Ppn with a few nar-

row dark scales cephalad of posterior bristles. Pleuron same color as meso-

notum. Legs. Relatively slender; anterior surface of hindfemur largely pale

whitish in basal 0.5, dark in apical 0.5. Wing. Plume scales on Ro, Ra mod-

erately broad clavate. Abdomen. Terga III-VII always with large basolateral

pale spots, from base to over 0.5 of segment; sterna nearly as dark as dorsal

surface of terga.

MALE (Fig. 23). As described by Colless (1965: 276-7); in general similar

to female except for the absence of basolateral pale spots on the abdominal

terga, differing from other members of the variatus complex in the following.

Palpus, Slender; basal fingerlike processes strong and relatively long, pro-

jecting beyond apex of segment 1; segments 4 and 5 moderately plumose. Pro-

62 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

boscis. Dorsal upright setae numerous and dense, extending from basal 0.295

to apex of labium; labial basal bristles relatively long, weak, hairlike. Anten-

na. Modified tuft of F-5 distinctive, composed of 16-20 scales, 8-10 scales in

dorsal group dark, broad, blunt tipped, followed by lateral group of 6-10 scales

which are broad and dark at base, distally narrow, pale and wavy in outline;

F-8 with a typical J-hooked tuft of fused setae; F-9 with 4,5 yellowish brown

to dark blade-like scales; F-10 with 3-5 dark bristlelike setae.

MALE GENITALIA (Fig. 23). As described and figured by Colless (1965:

276-7) and Bram (1967a: 62-3), differing from other members of the variatus

complex in the following features. Basimere, Slender and relatively small;

inner tergal surface evenly concave, with a linear row of 4 submarginal setae,

sometimes 3 or 5; marginal setae very weak and sparse. Subapical Lobe.

Rods a-c thin, smoothly curved, proximally completely overlapped from base

to beyond 0.5 of the length, distally more or less separated; setae d-/ in distal

part 3; leaflets 21,g9 broad, subequally long; seta k moderate. Distimere.

More or less swollen beyond middle, distally tapered into a blunt point; sub-

apical claw slender and short. Phallosome, Apical dorsal beaklike process

very slender, short and weakly sclerotized. Pyroctiger. Crown rather small;

apex of paraproct broadly rounded, without differentiated apical lateral and

sternal lobes; cercal setae 2,3, minute and rather inconspicuous.

PUPA (Fig. 23). Abdomen: 2.2 mm. Paddle: 0.6 mm. Trumpet: 0.62

mm; index 9-10. Generally yellowish white, with indefinite darkened area;

distinguished from other members of the variatus complex by the following fea-

tures. Trumpet, Slender; meatus uniformly cylindrical from base to 0.75 of

length; apical 0.25 more or less swollen; pinna rather broad and long, without

slit extending to meatus. Cephalothorax, Posterior margin of mid-dorsal

ridge dorsad of base of trumpet strongly folded and very irregular; setae 8, 9-C

double. Abdomen, Seta 5-IV 5-8 branched; 5-V usually 4 branched (3-5); 6-III-

V usually triple (2-3); 6-V usually double (2-3). Paddle, Inner and outer parts

pale except for darkened area along midrib and apex, midrib very strong and

dark.

LARVA (Fig. 24). Head: 0.70 mm. Siphon: 1.4 mm; index 8. Saddle:

0.3 mm; siphon/saddle ratio 4-5. As described by Colless (1965: 277), the

following characters are diagnostic. Head. Seta 4-C very short, minute,

less than 0.9 of the distance between bases of the pair; 9-C strong, 2-3 times

as long as 8-C, located near base of antennal prominence cephalad of 8-C; 14-C

3,4 branched. Antennal shaft largely pale, with relatively few and weak spi-

cules. Thorax. Spiculation absent; seta 3-P 3,4 branched, sometimes double.

Abdomen. Setae 6-III-VI 4,5 branched. Comb scales 37-44, scales on posteri-

or row elongate, twice as long as scales on anterior row, middle scales inter-

mediate; apical fringe normal. Saddle same color as siphon; seta 2-X 4 bran-

ched; 4-X with 6 pairs of branched setae, sometimes 4.5-5. Szphon. Slender,

moderately long and usually yellowish white; distal pecten teeth with 1,2 strong

basal denticles and 7-9 graded denticles distally; subventral tufts weak, short,

3,4 branched each, as long as or slightly longer than siphonal width at points of

attachment. 7

TYPE-DATA. Holotype “ (56/33/11) with associated pupal and larval skins,

slides of antenna (CH9) and genitalia (CT 301), SINGAPORE, March 26, 1956,

D. H. Colless (ANIC).

DISTRIBUTION. Thailand, Vietnam, Malaysia, Singapore, Indonesia and

the Philippines; also reported from India. 62 specimens examined: 36“, 179,

9 L; 13 with associated immature skins (3 p, 10 lp).

THAILAND. Sakon Nakhon; Bangkok: Bang Khen; Bang Kapi; Thon Buvi:

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 63

Bangkok Noi; Chon Buri: Sri Racha; Bang Lamung; Chanthaburi: Ban Si

Phraya; Phangnga: Khao Sung; Narathiwat: Kok Kien; Ban Tone; 130, 39, 7

je a Se

VIETNAM. Conson; Pleiku; Kontum; 130, 1 lp.

MALAYSIA. Peninsular Malaysia - Pahang: Tioman; Perak: Trong; 4~,

149, 2p, 3 lp; Malaysia - Sabah: Beaufort; 2 L.

SINGAPORE. 3¢, 1 lp.

INDONESIA. Sumatra: Bengkoelen, Dermajoe; 1.

PHILIPPINES. Mindoro: San Jose; 2, 1 lp.

Additional records from the literature: INDIA. Assam: Golaghat (Colless

1965: 277).

TAXONOMIC DISCUSSION. The modified tuft of antennal flagellomere 5 inthe

male of macdonaldi is superficially similar to variatus, cubitatus and whartoni

but is quite different from these species in the details as described and illus-

trated. There is a strong resemblance in this character between macdonaldi

and the typical fyaudatrix from the Papuan subregion, which I have also exa-

mined for comparison. All stages of macdonaldi except the females can be

readily separated from the rest of the vaviatus complex as indicated in the

keys and as described above. The females are extremely similar to whartoni

and inculus as pointed out by Colless (1965: 277) but appears to be distinct

from the latter 2 species in the details of the cibarial armature.

Culex macdonaldi as interpreted here is exclusive of the ''Gunong Tebu

form" of Colless (1965: 277). The latter was previously assigned to mac- ©

donaldi by this author, but present study has shown that it is distinct from it

in the adult, male genitalia and larval stage as described below under pazvoji.

BIONOMICS. The breeding sites of macdonaldi are restricted to shaded

fresh water ground pools in marsh or swamps near tidal zones at sea level

along the coast (Colless 1965). Several adults have been collected while rest-

ing among vegetation near the breeding sites and only a few from light traps.

22. CULEX (LOPHOCERAOMYIA) PAIROJI NEW SPECIES

(Figs. 25, 26)

Culex (Lophoceraomyia) macdonaldi (Gunong Tebu form), Colless 1965: 276

ipa),

FEMALE. As described for macdonaldi, differing from it particularly in

the following features. Head, Lateral patch of broad scales of vertex pale

brown to nearly white, very distinct. Cibarial Armature, Cuibarial teeth

longer, more numerous, 90-60, all subequal in length and size. Thorax,

Mesonotal disc with acrostichal bristles extending from anterior promontory

to the level of wing base. Wing. Plume scales on Ro, Rg relatively broader.

MALE (Fig. 25). As in female; almost identical to macdonaldi in the de-

tails of palpal, labial and antennal characters, differing from it in the presence

of acrostichal bristles on mesonotal disc and in having longer dorsal upright

setae on proboscis, the longest ones 4 times as long as labial width at points

of insertion.

MALE GENITALIA (Fig. 25). Essentially as described for macdonaldi,

differing from it particularly in the following. Basimere. Inner tergal sur-

face with a prominent row of 6,7 very strong submarginal setae, somewhat

resembling quadripalpis; marginal setae stronger, 6-8, about 0.25 of submar-

ginals. Phallosome, Apical dorsal beaklike process stouter, darker and well

64 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

sclerotized.

PUPA. Essentially as described and figured for macdonaldi, differing

particularly in the following. Abdomen. Seta 5-V usually double, rarely triple

or more branched; 4-VIII usually triple or 4 branched. Paddle. Inner part

lightly to strongly infuscated, outer part pale except for darkened areas along

midrib.

LARVA (Fig. 26). Head: 0.7mm. Siphon: 1.9-2.1 mm; index 10-11.

Saddle: 0.36 mm; siphon/saddle ratio 6-7. Extremely similar to macdonaldi,

differing from it particularly in the following. Head, Seta 9-C of the same

magnitude as 8-C, located at short distance cephalad of the latter. Thorax,

Seta 3-P usually double (2-3). Siphon. Relatively more slender, with greater

index and siphon/saddle ratio, median dark ring usually present; pecten teeth

barbed with evenly fine denticles, basal denticles weaker and finer than distal

denticles.

TYPE-DATA. Holotype “ (SS 32-106) with associated pupal and larval skin

and slide of genitalia, Khao Yai National Park, Nakhon Ratchasima, THAILAND,

deep shaded jungle swamp, 800-900 m, November 1, 1967, S. Sirivanakarn

(USNM); Allotype (SS 32-113) with associated pupal and larval skins; Para-

types: 2 lp (SS 32-110,111), 3 po (SS 32-100, 101,112), 5c (SS 31, SS 34);

42 (SS 28, SS 31, SS 34), same data as holotype (USNM). This species is dedi-

cated to Mr. Pairoj Suwanakorn, Director of Khao Yai National Park, Thailand

Ministry of Agriculture.

DISTRIBUTION. Thailand, Malaysia, Singapore and Indonesia. 76 speci-

mens examined: 280", 169, 32 L; 12 with associated immature skins (4 p, 8 Ip).

THAILAND. Nakhon Ratchasima: Khao Yai National Park; 21%, 129, 28 L,

3p, 6 lp.

MALAYSIA. Peninsular Malaysia - Pahang: Mela K. Lipis; Benta K.

Lipis; Perak: C. Highlands Rd.; 20, 29, 1 L, 1 p; Malaysia - Sabah: Tawau;

Kota Kinabalu; Beaufort; 2“, 3 L.

SINGAPORE. McRitchie Reservoir; Nee Soon Rifle Range; 3c", 29.

INDONESIA. Java: Surabaja; Sumatra: Dermajoe; 2.

Additional records from the literature. PENINSULAR MALAYSIA. Treng-

gsanu: Gunong Tebu (Colless 1965: 277; as macdonaldi).

TAXONOMIC DISCUSSION. The above descriptions of pairoji have been

based on the study of the specimens from Nakhon Ratchasima, Thailand. The

male of paivoji apparently agrees well with the ''Gunong Tebu form" of m™ac-

donaldi (Colless 1965: 277). Recently,a number of pairoji specimens have also

been discovered in Malaya, Singapore and Indonesia, indicating that it is a

widespread species. Culex pairoji appears to be restricted to high inland ele-

vations in mountainous areas except in Malaya and Singapore where it has also

been found along the coast in sympatry with the macdonaldi populations.

Culex patiroji is identical to macdonaldi in the modified tuft of male antenna,

but is distinct from the latter in the adults by the presence of acrostichal bris-

tles on the mesonotal disc and by the presence of a prominent row of 6,7 strong

submarginal setae on the basimere of the male genitalia. The female cibarial

armature and the larva are also distinct from macdonaldi as indicated in the

above description, but the pupa can not be separated from that species with

certainty.

BIONOMICS. The immatures and the adults of pairoji from the type-

locality in Thailand were collected from a large jungle swamp under heavy rain

forest shade at an elevation between 800-900 m. The water was fresh, contain-

ing numerous algae, fallen leaves and other aquatic vegetation. In Malaya and

Singapore, the immatures were collected from pools at the edge of a stream,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 65

swamp and marshy depression from sea level to about 30 m. The adults from

Gunong Tebu, Trengganu were collected at a high elevation (Colless 1965).

Several adults in Thailand were collected by a sweep net and a sucking tube

while resting under leaves of plants near the breeding site. The female does

not appear to bite man.

alphus complex

23. CULEX (LOPHOCERAOMYIA) ALPHUS COLLESS

(Migs 21,28)

Culex (Lophoceraomyia) alphus Colless 1965: 283 (o*, ?, L*); Bram 1967a:

53 (oe Dee

FEMALE. Wing: 2.6mm. Forefemur: 1.3 mm. Proboscis: 1.5 mm.

Small, blackish species, with general facies similar to most members of the

Fraudatrix Subgroup. Head, Vertex largely covered with broad, dusky decum-

bent scales which are pale whitish along upper eye margin, forming a distinct

ocular line; narrow decumbent scales sparse, restricted to dorsal midline in

center; erect scalesvery Slender. Proboscis with 6 strong labial basal setae

which are as long as palpus. Cibarial Armature (Fig. 27). As figured; ci-

barial bar with an even concave row of 50-60 subequal teeth which are apically

blunt or truncate. Thorax, Mesonotal integument dark brown, scales narrow,

blackish. Ppn bare on anterior upper surface; posterior bristles 2,3. Pleuron

same color as mesonotum; ppl bristles 2; 1 lower mep bristle absent. Legs.

Slender; anterior surface of hindfemur silvery white in basal 0.5, dark in

apical 0.5. Wing. Scales minute and rather scanty on all veins. Abdomen.

Terga dark, without basolateral pale spots; sterna yellowish scaled.

MALE (Fig. 27). In general similar to female, with diagnostic features

as described and figured by Colless (1965: 283-5). Palpus. Rather short,

very slender, as long as or slightly longer than proboscis; basal fingerlike pro-

cesses well developed, as long as the distal part of segment 1; segments 4 and

. Slightly upturned, with several weak and short bristles. Proboscis. Slender;

dorsal upright seta strong, numerous, extending from 0.25 from base to apex

of labium, longest setae 4,5 times as long as labial width at points of insertion.

Antenna, Flagellar whorls weakly to moderately plumose; F-5 with a large

characteristic tuft of 12-18 dark scales, 6-8 scales in dorsal group longest,

blunt or point tipped, as long as the combined length of the next 4 flagellomeres,

followed laterally by 2-4, narrower, shorter, pointed scales and ventrally by 3-6,

subapically swollen scales of increasing length; F-6, 7 with crumpled tufts of

curled setae as described in most forms; F-8 with normal J-hooked tuft; F-9

with 3,4 yellowish bladelike scales; F-10 with 2-4 dark, strong, bristlelike

setae.

MALE GENITALIA (Fig. 27). Rather small; generally similar to the mem-

bers of the rubithoracis and quadripalpis complexes, distinctive in the following

features. Segment IX. Tergal lobe with 2 very strong and long setae; sternum

usually bare, sometimes with 1 seta towards caudal margin. Basimere. Nor-

mal; with a row of 4 strong submarginal setae and a few weak marginal setae

on inner tergal surface; lateral tergal surface with a group of several rather

_ strong setae laterad of the submarginals. Subapical Lobe. Rodlike setae a-c

stout, subequal and smoothly curved or straight; setae d-f 5-6, rather broad,

flattened, bladelike, very distinct; leaflets 84> 89 well developed; seta /” strong,

66 Contrib. Amer. Ent. Inst., vol. 18, no. 4, 1977

Phallosome, Apical dorsal beaklike process, short, small, sustaining an

acute angle with the main stem. Pyvroctigery. Crown small, largely composed

of fine, spinelike spicules; paraproct with a small short apical lateral and

sternal lobes, the latter apically rounded; cercal setae 3, distinct.

PUPA (Fig. 27). Abdomen: 2.2 mm. Paddle: 0.55 mm. Trumpet:

0. 52-0.59 mm; index 8-9. Cephalothorax and abdomen more or less uniform-

ly brownish. Trumpet, Uniformly slender, long, darker than underlying in-

tegument; pinna broad, with slit extended to meatus. Cephalothorax. Seta 8-C

double, strong, 2,3 times longer than 9-C; 9-C triple. Metanotum. Seta 10-C

always single. Abdomen. Most setae rather weak and short; seta 5-IV usually

triple (3-4); 5-V double; 5-VI single; 6-IV-VI usually triple (2-4); 9-VII double;

4-VIII usually 4 branched (3-5); 9-VIII 4,5 branched. Paddle. Inner and outer

parts pale except for areas adjacent to midrib, midrib strong and dark.

LARVA (Fig. 28). Head: 0.55 mm. Siphon: 1.1 mm; index 8-9. Saddle:

0.3 mm; siphon/saddle ratio about 4. Relatively small sized; complete chaeto-

taxy as figured, distinctive in the following. Head, Usually brownish; seta 4-C

short, forked into 4 branches; 5-C rather short, 0.5-0.7 of length of 6-C, both

setae double; 14-C double. Thorax. Spiculation absent; seta 3-P 4-7 branched;

4-P double; 14-P single; 1-M 0.5 of 3-M. Abdomen. Seta 7-I single; 6-I

eriple; 6-II double; 6-III triple; 6-IV-VI 4,5 branched. Comb scales about 27,

posterior scales elongate, 2,3 times as long as scales on anterior row; apical

fringe normal. Saddle seta 2-X usually with 3 short branches and 1 long branch.

Siphon, Slender, moderately long, uniformly brownish, concolorous with head

capsule; pecten teeth characteristic, 9-11 in number all apparently simple or

with inconspicuous fringe of extremely fine denticles; subventral tufts short,

each 3-5 branched, as long as siphonal width at points of attachment; median

caudal filament of spiracular apparatus well developed.

TYPE-DATA. Holotype o (0477/6) with associated pupal and larval skins

and slide of genitalia (CT 551), Kg. Sijangkang, Selangor, Malaya | MALAY-

SIA], January 21, 1958, D. H. Colless (ANIC).

DISTRIBUTION. Thailand and Malaysia. 10 specimens examined: 3c,

32, 4 L; 5 with associated immature skins (4 p, 1 lp).

THAILAND. Narathiwat; 1“ (only antenna and genitalia).

MALAYSIA. Peninsular Malaysia - Selangor; Kg. Sijangkang (type-locali-

ty); Banting; Trengganu: Dugun; Johore: Kota Tinggi; 1“, 22, 2 L, 3p;

Malaysia: Sabah: Kota Kinabalu; Beaufort; 1%, 12, 2 L, 1p, 1 lp.

Additional records from the literature. SINGAPORE (Colless 1965: 285).

TAXONOMIC DISCUSSION. Culex alphus can generally be distinguished

from the other members of the Fraudatrix Subgroup in all stages by the rela-

tively small size; in the females by the presence of 6 long labial basal setae of

the proboscis (usually 2 in all others); the male by the characteristic tuft of

antennal flagellomere 5; the male genitalia by the presence of 2 very strong

and long setae of tergal lobe of segment IX and by other features of the basi-

mere and the phallosome as described and figured; the pupa by having setae

10-C and 5-VI single and in the larva by the elongate posterior comb scales

and the simple or fine denticulated pecten teeth of the siphon.

Culex alphus apparently combines features of the variatus and the rubi-

thorvacis complexes. It resembles the variatus complex in the development

of the modified tuft of the male antennal flagellomere 5, but its larva is similar

to the vubithoracis complex in having seta 3-P 4-7 branched and seta 7-I

single. The male genitalia is somewhat intermediate between the 2 complexes

but is apparently more similar to the rubithoracis complex than to the variatus

complex.

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 67

BIONOMICS. Culex alphus has been seldom encountered and is apparently

rare. The breeding sites are pools, ponds, wells and forest swamps at sea

level along the coast. Larvae recorded by Colless (1965: 285) were collected

from a shaded well of rather peaty water in association with the specimens of

whartoni. The adults largely came from rearing the pupae or larvae and a few

were collected in the field.

MA MMILIFER GROUP

FEMALE. As described for the subgenus; generally distinguished from

the Fraudatrix Group by darker coloration, absence of basolateral pale spots

or basal pale bands on abdominal terga and by the following features. Head,

Decumbent scales on dorsal and central areas of vertex largely narrow, cla-

vate or linear; erect scales usually completely dark, sometimes partially pale

bronzy or yellowish in center. Proboscis with 4 labial basal setae. Cibarial

Armature, Cibarial teeth relatively few, varying from 15-40. Thovax, Meso-

notal integument deep brown to black; scales denser and entirely blackish.

Scale patch on pleuron usually absent, sometimes present on Ppl, upper corner

and posterior border of stp. Wing, All veins with more numerous scales;

scales on veins Ro, Rg narrow, linear. Abdomen, Terga entirely dark; baso-

lateral pale spots or Baal pale bands absent.

MALE. Essentially similar to female in general external features; distin-

suished from the Fvaudatrix Group by the following. Palpus, Slender, usually

as long as proboscis, sometimes longer or reduced to about 0. 50-0. 75 of pro-

boscis length; basal fingerlike processes absent; segments 4 and 5 very weakly

plumose or with relatively few bristles on lateral and mesal surfaces. Pvo-

boscis., Labial basal setae weak, hairlike and short; false joint absent or not

clearly marked by flexion; dorsal upright setae in distal portion of labium com-

pletely absent or not developed. Antenna, Pedicel with a very distinct, nipple-

like spiculose prominence on inner dorsal surface, sometimes not developed

(certain New Guinea forms); modified tufts of scales and setae usually present

on F-5-8 or F-5-9, sometimes absent; modified tuft of F-5 usually small, in-

conspicuous, composed of short, acute setae; sometimes large, distinct, with

some broad, acute or blunt tipped scales; F-6 to F-8 usually with similar tufts

of dark setae as described for the Fraudatrix Group, sometimes reduced or

modified; F-9 with or without a tuft of long, bristlelike setae; F-10 without any

modified setae.

MALE GENITALIA. Segment IX. Sternum usually with a row of strong

setae or scales towards posterior caudal margin, rarely absent. Basimere.

Usually small, conical, sometimes modified; submarginal setae usually well

developed, forming a prominent row parallel to tergomesal margin, sometimes

absent or not differentiated. Subapical Lobe. Varied; all specialized setae

and leaflets in proximal and distal parts usually present, well developed, some-

times strongly modified. Distimere. Sickle-shaped, usually tapered and re-

curved towards apex, sometimes apex expanded or modified; subapical portion

with or without distinct crest of fine spicules on dorsal surface. Phallosome.

Dorsal lobe of lateral plate usually with a large, partially denticulate external

process and a simple, spinelike internal process; ventral lobe strongly reduced

or poorly developed. Pyroctiger. Crown large, composed of several coarse

spicules and numerous spinelike spicules; paraproct with a short, rounded api-

cal lateral lobe only; cercal setae usually 3, 4.

PUPA. Trumpet. Varied from short, bell-shaped or funnel-shaped, 0. 25-

68 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

0.40 mm (pitcher plant forms) to long, cylindrical, 0.60-0.80 mm; pinna usu-

ally without slit extending to meatus. Cephalothorax., Seta 8-C usually 1-3

branched, 9-C usually double, sometimes single or triple. Metanotum, Seta

—11-C single or double. Abdomen. Seta 5-IV 1-3 branched, sometimes 4,5

branched; 5-V single or double; 5-VI usually single, sometimes double; 6-III-

VI usually 1-3 branched; sometimes 4-6 branched; 9-VII usually shorter than

9-VIII. Paddle. Outer and inner parts usually pale, midrib weak, pale to

strong and pigmented.

LARVA. Varied, depending on types of breeding habitat. In general

similar to the Fraudatrix Group. Head. Usually broader than long, sometimes

as long as broad (certain pitcher plant forms); seta 4-C as long as or longer

than distance between bases of the pair; 5-C usually double, long, sometimes

3 or more branched and short; 6-C 1-3 branched. Antenna as long as or shorter

than head; spicules usually numerous, strong, sometimes reduced to completely

absent; seta 1-A large, plumose or sometimes considerably reduced; seta 2,

3-A usually subapical, sometimes apical. Thovax. Spiculation usually absent,

sometimes present, moderately to strongly developed; seta 3-P usually single;

7-P usually double, rarely single or triple; 8-P single, subequal to 7-P, or

sometimes strongly reduced; 14-P usually double, rarely single; 1-M strong,

as long as 3-M, sometimes shorter. Abdomen. Spiculation same as thorax;

live or whole mounted specimens without pattern of dark and light bands; setae

6-I, II usually triple, sometimes double or 4,5 branched; 7-I usually double,

sometimes single; 6-III-VI double or triple. Comb scales numerous, some-

times few, all subequal or differentiated in size; apical fringe normal, rarely

modified; seta 2-VIII always single. Saddle usually lightly spiculated on pos-

terior caudal margin, sometimes heavily spiculated. Szphon. Usually long,

slender as in the Fvaudatrix Group, sometimes considerably reduced (certain

pitcher plant forms); pecten teeth usually 10-15, sometimes few, 2-4; subven-

tral tufts usually strong and prominent, 4 pairs, sometimes 6 to 9; median

caudal filament absent or not developed except for certain ground pool forms.

DISCUSSION. The Mammilifer Group as interpreted here essentially cor-

responds to the original group C (Cyathomyia or Mammilifer) of Edwards (1932:

196) and group B (Mammilifer Group) of Colless (1965: 264). In the Oriental

region, the group includes all species which exhibit a distinct spiculose pro-

minence on the pedicel of the male antenna. Culex wilfredi, previously as-

signed to this group (Colless 1965: Bram 1967a), is here considered as repre-

senting a distinct lineage or group, the Wilfredi Group. The Mammilifer Group

is the most diverse major phyletic line in the subgenus. It exhibits a rather

distinctive pattern of distribution, being confined largely to the Indomalayan

parts of the Oriental region. Elsewhere, it has been reported only from the

Papuan part of the Australasian region where 4 species of 3 subgroups are

represented (Sirivanakarn 1968).

On the basis of the breeding sites and the morphology of all stages, the 32

Oriental species of the Mammilifer Group recognized in this study fall into 2

well marked subgroups: Mammilifer and Brevipalpus (or By and Bo of Colless

1965: 264-5). The Mammilifer Subgroup comprises 22 species whose breeding

habitats are usually tree holes, bamboos and rock pools or sometimes leaf

axils of pandanus and ground pools. The Brevipalpus Subgroup contains 10

species whose breeding habitats are exclusively pitcher plants. Although the

members of the Brevipalpus Subgroup exhibit some overlap with those of the

Mammilifer Subgroup in male characters, its separation from the latter on the

basis of the restricted breeding site and certain features of the male and lar-

val characters, as given by Colless (1965), is apparently justified and is fol-

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 69

lowed here.

KEYS TO SUBGROUPS AND SPECIES

FEMALES!

His Cibarial teeth coarse, short, widely spaced, usually 15-25; lower

mep bristle absent (Brevipalpus Subgroup). ..........6-. q

Cibarial teeth usually slender, elongate, closely spaced, 30-40;

lower mep bristle usually present, (except uniformis, bengal-

ensis, flavicornis and lasiopalpis) (Mammilifey Subgroup). ... 2

Mammilifer Subgroup

2{4), Lower mep bristle absent.

uniformis, bengalensis, flavicornis and lasiopalpis

LOWE? WEP Dristle Presents. ietCwea ee eee eare e eees 3

3(2). Vertex with a broad, pale, distinct ocular line; ground pool

PRECCOR Gai A: OF us eo Pee het Sree Ma aia near ee Ce eg a ise 4

Vertex with or without a narrow, pale, distinct ocular line;

breeders in tree holes, bamboos, rock pools or leaf axils. ... 95

4(3). Cibarial teeth short, coarse, 18 in number. ........ pholeter

Cibarial teeth elongate, relatively slender, 35 in number.

mammilifer and wardi

5(3). Anterior mesepimeron with a row of 5,6 weak setae dorsad of lower

IED ITA BU Ce ad: ORS Se ek as kai cee Se lavatae

Anterior mesepimeron without above setae. ........6..4.. 6

6(5). Relatively small species; wing length usually less than 3.0 mm.

impostor, traubi, demissus and ganapathi (inseparable, use keys

to male, pupa and larva).

Relatively larger species; wing length usually more than 3.0 mm.

spiculosus, minor, bicornutus, bandoengensis, tuberis, kunhsi,

peytoni and eukrines (inseparable, use keys to males, male

geenitalia, pupae and larvae).

Brevipalpus Subgroup

Tt): Upper corner and posterior border of stj with a distinct patch of

pale scales) (cuvtipalpis* complex)... curtipalpis

and sumatranus

Upper corner and posterior border of stp without distinct patch

Of Pale BCATOR MA AOS IS Se i ee Se ee eae 8

RON pas ae Mie tp rated

icomptus and crassicomus are unknown.

8(7).

9(8).

10(9).

Pane

3(2).

4(3).

(4).

6(5).

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Erect scales:of vertex entively Garks cnt ok) oS oes ees 9

Erect scales of vertex pale golden or bronzy in center, dark on

posterolateral area (bvevipalpus complex).

brevipalpus, lucaris, eminentia and acutipalus (inseparable, use

key to males and male genitalia).

Relatively small species, wing length not exceeding 3.0 mm; 18-25

Ce OGL ly eer erie aledeur eines sy eratey oes a) Si eee 10

Relatively large species, wing length more than 3.0 mm; 30 cibarial

tee ICMECNT COMPLIES ohistd oc iivialiamel Gees Wie ea jenseni

Minute species; 18-20 cibarial teeth (hewitti complex). . . hewitti

Relatively larger species; 23 or 25 cibarial teeth (mavalis complex).

navalis

and coerulescens

MALES AND MALE GENITALIA

Palpus as long as or longer than proboscis. .......+4.+e+-6-. 2

Palpus 0.50-0.75 of proboscis length (Brevipalpus Subgroup,

BAA acs email Pha eae pee ole of ty pia ete) > peseee atl! atcqnls 26

Inner tergal surface of basimere with 1,2 submarginal setae or

none; leaflet g, of subapical lobe absent or not developed

(Brevuipolpus Subarogi <4 Part). aus sd eG eei alee eee 24

Inner tergal surface of basimere with 1 or more rows of at least

4 submarginal setae; leaflet 8 of subapical lobe present,

well developed (Vammilifer Subgroup). .........-+.+.-. a

Mammilifer Subgroup

Palpal segments 2 and 3 with prominent lateral rows of numerous

fine hairlike setae. (flavicornis complex). .......... 4

Palpal segments 2 and 3 without prominent lateral rows of

numerous fine hairlike setae as above. ............. 9)

Basimere large, strongly swollen at base; apex of distimere

VGC ip iGrt Cig sn) o0e hi roe eee cla a as ed ee ee flavicornis

Basimere small, slender, normal; apex of distimere not modified

a ee he ae ng eR ee ks lasiopalpis

Antennal flagellomeres without modified tuft of scales or setae

Gipasion:coniples a Save tcc ar Mead aan we ws impostor

Antennal flagellomeres with modified tufts of scales or setae on

GG Oae Te Oia wig Ler oe Os Oe a a ee Sa 6

Internal process of phallosome long, reaching or usually projecting

beyond apex of external process; external process with a broad,

spiculose or denticulate apical lobe (Figs. 30, 32)....... 7

Internal process of phallosome shorter, not projecting beyond apex

of external process; external process with a smaller, elongate,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 71

spiculose apical lobe sternad (Figs. 39, 43). ......... 12

7(6). Antennal flagellomere 9 without a tuft of modified setae (tvaubi

COMDTER I Rh eee ae iy RSA Re 3, RE ATE 8

Antennal flagellomere 9 with a tuft of modified bristlelike

Sotae (WANN CY COMPS BP OE EOS es 10

8(7). Modified tufts of antennal F-5-7 rudimentary or weakly developed;

antennal F-8 with smoothly curved tuft of fused setae (Fig.

Saye SOP EY RS. QO IE ie Bee RE ee Ba: lavatae

Modified tufts of antennal F-5-7 well developed and distinct; F-8

with sigmoid or J-hooked tuft of fused setae. .......... 9

9(8). Modified tuft of antennal F-5 with some broad, blunt-tipped scales;

basimere with a row of 4 submarginal setae (Fig. 32).

uniformis

Modified tuft of antennal F-5 with narrow, acute scales only; basi-

mere with a row of 6,7 submarginal setae (Fig. 30)... . tvraubi

10(7). Palpal segments 4 and 5 shortened, segment 4 about 0.5 of

BOOMIENE By 6 rites be PW ee eo a BEE PA Re gs wardi

Palpal segments 4 and 5 normal, subequal in length. ....... 11

11(10). Modified tuft of antennal F-5 with some long, broad, blunt-tipped

scales; rod a of subapical lobe strongly curved, distally separ-

ated from rods b, c by a wide gap (Fig. 34). .... mammilifer

Modified tuft of antennal F-5 with short, acute flattened setae only;

rod a of subapical lobe gently curved, largely overlapping

with rods band CE -CPig eS BI. as Be oe eZ demissus

12(6). Antennal F-9 without modified bristlelike setae. ......... 13

Antennal F-9 with modified bristlelike setae. ........... 14

13(12). Modified tuft of antennal F-5 distinct, composed of 5,6 narrow,

acute scales, as long as the next 2,3 flagellomeres: F-8 with

a slender, gently hooked tuft (Fig. 39). ......... ganapathi

Modified tuft of antennal F-5 indistinct, composed of 4-7 short,

acute setae as long as 1 flagellomere; F-8 with a thick, more

weakly hoékedumtt(Tie. 42lyo ure uted bo ep iy ieely spiculosus

14(12). Modified tuft of antennal F-5 conspicuous, with some broad blunt-

tipped scales (prolefer complex)... 363 eG se ek ea pholeter

Modified tuft of antennal F-5 small, inconspicuous, with short

setae only OF BDRSBEs 2G Pore ee kon ae ee eS 15

15(14). Modified tuft of antennal F-5 with a small tuft of 3-5 very short

spinelike setae (Fig. 43) (minor complex, in part)....... 16

Modified tuft of antennal F-5 absent or not developed. ...... 22

16(15). Basimere with 15-17 submarginal setae, in 2,3 prominent rows

strongly diverging laterad on lateral tergal surface (Fig. 45).

bandoengensis

Basimere with 4-7 submarginal setae in 1 row, more or less parallel

iO TOPSOMOSAl HIRT e6e 6.8 ee oo ee ee 17

17(16).

18(17).

19(18).

20(19).

21(20).

22(15).

2a(a2).

24(2).

Contrib. Amer. Ent. Inst., vol. 138, no. 4, 1977

Distal part of subapical lobe elongate and hairy (Fig. 45). . tuberis

Distal part of subapical lobe not-as above. . ...°. 5 6. 2s owe 18

Proboscis strongly swollen in middle, with many strong, prominent

setae on lateral and ventral surfaces. ........ bicornutus

Proboscis uniformly thick, without numerous strong setae on

lateral and ventral surfaces as above. ......... ee ee cae)

Rod a of subapical lobe markedly curved in middle, distally

separated from rods 0,c by a wide gap; basimere with 6,7 very

strona submiarginal setae: (Fic.. 43)uicigee euetiad Ca vieoks- minor

Rod a of subapical lobe gently curved or straight and largely over-

lapped with rods 0, c; basimere with 4-7 weaker submarginal

Serpe jiies. 40249) bce aoe Bos Ores Di dbek eee ON 20

Antennal F-9-11 with a few to several short or long setae distad

of normal whorls (Figs. 7) AoW Jonmin bocteus i ewes 21

Antennal F-9-11 without any setae distad of normal whorls. . kuhnsi

Basimere with 4 submarginal setae; F-9-11 with several long setae

distad of normal whorlsh a. 05. . ies eo ae CVASSiCOMUS

Basimere with 6,7 submarginal setae; F-9-11 with 1,2 short

setae distad Of normal whorls... 6-4 6 ose le ee es incomptus

Internal process of phallosome strong and long, its apex reaching

base of apical lobe of external process; external process slender

with several strong denticles (Fig. 49)(7inor complex, in part).

bengalensis

Internal process of phallosome weak and rather short, its apex not

reaching base of apical lobe of external process; external process

broad, with or without a small number of weak denticles (pey-

LON? COMPIER). ca a as is a ee 23

Modified tufts of antennal F-6, 7 very weakly developed and rather

indistinct; subapical lobe of basimere with numerous fine hair-

like setae associated with usual specialized setae on mesal

SUTiaCe (ies OF) he es eukrines

Modified tufts of antennal F-6, 7 well developed and distinct;

subapical lobe of basimere without any fine hairlike setae

associated with usual specialized setae as above (Fig. 50).

peytoni

Brevipalpus Subgroup

Antennal F-5 with a distinct modified tuft of 4,5 narrow, acute

scales; F-6 with a crumpled tuft of twisted or curled setae

(laa SteGmnlesynn oe. Soames GS Saga ee tae eda ie. 25

Antennal F-5 without modified tuft of setae or scales; F-6 with 4,5

subapically swollen scales in a row (Fig. 61) (hewitti complex).

hewitti

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region te

25(24). Roda of subapical lobe stout, strongly curved and distally separated

from rods. b,c bya wide @apr CP ie. BE). iw see Sh heeds navalis

Rod a of subapical lobe slender, more or less straight and largely

overlapping with rods O70 (Fie. 60). 0 oe Ge coerulescens

26(1). Distimere normal, distally tapered into a recurved point; basimere

small, conical; subapical lobe small, with only a few accessory

setae in group d-f (Fig. 63) (jenseni complex). ...... jenseni

Distimere with modified expanded apex; basimere relatively large

and broad; subapical lobe large, usually with several accessory

SCtaG 1 BROUD dei nk ne ew SaaS ee Sa ee 27

27(26). Subapical lobe with numerous accessory setae in group d-f; leaflet ¢

not developed or absent ®revipalpus complex). ........ 2

Subapical lobe with only a few accessory setae in group d-/; leaflet

81 present, very broad, heart-shaped (curtipalpis complex). . 31

28(27). Antennal F-9 with a weak tuft of 4,5 bristlelike setae. ...... 29

Antennal F-9 with a strong tuft of 10 or more flattened setae.

eminentia

29(28). Seta h of subapical lobe very strong and long, of the same magnitude

as proximal vods. (Wie.b5), 6 4645. ile EN Sha brevipalpus

Seta h of subapical lobe weaker and shorter than proximal rods or

sometimes reduced, inconspicuous (Figs. 67, 68). ...... 30

30(29). Antennal F-8 with a strong modified tuft; setae of subapical lobe

as .Getauicd ane Pigg GO hiyete vse seo al aa eek Skt eds lucaris

Antennal F-8 with weaker modified tuft; setae of subapical lobe as

COPATIER IN LOG OO a Se Art ee ig ae han acutipalpus

31(27). Antennal F-5-9 with distinct tufts of modified setae. . . curtipalpis

Antennal F-5-9 without distinct tufts of modified setae. . sumatranus

PUPAE!

t. Trumpet long, cylindrical, 0.5-0.7 mm; breed in tree holes,

bamboos, rock pools, leaf axils or ground pools (Vammilifer

Suberoup)... 3.0 daaeiewadtiod sae, es ee a ee 2

Trumpet short, bell-shaped or funnel-shaped, 0.2-0.4 mm;

breed in pitcher plants (Brevipalpus Subgroup). ........ 19

Mammilifer Subgroup

a(4). Pinna of trumpet with slit extending to meatus; setae 6-III-VI 4, 5

branched (except pholeter); usually breed in ground pools or

sometimes rock pools. ....... Sy Rib cal ig ae aa Oe ae ie ee 2

lL incomptus and cvassicomus are unknown.

(4).

6(4).

T(2):

8(7).

9(8).

10(9).

11(10).

12(10).

13(7).

14(13).

15(14).

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Pinna of trumpet without slit extending to meatus; setae 6-III-VI

usually 1-3 branched; breed in tree holes, bamboos, rock pools

Oy “ear axils Ssh oO Or aes Se rae joss ea aed ie if

Seta 5-C double, strong, about 2 times as long as 7-C. . . . pholeter

Seta 5-C 4-6 branched, as long as or shorter than 7-C....... 4

Seta 8-C usually 3,4 branched; 5-IV double or triple; apex of paddle

PO MAES HO POU ACG ee ig tags Sern ee 5)

Seta 8-C usually double; 5-IV 5, 6 branched; apex of paddle

PONG rae Bere On Te aN ate, Ere poe Mite Pe Oe es! a: Sa 6

Seta 10-C 3,4 branched; apex of paddle pointed. ..... flavicornis

Seta 10-C double; apex of paddle rounded. ........ lasiopalpis

Seta 4-VIII usually triple; 6-IV-VI usually 5 branched. . mammiilifer

and wardi

Seta 4-VIII double; 6-IV-VI usually 4 branched. .... bengalensis

Petit tt, Woualiy meee Sere ew be SO Se ee ae ee ae 8

Ar re hah Gea Se a a ee ele be ee 13

Seta 10-C 3,4 branched; 8-C 3,4 branched. .......... kuhnsi

Seta 10-C double; 8-C usually double. ..........2...4.. 9

Seta 5-C strong and long, about 2 times as long as 7-C; trumpet

very long and thin; tindex 19-20. ose oe eS ganapathi

Seta 5-C weaker, as long as or shorter than 7-C; trumpet

moderately long and thick, index 7-10............. 10

Seta 0-IV triple; 5-IV-VI as long as or shorter than segment

TOTO Wiese Rohe hey eh hepa eine gree AL AOL Re 11

Seta 5-IV single; 5-IV-VI 1.5-2.0 times longer than segment

PO ae eee tye a te Pe CES Se eS 12

Seta 0-V single; 6-IV, V usually double. .......... spiculosus

Seta 5-V double or triple; 6-IV, V usually single. ....... tvaubi

All setae strong and long; 6-III-V usually triple; 5-IV-VI usually

2 times longer than segment following. ......... eukrines

All setae relatively weaker and shorter; 6-III-V usually double;

o-IV-VI 1.5 times longer than segment following. .... peytoni

peta 10-C usually single and weak. 4.2.95. 6 ee ee zmpostor

Seta 10-Cusually double and strong ati. ek ek ee 14

Paddle with broadly rounded apexsy. Goo ee ae Se Pe 15

Paddle with acuminate or pointed apex. ........... tuberis

Bet ory Weta ye rae re a a Pe Oe ae 16

Oe ih ry eee a ek ne ema eg oe ae pane ee gg 17

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 79

16(15). Seta 9-VII 5,6 branched; 5-V, VI single, strong, nearly 2 times

longer than segment following. ...........6.-. demissus

Seta 9-VII 3,4 branched; 5-V, VI double, weak, slightly longer or

shorter than segment following. ............. lavatae

17(15). Seta 5-C 5,6 branched; length of trumpet 0.5 mm, index 8.

uniformis

Seta 5-C 2-4 branched; length of trumpet 0.7 mm or more, index

at dpaet ia siaed 206.0300 Na Giee oe ROTO Se eee 18

18(17).. Seta: 8-GCausvally tripicnn 1s aren bos BAL hs bandoengensis

Seta 3-C: doubles 12 sities erlereie By Pee FA eee minor

Brevipalpus Subgroup

19(1). Trumpet elongate, funnel-shaped, 0.4 mm in length....... 20

Trumpet short, bell-shaped, 0.2-0.3 mminlength........ ae

20(19).. Seta J-G-einele; 10-CrcinGlens Yuk uchew setae th Dipti ada te ts jenseni

Seta 5-C 5,6 branched, 10-C 3,4 branched. ........... 21

2E20), . BOR Bel OV SINGS aay se el aie al wh os (Le hewitti

Seta 5-IV, V usually triple and double respectively. ....... ae

22(21). Seta 8-C subequal to 9-C; 12-C 3,4 branched. ........ navalis

Seta 8-C distinctly longer than 9-C; 12-C double. . . . coerulescens

23(19). Seta 9-VIII single, 0.5-1.0 of paddle length............ 24

Seta 9-VIII 4 branched, 0.25 of paddle length. .... . curtipalpis

: sumatyranus

24(23). Most setae very strong and long; seta 5-C 3-4 times as long as

trumpet; 5-IV-VI 0.1-1.5 times as long as segment following;

9>VTt 0; Sof -paddie lenothiisn eas ep eet eo brevipalpus

and acutipalus

Most setae relatively weaker than above; seta 5-C 1-2 times

as long as trumpet; 5-IV-VI 0.5-1.0 as long as segment

following; 9-VIII 0.5 of paddle length........ eminentia

and lucavis

LARVAE!

1. Setae 2, 3-A placed subapically; habitats: tree holes, bamboos,

rock pools, leaf axils or sometimes ground pools (Vammilifer

Subbe nowy.) ort 66 RE LA a se ey 2

Setae 2,3-A placed apically; habitats: pitcher plants (Brevipalpus

Ser OU eee se a ee Ve Shige eT oe aS 19

1 incomptus and cvassicomus are unknown.

Pais

6(5).

7(6).

- 8(7).

9(6).

10(9).

11(10).

