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+ Contributions

of the

American Entomological Institute

Volume 9, 1972-1973

CONTENTS

No. 1. Huang, Yiau-Min. Contributions to the mosquito fauna of

Southeast Asia. XIV. The subgenus Stegomyia of Aedes in

Southeast Asia. I. The scutellaris group of species. October

ai, 1972. 109 pages.

. 2. Reinert, John F. Contributions to the mosquito fauna of

Southeast Asia. XV. Genus Aedes Meigen, subgenus Ayurakitia

Thurman. October 27, 1972. 42 pages.

. 3. Zavortink, Thomas J. Mosquito studies (Diptera, Culicidae).

XXIX. A review of the subgenus Kerteszia of Anopheles.

pages 1-054.

Adames, Abdiel J. and Pedro Galindo.

(Diptera, Culicidae). XXX. pages 55-61.

Mosquito studies

May 4, 1973.

. 4. Valencia, Jose D. Mosquito studies (Diptera, Culicidae).

XXXI. A revision of the subgenus Carrollia of Culex. May 12,

1973.

. 9. Reinert, John F. Contributions to the mosquito fauna of

Southeast Asia. XVI. Genus Aedes Meigen, subgenus

Aedimorphus Theobald.in Southeast Asia. May 12, 1973.

This is an irregular-appearing series,

published by the

AMERICAN ENTOMOLOGI CAL INSTITUTE

5950 Warren Road

Ann Arbor, Michigan 48105, USA

Volumes 1-7 available in individual numbers

Volumes 8 and following supplied only as

complete cloth-bound volumes

Contributions

of the

American Entomological Institute

Volume 9, Number 1, 1972

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA. XIV.

THE SUBGENUS STEGOMYIA OF AEDES IN SOUTHEAST ASIA.

I - THE SCUTELLARIS GROUP OF SPECIES.

Yiau- Min Huang

CONTENTS

ENO LION 6 be a as a ee ee e's

THE

AEDES STEGOMYIA) SCUTELLARIS GROUP IN

POO Pe ee ga ey a ee ee eee eee

KEYS TO THE SPECIES DEALT WITH IN THIS REVIEW

Pigee NO Fea ee age ac se a ee SO eR es ee ‘

Male Terminalia . sek. s eg is at Sa ae ae hee

Pee eS a gta oe ee See Roe oe a eee Rie ee eee

Fourth Stace Larvae a 4 i a eS Cr ee

DESCRIPTIONS OF THE SPECIES

albopictus subgr ne

albopictus (Skuse

downs? BONAIY G@ Inev ai ge SE. ws ee nec eee

flavOpicius Vamaae oo. us sw 8 Bee a igre om ee ek

HOowalbOptCIAS Barraue 2. sess se ew hs sen ete ah eae

patriciae MaUinely oka es es ple se ighte eta a Was Re

PSCUARIOODICTIUS LOT), go 5 she cue eG Bek 6 + ue ie. Ge a

seatoi Huang. ... of eee es SOW ae ee oe Ee aS oe

subalbopictus Barraud. kG ee ee Sak Pal ah te eva ea

scutellaris: subgroup

POC ti a kw OS ee ee Se Ae I ek eA

GioFenSiSs HONNG WEPSter 6 ae ees ee we ee ee

andrew si Bawards «oe ee ee ap a ae Gah eran pee ae

ee ee i ee Sa aes ake ws OR Se aoe eS

TESS COC SS i eae 8 ee Ae a a wo

Pauline? Gione © Farner .. 06k ee es Ha ay Cae ce ea eee

Viger Sear © rae el a ee ee

scutellavis (Walker) ...... oe wh dices, ia ae ex Seca eo

BCENOW LIEDGEMENTS slic ee ee ee ee as 6 ee ee ee

TERA TUR CIB cee See eS 8 ewe se eee fe ee es

APPENDIX I. PRESENT STATUS OF THE AEDES (STEGOMYIA)

SCUTELLARIS GROUP OF SPECIES IN SOUTHEAST

Rota be BOR se ek We ee ee eee ae

APPENDIX II. DISTRIBUTION LIST OF THE SOUTHEAST ASIAN

SPECIES OF THE AEDES STEGOMYIA)

SCUTELLARIS GROUP 22.505 6 ees ore Se gle ee

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. XIV.

THE SUBGENUS STEGOMYIA OF AEDES IN SOUTHEAST ASIA!

I - THE SCUTELLARIS GROUP OF SPECIES

Yiau- Min Huang”

INTRODUCTION

The Aedes scutellaris group of species is the most dominant and

complex in the subgenus Sfegomyia as shown by the number of species and

the variety of types. It has not been studied as a unit since Edwards (1932)

and sound revision of the entire group is badly needed.

In Southeast Asia the group is comparatively poorly known, the only

extensive study previously made in the Oriental region being that of Barraud

(1934). In fact, Barraud's work was restricted to India which is the western

part of the Oriental region and is not in the SEAMP area. More recently the

faunae of individual islands have been examined by Bohart & Ingram (1946b)

for Okinawa and by Knight & Hull (1952) for the Philippine Islands.

Inadequate material and insufficient descriptions in the past have led

to confusion and misidentification in this area, as pointed out by Huang (1968).

In order to clarify the situation and in view of the present day interest

in the subgenus, because of its medical importance, it has been decided to

give a detailed description of all species and the present review is the first

of a series which it is hoped will eventually complete the task. Subgeneric

characters and classification of the species groups will be discussed in a final

paper.

The present paper deals with 16 species of the scutellaris group of

which 12 are definitely known to occur in the Southeast Asia area, 1 species

may occur and 3 others, which are unlikely to be found, are treated here for

comparison.

This study has been based primarily on specimens accumulated by the

Southeast Asia Mosquito Project. Additional material was borrowed from the

following institutions: Bernice P. Bishop Museum; United States National

Museum; Field Museum of Natural History; University of Utah; Cornell

University; Johns Hopkins School of Hygiene and Tropical Medicine; California

Academy of Science; Academy of Natural Science, Philadelphia; Medical

Zoology Laboratory, Institute for Infectious Disease, University of Tokyo;

British Museum (Natural History) and the Instituut voor Tropische Hygiene,

Amsterdam.

All the type specimens of the included species which are in the British

Museum (Natural History), the United States National Museum and the Medical

Zoology Laboratory, Institute for Infectious Diseases, University of Tokyo

have been seen and studied by me.

Fi2h id mune

This work was supported by Research Contract No. DA-49-193-MD- 2672

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

of the Surgeon General, Washington, D.C.

2 Southeast Asia Mosquito Project, Department of Entomology, Smithsonian

Institution, Washington, D.C. 20560.

2 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

The nomenclature chosen for the chaetotaxy of the larva and pupa and

the terminology of structural parts of the adult as used in this paper largely

follows that of Belkin (1962) with subsequent modification by SEAMP.

An asterisk (*) following the abbreviations used (¢ = male, ? = female,

P = pupa, L = larva and E = egg) indicates that all or some portion of that

sex or Stage is illustrated. Abbreviations used for the references to the Lit-

erature conform to the World List of Scientific Periodicals, 3rd ed., Aca-

demic Press, 1952.

Distribution records are indicated as follows: Country names are in

capital letters, where known, administrative divisions are in italics and

place names have the first letter capitalized. Place names which could not

be located in the Gazetteers available are spelled according to the labels on

the specimens and are placed within inverted commas. 1 = larval skin, p =

pupal skin, L = whole 4th instar larva. |

The information on the breeding habitats and the distribution presented

in this paper are entirely based on the specimens which have been examined

by me.

All the known stages of the 16 species of the scutellaris group from

Southeast Asia and its adjacent areas are redescribed and illustrated and a

number of previously unknown stages are described for the first time. The

Philippine species previously described as scutellavis (Walker) is rede-

scribed here under a new name and 3 subspecies are elevated to species

rank. Several species are recorded for the first time from particular terri-

tories. Keys to the identification of the species are provided. The informa-

tion on the present status of the scutellaris group in Southeast Asia and its

distribution are Summarized in appendices I and II.

The term "Southeast Asia" as used in this review does not exactly

correspond to any of the world mosquito faunal areas as defined by Belkin

(1962). It comprises the following area: The Ryukyu Islands (Amami,

Okinawa, Miyako, Ishigaki, Iriomote), Taiwan, The Pescadores, Hainan,

China (South of the Yangtze Kiang), The Philippines, North Vietnam, South

Vietnam, Indonesia (the eastern boundary is essentially that of Lee &

Woodhill (1944) as shown in Lee's Atlas of Mosquito Larvae), Cambodia,

Laos, Thailand, Malaysia, The Andaman Islands, The Nicobar Islands,

Burma and Assam. This area falls approximately within 11 degrees South |

to 27 degrees North Latitude and 87 to 130 degrees East Longitude (MAP I).

Bangladesh (East Pakistan) has not been included in this review be-

cause of the total lack of material in all collections.

In order to verify distributional records, all available specimens,

even if from beyond the confines of Southeast Asia, have been examined, as

may be seen in the case of albopictus (MAP II).

THE AEDES STEGOMYIA) SCUTELLARIS GROUP IN SOUTHEAST ASIA

DEFINITION OF THE GROUP. The Southeast Asia scutellaris group

is characterized by the following combination of characters.

MALE. Head. Proboscis dark scaled, with or without pale scales on

ventral side, as long as or longer than fore femur; palpus dark, slightly

shorter to longer than proboscis, with white basal band on each of segments

2-5; those on segments 4-5 incomplete dorsally;segments 4-5 subequal, slen-

der, upturned, and with only a few short hairs; antenna plumose, slightly to

considerably shorter than proboscis; clypeus bare; torus covered with white

scales except on dorsal side; decumbent scales of vertex all broad and flat;

erect forked scales dark, not numerous, restricted to occiput; vertex with

median stripe of broad white scales, with broad dark ones on each Side inter-

rupted by a lateral stripe of broad white scales followed by a patch of white

broad scales ventrally. Thorax. Scutum with narrow dark scales and a

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 3

prominent median stripe of similar white ones; the median stripe usually

narrows Slightly posteriorly and forks at beginning of prescutellar space;

sometimes this median stripe is broken at middle of scutum and is followed

by an inverted Y-shaped marking which forks at beginning of prescutellar

space; there is on each side a posterior dorsocentral line of narrow white to

yellowish scales which does not reach to middle of scutum and an incomplete,

not clearly defined, to complete and well developed supraalar white line; with

or without a few narrow white scales on various areas of scutum; acrostichal

bristles absent; dorsocentral bristles present; scutellum with broad white

scales on all lobes and with a few broad dark ones at apex of mid lobe; ante-

rior pronotum with broad white scales; posterior pronotum with a patch or a

stripe of broad white scales and some dark narrow ones dorsally; paratergite

with broad white scales; postspiracular area with or without scales; subspi-

racular area with or without scales; patches of broad white scales on pro-

pleuron, on upper and lower portions of sternopleuron and on upper and lower

portions of mesepimeron; mesepimeral scale patches connected or separated;

lower mesepimeron without bristles; metameron bare. Wing. With dark

scales on all veins, with or without a minute basal spot of white scales on

costa. Halter. With dark scales.* Legs. Coxae with patches of white

scales; knee-spots present on all femora; fore and mid femora dark, with or

without pale scales scattered anteriorly, paler posteriorly; sometimes mid

femur with a median white line on anterior surface; hind femur anteriorly

with a broad white longitudinal stripe which widens at base and is separated

from apical white scale patch; fore and mid tibiae dark anteriorly, paler pos-

teriorly; hind tibia dark; fore and mid tarsi with basal white bands on tarso-

meres 1-3; hind, tarsus with basal white bands on tarsomeres 1-4, tarso-

‘mere 5 all white, “ or sometimes with a few dark scales at tip on ventral

side; fore and mid legs with tarsal claws unequal, larger one toothed, smal-

ler one simple; hind leg with tarsal claws equal, simple. Abdomen. Abdo-

minal segment I with white scales on laterotergite; tergum I with or without

a median patch of white scales; terga II-VI with lateral white spots only or

with complete or incomplete transverse white basal or sub-basal bands as

well; lateral spots may be connected or not with the dorsal bands; tergum

VII with lateral white spots only; sterna I-VI with basal white bands to all

white; sternum VIII largely to. entirely covered with white scales. Termi-

nalia. Basimere 2 to 3.5 times as long as wide; its scales restricted to

dorsolateral, lateral and ventral areas; with few to a patch of hairs on the

basomesal area of dorsal surface; mesal surface membranous; claspette well

_ developed, with numerous setae and some specialized ones; distimere simple,

elongate, 0.75 to as long as basimere, with a spiniform process at apex to

some distance from the tip and with a few hairs; aedeagus with a distinct

sclerotized lateral toothed plate on each side; paraproct without teeth; cercal

setae absent; ninth tergum in middle rounded, truncated, or produced into a

lobe or a median projection with a hairy lobe on each side.

FEMALE. Essentially as in male, differing in the following respects:

palpus 4-segmented, short, 0.2 of proboscis, with white scales on apical

half or more. Fore and mid legs with tarsal claws equal, simple. Abdomi-

nal segment VIII largely to entirely retracted; sternum VII with conspicuous

rounded lateral lobe; post- genital plate with shallow notch; cerci short and

broad; 3 spermathecae, 1 larger than other 2.

J A. flavopictus, a Palearctic species has pale scales in this position.

2 One species thensilli) which does not occur in Southeast Asia has the

apical half of tarsomere 5 dark.

4. Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

PUPA. Cephalothorax. Trumpet short, 3 to 3.5 times as long as

wide at the middle; hair 1,3-C single, longer than 2-C; 6-C single, stout,

stouter than 7-C; hair 10-C with 2-5 branches, mesad and caudad of 11-C;

11-C single. Abdomen. Hair 1-I well developed, with more than 10 branches,

dendritic; hair 2-I single; hair 3-I single, long; hair 2,3-I not widely sepa-

rated, distance between them as distance between 4,5-I; hair 1-II branched;

hair 2-IV-V mesad of hair 1. Faddle. Margins with fringe; apex roundedor

produced; hair 1-P single.

LARVA. Head, Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; head hair 4-C well developed,branched,

closer to 6-C than 5-C, cephalad and mesad of 6-C; hair 5, 8,9,13-C single;

mentum with 9-14 teeth on each side. Thorax. Hair 2,6-P single;5,7-M

Single; 6-M with 2-4 branches; 9-M with 2-3 branches; 9-T with 2-3

branches. Abdomen. Combof 6-14 scales, in a single row, each scale

with strong or fine denticles or a delicate fringe at base of apical spine;

pentad hair 2-VII distant from 1-VII; 2,4-VII single; siphon short, less

than 2 to 2.5 times as long as wide, acus absent or small; pecten teeth 3-21

in number, evenly spaced, each tooth with 1-4 distinct basal denticles;

1-S with 2-5 branches, inserted beyond last tooth; saddle incomplete or

complete; marginal spicules very small and inconspicuous; 3-X single; ven-

tral brush with 4 pairs of hairs on grid, 4a-X and 4b-X single, 4c-X and 4d-

X with 1-2 branches; without precratal tufts; anal papillae varied.

DISTRIBUTION. The Southeast Asia scutellavis group is mainly con-

fined to the Oriental and Indomalayan areas, with extensions into the southern

art of the Palearctic and the western part of the Papuan area. Aedes

(SStegomyia) albopictus (Skuse) is also known in the Malagasy area, the Bonin

Islands, the Mariana Islands and the Hawaiian Islands.

TAXONOMIC DISCUSSION. The group has been differently interpret-

ed. Edwards (1932) divided the subgenus Stegomyia into four groups which

he designated A, B, C and D. In "Group C (scutellaris group)" he included

10 species from the Oriental and Australasian regions, Crete and Africa.

Knight & Rozeboom (1946) removed A. albolineatus (Theobald) from Group C

and defined a fifth group for it and its relatives and this was designated Group

E (albolineatus group) by Knight & Hurlbut (1949). The latter authors sub-

divided the scutellaris group into 3 subgroups known as Subgroup I. scutellaris

s. Str., Subgroup II. albopictus and Subgroup III. mediopunctatus. Mattingly

(1965) transferred mediopunctatus from Group C to Group B. The term "the

scutellaris group" as used by Farner & Bohart (1945) and others has, in fact,

referred to practically the same complex of species as Knight & Hurlbut's

Subgroup I. The scutellaris group of the present paper comprises both

Knight & Hurlbut's Subgroups I. and II.

At the present, 12 species of the scutellaris group are found within

the Southeast Asia area and 1 additional species may also be present. In ad-

dition, there is one form of scutellaris known but not named from Andaman

Islands (Barraud 1928, 1934). This awaits review when more adequate ma-

terial is available.

The Southeast Asian scutellaris group can be divided further into 2

subgroups, the albopictus subgroup and the scutellaris subgroup. (1) The

albopictus subgroup is characterized by having the supraalar white line not

clearly defined and with only narrow scales over the wing root. It is repre-

sented by 6 species, albopictus (Skuse), downsi Bohart & Ingram, novalbo-

pictus Barraud, patriciae Mattingly, pseudalbopictus (Borel) and seatoi

Huang. Included also in this subgroup is 1 Oriental species, subalbopictus

Barraud from India, which may eventually be found in Southeast Asia. Ithas

been recorded from Hainan Island by Stone, Knight & Starcke (1959), but I

have seen no specimens. In addition, 1 Palearctic species, flavopictus

Yamada from Japan and Korea, which is not found in Southeast Asia is treat-

ed here for comparison. (2) The scutellavis subgroup is characterized by

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia S

having the supraalar white line complete and well developed, with broad flat

scales over the wing root and toward scutellum. It is represented by 6 spe-

cies, alcasidi n. sp., alorensis Bonne-Wepster, andrewsi Edwards, malay-

ensis Colless, paullusi Stone & Farner and viversi Bohart & Ingram. In-

cluded also in this subgroup is 1 western Pacific island species, hensilli

Farner from Ulithi Island, W. Carolines and 1 Papuan species, scutellaris

(Walker) from Aru (Aroe) Islands, Ceram, Ambon Island and New Guinea.

These 2 species are not found in Southeast Asia and are treated here for com-

parison.

Based on the present collection data, all the members of the albopictus

subgroup in Southeast Asia occur in the Oriental area of Belkin (1962). A.

albopictus has the widest distribution throughout the entire Southeast Asia

area and beyond, as shown in MAP II, and pseudalbopictus is also known to

occur in the Indomalayan area (Malaya and Java). All members of the

scutellaris subgroup, except viversi from the Oriental area, are Indomalay-

an in distribution. A. malayensis is also known to occur in the Oriental area

(Thailand, Cambodia, Vietnam and Taiwan) and paullusi extends into the west-

ern fringe of the Papuan area (Ambon I. ). :

The pupae and larvae do not seem to have subgroup characters. The

larvae of albopictus proper are extremely similar to and difficult to separate

from those of alcasidi, malayensis, riversi and scutellaris which are ina

different subgroup. This indicates that the two subgroups are closely related

and should be recognized under one species group, the scutellavis group.

In the identification of the species of the scutellavis group, the adult

stages appear to be more promising than the immature stages. However, it

must be remembered that specific differences between the members of this

group tend to be very slight. Some members are highly variable in both

adult ornamentation and in the immature stages. Although the males of all

Species can be recognized on the basis of morphological features, the females

and the immatures are extremely difficult or impossible to distinguish in

many instances. The male terminalia of all species are distinct and the most

diagnostic feature of all is the claspette of the basimere. In dealing with

these, special preparations must be made and care taken to study both later-

al and mesal views of the dissected claspette as well as undissected views.

It is now difficult and sometimes impossible to say what former work-

ers were dealing with when they called a species 'albopictus". This applies

especially in the case of larvae of the different species which this review

will show are so closely related that misidentification has probably largely

been the rule. The fact that so many species of the group are known to share

the same habitat adds considerably to the problem. Surveys based on the

identification of a single larvae obviously lend themselves to criticism on this

account,

BIOLOGY. The immature stages have been found mainly in tree holes,

bamboo stumps, coconut shells and artificial containers. Some species have

also been found in rock holes and a few have also been found in leaf axils.

Females of 5 species, albopictus, downsi, seatoi, riversi and malayensis

are known to bite man and paullusi has been taken biting buffalo.

MEDICAL IMPORTANCE. The scutellaris group is one of the most

important groups of Stegomyia from the standpoint of transmission of pathogens.

A, albopictus isan important vector of dengue virusin Southeast Asia. It has

been incriminated in the transmission of dengue virus during an outbreak of

hemorrhagic fever in Singapore (Chan et al., 1971) and dengue virus has been

isolated from wild-caught albopictus from Koh Samui, Thailand (Gould et al.,

1968, 1970). A. albopictus from India can transmit chikungunya virus in the

laboratory (Rao et al., 1964) and scutellaris from New Guinea has been in-

criminated as a vector of dengue virus (Mackerras, 1946). Some members

of the scutellaris group are efficient vectors of non-periodic filariasis in the

South Pacific (Belkin, 1962), and 3rd stage larvae of Dirofilaria spp. have

6 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

been found in albopictus in Thailand (Harinasuta et al., 1970). The part

Se by —. in the transmission of pathogens is summarized in Table

I (page 18).

KEYS TO THE SPECIES DEALT WITH IN THIS REVIEW

The scutellaris group of species, including those not so far recorded

from Southeast Asia, can be distinguished from other Stegomyia by the fol-

lowing combination of characters: palpi with white scales; scutum with along

median longitudinal white stripe extending from anterior margin to about level

of wing root; scutellum with broad white scales on all lobes; hind tarsus with

Rime white bands on tarsomeres1-4, tarsomere 5 with basal white band or

all white.

MALES AND FEMALES

1. Supraalar white line incomplete, not clearly

defined and with only narrow scales over

WS OGG fone ea ee ok We eae Sale ee ee eS 2

Supraalar white line complete and well devel-

oped, with broad flat scales over wing

root and:toward scutellum (Fie.:21A).0. osce wee ew a 9

2(1). Scutum with patch of broad flat white scales

on lateral margin just before level of wing

MOO Pigs ako: ceeqp et sere ale | oleae ike w wou ars) DS 3

Sevlune wiloursach pater. Gtaciecw mle hh ees HA er acs ; 4

3(2). Scutum with small white patch of narrow

scales at scutal angle; tergum I witha

large median patch of white scales

(Piges Te Ag de. Ces aa exeredsd eacw esi be wes seatoi Huang

Scutum and tergum I without such patches

ef seales, (Piess1Ay B,D) grar ecw deers albopictus (Skuse)

4(2). Scutum with patch of narrow curved yellow-

ish scales on lateral margin just before

LEVEL OF WIN TOOL i: nh 67 a wae te gery eek ae errs 5)

PRESeBe ales While ie sot a ee EE eee eR ee IS A 8

5(4). Fore and mid femora with some pale scales

scattered on anterior surface ...... novalbopictus Barraud

6(5). Subspiracular area with scales; hind femur

anteriorly with broad white longitudinal stripe

which widens at base and occupies at least

basal 3/4; hind tarsomere 4 with at most

basal 2/3-3/4 white banded (Figs. 20M, O)......... 7

Subspiracular area without scales; white longi-

tudinal stripe confined to basal 3/5 of femur;

hindtarsomere 4 with basal 5/6-9/10 white

banded (Fig. 20N) sie se ele weeks downsi Bohart & Ingram

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 7

7(6). Scutum with patch of narrow curved golden

yellowish scales on lateral margin just

before level of wing root; halter with pale

SEARO Se Sera e: gery eR ate big! gg flavopictus Yamada!

These scutal scales pale yellowish; halter

without Dale Seales od eas, ok AS patriciae Mattingly

8(4). Scutum with patch of broad dark scales on

each side of prescutellar space between

prescutellar white line and postdorso-

central white line; postspiracular area

with scales (Figs. BO Eg Pes . pseudalbopictus (Borel)

Scutum without such patch of scales; post-

spiracular area without scales (Fig.

ZOD IR A ak eg SE Lt cianing subalbopictus Barraud

9(1). Abdomen with lunate lateral white spots

only; wing without a minute basal spot :

of white scales on costa (Fig. 20D) ..... andrewsi Edwards

Abdomen with some complete pale bands

or with indications of such bands on

terga; wing with a minute basal spot of

while: Scales Off COStAs a eo bw eee bee ie? era eee 10

10(9). Mid femur with a median white line on

anterior surface (Fig. 20K) ..... d diame ie wrghie gig ek 11

Mid femur without such line 2 o. sndin. e retelics ieee. 12

11(10). Scutum with a few narrow white scales on

lateral prescutal area and on scutal

angle area (Fig. 20J) .... ‘ paullusi Stone & Farner

Scutum without any narrow white scales

in these WOSTMONS 8G eos ae ee alorensis Bonne-Wepster

12(10). Hind raveomiene Dall whites es mere,” pinta ieee ‘ i3

Hind tarsomere 5 with basal 1/ 2 white 9

(Fis. B10), eee eh ees % gob cael Bi aie an hensilli Farner

13(12). Hind tarsomere 3 with basal 2/5 white;

hind tarsomere 4 with basal 2/3 hilo encals ho-tictiae. . 14

Hind tarsomere 3 with basal 1/2 white;

hind tarsomere 4 with basal 3/4 white

(Fie. ZU) ow. acs tecatiows eee aie eo. ae alcasidi n. sp.

(Pisy Be os al a ee ee .. scutellaris (Walker)

14(13). Hind tarsomere 1 with basal 1/4 white;

hind tarsomere 2 with basal 1/3 white

(Fig. 31D)... fea malayensis Colless

Hind tarsomere 1 with basal 1/ 5 white;

hind tarsomere 2 with basal 1/4 white

(Pie: S18) 5 se Bere eae oe ow &% Hiverst Bohart’ G Ingram

The female of novalbopictus Barraud is unknown.

Ws ee, ks

Palearctic species.

2 Western Pacific species.

Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

MALE TERMINALIA

1. Tergum IX with a median lobe or

BOON. 6 se eC eee ee eee we Se ee eee eS. 2

Tergum IX with middle broadly rounded

One se eS ee ee 8 eR EOS Pee ee 8 8 8

2(1). Tergum IX with conspicuous horn- like

median projection (Fig. 2C)....... .. albopictus (Skuse)

Tergum IX with middle part abe aobse

MILO @AODE owe. be ees Oe ee ee ee EOL So eS aes. 3

3(2). Tergum IX with middle part produced

into large lobe with apex serrated. .......2.2«.sese-. 4

mee OF 00e NOt SErrated «vg wid bi oe a ee 9)

4(3). Basimere 2.5 times as long as wide;

claspette large, apical angle reaching

to 0.8 of basimere; claspette with stem

at base and lateral arms rather slender

in lateral aspect (dissected claspette)

(Pies, 20, CB). ea Re eee eee es downsi Bohart & Ingram

Basimere 3 times as long as wide; clasp-

ette large, fan-shaped, apical angle

reaching to 0.75 of basimere; clasp-

ette with stem at base and lateral arms

widened in lateral aspect (dissected)

Pee Oe a a ee ee ee ees flavopictus Yamada

5(3). Tergum IX with large median lobe; clasp-

ette long, slender, reaching to 0.75 of

basimere, expanded portion facing

WSC OO I GC je a eo 6 RE de REG GF seatoi Huang

Tergum IX with rounded median lobe;

claspette large, broad, reaching to

about 0.5 of basimere, expanded portion

OPUS. A) DOEISION. 6s Fs vo woe 6 # ote aw US ore 6

6(5). Specialized setae on sternal side of expanded

portion of claspette spine-like, curved,

with sharply pointed tips and varying

lemetis (Pies. 9C, ety. St ae novalbopictus Barraud

These setae blade-like or clubbed, without

eliarply pointed tips. 6b vo. 6 « s a ee Taw | it ee 7

7(6). Claspette with broad stem and lateral distal

angle turned through 90° in lateral aspect

(dissected) (Fig. 18C) . we eee... Subalbopictus Barraud

Claspette with narrower stem and without a

90° lateral distal angle (Fig. 11C) .... patriciae Mattingly

8(1). Claspette complex with numerous setae on

expanded distal part, each seta on dis-

tinct cone, a tergal mesal finger-like

process bearing 6 modified setae at

tis Ee Pe) ace a a a 8 ee alorensis Bonne-Wepster

CIASDEUS BiMEIO toi ee ia 8 ee 8 ee « ai gee Bite 9

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia

9(8). Tergum IX with middle truncated;

claspette truncate, with a distinct

oval face at apex, with numerous

setae and several long, stout ones

on tergal side and with 4 spine-like

setae on sternal side of oval face

Cs, se agra! al ae ie reat a aan a dee paullusi Stone & Farner

Tergum IX with middle rounded;

claspette with apex more or less

cylindrica? ues. decd « penal svile i earieetetieieniba

10(9). Claspette long, slender, reaching to

0.7 of basimere, with 1 widened

specialized spine-like seta and nu-

merous setae distal to it (Fig. 183C). . pseudalbopictus

Claspette rather short, with several

widened specialized setae and nu-

merous setae. .... iM A wie, ee Baie Ae SOAS

11(10). Claspette with distal expanded part

square in lateral aspect (dissected),

sternal and tergal sides more or

less parallel, apicosternal angle

PYOSCNE he shies os le le BO ee é ere helh aichzeeie whe sows TBE:

Claspette with distal expanded part sub-

triangular in shape in lateral aspect

(dissected), sternal and tergal sides

not parallel but tapering, without apico-

stern@lsaneie «ar wccwl, eacikiwi ae «08S albbeg hs erate.

12(11). Claspette with 4-5 modified setae in a

row on apicosternal angle, with several

distinctly long and stout setae in apico-

tereal area (ie. 250) 60d 6 ae we oe andrewsi Edwards

Claspette with 5-6 modified setae set ona pro-

minence close to apicosternal angle area,

without any distinctly long and stout setae in

apicotergal area(Figs. 35C, 25A, B) ... scutellaris (Walker)

13(11). Apicotergal area of claspette with several

distinct] y-lome Setae), «igiszis- @suivieceid slivce sere SRee AES

Apicotergal area without any such setae ........2e.e. P

14(13). Claspette with 6-7 modified setae in a row

at center of sternal side and occupying

about 1/ 3:08 464 iges : 2265-260) sacs ts artercnice alcasidi n. sp.

Claspette usually with 7 modified setae,

basal one often rather smaller, ina

row, ‘set on a Slight prominence at

center of sternal side and occupying

about 2/5-of ib (Pies..88C 7:2, Hibbs oisixie do. hensilli Farner

15(13). Claspette with 7-10 modified setae forming

a prominant row at center of sternal

side and occupying about 1/2 of it

LF ices: 26y deed wor fis Che le aw ee malayensis Colless

10 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

Claspette with 6-8 modified setae at center

of sternal side, closer to sternal angle

area than to apicotergal angle area and

are about 2/5 of it (Figs. 26A, B,

32C

pi oocugl Gay weds Su Than. Sereda: pam Rega eee vyiveryst Bohart & Ingram

PUPAE

1. Hair 9-II-V strongly developed, thickened,

much stouter than 9-II (Figs. 16A, B) ......

Hair 9-III-V not strongly developed, slender,

about Same macnitude ac OrIl nei we So. a

2(1). Hair 9-VI much stouter than 9-V, at least

twice aS long as 9-V....... A ea

Hair 9-VI about as thick as 9-V, less than

IMBGS 26 JONG Ores tates “ad ett eet Poe 8

3(2). Hair 9-VI usually single and barbed; 9-VII

usually single and barbed or with 2

branches at tip; 9-VIII usually with 2

main stems (1-2) reaching beyond paddle

fringe, each with lateral branches of

Oo — 2O= Oy, JO. ~ 8 78

varying length (Figs. 4A, B) ..... downsi Bohart & Ingram

Hair 9-VI single and simple; 9- VI single

and barbed; 9- VII usually with 2 main

stems, barbed, not reaching beyond

fringe of paddle (Figs. 18A, B) .... subalbopictus Barraud

4(2). Hair 9-VII single, stout and barbed or

SPIE ALU © psx igo hoe ets eNO a i ane Sa aa Te ta ha ae oe 5)

Hair.2-V il Single, seiimpie. Fee Se a es ee Na a ta ke ae 8

5(4).. Hain 6-O auch shorter dian tO ee ee ST eee 6

HainG6-C abow. ae. lone Ga te a i a ee Sek ees 7

6(5). Paddle margins with rather short fringe;

hair 9-VIII single, strong, barbed

CES ORG? EB). «gutta gata ed oe ea ee Ce novalbopictus Barraud

Paddle margins with long fringe; hair

9-VIII usually with 2 main stems (1-

2), each barbed, reaching beyond

fringe of paddle (Figs. 6A, B)........ flavopictus Yamada

7(5). Hair 6-C much stouter than 7-C and

usually slightly longer; hair 1-II usu-

ally with 10-11 branches rising from

a common stem at base; 9- VIII usu-

ally with 2 main stems, reaching be-

yond fringe of paddle, each with lateral

branches of varying length; paddle mar-

gins fringed close to base, on more

than apical 3/4 of paddle (Figs.

C9 Ag Sele taeda) eke’ hie Bee ae paullusi Stone & Farner

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 11

Hair 6-C usually about as long as 7-C;

hair 1-II branched, without a common

stem at base; paddle margins with

fringe on less than apical 3/4 of pad-

dle. (Figs, CAA ccd ion ints a.aiow'ven by yp) UCESIAL ISP fin part)

(Figs. 35A, B). « «0-0 «+. «, Seutellaris (Walker). (in part)

8(4). Hair 9- VII usually with 2 (1-2) branches,

each barbed, not reaching beyond fringe

of paddle (Figs. LIA) Bo ee ; patriciae ae

Hair 9-VIII reaching beyond fringe of paddle . ia aire ch Sones 9

9(8). Hair 6-C; aboutl/2.as TOneA8 THO cscdi-< fear lw bar einen elie etneeie: + 10

Hair 6-C about 3/4 the length of 7-C to

aboul as Jose 26 foG- es rela fe weal Ks Sehnert aie Ga Beret 11

10(9). Hair 9- VIII with 2 branches (Figs. 13A, B). pseudalbopictus (Borel)

Hair 9-VII usually single (1-2) (Figs.

DA. BY ala bald Balms ie te ee « Bol GinObestye se ise)

11(9). Hair 6-C about 3/4 the length of 7-C........ elt ie 12

Hair 6-C about as long as 7-C

Wiggs aP2QAs - BY) gc%. 6 dite Shi te eae? ei Aw ee ot SOLCESIGE Me Bp, ia sp

Figs. SMG ds) we t< <8 ~~... sceutellaris (Walker) (in part

12(11). Hair 9-VIII usually with single main

stem or divided into 2 and lateral

branches of varying length; hair L-TT

with many primary and secondary

branches (Figs. 27A, B). . ‘

Hair 9-VIII usually with 2 branches 1 3)"

barbed; hair 1-II with very few secon-

dary branches (Figs. 32A, B). . viversi Bohart & Ingram

vite malayensis Colless

FOURTH STAGE LARVAE

1. Comb scale with prominent denticles at

base of apical spine; hair 2-VIL usu-

ally with 6 (5-8) branches (Pig. 2 (isis. aialecenarn, veearot Huang

Comb scale without such denticles........ 6:46 fe sisi es 2

2(1). Siphon acus present; pecten teeth 3-6,

each tooth short and stout; usually

with 3-4 basal denticles (Fig. 14). . . pseudalbopictus (Borel)

Siphon acus absent; at least 8 pecten

teeth, each tooth long, at least 4

times: as: long aS Wide: 44 lcniietPone agada BO ws es pe & 3

3(2). Saddle complete. ......... pphiPea + imiwh Gakcrecaee7e ke teats 4

Saddle incompletes 6 iin his wigan te Gay an wee eee esaaten fag, 6

4(3). Hair 14-P with 5-7 branches; hair 2-VII

with 4-5 branches; comb scale and

pecten tooth rather narrow and slender,

with sharply pointed tips (Fig. 7) ...... flavopictus Yamada

Hair 14-P with 2-3 branches; hair 2-VII

with 2-3 branches; comb scale and

pecten tooth rather broad and stout. ......... ce agul 3)

12 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

9(4). Hair 2-X 2-branched; pecten teeth each

usually with 3 (2-3) basal denticles;

1-S inserted beyond last tooth and

yentrad of teéth (Fig. 10). Oo

Hair 2-X usually single, when 2-branched

one much smaller than the other;

pecten tooth usually with 2 (2-3) basal

denticles; 1-S inserted well beyond last

tooth and in line with teeth (Fig. 19)

6(3). Hair 2-X 3-branched; hair 2-VII usu-

ally with 3 (2-3) branches; pecten tooth

with 1 large and occasionally 1-2

very small basal denticles (Fig. 30) ..

7(6). Hair 1-VII usually with 4 (3-4) branches,

short, less than twice as longas5-VIL...

Hair 1-VII usually with 2 (2-3) branches,

long, at least 2.5 times as long aso-VII..

8(7). Hair 2-VII with 3-4 branches ........ ‘

Hair 2-VII usually single (1-2) (Fig. 3)

9(8). Comb scale with free portion widened at

base and sharply pointed at tip

novalbopictus Barraud

subalbopictus Barraud

paullusi Stone & Farner

Bate Fo Br ae te a ee ee ee ee 7

POE ae 10

aa te ae 9

ca lbopictus (Skuse)

Pig are eer as age ee be ta downsi Bohart & Ingram

Comb scale with free portion rather

slender, nearly parallel-sided from

base and at least as long as attached

portion (Pie. 22 ey es oe eS

10(7). Hair 1-VIL with 2-3 branches; siphon about

twice as long as wide; pecten teeth 10-21

in number, closely arranged in a line;

1-S usually inserted beyond middle of

siphon; comb scale sometimes with

patriciae Mattingly

apical spine split at tip (Fig. 33) ... viversi Bohart & Ingram

Hair 1-VII usually with 2 long branches

(2-3), when 3-branched then one much

smaller than other two; siphon about 2.5

times as long as wide; pecten teeth 10-

16 in number; 1-S inserted at middle

or before middle of siphon. ........

11(10). Hair 1-S usually inserted at middle of

siphon; pecten teeth 10-14 in number,

each with 2-4 basal denticles (Fig.

PN gale pa ge ie ia cak ae Oe a Sak eae

Hair 1-S usually inserted before middle

of siphon ae BO Sh oy ee aes beta te

Pie 0 a gla a ee Ore a mo

aera ee e's 11

malayensis Colless

alcasidi n. sp.

scutellaris (Walker)

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 13

AEDES STEGOMYIA) ALBOPICTUS (SKUSE)

(Fig. 1, &; 2, o terminalia, pupa; 3, larva)

Culex albopictus Skuse 1894, Indian Mus. Notes 3(5):20 (9).

Stegomyia scutellaris (Walker), Theobald 1901, Mon. Cul. 1:298 (<*, 2*);

Leicester 1908, Cul. Malaya 3(3):86 (%, 2) (misidentifications).

ee Hs Ages samarensis Ludlow 1903, J. N.Y. ent. Soc. 11:138

J, 2)

Stegomyia lamberti Ventrillon 1904, Paris Mus. Bull. 10:552 (¢, 9).

Stegomyia nigritia Ludlow 1910, Canad. Ent. 42:194 (¢).

Stegomyia quasinigritia Ludlow 1911, Psyche 18:129 (¢).

Aedes (Stegomyia) albopictus (Skuse), Edwards 1917, Bull. ent. Res. 7:209

(synonymized samarensis); Dyar & Shannon 1925, Insec. Inscit.

menst. 13:74 (synonymized nigrvitia and quasinigritia); Barraud 1931,

Indian J. med. Res. 19:222 (o«*); Edwards 1932, Genera Insect. , Fasc.

194:164 (synonymized lamberti); Bonne-Wepster & Brug 1932, Geneesk.

Tijdschr. v. Ned.-Ind. 2:73 (o*, L*); Barraud 1934, Fauna Brit.

India 5:233 (o*, 9, L*); Bohart & Ingram 1946, U.S. Navmed. 1055:5,

35, 64 (oc *, 9*, L*); LaCasse & Yamaguti 1950, Mosq. Fauna Japan

and Korea :111 (o*, 2*, P*, L*); Knight & Hull 1952, Pacif. Sci. 6(2):

176 («*, 2, L); Bonne-Wepster 1954, Spec. Publ. R. trop. Inst.

Amsterdam 111:81 (o, ?*, L*); Bohart 1956, Insects Micronesia 12(1):

57 (o*, 9, L); Hara 1957, Jap. J. exp. Med. 27:65 (2*); Belkin 1962,

Mosq. South Pacific 1:456 (o*, 9, P*, L*); Huang 1968, Proc. ent. Soc.

Wash. 70(4):298 (o*, 2*, P*; Neotype designated).

MALE. Head. Proboscis dark scaled, as long as fore femur; palpus

dark, longer than proboscis, with white basal band on each of segments 2-5;

those on segments 4-5 incomplete dorsally; segments 4-5 subequal, slender,

upturned, and with only a few short hairs; antenna plumose, slightly shorter

than proboscis. Thorax. Scutum with narrow dark scales and a prominent

median stripe of similar white ones, which narrows slightly posteriorly and

forks at beginning of prescutellar space; on each side a posterior dorso-

central white line which does not reach to middle of scutum; a patch of broad

flat white scales on lateral margin just before level of wing root and few nar-

row curved white scales over wing root; posterior pronotum with a large

patch of broad white scales and some dark narrow ones dorsally; postspirac-

ular area without scales; subspiracular area with white scales; mesepimeral

scale patches connected forming a V-shaped white scale patch, the open end

of 'V' directed backwards. Wing. With dark scales on all veins except for

minute basal spot of white scales on costa; first forked cell 1.5 times as long

as its stem. Halter. With dark scales. Legs. Fore and mid femora dark

anteriorly, paler posteriorly; hind femur anteriorly with a broad white stripe

which widens at base andis narrowly separated from apical white scale patch;

fore and mid tarsi with basal white bands ontarsomeres 1-2; hind tarsus with

basal white bands on tarsomeres 1-4, the ratio of length of white band to total

length of tarsomeres 1-4 is 1/3, 2/5, 1/2 and 2/3, tarsomere 5 all white, or

sometimes with a few dark scales at tip on ventral side. Abdomen. Abdomi-

nal segment I with white scales on laterotergite; terga II-VI each with a bas-

al white band which widens laterally and with lateral white spots which do not

connect with the basal bands; terga II and VIL with lateral white spots only, or

sometimes tergum II also with a median white spot; sterna II-III largely cov-

ered with white scales; IV-VI each with a basal white band; sternum VII large-

ly covered with white scales. Terminalia. Basimere relatively short and

broad, twice as long as wide; with a patch of hairs on basomesal area of dor-

sal surface; mesal surface extensively membranous; claspette large, mush-

room-like, with numerous setae and with several widened specialized setae

14 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

on meSsal side and a few widened specialized curved ones on apical angle of

expanded distal part; distimere simple, elongate, apex somewhat swollen

and with some hairs; with a spiniform process at apex; tergum IX with con-

Spicuous horn-like median projection and with a hairy lobe on each side.

FEMALE. Essentially as in male, differing in the following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Abdominal

pale basal bands present on terga II-VI; segment VII large retracted, ster-

num VII with conspicuous rounded lateral lobe; post- genital plate with shal-

low notch; cerci short and broad; 3 spermathecae, 1 larger than other 2.

PUPA. Cephalothorax. Trumpet short, 3 times as long as width at

middle; both hair 1,3-C single, longer than 2-C; 2-C usually single (1-2);

4-C usually double (1-2); 5-C usually 3- branched (2-3); 6-C single, stout,

Shorter than 7-C; 7-C usually single (1-2); 10-C usually with 2-3 branches;

mesad and caudad of 11-C; 11-C single. Abdomen. Hair 1-I well developed,

with more than 10 branches, dendritic; 2-I single; 3-I single, long; 2, 3-I not

widely separated, distance between them as distance between 4, 5-I; hair 1-II

usually with 4-8 branches; 2-II laterad of 3-II; hair 2-IV, V mesad of hair 1-

IV,V; hair 1-I usually with 2-3 branches (2-5); hair 3-II, III single, shorter

than segment II; hair 5-IV-VI single, or sometimes 5-IV, V double, not

reaching beyond posterior margin of following segment; hair 9-I-VI small,

Single, simple; 9- VII, VII stouter than preceding ones; hair 9-VII single,

simple; hair 9- VIII usually single (1-2), barbed, reaching beyond fringe of

paddle. Paddle. Margins with fringe; hair 1-P single; 2-P sometimes pres-

ent.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A in-

serted near middle of shaft, single; inner mouth brushes pectinate at tip; head

hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad and

mesad of 6-C; hair 5, 8,.9,13-C single; 6-C single or double; 10-C usually

single (1-2); 7-C usually with 2-3 branches; 11-C usually 3-branched (3-4);

12-C usually double; 14-C usually with 3-4 branches; 15-C usually double (2-

3); mentum with 10-12 teeth on each side. Thorax. Hair 1-P with 3-4

branches; 2-P single; 3-P double; 4-P usually 3-branched (2-3); 5-P usually

single, rarely double; 6-P single; 7-P usually double (2-3); 9-P single; 11-P

usually double (1-2); 14-P 3-branched; 5, 7-M single; 6-M usually 3-branched

(3-4); 8-M with 4-5 branches; 9-M usually double, rarely 3-branched; 10, 12-

M single, long stout; 11-M single, small; 7-T usually with 4-5 branches; 9-T

double; 10,11-T similar to those on mesothorax; 12-T much reduced. Abdo-

men. Hair 6-I usually 3-branched (2-4); 7-I single; 6-II usually 3- branched

(2-3); 7-II usually 3-branched (2-3); 6-III-V double; 6-VI single; 1-VII usually

4-branched (3-4); 2-VIL usually single (1-2); comb of 8-12 scales, rarely 6,

in a single row, each scale with fine denticles or fringes at base of apical

Spine; pentad hair 2,4-VII single; 1-VII with 3-5 branches; 3-VIII with 5-7

branches; 5-VII with 3-4 branches; siphon short, about twice as long as wide,

acus absent; pecten teeth 8-14 in number, evenly spaced, each tooth with 2

main basal denticles; 1-S with 2-4 branches, inserted beyond last tooth and

in line with teeth; saddle incomplete; marginal spicules very small and incon-

Spicuous; 1-X 2-branched; 2-X 2-branched, rarely single; 3-X single; ventral

brush with 4 pairs of hairs on grid, each hair single; no precratal tufts; anal

papillae about 3 times as long as saddle, sausage-like.

TYPE DATA. Culex albopictus Skuse, type females, non-existent;

type locality: Calcutta, INDIA; Aedes Stegomyia) albopictus (Skuse), Neo-

type male (No. 1-14-104) with associated pupal skin and terminalia slide

(68/1054), Neo-allotype female (No. 1-14-15) (designated by Y.M. Huang,

1968) in U.S. National Museum, Washington, D.C.; type locality: Botanical

Garden, Calcutta, Bengal, INDIA, 3-VI-1967 (S. Ramalingam, E.D. Abraham

& E.S. Abraham). 7

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia 15

DISTRIBUTION. 12,300 specimens examined: 3, 2750, 4, 2849, 576

4 eh 4? terminalia, 182 L; 2,364 individual rearings (1, 615 1,

, 364 p).

PHILIPPINES. Luzon; Leyte; Mindoro; Samar; Mindanao; Jolo;

Culion; Busuanga; Palawan; Negros; 1,015, 1,47392, 55 & terminalia, 655

individual rearings (465 1, 655 p).

RYUKYU ISLANDS. Iriomote; Ishigaki; Miyako; Okinawa; Amami;

107c, 1819, 8 o terminalia, 184 individual rearings (184 1, 184 p).

TAIWAN. Taipei; Hsin-Chu; Ping-tung; Orchid Island; 9°, 169, 1 ¢

terminalia, 14 individual rearings (141, 14 p). — |

HONG KONG. Hong Kong; New Territories: Taipokau; Kowloon; 19,

249, 2 terminalia.

CHINA. Kwangtun: Kwangtung; Canton; Fukien: Fukien; Fuchow;

Chekiang: Hangchow; Kiangsu: Shanghai; Nanking; Hopeh: Peking; Hainan; 6¢,

169, 1 o terminalia.

VIET NAM. Con Son; Kontum; Gia Dinh: Gia Dinh; Tan San Nhut;

Saigon; Tay Ninh; Bien Hoa; Long Khanh: Gia Ray; Binh Duong: Lai Khe; Vinh

Long; Khanh Hoa: Duc My; Nha Trang; Quang Tri: An Khe; Da Nang: Danang;

Spanish Point; Thua Thien: Phu Bai; Quang Nam; Quang Duc: Dak Song; Darlac:

Ban Me Thuot; Tuyen Duc: Fyan; Da Lat: Dalat; 'NLong Binh; Chu Lai;"' 76¢,

2019, 140 terminalia, 13 individual rearings (13 1, 12 p). .

LAOS. Champassak: Sedone; Pakse; Vientiane; ''Ban Van Heue;" llc,

112.

CAMBODIA. Kandal: Phnom-Penh; Oudong; Kompong Speu; Kampot:

Sihanoukville; Takeo: Prey Phdau; ''Ari-Ksatr; Ari-Gsatr; Chrin Chang Phnom

Penh;'' 17c, 189.

THAILAND. Nakhon Si Thammarat; Khon Kaen; Nan; Phangnga;

Lampang; Prachinburi; Ranong; Nonthaburi; Chon Buri; Surat Thani; Ko Samui;

Trat; Prachuap Kiri Khan; Phuket; Phra Nakhon; Nakhonsawan; Ang Thong;

Maehong Son; Chiang Mai; Nakhon; Nayok; Nakhon Ratchasima; Loei; Lampoon;

Nakhonayor; Chumphon; Trang; Kanchanaburi; Songkhla; Chanthaburi; Rayong;

Tak; Yala; 8840, 9369, 330c¢ terminalia; 29 terminalia, 136 L, 902 individual

rearings (563 1, 902 p).

BURMA. Pegu Division: Rangoon; Shan State: Aung Ban; 350, 439, 3c

terminalia, 50 individual rearings (41, 50 p).

MALAYSIA. West Malaysia: Perak; Perlis; Kedah; Selangor; Johore;

Trengganu; Pahang; Kelantan; 640¢, 8179, 280 terminalia, 192 terminalia, 46

L, 402 individual rearings (285 1, 402 p). East Malaysia: Sabah- Kalabakan;

Sandakan; Tawau; Sawah Tuaran; Berbuloh Barat Labuan; Kota Belud; Saban

Semporna; Lipasli papar; Penampang Jesselton; Keningan; Limbuak Banggi;

Lingkabau; 250°, 259. Savawak- Kuching; Pang Kalan Tebang; 6c, 89.

SINGAPORE. 820, 879, 1c terminalia, 86 individual rearings (42 1,

86 p).

INDONESIA. Sumatra: Atjer; Benkoelen; Padang Panzang; Fort de

Kock; Singkarak; Deli; Tjoorab; Kaban Djahe; Nias. Java: Maos; Boueloeng;

Tjandfoer; Tjisarua; Tjilatjap; Malang; Batavia; Kapetakan; Bogor; Bataria;

Buitenzorg; Soerabna. Kalimantan: Kandangan; Bandjermasin; Tavakan.

Celebes: Kalawara; Bintaoena; Mamaedfre; Molino; Makassar; Kabaena.

Lesser Sunda Islands: Bali; Flores; Alor. Timor; Ceram; 123¢, 1489, a7

terminalia.

INDIA. Bengal: Calcutta; Darjeeling- Sukna; Jalpaiguri- Old Jalpai-

guri; Bihar; Patna- Bihar City; Purnea; Darbhanga- Pusa; Madras: Nilgiri-

Coonoor; Nilgiri Hills; Delhi; Kanara; Karwar; Canar: N. Canar; Central

Province; Sambalpur; Assam: Assam; Lakhimpur- Dibrugarh; Doom Dooma;

Sibsagar- Jorhat; '"Chabua; Misamari;"' 1490, 1699, 21¢ terminalia, 19

terminalia, 59 individual rearings (45 1, 59 p).

W. Pakistan. Lahore; 2, 59.

CEYLON. Colombo; Central Province: Peradeniya; 'Suduganga;

Diya'wa; Pundaluoya;"' 10c, 149.

16 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

NEPAL. Hetaura, 2,000 ft; 3c.

JAPAN. Honshu: Tokyo; Yokohama; Kyoto; Kiushiu: Nagasaki; 9c,

10°, 1c terminalia.

BONIN ISLANDS. 49.

HAWAIIAN ISLANDS. Oahu: Mokapu Point; Hawaii: Kona Coast; 42,

84c% terminalia.

CHAGOS ISLANDS. Pevros Banhos; Diamont Island; 19°.

SEYCHELLES. Mahe; Praslin; Platte; Capucin; Silhouette; Victoria;

Dennis; 37%, 609.

LA REUNION. 20, 39.

MAURITIUS. 1c, 59.

MADAGASCAR, 3c, 192.

MARIANA ISLANDS. Guam; Saipan; 4c, 49.

REMARKS. A great number of specimens of this species have been

examined and as it has been collected from many places and administrative

divisions in some countries, such as Thailand, Philippines, Ryukyu Islands

and Malaya, detailed locality records are therefore not presented here but

are available at SEAMP. :

TAXONOMIC DISCUSSION. A. albopictus is one of the commonest

species in Southeast Asia. The adult can easily be distinguished from all

other members of the scutellavis group in this area by having the scutum

with a patch of broad flat white scales on the lateral margin just before the

level of the wing root, no small white patch on the scutal angle area, and ab-

dominal tergal white bandings basal and not connected with the lateral white

spots. Tergum IX of the male terminalia has a conspicuous horn- like median

projection, thus differing from all other species that have been described in

this group.

A. albopictus is a member of the albopictus subgroup, having the

supraalar white line not clearly defined, with only narrow scales over the

wing root, and the abdominal tergal markings basal and not connected with

the lateral markings; it can thus easily be distinguished from all other mem-

bers of the scutellaris subgroup. However, the immature stages of this spe-

cies are extremely similar to those of alcasidi, malayensis, riversi and

scutellaris which belong to the scutellavris subgroup. In Southeast Asia, the

larvae of albopictus are often found in association with those of alcasidi,

malayensis and riversi in the field. Thus great care must be taken in identi-

fying them. The larva of albopictus can be distinguished from alcasidi,

malayensis and viversi by having abdominal hair 1-VII usually 4-, sometimes

3-branched, and always much shorter and stronger; in the 3 other species

hair 1-VI usually with 2 (2-3) long branches. The pupa of albopictus having

hair 6-C about 1/2 of 7-C, can also be distinguished from the 3 other species

which have hair 6-C usually much stouter than 7-C and at least 3/4 of 7-C to

about as long as 7-C.

BIOLOGY. The immature stages of albopictus have been found mainly

in tree holes, bamboo stumps and artificial containers in Philippines, Ryukyu

Islands, Taiwan, Viet Nam, Thailand, Malaysia, Burma and India. They

have also been found in coconut shells in the Philippines, Thailand and Malay-

sia, in rock holes in the Philippines and Thailand, in palm fronds in Malaysia

and in fallen abaca leaf and leaf axils in the Philippines. The immature stages

have been collected associated with malayensis in Singapore, Malaya, Thai-

land and Taiwan, with alcasidi in the Philippines, with viversi in Ryukyu

Islands, with pseudalbopictus in Thailand, Malaya, Burma and India, with

seatoi in Thailand, with downsi in Ryukyu Islands and with subalbopictus in

India.

Feng (1933) found the parasite Lankesteria culicis in one out of six

adult albopictus dissected in Woosung, China. According to his report the

pathogenicity of L. culicis to mosquitoes is that in light infection it does very

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia A7

little harm to the host but if the infection is a heavy one the larvae or pupae

frequently fail to hatch and die. Certainly this deserves further examination

and it may well find a place in biological control.

Gubler (1970a) found that males of albopictus mate readily with fe-

males of polynesiensis Marks and that they are strong competitors of poly-

nesiensis males for these females. A cage colony of polynesiensis was era-

dicated by the addition of albopictus males, at a ratio of 10 albopictus males

to 1 polynesiensis. The same author (1970b) reports that the competitive

displacement principle does apply to albopictus and polynesiensis under labo-

ratory conditions and suggested that a field trial should be considered.

Earlier reports on hybridization experiments between Aedes aegypti

(Linnaeus) and A. albopictus (Skuse) are conflicting. However, Leahy &

Craig (1967) found that the potential for hybridization between aegypti and

albopictus is extremely low and that at least five barriers act in sequence to

isolate these species, namely, (1). Mating behavior; (2) Structural incompa-

tibility; (3) Sperm inactivation; (4) Reduced oviposition and (5) Genetic in-

compatibility. These barriers make successful hybridization in the field

highly improbable and earlier reports of success may have been due to con-

tamination.

MEDICAL IMPORTANCE. A. albopictus is one of the most important

species from the standpoint of the transmission of pathogens. Table I (page

18) shows briefly what is known in this regard.

AEDES (SSTEGOMYIA) DOWNSI BOHART & INGRAM

(Figs. 4, o terminalia, pupa; 5, larva; 8B, claspette; 20N, hind leg)

Aedes (Stegomyia) downsi Bohart & Ingram 1946, J. Wash. Acad. Sci. 36(2):

51 (o*, 9, L*); Bohart & Ingram 1946, U.S. Navmed 1055:64 (c, 2, L).

Aedes (SStegomyia) flavopictus downsi Bohart & Ingram, Bohart 1953, Proc.

ent. Soc. Wash. 55:184 (to ssp. status).

MALE. Head, Proboscis dark scaled, without any pale scales on

ventral side, longer than fore femur; palpus dark, as long as proboscis, with

white basal band on each of segments 2-5; bands on segments 4-5 incomplete

dorsally; segments 4-5 subequal, slender, upturned, and with only a few

short hairs; antenna plumose, shorter than proboscis. Thorax. Scutum with

narrow dark scales and a prominent median stripe of similar white ones, me-

dian stripe narrows slightly posteriorly and forks at beginning of prescutellar

space; prescutellar line with some pale yellowish scales; on each side a pos-

terior dorsocentral pale yellowish line which does not reach to middle of scu-

tum; posterior scutal lines usually rather dull and indistinct; a patch of narrow

curved yellowish scales on lateral margin just before level of wing root and a

few narrow curved pale yellowish scales over wing root; posterior pronotum

with a patch of broad white scales and some dark narrow ones dorsally; post-

Spiracular area without scales; subspiracular area without scales; mesepime-

ral scale patches connected in the form of a 'V', the open end of 'V' directed

backwards. Wing. With dark scales on all veins except for minute basal spot

of white scales on costa; first forked cell 1.5 times as long as its stem.

Halter. With dark scales. Legs. Fore and mid femora dark anteriorly,

paler posteriorly; hind femur anteriorly with a broad white stripe which widens

at base and on about basal 3/5; fore and mid tarsi with basal white bands on

tarsomeres 1-2; hind tarsus with basal white bands on tarsomeres 1-4, ratio

of length of white band to total length of tarsomeres 1-4 is 1/5, 1/4, 1/3-2/5,

5/6-9/10, tarsomere 5 all white or sometimes with a few dark scales at tipon

ventral side. Abdomen. Abdominal segment I with white scales on latero-

tergite; terga IV-VI each with a basal white band and lateral white spots which

18 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

TABLE I. VERTEBRATE PATHOGENS ASSOCIATED WITH AEDES

(SSTEGOMYIA) ALBOPICTUS: (SKUSE

(RT RR Ea nS SE ey tC SE RSA RES SS a A SR A SAE

en a

zmmitis France laboratory |can be infected 1938

Divofilaria rom wild- |srd stage larvae

Spp. Taide ae eee eee al. (1970

gts ak ebonbeeeattts ducks Wettony (1

lophurae U.S.A. laboratory |mit to ducks. Jeffery (1944

Plasmodium in can be infected & trans-iRussell &

gallinaceum india __ laboratory |mit to fowls

pir" us. hasten et eae Ee

allax U.S.A. laboratory |mit to pigeons

Dengue

hee certs in can transm1l Philip

U.S.A. laboratory {hamster to hamster Smadel (1943

Tamiinad Strain [india laboratory [chick to chick [Varma (1960).

Tamilnad Strain (|India laboratory |chick to chick Varma (1960

virus [india laboratory fmouse to mouse (964)

virus India laboratory |mouse to mouse 1964 :

alitis virus U.S.A. _jlaboratory |chick to chick _ 1965 te

type- 2 Thailand icaught isolation of virus 1968, 1970) |

Dengue virus rom wild- |jvirus recovered during |Russell et al.

ic fever

type-2 [Singapore leaught’ [isolation of virus (Chan (1965) _

type- 2 Singapore |caught isolation of virus Chan (1965

Dengue virus rom wild- |jvirus recovered during (Chan et al.

type-1, 2 Singapore |caught epidemic hemorrhagic |(1971)

fever

* These authors reported successful transmission with Aedes scutellaris

but it seems certain that they were working with albopictus.

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 19

do not connect with basal bands; terga II, III, VII with lateral white spots

only; or sometimes tergum III also with basal white band; abdominal basal

bands usually rather weak or incomplete; sterna II-VI with basal white bands;

sternum VII largely covered with white scales. Terminalia. Basimere 2.5

times as long as wide; with patch of hairs on basomesal area of dorsal sur-

face; mesal surface membranous; claspette large, somewhat fan-shaped, api-

cal angle reaching to 0.8 of basimere, with numerous setae and several

widened specialized ones on mesal side of expanded distal part; distimere

simple, elongate, as long as basimere; with a spiniform process and a few

hairs at apex; tergum IX with middle part produced into a large lobe, apical

margin of lobe sharply serrate and with a hairy lobe on each side.

FEMALE. Essentially as in male, differing in the following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Abdominal

basal bands on terga II-VII, or as in male; segment VIII largely retracted.

PUPA. Cephalothorax. Trumpet about 3 times as long as width at

middle; hair 1,3-C single; longer than 2-C; 2-C single; 4-C usually double;

5-C usually double (2-3); 6-C single, stout, shorter than 7-C; 7-C usually

single (1-2); 10-C usually with 2-3 branches, mesad and caudad of 11-C;

11-C single. Abdomen. Hair 1-I well developed, with more than 10 branches,

dendritic; 2-I single; 3-I single, long; 2,3-I not widely separated, distance

between them as distance between 4, 5-I; hair 1-II usually with 8-9 branches

(3-12); 2-II laterad of 3-II; hair 2-IV, V mesad of hair 1-IV, V; hair 1-III usu-

ally 3-branched (2-6); hair 3-II, III single, shorter than segment II]; hair 5-

IV-VI single, or hair 5-IV, V double, not reaching beyond posterior margin

of following segment; hair 9-I-V small, single, simple; 9- VI, VIL stouter than

preceding ones; hair 9-VI single and barbed; 9-VII usually single, barbed, or

with 2 branches at tip; 9-VIII usually with 2 main stems (1-2), each with lat-

eral branches of varying length, reaching beyond fringe of paddle. Faddle.

Margins with fringe; hair 1-P single.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; inner mouth brushes pectinate at tip;

head hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad

and mesad of 6-C; 5, 8,9,13-C single; 6-C single or double; 10-C single or

double; 7-C with 2-3 branches; 11-C with 2-4 branches; 12,15-C with 2-3

branches; 14-C with 3-5 branches; mentum with 13-14 teeth on each side.

Thorax. Hair 1-P 3-branched; 2-P single; 3-P double; 4-P 3-branched; 5,

6-P single; 7-P double; 9-P usually single (1-2); 11-P usually single (1-2);

14-P with 3-4 branches; 5, 7-M single; 6-M with 3-4 branches; 8-M with4-5

branches; 9-M with 2-3 branches; 10,12-M single, long, stout; 11-M single,

small; 7-T with 4-6 branches; 9-T usually double (2-3); 10,11-T similar to

those on mesothorax; 12-T much reduced. Abdomen. Hair 6-I 3-branched;

7-I single; hair 6-II with 2-3 branches; 7-II 3-branched; 6-III-V double; 6-VI

single; 1-VII 4-branched; 2-VII 3-branched; comb of 8-12 scales in a single

row, each scale with fine denticles at base of apical spine; pentad hair 2, 4-

VIII single; 1-VII with 4-5 branches; 3-VIII with 5-6 branches; 5-VII with

4-6 branches; siphon short, about twice as long as wide, acus absent; pecten

teeth 10-14 in number, evenly spaced, each tooth with 3 (1-3) basal denticles;

1-S with 4-5 branches, inserted beyond last tooth and in line with teeth; saddle

incomplete; marginal spicules very small and inconspicuous; 1-X 2-branched

(2-3); 2-X 2-branched; 3-X single; ventral brush with 4 pairs of hairs on grid,

each hair single; no precratal tufts; anal papillae about 1.5 times as long as

saddle, dorsal pair longer than ventral pair.

TYPE DATA. Aedes (Stegomyia) downsi Bohart & Ingram, holotype

~ male in U.S. National Museum, Washington, D.C.; type locality: Chizuka,

Okinawa (RYUKYU-RETTO), IX-1945 (R. Bohart & R. Ingram). Paratypes:

3 males, 4 females, with same data as holotype; 2 males, 2 females, Chizuka,

Okinawa, VIII-1945; 5 males, 13 females, Shana Wan, Okinawa, IX-X-1945;

2 males, Heddo, Okinawa, IX-1945; 1 male, Hentona, Okinawa, IX-1945

20 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

in U.S. National Museum; 2 males, 1 female, Shana Wan, Okinawa, IX-1945

in British Museum (Natural History). Paratype larvae, all from Okinawa:

8 larvae on 7 slides, Shana Wan, [X-1945; 1 larva on slide, Shana Wan, 29-

IX-1945, ex tree hole (this larva appears to be albopictus); 2 larvae on 1 slide,

Hedo, IX-1945 in U.S. National Museum. All specimens collected by Bohart

& Ingram.

DISTRIBUTION. 118 specimens examined: 210, 439, 16 terminalia,

10 L, 14 individual rearings (14 1, 14 p).

RYUKYU ISLANDS. Okinawa: Chizuka; 80, 89, 6c terminalia;

Shana Wan; 8c, 159, 50 terminalia, 8 L; Heddo; 2c, 2¢ terminalia, 2 L;

Hentona; 1c, 1c terminalia (VIII-X-1945, all collected by Bohart & Ingram).

Iriomote: (25-V-1968, A.B. Silagan), 19, 1 individual rearing (1 1, 1 p);

Yabu village (XII-1968, A.B. Silagan), 59, 5 individual rearings (5 1, 5 p).

Yaeyama: Inoto (7-X-1968, G. Takaesu), 59, 5 individual rearings (5 1, 5 p);

Ishigaki city, Shiraho village (I-1970, I.V. Villanueva), 1¢, 29, 1o¢ termi-

nalia, 3 individual rearings (3 1, 3 p). Ishigaki-Jima: (X-XI-1961, Sasa),

1c, 59, 1o terminalia. Amami: (V-1962, Sasa), 29.

TAXONOMIC DISCUSSION. A. downsiis a member of the albopictus

subgroup. The adult differs from albopictus, seatoi, pseudalbopictus, sub-

albopictus and novalbopictus by having the scutum with a patch of narrow,

curved, yellowish scales on lateral margin just before level of wing root, and

fore and mid femora without pale scales scattered on anterior surface. It is

very similar to flavopictus and patriciae, but can easily be distinguished from

both by having subspiracular area without scales, hind femur anteriorly witha

white stripe on basal 3/5 and hind tarsomere 4 with basal 5/6- 9/10 white band-

ed; in flavopictus and patriciae the subspiracular area bears scales, hind fe-

mur anteriorly has a white stripe on at least 3/4 andhindtarsomere 4 has basal

2/3-3/4 white banded at most. The maleterminalia of this species have center —

of tergum IX produced into a large lobe, and apical margin of lobe serrate, and

can thus easily be distinguished from all other species except flavofictus. The

similarity between these two forms is so close that one would be inclined to

regard downsi as a subspecies of flavopictus. However, it can be separated

from flavopictus by the diagnostic characters mentioned in the key. In addi-

tion, the immature stages of downsi are markedly differentiated from flavo-

pictus. The larva of downsi has the saddle incomplete, hair 14-P with 3-4

branches, hair 11-M and 11-T usually single (1-2); in flavopictus the saddle

is complete, hair 14-P has 5-7 branches, hair 11-M is double and 11-T

double or 3-branched. The pupa of downsi has hair 9-VI much stouter than

9-V, at least twice as long as 9-V, hair 9-VI usually single and barbed; in

flavopictus hair 9-VI about same magnitude as 9-V, always single and simple.

Based on the morphological difference in all stages of these two forms, I be-

lieve that downsi should be recognized as a distinct species.

The larva of downsi is very similar to patriciae but can be separated

by having a comb Scale with the free portion widened at base and sharply

pointed at tip; in patriciae the free portion of a scale is rather slender and

nearly parallel- sided from base. The pupa of downs? is very similar to

subalbopictus in having hair 9-VI much stouter than 9-V, but can be separated

from it by having 9-VI usually single and barbed, hair 9-VIII usually with 2

main stems (1-2), each with lateral branches of varying length and reaching

beyond fringe of paddle. In subalbopictus hair 9-VI is single and simple,

hair 9-VIII usually with 2 main stems, barbed and not reaching beyond fringe

of paddle,

A. downsi is apparently restricted to the Ryukyu Islands. The imma-

ture stages are often found in association with albopictus in the field. Great

care must therefore be taken in identifying them. The larva of downsi can

be distinguished from albopictus by having hair 2-VII with 3-4 branches,

whereas in albopictus this hair is usually single (1-2). The pupa of downsi

is easily distinguished from albopictus by having hair 9-VI much stouter than

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 21

9-V, usually single and barbed, hair 9-VII usually single and barbed or with

2 branches at tip, 9-VII usually with 2 main stems (1-2), each with lateral

branches of varying length, reaching beyond fringe of paddle; in albopictus

hair 9-VI is about the same magnitude as 9-V, always single and simple; 9-

VII is single and simple, 9-VIII usually single (1-2), barbed and reaching be-

yond fringe of paddle.

BIOLOGY. The larvae of downsi have been found mainly in taro leaf

axils and in tree holes; once in cut bamboo and once in banana axil in Okinawa.

The immature stages from Yaeyama and Iriomote were found mainly in tree

holes and once in an artificial container. The specimens from Ishigaki- Jima

were found in taro leaf axils and the specimens from Amami in bamboo stumps.

The adult females have been taken biting in Okinawa. The immature stages

were associated with albopictus and riversi.

AEDES STEGOMYIA) FLAVOPICTUS YAMADA

(Figs. 6, o& terminalia, pupa; 7, larva; 8A, claspette; 200, hind leg)

Aedes Stegomyia) flavopictus Yamada 1921, Annot. zool. jap. 10:52 (c*, 2);

LaCasse & Yamaguti 1950, Mosq. Fauna Japan and Korea :116 (c*,

o*, P*, L*); Sasa & Kano 1951, Jap. J. exp. Med. 21:112 (L*);

Asanuma & Nakagawa 1953, Misc. Rep. Res. Inst. nat. Resourc.

Tokyo No. 31:87 (P*); Hara 1957, Jap. J. exp. Med. 27:65 (?*).

MALE. Head. Proboscis dark scaled, without any pale scales on

ventral side, slightly longer than fore femur; palpus dark, slightly longer

than proboscis, with white basal band on each of segments 2-5; those on seg-

ments 4-5 incomplete dorsally; segments 4-5 subequal, slender, upturned,

and with only a few short hairs; antenna plumose, shorter than proboscis.

Thorax. Scutum with narrow dark scales and a prominent median stripe of

Similar white ones, stripe narrows slightly posteriorly and forks at beginning

of prescutellar space; prescutellar line with some pale yellowish scales; on

each side a posterior dorsocentral pale yellowish line which does not reach to

middle of scutum; a patch of narrow curved golden yellowish scales on later-

al margin just before level of wing root and a few narrow curved yellowish

scales over wing root; sometimes a few narrow pale yellowish scales on scu-

tal angle area; scutellum with broad white scales on all lobes, without, or

sometimes with 1-2 broad dark ones at apex of mid lobe; posterior pronotum

with large patch of broad white scales and some dark narrow ones dorsally;

postspiracular area without scales; subspiracular area with pale scales; mes-

epimeral scale patches connected forming a V-shaped white- scaled patch, the

open end of 'V' directed backwards. Wing. Dark scales on all veins except

for minute basal spot of white scales on costa; first forked cell 1.8 times as

long as its stem. Halter. With pale scales. Legs. Fore and mid femora

dark anteriorly, paler posteriorly; hind femur anteriorly with a broad white

stripe which widens at base and on about basal 3/4; fore and mid tarsi with

basal white bands on tarsomeres 1-2; hind tarsus with basal white bands on

tarsomeres 1-4, the ratio of length of white band to total length of each tarso-

mere is 1/4, 1/3, 2/5-1/2, 2/3, tarsomere 5 all white except tip dark on

ventral side. Abdomen. Abdominal segment I with white scales on latero-

tergite; terga III- VI each with a basal white band and lateral white spots which

do not connect with basal band; terga II and VIL with lateral white spots only,

or sometimes tergum II with median white spot; sterna III- VI with basal white

bands; sternum VIII largely covered with white scales. Terminalia. Basi-

mere 3 times as long as wide; with a patch of hairs on basomesal area of

dorsal surface; mesal surface membranous; claspette large, fan-shaped, the

apical angle reaching to 0.75 of basimere, with numerous setae and several

22 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

widened specialized ones on mesal side of expanded distal part; distimere

simple, elongate, as long as basimere, with a spiniform process and a few

hairs at apex; tergum IX with center produced into a large lobe, apical mar-

gin of lobe serrate and with a hairy lobe on each side.

FEMALE. Essentially as in male, differing in the following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Abdominal

basal bands on terga IN-VU; segment VIII completely retracted.

PUPA. Cephalothorax. Trumpet about 4 times as long as width at

middle; hair 1,3-C single; 3-C much longer than 2-C; 2-C single; 4-C usually

single (1-2); 5-C usually double (2-5); 6-C single, stout, shorter than 7-C;

7-C usually single (1-2); 10-C usually with 2-3 branches, mesad and caudad

of 11-C; 11-C single. Abdomen. Hair 1-I well developed, with more than10

branches, dendritic; 2-I single; 3-I single, long; 2, 3-I not widely separated,

distance between them as distance between 4, 5-I; hair 1-II with 8-10 branches;

hair 2-II laterad of 3-II; hair 2-IV,V mesad of 1-IV,V; hair 1-III usually with

3-4 branches; hair 3-II, III single, shorter than segment III; hair 5-1V-VI

Single, not reaching beyond posterior margin of following segment; hair 9-I-

VI small, single, simple; 9-VII, VII stouter than preceding ones; hair 9- VI

usually single and barbed; hair 9- VII usually with 2 main stems (1-2), each

barbed, reaching beyond fringe of paddle. Paddle. Margins with fringe;

hair 1-P single.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A in-

serted near middle of shaft, single; inner mouth brushes pectinate at tip; head

hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad and mes-

ad of 6-C; hair 5, 8,9, 13-C single; 6-C single or double; 10-C usually single

(1-2); 7-C 3-branched; 11-C with 3-4 branches; 12-C double; 14-C with 4-6

branches; 15-C double; mentum with 10-11 teeth on each side. Thorax. Hair

1-P 4-branched; 2-P single; 3-P 3-branched; 4-P 4-branched; 5, 6-P single;

7-P double; 9-P double; 11-P usually double (2-3); 14-P with 5-7 branches;

5, 7-M single; 6-M 4-branched; 8-M 4-branched; 9-M with 2-3 branches; 10,

12-M single, long, stout; 11-M double, small; 7-T with 4-5 branches; 9-T

usually double; 10,11-T similar to those on mesothorax, or sometimes 11-T

3-branched; 12-T much reduced. Abdomen. Hair 6-I with 3-4 branches; 7-I

single; hair 6-II-V double; 7-II 3-branched; 6-VI single; 1, 2-VII with 4-5

branches each; comb of 8-10 scales in a single row, each scale with fine den-

ticles at base of apical spine; pentad hair 2,4-VIII single; 1-VII with 7-8

branches; 3-VIII with 6-8 branches; 5-VII with 8-9 branches; siphon short,

about twice as long as wide, acus absent; pecten teeth 10-14 in number, even-

ly spaced, each tooth usually with 2 (2-3) basal denticles; 1-S with 3-5 branches,

inserted beyond last tooth and in line with teeth; saddle complete; marginal

spicules very small and inconspicuous; 1-X with 2-4 branches; 2-X 2-branched;

3-X single; ventral brush with 4 pairs of hairs on grid, each hair single ex-

cept the 2 proximal ones double; 4d-X much shorter and smaller than others;

no precratal tufts; anal papillae about 3 times as long as saddle, sausage-

like.

TYPE DATA. Aedes Stegomyia) flavopictus Yamada, lectotype male

ea by Y.M. Huang, 1969), lectotype male terminalia mounted on slide

YMH- '69-80) in Medical Zoology Laboratory, Institute for Infectious Dis-

eases, University of Tokyo, Tokyo, Japan; type locality: Shiba, Tokyo, JAPAN,

20-IV-1916 (S. Yamada). Syntypes: 2 males, 1 female, with same data as

lectotype; 1 female, Inage, near Tokyo, 17-V-1916 (S. Yamada); 1 female,

Shiba, Tokyo, 14-V-1921 (S. Yamada) (misidentification, it is albopictus) in

Medical Zoology Laboratory, Institute for Infectious Diseases, University of

Tokyo, Tokyo.

DISTRIBUTION. 59 specimens examined: 200, 209, 11c terminalia,

4 individual rearings (41, 4p).

JAPAN. Honshu: Tokyo- (12-I1X-1915, S. Yamada), 19; Shiba (20-

IV-1916, S. Yamada), 3c, 19, 1c terminalia; Inage (17-V-1916, S. Yamada),

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 23

19; Chiba (VII-1949, 406 MGL), 10, 49, 1o¢ terminalia; Kyoto- (IX-X-1947,

207 MSD), 1c, 19, 1¢ terminalia; (VII-1949, 207 MSD), 2c, 19, 1o termi-

nalia; Yodo (VIII-1947, 207 MSD), 19; Nagaoka (24-VII-1953, Y. Shogaki & J.

McClendon), 1c, 1c terminalia; Saitama, Sagiyama (VI-1957), 3c, 19;

Shizuoka, Misakubo (VIII-1969, M. Sawada & A. Yoshii), 2°, 29, 1o termi-

nalia, 4 individual rearings (41, 4 p); Hokkaido: Rubeshibe (29-VII-1917, S.

Yamada), 2c, 29, 1c terminalia. )

KOREA. (K.W. Lee), 5c, 59, 40 terminalia.

TAXONOMIC DISCUSSION. A. flavopictus is a Palearctic species

of the albopictus subgroup. The adult is very similar to downsi and patriciae

in having the scutum with a patch of narrow, curved, yellowish scales on lat-

eral margin just before level of wing root and fore and mid femora without

some pale scales scattered on anterior surface. It is closer to patriciae than

to downsi in having the subspiracular area with scales, hind femur anteriorly

with a white stripe on at least basal 3/4 and hind tarsomere 4 with basal 2/3-

3/4 white banded at most. It can be distinguished from patriciae by having

the scutum with a patch of narrow, curved, golden yellowish scales on lateral

margin just before level of wing root, halter with pale scales; in patriciae

the scutal scales are pale yellowish and the halter lacks pale scales.

The male terminalia of flavopictus are extremely similar to downsi but

differing in having the basimere 3 times as long as wide; claspette large, fan-

shaped, apical angle reaching to 0.75 of basimere; claspette with stem wid-

ened at base and both arms rather broad and stout in lateral aspect (dissected

claspette); in downsi basimere rather short and broad, 2.5 times as long as

wide; claspette large, apical angle reaching to0.8of basimere, claspette with

stem and both arms rather slender in lateral aspect (dissected claspette).

The larva of flavopictus is very similar to novalbopictus and sub-

albopictus in having the saddle complete. It is easily distinguished from both

by having hair 14-P with 5-7 branches, hair 2-VI with 4-5 branches, comb

scale and pecten tooth rather narrow and slender with sharply pointed tips;

in novalbopictus and subalbopictus hair 14-P with 2-3 branches, hair 2-VI

with 2-3 branches, comb scale and pecten tooth rather broad and stout. The

pupa of flavopictus is very similar to novalbopictus but differs in having the

paddle margins with long fringe, hair 9-VIII usually with 2 main stems (1-2),

each barbed, reaching beyond fringe of paddle; in movalbopictus paddle mar-

gins with rather short fringe, hair 9-VIII single, strong and barbed.

A. flavopictus is restricted to the Palearctic Region and does not oc-

cur in Southeast Asia. It is known from Japan and Korea. In Japan, where

both albopictus and flavopictus are present, there is little doubt that the im-

mature stages of flavopictus will be found in association with albopictus in

the field. Thus, great care must be taken in identifying them. The larva of

flavopictus can easily be distinguished from albopictus by having hair 14-P

with 5-7 branches, hair 2-VII with 4-5 branches, hair 1,5-VIII with 7-9

branches; in albopictus hair 14-P 3-branched, hair 2-VII usually single (1-2)

and hair 1,5-VIII with 3-5 branches. The pupa of flavopictus can easily be

distinguished from albopictus by having hair 9-VII single and barbed, 9-VIII

usually with 2 main stems (1-2), each barbed, reaching beyond fringe of pad-

dle; in albopictus hair 9-VII single and simple, 9-VIII usually single (1-2),

barbed and reaching beyond fringe of paddle.

BIOLOGY. The immature stages of flavopictus have been found main-

ly in bamboo stumps, once in a rubber tire and once in a cement tank in

Japan.

24 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

AEDES STEGOMYIA) NOVALBOPICTUS BARRAUD

(Figs. 8C,D, claspette, o tergum IX; 9, o terminalia, pupa;

10, larva; 20B, thorax)

Aedes Stegomyia) novalbopictus Barraud 1931, Indian J. med. Res. 19:224

_ (&*); Barraud 1934, Fauna Brit. India 5:237 (o*, L*).

MALE. Head. Proboscis dark scaled, with some pale scales on

ventral side, slightly longer than fore femur; palpus dark, as long as pro-

boscis, with white basal band on each of segments 2-5; those on segments 4-

5 incomplete dorsally; segments 4-5 subequal, slender, upturned, and with

only a few short hairs; antenna plumose, shorter than proboscis. Thorax.

Scutum with narrow dark scales and a prominent median stripe of similar

white ones; stripe rather narrow, forking at beginning of prescutellar space;

on each side a posterior dorsocentral pale line which does not reachto mid-

dle of scutum; a patch of narrow curved pale yellowish scales on lateral mar-

gin just before level of wing root and a few narrow curved pale scales over

wing root; sometimes a few narrow pale scales on anterior prescutal area;

posterior pronotum with a large patch of broad white scales and some dark

narrow ones dorsally; postspiracular area without scales; subspiracular area

with pale scales; upper sternopleural scale patch does not reach to anterior

corner of sternopleuron; mesepimeral scale patches connected forming a V-

shaped white patch, the open end of 'V' directed backwards. Wing. Dark

scales on all veins except for minute basal spot of white scales on costa; first

forked cell 1.7 times as long as its stem. Halter. With dark scales. Legs.

Fore and mid femora dark with some pale scales scattered anteriorly, more

so on mid than on fore femur, paler posteriorly; hind femur anteriorly with

a broad white stripe which widens at base and is separated from apical white

scale patch; fore and mid tarsi with basal white bands on tarsomeres 1- 2;

hind tarsus with basal white bands on tarsomeres 1-4, the ratio of length of

white band to total length of tarsomere is 1/5, 1/4, 1/3 and 1/2, tarsomere

5 all white. Abdomen. Abdominal segment I with white scales on lateroter-

ite; terga II-VI with lateral spots; terga I and I each with median white spot

holotype male) or not; terga II-VI each with basal white band; lateral spots

not connected with basal bands; tergum VII with lateral white spots only;

sterna I-VI with all white scales. Terminalia. Basimere 2.5 times as long

as wide; with few (1-3) hairs on basomesal area of dorsal surface; mesal sur-

face membranous; claspette with broad stem, with numerous setae and sever-

al widened specialized curved ones on sternal side of expanded distal part;

these widened specialized curved setae with sharply pointed tips and vary-

ing in length; distimere simple, elongate,as long as basimere; with a spini-

form process and a few hairs at apex; tergum IX with center produced into

a rounded lobe and with a hairy lobe on each side.

FEMALE. Unknown.

PUPA. Cephalothorax. Trumpet short, 3 times as long as width at

middle; hair 1,3-C single, longer than 2-C; 2-C single; 4-C double; 5-C 3-

branched; 6-C single, stout, shorter than 7-C; 7-C double; 10-C 3-branched,

mesad and caudad of 11-C; 11-C single. Abdomen. Hair 1-I well developed,

with more than 10 branches, dendritic; 2-I single; 3-I single, long; 2, 3-I not

widely separated, distance between them as distance between 4, 5-I; hair 1-II

usually with 12 branches (12-13); 2-II laterad of 3-II; hair 2-IV, V mesad of

hair 1-IV, V; hair 1-III with 2-3 branches; hair 3-II, III single, shorter than

segment III; hair 5-IV-VI single, not reaching beyond posterior margin of fol-

lowing segment; hair 9-I-VI small, single, simple; 9-VII, VIII stouter than

preceding ones; hair 9-VII stout, single and barbed or split at tip; hair 9-VHl

with strong main stem and lateral branches of varying length. Paddle. Mar-

gins with rather short fringe; hair 1-P single. |

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 25

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; head hair 4-C well developed, branched,

closer to 6-C than 5-C, cephalad and mesad of 6-C; hair 5,6, 8, 9,10, 13-C

Single; 7,12,14,15-C double; mentum with 12-13 teeth on each side. Thorax.

