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Contributions

of the

American Entomological Institute.

No.

as

Volume 11, 1974-1976

CONTENTS

Reinert, John F. Medical Entomology Studies - 1.

A new interpretation of the subgenus Verrallina of

the genus Aedes (Diptera: Culicidae). 249 pages.

December 12, 1974.

Morse, John C. A phylogeny and revision of the

caddisfly genus Ceraclea (Trichoptera: Leptoceridae).

97 pages. April 17, 1975.

Young, Daniel K. A revision of the family Pyrochroidae

(Coleoptera: Heteromera) for North America based on

the larvae, pupae, and adults. 39 pages. Nov. 24, 1975.

Cook, David R. Contributions to the water mite fauna

of North America. May 28, 1976.

New North American species of the genus Arrenurus,

mostly from Florida (Acarina: Arrenuridae)

Pages 1-56.

North American species of the genus Koenikea

(Acarina: Unionicolidae). Pages 59-100.

North American species of the genus Mideopsis

(Acarina: Mideopsidae). Pages 101-148.

This is an irregular appearing series,

published by the

AMERICAN ENTOMOLOGICAL INSTITUTE

5950 Warren Road

Ann Arbor, Michigan 48105, USA

Volumes 1-11 available in cloth-bound volumes

Contributions

of the

American Entomological Institute

Volume 11, Number 1, 1974

MEDICAL ENTOMOLOGY STUDIES - I.

A NEW INTERPRETATION OF THE SUBGENUS

VERRALLINA OF THE GENUS AEDES

(DIPTERA: CULICIDAE).

by

John F. Reinert

il

CONTENTS

ABSTRACT. 2.0.2 ee ee a RN ee 1

INTRODUCTION on 6 6 oe a ee. ee ea eS 1

GENUS AEDES, SUBGENUS VERRALEINA (ca ke ee eee ee. 3

KEYS TO SECTIONS AND SERIES OF VERRALLINA, ..... 16 1 ete ee 11

DESCRIPTION OF SHCTIONA®. Jy... A ek bee ee ee nee 13

OIE | a a a a oe ee 14

SeRIRSd 25. oo Me a ies. 14

Ollie 6 a a a ee ee ae 15

DESC Ri TION OF SECTION Be ee a a ws 15

LIST OF SPECIES IN SECTIONS AND SERIES... ..3..... So ow ee 16

SPRCINS TREA MENT 2 2 ee ee 18

WbGis barradG 6. 6 se a ae a ee 18

WdUSfs EAMOON fe a ee ee ae a wk 18

G2VeStiS BArraue: ©. Sue eS eee a 20

andamanensis Edwards .......2+.e+.-s oe ee 20

ONAGIIGIICHS1S GIOUD. 6 ie a a ee re as a1

OEVIUS Barvaud., 6 ke eek ae a ee ea 22

Diplo? PNCCUA 0 a ea a a es 22

CGT Py OSV IGS TATIOON: 6c a a ee ee 28

Cie) BOWwarOs 6h a Re ee 28

Beis tear Pau: 8 a a a a ee ae 34

Cie Ss ert ouG a a a RO i a a ee es 34

comatus Barraud ..... a a a 30

CONMLOSUS NEW SNCCICS 6 6 6 a ve ke ee eee ws 36

CONSON esis ROME. 2 a ee a re a 38

PVCs DerinadG, 2 6 ae a a a ew 38

CVPICIIOts OW OTUs 9 ee ks ne i ee 39

Ou Dyae aid ShannOh |. 6 a2. oe a ee ek rs.

PUI ae A a a a ke we 42

TO EES oi ee 43

gibbosus Delfinado. .... de ee AA,

Wangieryls AGO 2 ne a ee ee ee eee 49

harrisonicus new species.......4..2-eee- ae a 46

PHOCV INS FON VOG a a a lg ee 48

MAC COVAOTITS (LLOICOStEN)? op ee Pe AI ae ee 52

indicus (INECOMAIG) Wk a Ge ew ee 53

FORIISONT OttOONs 6 8 ae ee a a ew a 08

GOROVCHSTS MEW SPECICS 6 ee re ee ee as Oe ee 08

WINROCHSTS SLONG aNG MNIGNE so a a ee ee as 61

IGPU DeHIIS DOU WIAGO 2. ee ee ee ee a er ee oe es 62

beveestey? BUWaras 5 4250 ee ee or a a ee 63

LUOUOTIS TRACY AUG i eh se He eae ae oe ee es 66

macrodixoa Dyar AiG BHATNON. ff oe we ee es ee ee 67

ili

CONTENTS

margarsen Dyar and: Shannon. . . 44444 6 eee a ee: 67

neomacrodixod Kine and Hoogsiraal . -. 6 a. 68

NIGVOIAY SiS (LMGIOW <6 a5 eo ke i eee; 68

NOOUROMIS Yamada <P oe ee ee: 69

notabilis: Delfinado: <2 eco ee eee. i 70

pahans? Delginado., 3. 6 ee ee ee ee a

panayens?s LUCIOW (a. we a a ee ew ee ee ee 71

petroélephanius Wijesundara. , 22>... FO a a, le

philipbinensis Deliinad® ...44..4 a. we Pee Se ee. 73

pPrnomuUs WIG 6k se a re ee Pe se. ve

PIpRiNG BORAT. soe 6 as ies ee eS 73

PYIOEHONCNSIS BYUS a ek ne A ee OR. ME ging e 74

pseudomediojasciatus (Theobald), ... .. .. 38 1s ee. 15

Ppseudovavicids Wew SPECiGS . 2. ee a ee, 76

VOMOLN SOIL DOW SPCCIOS. oo. ck sme 6 oe ate ee es ae oe a. 78

VOIERATIAUO oe coe ee sw ae 80

VOVUS ONIN so ea ae ete a ee we 81

SHUQVENTIS NEW SPeCiGS. . ek ks we oe ee, ee 82

SCCWlais WeNnON 5. 4k a a ee ee 84

SOMME MEW SPCCICS -¢ e a a a a ee a 85

LOYVOSUS DEMINACO. . nee aha os le ew ee 87

WHOS IVCOOAI) 88

DOCS Ss BOYER oe a ee ee 91

VaVields (belcester): 2 0 ee. 92

OIVIES (LEIGERION)) oe a a a 92

NOVOUIT DOWALUS 66. 6s foe a ek a ee ce 95

WUSGT, PACTAUG ove 6 cues 1 6 uke ee ee a ee 96

BORTOW LEDGMENTS <6. a a a ee re ee. 101

LITE RA FORE CUT BD 6 cg wae er a a ee ee 101

LAST OF FIGURES os ee is ae i a ee ee ee a ee 106

Ligt.OF FIGURE ABBREVIATIONS oo. 5. 2c6. ook oe we a 108

PUR ES oa a bad abeck a a ak, a es ea 109

BPP EN ES 5 ek ak aie ave ls ges ee a ke ee ee ae 175

TABLE 1. Current taxonomic status of the subgenus Verrallina ... 177

TABLES 2-14. Records of the branching of the setae on the pupae

of Aedes (Vervaliimg) SPECIES 6 6 ook oc a A ee ae 181

TARE 2. UGH ek as a ee ae 181

TABLE 3. DOOY oa ea a es a a 183

TABLE 4. COVIICIIEE er ee a eat ee 185

TABLE 5. CHEV OO ig ee ee a ie Be 187

TABLE ©. O1000SHS (Co. ee Ge sa ol a ee 189

TABLE 7. weCeris OO ee es ee goa a TR er ae a et a 191

TABLE 8. WiGIEUR hE ae a we ee ee 193

TABLE 9 LAPIROWNIO Gee a WO ae aes 195

TABLE LOD teveestert. eas Re i eS rs 197

1V

CONTENTS

TABLE 21, BOUOIES, i uke eee OO OS SR SEG. 199

TABUE LZ. . HOvOSUS Js wo. oe SER a SR a Coes, 201

Tee 1s... WIEads Sn ee a ee a ER MEIN, 203

TABI 94, GO a ew eo ee wl ek BN BO 205

TABLES 15-27. Records of the branching of the setae on the larvae

OF Actes (Verve iiiie) SROCIES, oo eae eo we CONS es. 207

ee ae a etek ee ee a AP 207

TAS UG. eer oe Bee ee oe RN, CS 210

Tee tt Cee Sk eo ww kw oe vn SR, EP. 213

TA 2G). CVPIOIIUIS 6 Ge Gy an ew 8 a oe SHEA ORO, 216

Ec ep Oe oe ae tk ee cage ew oe SS, SS 219

es 8s aw te ce oe ei ee ER SP 222

ag RIG oa in ee ol te a SPOT. 225

Te. Be ee a we we CS BS SHOT oN 4. 228

i, FO oh an ts so ne ce WR OAR PO A 231

ae a OOS oa ek a ae wk ew ao So TERT Cie 234

a ee a lg a i oe ee ein co ce ow CRE UL RE 237

TAS 2h. Ie on ee we es ke PRS GIRS PAPO RE. 240

a ak ek vik teste ce ce EN Ww ew we OE CG. 243

CONSPECTUS OF TAXONOMIC CHANGES, 4.04. 46 28S ee Be. 246

MEDICAL ENTOMOLOGY STUDIES - I.

A NEW INTERPRETATION OF THE SUBGENUS

VERRALLINA OF THE GENUS AEDES

(DIPTERA: CULICIDAE)!.

By

John F. Reinert

‘ ABSTRACT

The subgenus Vervallina Theobald is redescribed and compared to related

subgenera of Aedes Meigen. A new interpretation of the subgenus is given and

is based primarily on the unusual development of the male and female genitalia.

Neomacleaya Theobald is reduced to a synonym of Verrallina, New informa-

tion on biology, distribution, descriptions and corrections for a number of

species is presented. Seven new species, comosus, harrisonicus, johorensis,

pseudovarietas, ramalingami, sabahensis and sohni, are described and illus-

trated. A number of stages and structures are described and illustrated for

the first time. New synonyms in this paper are: butleri Theobald (=female

only of seculatus Menon); incertus Edwards (= dermajoensis Brug); carmenti

Edwards (= cevamensis Brug); nobukonis Yamada (= ishigakiensis Bohart);

varus Delfinado (= khmerus Klein); uncus (Theobald) (= hirsutipleura Barraud)

and yusafi Barraud (= sSiamensis Delfinado). Aedes adustus Laffoon is resur-

rected from synonymy. Lectotypes for cartevi Wijesundara, cevamensis Brug,

hirsutipleuva Barraud, mediofasciatus Theobald and petroelephantus Wijesun-

dara are designated.

INTRODUCTION

There has been considerable confusion as to the proper taxonomic status

of Aedes Meigen, Neomacleaya Theobald and Vervallina Theobald and their

included species. Verrallina was originally described by Theobald (1903: 295)

as a genus with 3 included species of which only one, butlevi Theobald, remains

in this taxon, the other 2 species have been transferred to other genera (nigra

Theobald to Ficalbia Theobald; smithii Coquillett to Wyeomyia Theobald).

Theobald later (1907: 238) described a new genus, Neomacleaya, for indica, a

new species. In 1908 Leicester described his new genus, Aioretomyia, and

IThis work was supported in part by Research Contracts DA-49-193-MD-2672

and 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.

2uajor, 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. 11, no. 1, 1974

placed 5 new species in it.

Edwards (1913: 229) synonymized the genera Aioretomyia, Neomacleaya

and Verrallina with the genus Aedes Meigen. He later (1917: 221) placed the

species included in the above 3 genera and several additional species in the

subgenus Aedes Meigen of the genus Aedes, All subsequent authors followed

Edwards' 1917 arrangement of the subgenus Aedes until 1962. Some of the

more notable articles were Barraud (1928, 1931, 1934), Edwards (1922a,

1922b, 1924, 1932), Dyar and Shannon (1925), Laffoon (1946), King and Hoog-

straal (1947), Wijesundara (1951), Knight and Hull (1951, 1953) and Stone,

Knight and Starcke (1959).

Belkin (1962: 412) resurrected Vervallina as a distinct subgenus of Aedes

and provisionally included Neomacleaya and Aioretomyia as synonyms. He

stated, and I concur, that the South Pacific species of Verrallina had little in

common with the type species (cinereus Meigen) of the subgenus Aedes,

Delfinado (1967) resurrected Neomacleaya as a subgenus containing primarily

Oriental species previously included in Aedes, Aioretomyia, Neomacleaya and

Vervallina; however, she mentioned Neomacleaya was closely related to both

Verrallina and Aedes sensu stricto. Huang (1968) followed Belkin's 1962 publi-

cation and assigned the species from New Guinea to the subgenus Vervallina

(including neomacrodixoa King and Hoogstraal and panayensis Ludlow). Thus

in 1968 Delfinado (30 April) and Huang (10 July), in independent work, each

determined the species neomacrodixoa and panayensis as belonging to the sub-

genus each was working with. This example demonstrates that the 2 subgenera

were poorly defined. Klein (1973) followed Delfinado's 1967 and 1968 works in

treating the Cambodian species of Neomacleaya.

A new interpretation and description of the subgenus Vervallina is presented

here. The subgenus, by this new concept, is separated into Sections A and B

and Section A divided into Series I, Il and III. This new definition of the sub-

genus resulted primarily from a study of the subgenera of the genus Aedes.

During this study it became apparent that there were a number of inconsis-

tencies, incorrect indentifications and synonymous species in earlier publica-

tions dealing with the subgenus Neomacleaya. The evolution of the present

interpretation of the Verrvallina resulted from the examination of the types of

most of the included species and specimens of all except 2 species. Material

of cunninghami Taylor and spermathecus Wijesundara was not available for

examination. All specimens and types in the United States National Museum

(USNM) including the Southeast Asia Mosquito Project (SEAMP) and Medical

Entomology Project (MEP) collections, many specimens and most types in the

British Museum (Natural History) (BMNH) and numerous specimens and types

from other museums and individuals were examined by me during this study.

Little new information on species in Series I of Section A can be added

here to the presentations given by King and Hoogstraal (1947), Belkin (1962)

and Huang (1968). However, considerable new information on synonyms,

biology, distribution, previously undescribed stages, corrections to published

work and several new species are added to Series II and III of Section A and

Section B. Unfortunately the adults of a number of species of these sections

and series are very similar in habitus and can not be separated with certainty

except by genitalia preparations. Other factors hampering the identification

of species are: lack of undamaged and sufficient numbers of specimens

(several species are represented by only the type or a very few specimens

which are badly rubbed, have vital parts broken off and missing, or repre-

sented now only by a genitalia mount, the adult body being lost); original and

subsequent descriptions and illustrations of a number of species are not suf -

Reinert: New Interpretation of Vervallina 3

ficient to separate them from other morphologically similar species; associa-

tion of sexes of several species is only presumptive and needs confirmation;

some species show adult sexual dimorphism; and the immatures of a signifi-

cant number of species remain to be found. The current taxonomic knowledge

of the subgenus Vervallina is summarized in Table 1. Many of the taxonomic

problems within the Vervallina such as species group classification, species

synonymy, correct association of sexes and knowledge of immatures can be

solved only when progeny rearings become available.

Abbreviations used in references to literature conform to the BIOSIS List

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

phia, 1972. In the synonymy sections, an asterisk following the abbreviations

used (2 =female, “ = male, P = pupa, L = larva) indicates that at least some

portion of that sex is figured. The abbreviations used in distribution sections

are the same as in the synonymy but with the following additions, p = pupal

skin, 1 = larval skin and E = egg. The number of specimens used in determin-

ing range and mode seta branching in pupal and larval descriptions and tables

is included in parentheses after the table headings. In pupal descriptions and

tables the number of branches on abdominal seta 1-I was measured on the basal

third of the seta. A dash (-) on pupal and larval tables indicates the seta was

broken off and missing from all specimens examined or the seta showed no

mode. 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 prov-

ince follows the last place name of the province in the distribution sections.

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

Official Standard Names Gazetteers prepared by the Geographic Names Divi-

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

Department of the Interior: Cambodia (1971), Indonesia (1968), Malaysia

(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

collection sheets and specimen labels of the specimens that I examined. Much

of this information is reported here for the first time.

Nomenclature and chaetotaxy used for females, males, male genitalia,

pupae and larvae 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 VERRALLINA THEOBALD 1903

Logotype: Aedes butleri Theobald

Neomacleaya Theobald 1907, Monogr. Culicidae 4: 238. Haplotype: Neo-

macleaya indica Theobald. NEW SYNONYM.

Aioretomyia Leicester 1908, Culicidae Malaya p. 185. Logotype: Azoreto-

myia varietas Leicester.

The following description of the subgenus is based on specimens of 91

4 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

species examined and the published descriptions of the remaining 2 species not

seen. |

FEMALE. Head. Antenna dark brown, 0.91-1.42 length of proboscis,

pedicel brown with a few short fine hairs and usually a few small scales mesally,

flagellomere 1 with basal area pale and with several small dark scales; clypeus

brown, bare; maxillary palpus dark brown scaled, 4 segmented, 0.11-0.21

length of proboscis; proboscis dark brown scaled, 0.84-1.55 length of femur J;

eyes contiguous (Section A) or separated in front (Section B and mccormicki

Belkin); ocular setae dark and well developed; scales on head broad and decum-

bent except for a few narrow curved pale ones along coronal suture (most species

of Series III of Section A) and a few semi-erect forked scales near ocular line in

Section B, a few narrow scales on ocular line; occiput with several to numerous

erect forked scales, decumbent scales broad and/or narrow. Thorax. Scutal

integument usually dark; scutum covered with narrow curved dark scales, some

species also with narrow curved pale scales forming small patches usually on

following areas: median anterior promontory, anterior and lateral scutal fos-

sal, supra-alar and occasionally on margins of prescutelar space; scutellum

with a patch of narrow curved scales on each lobe; prescutellar space bare, a

few species with a narrow median cephalic stripe of scales; dark setae on the

following areas: 2-10 median anterior promontory, numerous anterior acro-

stichal, several to numerous posterior acrostichal, numerous dorsocentral

(anterior and posterior), scutal fossal (3-9 anterior, 2-8 lateral and usually

1-2 posterior), numerous supra-alar, 4-8 posterior medial scutal, 1 postalar

callar and scutellar (several long and short ones on both lateral and median

lobes); pleural integument brown, usually dark; antepronota widely separated,

broad scales present or absent, 7-28 short and long setae; postpronotum with

narrow curved scales dorsally (a few species with these scales apparently ab-

sent) in Section A, or scales absent in Section B, 3-10 posterior dark setae,

some species also with a few short fine hairs; propleuron with a patch of broad

white scales, 6-21 setae; prosternum bare except for adustus Laffoon, anda -

manensis Edwards, margarsen Dyar and Shannon and sohni new species, which

have a small patch of short fine setae on median dorsal area; paratergite bare except

indicus which has afew narrow pale scales; subspiracular area and mesomeron bare;

postspiracular area with or without scales, 2-12 setae; mesepisternum with an

upper and a lower patch of broad scales, 2-7 upper and 6-19 posterior setae,

some species also with a few short fine hairs on anterior, dorsal and ventral

areas; prealar knob usually without scales, 4-22 setae; mesepimeron with a

patch of broad white scales near center, 5-33 setae dorsad of scales patch,

Section A with a few (2-3) to numerous fine golden hairs caudad of scale patch,

hairs in some species restricted to lower posterior margin of scale patch while

others have numerous hairs along entire posterior margin of scale patch and

extend to near ventral margin of sclerite; metameron bare, however, short

fine hairs and small pale scales are present in gibbosus Delfinado and members of

the andamanensis Group(see discussion of adustus). Legs, Coxae I-III each with

several setae and a patch of pale scales, I also with brown scales; trochanters

I-III each with a few short setae and small scales; femora I and II with anterior

surfaces brown scaled, II occasionally with a pale scaled ventral stripe, I with

posterior surface brown scaled with a pale dorsal longitudinal stripe, II with

posterior surface with a pale ventral longitudinal stripe, III with anterior and

posterior surfaces pale scaled except for an anterodorsal and posterodorsal

longitudinal brown scaled stripe; tibiae I-III brown scaled, I and II occasionally

with a longitudinal pale scaled stripe on posterior surfaces; tarsi I-III brown

scaled; posttarsi I-III with ungues variable, III] with ungues simple in Section A

Reinert: New Interpretation of Vervallina 9)

and toothed in Section B. Wing. Dorsal and ventral veins with brown scales;

alula with scales on margin, scales narrow to moderately broad; upper calyp-

ter with several setae on margin; 1-3 remigial setae. Halter. Pedicel pale;

capitellum covered with pale and/or dark scales. Abdomen. Tergal scaling

variable, all dark scaled or with pale scaled patches and bands, usually with

a laterobasal pale scaled patch, laterotergite of I with a patch of broad pale

scales; sternal scaling variable, from completely pale to dark scaled, usually

with pale scaled basal bands and apical areas dark scaled; terga and sterna

with numerous setae, mostly along posterior margins. Genitalia. Tergum VIII

moderately pigmented, wide, base usually slightly concave, apex gently rounded

or slightly concave, numerous short and moderately long setae scattered over

apical 0.28-0.88, basolateral seta present or absent, some species with a

small bulla in this location, covered with minute spicules, numerous broad

scales covering apical 0.50-0.84, basal 0.1-0.7 retracted into segment VII,

VIlI-Te index 0.35-0.79, VIII-Te/IX-Te index 1. 67-6. 82; sternum VIII moder-

ately pigmented, wide, base concave mesally, apex with a shallow to moder-

ately deep median indentation with a small lobe on each side of midline, setae

on lobe thin (Section A) or stout (Section B), numerous short and moderately

long setae scattered over most of surface, covered with minute spicules,

numerous broad scales covering much of apical 0. 80-0.96, apical interseg-

mental fold unpigmented, VIII-S index 0. 52-0. 80; tergum IX moderately to

heavily pigmented, wide, short and usually ribbon-like, setae absent (several

setae present in gibbosus and yusafi Barraud), covered with minute spicules,

IX-Te index 0.12-0.49; insula small, unpigmented, with 3-7 small tuberculi

(usually each with a minute spicule), or insula absent (Section B) and replaced

by median apical portion of lower vaginal sclerite; lower vaginal lip moderately

to heavily pigmented, narrow, with or without spicules, with a small caudally

projected heavily pigmented median apodeme, lower vaginal sclerite very

lightly to heavily pigmented, variable in shape and number, with or without

spicules; upper vaginal lip moderately to heavily pigmented, variable in shape

and development, may have median posterior area developed into a vertical

shield (Section A, Series III) or a horizontal shield (Section B), with or without

spicules, upper vaginal sclerite moderately to heavily pigmented, variable in

shape and development, very large, often nearly covering roof of vagina;

spermathecal eminence heavily pigmented, shallow to deep in depth, simple

to complex, with or without spiny excrescences, without (Section A) or with

(Section B) a heavily pigmented spine-lined pouch; postgenital lobe short, narrow

to moderately broad, apex rounded or with a small to moderately deep median

indentation, 5-29 setae on each side of midline, covered with small spicules,

dorsal PGL index 0.47-1.29, ventral PGL index 0.70-1.96; peri-anal mem-

brane with minute spicules; cercus triangular in shape, moderately long,

broad at base, apex acute, covered with minute spicules, dorsal surface with

a number of short and moderately long setae scattered over most of the sur-

face, a few long ones at apex, ventral surface with a number of short and

moderately long setae along outer margin becoming more numerous distally,

none to several broad scales on dorsal surface, cercus index 1. 76-3. 86,

cercus/dorsal PGL index 2.42-4.63; 3 seminal capsules, 1 large, 1 medium

and 1 small in size, heavily pigmented, elliptical or ovoid, with a narrow or

broad neck, several small seminal capsule pores near orifice, base of acces-

sory gland duct unpigmented to moderately pigmented.

MALE. Usually similar to female in general habitus but with the following

differences. Head, Antenna plumose, 0.76-1.27 length of proboscis; maxillary

palpus short, 0.09-0.18 length of proboscis; proboscis 1. 01-1. 56 length of fe-

6 Contrib. Amer. Ent. Inst., vol. 11,.no. 1, 1974

murI. Thorax. Most setal groups usually with fewer setae. Legs. Posttarsi

variable, III with ungues simple in Section A (except prioekaensis Brug which

has one of the ungues with a small tooth) and ungues toothed in Section B. Abdo-

men. Terga with lateral setae rather short; tergum VII wide with a number of

long caudally directed setae. Genitalia. Tergum IX moderately to heavily pig-

mented, mesal area narrow, setae absent; gonocoxite short, broad, heavily

pigmented, mesal surface membranous with minute spicules, dorsal, lateral

and ventral surfaces with short to long setae, many species with dorsoapical

and/or ventroapical lobes or projections, some species with spiniforms sub-

apically on sternomesal area, few to numerous scales on lateral and ventral

areas; gonostylus short to moderately long, moderately to heavily pigmented,

variable in shape but usually with basal portion somewhat expanded and with a

few setae, apical portion narrow and curved, gonostylar claw absent; basal me-

gal lobe variable in development, consists of a plate situated on membranous

basal mesal area of gonocoxite and is strongly fused along sternal margin to

tergomesal margin of scleritized ventral surface of gonocoxite, plate connected

with its mate by a narrow band covered with spicules and located ventrad of

apical portion of prosophallus, with several to numerous short to moderately

long thin setae, some groups also with lobes, projections or spiniforms; proc-

tiger with paraproct variously developed, many species with paraproct modified

into short or long, heavily pigmented, free arms, other species with paraproct

developed into broad contiguous heavily pigmented plates, sternobasal area pro-

duced into a small lobe which extends ventrad and articulates at a point on the

dorsal surface of the parameral apodeme near base of opisthophallus, base of

proctiger fused with tergum X, cercal setae absent; tergum X moderately to

heavily pigmented, fused with tergum IX; phallosome complex, opisthophallus--

consists of a lightly to heavily pigmented tergal transverse bridge between

parameral apodemes, located dorsad of phallus and prosophallus and ventrad

of proctiger, phallus--consists of a pair of heavily pigmented, narrowly sep-

arated, caudally produced arms which are fused together by a narrow band,

base extends laterally and fused to basal portion of prosophallus, apex of

phallus formed into a tergal penis filament which extends cephalad, prosophal-

lus--consists of a pair of moderately to heavily pigmented arms or plates with

bases widely separated, base attached to paramere, located laterad of phallus,

paramere--heavily pigmented, short and small (Section A) or very long and

moderately broad (Section B), parameral apodeme--moderately to heavily pig-

mented, short to long, outer surface broadly attached to tergobasal apodeme of

gonocoxite; sternum IX heavily pigmented throughout, large, broad, several

short setae in a patch near median caudal area, most species of Section B

and several species of Series III of Section A also with a few broad scales.

PUPA. Chaetotaxy and other features are summarized as follows. Cephalo-

thorax. Setae 1-3-CT approximately equally developed; 1-CT single to 9)

branched; 2-CT single to 4 branched; 3, 6-CT single to triple; 4-CT single to 9

branched; 5-CT double to 12 branched; 7-CT double to 9 branched; 8-CT double

to 11 branched; 9-CT single to 6 branched. Metanotal plate, Seta 10-CT double

to 13 branched; 11-CT single; 12-CT double to 8 branched. Respiratory trum-

pet. Moderately to heavily pigmented; hair-like spicules on inner surface of

meatus; index 3.00-5.67. Abdomen. Setae 0-II, VII-VIIl single; 0-III-VI

single or double; 1-I dendritic, well developed with 10-38 branches on basal

third; 1-II triple to 37 branched; 1-II double to 14 branched; 1-IV-V single to

10 branched; 1-VI double to 8 branched; 1-VII single to 6 branched; 2-II-VUl

stout in many species; 2-I single or double; 2-0-Vi single; 3-I double to 8

branched; 3-II single or double; 3-III single; 3-IV double to 10 branched; 3-V

Reinert: New Interpretation of Vervallina 1

single to 7 branched; 3-VI single to 6 branched; 3-VII double to 9 branched;

4-I triple to 12 branched; 4-II double to 11 branched; 4-III single to 8 branched;

4-IV double to 5 branched; 4-V triple to 11 branched; 4-VI double to 12

branched; 4-VII double to 7 branched; 4-VIII single to 5 branched; 5-I-III double

to 9 branched; 5-IV-V single to triple; 5-VI single to 4 branched; 5-VII single

to 6 branched; 6-I single to triple; 6-II single or double; 6-III-V single to 10

branched; 6-VI single to 4 branched; 6-VII double to 8 branched; 7-I, IV single

to 6 branched; 7-II single to 7 branched; 7-III single to 6 branched; 7-V single

to 11 branched; 7-VI-VII single to triple; 8-III, VII single to 6 branched; 8-IV

single to 5 branched; 8-V single to 7 branched; 8-VI double to 5 branched; 9-I

single to triple; 9-II-VI single; 9-VII single to 4 branched, weakly developed;

9-VIII single to 6 branched; 10-III single to 10 branched; 10-IV single to 7

branched; 10-V, VII single to 4 branched; 10-VI single or double; 11-III single

or double; 11-IV single; 11-VI single or double; 11-VI single to 4 branched;

14-III-VI single; 14-VII-VIII single or double. Paddle. Ovoid; minute serra-

tions on basal area of outer margin; minute spicules on apical portions of outer

and inner margins; midrib lightly to moderately pigmented and nearly reaches

or reaches apex; 1-P usually single, occasionally double; index 1.14-1.70.

LARVA. Chaetotaxy and other features summarized as follows. Head.

Moderately pigmented; setae 0, 1, 3, 18-C single; 4-C double to 10 branched;

5-C single to 9 branched; 6-C single to 7 branched; 7-C double to 15 branched;

8, 13-C single to 6 branched; 9-C double to 9 branched; 10, 11-C double to 8

branched; 12-C single to 10 branched; 14-C single or double; 15-C with 4-11

branches; 6-MP single or double; 1-C well developed, long and thickened; 4-C

small and weakly developed; 5-7-C long, well developed and barbed; 4-C

slightly cephalad of 6-C and mesad of 5, 6-C; 6-C laterad and well cephalad of

o-C; 7-C cephalad and well laterad of 6-C; dorsomentum heavily pigmented,

with 17-45 teeth; mouth brush with mesal hairs pectinate apically. Antenna.

Short to moderately long, usually moderately pigmented, scattered spicules

over most of shaft, usually more numerous on basal area and longer on outer

area near middle; seta 1-A short to long, attached near middle of shaft, and

branches more or less flattened; 2-A long, with a subapical constriction; 3-A

short, 0.20-0.36 length of 2-A; 4-A moderately long, 0.37-0. 74 length of 2-A;

5-A with basal portion flattened, pigmented and with a dorsal spine (see Fig.

02), apical portion unpigmented; 6-A peg-like, short, 0.22-0.39 length of 2-A.

Thorax. Seta 0-P with 4-23 branches; 1, 7, 14-P single to triple; 2, 3-P

single to 5 branched; 4-P single to 9 branched; 5, 6-P single; 8, 10, 12-P

single or double; 9-P single to 7 branched; 1-M double to 8 branched; 2-M

double to 7 branched; 3-M single to 4 branched; 4-M triple to 13 branched;

o-M single or double; 6-M triple to 9 branched; 7, 10, 12-M single; 8-M with

4-11 branches; 9-M with 6-12 branches; 11-M single to triple; 13-M with

7-40 branches; 14-M with 4-20 branches; 1-T single to 7 branched; 2, 7-T

with 4-10 branches; 3-T with 4-21 branches; 4-T double to 11 branched; 5-T

single or double; 6-T single to 6 branched; 8-T with 7-27 branches; 9-T double

to 4 branched; 10-T single; 11-T single to 4 branched; 12-T single to triple;

13-T with 6-23 branches. Abdomen. Setae O-II-VIII single; 1-I single to

triple; 1-II single to 5 branched; 1-III, VII double to 13 branched; 1-IV triple

to 15 branched; 1-V triple to 13 branched; 1-VI double to 11 branched; 1-VIII

double to 10 branched; 2-I-VII single; 2-VIII double to 5 branched; 3-I with

4-13 branches; 3-II triple to 16 branched; 3-III double to 13 branched; 3-IV

triple to 12 branched; 3-V double to 8 branched; 3-VI double to 10 branched;

3-VII double to 12 branched; 3-VII with 4-11 branches; 4-I with 7-27 branches;

4-II with 4-18 branches; 4-III double to 6 branched; 4-IV double to 5 branched;

8 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

4-V with 4-15 branches; 4-VI double to 11 branched; 4-VII double to 7 branched;

4-VIII single to 4 branched; 5-I double to 9 branched; 5-II double to 7 branched;

5-I0 single to 6 branched; 5-IV single to 4 branched; 5-V single to 5 branched;

5-VI single to 7 branched; 5-VII double to 10 branched; 5-VIII with 5-15 branches;

6-I single to triple; 6-II single to 6 branched; 6-III-IV single; 6-V, VI single or

double; 6- VII with 5-18 branches; 7-I single or double; 7-II double to 12 branched;

7-ILl with 5-15 branches; 7-IV double to 17 branched; 7-V with 4-17 branches;

7-VI double to 13 branched; 7-VII single to 6 branched; 8-II double to 4 branched;

8-II single; 8-IV, V, single or double; 8-VI double to 9 branched; 8-VII triple

to 14 branched; 9-I double to 7 branched; 9-II single to 4 branched; 9-III, VI

single or double; 9-IV, V, single; 9-VII single to 5 branched; 10-I double to 8

branched; 10-II, VI single to 4 branched; 10-II, IV, VII single to 6 branched; 10-

V single to triple; 11-I, III-V, VII single to 5 branched; 11-II double to 6

branched; 11-VI double to 5 branched; 12-II, VI single to 6 branched; 12-II, IV

double to 5 branched; 12-V single to 4 branched; 12-VII single to 10 branched;

13-I single to triple; 13-II with 4-24 branches; 13-III triple to 11 branched; 13-

IV double to 11 branched; 13-V double to 13 branched; 13-VI with 9-62 branches;

13-VIl with 4-18 branches; 14-II, IV, V, VIII single; 14-III, VI single or double;

14-VII single to triple; VIII with comb composed of 7-17 scales arranged ina

more or less curved irregular row (cuccioi Belkin with scales in 2 rows, one

row very short), scales long with a stout median apical spine and small denticles

on laterobasal areas or moderately long with uniformly developed denticles on

lateral and apical areas; 1-X moderately long, single to triple, attached near

middle of posterior area of saddle; 2-X moderately long (long in cuccioi), single

to 11 branched (usually multiple branched); 3-X very long, single; ventral brush

usually with 10 setae on grid and 2 precratal shorter ones, all multiple branched,

however, yusafi has 8 setae on grid and 1-2 precratal ones, indicus and nigro-

tarsus (Ludlow) have 12-14 setae on grid and 2-3 precratal ones and latipennis

Delfinado has 15-18 setae on grid and 3-4 precratal ones; saddle moderately

pigmented, incompletely rings segment X, acus absent; 4 anal papillae, usually

moderately long (short in butlevi and very long in cucczoz, hamistylus Laffoon

and robertsi Laffoon). Siphon. Moderately pigmented, moderately long, index

1.58-5.00, acus well developed; pecten on basal 0.49-0.65 (on basal 0. 67-0. 83

in cuccioi, hamistylus, robertsi, similis (Theobald) and trispinatus King and

Hoogstraal), composed of 6-17 teeth (johnsoni Laffoon with 14-20 teeth but when

17 or more present the basal 2-3 are much reduced), apical 1-5 teeth longer

and wider spaced than remainder of teeth, each tooth long, slender and with

1-3 small stout ventral denticles near base, distal 1-4 teeth occasionally with-

out denticles; seta 1-S short, double to 10 branched, base attached on basal

0. 63-0. 83 of siphon distad of last pecten tooth (attached within pecten in

cuccioi, similis and trispinatus);2, 6, 7, 9-S single; 8-S single to 8 branched,

usually multiple.

EGG. The eggs of only 6 species of Verrallina are known including those

of dux which are described here for the first time. Huang (1968) illustrated

and gave measurements for 4 species of the subgenus--carmentt Edwards,

funereus (Theobald), lineatus (Taylor) and pavasimilis King and Hoogstraal.

Egg measurements (in microns) of these 5 species are: dux 637-730 by 154-

179; lineatus 756 by 206; funereus 806 by 268; carmenti 845 by 140 and para -

similis 1040 by 200. Moriya et al. (1973) published a description and scanning

electron microphotograph of the egg of nobukonis Yamada but did not give the

egg measurements.

TAXONOMIC DISCUSSION. The subgenus Verrallina is characterized and

separated from the other subgenera of Aedes by the following: in the adults by

Reinert: New Interpretation of Verrallina 9

the combination of (1) head with decumbent scales of vertex all broad or with

only a double row of narrow curved scales on coronal suture, (2) antennal

pedicel with several short fine hairs and usually a few small broad scales

mesally, (3) dorsocentral (anterior and posterior) and acrostichal (anterior

and posterior) setae numerous, species in Section B usually with only a few

posterior acrostichals, (4) scutum and scutellum with only narrow curved

scales, (5) remigial setae present, (6) subspiracular area and paratergite

bare (indicus with a few pale scales on paratergite), (7) mesepimeron without

anterior lower setae, however, all species except the 12 species of Section B,

have a few to numerous fine hairs caudad of scale patch, (8) propleuron with

4-21 setae and (9) male maxillary palpus short (0. 09-0.18 length of proboscis);

in the female genitalia by the combination of (1) heavily pigmented spermathecal

eminence, (2) presence of lower vaginal sclerite and (3) presence of 3 heavily

pigmented seminal capsules, each of a different size; in the male genitalia by

the combination of (1) complex development of phallosome which consists of

several component parts (see below), (2) gonostylus without gonostylar claw,

(3) sternum IX heavily pigmented throughout, (4) tergum IX without setae and

(5) development of proctiger; in the pupae by the combination of (1) setae 1-3-

CT approximately equally developed, (2) 6,9-VII both weakly developed, (3)

9-VIII single to 6 branched, (4) 1-P single, occasionally double and (5) paddle

without fringe of hair-like spicules; and in the larvae by the combination of (1)

arrangement and development of head setae 4-7-C, (2) antenna with spicules

and seta 2-A with a subapical constriction, (3) pecten with 6-20 (usually 12-16)

teeth, apical 1-5 teeth wider spaced than remainder, (4) acus present on siphon,

absent on saddle and (5) ventral brush usually with 10 setae on grid and 2 pre-

cratal shorter ones, all multiple branched.

All stages of the species of Verrallina can be separated to either Section A

or B but only the female and male genitalia can be readily divided into series.

Species in Series II of Section A appear to occupy an intermediate position

between those in Series I and III and possess morphological features of the other

2 series. Much additional work is needed in establishing species group cate-

gories and the investigation of species complexes. In order to solve these

problems, considerable additional specimen collections of individual and progeny

reared adults, with their associated immature skins, are required.

Adults. of the subgenus Vervallina are similar in habitus to those of the sub-

genus Aedes especially cinereus, however, they differ significantly in the

development of both the male and female genitalia. Species of the subgenus

Aedes differ from the adults of the subgenus Vervallina by the combination of

the following: antepronotum and paratergite with narrow scales (only indicus

of Verrallina bears scales on the paratergite); vertex of head with a broad

median area covered with narrow curved scales; male with a greater number

of setae in antennal whorls; abdominal terga of male with numerous long curved

setae on lateral margins; and mesepimeron without fine hairs caudad or ven-

trad of the scale patch.

The short maxillary palpus of the male is similar to those of the following

subgenera of Aedes: Aedes Meigen, Bothaella Reinert, Cancraedes Edwards,

Christophersiomyia Barraud, Geoskusea Edwards, Huaedes Huang, Lepto-

somatomyia Theobald, Nothoskusea Dumbleton, Pavaedes Edwards and Rhino-

skusea Edwards. Both males and females of Vervallina can be separated from

those of the other subgenera of Aedes by a combination of the above mentioned

characters.

Barraud (1928) first recognized the importance of the female genitalia of

this taxon in separating the included species. The female genitalia of Verrallina

10 Contrib. Anver. Ent. Inst., vol. 11, no. 1, 1974

are very distinctive from all other subgenera of Aedes by the above features.

Other characters that are unusual and can be used to separate species of

Verrallina from many other Aedes are tergum IX wide, usually ribbon-like,

usually without setae except gibbosus and yusafi (some species, such as clava-

tus Barraud, quadrifolium Brug and vallistris Barraud occasionally with a single

short seta on one side); tergum VIII usually very wide; cerci triangular in

shape; insula small, unpigmented with 3-7 tuberculi, each usually with a minute

spicule, or insula absent (Section B) and replaced by median apical portion of

lower vaginal sclerite; and upper vaginal sclerite very large and in many species

covering most of vaginal roof. Each currently recognized species of Vervallina

except members of Section A, Series I, which have very uniform genitalia,

possesses distinguishing features of the female genitalia which separates it

from other species of the subgenus. However, it was found during this study

that the shape of tergum IX was useful in separating a number of species in

Series I. Noticeable similarities of the female genitalia were also observed

between members of related species within species groups.

The male genitalia of Vervallina are very distinct from the other subgenera

of Aedes especially in the complex development of the phallosome which is

divided into the following components: opisthophallus, phallus (with its tergal

penis filament), prosophallus, paramere and parameral apodeme. Several of

the structure are fused at the joints and have no definite sutures (Reinert

1973b). The complex arrangement and partial fusion of the integral parts of

the phallosome of Vervallina appear to be similar to forms found in the dixine

genera Nothodixa Edwards and Pavadixa Tonnoir (these 2 genera discussed and

described by Belkin 1968). Other structures and developments of the genitalia

of this subgenus which distinguish many of the species from a number of sub-

genera of Aedes follows: tergum IX without setae; proctiger with paraprocts

highly modified into short or long free arms or broad contiguous plates, cercal

setae absent; basal mesal lobe modified into a setose lobe which is strongly

fused along sternal margin with tergomesal margin of the ventral surface of

gonocoxite, many species (Section A, Series II and III and Section B) with stout

spiniforms and/or projections; sternum IX heavily pigmented throughout, most

species of Section B and several species of Series III of Section A with broad

scales; gonostylus without a gonostylar claw; and gonocoxite with dorsoapical

and/or ventroapical lobes or projections. The male genitalia of each species

are very distinct; however, the genitalia of species in Series I of Section A

appear to have undergone less differentiation and are more uniform than the

other series and section.

Pupae of Verrallina are very similar in chaetotaxy and most species are

difficult to separate from each other.

The placement and development of setae in the larvae of Verrallina are

fairly uniform within species groups; however, most species possess distin-

suishing features in this stage. A number of species have characters (as

pointed out in the subgeneric and species descriptions) that easily separate

them from other species within the subgenus. Larvae of Verrailina are simi-

lar to those of Aedimorphus Theobald but can be separated from them by the

positions of head setae 4-6-C and the position of the pecten as given by Reinert

(1973a: 11). The subapical constriction of antennal seta 2-A is consistant for

all members of the subgenus Vervallina; however, a number of species of sub-

genera Aedimorphus, Edwardsaedes Belkin, Finlaya Theobald, Neomelanicon-

ion Newstead and Ochlerotatus Lynch Arribalzaga also possess this feature.

The occurrence of precratal setae of the ventral brush is also shared by the

species in the other subgenera (i.e., Aedes, Aedimorphus, Edwardsaedes,

Mucidus Theobald, Neomelaniconion and Ochlerotatus) which inhabit temporary

Reinert: New Interpretation of Verrallina 11

eround pools.

DISTRIBUTION. The subgenus Veryvallina occupies a distribution primarily

in the Oriental Zoogeographical Region but with a short extention north to Kyushu,

Japan and east into the Carolines, New Guinea and South Pacific Islands and

south into northeastern Australia. The ranges of the sections and series of the

subgenus are outlined on the map in Figure 1.

MEDICAL IMPORTANCE. Very little has been reported concerning the

investigation of Verrallina species for pathogens even though a number of

species have been recorded feeding on man and other animals. The inability to

separate the adults of many species by habitus characters may partially account

for the few published accounts of attempts to isolate pathogens from members

of this subgenus.

BIOLOGY. The immature stages, for species which have the biology known,

prefer temporary, unmoving water in shaded pools located in forested areas.

Some species, notably butlervi, prefer brackish water in coastal mangrove

swamps. :

Females of a number of species readily bite man and domestic animals

usually during the daytime and in shaded forested areas. Some species are

serious pests. Adults of many species have been collected in light traps.

The bionomics of Vervallina species have been summarized by Steffan (1966),

Delfinado (1967, 1968), Huang (1968) and Klein (1973). A large number of spe-

cies, however, have the bionomics of the immatures and/or the adults unknown

or only imperfectly known.

KEYS TO SECTIONS AND SERIES OF VERRALLINA

ADULTS!

Mesepimeron with a few to numerous fine hairs caudad of scale patch; post-

tarsi III with ungues simple2, eyes contiguous in fronte; Section A

Mesepimeron without hairs caudad of scale patch; posttarsi III with ungues

toothed: eyes separated im front. 07.02. ak eee oe eS Section B

FEMALE GENITALIA

1. Spermathecal eminence with a cephalad produced pouch; insula absent but

replaced by ''U'' shaped median apical projection of lower vaginal

selerite; without tuberculin 3476 SR ee ee es ee Section B

Spermathecal eminence without a pouch; insula small, unpigmented and

with @ few faberculi Gection- Aye. ow a eae eS ee ee 2

2. Upper vaginal lip with median posterior area produced into a large vertical

shield caudad of spermathecal eminence; spermathecal eminence deep,

ladults are easily separated to section but show no consistant differences

between series of Section A.

2See footnote number 2 on page 13.

3See footnote number 3 on page 13.

12 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

complex and always with numerous well developed spiny excrescences.

Series III (indicus Series)

Upper vaginal lip without a shield; spermathecal eminence shallow, simple,

without spiny excrescences (except some members of Series II)... 3

3. Lower vaginal sclerites moderately pigmented, with spicules.

Series II (butleri Series)

Lower vaginal sclerites very lightly pigmented, without spicules.

Series I (carmenti Series)

MALE GENITALIA

1. Prosophallus long, phallus short and 0.5 or less length of prosophallus;

yWarcnmere Very 1Ong ge SS, eR So Pes Section B

Prosophallus and phallus approximately equal in length; paramere short

CSeGt Gi Aa ie ee ae a ee ee ss ee 2

2. Gonocoxite without dorsoapical or ventroapical lobes or projections, with

one or more subapical spiniforms on sternomesal area.

Series I (carmenti Series)

Gonocoxite with dorsoapical and/or ventroapical or subapical lobes or pro-

jections, subapical spiniforms absent-. .........2.26..06.224.- 3)

3. Prosophallus with apical 0.4-0.5 produced into several lightly pigmented,

leaf-like structures or a single short broad plate; gonostylus attached at

ADEN.OF PONNCOKILO. Sucka Fo a Yad Sei a Series II (butleri Series)

Prosophallus a moderately long to long, heavily pigmented, narrow arm;

gonostylus attached subapically on gonocoxite.

Series III (indicus Series)

PUPA

Respiratory trumpet with basal 0.4 somewhat expanded; ocular plate with well

developed CUlicdlar facets i. ake a ee RS ea Section B

Respiratory trumpet with basal 0.4 usually not expanded, uniform width

throughout or with apical area expanded, ocular plate with or without a

small area of poorly to moderately developed cuticular facets. . Section A

FOURTH STAGE LARVA

1. Antennal seta 1-A single or double? ; head seta 7-C double to 4 branched?.

Section B

Antennal seta 1-A triple to 9 branched; head seta 7-C with 5-15 branches

(Section Adis snc ta Ge Ga EO ERS FG De OG 2

lsee footnote number 3 on page 14.

2See footnote number 1 on page 16.

Reinert: New Interpretation of Vervallina 13

2. Comb scales with uniformly developed denticles on lateral and apical

SrO@Rs °F WAS IP Hey i peek FGSE ea Series I (carmenti Series)

Comb scales with a stout median apical spinel and with small denticles

on laterobasal areas. ... Series Il and III (butleri and indicus Series)

DESCRIPTION OF SECTION A AND SERIES I, II AND II

SECTION A: Adults. Posttarsi III with ungues simple2; eyes contiguous in

fronts; vertex without erect or semi-erect scales, occiput with

erect scales; mesepimeron with a few to numerous fine hairs

caudad of scale patch; postpronotum with narrow curved scales

on upper areat; antepronotum without scales except for a very

few species.