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Mammilifer Subgroup

Thorax and abdomen heavily spiculate, spicules large, setiform

and very distinct under. low mapnification: .....°. 2. 6 eee 3

Thorax and abdomen unspiculate or lightly spiculate, spicules

weak. dotlike. nol distinct as above, 62.0. yo. a i ee 5)

Spicules of thorax and abdomen branched; ventral surface of head

capsule with a transverse band of numerous spicules. .. kuhnsi

Spicules of thorax and abdomen unbranched; ventral surface of head

capsule without transverse bands of spicules. ......... 4.

Seta 5-C 3-5 branched; 7-P triple; posterior caudal margin of saddle

with numerous strong, spinelike spicules. ....... uniformis

Seta 5-C double; 7-P usually double; posterior caudal margin of

saddle lightly spiculate, spicules not as strong as above.

spiculosus

Setae 1-III-VI single; seta 8-P usually minute, inconspicuous, of the

same magnitude as.seta 14-P... 28 OS pholeter

Setae 1-III- VI double or more branched; seta 8-P strong, distinct,

nearly subequalto 7-P.. « 6.2 4k. a en ee 6

Seta 8-P double; head capsule, siphon and saddle pale yellowish

WHIEC: ao ieee yn Se a a a ee ee 7

Seta 8-P single; head capsule, siphon and saddle dark yellow to

POW ee es er ae ies Sk ae Se a ee a ee 9

eas se als es Be ee oe ee PO, eas Seca ei a 8

Seta 14-P double; 1-M longer than 3-M; 1-T longer than 2-T, of

the sammie maenitude as O-T yo. 6k eee Si oa bengalensis

Siphon with 3 pairs of subventral tufts, comb scales 30-50.

mammilifer, wardi and lasiopalpis

Siphon with 4 pairs of subventral tufts, comb scales 50 or more.

flavicornis

Setae 6-I, II double; siphon with 3 pairs (total 6) of subventral

PUES eres ee) hae hak Ge ea a te ee Woe a a aes Che eas fs 10

Setae 6-I, II triple or with 2 long, lateral branches and 1 weak median

branch; siphon with 4 pairs (total 8) of subventral tufts. .... i

Comb scales with round apical fringe of evenly fine spicules; thorax

and abdomen moderately spiculate......... bandoengensis

Comb scales with pointed apical fringe, or terminated into a strong

median spine; thorax and abdomen lightly spiculate. ..... 11

Pecten teeth barbed with graded denticles (Fig. 44). . .. bicornutus

Pecten teeth barbed with 1-3 strong basal denticles and 7, 8 graded

denticles distally (Pie. Ge). eh a kk aw sere he en minor

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region ca

12(9). Seta 1-VIII weak, short, less than 0.5 of 5-VIII, double or triple.

tuberis

Seta 1-VIII as long as or subequal to 5-VIII, 4,5 branched. ... 13

13(12). Seta 5-C 3,4 branched; thorax and abdomen usually lightly to moder-

ately spiculated, spicules visible under 10X objective. .... 14

Seta 5-C double; thorax and abdomen usually unspiculated or lightly

SPicu lakes gs hcg May Hetes wire ewer ae Se On ale Pas 15

14(13). Seta 6-VI usually single, longer than 6-III-V; posterior comb scales

with apical fringe of evenly fine spicules. ......... traubi

Seta 6-VI double, as long as 6-III-V; posterior comb scales with

apical fringe terminated into a strong median spine. ... Jlavatae

15(13). Subventral tufts of siphon dark, strong, 4-5 times as long as

siphonal width at points of attachment, usually 4 branched. . 16

Subventral tufts of siphon pale, weak, 1-2 times as long as siphonal

width at point of attachment, double ortriple. ......... 17

16(15). Seta 1-C usually swollen at middle or distally forked into 2,3 apical

spines; lateral setae of thorax and abdomen pale or moderately

pigmented; antennal shaft, siphon and saddle usually pale yellow.

ganapathi

Seta 1-C slender, single, spiniform; lateral setae of thorax and

abdomen dark; antennal shaft, siphon and saddle dark brown.

tmpostor

17(15). Head, siphon and saddle dark brown; siphon very long, index 14-19;

subventral tufts very weak, inconspicuous and widely spaced.

demissus

Head, siphon and saddle pale yellow; siphon relatively shorter,

index 7-9; subventral tufts distinct and more closely spaced. . 18

18(17). Seta 6-C single; spicules of antennal shaft weak and fine. . eukvrines

: Seta 6-C double; spicules of antennal shaft stronger. ... . peytoni

Brevipalpus Subgroup

19(1). Siphon longer than saddle; subventral tufts strong, prominent, 6-9

pains (otal 32-13 )6 i eee oe ee a ee 20

Siphon shorter than saddle; subventral tufts very weak, 3 pairs

(totae eos Ce See es OO ee ee curtipalpis

and sumatvranus

20(19). Siphon very slender and thin; seta 1-A strong, with 10-15 pectinate

branchese) eee Be eee ee ees ee ee Ba rae 21

Siphon broad and thick; seta 1-A weak, with3-6 simple branches. . . 23

21(20). Seta 5-C single or double; siphon brownish, lightly swollen in middle

orellipticabinshape: 4. &s St ee ee a Se 22

Seta 0-C 6-9 branched; siphon pale yellow, tubular, or gradually

tapered towards apexioy ip. OP A ee. gee, CO hewitti

78 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

22(21). Antennal shaft entirely pale or cream-colored; seta 5-C single.

navalis

Antennal shaft dark brown; seta 5-C double. ...... coerulescens

23(20). Seta 1-C dark, stout, spiniform; 6-C triple, subequal to 5-C, both in

normal position uae ies ae guage aera amar cra nett | err hheGtsk eee qensent

Seta 1-C pale, distally iene ca: 6-C single, longer than 5-C,

bot Diner Cleon ad. eS ee ae he ee ey ea es 24

24(23). Seta 7-II strong and dark, subequal to 7-I............. 25

Seta 7-II weak and pale, considerably shorter than 7-I. . lucaris

25(24). Seta 8-P usually double; siphon with 8-9 pairs of subventral tufts.

brevipalpus

Seta 8-P usually triple; siphon with 6-8 pairs of subventral tufts. .26

26(25). Ventral brush (4-X) usually with 5 pairs of setae; subventral tufts of

siphon usually Ot: PAalre ho... taki ae iw tee dee we aie flats acutipalus

Ventral brush (4-X) usually with 6 pairs of setae; subventral tufts of

siphonsially 1-9. Oairsers Sie PAI VS eminentia

Mammilifer Subgroup

The Mammilifer Subgroup essentially conforms to the description of Col-

less (1965: 264). It is chiefly characterized by the following characters: in

the adults by the erect scales on vertex of head entirely dark and the presence

of 1 lower mep bristle (except uniformis, bengalensis, flavicornis and lasio-

palpis); the female cibarial armature by cibarial bar with 30-40 subequally long

teeth (except pholeter); the male by palpus usually as long as or longer than

proboscis; the male genitalia by (1) basimere normal, slender, conical, with

4 or more strong submarginal setae in a prominent row or rows; (2) subapical

lobe as described for the group; leaflet g, well developed; (3) distimere slender,

strongly tapered and recurved towards apex and (4) internal process of the

phallosome usually strong, prominent or well developed; the pupa by trumpet

very long, more or less uniformly cylindrical, 0.6-0.8 mm in length and in

the larva by (1) setae 2,3-A placed subapically; (2) siphon slender, long, dis-

tally tapered, 4-7 times as long as saddle and (3) subventral tufts of siphon

3,4, pairs (total 6-8). Breeding habitats: usually tree holes, bamboos and

rock pools, sometimes leaf axils and ground pools.

DISCUSSION. The Mammilifer Subgroup is represented in the Oriental

region by 22 species. The majority of these have been described from South-

east Asia where the subgroup is better represented than anywhere else. The

subgroup is highly complex and taxonomically difficult mainly because many

of the forms involved are only slightly different from one another and show

considerable overlap in their breeding sites and distribution. The identity of

most species can be determined with certainty by examining the details of the

male antennae and genitalia, larval and pupal stages. Special caution should

be taken in using the characters given in the keys and descriptions.

To illustrate the evolutionary pattern as exhibited by the male antennal

modified tufts, palpus and genitalia, the 22 Oriental species of the Mammilifer

Subgroup are classified into 8 complexes: impostor, traubi, mammilifer,

ganapathi, minor, peytoni, pholeter and flavicornis. On the basis of the male

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 79

phallosome alone, the first 7 complexes appear to fall into 2 major lines: (1)

mammilifer line with tmpostor, traubi and mammilifer complexes and (2)

minor line with ganapathi, minor, peytoni and pholeter complexes. Each of

these complexes and its included species are briefly characterized as follows.

(1) impostor complex. This complex is represented only by impostor.

It is characterized in the male by: (1) palpus 0.75 of proboscis length and (2)

absence of modified tufts of scales and setae on the antennal flagellum; in the

male genitalia by (1) phallosome with internal process as long as or longer than

external process and (2) external process slender, narrow in tergal aspect and

with a broad, densely spiculose apical lobe.

(2) tvaubi complex. This complex is represented by traubi, uniformis and

lavatae, It is characterized in the male by: (1) palpus as long as or slightly

longer than proboscis and (2) modified tufts of scales and setae present on an-

tennal flagellomeres 6-8 or 5-8; in the male genitalia by (1) phallosome with

internal process longer than external process and (2) external process as in

ztmpostor complex.

(3) mammilifer complex. This complex includes mammilifer, wardi and

demissus, It is similar to the tvaubi complex in the male phallosome but dif-

fers from it in the presence of modified tuft of bristlelike setae on antennal

flagellomere 9 in addition to those on flagellomeres 5-8.

(4) ganapathi complex. This complex consists of ganapathi and spiculosus.

It is essentially similar to the tvaubi complex in the male palpus and antenna

but differs from it particularly in the following characters of the male phallo-

some: (1) lateral plate with internal process distinctly shorter than external

process; (2) external process broader in tergal aspect and (3) apical lobe of

external process small, placed sternad and with distinct basal stalk.

(5) minor complex. The minor complex is the largest, comprising 8

species, including minor, bicornutus, bandoengensis, tuberis, kuhnsi, crassi-

comus, incomptus and bengalensis. It is similar to the mammilifer complex in

having modified tufts of scales and setae present on antennal flagellomeres 5-9

and palpus as long as or slightly longer than proboscis but the male phallosome

resembles the ganapathi complex except for the apical lobe of the external

process, which is larger, with or without distinct basal stalk. |

(6) peytoni complex. This complex is represented by peytoni and eukrines.

It is differentiated from the minor complex by (1) absence of distinct modified

tuft on antennal flagellomere 5; (2) external process of phallosome larger and

broader; (3) internal process smaller, much shorter than external process

and (4) apical lobe of external process very minute, rudimentary and weakly

Sspiculate.

(7) pholeter complex. This complex includes only pholeter. It is similar

to the peytoni complex in the male phallosome, but differs from the latter in

having a distinct scale tuft on antennal flagellomere 5.

(8) flavicornis complex. This complex is represented by flavicornis,

lasiopalpis and vaghavanii. It is differentiated from all other complexes in

the male by (1) palpus longer than proboscis by more than the length of seg-

ment 5; (2) palpal segments 2 and 3 with prominent lateral rows of numerous

fine setae and (3) modified tufts present on antennal flagellomeres 5-9; in the

male genitalia by (1) distimere with a normal or modified, expanded apex; (2)

phallosome with internal process shorter than external process and (3) apical

lobe of external process very small or rudimentary.

80 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

ztmpostor complex

24. CULEX (LOPHOCERA OMYIA) IMPOSTOR NEW SPECIES

(Fig. 29)

FEMALE. Wing: 2.9mm. Forefemur: 1.5mm. Proboscis: 1.8 mm.

Small to medium sized species; in general conforms to the description of the

Mammilifer Group and Subgroup. Head, Decumbent scales in center of ver-

tex very narrow; linear and predominantly pale whitish; erect scales slender,

entirely black; lateral patch of broad scales pure white, very distinct. Palpus

slender, dark scaled, 0.2 of proboscis length. Proboscis dark, slender;

labial basal setae weak, 0.5 of palpal length. Cibarial Armature. Cibarial

bar with a close-set row of about 30 elongate teeth, 3,4 median teeth fine and

apically pointed, lateral teeth coarser and apically blunt or abruptly pointed.

Thorax. Mesonotal integument black; mesonotal scales narrow, entirely black

and moderately dense. Pleuron same color as mesonotum; 1 lower mep bristle

present. Legs, Anterior surface of hindfemur with a broad white stripe ex-

tending from base to apex. Wing. Plume scales on veins Ro, Rg narrow, line-

ar. Abdomen. Terga entirely dark or black; sterna pale whitish.

MALE (Fig. 29). Essentially similar to female in general external char-

acters. Palpus. Rather short, 0.75-0.80 of proboscis length; segments 4

and 5 strongly reduced in length, with only a few bristles and setae. Probos-

cis. Slender, uniformly thin; labial basal setae weak, short and pale. Antenna,

Pedicel with a spiculose nipplelike prominence projecting outwards on inner

dorsal surface; flagellar whorls weakly to moderately plumose, modified tufts

of scales and setae completely absent.

MALE GENITALIA (Fig. 29). As figured, essentially as described for the

Mammilifer Subgroup. Segment IX. Tergal lobe very small, bearing 2 weak

setae; sternum with 2 strong setae towards caudal margin. Basimere, Small,

slender, conical, 0.2 mm in length; inner tergal surface with a row of 4 or

sometimes 3, moderately strong submarginal setae which are rather widely

spaced. Subapical Lobe. Mesal surface with several weak setae at base of

setae d-f; rodlike setae a-c slender, gently curved, all terminated into a fine

hook apically; setae d-f consist of 3 acute blades, all apparently bent upwards;

leaflet go inform of a strong acute blade, distad of setae d-f; leaflet £, narrow,

lanceolate, as long as leaflet 99; seta h strong and long. Distimevre. Slender,

normal, as long as basimere; dorsal subapical crest of spicules not developed;

1 dorsal and 1 ventral tiny seta present, dorsal one rather inconspicuous; sub-

apical claw slender and short. Phallosome. Internal process of apical dorsal

lobe of lateral plate very strong and long, usually projecting beyond apex of

external process, its base provided with variable number of strong denticles;

external process narrow in tergal aspect, with 1,2 rows of 10-12 strong denticles

on tergal surface and a broad, densely spiculose apical lobe, its inner surface

with variable number of teeth. Pyroctiger, Apical crown large, composed of

4-7 flattened and apically blunt spicules laterally and numerous finer spicules

mesally; paraproct and cercal sclerite well sclerotized; cercal setae 2-4.

PUPA (Fig. 29). Abdomen: 2.7mm. Paddle: 0.62 mm. Trumpet: 0.60

mm; index 10-12. Cephalothorax and abdomen more or less uniformly brown.

Trumpet. Deep brown and uniformly cylindrical; meatus weakly spiculate.

Cephalothorax., Seta 1-C single or double; 5-C as long as 7-C, 3,4 branched;

8-C weak, subequal to 9-C, double. Metanotum, Seta 10-C weak, usually

single (1-2); 11-C strong, double; 12-C subequal to 10-C, double. Abdomen,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 81

Seta 5-IV double or triple, 1.5 times as long as segment following; 5-V single,

very long, about twice longer than segment following; 5-VI single, short, as

long as or shorter than segment following; 6-III-VI usually double (1-2); 9-VII

with 4 pectinate branches; 4-VIII single; 9-VIII with 8 pectinate branches.

Paddle. Very pale to almost transparent except for external basal margin of

outer part and midrib which are lightly pigmented.

LARVA. Head: 0.70 mm. Siphon: 1.8 mm; index 15. Saddle: 0.25 mm;

siphon/saddle ratio 8-9. In general as figured for traubi (Fig. 31). Head,

Pigmentation variable, generally yellowish, with pattern of dark brown areas,

seta 1-C dark, slender and moderately long; 4-C single, 2,3 times as long as

distance between bases of the pair; 5,6-C dark, as long as antenna, double.

Antenna shaft entirely brownish; spicules weak and rather sparse. Thorax.

Spiculation absent or poorly developed; seta 3-P single, about 0.5 of 1,2-P;

4-P double; 7-P double; 8-P single; usually subequal to 7-P, sometimes strong-

ly reduced to 0.25 of the latter; major setae of mesothorax and metathorax

dark. Abdomen. Spiculation absent; setae 6, '7-I and 6-II very dark and stout;

6-I, II triple; 7-I double; 6-III-VI usually double (2-3). Comb scales about 40,

in broad oval patch; posterior scales elongate, with fringe of evenly finespic-

ules. Saddle dark brown; posterior caudal margin very weakly spiculate; seta

2-X usually triple (2-3); ventral brush dark, with 6 pairs of branched setae.

Siphon. Very slender, dark brown, pecten teeth 9-10, each with 2,3 strong,

widely spaced basal denticles and an inconspicuous distal fringe of several fine

denticles; subventral tufts moderately long, widely spaced, double or triple

each, 2-4 times as long as siphonal width at point of attachment.

TYPE-DATA. Holotype “ (S 494-11) with associated pupal and larval skins

and slide of genitalia, Sapulut, Pensiangan, Sabah, MALAYSIA; tree holes,

rain forest, 600 feet (approx. 180 m), April 12, 1970, Sulaiman B. Omar and

Chia Y. Wang (USNM). Allotype 2 (S 488-12) with associated pupal and larval

skins (USNM); paratypes: 3c (S 486-10, S 488-13, S 494-12) with associated

pupal and larval skins; 9c, 42 (S 486, S 488, S 489, S 498), 1 lp (S 493-21),

same data as holotype, April 11-12, 1970 (USNM).

DISTRIBUTION. Known only from Sabah and Sarawak, Malaysia; 43 speci-

mens examined: 370%, 62, 1 1p; 5 with associated lp skins.

MALAYSIA. Sabah: Sapulut, Pensiangan (type-locality); 130°, 59, 6 lp.

Sarawak: Kapit; 240°, 12 (collected by K. J. Frogner, 1971).

TAXONOMIC DISCUSSION. Culex impostor is strongly differentiated from

all other members of the Mammilifer Subgroup in the male by the complete

absence of modified tufts of setae on the flagellomeres of the antenna. Because

of the absence of these antennal characters and the relatively short palpus, the

males of impostor are superficially similar to certain members of the subgen-

us Eumelanomyia. A detailed study of the male, including the genitalia, how-

ever, indicates that it exhibits characteristic features of the Mammailifer Sub-

group of Lophoceraomyia in having a distinct spiculose prominence on the an-

tennal pedicel and in the type of the phallosome. Among the members and

complexes of the Mammulifer Subgroup, impostor apparently shows a strong

affinity with the traubi and mammilifer complexes. It can be readily separated

from the members of these complexes in all stages as indicated in the keys and

as described above.

BIONOMICS. All of the adults and immatures of impostor were collected

from jungle tree holes at an elevation of about 180 m. It has been encountered

on several occasions and appears to be common in Sarawak and Sabah.

82 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

traubi complex

25. CULEX (LOPHOCERAOMYIA) TRAUBI COLLESS

(Figs. 30, 31)

Culex (Lophoceraomyia) traubi Colless 1965: 295 (o*, 9, L); Bram 1967a: 108

(o*, 2, L*).

FEMALE. Wing: 2.6 mm. Forefemur: 1.4 mm. Proboscis: 1.6 mm.

Small species, exceedingly similar to impostor, differing slightly in the fol-

lowing features. Head, Narrow decumbent scales of vertex broader on upper

eye margin, forming a distinct ocular line, scales in center of vertex narrow,

linear and predominantly dark; lateral patch of broad whitish scales broader,

more conspicuous. Labial basal setae of proboscis longer, about 0.75 of pal-

pal length. Cibarial Armature. Cibarial teeth narrower, finer, all with

pointed apices. Thorax. Mesonotal scales dark and somewhat glossy in ap-

pearance. Legs. Anterior surface of hindfemur largely pale whitish in basal

0.5, continued as broad pale stripe towards apex.

MALE (Fig. 30). As described by Colless (1965: 295) and Bram (1967a:

108); with the following diagnostic characters. Palpus, As long as or slightly

shorter than proboscis; segments 4,5 with only a few dark bristles. Proboscis.

Uniformly slender and thin from base to apex. Antenna. Flagellar whorls

moderately to strongly plumose; modified tufts of scales and setae present,

restricted to flagellomeres 5-8; F-5 with a small tuft of 5,6 narrow acute

golden scales, preceded dorsally by 2,3 long normal hairs, dorsalmost scales

longest, as long as the next 2 flagellomeres, rest gradually shorter and narrow-

er; F-6, 7 with moderately strong crumpled tufts of curled and twisted setae;

F-8 with a prominent tuft of 5,6 fused setae which are weakly curved subapi-

cally or not in form of a typical J-hook; F-9 apparently without any modified

setae.

MALE GENITALIA (Fig. 30). As figured. Segment IX. Tergal lobe with

3,4 weak and short setae; sternum with 2-8 strong setae in a transverse row

towards caudal margin. Basimere, Slender, small, about 0.22 mm in length;

inner tergal surface with a prominent row of 6-10 strong submarginal setae

which are closely spaced, parallel to tergomesal margin; marginal setae

rather strong, 15-20 in number. Subapical Lobe, Rodlike setae a-c subequal,

smoothly curved, largely overlapping in distal portion; setae d-f consist of 4

short pointed blades and 2 or more fine hairlike setae which are densely

packed; leaflet g5 in form of a long gently curved blade; leaflet g; broad, acu-

minate; setah moderately strong. Distimere. Dorsal subapical crest of fine

spicules absent. Phallosome. Very similar to impostor; internal spinelike

process very strong and long, projecting beyond apex of external process, its

base without any denticles; external process with a broad spiculose apical lobe

and 1-3 rows of several strong denticles on upper tergal surface, inner sur-

face without any denticles. Pyroctiger, Apical crown composed of numerous

dark, coarse spinelike spicules; cercal setae 3,4.

PUPA (Fig. 30). Abdomen: 2.2 mm. Paddle: 0.62 mm. Trumpet:

0.52 mm; index 8-10. Pigmentation of cephalothorax and abdomen variable

from yellowish to brownish. Trumpet. Moderately long, dark brown; meatus

strongly spiculate. Chaetotaxy as figured, the following setae are diagnostic.

Cephalothorvax. Seta 1-C usually double; 5-C usually triple, as long as 7-C;

8,9-C double. Metanotum. Seta 10-C double, as long as 11-C; 11-C single;

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 83

12-C usually double (2-3). Abdomen. Seta 5-IV usually triple (3-5), shorter

than segment following; 5-VI single or double, subequal to or shorter than 5-IV,

V; 6-I0 double; 6-IV-VI single; 9-VII usually triple (2-4); 4-VIII double; 9-VIII

6,7 branched. Paddle. Relatively narrow, entirely pale except for lightly

darkened midrib.

LARVA (Fig. 31). Head: 0.70 mm. Siphon: 1.4-2.0 mm (average 1.8

mm); index 9-11. Saddle: 0.30 mm; siphon/saddle ratio 5,6. As character-

ized for the Mammilifer Subgroup, with the following diagnostic features.

Head, Seta 1-C dark, slender, spiniform or sometimes lightly swollen at

base, with or without lateral accessory spines; 5-C usually triple (2-5); 6-C

usually double (2-3). Antennal shaft usually entirely dark brown, sometimes

lighter. Thovax, Lightly spiculate, spicules minute, rather inconspicuous;

seta 8-P strong, subequal to 7-P, single; 7-P double; 14-P double; 1-M as

long as 3-M, both setae single. Abdomen. Spiculation absent; setae 6-I, II

triple, 7-I double; 1-III-VI usually double (2-3); 6-III-V usually double (2-3);

6-VI usually single (1-2) and very long, 2 times as long as 6-III-V; 1, 4-VII

single; 7-VII usually 4 branched (3-4). Comb scales 32-45, posterior scales

longer than anterior ones, all with apical fringes of evenly fine spicules.

Saddle brownish, posterior caudal margin lightly to moderately spiculate;

seta 2-X double; anal gills at least 2 times as long as saddle. Siphon, Slender,

yellowish to dark brown; length extremely variable; pecten teeth 12, slender,

each tooth with 1,2 large basal denticles and 4, 6 fine distal denticles, latter

rather inconspicuous; subventral tufts strong, 4 pair in number, first 2 proxi-

mal pair double or triple, 4,5 times as long as siphonal width at point of at-

tachment; next 2 distal pair much shorter, 4-6 branched each, 1,2 times as

long as siphonal width; median caudal filament of spiracular apparatus not

developed or absent.

TYPE-DATA. Holotype “ with associated pupal and larval skins (0453/C44),

slides of antenna (CH 162) and genitalia (CT 586), Ulu Gombak, Selangor,

Malaya | (Peninsular Malaysia), MALAYSIA], 10 May 1958, D. H. Colless

(ANIC).

DISTRIBUTION. Thailand, Malaysia and Indonesia. 105 specimens exa-

mined: 530°, 402, 12 L; 68 with associated immature skins (12 p, 56 lp).

THAILAND. Mae Hong Son: Doi Chang; Nakhon Nayok: Khao Yai; Ranong:

Koh Chang; Trang: Muang; 4[, 5 L, 2p, 2 lp.

MALAYSIA. Peninsular Malaysia - Selangor: Ulu Gombak; Ulu Langat;

Ampang, F. R., Bt. Ulu Bakau; Kg. Tanjong Robok; Bt. Kutu; Selangor-

Pahang: The Gap; Pahang: Gunong Benom; Cameron Highlands; Perak: Kg.

Jalong; Perlis; Bt. Bintang F. R.; 230, 279, 7L, 9p, 291lp. Malaysia (N.

Borneo) - Sabah: Tawau; Mt. Kinabalu; Beaufort; Tenom; Pensiangan; 25,

13933 p,; 25 Ip. |

INDONESIA. Sumatra: Bengkoelen; 1% (Thienemann, 28 Jan. 29).

TAXONOMIC DISCUSSION. Culex traubi is one of the most common spe-

cies of the Mammilifer Subgroup and is probably more widespread than the

above records indicate. It can be recognized in the male by (1) the absence of

modified setae on antennal flagellomere 9; (2) the form of the modified tuft of

antennal flagellomere 8 and (3) the small, yellowish or golden tuft of 5, 6

narrow, acute scales of antennal flagellomere 5; the male genitalia by (1) the

very long internal process of the phallosome; (2) the prominent row of 6-10

strong submarginal setae of the basimere and (3) the gently curved and largely

overlapped setae a-c of the subapical lobe; the pupa by (1) the dark brown and

strongly spiculate trumpet; (2) the single seta 11-C and (3) the relatively short

setae 0-IV-VI which are usually 3,2 and 1-2 branched respectively, and in the

84 Contrib. Amer. Ent: Inst:,°vol.. 13; no. 4; 1977

larva by (1) seta 5-C usually triple; (2) setae 1-III-VI usually double; (3) the

usually single and long seta 6-VI; (4) the strong subventral tufts of the siphon

and (5) the usually dark brown pigmentation of the saddle and siphon.

The tvaubi larva is very variable in the length and color of the siphon and

branches of abdominal setae 1-III-VI, but these have not been found to be cor-

related with any differences in the male and pupa, indicating that probably only

a single form is involved.

The relationship of tvaubiis apparently closest to uniformis from Sri Lanka

and lavatae from the Philippines on the basis of the male phallosome and anten-

na, and together with the latter 2 species appears to fall into a complex. It

can be readily separated from uniformis and lavatae as indicated in the keys

and as inthe descriptions of the latter 2 species.

BIONOMICS. The immatures of trvaubi have been most frequently collected

from tree holes in the jungle at a broad range of elevation from 70-1, 700 m.

In several recent collections in Malaysia, they were also found breeding in bam-

boo stumps or fallen bamboos lying on the ground as frequent as in tree holes

without showing preference for either type of breeding habitat. All of the lar-

val collections from Thailand and most of the collections from Sabah came from

tree holes. The majority of the adults came from rearing the pupae and larvae

and only a few from sweeping among leaves of plants near the breeding sites.

The adults possibly attack man on occasion (Colless 1965: 296). No data are

available on their feeding behavior.

26. CULEX (LOPHOCERAOMYIA) UNIFORMIS (THEOBALD)

(Figs. 32, 33)

Lophoceratomyia uniformis Theobald 1905a: 245 (o*, 9).

Culex (Lophoceratomyia) uniformis (Theobald), Edwards 1922a: 281 (0, key);

Barraud 1924: 24 (“, 2); Barraud and Covell 1928: 678 (2 buccopharyngeal

armature); Edwards 1928: 276 (L): Edwards 1932: 197 (taxonomy); Bar-

raud 19345 3878 (o*, 2

Culex (Lophoceraomyia) uniformis (Theobald), Mattingly 1949: 226 (L, key);

Stone, Knight and Starcke 1959: 235 (catalog).

FEMALE. Wing: 3.2 mm. Forefemur: 1.6 mm. Proboscis: 1.9 mm.

Medium-sized species; in general as described for tyvaubi, differing from it in

the larger size and in the following characters. Head, Vertex with a broader

ocular line of pale decumbent scales contrasting sharply with dark scales in

center and the posterolateral area. Palpus and proboscis relatively thicker.

Cibarial Armature, Cibarial teeth stronger; lateral teeth with blunt or truncate

apices. Thorax, Lower mep bristle absent.

MALE (Fig. 32). Differing from tvaubi as described for the female and in

the following features. Palpus, Longer, exceeding proboscis by the full length

of segment 5; segments 4 and 5 with more bristles on lateral and mesal sur-

faces. Antenna. Flagellar whorls moderately long plumose; F-9 with a larger,

more conspicuous tuft of 6,7 scales, 1-3 dorsalmost scales longest, dark, dis-

tally broad and apically pointed or blunt, as long as the next 3 flagellomeres,

followed laterally by 4,5 very narrow, curved, acute, yellowish scales of de-

creasing length; F-6,7 with stronger tufts of curled setae; modified tuft of F-8

apparently thicker and distinctly curved in apical portion.

MALE GENITALIA (Fig. 32). As figured, differing from tvaubz in the fol-

lowing features. Basimere, Inner tergal surface with a row of 4 strong sub-

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 89

marginal setae; marginal setae sparse and weak. Subapical Lobe, Leaflet g,

much larger, longer and striated. Phallosome. Apical lobe of external

process of lateral plate more finely spiculose, denticles on tergal surface of

external process weaker and apparently fewer.

PUPA. As figured and described for trvaubi, differing from it in the follow-

ing features. Trumpet, Relatively shorter and stouter, about 0.5 mm in

length, index 8. Cephalothorax, Seta 1-C stronger, as long as 5-C, usually

triple; 5-C usually 5,6 branched (3-6); 8-C double or triple, sometimes more

branched. Metanotum. Seta 11-C double. Abdomen, Setae 5-IV, V 1.5-2.0

times as long as segment following; 5-IV usually double (2-3); 5-V single or

double; 5-VI single, as long as segment following; 6-IV-VI weaker, usually

double (1-2).

LARVA (Fig. 33). Head: 0.72 mm. Siphon: 1.6 mm; index 9. Saddle:

0.35 mm; siphon/saddle ratio 4,5. As figured; strongly differentiated from

traubi in having thorax and abdomen densely spiculose and in the following

features. Head, Most setae stronger, more or less flattened and dark; seta

4-C flattened, pectinate; 5-C 3-5 branched; 6-C double or triple, sometimes

4 branched. Antenna shaft pale cream-colored. Thorax. Spicules strong,

hairlike, unbranched and extremely dense all over; seta 7-P triple; 8-P minute,

rather inconspicuous, 4,5 branched; 1-M stronger and longer than 3-M. Abdo-

men. Spiculation as in thorax; setae 1-III-VI strong, dark, flattened, 4,5

branched; 6-III-VI triple. Saddle densely spiculate; lateral caudal margin with

numerous strong spines; seta 2-X 3,4 branched. Siphon. Moderately long,

pigmentation yellowish; pecten teeth 12-14, larger teeth with 2,3 graded basal

denticles and 1,2 finer distal denticles, latter sometimes not developed; sub-

ventral tufts subequally strong, 4, 6 branched each; first proximal pair placed

among pecten, 2,3 times as long as siphonal width at point of attachment, the

remaining pairs gradually shorter.

TYPE-DATA. Lectotype “* with attached genitalia mount and slide of

head, Peradeniya, CEYLON | SRI LANKA], May 1902, collected by Green

(BMNH; present selection).

DISTRIBUTION. Known only from India and Sri Lanka. 452 specimens

examined: 216, 230%, 6 L; 7 with associated immature skins (1 p, 6 lp).

INDIA. Bombay: Trombay; Malabar; Pudupadi; 2.

SRI LANKA. Central Province: Kandy District, Peredeniya (type-locality); .

Madugoda; Udawattekele; Wakarwatte; Uva Province: Monaragala District;

Southern Province: Kanneliya; Western Province: Singharaja Forest; Mora-

pitiya; Kalatuwawa; Labugama Reservoir; 214, 2302, 6 L, 1p, 6 lp.

Additional records from the literature. INDIA: Andaman Islands; Madras,

Nilgiri Hills (Barraud 1924: 43).

TAXONOMIC DISCUSSION. Culex uniformis appears to be restricted to

south India and Sri Lanka and probably does not occur anywhere else in the

Oriental region. In Sri Lanka it is one of the 4 known representatives of the

Mammilifer Subgroup. Of all stages of uniformis, the larva is very diagnostic,

especially in the development of the remarkably strong and dense spicules on

the thorax and abdomen. The only other species which exhibit strong spicula-

tion similar to the uniformis larvae are spiculosus and kuhnsi, but detailed

study has shown that each is distinct and can be readily separated from the

others by features indicated in the key and descriptions. Because of the simi-

larity in their larvae, uniformis has been confused in the past with spiculosus

and kuhnsi, The records of uniformis by Delfinado (1966: 114) and Baisas

(1974: 121, as subspecies mercedesae) from the Philippines are, in fact,

those of kuhnsi and the record by Chu (1958: 109-13) from Hainan Island, China,

86 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

is probably that of spiculosus.

BIONOMICS. Culex uniformis is apparently the most common Lopho-

cevaomyia species in Sri Lanka. All larval collections from this area came

from jungle tree holes. In India, it was collected from rock pools (Barraud

1934: 374).

AT CULEX (LOPHOCERA OMYIA) LAVA TAE STONE AND BOHART

(Fig. 32)

Culex (Lophoceraomyia) lavatae Stone and Bohart 1944: 220 (o"*); Delfinado

1966: 107 (o'*, 9); Baisas 1974: 112 (o*, 2, P*, L*).

Culex fidelis Dyar 1920: 180 (in part).

FEMALE. Wing: 2.8mm. Forefemur: 1.4mm. Proboscis: 1.8 mm.

Extremely similar to tvaubi, uniformis and other members of the Mammilifer

Subgroup, differing particularly in the following characters. Cibarial Arma-

ture. Cibarial teeth more numerous, about 40; 10 median teeth narrow, with

finely pointed apices, lateral teeth coarser with abruptly pointed or blunt

apices. Thorax, Mep with a row of 5,6 pale, weak, hairlike setae adjacent to

posterior border of sip, clearly dorsad of lower mep bristle; ppm with a few to

several scattered, narrow, linear, pale scales cephalad of posterior bristles.

Legs, Wing and Abdomen, Essentially similar to traubi,

MALE (Fig. 32). In general as described for female. Palpus, Usually

as long as proboscis, sometimes slightly shorter or longer; segment 3-5 with

a few short bristles on its apices. Antenna, Flagellum moderately to densely

long plumose; modified tuft of setae present only on flagellomeres 5-8; F-95

with a minute inconspicuous tuft of 3 very short yellowish setae among dense

whorl of long hairs; F-6, 7 with poorly developed tufts of few weak curled

setae; F-8 with a prominent tuft of 4,5 stout, long, gently curved setae which

are proximally fused, distally spread out, as long as the combined length of 2-3

flagellomeres.

MALE GENITALIA (Fig. 32). As figured; exceedingly similar to fraubi in

the phallosome, proctiger and distimere, differing particularly in the following.

Basimere, Inner tergal surface with a row of 5,6 submarginal setae; marginal

setae very weak and rather sparse. Subapical Lobe, Accessory setae in group

d-f stronger, more flattened, 2 with distinct serrate margin in subapical por-

tion; leaflet g; short, narrow, lanceolate.

PUPA. Abdomen: 2.7mm. Paddle: 0.70mm. Trumpet: 0.55 mm;.

index 10. As figured for tvaubi (Fig. 30). Cephalothorax and abdomen deep

yellow to brown. Trumpet, Dark brown and moderately long. Cephalothorax,

Seta 1-C usually triple (2-3); 5-C usually 4-branched (3-5); 8, 9-C usually

double (1-2). Metanotum, Seta 10-C usually double (2-3); 11-C double; 12-C

double or triple. Abdomen. Seta 5-IV usually triple (3-4), 1.0-1.5 times as

long as segment following; 5-V double, as long as 5-IV; 5-VI double, 0.9 of

the length of segment following; 6-III, IV usually double; 6-V-VI single; 9-VII

usually 4 branched (3-5); 9-VIII 7-10 branched. Paddle, Pale whitish to al-

most transparent except for yellow midrib and external margin of outer part.

LARVA. Head: 0.65 mm. Siphon: 1.7 mm; index 10. Saddle: 0.34 mm;

siphon/saddle ratio 4-6. Very similar to and as figured for tvaubi (Fig. 31),

differing from it particularly in the following. Head, Seta 1-C more slender,

with or without accessory lateral spines. Thovax, Lightly spiculate, spicules

apparently stronger and denser, more distinct. Abdomen, Setae 1-III-VI

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 87

stronger, 0.5-1.0 of 6-III-VI; 6-III, IV usually triple (2-3); 6-V double or

triple; 6-VI usually double (1-2). Comb scales more numerous, 50 or more.

Saddle with several strong sharp spicules on posterior caudal margin. Siphon,

Subventral tufts 4,5 branched each, first proximal pairs strongest, about 2

times as long as siphonal width at point of attachment, next 3 pairs gradually

shorter and weaker.

TYPE-DATA. Holotype “*, Los Banos, (Luzon), PHILIPPINES, 28 Jul

1915 (USNM; Cat. No. 56984).

DISTRIBUTION. Malaysia (N. Borneo) and the Philippines. 91 specimens

examined: 50, 372, 34 L; 57 with associated immature skins (3 p, 54 lp).

MALAYSIA. Sabah; Sandakan Bay, Sapagaya Lumber; 1.

PHILIPPINES. Luzon: Los Banos (type-locality); Baguio; Munoz (central

Luzon); San Jose; Subic Naval Base; Zambales; Mt. Makiling; Laguna; Dalton,

N. Vizcaya; Mindoro: San Jose; Palawan: Princesa; Mt. Malino; Irahnan

River; Samar: Osmena; Leyte: Tacloban; Jolo Island: Indanan; 490", 379,

34 L, 3p, 54 lp.

TAXONOMIC DISCUSSION. Culex lavatae is one of the most common

Lophoceraomyia in the Philippines. The additional new record of lavatae from

Sabah based on a male is probably correct as this specimen agrees well with

those from the Philippines in every feature of the antenna and genitalia. Al-

though lavatae is widespread among the Philippine Islands, there is, however,

no indication of local differentiation among the material I have examined.

All stages of lavatae can be readily separated from other members of the

Mammilifer Subgroup as indicated in the above diagnosis and in the keys. Its

affinity is closest to trvaubi from which it can be differentiated in the male

antenna by the reduction in the size and the number of setae of the modified

tufts of flagellomeres 5-7, in the pupa by the double seta 11-C and in the

larva by the double seta 6-VI and the stronger spicules on the posterior caudal

margin of the saddle. The adults of both sexes of lavatae are distinct from all

other members of the Mammilifer Subgroup in the presence of a row of weak,

hairlike setae dorsad of the lower ™ep bristle.

BIONOMICS. The immatures of lavatae largely came from the collections

made in tree holes in the mountain areas of the Philippine Islands. In Luzon,

larvae have also been found in discarded tin cans; wood barrel and used tires.

The elevation ranges from 100 to over 1000 m. Most adults from the Philip-

pines were obtained from rearing the pupae and larvae. The single male from

Sabah, Malaysia, apparently came from a general field catch.

mammilifer complex

28. CULEX (LOPHOCERAOMYIA) MAMMILIFER (LEICESTER)

(Figs. 34, 35)

Lophoceratomyia mammilifer Leicester 1908: 128 (c, 9).

Culex (Lophoceratomyia) mammilifer (Leicester), Edwards 1922a: 281 (<,

key); Barraud 1924: 43 (o*); Edwards 1928: 276 (L); Edwards 1932: 198

(taxonomy); Barraud 1934: 374 (o'*, L); Baisas 1935: 174 (o°*).

Culex (Lophoceraomyia) mammilifer (Leicester), Bohart 1945: 75 (distribu-

tion); Mattingly 1949: 227 (L, key); Colless 1965: 287 (o*, 9, L*);

Delfinado 1966: 109 (“*, P*, L*); Bram 1967a: 93 (o°*, L*, P); Baisas

1974: 114 (“, 2, P, L).

Culex (Lophoceratomyia) chiungchungensis Hsu 1963: 231 (o*, L*); Colless

88 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

1965: 287 (synonymy).

FEMALE. Wing: 2.8-3.0 mm. Forefemur: 1.3-1.4 mm. Proboscis:

1.6mm. Essentially conforming to the Mammilifer Group and Subgroup;

usually small, slender species, with the following diagnostic features. Head,

Vertex with rather broad, ovate, pale, decumbent scales forming distinct ocular

line anteriorly; decumbent scales in center narrow, linear and usually entirely

dark; erect scales slender, entirely dark; lateral patch of broad scales whitish,

very distinct. Strong labial basal setae of proboscis about 0.9 of palpal length.

Cibarial Armature (Fig. 34). Cibarial bar with a concave row of about 34 elon-

gate teeth, 4 median teeth fine and apically pointed, lateral teeth coarser,

apically blunt or abruptly pointed. Thorax, One lower mep bristle present.

Legs, Anterior surface of hindfemur with a broad pale stripe extending from

base to almost at apex. Abdomen, Terga entirely blackish,sterna pale whitish.

MALE (Fig. 34). In general as described for female. Palpus, Longer

than proboscis by 0.5 to full length of segment 5; segments 4, 5 upturned, with

a few short bristles on lateral and mesal surfaces. Pyroboscis, Very slender

and thin. Antenna, Flagellum moderately plumose; modified tufts of scales

and setae present on flagellomeres 5-9; F-5 usually with a distinct tuft of 6-8

dark scales, 3,4 dorsalmost scales broad, acuminate or apically blunt, as

long as the next 2,3 flagellomeres, followed laterally by 3,4 narrow, shorter,

acute scales of decreasing length, sometimes composed of narrow acute

scales only; F-6,7 with strong, crumpled tufts of curled setae; F-8 witha

typical J-hooked tuft of 6,7 fused setae, some of which are distinctly swollen

before tapering into sharp recurved points; F-9 with a prominent tuft of 5-7

dark, long, flattened, bristlelike setae.

MALE GENITALIA (Fig. 34). Segment IX. Tergal lobe small, bearing 3

short and weak setae; sternum with an irregular row of 7, 8 strong setae to-

wards caudal margin. Basimere, Inner tergal surface with a prominent row

of 5-8 strong, sinuous submarginal setae; marginal setae 4-6, all weak, short,

widely spaced. Subapical Lobe, One of the 3 rodlike setae a-c usually curved

in middle, separating from the other 2 rods by a distinct gap, sometimes

straight and overlapping with other 2 rods; setae d-f consist of 1 curved

blade and 2,3 hairlike setae; leaflet go in form of a long, stout blade or rod;

leaflet gj broad, acuminate; seta i strong. Distimere, Slender, with distinct

crest of fine spicules extending from apex to near middle of curvature; ventral

tiny seta present, dorsal one absent; subapical claw short and small. Phallo-

some. Strongly resembling zmpostor and members of the traubi complex;

internal process of lateral plate long, projecting beyond apex of external pro-

cess; external process slender in tergal aspect, with only a few weak denticles

on lower tergal surface, sometimes practically bare; apex with a spinose lobe

which is broad in lateral aspect and apparently not constricted at base. Proc-

tiger. As figured, essentially similar to other members of the Mammilifer

Subgroup; cercal setae 3, 4.

PUPA (Fig. 34). Abdomen: 2.4mm. Paddle: 0.60 mm. Trumpet: 0.65

mm; index 12-15. Cephalothorax and abdomen yellowish white or cream-

colored, rather paler than most forms. Trumpet, Darkened, uniformly

cylindrical and relatively long; pinna with distinct slit extending to meatus.