Hair 1-P 3-branched; 2-P single; 3-P double; 4-P double; 5,6-P single; 7-P

double; 9,11-P single; 14-P 3-branched; 5, 7-M single; 6-M 3-branched; 8-M

4-branched; 9-M double; 10,12-M single, long, stout; 11-M single, small;

7-T with 4-5 branches; 9-T double; 10,11-T similar to those on mesothorax;

12-T much reduced. Abdomen. Hair 6-I double; 7-I single; 6-II double; 7-II

3-branched; 6-III single; 6-IV, V double; 6-VI single; 1-VII 4-branched; 2-VII

double; comb of 8-10 scales in a single row, each scale with fine denticles

or fringe at base of apical spine; pentad hair 2,4-VIII single; 1- VIII 3- branched;

3-VII with 6-7 branches; 5-VIII with 4-5 branches; siphon short, about twice

as long as wide, acus absent: pecten teeth 13-14 in number, evenly spaced,

each tooth usually with 3 (2-3) basal denticles; 1-S 4-branched, inserted be-

yond last tooth and ventrad of teeth; saddle complete; marginal spicules very

small and inconspicuous; 1, 2-X 2-branched; 3-X single; ventral brush with 4

pairs of hairs on grid, each hair single; no precratal tufts; anal papillae

about 3 times as long as saddle, sausage-like.

TYPE DATA. Aedes Stegomyia) novalbopictus Barraud, type male

(Y 534) in British Museum (Natural History), London; type locality: Pusa,

Bihar, INDIA, 27-VII-1916 (S.K.S.).

DISTRIBUTION. 20 specimens examined: 9c, 7c terminalia, 2 in-

dividual rearings (2 1,

INDIA. Bihar: Darbhanga, Pusa- (27-VII-1916, S.K.S.), 1¢, 1¢

terminalia; (15-IX-1921, Shaffi), 1o; (11-1931, Barraud), 3c, 3¢ terminalia;

(6- VO- 1921), 14; Madras: Nilgiri, Coonoor, Nilgiri Hills (29-V-1969, B.N.

Mohan), 2c, 2¢ terminalia, 2 individual rearings (21, 2p).

THAILAND. Chiang Mai: Huey Keo (4-II-1953, D.C. Thurman Jr.)

1h, 1o terminalia.

y DEMARIS. There are 3 larval and pupal skin slides (no. 59/mn

107/35 ; 119/59) in B.M., all marked India, Koti, Kasauli area, 1-183. No

anadetited adults of these slides were found and I have not included this mate-

rial in the larval and pupal descriptions.

TAXONOMIC DISCUSSION. The adult of novalbopictus is very similar

to patriciae, downsi and flavopictus in having scutum with a patch of narrow,

curved, yellowish scales on lateral margin just before level of wing root. It

can easily be distinguished from them in having fore and mid femora with

some pale scales scattered on anterior surface; in patriciae, downsi and

flavopictus fore and mid femora lack such pale scales. The male terminalia

of this species are very Similar to patriciae and subalbopictus in having mid-

dle part of tergum IX produced into a rounded lobe, claspette large and broad,

reaching to about 0.5 of basimere, expanded portion of claspette horizontal

in position. It is easily distinguished from both by having the specialized

setae on the sternal side of expanded portion of claspette spine-like, curved,

with sharply pointed tips and of varying lengths; in patriciae and subalbopictus

these setae are blade-like or clubbed, without sharply pointed tips.

The larva of novalbopictus is very similar to subalbopictus but differs

in having hair 2-X 2-branched, pecten tooth usually with 3 (2-3) basal denti-

cles, 1-S inserted just beyond last tooth and ventrad of teeth; in subalbopictus

hair 2-X usually single, sometimes 2-branched with one much smaller than

the other, pecten tooth usually with 2 (2-3) basal denticles, 1-S inserted well

beyond last tooth and in line with teeth. The pupa of novalbopictus is very

similar to flavopictus but differs in having paddle margins with rather short

fringe, hair 9- VII single, strong and barbed; in flavopictus paddle margins

with long fringe, hair 9- VIII usually with 2 main stems (1-2), each barbed,

reaching beyond fringe of paddle.

26 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

A, novalbopictus, anOriental species of the albopictus subgroup, is pre-

sently known from India and northern part of Thailand, In Southeast Asia, it is

known from only Chiang Mai, Thailand and is recordedhere for the first time

from this country.

BIOLOGY. The immature stages of novalbopictus have been found in

tree holes and bamboo internodes in India,

AEDES STEGOMYIA) PATRICIAE MATTINGLY

(Figs. 11, * terminalia, pupa; 12, larva; 20M, hind leg)

Aedes Skate aa flavopictus Yamada, Barraud 1931, Indian J. med, Res,

9:224 (o*); Barraud 1934, Fauna Brit. India 5:239 (o*, 9, L*)

eae)

Aedes ie patriciae Mattingly 1954, Ann, trop. Med. Parasit, 48:

262 (, 9, P*, L).

MALE. Head. Proboscis dark scaled, with a few pale scales on

ventral side, slightly longer than fore femur; palpus dark, as long as pro-

boscis, with white basal band on each of segments 2-5; those on segments

4-5 incomplete dorsally; segments 4-5 subequal, slender, upturned, and

with only a few short hairs; antenna plumose, shorter than proboscis. Tho-

vax. Scutum with narrow dark scales and a prominent median stripe of sim-

ilar white ones; stripe narrowed slightly posteriorly and forked at beginning

of prescutellar’ Space; on each side a posterior dorsocentral pale line which

does not reach to middle of Sscutum; a patch of narrow curved pale yellowish

scales on lateral margin just before level of wing root and a few narrow

curved pale yellowish scales over wing root; sometimes a few narrow pale

scales on anterior prescutal area and on scutal angle area; posterior prono-

tum with large patch of broad white scales and some dark narrow ones dor-

sally; postspiracular area without scales; subspiracular area with pale scales;

upper sternopleural scale patch does not reach to anterior corner of sterno-

pleuron; mesepimeral scale patches connected forming a V-shaped white

patch, the open end of 'V' directed backwards. Wing. Dark scales on all

veins except for minute basal spot of white scales on costa; first forked cell

twice as long as its stem. Halter. With dark scales. Legs. Fore and mid

femora dark anteriorly, paler posteriorly; hind femur anteriorly with broad

white stripe which widens at base and is separated from apical white scale

patch; fore and mid tarsi with basal white bands on tarsomeres 1-2; hind tar-

sus with basal white bands on tarsomeres 1- 4, the ratio of length of white

band to total length of tarsomere is 1/4, 1/3, 2/5-1/2 and 2/3-3/4, tarso-

mere 5 all white. Abdomen. Abdominal segment I with white scales on lat-

erotergite; terga II-VI with lateral spots; tergum I with (holotype male) or

without madian white spot; tergum II with (holotype male) or without basal

white band; terga II-VI each with a basal white band; the lateral spots do not

connect with basal bands; tergum VI with lateral white spots only; sterna I0-

VI with basal white bands; sternum VIII largely covered with white scales.

Terminalia. Basimere 3 times as long as wide; with a patch of hairs on baso-

mesal area of dorsal surface; mesal surface membranous; claspette large,

reaching to 0.5 of basimere, with numerous Setae and several widened spe-

cialized clubbed ones on sternal side of expanded distal part; distimere sim-

ple, elongate, 0.8 as long as basimere, with a spiniform process and a few

hairs at apex; tergum IX with center produced into a rounded lobe and with a

hairy lobe on each side.

FEMALE. Essentially as in male, differing inthe following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Abdominal

basal bands on terga II- VU; segment VIII largely retracted.

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia 27

PUPA. Cephalothorvax. Trumpet short, 3 times as long as width at

middle; hair 1,3-C single, longer than 2-C; 2-C single; 4-C usually single

(1-2); 5-C usually 3-branched (2-3); 6-C single, stout, shorter than 7-C; 7-C

usually single (1-2); 10-C usually 3-4 branched, mesad and caudad of 11-C;

11-C single. Abdomen. Hair 1-I well developed, with more than 10 branches,

dendritic; 2-I single; 3-I single, long; 2,3-I not widely separated, distance

between them as distance between 4,5-I; hair 1-II usually with 5-7 branches

(4-9); 2-II laterad of 3-II; hair 2-IV, V mesad of 1-IV, V; hair 1-II usually

with 1-2 branches, rarely with 3; hair 3-II,II. single, shorter than segment

III; hair 5-IV-VI single, short, not reaching beyond posterior margin of fol-

lowing segment; 9-I-VI small, single, simple; 9- V0, VOI stouter than pre-

ceding ones; hair 9-VII stout, single, simple; 9- VII usually with 2 main

stems (1-2), barbed, not reaching beyond fringe of paddle. Paddle. Mar-

gins with fringe; hair 1-P single.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; inner mouth brushes not pectinate at

tip; head hair 4-C well developed, branched, closer to 6-C than 5-C, cepha-

lad and mesad of 6-C; 5,6, 8,9,13-C single; 7,12-C usually double (2-3); 10-C

usually double (1-2); 11-C usually 3-branched (3-4); 14-C usually with 4-5

branches (4-6); 15-C usually 3-branched (2-3); mentum with 11-12 teeth on

each side. Thorax. Hair 1-P 3-branched; 2-P single; 3-P double; 4-P 3-

branched; 5, 6-P single; 7-P double; 9-P double; 11-P usually double (1-2);

14-P 3-branched; 5,7-M single; 6-M 3-branched; 8-M 4-branched; 9-M 3-

branched; 10,12-M single, long, stout; 11-M single, small; 7-T usually 5-

branched (5-6); 9-T usually double (2-3); 10,11-T similar to those on meso-

thorax; 12-T much reduced. Abdomen. Hair 6-I 3-branched; 7-I single; 6-II

double; 7-II 3-branched; 6-III-V double; 6-VI single; 1-VII usually 4-branched

(3-4); 2-VII usually with 3-4 branches; comb of 8-12 scales in a Single row,

each scale with fine denticles or fringe at base of apical spine; comb scale

with free portion rather slender, nearly parallel- sided from base and as long

as attached portion; pentad hair 2,4-VIII single; 1-VII with 4-5 branches;

3-VIII with 5-6 branches; 5-VII with 4-6 branches; siphon short, about twice

as long as wide, acus absent; pecten teeth 9-14 in number, evenly spaced,

each tooth with 2-3 basal denticles; 1-S with 3-4 branches, inserted well be-

yond last tooth and in line with teeth; saddle incomplete; marginal spicules

very small and inconspicuous; 1-X 2-branched; 2-X 2-branched; 3-X single;

ventral brush with 4 pairs of hairs on grid, each hair single except the 2

proximal ones usually double (1-2), 4d-X much shorter and smaller than

pepe no precratal tufts; anal papillae about 3 times as long as saddle, sau-

sage- like.

TYPE DATA. Aedes Gtegomyia) patriciae Mattingly, holotype male,

allotype female in British Museum (Natural History), London; type locality:

7,000 ft. on Krol Mountain, near Solan, Western Himalayas, INDIA, 20-VI-

1923 (Barraud). Paratypes: 4 males, 4 females, with same data as holotype,

in British Museum.

DISTRIBUTION. 87 specimens examined: 250, 169, 21c terminalia,

2L, 18 individual rearings (5 1, 18 p).

INDIA. "Western Himalayas, Krol Mountain" (20-VII-1923, Bar-

Sine ks Ail 1o terminalia; Punjab: Rawalpindi, Murree, W. Kimal (1922,

Gill), “to, 22,

THAILAND. Chiang Mai: (4-IV-1953, Thurman), 3c, 49, 4c ter-

minalia; Doi Suthep (15-VIII-1963, Sahem), 1c, 19, 1c terminalia; Banchang

Khiam (30-Il-1970, SEATO), 1c, 1¢ terminalia, 1 individual rearing (1 p):

Songkhla: Ton Nga Chang (26-III-1965, Kol), 2%, 2c terminalia, 2 individual

rearings (2 p); Tak: Khao Salak Phra (5-VII-1965, Sumeth), 11¢, 59, llc

terminalia, 2 L, 15 individual rearings (51, 15 p).

VIET NAM. Quang Tri: An Khe (5-XI-1967, Neal), 1c, 1c termi-

nalia.

28 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

REMARKS. There are 4 slides of larval and pupal skins and 4 of

whole 4th instar larvae in B.M., all with the same data as holotype, but 3

of the larval and pupal skins (Nos. 2439, 2450, 2451) are dated 4-VII-1930

and the remaining slide (No. 2512) is dated 16-VIII-1930. Unfortunately, no

associated adults of these slides can be found. I have not used this material

in the larval and pupal descriptions since its true identity is not known.

TAXONOMIC DISCUSSION. A. patriciae isa member ofthe albopictus

subgroup. The adult is very similar to flavopictus, a Palearctic species of

the albopictus subgroup, but can be separated from it by having the scutum with

a patch of narrow, curved, pale, yellowish scales on the lateral margin just be-

fore level of wing root and the halter without pale scales; in flavopictus these

scutal scales are golden yellowishin color and the halter is pale scaled.

In Southeast Asia, patriciae greatly resembles pseudalbopictus, for

which it can easily be mistaken. Thus, great care must be taken in identi-

fying them. The adult of patriciae can easily be distinguished from pseud-

albopictus by having the scutum without a patch of broad dark scales on each

side of prescutellar space between prescutellar white line and postdorso-

central white line.

The male terminalia of patriciae arevery similar to subalbopictus but

can be separated from it by tergum IX having the center produced into a

rounded lobe, claspette enlarged without a 90 degree lateral distal angle in later-

al aspect (dissected niaenettel and with numerous setae and several widened

specialized clubbed ones on sternal side of expanded distal part; in sub-

albopictus tergum IX is produced centrally as a wide, curved lobe, claspette

has a broad stem and a 90 degree lateral distal angle in lateral aspect (dis-

sected claspette) and is provided with numerous setae and several widened

specialized blade-like ones on sternal side of distal part, all the setae with

hooked tips and about same length as widened specialized blade-like ones.

The larvaof patriciae isvery similar to downsi but can be separated from

it by having comb scale with free portion rather slender, nearly parallel- sided

from base; in downsi the free portion of scale is widened at base and sharply

pointed at tip. The pupa of patriciae resembles pseudalbopictus but can be sepa-

rated from it by having hair 9-VIII usually with 2 (1-2) branches, each barbed and

not reaching beyond fringe of paddle; in pseudalbopictus hair 9- VIII has 2

branches, each barbed and reaching beyond the fringe of paddle.

A. patriciae apparently is confined to the Oriental Region. It is pre-

sently known from India, Thailand and Viet Nam. In Southeast Asia it is re-

ported here for the first time from Thailand (Chiang Mai, Songkhla, Tak) and

Viet Nam (An Khe).

BIOLOGY. The immature stages of patriciae have been found mainly

in tree holes in Thailand. The pupa from Chiang Mai, Banchang Khiam was

found in a stump hole. The male from Viet Nam was caught in a light trap.

AEDES STEGOMYIA) PSEUDALBOPICTUS (BOREL)

(Figs. 13, o terminalia, pupa; 14, larva; 20C,I, thorax)

Stegomyia sree eigenen Borel 1928, Arch. Inst. Pasteur Indo-Chinie 7:85

(o*, os L* °

Aedes Stegomyia) pseudalbopictus (Borel), Barraud 1931, Indian J. med.

Res. 19:223 (o*); Bonne-Wepster & Brug 1932, Geneesk. Tijdschr.

v. Ned.-Ind. 2:81 (o*, L*); Barraud 1934, Fauna Brit. India 5:235

(o*, a L).

MALE. Head. Proboscis dark scaled, with a few pale scales on ven-

tral side, as long as fore femur; palpus dark, longer than proboscis, with

white basal band on each of segments 2-5; those on segments 4-5 incomplete

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 29

dorsally; segments 4-5 subequal, slender, upturned, and with only a few

Short hairs; antenna plumose, slightly shorter than proboscis. Thorax. Scu-

tum with narrow dark scales and prominent median stripe of similar white ones;

median stripe rather broad reaching from anterior margin to middle of scutum

where it becomes very narrow or broken and is followed by an inverted Y-

shaped marking which forks at beginning of prescutellar space; on each side

a posterior dorsocentral white line which does not reach to middle of scutum;

a patch of broad dark scales on each side of prescutellar space, between pre-

scutellar white line and posterior dorsocentral white line; a patch of narrow

curved white scales on lateral margin just before level of wing root and a few

narrow curved scales over wing root; posterior pronotum with a larger patch

of broad white scales and some dark narrow ones dorsally; postspiracular

area with white scales; subspiracular area with pale scales; upper sterno-

pleural scale patch reaches to anterior corner of sternopleuron; mesepimeral

scale patches connected forming a V-shaped white patch, the open end of 'V'

directed backwards. Wing. Dark scales on all veins except for minute basal

spot of white scales on costa; first forked cell 1.5 times as long as its stem.

Halter. With dark scales. Legs. Fore and mid femora dark anteriorly,

paler posteriorly; hind femur anteriorly with broad white stripe which widens

at base and is Separated from apical white scale patch; fore and mid tarsi with

basal white bands on tarsomeres 1-2; hind tarsus with basal white bands on

tarsomeres 1-4, the ratio of length of white band to total length of tarsomere

is 1/3. 2/5. 1/2 and 2/3, tarsomere 5 all white. Abdomen. Abdominal seg-

ment I with white scales on laterotergite; terga II-VI each with a basal white

band and lateral white spots, or sometimes tergum II with lateral white spots

only; lateral spots do not connect with basal bands; tergum VII with lateral

white spots only; sterna III-VI with basal white bands; sternum VIII largely

covered with white scales. Terminalia. Basimere 3 times as long as wide,

with a patch of hairs on basomesal area of dorsal surface; claspette long and

slender, reaching to 0.7 of basimere, with 1 widened specialized spine-like

seta and numerous setae distal to it; distimere simple, elongate, as long as

basimere; with a spiniform process some distance from tip and with a few

hairs; tergum IX nearly flat at middle, with a hairy lobe on each side.

FEMALE. Essentially as in male, differing inthe following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Abdominal

basal pale bands on terga II-VI; segment VIII largely retracted.

PUPA. Cephalothorax. Trumpet 3.5 times as long as width at middle;

1,3-C single, longer than 2-C which is single; 4-C single or double; 5-C sin-

gle, or double; 6-C single, stout, shorter than 7-C; 7-C single or double; 10-

C usually 2-branched, mesad and caudad of 11-C; 11-C single. Abdomen.

Hair 1-I well developed, with more than 10 branches, dendritic; 2-I single;

3-I single, long; 2,3-I not widely separated, distance between them as dis-

tance between 4, 5-1; hair 1-II usually with 3-6 branches; 2-II laterad of 3-1;

hair 2-IV,V mesad of 1-IV, V; hair 1-II usually 2-branched; hair 3-II, III sin-

gle, shorter than segment III; hair 5-IV-VI single, not reaching beyond pos-

terior margin of following segment; 9-I-VI small, single, simple; 9- VI, VII

stouter than preceding ones; hair 9-VII single, simple; 9- VIII with 2 branches,

each barbed, reaching beyond fringe of paddle. Paddle. Margins with fringe;

hair 1-P single.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A in-

serted near middle of shaft, single; inner mouth brushes pectinate at tip; head

hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad and

mesad of 6-C; 5,8, 9,13-C single; 6-C usually double; 7-C with 2-4 branches;

10-C single or double; 11-C with 2-4 branches; 12-C with 2-3 branches; 14-C

with 3-4 branches; 15-C double; mentum with 9-10 teeth on each side. Thorax.

Hair 1-P 3-branched; 2-P single; 3, 4-P double; 5,6-P single; 7-P double;

9-P single; 11-P single; 14-P double; 5, 7-M single; 6-M with 2-3 branches;

8-M with 3-4 branches; 9-M 2-branched; 10,12-M single, long, stout; 11-M

30 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

Single, small; 7-T usually with 3-4 branches; 9-T double; 10,11-T similar to

those on mesothorax; 12-T much reduced. Abdomen. Hair 6-I with 2-3

branches; 7-I single; 6-II with 2-3 branches; 7-II double; 6-III double; 6-IV, V

single or double; 6-VI single; 1-VII usually 2-branched; 2-VII single; comb of

6-8 scales in a single row, each scale with fine denticles or fringe at base of

apical spine; pentad hair 2,4-VII single; 1-VII with 2-4 branches; 3-VII 3-

branched; 5-VIII with 2-3 branches; siphon short, less than twice as long as

wide, acus present, small; pecten teeth 3-6 in number, usually evenly spaced,

each tooth short and stout, with 3-4 basal denticles; 1-S with 3-4 branches,

inserted beyond last tooth and in line with teeth; saddle incomplete; marginal

spicules very small and inconspicuous; 1-X 2-branched; 2-X 2-branched; 3-X

Single; ventral brush with 4 pairs of hairs on grid, each hair single; 4d-X

much smaller than others; no precratal tufts; anal papillae 3 times as long as

saddle, sausage-like.

TYPE DATA. Stegomyia pseudalbopictus Borel, type location unknown;

type locality: Terre Rouges, COCHIN CHINA (Borel).

DISTRIBUTION. 681 specimens examined: 1740, 1472, 66c termi-

nalia, 35 L, 21, 2p, 152 individual rearings (99 1, 152 p).

INDIA. Bengal: Darjeeling Dist. - Sukna (II-1967, Ramalingam's

team), 33c, 119, 2 terminalia, 2 L, 10 individual rearings (71, 10 p);

Tindharia (III- 1967, Ramalingam's team), 4c, 32, 6 L, 4 individual rearings

(41, 4p); Mungpoo (III-1967, Ramalingam's team), 7c, 129, 2 terminalia,

6 individual rearings (1 1, 6 p); Pashok (II-1967, Ramalingam's team), 2c.

BURMA. Shan State: Aung Ban (X-1965, de Meillon), 5c, 492, 4c

terminalia, 21, 6p.

VIET NAM. Thua Thien: Phu Bai (VII-1965, R.T. Holway), 1°,

1o terminalia.

MALAYSIA. West Malaysia: Selangor- (XIl-1955, J.A. Reid), 5c,

49, 3c terminalia, 9 individual rearings (6 1, 6 p); Ulu Gombak (XII-1965,

Ramalingam's team), 1 L; ([X-1966, Ramalingam's team); 1c; Ulu Langat,

F.R. (X-1966, Ramalingam's team), 1c, 1c terminalia; (VI-1968, Rama-

lingam's team), 60, 49, 4 L, 2 individual rearings (21, 2p); Bt. Kutu (V-

1968, Ramalingam's team), 19; Pevak- Pulai (X-1967), 1 L; Chior F.R. (X-

1967), 3c, 39, 3 individual rearings (1 1, 3 p); Kg. Kuala Dipang (X-1967),

9%, 79, 2 individual rearings (2 p); Kg. Jalong (X-1967), 1c, 59, 1 individual

rearing (1 p); Lahat (X-1967), 2c’, 99, 1o terminalia, 11 individual rearings

71, 11 p); Lasah (X-1967), 2¢, 1o terminalia, 1 L, 1 individual rearing (1

p); C. Highlands Road (V0-1968), 5c, 59, 2 L, 3 individual rearings (3 1, 3

p); Chenderiang (X-1968), 1c; Kuala Kangsar (VI-1968), 29; Trong (VI-1968),

19, 1 L; all collected by Ramalingam's team; Pahang- Bentong Rd. (III- 1967),

1c; Kuala Lipis (IV-1967), 4°, 109, 1o¢ terminalia; Kuala Lipis, Pdg. Tungku

(IV-1967), 19, 1 individual rearing (1 1, 1 p); all collected by Ramalingam's

team; Perlis- Kg. Wang Tangga (IX-1967), 22; Kg. Wang Kelian (IX- 1967);

20°, 19; Kedah- Sintok F.R. (I[X-1967), 59; Kelantan- Gua Musang (IV-1967),

2°; Bertam (IV-1967), 2c, 1o¢ terminalia; Pasir Mas (X-1968), 2c, 59, 1 L,

6 individual rearings (6 1, 6 p); Rantau Panjang (X-1968), 19, 1 individual

rearing (11, 1 p); Machang (X-1968), 1c, 1 individual rearing (1 1, 1 p); Kota

Bharu (X-1968), 2c’, 49, 6 individual rearings (61, 6 p); all collected by

Ramalingam's téam.

THAILAND. Nakhon Nayok: (VI-1964, Kol & Sumeth), 2c, 2c ter-

minalia; Kanchanaburi: Huai Lin Thin (V-1965, peyton & Sumeth), 60, 4c

terminalia, 2 L; Huai Mae Nam Noi (V-1965, Peyton), 7°, 59, 7% terminalia,

6 individual rearings (41, 6 p); Huai Bong Ti (VI-1965, Peyton), 19, 1 indi-

vidual rearing (1 p); Nakhon Si Thammarat: Ban Saikae (VI-1966; Peyton's

team), 1c, 1o¢ terminalia, 1 individual rearing (1 p); Mae Hong Son: Ban Mae

Ho Nua (IX-1966, Kol), 59, 5 individual rearings (1 1, 5 p); Nan: Ban Wang

Mo (VIM-1966, Somboon), 40°, 49, 4c’ terminalia, 5 individual rearings (1 1,

5 p); Ban Pha Man (VII-1966, Chaliou), 4°, 4¢ terminalia, 3 individual

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia 31

rearings (3 p); Ban Pha Hang el ott 5 Somboon), 1c, 1¢ terminalia, 2 L;

Phangnga: ra Pak Chaung (X-1966, Kol), 19, 1 individual rearing (1 1, 1 p);

Tak Khet (X-1966, Chaliou), 19, 1 individual rearing (1 p); Surat Thani: Koh

Samui- Ban Li Pa Noi (IX-1967, Chaliou), 2c, 19, 2c terminalia, 3 individual

rearings (11, 3 p); Ban Lipa Noi (XII-1968-I-1969, Kol's team), 3c, 19, 2c

terminalia, 4 individual rearings (21, 4 p); Lampang: Ban Pha Daeng (IV-

1967, Somboon), 20, 22, 1o% terminalia, 1 individual rearing (1 p); Ban Na

Kiang (IV-1967, Kol), 29, 3 L, 2 individual rearings (21, 2 p); Huai Mae

Yuak (V-1968, Harrison), 17, 62, 16 terminalia, 2 L, 21 individual rear-

ings (161, 21 p); Huai Mae Phlung (V-1968, Kol & Harrison), 29, 1 individual

rearing (1 1, 1 p); Nakhon Sawan: Ban Nua Sathrni (XI-1968, Kol's team), 29,

2 individual rearings (21, 2p); Ang Thong: Ban Sang Thong (IV-1969, Kol's

team), 2c’, 19, 1o terminalia, 4 individual rearings (41, 4 p); Chiang Mai:

Ban Rong Rua Taeng (X-1969, Kol's team), 19, 1 individual rearing (1 1, 1 p);

Ban Kea Lek Noi (X-1969, Kol's team), 5c, 39, 1c terminalia, 1 L, 8 indi-

vidual rearings Mf 1, 8 p); Banka (X-1969, Kol's team), 9c, 59, 6 L, 11 indi-

vidual rearings (41, 11 p); Ban Rong Wua Daeng (II-1970, SEATO), 3c, 39,

30 terminalia, 6 individual rearings (5 1, 6 p); Ban Nong Pa Seet (I-1970,

SEATO), 19, 1 individual rearing (1 1, 1 p).

JAVA. Lembang, #11839, 1c; Tjandfoer, #7603, 1c.

TAXONOMIC DISCUSSION. A. pseudalbopictus is a member of the

albopictus subgroup. The adultis very similar to downsi, flavopictus, patriciae

and subalbopictus, in having the scutum with apatchof narrow, curved, white to

yellowish scales on lateral margin just before level of wing root, fore and mid fe-

mora without some pale scales scattered on anterior surface. However, pseud-

albopictus caneasily be distinguished from them by having the scutum with a patch

of broad dark scales on each side of prescutellar space, between prescutellar

white line and postdorsocentral white line, postspiracular area with scales. The

male terminalia of this species are markedly different from all other members of

the albopictus subgroup by having tergum IX nearly flat at middle, claspette

long and slender, reaching to 0.7 of basimere, 1 widened specialized spine-like

seta and numerous setae distal to it and the spiniform of the distimere placed

some distance from the tip.

The larva of pseudalbopictus is quite similar to albopictus and also

shares some similarities with alcasidi, malayensis and scutellaris, the mem-

bers of the other subgroup. It can easily be distinguished from all other

members of the group by having siphon acus present, pecten teeth 3-6 in

number, each tooth short and stout and usually with 3-4 basal denticles. The

pupa of pseudalbopictus is very similar to albopictus and patriciae but can be

separated from them by having hair 9- VIII with 2 branches, each barbed,

reaching beyond fringe of paddle; in albopictus hair 9-VIII is usually single

(1-2); when it is 2-branched, then only the male pupa can be separated from

pseudalbopictus by having the male genital lobe short and broad, about as

long as wide, whereas in pseudalbopictus it is rather long and narrow, longer

than wide; in patriciae hair 9-VIII usually with 2 (1-2) branches, each barbed,

not reaching beyond fringe of paddle.

A. pseudalbopictus apparently is a common species in the Oriental

Region and extends into the western part of the Indomalayan area. It is

presently known from India, Burma, Thailand, Viet Nam, W. Malaysia and

Java. The species is recorded here for the first time from Burma, Thailand

and Java.

BIOLOGY. The immature stages of pseudalbopictus have been found

mainly in bamboo stumps in Burma, India, Malaysia and Thailand. It has

also been found in fallen split bamboo, coconut shells and artificial containers

in Malaysia and in bamboo internodes, bamboo cups, bamboo pots, split bam-

boos and tree holes in Thailand. The immature stages were associated with

albopictus.

32 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

AEDES (SSTEGOMYIA) SEATOI HUANG

(Figs. 15, o«; 16, o terminalia, pupa; 17, larva)

Aedes Stegomyia) seatoi Huang 1969, Proc. ent. Soc. Wash. 71(2):234 (o*,

2, P, L*).

MALE. Head. Proboscis dark scaled, without any pale scales on

ventral side, as long as fore femur; palpus dark, longer than proboscis, with

white basal band on each of segments 2-5; those on segments 4-5 incomplete

dorsally; segments 4-5 subequal, slender, upturned, and with only a few short

hairs; antenna plumose, slightly shorter than proboscis. Thorax. Scutum

with narrow dark scales and a prominent median stripe of similar white ones;

stripe reaches from anterior margin to middle of scutum where it becomes

very narrow or broken and is followed by an inverted Y-shaped marking which

forks at beginning of prescutellar space. There is on each Side of this: (1) a

posterior dorsocentral white line which does not reach to middle of scutum

and which sometimes becomes very narrow or broken at level of wing root,

(2) a small white patch of similar scales at a short distance anterior to posterior

dorsocentral white line, (3) a few narrow white scales on anterior prescutal

area and some narrow white ones on scutal angle area where they form a

small white patch, (4) a patch of broad flat white scales on lateral margin just

before level of wing root and a few similar scales on posterior portion of

Supraalar area. There is no complete supraalar line of broad white scales;

posterior pronotum with a large patch of broad white scales and some dark

narrow ones dorsally; postspiracular area without scales; subspiracular area

with pale scales; mesepimeral scale patches connected forming a V-shaped

patch, the open end of 'V' directed backwards. Wing. Dark scales on all

veins except for minute basal spot of white scales on costa; first forked cell

1.5 times as long as its stem. Halter. With dark scales. Legs. Fore and

mid femora dark with some pale scales scattered anteriorly, more so on mid

than on fore femur, paler posteriorly; hind femur anteriorly with a broad

white stripe which widens at base and is narrowly separated from the apical

white scale patch; fore and mid tarsi with basal white bands on tarsomeres I-

3; hind tarsus with basal white bands on tarsomeres 1-4, the ratio of length of

white band to total length of tarsomere is 2/5, 2/5, 1/2 and 2/3, tarsomere 5

all white, or sometimes with a few dark scales at tip on ventral side. Abdo-

men. Abdominal segment I with white scales on laterotergite, tergum I with

a large median patch of white scales; terga II-VI each with a basal white band

which widens laterally except on tergum II where it widens in middle; all seg-

ments with lateral white spots which are not connected with basal bands; ter-

gum VII with lateral white spots only; sterna I-III] largely covered with white

scales; IV-VI each with a basal white band; sternum VII largely covered with

white scales. Terminalia. Basimere 3 times as long as wide, with a patch

of hairs on basomesal area of dorsal surface; claspette long, reaching to 0.75

of basimere, with numerous setae and several widened specialized curved

ones on mesal side of slightly expanded distal part; distimere simple, elongate,

0.75 as long as basimere; with a spiniform process at apex and with some

hairs; tergum IX with middle part produced into a large rounded lobe and with

a small hairy lobe on each side.

FEMALE. Essentially as in male, differing in the following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Abdominal

tergum II with basal white band also widening laterally; abdominal basal bands

on terga II-V0; segment VUI largely retracted.

PUPA. Cephalothorax. Trumpet short, 3 times as long as width at

middle; hair 1,3-C single, longer than 2-C; 2-C usually double; 4-C usually

double; 5-C usually 4-branched (4-5); 6-C single, stout, shorter than 7-C; 7-C

usually double; 10-C usually 4-branched, mesad and caudad of 11-C;11-C

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 33

single. Abdomen. Hair 1-I well developed, with more than 10 branches,

dendritic; 2-I single; 3-I single, long; 2,3-I not widely separated, distance

between them as distance between 4, 5-I; hair 1-II usually with 8-10 branches;

2-II laterad of 3-II; hair 2-IV, V mesad of 1-IV, V; hair 1-III usually with 3-4

branches; hair 3-II, III single, shorter than segment III; hair 5-IV-VI single

or double, not reaching beyond posterior margin of following segment; 9-I, II

small, single, simple; 9-III-VIII stouter than preceding ones; hair 9-II- VII

strongly developed, thickened; 9- VII usually single and barbed; 9-VIUI with a

strong main stem and lateral branches of varying length. Paddle. Margins

with rather short fringe on apical half; hair 1-P single.

LARVA. Head, Antenna 0.5 length of head, without spicules; 1-A in-

serted near middle of shaft, single; inner mouth brushes pectinate at tip; head

hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad and

mesad of 6-C; hair 5, 6, 8, 9,13-C single; 7,10,12-C usually double; 11-C usu-

ally 3-branched (3-5); 14-C with 3-4 branches; 15-C usually with 2-3 branches;

mentum with 10-12 teeth on each side. Thorax, Hair 1-P with 4-5 branches;

2-P single; 3-P 3-branched; 4-P 5-branched; 5-P with 3-4 branches; 6-P sin-

gle; 7-P with 2-3 branches; 9-P usually double (1-2); 11-P single; 14-P usu-

ally 7-branched (5-9); 5, 7-M single; 6-M 3-branched; 8-M with 4-5 branches;

9-M with 2-3 branches; 10,12-M single, long, stout; 11-M single, small; 7-T

usually with 4-5 branches; 9-T double; 10,11-T similar to those on mesotho-

rax; 12-T much reduced; basal spine of meso- and metapleural hairs stout

and straight or slightly curved. Abdomen. Hair 6-I with 3-4 branches; 7-I

single; 6-II usually 3-branched (2-3); 7-II 3-branched; 6-III-V double; 6- VI

single; 1-VII usually 5-branched; 2-VII usually 6-branched (5-8); comb of 6-10

scales in a single row, each scale with prominent denticles at base of apical

Spine; pentad hair 2,4-VII single; 1-VII usually 6-branched (5-7); 3- VIII with

4-6 branches; 5-VIII usually 7-branched (5-8); siphon short, less than twice

as long as wide, acus absent; pecten teeth 8-12 in number, evenly spaced,

each tooth with 2-4 basal denticles; 1-S 4-branched, inserted beyond last

tooth and in line with teeth; saddle incomplete; marginal spicules very small

and inconspicuous; 1-X usually 2-branched (2-4); 2-X 2-branched; 3-X single;

ventral brush with 4 pairs of hairs on grid, each hair usually single, some-

times, however, 1 or 2 proximal ones double; no precratal tufts; anal papillae

longer than saddle, lanceolate.

TYPE DATA. Aedes Stegomyia) seatoi Huang, holotype male, with

associated larval and pupal skins and terminalia on a slide, allotype female,

with associated larval and pupal skins, in U.S. National Museum, Washington,

D.C.; type locality: Bangphra, Chon Buri, THAILAND, 27-IX-1968 (Kol's

team). Paratypes: same locality and collectors as holotype; 2 males, with

associated larval and pupal skins and terminalia slides, 1 male, with associ-

ated pupal skin and terminalia slide, 23-IX-1968; 2 females, with associated

larval and pupal skins, 17-IX-1968, in U.S. National Museum.

DISTRIBUTION. 454 specimens examined: 1180, 1102, 300 terminalia,

48 L, 76 individual rearings (721, 76 p).

THAILAND. Chon Buri: Bangphra (IX-1968), 150, 79, 150 termi-

nalia, 5 L, 24 individual rearings (23 1, 24 my Khao Mai Kaeo (X-1963), 1¢,

1o terminalia; Kanchanaburi: Hinlub village (VII-1964, Sumeth), 1c, 1c ter-

minalia, 1 individual rearing (1 p); Nakhonsawan: Ko Klang Daet (XI-1968),

oo, 49, 50 terminalia, 9 individual rearings (9 1, 9 p); Ban Phanom Set (XI-

1968), 5c, 19, 5c terminalia, 1 L, 4 individual rearings (3 1, 4 p); Ban Ta

Khian Luan (XI-1968), 3c, 49, 3c terminalia, 7 individual rearings (71, 7 p);

Ko Yuan (XI-1968), 19, 12 L, 1 individual rearing (1 1, 1 p); Khao Kop (XI-

1968), 1 L; Ang Thong: Ban Phothong (IV-1969), 10c, 39, 6 L, 11 individual

rearings ({1 1, 11 p); Ban Bang Chaocha (II-1969), 10c¢, 19, 20 L, 10 indi-

vidual rearings (9 1, 10 p); Ban in Pramun (IV-1969), 4c, 4 individual rear-

ings (41, 4p); Ban Bang Thong (IV-1969), 1c, 1 individual rearing (1 1, 1 p);

Ban Sang Thong (IV-1969), 1c, 2 individual rearings (21, 2 p); Amphao Murui

34 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

(11-1969), 1 L; Chiang Mai: Ban Pasak Khwang (X-1969), 2c, 2 L, 2 indi-

vidual rearings (21, 2 p); Savaburi: Tambol Huapluak, Ban Nam Tone, Mulo

(11-1970, Prajims' team), 7<, 379. Two progeny rearings: NO. (3)- 29¢,

379; NO. (5)- 240, 159. Except as indicated, all specimens were collected

by Kol's team.

TAXONOMIC DISCUSSION. A. seatoi, a member of the albopictus

subgroup, is a very clearly marked species in all stages. The adult is very

Similar to albopictus in having the scutum with a patch of broad flat white

scales on lateral margin just before level of wing root. It can easily be re-

cognized, however, in having the scutum with a small white patch of narrow

scales on scutal angle area and abdominal tergum I with large median patch

of white scales; in albopictus the scutum is without such a patch of scales on

scutal angle area and abdominal tergum I lacks a large median patch of white

scales. The male terminalia of this species are also very similar to albo-

pictus but can easily be distinguished by having tergum IX with middle part

produced into a large lobe and not the conspicuous horn-like median projection

of albopictus.

The larva of seatoi shows some resemblance to albopictus and flavo-

pictus. It can easily be distinguished from all other members of the group by

having comb scales with prominent denticles at base of apical spine and hair

2-VII usually with 6 (5-8) branches. The pupa of seatoz is very similar to

downsi but can easily be separated from it by having hair 9-III-V strongly de-

veloped, thickened, much stouter than 9-II and hair 9- VII with a strong main

stem and lateral branches of varying length; in downsi hair 9-III-V is not so

strongly developed, slender and of about the same magnitude as 9-II, hair

9-VIII usually has 2 main stems (1-2), each with lateral branches of varying

length and reaching beyond fringe of paddle.

A. seatoi is presently known only from Thailand. The immature

stages are often found in association with albopictus and aegypti in the field.

In addition, the larva having the aegypti type of comb scales can easily be

misidentified and great care must be taken in this regard. It can easily be

separated from albopictus by having the comb scales with prominent denticles

at base of apical spine, whereas in albopictus the comb scales have only very

fine denticles in this position. .The pupa of seatoz is easily distinguished from

albopictus by having hair 9-III-V strongly developed, thickened, much stouter

than 9-II.

The immature stages of seatoi are markedly different from aegypit,

which belongs to a different group. The larva of seatoi can easily be sepa-

rated from aegypti by having hair 14-P usually with 7(5-9) branches, hair

1-VII usually 5-branched, 2-VII usually with 6(5-8) branches, ventral brush

with 4 pairs of hairs on grid, each hair usually single, sometimes 1 or 2 of

the proximal ones double; in aegypti hair 14-P usually with 2-3 branches,

1-VII usually 2-branched, 2-VII usually single, ventral brush with 5 pairs of

hairs on grid, each hair branched. The pupa of seatozi can easily be sepa-

rated from aegypti by having the paddle margins with fringe of hair-like spic-

ules; in aegypti the margins bear distinct denticles and lack a fringe of deli-

cate hairs.

BIOLOGY. The immature stages of seatoi have been collected mainly

in bamboo pots and bamboo cups which were placed in orchard plantations, in

villages and in mangrove forests in Thailand. The specimens from Kanchana-

buri, Hinlub village and Chon Buri, Khao Mai Kaeo were found in banana trees

and the larvae from Chiang Mai, Ban Pasak Khwang were found in a bamboo

stump. The females have been taken biting man in Saraburi, Thailand. The

immature stages were associated with aegypti, albopictus and Armigeres sp.

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 35

AEDES STEGOMYIA) SUBALBOPICTUS BARRAUD

(Figs. 18, o terminalia, pupa; 19, larva; 20A,L, thorax, hind leg)

Aedes (SStegomyia) subalbopictus Barraud 1931, Indian J. med. Res. 19:225

(o*); Barraud 1934, Fauna Brit. India 5:238 (c*).

MALE. Head. Proboscis dark scaled, sometimes with a few pale

scales on ventral side, as long as fore femur; palpus dark, as long as probos-

cis, with white basal band on each of segments 2-5; those on segments 4-5 in-

complete dorsally; segments 4-5 subequal, slender, upturned and with only a

few short hairs; antenna plumose, shorter than proboscis. Thorax. Scutum

with narrow dark scales and a prominent median stripe of similar white ones,

which narrows abruptly a short distance in front of prescutellar bare space

and forks at beginning of prescutellar space; on each side a posterior dorso-

central pale line which does not reach to middle of scutum; a patch of narrow

curved pale scales on lateral margin just before level of wing root and a few

narrow curved pale scales over wing root; posterior pronotum with a large

patch of broad white scales and some dark narrow ones dorsally; postspirac-

ular area without scales; subspiracular area with pale scales; upper sterno-

pleural scale patch reaches to anterior corner of sternopleuron; mesepimeral

scale patches connected forming a V-shaped white patch, the open end of 'V'

directed backwards. Wing. Dark scales on all veins except for a minute

basal spot of white scales on costa; first forked cell twice as long as its stem.

Halter. With dark scales. Legs. Fore and mid femora dark anteriorly,

paler posteriorly; hind femur anteriorly with a broad white stripe which widens

at base and is separated from apical white scale patch; fore and mid tarsi with

basal white bands on tersomeres 1-2; hind tarsus with basal white bands on

tarsomeres 1-4, the ratio of length of white band to total length of tarsomere

is 1/4, 1/4, 1/3 and 3/5, tarsomere 5 all white. Abdomen. Abdominal seg-

ment I with white scales on laterotergite; terga II-VI with lateral spots only

(holotype male); or terga III-VI with some pale scales on basal area forming

an incomplete band or indistinct bands at middle and with lateral white spots;

lateral spots do not connect with basal incomplete bands; terga II and VII with

lateral white spots only; sterna III-VI with basal white bands; sternum VII

largely covered with white scales. Terminalia. Basimere 3 times as longas

wide; with a patch of hairs on basomesal area of dorsal surface; mesal sur-

face membranous; claspette with broad stem, with numerous setae and several

widened specialized blade-like ones on sternal side of distal part; all setae

with hooked tips and about same length as specialized blade-like ones; there

is a 90 degree lateral distal angle (dissected claspette); distimere simple, e-

longate, as long as basimere; with a spiniform process and a few hairs near

apex; tergum IX with middle part forming a wide curved lobe and with a hairy

lobe on each side.

FEMALE. Essentially as in male, differing inthe following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Abdominal

basal bands incomplete at middle on terga II-VU; segment VII largely re-

tracted.

PUPA. Cephalothorax. Trumpet short, 3.5 times as long as width at

middle; hair 1,3-C single, longer than 2-C; 2-C single; 4-C usually double

(1-2); hair 5-C usually 4-branched (2-6); 6-C single, stout, shorter than 7-C;

7-C usually single (1-2); 10-C usually 3-branched (3-5), mesad and caudad of

11-C; 11-C single. Abdomen. Hair 1-I well developed, with more than 10

branches, dendritic; 2-1 single; 3-I single, long; 2, 3-I not widely separated,

distance between them as distance between 4,5-I; hair 1-II usually with more

than 10 branches (5-12); 2-II laterad of 3-II; hair 2-IV, V mesad of 1-IV, V;

hair 1-III usually with 3-4 branches (3-5), rarely double; hair 3-II, III single,

shorter than segment III; 5-IV-VI usually single (1-2), not reaching beyond

36 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

posterior margin of following segment; 9-I-V small, single, simple; 9-VI- VII

stouter than preceding ones; hair 9-VI stout, single and simple; 9- VIL stout,

single and barbed; 9- VIII usually with 2 main stems, barbed, not reaching be-

yond fringe of paddle. Paddle. Margins with fringe; hair 1-P single.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; head hair 4-C well developed, branched,

closer to 6-C than 5-C, cephalad and mesad of 6-C; hair 5, 6, 8, 9, 13-C single;

7,10-C single or double; 12-C double; 11-C usually double (2-3); 14,15-C usu-

ally double (2-4); mentum with 11-12 teeth on each side. Thovax. Hair 1-P

with 3-4 branches; 2-P single; 3-P double; 4-P with 2-3 branches; 5, 6-P sin-

gle; 7-P double; 9-P single; 11-P usually single (1-2); 14-P double; 5, 7-M sin-

gle; 6-M 3-branched; 8-M with 3-4 branches; 9-M usually double (2-3); 10, 12-

M single, long, stout; 11-M single, small; 7-T with 3-4 branches; 9-T double;

10,11-T similar to those on mesothorax; 12-T much reduced. Abdomen.

Hair 6-I with 2-3 branches; 7-I single; 6, 7-II double; 6-III-V usually double

(1-2); 6-VI single; 1-VII usually 3-branched; 2-VII 3-branched; comb of 8-10

scales in a single row, each scale with fine denticles or fringes at base of

apical spine; pentad hair 2,4-VIII single; 1-VUI with 3-4 branches; 3-VIII with

4-5 branches; 5-VIII usually 4-branched (3-5); siphon short, about twice as

long as wide, acus absent; pecten teeth 8-14 in number, evenly spaced, each

tooth usually with 2 (2-3) basal denticles; 1-S with 3-5 branches, inserted

well beyond last tooth and in line with teeth; saddle complete; marginal spic-

ules very Small and inconspicuous; 1-X 2-branched; 2-X usually single, some-

times 2-branched, one much smaller than other; 3-X single; ventral brush

with 4 pairs of hairs on grid, each hair single; no precratal tufts; anal papil-

lae about 3 times as long as saddle, sausage-like.

TYPE DATA. Aedes Stegomyia) subalbopictus Barraud, type male

(Y 149) in British Museum (Natural History), London; type locality: Bombay

Decean, Belgaum, INDIA, VIII-1921 (Barraud).

DISTRIBUTION. 39 specimens examined: 6c, 59, 60 terminalia, 11

individual rearings (11 1, 11 p).

INDIA. Bombay: Belgaum- Bombay Deccan (VUI-1921, Barraud),

1o, 1¢ terminalia; Madras: Nilgiri- Coonoor, Nilgiri Hills (14-IV-1969, B.

N. Mohan), 2c, 2c terminalia, 2 individual rearings (21, 2p); (29-V-1969,

B.N. Mohan), 22, 3 individual rearings (3 1, 3 p), one slide #101 (1 1, 1 p)

without adult; Sim's Park (1968, B.N. Mohan), 3¢, 19, 3o¢ terminalia, 4 in-

dividual rearings (41, 4 p); Bengal: Darjeeling Dist. - Sukna (III- 1967,

Ramalingam's team), 19, 1 individual rearing (1 1, 1 p).

TAXONOMIC DISCUSSION. The adult of subalbopictus is very similar

to pseudalbopictus and patriciae. It can easily be distinguished from pseud-

albopictus by the diagnostic characters mentioned in the key andfrom patriciae

by having the scutum with a patch of narrow curved white scales on lateral

margin just before level of wing root, abdominal basal bands usually incom-

plete or indistinct at middle, and hind tarsomere 3 with basal 1/3 white banded

and tarsomere 4 with basal 3/5 white banded; in patriciae the scutal patch is

pale yellowish in color, the abdominal basal bands are always complete on

terga II-VI, hind tarsomere 3 with basal 2/5-1/2 white banded and tarsomere

4 with basal 2/3-3/4 white banded. The male terminalia of this species are

very similar to patriciae but can be separated from it by the diagnostic char-

acters mentioned under the discussion of patriciae.

The larva of subalbopictus is very similar to novalbopictus but can be

separated from it by the diagnostic characters mentioned under the discus-

sion of novalbopictus. The pupa of subalbopictus is very similar to downsi

but can be separated from it by the diagnostic characters mentioned under

the discussion of downsi,

A. subalbopictus, an Oriental species of the albopictus subgroup, is

here recorded from India only. I have not seen any of the material mentioned

by Stone et al. (1959) as coming from Hainan Island,

Huang: Aedes (SStegomyia) scutellaris group in Southeast Asia 37

BIOLOGY. The larvae of subalbopictus have been found in tree holes,

bamboo internodes and bamboo stumps in India. The immature stages were

associated with pseudalbopictus and albopictus.

AEDES GTEGOMYIA) ALCASIDI N. SP.

(Figs. 21, o; 22, o terminalia, pupa; 23, larva; 26C, claspette)

Aedes Stegomyia) scutellaris (Walker), Knight & Hull 1952, Pacif. Sci. 6(2):

180 (o*, 2, L) (misidentification).

This species is named for Dr. Godofredo L. Alcasid, Department of

Education, National Museum, Manila, Philippines, in appreciation of his

interest in the mosquitoes which has greatly furthered our knowledge of the

Culicidae in the Philippines.

MALE. Head. Proboscis dark scaled, with pale scales on ventral

side, longer than fore femur; palpus dark, as long as proboscis, with white

basal band on each of segments 2-5; those on segments 4-5 incomplete dorsal-

ly; segments 4-5 subequal, slender, upturned, and with only a few short hairs;

antenna plumose, shorter than proboscis. Thorax. Scutum with narrow dark

scales and a prominent median stripe of similar white ones; stripe narrows

slightly posteriorly and forks at beginning of prescutellar space; on each side

a posterior dorsocentral pale line which does not reach to middle of scutum;

a supraalar line of broad white scales present; posterior pronotum with nar-

row dark scales on upper portion and broad white scales on lower portion

forming a stripe instead of a patch; postspiracular area without scales; sub-

Spiracular area with or without white scales; mesepimeral scale patches nar-

rowly separated. Wing. Dark scales on all veins except for minute basal

spot of white scales on costa; first forked cell 1.5 times as long as its stem.

Halter. With dark scales. Legs. Fore and mid femora dark anteriorly,

paler posteriorly; hind femur anteriorly with a broad white stripe which

widens at base and is narrowly separated from apical white scale patch; fore

and mid tarsi with basal white bands on tarsomeres 1-2; hind tarsus with basal

white bands on tarsomeres 1-4, the ratio of length of white band to total length

of tarsomere is 1/3, 2/5, 1/2 and 3/4, tarsomere 5 all white. Abdomen.

Abdominal segment I with white scales on laterotergite; tergum II dark dor-

sally, with lateral white spots only, or Sometimes with small median spot as

well;terga III- VI each with sub- basal white band which is connected to lateral

spots, sometimes tergum III with sub-basal median-spot and with lateral spots

which are turned dorsomesally; tergum VII with lateral white spots only; ster-

num VIII entirely covered with white scales. Tervminalia. Basimere 3.5

times as long as wide, with a patch of hairs on basomesal area of dorsal sur-

face; mesal surface membranous; claspette simple, with distal expanded part

subtriangular in shape, sternal and tergal sides not parallel but tapering,with

6 or 7 modified setae in a row on center of sternal side and occupying about

1/3 of it; apicotergal area with several distinctly long setae; distimere sim-

ple, elongate, as long as basimere, with a spiniform process and a few hairs

near apex; tergum IX with middle rounded and with a hairy lobe on each side.

FEMALE. Essentially as in male, differing inthe following respects:

palpus 0. 2 length of proboscis, with white scales on more than apical half.

Wing with first forked cell about twice as long as its stem. Abdominal terga

II-III always dark dorsally, with lateral white spots which are turned dorso-

mesally; sometimes tergum III with sub-basal median spot as well; terga IV-

VII each often with sub-basal white band which is connected to lateral spots or

sometimes tergum IV with sub-basal band incomplete at middle; segment VIII

entirely retracted.

38 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

, PUPA. Cephalothorax. Trumpet 3.5 times as long as width at

middle; hair 1,3-C single, longer than 2-C; 2-C usually single (1-2); 4-C

usually double (1-2); 5-C usually double; 6-C single, about as long as and

much stouter than 7-C; 7-C usually single (1-2); 10-C usually with 3-5

branches, mesad and caudad of 11-C; 11-C single. Abdomen. Hair 1-I well

developed, with more than 10 branches, dendritic; 2-I single; 3-I single, long;

2, 3-1 not widely separated, distance between them as distance between 4, 5-];

hair 1-II usually with 7-11 branches; hair 2-IV, V mesad of 1-IV, V; 1-III usu-

ally 2-3 branched; 1-IV usually double (1-2); 3-II, III single, shorter than seg-

ment III; hair 5-IV-VI single, not reaching beyond posterior margin of follow-

ing segment; 9-VII single and barbed or split at tip; 9- VIII usually with 2

main stems (1-2), barbed, reaching beyond fringe of paddle. Paddle. Mar-

gins with fringe; hair 1-P single.

LARVA. Head, Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; inner mouth brushes pectinate at tip;

head hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad

and mesad of 6-C; hair 5,6, 8,9,13-C single; 7-C usually 3-branched (2-3);

10,12-C usually double; 11-C usually 4-branched; 14-C with 2-4 branches;

15-C usually double (2-3); mentum with 11-12 teeth on each side. Thorax.

Hair 1-P usually 3-branched; 2-P single; 3-P double; 4-P 2-branched; 5, 6-P

single; 7-P double; 9-P single; 11-P usually double; 5, 7-M single; 6-M 3-

branched; 8-M with 4-5 branches; 9-M 3-branched; 10,12-M single, long,

stout; 11-M single, small; 7-T with 4-6 branches; 9-T usually 3-branched

(2-3); 10,11-T similar to those on mesothorax; 12-T much reduced. Abdo-

men. Hair 6-I 3-branched; 7-I single; 6-II with 2-3 branches; 7-II with 2-3

branches; 6-III- V double; 6-VI single; 1-VI usually 2-branched (2-3), long;

2-VII usually single (1-2); comb of 8-12 scales in a single row, each scale

with fine denticles or fringes at base of apical spine; sometimes 2-4 comb

scales connected at base; pentad hair 2, 4-VIII single; 1,5-VII with 3-5

branches; 3-VIII with 5-7 branches; siphon about 2.5 times as long as wide,

acus absent; pecten teeth 10-16 in number, evenly spaced, each tooth with 1

large and 1-3 small basal denticles; 1-S with 3-4 branches, inserted beyond

last tooth and usually before middle of siphon; saddle incomplete; marginal

spicules very small and inconspicuous; 1-X 2-branched; 2-X 2-branched; 3-X

single; ventral brush with 4 pairs of hairs on grid, each hair single except

the 2 proximal ones usually double (1-2), sometimes 4 proximal ones double;

no precratal tufts; anal papillae about 2.5 times as long as saddle, sausage-

like.

TYPE DATA. Holotype male (11-9) with associated larval and pupal

skins and terminalia slide (70/218), Dalton Pass, Nueva Ecija, Luzon,

PHILIPPINES, collected as a larva in a small tree hole, 3 ft. above ground

level, partially shaded, in a secondary rain forest, altitude 3500 ft., 6-VI-

1969 (Huang & Peyton). Allotype female (11-6) with associated larval and

pupal skins, with same data as holotype. Holotype and allotype deposited in

U.S. National Museum. Paratypes: 16 males, 19 females as follows: 12

males (11-1, 12,13; 9-4, 7,8; 8-4,5; 7-5, 6,8, 9) with associated larval and pu-

pal skins and terminalia slides; 4 males (11-104; 9-100; 8-100; 7-113) with

associated pupal skins and terminalia slides; 3 females (11-7; 8-1; 7-3) with

associated larval and pupal skins and terminalia slides; 11 females (11-4, 5,

8,10,11; 9-2,5,6,9; 7-1, 7) with associated larval and pupal skins; 5 females

(11-101, 102, 103; 8-101; 7-111) with associated pupal skins, all with same data

as holotype. Deposited in U.S. National Museum and British Museum.

DISTRIBUTION. 3,670 specimens examined: 539, 9209, 192 termi-

nalia, 169 terminalia, 1, 206 individual rearings (7971, 1, 206 p).

PHILIPPINES. Luzon: La Union- Agoo (IV-1945, J.G. Franclemont)

39; San Fernando (V-VI-1945, A.B.Gurney), 16<, 129, 7o terminalia; Calong-

boyan (VI-1945, A.B.Gurney), 40°, 49, 1¢ terminalia; Bacqui (VI-1945, A.B.

Gurney), 2c’, 89, 1¢ terminalia; Camansi (VI-1945, A.B.Gurney), 89.

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 39

Bataan- Pandan R. (VII-1931, W.V. King), 19; Pangasinan- Bayamloang (V-

1904), 19; San Fabian (I-1945, A.B. Gurney), 19. Albay- Malinao (VII-1964,

M. Delfinado), 19; Subic Bay (VI-1945, Rozeboom, Knight & Laffoon), 19.

Luzon: (V-1945, 32 MSU), 1c, 39; (V-1945, n32 MSU), 1o¢ terminalia (#662b)

only without adult. Batangas - Calatagan Oak cre , Alcasid's team), 3c,

59, 1o terminalia, 2 individual rearings (21, 2p). Nueva Vizcaya- Dalton

(VI-1967-IX-1968, Alcasid's team), 2c’, 62, 2c’ terminalia, 4 individual rear-

ings (41, 4p); Malete (VI-1969, Huang & Peyton), 36, 862, 30c terminalia,

2° terminalia, 114 individual rearings (78 1, 114 p); Aritao (VI-1969, Huang

& Peyton), 14°, 272, 8c terminalia, 41 individual rearings (29 1, 41 p).

Nueva Ecija-Dalton Pass (VI-1969, Huang & Peyton), 740, 1129, 62 termi-

nalia, 149 terminalia, 185 individual rearings (121 1, 185 p); Kaointalan (VI-

1969, Huang & Peyton), 9c, 479, 6c terminalia, 55 individual rearings (38 1,

55 p). Mountain Prov. - Lagawe (VI-1969, Huang & Peyton), 5c, 59, ao ter-

minalia, 10 individual rearings f 1, 10 p). Laguna- Los Banos (XI-1912),

49; (XI-1915), 12; Mt. Makiling (VI-XII-1967, Alcasid's team), 9c, 49, 8¢

terminalia, 4 individual rearings (3 1, 4 p); haere Huang & Peyton), 3c,

89, 11 individual rearings (21, 11 p); Pakil (VI-1968, Alcasid's team), 3c,

69, 2c terminalia, 5 individual rearings (5 1, 5 p); (VH-1969, Huang & Peyton)

22, 2 individual rearings (11, 2 p); Pangil (VI-V0-1969, Huang & Peyton),

231c, 3279, 17% terminalia, 549 individual rearings (364 1, 549 p); Lumban

pacer tes: Huang & Peyton), 2c, 59, 2c’ terminalia, 7 individual rearings

61, 7p); Kapatalan (VII-1969, Huang & Peyton), 20c, 159, 5c terminalia,

35 individual rearings (21 1, 35 p); Laguna (VIII-1967-IV-1968, Alcasid's

team), 1c, 19, 1c terminalia. Samar: San Antonio (XII-1944, J.H. Paullus),

22; Oras (I-1906, Gregory), 1¢, 1o terminalia; Osmena (VIII-1945, Rozeboom,

Knight & Laffoon), 50, 129, 3¢ terminalia. Leyte: Tacloban (VII-VII-1945,

H.R. Roberts), 49, 1 individual rearing (11, 1 p); (VIII-1945, Rozeboom,

Knight & Laffoon), 19; Dagami, Mt. Lobi (VIII-1945, H.R. Roberts), 19;

Baybay (VI-1945, Rozeboom, Knight & Laffoon), 19; Carigara nner ES.

Ross), 1o, 19, 1h terminalia; Abuyog (XI-1944, O.H. Graham), 19; Jinamoc

I. (Nav. Med.Sch. 113), 19. Mindoro: San Jose (III-IV-1945, E.S. Ross), 6c,

219, 3c terminalia; Oriental- Victoria, Alcate (VI-1969, Huang & Peyton),

123, 309, 41 individual rearings (381, 41 af (VII-1969, Harrison & Kol), 8¢,

239, 2% terminalia, 30 individual rearings (15 1, 30 p); Victoria, San Pedro

(VII-1969, Huang & Peyton), 1c, 1 individual rearing (11, 1 p); Naujan, San

Augustin hersignk eo Huang & Peyton), 350°, 649, 2c terminalia, 98 individual

rearings (641, 98 p). Mindanao: Kabakan (V-1945, R. Hoe a 19, 1¢

terminalia; Zamboanga (IX-1945, Rozeboom, Knight & Laffoon), 4c, 792, 2¢

terminalia; Pasanonco (IX-1945, Rozeboom, Knight & Laffoon), 1c, 39.

Palawan: Irahnan R. (VI-1945, Rozeboom, Knight & Laffoon), 100, 89, 9c

terminalia; Tacburos (VI-1945, Rozeboom, Knight & Laffoon), 19; Iwahig (VI-

1945, Rozeboom, Knight & Laffoon), 29; (XI-1968, Alcasid's team), 2c, 69,

8 individual rearings (2 l, 8p); F. A. W. 10 Camp (VI-1945, Rozeboom,

Knight & Laffoon), 3c, 39, 3c terminalia; Puerto Princesa (V-VI-1945,

Rozeboom, Knight & Laffoon), 2c, 119, 20 terminalia; (IX-1945, 19th MGL),

1¢ terminalia (Lot. P2BO- (3-4) o gen. #1c, C.N.H.M.) only without adult;

bw zen _~yCamarins Norte (X-1968, Alcasid's team), 19, 1 individual rearing (1 p);

4g Palawan; 1°. Philippine Islands (J.H. Paullus), 1o terminalia (#83 Sta. 3,

45-VI-12) only without adult; Philippines, APO 321 (V-1945, E.S. Ross), 60.

Calicoan I.: (I-1945, J.H. Paullus), 2c, 69, 2c terminalia. Basilan I.:

Isabela (1945, Rozeboom, Knight & Laffoon), 12. Basbas I.: (IV-1967; M.

Delfinado), 2c, 19, 2c terminalia, 2 individual rearings (2 p). Sulu Arch.:

ores M. Delfinado), 1o, 1c terminalia. Sanga Sanga I. : Lapit- Lapit

IV-1967, M. Delfinado), 1c.

TAXONOMIC DISCUSSION. A. alcasidi is a member of the

scutellaris subgroup. The adult differs from all the other members

of the albopictus subgroup by having the supraalar white line complete

and well developed, with broad flat scales over wing root and toward the

40 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

scutellum. It is very similar to malayensis, riversi and scutellaris in having

the mid femur without a median white line on anterior surface, wing with mi-

nute basal spot of white scales on costa and hind tarsomere 5 all white. The

adult is indistinguishable from scutellaris except for the male terminalia; it

can be separated from malayensis and riversi by having hind tarsomere 3 with

basal half white banded and hind tarsomere 4 with basal 3/4 white banded; in

malayensis and riversi hind tarsomere 3 has the basal 2/5 white banded and

hind tarsomere 4 has the basal 2/3 white banded.

The male terminalia of alcasidi are very similar to hensilli, malay-

ensis and viverysi in having the distal expanded part of claspette subtriangular

in shape in lateral aspect (dissected claspette), sternal and tergal sides not

parallel but tapering and without an apicosternal angle. It is closer to hensillz

than to malayensis and viversi because of the presence of several distinctly

long setae on the apicotergal area of the claspette; it can, however, be sepa-

rated from hensilli in having claspette with 6 or 7 modified setae in a row on

center of sternal side occupying about 1/3 of it; in hensilli the claspette usu-

ally has 7 modified setae, the basal one often rather smaller, set in a row on

a slight prominence on center of sternal side and occupying about 2/5 of it.

The larva of alcasidi is very similar to albopictus, malayensis , riversi

and scutellavis in having no siphon acus, saddle incomplete, hair 2-X 2-

branched and 2-VII usually single (1-2). It is closer to malayensis and

scutellaris in having hair 1-VII usually with 2 long branches (2-3), or, when

3-branched then one much smaller than other two, siphon about 2.5 times as

long as wide, pecten teeth 10-16 in number and 1-S inserted at or before mid-

dle of siphon. The larva of alcasidi is indistinguishable from scutellaris and

can only be separated from malayensis by having hair 1-S usually inserted

before middle of siphon instead of at middle; pecten teeth 10-14 in number,

each tooth with 2-4 basal denticles.

The pupa of alcasidi is very similar to albopictus, pseudalbopictus,

malayensis, vYiversi and scutellaris in having hair 9-VI about same magnitude

as 9-V, 9-VII single and simple and 9- VIII reaching beyond fringe of paddle.

It is closer to malayensis, riversi and scutellaris in having hair 6-C about

3/4 the length to about as long as 7-C. The pupa of alcasidi is indistinguish-

able from scutellaris,. It can be separated from malayensis and rivers? by

the diagnostic characters mentioned in the key.

Although the immature stages of alcasidi are so similar to scutellavis,

the male terminalia differ markedly and can easily be distinguished from

those of scutellaris by having the claspette with distal expanded part subtri-

angular in shape in lateral aspect (dissected claspette), sternal and tergal

sides not parallel but tapering and without an apicosternal angle; in scutellaris

the claspette has the distal expanded part square in shape in lateral aspect

(dissected claspette), sternal and tergal sides more or less parallel and the

apicosternal angle present. A. alcasidi has been mistaken in the past for

scutellaris (Knight & Hull, 1952) and for malayensis (Colless, 1962).

A. alcasidi is apparently restricted to the Philippines. The immature

stages greatly resemble albopictus and since the two often occur in the same

breeding places, care must be taken in identification. The discussion under

albopictus deals with this matter.

BIOLOGY. The immature stages of alcasidi have been collected

mainly in tree holes and bamboo stumps in the Philippines. It has also been

found in stump holes, coconut stumps, coconut husks, coconut spathes, coco-

nut shells; artificial containers and a shelf fungus in the Philippines. The

immatures were associated with albopictus.

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 41

AEDES STEGOMYIA) ALORENSIS BONNE-WEPSTER & BRUG

(Figs. 20G,H, 2 abdomen; 24B, - terminalia)

Aedes (Stegomyia) variegatus var. alorensis Bonne-Wepster & Brug 1932,

Geneesk. Tijdschr. v. Ned.-Ind. 72 (Bijblad a : 92 (o*); Stone &

Farner 1945, Proc. biol. Soc. Wash. 58: 161 (to sp. status; men-

tioned only); Marks 1954, Bull. Br. Mus. (nat. Hist.) Ent. 3(10):

383 (o'*, 9*) (taxonomy).

MALE. Head, Proboscis dark scaled, with a few pale scales on

ventral side, slightly longer than fore femur; palpus dark, slightly shorter

than proboscis, with white basal band on each of segments 2-5; those on seg-

ments 4-5 incomplete dorsally; segments 4-5 subequal, slender, upturned,

and with only a few short hairs; antenna plumose, shorter than proboscis.

Thorax.. Scutum with narrow dark scales and prominent median stripe of

Similar white ones; stripe narrows Slightly posteriorly and forks at beginning

of prescutellar space; on each side a posterior dorsocentral pale line which

does not reach to middle of scutum; a supraalar line of broad white scales

present; posterior pronotum with narrow dark scales on upper portion and

with broad white scales on lower portion forming a white stripe instead of a

white patch; postspiracular area without scales; subspiracular area witout

scales; mesepimeral scale patches narrowly connected. Wing. Dark scales

on all veins except for minute basal spot of white scales on costa; first forked

cell 1. 2 times as long as its stem. Halter. With dark scales. Legs. Fore

femur dark anteriorly, paler posteriorly; fore tarsus with basal white bands

on tarsomeres 1-2; mid femur with median white line on anterior surface;

the rest of mid leg broken off; hind femur anteriorly with broad white stripe

which widens at base and is narrowly separated from apical white scale patch;

hind tarsus with basal white bands on tarsomeres 1-4, the ratio of length of

white band to total length of tarsomere is 1/4, 1/3, 2/5 and 2/3; tarsomere 5

all white. Abdomen. Abdominal segment I with white scales on laterotergite;

tergum II with sub-basal white band and with lateral spots; spots do not connect

with sub-basal band; terga III- VI each with sub-basal white band which is con-

nected to lateral spots; sterna II-IV largely covered with white scales; sterna

V-VI with basal white bands. Terminalia. Basimere about 3 times as long

as wide; with a patch of hairs on basomesal area of dorsal surface; mesal sur-

face membranous; claspette complex, with numerous setae on expanded distal

part, each seta distinctly on a separate cone, a tergomesal finger-like pro-

cess which bears 6 modified setae at tip; distimere simple, elongate, as long

as basimere, with a spiniform process and a few hairs at apex; tergum IX

with middle truncates and with a hairy lobe on each side.

FEMALE. Essentially as in male, differing in the following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Fore leg

broken off; only femur of mid leg remains (as in male); hind femur, tibia and

tarsomere 1 only remain (as in male). Abdominal tergum II with median spot

and lateral spots which are turned dorsomesally; tergum VII with complete

sub-basal white band; segment VIII largely retracted.

PUPA and LARVA. Unknown.

TYPE DATA. Aedes Stegomyia) variegatus var. alorensis Bonne-

Wepster & Brug, type male in British Museum (Natural History), London;

type locality: Kalabaki, Alor Island (LESSER SUNDA ISLANDS) (near Timor),

7-I-1923 (S.L. Brug & H. de Rook).

DISTRIBUTION. 3 specimens examined: 1c, 19, 1c terminalia.

INDONESIA. Lesser Sunda Islands; Alor Island, Kalabaki (7-I-1923,

S.L. Brug & H. de Rook), 1, 1o terminalia; Alor Island (30-IlI-1926, Van

Beek), 19 (#6020).

TAXONOMIC DISCUSSION. A. alorensis is very similar to paullusi

in having mid femur with median white line on anterior surface and wing with

42 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

minute basal spot of white scales on costa. It differs, however, in having

the scutum without any white scales on lateral prescutal area and on scutal

angle area.

The male terminalia of alorensis have a complex claspette with numer-

ous setae each on a separate cone on the expanded distal part, a tergal mesal

finger-like process which bears 6 modified setae at tip, not 2 as described

by Bonne-Wepster & Brug (1932) and Marks (1954). It therefore differs from

all other species that have been described in this group. A. alorensis is an

Indomalayan species of the scutellavis subgroup. It is presently known from

Alor Island only.

BIOLOGY. Unknown.

AEDES STEGOMYIA) ANDREWSI EDWARDS

(Figs. 20D, ? abdomen; 24A, ¢ terminalia)

Aedes (SStegomyia) variegatus var. andrewsi Edwards 1926, Bull. ent. Res.

17: 103 (o*); Stone & Farner 1945, Proc. biol. Soc. Wash. 58: 159

(to sp. status); Marks 1954, Bull. Br. Mus. (nat. Hist.) Ent. 3(10):

383 (o*, 2*) (taxonomy).

MALE. Head. Proboscis dark scaled, with a few pale scales on ven-

tral side, slightly longer than fore femur; palpus dark, slightly shorter than

proboscis, with white basal band on each of segments 2-5; those on segments

4-5 incomplete dorsally; segments 4-5 subequal, slender, upturned, and with

only a few short hairs; antenna plumose, shorter than proboscis. Thorax.

Scutum with narrow dark scales and prominent median stripe of similar white

ones; stripe narrows slightly posteriorly and forks at beginning of prescutellar

space; on each side a posterior dorsocentral pale yellowish line which does not

reach to middle of scutum; a supraalar line of broad white scales present;

posterior pronotum with narrow dark scales on upper portion and with broad

white scales on lower portion forming a white stripe instead of a white patch;

postspiracular area without scales; subspiracular area with white scales; mes-

epimeral scale patches connected forming a V-shaped white scale patch; the

open end of 'V' directed backwards. Wing. Dark scales on all veins, without

minute basal spot of white scales on costa; first forked cell 1.5 times as long

as its stem. Halter. With dark scales. Legs. Fore and mid femora dark

anteriorly, paler posteriorly; hind femur anteriorly with broad white stripe

which widens at base and is narrowly separated from apical white scale patch;

fore and mid tarsi each with basal white band on tarsomere 1; hind tarsus

with basal white bands on tarsomeres 1-4, the ratio of length of white band to

total length of tarsomere is 1/5, 1/4, 1/3 and 1/2, tarsomere 5 all white.

Abdomen. Abdominal segment I with white scales on laterotergite; terga II-VI

with lunate lateral white spots only; sterna I-IV entirely white scaled; sterna

V-VI with basal white bands. Terminalia. Basimere about 3 times as long

as wide; with patch of hairs on basomesal area of dorsal surface; mesal sur-

face membranous; claspette simple, with 4 or 5 modified setae in a row on

apicosternal angle, with several rather long and stout setae on apicotergal

area; distimere simple, elongate, as long as basimere, with a spiniform pro-

cess and a few hairs at apex; tergum IX with middle rounded and with a hairy

lobe on each side.

FEMALE. Essentially as in male,differing in the following respects:

proboscis all dark scaled; palpus 0. 2 length of proboscis, with white scales

on apical half. Wing with first forked cell about twice as long as its stem.

Abdominal terga II-VII with lunate lateral white spots only; segment VIII large-

ly retracted.

PUPA and LARVA. Unknown.

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 43

TYPE DATA. Aedes Stegomyia) variegatus var. andrewsi Edwards,

type male in British Museum (Natural History), London; type locality:

CHRISTMAS ISLAND (south of Java), II-1902 (Dr. haan.

DISTRIBUTION. 12 specimens examined: 2c, 89, 2c terminalia.

CHRISTMAS ISLAND (south of Java) (III-1902, Dr. Durham), 2¢,

2s terminalia; (X-1910, Dr. R. Kirkpatrick), 19; North part of Island (1-1898,

Dr. C.W. Andrews), 19; Flying Fish Cove ([IX-X-1908, Dr. C.W. Andrews),

29; (11-1933, F. Harms), 49.

TAXONOMIC DISCUSSION. A. andrewsi isa clearly marked species in

the adult stage. Itcan easily be distinguished from all other members of the

group by having the supraalar white line complete and well developed, with

broad flat scales over wing root and toward scutellum, wing without minute

basal spot of white scales on costa, mid femur without median white line on

anterior surface and abdomen with lunate lateral white spots only.

The male terminalia of andrewsi arevery similar to scutellavis in having

claspette with distal expanded part square in shape in lateral aspect (dissected

claspette), sternal and tergal sides more or less parallel and apicosternal angle

present. It can easily be separated from scutellaris by having claspette with 4 or

5 modified setae in arow on apicosternal angle and with several distinctly long and

stout setae on apicotergal area; in scutellaris the claspette has 5 or 6 modified se-

tae set ona prominence close to apicosternal angle area and lacks long and stout

setae onapicotergal area. A. andrewsi is anIndomalayan species ofthe scu-

tellaris subgroup. Itis presently known from Christmas Island only.

BIOLOGY. Unknown.

AEDES STEGOMYIA) HENSILLI FARNER

(Figs. 25C,D,E, o terminalia; 31C, hind leg)

Aedes (Stegomyia) hensilli Farner 1945, Proc. biol. Soc. Wash. 58: 59 (c, 9,

L); Bohart & Ingram 1946, U.S. Navmed 1055 :25 (o'*, 9¢*, L); Marks

1954, Bull. Br. Mus. (nat. Hist.) Ent. 3(10): 383, Pl. 18 (o*, 9*);

Bohart 1956(1957), Insects of Micronesia, 12(1): 48 (“*, 2*, L*).

Aedes (SStegomyia) scutellaris hensilli Farner, Colless 1962, Proc. Linn.

Soc. N.S.W. 87: 314 (to ssp. status).

MALE. Head. Proboscis dark scaled, slightly longer than fore femur;

palpus dark, as long as proboscis, with white basal band on each of segments

2-5; those on segments 4-5 incomplete dorsally; segments 4-5 subequal, slen-

der, upturned and with only a few short hairs; antenna plumose, shorter than

proboscis. Thorax. Scutum with narrow dark scales and prominent median

stripe of similar white ones; stripe narrows slightly posteriorly and forks at

beginning of prescutellar space; on each side a posterior dorsocentral pale

line which does not reach to middle of scutum; a supraalar line of broad white

scales present; posterior pronotum with narrow dark scales on upper portion

and with broad white scales on lower portion forming a white stripe instead

of a white patch; postspiracular area without scales; subspiracular area with-

out scales; mesepimeral scale patches separated. Wing. Dark scales on allveins

except for minute basal spot of white scales on costa; first forked cell 1.2 times

as long as its stem. Halter. With dark scales. Legs. Fore and mid femora

dark anteriorly, paler posteriorly; hind femur anteriorly with broad white

stripe which widens at base and is separated from apical white scale patch;

fore and mid tarsi with basal white bands on tersomeres 1-2; hind tarsus with

basal white bands on tarsomeres 1-5, the ratio of length of white band tototal

length of tarsomere is 1/5, 1/4, 1/3, 1/2 and 1/2. Abdomen. Abdominal

segment I with white scales on laterotergite; tergum II dark dorsally, with

lateral white spots only; terga II-VI each with sub-basal white band which is

Ad Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

connected to lateral spots; tergum VII with lateral white spots only; sternum

VIII largely covered with white scales. Terminalia. Basimere 3.5 times as

long as wide; with patch of hairs on basomesal area of dorsal surface; mesal

surface membranous; claspette simple, with distal expanded part more or

less subtriangular in shape, usually with 7 modified setae, the basal one often

rather smaller, set in a row on a Slight prominence at center of sternal side

and occupying about 2/5 of it, apicotergal area with several distinctly long

setae; distimere simple, elongate, as long as basimere, with a spiniform

process and a few hairs near apex; tergum IX with middle rounded and with

a hairy lobe on each side.

FEMALE. Essentially as in male, differing in the following respects:

palpus 0. 2 length of proboscis, with white scales on more than apical half.

Abdominal tergum III usually dark dorsally, with lateral white spots only, or

sometimes with a sub-basal median spot as well; terga IV- VII each with a sub-

basal white band, or sometimes with sub-basal incomplete bands; segment

VU largely retracted. |

PUPA. Unknown.

LARVA. Not known with certainty.

TYPE DATE. Aedes (Stegomyia) hensilli Farner, holotype male in

U.S. National Museum, Washington, D.C.; type locality: Ulithi Is., W.

CAROLINES, XII-1944 (George Hensill). Paratypes: 8 males, 6 females,

UlithiIs., W. Carolines, XTI-1944 (G. Hensill) in U.S. National Museum.

DISTRIBUTION. 31 specimens examined: 150, 892, 80 terminalia.

W. CAROLINES, Ulithi Is. (XII-1944, George Hensill), 150, 89,

80° terminalia.

REMARKS. There are 10 whole 4th instar larvae on 5 slides in U.S.

National Museum, all marked W. Carolines, UlithilIs., (XII-1944, George

Hensill). No associated larval skins and pupal skins were found and I have

not used this material for larval and pupal descriptions. |

TAXONOMIC DISCUSSION. The adult of hensilli is very similar to

alcasidi, malayensis, riversi and scutellaris in having the mid femur with-

out a median white line on anterior surface and wing with a minute basal spot

of white scales on costa. It can easily be distinguished from them, however,

by having hind tarsomere 5 with basal 1/2 white banded instead of all white.

The male terminalia of hensilli are very similar to alcasidi, malay-

ensis and rviversi in having claspette with distal expanded part subtriangular

in shape in lateral aspect (dissected claspette), sternal and tergal sides not

parallel but tapering and without apicosternal angle. It is closer to alcasidi

than to malayensis and riversi in having apicotergal area of claspette with

several distinctly long setae. However, hensilli can be separated because

the claspette usually has 7 modified setae, the basal one often rather smaller,

set in a row on a Slight prominence at center of sternal side and occupying

about 2/5 of it; in alcasidi claspette has 6 or 7 modified setae in a row at

center of sternal side and occupying about 1/3 of it.

Colless (1957) reported a "scutellavis''form from Singapore as A.

hensilli Farner. In 1962, he described this Malayan form as A. (S.) s.

malayensis subsp. on the basis of laboratory hybridization experiments be-

tween this Malayan form and scutellaris from New Guinea. At the sametime,

he also considered that A. hensilli of the Carolines is a subspecies of A.

scutellaris (Walker). Apparently he did not recognize the morphological dif-

ferences between hensilli and malayensis in his paper (Colless, 1962). In

fact, the male terminalia of hensilli are quite different from malayensis and

can easily be identified by having the apicotergal area of claspette with sever-

al distinctly long setae, claspette usually with 7 modified setae, the basal

one often rather smaller, in a row set on a slight prominence at center of

sternal side and occupying about 2/5 of it; in malayensis the apicotergal area

lacks long setae and claspette has 7-10 modified setae on center of sternal

side and occupying about 1/2 of it.

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 45

The male terminalia of hensilli, particularly the shape of the claspette

are strikingly different from scutellaris and can easily be recognized by hav-

ing claspette with distal expanded part subtriangular in shape in lateral as-

pect (dissected claspette), sternal and tergal sides not parallel but tapering

and without apicosternal angle; in scutellavis the distal expanded part is

square in shape in lateral aspect (dissected claspette), sternal and tergal

sides are more or less parallel and apicosternal angle is present.

A. hensilli is a Western Pacific Is. species of the scutellaris sub-

group. It is presently known only from Ulithi Island, W. Carolines, and rec-

ords from other localities need a critical examination.

BIOLOGY. The larvae of hensilli were found in empty coconut shells,

tree holes, and to some extent in artificial containers such as tin cans as

well as discarded drums, barrels, and bottles used by natives (Farner, 1945).

AEDES GTEGOMYIA) MALAYENSIS COLLESS

(Figs. 26D,E, claspette, 9° terminalia; 27, o terminalia, pupa;

28, larva; 31D, hind leg)

Aedes Stegomyia) hensilli Farner, Colless 1957, Med. J. Malaya 12(2): 464

(o*) (misidentification).

Aedes (Stegomyia) scutellaris malayensis Colless 1962, Proc. Linn. Soc. N.

S.W. 87: of (s*, 9); Huang 1971, Proc. ent. Soc. Wash. 73(1): 2 (*,

PP Pe

MALE. Head, Proboscis dark scaled, sometimes with pale scales

on ventral side, slightly longer than fore femur; palpus dark, as long as pro-

boscis, with white basal band on each of segments 2-5; those on segments 4-

39 incomplete dorsally; segments 4-5 subequal, slender, upturned, and with

only a few short hairs; antenna plumose, shorter than proboscis. Thorax.

Scutum with narrow dark scales and prominent median stripe of similar white

ones; stripe narrows slightly posteriorly and forks at beginning of prescutel-

lar space; on each side a posterior dorsecentral white line which does not

reach to middle of scutum; a supraalar line of broad white scales present;

posterior pronotum with narrow dark scales on upper portion and with broad

white scales on lower portion forming a white stripe instead of a white patch;

postspiracular area without scales; subspiracular area without scales; mes-

epimeral scale patches well separated, sometimes narrowly connected. Wing.