Female geniialia, Insula small, unpigmented, with 3-7 small

tuberculi; sternum VIII with numerous thin setae on pair of

median apical lobes; upper vaginal lip with or without median

posterior area produced into a vertical shield caudad of sper-

mathecal eminence; lower vaginal sclerite very lightly to heavily

pigmented, shape and number of plates variable but always

positioned on floor of vagina and not attached to lower vaginal

lip, with or without spicules; spermathecal eminence shallow to

deep, simple to complex, with or without spiny excrescences,

without a pouch.

Male genitalia. Phallosome with prosophallus and phallus

approximately equal in length, paramere short, parameral

apodeme short to moderately long.

Pupa. Respiratory trumpet usually with basal area not expanded,

but expanded apically or with uniform width throughout; ocular

plate with or without area of poorly to moderately developed

cuticular facets.

Larva, Antennal seta 1-A triple to 9 (usually 4-6) branched;

head seta 7-C with 5-15 (usually 7-14) branches; thoracic seta

4-P triple to 6 (usually 4-6) branched.

1see footnote number 2 on page 15.

2Male of prioekanensis with a small tooth on one ungue.

3ives of mccormicki separated in front.

A few species of Series I did not have scales on the postpronotum but these

may have been rubbed off in the specimens examined.

14 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

Series I.

Series I.

Female genitalia. Upper vaginal lip with median

posterior area narrow and not produced into a shield;

lower vaginal sclerite with a pair of very lightly pig-

mented plates connected by a small apical band, with-

out spicules, all species with uniform shape; spermathe-

cal eminence shallow, simple, composed of a pair of

comma-shaped plates, without spiny excrescences.

Male genitalia. Gonocoxite without dorsoapical or

ventroapical lobes or fleshy projections, with one or

more subapical spiniforms on sternomesal area;

basal mesal lobe consists of a small broad plate with

a number of short to moderately long thin setae;

prosophallus with apical 0. 5-0.6 produced into several

lightly pigmented flattened contiguous leaf-like struc-

tures; gonostylus attached to apex of gonocoxite (quad-

rispinatus King and Hoogstraal and vavriabilis Huang

attached slightly subapically).

Pupa. Respiratory trumpet with a more or less uni-

form width throughout (a few species with apical por-

tion expanded); bases of setae 2-IV-VI (in most spe-

cies) more cephalad than bases of setae 4-IV-VI; seta

9-VIII usually triple to 6 branched.

Larva, Comb scales with uniformly developed denti-

cles on lateral and apical areas; seta 2-X usually single

to triple! | usually with one branch longer than others¢.

Female genitalia. Upper vaginal lip with median pos-

terior area moderately broad and not developed into a

vertical shield; lower vaginal sclerite with a pair of

lightly to moderately pigmented plates, usually ovoid,

with spicules; spermathecai eminence shallow, simple,

with or without poorly developed small spiny excre-

scences.

Male genitalia, Gonocoxite occasionally without but

usually with dorsoapical or ventroapical lobes or fleshy

projections, without subapical spiniforms? on sterno-

mesal area; basal mesal lobe consists of a small plate

with several short to moderately long thin setae and

one or more stout spiniforms or lobes; prosophallus

with apical 0.4-0.5 produced into several lightly pig-

mented flattened contiguous leaf-like structures or a

single short broad plate; gonostylus attached to apex

of gonocoxite.

Seta 2-x triple to 4 branched in sentanius King and Hoogstraal, 6-7 branched

in lineatus and 8 branched in variabilis.

2Branches of seta 2-X approximately equal in length in foliformis King and

Hoogstraal, lineatus and variabilis,

3 Aedes butleriand incertus Edwards with spiniforms but also with apical

projections.

Reinert: New Interpretation of Vervallina 15

Pupa. Respiratory trumpet as for Series I; bases of

setae 2, 4-IV-VI as for Series I; seta 9- VIII usually

Single-+.

Larva, Comb scales with a stout median apical spine2

and with smaller denticles on laterobasal areas; seta

2-X with 4-11 (usually 6 or more) branches which are

approximately equal in length.

Series III. Female genitalia. Upper vaginal lip with median

posterior area developed into a large vertical shield

caudad of spermathecal eminence; lower vaginal

sclerite heavily pigmented, shape and number of

plates variable, usually with spicules; spermathecal

eminence deep, complex, with numerous well developed

Spiny excrescences.

Male genitalia, Gonocoxite with one or more well

developed dorsoapical and/or ventroapical lobes or

fleshy projections, without subapical spiniforms on

sternomesal area; basal mesal lobe consists of a

large plate with several to numerous short to moder-

ately long thin setae and with 2 or more stout spiniforms

and/or one or more large lobes or projections; pro-

sophallus consists of a moderately long to long narrow

heavily pigmented arm; gonostylus attached subapically

to gonocoxite.

Pupa. Respiratory trumpet with apical portion expanded;

bases of setae 2-IV-VI (in most species) more caudad

than bases of setae 4-IV-VI; seta 9-VIII usually single

or double.

Larva. As for Series II.

DESCRIPTION OF SECTION B

SECTION B: Adults. Posttarsi III with ungues toothed; eyes separated in front;

vertex with a few semi-erect scales, occiput with erect scales; .

mesepimeron without hairs caudad or below scale patch; postpro-

notum without scales; antepronotum with broad scales.

Female genitalia. Insula absent but replaced by a ''U'' shaped

median apical projection of lower vaginal sclerite, without tuber-

culi; sternum VIII with numerous very stout setae on pair of

median apical lobes; upper vaginal lip with median posterior area

produced into a large horizontal shield extending over or around

spermathecal eminence; lower vaginal sclerite heavily pigmented,

formed into a pair of ribbon-like plates around floor of vagina,

attached to lower vaginal lip throughout length, fused at apex and

developed into a median, "'U'' shaped projection, plates without

14 edes lugubris Barraud with seta 9-VIII triple to 6 branched.

24 edes butleri with 1-2 slightly longer and stouter median apical denticles.

16

CONRHRwWN re

azureosquamatus Bonne-Wepster** 14.

Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

spicules; spermathecal eminence shallow, simple, without spiny

excrescences, with a heavily pigmented cephalad spine-lined

pouch.

Male genitalia, Phallosome with prosophallus very long, phallus

short and 0.5 or less length of prosophallus, paramere very long, —

parameral apodeme very long.

Pupa. Respiratory trumpet with basal 0.4 somewhat expanded;

ocular plate with an area of well developed cuticular facets; bases

of setae 2-IV-VI more cephalad than bases of setae 4-IV-VI.

Larval, Antennal seta 1-A single or double; head seta 7-C

double to 4 branched; thoracic seta 4-P single to triple.

LIST OF SPECIES IN SECTIONS AND SERIES

SECTION A

SERIES I (carmenti Series)

obsoletus Huang***

bifoliatus King and Hoogstraal**

carmenti Edwards**

cuccioi Belkin**

embiensis Huang**

foliformis King and Hoogstraal**

funereus (Theobald)*

killertonis Huang***

leilae King and Hoogstraal**

lineatus (Taylor)*

mccormicki Belkin**

milnensis King and Hoogstraal**

multifolium King and Hoogstraal**

15.

16.

ee

18.

parasimilis King and Hoogstraal**

pipkini Bohart**

quadrifolium Brug*

quadrispinatus King and

Hoogstraal**

reesi King and Hoogstraal**

sentanius King and Hoogstraal**

similis (Theobald)*

simplus King and Hoogstraal**

trispinatus King and Hoogstraal**

vanapus Huang***

variabilis Huang***

1A4edes consonensis Reinert is somewhat aberrant for Section B in that one

larva has seta 1-A triple on one side, 7-C with 7 branches on one side and

4-P with 4 branches on one side.

*

EK

2K

= Specimens of this species examined.

= Holotype or lectotype examined.

wes Paratypé examined.

(?) = Assignment to series uncertain.

orm & be

our Whe

Reinert: New Interpretation of Vervallina 17

SERIES II (utlevi Series)

butleri Theobald** 6. lugubris Barraud*

cunninghami Taylor (?) 7. panayensis Ludlow**

dux Dyar and Shannon** 8. prioekanensis Brug**

incertus Edwards** 9. pseudodiurnus (Theobald)*

iviomotensis Tanaka and 10. varus Delfinado**

Mizusawa***

SERIES III (indicus Series)

adustus Laffoon** 25. neomacrodixoa King and

agrestis Barraud** Hoogstraal**

andamanensis Edwards** 26. nigrotarsis (Ludlow) **

atriisimilis Tanaka and 27. notabilis Delfinado**

Mizusawa *** 28. nubicolus Laffoon**

atrius Barraud* 29. petroelephantus Wijesundara**

campylostylus Laffoon** 30. philippinensis Delfinado*

cautus Barraud** ; 31. phnomus Klein**

clavatus Barraud* 32. protuberans Delfinado**

comatus Barraud** 33. pseudomediofasciatus

comosus Reinert** (Theobald)*

cretatus Delfinado** | 34. pseudovarietas Reinert**

cyrtolabis Edwards* 35. vami Barraud**

gibbosus Delfinado** 36. sabahensis Reinert**

harrisonicus Reinert** 37. seculatus Menon**

hispidus Delfinado** 38. singularis (Leicester) **

indicus (Theobald)** 39. sohni Reinert**

johnsoni Laffoon** 40. spermathecus Wijesundara

johorensis Reinert** 41. torosus Delfinado**

kRompongus Klein** 42. uncus (Theobald)*

lankaensis Stone and Knight** 43. vallistris Barraud*

latipennis Delfinado** 44, varietas (Leicester)**

leicesteri Edwards ** 45. virilis (Leicester)**

macrodixoa Dyar and Shannon** 46. yerburyi Edwards*

margarsen Dyar and Shannon**

SECTION B

abditus Barraud** 7. pahangi Delfinado**

consonensis Reinert** 8. vamalingami Reinert**

fragilis (Leicester)** 9. vrobertsi Laffoon***

hamistylus Laffoon** 10. stungus Klein**

indecorabilis (Leicester) ** 11. wuniformis (Theobald)*

nobukonis Yamada* 12. yusafi Barraud**

18 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

SPECIES TREATMENT

AEDES (VERRALLINA) ABDITUS BARRAUD

(Figs. 2, 22)

The following information on the holotype female supplements the descrip-

tions of Barraud (1931: 615, 1934: 282).

FEMALE. Head, Antenna 1.42 length of proboscis, pedicel brown with a

few small brown scales and short fine brown hairs mesally; maxillary palpus

0.2 length of proboscus; proboscus 0.84 length of femur I; eyes separated in

front. Thorax, Scutum nearly completely denuded of scales but a few narrow

curved reddish-black scales remain, Similar scales on each lobe of scutellum;

antepronotum with scales rubbed off but scars present; postpronotum without

scales, 3-4 dark posterior setae; postspiracular area without scales, 3-4

dark setae; subspiracular area, paratergite, mesomeron and metameron

bare; prealar knob without scales, 4-6 dark setae; mesepimeron with a large

patch of broad white scales near center, 5-6 dark setae dorsad of scale patch.

Legs. Posttarsi I-III each with 2 ungues, equal and each with a tooth. Abdo-

men. Terga I-V each dark brown scaled and with a narrow lateral stripe of

creamy-white scales; sterna II-V each creamy-white scaled; remainder of ab-

domen removed with genitalia. Genitalia. As in Figure 2.

DISCUSSION. The adult habitus of abditus is similar to that of indecorvabilis

but differs as follows: abditus with a lateral narrow stripe of creamy-white

scales on abdominal terga and sterna with creamy-white scales, while indecor-

abilis possesses a lateral broad stripe of whitish scales on abdominal terga

and sterna with creamy-brown scales.

Female genitalia of abditus are illustrated in order to show all structures

including the lower vaginal lip and sclerites, terga VIII and IX, sternum VIII

and seminal capsules. Distinctive parts of the genitalia of this species are

compared to the other members of Section B in Figures 21 and 22.

AEDES (VERRALLINA) ADUSTUS LAFFOON

(Figs. 25, 36, 48, 58, 60)

FEMALE. No differences could be found between the adults and the geni-

talia of adustus and andamanensis.

PUPA (Fig. 36). Chaetotaxy as figured and recorded in Table 2. Ocular

plate with moderately developed cuticular facets. Respiratory trumpet.

Moderately pigmented; hair-like spicules on distal 0.38-0.51 of inner surface;

index 3.47-4.22, average 3.85. Abdomen. Setae 6, 9-VII weakly developed;

4-VIII single to 4 branched, usually double; 9-VIII single. Paddle. Ovoid;

small serrations on distal portion of basal 0.61-0.63 of outer margin; tiny

spicules on apical 0.37-0.39 of outer and apical 0.12-0.21 of inner margins;

midrib nearly reaches apex; seta 1-P single; index 1.32-1.61, average 1.51.

LARVA (Fig. 48). Chaetotaxy as figured and recorded in Table 15.

Head, Seta 4-C with 5-8 branches; 5-C with 6-10 branches; 6-C with 4-7

branches; 7-C with 10-14 branches; 12-C with 4-9 branches; dorsomentum with

33-35 teeth. Antenna. Moderately long, moderately pigmented, with several

scattered small spicules, more numerous on basal area; seta 1-A with 4-6

Reinert: New Interpretation of Vervallina 19

branches, moderately long, branches thickened; 2-A long, with a subapical

constriction; 3-A short, 0.29-0.39 length of 2-A; 4-A moderately long, 0. 50-

0.69 length of 2-A; 6-A short, 0.26-0.37 length of 2-A. Thovax,. Seta1-P

long and single. Abdomen. Seta 1-VIII with 4-6 branches; 3-VIII with 5-11

branches; 5-VIII with 5-8 branches; 2-X with 6-12 branches, all approximately

equal in length; ventral brush with 9-11 setae on grid and 2-3 precratal shorter

ones, all multiple branched, usually with 10 setae on grid and 2 precratal ones;

saddle moderately pigmented, incompletely rings segment X, acus absent; 4

moderately long to long anal papillae; comb with 7-15 (usually 10) scales

arranged in a single curved irregular row, each scale with a stout median

apical spine and small denticles on laterobasal areas. Siphon. Moderately

pigmented; acus well developed; index 2.'73-3.50, average 3.11; pecten on

basal 0. 56-0. 59 of siphon, composed of 14-18 (usually 15) teeth, apical 2-4

teeth longer and wider spaced than remainder of teeth, each tooth long, slender

and with 1-2 small stout ventral denticles near base; seta 1-S short, triple to

) branched, base attached on basal 0. 66-0. 76 of siphon distad of last pecten

tooth.

DISCUSSION. Aedes adustus, previously reduced to a synonym of anda-

manensis by Delfinado (1967: 9), is here resurrected from synonymy on

characters of the male genitalia. This species belongs to a species complex

(i.e. adustus, andamanensis, johnsoni, margarsen and sohnt) in which the

members have previously been difficult to separate from each other. The

types of these 5 species have been examined. Aedes adustus is separated

from the other members of the complex by the presence of a short spine-like

apical sternomesal projection on the basal mesal lobe (Fig. 25). The develop-

ment of the basal mesal lobe of adustus is intermediate between andamanensis

and sont, both of which lack an apical sternomesal projection, and johnsoni

and margarsen, both of which have a long, slightly curved, knife-like, apical

sternomesal projection. Aedes adustus occupies much of the same geographi-

cal range as andamanensis, however, all specimens of this group examined

from Sabah were adustus (except for the holotype male of sohni). Since females

of the above 2 species can not be separated with certainty they are recorded as

andamanensis Group.

Adults of this complex (andamanensis Group) are easily distinguished by

the presence of fine hairs on the metameron from the other species in the

subgenus except gibbosus which also has hairs and a few scales on this struc-

ture. Aedes gibbosus, however, is easily separated from this group by fea-

tures of the male genitalia, absence of scales on the antepronotum and post-

spiracular area and the absence of setae on the prosternum while members of

the andamanensis Group have scales on the antepronotum and postspiracular

area and setae on the upper mesal area of the prosternum (setae absent in

johnsont).

Illustrations are given of tarsomeres 5 and posttarsi of the male (Fig. 58)

and the female (Fig. 60).

The pupa and larva of adustus are described for the first time and are

similar to those of andamanensis.

DISTRIBUTION. 168 specimens examined: 559, 540°, 33 L,21 with

associated skins (16 p, 51).

CAMBODIA. Kompong Sela, Stung Chral; 2.

MALAYSIA. Kedah, Kg. Peng Besar, Kuala Nerang; 1, Kelantan, Ke.

Siupol, Mochang; 3. Malacca, Taugea Batu; 50°. Negri Sembilan, Kg. Kuala

Johol, Kg. Tengah, Tampin; 3°. Perlis, Ke. Gunong; 1% pl, 20°. Sabah, Apin,

Beaufort, Jesselton, Keningau, Kota Belud, Bandau, Kudat, Labuk and Sugut,

20 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

Papar, Pulau Labuan, Rangan, Sandakan, Mount Pyramid, Mount Tiger, Kg.

Gading, Tawan, Tenam; 12 pl, 22 p, 462, 20% pl, 10 p, 120°, 33 L.

PHILIPPINES. Balabac Island, Cape Melville; 62 and 5c (paratypes), 1°

(holotype) and 10° with asymmetrical genitalia (see Laffoon 1946: 234, 236).

THAILAND. Chiang Mai, Ban Rong Rua Taeng; 1“ p. Khon Kaen, Tham

Pho Ti Yan; 1%. Krabi, Ban Phru Toei; 6% p. Phangnga, Pak Chaung; 10 p.

Prachin Buri, Ban Bu Phram, Ban Tub Lan; 1“ pl, 4& p. Songkhla, Amphoe

Haad Yai, Ton Nga Chang Waterfalls; 1° p.

BIOLOGY. In Malaysia immatures were collected from a small wheel rut

and from a marshy depression with buffalo footprints; from colored, fresh,

unmoving, temporary water in partially shaded areas located in nipa palm and

rubber plantations; and at elevations from sea level to 213 m (most often at

sea level). Adults were taken biting humans (on legs) between 1900-2000 hours

(1 April 1967) and resting in nipa palms between 1000-1100 hours (27 March

1967).

In Thailand immatures were collected from small ground pools, a small

ditch and elephant footprints; from clear or turbid, fresh, unmoving, tempor-

ary water in partially to heavily shaded areas located in secondary scrub, in

hilly and valley terrain; and at elevations from 80-330 m.

AEDES (VERRALLINA) AGRESTIS BARRAUD

(Fig. 3)

DISCUSSION. Only the female genitalia, which were mounted in Canada

balsam on a microscope slide, remains of the holotype in the BMNH (Peter

Mattingly, 21 May 1973, personnal communication). Therefore, since

additional specimens have not been collected, no additional information on

the adult habitus can be added to the original description of Barraud (1931).

The female genitalia have been examined and are illustrated (Fig. 3) showing

for the first time the development of the lower vaginal lip and lower vaginal

sclerite. The latter consists of a large, heavily pigmented plate with a deep

median apical indentation. The development of the spermathecal eminence is

unique for this species.

AEDES (VERRALLINA) ANDAMANENSIS EDWARDS

(Figs. 23, 25, 56, 58, 60)

DISCUSSION. Aedes andamanensis belongs to a group of closely related

species (i.e. adustus, johnsoni, margarsen and sohni) but can be separated

from the other members of the complex by characters of the male genitalia

(Figs. 25 and 56) and from all except adustus by its geographical distribution

(see discussion under adustus).

Illustrations are given of the tarsomeres 5 and posttarsi of the male

(Fig. 58) and the female (Fig. 60).

The male genitalia illustration of andamanensis by Delfinado (1967, Fig. 2)

is adustus,

Additional distribution and biology information is given below.

DISTRIBUTION. 63 specimens examined: 40c, 16 with associated skins

(9p, 71).

INDIA. Andaman Islands; 10 (holotype).

INDONESIA. Java, Padaberang; 10%. Sumatra, Atchin, Kotta Tjane; 1°.

Reinert: New Interpretation of Verrallina 74

MALAYSIA. Kedah, Kg. Peng Besar, Kuala Nerang; 1% p, 10%. Kelantan,

Kg. Siupol, Mochang, Pasir Puteh, Selising; 2°. Malacca, Malacca, Taugga

Batu; 1° pl, 50%. Negri Sembilan, Kg. Kuala Johol, Kg. Marjid Tengah,

Tampin; 2°. Pahang, Bt. Betong, Kuala Lipis, Merapah, Gna Mueang; 1° p,

1c’. Perlis, Bukit Pintang Forest Reserve, Kg. Gunong; 1° p, 3°,

THAILAND. Chiang Mai, Amphoe Maerim, Ban Rong Rua Taeng; 2° p.

Kanchanaburi, Ban Sai Yok; 1% p. Khon Kaen, Pha Dong Larn, Tham Pho

Ti Yan; 50°. Kvabi, Ban Phru Toei; 5° pl. Nakhon Ratchasima, Ban Wang

Mut; 1 p. Phangnga, Pak Chaung; 2°. Pvrachin Buri, Ban Bu Phram; 1° pl,

20" p,

BIOLOGY. The preferred habitat of immatures appears to be small, fresh

water, flood and ground pools in partial to heavily shaded areas. In Thailand

immatures were collected from small and large flood pools, a small ditch,

small animal footprints, elephant footprints, large wheel ruts, flood pools in

dry stream beds; from clear or turbid, fresh, unmoving, temporary water in

partially to heavily shaded areas (once from an unshaded area); in secondary

scrub forests and once each from the following: secondary rain forest, rubber

plantation, and a fruit, banana and pineapple farm; and at elevations from 80-

350 m (most often at 100-350 m).

In Malaysia immatures were collected from small and large ground and

flood pools, small pools ina dry stream, small buffalo footprints, anda

small wheel rut; from clear or colored, fresh, unmoving, temporary water

in partially to heavily shaded areas (3 times from unshaded areas) located

in secondary rain forests; and at elevations from sea level to 61 m.

AEDES (VERRALLINA) ANDAMANENSIS GROUP

DISCUSSION. The females of andamanensis and adustus can not be separ-

ated by either habitus or genitalia features. These 2 species also occupy much

the same geographical areas. Aedes andamanensis has not been collected

from Sabah, Malaysia, therefore all females of this group from this area are

considered to be adustus. The female of sohni, which has been collected from

Sabah and southern Thailand, is not known so that the exact status of the fe-

males from Sabah are also in doubt. The females of the andamanensis Group

from Cambodia, Thailand and West Malaysia are listed below. Many addition-

al specimens of progeny reared adults with associated immature skins from a

number of different localities within the range of these 2 species will be re-

quired in order to understand this species complex.

DISTRIBUTION. 181 specimens examined: 872, 15 L, 47 with associated

skins (15 p, 32 1).

CAMBODIA. Kompong Sela, Stung Chral; 29.

MALAYSIA. Kedah, Kg. Peng Besar, Kuala Nerang; 19°. Kelantan, Pasir

Puteh, Selising; 12 pl, 22, 7 L. Malacca, Malacca, Taugga Batu; 6¢ pl, 89,

6L. Negri Sembilan, Kg. Kuala Johol, Kg. Marjid Tengah, Tampin; 6%, 2 L.

Pahang, Merapoh; 2°. Pevak, Chior Forest Reserve; 12. Perlis, Kg. Gunong,

Kg. Prok Buah; 4@ pl, 1° p, 1°.

THAILAND. Khon Kaen, Pha Dong Larn, Tham Pho Ti Yan; 32. Krabi,

Ban Phru Toei; 72 pl. Nakhon Ratchasima, Ban Wang Nam Khea; 6 pl, 12.

Nakhon Si Thammarat, Amphoe Chawang, Banna Nabon, Ban Tan; 3°. Nan,

Ban Ta Loc, Pha Dang Khawi; 2? pl, 19. Phangnga, Pathum; 12 p, 5°. Pra-

chin Buri, Ban Bu Phram, Ban Tub Lam; 62 pl, 92 p, 1°. Songkhla, Amphoe

Haad Yai, Ton Nga Chang Waterfalls; 42 p, 22. Trang, Trang; 19.

22 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

BIOLOGY. In Thailand females were collected biting cattle and horses.

AEDES (VERRALLINA) ATRIUS BARRAUD

(Pig. 23, 60)

DISCUSSION. Tarsomeres 5 and posttarsi of the female (Fig. 60) and ter-

sum IX of the female genitalia (Fig. 23) are illustrated for the first time.

Many of the following specimens represent new distribution records in

Thailand. Additional information on the biology of this species is also given.

DISTRIBUTION. 73 specimens examined: 239, 17%, 11, 4L, 19 with

associated skins (10 p, 91).

THAILAND. Chiang Mai, Chiang Mai, Doi Sutep; 22. Chon Buri, Amphoe

Bang Lamung, Khao Mai Keo, Khao Wai; 12. Lampang, Ban Pak Hoi, Ban

Pha Mon, Ban Rai Na Dieo; 12 p, 42, 1% pl, 30°. Nakhon Nayok, Amphoe

Muang, Khao Tha Kor, Pha Kluay Mai Waterfalls; 32 p, 5°, 1% pl, 1¢% p, 3c,

2L. Nakhon Ratchasima, Amphoe Pak Chong, Dong Chao Thong, Kong Koa

Waterfalls, Lam Nam Tha Klong; 22, 1%, 11, 2 L. Nan, Ban Pha Mon; 1c.

Phuket, Khao Prathiu; 1¢ pl, 12, 3& pl. Tak, Khao Salak Phra; 12 pl, 12 p,

19). 26 pl...

BIOLOGY. In Thailand immatures were collected from small and large

flood pools in dry stream beds and small rain water puddles containing clear

or turbid, fresh, unmoving, temporary water with dead leaves, in partially

to heavily shaded areas located in primary and secondary rain forests and a

secondary deciduous forest and at elevations from 85-823 m (most often at

120-250 m), Adults were collected biting man in a forest between 1515 and

1545 hours.

AEDES (VERRALLINA) BUTLERI THEOBALD

(Figs. 4, 26, 37, 49, 56, 58, 60, 63)

Aedes (Aedes) seculatus Menon 1950, 2 only synonym carteri Wijesundara 1951,

Ceylon J. Sci. 24: 176 (2*) (see discussion section of seculatus). NEW

SYNONYM. ,

Aedes butleri is the type species for the subgenus Vervallina and the

representative species for Series II, Section A, and is therefore completely

redescribed below. :

FEMALE (Fig. 63). Head, Antenna dark brown, 1.01-1.11 length of

proboscis, pedicel dark brown with a few small dark scales and short fine

hairs mesally, flagellomere 1 with basal 0.25 pale and with several small

dark scales near middle, flagellar whorls with 6 dark setae; clypeus dark

brown, bare; maxillary palpus blackish-brown scaled, 4 segmented, 0.17-

0.21 length of proboscis; proboscis blackish-brown scaled, 1.17-1.24 length

of femur I; eyes contiguous; ocular setae blackish-brown and well developed;

scales on head all broad and decumbent except upper interocular space which

has 6-8 narrow curved white scales (some specimens with these scales pale

brown) and occiput with a patch of short erect forked scales, mesal ones

dark and lateral ones paler; vertex blackish-brown scaled with a double row

of broad white scales on midline extending anteriorly nearly to ocular line

(some specimens with only a few white scales); lateral surface blackish-

Reinert: New Interpretation of Verrallina 20

brown scaled with a narrow stripe of white scales extending from area in front

of antepronotum to ocular line. Thovax. Scutal integument dark brown; scu-

tum covered with narrow curved reddish-black scales except for a small patch

of narrow curved white scales on median anterior promontory area; prescutel-

lar space bare; scutellum with a patch of narrow curved reddish-black scales

on each lobe; dark reddish-black setae on the following areas: 2-4 median

anterior promontory, numerous acrostichal (anterior and posterior), numerous

dorsocentral (anterior and posterior), scutal fossal (4-6 anterior, 4-5 lateral

and 2 posterior), numerous supra-alar, 5-6 posterior medial scutal, 1 postalar

callar and scutellar (5-6 long and 3-4 short ones on lateral lobe, 5-7 long and

4-5 short ones on median lobe); pleural integument dark brown; antepronota

widely separated, scales absent, 11-17 long and short dark setae; postpronotum

with narrow curved reddish-black scales on upper area, 5-8 posterior dark

setae; propleuron with a patch of broad white scales, 9-12 dark setae; pro-

sternum, subspiracular area, paratergite, mesomeron and metameron bare;

postspiracular area without scales, 3-8 dark setae; mesepisternum with an

upper and a lower patch of broad white scales, several cephalic scales of

upper patch show a brownish tinge, 4-5 upper and 14-16 posterior dark setae,

a few (3-6) short fine pale hairs forming a small patch cephalad of upper

scale patch near subspiracular area; prealar knob without scales, 9-12 dark

setae; mesepimeron with a patch of broad white scales near center, 13-19

setae (several hair-like) dorsad of scale patch, several fine golden hairs on

areas caudad and below scale patch but not extending over lower area. Legs,

Coxae I-III each with several dark setae, I with broad brown scales and a

small dorsal patch of white ones on anterior surface, II and III each witha

patch of broad white scales on outer area of anterior surface; trochanters I-III

each with broad pale brown scales; femora I and II each with anterior surface

dark brown scaled, I with a broad posterodorsal longitudinal stripe of pale

brownish scales, II with a broad posteroventral longitudinal stripe of pale

brownish scales, remainder of posterior surfaces of I and II dark brown

scaled, III dark brown scaled with an anteroventral broad longitudinal stripe

of creamy colored scales from base to near apex, posterior surface with a

ventral broad longitudinal pale brownish stripe; tibiae I-III and tarsi I-III

blackish-brown scaled; posttarsi I-III (Fig. 60) each with 2 ungues, I and II

with ungues equal, each with a tooth, III with ungues equal, both simple.

Wing. Dorsal and ventral veins with blackish-brown scales; alula with

several moderately broad dark brown scales on margin; upper calypter with

several pale brown setae on margin; 1 remigial seta. Halter. Pedicel pale;

capitellum dark brown scaled. Abdomen. Terga blackish-brown scaled,

laterotergite of I with a rectangular patch of white scales, II-VII each with a

lateral white scaled spot, extreme lateral portion of spot basal; sterna brown

scaled each with a laterobasal white scaled spot, V-VI also each with a few

pale scales on mesal areas; terga and sterna with numerous golden setae,

mostly along posterior margins. Genitalia (Fig. 4). Tergum VIII moderately

pigmented, wide, base and apex both nearly straight, lateral margins slightly

concave, several moderately long stout setae apically and a few short ones

scattered over apical 0.40-0.57, basolateral seta present, covered with

minute spicules, numerous broad scales covering apical 0. 70-0.72, basal

0. 10-0. 35 retracted into segment VII, VIII-Te index 0.59-0.65, VIII-Te/IX-

Te index 3.14-3.75; sternum VIII moderately pigmented, wide, base concave,

apex with a moderately deep median indentation with a small lobe on each side

of midline, numerous short and moderately long setae scattered over apical

0.80-0.87, setae somewhat stouter at apex and on median lobes, also more

24 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

numerous on the latter, covered with minute spicules, numerous broad scales

covering apical 0. 83-0.87 except for a small median apical area, apical inter-

segmental fold unpigmented, VIII-S index 0. 65-0. 68; tergum IX moderately

pigmented, wide, short and band-like, base with a short broad median indenta-

tion, apex slightly concave, setae absent, covered with minute spicules, IX-Te

index 0.32-0. 38; insula unpigmented, short, covered with minute spicules,

4-5 small tuberculi, each with a minute spicule; lower vaginal lip moderately

pigmented, narrow, covered with minute spicules, with a small caudally pro-

jected heavily pigmented median apodeme, lower vaginal sclerites composed .

of a basal pair of oblong moderately pigmented structures covered with small

spicules on dorsal surface and situated on an elevated portion of the membrane

extending over area between the vaginal lip, and a small subapical heavily pig-

mented transverse bar; upper vaginal lip moderately to heavily pigmented,

narrow to moderately broad, posterolateral area produced into a small lobe,

posterior and posterolateral lobe covered with minute spicules, upper vaginal

sclerite very large, well developed, base attached to basal half of lateral area

of upper vaginal lip and with a heavily pigmented apodeme on posterobasal area,

sclerite bifurcate with anterior branch wide, moderately pigmented and attached

to cephalic portion of spermathecal eminence, posterior branch wide, heavily

pigmented with a caudally produced, heavily pigmented, mesally curved arm

connected with its mate by a small membranous strip which forms a crown

around the spermathecal eminence; spermathecal eminence heavily pigmented,

shallow in depth, composed of a pair of comma shaped lateral structures con-

nected caudally and produced into a fleshy very lightly pigmented cephalic pro-

jection, moderately long spiny excrescences on lateral and cephalic areas;

postgenital lobe short, wide, apex broad with a small median indentation, 14-

19 setae on each side of midline, completely covered with small spicules, dor-

sal PGL index 0.87-1.00, ventral PGL index 1.11-1.21; peri-anal membrane

with small spicules; cercus triangular in shape, moderately long and broad at

base, apex acute, completely covered with small spicules, dorsal surface with

a number of long and moderately long setae and a few short ones on apical 0. 91-

0.94, ventral surface with a few short setae along outer area, a number (12 or

more) of broad scales on dorsal surface, cercus index 2.38-2.72, cercus/

dorsal PGL index 3. 03-3.43; 3 seminal capsules, 1 large, 1 medium and 1

small in size, heavily pigmented, spherical, each with a short pigmented neck,

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

duct moderately pigmented.

MALE. Similar to female in general habitus but with the following differ-

ences. Head, Antenna plumose, 0.97-1.09 length of proboscis; maxillary

palpus 0.11-0.14 length of proboscis; proboscis 1.09-1.11 length of femur I;

vertex with only 2-3 broad white scales on midline. Thorax. Scutellar setae

(3-4 long and 3-4 short ones on lateral lobe, 4 long and 4-5 short ones on

median lobe); 9-11 antepronotal setae; 4 postpronotal setae; 4-6 propleural

setae; 2-3 postspiracular setae; 5-7 prealar knob setae; mesepimeron with 6-8

setae dorsad of scale patch. Legs. Posttarsi I-III (Fig. 58) each with 2 ungues,

I with ungues unequal, each with a tooth, II with ungues unequal, larger ungue

simple and smaller ungue with a tooth, III with ungues equal, both simple.

Abdomen. Tergum VIII (Fig. 56) wide, caudal lateral area produced into a lobe

bearing a patch of very long stout curved setae and median caudal area bearing

a patch of moderately long to long stout setae. Genitalia (Fig. 26). Tergum IX

moderately to heavily pigmented, band-like with median caudal margin slightly

concave and strongly fused to tergum X, setae absent, narrowly connected

laterally to sternum IX; gonocoxite short, broad, with ventral surface strongly

Reinert: New Interpretation of Vervallina 29

produced sternally, heavily pigmented, mesal surface membranous with minute

spicules, dorsal surface with a few very long stout setae on lateral portion of

apical area, ventral surface with apex slightly produced caudally and bears 2

long broad heavily pigmented longitudinally striated setae with bases nearly

contiguous and apices bluntly pointed and one longer narrower stout lanceolate

seta slightly ventrad of other 2, mesal subapical area with a small moderately

to heavily pigmented bulla which bears a moderately long sternal arm, a short

proximal conical tergal projection and a small distal lump, one moderately

long seta between bulla and 2 broad apical striated setae, remainder of ventral

surface with numerous short thin setae and a few broad scales, lateral surface

with several long stout setae on apical area and a few broad scales on basal

area; gonostylus moderately pigmented, long (approximately 1.1 length of

gonocoxite), basal area formed into a large lobe with minute spicules and 8-10

short,moderately long and long stout setae, apical portion bifid, produced into

a ventral shorter striated blade-like structure with a few minute spicules, and

a dorsal narrow arm which is thicker distally and terminates in a short pointed

hook, distal area bears 3-9 short thin setae and basal area of arm bears 3

moderately long stout setae, attached to apex of gonocoxite; basal mesal lobe

lightly to moderately pigmented, consists of a small basal lobe bearing 8-13

short thin setae, each of which arise from a tubercule, and a small caudal

lightly pigmented indistinct area on mesal membrane of gonocoxite bearing

1-3 very short setae, lobe connected mesally with its mate by a narrow moder-

ately pigmented band covered with minute spicules and located ventrad of

prosophallus and phallus; proctiger continuous tergally, moderately pigmented,

paraproct not separated but restricted to a very heavily pigmented band along

the sternal margin of proctiger, apex with 3-6 small heavily pigmented teeth

and base with a small short sternal projection which articulates at a point on

the dorsomesal surface of the parameral apodeme near base of opisthophallus,

cercal setae absent; tergum X moderately to heavily pigmented, strongly fused

to caudal margin of tergum IX and base of proctiger, margins indistinct but

fusion lines somewhat wrinkled; phallosome complex, opisthophallus--consists

of a moderately pigmented tergal transverse bridge between the parameral

apodemes, lateral surface curved ventrally, caudal margin with outer area

produced into a cup-like lobe, base attached to a dorsomesal extension of the

caudal portion of parameral apodeme, located dorsad of phallus and prosophallus

and ventrad of proctiger, phallus--consists of a pair of moderately long heavily

pigmented arms which are joined near their center by a narrow sternal bridge,

apices of arms contiguous giving an A-shaped appearance, base of arm attached

to inner tip of paramere, tergal area of arm basad of sternal bridge strongly

fused to inner sternobasal portion of prosophallus, apex of phallus with a

loosely attached tergal penis filament which consists of a small moderately

pigmented U-shaped structure and is produced cephalad, prosophallus--con-

sists of a bulbous base which is fused to tergobasal area of phallus and tergo-

mesal portion of paramere, apical 0.4-0.5 produced into several lightly pig-

mented flattened contiguous leaf-like structures with apices fused into a single

flattened plate which is positioned between laterosternal surface of opistho-

phallus and lateral margin of phallus, prosophallic arm approximately equal in

length to phallic arm, paramere--heavily pigmented, moderately long, approxi-

mately 0.67 length of phallic arm, articulates at a point near middle of outer

margin with caudal area of parameral apodeme, parameral apodeme--moder-

ately to heavily pigmented, moderately long, outer surface broadly attached to

tergobasal apodeme of gonocoxite; sternum IX heavily pigmented throughout,

large, broad with apicolateral areas rounded, 6-11 short and moderately long

26 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

setae in a small patch near middle of caudal margin.

PUPA (Fig. 37). Chaetotaxy as figured and recorded in Table3. Ocular

plate without or with a small area of very poorly developed cuticular facets.

Metanotal plate and areas of cephalothorax and first 3-4 abdominal segments

heavily pigmented. Respiratory trumpet. Heavily pigmented; hair-like spicules

on distal 0. 75-0. 82 of inner surface; index 4.13-5.60, average 4.83. Abdomen.

Setae 6, 9-VII weakly developed; 4-VIII triple to 5 branched; 9-VIII single.

Paddle. Ovoid; minute serrations on distal portion of basal 0.62 of outer mar-

gin; minute spicules on apical 0.38 of outer and apical 0.3 of inner margins;

midrib does not reach apex; seta 1-P moderately long, single; index 1.17-1. 58,

average 1.33.

LARVA (Fig. 49). Chaetotaxy as figured and recorded in Table 16. Head,

Seta 4-C triple to 6 branched; 5-C double to 4 branched; 6-C triple to 4

branched; 7-C with 5-7 branches; 12-C with 4-5 branches; dorsomentum with

21-26 teeth. Antenna, Moderately long, lightly pigmented, with several small

scattered spicules; seta 1-A with 6-9 branches, moderately long; 2-A long,

with a subapical constriction; 3-A short, 0.22-0.27 length of 2-A; 4-A moder-

ately long, 0.40-0.56 length of 2-A; 6-A short, 0.22-0.29 length of 2-A.

Abdomen. Seta 1-VIII triple to 5 branched; 3-VIII with 5-11 branches; 5-VIII

with 9-15 branches; 2-X with 6-8 branches all approximately equal in length;

ventral brush with 10-11 setae on grid and 2 precratal shorter ones, all

multiple branched; saddle moderately pigmented, incompletely rings segment

X, acus absent; 4 short anal papillae; comb with 9-14 (usually 12) scales

arranged in a single curved irregular row, each scale with 1-2 slightly stouter

and longer median apical spines and smaller denticles on laterobasalareas.

Siphon. Moderately pigmented; acus well developed; index 2.32-2.82, average

2.53; pecten on basal 0. 52-0. 56 of siphon, composed of 9-12 (usually 10) teeth,

apical 2-3 teeth longer and wider spaced than remainder of teeth, each tooth

long, slender and with 1-2 small stout ventral denticles near base; seta 1-S

short, 4-7 branched, base attached on basal 0. 64-0. 71 of siphon distad of last

pecten tooth.

DISCUSSION. Aedes butleri belongs to a species group with dux, iriomo-

tensis and lugubris. These species are easily separated by features of the

male and female genitalia but are more difficult to distinguish by adult habitus

features. The following features can be used to separate adult females of the

group: butleri possesses abdominal terga II-VII with laterobasal white scaled

patches, scutellum with narrow curved dark brown scales on each lobe; ivio-

motensis possesses abdominal terga II-VII with laterobasal white scaled patches,

scutellum with narrow curved scales on each lobe, scales dark brown on lateral

lobes, median lobe dark brown scaled with a small median apical patch of

white scales (not reported in the original description by Tanaka and Mizusawa

1973: 633, but present in the 10 paratypes examined); lugubris possesses ab-

dominal terga II- VII with laterobasal white scaled patches, II also with a tergo-

basal white scaled band connected with laterobasal white patch (band on a few

specimens incomplete mesally), scutellum with narrow curved dark brown

scales on each lobe; and dux possesses abdominal terga II-VII with laterobasal

creamy-brown scaled patches (terga vary from completely dark brown scaled

to creamy-brown scaled patches present, some specimens with laterobasal

pale scaled patches reduced or absent on II-V), scutellum with narrow curved

dark brown scales on each lobe. The female genitalia of dux, lugubris and

iriomotensis are similar in the development of the spermathecal eminence,

the later species being somewhat intermediate in development between the

other 2 species.

Reinert: New Interpretation of Verrallina 27

The description and illustration of butlevi male genitalia by Edwards in

Barraud (1934: 296) belong to lugubris, as suggested by Laffoon (1946: 244),

is here confirmed.

Laffoon (1946: 244) and Huang (1968: 17) report the male midleg posttarsus

of butleri as having the ungues unequal in size and both simple. I have examined

specimens from Malaysia (type-locality), Philippine Islands, Singapore and

Thailand and find the midleg posttarsus to always be unequal in size with the

larger ungue simple and the smaller ungue with a tooth. The tooth on the

smaller ungue is small and may have been overlooked by previous workers.

The specimens from Brunei represent a new record for that country.

TYPE-DATA. The lectotype female of butlerz in the BMNH bears the

following information on the labels: Klang jungle, Sept./ on the underside of a

paper circle on which the adult is pinned]; 28.10.99, Selangor, A. L. Butler;

K. Selangor (mangroves) swarming; Aedes Butleri (Type) Theobald; Aédes

butleri Theobald, Hololectotype, P. F. Mattingly, 26: viii: 1955. There isa

descrepency in the above data in the month of collection (given as Sept. on one

label and as Oct. on another label). Aedes butleri was first described by

Theobald (1901: 230) from 3 female syntypes. Knight and Hull (1953: 476)

noted one of the females was not butlervz but probably a species in the subgenus

Skusea Theobald. Mattingly (1956: 794) selected the above lectotype; a para-

type with the following data: 28.10.99, Selangor, A. L. Butler, Klang, very

common & troublesome; and labeled the third specimen as a paratype (label

data as follows: 28.10.99, Selangor, A. L. Butler) and transferred it to

Skusea, confirming the identification of Knight and Hull.

DISTRIBUTION. 368 specimens examined: 202°, 71°, 2p, 41, 28L,

34 with associated skins (7 p, 271).

BRUNEI. Temburong District; 42 pl, 272, 1°, 12 L.

INDONESIA. Borneo (Southeast), Tanah Grogot; 32, 1° (paratype of umbro-

sis). Celebes, Loemae;1¢. Cevam;12. Java, Batavia, Kemajoran; 99.

MALAYSIA. Kedah, Kg. Sungei Lintang; 12 pl, 32. Pevak, Maxwell's

Hill; 42. Perlis, Kg. Gunong; 22 pl. Sabah, Beaufort, Jesselton, Kuala Abai,

Kudat, Labuan, Sandakan, Tawau, Tuaran; 139, 20°. Selangor, Banting, Carey

Island, Kampong Bandar, Klang, Kuala Lumpur, Kuala Selangor, Rantau Pan-

jang, Sg. Bertek, Telak Gong; 12 p, 389, 19 (lectotype of butleri), 2% pl, 5c

bp, 4c.

PHILIPPINES. Basbas Island; 12. Culion Island; 12, 3L, 11. Leyte,

Palo, Puerto Princesa, Tacloban; 6 pl, 539, 1° pl, 41c’, 2p, 7h, 31. Pala-

wan Island, Iwahig Penal Colony, Tinabog; 2°.

SINGAPORE. Macritchie Reservoir; 12. West coast road between 8 and 9

miles; 42 pl, 172, 4% pl, 1% p, 40°, 5L.

SRI LANKA. Kalutava District, Matugama; 19 (cotype of carteri). Southern

Province, Potwatte Ganga at Denipitiya; 12 (specimen of cartevi).

THAILAND. Chanthaburi, Ban Bo Phu; 1. Ranong, Khao Muang Lak,

Muang; 32 pl, 22, 80°, 1 L. Udorn Thani, Nong Bau; 18.

BIOLOGY. In Malaysia immatures were collected from a ground pool, hoof-

prints along coast, small ditch, ground pool surrounded by nipa palms, small

marshy depressions; from clear, colored and turbid, fresh or brackish, un-

moving, temporary water in partially shaded areas located in secondary man-

groves, costal villages, villages with coconut palms and an orchard plantation;

and at elevations from sea level to 45 m.

Immatures in Thailand were collected from small and large ground pools

containing clear, fresh or brackish, unmoving, temporary water in partially or

heavily shaded areas located in primary mangroves and at elevations from 3-

28 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

26 m.

Immatures from Brunei were collected from a small ground pool containing

clear, fresh, unmoving, temporary water in a heavily shaded area in a village

and at sea level.

Collections of immatures from the Philippines were made from mangrove

swamps, swamp rain pools, pool surrounded by nipa palms and a puddle with

leaves; from brackish and fresh, unmoving water in partially shaded areas;

and at elevations at and near sea level.

In Singapore immatures were collected from small ground pools and a ground

pool at base of a coconut palm; from colored, brackish, unmoving, temporary

water in partially shaded areas located in an open swamp and a costal village;

and at sea level.

Adults have been collected biting man in costal villages, a secondary ever-

green forest and a rain forest in Singapore. In the Philippines adults were

taken biting man in mangrove Swamps and coconut groves. Adults were col-

lected in Malaysia biting man in mangroves, coconut groves, secondary rain

forests and a village with nipa palms.

AEDES (VERRALLINA) CAMPY LOSTY LUS LAFFOON

(Figs. 23, 58, 60)

DISCUSSION. The following specimen from New Guinea is the first record

of this species outside of the Philippine Islands. This female bears the follow-

ing information on the labels: NETH. NEW GUINEA, Hollandia, I: 2: 1945,

Lot # 496-G, adults biting Sago swamp, 19th Med. Gen. Lab. U. S. Army,

L. W. Saylor collector, genitalia preparation number T 73.463.

Structures of campylostylus illustrated for the first time are: male post-

tarsi (Fig. 58), female posttarsi (Fig. 60) and tergum IX of the female geni-

talia (Fig. 23).

DISTRIBUTION. 82 specimens examined: 342, 260°, 2p, 21, 6L, 6 with

associated pupal and larval skins.

INDONESIA. New Guinea, West Irian, Hollandia; 192.