Complete chaetotaxy as figured, the following setae are diagnostic. Cephalo-

thorax, Seta 1-C usually 5-branched (5-6); 5-C weaker than 7-C, usually 6

branched (5-8); 8-C usually double (1-2), 9-C usually triple (2-3). Metanotum,

Setae 10,11-C double, 11-C sometimes single. Abdomen, Seta 5-IV usually

5 branched (4-7), shorter or as long as segment following; 5-V, VI usually

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 89

double (1-2), as long as or slightly longer than segment following; 6-III-VI

usually 4 branched (3-7); 9-VII usually triple (2-3); 4-VIII usually triple (2-3);

9-VIII usually 6 branched (5-8). Paddle, Pale whitish to almost transparent,

midrib lightly to moderately infuscate.

LARVA (Fig. 35). Head: 0.70 mm. Siphon: 1.2-1.9 mm; index usually

10,11 (7-12). Saddle: 0.27 mm; siphon/saddle ratio 5. Complete chaetotaxy

as figured; distinctive particularly in the follownmg. Head, Integument varying

from pale whitish to yellowish; seta 1-C dark, slender, simple, spiniform;

4-C single, 1.5-2.0 times as long as distance between bases of the pair; 5-C

usually double (2-3); 6-C double. Antennal shaft entirely pale except for basal

dark ring. Thorax, Spiculation absent or not developed; seta 7-P usually

double (2-3); 8-P double, subequal to 7-P; 14-P single; 1-M single, as long as

or longer than 3-M; all major setae in pleural group lightly or moderately pig-

mented. Abdomen, Spiculation absent; setae 6-I, II triple; 7-I double; 1-III-VI

usually triple (3-4); 6-III usually 4 branched (3-4); 6-IV-VI 3, 4 branched,

sometimes 9. Comb scales 30-40, more or less subequal in size, each with

rounded apical fringe of evenly fine spicules. Saddle pale yellowish, some-

times darker; seta 2-X with 1 short and 1 long branch; 4-X with 6 pairs of

setae; anal gills about as long as saddle. Siphon. Relatively long and thin;

pigmentation varying from pale whitish to light yellowish, sometimes darkened

in the middle; pecten teeth 10-16, larger teeth barbed with 6, 7 graded denticles,

basal denticle not distinctly differentiated; subventral weak, subequal, usually

3 pairs, sometimes 4; single or double each, as long as or slightly longer than

siphonal width at point of attachment; median caudal filament of spiracular

apparatus developed.

TYPE-DATA. Lectotype “* with slide of antenna and genitalia, Raub,

Pahang, Malaya | MALAYSIA], larva from pools in jungle path, 5 Jan 1904,

G. F. Leicester (BMNH; selection of Colless 1965: 287).

DISTRIBUTION. India, Sri Lanka, China, Thailand, Malaysia, Indonesia

and the Philippines. 200 specimens examined: 1190, 499, 32 L; 72 with

associated immature skins (13 p, 59 lp).

SRI LANKA. Suduganga, 20, 19.

THAILAND. Chiang Mai: Doi Sutep, Suan Kwi Nin; Chiang Dao Mtn. ;

Lampang: Ngao; Ban Rong Na; Ban Pha Daeng; Nakhon Ratchasima; Ban Tha

Maprang; Kanchanaburi: Huai Mae Nam Noi; Ban Sai Yok; Huai Bong Ti; Khao

Sung; Chanthaburi: Khao Sai Dao; Nakhon Si Thammarat; Chaung Khao; Ranong:

Khao Hin Chang; Phuket: Khao Prathin; Ban Khian; Yala: Yala Bong; Navathi-

wat: Khao Lau; 300, 59,18 L,. 9p, 12 Ip.

MALAYSIA. Peninsular Malaysia - Selangor: Rantau Panjang; Ulu Gom-

bak; Pahang: Gunong Benom; Merapoh; Mela K. Lipis; Pevak: Chior F. R.;

22%, 162, 2p, 6 lp. Malaysia (N. Borneo) - Sarawak: Kuching; Sabah: Tawau;

Kota Kinabalu; Sandakan Bay, Sepilak F. R.; Beaufort; 140°, 89, 5 lp.

INDONESIA. Sumatra: Bengkoelen; Java: Djakarta; 6.

PHILIPPINES. Luzon: Subic Naval Base; Malawin Creek, Mt. Makiling;

Mindoro: San Jose; Palawan: Iwahig; Iraknam; Balshahan River; Leyte:

Dagami; Lagolago Baybay; Samar: Osmena; Mindanao: Gingoog, Ori Missa-

mis; Lanao, Kolambugan; Davao; 450, 192, 5L, 1p, 36 lp.

Additional records from the literature. INDIA, North Bengal; Andaman Is. ;

Malabar Coast (Barraud 1934: 374); CHINA, Hainan Island (as chiungchungen-

sis, Hsu 1963: 231).

TAXONOMIC DISCUSSION. Culex mammilifer is one of the most common

members of the Mammilifer Subgroup and is apparently widespread throughout

Southeast Asia and adjacent areas. It can be readily separated from other

90 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

members of the Mammilifer Subgroup in the male, pupa and larva as indicated

in the keys and as described above. The female is difficult to distinguish, but

with practice it can be recognized by the predominantly broad, pale decumbent

scales on the anterior margin of the vertex and by the presence of a lower mep

bristle.

Most of the mammilifer males from various parts of Southeast Asia essen-

tially agree with the type and topotypic material from Malaysia except in the

Philippines where they are very variable and appear to be differentiated into 2

distinct local forms. One of these corresponds well with the typical form in

having the modified tuft of antennal flagellomere 5 consisting of some dark broad

scales and modified tuft of antennal flagellomere 9 consisting of 5 normal bris-

tlelike setae. This form is apparently widespread throughout the Philippines.

The other is known only from Palawan island. It differs markedly from the

type form in having only narrow acute scales in the modified tuft of antennal

flagellomere 5 and in having modified tuft of antennal flagellomere 9 composed

of 6,7 very strong, flattened bristlelike setae. In the male genitalia, the Pala-

wan form differs slightly from the typical form only in having the rodlike setae

a-c of the subapical lobe straight and largely overlapping in the distal portion.

No associated immature stages of the Palawan males are available for further

comparison. Thus, the Palawan form is provisionally treated here as a geo-

graphical segregate of mammilifer although there is a possibility that it may

represent a distinct species.

BIONOMICS. Culex mammilifer is one of the few members of the Mammi-

lifer Subgroup which utilizes ground pools as its principal breeding habitat.

The pupae and larvae have been frequently collected from small pools, particu-

larly puddles and foot prints along stream margins under heavy forest shade.

On occasions, they have also been found in rock pools, leaf axils of nipa palm

and cut bamboos or bamboo stumps lying close to the ground. These habitats

were reported from a broad range of elevation. In Malaysia most of the larval

collections came from near the tidal zone along the coast and only a few from

inland forests (Colless 1965: 289). The adults mostly came from rearing

larvae or pupae but some were also caught by sweeping amongst foliage near

the breeding sites. The females are not known to bite man.

29. CULEX (LOPHOCERA OMYIA) WARDI NEW SPECIES

(Pie. 3G)

FEMALE. Exceedingly similar to mammilifer, differing slightly in the

following features. Cibarial Armature, All cibarial teeth finer and distally

tapered into sharp apices. Legs, Anterior surface of hindfemur with a

broader longitudinal whitish stripe from base to apex.

MALE (Fig. 36). In general as in female, differing from mammilifer par-

ticularly in the following characters. Palpus, Shorter, as long as or slightly

shorter than proboscis; segments 4,5 reduced in length, segment 4 about 0.9

of segment 5. Proboscis., Apical 0.50-0.75 with prominent rows of several

short setae on lateral and ventral surfaces. Antenna, F-5 with a minute and

rather inconspicuous tuft of 5-7 short, yellow acute scales, as long as next 2,

3 flagellomeres, followed ventrally by a brushlike tuft of 3,4 long normal

setae; F-9 with a dark tuft of about 7 strong bristlelike setae.

MALE GENITALIA (Fig. 36). Extremely similar to mammilifer in the

phallosome and proctiger, differing from it in the following features. Basi-

mere. Inner tergal surface with a dense double row of 10-12 strong submar-

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 91

ginal setae. Subapical Lobe, Roda strongly curved at middle, separating

from rods 0, c by a wide gap; setae in group d-f absent or represented by one

short fine seta; leaflet 89 rodlike, wavy in outline, with apex terminating into

a sharp point; leaflet 81, moderately broad. Distimere, Median curvature

with a distinct hump on dorsal margin; subapical dorsal surface with more

distinct crest of strong spicules.

PUPA. As described and figured for mammilifer, differing from it particu-

larly in having apex of paddle slightly produced into a blunt point and generally

in darker trumpet, cephalothorax and abdomen.

LARVA. Indistinguishable from mammilifer.

TYPE-DATA. Holotype “ (212-112) with associated pupal and larval skins

and slide of antenna, palpus, proboscis and genitalia, Kanneliya, Galle District,

Sabavagamuwa Province, SRI LANKA, rock pool, elevation 245 m, 9 July 1975,

FE. L. Peyton and Y.-M. Huang (USNM); Allotype 2 (212-6) with associated pupal

and larval skins, same data as holotype (USNM); Paratypes: 2 lpo (212-5, 10);

9 po (212-100, 105, 106, 108, 109); 5 Ip? (212-3, 7, 8, 9, 12); 2 p? (212-103,

107), same data as holotype and allotype; 1 lpo’ (190-1); 2 lpo’ (197-2, 5); 2 po

(197-100, 103); 1 Ipc (197-3); 2 p? (197-101), same locality and collectors as

collection No. 212, 8-9 July 1975 (to be deposited in BMNH, Bishop Museum

(BPBM) and USNM). This species is named in honor of Dr. Ronald A. Ward,

Walter Reed Army Institute of Research, Washington, D. C.

DISTRIBUTION. Known only from Sri Lanka. 75 specimens examined:

390°, 362, 75 with associated immature skins (31 p, 44 Ip).

SRI LANKA. Sabaragamuwa Province: Galle Dist., Kanneliya; Kalutara

Dist., Morapitiya; Ratnapura Dist; Vaddgala; Western Province: Colombo

Dist., Labugama; Waga; 390°, 369, 31 p, 44 lp.

TAXONOMIC DISCUSSION. Culex wardi is known from several specimens

recently collected in the southern part of Sri Lanka. It is most closely related

to mammilifer but is distinct from it in features of the male antenna, palpus

and genitalia as indicated above. These differences are striking and constant

among the material Ihave examined. The female and pupa exhibit slight dif-

ferences as noted above, but the larva is indistinguishable from that of mam-

milifer.

BIONOMICS. This is apparently one of the common species of the sub-

genus in Sri Lanka. As in mammilifer, the immatures have frequently been

collected in ground pools and rock pools under heavy shade in forests. On

one occasion, it was also reported from a bamboo stump. Collections were

made at elevations from 200 to 300 m.

30. CULEX (LOPHOCERAOMYIA) DEMISS US COLLESS

(Figs. 37, 38)

Culex (Lophoceraomyia) demissus Colless 1965: 296 (0'*).

Culex (Lophoceraomyia) fuscosiphonis Bram and Rattanarithikul 1967: 14

(o*, 2, L*); Bram 1967a: 87 (“*, 9, L*). NEW SYNONYMY.

FEMALE. Wing: 3.0mm. Forefemur: 1.5 mm. Proboscis: 1.7 mm.

Exceedingly similar to traubi, uniformis and mammilifer, differing slightly

in the following features. Head, Vertex with 1,2 rows of narrow clavate

_ decumbent scales which are predominantly pale, forming a narrow ocular

line; lateral patch entirely dark or sometimes grayish. Cibarial Armature.

As in mammilifer; cibarial bar with 30-32 teeth. Thorax. Upper surface of

92 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

ppn with some scattered dark narrow scales cephalad of posterior bristles.

One lower mep bristle present, weak and pale. Abdomen, Sterna dark gray-

ish or slightly paler than terga.

MALE (Fig. 37). In general as infemale. Palpus, As long as proboscis.

“Antenna, Flagellum moderately plumose; modified tufts of setae present on

F-5-9; F-5 with a small and inconspicuous tuft of 3,4 short acute setae, as long

as 1 flagellomere in length, preceded dorsally by 1 long normal seta and 2

other fine setae; F-6, 7 with weak crumpled tufts of curled and apically twisted

setae; F-8 with a smoothly curved tuft of 6 dark, fused setae; F-9 with 3-6 dark,

bristlelike setae; F-10 with or without 1 long, hairlike seta distad of normal

whorls.

MALE GENITALIA (Fig. 37). Basimere. Inner tergal surface with 2

irregular rows of 10 strong submarginal setae associated with several other

short setae; marginal setae numerous, strong, as in tvaubi, Subapical Lobe.

Rods a-c largely overlapping in distal portion; leaflet gj] narrow, acuminate;

leaflet 25 in form of a broad pointed blade; seta h 1, sometimes 2. Distimere,

Saba piel dorsal surface with a weak crest of few sharp spicules, not extending

to middle of curvature. Phallosome, External process with 1,2 rows of strong

denticles on tergal surface; internal process prominent, projecting beyond api-

cal lobe of external process. Proctiger. Essentially as described for traubi

and mammulifer. |

PUPA (Fig. 38). Abdomen: 2.5mm. Paddle: 0.65 mm. Trumpet: 0.5-

0.7 mm; index 12-14. Cephalothorax and abdomen dark brown or deep yellow.

Trumpet, Slender and long, dark brown, much darker than underlying integu-

ment; meatus weakly spiculate or smooth; pinna without slit extending to meatus.

Complete chaetotaxy as figured, the following are diagnostic. Cephalothorax,

Seta 1-C usually double (2-3); 5-C 3,4 branched, as long as 7-C; 8, 9-C double.

Metanotum, Setae 10,11-C usually double (1-3). Abdomen. Seta 5-IV usually

double or triple (2-4), 1.5 times as long as segment following; 5-V single, 2

times as long as segment following; 5-VI single, as long as segment following;

6-III, IV double; 6-V single or double; 6-VI usually single (1-2); 9-VII 4,5

branched; 4-VIII double; 9-VIII usually 9 branched (6-10). Paddle, Very

broad, external basal margin of outer part distinct, dark-pigmented, midrib

moderately pigmented, inner part lightly darkened in apical portion, the rest

pale whitish. 7

LARVA (Fig. 38). Head: 0.74 mm. Siphon: 2.0-2.2 mm; index 18-19.

Saddle: 0.27 mm; siphon/saddle ratio 8-9. Essentially conforms to the Mam-

milifer Subgroup, with the following diagnostic features. Head, Pigmentation

yellowish with considerable amount of brownish tinge; seta 1-C dark, slender

and simple; 5,6-C double. Antennal shaft entirely dark. Thorax, Spiculation

absent or sometimes present, very poorly developed; 7-P double; 8-P single,

varying from as long as 3-P to nearly as long as 7-P. Abdomen, Spiculation

absent; setae 6-I, II and 7-I very dark, strong and stout; 6-I, II triple; 7-I

double; 6-III-V triple; 6-VI double or triple; 1-III-V usually triple (3-4); 1-VI

4,5 branched; the remaining setae very weak and rather inconspicuous. Comb

scales narrow, numerous, about 47; posterior scales elongate, with even

fringe of fine spicules largely restricted to apex. Saddle dark brown, concolor-

ous with siphon, posterior caudal margin lightly spiculate, spicules minute and

very fine; seta 2-X double. Siphon. Very dark brownish, slender and long;

distal portion lightly tapered or uniformly thick; 10 pecten teeth, larger distal

teeth with 1,2 strong basal denticles and 3, 4 weak distal denticles; subventral

tufts relatively weak and short, 4 pairs, all of which are subequal, double or

triple, as long as or slightly longer than siphonal width at points of attachment;

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 93

median caudal filament not developed.

TYPE-DATA. (1) Culex (L.) demissus Colless, Holotype o* with slides of

antenna (CH 164) and genitalia (CT 585), Ulu Gombak, Selangor, Malaya | MA-

LAYSIA], 13 May 1957, D. H. Colless (ANIC); (2) Culex (L.) fuscosiphonis

Bram and Rattanarithikul, Holotype o* (PU4-102) with slide of antenna and

eenitalia, Muang, Phatthalung Province, THAILAND, from a tree hole ina

primary forest, 15 Oct. 1964, S. Chumchulcherm (USNM, No. 69185).

DISTRIBUTION. Known only from Thailand and Peninsular Malaysia.

21 specimens examined: 3c, 49, 14 L; 5 with associated immature skins (4 p,

1 lp).

THAILAND. Khon Kaen: Phu Wiang; Trang: National Park Forest;

Phatthalung: Muang; Nakhon Si Thammarat; Ban Thuen Lek; Ban Sai Kae; 2“,

4°, 14L, 4p, 1 lp. ;

MALAYSIA. Peninsular Malaysia - Selangor: Ulu Gombak (type-locality)

1° (holotype).

TAXONOMIC DISCUSSION. I have checked the males of fuscosiphonis

Bram and Rattanarithikul (1967: 14-6) against the holotype male of demissus

Colless (1965: 296-7) and found that they agree in every essential detail of the

antenna and genitalia. On this basis, it appears that the 2 species are con-

specific and as proposed above, I am synonymizing fuscosiphonis with demissus,

Culex demissus resembles mammilifer in the type of male phallosome and

the presence of modified tufts of setae on flagellomeres 5-9 of the male antenna

and with the latter apparently falls into a complex (the mammilifer complex).

It can be readily separated from mammialifer in all stages as indicated in the

keys and as described above. The whole larvae from Trang and Nakhon Si

Thammarat and Patthalung in southern Thailand are essentially similar, but

those from Khon Kaen in northeastern Thailand are slightly different in having

a shorter siphon and stronger siphonal tufts. It appears that the Khon Kaen

larvae probably belong to a different species. However, in the absence of

associated adult males, this record cannot be definitely confirmed.

BIONOMICS. Culex demissus is apparently rare. The specimens from

Thailand all came from the collections made in tree holes in forests. The

single male from Peninsular Malaysia was reared from a larva found ina

jungle tree hole (Colless 1965: 297).

ganapathit complex

31. CULEX (LOPHOCERAOMYIA) GANAPA THI COLLESS

(Figs. 39, 40)

Culex (Lophoceraomyia) ganapathi Colless 1965: 294 (o*, 2, L*); Bram 1967a:

00 (aeons

FEMALE. Wing: 2.9mm. Forefemur: 1.4mm. Proboscis: 1.7 mm.

Rather small, slender species; in general extremely similar to tvaubi; differ-

ing slightly in paler coloration and in the following. Head, Narrow decumbent

scales in center of vertex predominantly pale; erect scales more slender;

lateral patch of broad scales more conspicuous. Cibarial Armature, Cibarial

teeth coarser, 30-32. Thorax, Mesonotal integument paler; scales finer,

darker than underlying integument. Upper surface of ppn with some scattered

narrow dark scales cephalad of posterior bristles. Pleuron paler than mesono-

tum; lower mep bristle stronger. Legs, Anterior surface of hindfemur with

94 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

narrower pale stripe, not extending dorsad in basal 0.9.

MALE (Fig. 39). As described by Colless (1965: 294) and Bram (1967a:

90). Palpus, Equal to or slightly longer than proboscis. Pyvoboscis, Slender

and uniformly thin. Antenna, Pedicel with a rather broadly blunt spiculose

prominence on inner dorsal surface; flagellar whorls weakly to moderately

plumose; F-5 with a distinct tuft of 5,6 narrow scales, 1-3 dorsalmost scales

dark, lanceolate, as long as next 2,3 flagellomeres; followed laterally by 3, 4

yellowish brown or golden scales of decreasing length; F-6, 7 with strong

crumpled tufts of curled setae, as in tvaubi; F-8 with a slender, subapically

sinuous or gently hooked tuft of 7 fused setae; F-9 without differentiated bristle-

like setae.

MALE GENITALIA (Fig. 39). As figured. Segment LX, Tergal lobe with

2-4 tiny setae; sternum usually with 2-4 strong setae towards caudal margin.

Basimere. Inner tergal surface with a row of 4-6 moderately strong submar-

ginal setae, rather widely spaced; marginal setae weak and sparse. Subapical

Lobe, Rodlike setae a-c subequal, largely overlapping; setae d-f consist of

3,4 curved blades; leaflets g,, g) moderately strong; seta strong and apically

hooked. Distimere, Subapical dorsal surface without crest of spicules, sub-

apical claw very short and small. Phallosome, External process relatively

broad in tergal aspect, upper tergal surface with 15-17 strong denticles in 2, 3

close-set rows, apical spiculose lobe elongate, with distinct basal stalk, ap-

parently situated sternad; internal process very strong, but not projecting be-

yond apex of external process. Proctiger. Apical crown large or medium-

sized, cercal setae 3.

PUPA (Fig. 39). Abdomen: 2.4mm. Paddle: 0.55 mm. Trumpet: 0.7

mm, index 15-20. Cephalothorax and abdomen varying from cream-colored to

yellow with variable amount of brownish tinge. Trumpet, Dark, thin and very

long, apparently longest among the members of the Mammilifer Subgroup;

meatus very weakly spiculate or smooth, especially in apical 0. 5 distad of

median annulation; pinna very narrow, without slit extending into meatus.

Complete chaetotaxy as figured, the following setae are diagnostic. Cephalo-

thorax. Seta 5-C remarkably strong and long, at least 2 times as long as 7-C,

usually triple (2-4); 8,9-C usually double (2-3). Metanotum, Seta 10-C usually

double (1-3); 11-C always single. Abdomen, Seta 3-I-III usually single (1-2);

5-IV-VI very strong, 1.5-2.0 times as long as segment following; 5-IV usually

double (2-3); 5-V single or double; 5-V always single; 6-IIJ, IV weak, usually

triple (2-3), 6-V, VI stronger, usually double, sometimes triple; 9-VII, VIII

strong, subequal, 9-VII 3-7 branched; 9-VIII 7-12 branched; 4-VIII usually

double (1-3). Paddle, Entirely pale whitish; midrib lightly to moderately

darkened.

LARVA (Fig. 40). Head: 0.70 mm. Siphon: 1.3-2.0 mm; index 8-11.

Saddle: 0.3 mm; siphon/saddle ratio 5. Complete chaetotaxy as figured, dis-

tinctive in the following. Head, Cream-colored or yellowish; seta 1-C simple

or with lateral spines, lightly to strongly swollen in middle before tapering into

a sharp point, sometimes forked into 2,3 apical spines; 5,6-C double. Anten-

nal shaft pale in the middle, darkened at extreme base and beyond seta 1-A.

Thorax, Spiculation light to moderate, spicules minute, usually not very

dense; seta 7-P usually double (2-3); 8-P single, subequal to 7-P. Abdomen,

Spiculation absent; setae 6-I, II and 7-I dark, stout; 6-I, II triple; 7-I double;

1-IJI-VI usually triple (2-3); 6-III-VI triple. Comb scales 32; posterior scales

larger than anterior ones, all with rounded apical fringe of evenly fine spicules.

Saddle same color as siphon or sometimes darker; posterior caudal margin

lightly spiculate; seta 2-XK double. Siphon, Variable in length; pigmentation

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 95

usually yellowish, concolorous with head capsule, sometimes brownish; pecten

teeth 11-16, larger distal teeth with 7-10 graded denticles, basal denticles

usually not strongly differentiated, subventral tufts 4 pairs, sometimes 4. 5-5

pairs; 3 proximal pairs strong 3-5 branched each, most proximal pair longest,

4,5 times as long as siphonal width at points of attachment, next 2 pairs gradu-

ally shorter; most distal pairs shortest, 4,5 branched, about 0.5 of the length

of the next proximal pair.

TYPE-DATA. Holotype “ with associated larval and pupal skins (0643/3)

and slide of genitalia (CT 582), Ulu Langat, Selangor, Malaya | MALAYSIA],

10 May 1958 (ANIC).

DISTRIBUTION. Thailand and Malaysia. 403 specimens examined: 226,

1589, 19 L; 293 with associated immature skins (181 p, 112 lp).

THAILAND. Chiang Mai: Doi Sutep; Mae Hong Son: Ban Pa Chi; Ban Huey

Yang; Tak: Doi Sam Sao; Khao Salak Phra; Huey Lang Saeng; Nakhon Ratchasi-

ma: Khao Yai; Nakhon Nayok: Khao Yai; Khao Sing To; Kaeng Sa-An>; Kanchana-

buvi: Huey Lin Tin; Huey Mae Nam Noi; Ban Sai Yok; Khao Saeng; Chanthaburi:

Khao Hin Paeng; Khao Sai Dao; Ranong: Kraburi; Ban Chatri; Ban Bang Hin;

Khlong Baeng Man; Ban Sami; Khlong Bang Yang; Phangnga: Pak Chaung; Thap

Wen; Pathum, Nam Tai; Tang Mai; Trang: Muang; Phatthalung: Muang; Nak-

hon Si Thammarat; Ban Thuan Lek; Narathiwat; Khau Lau; 178s", 1039, 6 L,

174 p, 79 lp.

MALAYSIA. Peninsular Malaysia - Selangor: Ulu Langat (type-locality);

Ulu Gombak; The Gap; Ulu Klang; Ampang F. R. Pahang: Bentong Rd. ; Gunong

Benom; Sungai Temau; Kelantan: Bertram; Perak: Cameron Highlands; 480",

002, 9L, Tp, 32 lp. Malaysia - Sabah: Pensiangan; 4 L, 1 lp (adult ¢ lost).

TAXONOMIC DISCUSSION. Culex ganapathi is one of the most common

forms of the Mammilifer Subgroup with its distribution restricted to Malaysia

(particularly Peninsular Malaysia) and Thailand. The male antenna and geni-

talia appear to be similar to szemaonensis as described and figured by Wang

and Feng (1964: 37-41) from Yunnan, China. However, as I have not examined

the type of the latter, it is not possible to determine whether both forms are

conspecific.

The male, pupa and larva of ganapathi can be easily confused with tvaubi

and the members of the minoy complex. The male strongly resembles traubi

in the development of the modified tufts of antennal flagellomeres 5-8, but is

different from the latter in having darker, broader and longer acute scales

in the modified tuft of flagellomere 5 and in the shape of the modified tuft of

flagellomere 8. The male genitalia resemble members of the mznor complex

from which it is differentiated by the presence of 4-6 moderately strong sub-

marginal setae on the basimere and the largely overlapped rodlike setae of

the subapical lobe. The pupa can be readily recognized by the very slender

and long trumpet; the remarkably strong seta 5-C and the usually single setae

3-I-III. The larva resembles tvaubi in most features of the chaetotaxy but can

be distinguished from it by the paler antenna, siphon and saddie, double seta

o-C, triple seta 6-VI and by the poorly differentiated denticles of the pecten

teeth.

All larvae of ganapathi essentially conform to the above description except

for the specimens from Sabah, which differ from the usual form in having denser

spicules on the thorax, a darker siphon, presence of 5 pairs of the siphonal

tufts and in the development of strong basal denticles of the pecten teeth. Un-

fortunately, the male associated with this larval form is not available for this

study. This form may prove to be distinct, but pending further knowledge of

the male, they were tentatively identified as ganapathi,

96 Contrib.: Amer. Ent. Inst., vol. 13, no. 4, 1977

BIONOMICS. The immatures of ganapathihave been collected from bam-

boos and tree holes. They were found more frequently in bamboo than in tree

holes. In Thailand, it has also been reported from pandanus axils and ground

pools and in Peninsular Malaysia from artificial containers. The records from

ground pools are probably in error, perhaps due to contamination. In Pen-

insular Malaysia, ganapathi has been frequently found in association with

minor (Colless 1965: 295). All adults apparently came from individual or

mass rearing the larvae.

32. CULEX (LOPHOCERA OMYIA) SPICULOSUS

BRAM AND RATTANARITHIKUL

(Figs. 41, 42)

Culex (Lophoceraomyia) spiculosus Bram and Rattanarithikul 1967: 3 (o*, &,

bo): Bran be6ia- 105 (67, 2, GF).

Culex (Lophoceraomyia) hui Lien 1968: 227 (*, 2, L*). NEW SYNONYMY.

FEMALE. Wing: 3.2 mm. Forefemur: 1.6 mm. Proboscis: 1.9 mm.

Medium-sized species with external characters as described for most members

of the Mammilifer Subgroup, differing from the preceding species in relatively

larger size and slightly in the following. Head, Decumbent scales of vertex

entirely narrow, linear and predominantly yellowish brown; lateral patch of

broad scales dingy white; erect scales rather coarser, usually entirely dark,

sometime partially pale yellowish on posterolateral area. Cibarial Armature.

As figured for mammilifer, cibarial teeth about 35, 8-10 median teeth narrow

and apically pointed, lateral teeth coarser, with truncate or blunt apices.

Thorax, Mesonotal integument and scales blackish. Lower mep bristle pre-

sent. Legs, Anterior surface of hindfemur with a broad whitish stripe, as

in traubi, Wing. Scales on wing veins rather dense. Abdomen, Terga black

scaled, sterna pale whitish or light yellowish.

MALE (Fig. 41). Similar to tvaubi and ganapathi in having modified anten-

nal tufts developed only on flagellomeres 5-8, differing chiefly in the following

features. Palpus. Longer, exceeding proboscis length by 0.5 to full length of

segment 5. Antenna, Flagellum densely long plumose; F-9 with a minute tuft

of 4-7 dark, short, hairlike setae, all subequal, as long as 1 flagellomere in

length; F-6,7 with slender or weakly developed crumpled tufts of curled setae;

F-8 with a prominent tuft of 5,6 fused setae, which is straight or weakly

curved in distal portion; F-9 usually without any modified bristlelike seta,

sometimes with 1 weak seta among several long normal setae.

MALE GENITALIA (Fig. 41). As figured; exceedingly similar to traubi

and ganapathi, differing in the following details. Basimere, Inner tergal sur-

face with a prominent row of 6-8 moderately strong submarginal setae and 2,3

rows of several weak marginal setae, submarginals closely spaced, in a linear

or irregular row. Subapical Lobe. Distal part with several weak, short, hair-

like setae associated with setae d-f; seta h remarkably strong and long; rod c

clearly separated from @, 0 distad; leaflets gy and g» as in traubi and ganapathi,

Phallosome, As in ganapathi; internal process distinct, shorter than external

process; external process with a distinct spiculate apical lobe sternad and

several strong denticles on tergal surface. Proctiger. Crown large; cercal

setae usually 4.

PUPA. Abdomen: 2.7mm. Paddle: 0.72 mm. Trumpet: 0.60 mm;

index 9-10. As figured for traubi (Fig. 30). Pigmentation of cephalothorax

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region of

and abdomen brownish or yellowish with considerable amount of brownish

tinge. Trumpet, Dark brown, relatively thick and long; meatus strongly

spiculate distad of median annulation, as in fvaubi; pinna broad and moderately

oblique. Diagnostic chaetotaxy as indicated in the key and as in the following.

Cephalothovax, Seta 1-C usually double (2-3) and weak; 5-C usually triple

(2-5), as long as 7-C; 8,9-C double. Metanotum, Seta 10-C usually double

(1-3); 11-C single. Abdomen, Setae 3-I-III single or double; segment II usu-

ally with 1 extra seta (seta 11 of Belkin 1962) on ventral surface; 5-IV 2-4

branched, as long as segment following; 5-V, VI single, 5-V 1.5 times as long

as segment following; 5-VI 0.5 of segment following; 6-III- VI weak, subequal,

usually double (2-3); 9-VII 3-5 branched, shorter than 9-VIII; 9-VIII 5-8 bran-

ched and strongly pectinate. Paddle, Rather broad, yellowish white, midrib

strong and moderately pigmented.

LARVA (Fig. 42). Head: 0.78 mm. Siphon: 1.4-1.9 mm; index 7-8.

Saddle: 0.34 mm; siphon/saddle ratio 4-5. Strikingly differentiated from all

members of the Mammilifer Subgroup except uniformis and kuhnsi by the

heavily spiculose thorax and abdomen; complete chaetotaxy as figured; differ-

ing from the latter 2 species as indicated in the key and in the following addi-

tional features. Head, No transverse band of spicules at level of ocular bulb

on ventral surface; seta 1-C slender, simple and long; 5,6-C double. Antenna

usually entirely dark, sometimes partially pale in middle, spicules rather fine.

Thorax, Spiculation usually strong and dense, sometimes reduced; individual

spicules hairlike and unbranched; seta 8-P strong, subequal to 7-P, single;

7-P usually double (2-3). Abdomen. Spiculation as in thorax; setae 6-III usu-

ally triple (8-5); 6-IV-VI usually double or triple (2-4); 1-III-VI strong, about

0.5 of 6-III-VI, double or triple (2-4). Comb scales about 40. Saddle dark

brown; posterior caudal margin lightly to moderately spiculate, individual spi-

cule short and weak; seta 2-X single. Siphon, Yellowish to dark brownish,

length variable; pecten teeth 10-15, large distal teeth with 1 strong basal

denticle and 6, 7 fine distal denticles; subventral tufts moderately long, 4 pairs,

first 3 proximal pairs usually 4 branched each (2-4), 2,3 times as long as

siphonal width at point of attachment, most distal pair 4,5 branched, 1.5

times as long as siphonal width at point of attachment.

TYPE-DATA. (1) Culex (L.) spiculosus Bram and Rattanarithikul, Holo-

type o* (00272-3) with associated larval and pupal skins and slide of antenna

and genitalia, Doi Sam Sao, Tak, THAILAND, bamboo stump, 30 July 1965,

Sumeth Chumchulcherm (USNM; No. 69181); (2) Culex (L.) hui Lien, Holotype

So (61864,4) with associated larval and pupal skins, Chukou, Fanlu, Chia Hsien,

TAIWAN, bamboo stump, 14 May 1961, J. C. Lien (Taiwan Provincial Institute

of Infectious Diseases, Taipeh, Taiwan, Republic of China).

DISTRIBUTION. Thailand, Burma, Peninsular Malaysia and Taiwan. 187

specimens examined: 64, 32°, 91 L; 67 with associated immature skins (19

p, 48 lp).

THAILAND. Chiang Mai: Doi Sutep; Ban Khun Klang; Tak: Doi Sam Sao

(type locality); Khao Salak Phra; Nakhon Nayok: Khao Yai; Chanthaburi: Khao

Sai Dao;'580,. 299, 91 L, 29 p, 36. lp.

BURMA. Rangoon; 1c.

MALAYSIA. Peninsular Malaya - Perak: Cameron Highlands Rd. ; 4c,

62, 10 lp.

TAIWAN. Chia Hsien: Chukuo Fanlu; 1, 12, 2 lp (as hui, Lien 1968).

TAXONOMIC DISCUSSION. The proposed synonymy of hui Lien (1968: 227)

from Taiwan with spiculosus Bram and Rattanarithikul (1967: 3) from Thailand

has been based on the comparison of the specimens in the type-series. In

98 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

checking 2 paratypes of hui against the type and several other topotypic speci-

mens of spiculosus, no difference has been found except for the spiculation of

the larva which is better developed in spiculosus than in hui,

Culex spiculosus is morphologically intermediate between ganapathi and

traubit. The male phallosome of spiculosus resembles that of ganapathi but

the larva and pupa are similar to tvaubi in most features of the chaetotaxy.

The larva is most diagnostic, particularly in the development of dense spicula-

tion on the thorax and abdomen which is superficially very similar to those of

uniformts from Sri Lanka and kuhnsi from the Philippines. It is quite probable

that the record of uniformis from Hainan Island, China by Chu (1958: 109-13)

is in fact that of spiculosus,

BIONOMICS. Culex spiculosus is restricted to high elevations in inland

mountainous areas. The immatures have been frequently collected from bam-

boo stumps and occasionally from tree holes under heavy shade of tropical for-

ests. Most of the adults came from rearing the larvae and pupae. In Thailand,

several were also caught while resting on vegetation near the breeding sites.

minor complex

33. CULEX (LOPHOCERAOMYIA) MINOR (LEICESTER)

(Figs. 43, 44)

Lophoceratomyia minor Leicester 1908: 126 (o, 9).

Culex (Lophoceratomyia) minor (Leicester), in part, Edwards 1917: 227

(", key); Edwards 1922a: 282 (key); Barraud 1924: 44 (o*, 2); Borel

1930: 356 (“*, &, L*); Edwards 1928: 276 (key); Edwards and Given

1928: 352 (L); Edwards 1932: 198 (taxonomy); Barraud 1934: 370 (o*, 9);

Brug and Bonne-Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) minor (Leicester), Mattingly 1949: 227 (L, key);

Colless 1965: 289 (“*, @, L*); Delfinado 1966: 111 (c"*, 2); Bram 1967a:

OG Oe ES Na pare..

Culex (Lophoceratomyia) plantaginis Barraud 1924: 46 (o'*); Barraud 1934:

372 (o*, in part). NEW SYNONYMY.

Culex (Lophoceratomyia) nolledoi Baisas 1935: 170 (o“*); Colless 1965: 289

(Synonymy).

FEMALE. Wing: 2.9-3.8 mm (average 3.4 mm). Forefemur 1.4-1.9

mm (average 1.8 mm). Proboscis: 1.7-2.2 mm (average 2.1mm). Usually

medium size (wing length over 3.0 mm), sometimes small or rather large; in

general very similar to most members of the Mammilifer Subgroup from

which it cannot be distinguished with certainty. Head, Decumbent scales on

dorsum of vertex very narrow, linear and pale along upper eye margin, pre-

dominantly dark or yellowish brown in center and on occiput; erect scales

entirely blackish and rather coarse; lateral patch of broad scales whitish or

grayish, fairly distinct. Palpus black scaled, 0.2 of proboscis length, seg-

ment 4 longest, about 3 times as long as the combined length of segment 1-3.

Proboscis black and relatively thick; longest labial basal seta about 0.5 of pal-

pal length. Cibarial Armature. Cibarial bar with a concave row of 35, 36

teeth; 8 median teeth narrow, fine with sharply pointed apices, lateral teeth

coarser with blunt or truncate apices. Thorax, Integument of mesonotum

deep chestnut brown or black; scales rather narrow, short, fine and entirely

black; dorsocentral, supra-alar and scutellar bristles very strong. Upper

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 99

surface of ppn without any setae or scales cephalad of posterior bristles.

Pleuron paler or same color as mesonotum; scales practically absent; pp/ with

1,2 strong dark bristles and a few other weak setae; 1 lower mep bristle pre-

sent. Legs, Anterior surface of hindfemur with moderately broad pale

stripe from base to apex, rest of legs blackish, without any marked coloration.

Wing. Scales on all wing veins very narrow, moderately to very dense. Abdo-

men, All terga black scaled; sterna usually pale yellowish or whitish, some-

times as dark as terga.

MALE (Fig. 43). As described by Colless (1965: 289-91). Palpus,

Slender, varying from as long as or longer than proboscis by 0.5-1.0 of seg-

ment 5: segments 4 and 5 and apex of segment 3 with only a few short bristles.

Proboscis, Uniformly thick throughout entire length, setae widely scattered

and sparse, largely restricted to lateral surface. Antenna, Spiculose promi-

nence on inner dorsal surface of pedicel strong, distinct, with apex more or

less pointed; flagellar whorls moderately to densely long plumose; modified

tuft of setae developed on flagellomeres 5-9; F-5 with a small, rather incon-

spicuous tuft of 3-5 yellow or golden spinelike setae which are subequal, as

long as 1 flagellomere, preceded dorsally by 2-4 long normal setae; F-6 with

a strong crumpled tuft of heavily curled and subapically twisted setae; F-7 with

a mesal tuft of 5-7 narrow, short, acute spinelike setae in addition to 2 other

usual tufts on lateral and ventral surfaces; F-8 with a thick, sigmoid or J-

hooked tuft of 6,7 fused setae and a close-set row of 6,7 short bladelike setae

laterad; F-9 with a tuft of 3, 4 dark bristlelike setae.

MALE GENITALIA (Fig. 43). Segment Lx, Tergal lobe bears 3-5 mod-

erately strong setae; sternum with 1,2 irregular rows of 4-10 strong setae

towards caudal margin. Basimere. Well sclerotized; inner tergal surface

with 6,7 strong sinuous submarginal setae in a close-set row which is slightly

diverging towards lateral tergal surface; 6-9 marginal setae weak, widely

spaced. Subapical Lobe, Most proximal rod or seta a strongly curved in mid-

dle, separated from setae b, c by a wide gap; setae 0, c gently curved or

straight, with bases overlapped with those of setae d-/f in distal part; leaflet

g 1 broad, apex acuminate; & long, bladelike; seta i strong. Distimere,

Normal; subapical dorsal surface without crest of spicules; subapical claw

short and small; dorsal and ventral tiny seta present, dorsal seta distad of

ventral. Phallosome, Well sclerotized; internal process strong, simple,

curved tergad and not projecting beyond apex of external process; external

process with 2,3 rows of 15-20 strong denticles on upper tergal surface and a

spiculose apical lobe sternad, inner surface with variable number of toothlike

marks. Proctiger. Crown dark, large, consisting of numerous strong spine-

like spicules; paraproct well sclerotized and moderately pigmented; cercal

sclerite broad, poorly or moderately sclerotized and pale; cercal setae 3-5,

distinct.

PUPA (Fig. 43). Abdomen: 3.2 mm. Paddle: 0.70 mm. Trumpet: 0.80

mm; index 12. Cephalothorax and abdomen varying from pale, cream-colored,

yellowish to dark brownish. Trumpet. Dark brown, rather thick and long;

apical portion distad of proximal annulation of meatus weakly to moderately

spiculated; pinna moderately broad oblique, without slit extending to meatus.

Complete chaetotaxy as figured, diagnostic features as in the key. Cephalo-

thorax, Seta 1-C usually double (2-4); 5-C subequal to 7-C, usually double

(2-4); 7-9-C double. Metanotum,. Setae 10-C usually double (1-3); 11-C usu-

ally double (1-2). Abdomen, Setae 3-I-III usually double (1-2); 5-IV, V 1. 5-

2.0 times as long as segment following; 5-IV usually double (2-4); 5-V usually

double (1-2); 5-VI slightly shorter than segment following, usually single (1-2);

100 Contrib. Amer. Ent. Inst., vol. 18, no. 4; 1977

6-III-VI usually double (1-3); 9-VII subequal to 9-VIII, usually triple (2-4);

9-VIII 6-9 branched. Paddle, Very broad and pale, basal external margin

of outer part distinct, apical margin indistinct, midrib lightly pigmented;

setae 1,2-P present; 1-P strong, dark; 2-P weak, minute, rather inconspicu-

ous.

LARVA (Fig. 44). Head: 0.78 mm. Siphon: 1.7 mm; index 8,9. Saddle:

0.30 mm.; siphon/saddle ratio 5. Complete chaetotaxy as figured, diagnostic

characters as in the key; setae very variable in color, length and thickness.

Head, Yellowish to light brownish; seta 1-C simple, slender and moderately

long; 5-C usually double (1-3); 6-C double. Antenna variable in color, usually

pale in the middle, dark at base and beyond insertion of seta 1-A, or some-

times uniformly dark; spicules strong and dark. Thorax, Unspiculate or

lightly spiculate; spicules minute, dotlike, widely spaced and rather indis-

tinct; setae 1-3-P single; 4-P usually double (2-3); 5,6-P single; 7-P double,

8-P single, subequal to 7-P; 14-P double. Abdomen, Spiculation absent ex-

cept on segment VIII; setae 6-I, II and 7-I usually very dark and stout, all

double; 6-III-VI double or triple; 1-III-VI and 13-III-V usually triple (2-4),

varying from 0.25-0.75 of length of 6-III-VI. Comb scales 35-50; anterior

scales with normal fringe of evenly fine spicules; posterior scales elongate,

with apical fringe terminated into a stout median spine. Saddle yellowish to

brownish, posterior caudal margin lightly spiculate; seta 2-X triple; anal gills

1.5 times as long as saddle length. Siphon. Very slender, more or less

tapering distally; pigmentation variable, usually yellow, concolorous with head

capsule, sometimes light brown; pecten teeth 11-16, larger distal teeth barbed

with 7,8 denticles, 1,2 basal denticles usually enlarged and strongly differenti-

ated from distal denticles; subventral tufts weak and rather short, 3 pairs,

usually 3,4 branched (3-5) each; first proximal pair strongest, 1.0-1.5 times

as long as siphon width at point of attachment, the remaining distal pairs gradu-

ally shorter; median caudal filament of spiracular apparatus present, well

developed.

TYPE-DATA. (1) Lophoceratomyia minor Leicester, lectotype “* with

attached antenna and genitalia mount, Ulu Klang, Kuala Lumpur, Selangor,

Malaya | MALAYSIA], 14 October 1903, G. F. Leicester (BMNH; selection of

Colless 1965: 289); (2) Culex (Lophoceratomyia) plantaginis Barraud, Lecto-

type ¢* with slide of antenna and genitalia, marked as type by Barraud, Koti,

near Kalka (Kalka-Simla road), Himalaya foothills, INDIA, August 1923, P. J.