With dark scales on all veins except for minute basal spot of white scales on

costa; first forked cell 1.5 times as long as its stem. Halter. With dark

scales. Legs. Fore and mid femora dark anteriorly, paler posteriorly;

hind femur anteriorly with a broad white stripe which widens at base and is

narrowly separated from apical white scale patch; fore and mid tarsi with

basal white bands on tarsomeres 1-2; hind tarsus with basal white bands on

tarsomeres 1-4, the ratio of length of white band to total length of tarsomere

is 1/4, 1/3, 2/5 and 2/3, tarsomere 5 all white or sometimes with a few

dark scales on apical ventral side. Abdomen. Abdominal segment I with

white scales on laterotergite; tergum II dark dorsally, with lateral white

spots only or sometimes with median spot as well; terga II-VI each with sub-

basal white band which is connected to lateral spots, sometimes tergum III

with a sub~basal median spot and with lateral spots which are turned dorso-

mesally; tergum VII with lateral white spots only; sternum VIII largely cov-

ered with white scales. Terminalia. Basimere 3.5 times as long as wide;

with a patch of hairs on the basomesal area of dorsal surface; mesal surface

membranous; claspette simple, with distal expanded part subtriangular in

Shape, sternal and tergal sides not parallel but tapering, with 7-10 modified

setae forming a prominent row on center of sternal side and occupying about

46 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

1/2 of it; distimere simple, elongate, as long as basimere, with a spiniform

process and a few hairs near apex; tergum IX with middle rounded and with

a hairy lobe on each side.

FEMALE. Essentially as in male, differing inthe following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Wing with

first forked cell about twice as long as its stem. Abdominal tergum I Ssome-

times with median spot; tergum II always dark dorsally, with lateral spots

which are turned dorsomesally; tergum III often dark dorsally, with lateral

spots only, or sometimes as in male; terga IV-VII each often with complete

sub- basal transverse white band which is connected to lateral spots, or

sometimes with incomplete sub-basal band on tergum IV; segment VII large-

ly retracted; sternum VII with conspicuous rounded lateral lobe; post- genital

plate with shallow notch; cerci short and broad; 3 spermathecae, 1 larger

than other 2.

PUPA. Cephalothorax. Trumpet 3 times as long as width at middle;

hair 1,3-C single, longer than 2-C; 2-C usually single (1-2); 4-C usually

single (1-2); 5-C usually 2-branched (2-3); 6-C single, much stouter than

7-C, slightly shorter than 7-C; 7-C usually single (1-2); 10-C usually with

3-4 branches, mesad and caudad of 11-C; 11-C single. Abdomen. Hair 1-1

well developed, with more than 10 branches, dendritic; hair 2-II laterad of

3-II; 2-I single; 3-I single, long; 2,3-I not widely separated, distance be-

tween them as distance between 4,5-I; hair 1-II usually with 7-12 branches;

hair 2-IV, V mesad of 1-IV, V; hair 1-III usually with 2-8 branches; 1-IV usu-

ally double (1-2); 3-II, III single, shorter than segment ITI; 5-IV-VI single,

not reaching beyond posterior margin of following segment; hair 9-VII single,

simple; 9-VIII usually with a strong main stem (1-2) and lateral branches of

varying length. Paddle. Margins with fringe; hair 1-P single; 2-P some-

times present.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; inner mouth brushes pectinate at tip;

head hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad

and mesad of 6-C; hair 5, 6,8,9,13-C single; 7,12-C double; 10-C usually

single (1-2); 11-C usually 3-branched (3-4); 14, 15-C usually double (2-3);

mentum with 11-12 teeth on each side. Thorax. Hair 1-P usually 3-branched

(2-3); 2-P single; 3-P double; 4-P 2-branched; 5,6-P single; 7-P double;

9-P single; 11-P usually double (1-2); 5, 7-M single; 6-M 3-branched; 8-M

with 4-5 branches; 9-M usually 3-branched (2-3); 10,12-M single, long, stout;

11-M single, small; 7-T usually 5-branched (5-7); 9-T usually double (2-3);

10, 11-T similar to those on mesothorax; 12-T much reduced. Abdomen.

Hair 6-I usually 4-branched (3-4); 7-I single; 6-II usually 3- branched (2-3);

7-II usually 3-branched (2-3); 6-II-V double; 6-VI single; 1-VII usually 2-

branched (2-3), long; 2-VII usually single; comb of 8-12 scales in a single

row, each scale with fine denticles or a fringe at the base of the apical spine;

sometimes a few (2-4) comb scales connected at base; pentad hair 2-VII

distant from 1-VII; 1,5-VIII with 3-4 branches; 3-VIII with 5-9 branches;

2,4-VIII single; siphon short about 2.5 times as long as wide, acus absent;

pecten teeth 10-14 in number, evenly spaced, each tooth with 2-4 basal den-

ticles; 1-S with 4-5 branches, inserted beyond last tooth and usually at mid-

dle of siphon; saddle incomplete; marginal spicules very small and inconspic-

uous; 1-X 2-branched; 2-X 2-branched; 3-X single; ventral brush with 4 pairs

of hairs on grid, each hair single except the 2 proximal ones usually double

(1-2), sometimes 4 proximal ones double; no precratal tufts; anal papillae

long and about 4 times as long as saddle, sausage-like.

TYPE DATA. Aedes Stegomyia) scutellaris malayensis Colless,

holotype male, allotype female (both from laboratory colony) in the Australian

National Insect Collection, Canberra; type locality: Palau Hantu, Keppel

Harbour, SINGAPORE (Colless 1962). Paratypes: 4 males, 4 females in U.

S. National Museum, 2 males, 2 females in British Museum (Natural History)

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia 47

London, all bearing same data as holotype male and allotype female A. s.

malayensis, Ex. Lab. Colony from Singapore, 1962).

DISTRIBUTION. 3,332 specimens examined: 710c, 8089, 3150 ter-

minalia, 242 terminalia, 82L, 21, 2p, 732 individual rearings (657 1, 732 p).

SINGAPORE. (1962, Ex. Lab. Colony, Colless), 9°, 69, 7% ter-

minalia; (I-1956), 1c, 1¢ terminalia; Pulau Hantu (HI-XII-1969-I-1970, W.

T. Chellappah), 231¢, 2059, 590 terminalia, 179 terminalia, 12 L, 257 in-

dividual rearings (251 1, 257 p); Pulau Blakang Mati (IV-1968, Ramalingam &

James), 1c, 59, 2 individual rearings (2p).

MALAYSIA. West Malaysia: Pahang- Kuantan (VIII-1966, Rama-

lingam & Ramakoishnan), 19; ([IX-1968, Wilson, Chia & Sulaiman), 1¢, 79,

2 individual rearings (21, 2p); Pulau Tioman (V-1966, Ramakoishnan), 33¢,

329, 25¢ terminalia, 8 L, 14 individual rearings (61, 14 p); Kg. Lamir (IX-

1968, Sulaiman, Chia & Wilson), 12; Kg. Cherating (X-1968, Chia & Seguan),

80°, 229, 1h terminalia, 3 L, 24 individual rearings (241, 24 p); Tg. Gelang

hovers Sulaiman & Chia), 120, 109, 6 terminalia, 15 individual rearings

121, 15 p). Trengganu-Pulau Perhentian Besar (V-1967, Ramachandran),

2%, 39, 1o¢ terminalia, 4 individual rearings (3 1, 4 p); Dungun (X-1968,

Sulaiman, Chia & Seguan), 3c, 72, 2c terminalia, 6 individual rearings (6 l,

6 p). Penang Is. -Telok Kumbor (I-1969, James, Chia & Sulaiman), 16¢,

372, 7c terminalia, 7 L, 29 individual rearings (26 1, 29 p); Pulau Perok,

West of Penang (IV-1949, L.A. Gibson Hill), 1¢, 1o¢ terminalia. Langkawi

Is. - (VII-1958, R. Traub), 19. "West Pulau, Penins, Coast Jarak (IV-1932,

E. Seimund), 49; Malacca Str. Pulau Jarak (KI-1958, W.W. MacDonald),

190°, 179, 12% terminalia, 27 individual rearings (271, 27 p)."

THAILAND. Siam (VIII-1933, O.R. Causey), 3c, 19, 3c termi-

nalia. Chon Buri: Khao Phuthabath (VII-1964, Kol & Sumeth), lo, 1o termi-

nalia; Top Mountain (VII-1964, Kol & Sumeth), 3c, 29, 3c terminalia; Ko Sz

Chang Is., Chagka Phong's cave (VII-1964, Sumeth), 2c, 20 terminalia; Ko

Si Chang Is., Thatanavong (VII-1964, Kol), 12; Ko Si Chung (VH-1964, Kol

& Sumeth), 19. Songkhla: Ton Nga Chang (III-1965, Peyton), 1c, 1¢ termi-

nalia, 1 individual rearing (1 p). Chanthaburi: Ban Laem Sing (XI-1965,

Sumeth), 1¢, 1c terminalia, 1 individual rearing (1 p). Nonthaburi: Bankok

(V-1968, ARU), 580, 309, 27% terminalia, 59 terminalia; (IJ-IV-1969,

SEATO), 860, 809, 42¢ terminalia, 97 individual rearings (89 1, 97 p).

Surat Thani: Ko Samui (X-1967, XII-1968, I-1969, SEATO), 22, 439, 22¢

terminalia, 60 individual rearings (311, 60 p). Tvat: Ko-Chang (XII-1967,

eit 1c, 39, 4 individual rearings i p). Phuket: Ban Huai Luk (11-1968,

Kol), 1o%, 392, 4 individual rearings (4 p). Prachuap Khiri Khan: Klong Van

Hill (11-1964, Kol), 1c, 22, 1c terminalia; Bo-Pia (IV-1968, SEATO), 42c,,

802, 29% terminalia, 20 L, 60 individual rearings (55 1, 60 p); Huai Yang

Phrach Khwa (IV-1968, SEATO), 3c, 119, 3c terminalia, 1 L, 13 individual

rearings (13 1, 13 p); (SEATO's Insectory material, III-IV-1970), 390, 479,

380 terminalia, 2? terminalia, 82 individual rearings (821, 82 p); (VIII-1970),

830, 1059, 2c terminalia, 31 L, 29 individual rearings (291, 29 p).

CAMBODIA. Kampot: Sihanoukville (XII-1966, J.M. Klein), 2c,

39. Kandal: Phnom-Penh, Ari Ksatr (VII-1967, J.M. Klein), 6°, 79, 2¢

terminalia; "Ari Gsatr" (V-1967, M. Delfinado), 19.

VIET NAM. Con Son: (VIII-1966, 20th PMU), 11¢, 189, 9% termi-

nalia, 21, 2p; (XI-1966, R. Hochman), 6c, 49, 6c terminalia. Da Nang:

Spanish Point (XII-1966, R.A. Woiff), 19. Khanh Hoa: Cam Ranh Bay (V-

1967, R. Hochman), 59. Binh Dinh: An Khe (IV-1967, R. Hochman), 19.

"Quang Tin, Nuoc Man" (V-1967, R. Hochman), 19.

TAIWAN. Orchid Island (V-1969, MAPS), 1c, 1c terminalia, 1

individual rearing (1 1, 1 p).

TAXONOMIC DISCUSSION. A. malayensis is a member of the

scutellaris subgroup. The adult is very similar to alcasidi, riversi and

scutellaris. in having the mid femur without median white line on anterior

48 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

surface, wing with minute basal spot of white scales on costa, hind tarsomere

5 all white. It is closer to viversz than to alcasidi and scutellaris in having

hind tarsomere 3 with basal 2/5 white banded and hind tarsomere 4 with basal

2/3 white banded. It can be separated from viversi by having hind tarsomere

1 with basal 1/4 white banded and hind tarsomere 2 with basal 1/3 white band-

ed; in viversi hind tarsomere 1 with basal 1/5 white banded and hind tarso-

mere 2 with basal 1/4 white banded.

The male terminalia of malayensis are very Similar to alcasidi,

hensilli and vriversi in having claspette with distal expanded part subtriangu-

lar in shape in lateral aspect (dissected claspette), sternal and tergal sides

not parallel but tapering and without an apicosternal angle. It is closer to

riversi than to alcasidi and hensilli because the apicotergal area of claspette

is without any distinctly long setae. It can be separated from viversi by hav-

ing claspette with 7-10 modified setae forming a prominent row on center of

sternal side and occupying about 1/2 of it; in rviversi claspette has 6-8 modi-

fied setae on center of sternal side, closer to sternal angle area than to api-

cotergal angle area and occupying about 2/5 of it.

The larvaof malayensis isvery similar to albopictus, alcasidi, viversi

and scutellaris inhaving no siphon acus, saddle incomplete, hair 2-X 2-branched

and 2- VII usually single (1-2). Itis closer to alcasidi and scutellaris but can be

separated from both by having hair 1-S usually inserted at middle of siphon,

pecten teeth 10-14 in number, each tooth with 2-4 basal denticles; in alcasidi

and scutellaris hair 1-S is usually inserted before middle of siphon. The

pupa of malayensis is very similar to albopictus, pseudalbopictus, alcasidi,

riversi and scutellaris in having hair 9-VI of about the same thickness as

9-V, less than twice as long as 9-V, hair 9-VII single and simple and 9- VHUI

reaching beyond fringe of paddle. It is closer to viversz than to any other

species in having hair 6-C much stouter than 7-C and about 3/4 of 7-C. It

can be separated from viversi by having hair 9- VOI usually with a strong

main stem (1-2) and lateral branches of varying length and hair 1-II with

many primary and secondary branches; in viversi 9- VIII usually has 2 main

stems (1-2), barbed, reaching beyond fringe of paddle and hair 1-II with very

few secondary branches.

A. malayensis is highly variable in both adult ornamentation and in

the immature stages. However, certain characters of the male terminalia

such as the shape of the claspette and the degree of development of modified

setae on the claspette are constant and unique. The shape of the claspette is

strikingly different from that of scutellaris, from which it can easily be sep-

arated by having the claspette with distal expanded part subtriangular in shape

in lateral aspect (dissected claspette), sternal and tergal sides not parallel

but tapering and without an apicosternal angle; in scutellaris the claspette has

distal expanded part square in shape in lateral aspect (dissected claspette),

sternal and tergal sides more or less parallel and an apicosternal angle is

present,

Colless (1962) considered malayensis to be a subspecies of scutellaris

on the basis of the laboratory hybridization experiments between the Malayan

form from Singapore and scutellaris from New Guinea. He stated in his

paper that the cross appeared to be a complete success in both directions.

However, I have seen his hybrids in the BM., namely, 14 adults (6c, 89)

and in the USNM. (2c") and all were from a one-way cross A. s. malayensis

(M.) X A. s. scutellaris (F.). Further hybridization experiments are de-

sirable.

A. malayensis is apparently a common species in the Southeast Asia

area. It is presently known from Taiwan, Viet Nam, Cambodia, Thailand,

W. Malaysia and Singapore. In Southeast Asia the immature stages are often

found in association with albopictus in the field and great care must be taken

in identifying them. The larva and pupa of malayensis can be distinguished

ye ie tenga by the diagnostic characters mentioned under the discussion

of the latter.

Huang: Aedes (Stegomyia) scutellavis group in Southeast Asia 49

BIOLOGY. The immature stages of malayensis have been collected

mainly in tree holes and spathes in Singapore and in rock pools in Malaya.

They have also been found in a bamboo stump, a coconut shell and artificial

containers in Malaya and in rock holes, rock pools, water jars and bamboo

cups in Thailand. The specimen from Orchid Island, Taiwan was found ina

tree hole while the specimens from Con Son, Viet Nam were found in coco-

nut shells. The immature stages were associated with albopictus in Singapore,

Malaya, Thailand and Taiwan. The adult females have been taken biting man

in Bo-Pia, Prachuap Khiri Khan, Thailand.

AEDES STEGOMYIA) PAULLUSI STONE & FARNER

(Figs. 20E, F,J,K, 9 abdomen, thorax, mid leg;

29, & terminalia, pupa; 30, larva)

Aedes Stegomyia) paullusiStone & Farner 1945, Proc. biol. Soc. Wash. 58:

155 (o*, 9); Knight & Hull 1952, Pacif. Sci. 6(2): 178 («*, 9, L); Marks

1954, Bull. Br. Mus. (nat. Hist.) Ent. 3(10): 376, 383, Pl. 18 (c*,

ee Bonne-Wepster 1954, Spec. Publ. R. trop. Inst. Amsterdam,

: 84,

MALE. Head, Proboscis dark scaled, with pale scales on ventral

side, longer than fore femur; palpus dark, as long as proboscis, with white

basal band on each of segments 2-5; those on segments 4-5 incomplete dor-

sally; segment 5 with white basal band on ventral side unusually long, occu-

pying about basal 2/3 or more; segments 4-5 subequal, slender, upturned,

and with only a few short hairs; antenna plumose, shorter than proboscis.

Thorax. Scutum with narrow dark scales and prominent median stripe of

Similar white ones, stripe narrows posteriorly and forks at beginning of pre-

scutellar space; on each side a posterior dorsocentral white line which does

not reach to middle of scutum; a few narrow white scales on lateral prescutal

area and on scutal angle area; a supraalar line of broad white scales present;

posterior pronotum with narrow dark scales on upper portion and with broad

white scales on lower portion forming a white stripe instead of a white patch;

postspiracular area without scales; subspiracular area without scales; mes-

epimeral scale patches separated or sometimes narrowly connected. Wing.

Dark scales on all veins except for minute basal spot of white scales on costa;

first forked cell 1.5 times as long as its stem. Halter. With dark scales.

Legs. Fore femur dark anteriorly, paler posteriorly; mid femur with median

white line on anterior surface; hind femur anteriorly with a broad white stripe

which widens at base and is narrowly separated from apical white scale patch;

fore and mid tarsi with basal white bands on tarsomeres 1-2; hind tarsus with

basal white bands on tarsomeres 1-4, the ratio of length of white band tototal

length of tarsomere is 1/3, 2/5, 1/2 and 2/3, tarsomere 5 all white. Abdo-

men. Abdominal segment I with white scales on laterotergite; tergum II dark

dorsally, with lateral white spots only; terga II-VI each with sub- basal white

band which narrows dorsally and turns abruptly caudad at lateral margin and

ends there near the large oblique lateral spot or attached to it; sterna II-VI

with basal white bands. 'Terminalia.. Basimere about 3.5 times as long as

wide; with a patch of hairs on basomesal area of dorsal surface; mesal sur-

face membranous; claspette simple, truncate, with numerous setae, with

several long, stout setae on tergal side and 4 spine-like setae on sternal

side; distimere simple, elongate, as long as basimere, with a spiniform pro-

cess and a few hairs near apex; tergum IX with middle truncated and with a

hairy lobe on each side.

FEMALE. Essentially as in male, differing inthe following repsects:

palpus 0. 2 length of proboscis, with white scales on more than apical half.

50 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

Wing with first forked cell about twice as long as its stem. Abdominal terga

II- VI each with sub-basal white band which narrows dorsally and turns abrupt-

ly caudad at lateral margin and there ends near or attached to lateral white

spot which is turned dorsomesally; sometimes tergum II without complete

band and sometimes with lateral spots only; tergum VII with band broken on

each side of median patch; segment VIII largely retracted.

PUPA. Cephalothorax. Trumpet 3.5 times as long as width at mid-

dle; hair 1,3-C single, longer than 2-C; 2-C single; 4-C single or double; 5,

6-C single; 6-C much stouter and slightly longer than 7-C, 7-C usually double

(1-2); 10-C usually with 3-4 branches, mesad and caudad of 11-C; 11-C sin-

gle. Abdomen. Hair 1-I well developed, with more than 10 branches, den-

dritic; 2-I single; 3-1 single, long; 2,3-I not widely separated, distance be-

tween them as distance between 4,5-I; hair 1-II usually with 10-11 branches;

2-Il mesad of 3-II; 2-IV, V mesad of 1-IV, V; hair 1-II,IV usually double (1-

2); 3-II, III single, shorter than segment III; hair 5-IV-VI single, not reach-

ing beyond posterior margin of following segment; hair 9-VII single and barb-

ed or split at tip; hair 9- VIII usually with 2 main stems, barbed, reaching

beyond fringe of paddle. Paddle. Margins with fringe; hair 1-P single.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A in-

serted near middle of shaft, single; inner mouth brushes pectinate at tip; head

hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad and

mesad of 6-C; hair 5,6,8,9,10,13-C single; 7,12,15-C usually double; 11-C

usually 5-branched (4-5); 14-C usually 3-branched (2-3); mentum with 9-10

teeth on each side. Thovax. Hair 1-P 3-branched; 2-P single; 3-P double;

4-P 2-branched; 5,6-P single; 7-P double; 9-P single; 11-P single; 5, 7-M

single; 6-M with 3-4 branches; 8-M 4-branched; 9-M 3-branched; io, 12-M

single, long, stout; 11-M single, small; 7-T usually 5-branched (5-6); 9-T

usually double; 10,11-T similar to those on mesothorax; 12-T much reduced.

Abdomen. Hair 6-1 with 3-4 branches; 7-I single; 6-II 3-branched; 7-II 3-

branched; 6-III-V double; 6-VI single; 1-VII usually 3-branched; 2-VII usually

3-branched (2-3); comb of 8-10 scales in single row, each scale with fine

denticles or fringes at base of apical spine; pentad hair 2, 4-VIII single; 1-VUI

with 3-4 branches; 3- VIII with 5-7 branches; 5-VIII with 3-5 branches; siphon

short, about twice as long as wide, acus absent; pecten teeth 9-13, evenly

spaced, each tooth with 1 large and occasionally 1 or 2 very small basal den-

ticles; 1-S with 3-4 branches, inserted beyond last tooth and in line with

teeth; saddle incomplete; marginal spicules very small and inconspicuous;

1-X 2-branched; 2-X usually 3-branched; 3-X single; ventral brush with 4

pairs of hairs on grid, each hair single; no precratal tufts; anal papillae about

2.5 times as long as saddle, sausage-like.

TYPE DATA. Aedes (Stegomyia) paullusi Stone & Farner, holotype

male in U.S. National Museum, Washington, D.C.; type locality: San Antonio,

Samar, PHILIPPINE ISLANDS, 6-XII-1944 (J.H. Paullus, 51). Paratypes:

1 male, N'goles, Calicoan Island, Philippine Islands, 27-I-1945 (J.H. Paullus,

103); 1 female, with same data as holotype; 1 female, Baras, Calicoan Island,

Philippine Islands, 24-I-1945 (J.H. Paullus, 100); 1 female, small island

near Calicoan Island, Philippine Islands, 12-II-1945 a H. Paullus, 110); 1

female, Abuyog, Leyte, Philippine Islands, XI-1944 (O.H. Graham) in U.S.

National Museum; 2 males (# 6053), Taroena, Sangir Islands, IM-1928 (S.L.

Brug & de Rook) in British Museum (Natural History), London.

DISTRIBUTION. 659 specimens examined: 201<, 2079, 104c termi-

nalia, 42 terminalia, 6 L, 11, 1 p, 68 individual rearings (671, 68 p).

PHILIPPINES. Samar: San Antonio (XII-1944, J.H. Paullus), 3c,

4°, 2¢ terminalia; Pintanahon (IV-1945, Rozeboom, Knight & Laffoon), 2c,

1¢ terminalia, 4 L, 2 individual rearings (21, 2 p); Bulusao (V-1945, Roze-

boom, Knight & Laffoon), 2c, 19, 2¢ terminalia, 2 individual rearings (2 1,

2p); Ducong (V-1945, Rozeboom, Knight & Laffoon), 19, 1 individual rearing

(11, 1 p); Osmena (IV-IX-1945, Rozeboom, Knight & Laffoon), 30c¢, 279, 19¢

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 51

terminalia, 21 individual rearings (21 1, 21 p); Tank Farm (IV-1945, Roze-

boom, Knight & Laffoon), 19; Samar (III-1945, Rozeboom, Knight & Laffoon),

1¢, 1o terminalia; Leyte: Abuyog (XI-1944, O.H. Graham), 12; Tacloban-

een n H.R. Roberts), 9c, 279, 60% terminalia, 6 individual rearings

61, 6p); (V-1945, E.S. Ross), 60, 39, 3c terminalia. Dagami- (VII-1945,

H.R. Roberts), 20c, 169, 19% terminalia, 3 individual rearings (3 1, 3 p);

(VII-1945, Rozeboom, Knight & Laffoon), 19. Palo (V-1945, H.R. Roberts),

19; Lagolago Baybay (I-II-1945, H.R. Roberts), 50, 29, 3c terminalia; Mt.

Lobi 1000' (VIII-1945, Rozeboom, Knight & Laffoon), 20; Carigara (XI-1944,

E.S. Ross), 2c, 22; Mahaplag (VII-1964, M. Delfinado), 7", 72, 50 termi-

nalia. Mindoro: San Jose (III-V-1945, E.S. Ross), 40%, 492, 3c terminalia;

Calicoan: Baras (I-1945, J.H. Paullus), 19; (11-1945, J.H. Paullus), 19;

N'goles (I-1945, J.H. Paullus), 1c, 1o¢ terminalia. Palawan: Irahuan R.

(VI-1945, Rozeboom, Knight & Laffoon), 39, 1 individual rearing (11, 1 p);

Puerto Princesa (VI-1945, Roseboom, Knight & Laffoon), 10°; Quezon (XII-

1967-II-1968, Alcasid's team), 29; Iwahig (XI-1968, Mantubig), 29, 2 indi-

vidual rearings (11, 2 p); Balabac I., Cape Melville (VI-1945, Rozeboom,

Knight & Laffoon), 19. Mindanao: Lanao- Dapau, Porque R. (IV-1931, W.V.

King), 2%, 49, 1o terminalia; Kolambugan, Titunod (V-1931, W.V. King),

60°, 49, 3h terminalia. San Ramon (IX-XI-1945, Rozeboom, Knight &

Laffoon), 3%, 119, 1o% terminalia, 9 individual rearings (9 1, 9 p); Pasanonco

(IX-1945, Rozeboom, Knight & Laffoon), 2c’, 59, 4 individual rearings (4 1,

4p); Parang (VI-1945, J.H. Paullus), 2c’, 49; Kabakan (V-1945, R. Staples),

13c, 9c terminalia; (V-XI-1945), 4c’, 79; (1945, Rozeboom, Knight & Laffoon),

30°, 69; Jolo Jolo I. (IX-1945, Rozeboom, Knight & Laffoon), 2c, 19, 1 in-

dividual rearing (11, 1 p); Philippine Islands (V-1945), 20, 39; (V-1946), 8c,

a 7s terminalia, 49 terminalia; (VI-1945, Rozeboom, Knight & Laffoon),

Co.

MALAYSIA. West Malaysia: Selangor- 15 ml. Ulu Gombak (III-

IV-1959, W.W. MacDonald), 5c, 79, 5c terminalia, 10 individual rearings

ar 1, 10 p); Ulu Gombak 16 (XI-1965, Ramalingam's team), 1 L; Bt. Kutu

V-1968, Ramalingam's team), 192; Pahang- Gunong Benom (IX-1968, Rama-

lingam), 1o, 79, 1c terminalia, 1 L, 2 individual rearings (21, 2p). East

Malaysia: Sabah- Kudat (VI-VOI-1966, F.Y. Cheng), 29; Balembangan Is.,

Kok Simpul (X-1965, F.Y. Cheng), 2c, 19, 2% terminalia, 1 L, 1 p, 1 indi-

vidual rearing (1 1, 1 p). -

INDONESIA. Sangir Islands: Taroena (III-1928, S.L. Brug & de

Rook), 110, 49, 2c terminalia; (27-I-1942), 180, 99, 1c terminalia.

Celebes: Lindoemeer (II-1937, Brug & Tesch), 2c, 29, 1c terminalia;

Halino (VI-1937, Brug), 2c’, 29; Kalawara (I-II-1937, Brug), 4°, 59, 2c

terminalia; Kabaena- (V-1935, Brug), 3o, 1o terminalia; (VI-1937, Brug),

19. Ambon Island- (27-1-1938), 7, 49; Waai (XII-1965-1966, A.M.R.

Wegoner), 3c’, 19, 3c terminalia, 3 individual rearings (3 1, 3 p).

TAXONOMIC DISCUSSION. A. paullusi is a member of the scutellaris

subgroup. The adult is very similar to alorensis in having the mid femur with

a median white line on anterior surface and wing with minute basal spot of

white scales on costa. It can easily be separated from alorensis by having a

few narrow white scales on lateral prescutal area and on scutal angle area;

in alorensis there are no such scales. When scutal markings are rubbed off, —

paullusi can easily be mistaken for alorensis but can still be recognized by

the characteristic abdominal bandings.

The male terminalia of paullusi have the claspette truncate, with the

apical surface distinctly oval in shape, bearing numerous Setae and several

long, stout ones on tergal side and with 4 spine-like setae on sternal side, |

thus differing from all other species that have been described in this group.

The larva of paullusi is very similar to alcasidi, malayensis, viverst

and scutellaris but can be distinguished from them by having hair 2-X 3- branched,

2-VII usually with 3 (2-3) branches; pecten tooth with 1 large and occasionally

52 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

1 or 2 very small basal denticles; in alcasidi, malayensis, riversi and

scutellaris hair 2-X 2-branched, 2-VII usually single (1-2). The pupa of

paullusi resembles alcasidi and scutellaris but can be separated from both

by having hair 6-C usually slightly longer than 7-C, hair 1-II usually with 10-

11 branches, which rise from a common stem at base, paddle margins with

fringe extending close to base and occupying more than apical 3/4 of paddle;

in alcasidi and scutellaris hair 6-C usually about as long as 7-C, hair 1-II

branched, usually without a distinct common stem at base, paddle margins

with fringe on less than apical 3/4 of paddle.

A. paullusi is apparently a common species in the Indomalayan area

and extends into the western fringe of the Papuan area. It is presently known

from the Philippines, Sangi Islands, Taroena, Celebes, Kabaena, Sabah, W.

Malaysia and Ambon.

BIOLOGY. The immature stages of paullusi have been found mainly

in rock holes, coconut shells, coconut husks, coconut fronds and bamboo

stumps in the Philippines. It has also been found in a tree hole, a hole ina

log and in a fallen abaca leaf in the Philippines. The immature stages were

found in a small artificial container in rainforest in Pahang, Malaysia, The

specimens from Lindoemeer, Celebes were found in bamboo stumps and

those from Waai, Ambon Island in coconut shells. The adult females have

been taken biting buffalo in a village in Kudat, Sabah

AEDES STEGOMYIA) RIVERSI BOHART & INGRAM

(Figs. 26A, B, claspette; 31A,B,E,G,H, 9? abdomen, hind leg, &* abdomen;

32, o terminalia, pupa; 33, larva)

Aedes (Stegomyia) riversi Bohart & Ingram 1946, J. Wash. Acad. Sci. 36:

90 (o*, 29, L*); Bohart & Ingram 1946, U.S. Navmed 1055 :65 (c*, 9,

a Aan, 1954, Bull. Br. Mus. (nat. Hist.) Ent. 3(10): 383, Pl.

18 (o*, 2*).

MALE. Head. Proboscis dark scaled, with pale scales on ventral

side, longer than fore femur; palpus dark, slightly shorter than proboscis,

with white basal band on each of segments 2-5; those on segments 4-5 incom-

plete dorsally; segments 4-5 subequal, slender, upturned and with only a few

short hairs; antenna plumose, shorter than proboscis. ‘Thorax. Scutum with

narrow dark scales and prominent median stripe of similar white ones; stripe

narrows Slightly posteriorly and forks at beginning of prescutellar space; on

each side a posterior dorsocentral yellowish line which does not reach to

middle of scutum; a supraalar line of broad white scales present; posterior

pronotum with narrow dark scales on upper portion and with broad white

scales on lower portion forming a white stripe instead of a white patch; post-

spiracular area without scales; subspiracular area without scales; lower mes-

epimeral scale patch large and connected to upper mesepimeral scale patch.

Wing. Dark scales on all veins except for minute basal spot of white scales

on costa; first forked cell 1.5 times as long as its stem. Halter. With dark

scales. Legs. Fore and mid femora dark anteriorly, paler posteriorly; hind

femur anteriorly with broad white stripe which widens at base and is sepa-

rated from apical white scale patch; fore and mid tarsi with basal white bands

on tarsomeres 1-2; hind tarsus with basal white bands on tarsomeres 1-4,

the ratio of length of white band to total length of tarsomere is 1/5, 1/4, 2/5

and 2/3, tarsomere 5 all white or sometimes with a few dark scales on api-

cal ventral side. Abdomen. Abdominal segment I with white scales on later-

otergite; tergum II dark dorsally, with lateral white spots only; tergum II

dark dorsally, with lateral white spots only or sometimes with median spot

as well; terga IV-VI each with sub-basal white band which is rather narrow

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 53

and is connected to the large lateral spots which are turned dorsomesally,

sometimes terga IV-VI each with sub-basal narrow band at middle or with

median spot and with lateral spots which are turned dorsomesally, or some-

times terga IV-VI dark at middle with lateral spots which are turned dorso-

mesally as in the form of incomplete bands; tergum VI with lateral white

spots only; sternum VII largely covered with white scales. Terminalia.

Basimere 3.5 times as long as wide; with a patch of hairs on basomesal area

of dorsal surface; mesal surface membranous; claspette simple, with distal

expanded part subtriangular in shape, sternal and tergal sides not parallel

but tapering, with 6-8 modified setae on center of sternal side, closer to

sternal angle area than to apicotergal angle area and occupying about 2/5 of

it; distimere simple, elongate, as long as basimere, with a spiniform pro-

cess and a few hairs near apex; tergum IX with middle rounded and with a

hairy lobe on each side.

FEMALE. Essentially as in male, differing inthe following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Wing with

first forked cell about twice as long as its stem. Abdominal terga II-III al-

ways dark dorsally, with lateral white spots only; tergum IV with incomplete

sub-basal band only or sometimes with small median spot as well; terga V-

VII each with sub-basal white band which is rather narrow and is connected

to lateral spots which are turned dorsomesally, sometimes terga V-VII with-

out complete bands or sometimes terga V-VII with incomplete sub-basal

bands only; segment VIII largely retracted.

PUPA. Cephalothorax. Trumpet 3 times as long as width at middle;

hair 1,3-C single, longer than 2-C; 2-C single; 4-C usually double; 5-C dou-

ble; 6-C single, much stouter than 7-C, slightly shorter than 7-C; 7-C usu-

ally single (1-2); 10-C usually with 2-4 branches, mesad and caudad of 11-C;

11-C single. Abdomen. Hair 1-I well developed, with more than 10 branches,

dendritic; 2-I single; 3-I single, long; 2,3-I not widely separated, distance

between them as distance between 4, 5-I; hair 1-II usually with 7-12 branches;

2-II mesad of 3-II; hair 2-IV, V mesad of 1-IV, V; 1-III usually double (2-6);

1-IV usually double (1-2); hair 3-II, III single, shorter than segment IIT; hair

o-IV-VI single, not reaching beyond posterior margin of following segment;

9-VII single, simple; 9- VIII usually with 2 main stems (1-2), barbed, reach-

ing beyond fringe of paddle. Paddle. Margins with fringe; hair 1-P single.

LARVA. Head. Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; inner mouth brushes pectinate at tip;

head hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad

and mesad of 6-C; hair 5,6, 8,9,10,13-C single; 7,12,14-C usually double;

11-C usually with 3-4 branches; mentum with 11-12 teeth on each side.

Thorax. Hair 1-P with 2-3 branches; 2-P single; 3-P double; 4-P 2-branched;

5,6-P single; 7-P double; 9-P single; 11-P single; 5, 7-M single; 6-M 3-

branched; 8-M 5-branched; 9-M 3-branched; 10,12-M single, long, stout;

11-M single, small; 7-T with 4-5 branches; 9-T usually double; 10,11-T

Similar to those on mesothorax; 12-T much reduced. Abdomen. Hair 6-I

3-branched; 7-I single; 6-II with 2-3 branches; 7-II double; 6-III-V double;

6-VI single; 1-VII usually with 2-3 branches; 2-VI single; comb of 10-14

scales in a single row, each scale with fine denticles or fringes at base of

apical spine; comb scales often with apical spine split at tip; sometimes 2-4

comb scales connected at base; pentad hair 2, 4-VIII single; 1,5-VIII with 3-4

branches; 3-VIUI with 5-7 branches; siphon short, about twice as long as wide,

acus absent; pecten teeth 10-21 in number, evenly spaced, each tooth with 1

large and 1-2 small basal denticles; 1-S with 3-4 branches, inserted beyond

last tooth and beyond middle of siphon; saddle incomplete; marginal spicules

very small and inconspicuous; 1-X 2-branched; 2-X 2-branched; 3-X single;

ventral brush with 4 pairs of hairs on grid, each hair single except 2 proxi-

mal ones usually double (1-2); no precratal tufts; anal papillae about twice as

long as saddle, sausage- like.

54 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

TYPE DATA. Aedes (Stegomyia) riversi Bohart & Ingram, holotype

male in U.S. National Museum, Washington, D.C.; type locality: Chizuka,

Okinawa (RYUKYU-RETTO), IX-1945 (R. Bohart & R. Ingram). Paratypes

(except as indicated, all collected by R. Bohart & R. Ingram): 6 males, 9 fe-

males, Chizuka, Okinawa, IX-1945; 1 male, 3 females, Chizuka, Okinawa,

IX-6-1945; 2 females, Chizuka, Okinawa, VIII- 24-1945; 2 males, 1 female,

Chizuka, Okinawa, VIII-1945; 1 male, 6 females, Hentona, Okinawa, IX-1945

(C.L. Harnage); 3 males, 2 females, Hentona, Okinawa, IX-1945; 1 male, 5

females, Shana Wan, Okinawa, IX-1945; 1 female, Shana Wan, Okinawa, IX-

4-1945 in U.S. National Museum; 1 female, Chizuka, Okinawa, IX-1945 in

British Museum (Natural History), London. Also paratype larvae: 6 larvae

on 3 slides, Chizuka, Okinawa, VII-IX-1945; 4 larvae on 2 slides, Shana

Wan, Okinawa, IX-13-1945 in U.S. National Museum.

DISTRIBUTION. 807 specimens examined: 910, 2139, 65< terminalia,

99 terminalia, 35 L, 197 individual rearings (197 1, 197 p).

RYUKYU ISLANDS. Okinawa: Chizuka (VII-IX-1945, R. Bohart &

R. Ingram), 10c, 209, 8% terminalia, 6 L; Hentona (IX-1945, C.L. Harnage)

4c, 99, 2c terminalia; ([X-1945, R. Bohart & R. Ingram), 4c, 29, 3c ter-

minalia; Shana Wan (IX-1945, R. Bohart & R. Ingram), 2c, 69, 1c termi-

nalia, 4 L; Iviomote: Shirahama (V-1968, A.B. Silagan), 29, 2 individual

rearings (21, 2p); 15 ml. N. of Ohara Village (XII-1968, M. Nakama), 29,

2 individual rearings (21, 2p); 2 ml. W. of Yabu Village (XII-1968, M.

Nakama), 2c, 129, 1c terminalia, 14 individual rearings (141, 14 p); (XII-

1968, A.B. Silagan), 4c, 29, 3o terminalia, 6 individual rearings G i, 6p):

2ml. S. of Yabu (XII-1968, M. Nakama), 1c, 29, 1o terminalia; Komi (II-

1970, I.V. Villanueva), 1°, 39, 4 individual rearings (41, 4 p); Yaeyama:

Kabira (X-1968, G. Takaesu), 29, 2 individual rearings (21, 2p); (X-1968;

A.B. Silagan), 3c, 89, 2% terminalia, 11 individual rearings (11 1, 11 p);

Yarabu Dake (X-1968, G. Takaesu), 4c, 149, 2c’ terminalia, 18 individual

rearings (181, 18 p); (X-XII-1968, A.B. Silagan), 5c, 259, 2c terminalia,

30 individual rearings (30 1, 30 p); Haskino (X-1968, G. Takaesu), 119, 11

individual rearings a 1, 11 p); Inota (X-1968, G. Takaesu), 49, 4 individual

rearings (41, 4 p); (XII-1968, A.B. Silagan), 2c, 19, 2% terminalia, 3 indi-

vidual rearings (3 1, 3 p); (XII-1968, M. Nakama), 19, 1 individual rearing

(1 1, 1 p); Ishigaki- Yarabu Dake (I-1970, I.D. Cocklin), 1c, 29, 1o¢ termi-

nalia, 3 individual rearings (3 1, 3 p); (I-1970, I. V. Villanueva), 3c, 79, 3¢

terminalia, 10 individual rearings (16 1, 10 p); (I-1970, T.S. Bolingust), 9c,

219, 7 terminalia, 30 individual rearings (301, 30 p); (I-1970, A.B. Comp),

2°, 2 individual rearings (21, 2 p); Inota III (I-1970, I.V. Villanueva), 8¢,

139, 60% terminalia, 21 individual rearings (21 1, 21 p); (I-1970, T. S.

Bolingust), 3c, 39, 6 individual rearings (61, 6 p); 5 ml. N. of Arakawa

(I-1970, I.V. Villanueva),19, 1 individual rearing (1 1, 1 p); Shirako (I-1970,

T.S. Bolingust), 4c, 129, 4c terminalia, 9° terminalia, 16 individual rear-

ings (161, 16 p); Yaeyama (I-1970), 25 L. Ishigaki-Jima: (X-XI-1961,

Sasa), 9c, 229, 6c terminalia. Amami: (V-1962, Sasa), 120, 49, 11¢ ter-

minalia.

TAXONOMIC DISCUSSION. A. rviversi is a member of the scutellaris

Subgroup. The adult is very similar to alcasidi, malayensis and scutellaris

in having the mid femur without median white line on anterior surface, wing

with minute basal spot of white scales on costa and hind tarsomere 5 all white,

It is closer to malayensis than to alcasidi and scutellaris in having hind tarso-

mere 3 with basal 2/5 white banded and hind tarsomere 4 with basal 2/3 white

banded. It can be separated from malayensis by having hind tarsomere 1 with

basal 1/5 white banded and hind tarsomere 2 with basal 1/4 white banded; in

malayensis hind tarsomere 1 has basal 1/4 white banded and hind tarsomere

2 with basal 1/3 white banded.

The male terminalia of viversi are very similar to alcasidi, malay-

ensis and hensilli in having claspette with distal expanded part subtriangular

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia 5}9)

in shape in lateral aspect (dissected claspette), sternal and tergal sides not

parallel but tapering and without apicosternal angle. It is closer to malay-

ensis than to alcasidi and hensilli in having apicotergal area of claspette

without distinctly long setae. It can be separated from malayensis by having

the claspette with 6-8 modified setae on center of the sternal side, closer to

sternal angle area than to apicotergal angle area and occupying about 2/5 of

it; in malayensis the claspette has 7-10 modified setae forming a prominent

row on center of sternal side and occupying 1/2 of it.

The larva of riversi resembles albopictus, alcasidi, malayensis and

scutellaris in having no siphon acus, saddle incomplete, hair 2-X 2-branched

and 2-VII usually single (1-2). It can be separated from them by having hair

1-VIL with 2-3 branches, siphon about twice as long as wide, pecten teeth 10-

21, closely arranged in a line, hair 1-S usually inserted beyond middle of

Siphon and comb scales sometimes with apical spine split at tip. The pupa of

riversi is very Similar to albopictus, pseudalbopictus, alcasidi, malayensis

and scutellaris in having hair 9-VI about same thickness as 9-V, less than

twice as long as 9-V, hair 9-VII single and simple, hair 9-VIII reaching be-

yond fringe of paddle. It is closer to malayensis than to any other species

in having hair 6-C much stouter than 7-C and about 3/4 as long as 7-C. It

can be separated from malayensis by having hair 9-VIII usually with 2 main

stems (1-2), barbed, reaching beyond fringe of paddle, hair 1-II with very

few secondary branches; in malayensis hair 9-VII usually with a strong main

stem (1-2) and lateral branches of varying length, hair 1-II with many pri-

mary and secondary branches.

A. riversi is apparently restricted to the Ryukyu Islands. The imma-

ture stages are often found in association with albopictus and downsi in the

field. Great care must be taken in identifying them. The larva and pupa of

riversi can be separated from albopictus by the diagnostic characters men-

tioned under the discussion of albopictus. The immature stages are markedly

different from downsi, The larva of viversi can be separated from downsi by

having hair 1-VII usually with 2 (2-3) branches, long, at least 2.5 times as

long as 5-VII and hair 2-VII usually single; in downsi hair 1-VII usually has

4 branches which are short and less than twice as long as 5-VII and hair 2-VII

3-branched. The pupa of viversi can easily be distinguished from downsi by

having hair 9-VI slender, about same thickness as 9-V, less than twice as

long as 9-V, single, simple and hair 9-VII single and simple; in downsi hair

9-VI stout, much stouter than 9-V, at least twice as long as 9-V, usually

ee and barbed, hair 9-VII usually single and barbed or with 2 branches

at tip.

BIOLOGY. The larvae of viversi have been found mainly in rock

holes, tree holes and cut bamboo in Okinawa. The immature stages from

Ryukyu Islands, Yaeyama and Iriomote were found mainly in tree holes. The

specimens from Ishigaki- Jima were found in tree holes and rock holes. The

immature stages were associated with aibopictus and downsi. The adult fe-

males have been taken biting man in Okinawa.

AEDES (SSTEGOMYIA) SCUTELLARIS (WALKER)

(Figs. 25A,B, claspette; 31 F, hind leg; 34, ¢;

35, & terminalia, pupa; 36, larva)

Culex pen Doleschall 1858, (non Schrank, 1781), Nat. Tijd. Ned.-Ind.

Culex scutellaris Walker 1859, Proc. Linn. Soc. Lond. 3: 77 (2).

Culex zonatipes Walker 1861, Proc. Linn, Soc. Lond, 5: 229 (9).

Aedes (SStegomyia) scutellaris (Walker), Edwards 1932, Genera Insectorum,

Fasc. 194 :165 (synonymized Culex variegatus and Culex zonatipes);

56 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

Taylor 1934, Proc. Linn, Soc. N.S.W. 59:235; Stone 1947, Proc.

ent. Soc. Wash. 49(3): 85; Penn 1949, Pacif. Sci. 3: 56 (P*);

Marks 1954, Bull. Br. Mus. (nat. Hist.) Ent. 3(10): 383, Pl. 18

(o°*, 2*); Bonne-Wepster 1954, Spec. Publ. R. trop. Inst.

Amsterdam 111: 79 (¢, 9*, L*); Belkin 1962, Mosq. South Pacific,

Vol. I: 474-475 & Vol. II: 331 (o*, 2); Huang 1969, Proc. ent. Soc.

Wash, 71(4): 472 (o*) (topotypic <).

Aedes (Stegomyia) scutellaris scutellaris (Walker), Colless 1962, Proc.

Linn. Soc. N.S.W. 87: 313 (o*, 2) (to ssp. status).

MALE. Head. Proboscis dark scaled, slightly longer than fore fe-

mur; palpus dark, as long as proboscis, with white basal band on eachof seg-

ments 2-5; those on segments 4-5 incomplete dorsally; segments 4-5 subequal,

slender, upturned, and with only a few short hairs; antenna plumose, shorter

than proboscis. Thovax. Scutum with narrow dark scales and prominent

median stripe of similar white ones; stripe narrows slightly posteriorly and

forks at beginning of prescutellar space; on each side a posterior dorsocen-

tral pale yellowish line which does not reach to middle of scutum; a supra-

alar line of broad white scales present; posterior pronotum with narrow dark

scales on upper portion and with broad white scales on lower portion forming

a white stripe instead of a white patch; postspiracular area without scales;

subspiracular area with or without scales; mesepimeral scale patches nar-

rowly connected, sometimes well separated. Wing. Dark scales on allveins

except for minute basal spot of white scales on costa; first forked cell 1.5

times as long as its stem. Halter. With dark scales. Legs. Fore and mid

femora dark anteriorly, paler posteriorly; hind femur anteriorly with broad

white stripe which widens at base and is separated from apical white scale

patch; fore and mid tarsi with basal white bands on tarsomeres 1-2; hind tar-

sus with basal white bands on tarsomeres 1-4, the ratio of length of white

band to total length of tarsomeres is 1/3, 2/5, 1/2 and 3/4, tarsomere 5 all

white. Abdomen. Abdominal segment I with white scales on laterotergite;

tergum II always dark dorsally, with lateral white spots only; terga II-VI

each with sub-basal white band which is connected to lateral spots, some-

times tergum III with sub-basal median spot and with lateral spots which are

turned dorsomesally; tergum VII with lateral white spots only; sternum VIII

largely covered with white scales. Terminalia. Basimere 3.5 times as long

as wide; with patch of hairs on basomesal area of dorsal surface; mesal sur-

face membranous; claspette simple, with distal expanded part square in shape,

sternal and tergal sides more or less parallel, apicosternal angle present

and with 5 or 6 modified setae close to apicosternal angle area; distimere

simple, elongate, as long as basimere, with a spiniform process and a few

hairs near apex; tergum IX with middle rounded and with a hairy lobe on each

Side.

FEMALE. Essentially as in male, differing inthe following respects:

palpus 0. 2 length of proboscis, with white scales on apical half. Wing with

first forked cell about twice as long as its stem. Abdominal terga II-III al-

ways dark dorsally, with lateral white spots which are turned dorsomesally;

sometimes tergum III with sub-basal median spot as well; terga IV- VI each

often with sub-basal white band which is connected to lateral spots or some-

times tergum IV with sub-basal median spot and with lateral spots which are

turned dorsomesally; or sometimes tergum IV with incomplete sub-basal

band only; segment VIII entirely retracted.

PUPA. Cephalothorax. Trumpet 3.5 times as long as width at mid-

dle; hair 1,3-C single, longer than 2-C; 2-C usually double (1-2); 4-C usu-

ally double (1-2); 5-C usually 2-branched (2-3); 6-C single, much stouter than

7-C, about as long as 7-C; 7-C usually double (1-2); 10-C usually with 2-4

branches, mesad and caudad of 11-C; 11-C single. Abdomen. Hair 1-I well

Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 57

developed, with more than 10 branches, dendritic; 2-I single; 3-I single,

long; 2, 3-I not widely separated, distance between them as distance between

4,5-I; 1-II usually with 7-9 branches; hair 2-IV, V mesad of 1-IV, V; 1-III

usually with 2-6 branches; 1-IV usually double (2-3); hair 3-II, III single,

shorter than segment III; 5-IV-VI single, not reaching beyond posterior mar-

gin of following segment; 9-VII single, simple or split at tip; 9- VIII usually

with 2 main stems (2-3) and barbed, reaching beyond fringe of paddle. Pad-

dle. Margins with fringe; hair 1-P single.

LARVA. Head, Antenna 0.5 length of head, without spicules; 1-A

inserted near middle of shaft, single; inner mouth brushes pectinate at tip;

head hair 4-C well developed, branched, closer to 6-C than 5-C, cephalad

and mesad of 6-C; hair 5,6,8,9,10,13-C single; 7,11-C usually with 2-3

branches; 12,14-C double; 15-C usually 3-branched (2-3); mentum with 11-12

teeth on each side. Thovax. Hair 1-P usually 3-branched (2-3); 2-P single;

3-P double; 4-P 2-branched; 5,6-P single; 7-P double; 9-P single; 11-P usu-

ally single (1-2); 5, 7-M single; 6-M 3-branched; 8-M with 4-5 branches; 9-M

3-branched; 10,12-M single, long, stout; 11-M single, small; 7-T usually 5-

branched (5-6); 9-T usually double (2-3); 10,11-T similar to those on meso-

thorax; 12-T much reduced. Abdomen. Hair 6-I usually 4-branched (3-4);

7-I single; 6-II usually 3-branched; 7-II usually with 2-3 branches; 6-III-V

double; 6-VI single; 1-VII usually 2- branched 2-3), long; 2-VI usually sin-

gle; comb of 9-12 scales in a single row, each scale with fine denticles or

fringes at base of apical spine; pentad hair 2-VIII distant from 1-VII; 1, 5-VIil

3-branched; 3-VIII with 5-6 branches; 2,4-VIII single; siphon about 2.5 times

as long as wide, acus absent; pecten teeth 11-12 in number, evenly spaced,

each tooth with 1-3 basal denticles; 1-S with 3-4 branches, inserted beyond

last tooth and usually before middle of siphon; saddle incomplete; marginal

spicules very small and inconspicuous; 1-X 2-branched; 2-X 2-branched; 3-X

single; ventral brush with 4 pairs of hairs on grid, each hair single except 1

or 2 proximal ones usually double (2-3); no precratal tufts; anal papillae 2.5

times as long as saddle, sausage-like.

TYPE DATA. Culex scutellaris Walker, type female in British Mu-

seum (Natural History), London; type locality: ARU (AROE) ISLANDS. Culex

zonatipes Walker, type female in British Museum (Natural History); type lo-

cality: Dorey, NEW GUINEA.

DISTRIBUTION. 163 specimens examined: 56c, 249, 400 terminalia,

4° terminalia, 31 individual rearings (81, 31 p).

INDONESIA. Moluccas: Aru (Aroe) Islands., 19; Aroe Islands,

Dobo (23-I-1932, Brug & de Rook), 10, 2c terminalia; Cevam- Sawaai (XII-

1931, Brug & de Rook), 2c; Nhust Sawai (VII-1931,-Brug & de Rook), 2c, 2c

terminalia; Ambon I., Waai (1966,A.M.R. Wagner), 10°, 22, 10% termi~

nalia, 29 terminalia, 11 individual rearings : 1, 11 p); West New Guinea -

Dorey, 19; Fak Fak (I-1932, Brug & de Rook), 1°, 1¢ terminalia; Cyclops

Mt. Sobron 930 ft. (V-VI-1936, L.E. Cheesman), 1c, 29, 1o terminalia;

Cyclops 1000 ft. (III-1945, Rozeboom, Knight & Laffoon), 20°, 2c terminalia;

Bougainville Bay (I-1945, Knight), 1o, 1o¢ terminalia; Hollandia (1962, ex.

Lab. Colony, Colless), 2c, 2c terminalia.

NEW GUINEA. Goodenough Is. (VI-1944, E.S. Ross), 1c, 1o ter-

minalia; Tanah Merah Road (II-1945, Schultz & Rozeboom), 39; Lake Sentani

(I1I-1945, Schultz), 1¢, 1o terminalia; New Guinea NE, Alexishafen (1944,

Johnson), 4c, 29, 4c terminalia; (XI-1964, W.A. Steffan), 200, 89, 50 ter-

minalia, 13 individual rearings (3 1, 13 p); Lae, Botanical Garden, 5-10 m.

(IV-1965, W.A. Steffan), 1c’, 29, 1o terminalia, 19 terminalia, 2 individual

rearings (2 p); Oomsis, 24 km. W. Lae (IV-1965, W.A. Steffan), 1¢, 1° ter-

minalia; Kaisinik, 1000 m. (IV-1965, M. Sedlacek), 19, 1 individual rearing

(11, 1 p); Madang, 0-5 m. (XI-1964, W.A. Steffan), 50, 29, 50 terminalia,

1? terminalia, 3 individual rearings (1 1, 3 p); New Guinea, SE, Cape Killer-

a ‘i m. (V-1965, W.A. Steffan), 1o, 1c terminalia, 1 individual rearing

p).

08 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

TAXONOMIC DISCUSSION. A. scutellaris isa Papuan species of the

scutellaris subgroup. The adultis very similar to alcasidi, malayensis and

riversi in having the mid femur without a median white line on anterior surface,

wing with minute basal spot of white scales on costa, hind tarsomere 5 all white.

The adult of scutellaris cannot be separated from alcasidi except by the male ter-

minalia but can be separated from malayensis and riversi in having hind tarso-

mere 3 with basal 1/2 white banded and hind tarsomere 4 with basal 3/4 white band-

ed; in malayensis and riversi hind tarsomere 3 has basal 2/5 white banded and

hind tarsomere 4 has basal 2/3 white banded.

The male terminalia of scutellaris are very similar to andrewsi in

having claspette with distal expanded part square in shape in lateral aspect

(dissected claspette), sternal and tergal sides more or less parallel and api-

costernal angle present. Itdiffers from andrewsi in having claspette with 5 or

6 modified setae, set on a prominence, close to apicosternal angle area and

being without several distinctly long and stout setae on apicotergal area; in

andrewsi the claspette has 4 or 5 modified setae in a row on apicosternal an-

gle and several distinctly long and stout setae on apicotergal area.

The larva of scutellaris resembles albopictus, alcasidi, malayensis

and viversi, but is closer to alcasidi and malayensis in having hair 1-VII usu-

ally with 2 long branches (2-3), when 3-branched then one much smaller than

the other two, siphon about 2.5 times as long as wide, pecten teeth 10-16 and

1-S inserted at middle or before middle of siphon. It is indistinguishable

from alcasidi and can only be separated from malayensis by the diagnostic

characters mentioned under the discussion of that species.

The pupa of scutellavis is very similar to albopictus, pseudalbopictus,

alcasidi, malayensis and riversi in having hair 9-VI about same magnitude

as 9-V, 9-VII single and simple, 9-VIII reaching beyond fringe of paddle. It

is closer to alcasidi, malayensis and riversi in having hair 6-C about 3/4

length of 7-C to about as long as 7-C. The pupa is indistinguishable from

alcasidi and can be separated from malayensis and riversi by the diagnostic

characters mentioned in the Key.

Although the immature stages of scutellavis are so similar to alcasidi

the male terminalia are markedly different by having the claspette with distal

expanded part square in shape in lateral aspect (dissected claspette), sternal

and tergal sides more or less parallel and apicosternal angle present; in

alcasidi claspette with distal expanded part subtriangular in shape in lateral

aspect (dissected claspette), sternal and tergal sides not parallel but tapering

and without apicosternal angle.

A. scutellaris is a Papuan, alcasidi a Philippine, hensilli a Micro-

nesian, malayensis a Southeast Asian and viversi a Ryukyu species. All of

them are extremely variable and difficult to separate in all stages except for

the male terminalia. The diagnostic characters are summarized in Table II.

A. scutellaris is apparently restricted to the East of Lee & Woodhill's

Line. It is presently known from Ceram, Ambon, Aru Islands and New

Guinea in the Papuan area.

BIOLOGY. The immature stages of scutellaris have been collected

from coconut shells and artificial containers.

ACKNOWLEDGEMENTS

I wish to express my Sincere appreciation to Dr. Botha de Meillon and

Dr. Alan Stone for the helpful assistance and valuable consultations through-

out this study and also for a critical review of the manuscript.

I am most grateful to Dr. P. F. Mattingly, Department of Entomology,

British Museum (Natural History), London, for several types and other ma-

terial in the British Museum; Dr. M. Sasa, Director, The Institute of Medical

Science, The University of Tokyo, for Yamada's syntype material of flavo-

pictus; Dr. W.A. Steffan, Department of Entomology, Bishop Museum,

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia og

TABLE Il. CHARACTERS FOR SEPARATING SOME SPECIES CLOSELY

RELATED TO AEDES (SSTEGOMYIA) SCUTELLARIS (WALKER)

Species

latcasidi| hensitti| scutellaris

Diagnostic

Characters

Hind tarsomere 4

with more than basal

0. 7 white

Hind tarsomere 5

with apical half

all dark

Claspette with

sternal and tergal

sides parallel

Apicosternal angle

present

Apicotergal area of

claspette with several

distinctly long setae

Modified setae on,

the sternal side

Modified setae closer

to sternal angle area

than to apicotergal

angle area

Modified setae set

on a prominence

Modified setae usually

more than 7 in number,

forming a prominent row

Modified setae occupy

less than half of the

sternal side

Main geographical Philip- |Micro- |Southeast em

distribution pines |nesia Asia ukyus} Papua

X = Has the character

- = Does not have the character

60 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

Honolulu, Hawaii, for the New Guinea material of scutellaris; Dr. A. Stone,

Agriculture Research Service, U.S. Department of Agriculture, for the type

material of hensilli and other species in U.S. National Museum; Mr. W. T.

Chellappah, Department of Parasitology, Faculty of Medicine, University of

Singapore, Singapore, for topotypic material of malayensis. The Thailand

material from Dr. D.J. Gould and his staff of the SEATO Medical Research

Laboratory, Bangkok; the Philippine material from Dr. G.L. Alcasid and

his staff, Department of Education, National Museum, Manila; the Ryukyus

material from Major R.W. Intermill, Chief, Entomology Division, U.S.

Army Medical Center, Ryukyu Islands and the Malayan and Indian material

from Dr. S. Ramalingam and his staff, Department of Parasitology, the

University of Malaya, Kuala Lumpur, are acknowledged with sincere appre-

ciation. I wish to thank also the following Institutions for the loan of materi-

al: Bernice P. Bishop Museum, United States National Museum; Field Muse-

um of Natural History; University of Utah; Cornell University; Johns Hopkins

School of Hygiene & Tropical Medicine; California Academy of Science;

Academy of Natural Science, Philadelphia; Medical Zoology Laboratory,

Institute for Infectious Disease, University of Tokyo; British Museum (Natural

History) and the Instituut voor Tropische Hygiene, Amsterdam.

I also wish to express my gratitude to Mr. Vichai Malikul of Southeast

Asia Mosquito Project for his help in making the drawings and to Miss Virginia

Ford, SEAMP, for her help in rearing and preparing the specimens, and to

Miss Helle Starcke, who rendered editorial assistance and typed the manuscript

for offset reproduction; and finally to my parents and friends for their kind

encouragement,

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Huang: Aedes Stegomyia) scutellaris group in Southeast Asia 67

APPENDIX I. PRESENT STATUS OF THE AEDES GTEGOMYIA)

SCUTELLARIS GROUP OF SPECIES IN SOUTHEAST ASIA

SPECIES ire ale BIOLOGY

a a a

ALBOPICTUS SUBGROUP

albopictus X* | X*| K*) K* Larval habitats known

Female bites man

downs pela ace | = Larval habitats known

Female bites man

novalbopictus Bae ee eee Larval habitats known

patriciae ee eee Larval habitats known

pseudalbopictus ee eae Larval habitats known

seatot pxefac ce ar] = Larval habitats known

Female bites man

subalbopictus** eee te aie Larval habitats known

SGUT ELLA Resueenane Bas 2

alcasidi X*| X* | X*| X* Larval habitats known

imal xvas] -|- [| unknown

nical Lxelxe] -|- | - | uninown

malayensis X* X* Larval habitats known

Female bites man

paullusi X*| X* Larval habitats known

Female bites buffalo

viverst x* perl xe] = Larval habitats known

Female bites man

X* = Stage or sex described and illustrated.

- = Stage or sex unknown.

X = Stage or sex described.

eae

Species may also be present in Southeast Asia.

68 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

APPENDIX Il.

SPECIES

DISTRIBUTION LIST OF THE SOUTHEAST ASIAN SPECIES

SOUTHEAST ASIA

5 .

mM ee

= d¢ 4%

ss I ft od

So 4 - 8 fi

= O'R ‘

cs ~ ee ae

c ee = oo 2 & E

= DAs 3 os = ® Ww ve

oA Mg z oe fe a @

GS 8 yp eee beg taod ad

aoe eg eh ee eS a4 oO oe

PeREOMMFPHOHRFAKTANRONDHNSM

ALBOPICTUS SUBGROUP

albopictus

downst

novalbopictus

patriciae

pseudalbopictus X

seatoi

mM [i OI

subalbopictus ** ?

SCUTELLARIS SUBGROUP

alcasidi

alorensis

_andrewsi

malayensis

paullust

Viverst

X X KX X

X = Areas from which specimens were available for

examination

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia 69

OF THE AEDES STEGOMYIA) SCUTELLARIS GROUP

Po OTHER LOCALITIES

Pp e- D a-

cage Paci cif.

Oriental Is. |Papuan | Malagasy Is.

Ler cence amen men seer rere ma ter hess erreur coerce ca errno nearer srg eee

Ds oO)

& 5

cs =. op 2 W

a So i a) ro} ce)

3 ae EIS & a

—_ ee ae, aS rr

DM wn Oo |G op) a ai @

D> oO ce 5 | GS Ne

Ct @ S Z rein xa 2 | &

a ok ae Bet ee alae aa 8 3/8

S 3 a ® ag @ Sag = ae ea Ge

Bp CM eee ee ae, S abe 55% oF

-PR sk aS Ss Has £\6 3 eS a Sle

SFnnarn OH dt OHD O|n & Aa vq ele

SS RR AR A LR A SE NN

= Species which may also occur in Southeast Asia.

= Species recorded from the area but have not been

seen by the author.

MATAGY SIHL AD GHUYAAOO VaeV VISV LSVAHHLNOS HHL “I dVWN

wa VISV LSVaHLBOS

SvwisivHo

3uo avons

SONV1SI

0 S| NVAVONY

BY fi

yotin ONVIHO> ‘ E i]

; Akt vusooos” >

uspy 30/ yin9d

wIWHSY® _/

“4

‘youynd au, Aq uaas suawioads uo pasog “spupjs| uDIIOMDH Bbulpnjoxa

‘(asnys) Snyoidog|o (DIAWOHaIS ) Sapay 40 uolsNquysig jod!IydD4sboay ----

Po)

ie. Viov- SESVIRLEOS

SVWLSINHO

“a

oes

w3nind want

z Xs S| NVAVONY

~aiao8 wv)

‘ y 2

Neyo geal

ONvrivHas, &

eva] ONVIHO > ts

‘ s 3 wasonoe?) ©

IER in

Sea ? et tatet

> bv Ls

Tita SEC,

Caer |

NGiy ae

MINHSY) _/

-N.

Neotype

Aedes ( Stegomyia )

ictus (Skuse) Cali Mealhale

albop

Fig. 2

ee Barf

Near ti

ae aNd

ee \\ ites

\ Is A

] 4 Si i

oS" -

paraprocts

/__— IX thtergum |

Neotype

Aedes (Stegomyia) albopictus (Skuse)

Aedes (Stegomyia) albopictus (Skuse )

7 ee

Boe

Bee aes ee

NESE RET

~ - - iz he :

- = = it $

~ he if we se

= ~ < ar ~ iz

SSeS SESE é

See . > y gs 8

) :

hah agit! hai

/ Bohart & Ingram

(Stegomyia ) down

Aedes

Aedes (Stegomyia) flavopictus Yamada

terga/ aspect

downs!

flavopictus

pictus Barraud

novalbo

gomyia )

(Ste

Aedes

Aedes (Stegomyia) novalbopictus Barraud

—.

(—)

=—

phe dM bbe hace HES

Sooo

i /

seo

gel) Soe

Sac

0-2

Aedes (Stegomyia) patriciae Mattingly

(Stegomy/a ) patriciae Mattingly

Aedes

Aedes (Stegomyia ) pseuda/bopictus (Bore!)

Aedes (Stegomyia) seatoi Huang ihuMhelibal,

Aedes (Stegomyia ) seatoi Huang

Tee

32 |

A A

Aedes (Stegomyia) seato/ Huang

Aedes (Stegomyia) subalbopictus Barraud

Aedes (Stegomy/a) subalbopictus Barrdud

alorensis paullusi andrewsi

ONML

subalbopictus

flavopictus patriciae

hind leg

Aedes ( Stegomyia) alcasidi n.sp.

AS}

gomyia )

( Ste

Aedes

]

2 vi

6 4 \ 10 431248 ‘

3 1

Aedes (Stegomyia ) alcasid/ n.sp. Malhul

Aedes (Stegomyia) alorensis Bonne-Wepster & Brug Aedes (Stegomyia)andrewsi Edwards

scutel/aris

hensilli

Aedes (Stegomyia ) hensilli Farner

Fig. 26

ip FZ,

VI Gp f°

Rae eae

VE 7

EES

PLY,

Wie,

447404

PLE Bags

57.

K7,

yes 4

iy: SIZ ZZ

alcasid/

y714

cerc/

sy postgenita/

plate

Spermathecae

Aedes (Stegomyia) malayensis Colless

War

rivers!

Tips

RPT AT

KIT

79 labs Sy,

malayensis

that fitabihul,

Aedes (Stegomyia) malayensis Colless

Aedes (Stegomyia ) malayensis Colless

Aedes (Stegomyia ) paullusi Stone & Farner

Aedes (Stegomyia) paullusj Stone & Farner

Aedes (Stegomyia) riversi Bohart & Ingram

scute/laris malayensis

rivers! hensil

Bf ifsi ALA

Nf) AAR he eRe

Len, Wilh) VV) Beate

ti & ELSA

Aedes (Stegomyia) riversi Bohart & Ingram

KOS

Topotypie male (Aroe /s/lands ) aha: eathals

Aedes (Stegomyia) scutellaris ( Walker )

Aedes (Stegomyia) scutellaris (Walker)

Aedes (Stegomyia) scutellaris (Walker )

108 Contrib. Amer. Ent. Inst., vol. 9, no. 1, 1972

INDEX

Names of valid taxa are in roman type; synonyms are in italic type.

Italic numerals refer to the principal text references. Roman numerals refer

to secondary text references; the suffix ''k'’ indicates mentioning in a key and

the suffix ''t'' in a table.

Roman numerals in parentheses without a suffix

refer to the figures and with the suffix ''m" to a map.

aegypti (Linnaeus)

albolineatus (Theobald)

albolineatus grou

albopictus (Skuse)

albopictus subgroup

alcasidi n. sp.

alorensis Bonne- Wepster

andrewsi Edwards

Armigeres

downsi Bohart & Ingram

flavopictus Yamada

hensilli Farner

lambertt Ventrillon

malayensis Colless

mediopunctatus subgroup

nigritia Ludlow

novalbopictus Barraud

patriciae Mattingly

paullusi Stone & Farner

polynesiensis Marks

pseudalbopictus (Borel)

quasinigritia Ludlow

riversi Bohart & Ingram

samarensis Ludlow

scutellaris (Walker)

scutellaris group

scutellaris subgroup

seatoi Huang

17, 34

274, 5, Gk, 8k, dik, 42k./13; 14.46;

17, 18t.*20, 21. 22; 23.31, 34-37, 40,

48, 49, 55, 58, 67t, 68t (1, 2, 3, 2m)

#, 6s 16, 20, 28, 26, 28, Sis 34, 36,

39, 67t, 68t

5, 7k, 9k, 11k, 12k, 16, 31, 37, 39, 40,.

44, 47, 48, 51, 52, 54, 55, 58, 59t,

67t, 68t (21, 22, 23, 26)

5, 7k, 8k, 41, 42, 51, 67t, 68t (eo, )

5, 7k, 9k, 42, 43, 58, 67t, 68t (20, 24

4, 5, 6k, 8k, 10k, 12k, 16, 17, 19, 20,

21, 23, 25, 28, 31, 34, 36, 55, 67t,

68t (4, 5, 8, 20)

3, 4, 7k, 8k, 10k, 11k, 20, 21, 22, 23,

25, 28, 31, 34 (6, 7, 8, 20)

3, 5, 7k, 9k, 40, 43, 44, 45, 48, 54,

55, 58, 59t (25, 31)

5, 7k, 9k, 11k, 12k, 16, 31, 40, 44,

45, 46, 47, 48, 49, 51, 52, 54, 55, 58,

59t, 67t, 68t (26, 27, 28, 31)

4, 6k, 7, 8k, 10k, 12k, 20, 23, 24, 25,

26, 36, 67t, 68t (8, 9, 10, 20)

4, Tk, 8k, 11k, 12k, 20, 23, 25, 26,

27, 28, 31, 36, 67t, 68t (11, 12, 20)

5, 7k, 9k, 10k, 12k, 41, 49, 50, 51,

52, 67t, 68t (20, 29, 30)

4, 5, 7k, 9k, 11k, 16, 20, 28, 30, 31,

36, 37, 40, 48, 55, 58, 67t, 68t (13,

14, 20)

5, 7k, 10k, 11k, 12k, 16, 21, 40, 44,

47, 48, 51, 52, 54, 55, 58, 59t, 67t,

68t (26, 31, 32, 33)

9 4..5, Te Selik, 12k, 16, 18,. 31,

40, 43, 44, 45, 47, 48, 51, 52, 54, 55,

57, 58, 59t (25, 31, 34, 35, 36)

1, 2, 4, 5, 6, 16, 67t, 69t

4, 5, 16, 39, 42, 43, 45, 47, 51, 54,

58, 67t, 68t

4, 5, 6k, 8k, 10k, 11k, 16, 20, 32, 33,

34, 67t, 68t (15, 16, 17)

Huang: Aedes (Stegomyia) scutellaris group in Southeast Asia 109

Stegomyia 1, 4, 5, 6

subalbopictus Barraud 4, 7k, 8k, 10k, 12k, 16, 20, 23, 25,

50} a1, 35... 36, a7, - Cit, Bat (18, 19,

variegatus Doleschall 5)5)

Zonatipes Walker 5)9)

Contributions

of the

American Entomological Institute

Volume 9, Number 2, 1972

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA. - XV

Genus Aedes Meigen, Subgenus Ayurakitia Thurman

John F. Reinert

CONTENTS

ABSTRACT ee ee ee ee |

INTRODUCTION: |. mua iy © oc. Were tae Sp le. eee, op ee tie ene

GENUS AEDES MEIGEN, SUBGENUS "AY URAKITIA THURMAN ...

KEYS TO THE SPECIES OF AEDES (AYURAKITIA) ...... 40.

ADULTS . APO NR Pre BE SR er ee a

MALE GENITALIA. en Ay: SEAS Seg Da) hee ees Osea ae emer

PANE iia. cn ea ce oR ice ey Oe SE igs ik ie el Jeng wa bai DQ nie) mere

AEDES (AYURAKITIA) PEYTONI NEW SPECIES

POUR Eo PAG A A ee os dig ge bee eee een 6

AEDES (AYURAKITIA) GRIFFITHI D. THURMAN .......00-

ACKNOWLEDGEMENTS ..... eee Fe | callie Sheet =

Le OE CRB ae ta eet eee ee ee eer ee .

EUS OP PIU screen hese eee te ndite @ Vere ace eee ere cece eens

Fes as were sae he ahaa eee wena Dat eeneiew etre he dws oe Flo ahe we

APPENDICES

TABLE 1. Record of the Branching of the Setae on the

Pupae of Aedes (Ayurakitia) peytoni .....

TABLE 2. Record of the Branching of the Setae on the

Pupae of Aedes (Ayurvakitia) griffithi .....

TABLE 3. Record of the Branching of the Setae on the

Larvae of Aedes (Ayurakitia) peytoni.....

TABLE 4. Record of the Branching of the Setae on the

Larvae of Aedes (Ayurakitia) griffithi ....

ATIARARHONWRH

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA, - XV.

GENUS AEDES MEIGEN, SUBGENUS AYURAKITIA THURMAN?

John F. Reinert“

ABSTRACT

The subgenus Ayurakitia Thurman of Aedes is defined, discussed and

compared to related subgenera. Descriptions, illustrations and keys are given

for the known stages of the 2 included species, peytoni, new species and

griffitht Thurman.

INTRODUCTION

The taxonomic history of Ayuvakitia is brief. Avyurakitia was origin-

ally erected as a monotypic genus by D.C. Thurman, Jr. in 1954. E.H.B.

Thurman (1959: 74) retained the single species, griffithi Thurman, in the genus

but Mattingly (1971: 31) in note number 28 of his key to the world genera of

mosquitoes, reduced Ayuvakitia to subgeneric rank in the genus Aedes Meigen.

In the present paper I have retained Ayurakitia as a subgenus of Aedes since

it fits well within the genus in all aspects except the absence of postspiracular

bristles in the adults. This feature is also shared by Kompza Aitken, a North

American monotypic subgenus of Aedes. The subgenus as defined possesses

characters in all known stages that distinguishes it from the other subgenera

of Aedes. A new Species, peytoni, is placed in Ayurakitia and is described

herein. :

During the course of this revision, I examined all specimens and

types of Ayurakitia in the United States National Museum (Natural History)

and the British Museum (Natural History).

Abbreviations used in references to literature conform to the World

List of Scientific Periodicals, 4th edition, Butterworths, Washington, 1963.

In the synonymy sections, an asterisk following the abbreviations used (2 =

female, o* = male) indicates that at least some portion of that sex is figured.

The abbreviations used in distribution sections are the same as in the synon-

ymy but with the following additions: P = pupa, L = larva, p = pupal skin and

1] = larval skin. Pupal descriptions, tables and key utilize the following ab-

breviations: C = cephalothorax; P = paddle; and I- VIII = abdominal segments

1 through 8. In larval descriptions the range of hair branching is given and

the following abbreviations signify: A = antenna; C = head; I-VI, X = abdo-

minal segments 1 through 8 and 10; M = mesothorax; P = prothorax; S = siphon;

and T = metathorax. When possible 10 specimens were used in determining

the range, mode and mean hair branching in pupal and larval descriptions and

tables. In the pupal descriptions, the number of branches on abdominal hair

1 This work was supported in part by Research Contract No. DA-49-193-MD-

2672 from the U.S. Army Medical Research and Development Command,

Office of the Surgeon General and carried out at the Southeast Asia Mosquito

Project, Smithsonian Institution, Washington, D.C. 20560.

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

Walter Reed Army Institute of Research, Walter Reed Army Medical Center,

Washington, D.C. 20012.

2 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

1-I is measured on the basal 0.33 of the hair. Measurement scales on the

illustrations are in millimeters, Distribution records are indicated as follows:

countries are in capital letters; provinces are in italics; and place names have

the first letter capitalized. The number of specimens examined from each

province follows the last place name of the province in the distribution sec-

tions. The spelling of provinces (changwats) and locality names in Thailand

is taken from the Official Standard Names Gazetteer No. 97 prepared by the

Office of Geography, Department of the Interior, Washington, D.C., April

1966. Locality names which do not appear in the gazetteer are spelled accord-

ing to the data sheets and labels on the specimens.

The nomenclature and chaetotaxy used for females, males and male

genitalia follow Knight (1970) and Knight and Laffoon (1970a, 1970b, 1971) and

those for the pupae and larvae follow Belkin (1962). The terminology of the

female genitalia is taken from Coher (1948).

GENUS AEDES MEIGEN

SUBGENUS AYURAKITIA THURMAN

Ayurakitia Thurman 1954, J. Wash. Acad. Sci. 44: 198.

Orthotype:. Ayurakitia griffithi Thurman.

Aedes (Ayurakitia) Thurman, Mattingly 1971, Contr. Am. ent. Inst. 7(4): 31

(Reduced Ayurakitia to subgeneric rank in the genus Aedes).

The two species assigned to the subgenus share the following combi-

nation of characters.

FEMALE. dead. Antenna approximately 0.85 length of proboscis,

pedicel with a few small broad brown scales mesally, flagellomere 1 witha

few small brown scales; clypeus bare; maxillary palpus approximately 0.18

length of proboscis; proboscis approximately 1.05 length of femur I; eyes con-

tiguous in front; vertex covered with broad decumbent golden scales with a

band of overlapping broad silvery scales around the dorsal margin of the eyes

which is separated in the middle by a small triangular patch of broad dark

scales; occiput with narrow curved decumbent golden scales; numerous dark

erect forked scales on occiput and vertex extending anteriorly to the silvery

band of scales; ocular setae well developed and located along posterior margin

of band of silvery scales. Thorax. Scutum covered with narrow curved red-

dish-black scales with prescutellar space bare; scutellum with a small patch

of narrow curved dark scales on each lobe in addition to a few broad ones on

the median lobe; following bristles present and well developed: median ante-

rior promontory, acrostichal, dorsocentral (anterior and posterior), scutal

fossal (3-4 anterior, 1-2 lateral and 1 posterior), 17-29 supra-alar, 1 post-

alar callar and scutellar (3-4 long and 1-2 short ones on each lateral lobe and

4 long and 1-3 short ones on median lobe); antepronota normal sized and widely

separated, 7-10 bristles on each side; postpronotum with 2 posterior bristles,

upper one short and lower one long; propleuron with a patch of overlapping

broad silvery scales, 4-7 bristles; mesepisternum with an upper and posterior

patch of broad overlapping silvery scales, 1 moderately long upper and 1 long

lower bristle and 3-6 short lower bristles ventral to lower scale patch; pre-

alar knob with 3-4 short bristles; mesepimeron with a patch of overlapping

broad silvery scales, a small patch of 5-8 short bristles on upper area, 2

bristles on lower anterior area, upper bristle short and lower one long; other

pleural areas bare. Legs. Femora I-III brown scaled with an apical silvery

scaled spot and a golden scaled longitudinal stripe on anterior surfaces, the

stripe is dorsal on I and ventral on II and III, II and III also with a dorsal sil-

very spot which extends onto anterior and posterior surfaces and is 0.30 from

apex, posterior surface of I and II golden scaled with a longitudinal brown

stripe which is ventral and on apical 0.75 of I and dorsal and on apical 0. 70 of

Reinert: Aedes (Ayurakitia) in Southeast Asia 3

II, posterior of III brown scaled with a ventral patch of golden scales on basal

0.50; tibiae I-III dark brown scaled with a posterior median longitudinal golden

scaled stripe which extends from base to apex on I and II, and from basal 0. 30

to apical 0. 25 on III; tarsi I-III dark brown scaled with basal white scaled bands

on tarsomeres 1-2 of I, on tarsomeres 1-3 of II and on tarsomeres 1-4 of III;

posttarsi I-III each with 2 ungues, I and II with ungues equal, each with a tooth;

III with ungues equal and simple. Wing. Upper calypter with 13-23 hairs; 2

remigial bristles. Abdomen. Terga dark brown scaled with silvery scaled

lateral spots, II-VI also each with a dorsobasal golden scaled band; sterna

each golden scaled with a narrow apical brown scaled band. Genitalia. Ter-

gum VII with numerous broad scales, 0.85 - 0.95 retracted into segment VII

and only apex visible dorsally; apex broad and only slightly rounded apically;

sternum VII with numerous broad scales and a small median apical indenta-

tion; tergum IX membranous, moderately long, covered with minute setae and

divided into 2 partially pigmented apical lobes each bearing 2-4 short bristles;

cerci retracted and only apical 0. 20 visible dorsally, each short, broad, flat

and broadly rounded apically, tergal surface completely covered with minute

spicules, short bristles and broad scales, 4-6 long stout bristles along margin

at apex, sternal surface covered with minute setae and 4-7 short bristles along

apical and distal 0.40 of outer area; postgenital plate with a deep median notch

forming 2 lobes each bearing 3-6 short bristles, entire surface covered with

minute setae; posterior cowl membranous and covered with tiny setae; anterior

and posterior sigma each narrow, lightly pigmented and covered with minute

setae; atrial plate moderately developed and lightly pigmented; insula tongue-

like, membranous, covered with minute setae and with 6 long thin bristles on

anterior 0.25; 3 pigmented, spherical spermathecae, 1 large and 2 slightly

smaller ones.

MALE. Similar to female in general habitus. Head. Antenna with

hair tufts directed mainly dorsally and ventrally, 0.75 - 0.84 length of probos-

cis; maxillary palpus 0.87 - 0.95 length of proboscis, segment 1 tiny, seg-

ments 2 and 3 long, segment 4 approximately 0.33 length of segment 3 and with

4 moderately long bristles dorsoapically, segment 5 short and downturned with

3-4 moderately long and 5-7 short bristles at apex. Thorax. Scutum with 15-

22 supra-alar bristles; antepronotum with 7-10 bristles; postpronotum with 2-

3 bristles; propleuron with 4-8 bristles; mesepisternum with 1 moderately long

upper bristle and 1 long and 3-8 short lower bristles; prealar knob with 3-5

bristles; mesepimeron with 5-6 bristles on upper area and 1 long lower bristle.

Legs. Posttarsi I-III each with 2 ungues, I with ungues unequal, larger one

with a tooth, II with ungues unequal and simple, III with ungues equal and sim-

ple. Wing. Upper calypter with 14-20 hairs. Abdomen. Terga brown scaled

with patterns of golden scales. Genitalia. Tergum IX with apical margin bi-

lobed with 1-6 bristles on each lobe, basal margin with a deep median indenta-

tion, entire surface covered with minute spicules; gonocoxite long and narrow,

dorsal and ventral surfaces with scattered long and moderately long bristles,

ventral surface with 2 irregular rows of 14-20 stout moderately long bristles

along sternomesal margin near middle entire gonocoxite covered with minute

Spicules; gonostylus with distal U. 73 expanded into a large bluntly pointed lat-

eral lobe and a slightly longer, narrow mesal arm which bears an apical flat-

tened gonostylar claw, lateral lobe with short and long hair-like spicules and

short setae scattered over sternal and tergal surfaces; basal mesal lobes con-

nected basally, basal 0.50 attached to mesal membrane of gonocoxite and dis-

tal 0.50 extended as a lobe with long subapical bristles and several shorter

ones scattered over remainder of area, entire surface covered with small hair-

like spicules; proctiger long, paraproct pigmented and bluntly pointed at apex,

cercal setae absent; phallosome with aedeagus divided into 2 lateral plates

which are connected basally, each plate with several long longitudinal lateral

teeth with tergally curved apices and covered with a very lightly pigmented

dorsal flap which bears scattered hair-like spicules on the sternal surface,

4 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

paramere long, approximately 0.80 length of lateral plate, parameral apodeme

broad basally and tapering into a long narrow distal arm; sternum IX large,

entire surface covered with minute spicules, several bristles near center of

posterior margin.

PUPA. Cephalothorax. Hair 6-C single or double and small; 7-C

double and very long. Respiratory trumpet. Moderately pigmented with nu-

merous hair-like spicules on inner surface; index 2.55 - 4.00. Abdomen.