PHILIPPINE ISLANDS. Balabac Island, Cape Melville Te bo, 4, (para-

types). Mindanao, San Ramon; 12, 50, 3 L (paratypes). Mindoro, San Jose;

1% (paratype). Palawan, Balsahan River, Irahuan River; 1° pl (holotype), 42

pl, 199, 120°, 2 pl, 2 L (paratypes), Quezon, Panitian; 22, 1° pl, 1¢.

BIOLOGY. The new material from Palawan was collected as immatures

from a ground pool (1° pl) and a rock pool (22, 1° skins lost).

The female from Hollandia was collected biting man in a Sago swamp.

AEDES (VERRALLINA) CARMENTI EDWARDS

(Figs. 5, 27, 38, 50, 58, 60)

Aedes (Aedes) ceramensis Brug 1934, Bull. Entomol. Res. 24: 512 (¢ *).

NEW SYNONYM.

Aedes carmenti is the representative species for Series I, Section A, and

is therefore completely redescribed below.

FEMALE. Head. Antenna dark brown, 0. 76-0. 83 length of proboscis,

pedicel dark brown with a few small dark scales and short fine hairs mesally,

flagellomere 1 with basal 0.3 pale and with several small dark scales near

Reinert: New Interpretation of Vervallina 29

middle, flagellar whorls with 6 dark setae; clypeus dark brown, bare; maxillary

palpus blackish-brown scaled, 4 segmented, 0.11-0.13 length of proboscis;

proboscis blackish-brown scaled, 1.50-1.55 length of femur I; eyes nearly con-

tiguous; ocular setae blackish-brown and well developed; scales on head all

broad blackish-brown and decumbent except a row of narrow curved pale brown

scales on ocular line, a narrow stripe of broad white scales on lateral surface

extending from ocular line to a point anterior of antepronotum, and numerous

moderately long dark brown erect forked scales on occiput. Thorax, Scutal

integument dark brown; scutum covered with narrow curved reddish-black

scales except for narrow curved golden-white scales forming a small patch on

median anterior promontory area, anterior scutal fossal area, supra-alar area

near base of wing and along anterior and lateral margins of prescutellar space;

prescutellar space bare; scutellum with a patch of narrow curved reddish-black

scales on each lobe; dark blackish-brown setae on the following areas: 4-5

median anterior promontory, numerous acrostichal (anterior and posterior),

numerous dorsocentral (anterior and posterior), scutal fossal (3-6 anterior,

6-7 lateral and 2 posterior), numerous supra-alar, 7-8 posterior medial scutal,

1 postalar callar and scutellar (8-11 long and 4-6 short ones on lateral lobe,

6-10 long and 5-7 short ones on median lobe); pleural integument dark reddish-

brown; antepronota widely separated, scales absent, 18-28 long and short dark

setae; postpronotum with narrow curved reddish-black scales on upper area

and a few moderately broad brown ones below, 6-10 long and short dark

posterior setae and a patch of short hairs ventrad of scale patch and cephalad

of posterior setae; propleuron with a patch of broad white scales, 15-19 long

and short dark setae; prosternum, subspiracular area, paratergite, mesomeron

and metameron bare; postspiracular area with 6-12 setae; mesepisternum with

an upper patch of broad pale brown scales (a few posterior ones white) and a

lower patch of broad white scales, 4-7 upper and 15-19 posterior dark setae,

a patch of short fine golden hairs cephalad of upper scale patch near subspiracu-

lar area; prealar knob with 11-15 dark setae; mesepimeron with a patch of

broad white scales near center, 21-31 setae dorsad of scale patch, several

brown hairs along posterior and ventral margins of scale patch becoming more

numerous posteroventrally but not extending onto lower 0.25 of sclerite. Legs,

Coxae I-III each with several dark setae, I with anterior surface with broad

brown scales and a small dorsal and a ventral patch of white scales, II and HI

each with a patch of broad white scales on outer area of anterior surface; tro-

chanters I-III. each with broad pale brown scales; femora I-II each with anterior

surface dark brown scaled, III with anterior surface white scaled with a dorsal

longitudinal stripe of dark brown scales extending from base to apex and forming

a narrow apical band, I with posterior surface pale scaled with a ventral longi-

tudinal stripe of dark brown scales extending from base to apex, IJ and III each

with posterior surface pale scaled with a narrow dorsal longitudinal stripe of

dark brown scales extending from base to apex, stripe broader distally and

forms a band on apical 0.12-0.20; tibiae I-III and tarsi I-III each blackish-brown

scaled; posttarsi I-III (Fig. 60) each with 2 ungues, I with ungues equal, each

with a tooth, II with ungues equal, one with a tooth, III with ungues equal, both

simple. Wing. Dorsal and ventral veins with blackish-brown scales; alula with

several narrow brown scales along margin; upper calypter with several pale

hairs on margin; 1-2 remigial setae. Halter. Pedicel pale; capitellum brown

scaled with pale scales dorsally. Abdomen, Terga blackish-brown scaled,

laterotergite of I with a rectangular patch of white scales, II-VII each with a

lateral white scaled spot, extreme lateral portion of spot basal; sterna dark

brown scaled, I[-VII each with a laterobasal white scaled spot; terga and sterna

30 Contrib. Amer. Ent, Inst.; vol./11, no. 1, 1974

with numerous golden setae, mostly along posterior margins. Genitalia (Fig.

d). Tergum VII moderately pigmented, wide, base and apex each slightly

concave, several short and moderately long setae on apical 0.46-0.60, a few

long setae along apical margin, basolateral seta absent but a small bulla in

this location, numerous broad scales covering apical 0.74-0. 83, covered with

minute spicules, basal 0.3-0.5 retracted into segment VII, VIII-Te index

0.63-0.70, VIII-Te/IX-Te index 3. 68-4. 09; sternum VIII moderately pigmented,

wide, base slightly concave, apex with a moderately deep median indentation

with a very small lobe on each side of midline, numerous short and moderately

long setae scattered over apical 0. 86-0. 83, covered with minute spicules,

numerous broad scales covering most of apical 0.90-0.94, apical interseg-

mental fold unpigmented, VIII-S index 0. 75-0. 80; tergum IX moderately pig-

mented, wide, short, band-like, base with a short median indentation, setae

absent, covered with minute spicules, [X-Te index 0.33-0. 39; insula unpig-

mented, short, covered with minute spicules, 4-5 small tuberculi, each with

a minute spicule; lower vaginal lip moderately pigmented, narrow, covered

with minute spicules, lower vaginal sclerite composed of a pair of very lightly

pigmented small plates, without spicules and somewhat wrinkled; upper vaginal

lip lightly to moderately pigmented, moderately broad laterally and narrow

posteriorly, posterolateral margin evenly rounded and not lobed, minute

spicules on posterior margin, upper vaginal sclerite moderately pigmented,

well developed, base attached to basal 0.5 of lateral area of upper vaginal lip,

sclerite bifurcate with anterior branch small, lightly pigmented and attached

to cephalic portion of spermathecal eminence, posterior branch wide with a

caudally produced, lightly pigmented, mesally curved arm connected with its

mate and forming a crown around the spermathecal eminence; spermathecal

eminence heavily pigmented, shallow in depth, composed of a pair of comma

shaped lateral structures connected caudally and produced into a fleshy very

lightly pigmented small cephalic projection, spiny excrescences absent; post-

genital lobe short, wide, apex broad with a moderately deep median indenta-

tion, 11-15 setae on each side of midline, completely covered with minute

spicules, dorsal PGL index 0. 88-1. 08; peri-anal membrane with minute

spicules; cercus triangular in shape, moderately long and broad at base,

apex acute, base concave, completely covered with small spicules, dorsal

surface with a number of short and moderately long setae on apical 0. 84-0. 90

and a few (1-9) broad scales, a few long setae at apex, ventral surface with a

few short and moderately long setae along outer margin and apical area,

cercus index 2.64-2.97, cercus/dorsal PGL index 3. 46-3. 84; 3 seminal cap-

sules, 1 large, 1 medium and 1 small in size, heavily pigmented, spherical,

each with a short narrow heavily pigmented neck, a few tiny seminal capsule

pores near orifice, base of accessory gland duct moderately pigmented.

MALE. Similar to female in general habitus but with the following differ-

ences. Head, Antenna plumose, 0. 76-0. 84 length of proboscis; maxillary

palpus 0.11-0.12 length of proboscis; proboscis 1.39-1.56 length of femur I.

Thorax. Median anterior promontory with 3-4 setae; scutal fossal setae (2-5

anterior, 2-4 lateral and 1 posterior); 6 posterior medial scutal setae; 14-2]

antepronotal setae; 4-6 postpronotal setae; 14-18 propleural setae; 5-7 post-

spiracular setae; 8-10 prealar knob setae; mesepimeron with 14-18 setae

dorsad of scale patch. Legs. Posttarsi I-III (Fig. 58) each with 2 ungues, I

and II with ungues unequal, each with a tooth, III with ungues equal, both

simple. Genitalia (Fig. 27). Tergum IX moderately pigmented, caudal

margin concave mesally, strongly fused to tergum X, setae absent, narrowly

connected laterally to sternum IX; gonocoxite short, broad, heavily pigmented,

Reinert: New Interpretation of Verrallina 31

mesal surface membranous with minute spicules, dorsal and ventral surfaces

both with a number of short and a few moderately long setae scattered over

most of surfaces, a few long stout setae on apical 0.6 of outer margin of dor-

sal surface and on lateral surface, ventral surface also with a short heavily

pigmented spiniform on apicomesal area, this seta arises from a short tuber-

cule, a short stout seta basad of tubercule and a moderately long seta distad of

tubercule, several broad scales on basal area becoming more numerous lateral-

ly and extending onto lateral surface; gonostylus moderately pigmented, moder-

ately long (approximately 0.68 length of gonocoxite), basal 0.4 expanded and

bearing 10-12 short setae, apical 0.6 narrow, curved and tapered to a point,

apex slightly recurved; basal mesal lobe lightly to moderately pigmented, con-

sists of a somewhat triangular plate situated on basal mesal area of gonocoxite

and bears 7-13 short setae and a small apical lobe bearing 8-11 short setae,

sternal margin of plate strongly fused to basomesal margin of ventral surface

of gonocoxite, basal mesal lobe connected mesally to its mate by a narrow

moderately pigmented band covered with minute spicules and located ventrad

of prosophallus and phallus; proctiger continuous tergally, moderately pig-

mented, paraproct indistinct, not separated or free, base with a small short

sternal projection which articulates at a point on the dorsomesal surface of

the parameral apodeme near base of opisthophallus, cercal setae absent;

tergum X moderately pigmented, strongly fused to median caudal margin of

tergum IX and base of proctiger, margins indistinct but fusion lines somewhat

wrinkled; phallosome complex, opisthophallus--consists of a moderately pig-

mented tergal narrow transverse bridge between the parameral apodemes,

lateral surface curved ventrally, caudal margin slightly concave to truncate,

base attached to a dorsomesal extension of the caudal portion of parameral

apodeme, located dorsad of phallus and prosophallus and ventrad of proctiger,

phallus--consists of a pair of moderately long heavily pigmented arms which

are joined near their center by a narrow sternal bridge, apices of arms con-

tiguous giving an A-shaped appearance, base of arm attached to inner tip of

paramere, tergal area of arm (basad of sternal bridge) strongly fused to inner

sternobasal portion of prosophallus, apex of phallus with a loosely attached

tergal penis filament which consists of a large moderately pigmented U-

shaped structure and is produced cephalad, prosophallus--consists of a bulbous

base which is fused to tergobasal area of phallus and tergomesal portion of

paramere, apical 0.5-0.6 produced into several lightly pigmented flattened

contiguous leaf-like structures with apices fused into a single plate which ex-

tends caudad of posterior margin of opisthophallus, base is positioned be-

tween laterosternal surface of opisthophallus and lateral margin of phallus,

prosophallic arm approximately equal in length to phallic arm, paramere--

heavily pigmented, short, approximately 0.46 length of phallic arm, articu-

lates at a point near middle of outer margin with caudal area of parameral

apodeme, parameral apodeme--moderately pigmented, moderately long, outer

surface broadly attached to tergobasal apodeme of gonocoxite; sternum IX

moderately pigmented throughout,: large, broad, apicolateral areas rounded,

9-12 short and moderately long setae in an enlongate patch near middle of

caudal margin.

PUPA (Fig. 38). Chaetotaxy as recorded in Table 4. Ocular plate with

area of moderately developed cuticular facets. Respivatory trumpet. Heavily

pigmented; hair-like spicules on distal 0. 54-0. 60 of inner surface; index 3.19-

4,28, average 3.81. Abdomen. Setae 6, 9-VII weakly developed; 4-VIII double

to 4 branched; 9-VIII triple to 6 branched. Paddle. Ovoid; minute serrations

on distal portion of basal 0.57 of outer margin; minute spicules on apical 0. 43

32 | Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

of outer and apical 0.3 of inner margins; midrib does not reach apex; seta 1-P

moderately long, single or double; index 1.14-1.55, average 1.38.

LARVA (Fig. 50). Chaetotaxy as figured and recorded in Table 17. Head.

Seta 4-C with 6-10 branches; 5-C triple to 7 branched; 6-C triple; 7-C with 8-

11 branches; 12-C triple to 6 branched; dorsomentum with 36-41 teeth. Anten-

na. Moderately long, lightly to moderately pigmented, with small spicules

scattered over shaft, more numerous on basal area; seta 1-A triple to 5

branched; 2-A long, with a subapical constriction; 3-A short, 0.20-0.31 length

of 2-A; 4-A moderately long, 0.57-0. 74 length of 2-A; 6-A short, 0.27-0.35

length of 2-A. Abdomen. Seta 1-VIII with 5-8 branches; 2-X triple, one

branch longer than other 2; ventral brush with 10 setae on grid and 2 precratal

shorter ones, all multiple branched; saddle moderately pigmented, incompletely

rings segment X, acus absent; 4 moderately long anal papillae; comb with 10-

14 (usually 12-14) scales arranged in a single curved irregular row, each

scale with small uniformly developed denticles on lateral and apical areas.

Siphon. Moderately pigmented; acus well developed; index 3.00-3.49, aver-

age 3.29; pecten on basal 0. 59-0. 64 of siphon, composed of 12-16 (usually

13-15) teeth, apical 3 teeth longer and wider spaced than remainder of teeth,

each tooth long, slender and with 1-2 small stout ventral denticles near base,

distal 2-4 teeth with or without denticles; seta 1-S short, double to 4 branched,

base attached on basal 0. 70-0. 72 of siphon distad of last pecten tooth.

DISCUSSION. Adults of carvmenti show considerable variation in habitus

and are superficially very similar to a number of other species of Series I,

Section A. Characters for separating this species from related forms are

given in a key by Huang (1968). The male genitalia of carmenti and embiensis

appear to be identical (see discussion section of embiensis).

Upon comparison of the 2 cotype males of cevamensis with the lectotype

and numerous specimens of carmenti from Guadalcanal, Solomon Islands, I

found no difference in the male habitus or genitalia and therefore consider

them conspecific. Aedes ceramensis was described as a new species by Brug

(1934: 512) from 2 males collected from Ceram. King and Hoogstraal (1947:

115) synonymized cevamensis with carmenti; however, Belkin (1962: 417)

considered the 2 to be distinct species. Huang (1968: 22) followed Belkin's

determination of the species. The figure of the male genitalia by Brug (1934:

512) is a distorted view, especially of the basal mesal lobe. Since Belkin

apparently did not examine the cotype specimens of cevamensis and relied

only on the published figure he-was mislead into believing they were different

from carmenti.

The female genitalia of carmenti and the other members of Series I, Sec-

tion A, are very uniform in development and many species do not display any

consistant differences with which to separate them; however, the shape of ter-

gum IX is useful in distinguishing a number of species. Figure 24 illustrates

the difference in shape of tergum IX of several species in Series I.

Tarsomeres 5 and posttarsi of several species of Series I are illustrated

(Figs. 58, 59, 60, 61, 62) for the first time and can be compared to other

members of the series.

Larvae of carmenti and most other members of Series I (funerus, mccor-

micki, multifolium, parasimilis, sentanius and similis) have setae 2-X with

one branch noticeably longer than the other branches. Three species (/foli-

formis, lineatus and variabilis), however, have the branches of setae 2-X

approximately equal in length, a feature common to members of Series II and

III. Seta 2-X is single in cuccioi and double in trispinatus.

TYPE-DATA. Type female in BMNH bears the following information on

Reinert: New Interpretation of Vervallina 33

the labels: Aedes (Aedes) carmenti Edw., Type H. T., Solomon Is.: Guadal-

canal I., Dr. A. G. Carment, B. M. 1923-610. I have examined the type and

it has the abdomen and genitalia missing and is pinned with a minuten pin

through the venter of the thorax and extended through the scutum.

Edwards (1924: 388) did not report the designation of a type in his original

description of carmenti but he recorded numerous females taken in brush and in

native houses by Dr. A. G. Carment at Maravovo, Guadalcanar| sic] I., Solo-

mon Is.,vi. 1923 and one female collected by G. F. Hill on Palm Island, N.

Queensland. King and Hoogstraal (1947: 115) reported the following for carmenti

"Type female, with numerous other specimens, from Maravova, Guadalcanal,

Solomon islands, and one female from Palm Island, North Queensland, Austra-

lia. '’ Belkin (1962: 416) recorded the following for this species "TYPE: lect-

type £, marked as type by Edwards, Maravove, Guadalcanal, Solomon Islands,

taken in brush or native house, June 1923, A. G. Carment; by present selec-

tion (BMNH)."' Huang (1968: 18) mistakenly took the statement by King and

Hoogstraal (1947) as a lectotype designation.

Lectotype designation is here made for one of the cotype males of ceramen-

sis and it bears the following information on the adult labels: Ceram, Oewin,

3.1.1932, De Rook [ collector], B.M. 1934-45, 2697 [ collection number], co-

type, Larvae in swamp of virgin forest, Aédes (Aédes) ceramensis Brug, LEC-

TOTYPE & selected by J. F. Reinert, July 1974. Paralectotype designation is

made for the other cotype male which possesses the following label data: Ceram,

Warasiwa, Brug & De Rook [ collectors], 2696 [ collection number], cotype.

The lectotype is in good condition but it has the 3 left legs and the left wing mis-

sing. One wing, a thoracic fragment and genitalia remain of the paralectotype.

The genitalia are mounted on small points which are attached to the adult pins,

those of the lectotype are in good condition but those of the paralectotype are

smashed and distorted and were probably used by Brug in making his Figure 8b.

Both types are in the BMNH.

DISTRIBUTION. 415 specimens examined: 3229, 570°, 3p, 61, 13 L,

7 with associated pupal and larval skins.

AUSTRALIA (and Trust Territories). Admiralty Islands, Los Negros; 662.

Bougainville Island (Solomon Islands), Empress, Augusta Bay; 432, 180, 5 L.

New Britain, Cape Hoskins; 292. New Guinea, Papua, Ladova, Milne Bay area;

139, 30%, 2 pl. Queensland, Palm Island; 19.

INDONESIA. Ceram, Oewin, Warasiwa; 2 (lectotype and paralectotype of

cevamensis). New Guinea, Hollandia, Lake Sentani, Tor River; 6@ pl, 419,

ae 3S, BAO Te,

SOLOMON ISLANDS (British Protectorate). Guadalcanal Island, Belasuma

River, Koli Ar., Teneru Ar.; 1012, 17, 1 pl. New George, Munda; 21°, 12

(lectotype), 130°, 2 L.

BIOLOGY. Immatures were collected from water ina grassy road rut in

New Guinea.

In the Solomons the immatures are apparently adapted to breeding in

shallow, heavily shaded, jungle pools which are frequently flushed (Belkin

1962: 417). Additional information on the biology of this species in the Solo-

mons is given by Belkin (1962: 417) and in New Guinea by Huang (1968: 21).

Females in the Solomons attacked man voraciously in forested areas during

the daytime (Belkin 1962: 418). Similar feeding habits of the females were

observed by Huang (1968: 21) for this species in New Guinea.

34 Contrib; Amer. Ent. Inst. ,° vol. 11, no..1; 1974

AEDES (VERRALLINA) CAUTUS BARRAUD

(Fig. 6)

The following information is based on the lectotype male, paratype (cotype)

female and one other paratype female and supplements the descriptions of

Barraud (1928: 368, 1934: 288).

FEMALE. Head, Antenna 1.08 length of proboscis, pedicel brown with a

few small brown scales and short fine brown hairs mesally; maxillary palpus

0.18 length of proboscis; proboscis 0.98 length of femur I; eyes contiguous in

front. Thorax. Scutum covered with narrow curved reddish-black scales ex-

cept for small patches of narrow curved creamy-white scales on median an-

terior promontory area, anterior scutal fossal area and on supra-alar area

above paratergite; scutellum with narrow curved reddish-black scales on each

lobe in specimen labeled cotype and a few creamy-white scales mixed with

dark ones in specimen labeled paratype; antepronotum with scales absent,

12-13 dark setae; postpronotum with narrow curved reddish-black scales on

upper area, 4-7 dark posterior setae; propleuron with a patch of broad white

scales, 9-10 setae; prosternum, subspiracular area, paratergite, meso-

meron and metameron bare; postspiracular area with scales absent, 9-6 dark

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

scales, 4 upper and 9-10 posterior setae, lower ones shorter; mesepimeron

with a large patch of broad white scales near center, 7-10 golden setae dorsad

of scale patch, 2-4 small golden hairs near lower posterior corner of scale

patch. Abdomen. Sterna white scaled with a few apical brown scales.

MALE. Head, Antenna 0.88 length of proboscis; maxillary palpus 0.11

length of proboscis; proboscis 1.02 length of femur I.

DISCUSSION. ‘The adult habitus of cautus is very similar to pseudomedio-

fasciatus and sabahensis, new species. There appears to be no consistant

differences between the habitus of the adults of the former 2 species; however,

distinctive differences exist between the genitalia of both males and females

(see discussion section of sabahensis for separation of it from cautus and

pseudomediofasciatus). |

The above additions to the descriptions of Barraud (1928, 1934) are given

because the female description of Delfinado (1967: 13) was based on a mixture

of females of cautus and sabahensis. Her illustration (Fig. 18) of cautus is

also that of sabahensis. The present illustration (Fig. 6) was drawn from the

female specimen labeled cotype of cautus, Delfinado (1967: 13) reported this

species from Thailand and North Borneo; however, after examination of these

specimens the Thailand females were found to be a mixture of andamanensis

Group, atrius, vallistris and varietas andthe North Borneo specimen was saba-

hensis. No material of cautus from Thailand, Malaysia or North Borneo was

found in the USNM, SEAMP or BMNH collections. This species is probably

restricted to the Indian area. :

DISTRIBUTION. 3 specimens examined: 22, 1.

INDIA. Bombay Deccan, Tavargatti; 12 (cotype), 12 (paratype), 1% (lecto-

type).

AEDES (VERRALLINA) CLAVATUS BARRAUD

(Figs? 235.39, 58° 60)

PUPA (Fig. 39). The following description and Table 5 are the first de-

scription of the pupa of this species. Pupal skins from females have heavily

Reinert: New Interpretation of Verrallina 20

pigmented areas on the cephalothorax, abdominal segments and base of paddles,

while those of males lack heavily pigmented areas on the abdominal segments.

Respiratory trumpet. Moderately pigmented; index 3.21-3.88, average 3.61.

Abdomen. Setae 6, 9-VII weakly developed; 4-VIII double or triple; 9-VIII

double to 4 branched. Paddle. Ovoid; minute serrations on distal portion of

basal 0.6 of outer margin; minute spicules on apical 0. 4 of outer and apical

0.28 of inner margins; midrib does not reach apex; seta 1-P moderately long,

single; index 1.32-1.58, average 1.4.

DISCUSSION. The male genitalia are very distinctive in having the proc-

tiger divided into a long stout caudally produced tergal arm and a moderately

long stout mesally produced sternal arm which is directed over the lateral por-

tion of the phallosome and is difficult to see in an undissected preparation.

The proctiger articulates with the parameral apodeme at a point on the base of

the sternal arm.

The following structures of clavatus are illustrated for the first time:

male posttarsi (Fig. 58), female posttarsi (Fig. 60) and tergum IX of the fe-

male genitalia (Fig. 23). Tergum IX of thefemale genitalia occasionally with

a single short seta on one side.

Additional information on the distribution and biology of this species is

listed below.

DISTRIBUTION. 33 specimens examined: 17°, 6, 10 with associated

pupal skins.

CAMBODIA. Kompong Sela, Stung Chral; 49.

SOUTH VIETNAM. Kon Tum; 22. Pleiku, Plei Djereng; 29.

THAILAND. Chon Buri, Amphoe Bang Lamung, Khao Mai Keo, Khao Wai;

12, 20°. Nakhon Ratchasima, Khlong Phai, Pak Thong Chai; 42 p, 1% p.

Nakhon Si Thammarat, Amphoe Chawang, Khao Luang, Ban Tha Phon; 18.

Ranong, Khao Chatri; 1“ p. Satum, Khlong Baraket; 22 p, 2% p. Trang, Am-

phoe Muang, Tong Ka Beau; 18.

BIOLOGY. In Thailand immatures were collected from crab holes, a

flood pool and stream pools in dry streams (filled with rain water); from

clear or colored, fresh, unmoving, temporary water in heavily shaded areas;

located in mountainous and valley terrain, in primary and secondary rain

forests; and at elevations from 75-540 m (most often at 75-80 m). Adults

were taken in biting collections between 1100-1200 hours and in light traps.

In South Vietnam adults were collected from light traps.

AEDES (VERRALLINA) COMATUS BARRAUD

(Fig. 3)

The following information is based on the holotype female and supplements

the original description of Barraud (1931: 615).

FEMALE. Head, Antenna 0.94 length of proboscis; maxillary palpus

0.18 length of proboscis; proboscis 1.22 length of femur I. Thovax. Follow-

ing setae present: 4 median anterior promontory; numerous acrostichals

(anterior and posterior); numerous dorsocentrals (anterior and posterior);

scutal fossal (5 anterior, 4-5 lateral and 1-2 posterior); numerous supra-

alar; 6 posterior medial scutal; 1 postalar callar; scutellar (4 or 6 long and

3-4 short ones on lateral lobe, 6 long and 4 short ones on median lobe); 15 or

17 antepronotal; 5-6 postpronotal; 20 propleural; 5 or 8 postspiracular; 5 upper

and 15 or 19 posterior mesepisternal; 7 on prealar knob; and mesepimeron

with 18-19 dorsad of scale patch and numerous hairs caudad of and below scales.

36 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

Prosternum, subspiracular area, paratergite, mesomeron and metameron

bare. Wing. Remigium with 1 brown seta. Genitalia (Fig. 3). Distinctive,

as illustrated. |

DISCUSSION. The female genitalia of comatus are distinctive for the

species. The genitalia are illustrated, the lower vaginal lip and sclerites for

the first time.

DISTRIBUTION. Known only from the holotype and type-locality.

AEDES (VERRALLINA) COMOSUS NEW SPECIES

(Fig. 7)

FEMALE. Head, Antenna dark brown, 0.87 length of proboscis, pedicel

dark brown with a few small dark scales and short fine brown hairs mesally,

flagellomere 1 with basal 0.23 pale and with several small brown scales near

middle; clypeus dark brown, bare; maxillary palpus dark brown scaled, 0.2

length of proboscis; proboscis dark brown scaled, 1.18 length of femur I;

eyes contiguous; ocular setae blackish-brown and well developed; scales on

head all broad, blackish-brown and decumbent except for a few narrow

curved white scales along coronal suture and on upper area of interocular

space, broad white scales on occiput, similar scales forming a stripe on area

in front of antepronotum and extending to ocular line, a few broad pale scales

on postgena, and a number of dark brown erect forked scales on occiput.

Thorax. Scutal integument dark reddish-brown; scutum somewhat rubbed

mesally, but otherwise covered with narrow curved dark reddish-brown

scales except for narrow curved golden scales on the following: small

patch on median anterior promontory area, small patch on anterior scutal

fossal area, patch on supra-alar area and a small patch on posterior medial

scutal area; scutellum with a patch of narrow curved golden-brown scales on

each lobe; prescutellar space bare; dark reddish-black setae on the following

areas: 5 median anterior promontory, numerous acrostichal (anterior and

posterior), numerous dorsocentral (anterior and posterior), scutal fossal

(6-8 anterior, 4 lateral and 1 posterior), numerous supra-alar, 4 posterior

medial scutal, 1 postalar callar and scutellar (4-5 long and 3-4 short ones

on lateral lobe, 5 long and 4 short ones on median lobe); pleural integument

dark reddish-brown; antepronota widely separated, scales absent, 10-13 long

and short dark setae; postpronotum with a few narrow curved reddish-brown

scales on upper area and a few broad reddish-brown ones below, 4-5 dark

posterior setae, a few short fine brown hairs cephalad of setae; propleuron

with a patch of broad white scales, 23-25 setae; prosternum, subspiracular

area, paratergite, mesomeron and metameron bare; postspiracular area

without scales, 6 brown setae; mesepisternum with a large upper and a small

lower patch of broad white scales (a few scales in upper patch with a brown

tinge), 4 upper and 8-10 posterior setae, several short fine brown hairs

cephalad of upper scale patch and others ventrad of lower scale patch; prealar

knob without scales, 8 brown setae; mesepimeron with a patch of broad white

scales near center, 16 brown setae dorsad of scale patch, numerous short

fine hairs caudad of and below scale patch but not extending onto lower 0.25

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

scales and a small dorsal patch of white ones on anterior surface, II and III

each with a patch of broad white scales on anterior surface; trochanters I-III

each with several pale brown scales; femora I and II each with anterior surface

brown scaled, posterior surfaces brown scaled with a longitudinal stripe of

Reinert: New Interpretation of Vervrallina 37

white scales, stripe dorsal on I and ventral on II, III with anterior and posteri-

or surfaces brown scaled both with a ventral longitudinal white scaled stripe

extending from base to near apex, II and III each with a few lateroapical white

scales on anterior surface; tibiae I-III and tarsi I-III each dark brown scaled;

posttarsi I-III (Fig. 7) each with 2 ungues, I and II each with ungues equal,

each with a tooth, III with ungues equal, both simple. Wing. Dorsal and ventral

veins with dark brown scales; alula with a few narrow brown scales on margin;

upper calypter with several brown setae on margin; 2 remigial setae. Halter.

Pedicel pale; capitellum with pale brown scales. Abdomen. Terga dark brown

scaled, I-VI each with a large laterobasal patch of white scales (remainder of

abdomen removed with genitalia); sterna each white scaled with an apical band

of brown scales; terga and sterna with several brown setae, mostly along

posterior margins. Genitalia (Fig. 7). Tergum VIII moderately pigmented,

wide, base slightly concave, apex convex, a number of short and moderately

long thin setae at apex and similar ones scattered over apical 0.36, basolateral

seta present, covered with minute spicules, numerous broad scales covering

apical 0.75, basal 0.45 retracted into segment VII, VIII-Te index 0.57, VIII-

Te/IX-Te index 2.98; sternum VII moderately pigmented, wide, base concave

mesally, apex with a shallow median indentation with a small lobe on each

side of midline, numerous moderately long setae apically, more numerous

and slightly stouter on apical lobes, numerous short and a few moderately

long setae scattered over apical 0.93, covered with minute spicules, numerous

broad scales covering much of apical 0.8, apical intersegmental fold unpig-

mented, VII-S index 0.65; tergum IX moderately pigmented, wide, short,

base with median area concave, apical area straight, setae absent, covered

with minute spicules, [IX-Te index 0.32; insula unpigmented, short, covered

with minute spicules, 5 small tuberculi, each with a tiny spicule; lower vaginal

lip heavily pigmented, narrow, a few spicules on outer margin and on hinge

area, with a small caudally projecting moderately pigmented median apodeme,

lower vaginal sclerite composed of a pair of moderately long and moderately

broad heavily pigmented plates covered with minute spicules and situated on

median basal area of the vaginal floor; upper vaginal lip heavily pigmented,

complex, wide, posterior margin formed into a large upright shield caudad of

spermathecal eminence, posterolateral corners formed into sharply rounded

lobes, upper vaginal sclerite heavily pigmented, extremely large and complex,

base attached along entire lateral margin of upper vaginal lip, median lateral

area formed into a large scoop-like structure, a second plate caudad and

extends around the posterior of the spermathecal eminence, moderately to

heavily pigmented and broad basally; spermathecal eminence heavily pigmented,

large, deep in depth, U-shaped in dorsal outline, numerous long spiny excre-

scences attached over entire dorsal surface except the apex; spermathecal

eminence, upper vaginal lip and upper vaginal sclerite cover nearly entire

upper vaginal area with heavily pigmented structures; postgenital lobe short,

narrow, apex with a moderately deep median indentation, 16 setae on each

side of midline, completely covered with minute spicules, dorsal PGL index

1.0, ventral PGL index 1.12; peri-anal membrane with minute spicules; cer-

cus triangular in shape, moderately long, broad at base, apex acute, complete-

ly covered with small spicules, dorsal surface with a number of moderately

long and a few short setae scattered over apical 0.9, ventral surface with

several short and moderately long setae on outer 0.5, 1-2 broad scales,

cercus index 2.95, cercus/dorsal PGL index 3.02; 3 seminal capsules, 1

large, 1 medium and 1 small in size, heavily pigmented, spherical, several

small seminal capsule pores near orifice, base of accessory gland duct un-

38 Contrib. Amer. Ent. Inst.; vol. 11, no. 1, 1974

pigmented.

MALE, PUPA, LARVA AND EGG. Not known.

DISCUSSION. Aedes comosus is similar to comatus in both adult habitus

and the female genitalia. It possesses the following combination of characters:

antepronotum with 10-13 setae, propleuron with 23-25 setae and mesepimeron

with 16 setae dorsad of scale patch which differs from comatus which has:

antepronotum with 15-17 setae, propleuron with 20 setae and mesepimeron

with 18-19 setae dorsad of scale patch.

The female genitalia of comosus differ from comatus in the shape or the

lower vaginal sclerite, development of the spermathecal eminence and the

number, position and development of the spiny excrescences on the spermathe-

cal eminence.

TYPE-DATA. Holotype female (genitalia mounted on a slide in Canada

balsam) with the following collection data: MALAYSIA, Sabah, Kota Belud,

6 April 1965, collection number 306, F. Y. Cheng collector, SEAMP acces-

sion number 35, genitalia preparation number T73.439. The holotype is

deposited in the USNM and is in good condition.

DISTRIBUTION. Known only from the type-locality.

BIOLOGY. Not known. |

AEDES (VERRALLINA) CONSONENSIS REINERT

(Figs. 28, 40, 51)

DISCUSSION. Sternum IX (Fig. 28) of the male genitalia has 17 short and

moderately long setae on the median posterior area and several small scars

basad of the setae. These scars are presumably from scales (which had

been rubbed off) since several other species in Section B possess scales on

this area.

The larvae of consonensis are somewhat aberrant from the other members

of Section B in having most setae with more branches. Antennal seta 2-A has

a subapical constriction (Fig. 51). Larval seta 18-C (see Hochman and

Reinert 1974) is small and single.

Pupae (Fig. 40) of this species are normal for Section B.

Aedes consonensis is described and compared to the other members of

Section B by Reinert (1973b).

AEDES (VERRALLINA) CRETATUS DELFINADO

(Figs. 23, 58)

DISCUSSION. Additional information taken from the collection data sheet

of the holotype is given in the type-data section below.

Biology and distribution data,for cretatus are presented. The specimens

from Malaysia represent a new record for this country.

Tergum IX of the female genitalia (Fig. 23) and tarsomeres 5 and post-

tarsi of the male (Fig. 58) are illustrated for the first time.

TYPE-DATA. The collection data sheet for the holotype male of cretatus

contains the following information: THAILAND, Khon Kaen, Tham Pholi

Yan, 26 November 1962, Sahem Esah collector, resting in heavy forest

vegetation, collection number T-1822-5. The label on the holotype specimen,

however, has printed Tham Pho Thi Yan as the collection locality. Neither

spelling of the locality could be found in the gazetteer to Thailand.

Reinert: New Interpretation of Vervallina 39

DISTRIBUTION. 71 specimens examined: 209, 16%, 3p, 9 L, 13 with

associated skins (3 p, 101).

CAMBODIA. Kompong Speu, Kirirom; 192, 2%.

MALAYSIA. Selangor, Ulu Gombak; 5? pl, 4% pl, 30°, 5 L.

THAILAND. Chiang Mai, Amphoe Muang, Huey Chang Kien; 12 (paratype).

Chon Buri, Khao Mai Keo; 1%. Khon Kaen, Tham Pholi Yan; 1“ (holotype),

1c" (paratype). Lampang, Ban Pa Khoi, Ban Pha Dang, Ban Rai Na Dieo; 1c

(paratype), 22 p, 42, 20°, 1p. Nakhon Nayok, Kaeng Saquan area, Sarica

Village, Khao Yai; 2°, 1p, 4L. Nakhon Ratchasima, Amphoe Pak Chong,

Khao Tha Kor, Khao Yai, Lam Nam Tha (along klong); 12 (allotype), 12 and

1“ (paratypes). Tak, Khao Salak Phra; 12 p and 192 (paratypes), 12 pl, 1 p.

All the above specimens are in the USNM.

BIOLOGY. Immatures in Thailand were collected from a small flood pool,

a small and a large pool in a dry stream bed, and on the margin of a stream;

from clear, fresh, unmoving, temporary water in partially shaded areas

located in secondary deciduous forests (3 times) and a primary rain forest

(once) in mountainous terrain; and at elevations from 280-700 m. In Malaysia

immatures were collected from a small ground pool containing colored, fresh,

unmoving, temporary water in a partially shaded area located in a secondary

rain forest in mountainous terrain and at an elevation of 152 m.

Adult females in Thailand have been collected biting man in forested areas at

the following times 0900-1000, 1500-1630 and 1800-1900 hours and males have

been found resting in forest vegetation.

AEDES (VERRALLINA) CYRTOLABIS EDWARDS

(Figs. 41, 52, 58)

FEMALE. The following characters are given to supplement the description

of the female by Klein (1973: 3). Thovax. Following setae present: 4-5 median

anterior promontory; numerous acrostichal (anterior and posterior); numerous

dorsocentral (anterior and posterior); scutal fossal (3-5 anterior, 3-6 lateral

and 1 posterior); numerous supra-alar; 5-6 posterior medial scutal; 1 postalar

callar; and scutellar (4-6 long and 4-6 short ones on lateral lobe, 4-6 long and

4-5 short ones on median lobe); 11-15 antepronotal; 4-6 postpronotal; 15-18

propleural; 4-7 postspiracular; mesepimeron with 4-6 upper and 11-18 posteri-

or, lower ones shorter; 13-22 on prealar knob; mesepimeron with 16-24 dorsad

of scale patch; prosternum, subspiracular area, paratergite, mesomeron and

metameron bare.

PUPA (Fig. 41). Chaetotaxy based on a single skin with associated adult.

Cephalothorax. Setae 1, 2-CT double or triple; 3-CT single or double; 4-CT

with 7 branches; 5, 7-CT with 7 or 9 branches; 6-CT double; 8-CT with 10-11

branches; 9-CT triple or 4 branched. Respiratory trumpet. Moderately pig-

mented; index 3.30. Metanotal plate. Seta 10-CT with 6 branches; 11-CT

single; 12-CT with 5-6 branches. Abdomen, Setae O-II, VI, VIII single;

0-II-VI double; 1-I with 13-16 branches on basal third; 1-II triple; 1-III with

8 branches; 1-IV with 7 branches; 1-V with 6-7 branches; 1-VI with 7-8

branches; 1-VII with 5 branches; 2-I-VII single and stout; 3-I with 7 branches;

3-II, IM single; 3-IV with 7-8 branches; 3-V with 5-6 branches; 3-VI with 5

branches; 3-VII with 5 or 8 branches; 4-I with 7-8 branches; 4-II, VI with 6-7

branches; 4-III with 6 branches; 4-IV triple or 5 branched; 4-V with 6 or 8

branches; 4-VII with 5 branches; 5-I, VII with 5 branches; 5-II with 7-8

branches; 90-III with 6-8 branches; 5-IV-VI double; 6-I triple; 6-II single; 6-II,

40 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

VIL with 6 branches; 6-IV, VI with 4 branches; 6-V with 5 branches; 7-I with 6

branches; 7-II triple or 6 branched; 7-III, IV with 5 branches; 7-V with 8 or 10

branches; 7-VI triple; 7-VII double; 8-III, IV triple or 4 branched; 8-V, VI with

4 branches; 8-VII with 4 or 6 branches; 9-I, VIII single or double; 9-II-VI single;

9-VII double; 10-I single; 10-II, VI double; 10-II[ with 5-6 branches; 10-IV with

6 branches; 10-V, VII with 4 branches; 11-I, ITI-VI single; 11-II double; 11-VII

double or triple; 14-III-VI single; 14-VII, VIII single or double. Paddle, Ovoid;

small serrations on distal portion of basal 0.66 of outer margin, minute proxi-

mally and larger distally; tiny spicules on apical 0.34 of outer and apical 0.28

of inner margins; midrib reaches apex; seta 1-P single; index 1.63.

LARVA (Fig. 52). Chaetotaxy as figured and recorded in Table 18. Descrip-

tion based on one skin with associated adult from Sabah and 5 badly damaged

skins from Singapore (description by Edwards and Given, 1928, was based on the

latter 5 specimens). Head. Seta 4-C with 4-8 branches; 5-C double to 5

branched; 6-C double or triple; 7-C with 7-14 branches; 12-C with 4-8 branches;

dorsomentum with 36-37 teeth. Antenna. Moderately long, moderately pig-

mented, with several scattered small spicules, more numerous on basal area;

seta 1-A with 4-6 branches, moderately long, branches somewhat thickened;

2-A long, with a subapical constriction; 3-A short, 0.19-0.28 length of 2-A;

4-A moderately long, 0.38-0.52 length of 2-A; 6-A short, 0.24-0.33 length of

2-A. Abdomen. Seta 1-VIII with 6-9 branches; 3-VIII with 7-11 branches;

5- VIII with 8-13 branches; 2-X with 6-8 branches; ventral brush with 10 setae

on grid and 1-2 precratal shorter ones, all multiple branched; saddle moder-

ately pigmented, incompletely rings segment X, acus absent; 4 moderately

long to long anal papillae; comb with 8-12 (usually 10) scales arranged ina

single curved irregular row, each scale with a stout median apical spine and

small denticles on laterobasal areas. Siphon. Moderately pigmented; acus

well developed; index 1. 83-2.30, average 2.14; pecten on basal 0.55-0. 57 of

siphon, composed of 11-13 (usually 12) teeth, apical 2-4 teeth wider spaced

than remainder of teeth, apical 6-9 teeth long and slender, each tooth with 1-2

small stout ventral denticles near base; seta 1-S short, 5-8 branched, base

attached on basal 0. 64-0. 65 of siphon distad of last pecten tooth.

DISCUSSION. The above is the first description and illustration of the pupal

stage of cyvtolabis. The description and illustration given for this species by

Delfinado (1967) are of a Finlaya, probably a species in the niveus Group.

Pupal skin NY57-102, described and illustrated by Delfinado (1967: 17, Fig. 7)

for cyrtolabis, does not belong to the adult male even though the adult and slide

labels bear the same collection number and information. The following data

were used in determining the above error: adult specimen of cyrtolabis from

Thailand is a male but the pupal skin is from a female; data sheet and rearing

record for collection NY57 indicates 4 species were associated in this collec-

tion--Culex (Lophoceraomyia ) species (one male with pupal and larval skins),

Aedes (Finlaya) species (probably prominens; 2 males with pupal skins, one

male with larval skin), Aedes (Finlaya) niveus Group (one female with pupal

skin) and Aedes (Aedes) probably new species (one male with pupal skin)

[adult is cyrtolabis |; pupal skin NY57-102 fits well with members of the ”iveus

Group of Finlaya, however, it is atypical of other species of Verrallina,

especially in the development of setae 3-C, 9-VII and VIII and the respiratory

trumpet; pupal skin NY57-102 differs significantly from the associated pupal

skin of the female of cyrtolabis from Sabah which also has an associated

larval skin which compares very well with the larval skins from Singapore;

and the immature habitat of this collection (NY57) was water in a treehole

which is characteristic of Finlaya but quite different from the ground pool

Reinert: New Interpretation of Verrallina 41

inhabiting Verrallina. From the above information I believe the pupal skin

(NY57-102) belongs to a species in the Aedes (Finlaya) niveus Group and was

mistakenly labeled with the same number as the adult male of cyrtolabis.

The larva is redescribed from a skin (with an associated pupal skin and

adult) collected in Malaysia and 5 larval skins from Singapore. The descrip-

tion and illustration of cyrtolabis by Edward and Given (1928: 344-345) were

sketchy and incomplete. The larva was therefore illustrated and completely

described here.

The adult habitus and male and female genitalia of cyvtolabis indicate the

species belongs to the uncus Group of species. Tarsomeres 5 and posttarsi of

the male are illustrated (Fig. 58) for the first time.

DISTRIBUTION. 29 specimens examined: 119, 11°, 51, 1 with associated

pupal and larval skins.

CAMBODIA. Kompong Sela, Phnom Praung; 62, 7°. Kompong Som, Tuk

Sap; 39: 2%

MALAYSIA. Sabah, Beaufort, Sipitang; 12 pl, 12, 1°.

SINGAPORE. 951.

THAILAND. Nakhon Nayok, Amphoe Muang, Khao Yai; 1“.

BIOLOGY. The larva from Malaysia was collected from a small, heavily

shaded ditch containing fresh, clear, temporary, unmoving water located ina

secondary rain forest and at an elevation of 15 m.

AEDES (VERRALLINA) DUX DYAR AND SHANNON

(Figs. 23, 51, 58, 60)

EGG (Fig. 51). Shape. Fusiform; both anterior and posterior ends with

a gradual taper, the latter more acute; greatest diameter near middle. Size.

Length 637-730 microns; width at widest point 154-179 microns. Color.

Dark brown. Chovrion. Cleared chorion viewed with transmitted light shows a

reticulation composed of polygonal and irregular sided cells with cell walls

raised; surface sculpturing well defined on ends and faint on middle 0. 5.

DISCUSSION. Mattingly (1956: 795) reported the Aedes sp. ? No. 50 of

Borel (1930: 287) appeared to be Aedes dux. After examining a number of

specimens of dux from Saigon and comparing them with the description and

illustrations of Borel's Aedes sp. ? No. 50, Iam confident Mattingly's

appraisal of this species was correct.

The following structures of dux are illustrated for the first time: male

posttarsi (Fig. 58), female posttarsi (Fig. 60), tergum IX of the female geni-

talia (Fig. 23). 3

The eggs of dux, previously undescribed, resemble those of the other

described species of Vervallina. This description is based on 9 mature eggs

extracted from the abdomen of a specimen in the USNM which possesses the

following collection data on the labels: PHILIPPINE ISLANDS, Luzon Island,

Rizal, Camp Nichols, 3 July 1923, Pvt. Carraway collector, genitalia prepara-

tion number T73.231. Nomencalture used in the description follows Kalpage

and Brust (1958) and the chorion preparation method used is that of Craig

(1955).

Additional information on dux is included in the discussion section of butleri,

DISTRIBUTION. 322 specimens examined: 2289, 41°, 3 L, 171, 9 E,

15 with associated skins (6 p, 9 1).

INDONESIA. Java, Angke, Kopetakan, Semarang, Tijilintjing; 159, 3c.

MALAYSIA. Perlis, Kg. Sangei Padang, Kg. Sungai Bhorn; 32 pl, 32 p,

42 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

29,10 pl.

PHILIPPINES. Leyte Island, Dulag, Palo; 2°, 1°. Luzon Island, Manila,

Manila; 322, 1%. Pampanga, Camp Stotsenberg; 502. Pangasinan, San Fabian;

52, 8c. Rizal, Camp Nichols; 37%, 2¢, 9 E. Zambales, Subic Bay; 19 pl, 99,

50°, 2L, 171. Mindanao Island, Zamboanga, Zamboanga City; 22, 2°. Min-

doro Island, Caminawit Point; 12, 1°. Samar Island, San Antonio; 19.