Barraud (BMNH; present selection); (3) Culex (Lophoceratomyia) nolledoi

Baisas, Type o, 2 (non-existent), Kolambugan, Lanao Province, Mindanao,

PHILIPPINES, 31 July 1934, Francisco Guinto (Bureau of Health, Manila;

apparently lost or destroyed).

DISTRIBUTION. India, Thailand, Malaysia, Indonesia and the Philippines.

676 specimens examined: 332%, 2719, 72 L; 359 with associated immature

skins (147 p, 212 lp).

INDIA. Western Himalayas: Koti near Kalka; 1° (as plantaginis Barraud);

Andaman Is.; 2%.

THAILAND. Nakhon Ratchasima; Koraj, Pachong; Khao I-Phrom; Nakhon

Nayok: Huey Tha Thong; Chon Buri: Svriracha; Chanthaburi: Ban Chak Yai;

Khao Hin Phaeng; Khao Sai Dao; Khlong Ka Ting; Tvat; Koh Chang; Ban Cham

Rung; Phattalung: Muang; Ranong: Kraburi: Khlong Bang Yang; Nakhon Si

Thammarat; Ban Thuen Lek; Tvang: Gun Tung; Songkhia; Tone Nga Chang;

1030, 89%) 1o'L, 101 p, 69 Ip.

MALAYSIA. Peninsular Malaysia-Selangor: Ulu Gombak; Ulu Langat;

Bt. Ulu Bakau; Bt. Kutu; Pahang; Pulau Tioman; Bentong Rd., M. S.; Kg.

Sirivanakarn: Subgenus Lophoceratomyia in the Oriental Region 101

Janda Baik; Gunong Benom; Sungai Temau; Fraser's Hill; Perak; Kg. Sq. Itek;

Perlis: Kg. Wang Kelian; Kedah; Kg. Bagan; Malaysia-Savawak; Tebangan;

Sabah; Mt. Kinabalu; Beaufort; Tenom; 110, 1122, 31 L, 38 p, 75 lp.

INDONESIA. Kalimantan: Tarakan; Java: Dijakarta; West Irian: Humboldt

Bay; 62,52. |

PHILIPPINES. Luzon: Tayabas; Trinidad Mt. Province; Dalton, N.

Vizcaya; Subic Naval Base; Mindoro: San Jose; Palawan; Mt. Malinao; Irah-

nan River; Iwahig; Puerto Princesca; Leyte: Lagolago Baybay; Camarines Sur,

Mt. Isarog; Samar: Osmena; Calaccad; Sohoton; Jolo Jolo Is, ; 110, 659,

2 1s, 8 p, 08 1p.

Additional records from the literature: BURMA (Barraud 1934).

TAXONOMIC DISCUSSION. Culex minor has been recorded from Malaysia,

southern Thailand, Indonesia and the Philippines where it is one of the most

common forms of the Mammilifer subgroup. Elsewhere in Southeast Asia and

other parts of the Oriental region, it appears to be completely replaced by

bicornutus to which it is most closely related. The proposed synonymy of

plantaginis Barraud from India with minor is only tentative but appears to be

correct since the type-male which I examined and designated as a lectotype per-

fectly agrees with minor in the characters of antenna, proboscis and genitalia.

As described by Barraud (1924: 46; 1934: 372) and subsequently discussed by

Colless (1965: 291), plantaginis differs from minor in having basal pale bands

on the abdominal terga. However, as the type was mounted in balsam and

other specimens in the type-series were found to be a mixture of different

species, including bicornutus, it has not been possible to evaluate the taxono-

mic importance of this character. Further study of additional topotypic ma-

terial is required in establishing the identity of plantaginis, but for the pre-

sent, it appears justified to consider it as a synonym of minor. There is also

a possibility that it may be conspecific with bicornutus, but without additional

material, this problem cannot be resolved. I have examined the larvae from

Nepal, tentatively identified as plantaginis by Peters and Dewar (1956: 44)

and found them to be Culex (Culex) whitei.

The adult males of minor which I have examined from several localities

listed in the distribution data agree well with the type and topotypic specimens

from Malaya. The larvae and pupae are subject to considerable variation in

size, pigmentation and chaetotaxy. Those from rock pools are usually darker

and exhibit stronger setae than the ones from tree holes or bamboos but all

essentially conform to the diagnostic chaetotaxy as described and illustrated.

The larvae from rock pools usually differ from those in container habitats in

having setae 1 and 6-III-VI stronger and more branched. The development of

1,2 stout basal denticles in the larval pecten teeth as described by Colless

(1965: 290) is especially very diagnostic of the minor populations in Malaya

and Sabah but has been found to be subject to a considerable variation, over-

lapping with bicornutus in the populations from Thailand and the Philippines.

In all the material from the Philippines which I have examined, the males are

typical of minor but the larvae show evenly graded denticles of the pecten

teeth as in bicornutus,

Among the members of its complex, minor most closely resembles bicornu-

tus and bandoengensis in all stages. It can be separated from bicornutus with

certainty only in the male by the uniformly thick proboscis and the absence of

prominent lateral and ventral rows of several strong setae in the basal 0. 2-

0.9 of the labium; from bandoengensis in the male genitalia by the presence of

6,7 submarginal setae in a single row on the inner tergal surface of the basi-

mere (15-17 in double row in bandoengensis).

102 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

BIONOMICS. Culex minor has been encountered in practically all collec-

tions from the inland jungle in various parts of Southeast Asia. The adults

and immatures are very abundant and have frequently been collected in num-

bers. The principal breeding sites include bamboos and tree holes. In the

Philippines, it has also been found breeding in rock pools, but elsewhere, in

Thailand and Malaysia, it was reported to utilize bamboos and tree holes in

preference to rock pools (Bram 1967a: Colless 1965). In Malaya, Colless

(loc, cit.) noted that the adults appeared to attack man on occasion, otherwise

nothing is known about its biology and medical importance.

34. CULEX (LOPHOCERA OMYIA) BICORNUTUS (THEOBALD)

(Fig. 44)

Lophoceratomyia bicornuta Theobald 1910: 25 (o'*).

Culex (Lophoceratomyia) mammilifer of Barraud 1924: 43; Edwards 1932: 198;

Barraud 1934: 374; Stone, Knight and Starcke 1959: 234.

Culex (Lophoceratomyia) plantaginis Barraud (in part), Barraud 1924: 46 (o*);

Barraud 1934: 372 (c‘*). :

Culex (Lophoceraomyia) minor (Leicester), (in part), Stone, Knight and

Starcke 1959: 234 (catalog); Bram 1967a: 97 (o*, 9, L*).

Culex (Lophoceraomyia) bicornutus (Theobald), Colless 1965: 291 (o*, °, L*).

FEMALE. As described for minor from which it is virtually indistinguish-

able.

MALE. Exceedingly similar to minor, differing from it in the following

characters. Palpus, Rather longer, usually exceeding proboscis length by

0.5-1.0 of segment 5. Pyvroboscis, Relatively thick and moderately to strongly

swollen from 0.2-0.5 of the length from base, lateral and ventral surfaces of

swollen portion with prominent rows of numerous strong, curved setae. <An-

tenna, As figured for minor (Fig. 43); F-7 with a stronger and more promi-

nent mesal tuft of dark, flattened spinelike setae which are well separated

from 2 other usual tufts on ventral and lateral surfaces.

MALE GENITALIA. Essentially as described and figured for minor (Fig.

43) to which it is extremely similar in practically every detail of the basimere,

subapical lobe, distimere, phallosome and proctiger.

PUPA. As described and figured for minor (Fig. 43) to which it is essen-

tially similar in the detailed chaetotaxy. Cephalothorax and abdomen varying

from extensively pale yellowish in specimens from pandanus axils to brownish

in the specimens from rock pools or tree holes.

LARVA (Fig. 44). As described and figured for minor to which it is

essentially similar in the detailed chaetotaxy, differing slightly in the follow-

ing. Thorax and Abdomen, Spiculation very variable; those from rock pools

and tree holes lightly spiculate, spicules minute and sparse as in mnor; those

from pandanus axils heavily spiculate; spicules dense, hairlike, nearly ap-

proaching the conditions noted in spiculosus; setae 1 and 6-III-VI relatively

stronger and longer; 6-III-VI usually triple, rarely double. Szphon, Larger

distal pecten teeth with 6, 7 denticles of graded sizes (Fig. 44), 1,2 basal

denticles apparently not differentiated from distal denticles; subventral tufts

as in minor, sometimes stronger and longer.

TYPE-DATA. Type & (lost), Dawna Hills, nr. Kawkareik, BURMA (LU).

DISTRIBUTION. India, Sri Lanka, Thailand, Vietnam, Peninsular Malay-

sia, Ryukyus (Japan) and Hainan (China). 1,979 specimens examined: 743¢,

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 103

7612, 475 L; 1,153 with associated immature skins (512 p, 641 Ip).

INDIA. Bombay: Kawar; 40°, 29.

SRI LANKA. Central Province: Imbulpitiya; Sabavagamuwa Province:

Galle Dist., Kanneliya; Western Province: Colombo Dist. ,Labugama Reservoir;

36¢,.. 502... 8.1. B.4p,

THAILAND. Mae Hong Son: Ban Pha Chi; Ban Mae Tia; Doi Chang; Ban

Mae Ho Nua; Ban Hua Yang; Lampang: Ban Pha Daeng; Chiang Mai: Doi Sutep;

Huey Kaew: Chiang Dao; Huey Pao; Huey Payao; Ban Sap O Nok; Wat Phra

That; Ban Doi Sutep; Ban Than Kleap; Ban Mae Sa; Doi Khun Tan; Ban Pha

Miang; Tak: Doi Sam Sao; Ubon Ratchathani: Pibul Muang Saharn; Nakhon

Ratchasima; Koraj, Pachong; Kabin Buri; Khao Salak Dai; Khao I-Phram;

Nakhon Sawan; Nakhon Nayok: Huey Tha Krong; Khao Yai; Huey Sai Noi; Sava-

buri; Chon Buri: Bang La Mung; Khao Mai Kaeo; Sriracha, Bang Phra; Kan-

chanaburi: Ban Sai Yok; Chanthaburi: Ban Chak Yai; Ban Laem Sing; Khlong

Ka Ting; Ban Cham Rung, Ban Sapan Hin; Tvat; Koh Chang; Prachuap Khiri

Khan; Tub Sakae; Chumphon: Ban Wang Phi; Ranong: Kraburi; Khlong Set

Takuart; Kvabi: Ban Mai Kaen Tai; Phangnga: Pathum;Nam Tai: Sipsi Hon;

Surat Thani: Koh Samui; Khao Yai; Khao Plu; Ron Pibul; Phuket: Ban Huey

Luk; Khao Prathui; Khao Prathin; Nakhon Si Thammarat; Phu Wiang; Ban

Thuan Lek; Khao Luang; Phatthalung: Muang; Songkhla: Boripat Water Falls;

Tone Nga Chang; Trang: Muang; Narathiwat: Khau Lau; Yala; Yala Bong;

0260°, 5409, 411 L, 496 p, 437 lp.

VIETNAM. Con Son; 6c, 19, 5 L.

MALAYSIA. Peninsular Malaysia-Selangor: Ulu Gombak; K. Kubu-Gap

Rd. ; Pacific Tin K. Sel.; Banting; Selangor-Pahang: The Gap; Pahang: Pulau

Tioman; Gunong Benom; Fraser's Hill; Perak: Kg. Sg. Itek; Cameron High-

lands Rd.; Bintang Hijau F. R.; Maxwell's Hill; Penang Is.; 117%, 1142, 46 L,

16 p, 94 lp. :

CHINA. Hainan Is.; 10.

JAPAN. Ryukyus (Okinawa): Yaeyama; Iriamote; 44°, 542, 5 L, 101 lp.

Additional records from the literature: Botel Tobago near Formosa

(Taiwan); lower BURMA; (Colless 1965). _

TAXONOMIC DISCUSSION. Culex bicornutus is exceedingly similar to

minor in all stages. It can only be separated from the latter by the male

proboscis which differs strikingly in being swollen in the basal 0. 2-0. 5 and in

the presence of prominent rows of several strong curved setae on its lateral

and ventral surfaces. These features are remarkably constant in all material

examined except in central and northern parts of Thailand where they are vari-

able and occasionally exhibit overlap with the normal condition noted in the typi-

cal minor, It is quite possible that some of the Thailand specimens which I

have identified with minor are actually bicornutus.

In spite of their close resemblance, bicornutus is undoubtedly distinct from

minor on the basis of their distribution and breeding site. Over most parts of

Southeast Asia and adjacent areas, bicornutus has been found to be more com-

mon and widespread than minor, Its occurrence is apparently restricted to the

north, extending from India, Sri Lanka eastwards to southern China, northern

and central Thailand, Okinawa and perhaps also Taiwan. I have never seen

bicornutus specimens from Sabah, Indonesia or the Philippines where minor

is predominant. In Sri Lanka, Hainan and Okinawa, the Ryukyus, however, it

appears to completely replace minor. In Thailand and Malaysia, both forms

are generally or closely overlapping in their occurrence and breeding sites.

The differences in the type of the breeding sites between the 2 forms also

appear to be rather striking, as noted by Colless (1965); bicornutus exhibits a

104 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

tendency to breed in rock pools instead of bamboos or tree holes as opposed to

minor which usually breeds in container habitats in preference to rock pools

or other types of ground pools. In addition, bicornutus has also been found

capable of utilizing a broader spectrum of breeding sites than minor, Included

among them are pandanus axils, ground pools and artificial containers. On

this basis, Iam in full agreement with Colless (1965) in treating bicornutus

as a distinct species instead of relegating it to a form or synonym of minor as

proposed by Bram (1967a).

In Thailand and Peninsular Malaysia, the larvae of bicornutus exhibit con-

siderable variation in the spiculation of the thorax and abdomen, depending on

the type of breeding sites; in those from pandanus axils these features are

better developed than the ones from rock pools and other breeding habitats.

Certain abdominal setae are also variable in the length and in the number of

branches as noted in minor, indicating that bicornutus is probably a composite

of several different ecotypes.

BIONOMICS. The majority of the immatures of bicornulus from southern

Thailand and Peninsular Malaysia were collected in rock pools at stream mar-

gins under heavy or partial shade of dense tropical forest and usually contained

numerous decayed fallen leaves. Occasionally, they have also been found in

tree holes, log holes and bamboo stumps close to the ground, stream pools,

wells, puddles, footprints, clay or bamboo pots, coconut shells and pandanus

axils. In northern Thailand, it has been reported to utilize rock pools as fre-

quently as tree holes or bamboos. In the Ryukyus, most of the larval collec-

tions were from rock pools and only a few from tree holes. The immatures

recently collected in Sri Lanka largely came from rock pools. Both pupae and

larvae were abundant and have frequently been taken in numbers. The numer-

ous adults have been obtained largely from rearing the pupae and larvae.

Nothing is known about the adult biology.

oo. CULEX (LOPHOCERAOM VIA) BANDOENGENSIS BRUG

(Fig. 45)

Culex (Lophoceratomyia) minor var. bandoengensis Brug 1939: 112 (*).

Culex (Lophoceraomyia) minor (Leicester) of Stone, Knight and Starcke 1959:

234 (catalog). |

Culex (Lophoceraomyia) bandoengensis Brug, Colless 1965: 292 (o*, &, i).

FEMALE. Exceedingly similar to minor and bicornutus from which it can

not be distinguished with certainty, differing slightly in having the narrow de-

cumbent scales of the vertex finer, the erect scales more slender and less

numerous. :

MALE. In general differing from minor and bicornutus as described for

female. Palpus, Slender and relatively long, exceeding proboscis by full

length of segment 5. Proboscis, Uniformly thick, as in minor. Antenna,

Modified tufts of F-5-9 as described for minor and bicornutus, differing slight-

ly in having the mesal tuft of F-7 more weakly developed, composed of 4, 5

narrow, pale spinelike setae.

MALE GENITALIA (Fig. 45). As figured and described by Colless (1965:

292), differing from minor and bicornutus as in the following. Basimere,

Inner tergal surface with a double or triple row of 15-17 very strong sinuous

submarginal setae. Subapical Lobe, Rod a always separated from rods De

by a wide gap, but more strongly curved and sometimes distinctly angulate or

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 105

elbowed at middle; leaflet g; very broad, usually longer than bladelike leaf-

let g9; setae d-f relatively stronger and more distinct. Distimere, Rather

short, usually more or less straight with slightly recurved apex; subapical

claw short and strongly inflated distally. Phallosome and Proctiger. Es-

sentially as described for minor and bicornutus,

PUPA. As figured for minor to which it is exceedingly similar in general

and detailed chaetotaxy, differing particularly in having seta 8-C usually 3, 4

branched (2-4).

LARVA. As figured for ™inor to which it is exceedingly similar, differing

particularly in the following. Abdomen, Setae 6-III-VI usually triple (2-3).

All comb scales with normal fringe of evenly fine spicules. Szphon, Distal

teeth of pecten always barbed with 6,7 denticles of graded sizes.

TYPE-DATA. Holotype “* with attached antenna and genitalia mount,

Bandoeng (Bandung), Java, Dutch East Indies | INDONESIA], reared from

larva collected in bamboo stump, elevation 800 meters, 2 Jan. 1934, S. L.

Brug (BMNH).

DISTRIBUTION. Peninsular Malaysia and Indonesia. 39 specimens ex-

amined: 180%, 1392, 7 L; 13 with associated immature skins (2 p, 11 lp).

INDONESIA. Java: Bandung (type-locality); Bogor, Botanical Garden;

Ceram; 10, 39.

Additional records from the literature: PENINSULAR MALAYSIA-Selangor:

Fraser's Hill; Perak: Maxwell Hill (Colless 1965: 293).

TAXONOMIC DISCUSSION. Culex bandoengensis is very closely related to

minor and bicornutus, It can be separated from the latter 2 species particular-

ly by the presence of more numerous submarginal setae in double or triple rows

on the basimere of the male genitalia and by the even fringe of the larval comb

scales. 3

Based on the present records, bandoengensis appears to be restricted to

Peninsular Malaysia and Indonesia. The new record from Ceram is based ona

male collected by Brug and de Rook in 1931. This, together with the recent

collection of the specimens from Java and Peninsular Malaysia, indicates that

it is widespread, probably occurring throughout these areas.

BIONOMICS. As in minor, bandoengensis has been frequently found to

utilize bamboos and tree holes as the breeding sites. All of the adults from

Java were reared from larvae collected in bamboos or tree holes. In Malaya,

the immatures were also collected in rock pools in association with numerous

specimens of bicornutus. On occasion, they were also collected from discarded

oil drums and tin cans. All of these collections were made in or near the forest

at a high elevation in the mountainous region.

36. CULEX (LOPHOCERAOMYIA) TUBERIS BOHART

(Fig. 45)

Culex (Lophoceraomyia) tuberis Bohart 1946: 42 (o'*); Bohart and Ingram 1946:

14(os, Go op It): Bohart 1953: lea (ots Pie

FEMALE. Specimens not available for this study; the following description

is quoted from Bohart (1953: 185): "Agreeing with male in general coloration.

Pleuron sometimes greenish, scutal integument medium to dark brown, scales

of abdominal tergites all dark. Palpus about one-fifth as long as proboscis,

torus with a small but sharp "'point" at the upper inner angle corresponding to

the pronounced knob of the male, no lower mesepimeral bristle."

106 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

MALE. Essentially similar to minor and other members of the minor

complex in antennae, palpal and labial characters, differing in the absence of

a lower mep bristle, paler coloration and slightly in the following features.

Head, All narrow decumbent scales of vertex predominantly pale, beige or

yellowish white and relatively broad along upper eye margin; erect scales

largely pale light brown. Palpus, Exceeding proboscis length by a little more

than the length of segment 5. Proboscis. Uniformly thick, basal 0.2-0.95 not

swollen. Antenna. As described and figured for minor; F-5 with a small in-

conspicuous tuft of 5 pale narrow, short acute scales or setae, about as long

as one flagellomere, followed laterally by 1,2 longer and broader scales; F-7

with a rather weak mesal tuft of 4,5 acute setae in addition to the usual tufts

of curled setae on ventral and lateral surfaces; F-9 with a strong tuft of 5, 6

dark flattened bristlelike setae.

MALE GENITALIA (Fig. 45). As figured. Segment LX, Tergal lobe with

3 weak setae; sternum without any setae or scales towards caudal margin.

Basimere, Normal; inner tergal surface with 6-8 moderately strong submar-

ginal setae in a row parallel to tergomesal margin, median area laterad of

submarginals with a prominent patch of numerous shorter setae. Subapical

Lobe. Rodlike setae a-c largely overlapping from bases to apices; leaflet 84

broad, long with acuminate apex; seta moderately strong and long; leaflet go

and setae d-f situated on a hairy elongate stem projecting mesad. Distimere.

Normal, slightly curved in middle, distally tapering into a small truncate apex;

subapical claw very short and small; ventral minute seta present, rather long;

dorsal one absent or not conspicuous. Phallosome. As in minor and bicornu-

tus except for relatively shorter internal process; external process with a dis-

tinct spiculose apical lobe and only a few denticles on lower tergal surface.

PUPA. Abdomen: 2.7mm. Paddle: 0.75 mm. Trumpet: 0.6 mm; in-

dex about 15. As figured for minor (Fig. 43). Cephalothorax and abdomen

whitish or cream-colored. Trumpet. Dark, long and rather thin. All setae

well developed, as in minor and bicornutus; the following are diagnostic. Ce-

phalothorax., Seta 7-C single or double; 8-C double; 9-C usually single (1-2).

Metanotum, Seta 10-C triple; 11-C double. Abdomen, Setae 1-TIII-VII double

or triple; 5-IV-VI single, 5-IV short; 5-V, VI 1.0-1.5 times as long as seg-

ment following; 6-III-VI usually double (1-2); 4, 5-VII single; 9-VII double or

triple, weakly pectinate; 9-VIII usually 4 branched (3-4), weakly pectinate.

Paddle: Very broad with characteristic elongate apex which is more or less

pointed or acuminate.

LARVA. Head: 0.76 mm. Siphon: 1.5 mm; index 8. Saddle: 0.30 mm;

siphon/saddle ratio 5. Head capsule, antenna, siphon and saddle pale, cream-

colored or yellowish white. In general as figured for minor (Fig. 44) with the

following distinctive features. Head, Seta 6-C single; 7-C 4, 5 branched.

Thorax. Spiculation entirely absent; seta 1-M short, 0.25-0.50 of 3-M. Abdo-

men. Spiculation absent; setae 6-I, II with 2 long lateral branches and 1 short

median branch; 7-II double; 6-III-VI usually double (1-3); 1-III-VI single or

double. Comb scales numerous, 60-70, all with normal fringe of evenly fine

spicules. Saddle very weakly spiculate on posterior caudal margin; seta 2-X

double; anal gills fusiform, as long as saddle. Siphon. Relatively thick and

straight; pecten teeth 11-13, larger distal teeth barbed with 2,3 distinct basal

denticles and 6,7 very fine distal denticles; subventral tufts 4 pairs, usually

triple (2-3) each, first proximal pairs 1.5 times as long as siphonal width at

point of attachment, the remaining pairs gradually shorter; median caudal fila-

ment of spiracular apparatus absent or not developed.

TYPE-DATA. Holotype “* with slide of antenna and genitalia, Chizuka,

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 107

Okinawa, Ryukyu Retto, | JAPAN], reared from larva collected in a rock hole

along a stream, Sept. 1945, R. Bohart and R. Ingram (USNM, No. 57760).

DISTRIBUTION. Okinawa, the Ryukyus and Chiang Mai, Thailand. 8 speci-

mens examined: 50, 3 L, 2 pupal skins.

THAILAND. Chiang Mai: Doi Sutep; 2% (M481-8; M559).

JAPAN. Ryukyus-Okinawa; Chizuka; E. Taira; 3c (including holotype);

> hs ome

TAXONOMIC DISCUSSION. The above description of the male of tuberis

has been based on the holotype and a few paratypes originally designated by

Bohart (1946: 42-3) and those of the pupa and larva have been based on the

specimens associated with the males subsequently described by the same author

(Bohart 1953: 185-6). In addition, I have also discovered 2 males of tuberis

among the specimens of minory in the collection of E. B. and D. C. Thurman

from Chiang Mai, Thailand. These males differ slightly from the Okinawan

Specimens in having finer and longer scales in the modified tuft of antennal

flagellomere 5 but are essentially similar to the latter in every detail of the

male genitalia.

Except for the specific diagnostic features as noted above and as indicated

in the keys, the male, male genitalia, pupa and larva of ftuberis are extremely

similar to those of minor and bicornutus with which it evidently falls into the

same complex. This, together with the presently known distribution, seems

to suggest that fuberis probably originated in Southeast Asia as a derivative of

minor and subsequently spread into the southern Ryukyus.

BIONOMICS. In Okinawa, the larvae and pupae of tuberis were collected

by Bohart (1953) in deep rock holes at the side of a cliff in association with

specimens of Uvanotaenia stonei Bohart and Ingram. All adults apparently

came from rearing larvae collected in the field. The 2 males from Thailand

were labelled as coming from sweeping with net.

37. CULEX (LOPHOCERA OMYIA) KUHNSI KING AND HOOGSTRAAL

(Fig. 46)

Culex (Lophoceraomyia) kuhnsi King and Hoogstraal 1955: 1 (o"*, 2, L*);

Sirivanakarn 1968: 75 (o"*, 9, P*, L*).

Culex (Lophoceraomyia) uniformis of Delfinado 1966: 114 (P*; L*; misidenti-

fication).

Culex (Lophoceraomyia) uniformis ssp. mercedesae Baisas 1974: 65 (o“*, &,

P, L*). NEW SYNONYMY.

FEMALE. Wing: 3.0 mm. Forefemur: 1.7mm. Proboscis: 1.8 mm.

Medium-sized, blackish species, exceedingly similar to minor and most mem-

bers of the minor complex, differing slightly in the following. Head, Narrow

decumbent scales of vertex entirely or predominantly pale whitish, scales on

upper eye margin relatively broad, clavate, forming a distinct ocular line;

erect scales entirely dark; lateral white patch of broad scales distinct. Cij-

barial Armature, As described for minor, Thorax. Lower mep bristle

present, usually 1, sometimes 2. Legs. Entirely dark or black scaled except

for the anterior surface of hindfemur which is broadly pale whitish in basal

0.25, gradually narrowed in apical portion.

MALE. As figured and described by King and Hoogstraal (1955: 1-4) and

Sirivanakarn (1968: 96-9); essentially similar to minor in antennal, palpal and

labial characters, differing slightly in the following. Antenna, F-5 with 6-9

108 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

short, narrow, acute scales; modified tufts of F-6,7 variable from weakly to

strongly developed; F-7 with a mesal row or tuft of weak acute setae in addi-

tion to other 2 tufts of curled setae on ventral and lateral surfaces; F-8 with a

weakly hooked tuft of 6,7 fused setae; F-9 with or without 2,3 weak bristlelike

setae.

MALE GENITALIA (Fig. 46). Essentially conforming to the minor com-

plex in the phallosome. Segment IX, Tergal lobe with 2,3 weak and short

setae; sternum with a transverse row of 6,7 strong setae towards caudal mar-

gin. Basimere, Normal; inner tergal surface slightly convex, with 3,4 rela-

tively weak submarginal setae in a row parallel to tergomesal margin, most

basal submarginal setae usually weaker than other 2,3 submarginals; short

setae laterad of submarginal moderately strong and dense. Subapical Lobe,

Rodlike setae d-c subequal, largely overlapping from bases to apices; leaflet

g1 broad, striated; seta 2 very strong, long and apically hooked. Distimere.

Normal. Phallosome. Internal process simple, stout and long, not projecting

beyond apex of external process; external process broad, with constricted api-

cal spiculose lobe sternad, tergal surface with several strong denticles in 2, 3

irregular rows, inner surface with variable number of toothlike marks. Proc-

tiger, Apical crown large; paraproct and cercal sclerite well sclerotized and

pigmented; cercal setae 3.

PUPA. Abdomen: 2.7mm. Paddle: 0.72 mm. Trumpet: 0.78 mm; in-

dex 10-12. In general as figured for minor (Fig. 43); cephalothorax and abdo-

men brownish or strongly yellowish. Trumpet, Slender, dark brown; apical

portion of meatus distad of annulation lightly spiculate. All setae developed,

the following are diagnostic. Cephalothorax, Seta 5-C stronger and longer

than 7-C, usually double (2-3); 8,9-C usually triple (3-4). Metanotum. Seta

10-C 3, 4 branched; 11-C always single. Abdomen. Setae 3-I-III single or

double; 5-IV, V single or double, subequal, 2 times as long as segment follow-

ing; 5-VI shorter, always single; 6-III-VI usually single (1-2); 9-VII 4-branched,

strongly pectinate; 9-VIII strong, usually with 10 strongly pectinate branches.

Paddle, Broad and entirely pale whitish, contrasting sharply with color of

abdomen; midrib weak, lightly pigmented.

LARVA (Fig. 46). Head: 0.74 mm. Siphon: 1.2 mm; index 9. Saddle:

0.26 mm; siphon/saddle ratio 5. Superficially very similar to uniformis and

spiculosus in having thorax and abdomen heavily spiculated, differing from the

latter 2 species and from all other members of the Mammilifer Subgroup in

the following characters. Head, Strongly yellowish to brownish; ventral sur-

face at level of ocular bulge’ with a broad band of numerous spicules; seta 1-C

dark, basally swollen, distally simple or forked into 2,3 apical spines, or

sometimes with 1 lateral spine on each side of large median spine; 5-C 2-4

branched; 6-C 1-3 branched; 12,13-C remarkably flattened and very dark; 12-C

3,4 branched; 13-C 4-6 branched. Antenna slender, relatively short and large-

ly pale yellowish; spicules numerous, rather fine and sharp. Thorax. Spi-

cules densely packed, hairlike, 4-6 branched; seta 8-P minute, similar to

spicules, usually single (1-2); 14-P usually single, sometimes double. Abdo-

men, Spiculation as in thorax; setae 6-I, II and 7-I dark, stout; 6-I, II usually

triple, sometimes 4-branched; 7-I double; 6-III-VI usually triple (8-4), medi-

an branch weaker and shorter than lateral branches; 1-III-VI strong, 0.5 of

length of 6-III-VI, double or triple. Comb scales about 40, scales on anterior

rows small, short, with normal fringes of spicules, those on posterior row

enlarged, terminating into a stout apical spine. Saddle usually dark brown

and strongly spiculate; posterior caudal margin densely packed with numerous

heavy spines; seta 2-X 3,4 branched, all branches subequally long; anal gills

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 109

slender, 2 times as long as saddle length. Siphon. Slender, moderately long,

yellowish or concolorous with head capsule; pecten teeth 15-20, denticulation

strongly differentiated, basal teeth barbed with 2-4 strong denticles, median

teeth with 1,2 denticles, 2-4 distal teeth usually simple; subventral tufts

strong, 4 pairs, 4,5 branched each, first 1,2 proximal tufts longest, 3,4 times

as long as siphonal width at point of attachment, next 2 distal tufts gradually

reduced in length; median caudal filament of spiracular apparatus absent or

not developed.

TYPE-DATA. (1) Culex (L.) kuhnsi King and Hoogstraal; Holotype o*

(766C) with slide of antenna and genitalia, Doromena, Hollandia (Kota Baru),

Netherlands New Guinea | West Irian, INDONESIA] reared from larva from a

shaded tree hole in open coastal hillside woods, 24 Feb. 1945, H. Hoogstraal

and W. H. Christ (USNM); (2) Culex (L.) uniformis ssp. mercedesae Baisas,

Type not specified; Mt. Makiling, Luzon, PHILIPPINES (LU).

DISTRIBUTION. Sabah, Malaysia; Philippines; Ceram, Indonesia and New

Guinea (including West Irian, Indonesia and Papua New Guinea). 93 specimens

examined: 440°, 222, 27 L; 41 with associated immature skins (7 p, 34 lp).

MALAYSIA. Sabah: Tawau; Keningau; 6c.

PHILIPPINES. Luzon; Bataan Province, Paysawan; Mt. Makiling; Subic

Bay (Naval Base); Calaca; Baguio; Dalton, N. Vizcaya; Mindoro: San Jose;

Negros: Fabrica Occidental; Leyte: Balinsasa Yao; Lago Lago Bay; Mindanao:

Kolambugan; Davao, Cotabato; Wenes; Mt. Apo; Jolo; Indanan; Mampallan,

Tali Pao; 380, 222, 26 L, 7p, 34 lp.

INDONESIA. West Irian: Kota Baru (Hollandia), Doramena; Cevam; 4%,

t Leal,

Additional records from the literature. PAPUA NEW GUINEA: Sepik

District, Vanimo (Sirivanakarn 1968: 96-100).

TAXONOMIC DISCUSSION. Culex kuhnsi is very widespread, with a

range extending from New Guinea to the Philippines and Sabah. Because of

the densely spiculose larva, kuhnsi has been repeatedly confused in the past

with uniformis,. As discussed under the latter, the Philippine records of uni-

formis by Delfinado (1966: 114) and Baisas (1974: 121, as subsp. mercedesae)

are in error, due to misidentifications. It is also possible that mindanaoensis

Baisas (1938) is in fact kuhnsi. However, as the type of the former no longer

exists the point cannot be settled. In comparing the Philippine specimens and

others currently available with the type and topotypic material from New

Guinea, I found them extremely similar in all stages and on this basis Iam

convinced that they all belong to kuhnsi.

All stages of kuhinsi are very variable, but there is no indication of local

differentiation among the material examined. The males from the Philippines

and Sabah differ slightly from those in New Guinea, particularly in the presence

or absence of weaker and fewer bristlelike setae in the modified tuft of anten-

nal flagellomere 9. The larva exhibits considerable individual variation in the

branching of setae 1,5,6-C and 6-I,II. The Philippine larvae differ slightly

from those in New Guinea in having the posterior comb scales terminating into

a stronger and longer median apical spine, otherwise both forms agree per-

fectly in general and in the detailed chaetotaxy. The pupae are also very vari-

able in pigmentation, but all apparently conform to the diagnostic chaetotaxy

as indicated in the above diagnosis.

BIONOMICS. The immatures of kuhnsi have been frequently collected from

tree holes and occasionally from bamboos at a broad range of elevation. In

the Philippines, they were also found in rock pools, discarded tin cans and

foot prints along stream margins and forest trails. In Sabah, only adults were

110 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

collected, presumably by sweeping with a net among foliage in the forest.

38. CULEX (LOPHOCERAOMYIA) CRASSICOMUS COLLESS

(Fig. 47)

Culex (Lophoceraomyia) crassicomus Colless 1965: 296 (o*).

FEMALE. Unknown.

MALE (Fig. 47). Based on the holotype. Essentially similar to other

members of the minor complex. Head. Narrow decumbent scales of vertex

largely pale beige, very narrow, linear and fine in center, slightly broader

on ocular line; erect scales slender and entirely blackish. Palpus. Slightly

longer than proboscis by a little less than the length of segment 09. Proboscis,.

Uniformly thick. Antenna. As described and figured by Colless (1965: 296);

spiculose prominence on pedicel more or less blunt, not clearly projecting

outwards; modified tufts of setae and scales present on F-5 to F-9; F-9 with

a small tuft of 2-4 dark, narrow, acute, spinelike scales about as long as the

next 2 flagellomeres, preceded to dorsally by 1,2 long normal setae; F-6, 7

with usual strong, dark, crumpled tufts of curled setae as in other species;

mesal tuft or row of short, narrow setae of F-7 poorly developed or indistinct;

F-8 with a relatively thick dark tuft of 7,8 fused setae in form of a typical J-

hook rather than sigmoid; modified tuft of F-9 with 4,5 dark, flattened bristle-

like setae and 3,4 other finer setae; middle portion of F-9 to F-11 with several

scattered hairlike setae as long as one flagellomere distad of normal whorls

of long setae.

MALE GENITALIA (Fig. 47). As described and figured by Colless (1965:

296). Segment IX. Tergal lobe with 2 weak setae; sternum with an irregular

transverse row of 5,6 strong setae on caudal margin. Basimere. Normal;

inner tergal surface with 4,5 submarginal setae in a linear row parallel to

tergomesal margin; marginal setae weak and rather sparse; short setae later-

ad of submarginals on outer surface weak and relatively few in number. Sub-

apical Lobe. Rodlike setae a-c largely overlapping; seta a@ thick, distinctly

broadened and strongly curved upwards in apical portion; leaflet g; narrow,

acuminate; seta h moderately long; setae d-f consist of 2,3 narrow bladelike

setae; leaflet g9 long, rodlike and strongly bent in middle. Distimere. Slender,

normal. Phallosome, Internal process strong, but its apex not projecting be-

yond apex of external process; external process with distinct spiculose lobe

sternad, its upper tergal surface with only few denticles or tubercles.

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype “* with slides of antenna (CH 152) and genitalia

(CT 571), Ulu Gombak, Selangor, Malaya | MALAYSIA], March 1956, D. H.

Colless (ANIC).

DISTRIBUTION. Known only from Selangor and Sabah, Malaysia. 3° ex-

amined.

MALAYSIA. Peninsular Malaysia - Selangor: Ulu Gombak (type-locality);

10° (holotype); Malaysia - Sabah: Tawau; 2c.

TAXONOMIC DISCUSSION. In addition to the holotype, I observed 2 other

males of cvassicomus in the BMNH. They were collected by D. H. Colless at

Tawau, Sabah, during February 1960. One of these agrees perfectly with the

holotype but the other differs in having fewer and thicker curled setae on the

flagellomeres 6,7 and in the complete absence of scattered setae in the middle

of antennal flagellomeres 9-11. In the male genitalia, the phallosome of the

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region LT

second male also differs from the typical specimen in having more numerous

denticles on the upper tergal surface of the external process. It is possible

that this specimen probably represents a distinct species, but pending further

study of additional material, it is now tentatively identified with cvassicomus,

The male of cvassicomus can be separated from other members of the

minor complex in the antenna by the form of the modified tufts of flagellomeres

o and 8 and the presence of several scattered setae distad to the normal whorls

of flagellomeres 9-11; in the male genitalia by the presence of 4, 5 submarginal

setae on the basimere, the characteristic development of rodlike seta a of the

subapical lobe and by the presence of only a few denticles or tubercles on the

external process of the phallosome. The male genitalia of Ccvassicomus are

apparently more similar to bengalensis and peytoni than to any other species

in the Mammiilifer Subgroup, suggesting that they are closely related.

BIONOMICS. The males of cvassicomus were collected by sweeping

among foliage (Colless 1965: 296). The breeding site and adult biology are un-

known.

39. CULEX (LOPHOCERA OMYIA) INCOMPTUS

BRAM AND RATTANARITHIKUL

(Fig. 48)

Culex (Lophoceraomyia) incomptus Bram and Rattanarithikul 1967: 16 (o*);

Bram 1967a: 81 (o*).

FEMALE. Unknown.

MALE (Fig. 48). Exceedingly similar to other members of the minor

complex, differing slightly in the following. Antenna, Flagellar whorl weakly

plumose or with relatively fewer normal long setae; F-5 with a small, distinct

tuft of 5,6 dark, narrow, acute scales, as long as the next 2 flagellomeres,

preceded dorsally by 1 normal long seta, 4,5 dorsal scales subequal, 1,2

ventralmost scales slightly longer; F-6 with a strong crumpled tuft of several

curled setae; F-7 with a lateral combed-tuft of several curled setae as in

most forms, a slender ventral tuft of weak fused setae and a weak mesal tuft

(or row) of fine acute setae; F-8 with a slender, smooth J-hooked tuft of 7, 8

fused setae, some of which are distinctly swollen at about middle before taper-

ing into fine points; F-9 with 2-4 dark, slender bristlelike setae; F-8-11 with

2,3 very short setae at middle of flagellomere distad of bases of normal

whorls.

MALE GENITALIA (Fig. 48). Segment IX. Tergal lobe with 3-5 moder-

ately strong setae; sternum with 1 strong seta towards caudal margin at mid-

dle. Basimere. Normal; submarginal setae moderately strong, 6,7 in num-

ber, ina linear row accompanied by a number of shorter setae along tergo-

mesal margin. Subapical Lobe, Rodlike setae a-c subequal and largely over-

lapping; leaflet g; well developed; seta 4 moderately strong; 1,2 weaker setae

present laterad and mesad of seta /; leaflet go strong, rodlike; setae d-f weak,

hairlike. Distimere. Normal; subapical dorsal surface with a weak crest of

very fine spicules, somewhat similar to mammilifer, Phallosome. Well

sclerotized; internal process strong, not projecting beyond apex of external

process, the latter with several strong denticles on upper tergal surface and

a distinct spiculose apical lobe sternad; inner surface with a number of distinct

toothlike marks proximally. Proctiger., Essentially similar to other members

of the minor complex; cercal setae 2, 3.

112 Contrib. Amer.’ Ent, Inst. vol,- 13, noo 4, 1977

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype “* (M405-4) with slides of antenna and genitalia,

Doi Sutep, Chiang Mai, THAILAND, 7 Jan. 1953, D. C. and E. B. Thurman

(USNM; No. 69186).

DISTRIBUTION. Known only from Chiang Mai, Thailand. 40 examined.

THAILAND. Chiang Mai: Doi Sutep (type-locality); Doi Tad Fah; 4% (in-

cluding holotype).

TAXONOMIC DISCUSSION. Culex incomptus can be readily separated from

other members of the minor complex in the male antenna by the presence of 2,

3 short setae distad of normal whorls of flagellomeres 8-11 and by the differ-

ences in the modified tufts of flagellomeres 5-8. The male genitalia of zmcomp-

tus resemble mammilifer in the presence of a fine crest of spicules on the dor-

sal subapical surface of the distimere but its phallosome is similar to minor

and other members of the minor complex. Based on the latter feature, incomp-

tus is here assigned to the minor complex.

BIONOMICS. The males of incomptus, including the holotype, were caught

while resting among vegetation in a forest at a very high elevation in the moun-

tain region. The associated larvae, which were lost, were collected from a

tree hole (Bram and Rattanarithikul 1967: 17).

40. CULEX (LOPHOCERA OMYIA) BENGA LENSIS BARRAUD

(Fig. 49)

Culex (Lophoceratomyia) minor var. bengalensis Barraud 1934: 371 (co),

Culex (Lophoceraomyia) bengalensis Barraud, Colless 1965: 293 (“*, 2, L);

Bram 1967a: 83: (o*, 2, L*).

FEMALE. Wing: 3.0 mm. Forefemur: 1.3 mm. Proboscis: 1.5 mm.

Differing from other members of the minor complex particularly in the absence

of a lower mep bristle, relatively slender size and in having decumbent scales

of vertex broad clavate along upper eye margin.

MALE (Fig. 49). Differing from other members of the minor complex as

described for female and as in the following. Palpus. Exceeding proboscis

length by at least full length of segment 5; segments 4 and 5 apparently with

more numerous strong bristles on lateral and mesal surfaces. Pyroboscis,

Uniformly slender and thin. Antenna, Flagellar whorls densely plumose;

distinct modified tufts of setae developed only on F-6-9; F-5 usually without

any modified setae or scales, sometimes with 2,3 inconspicuous short fine

hairlike setae among the dense row of normal long setae; F-6, 7 with crumpled

tufts of curled setae as in mammilifer or minor; F-8 with a slender typical

sigmoid hooked tuft of 6 fused setae; F-9 with 3,4 weak bristlelike setae.

MALE GENITALIA (Fig. 49). Essentially conforming to the minor com-

plex. Segment Lx, Sternum usually without any setae, sometimes with a

transverse row of 4 strong setae towards caudal margin. Basimere, Very

slender, inner tergal surface with a row of 4,5 moderately strong submar-

ginal setae parallel to tergomesal margin; marginal setae weak and sparse;

short setae laterad of submarginals weak, short and rather few in number.

Subapical Lobe. Rodlike setae a-c slender, largely overlapping from base to

apices; rod a distally expanded and curved upwards as in cvassicomus; leaflet

g1 club-shaped; seta i moderately strong; leaflet go rodlike, straight; setae

d-f slender, simple, bladelike. Distimere. Subapical dorsal surface without

crest of fine spicules. Phallosome. Internal process strong and long but not

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 113

projecting beyond apex of external process, latter with a constricted apical

spiculose lobe sternad and 6-10 strong denticles on its upper tergal surface,

inner surface with or without toothlike marks. Pyroctiger. As in minor.

PUPA (Fig. 49). Abdomen: 2.3mm. Paddle: 0.63 mm. Trumpet: 0. 62

mm; index 12-15. Cephalothorax and abdomen pale creamy or yellowish white.