Hairs 1-I with 17-34 branches on basal 0.33, 1-II with 5-14 branches, 1-III

double to 7 branched; 5-I single to 4 branched, 5-II,III double to 7 branched,

0-IV-VI single to double, 5-VII single to 4 branched; 6-VIL with 4-11 branches;

9-I-VI single, 9-VII well developed, double to 7 branched, 9-VII well devel-

oped, 4-20 branched. Paddle. Elliptical in shape; with minute spicules on

distal 0.50 and minute serrations on basal 0.50 of outer margin; midribreaches

and occasionally somewhat projects beyond apical margin of paddle; hair 1-P

long, stout, with apical portion smoothly recurved, single; index 1.36 - 1.92.

LARVA. Head. Mouth brushes normal, non-pectinate; hairs 4-7-C

long and barbed; 4-C with 5-15 branches; 5-C with 5-16 branches; 6-C with 6-

12 branches; 7-C with 9-17 branches; 8,9,11-14-C stellate; 8-C single to 9

branched; 9-C with 5-17 branches; 10-C stellate or barbed, double to 4 branch-

ed; 11-C with 15-22 branches; 12-C double or triple; 13-C with 4-7 branches;

14-C with 4-14 branches; 15-C double or triple; basal maxillary hair stellate,

triple to 7 branched; mental plate with 21-29 teeth. Antenna. Long, slightly

incurved, lightly pigmented, short stout spicules scattered over entire shaft,

most numerous on basal 0.50; hair 1-A moderately long, barbed, 5-8 branched,

inserted 0.39 - 0.47 from base; 2-A very long; 3-A short, 0.18 - 0.33 length

of 2-A; 4-A long, 0.49 - 0.72 length of 2-A; 5-A short, flat; 6-A short, coni-

cal in shape. Thorax. Hairs 0,1,8,9-P stellate, 3-P barbed or stellate, 4-7-

P barbed; 1,13,14-M stellate, 3,5-10,12-M barbed; 1,3,8,13-T stellate, 7,9,

10,12-T barbed. Abdomen. Hairs 1-I-VUI, 2,4,13-I-VU, 3,6,8-VU, 7-I-

VI, 10-VI, VII and 11-I stellate; 1-X, 2,4-VIII, 3-Il, VOI, 5-II-VII and 6-I-

VI, VIII barbed, 1-X long, stout, single to 5 branched and inserted at middle of

posterior margin of the saddle; 2-X moderately long, 6-13 branched; 3-X long,

single; comb with 19-36 scales arranged in 2 rows, posterior row shorter with

less than 1/3 of the scales, each scale with a long, stout, pointed, median spine

with short, stout denticles laterally on basal 0.75; saddle heavily pigmented,

incompletely rings segment X, with numerous short, stout, heavily pigmented

spicules scattered over entire surface, spicules large along entire posterior

margin, acus absent; ventral brush with 7-8 hairs on grid and no precratal

ones, anterior 2 hairs very short and single or occasionally double, following

2 hairs moderately long and single or double, remaining hairs long and double

to 4 branched; 4 anal papillae, moderately long and narrow with bluntly rounded

apices. Siphon. Heavily pigmented, very dark beyond middle with basal 0.45

and apical 0.10 paler, basal 0.50 - 0.60 covered with small short ridges of

spicules; index 3.30 - 5.00; pecten composed of 5-12 evenly spaced teeth with

lateral denticles and on basal 0.43 of siphon; hair 1-S long, barbed and mul-

tiple branched, base inserted distal to last pecten tooth and 0.38 - 0.50 from

base of siphon; hair 2-S short, stout, heavily pigmented with apex recurved.

EGG. Not known.

DISTRIBUTION. Only 2 species of Ayurakitia are known and they have

been collected from valleys in mountainous areas of western Thailand. The

range of the subgenus may well extend into the mountain ranges of eastern

Burma and northern Malaysia since similar habitats and climates occur in

these areas.

The known distributions of peytoni and griffithi are plotted on the map

in Figure l.

TAXONOMIC DISCUSSION. The subgenus Ayuvakitia possesses char-

acters in the female, male, pupa and larva that allows it to be separated from

the other subgenera of Aedes. The most striking characteristic is the absence

Reinert: Aedes (Ayurakitia) in Southeast Asia 5)

of postspiracular bristles in the adults which is unusual for the genus Aedes

and is found elsewhere only in the North American monotypic subgenus Kompia

Aitken. However, these 2 subgenera differ in many features in all stages.

Kompia, for example, has the adults with a few posterior acrostichal bristles;

prealar knob with numerous bristles; mesepimeron without lower bristles;

broad silvery scale patches on the following areas -- antepronotum, postpro-

notum, postspiracular, subspiracular and prosternum; male maxillary palpus

much like that of Aedimorphus Theobald, Ochlerotatus Lynch Arribalzaga,

Aztecaedes Zavortink, Protomacleaya Theobald, Abvaedes Zavortink and some

Finlaya Theobald; male genitalia with the aedeagus composed of a simple tube,

basal mesal lobe developed into a claspette with a large flattened, curved

bristle at apex, proctiger with cercal setae, and gonostylus long, narrow with

an apical gonostylar claw; pupae with abdominal hairs 9-II-VI well developed

into stout, heavily pigmented bristles; larvae with the ventral brush of abdom-

inal segment X with 13-14 hairs, anterior 1-2 hairs short, single to triple and

anterior to grid, posterior 2 hairs short with 11-14 branches, remainder of

hairs long with 1-4 branches; larval antenna short and with only a few minute

spicules; larval head hair 4-C multiple, short mesad and even with 6-C, 5-C

directly posterior to 6-C, 5-C and 6-C long and single, 7-C with 4-8 branches.

Kompia shows the closest relationship to Abvaedes, especially in the larval

and pupal stages, and does not appear to be related to Ayuvakitia.

Other important features of the adults of Ayuvakitia are: head with

vertex covered with decumbent broad scales and a band of overlapping silvery

scales around the dorsal margins of the eyes; numerous erect forked scales

on occiput and vertex; scutum with acrostichal bristles absent, dorsocentral

(anterior and posterior) bristles well developed and mesepimeron with 1 lower

bristle in the male and 2 lower ones in the female; and posttarsi of females

each with 2 ungues, I and II with ungues equal in size and each with a tooth

and III with ungues equal in size and simple. These characteristics may be

shared in part, but not in combination, by other subgenera of Aedes, The

structure of the male palpus of Ayuvakitia has a striking resemblance to that

of members in the subgenus Diceromyia Theobald.

Female genitalia of Ayurakitia are similar to those of Aedes nummatus

Edwards, Udaya Thurman and Aedes subgenera Diceromyia and Stegomyia

Theobald. They can be separated from these, however, by the deep median

indentation of the postgenital plate and the presence of long thin bristles on the

insula.

Male genitalia of Ayurakitia are most similar to those of Aedes num-

matus which is being transferred from Aedimorphus to Diceromyia. (Reinert

in press). The aedeagus is similarly developed in both nummatus and Ayura-

kitia especially the dorsal flap which possesses hair-like spicules on the ster-

nal surface. A similar aedeagus with a dorsal flap is found in Udaya, Aedes

subgenera Diceromyia, Aedimorphus and Stegomyia meronephada (Dyar and

Shannon) and vittatus (Bigot), but these all lack the spicules sternally on the

dorsal flap. Development of the parameres of the phallosome in Ayurakitia

resembles that of the subgenera Diceromyia, Aedimorphus and Neomelanico-

nion Newstead and the parameral apodemes are similarly developed in num-

matus and Aedimorphus. The proctiger of Ayurakitia resembles that of num-

matus, Aedes (Neomelaniconion), Aedes (Aedimorphus) vexans Group, and

some Aedes (Stegomyia) in being long, narrow and bluntly pointed apically but

can be separated from the first 2 by the lack of apical teeth. The apically ex-

panded gonostylus is superficially like Aedimorphus, some Diceromyia and

vittatus and the gonocoxite is long, narrow, without lobes and with modified

bristles on the sternomesal margin resembling conditions found in some

Finlaya and Diceromyia. Genitalia of Ayurakitia can be separated from those

of all the other subgenera of Aedes by the shape of tergum IX, development of

the basal mesal lobes, and a combination of the other features mentioned above

especially the development of the phallosome.

6 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

The most distinctive features of the pupae of Ayuvakitia taken singularly

or in combination separates this subgenus from the other subgenera of Aedes,

These features are: cephalothoracic hair 7-C bifid and very long, approxi-

mately equal in length to the width of abdominal segment VII; hair 6-C small;

abdominal hairs 5-IV-VI very long, each equal to or exceeds the combined

length of the following 2 abdominal segments; abdominal hairs 9-VII, VII well

developed and barbed, 9-VII double to 7 branched, 9-VIII with 4-20 branches;

and paddle elliptical in shape, midrib reaching and somewhat projecting be-

yond apical margin and hair 1-P single with apical portion smoothly recurved

into a hook.

Larvae of Ayurakitia are distinguished from other subgenera of Aedes

by the following combination of characters: head hairs 4,5,6,7-C multiple

branched, well developed and approximately equal in length, 4-C with branches

slightly shorter than 5,6, 7-C; 5-C posterior in position to 6-C and posterior

and mesal to 7-C; 4-C mesad and equidistant from 5-C and 6-C; 7-C posterior

and slightly mesad to base of antenna; antenna long with numerous stout spic-

ules scattered over entire surface; antennal hair 1-A multiple branched, barb-

ed and branches extend beyond apex of antennal shaft; head, thorax and abdo-

men with numerous stellate hairs, this feature, however, is shared by many

Species of mosquitoes that breed in water in plant-containers; and development

of the ventral brush on abdominal segment X. The head hair arrangement and

development are similar to some species of Diceromyia in Africa but these

species can be distinguished from Ayuvakitia on other characters mentioned in

the subgeneric description.

MEDICAL IMPORTANCE. Nothing is known about the medical signif-

icance of the species in the subgenus.

BIOLOGY. The larval habitat is water in small plant-containers.

Immatures have been collected from clear or colored fresh water in small and

large Pandanus axils, banana axils and bamboo internodes; usually in heavily

or partially shaded areas located in primary and secondary rain forests, sec-

ondary deciduous forests, rubber plantations, bamboo groves and once each

from a secondary open swamp and a rice field; sites were in valleys located in

mountainous areas. Larvae have been collected in association with those of

several species of Aedes, Anopheles, Culex, Malaya, Orthopodomyia, Topo-

myia and Tripteroides that inhabit the water found in Pandanus and banana ax-

ils and bamboo internodes.

Adults have been collected resting on tree trunks and vegetation in

jungle valleys located in mountainous areas.

KEYS TO THE SPECIES OF AEDES (AYURAKITIA)

ADULTS

Occurring in mountains of Thailand south of '

the Isthmus of Kra @ e e@ e e e @ @ e e e se @ @ e e @ e @ @ peytont

Occurring in mountains of Thailand north of

RG TRUS NE IEP a5 ate uate tener x Meth fe 0 Fmt Ae 8 Be griffithi

MALE GENITALIA

Tergum IX bilobed with 4-6 bristles on each

lobe; basal mesal lobe with 10-12 short

bristles on distal 0.50 in addition to 2 long

BUN ADIGAL OPE eo. ee ah ae) aida Gel eles rte be ee eee ee See peytont

Reinert: Aedes (Ayurakitia) in Southeast Asia 7

Tergum IX bilobed with 1-3 bristles on each

lobe; basal mesal lobe with 6-9 short

bristles on distal 0.50 in addition to 2 long

Subspical.one sun. 4% mei tea te eniges tance ve recaels iat etal ee oe griffithi

PUPAE

Abdominal hair 9- VII with 4-8 branches;

abdominal hairs 5-IV, VI single © «2 «2 6 6 000s 6 6's peytoni

Abdominal hair 9-VII with 13-20 branches;

abdominal hairs 5-IV, VI doubles -.:..4 6 coeeelere Res griffithi

FOURTH STAGE LARVAE

Head hair 5-C with 5-9 branches; prothoracic

hair 9-P with 9-17 branches; abdominal

hain. ie eingle. on doutle Galipas mise vite Bel Glield): «adie peytont

Head hair 5-C with 11-16 branches; prothoracic

hair 9-P double to 5 branched; abdominal

hair 1-X with 4-5 branches Cc) e @ e o @ @ @ e e e e@ e e e 6 griffithi

AEDES (AYURAKITIA) PEYTONI NEW SPECIES

(Figs. 2, 3, 4, 6, 8, 10)

FEMALE (Fig. 2). Head. Antenna dark brown, approximately 0. 85

length of proboscis, pedicel light brown with a few small broad brown scales

and a few short brown hairs mesally, flagellomere 1 with basal 0. 85 pale

brown with a few small brown scales; clypeus brown, bare; maxillary palpus

brown scaled, approximately 0.18 length of proboscis; proboscis golden scaled

with basal 0.07 and apical 0.30 dark brown scaled, a median dorsal longitudi-

nal dark brown scaled stripe from base to apex and a few brown scales scat-

tered over ventral surface, approximately 1.05 length of femur I; vertex cover-

ed with broad decumbent golden scales with a band of overlapping broad silvery

scales around the margin of the eyes, band is separated in the middle by a

small triangular patch of broad brown scales; lateral surface covered with

broad golden scales; occiput with narrow curved golden scales; numerous dark

brown erect forked scales on occiput and vertex extending anteriorly to the

Silvery band of scales. Thorax. Scutal integument orange- yellow colored;

scutum covered with narrow curved reddish-black scales with prescutellar

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

scales on each lobe in addition to a few broad reddish-black scales on me-

dian lobe; median anterior promontory, acrostichal, dorsocentral (ante-

rior and posterior), scutal fossal (3-4 anterior, 2 lateral and 1 posterior), 17-

21 supra-alar, 1 postalar callar and scutellar (3 long and 1-2 short lateral and

4 long and 1 short median) bristles reddish- black and well developed; pleural

integument orange- yellow; antepronotum with 9-10 dark bristles; postpronotum

with 2 posterior dark bristles, upper one short and lower one long; propleuron

with a patch of overlapping broad silvery scales, 4-5 dark and golden bristles;

paratergite, postspiracular and subspiracular areas bare; mesepisternum with

an upper and a posterior patch of broad overlapping silvery scales, 1 moder-

ately long upper and 1 long dark and 5-6 short golden lower bristles; prealar

knob with 3 short dark bristles; mesepimeron with a patch of overlapping broad

8 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

Silvery scales on upper area, a small patch of 7-8 short dark and golden

bristles on upper area, 2 bristles on lower anterior area, upper one short

and golden and lower one long and dark; other pleural areas bare. Legs.

Coxae I-III each with several brown and golden bristles, I with broad brown

scales on anterior surface and a patch of broad golden scales dorsally, II, II

each with broad golden scales anteriorly and a small lower posterior patch of

brown scales; trochantersI-III each with a few short dark bristles, a patch of

broad golden scales and a small spot of broad brown scales; femora I-III each

brown scaled with an anterodorsal silvery scaled spot at apex, a basal golden

scaled band and a longitudinal golden stripe on anterior surface, stripe be-

comes narrower distally and is located dorsally on I and ventrally on II and III,

II and III also with a dorsal silvery scaled spot which extends onto anterior and

posterior surfaces and is located 0.30 from apex, posterior surface of I andII

golden scaled with a longitudinal brown stripe which is ventral and on apical

0.75 of I and dorsal and on apical 0.70 of II, I also with a small posterodorsal

silvery scaled spot at apex, III with posterior surface brown scaled with aven-

tral patch of golden scales extending from base to middle, patch broad at base

and tapering to a point distally; tibiae I, Il each dark brown scaled with a

posteromedian longitudinal golden stripe from base to apex, II also with a

ventrobasal golden spot, III dark brown scaled with a posteromedian gold-

en stripe extending from basal 0.30 to distal 0.25, also a basal golden

Spot ventrally and extending dorsally onto anterior and posterior surfaces;

tarsi I-III dark brown scaled, I with basal white bands on tarsomeres 1,

2, a posteroventral white longitudinal stripe on basal 0.50 of tarsomere 1,

II with broad basal white bands on tarsomeres 1-3, a posteroventral white lon-

gitudinal stripe on tarsomere 1 and basal 0.50 of tarsomere 2, III with broad

basal white bands on tarsomeres 1-4; posttarsi I-III each with 2 ungues, I, II

each with ungues equal and each with a tooth, III with ungues equal and simple.

Wing. Dorsal and ventral veins covered with brown scales; costa with golden

scales on basal 0. 25 of its posterior margin dorsally and ventrally; alula with

moderately broad brown scales along fringe; upper calypter with 13-21 hairs;

2remigial bristles. Halter. Pedicel pale, capitellum brown scaled. Abdo-

men. Terga dark brown scaled, I with a rectangular patch of silvery scales

on laterotergite, II-VI each with a median silvery scaled spot on lateral sur-

face, VII with a dorsobasal silvery scaled spot, II with a dorsobasal narrow

golden scaled band and a few brown scales intermixed with golden ones, II,

IV each with a dorsobasal broad golden band and a pair of indistinct dorsal ad-

median golden scale patches, V, VI each with a broad dorsobasal golden band

which is narrow medially and broad laterally, dorsolateral margins of this

band extend to posterior margins of terga; sterna each golden scaled with a

narrow apical brown scaled band, brown band broader mesally and does not

reach lateral margins on VI, V0; terga and sterna with numerous golden bris-

tles, mostly along posterior margins. Genitalia (Fig. 4). Tergum VII 0.85

retracted into segment VI and visible dorsally, apex broad and only slightly

rounded apically; sternum VII with a small median apical indentation; tergum

IX with 2 partially pigmented lateral lobes each bearing 3-4 short bristles,

entire surface covered with minute setae; cerci retracted and only apical 0. 20

visible dorsally, each cercus short, broad, flat and broadly rounded apically,

tergal surface covered with scales, short bristles and minute spicules, 4-5

long stout bristles along lateral margin at apex, sternal surface with 4-7 short

bristles along apical and distal 0.40 of outer area and minute setae scattered

over entire area; postgenital plate broad, a deep median indentation forming

2 lobes, 3-5 short bristles on each lobe, entire surface covered with minute

setae; posterior cowl membranous and covered with tiny setae; anterior cowl,

anterior and posterior sigma each narrow andcovered with minute setae; atrial

plate moderately developed and lightly pigmented; insula tongue- like, membra-

nous, covered with minute setae and with 6 long thin bristles on anterior 0. 25;

3 pigmented, spherical spermathecae, 1 large and 2 slightly smaller ones.

Reinert: Aedes (Ayurakitia) in Southeast Asia 9

MALE (Fig. 3). Similar to female in general habitus. Head. Antenna

with hair tufts directed mainly dorsally and ventrally, approximately 0. 84

length of proboscis; maxillary palpus with segments 1, 2,5 brown scaled, seg-

ment 3 golden scaled with apical 0. 20 brown scaled and a few brown scales

near base, segment 4 brown scaled with a few golden scales laterobasally and

with 4 moderately long golden bristles dorsoapically, approximately 0.35 length

of segment 3, segment 5 short and downturned with golden bristles at apex, of

which 3-4 are moderately long and 5-7 are short, segments 3,4 long and ap-

proximately equal in length, overall length approximately 0.95 of proboscis;

proboscis with numerous brown scales intermixed with golden ones. Thorax.

Scutum with 1-2 lateral scutal fossal bristles and 19-22 supra-alar bristles;

antepronotum with 7-9 bristles; postpronotum with 2-3 bristles; mesepister-

num with 1 moderately long upper bristle and 1 long and 4-5 short lower ones;

prealar knob with 3-4 bristles; mesepimeron with 5-6 bristles on upper area

and only 1 lower bristle. Legs. Posttarsi I-III each with 2 ungues, I with un-

gues unequal, larger one with a tooth, II with ungues unequal and simple, III

with ungues equal and simple. Wing. Upper calypter with 14-18 hairs. Ad-

domen. Tergum I brown with laterotergite silvery scaled; terga II-VI each

with dorsal surface golden scaled with a posteromedian triangular brown scaled

patch which has the apex pointing anteriorly, lateral surface brown scaled with

a large median patch of silvery scales and golden scales basally; terga VII, VIII

each brown scaled with a small median basal patch of golden scales and a pair

of admedian silvery spots dorsally; sterna as in female except VIII which is

brown scaled with a basal pair of large silvery spots. Genitalia.(Fig. 6). Ter-

gum IX bilobed with 4-6 bristles on each lobe, anterior margin with a deep

median indentation, entire surface covered with minute spicules; gonocoxite

long and narrow, dorsal surface with scattered long and a few moderately long

bristles from near base to near apex, a number of short bristles along tergo-

mesal margin from near base to near apex, lateral and ventral surfaces with

numerous scales, ventral surface with 8-11 long and moderately long bristles

on distal 0.50 and 3-4 long bristles on basal 0.50, a few short bristles on basal

0.25 mainly on mesal area, 2 irregular rows of 14-20 stout moderately long

bristles on sternomesal margin near middle, entire surface covered with mi-

nute spicules; gonostylus with pedicel short and moderately broad, distal 0. 73

expanded into a large bluntly pointed lateral lobe with a short seta at apex and

a slightly longer, narrow mesal arm which is bulbous distally with an apical

flattened gonostylar claw with apex blunt and recurved, lateral lobe withdistal

0. 80 covered with short and long hair-like spicules, 11-19 short setae scattered

over sternal surface, 1-4 similar setae on tergal surface near apical margin and

3-5 short setae on tergal surface near base of mesal arm; basal mesal lobe with

basal 0.50 attached to mesal margin of gonocoxite and distal 0.50 producedas a

lobe with 2 long subapical bristles, 9-11 short bristles scattered over remainder

of distal 0.50 and 1 short bristle ona small tubercle near middle of mesal margin,

entire surface covered with small hair-like spicules; proctiger long, paraproct

pigmented and bluntly pointed apically, cercal setae absent; phallosome with aede-

agus with 2 lateral plates connected basally, each plate with 9-10 long, lateral

teeth with tergally curved apices and covered with avery lightly pigmented dorsal

flap which has scattered hair-like spicules on sternal surface, paramere long,

approximately 0.80 length of lateral plate, parameral apodeme broad basally

and tapering into a long narrow distal arm; sternum IX large, entire surface

covered with minute spicules, 2 bristles near center of posterior margin.

PUPA (Fig. 8). Chaetotaxy as figured and recorded in Table 1.

Respiratory trumpet. Moderately pigmented; numerous hair-like spicules on

inner surface; index 3.55 - 4.00, average 3.81. Abdomen. Hairs 5-IV, VI

very long and single; 5-V very long and usually single, double in one specimen;

9-VII well developed and barbed, double to 4 branched; 4-VIII single and occa-

sionally double; 9-VII well developed and barbed, 4-8 branched. Paddle. El-

liptical in shape; with minute serrations on basal 0.50 and minute spicules on

ip Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

distal 0.50 of outer margin; midrib reaches and somewhat projects beyond

apical margin of paddle; hair 1-P long, stout, with apical portion smoothly re-

curved, Single; index 1.58 - 1.92, average 1.73.

LARVA (Fig. 10). Chaetotaxy as figured and recorded in Table 3.

Head, Hair 4-C long and barbed with 5-7 branches; 5-C long and barbed with

0-9 branches; 6-C long and barbed with 6-9 branches; 14-C stellate with 7-14

branches; basal maxillary hair stellate with 5-7 branches; mental plate with

25-29 teeth, usually with 26 teeth. Antenna. Hair 1-A attached 0.39 - 0.42

from base. Thovax. Hair 0-P stellate with 18-25 branches; 1-P stellate with

16-21 branches; 5-P barbed and double; 9-P stellate with 9-17 branches; 1-M

stellate with 20-26 branches; 4-M barbed and single; 14-M stellate with 19-27

branches; 1-T stellate with 20-31 branches; 3-T stellate with 19-28 branches.

Abdomen, 1-1 stellate with 23-34 branches; 2-I stellate with 13-17 branches;

4-I stellate with 16-18 branches; 6-I barbed with 6-7 branches; 10-I single;

1-II stellate with 19-27 branches; 2-II stellate with 13-19 branches; 4-II stel-

late with 17-24 branches; 13-II stellate with 17-24 branches; 1-IIL stellate with

20-25 branches; 2-III stellate with 10-17 branches; 4-III stellate with 16-20

branches; 7-III stellate with 17-20 branches; 13-III stellate with 15-19 branches;

1-IV stellate with 24-28 branches; 2-IV stellate with 11-16 branches; 4-IV stel-

late with 18-22 branches; 7-IV stellate with 20-24 branches; 13-IV stellate with

14-20 branches; 1-V stellate with 22-30 branches; 2-V stellate with 12-16

branches; 4-V stellate with 18-22 branches; 7-V stellate with 16-23 branches;

13-V stellate with 14-19 branches; 1-VI stellate with 19-30 branches; 2-VI

stellate with 13-23 branches; 4-VI stellate with 20-27 branches; 7-VI stellate

with 15-19 branches; 10-VI stellate with 9-12 branches; 13-VI stellate with 14-

17 branches; 1-VII stellate with 21-27 branches; 2-VIL stellate with 14-18

branches; 3-VIL stellate with 20-23 branches; 6-VIL stellate with 10-15 branches;

8-VII stellate with 18-25 branches; 10-VII stellate with 7-14 branches; 13-VI

stellate with 10-13 branches; 1-X long, barbed usually double with one branch

long and second branch approximately half as long, occasionally long and sin-

gle and once long and triple; 2-X moderately long with 6-9 branches; comb with

28-36 scales arranged in 2 rows. Siphon. Index 4.38 - 5.00.

TYPE DATA. Type series includes holotype female, allotype and

paratypes (9 females, 10 males, 26 whole larvae). Aedes (Ayurakitia) peytoni,

holotype female with associated larval and pupal skins, THAILAND, Phangnga,

Sip Si Hon (approximately 5 miles north of Rhung Nga), 20 October 1966, E.L.

Peyton collector, collection number 01782-1, SEAMP accession number 84,

collected as a larva from colored water in an unshaded large Pandanus axil lo-

cated in a rice field in a valley between mountains at an altitude of 100 feet;

allotype male with associated pupal skin, THAILAND, Phangnga, Sip Si Hon,

20 October 1966, E.L. Peyton collector, collection number 01781-103, SEAMP

accession number 84, collected as a pupa from colored water in an unshaded

large Pandanus axil located in a secondary rain forest in mountainous terrain

at an altitude of 100 feet; paratype 1 whole larva, same data as holotype, col-

lection number 01782; paratypes 2 males with associated pupal skins, collec-

tion number 01781-104 and 01781-106 and 3 whole larvae, collection number

01781, same data as allotype; paratypes 2 females with associated larval and

pupal skins, collection numbers 01699-6, 01699-12, 1 female, collection num-

ber 01699-11, and 2 whole larvae, collection number 01699, THAILAND,

Phangnga, Pathum, 15 October 1966, Kol Mongkolpanya collector, collected

as larvae from colored water in an unshaded large Pandanus axil located ina

rubber plantation in a valley of mountainous terrain at an altitude of 150 feet;

3 males with associated pupal skins, collection numbers 01701-100, 01701-101,

01701-104 and 1 whole larva, collection number 01701 with same data as col-

lection number 01699 except E. L. Peyton collector; paratypes 3 females and 1

male with associated larval and pupal skins, collection numbers 01704-2,

01704-3, 01704-4, 01704-5, 1 male, collection number 01704-17, 12 whole

larvae, collection number 01704, 1 female with associated larval and pupal

Reinert: Aedes (Ayurakitia) in Southeast Asia 11

skins, collection number 01700-1 and 1 whole larva, collection number 01700,

same data as collection number 01699 except Somboon Maneechai collector;

paratype 1 male with associated pupal skin, same data as allotype except Kol

Mongkolpanya collector, 18 October 1966 and at an altitude of 220 feet; para-

types 1 male with associated larval and pupal skins, collection number 01820-1

and 2 whole larvae, collection number 01820, same data as allotype except lo-

cality Nam Tai, 22 October 1966, Kol Mongkolpanya collector, and at an alti-

tude of 450 feet; paratypes 1 male and 1 female with associated larval and pupal

skins, collection numbers PG 1-30, PG 1-32, THAILAND, Phangnga, Ampur

Tye Murng, 33 km from Tha Gua Pha, 5 October 1964, E.L. Peyton, Kol

Mongkolpanya and Sumeth Chunchunchem collectors, collected as larvae from

water in a Pandanus axil 6 feet above ground surface in a dense secondary

growth on a mountain side 400 yards from Andaman Sea at an elevation of 245

feet; paratypes 1 female with associated larval and pupal skins, collection num-

ber TG 61-30 and 2 whole larvae, collection number TG 61, THAILAND, Trang,

Ampur Muang, National Park at Trang, 9 October 1964, E.L. Peyton collec-

tor, collected as larvae from water in a Pandanus axil; paratypes 2 whole lar-

vae, collection number 02176, THAILAND, Ranong, Ban Phon Rang, 19 July

1967, Kol Mongkolpanya collector, collected from water in a small Pandanus

axil. All type material is deposited in the U.S. National Museum (Natural

History), Washington, D.C., except 1 male and 1 female paratypes which will

be deposited in the British Museum (Natural History), London, England and 1

male and 1 female paratypes which will be sent to Department of Entomology,

SEATO Medical Research Laboratory, Bangkok, Thailand.

DISTRIBUTION. Occurs in mountainous areas of Thailand south of the

Isthmus of Kra. Material examined: 11, 109, 32 L, 1 p, 21, 18 with asso-

ciated skins (6 p, 121).

THAILAND. Nakon Si Thammarat; Ban Sai Koe; Ban Thuan Lek;

Khao Luang; 5 L. Phangnga: Nam Tai; Pathum; Sip Si Hon; Tye Murng, Tha

Gua Pha; llc, 99, 19 p, 22 L, 121. Ranong: Ban Phon Rang; 2L. Tvang:

Muang, National Park at Trang; 19, 1 p, 3 L, 21.

TAXONOMIC DISCUSSION. The female habitus is described from the

holotype and the female genitalia is from a paratype. The abdominal markings

of the female paratypes are variable and some differ from the holotype. Abdo-

minal terga II, II, V in several paratypes, have only a basal golden scaled band

dorsally and no scattered golden scales or lateral extensions of the bands.

Paratypes also differ from the holotype as follows: scutum with 2-3 median an-

terior promontory bristles and 17-24 supra-alar bristles; scutellum with 3-4

long and 1-2 short bristles on lateral lobes and 4 long and 1-2 short bristles

on median lobe; antepronotum with 8-10 bristles; mesepisternum with 1 long

and 4-6 short bristles in lower patch; mesepimeron with 6-8 bristles on upper

area; and wing with 13-21 hairs on upper calypter.

The females of peytoni are very similar to those of griffithi and there

appear to be no clear cut differences with which to separate them except geo-

graphically in that gviffitht occupies a range in the mountains of Thailand north

of the Isthmus of Kra while peytoni is found in the mountains south of the Isth-

mus of Kra. Some characters, however, have different ranges, even though

the ranges partially overlap, but when used in combination serve to separate

Some specimens. These features are discussed under gviffithi.

The male habitus of peytoni and griffithi are also very similar. There

are, however, differences in the male genitalia as follows, in peytoni: tergum

IX is bilobed with 4-6 bristles on each lobe; basal mesal lobe with 2 long sub-

apical bristles and 10-12 short bristles over remainder of distal 0.50; and

aedeagus composed of 2 lateral plates each with 9-10 teeth; while in gviffithi

tergum IX is bilobed with 1-3 bristles on each lobe; basal mesal lobe with 2

long subapical bristles and 6-9 short bristles over remainder of distal 0.50;

and aedeagus composed of 2 lateral plates each with 7-8 teeth.

12 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

The pupa of peytoni has the following distinctive features: hair 9-VII

with 4-8 branches; hairs 5-IV, VI single; trumpet index 3.55 - 4.00; and pad-

dle index 1.58 - 1.92; while griffithi has: hair 9- VII with 13-20 branches;

hairs 5-IV, VI double; trumpet index 2.55 - 3.44; and paddle index 1.36 - 1.52.

The larva of peytoni superficially resembles that of griffithi but can

easily be separated from it by the above mentioned features. Some of the most

distinctive characteristics of peytoni are: hair 4-C with 5-7 branches; hair

o-C with 5-9 branches; hair 8-C single to triple; hair 9-P with 9-17 branches;

branching of abdominal hairs 1, 2, 4,13-I-VU; and hair 1-X single to double;

while griffithi larva has: hair 4-C with 9-15 branches; hair 5-C with 11-16

branches; hair 8-C with 5-9 branches; hair 9-P double to 4 branched, once 5

branched; and hair 1-X with 4-5 branches.

This species is named for Mr. E.L. Peyton in recognition of his valu-

able work on mosquitoes in Southeast Asia.

BIOLOGY. The immature habitat is water in small plant-containers.

Immatures were collected from clear or colored fresh water in small and

large Pandanus axils and once from bamboo internodes; in heavy, partial or

unshaded areas located in primary and secondary rain forests, rubber planta-

tions, bamboo groves, and once each from a rice field and a secondary open

Swamp; sites were in valleys located in mountainous areas at elevations of 33

to1,000 feet, most often from elevations of 100 to 150 feet. Larvae were col-

lected in association with the following species of mosquitoes: Aedes poicilus

(Theobald), Anopheles sintonoides Ho, Culex (Lophocervaomyia) species,

Malaya genurostris (Leicester), Ovrthopodomyia albipes Leicester, Topomyia

inclinata Thurman and Tripteroides aranoides (Theobald).

AEDES (AYURAKITIA) GRIFFITHI D. THURMAN

(Figs. 3, 5, 7, 9, 11)

Ayurakitia griffitht Thurman 1954, J. Wash. Acad. Sci. 44: 198 (o*, 9);

Thurman 1959, Univ. Md. Agri. exp. Sta. Bull. A-100, p. 75, 158

(o*, 2); Stone et al. 1959, Thomas Say Found. 6: 211.

Aedes (Ayurakitia) Thurman, Mattingly 1971, Contr. Am. ent. Inst. 7(4): 31

(griffithi not mentioned but included by implication).

The adults of griffithi are very similar to those of peytonz but some

characters show a different range of values, though partially overlapping, when

used in combination serve to identify some specimens. These ranges and oth-

er variable characters for the presently known adult specimens of griffithi

(170, 82) and peytoni (11°, 102) are listed below.

FEMALE griffithi peytoni

Number supra-alar bristles 18-29 17-24

Number antepronotal bristles 19: 8-10

Number propleural bristles 4-7 4-5

Number lower mesepisternal bristles : 4-6 Sat

Number upper mesepimeral bristles saa 6-8

Number upper calypter hairs 14-23 13-21

Reinert: Aedes (Ayurakitia) in Southeast Asia ig

MALE griffitht peytoni

Number supra-alar bristles 15-20 19-22

Number antepronotal bristles 9-10 1-9

Number propleural bristles 4-6 o-8

Number lower mesepisternal bristles 4-9 5-6

Number prealar knob bristles 4-5 3-4

Number upper calypter hairs 17-20 14-18

The following account of the female and male of griffithi contains

characters that shows a range of variation not seen in peytoni. The male gen-

italia, pupa and larva show characters that differ from peytoni.

FEMALE (Fig. 3). Head. Maxillary palpus brown scaled in some

specimens but others have a few golden scales at base of apical segment and

scattered over dorsal surface; proboscis in some specimens with only a few

brown scales dorsally and without a stripe while other specimens have a defi-

nite brown dorsomedian brown stripe. Abdomen. TergaTII-V each with a nar-

row dorsobasal golden scaled band, VI with a dorsobasal golden band, narrow

medially and extending posteriorly along lateral margins, VII brown scaled,

some specimens with a few golden scales dorsobasally, one specimen with

dorsal markings similar to tergum VI, VIII with a large dorsobasal silvery

scaled spot. Genitalia (Fig. 5). Tergum VII 0.95 retracted into segment VII;

tergum IX with 2-4 bristles on each lateral lobe; cerci with 5-6 long stout bris-

tles along lateral margin at apex, sternal surface with 4-6 short bristles along

apical and distal 0.40 of outer area; postgenital plate with 4 short bristles on

each lobe.

MALE (Fig. 3). Head. Antenna approximately 0.75 length of probos-

cis; maxillary palpus approximately 0.87 length of proboscis. Genitalia (Fig.

7). Tergum IX with 1-3 bristles on each lobe; gonocoxite with 15-19 stout

moderately long bristles on sternomesal margin; gonostylus with lateral lobe

with 12-18 scattered short setae on sternal surface, 1-3 similar setae on ter-

gal surface near base of mesal arm; basal mesal lobe with distal 0.50 pro-

duced into a lobe with 2 long subapical bristles, 5-8 short bristles scattered

over remainder of distal 0.50 and 1 short bristle on a small tubercle near

middle of mesal margin; phallosome with 7-8 teeth on each lateral plate of the

aedeagus; sternum IX with 2-4 bristles near center of posterior margin.

PUPA (Fig. 9). Chaetotaxy as figured and recorded in Table 2. _

Respiratory trumpet. Moderately pigmented; index 2.55 - 3.44, average 3.08.

Abdomen. Hairs 5-IV-VI very long and double; 9- VII well developed and barb-

ed, 4-9 branched; 4-VIII double to 4 branched; 9-VIII well developed and barb-

ed, 13-20 branched. Paddle. Elliptical in shape; with minute serrations on

basal 0.50 and minute spicules on distal 0.50 of outer margin; midrib reaches

and somewhat projects beyond apical margin of paddle; hair 1-P long, stout,

tei apical portion smoothly recurved, single; index 1.36 - 1.52, average

- 48.

LARVA (Fig. 11). Chaetotaxy as figured and recorded in Table 4.

Head. Hair 4-C long and barbed with 9-15 branches; 5-C long and barbed with

11-16 branches; 6-C long and barbed with 9-12 branches; 14-C stellate with

4-7 branches; basal maxillary hair stellate and triple to 4 branched; mental

plate with 21-23 teeth. Antenna. Hair 1-A attached 0. 42 - 0.47 from base.

Thorax. Hair 0-P stellate with 11-17 branches; 1-P stellate with 7-11 branches;

o0-P barbed and triple to 4 branched; 9-P stellate and double to 5 branched;

1-M stellate with 8-13 branches; 4-M single to double; 14-M stellate with 7-10

branches; 1-T stellate with 8-11 branches; 3-T stellate with 11-17 branches.

Abdomen. Hair 1-I stellate with 10-14 branches; 2-I stellate with 8-10

branches; 4-I stellate with 11-15 branches; 6-I barbed with 4-5 branches; 10-I

double; 1-II stellate with 9-12 branches; 2-II stellate with 5-8 branches; 4-II

stellate with 12-14 branches; 13-II stellate with 9-12 branches; 1-III stellate

14 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

with 10-12 branches; 2-III stellate with 4-8 branches; 4-III stellate with 12-15

branches; 7-II stellate with 12-16 branches; 13-III stellate with 7-11 branches;

1-IV stellate with 12-14 branches; 2-IV stellate with 5-8 branches; 4-IV stel-

late with 13-17 branches; 7-IV stellate with 12-17 branches; 13- -IV stellate with

6-10 branches; 1-V stellate with 13-19 branches; 2-V stellate with 6-9 branches;

4-V stellate with 15-18 branches; 7-V stellate with 11-15 branches; 13-V stel-

late with 7-9 branches; 1-VI stellate with 15-18 branches; 2-VI stellate with

6-8 branches; 4-VI stellate with 16-18 branches; 7-VI stellate with 8-13

branches; 10-VI stellate with 3-4 branches; 13- -VI stellate with 8-11 branches;

1-VII stellate with 16-19 branches; 2- VII stellate with 5-8 branches; 3-VII stel-

late with 13-19 branches; 6-VII stellate with 5-8 branches; 8- VII stellate with

10-16 branches; 10- VII stellate triple to 6 branched; 13- -VIl stellate with 7-10

branches; 1-X long, barbed with 4-5 branches, each branch approximately

equal in length; 2-X moderately long with 10-13 branches; comb with 19-27

scales arranged in 2rows. Siphon. Index 3.30 - 3.84.

TYPE DATA. Ayurakitia griffithi D. Thurman, holotype male,

THAILAND, Chiengmai, Doi Sutep Mountain Range, Doi Chom Cheng Moun-

tain, Lemmon Cabin, 4 January 1953, collector Deed C. Thurman, Jr., col-

lected with a net while the mosquito was resting on the side of a tree growing ©

in a damp, cool, shady mountain valley around 3,000 feet elevation, U.S.N.M.

Type No. 62022; allotype, THAILAND, Chiengmai, Doi Sutep Mountain Range,

Doi Chom Cheng Mountain, 14 February 1953, Monop Rattanopradith collector,

on tree; 1 female paratype with same data as allotype; 1 male paratype with

same data as allotype except Deed C. Thurman, Jr. collector; 1 female para-

type same data as allotype except Tad Muey Falls added; 1 paratype male with

same data as holotype; 2 male and 1 female paratypes with same data as holo-

type except Sapria Valley, 24 February 1953, Deed C. and Ernestine B.

Thurman collectors, all the above specimens are in the U.S. National Museum

(Natural History) with the exception of one of the last mentioned male paratypes

and its location is unknown. One male paratype with the same data as the allo-

type is in the British Museum (Natural History).

DISTRIBUTION. Occurs in mountainous areas of Thailand north of

the Isthmus of Kra. Material examined: 17, 89, 1 P, 17 L, 11 with asso-

ciated skins (5 p, 61). All type material except 1 paratype was examined.

THAILAND. Chiang Mai: Buak Ha; Doi Sutep Mountain Range, Doi

Chom Cheng Mt.; Doi Sutep Huey; 16, ser 1 P, 10 p, 10 L, 61. Kanchana-

buri: Bo Ploy, Hin-Lub Village; io, 7 L,

TAXONOMIC DISCUSSION, The ee genitalia, pupa and larva are

discussed and compared in the discussion section of peytoni and the adults are

compared above.

The larva of griffithit tends to have more branches per hair on the head

and abdominal segment X and less branches per hair on the thorax and abdomi-

nal segments I- VIII than that of peytoni.

BIOLOGY. Immatures were collected from clear fresh water in the

axils of bananas and Pandanus; in heavily shaded areas located in secondary

deciduous forests; sites were in valleys located in mountainous areas; and at

an elevation of 3,920 feet in Chiang Mai Province. Larvae were collected in

association with the following species of mosquitoes: Aedes formosensis

Yamada, Topomyia inclinata Thurman and aenea Thurman. Adults were col-

lected resting among vegetation along a stream in Chiang Mai Province by

SEATO Laboratory personnel. Thurman (1954: 200) first collected this spe-

cies in Chiang Mai Province as adults which were resting on tree trunks ina

damp, cool, shady jungle valley in the mountains at an elevation of 3,000 feet.

Reinert: Aedes (Ayurakitia) in Southeast Asia 15

ACKNOWLEDGEMENTS

I am grateful to Dr. Botha de Meillon, Principal Investigator, South-

east Asia Mosquito Project (SEAMP) and Lieutenant Colonel Bruce F.

Eldridge, Chief of the Department of Entomology, Walter Reed Army Institute

of Research, for critically reading the manuscript. Special thanks are given

to the personnel of the Southeast Asia Treaty Organization Medical Research

Laboratory, Bangkok, Thailand, for the collection and preparation of many

specimens. Gratitude is expressed to Dr. Alan Stone, U.S. National Museum

(Natural History) and Dr. P. F. Mattingly, British Museum (Natural History),

for making available types and other specimens. Appreciation is expressed

to Miss Sheila Ford, Mr. Vichai Malikul and Mrs. Shuling Tung, SEAMP, for

preparing the illustrations and Miss Helle Starcke for typing the manuscript

oe na reproduction. I am especially grateful to my wife, Mollie, for typing

e drafts.

LITERATURE CITED

BELKIN, J.N.

1962. The mosquitoes of the South Pacific. Univ. Calif. Press,

Berkley, 2 vols. 608 and 412 p.

COHER, E.I.

1948(1949). A study of the female genitalia of Culicidae: With partic-

a Seas to characters of generic value. Ent. Am. 28(3):

75-112.

KNIGHT, K.L.

1970. A mosquito taxonomic glossary I. Adult head (external).

Mosq. Syst. Newsletter 2(1): 23-33.

KNIGHT, K.L., and J.L. LAFFOON.

1970a. A mosquito taxonomic glossary III. Adult thorax. Mosq. Syst.

Newsletter 2(3): 132-146.

KNIGHT, K.L., and J.L. LAFFOON.

1970b. A mosquito taxonomic glossary IV. Adult thoracic appendages.

Mosq. Syst. Newsletter 2(4): 165-177.

KNIGHT, K.L., and J.L. LAFFOON.

1971. A mosquito taxonomic glossary V. Abdomen (except female

genitalia). Mosq. Syst. Newsletter 3(1): 8-24.

MATTINGLY, P. F.

1971. Contributions to the mosquito fauna of Southeast Asia. XII.

Illustrated keys to the genera of mosquitoes (Diptera, Culicidae).

Contr. Am. ent. Inst. 7(4): 1-84.

STONE, A., K.L. KNIGHT, and H. STARCKE.

1959. A synoptic catalog of the mosquitoes of the world (Diptera,

Culicidae). Thomas Say Found. 6: 1-358.

THURMAN, D.C., JR.

1954. Avyurakitia, a new genus of mosquito from northern Thailand

(Diptera: Gulicidae), J. Wash. Acad. Sci. 44(6): 197-200.

16 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

THURMAN, E.H.B.

1959. A contribution to a revision of the Culicidae of northern

Thailand. Univ. Md. Agri. exp. Sta. Bull. No. A-100, 182 p.

LIST OF FIGURES

1. Distribution of the species of Ayurakitia

2. Aedes (Ayurakitia) peytoni - adult morphology

3. Aedes (Ayurakitia) peytoni and griffitht - adult morphology

4, Aedes (Ayurakitia) peytoni - female genitalia

5. Aedes (Ayurakitia) griffitht - female genitalia

6. Aedes (Ayurakitia) peytoni - male genitalia

7. Aedes (Ayurakitia) griffitht - male genitalia

8. Aedes (Ayurakitia) peytoni - pupa

9, Aedes (Ayurakitia) griffitht - pupa

10. Aedes (Ayurakitia) peytoni - larva

11. Aedes (Ayurakitia) griffithi - larva

“UR Fig. 1

LAOS

THAILAND

LY)

(

z CAMBODIA

Q S,

bee >

So ae

Ithmus of Kra

any A :

S | B

, e

° S

ab aa M@ GRI/FFITHI/

‘ “st @ PEYTON/

MALAYSIA

ieee

peytoni

peytoni griffith

CERCUS

POSTGENITAL

PLATE

' dorsal view

TERGUM IX

INSULA

ventral view

SPERMATHECAE

TERGUM VIII

STERNUM VIII

7 -— 0-1 4

peyton

@" ily \

UPA

Wildes (| As \ i

WAL GTS UK SAX: Ne Bi a>! 4

Wel Guys A K A] cs

Wipe CL 4G QMS oi

SN WAMAAAS OCD SS :

\ WL Li SS “ii | An

VIYT EL ELV: WN H

f Wabi 4 SOS) M, fei\\\ tH)

WW OGL SLLL TS Le if SQV Qn "

WU L/LEEL EDI SNA AY

WU IELLIA CONG RRA |

iy Vb ip toes SSD SY WW

ip SS

WY YS {Az ADEA SS SWS \

WIRE LOLIAE SSS Nd \

WINGO lG Ee Ss

y/ Wig) 7 STEALER SSNS YA nn

IOSD, BY A A IQS AQ

TAL ope SER, \ SSN QA.

Mies: C \\\

ny Wp AGL AG

AY MOA

We ER SN, j

\! jie ay Ny um WN SY Sh IN Wh Yq

ds , i MeN 1

17, RN SSSA HH! \ mh

ANN SSIN WT \ tlh f NT I

SENT | tl Mi ited tt

NES ty / RST TL

oS SAN

i \ yl Wi NUNN ty

I yy, ANUVCRNCR BHH E Io Mi i my ty

\\ Wh yl

Hl Wren yp Ly)

Wy diy My

}! Wi Mia) Maly

Ui Mi eal ip hi

Ui ! UU,

TERGUM

r—0-] ——4

dorsal view

ventral view

TERGUM VIII Snel?

STERNUM VIII

grittithi

TERGUM IX

GONOSTYLUS

ventral view

STERNUM IX

BASAL MESAL

LOBES

dorsal view

peytoni

TERGUM IX

GONOSTYLUS

dorsal view

STERNUM IX

SAL MESAL

LOBES

dorsal view

PHALLOSOME

ventral view

griftithi

arg

SG,

seu eaears

Bsitseeas

Fig.8

griffithi

28 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

TABLE 1. Record of the Branching of the Setae on the Pupae of

Aedes (Ayurakitia) peytoni

Hair Range Mode Mean Hair Range Mode Mean

Cephalothorax Abdomen II

1 2-4 3 3.1 0 1 1 1

2 2-3 2 Avid 1 6-14 9 So. 9

3 3-5 4 2 1 1 1

4 a3 3 3.4 3 1 1 1

5 3-5 4 4.1 4 4-9 4, D.3

6 1 1 1 9) 3-7 4 5

7 2 2 2 6 1 1 1

8 2-4 3 3.1 7 2-5 2 2.9

9 1-2 1 1,1 9 1 1 1

Metanotum 11 1-2 1 oe

10 3-8 5) d.1 Abdomen III

11 1 1 1 0 1 1 1

12 2-5 3 3.1 1 4-7 a 4.7

Abdomen I 2 1 1 1

1 17-34 24 25.6 3 1 1 1

2 1 1 1 4A 2-6 4. 3.8

3 1 1 1 5 25 L 3.7

4 4-8 5) 0.6 6 2-3 2 2.3

5 1-4 2 2.3 7 3-5 4 3.9

6 1 1 1 8 3-7 9) 4.6

7 1-2 1 1.1 9 1 1 1

9 1 1 1 10 2 2 2

11 1-2 2 1.9 11 1 1 1

14 1 1 1

Reinert: Aedes (Ayurakitia) in Southeast Asia 29

TABLE 1. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen IV Abdomen V (Cont. )

0 1 1 1 11 1 i 1

1 ard 9) 3.8 14 1 1 1

2 1 1 1 Abdomen VI

3 2-5 4 4.1 0 1 1 1

4 1 1 1 1 i=6 3 3k

5 1 1 1 2 1 1 1

6 1-3 2 2.7 3 1-3 1 1.9

7 2-6 3 3.1 4. 1-3 2 2

8 376 9) ae | 5 1 1 1

9 1 1 1 6 1-4 3 2. 4

10 1-2 2 1.9 7 1-2 1 Lak

11 1 1 1 8 3-8 4 4.9

14 ! 1 1 9 1 1 1

Abdomen V 10 1-4 2 2.7

0 1 1 1 11 1 1 1

1 2-6 5 4 14 1 1 1

2 1 1 1 Abdomen VII

3 1-3 1 1.9 0 1 1 1

4 2-3 2 2.2 1 i-D 2 2.5

9) 1-2 1 | 2 1 1 t

6 1-3 2 Lit 3 1-6 4 3.3

7 3-7 5) 4.9 4 2-3 2 2.2

8 2-7 4 4.1 5 1-4 2 2.3

9 1 1 1 6 4-11 6 6.1

10 1-2 2 bet q I-2 1 1,2

30 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

TABLE 1. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen VII (Cont. ) Abdomen VII

8 4-10 5 0.8 0 1 1 1

9 2-4 3 2.9 4 1-2 1 1.4

10 1-4 3 2.6 9 4-8 6 0.9

11 1-4 2 2 14 1-2 2 1.6

14 1-2 1 Ls Paddle

TABLE 2,

Reinert: Aedes (Ayurakitia) in Southeast Asia

Record of the Branching of the Setae on the Pupae of

Aedes (Ayurakitia) griffithi

Hair Range Mode Mean Hair Range Mode Mean

oO oN Oo OF FP WO po F

oN oo on FP ©C LO FK&

—_

bansk

Cephalotho rax

2-3 3

1-2 2

4-6 5)

2-4 3

1-2 1

2 2

2-7 4

1-3 2

Metanotum

2-10 3

1 1

2-4 3

Abdomen I

19-26 24

1-2 1

1 1

4-7 4

1-4 2

1 1

1-3 1

1 1

2.6

1.8

onrnren oa FF CO DPD KF OS

Abdomen II

1-2

= & Oo KF OO OO - & C

Abdomen III

— -=& po —|§ WHO KL PH HO WS KF KF CO -

32 | Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

TABLE 2. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen IV Abdomen V (Cont. )

0 1 1 1 11 1 1 1

1 2-4 3 3 14 1 1 1

2 1 1 1 Abdomen VI

3 2-5 5 3.8 0 1 1 1

4, ts 1 1 1 2-4 3 yal

5 2 2 2 2 1 1 1

6 1-3 1 a0 3 2-3 2 2.1

7 2-5 3 3.6 4 2-4 2 2.6

8 3-7 3 4 5 2 2 2

9 1 1 1 6 1-4 2 Bae

10 2-3 2 2.1 fi 1-2 1 tot

11 1 1 1 8 3-6 5 4.7

14 1 1 1 9 1 1 1

Abdomen V 10 1-3 2 1.9

0 1 1 1 11 1 1 1

1 1-4 3 3 14 1 1 1

2 1 1 1 Abdomen VII

3 2-3 2 2.3 0 1 1 1

4, 1-3 2 2.3 1 1-4 2 2.3

5 2 2 2 2 1 1 1

6 1-3 2 1.8 3 2-6 5) 4.1

7 4-8 6 0.6 & 2-4 2 2.6

8 4-7 5 D3 5 2-3 3 2.6

9 1 i 1 6 6-9 7 6.8

10 1-3 2 1.8 7 1-3 2 1.9

Reinert: Aedes (Ayurakitia) in Southeast Asia 33

TABLE 2. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen VII (Cont. ) Abdomen VIII

8 3-9 5) 5.8 0 1 1 1

9 4-7 5) 5.6 4. 2-4 2 2.6

10 1-3 2 2 9 13-20 18 I eek

11 1-4 1 1.8 14 1-4 1 act

14 1-4 2 2.2 Paddle

34 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

TABLE 3. Record of the Branching of the Setae on the Larvae of

Aedes (Ayurakitia) peytoni

Hair Range Mode Mean Hair Range Mode Mean

Antenna (A) Prothorax (P) (Cont. )

1 0-8 6 2.9 7 3-5 3 3.9

Head (C) 8 11-15 11 12.5

0 1 1 1 9 9-17 10 11.4

1 1 1 1 10 1-2 1 it

3 1 1 1 11 2-5 4 3.3

4: a- 7 7 6.3 12 1 1 1

5) a-9 8 12 14 1 1 1

6 6-9 7 7.3 Mesothorax (M)

7 9-14 11 11.5 1 20- 26 21 21.8

8 1-3 2 YF 2 2-3 3 2.9

9 oul Be 7 ote, 3 1 1 1

10 2-4 2 2.3 4 1 1 1

11 15-21 19 18.4 5) 1 1 1

12 2-4 2 2.2 6 6-8 7 6.8

13 4-7 6 D. t 7 1 1 1

14 7-14 7 8.2 8 7-12 9 8.9

15 2 2 2 9 8-10 9 9.1

bmh ae 4) 0.6 10 1 1 1

Prothorax (P) 11 1-2 1 ste

0 18-25 20 21.6 12 1 1 1

1 16-21 17 17.9 13 11-14 11 11.9

2 1 1 1 14 19-27 23 22.3

3 7-16 12 i ees Metathorax (T)

4 1 1 1 1 20-31 23 24.4

5 2 2 2 2 1 1 1

6 1 1 1 3 19-28 23 22.6

TABLE 3.

Reinert: Aedes (Ayurakitia) in Southeast Asia

(Continued)

Hair Range Mode Mean Hair Range Mode Mean

oOo oa NI OD OO eS

orn oa FF CO DW F

— >

o ~_ &

Metathorax (T) (Cont. )

2-3 2

2-4 3

1 1

8-15 10

6-11 8

6-8 7

1 1

1-2 1

1 1

20- 29 23

Abdomen I

23-34 25

13-17 15

1 1

16-18 17

1-2 2

6-7 6

1-2 1

1 1

12-15 14

17-24 19

Abdomen II

1 1

19-27 24

13-19 16

oOo wore oa _ Ww

ownonrenwsk# ft! Ww ND FF O

Abdomen II (Cont. )

1 L

17-24 19

1 1

4-5 &

14-16 16

1 1

2-3 3

1 1

17-24 19

1 1

Abdomen III

1 1

20-25 25

10-17 11

1 1

16-20 19

1 1

1-2 2

17-20 19

1 1

3-4 3

1 1

1929

36 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

TABLE 3. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen III (Cont. ) Abdomen V (Cont. )

13 15-19 17 16.8 6 2 2 2

14 1 1 1 7 16- 23 16 18.3

Abdomen IV 8 1 1 1

0 1 1 1 9 1 1 1

1 24-28 28 26.1 10 1 1 1

2 11-16 13 13 11 2-4 3 2.9

3 1 1 1 12 1 1 1

4 18-22 18 18.9 13 14-19 15 16

5 1 1 1 14 1 1 1

6 2 2 2 Abdomen VI

7 20- 24 21 21.7 0 1 1 1

8 1 1 1 1 19-30 28 28.8

9 1 1 1 2 13-23 15 16.3

10 1 1 1 3 1 i 1

11 2-4 2 2.6 4 20-27 23 22.8

12 1 1 1 5 1 1 1

13 14-20 L7 16.6 6 2 2 2

14 1 1 1 7 15-19 19 17.4

Abdomen V 8 2-3 3 2.6

i. 1 1 9 1 1 1

= bo

f= Oo

ao ©

igs bo

oo fee)

Les bo

» s

for)

—_

= ©

Lw) io)

t 1

oo eat

)

—_

(St) iw)

—_

an)

e e

ol H> (St) iw) Leroy)

oe)

1 —

INC) 1

bo iw)

—_

(oe) bs

—_

ite)

e ®

_—

—_

(St) Ww)

-—

—_

(op) =

j—

o1 fat

1 1 1 14 | 1 ct

Reinert: Aedes (Ayurakitia) in Southeast Asia 37

TABLE 3. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen VII Abdomen VUI

0 1 1 1 0 1 1 1

1 21-27 25 24.4 1 17-24 L7 19

2 14-18 16 15.9 2 1 1 1

3 20- 23 20 20.9 3 6-8 7 6.8

4 14-19 un 15.6 4 1 1 1

5 1 1 1 5 5-11 6 6.9

6 10-15 13 12.1 14 2-3 2 1.9

7 1 1 1 Abdomen X

8 18-25 18 19.7 1 1+3 2 1.8

9 3-5 4 é, 2 6-9 (3 y

10 7-14 9 9.3 3 1 1 1

11 37-5 3 3.8 Siphon

12 1 1 1 1 4-7 5 0.4

13 10-13 13 11,6 2 1 1 1

—

—_

38 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

TABLE 4. Record of the Branching of the Setae on the Larvae of

Aedes (Ayurakitia) griffithi

Hair Range Mode Mean Hair Range Mode Mean

Antenna (A) Prothorax (P) (Cont. )

1 93 6 6.4 i 3-4 3 3.4

Head (C) 8 7-11 8 8.8

0 1 1 1 9 2-5 e 4.3

1 1 1 1 10 1 1

3 1 1 1 11 2-6 4 4

4, 9-15 11 12.3 12 1 1 1

a 11-16 12 12.6 14 1 1 1

6 9-12 10 10.6 - Mesothorax (M)

7 12-17 16 14,5 1 8-13 13 11.2

8 5-9 7 6.8 2 2-4 3 2.9

9 11-17 15 14.7 3 1 1 1

10 2-4 3 3.3 4 1-2 2 1.9

11 15-22 16 18.1 5 1 1 1

12 2-3 3 2.6 6 7-10 8 8.1

13 4-6 5 4,7 1 1 1 1

14 4-7 6 2 8 1-9 9 8.4

15 2-3 3 2.6 9 6-10 10 8.8

bmh 3-4 3 3.4 10 1 1 1

Prothorax (P) 11 1-3 2 2.2

0 11-17 15 14.3 12 1 1 1

1 q-13 11 12.8 13 6-16 8 8.8

2 1 1 1 14 7-10 9 9.8

3 8-10 9 8.9 Metathorax (T)

4A 1-2 1 1.4 1 8-11 9 8.8

5; 3-4 3 aril 2 ‘ 1 1

6 1 1 1 3 11-17 11 14

TABLE 4,

Reinert: Aedes (Ayurakitia) in Southeast Asia

(Continued)

Hair Range Mode Mean Hair Range Mode Mean

o Oo NN GD HO fe

oN OD oO BP | DP

—_ =

Oo - Oo

Metathorax (T) (Cont. )

2-4 3

2-5 3

1 1

9-12 9

7-10 9

ts 8

1 t

1-3 1

1 1

16-22 18

Abdomen I

10-14 10

8-10 8

1 1

11-15 14

2 2

4-5 5)

1 1

1-2 1

2 2

10-14 11

9-14 11

Abdomen II

1 1

9-12 9

0-8 6

2.7

3.4

no wo NHN OO HO Ee ww

eo ma N Oo OF FPF OO PBF OS

Abdomen II (Cont. )

1-2 1

12-14 13

1 1

4-5 5)

13-18 15

1 1

2-4 3

1 1

9-12 9

1 1

Abdomen III

1 1

10-12 11

4-8 6

1 1

12-15 13

1 1

2 2

12-16 14

1 1

1 4

2-3 3

1 1

1.3

12.8

4.7

3.3

9.6

40 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

TABLE 4. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen III (Cont. ) Abdomen V (Cont. )

13 7-11 8 8.6 6 2 2 2

14 1 1 1 7 11-15 12 12.6

Abdomen IV 8 1 1 1

0 1 1 1 9 1 1 1

1 12-14 13 12.9 10 1 1 1

2 5-8 6 6.4 11 3-4 3 3.4

3 1 1 1 12 1 t 1

4 13-17 15 14.8 13 T-9 7 7.6

5 1 1 1 14 1 1 i

6 2 2 2 Abdomen VI

7 12-17 13 13.6 0 1 1 i

8 1 1 1 1 15-18 16 16.5

9 1 1 1 2 6-8 7 7.1

10 1 1 1 3 1 1 1

11 2-6 3 3.5 4 16-18 16 16.5

12 1 1 1 5 1 1 1

13 6-10 7 7.8 6 2 2 2

14 1 1 1 7 8-13 il 11

Abdomen V 8 2-3 2 2.4

0 1 1 1 9 1 1 1

1 13-19 14 14.6 10 3-4 4 3.6

2 6-9 6 ie 11 3-4 4 3.6

3 1-2 1 1.3 12 1 1 1

4 15-18 15 15.9 13 8-11 9 8.7

5 1 1 1 14 1 1 1

Reinert: Aedes (Ayurakitia) in Southeast Asia

TABLE 4,

(Continued)

Hair Range Mode Mean Hair Range Mode Mean

ao moO NN OO oO FP CF poe rF oo

—>~_ —- —_ ee

> CO DPB F- OO

Abdomen VII

1 1

16-19 18

0-8 8

13-19 15

10-15 13

1 1

o-8 6

1 1

10-16 13

O76 6

3-6 4

3-5 3

1 1

7-10 8

1 1

Gor a 60) pn =) ©

Abdomen VII

1 1

14-20 15

1 1

8-11 9

1 1

7-10 8

2-3 2

Abdomen X

4-5 4

10-13 10

1 1

Siphon

6-7 6

1 1

42 Contrib. Amer. Ent. Inst., vol. 9, no. 2, 1972

INDEX

Italicized pages are those which begin the primary treatment of the

taxon. Numbers in parenthesis refer to the figures illustrating the species

in question.

Abraedes 5

Aedes 1, 2, 4, 5, 6.

Aedimorphus Ds

aenea, Topomyia 14,

albipes, Orthopodomyia 12.

Anopheles 6.

aranoides, Tripteroides 12.

Ayurakitia dye, 4, Sy Gh oly 12, BAe 16, 17.

Aztecaedes as

Culex 6.

Diceromyia 5, 6.

Finlaya D's

formosensis 14,

genurostris, Malaya 12.

eriffithi 2, 6, YT, b44-1 2,15, 14, 16 Cs 94

Tey alt,

inclinata, Topomyia 12, 14.

Kompia Ea

Lophoceraomyia, Culex 12.

Malaya 6.

meronephada De

Neomelaniconion D.

nummatus De

Ochlerotatus De

Orthopodomyia 6.

Pandanus 6)4h0,. 11, a2, 14.

peytoni 1, 4, 6; 7, 10, 12; 12, 13, 14,. To, 28,

3412,.3,. 4,°6,.8, 10)

poicilus 12,

Protomacleaya D.

sintonoides, Anopheles 12.

Stegomyia Dis

Topomyia 6.

Tripteroides 6.

Udaya De

vexans Group os

vittatus De

Contributions

of the

American Entomological Institute

Volume 9, Number 3, 1973

MOSQUITO STUDIES (Diptera, Culicidae)

XXIX. A review of the subgenus Kerteszia of Anopheles.

By Thomas J. Zavortink

XXX. A new subgenus and species of Culex from Colombia.

By Abdiel J. Adames and Pedro Galindo

oo tee |

Bee) Si

fae,

% :

ie af is

MOSQUITO STUDIES (Diptera, Culicidae)

XXIX. A REVIEW OF THE SUBGENUS KERTESZIA

OF ANOPHELES'

Thomas J. Zavortink”

INTRODUCTION

Kerteszia is a small, Neotropical subgenus of Anopheles. Although several species

of this subgenus are important, even primary, vectors of human malarias and other

species are suspected vectors, Kerteszia is appallingly poorly known systematically.

Because of the dearth of specimens available for study, the present paper can be

little more than a preliminary review laying the groundwork for a future, more com-

prehensive study.

Only 1146 specimens, 97 males, 64 male genitalia, 538 females, 294 larvae and

153 pupae, were examined during this study; included were 155 individual rearings

(100 larval, 28 pupal and 27 incomplete). Most of the adults studied, including the

reared ones, are in very poor condition, either broken, denuded, shriveled, fungused,

decomposed or imbedded in adhesive, and are not of research quality. Many of the

skins of the immature stages are poorly preserved or poorly mounted and almost all

whole larvae examined died during the attempt to rear them and are rotted, mis-

shapen and discolored. Only the Colombian and Panamanian populations of neivai

and the Trinidadian populations of bellator and homunculus are represented by even

modest series of adults and immatures, and only 2 of these populations, the Panama-

nian neivai and the Trinidadian bellator, are topotypic or nearly so.

Classical, comparative morphological taxonomic procedures were used. Many of

the conclusions reached must be considered as tentative only, for in most cases

neither the quantity nor quality of material available for study is adequate for test-

ing the hypotheses about the number of species and their diagnostic characters that

are summarized in the keys. The descriptions are based on a few individuals from,

if possible, one population, as indicated under the species; the organization, termi-

nology and abbreviations used in the descriptions follow Belkin (1962) in large part.

When available, the wing, hindtarsus, male genitalia, pupa and larva of each species

are illustrated; all these illustrations, including those of the immature stages, are

based on a single, representative specimen only. The collection data for all speci-

mens examined are given for each species and the localities are marked with solid

symbols on the outline maps. Distribution records from the literature are listed

separately and the localities of the more reasonable, reliable or better documented

‘Contribution from project “Mosquitoes of Middle America” supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478.

* Department of Biology, University of California, Los Angeles, California 90024.

a Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

records are marked with open symbols on the maps. Many literature records, even

modern ones appearing after Komp’s 1937 revision of Kerteszia, are undoubtedly

based on misidentifications or the uncritical repeating of previous incorrect litera-

ture records.

I thank John N. Belkin for reading and criticizing the manuscript; Willis W. Wirth

for arranging the loan of specimens from the National Museum of Natural History

[USNM]; Peter F. Mattingly for loaning all Kerteszia, including the lectotype of

Anopheles lutzii Theobald, 1901 in the British Museum of Natural History [BMNH];

Alfonso Diaz Najera for loaning specimens in the Instituto de Salubridad y Enferme-

-dades Tropicales [ISET]; Sebastiao H. Xavier for loaning specimens in the Instituto

de Endemias Rurais [IER]; Pablo Cova Garcia for loaning specimens in the Division

de Endemias Rurales [DERM]; William A. Powder and Sandra J. Heinemann for

preparation of the material collected for the project “Mosquitoes of Middle Amer-

ica’; L. Margaret Kowalcyzk and Nobuko Kitamura for preparation of the prelimi-

nary drawings and final illustrations; Caryle Stallard for typing a portion of the

rough draft; and Angeliki Demos for typing the remainder of the rough draft and

preparing the text copy for lithoprinting.

SYSTEMATICS

I am recognizing 10 species of Kerteszia. Two of these, boliviensis, the type spe-

cies of the subgenus, and the unnamed species 10, are known only as females; the

remaining 8 species are known, at least to a limited extent, in all stages except some-

times the pupa.

Anopheles neivai, the dominant or only species throughout the northern portion

of the range of the subgenus, is differentiated from all other species by several

derived characters, especially conspicuous of which are the form of the ventral lobe

of the claspette of the male genitalia, the short, weakly developed hair 4a-X and

absence of hair 8-S in the larva, and the absence of hair 2-P in the pupa. Anopheles

pholidotus and lepidotus are a pair of closely related species, the former known

from only western Panama and western Venezuela and the latter from the east side

of the Andes in Colombia and Bolivia, that share several derived or unusual features,

such as apical transverse bands of outstanding scales on the distal tergites and a

large, vertical patch of scales on the mesepimeron in the adult, and the nonpalmate

hair 1-VII and moderately long hair 6-VI in the larva. Both these species, but espe-

cially pholidotus, share a number of derived features in the immatures (pupa of

lepidotus unknown) with neivai, and are apparently related to it. Conspicuous larval

features possessed by these 3 species are the nonpalmate hair 1-I and the uniformly

long pecten teeth. Additional derived features shared by pholidotus and neivai are

the long, wavy, filamentous marginal spicules on the pupal paddle, and the small,

brushlike palmate hairs and the fringe of spinules extending to near the apex of both

the inner and outer edges of the pecten teeth in the larva. Anopheles pholidotus and

lepidotus share the possession of abdominal scales with boliviensis and the presence

of a white subcostal spot on vein R,,3 and a submedian white scaled line on R445

(not always present) with boliviensis and other species mentioned below.

Anopheles bambusicolus is, in many respects, the most unusual Kerteszia; unique

features are the usually reduced number and size of the white spots on the costal

vein of the wing and the entirely white scaled hindtarsal segment 5 in the adult, the

nearly glabrous ventral claspette lobe in the male genitalia, the short hair 9-V and

Zavortink: Subgenus Kerteszia 3

broad, elliptic paddle in the pupa, the large palmate hairs with long, lanceolate

leaflets and short hairs 6-VI and 5-7-C in the larva, and, of course, its bambusicolous

habits. The affinities of bambusicolus may be with neivai, pholidotus and lepidotus;

characteristics shared by all these species are a single patch of scales on the mese-

pimeron, the absence of leaflets on the aedeagus, the presence of marginal spicules

distad of the external buttress on the pupal paddle, and a weakly developed hair 4-C

in the larva. In addition, bambusicolus shares with neivai a small patch of scales on

the mesepimeron and the absence of a median or submedian white scaled line on

vein R445 in the adult; with pholidotus and lepidotus a shortened hair 6-VI in the

larva; with pholidotus a more or less similarly shaped and spiculed ventral claspette

lobe; and with lepidotus a short hair 11-P and enlarged palmate hairs. Some of the

morphological peculiarities of bambusicolus are found also in Nyssorhynchus, which

suggests that bambusicolus is the most primitive Kerteszia.

Anopheles homunculus, cruzii, bellator and laneanus, the 4 remaining species of

Kerteszia in which the male genitalia are known and the immatures are at least

partly known, form a group characterized, but not always absolutely separated from

the previously considered species, by the presence of 2 patches of scales on the

mesepimeron and a short to long, median or submedian white scaled line on vein

R445 in the adult, the presence of leaflets on the aedeagus in the male genitalia

(leaflets secondarily lost in some individuals of homunculus), the absence of mar-

ginal spicules distad of the external buttress on the pupal paddle (pupae of cruzii

and laneanus unknown), and the strong hair 4-C, palmate hair 1-I and alternating

long and short pecten teeth with spinules restricted to the basal portion of the outer

edge in the larva (the last character not discernable in specimens of Janeanus and

cruzii available for study). These 4 species fall into 2 pairs: homunculus and cruzii,

distinguished by the predominantly dark large acrostichal and dorsocentral setae of

the adult, the shape of the ventral claspette lobe of the male genitalia, and the pre-

dominantly basally branched hair 5-II-V of the larva; and bellator and laneanus,

characterized by the predominantly pale large acrostichal and dorsocentral setae, the

shape of the ventral claspette lobe, and the predominantly plumose hair 5-II-V in

the larva. Anopheles homunculus and cruzii are very similar in external adult fea-

tures and apparently can be reliably separated only by the male genitalia (larva of

cruzii not studied by me and pupa unknown). Which of these species is the more

derived is not obvious to me at this time: the hypertrophied ventral claspette lobe

and reduced leaflets of the male genitalia of homunculus are certainly derived fea-

tures, but this species is apparently uncommon except locally throughout its exten-

sive South American range; cruzii, on the other hand, is a common, dominant

Kerteszia throughout its range in southeastern Brazil and occurs in a greater variety

of habitats, including harsher, drier ones, than homunculus. Anopheles bellator and

laneanus are easily separated by external adult features as well as by the male genita-

lia (larva of Janeanus not studied by me and pupa unknown). Anopheles bellator is

unquestionably the more derived of these 2 species: it is characterized by derived

morphological features in the adult and is the more common and widespread spe-

cies.

The affinities of boliviensis cannot be determined in the absence of males and im-

matures. This species possesses abdominal scales, as do pholidotus and lepidotus,

and has only a single, small, upper patch of scales on the mesepimeron, as do neivai

and bambusicolus. However, the ornamentation of the scutum, hindleg and wing,

a Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

including even the unusual occurrence of white scales in the basal part of vein M, is

almost identical to that of laneanus and unlike that of neivai, bambusicolus, pholi-

dotus and lepidotus. The single female of species 10 is so badly damaged that most

of the morphological characteristics of even the adult stage are unknown.

The bromelicolous species of Anopheles have been recognized as a distinct group

since the works of Knab (1913:15-17) and Dyar and Knab (1917:38-40) and their

relationship to Nyssorhynchus has been known since the early studies of Root

(1922a:322; 1922b:388; 1923:271, 277-278). Edwards (1932:44,46), following

Christophers (1924:42), included Kerteszia in the subgenus Nyssorhynchus but

Dyar (1928:467-470) and Komp (1937:492-529; 1942:8-9, 162-165) treated it ds a

distinct subgenus. For the time being I am following the currently accepted classifi-

cation, as reflected in Stone, Knight and Starcke (1959), and am recognizing this

monophyletic group as a distinct subgenus of Anopheles. However, the relationship

of Kerteszia to Nyssorhynchus may prove to be so close that the former will again

be recognized as only a species group of the latter. Little can be done to clarify this

Situation at the present time because the systematics of Nyssorhynchus are too

poorly known and hopelessly confused.

BIONOMICS AND MEDICAL IMPORTANCE

With one exception the species of Kerteszia normally breed in the water that col-

lects in the leaf axils of terrestrial and epiphytic bromeliads. The single exception is

bambusicolus, which normally breeds in unbroken bamboo internodes. In Trinidad,

where the ecological relationships between sympatic species of Kerteszia have been

thoroughly studied, homunculus and bellator are associated with different brome-

liad species in that part of the island where their distributions overlap, homunculus

breeding in small, shade-tolerant species near the forest floor and bellator in large,

heliophilous species in the canopy. It is possible that similar associations operate to

reduce competition between bromelicolous species of Kerteszia in other areas where

2 or more are sympatric.

The adults of Kerteszia are strongly anthropophilic, biting especially during the

evening but also sometimes in the shade during the day, throughout the night and in

the morning. The periods of adult activity, the microdistributions of individuals and

demes, and possibly even the macrodistributions of populations and species are de-

termined, at least to some extent, by humidity. For example, in Trinidad homuncu-

lus requires wetter conditions than bellator: it is largely restricted to the part of the

island with the highest rainfall, it occupies the lower strata of the forest and its ac-

tivity is more closely limited to the moister periods of the day.

The populations of Kerteszia that attain very high densities are of considerable

importance as pests and vectors of human, and probably also animal, diseases. The

Colombian population of neivai, the Brazilian and Trinidadian populations of bell-

ator and homunculus, and the Brazilian population of cruzii are important, proven

vectors of human malarias, and the Ecuadorian population of “‘boliviensis’’, the

Guyanan population of bellator, and the Bolivian population of Janeanus are sus-

pected vectors. Anopheles neivai has also been found naturally infected with the

viruses of yellow fever, Guaroa and possibly also Venezuelan equine encephalomy-

elitis and Ilheus; “‘boliviensis’’ with the viruses Anopheles A and Anopheles B and

the eges of Dermatobia hominis (Linnaeus); and, bellator with the filarial worm Wu-

chereria bancrofti (Cobbold). Anopheles cruzii is also a proven vector of monkey

malarias.

Zavortink: Subgenus Kerteszia 5

DISTRIBUTION

The known distribution of Kerteszia extends south from the State of Veracruz in

Mexico through Central America and Atlantic South America, along the Andes and

along the coast, to the States of Misiones in Argentina and Rio Grande do Sul in

Brazil; it also extends south along the Pacific Coast of South America to the State of

El Oro, Ecuador. The subgenus is conspicuously absent from all islands of the West

Indies except Trinidad and from most of the vast expanse of the Amazon basin in

South America.

TAXONOMIC TREATMENT

Subgenus KERTESZIA Theobald

1905. Kerteszia Theobald, 1905b:66. TYPE SPECIES: Kerteszia boliviensis Theobald, 1905,

Bolivia; monobasic.

1918. Dendropaedium Dyar and Knab, 1918:141. TYPE SPECIES: Anopheles cruzii Dyar &

Knab, 1908 (=Anopheles lutzii Theobald, 1901), Brazil; monobasic. Synonymy with

Kerteszia by Christophers (1924:42).

Anopheles (Kerteszia) of Dyar (1918:148; 1925b:193; 1928:467-470); Dyar and Knab (1918: 140-

141); Komp (1937:492-529; 1942:8-9, 162-165); Lane (1939:17-19; 1953:278-288); Vargas

(1943:60-63); Senevet (1958:141-144); Stone, Knight and Starcke (1959:35-36); Forattini

(1962:433-462).

Kerteszia of Theobald (1905b:66; 1907:117; 1910:74).

Anopheles (Dendropaedium) of Dyar and Knab (1918:141); Dyar (1918:145-146; 1925a:25-27).

Anopheles (Nyssorhynchus) Kerteszia Group of Christophers (1924:42); Edwards (1932:44,46).

Anopheles (Nyssorhynchus) Dendropaedium Group of Root (1922a:322; 1922b:388; 1923:271,

277-278).

Anopheles (Anopheles) in part of Bonne and Bonne-Wepster (1925:504-507,533).

Anopheles in part of Theobald (1901:177-178); Giles (1902:303-304); Dyar and Knab (1917:38-

40); Howard, Dyar and Knab (1917:985-988); Dyar (1923:72); Cova Garcia (1961:57-62, 104-

108, 121-136).

Myzomyia in part of Theobald (1905a:8; 1907:41-43; 1910:16,17).

Laverania in part of Theobald (1902:183).

Nyssorhynchus in part of Blanchard (1905:211).

FEMALES. Head: Interocular space narrow. Vertex and occiput with numerous

short, broad erect scales, these dark brown to black except for white in median ante-

rior patch and sometimes in partial or complete orbital line or small lateral patch;

frontal tuft moderately conspicuous, white. Proboscis longer than forefemur; entire-

ly dark scaled. Palpus subequal in length to proboscis; predominantly dark scaled,

with dorsal white patch at apex of some or all of segments 2-5; at least segments 1

and 2 with scales outstanding. Torus without scales; flagellar segment 1 with small,

dark scales; proximal flagellar segments with bristles on inner side greatly reduced in

size. Thorax: Scutum shortened and arched. Integument of scutum pruinose with 2

pairs of darker, nonpruinose longitudinal stripes when viewed at an angle from the

front, as follows: (1) narrow, submedian stripe laterad of acrostichal bristles extend-

ing from anterior promontory to prescutellar space, and (2) broad, irregular, sub-

dorsal strip laterad of dorsocentral and lateral prescutellar bristles extending from

6 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

anterior promontory to scutellum. Acrostichal and dorsocentral bristles numerous,

strongly developed, moderately long to long. Scutum with light scales (1) in median

and lateral patches on anterior promontory, (2) in lateral prescutal line, (3) in ante-

alar and supraalar areas, (4) sometimes scattered in fossal or posterior fossal area,

and (5) sometimes scattered in anterior portion of acrostichal and dorsocentral ar-

eas. Scutellum without scales or with a few scales medially. Pleuron with bristles on

apn, ppl, sp, stp, pra and upper mep, ppl bristles very few (1 or 2); stp bristles very

few (1-4), arising in upper scale patch. Pleuron with scales on apn, in upper patch or

line on stp, in small lower patch on stp, on pra, in 1 or 2 patches on mep, and some-

times on sp. Legs: Forecoxa and midcoxa with numerous scales. Hindfemur without

outstanding scales distally. Knee spots absent. Legs dark scaled with conspicuous,

complicated light markings, principally complete or broken longitudinal lines or

streaks on femora and tibiae and complete or incomplete transverse bands on tarsi;

hindtarsal segments 3-5 never all entirely white. Wing: Dark scaled with distinct light

markings. Vein R-R, and usually also vein C with basal, humeral, presectoral, sec-

toral, subcostal and preapical light spots. Veins R;, M and Cu, distad of crossveins,

Cu, and 1A predominantly or entirely dark scaled. Dark scales not grouped into

spots. Fringe dark scaled with 2-5 light spots. Haltere: Knob dark scaled with white

scales in basal anterior portion. Abdomen: Tergites and sternites II-VII with or with-

out obvious scales. Tergites without conspicuous caudolateral tufts of outstanding

scales. Buccopharyngeal Armature: Present, not studied.

MALES. Essentially as in females except for sexual characters. Head: Palpus with

dorsal white patch at apex of segments 4 and 5 and often also segments 2 and 3;

segments 3 and 4 with scales entirely or predominantly decumbent. Wing: Light

scaling of veins behind R-R, often reduced in extent or indistinct.

MALE GENITALIA. Segment [X: Tergite without lobes. Sternite with strong,

median longitudinal ridge; distal margin usually concave laterad of ridge. Sidepiece:

Moderately long, cylindrical to long conical, curved inward. Tergomesal surface

membranous from base to near apex. Basal tergomesal area with 1 very long, strong,

curved, blunt parabasal spine arising from a large tubercle; distal tergomesal surface

with 2 long, strong, apically flattened accessory spines arising about 0.6-0.7 distance

from base of sidepiece; mesal surface with 1 long, strong internal spine arising about

0.4-0.5 distance from base of sidepiece; lateral surface with numerous bristles and

scales. Claspette: Divided into well developed, paired dorsal and ventral lobes. Dor-

sal lobe with 2 groups of 3-5 very long, slender, contorted, flattened setae. Ventral

lobe usually expanded laterally distally; with few to numerous spicules on mesal

margin and usually also sternal surface. Clasper: Long, slender. Spiniform short.

Phallosome: Aedeagus long, slender, nearly straight; with or without 1 pair of sim-

ple, slender, subapital leaflets directed cephaloventrad. Proctiger: Anal lobe broad.

Paraproct weakly sclerotized.

PUPAE. Setae generally weakly to moderately developed, short, few branched

and lightly pigmented. Trumpet: ‘“‘Culicine” in form, not strongly flared and not

deeply cleft. Abdomen: Hair O-II-VII weakly developed, short to moderately long,

usually single. Hair 1-II-VII weakly developed, short to moderately long, usually

1-4b or 4f (1-5). Hair 5-III-VII weakly developed, short to moderately long, 1-5f or

5b or weakly plumose. Hair 9-I weakly developed, short; 9-IV, V varied, short, blunt

and peglike to long, pointed and spinelike; 9-VI-VIII long, pointed, spinelike, lightly

pigmented, strongly barbed on outer edge except in bambusicolus. Terminal Seg-

ments: Apex of male genital lobe with mammillate protuberance.

Zavortink: Subgenus Kerteszia 7

LARVAE. Head: Inner clypeals (2-C) widely spaced, single, simple or weakly plu-

mose apically. Outer clypeals (3-C) single, simple or with a few strong, irregular

barbs apically. Posterior clypeals (4-C) laterad of outer clypeals, weakly to moder-

ately developed, single, simple or with a few strong, irregular barbs apically. Hairs

5-7-C usually moderately long to long, single, simple, weakly plumose apically or

with a few strong, irregular barbs apically; short to moderately long in bambusi-

colus. Hairs 8-10-C short to moderately long, usually single and simple, rarely 2,

3f or 2, 3b. Hair 11-C long, simple, with a few strong, irregular barbs apically or

strongly plumose apically. Antenna: Hair 1-A short, single, simple, arising on dorso-

lateral surface in basal portion of shaft. Thorax: Hairs 1-P and 3-T not palmate. Hair

9 and/or 10-P,M,T weakly to very weakly plumose. Hair 11-P weakly to strongly de-

veloped, short to long. Abdomen: Hair O-II-VIII weakly developed, short, single.