Philippine Islands, no locality given; 10°.

SOUTH VIETNAM. Binh Dinh, Qui Nhon; 42. Gia Dinh, Khanh Hoi (Sai-

gon Port), Saigon, Tan Son Nhut (U. S. Army Tent Camps Alpha and Bravo);

22 pl, 329, 20 pl, 100, 1 L. Khonh Hoa, Cam Ranh, Nha Trang; 2°, 1¢

Phong Dinh, Can Tho; 1°. Thua Thien, Phu Bai; 12.

THAILAND. Chanthaburi, Wat Khao Phloi Waen; 12. Phra Nakhon, Bangkok;

82, 30°. Samut Prakan; 22 p, 1¢ p. Songkhla; 12. Thailand, no locality given;

1%.

BIOLOGY. In Malaysia immatures were collected from small animal

hoofprints near the coast, buffalo hoofprints and ground pools containing

colored or clear, fresh, unmoving, temporary water in unshaded areas

located in and near villages.

Immatures were collected from ground pools (Thailand), footprint in

ricefield, buffalo wallow, duckweed pool (Philippines), marshy depressions,

wheel track, footprints and ground pools (South Vietnam).

Adults were taken in light traps in South Vietnam.

AEDES (VERRALLINA) EMBIENSIS HUANG

(Fig. 58)

DISCUSSION. Upon examining the holotype male of embiensis I found

posttarsi I and II with ungues unequal in size, each with a tooth and III with

ungues equal in size, both simple (Fig. 58), which differs from Huang (1968:

25) who reported only the larger ungues of I and Il as bearing teeth. The

basal mesal lobe of the genitalia possessed 7 (9 on other side) moderately

long setae on apical area and 11 (10 on other side) short setae on basal portion

which also differs from Huang who recorded 9 setae on the mesal lobe (apical

area of basal mesal lobe). These features are like carmenti and when the

genitalia of embiensis were compared to the former species no consistant

differences could be found. The adults of these 2 species are also very similar

in habitus, however, embiensis differs by possessing poorly developed white

scaled dorsal bands on abdominal terga III-V and a few more hairs ventrad of

scale patch of mesepimeron.

Several features indicate embiensis and carmenti may be conspecific,

some of which are: no apparent differences between the male genitalia; male

posttarsi are identical; adults are very similar in habitus, except abdominal

banding; tergal abdominal scaling variable in other species of the subgenus;

and type-locality of embiensis is within the geographical range of carmentzt,

The exact taxonomic status of embiensis, however, I feel should await the

collection of progeny reared specimens, both male and female, with associ-

ated immature skins from the type-locality.

Reinert: New Interpretation of Vervallina 43

AEDES (VERRALLINA) FRAGILIS (LEICESTER)

(Figs. 21, 29)

The following information supplements the descriptions of Leicester (1908:

198), Delfinado (1968: 15) and Klein (1973: 5).

FEMALE. Based on the holotype of malayi and females from Cambodia

and Malaysia. Head, Antenna 1.22-1.26 length of proboscis; maxillary palpus

0.17-0.18 length of proboscis; proboscis 0.96-0.98 length of femur I; eyes

separated in front with broad white scales on interocular space. Thorax. Fol-

lowing setae present: 2 median anterior promontory; numerous acrostichal

(anterior and posterior); numerous dorsocentral (anterior and posterior);

scutal fossal (3 anterior and 3-4 lateral); numerous supra-alar; 4 posterior

medial scutal; 1 postalar callar; scutellar (3 long and 2-3 short ones on lateral

lobe, 4 long and 3 short ones on median lobe); 7-10 antepronotal; 3-4 post-

pronotal; 7-8 propleural; 2-4 postspiracular; 3-4 upper and 7-8 posterior

mesepisternal; 5-6 on prealar knob; and mesepimeron with 5-6 dorsad of

scale patch. Prosternum, subspiracular area, paratergite, mesomeron and

metameron bare. Wing. Remigium with 1 seta. Halter. Pedicel pale;

capitellum brown scaled. Genitalia (Fig. 21). As illustrated.

MALE. Based on the lectotype of fragilis. Head. Antenna plumose, 1.09

length of proboscis; maxillary palpus short, 0.09 length of proboscis; eyes

separated infront. Thorax. Following setae present: 2 median anterior

promontory; numerous acrostichal (anterior and posterior); numerous dorso-

central (anterior and posterior); scutal fossal (2-3 anterior and 3 lateral);

numerous supra-alar; 4 posterior medial scutal; 1 postalar callar; scutellar (4

long ones on lateral and median lobes, based on size of setal bases); 7-8

antepronotal; 2 postpronotal; 5-6 propleural; 2 postspiracular; and mesepimeron

with 5 dorsad of scale patch. Prosternum, subspiracular area, paratergite,

mesomeron and metameron bare. Wing. Remigium with 1 seta. Genitalia

(Fig. 29). Phallosome complex, opisthophallus consists of a moderately pig-

mented tergal transverse bridge between parameral apodemes, median

caudal margin produced into a long broad tapering lobe with apex sharply

pointed and extends caudad to near apices of prosophallic arms, a small

pointed tongue-like lobe situated on tergal surface near middle and projected

caudally; sternum IX with 9-10 setae and 5-6 broad scales near center;

remainder of genitalia as in the illustration.

DISCUSSION. The holotype male genitalia of fragilis are completely

illustrated showing the complex arrangement of the phallosome. Edwards

(1917: 222) partially illustrated the male genitalia of this species for the first

time but did not show the basal mesal lobe and only drew part of the phallosome.

Delfinado (1968: Fig. 8) redrew Edward's illustration.

The inclusion of malayias a synonym of fragilis by Delfinado (1968: 15)

appears to be correct but final determination should await reared specimens of

both sexes with associated immature skins or sibling reared males and

females.

Male adult habitus features of fragilis are similar to those of consonensis

(see Reinert 1973b) but these 2 species can be separated as follows--fragilis

possesses: antenna 1.09 length of proboscis; maxillary palpus 0.09 length

of proboscis; postpronotum with 2 posterior setae; and mesepimeron with 5

setae dorsad of scale patch, while consonensis possesses: antenna 0. 82

length of proboscis; maxillary palpus 0.195 length of proboscis; postpronotum with

5 posterior setae; and mesepimeron with 11 setae dorsad of scale patch.

Ag Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

TYPE-DATA. The holotype female of malayi in the BMNH possesses the

following label information in addition to that listed by Delfinado (1968: 16):

20/12/03, Mosq. 53, Dr. G. F. Leicester, 1912-350. I remounted the geni-

talia in Canada balsam and gave the slide preparation number T73. 596.

The holotype of malayi is in fair condition and is as follows: scutum partially

rubbed; right legs I and II and tarsomeres 2-5 of II missing; and right wing

missing. The genitalia are in good condition and mounted on a microscope

slide.

The male lectotype of fvagilis is in poor condition and is as follows: left

antenna and left palpus missing; pleural areas rubbed; all legs except right

femur and tibia of II] missing (parts of 2 legs mounted on slide with genitalia);

left wing missing; and abdominal terga II-V not attached to specimen but glued

to the paper stage attached to the pin. Male genitalia were dissected, mounted

in Canada balsam on a microscope slide and are in good condition.

DISTRIBUTION. 4 specimens examined: 392, 1°.

CAMBODIA. Kompong Som, Tuk Sap; 19.

MALAYSIA. Johore, Panti Forest Reserve; 1°. Selangor, Kuala Lumpur;

12 (holotype of malayi), 1% (lectotype of fragilis).

BIOLOGY. One female from Johore, Malaysia was collected from a biting

collection on a clear day in a partially shaded area of a secondary rain forest

in hilly terrain, on 5 September 1969 at 1300 hours and at an elevation of 76 m.

AEDES (VERRALLINA) GIBBOSUS DELFINADO

(Figs. 25, 53, 58)

PUPA. The pupa is completely described here since the original descrip-

tion was incomplete and based on a single specimen. Chaetotaxy as recorded

in Table 6. Respiratory trumpet. Moderately pigmented; index 3.18-5. 00,

average 4.26. Metanotal plate. Lateral portion heavily pigmented. Abdomen.

Setae 6, 9-VII weakly developed; 4-VIII double or triple; 9-VIII single. Paddle.

Ovoid; minute serrations on distal portion of basal 0.63 of outer margin;

minute spicules on apical 0.37 of outer and apical 0.27 of inner margins; mid-

rib does not reach apex; seta 1-P moderately long, single; index 1.3-1.7,

average 1.51.

LARVA (Fig. 53). Chaetotaxy as figured and recorded in Table 19. Head,

Seta 4-C with 5-10 branches; 5-C triple to 6 branched; 6-C triple to 5 branched;

7-C with 9-15 branches; 12-C with 6-8 branches; dorsomentum with 35-40

teeth. Antenna. Moderately long, lightly to moderately pigmented, with

several small scattered spicules; seta 1-A triple to 5 branched, moderately

long; 2-A long, with a subapical constriction; 3-A short, 0.26-0.36 length of

2-A; 4-A moderately long, 0. 51-0. 70 length of 2-A; 6-A short, 0.29-0.38

length of 2-A. Abdomen. Seta 1-VII with 5-10 branches; 3, 5-VIII with 6-11

branches; 2-X with 6-10 branches; ventral brush with 10 setae on grid and 2

precratal shorter ones, all multiple branched; saddle moderately pigmented,

incompletely rings segment X, acus absent; 4 moderately long anal papillae;

comb with 7-12 (usually 10) scales arranged in a single curved row, each

scale with a stout median apical spine and small denticles on laterobasal areas.

Siphon. Moderately pigmented; acus well developed; index 2.36-3.16, aver-

age 2.88; pecten on basal 0. 49-0. 53 of siphon, composed of 9-12 (usually 10)

teeth, apical 2-3 teeth longer, smooth and wider spaced than remainder each

of which has a small stout ventral denticle near base; seta 1-S short, double or

4 branched, base attached on basal 0. 63-0. 66 of siphon distad of last pecten

Reinert: New Interpretation of Verrallina 45

tooth.

DISCUSSION. The holotype data are amended as follows (information from

collection data sheet and specimen label): holotype male (genitalia mounted on

slide) with associated pupal skin, THAILAND, Songkhia, Amphoe Haad Yai,

near Ton Nga Chang Waterfalls, 6 km from weather station to the main road,

27 May 1965, collection number SL-87-138, USNM type No. 69211, Kol Mong-

kolpanya collector, pupal slide preparation number 18148, collected as a pupa

from a large (8 feet x 30 feet) pool in a dry stream bed, water containing

numerous leaves and sticks with moderate shade from bamboo. Holotype

desposited in the USNM.

The present description and illustration are the first of the fourth stage

iarva of this species and are based on 4 reared specimens with associated skins.

Tarsomeres 5 and posttarsi of the male are also illustrated for the first time

(Fig. 58).

Female genitalia of gibbosus are very distinctive, especially in the develop-

ment of the lower vaginal sclerite and the shape and presence of several setae

on tergum IX (Fig. 25).

Adults of gibbosus are easily recognized by the presence of fine hairs and

scales on the metameron. Other species with this feature and occuring within

the distribution range of gibbosus are members of the andamanensis Group,

however, these species also possess scales on the antepronotum and post-

spiracular area and setae on the median dorsal area of the prosternum.

DISTRIBUTION. 77 specimens examined: 229, me 4 1, 24 with associated

skins (10 p, 131, 11 incomplete).

INDONESIA. Java, Pagar Dewa; 1 genitalia Gidanta on slide) paratype.

MALAYSIA. Negeri Sembilan, Ulu Serting Forest Reserve; 12 pl. Pevak,

Choir Big Game Forest Reserve; 12 pl, 42 p, 39, 5c p. Sabah, Beaufort,

Kudat, Tenom, Mandalom Forest Reserve, Sipitang Forest Reserve, Keningau,

Tuaran; 42 pl, 59, 40° pl, 20°, 4 L. Selangor, Rantaw Panjang; 12 genitalia

(mounted on slide) paratype.

THAILAND. Krabi, Ban Phru Toei; 1% pl. Phangnga, Khao Sung; 12 pl,

1° 1. Songkhla, Amphoe Haad Yai, near Ton Nga Chang Waterfalls; 192 pl, 1° p

(holotype).

BIOLOGY. In Thailand immatures were collected from large stream pools

in dry stream beds (filled with rain water) and a swampy area; from clear or

turbid, fresh, unmoving, temporary water in moderately shaded areas located

in a secondary rain forest, a bamboo grove and a secondary scrub area; and

once at an elevation of 80 m.

Immatures in Malaysia were collected from small and large flood and ground

pools and a large swampy area; from clear, fresh, unmoving, temporary water

in moderately and heavily shaded areas located in secondary rain forests; and

at elevations from 30-260 m. One female was taken in a biting collection.

AEDES (VERRALLINA) HAMISTYLUS LAFFOON

(Figs. 21, 58, 60)

DISCUSSION. The female posttarsi and tarsomeres 5 (Fig. 60) and tergum

IX (Fig. 21) are illustrated for the first time. Male posttarsi and tarsomeres 5

are shown in Figure 58.

DISTRIBUTION. ‘11 specimens examined: 499, 17°, 5 L.

MALAYSIA. Sabah, Lahan Datu, Kg. Bukit Blachan; 19.

PHILIPPINES. Balabac Island, Cape Melville; 46¢, 12° and 5 L (para-

} &

46 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

types), 1° (holotype). Mindoro, San Jose; 2¢ and 4c (paratypes).

BIOLOGY. One female from Malaysia was collected from a biting col-

lection on a clear day, in a partially shaded area of a secondary rain forest,

on 20 May 1970 at 1440 hours and at an elevation of 30 m.

AEDES (VERRALLINA) HARRISONICUS NEW SPECIES

(Fig. 8)

FEMALE. Head, Antenna dark brown, 0.97-0.98 length of proboscis,

pedicel brown with a few small dark scales and short fine brown hairs mesally,

flagellomere 1 with basal 0.55 pale and with a few small dark scales; clypeus

dark brown, bare; maxillary palpus dark brown scaled, 4 segmented, 0.15-

0.17 length of proboscis; proboscis dark brown scaled, 1.02-1.10 length of

femur I; eyes contiguous; ocular setae dark brown; scales on head broad,

dark brown and decumbent except for narrow curved golden scales along

coronal suture and a few on ocular line which become more numerous mesally,

a stripe of broad white scales on lateral surface extending from area in front

of antepronotum to ocular line and postgena with similar scales; occiput with

a number of short brown erect forked scales and a few decumbent broad white

scales. Thorax. Scutal integument dark reddish-brown; scutum covered with

narrow curved reddish-brown scales except for a few narrow curved golden

scales on median anterior promontory area, anterior scutal fossal area and

‘supra-alar area; scutellum with narrow curved reddish-brown scales on each

lobe; dark brown setae on the following areas: 3-4 median anterior promon-

tory, numerous acrostichals (anterior and posterior), numerous dorsocentrals

(anterior and posterior), scutal fossal (3-5 anterior, 5-7 lateral and 2 posteri-

or), numerous supra-alar, 5-6 posterior medial scutal, 1 postalar callar and

scutellar (5-6 long and 3-4 short ones on lateral lobe, 5-7 long and 4-6 short

ones on median lobe); pleural integument dark reddish-brown; antepronotum

without scales, 9-13 dark setae; postpronotum with narrow curved reddish-

brown scales dorsally, 5-6 posterior brown setae; propleuron with a patch of

broad white scales, 15-18 brown setae; prosternum, subspiracular area,

paratergite, mesomeron and metameron bare; postspiracular area without

scales, 5-7 brown setae; mesepisternum with an upper and a lower patch of

broad white (with a brownish tinge) scales, 4-5 upper and 10-16 posterior

setae, lower ones shorter, a few short fine golden hairs on anterodorsal area;

prealar knob without scales, 9-15 brown setae; mesepimeron with a patch of

broad white scales near center, 19-33 golden-brown setae dorsad of scale patch,

numerous long golden hairs caudad of scale patch and extending below it to near

lower margin of sclerite. Legs, Coxae I-III each with several brown setae, I

with broad brown scales and a small dorsal patch of white ones on anterior

surface, II and III each with a patch of broad white scales on anterior surface;

trochanters I-III each with a few broad white scales; femora I and II each with

anterior surface brown scaled, posterior surface brown scaled with a longi-

tundinal white scaled stripe from base to near apex, stripe dorsal on I and

ventral on II, III with anterior and posterior surfaces mostly white scaled and

with a dorsal brown scaled longitudinal stripe from base to apex, forming a

narrow apical brown band; tibiae I-III and tarsi I-III brown scaled; posttarsi

I-III each with 2 ungues, I and II with ungues equal, each with a tooth, II with

ungues equal, both simple. Wing. Dorsal and ventral veins with brown

scales; alula with several moderately broad brown scales on margin; upper

calypter with several brown setae on margin; 1-2 brown remigial setae.

Reinert: New Interpretation of Vervallina 47

Halter. Pedicel pale; capitellum creamy-white scaled. Abdomen. Terga

brown scaled with a laterobasal white scaled patch on I-VII; sterna creamy-

white scaled with a few pale brown scales apically; terga and sterna with numer-

ous golden setae, mostly along posterior margins. Genitalia (Fig. 8). Tergum

VIII lightly to moderately pigmented, very wide, base very slightly concave,

apex gently convex, a number of short and moderately long setae on apical

0.28-0.43, several along apical margin long, numerous broad scales covering

apical 0.63-0.68, basolateral seta present, basal 0. 35-0.40 retracted into

segment VII, VIII-Te index 0.35-0.42, VIII-Te/IX-Te index 1.67-2.27; ster-

num VIII lightly to moderately pigmented, wide, base nearly straight, apex

with a small median indentation with a small lobe on each side of midline,

numerous moderately long setae apically, numerous short and a few moderately

long setae scattered over apical 0.92-0.94, numerous broad scales covering

most of area, apical intersegmental fold unpigmented, VIII-S index 0. 59-0. 63;

tergum IX lightly pigmented with a triangular moderately pigmented plate on

each side of midline, wide, setae absent, covered with minute spicules, IX-Te

index 0.43-0.49; insula unpigmented, short, covered with minute spicules,

4-5 tiny tuberculi, each with a small spicule; lower vaginal lip heavily pig-

mented, median area narrow, lateral areas wide and wrinkled, basal area

bulbous and wrinkled, lower vaginal sclerite composed of a large median

heavily pigmented wrinkled plate which is incompletely divided along most of

midline except basal portion; upper vaginal lip heavily pigmented, wide through-

out, forming nearly right angles at posterolateral margin, developed into a

broad shield caudad of spermathecal eminence, upper vaginal sclerite heavily

pigmented, large, extending mesad from lateral portions of vaginal lip and

covering much of the vaginal roof; spermathecal eminence heavily pigmented,

large, dorsal surface with a long basal extension covered with closely matted,

moderately long, branched spiny excrescences, ventral outline heart-shaped;

postgenital lobe short, moderately broad, apex with a small median indenta-

tion, 21-29 setae on each side of midline, completely covered with minute

spicules, dorsal PGL index 0.47-0.68, ventral PGL index 0. 70-0. 79; peri-

anal membrane with minute spicules; cercus triangular in shape, moderately

long, broad at base, apex acute, completely covered with minute spicules,

dorsal surface with numerous short and moderately long setae over apical

0.90-0.94, a few long setae at apex, 0-3 broad scales, ventral surface with

short and moderately long setae on outer margin and becoming more numer-

ous apically, cercus index 2.00-2.42, cercus/dorsal PGL index 3.25-4.48; 3

seminal capsules, 1 large, 1 medium and 1 small in size, heavily pigmented,

ovoid, each with several small seminal capsule pores near orifice, base of

accessory gland duct unpigmented.

MALE, PUPA, LARVA AND EGG. Not known.

DISCUSSION. There appears to be no consistant differences between the

adult habitus of harrisonicus and the other members of the wncus Group.

Female genitalia of harrisonicus are similar to the other members of the

uncus Group but can easily be separated from them by the development of

the lower and upper vaginal lips, shape of the area covered with spiny

excrescences of the spermathecal eminence, and development of the lower

vaginal sclerite.

There is a possibility that havvrisonicus may be the female of one of the

previously described members of the uwncus Group for which only the male is

known, however, it is impossible at present to assign it to an existing species

because the adults of the group exhibit no noticeable differences and the im-

matures of the new species are unknown. Since harrisonicus is separated

48 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

geographically from the species with unknown females (nubicolus, protuberans

and torosus) it is best at present to describe it as new.

This species is dedicated to Bruce A. Harrison, Department of Entomology,

Walter Reed Army Institute of Research, Washington, D. C., in recognition of

his work on mosquitoes in Southeast Asia.

TYPE-DATA. Holotype female, INDONESIA, Java, Megamendg Mts.,

10 January 1909, Bryant and Palmer collectors, genitalia preparation number

T73. 388; 7 female paratypes, genitalia preparation numbers T73.400, T73.401,

T73.402, T73.568, T73.569, 46-II-14a (no adult), 46-III-10e, each with the

following label data, INDONESIA, Java, Mt. Gede, 1909, Bryant and Palmer

collectors. The holotype and 6 paratypes are deposited in the USNM, and one

paratype will be deposited in the BMNH.

The holotype is in good condition, though somewhat rubbed, and all para-

types are moderately to heavily rubbed, most also with several legs missing.

DISTRIBUTION. Known only from localities mentioned in type-data.

BIOLOGY. Nothing is known about the biology of this species.

AEDES (VERRALLINA) INCERTUS EDWARDS

(Figs. 9, 25)

Aioretomyia Taeniata Leicester 1908, Culicidae Malaya 3: 190 (2) (Name pre-

occupied).

Aedes(Aedes) incertus Edwards 1922a, Indian J. Med. Res. 10: 268 (nom.

nov. for Taeniata Leicester 1908, non Wiedemann 1828).

Aedes (Aedes) dermajoensis Brug 1831, Tijdschr. Entomol. 74: 250 (o*).

NEW SYNONYM.

FEMALE. Head, Antenna dark brown, 1.02-1.10 length of proboscis,

pedicel light brown with a few small brown scales and short fine brown hairs

mesally, flagellomere 1 with basal 0.4 pale and with several small brown

scales; clypeus dark brown, bare; maxillary palpus blackish-brown scaled,

0.14-0.15 length of proboscis; proboscis blackish-brown scaled, 1.06-1.12

length of femur I; eyes contiguous; ocular setae dark brown and well developed;

scales on head broad, dark brown and decumbent except for narrow curved

golden-brown scales along coronal suture and a few on ocular line which be-

come more numerous mesally, a stripe of broad white scales on lateral sur-

face extending from area in front of antepronotum to ocular line and postgena

with similar scales; occiput with a number of short brown erect forked scales

and a few decumbent broad white scales. Thorax, Scutal integument reddish-

brown; scutum covered with narrow curved reddish-black scales except for

narrow curved golden-white scales forming small patches (occasionally only a

few golden-white scales) on median anterior promontory area, anterior scutal

fossal area, supra-alar area and a few on basal area of median lobe of scutel-

lum; scutellum with pale brown to golden narrow curved scales on each lobe;

dark brown setae on following areas: 3 median anterior promontory, numerous

acrostichals (anterior and posterior), numerous dorsocentrals (anterior and

posterior), scutal fossal (3-6 anterior, 2-3 lateral and 1-2 posterior), numer-

ous supra-alar, 6 posterior medial scutal, 1 postalar callar and scutellar

(3-4 long and 2-4 short ones on lateral lobe, 5-6 long and 3-5 short ones on

median lobe); pleural integument reddish-brown; antepronotum without scales,

8-13 brown setae; postpronotum with narrow curved reddish-black scales on

upper area, 3-5 posterior brown setae; propleuron with a patch of broad white

Reinert: New Interpretation of Vervallina 49

scales, 12-18 brown setae; prosternum, subspiracular area, paratergite,

mesomeron and metameron bare; postspiracular area without scales, 2-4 brown

setae; mesepisternum with an upper and a lower patch of broad white scales,

3-6 upper and 10-16 posterior brown setae, lower ones shorter and golden, a

few short fine golden hairs on anterodorsal area; prealar knob without scales,

6-9 brown setae; mesepimeron with a patch of broad white scales near center,

10-17 golden setae dorsad of scale patch, several short fine golden hairs along

posterior margin and posteroventral corner of scale patch. Legs, Coxae I-III

each with several brown setae, I with broad brown scales and a small dorsal

and ventral patch of broad white scales on anterior surface, II and III each with

a patch of broad white scales on anterior surface; trochanters I-III each with a

few white and pale brown scales; femora I and II each with anterior surface

brown scaled, posterior surface brown scaled with a longitudinal white scaled

stripe from base to near apex, stripe dorsal on I and ventral on II, II with

anterior and posterior surfaces mostly white scaled and with a dorsal brown

scaled longitudinal stripe from base to apex, forming a narrow apical brown

scaled band, II and III each with a few anterior apical pale scales; tibiae I-III

brown scaled, I and II each with a posterior longitudinal white scaled stripe,

indistinct on I; tarsi I-iII each brown scaled; posttarsi I-III each with 2 ungues,

I and II with ungues equal, each with a tooth, III with ungues equal, both simple.

Wing. Dorsal and ventral veins with brown scales; alula with several narrow

brown scales on margin; upper calypter with several brown setae on margin;

1-2 brown remigial setae. Halter. Pedicel pale; capitellum with creamy-white

lateral scales and a few brown dorsal ones. Abdomen. Terga brown scaled

with a laterobasal white scaled patch on I-VII, dorsal pale scaled markings

extremely variable, ranging from well developed white scaled basal bands

(not reaching laterobasal white scaled patches) on II-VI to dorsal areas all

brown scaled; sterna each brown scaled with a large triangular laterobasal

white scaled patch which connects mesally on base of some sterna; terga and

sterna with numerous golden setae, mostly along posterior margins. Genitalia

(Figs. 9 and 25). The female genitalia of the lectotype of incertus are illustra-

ted. The genitalia of this species are very uniform but vary from specimen to

specimen in the number of short hair-like spicules on the basal area of the

spermathecal eminence. The number of spicules range from a few to many,

the lectotype having an intermediate number.

MALE. Similar to female in general habitus but with the following differ-

ences. Head, Antenna plumose, 0.87-0.95 length of proboscis; maxillary pal-

pus 0.10-0.11 length of proboscis; proboscis 1.27-1.31 length of femur I.

Thorax, Scutal fossal setae (2-4 anterior, 3-4 lateral and 1 posterior); 10-13

propleural setae; 3-4 upper and 8-12 posterior mesepisternal setae; 5-8 prealar

knob setae; mesepimeron with 9-12 setae dorsad of scale patch. Legs. Post-

tarsi each with 2 ungues, I and II each with ungues unequal, each with a tooth,

II with ungues equal, both simple. Abdomen. Terga I-VII each with a small

laterobasal white scaled patch, dorsal white scaled markings variable, ranging

from all dark dorsally to a well developed white scaled basal band which is con-

tinuous with laterobasal white scaled patch. Genitalia. As illustrated by Del-

finado (1967, Fig. 8) and Klein (1973, Fig. 1). The tergoapical projection of

the gonocoxite varies from sharply pointed to bluntly pointed (Fig. 25).

PUPA. The pupa is completely described here since the original description

was incomplete. Chaetotaxy as recorded in Table 7. Respivatory trumpet.

Heavily pigmented; index 3.91-5.29, average 4.62. Abdomen. Setae 6, 9-VII

weakly developed; 4- VIII double to 5 branched; 9-VIII single. Paddle. Ovoid;

minute serrations on distal portion of basal 0.67-0.72 of outer margin; minute

50 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

spicules on apical 0. 40-0. 49 of outer and apical 0.15-0.23 of inner margins;

midrib does not reach apex; seta 1-P moderately long, single; index 1. 50-1. 72,

average 1.6.

LARVA. Chaetotaxy as recorded in Table 20. Head. Seta 4-C with 5-8

branches; 5-C double to 4 branched; 6-C double or triple; 7-C with 8-12 branches;

12-C with 6-9 branches; dorsomentum with 34-38 teeth. Antenna. Moderately

long, lightly pigmented, with several small scattered spicules, more numerous

on basal area; 2-A long, with a subapical constriction; 3-A short, 0.2-0.3 length

of 2-A; 4-A moderately long, 0.54-0.70 length of 2-A; 6-A short, 0.28-0. 39

length of 2-A. Abdomen. Seta 1-VUI with 5-9 branches; 3-VII with 5-10 bran-

ches; 5-VIII with 7-12 branches; 2-X with 6-10 branches; ventral brush with

9-10 setae on grid and 2-3 precratal shorter ones, all multiple branched; saddle

moderately pigmented, incompletely rings segment X, acus absent; 4 moderate-

ly long anal papillae; comb with 8-11 (usually 10) scales arranged in a single ir-

regular row, each scale with a stout median apical spine and small denticles on

laterobasal areas. Siphon. Moderately pigmented; index 2.51-3.06, average

2.71; pecten on basal 0. 51-0. 56 of siphon, composed of 8-12 (usually 10) teeth,

apical 2-3 teeth longer and wider spaced than remainder of teeth, each tooth

long, slender and with a small stout ventral denticle near base; seta 1-S short,

triple to 5 branched, base attached on basal 0. 64-0. 65 of siphon distad of last

pecten tooth.

DISCUSSION. The lectotype female of incertus has abdominal terga II-VI

(remainder of abdomen missing) each brown scaled with a narrow tergobasal

white scaled band and a small laterobasal white scaled patch which is not con-

nected to the white scaled band. The holotype male of dermajoensis has ab-

dominal terga I-IV each brown scaled with a small laterobasal white scaled

patch. The male genitalia of the latter have a sharply pointed tergoapical

projection of the gonocoxite (as in Delfinado 1967, Fig. 8). Two males from

Sabah, Malaysia have identical genitalia to the type of dermajoensis but one

has well developed basal white scaled bands on abdominal terga II-III and the

other one has bands on terga II-VI. One male from Krabi, Thailand and 3

males from Cambodia have well developed basal white scaled bands onabdominal

terga III-V and a few tergobasal white scales on II, one Cambodian specimen

also with a band on VI, 2 additional Cambodian specimens have white bands on

terga I[-VII, and the genitalia of all 5 are identical to dermajoensis except

the tergoapical projection of the gonocoxite which is bluntly pointed (as in

Klein 1973, Fig. 1). Two males from Songkhla, Thailand have narrow basal

white scaled bands on abdominal terga III-VI, a few tergobasal white scales

on II and genitalia identical to dermajoensis. One male from the type-locality

of dermajoensis has abdominal markings like dermajoensis but genitalia like

the Cambodian males. A collection from Selangor, Malaysia contained 8 males

all of which had genitalia similar to the holotype of dermajoensis but with basal

white scaled bands on the following terga: III-V on 2 males, III-IV on 3 males,

and no bands on 3 males. Four females from Cambodia, one female from

Phangnga, Thailand and one female from Perak and 3 females from Selangor,

Malaysia have abdominal markings similar to the lectotype of incertus. Three

females from Songkhla, Thailand and one female from Sabah, Malaysia have

abdominal terga each brown scaled with a small laterobasal white scaled patch

and only 2-6 white scales tergobasally on terga III and IV, the Sabah specimen

also with 1-3 white scales on Il, however, 3 females from this same locality,

one female from Narathiwat, Thailand and 4 females from Sabah, Malaysia,

lacked tergobasal white scales. The above specimens demonstrate that the

abdominal tergal markings are extremely variable and range from well

Reinert: New Interpretation of Verrallina ol

developed tergobasal white scaled bands to lacking tergal white scales in both

incertus and dermajoensis. The tergoapical projection of the gonocoxite is

variable in the male genitalia of dermajoensis (Fig. 25). I can not find any

differences between the immatures of incertus females and immatures of der-

majoensis males. The larva and pupa of incertus are redescribed above.

From the above evidence I believe dermajoensis to be conspecific with incertus.

Upon examination I found the pupal specimens of dermajoensis, studied and

labeled by Delfinado (1967), to have seta 6-VII 4-5 branched (not single), 7-V

triple to 7 branched (not single) and cephalothoracic seta 10-C double to 4

branched (not 5-6. branched). These setae conform to the branching of the

setae on incertus and therefore support the above synonymy. In addition, the

females of incertus and males of dermajoensis from Thailand reported by

Delfinado (1967) came from the same collection.

Female genitalia of the above specimens are very uniform but vary in the

number of small hair-like spicules on the basal area of the spermathecal

eminence. The genitalia are similar to those of varus and panayensis (see

discussion of varus).

Aedes incertus belongs to a species group with panayensis, prioekanensis

and varus. The female and male genitalia of this group show many similarities

in their development.

TYPE-DATA. The lectotype female of incertus in the BMNH possesses the

following adult label information: A. Taeniata| written on top of paper stage |,

Jungle 5th mile Gombak Rd., Kuala Lumpur, Midday, 29/2/04 [ written on

underside of paper stage]; co-type; Kuala Lumpur, Fed. Malay States,

Dr. G. F. Leicester, 1912-350; 2 lectotype, M. Delfinado, 1960; SEAMP

Acc. No. 374; genitalia preparation number T73.561. The holotype male of

dermajoensis contains the following adult label information: Aedes (Ae.)

dermajoensis n. sp. o TYPE Brug; 18.153; Sumatra: Benkulen, xi-xii. 1929,

Col. S. L. Brug, B. M. 1930-511; From larva in pool in virgin forest; SEAMP

Acc. No. 376. Brug's field notes for this specimen differ somewhat from the

label data and are as follows: Ae. dermajoensis, 18.153 type Br. Mus.| 18.

153 is the specimen collection number], Marsh in low primary forest (Breed-

ing place H) [ translation kindly supplied by P. F. Mattingly], de Rook | col-

lector], Dermajoe (Benkoelen), Nov.-Dec. 1929. The locality Benkoelen was

spelled differently on the label attached to the holotype.

The lectotype female of incertus is in very poor condition and is as follows:

only a fragment of thorax remains on a minuten pin; abdominal segments I-VI

glued to paper stage; right and left midlegs and right hindleg glued to paper

stage; the genitalia were mounted on a celluloid point but I remounted the genit-

alia in Canada balsam on a microscope slide (genitalia in good condition); re-

mainder of specimen missing.

The holotype male of dermajoensis is in poor condition and is as follows:

most of head missing, including appendages; thorax pinned through pleural

areas of both sides with a minuten pin, pleural areas somewhat rubbed; ab-

dominal segments I-IV and part of V intact and in good condition; right foreleg

intact, left midleg mounted in balsam on a celluloid point with genitalia, re-

maining legs missing; both wings in fair condition; genitalia mounted in a drop

of balsam on a celluloid point and in good condition.

DISTRIBUTION. '73 specimens examined: 269, 21°, 1 pl, 14 with asscoi-

ated skins (4 p, 101).

CAMBODIA. Kompong Sela, Stung Chral; 32, 30°. Kompong Som,

Sihanoukville; 12. Siem Reap, Angkor Forest; 2.

INDONESIA. Sumatra, Dermajoe (Benkoelen); 1° (holotype of dermajoensis),

ee. ic.

o2 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

MALAYSIA. Pevak, Tanjong Tualang Forest Reserve; 12. Sabah, Beau-

fort; 52, 10 pl, 10°. Selangor, Kuala Lumpur; 19 (lectotype of incertus), Tem-

pler Park;.3%,.1¢ pl, 1¢ p,-1¢.

THAILAND. Kyvrahi, Ban Phru Toei; 1% p. Navathiwat, Amphoe Waeng,

Soi Teo; 22 pl. Phangnga, Nam Tai; 12. Songkhla, Amphoe Haad Yai, Ton

Nga Chang Waterfalls; 62 pl, 12, 20% p, 1 pl.

BIOLOGY. In Thailand immatures were collected from small and large

flood pools in dry stream beds and a small seepage pool; from clear or turbid,

fresh, unmoving, temporary water in partially shaded areas; located in secon-

dary rain forests; and at elevations from 46-80 m.

In Malaysia immatures were collected from small flood and ground pools

containing clear, fresh, unmoving, temporary water, in partially and heavily

shaded areas located in secondary rain forests in hilly and valley terrains, and

at elevations of 30-152 m. In Perak a female was taken biting man in a sec-

ondary rain forest between 0800-1050 hours.

AEDES (VERRALLINA) INDEC ORABILIS (LEICESTER)

(Fig. 21)

DISCUSSION. Aedes indecorabilis was described by Leicester (1908: 200)

from 2 cotypes, a male and a female, which he stated (page 201) were "Bred

from larvae taken in a small jungle pool near Kuala Lumpur.'' Only the 2

damaged adults remain, the associated immature skins could not be located

and are presumed lost. The cotype male possesses the following information,

in Leicester's handwriting, on the underside of the paper stage holding the

adult: Larva fr. pool in jungle patch Gombak Rd. 5 miles fr. Kuala Lumpur,

29/2/04; however, the female cotype contains the following handwritten infor-

mation on the underside of the paper stage: Jungle 5th mile Gombak Rd. ,

11/2/04. The above label data indicate that both the cotypes were collected

in the same locality but on different days in February 1904 and no mention is

given of the female being reared. The adults of both cotypes are badly dam -

aged but the habitus and chaetotaxy of both are very similar. I am therefore

considering that the male and female cotypes are conspecific. Delfinado

(1967: 26) selected the male cotype as lectotype and redescribed and illustrated

both the male and female. Klein (1973: 6) also redescribed and illustrated

specimens from Cambodia which he attributed to this species. I have com-

pared Klein's Cambodian specimens with the male and female cotypes of

indecorabilis and can find no apparent differences in the adult habitus and

male genitalia; however, the female genitalia (Fig. 21) of the Cambodian

specimens are markedly different from the cotype female (Fig. 21) of indecor-

abilis and are very similar to those of stungus (Fig. 21). The adults of stun-

gus differ in abdominal marking from these specimens. Klein's female speci-

mens therefore may belong to a new species. The correct association of sexes

of indecovabilis can not be solved with the present specimens and must wait

until sibling reared males and females become available.

DISTRIBUTION. 6 specimens examined: 19, 5c.

CAMBODIA. Kompong Sela, Phnom Praung; 2% (3° examined by me from

this locality, sent by Dr. Klein, do not belong to this species--see discussion

above).

MALAYSIA. Selangor, Kuala Lumpur; 1° (lectotype), 1 (paralectotype).

THAILAND. Nakhon Si Thammarat, Amphoe Chawang, Ban Tha Phon; 2c

genitalia.

Reinert: New Interpretation of Verrallina o3

BIOLOGY. The 2 males from Thailand were taken in a light trap situated

in light forest vegetation on 21 September 1962 between 1800 and 0600 hours

(collection number T-1593).

Leicester (1908: 201) reports the types were reared from larvae taken ina

small jungle pool in Malaysia. In Cambodia Klein (1973: 6) found larvae in

jungle pools.

AEDES (VERRALLINA) INDIC US (THEOBALD)

(Figs. 10, 30, 44, 54, 58, 60, 64)

Aedes indicus is the representative species for Series III, Section A, and

is therefore completely redescribed below.

FEMALE (Fig. 64). Head. Antenna dark brown, 0.99-1.03 length of pro-

boscis, pedicel dark brown with a few small dark scales and short fine dark

hairs mesally, flagellomere 1 with basal 0.65 pale and with several small

dark scales near middle, flagellar whorls with 6 dark setae; clypeus dark

brown, bare; maxillary palpus blackish-brown scaled, 4 segmented, 0.16-

0.20 length of proboscis; proboscis blackish-brown scaled; 1.14-1.26 length

of femur I; eyes nearly contiguous; ocular setae dark brown and well developed;

scales on head broad, dark brown and decumbent except interocular space,

ocular line and occiput each of which has narrow curved white scales, similar

scales on vertex forming a single row on each side of coronal suture from

occiput to near ocular line, occiput with a number of short brown erect forked

scales and a few broad white scales along anterior margin of narrow scaled

area, a narrow stripe of broad white scales on lateral surface extending from

area in front of antepronotum to ocular line, postgena with broad creamy

colored scales. Thorax. Scutal integument dark brown; scutum covered with

narrow curved dark reddish-brown scales except for narrow curved white

scales on median anterior promontory area, anterior scutal fossal area, supra-

alar area and along lateral margins of prescutellar space; prescutellar space

bare; scutellum with a patch of narrow curved white scales on each lobe, a few

similar brown scales basally on median lobe; dark reddish-black setae on the

following areas: 2-4 median anterior promontory, numerous acrostichals

(anterior and posterior), numerous dorsocentrals (anterior and posterior),

scutal fossal (3-6 anterior, 5-7 lateral, 1-3 median and 2-5 posterior), numer-

ous supra-alar, 5-7 posterior medial scutal, 1 postalar callar and scutellar

(8-11 long and 3-5 short ones on lateral lobe, 5-11 long and 3-5 short ones on

median lobe); pleural integument dark brown; antepronota widely separated,

without scales, 14-23 long and short dark setae; postpronotum with narrow

curved brown scales on upper area and several similar white ones on posterior

area cephalad of setae, 5-8 posterior dark setae; propleuron with a patch of

broad white scales, 8-13 setae; prosternum, subspiracular area, mesomeron

and metameron bare; postspiracular area with several narrow curved white

scales, 6-10 dark setae; paratergite with a few narrow curved white scales

laterally (rubbed off in some specimens); mesepisternum with an upper and a

lower patch of broad white scales, 3-4 upper and 9-15 posterior dark setae;

prealar knob with 1-3 long moderately broad scales (2 specimens), 10-14

dark setae; mesepimeron with a patch of broad white scales near center, dor-

sal ones longer and somewhat narrower, 10-18 setae (several hair-like) dor-

sad of scale patch (a few long scales also usually dorsad of setae), 2-4 small

golden hairs along posterior margin of scale patch. Legs; Coxae I-III each

with several dark setae, I with broad brown scales and a small dorsal patch

o4 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

of white ones on anterior surface, II and III each with a patch of broad white

scales on outer area of anterior surface; trochanters I-III each with broad

pale scales; femora I and II each with anterior surface brown scaled, I with

scales paler brown colored, II with a narrow longitudinal anteroventral pale

stripe, III with most of anterobasal area pale scaled, I-III each with a few

white apical anterior and posterior scales, I-III with posterior surfaces mainly

pale scaled, I with a narrow longidudinal posteroventral brown scaled stripe,

II and III each with a posterodorsal longitudinal brown scaled stripe which is

narrow basally and broader apically; tibiae I-III each dark brown scaled, I

and III each with a broad posteroventral longitudinal pale scaled stripe, II with

posterior surface pale scaled; tarsi I-III each dark brown scaled; posttarsi I-

III (Fig. 60) each with 2 ungues, I and II with ungues equal, each ungue with a

tooth, III with ungues equal, both simple. Wing. Dorsal and ventral veins

with dark brown scales; alula with several moderately broad and a few narrow

brown scales on margin; upper calypter with several pale setae on margin;

1-3 remigial setae. Halter. Pedicel pale; capitellum creamy-white scaled

with dorsal ones pale brown. Abdomen. Terga blackish-brown scaled with

white markings as follows: I with a few dorsomedian white scales and latero-

tergite covered with white scales, II-VI each with a narrow laterobasal white

scaled patch, II-VII each with a dorsomedian transverse white scaled band

which extends cephalad and attached to base on lateral margin, bands joined to

lateral white scaled patches on IV-VII, bands incomplete on tergomesal areas

of V-VII, VIII with a number of scattered white scales; sterna brown scaled

with laterobasal pale scaled areas; terga and sterna with numerous golden

setae, mostly along posterior margins. Genitalia (Fig. 10). Tergum VII

moderately pigmented with a lightly pigmented median basal area, wide, base

slightly concave, apex gently convex, a number of short and moderately long

thin setae at apex and similar ones scattered over apical 0. 50-0. 67, basolater-

al seta present, covered with minute spicules, numerous broad scales cover-

ing apical 0.65-0.75, basal 0.4-0.6 retracted into segment VI, VII-Te index

0, 54-0.57, VIII-Te/IX-Te index 5.48-6.00; sternum VIII moderately pigmented,

wide, base concave mesally, apex with a shallow median indentation with a

small lobe on each side of midline, numerous moderately long setae apically,

more numerous and slightly stouter on apical lobes, numerous short and a few

moderately long setae scattered over apical 0. 85-0. 88, covered with minute

spicules, numerous broad scales covering most of apical 0.77-0. 87, apical

intersegmental fold unpigmented, VIII-S index 0. 61-0. 62 tergum IX moderately

pigmented, very wide and short, ribbon-like, setae absent, covered with min-

ute spicules, [X-Te index 0.16-0.17; insula unpigmented, short, covered with

minute spicules, 5-6 small tuberculi, each with a small spicule; lower vaginal

lip heavily pigmented, narrow, without spicules except a few on outer margin

of hinge area, with a small caudally projecting heavily pigmented median apo-

deme, lower vaginal sclerite composed of a pair of sigmoid-shaped heavily

pigmented structures situated lateromesad on an elevated portion of the mem-

brane extending over area between the vaginal lip; upper vaginal lip heavily

pigmented, complex, lateral surfaces forming wide parallel plates which pro-

duce sharp posterior angles and curve sternally and then mesad to form a

continuous band, a strong apodeme extends mesad from near midpoint of dor-

sal surface of lateral area and forms a large continuous upright shield sur-

rounding posterior portion of spermathecal eminence, spicules only along pos-

terior margin of upper vaginal lip, upper vaginal sclerite heavily pigmented,

extremely large and complex, base attached along entire lateral margin of

upper vaginal lip except for a short posterior area, a large caudal arm extends

Reinert: New Interpretation of Vervrallina o9

mesad and forms a crown around the posterior portion of the spermathecal

eminence, posterior area of sclerite not pigmented, cephalic arm very large

forming a continuous broad band around anterior of spermathecal eminence,

wrinkled, fenestrated and with a mesal transverse slit; spermathecal emin-

ence heavily pigmented, large, deep in depth, ovoid in dorsal outline, long

spiny excrescences with lateral projections attached to cephalic area; sper-

mathecal eminence, upper vaginal lip and upper vaginal sclerite cover nearly

entire upper vaginal area with heavily pigmented structures; postgenital lobe

short, narrow, apex with a moderately deep median indentation, 8-15 setae on

each side of midline, completely covered with small spicules, dorsal PGL

index 0.94-1.29, ventral PGL index 1.06-1.44; peri-anal membrane with

minute spicules; cercus triangular in shape, moderately long, broad at base,

apex acute, completely covered with small spicules, dorsal surface with a

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

ventral surface with several short and moderately long setae on outer 0.4, a

number (14 or more) broad scales scattered over entire dorsal surface, cer-

cus index 3.33-3.86, cercus/dorsal PGL index 3. 00-4. 63; 3 seminal cap-

sules, 1 large, 1 medium and 1 small in size, heavily pigmented, elliptical,

each with a moderately broad neck, several small seminal capsule pores near

orifice, base of accessory gland duct unpigmented to lightly pigmented.

MALE. Similar to female in general habitus but with the following differ-

ences. Head. Antenna plumose, 10-11 setae in whorls, 1.03-1.14 length of

proboscis; maxillary palpus 0.14-0.18 length of proboscis; proboscis 1. 06-

1.21 length of femur I; vertex without narrow scales on coronal suture. Thorax,

Scutellar setae (4-7 long and 2-4 short ones on lateral lobe, 4-8 long and 2-4

short ones on median lobe), 7-11 antepronotal setae; 4-5 postpronotal setae;

7-8 propleural setae; 2-3 postspiracular setae; 5-6 prealar knob setae; mese-

pimeron with 5-8 setae dorsad of scale patch and 1-2 hairs posterior to patch.