Trumpet. Dark, slender, long, uniformly cylindrical; pinna with distinct slit

extending into meatus. Chaetotaxy as figured. Cephalothorax, Seta 1-C usu-

ally 4,5 branched; 5-C subequal to 7-C, usually 5 branched (4-7); 8-C usually

double (2-3); 9-C double or triple. Metanotum, Seta 10-C usually double

(1-3); 11-C double. Abdomen, Setae 5-IV-VI as long as or slightly longer

than segment following; 5-IV 4-6 branched; 5-V, VI double; 6-III- VI usually

4 branched (3-5); 9-VII usually triple (2-4); 9-VIII usually 5, 6 branched (4-7).

Paddle, Broad, pale whitish or almost transparent; midrib weak, lightly

pigmented.

LARVA. Head: 0.65 mm. Siphon: 1.7 mm; index 9. Saddle: 0.27 mm;

siphon/saddle ratio 6. As figured for mammilifer (Fig. 34), with the follow-

ing diagnostic features. Head, Pale creamy or yellowish white; setae 5, 6-C

double. Antenna largely pale, same color as head capsule; spicules numerous

and strong. Thorax, Spiculation absent; seta 7-P usually double (2-3); 8-P

double, subequal to 7-P; 14-P double. Abdomen. Setae 6-I, II triple; 6-III-VI

usually triple (3-4); 1-III-VI usually triple (2-4). Comb scales 40-50, all of

which are rather minute, subequal, with normal or rounded apical fringe of

evenly fine spicules. Saddle very lightly spiculate on posterior caudal mar-

gin; seta 2-X double. Szphon. Slender, same color as head; pecten teeth 12,

13 all apparently barbed with 7,8 graded denticles, basal denticles not differ-

entiated from distal ones; subventral tufts 3 pairs, widely spaced, double or

triple each, most proximal pair twice as long as siphonal width at point of

attachment, the remaining distal pairs gradually shorter; median caudal fila-

ment of spiracular apparatus well developed.

TYPE-DATA. Lectotype <* (only antenna (1451) and genitalia (1452) slides,

the remainder of the specimen is completely lost), Nongpoh, Shillong, Assam:

INDIA, June 1922. P. J. Barraud (BMNH; selection of Colless 1965: 293).

DISTRIBUTION. India, Thailand, Malaysia and Indonesia; also reported

from Hainan, China. Not known elsewhere. 71 specimens examined: 41°,

222, 8 L; 28 with associated immature skins (15 p, 13 lp).

INDIA. Assam: Shillong, Nongpoh (type-locality); 10° (lectotype).

THAILAND. Nan; Ranong: Muang; Kraburi; Phangnga: Tang Mai; Phuket:

Khao Prathin; Kvabi: Ban Mai Kaen Tai; Khao Chaung; Narathiwat: Khau Lau;

lOc, 32, 41,6 p, 1 ip.

MALAYSIA. Peninsular Malaysia - Pahang: Gunong Benom; 260, 199,

7p, 12 lp. Malaysia - Sabah: Tawau; Mt. Kinabalu; 2c, 4L.

INDONESIA. Kalimantan: Tarakan; 2%. .

Additional records from the literature: CHINA. Hainan (Chu 1957: 145-63

as var. of minor); MALAYSIA. Selangor: Ulu Gombak; Trengganu: Gunong

Tebu (Colless 1965: 293-4); INDIA. North Bengal: Sukna and Marianbarrie

Tea Estate (Barraud 1934: 372, as var. of minor).

TAXONOMIC DISCUSSION. Culex bengalensis is differentiated from the

preceding species in the minor complex by the combination of the male anten-

nal characters and genitalia as described above and as indicated in the key.

The male genitalia are exceedingly similar to cvasstcomus from which it can

be separated by the presence of more numerous denticles on the upper tergal

surface of the external process of the phallosome. Both the pupa and larva

of bengalensis exhibit considerable overlap and extreme similarity to mam-

114 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

mtlifer in general and in the detailed chaetotaxy. It can be separated from the

latter in the larva by the double seta 14-P and the stronger and longer subven-

tral tufts of siphon. No clear-cut characters have been found to separate the

pupa of bengalensis from that of mammilifer except for the usual number of

branches of a few setae as given in the key.

All the above records of bengalensis, including those from the literature

are valid except from Hainan, China by Chu (1957; 1958), which is doubtful

and requires confirmation.

BIONOMICS. The immatures of bengalensis came largely from collections

made in general ground pools (including stream pools), foot prints, puddles

and wheel tracks at the edges of small streams or rivers in mountainous areas

at an elevation ranging from 500-1,500 m. Ona few occasions, they were also

collected from rock pools and holes in fallen logs lying on the ground. The lar-

vae and pupae from stream pools have frequently been found in association with

specimens of mammilifer, All adults apparently came from rearing the pupae

and larvae.

peytoni complex

41. CULEX (LOPHOCERAOMYIA) PEYTONI

BRAM AND RATTANARITHIKUL

(Figs. 50, 51)

Culex (Lophoceraomyia) peytoni Bram and Rattanarithikul 1967: 7 (“*, 2, L*);

Bram 1967a: 100 (“*, 2, L*).

FEMALE. Wing: 3.2 mm. Forefemur: 1.6 mm. Proboscis: 1.9 mm.

Medium-sized, blackish species, exceeding similar to the members of minor

complex from which it can not be distinguished with certainty. Head, Narrow

decumbent scales of vertex very fine, pale, whitish along upper eye border,

forming a narrow ocular line, scales in center predominantly blackish or

sometimes pale yellowish; all erect scales blackish; lateral white patch minute,

rather indistinct. Cibarial Armature. Cibarial bar with about 40 elongate

teeth in a concave row. Thovax, Mesonotal scales blackish and rather

coarse; 1 lower mepbristle. Legs, Wing and Abdomen, Essentially similar

to minor and its closely related species.

MALE (Fig. 50). Extremely similar to bengalensis, differing particularly

in the following details. Antenna. Modified tuft of F-5 absent or poorly de-

veloped, with or without 1,2 short, fine hairlike setae as long as the combined

length of the next 2 flagellomeres among several normal long setae, sometimes

with only 1 long, very narrow acute scale; modified tuft of F-6, 7 poorly or

well developed; F-7 with a distinct mesal row of 6-8 short, straight or distally

curved setae in addition to crumpled tufts of curled setae on lateral and ven-

tral surfaces; F-8 with a slender or thick tuft of 4-6 fused setae which are

rather straight or slightly curved subapically, but not strongly hooked or sig-

moid as in most forms; F-9 with 2-4 weak or strong bristlelike setae; setae

distad or normal whorls of flagellomeres 9-11 practically absent.

MALE GENITALIA (Fig. 50). Exceedingly similar to bDengalensis and

cvassicomus; differing in the following features. Segment LX, Sternum al-

ways with a transverse row of 7,8 strong setae towards caudal margin. Basi-

mere. Inner tergal surface usually with 4 relatively weaker submarginal setae

in a row parallel to tergomesal margin, sometimes 5; marginal setae and

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 415

setae laterad of the submarginals strongly reduced and very sparse. Subapical

Lobe, Setae d-f well developed and more conspicuous, 3,4 in number; leaflet

g1 broad, striated and acuminate. Phallosome, Poorly sclerotized; internal

process rather weak and short; external process very broad, upper tergal sur-

face irregular, with a few to several weak denticles; apical tergal portion

strongly expanded or produced into a blunt point, projecting tergad in lateral

aspect; apical lobe (or knob) reduced and weakly spiculate.

PUPA (Fig. 50). Abdomen: 2.4mm. Paddle: 0.61 mm. Trumpet: 0.60

mm; index 8-10. Exceedingly similar to ganapathi, differing from it in the

following combination of characters. Cephalothorax and abdomen usually yel-

lowish to dark brownish, sometimes lighter. Tvumpet, Dark brown and re-

latively thick; apical portion of meatus distad of proximal annulation moder-

ately spiculate; pinna without slit extended to meatus. Cephalothorax, Seta

1-C double or triple; 5-C usually double, dark, as long as 7-C; 8,9-C usually

double (1-3). Metanotum. Seta 10-C double; 11-C always single. Abdomen,

Setae 3-I-III single; 3-III long and dark; 5-IV-VI usually single (1-2) and re-

markably long; 5-IV, V 1. 5-2.0 or more of segment following; 5-VI as long as

or Slightly longer than segment following; 6-III-VI weak, subequally long, usu-

ally double (2-3); 9-VII 2-4 branched; 9-VIII 6-9 branched. Paddle, Very

broad and pale.

LARVA (Fig. 51). Head: 0.78 mm. Siphon: 1.6 mm; index 8,9. Saddle:

0.30 mm; siphon/saddle ratio 5,6. Essentially conforming to the Mammilifer

Subgroup with general and detailed chaetotaxy resembling ganapathi, differing

from it in the following combination of characters. Head, Integument yellow-

ish white with or without dark brownish areas; seta 1-C slender, simple,

dark, spiniform; 5, 6-C always double; 7-C 8,9 branched. Antenna usually

pale yellowish, sometimes darkened; spicules moderately strong. Thorax,

Lightly spiculate; spicules minute, dotlike and moderately dense; seta 7-P

double; 8-P single, subequal to 7-P; 14-P always double. Abdomen. Spicula-

tion absent; setae 6-I, II and 7-I usually stout and dark, sometimes pale; 6-I, II

triple; 7-I double; 6-III-VI subequal, usually double, sometimes triple; 1-III-VI

variable in length; 1-III usually triple (2-3); 1-IV-VI usually double (2-3).

Comb scales 35-40, all subequal in size and with normal apical fringe of evenly

fine spicules. Saddle lightly to moderately infuscated; posterior caudal mar-

gin lightly spiculate; seta 2-X usually double (2-3); anal gills 1-3 times as

long as saddle length. Siphon. Usually yellowish, sometimes darker, but not

brownish; pecten teeth 11-14; large distal teeth with 1,2 strong, basal denticles

and 6,7 fine, graded, distal denticles; subventral tufts moderately strong, 4

pairs; first 3 proximal pairs double or triple, 1.5-2.0 times as long as siphonal

width at point of attachment; most distal pair shortest, 3-5 branched; median

caudal filament of spiracular apparatus not developed.

TYPE-DATA. Holotype “* (PU8-45) with associated larval and pupal

skins and slide of antenna and genitalia, Patthalung, THAILAND, tree hole,

12 feet (approximately 4 m) above the ground, 15 Oct 1964, K. Mongkolpanya

(USNM; No. 69182).

DISTRIBUTION. Thailand, Vietnam, Malaysia, Indonesia, Andaman

Islands (India). 124 specimens examined: 570", 32°, 35 L; 69 with associated

immature skins (27 p, 42 lp).

THAILAND. Mae Hong Son: Doi Chang; Chiang Mati: Doi Sutep; Nan: Ban

Wang Mo; Khon Kaen: Phu Wiang; Chanthaburi: Khao Hin Phaeng; Chumpon:

Ban Wang Phi; Ranong: Koh Chang; Ban Bang Hin; Khlong Bang Man; Phangnga:

Tak Khet; Tang Mai; Thap Wen; Nakhon Si Thammarat: Ban Thuan Lek; Ban

Sai Kae; Trang: Muang, National Park; Ban Kan Tang Tai; Phatthalung: Muang

116 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

(type-locality); Navathiwat: Waeng; 250, 149, 35 L, 23 p, 17 lp.

VIETNAM. Pleiku, Plei Djereng; Kontum; 7°, 29.

MALAYSIA. Peninsular Malaysia - Pahang: Mela K. Lipis; The Gap;

Perlis: Kg. Wang Tangga; Padang Besar: Kg. Wang Kelian; Kedah: Sintok,

F. R202... 199, 4 D...25. (p.

INDONESIA. Java: Ciran Jang, Ciloto; Sumatva: Bengkoelen; 4c, 19,

2 lp.

INDIA. Andaman Islands; 1°“ (slide of antenna and genitalia only).

TAXONOMIC DISCUSSION. Culex peytoniand eukrines are very closely

related and apparently fall into a complex which combines the features of the

minor complex in the male antenna and genitalia and of the ganapathicomplex .

in the pupa and larva. Except for the additional details described above, the

males and larvae of peytoni essentially agree with the original descriptions

and figures by Bram and Rattanarithikul (1967: 7-9). All stages of peytoniare

very variable. Caution should be taken in separating the males of peytoni

from those of bengalensis and crassicomus, and the larvae and pupae from

those of ganapathi, The most diagnostic features of the male genitalia of

peytoni are the well developed setae d-f of the subapical lobe; the very broad

and poorly sclerotized external process of the phallosome and the weakly spicu-

late apical lobe of the external process. The pupa resembles ganapathi in the

detailed chaetotaxy but differs from it in having thicker trumpet, shorter and

weaker setae 5-C and 6-III-VI. The larva differs from ganapathi particularly

in the simple and more slender seta 1-C and in having stronger basal denticles

of the pecten teeth.

In addition to the records from Thailand, peytoni is also known from other

neighboring countries, as noted above, indicating that it is one of the very

widespread species in the Mammilifer Subgroup.

BIONOMICS. The common breeding sites of peytoni are either tree holes

or bamboos in tropical rain forests. Ona few occasions, the larvae have also

been found in fallen coconut shells. Bram and Rattanarithikul (1967: 9) noted

that it was once collected in a rock hole. These breeding sites were located

at a broad range of elevation.

42. CULEX (LOPHOCERAOMYIA) EUKRINES

BRAM AND RATTANARITHIKUL

(Fig. 52)

Culex (Lophoceraomyia) eukrines Bram and Rattanarithikul 1967: 11 (o*, &,

L*); Bram. 1967a: 85.(¢*,.2, L*).

FEMALE. As described for peytoni to which it is almost identical, differ-

ing slightly in having narrow decumbent scales of vertex largely pale yellow,

forming a broader ocular line on anterior margin and erect scales varying from

partially yellowish brown or golden to entirely black. Cibarial Armature,

Similar to peytonifrom which it is indistinguishable.

MALE (Fig. 52). Exceedingly similar to peytoni, differing chiefly in the

following. Antenna. Flagellar whorls more densely plumose; F-5 with or

without 1, 2 fine hairlike setae among several normal long setae; F-6, 7 with

strongly reduced or poorly developed tufts of curled setae; mesal setae of F-7

finer and longer, projecting beyond F-8; modified tuft of F-8 more slender,

smoothly curved or gently hooked subapically; F-9 with 4, 5 weak bristlelike

setae.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 117

MALE GENITALIA (Fig. 52). As figured, almost identical to peytoni,

differing slightly in the following. Basimere, Usually with 5 submarginal

setae in an irregular row, most basal submarginal weak and short, the re-

maining setae strong, subequal in length. Swubapical Lobe. Rodlike setae a-c

stronger; distal part with several long fine hairlike setae which are densely

packed at bases of setae d-f and leaflet 85. Phaliosome, Well sclerotized;

external process moderately broad with several strong, dark denticles on

upper tergal surface.

PUPA. As described and figured for peytoni (Fig. 50), differing from it

particularly in the following characters. Cephalothorax. Seta 1-C 4 branched;

0-C 3,4 branched. Abdomen, Setae 3-I, II usually single, sometimes double;

6-III-V usually triple (2-4); 9-VIII 5-7 branched.

LARVA (Fig. 52). Differing from peytoni in the following combination of

characters. Head, Seta 5, 6-C flattened and relatively short; 5-C double;

6-C always single; 7-C weaker and shorter, 4-9 branched. Antenna shaft with

considerably weaker and fewer spicules. Thorax. Spiculation absent or in-

conspicuous; seta 7-P usually double, sometimes single. Abdomen. Seta 6-I

usually triple, sometimes 4 branched; 6-III, IV usually triple (2-4); 6-V, VI

double; 1-III-VI weak and short, 3,4 branched. Seta 2-X of saddle usually

double, sometimes single. Siphon. Relatively thick, covered with numerous

minute, dotlike spicules; distal pecten teeth with weaker basal denticles; sub-

ventral tufts weaker and shorter, usually 4 pairs, sometimes with 1 additional

tuft distad of pecten; pairing very irregular, 3,4 branched each; most proxi-

mal pair longest, about 1.5 times as long as siphonal width at point of attach-

ment, the remaining pairs gradually shorter.

TYPE-DATA. Holotype o* (0181-11) with associated larval and pupal

Skins and slide of antenna and genitalia, Huey Bong Ti, Kanchanaburi, THAI-

LAND, rock hole, 2 June 1965, K. Mongkolpanya (USNM; No. 69183).

DISTRIBUTION. Known only from Thailand. 146 specimens examined:

280°, 232, 95 L; 48 with associated immature skins (31 p, 17 lp).

THAILAND. Nan: Doi Chick Chong; Ban Pha Man; Kanchanaburi: Sai

Yok Waterfall; Ko Chum Rong Village; Huey Mae Nam Noi: Ban Sai Yok; Huey

Bong Ti (type-locality); Khao Saeng; Khao Nga Chang; Chanthaburi: Khao Wong,

dha, Mat: Phanonsa:: 269-239-951 81 ip li Ip.

TAXONOMIC DISCUSSION. Culex eukrines is particularly known from

Kanchanaburi, Thailand where the types and numerous topotypic specimens

were originally and subsequently collected. Its distribution is very localized

and apparently more restricted than that of peytoni. Except for records from

Thailand, this species is not known elsewhere in Southeast Asia.

Although the males of eukrines and peytoni slightly differ from each other

in the antenna and genitalia, their differences in the larvae are clear-cut and

constant, as indicated in the key and in the above description. The most

diagnostic features of the eukrines larvae are the single flattened seta 6-C and

the reduction in size of the spicules of the antennal shaft. In the males,

eukrines differs from peytoni in the reduction of the modified tufts of the anten-

na and in the presence of several densely packed long hairlike setae associated

with seta d-f of the subapical lobe of the basimere of the genitalia. The differ-

ence in the breeding sites of eukrines and peytoni are also well marked; some-

what resembling that between those of minor and bicornutus; eukrines appar-

ently exhibits a tendency to breed in rock holes or pools instead of bamboos

or tree holes, whereas the reverse is true of peytoni.

All of the specimens of eukrines which I have examined essentially con-

form to the above descriptions except for a single larva from Ban Pha Man in

118 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Nan (Coll. 01439, from a crab hole) which differs from the typical form in hav-

ing strong spicules on the antenna and in having 1, 2 simple or undenticulate

pecten teeth distad of normal teeth of the siphon. Without the associated adult

male, this larva is tentatively identified with ewkrines although it is possible

that it may represent a distinct species.

BIONOMICS. The immatures of eukvines have been frequently collected

from rock holes or pools along the margin of mountain streams under heavy

or partial shade of tropical forest. Of the 14 larval collections made in Thai-

land, 8 were from rock pools, 2 from tree holes and one each from bamboo, a

coconut shell, a wheel track and a crab hole. The larvae from rock pools were

abundant and have been collected in numbers at several localities in Kanchana-

buri. All adults apparently came from rearing the larvae or pupae.

pholeter complex

43. CULEX (LOPHOCERA OM YIA) PHOLETER

BRAM AND RATTANARITHIKUL

(Figs. 53, 54)

Culex (Lophoceraomyia) pholetey Bram and Rattanarithikul 1967: 13 (Sx, 2,

L*); Bram 1967a: 103 (o*, 9, L*).

FEMALE. Wing: 2.7mm. Forefemur: 1.5mm. Proboscis: 1.8 mm.

Relatively small species with general external features exceedingly similar to

mammilifer and bengalensis, differing in the following characters. Head. De-

cumbent scales on anterior margin of vertex relatively broader, pale whitish,

forming a broad ocular line; narrow decumbent scales restricted to center,

rather coarse, crescent-shaped and predominantly dark; erect scales usually

entirely dark, sometimes partially pale yellowish on posterolateral area;

lateral patch of broad scales whitish. Cibarial Armature. Very distinctive;

cibarial bar with a concave row of about 18 teeth, 4 median teeth narrow,

tapering into sharp points, remaining lateral teeth broad, flattened, apically

blunt or truncate. Thorax. One lower mep bristle present. Wing. Scales on

all veins more numerous and rather dense. Abdomen. Sterna yellowish white

scaled, contrasting sharply with entirely black scales on terga.

MALE. Very similar to mammilifer, differing from it in the following

characters. Palpus. Rather short, varying from slightly shorter to as long as

proboscis; segments 4,5 reduced in length, each with only a few short bristles.

Antenna, Flagellar nore weakly to moderately plumose; modified tuft of F-5

composed of 3-5 dark scales which are variable from narrow, linear or lanceo-

late to broad, abruptly acuminate, as long as the combined length of the next

2,3 flagellomeres; dorsalmost scale usually broad, apically blunt or pointed;

remaining scales narrow, hairlike or lanceolate; modified tufts of F-6-8 well

developed, as in mammilifer; F-9 with 3,4 weaker bristlelike setae.

MALE GENITALIA (Fig. 53). As described and figured by Bram and

Rattanarithikul (1967: 13); essentially conforming to the Mammilifer Subgroup

with the following diagnostic features. Segment IX, Tergal lobe very poorly

developed, bearing 1-3 tiny setae; sternum with a transverse row of 5 strong

setae towards caudal margin. Basimere, Inner tergal surface with a linear

row of 3 submarginal setae near base; marginal setae and other short setae

laterad of submarginals and subapical lobe very sparse and weak. Subapical

Lobe. Rodlike seta a short, distally tapered into a sharp point, its length 0. 50-

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 119

0.75 of rods 0,c; the latter very slender, rather long and apically hooked;

setae d-f rather narrow and weak; leaflet &; very broad, long and striated, its

apex blunt or acuminate; seta h moderately strong. Distimere. Dorsal sur-

face with distinct ridges or folds distad of median curvature; dorsal subapical

crest of spicules absent. Phallosome. Poorly sclerotized, as in peytoni;

internal process rather short or markedly reduced in size; external process

broad, with 2 irregular rows of several distinct denticles on upper tergal sur-

face; apical spiculose lobe (or knob) very small and short. Proctiger. As in

most forms, 2 cercal setae present.

PUPA (Fig. 53). Abdomen: 2.7mm. Paddle: 0.6 mm. Trumpet: 0.55

mm; index 10-15. Cephalothorax and abdomen pale whitish to yellowish.

Trumpet, Dark, slender; apical portion distad of proximal annulation of mea-

tus strongly spiculate; pinna with distinct slit extended to meatus. Complete

chaetotaxy as figured, distinctive in the following. Cephalothorax.. Seta 5-C

very strong, 2-3 times as long as 7-C, usually double (2-3); 8-C double; 9-C

usually double (1-2). Metanotum. Seta 10-C usually double (2-3); 11-C double.

Abdomen, Setae 3-I-III double; 5-IV usually double (2-3), 1.5 times as long

as segment following; 9-V, VI single, very strong, 2 times as long as segment

following; 6-III-VI usually double (2-3); 9-VII usually triple (3-4); 9-VIII strong,

twice longer than 9-VII, 4-8 branched, 3,4 median branches reaching 0.5 of

paddle length. Paddle. Broad, pale whitish to almost transparent; midrib

weak and lightly pigmented.

LARVA (Fig. 54). Head: 0.78 mm. Siphon: 1.2-1.8 mm; index 7.

Saddle: 0.30 mm; siphon/saddle ratio 5. As figured by Bram and Rattanarithi-

kul (1967: 8) and Bram (1967a: 104); in general similar to most forms, but with

the following diagnostic features. Head, Integument pale yellowish or cream-

colored; seta 1-C simple, pale yellow or dark; 4-C variable from 1.2-2.5

times as long as distance between bases of the pair; 5,6-C strong, subequal,

double; 7-C 5-8 branched. Antennal shaft pale, cream-colored or slightly

darkened; spicules strong and dark. Thorax, Spiculation absent; seta 7-P

usually double, sometimes triple; 8-P minute, 2-5 branched, of the same

magnitude as 14-P; the latter usually double, sometimes triple. Abdomen,

Spiculation absent; setae 6-I, II and 7-I dark, stout; 6-I, II usually with 2 long

lateral branches and 1 short median branch, sometimes median branch absent

or not developed; 7-I double; 6-III-VI usually double (1-3); 1-III-VI variable

in length, usually single or double, sometimes triple. Comb scales rather

numerous, usually 60 or more, sometimes fewer; all scales with normal api-

cal fringe of evenly fine spicules. Saddle weakly spiculate on posterior caudal

margin; pale yellowish, sometimes slightly darker; seta 2-X usually double

(2-3), shorter branch nearly as long as longer branch; 2 proximal pairs of setae

of ventral brush more or less detached from grid; anal gills as long as saddle.

Siphon. Moderately long, pale yellowish or same color as head capsule; pecten

teeth 8-10, larger distal teeth with 1,2 strong basal denticles and 5, 6 finer

distal denticles; subventral tufts 4 pairs, each usually double (1-2), sometimes

4,9 branched, length variable, usually weak and short, sometimes very strong

and long; most proximal pair distad of pecten as long as siphonal width at point

of attachment, the remaining pairs gradually reduced in length; median caudal

filament of spiracular apparatus present.

TYPE-DATA. Holotype “* (00416-2) with associated larval and pupal

skins and slide of antenna and genitalia, Khao Mai Ha Wa, Chon Buri: THAI-

LAND, from a crab hole in a mountainous secondary rain forest, 19 July 1965,

E. L. Peyton (USNM; No. 69184).

DISTRIBUTION. Known only from Thailand. 88 specimens examined:

120 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

140°, 72, 67 L; 18 with associated immature skins (13 p, 5 lp).

THAILAND. Chon Buri, Khao Mai Ha Wa (type-locality). Bang La Mung,

Khao Mai Kaeo; Khao Sai Dao; Nakhon Ratchasima: Khao Suan Hom; Khai Phai;

14°, 72, 41 L, 13 p, 5 1p; Ranong: Ban Chatri; Phangnga: Thap Wen; Nam

Tai; Tang Mai; 26 L.

TAXONOMIC DISCUSSION. Culex pholeter is apparently very variable, but

can be readily recognized in the male, male genitalia, pupal and larval stages

by the characters as given in the keys and as indicated in the above diagnosis.

In addition, pholeter also appears to be rather distinct ecologically. It differs

from almost all other species of the Mammilifer Subgroup in having a tendency

to breed in crab holes in preference to other types of breeding sites, including

general ground pools. Most of the larvae from Thailand came from crab holes

except for those from Ranong and Phangnga, which were collected in cut bamboo

lying on the ground and bamboo stumps. The material from bamboo differs

slightly or strongly from that in the crab holes in the darker pigmentation and

in having stronger and more numerous branched subventral tufts of the siphon.

It is likely that more than one form is involved. However, as the bamboo

material consists only of whole larvae, they are here tentatively identified as

pholeter, pending further study of associated males when they become avail-

able.

Because of the distinctive male characters, including the genitalia, pholeter

appears to belong to a distinct complex of the Mammilifer Subgroup. Its af-

finity is not clear and cannot be accurately determined, but it appears to be

intermediate between the mammilifer and peytoni complexes. In the male,

the modified tuft of antennal flagellomere 5 somewhat resembles that of mam~

milifer but is less developed. The male genitalia is more or less similar to

peytoni but the internal process of the phallosome is more strongly reduced in

length and size. In the immatures, the pupa resembles mammialifer in the

trumpet by having pinna slit extended to meatus and double seta 11-C but is

similar to peytoni in the development and branching of seta 0-IV-VI. The lar-

ya is the most diagnostic stage, particularly in having seta 8-P strongly re-

duced to the magnitude of seta 14-P; otherwise it resembles peytoni in general

and detailed chaetotaxy.

BIONOMICS. The larvae and pupae of pholeter from Chon Buri and Nak-

hon Ratchasima were mostly collected in crab holes on the bank of mountain

streams at elevations of 150-1,400 m. On occasion, they were also taken in

stream pools, tree holes and foot prints. Of the 20 larval collections made at

the above localities; 13 were from crab holes, 4 from stream pools, 2 from

tree holes and one from a foot print. Most of the larvae from Ranong and

Phangnga were collected in split bamboos lying on the ground and only a few

from bamboo stumps, at an elevation of 100-200 m.

flavicornis complex

44, CULEX (LOPHOCERAOMYIA) FLAVICORNIS BARRAUD

(Figs. 55, 56)

Culex (Lophoceratomyia) flavicornis Barraud 1924: 45 (o*, 2); Edwards 1932:

198 (taxonomy); Barraud 1934: 375 (o*, 9).

FEMALE. Wing: 4.7mm. Forefemur: 2.3 mm. Proboscis: 2.7 mm.

As described by Barraud (1924: 45; 1934: 375-6); in general essentially con-

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 121

forming to the Mammilifer Subgroup with the following diagnostic features.

Size medium to large. Head, Decumbent scales of vertex entirely narrow

and pale yellowish; erect scales slender, entirely blackish and moderately

dense. Czibarial Armature, Cibarial bar with a concave row of 50-60 narrow,

elongate teeth which are subequal in size and length and distally tapered into a

fine point. Thorax, Mesonotal scales predominantly blackish except for some

pale yellowish ones on extreme anterior promontory. Lower ™eép bristle ab-

sent. Legs, Entirely blackish except for anterior surface of hindfemur which

is entirely pale whitish in basal 0.4-0.5, continuing as narrow pale stripe in

apical 0.5-0.6. Wing. Scales on veins Ro, Rg and R4,5 narrow, linear. Ab-

domen, Terga entirely dark; sterna pale yellowish.

MALE (Fig. 55). In general as in female, differing in smaller size. Pal-

pus. Long, exceeding proboscis length by about 0.75 of the length of segment

0; segments 2,3 with several rows of numerous long, hairlike setae on ventral

and mesal surfaces, longest ones about 4,5 times as long as segment width at

point of insertion; lateral surface of segment 3 with a distinct row of 15-20 dark,

long hairlike setae extending from base to apex; segments 4 and 5 upturned,

with several short, weak setae and a few bristles; segment 5 elongate, about 2

times as long as segment 4. Proboscis, Lightly swollen in basal 0.2-0.5

and with a double row of 12-16 strong setae on ventral surface. Antenna.

Pedicel with a strong spiculose prominence on inner dorsal surface; flagellar

whorls weakly to moderately plumose; modified tufts of scales and setae pre-

sent on F-0 to F-9; F-5 with a large, brushlike yellowish or golden tuft of 12-

15 equally long fine hairlike setae which are as long as the combined length of

the next 5, 6 flagellomeres; F-6, 7 with strong, dark, crumpled tufts of curled

setae as described for most forms; F-8 with a strong sigmoid hooked tuft as

in mammilifer and minor; F-9 with a dark tuft of 4-6 strong bristlelike setae;

proximal portions of F-9-11 with 1-3 short, weak setae distad of whorls.

MALE GENITALIA (Fig. 55). Segment LX, Tergal lobe with 3,4 relatively

strong setae; sternum with several strong setae towards caudal margin. Basi-

mere, Rather large, about 0.30 mm in length; basal portion broad, outer ter-

gal margin strongly convex or swollen, apical portion narrow; inner tergal mar-

gin (or surface) strongly concave, with a close-set row of 8,9 strong, flattened

submarginal setae; marginal setae 5,6, strong, about 0.2 of submarginals,

placed among the latter; short setae laterad of submarginals weak and sparse.

Subapical Lobe, Prominent; rod a very thick with characteristic expanded

twisted apex resembling a hammer head; rods b,c slender and apically hooked,

distal portion of 0 somewhat wavy or kinked; leaflet go with characteristic

striation, resembling a tree leaf; seta 2 moderately strong; leaflet 81 rodlike,

distally curved and apically hooked; setae in group d-f represented by one fine

blade, placed close to leaflet g1. Distimere. Very thick, with modified,

broadly expanded truncate apex; median curvature pronounced; subapical dor-

sal surface recurved, with distinct crest of fine spicules; ventral tiny seta

present, dorsal one absent; claw small, slender, placed almost at tip of the

recurved apex. Phallosome. Well sclerotized; lateral plate with a short,

simple internal process and a weakly, denticulate external process, the latter

with a distinct spiculose apical lobe sternad and 15-17 weak denticles in 2,3

irregular rows on upper tergal surface. Proctiger. Apical crown medium

sized, with several flat and blunt spicules laterally and numerous finer spine-

like spicules mesally (or internally); paraproct and cercal sclerite moderately

broad, well sclerotized; 3 minute cercal setae.

PUPA (Fig. 56). Abdomen: 2.9mm. Paddle: 0.90 mm. Trumpet: 0.75

mm; index 10. Cephalothorax brownish on wing, leg and labial cases, head

122 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

shield and area along mid-dorsal ridge, pale yellowish on other areas; metano-

tum brownish; abdomen largely brownish on segments I-III, pale cream-colored

on segments IV-VIII. Trumpet. Dark brown, cylindrical and long; pinna with

slit extending to meatus. Complete chaetotaxy as figured; in general conform-

ing to the Mammilifer Subgroup, distinctive in the following. Cephalothorax,

Seta 1-C usually 4 branched (3-5); 5-C usually 5 branched (3-6), subequal to

7-C; 8-C 2-5 branched; 9-C usually triple (1-3). Metanotum, Seta 10-C

usually 3,4 branched (2-4); 11-C always double. Abdomen, Setae 3-I-III

double; 5-IV-VI double, 5-IV, V subequal, 1.5 times as long as segment follow-

ing, 5-VI slightly shorter; 6-III-VI usually 3,4 branched (2-5); 9-VII, VIII weak,

subequal; 9-VII usually double or triple (2-4); 9-VIII usually 6,7 branched (5-

8); 4-VIII double. Paddle. Broad, pale whitish, with characteristic bluntly

pointed apex, somewhat resembling a leaf; midrib weak and pale; apical por-

tion minutely spiculate; setae 1,2-P present, distinct.

LARVA (Fig. 56). Head: 0.75 mm. Siphon: 1.9 mm; index 11; saddle:

0.36 mm; siphon/saddle ratio 5.5-6.0. Essentially conforming to the Mam-

milifer Subgroup, distinctive in the following features. Head, Setae 5,6-C

double; 14-C double or triple. Antennal shaft yellowish brown with numerous

strong spicules. Mental plate with 8,9 lateral teeth on each side of median

tooth. Thovax. Lightly spiculate, sometimes spiculation not developed or in-

distinct; setae 7, 8-P subequally strong, double; 14-P single; 1-M as long as

or longer than 3-M, single; 8,9-M 4, 5 branched; 1-T longer than 2-T, single;

7-T 5-7 branched; 9-T 5 branched; 13-T 4,5 branched. Abdomen, Spiculation

absent; 6-I, II and 7-I dark, stout; 6-I, II usually triple (2-3); 7-I usually double

(1-2); 6-III-VI triple; 1-IJI-VI strong, nearly as long as 6-III-V, usually 4

branched (3-4); 13-III-V shorter than 1-III-VI, 4 branched. Comb scales nu-

merous, 50-60, all apparently subequal, with normal fringe of evenly fine

spicules. Saddle yellowish brown, posterior caudal margin weakly spiculate;

seta 2-X double; anal gill shorter or as long as saddle. Siphon. Relatively

long and slender, distally slightly tapered; yellowish or same color as saddle;

pecten teeth 13-18, median teeth with a strong basal denticle and 3-5 finer

distal denticles; 4,5 distal teeth rather widely spaced, with or without a fine

barb of 3-5 graded denticles; subventral tufts 4 pairs, all weak, subequal,

usually double, 1.0-1.5 times as long as siphonal width at point of attachment;

median caudal filament of spiracular apparatus well developed.

TYPE-DATA. Lectotype “*, Nilgiri Hills, Kasauli, Madvas, INDIA,

October 1915, Khazan Chand (BMNH; selection of Bram 1967b: 328).

DISTRIBUTION. Known only from India. 10 specimens examined: 6°,

4°: 6 with associated immature skins (1 p, 5 lp).

INDIA. Madras; Kasauli; Nilgiri Hills; 60 (including lectotype), 49, 1p,

5 lp. Also reported from Kodaikanal, Palni Hills by Barraud (1934: 376).

TAXONOMIC DISCUSSION. Culex flavicornis is strongly differentirated

from all other species in the Mammilifer Subgroup in the male by the promi-

nent rows of numerous fine setae and bristles on the ventral, lateral and mesal

surfaces of palpal segments 2 and 3 and the large, yellowish or golden brush-

like tuft of long setae on antennal flagellomere 5; and in the male genitalia by

the shape of the basimere, the modified apex of distimere, the setae and leaf-

lets of the subapical lobe, the reduction of the internal process of the phallo-

some and the relatively coarse spicules of the proctiger crown. The female

is generally very similar to most species but can be recognized by the absence

of a lower mesepimeral bristle and by the presence of 50-60 narrow fine teeth

in the cibarial armature. The pupa somewhat resembles fuberis in having the

apex of the paddle more or less pointed but differs from it in the darker pig-

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 123

mentation of cephalothorax, pinna of trumpet with slit extending to meatus,

setae 5-IV-VI double and 6-III-VI usually 3,4 branched. The larva closely

resembles mammilifery in having seta 8-P double and 14-P single but differs

from it in the darker pigmentation, setae 1-III-VI usually 4 branched and cs auiee

with 4 pairs of subventral tufts.

Within the Mammilifer Subgroup, flavicornis is apparently most closely

similar to vaghavanii Rahman, Choudhury and Kalra (1968 (1969): 28-34). Both

were described from the same locality in the Nilgiri Hills, Madras, India.

However as I have not seen the specimens of the latter, it is not possible to

determine whether or not it is really distinct from flavicornis, From the

figures and descriptions by the above authors, vaghavanii apparently differs

from flavicornis in the male genitalia by having unmodified apex of distimere,

different shape of leaflet £1 of the subapical lobe and in the larva by the heavily

spiculose thorax and abdomen. It is possible that the larva of vaghavanii is

incorrectly associated and belongs to uniformis,

BIONOMICS. Culex flavicornis is restricted to high elevations in moun-

tainous areas. At Palni Hills, Kodaikanal, India, it was recorded from an

elevation of 7, 000 ft. (approximately as 300 m) (Barraud 1934: 376). Inthe

Nilgiri Hills, Madras, the larvae have been collected from rock pools, con-

taining numerous fallen leaves under heavy shade of a tropical forest. They

were found in association with larval specimens of Culex (Culiciomyia) pallido-

thorax Theobald.

49. CULEX (LOPHOCERAOMYIA) LASIOPALPIS NEW SPECIES

(Fig. 57)

FEMALE. Wing: 2.18 mm. Forefemur: 1.44 mm. Proboscis: 1.7 mm.

Small, slender, blackish species; in general similar to flavicornis in the ab-

sence of a lower mep bristle, differing in smaller size and in the following

features. Head, Narrow decumbent scales on anterior margin of vertex

broader and pale whitish, scales in the center blackish. Cibarial Armature.

Cibarial bar with a concave row of 30-34 teeth, lateral ones apparently coarser

or more flattened with abruptly pointed apices.

MALE (Fig. 57). Essentially similar to flavicornis in the antennal modi-

fied tufts and in the presence of numerous densely packed hairlike setae on

lateral, ventral and mesal surfaces of palpal segments 2 and 3, differing from

it as follows. Palpus, Shorter, exceeding proboscis by about 0.5 of segment

0; segments 4,5 subequally long. Proboscis, Uniformly thin with fewer setae

on ventral surface. Antenna. F-5 with a smaller yellowish or golden brush-

like tuft of 8-10 long setae.

MALE GENITALIA (Fig. 57). As figured, differing from /lavicornis strik-

ingly in the following features. Basimere, Normal, slender, conical, about

0.2 mm in length; submarginal seta weaker and shorter, 5-7 in number. Sub-

apical Lobe, Rod a distally tapering into a point; rodlike seta 2) more slender;

setae in group d-f consist of 1,2 fine setae, 1 strong, serrate blade and 2

other shorter and simple blades: leaflet £; Simple, broad lanceolate, apical

foliate portion unstriated. Distimere. Slender, normal, apex unmodified:

dorsal subapical crest absent. Phallosome. Apical spiculose lobe of external

process reduced or poorly developed; upper tergal surface of external process

with about 15 strong denticles in a dense triple row. Proctiger. Apical

crown smaller and with fewer spicules; paraproct and cercal scerite narrower.

PUPA (Fig. 57). Abdomen: 2.2 mm. Paddle: 0.52 mm. Trumpet: 0.57

124 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

mm; index 10-12. Complete chaetotaxy as figured, differing from /lavicornis

in the following. Trumpet. Thinner. Metanotum. Seta 10-C always double.

Abdomen. Seta 5-IV usually triple (2-4); 6-II-VI usually 4 branched (4-6).

_ Paddle. Apex evenly rounded or slightly produced.

LARVA. Head: 0.65 mm. Siphon: 1.44 mm; index 10. Saddle: 0.29

mm; siphon/saddle ratio 5. As figured and described for flavicornis, differ-

ing from it in the following features. All setae apparently weaker. Thorax,

Spiculation absent. Abdomen. Spiculation absent; seta 1-II triple; 1-IV-VI 4

branched. Comb scales 40-50. Siphon. Shorter with lower index and siphon/

saddle ratio; pecten teeth 10-12, all with fine barb of 6,7 graded denticles;

subventral tufts weaker, 3 pairs, more widely spaced, as long as or slightly

longer than siphon width at point of insertion.

TYPE-DATA. Holotype “ (126-8) with associated pupal and larval skins

and slide of antenna and genitalia, Matale, Matale District. Central Province,

SRI LANKA, foot print in seepage, elevation 915 m, 26 June 1975, E. L. Pey-

ton and Y.-M. Huang (USNM). Allotype 2 (126-9) with associated pupal and

larval skins and slide of cibarial armature (USNM); paratypes: 1 lp (126-6),

2 po (126-101, 117), 1 p? (126-104), same data as holotype; 1 lp (220-3), 6

L (220); 1 lpo’ (221-8), 4 L (221), Kanneliya, Galle District, Sabaragamuwa

Province, elephant foot print in marshy depression, elevation 245 m, 10 July

1975, other data as in collection No. 126 (USNM, BMNH).

DISTRIBUTION. Known only from Sri Lanka. 18 specimens examined:

50°, 32, 10 L; 8 with associated immature skins (3 p, 5 lp), as indicated in the

type-data. :

TAXONOMIC DISCUSSION. Culex lasiopalpis is provisionally recognized

as distinct from vaghavanii Rahman, Choudhury and Kalra (1968 (1969): 28-34)

from the Nilgiri Hills, Madras, India, pending a thorough comparison with the type-

male of the latter. The male of lasiopalpis apparently agrees well with the

description and figure of vaghavanii in the features of antenna and palpus and

in several details of the genitalia, but differs from it in having simple, broad

lanceolate leaflet g; of the subapical lobe (broad, triangular in vaghavanit) and

in the presence of 5-7 submarginal setae on the basimere (8-9 in vaghavanii).

In the immature stage, the pupa of lasiopalpis differs from the description and

figure of vaghavanii in the evenly rounded apex of the paddle (pointed in vagha-

vanit); the larva differs in the absence of the spiculation on the thorax and abdo-

men and in the presence of 3 pairs of the subventral tufts of the siphon (4 pairs

in vaghavanii). As discussed under flavicornis, the association of the adults

and immature stages of vaghavanii may possibly be incorrect and certain fea-

tures of its male genitalia were perhaps erroneously described. A detailed

study of the type and additional topotypic specimens of vaghavanii are critically

needed to definitely establish its identity.

Culex lasiopalpis is most closely related to flavicornis and evidentally falls

into a complex with the latter species. It can be readily separated from /flavi-

cornis in the male, pupa and larva by the features as given in the keys and in

the above description.

BIONOMICS. Culex lasiopalpis is apparently rare. In Sri Lanka the imma-

tures have been taken twice, from animal foot prints in a seepage and from a

marshy depression under partial shade of tropical rain forests, both in moun-

tainous areas at elevations of 915 m and 245m. All adults came from rearing

pupae and larvae.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 125

Brevipalpus Subgroup

The Brevipalpus Subgroup is characterized in the adults by the color of

erect scales of vertex varying from entirely dark to partially pale golden or

bronzy in center, dark on posterolateral areas; lower mep bristle absent; in

the female cibarial armature by the presence of 15-25 coarse teeth (except

jensent); in the male by the relatively short palpus which is usually from 0. 50-

0.75 of proboscis length, although sometimes as long as or slightly longer

than proboscis; in the male genitalia by (1) basimere varying from small,

slender, conical to large, stout or modified, without prominent row of sub-

marginal setae; (2) setae of subapical lobe as described for the group or

strongly differentiated in number and shapes; leaflet § 1 usually absent, some-

times present; (3) distimere normal or with strongly modified apex; (4) inter-

nal process of phallosome poorly or well developed; in the pupa by trumpet

relatively short, bell-shaped or funnel-shaped, varying from 0.2-0.4 mm in

length and in the larvae by (1) seta 2, 3-A placed apically on the antenna; (2)

setae 16,17-C absent; (3) siphon slender or thick, length varied from shorter

than to 2-3 times as long as saddle and (4) subventral tufts of siphon usually

6-8 pairs (total 16), sometimes 3. Breeding habitats: exclusively pitcher

plants.