Hair 1-I or II-VI or VII palmate. Hair 2-II-VI usually single and simple, strongly de-

veloped and long on IV. Hair 6-III-V long, single, weakly to moderately plumose.

Spiracular Lobe: Pecten teeth (18-23) all long or alternate teeth in middle of row

shortened. Spiracular apparatus small; median dorsal and dorsolateral lobes without

specialized process. Hair 1-S single, 2, 3f or 2, 3b. Anal Segment: Ventral brush with

9 pairs of hairs.

KEYS TO SPECIES

FEMALES

(Species /0 not included)

ie Tergites and sternites II-VII with numerous scales : az

Tergites and sternites II-VII without obvious scales. . . ...... .4

2(1). Hindtarsal segment 2 with broad white band in apical 0.5-0.7; mesepimeron

with small upper patch of scales; scales of distal tergites not outstanding

and not forming transverse apical bands. . ... . . . 9. boliviensis

Hindtarsal segment 2 with narrow white band in apical 0.1-0.2; mesepimeron

with large, curved patch of scales extending ventrad from bristles to below

middle of segment; scales of distal tergites predominantly outstanding and

apgresated: inte transverse apica: Bands «ee; eee aS

3(2). Scales of proximal tergites predominantly narrow to moderately broad

. . 2. pholidotus

Scales of silat tergites eredorsinantly moderately broad to broad. ,

3. lepidetus

4(1). Mesepimeron with only upper patch of scales; vein R445 with light scales

absent or restricted to small basal spot. . .. . fag

Mesepimeron with both upper and middle patches of scales: 1 vein nR, 45 with

light scales in basal spot and short to long, median or submedian line . °°.-6

5(4). Hindtarsal segment 5 with broad white band in apical 0.4-0.6; vein C with

presectoral, sectoral and subcostal light spots usually present, moderately

large to large; palpus with scales slightly to conspicuously outstanding

6(4).

7(6).

8(7).

3(2).

4(3).

Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

on proximal portion of segment 3, decumbent on distal portion of seg-

ment 3 and onsegment4. .. . . . L. neivai

Hindtarsal segment 5 entirely white neaiods. vein Cc with sirenectaral: sectoral

and subcostal light spots usually absent, small when present; palpus with

scales conspicuously outstanding on segments 3 and 4. dniasienecrs: jl x

. 4. bambusicolus

Hindtarsal segments 2 and 3 with narrow white band in apical 0.1-0.3, seg-

ment 4 with narrow white band in apical 0.2-0.3, and segment 5 usually

entirely dark scaled. . . . 7. bellator

Hindtarsal segments 2-5 with broad white band i in n apical 0. 4- 5 ea a ee

Larger acrostichal, dorsocentral and middle scutellar bristles predominantly

pale; anterior 0.3-0.4 of acrostichal and dorsocentral areas and middle of

scutellum with a few white scales; vein M entirely or partially white scaled

basad of level of furcation in veinCu .. . . . &. laneanus

Larger acrostichal, dorsocentral and middle scutellar bristles predominantly

dark; acrostichal and dorsocentral areas without scales and middle of

scutcium without scales or with a few dark scales; vein M dark scaled

Dasacdat level of turcationin yeir-Cuens elec el kok eS oui eos 48

Palpus with scales slightly to moderately outstanding on segment 3 and

slightly outstanding to decumbent on segment 4; palpus with white patch

at apex of segments 3-5, segments 4 or 5 or segment 4, the patch largest

on 4 when present on more than 1 segment; abdominal tergites tan to

dark brown or blackish. . . . . . . 3S. homunculus

Palpus with scales entirely or predominantly decumbent on segments 3 and

4, sometimes slightly outstanding at base of segment 3; palpus with white

patch at apex of segments 3-5, the patch on 3 subequal in size to or larger

than patch on 4; abdominal tergites sient to conspicuously reddish

. 6. cruzii

MALE GENITALIA

(9. boliviensis and species 10 unknown)

Alosms without leat ete tn ete ing he gat Bae cpees he, teckel so 2

medecqeue with 6ydw of leaflets <e-.ko med aed Se aie seek ae ew oO

Ventral lobe of claspette glabrous except for 4-10 strong spicules and a few

weak spicules along mesal margin. . . : . . . 4. bambusicolus

Ventral lobe of claspette moderately to Aico spiculose, with many of the

spicules arising from ceneral mari Aced 4 tuisei.04) Hoste en eae Ce we wD

Tergite VIII without numerous broad scales medially. . . ..... .4

Tergite VIII with numerous broad scales medially . ........ 2.9

Lateral expansion of ventral lobe of claspette very narrow, inconspicuous,

not curved ventrad posteriorly and not retrorsely produced anteriorly;

Zavortink: Subgenus Kerteszia 9

internal spine of sidepiece flattened and conspicuously broadened at apex

. 1. neivai

Lateral ‘expansion of ventral lobe of ‘claspette broad, conspicuous, curved

ventrad posteriorly and produced into sharp retrorse point anteriorly;

internal spine of sidepiece flattened and slightly broadened at apex

5. homunculus

5(3). Lateral expansion of ventral lobe of claspette rounded and not retrorsely

produced anteriorly; internal spine of sidepiece flattened at apex a: %

$i pholidotus

Lateral eae of ental ae ar claspette Salina: and broadly emar-

ginate and produced into blunt retrorse lobe anteriorly; internal spine of

sidepiece apparently not flattened atapex . .... . . .3. lepidotus

6(1). Lateral expansion of ventral lobe of claspette produced into sharp retrorse

point anteriorly; leaflets of aedeagus weak, short or long. 5. homunculus

Lateral expansion of ventral lobe of claspette not retrorsely produced ante-

riorly or with blunt retrorse lobe; leaflets of aedeagus strong, long . . .7

7(6). Lateral expansion of ventral lobe of claspette more or less rounded to sinu-

ous-margined, not curved ventrad posteriorly, sometimes produced into

blunt retrorse lobe anteriorly . . . . 6. cruzii

Lateral expansion of ventral lobe of claspette rounded, ounved ventrad pos-

teriorly, not retforsely produced. anteriorly «20 4.404... ete. 8

8(7). Ventral lobe of claspette densely spiculose only mesally. . . . 7. bellator

Ventral lobe of claspette very densely spiculose except laterally . 8. laneanus

PUPAE

(3. lepidotus, 6. cruzii, 8. laneanus, 9. boliviensis and species 10 unknown)

ke Hair 9-V short, blunt and peglike, similar to 9-III; outer edge of paddle

distad of external buttress with short marginal spicules; paddle elliptic,

relatively broad, index 1.4 . . . . 4. bambusicolus

Hair 9-V moderately long to long, pointed ‘and spinelike, dissimilar to 9-IIT;

outer edge of paddle distad of external buttress with long filamentous

marginal spicules or without any spicules; paddle obovate, relatively

PALTOW AMON Fuh oe ial ae aka remem Nes) ata a gS

2(1). Outer edge of paddle distad of external buttress with long, wavy, filamen-

tous marginal spicules; paddle usually more or less obliquely truncate. . 3

Outer edge of paddle distad of external buttress without marginal spicules;

paddle usually not obliquely Wiuncate 63s a we OO ee Ge

3(2). Marginal spicules along distal portion of external buttress of paddle pre-

dominantly toothlike, short, few, widely spaced, and not extending basad

into proximal 0.5 of paddle; hair 1-P weakly to moderately developed,

moderately long; 2-P absent; 1-VII arising near caudal border of segment

. 1. neivai

10 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

Marginal spicules along distal portion of external buttress of paddle filamen-

tous, moderately long, numerous, closely spaced, and extending basad

into proximal 0.5 of paddle; hair 1-P strongly developed, short; 2-P pre-

sent; 1-VII arising cephalad of caudal border of segment. . 2. pholidotus

4(2). Paddle unpigmented to very weakly pigmented, lighter than posterior ab-

dominal segments; marginal spicules along distal portion of external but-

tress of paddle moderately long, numerous and closely spaced. :

; : a hariuncalns

Paddie weakly to rongly pigmented, “darker ‘than posterior abdominal seg-

ments; marginal spicules along distal portion of external buttress of pad-

dle short, few atic widely spaced 2) OO ee PO 2 7, Dellator

LARVAE

(9. boliviensis and species 10 unknown; 6. cruzii

and 8. laneanus not included)

l. Hair 6-VI short; palmate hairs large, with long, lanceolate leaflets; 5-7-C

short to moderately lorig 2013 . . 4. bambusicolus

Hair 6-VI moderately long to long: palmate hans staal to moderate, with

short to moderately ie siherss or blunt leaflets; 5-7-C age long

tOONe SS OP es i er a ee CZ

2(1). Pecten teeth all long, with spinules usually extending to near apex of both

internal and external edges; hair 1-I not palmate, 1-4b or 4f; hair 4-C

weaker than 2-00.52... ss

Pecten teeth alternating long and short “medially, with Singles usually re-

stricted to basal portion of external edge; hair 1-I palmate; 4-C as strong

ao OF stromuer dimmer ee ee ae ce oe Oe

3(2). Hair 6-VI long, plumose, similar to 6-IJI-V; hair 1-VII palmate; most caudal

hair of ventral brush (4a-X) weakly developed, shorter than anal saddle,

usually simple . . . . . 1. neivai

Hair 6-VI moderately — with a ae long Bianchion® in basal portion, dis-

similar to 6-III-V; hair 1-VII not palmate, 1-4b or 4f; most caudal hair of

ventral brush (4a-X) moderately developed, usually longer than anal sad-

dle distinctly pectinate-or pluitioses 2/4 Fo 8 Se,

4(3). Hair 11-P strongly developed, long; 3-C moderately long, moderately de-

veloped, not fusiform; palmate hairs small . . . . . . 2. pholidotus

Hair 11-P weakly developed, short; 3-C short, strongly developed, fusiform;

palmate hairs moderate .... ; . . . Jd. lepidotus

5(2). Most caudal hair of ventral brush (4a-X) usually weakly developed and short-

er or slightly longer than anal saddle; hair 3-C much stronger than 2-C;

hair 5-II-V usually branched near base only; 4-A usually single :

28 Ds homunculus

Most caudal fair of ventral brush moderately developed and much longer

Zavortink: Subgenus Kerteszia 11

than anal saddle; hair 3-C slightly stronger than 2-C; hair 5-II-V usually

distinctly plumose; 4-A usually forked apically .. . . . . 7. bellator

I. Anopheles (Kerteszia) neivai Howard, Dyar & Knab

Figs. 1,3-7

1913. Anopheles neivai Howard, Dyar and Knab, 1913:plate 41, fig. 8, and plate 130, fig. 461.

TYPE: Lectotype female (no. 344.1) with slide of associated larval head capsule and

pupal skin, Fort San Felipe, Porto Bello [Portobelo] Bay, Colon, Panama, larva collected

in bromeliad leaf axil, 2 June 1908, A.H. Jennings [USNM, 20440; designation by Stone

and Knight, 1956:279]. Synonymized with Jutzii Theobald, 1901 (as bellator) by Dyar

(1923:72); considered as variety of lutzii (as bellator) by Christophers (1924:42); con-

sidered as race of lutzii (as cruzii) by Dyar (1925a:26); resynonymized with lutzii (as

cruzii) by Dyar (1925b:193); resurrected to specific rank by Komp (1937:502).

1917. Anopheles hylephilus Dyar and Knab, 1917:38-40. TYPE: Lectotype female, Gatun,

Canal Zone, Panama, Feb 1917, L.H. Dunn (W-38) [USNM; designation by Belkin,

Schick and Heinemann, 1965:44]. Synonymized with lutzii Theobald, 1901 (as bella-

tor) by Dyar (1923:72); considered as variety of lutzii (as bellator) by Christophers

(1924:42); synonymized with neivai (as cruzii race neivai) by Dyar (1925a:26); resur-

rected as variety of lutzii (as bellator) by Edwards (1932:46); resynonymized with nei-

vai by Komp (1937:521).

Anopheles (Kerteszia) neivai of Komp (1937:502-503,509,521; 1942:77-78,129-130,163-164);

Senevet and Abonnenc (1938:504-509); Lane (1939:19; 1953:283-284); Levi-Castillo (1945:

118-128); Trapido, Galindo and Carpenter (1955:533-539); Vargas and Martinez Palacios

(1956:132-135,141); Trapido and Galindo (1957:132,133); Stone, Knight and Starcke (1959: -

35-36); Cerqueira (1961:128); Forattini (1961:31-32; 1962:447-449); Aragao (1964:74-75);

Galindo et al. (1966:393); Lee and Sanmartin (1967:778-78 1); Morales-Ayala (1971:139).

Anopheles neivai of Dyar and Knab (1917:40); Howard, Dyar and Knab (1917:986-988); Gab-

aldon and Cova Garcia (1952:183); Galindo and Trapido (1955:546); de Rodaniche, Galindo

and Johnson (1957:683 684); Cova Garcia (1961:58-59,105,135-136,164).

Anopheles (Dendropaedium) neivai of Dyar (1918:146).

Anopheles (Nyssorhynchus) neivai of Root (1922b:391).

Anopheles (Nyssorhynchus) bellator var. neivai of Christophers (1924:42).

Anopheles (Dendropaedium) cruzii race neivai of Dyar (1925a:26-27).

Anopheles (Dendropaedium) hylephilus of Dyar (1918:146).

Anopheles (Nyssorhynchus) bellator var. hylephilus of Christophers (1924:42); Edwards (1932:

46).

Anopheles (Kerteszia) cruzii in part of Dyar (1925b:193; 1928:468-469); Lane (1939: 18-19).

Anopheles (Nyssorhynchus) bellator var. cruzii in part of Edwards (1932:46).

Anopheles cruzii in part of Dyar and Knab (1908:53).

Anopheles bellator in part of Dyar (1923:72).

Anopheles (Anopheles) bellator of Bonne and Bonne-Wepster (1925 :504-507).

Anopheles (Kerteszia) anoplus in part of Komp (1937:515).

FEMALE (figs. 3-5). Wing: 2.93 mm. Proboscis: 2.14 mm. Forefemur: 1.63

mm. Abdomen: about 2.3 mm. Head: Integument brown to dark brown. Proboscis

1.3-1.4 length of forefemur. Palpus with small to moderate white patch at apex of

segment 4 and sometimes at apex of segment 5; scales slightly to conspicuously out-

12 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

standing on proximal portion of segment 3, decumbent on distal portion of segment

3 and on segment 4. Thorax: Integument of scutum light to dark brown. Larger

acrostichal, dorsocentral and middle scutellar bristles predominantly dark. Acrosti-

chal and dorsocentral areas and scutellum without scales. Mep with small upper

patch of scales. Legs: Hindfemur with dark scaled line along lower anterior or

ventral surface. Hindtarsal segment 1 white scaled dorsally from 0.2 to 0.3-0.4,

segments 2-4 with broad white band in apical 0.5-0.7 and segment 5 with broad

white band in apical 0.4-0.6. Wing: Vein C with presectoral, sectoral and subcostal

light spots usually present and moderately large to large; with 1 subcostal light spot.

Vein R243 without subcostal light spot. Vein R445 with light scales absent or re-

stricted to small basal spot. Vein M dark scaled basad of level of furcation in vein

Cu. Vein Cu usually with subbasal light spot. Vein 1A usually entirely dark scaled

basally. Apical fringe spot large, conspicuous, undivided; additional inconspicuous

fringe spots usually at apex of veins M344, Cu,, Cu, and 1A. Abdomen: Tergites

light tan to dark brown. Tergites and sternites II-VII without obvious scales.

MALE. Essentially as in female except for sexual characters.

MALE GENITALIA (fig. 6). Segment VII: Tergite without numerous broad

scales medially. Sidepiece: Internal spine flattened and conspicuously broadened

apically. Claspette: Ventral lobe densely spiculose over entire surface; lateral expan-

sion very narrow, inconspicuous, not curved ventrad posteriorly and not retrorsely

produced anteriorly. Phallosome: Aedeagus without leaflets.

PUPA (fig. 6). Abdomen: about 2.6 mm. Trumpet: 0.36 mm. Paddle: 0.69 mm.

Cephalothorax: Very weakly to moderately pigmented, lighter ventrally. Trumpet:

Yellow to amber. Abdomen: Very weakly to moderately pigmented, lighter poste-

riorly. Hair 1-VII arising near caudal border of segment. Hair 9-IV short and fine,

short, blunt and peglike, or moderately long, pointed and spinelike; 9-V usually

moderately long to long, pointed, spinelike, dissimilar to 9-III and similar, although

shorter and weaker, to 9-VI. Paddle: Obovate; usually more or less obliquely trun-

cate; relatively narrow, index usually 1.6-1.8 (1.5-1.9). Unpigmented or very weakly

pigmented, as light as or lighter than posterior abdominal segments. Marginal spic-

ules along distal portion of external buttress usually predominantly toothlike, short,

few, widely spaced, not extending basad into proximal 0.5 of paddle; outer edge

distad of external buttress with long, wavy, filamentous marginal spicules. Hair 1-P

weakly to moderately developed, moderately long. Hair 2-P absent.

LARVA (fig. 7). Head: 0.62 mm. Anal Saddle: 0.24 mm. Head: Weakly to mod-

erately pigmented, collar darker. Hair 3-C moderately long, moderately developed,

not fusiform; as strong as or slightly stronger than 2-C. Hair 4-C weaker than 2-C.

Hairs 5-7-C moderately long to long. Antenna: Weakly to moderately pigmented,

apex usually slightly darkened. Shaft with a few large, conspicuous spicules on

inner and lower surfaces distad of level of hair 1-A. Hairs 2,3-A moderately long,

subequal in length. Hair 4-A usually single. Thorax: Epidermis usually unpigment-

ed, rarely weakly to moderately pigmented, blackish. Hair 11-P strongly devel-

oped, long. Abdomen: Epidermis usually unpigmented, rarely weakly to moder-

ately pigmented, blackish, on I-VIII. Hair 1-I not palmate, 1-3b or 3f; hair 1-VII

palmate. Palmate hairs small, brushlike in form, the leaflets contracted; leaflets

short, narrow, blunt. Hair 5-II-V usually branched near base only; 5-VII short to

moderately long. Hair 6-VI long, plumose, similar to 6-III-V. Hair 9-IV-VI usually

branched near base only. Spiracular Lobe: Pecten teeth all long, with spinules usu-

ally extending to near apex of both internal and external edges. Hair 1-S usually

Zavortink: Subgenus Kerteszia 13

single (1-3f). Hair 8-S absent. Anal Segment: Hair 4a-X weakly developed, shorter

than anal saddle, usually simple.

SYSTEMATICS. Hairs 8-S on the spiracular lobe of the larva and 2-P on the

paddle of the pupa are absent in this species, thus establishing the ontogenetic

homology of these hairs. The description and drawings of neivai are based on topo-

typic and other material from Panama.

BIONOMICS. The immatures of this species are usually found in leaf axils of

terrestrial and epiphytic bromeliads and rarely in leaf axils of aroids and treeholes.

Females commonly bite humans, particularly in the evening hours. Trapido and

Galindo (1957:132,133) reported that neivai is a predominantly arboreal species

in rain forest in Panama and a predominantly ground level species in deciduous

forest.

A. neivai is the primary vector of human malaria in the coastal area south of

Buenaventura, Colombia (Lee and Sanmartin, 1967:778-781). It has been found

infected with yellow fever virus in Panama (de Rodaniche, Galindo and Johnson,

1957:683,684) and Guaroa virus in Colombia (Lee and Sanmartin, 1967:778-781).

This species was included in a mixed pool found infected with Venezuelan equine

encephalomyelitis, Ilheus and Guaroa viruses in Panama (Galindo et al, 1966:393).

DISTRIBUTION (fig. 1). Southern Mexico to Ecuador and French Guiana; record-

ed also from Bolivia, northern Brazil and Peru.

Material Examined: 485 specimens; 64 males, 23 male genitalia, 118 females,

176 larvae, 104 pupae; 104 individual rearings (64 larval, 22 pupal, 18 incomplete).

BRITISH HONDURAS. Cayo: Guacamallo, 27 Sept 1963, D.J. Lewis, 5 F [BMNH].

COLOMBIA. Choco: Condoto, 19 May and 4 Oct 1913, H.G.F. Spurrell, 2 F [BMNH]. Valle:

Buenaventura, 1938, 5 F [USNM]. Leticia Rio Raposo (ca. 40 km E RFVFS), 9 June 1966, P.A.

Orjuela (COL 266), 1 L; same data (COL 279), 1 IpF (279-12), 2 L; same data (COL 284), 1 L

[UCLA]. Rio Raposo, near mouth (ca. 20 km S Buenaventura), 2 Mar 1965, V.H. Lee (COL 58),

1 IpM (58-17), 1 IpF (58-16), 1 pM (58-107), 1 M gen, 7 L; 31 Mar 1965, V.H. Lee (COL 88),

1 L; 14 Sept 1965, V.H. Lee (COL 92), 1 L; 18 Nov 1965, V.H. Lee (COL 135), 1 IpM (135-10),

1 pF (135-104), 2 L; 4 Apr 1966, P.A. Orjuela (COL 217), 2 IpM (217-13,16); same data (COL

218), 1 L; same data (COL 219), 1 pM (219-10), 1 L; 18 Apr 1966, P.A. Orjuela (COL 227), 1 pF

(227-100); same data (COL 229), 1 Ip (229-12); 20 Aug 1966, P.A. Orjuela (COL 289), 1 IpF

(289-20); same data (COL 293), 1 lpM (293-21), 1 M gen [UCLA]. Rockefeller Foundation Virus

Field Station (Raposo), 23 Feb 1965, V.H. Lee (COL 57), 1 IpF (57-22), 1 pM (57-105), 1 M

gen, 2 L; 4 Nov 1965, V.H. Lee (COL 125), 1 IpM (125-12); same data (COL 126), 1 IpM (126-

10); same data (COL 127), 1 L; 26 Mar 1966, P.A. Orjuela (COL 200), 1 IpF (200-10); 18 Apr

1966, P.A. Orjuela (COL 232), 1 L; same data (COL 236), 1 lp (236-11), 1 L; 28 Apr 1966,

P.A. Orjuela (COL 237), 5 L; 2 May 1966, P.A. Orjuela (COL 240), 1 L; 20 Aug 1966, P.A.

Orjuela (COL 294), 1 L [UCLA]. San Francisco (ca. 20 km E RFVFS), 9 May 1966, P.A.

Orjuela (COL 246), 1 L; 11 May 1966, P.A. Orjuela (COL 253), 1 IpM (253-16), 1 lpF (253-12),

1 P, 2 L; 12 May 1966, P.A. Orjuela (COL 254), 1 IpF (254-14), 1 Ip (254-16), 1 IM (254-11),

1 M gen, 5 L; same data (COL 255), 1 IpM (255-11), 1 L; same data (COL 261), 1 L [UCLA].

COSTA RICA. Cartago: Cervantes, 9 Nov 1962, J.N. Belkin, C.L. Hogue and W.A. Powder

(CR 13), 2 1IpF (13-301,302), 1 L [UCLA]. Estrella, C. Picado, 1 F [USNM]. Orosi, Jan 1938,

H.W. Kumm, 1 M gen [USNM]; 2 F, 1 L [USNM]; (2 km N), 7 Dec 1962, C.L. Hogue and W.A.

Powder (CR 83), 1 IpM (83-604), 2 IpF (83-602,605), 1 Ip (83-603), 1 M gen, 1 L [UCLA].

Heredia: Finca La Selva (Puerto Viejo de Sarapiqui), 6 Aug 1971, A. Berrios Arias (CR 403),

2 1pM (403-10,12), 1 L; 7 Aug 1971, A. Berrios Arias (CR 417), 1 L; 8 Aug 1971, D.W.

Heinemann (CR 422), 1 pF (422-101); same data (CR 426), 1 L;27 Aug 1971, D.W. Heinemann

(CR 500), 1 IpM (500-11), 2 IpF (500-10,12), 4 L; same data (CR 502), 1 Ip (502-10); 28 Aug

1971, D.W. Heinemann (CR 504), 1 L [UCLA]. Puerto Viejo de Sarapiqui, 9 Sept 1965, W.L.

Hjort (CR 227), 1 IpM (227-10) [UCLA]. Limon: Guapiles (10 km SW), 11 Sept 1965, W.L.

14 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

Hjort (CR 230), 2 L; same data (CR 231), 1 L [UCLA]. Department Unknown: Las Concavas,

1 lp (1507) [USNM] . Locality Unknown: C. Picado (1591), 1 M, 1 M gen [USNM]; 1 L [UCLA].

ECUADOR. Bolivar: Balzapamba, 22 July 1938, Hanson, 3 M, 1 M gen, 2 F [UCLA]. Guayas:

Guayaquil, F. Campos Ribadeneira, 1 F (paralectotype of hylephilus) [USNM]. Los Rios: Mont-

alvo, 8 Feb 1966, J.N. Belkin and E. Gerberg (ECU 123), 1 L [UCLA]. Pichincha: Tandapi, 3 F

[USNM]. Locality Unknown: 1 Aug 1938, Hanson, 1 M; 15 Aug 1938, 1 M, 1 M gen, 3 F; 15

Sept 1938, 1 F;5 F [UCLA].

FRENCH GUIANA. Inini: Saut Tigre, 21 Mar 1967, R.X. Schick and Chatanay (FG 172),

1 pM (172-100), 1 M gen [UCLA].

GUATEMALA. Guatemala: Guatemala City, 1 wing [UCLA].

GUYANA. Essequibo: Mazaruni, W.H.W. Komp (KO 121A-8), 2 F [UCLA]. Potaro, May

1910, L.D. Cleare, 1 F [BMNH].

? HONDURAS. Locality Unknown: P.A. Woke (893), 2 M, 1 M gen [UCLA].

MEXICO. Chiapas: Naranjo, Palenque, 25 Mar 1958, A. Diaz Najera, 4 L [ISET]. Santa Rosa,

Independencia, Comitan de Dominguez, 14-22 Nov 1952, J. Williams, 4 F [ISET].

NICARAGUA. Zelaya: Bluefields (1-3 km W Cemetery), 23 Nov 1971, D. and K. Schroeder

(NIC 82), 1 F; same data (NIC 83), 1 IpM (83-20), 1 M gen; 24 Nov 1971, D. Schroeder (NIC

99), 1 IpM (99-11), 1 IpF (99-10), 1 Ip (99-12), 1 M gen, 1 L; (4 km S), 27 Nov 1971, D. and

K. Schroeder (NIC 121), 1 IpF (121-10) [UCLA].

PANAMA. Bocas del Toro: Isla Colon, Big Creek, 11 Apr 1964 (PA 656), 1 L [UCLA].

Kaysan (Chiriquicito), 18 Apr 1963 (PA 229), 1 L; same data (PA 230), 1 L [UCLA]. Wenan

(Almirante), 30 Apr 1963 (PA 293), 1 lp (293-106) [UCLA]. Canal Zone: Cane, 12 Jan 1914,

1 F [USNM]. Fort Davis, 23 Oct-20 Nov 1951, S.J. Carpenter, 7 F; 10 Jan 1951, 4 F [UCLA].

Fort Randolph, 14 July 1924, C.H. Bath, 3 F; 30 Nov 1925, C.H. Bath, 3 F; (fire tower near),

14 July 1924, C.H. Bath, 1 F [USNM]. Gatun, Feb 1917, L.H. Dunn (W-38), 1 F (lectotype of

hylephilus); 28 Oct 1919, A. Tully, 1 F [USNM]. Majagual, 23 Dec 1922, J.B. Shropshire, 1 M

[USNM]. Margarita, 3 Feb 1923, J.B. Shropshire, 1 M [USNM]. Mojinga Swamp, 2 Jan 1937,

W.H.W. Komp, 2 M gen [USNM]. Peluca, 1 M gen [USNM]. Locality not specified, D.P. Curry,

6 M, 5 F [USNM]; 1933, D.P. Curry, 1 M [BMNH]. Colon: Caldera Island, Portobelo Bay,

20 Jan 1908, A.H. Jennings (156.3), 1 F with slide of wing (553) (paralectotype of neivai

no. 20440) [USNM]. Portobelo, Aug 1923, H.G. Dyar and R.C. Shannon, 2 M, 1 F [USNM];

4 Dec 1963 (PA 579), 1 IpF (579-130), 1 L; 9 Dec 1963 (PA 599), 2 L [UCLA]. Darien:

Cerro Mali (summit), 26 May 1963 (PA 363), 1 lpM (363-104), 4 lpF (363-105,117,124,127),

2 pF (363-115,116), 1 P, 3 L; (3 km W summit), 23 May 1963 (PA 357), 1 Ip (357-112), 2 L;

same data (PA 358), 1 lpM (358-101); (6 km W summit), 24 May 1963 (PA 359), 4 IpM

(359-121,123,125,131), 4 IpF (359-117,124,126,127), 4 lp (359-107,122,130,135), 2 pM (359-

110,134), 1 pF (359-109), 1 M, 1 M gen, 10 L; 1 June 1963 (PA 371), 5 IpF (371-106,108,118,

122,127), 2 lp (371-109,130), 1 pM (371-101), 1 IF (371-105), 9 L; 3 June 1963 (PA 373),

2 IpM (373-106,108), 1 IpF (373-105) [UCLA]. La Laguna, 10 July 1963 (PA 451), 1 pF

(451-107) [UCLA]. Morti Hydrographic Station, 1 Dec 1966, O.G.W. Berlin and R. Hinds (PA

964), 1 pM (964-105), 1 M gen; same data (PA 966), 1 lpM (966-20); 2 Dec 1966, O.G.W. Berlin

and R. Hinds (PA 971), 1 IpF (971-30); 4 Dec 1966, O.G.W. Berlin and Linards (PA 980), 1 lpF

(980-20), 1 pF (980-110); 7 Dec 1966, O.G.W. Berlin and R. Hinds (PA 984), 1 pM (984-101);

same data (PA 986), 1 lpM (986-10); same data (PA 990), 2 IpF (990-11,12), 1 lp (990-10),

2 pM (990-101,102), 1 pF (990-100), 1 M gen [UCLA]. Tacarcuna Mts. (1525 m), 25 May-

7 June 1963 (PA 930), 1 IpM (930-104) [UCLA]. Tacarcuna River Valley, 27 June 1963 (PA

430), 1 IpM (430-104); 5 July 1963 (PA 435), 1 pM (435-105), 1 pF (435-106), 1 M gen

[UCLA]. Panama: Candelaria Hydrographic Station, 7,8 May 1966, A. Quinonez (PA 942), 2 F

[UCLA]. La Zumbadora, 8 Feb 1963 (PA 78), 1 IpM (78-102), 1 IpF (78-101), 1 M gen, 1 L

[UCLA]. Pacora, 28 Sept 1941, 1 F [USNM]. Locality Unknown: 1 M, 1 L [UCLA], 1 M gen

[USNM].

SURINAM. Suriname: Paramaribo, J. Bonne-Wepster, 1 M, 1 M gen, 1 F [USNM].

VENEZUELA. Delta Amacuro: Manoa Woods, Orinoco River, 10 Jan 1910, F.L. de Verteuil,

1 F (paralectotype of hylephilus) [USNM].

LOCALITY UNKNOWN. 2 L (11146-x, -y) [USNM].

Zavortink: Subgenus Kerteszia 15

Additional Records from the Literature

BOLIVIA (Stone, Knight and Starcke, 1959:35).

BRAZIL. Amapa (Forattini, 1961:31-32). Amazonas (Cerqueira, 1961:128).

COSTA RICA. Alajuela (Galindo and Trapido, 1955:546). Puntarenas (Trapido, Galindo and

Carpenter, 1955:533).

ECUADOR. EI! Oro, Esmeraldas, Manabi (Levi-Castillo, 1945:127).

EL SALVADOR (Lane, 1953:284; Stone, Knight and Starcke, 1959:35).

MEXICO. Veracruz (Howard, Dyar and Knab, 1917:988).

PANAMA. Chiriqui (Gabaldon and Cova Garcia, 1952:183). Veraguas (Trapido, Galindo and

Carpenter, 1955:533).

PERU (Stone, Knight and Starcke, 1959:35). Loreto (Morales-Ayala, 1971:139).

VENEZUELA. Amazonas, Bolivar, Tachira (Cova Garcia, 1961:164).

2. Anopheles (Kerteszia) pholidotus Zavortink, sp. n.

Figs. 1,3,4,8,9

TYPES: Holotype male (PA 173-110) with slides of associated larval and pupal skins and

genitalia, La Zorra, Bocas del Toro, Panama, 1340 m, larva from leaf axil of terrestrial brome-

liad, 5 Apr 1963, A. Quinonez [USNM]. Allotype female, Caldera-Chiriqui Trail, Bocas del Toro,

Panama, 1000 m, biting human in upper canopy of deep forest, 31 Oct 1955, P. Orguela [USNM].

Paratypes: 1 |\pM (PA 173-104) with slide of genitalia, 1 lp (PA 173-111), 7 L, same data as

holotype (PA 173); 1 L, same data as holotype except collected 6 Apr 1963 in leaf axil of

epiphytic bromeliad (PA 176) [UCLA] ; 1 F, same data as allotype [USNM].

Anopheles (Kerteszia) boliviensis of Anduze (1943:191, ?); Levi-Castillo (1949:16, in part, ?);

Trapido and Galindo (1957:128); Stone, Knight and Starcke (1959:35, in part); Aragao (1964:

76-77, in part, ?).

Anopheles boliviensis of Cova Garcia (1961:57-58,104,134-135,162, in part, ?).

FEMALE (figs. 3,4). Wing: 3.40 mm. Proboscis: 1.90 mm. Forefemur: 1.74 mm.

Abdomen: about 2.1 mm. Head: Integument dark brown. Proboscis 1.1 length of

forefemur. Palpus with moderate white patch at apex of segments 3-5 and smaller

white patch at apex of segment 2; scales entirely or predominantly decumbent on

segments 3 and 4, sometimes slightly outstanding at base of segment 3. Thorax:

Integument of scutum dark brown. Larger acrostichal and dorsocentral bristles pale

and dark; larger middle scutellar bristles dark. Anterior 0.3-0.4 of acrostichal and

dorsocentral areas with a few white scales and middle of scutellum with a few dark

scales. Mep with large, curved patch of scales extending ventrad from bristles to

below middle of segment. Legs: Hindfemur with dark scaled line along lower ante-

rior or ventral surface. Hindtarsal segment 1 without white scaling dorsally, segment

2 with narrow white band in apical 0.1-0.2, segment 3 with broad white band in

apical 0.6, segment 4 with broad white band in apical 0.8 and segment 5 with broad

white band in apical 0.4. Wing: Vein C with large presectoral, sectoral and subcostal

light spots; with 1 subcostal spot. Vein R43 with subcostal light spot. Vein Rais

with or without light scales in separate small basal spot and short submedian line

extending from 0.2 to 0.3-0.4. Vein M dark scaled basad of level of furcation in

vein Cu. Vein Cu with or without subbasal light spot. Vein 1A entirely dark scaled

basally. Apical fringe spot apparently moderate, conspicuous, undivided; additional

inconspicuous fringe spots usually at apex of veins Mi42, M344, Cu, and Cu,.

Abdomen: Tergites light tan to brown. Tergites II-VII with numerous dark brown

16 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

to black scales; scales of proximal tergites predominantly narrow to moderately

broad; scales of distal tergites predominantly outstanding and aggregated into trans-

verse apical bands. Sternites II-VII with white scales medially; more distal sternites

with additional dark scales apically.

MALE. Essentially as in female except for sexual characters.

MALE GENITALIA (fig. 8). Segment VIII: Tergite with numerous broad scales

medially. Sidepiece: Internal spine flattened and slightly broadened apically. Clasp-

ette: Ventral lobe moderately spiculose except laterally; lateral expansion broad,

conspicuous, rounded, sometimes curved ventrad posteriorly, not retrorsely pro-

duced anteriorly. Phallosome: Aedeagus without leaflets.

PUPA (fig. 8). Abdomen: about 2.8 mm. Trumpet: 0.41 mm. Paddle: 0.79 mm.

Cephalothorax: Very weakly to moderately pigmented, lighter ventrally. Trumpet:

Yellow to amber. Abdomen: Very weakly to moderately pigmented, lighter poste-

riorly. Hair 1-VII arising cephalad of caudal border of segment. Hair 9-[V, V mod-

erately long, pointed, spinelike, slightly longer and stronger on V, dissimilar to 9-III

and 9-VI. Paddle: Obovate; more or less obliquely truncate; relatively narrow, index

1.8. Unpigmented or very weakly pigmented, as light as or lighter than posterior

abdominal segments. Marginal spicules along distal portion of external buttress fila-

mentous, moderately long, numerous, closely spaced, extending basad into proximal

‘0.5 of paddle; outer edge distad of external buttress with long, wavy, filamentous

marginal spicules. Hair 1-P strongly developed, short. Hair 2-P present.

LARVA (fig. 9). Head: 0.66 mm. Anal Saddle: 0.26 mm. Head: Weakly pig-

mented, collar darker. Hair 3-C moderately long, moderately developed, not fusi-

form; slightly to considerably stronger than 2-C. Hair 4-C weaker than 2-C. Hairs

5-7-C moderately long to long. Antenna: Weakly pigmented, apex not darkened.

Shaft with relatively small, inconspicuous spicules. Hair 2-A moderately long, 3-A

long. Hair 4-A usually single. Thorax: Epidermis unpigmented. Hair 11-P strongly

developed, long. Abdomen: Epidermis unpigmented. Hair 1-I, VII not palmate, 1-4b

or 4f. Palmate hairs small, brushlike in form, the leaflets contracted; leaflets short,

narrow to moderately broad, pointed or blunt. Hair 5-II-V usually branched near

base only; 5-VII short to moderately long. Hair 6-VI moderately long, with a few

long branches in basal portion, dissimilar to 6-III-V. Hair 9-IV-VI usually single.

Spiracular Lobe: Pecten teeth all long, with spinules usually extending to-near apex

_of both internal and external edges. Hair 1-S single. Hair 8-S present. Anal Segment:

Hair 4a-X moderately developed, usually longer than anal saddle, distinctly pecti-

nate or plumose.

SYSTEMATICS. Anopheles pholidotus and the closely related lepidotus possess

abdominal scales in the adult, and, as a consequence, they have been confused with

boliviensis in the past. These species differ from boliviensis, however, in several

striking features of the adult, as pointed out in the key and descriptions. The spec-

imens of pholidotus from Venezuela differ from the Panamanian ones described

above in the adult by the possession of light and dark scales on the middle of the

-scutellum and the small, inconspicuous, divided apical fringe spot, in the pupa by

the more strongly pigmented cephalothorax, trumpet and abdomen and in the lar-

va by the more strongly pigmented head capsule and antennal apex.

BIONOMICS. The immatures of this species have been collected in leaf axils of

terrestrial and epiphytic bromeliads and females have been captured biting humans

in the upper canopy of deep forest.

DISTRIBUTION (fig. 1). Mountains of western Panama and western Venezuela.

Material Examined: 25 specimens; 3 males, 3 male genitalia, 3 females, 12 larvae,

4 pupae; 4 individual rearings (3 larval, 1 incomplete).

Zavortink: Subgenus Kerteszia 17

PANAMA. Bocas del Toro: Caldera-Chiriqui Trail and La Zorra, type series, cited above.

VENEZUELA. Barinas: Altamira, 1 lpM, 1 M gen (1981-2) [DERM]. Merida: Merida, 10

June 1938, P.J. Anduze, 1 F [USNM].

3. Anopheles (Kerteszia) lepidotus Zavortink, sp. n.

Figs. 1,3,4,16

TYPES: Holotype male (B) with slides of associated larval skin and genitalia, Restrepo, Meta,

Colombia, larva from leaf axil of bromeliad, Dec 1935, E. Osorno-Mesa [USNM]. Allotype

female, Cuchillo, E of Villavicencio, Meta, Colombia, 16 July 1943 [USNM]. Paratypes: 1 IM

(A) with slide of genitalia, same data as holotype [USNM]; 1 F, Buena Vista (near), Meta,

Colombia, 4 June 1942, W.H.W. Komp [USNM]; 15 F, Restrepo, Meta, Colombia, with the

following additional information—Aug 1935, W.H.W. Komp, 4 F [USNM], 2 F [BMNH]; 1935,

W.H.W. Komp, 3 F [USNM]; 20 Nov 1936, W.H.W. Komp, 1 F [USNM]; W.H.W. Komp (KO

121A-10), 2 F [UCLA] ;3 F [USNM].

Anopheles (Kerteszia) boliviensis of Komp (1936:68, in part; 1937:503-504); Komp and Osorno-

Mesa (1936:415-419); Lane (1939:18, in part; 1953:279-281); Roca-Garcia (1944: 160-169,

in part, ?); Levi-Castillo (1945:128-139,150, in part, ?; 1949:16, in part); Rachou (1958:

146, in part, ?); Stone, Knight and Starcke (1959:35, in part); Forattini (1962:461-462);

Aragao (1964:76-77, in part); Morales-Ayala (1971:139, in part, ?).

Anopheles boliviensis of Bates (1943:23, in part; 1944:168,169, in part; 1945:24, in part);

Gabaldon and Cova Garcia (1952:189, in part).

FEMALE (figs. 3,4). Wing: 3.55 mm. Proboscis: 2.06 mm. Forefemur: 1.86 mm.

Abdomen: about 2.2 mm. Head: Integument dark reddish brown to brown. Probos-

cis 1.1 length of forefemur. Palpus with moderate white patch at apex of segments

2-5; scales entirely or predominantly decumbent on segments 3 and 4, sometimes

slightly outstanding at base of segment 3. Thorax: Integument of scutum dark red-

dish brown. Larger acrostichal and dorsocentral bristles pale and dark; larger middle

scutellar bristles dark. Anterior 0.3-0.4 of acrostichal and dorsocentral areas with a

few white scales and middle of scutellum with a few white and dark scales. Mep

with large, curved patch of scales extending ventrad from bristles to below middle

of segment. Legs: Hindfemur with dark scaled line along lower anterior or ventral

surface. Hindtarsal segment 1 without white scaling dorsally, segment 2 with narrow

white band in apical 0.1-0.2, segment 3 with broad white band in apical 0.4-0.7,

segment 4 with broad white band in apical 0.7-0.8 and segment 5 with broad white

band in apical 0.4-0.8. Wing: Vein C with presectoral, sectoral and subcostal light

spots usually present and moderately large to large; with 1 subcostal spot. Vein

R43 usually with subcostal light spot. Vein R44; usually without light basal spot

or submedian line. Vein M dark scaled basad of level of furcation in vein Cu. Veins

Cu and 1A dark scaled basally. Apical fringe spot small, inconspicuous, sometimes

divided into 2 separate spots; additional inconspicuous fringe spots usually at apex

of veins M,42, M344, Cu, and Cu,. Abdomen: Tergites brown. Tergites II-VII with

numerous dark brown to black scales and sometimes a few white scales; scales of

proximal tergites predominantly moderately broad to broad; scales of distal tergites

predominantly outstanding and aggregated into transverse apical bands. Sternites

II-VII with white scales medially; more distal sternites with additional dark scales

apically.

MALE. Essentially as in female except for sexual characters.

18 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

MALE GENITALIA (fig. 16). Segment VII: Tergite with numerous broad scales

medially. Sidepiece: Internal spine apparently not flattened at apex. Claspette:

Ventral lobe moderately spiculose except near lateral margin; lateral expansion

broad, conspicuous, shallowly and broadly emarginate, not curved ventrad poste-

riorly, produced into blunt retrorse lobe anteriorly. Phallosome: Aedeagus without

leaflets.

PUPA. Unknown.

LARVA (not illustrated). Imperfectly known. In general similar to pholidotus,

but with the following differences. Head: Hair 3-C short, strongly developed, fusi-

form; much stronger than 2-C. Thorax: Hair 11-P weakly developed, short. Abdo-

men: Palmate hairs moderate, not particularly brushlike, the leaflets spreading;

leaflets moderately long, moderately broad, pointed. Hair 5-II-V distinctly plumose;

~ 5-VII short. Hair 9-I[V-VI usually branched near base only. Spiracular Lobe: Pecten

teeth with spinules possibly restricted to basal portion of external edge.

SYSTEMATICS. Although lepidotus closely resembles pholidotus or is possibly

even indistinguishable from it in the adult and male genitalia, | am recognizing it as

a distinct species on the basis of the larval differences indicated in the key and de-

scription. The holotype and paratype males of /epidotus with associated larval skins

and genitalia are the specimens described as boliviensis by Komp and Osorno-Mesa

(1936).

BIONOMICS. The immatures of lepidotus are found in bromeliad leaf axils. Of

the 204 females of Kerteszia with scales on the abdomen from Colombia in the

UCLA and USNM collections, 174 or 85% are this species; as a consequence, I

believe that most information on biology and medical importance of “‘boliviensis”’

in the Department of Meta, Colombia, found in the literature pertains to lepidotus

and, for that reason, it is summarized here. Komp (1936:68) reported that females

of “boliviensis’’ were common in the jungle where they frequently bit humans

during the day and were taken in large numbers on horses in the evening. Bates

(1944:168,169; 1945:24) reported that females were most common from April to

October, were most active in the late afternoon and early evening and were more

frequently encountered in the upper levels of forest (79% of 314) than at ground

level.

In Colombia “boliviensis’’ has been found infected with the viruses Anopheles A

and Anopheles B (Roca-Garcia, 1944:160-169) and found carrying the eggs of

Dermatobia hominis (Linnaeus) (Bates, 1943:23). In Ecuador it was suspected to

be a vector of human malaria by Levi-Castello (1945: 137).

DISTRIBUTION (fig. 1). Eastern slope of the Andes in Colombia and Bolivia;

possibly also Brazil, Guyanas, Paraguay and Venezuela. :

Material Examined: 181 specimens; 2 males, 2 male genitalia, 175 females, 2

larvae; 2 incomplete individual rearings.

BOLIVIA. Cochabamba: El Palmar, Chapare, 2 May 1944, Torres Munoz, 1 F [USNM].

COLOMBIA. Meta: Buena Vista, Cuchillo and Restrepo, type series, see above. Locality Un-

known: J.A. Kerr, 157 F [UCLA].

Additional Records from the Literature

Some of the records of ‘‘boliviensis’’ listed under that species may pertain to lepidotus.

4. Anopheles (Kerteszia) bambusicolus Komp

Figs. 1,3,4,10,11

Zavortink: Subgenus Kerteszia 19

1937. Anopheles (Kerteszia) bambusicolus Komp, 1937:515-518. TYPE: Lectotype female,

La Union, Meta, Colombia, reared from larva collected in unbroken bamboo internode,

9 Sept 1935, J. Boshell [USNM, 53075; designation by Stone and Knight, 1956:276].

1967. Anopheles (Kerteszia) bambusicola Stone, 1967:200. Junior objective synonym (unjus-

tified emendation) of bambusicolus Komp, 1937.

Anopheles (Kerteszia) bambusicolus of Lane (1939:17-18; 1953:281-282); Coutinho (1946:150-

152); Levi-Castillo (1949:16); Rachou and Ferreira Neto (1950:303-305); Komp (1956:40-

41); Bejarano (1957:317); Stone, Knight and Starcke (1959:35); Forattini (1962:462); Garcia

and Ronderos (1962:149-150); Aragao (1964:78-80).

Anopheles (Kerteszia) bambusicola of Morales-Ayala (1971:139).

Anopheles bambusicolus of Gabaldon and Cova Garcia (1952:188-189).

Anopheles bellator in part of Komp (1936:68).

FEMALE (figs. 3,4). Wing: 3.43 mm. Proboscis: 2.18 mm. Forefemur: 1.73 mm.

Abdomen: about 2.2 mm. Head: Integument dark reddish brown to dark brown.

Proboscis 1.2-1.3 length of forefemur. Palpus with small white patch at apex of seg-

ment 4; scales conspicuously outstanding on segments 3 and 4. Thorax: Integument

of scutum reddish brown to brown. Larger acrostichal, dorsocentral and middle scu-

tellar bristles predominantly dark. Acrostichal and dorsocentral areas and scutellum

without scales. Mep with small upper patch of scales. Legs: Hindfemur with dark

scaled line along lower anterior or ventral surface. Hindtarsal segment 1 white scaled

dorsally from 0.1 to 0.3-0.5, segments 2 and 3 with broad white band in apical 0.7-

0.8, segment 4 with broad white band in apical 0.5-0.6, and segment 5 entirely

white scaled. Wing: Vein C with presectoral, sectoral and subcostal light spots usu-

ally absent, small when present; with 0, 1 or 2 subcostal spots. Vein R243 without

subcostal light spot. Vein R45 with light scales restricted to small basal spot. Vein

M dark scaled basad of level of furcation in vein Cu. Vein Cu with subbasal light

spot. Vein 1A dark scaled basally. Apical fringe spot not developed; inconspicuous

fringe spots at apex of veins M3,, and Cu,. Abdomen: Tergites brown to dark

brown. Tergites and sternites II-VII without obvious scales.

MALE. Essentially as in female except for sexual characters.

MALE GENITALIA (fig. 10). Segment VII: Tergite without numerous broad

scales medially. Sidepiece: Internal spine flattened and slightly broadened apically.

Claspette: Ventral lobe glabrous except for 4-10 strong spicules and a few weak

spicules along mesal margin; lateral expansion broad, prominent, rounded, not cur-

ved ventrad posteriorly and not retrorsely produced anteriorly. Phallosome: Ae-

deagus without leaflets.

PUPA (fig. 10). Abdomen: about 2.8 mm. Trumpet: 0.37 mm. Paddle: 0.71 mm.

Cephalothorax: Moderately pigmented, lighter ventrally. Trumpet: Possibly golden

brown. Abdomen: Moderately pigmented, lighter posteriorly. Hair 1-VII arising on

caudal border of segment. Hair 9-IV, V short, blunt, peglike, similar to 9-III and

dissimilar to 9-VI. Paddle: Elliptic; not obliquely truncate; relatively broad, index

1.4. Unpigmented or very weakly pigmented, lighter than posterior abdominal seg-

ments. Marginal spicules along distal portion of external buttress toothlike, short,

few, widely spaced, not extending basad into proximal 0.5 of paddle; outer edge

distad of external buttress with short marginal spicules. Hair 1-P strongly developed,

short. Hair 2-P present.

LARVA (fig. 11). Head: 0.63 mm. Anal Saddle: 0.29 mm. Head: Weakly (?) to

strongly pigmented, collar darker, ocular area lighter. Hair 3-C moderately long,

weakly to moderately developed, not fusiform; weaker or stronger than 2-C. Hair

20 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

4-C weaker than 2-C. Hairs 5-7-C short to moderately long. Antenna: Weakly (?)

to strongly pigmented, distal portion darkened. Shaft with relatively small, incon-

spicuous spicules. Hairs 2,3-A short to moderately long, subequal in length. Hair

4-A usually single. Thorax: Integument very dark according to Komp (1937:518).

Hair 11-P weakly developed, short. Abdomen: Integument very dark according to

Komp. Hair 1-I, VII with numerous simple branches, subpalmate with numerous

slightly flattened branches, or palmate. Palmate hairs large, usually not at all brush-

like, the leaflets widely spread; leaflets long, moderately broad to broad, lanceolate.

Hair 5-II-V single, plumose, or branched near base only; 5-VII short. Hair 6-VI short,

with a few long branches near base or short branches on main shaft distally, dis-

similar to 6-III-V. Hair 9-IV-VI 1-4b, the branches usually arising near base only.

Spiracular Lobe: Pecten teeth all long or a few shortened, with spinules varied, ex-

tending to apex or restricted to basal portion, occurring on both internal and ex-

ternal edges or restricted to external edge. Hair 1-S single or 2,3f. Hair 8-S present.

Anal Segment: Hair 4a-X moderately to strongly developed, much longer than anal

saddle, pectinate or plumose.

SYSTEMATICS. One of the 3 females of bambusicolus examined has no presec-

toral, sectoral or subcostal light spots on the wing, one has a moderate subcostal

spot, and one has 2 small subcostal spots; the only male examined has a small sec-

toral spot and 2 large subcostal spots.

BIONOMICS. The immatures of bambusicolus are found in unbroken bamboo

internodes; Komp (1956:40) reported finding them 35 feet above the ground. One

female examined was taken in a biting-landing collection in association with lepido-

tus.

DISTRIBUTION (fig. 1). Eastern slope of Andes in Colombia; recorded also

from Argentina, Bolivia, Brazil, Ecuador, Guyanas, Peru and Venezuela.

Material Examined: 15 specimens; 1 male, 3 male genitalia, 3 females, 7 larvae,

1 pupa; 1 larval individual rearing.

COLOMBIA. Meta: La Union, 9 Sept 1935, J. Boshell, 1 F, 4 L (paralectotypes no. 53075)

[USNM]. Villavicencio, 1944, M. Bates, 1 lpF (72-1), 2 L [USNM]. Villavicencio (115), 1 M,

1 M gen [UCLA]. Locality not specified, 2 M gen (464.4, 467.2) [UCLA]. Locality Unknown:

J.A. Kerr, 1 F [UCLA].

Additional Records from the Literature

ARGENTINA. Misiones (Bejarano, 1957:317; Garcia and Ronderos, 1962:149-150).

BOLIVIA (Levi-Castillo, 1949:16).

BRAZIL. Parana (Coutinho, 1946:150-152). Santa Catarina (Rachou and Ferreira Neto, 1950:

303-305).

ECUADOR. Napo-Pastaza, Santiago-Zamora (Gabaldon and Cova Garcia, 1952:188-189).

GUIANAS (Levi-Castillo, 1949:16).

PERU. Amazonas, Cuzco, Madre de Dios, San Martin (Gabaldon and Cova Garcia, 1952:188-

189). Loreto (Morales-Ayala, 1971:139).

VENEZUELA (Levi-Castillo, 1949:16).

5. Anopheles (Kerteszia) homunculus Komp

Figs. 2-4,12,13

1937. Anopheles (Kerteszia) homunculus Komp, 1937:509-513. TYPE: Lectotype male (no. 3)

with slides of associated larval skin and genitalia, Restrepo, Meta, Colombia, larva collect-

ed in bromeliad leaf axil, 9 Sept 1935, W.H.W. Komp [USNM, 53073; designation by

Stone and Knight, 1956:278].

Zavortink: Subgenus Kerteszia ZA

1937. Anopheles (Kerteszia) anoplus Komp, 1937:514-515. TYPE: Holotype male with slides

of associated larval skin and genitalia, Restrepo, Meta, Colombia, larva collected in bro-

meliad leaf axil, Dec 1936, E. Osorno-Mesa [USNM, 53074]. Synonymy with homun-

culus by Lane (1953:287).

Anopheles (Kerteszia) homunculus of Lane (1939:19; 1953:287-288); Downs and Pittendrigh

(1946:49,55,56); Pinotti (1948:694-695); Pittendrigh (1948:59; 1950a:457-468; 1950b:43-

63; 1950c:64-77); Levi-Castillo (1949:16-17); Lima (1952:401-404); Veloso, Moura and Klein

(1956:519); Martins (1958:429-430); Rachou (1958:149); Stone, Knight and Starcke (1959:

35); Forattini (1962:441); Aragao (1964:86-87); Ferreira (1964:335,338,340); Morales-Ayala

(1971:139).

Anopheles humunculus of Senior-White and Lewis (1951:151); Gabaldon and Cova Garcia (1952:

191); Cova Garcia (1961:61-62,107-108,133-134,162,164); Forattini, Rabello and Cotrim

(1970:12).

Anopheles (Kerteszia) anoplus of Komp (1937:514-515, in part); Lane (1939:17).

Anopheles bellator in part of Komp (1936:68).

FEMALE (figs. 3,4). Wing: 2.80 mm. Proboscis: 2.15 mm. Forefemur: 1.54 mm.

Abdomen: about 2.1 mm. Head: Integument dark reddish brown to dark brown.

Proboscis 1.3-1.4 length of forefemur. Palpus with small to moderate white patch

at apex of segments 3-5, segments 4 and 5 or only segment 4, the patch largest on

segment 4 when present on more than | segment; scales slightly to moderately out-

standing on segment 3 and slightly outstanding to decumbent on segment 4. Tho-

rax: Integument of scutum reddish brown. Larger acrostichal, dorsocentral and mid-

dle scutellar bristles predominantly dark. Acrostichal and dorsocentral areas and

scutellum without scales. Mep with small upper and larger middle patches of scales.

Legs: Hindfemur without dark scaled line or with weak, broken dark scaled line

along lower anterior or ventral surface. Hindtarsal segment 1 white scaled dorsally

from 0.2 to 0.3-0.5, segments 2-4 with broad white band in apical 0.5-0.7 and seg-

ment 5 with broad white band in apical 0.4-0.7. Wing: Vein C with moderately

large to large presectoral, sectoral and subcostal light spots; with 1 subcostal spot.

Vein R243 without subcostal light spot. Vein R445 with light scales in small basal

spot and moderately long to long submedian line extending from 0.2-0.4 to 0.5-0.8.

Vein M dark scaled basad of level of furcation in vein Cu. Vein Cu with subbasal

light spot. Vein 1A dark scaled. basally. Apical fringe spot large, conspicuous, un-

divided; additional inconspicuous fringe spots at apex of veins M344, Cu,, Cu,

and 1A. Abdomen: Tergites tan to dark brown or blackish. Tergites and sternites

II-VII without obvious scales.

MALE. Essentially as in female except for sexual characters.

MALE GENITALIA (fig. 12). Segment VIII: Tergite without numerous broad

scales medially. Sidepiece: Internal spine flattened and slightly broadened apically.

Claspette: Ventral lobe densely spiculose except near lateral edge; lateral expansion

broad, conspicuous, more or less sinuous-margined, curved ventrad posteriorly and

produced into sharp retrorse point anteriorly. Phallosome: Aedeagus with or with-

out leaflets; if present, leaflets usually weak, short or long.

PUPA (fig. 12). Abdomen: about 2.3 mm. Trumpet: 0.31 mm. Paddle: 0.66 mm.

Cephalothorax: Very weakly to moderately pigmented, lighter ventrally. Trumpet:

Yellow to amber. Abdomen: Very weakly to moderately pigmented, lighter poste-

riorly. Hair 1-VII arising on caudal border of segment. Hair 9-[V moderately long

to long, pointed, spinelike; hair 9-V long, pointed, spinelike, dissimilar to 9-III and

similar, but sometimes weaker and shorter, to 9-VI. Paddle: Obovate; not obliquely

ZZ Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

truncate; relatively narrow, index 1.6-1.9. Unpigmented to very weakly pigmented,

lighter than posterior abdominal segments. Marginal spicules along distal portion

of external buttress filamentous, moderately long, numerous, closely spaced, not

extending into proximal 0.5 of paddle; outer edge distad of external buttress with-

out spicules. Hair 1-P strongly developed, short. Hair 2-P present.

LARVA (fig. 13). Head: 0.54 mm. Anal Saddle: 0.20 mm. Head: Weakly to

moderately pigmented, collar darker. Hair 3-C moderately long, moderately to

strongly developed, not fusiform; much stronger than 2-C. Hair 4-C as strong as

or stronger than 2-C. Hairs 5-7-C moderately long to long. Antenna: Weakly to

moderately pigmented, apex not darkened. Shaft with relatively small, inconspic-

uous spicules. Hairs 2,3-A moderately long to long, subequal in length. Hair 4-A

usually single. Thorax: Epidermis moderately to strongly pigmented, blackish.

Hair 11-P strongly developed, long. Abdomen: Epidermis moderately to strongly

pigmented, blackish, on entire dorsal surface of segments III and VI-VIII and mid-

dorsally on I,II and IV. Hair 1-I, VII palmate. Palmate hairs small to moderate,

usually not particularly brushlike, the leaflets spreading; leaflets moderately long,

moderately broad, pointed. Hair 5-II-V usually branched near base only; 5-VII long.

Hair 6-VI long, plumose, similar to 6-III-V. Hair 9-IV-VI usually double with 1

branch shortened, plumose or pectinate. Spiracular Lobe: Pecten teeth alternating

long and short medially, with spinules usually restricted to basal portion of external

edge. Hair 1-S single. Hair 8-S present. Anal Segment: Hair 4a-X usually weakly de-

veloped, shorter or slightly longer than anal saddle, pectinate or plumose.

SYSTEMATICS. The description and drawings of homunculus are based on speci-

mens from Trinidad. The scanty material of homunculus from Colombia that I have

seen differs from that from Trinidad in the female by the reduced size or absence

of the presectoral, sectoral and subcostal light spots on vein C, the usual absence of

a light fringe spot at the apex of vein Cu,, the usual absence of extensive white

scaling on midtarsal segment 3 and the usually more decumbent scales on the palpus

and in the larva by the finer and less conspicuously barbed hair 3-C and the longer

hair 4a-X. The taxonomic significance of these differences cannot be determined

until much more material is available.

BIONOMICS. The immatures are found in the leaf axils of epiphytic and terres-

trial bromeliads. Adults are attracted to lights and females bite humans. Pittendrigh,

in a series of excellent papers (1948,1950a,b,c,), compared the ecologies of homun-

culus and bellator in Trinidad and found that homunculus characteristically inhabit-

ed areas of higher humidity than the latter species and, as a consequence, it occurred

in the wetter parts of the island and at lower levels in the forest. This same ecolog-

ical relationship between homunculus and bellator holds in southeastern Brazil,

where homunculus inhabits the dense, humid primary forest of the higher, wetter,

inland areas. (Pinotti, 1948:694-695; Veloso, Moura and Klein, 1956:519: Aragao,

1964:86-87).

A. homunculus is an important vector of human malaria locally in southeastern

Brazil (Rachou, 1958:149; Forattini, 1962:554) and in Trinidad (Pittendrigh, 1948:

59; Forattini, 1962:554).

DISTRIBUTION (fig. 2). Eastern slope of Andes in Colombia and Bolivia, Trini-

dad; recorded also from southeastern Brazil, Guianas, Peru, Surinam and Venezuela.

Material Examined: 155 specimens; 9 males, 11 male genitalia, 77 females, 36

Zavortink: Subgenus Kerteszia zd

larvae, 22 pupae; 21 individual rearings (17 larval, 2 pupal, 2 incomplete).

BOLIVIA. Cochabamba: El Palmar, Chapare, 30 Apr 1944, Torres Munoz, 2 F [USNM].

COLOMBIA. Meta: Restrepo, 9 Sept 1935, W.H.W. Komp, 1 F (No. 1), 1 Ip (No. 4) (para-

lectotypes No. 53073) [USNM]. Villavicencio, 9 July 1965, E. Osorno-Mesa et al. (COB-67),

1 lpM (67-30); 1944, M. Bates, 1 P, 5 L; June 1944, 1 F; 1 F [UCLA]. Locality Unknown:

J.A. Kerr, 2 F [UCLA]. :

TRINIDAD. Caroni: Tabaquite, 18 May 1914, J.R. Dickson, 1 F [BMNH]. Nariva: Charuma

Forest, 27 Aug 1964, A. Guerra (TR 644), 1 F [UCLA] ; Oct 1954, T.H.G. Aitken, 3 F [BMNH].

St. Andrew: Caratal Rd., 3 Dec 1964, F. Powdhar (TR 866), 1 F [UCLA]. Coalmine Rd., 24 June

1965, F. Powdhar (TR 1227), 1 pF (1227-100) [UCLA]. Cumaca, 3 Sept 1964, A. Guerra (TR

654), 3 F; 18 Feb 1965, A. Guerra (TR 1010), 5 F [UCLA]. Cumaca Rd., 14 Jan 1965, A.

Guerra (TR 943), 1 F [UCLA]. Cunaripa, 11 June 1964, A. Guerra (TR 470), 1 F; 12 June

1964, A. Guerra (TR 471), 1 F [UCLA]. El Quemado Rd., 3 July 1964, F. Powdhar (TR 540),

1 L [UCLA]. Heights of Oropuche, 5 Apr 1925, F.W. Urich, 1 F [BMNH]. Mara Forest, Valencia,

26 Dec 1930, M.V. Beattie, 1 F [BMNH]. Mt. Becke, 29 Apr 1965, A. Guerra (TR 1132), 2 F;

same data (TR 1136), 1 lpM (1136-20), 1 L; same data (TR 1137), 1 F; same data (TR 1141),

1 F; same data (TR 1143), 3 F [UCLA]. Mt. Harris, 23-31 July 1924, C.L. Withycombe, 2 F

[BMNH]; 16 July 1964, F. Powdhar (TR 564), 2 L [UCLA]. Nestor Village, 12 June 1964,

A. Guerra (TR 484), 1 F [UCLA]. Sangre Grande, 1944, 1 M, 1 M gen; 1946, 3 F [USNM].

Tamana, June 1944, 3 M gen [USNM]. Turure Forest, 24 Dec 1964, F. Powdhar (TR 901),

1 IpF (901-132); 30 Apr 1966, A. Guerra (TR 1508), 3 IpF (1508-20-22); 30 July 1966, F.

Powdhar (TR 1572), 1 IpF (1572-30), 2 L; Sept 1966, F. Guerra (TR 1617), 1 F [UCLA].

Turure Rd., 9 Apr 1966, A. Guerra (TR 1497), 1 lpF (1497-20) [UCLA]. Turure Trace, 2 Apr

1966, A. Guerra (TR 1491), 2 IpF (1491-20,23), 1 lp (1491-21), 2 L [UCLA]. Valencia, 22

Aug 1945, 1 lpM (5-T-44) [USNM]. Valencia District, 1 M gen [USNM]. St. George; Arena

Forest, 10 Aug 1965, A. Guerra (TR 1309), 1 lpM (1309-30), 1 M gen; Dec 1965, TRVL (TR

1441), 1 F [UCLA]. Aripo Valley, 15 Apr 1965, A. Guerra (TR 1112), 1 F [UCLA]. Blanch-

isseuse Rd. (mile post 11), 2 Sept 1945, 4 F [USNM]. Brasso Seco, 2 Apr 1964, A. Guerra

(TR 262), 1 L; same data (TR 264), 1 lpM (264-153), 2 IpF (264-137,200), 1 M gen; 18 Mar

1965, A. Guerra (TR 1046), 1 IpF (1046-11), 1 pF (1046-10), 1 L [UCLA]. Heights of Guanapo,

26 Mar 1964, A. Guerra (TR 259), 2 F; 22 Apr 1965, A. Guerra (TR 1127), 1 F; same data (TR

1129), 1 F [UCLA]. Las Lapas Trace, 3 Apr 1964, A. Guerra (TR 279), 1 L; same data (TR

281), 4 F [UCLA]. La Lujha Rd., 11 Mar 1965, A. Guerra (TR 1036), 3 F [UCLA] . Spring Hill

Estate, 26 Aug 1957, T.H.G. Aitken, 2 M, 2 M gen; 13 May 1965, A. Guerra (TR 1155), 5 F;

5 Mar 1966, A. Guerra (TR 1476), 1 IpM (1476-20), 1 M gen; 5 Aug 1966, T.H.G. Aitken (TR

1574), 1 F [UCLA]. St. Patrick: La Brea, 24 Feb 1914, J.R. Dickson, 1 F, 1 L [BMNH]. Lo-

cality Unknown: 1949, 1 M gen [USNM].

Additional Records from the Literature

BRAZIL. Parana (Forattini, Rabello and Cotrim, 1970:12). Santa Catarina, Sao Paulo (Aragao,

1964:86).

GUIANAS (Levi-Castillo, 1949:16-17).

PERU. Loreto (Morales-Ayala, 1971:139).

? SURINAM (Stone, Knight and Starcke, 1959:35).

VENEZUELA. Bolivar, Monagas, Portuguesa, Tachira, Trujillo (Cova Garcia, 1962:164). Sucre

(Gabaldon and Cova Garcia, 1952:191).

6. Anopheles (Kerteszia) cruzii Dyar & Knab

Figs. 2-4, 16

1901. Anopheles lutzii Theobald, 1901:177-178. TYPE: Lectotype female, Rio de Janeiro

[Guanabara], Brazil, 4 July 1899, A. Lutz [BMNH; designation by Belkin, 1968:10].

Junior primary homonym of lutzii Cruz, 1901.

24 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

1908. Anopheles cruzii Dyar and Knab, 1908:53 (30 Oct). Nomen novum for lutzii Theobald,

1901.

1908. Myzorhynchella adolphoi Neiva, 1908:457 (30 Nov). Nomen novum for lutzii Theobald,

1901.

1950. Anopheles (Kerteszia) montemor Correa, 1950:53-54. TYPE: Holotype male (102) with

slides of associated larval and pupal skins (15, G8D1) and genitalia (15, G8D1), Caragua-

tatuba, Sao Paulo, Brazil, 1946, G.R. Ramalho and J. Germano [FH; see Belkin, Schick

and Heinemann, 1971:7]. Synonymy with lutzii Theobald, 1901 (as cruzii) by Barretto

and Coutinho (1951:177-179).

Anopheles (Kerteszia) cruzii of Dyar (1928:468-469, in part); Lane (1939:18-19, in part); Correa

and Cerqueira (1944:111,114,115); Levi-Castillo (1949:16); Barretto and Coutinho (1951:

177-179); Lima (1952:401-404); Veloso, Moura and Klein (1956:520); Forattini (1962:437-

439); Aragao (1964:88-94); Ferreira (1964:335,338,344-345); Forattini, Lopes and Rabello

(1968:136-172); Deane et al. (1970:647); Morales-Ayala (1971:139).

Anopheles (Kerteszia) cruzii cruzii of Lane (1951:336; 1953:285-286); Martins (1958:429-430);

Rachou (1958:171-178); Stone, Knight and Starcke (1959:35); Garcia and Ronderos (1962:

151).

Anopheles cruzii of Dyar and Knab (1908:53, in part); Gabaldon and Cova Garcia (1952:189);

Cova Garcia (1961 :60-61,107,132-133,164).

Anopheles (Dendropaedium) cruzii in part of Dyar (1918:146; 1925a:26).

Anopheles (Nyssorhynchus) bellator var. cruzii of Christophers (1924:42); Edwards (1932:46,

in part).

Anopheles lutzii of Giles (1902:303-304).

Laverania lutzii of Theobald (1902:183).

Myzomyia lutzii in part of Theobald (1905a:8; 1907:41,42, 1910:16,18); Peryassu (1908:59,

78-80, 328-329 ,359).

Nyssorhynchus lutzii of Blanchard (1905:211).

Anopheles bellator in part of Dyar (1923:72).

Anopheles boliviensis in part of Knab (1913:15-17); Dyar and Knab (1917:40); Howard, bya

and Knab (1917:988).

FEMALE (figs. 3,4). Wing: 2.92 mm. Proboscis: 2.02 mm. Forefemur: 1.56 mm.

Abdomen: about 1.7 mm. Head: Integument dark reddish brown to dark brown.

Proboscis 1.2-1.3 length of forefemur. Palpus with small to moderate white patch

at apex of segments 3-5, the patch on segment 3 subequal in size to or larger than

patch on segment 4; scales entirely or predominantly decumbent on segments 3 and

4, sometimes slightly outstanding at base of segment 3. Thorax: Integument of scu-

tum dark reddish brown to dark brown. Larger acrostichal, dorsocentral and middle

scutellar bristles usually predominantly dark. Acrostichal and dorsocentral areas

without scales; scutellum without scales or middle with a few dark scales. Mep

with small upper and larger middle patches of scales. Legs: Hindfemur without

dark scaled line on lower anterior or ventral surface. Hindtarsal segment 1 white

scaled dorsally from 0.2 to 0.5-0.7 and segments 2-5 with broad white band in

apical 0.4-0.6. Wing: Vein C with presectoral, sectoral and subcostal light spots

usually present and moderately large to large; with 1 subcostal spot. Vein R243

without subcostal light spot. Vein R445 with light scales in small basal spot and

long median line extending from 0.2-0.3 to 0.6-0.8. Vein M dark scaled basad of

level of furcation in vein Cu. Vein Cu with subbasal light spot. Vein 1A usually

dark scaled basally. Apical fringe spot small to moderate, inconspicuous to con-

spicuous, undivided; additional inconspicuous fringe spots usually at apex of veins

M344, Cu,, Cu, and 1A. Abdomen: Tergites slightly to conspicuously reddish.

Tergites and sternites II-VII without obvious scales.

Zavortink: Subgenus Kerteszia 25

MALE. Not available for study.

MALE GENITALIA (fig. 16). Segment VIII. Tergite probably without numerous

broad scales medially. Sidepiece: Internal spine flattened and slightly broadened

apically. Claspette: Ventral lobe densely spiculose except laterally; lateral expansion

broad, conspicuous, more or less rounded to sinuous-margined, not curved ventrad

posteriorly, not retrorsely produced anteriorly or with blunt retrorse lobe. Phallo-

some: Aedeagus with strong, long leaflets.

PUPA. Unknown.

LARVA (not illustrated). Association with adult uncertain. Apparently very sim-

ilar to homunculus, possibly distinguished from it by a reddish epidermal pigment

and a more weakly pigmented anal saddle.

SYSTEMATICS. I have seen 3 whole larvae from Brazil that may be cruzii since

they run to it in Forattini’s key (1962:492) and differ in several respects from

larvae of homunculus, bellator and laneanus that I have seen. However, one of

these larvae, from Rio de Janeiro, differs in several important features from the

Other 2, from Brusque, Santa Catarina, and I doubt that it is conspecific with them.

None of these larvae can be completely studied because the terminal segments have

not been cut and mounted laterally, thus obscuring all details of the pecten teeth

and ventral brush.

BIONOMICS. The immatures are found in the leaf axils of bromeliads. Females

readily bite humans, especially during the evening and night. Veloso, Moura and

Klein (1956:520) and Aragao (1964:88-94) report that cruzii is restricted less by

environmental factors than homunculus and bellator in southeastern Brazil and

that it occurs in a variety of habitats from the coast into the mountains.

A. cruzii is a primary vector of human malaria in southeastern Brazil (Rachou,

1958:171-178; Forattini, 1962:554). It is also a natural vector of monkey malaria

in that region (Deane et al., 1970:647).

DISTRIBUTION (fig. 2). Southeastern Brazil; recorded since Komp (1937) also

from Argentina, Bolivia, northern and western Brazil, Colombia, Costa Rica, Ecua-

dor, Guianas, Panama, Peru and Venezuela.

Material Examined: 32 specimens; 2 male genitalia, 27 females, possibly 1-3

larvae.

BRAZIL. Guanabara: Rio de Janeiro, 4 July 1899, A. Lutz, 3 F (lectotype and paralectotypes

of lutzii Theobald, 1901) [BMNH]; M.F. Boyd, 1 F [BMNH], ? 1 L [USNM]. Santa Catarina:

Brusque, 2 M gen, ? 2 L [IER]. Locality unknown, 10 F [IER]. Sao Paulo: Barueri, June 1932,

J. Lane, 7 F [BMNH]. Juquia, 27 Jan 1932, J. Lane, 4 F [BMNH]. Locality Unknown: A. Lutz,

2 F [BMNH].

Additional Records from the Literature

ARGENTINA. Misiones (Garcia and Ronderos, 1962:151).

BOLIVIA (Levi-Castillo, 1949:16).

BRAZIL. Acre, Amazonas, Bahia, Espirito Santo, Minas Gerais, Para, Parana, Pernambuco,

Rio de Janeiro, Rio Grande do Sul, Sergipe (Ferreira, 1964:336-337; Aragao, 1964:88).

COLOMBIA (Levi-Castillo, 1949: 16).

COSTA RICA (Lane, 1953:286).

ECUADOR (Lane, 1953:286).

GUIANAS (Levi-Castillo, 1949:16; Lane, 1953:286).

PANAMA (Lane, 1953:286).

PERU. Loreto (Morales-Ayala, 1971:139).

VENEZUELA. Delta Amacuro (Cova Garcia, 1961:164).

26 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

7. Anopheles (Kerteszia) bellator Dyar & Knab

Figs. 2,4,14,15

1906. Anopheles bellator Dyar and Knab, 1906:160. TYPE: Lectotype male (no. 44.1) with

slides of associated pupal skin, genitalia (314) and foretarsus (659), Pitch Lake, La Brea,

St. Patrick, Trinidad, reared from larva collected in leaf axil of century plant, 8 July

1905, A. Busck [USNM, 10027; designation by Stone and Knight, 1956:276; see Belkin,

Schick and Heinemann, 1965:67]. Synonymized with lutzii Theobald, 1901 (as bellator)

by Dyar (1923:72); considered as variety of lutzii (as bellator) by Christophers (1924:

42); resurrected to specific rank by Dyar (1925a:25-27); considered as variety of lutzii

(as bellator) by Edwards (1932:46); resurrected to specific rank by Komp (1937:506-

508).

1925. Anopheles (Dendropaedium) bellator race bromelicola Dyar, 1925a:27. TYPE: Lecto-

type female, Manoa Woods, Orinoco River, Delta Amacuro, Venezuela, 10 Jan 1910,

F.L. de Verteuil [USNM; designation by Stone and Knight, 1956:276]. Synonymy with

bellator by Dyar (1928:469).

Anopheles (Kerteszia) bellator of Dyar (1928:469-470); Komp (1937:506-508; 1942:75-76, 127-

128, 164-165); Lane (1939:18; 1953:282-283); Rozeboom, Fox and Laird (1941:114); Roze-

boom and Laird (1942:89-91); Simmons and Aitken (1942:99-101); Ross and Roberts (1943:

37-38); Downs and Pittendrigh (1946:47-64); Pinotti (1948:694-695); Pittendrigh (1948:58,

59; 1950a:457-468; 1950b:43-63; 1950c:64-77); Rachou (1956:267,272; 1958:171-178);

Veloso, Moura and Klein (1956:519-520); Charles (1959:160-166); Stone, Knight and Starcke

(1959:35); Forattini (1962:441-445); Aragao (1964:80-86); Ferreira (1964:338, 344-345).

Anopheles (Dendropaedium) bellator of Dyar (1918:145; 1925a:27).

Anopheles (Nyssorhynchus) bellator of Christophers (1924:42, in part); Edwards (1932:46, in

part).

Anopheles bellator of Theobald (1910:86); Dyar and Knab (1917:40); Howard, Dyar and Knab

(1917:985-986); Dyar (1923:72, in part); Downs, Gillette and Shannon (1943:19-20,34-42);

Senior-White and Lewis (1951:151); Gabaldon and Cova Garcia (1952:189); Cova Garcia

(1961:59-60,106,131-132,164).

Myzomyia lutzii of Theobald (1903:51; 1905a:8, in part; 1907:41,42, in part; 1910:16,18, in

part).

Anopheles cruzii of de Verteuil and Spence (1937:453-454).

FEMALE (fig. 4). Wing: 2.69 mm. Proboscis: 1.90 mm. Forefemur: 1.40 mm.

Abdomen: about 2.0 mm. Head: Integument dark brown. Proboscis 1.3-1.4 length

of forefemur. Palpus with small to moderate white patch at apex of segment 4 or

segments 4 and 5, the patch on 4 larger when 2 are present; scales slightly to con-

spicuously outstanding on proximal portion of segment 3, decumbent on distal

portion of segment 3 and on segment 4. Thorax: Integument reddish brown to

dark brown. Larger acrostichal and dorsocentral bristles predominantly pale; larger

middle scutellar bristles predominantly dark. Acrostichal and dorsocentral areas

without scales, scutellum without scales or middle with a few dark scales. Mep

with small upper and larger middle patches of scales. Legs: Hindfemur usually

with dark scaled line along lower anterior or ventral surface. Hindtarsal segment

1 usually without white scales dorsally, segments 2 and 3 with narrow white band

in apical 0.1-0.3, segment 4 with narrow white band in apical 0.2-0.3 and segment

5 usually entirely dark scaled. Wing: Vein C with presectoral, sectoral and sub-

costal light spots usually present and moderately large to large; with 1 subcostal

spot. Vein R243 without subcostal light spot. Vein R445 with light scales in small

basal spot and long median line extending from 0.2-0.3 to 0.7-0.8. Vein M dark

Zavortink: Subgenus Kerteszia 27

scaled basad of level of furcation in vein Cu. Vein Cu with subbasal light spot.

Vein 1A dark scaled basally. Apical fringe spot moderate to large, conspicuous,

undivided; additional inconspicuous fringe spots usually at apex of veins M344,

Cu, and 1A. Abdomen: Tergites light brown to dark brown, blackish or reddish.

Tergites and sternites II-VII without obvious scales.

MALE. Essentially as in female except for sexual characters.

MALE GENITALIA (fig. 14). Segment VIIT: Tergite without numerous broad

scales medially. Sidepiece: Internal spine flattened and slightly broadened apically.

Claspette: Ventral lobe densely spiculose mesally; lateral expansion broad, promi-

nent, rounded, curved ventrad posteriorly, not retrorsely produced anteriorly.

Phallosome: Aedeagus with usually strong, lorig leaflets.

PUPA (fig. 14). Abdomen: about 2.5 mm. Trumpet: 0.31 mm. Paddle: 0.69 mm,

Cephalothorax: Weakly to strongly pigmented, lighter ventrally. Trumpet: Yellow

to amber. Abdomen: Weakly to strongly pigmented, uniform or lighter posteriorly.

Hair 1-VII arising on caudal border of segment. Hair 9-[V moderately long, pointed,

spinelike; 9-V moderately long to long, pointed, spinelike, dissimilar to 9-III and

similar, but shorter and weaker, to 9-VI. Paddle: Obovate; usually not obliquely

truncate; relatively narrow, index 1.6-1.8. Weakly to strongly pigmented, but always

darker than posterior abdominal segments. Marginal spicules along distal portion of

external buttress toothlike or filamentous, short, few, widely spaced, usually not

extending basad into proximal 0.5 of paddle; outer edge distad of external buttress

without spicules. Hair 1-P strongly developed, short. Hair 2-P present.

LARVA (fig. 15). Head: 0.49 mm. Anal Saddle: 0.20 mm. Head: Very weakly

to moderately pigmented, collar darker. Hair 3-C moderately long, moderately

developed, not fusiform; slightly stronger than 2-C. Hair 4-C as strong as or strong-

er than 2-C. Hairs 5-7-C moderately long to long. Antenna: Very weakly to moder-

ately pigmented, apex not darkened. Shaft with relatively small, inconspicuous

spicules. Hairs 2,3-A long, subequal in length. Hair 4-A usually forked apically.

Thorax: Epidermis weakly to strongly pigmented, blackish. Hair 11-P moderately

to strongly developed, moderately long to long. Abdomen: Epidermis weakly to

strongly pigmented, blackish, on entire dorsal surface of segments I-VIII or III and

VI-VIII. Hair 1-I, VII palmate. Palmate hairs small, usually not particularly brush-

like, the leaflets spreading; leaflets moderately long, moderately broad, pointed.

Hair 5-II-V usually distinctly plumose; 5-VII long. Hair 6-VI long, plumose, similar

to 6-III-V. Hair 9-IV-VI usually plumose or pectinate. Spiracular Lobe: Pecten

teeth alternating long and short medially, with spinules usually restricted to basal

portion of external edge. Hair 1-S usually 2, 3b or 2, 3f, sometimes single. Hair

8-S present. Anal Segment: Hair 4a-X moderately developed, much longer than

anal saddle, pectinate or plumose.

SYSTEMATICS. The description and drawings of this species are based largely

on material from Trinidad. The few larvae of bellator from Guyana that have been

examined are much more intensely pigmented that those from Trinidad or Bahia,

Brazil.

BIONOMICS. The immatures are found in the leaf axils of terrestrial and epi-

phytic bromeliads. Females commonly bite humans, especially in the evening.

Pittendrigh (1948, 1950a, b, c) reported that bellator preferred regions of lower

humidity than homunculus in Trinidad and, therefore, was found in drier areas

of the island and higher levels in the forest. Pinotti (1948:694-695), Veloso, Moura

and Klein (1956:519-520) and Aragao (1964:81-86) also report that bellator occurs

in less humid regions than homunculus in southeastern Brazil and that it is the

28 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

species found in open places and secondary forest in the relatively dry area along

the coast.

A. bellator is a primary vector of human malaria in southeastern Brazil (Rachou,

1958:171-178; Forattini, 1962:554) and Trinidad (de Verteuil and Spence, 1937:

453-454; Rozeboom and Laird, 1942:89-91; Downs, Gillette and Shannon, 1943:

40-42) and is thought to be a vector in Guyana (Charles, 1959:160-166). It has

also been found infected with Wuchereria bancrofti (Cobbold) in southeastern Brazil

(Rachou, 1956:267, 272).

DISTRIBUTION (fig. 2). Coastal areas from eastern Venezuela to southeastern

Brazil.

Material Examined: 196 specimens; 15- males, 18 male genitalia, 84 females, 57

larvae, 22 pupae; 22 individual rearings (15 larval, 4 pupal, 3 incomplete).

BRAZIL. Bahia: Caravelas, Jan 1931, N.C. Davis, 1 M [USNM], 1 F, 4 L [BMNH]. Muriqueira

(Gois Calmon), 26 Apr 1929, R.C. Shannon, 1 M, 1 M gen, 2 F [USNM].

GUYANA. Demerara: St. Cuthbert’s Mission (1.5-3.0 km W), 17 Nov 1967, P. Rauch and R.