Legs. Posttarsi I-III (Fig. 58) each with 2 ungues, I and II each with ungues

unequal, larger one with a tooth, III with ungues equal, both simple. Abdo-

men. Terga with laterobasal white patches which usually extend slightly onto

dorsal surfaces. Genitalia (Fig. 30). Tergum IX heavily pigmented, band-

like with lateral areas broad, cephalic margin evenly concave, outer caudal

margin fused to tergum X, setae absent, broadly connected laterally to ster-

num IX; gonocoxite short, broad, heavily pigmented, mesal surface membran-

ous with minute spicules, dorsal surface with apex extended into a long flat-

tened broad lobe with 4-5 short setae in a small patch on basomesal area, a

number of short and long stout setae on apical 0.8, ventral surface with a

number of short and long stout setae on apical 0.6, a number of broad scales

on ventral and lateral surfaces of gonocoxite; gonostylus moderately pigmented,

moderately long (approximately 0.73 length of gonocoxite minus apical lobe),

narrow, curved, base with a small lobe, apex pointed and tip recurved, 1-2

short thin setae subapically, attached subapically on gonocoxite; basal mesal

lobe moderately pigmented, consists of a basal lobe with 18-19 dorsal short

thin setae and 15-17 short stout flattened acuminate setae on mesal surface,

a broad flat extension onto mesal membrane of gonocoxite which bears 8-9

short fine setae and minute spicules, and an apical long curved arm which

resembles the gonostylus, lobe connected mesally with its mate by a narrow

moderately pigmented band covered with minute spicules and located ventrad

of apical portion of prosophallus and phallus; proctiger with paraproct formed

into a heavily pigmented long slender slightly curved acuminate arm with

basal area expanded and connected to tergum IX, base also with a short project-

tion which extends ventrad and articulates at a point on the dorsomesal surface

o6 | Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

of the parameral apodeme near base of opisthophallus, base of proctiger

lightly fused with tergum X, cercus membranous and without cercal setae;

tergum X heavily pigmented, consists of a small rectangular plate laterad of

paraproct base and extended ventrally from caudal margin of tergum IX; phal-

losome complex, opisthophallus--consists of a lightly to moderately pigmented

tergal transverse bridge between parameral apodemes, caudal margin slightly

indented mesally, sternobasal area attached to tergobasal portion of outer area

of prosophallus, located dorsad of phallus and prosophallus and ventrad of proc-

tiger, phallus--consists of a pair of moderately long, heavily pigmented, nar-

rowly separated, caudally produced arms which are fused together by a narrow

basal plate which extends laterally and is fused to tergobasal portion of pro-

sophallus, apex of phallus formed into a tergal penis filament which consists

of a small lightly pigmented and sclerotized lobe produced cephalad, proso-

phallus--consists of a pair of moderately long widely separated acuminate

arms, each arm with distal portion heavily pigmented and slightly curved,

proximal portion lightly pigmented, formed into abroadly rounded area which

is attached to tergomesal portion of paramere, located laterad and slightly

ventrad of phallus, prosophallic arm approximately equal in length to phallic

arm, paramere--heavily pigmented, approximately equal in length to phallic

arm, articulates at a point approximately 0.65 from base with caudal area of

parameral apodeme, parameral apodeme--moderately to heavily pigmented,

moderately long, outer surface broadly attached to tergobasal apodeme of

gonocoxite; sternum IX heavily pigmented throughout, large, broad, latero-

apical areas rounded, 15 short and moderately long setae in an elongate patch

along caudal margin.

PUPA (Fig. 44). Chaetotaxy as figured and recorded in Table 8. Ocular

plate with moderately developed cuticular facets. Respivatory trumpet.

Heavily pigmented; hair-like spicules on distal 0. 57-0. 60 of inner surface;

index 2. 53-3.45, average 2.88. Abdomen. Setae 6, 9-VII weakly developed;

4-VIII single to triple; 9- VIII single or double. Paddle, Ovoid; minute serra-

tions on most of basal 0. 68-0.72 of outer margin; minute spicules on apical

0. 35-0. 38 of outer and apical 0.13-0.15 of inner margins; midrib does not

reach apex; seta 1-P moderately long, single; index 1.43-1.68, average 1.56.

LARVA (Fig. 54). Chaetotaxy as figured and recorded in Table 21. Head.

Seta 4-C double to 5 branched; 5-C triple; 6-C double to 4 branched; 7-C with

7-12 branches; 12-C with 4-9 branches; dorsomentum with 35-40 teeth. An-

tenna. Moderately long, moderately pigmented, with several small scattered

spicules, more numerous on basal area; seta 1-A double or triple, moderately

long, branches thickened; 2-A long, with a subapical constriction; 3-A short,

0. 19-0. 26 length of 2-A; 4-A moderately long, 0.31-0.45 length of 2-A; 6-A

short, 0.20-0.26 length of 2-A. Abdomen. Seta 1-VIII double to 5 branched;

3- VIII with 6-10 branches; 5-VIII with 7-12 branches; 2-X with 9-11 branches,

all branches approximately equal in length; ventral brush with 12-14 setae on

erid and 2-3 precratal shorter ones, all multiple branched; saddle moderately

pigmented, incompletely rings segment X, acus absent; 4 moderately long

anal papillae; comb with 8-12 (usually 10) scales arranged in a single curved

irregular row, each scale with a stout median apical spine and small denticles

on laterobasal areas. Siphon. Moderately pigmented; acus well developed;

index 1. 68-2.37, average 1. 88; pecten on basal 0. 56-0. 65 of siphon, com-

posed of 11-15 (usually 12) teeth, apical 2 teeth longer and wider spaced than

remainder of teeth, each tooth long, slender and with 1-3 small stout ventral

denticles near base; seta 1-S short, triple to 5 branched, base attached on

basal 0.70-0.77 of siphon distad of last pecten tooth.

Reinert: New Interpretation of Vervallina o7

DISCUSSION. Delfinado (1967: 4, 25) reported Thailand as a new country

record for indicus based on 2 females. After examining the 2 Thailand speci-

mens and comparing them with the types of indicus, nigrotarsis and phnomus

it was apparent these 2 females did not belong to either indicus or nigrotarsis.

The above Thailand record for indicus is therefore incorrect.

The adult habitus of the 2 Thailand specimens compares well with that of

the type of phnomus, a species for which the female is unknown. There appears

to be no consistent difference in the adult habitus of the 3 included species in

this group (e.g. indicus, nigrotarsis and phnomus) except the absence of scales

on the paratergite of nigrotarsis and phnomus; however, these scales are mis-

sing (rubbed off?) in some specimens of indicus. These 3 species are easily

distinguished from each other and from other species of the subgenus by char-

acters of both the male and female genitalia.

Since the adults of the 2 females from Thailand can not be separated with

certainty, using habitus features, from indicus, nigrotarsis or phnomus, the

female genitalia are different from those of indicus and nigrotarsis, the fe-

male is not known for phnomus, and the collection sites of the 2 Thailand

specimens are close geographically and are apparently similar in climatic

and ecological conditions to the type locality of bhnomus, I am provisionally

assigning these 2 Thailand females to phnomus. This of course can not be

confirmed until progeny reared males and females with associated immature

skins are obtained.

After examining the types of both indicus and mediofasciata I agree with

Edwards (1922b: 468) that the 2 are conspecific.

TYPE-DATA. The holotype female of indicus bears the following label

data: Neomacleaya indica Type 2 F. V. T.; India, S. Christophers; Nov.

Spec., Nov. gen. nr. Macleaya, (a drawing of vertex of head and scutellum);

27[ on circular paper pining stage]; Received from F. V. Theobald, 1907-29;

T73.211 Term. [ my genitalia preparation number |; SEAMP Accession No.

374. The holotype, deposited in the BMNH is in good condition (head and one

leg are broken off and glued to the circular paper stage) and the genitalia are

mounted in Canada balsam on a microscope slide.

Theobald (1907: 544) originally described Skusea mediofasciata from 3 fe-

male and one male syntypes which are deposited in the BMNH. The male,

mounted in Canada balsam on 2 microscope slides (male genitalia and abdomen

on one slide and remaining parts mounted on second slide), is hereby designa-

ted lectotype and bears the following information on the slide labels: Skusea

mediofasciata n. sp. %, India, Received from F. V. Theobald, 1907-29,

SEAMP Accession No. 374; allolectotype is designated for one female and

bears the following data on the adult labels: Skusea mediofasciata (Type) F. V.

T., 27. F., India, S. Christophers, Received from F. V. Theobald, 1907-29. ,

SEAMP Accession No. 374; paralectotype designation is made for 2 females,

one with same label data as allolectotype except absence of 27.F., and the

other female with the following label information: India, S. Christophers, 18,

Received from F. V. Theobald, 1907-29., SEAMP Accession No. 374 (this

specimen I believe to be part of the original type series because of the same

handwritten Christophers label, the received from Theobald label and the

same type of circular paper card mount as the other 2 females; however, this

specimen does not bear the identification label of Theobald). The abdomens

and genitalia of all 3 females are missing. _

DISTRIBUTION. 73 specimens examined: 329, 140, 4pl, 11, 3 L, 8 with

associated skins (71, 11 incomplete).

INDIA. 12 (holotype of indicus), 3°, 1% (1¢% lectotype, 1° allolectotype, -

08 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

and 22 paralectotypes of mediofasciata). Bihar, Pusa; 52, 1%. Delhi,

Roshanara Gardens; 3°, 1%. Punjab, Amritsar, Karnal; 4@ pl, 142, 30 pl,

idk 7¢. 4 pk bk 8. :

SRI LANKA. 12. Patanthaw; 19.

BIOLOGY. Immatures were collected in India from water in rain-filled

ditches and open pools (Barraud 1934: 286).

AEDES (VERRALLINA) JOHNS ONI LAFFOON

(Pigs. 22, 25, 59, 60)

DISCUSSION. Aedes johnsoni had previously been separated from the

other members of a species complex (i.e. adustus, andamanensis, margarsen,

and sohni) by characters of the male genitalia and geographical distribution.

Adults of johnsoni can now be easily distinguished from the other species of

the group by the absence of setae (Fig. 22) on the prosternum (see discussion

under adustus). Aedes johnsoni is most similar to margarsen and at present

can only be separated with certainty by the prosternal feature. When the pro-

sternum was used to separate the adults of these 2 species it was found a large

number of the paratypes of johnsoni possessed setae on this structure and

therefore belong to margarsen. With this finding, the geographical ranges of

these 2 species now overlap.

The basal mesal lobe of the male genitalia of johnsoni is illustrated (Fig. 25)

and compared to the other members of the group. Illustrations are given for

the first time of tarsomeres 5 and posttarsi of the male (Fig. 59) and the fe-

male (Fig. 60).

DISTRIBUTION. 25 specimens examined: 199, 6c.

PHILIPPINES. Basilan Island, Isabela; 3° (paratypes). Mindanao,

Pasanonco; 22, 2% (paratypes) and 1“ (holotype), San Ramon; 8°, 2% (para-

types), Kabakan; 62, 1.

The above specimens were the only ones of johnsoni remaining in the USNM

when the newly found character (presence or absence of setae on the prosternum)

was used. Thirty-seven female and 33 male paratypes of johnsoni had setae

present on the prosternum and are therefore transferred to margarsen (see

distribution of margarsen).

AEDES (VERRALLINA) JOHORENSIS NEW SPECIES

(Figs. 11, 61, 65)

FEMALE (Fig. 65). Head, Antenna dark brown, 0.96-1.10 length of pro-

boscis, pedicel mostly dark brown with a few small dark brown scales and

short fine hairs mesally, flagellomere 1 with basal 0.35-0. 50 pale and with

several small brown scales near middle; clypeus dark brown, bare; maxillary

palpus blackish-brown scaled, 4 segmented, 0.16-0.17 length of proboscis;

proboscis blackish-brown scaled, 1.11-1.24 length of femur I; eyes contigu-

ous; ocular setae blackish-brown and well developed; scales on head all

broad, blackish-brown and decumbent except for narrow curved white scales

forming a double row along coronal suture, a few on occiput and along ocular

line, and a narrow stripe of broad white scales extending from area in front

of antepronotum to ocular line, postgena with a few broad pale brown scales;

occiput with numerous brown erect forked scales. Thovax. Scutal integument

dark brown; scutum covered with narrow curved reddish-brown scales except

Reinert: New Interpretation of Vervallina o9

for the following areas which have narrow curved creamy-white scales; a

patch on anterior scutal fossal area and extending onto lateral scutal fossal

area, a small patch at scutal angle, a large patch on supra-alar area, and

along lateral and anterior margins of prescutellar space; prescutellar space

bare except for a small median cephalad area with scales; scutellum with

narrow curved creamy-white scales on each lobe; dark reddish-black setae

on the following areas: 5-7 median anterior promontory, numerous acrostichal

(anterior and posterior), numerous dorsocentral (anterior and posterior),

scutal fossal (5-9 anterior, 5-7 lateral and 1-2 posterior), numerous supra-

alar, 6-8 posterior medial scutal, 1 postalar callar and scutellar (6-11 long

and 4-6 short ones on lateral lobe, 5-8 long and 6-9 short ones on median

lobe); pleural integument dark brown; antepronota widely separated, scales

absent, 20-28 short and long dark setae; postpronotum with narrow curved

reddish-brown scales on upper area and a few broad white ones below, 6-8

posterior dark setae, a few short hairs on area cephalad of lower seta; pro-

pleuron with a patch of broad white scales, 16-26 setae; prosternum, sub-

spiracular area, paratergite, mesomeron and metameron bare; postspiracular

area without scales, 5-13 brown setae; mesepisternum with a large upper and

a small lower patch of broad white scales, 5-8 upper and 14-21 posterior

setae, a patch of short hairs cephalad of lower scale patch and another patch

cephalad of upper patch next to subspiracular area; prealar knob without

scales, 15-29 brown setae; mesepimeron with a patch of broad white scales

near middle, 24-55 setae dorsad of scale patch (several hair-like), numerous

moderately long fine golden hairs posterior of scale patch and extending to

near lower margin of sclerite. Legs. Coxae I-III each with several setae, I

with broad brown scales and a small dorsal patch of white ones on anterior

surface, II and III each with a patch of broad white scales on anterior surface;

trochanters I-III each with broad white scales; femora I and II with anterior

surfaces brown scaled except for a narrow ventral longitudinal stripe of white

scales on II, posterior surfaces of I and II each brown scaled with a longitudin-

al stripe of white scales, stripe dorsal on I and ventral on II, anterior and

posterior surfaces of each white scaled with a dorsal longitudinal stripe of

brown scales, stripe becomes broader distally and forms an apical band, I-III .

each with a few lateroapical white scales on anterior and posterior surfaces;

tibiae I-III each brown scaled, I and II also with a posterior longitudinal white

stripe, stripe ventral on I and mesal on II; tarsi I-III brown scaled; posttarsi

I-III (Fig. 61) each with 2 ungues, I and II with ungues equal each with a tooth,

III with ungues equal, both simple. Wing. Dorsal and ventral veins with dark

brown scales; alula with several moderately broad brown scales on margin;

upper calypter with several brown setae on margin; 2-3 remigial setae. Halter.

Pedicel pale; capitellum dark brown scaled. Abdomen. Terga dark brown

scaled, laterotergite of I with a rectangular patch of white scales, II-VII each

with a large laterobasal patch of white scales; sterna II-VII each white scaled

with a narrow apical band of brown scales, VIII brown scaled; terga and sterna

with a number of golden-brown setae, mostly along posterior margins. Geni-

talia (Fig. 11). Tergum VIII moderately to heavily pigmented, wide, base

slightly concave, apex very slightly concave, a number of short, moderately

long and long setae at apex (a number of them stout), short and moderately

long thin setae scattered over apical 0.42-0.47, basolateral seta minute,

covered with minute spicules, numerous broad scales covering apical 0. 71-

0.77, basal 0.45-0.50 retracted into segment VII, VIII-Te index 0. 49-0. 52,

VIII-Te/IX-Te index 3. 13-3. 26; sternum VIII moderately to heavily pigmented,

wide, base slightly concave mesally, apex with a small median indentation with

60 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

a small lobe on each side of midline, numerous short and moderately long

setae apically, more numerous and slightly stouter on apical lobes, numerous

short and a few moderately long setae scattered over apical 0.78-0.94, covered

‘with minute spicules, numerous broad scales covering most of apical 0. 85-0. 96,

apical intersegmental fold unpigmented, VIII-S index 0. 57-0. 66; tergum IX mod-

erately pigmented, wide and short, setae absent, covered with minute spicules,

IX-Te index 0.25-0. 28; insula unpigmented, short, covered with minute spicules,

5-6 small tuberculi, each with a minute spicule; lower vaginal lip heavily pig-

mented, narrow, a few spicules on hinge area, with a small caudally projected

moderately pigmented median apodeme, lower vaginal sclerite composed of -

small elongated heavily pigmented plates situated on median basal area of vagin-

al floor; upper vaginal lip heavily pigmented, moderately broad, posterolateral

areas produced into small rounded lobes, median posterior area produced into

a small shield caudad of spermathecal eminence, upper vaginal sclerite pro-

duced into a moderately broad heavily pigmented transverse band which pro-

jects mesad from basal area of upper vaginal lip and forms a continuous crown

around the posterior margin of the spermathecal eminence; spermathecal

eminence heavily pigmented, large, moderately deep in depth, dorsal outline

somewhat ovoid with basal area flat, short spiny excrescences on basal and

basolateral areas, a few longer and stouter ones at basolateral corners; post-

genital lobe short, moderately broad, apex with a small median indentation, ~

20-25 setae on each side of midline, completely covered with minute spicules,

dorsal PGL index 0. 58-0. 69, ventral PGL index 0.76-0.91; peri-anal membrane

with minute spicules; cercus triangular in shape, moderately long, broad at

base, apex acute, completely covered with minute spicules, dorsal surface

with a number of moderately long and a few short setae scattered over apical

0.84-0.90, 2-3 long setae at apex, ventral surface with several short anda

few moderately long setae on outer 0.40-0.55, 2-6 broad scales on dorsal

surface, cercus index 2.58-2.97, cercus/dorsal PGL index 3. 55-3. 89; 3

seminal capsules, 1 large, 1 medium and 1 small in size, heavily pigmented,

elliptical, each with a number of small seminal capsule pores near orifice,

base of accessory gland duct lightly pigmented.

MALE, PUPA, LARVA AND EGG. Not known.

DISCUSSION. Adult females of johorensis are very similar in habitus to

members of the uwncus Group. They differ in having the lateral white scaled

patches of the abdominal terga larger in size. The description of the adult of

agrestis by Barraud (1931: 613) also appears to be similar to johorensis; how-

ever, a few differences appear to be present between the 2 species. The adult

body of the type of agrestis is lost so that these features can not be compared.

The female genitalia are most similar to those of petroelephantus. They

differ from those of this species, however, in the development of the lower

vaginal sclerite, upper vaginal lip, spermathecal eminence and tergum IX

(compare Figs. 11 and 15). The lower vaginal sclerite is similar to agrestis

but differs strongly in the other genitalia features (compare Figs. 3 and 11).

TYPE-DATA. Holotype female (genitalia mounted on a slide in Canada

balsam) with the following collection data: MALAYSIA, Johore, Pontian

Besar, 3 August 1969, Sulaiman bin Omar and Chia Yien Wang collectors,

collection number 2860, genitalia preparation number T74.66, MEP accession

number 409, collected as an adult in a biting collection on a clear day, among

coconut palms in a village, in partial shade at 1010 hours and at sea level; 6

female paratypes, same data as holotype except 3 with genitalia preparation

numbers 74/398, 74/399 and 74/448; and one female paratype genitalia (adult

lost), INDONESIA, Sumatra, Djambi, Moearatebo, XI. 1922-II. 1923, de Rook

Reinert: New Interpretation of Verrallina | 61

collector, collection number 9812, Ae. (Ae.) Sp. 6, 2 hypopygium, imago neg.

Holotype and 5 paratypes deposited in the USNM. One paratype will be de-

posited in the BMNH and one paratype will be deposited in the Shivaji Ramaling-

gam Collection at the University of Malaya. All types in excellent condition

except the adult body is missing for the Indonesian specimen.

DISTRIBUTION. Known only from the localities in Indonesia and Malaysia

where the type-specimens were collected.

BIOLOGY. As listed in the type-data.

AEDES (VERRALLINA) LANKAENSIS STONE AND KNIGHT

(Big, 12)

The following information on the female is based on a specimen from Col-

ombo, Sri Lanka and supplements the very brief description of Wijesundara

(1951: 174).

FEMALE. Head, Antenna dark brown, 1.12 length of proboscis, pedicel

brown with a few small brown scales and short fine hairs mesally; clypeus

brown, bare; maxillary palpus dark brown scaled, 0.18 length of proboscis;

proboscis dark brown scaled, 1.06 length of femur I; scales on head broad

dark brown and decumbent except a few narrow curved brown scales on coronal

suture and a stripe of broad white scales on lateral surface extending from in

front of antepronotum to ocular line; occiput with a number of short brown erect

forked scales, a few broad white decumbent scales and a number of narrow

curved brown ones. Thorax. Scutal integument dark brown; scutum covered

with narrow curved reddish-brown scales and with a few paler ones on median

anterior promontory and anterior scutal fossal areas; prescutellar space bare;

scutellum with a patch of narrow curved reddish-brown scales on each lobe;

dark brown setae on following areas: 2 median anterior promontory, numer-

ous acrostichals (anterior and posterior), numerous dorsocentrals (anterior

and posterior), scutal fossal (4-5 anterior, 3-4 lateral and 1 posterior), nu-

merous supra-alar, 6 posterior medial scutal, 1 postalar callar and scutellar

(6 long and 2-3 short ones on lateral lobe, 5 long and 4 short ones on median

lobe); pleural integument dark brown; antepronotum without scales, 13-14 dark

setae; postpronotum with narrow curved brown scales on upper area, 4-5 dark

posterior setae; propleuron with a patch of broad white scales, 9-10 setae;

prosternum, subspiracular area, paratergite, mesomeron and metameron

bare; postspiracular area without scales, 3-4 dark setae; mesepisternum with

an upper and a lower patch of broad white scales, 3 upper and 11-12 posterior

setae, lower ones shorter and pale; prealar knob with 8-9 dark setae; mese-

pimeron with a patch of broad white scales near center, 11 setae dorsad of

scale patch, several thin golden hairs along posterior and lower posterior

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

broad brown scales and a small dorsal and a small anteroventral patch of white

scales, II and III each with a patch of broad white scales on anterior surface;

femora I and II each with anterior surface brown scaled, posterior surface

brown scaled with a longitudinal white scaled stripe, ventral on I and dorsal

on II, II with most of anteroventral and posteroventral areas white scaled,

remainder brown scaled; tibiae I-III brown scaled, I also with a posteroventral

longitudinal white scaled strip, II also with a posteromedjan longitudinal white

scaled stripe; tarsi I-III each brown scaled; posttarsi I-III each with 2 ungues,

I and II with ungues equal, each with a tooth, III with ungues equal, both simple.

Wing. Dorsal and ventral veins with brown scales; alula with several narrow

62 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

dark brown scales on margin; upper calypter with several dark setae on margin;

2 remigial setae. Halter. Pedicel pale; capitellum brown scaled. Abdomen.

Terga each dark brown scaled with a large laterobasal white scaled patch; sterna

each white scaled with a narrow apical brown scaled band; terga and sterna with

numerous brown setae, mostly along posterior margin. Genitalia (Fig. 12).

As illustrated; spermathecal eminence long and narrow; upper vaginal sclerite

with a broad anteromesal extension; and lower vaginal sclerite small, moder-

ately pigmented and spiculate.

DISCUSSION. The female genitalia are illustrated (Fig. 12) and show for the

first time the lower vaginal lip and sclerite, terga VIII and IX, sternum VUI

and seminal capsules. The male genitalia were described and illustrated by

Edwards (1917: 221).

DISTRIBUTION. 2 specimens examined: 19, 1c.

SRI LANKA. Colombo; 1“ (holotype) and 18.

AEDES (VERRALLINA) LATIPENNIS DELFINADO

(Figs. 13, 61)

FEMALE. Thovax. Mesepimeron with numerous fine hairs caudad of scale

patch and extending to near lower margin of sclerite. Legs. Posttarsi I-III

(Fig. 61) each with 2 ungues, I and II with ungues equal in size, each with a

tooth, IT] with ungues equal, both simple. Genitalia (Fig. 13). Tergum VIII

moderately pigmented, wide, base slightly concave, apex broadly convex,

basolateral seta present, index 0.41-0.42; sternum VIII moderately pigmented,

wide, apical margin with a small lobe on each side of midline and a larger

lobe on lateroapical area, index 0. 53-0. 56; lower vaginal sclerite formed into

2 large, heavily pigmented, reticulated plates which are spiculate and narrowly

separated.

PUPA. Chaetotaxy as recorded in Table 9. Ocular plate with a small

area of moderately developed cuticular facets. Respiratory trumpet, Moder-

ately pigmented, hair-like spicules on distal 0.73-0.90 of inner surface; index

3.73-4.33, average 4.12. Abdomen. Seta 6, 9-VII weakly developed; 4- VIII

triple to 4 branched; 9-VIII single. Paddle. Ovoid; minute serrations on most

of basal 0. 67-0. 69 of outer margin, more strongly developed on distal area;

minute spicules on apical 0.31-0.43 of outer and apical 0. 08-0.15 of inner

margins; midrib does not reach apex; seta 1-P moderately long, single; index

1.41-1.67, average 1.57. | )

LARVA. Chaetotaxy as recorded in Table 22. Head. Seta 4-C with 4-8

branches; 5-C triple; 6-C single to triple; 7-C with 7-11 branches; 12-C with

4-8 branches; dorsomentum with 39-40 teeth. Antenna. Moderately long,

moderately pigmented, with a number of small scattered spicules; seta 1-A

triple to 5 branched, moderately long; 2-A long, with a subapical constriction;

3-A short, 0.27-0.33 length of 2-A; 4-A moderately long, 0.44-0.61 length of

2-A; 6-A short, 0.22-0.30 length of 2-A. Abdomen. Seta 1-VIII triple to 5

branched; 3-VIII triple to 6 branched; 5- VIII with 5-8 branches; 2-X with 5-7

branches; ventral brush with 15-18 setae on grid and 3-4 precratal shorter

ones, all multiple branched; saddle moderately pigmented, incompletely rings

segment X, acus absent; 4 moderately long anal papillae; comb with 8-12

(usually 12) scales arranged in a single curved irregular row, each scale with

a stout median apical spine and small denticles on lateral areas. Siphon.

Moderately pigmented; acus well developed; index 1. 77-2.33, average 1.97;

pecten on basal 0. 64-0. 68 of siphon, composed of 12-16 (usually 14) teeth,

Reinert: New Interpretation of Vervallina 63

apical 2-3 teeth longer and wider spaced than remainder of teeth, each tooth

with 1-2 small stout ventral denticles near base; seta 1-S short, 5-8 branched,

base attached on basal 0. 71-0. 74 of siphon distad of last pecten tooth.

DISCUSSION. Aedes latipennis belongs to the wncus Group of species (see

discussion under uncus). The male and female genitalia are very distinctive

and readily separated from the other species of the group and subgenus. The

paraproct of the male genitalia is unusually developed and is long, broad, flat

and with 2-3 long setae. The female genitalia have sternum VIII with latero-

apical lobes which is distinctive and unusual for the subgenus. The larvae

are easily separated from other members of the subgenus in the development

of the ventral brush of segment X which has 15-18 setae on the grid and 3-4

precratal setae. The larva and pupa are fully described above.

TYPE-DATA. The type-data of the holotype of latibennis, taken from the

collection data sheet, are amended as follows: male with associated pupal and

larval skins, THAILAND, Songkhla, Amphoe Haad Yai, 2 km from weather

station below Ton Nga Waterfalls, 26 March 1965, E. L. Peyton collector,

collection number SL-84-11, collected as a larva from water in a large tem-

porary flood pool in a rocky depression near a stream, in partial shade, at

an altitude near 250 feet, and ina primary forest. The allotype and paratypes

with same data as holotype except collection numbers which are in SL-82

series. The paratype male also has a collection number of SL-84 and is from

the holotype locality and not from Nakhon Si Thammarat.,

DISTRIBUTION. 22 specimens examined: 492, 20, 6 L, 5 with associated

pupal and larval skins.

THAILAND. Pyvachin Buri, Ban Bu Phram; 22 pl, 1L. Songkhla, Amphoe

Haad Yai, Ton Nga Waterfalls; 1“ pl (holotype), 12 pl (allotype), 12 pland1¢

(paratypes), 5 L.

BIOLOGY. Larvae were collected from a small ground pool and from a

large flood pool in a rocky depression near a stream; from clear, fresh,

unmoving, temporary water in partial shade; once in a primary forest and

once in a secondary scrub area in a valley; and at elevations from 76-150 m.

AEDES (VERRALLINA) LEICESTERI EDWARDS

(Figs. 14, 42, 56, 61)

FEMALE. Head. Antenna dark brown, 0.96-1.01 length of proboscis,

pedicel brown with a few small brown scales and short fine hairs mesally,

flagellomere 1 with basal 0.4 pale and with several small brown scales near

middle; clypeus dark brown, bare; maxillary palpus blackish-brown scaled,

4 segmented, 0.14-0.15 length of proboscis; proboscis blackish-brown scaled,

1.14-1.17 length of femur I; eyes nearly contiguous in front; ocular setae

dark brown and well developed; scales on head all broad and decumbent except

for a double row of narrow curved white scales along coronal suture, a few

similar ones on occiput and occiput with a patch of dark brown erect forked

scales; vertex blackish-brown scaled; lateral surface blackish-brown scaled

with a narrow stripe of white scales extending from area in front of antepro-

notum to ocular line; occiput with a few broad white scales mixed with narrow

scales. Thorax. Scutal integument brown; scutum covered with narrow curved

reddish-black scales except for a small patch of narrow: curved white scales on

median anterior promontory area and anterior scutal fossal area; prescutal

space mostly bare, however, with a small median cephalad patch of narrow

curved dark scales; scutellum with a patch of narrow curved reddish-black

64 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

scales on each lobe; dark reddish-black setae on the following areas: 4-5 medi-

an anterior promontory, numerous acrostichal (anterior and posterior), numer-

ous dorsocentral (anterior and posterior), scutal fossal (4-7 anterior, 3-6

lateral and 1-2 posterior), numerous supra-alar, 6 posterior medial scutal, 1

postalar callar and scutellar (5-7 long and 4-5 short ones on lateral lobe, 5-6

long and 4-6 short ones on median lobe); pleural integument brown; antepronota

widely separated, scales absent, 10-14 dark setae; postpronotum with narrow

curved reddish-black scales on upper area, 5-7 dark posterior setae and a

patch of short fine hairs cephalad of setae; propleuron with a patch of broad white

scales, 19-19 brown setae; prosternum, subspiracular area, paratergite, meso-

meron and metameron bare; postspiracular area without scales, 5-8 dark setae;

mesepisternum with a large upper patch of broad pale scales with a brown hue

and a small lower posterior patch of broad white scales, 4-7 upper and 15-19

posterior setae, a patch of short fine hairs cephalad of upper scale patch near

subspiracular area; prealar knob without scales, 14-19 dark setae; mesepimeron

with a patch of broad white scales near center, 27-29 setae dorsad of scale

patch, numerous fine golden hairs cephalad of scale patch and extending to near

lower margin of sclerite. Legs. Coxae I-III each with several setae, I with

broad brown scales and a small dorsal patch of white ones on anterior surface,

II and III each with a patch of broad white scales on anterior area; trochanters

I-III each with broad pale scales; femora I and II with anterior surfaces brown

scaled, II also with a longitudinal stripe of pale scales on anteroventral area,

I and II with posterior surfaces brown scaled with a longitudinal white scaled

stripe, stripe dorsal on I and ventral on II, III dark brown scaled with an antero-

ventral and a posteroventral stripe of white scales, stripe broad at base and

narrower apically; tibiae I-III each brown scaled, I with a posteroventral longi-

tudinal white scaled stripe from base to apex, II with posterior surface white

scaled; tarsi I-III brown scaled; posttarsi I-III (Fig. 61) each with 2 ungues,

I and II with ungues equal, each with a tooth, III with ungues equal, both simple.

Wing. Dorsal and ventral veins with dark brown scales; alula with several

moderately broad dark brown scales on margin; upper calypter with several

setae on margin; 1 remigial seta. Halter. Pedicel pale; capitellum dark brown

scaled. Abdomen. Terga blackish-brown scaled, laterotergite of I with a rec-

tangular patch of broad white scales, II-VII each with a large laterobasal patch

of broad white scales, patches extend onto dorsolateral areas of V-VII; sterna

white scaled each with a narrow apical band of broad brown scales, VIII brown

scaled; terga and sterna with numerous golden or brown setae, mostly along

posterior margins. Genitalia (Fig. 14). Tergum VIII moderately pigmented,

wide, base very slightly concave, apex slightly convex, a number of short,

moderately long and long setae at apex, a number of setae stout, several short

and moderately long setae on apical 0.33-0.42, basolateral seta absent, covered

with minute spicules, numerous broad scales on apical 0. 68-0.76, basal 0. 40-

0.45 retracted into segment VII, VII-Te index 0.48-0.50, VIII-Te/IX-Te index

3.97-4.17; sternum VIII moderately pigmented, wide, base slightly concave

mesally, apex with a shallow median indentation with a small lobe on each side

of midline, numerous moderately long and short setae apically, more numerous

and slightly stouter on apical lobes, numerous short and a few moderately long

setae scattered over apical 0.88-0.91, covered with minute spicules, numerous

broad scales covering most of apical 0.68-0.76, apical intersegmental fold

unpigmented, VIII-S index 0. 62-0. 67; tergum IX moderately pigmented, wide,

base concave mesally, setae absent, covered with minute spicules, index

0.19-0.20; insula unpigmented, short, covered with minute spicules, 4-6 small

tuberculi, each with a minute spicule; lower vaginal lip heavily pigmented,

Reinert: New Interpretation of Vervallina 65

narrow, a few spicules on hinge area, with a small caudally projected moder-

ately pigmented median apodeme, lower vaginal sclerite broad with a moder-

ately deep median apical indentation and usually 1-3 small holes along distal

portion of midline, base with a small median indentation; upper vaginal lip

heavily pigmented, moderately broad, posterolateral areas rounded, median

posterior area formed into a large upright shield caudad of spermathecal em-

inence, upper vaginal sclerite heavily pigmented, large, complex, forms a

band from upper vaginal lip to and caudad of spermathecal eminence; spermathe-

cal eminence heavily pigmented, large, somewhat diamond-shaped in dorsal

outline, ventral outline heart-shaped, spiny excrescences, short, with lateral

projections and along laterobasal and basal areas; postgenital lobe short, mod-

erately broad, apex with a small median indentation, 11-19 setae on each side

of midline, completely covered with minute spicules, dorsal PGL index 0. 6-

0.7, ventral PGL index 0. 88-1.03; peri-anal membrane with minute spicules;

cercus triangular in shape, moderately long, broad at base, apex acute, com-

pletely covered with small spicules, dorsal surface with a number of moder-

ately long and a few short setae scattered over apical 0. 89-0.92, 2 longer

ones at apex, ventral surface with several short and a few moderately long

setae on outer 0.4-0.5, 1-6 broad scales on dorsal surface, cercus index

2.40-2.79, cercus/dorsal PGL index 4. 04-4.54; 3 seminal capsules, 1 large,

1 medium and 1 small in size, heavily pigmented, spherical, each with a broad

neck, several seminal capsule pores near orifice, base of accessory gland duct

lightly pigmented.

MALE. Similar to female in general habitus but with the following differ-

ences. Head. Antenna plumose, 0.81-0.88 length of proboscis; maxillary

palpus 0.12-0.14 length of proboscis; proboscis 1. 04-1.16 length of femur I.

Thorax. Antepronotum with 8-12 setae; prealar knob with 8-14 setae; mese-

pimeron with 18-24 setae dorsad of scale patch. Legs. Posttarsi I-III

(Fig. 56) each with 2 ungues, I and II with ungues unequal, larger ungue of

each with a tooth, III with ungues equal, both simple. Abdomen. Tergum VIII

brown scaled with a laterobasal patch of broad white scales. Genitalia. As

figured by Delfinado (1968, Fig. 8).

PUPA (Fig. 42). Chaetotaxy as figured and recorded in Table 10. Ocular

plate with moderately developed cuticular facets. Respivatory trumpet.

Heavily pigmented; hair-like spicules on distal 0.75-0.77 of inner surface;

index 2.95-3.60, average 3.2. Abdomen. Setae 6, 9-VII weakly developed;

4-VIII double to 5 branched; 9-VIII single. Paddle. Ovoid; minute serrations

on most of basal 0. 70-0.72 of outer margin; minute spicules on apical 0. 28-

0.30 of outer and apical 0.19-0.24 of inner margins; midrib does not reach

apex; seta 1-P moderately long, single or double; index 1.37-1.50, average

1.42.

LARVA (Fig. 56). Chaetotaxy as figured and recorded in Table 23. Head,

Seta 4-C with 5-7 branches; 5-C triple; 6-C missing on all specimens; 7-C

with 8-12 branches; 12-C triple to 7 branched; dorsomentum with 35 teeth.

Antenna. Moderately long, moderately pigmented, with several scattered

small spicules, more numerous on basal area; seta 1-A with 4-5 branches,

moderately long, branches thickened; 2-A long, with a subapical constriction;

3-A short, 0.19-0.23 length of 2-A; 4-A moderately long, 0.42-0.47 length of

2-A; 6-A short, 0.29-0.32 length of 2-A. Abdomen. Setae 1, 2-VIII on small

common basal plate; 1-VIII with 5-7 branches; 3-VIII with 8-10 branches; 5-VIII

with 7-13 branches; 2-X with 6-8 branches, each approximately equal in length;

ventral brush with 10-11 setae on grid and 2 precratal shorter ones, all mul-

tiple branched; saddle moderately pigmented, incompletely rings segment X,

66 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

acus absent; 4 moderately long anal papillae; comb with 8-12 (usually 10) scales

arranged in a single curved irregular row, each scale with a stout median api-

cal spine and small denticles on laterobasal areas. Siphon. Moderately pig-

mented; acus well developed; index 2.21-2.47, average 2.31; pecten on basal

0. 58-0. 63 of siphon, composed of 11-15 (usually 14) teeth, apical 2-4 teeth

longer and wider spaced than remainder of teeth, each tooth long, slender

and with 1-2 small stout ventral denticles near base; seta 1-S short, with 9-10

branches, base attached on basal 0.65-0. 71 of siphon distad of last pecten tooth.

DISCUSSION. Aedes leicesteri belongs to the wncus Group of species. The

adults are very similar to the other members of the group in habitus features

and therefore can not be distinguished from them with certainty. The female

and male genitalia of leicesteri, however, are quite distinctive and easily

separated from the other members of the group and subgenus. The shape of

the lower vaginal sclerite of the female genitalia and the development of the

gonocoxite projections and gonostylus of the male genitalia are the most strik-

ing differences from the other species of the group.

Pupae of this species are very similar to the other members of the group.

The larvae are also similar to the other species of the wzncus Group but differ

in the very long anal papillae and the shortness of antennal seta 3-A.

The female, female genitalia, pupa and larva are described and illustrated

for the first time. Male posttarsi are illustrated for the first time and addi-

tional information on the adult male is presented.

DISTRIBUTION. 30 specimens examined: 692, 80°, 10 with associated

skins (4 p, 61).

MALAYSIA. Selangor, Templer Park; 5¢ pl, 12 p, 1¢% pl, 3% p, 3c; Kuala

Lumpur, Ampang jungle; 1 (holotype).

BIOLOGY. Immatures were collected in Malaysia from small ground pools

containing clear, fresh, unmoving, temporary water in heavily shaded areas

located in a secondary rain forest in hilly terrain and at an elevation of 152 m.

AEDES (VERRALLINA) LUGUBRIS BARRAUD

(Figs. 23, 43, 61)

PUPA (Fig. 43). Chaetotaxy as figured and recorded in Table1l. The

description and table are based on 5 skins and represent the first description

of the pupa of this species. Cephalothorax and abdominal segments with

heavily pigmented areas. Respiratory trumpet. Moderately pigmented; index

4,.75-5.67, average 5.11. Abdomen. Setae 6, 9-VI weakly developed; 4-VII

double or triple; 9-VIII triple to 6 branched. Paddle. Ovoid; minute serra-

tions on distal portion of basal 0.49 of outer margin; minute spicules on api-

cal 0.51 of outer and apical 0.32 of inner margins; midrib reaches apex, how-

ever, apical portion lightly pigmented; seta 1-P moderately long, single or

double; index 1.20-1.36, average 1.27.

DISCUSSION. See discussion section of butleri.

The description and illustration of butleri male genitalia by Edwards in

Barraud (1934: 296) belong to lugubris, however, the description of the female

was attributed to the correct species.

The posttarsi (Fig. 61) and tergum IX (Fig. 23) of the female of lugubris

are illustrated for the first time.

DISTRIBUTION. 15 specimens examined: 62, 40, lp, 4 with associated

pupal skins.

if

F

ae

Reinert: New Interpretation of Verrallina 67

MALAYSIA. Selangor, Kuala Selangor, Rantau Panjang; 32 p, 22, 1° p,

ao, kp.

THAILAND. Tvat(?); 12 (one female with genitalia preparation containing

the following data: STM-1-A, Sept. 1963, Thailand, probably Trat Province).

BIOLOGY. In Malaysia pupae were collected from fresh, colored, unmov-

ing water in small marshy depressions, in a partially shaded area located ina

coconut orchard. Females in this country were collected biting man.

AEDES (VERRALLINA) MACRODIXOA DYAR AND SHANNON

(Figs. 23, 59, 61)

DISCUSSION. For a discussion of macrodixoa and comparison with neo-

macrodixoa see Laffoon (1946: 238) and King and Hoogstraal (1947: 124).

Illustrations are presented for the first time of the tarsomeres 5 and

posttarsi of the male (Fig. 59) and the female (Fig. 61) and tergum IX of the

female genitalia (Fig. 23).

DISTRIBUTION. 29 specimens examined: 198, 10.

PHILIPPINES. Leyte, Tacloban; 32, 3c°. Luzon, Tayabas, Infanta; 1“

(holotype). Mindanao, Kabakan, Mercedes, Parang (Ludlow Barracks); 139,

30°. Mindoro, San Jose; 32, 3c. |

BIOLOGY. Immatures have been collected from rain pools and adults col-

lected resting in military foxholes.

AEDES (VERRALLINA) MARGARSEN DYAR AND SHANNON

(Figs. 22, 23, 25, 59)

DISCUSSION. Aedes margarsen belongs to the andamanensis Group of

species. It is very similar to johnsoni and is discussed under that species.

Adults are, however, readily distinguished from johnsoni by the presence of

setae on the upper mesal area of the prosternum (Fig. 22) while these are

absent in johnsoni,

Tergum IX of the female genitalia (Fig. 23), basal mesal lobe of the male

genitalia (Fig. 25) and tarsomeres 5 and posttarsi of the male (Fig. 59) are

illustrated for the first time. :

A label on the lectotype male gives the genitalia preparation as slide 36. IX.

10d. This information had not previously been published.

DISTRIBUTION. 192 specimens examined: 722, 61°, 29 individual rear-

ings with pupal and larval skins.

PHILIPPINES. Luzon Island, Camp Eldridge, 1% (lectotype) and 49 and

30° (paratypes); La Union, Anastacio, 8°, 80°; Zambales, Subic Bay, 22, 1°;

Clark Air Force Base (?), 212 pl, 8c pl, 60° (Cl 337, Baisas). Mindoro,

Kabakan, 139, 90°; San Jose, 12, 1°; San Ramon, 12, 1°.

The following specimens are from the paratype series of johnsoni but be-

long to margarsen.,

PHILIPPINES. Basilan Island, Isabela, 3%. Leyte, Tanauan, 22. Min-

danao, Pasanonco, 182, 60°; San Ramon, 3, 120°; Zamboaga, 2°.

All the above specimens are in the USNM.

68 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

AEDES (VERRALLINA) NEOMACRODIXOA KING AND HOOGSTRAAL

(Figs. 23, 61)

DISCUSSION. Belkin (1962: 422, Fig. 294) described and illustrated, from

4 whole larvae, what he termed an undetermined species and subgenus of

Aedes from Bougainville, Solomon Islands. After examining the above speci-

mens (kindly loaned to me by Dr. John N. Belkin) and comparing them with a

larval skin of Aedes neomacrodixoa from Papua, New Guinea, I found the 2 to

be conspecific. Label information on these 4 larvae is-as follows: Bougain-

ville, Solomon Is., May 10, 1944, A. B. Gurney, 372, Ground pools, Aedes

(?) sp. 1 det. J. N. Belkin. This record extends the northeastern range of

neomacrodixa to the Solomon Islands. Belkin in his discussion of the above

species stated the larvae possessed some features of species in the subgenera

Verrallina and Aedimorphus but they could not be placed in any recognized sub-

genus. Several circumstances probably prevented Belkin from recognizing

that his larvae belonged to the subgenus Vervallina, they are: (1) his study

included only species in Section A, Series I (neomacrodixoa belongs to Section

A, Series III), (2) larvae of neomacrodixa were not described until later (Huang

1968: 42), and (3) many species of the subgenus had the immatures unknown.

Sloof (1972: 178) also reported larvae of this species from Sipozae Island,

Choiseul, Solomon Islands.

Tergum IX (Fig. 23) of the female Sabena and female posttarsi (Fig. 61)

are illustrated for the first time.

DISTRIBUTION. 56 specimens examined; 212, 280, 4 L, 2 with associ-

ated skins (1 p, 11).

AUSTRALIA (Trust Territories). New Guinea, Northeast, Gusika, Fin-

schhafen area; 132 and 150" (paratypes), 22, 3°. Papua, Vanapa River, Port

Moresby; 1% pl, 1% p. Solomon Islands, Bougainville; 4 L.

INDONESIA. Celebes, Kalawara; 12, 20°. Cevam, Noehoehai, Wahaai,

Warasiwa; 42, 40°. New Guinea, West Irian, Hollandia, 1“ (holotype), Cyclops

Mountains, Hollandia area, 12 and 1“ (paratypes).

BIOLOGY. A larva from Ceram was collected from a pool located ina

virgin forest.

AEDES (VERRALLINA) NIGROTARSIS (LUDLOW)

(Figs. 3, 59, 61)

DISCUSSION. A discrepancy was noted between the descriptions and illus-

trations of the lower vaginal sclerites of the female genitalia of nigrotarsis as

given by Laffoon (1946) and Delfinado (1968). After examining the holotype

genitalia in the USNM it was apparent Laffoon's account was correct. A des-

sciption and illustration (Fig. 3) of this structure from the holotype follows.

The lower vaginal sclerites consists of a large heavily pigmented sclerite with

eranular appearance and a median apical jagged indentation, attached basally

to cephalic margin of upper vaginal sclerite, and a pair of small moderately

pigmented, sigmoid shaped apical sclerites. The single male from Calicoan

Island, Philippines, reported by Delfinado (1968: 29) is a species of the sub-

genus Lorvania Belkin.

Tarsomeres 5 and posttarsi of the male (Fig. 59) and the female (Fig. 61)

are illustrated for the first time.

DISTRIBUTION. 86 specimens examined: 469, 40c".

PHILIPPINES. Leyte, Abuyog, Jinamic Island, Leyte Gulf area, Tacloban;

Reinert: New Interpretation of Verrallina 69

82, 17%. Luzon, Rizal, Camp Nichols, Clark Airforce Base, Infanta, Manila, Subic

Bay, Wack-Wack; 262, 130°. Mindanao, SanRamon, Zamboanga;22¢, 50°. Samar,

Osmena; 6, 4. 3

SOUTH VIETNAM. Binh Dinh, AnKhe; 22, 1% (these 3 specimens are in the USNM

and represent the first country record for nigrotarsis outside the Philippines).

BIOLOGY. In the Philippines immatures were collected from water in

grassy, temporary, ground pools and adults were taken in light traps and ata

light in a military tent. Adults were collected in light traps in South Vietnam.

AEDES (VERRALLINA) NOBUKONIS YAMADA

(Figs. 22, 61)

Aedes (Aedes) ishigakiensis Bohart, 1956. Bull. Brooklyn Entomol. Soc. 51:

32 (o*, 9*, P, L*). NEW SYNONYM.

The following information based on the types of nobukonis and ishigakiensis

supplements the descriptions of Yamada (1932), Sasa (1949), La Casse and Yamaguti

(1950), Bohart (1956) and Delfinado (1968).