DISCUSSION. The Brevipalpus Subgroup comprises several highly special-

ized forms whose breeding habitats are restricted to pitcher plants. Certain

members of this subgroup, including particularly jenseni, navalis and coeru-

lescens are morphologically closer to the members of the minor complex than

any others in the Mammilifer Subgroup. The group as a whole appears to be

an offshoot from the latter through specialization in the breeding habitats. In

the Oriental region, the Brevipalpus Subgroup is largely restricted to Malaysia

(including Peninsular Malaysia and Borneo), Singapore and other neighboring

islands in Indonesia with only a few extending into the north in Thailand, Viet-

nam and Hong Kong.

All 10 species in the Brevipalpus Subgroup are apparently well known except

for sumatranus Brug which had not, until recently been transferred from Neo-

culex to Lophoceraomyia by Sirivanakarn (1971). They can be readily classi-

fied into 5 more or less distinct complexes: navalis, hewitti, jenseni, brevi-

palpus and curtipalpis as characterized below. |

(1) navalis complex, represented by navalis and coerulescens, It is

characterized in the male by (1) palpus as long as or longer than proboscis

and (2) modified tufts of scales and setae on antennal flagellomeres 5-9 well

developed, as described for the Mammilifer Group; in the male genitalia by

(1) basimere small, conical, with 1 strong submarginal seta or none: (2) sub-

apical lobe with 3 rodlike setae in group a-c, 3 bladelike setae in group d-f, and

1 basal seta (1); leaflet g1 absent and (3) distimere slender, normal; in the

pupa by (1) trumpet funnel-shaped, 0.40-0.45 mm in length; (2) seta 8-C weak,

double; (3) seta 5-IV 2-4 branched and (4) 9-VIII 7,8 branched; in the larva by

(1) antenna nearly as long as head length; (2) seta 1-A strongly plumose, 15 or

more branched; (3) seta 1-C dark, spiniform; (4) setae 5, 6-C long, single or

double; (5) comb scales numerous, 30-35; (6) siphon slender, lightly swollen

in middle, 2-3 times as long as saddle and (7) subventral tufts of siphon 7, 8

pairs.

(2) hewitti complex, including only hewitti. It is characterized by the male

by (1) palpus 0.9 of proboscis length; (2) modified tufts of antennal flagello-

meres 0-9 minute or rudimentary; and (3) F-6 with a characteritstic tuft of 2-4

126 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

subapically swollen setae; in the male genitalia by (1) basimere and subapical

lobe as in vavalis complex except for the absence of submarginal seta and (2)

distimere thick, with modified truncate apex; in the pupa by (1) trumpet as in

navalis complex; (2) seta 8-C usually triple; (3) 5-IV long, single and (4) 9-VIII

8 branched; in the larva by (1) antenna nearly as long as head; (2) seta 1-A

weakly plumose, 9-11 branched; (3) seta 1-C pale, spiniform; (4) setae 5-C

5-9 branched, 6-C single; (5) comb scales 50; (6) siphon slender, distally

tapered, about 4 times as long as saddle and (7) subventral tufts of siphon 7

pairs.

(3) jensenit complex, represented only by jenseni. It is intermediate,

resembling the navalis complex in the male, including the genitalia, but is

similar to the brevipalpus complex in the pupa and larva. It is characterized

in the male particularly by palpus 0.6-0.7 of proboscis length; in the male

genitalia by subapical lobe and distimere as in navalis complex; in the pupa by

(1) trumpet somewhat oval or oblong in shape, 0.3-0.4 mm in length; (2) seta

8-C single; (3) seta 5-IV single and (4) 9-VIII 5-7 branched; in the larva by (1)

seta 1-A weak, 3-6 branched; (2) seta 1-C dark, stout, spiniform; (3) setae 5,

6-C long, each 2-4 branched; (4) comb scales 24-27 in number; (5) siphon thick,

relatively short, 1.7-1.8 of saddle length and (6) subventral tufts very strong,

6, 7 pairs.

(4) brevipalpus complex, represented by brevipalpus, lucaris, eminentia

and acutipalus. It is characterized in the male by (1) palpus 0. 50-0. 75 of

proboscis length and (2) modified tufts on antennal flagellomeres 5-9 well

developed, as in navalis and jenseni complexes; in the male genitalia by (1)

basimere broad, stout, somewhat rectangular in lateral aspect, submarginal

seta not developed; (2) subapical lobe broad, strongly differentiated, with nu-

merous strong setae associated with rods a-c and setae d-h, leaflets absent

and (3) distimere with modified truncate apex; in the pupa by (1) trumpet bell-

shaped, 0.25-0.30 mm in length; (2) seta 8-C strong, single; (3) seta 5-IV

long, single and (4) seta 9-VIII long, single; in the larva by (1) antenna short,

about 0.5 of head length; (2) seta 1-A weak, 4-6 branched; (3) seta 1-C pale,

distally filamentous; (4) setae 5,6-C markedly flattened and short, placed

cephalad, 5-C 2-4 branched, 6-C single; (5) comb scales 20-30; (6) siphon

thick, 2-3 times as long as saddle and (7) subventral tufts strong, 6-9 pairs.

(5) curtipalpis complex, represented by curtipalpis and sumatranus, It

is characterized in the male by (1) palpus 0.5 of proboscis length and (2) modi-

fied tufts of scales and setae present on antennal flagellomeres 5-9 or com-

pletely absent; in the male genitalia by (1) basimere broad, stout, somewhat

oval, without differentiated submarginal setae; (2) subapical lobe with a broad

external leaflet (g,) in addition to usual specialized setae as in navalis com-

plex and (3) distimere thick, with modified truncate apex; in the pupa by (1)

trumpet short, small, typically bell-shaped, 0.3 mm in length; (2) seta 8-C

single; (3) seta 5-IV single, moderately long and (4) seta 9-VIII 4, 5 branched;

in the larva by (1) antenna short, about 0.5 of head length; (2) seta 1-A weak,

3, 4 branched; (3) seta 1-C short, stout, pale, spiniform, placed on distinct

tubercle; (4) setae 5,6-C short, fine, placed cephalad, each 2-4 branched;

(5) comb scales 5-8 and (6) siphon markedly reduced, about 0.75 of saddle and

(7) subventral tufts of siphon weak, 3 pairs.

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 127

navalis complex

46. CULEX (LOPHOCERA OMYIA) NA VA LIS EDWARDS

(Figs. 58, 59)

Culex (Lophoceratomyia) navalis Edwards 1926: 121 (o'*, L); Edwards 1928:

277 (**); Edwards and Given 1928: 353 (L*); Edwards 1932: 198 (taxono-

my); Brug and Bonne-Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) navalis Edwards, Mattingly 1949: 226 (L*); Stone,

Knight and Starcke 1959: 235 (catalog); Colless 1965: 304 (o"*, 9, L).

FEMALE. Wing: 3.0 mm. Forefemur: 1.6mm. Proboscis: 1.9 mm.

Small, slender, blackish species; essentially conforming to the Brevipalpus

Subgroup characters. Head, Decumbent scales of vertex entirely narrow,

linear, scales on upper anterior margin pale whitish, forming a distinct ocular

line; scales in the center yellowish brown or golden to black; all erect scales

black; lateral patch of broad scales whitish or grayish. Czibarial Armature

(Fig. 58). Cibarial bar with concave row of about 20 stout teeth; 3-4 median

teeth narrow with apices tapered into sharp point; lateral teeth widely spaced

with abruptly pointed or truncate apices. Thovax, Pleuron without scale

patches; lower mep bristle absent; pp/ with 2 strong dark bristles and a few

yellowish short setae. Legs, Anterior surface of hindfemur with a distinct

white stripe extending from base to apex, rest entirely blackish. Wing.

Scales very narrow, linear and moderately dense. Abdomen. Terga entirely

black scaled; sterna pale yellowish or silvery white scaled.

MALE (Fig. 58). Exceedingly similar to members of Mammilifer Sub-

group, differing slightly in the following characters. Palpus. Usually as long

as proboscis, sometimes slightly longer. Pyroboscis, Uniformly thin and

slender. Antenna, Spiculose prominence of pedicel strong and distinct; flagel-

lar whorls moderately to strongly plumose; F-5 with a distinct yellowish or

golden tuft of 4,5 long, narrow, somewhat lanceolate scales, as long as the

next 2 flagellomeres, preceded dorsally by 2 short fine, hairlike setae; F-6, 7

with strong, dark, crumpled tufts of curled setae as in members of the Mam-

milifer Subgroup; F-8 with a typical sigmoid hooked tuft of 6, 7 fused setae;

F-9 with a prominent dark tuft of 6,7 strong bristlelike setae; F-9-11 without

any setae distad of normal whorls of long setae.

MALE GENITALIA (Fig. 58). As figured. Segment IX. Tergal lobe very

poorly developed, bearing 1,2 minute, hairlike setae; sternum with 6 strong

setae in an irregular transverse row towards caudal margin. Basimere,

Slender, normal, conical, about 0.2 mm in length; inner tergal surface with

or without 1 strong submarginal seta near base; marginal setae sparse and

weak; stronger bristles largely restricted to lateral tergal surface, median

lateral area with relatively small number of short and weak setae. Subapical

Lobe. Distinctive; rodlike seta a stout, strongly curved in middle, separated

from rods b,c by a wide gap; setae b, c overlapped at bases with those of

setae d-f on a distinct lobe distad; leaflet 21 absent or replaced by a weak,

short seta distad of 895 seta h weak and rather short; leaflet 85 Slender, blade-

like, nearly as long as setae b,c; setae in group d-f consist of 2 narrow and

fine blades, accompanied by variable number of rather indistinct fine hairlike

setae. Distimere. Slender, simple, about 0.14 mm in length, distal portion

slightly or strongly swollen before tapering to a short, recurved apex; sub-

apical claw very short and small; dorsal and ventral subapical setae rather

128 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

strong, subequal, opposite each other. Phallosome. Unpigmented, poorly

sclerotized; internal process slender, simple, much shorter than external

process, the latter with a constricted spiculose apical lobe (or knob) sternad,

its upper tergal surface with 15 or fewer weak denticles. Proctiger. Apical

crown medium-sized, largely consisting of strong, spinelike spicules; 3-6

cercal setae.

PUPA (Fig. 58). Abdomen: 2.3 mm. Paddle: 0.52 mm. Trumpet: 0.45

mm; index 8. Cephalothorax and abdomen usually uniformly yellowish white,

with or without indefinite brownish areas. Tvumpet, Dark brown, relatively

short, more or less resembling funnel in shape, meatus uniformly cylindrical,

pinna usually flared. Complete chaetotaxy as figured, most setae relatively

weak and short. Cephalothorax, Seta 5-C weak, as long as or shorter than

7-C, 5,6 branched; 8,9-C weak, double. Metanotum. Seta 10-C 3,4 branched;

11-C usually double (1-2); 12-C usually triple (3-4). Abdomen, Setae 3-I-III

usually double (1-2); 5-IV-VI slightly shorter or longer than segment following;

5-IV usually triple (2-4); 5-V usually double (1-2); 5-VI usually single (1-2);

6-III- VI double or triple; 9-VII usually 4 branched (3-5); 9-VIII apparently

placed near caudolateral angle, usually with 8 strongly plumose branches (7-9).

Paddle. Very pale to almost transparent; moderately broad and long; apical

margin rounded and rather indistinct; midrib lightly yellowish and distinct; api-

cal setae 1,2-P minute, very inconspicuous.

LARVA (Fig. 59). Head: 0.65 mm. Siphon: 0.60 mm; index 5,6. Saddle:

0.26 mm; siphon/saddle ratio 2,3. As figured; in general similar to the mem-

bers of the Mammilifer Subgroup with the following diagnostic features. Head,

Integument pale, cream-colored; seta 1-C dark, slender, spiniform and rela-

tively long, nearly as long as distance between bases of the pair; 4-C very fine,

single or double; 5-C usually single, sometimes double; 6-C always single;

7-C weakly plumose, 5,6 branched. Antennal shaft largely pale, with several

strong, spinelike spicules; seta 1-A with about 15 or more pectinate branches;

2,3-A placed apically. Mental plate with 8 lateral teeth on each side of medi-

an tooth. Thovax., Spiculation not developed or absent; seta 1-P double; 3-P

usually triple (3-4); 4-P usually 6 branched (5-7); 7-P usually triple (2-3);

8-P usually double (1-2), subequal to 7-P; 1-M minute, single, about 0.20 of

3-M; 8, 9-M 7-9 branched; 7,9-T 8-10 branched. Abdomen. Setae 6-I, IT usu-

ally with 5(4-5) and 4 (3-4) strongly pectinate branches; 7-I single; 6-III-VI

usually double, sometimes triple; 1-III-VI weak, short, usually triple (2-4);

1-VII 3,4 branched. Comb scales narrow, elongate, about 34, all with normal

apical fringe of evenly fine spicules. Saddle yellowish to brownish, posterior

caudal margin not spiculate; setae 1,2-X single; ventral brush dark, composed

of 6 pairs of setae, all usually inserted within grid, most proximal pair some-

times detached; anal gills long, fusiform, 2-4 times as long as saddle. Siphon.

Yellowish to dark brown, moderately long, slender, more or less fusiform in

shape; pecten teeth 6-8, larger distal teeth narrow, elongate and sharply

pointed with 6,7 fine denticles of graded size; subventral tufts remarkably

strong and dense, 7,8 pairs, 5,6 branched each, closely spaced, forming a

prominent double row; first 3,4 proximal pairs longest, subequal, 3,4 times

as long as siphonal width at point of attachment, next 3,4 pairs gradually re-

duced in length; dorsal and ventral valves of spiracular apparatus poorly differ-

entiated, bearing a strong, hooked seta (9)on its apex; median caudal filament

absent. |

TYPE-DATA. Lectotype “* (marked as type), reared from pitcher plants,

SINGAPORE, 1926? D. H. C. Given (BMNH; present selection).

DISTRIBUTION. Known from Singapore, Malaysia and Indonesia. 68 speci-

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 129

mens examined: 370, 192, 12 L; 8 with associated immature skins (1 p, 7 lp).

SINGAPORE. 17° (including lectotype), 62, 3 L, 4 1p;3 0 (A. R. Barr

collection).

MALAYSIA. Peninsular Malaysia - Selangor: Pacific Tin, K. Sel; Kg.

Tanjong Rabok; Pahang: Pekan Rd.; Kedah; Sintok F. R.; 26, 139, 5L,

1D, ao. Ube

INDONESIA. Kalimantan: Tarakan; 1, 4 L.

TAXONOMIC DISCUSSION. Culex navalis can be readily separated from

all other pitcher plant species except coevulescens in the male, including the

genitalia, pupa and larva by the characters as indicated in the Keys and in the

above descriptions. It can be distinguished from coerulescens particularly by

the development of the modified tufts of the male antenna, the setae of the sub-

apical lobe of the basimere of the male genitalia, the color of antenna of the

larva and by a few other features indicated in the keys and descriptions. Cau-

tion should be taken in separating the 2 species. Because of the absence of the

lower mesepimeral bristle, the adults of navalis can be easily mistaken for

bengalensis and other species of the Mammilifer Subgroup. Except for the

differences in the detail of the modified tuft of the male antenna and genitalia,

a great deal of similarity in the adult morphology is evident between ”avalis

and the members of the minor complex of the Mammailifer Subgroup. This

evidence suggests that it was probably directly or indirectly derived from cer-

tain forms of the latter subgroup through specialization in the breeding habitat.

BIONOMICS. Culex navalis is apparently not uncommon as it has been taken

on several occasions at the various localities within its reported range. In

Singapore, the larvae were collected from Nepenthes ampullaria (Barr and

Chellapah 1963: Colless 1965). All collections recently made in Peninsular

Malaysia were also reported from this pitcher plant species. They were found

in association with the larvae of coerulescens, hewitti and acutipalus., In

western Borneo, it was reported to be taken from Nepenthes bicalcarata

and vaffleziana (Brug 1934). All adults apparently came from rearing the

larvae or pupae.

47. CULEX (LOPHOCERAOMYIA) COERULESCENS EDWARDS

(Fig. 60)

Culex (Lophoceratomyia) coerulescens Edwards 1928: 277 (o'*, ); Edwards

and Given 1928: 355 (L); Edwards 1932: 197 (taxonomy); Brug and Bonne-

Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) coerulescens Edwards, Mattingly 1949: 226 (L); Stone,

Knight and Starcke 1959: 232 (catalog); Colless 1965: 305 (o“*, 2, L).

FEMALE. Exceedingly similar to mavalis from which it can not be dis-

tinguished with certainty, differing slightly in having finer and largely pale

decumbent scales in the center of vertex. Cibarial Armature, Apparently in-

distinguishable from “avalis,

MALE (Fig. 60). Essentially similar to mavalis, differing particularly in

the following details. Palpus., Length usually equal to proboscis. Antenna,

Pedicel apparently with weaker spiculose prominence on its inner dorsal sur-

face; modified tuft of F-5 reduced, rather inconspicuous, consists of only a

few short fine hairlike setae among several normal long setae; modified tufts

of F-6, 7 reduced or more weakly developed; F-8 with a smaller tuft of 4,5

fused setae which are gently curved, or not strongly hooked; F-9 with 4 rela-

130 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

tively weak bristlelike setae; F-9-11 with or without 1-3 short setae distad of

normal whorls.

MALE GENITALIA (Fig. 60). Exceedingly similar to navalis, differing

from it particularly in the following features. Basimere, Inner tergal surface

always with 1 strong submarginal seta; marginal setae relatively stronger or

more conspicuous. Subapical Lobe, Roda rather slender and straight, more

or less overlapped with rods b,c in distal portion; setae d-f stronger, longer,

more or less flattened, bladelike, 3,4 in number; hairlike setae associated with

setae d-f stronger, very distinct and apparently more numerous. Dislimere,

Slender, distally tapering to a blunt, gently recurved point, apical portion not

swollen distad of median curvature; dorsal subapical seta distad of ventral.

Phallosome. External process of lateral plate apparently without distinct api-

cal spiculose lobe (or knob) sternad. Pyroctiger. Cercal setae 3.

PUPA. As figured and described for vavalis (Fig. 58), differing from it

in the following characters. Metanotum, Seta 12-C double. Abdomen. Seta

9-VII usually 5,6 branched (4-6). Paddle, Relatively smaller and shorter,

about 0.48 mm in length.

LARVA (Fig. 60). Extremely similar to navalis, differing from it in the

following characters. Head, Seta 5-C always double. Antennal shaft entirely

dark brown; spicules strong, fewer, more or less restricted to lateral surface

below insertion of seta 1-A. Thovax. Seta 1-P always single; 7-P double; 1-M

longer, almost as long as 3-M. Abdomen. Seta 7-I double, rarely single.

Saddle dark brown, same color as antenna; seta 1-X weaker, double. Siphon.

Dark brown, same color as antenna and saddle; subventral tufts relatively

stronger and more pigmented.

TYPE-DATA. Holotype “*, Kuching, Savawak, Borneo, {| MALAYSIA],

1914, J. C. Moulton (BMNH).

DISTRIBUTION. Known only from Malaysia (Peninsular Malaysia and

Borneo) and Singapore. 21 specimens examined: 10°, 892, 3 L; 5 with associ-

ated lp skins.

MALAYSIA. Peninsular Malaysia - Selangor: Pacific Tin, K. Sel.; Kg.

Tanjong Rabok; Johore: Kota Tinggi; 7%, 69, 1 L, 3 lp. Malaysia - Sarawak:

Kuching; 3c.

SINGAPORE. 29, 2 L, 2 lp.

TAXONOMIC DISCUSSION. Culex coerulescens is extremely similar to —

navalis in all stages and evidently falls into a complex with the latter species.

It can be differentiated from “”davalis in the male by the details of the modified

tufts of the antenna, as described above; in the male genitalia by the details of

the setae of the subapical lobe and the shape of the distimere; in the pupa by

the reduction in size and length of the paddle and in the larva by the dark an-

tenna; the single seta 1-P; the double seta 7-P and the double seta 7-I. Both

species overlap in distribution and breeding site, but there is apparently no

indication of intergradation in their specific diagnostic features among the

material which I have examined.

BIONOMICS. As in navalis, coerulescens has also been found to utilize

Nepenthes ampullaria and other pitcher plant species as breeding sites. It

appears to be rare and less common than navalis,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 131

hewitti complex

48. CULEX (LOPHOCERA OMYIA) HEWITTI (EDWARDS)

(Figs. 61, 62)

Lophoceratomyia hewitti Edwards 1914a: 80 (“, ).

Culex (Lophoceratomyia) hewitti (Edwards), Edwards 1928: 277 (o*); Edwards

and Given 1928: 355 (L*); Brug and Edwards 1931: 261 (distribution);

Brug and Bonne-Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) hewitti (Edwards), Stone, Knight and Starcke 1959:

233 (catalog); Colless 1965: 303 (“*, 9, L).

FEMALE. Wing: 2.5mm. Forefemur: 1.3 mm. Proboscis: 1.7 mm.

Small or minute species; in general extremely similar to navalis and coerules-

cens, differing in smaller size and in the following features. Head, Decum-

bent scales on ocular line of vertex broader, extending to posterolateral areas;

narrow linear decumbent scales restricted to dorsal midline in center; erect

scales finer, usually entirely blackish, sometimes distally pale yellowish;

palpus more slender, segmentation not clearly marked. Cibarial Armature

(Fig. 61). Cibarial bar with 18-20 short teeth, 4 median teeth narrow, fine,

apically pointed; lateral teeth very coarse, widely spaced, dark at bases, pale

distally, with apices abruptly pointed, truncate or blunt.

MALE (Fig. 61). As in female except for smaller size. Palpus, Very

slender, distinctly shorter than proboscis by about the length of segment 9:

segments 4,5 markedly reduced in length. Pyroboscis, Very slender and thin.

Antenna, Flagellar whorls weakly to strongly plumose, normal long setae

rather few or sparse; modified tufts of scales or setae poorly developed; F-5

apparently without any conspicuous modified setae or scales, sometimes with

2,3 undifferentiated, shortened setae among normal long setae; F-6 witha

characteristic tuft of 2-4 dark, narrow, subapically swollen budlike scales

which are equally long and apically drawn out into fine points; F-7 with a

markedly reduced comb-shaped tuft of 3,4 short, subapically swollen setae

and a thick dark tuft of fused, subapically twisted setae; F-8 with relatively

small, gently hooked tuft of 5,6 fused setae; F-9 with or without 4 dark, weak

bristlelike setae; F-9-11 usually with 1 short seta distad of whorls.

MALE GENITALIA (Fig. 61). Small or minute. Segment IX, Tergal lobe

poorly developed, bearing 2,3 short fine setae; sternum with a transverse row

of 5 strong setae caudad. Basimere, Broad, conical, about 0.17 mm in

length, sometimes shorter; inner tergal surface without differentiated submar-

ginal setae; lateral tergal surface with numerous strong setae and bristles.

Subapical Lobe, Rodlike setae a-c largely overlapping, a,b stout, subequal,

c very slender, all with hooked apices; leaflet 8; replaced by a short, hairlike

seta, subequal to seta 1; setae d-f narrow, bladelike, 5,6 in number; leaflet

§9 more or less similar to setae in group d-f, but broader and longer. Disti-

mere, Rather thick, as long as basimere, with strongly modified recurved

apex; dorsal subapical surface irregular, with a few toothlike spicules; sub-

apical claw slender and relatively long, dorsal and ventral subapical setae

present, distinct, placed opposite each other. Phallosome, Poorly sclero-

tized; internal process short, simple, spinelike; external process distally

tapered in lateral aspect, with several minute denticles or spicules largely

restricted to its apex, apical lobe (or knob) not differentiated. Proctiger.

Essentially similar to other species in the Mammilifer Group; apical crown

132 Contrib. Amer. Ent.<Inst.,° vol. 13, no. 4, 1977

medium-sized; composed of several strong and weak spinelike spicules; cercal

setae 3-5.

PUPA (Fig. 61). Abdomen: 2.1mm. Paddle: 0.36 mm. Trumpet: 0.40

mm; index about 8. Similar to mavalis in the shape and length of trumpet and

detailed chaetotaxy, differing in darker pigmentation and in the following char-

acters. Cephalothorax, Seta 8-C usually triple (2-4); 9-C usually double or

triple (2-4). Abdomen, Seta 3-II, III single; 3-III strong, slightly longer than

segment following; 5-IV-VI remarkably strong and long, all single; 5-V, VI at

least 2 times as long as segment following; 9-VIII large, almost as long as

paddle length, usually 8 branched (7-9). Paddle. Markedly reduced in width

and length; setae 1,2-P minute, rather inconspicuous.

LARVA (Fig. 62). Head: 0.52 mm. Siphon: 1.0 mm; index 9-10. Saddle:

0.26 mm; siphon/saddle ratio 4. In general similar to navalis and coerulescens,

differing as follows. Head, Integument pale yellowish; seta 1-C long, slender,

spiniform and pale; 4-C weak, 3,4 branched, situated near dorsal midline; 5-C

weak, as long as 7-C, situated at same level as 6-C,mesad, usually with 6

pectinate branches (5-9); 6-C long, single; 7-C 7, 8 branched; 8-C long, single

(1-2). Antenna entirely pale, about 0.75 of head length; spicules numerous

and fine; seta 1-A with 9-11 short plumose branches; 2, 3-A apical or slightly

subapical. Thorax, Seta 1-P single and long; 2,3-P weak, subequal and single;

4-P double; 7, 8-P double; 8,9-M 5 branched; 7,9-T 5-7 branched. Abdomen,

Setae 6-I, II usually triple (3-5); 7-I double; 6-III- VI double; 1-III-VI weak,

double or triple; 1-VII single, strongly pectinate. Comb scales numerous,

minute, about 50. Saddle pale yellowish. Siphon, Slender, distally tapering,

same color as head capsule and saddle; pecten teeth 5-9; larger distal teeth

narrow, elongate, with 12-15 fine, undifferentiated denticles; subventral tufts

strong, 6, 7pairs, eachusually 5branched (3-6), forming a prominent double

row between 0. 295-0. 75 of total siphon length; 5, 6 proximal pairs long, subequal,

4,9 times as long as siphonal width at point of attachment; 1, 2 distal pairs

markedly or gradually reduced in length.

TYPE-DATA. Lectotype “* (one of 2 males in the type-series described

by Edwards 1914a: 80), Matang, Sarawak, N. Borneo [ MALAYSIA], July 1908,

J. Hewitt (BMNH; present selection).

DISTRIBUTION. Known only from Malaysia, Singapore and Sumatra,

Indonesia. 88 specimens examined: 460, 289, 14 L; 29 with associated im-

mature skins (14 p, 15 lp).

MALAYSIA. Peninsular Malaysia - Selangor: Pacific Tin, Butang; Kg.

Tanjong Rabok; Pahang: Pekan Rd.; Johore: Kota Tinggi; 330° 2192, 6L, 12 p,

9 lp. Malaysia - Sarawak: Matang (type-localitv); Mt. Dulit; 20".

SINGAPORE. Lornil Rd.; Pearce Reservoi Nee Soon Rifle Range; 2c,

02, 3 L, 2 p, 3 lp; unspecified locality; 40° (A. k. Barr collection).

INDONESIA. Sumatva; Dermajoe, Air Prioekan; 5c, 22, 5 L, 3 lp.

TAXONOMIC DISCUSSION. Culex hewitti apparently falls into a distinct

complex of the Brevipalpus Subgroup. It appears to resemble navalis and co-

evulescens more than other known pitcher plant species. It can be readily

recognized in the adults by the relatively small or rather minute size, in the

male by the peculiar modified scale tuft of antennal flagellomere 6 and the re-

duction of the modified tufts of antennal flagellomeres 7-9; in the genitalia by

the details of the setae of the subapical lobe, the modified recurved apex of the

distimere and the shape of the external process of the phallosome; in the pupa

by the elongate, funnel-shaped trumpet; the remarkably strong, single setae 5-

IV-VI of the abdomen and the reduction in size and length of the paddle and in

the larva by the branching and position of setae 4,5, 6 and 8-C; the branching

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 133

of setae 1, 3,4, 7 and 8-P and the features of the siphon.

The male antenna and genitalia of the type of hewitti originally designated

by Edwards were lost. I have, however, examined 2 other males in the origin-

al type-series of the author and here selected one of these as a lectotype.

BIONOMICS. Culex hewitti is apparently as common as vavalis and has

usually been found in association with the latter in the same species of pitcher

plant (Nepenthes ampullaria). In Singapore, the immatures have also been

found with specimens of curtipalpis in Nepenthes gracilis (Barr and Chellapah

1963).

jenseni complex

49. CULEX (LOPHOCERA OMYIA) JENSENI (DE MEIJERE)

(Figs. 63, 64)

Cyathomyia jenseni de Meijere 1910: 922 (“*, 2, P*, L*).

Culex (Lophoceratomyia) jenseni (de Meijere), Edwards 1928: 279 (o*);

Edwards and Given 1928: 356 (L); Brug and Edwards 1931: 261 (distribu-

tion); Brug and Bonne-Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) jenseni (de Meijere), Mattingly 1949: 226 (L); Stone,

Knight and Starcke 1959: 234 (catalog); Colless 1965: 303 (o"*, 2, L).

FEMALE. Wing: 3.5mm. Forefemur: 1.8mm. Proboscis: 2.1 mm.

In general exceedingly similar to the mavalis complex, differing slightly in

larger size and in the following. Head, Narrow linear decumbent scales of

vertex relatively dense, more numerous and entirely pale whitish; erect scales

coarser, pale golden or yellowish in center and on anterior margin, dark

brownish on posterolateral areas; lateral patch of broad scales whitish, very

distinct. Cibarial Armature (Fig. 63). Essentially similar to mammilifer,

minor and most members of the Mammilifer Subgroup; cibarial bar with 30 or

more narrow elongate teeth which are subequal in size and length, closely

spaced and distally tapering into sharp points. Thorax. Mesonotal scales

relatively dense, vestiture smooth. Pleuron with or without a few pale trans-

lucent scales on upper corner and posterior border of sip.

MALE (Fig. 63). In general as in female, with the following distinctive

features. Antenna. Spiculose prominence of pedicel moderately to strongly

developed; flagellar whorls densely long plumose; F-5 with a distinct tuft of

2,3 long, narrow golden acute scales as long as or Slightly longer than the

next 4 flagellomeres; modified tufts of F-6, 7 weakly developed; F-6 with 2

short, fine hairlike setae and 1 dark, weakly curled flattened seta; F-7 witha

lateral tuft of 5 dark, weakly curled setae and a small ventral tuft of 3 fused,

subapical, twisted or kinky setae. F-8 with a smoothly curved or typically J-

hooked tuft of 6,7 fused setae; F-9 with 3 long poorly differentiated bristlelike

setae; F-9-11 without any short setae distad of normal whorls.

MALE GENITALIA (Fig. 63). Very similar to navalis and coerulescens,

differing particularly in the following features. Segment IX, Tergal lobe with

3,4 longer hairlike setae; sternum with a transverse row of 6 strong setae

caudad. Basimere,. Relatively larger, about 0.22 mm in length; submarginal

setae absent. Subapical Lobe, Broad; rodlike setae @-c straight or smoothly

curved and largely overlapping; rod @ strong with abruptly pointed apex, b,c

more slender and longer, with hooked apices; seta 2 very strong and long,

placed close to the base of rod a; leaflet g] absent or represented by a short

weak, hairlike seta; setae d-fconsist of 3 short narrow blades and 1 hairlike

seta; leaflet go in form of a long stout blade. Distimere, More slender,

134 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

typically sickle-shaped with distal portion tapered to a slender, recurved apex;

ventral subapical seta stronger and more distinct than dorsal; subapical claw

short, apically broad and rounded. Phallosome, External process of lateral

plate tapered to a round, apical spiculose lobe, upper tergal surface without

any distinct teeth or denticles; internal process short, slender, about 0.5 of

external process in length.

PUPA (Fig. 63). Abdomen: 2.5mm. Paddle: 0.60 mm. Trumpet: 0.3-

0.4 mm; index 3-4. Rather similar to brevipalpus complex, with the follow-

ing diagnostic features. More or less uniformly yellowish with light brownish

tinge on cephalothorax and metanotum. Trumpet, Brownish, short, stout,

sausage-like, apex slightly projecting beyond margin of middorsal ridge in

flat preparation. Cephalothorax, Seta 1-C single; 5-C very strong and long,

single; 7-9-C single, subequal. Metanotum. Seta 10-12-C single; 12-C some-

times double. Abdomen, Seta 1-II weakly plumose, 5, 6 branched; 3-I-III

single; 5-IV-VI single, as long as or slightly shorter than segment following;

6-III-VI single; 9-VII weak, double or triple; 9-VIII 5-7 branched, subequal to

or slightly longer than 9-VII. Paddle. Broad and pale; outer margin distinct,

midrib lightly pigmented; setae 1,2-P indistinct or not developed.

LARVA (Fig. 64). Head: 0.78 mm. Siphon: 0.65 mm; index 3. Saddle:

0.38 mm; siphon/saddle ratio 1.7-1.8. Rather similar to brevipalpus complex

with the following diagnostic features. Head, Yellowish or pale cream-

colored; seta 1-C dark, long, stout, spiniform; 5-7-C dark, subequally long;

5, 6-C usually triple (2-5); 7-C 8 branched. Antenna straight, proximally

swollen and entirely pale, or same color as head capsule; spicules minute,

sparse and rather inconspicuous; seta 1-A short, weak, 3-6 branched, situated

at 0.75 of the length from base; 2, 3-A dark, strong and long. Mental plate

narrow, with 8,9 equally small lateral teeth on each side of a large median

tooth. Thorax. Setae 1,2-P single; 3-P double; 4-P 3,4 branched; 5, 6-P

single or double, sometimes triple; 7-P double or triple; 8-P double; 14-P

double; 5-M 2-7 branched. Abdomen. Setae 6-I, II dark, 3,4 branched; 7-I

double; 6-III-VI double; all other setae weak and short. Comb scales 24-30,

aggregating in 2,3 rows; apical fringe of posterior scales more or less pointed

or terminated into a strong median spine, latter sometimes not differentiated.

Saddle same color as head capsule; spiculation weakly developed or absent;

seta 2-X single or double; ventral brush dark, with 6 pairs of triple or more

branched setae. Siphon, Short, thick, distally slightly tapered, sometimes

more or less cylindrical; pigmentation pale, concolorous with head capsule

and saddle; pecten teeth 4-8, widely spaced, with fine barb of several minute

denticles; siphonal tufts 8-10 pairs, first 5,6 subventral pairs very dark,

strong, 4-6 branched each, 3,4 times as long as siphonal width at point of

attachment; 2-4 weaker pairs strongly reduced, 3,4 branched; more proximal

1-3 pairs placed dorsolaterally distad of pecten from 0.4-0.7 of siphon, most

distal pair subapical, in line with the strong subventral pairs.

TYPE-DATA. Lectotype “ (No. 32) with attached genitalia mount, Tjibodas,

Bogor (Buitenzorg), Java | INDONESIA], in Nepenthes pitcher, 1909, Dr. Jen-

sen (Zoological Museum, Amsterdam University; selection of Bram 1967b:

328).

DISTRIBUTION. Known only from Malaysia (Sabah), Singapore and Indo-

nesia (Sumatra and Java). 72 specimens examined: 150°, 39, 54 L, 8 with

associated lp skins.

MALAYSIA. Sabah: Mt. Kinabalu; 54 L.

SINGAPORE. 1¢.

INDONESIA. Sumatra: Batang, Palupuk; 10°; Java: Bogor (Buitenzorg);

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 135

Tjibodas (Botanical Garden); Mt. Salak; 130°, 39, 8 lp.

TAXONOMIC DISCUSSION. Culex jenseni is an interesting pitcher plant

form which exhibits a bizarre combination of characters of the Mammilifer

Subgroup in the female cibarial armature, the ”avalis complex in the male

antenna and genitalia and the brevipalpus complex in the relatively short male

palpus, the chaetotaxy and general features of the pupa and larva. It can be

readily recognized and separated from other pitcher plant species in all stages

by the characters as given in the above diagnosis and as illustrated. Because

of several distinctive features, jenseni apparently falls into a distinct complex

of the Brevipalpus Subgroup.

Several topotypic males of jenseni from Java and a few other males from

Sumatra and Singapore which I have thoroughly re-examined in the collections

of the Instituut voor Tropische Hygiene, Amsterdam and the BMNH agree well

with the previous descriptions by Edwards (1928: 279) and Colless (1965: 303).

I have not seen the larvae from Singapore described by Edwards and Given

(1928: 356), but judging from the description of these authors, they apparently

conform to the topotypic specimens described above. The numerous whole

larvae recently collected from Sabah, Malaysia, which I have tentatively

identified with this species are, however, variable and appear to be locally

differentiated into 2 forms. One of these almost corresponds with the typical

form in the detailed chaetotaxy but differs from it in having posterior comb

scales terminating into a stout median apical spine, setae 5, 6-P usually double

or triple and in the presence of 2 or 3 additional pairs of weak dorsolateral

tufts from basal 0.40-0. 75 of the siphon. The other differs from the typical

form rather constantly in having setae 5 and 6-C and setae 6-III-V] single. It

is probable that 2 or more forms are involved and that at least one of these

may be recognized as distinct. However, in the absence of correlated male

specimens, it seems best to consider this larval material only as forms of

jenseni or the jenseni complex for the present.

BIONOMICS. Culex jenseni is apparently restricted to high elevations in

mountainous areas where it is fairly common and has been found on several

occasions. At the type-locality on Tjobidas Mountain and Bogor, Java, the

larvae were collected from unknown pitcher plant species at an elevation

ranging from 1,500-1,600 m. At Mt. Kinabalu, Sabah, the larvae were taken

from Nepenthes villosa at an elevation of about 2,500 m (8,200 ft.). In Singa-

pore, the larvae were collected from pitcher plants without further particulars

(Edwards and Given 1928: 356).

brevipalpus complex

50. CULEX (LOPHOCERAOMYIA) BREVIPALPUS (THEOBALD)

(Figs. 65, 66)

Lophoceratomyia brevipalpus Theobald 1905b: 96 (o"); Theobald 1907: 477 (<);

Leicester 1908: 129 (); Stone 1957: 172 (nomenclature).

Culex (Lophoceratomyia) eminentia (Leicester), in part, Edwards 1928: 278

(o*); Edwards and Given 1928: 355 (L*).

Culex (Lophoceraomyia) brevipalpus (Theobald), Colless 1965: 301 (o*, 2, L).

FEMALE. Wing: 2.8mm. Forefemur: 1.5mm. Proboscis: 1.8 mm.

Medium or small size, in general conforming to the Brevipalpus Subgroup with

the following distinctive features. Head, Decumbent scales on dorsum of ver-

136 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

tex narrow, linear and entirely dark or partially pale whitish along upper eye

border, dark brownish in the center; erect scales coarse, usually predomin-

antly pale bronzy or golden, sometimes entirely dark brown; lateral patch of

broad scales grayish to blackish, not distinct. Cibarial Armature (Fig. 65).

Cibarial bar narrow, with 15-25 teeth; 2,3 median teeth narrow, elongate,

distally tapering into sharp points, lateral teeth coarser, shorter, more wide-

ly spaced with blunt, abruptly pointed or truncate apices. Thovax, Mesonotal

integument deep chestnut brown; scales narrow, fine and blackish. Apn witha

few to several dark broad scales which sometimes form a distinct patch.

Pleural integument paler than or as dark as mesonotum; pp/ without any scales;

upper corner and posterior border of stp usually with some scattered pale

scales; lower ep bristle absent. Legs and Wing. As described for the Mam-

milifer Group, apparently without any distinctive features. Abdomen, Terga

and sterna dark scaled; terga II-IV usually slightly pale beige on lateral mar-

gin.

MALE (Fig. 65). In general similar to female with the following distinc-

tive characters. Palpus, About 0.5 of the length of proboscis, segments 3, 4

without any apical setae or bristles, segment 5 with 1 short apical spinelike

seta. Proboscis, Rather slender and thin in basal 0.5, apical 0.5 gradually

thickened. Antenna. Spiculose prominence of pedicel well developed, distinct;

flagellar whorls moderately to strongly plumose; modified tufts of scales and

setae present on F-5 to F-9; F-5 with a distinct tuft of 5 narrow, acute scales;

dorsalmost scales usually dark, as long as next 3 flagellomeres, the remain-

ing scales yellowish, golden or brownish, gradually increased in length, some-

times all scales pale yellowish; F-6 with a crumpled tuft of 5 dark weakly

curled setae; F-7 with a lateral comb-shaped tuft of 10 distally kinked setae

and a ventral tuft of 4 subapically twisted fused setae; F-8 with a slender and

weak J-hooked tuft of 5, 6 fused setae; F-9 with 2-4 long and strong bristlelike

setae.

MALE GENITALIA (Fig. 65). Segment IX. Tergal lobe poorly developed,

bearing 3,4 tiny setae; sternum with 4-10 strong setae in a transverse row

caudad. Basimere, Rather broad, more or less rectangular or somewhat

square in shape; its length about 0.22 mm, maximal width about 0.75 of the

length; inner and lateral tergal surface practically bare or with only a small

number of short, minute setae; submarginal setae absent; strong setae and

bristles restricted to outer tergal surface. Subapical Lobe, Details as fig-

ured; very broad; with several strong, flattened accessory bladelike setae

in group d-f densely packed at bases of usual rodlike setae @~c and leaflet g9;

rod a stout, short, not clearly differentiated from other bladelike setae; rods —

b,c distinct, stout, subequally long and apically hooked; leaflet g; absent;

seta 2 very long, flattened, rodlike, as long as or longer than rods 0,Cc; setae

in group d-f all strong, bladelike; leaflet go strong, long, markedly flattened

and subapically bent, somewhat resembling a boot in shape. Distimere,

Sickle-shaped, with characteristic modified truncate apex; ventral subapical

seta strong, spiniform; dorsal subapical seta weak, short, setiform or some-

times spiniform; subapical claw short, flattened and apically blunt. Phallo-

some, External process slender, with a distinct apical spiculose lobe (or

knob) sternad and variable number of weak denticles at base of lobe, upper

tergal surface with or without a few weak denticles at some distance below

base of apical lobe; internal process shorter than external process. Pvroctiger,

As described for the Mammilifer Group; cercal setae 2.

PUPA (Fig. 65). Abdomen: 2.5mm. Paddle: 0.65 mm. Trumpet: 0.3

mm; index 3,4. Cephalothorax and abdomen brownish yellow, sometimes

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 137

lighter or yellowish white. Trumpet, Dark brown, short, stout, sausage-like.

Complete chaetotaxy as figured, rather similar to jenseni, differing as follows.

Cephalothorax, Seta 9-C single, strong and long, about 3 times as long as

trumpet; 8-C strong, 3 times as long as 9-C. Metanotum, Setae 10-12-C

always single. Abdomen, Seta 1-II weaker, 3,4 branched; 6-I, II very strong

and long; 9-IV, V strong, 1.5 times as long as segment following; 9-VII, VIII

single and strong; 9-VIII almost as long as paddle. Paddle. Midrib strong and

dark pigmented.

LARVA (Fig. 66). Head: 0.67 mm. Siphon: 0.90 mm; index 4.7. Saddle:

0.32 mm; siphon/saddle ratio 3. In general very similar to jenseni, Head,

Seta 1-C pale, distally filamentous; 4-C close together near dorsal midline;

o-7-C relatively short, flattened, strongly plumose, situated more cephalad;

o-C 2-4 branched; 6-C single; 7-C 4-6 branched. Antennal shaft practically

bare or with a few to several minute, inconspicuous spicules; setae 2, 3-A

weak short and pale. Mental plate very dark and broad, with 2,3 large and 4,5

small lateral teeth on each side of median tooth. Mouth brush strong, rather

coarse and short. Thorax, Setae 3,5,6-P single; 4-P double or triple; 7,8-P

usually double (1-3); 14-P single or double; 5-M 4,5 branched. Abdomen.

Seta 6-I 4-6 branched; 6-II triple; 7-I usually triple (3-5); 7-II strong, subequal

to 7-I, usually triple (2-5); 6-III-VI double or triple; setae 1, 13-III-VI and 1,4,

7, 10,12 and 13-VII single and strongly flattened. Comb scales 20-30, all with

apically rounded fringe of evenly fine spicules. Setae 1, 2,4, 5-VIII single.

Saddle seta 2-X with 1 short and 1 long branch. Siphon. Pecten teeth 2-4;

subventral tufts at least 7 pairs, sometimes 8,9, all in a prominent double

row on lateral ventral surface; 6,7 strong proximal pairs 4-6 branched, 3, 4

times as long as siphonal width at point of attachment.

TYPE-DATA. Holotype “with slide of genitalia, SINGAPORE, 1902, Hun-

garian National Museum, Budapest (HNM).

DISTRIBUTION. Known only from Malaysia and Singapore. 62 specimens

examined: 260%, 20°, 16 L, 21 with associated lp skins.