Hansell (GUY 68), 5 L; same data (GUY 70), 3 L [UCLA]. Essequibo: Barima River, Koriabo,

10 Aug 1901, G.C. Low, 3 F [BMNH]. Kaieteur Plateau, 8-11 Aug 1959, W. Adams, 1 F

[BMNH] . Potaro, May 1910, L.D. Cleare, 1 F [BMNH].

TRINIDAD. Nariva: Biche, 9 July 1964, F. Powdhar (TR 554), 1 IpF (554-126), 1 L; same

data (TR 555), 1 L [UCLA]. Bush Bush Forest, Nariva Swamp, 8 Sept 1964, TRVL (TR 664),

1 lp (664-116); same data (TR 665), 1 lpM (665-118), 1 M gen; same data (TR 667), 1 IpF

(667-103); same data (TR 669), 1 L; 3 Nov 1964, TRVL (TR 803), 2 L; 10 Nov 1964, TRVL

(TR 820), 4 L [UCLA]. Charuma Forest, 27 Aug 1964, A. Guerra (TR 636), 2 L; same data

(TR 644), 1 F [UCLA]. Saddle Hills-Brasso Venado, 21 June 1964, R. Manuel and R. Martinez

(TR 513), 3 F [UCLA]. St. Andrew: Caratal Rd., 3 Dec 1964, F. Powdhar (TR 866), 1 F

[UCLA]. Coryal, 18 June 1964, A. Guerra (TR 497), 1 pM (497-102), 1 M gen; same data

(TR 498), 2 F [UCLA]. Cumaca, 22 Oct 1964, A. Guerra (TR 784), 1 F [UCLA]. Gunapo

or Sangre Grande, Nov or Dec 1936, 2 M, 2 M gen [USNM]. Mt. Becke, 29 Apr 1965, A. Guerra

(TR 1137), 3 F [UCLA]. Mt. Harris, 23-31 July 1924, C.L. Withycombe, 7 F [BMNH]; 16

July 1964, F. Powdhar (TR 572), 1 L; same data (TR 573), 1 F [UCLA]. Mt. Tamana, 19 June

1964, A. Guerra (TR 510), 1 IpM (510-119), 4 IpF (510-102,117,131,153), 1 M gen, 2 L;

same data (TR 512), 7 F [UCLA]. Sangre Grande, 1946, 2 M, 2 M gen, 1 F [USNM]. Tamana,

1944, 2 M, 1 M gen, 5 F; June 1945, 1 F [USNM]. Turure Forest, Aug 1966, F. Guerra (TR

1616), 1 F; Sept 1966, F. Guerra (TR 1617), 1 F [UCLA]. Turure River, Valencia, 5 Apr 1925,

F.W. Urich, 1 F [BMNH]. St David: Matelot, 12 Mar 1964, A. Guerra (TR 176), 2 IpF (176-

118,137), 1 pF (176-111), 1 L; same data (TR 178), 1 IpM (178-120), 1 M gen [UCLA]. Sans

Souci, 5 Mar 1964, A. Guerra (TR 141), 1 Ip (141-156); same data (TR 142), 1 IpM (142-122),

1 M gen [UCLA]. Toco, 6 Mar 1964, A. Guerra (TR 157), 1 lpF (157-107), 2 L; same data

(TR 159), 1 pF (159-166) [UCLA]. St. George: Arena Forest, 17 Aug 1965, A. Guerra (TR

1334), 1 pF (1334-101) [UCLA]. Blanchisseuse, 9 Apr 1964, A. Guerra (TR 291), 1 L [UCLA].

Guanapo, Arima, 2 Feb 1938, E. de Verteuil, 1 F [BMNH]. La Holquetta, Guanapo, 21 Jan

1938, E. de Verteuil, 1 F [BMNH]. Monos Island, Sept 1936, 1 M, 1 M gen, 2 F [USNM].

Talparo, 26 June 1964, A. Guerra (TR 530), 2 IpF (530-106,116), 1 lp (530-154), 1 L; same

data (TR 531), 3 F [UCLA]. St Patrick: Point Fortin, 30 Aug 1945, 8 L [USNM]. County

Unknown: Central Range, 22-31 July 1924, C.L. Withycombe, 2 F [BMNH]. Malestafemolo,

7 Feb 1938, E. de Verteuil, 1 M [BMNH]. Rio Claro, Navet, 20 Aug 1964, A. Guerra (TR 619),

1 F [UCLA]. Locality Unknown: June, A. Busck, 1 F (paralectotype of bellator no. 10027);

F.W. Urich, 1 wing (no. 554); W.H.W. Komp, 12 F, 6 M gen [USNM].

VENEZUELA. Delta Amacuro: Manoa Woods, Orinoco River, 10 Jan 1910, F.L. de Verteuil,

1 F (paralectotype of bromelicola) [USNM]. Monagas: Caripito, July 1937, P.J. Anduze, 1 F

[USNM].

Additional Records from the Literature

BRAZIL. Espirito Santo, Guanabara, Paraiba, Parana, Pernambuco, Rio de Janeiro, Rio Grande

Zavortink: Subgenus Kerteszia 29

do Sul, Santa Catarina, Sao Paulo (Gabaldon and Cova Garcia, 1952:189; Aragao, 1964:80).

SURINAM (Gabaldon and Cova Garcia, 1952:189; Forattini, 1962:445).

VENEZUELA. Sucre (Gabaldon and Cova Garcia, 1952:189).

8. Anopheles (Kerteszia) laneanus Correa & Cerqueira

Figs. 2,4,16

1944. Anopheles (Kerteszia) laneanus Correa and Cerqueira, 1944:112-114. TYPE: Holotype

male (383. H. 13) with slide of genitalia (640), Campos do Jordao, Serra da Mantiqueira,

Sao Paulo, Brazil, J. and F. Lane [FH, 2226 in tube 2227; see Belkin, Schick and

Heinemann, 1971:7]. Synonymized with lutzii Theobald, 1901 (as cruzii) by Correa

(1950:53); resurrected by Barretto and Coutinho (1951:177-179); reduced to subspe-

cies of lutzii (as cruzii) by Lane (1951:336); elevated to specific rank by Forattini

(1962:445).

Anopheles (Kerteszia) laneanus of Martinez (1949:13-15); Barretto and Coutinho (1951:177-.

179); Forattini (1962:445-447); Aragao (1964:77-78).

Anopheles (Kerteszia) cruzii laneanus of Lane (1951:336; 1953:286-287); Martinez and Prosen

(1953:28-29); Bejarano (1957:317); Stone, Knight and Starcke (1959:35); Garcia and Ron-

deros (1962:152).

Anopheles (Kerteszia) cruzii of Dyar (1928:468-469, in part); Komp (1937:504-506); Correa

(1950:53).

Anopheles (Dendropaedium) cruzii in part of Dyar (1918:146; 1925a:26).

Anopheles boliviensis in part of Knab (1913:15-17); Dyar and Knab (1917:40); Howard, Dyar

and Knab (1917:988).

Anopheles bellator in part of Dyar (1923:72).

FEMALE (fig. 4). Wing: 3.67 mm. Proboscis: 2.17 mm. Forefemur: 1.76 mm.

Abdomen: about 2.1 mm. Head: Integument dark brown. Proboscis 1.2-1.4 length

of forefemur. Palpus with moderate white patch at apex of segments 3-5; scales

entirely or predominantly decumbent on segments 3 and 4, sometimes slightly out-

standing at base of segment 3. Thorax: Integument brown to dark reddish brown.

Larger acrostichal, dorsocentral and middle scutellar bristles predominantly pale.

Anterior 0.3-0.4 of acrostichal and dorsocentral areas and middle of scutellum with

a few white scales. Mep with small upper and larger middle patches of scales.

Legs: Hindfemur without dark scaled line along lower anterior or ventral surface.

Hindtarsal segment 1 white scaled dorsally from 0.2 to 0.5-0.6, segments 2, 4 and

5 with broad white band in apical 0.4-0.6 and segment 3 with broad white band in

apical 0.6-0.7. Wing: Vein C with presectoral, sectoral and subcostal light spots

usually present and moderately large to large; with 1 subcostal light spot. Vein

R,43 with or without subcostal light spot. Vein R4,5 with light scales in small

basal spot and long median line extending from 0.2-0.3 to 0.7-0.9. Vein M entirely

or partially light scaled basad of level of furcation in vein Cu. Vein Cu with sub-

basal light spot. Vein 1A with 1 or 2 light spots basally. Apical fringe spot small

to moderate, usually conspicuous, undivided; additional inconspicuous fringe spots

usually at apex of veins M344, Cu,, Cu, and 1A. Abdomen: Tergites tan to dark

brown or reddish. Tergites and sternites II-VI] without obvious scales.

MALE. Essentially as in female except for sexual characters.

MALE GENITALIA (fig. 16). Segment VIII: Tergite probably without numer-

ous broad scales medially. Sidepiece: Internal spine flattened and slightly broad-

30 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

ened apically. Claspette: Ventral lobe very densely spiculose except laterally; lat-

eral expansion broad, prominent, rounded, curved ventrad posteriorly, not retrorse-

ly produced anteriorly. Phallosome: Aedeagus with strong, long leaflets.

PUPA. Unknown.

LARVA (not illustrated). Imperfectly known. Apparently very similar to bella-

tor, possibly distinguished from it by the longer hair 14-P that is not branched

near base.

SYSTEMATICS. I have seen one topotypic male of laneanus with an associated

larval skin. The skin is poorly mounted and cannot be studied or described in de-

tail; it appears, however, to be very similar to bellator and runs to the vicinity of

that species in the key. It does not exhibit any of the diagnostic characteristics

of the larva ascribed to laneanus (strongly barbed or plumose hairs 2-4, 11-C) by

Forattini (1962:445-447). The identity of the larva described by Forattini cannot

be determined at this time.

BIONOMICS. The immatures are found in leaf axils of bromeliads. This species

was suspected to be a vector of human malaria in Bolivia by Martinez and Prosen

(1953:28-29).

DISTRIBUTION (fig. 2). Southeastern Brazil; recorded also from Argentina and

Bolivia.

Material Examined: 19 specimens; 3 males, 2 male genitalia, 13 females, | larva;

1 incomplete individual rearing.

BRAZIL. Rio de Janeiro: Agulhas Negras (Itatiaia), 14 Mar 1941, L. Gomes, 1 F; 20 Mar

1941, L. Gomes and I. Luz, 1 F [UCLA]. Sao Paulo: Campos do Jordao, Nov 1936, J. Lane,

1 IM (742-13); 20 Mar 1937, P.C.A. Antunes, 1 M, 3 F [USNM]; J. Lane, 3 F [USNM], 2 F

[UCLA]. Salesopolis (Boracea), 22 May 1947, Travassos, Travassos F. and Vanzolini, 1 M, 1 M

gen, 2 F [USNM]. Sao Paulo, A. Lutz, 1 F; 1935, P.C.A. Antunes, 1 M gen [USNM].

Additional Records from the Literature

ARGENTINA. Misiones (Bejarano, 1957:317; Garcia and Ronderos, 1962:152).

BOLIVIA. Cochabamba, La Paz (Martinez, 1949:13-15). Santa Cruz (Martinez and Prosen,

1953:28-29).

9, Anopheles (Kerteszia) boliviensis (Theobald)

Figs. 2,4

1905. Kerteszia boliviensis Theobald, 1905b:66-68. TYPE: Holotype female, Songo [Zongo,

La Paz], Bolivia, M. Biro [HNM; see Belkin, 1968:9]. Synonymized with lutzii Theo-

bald, 1901 (as boliviensis) by Knab (1913:16); resurrected by Dyar and Knab (1918:

140-141).

Anopheles (Kerteszia) boliviensis of Dyar (1918:148; 1928:467); Komp (1936:68, in part); Lane

(1939:18, in part); Anduze (1943:191, ?); Roca-Garcia (1944:160-169, in part, ?); Levi-Cas-

tillo (1945:128-139,150, in part, ?; 1949:16, in part); Rachou (1958:146, in part, ?); Stone,

Knight and Starcke (1959:35, in part); Aragao (1964:76-77, in part); Morales-Ayala (1971:

139, in part, ?).

Anopheles (Nyssorhynchus) boliviensis of Christophers (1924:42); Edwards (1932:46).

Kerteszia boliviensis of Theobald (1907:118-120; 1910:74); Dyar and Knab (1918:140-141);

Stone (1957:171).

Anopheles boliviensis in part of Knab (1913:15-17); Dyar and Knab (1917:40); Howard, Dyar

and Knab (1917:988); Bates (1943:23; 1944:168,169; 1945:24); Gabaldon and Cova Garcia

(1952:189); Cova Garcia (1961:57-58,104,134-135,162, ?). .

Anopheles (Dendropaedium) cruzii in part of Dyar (1925a:26).

Zavortink: Subgenus Kerteszia 31

FEMALE (fig. 4). Wing: 3.95 mm. Proboscis: 2.44 mm. Forefemur: 2.02 mm.

Abdomen: about 2.5 mm. Head: Integument dark reddish brown to dark brown.

Proboscis 1.2-1.4 length of forefemur. Palpus with moderate white patch at apex

of segments 3-5; scales slightly to conspicuously outstanding on proximal portion

of segment 3, decumbent on distal portion of segment 3 and on segment 4. Thorax:

Integument reddish brown to dark reddish brown or brown. Larger acrostichal and

dorsocentral bristles predominantly pale; larger middle scutellar bristles dark. Ante-

rior 0.3-0.4 of acrostichal and dorsocentral areas with a few white scales; middle -

of scutellum with a few dark scales. Mep with small upper patch of scales. Legs:

Hindfemur with complete or broken dark scaled line along lower anterior or ven-

tral surface. Hindtarsal segment 1 white scaled dorsally from 0.1-0.2 to 0.3-0.7,

segments 2, 4 and 5 with broad white band in apical 0.5-0.7 and segment 3 with

broad white band in apical 0.7-0.8. Wing: Vein C with presectoral, sectoral and

subcostal light spots usually present and moderately large to large; with 1 sub-

costal light spot. Vein R,43 with subcostal light spot. Vein R44; with light scales

in small basal spot and long median line extending from 0.2-0.3 to 0.7-0.8. Vein

M entirely or predominantly light scaled basad of level of furcation in vein Cu.

Vein Cu with subbasal light spot. Vein 1A usually with 1 light spot basally. Api-

cal fringe spot small to moderate, inconspicuous to conspicuous, undivided; addi-

tional inconspicuous fringe spots usually at apex of veins M344, Cu,, Cu, and 1A.

Abdomen: Tergites tan to dark brown. Tergites II-VII with numerous narrow to

moderately broad, dark brown to black scales; scales of distal tergites not out-

standing, and not forming transverse apical bands. Sternites II-VII with white scales

medially; more distal sternites sometimes with additional dark scales apically.

MALE, PUPA, LARVA. Unknown.

SYSTEMATICS. This species is known only in the adult female; the larva and

male described as boliviensis by Komp and Osorno-Mesa (1936) are actually lepi-

dotus. The description and drawings of boliviensis are based largely on material

from Colombia and Peru.

BIONOMICS. Some of the information on ‘‘boliviensis’’ summarized under lepi-

dotus may actually pertain to this species.

DISTRIBUTION (fig. 2). Eastern slope of Andes from Colombia to Bolivia; re-

corded also from Brazil, Guyanas, Paraguay and Venezuela.

Material Examined: 38 females.

BOLIVIA. Cochabamba: Chimore, Chapare, 29 Apr 1944, Torres Munoz, 1 F [USNM].

COLOMBIA. Meta: Restrepo, W.H.W. Komp (KO 121A-10), 1 F [UCLA]. Villavicencio,

Forzosa Bosque, 7 July 1943, 1 F [USNM]. Locality Unknown: J.A. Kerr, 28 F [UCLA].

PERU. ? Cajamarca: Huascaray, 22 Sept 1911, C.H.T. Townsend, 4 F [USNM]. Rio Charape,

13 Sept 1911, C.H.T. Townsend, 3 F [USNM].

Additional Records from the Literature

BRAZIL (Levi-Castillo, 1949:16; Rachou, 1958:146).

BOLIVIA. Beni, Cochabamba, La Paz (Gabaldon and Cova Garcia, 1952:189).

COLOMBIA. Cundinamarca, Meta (Gabaldon and Cova Garcia, 1952:189).

ECUADOR. Napo-Pastaza, Santiago-Zamora (Levi-Castillo, 1945:150; Gabaldon and Cova

Garcia, 1952:189).

GUIANAS (Levi-Castillo, 1949:16).

PARAGUAY (Levi-Castillo, 1949:16).

PERU. Amazonas, Cuzco, Madre de Dios, San Martin (Gabaldon and Cova Garcia, 1952:

189). Ayacucho, Loreto (Morales-Ayala, 1971:139).

VENEZUELA. Merida (Anduze, 1943:191; Cova Garcia, 1961:162).

32 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

10. Anopheles (Kerteszia) sp., Auyan-Tepui Mesa form

Fig. 1

Anopheles (Kerteszia) sp. of Anduze (1941:823; 1942:435).

This form is known by a single female from Auyan-Tepui Mesa, Bolivar, Vene-

zuela (as Oyantepuy, June 1937, P.J. Anduze) in the USNM that apparently resem-

bles neivai in most respects (distribution of scales on mep unknown), but differs

from that species by its larger size, the presence of a patch of white scales at the

apex of palpal segment 3 and by the narrower apical white band on hindtarsal

segments 1-4. This form almost certainly represents a distinct species, but since

the only specimen is badly damaged, I am not naming or describing it at this time.

REFERENCES CITED

Anduze, Pablo J.

1941. Primer informe sobre entomologia medica del Estado Bolivar (Venezu-

ela). Serie 1. La fauna culicidiana. Descripcion de tres especies nuevas (Dip-

tera, Culicidae). Rev. Sanid. Asist. Soc. 6:812-836.

1942. Sobre la morfologia de la armadura bucofaringea de algunos representantes

venezolanos del subgenero Kerteszia (Diptera: Culicidae). Rev. Sanid. Asist.

Soc. 7:435-436.

1943. Estudios de entomologia medica en el Estado Merida-Venezuela. La fauna

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2:189-196.

Aragao, Mario B.

1964. Distribuicao geografica e abundancia das especies de Anopheles ( Kerteszia)

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Barretto, M. Pereira and J.O. Coutinho

1951. Sobre o Anopheles (Kerteszia) cruzii Dyar & Knab e 0 Anopheles ( Kertes-

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15:177-180.

Bates, Marston

1943. Mosquitoes as vectors of Dermatobia in eastern Colombia. Entomol. Soc.

Am., Ann. 36:21-24.

. 1944. Observations on the distribution of diurnal mosquitoes in a tropical forest.

Ecology 25:159-170.

1945. Observations on climate and seasonal distribution of mosquitoes in eastern

Colombia. J. Anim. Ecol. 14:17-25.

Bejarano, Juan F.R.

1957. Distribucion geografica de ““Anophelini” de la Republica Argentina. Rev.

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Belkin, John N.

1962. The mosquitoes of the South Pacific (Diptera, Culicidae). Vol. 1. Berkeley,

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69 p.

Zavortink: Subgenus Kerteszia 33

Belkin, John N., R.X. Schick and S.J. Heinemann

1965. Mosquito Studies (Diptera, Culicidae). V. Mosquitoes originally described

from Middle America. Am. Entomol. Inst., Contrib. 1(5). 95 p.

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Blanchard, Raphael

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1961. Distribuicao geografica dos mosquitos da Amazonia (Diptera, Culicidae,

Culicinae). Rev. Bras. Entomol. 10:111-168.

Charles, L.J.

1959. Observations on Anopheles (Kerteszia) bellator D. & K. in British Guiana.

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Christophers, S. Rickard

1924. Provisional list and reference catalogue of the Anophelini. Indian Med.

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Correa, Renato R.

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1956. Type specimens of mosquitoes in the United States National Museum:

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1901. A monograph of the Culicidae or mosquitoes. Vol. 1. Lond., Br. Mus.

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Fig. 1

KERTESZIA

Gers NS,

KS) Sa 3

. Tropic of Capricory

bambusicolus

1000

STATUTE MILES

KERTESZIA

NORTH & SOUTH AMERICA

KILOMETERS

1 2000

1000

STATUTE MILES

KERTESZIA Fig.3

neivai

PAIS TS

Darien

Panama

in Hine me \ . » gy sa S: wh:

ii HAN WATER Nh Ai i mn i ia k =a NAN oa WwW

pholidotus

Bocas del Toro

Panama

ey i ih My ii y — Se, ae Gh

mn UN Te TW or \\\ SS

Hii ii HA Ah “8 a aA ee \\\\es

lepidotus SESS SE

Meta

Colombia

Minin

m Hii Hii

Nii \

Nii iin emi

ANY ii NN oem, bene

et +“

bambusicolus

Meta

Colombia i

1m ee

Wit iy _——

m Hi ni Bilin ni ii i ATS aRaRRRANN RH

LiWidiniaiuuiluuuinll

Hil

homunculus

TR 654 ¥

St. Andrew me

Trinidad =

" " Tis y S : te Wes > ra re

ee Sa,

Wiiiiblidiishiini RC: eae a

cruzil

Sao Paulo

Brazil

Ss —

AW \

i, _

bidididibinnan A, eae TN

KERTESZIA Fig. 4

htc —

Essequibo

Guyana

ih iii nt a

laneanus

Sao Paulo

Brazil

iM ~

\ TS ‘ SR aa

ND lines ss AK. §

Hii ih i nl wilh WN Mi ii i hi iN WY iy: a STN ¥ rene 5 WW: W

E3Ba

SAB

EAB}

—sa

—s

boliviensis

Cochabamba

Bolivia

\\t

‘io

A) ra

iia

VVevye

pholidotus Bocas del Toro

Panama | | | |

lepidotus ge.

bambusicolus mas

Fiisndplis TR 4

Trinidad |

cruzii Sao Paulo

Brazil | | |

Trinidad | | |

bell ator rete

laneanus es eee

Brazil | | | |

bolivicner Cochabamba

Bolivia | | |

FORE

MID

HIND

Fig.5

KERTESZIA

Cunal Zone

Panama

basal

humeral presectoral

SSS SSS

0 SS

SEZ gy fe g

Zo zp:

PA 363, 373, 964, 990

Darien

Panama

sectoral

AARTRAR

MANNA VN

subcostal

preapical

"Weruvbe

Fig. 6

KERTESZIA

i ie

. KERTESZIA

3 Gf

pon

/ (Ui

4 saggy? 11

: ag / ; | FF

neival

PA 363

Darien

Panama

Fig.8

KERTESZIA

\ |

SS WA

\\\

\ SD y : |

‘ “ )

~\ Af | L :

NY \ ' Ww

YY : 4 \ , ,

\ \\ Be tee ne

\ LAN 2 4

QW AF

. , ° A, . . det Z ; 3

, sia

e N

- KERTESZIA | \. | Fig.9

oy >

A ie}

a1 9

és, Cc

1 f 2

1 bmh

{

7 6| 5 13

° 1.0

ery : | pholidotus

PA 173

Bocas del Toro

Panama

tj a

Z, ——

Fig. 10

KERTESZIA

Meta

Colombia

ambusicolus

KERTESZIA

bambusicolus

Fig.12

KERTESZIA

TR 1046

KERTESZIA

homunculus

TR 564

St. Andrew

Trinidad

Fig. 14

KERTESZIA

2 rt » @

‘le 3 =

=> I

yf |!

; “py

yf | Ul

($s

ee Pees r, 4

x At 4 i

TR 178, 665 Pi leah

St. David, Nariv

Trinida

KERTESZIA

bellator

TR 803

Nariva

Trinidad

Fig. 16

KERTESZIA

laneanus

Sao Paulo

Brazil

54 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

FIGURES

Distribution of Kerteszia species

Wings of Kerteszia species

Wings and hindtarsi of Kerteszia species

Anopheles (K.) neivai; adult morphology

Anopheles (K.) neivai; male genitalia and pupa

Anopheles (K.) neivai; larva |

Anopheles (K.) pholidotus; male genitalia and pupa

Anopheles (K.) pholidotus; larva

10. Anopheles (K.) bambusicolus; male genitalia and pupa

11. Anopheles (K.) bambusicolus, larva

12. Anopheles (K.) homunculus; male genitalia and pupa

13. Anopheles (K.) homunculus, larva

14. Anopheles (K.) bellator; male genitalia and pupa

15. Anopheles (K.) bellator; larva

16. Anopheles (K.) lepidotus, cruzii and laneanus; male genitalia

OW NAAKRWN >

SYSTEMATIC INDEX

adolphoi, 24

Anopheles, 1, 4

anoplus, 21

Auyan-Tepui Mesa form, 32

bambusicola, 19

bambusicolus, 2, 3, 4, 6, 7, 8k, 9k, 10k, 18-20

bellator, 1,3, 4, 8k, 9k, 10k, 11k, 11, 22, 25, 26-29, 30

boliviensis, 2, 3, 5, 7k, 8, 9, 10, 16, 30-31

boliviensis of authors, 4, 18, 31

bromelicola, 26

cruzii, 3, 4,5, 8k, 9k, 9, 10, 11, 23-25, 29

Dendropaedium, 5

homunculus, 1, 3, 4, 8k, 9k, 10k, 20-23, 25, 27

hylephilus, 11

laneanus, 3, 4, 8k, 9k, 9, 10, 25, 29-30

lepidotus, 2, 3, 4, 7k, 9k, 9, 10k, 16, 17-18, 20, 31

lutzii Cruz, 23

lutzii Theobald, 2,5, 11, 23, 24, 26, 29, 30

Kerteszia, 1, 2, 3, 4, 5, 5-7, 18

Kerteszia sp. 10, 2, 4, 7, 8, 9, 10, 32

montemor, 24

neivai, 1, 2, 3, 4, 8k, 9k, 10k, 11-15, 32

Nyssorhynchus, 3, 4

pholidotus, 2, 3, 4, 7k, 9k, 10k, 15-17, 18

MOSQUITO STUDIES (Diptera, Culicidae)

XXX. A NEW SUBGENUS AND SPECIES OF CULEX

FROM COLOMBIA’

Abdiel J. Adames and Pedro Galindo’

During biomedical studies conducted by the Office of. Interoceanic Canal Studies,

LTC Bruce F. Eldridge and his associates collected a new species of Culex, which

Dr. John N. Belkin and ourselves consider to belong to a hitherto undescribed

higher taxon of the genus Culex herewith described as a new subgenus. We take

pleasure in naming this taxon after Dr. John N. Belkin, whose numerous and bril-

liant contributions to the morphology and systematics of the Culicidae of the

World have led to a better understanding of this important group of insects. We

are also pleased to designate the new species after LTC Bruce Eldridge whose well

organized surveys along the proposed interoceanic canal routes 17 and 25 yielded

material from which basic knowledge on the Culicidae of Middle America has been

derived.

The presentation and description of the new taxa in general follow Belkin (1962).

For the illustrations we are indebted to N. Kitamura and L.M. Kowalczyk, and for

the text copy for lithoprinting to A. Demos.

BELKINOMYIA, new subgenus

TYPE SPECIES: Culex (Belkinomyia) eldridgei, n. sp.

FEMALE. Head: Eyes not distinctly separated above base of antennae. Decum-

bent scales on vertex narrow, with a group of linear scales toward sides and a patch

of broad scales laterally. Erect scales numerous, forked, extending toward sides.

Clypeus bare. Proboscis slightly longer than forefemur. Palpus very short, 3-seg-

mented, segments 1 and 2 ankylosed. Antenna slightly longer than proboscis; flagel-

lar whorls moderately long, decreasing in length distally; torus with a few short hairs

in anterior mesal surface. Thorax: Mesonotum and scutellum with narrow, curved

decumbent scales, except for long narrow inner dorsocentral bare lines. Acrostichal

bristles absent on disc. Dorsocentrals and marginal scutellars strongly developed.

Paratergite bare. Pleuron with scales on.ppn and stp; bristles present on apn, ppn,

‘Contribution from project “Mosquitoes of Middle America” supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478. This work was supported in part by Grant AI-02984

from the National Institute of Allergy and Infectious Disease and by Contract DADA-17-67-7020,

U.S. Army Medical Research and Development Command.

* Gorgas Memorial Laboratory, Apartado 6991, Panama 5, Panama.

56 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

ppl, and pra; a single strong bristle on Jmep. Base of haltere with a few microsetae.

Legs: Claws simple. Wing: All scales squamous, moderately broad. Abdomen: Later-

otergite with many short to moderately long bristles.

MALE. Essentially similar to female except for sexual characters. Sexual dimor-

phism of head appendages not marked. Palpus very short. Antenna distinctly longer

than proboscis; flagellar whorls essentially as in female but slightly longer; segments

12 and 13 more elongate than others; torus swollen. Claws of foreleg and midleg

unequal, simple. Hindclaws as in female.

MALE GENITALIA. Segment IX: Tergite with a pair of small hairy lobes. Side-

piece: Subapical lobé somewhat as in the genus Deinocerites; with 4 differentiated

setae, 3 of which are spiniform and the other foliform. Clasper: Unique for the

genus Culex, short and broad with an apical forked spiniform as in Deinocerites.

Phallosome: Basal part of lateral plate (Belkin et al. 1970:78; “basal hook” of

authors) mostly membranous, narrowly sclerotized along lateral margin; membra-

nous areas bearing some spines. Proctiger: Crown of paraproct comblike, with a

row of short and heavy teeth.

PUPA. Cephalothorax: All hairs present, variously developed. Hair 5-C subequal

in length to 4+C and about 0.33 of trumpet. Hair 8-C multibranched, the longest

hair on cephalothorax, inserted at a level caudad of base of trumpet. Metanotum:

Hair 10-C longer than 11-C, with multiple weak branches. Abdomen: Hair 1-II, [Il

with 20 or more branches, slightly resembling float hair. Hair 2-IJI-VI nearly mar-

ginal; hair 2-III laterad of 1-III; hair 5-IV-VI weakly developed, barely extending

to middle of succeeding tergite; hair 9-VII shorter than 3-VII and less than 0.5 of

tergite VIII; hair 1-I[X small, single. Paddle: Hairs 1,2-P distinct, single.

FOURTH INSTAR LARVA. Head: Head capsule without spicules or lateral ex-

pansion on each side caudad of antenna. Labrum well differentiated dorsally. Hairs

2,3-C apparently not developed; hairs 4-6-C closely grouped together and caudad

of level of 7-C; hair 6-C single. Thorax: Hairs 1-3-P on a distinct tubercle; 1,2-P

long, single; 3-P with 4-6 weak branches. Abdomen: Comb in a patch of 4 irregular

rows. Siphon: Pecten teeth simple. One short dorsolateral tuft distad of ventro-

lateral tufts. Hair 2-S distinct, slightly curved, without a recurved tooth on shaft.

Anal Segment: Ventral brush (4-X) with 6 pairs of hairs on grid.

Culex (Belkinomyia) eldridgei, n. sp.

Figs. 1,2

TYPES: Holotype male with slides of associated larval and pupal skins and genitalia (OICS-

0197-2), Curiche, El Choco, Colombia, larva from crabhole, 6 August 1967, B.F. Eldridge

[USNM]. Allotype female with slide of associated larval and pupal skins (OICS-0197-3), same

data as holotype [USNM]. Paratypes: 7 1pM (OICS-0197-6,7,8,15,16,17,18), 3 1pF (OICS-0197-

9-11), 3 M (OICS-0197-1,4,14), 1 F (OICS-0197-5), same data as holotype [GML, UCLA].

FEMALE. Wing: 2.28 mm. Proboscis: 1.54 mm. Forefemur: 1.16 mm. Abdomen:

about 1.60 mm. Small; mesonotum dark brown to black, contrasting with lighter

portions of pleuron; abdomen and legs without distinct ornamentation. Head: De-

cumbent and erect scales of vertex. brown. Orbitals and interobitals strongly devel-

oped, 6 pairs of upper orbitals, heavier, longer and more widely spaced than lower.

Clypeus dark brown. Proboscis about 1.20 of forefemur, dark scaled, with numerous

bristles. Palpus short, about 0.2 of proboscis; segment 2 about 0.40 of segment 3,

Adames & Galindo: New Culex Subgenus 57

both dark scaled. Antenna about 1.20 of proboscis. Torus dark brown; flagellar

whorls of segments 2-13 moderately long, progressively slightly shortened distad,

last whorl distinctly shorter than segment. Thorax: Integument dark brown. Meso-

notum and scutellum with auburn to dark scales; inner dorsocentral bare lines

extending from anterior margin to 0.75. Acrostichal bristles absent on disc, slightly

developed on anterior promontory; 3 posterior fossals and 1 parascutellar. Pleuron

pale brown to brown. Apn, ppn, ppl, psp, upper stp, pra and upper 0.67 of mep

darker than lower stp and basal 0.33 of mep and meron; ppn with a few bristles on

dorsocaudal angle and with narrow curved scales on upper part; ppl with a few (6)

bristles, stp with bristles arranged in a single continuous curved row along dorsal and

caudal margins and with a few scattered broad scales among bristles; umep with a

few small and poorly developed bristles. Legs: Coxae with scales and setae; anterior

surface of forecoxa largely covered with scales and bristles, basolateral area with a

few bristles; midcoxa with scales anterolaterally and with 2 rows of setae laterally,

the posterior stronger; hindcoxa with a few strong bristles on anterobasal area.

Trochanter with scales and bristles. Forefemur and midfemur predominantly dark

scaled; antero and posterolateral surfaces of hindfemur whitish. All tibiae and tarsi

dark scaled. Wing: Veins entirely dark scaled. Haltere: Stem pale brown, knob

entirely dark scaled. Abdomen: Tergites II-VII completely dark scaled. Sternites

with creamy scales.

MALE. Wing: 1.98 mm. Proboscis: 1.35 mm. Forefemur: 1.04 mm. Abdomen:

(not including genitalia) 1.54 mm. Palpus, about 0.2 of proboscis, apparently 3-

segmented, segment 1 without scales, partly ankylosed with 2; segment 3 about

2.0 of 2, both dark scaled. Antenna about 1.40 of proboscis.

MALE GENITALIA (fig. 1). Segment [X: Forming a complete ring. Lobes of

tergite round, well separated, each lobe articulated ventrally with a lobe of the

basolateral sclerotization of proctiger; sternite small, bare. Sidepiece: Conical, with

a few strong and numerous small bristles. Subapical lobe at about 0.67 from base,

projecting dorsad; with 4 differentiated setae, 3 of which are spiniform (a basad of

b and c laterad of b), the other longer, foliform and at base of distal margin of lobe;

tergal surface with a large patch of hairs slightly shorter than the longest hair of

subapical lobe. Clasper: Short and broad; inner margin convex and with broad distal

expansion; outer margin convex. Apex with a heavy forked spiniform; 2 subapical

setae, one ventral, the other subdorsal and longer. Phallosome: Lateral plate, in side

view, with basal part consisting of a very strongly sclerotized hook with apex

projecting sternad (basal hook of authors); a moderately sclerotized area with a few

spines basad of marginal sclerotization of hook; a large moderately sclerotized area

bearing lateral spines extending dorsad of hook and joined to distal part by a poorly

sclerotized band; distal part very strongly sclerotized, much shorter than basal part,

with a sinuous apical border projecting dorsad into a short rounded horn. Proctiger:

Strongly developed; paraproct strongly sclerotized; basolateral sclerotization with

conspicuous tergal lobe and without free sternal process. Cercal setae 2,3.

PUPA (fig. 1). Abdomen: 2.31 mm. Trumpet: 0.38 mm. Paddle: 0.49 mm.

Cephalothorax: Integument light brown; hair 6-C markedly shorter than 7-C; hairs

6,7-C very close together, hair 8-C at least 5-branched (5-8). Trumpet: Not placed

on distinct tubercle; moderately long; index about 6.0-7.0; integument light brown,

darker on tracheoid; pinna about 0.25-0.28 of total length. Metanotum: Hairs 10,

11-C removed from 12-C but moderately close together. Abdomen: Integument

lightly pigmented. Hair 3-I double or triple; hair 6-II-VI usually single; hair 9-II-VI

caudolaterad of 6; hair 9-VII usually with more than 3 branches (3-6); hair 9-VIII

58 Contrib. Amer. Ent. Inst., vol. 9, no. 3, 1973

at least 4-branched (4,5), weakly developed, distinctly shorter than length of tergite

VIII; hair 4-VIII usually single (1,2). Paddle: Longer than wide; without any pig-

mented spot. Midrib well developed; external buttress slightly developed; apex

smoothly rounded, margins without spicules.

FOURTH INSTAR LARVA (fig. 2). Head: 0.66 mm. Siphon: 1.06 mm. Head:

Integument yellowish. Head capsule width about 1.2 of length. Mental plate well

developed, with 10 lateral teeth, median tooth not shouldered. Collar moderately

developed, narrow. Maxillary suture complete, extending caudolaterad to collar.

Posterior tentorial pit a short distance from caudal border. Hair 1-C strong, straight;

4-C moderately long, weak, single or double, cephalomesad of 5 and mesad of 6;

hairs 5,6-C strongly developed, 5-C usually with 3 branches (3,4), 6-C slightly longer

than 5-C (1.14); hair 7-C moderately long, with 3-5 barbed branches; 11-C with 3-5

moderately long, weak branches; 13-C closer to 11-C than 12-C; basal maxillary hair

(bmh) small, single or double; 16,17-C not developed. Antenna: with numerous

spicules in basal part. Hair 1-A inserted about 0.75 from base, with 12-16 barbed

branches. Thorax: Without any apparent outstanding features; integument glabrous.

Hair 14-P usually double (1,2); O-P, 1,13,14-M, 1,3,13-T with short weak branches,

not stellate. Abdomen: Integument glabrous. Hairs 1,2,4,11,13-I, 1,2,4,9,13-H, 1,

2,5,7,9,13-II-VI, 1,2,5,8,10,13-VI unbranched or with branches of varied length

and thickness, never stellate; 6-I,II long, double, branches unequal; 7-I long, single;

7-II short, with 4-6 branches; 6-III-V moderately long, double, branches unequal;

6-VI single or double. Segment VIII: Comb scales spatulate, apex fringed. Hair 1-Il

with 3,4 branches; 5-VIII double or triple; 1,2-VIII not on tubercle or sclerotized

plate. Siphon: Integument yellowish. Moderately long; index about 6.0-6.5. Acus

distinct, attached. Pecten extending to 0.33. Hair 1-S distinctly caudad of last

pecten tooth; 4 pairs of moderately long subventral hairs (1,la-S), the apical short-

est. Anal Segment: Integument yellowish. Saddle complete. Acus not developed.

Caudal margin without spines. Hair 1-X short, weak, double or triple; hair 4f-X

short, shorter than 1-X, usually single.

SYSTEMATICS. Belkinomyia can best be separated from other subgenera of

Culex in the male. The very short palpus, the sparsely plumose antenna and the mor-

phology of the clasper and of the subapical lobe of the sidepiece separate this new

subgenus from all other Culex. The nonplumose antenna, the appendages of the

subapical lobe of the sidepiece and the bifid terminal spine of the clasper appear to

indicate a relationship between Belkinomyia and the genus Deinocerites. In the fe-

male, the following combination of characters separates Belkinomyia from most

other subgenera of Culex: (1) lack of acrostichal bristles, (2) presence of dorso-

central bare lines, (3) poorly developed propleural bristles, and (4) moderately

broad scales on veins R2 and R3. In the pupa, the position of hair 2-III, which

in Belkinomyia is laterad of 1-III, separates it from most other Culex. The short

hair 9-VII also serves to distinguish Belkinomyia, as it is shorter than 3-VII and less

than 0.5 the length of segment VIII, while in most other Culex the reverse is true.

The larva apparently does not show any particular character which can by itself

separate Belkinomyia from most other subgenera of Culex. However, it can be

separated from other individual American subgenera as follows: (1) from Culex by

prothoracic hair 3-P being weaker and shorter than 1-P, less than 0.5 of its length,

(2) from Neoculex by the presence of a pair of short dorsolateral tufts above the

ventrolateral tufts on the siphon, (3) from Lutzia by the mouthbrushes being normal

in position, numerous and filamentous, (4) from Carrollia by the absence of hair

Adames & Galindo: New Culex Subgenus 59

2-C, (5) from Melanoconion and Mochlostyrax by siphonal hair 2-S being slightly

curved and simple, (6) from Microculex and Anoedioporpa by the ventral brush

with 6 pairs of hairs, (7) from Aedinus by the simple pecten teeth, (8) from

Micraedes by the anal saddle without long caudal marginal spicules, and (9) from

Tinolestes by the absence of spicules on the head capsule and the presence of only

1 pair of dorsolateral hairs on the siphon.

BIONOMICS. The immature stages occur in crabholes with water of low salinity

in association with Culex latisquama (Coquillett, 1906) and Deinocerites curiche

Adames, 1971. The adults rest in crabholes.

DISTRIBUTION. C. eldridgei is known by the authors to occur only at the type

locality, in Curiche and nearby Coredo, in the Department of El Choco, along the

Pacific coast of Colombia. Its distribution may possible extend farther south to

Buenaventura. Material examined: 32 specimens; 24 males, 8 females; 12 larval

rearings.

COLOMBIA. El Choco, Curiche, B.F. Eldridge (OICS-0197), (GML-03827-33) [USNM, GML,

UCLA]. El Choco, Curiche, Coredo, A.J. Adames and A. Quinonez (COA-5) [UCLA].

REFERENCES CITED

Belkin, John N.

1962. The mosquitoes of the South Pacific (Diptera, Culicidae). Vol. 1. Berkeley,

Univ. Calif. Press. 608 p.

Belkin, John N., S.J. Heinemann and W.A. Page

1970. The Culicidae of Jamaica (Mosquito Studies. XXI). Am. Entomol. Inst.,

Contrib. 6(1). 458 p. (Also published as Inst. Jam., Bull. Sci. Ser. 20)

FIGURES

1. Culex (Belk.) eldridgei; male genitalia and pupa

2. Culex (Belk. ) eldridgei; larva

Curiche

El Choco

Colombia

BELKINOMYIA

eldridgei

Curiche

El Choco

Or Colombia VII

WHAM

ee ‘ bs s

€

- ee

Contributions

of the

American Entomological Institute

Volume 9, Number 4, 1973

MOSQUITO STUDIES (Diptera, Culicidae)

XXXII. A revision of the subgenus Carrollia of Culex

By Jose D. Valencia

CONTENTS

INTRODUCTION .. . :

MATERIAL AND METHODS

TAXONOMIC CHARACTERS

SYSTEMATICS

BIONOMICS . . :

TAXONOMIC TREATMENT

Subgenus Carrollia

Keys to Groups and Species

Bihaicolus Group. .

1. Culex (Car.) bihaicolus

2. Culex (Car.) guerreroi .

3. Culex (Car.) rausseoi

3a. Culex (Car.), Panama form .

4. Culex (Car.) metempsytus .

5. Culex (Car.) infoliatus

Iridescens Group . ,

Urichii Subgroup. . .

6. Culex (Car.) at

7. Culex (Car.) anduzei

Iridescens Subgroup.

8. Culex (Car.) bonnei

8a. Culex (Car.), Bahia form

9. Culex (Car.) secundus .

9a. Culex (Car.), Espirito Santo foun ,

10. Culex (Car.) iridescens

11. Culex (Car.) soperi . Ca

12. Culex (Car.) babahoyensis .

13. Culex (Car.) antunesi .

14. Culex (Car.) cerqueirai

15. Culex (Car.) wilsoni

16. Culex (Car.) kompi .

REFERENCES CITED

FIGURES ..

TABLE OF DISTRIBUTIONS :

INDEX TO SCIENTIFIC NAMES

MOSQUITO STUDIES (Diptera, Culicidae)

XXXI. A REVISION OF THE SUBGENUS CARROLLIA OF CULEX’

Jose D. Valencia”

INTRODUCTION

Carrollia, precinctive to the New World tropics, is probably the most distinctive

subgenus of Culex and includes the only species with conspicuous metallic orna-

mentation in the adults in the entire genus. When I undertook this revision in 1969,

the 14 described nominal species assigned to the subgenus were differentiated

almost entirely on the basis of male genitalic characters and no larvae or pupae

were adequately known for any species (Lane, 1953:496-498; Levi-Castillo, 1953:

92-94). My work stimulated other investigators to reexamine the group and recently

2 additional nominal species have been described from Venezuela and 2 other

species have been redescribed from Brazil. In this revision, I am recognizing 16

species in the subgenus, with the reduction of 2 formerly regarded as taxonom-

ically valid to synonymy and the description of 2 new species. For 14 of these

species all the stages are associated through individual rearings and for kKompi and

cerqueirai the association of the pupa and larva, respectively, are presumptive only.

In addition, | am recognizing 3 unnamed forms and it is very probable that there are

2 or 3 other species in the subgenus which I have tentatively identified as the previ-

ously described antunesi and wilsoni. These unresolved taxonomic problems are pri-

marily due to the lack of material from Brazil.

I am indebted to John N. Belkin for encouragement and guidance in this study

and in the preparation of the manuscript; to Abdiel J. Adames, J. Hal Arnell

and Thomas J. Zavortink for assistance and stimulating discussions; and to Sandra

J. Heinemann and William A. Powder for the preparation of material. For the

loan of material I thank: Sebastiao H. Xavier, Centro de Pesquisas ‘‘Rene Rachou”,

Belo Horizonte, Brazil; Pablo Cova Garcia, Division de Endemias Rurales, Mara-

cay, Venezuela; Pierre Fauran, Institut Pasteur de la Guyane Francaise; and Willis

W. Wirth, U.S. National Museum of Natural History. I thank Felipe Martin, Fac-

ultad de Agronomia, Maracay, Universidad Central de Venezuela, for providing

laboratory facilities at Rancho Grande field station; Lacenio Guerrero, Division

de Malariologia y Endemias Rurales, Maracay, Venezuela, for providing transpor-

‘Contribution from project “Mosquitoes of Middle America” supported by U.S. Public Health

Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command

Research Contract DA-49-193-MD-2478. Based on PhD dissertation submitted to the University

of California, Los Angeles; research conducted during tenure of a fellowship from the University

of Chile and University of California Cooperative Program.

* Department of Biology, University of California, Los Angeles, California 90024. Present ad-

dress: Departamento de Biologia, Facultad de Ciencias, Universidad de Chile, Santiago, Chile.

2 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

tation; Juan Pulido and Jose Clavijo for field and laboratory assistance. | am in-

debted to Nobuko Kitamura for the preparation of the illustrations, to Caryle

Stallard for typing the preliminary draft and for editorial assistance. and Angeliki

Demos for the preparation of the text copy for lithoprinting.

MATERIAL AND METHODS

MATERIAL. All together 6,238 specimens were examined for this study, 1,340

males, 939 females, 2,675 larvae and 1,285 pupae; included among these were 958

individual rearings (540 larval, 282 pupal, 136 incomplete) of 14 of the 16 recog-

nized species. The bulk of this material was collected for the project “Mosquitoes

of Middle America’ (Belkin, Schick et al., 1965, 1967) and is deposited at the

University of California, Los Angeles [UCLA]. The only other major source of

material was the U.S. National Museum of Natural History [USNM]. Shared by

these institutions is a small number of important specimens from the collection of

W.H.W. Komp, which is unfortunately partially lacking in detailed data and has a

complicated numbering system (see Schick, 1970:4-5). A few valuable specimens

were borrowed from the Centro de Pesquisas Rene Rachou, Brazil [BH], the Divi-

sion de Endemias Rurales, Venezuela [MDM], and the Institut Pasteur, French

Guiana [PIG]. In July and August 1969, I collected additional material and made

field observations in northcentral Venezuela. I examined the types of babahoyensis,

bonnei, metempsytus, urichii [USNM] ; manaensis [PIG] ; and rausseoi [MDM]. J.N.

Belkin examined the types of anduzei, iridescens, soperi [FH]; and infoliatus [ITH].

I am designating here a lectotype for bihaicolus [USNM]. The types of the following

species were not seen: antunesi [BH] ; mathesoni [IHC]; guerreroi [MDM]; wilsoni

and secundus [LU].

TAXONOMIC PROCEDURES. I have followed the classical comparative morph-

ological taxonomic procedures as adapted to mosquito systematics by Belkin,

Schick et al. (1965:10; 1967:10-11). Species were first segregated on the basis

of a series of constant correlated morphological features in all stages, obtained

in most instances from the type localities in case of previously described forms.

They were then compared minutely for shared derived constant correlated fea-

tures to determine the primary phyletic lines and these were assembled into

groups, subgroups and complexes, on the basis of degree of similarity and di-

vergence (see chapter on Taxonomic Characters). In the final interpretation of

evolution of the subgenus, data from habitats and distribution were taken into

account.

DESCRIPTIONS. The terminology and abbreviations used in the descriptions

of the taxa in general follow Belkin (1962) except for a few modifications pro-

posed by Belkin (1968a:49), Belkin, Heinemann and Page (1970:78) and Knight

and Laffoon (1970a, 1970b), and a number of special terms which are explained

in the chapter on taxonomic characters and are labelled on the figures. Except

as stated for the adults of some species, all measurements were made on slide-

mounted specimens; further details and special problems in measurements are

given in the chapter on taxonomic characters. Descriptions are composite when-

ever more than 1 specimen was available. The method of presentation of data

in general follows that of Belkin (1962). For the chaetotaxy of the immature

stages the first figure(s) given following the hair number is the mode (frequency

of at least 75%), followed by the range in the sample (usually 5-10 specimens,

Valencia: Subgenus Carrollia of Culex 3

exact number specified in the description). The diagnostic features were check-

ed on all the available material; any departure from these is noted in the sec-

tions on Systematics under each species or group.

SYSTEMATICS. The diagnostic taxonomic features, probable affinities, indi-

vidual and geographic variation, and nomenclatural problems are discussed for

every species insofar as the available material permits. For groups and subgroups

this section is combined with those on bionomics and distribution in a Discus-

sion in which an attempt is made to synthesize the information from the indi-

vidual species and to determine the evolutionary trends in that group or sub-

group and its affinities with others. Much of this, of course, is speculation be-

cause of the paucity of data which may be subject to different interpretations.

BIONOMICS. The meager data on bionomics are derived primarily from the

record forms used in the project “Mosquitoes of Middle America” and a few

literature reports. They are largely restricted to data on the breeding sites of

the immature stages and biting records of females.

DISTRIBUTION. Only synoptic distributional data are given: country, major

political subdivision, locality, collection number (when available; if not available,

then collector’s name) and depository; reliable literature records are included.

Full data for every specimen examined are entered in a permanent ledger and

will be available upon request to interested parties on microfilm or other de-

sired reproduction method from the Department of Biology, University of Cali-

fornia, Los Angeles. On the maps, the probable distribution of each species is

plotted by circumscribing the entire area of individual spot localities. Overlap-

ping distributions of different species do not necessarily indicate complete sym-

patry as the species in question may occur at different elevations and in different

plant formations.

ILLUSTRATIONS. The full chaetotaxy of the immature stages of every spec-

ies is figured, with the modal hair branching determined as noted in the para-

graph on descriptions. For the adults pertinent details of the external morph-

ology of the female and head and claws of the male of a representative species

of the Bihaicolus Group, Urichii Subgroup and the Iridescens Subgroup are fig-

ured from a “typical” specimen, whenever possible from the type locality. The

male genitalia drawings of every species are usually based on several specimens,

from the type locality whenever available. The figures of the cibarial armature

of females are based usually on a limited number of specimens and may not be

entirely reliable.

TAXONOMIC CHARACTERS

The following morphological features of the adults, larvae and pupae proved

to be particularly useful for the diagnosis of species and as supraspecific char-

acters for the determination of their affinities. For the recognition of the phy-

letic lines and their interrelationships it is essential to establish which of the

states of each significant character is primitive and which is derived and whether

or not the derived state has occurred only once in the evolution of the group.

As noted by Adames (1971:3), because of the lack of paleontological evidence

in mosquitoes, reliance has to be placed for the identification of the primitive

state primarily on the trend within the taxon by comparison with related taxa

of the same rank. In this case the taxa involved are the other subgenera of Cu-

4 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

lex, primarily of the Melanoconion Section (see the chapter on Systematics).

In the following discussion I have indicated whenever possible the primitive state

of the various characters on this basis. An even greater problem is the deter-

mination of the monophyletic origin (true homology) of a derived state, which

is necessary for the recognition of true phyletic lines. The only criterion that

I could use with the available material was that of extreme similarity of the de-

rived states in the taxa being compared, as for example the position and de-

velopment of hair 9-C on the head capsule of the larvae of all species of the

Iridescens Group. When derived states of several characters in different stages

or different body parts of the same stage are correlated in this manner then

the monophyletic origin of the taxa involved is almost certain. Although close

correspondence of derived states of a single character appears to be reliable cri-

terion of true homology in most instances involving elaboration of new struc-

tures or shifts in position it is often useless in those dealing with reduction or

loss of structures.

ADULTS

HEAD. The most significant taxonomic features on the head are in the devel-

opment of the scales of the vertex.

Decumbent scales.—The trend is from a primitive narrow anterior band of dark

broad scales and numerous light narrow scales ‘to practically entirely light broad

scales. The color of the broad decumbent scales is quite variable and often dif-

ficult to determine because of iridescence; in many instances scales that appear

light from a dorsal aspect show a very dark blue iridescence from an anterior

view.

Erect scales._The trend is from very long slender ones extending forward to

near the orbital bristles to derived shorter, broader ones confined to the pos-

terior part of the vertex and occiput; the primitive color was probably also dark,

but is pale in most species.

Appendages.—The proboscis/forefemur ratio shows no consistent trend in the

subgenus; it was probably primitively near 1.0 in both sexes and shows an in-

crease in at least 2 separate phyletic lines. The labium primitively has no elon-

gate hairs except at the extreme base (basal bristles). The palpus/proboscis ratio

is fairly uniform throughout the subgenus in the females except for a marked

increase in kKompi and independent decrease in infoliatus and possibly iridescens;

there is a great deal more variation in the ratio in the males as can be expected

and this occurs in several independent lines. The antenna/proboscis ratio shows

a definite tendency towards an increase from the primitive condition of less than

1.0 in both sexes.

THORAX. There is a marked derived trend in the reduction in the mesono-

tal chaetotaxy from the presence of a few anterior acrostichals, a complete row

of anterior and posterior dorsocentrals, numerous prescutellars and some lateral

prescutals to the nearly complete absence of all of these. The primitive scala-

tion of the mesonotum is sparse and largely composed of nearly straight linear

dark scales; in the derived condition the scales are denser, broader and more

strongly curved and often paler in the posterior part. The scutellar scales on the

midlobe and lateral lobes are primitively linear, and narrow curved to broad in

the derived condition. The pleuron exhibits several obviously derived features:

Valencia: Subgenus Carrollia of Culex 5

(1) darkening and development of a broad pruinose stripe; (2) reduction in the

upper stp bristles; and (3) development of broad scales on apn and lower stp.

LEGS. The legs show relatively few significant taxonomic characters other than

specific differences in the extent of the light basal scaling of the femora. The

metallic spots of the femora show a great deal of individual variation and rela-

tively few specific differences in location, size and intensity of iridescence; on-

ly in the Urichii Subgroup is there a marked derived condition in the develop-

ment of a metallic spot on the forefemur. The large field of plantar setae on

segment 5 and a smaller on segment 4 of the foretarsus and midtarsus of the

Bihaicolus Group is probably a primitive condition. The loss of the tooth on

the larger claw of the midtarsus of males of the Iridescens Subgroup appears

to be a derived state.

WING. The only significant taxonomic character noted on the wing is the

type of plume scales on Rs, R243, R2, R3 and M. In the primitive condition

these scales are long and linear and occur on all these veins; in the derived con-

dition they are shorter and broader and are more restricted in distribution, be-

ing replaced by scales hardly differentiated from squame scales on R, and R3.

ABDOMEN. There appear to be few significant taxonomic characters on the

pregenital segments of the abdomen. The following character states I have in-

terpreted as being primitive: (1) abdomen subcylindrical, not distinctly compress-

ed; (2) dorsal part of tergite I with few scales; (3) basal tergal light bands not

developed; and (4) distal sternites without dark apical bands. The shape and po-

sition of the lateral metallic markings may show group differences but I did not

study these in detail. There are also distinct specific differences in the size of

these markings on the proximal segments; these are obscured to some extent

by considerable individual variation.

CIBARIAL ARMATURE. The female cibarial armature was not studied in suf-

ficient detail and in a sufficient number of specimens to determine whether or

not reliable taxonomic characters are present. However, it appears that there are

at least good specific differences. I am proposing the term cibarial dome for

the hemispherical internal projection from the anterior (dorsal) wall of the ci-

barium just proximad of its junction with the pharynx. It protrudes into the prox-

imal part of the pharynx (fig. 9). This structure is found throughout the Melanoco-

nion Section of Culex as well as in the genus Deinocerites.

FEMALE GENITALIA. The only group difference noted in the female gen-

italia was the presence of an atrial plate in the Bihaicolus Group; presumably

this is the primitive condition in the subgenus. There appear to be specific dif-

ferences in the number of setae on tergite IX, cercus, postgenital plate and in-

sula but the sample studied was insufficient to evaluate the significance of these

differences.

MALE GENITALIA. Previously described species have been characterized princ-

ipally on male genitalic characters without adequate analysis of the considerable

individual and population variation or much regard to the homology of the struc-

tures described. I have attempted to determine these homologies and for des-

criptive purposes have proposed several new terms as indicated below.

Sidepiece.—In the primitive state the sidepiece is elongate, more or less coni-

cal and without scales; the development of the short ovoid shape and of the

scales may have been derived independently in several lines.

Subapical lobe.—Characteristic of the subapical lobe in the subgenus are 2 de-

rived trends: (1) reduction and loss of the distal division probably independently

6 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

in several lines; and (2) development of an accessory division at or near the base

of the proximal division, possibly more than once. In the Bihaicolus Group the dis-

tal division is represented by a reduced lobe adjacent to the proximal division

(fig. 10) or is absent; in the Urichii Subgroup it is small and widely separated

from the proximal division (figs. 20,22); and in the Iridescens Subgroup it is

represented distad of the proximal division by a small bare proturberance (figs.

24,28), except in secundus in which it is apparently completely absent (fig. 26).

The proximal division normally bears 2 apical specialized setae, a and b (fig.

10); occasionally a short subapical seta is more or less differentiated, it is arbi-

trally designated seta c although it may not be homologous with seta c on the

proximal division in the subgenus Culex. The accessory division is developed on-

ly in the Iridescens Subgroup (see) and may have arisen independently in each

of the complexes of the group from different parts on or near the base of the

proximal division.

Clasper.—Characteristic of Carrollia is the elaboration of the clasper. In the

primitive state (Bihaicolus Group) it is simple, parallel-sided or slightly tapered

apically. As in some groups of the Melanoconion Section of the genus it has:

(1) a snout, an external apical recurved process associated with the spiniform;

(2) a ridged crest, externally before the beak; and (3) in the distal part 3 setae

which are here designated from the apex basad as seta a (the usual thickened

spiniform) and the simple hairlike setae b and c (fig. 10). The derived states

are: (1) the development of a distal head with an outer curvature and inner pre-

apical lobe (fig. 26); (2) the modification of the crest to a flat sclerite; (3) the

thickening or flattening of seta b which now may appear as the spiniform; and

(4) the reduction of seta a to a hairlike state.

Phallosome.—The lateral. plate of the aedeagus is of the usual Melanoconion

type with a so-called basal hook. The primitive type of lateral plate appears to

be broad, with a distinct sternal spine, a conspicuous lateral external spine and a

simple apex (fig. 10). The derived conditions are: (1) narrowing of the plate;

(2) loss of the external spine; (3) loss of the sternal spine; and (4) development

of a long caudal process from the apex (fig. 26).

Proctiger.—_The proctiger is also of the usual Melanoconion type. The primi-

tive states are probably as follows: (1) the basal part broad and distal elongate;

(2) paraproct sclerotization without basal mesal sternal lobe; (3) crown of para-

proct with several flattened teeth; and (4) several cercal setae. The only phylo-

genetically significant derivation is the development of a mesal sternal lobe (fig.

26) at the base of the paraproct sclerotization. All the other derived states in-

volve reduction of crown teeth, reduction or loss of cercal setae and shorten-

ing or broadening of the distal part of the paraproct and have occurred inde-

pendently in several lines.

PUPAE

The pupae of the subgenus are remarkably uniform, with only a few phylo-

genetic characters that are correlated with those of the adults and larvae. Al-

though there are some differences in the chaetotaxy of most species the indi-

vidual and sometimes geographical variations are so great that they are difficult

to use for diagnosis. In the descriptions I have given data on the mode and range

of branching for all the cephalothoracic and metanotal hairs and for selected

Valencia: Subgenus Carrollia of Culex e

abdominal hairs which may prove to be useful for future work with larger sam-

ples than available for this study.

CHAETOTAXY. The most significant phylogenetic characters in the pupal chae-

totaxy are the positions of hair 2 on abdominal segment II and hair 3 on seg-

ment VI, and the development of cephalothoracic hair 5. The primitive position

of hair 2-II laterad of 3-II in nearly the entire tribe Culicini is preserved in the

Bihaicolus Group; the derived position mesad of 3-II is found in all members

of the Iridescens Group. The position of hair 3-VI is extremely varied in the

Culicini but appears to be primitively laterad of hair 1 in the Melanoconion Sec-

tion of Culex; this position is found in the Iridescens Group, while in the Bi-

haicolus Group 3-VI is mesad of hair 1. Cephalothoracic hair 5-C is elongated

in the Bihaicolus Group and the Urichii Subgroup while it is short in the Iri-

descens Subgroup; the latter is probably the derived condition in the subgenus.

The only other hairs that appear to show significant phylogenetic differences are

abdominal hairs 3-II,III, 1-IJI and 5-VI. Specific differences have been noted pri-

marily in the development of abdominal hairs 1-IV, 3-IV, 4-VI, 5-IV,V, 6-III-

V and 9-VII, VIII.

TRUMPET. As in most other Culicini a short broad flared trumpet with a

relatively large pinna is the primitive condition in Carrollia, retained in the Bi-

haicolus Group and to a lesser extent in the Urichii Subgroup. In the Iridescens

Subgroup the derived states involve elongation and narrowing of the trumpet and

reduction of the size of the pinna.

PADDLE. The paddle in Carrollia presents no significant taxonomic charact-

ers other than changes in shape. There is a tendency for the loss of hair 2-P

independently in several lines and even the reduction of 1-P.

FOURTH INSTAR LARVAE

In the larvae of Carrollia there are many excellent constant phylogenetic char-

acters that are concordant with similar characters of the adults and to a lesser

extent of the pupae. There are also good specific differences but the hair branch-

ing often shows considerable individual as well as population variation. In the

descriptions I have given data on the mode and range of branching for all ceph-

alic and thoracic hairs and for selected abdominal hairs. Further study of these in

larger samples may reveal additional reliable specific differences.

HEAD. A striking feature of Carroillia is the anterior position of cephalic hairs

5,6-C which is probably the primitive condition in the Melanoconion Section of

the genus but may also be related to the shape of the head capsule and the con-

fined habitat utilized for breeding sites. Another unusual primitive feature is the

retention and strong development of hair 2-C in the entire subgenus, known else-

where in the tribe only in the genus Deinocerites. These 2 groups also share the

loss of hair 3-C, rarely represented by a spicule. The primitive normal position

of hair 9-C is retained in the Bihaicolus Group but this hair is removed to near

the collar in all members of the Iridescens Group, a unique feature in the tribe

Culicini. There is also a definite derived trend for the caudal migration of hairs

12,14-C in the Iridescens Subgroup. Characteristic of the subgenus is the derived

condition of hair 11-C which is relatively poorly developed and located farther

ventrad from the base of the antenna than in any other group in the tribe. Spe-

cific differences are most evident in the development of hairs 5-7-C. The anten-

8 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

na is very uniform throughout the subgenus and is of the primitive type with-

out differentiation of a wider basal part. Maxillary hair 5-MP of all members of

the Iridescens Group is greatly thickened, obviously a derived condition from

the simple seta of the Bihaicolus Group.

THORAX. The taxonomic characters of the thorax are primarily useful at the

specific level. The following derived states are evident however at the subgroup

level: (1) in the Iridescens Subgroup, spicules on the prothorax, enlargement and

substellate to stellate development of hairs O-P, 14-M and 13-T; and (2) in the

Urichii Subgroup, thickening and enlargement of hairs 8-P and 13-T.

ABDOMEN. There appear to be no significant major group characters except

for the following hairs (derived states): (1) in the Iridescens Group, dendritic

hair 11-I, short dendritic hair 3-VI; and (2) in the Iridescens Subgroup, enlarge-

ment of hair 1-IJ. The conspicuous enlargement of hair 1-VII is found also in

some other container-breeding groups of the Melanoconion Section of Culex and

in Deinocerites. The strong development of hair 6 on segments III-VI is prob-

ably a primitive character as it occurs in some groups of the Melanoconion Sec-

tion, in the subgenus Culex and in Deinocerites. The most significant characters

at the level of complexes and species were found to be in the development and

branching of hairs 7-II, 6-III-V, 1-III-V, 3-IJ-V, and 13-III-V.

SEGMENT VIII. The type and arrangement of comb scales divides the sub-

genus into the 2 principal groups. The single row of long spinelike scales is un-

doubtedly a derived condition as the patch of short apically fringed scales is

characteristic of Deinocerites and all primitive forms in the other subgenera of

Culex.

SIPHON. A derived character of the Urichii Subgroup is the presence of elon-

gate spicules on the siphon. The siphon index is useful for the diagnosis of sev-

eral species but shows considerable individual and geographical variation in many

species. In the ventral siphonal tufts there is a derived trend in the reduction

in the number of hairs and in the number of branches in individual hairs; the

primitive condition is retained in the Urichii Subgroup. The shape of the pec-

ten teeth is varied but shows no consistent group characters; there is a great

deal of individual variation in the number of pecten teeth in most species.

ANAL SEGMENT. A very important group character, as in other mosquito

larvae, is the development of the ventral brush. The primitive condition of a

large number of hairs, and these with numerous branches, is retained in the Urichii

Subgroup. In the Bihaicolus Group the number of hairs is large but their branch-

ing is reduced. In the Iridescens Subgroup there is a reduction in the number

of hairs but relatively little in their branching. The primitive state of hair 1-X

is probably an unbranched condition and that of 2-X a large multiple fan-shaped

hair; both of these are a characteristic condition in the Urichii Subgroup; the

derived branching of 1-X and reduction of 2-X are found in the Bihaicolus Group

and the Iridescens Subgroup. There are no significant taxonomic characters on

the saddle except in guerreroi. The gills are quite varied in the different species

but there is no detectable phylogenetic trend.

SYSTEMATICS

TAXONOMIC HISTORY. Although the taxonomic history of Carrollia is rather

simple it is necessary to summarize it here as no previous revision of the group

Valencia: Subgenus Carrollia of Culex 9

has been made. Adolpho Lutz (1905:81-82) described Carrollia as a distinct genus

with iridescens as the only included species. The combination Carollia irides-

cens appeared earlier as a nomen nudum (Lutz, 1904:16) and this was repeated

by Blanchard (1905:634). Coquillett (1906a:61) described urichii in the genus

Melanoconion and retained it in this genus in his classification of the mosqut-

toes of North and Middle America (Coquillett, 1906b:23). Dyar and Knab (1906:

223) and Dyar (1906:18) transferred urichii to Mochlostyrax on the basis of

larval characters. Subsequently Dyar and Knab (1909:101) recognized the affin-

ities of urichii and transferred it to Carrollia which they treated as a full genus

following Peryassu (1908: 167-169,343) who had redescribed the adults and pro-

vided a brief description and figure of the larva of Carrollia iridescens.

Considerable confusion occurred during this early period of very intensive and

competitive taxonomic studies on mosquitoes. Unaware of Lutz’s paper, Theobald

(1907:206-209) redescribed the genus Carrollia and C. iridescens, but these re-

descriptions should not be considered as proposals of new taxa (see Belkin, 1971:

42). Theobald’s description of the scutellar scaling, repeated in a later key (1910:

113), was midleading and he probably had at least 2 species included under iri-

descens. Theobald (1910:455,458) did not recognize the affinity of urichii but

questioned its inclusion in Melanoconion, suggesting that it might be more prop-

erly placed in Janthinosoma. At about the same time further confusion arose

with the report of Culex (Carrollia) iridescens from the Canal Zone by Busck

(1908:70). The identification, made by Dyar and Knab (Busck, 1908:50), was

erroneous and actually 2 other species, described later, were involved. This mis-

identification persisted in Howard, Dyar and Knab (1915:461-466) who treated

Carrollia as a genus, redefined it and provided the first keys to the adults, male

genitalia and larvae of urichii and ‘‘iridescens’’ together with detailed descrip-

tions of the species including figures of the male genitalia and larvae. Dyar (1918:

108) reduced Carrollia to a subgenus of Culex and suggested that urichii might

be placed in a separate subgenus but he did not recognize the misidentification

of iridescens from Panama.

The Bonnes were the first investigators to make significant additions to the

knowledge of the subgenus. They proposed the name secundus for the misiden-

tified “‘iridescens’’ from Panama and described infoliatus but retained incorrectly

“iridescens’’ for the Surinam species (Bonne-Wepster and Bonne, 1920:170-171).

Dyar quickly followed with the description of metempsytus from Costa Rica and

proposed bonnei for the ‘“‘iridescens’’ from Surinam (Dyar, 1921:154155). The

latter was not accepted by Bonne and Bonne-Wepster (1925:196-207), who re-

viewed all the previously described species. The unjustified replacement name

Carrollella of Lutz (1921:161) was not noted either by the Bonnes or by Dyar

(1925:176) who incorrectly placed amazonensis Lutz, 1905 in the subgenus.

Dyar and Nunez Tovar described bihaicolus as a new species in 2 separate pub-

lications (1927:4-5, Spanish; 1928:91, English) just prior to the appearance of

Dyar’s monograph on the mosquitoes of the Americas (Dyar, 1928). In the latter

work Dyar accepted Carrollella as a replacement for the subgeneric name Carrollia,

removed amazonensis from the subgenus and considered all the previously described

7 nominal species to be valid taxonomically except bonnei which he retained as a

synonym of iridescens following Bonne and Bonne-Wepster (1925). Dyar’s (1928:

270) unfortunate error in indicating the presence of only narrow curved scales on

the vertex of the head of Carrollella was followed by Edwards (1932:220), who

also accepted Dyar’s taxonomic treatment but reverted to the use of Carrollia, and

10 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

also later by Lane (1953:496), probably directly from Edwards. The keys in Dyar’s

monograph provided little help in the identification of species.

The most recent comprehensive treatment of Carrollia, by Lane (1953:496-

514), was very superficial as it did not include a description of the adult morph-

ology of even 1 species and provided a more or less workable key to male gen-

italia only. All the 12 nominal species described to that date were treated as

taxonomically valid, including in addition to those previously mentioned in this

review: anduzei Cerqueira & Lane, 1944; antunesi Lane & Whitman, 1943; math-

soni Anduze, 1942; soperi Antunes & Lane, 1937; and wilsoni Lane & Whitman,

1943. The diagnoses of these additional species by the original authors were based

almost entirely on male genitalia except for antunesi and soperi. Other than the

original descriptions, the only significant taxonomic contributions during this per-

iod were those of Antunes and Ramos (1939) on the iridescens complex, with

a designation of the neotype of iridescens and the resurrection of bonnei; Sene-

vet and Abonnenc (1939; 1958) on the species from French Guiana; a few distribu-

tional records listed under the individual species in the taxonomic treatment; and an

unwarranted synonymy of infoliatus with bihaicolus by Komp (1936:64; 1956:39).

Subsequent to Lane’s review, 2 additional species have been described, baba-

hoyensis Levi-Castillo, 1953 and manaensis Floch & Fauran, 1955. Stone and

Knight (1957) designated lectotypes for bonnei and metempsytus. In the world

catalog (Stone, Knight and Starcke, 1959:282-283) all 14 nominal species were

considered to be taxonomically valid. Recent papers dealing with Carrollia are

those of Casal and Garcia (1968) reporting soperi from Argentina and indicat-

ing several characters useful in the separation of females; Belkin (1968; 1971)

on the type specimens and designation of lectotypes; the description of 2 new

species from Venezuela, guerreroi and rausseoi, by Cova Garcia, Sutil and Pulido

(1971; 1972); and an adequate redescription of anduzei and a confused one of

secundus from Brazil by Barata and Cotrim (1972).

PROPOSED CLASSIFICATION. The species recognized in this revision fall in-

to 2 clearly marked major groups on at least 2 correlated characters each in ex-

ternal adult features, male genitalia, pupal chaetotaxy and larval features. Only

generalities on the groups and their components are briefly reviewed here; de-

tails are presented in the discussions of the groups, subgroups. and individual

species.

The Bihaicolus Group of 5 named species (bihaicolus, guerreroi, rausseoi, me-

tempsytus, infoliatus) and the unnamed sp. 3a from Panama has undoubtedly

retained more primitive features in all stages and is closer to the ancestral stock

of the subgenus. It is characterized primarily by these primitive features: (1) in

the adults, strongly developed dorsocentral bristles; pleuron pale; (2) in the male

genitalia, lateral plate of phallosome with subapical external spine and not pro-

duced caudad; clasper simple; (3) in the pupae, abdominal hair 2-II laterad of

3-II; hair 3-VI mesad of 1-VI; and (4) in the larvae, head hair 9-C in normal po-

sition; comb scales apically fringed and in a patch. The group is so compact that

it is not divided into subgroups. However, 2 phyletic lines are discernible largely

on differences in the development of the subapical lobe of the male genitalia. The

Infoliatus Complex (infoliatus, metempsytus) has lost the distal division of the

subapical lobe which is retained in the Bihaicolus Complex (bihaicolus, guerreroi,

rausseoi). Species 3a is known in the larva only and cannot be placed in either

complex at this time. The species are poorly differentiated except in male genitalia.

On general adult features, only infoliatus is readily separated from the others. The

Valencia: Subgenus Carrollia of Culex 11

pupae are extremely similar and the diagnostic characters of the larvae are rather

tenuous. Two species (bihaicolus, infoliatus) are widespread dominant modern

forms; metempsytus is moderately common in a more restricted area; and guerreroi,

rausseoi and probably sp. 3a are relict forms.

The Iridescens Group of 11 named species, 2 unnamed forms and possibly 3

additional forms is characterized primarily by derived features: (1) in the adults,

dorsocentral bristles not developed on mesonotal disc; pleuron with pruinose stripe;

(2) in the male genitalia, lateral plate of phallosome without subapical external

spine and with a long caudal process; clasper complex; (3) in the pupae, abdominal

hair 2-II mesad of 3-II; hair 3-VI laterad of 1-VI; and (4) in the larvae, head hair

9-C removed caudad; comb scales elongate, spinelike and in a single row.

The Iridescens Group is divided into 2 very distinct major phyletic lines. The

small Urichii Subgroup of 2 poorly differentiated species, the dominant wide-

spread urichii and the restricted anduzei, is characterized by a combination of

derived and primitive features: (1) in the adults, upper sternopleurals present;

hindtarsal segment 4 with a broad basal white ring; (2) in the male genitalia,

distal division of the subapical lobe present; caudal process of lateral plate of

phallosome curved ventrad; seta b of clasper simple; (3) in the pupae, cephalo-

thoracic hair 5-C strongly developed; and (4) in the larvae, ventral brush with

at least.5 pairs of hairs; prothoracic hair 8-P spikelike.

The Iridescens Subgroup is the largest and most diverse major phyletic line

in the subgenus. It is characterized primarily by derived features in all stages:

(1) in the adults, upper sternopleurals absent; hindtarsal segments entirely dark

scaled; (2) in the male genitalia, distal division of the subapical lobe represented

by only a knoblike protuberance or completely absent; caudal process of lateral

plate of phallosome straight or curved laterad; (3) in the pupae, cephalothoracic

hair 5-C weak; and (4) in the larvae, ventral brush usually with only 4 pairs of

hairs; prothoracic hair 8-P weak or stellate. Several phyletic lines are evident with-

in this subgroup but their relationships are obscure. The derived male genitalic

features characterizing these different lines are not concordant with the mosaic

of primitive and derived features of the pupae and larvae. For the present, I am

dividing the subgroup into 2 complexes. The Iridescens Complex (bonnei, sec-

undus, iridescens, sp. 8a, sp. 9a) consists of a single phyletic line that is clearly

marked by the strongly differentiated preapical lobe on the clasper of the male

genitalia and the presence of erect scales on anterior part of the vertex of the

head of the adults. The Antunesi Complex lacks these features and consists of

4 apparently separate phyletic lines: Soperi, Babahoyensis and Kompi, each with

a single species, and Antunesi with at least 3 species (antunesi, cerqueirai, wil-

soni) and possibly 3 additional forms.

DISTRIBUTION (figs. 2-5). The subgenus Carrollia is precinctive to the New

World tropics. Its northernmost record is from the State of Vera Cruz in south-

ern Mexico and its southern limit appears to be in the States of Sao Paulo and

Parana in Brazil and the Province of Missiones in northeastern Argentina. The

subgenus is not known from the Caribbean islands and on the Pacific coast of

South America it has not been reported south of the Guayas River Basin in

Ecuador. The distribution is not continuous through the range but is limited to

areas of tropical, subtropical and cloud forests from near sea level to as high

as 2,000 meters.

The species of Carrollia are concentrated on the northern margins of South

America where all the groups, subgroups and complexes are represented by 14

12 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

named species and by sp. 3a. As noted under the discussion of the systematics

of antunesi and wilsoni, their populations in this area may not be conspecific

with topotypic southern populations and may be represented by 2 or 3 unnamed

forms. In the vast Amazon Basin only 4 species are known, infoliatus of the

Bihaicolus Group, urichii and anduzei of the Urichii Subgroup and bonnei of

the Iridescens Complex, but it is probable that 1 or more members of the An-

tunesi Complex remain undiscovered here. South of the Amazon Basin the Bi-

haicolus Group and the Urichii Subgroup are absent and only 6 species or forms

have been reported, all apparently derived precinctives, 3 in the Iridescens Com-

plex (iridescens, sp. 8a, sp. 9a) and 3 in the Antunesi Complex (soperi, antunesi,

wilsoni).

AFFINITIES. Carrollia has all the characteristics diagnostic of the genus Culex

as currently interpreted: (1) in the adults, pulvilli developed; female genitalia with

a short cercus; sidepiece of the male genitalia with a subapical lobe and para-

proct with a crown of teeth; (2) in the pupae, hair 9-VIII sternal and removed

from caudolateral angle; and (3) in the larvae, siphon with numerous accessory

subventral tufts; ventral brush of anal segment strongly developed.

The relationship to the Melanoconion Section of Culex (subgenera Aedinus,

Anoedioporpa, Melanoconion, Micraedes, Microculex, Tinolestes) is evident in the

following features: (1) in the adults, vertex of head with at least an anterior

border of broad scales; phallosome of the male genitalia with a simple “hooked”

lateral plate; female with a cibarial dome; (2) in the larvae, siphon with distal

accessory hairs reduced in size. The pupae of this section are not known suffic-

iently to distinguish them from other sections of the genus.

Carrollia shows some affinities also with other genera of the tribe Culicini.

With Deinocerites it shares the retention of head hair 2-C and the similar dif-

ferentiation of the distal accessory hairs of the siphon of the larva; and the pres-

ence of the cibarial dome in the female. The cibarial dome is also differentiated

in Galindomyia, currently considered to be a distinct genus but probably merely

an aberrant subgenus of Culex of the Melanoconion Section. Because of these

annectent features, Carrollia appears to be an offshoot from a very primitive stock

of the genus Culex. |

Carrollia is differentiated from the other subgenera of Culex: (1) in the adults

by the development of metallic spots on the femora of the legs and on the ab-

dominal tergites, (2) in the larvae by the retention of hair 2-C on the head. As

noted above, the pupae of other subgenera of the Melanoconion Section are too

poorly known to establish distinguishing characters.

EVOLUTION. Although our present knowledge of the systematics, distribu-

tion, bionomics and affinities of Carrollia is fragmentary some general aspects

of the evolution of the subgenus may be discerned.

The subgenus probably originated from a stock of the Melanoconion Section

of Culex which possessed many primitive features, some apparently annectent

with the genus Deinocerites. The place of origin was undoubtedly somewhere in

the northern margin of the present South American continent, possibly at the

northern end of the Andean Cordilleras, where all the groups, subgroups and

complexes are presently found and where several relicts are known. Subsequent

radiation appears to have occurred largely in the same general area.

The first division was undoubtedly into the Bihaicolus Group stock which re-

tained primitive character states and the Iridescens Group stock with derived fea-

tures. The Bihaicolus Group stock probably occupied the west end of the distribu-

Valencia: Subgenus Carrollia of Culex 13

tion of the subgenus at that time and the Iridescens Group stock the eastern part.

The Iridescens Group was then divided into the Urichii Subgroup stock, probably

also in the west, and the Iridescens Subgroup stock. The splitting of the Bihaicolus

Group into the 2 complexes may have taken place at the same time. All these events

were probably coincidental with major geologic changes in this unstable marginal

continental area.

Subsequent radiation and speciation in the Bihaicolus Group (fig. 2) took place

largely in the northwest corner of South America with a major invasion of Central

America and only 1 penetration southward into the Guiana Shield and the Ama-

zon Basin. The Urichii Subgroup (fig. 3), which retained a number of primitive

features, expanded its range into the Guiana Shield and the Amazon Basin with

only 1 derivative. The evolution of the highly derived Iridescens Subgroup is not

at all clear but it appears that the subgroup was fragmented into several phyletic

lines, again in northern South America. Three of these lines, the Iridescens Com-

plex and the Soperi and Antunesi lines of the Antunesi Complex spread into

South America south of the Amazon Basin.

Speciation in Carrollia appears to have occurred primarily by allopatric differ-

entiation of populations. However, there are suggestions of introgression involved

in the formation of some species, sp. 3a, sp. 9a, soperi, cerqueirai and possibly

kompi. The obligate association of Carrollia with humid forests has played a ma-

jor part in the speciation and present distribution of the subgenus. These forests

are now widely disjunct except in the Amazon Basin and their geographical and

altitudinal distributions have undoubtedly varied in the past.

BIONOMICS

Species of Carrollia are silvan mosquitoes, restricted to humid areas of tropical,

subtropical and cloud forests. The highly ornamented adults are diurnal in activ-

ity but have been seldom collected in nature as they are apparently not attracted

to humans or to the baits used in traps. The immature stages are found in various

types of container habitats, natural and artificial, but have been found occasionally

in water on the ground, probably containing decaying leaves or other organic

matter.

ADULTS. Practically nothing is known of the blood-feeding habits of females

of Carrollia. The only record of feeding on humans is that of Levi-Castillo (1953c:

163) for babahoyensis. There are a few reports of collections of adults, presum-

ably females, on humans or in animal-baited traps: infoliatus (Cerqueira, 1961:

136); urichii (Komp, 1936:64; Kumm and Novis, 1938:503, agouti baited; Cer-

queira, 1961:136) and soperi (Antunes and Lane, 1937:22). A few species have

also been found in unbaited traps, by sweeping, or collected in an unspecified

manner as adults: bihaicolus, infoliatus (Antunes, 1937:79); urichii; anduzei (Barata

and Cotrim, 1972:30); bonnei; soperi (Casal and Garcia, 1968:97).

IMMATURE STAGES. The recorded natural breeding sites of Carrollia include

flower bracts of Heliconia and leaf axils of Dieffenbachia (both only for bihaicolus),

leaf axils of bromeliads (for iridescens only, Forattini, 1965:34); fallen leaves

and spathes and fronds of palms (very frequently), fallen nuts and fruits (occasion-

ally), treeholes and cut or broken bamboo (very frequently). Several species, es-

pecially dominant widespread ones, are often found in artificial containers of all

types, wooden, metal, glass, rubber and even concrete and leather, and have also

14 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

been recovered from bamboo or wooden traps. There are a few records of breed-

ing in ground waters: urichii (rockholes), secundus (ground seepage), iridescens

(ground pools). The dominant species utilize a wide variety of habitats while the

relict ones appear to be confined to only 1 or 2 natural types but are also some-

times found in artificial containers (guerreroi).

It is not uncommon to find several sympatric species of Carrollia breeding in

the same habitat. These are usually members of different groups or subgroups

but occasionally one finds members of the same complex together. Associated

with species of Carrollia are many species of other groups of container breeders.

At least 41 such associates have been recorded, representing nearly every sym-

patric group of container breeders: Aedes (Howardina) ioliota Dyar & Knab, 1913,

A. (Ochlerotatus) fluviatilis (Lutz, 1904), A. (Protomacleaya) daryi Schick, 1970,

A. (P.) homoeopus Dyar, 1922, A. (P.) impostor Schick, 1970, A. (P.) insolitus

(Coquillett, 1906), A. (P.) podographicus Dyar & Knab, 1906, A. (P.) terrens

(Walker, 1856); Anopheles (A.) eiseni Coquillett, 1902, An. (Kerteszia) neivai

Howard, Dyar & Knab, 1913; Culex (Anoedioporpa) conservator Dyar & Knab,

1906, C. (An.) sp.; Culex (C.) chidesteri Dyar, 1921, C. (C.) corniger Theobald,

1903, C. (C.) mollis Dyar & Knab, 1906, C. (C.) quinquefasciatus Say, 1823,

C. (Lutzia) sp., C. (Melanoconion) sp., C. (Microculex) kukenan Anduze, 1942,

C. (Microc.) stonei Lane & Whitman, 1943; Haemagogus (H.) iridicolor Dyar,

1921, H. (Stegoconops) equinus Theobald, 1903, H. (S.) soperi Levi-Castillo 1955,

H. (S.) spegazzinii falco Kumm, Osorno-Mesa & Boshell Manrique, 1946; Limatus

asulleptus (Theobald, 1903), L. durhamii Theobald, 1901; Orthopodomyia albi-

costa (Lutz, 1904), O. fascipes (Coquillett, 1905); Sabethes (Sabethinus) undosus

(Coquillett, 1906), Sabethes spp.; Toxorhynchites (Lynchiella) spp.; Trichopro-

sopon (Ctenogoeldia) magnum (Theobald, 1905), 7. (Runchomyia) ulopus (Dyar

& Knab, 1906), T. (T.) digitatum (Rondani, 1848), T. (T.) pallidiventer (Lutz,

1905); Wyeomyia aporonoma Dyar & Knab, 1906, W. arthrostigma (Lutz, 1905),

W. complosa Dyar, 1928, W. felicia (Dyar & Nunez Tovar, 1927), Wyeomyia spp.;

Corethrella spp.