FEMALE. Head, Antennal1.11-1.12 length of proboscis; maxillary palpus 0. 15-

0.17 length of proboscis; proboscis 0. 97-0. 98 length of femur I; eyes separated infront

with white scales on interocular space. Thorax. Following setae present: 2 median

anterior promontory; numerous acrostichal (anterior and posterior); numerous dor-

socentral (anterior and posterior); scutal fossal (3-4 anterior, 4-7 lateral, 1 median

and 1 posterior); numerous supra-alar; 4-6 posterior medial scutal; 1 postalar callar;

scutellar (3-6 long and 4-5 short ones on laterallobe, 4-5 long and 3-5 short ones on

median lobe); 7-9 antepronotal; 3-4 postpronotal; 7-10 propleural; 2-4 postspiracular;

3 upper and 6-10 posterior mesepisternal, lower ones shorter; 6-11 on prealar knob;

and mesepimeron with 6-9 dorsad of scale patch. Prosternum, subspiracular area,

paratergite, mesomeronand metameronbare. Postspiracular area with afew broad

white scales (rubbed off inseveral specimens). Genitalia (Fig. 22). As illustrated.

MALE. Head, Antenna plumose, 1.1 length of proboscis; maxillary palpus

0.17 length of proboscis; proboscis 1.04 length of femur I.

EGG. The following account of the egg is taken from Moriya etal. (1973:

o2, 54). The egg chorion has a surface sculpturing consisting of a net-like pat-

tern of cells. Each cell has a diameter of about 10 microns and under higher

magnification is composed of 5-10 convex structures, each of which also has a

net-like sculpturing. The authors give a scanning electron microphotograph of

the surface sculpturing of the egg (page 52).

DISCUSSION. Male and female paratypes of ishigakiensis were kindly com-

pared for me by Dr. Kazuo Tanaka with 5 syntype specimens (one male without

apical abdomenal segments and genitalia and 4 females) of nobukonis. Dr. Tanaka

also compared ishigakiensis with several males and females of nobukonis in Dr.

Manabu Sasa's collection. No differences were detected between the 2 species

and I am therefore considering them to be conspecific.

La Casse and Yamaguti (1950: 102) mistakenly reported pale broad scales

on the paratergite.

Aedes nobukonis is easily separated from the other members of Section B

by the distinctive male and female genitalia (Fig. 22) and in the adult from

all species except yusafi (see Reinert 1973b) by the presence of white scales

on the postspiracular area. Aedes yusafi differs from this species by the

very large laterobasal white scaled patches on the abdominal terga and 5-8

posterior pronotal setae compared to the small white patches and 3-4

70 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

posterior pronotal setae in nobukonis.

Tarsomeres 5 and posttarsi of the female are illustrated (Fig. 61) for the

first time. |

DISTRIBUTION. 44 specimens examined: 129, 4°, 6p, 101, 12 L.

JAPAN. Ryukyu Islands, Ishigaki Island, Mt. Banna; 1“ (holotype of

ishigakiensis), 122, 3%, 6p, 101, 12 L (paratypes of ishigakiensis).

Tanaka (1971: 4) recently recorded this species from Sakishima Gunto in

the Ryukyu Islands.

AEDES (VERRALLINA) NOTABILIS DELFINADO

(Figs. 13, 61)

The ent information supplements the original description of Delfinado

(1967: 28).

FEMALE. Thorax, Antepronotum with 14-18 dark setae; postpronotum

with numerous narrow curved reddish-brown scales on upper area, 4-6 dark

posterior setae; propleuron with broad white scales, 13-15 setae; prosternum,

subspiracular area, paratergite, mesomeron and metameron bare; postspiracu-

lar area with 4-6 brown setae; mesepisternum with an upper and a lower patch

of broad pale scales, 4-5 upper and several posterior setae; prealar knob with

9-12 dark setae; mesepimeron with a moderately large patch of broad white

scales near center, 14-19 setae dorsad of scale patch, 7-11 hairs caudad of

scale patch. Legs. Femora I and II each with anterior surface brown scaled,

I and II each with posterior surface brown scaled with a broad white scaled

longitudinal stripe on basal 0.9, I with stripe dorsal and II with stripe ventral,

III white scaled with a narrow dorsal brown scaled longitudinal stripe on anteri-

or and posterior surfaces from base to apex, stripe broader distally and rings

apical 0.1; tibiae I-III each brown scaled, I and II each also with an indistinct

posteriomedian white scaled longitudinal stripe; tarsi I-III each brown scaled;

posttarsi I and II (Fig. 61) each with 2 ungues, equal in size, each witha

tooth, III of all specimens missing. Abdomen. Terga each brown scaled with

a large laterobasal white scaled patch which extends slightly on to tergum;

sterna each white scaled with a narrow apical brown scaled band. Genitalia

(Fig. 13). Insula with 4 small tuberculi; lower vaginal lip heavily pigmented

and covered with small hair-like spicules; lower vaginal sclerite composed of

a pair of moderately pigmented ovoid plates with spicules (illustrated here for

the first time); and tergum IX wide, short and ribbon-like, index 0.24.

Shape of spermathecal eminence distinctive for notabilis.

DISCUSSION. The holotype data are amended as follows: THAILAND,

Chon Buri, Amphoe Bang Lamung, Khao Mai Keo, 7 October 1963, collection

number CL-1, collector Kol Mongkolpanya, genitalia preparation number 30.

iv.66, USNM type catalog number 69214, collected in a human biting collec-

tion. The above information is taken from the collection data sheet and adult

labels.

Posttarsus III of all 4 specimens examined was missing but probably con-

sists of 2 ungues which are equal in size and simple. Delfinado (1967: 28)

mistakenly reported the holotype posttarsus III was composed of 2 ungues

which were equal in size and each with a tooth. The holotype has all legs

broken off except the femur and tibia of the left hindleg; however, most of

the parts from the fore- and midlegs are glued to the paper specimen point.

The right foreleg is glued on the paper point, under the specimen, near the

normal position of the hindleg (both ungues equal in size and with a tooth) and

Reinert: New Interpretation of Verrallina 71

could have been mistaken by Delfinado for the hindleg; however, the femur and

tibia are marked as the foreleg in the above description (taken from the addi-

tional specimens). The sterna are also ornamented differently than the origin-

al description (see above).

The 2 basal hairy lobed structures of the postatrial plate of Delfinado

(1967: 28, Fig. 18) are the basal portion of the lower vaginal lip. This struc-

ture in Delfinado's holotype preparation is somewhat damaged. The lower vag-

inal lip, lower vaginal sclerites, terga VIII and IX and sternum VIII are illus-

trated (Fig. 13).

DISTRIBUTION. 4 specimens examined: 49.

THAILAND. Chon Buri, Amphoe Bang Lamung, Khao Mai Keo; 12 (holo-

type), 22. Nakhon Si Thammavat, Amphoe Chawang, Nabon, Ban Tha Phea; 19.

BIOLOGY. The holotype and 2 other females were taken in daytime human

biting collections. One female was collected from a light trap in a forest con-

taining light vegetation.

AEDES (VERRALLINA) PAHANGI DELFINADO

(Fig. 22)

DISCUSSION. The female genitalia of pahangi are similar to those of

abditus, These 2 species can be separated by the development of the median

portion of the upper vaginal lip, shape of tergum IX and the wrinkled appear-

ance of the seminal capsules in pahangi,

The median area of the upper vaginal lip and tergum IX of the female

genitalia are illustrated (Fig. 22), the latter structure for the first time.

These structures are compared to those of the other members of Section B

(Figs. 21 and 22).

DISTRIBUTION. 3 specimens examined: 39.

MALAYSIA. Pahang, Kuala Terenggan; 12 (holotype), 1° (paratype).

Selangor, Templer Park; 19.

BIOLOGY. One female in Malaysia was collected from a biting collection

on a clear day, in a heavily shaded area of a secondary rain forest in a valley,

on 3 February 1970 at 1300-1400 hours and at an elevation of 152 m.

AEDES (VERRALLINA) PANAYENSIS LUDLOW

(Figs. 25, 59)

DISCUSSION. Delfinado (1968: 32) recorded this species from Leyte (2

males which are butlevi and 2 females which could not be located) and Puerto

Princesa, Palawan (4 females which are butleri and one female which could

not be found), but these records were based on misidentifications as indicated

by the correct determinations included in the parentheses. Laffoon (1946:

242) selected a lectotype for this species and gives information about the type-

series.

Female genitalia (Fig. 25) of panayensis are similar to those of incertus

and varus (see discussion of varus).

Tarsomeres 95 and posttarsi of the male are illustrated (Fig. 59).

DISTRIBUTION. 202 specimens examined: 1489, 54c,

INDONESIA. Alor, Kalabahi; 12. Celebes;10. Ceram, Piroe; 192, 2°.

Kalimantan, Poeroek Tjahoe; 1°. Moluccas, Morotai Island, Point Gila;

Poepliara, Regoha; 42. New Guinea, Schouten Islands, Aeoki Island in

72 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

Padaido Group; 249, 1°. | |

MALAYSIA. Sabah, Kuala Penyu, Tawau, Semporna; 92, 5c.

PHILIPPINES. Basbas Island;.42. Mindanao, Zamboanga; 12. Palawan,

Balabac Island, Cape Melville; 12, 1°, Culion Island, San Pedro; 12, 2c.

Panay, Tloilo; 292 and 20c (paratypes), 1° (lectotype), 22, 1%. Samar,

Calicoan Island; 1°. Philippine Islands, no locality given; 89, 4.

There are 62 females and 15 males in the USNM with the following label

information: Jola Jola, P.I., Maj. Ewing Coll., Thru Miss Ludlow. These

specimens were apparently incorrectly labeled according to Laffoon (1946:

243).

BIOLOGY. The specimens from Malaysia were reared from immatures

whose exuviae were apparently lost. These immatures were collected from a

small ground pool containing clear, fresh, unmoving temporary water in an

unshaded area located in a coastal village at sea level and with coconut palms.

Females from Malaysia were collected from a biting collection on a clear

day, in a partially shaded area of a secondary rain forest in a plain terrain,

on 1 June 1970 at 1030-1100 hours and at an elevation of 15 m.

AEDES (VERRALLINA) PETROELEPHANTUS WIJ ESUNDARA

(Fig. 15)

The following information is based on the lectotype female and 2 paralecto-

type females and supplements the brief description of Wijesundara (1951: 176).

FEMALE. Head. Antennal pedicel with a few small brown scales and short

fine brown hairs mesally; vertex with a few narrow curved white scales along

coronal suture; eyes contiguous in front. Thorax. Scutum covered with nar-

row curved reddish-brown scales except for small patches of narrow curved

white scales on median anterior promontory and anterior scutal fossal areas;

scutellum with narrow curved reddish-brown scales on each lobe; prescutellar

space bare; numerous anterior and posterior acrostichal and dorsocentral

setae; antepronotum with scales absent, 19-21 brown setae; postpronotum

with narrow curved reddish-brown scales on upper area, 6 brown posterior

setae; propleuron with a patch of broad white scales, 19-20 brown setae;

prosternum, subspiracular area, paratergite, mesomeron and metameron

bare; postspiracular area with scales absent, 4-5 brown setae; mesepisternum

with a large upper and a smaller lower patch of broad white scales, 5-6 upper

and 13-14 posterior setae, lower ones shorter; mesepimeron with a large

patch of broad white scales near center, 20-21 setae dorsad of scale patch,

several small golden-brown hairs along posterior margin and lower posterior

corner of scale patch. Genitalia (Fig. 15). Spermathecal eminence and lower

vaginal sclerites with characteristic shapes.

DISCUSSION. Wijesundara (1951: 176) described petroelephantus from 3

female cotypes which were later sent to the BMNH. Unfortunately these adult

specimens, as well as other adult types in this shipment, were severely

damaged in transit so that presently only fragments of the adults remain.

~The cotypes of this species are represented now by only 2 pinned specimens

and 3 genitalia preparations (each in good condition and mounted in Canada

balsam on a microscope slide). Lectotype designation is made for one of the

cotype females with genitalia preparation and possesses the following label

information: Aéd. (Aéd.) sp. new, Kurunegala, 16-12-22, Med. Entomologist,

Ceylon, 2 Hypo. No. 85, cotype, Aédes (Aédes) petroelephantus Wijesundara

LECTOTYPE £&, selected by J. F. Reinert, 10 July 1974. Paralectotypes are

Reinert: New Interpretation of Vervallina 73

selected for the other 2 female cotypes which bear the same label data as the

lectotype except for the genitalia preparation numbers--84 and 86. The adult

of specimen No. 84 is lost and only the genitalia mount remains.

The adult of the lectotype is in very poor condition with only a twisted and

rubbed thorax and head ona pin. The genitalia were dissected and remounted

(10 July 1974) in Canada balsam and are in good condition.

AEDES (VERRALLINA) PHILIPPINENSIS DELFINADO

DISCUSSION. Delfinado (1968: 34) records the allotype female (with geni-

talia and tarsal claws on slides) deposited in the USNM. This type could not be

located in neither the USNM nor SEAMP collections.

DISTRIBUTION. One additional female of philippinensis was found among

unidentified Aedes in the USNM collection and at present represents the only

specimen of this species in this collection. The specimen bears the following

information on the labels: P. I., Leyte, Mahaplag, 11. VII. 64, Light trap,

M. Delfinado collector, genitalia preparation number T73. 542, Det. J. F.

Reinert.

AEDES (VERRALLINA) PHNOMUS KLEIN

DISCUSSION. The 2 female specimens recorded as indicus from Thailand

by Delfinado (1967: 25) were incorrectly identified and are here provisionally

assigned to phnomus (see taxonomic discussion section of indicus). Her Figure

18 also belongs to this species.

DISTRIBUTION. The following information was taken from the collection

data sheets for the 2 Thailand specimens: 12, collection number U-72, THAI-

LAND, Nakhon Ratchasima, Amphoe Muang, Pak Thong Chai, Nong Nam Keo,

30 May 1963, 1930-2000 hours, biting a buffalo and 12, collection number U-555,

THAILAND, Chon Buri, Amphoe Bang La Mung, Khao Mai Kaeo, 14 August

1963, 1900-2000 hours, biting collection.

I have also examined the holotype of phnomus from Chrui Changwar, Kandal,

CAMBODIA.

AEDES (VERRALLINA) PIPKINI BOHART

(Figs. 24, 61)

DISCUSSION. Bohart and Ingram (1946: 29) reported a single female Aedes

from Truk but they were unable to identify it. I have compared this specimen

with the holotype female and paratypes of Aedes pipkini and have determined

it to be conspecific with them. The adult (genitalia mounted in Canada balsam

on a slide) is in the USNM and contains the following information on the labels:

Moen I., Truk, XI-28-45, caught resting in occupied tent at 0700, R. L. In-

gram, 807.

Tergum IX of the female genitalia (Fig. 24) and female posttarsi (Fig. 61)

are illustrated for the first time.

74 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

AEDES (VERRALLINA) PRIOEKANENSIS BRUG

(Fig. 25)

The following information based on the holotype and specimen number 18. 220

supplements the descriptions of Brug (1931: 249) and Delfinado (1968: 34).

MALE. Head (head description taken from specimen number 18.220). An-

tenna brown, plumose, 1.0 length of proboscis, pedicel dark brown; clypeus

dark brown, bare; maxillary palpus dark brown scaled, 0.11 length of proboscis;

proboscis dark brown scaled, 1.23 length of femur I; eyes contiguous in front;

ocular setae dark brown; scales on head broad, dark brown and decumbent ex-

cept for a few narrow curved white scales along coronal suture, a small patch

of broad white scales on lateral surface in front of antepronotum next to ocular

line and a small patch of similar ones on postgena; occiput with several short

brown erect forked scales and a few decumbent broad white scales. Thorax,

Scutal integument reddish-brown; scutum covered with narrow curved reddish-

brown scales except for a few narrow curved white scales on median anterior

promontory area, anterior scutal fossal area, supra-alar area near base of

wing, median anterior area of prescutellar space and on median lobe of scutellum

(lateral lobes with narrow curved reddish-brown scales); dark brown setae on

following areas: 2 median anterior promontory; numerous acrostical (anterior

and posterior); numerous dorsocentral (anterior and posterior); scutal fossal

(3 anterior, 2-3 lateral and 1 posterior); numerous supra-alar, 4 posterior

medial scutal, 1 postalar callar and scutellar (3 long and 2-3 short ones on

lateral lobe, 4 long and 3 short ones on median lobe); pleural integument

dark brown; antepronotum with 7-8 brown setae; postpronotum with a few

narrow curved reddish-brown scales dorsally, 3 brown posterior setae; pro-

pleuron with several broad white scales, 11-14 setae; prosternum, subspiracular

area, paratergite, mesomeron and metameron bare; postspiracular area with

6 brown setae; mesepisternum damaged but with an upper and a lower patch of

broad white scales, several setae; prealar knob with 7-8 brown setae;

mesepimeron with a patch of broad white scales near center, 10-12 brown

setae dorsad of scale patch, a few short fine golden hairs caudad of lower

posterior corner of scale patch. Legs. Coxae I-III each with several brown

setae, I with broad brown scales and a small dorsal patch of broad white scales

on anterior surface, II and III each with a patch of broad white scales on anteri-

or surface; trochanters I-III each with a few pale brown scales; femora I and II

each with anterior surface brown scaled, posterior surface brown scaled with

a longitudinal pale scaled stripe from base to near apex, stripe dorsal on I and

ventral on II, II with anterior and posterior surfaces each mostly white scaled

and with a dorsal brown scaled longitudinal stripe from base to apex forming a

narrow apical brown scaled band; tibiae I-III and tarsi I-III brown scaled; post-

tarsi I-III each with 2 ungues, I and II each with ungues unequal, larger ungue

with a tooth and smaller one simple, III with ungues equal, one ungue with a

small tooth and other one simple. Wing. Dorsal and ventral veins with brown

scales; alula badly rubbed in both specimens but specimen 18.220 with a few

narrow brown scales on margin; upper calypter with several dark setae on mar-

gin. Halter. Pedicel pale; capitellum white scaled. Abdomen. Terga I-V

each brown scaled with a small laterobasal patch of white scales (remainder of

abdomen removed with genitalia); sterna brown scaled with pale basal scales;

terga and sterna with numerous golden-brown setae, mostly along posterior

margins. Genitalia (Fig. 25). Complete genitalia as illustrated by Delfinado

(1968: Fig. 8).

DISCUSSION. Aedes prioekanensis male adults (females unknown) are very

Reinert: New Interpretation of Vervallina 79

similar to those of incertus and varus. The male genitalia of the 3 species are

easily distinguishable by the development of the apical portion of the basal mesal

lobe (prioekanensis has a thin seta, incertus has a thick spiniform seta, and

varus has a fleshy projection with a frayed apex). The gonocoxite of the male

genitalia of prioekanensis and incertus also have an apicomesal projection

while this projection is absent in varus,

Posttarsus III of prioekanensis has 2 ungues which are equal in size and one

with a small tooth which differs from the males of both varus and incertus which

have the ungues of posttarsi III simple. The presence of a tooth on the ungue of

posttarsus III is very unusual for Section A but normal for Section B.

The abdominal terga of prioekanensis are brown scaled with small latero-

basal white scaled patches and posttarsus III, which had previously been

mounted, has ungues equal and only one with a small tooth. Delfinado (1968:

34) mistakenly reported the terga as wholly dark brown and both ungues of

posttarsus III toothed.

TYPE-DATA. Adult holotype specimen bears labels with the following

data: Aedes (Aedes) dermajoensis, n. sp.. VAR. prioekanensis, “ TYPE,

Brug; 18.219 | collection number]; Sumatra: Benkulen, xi-xii. 1929, Col.

Ss. L. Brug, B. M. 1930-511 | a BMNH printed label]; Larva in pool in virgin

forest; SEAMP accession number 376; and genitalia preparation number T73.

062. Type deposited in the BMNH. Brug's field collection notes for this speci-

men record the following information: de Rook | collector], Dermajoe (Ben-

koelen), Nov.-Dec. 1929, 18.219 type Br. Mus., Ae. prioekanensis, 18.219-

20 (20°) | 2 males from the collection]. Brug (1931: 249), however, reported

the following in the original description of this species 'Described from one

specimen bred from a larva found in a swamp in a virgin forest near Air

Prioekan.'' The second male, 18.220, recorded in Brug's field collection

notes is also in the BMNH and bears the following information on the adult

labels: 18.220; Co-type Ae. prioekanensis; and SEAMP accession number 30.

The holotype is in poor condition and is as follows: head and all appendages

missing; pinned through pleura of thorax with a minuten pin; all legs on left side

and right tarsus I mounted on a small celluloid point attached to pin; and left

wing missing. Genitalia remounted in Canada balsam on a microscope slide

from a celluoid point attached to pin. The specimen (18.220) labeled co-type

has the head and its appendages in good condition.

DISTRIBUTION. 2 specimens examined: 2c.

INDONESIA. Sumatra, Dermajoe (Benkoelen); 1“ (holotype), 1¢.

AEDES (VERRALLINA) PSEUDOMEDIOFASCIATUS (THEOBALD)

(Figs. 16, 59, 61)

The following information supplements the descriptions of Theobald (1910:

489) and Barraud (1928: 367).

FEMALE. Head, Antenna 1.11-1.14 length of proboscis, pedicel with a

few small brown scales and short fine hairs mesally; maxillary palpus 0. 17-

0.18 length of proboscis; proboscis 0.95-0.98 length of femur I. Thovax.

Antepronotum with 9-16 dark setae, scales absent; postpronotum with narrow

curved reddish-black scales on upper area, 5-7 dark posterior setae; propleuron

with a patch of broad white scales, 10-12 dark setae; prosternum, subspiracu-

lar area, paratergite, mesomeron and metameron bare; postspiracular area

without scales, 2-5 dark setae; mesepisternum with a large upper and a

small lower patch of broad white scales, 4-5 upper and 8-10 posterior setae,

76 Contrib: Amer, Ent. ‘ast, vol 11, no'1;1974

lower ones shorter; mesepimeron with a large patch of broad white scales near

center, 10-14 dark setae dorsad of scale patch, 3-5 short fine hairs near lower

posterior corner of scale patch. Abdomen. Sterna white scaled each witha

narrow apical band of brown scales. Genitalia (Fig. 16). As illustrated.

MALE. Head. Antenna 0.97-1.02 length of proboscis; maxillary palpus

0.10-0.13 length of proboscis; proboscis 1. 03-1.14 length of femur I. |

DISCUSSION. No consistent difference in adult habitus characters could be

found between pseudomediofasciatus and cautus; however, marked differences

exist between the genitalia of both females and males of the 2 species. These 2

species are also similar to sabahensis, new species, and can be separated

from it by features of abdominal markings and the propleuron.

Tarsomeres 5 and posttarsi of the male (Fig. 59) and the female (Fig. 61)

are illustrated for the first time.

DISTRIBUTION. 9 specimens examined: 49, 5c".

SRI LANKA. Central, Kandy District, Peradeniya; 12. North Central,

Anuradhapura District, Hunuwilagama, Wilpattu; 2°. Northern, Vavuniya

District, Parayanalankulan, 25 miles NW. of Medawachchiya; 22, 3%. Sabava-

gamuwa, Ratnapura District, Uggalkaltota; 12.

BIOLOGY. Adults were collected at elevations of 30 and 60 m.

AEDES (VERRALLINA) PSEUDOVARIETAS NEW SPECIES

(Figs. 31, 59)

MALE. Head. Antenna plumose, brown, 0.96 length of proboscis, pedicel

dark brown; clypeus dark brown, bare; maxillary palpus blackish-brown scaled,

0.12 length of proboscis; proboscis blackish-brown scaled, 1.15 length of femur

I; eyes contiguous; ocular setae blackish-brown and well developed; scales on

head all broad, blackish-brown and decumbent except for a few narrow curved

white scales along coronal suture, a narrow stripe of broad white ones extend-

ing from area in front of antepronotum to ocular line, and a small patch of dark

erect forked scales on occiput. Thovax. Scutal integument dark reddish-brown;

scutum covered with narrow curved dark reddish-brown scales; prescutellar

space bare except for a small median cephalad area of narrow curved scales;

scutellum with a patch of narrow curved reddish-brown scales on each lobe;

dark reddish-black setae on the following areas: 2 median anterior promontory,

numerous acrostichal (anterior and posterior), numerous dorsocentral (anterior

and posterior), scutal fossal (4 anterior and 3 lateral), numerous supra-alar,

4 posterior medial scutal, 1 postalar callar and scutellar (4 long and 2-3 short

ones on lateral lobe, 4 long and 3 short ones on median lobe); pleural integu-

ment brown; antepronota widely separated, scales absent, 7 long and short dark

setae; postpronotum with a few narrow curved reddish-brown scales on upper

area, 3 posterior dark setae; propleuron with a patch of broad white scales,

several setae; prosternum covered; subspiracular area, paratergite, mesomeron

and metameron bare; postspiracular area within a fold of the pleuron and not

visible; mesepisternum with a large upper and a small lower patch of broad

white scales, 3 upper and 7 posterior setae; prealar knob without scales, 5-6

dark setae; mesepimeron with a patch of broad white scales near center, 5-6

setae dorsad of scale patch, 2-4 short fine hairs near lower posterior corner

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

scales and a small dorsal patch of broad white ones on anterior surface, II and

III each with a patch of broad white scales on anterior surface; trochanters I-

III each with a few pale brown scales; femora I and II each with anterior surface

Reinert: New Interpretation of Vervallina 717

brown scaled, II also with an indistinct ventral longitudinal stripe of pale

scales, posterior surfaces of I and II brown scaled with a longitudinal pale

stripe, stripe dorsal on I and ventral on II, III with anterior and posterior

surfaces brown scaled with a ventral longitudinal stripe of white scales,

broader at base and tapering apically; tibiae I-III and tarsi I-III each dark brown

scaled; posttarsi I-III (Fig. 59) each with 2 ungues, I and II each with ungues

unequal, larger one with a tooth, smaller one simple, III with ungues equal,

both simple. Wing. Dorsal and ventral veins with dark brown scales; alula

with a few narrow brown scales on margin; upper calypter with several brown

setae on margin; 1 remigial seta. Halter. Pedicel pale; capitellum dark

brown scaled. Abdomen. Terga blackish-brown scaled, I-V each with a small

laterobasal patch of white scales (remainder of abdomen removed with genitalia);

sterna with basal areas pale scaled and apical areas brown scaled; terga and

sterna with several setae, mostly along posterior margins. Genitalia (Fig. 31).

Tergum IX moderately to heavily pigmented, band-like with lateral areas some-

what broader, outer caudal margin fused to tergum X, setae absent, broadly

connected laterally to sternum IX; gonocoxite short, broad, heavily pigmented,

mesal surface membranous with minute spicules, dorsal surface with apex

extended into a long narrow pointed mesally curved arm with a moderately

long stout seta near middle of dorsal area, a short heavily pigmented subapical

Spine also on dorsal surface, a long stout mesally curved seta at base of arm,

a few moderately long and long setae on dorsal surface mainly along mesal mar-

gin of gonocoxite, ventral surface with a long flat broad heavily pigmented api-

cal projection with a wrinkled and bluntly pointed apex, several moderately

long setae and a few long stout ones mainly on mesal surface of gonocoxite, a

few long setae on lateral surface, a number of broad scales on lower dorsal,

lateral and most of ventral surfaces; gonostylus heavily pigmented, long,

strongly curved near middle, apical 0.27 bifurcated, outer arm of furcation

longer, 4-5 moderately long setae on mesal margin of area between median

bend and furcation, gonostylus attached subapically on gonocoxite; basal mesal

lobe moderately pigmented, consists of a broad basal area with 3 long heavily

pigmented spiniforms (mesal one shorter than other 2) and a narrower apical

extension on mesal area of gonocoxite which projects slightly past the apex

and is covered with numerous short and moderately long thin setae, however,

setae more numerous on basal area, covered with small spicules, lobe con-

nected mesally with its mate by a moderately broad band which is covered with

small spicules and is located ventrad of apical portion of prosophallus and

phallus; proctiger with paraproct formed into a heavily pigmented long slender

acuminate arm with a sharp bend near basal area, base with a short projection

which extends ventrad and articulates at a point on the dorsomesal surface of

the parameral apodeme near base of opisthophallus, base of proctiger lightly

fused with tergum X, cercus membranous and without cercal setae; tergum X

moderately pigmented, consists of a small indistinct plate laterad of paraproct

base and extends ventrally from caudal margin of tergum IX; phallosome com-

plex, opisthophallus--consists of a lightly to moderately pigmented tergal trans-

verse bridge between parameral apodemes, caudal margin slightly indented me-

sally, sternobasal area attached to tergobasal portion of outer area of prosophal-

lus, located dorsad of phallus and prosophallus and ventrad of proctiger, phallus--

consists of a pair of moderately long, heavily pigmented, narrowly separated,

caudally produced arms with apical area curved dorsad, arms fused together

by a basal plate which extends laterally and is fused to basal portion of pro-

sophallus, prosophallus--consists of a pair of moderately long widely separated

arms with distal portion heavily pigmented and curved, proximal portion moder-

78 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

ately pigmented and formed into a broadly rounded area which is attached to ter-

somesal portion of paramere, located laterad of phallus, prosophallic arm

approximately equal in length to phallic arm, paramere--heavily pigmented,

approximately equal in length to phallic arm, articulates at a point approximate-

ly 0.6 from base with caudal area of parameral apodeme, parameral apodeme--

moderately to heavily pigmented, moderately long, outer surface broadly

attached to tergobasal apodeme of gonocoxite; sternum IX heavily pigmented

throughout, large, broad, lateroapical areas rounded, 18 short and moderately

long setae in an elongate patch near caudal margin.

FEMALE, PUPA, LARVA AND EGG. Not known.

DISCUSSION. The adult male habitus of pseudovarietas and varietas are

very similar and no consistent differences could be found with which to separate

the 2 species. They do, however, occupy different geographical areas.

The male genitalia of the 2 species differ and are easily separated. The

male genitalia of pseudovarietas differ most noticeably from those of varietas

by the following features: shape of the ventroapical projection of the gonocoxite,

development of the basal mesal lobe, and with fewer setae on sternum IX (com-

pare Fig. 31 and Delfinado's 1968 Fig. 10).

TYPE-DATA. Holotype male (genitalia mounted on a slide in Canada bal-

sam) with the following collection data: MALAYSIA, Sabah, Kudat, Matunggong,

30 April 1970, Samuel Wilson James and Chia Yiew Wang collectors, collection

number S-697, genitalia preparation number 74/379, MEP accession number

409, collected as an adult on a clear day, among coconut trees in a village, in

partial shade at 1100 hours and at an elevation of 61 m. Holotype is in good

condition and is deposited in the USNM.

DISTRIBUTION. Known only from the type-locality.

BIOLOGY. As listed in the type-data.

AEDES (VERRALLINA) RAMALINGAMI NEW SPECIES

(Figs. 32, 59)

MALE. Head. Antenna with flagellomeres dark brown basally, plumose,

1.16 length of proboscis, pedicel pale brown; clypeus brown, bare; maxillary

palpus dark brown scaled, 0.17-0.18 length of proboscis; proboscis dark brown

scaled, 1.08-1.14 length of femur I; eyes separated in front; several long

blackish-brown ocular setae; scales on head all broad, decumbent and dark brown

except for a few broad white ones on ocular line and a patch of similar ones an-

terodorsally on lateral surface next to eye and at level with antepronotum, and

a few short erect forked blackish-brown scales on occiput. Thorax. Scutal

integument brown; scutum covered with narrow curved dark reddish-brown scales,

scutellum with a few similar scales on each lobe; prescutellar space bare;

blackish-brown setae on following areas: 4 median anterior promontory, sev-

eral acrostichal (anterior and posterior), numerous dorsocentral (anterior and

posterior), scutal fossal (2 anterior, 2 lateral and 1 posterior), numerous supra-

alar, 4 posterior medial scutal, 1 postalar callar and scutellar (3-4 long and 1

short on lateral lobe; 3-4 long and 1-2 short on median lobe); pleural integument

pale brown; antepronotum with a few broad dusky-white scales, 7-8 dark brown

setae; postpronotum with 3-4 dark brown setae; propleuron with a few broad

white scales, 4-6 dark brown setae; prosternum, subspiracular area, parater-

site, mesomeron and metameron bare; postspiracular area with 2 dark brown

setae; mesepisternum with an upper and a small lower patch of broad white

scales, 1-2 upper and 5-7 posterior dark brown setae; prealar knob with 5 dark

Reinert: New Interpretation of Vervallina 79

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

4-5 golden-brown setae dorsad of scale patch. Legs. Coxae I-III each with

several dark brown setae, I with broad pale brown scales and a small dorsal

patch of white ones on anterior surface, II and III each with a patch of broad

white scales on anterior surface; trochanters I-III each with broad pale

scales; femora I and II each with anterior surface dark brown scaled, I and II

each with posterior surface brown scaled with a white scaled longitudinal stripe

from base to apex, stripe dorsal on I and ventral on II, III brown scaled with an

anteroventral and a posteroventral white scaled stripe which is broad basally

and narrow apically; tibiae I-III and tarsi I-III each dark brown scaled; post-

tarsi I-III (Fig. 59) each with 2 ungues, I, II and III each with ungues unequal,

each ungue with a tooth. Wing. Dorsal and ventral veins with dark brown

scales; alula with several narrow to moderately broad brown scales on margin;

upper calypter with several brown hairs on margin; 1 remigial seta. Halter.

Pedicel pale; capitellum dark brown scaled. Abdomen. Terga dark brown

scaled, I with a patch of broad white scales on laterotergite; sterna brown

scaled; terga and sterna with numerous dark setae, mostly along posterior

margins. Genitalia (Fig. 32). Tergum IX heavily pigmented, formed into a

narrow band mesally and expanded laterally into a small caudal lobe which is

fused with tergum X, setae absent, connected laterally to sternum IX by a

narrow heavily pigmented band; gonocoxite short, broad, heavily pigmented,

tergobasal portion of mesal surface membranous with minute spicules, dorsal

surface with apex extended into a small strongly sclerotized lobe which bears

2-3 long stout setae, 5-8 moderately long and long stout setae on distal 0.6 of

area basad of apical setae, ventral surface produced into a large apicomesal

lobe which is heavily pigmented on both tergal and sternal areas, tergal area

of lobe with a few short thin setae along outer margin, sternal area of lobe with

a number of short thin setae on apical 0.4, a few broad scales on ventral and

lateral surfaces of gonocoxite, lateral surface also with a few long stout setae;

gonostylus heavily pigmented, moderately long, approximately 0.68 length of

gonocoxite, basal 0.5 somewhat swollen, apical 0.5 bifid, outer arm with api-

cal portion somewhat expanded and with 4-5 short setae and 4-5 small stout

spiniforms, apex rounded, inner arm with proximal 0.52 swollen and with 5

short thin setae, distal 0.48 narrow, curved and with apex recurved; basal me-

sal lobe broad, lightly to moderately pigmented, covered with small spicules,

11-12 short thin setae on basomesal area, 5-7 short thin setae on mesal mar-

gin, caudal area with 5 short thin setae, caudomesal portion expanded into a

small apical hump and a short mesal heavily pigmented pointed projection with

a short seta at base (structure resembles a bird's head), sternomesal and cau-

dal margins of basal mesal lobe fused with gonocoxite, lobe connected mesally

with its mate by a moderately pigmented broad band which forms a shallow

trough ventrad of apical portions of opisthophallus and prosophallus; proctiger

with paraproct formed into a long narrow heavily pigmented curved arm, base

with a very short rounded projection which extends ventrad and articulates at

a point on the dorsal surface of parameral apodeme near base of opisthophallus,

base of proctiger strongly fused with tergum X, cercuSs membranous and with-

out cercal setae; tergum X heavily pigmented, formed into a small plate be-

tween proctiger and caudal lateral portion of tergum IX; phallosome complex,

opisthophallus--consists of a moderately pigmented tergal transverse bridge

between the parameral apodemes, caudal margin produced into a long lobe

which extends beyond apex of prosophallus and is broad at the base and tapers

to a pointed apex, base attached to a dorsomesal extension of the caudal portion

of parameral apodeme, located dorsad of phallus and prosophallus and ventrad

80 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

of proctiger, phallus--consists of a pair of short heavily pigmented narrowly

separated arms which are fused at their bases, sternolateral area of base ex-

tended laterally into a moderately long arm which is fused to basomesal portion —

of prosophallus, apex of phallus formed into a tergal penis filament which con-

sists of a pair of bars extending cephalad, bars connected near base by a broad

transverse bridge, apical portion of bars curved dorsad, prosophallus--consists

of a pair of long heavily pigmented acuminate arms which have their bases widely

separated and their apices contiguous, base of arm attached to mesal portion of

paramere, located laterad of phallus, prosophallic arm 2. 44 length of phallic

arm, paramere--heavily pigmented, broad, long, approximately 1.8 length of

phallic arm, articulates near middle with caudal area of parameral apodeme,

parameral apodeme--moderately to heavily pigmented, very long, approximately

1.64 length of paramere, outer surface narrowly attached to tergobasal apodeme

of gonocoxite; sternum IX heavily pigmented throughout, large, broad, apico-

lateral margin evenly rounded, 6-7 short and moderately long setae near center

of caudal margin (holotype also with one broad scale).

FEMALE, PUPA, LARVA AND EGG. Not known.

DISCUSSION. Aedes ramalingami belongs to Section B and the adults can

be readily distinguished from the other species of this section except pahangi,

robertsi and uniformis by the dark scaled terga. From the latter 3 species it

is separated by having only 4-5 setae dorsad of scale patch on mesepimeron

while the other 3 species possess 7 or more setae.

The male genitalia of vamalingami differ strikingly from the other species

of Section B in the development of the gonostylus and basal mesal lobe.

This species is dedicated to Dr. Shivaji Ramalingam, Faculty of Medicine,

University of Malaya, Kuala Lumpur, Malaysia, in recognition of his work on

the Malayian Culicidae and the numerous mosquito specimens he has provided

to the SEAMP and MEP collections.

TYPE-DATA. Holotype male, MALAYSIA, Sabah, Beaufort, Kg. Maraba,

0 April 1970, Samuel W. James and Chia Yiew Wang collectors, collection

number S-382, genitalia preparation number T73.555, SEAMP accession num-

ber 300, adult collected resting in bushes of a rubber estate, elevation 76 m

and at 1350 hours; one paratype male, same data as holotype except, collec-

tion number S-383, genitalia preparation number 55467, and collected at 1410

hours. Holotype and paratype deposited in the USNM.

DISTRIBUTION. Known only from the type-locality.

BIOLOGY. As recorded in the type-data.

AEDES (VERRALLINA) RAMI BARRAUD

(Fig. 59)

The following information is based on the holotype male and supplements

the very brief description of Barraud (1928: 370).

FEMALE. Head. Antenna 1.07 length of proboscis, pedicel brown with

a few small brown scales and short fine hairs mesally; maxillary palpus 0.14

length of proboscis; proboscis 1.1 length of femur I; eyes contiguous in front.

Thorax. Scutum covered with narrow curved reddish-brown scales; scutellum

with narrow curved reddish-brown scales on each lobe; dark setae on the fol-

lowing areas: 4 median anterior promontory, numerous acrostichal (anterior

and posterior), numerous dorsocentral (anterior and posterior), scutal fossal

(4-5 anterior, 4 lateral and 1 posterior), numerous supra-alar, 1 postalar

callar and scutellar (4-5 long and 3-4 short ones on lateral lobe, 6 long and 5

Reinert: New Interpretation of Vervallina 81

short ones on median lobe); antepronotum with scales absent, 13-14 brown

setae; postpronotum with narrow curved reddish-brown scales on upper area,

5 dark posterior setae; propleuron with a patch of broad white scales, 15

setae; prosternum, subspiracular area, mesomeron and metameron bare; post-

spiracular area with scales absent, 3 brown setae; mesepisternum with a large

upper and a small lower patch of broad white scales, 4 upper and 10-11 posteri-

or setae, lower ones shorter; mesepimeron with a large patch of broad white

scales near center, 14 setae dorsad of scale patch, several small hairs around

lower posterior corner of scale patch. Legs, Posttarsi I-III (Fig. 59) each

with 2 ungues, I and II each with ungues unequal, larger one with a tooth and

smaller one simple, III with ungues equal, both simple. Wing. Veins brown

scaled; alula with narrow brown scales on margin; 2 remigial setae. Abdomen.

Missing except for genitalia which are mounted on a microscope slide.

DISCUSSION. Aedes rami is apparently a rare species since it is still

represented by only the holotype specimen and no additional specimens have

been collected. It is easily recognized by the male genitalia as figured by

Barraud (1928, Figs. 23, 24).

Tarsomeres 5 and posttarsi are illustrated (Fig. 59) for the first time.

AEDES (VERRALLINA) RARUS DELFINADO

(Figs. 23, 25)

Aedes (Neomacleaya) khmerus Klein 1973, Contrib. Am. Entomol. Inst.

(Ann Arbor) 10(1): 7 (Q*, &*). NEW SYNONYM.

The following information, based on the types of varus and khmerus and the

specimen from Indonesia, supplements the descriptions of Delfinado (1968: 35)

and Klein (1973: 7).

FEMALE. Thovax. Following setae present: 3-4 median anterior prom-

ontory; numerous acrostichal (anterior and posterior); numerous dorsocentral

(anterior and posterior); scutal fossal (5-8 anterior, 4-5 lateral and 1-2 posteri-

or); numerous supra-alar; 4-6 posterior medial scutal; 1 postalar callar; scu-

tellar (4-5 long and 3-5 short ones on lateral lobe, 5 long and 3-5 short ones on

median lobe); 10-15 antepronotal; 4-6 postpronotal; 11-16 propleural; 5-9 post-

spiracular; 4-7 upper and 12-16 posterior mesepisternal, lower ones shorter;

9-11 on prealar knob; and mesepimeron with 12-18 dorsad of scale patch, 2-6

short fine hairs caudad of lower margin of scale patch. Prosternum, sub-

spiracular area, paratergite, mesomeron and metameron bare. Genitalia

(Figs. 23 and 25). As illustrated.

MALE. Head. Antenna 0.98-1.17 length of proboscis; maxillary palpus

0.13 length of proboscis; proboscis 1.22-1.25 length of femur I. Thorax.

Following setae present: 2-3 median anterior promontory; numerous acrostich-

al (anterior and posterior); numerous dorsocentral (anterior and posterior);

scutal fossal (4-5 anterior, 3-5 lateral and 1 posterior); numerous supra-alar;

4-5 posterior medial scutal; 1 postalar callar; scutellar (3-5 long and 1-2 short

ones on lateral lobe, 4 long and 2-3 short ones on median lobe); 7-10 antepro-

notal; 3-6 postpronotal; 9-14 propleural; 2-5 postspiracular; 2-5 upper and 8-

14 posterior mesepisternal, lower ones shorter; 6-9 on prealar knob; and

mesepimeron with 8-12 dorsad of scale patch, 2-4 short fine hairs caudad of

lower margin of scale patch. Prosternum, subspiracular area, paratergite,

mesomeron and metameron bare.

DISCUSSION. I have compared the holotype of varus and the allotype of

82 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

khmerus and find them to be conspecific. The adult habitus of the holotype and

paratypes of kimerus also agree well with those of the holotype of varus and the

allotype of kamerus. Klein (1973: 7) did not examine the holotype of varus and

therefore did not see the male genitalia, which are identical for varus and

kRhmerus.,

This species is apparently infrequently collected, as denoted by the very

few specimens, even though the distribution ranges from Cambodia to Sabah

and to Sumatra.

Female genitalia (Fig. 23) of vavus are very similar to those of incertus

but lack the short hair-like spicules on the basal area of the spermathecal

eminence. They also resemble those of panayensis but have an evenly rounded

shield caudad of the spermathecal eminence while this shield is emarginate in

panayensis.

Male genitalia of varus, incertus and prioekanensis are similar and are

compared in the discussion section of the latter species.

DISTRIBUTION. 11 specimens examined: 89, 3c.

CAMBODIA. Kompong Speu, Sre Klong; 1¢ (holotype of Rkmerus), 52

(paratypes of Ramerus), 22; Kirirom Hills; 1% (allotype of Rhmerus).

MALAYSIA. Sabah (North Borneo), Tawau; 1% (holotype of varus).

INDONESIA. Sumatra, Indragiri; 1.

AEDES (VERRALLINA) SABAHENSIS NEW SPECIES

(Figs. 17, 62, 66)

FEMALE (Fig. 66). Head. Antenna dark brown, 1.06-1.19 length of

proboscis, pedicel dark brown with a few small brown scales and short fine

brown hairs mesally, flagellomere 1 with basal 0.4 pale and with a few small

brown scales; clypeus dark brown, bare; maxillary palpus dark brown scaled,

0. 18-0. 19 length of proboscis; proboscis dark brown scaled, 1.00-1.03 length

of femur I; eyes contiguous in front; several dark ocular setae; vertex cov-

ered with broad blackish-brown decumbent scales except for a few narrow curved

whitish scales along coronal suture, a few similar scales on mesal portion of

ocular line, lateral surface of head covered with broad blackish-brown decum-

bent scales except for a dorsal stripe of broad white scales extending from in

front of antepronotum to eye margin; occiput with numerous dark brown erect

forked scales and several narrow curved and a few broad white decumbent

scales. Thorax. Scutal integument dark brown; scutum covered with narrow

curved reddish-black scales except for narrow curved white scales on median

anterior promontory area, anterior scutal fossal area, supra-alar area above

paratergite and base of wing, and a few along lateral margins of prescutellar

space; scutellum with a patch of narrow curved golden-brown scales on each

lobe; dark reddish-black setae on following areas: 3-4 median anterior prom-

ontory, numerous acrostichal (anterior and posterior), numerous dorsocentral

(anterior and posterior), scutal fossal (3-5 anterior, 4-5 lateral and 2 posterior),

numerous supra-alar, 5-6 posterior medial scutal, 1 postalar callar and scu-

tellar (5-6 long and 3-4 short ones on lateral lobe, 5-6 long and 2-4 short ones

on median lobe); pleural integument dark brown; antepronota widely separated,

scales absent, 12-15 long and short dark setae; postpronotum with narrow

curved reddish-black scales on upper area, 5 posterior dark setae; propleuron

with a patch of broad white scales, 14-21 dark setae; prosternum, subspiracu-

lar area, paratergite, mesomeron and metameron bare; postspiracular area

without scales, 3-7 dark setae; mesepisternum with a large upper and a small

Reinert: New Interpretation of Vervallina 83

lower patch of broad white scales (several scales in upper patch with a brown

hue), 4-5 upper and 8-10 posterior dark setae, lower ones shorter; prealar

knob without scales, 7-9 dark setae; mesepimeron with a patch of broad white

scales near center, 12-15 dark setae dorsad of scale patch, 3-5 small golden

hairs near lower posterior edge of scale patch. Legs. Coxae I-III each with

several brown setae, I with broad dark brown scales on anterior surface and a

patch of broad white scales dorsally and ventrally of brown ones, II and III each

with a small patch of broad white scales on anterior surface; trochanters I-III

each with a few short golden setae and a few brown and white scales; femora I

and II each with anterior surface dark brown scaled, I with posterior surface

brown scaled with a ventral longitudinal white scaled stripe from base to apex,

II and III each with posterior surface and also anterior surface of III white

scaled with a narrow dorsal longitudinal brown scaled stripe from base to apex,

stripe forming a narrow brown scaled band at apex; tibiae I-III and tarsi I-III

each dark brown scaled; posttarsi I-III (Fig. 62) each with 2 ungues, I and II

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

Dorsal and ventral veins with dark brown scales; alula with several moderately

broad and a few narrow brown scales on margin; upper calypter with a number

of brown setae on margin; 1-3 remigial setae. Halter. Pedicel pale; capitel-

lum brown scaled. Abdomen. Terga each blackish-brown scaled with a latero-

basal white scaled patch; sterna II-VII each brown scaled with a narrow basal

band of creamy-white scales which becomes broader laterally; terga and sterna

with numerous golden setae, mostly along posterior margins. Genitalia (Fig.