MALAYSIA. Peninsular Malaysia - Johore: Kota Tinggi; Pahang: Pekan

Rd.; Trengganu: Kemaman; 180", 209, 12 L, 18 lp. Malaysia - Sabah: Jessel-

ton: 1c,

SINGAPORE. Kg. Chantek Bahru; Swiss Club; other unspecified localities;

10, @ ©. 3 ip.

TAXONOMIC DISCUSSION. Culex brevipalpus and the next 3 species,

lucaris, eminentia and acutipalus of the brevipalpus complex, are very similar,

forming a compact monophyletic group which exhibits its strongest affinity with

the jenseni complex in pupal and larval morphology. As noted by Colless

(1965), morphological differences among the members of this complex are

exceedingly slight, but each is apparently distinct and can be differentiated

from the others by the subtle details of the modified tufts of the male antenna,

the setae of the subapical lobe of the genitalia, and the usual features of larval

and pupal chaetotaxy as indicated in the keys, figures and descriptions. The

males of brevipalpus differ from eminentia and acutipalus in the modified tufts

oi antennal flagellomeres 5, 8 and 9, but are extremely similar to those of

lucaris. It can be separated from the latter by the stronger and longer seta h

and the absence of numerous short hairlike setae at the base of seta go of the

subapical lobe of the genitalia. The pupae are indistinguishable from acutipal-

us, but can be separated from lucaris and eminentia by the stronger and longer

setae 0 and 8-C, 5-IV-VI and 9-VII-VII. The larvae differ from eminentia in

having seta 4-C more closely spaced near the dorsal midline, stronger setae

1-4-P and by the usually double setae 7 and 8-P; from lucaris and acutipalus in

138 Contrib. Amer. Ent. Inst., vol. 13,:.no. 4, 1977

having stronger seta 7-II which is of the same magnitude as setae 6-I-II or

7-1,

BIONOMICS. The larvae of brevipalpus have been collected either from

Nepenthes ampullaria or gracilis in open sunlight or under partial shade of

secondary tropical forest at an elevation ranging from sea level to about 30

m. In Singapore, brevipalpus has been reported to be more common than

eminentia and acutipalus (Colless 1965: 301). Of 6 collections recently made

in Peninsular Malaysia, the larvae and reared adults were once found in as-

sociation with the specimens of eminentia, The strong filaments of the mouth

brush and the heavy teeth of the maxilla, mandible and mental plate of the lar-

vae suggest that it probably feeds by chewing on dead insects trapped in the

pitcher plants.

01. CULEX (LOPHOCERA OMYIA) LUCARIS COLLESS

(Fig. 67)

Culex (Lophoceraomyia) lucaris Colless 1965: 299 (o"*); Bram 1967a: 79 (C*).

FEMALE. Exceedingly similar to brevipalpus from which it can not be

distinguished with certainty, differing slightly in the following: Thorax, Ppl

usually with a few pale scales on anterior surface; upper corner and posterior

border of stif usually with more numerous pale translucent scales aggregated

into distinct patches. :

MALE (Fig. 67). As described and figured for brevipalpus, differing from

it particularly in the following features. Palpus, Relatively longer, its length

usually 0.60-0.75 of proboscis. Antenna, F-8 with a smoothly curved J-

hooked tuft of fused setae; F-9 with a stronger tuft of 4 bristlelike setae.

MALE GENITALIA (Fig. 67). Exceedingly similar to brevipalpus, differ-

ing from it in the following features. Subapical Lobe. Seta h shorter and

weaker, bristlelike, as long as or shorter than rodlike setae b,c; base of seta

g9 prominent, with numerous conspicuous hairlike setae; setae in group d-f

shorter and narrower.

PUPA. As figured and described for brevipalpus (Fig. 65), differing from

it in having most setae considerably shorter and weaker and in the following.

Cephalothorax, Seta 5-C weaker and shorter, about 1.25 of trumpet length.

Abdomen. Setae 5-IV-VI weaker, shorter than or as long as segment follow-

ing; setae 6-III-VI weaker, usually double or triple, sometimes single; 9-VII

much weaker, usually double or triple, sometimes single or 4 branched; 9-VIII

shorter, about 0.5 of paddle length. Paddle. Midrib weak and pale.

LARVA. As figured and described for brevipalpus, differing from it in the

following features. Head, Seta 4-C more widely spaced. Thovax, Seta 4-P

usually double (2-3); 7-P usually single (1-2); 8-P single or double. Abdomen.

Seta 7-II very weak and pale, similar to 7-III-VI, 5, 6 branched; 6-III-VI

double or triple. Siphon, Subventral tufts usually 7 pairs, sometimes 8 or 9.

TYPE-DATA. Holotype “ with slide of antenna (CH 70) and genitalia (CT

663), Ulu Pandan, SINGAPORE, 7 July 1957, D. H. Colless (ANIC).

DISTRIBUTION. Known only from Thailand, Peninsular Malaysia and

Singapore. 44 specimens examined: 230, 162, 5 L; 21 with associated imma-

ture skins (3 p, 18 lp).

THAILAND. Ranong: Hard Sam Pan; Ban Phong Rang; 30, 2 L, 1p, 1 lp.

MALAYSIA. Peninsular Malaysia -Selangor: Ulu Klang; 29, 1 p.

SINGAPORE. Unspecified locality; Ulu Pandan; Pulau Blakang Mati; 20¢,

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 139

149 8eice kip;

TAXONOMIC DISCUSSION. As discussed under brevipalpus, morphological

differences between lucaris and the other 3 members of the brevipalpus complex

are only slight and caution should be taken in separating them. The most dis-

tinctive features of the males of lucaris are: the relatively long palpus which

is Slightly more than 0.9 of the proboscis length; the shape of the modified tuft

of antennal flagellomere 8 which is smoothly curved or of a typical J-shape

rather than sigmoid and the presence of numerous hairlike setae at the base

of the boot-shaped seta 85 of the subapical lobe of the genitalia. In the imma-

tures, the pupa is indistinguishable from eminentia but can be differentiated

from brevipalpus and acutipalus by the weaker and shorter setae of the cephalo-

thorax and abdomen and the usually double or triple setae 6-III- VI and 9-VII.

The larva is separated from brevipalpus and eminentia by the considerably

weaker and shorter seta 7-II, from acutipalus by the more widely spaced seta

4-C and by a few other usual features as described under the latter species.

BIONOMICS. As in brevipalpus, lucaris usually occurs in the open secon-

dary tropical forest at a low elevation ranging from sea level to about 60 m.

In Singapore, the larvae have been collected from Nepenthes gracilis, rafflezi-

ana and other unidentified pitcher plant species along the coast as well as

towards the inland hills. Larvae from a single collection in Peninsular Malay-

sia were found in Nepenthes gracilis in association with specimens of eminentia.

In Thailand, all collections were made from pitcher plant species without fur-

ther particulars. Most of the adults came from rearing larvae or pupae. In

Singapore, Colless (1965: 299) noted that the adults were taken while resting

by day in an isolated patch of rather open forest.

02. CULEX (LOPHOCERAOMYIA) EMINENTIA (LEICESTER)

(Fig. 67)

Lophoceratomyia eminentia Leicester 1908: 131 (<*).

Culex (Lophoceratomyia) eminentia (Leicester), in part, Edwards 1917: 227

(, key); Edwards 1928: 278 (o‘*); Edwards and Given 1928: 355 (L*);

Edwards 1932: 198 (taxonomy); Brug and Bonne-Wepster 1947: 186

(distribution).

Culex (Lophoceraomyia) eminentia (Leicester), Stone, Knight and Starcke

1959: 233 (catalog, in part); Colless 1965: 298 (<**, @, L).

FEMALE. As described for brevipalpus and lucaris from which it is in-

distinguishable.

MALE (Fig. 67). In general as in female, differing from brevipalpus and

lucaris particularly in the following features. Palpus. Longer, about 0.75-

0.90 of proboscis length. Antenna. Modified tuft of F-5 larger, very conspi-

cuous, composed of 6-10 bright yellow or golden, broad lanceolate scales

which are as long as the combined length of next 4 flagellomeres; F-6 with a

stronger tuft of 9-10 curled setae; F-8 with a sigmoid-hooked tuft of 6, 7

fused setae; modified tuft of F-9 large, prominent, consisting of 10-12 long,

strong, flattened setae which are more or less fused proximally.

MALE GENITALIA (Fig. 67). Essentially similar to bvevipalpus and

lucaris, differing particularly in the following features. Subapical Lobe, Seta

h absent or replaced by a weak short, inconspicuous seta; base of boot-shaped

seta £5 in distal portion without distinct long, hairlike setae; bladelike setae

d-fand other accessory flattened setae at bases of rods a-c apparently fewer

140 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

and stronger.

PUPA. Extremely similar to /ucayvis in having most setae relatively weak

and short and in diagnostic chaetotaxy as described under the latter species.

LARVA. As described and figured for brevipalpus, differing from it and

from other members of the brevipalpus complex in the following combinations.

Head, Seta 4-C more widely spaced. Thorax, Setae 1-8-P relatively weak

and short, 7,8-P double or triple. Abdomen. Seta 7-II double, as strong as

or subequal to 6-I,II and 7-I. Siphon. Subventral tufts 7, 8 pairs.

TYPE-DATA. Lectotype “*, Kuala Lumpur, (Selangor), Malaya | MALAY-

SIA], Jungle patch, 5th mile, Gombak Rd., 5 December 1903, G. F. Leicester

(BMNH; selection of Colless 1965: 298).

DISTRIBUTION. Known only from Malaysia and Singapore. 31 specimens

examined: 280, 22, 1 L; 2 with associated immature skins (1 p, 1 lp).

MALAYSIA. Peninsular Malaysia - Selangor: Kuala Lumpur (type-

locality), Ulu Klang; Trengganu: Kemaman; 5c, 19. Malaysia - Sarawak:

Kuching; Sabah: Kota Kinabalu; 4c.

SINGAPORE. Pulau Blakang Mati; 10, 12, 1L, 1p, 1 lp; unspecified

locality; 180° (A. R. Barr collection).

TAXONOMIC DISCUSSION. The males of eminentia which I have examined

all agree well with the description by Colless (1965: 298). The most distinc-

tive features are the relatively long palpus which is the longest among the

members of the brevipalpus complex; the large yellowish or golden tuft of

flagellomere 5 and the prominent tuft of 10-12 dark, long, flattened setae of

flagellomere 9 of the antenna and the absence of seta 2 (basal seta of Colless)

on the subapical lobe of the basimere of the genitalia. The pupa is indistin-

cuishable from lucaris, but is distinct from brevipalpus and acutipalus in

having most setae of cephalothorax and abdomen relatively shorter and weaker.

The larva closely resembles brevipalpus more than lucaris and acutipalus, It

can be separated from Drevipalpus by the more widely spaced seta 4-C and the

relatively weaker and short setae 1-8-P.

BIONOMICS. Culex eminentia is apparently less common than the other 3

members of the bDrevipalpus complex. As in brevipalpus and lucaris, its oc-

currence is restricted to a lowland, open, scrub or secondary forest at ele-

vations from sea level to about 60 m. In Peninsular Malaysia and Singapore,

the larvae were collected 3 times from Nepenthes ampullaria and once from

gracilis, In Singapore, Barr and Chellapah (1963) reported that the larvae

were most frequently found in N. gracilis,

58. CULEX (LOPHOCERAOMYIA) ACUTIPALUS COLLESS

(Fig. 68)

Culex (Lophoceraomyia) acutipalus Colless 1965: 299 (“*, 9, L).

FEMALE. As described by Colless (1965: 299-300), differing slightly

from brevipalpus, lucavis and eminentia in the following features. Head.

Narrow decumbent scales of vertex entirely blackish; lateral patch of broad

scales at side of eye blackish; erect scales darker, dull golden brown to dark

brownish. Thorax. Upper corner and posterior border of stf with some

scattered pale scales not aggregating into a definite patch, sometimes scales

entirely absent.

MALE (Fig. 68). Differing from other 3 members of the brevipalpus com-

plex as described for female and in the following sexual characters. Palpus.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 141

Relatively shorter, about 0.5 or less of proboscis length. Antenna. Modified

tufts of F-0-9 markedly reduced and less conspicuous; F-5 with 4, 5 narrow

golden or yellowish brown lanceolate scales; F-8 with a slender sigmoid

hooked tuft of 4, 5 fused setae; F-9 with 3,4 relatively weak bristlelike setae

which are not strongly differentiated from long normal setae.

MALE GENITALIA (Fig. 68). As figured, differing from brevipalpus,

lucavis and eminentia particularly in the following. Basimere, Narrower

or not strongly compressed; inner surface laterad of tergomesal margin with

more numerous short setae. Subapical Lobe. Accessory bladelike setae in

group d-f associated with seta a-c more numerous and densely packed; seta /

(basal seta of Colless) present, relatively weak and short, its length not as

long as or longer than rodlike setae a-c; base of boot-shaped seta 89 apparently

without any long, fine, hairlike setae.

PUPA. As described and figured for brevipalpus from which it is insepar-

able; differing from lucaris and eminentia particularly in having most setae of

cephalothorax and abdomen stronger and longer.

LARVA. As described and figured for brevipalpus, differing from it and

from lucaris and eminentia by the following combination of characters. Head.

Seta 4-C situated closer near dorsal midline; 1-C more slender. Thorax.

Seta 8-P usually triple (2-3); 14-P double or triple, rarely single. Abdomen.

Seta 7-II weaker and shorter than 6-I, II or 7-I, 5,6 branched, its length of the

same magnitude as 7-III-VI. Siphon. Subventral tufts 6,7 pairs.

TYPE-DATA. Holotype “ with associated pupal and larval skins (58/25/2)

and slides of antenna (CH 192) and genitalia (CT 666), Swiss Club, Kg.

Chantek Bahru, SINGAPORE, 28 November 1958, D. H. Colless (ANIC).

DISTRIBUTION. Known only from Peninsular Malaysia and Singapore.

27 specimens examined: 160°, 102, 1 L; 10 with associated immature skins

(2: p; <6: Ip).

MALAYSIA. Peninsulay Malaysia - Selangor: Pacific Tin, K. Sel;

Malacca: Malacca; 2c, 52, 2 lp.

SINGAPORE. Kg. Chantek Bahru: Swiss Club (type-locality); Ulu Pandan;

Pulau Blaklang; 110°, 59, 1 L, 4p, 4 lp; unspecified locality; 3c (A. R. Barr

collection).

TAXONOMIC DISCUSSION. Of all stages of acutipalus, the male is the

most distinct. It can be readily separated from the other 3 members of the

brevipalpus complex by the reduction of the antennal modified tuft and the

more numerous accessory setae (d-/) of the subapical lobe of the genitalia.

The pupa exhibits no clear-cut difference from that of brevipalpus, but can be

separated from lucaris and eminentia by the remarkably stronger setae 5-C

and the stronger setae 6-I, II, 5-IV-VI, 6-III-VI and 9-VII-VIII. The larva is

most similar to lucavis from which it can be separated by the more closely

spaced seta 4-C and by the presence of 6 or 7 pairs of siphonal tufts.

BIONOMICS. As in other members of the brevipalpus complex, acutipalus

has also been reported from the open scrub or secondary forest at elevations

from sea level to about 30 m. In Singapore, Colless (1965: 300) noted that

adults are quite commonly found resting on vegetation in the vicinity of breed-

ing places, but were not taken with mammalian baits. Little is known about

which species of the pitcher plants are most frequently utilized as breeding

sites. All of the 3 collections recently made in Singapore came from Nepenthes

ampullaria,

142 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

curtipalpis complex

54. CULEX (LOPHOCERAOMYIA) CURTIPA LPIS (EDWARDS)

(Figs. 69, 70)

Lophoceratomyia curtipalpis Edwards 1914b: 127 (¢).

Culex (Lophoceratomyia) curtipalpis (Edwards), Edwards 1921: 78 | as synonym

of C. (Lophoceratomyia) jenseni (de Meijere)]; Edwards 1928: 279 (o"*;

revalidated); Edwards and Given 1928: 357 (P, L*); Brug and Edwards

1931: 261 (distribution); Brug and Bonne-Wepster 1947: 186 (distribution).

Culex (Lophoceraomyia) curtipalpis (Edwards), Stone, Knight and Starcke

1959: 232 (catalog); Colless 1965: 301 (o*, 9, L): Bram 1967a: 76 (o*,

GL, in part).

FEMALE. Wing: 3.2 mm. Forefemur: 1.8mm. Proboscis: 2.0 mm.

Medium-sized, dark brown to black; in general conforming to the subgroup

characters, distinctive in the following. Head. Narrow decumbent scales of

vertex very fine, largely yellowish brown or black in center, pale whitish

along upper eye border, forming a distinct ocular line; lateral patch of broad

scales whitish, very distinct; erect scales usually predominantly dark brown

to black except for a few pale golden ones on lateral area; sometimes entirely

golden or yellowish brown. Cibarial Armature (Fig. 69). Cibarial bar moder-

ately broad, with a concave row of 24-30 coarse, short teeth which are rather

widely spaced, a few median teeth narrow, terminated into fine apices, lateral

teeth larger, flattened, abruptly pointed or truncate apically. Thorax, Meso-

notal integument deep chestnut brown to almost black. Apn with a small but

distinct patch of some broad pale or dark scales; pp/ with or without some

broad pale scales; upper corner and posterior border of sff with a conspicuous

vertical patch of loosely packed broad pale scales; lower mep bristle absent.

Legs and Wing. Essentially similar to the rest of the Mammuilifer Group.

Abdomen. Terga entirely black scaled except for lateral margin which is

slightly paler; sterna pale yellowish scaled.

MALE (Fig. 69). In general as in female with the following distinctive

features. Palpus. Short, about 0.5 of proboscis length. Pyvoboscis. Slender

and uniformly thin in basal 0.5, gradually thickened in apical 0.5. Antenna.

Pedicel with a blunt spiculose prominence on inner dorsal surface; flagellar

whorls moderately plumose; modified tufts of F-5-9 present, poorly developed

or strongly reduced in size, most of which are apparently restricted to ventral

and mesal surfaces; F-5 with a small tuft of 5,6 modified setae, 3,4 setae on

lateral dorsal surface short, slightly longer than the length of one flagellomere,

followed ventrally by 2 longer, flattened, scalelike setae which are about as

long as the combined length of next 4 flagellomeres; F-6 with a small tuft of 4

_ weakly curled setae; F-7 with a lateral comb-shaped tuft of 4,5 slender, sub-

apically hooked setae and a mesal tuft of 2-4 straight acute setae; F-8 with a

weakly curved tuft of 3-5 stout, fused setae; F-9 with 2-5 weak bristlelike

setae.

MALE GENITALIA (Fig. 69). As figured. Segment IX. Tergal lobe poor-

ly developed, bearing 2,3 short, weak setae; sternum with a transverse row

of 5,6 strong setae towards caudal margin. Basimere. Stout, broad oval-

shaped, about 0.25 mm in length; inner tergal surface lightly convex, with

several subequally strong setae on tergomesal margin and outer area laterad

of subapical lobe. Subapical Lobe. Prominent, elongate. projecting mesad,;

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 143

rodlike setae a-c rather short, stout, subequally long; seta a (most ventral)

strongest, distally curved upwards; b-c slender and slightly curved downwards:

setae in group d-fconsist of 3 strong, sharply bent blades and 2 smoothly

curved blades; seta gy (or leaflet) not developed or replaced by one of the setae

in group d-f; leaflet g; very broad, heart-shaped, distad of setae d-f; seta h

moderately long, closely adjacent to leaflet g,; numerous fine hairlike setae

present basad and mesad of leaflet 81. Distimere, Rather thick with modified,

expanded truncate apex; subapical claw short, small and apically blunt, situ-

ated close to the recurved point; dorsal and ventral subapical seta tiny, oppo-

site each other. Phallosome. Strongly sclerotized; internal process shorter

than external process; latter broad, with distinct spiculate apical lobe (or

knob) and 1-3 rows of 15 or more strong denticles on upper tergal surface. 7

Proctiger, Apical crown well developed, medium-sized, composed of several

strong spinelike spicules; cercal setae 3-5.

PUPA (Fig. 69). Abdomen: 2.9mm. Paddle: 0.5mm. Trumpet: 0.28

mm; index 2. Cephalothorax and abdomen usually dark brown, sometimes pale

cream-colored; complete chaetotaxy as figured, most setae weakly developed,

distinctive in the following. Trumpet. Dark brown, very short, bell-shaped,

apex not reaching beyond posterior margin of mid-dorsal ridge in flat prepara-

tion. Cephalothorax, Setae 1-9-C weakly developed; 1-C single; 2, 3-C triple;

4-C double; 5-C double; 6-C double; 7-9-C single. Metanotum. Seta 10-C usu-

ally triple (2-3); 11-C single; 12-C single or double. Abdomen, Seta 2-I 3,4

branched; 3-I 1-3 branched, weak, as long as 2-I, situated close to 4, 5-I; seg-

ment II with or without a spiniform seta (10-II?) on ventral surface; 1-II-VII

very weak, double or triple; 3-II usually triple (2-3); 3-III dark, strong, single;

6-III- VI weak, short, usually double; 5-IV dark, strong, as long as segment

following, usually single (1-2); 5-V, VI single, short, about 0.3 of segment

following; 9-III-VI minute, single, somewhat spiniform; 9-VII dark, strong,

double or triple; 9- VIII dark, stronger and longer than 9-VII, 4, 5 branched.

Paddle, Rather short and largely pale except at extreme base; apex lightly to

strongly emarginate; midrib poorly developed; seta 1-P present; 2-P absent.

LARVA (Fig. 70). Head: 0.60 mm. Siphon: 0.24 mm; index 1.5. Sad-

dle: 0.30 mm; siphon/saddle ratio 0.8. Complete chaetotaxy as figured;

pigmentation pale whitish with variable amount of greenish tinge; strikingly

different from the larvae of other pitcher plant species in having siphon short-

er than saddle and in the following. Head, Length greater than maximal width,

more or less resembling an Aedes; labrum narrow; seta 1-C short, stout,

apically blunt or abruptly pointed, situated on a distinct tubercle; 4-C usually

single, 2 times as long as distance between bases of the pair; 5, 6-C weak,

cephalad, usually triple, each (2-4); 7-C subequal to 5,6-C, double or triple ‘

8-9-C minute, cephalad; 10,11-C weakly developed; 13-C strong and long, usu-

ally double or triple (1-3). Antenna pale, very slender, uniform in width,

about 0.5 of head length; spicules absent; seta 1-A weak, 3,4 branched, situ-

ated slightly beyond the midpoint of antennal shaft; 2-6-A very weak, all

placed apically. Mental plate with 6,7 lateral teeth on each side of a small

median tooth. Mouth brush relatively short and rather coarse. Thorax,

Spicules absent; seta 4-P usually single, sometimes double; 7-P single or

double; 8-P extremely minute, rather inconspicuous, single; 14-P single; 5-M

single; 8, 9-M 3,4 branched; 7,9-T 3,4 branched. Abdomen. Not spiculate,

_ all setae except setae 6-I-VI, 7-I, II and 13-VII extremely weak and inconspicu-

ous; 6-I-III usually triple (2-3); 6-IV-VI double; 7-I, II both as strong as seta

6, double; 13-VII double, remarkably strong, of the same magnitude as setae

6 or 7. Comb scales 4-10, ina single row, all broad, subequal, with rounded

144 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

apical fringe of evenly fine spicules; setae 1,2 and 4-VIII minute, single or

double; 3 and 5-VIII dark, strong, flattened; 3-VIII double or triple; 5-VIII

single. Saddle broad; seta 2-X dark, with 5,6 strong branches; 4-X (ventral

brush) with 6 pairs of setae; anal gills fusiform, as long as saddle. Siphon.

Short, stubby and small, pecten teeth short, broad, 2-4, all simple; subven-

tral tuft 3 pairs; first 2 proximal pairs strong, subequal, usually 4, 5 branched

(3-4), as long as or slightly longer than siphonal width at point of attachment;

most distal pair weaker and shorter, 5 branched; dorsal valve of spiracular

apparatus broad with a strong hooked seta (seta 9?); median caudal filament

absent.

TYPE-DATA. Holotype “* (marked as type by Edwards) with slide of

genitalia, Kuching, Sarawak, | MALAYSIA], 6 March 1914, J. C. Moulton

(BMNH).

DISTRIBUTION. Thailand, Malaysia and Singapore; also reported from

Sumatra and Kalimantan (Borneo), Indonesia. 185 specimens examined:

113°, 649, 8 L; 39 with associated immature skins (14 p, 25 lp).

THAILAND. Ranong; Hard Sam Pan; Ban Phon Rang; Patthalung: Muang;

60, 32, 11, 3p, 6 Ip.

MALAYSIA. . Peninsular Malaysia - Malacca: Malacca; Johore: Kota

Tinggi; Trengganu; Kemaman; 28%, 179, 7L, 2p, 13 1p. Malaysia - Sarawak:

Kuching (type-locality); Sabah: Kota Kinabalu; 16.

SINGAPORE. Mandai Rd.; other unspecified localities; 520°, 422, 9p, 6

lp; 11% and 22 (A. R. Barr collection).

Additional records from the literature. INDONESIA. Sumatra: Padang

(Brug and Edwards 1931: 261); Kalimantan (Brug and Bonne-Wepster 1947:

186).

TAXONOMIC DISCUSSION. Culex curtipalpis is apparently one of the most

common pitcher plant forms in Malaysia and Singapore with the range extending

north as far as southern Thailand. The specimens collected by H. H. Stage,

P. J. Santana and others in Vietnam, which were recorded by Bram (1967a:

77) as curtipalpis are incorrect. I have reexamined these specimens and found

them to be sumatranus. Both species are exceedingly similar in all stages

except for the development of the modified tufts of the male antenna, which are

present in curtipalpis but entirely absent or largely degenerate in sumatranus.,

All stages of curtipalpis except for the female are most distinctive and can

readily be recognized by several features as indicated in the keys and as given

in the above diagnosis. The female is generally very similar to the members

of the brevipalpus complex, from which it can be separated by the presence of

broad, loosely packed vertical patch of pale scales on the upper corner and

posterior border of sternopleuron and by the more numerous cibarial teeth.

Most of the diagnostic features of curtipalpis are remarkably constant and

there is no indication of local or geographical differentiation among the materi-

al examined.

BIONOMICS. The immatures of curtipalpis have been frequently collected

from Nepenthes gracilis in open or partial shaded scrub or secondary rain

forest at sea level to about 40 m (Barr and Chellapah 1963). At Mandor, near

Pontianak, western Borneo (Kalimantan), it was also reported from Nepenthes

mirabilis (Brug 1934: 150). Of the 4 larval collections recently made in Pen-

insular Malaysia, 3 were from N. gvacilisand one from ampullaria, In Sabah,

and Sarawak, numerous adults were collected while resting on leaves of plants

under heavy shade of tropical forest. In Singapore, Colless (1965: 302) noted

that the adults were found resting near the breeding places and exhibited no

attempt to feed on bait animals.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 145

99. CULEX (LOPHOCERAOMYIA) SUMATRANUS BRUG

(Fig. 70)

Culex (Culex) sumatranus Brug 1931: 248 (3, L); Brug and Bonne-Wepster

1947: 187 (distribution).

Culex (Neoculex) sumatranus Brug, Brug and Edwards 1931: 261 (distribution);

Stone, Knight and Starcke 1959: 229 (catalog).

Culex (Lophoceraomyia) curtipalpis (Edwards), Bram 1967a: 76 (in part).

Culex (Lophoceraomyia) sumatranus Brug, Sirivanakarn 1971: 62 (taxonomy).

FEMALE. Wing: 2.7mm. Forefemur: 1.5 mm. Proboscis: 1.8 mm.

Extremely similar to curtipalpis, differing slightly in smaller size, paler

coloration and in the following. Head, Erect scales entirely pale yellowish

brown or golden. Thovax, Mesonotal integument pale brown or reddish brown.

MALE (Fig. 70). As described for curtipalpis, differing from it particular-

ly in the following. Antenna. Flagellar whorls weakly plumose or composed

of relatively fewer long setae; modified tufts of setae and/or scales on F-5 to

F-9 entirely absent or not developed; F-7 sometimes with a rudimentary or

inconspicuous tuft of 4 very short setae on mesal surface.

MALE GENITALIA. As described and figured for curtipalpis from which

it can not be distinguished with certainty, differing slightly in having relative-

ly smaller basimere and less expanded modified apex of distimere.

PUPA. Essentially as described and figured for curtipalpis from which it

differs slightly in smaller size and in pale, cream-colored cephalothorax and

abdomen.

LARVA. As described and figured for curtipalpis from which it is virtual-

ly indistinguishable.

TYPE-DATA. Holotype o* (18.140, marked as type by Brug 1931) with

slide of antenna and genitalia, Dermajoe, Benkoelen, Sumatra, | INDONESIA ],

Nov. -Dec., 1929, S. L. Brug (BMNH).

DISTRIBUTION. Hong Kong, Vietnam, Cambodia and Indonesia. 249

specimens examined: 560°, 459, 3 L, 68 p, 77 1; 26 with associated immature

skins (3 p, 23 lp).

HONG KONG. 4°, 49, 31 (associated with the adults).

VIETNAM. Saigon; Bien Hao; Kontum; Baria; Thoutum; Djiring; Danang;

ASD Ato. 3) 43 68 py THE 23 Ip:

CAMBODIA. Phu Quoc Island; 3° (genitalia only).

INDONESIA. Sumatra: Dermajoe, Benkoelen (type-locality), 10 (holotype).

TAXONOMIC DISCUSSION. Culex sumatranus was very poorly known until

I re-examined the male holotype at the BMNH and transferred it from Neoculex

to Lophoceraomyia (Sirivanakarn 1971: 62-85). As pointed out by Barraud

(1934: 351, footnote), there has been some confusion in associating the larvae

and adults of this species. Brug's description and figure of sumatranus lar-

vae (Brug 1931: 248) were apparently in error. However, judging from his

figure, the specimens probably belong to hewitti which, on the basis of its

occurrence and breeding habitat might be found with sumatranus in the same

pitcher plant. The association of the males and larvae of sumatranus by R. B.

Jackson in Hong Kong (in Barraud's footnote, p. 351) is undoubtedly correct.

I have checked this material against the type-male and numerous males with

associated larvae from Vietnam and found them agreeing well with the brief

diagnosis by Barraud (loc. cit. ).

All stages of sumatranus are exceedingly similar to and indistinguishable

146 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

from curtipalpis except for the male which differs from the latter rather strik-

ingly in the absence of the modified tufts of the male antenna. Both species

have been reported from Sumatra (Brug and Edwards 1931: 261) but have not

yet been found occurring together in Vietnam and Hong Kong.

BIONOMICS. The single collection of the male type (Brug 1931: 248-9) and

numerous collections of the reared specimens of sumatranus from Vietnam as

well as a few from Hong Kong and Cambodia were from pitcher plants, species

unknown. All of these collections were made at a low elevation along the coast.

WILFREDI GROUP

The Wilfredi Group is a complex of 3 closely similar species: wilfredi

Colless, pilifemoralis Wang and Feng and hirtlipalpis n. sp. All stages of the

Wilfredi Group are characterized below in the description of wilfredi. The

group is strongly differentiated from the other 2 major groups: Fvaudatrix and

Mammilifer, especially in the male by apical 0.25-0. 50 of forefemur with a

dense tuft of several strong setae or bristles on anterior surface and in the

male genitalia by the large compact, barrel-shaped and heavily denticulate

phallosome. Both features are unique but the latter, somewhat resembles

those of the /enuipalpis Subgroup of Eumelanomyia (Sirivanakarn 1971: 47-8).

In the general external characters of the adults, the females are apparently

more similar to the Mammilifer Group than to the Fvaudatrix Group in having

the decumbent scales of vertex of the head and the scales on wing veins Ro and

R3 narrow, linear and fine. The males are similar to the Fvaudatrix Group in

the absence of nipplelike prominence on the antennal pedicel, the presence of

modified tufts of scales and setae on antennal flagellomeres 5-10, the develop-

ment of broad, bladelike scales of the modified tuft of antennal flagellomere

9 and in the presence of a distinct false joint at 0.25 from the base of probos-

cis but share with the Mammiailifer Group in the absence of the basal fingerlike

processes of palpus and the absence of dorsal upright setae in the distal por-

tion of proboscis. Except for the unique phallosome, all other features of the

male genitalia, including particularly the proctiger crown are essentially simi-

lar to most members of the Mammilifer Group. The known pupae and larvae,

which occur primarily in general ground pools, both exhibit much overlap with

the Fraudatrix and Mammilifer groups in general and detailed chaetotaxy.

The pupae are rather similar to the Fraudatrix Group and certain ground pool

forms of the Mammilifer Group in having pinna of the trumpet with a slit ex-

tending into the meatus, setae 5-IV 3-6 branched and 6-III-VI usually 4, 5

branched. The larvae show a strong affinity with the Mammilifer Group in

having seta 4-C and seta 1-M relatively strong and long, and seta 8-P single,

but share with most members of the Fraudatrix Group in having seta 14-P

Single.

The Wilfredi Group is known only from Peninsular Malaysia, Thailand,

Vietnam and southern China.

Sirivanakarn: Subgenus Lophocevraomyia in the Oriental Region 147

KEYSTO SPECIES!

FEMALES

(Separation not reliable, see keys to males, pupaeand larvae)

ds Decumbent scales of vertex entirely narrow, linear and fine.

wilfredi

Decumbent scales of vertex relatively broad clavate on anterior dorsal

margin, narrow and Tine) in center) ..5. 2aesles pilifemoralis

MALES

i. Palpal segment 3 with prominent rows of numerous setae in apical

DuSe ik + Omit Abed een ie ud ous hirtipalpis

Palpal segment 3 without prominent rows of numerous setae in apical

2(1). Proboscis with 2-4 long setae on ventral surface distad of false joint;

anterior surface of forefemur with 10-12 strong setae forming

prominent: tuftin apreai0; 25-O0.50.m, elivel. 6a pilifemoralis

Proboscis without any long setae on ventral surface; anterior surface

of forefemur usually with 20 or more strong setae forming promi-

pent taht In Spies] wes: GRMe res bo. MEI Wa RA wilfredi

MALE GENITALIA

(Inseparable, see keys to males, pupaeand larvae)

PUPAE

1. Seta .6-C usually 40 hranched:-o..csih ome ened wens WOR oie wilfredi

Seta GC mamely Goubiesn. becca euch Glow rau seh he pilifemoralis

LARVAE

1, Setae 6-III-VI 3,4 branched; seta 2-X usually triple. ..... wilfredi

Setae 6-III- VI 5 branched; seta 2-X usually double. .. pilifemoralis

06. CULEX (LOPHOCERAOMYIA) WILFREDI COLLESS

(Pies. Thott, ta)

Culex (Lophoceraomyia) wilfredi Colless 1965: 297 (o*); Bram 1967a: 110

(o*, 9; L*; in part).

lFemale, pupa and larva of hirtipalpis are unknown.

148 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

FEMALE. Wing: 3.0 mm. Forefemur: 1.6mm. Proboscis: 1.9 mm.

Medium-sized, general coloration brownish to blackish; in general essentially

conforming to the description of the subgenus with the following diagnostic fea-

tures. Head, Decumbent scales on dorsum of vertex entirely narrow, linear

and fine, scales on upper eye margin pale whitish, forming a narrow distinct

ocular line; erect scales narrow, slender and entirely dark brown; lateral

patch of broad scales paler whitish to grayish. Proboscis with 4 labial basal

setae, 2 lateral ones about 0.5 of palpal length. Cibarial Armature (Fig. 71).

Cibarial bar with a concave row of 50 or more narrow, elongate teeth which

are closely spaced, subequal and abruptly pointed apically. Thorax. Mesono-

tal integument reddish to dark brown; scales narrow, moderately dense, en-

tirely dark brown or almost black. Apu with or without a few pale scales;

pbpn with about 10 or less scattered narrow, scalelike setae cephalad of pos-

terior bristles. Pleuron same color as mesonotum; pp/ with 3 bristles and

0-7 weak setae, scales absent; stp with or without a few scattered pale scales

on its upper corner; 1,2 lower mep bristles present. Legs. Anterior surface

of hindfemur with a broad longitudinal whitish stripe extending from base to

apex, rest entirely dark. Wing. Scales on veins Ro, Rg narrow, linear. Abdo-

men. Terga entirely black scaled; sterna pale yellowish scaled.

MALE (Fig. 73). Sexually dimorphic, particularly in having a dense tuft

of about 20 or more strong bristlelike setae in apical 0.5 of anterior surface

of forefemur. Palpus. Slender, exceeding proboscis by about full length of

segment 5; segment 2 with a dense row of several dark, short setae on lateral

and mesal surfaces; segment 3 very thin, with several tiny setae largely

restricted to basal 0.25 on ventral surface, its apex bears 3-5 bristles; seg-

ments 4,5 very weakly plumose, bristles sparse or relatively few. Proboscis,

Uniformly thick; labial false joint present at 0.25 from base, ventral setae

distad of joint absent; labial basal setae weak, short, hairlike, about 10 in

number. Antenna. Pedicel without distinct spiculose prominence on inner

dorsal surface; flagellar whorls densely long plumose; modified tufts of setae

and scales present on F-5-10; F-5 with a large yellowish or brownish tuft of

8-12 narrow scales which are subequally long, flattened in basal 0.25, linear,

hairlike in apical 0.75, as long as the combined length of the next 5-7 flagello-

meres, preceded dorsally by 2-5 long normal setae and followed ventrally by

o-7 short acute setae; F-6 with a large crumpled dark brownish tuft of about

15 curled setae; F-7 with a ventral comb-shaped, dark brownish tuft of 8-10

weakly curled setae and a thick dark matted tuft of 5-7 subapically twisted

setae on mesal surface; F-8 with a thick dark weakly hooked tuft of 7-10 fused

setae, 3,4 of which are shortened and slightly swollen subapically, the re-

maining longer, distally tapered and smoothly curved; F-9 with 3-5 narrow

yellowish bladelike scales on lateral surface and several strong bristlelike

setae on ventral and mesal surfaces; F-10 usually with 3-4 dark, long bristle-

like setae, sometimes absent or not well differentiated from normal long setae.

MALE GENITALIA (Fig. 71). Segment LIX. Tergal lobe small, bearing 3, 4

moderately strong setae; sternum with 10-17 strong setae in an irregular trans-

verse row towards caudal margin. Basimere. Slender, conical, about 0.30

mm in length; inner tergal surface with a linear row of 5, 6 strong submargin-

al setae and a few other weaker setae basad; marginal setae 7-10, about 0.5

of length of submarginals. Subapical Lobe. All usual specialized setae and

leafiets present; rodlike setae a-c subequal, largely overlapping; setae d-f

strong, 5,6 in number, all flattened, bladelike, weakly bent dorsad; leaflet go

narrow, slightly longer and broader than setae d-f; leaflet gy lanceolate,

shorter or subequal to leaflet 893 seta strong, situated close to bases of

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 149

setae a-c. Distimere. Slender, normal, distally tapered into a slender re-

curved apex; subapical claw small; ventral and dorsal subapical seta present,

minute, situated opposite each other. Phallosome. Large, dark, compact,

barrel-shaped and strongly sclerotized, tergal surface heavily denticulate;

lateral plate rectangular in lateral aspect, with a short strongly denticulate

internal process basad. Pyroctiger. Apical crown large, composed of several

coarse spinelike spicules; paraproct and cercal sclerite broad, well sclero-

tized; cercal setae 3,4.

PUPA (Fig. 71). Abdomen: 2.6mm. Paddle: 0.65 mm. Trumpet:

0.62 mm; index 10-12. Cephalothorax and abdomen more or less uniformly

pale cream-colored or slightly darkened. Tvumpet, Slender, long, apex

lightly to moderately widened; pinna with slit extending to meatus. Complete

chaetotaxy as figured; all setae well developed. Cephalothorax, Seta 1-C

usually 4 branched (4-6); 3-C usually triple (2-4); 5-C usually 6 branched (5-8);

shorter or subequal to 7-C; 8-C usually 4,5 branched (3-5); 9-C usually triple

(3-5). Metanotum, Setae 10,11-C double; 12-C usually 4 branched (3-4). Ab-

domen. Seta 1-II dendritic or multibranched; 1-III, IV usually 9 branched (7-

11); 1-V usually 6 branched (5-8); 1-VI usually 5 branched (4-7); 1-VII usually

4 branched (3-5); 3-I-III double; 5-IV usually 4 branched (3-6), as long as seg-

ment following; 5-V usually double (2-3), about 1.5 times as long as segment

following; 5-VI double, subequal to 5-IV; 6-III-VI usually 4,5 branched (3-6);

9-VII usually triple (2-3); 9-VIII 4-7 branched. Paddle. Broad, pale whitish

to almost transparent; midrib weak, lightly pigmented; setae 1,2-P present,

minute.

LARVA (Fig. 72). Head: 0.78 mm. Siphon: 1.8 mm; index 9-10. Sad-

dle: 0.34 mm; siphon/saddle ratio 5. Generally similar to most members

of the Fraudatrix and Mammilifer groups, complete chaetotaxy as figured.

Head, Pale yellowish; seta 1-C relatively long, slender, spiniform, its

length usually as long as distance between bases of the pair, or sometimes

slightly shorter; 4-C single, strong, about 2 times as long as distance between

bases of the pair; 5,6-C strong, subequal, reaching well beyond mouth brush;

5-C usually double (2-3); 6-C double; 7-C usually 6 branched (5-7); 10-C

double; 11-C 4,5 branched; 13-C usually 6 branched (4-7); 14-C 2-6 branched;

16,17-C developed. Antenna relatively long, nearly as long as head; pigmen-

tation entirely pale as head capsule; spicules numerous, rather fine; seta 1-A

large, strongly plumose; 2,3-A strong, rather long, bristlelike, situated sub-

apically. Mental plate with 6,7 lateral teeth on each side of median tooth.

Thorax. Spicules absent; seta 3-P single; 4-P double; 7-P usually double,

sometimes triple; 8-P single, subequal to 7-P; 14-P single; 1-M long, usually

single (1-2), subequal to 3-M; 8,9-M 4,5 branched; 7-T usually 6 branched;

9-T 5,6 branched; 13-T usually 5 branched (4-7). Abdomen. Spicules absent;

setae 6-I, II triple; 7-I double; 6-III-V usually 4 branched (3-4); 6-VI usually

triple (3-4); 1-III-VI usually 4 branched (3-4); 1-VII usually 5 branched (5-7);

1, 5-VIII 4 branched; 2-VIII single; 3-VIII usually 7 branched (6-7). Comb

scales numerous, 44-50, all apparently subequal, with rounded apical

fringe of evenly fine spicules. Saddle complete, lightly spiculate on posterior

caudal margin; seta 1-X 3,4 branched; 2-X usually triple (2-3); anal gill slen-

der, fusiform, 1.5 times as long as saddle. Siphon. Slender, long, lightly

tapered in distal portion; pale yellowish, concolorous with head capsule and

saddle; pecten teeth 10-12, 4,5 distal teeth with a fine barb of 5-7 graded

denticles; subventral tufts 4 pairs, all weak, subequal, usually double (1-2),

slightly shorter than siphonal width at point of attachment; median caudal fila-

ment of spiracular apparatus dark, very distinct.

150 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

TYPE-DATA. Holotype “* with slides of antenna (CH 190) and genitalia

(CT 646), Malaya | MALAYSIA], date and collector not specified (ANIC).

DISTRIBUTION. Known only from Peninsular Malaysia and Thailand. 152

specimens examined: 680", 339, 51 L; 42 with associated immature skins

(11: pjn81Tp). |

THAILAND. Chiang Mai: Doi Sutep; Lampang: Ngao; Tak: Doi Sam Sao;

Nakhon Ratchasima: Ban Tha Ma Prang; Nakhon Nayok: Khao Yai, Khao

Kheow; Chanthaburi: Khao Sai Dao; 490", 9°, 47 L, 11 p, 18 lp.

VIETNAM. Ban Me Thuot; 1.

MALAYSIA. Peninsular Malaysia. Locality unspecified; 10° (holotype);

Pahang: Gunong Benom; Cameron Highlands; Perak: Cameron Highlands Rd. ;

17%, 242, 4L, 13 lp.

TAXONOMIC DISCUSSION. Culex wilfredi was tentatively assigned to the

Mammilifey Subgroup of the Mammilifer Group by Colless (1965: 297-8), an

interpretation subsequently followed by Bram (1967a: 110-3). Although wil-

fredi apparently shares with the Mammilifer Group several features of the

male and larva as pointed out by the latter author and as indicated in the above

discussion of the Wilfredi Group, the unique male features, including particu-

larly the extraordinary type of the phallosome, the peculiar development of a

prominent tuft of several strong setae on the forefemur, the absence of distinct

spiculose prominence on the antennal pedicel and a few other features shared

with the Fraudatrix Group, strongly suggest that it represents a distinct lineage

of Lophoceraomyia. On this basis, wilfredi and its closely similar forms,

pilifemoralis and hirtipalpis, are considered as belonging to a separate group

or Wilfredi Group. For the separation of wilfredi from pilifemoralis and

hirtipalpis, see the keys and the description of the latter 2 species.