TAXONOMIC TREATMENT

Subgenus CARROELIA Lutz

1904. Carollia Lutz, 1904:16. Nomen nudum.

1905. Carrollia Lutz, 1905:81-82. TYPE SPECIES: Carrollia iridescens Lutz, 1905, Brazil;

monobasic.

1921. Carrollella Lutz, 1921:163. TYPE SPECIES: Carrollia iridescens Lutz, 1905, Brazil.

Unjustified emendation of Carrollia Lutz, 1905 believed to be preoccupied by Carollia

Gray, 1838, Carolia Cantraine, 1838 and Carolia Gray, 1867.

Culex (Carrollia) of Dyar (1918:108; 1925:176); Bonne-Wepster and Bonne (1920:170-171);

Bonne and Bonne-Wepster: (1925:196-207); Edwards (1932:220-221); Antunes and Ramos

(1939:376-385); Lane (1939:81-83; 1953:496-514); Senevet and Abonnenc (1939:123-129;

1958:287-293); Lane and Whitman (1943:389-397); Cerqueira and Lane (1944:216-220);

Stone and Knight (1957:44,53,58-59); Stone, Knight and Starcke (1959:282-283); Cerqueira

(1961:136); Fauran (1961:44); Cova Garcia, Sutil and Rausseo (1966a:28,111-117; 1966b:

42-4484 ,141,231-235 343-344); Casal and Garcia (1968:97-98); Barata and Cotrim (1972:

17-30).

Valencia: Subgenus Carrollia of Culex 15

Culex (Carrollella) of Dyar (1928:280-285); Komp (1936:64; 1956:39); Antunes (1937:78).

Carrollia of Theobald (1907:206-209; 1910:113); Peryassu (1908:34,44,167,169,343); Dyar and

Knab (1909:101); Surcouf and Gonzales-Rincones (1911:144); Howard, Dyar and Knab

(1915:461-466); Townsend (1934:489-490).

Carrollella of Martini (1931:217); Shannon (1931a:8; 1931b:137).

Carollia of Blanchard (1905:634).

Mochlostyrax in part of Dyar and Knab (1906:223).

Melanoconion in part of Coquillett (1906:23).

FEMALES. Small to medium-sized; mesonotum light brown to dark brown

or black; pleuron pale to dark; midfemur with 2 pale metallic spots, hindfemur

and sometimes forefemur with 1 similar spot; tarsi all dark or hindtarsal segment

4 with broad basal pale ring (Urichii Subgroup); pregenital abdominal segments

subcylindrical (Bihaicolus Group) or distinctly compressed, tergites II-VI] with

pale metallic lateral spots. Abdominal pale metallic spots basically silvery white

with varying intensity of violet reflections; femoral markings silvery white to

golden with similar iridescence. Dark scales of head, labium, palpus, legs and

abdomen almost always with strong deep blue, greenish blue or purple metallic

iridescence. Head: Eyes contiguous above antennal bases, interocular space not

developed. Decumbent scales of vertex broad on an anterior transverse band of

varying width, interrupted on midline in Urichii Subgroup; narrow decumbent

scales in the middle posteriorly on vertex and on occiput or absent; no indica-

tion of a frontal tuft, rarely a pair of setae below interorbital bristles; orbital

line without scales; a patch of broad decumbent light scales laterally continuous

with band of broad decumbent scales of vertex. Erect scales forked, elongate;

numerous to relatively few; extending to orbital bristles or confined to poster-

ior part of vertex and occiput. One pair of strongly developed interorbital (fron-

tal) bristles and a continuous row of shorter orbitals; a few very short hairs or

hairlike scales sometimes present caudad of orbitals. Clypeus prominent, bare.

Proboscis slender, distinctly shorter than abdomen; subequal to forefemur to dis-

tinctly longer; labium slightly dilated and flattened apically, dark scaled, some-

times with pale scales ventrally and rarely also dorsally; labellum small, with

scales on basal part. Palpus usually short, 0.16-0.25 of proboscis, rarely nearly

0.5 (kompi); usually 3-segmented, sometimes with a minute apical 4th segment,

palpifer not differentiated from basal segment; segment 3 elongate; all 3 seg-

ments with dark scales. Antenna subequal to proboscis or distinctly longer; scape

poorly differentiated; torus (pedicel) normal, usually with 3-14 short setae on

anterior mesal surface; flagellar segment 1 less than 1.5 of segment 2, with or

without distinct submedian whorl of moderately long bristles; segments 2-13 sub-

equal, with distinct basal whorl of 8-10 long bristles, progressively shorter on

distal segments. Thorax: Mesonotal integument shiny, pigmentation varied from

black to light brown; vestiture of linear to narrow curved scales varied from sparse

to dense, usually dark but sometimes predominantly pale. Prescutellar space bare

in distal half or less. Scutellar scales linear to broad, varied in color. Paratergite

bare. Acrostichals usually absent on disc, a few short ones caudad of anterior

promontory usually present, rarely (infoliatus) extending to level of scutal angle

or even beyond but never reaching prescutellar space; dorsocentrals present (Bi-

haicolus Group) or absent on disc; humerals present; prescutals (fossals) present

(Bihaicolus Group) or absent; prescutellars nearly always present; antealars and

supraalars always present; scutellars strongly developed; 1 parascutellar bristle.

Pleural integument pale, greenish or whitish, and dull (Bihaicolus Group), or

16 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

darkened in upper part and with a broad diagonal pruinose stripe extending from

ppn to metapleuron, margined with dark below; scaling restricted. Apn with long

and short bristles and sometimes broad dark scales; ppn with a few short upper

bristles forming a more or less continuous line with stronger posterior bristles,

upper part with or without narrow scales, lower always bare; ppl with a few

strong bristles; psp usually bare, sometimes with a few broad or very narrow

short pale scales; pst bare; stp with or. without upper bristles, lower posterior

bristles always present, scales present or absent; pra with a few bristles; 1 strong

lower mep bristle (rarely absent) and several weaker upper mep bristles always

present, scales usually absent; meron, metameron and metapleuron bare. Legs:

Coxae with pale integument and largely pale scaling; forecoxa with large anter-

ior patch of scales and a few bristles near lower margin, basolateral area with

or without scales and bristles; midcoxa with anterolateral patch of scales and

bristles; hindcoxa with anterolateral patch of scales and a few short bristles near

lower margin, sometimes a separate smaller scale patch near base, posterior sur-

face usually without scales but with a dorsoventral row of moderately strong

bristles. Femora varied in length; forefemur usually distinctly shorter or subequal

to proboscis; midfemur slightly swollen and longest; hindfemur more slender than

others. Forefemur pale scaled at base for a varied distance; anterior surface dark

scaled except for a few pale scales apically and sometimes a poorly differenti-

ated pale metallic preapical spot (Urichii Subgroup); posterior surface varied from

largely pale to largely dark. Midfemur pale scaled at base for varied distance;

anterior surface dark scaled except sometimes for a few pale scales apically and

always with 2 pale metallic spots near dorsal margin, 1 submedian and the other

preapical, rarely the 2 spots connected by metallic streak; posterior surface usu-

ally largely dark. Hindfemur extensively pale at base; anterior surface with sil-

very white to pale golden ventral streak, remainder dark except for a light me-

tallic subdorsal postmedian or preapical streak or elongate spot, sometimes con-

nected to ventral light streak, usually a few apical pale scales; posterior surface

pale scaled for variable distance from base. Knee spots poorly developed. Tibiae

usually entirely dark scaled; midtibia slightly longer than others, hindtibia short-

est. Tarsi entirely dark scaled or hindtarsal segment 4 with broad basal white

or whitish ring (Urichii Subgroup); scaling rather shaggy; basal segment of fore-

tarsus and midtarsus shorter than respective tibiae, that of hindtarsus slightly

longer than hindtibia; segment 5 of foretarsus subequal to segment 4 or slightly

shorter, that of midtarsus slightly shorter; segment 5 and to a lesser extent seg-

ment 4 of foretarsus and midtarsus with a large distal field of specialized plan-

tar bristles; segment 5 of hindtarsus about 0.5 of segment 4; pretarsi small, not

deeply inserted into tarsal segments 5; claws all simple, without basal spicules,

those of foreleg and midleg moderate and subequal, those of hindleg much short-

er; empodium spiculose; pulvilli short and spiculose. Wing: Membrane with con-

spicuous microtrichia. Veins entirely dark scaled; dorsally all veins with short

broad squame scales except for long linear or relatively short narrow plume scales

on Rs, R243, M and sometimes on R, and R3 (Urichii Subgroup). Base of vein

R_ usually with a few (remigial) bristles; cell R, longer than its stem; vein 1A

ending well distad of furcation of Cu; plical vein without scales on base of ven-

tral surface; fringe normal, dark; alula with even row of long hairlike scales on-

_ly; upper calypter with even row of long bristles. Haltere: Stem pale; at least

distal part of stem and all of knob with dark scales. Abdomen: Pregenital seg-

ments subcylindrical (Bihaicolus Group) or more or less distinctly compressed.

Valencia: Subgenus Carrollia of Culex 17

Tergite I with a more or less extensive scale patch dorsally; laterotergite with

numerous hairs and often with a few broad scales. Tergites II-VII predominantly

dark scaled and with conspicuous lateral light metallic spots of varied size and

position; white or whitish basal transverse bands present only in Iridescens Sub-

group; distal and lateral margins with bristles more prominent laterally. Sternites

II-VII with pale scales only or IV-VII with apical dark bands; apical margins with

long bristles. Cibarial Armature: Cibarial bar with a single row of 3-10 teeth

rounded or truncate apically, and sometimes with filamentous spicules alternat-

ing with teeth; lateral flange slender, with or without projecting lobe. Cibarial

dome strongly developed, with numerous closely packed minute imbricate den-.

ticles of varied shapes.

FEMALE GENITALIA. Small, inconspicuous. Segment VIII partially retracted

into segment VII, densely covered with scales and bristles; sternite longer than

tergite, caudal margin deeply emarginate, bristles moderately developed. Tergite

IX narrow, poorly sclerotized except sometimes for basal margin; dorsolateral

bristles always present. Cercus short and very broad, more or less rounded api-

cally; with more or less numerous setae in distal part. Postgenital plate promi-

nent, projecting beyond cercus; more or less rounded apically; with numerous

setae, primarily distally; base of plate distant from cowl. Cowl strongly sclero-

tized, articulating laterally with tergite IX; atrial plate present (Bihaicolus Group)

or absent. Sigma membranous, not differentiated. Insula largely membranous, with

a patch of 15-30 setae. Spermathecae 3, very large and subequal.

MALES. Essentially similar to females except for usual sexual characters. Lab-

ium with false joint distad of middle; with or without long bristles near base

and/or distally; often with pale scales ventrally. Palpus porrect, very slender, from

0.31 to nearly as long as proboscis; predominantly dark scaled but sometimes

with pale scales on mesoventral surface of segment 3; segments 1 and 2 without

scales ventrally; segments 4 and 5 with very few bristles. Antenna from slightly

longer than proboscis to distinctly shorter; flagellar whorl bristles very long and

numerous; flagellar segments 12 and 13 elongate, segment 13 slightly longer and

with short, sparse basal whorl. Claws of foreleg and midleg enlarged and uneven;

larger anterior claw of foreleg always and that of midleg sometimes (Bihaicolus

Group, Urichii Subgroup) with a submedian tooth and with a very small pul-

villus; posterior claw of foreleg and midleg always simple and without pulvillus;

hindclaws small, simple and equal as in females but with very short pulvilli.

MALE GENITALIA. Small, inconspicuous; basic structure as in the Melano-

conion Section of the genus. Segment VIII: Deeply retracted into segment VII;

proximal part poorly sclerotized and without setae or scales; distal part densely

covered with scales. Tergite broader than long; caudal border rounded laterally,

with or without median emargination; distal part with 3-6 or more rows of strong

setae. Sternite usually subquadrate; caudal border truncate, with 1 complete and

sometimes | or 2 additional incomplete rows of setae. Segment IX: Tergite short

except laterally; connected to sternite by very narrow sclerotized bar; lobes ap-

proximated or widely separated, small, simple, conical or flattened and round-

ed, with 2-12 setae directly largely laterad. Sternite without setae: variously de-

veloped. Sidepiece: Elongate and conical to short and ovoid; scales present or

absent; with or without conspicuous spicules. Dorsal surface with long bristles

Sparse, usually restricted to lateral half; usually a differentiated apicotergal seta

near base of clasper. Ventral surface with numerous long bristles, many longer

than clasper; frequently a small, rounded apicosternal process near base of clasp-

18 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

er, with long spicules and short setae. Subapical Lobe: Varied in position and

development. Proximal division always present; always with an elongate colum-

nar process bearing 2 flattened apical specialized setae (a and b) which are often

contorted, and sometimes 1 differentiated subapical or apical relatively simple

seta (c), additional setae sometimes present on stem and base of process. Distal

division either: (1) completely absent, (2) represented by a small protuberance

with 1 or 2 depressions but usually bare, or (3) a small lobe with 1 or more

specialized apical setae adjacent to proximal division or widely separated from

it. An accessory division frequently developed near base of proximal division as

a simple long digitiform process, as a complex lobe with broad dorsal expan-

sion or as 2 separate columnar processes. Clasper: Slender; strongly curved dor-

sad distally; shorter to longer than sidepiece. Variously developed from simple

and more or less uniform in width or gradually tapered to conspicuously mod-

ified with a narrow basal part and a distal head with an outer preapical curva-

ture and/or a projecting inner preapical lobe. Snout always developed. Crest ridged

or a flat sclerite. Three distal setae always developed; seta a (= spiniform) as-

sociated with beak, simple or thickened; seta b in normal position or on the

inner lobe, simple, thickened or flattened, seta c in normal position and always

simple. Phallosome: Aedeagus simple, composed of a pair of lateral plates with

a submedian tergal bridge of varied degrees of sclerotization. Lateral plate with

basal “‘hook’’; distal part with or without sternal spine, external spine or caudal

process. Paramere simple; basal piece complex. Proctiger: Strongly developed; bas-

al part very broad, distal part narrowed, usually sharply. Paraproct sclerotiza-

tion broad at base, with or without a mesal sternal lobe; distal part narrow;

crown with a single row of a few flattened, usually blunt, teeth. Basolateral scle-

rotization crescentic, projecting conspicuously dorsad under lobe of tergite IX.

Basal sternal process not developed. Cercal sclerite poorly developed, usually mem-

branous; cercal setae 1-3 or absent.

PUPAE. Cephalothorax: ‘Integumentary sculpturing tubercular. All hairs pres-

ent, variously developed; 1-C moderate, single or double; 2,3-C approximated,

subequal or 3-C longer; 4-C always shorter than 5-C, single to multiple; 5-C short

to very long; 6,7-C approximated, 6-C always shorter and anterior; 8,9-C well

caudad of trumpet base and moderately approximated. Trumpet: Not placed on

distinct tubercle; inserted closer to wing base than middorsal line; short to very

long; tracheoid in at least basal 0.25; pinna large to small, minute in kKompi. Met-

anotum: Integumentary sculpturing tubercular; hairs 10-12-C moderately devel-

oped, double to multiple. Abdomen: Integumentary sculpturing finely imbricate;

spiracular sensilla (rudimentary spiracles) well developed. Most hairs with few

branches or single. Hair 1-I (float hair) with 5-15 primary branches and 40-120

secondary branches, strongly pigmented, barbed or secondarily dendritic; 1-II al-

ways close to midline, frequently multiple; 1-IIJ-VII removed laterad, usually weak,

single or with a few branches, 1-III sometimes thickened and enlarged. Hair 2

always small and single; 2-II mesad or laterad of hair 3; hair 2-III-VII always

mesad of hair 1. Hair 3-VI mesad or laterad of hair 1; hair 3-II,III often thick-

ened and enlarged. Hair 5-IV,V nearly always long and single. Hair 6 poorly

developed, single or branched. Hair 9-II-VI normal in position and single; 9-

VII, VIII usually strongly developed. Terminal Segments: Hair 1-IX not develop-

ed. Median caudal lobe about half as long as broad. Female genital lobe sub-

equal to or slightly longer than median caudal lobe; cercal lobe poorly different-

lated. Male genital lobes (sidepieces) about twice as long as median caudal lobe.

Valencia: Subgenus Carrollia of Culex 19

Paddle: Usually ovate, always longer than broad; midrib well developed; external

buttress moderately developed; external margin with weak short spicules on ex-

ternal buttress, remainder and all of inner margin usually without distinct spic-

ules. Hair 1-P weak, single or double, rarely absent; 2-P usually absent.

FOURTH INSTAR LARVAE. Head: Head capsule subquadrate; usually slight-

ly wider than long and usually widest just caudad of antenna; ocular bulge not

developed; ventral surface flat, anterior dorsal surface moderately convex (Bihai-

colus Group) or flat (Iridescens Group); integument smooth or finely sculptured.

Labrum poorly differentiated dorsally, its anterior margin slightly concave; men-

tal plate normal, with 8-11 teeth on each side of median tooth; labial plate very

long and narrowed anteriorly; maxillary suture complete, projecting caudad of

posterior tentorial pit but not reaching collar; collar poorly developed. Mouth-

brushes with numerous filaments, some pectinate; mandible and maxilla normal.

Hair 0-C in normal position, simple; 1-C arising from a lateral process of labrum,

strongly developed and pigmented, usually curved ventromesad; 2-C always de-

veloped and single; 3-C nearly always absent but rarely represented by a spic-

ule; 4-6-C always in anterior 0.3 of head capsule; 4-C short and usually mul-

tiple; 5,6-C both moderately long and barbed, 5-C always multiple and usually

shorter than 6-C, latter usually with 2 or 3 branches, rarely single or multiple;

7-C multiple, usually barbed, shorter than 5,6-C; hairs 8,10-C in normal position;

9-C normal in development and position near level of 8,10-C (Bihaicolus Group)

or removed far caudad to near collar and more or less stellate; 11-C short, with

few to numerous branches, removed far caudad ventrally from antennal base; 12-

C long, single or double; 13-C long, always single; 14,15-C short, with a few

branches or single, position varied; 16,17-C not developed. Maxillary hair 5-MP

a simple bristle near apex (Bihaicolus Group) or greatly thickened, brush-tipped

and arising near the middle of maxilla; 6-MP short, single or branched. Antenna:

About 0.30-0.60 of head capsule length; shaft very slender and nearly uniform

in diameter, straight or slightly curved mesad; integument smooth or with very

minute inconspicuous spicules (exaggerated in figs.). Hair 1-A inserted usually in

middle third of shaft, short, not reaching apex of shaft, branches few or rarely

absent; 2-6-A short, not markedly modified. Thorax: Integument smooth or with

spicules in anterior part of prothorax. All hairs present except 13-P. Prothorax:

hair O-P very short to moderate, always multiple, sometimes dendritic or sub-

stellate; 1-3-P on strongly developed common tubercle; 1-P long, usually single;

2-P usually distinctly shorter than 1-P but sometimes subequal, single or branched;

3-P always shorter than 1-P, from 0.25 to about 0.67, and usually branched;

4-7-P all strongly developed, usually all on separate tubercles but those of 5,6-P

sometimes fused, 5-P always and 6-P nearly always single; 8-P varied in devel-

opment, with or without tubercle; prothoracic pleural group normal, 9,10-P sin-

gle or with a few branches; 12-P nearly always single; 14-P usually single. Meso-

thorax: hairs 1-4-M relatively weak, without tubercles, 3-M usually longest; 5-

M always single, very strong and with tubercle; 6,7-M both very strong and near-

ly always single, on common tubercle; 8-M very strong and multiple on large

tubercle; mesothoracic pleural group normal, 9-M always multiple, 10-M nearly

always single, 12-M always single; 13-M always short, multiple or dendritic; 14-

M short and multiple to more or less stellate. Metathorax: hairs 1-6-T relatively

poorly developed, without tubercles; 5,6-T nearly always single; 7-T very strong,

multiple and with large tubercle; 8-T short and dendritic; metathoracic pleural

group normal, tubercle with short or long spines, 9-T double to multiple, 10-T

20 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

nearly always and 12-T always single; 13-T varied, short dendritic, more or less stel-

late, or long and single. Abdomen: Integument without spicules. Chaetotaxy nor-

mal for genus. Hair 1-II minute to large, 1-III-VI usually strongest dorsal hair

of its segment and single to multiple, 1-VII very strongly developed and surpass-

ing base of siphon; 3-II-V sometimes strongly developed; 6-I-VI all strong, 6-I,

II always branched, 6-III-V single or double, 6-VI always single; 6-VII short, us-

ually dendritic; 7-I similar to 6-I and single or double, 7-II-VI usually all short

and multiple, 7-II rarely long and single (kompi); 13-I small and with few branch-

es, 13-I1,VI,VII short dendritic or multiple, 13-III-V usually strongly developed

and subequal to hair 1-III-V; other hairs usually small and weak. Segment VIII:

Comb varied; scales short, spatulate, apically fringed and in a patch (Bihaicolus

Group) or spinelike and in 1 more or less regular row. Hair 1-VIII always mul-

tiple, short to moderate; 2,4-VIII long and thin, 4-VIII always and 2-VIII near-

ly always single; 3-VIII strongly developed, multiple, barbed; 5-VIII moderately

strong, usually barbed. Siphon: Short to long; index varied from about 2.2 to

about 12.0; integumentary sculpturing imbricate, with indistinct to rather con-

spicuous fine spicules; usually darkened on basal ring; acus present, attached;

valves short; tracheae large; stirrup-shaped piece short, simple; median caudal fil-

ament apparently not developed. Pecten restricted to basal 0.5; teeth short, broad,

with or without denticles or fringes, or elongate, spinelike and with or without

marginal fringe. Siphonal hairs 1,la-S arranged in an irregular double row of 8-

16 long, strong, usually multiple barbed ventral tufts and 2 very short multiple

accessory tufts, 1 lateral (lad) and the other subventral (lav); occasionally a

third small subventral tuft present; 2-S short, simple, single, arising from mem-

brane distad of sclerotized part of siphon; 6,7,9-S simple and short; 8-S short

and branched. Anal Segment: Saddle completely ringing segment, short, acus not

developed; integument smooth or finely imbricate; spicules when present restrict-

ed to caudolateral angle. Gills equal; varied from short and rounded to long,

pointed or rounded. Hair 1-X varied, single and moderate or multiple and short;

2-X always strong and branched; 3-X always strong and single; 4-X (ventral brush)

strongly developed, hairs all on poorly developed grid without lateral bar, vary-

ing from 4-6 pairs, single to multiple.

KEYS TO GROUPS AND SPECIES

ADULTS

(Females of 3a. Panama form, 14. cerqueirai and 15. wilsoni unknown; ©

8a. Bahia form and 9a. Espirito Santo form not included)

l. Pleuron pale, whitish to greenish, and without broad diagonal pruinose

stripe; dorsocentral bristles strongly developed on disc of mesono-

tum (Bihaicolus Group) . . . 2

Pleuron darkened to near base of coxae and with a broad diagonal pru-

inose stripe from ppn to metapleuron; dorsocentral bristles not de-

veloped on disc of mesonotum (IridescensGroup) . ...... .9

Bihaicolus Group

2(1). Froboscis about 1:5 ‘of foréremurs °F FP, PY PO. “DO gyerrerol

| Proposes 1. 3.0r ess Ol Toreisin fe ee 8

3(2).

4(3).

5(4).

6(5).

7(4).

8(7).

9013:

10(9).

11(9).

1 LA 1).

Valencia: Subgenus Carrollia of Culex 21

Hindfemur with light metallic streak very narrow, at least 0.25 of fe-

mur length and confluent with light scaling anteriorly and/or pos-

teniorly Ss . . . +. infoliatus

Hindfemur with hort discret streak, fioderately broad: OTS “or less

of femur length and not connected to light scaling anteriorly or

ORI CHIORIY Es FRE BOG. Sn ch te ese, Cae ote)

Males fcee also Key to Wale Genitalia). pe ee oe ne ees, YS

Females (separation of species tenuous)! Ui ey Gey Le aT

Palpus about 0,33 of proboscis length... 5°... 6 4 1°. Se, 3, esseal

Palpus 0.7 or more of proboscis length: fae 4 ON ee ®

Abdominal sternites all uniformly pale scaled. . . . . . J. bihaicolus

Abdominal sternites IV-VII with conspicuous apical dark bands .

. 4. metempsytus

Abdominal sternites IV-VII with distinct aut dark bands; large cibar-

ial teeth more.taan 8 u-- 4 .. . . 4. metempsytus

Abdominal sternites all utiformily vale scaled or distal with only a few

apical dark scales not forming distinct bands; large cibarial teeth few-

Be ce de ear Nb i Ue er ee gre ver ee Gs fh a ta

Erect scales of vertex of head yellowish to white; abdominal sternites

entirely pale scaled; flange of cibarial bar with conspicuous lobe.

. . . . IL. bihaicolus

rest scales i” vertex of head degks vabdoniinal sternites V-VII usually

with some apical dark scales; flange of cibarial bar without conspic-

WOH Aes ee a ig cei ed ae aad eee Ae ae

Iridescens Group

Hindtarsal segment 4 with a broad basal white or whitish ring; upper

stp with 3 or 4 bristles (Urichii Subgroup) . . ; 10

Hindtarsal segment 4 entirely dark scaled; upper stp without bristles

(iVideacone Sere ges a ee a gee pie ae ge 4 rey Mae

Urichii Subgroup

Males. 6 foci ele Wick Se ee eee ee ee Re le Genii

Females, Gasitcy 0G ev BUC! oO SOC A em 7 enduzel

Iridescens Subgroup

Long slender erect scales present on anterior part of vertex to near orbi-

tal bristles; narrow decumbent scalesnumerous . . Pee nal: (..

Anterior part of vertex without erect scales, these confined to posterior

part and/or occiput; narrow decumbent scales feweor absent. «..,.. ,..44

Midlobe of scutellum with short broad dark scales in addition to a few

linear scales; male labium without long hairs distad of false joint

8. bonnei

Midlobe of soutetiunn with ae or : Jone saderiol: wide curved scales;

male labium with at least | pair of long hairs distad of false joint . 13

LSC42).

14(11).

15(14).

16(15).

17(16).

[3¢13).

19(18).

20(19).

Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

Midlobe of scutellum with linear dark scales; mesonotal vestiture pre-

dominantly dark bronzy on disc; male labium with a single pair of

long subapical hairs; male palpus entirely dark scaled . . .9. secundus

Midlobe of scutellum with long moderately wide curved pale scales;

mesonotal vestiture predominantly light bronzy to dark coppery on

disc; male labium with numerous long hairs distad of false joint; male

palpus with white scaled line on ventromesal surface . . 10. iridescens

Lateral lobe of scutellum with linear scales. . . . .. . . Jd. soperi

bates) lose witli road scales ys cui es a Se eee! OS

Females . . . oS py 6

Males (separation . soscies ieaacus: see Key to Male Genuine 2) ens

Palpus about 0.45-0.50 of proboscis; a ne of short setae below inter-

orbital bristles . . : Lieto. 226, eompt

Palpus about 0.19-0. 22 of proboscis: no ‘setae below interorbital bris-

tles (separation of species very tenuous) ci seee Soir we aT

Mesonotal and pleural integument black or very dark brown; lower edge

of dark part of pleuron sharply marked off on sternopleuron . .

SG: i2. hopakopenss

Mesarotal ae pea pee ihe brew to dark brown; lower

edge of dark part of pleuron not sharply marked off on sternopleuron

13. antunesi

A pair of short setae below interorbital bristles . . . . . . 16. kompi

Nosseiacecloy anicrorbitabbiisticssse Sec as SR ee YD

Mesonotal and pleural integument black or very dark brown; lower edge

of dark part of pleuron sharply marked off on sternopleuron .

f2. b ialoycadk

Mesonotal and pleural intemanedt light oe to dark brown; lower

edge of dark part of pleuron not sharply marked off on sternopleuron

eee ei A : 20

Lower surface of labium uniformly distinctly whitish from base to near

AEN . 5 : . . 15. wilsoni

Lower surface ‘of labium indistinctly pale, usnally only on distal part

.13. antunesi; 14. cerqueirai

MALE GENITALIA

(3a. Panama form unknown; 8a. Bahia form

and 9a. Espirito Santo form not included)

Lateral plate of phallosome with subapical external spine, its apex with-

out long caudal process; clasper simple, seta b in normal position

(Bihaicolus Group) . . oe

Lateral plate of phallosome without subapical ternal one. its ae

developed as a long caudal process; clasper apex modified, seta b dis-

placed apically or on a prominent lobe (Iridescens Group) . . . . .6

2(1).

a2).

4(2).

5(4).

6(1).

7(6).

8(6).

Valencia: Subgenus Carrollia of Culex 23

Bihaicolus Group

Subapical lobe of sidepiece without any indication of a distal division. . 3

Subapical lobe of sidepiece with distal division represented by at least

I spetialized seta grising from a procese Fe Pe Pe A Se

Subapical lobe of sidepiece with more than 30 long setae on stem and

base of proximal division; distal part of proctiger parallel-sided in dor-

sal aspect; cercal setae present. . . . ». infoliatus

Subapical lobe of sidepiece with about 43 short setae on stem and base

of proximal lobe; distal part of proctiger tapering in dorsal aspect;

cercarsetae absent 4 2 SP Pe TS at etempsy tus

Distal division of subapical lobe of sidepiece relatively broad and with

2 specialized setae with recurved apex. . ... . . . J. bihaicolus

Distal division of subapical lobe of sidepiece represented by a slender

fingerlike process with only 1 specialized seta with simple apex . . .5

Snout of clasper longer than seta a (spiniform); specialized setae a and b

of proximal division of subapical lobe unequal in size and relatively

simpte, scta c not differentiated 2 . =. . . 3. rausseoi

Snout of clasper shorter than seta a (spiniitori). specialized setae a and

6 of proximal division of subapical lobe subequal and highly contorted,

Seta € Cileren eter eo ee ge a eee

Iridescens Group

Distal division of subapical lobe of sidepiece distinctly developed and

widely separated from proximal division, accessory division not de-

veloped; caudal process of lateral plate of phallosome curved ventrad

distally; seta b of clasper simple, not thickened or flattened ei

Subgroup) . . Ae

Distal division of subapical lobe ‘of sidepiece feprésenited by” a small

tubercle or apparently completely absent, accessory division develop-

ed near base of proximal division; caudal process of lateral plate of

phallosome straight or curved laterad distally; seta b of clasper thick-

ened or Tlattened (ndescens Suberoup) <: 47% ee ae ee ee

Urichii Subgroup

Distal division of subapical lobe with 4 specialized flattened setae on

a process and a group of about 6-20 specialized flattened setae proxi-

mad’ oF process . . . 6. urichii

Distal division of subapical lobe with only | specialized bristlelike seta

on digitifomnpiactssi: 4 12-6 ies heysigh Sie Gok tox niet a7, CEE

Iridescens Subgroup

Clasper with strongly differentiated elongate spiculose preapical lobe with

flattened seta 5; lateral plate of phallosome without sternal spine . .9

Clasper with poorly differentiated broad nonspiculose preapical lobe with

thickened seta b; lateral plate of phallosome with sternal spine . . 11

9(8).

10(9).

11(8).

I2¢11):

13¢ 72):

14(12).

15(14).

2(1).

Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

Accessory division of subapical lobe of sidepiece long columnar and with

4 apical or preapical elongate flattened specialized setae. . 8. bonnei

Accessory division of subapical lobe of sidepiece not columnar and with-

out flattened specialized sefae.. 3's, Je 4c Ske es 1

Accessory division of subapical lobe distinctly separated from proximal

division, digitiform and with 1 specialized bristlelike apical seta and

a few simple setae at base; apicotergal specialized seta of sidepiece

not developed . . . . .9. secundus

Accessory division of subapical lobe very broad a joined to proximal

division, with several apical marginal setae; apicotergal specialized seta

of sidepiece developed. . ........ .. . ~ L0. iridescens

Accessory division of subapical lobe composed of 2 widely separated

columnar processes . . . . . . . 16. kompi

Accessory division of subapical lobe consisting of a single, simple or

Conipies recess i Pe oO eee ee a es

Accessory division of subapical lobe complex, with large dorsal expanded

process... 3 4 Se ME ee eee

Accessory division of suns taal lobe digitiformn, qwihout dorsal expansion

Sidepiece short, subovate; dorsal process of accessory division of sub-

apical lobe with flattened specialized setae . . . . . . Il. soperi

Sidepiece elongate, subcylindrical; dorsal process of accessory division

of subapical lobe with very long simple bristles . . 12. babahoyensis

Accessory division of subapical lobe without setae . . . . J3. antunesi

Accessory division of subapical lobe with setae 2... a. Oe we dS

Accessory division of subapical lobe with no more than 16 setae, largely

in distal hialf.s Gy. . . . . L4. cerqueirai

Accessory division of urea Jobe with at jeast 24 sete extending

Hite ase! Tite! ye ea a oA anos) hanadieti aes rea LO. WHSORT

PUPAE

(3a. Panama form and 8a. Bahia form unknown;

9a. Espirito Santo form not included)

Abdominal hair 2-II laterad of 3-II; hair 3-VI mesad of 1-VI (Bihaicolus

Group). .. ne

Abdominal hair 241 eeia i # Il: at 3-VI ee of 1-VI fitdeccens

CSPOUP) <M) A ea pe ah ait ‘ 0) Sn ie BS

Bihaicolus Group

Abdominal hairs 6-III-V usually all branched . :

. 2. guerreroi; 3. rausseoi; i 4, metempsytus

Abdominal hairs 6-Ill VV usually all single. ;

32):

4(1).

5(4).

6(4).

7(6).

8(7).

9(8).

10(9).

11(10).

Valencia: Subgenus Carrollia of Culex 25

Abdominal hair 5-VI weak, shorter than 4-VI; hair 1-I[V about twice as

long as3-IV. . . . IL. bihaicolus

Abdominal hair 5-VI strong longer hare 4 VI: ait 1-IV subequal to 3-IV

Anaesie ual nse ADSM eaters 5. infoliatus

Iridescens Group

Cephalothoracic hair 5-C strong, at least 0.67 of trumpet length (Urichi

Subgroup) . . resi.

Cephalothoracic hair SC weak, “usually only ¢ 0, 5 of trumpet length or

shorter (Iridescens Subgroup) . ; ge 3 hee . 6

Urichii Subgroup

Abdominal hair 5-VI relatively weak and short, not extending beyond

caudal border of tergite VI]. . . . . . 6. urichii

Abdominal hair 5-VI very strong and long, extending well beyond caudal

border of terete Vibo<G shou. ; i is ghee, 7 eet

Iridescens Subgroup

Trumpet fusiform, with minute pinna; abdominal hair 9-VIII more than

twice as long as paddle. Hog Caen oe. afel6. kompi

Trumpet truncate apically, with nottial pinna: abdominal hair 9-VIII

at most shently fonger than paddle = ole ks. ee

Trumpetdndexisatdeast 15203 Was) Feed. YOR ek Tenis £3; ween

Weumpet mdex Jess than [4G aa See GS 8

Trumpet strongly flared apically and with broad meatus. . 14. cerqueirai

Trumpet at most only slightly flared apically, meatus moderate. . . .9

Trumpet index 10.0 or more pth Tee ar eS ee EY

9. secundus; 12. babahoyensis; 13. antunesi

Teinees index less than 8.0 igen Wetepi neg s Sey eae ae a eye

Abdominal hair 9-VII weak and without barbs ... . . . JJ. soperi

Abdominal hair 9-Viletrone and barbed: c: wersaiitivds th wre ie oe odd

Abdominal hairs 5-IV,V relatively short, not reaching caudal border of

next tergite . . . . 10. iridescens

Abdominal hairs 5-IV Vv ee lone extending wel: beyond caudal border

of next terettiers: fi; 2 : cig oo 6G, ome

FOURTH INSTAR LARVAE

(Sa. Bahia form and 9a. Espirito Santo

form not included)

Head hair 9-C short, inconspicuous, normal in position at level of 10-C;

comb scales apically fringed, more or less spatulate, and arranged in

a patch of 2-5 irregular rows (Bihaicolus Group). ....... .2

2(1).

3(2).

4(2).

5(4).

6(4).

7(1)-

8(7).

9(7).

10(9).

11(10).

Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

Head hair 9-C large, conspicuous, near collar; comb scales elongate, spine-

like, and arranged in 1 more or less regular row, rarely a few scales

displaced into second: rew (iridescens Group) five. wT

Bihaicolus Group

Ventral brush with at least 12 hairs, the majority except the proximal

pair usually single. . . is

Ventral brush hag with 10 hae rarely with re ‘the majority branched

Siphonal hair lad-S in proximal half of siphon. . . . . . J. rausseoi

Siphonal hair lad-S in distal half of siphon. . ... 3a. Panama form

Abdominal hairs 6-III-V all single. . . Rie OU ee: . emeaiod

Abdominal hairs 6-III-V usually all branched a. aeAeE CREO, OE, Roa esO

Head hair 6-C single. . . Pie apres ee Ben oes COMES

Head hair 6-C double or triple . Be RT ee hake eth ngs gs ge: i a RO TREEES

Anal saddle with unsclerotized area on caudolateral border; metathoracic

hair 3-T usually double or triple; abdominal hair 3-III usually single

OF Goulie. 2 : . .2. guerreroi

Anal saddle completely sclerotized on 1 candolateral border; metathoracic

hair 3-T usually with at least 5 branches; abdominal hair 3-III usually

atdoast-triple is sation Poh eee wet seo. <ehin Lo feanetempsytus

Iridescens Group

Ventral brush with at least 5 pairs of hairs, usually with 6 pairs; pro-

thoracic hair 8-P similar in development to 7-P, usually double or

triple (Urichii Subgroup). . . .8

Ventral brush with 4 pairs of hairs, rarely with 9 hairs: prothoracic

hair 8-P different in development from 7-P, multiple, weak or stel-

bate divide seems Subs(Oup) 2. Sa es eee

Urichii Subgroup

Siphon with at least 7 pairs of ventral tufts, usually with 8 pairs or 15

hairs; abdominal hair 1-IV reaching level of rudimentary spiracle of

Ol segment Vio. as . . 6. urichii

Siphon with 6 pairs of ventral tufts: abdominal hair 1-1V shorter, not

fegentae posterior border Of SevMIent Vo... nie sec niet 7 anduzei

Iridescens Subgroup

Abdominal hair 7-II long, similar in development to 7-I . . . 16. kompi

Abdominal hair 7-II short, weak, different in development from 7-I . 10

Abdominal hairs 6-IIJ-V apparently alldouble. . . . . . . JJ. soperi

Abdominal hairs 6-III-V predominantly single. . . ....... «211

Pecten teeth apparently with distinct marginal denticles. . 10. iridescens

Pecten teeth with indistinct marginal spicules or spinelike . . . . . 12

Valencia: Subgenus Carrollia of Culex an

12(11).. Abdominal hair 1-VII single; prothoracic hair 8-P weak, its branches

smooth, its basal tubercle small; pecten teeth broad and with indis-

tinct iharsinal spicules: . yt; Nae

Abdominal hair 1-VII usually doable: prOthoracic ae 8&P ceconi its

branches barbed, its basal tubercle large; pecten teeth largely spinelike

13(12). Siphon index usually less than 4.0; head hair 1-C very long and slender

and curved mesad on apex . . . FSS SC AVOnne

Siphon index usually more than 6.0; “head hair 1-C shorter, stouter and

cutved mesad Near Basen. Pec 7 Ray i Pe i Se, seen

14€12). °’Siphen*index “more tham9.0: 22" 86 Ak Soe ee eS wibsoni

Siphon index Tess tran GAy rE ee ee ae ee ee See

15(14). Head hair 5-C usually with about 15 branches; 7-C with about 10 branches

. . 14. cerqueirai

Head hair 5-C ustially ‘with less than. 10 branches: TL with less than 6

branches 2s one, ieee ie Sine Seer ae EE A

16(15). Prothoracic hair 8-P usually with 14 or more branches . . 13. antunesi

Prothoracic hair 8-P usually with not more than 12 branches . . .

i Nshenierisis

BIHAICOLUS GROUP

FEMALES. Head: Band of broad decumbent scales of vertex complete, nar-

row or moderately broad; narrow decumbent scales of vertex and occiput very

numerous; erect scales very long and numerous, reaching orbital bristles. Pro-

boscis at least as long as forefemur, usually longer, 1.0-1.7. Palpus not more than

0.25 of proboscis. Antenna shorter than proboscis. Thorax: Mesonotal integu-

ment light brown to black; vestiture of dark bronzy linear, only slightly curved

scales sparse throughout, inner dorsocentral line bare to beyond distal half. Scu-

tellar scales all linear as on mesonotum. Anterior acrostichals sometimes devel-

oped on disc. Anterior and posterior dorsocentral and prescutellar bristles all

well developed and in a continuous line; prescutals present. Pleural integument

always light, greenish or whitish. Upper stp bristles strong and in a curved row

joining lower bristles. Legs: Hindtarsus entirely dark scaled. Segments 5 and 4

of foretarsus and midtarsus with an apical field of plantar bristles. Wing: Plume

scales long and linear, present on R, and R;3 as well as on Rs, R243 and M.

Abdomen: Pregenital segments subcylindrical; tergites without basal dorsal light

bands. Cibarial Armature: Denticles of cibarial dome triangular, rounded or trifid.

MALES. General characters as in the females. Proboscis distinctly longer than

forefemur. Palpus and antenna shorter than proboscis, former as short as 0.33

(rausseoi). Anterior claw of both foretarsus and midtarsus with premedian tooth.

Plantar bristles of foretarsus and midtarsus not developed.

MALE GENITALIA. Segment VIII: Tergite without median emargination; with

4 or 5 rows of setae. Sternite with 2 incomplete rows of setae distally. Segment

IX: Tergite largely membranous dorsally; lobes widely separated, more or less

28 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

conical. Sternite long but without sclerotized caudal projection between sidepieces.

Sidepiece: Conical, mesal surface usually concave. Apicosternal process poorly

developed. Subapical Lobe: Accessory division not developed. Distal division small

and approximated to proximal or absent. Clasper: Simple, without differentiated

head; shorter than sidepiece; only moderately curved dorsad; parallel-sided or

slightly tapered to apex. Preapical lobe not developed; seta a developed as nor-

mal thickened spiniform; seta b simple, in normal position. Crest ridged. Lateral

Plate: Broad; sternal and external spines both developed; caudal process not de-

veloped. Proctiger: Paraproct sclerotization without mesal sternal lobe.

PUPAE. Cephalothoracic hair 5-C always very strong and at least as long as

trumpet. Trumpet short, index usually less than 7.0; pinna always large. Abdom-

inal hair 2-II laterad of 3-II; hair 3-VI mesad of 1-VI; hair 3-IJ always weak; 1-

III always strong and longest hair of segment and 3-III always weak; 5-VI usually

weak.

LARVAE. Head capsule with anterior dorsal surface moderately convex so

that hairs 5,6-C appear anterior to antennal base in dorsal aspect; integument

without sculpturing visible at 100X; hair 9-C normal in development and posi-

tion near level of 8,10-C; hair 12-C about halfway between antennal base and

collar; 14-C near anterior ventral border of head capsule. Maxillary hair 5-MP

a simple bristle. Anterior part of prothorax without spicules; prothoracic hair

Q-P always short or minute, 8-P varied; mesothoracic hair 14-M very small and

dendritic; metathoracic hair 13-T short and multiple. Abdominal hair 1-II short

or minute; 3-VI usually moderate, single to quadruple; 11-I with a few branches.

Comb scales short, apically fringed, more or less spatulate, and arranged in a

patch of 2-5 rows; hair 1-VIII varied in development. Integument of siphon with-

out elongate spicules visible at 100X; pecten teeth scalelike or spinelike; ven-

tral siphonal tufts usually 5 or 6 pairs; siphon index usually less than 4.0. Hair

1-X branched; 2-X with a few subequal branches; ventral brush usually with 5

or 6 pairs of hairs.

DISCUSSION. Nearly every character state listed above for all stages in the

diagnosis of the group represents the primitive condition for the subgenus, as

determined in the chapter on taxonomic characters. Within the group itself some

features show derived states as indicated by their range of expression. The group

occupies the northernmost portion of the known distribution of Carrollia, ex-

tending into southeastern Mexico in the north and with a southern limit in the

Amazon Basin.

The group is so compact that there is no justification for dividing it into sub-

groups. However, solely on the development of the subapical lobe of the male

genitalia, 2 phyletic lines may be recognized: the Bihaicolus Complex of 3 spec-

ies and the Infoliatus Complex of 2 species. This division is supported to some

extent by the ornamentation of abdominal sternites III-VII. The unnamed Panama

form (sp. 3a), known in the larval stage only, cannot be placed with certainty

in either complex at this time.

The Bihaicolus Complex is characterized by the retention of the distal divi-

sion and a relatively short proximal division in the subapical lobe. These are un-

doubtedly primitive character states and on this basis as well as the presence of

6 pairs of hairs in the ventral brush of the larva of rausseoi, the Bihaicolus Com-

plex can be considered closer to the original stock of Carrollia than the Infoliatus

Complex. Abdominal sternites III-VII are either light or with only scattered api-

cal dark scales. The Bihaicolus Complex is more northern in distribution, not

Valencia: Subgenus Carrollia of Culex ae

reaching the Amazon Basin, but included entirely in its central range is metempsy-

tus of the Infoliatus Complex. Two of the species, guerreroi and rausseoi, are

relicts with very restricted known distributions in the coastal range of north-

central Venezuela and bihaicolus is the dominant derived form with a distribu-

tion encompassing the entire range of the complex.

The Infoliatus Complex is characterized by the loss of the distal division and

the elongation of the proximal division of the subapical lobe, both derived states.

Abdominal sternites IIJ-VII have distinct apical dark bands. The distribution of

the complex is more southern and the 2 species are completely allopatric; metem-

psytus, as noted above, occurring entirely within the central range of the Bihai-

colus Complex, and infoliatus occupying an extensive area in the Orinoco Basin,

the Guianas and the Amazon Basin. Culex infoliatus exhibits more derived states

than metempsytus, particularly in the development of the metallic streak on the

hindfemur (unique in the group), and in the reduction in the branching of hair

6 on abdominal segments HI-V of both the larva and the pupa (shared with Di-

haicolus). On this basis it appears to be the most derived species in the entire

Bihaicolus Group.

As noted on page 38, the Panama form (sp. 3a) shows some resemblance to

rausseoi and possibly introgression with metempsytus. This form may prove to

be an annectent relict between the 2 complexes or a relict of either complex.

Although male genitalic characters distinguish the 2 complexes and are diag-

nostic of every species in the group, the separation of the species in other stages

is rather tenuous. In the keys to these stages I have had to use diagnostic features

rather than phyletic ones as the latter are either not apparent or difficult to

interpret.

On external features of the adults, only guerreroi and infoliatus are readily

diagnosed in both sexes and rausseoi in the male, as indicated in the key. In the

pupal stage, guwerreroi, rausseoi and metempsytus are apparently indistinguishable

and the key characters for the separation of bihaicolus from infoliatus are not

entirely reliable. The separation of the fourth instar larvae is equally difficult.

The close resemblance of the adults in external features has been responsible

for many misidentifications of bihaicolus, metempsytus and infoliatus by several

investigators in the past.

The group as a whole shows nearly the entire range of breeding sites known

for the subgenus. The dominant species (bihaicolus, metempsytus, infoliatus) have

a broad spectrum of breeding sites and the relict ones (guerreroi, rausseoi and

sp. 3a) appear to be restricted to 1 or 2 types. The only records of adults col-

lected in nature are 1 female and 2 males of bihaicolus, and 2 males and an un-

specified number of females of infoliatus, the latter attracted to human or animal

bait.

I. Culex (Car.) bihaicolus Dyar & Nunez Tovar

Figs. 2,6,9-1 1

1927. Culex (Carrollia) bihaicolus Dyar and Nunez Tovar, 1927:4-5; 1928:91. TYPE: Lecto-

type by present designation, male, 1 of 4 specimens bearing type labels, Ocumare de

la Costa [incorrectly Maracay], Aragua, Venezuela, 5 July 1927, M. Nunez Tovar

[USNM]. Contrary to the interpretation of Stone and Knight (1957:44) the 4 spec-

imens bearing the blank printed type labels and handwritten labels ‘“Bihai’’ are un-

30 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

doubtedly syntypes which were incorrectly labelled Maracay, the city of residence of

Nunez Tovar where they may have been reared from larvae collected in Ocumare de

la Costa. That Nunez Tovar was in Ocumare de la Costa on 5 July 1927 is documented

by 2 specimens of Deinocerites melanophylum Dyar & Knab, 1907 bearing these data

(Belkin and Hogue, 1959:434). Other material collected by Nunez Tovar in various

localities in Venezuela is similarly mislabelled Maracay in the USNM, including 4 spec-

imens of D. melanophylum (loc. cit.).

Culex (Carrollia) bihaicolus of Edwards (1932:221); Galindo, Carpenter and Trapido (1951:

104,105,111,113,126); Levi-Castillo (1952:553; 1953a:35); Galindo and Blanton (1955:73);

Vargas (1956:24); Stone and Knight (1957:44); Stone, Knight and Starcke (1959:282); Belkin,

Schick and Heinemann (1965:76); Cova Garcia, Sutil and Rausseo (1966a:28,117; 1966b:

42,84,141,233,343); Barreto and Lee (1969:433); Cova Garcia, Sutil and Pulido (1971:205).

Culex (Carrollia) bihaicolus in part of Lane (1939:81; 1953:509).

Culex (Carrollella) bihaicolus of Dyar (1928:284,285 534).

Culex (Carrollela) infoliatus in part of Komp (1936:64; 1956:39).

FEMALE (figs. 6,9). Wing: 3.7 mm. Proboscis: 2.7 mm. Forefemur: 2.2 mm.

Abdomen: about 3.2 mm. In general similar to other species of group, especially

guerreroi and rausseoi; diagonostic characters as in the key. General characters as

for the group; description based on 18 topotypic and 30 other specimens from

Venezuela. Head: Band of broad decumbent scales of vertex relatively narrow,

scales coppery with bluish to greenish iridescence; narrow decumbent scales yellow-

ish white, very numerous; erect scales yellowish white, concolorous with narrow

decumbent scales; lateral patch of scales silvery with light bluish iridescence. Pro-

boscis 1.04-1.23 of forefemur. Labium entirely dark, scales dark brown to black and

with bluish green iridescence. Palpus 0.23-0.25 of proboscis; scales similar to those

of labium. Antenna 0.89-0.91 of proboscis; torus with 15-20 setae. Thorax: Meso-

notal integument dark brown, strongly contrasting with light greenish pleuron. An-

terior promontory with 2 or 3 bristles; prescutal bristles 5 or 6; all mesonotal bris-

tles dark. Midlobe of scutellum with 6-8 long bristles and numerous dark linear

scales. Apn with 10-15 bristles; ppn with 4 or 5 narrow dark scales and 4-6 posterior

bristles; ppl bristles 5 or 6; stp with about 15 bristles in curved line; pra bristles 8-10;

umep bristles 5. Legs: Coxal scales whitish. Forefemur pale scaled to apex of ventral

surface and with pale line on dorsal margin extending to about 0.5 from base; with-

out light metallic spot. Midfemur pale scaled at extreme base only, chiefly on pos-

terior surface; light metallic spots small and discrete. Hindfemur pale scaled ventral-

ly on both anterior and posterior surfaces to about 0.5-0.6 from base; light metallic

marking elongate, about 0.15 of femur length, discrete, not connected to ventral

pale scaling. Knee spots slightly indicated by a few pale scales on apex of all femora.

Abdomen: Light metallic spots of tergites large, with pale violet iridescence, basal

on II-V and submedian on VI,VII; marginal bristles of tergites golden. Sternites uni-

formly light golden, without apical dark bands or scales; apical bristles golden. Cibar-

ial Armature: Cibarial bar with rounded lobe on lateral flange; cibarial teeth 3-

5; cibarial dome with trifid denticles. Genitalia: Tergite IX with 3 or 4 lateral

bristles; insula with 10-15.

MALE (fig. 6). Wing: 3.2 mm. Proboscis: 2.5 mm. Forefemur: 2.0 mm. Ab-

domen: about 2.5 mm. In general similar to other species of the group; diagnostic

characters as in the key. Essentially similar to the female except for sexual differ-

ences and more prominent knee spots. Description based on 10 topotypic and 26

other specimens from Venezuela. Palpus 0.76-0.83 of proboscis length; 3 or 4 con-

Valencia: Subgenus Carrollia of Culex 31

spicuous bristles on segment 5. Tooth of anterior claw of foretarsus and midtarsus

blunt and arising before middle.

MALE GENITALIA (fig. 10). Lobe of tergite IX usually with 4 (3-6) bristles.

Sidepiece without scales. Proximal division of subapical lobe with large protu-

berant base bearing numerous fine apically curved bristles; stem elongate and

with only a few bristles; setae a and b subequal, recurved apically but not con-

torted. Distal division moderately developed as a rounded lobe with 2 apically

recurved thickened setae. Clasper not markedly tapered distally; spiniform long-

er than snout. Distal part of proctiger narrow, more or less parallel-sided; para-

proct crown usually with 4 (3-8) teeth; cercal setae absent.

PUPA (fig. 10). Abdomen: 3.3 mm. Trumpet: 0.65-0.75 mm. Paddle: 0.85-

0.95 mm. Very similar to infoliatus; diagnostic characters as in the key. General

chaetotaxy based on 10 reared specimens from Venezuela. Cephalothorax: Pig-

mentation light yellow; sculpturing weak, without definite pattern. All hairs ex-

cept 5-C shorter than trumpet; 1(1;1-2) slightly longer than hair 7; 2(2,3;1-3)

shorter than hair 3; 3(2;1-2); 4(1,2;1-3) about 0.5 of hair 7; 5(1;1-2) sometimes

finely barbed; 6(1,2;1-4) very short; 7(2;1-3); 8(1) longer than hair 9; 9(1-2).

Trumpet: Distinctly widened distally; index about 4.4 (3.8-6.0); pinna about 0.33.

Pigmentation light brown. Metanotum: Pigmentation light yellow; sculpturing weak;

hair 10(1,2) longer than hairs 11,12; 11(23;1-3); 12(1;1-2). Abdomen: Pigmen-

tation yellow to light brown, anterior mesal areas of tergites III,IV slightly dark-

er; sculpturing moderately developed. Segment I: hair 1 (about 45; 37-48). Seg-

ment II: hair 1(6,7;3-8) forked and longest hair of segment; 3(1,1-2); 4(4-6)

forked and small; 5(1). Segment III: hair 1(1;1-2) surpassing caudal border of

following tergite; 3(1,1-2); 4(3,4;2-5); 5(1,2) shorter than hair 3. Segment IV:

hair 1(131-2); 3(2,3;2-5) shorter than hair 6; 4(2;1-3), 5(1) strong, always reach-

ing level of rudimentary spiracle VI. Segment V: hair 1(1) longer than hair 6;

3(231-2); 4(3,4;3-5); 5(1) strong, always reaching level of rudimentary spiracle

VII. Segment VI: hair 1(1) reaching level of rudimentary spiracle VII; 3(1,2;

1-3); 4(3;2-4) subequal to hair 3; 5(2,3;2-4) very short. Segment VII: hair 1(1;

1-3) longer than hair 4; 3(2,3;1-3) shorter than hair 4; 4(1;1-2); 5(3,4;2-5) sub-

equal to hair 6; 6(4;2-4); 9(3,4;2-7) barbed, subequal to hair 9-VIII. Segment

VIII: hair 4(1;1-2) fine, subequal to hair 4-VII: 9(3,4;3-6).

LARVA (fig. 11). Head: 0.95 mm. Siphon: 0.90 mm. Anal Saddle: 0.30 mm.

Very similar to infoliatus; diagnostic characters as in the key. General chaetotaxy

based on 8 reared and 2 other specimens from Venezuela. Head: Integument light

yellow to brown; collar darkened. Mental plate with 9-11 lateral teeth. Hair 4

(4,5;3-6); 5(7,8;4-8); 6(1); 7(5,6;4-8); 8(1) shorter than hair 7; 9(2,3;2-4) short-

er than hair 8; 10(2;1-2); 11(3,4;2-5); 12(€1); 14(131-2); 15(3,4;2-4). Antenna:

About 0.33 of head length; hair 1(2,3;1-3). Thorax: Prothorax: hair 0(6,7;6-8)

minute; 1(1;1-2); 2(1); 3(2;1-2); 4(2); 7(2;2-3); 8(2-3) moderately long, with spike-

like barbed branches; 9(1,2;1-3); 10(1;1-2); 11(4,5;2-5); 14(1;1-2). Mesothorax:

hair 1(2,3;1-3); 2(2,331-4); 3(131-2); 4(2,3;1-4); 8(7,8;7-10); 9(6;6-8); 11(1-4); 13

(7-9;6-9); 14(11-16). Metathorax: hair 1(2,3;1-3); 2(2;1-2); 3(4,5;3-5); 4(2-3); 6

(131-2); 7¢(7-10); 8(6-9;5-10); 9(3-5); 11(2,3;1-3); 13(6-8;6-11). Abdomen: Seg-

ment I: hair 1(4-6;2-6) very short; 6(2;2-3); 7(1). Segment II: hair 1(2;1-2) very

short; 3(2,3;1-3) short; 6(2;2-3). Segment III: hair 1(2-3) long, strong; 3(1-2)

moderate; 6(1); 13(3) long, strong and subequal to hair 1. Segment IV: hair 1

(2-3) strong, longer than hair 1-III; 3(2-4;2-5) short; 6(1); 13(3) subequal to hair

1. Segment V: hair 1(2-3) strong, usually shorter than hair 1-I[V; 3(1;1-2) sub-

32 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

equal to hair 1; 6(1); 13(3;3-4) subequal to hair 1. Segment VI: hair 1(2;2-3)

very strong, subequal to or longer than hair 1-IV; 6(1). Segment VII: hair 1

(2;1-3). Segment VIIT: Comb scales (29-54) distinctly spatulate and conspicuous-

ly fringed; hair 1(2,3;2-4) about 0.5 of hair 2; 3(5-7;5-8); 5(1;1-2) slightly longer

than hair 1. Siphon: Pigmentation light yellow; sculpturing weak. Index about

2.5-3.0. Pecten teeth (7-10) short, with distinct fringes on ventral margin. Ventral

siphonal tufts (10-12) usually triple (3 or 4). Anal Segment: Saddle shallowly

emarginate laterally; pigmentation light yellow; sculpturing weak except caudo-

dorsad. Hair 1(2,3;2-4); 2(5,6;4-7). Ventral brush with 5 pairs of hairs, all usu-

ally double (2,1-2). Gills slightly more than 2.0 of dorsal saddle length.

SYSTEMATICS. Culex bihaicolus is the dominant widespread species of its

complex which includes also the relict guerreroi and rausseoi. It is interpreted

here as the most derived species of the complex because of the reduction in the

branching of hair 6 on abdominal segments III-V of both the larva and the pupa

(shared with the derived nominate species of the Infoliatus Complex) and the

single head hair 6-C of the larva (unique in the group).

The adults of bihaicolus are superficially very similar to other species of the

group and were often confused with infoliatus in the past (see taxonomic refer-

ences under infoliatus). Komp (1936:64; 1956:39) incorrectly synonymized the

2 species. As noted in the key and description, adults of infoliatus are readily

separated from bihaicolus by the long metallic streak on the hindfemur and con-

spicuous apical dark bands on the abdominal sternites.

The key characters for the separation of the adults of bihaicolus from guerreroi

and rausseoi on external features are quite tenuous, except for the male of raus-

seoi. However the male genitalia of the 3 species are very distinct and diagnostic.

As indicated above, the pupae and larvae of bihaicolus and infoliatus are very

similar and share a number of derived character states not present in other mem-

bers of the group. The separation of the species in the pupal stage by the key

characters may not be entirely reliable but it appears to be clearcut in the larva.

In the large sample available I have found only a normal range of variation

in all stages and have not detected any geographical population differentiation

over the wide range of bihaicolus. There is also no indication of any significant

differences in specimens from the wide variety of breeding sites.

BIONOMICS. Culex bihaicolus has an extremely wide range of breeding sites,

approached in the subgenus only by that of urichii. It is the only species of

Carrollia breeding in the flower bracts of Heliconia and the leaf axils of Dieffen-

bachia. In 47 collections, immature stages have been recorded from the follow-

ing natural sites: Heliconia (5); Dieffenbachia (1); fallen leaves, palm spathes and

fronds (10); fallen cacao pods and coconut shells (3); treeholes (12); and cut or

broken bamboo (8). They were also common in various types of artificial con-

tainers (10), including once each in a hole in a cement bridge and in a wooden bowl

trap.

Culex bihaicolus is equally catholic in its association with other container-breed-

ing mosquitoes. It has been found with 5 species of Carrollia: metempsytus (2),

rausseoi (5), urichii (2), secundus (1), bonnei (1); and with 20 other species:

Aedes (H.) ioliota (5), A. (O.) fluviatilis (2), A. (P.) daryi (1), A. (P.) impostor

(1); Anopheles (An.) eiseni (4), An. (K.) neivai (2); Culex (Anoed.) conservator

(1), C. (C) corniger (3), C. (C.) mollis (3), C. (C.) quinquefasciatus (1), C. (Microc.)

kukenan (2); Haemagogus (S.) equinus (1), H. (S.) soperi; Limatus asulleptus (7),

L. durhamii (3); Orthopodomyia fascipes (2); Sabethes sp. (1); Toxorhynchites

Valencia: Subgenus Carrollia of Culex 33

sp. (2); Trichoprosopon (T.) digitatum (4); Corethrella sp. (1).

Although the immature stages are quite common, adults of bihaicolus have been

rarely collected in nature. I collected a single female flying over a tin can contain-

ing immature stages (VZ 141); 2 males from Costa Rica [USNM] may have been

obtained also in the adult stage.

DISTRIBUTION (fig. 2). From southeastern Mexico through Central America

to Pacific drainage in Ecuador and Caribbean drainage in Colombia and Vene-

zuela at elevations of 30-1600 m.

Material examined: 2,204 specimens; 658 males, 224 females, 909 larvae, 413

pupae; 284 individual rearings (188 larval, 62 pupal, 34 incomplete).

COLOMBIA. Antioquia: Providencia, Anori R. valley, C. Porter [UCLA]. Valle: Rio Raposo

(COL 3) [UCLA].

COSTA RICA. Cartago: Atirro (CR 388) [UCLA]. Suiza de Turrialba [USNM]. San Jose:

San Isidro, T. Aitken [USNM].

ECUADOR. F'smeraldas: Quininde, Levi-Castillo [USNM]. Los Rios: Balzar, Hanson [UCLA].

Valencia (ECU 112) [UCLA].

GUATEMALA. Alta Vera Paz: Trece Aguas (GUA 33) [UCLA].

MEXICO. Veracruz: Cordoba (MEX 34,41,42,44) [UCLA]. Rio Metlac (MEX 441,442)

[UCLA].

PANAMA. Darien: Alturas de Nique (PA 1059) [UCLA]. Cerro Mali (PA 364-366,370,377)

[UCLA]. La Laguna (PA 450,451) [UCLA]. Rio Pucro headwaters (PA 375) [UCLA]. Rio

Tacarcuna valley (PA 404,431 ,432,435 ,436) [UCLA] . Panama: Campana, S.J. Carpenter [UCLA].

Cerro Azul (PA 58) [UCLA]. Cerro El Jefe, P. Galindo [UCLA]. Cerro La Victoria, S.J. Carpenter

[UCLA].

VENEZUELA. Aragua: Choroni, road to (VZ 359) [UCLA]. Guamita, Nunez Tovar [USNM].

Maracay, Nunez Tovar [USNM]. Ocumare del la Costa, Nunez Tovar, type series [USNM];

Ocumare de la Costa, road to (VZ 186,280,310,329) [UCLA]. Rancho Grande (VZ 140,141,

156-158,255,296,303) [UCLA]. Tio Julian, Nunez Tovar [USNM]. Turmero, Nunez Tovar

[USNM]. Carabobo: Mariara, road to (VZ 246) [UCLA]; Nunez Tovar [USNM]. Miranda:

Cerro el Pipe (VZ 162) [UCLA].

Additional Record From the Literature

MEXICO. Chiapas (Vargas, 1956:24).

2. Culex (Car. ) guerreroi Cova Garcia, Sutil & Pulido

Figs. 2,9,12,13

1971. Culex (Carrollia) guerreroi Cova Garcia, Sutil and Pulido, 1971:203-210. TYPE: Holo-

type male (1677-2) with slides of male genitalia and associated larval and pupal skins,

Colonia Tovar, Aragua, Venezuela, date and collector not specified [MDM].

FEMALE (fig. 9). Wing: 3.4 mm. Proboscis: 2.9 mm. Forefemur: 1.9 mm.

Abdomen: about 3.1 mm. Differentiated from all other members of the group

by the very long proboscis, about 1.5 of forefemur length, otherwise very sim-

ilar to rausseoi and bihaicolus. In general similar to bihaicolus and differing from it

in the following features. Description from 8 topotypic specimens from Venezuela.

Head: Band of broad decumbent scales of vertex narrowed on midline, scales darker

in middle and caudolaterad, white laterally behind orbitals before joining white

lateral spot; narrow decumbent scales whitish, less numerous; erect scales all dark.

Proboscis 1.53 of forefemur. Palpus 0.17-0.20 of proboscis. Antenna 0.89-0.90 of

34 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

proboscis; torus with 6-12 setae. Thorax: Pleural integument light green to gray.

Anterior promontory with 2-4 bristles; prescutal bristles 2 or 3. Apn usually with

10 bristles and 2-4 light scales; ppn with 2 or 3 narrow dark scales and 3 or 4 pos-

terior bristles; stp with 11 or 12 bristles in curved row and 2 or 3 light scales; pra

bristles 3-5; umep bristles 8-11. Legs: Coxal integument light green. Hindfemur

more extensively light scaled, to near apex; light metallic marking closer to light

scaling of anterior surface but usually not connected to it. Abdomen: Marginal

bristles of tergites darker, bronzy. Sternites IV-VII with light brown apical scales

not forming distinct bands. Cibarial Armature: Cibarial bar with narrow lateral

flange without lobe; teeth 5 or 6; cibarial dome with triangular denticles. Genitalia:

Tergite IX with 4 or 5 lateral bristles; insula with about 25.

MALE. Wing: 3.3 mm. Proboscis: 2.9 mm. Forefemur: 1.9 mm. Abdomen:

about 3.1 mm. Differentiated from all other members of the group by the very

long proboscis, at least 1.5 of forefemur length, otherwise very similar to raus-

seoi and metempystus. Essentially similar to the female except for usual sexual

differences. Proboscis 1.53-1.70 of forefemur. Palpus 0.53-0.64 of proboscis. An-

tenna 0.73-0.74 of proboscis. Specimens studied: 6 from the type locality in

Venezuela.

MALE GENITALIA (fig. 12). Lobe of tergite IX usually with 6 (4-9) bristles.

Sidepiece with scales. Proximal division of subapical lobe with simple broad base

bearing numerous fine simple bristles; stem short and with only a few bristles;

setae a and b relatively short and markedly contorted. Distal division developed

as a fingerlike process bearing a single saberlike apical seta. Clasper slightly tap-

ered distally; spiniform longer than snout. Distal part of proctiger relatively broad

and more or less parallel-sided; paraproct crown usually with 11 (8-11) strong

teeth; cercal setae absent.

PUPA (fig. 12). Abdomen: 3.50 mm. Trumpet: 0.75 mm. Paddle: 0.90 mm.

Apparently indistinguishable from rausseoi and metempsytus,; diagnostic charact-

ers from other members of the group as in the key. General chaetotaxy based

on 9 specimens from Venezuela. Cephalothorax: Pigmentation light yellow to

brown; sculpturing moderate. All hairs distinctly shorter than trumpet except 5-

C which is subequal; 1(2;2-3) longer than hair 4; 2(2;2-3) subequal to hair 6;

3(2;2-3) longer than hair 6; 4(3;2-5); 5(2,3;2-4); 6(2;1-3); 7(4;3-4) subequal to

hair 1; 8(1) longer than hair 9; 9(1,2). Trumpet: Moderately widened distally;

index about 6.1 (5.0-6.8); pinna about 0.25. Pigmentation light brown. Meta-

notum: Pigmentation yellow to light brown; sculpturing moderate; hair 10(1;

1-2) longer than hairs 11,12; 11(2;1-3); 12(2;1-2). Abdomen: Pigmentation yel-

low to light brown, anterior mesal areas of tergites III-V slightly darker; sculp-

turing moderate. Segment I: hair 1(25-33). Segment II: hair 1(2,3;1-4) reaching

level of rudimentary spiracle III; 3(2;1-2) subequal to hair 5; 4(5,6;3-7) short;

5(1,2). Segment III: hair 1(1,2;1-3) strong and surpassing caudal margin of ter-

gite IV; 3(2;1-2) subequal to hair 5; 4(3,4;2-5) short and forked; 5(2). Segment

IV: hair 1(1,2) weak but surpassing level of rudimentary spiracle V; 3(2,3;2-4);

4(2,3) short; 5(1) surpassing level of caudal margin of tergite V. Segment V:

hair 1(1) longer than hair 4; 3(2,3) weak but longer than hair 6; 4(3,4;2-4);

5(1) surpassing level of caudal margin of tergite VI. Segment VI: hair 1(1) reach-

ing level of rudimentary spiracle VII; 3(2;1-3); 4(3;3-4) subequal to hair 3; 5(1;

1-3). Segment VII: hair 1(1;1-3) nearly reaching alveolus of hair 4-VIII; 3(2;1-

3); 4(2;2-3); 5(3,4) very short; 6(3,4;2-5); 9(3,4;2-4) barbed and subequal to

hair 9-VIII. Segment VIII: hair 4(2;1-2); 9(3,4;3-5).

Valencia: Subgenus Carrollia of Culex 35

LARVA (fig. 13). Head: 1.00 mm. Siphon: 1.00 mm. Anal Saddle: 0.30 mm.

Very similar to metempsytus,; diagnostic characters as in the key. General chaeto-

taxy based on 1 reared and 6 whole specimens from Venezuela. Head: Integu-

ment light yellow to brown; collar darkened. Mental plate with 8-10 lateral teeth.

Hair 4(4;2-5); 5(7,8;7-9); 6(2); 7(4,5;4-7); 8(1;1-2) shorter than hair 7; 9(4,5;3-

6) very short; 10(2;1-2); 11(4;4-5); 12(2;1-3); 14(1); 15(2-3). Antenna: About

0.40 of head length; hair 1(3;2-3). Thorax: Prothorax: hair 0(6;5-8) minute; 1

(1); 2(1); 3(2;1-2); 4(2;2-3); 7(2;2-4); 8(4,5) strongly developed and barbed; 9

(131-2); 10(1;1-2); 11(2,3;2-4); 14(1). Mesothorax: hair 1(2-3); 2(2); 3(1); 4(2-

3); 6(131-2); 8(4,5;4-8); 9(6;5-8); 10(1;1-2); 11€1-2); 13(9-14); 14(8-15). Meta-

thorax: hair 1(3;2-3); 2(2;2-3); 3(3,4;2-4); 4(2-3); 6(1); 7(5,6;4-8); 8(10;8-11);

9(3-5 53-8); 10(1;1-2); 11(1;1-2); 13(8-10). Abdomen: Segment I: hair 1(3-5;3-6)

very short; 6(3;2-5); 7(1,2;1-3). Segment II: hair 1(3,4;3-5) very short; 3(2;1-

2) short; 6(3,4;2-4). Segment III: hair 1(3;2-4) strong and long; 3(1;1-2) long

but shorter than hair 1; 6(2;2-3); 13(4;3-5) subequal to hair 1. Segment IV: hair

1(3;3-4) slightly longer than hair 1-III; 3(2;1-3) short; 6(2); 13(3;2-4) shorter

than hair 1. Segment V: hair 1(3) subequal to hair 1-IV; 3(1) strong, slightly

shorter than hair 1; 6(2;1-3); 13(3-4). Segment VI: hair 1(3) slightly longer than

hair 1-III; 6(1-3). Segment VII: hair 1(3). Segment VIII: Comb scales (16-39)

distinctly spatulate and conspicuously fringed; hair 1(2-3;1-3) strong, barbed and

subequal to hair 5; 3(6-8;5-9); 5(2;2-3). Siphon: Pigmentation light yellow to

brown; sculpturing weak. Index about 2.9-3.1. Pecten teeth (5-10) short, with

distinct fringes on distal part of ventral margin. Ventral siphonal tufts (10-12)

usually triple (2-4). Anal Segment: Saddle with unsclerotized indendation from

caudolateral border; pigmentation light yellow; sculpturing weak except caudo-

dorsad. Hair 1(5,6;3-6); 2(4;3-5). Ventral brush with 5 pairs of hairs; all except

the 2 proximal pairs usually double (1-2). Gills usually not much more than 2.0

of dorsal saddle length.

SYSTEMATICS. Several primitive character states and the restricted distribu-

tion indicate that guerreroi is an ancient relict of the Bihaicolus Complex. In

the adults the erect scales of the vertex are dark and the proctiger of the male

genitalia is short and broad and bears a crown of numerous teeth on the para-

proct and the snout of the clasper is short. In the pupa and larva abdominal

hair 6-IJI-V is short and branched, and in the larva head hair 6-C is branched;

these character states are shared with rausseoi and metempsytus. However, it has

a derived feature, unique in the subgenus, in the elongation of the proboscis.

Culex guerreroi is readily separated from the other members of the complex

and group by the male genitalia. Although the elongation of the proboscis in

both sexes is apparently diagnostic of guerreroi, there is considerable variation

in this character but no overlap with other species of the group in my small

sample. I have found no reliable character to separate the pupa of guerreroi from

those of rausseoi and metempsytus as there is considerable variation in the chaet-

otaxy of all 3 species in this stage. The separation of the larvae of guerreroi

and metempsytus on the key characters may be difficult and not entirely reli-

able as the unsclerotized indentation on the anal saddle of guerreroi is quite var-

iable and often obscure and the chaetotaxy of both species shows considerable

individual variation.

Although guerreroi occurs in the same general area in the coastal range of

northcentral Venezuela as rausseoi, the 2 species have been found at different

elevations, guerreroi at 1600-1900 m and rausseoi at 1200-1300 m. There may

36 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

be an ecological barrier separating the 2 species that is associated with the dif-

ference in elevation.

BIONOMICS. The only recorded natural breeding site of guerreroi is a tree-

hole (type collection). The other 5 known collections of immature stages are all

from tin cans, glass jars, a tire, a shoe and a wash basin in a trash dump. Associ-

ated with guerreroi in these artificial containers were: Culex (C.) quinquefasciatus

(1), C. (Lutzia) sp. (1) and Trichoprosopon (Ct.) magnum (1). No adults have

been found in nature.

DISTRIBUTION (fig. 2). Known at present only from northcentral Venezuela,

at elevations of 1600-1900 m.

Material examined: 66 specimens; 7 males, 10 females, 30 larvae, 19 pupae;

14 individual rearings (4 larval, 7 pupal, 3 incomplete).

VENEZUELA. Aragua: Colonia Tovar (VZ 390,394,397-399) [UCLA].

3. Culex (Car. ) rausseoi Cova Garcia, Sutil & Pulido

Figs. 2,9,14,15

1972. Culex (Carrollia) rausseoi Cova Garcia, Sutil and Pulido, 1972:207-209. TYPE: Holotype

male (1951-1) with associated larval and pupal skins, Campamento Rangel, Tejerias,

Aragua, Venezuela, date and collector not specified [MDM].

FEMALE (fig. 9). Wing: 4.1 mm. Proboscis: 2.9 mm. Forefemur: 2.5 mm.

Abdomen: about 2.9 mm. Extremely similar to bihaicolus and metempsytus; di-

agnostic characters as in the key. Differing from bihaicolus in the following fea-

tures. Description based on 24 specimens from near type locality in Venezuela.

Head: Band of broad decumbent scales of vertex broader, scales darker and with

deeper blue iridescence; narrow decumbent scales whitish, less numerous; erect

scales of vertex dark bronzy to black, those of occiput tan; lateral patch with

white scales. Proboscis 1.00-1.20 of forefemur. Palpus 0.20-0.25 of proboscis.

Antenna 0.88-0.91 of proboscis; torus with 10-14 bristles. Thorax: Anterior pro-

montory with 3 or 4 bristles; prescutal bristles 3 or 4. Apn bristles 6-8; ppn with

4-6 posterior bristles but without scales; ppl bristles 3-6; stp with 10-15 bristles

in a curved line, often 1 or 2 light scales cephalad of bristles; pra bristles 3-6;

umep bristles 3-6; Imep bristle sometimes absent. Legs: Knee spots usually more

prominent. Abdomen: Bristles of tergites and sternites somewhat darker. Ster-

nites V-VII usually with some apical dark scales. Cibarial Armature: Cibarial bar

without distinct lateral lobe; broad cibarial teeth 5 or 6, alternating with long

filamentous spicules. Genitalia: Tergite IX with 4-6 lateral bristles; insula with

18-20.

MALE. Wing: 3.0 mm. Proboscis: 2.4 mm. Forefemur: 2.0 mm. Abdomen:

about 2.3 mm. Differentiated from all other members of the group by the very

short palpus, about 0.33 of proboscis. Essentially similar to the female except

for usual sexual differences. Specimens studied: 23 from near type locality in

Venezuela.

MALE GENITALIA (fig. 14). Lobe of tergite IX usually with 8 (5-12) bris-

tles. Sidepiece with scales. Proximal division of subapical lobe with simple broad

base bearing numerous fine simple bristles; stem short, not markedly differenti-

ated from base and bearing several bristles; setae a and b not contorted, a short

and with simple pointed apex, b about twice as long and with recurved apex.

Valencia: Subgenus Carrollia of Culex 37

Distal division developed as a small fingerlike process bearing a single thicken-

ed apical seta with simple attenuate apex. Clasper distinctly tapered distally;

spiniform shorter than snout, latter very strongly developed. Distal part of proct-

iger relatively broad and distinctly rounded apically; paraproct crown usually

with 7 (6-8) teeth; cercal setae absent.

PUPA (fig. 14). Abdomen: 3.65 mm. Trumpet: 0.75 mm. Paddle: 0.85 mm.

Apparently indistinguishable from guerreroi and metempsytus; diagnostic char-

acters from other members of the group as in the key. General chaetotaxy based

on 10 specimens from Venezuela. Cephalothorax: Pigmentation light yellow; sculp-

turing moderate. All hairs distinctly shorter than trumpet except 5-C which is

slightly longer; 1(2;2-4) longer than hair 4; 2(2;2-3) shorter than hair 6; 3(2;

2-3) subequal to hair 6; 4(3,4;2-4) longer than hair 7; 6(2;1-3); 8(1,2) longer

than hair 9; 9(2;1-2). Trumpet: Moderately widened distally; index about 4.1

(3.8-5.0); pinna about 0.25. Pigmentation light brown. Metanotum: Pigmentation

yellow to light brown; sculpturing moderate; hair 10(2,3;1-3) longer than hairs

11,12; 11(2); 12(2,3;1-3). Abdomen: Pigmentation light yellow to brown, an-

terior mesal areas of tergites III-V darker; sculpturing moderate. Segment I: hair

1(35-60). Segment II: hair 1(8-10;6-10) forked; 3(2;1-2); 4(6,7;3-7) short; 5(2;

1-3), subequal to hair 3. Segment III: hair 1(2,3;1-4) strong, reaching caudal

margin of tergite IV; 3(2;2-4) longer than hair 6; 4(2,3;1-4); 5(2,3;2-4). Seg-

ment IV: hair 1(3,4;2-5) surpassing level of rudimentary spiracle V; 3(3,4;3-9)

shorter than hair 6; 4(2;2-3); 5(1;1-2) strong, reaching level of rudimentary spi-

racle VI. Segment V: hair 1(2,3;1-4) reaching level of rudimentary spiracle VI;

3(3,4;2-5) shorter than hair 6; 4(4-6;2-6) shorter than hair 5; 5(1;1-2) strong,

reaching level of rudimentary spiracle VII. Segment VI: hair 1(1;1-2) shorter than

hair 6; 3(2;2-4); 4(4,5;3-5) shorter than hair 3; 5(4,5;3-10). Segment VII: hair

1(1,231-3); 3(3;2-4) subequal to hair 6; 4(2;2-3), shorter than hair 1; 5(2,3;2-6)

very short; 6(4,5;4-6); 9(5,6;3-9) strong, barbed and shorter than hair 9-VIII.

Segment VIII: hair 4(2;1-3); 9(8;6-9) strong and barbed.

LARVA (fig. 15). Head: 1.00 mm. Siphon: 0.90 mm. Anal Saddle: 0.25 mm.

Very similar to species 3a., Panama form; diagnostic characters as in the Key.

General chaetotaxy based on 10 reared specimens from Venezuela. Head: Inte-

gument light yellow to brown; collar brown. Mental plate with 10 lateral teeth.

Hair 4(4;3-5); 5(5,6;5-10); 6(2;2-3); 7(6-8;5-8); 8(1;1-2) subequal to 7-C; 9(4,5:

3-7) shorter than hair 8; 10(2;1-2); 11(3,4;3-5); 12(2;2-3); 14(1,2;1-3) more dis-

tant from anterior border of head capsule than in other species of group; 15

(332-4). Antenna: About 0.4 of head length; hair 1(3,4;2-4). Thorax: Prothorax:

hair 0(12-14;10-14) short; 1(1); 2(1); 3(1,2;1-3) relatively short; 4(2); 7(2;2-3);

8(4,5;4-6) moderately developed, barbed; 9(1;1-2); 10(1); 11(4;3-4); 14(1). Meso-

thorax: hair 1(2,3;2-4); 2(2,3;2-5); 3(1); 4(3;2-4); 8(5-7;5-9); 9(5,6;5-7); tT

13(10-18); 14(24-30). Metathorax: hair 1(4,5;2-6); 2(3,4;2-4); 3(5,6:4-7); 4(2,3:

2-4); 6(1;1-2); 7(6-9;5-10); 8(10-14); 9(2,3;2-5); 11(2;2-3); 13(8-16). Abdomen:

Segment I: hair 1(5,6;3-8) minute; 6(2,3); 7(1;1-2). Segment II: hair 1(4-6;3-6)

Short; 3(2,3;2-4) short; 6(3;2-3). Segment III: hair 1(3,4;3-8) short to moderate;

3(1,2;1-3) longer and stronger than hair 1; 6(2,3;1-4); 13(3,4;3-5) relatively short

and weak but longer than hair 1. Segment IV: hair 1(3,4;3-5) moderately long

but rather weak; 3(3;3-5) usually forked, relatively short; 6(1,2;1-3); 13(3,4;3-

5) subequal to hair 1. Segment V: hair 1(3;2-4) longer than hair 1-IV; 3(1) sub-

equal to hair 1; 6(1); 13(3;3-5) shorter than hair 1. Segment VI: hair 1(3;2-4)

subequal to hair 1-IV; 6(1). Segment VII: hair 1(3;3-4). Segment VIII: Comb

38 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

scales (16-28) only slightly dilated and lightly fringed; hair 1(3;3-4) about 0.5

or less of hair 2; 3(6,7;6-9); 5(3,4;2-4) distinctly longer than hair 1. Siphon:

Pigmentation light yellow to brown; sculpturing weak. Index about 2.8-3.2. Pec-

ten teeth (3-7) elongate, spinelike and with a few indistinct spicules on ventral

margin. Ventral siphonal tufts (10;10-14) usually quadruple (3-5); proximal ac-

cessory tuft (lad) in basal 0.5 of siphon, distal varied in position but seldom sub-

apical. Anal Segment: Saddle pigmentation light yellow to brown; sculpturing

weak except caudodorsad. Hair 1(4,5;3-6); 2(3,4;3-5). Ventral brush with 6 pairs

of hairs all except the proximal single and without barbs. Gills about 3.0 of dor-

sal saddle length, pointed.

SYSTEMATICS. Culex rausseoi is another relict species in the Bihaicolus Com-

plex. Its distribution is nearly as restricted as that of guerreroi and it possesses

all the primitive character states of the latter except for the snout of the clasper

which is very strongly developed. Another primitive character state (shared with

sp. 3a) is the retention of 6 pairs of hairs in the ventral brush of the larva; the

branching of these hairs, however, is reduced. A unique and very striking feature

of rausseoi is the very short palpus of the male, which is about half as long as in the

other species of the group.