17). Tergum VIII moderately pigmented, wide, base slightly concave, apex

broadly convex, several long stout setae along apical margin and several mod-

erately long ones on apical 0.33-0.36, basolateral seta present, covered with

minute spicules, numerous broad scales covering apical 0. 82-0. 84, VIII-Te

index 0.43-0.44, VIII-Te/IX-Te index 5. 53-6. 82; sternum VIII moderately

pigmented, wide, base concave mesally, apex with a shallow median indentation

with a small lobe on each side of midline, numerous moderately long setae

apically, numerous short and a few long setae scattered over apical 0. 87-0. 92,

covered with minute spicules, numerous broad scales covering most of apical

0.82-0.85, apical intersegmental fold unpigmented, VIII-S index 0. 58-0. 64;

tergum IX moderately pigmented, very wide and short, ribbon-like, setae

absent, covered with minute spicules, [X-Te index 0.12-0.16; insula unpig-

mented, short, covered with minute spicules, 4 small tuberculi, each with a

tiny spicule; lower vaginal lip heavily pigmented, narrow, a few minute spicules

on outer part of hinge area, with a small caudally projecting heavily pigmented

median apodeme, lower vaginal sclerite composed of a pairof long slender

heavily pigmented plates which are connected basally, with minute spicules

(more numerous and longer on apical area), apical portion slightly expanded

and slightly cupped; upper vaginal lip heavily pigmented, posterolateral corn-

ers Slightly produced, median posterior area produced into a shield caudad of

spermathecal eminence, upper vaginal sclerite heavily pigmented, large and

complex, base attached along entire lateral inner margin of upper vaginal lip

and covering most of vaginal roof; spermathecal eminence heavily pigmented,

large, deep in depth, triangular in dorsal outline, short fine branched spiny

excrescences attached to most of dorsal area; postgenital lobe short, apex

with a moderately deep median indentation, 14-24 setae on each side of mid-

line, completely covered with minute spicules, dorsal PGL index 0. 64-0. 69,

ventral PGL index 0. 81-0.93; peri-anal membrane with minute spicules; cer-

cus triangular in shape, moderately long, broad at base, apex acute, com-

pletely covered with minute spicules, dorsal surface with a number of moder-

84 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

ately long setae over most of apical 0.75-0.78, ventral surface with a number

of short and moderately long setae along outer area, a few long setae at apex,

scales absent, cercus index 2.44-2.63, cercus/dorsal PGL index 3. 18-3. 33; 3

seminal capsules, 1 large, 1 medium and 1 small in size, heavily pigmented,

elliptical, each with a neck and with several small seminal capsule pores

near orifice, base of accessory gland duct unpigmented.

MALE, PUPA, LARVA AND EGG. Not known.

DISCUSSION. Aedes sabahensis is similar in adult habitus to cauius and

pseudomediofasciatus; however, it can be separated from these 2 species by the

abdominal markings and the number of setae on the propleuron. Features of

the female genitalia readily distinguish each of the 3 species. The female

genitalia of sabahensis are also easily separated from all other species of

the subgenus in the development of the lower vaginal sclerites which are repre-

sented by a pair of long narrow spiculate plates, joined basally and slightly ex-

panded apically.

Delfinado (1967: 13) mistakenly described and illustrated the female of

sabahensis as cautus. The adults of these 2 species, as mentioned above,

are similar in adult habitus but possess markedly different genitalia.

TYPE-DATA. Holotype female, MALAYSIA, Sabah, Beaufort, Kg.

Bingkul, 1 April 1970, Sulaiman bin Omar and Chia Yiew Wang collectors,

collection number S-318, genitalia preparation number T73.487, SEAMP

accession number 300, adult collected biting man at 1900-2000 hours in a vil-

lage with numerous nipa palms and at an elevation of 30 m; one paratype fe-

male, same data as holotype except genitalia preparation number 54374; one

paratype female, MALAYSIA, Sabah, Kengingau, 16 August 1956, collection

number US-79, genitalia preparation number T73.389, SEAMP accession num-

ber 24, collected from a horse-baited trap; one paratype female, MALAYSIA,

Sabah, Papar, 26 July 1965, F. Y. Cheng collector, collection number 101,

genitalia preparation number 10-III-66 (identified as cautus by Delfinado); and

one paratype female, MALAYSIA, Sabah, Forest camp 9.8 km southwest of

Tenom, 18 December 1962, Y. Hirashima collector, genitalia preparation

number T73.430. The holotype and 3 paratypes are deposited in the USNM,

and one paratype (T73. 389) will be deposited in the BMNH.

DISTRIBUTION. Known only from localities mentioned in the type-data.

BIOLOGY. Females were taken biting man (bare leg catch) in a village

with numerous nipa palms, at 30 m elevation, and at 1900-2000 hours. One

female was collected in a horse-baited trap.

AEDES (VERRALLINA) SECULATUS MENON

DISCUSSION. I have examined the male and female cotypes of Aedes

carteri and agree with Stone's (1956: 339) synonymy of this species with

seculatus; however, only the males are conspecific. The female cotype and

an additional female of carteri are synonymous with butleri. The adult female

cotype of carteri is very badly damaged (only part of the head and part of the

thorax remain on the pin and no genitalia mount could be found) and could not

be identified; however, a female genitalia, No. 187 (assumed by me to be

the female from Denipitiya mentioned by Wijesundara (1951: 176)), mounted in

Canada balsam on a slide, and labeled as a paratype of cavteri can be deter-

mined as butleri. Figure 5 of Wijesundara (1951) appears to have been drawn

from the genitalia slide No. 187. Since the adult fragment of the female cotype

does not possess any remaining characters that can be used to identify it to

Reinert: New Interpretation of Verrallina 85

species I assume it to be conspecific with the Denipitiya female, the genitalia

of which are identical to those of the lectotype of butleri. The adult male co-

type of cartervi is also heavily damaged (only a thoracic fragment remains on

the pin) but the male genitalia, mounted in Canada balsam on a slide, clearly

indicate this species is identical to the type male of seculatus, which I have

also examined. The male cotype of cartevi in the BMNH is hereby selected

lectotype and bears the following data on the adult and genitalia slide mount

labels: type &, Hypo. No. 102, Matugama R. H.| Rest House], 2.9.21, Rain-

water puddle in dense shade, pupa & larva skins, Med. Entomologist, Ceylon.

The female cotype of cartevi possesses the same data on the labels as the co-

type male except the following: 29.8.21. and Hypo. No. 151 and the remaining

female genitalia, mounted on a slide, contain the following information--para-

type, 187, Aédes (Aédes) carteri Wijesundara, are transferred to synonymy

with butleri.

AEDES (VERRALLINA) SOHNI NEW SPECIES

(Figs. 33, 56)

MALE. Head. Antenna plumose, brown with pale areas, 0. 85-0. 99

length of proboscis, pedicel dark brown, flagellomere 1 with a few small

brown scales near middle; clypeus dark brown, bare; maxillary palpus dark

brown scaled, 4 segmented, 0.12 length of proboscis; proboscis dark brown

scaled, 1.13 length of femur I; eyes contiguous; ocular setae blackish-brown

and well developed; scales on head all broad, blackish-brown and decumbent

except for a few narrow curved white scales along coronal suture and upper

interocular space, a stripe of broad white scales extending from area in front

of antepronotum to ocular line, and a patch of dark brown erect forked scales

on occiput. Thorax. Scutal integument dark reddish-brown; scutum covered

with narrow curved reddish-brown scales except for narrow curved white

scales on the following: small patch on median anterior promontory area,

small patch on anterior scutal fossal area and extending caudally onto

lateral scutal fossal area, a few along scutal ridge and on acrostichal area,

patch on supra-alar area and a few on posterior medial scutal area; scutellum

with a patch of narrow curved pale brown scales on each lobe; prescutellar

space bare; reddish-black setae on the following areas: 6-10 median anterior

promontory, numerous acrostichal (anterior and posterior), numerous dorso-

central (anterior and posterior), scutal fossal (7-9 anterior, 4-5 lateral and

1-2 posterior), numerous supra-alar, 6 posterior medial scutal, 1 postalar

callar and scutellar (4-5 long and 4-8 short ones on lateral lobe, 4 long and

6-8 short ones on median lobe); pleural integument brown; antepronota widely

separated, a few broad white scales, 9-13 long and short dark setae; postpro-

notum with a few narrow curved reddish-brown scales on upper area, 4-6 dark

posterior setae, a few short dark hairs cephalad of setae; propleuron with a

patch of broad white scales, 16-18 setae; prosternum with a patch of short

brown setae on the upper mesal area; subspiracular area, paratergite and

mesomeron bare; postspiracular area with a small patch of broad white scales

caudad of setae, 2-4 brown setae; mesepisternum with a large upper and a

small lower patch of broad white scales, 3-5 upper and 12-14 posterior brown

setae, a few short fine pale golden hairs dorsad and ventrad of lower scale

patch; prealar knob without scales, 9 brown setae; mesepimeron with a patch

of broad white scales near center, 10-11 setae dorsad of scale patch, several

short fine golden hairs caudad of scale patch; metameron with several small

86 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

broad white scales and several short fine pale hairs. Legs. Coxae I-III each

with several brown setae, I with broad brown scales and a small dorsal and a

ventral patch of broad white scales on anterior surface, II with a patch of broad

white scales on anterior surface, III with a ventral and a dorsal patch of broad

white scales on anterior surface; trochanters I-III each with broad pale scales;

femora I and II each with anterior surface brown scaled, posterior surface with

a longitudinal white scaled stripe from base to near apex, stripe dorsal on I

and ventral on II, III brown scaled with a ventral longitudinal white stripe from

base to near apex on both anterior and posterior surfaces; tibiae I-III and tarsi

I-III each dark brown scaled; posttarsi I-III (Fig. 56) each with 2 ungues, I and

Il each with ungues unequal, each with a tooth, II with ungues equal, both

simple. Wing. Dorsal and ventral veins with blackish-brown scales; alula

with several narrow brown scales on margin; upper calypter with several

brown setae on margin; 1 remigial seta. Halter. Pedicel pale; capitellum

dark brown scaled. Abdomen. Terga dark brown scaled, I-VI each with a

small laterobasal patch of white scales (remainder of abdomen removed with

eenitalia); sterna white scaled with a few pale brown scales along posterior

margins of several sterna; tergaand sterna with several brown setae, most

numerous along posterior margins. Genitalia (Fig. 33). Tergum IX heavily

pigmented, band-like with lateral areas broader than median area, outer caudal

margin fused to tergum X, setae absent, broadly connected laterally to sternum

IX; gonocoxite short, broad, heavily pigmented, mesal surface membranous

with minute spicules, dorsal surface with apex projected into a long moderately

broad tapering flattened lobe with apical area bifid, 4-5 long stout setae along

outer proximal area, a few short and long setae on apical area of dorsal sur-

face of gonocoxite, ventral surface projected into a broad flattened mesal area

with a rough apical lobe, numerous short stout setae on flattened area, several

long stout setae on apical 0.65 of remainder of gonocoxite, ventral surface also

with an outer apical short rounded arm which bears 2-3 long heavily pigmented

flattened pointed spines, lateral surface with a few long setae, broad scales on

outer area of dorsal surface and on lateral and ventral surfaces; gonostylus

long (approximately 1.12 length of gonocoxite minus apical lobe), narrow,

curved, base with a small lobe bearing 2 moderately long setae, apex slightly

recurved, attached subapically on gonocoxite; basal mesal lobe moderately

pigmented, consists of a basal flattened area bearing 2-3 moderately long

heavily pigmented spiniforms, 2-3 short lightly pigmented spiniforms and num-

erous short thin setae, a narrow distal extention on gonocoxite along mesal

margin to apex which bears several moderately long thin setae and terminates

in a long slender rounded arm with a short stout subapical spine (holotype with

spine asymmetrical, one side as above and other side with a long stout setae

attached near base of arm instead of a spine), lobe connected mesally with its

mate by a narrow moderately pigmented band covered with minute spicules

and located ventrad of apical portion of prosophallus and phallus; proctiger

with paraproct formed into a heavily pigmented long slender curved bluntly

pointed arm with basal area slightly expanded and connected to tergum IX,

base with a short projection which extends ventrad and articulates at a point

on the dorsomesal surface of the parameral apodeme near base of opistho-

phallus, cercus membranous and without cercal setae; tergum X moderately

to heavily pigmented, indistinct from tergum IX; phallosome complex, opistho-

phallus--consists of a moderately pigmented tergal transverse bridge between

parameral apodemes, caudal margin nearly straight, sternobasal area attached

to tergobasal portion of prosophallus, located dorsad of phallus and prosophal-

lus and ventrad of proctiger, phallus--consists of a pair of moderately long,

Reinert: New Interpretation of Vervallina 87

heavily pigmented, narrowly separated, caudally produced arms which are

fused together by a narrow basal plate which extends laterally and is fused to

basal portion of prosophallus, apex of phallus formed into a tergal penis fila-

ment which consists of a small lightly pigmented lobe produced cephalad,

prosophalius--consists of a pair of moderately long heavily pigmented widely

separated arms, apex blunt, basal portion somewhat expanded and attached to

mesal portion of paramere, located laterad and slightly ventrad of phallus,

prosophallic arm approximately equal in length to phallic arm, paramere--

heavily pigmented, approximately equal in length to phallic arm, articulates

at a point approximately 0.65 from base with caudal area of parameral apodeme,

parameral apodeme--moderately to heavily pigmented, moderately long, outer

surface broadly attached to tergobasal apodeme of gonocoxite; sternum IX

heavily pigmented throughout, large, broad, lateroapical areas rounded, 25-31

short and moderately long setae in an elongate patch along caudal margin, 1

broad scale on holotype.

FEMALE, PUPA, LARVA AND EGG. Not known.

DISCUSSION. Aedes sohni belongs to the andamanensis Group of species.

The adult habitus is very similar to the other members of the group and can

not be: separated from them with certainty (except johnsoni which lacks setae

on the prosternum). The male genitalia, however, are easily distinguished

from the other species by the development of the basal mesal lobe, gonocoxite

with sternoapical portion expanded, thickening of the paraproct and the develop-

ment of the gonostylus. See discussion section of adustus for additional infor-

mation.

This species is dedicated to Young T. Sohn, scientific illustrator for the

Medical Entomology Project, for preparing the many fine detailed illustrations

included in this and other articles.

TYPE-DATA. Holotype male (genitalia mounted on a slide in Canada bal-

sam) with the following collection data: MALAYSIA, Sabah, Sapulut, Pensiangan,

11 April 1970, collection number S-481, Sulaiman ben Omar and Chia Yiew

Wang collectors, SEAMP accession number 300, genitalia preparation number

74/459, collected as an adult on a clear day, ina secondary rain forest, ina

partially shaded area by the side of a river, at 1225 hours, and at an elevation

of 168 m; and one paratype male with the following collection data: THAILAND,

Narathiwat, Amphoe Waeng, in the jungle 1 km from Soi Teo 5 in Naeng Set-

tlement, 22 February 1965, collection number NV76-12, Sumeth Chunchuncherm

collector, genitalia preparation number 74/487, collected as a pupa from a

small puddle (2 feet by 2 feet) containing slightly turbid water in a moderately

shaded area. Holotype and paratype deposited in the USNM and are in excellent

condition except 3 left legs missing from the holotype and the pupal skin of the

paratype is missing.

DISTRIBUTION. Known only from localities mentioned in the type-data.

BIOLOGY. As recorded in the type-data.

AEDES (VERRALLINA) TOROSUS DELFINADO

(Figs. 45, 48, 57)

PUPA (Fig. 45). Chaetotaxy as figured and recorded in Table 12. Des-

cription based on 2 pupal skins with associated male adults. Ocular plate with

a small area of moderately developed cuticular facets. Respiratory trumpet.

Moderately to heavily pigmented, hair-like spicules on distal 0. 76-0. 87 of

inner surface; index 3.31-3.39. Abdomen. Setae 6, 9-VII weakly developed;

88 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

4-VIII with 4-5 branches; 9-VIII single. Paddle. Ovoid; minute serrations on

distal portion of basal 0.63-0.71 of outer margin, more strongly developed on

distal area; minute spicules on apical 0. 30-0. 38 of outer and apical 0.10-0. 13

of inner margins; midrib does not reach apex; seta 1-P moderately long, single;

index 1. 56.

LARVA (Fig. 48). Chaetotaxy as figured and recorded in Table 24. Des-

cription based on 2 larval skins with associated adults. Head. Seta 4-C with

0-6 branches; 9-C double or triple; 6-C single; 7-C with 7-8 branches; 12-C

with 9-6 branches; dorsomentum with 37-38 teeth. Antenna. Moderately long,

moderately pigmented, with a number of scattered small spicules; seta 1-A

triple to 4 branched, moderately long; 2-A long, with a subapical constriction;

3-A short, 0.17-0.23 length of 2-A; 4-A moderately long, 0. 44-0. 57 length of

2-A; 6-A short, 0.28-0.32 length of 2-A. Thorax. Seta 6-T triple to 5

branched. Abdomen. Seta 1-VII 4 branched; 3, 5-VIII with 7-9 branches; 2-X

with 6-9 branches; ventral brush with 8-10 setae on grid and 2 precratal shorter

ones, all multiple branched; saddle moderately pigmented, incompletely rings

segment X, acus absent; 4 moderately long anal papillae; comb with 11-13

(usually 12) scales arranged in a single curved irregular row, each scale with

a stout median apical spine and small denticles on laterobasal areas. Siphon.

Moderately pigmented; acus well developed; index 2.13-2.31, average 2. 22;

pecten on basal 0. 59-0. 60 of siphon, composed of 14-15 (usually 14) teeth,

apical 2 teeth longer and wider spaced than remainder of teeth, each tooth long,

slender and with 1-2 small stout ventral denticles near base; seta 1-S short,

4-6 branched, base attached on basal 0. 69-0. 71 of siphon distad of last pecten

tooth.

DISCUSSION. The present descriptions and illustrations are the first for the

pupa and fourth stage larva and are based on the specimens from Prachin Buri

Province, Thailand.

Tarsomeres 5 and posttarsi of the male are illustrated (Fig. 57) for the

first time.

DISTRIBUTION. 8 specimens examined: 40", 2 with associated pupal and

larval skins.

THAILAND. Khon Kaen, Tham Pho Ti Yan; 2° (holotype and paratype,

USNM type No. 69216). Pvachin Buri, Ban Tub Lan; 20 pl.

BIOLOGY. The above larvae were collected from a small seepage pool

containing clear, fresh, unmoving, temporary water, in partial shade, located

in a secondary scrub area in a valley and at an elevation of 80 m.

AEDES (VERRALLINA) UNCUS (THEOBALD)

(Pies: 23, 55.57, 62)

Aedes (Aedes) hirsutipleura Barraud 1928, Indian J. Med. Res. 16: 370 (c*,

2*). NEW SYNONYM.

LARVA (Fig. 55). Chaetotaxy as figured and recorded in Table 25. Head.

Seta 4-C with 5-7 branches; 5-C double or triple; 6-C single; 7-C with 5-10

branches; 12-C with 7-10 branches; dorsomentum with 38-42 teeth. Antenna.

Moderately long, lightly pigmented, with several small scattered spicules;

seta 1-A triple to 6 branched, moderately long; 2-A long, with a subapical

constriction; 3-A short, 0.20-0.27 length of 2-A; 4-A moderately long, 0. 37-

0.53 length of 2-A; 6-A short, 0.26-0.30 length of 2-A. Abdomen. Seta 1-VIII

triple to 6 branched; 3-VIII with 6-11 branches; 5- VIII with 6-10 branches; 2-X

Reinert: New Interpretation of Vervallina 89

with 5-8 branches; ventral brush with 12 setae on grid and 2 precratal shorter

ones, all multiple branched; saddle moderately pigmented, incompletely rings

segment X, acus absent; 4 moderately long anal papillae; comb with 10-12 (usu-

ally 12) scales arranged in a single curved row, each scale with a stout medi-

an apical spine and small denticles on laterobasal areas. Siphon. Moderately

pigmented; acus well developed; index 1.77-2.15, average 1.95; pecten on |

basal 0. 54-0. 61 of siphon, composed of 13-16 (usually 15) teeth, apical 2 teeth

longer and wider spaced than remainder of teeth, each tooth long, slender and

with 1-2 small stout ventral denticles near base; seta 1-S short, 4-8 branched,

base attached on basal 0. 63-0.68 of siphon distad of last pecten tooth.

DISCUSSION. A single male (with genitalia preparation T73.255) in the

USNM from Mindanao reestablishes the range of wcus on this island. Materi-

al forming the previous record from Mount Apo, Mindanao (Bohart 1945: 66)

served as the type material of nubicolus, a new species described by Laffoon

(1946: 237).

Delfinado's record (1967: 32) of uncus larvae collected from axils of the

elephant ear plant (Colocasia) is a doubtful biological record. There are 2

male and 2 female specimens of wncus in the USNM from Palawan, P.I., and

each bears 2 labels. The upper label contains the following data--P 335,

Aédes uncus Theobald; data on the lower label are as follows--P355, 19MGL,

Elephant ear axil, Palawan, 2-X-45. Since the collection numbers (P335,

P355) on the 2 labels are different, it appears that there was a mix-up in at-

taching labels to the specimens and the lower labels do not belong to the

specimens of uncus, In addition, this would be an unusual habitat for the im-

matures of wncus, as well as other members of the subgenus. Other collec-

tions of uncus by the 19th Medical General Laboratory (19MGL) in the same

area and near the date were adult resting and biting collections and these

specimens are in a badly rubbed condition similar to those of the P335 collec-

tion. If the P335 specimens were reared from immatures they probably

would have been in better condition. Immature skins of wncus from the P335 or

other collections made by the 19MGL from Palawan are not in the USNM.

Borel's (1930: 291) description and figure of Aedes sp. ? no. 51 compare

very well with those of Aedes uncus and I am confident they belong to this

species.

The present description and illustration are the first of the fourth stage

larva of this species and are based on several whole larvae and reared speci-

mens with associated pupal skins and adults from Malaysia and several whole

larvae from the Philippines.

Barraud (1928: 370) originally described Aedes (Aedes) hirsutipleura from

one female and one male cotypes and 5 other females and 3 other males. The

2 cotypes and one female and one male (labeled as paratypes) are deposited

in the BMNH. The male cotype is hereby designated lectotype and bears the

following label data: Type, Golaghat, Assam, Jan. 1925, P. J. Barraud,

No. 2085, B. M. 1935-622., o genit. YMH-66-BM-3 | YMH = Yiau-Min

Huang], SEAMP Accession Number 374; allolectotype is designated for the

female cotype and bears the following data on the adult labels: same as lecto-

type except No. 2086 and ? genit. YMH-66-BM-6. Condition of the lectotype

follows: thorax split on right side, right wing missing, 3 legs on right side

broken off, 2 of these glued to cork stage, and genitalia mounted in Canada

balsam on a microscope slide.

Aedes uncus belongs to a group of closely related species (cyrtolabis,

harrisonicus, latipennis, leicesteri, macrodixoa, neomacrodixoa, nubicolus,

protuberans, singularis, torosus and virilis) all of which are extremely similar

90 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

in adult habitus and can not be separated from each other with certainty by

habitus features. The species of this group, however, are easily distinguish-

able by characters of both the male and the female genitalia. Members of

this species group share the following combination of characters: mesepistern-

um with several fine hairs on anterodorsal area; mesepimeron with numerous

fine hairs caudad of scale patch and extending to near lower margin of scler-

ite; male genitalia with both prosophallus and phallus very long, approximately

equal in length, the later very slender; and female genitalia with lower vaginal

sclerite large, usually with an apical median notch, spermathecal eminence

with a characteristic shape and the inner margin heart-shaped.

Tarsomeres 5 and posttarsi of the male (Fig. 57) and the female (Fig. 62)

and tergum IX of the female genitalia (Fig. 23) are illustrated for the first

time.

DISTRIBUTION. 321 specimens examined: 1462, 690, 4p, 30 L, 47 with

associated skins (22 p, 251).

CAMBODIA. Kompong Sela, Stung Chral; 92. Kompong Speu, Kirirom;

12;

INDONESIA. Java, Bali Bengkula; 12. Sumatra, Atchin, Kotta Tjane;

Dermajoe (Benkoelen); 32, 9.

MALAYSIA. Negri Sembilan, Bahau; 19? pl, 31°, 4c pl, 260%, 24 L.

Perlis, Kaki Bukit, Kubang Tiga; 12 pl, 1% pl. Sabah, Beaufort, Kg. Bingkul,

Kg. Maraba, Kuala Papar, Tenom, Tuaran; 212, 7%. Selangor, Kuala Lum-

pur, Templer Park; 2°, 1 L.

PHILIPPINES. Balabac Island, Cape Melville; 22, 1°. Culion Island;

1%. Mindanao Island, along Davao highway near Cotabato; 1*. Palawan

Island, Irahuan River Valley, Puerto Princesa; 169, 100, 5L.

SOUTH VIETNAM. Khanh Hoa, My Ca; 19.

THAILAND. Chiang Mai, Chiang Mai; 12, 1%. Chon Buri, Khao Mai Keo;

22. Khon Kaen, Tham Pho Ti Yan; 10. Nakhon Nayok, Amphoe Muang, Khao

Yai National Park, Sariga; 22. Nakhon Ratchasima, Khlong Phai, Amphoe

Pak Chong, Lam Nam Tha; 52. Songkhla, Amphoe Haad Yai, near Ton Nga

Chang Waterfalls; 162 p, 22, 6 p, 4p. Nakhon Si Thammarat, Banna Nabon;

12. Tvang, Amphoe Muang, Ban Kon Tn Thaningarm; 1c.

BIOLOGY. In Thailand immatures were collected from large stream pools

in a dry stream bed (filled with rain water) and a seepage pool; from colored

or turbid, fresh, unmoving, temporary water in moderately to heavily shaded

areas located in secondary rain forests and bamboo groves; and at an elevation

of 500 m. Adults were collected biting man in a forest during a rain shower

and resting in heavy forest vegetation.

Immatures in Malaysia were collected from small flood and ground pools

and a small swampy area; from clear, colored or turbid, fresh, unmoving,

temporary water in moderately to heavily shaded areas; located in rubber

plantations and secondary rain forests; and at elevations from sea level to

76m. Adults were taken biting man between 1900-2000 hours in a nipa palm

village, in a secondary rain forest at 1010 hours and in heavy shade in scrub

area of a rubber plantation between 1300-1440 hours.

Adults in the Philippines were collected biting man at 0900 hours ina

coconut-bamboo grove and biting man during the morning in a jungle.

One female was collected in a light trap in South Vietnam. The light trap

was situated in a beach environment near My Ca.

Reinert: New Interpretation of Verrallina 91

AEDES (VERRALLINA) VALLISTRIS BARRAUD

(Figs. 23, 46, 52, 57, 62)

PUPA (Fig. 46). The following description and Table 13 are the first

description of the pupa of this species. Respivatory trumpet. Moderately

pigmented; index 3.41-4.46, average 4.05. Abdomen. Setae 6, 9-VII weakly

developed; 4-VIII double; 9-VIII double or triple. Paddle. Ovoid; minute ser-

rations on distal portion of basal 0.6 of outer margin; minute spicules on apical

0.4 of outer and apical 0.21 of inner margins; midrib does not reach apex;

seta 1-P moderately long, single; index 1.31-1.44, average 1.38.

LARVA (Fig. 52). The following description and Table 26 are the first

description of the larva of vallistris and are taken from 2 damaged skins with

associated adults. Head, Seta 4-C with 9 branches; 5, 6-C triple; 7-C with

8-13 branches; 12-C with 7 branches; dorsomentum with 36-39 teeth. Antenna,

Moderately long, lightly pigmented, with several scattered small spicules,

most numerous on basal area; seta 1-A triple to 4 branched, moderately long;

2-A long, with a subapical constriction; 3-A short, 0.24-0.25 length of 2-A;

4-A moderately long, 0.56 length of 2-A; 6-A short, 0.24-0.25 length of 2-A.

Abdomen. Setae 1, 3-VIII with 6-8 branches; 5-VIII with 8-13 branches; 2-X

with 7-9 branches; ventral brush with 10 setae on grid and 2 precratal shorter

ones, all multiple branched; saddle moderately pigmented, incompletely rings

segment X, acus absent; 4 moderately long anal papillae; comb with 10 scales

arranged in a Single irregular row, each scale with a stout median apical spine

and small denticles on laterobasal areas. Siphon. Moderately pigmented;

acus well developed; index 1. 89-2.07; pecten on basal 0. 61-0. 62 of siphon,

composed of 9-11 (usually 11) teeth, apical 2-3 teeth longer and wider spaced

than remainder of teeth, each tooth long, slender and with 1-2 small stout

ventral denticles near base; seta 1-S short, double or triple, base attached on

basal 0. 72-0. 73 of siphon distad of last pecten tooth.

DISCUSSION. Many of the specimens reported as vallistris from Thailand

by Delfinado (1967: 33) were incorrectly identified. These records of this

species from Thailand are in error and were based on the following specimens

(correct identifications are in parentheses): Chiang Mai (M694-219), atrius);

Nakhon Nayok (NY-60°2, uncus; NY-141°, cretatus; no male found); Nakhon

Ratchasima (T-93389, uncus); Nakhon Si Thammarat (T-15769, wncus; T-16029,

notabilis); Ubol (T-8802, unidentifiable fragment on pin, no genitalia prepara-

tion); and Udorn (T-5292°, butleri). After examining the above list it is found

the only valid record of vallistris cited by Delfinado (1967: 33) remaining for

Thailand is from Chiang Mai Province. Aedes vallistris is similar in the adult

habitus to several species in the subgenus and positive identification should be

made from genitalia preparations. All of the specimens in the above list did not

have the genitalia prepared prior to the present study which could partially ex-

plain the previous misidentifications. From the illustration and discussion of

vallistris by Iyengar and Menon (1956: 791) this species is also reported from

Yala and Pattani in southern Thailand.

Tergum IX of the female genitalia is illustrated (Fig. 23) for the first time.

This structure occasionally has a single short seta on lateroapical margin of

one side. The posttarsi of the male (Fig. 57) and the female (Fig. 62) are il-

lustrated for the first time.

DISTRIBUTION. 37 specimens examined: 299, 40°, 2 with associated pupal

and larval skins.

CAMBODIA. Kandal, Phnom Penh, Arey Ksatr, Chrui Chang War; 89.

THAILAND. Chiang Mai, Ban San Khayom, Huey Chang Kien; 22 pl, 192, 4°.

92 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

BIOLOGY. Immatures in Thailand were collected from a large well con-

taining clear, fresh, unmoving, temporary water in a partially shaded area

located in a village and at an elevation of 300 m. Adults in Thailand were taken

biting man in bamboo forests and resting in a bamboo forest located at the mar-

gin of a rice field.

AEDES (VERRALLINA) VARIETAS (LEICESTER)

(Figs. 23, 62)

DISCUSSION. Delfinado (1968: 40) reported the male genitalia of the lecto-

type of vavrietas as missing from the celluloid mount on the adult pin. After

careful examination of the celluloid mount, which was strongly curved upward

and very dark, I was able to locate the genitalia. The genitalia were in poor

condition, but recognizable, and were remounted in Canada balsam on a micro-

scope slide bearing the designation T73.295. Delfinado also stated the lecto-

type label read "Jungle Pahang Rd 5th mile." It actually reads--Jungle Pahang

Rd 4th mile. For a discussion on the lectotype designation see Knight and Hull

(1953: 474) and Delfinado (1968: 40). .

The female posttarsi (Fig. 62) and tergum IX (Fig. 23) of the female geni-

talia are illustrated for the first time.

The females listed below compare well with the lectotype of varielas and

the description and illustration of Delfinado (1968) and are therefore provisional-

ly assigned to this species.

DISTRIBUTION. 26 specimens examined: 239, 3c.

INDONESIA. Sumatra, Atchin District, Kotta Tjane, Benkoelen District,

Dramajoe; 89, 2c.

MALAYSIA. Selangor, Kuala Lumpur, Segambut; 129, 1 (lectotype).

Trengganu, Kuala Brang, Kg. Payang; 12, Kg. Tapah; 19.

SINGAPORE. West coast road 8.5 miles; 19.

BIOLOGY. Adult females from Malaysia were taken biting man and the

female from Singapore was collected at 1900 hours in a resting collection near

sea level.

AEDES (VERRALLINA) VIRILIS (LEICESTER)

(Figs. 18, 34, 57, 61)

FEMALE. Head, Antenna dark brown, 0.91-0.95 length of proboscis,

pedicel dark brown basally and yellow apically, with a few small brown scales

and short fine brown hairs mesally, flagellomere 1 with basal 0.395 pale and

with a few small brown scales; clypeus dark brown, bare; maxillary palpus

dark brown scaled, 0.15 length of proboscis; proboscis dark brown scaled,

1.19-1.21 length of femur I; eyes contiguous in front; several dark ocular

setae; vertex covered with broad blackish-brown decumbent scales except for

narrow curved golden-white scales along posterior 0.5 of coronal suture, a

few similar scales on ocular line, lateral surface of head covered with broad

blackish-brown decumbent scales except for a dorsal stripe of broad white

scales extending from in front of antepronotum to eye margin; occiput with

numerous brown erect forked scales and narrow curved golden-white decum-

bent scales, a few scattered broad decumbent white scales. Thovax. Scutal

integument brown; scutum covered with narrow curved reddish-black scales

except for narrow curved golden-white scales forming a small patch on median

Reinert: New Interpretation of Verrallina 93

anterior promontory and anterior scutal fossal areas, a few similar scales on

lateral scutal fossal area; scutellum with a patch of narrow curved reddish-

black scales on each lobe; prescutellar space bare except for a median cephalic

extention of narrow curved reddish-black scales; dark reddish-black setae on

following areas: 4-5 median anterior promontory, numerous acrostichal

(anterior and posterior), numerous dorsocentral (anterior and posterior),

scutal fossal (4-7 anterior, 5-7 lateral and 1-2 posterior), numerous supra-

alar, 6 posterior medial scutal, 1 postalar callar and scutellar (5-6 long and

4-6 short ones on lateral lobe, 6 long and 5-6 short ones on median lobe);

pleural integument brown; antepronota widely separated, scales absent, 16-21

long and short dark setae; postpronotum with narrow curved reddish-black scales

on upper area, 6-7 posterior dark setae, a few short fine dark hairs cephalad of

setae; propleuron with a patch of broad white scales, 17-20 setae; prosternum,

subspiracular area, paratergite, mesomeron and metameron bare; postspiracu-

lar area with 9-11 dark setae, scales absent; mesepisternum with a large upper

patch of broad brown scales and a small lower patch of broad white scales,

several short fine golden hairs cephalad of and caudad of upper scale patch, 7-8

upper and 15-16 posterior brown setae; prealar knob with 17-19 dark setae,

scales absent; mesepimeron with a patch of broad white scales near center, 26-

29 light brown setae dorsad of scale patch, numerous small golden hairs caudad

of scale patch and extending from top of patch to near lower margin of sclerite.

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

scales on anterior surface and a small patch of broad white scales dorsally, II

and III each with a patch of broad white scales on anterior surface; trochanters

I-III each with several white scales; femur I with anterior surface brown

scaled and posterior surface white scaled with a longitudinal narrow brown

stripe from base to apex, II with anterior and posterior surfaces brown scaled

and with a longitudinal white scaled stripe from base to near apex, stripe nar-

row anteriorly and broad posteriorly, III with anterior and posterior surfaces

white scaled and with a dorsal longitudinal narrow brown stripe from base to

apex of anterior and posterior surfaces, stripe forming a narrow brown band

at apex; tibiae I-III each brown scaled, I and II each with a posterior longitu-

dinal white scaled stripe from base to apex, stripe posteroventral on J; tarsi

I-III brown scaled; posttarsi I-III (Fig. 61) each with 2 ungues, I and II with

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

Dorsal and ventral veins with dark brown scales; alula with several moderately

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

margin; 1 remigial seta. Halter. Pedicel pale; capitellum brown scaled with

white scales at apex. Abdomen. Terga dark brown scaled, each with a latero-

basal white scaled patch; sterna white scaled with brown scaled apical bands;

terga and sterna with numerous golden-brown setae, mostly along posterior

margins. Genitalia (Fig. 18). Tergum VIII moderately pigmented, wide, base

slightly concave, apex gently convex, a number of short and moderately long

setae on apical 0.35, several long setae along apical margin, numerous broad

scales covering apical 0.70-0.74, basolateral seta absent; sternum VIII moder-

ately pigmented, wide, base slightly concave, apex with a small median in-

dentation with a small lobe on each side of midline, a number of moderately

long setae apically, numerous short and moderately long setae scattered over

apical 0.90-0.92, numerous broad scales covering most of area, apical inter-

segmental fold unpigmented, VIII-S index 0. 56-0. 59; tergum IX moderately

pigmented, wide and short, ribbon-like, setae absent, covered with minute

spicules, IX-Te index 0.21-0.28; insula unpigmented, short, covered with

minute spicules, 5-6 small tuberculi, each with a minute spicule; lower vaginal

94 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

lip heavily pigmented, narrow, a few minute spicules on outer part of hinge

area, with a small caudad projecting heavily pigmented median apodeme,

lower vaginal sclerite composed of a large median heavily pigmented apically

wrinkled plate with small spicules and a narrow median indentation on apical

0.35; upper vaginal lip heavily pigmented, produced laterally, lateroposterior

corners forming small lobes, median posterior area produced into a shield

caudad of spermathecal eminence, upper vaginal sclerite heavily pigmented,

large and complex, covering most of vaginal roof; spermathecal eminence

heavily pigmented, large, deep in depth, dorsal surface with a long basal ex-

tension covered with closely matted moderately long branched spiny excres-

cences, ventral outline heart-shaped with inner portion developed into a pair

of very long slender cephalad produced arms; postgenital lobe short, moder-

ately broad, apex with a moderately deep median indentation, 15-21 setae on

each side of midline, completely covered with minute spicules, dorsal PGL

index 0.90-1.02, ventral PGL index 1.31-1.56; peri-anal membrane with

minute spicules; cercus triangular in shape, moderately long, broad at base,

apex acute, completely covered with minute spicules, dorsal surface with nu-

merous short and moderately long setae over apical 0.85-0.93, a few long

setae at apex, 0-3 broad scales, ventral surface with short and moderately

long setae on outer margin and becoming more numerous apically, cercus

index 2.98-3.13, cercus/dorsal PGL index 2. 66-2.84; 3 seminal capsules, 1

large, 1 medium and 1 small in size, heavily pigmented, ovoid, each with sev-

eral small seminal capsule pores near orifice, base of accessory gland ducts

unpigmented. i

MALE. Differs from the female by the following features. Head. Anten-

na plumose, 0.86-0.97 length of proboscis; maxillary palpus 0.12-0.13 length

of proboscis; proboscis 1.11-1.15 length of femurI. Thovax. Setal differences:

scutal fossal (7-10 anterior, 4-5 lateral), scutellum (4-7 long and 3-6 short

ones on lateral lobe, 4-7 long and 3-8 short ones on median lobe), 12-19 ante-

pronotal, 4-6 postpronotal, 16-25 propleural, 5-9 postspiracular, 7-16 on

prealar knob, 5-6 upper and 15-17 posterior mesepisternal, and 20-32 dorsad

of scale patch on mesepimeron. Legs. Posttarsi I-III (Fig. 57) each with 2

ungues, I and II each with ungues unequal, larger one with a tooth, smaller one

simple, III with ungues equal, both simple. Genitalia (Fig. 34). Distinctive as

illustrated.

DISCUSSION. Aedes virilis is very similar to the other members of the

uncus Group and no apparent differences could be found in the habitus by which

to separate them. The female genitalia of vivilis are similar to those of latz-

pennis and harrisonicus, however, they can be separated from latipennis by the

shape of sternum VIII which is without a posteriolateral lobe and the lower vag-

inal sclerite which has only a median indentation on the apical 0.35; from har-

visonicus by the shape of the lower vaginal sclerite and the narrow base of the

lower vaginal lip; and from all members of the uwncus Group by the spermathe-

cal eminence which has the inner portion produced into a pair of very long

slender arms.

The male genitalia of virilis were partially illustrated by Edwards (1917:

222) but his drawing showed little detail. Delfinado (1968, Fig. 22) copied

Edward's drawing. The genitalia were mounted on a celluloid point and were

twisted, overdecolorized and very difficult to examine. I therefore stained,

dissected and remounted the genitalia (preparation number T74. 26) in Canada

balsam on a microscope slide. The present illustration (Fig. 34) shows in

detail the unique features of the genitalia of this species. Some distinctive

features are: prosophallus very long, curved and slender; gonostylus long,

Reinert: New Interpretation of Vervallina 95

elbowed, with a moderately long slender projection near middle; and gono-

coxite with sternomesal surface with a median projection terminating in 2

apical claws (somewhat similar to dux).

The adult holotype male of vivilis is unique for the subgenus in the presence of

one long and one short seta on the median caudal area of the mesopostnotum.

This feature has not been recorded in any other species of the genus Aedes;

however, 2 genera (Heizmannia Ludlow and Haemagogus Williston) of the tribe

Aedini (tribe as defined by Belkin 1962) possess this feature. The series of

13 males and 15 females from Sabah do not possess setae on the mespostnotum

and I therefore consider their presence in the holotype to be an anomaly.

The above description is the first for the female of this species. The female

genitalia (Fig. 18) and posttarsi (Fig. 61) and the male posttarsi (Fig. 57) are

also illustrated for the first time.

TYPE-DATA. The holotype male in the BMNH contains labels which

possess the following data in addition to that given by Delfinado (1968: 41):

26/2/04; Mosq. 58; Dr. G. F. Leicester; and 1912-350.

The holotype adult is in very poor condition but the genitalia are in excellent

condition.

DISTRIBUTION. 30 specimens examined: 162, 14.

INDONESIA. Sumatra, Dermajoe (Benkoelen); 12 genitalia.

MALAYSIA. Sabah, Beaufort, Kg. Maraba, Sandakan; 159, 130%. Selangor,

Kuala Lumpur; 1° (holotype).

BIOLOGY. Adults were collected in Malaysia on a clear day resting in

bushes of a rubber estate at 1410 hours and at an elevation of 76 m.

AEDES (VERRALLINA) YERBURYI EDWARDS

(Figs. 19, 57, 62)

The following information supplements the descriptions of Edwards (1917:

222) and Wijesundara (1951: 174).

FEMALE. Head, Antenna 1.03-1.05 length of proboscis; oatiliees pal-

pus 0.19 length of proboscis; proboscis 1.08-1.13 length of femur I. Thorax,

Following setae present: 4-5 median anterior promontory; numerous acrostichal

(anterior and posterior); numerous dorsocentral (anterior and posterior); scutal

fossal (3-4 anterior, 4-6 lateral and 1-2 posterior); numerous supra-alar; 4

posterior medial scutal; 1 postalar callar; scutellar (6-7 long and 2-4 short ones

on lateral lobe, 5-6 long and 3-6 short ones on median lobe); 14-19 anteprono-

tal; 5-6 postpronotal; 15-17 propleural; 3-6 postspiracular; 4-6 upper and 8-11

posterior mesepisternal; 10-16 on prealar knob; and mesepimeron with 10-15

dorsad of scale patch, several short golden hairs near lower posterior corner

of scale patch. Prosternum, subspiracular area, paratergite, mesomeron

and metameron bare. Legs. Posttarsi I-III (Fig. 62) each with 2 ungues, I

and II with ungues equal, each with a tooth, III with ungues equal, both simple.

Wing. Remigium with 2 setae. Genitalia (Fig. 19). Very large and expanded.

MALE. Habitus similar to female. Head. Antenna plumose, 0.91 length

of proboscis; maxillary palpus 0.18 length of proboscis; proboscis 1.14 length

of femur I. Legs. Posttarsi I-II (Fig. 57) each with 2 ungues, I and II with

ungues unequal, larger one with a tooth, II with ungues equal, both simple.

Genitalia, Extremely large and expanded.

DISCUSSION. Adults resemble uncus but can be separated from this species

by the reduced number of golden hairs located caudad of the lower corner of the

mesepimeral scale patch.

96 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

DISTRIBUTION. 11 specimens examined: 88, 3c.

SRI LANKA. North Central, Anuradhapura District, Padaviya; 1.

Northern, Vavuniya District, Parayanalankulam, 25 miles NW. of Medawach-

chiya; 82, 2.

BIOLOGY. Adults were collected at an elevation of 55 m. 3

AEDES (VERRALLINA) YUSA FI BARRAUD

(Figs. 20, 22, 35, 47, 57, 62)

Aedes (Neomacleaya) siamensis Delfinado 1968, Contrib. Am. Entomol. Inst.

(Ann Arbor) 2(4): 36 (2*, “*, P, L*). NEW SYNONYM.

Aedes yusafi is the representative species for Section B and is therefore

completely redescribed below.

FEMALE. Head, Antenna dark brown, 1.11-1.21 length of proboscis,

pedicel dark brown with a few small pale brown scales mesally, flagellomere

1 with basal 0. 65-0. 80 pale and with several small brown scales near middle,

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

pus blackish-brown scaled, 4 segmented, 0.19-0.21 length of proboscis; pro-

boscis blackish-brown scaled, 1.05-1.19 length of femur I; eyes well separated

in front; ocular setae blackish-brown and moderately long; scales on head all

broad, decumbent and blackish-brown except for a few short erect forked

blackish-brown scales on occiput and a very few similar ones caudad of ocular

setae, interocular space covered with broad silvery scales, similar scales

extending caudally onto vertex for a very short distance and laterally a short

distance on ocular line, broad silvery scales also forming a broad stripe from

area in front of antepronotum to ocular line on upper lateral surface, and

postgenal scales pale brown. Thorax. Scutal integument dark brown; scutum

covered with narrow curved reddish-black scales except for a small patch of

narrow curved white scales on median anterior promontory area, a few similar

scales on anterior and lateral scutal fossal areas and at scutal angle; prescu-

tellar space bare; scutellum with a patch of narrow curved white scales on

each lobe, median lobe also usually with a few narrow curved brown scales

basally; dark reddish-black setae on the following areas: 2-4 median anterior

promontory, acrostichal (anterior and posterior), numerous dorsocentral (an-

terior and posterior), scutal fossal (3-6 anterior, 4-8 lateral and 1-2 posterior),

numerous supra-alar, 4-5 posterior medial scutal, 1 postalar callar and scutel-

lar (5-8 long and 3-5 short ones on lateral lobe, 4-6 long and 4 short ones on

median lobe); pleural integument dark brown; antepronota widely separated,

_ covered with broad silvery scales, 9-12 long and short dark setae; postprono-

tum with 5-8 posterior golden-brown setae; propleuron with a patch of broad

silvery scales, 6-8 golden setae; prosternum, subspiracular area, paratergite,

mesomeron and metameron bare; postspiracular area covered with broad sil-

very scales, 3-5 golden setae; mesepisternum with an upper and a lower patch

of broad silvery scales, 6-8 posterior golden setae; prealar knob with 6-12 }

golden setae; mesepimeron with a large patch of broad silvery scales near

middle, 8-11 golden setae dorsad of scale patch. Legs. Coxae I-III each with

several golden-brown setae and a patch of broad silvery scales, I and II each

with scales on anterior surface, III with a narrow band of scales on anterior

margin and forming a broad patch dorsally; trochanters I-III each with several

broad pale scales and a few short pale setae; femora I and II each with anterior

surface brown scaled and with white scales forming a narrow basal band, I with

Reinert: New Interpretation of Verrallina 97

white scales also on dorsobasal area, II also with a narrow longitudinal stripe

of white scales extending from base to near apex, III with anterior and posterior

surfaces white scaled with dorsoapical area brown scaled, I and II each with

posterior surface white scaled with a narrow longitudinal brown scaled stripe

from base to apex, stripe ventral on I and dorsal on II; tibiae I-III each dark

brown scaled, I also with a posteroventral longitudinal white scaled stripe ex-

tending from base to apex, II also with posterior surface white scaled; tarsi I-

III each dark brown scaled; posttarsi I-III (Fig. 62) each with 2 ungues, I, II

and III each with ungues equal, each with a tooth. Wing. Dorsal and ventral

veins with brown scales, costa also with a few white scales at base and remigi-

um with a few white scales on posterior margin; alula with several narrow

brown scales on margin; upper calypter with several golden hairs on margin; 1

remigial seta. Halter, Pedicel pale; capitellum dark brown scaled. Abdomen.