BIONOMICS. Culex wilfrediis restricted to mountainous areas at eleva-

tions from 500-1,500 m. The immatures have been frequently found in small

ground pools at stream margins or stream pools and occasionally in puddles,

marshy depressions, seepages and animal foot prints under partial or heavy

shade of primary tropical forest. In Perak, Peninsular Malaysia, one collec-

tion (No. 1574) was reported to come from a bamboo internode, but this is

probably an error due to contamination. The tree hole collection noted by

Bram (1967a: 113) in Thailand also appears to be an error as there has been

no record from this habitat in all collections from that area. The 2 type-males

from Malaysia (Malaya) noted by Colless (1965: 298) to be associated with a

collection in pitcher plants were probably from ground pools on the basis of

the present records. At Doi Sutep, Chiang Mai, Thailand, several adult males

were collected resting near the breeding sites, especially among veuetation

along stream margins.

07. CULEX (LOPHOCERA OMYIA) PILIFEMORA LIS WANG AND FENG

(Pio. 73)

Culex (Lophoceratomyia) pilifemoralis Wang and Feng 1964: 37 (o*),

Culex (Lophoceraomyia) wilfredi Colless, Bram 1967a: 110 (c*, 2, L*, in

part).

FEMALE. Wing: 2.8mm. Forefemur: 1.3 mm. Proboscis: 1.7 mm.

Extremely similar to wilfredi from which it cannot be distinguished with cer-

tainty, differing slightly in smaller size and in having broader decumbent

scales on ocular line of vertex.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 151

MALE (Fig. 73). In general as in female, differing from wilfredi particu-

larly in the following features. Palpus. As long as or slightly longer than

proboscis; segment 2 apparently without conspicuous rows of dark, short setae

on lateral and mesal surface. Proboscis. Labium with 2-4 long ventral setae

distad of false joint, these setae are 4,5 times as long as labial width at point

of attachment. Antenna. Flagellar whorls moderately plumose; modified tuft

of F-5 apparently smaller, shorter, whitish or yellowish, but not brownish;

composed of 10-12 narrow scales, dorsalmost 4-7 scales short, hairlike or

somewhat lanceolate, as long as the length of 1 flagellomere, next 4,5 scales

long, linear, as long as the next 4, 5 flagellomeres, followed ventrally by 4, 5

long, normal setae and mesally by a minute tuft of 6 acute setae. Legs, An-

terior surface of forefemur with a prominent tuft of 10-12 strong setae in api-

cal 0.25-0. 50.

MALE GENITALIA. Essentially as described and figured for wilfredi

from which it is virtually indistinguishable.

PUPA. Abdomen: 2.2mm. Paddle: 0.60 mm. Trumpet: 0.60 mm,

index 9. As described and figured for wilfredi, differing from it particularly

in having seta 8-C usually double, or sometimes triple.

LARVA. Based on 3 whole larvae tentatively associated with above pupa

and adults. Exceedingly similar to wilfredi in general and detailed chaetotaxy,

differing from it in the following. Head. Seta 1-C shorter, about 0.5 of the

distance between bases of the pair. Antennal shaft pale proximally, darkened

beyond insertion of seta 1-A. Abdomen, Setae 6-III-VI usually 5 branched,

sometimes 6; 1-V, VI 5,6 branched. Saddle seta 2-X usually double (2-3).

October 1958, Hsing-Hsiang Wang and Chung-Ying Feng (Dept. Parasitology,

Seventh Military Medical College, China).

DISTRIBUTION. Known only from northern Thailand and southern China.

30 specimens examined: 220", 52, 3 L; 8 with associated pupal skins.

CHINA. Yunnan, as indicated in the type-data (Wang and Feng 1964).

THAILAND. Chiang Mai: Doi Sutep; Lampang: Ngao; Ban Pho Daeng; 22",

Ors tue te

TAXONOMIC DISCUSSION. Culex pilifemoralis was originally described

by Wang and Feng (1964: 40) based on 2 males from Szemao, Yunnan, China.

Several males of this species were discovered among numerous specimens

previously identified as wilfredi by Bram (1967a: 110-3) from northern Thai-

land. In addition, a number of adults reared from pupae and a few whole lar-

vae have also been obtained from Lampang, Thailand. The assignment of the

Thailand material to this species is only provisional, pending a thorough ex-

amination of the type which was poorly described. In checking the males from

Thailand against the description and figure by Wang and Feng (loc. cit.), I

found them agreeing well in the features of the palpus, modified tufts of anten-

nal flagellomeres 5-9 (as segments 6-10 of the authors) and in the number of

strong setae on the anterior surface of forefemur. The only discrepancy is

in the antennal pedicel which was described by Wang and Feng (as torus) as

having a blunt prominence on the inner side. This feature is not developed in

male specimens from Thailand except for one slide which exhibits a slightly

swollen pedicel due to flat preparation.

Although pilifemoralis is exceedingly similar to wilfredi and hirtipalpis,

the differences from the latter 2 species in the male characters as indicated

in the key and as described above are constant. In northern Thailand, they

were found to be sympatric with one another without any indication of overlap

or intergradation in the male diagnostic features.

152 Contrib. Amer. Ent. Inst.., vol. 13, no. 4, 1977

BIONOMICS. As in wilfredi, pilifemoralis occurs at high elevations in

mountainous areas and is typically a ground pool breeder. The larvae and

pupae have been collected from stream pools, puddles and animal foot prints

in the vicinity of forest streams. Most adults were caught while resting on

vegetation near their breeding sites.

98. CULEX (LOPHOCERAOMYIA) HIRTIPALPIS N. SP.

(Fig. 73)

Culex (Lophoceraomyia) wilfredi Colless, Bram 1967a: 110 (o*, 2, L*, in

part).

FEMALE. Unknown.

MALE (Fig. 73). Exceedingly similar to wilfredi and pilifemoralis, differ-

ing in the following features. Palpus, Segment 3 with prominent lateral and

mesal rows of numerous hairlike setae which are very dense and gradually in-

crease in length towards distal portion, proximal setae 2,3 times as long as

segment width, distal ones twice as long as proximals. Pyroboscis. Ventral

setae distad of false joint absent, as in wilfredi. Antenna. Essentially as

described and figured for wilfredi. Legs, Anterior surface of forefemur with

more numerous strong setae, forming denser tuft from basal 0.25 to apex,

lateral dorsal surface also with a distinct tuft of numerous weaker setae in api-

cal 0.20. 3

MALE GENITALIA. As figured and described for wilfredi from which it is

virtually indistinguishable.

PUPA and LARVA. Unknown.

TYPE-DATA. Holotype “ (M411-7) with slides of antenna and genitalia,

Doi Sutep, near temple, Chiang Mai, THAILAND, 13 Jan. 1953, Prayoon Chait

and D. C. Thurman (USNM); paratypes: 3c (M481-3; M569-273; M462a) with

slides of antenna and genitalia, same locality as holotype (USNM).

DISTRIBUTION. Known only from Thailand. Specimens examined: 6.

THAILAND. Chiang Mai: Doi Sutep; 4“ (as indicated in the type-data); 1o

(No. 559, genitalia lost); Lampang: Ngao; 1c.

TAXONOMIC DISCUSSION. The males of hirtipalpis were discovered

among the numerous specimens identified as wilfredi by Bram (1967a: 110-3)

from Chiang Mai and Lampang, Thailand. The presence of the prominent rows

of long setae on palpal segment 3 and the more numerous setae on the anterior

surface of the forefemur in hirtipalpis are constant and diagnostic. There is no

indication that there is an overlap in these features with the males of either

wilfredi or pilifemoralis.

BIONOMICS. Culex hirtipalpis occurs at a high elevation in mountain

forests. At the type-locality in Chiang Mai and at Lampang, Thailand, the

males were collected resting in association with specimens of wilfredi and

pilifemoralis. The breeding sites are unknown, but most probably are ground

pools in dried up stream beds or along the margin of mountain streams.

Sirivanakarn: Subgenus Lophocevaomyia in the Oriental Region 153

ACKNOWLEDGMENTS

Iam grateful to Dr. Botha de Meillon for his encouragement and assistance

during the early phase of this study, and to E. L. Peyton and Ronald A. Ward

for continued support in completing the remainder of the work. I am indebted

to Donald H. Colless, Division of Entomology, CSIRO, Canberra, Australia;

Bruce F. Eldridge, Department of Entomology, Walter Reed Army Institute

of Research and Rampa Rattanarithikul, U. S. Army Medical Component -

SEATO, Bangkok, Thailand for valuable comments and suggestions in improv-

ing the manuscript; Phuangthong Malikul for preparing specimens and numer-

ous slides; John Lane and Thomas V. Gaffigan for curatorial assistance;

Chien C. Chang (Department of Entomology, North Carolina State University,

Raleigh) for preparing most of the illustrations; Thelma Ford Smith for com-

pleting the remainder of illustrations; Helle Starcke and Owilda J. R. Curtis

for typing preliminary drafts and Janet D. Rupp for final preparation of text

copy for lithoprinting.

I am also grateful to the following individuals and institutions for generous

support in sending the material (including some types) on loan or as gifts:

Douglas J. Gould, U. S. Army Medical Component - SEATO, Bangkok, Thai-

land; Shivaji Ramalingam, Department of Parasitology, University of Malaya,

Kuala Lumpur, Malaysia; Peter F. Mattingly, British Museum (Natural His-

tory), London, England; Donald H. Colless, Division of Entomology, CSIRO,

Australia; F. E. Baisas (deceased), Manila, and G. L. Alcasid, National Mu-

seum, Manila, the Philippines; A. Ralph Barr, School of Public Health, Uni-

versity of California, Los Angeles; B. N. Mohan, Coonoor, Madras and N.

L. Kalra, National Institute of Communicable Diseases, Delhi, India; T. Kuri-

hara, Teikyo University, Tokyo and Yoshito Wada, Nagasaki University, Naga-

saki, Japan; J. C. Lien, Taiwan Provincial Malaria Research Institute,

Nankang, Taipei; Bernice P. Bishop Museum, Honolulu, Hawaii; Institute

Voor Tropische Hygiene, Amsterdam, the Netherlands; Field Museum of

Natural History, Chicago; California Academy of Sciences, San Francisco, and

the Academy of Natural Sciences, Philadelphia.

Finally, Iam thankful to Ronald A. Ward for editing the manuscript.

154 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

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BREAD Het.

1941. An observation on the tree snake Passevita prasina. Malayan

Nat. J. 2: 3-4.

SENIOR-WHITE, R.

1927, Notes on Ceylon mosquitoes, -II. The larvae of the commoner

non-anopheline mosquitoes. Spolia Zeylan. Ceylon J. Sci. B.

14: 61-76.

SIRIVANAKARN, S.

1968. The Culex subgenus Lophoceraomyia in New Guinea and

Bismark Archipelago (Diptera: Culicidae). Pacific Insects

Mono. 17: 75-186.

1971. Contributions to the mosquito fauna of Southeast Asia. XI. A

proposed reclassification of Neoculex Dyar based principally

on the male terminalia. Contrib. Am. Entomol. Inst. (Ann

Arbor) 7(3): 62-85.

ey Contributions to the mosquito fauna of Southeast Asia. XIII.

The genus Culex, subgenus Eumelanomyia Theobald in South-

east Asia and adjacent areas. Contrib. Am. Entomol. Inst.

(Ann Arbor) 8(6): 1-86.

1973: Descriptions of two new species of Culex (Lophoceraomyia) with

notes on three other species from the Papuan subregion (Diptera:

Culicidae), J. Med. Entomol. 10: 212-6.

160

STONE, A.

1957.

1961.

1963.

1967.

1970.

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Notes on types of mosquitoes in the Hungarian National Museum

(Diptera, Culicidae). Ann. Entomol. Soc. Am. 50: 171-4.

A synoptic catalog of the mosquitoes of the world, Supplement I

(Diptera: Culicidae), Proc. Entomol. Soc. Wash. 63: 29-52.

A synoptic catalog of the mosquitoes of the world, Supplement II

(Diptera: Culicidae). Proc. Entomol. Soc. Wash. 65: 117-40.

A synoptic catalog of the mosquitoes of the world, Supplement III

(Diptera: Culicidae). Proc. Entomol. Soc. Wash. 69: 197-224.

A synoptic catalog of the mosquitoes of the world, Supplement IV

(Diptera: Culicidae). Proc. Entomol. Soc. Wash. 72: 137-71.

STONE, A. and R. M. BOHART.

1944,

Studies on mosquitoes from the Philippine Islands and Austral-

asia (Diptera: Culicidae). Proc. Entomol. Soc. Wash. 46:

205-25.

STONE, A. and K. L. KNIGHT.

1957.

Type specimens of mosquitoes in the United States National

Museum: IV, The genus Culex (Diptera, Culicidae). J. Wash.

Acad. Sci. 47: 42-59.

STONE, A., K. L. KNIGHT and H. STARCKE.

1959.

A synoptic catalog of the mosquitoes of the world (Diptera,

Culicidae). Thomas Say Found., Entomol. Soc. Am. 6, 358 p.

THEOBALD, F. V.

1901.

1905a.

1905b.

1907.

1910.

A monograph of the Culicidae or mosquitoes. Vol. I, Br. Mus.

(Nat. Hist.), London, 424 p.

Some new mosquitoes from Ceylon. J. Bombay Nat. Hist. Soc.

16: 237-50.

A catalogue of the Culicidae in the Hungarian National Museum

with descriptions of new genera and species. Am. Mus. Natl.

Hung. 3: 61-119.

A monograph of the Culicidae or mosquitoes. Vol. IV, Br.

Mus. (Nat. Hist.), London, 639 p., 16 pl.

Second report on the collection of Culicidae in the Indian

Museum, Calcutta, with descriptions of new genera and species.

Rec. Indian Mus. 4: 1-33.

WANG, HSING-SHIANG and FENG, CHUNG-YING.

1964.

Description of three new species of Culex (Lophoceratomyia).

Acta Zooltaxon. Sin. 1: 37-41 (in Chinese with English transla-

tion).

17.

16.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 161

LIST OF FIGURES

Male genitalia and modified tufts of antennal flagellomeres 7-8 of C.,

minutissimus and alorensis.

Pupa, female cibarial armature, male genitalia and modified tufts of anten-

nal flagellomeres 7-8 of C. infantulus.

Larva of C. infantulus.

Pupa, female cibarial armature, male genitalia and modified tuft of anten-

nal flagellomere 5 of C. cinctellus,

Larva of.C. cincieilns.

Male genitalia, modified tuft of antennal flagellomere 5 and female cibarial

armature of C. fullevi; male genitalia and modified tufts of antennal flagel-

lomeres 6-10 of C. seniovi.

Pupa, female cibarial armature and modified tufts of antennal flagellomeres

5and:8 of C. vabithovacis,

Larva of C. rubithoracis.

Pupa, male genitalia and modified tuft of antennal flagellomere 5 of C.

niger.

Larva of C. niger.

Male genitalia and modified tufts of antennal flagellomeres 95-10 of C.

gibbulus.

Male genitalia and modified tufts of antennal flagellomeres 5-10 of C.

inculus.

Pupa and larva of C. inculus,

Pupa, female cibarial armature, male genitalia and modified tuft of anten-

nal flagellomere 5 of C. quadripalpis.

Larva of C. quadripalpis.

Female cibarial armature, male genitalia and modified tuft of antennal

flagellomere 5 of C. aculeatus; male genitalia and modified tufts of anten-

nal flagellomeres 5-10 of C. pavaculeatus.

Male genitalia and modified tufts of antennal flagellomeres 5-10 of C.

aestivus.

Larva, male genitalia and modified tufts of antennal flagellomeres 5-10 of

C. rveidt.

162 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

19. Pupa, female cibarial armature, male genitalia and modified tufts of

antennal flagellomeres 5 and 8 of C. varviatus,

20. Larva of C.. variatus.

21. Male genitalia and female cibarial armature of C. josephinae and cubitatus.

modified tuft of antennal flagellomeres 5 and 8 of C. cubitatus.

22. Male genitalia and modified tuft of antennal flagellomere 5 of C. gvaci-

cornis; female cibarial armature, male genitalia and modified tuft of an-

tennal flagellomere 5 of C, whartoni.

23. Pupa, female cibarial armature, male genitalia and modified tuft of an-

tennal flagellomere 5 of C. macdonaldi.

24. Larva of C. macdonaldi,

20. Male genitalia and modified tuft of antennal flagellomere 5 of C. pairoji.

26. Larva of C. pairoji.

27. Pupa, female cibarial armature, male genitalia and modified tuft of an-

tennal flagellomere 5 of C, alphus.

28. Larva of C. alphus,

29. Pupa and male genitalia of C. impostor.

30. Pupa, male genitalia and modified tuft of antennal flagellomeres 5 and 8

of C. traubi.

31. Larva of Co. teaubi.

32. Male genitalia and modified tufts of antennal flagellomeres 5-8 of C.

uniformis and lavatae,

33. Larva of C. uniformis.

34. Pupa, female cibarial armature, male genitalia and modified tufts of

antennal flagellomeres 5-9 of C. mammilifer.

30. Larva of C. mammilifer.

36. Male genitalia, palpus, proboscis and modified tufts of antennal flagello-

meres 5-9 of C. wardi.

37. Male genitalia and modified tufts of antennal flagellomeres 5-10 of C.

demissus.

38. Larva of C. demissus.

39. Pupa, male genitalia and modified tufts of antennal flagellomeres 5 and 8

of C. ganapathi.

40.

41.

42.

43.

44.

49.

46.

47.

48.

49.

o0.

ol.

O2.

03.

04,

Oo.

o6.

Ov.

08.

09.

60.

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 163

Larva of C. ganapatht.

Male genitalia and modified tufts of antennal flagellomeres 5 and 8 of C.

spiculosus.

Larva of C, spiculosus,

Pupa, male genitalia and modified tufts of antennal flagellomeres 5-9 of

C. minor.

Larva of C, minor; denticulation of larval pecten teeth of C. bicornutus,

Male genitalia of C. bandoengensis and tuberis.

Larva, male genitalia and modified tufts of antennal flagellomeres 5-8 of

C. kuhnsi.

Male genitalia and antennal flagellomeres 5-11 of C. cvassicomus.,

Male genitalia and modified tufts of antennal flagellomeres 5-9 of C,

incomptus.

Pupa, male genitalia and modified tuft of antennal flagellomere 5 of C,

bengalensis.

Pupa, male genitalia and modified tufts of antennal flagellomeres 5-8 of

C. peytont.

Larva of C. peytoni.

Larva, male genitalia and modified tufts of antennal flagellomeres 5-8 of

C. eukrines,

Pupa and male genitalia of C, pholeter.

Larva of C, pholeter.

Male genitalia, palpus, proboscis, antennal pedicel and modified tufts of

antennal flagellomeres 9-9 of C. flavicornis.

Larva and pupa of C. flavicornis,

Pupa, male genitalia and modified tuft of antennal flagellomere 5 of C.

lasiopalpis.

Pupa, female cibarial armature, male genitalia and modified tufts of an-

tennal flagellomeres 5-9 of C. navalis,

Larva of C. navalis.

Larva, male genitalia and modified tufts of antennal flagellomeres 5-9 of

C. coerulescens.

164

61.

62.

63.

64.

65.

66.

67.

68.

69.

70.

12.

73.

Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

Pupa, female cibarial armature, male genitalia and modified tufts of an-

tennal flagellomeres 5-9 of C. hewitt.

Larva of C. hewitti.

Pupa, female cibarial armature, male genitalia and modified tufts of an-

tennal flagellomeres 5-9 of C. jenseni.

Larva of C. jenseni.

Pupa, female cibarial armature, male genitalia and modified tufts of an-

tennal flagellomeres 5-9 of C. brevipalpus.

Larva of C. brevipalpus.

Male genitalia and modified tufts of antennal flagellomeres 5-9 of C,

lucavis and eminentia.,

Pupa, male genitalia and modified tufts of antennal flagellomeres 5-9 of

C. acutipalus.

Pupa, female cibarial armature, male genitalia and modified tufts of

antennal flagellomeres 5-9 of C. curtipalpis.

Larva of C. curtipalpis; male antenna of C. sumatranus.

Pupa, female cibarial armature and male genitalia of C. wilfredi.

Larva of C. wilfredi.

Male antenna of C. wilfredi; male palpus, proboscis and forefemur of

C. wilfredi, pilifemoralis and hirtipalpis.

\d-f

C: o antenna

(i)

b crown dorsal beaklike process

dorsal lobe (DL)

subapical

lobe

submarginal

setae

ventral -

lobe :

0.1

marginal

setae

aN pee phallosome (lateral view)

N proctiger

IXT

basimere

So Oe eee et

o antenna

phallosome (lateral view)

ee | a alorensis

% uaa N

a CN

Hi etal eee

reticulate dorsal

14 beaklike process

phallosome

infantulus

Sy!

; SW“

— aS

= ze s: P

awk

pane er an

Me a 6 Shc.

eee J

+ et Bt.

2 o owt

—submarginal

setae

cinctellus

rithm oF iT: mM TT MTT

“ee

1S)

2 cibarial armature

phallosome (lateral view) ,

proctiger

phallosome (lateral view )

o antenna

seni/ori

Lae

3 f

5A > Sly

a SS Qt)

: ASS |

ae C2) }

? Sy a

e F j//

~- mee |

We 2. et

proctiger

Pe 0] ot

C. Aang

rubithoracis

proctiger

niger C. Chang

ects tel

cig rut ses

bay awe S's wis

beste feat

erased jr

SS ae

Titi eh te ie ae

Ta a a

Fig. 11

o& antenna

proctiger

phallosome

gibbulus

Fig, IZ

)

phallosome

(lateral view

proctiger

IX-T

& antenna

inculus

Tif KR

Yfy) HNN

ly \ \

on 4

proctiger

f iy, wih

Bee a wed,

Ide ee

ow,

oF o

ba {2

Bi. =

proctiger

phallosome (lateral view )

-————— 0.] ——__——"

aestivus

2 2

9 Pes : =

Bee! et

ROS

o& antenna

phallosome

(lateral view)

9 cibarial armature

phallosome

I 0.1 3

proctiger

Fig. 21

Uy MTT AN

Crvenmnen

2 cibarial armature

freee OQ) | ee

ee to ts me NN

josephinae

0.1

fii iN

Gj YY) CQ

\ Uf l mg |

fii

4 Eisen armature

“

eh ene a ne é

: <

od id

proctiger

0.1

phallosome (lateral view)

were e = By.

eas

> hdan

p> te a

phallosome

fat MES AU 8 peer eee

whartoni

9 cibarial armature

at epee on

3 Fig. 29

proctiger

phallosome (lateral view)

phallosome

pairoj!

phallosome (lateral view )

proctiger

alohus

C. Chang

13 10 i

13 e ;

rie

internal process

external process

phallosome

Impostor

#5. —spiculose

apical lobe

| : ~ L seta process

spiculose

apical lobe

proctiger

internal process

traubi

N ts

lee TET org

am Tee

3 4h),

Sj fro

N Hy}

14%

traub/

Fig. 32

® wd phallosome (lateral view)

abs

d ee ay brn

ty oe: : a.

£43

One bd A

4 4 J

pen ee

Seite

proctiger

phallosome (lateral view)

o’ antenna

0.1

proctiger

| /avatae C. Chang

<A ZEEE

Z/ij44}'

Chief)

5 Of

o” antenna

AE te its is iain Se sere gt %.- ae

submarginal Ne

setae

proctiger

——— O)|—_—

mammilifer

—_ pet Ewe 5 a —

A So. PR.

Se SS

fh 17

# MH

1 Li

ee hil

aa Qn SLZ/

o” antenna

phallosome

proctiger

C ee.

wardi 7

phallosome

eed sae

meee

~----

2°

— —"

eee

<——— =~, ra

==<-- ~—---"

“=. oy

ear en ee a

ane

Ci ss

s----"

“s

phallosome

(lateral view)

ISSUS

dem

6a ARAL RE tat

8 ee

Se “a

demissus

\ ~S . v4

—— a

Sey

Sd Ey

RZLLALLI

; 1

114

fl 2

Bis

ade

tac

epee

ganapathi

f a

Ele

ed:

ganapathi

proctiger

phallosome

(lateral view)

phallosome

fee Oo

rer ; "

LTS i

sre tay %; eyes

Wahcnan pate <

Oe Be

IX ST

spoiculosus

phallosome (lateral view)

o” antenna

= ~ — exe Xe = . oe 5 Ss “8

wr 2 = a & AY... 3 <7. 7;

x =k - 5 i A hy fj

, 1. ’ d\ y / |

4 y

Ni D '

proctiger

phallosome

IX-ST

phallosome (lateral view)

phallosome

dt ee

= P

are : wv: 4

bien et nal SPST eR

eee Se Te rs,

, \ Xe: :

eS ee

proctiger 0.1

tuberis

feeb () Sp enact

phallosome

TY TARY

Noth

Wit

kuhnsi

proctiger

phallosome (lateral view)

eS ae

oo"

“se

Crass/COMUS

proctiger

internal process

apietions apical lobe

»\—eX ternal

be process

phallosome

SSS

é SO a" ~

UMass WERE

. WE egee

vinegar Ut

BN EER LS Lt Da

ERS LNL Ly SE bay UT ;

(ev i lia aay Cra

o antenna

a ey

iIncomptus

on ~ i

(AS

ZE se

=) fe Ju fo} (f]

or!

proctiger

~ 4 ry 24°" °

ae raat Sgt etsy

PETIT AGT TOLL eohed TOD ys Red ae

yer "l 4a 4 #4 se

J, iat alts ty tl

i, L sat ey FE aye 2

Pryde

Layiy

i 4 iad

ae aE Tt,

ist

ac bengalensis .

Fig. 50

2 Who

= J

A — A

ugh WN "14 ae

0.1

= oy" iW 6

rt ttre, i "

Foe: ak seat mati

TTY

St fabs wuts

eukrines

proctiger

phallosome

Th Ayo 18 4

Bo a

CYC nee an Oe

MU s ne rey,

’

icc iid SY pholeter C.

a oe ee

4 3

7 f'S LZ} a

a———])

Fig. 99

Se ace, ~ 2

~F x

o antenna palpus & proboscis

antennal pedicel

proctiger

phallosome (lateral view)

flavicornis

SREY

> KR

2 SAX

4 N

43 W477

NS a

aS :

—

LAWN

\ NN ;

flavicornis

oN |

lasiopalpis

proctiger

UL, ARE.

(

phallosome

navalis

phallosome

proctiger

coeru/escens

2 cibarial armature

——_—_——— 0.]

proctiger

phallosome

hewitti

mature

9 cibarial ar

proctiger

one jenseni

proctiger

lateral view)

Y Y,

ae EES — previpalous

phallosome

“fy

/ucaris

My,

eminentia

“<M vv ual! ns

as t\)

SO OE * Sag

o& antenna

proctiger

acutipalus

- 2

“-seec-

phallosome (lateral view)

curtipalpis

o& antenna

Sumatranus

Vil 0.5

cS PT

14 ”)

a curtipalpis if

fom

| phallosome

0.1

phallosome (lateral view)

proctiger

wilfredi/

f Ay Uh i (

ve Gurchy}

UH 'f (9) R ON; y

iY ti, Wie i

te eS

i Has

iy i all

Wee, 2 RING

Ait 0.5 WAX N djs

sf) \\ GA hy Ky "i

AS ENN

| Ni Wi NY M

A sa ND

Mee Hi NAW

MONE ay Nae

i Le A

PN DS ANAT AD

ea ce

ring Ny tt A

Bee | Hs At

ZA ah

hh eo WN

v0 1 (OF i) AN i

aunty MN AND NG

aay oO” Hi NY

Zeek HK iy

a i . i

BRIA) "i NK

ZY: i nD

, WE 4) fi "4, \\ Hi)

va i wh

iis i “Ny

we AN) Ly \ (|

7/7 [BBW AA) wv:

fib wy Ky) ij

iN bi Whe

\ 4 (e, p

Sop YK, YO

427 \yfi 4% |

hy wie

forefemur A Rh

‘ Gop ut h/ /) |

A Gi y a i

5 All at

LY palpus ght

y ae

o” antenna tif proboscis ‘

wi/fred/ Wi

AR ADS ASA }

HN ny Ay

Dew DA\\e uy , f )

HNN int ANAT

di if Vi aah if RAYNE

HD Ad Aa

Wii v(t \ wild 0.5 AT ERAN iy!

2 a AY) OD} ry qin\ hit qu Ny) \

0.5 vi W i HM iN OK Ky Me iy ay

Lint Hy yl ROA OW ANKSTENN

He i A(t hi SI) I) 4 2 aN Ay wi l

rs many UMN viewed ‘4 Hh “i u(\ ih

Melia aie | AR SNS

h Ato Tee i Ri iH)

id M1 Yeh ares aye Of DEO

oA RNa Za He Sh

TB IRDA psse ny AY ARNO YY

i ae a ei

| I) ne “So ‘ f i iat

‘git TON BR rel He | NR

AR atin ia) om

gS ANNA Aen 7 VN

secs) SS Zi A Nt

Hi AGIA fad ra Be 44

3 & poe Soe, GN inlay

fin ny easy ZA A KK

a veo a) 3 Ak i

he iy, Wh a aN TR \ iy) ) \

oi is WN NE oO” SNS MWA

Wa Hh Ky ZF en oi ® int

“i NH ZZ sis ZN bY

fh i, aN i \ MN

Dry Web Syed ou ) yh: MU WU

oi Nhe SK te ‘aa

van ) ANY eae eR | a | 1f"e) Oh

i wie, 4 Zn oH Uf Za i) He

Ay" UR RUT ian 7) /] VAN ) tu

Ty mya AYA) S| DD NY

i ie Zi" ; no

A qd Cony NYU XU a

RN AN bday

Gp di, PN OU

Jif wi LA Pie

forefemur 4 def iN) 7 ~ BEN ti

Y a7") rye A RIT ae YUU

Hf! palpus 2m vim\\; Mi an

JOE M WHAT : W Wd)

fg)’ & BN palpus IAS

Zot 7 age 423 I y)

5 proboscis AUhUn & pana

ye oN forefemur proboscis My uin

Gh / I)! i Hh\))

Saad I, qin

id i "S

pilifemoralis hirtipalpis

iin ae SE EE «Sanne re

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 239

APPENDIX A: TABLE OF DISTRIBUTION AND BREEDING HABITATS

Legend

- Artificial container

- Bamboo

- Crab hole

- Ground pools (puddle, swamp, marsh, etc.)

- Leaf axils

- Pitcher plant

~ Rock pool

~ Tree hole

Distribution Symbols

= — indigeneous, endemic

APPENDIX A: TABLE OF DISTRIBUTION AND BREEDING HABITATS

DISTRIBUTION

Indonesia

c<)

On S

Zaks w e)

mR < a | m

me Bl.

fe G & a op

oa}

om a] s a S| 3

: 2, Vat (Ss

fs = wo | &

g z\a| 31.

= Oo, re Ie

FRAUDATRIX GROUP

GP 1. minutissimus

oe @

GP 2. alorensis

GP 3. infantulus

GP 5. cinctellus

Burma

Ogg Cc a

GP? 6. fulleri

G 7. vubithoracis C)

eo @

GP 10. inculus

GP 15. reidi

GP 18, cubitatus

MAMMILIFER GROUP

TH 24, impostor

Seana

TE. eo

mac [ ax meme | | | |

EET | a monmiier [@| [O[@

29, wardi

aoe

eae

mC)

GP,RP

APPENDIX A: TABLE OF DISTRIBUTION AND BREEDING HABITATS (Continued)

EXTRA

DISTRIBUTION aa ies per ey

On ; es

Ze oO} s

< Sion

a ay & o

f AQ 5 ro

i 3 elale| | # | &

age 4/¢= Slele| aller eto) 5 |

@ Q, o oo tee bd

alelals BS slei el elé6|S16l sla! &

3/5/2/8 Se) Be ee ee ey

Al(Ml|aAle | RAR ea ea hae de ee ae

| tH | so demisus | | | |@ a pe

tnt | | | Tiel el er

sages ee Ne Aa <a ee

vz minst | | | | dT | | fe | fem | dT | | ele

pe Be | Se ermssicomes | || hh OM sales ee eok

RELAC_| 99 incomptus | | eee ae | ee ee

GP,RP, TH] 40, bengalensis | @ OO ee

41. peyton |@ | hide ee eee

| @ewerines | | | eae ae

ae Pewee

el | a oe a Pe er

peeiees Te ee

Pa eae eee | eee eee

Pet] Migs fT eee ea

eee | se mete || | ee ee

ai eee eo Pe eee eee

Be | ee ements || Oe

Bae ols qemie< 3 LTS) || ee

| pp | se cups | | | |@| | |@j@@e@ |) | |i | | |

|e | 65, swmaioms | | | | @@l | | je | je | tt

WILFREDI GROUP

GP 56, wilfredi

ae Reeser eee See

ee, ee eee see

242 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

APPENDIX B: CURRENT TAXONOMIC CHANGES

NEW TAXA

EEE CO) ch nn na Ghd eto a. WORE OA eR SO a Oba laos, 8 hee ee yk a eh He

CLOGS SS EEE ON GD MN, re ST SE OER Bnei Pret as aime | era eens Oo eae ae ee ee

SPOT OF IES 2% he et Kees Ob te Ree tah en) ete Ge ie odbice, Ue fan ml wes ne ee

FEE A id) gta, end in, Wade ee) ae Nae oe ae a a Se Ray gle Ge Nga aod Hp ciew eng ue Oe

PT DOS LOU VERA: Shee. Se) SE a ene eo Re Rd Se fo teed a Fe Pokus Sie ae wc weed oe

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LECTOTYPE SELECTION

POTEET) ty = as soon tees Uy RE 9 NSA Re Se b oA J) EA, eau km IO ee ng Se ee

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LOE IIE Sloe eb aoe ae AE Sa ae eae ee ee EE ee Ge ES a SET

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 243

INDEX

Names of valid taxa are set in roman type, synonyms and invalid names

are italicized. Italicized numerals refer to the principal text references,

roman numerals to secondary text reference, with suffix k indicating diagno-

Sis given ina key. Numerals enclosed in parentheses refer to figures.

Acalleomyia 8

aculeatus 15, 16k, 18k, 19k, 21k,

30, 48-49, 50-52, (16)

acutipalus 70k, 73k, 75k, 78k, 126,

129, 137-139, 140-141, (68)

aestivus 1, 9, 15, 16k, 18k, 19, 20,

30, 50-51, (17)

alorensis 1, 9, 15, 16k, 17k, 19,

20, 22,24, 25-26, 27, 26, 30,

(1)

alphus 15, 16k, 18k, 19k, 21k,

65-67, (27,28)

alphus complex 9, 15, 18, 21, 65

ampullaria (Nepenthes) 129, 130,

133, 138, 140, 141, 144

atvatulus 40

bandoengensis 69k, 71k, 75k, 76k,

79, 101, 104-195, (45)

barkerii 1, 4, 9, 45, 47

Barraudius 8

bengalensis 13, 69k, 72k, 74k, 76k,

10,09, °121,° 712-714, 116, 448,

129, (49)

bernardi 1, 9

bicalcarata (Nepenthes) 129

bicornuta 102

bicornutus 1, 9, 69k, 72k, 75k,

76k, 79, 101, 102-104, 105-107,

117, (44)

bolii 10

brevipalpus 70k, 73k, 75k, 78k,

126, 135-138, 139, 140, 141,

(65, 66)

brevipalpus complex 9, 70, 73,

126, 134, 135, 137, 139, 140,

141, 144

Brevipalpus Subgroup 9, 68, 69,

10,. 12, T, 15, TG W2e-12

132, 135

chiungchungensis 87, 89

cinctellus 15, 16k, 17k, 19k, 21k,

30, 33-35, 36, 37, 54, (4, 5)

cinctellus complex 9, 15, 17, 21,

30,30

coerulescens 70k, 73k, 75k, 78k, 125,

129-130, 131-133, (60)

crassicomus 69, 72k, 73, 75, 79,

110-111, 113, 114, 116, (47)

cubitatus 15, 16k, 19k, 20k, 21k,

Si, ta7 oe, on, OO, Wieros, tr)

Culex (subgenus) 8, 101, 145

Culiciomyia 8, 10, 22, 24

curtipalpis 8, 69k, 73k, 75k, 77k,

126, 133, 142-144, 145, 146,

(69, 70)

curtipalpis complex 9, 69k, 73k,

126, 142

Cyathomyia 4, 68, 133

demissus 1, 9, 69k, 71k, 75k, 77k,

79, 91-93, (37, 38)

digoelensis 10

eminentia 70k, 73k, 75k, 78k, 126,

137, 138, 139-149, 141, (67)

eukrines 69k, 72k, 77k, 79, 116-118

(52)

Eumelanomyia 8-10, 81, 146

fidelis 86

flavicornis 9, 69k, 70k, 74k, 76k,

78, 79, 120-123, 124, (55, 56)

flavicornis complex 9, 70, 78, 79,

120

fraudator 99

fraudatrix 4, 49, 53, 55, 56, 63

Fraudatrix Group 9-15, 22, 29, 37,

39, 53, 67, 68, 146, 149, 150

Fraudatrix Subgroup 9, 16, 17, 19,

20, 22, 26,29, 30, 32, 33, 33,

37, 39, 41, 42, 45, 47, 55, 61,

fulleri 15, 16k, 18k, 19, 20, 30, 35,

36-37, (6)

fuscosiphonis 1, 9, 91, 93

244 Contrib. Amer. Ent. Inst., vol. 13, no. 4, 1977

ganapathi 9, 69k, 71k, 74k, 77k,

79, 93-96, 98, 115, 116,

(39, 40)

- ganapathi complex 9, 78, 79, 93,

116

gibbulus 15, 16k, 18k, 20, 30, 39,

41-42, (11)

gracicornis 1, 9, 15, 16k, 19k, 20,

31, 58-59, 60, (22)

gracilis (Nepenthes) 133, 138, 139,

140, 144

harpagophallus 9

hewitti 70k, 72k, 75k, 77k, 125,

129, J31-133,,.145,- (61, 62)

hewitti complex 9, 70, 72, 125,

131

hirtipalpis 1, 9, 146, 147k, 150,

168 162) (73)

hut 1, 9, 96,.97, 98

impostor 1, 7-10, 69k, 70k, 77k,

79, 89-81, 82, (29)

impostor complex 9, 70, 74, 78-80

incomptus 69, 72k, 73, 75, 79,

£11-172, (48)

inculus 15, 16k, 17k, 20k, 30,

43-44, 53, 63, (12, 13)

inculus complex 9, 15, 17, 20, 30,

infantulus 11, 15, 16k, 17k, 19k,

20k, 22-25, 26-29, 35, 39,

(2, 3)

jenseni 4, 70k, 73k, 75k, 78k,

125, 126, 133-135, 142, (63, G4)

jenseni complex 9, 70, 73, 126,

133, 137

josephinae 15, 16k, 19k, 20k, 21k,

1, 46-57, (21)

juxtapallidiceps 22, 24

kuhnsi 1, 69k, 72k, 74k, 76k, 79,

85, 97, 98, 107-110, (46)

kusaiensis 8

lasiopalpis 1, 9, 69k, 70k, 74k,

Tok, 10, 72, 1232124, (57)

lavatae 69k, 71k, 75k, 77k, 79, 84,

86-87, (32)

Lophoceraomyia 4-12

Lophoceratomyia 4, 15, 31, 33,

34, 36, 37, 40, 44-46, 49, 51,

03, 06, 57, 84, 87, 100, 102,

104-112, 120, 129, 129,131,

135, 139, 142, 150

lucaris 70k, 73k, 75k, 78k, 126, 137,

138-139, 140, 141, (67)

Lutzia 8

macdonaldi 15, 16k, 19k, 20k, 21k,

31, 61-63, 64, (23, 24)

mammilifer 13, 69k, 71k, 74k, 76k,

79, 87-99, 91-93, 102, 111-114,

118,.120, 121, 123, (34,.35)

mammilifer complex 9, 71, 78, 81,

87, 93, 120

Mammilifer Group 4, 9-13, 22,

67-69, 80, 81, 125, 131, 136,

146, 149, 150

Mammilifer Subgroup 9, 68-70, 73,

75, 76, 78-79, 80, 83, 85-90, 92,

94-97, 101, 108, 115, 116, 120-

123, 125, 127-129, .135,.150

Melanoconion 22, 24

mindanaoensis 1, 9, 109

minor f, 9, 69k;. (2k,. Tok, .76k, 19,

96, 98-102, 103-108, 111-114,

117, 121, (43, 44)

minor complex 9, 71, .72,. 76,79;

95, 98, 106, 110-113, 116, 125,

129

minutissima 22, 24

minutissimus 15, 16k, 17k, 19k, 20k,

22-25, 26-28, 35, (1)

Minutissimus Subgroup 9, 15-17,

19-22, 26

mirabilis (Nepenthes) 144

navalis 70k, 73k, 75k, 78k, 125,

127-129, 130-133, (58, 59)

navalis complex 9, 70, 72, 125-127

133, 135

Neoculex 125, 145

Nepenthes 129, 130, 133-135, 138-

141, 144

niger 15, 16k, 18k, 20k, 21k, 30,

39, 40-41, 42, (9, 10)

niggerima 22

nolledoi 98, 100

Oculiomyia 36

pachecoi 1, 9, 44, 46, 47

pairoji 1, 8, 9, 15, 16k, 19k, 20k,

21k, 31, 63-65, (25, 26)

Sirivanakarn: Subgenus Lophoceraomyia in the Oriental Region 245

pallidothorax (Culiciomyia) 123

paraculeatus 1, 9, 15,16k, 18k, 19,

20, 30, 49-50, 51, (16)

pavainfantulus 26, 27

Passerita 11 ,

peytoni 69k, 72k, 74k, 77k, 79, 111

114-116, 11%, 420, (50, 3)

peytoni complex 9, 72, 78, 79, 114,

120

philodendromyia 4

pholeter 13, 69k, 71k, 73k, 74k, 76k,

18, 19, 118-120, (58, 34)

pholeter complex 9, 78, 79, 118

pilifemoralis 146, 147k, 150-152,

(73)

plantaginis 1, 9, 98, 100-102

prasina (Passerita) 11

pseudorubithoracis complex 39

quadripalpis 1, 9, 15, 16k, 18k,

19k, 21k, 30, 44-47, 48, 51, 52,

63, (14, 15)

quadripalpis complex 9, 15, 16, 18,

20, 30, 44, 47, 49, 53, 65

raffleziana (Nepenthes) 139

raghavanii 9, 79, 123, 124

reid) 13, 15, 16k. 16k, 20k. 30,

44, 51-53, (18)

roubaudi 44, 46, 47

rubinotus-rima group 10

rubithoracis 15, 16k, 18k, 20k,

21k, 30, 37-40, 41, 42, 47,

(7, 8)

rubithoracis complex 9, 15, 18, 30,

37, 39, 40, 42, 44, 65, 66

seniori 15, 16, ‘17k, 19, 20, 30,

21-92; 35,. 35, (6)

seniori complex 9, 15, 17, 30, 31

spiculosus 1, 9, 69k, 71k, 74k,

76k, 79, 85, 86, 96-98, 102,

108, (41, 42)

stonei (Uranotaenia) 107

sumatranus 7, 8, 10, 69k, 73k, 75k,

77k, 125, 126, 144, 145-146,

(70)

sylvestris 44, 46, 47

szemaonensis 9, 95

taeniata 33, 34

traubi 69k, 71k, 74k, 77k, 79, 81,

82-84, 85, 86, 87, 91-93, 95-98,

(30, 31)

traubi complex 9, 71, 78, 79, 81,

tuberis 69k, 72k, 74k, 77k, 79,

105-107, 112, (45)

uniformis 4, 69k, 71k, 75k, 76k,

78, 79, 84-86, 91, 97, 98, 107,

108, 123, (32, 33)

uniformis mercedesae 1, 85, 107,

109

variata 53

variatus 15, 16k, 19k, 20k, 21k, 31

53-56, 57-63, (19, 20)

variatus complex 9, 15, 31, 53, 55,

58, 09, G1, 62, 66

villosa (Nepenthes) 135

-wardi 1, 9, 13, 69k, 71k, 74k, 76k,

79, 90-91, (36)

whartoni 15, 16k, 19k, 20k, 21k, 31

59-61, 638, 67, (22)

whitei (subgenus Culex) 101

wilfredi 9, 68, 146, 147k, 147-150,

131. Loe ATI, 2, 78)

Wilfredi Group 9, 12, 13, 68, 146,

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