Terga blackish-brown scaled, laterotergite of I with a large rectangular patch

of white scales, II-VII each with a large, laterobasal patch of white scales which

extend slightly onto tergal surface; sternum II white scaled; sterna III-VII white

scaled with a broad apical band of brown scales, VII with white scaled area re-

duced; terga and sterna with numerous golden setae, mostly along posterior

margins. Genitalia (Figs. 20 and 22). Tergum VIII moderately pigmented,

base slightly concave, apex straight, several short and moderately long setae

apically, numerous short setae scattered over apical 0. 77-0. 88, basolateral

seta absent, covered with minute spicules, numerous broad scales covering

apical 0.50-0.65, basal 0.2-0.7 retracted into segment VII, VIII-Te index

0.70-0.79, VIII-Te/IX-Te index 2.67-2.85; sternum VIII moderately pigmented

with apical 0.35-0.39 darker, base concave mesally, apex with a moderately

deep median indentation with a small lobe on each side of midline, numerous

short stout setae on lobes, numerous short and a few moderately long setae

scattered over entire surface except basal 0.09-0.14 and a small basolateral

area, covered with minute spicules, a number of broad scales forming a latero-

apical patch, apical intersegmental fold unpigmented, VIII-S index 0. 67-0. 73;

tergum IX moderately pigmented with basomedian area lightly pigmented, wide,

short, base concave, apex with a wide moderately deep median indentation and

with 1-4 (usually 3-4) short setae apically on each side of midline, covered

with minute spicules, IX-Te index 0.46-0.49; insula absent, however, median

apical U-shaped portion of lower vaginal sclerite apparently serves in the capa-

city of the insula since it is situated in its normal position and lies in the space

between the apical median lobes of VIII-S; lower vaginal lip lightly to moder-

ately pigmented, forming a narrow band along basal 0.75 of area, apical 0.25

not developed, base attached near midpoint of lateral portion of upper vaginal

lip, covered with small spicules, lower vaginal sclerite forming a heavily pig-

mented ribbon-like strip attached to lower vaginal lip, extending around lower

vaginal membrane and fused at apex forming a projected wrinkled U-shaped

structure, sclerite apparently replaces apical 0.25 of lower vaginal lip and

insula, base of sclerite articulates with the lateral wing-like projection of the

spermathecal eminence "pouch"; upper vaginal lip moderately pigmented, nar-

row, median posterior portion lightly pigmented as base and projected into a

large heavily pigmented cephalic shield which covers posterior portion of

spermathecal eminence, ventral surface of shield attached to caudal surface

of spermathecal eminence, upper vaginal sclerite moderately pigmented, very

large, well developed, base attached along entire lateral margin of upper vaginal

lip and with a heavily pigmented apodeme extending from lateroposterior mar-

gin of upper vaginal lip to anterior mesal area; spermathecal eminence heavily

pigmented, shallow in depth, egg-shaped in dorsal outline, a heavily pigmented

98 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

cephalad produced pouch attached to anterior tergal portion of spermathecal

eminence, pouch with inner surface lined with numerous hair-like spicules

and with a laterobasal wing-like structure on each side; postgenital lobe short,

narrow, apex flat to broadly rounded, 8-10 setae on each side of midline,

completely covered with small spicules, dorsal PGL index 0. 96-1.24, ventral

PGL index 1. 69-1.96; peri-anal membrane with minute spicules; cercus tri-

angular in shape, moderately long and broad at base, apex acute, completely

covered with small spicules, dorsal surface with a number of moderately long

and a few short setae on apical 0.91-0.95, ventral surface with a few short

setae along outer area, a number (12 or more) of broad scales on apical 0. 75-

0. 84 of dorsal surface, cercus index 3.10-3.28, cercus/dorsal PGL index

3.10-3.65; 3 seminal capsules, 1 large, 1 medium and 1 small in size, heavily

pigmented, nearly spherical, a few seminal capsule pores near orifice, base

of accessory gland duct lightly to moderately pigmented.

MALE. Similar to female in general habitus but with the following differ-

ences. Head. Antenna plumose, whorls with 8 setae, 1.15-1.27 length of

proboscis; maxillary palpus 0.19-0.17 length of proboscis; proboscis 1. 01-

1.11 length of femur I. Thovax. Scutellum with narrow curved brown scales

on each lobe, setae (4-5 long and 2-3 short ones on lateral lobe, 4 long and

2-3 short ones on median lobe); 5-9 antepronotal setae; 3-5 postpronotal setae;

4-5 propleural setae; 1-2 postspiracular setae; 4-5 posterior mesepisternal

setae; 4-7 prealar knob setae; 3-7 mesepimeral setae dorsad of scale patch.

Legs. Tibiae I-III each dark brown scaled; posttarsi I-III (Fig. 57) each with

2 ungues, I with ungues unequal, larger ungue with a tooth, II with ungues

unequal, larger ungue with a tooth, III with ungues equal, one with a tooth.

Wing. Costa and remigium without white scales. Abdomen. Terga with latero-

basal white scaled spots small; sterna II-VII pale scaled, VIII brown scaled.

Genitalia (Fig. 35). Tergum IX heavily pigmented, formed into a moderately

broad band mesally and expanded laterally on each side into a triangular lobe,

fused with tergum X, setae absent, connected laterally to sternum IX by a

narrow heavily pigmented band; gonocoxite short, broad, heavily pigmented,

tergobasal portion of mesal surface membranous with minute spicules, dorsal

surface with a few short and moderately long setae, 1-2 long setae near base

of gonostylus, ventral surface produced into a large apicomesal lobe which is

heavily pigmented on both sternal and tergal areas, apex expanded into a small

finger-like lobelet with a short seta near apex and 4-5 short setae at base,

sternal area of lobe with 2 long stout setae at base of outer margin and several

short setae over remainder of area, several broad scales on lateral and ventral

surfaces of gonocoxite; gonostylus heavily pigmented, moderately long, approxi-

mately 0.6 length of gonocoxite, basal 0.28 moderately broad, middle 0. 38

expanded and with 44-54 short setae, apical 0.34 narrow, recurved and with

apex very heavily pigmented, gonostylar claw absent, attached subapically to

gonocoxite; basal mesal lobe broad, moderately pigmented, covered with small

spicules, 26-28 very short setae near center of lobe, 2 heavily pigmented short

stout spine-like structures near middle of sternomesal margin, lobes connected

mesally by a broad band which forms a shallow trough ventrad of apical portion

of prosophallus, sternomesal and outer margins of basal mesal lobe fused with

gonocoxite; proctiger with paraproct heavily pigmented, formed into a long arm

which is moderately broad basally and with apical 0.45 acuminate and curved

slightly mesad, dorsal surface with a few small spicules on basal 0.4 anda

small heavily pigmented lump near middle at point where cercus attaches,

sternobasal area of paraproct formed into a small broad projection which ex-

tends ventrad and articulates at a point on the dorsomesal surface of the para-

Reinert: New Interpretation of Verrallina 99

meral apodeme near base of opisthophallus, proctiger strongly fused with ter-

eum X, cercus membranous and without cercal setae; tergum X heavily pig-

mented, formed into a plate between proctiger and caudolateral portion of ter-

sum IX, not connected mesally; phallosome complex, opisthophallus--consists

of a moderately pigmented tergal transverse bridge between the parameral apo-

demes, base attached to a dorsomesal extension of the caudal outer portion of

paramere apodeme, located dorsad of phallus and prosophallus and ventrad of

proctiger, phallus--consists of a pair of short heavily pigmented narrowly

separated arms which are fused at base, sternolateral area of base extended

into a long arm which is fused with tergomesal portion of prosophallus base,

apex of phallus formed into a tergal penis filament which consists of a pair of

long bars extending cephalad from phallus apex, bars connected by a narrow

transverse bridge near middle, apical 0.5 of bars curved dorsad and attached

to sternal surface of opisthophallus, prosophallus--consists of a pair of long

heavily pigmented acuminate arms which have their bases widely separated

and their apices contiguous, base of arm attached to tergomesal portion of

paramere and located ventrad and laterad of phallus, apical portion of arm

projecting caudad and dorsad of phallus, prosophalic arm approximately 2. 63

length of phallic arm (measured along midline of genitalia), paramere--heavily

pigmented, broad, long (approximately 2.32 length of phallic arm), articulates

at 2 points near middle with caudal area of parameral apodeme, most inner

point of apodeme attached to sternobasal area of prosophallic arm, parameral

apodeme--moderately to heavily pigmented, very long (approximately 1. 68

length of paramere), attached near middle of outer surface with tergobasal

apodeme of gonocoxite; sternum IX heavily pigmented throughout, large, broad,

strongly attached along lateral surfaces to sternomesal areas of gonocoxites,

2-8 short setae near center of caudal area.

PUPA (Fig. 47). Chaetotaxy as recorded in Table 14. Ocular plate with

area of well developed cuticular facets. Metanotal plate and areas of cephalo-

thorax heavily pigmented. Respivatory trumpet. Moderately pigmented; short;

hair-like spicules on distal 0.40-0.43 of inner surface; index 3. 00-4.138, aver-

age 3.58. Abdomen. Setae 6, 9-VII weakly developed; 4-VIII double to 4 bran-

ched; 9-VIII single or double. Paddle. Ovoid; serrations on most of basal 0.85

of outer margin, minute proximally and well developed distally; minute spicules

on apical 0.15 of outer and apical 0.09 of inner margins; midrib reaches apex;

seta 1-P weakly developed, moderately long, single; index 1.27-1.47, average

1.34.

LARVA (Fig. 57). Chaetotaxy as figured and recorded in Table 27 and is

based on 3 damaged, associated skins from the type-series of stamensis. Head,

Seta 4-C triple to 4 branched; 5-C single or double; 6-C double or triple; 7-C double to

4 branched; 12-C single to 4 branched; dorsomentum with 29-34 teeth. Antenna,

Short, lightly pigmented, with several small scattered spicules; seta 1-A short,

thickened, single; 2-A long, with a subapical constriction; 3-A short, 0.27-

0.31 length of 2-A; 4-A moderately long, 0. 56-0. 62 length of 2-A; 6-A short,

0. 22-0. 37 length of 2-A. Abdomen. Seta 1-VIII double or triple; 3-VIII with

4-8 branches; 5-VIII with 5-6 branches; 2-X triple or 4 branched, branches

approximately equal in length; ventral brush with 8 setae on grid and 2 precratal

shorter ones, all multiple branched; saddle moderately pigmented, incompletely

rings segment X, acus absent; 4 moderately long anal papillae; comb with 8-12

scales arranged in a single slightly curved irregular row each scale with a stout

median apical spine and small denticles on laterobasal areas. Siphon. Mod-

erately pigmented; acus well developed; index 1.58-2.08, average 1.82; pec-

ten on basal 0. 56-0. 58 of siphon, composed of 11-15 (usually 14) teeth, apical

100 - Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

1-2 teeth longer and wider spaced than remainder of teeth, each tooth long,

slender and with a small stout ventral denticle near base; seta 1-S short,

double to 4 branched, base attached on basal 0. 67-0. 68 of siphon distad of last

pecten tooth.

DISCUSSION. I have compared the female holotype of yusafi with the holo-

type male, allotype and paratypes of siamensis and determine them to be con-

specific. The female genitalia of this species are very distinctive in the

shape of the median portion of the upper vaginal lip and the presence of sev-

eral setae on tergum IX and therefore leaves no doubt about the synonymy or

association of sexes, especially since both males and females of siamensis

were reared from a batch of eggs from a wild caught female.

The holotype of stamensis was collected in 1967, according to the collec-

tion data sheet and adult label, but Delfinado (1968: 38) erroneously reported

it collected in 1965. | |

Two male paratypes listed by Delfinado (1968: 38) could not be located in

the USNM, SEAMP, BMNH or Bishop Museum and are presumed lost.

_Posttarsus II of the holotype male of stamensis has the larger ungue with

an abnormal double tooth and the second tooth on both the larger ungues of

posttarsi I and II (paratype) are actually hair-like spicules (see Fig. 26,

Delfinado 1968).

The male and female genitalia of all species in Section B are very distinc-

tive and possess excellent characters for separating the species. Figures 21

and 22 compare the spermathecal eminence "pouch", median area of the upper

vaginal lip and tergum IX of all the known members of this section. Tergum

IX of the female genitalia of yusafi is distinctive in possessing several setae

on each side of the midline.

Features of the adults of Section B are compared by Reinert (1973b: 257).

TYPE-DATA. Aedes (Aedes) yusafi Barraud, holotype female, genitalia

mounted in Canada balsam on a microscope slide, INDIA, Delhi, Roshanara

Garden, IV. 1914, S. R. Christophers collector, specimen number 2555, Mus.

Reg. 930, B. M. 1935-622, genitalia preparation number T73.580, biting

collection, and deposited in the BMNH. Aedes (Neomacleaya) siamensis Del-

finado, holotype male, associated pupal skin and genitalia mounted in Canada

balsam on microscope slides, THAILAND, Lampang, Ban Pong Tao, 21 April

1967, E. L. Peyton and Kol Mongkolpanya collectors, collection number 01998-

103, reared from an egg obtained from a female which had fed on man; allo-

type with associated pupal and larval skins and genitalia mounted in Canada

balsam on microscope slides, collection number 01998-3, other data same as

holotype; paratypes, one male with associated pupal and larval skins, collec-

tion number 01998-2, 8 males with associated pupal skins, collection numbers

01998-100, 01998-101, 01998-102, 01998-104, 01998-105, 01998-1007, 01998-

108, 01998-109, one female with associated pupal and larval skins, collection

number 01998-4, and one female with associated pupal skin, collection number

01998-106, other data same as holotype; paratypes, 12 females, collection

number 01974, same data as holotype except for the following: collected feed-

ing on man at 1000 hours, 21 April 1967, on a cloudy day near a medium size

fresh water ground pool situated beneath a bamboo clump at an elevation of

200 m, females isolated and eggs deposited from which the 01998 series of

specimens were reared (the above information about the types of siamensis was

taken from the collection data sheets); the above types of siamensis are de-

posited in the USNM, BMNH and Bishop Museum.

DISTRIBUTION. 58 specimens examined: 329, 100", 13 with associated

skins (10 p, 3 1).

Reinert: New Interpretation of Vervallina 101

INDIA. Delhi, Roshanara Garden; 1° (holotype of yusa/i).

THAILAND. Lampang, Ban Pong Tao; 1“ p (holotype of siamensis), 12 pl

(allotype of stamensis), 12 pl, 12 p, 122, 10 pl, 8 p (all paratypes of siamen-

sis), 162. 7

BIOLOGY. Numerous females in Thailand were collected biting man at

1000 hours, 21 April 1967, near a medium size ground pool containing fresh,

unmoving water, located beneath a clump of bamboo and at an elevation of

200m. The adults presumedly had emerged from the ground pool but no im-

matures could be found (E. L. Peyton, personal communication). Several of

the blooded females were isolated and eggs obtained. The type-series of sia-

mensis consists of specimens reared from these eggs and the above mentioned

biting females.

~ ACKNOWLEDGMENTS

Grateful appreciation is expressed to: Bruce F. Eldridge and Ronald A.

Ward, Walter Reed Army Institute of Research and E. L. Peyton, Medical

Entomology Project (MEP), for critically reading the manuscript; to Douglas

J. Gould, Southeast Asia Treaty Organization Medical Research Laboratory,

Bangkok, Thailand, and Shivaji Ramalingam, University of Malaya, Kuala

Lumpur, Malaysia, for the collection and preparation of many specimens; to

Peter F. Mattingly, British Museum (Natural History), London, Wallace A.

Steffan, Bernice P. Bishop Museum, Hawaii, J-M. Klein, Office de la Recherche

Scientifique et Technique Outre-Mer, Bondy, France, Kazuo Tanaka and

Edward S. Saugstad, U. S. Army Medical Laboratory (Pacific), Japan, and

John N. Belkin, University of California, Los Angeles, for the generous loans

of types and museum specimens; to Elizabeth N. Marks, Queensland Institute

of Medical Research, Brisbane, Australia, for providing notes on the types of

Aedes cunninghami; to Young T. Sohn, MEP, for preparing the numerous de-

tailed illustrations; to Robert H. Hochman, MEP, for helpful suggestions and

for making many of the larval and pupal setal counts; to Carolyn M. Emerick,

Virginia M. Ford, Thomas V. Gaffigan and Susan Hevel, MEP, for laboratory

assistance; and Janet D. Rupp, MEP, for typing the manuscript for offset re-

production.

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Oo: 202-62.

Reinert: New Interpretation of Vervallina 105

1974. Terminology and preparation techniques of the female genitalia

of aedine mosquitoes (Diptera: Culicidae). Mosq. Syst. 6: 46-

06,

SASA, M.

1949. Redescription of Aedes nobukonis Yamada, 1932, with desig-

nation of its subgenus. Jap. J. Exp. Med. 20: 409-11.

SLOOFF, R.

1972. | Mosquitoes collected in the British Solomon Islands Protector-

ate between March 1964 and October 1968 (Diptera: Culicidae).

Entomol. Ber. (Amst. ) 32: 171-81.

STEFFAN, W. A.

1966. A checklist and review of the mosquitoes of the Papuan sub-

region (Diptera: Culicidae). J. Med. Entomol. 3: 179-237.

STONE, A.

1956. Corrections in the taxonomy and nomenclature of mosquitoes

(Diptera, Culicidae). Proc. Entomol. Soc. Wash. 58: 333-44.

STONE, A., K. L. KNIGHT and H. STARCKE.

1959. A synoptic catalog of the mosquitoes of the world (Diptera, Culi-

cidae). Thomas Say Found., Entomol. Soc. Am. vol. VI, 358p.

TANAKA, K.

1071. Mosquitoes of the Ryukyu Islands. Jap. J. Sanit. Zool. 21: 4.

TANAKA, K. and K. MIZUSAWA.

1978. Two new species of the genus Aedes (Neomacleaya) from the

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THEOBALD, F. V.

1901. A monograph of the Culicidae or mosquitoes. Mainly compiled

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1907. A monograph of the Culicidae or mosquitoes. Mainly com-

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1910. A monograph of the Culicidae or mosquitoes. Mainly com-

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WIJESUNDARA, D. P.

1951. | Notesonspecies of the subgenus Aedes Meigen (sens. str. ) in Cey-

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173-9. 3

106 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

YAMADA, S.

1932. Nippon konchu zukan. Iconographia Insectorum Japonicorum,

1st Ed. Hokuryukan, Tokyo. 2241 p.

LIST OF FIGURES

1. Distribution of the species of Vervallina

2. Aedes (Verrallina) abditus--female genitalia

3. Aedes (Verrallina) agrestis, comatus and nigrotarsis --female genitalia

4. Aedes (Verrallina) butleri--female genitalia

0. Aedes (Verrallina) carmenti--female genitalia

6. Aedes (Verrallina) cautus--female genitalia

7. Aedes (Verrallina) comosus--female genitalia and tarsomeres 9 and post-

tarsi

8. Aedes (Verrallina) harrisonicus--female genitalia

9. Aedes (Verrallina) incertus--female genitalia

10. Aedes (Verrallina) indicus--female genitalia

11. Aedes (Verrallina) johorensis--female genitalia

12. Aedes (Verrallina) lankaensis--female genitalia

13. Aedes (Verrallina) latipennis and notabilis--female genitalia

14. Aedes (Verrallina) leicesteri--female genitalia

15. Aedes (Verrvallina) petroelephantus--female genitalia

16. Aedes (Verrallina) pseudomediofasciatus --female genitalia

17. Aedes (Verrallina) sabahensis--female genitalia

18. Aedes (Verrallina) virilis--female genitalia

19. Aedes (Verrallina) yerburyi--female genitalia

20. Aedes (Verrallina) yusafi--female genitalia

21. Aedes (Verrallina) fragilis, hamistylus, indecorabilis, stungus and uni-

formis--female genitalia

22. Aedes (Verrallina) abditus, nobukonis, pahangi, robertsi, and yusafi--

female genitalia; johnsoni and margarsen--female

prosterna

23. Aedes (Vervallina) andamanensis, atrius, campylostylus, clavatus, cretatus,

cyrtolabis, dux, lugubvis, macrodixoa, margarsen, neéo-

macrodixoa, varus, uncus, vallistris and varietas --

terga IX of female genitalia

24. Aedes (Verrallina) azureosquamatus, cuccioi, foliformis, funereus,

killertonis, lineatus, mccormicki, milnensis, multifoli-

um, pipkini, quadrifolium, quadrispinatus, sentanius,

trispinatus and variabilis --terga IX of female genitalia

25. Aedes (Verrallina) incertus, panayensis and varus--spermathecal emin-

ences of female genitalia; szbbosus--tergum IX of female

cenitalia; adustus, andamanensis, johnsoni and margar-

sen--basal mesal lobes of male genitalia; zwcertus and

prioekanensis--gonocoxite apicomesal lobes of male

genitalia

26. Aedes (Verrallina) butleri--male genitalia

27. Aedes (Verrallina) carmenti--male genitalia

28. Aedes (Verrallina) consonensis--male genitalia

29. Aedes (Verrallina) fragilis--male genitalia

30. Aedes (Verrallina) indicus--male genitalia

. Aedes (Verrallina) pseudovarietas--male genitalia

08.

o9.

60.

61.

62.

63.

64.

65.

66.

Reinert: New Interpretation of Verrallina 107

Aedes (Verrallina) vamalingami--male genitalia

. Aedes (Verrallina) sohni--male genitalia

. Aedes (Verrallina) virilis--male genitalia

Aedes (Verrallina) yusafi--male genitalia

. Aedes (Verrallina) adustus--pupa

Aedes (Verrallina) butleri--pupa.

. Aedes (Verrallina) carmenti--pupa

. Aedes (Vervallina) clavatus--pupa

. Aedes (Verrallina) consonensis--pupa

. Aedes (Verrallina) cyrtolabis --pupa

. Aedes (Verrallina) leicesteri--pupa

. Aedes (Verrallina) lugubris--pupa

. Aedes (Verrallina) indicus --pupa

. Aedes (Verrvallina) torosus--pupa

. Aedes (Verrallina) vallistris--pupa

. Aedes (Verrallina) yusafi--pupa

. Aedes (Verrallina) adustus and torosus--larvae

. Aedes (Vervallina) butleri--larva

. Aedes (Verrallina) carmenti--larva

. Aedes (Verrallina) consonensis--larva; dux--egg

. Aedes (Verrallina) cyrtolabis and vallistris--larvae

. Aedes (Verrallina) gibbosus--larva

. Aedes (Verrallina) indicus --larva

. Aedes (Verrallina) uncus --larva

Aedes (Verrallina) leicesteri--larva; leicesteri and sohni--male tarso-

meres 5 and posttarsi; andamanensis and butleri--

male terga VIII

Aedes (Verrallina) yusafi--larva; torosus, uncus, vallistris, virilis,

yerburyi and yusafi--male tarsomeres 5 and posttarsi

Aedes (Verrallina) adustus, andamanensis, butleri, campylostylus,

carmenti, clavatus, cretatus, cuccioi, cyrtolabis, dux,

embiensis, gibbosus, hamistylus and indicus --male

tarsomeres 5 and posttarsi

Aedes (Verrallina) johnsoni, macrodixoa, margarsen, mccormicki,

nigrotarsis, panayensis, pseudomediofasciatus,

pseudovarietas, quadrispinatus, vamalingami,

rami, veesi, sentanius and singularis --male tarso-

meres 5 and posttarsi

Aedes (Verrallina) adustus, andamanensis, atrius, azureosquamatus,

butleri, campylostylus, carmenti, clavatus, cucctiot,

dux, foliformis, hamistylus, indicus and johnsoni--

female tarsomeres 5 and posttarsi

Aedes (Verrallina) johorensis, latipennis, leicesteri, lugubris, macro-

dixoa, mccormicki, neomacrodixoa, nigrotarsis,

nobukonis, notabilis, pavasimilis, pipkinit, pseudo-

mediofasciatus and virilis --female tarsomeres 5 and

posttarsi

Aedes (Verrallina) quadrispinatus, reesi, sabahensis, sentanius, singularis,

trispinatus, uncus, uniformis, vallistris, vanapus, vari-

abilis, varietas, yerburyi and yusafi--female tarso-

meres 5 and posttarsi

Aedes (Verrallina) butleri--adult morphology —

Aedes (Verrallina) indicus--adult morphology

Aedes (Verrallina) johorensis--adult morphology

Aedes (Verrallina) sabahensis--adult morphology

108

BML

Gc

Gs

IX-S

IxX-Te

OP

CT

I- VIII

Wiel ivencl

Ee, ele se |

Ue a ate | roa

Hood eas

Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

LIST OF FIGURE ABBREVIATIONS

Female Genitalia

Accessory gland duct base LVS

Cercus PGL

Spiny excrescences on SE SCa

Line of attachment of peri- SCaP

anal membrane to dorsal SE

surface of PGL Tu

Hinge UVL

Insula UVS

Tergum IX VITI-S

Lower vaginal lip Vill-Te

Male Genitalia

Basal mesal lobe PaA

Gonocoxite Par

Gonostylus PH

Sternum IX PO

Tergum IX Ppr

Opisthophallus Pr

X-Te

Pupa

Cephalothorax MP

Abdominal segments I-VIIT P

aa

Larva

Antenna M

Head Pp

Comb scale jae

Dorsomentum S

Abdominal segments T

I-VII, X

ee ese bere aia ace, re ak nol ei pe

emis | neat | eA

Sell il

I

Hate

Slt acthcott aw

Lower vaginal sclerite

Postgenital lobe

Seminal capsule

Seminal capsule pore

Spermathecal eminence

Tuberculus

Upper vaginal lip

Upper vaginal sclerite

Sternum VIII

Tergum VIII

Parameral apodeme

Paramere

Phallus

Prosophallus

Paraproct

Proctiger

Tergum X

Metanotal plate

Paddle

Respiratory trumpet

Mesothorax

Prothorax

Pecten tooth

Siphon

Metathorax

gq uoljzeS

II Set4ag “Ww YoNDeS

{J set4ag “WwW UuoljzaSg

| S@14ag “YW UolyDaS

ventral

Vy.

Levante

Vyeuay

eg:

\ at as

Sonali i

\ : ;

PSU Nap eerie pues

NENU Dr ROEG at ere eal RC) Yove

Waal Neve hy lines a ;

v" Nid J °

PGE Heindipuce Lanza latas ts

VN

ween ae CYA LT QZ

OV irre

Oo AZ

Vay Ht secede A:

abditus

Vill-Te

fn i om

Bay

CLIK

i

e - ventral

=o:

=< ve =

SSS ee

SS mae = c= : Z 3

ae i

' M

yi

VT

ia t ip

nigrotarsis Vill-Te

O

N

ventral

/

)

\\

nN bill of

f AlN Te \

ane WPA Ne 8) \\

ANP A) \ |

1 \\}: ap ss 5) \

2

\

4G

i ( (3 \\

a a

Vill-Te

butlers

re

1 heeehy

/ —

WTS

} I ii

Be es

: )

Bi

Teta

jeg

/\ \ ee

ventral

\

carmenti

ids |

ik

gin,

ventral

SCa

0:2

its \ 5

YL CAN as

\ 2 ee

iN oA va X() (i fA 4 4 Wy f

toh Q\\ Ny i) M, i

a “) iy ) ( ewe WN hing i A Gh

\ WN ne i, a we a aie f ee Hk

\

VI WAS

AN

aad \ \" * ‘s Ae \ uw vy rey ee ey ae oe / fy fe

ON NSS SOE ama \s My \\ e are i! a: i Rabie oe opie LM / My: “4g

ey, ‘\ ee viel Ua seed kt Gane ets i SN Geaae a4 Re i Bh) ee.

AALS Ne) Nie Aire tae HM Ul nace Hai TS; WG PDI LE ty Sate EGP Od sg g

WN Sen 8 AN Se eye Cede s Vers cineoee : j pees ares eye, ae $ 4

IX-Te

cautus

ventral

Comosus

Vill-Te

Aarrisonicus

Viil-Te

incertus

Fig.10

jan es

Pecne te

tS

i):

i) | fees

Nee alae Vi etn

Am we Ween

% SS 1 ite

Re ele raion ao

Noe acaale niga aac

pa eats

S29 oe

er —Tu

ventral

\ Ae

AAAS

fia

indicus

Fig. 11

we

1A

ene

mn i" iN AN i

0.2

ventral

SCa

os ,

Ay cua (ey :

; iby } \

. S Br ar PSS ial bag

EY pee t

, Sys Ls er \ Uae iar ed Ne ty es uaa

ie Ver ay

Vill-Te

Johorensis

33 sal Ns

qr)

t

“ 4 Ree ~

wpe Y ae ‘ ER - thats ; A

ventral

a7 7; aha dently ta

, ’ Po eh.

a

ae

ica

Iptien date

iy - r

\ ‘ if

Saal WILE Li)!

lankaensis

t

(PR oun ety fil

OHV Al MY AANA

We nile

NY),

' \ :

} ff ke and Vitae BEY Aa x Tus

i 1c! bey pe WYf#. 1.1.) FAW \ ‘ \*\.

f { y'. jeaelale Hy qbehe BG ral\ 4 i ‘ yi

; Nk Me lf! A Bg ake ZANE I A A NAIRIN & ; M\

i nt Wh Ve UL Fb Wik IE AN vs if «( ie and PF AN, Mth

| ay bie ||| Bae N ERAN AE eS a NUN Y

x . Lae Reed, he if ef, be \ ; :

Ny VEAL © MN] 07S | CA A Cee

f DD be oy FU fii: fi AP OL! 1 8-\ iY! i; ; 5

yu by aay Atal at ! fil 1) , ae i]

. | i} . / / f f i

ay! Lh, _, | | j Wa \\r

!

\

Z

WW

i NS

/

.

WOT

ae

Lip

i

\ WW rf; vk | oy LUG va \\( A (\ ie

q V\WWAY A a KUN fo A iti | hy Fatty be

: DADE al . r ry ly id ae j ;

an\ A yt i

Vit! \) fl \ ba Ale MY, hil, ; bd

i | ag a Ae eres Vac eG INR GHP:

i \ \l i Vale dan pa Aba iphda tie | ie : : ;

1 | ' ips | (eel Pits 4

aN: Uc Nang er elf Wl er pa Valea epee . ae

\ aes lies) ray tS Pets Re Gl Tg

1 el yal be pae ee Cy Raph PS Bap eee.

= sR area eae ae

ventral

=

ea

—S, Ss = Z

es ee

ees

Late

ih we ‘ fis AY ! e% “in? r 1

\ n aati ue / f

Vy i) x K y } aN \

ff ( \ ‘

\\

iy i

I

\

—— See Ses

=> <~SSS = yar

Ss 20 Se

2 <= S aS

SS —— pak a GR

PTs SS -> ~S =

> ~—se este.

=< VSS

LALA

ct) 4 | '

POLINA iy.

i i F : |

|

\ |

' ‘ \

Tet

SS" EN

— =<

SSS % ==

Sa ==>

7 —S

Vill-Te

notabilis

5 oy Gee

cod BP ee

ar) eaten

\

\

io Vy ' “

fs:

Wavy

‘fs

j o/

Hf AY 0.3

pl arn : i

UWA i 8 NVI) LM), ‘ K

VOR ory 3 We is

i i CL) 15);

. Wy ly Wey)

Wil Wa Oy te 4

Won

4 ad BR YS c

ae eb SY ape

: Vat Ak.

EER: Os 1

yee ag

i,

}

yey

/

| U

WM Sac i

SYS if. AN

Oa.

Vill-Te

latipennis

Fig. 14

Sa a. SCa

: a

ventral

apace:

Sie Ca pete :

Chak re be iaeaaed WS haha rey et to by

oat

ah

Vitl- Te

leicesteri

| Meg thy boty

: ‘ it |

3 hu :

} | \ \ \ ] 4

| : phe ty IN By wees Y. Nae

ret |

| be penn UT os 4 \ {

See

LVS

em

ventral

SCa

| te A

We

: Vill-Te

ioe ee

ava Warn eee bi les] 4 iy

i NGA earl Al AN A Leal

ye “A i I nny e ty

|

iN ny |

0.2

1

) 8

Lope ay ed

Lots

SEX

uf \

eA Ney ee

\

ee Sea

\

va

Siam Ses Sate alma

IX-Te

pseudomediofasci/atus

—

ro

ee

—

ss

——s

_—

=

ae

Ses

as ae

=e

SSS

SS Fe Sia

£, a i as, 7s | ee

| davai i S ;

LIK WEA i. at Nyy WS

ate A C9 We

Vi ie

VII-S

, Sean ee me r

ae * oo )

i ; . “as P

Geel ae BY

Bess, Pile sess EN

ane ir. oe el

Ort Bee ce. KN

fe pay rae a

% iy ‘ee a

Ree: eae

Re ey §

Oxt ay Biba a

eA ee

re B08

eae ve

eID

0:2

\ tA \ : {hie Me AY | | |

ee Re

a aie / ‘ ve int) i th i AN i fh Mth ! ; i Hy

Ooi a Mb K SD atid Fg

NS A Sap M ur ve A Oa nee

ae ; ; We me ba ie

Nei nn

aan NN ny mM

“ “— i

™ ‘ ,

i

dorsal

9 y

Way

hp MID 3

Vill-Te

IX-Te

Sabahensis

Fig.18

SCa

Vill-Te

virilis

ET

fobs

a»

eH,

Bie,

AN

LED

et

ELL.

<x

NI

UVS

Mh.

,

VEhc

4]

vs

Ce aig yerburyi

SCa

Vill-Te

ventral

yusafi

i

hk td

VaR eH Tekan ened:

ha ita abel ss

ey SYS tes rt ay

fragilis hamistylus stungus

(Holotype ma/aya) (Paratype) (Holotype)

untformis indecorabilis indecorabilis

(of Klein, 1973 Cambodia) (Cotype of Leicester, 1908 Malaysia)

NRE a

Cae

Nee

WAY \\

abditus pahangi yusafti

(Holotype) (Holotype) (Holotype)

propleuron

prosternum

Coxa I.

?

margarsen

2.0

nobukonis robertsi johnsoni

(Paratype ishigakiensis) (Paratype)

FEMALE TERGUM IX Fig. 23

SECTION A, SERIES Il -é Il

Cea

a

OY oes

andamanensis rer: neomacroadixoa

ae) Zoos +———— Q.] ———4 + — 0.2

atrius ux rarus

+—— 0.2 r—_— 0.2 ——__

campylostylus ae uNCUS

r-—— 0.2—— aE SRS ESSE 9) 7 REE SY t————0.2 4

clavatus macrodixoa vallistris

cD

SUE 6 Oy eee pe iy gy RE CEES We

eA Y Sohn

cretatus margarsen varietas

FEMALE TERGUM 1X

SECTION A, SURIES

Se Oe 7 ai -———— ae 1———

0.1

azureosquamatus — /ineatus quadrifolium

+—_—__— 0] +—_—_— 0.]

CUCCI/O/ mecormiki sundae

Pe See 0.1% +—— 0.] ——H

foliformis milnensis sentanius

}-——_——-— 0.1 + 0.] ———+ 0.1

funereus multifolium trispinatus

one oo) ay

Sperry a

teeter: t+ 0.] ———+4

killer tonis pipkini eee.

Fig. 25

een kos

BML \ BML \ -

od ‘ ele ae

margarsen johnsoni adustus andamanensis

incertus

prioekanensis

| gibbosus

incertus

butleri

/

; hese]:

neler / finde

Racin ae

. popes

St PU J ef pet eae

Cae VA Me eee

_.—H

Ccarment/

dorsal

consonensis

Fig. 29

dorsal

fragilis

indicus

Fig. 31

IX,X-Te and Pr

pseudovarietas

dorsal

ramalingam!

1X,X-Te and Pr

eae os

mA HAWES

BML

tergobasal

Ko xe) Y . apodeme of

Ne Me Wy : gonocoxite

| JHPaA PH po

dorsal

IX-S IX,X-Te and Pr

virilis

dorsal

t+—__—0.] —_—_—_

IX, X-Te and Pr

WV Y

hi) yl wd Wulf

— eae.

| = 4

adustus

Fig.37

but/eri

carmentl!

Fig. 39

clavatus

consonens/s

Fig. 41

cyrtolabis

leicester/

Fig.43

lugubris

/CUS

d

in

Fig.45

Oy y, Gy <i i ii

Z Ld Wh

ae 5 ‘ 9

——

forosus

vallistris

yusafti

yy \

2: ie NI ) 2 \ \ \\

ES NYAS

Yep 3 \\

3

Yay,

‘Frm

-————. 0.]

adustus

4 Bale.

butleri

ventral

aw, INC TE

S< \ 4 fl Since

NU: AIT sy | eo Cea i

SSS af ; : f 13 : per

Y sete

carment/

Oras ed rT

oa Mayle

PT

ventral

Foose ee Ob

outline

EGG

+———- 0-05 ——“+4

chorion reticulation

dux

vallistris

Fig.93

ol

gibbosus

ventral

A Soe

indicus

UNCUS

2

N

YY]

a,

CAPE ei Y

we " a iN ms We " " ie i ay HH |

om 7 ne a

od Vill-Te o Vili-Te

andamanensis butlers

ap tas <i a or

forosus ¢

Fig.o7

Vill =

ty a hve ] aN

44 OS Ear:

4p a9 : OY, (2.

a ei o : hes 23 - z = me

x. a En as) 2

HN aie 4

Cyc *®

Fig. 58

MALE TARSOMERES 5 AND POSTTARSI

Yo

butleri dux

eae Ae | | ee ee

02——1

embiensis

Va SE Rs fe eye |

a eat ES AEST |

carment/ gibbosus

—

gf CII fo PP C Il

‘ ce per i

clavatus hamistylus

Vz a

Vg a

= ees,

A Sob

-/}—__ 0,2 ————__ indicus

Fig. 59

MALE TARSOMERES 5 AND POSTTARSI

panayensis sentanius

APS

=

| psedomediofasciatus ; 0.2 Singularis p

a a 5,

i eee a

azureosquamatus gts.

Fe Ce eee | eS

ZA. q

eS q

1 EY a) Seen

HII ASG;

butleri eae

(Ga

Q ee oe iy

ee ey

(es a

os a an,

: ‘ Sf ee

carmenti oy BENE Lata vj. SEAR AC Johnson

2S I T

a ee Seba tere

FT rama Sens

lugubris

pseudomediofasciatus

Rien pauline °

neomacrod/xoa Ene) SE Ae virilis A Sone

Fig. 62

guadrispinatus

rees/

sentanius

Vie Gel

7 ET

singularis

ae da

a= cu

FEMALE TARSOMERES 5 AND POST TARSI

CS er HI

uniformis

b> 1.0 —____

butleri

indicus

‘

AS

Ae)

~%

c

®

<

w

Q

wv

7a)

APPENDICES

Reinert: New Interpretation of Vervrallina 177

TABLE 1. Current taxonomic status of the subgenus Verrallina

Species ) oy 2G JG P G E

abditus * «x

adustus > xk 2 >k 6k >

agrestis * x

andamanensis * ek kk > > x

atriisimilis 2k ok 2k *k ek

atrius > > > > > >

azureos quamatus x >

bifoliatus * ek

butleri *k 2k 2k 2k 2 k 2k

campylostylus > kk > ek > 2k

carmenti 2k *k ** >k *K *k ek

cautus * * *k Ok

clavatus > > kk > kk

comatus * kk

COMOSUS >k 2K > >k

consonensis * ek 2k >

cretatus > > > 2k 2k >k Ae

Cuccioi * > > * > > >k 4k > >K

cunninghami x *

* = Stage has been described.

**k = Portion of stage has been figured. Many of the adult 2 and @ illustra-

tions are of the posttarsi only.

? = Not a positive association.

° = Female; °G = Female genitalia; o = Male; “G = Male genitalia; P = Pupa;

L = Larva; E = Egg.

178

Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

TABLE 1 (Continued).

Species 2 c 2G JG P L E

cyrtolabis >k kK * Ok *x *K

aux OK x OK xk eK x *K

embiensis x 2k

foliformis 2k 2k 2k 1k 1k 1k

fragilis x * 2K eK

funereus * * > kk x kk ek

gibbosus * ok 2k ek > kk

hamistylus ** x * 2k x

harvrisonicus * >

hispidus * sheik

incertus > >K kk kK > >K kK kk

indecorabilis * Ok «> *

indicus > x ok x x ek

tviomotensis *k >

johnsoni kk oe ne 4k xk 2k >k

johorensis > ok

killertonis * x 4x me

kompongus 2K ok

lankaensis * x ek kk

latipennis x * ae > 2k >

leicestev1 > >k > 2k k > * x

leilae K kK

lineatus ns 0k xk ok 2k x 4

lugubris * > ek kk sk

macrodixoa *k Ok 3k 1k ok

Reinert: New Interpretation of Vervallina 179

TABLE 1 (Continued).

Species 2 oy 2G JG P ie E

MAY ZAYSEN * OK KK 2k > ae

mccormicki x >k «> ok >k 2k

milnensis * *k 2k x

multifolium * 2k ae 2k * *

neomacrodixoa > kk x x x x

nigrotarsts > > > * x x

nobukonis x x 2k x x x x

notabilis > x

nubicolus * 2k

obsoletus * 2k ee

pahangi * 2k

panayensts * x > ** 2k

parasimilis x KK D «KD *k 6k 2k

petroelephantus Xk **

philippinensts * * x 2k

phnomus * 2k 2k Xk

prioekanensis ok oe

protuberans * ske3k

pseudodiurnus * 2k

pseudomediofasciatus +m > kk * Ok

pseudovarietas > x :

quadrifolium i KK KK > 1k

quadrispinatus kk 2k 2k ok * *

rvamalingamt 2k 4k

180

Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

TABLE 1 (Continued).

Species

yam

YaYUS

reest

robertsi

sabahensis

seculatus

sentanius

similis

simplus

singularis

sohni

spermathecus

stungus

LOYOSUS

trispinatus

uUnCUSs

uniformis

vallistris

vanapus

var iabilis

varietas

virilis

yerburyi

yusaft

2K K

2K K

2K

2K >K

2K

2K 2K

2K K

2K K

2K K

2K K

2K K

2K >K

2K K

2K K

2K K

2K >K

2K >K

2K

2K

KK

2K

2K >K

2K K

2K K

2K

Xk

ok

2K

2K

2K

2K K

2K K

2K K

2K >K

°G

2K

2K

2K

2K K

2K 2K

2K

2K

2K K

2K >K

2K K

2K

2K

2K K

2K >K

2K K

2K

2K

KK

2K K

2K kK

2K

2K

2K K

2K >K

2K

2K K

KK

> kK

2K >K

2K K

2K K

2K K

2K K

2K

2K

2K >K

2K K

2K >

2K >K

2K >K

2K K

2K

2K K

2K K

2K K

> kK

2K K

Reinert: New Interpretation of Vervallina 181

TABLE 2. Record of the branching of the setae on the pupae of

Aedes (Vervallina) adustus (7 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

| Cephalothorax Abdomen I (Cont. ) Abdomen III (Cont. )

1 2-4 2 10 1 1 9 1 1

2 2-3 2 11 1 1 10 3-5 3

3 1-3 2 Abdomen II 11 1 1

4 3-4 3 0 1 1 14 1 1

D 2-4 3 1 16-34 26 Abdomen IV

6 1 1 2 1 1 0 1 1

7 2-4 2 3 1 1 1 3-6 5

8 3-6 3 4 4-8 7 2 1 1

9 1-3 2 5) 3-5 4 3 4-6 6

Metanotal Plate 6 1 1 4 2-4 3

10 6-12 ie! 7 1-2 2 9) 2-4 2

11 1 1 9 1 1 6 1-3 1

12 2-95 4 Abdomen III 7 2-4 3

Abdomen I 0 1 1 8 1-4 2

1 16-25 21 1 5-12 7 9 1 1

2 1 1 2 1 1 10 2-4 3

3 2-4 2 3 1 1 11 1 1

4 5-8 8 4 1-3 2 14 1 1

5 4-8 6 5 4-5 4 Abdomen V

6 1-3 1 6 1-3 2 0 1 1

7 2-6 2 7 3-5 3 1 2-4 2

9 1 1 8 3-6 4 2 1 1

182 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

TABLE 2 (Continued).

Seta Range Mode Seta Range Mode Seta Range Mode

Abdomen V (Cont. ) Abdomen VI (Cont. ) Abdomen VII (Cont. )

3 1-4 2 4 3-6 &, 5) 1-2 i

4. 3-8 6 O 1-3 2 6 3-7 4

5) 1-3 2 6 1 1 7 1 1

6 1 1 i 1 1 8 1-4 3

7 3-7 5) 8 2-4 4. 9 1 1

8 2-9 2 9 1 1 10 1 1

9 1 1 10 i 1 11 | 1 1

10 1 1 11 1-2 1 14 1 1

11 1 i 14 1 1 Abdomen VIII

14 i, 1 Abdomen VII 0 1 1

Abdomen VI 0 1 1 4 2-3 2

0 1 1 1 1-2 1 ) 1-4 2

1 2-5 3 2 1 1 14 1 1

2 1 1 3 2-4 3 Paddle

3 2-4 2 4 1-4 2 1 1 1

Reinert: New Interpretation of Vervallina 183

TABLE 3. Record of the branching of the setae on the pupae of

Aedes (Verrallina) butleri (10 specimens)

Seta Range Mode Seta Range Mode Seta Range Mode

Cephalothorax Abdomen I (Cont. ) Abdomen III (Cont. )

1 2-95 3 10 1 1 7 1 1

2 1-2 2 11 1-2 1 10 3-6 4

3 1-3 2 Abdomen II 11 1 1

4 1-5 4 0 1 1 14 1 1

D 4-7 5 1 17-27 20 Abdomen IV

6 1 1 2 1 1 0 1 1

7 3-7 5 3 1 1 1 4-7 5

8 4-7 4 4 4-8 4 2 1 1

9 2-4 3 D 3-7 6 3 4-8 D

Metanotal Plate 6 1 1 a 2-4 2

10 7-13 7 7 3-6 a 5) 1-2 2

11 1 1 9 1 1 6 2-4 3

12 3-5 é Abdomen III 7 2-4 +

Abdomen I 0 1 1 8 2-4 3

1 14-23 20 1 4-10 8 9 1 1

2 1 1 2 1 1 10 3-5 -

3 3-5 4 3 1 1 11 1 1

4 6-9 6 4. 4-6 4 14 1 1

O 4-9 4 4) 4-7 4 Abdomen V

6 1 1 6 2-4 3 0 1 1

7 1-4 3 7 4-8 5 1 3-5 4

184 Contrib. Amer. Ent. Inst., vol. 11, no. 1, 1974

TABLE 3 (Continued).

Seta Range Mode Seta . Range Mode Seta Range Mode

Abdomen V (Cont. ) Abdomen VI (Cont. ) Abdomen VII (Cont. )

3. 3-5 4 4 4-8 5 5 2-4 4

& 4-7 6 D 2-4 a 6 4-8 &

D 1-3 2 6 2-4 3 7 1-3 2

6 2-3 2 7 1-3 2 8 2-4 4

ny 4-7 6 6. Baas 3 9 1-2 1

8 2-4 3 9 1 1 10 1-3 2

x, 1 i 10 1-2 1 11 1-2 1

10 2-4 2 11 1 1 14 1 1

11 1 1 14 1 1 Abdomen VIII

14 1 1 Abdomen VII 0 1 1

Abdomen VI 0 1 1 4 3-5 4

0 1 1 1 4-6 5 9 1 1

1 4-6 & 2 1 1 14 1 1

Reinert: New Interpretation of Vervallina

185

TABLE 4. Record of the branching of the setae on the pupae of

Aedes (Verrallina) carmenti (9 specimens)

Range Mode

Seta

Cephalothorax

1 2-3 2

2 2-3 2

3 1-2 1

4 3-5 3

5 5-10 6

6 1 1

7 2-7 5

8 6-10 7

9 3-6 3

Metanotal Plate

10

11

12

4-10 7

1 1

4-7 5