Culex rausseoi is readily separated from the other members of the complex

and group by the male genitalia. Although the males cannot be confused with

any other species of the complex, the females are very similar to bihaicolus and

metempsytus and the key characters may not be completely reliable. The pupa

cannot be differentiated from those of guerreroi and metempsytus. The larva is

separated from all described species of the complex and group by the ventral

brush but it is superficially very similar to that of the Panama form (sp. 3a).

As noted under guerreroi, there is no real sympatry between these 2 species

as rausseol occurs at a lower elevation (1200-1300 m) and there may also be

an ecological barrier separating them.

BIONOMICS. The immature stages of rausseoi have been found only in fallen

palm spathes (5) where they were associated with bihaicolus (5), Anopheles (An.)

eiseni (2), Limatus asulleptus (1), Trichoprosopon (T.) digitatum (1) and Wyeo-

myia felicia (1). No adults have been collected in nature.

DISTRIBUTION (fig. 2). Known at present only from northcentral Venezuela,

at elevations of 1200-1300 m.

Material examined: 257 specimens; 39 males, 38 females, 133 larvae, 47 pupae;

46 individual rearings (30 larval, 12 pupal, 4 incomplete).

VENEZUELA. Aragua: Campamento Rangel (1951-5) [MDM]. Rancho Grande (VZ 156,

157,255,303,304,311) [UCLA].

3a. Culex (Car.) sp., Panama form

Fig. 2

This form is known from a single collection of larvae in Cerro Mali, Tacarcuna

Mts., Darien Province, Panama (PA 929; TC-2) [UCLA]. Nearly all the larvae

in this collection have 6 pairs of hairs in the ventral brush, the majority of these

being long and simple. In this respect they resemble rausseoi, but they differ

from this species in having accessory siphonal hair Jad in the distal half of the

siphon and in other details of chaetotaxy. It is very probable that these larvae

represent a distinct undescribed species. In the same collection there are 2 indi-

Valencia: Subgenus Carrollia of Culex 39

vidual rearings of males (929-107,112) of more or less typical metempsytus and

a few larvae with the ventral brush tending toward the condition in the latter,

suggesting introgression between the 2 species. This form cannot be placed in

either complex at this time. It may prove to be an annectent relict between the

2 complexes or a relict of either complex.

4. Culex (Car.) metempsytus Dyar

Figs. 2,9,16,17

1921. Culex (Carrollia) metempsytus Dyar, 1921:154. TYPE: Lectotype male, Alajuela, Ala-

juela, Costa Rica, 4 July 1921, A. Alfaro [USNM, 24863; designation by Stone and

Knight, 1957:53].

Culex (Carrollia) metempsytus of Bonne and Bonne-Wepster (1925:204); Dyar (1925:176); Ed-

wards (1932:221); Galindo, Carpenter and Trapido (1951:126); Galindo and Blanton (1955:

73); Stone and Knight (1957:53); Stone, Knight and Starcke (1959:283); Belkin, Schick and

Heinemann (1965:13-14).

Culex (Carrollella) metempsytus of Dyar (1928:282-283).

Culex (Carrollia) metempsytus in part of Lane (1939:82; 1953:510-511).

FEMALE (fig. 9). Wing: 3.2 mm. Proboscis: 2.3 mm. Forefemur: 1.8 mm.

Abdomen: about 2.5 mm. Extremely similar to bihaicolus and rausseoi; diag-

nostic characters as in the key. In general similar to bihaicolus and differing from

it in the following features. Description based on 34 specimens from near type

locality in Costa Rica and 16 specimens from Panama. Head: Broad decumbent

scales of vertex somewhat darker; narrow decumbent scales whitish; erect scales

tan to bronzy; lateral patch white. Proboscis 1.08-1.30 of forefemur. Palpus 0.19-

0.20 of proboscis. Antenna 0.90-0.95 of proboscis. Torus with 6-10 setae. Tho-

rax: Mesonotal integument dark brown to black. Prescutal bristles 2-4. Midlobe

of scutellum with 4-6 long bristles. Apu with 5-9 bristles; ppn with 2 or 3 nar-

row dark scales and 3 or 4 posterior bristles; stp with 8-12 bristles in curved

row; pra bristles 3 or 4; umep bristles 4-6; Jmep bristle sometimes absent. Legs:

Light scaling of hindfemur usually more extensive, extending to near light me-

tallic streak on anterior surface but usually not merging with it. Knee spots some-

what more prominent. Abdomen: Bristles of tergites and sternites somewhat dark-

er. Sternites IV-VII with conspicuous dark apical bands. Cibarial Armature: Ci-

barial bar without any indication of a lobe on lateral flange; cibarial teeth a ser-

ies Of 10 expanded on apex alternating with thin filamentous processes; cibarial

dome with rounded minutely serrate denticles. Genitalia: Tergite IX with 2-4

lateral bristles; insula with 16-20.

MALE. Wing: 2.9 mm. Proboscis: 2.1 mm. Forefemur: 1.8 mm. Abdomen:

about 2.6 mm. Very similar to bihaicolus; diagnostic characters as in the key.

Similar to the female except for usual sexual characters. Proboscis 1.09-1.20 of

forefemur. Palpus 0.76-0.83 of proboscis. Antenna 0.75-0.76 of proboscis. Spec-

imens studied: 26 from near type locality in Costa Rica and 9 from Panama.

MALE GENITALIA (fig. 16). Lobe of tergite IX usually with 4 (3-8) bris-

tles. Sidepiece without scales. Proximal division of subapical lobe with basal part

narrow, not differentiated from very long stem which bears relatively few short

bristles; setae a and b contorted, b forked distally. Distal division not developed.

Clasper not markedly tapered distally; spiniform and snout subequal. Distal part

40 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

of proctiger very narrow and distinctly tapering distally, paraproct crown usu-

ally with 5 (4-7) teeth; cercal setae absent.

PUPA (fig. 16). Abdomen: 3.30 mm. Trumpet: 0.60 mm. Paddle: 0.85 mm.

Apparently indistinguishable from rausseoi and guerreroi; diagnostic characters from

other members of the group as in the key. General chaetotaxy based on 10 spec-

imens from Panama. Cephalothorax: Pigmentation light yellow; sculpturing mod-

erate. All hairs distinctly shorter than trumpet except 5-C which is slightly long-

er; 1(2,3) longer than hair 7; 2(3;2-4) subequal to hair 6; 3(2;2-3) subequal

to hair 4; 4(3,4;2-4) shorter than hair 7; 5(2,3;2-4); 6(1,231-3); 7(3,4;2-5); 8(2;

1-2) longer than hair 9; 9(2;2-4). Trumpet: Moderately widened distally; index

about 6.5 (5.5-7.0); pinna about 0.25. Pigmentation yellow to brown. Metano-

tum: Pigmentation light yellow; sculpturing moderate; hair 10(2;2-3) longer than

hairs 11,12; 11(2;2-4); 12(2,3;2-6). Abdomen: Pigmentation light yellow, anter-

ior mesal areas of tergites III-V darker; sculpturing moderate. Segment I: hair

1(36-65). Segment II: hair 1(5,6;3-10) surpassing level of rudimentary spiracle

III; 3(2,3); 4(6-8;5-10); 5(2;2-3) subequal to hair 3. Segment III: hair 1(1,2)

nearly reaching caudal margin of tergite IV; 3(2,3); 4(3,4;3-8); 5(2;2-5) very short.

Segment IV: hair 1(1,2;1-3) longer than hair 3; 3(4,5;3-9) subequal to hair 6;

4(2;1-2); 5(1;1-2) moderately long, surpassing caudal margin of tergite V. Seg-

ment V: hair 1(1,2;1-4) subequal to hair 4; 3(2,3;1-4) twice as long as hair 1;

4(5,6;5-8); 5(1;1-2) moderately long, surpassing caudal margin of tergite VI. Seg-

ment VI: hair 1(1;1-2) longer than hair 4; 3(3,4;2-4); 4(4-6;4-7); 5(3,4;2-6) short.

Segment VII: hair 1(2;1-3) subequal to hair 4; 3(4,5;3-6) subequal to hair 6;

4(3,4;2-5); 5(4,5;3-6) very short; 6(6,7;5-8); 9(2,3;2-6) strong, barbed and sub-

equal to hair 9-VIII. Segment VIII: hair 4(2;1-3); 9(3,4;2-6) strong and barbed.

LARVA (fig. 17). Head: 0.85 mm. Siphon: 0.85 mm. Anal Saddle: 0.20 mm.

Very similar to guerreroi; diagnostic characters as in the key. General chaetotaxy

based on 6 reared and 2 whole specimens from Panama. Head: Integument light

yellow; collar dark brown. Mental plate with 8-10 lateral teeth. Hair 4(3-5;3-7);

5(7-10); 6(2;2-4); 7(7,8;6-10); 8(3,4;3-5) short; 9(4,5;4-6) subequal to hair 6; 10

(332-4); 11(3;2-4); 12(2,3;2-4); 14(1,2;1-3); 15(3;2-4). Antenna: About 0.41 of

head length; hair 1(1;1-2). Thorax: Prothorax: hair 0(10-18) .very short; 1(1);

2(1); 3(2;1-2) relatively short; 4(2;2-3); 7(3;2-4); 8(4,5;2-5) moderate, barbed;

9(1;1-2); 10(€1); 11(4,5;3-6); 14(1;1-2). Mesothorax: hair 1(3,4;3-5); 2(2;1-3); 3

(151-2); 4(5,6;3-7); 8(5,6;4-6); 9(4-5); 11(1); 13(10-15); 14(18-22). Metathorax:

1(333-5); 2(3-5;2-5); 3(10-12;8-12); 4(3;1-3); 6(1-2); 7(4,5;3-6); 8(10-12;10-14);

9(1;1-2); 11(1); 13(8;6-10). Abdomen: Segment I: hair 1(8-12) minute; 6(2;2-

3); 7(1). Segment II: hair 1(4-6;3-6) minute; 3(5-8;3-8) usually short; 6(3;2-4).

Segment III: hair 1(3,4;3-6) strong, moderately long; 3(4,5;3-5) weaker and short-

er than hair 1; 6(2,3;2-6); 13(2,3;2-4) stronger and longer than hair 1. Segment

IV: hair 1(3,4;3-6) strong, slightly longer than hair 1-III; 3(5-8;2-8) usually short;

6(2,3;2-4); 13(3;3-6) subequal to hair 1. Segment V: 1(3;3-5) subequal to hair

1-IV; 3(1,2;1-3) usually slightly shorter than hair 1; 6(2,3;2-4); 13(2-4;2-6) sub-

equal to hair 1. Segment VI: hair 1(1,2;1-3) subequal to hair 1-V; 6(1,2;1-4).

Segment VII: hair 1(3,4;3-5) rather short, barely exceeding base of siphon. Seg-

ment VIII: Comb scales (16-28) moderately spatulate and with distinct marginal

fringes; all hairs relatively short; 1(4-6;4-7) about 0.5 or more of hair 2; 2(2;1-

3); 3(5,6;4-9); 4(1;1-2); 5(1-2) subequal to hair 1. Siphon: Pigmentation light

brown; sculpturing weak. Index about 2.2-3.0. Pecten teeth (4-7) spinelike, with

distinct fringes on ventral margin. Ventral siphonal tufts (8-10) usually triple (2-

Valencia: Subgenus Carrollia of Culex 41

4). Anal Segment: Saddle without distinct emargination; pigmentation light brown;

sculpturing weak except caudodorsad. Hair 1(4,5;4-6) very short; 2(3,4;2-4). Ven-

tral brush with 5 pairs of relatively short hairs, all except the proximal usually

double. Gills sausage-shaped, about 3.5 of dorsal saddle length.

SYSTEMATICS. Culex metempsytus differs conspicuously from its close rela-

tive, infoliatus, in the retention of the primitive character state of branched abdom-

inal hair 6-III-V in both the larva and the pupa (shared with the relict species of

the Bihaicolus Complex). For this reason I consider it to be closer to the ancestral

stock of the Infoliatus Complex.

Several authors misidentified adults of infoliatus and metempsytus in the past

(see taxonomic references under infoliatus). Although the male genitalia of me-

tempsytus and infoliatus are very similar, the minor differences used in the key

are quite constant and diagnostic. The adults of these 2 species are also readily

separated by the hindfemoral markings but some difficulty may be encountered

in distinguishing females of metempsytus by the key characters from those of

bihaicolus and particularly rausseoi. I have found no reliable characters to sep-

arate metemspytus from guerreroi and rausseoi in the pupal stage. As noted under

guerrerol, the key characters distinguishing the larva of this species from metemp-

sytus may not be reliable.

There is a great deal of individual variation in the chaetotaxy of the larva and

pupa but I have not found any significant population or ecological differentia-

tion in the large sample from the wide geographical and altitudinal range and

the various breeding sites.

Culex metempsytus is apparently completely allopatric from infoliatus and its

known distribution is entirely within the central part of the range of bihaicolus.

It has been found breeding in association with bihaicolus as well as sp. 3a. There

is a suggestion of introgression between the latter and metempsy tus.

BIONOMICS. The recorded breeding sites of metempsytus are fallen palm spathes

(11), broken or cut bamboo (5), treeholes (2) and wooden bowl traps (2). Assoc-

iated with metempsytus were 3 species of Carrollia: bihaicolus (2), sp. 3a (1)

and bonnei (1). The other associates included a representation of several con-

tainer-breeding groups: Aedes (P.) homoeopus, A. (F./ terrens (2); Culex (C.) mol-

lis (1); Haemagogus (H.) iridicolor (1); Limatus asulleptus (2), L. durhamii (1);

Sabethes sp. (1). All the adults I have seen have been reared and there are no

published records of adults collected in nature.

DISTRIBUTION (fig. 2). From Guatemala through Central America to Pacific

drainage in central Colombia, at elevations of 30-2000 m.

Material examined: 1,268 specimens; 119 males, 162 females, 714 larvae, 273

pupae; 257 individual rearings (136 larval, 75 pupal, 46 incomplete).

COLOMBIA. Valle: Rio Raposo (COL 65) [UCLA].

COSTA RICA. Alajuela: Alajuela, A. Alfaro (24863) [USNM]. Rio Barranca N of San Ramon

(CR 343) [USNM]. San Jose: Lourdes (CR 491,493) [UCLA].

GUATEMALA. Chimaltenango: Yepocapa (GUA 136,138) [UCLA].

PANAMA. Chiriqui: El Volcan, W.H. Komp [UCLA]. Rio Chiriqui Viejo, WH. Komp [USNM].

Darien: Alturas de Nique (PA 1057) [UCLA]. Cerro Mali (PA 347,355,364,376,377,382,929,931)

[UCLA].

5. Culex (Car.) infoliatus Bonne-Wepster & Bonne

Figs. 2,9,18,19

1920. Culex (Carrollia) infoliata Bonne-Wepster and Bonne, 1920:170-171. TYPE: Lectotype

42 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

male (4496) with genitalia slide (G.g. 2; BB 690), Dam, Surinam, Jan 1919 [ITH; desig-

nation by Belkin, 1968b:16].

Culex (Carrollia) infoliatus of Bonne and Bonne-Wepster (1925:205-207); Edwards (1932:221);

Senevet and Abonnenc (1939:125-126; 1958:288-290); Levi-Castillo (1925:553; 1953a:35);

Lane (1939:82; 1953:510); Stone, Knight and Starcke (1959:282); Fauran (1961:44); Belkin,

Schick and Heinemann (1965:65-66); Cova Garcia, Sutil and Rausseo (1966a:28,115; 1966b:

42-44,141,235 344); Belkin (1968b:16).

Culex (Carrollella) infoliatus of Dyar (1928:283-284); Boshell Manrique (1938:416).

Culex (Carrollella) infoliatus in part of Komp (1936:64; 1956:39).

Culex (Carrollia) bihaicolus of Cerqueira (1961:163); Forattini, Rabello and Cotrim (1970:36).

Culex (Carrollia) bihaicolus in part of Lane (1939:81; 1953:509).

Culex (Carrollella) bihaicolus of Antunes (1936:74); Boshell Manrique (1938:416).

Carrollia bihaicolus of Townsend (1934:490).

Carrollella bihaicolus of Martini (1931:217); Morales-Ayala (1971:143).

Culex (Carrollia) metempsytus of Levi-Castillo (1952:533; 1953a:35).

Culex (Carrollia) metempsytus in part of Lane (1939:82; 1953:510-511).

Culex (Carrollella) metempsytus of Komp (1936:64); Boshell Manrique (1938:416).

FEMALE (fig. 9). Wing: 3.6 mm. Proboscis: 2.3 mm. Forefemur: 2.1 mm.

Abdomen: about 2.1 mm. Differentiated from other species of the group by

the very long narrow light metallic streak of the hindfemur and the more ex-

tensive pale scaling of this segment. In general similar to bihaicolus and differ-

ing from it in the following features. Description and measurements based on

4 pinned specimens from French Guiana. Head: Broad decumbent scales of ver-

tex darker and with deeper blue iridescence;. lateral patch white. Proboscis 1.10-

1.20 of forefemur. Palpus about 0.17 of proboscis. Antenna 0.76-0.80 of pro-

boscis; torus with only 4-6 setae. Thorax: Mesonotal integument light brown,

not as strongly contrasting with pleuron. Anterior acrostichals often developed

to near level of scutal angle and rarely distad. Prescutal bristles 3 or 4; all meso-

notal bristles lighter. Midlobe of scutellum with only a few scales, sometimes

whitish. Apn bristles 12-14; ppn bristles 3 or 4, absent from upper part; upper

stp bristles 5, lower 6; pra bristles 6-10; upper mep bristles 5 or 6. Legs: Mid-

femur extensively pale ventrally on both surfaces. Hindfemur more extensively

pale; light metallic streak inconspicuous, very narrow and long, at least 0.25 of

femur length, and confluent with the light scaling anteriorly and/or posteriorly.

Abdomen: Sternites IV-VII with apical band of dark scales. Cibarial Armature:

Cibarial bar with less conspicuous lobe on lateral flange; cibarial teeth 10 or 11;

cibarial dome with rounded denticles. Genitalia: Tergite IX with 4 lateral bris-

tles; insula with about 20.

MALE. Wing: 3.5 mm. Proboscis: 2.4 mm. Forefemur: 2.2 mm. Abdomen:

about 2.5 mm. Differentiated from other species of the group and from bihai-

colus by the same features as the female. Proboscis 1.09-1.10 of forefemur. Pal-

pus 0.72-0.77 of proboscis. Antenna 0.85-0.91 of proboscis. Specimens studied:

8 from French Guiana. Measurements from dry specimens.

MALE GENITALIA (fig. 18). Lobe of tergite [IX usually with 5 (4-6) bris-

tles. Sidepiece without scales. Proximal division of subapical lobe with simple

broad base bearing numerous fine simple bristles; stem elongate, with only a

few bristles; setae a and b subequal, recurved apically but not contorted. Distal

division not developed. Clasper not markedly tapered distally; spiniform longer

than snout. Distal part of proctiger narrow, more or less parallel-sided; paraproct

crown usually with 6 (4-6) teeth; 1 pair of cercal setae present.

Valencia: Subgenus Carrollia of Culex 43

PUPA (fig. 18). Abdomen: 3.4 mm. Trumpet: 0.65 mm. Paddle: 0.75 mm.

Very similar to bihaicolus; diagnostic characters as in the key. General chaeto-

taxy based on 10 reared specimens from French Guiana. Cephalothorax: Pig-

mentation light yellow; sculpturing weak, without definite pattern. All hairs dis-

tinctly shorter than trumpet except 5-C which is subequal; 1(1;1-2); 2(2,351-3)

very short; 3(2,3;1-3) about 0.5 of hair 4; 4(2;2-3) subequal to hair 7; 5(2,1-3);

6(1;1-2) subequal to hair 2; 7(2-4); 8(1;1-2) longer than hair 9; 9(1-2). Trum-

pet: Moderately widened distally; index about 6.5 (5.0-6.5); pinna about 0.33.

Pigmentation light brown. Metanotum: Pigmentation light yellow; sculpturing weak;

hair 10(1-2) longer than hairs 11,12; 11(1-2); 12(1). Abdomen: Pigmentation

yellow, anterior mesal areas of tergites IJI,I[V slightly darker; sculpturing weak.

Segment I: hair 1(26-57). Segment II: hair 1(2-4;2-5) forked and longest hair

of segment; 3(1-2) subequal to hair 5; 4(3,4;3-6) short; 5(1;1-2). Segment III:

hair 1(1,2) strong, finely barbed, almost reaching caudal margin of tergite IV;

3(1-2) about 0.5 of hair 1; 4(2,3;1-4) short; 5(1,2;1-3) shorter than hair 3. Seg-

ment IV: hair 1(1;1-2) slightly longer than hair 3; 3(3,4;2-6); 4(3,4;2-5); 5(1)

strong, reaching level of rudimentary spiracle VI. Segment V: hair 1(1;1-2) long-

er than hair 3; 3(1,2;1-4); 4(3,4;2-6); 5(1) strong, reaching level of rudimentary

spiracle VII. Segment VI: hair 1(1;1-2) longer than hair 4; 3(23;1-2); 4(2,3;2-4);

5(1) strong and longer than hair 3. Segment VII: hair 1(1,2;1-3) fine and longer

than hair 4; 3(2,3;1-3); 4(1,2;1-3); 5(3;1-4) subequal to hair 6; 6(3,4;1-5); 9(Q,

3;1-3). Segment VIII: hair 4(1) shorter than hair 1-VII; 9(3;2-5) shorter than

paddle.

LARVA (fig. 19). Head: 0.95 mm. Siphon: 0.95 mm. Anal Saddle: 0.25 mm.

Very similar to bihaicolus; diagnostic characters as in the key. General chaeto-

taxy based on 5 reared and 5 other specimens from French Guiana. Head: In-

tegument light yellow; collar dark brown. Mental plate with 9,10 lateral teeth.

Hair 4(2;2-3); 5(6,7;5-8); 6(2;2-3); 7(6-8;4-8); 8(1;1-2) subequal to hair 7; 9(4;

3-4) shorter than hair 8; 10(131-2); 11(2,3;1-4); 12(1-2); 14¢€1;1-2); 15(3;2-4).

Antenna: About 0.31 of head length; hair 1(2;1-2). Thorax: Prothorax: hair 0

(8-10;6-10) minute; 1(1); 2(1); 3(1,2;1-3) relatively short; 4(2;1-2); 7(2;1-3); 8

(3,4;2-4) relatively short and weak; 9(1;1-2); 10(€1); 11(3,4;2-4); 12(1); 14(1).

Mesothorax: hair 1(2;2-3); 2(2,3); 3(1); 4(2;2-4); 8(7-9;5-9); 9(5,6;4-6); 10(1;1-

2); 11(€1); 13(8-12); 14(10-19). Metathorax: hair 1(2;1-2); 2(2;1-2); 3(3,4;3-6);

4(3;2-3); 7(6,7;6-9); 8(11,12;9-15); 9(2,3;1-5); 11(1); 13(5,6;5-7). Abdomen: Seg-

ment I: hair 1(1,2;1-4) minute; 6(2;2-3); 7(1). Segment II: hair 1(1;1-2) minute;

3(2,3) short; 6(2;2-3). Segment III: hair 1(1;1-2) moderate, weak; 3(2;1-2) short;

6(1); 13(2;2-3) stronger and longer than hair 1. Segment IV: hair 1(2;1-3) strong

and long; 3(3,4;2-4) short and weak; 6(1); 13(2,3) strong, subequal to hair 1.

Segment V: hair 1(2;1-3) strong, shorter than hair 1-IV; 3(1) long, weak; 6(1);

13(2,3) longer than hair 1. Segment VI: hair 1(2;2-3) intermediate in length be-

tween hairs 1-IV,V; 6(1). Segment VII: hair 1(2;2-3) stronger and longer than

hair 1-[V. Segment VIII: Comb scales (31-62) spatulate and conspicuously fringed;

hair 1(4,5;4-6) very short, about 0.33 of hair 2; 3(6,7;5-7); 5(1;1-2) more than

2.0 of hair 1. Siphon: Pigmentation light yellow to brown; sculpturing weak. In-

dex about 2.7-3.7. Pecten teeth (6-15) short, with conspicuous denticles on ven-

tral margin. Ventral siphonal tufts (10-12) usually quadruple (3-7). Anal Seg-

ment: Saddle shallowly emarginate laterally; pigmentation light yellow to brown;

sculpturing weak, only slightly more prominent caudodorsad. Hair 1(1,2;1-3); 2

(5,6;4-6). Ventral brush with 5 pairs of hairs, all except proximal pair usually

triple (2-4). Gills more than 3.0 of dorsal saddle length.

44 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

SYSTEMATICS. Culex infoliatus is interpreted here as the derived species of

its complex, whose only other member is metempsytus, because of the reduc-

tion in the branching of hair 6 on abdominal segments III-V of both the larva

and pupa (shared with the nominate species of the Bihaicolus Complex) and the

long metallic streak of the hindfemur of the adults (unique in the group). It is

probably the dominant species of the complex as suggested by its wide geograph-

ical range. 3 !

Adults of infoliatus have been misidentified as bihaicolus and metempsytus by

several authors in the past (see taxonomic references above). However, they are

readily distinguished from all members of the group by the ornamentation of

the hindfemur. As noted under metempsytus the minor key characters for the

differentiation of the male genitalia of these 2 species appear to be reliable. The

separation of metempsytus and bihaicolus in the pupal stage by the key char-

acters may not be entirely reliable but it appears to be satisfactory in the larva. 3

In the limited sample available I have found no significant population differ-

ences in any stage. Although the present distribution of infoliatus appears to be

primarily Amazonian, its original center may have been the Guiana Shield with

subsequent dispersal into the Amazon and upper Orinoco drainages. The absence

of infoliatus from Trinidad is probably real as container-breeding habitats have

been very extensively sampled on this island.

BIONOMICS. I have examined collections of immature stages only from the

Villavicencio area in Colombia and from Surinam and French Guiana. These were

made in treeholes (4), fallen palm spathes (3) and tin cans (2). The only recorded

associates were in a tin can in French Guiana: a species of Carrollia (bonnei),

C. (Microc.) stonei and Wyeomyia aporonoma. Because of the wide distribution

of infoliatus, the range of both breeding sites and associates is probably much

greater. Antunes (1937:79) reports the capture of 2 males resting on plants in

the Villavicencio area and Cerqueira (1961:136) an unspecified number of fe-

males attracted to human or animal bait in Manaus, Brazil.

DISTRIBUTION (fig. 2). Amazon and Orinoco basins and the Guianas at ele-

vations up to 600 m.

Material examined: 119 specimens; 29 males, 32 females, 45 larvae, 13 pupae;

6 individual rearings (5 larval, 1 incomplete).

BRAZIL. Para: Boa Vista, C.H.T. Townsend [USNM].

COLOMBIA. Meta: Retiro, W.H. Komp (KO 10-28) [UCLA]. Restrepo, W.H. Komp (1019,

1025,1040) [USNM] , (KO 31-36, 31-39) [UCLA]. Villavicencio (CV 1002, C-30) [UCLA].

FRENCH GUIANA. Inini: Maroni [UCLA]. Saut Tigre (FG 179,180) [UCLA].

PERU. Huanuco: Tingo Maria, Humbleton [USNM].

SURINAM. Dam (BB 621,739) [USNM] ; BB 690, lectotype [ITH].

Additional Records From the Literature

BRAZIL. Amazonas: Manaus. Para: Santarem (Cerqueira, 1961:136).

ECUADOR. Napo-Pastaza: Tena. Ila. Puyo (Levi-Castillo, 1952:553).

FRENCH GUIANA. Guyane: Cabassou. Crique Anguille. Inini: Balorou. Maroni. St. Elie.

St. Nazaire. Saut Canori. Saut Machicou. Saut Tigre (Fauran, 1961:44).

PERU. Junin: Chanchamayo (Martini, 1931:217; Morales Ayala, 1971:143).

VENEZUELA. Locality not specified (Anduze, 1941a:17).

IRIDESCENS GROUP

FEMALES. Head: Band of broad decumbent scales of vertex very narrow and

Valencia: Subgenus Carrollia of Culex 45

sometimes broken on midline (Urichii Subgroup) or very broad; narrow decum-

bent scales of vertex and occiput very numerous to apparently absent; erect scales

moderately long to short, from very numerous and reaching orbital bristles to

relatively few and restricted to posterior part of vertex and occiput. Proboscis

subequal to forefemur or slightly longer. Antenna subequal to proboscis to dis-

tinctly longer. Thorax: Mesonotal integument dark brown to black, vestiture of

dark bronzy to dark golden scales moderate to very dense, scales linear to nar-

row curved, latter more numerous in prescutellar space; inner dorsocentral line

completely scaled or unscaled at anterior end (Urichii Subgroup). Scutellar

scales linear to broad. Acrostichals absent except a few at anterior promontory.

Anterior and posterior dorsocentral bristles absent; prescutellars well developed

and numerous (Urichii Subgroup) or reduced; prescutals present (Urichii Sub-

group) or absent. Pleural integument always darkened to near base of coxae and

with a broad diagonal pruinose stripe from ppn to metapleuron. Upper stp bris-

tles absent or a few weak ones not forming curved row to lower bristles (Urichii

Subgroup). Legs: Hindtarsus entirely dark or with a broad basal light ring on

segment 4. Wing: Plume scales moderately long and narrow but not linear on

Rs, R243 and M; very broad and not differentiated from squame scales on R, and

R;. Abdomen: Pregenital segments more or less distinctly compressed; tergites

without (Urichii Subgroup) or with basal dorsal light bands. Cibarial Armature:

Denticles of cibarial dome triangular to rounded.

MALES. General characters as in the females. Proboscis subequal to forefemur

or distinctly longer. Antenna and palpus subequal to proboscis or distinctly short-

er. Anterior claw of foretarsus with tooth, that of midtarsus with or without a

tooth (Urichii Subgroup).

MALE GENITALIA. Segment VIII: Tergite with median emargination; with

3-6 or more rows of strong setae mixed with scales. Sternite with 1 complete

row of setae. Segment IX: Tergite more or less strongly sclerotized dorsally;

lobes widely separated or approximated, always flattened. Sternite short, poorly

sclerotized. Sidepiece: Conical to ovoid. Apicosternal process varied. Subapical

Lobe: Accessory division present or absent. Distal division small and widely sep-

arated from proximal or represented by a small protuberance or apparently com-

pletely absent. Clasper: Complex, with a more or less distinct head differenti-

ated from basal part; shorter to longer than sidepiece; moderately to strongly

curved dorsad; shape varied. Preapical lobe always developed and bearing seta

b which is variously developed; seta a developed as normal thickened spiniform

or hairlike. Crest ridged or a flat sclerite. Lateral Plate: Relatively narrow; ex-

ternal spine not developed; sternal spine present or absent; caudal process always

developed. Proctiger: Paraproct sclerotization with mesal sternal lobe.

PUPAE. Cephalothoracic hair 5-C varied in development and length. Trumpet

varied in length and development of pinna. Abdominal hair 2-II mesad of 3-II;

hair 3-VI laterad of 1-VI; hair 3-II usually strong; 1-IHII always weak and 3-Ill

always strong and longest hair of segment; 5-VI frequently strong.

LARVAE. Head capsule with anterior dorsal surface flattened so that hairs

5,6-C appear posterior to antennal base from dorsal aspect; integument with or

without sculpturing visible at 100X; hair 9-C removed caudad to near collar,

more or less stellate; 12,14-C varied in position. Maxillary hair 5-MP greatly thick-

ened and brush-tipped. Anterior part of prothorax with or without spicules; pro-

thoracic hairs 0,8-P varied in development; mesothoracic hair 14-M and meta-

thoracic hair 13-T varied in development. Abdominal hair 1-II minute to strong-

46 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

ly developed; 3-VI short, dendritic; 11-I dendritic. Comb scales elongate, spine-

like and arranged in 1 more or less regular row, rarely a few scales displaced in-

to second row; hair 1-VIII very short, multiple. Integument of siphon with or

without elongate spicules visible at 100X; pecten teeth scalelike or spinelike; ven-

tral siphonal tufts 4-8 pairs; siphon index varied. Hair 1-X single or branched;

2-X fan-shaped or asymmetrical; ventral brush varied.

DISCUSSION. Nearly all the character states listed above in the diagnosis of

all the stages of the Iridescens Group are derived. However, some primitive fea-

tures have been retained in several lines of the group. The most significant diag-

nostic features of the group are indicated in the keys. The group does not oc-

cur as far north as the Bihaicolus Group and extends southward beyond the

Amazon Basin into southern Brazil and adjacent northeastern Argentina.

There are 2 very distinct principal phyletic lines in the group: the compact

Urichii Subgroup of 2 very similar species and the complex Iridescens Subgroup

of 9 named species, 2 unnamed forms and possible 2 or 3 additional species.

For the diagnostic features and the systematics see the discussions under the re-

spective subgroups.

The 2 subgroups are sympatric in the northern part of the range of the group

and only the Iridescens Subgroup extends southward of the Amazon Basin.

The group as a whole shows nearly the entire range of breeding sites known

for the subgenus in its few widespread species but includes many species or forms

with restricted distribution and breeding sites. It has the same range of breed-

ing associates. Only a few adults have been collected in nature, principally in

the Urichii Subgroup.

URICHIT SUBGROUP

FEMALES. Band of broad decumbent scales of vertex very narrow and often

broken on midline; narrow decumbent scales of vertex and occiput very num-

erous; erect scales moderately long, very numerous and extending to orbital bris-

tles. Antenna distinctly longer than proboscis. Mesonotal vestiture moderately

dense, all dark, scales largely linear; inner dorsocentral line bare at anterior end;

scutellar scales linear; prescutellar bristles well developed; prescutals present; up-

per stp bristles present but not forming continuous row with lower bristles. Me-

tallic spots of femora golden to coppery but with dark violet iridescence, incon-

spicuous and variable in size; forefemur usually with small preapical spot. Hind-

tarsal segment 4 with a broad basal white or whitish ring. Abdominal tergites

without basal dorsal light bands; metallic spots small.

MALES. Palpus nearly as long as proboscis. Anterior claws of both foretarsus

and midtarsus with submedian tooth.

MALE GENITALIA. Median emargination of tergite VIII shallow. Lobes of

tergite IX widely separated. Sidepiece conical; without lateral scales; with short

setae on mesal surface; apicosternal process poorly developed. Accessory division

of subapical lobe not developed; distal division represented by distinct lobe with

1 or more specialized setae near apex of sidepiece. Clasper distinctly shorter than

sidepiece, only moderately curved dorsad; preapical lobe poorly developed; seta

b simple; seta a developed as normal thickened spiniform; crest ridged. Lateral

plate of phallosome with sternal spine; caudal process curved ventrad apically.

PUPAE. Cephalothoracic hair 5-C strong and usually at least 0.67 of trumpet

Valencia: Subgenus Carrollia of Culex 47

length. Trumpet short, index usually less than 7.0; pinna large or moderate.

LARVAE. Integument of head capsule with slight sculpturing visible at 100X,

particularly caudad; head hair 12-C about halfway between antennal base and

collar; 14-C near anterior ventral border of head capsule. Maxillary hair 5-MP

forked near middle. Anterior part of prothorax without spicules; prothoracic hair

QO-P always short and dendritic; 8-P similar in development to 7-P and usually

double or triple; mesothoracic hair 14-M very small and dendritic; metathoracic

hair 13-T single, strong and very long. Abdominal hair 1-II minute. Integument

of siphon sparsely covered with elongate spicules visible at 100X; distal pecten

teeth very long and slender. Hair 1-X single; 2-X fan-shaped, with numerous

branches; ventral brush (4-X) with at least 5 pairs of hairs, usually with 6.

DISCUSSION. The Urichii Subgroup is characterized by striking unique derived

features. In the adults hindtarsal segment 4 has a broad basal white ring and in

the fourth instar larvae prothoracic hair 8-P and metathoracic hair 13-T are thick-

ened. The subgroup also possesses some primitive character states not found even

in the Bihaicolus Group. In the male genitalia the distal division of the subapical

lobe is relatively better developed than elsewhere in the subgenus. In the larvae

there is a stronger development of the siphonal hairs and of the ventral brush

as well as hair 2-X of the anal segment. In other respects the Urichii Subgroup ap-

pears to be intermediate between the Bihaicolus Group and the Iridescens Sub-

group. In the adults the inner dorsocentral line is bare; prescutal and upper sterno-

pleural bristles are present; and the anterior claws of both the foretarsus and mid-

tarsus are toothed. In the pupae the trumpet is short and cephalothoracic hair

5-C strongly developed.

Because of this mosaic of characters and especially the unique features it is

tempting to elevate the subgroup to group rank. However the Urichii Subgroup

shares with the Iridescens Subgroup some basic correlated features in all stages:

(1) in the adults, darkened pleuron and reduction of dorsocentral bristles; (2)

in the male genitalia, caudal process on the lateral plate of the phallosome, para-

proct sclerotization with mesal sternal lobe, and seta b of the clasper displaced;

(3) in the pupae, position of hair 2 on abdominal segment II and hair 3 on seg-

ment VI; and (4) in the larvae, position of head hair 9-C and the type and ar-

rangement of the comb scales. The sharing of this combination of characters

clearly indicates a single phyletic line for the 2 subgroups and not an independ-

ent derivation of each subgroup from a common ancestor with the Bihaicolus

Group.

The subgroup is very compact. Its 2 component species, urichii and anduzei,

cannot be separated on external characters of the adults and the distinguishing

characters of the pupae and larvae are minor ones but apparently quite constant.

However male genitalic differences are clear cut.

The Urichii Subgroup is known from a wide area extending in the north to

Costa Rica and in the south to upper parts of the Amazon Basin in Brazil, Bolivia

and Peru. Culex urichii occupies the entire range and anduzei has a restricted sym-

patric distribution in the lower Amazon Basin.

The subgroup as a whole shows nearly the entire range of breeding sites and

associates known for the subgenus. The dominant urichii has been frequently col-

lected and shows a broad spectrum of breeding sites and associates but has not

been found in the flower bracts of Heliconia or leaf axils of Dieffenbachia. On

the other hand, immature stages of anduzei are known from only a few collec-

tions in treeholes, bamboo and tin cans and only a few breeding associates are

48 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

known for this species. Adults of both species have been collected in nature and

it appears that urichii is occasionally attracted to humans.

6. Culex (Car. ) urichii (Coquillett)

Figs. 3,7,9,20,21

1906. Melanoconion urichii Coquillett, 1906a:61. TYPE: Holotype female, St. Ann’s Valley,

Port-of-Spain, St. George, Trinidad, Jan 1906, F.W. Urich [USNM, 9141].

1942. Culex (Carrollia) mathesoni Anduze, 1942:47-49. TYPE: Holotype male genitalia only

(241), Penon de Parai-tepui, Bolivar, Venezuela, Nov 1940, P. Anduze [FH, 10111;

slide 2139]. NEW SYNONYM.

Culex (Carrollia) urichii of Dyar (1918:108; 1928:283,538); Bonne-Wepster and Bonne (1920:

171; 1923:124); Bonne and Bonne-Wepster (1925:197-200); Edwards (1932:221; 1934:663);

Komp (1936:64); Boshell-Manrique (1938:416); Kumm and Novis (1938:503,511); Senevet

and Abonnenc (1939:128; 1958:290-292); Lane (1939:83; 1953:511-513); Anduze (1941a:

17; 1941b:834); Cerqueira (1943:35; 1961:136); Cerqueira and Lane (1944:216-218); Anduze,

Pifano and Vogelsang (1947:16); Galindo, Carpenter and Trapido (1951:102,104,105,108,

126); Levi-Castillo (1952:553; 1953a:35); Galindo and Blanton (1955:73); Stone and Knight

(1957:59); Stone, Knight and Starcke (1959:283); Fauran (1961:44); Mattos and Xavier

(1965:280); Belkin, Schick and Heinemann (1965:70-71); Cova Garcia, Sutil and Rausseo

(1966a:28,11; 1966b:42,44,84,141,231,344); Barreto and Lee (1969:434); Forattini, Rabello

and Cotrim (1970:50); Morales-Ayala (1971:143).

Culex (Carrollella) urichii of Antunes (1937:78).

Carrollella urichii of Martini (1931:217).

Carrollia urichii of Dyar and Knab (1909:101); Urich (1913:528); Howard, Dyar and Knab

(1915:464-466).

Mochlostyrax urichii of Dyar (1906:18); Dyar and Knab (1906:223).

Melanoconion urichii of Coquillett (1906b:23); Theobald (1910:455,458).

Culex (Carrollia) mathesoni of Cerqueira and Lane (1944:218-219); Anduze, Pifano and Vogel-

sang (1947:16); Lane (1953:513); Stone, Knight and Starcke (1959:283); Fauran (1961:

44); Cerqueira (1961:136); Belkin, Schick and Heinemann (1965:76); Cova Garcia, Sutil and

Rausseo (1966a:28,113; 1966b: 141,344); Forattini, Rabello and Cotrim (1970:43).

FEMALE (figs. 7,9). Wing: 4.4 mm. Proboscis: 2.45 mm. Forefemur: 2.5 mm.

Abdomen: about 3.6 mm. Apparently indistinguishable from anduzei. General

characters as for the subgroup; description based on 29 topotypic specimens from

Trinidad. Head: Band of broad decumbent scales of vertex dark with bluish iri-

descence; narrow decumbent scales golden; erect scales golden; lateral patch white.

Proboscis 0.96-0.98 of forefemur. Labium largely dark scaled, indefinitely paler

ventrally. Palpus 0.21-0.22 of proboscis; entirely dark scaled. Antenna 1.06-1.12

of proboscis; torus with 8-10 setae. Thorax: Mesonotal and scutellar integument

very dark brown; scales dark bronzy; bristles dark bronzy except for some cop-

pery to golden prescutellars. Anterior promontory with 3-6 bristles; prescutals

2 or 3. Midlobe of scutellum with 6-8 long bronzy to coppery bristles, scales

bronzy; lateral lobe with 3 long bristles and bronzy scales. Pleuron usually en-

tirely darkened, including all of sternopleuron (lower part sometimes light), mer-

on and metameron; pruinose stripe very broad, greatly widened distally. Apn

with 5-8 bristles and usually numerous short broad erect black scales; ppn with

3 or 4 posterior bristles and several narrow bronzy scales; ppl bristles 4-6; stp

with 1-3 upper bristles and 4-6 lower posterior; pra bristles 6 or 7; umep bris-

tles 8-11. Legs: Coxal integument light, sometimes darkened at base; scales whit-

Valencia: Subgenus Carrollia of Culex 49

ish to light golden. Forefemur largely dark scaled; pale scaled at base, more ex-

tensively on posterior surface, and on a transverse posterior preapical band which

is extended basad dorsally for variable distance; anterior surface usually with a

variable metallic spot, often difficult to see. Midfemur light scaled at extreme

base, remainder dark scaled but scales along anteroventral margin with golden

iridescence and appearing light in some aspect; metallic spots basically golden

with deep violet reflections, extremely variable in size and shape and appearing

confluent with ventral golden streak. Hindfemur pale scaled except for dark api-

cal band and dorsal margin; metallic spot basically golden with deep violet iri-

descence, variable in size, located basad of apical dark band. Knee spots barely

indicated by a few pale apical scales on femora. Hindtarsal segment 4 with a

broad ring of white or whitish scales at base extending at least 0.5 but often

up to 0.8 of segment. Abdomen: Tergite I with an extensive patch of dark scales.

Light metallic spots of tergites basically silver with pale violet iridescence, of-

ten small on II,III and subbasal on IV,V; marginal bristles of tergites golden.

Sternites golden, IV-VII with black apical bands; apical bristles golden. Cibarial

Armature: Cibarial bar with moderately developed lobe on lateral flange; cibarial

teeth 3-5, very broad; cibarial dome with triangular denticles. Genitalia: Tergite

IX with 3-5 lateral bristles; insula with 18-20.

MALE. Wing: 4.4 mm. Proboscis: 2.9 mm. Forefemur: 2.5 mm. Abdomen:

about 3.8 mm. Indistinguishable from anduzei except on genitalic characters. Es-

sentially similar to the female except for usual sexual characters. Proboscis 1.1-

1.2 of forefemur. Palpus 0.86-0.89 of proboscis. Antenna 0.98-1.00 of probos-

cis. Specimens studied: 28 from the type locality in Trinidad.

MALE GENITALIA (fig. 20). Lobe of tergite IX usually with 6 (4-16) bris-

tles. Seta a of proximal division of subapical lobe with pointed curved apex;

seta b weakly dilated apically. Distal division with 2 short apically truncate and

2 long tapered flattened specialized setae on a short broad process and a group

of 6-20 flattened saberlike setae proximad of process. Clasper with poorly de-

veloped triangular preapical lobe. Distal part of proctiger relatively broad; para-

proct crown usually with 3 (2-6) teeth; cercal setae usually 1 pair (1-4).

PUPA (fig. 20). Abdomen: 3.25 mm. Trumpet: 0.65-0.70 mm. Paddle: 0.85

mm. Very similar to anduzei; diagnostic characters as in the key. General chaeto-

taxy based on 10 specimens from Trinidad. Cephalothorax: Pigmentation light

brown; sculpturing generally moderate but very distinct near trumpet base. All

hairs distinctly shorter than trumpet except 5-C which is usually subequal or at

least 0.67 of its length; 1(1;1-2); 2(2,3) subequal to hair 6; 3(2;2-3); 4(3,4;2-4)

subequal to hair 3; 5(2,3;1-4) barbed; 6(3,4;1-5) subequal to hair 2; 7(2;1-2);

8(1;1-2); 9(1;1-2). Trumpet: Moderately to strongly widened distally; index var-

lable, 5.3-7.5; pinna variable, 0.17-0.33. Pigmentation light brown. Metanotum:

Pigmentation yellow to light brown; sculpturing uneven, moderate; hair 10(2;

1-3) longer than hairs 11,12; 11(3,4;2-6); 12(2:1-2). Abdomen: Pigmentation yel-

low to light brown, anterior mesal areas of tergites II,TV darker; sculpturing mod-

erate. Segment I: hair 1(80-115) fan-shaped. Segment II: hair 1(22-38) longer

than hair 5; 3(1) strong, reaching level of hair 6-III; 4(8-12;6-12) very short;

5(1,2;1-3) about twice as long as hair 4. Segment III: hair 1(4,5;2-6); 3(1) strong,

reaching level of hair 6-IV; 4(3,4;2-5) forked; 5(2,3;1-4) about twice as long as

hair 4. Segment IV: hair 1(1,2) subequal to hair 1-V; 3(4,5;3-8); 4(2;1-4); 5(1)

strong, surpassing level of rudimentary spiracle VI. Segment V: hair 1(1,2) subequal

to hair 1-IV; 3(1;1-2); 4(5,6;3-8); 5(1) strong, surpassing level of rudimentary spiracle

50 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

VII. Segment VI: hair 1(1) weak and shorter than hair 6; 3(1,2;1-4) shorter than hair

1; 4(3,4;1-6) shorter than hair 3; 5(1) at most reaching level of rudimentary spiracle

VII. Segment VII: hair 1(1;1-2) longer than hair 4; 3(2,3;1-3); 4(2;1-2); 5(4-6;2-6)

subequal to hair 6; 6(8-10;5-15) short and dendritic; 9(3,4;3-6) barbed, shorter than

hair 9-VIII. Segment VIII: hair 4(1) shorter than hair 1-VII; 9(3,4) shorter than

paddle.

LARVA (fig. 21). Head: 0.95 mm. Siphon: 1.25 mm. Anal Saddle: 0.40 mm.

Very similar to anduzei; diagnostic characters as in the key. General chaetotaxy

based on 7 reared and 3 other specimens from Trinidad. Head: Integument yel-

low to brown; collar darkened. Mental plate with 9 or 10 lateral teeth. Hair

4(6,7;5-9); 5(8-10;6-10); 6(2,3;1-4); 7(4,5;4-8); 8(1;1-2) subequal to hair 7; 9

(5,6;5-7); 10(3;2-4); 11(3,4;2-5); 12(1); 14(1-2); 15@;2-4). Antenna: About 0.35

of head length; hair 1(3,4;3-6). Thorax: Prothorax: hair 0(30-40); 1(1); 2(1); 3

(3-4); 4(2); 7(4;3-6); 8(2;2-3); 9(1); 10(1); 11(5,6;4-6); 14(1). Mesothorax: hair

1(1;1-2); 2(2,3;1-3); 3(1); 4(2); 8(12;11-16); 9(10-12;8-12); 11(1); 13(20-50); 14

(27-40). Metathorax: hair 1(2;1-4); 2(2;2-3); 3(6,7;5-8); 4(6-8;4-10); 6(1); 7(8,

9;6-10); 8(25-50); 9(4;3-5); 11(1); 13(1). Abdomen: Segment I: hair 1(1;1-2)

minute; 6(2); 7(2). Segment II: hair 1(1;1-3); 3(1) moderate; 6(2). Segment III:

hair 1(1) very long, strong; 3(1;1-2) moderate; 6(1); 13(1) subequal to hair 1.

Segment IV: hair 1(1) very long, strong and usually slightly longer than hair

1-III; 3(3,4;2-5) short; 6(1); 13(1) subequal to hair 1. Segment V: hair 1(1) sub-

equal to hair 1-III; 3(1) long; 6(1); 13(1) subequal to hair 1. Segment VI: hair

1(1) weak and relatively short; 6(1). Segment VII: hair 1(1) very strong and

long, usually reaching middle of siphon. Segment VIII: Comb scales (17-22) with

lateral fringes distinct at base only; hair 1(8-10;8-16); 3(11,12;9-12); 5(2;2-3).

Siphon: Pigmentation light brown. Index about 4.0-5.0. Pecten teeth (8-11) with

indistinct fringes on ventral margin. Ventral siphonal tufts (14-16) with about

8 branches (6-10). Anal Segment: Saddle pigmentation light brown, darkened

dorsad. Hair 1(1); 2(12-16). Ventral brush with 6 pairs of hairs, all except the

proximal pair usually with at least 16 branches. Gills usually less than 2.0 of dor-

sal saddle length, tapered.

SYSTEMATICS. Culex urichii is very similar to anduzei, the other member of

the subgroup. I have not been able to separate the 2 species on any external

features of the adults. However, the male genitalia are quite distinctive and show

no overlap in the key characters. The larvae and pupae are very similar but ap-

parently readily separated by the relatively minor differences used in the keys.

A careful study of the male genitalia of some 80 specimens has revealed that

the reduced number of foliaceous setae on the distal division of the subapical

lobe, supposedly diagnostic of mathesoni, falls well within the wide range of var-

iation exhibited by typical urichii. I have therefore synonymized mathesoni with

urichii.

Culex urichii is the dominant species of the subgroup and has the widest known

distribution in the subgenus. It might appear therefore that it is the derived spec-

ies of the subgroup. However, because urichii has retained primitive character

States in the male genitalia (well developed distal division of the subapical lobe;

relatively simple clasper head; cercal setae), pupa (short hair 5-VI) and larva (more

numerous and more profusely branched hairs in the siphon and in the ventral

brush), I believe that it is closer to the ancestral stock than anduzei, which has

derived states in these characters.

BIONOMICS. Culex urichii has been collected from nearly as wide a range of

Valencia: Subgenus Carrollia of Culex 51

natural breeding sites as bihaicolus: treeholes (18), broken or cut bamboo (11),

fallen leaves and palm spathes (7), fallen nuts (1) and rockholes (1). It has also

been frequently found in a wide variety of artificial containers (15) and twice

in bamboo traps. Two species of Carrollia have been recorded as breeding assoc-

iates: bihaicolus (2) and bonnei (3); and 15 other species representing nearly

every sympatric group with similar breeding sites: Aedes (H.) ioliota (3), A. (O.)

fluviatilis (1), A. (P.) insolitus (4), A. (P.) podographicus (1); Anopheles (An.)

eiseni (4); Culex (Anoed.) sp. (1), C. (C.) corniger (1), C. (C.) mollis (5), C. (Mel.)

sp. (1); Limatus durhamii (1); Orthopodomyia albicosta (1), O. fascipes (2); Sa-

bethes undosus (3); Trichoprosopon (R.) ulopus (1); Wyeomyia complosa (1).

There are more records of urichii collected as adults in nature than for any

other species of Carrollia. | have seen 1 male and 2 females collected by T.H.G.

Aitken in Trinidad. Komp (1936:64) reported females attracted to humans in

the Villavicencio area in Colombia; Kumm and Novis (1938:503) captured adults

attracted to animal bait (agouti) on Marajo Island, Brazil; and Cerqueira (1961:

136) found females on human and animal bait in Manaus, Brazil.

DISTRIBUTION (fig. 3). From central Costa Rica through northern South

America to Pacific drainage in Colombia and southern drainages of the Amazon

basin in Peru, Bolivia and Brazil, at elevations of 20-800 m.

Material examined: 515 specimens; 120 males, 100 females, 176 larvae, 119

pupae; 96 individual rearings (50 larval, 15 pupal, 31 incomplete).

BRAZIL. Amazonas: Rio Paruari (FM 38) [USNM]. Para: near Belem (BRA 38) [UCLA].

COLOMBIA. Antioquia: Providencia, Anori R. valley, C. Porter [UCLA]. Meta: Balconcito

(KO 1-36) [UCLA]. Bosque Ocoa (C-24, CV 581,1014) [UCLA]. Restrepo, W.H. Komp (565,

566,585,598) [UCLA]; (10-40) [USNM]; (1000) [FH]. Retiro (KO 10-28) [UCLA]. Rio Gua-

cavia, W.H. Komp [UCLA]. Villavicencio, M. Bates (194) [USNM] ; (7438,7439) [FH].

COSTA RICA. Puntarenas: Esparta (CR 356) [UCLA].

FRENCH GUIANA. Guyane: Cabassou (FGC 3518,3902) [UCLA]. Cogneau (FGC 3384,3462,

3472) [UCLA]. Le Gallion (FGC 3138,3173,3375,3379) [UCLA]. Rochambeau (FGC 3536)

[UCLA] . Matouri (FGC 3382) [UCLA].

PERU. Loreto: Iquitos, R.C. Shannon [USNM].

PANAMA. Canal Zone: Barro Colorado, W.H. Komp [UCLA, USNM]. Madden (PA 707)

[UCLA]. Chiriqui: El Volcan, W.H. Komp [UCLA]. Darien: Morti (PA 948) [UCLA]. Panama:

Cerro Azul (94-W) [USNM]. Pacora (4897,4942,6992) [UCLA].

SURINAM. Marowijne: Moengo, H.H. Stage [USNM]. Locality not Specified; BB 641,642,

666,701 [USNM].

TRINIDAD. Nariva: Charuma Forest (TR 750) [UCLA]. Nariva Swamp, T. Aitken [UCLA].

Tabaquita, T. Aitken [UCLA]. St. Andrew: Coryal (TR 492) [UCLA]. Cumaca (TR 782,787,

940,1009,1144) [UCLA]. Guatepajaro road (TR 859) [UCLA]. Oropouche Swamp, T. Aitken

[UCLA]. Nestor (TR 475) [UCLA]. Turure, T. Aitken [UCLA]. Valencia old road (TR 1113)

[UCLA]. St. George: Chaguaramas (TR 49) [UCLA]. Guanapo Heights (TR 1119) [UCLA].

L’Orange road (TR 707) [UCLA]. St. Ann’s Valley, F.W. Urich, holotype of urichii [USNM,

9141]. St. Pats, T. Aitken [UCLA]. Verdant Vale (TR 676,677,822,1452) [UCLA].

VENEZUELA. Aragua: near Choroni (VZ 315) [UCLA]. Ocumare de la Costa, road to

(VZ 310) [UCLA]. Turiamo, W.H. Komp [USNM]. Bolivar: Penon de Parai-tepui, P. Anduze,

Foes of mathesoni [FH]. Monagas: Quiriquiri (KO 1-24) [UCLA]. Maturin, W.H. Komp

Additional Records From the Literature

BOLIVIA. Beni: Vaca Diez (Cerqueira, 1943:35).

BRAZIL. Amazonas: Manaus (Cerqueira, 1961:136). Rio Maues [FH, 4097] (Cerqueira, 1944:

218). Rio Paruari (Cerqueira, 1944:218). Goias: Anapolis, Inhumas (Mattos and Xavier, 1965:

280). Para: Curralinho (Cerqueira, 1961:136). Marajo Island (Kumm and Novis, 1938:503).

52 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

BRITISH GUIANA. Essequibo River (Edwards, 1934:663).

ECUADOR. Napo-Pastaza: Arajuno, Shell-Mera, Tena (Levi-Castillo, 1952:553).

FRENCH GUIANA. Guyane: Cabassou. Crique Anguille. Rochambeau (Fauran, 1961:44).

Inini: Balourou. Saut Canori. St. Elie (Fauran, 1961:44).

PERU. Huanuco: Rio Pachitea. Pasco: Puerto Bermudez (Martini, 1931:217; Morales-Ayala,

1971:143).

VENEZUELA. Aragua: Choroni, Guamitas. Ocumare de la Costa. Rancho Grande (Cova

Garcia, 1966b:84). Bolivar: Surukum. Uriven (Anduze, 1941b:834). Monagas: Caripito, Quiri-

quiri (Cova Garcia, 1966b:84).

7. Culex (Car.) anduzei Cerqueira & Lane

Figs. 3,9,22,23

1944. Culex (Carrollia) anduzei Cerqueira and Lane, 1944:219-220. TYPE: Holotype male,

Rio Maues, Amazonas, Brazil, Feb 1937, C. Worontzow [FH, 4099].

Culex (Carrollia) anduzei of Lane (1953:513-514); Stone, Knight and Starcke (1959:282); Cer-

queira (1961:136); Forattini, Rabello and Cotrim (1970:35); Belkin, Schick and Heinemann

(1971:27); Barata and Cotrim (1972:25-30).

FEMALE (fig. 9). Wing: 4.1 mm. Proboscis: 2.4 mm. Forefemur: 2.4 mm.

Abdomen: about 3.2 mm. Apparently indistinguishable from urichii on external

features. Description and measurements based on 5 pinned specimens from Belem,

Brazil. Head: Proboscis 1.00-1.18 of forefemur; palpus 0.20 of proboscis. Anten-

na 1.10-1.18 of proboscis; torus with 10-12 setae. Thorax: Anterior promontory

with 2-4 bristles; prescutals 2-4. Lateral lobe of scutellum with 3 or 4 bristles.

Apn with 4-6 bristles, broad scales apparently less numerous than in urichii; ppl

bristles 4; stp with 2 upper bristles and 5 or 6 lower posterior; pra bristles 6-8;

umep bristles 6. Cibarial Armature: Cibarial bar without lobe on lateral flange;

cibarial teeth 5. Genitalia: Tergite IX with 2 or 3 lateral bristles; insula with

20-24.

MALE. Wing: 3.5 mm. Proboscis: 2.6 mm. Forefemur: 2.2 mm. Abdomen:

about 3.1 mm. Indistinguishable from urichii except on genitalic characters. Es-

sentially similar to the female except for sexual characters. Proboscis 1.10-1.18

of forefemur. Palpus 0.92 of proboscis. Antenna 1.01-1.03 of proboscis. Spec-

imens studied: 5 from Belem, Brazil. Measurements from dry specimens. |

MALE GENITALIA (fig. 22). Lobe of tergite IX usually with 4 (4-6) setae.

Setae a and b of proximal division of subapical lobe both with dilated recurved

apex. Distal division represented only by a digitiform process bearing 1 heavy

bristlelike thickened apical seta. Clasper with large broad preapical lobe. Distal

part of proctiger relatively narrow; paraproct crown usually with 4 (3-5) teeth;

cercal setae usually 2 pairs (1 or 2).

PUPA (fig. 22). Abdomen: 3.20 mm. Trumpet: 0.65 mm. Paddle: 0.70 mm.

Very similar to urichii; diagnostic characters as in the key. General chaetotaxy

based on 9 specimens from Brazil. Cephalothorax: Pigmentation light brown;

sculpturing moderate. All hairs distinctly shorter than trumpet except 5-C which

is subequal; 1(1;1-2); 2(1;1-2) subequal to hair 6; 3(2;1-2); 4(3,4;1-4) longer than

hair 3; 5(1,2;1-3); 6(1,2;1-3); 7(2;1-2) twice as long as hair 4; 8(1); 9(1) longer

than hair 8. Trumpet: Moderately to strongly widened distally; index about 4.0

(3.6-5.0); pinna about 0.33. Pigmentation light brown. Metanotum: Pigmentation

Valencia: Subgenus Carrollia of Culex 53

yellow to light brown; sculpturing uneven, moderate; hair 10(5,4;1-5); 11(3,4;

2-6) shorter than hairs 10,12; 12(1,2;1-3). Abdomen: Pigmentation yellow to

light brown, anterior mesal areas of tergites III,I[V slightly darker; sculpturing

moderate. Segment I: hair 1(50-100) fan-shaped. Segment II: hair 1(15-20;14-

22) subequal to hair 3; 3(1) strong, reaching level of hair 6-III; 4(3-6;3-8); 5(1,

2;1-3) poorly developed. Segment III: hair 1(3-5;2-6); 3(1) strong, reaching level

of hair 6-IV; 5(2,3;1-4). Segment IV: hair 1(1,2) longer than hair 1-V; 3(3,4;3-

7); 4(2,3); 5(1) strong, surpassing level of rudimentary spiracle VI. Segment V:

1(1,2); 3(1,2) longer than hair 1; 4(4,5;4-6); 5(1) strong, surpassing level of rud-

imentary spiracle VII but shorter than hair 5-[V. Segment VI: hair 1(1;1-2) short-

er than hair 3; 3(2,3;1-4) subequal to hair 4; 4(3,4;2-4); 5(1) strong, surpass-

ing caudal margin of tergite VII. Segment VII: hair 1(1) reaching level of hair

4-VIII; 3(1,2) shorter than hair 5; 4(1,2); 5(4-6;2-6); 6(4-16) short; 9(4,5;3-10)

barbed, slightly shorter than hair 9-VIII. Segment VIII: hair 4(1;1-2); 9(4,5;3-

5) shorter than paddle.

LARVA (Fig. 23). Head: 1.00 mm. Siphon: 1.00 mm. Anal Saddle: 0.30 mm.

Very similar to urichii; diagnostic characters as in the key. General chaetotaxy

based on 1 reared and 9 other specimens from Brazil. Head: Integument light

brown; collar darkened. Mental plate with 9 or 10 lateral teeth. Hair 4(4,5;3-

6); 5(9,10;8-12); 6(3,4;2-4); 7(6;5-7); 8(1) subequal to hair 7; 9(6-8); 10(2,3;

1-3); 11(3;2-4); 12(1); 14(1-2); 15(4,5;3-6). Antenna: About 0.30-0.35 of head

length; hair 1(3,4;2-6). Thorax: Prothorax: hair 0(28-30); 1(1); 2(1); 3(3;2-4);

4(2); 7(4;3-5); 8(2); 9(1); 10(1); 11(3,4;2-4); 14(1). Mesothorax: hair 1(1); 2

(3,4;3-6); 3(1); 4(2); 8(8,9;6-14); 9(8,9;7-9); 11(1;1-2); 13(30-38); 14(20-30). Meta-

thorax: hair 1(2;1-2); 2(1-2); 3(4,5;4-6); 4(4-6); 6(1); 7(6,7;5-7); 8(30-40); 9¢4,

533-5); 11(1); 13(1). Abdomen: Segment I: hair 1(2;1-2) minute; 6(2); 7(2). Seg-

ment II: hair 1(1); 3(1) moderate; 6(2;1-2). Segment III: hair 1(1) long, strong;

3(1) moderate; 6(1); 13(1) subequal to hair 1. Segment IV: hair 1(1) subequal

to hair 1-II; 3(2;1-3) shorter than hair 3-III; 6(1); 13(1) subequal to hair 1.

Segment V: hair 1(1) subequal to hair 1-III,IV; 3(1) subequal to hair 3-III; 6

(1); 13(1) subequal to hair 1. Segment VI: hair 1(1) weak and relatively short;

6(1). Segment VII: hair 1(1) very strong and long but usually not reaching mid-

dle of siphon. Segment VIIT: Comb scales (12-14, rarely 9) with lateral fringes

distinct to near apex; hair 1(4,5;3-7); 3(10;8-11); 5(2;1-2). Siphon: Pigmentation

brown. Index about 3.6-4.2. Pecten teeth (7-10) with indistinct fringes on ven-

tral margin. Ventral siphonal tufts (11-12) with about 7 branches (5-8). Anal

Segment: Saddle pigmentation light brown. Hair 1(1); 2(10-14). Ventral brush

with 5 or 6 pairs of hairs, all except the proximal pair usually with at least 12

branches. Gills more than 2.0 of dorsal saddle length, tapered.

SYSTEMATICS. As noted under urichii, the other member of the subgroup,

the 2 species are very similar and cannot be separated at this time by any external

feature of the adults. There are, however, relatively minor but constant char-

acters distinguishing anduzei from urichii in the male genitalia, pupae and larvae

as indicated in the respective keys.

I am interpreting anduzei as the derived species of the group (see urichii), and

possibly segregated directly from uwrichii in recent times. Its distribution appears

to be limited to the lower Amazon Basin where it is completely sympatric with

the dominant widespread urichii. Nothing is known of the barrier separating the

2 species. Both breed in treeholes and bamboo and have been reported from

the same localities but there are no records of the association of these species

in the same breeding site.

54 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

BIONOMICS. The only reported natural breeding sites of anduzei are treeholes

(2) and cut or broken bamboo (1) (Barata and Cotrim, 1972:30). The small ser-

ies of associated adults was reared from a collection in tin cans, where anduzei

was associated with Culex (Anoed.) sp., Limatus asulleptus and Wyeomyia sp.

Cerqueira (1961:136) reports adults obtained by sweeping in Manaus, Brazil.

It appears likely that the collections of males and females of anduzei ‘‘no solo”’

by L. Gomes in 2 localities in the State of Para, Brazil (Barata and Cotrim, 1972:

30) were also made by sweeping.

DISTRIBUTION (fig. 3). Lower Amazon basin downstream from Manaus, at

elevations of 30-50 m.

Material examined: 37 specimens; 6 males, 4 females, 13 larvae, 14 pupae; 9

individual rearings (1 larval, 8 pupal).

BRAZIL. Amazonas: Rio Maues, C. Worontzow, holotype [FH]. Para: near Belem (BRA

54) [UCLA].

Additional Records From the Literature

BRAZIL. Amazonas: Rio Maues (4099-4120) [FH] (Cerqueira and Lane, 1944:220). Manaus

(Cerqueira, 1961:136). Para: Belem, Benevides, Jacarequara, Sao Domingos do Capim (Barata

and Cotrim, 1972:30).

IRIDESCENS SUBGROUP

FEMALES. Band of broad decumbent scales of vertex always broad to very

broad; narrow decumbent scales of vertex numerous to apparently absent; erect

scales relatively short, either numerous and extending to orbital bristles or few

and confined to caudal part of vertex and/or occiput. Antenna subequal to pro-

boscis or slightly shorter. Mesonotal vestiture very dense, with dark scales only

or sometimes with light coppery to dark golden scales also; inner dorsocentral

line entirely scaled; scutellar scales linear, narrow curved or broad; prescutellar

bristles poorly developed and sometimes apparently completely absent; prescu-

tals absent; upper stp bristles absent. Metallic spots of femora silvery with light

violet iridescence, conspicuous and discrete on midfemur; forefemur apparently

always without metallic spot. Hindtarsus entirely dark scaled. Abdominal ter-

gites with more or less distinct basal dorsal light bands.

MALES. Palpus subequal to proboscis or only about 0.72 of its length. An-

terior claw of foretarsus only with submedian tooth.

MALE GENITALIA. Median emargination of tergite VIII usually very deep.

Lobes of tergite IX moderately to closely approximated. Sidepiece ovoid or con-

ical; with lateral scales; without setae on mesal surface; apicosternal process well

developed. Accessory division of subapical lobe variously developed; distal divi-

sion represented by a small protuberance with 1 or 2 depressions but without

specialized setae, or apparently completely absent. Clasper subequal to or short-

er than sidepiece, usually strongly curved dorsad; preapical lobe moderately to

very strongly developed; seta b thickened or flattened; seta a developed as a

normal thickened spiniform or a simple bristle. Lateral plate of phallosome with

or without sternal spine; caudal process straight or curved dorsad or laterad apically.

PUPAE. Cephalothoracic hair 5-C weak and usually less than 0.5 of trumpet

length. Trumpet moderate to very long, index 6.8 to nearly 20.0; pinna mod-

erate to minute.

Valencia: Subgenus Carrollia of Culex 55

LARVAE. Integument of head capsule appearing smooth at 100X; head hair

12-C distinctly nearer collar than antennal base; 14-C usually nearer posterior

tentorial pit than anterior ventral border of head capsule, at most equidistant.

Maxillary hair 5-MP not forked. Anterior part of prothorax with spicules; pro-

thoracic hair O-P usually moderate and more or less stellate; 8-P different in de-

velopment from 7-P, multiple, weak or stellate; mesothoracic hair 14-M mod-

erate, multiple and sometimes more or less stellate; metathoracic hair 13-T branch-

ed, moderate to rather large and somewhat stellate. Abdominal hair 1-II mod-

erate to strong. Integument of siphon appearing smooth at 100X; distal pecten

teeth short. Hair 1-X branched, usually multiple; 2-X asymmetrical, with 1 long

and 2-4 shorter branches; ventral brush with no more than 4 pairs of hairs.

DISCUSSION. The Iridescens Subgroup exhibits more derived character states

in all stages than any other major phyletic line in the subgenus. Only a few of

the characters listed in the diagnosis have been retained in the primitive state:

(1) ornamentation of the legs, (2) conical sidepiece in some species, (3) branched

larval hair 6-III-[V in a few species. The most striking feature shared by all mem-

bers of the complex are in the male genitalia: (1) loss of the distal division and

elaboration of an accessory division in the subapical lobe of the sidepiece and

(2) modification of the distal part of the clasper.

The subgroup is very complex and it has not been possible to resolve it satis-

factorily largely because of the very limited material of the majority of the spec-

ies, especially of associated immature stages from Brazil.

I am recognizing 2 primary divisions in the subgroup, the Iridescens Complex

and the Antunesi Complex. This division is based largely on male genitalic char-

acters in the development of the head of the clasper and the presence or ab-

sence of a sternal spine on the lateral plate of the phallosome. The only support-

ing character for this division is found in the development of the erect and de-

cumbent scales on the vertex of the head. I have not found any concordant

features in the larvae or pupae. The 2 complexes are in general sympatric, with

some wide gaps in northeastern Brazil.

The compact Iridescens Complex of 3 named species (bonnei, secundus, iri-

descens) and 2 unnamed forms (sp. 8a, Bahia form; sp. 9a, Espirito Santo form)

is undoubtedly monophyletic. It is characterized in the male genitalia by a strongly

differentiated elongate spiculose lobe on the head of the clasper and its flattened

seta b, and by the lateral plate of the phallosome without sternal spine. The

sidepiece of the male genitalia is short and ovoid. The vertex of the head of the

adults has numerous narrow decumbent scales and the erect scales extend to near

the orbital bristles. The immature stages do not seem to possess any significant

phyletic characters, except perhaps in the development of hair 1-VII of the lar-

vae. The interrelations of the species of the complex are difficult to determine

and I have not attempted to do it because of inadequate material of iridescens

and sp. 8a and 9a. The central part of the range of the complex is occupied

by bonnei and sp. 8a which have similar and very distinctive male genitalia. Spec-

ies 8a may be only an isolated southern population of bonnei differing primar-

ily in the scaling of the midlobe of the scutellum but its taxonomic status can-

not be determined without additional material of adults and particularly imma-

ture stages. Culex secundus occupies the northern part of the range and is sym-

patric with bonnei in Colombia on both sides of the Cordillera Central of the

Andes; it shows more similarity with bonnei than with iridescens in the devel-

opment of the accessory division of the subapical lobe of the male genitalia and

56 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

the unbranched hair 1-VII of the larva. The taxonomic status of the Espirito

Santo form, sp. 9a, cannot be determined without additional material. It appears

to be related to secundus on male genitalic characters and may be nothing more

than a widely isolated population of this species. The poorly known iridescens,

the type species of the subgenus, has a restricted distribution at the southern end

of the range of the complex in southern Brazil and adjacent northeastern Ar-

gentina. It is sympatric with sp. 9a in the State of Espirito Santo, Brazil. It shows

more similarities with secundus than with bonnei in external features of the adults

and in the male genitalia and was probably the earliest segregate from the ancestral

form of the complex. Species of the Iridescens Complex utilize a wide range of

breeding habitats and have a wide range of associates. Except for 1 male and

1 female of bonnei captured in a trap there are no records of adults of this com-

plex collected in nature.

The Antunesi Complex is very diverse and poorly understood. It is character-

ized in the male genitalia by a poorly differentiated head on the clasper and its

unflattened seta b, and by the lateral plate of the phallosome with a sternal

spine. The sidepiece of the male is either elongate and conical or short and ovoid.

The vertex of the head of the adults has only a few or no narrow decumbent

scales and the erect scales are confined to its posterior part and/or occiput. The

immature stages show a variety of primitive and derived character states in the

different species. Most species of the complex appear to have relict distributions

and are known only from a few collections. It is very unlikely that the complex

is monophyletic as 4 more or less distinct lines may be recognized, each with

rather clearcut features. The Kompi line consists of a single species with a unique

and bewildering combination of primitive and derived character states in the dif-

ferent stages; kompi is only known from the Villavicencio area of Colombia where

it is sympatric with 2 species of the Antunesi line. The Babahoyensis and Soperi

lines, each with a single species, have superficially similar accessory divisions of

the subapical lobe of the male genitalia but differ conspicuously in the shape

of the sidepiece, scaling of the scutellum and in the immature stages. The 2 lines

have moderately wide distributions but occupy opposite ends of the range of

the complex: babahoyensis in the north in Nicaragua, Costa Rica and Ecuador

and soperi in southern Brazil and adjacent northeastern Argentina. The species

of the Antunesi line are obviously closely related because of similarity in exter-

nal features of adults, in male genitalia and larval and pupal characters. There

are at least 3 species in this line, all apparently rare and with relict or disjunct

distributions; cerqueirai is known from a single collection in northwestern Pan-

ama; wilsoni from 2 widely separated localities in the Villavicencio area of Colom-

bia and the State of Espirito Santo in Brazil; antunesi from Costa Rica, Panama

and Colombia, from French Guiana and from the States of Espirito Santo and

Rio de Janeiro in Brazil. As noted under the respective species, it is possible

that the northern populations of antunesi and wilsoni are not conspecific with

the typical southern populations and that there are 2 or 3 additional species in

the Antunesi line. Species of the Antunesi Complex appear to be almost entirely

restricted to breeding in bamboo but there is 1 record of babahoyensis from a

treehole. The breeding associates of the complex are also limited. The only rec-

ord of an adult captured in nature is for soperi.

It appears from the above considerations that early in the evolution of the Iri-

descens Subgroup there was a separation into several phyletic lines, only 1 of

which, the Iridescens Complex, was really successful and is now dominant. The

Valencia: Subgenus Carrollia of Culex 57

remaining lines, constituting the Antunesi Complex, are now largely relict although

in the past some of them, especially the Antunesi line and possibly the Soperi-

Babahoyensis lines, had wide distributions.

8. Culex (Car.) bonnei Dyar

Figs. 4,9,24,25

1921. Culex (Carrollia) bonnei Dyar, 1921:155. TYPE: Lectotype male (BB 550), Gansee,

Surinam, date not specified, C. Bonne and J. Bonne-Wepster [USNM, 24862; designation

by Stone and Knight, 1957:44]. Reduced to synonymy with iridescens by Bonne-

Wepster and Bonne (1920:170); resurrected to specific rank by Antunes and Ramos

(1939:381).

Culex (Carrollia) bonnei of Lane (1953:500-502); Stone and Knight (1957:44); Senevet and

Abonnenc (1958:288); Stone, Knight and Starcke (1959:282); Cerqteira (1961:156); Fauran

(1961:44); Belkin, Schick and Heinemann (1965:65); Forattini, Rabello and Cotrim (1970:

36).

Culex (Carrollia) bonnei in part of Antunes and Ramos (1939:381-383); Lane and Whitman

(1943 :397).

Culex (Carrollia) secundus of Levi-Castillo (1953a:35).

Culex (Carrollia) secundus in part of Barata and Cotrim (1972:23-24).

Culex (Carrollella) secundus in part of Komp (1936:64; 1956:39).

Culex (Carrollia) iridescens of Bonne-Wepster and Bonne (1920:170); Bonne and Bonne-Wepster

(1925:200-203); Komp (1936:64; 1956:39); Antunes (1937:78); Kumm and Novis (1938:

503,511); Senevet and Abonnenc (1939: 123-124,126,129,132); Levi-Castillo (1953a:35).

Culex (Carrollia) iridescens in part of Edwards (1932:221); Stone, Knight and Starcke (1959:

283).

Culex (Carrollella) iridescens in part of Dyar (1928:280-281,538).

Carrollia iridescens in part of Peryassu (1908:34,44,167-169,343).

FEMALE (fig. 9). Wing: 2.7 mm. Proboscis: 1.9 mm. Forefemur: 1.8 mm.

Abdomen: about 2.5 mm. Very similar to iridescens and secundus; diagnostic

characters as in the key. Differing from secundus primarily in the following fea-

tures. Description based on 2 specimens from Surinam, 2 from French Guiana,

3 from Colombia and 9 from Ecuador. Head: Broad decumbent scales of ver-

tex sometimes all pale. Proboscis 1.05-1.08 of forefemur. Palpus 0.20-0.21 of

proboscis. Antenna 0.94-1.05 of proboscis; torus with 3-5 setae. Thorax: Mid-

lobe of scutellum with numerous short broad dark bronzy scales in addition to

a few linear scales. Sternopleuron dark below level of coxae, usually completely.

Apn with 6-8 bristles; ppn bristles 3 or 4; stp with 4-6 lower posterior bristles;

pra bristles 4 or 5; umep bristles 6-8. Legs: Forecoxal scale patch without dark

scales. Forefemur with a longer pale streak on posterior surface, sometimes to

near apex. Abdomen: Dorsal basal tergal light bands less developed, usually on-

ly on VI,VII, small and dingy white. Cibarial Armature: Lobe of lateral flange

poorly developed; 4 median cibarial teeth; cibarial dome larger and with triang-

ular denticles. Genitalia: Tergite IX with 4-6 lateral bristles; insula with 20.

MALE. Wing: 3.2 mm. Proboscis: 2.3 mm. Forefemur: 2.0 mm. Abdomen:

about 2.5 mm. Very similar to iridescens and secundus; diagnostic characters as

in the key. Essentially similar to female except for usual sexual differences. Dif-

fering from secundus primarily in the following features. Proboscis 1.15 of fore-

femur; pale scaling of labium not as white; long subapical bristles not developed.

Palpus 0.78-0.84 of proboscis. Antenna 0.86-0.90 of proboscis. Specimens studied:

3 from Surinam, 6 from French Guiana, 4 from Colombia and 2 from Ecuador.

MALE GENITALIA (fig. 24). Lobes of tergite IX separated by distance sub-

58. Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

equal to width of 1 lobe at base; each lobe usually with 5 (3-7) bristles. Side-

piece ovoid; apicotergal seta strongly developed. Accessory division of subapi-

cal lobe not joined to proximal division, long columnar and with 4 apical or

preapical elongated flattened specialized setae. Proximal division elongate, curved,

with 2 or 3 simple bristles distally; setae a and b with dilated apex with recurved

sharp point, seta a longer. Distal division represented by a small protuberance.

Clasper subequal to sidepiece length; strongly curved dorsad; with long slender

basal stem and strongly modified head; with a prominent outer preapical curva-

ture and a long spiculose inner preapical lobe; crest apparently represented by

a flat sclerite; basal (lower) spicules of preapical lobe very long and hairlike, dis-

tal (upper) spicules short; snout moderate; seta a developed as a normal thick-

ened spiniform, seta b flattened, foliform. Lateral plate of phallosome without

sternal spine. Distal part of proctiger broad and rounded apically; paraproct crown

usually with 4 (3-5) teeth; cercal setae usually 2 pairs (1-3).

PUPA (fig. 24). Abdomen: 3.15 mm. Trumpet: 0.55 mm. Paddle: 0.70 mm.

Very similar to iridescens; diagnostic characters as in the key. General chaeto-

taxy based on 7 specimens from French Guiana. Cephalothorax: Pigmentation

light yellow to brown; sculpturing weak, without definite pattern. All hairs short-

er than trumpet and moderately developed, 1,4,5-C subequal, 7-C the longest;

1(1,2); 2(2); 3(2;1-2); 4(2;1-4); 5(2;1-4); 6(2;1-3) shorter than hair 2; 7(2;1-4);

8(2;1-2) shorter than hair 1; 9(2;1-2). Trumpet: Nearly uniform in width; index

6.8-7.8; pinna small. Pigmentation yellow to brown. Metanotum: Pigmentation

yellow to dark brown; sculpturing uneven, moderate; hair 10(2,3;2-4); 11(2,3;

2-5); 12(2,3;1-3) longer than others. Abdomen: Pigmentation light yellow to brown,

anterior mesal areas of tergites II-IV slightly darker; sculpturing weak to mod-

erate. Segment I: hair 1(70-110) fan-shaped. Segment II: hair 1(7;7-12) distinct-

ly longer than hair 5; 3(1) strong, surpassing level of rudimentary spiracle III;

4(6,7); 5(2;1-3). Segment III: hair 1(4;3-4); 3(1) strong, surpassing level of rud-

imentary spiracle IV; 4(2,3;2-5); 5(2,3;1-3). Segment IV: hair 1(2;2-5); 3(4-6);

4(1;1-2); 5(1) strong, surpassing level of rudimentary spiracle VI. Segment V:

hair 1(2;1-3) subequal to hair 3; 3(2;2-3); 4(5,6); 5(1) strong, surpassing level

of rudimentary spiracle VII. Segment VI: hair 1(1;1-2); 3(33;1-3); 4(4;2-5); 5(1)

strong, surpassing caudal margin of tergite VII. Segment VII: hair 1(1); 3(1;1-

2); 4(2;1-2); 5(4;3-5) subequal to hair 6; 6(6;4-7); 9(5;5-7) barbed. Segment VIII:

hair 4(2;1-2); 9(8;7-14) sometimes secondarily branched, distinctly shorter than

paddle.

LARVA (fig. 25). Head: 0.95 mm. Siphon: 0.95 mm. Anal Saddle: 0.35 mm.

Very similar to secundus; diagnostic characters as in the key. General chaeto-

taxy based on 7 reared specimens from French Guiana. Head: Integument light

yellow to brown; collar dark brown. Mental plate with 9 or 10 lateral teeth.

Hair 4(6;3-8); 5(9,10;7-12) with or without barbs; 6(2); 7(4-7) not barbed; 8

(1); 9(8,9;6-9); 10(1-2); 11(5); 12€1); 14(2); 15(2;2-3). Antenna: About 0.42 of

head length; hair 1(3;3-4). Thorax: Prothorax: hair 0(10-18) very short; 1(1);

2(1); 3(2;2-3); 4(2;1-2); 7(3;3-4); 8(3-8) short and weak; 9(1;1-2); 10(1); 11(1);

14(1). Mesothorax: hair 1(3;3-4); 2(1); 3(1); 4(231-2); 8(5,6;3-7); 9(6;5-6); 11

(1); 13(16-20); 14(7,8;6-9). Metathorax: hair 1(3,4;2-4); 2(1); 3(3,4;2-4); 4(3-

533-7); 5(2,3;2-4); 6(1); 7(6-8;4-8); 8(12-20); 9(5,6;3-6); 11(1); 13(2-4) variable

in development. Abdomen: Segment I: hair 1(4,5;3-5) very short; 6(2); 7(2).

Segment II: hair 1(1-2) moderate; 3(1;1-2) longer than hair 1; 6(2;1-2). Segment

III: hair 1(1) moderate, longer than hair 1-II; 3(1) longer than hair 1; 6(1); 13

Valencia: Subgenus Carrollia of Culex 59

(2;1-3) longer and stronger than hair 1. Segment IV: hair 1(1) longer than hair

1-III; 3(2) shorter than hair 1; 6(1); 13(2;1-2) shorter but stronger than hair 1.

Segment V: hair 1(1) longer and stronger than hair 1-IV; 3(1) strong but slight-

ly shorter than hair 1; 6(1); 13(2;2-3) shorter than hair 1. Segment VI: hair

1(1) subequal to hair 1-III; 6(1). Segment VII: hair 1(1). Segment VIII: Comb

scales (12-16) with lateral fringes indistinct; hair 1(6;5-6); 3(7-9); 5(3,4;2-4). Si-

phon: Pigmentation light brown. Index about 2.8-3.5. Pecten teeth (9-13) broad,

with indistinct fringes on ventral margin. Ventral siphonal tufts (10-12) usually

with 4 or 5 branches (3-6). Anal Segment: Saddle shallowly emarginate lateral-

ly; pigmentation light yellow to brown. Hair 1(3,4;2-5); 2(3,4;2-4). Ventral brush

with 4 pairs of hairs, majority usually with 4 or 5 branches (3-7). Gills about

2.0 of dorsal saddle length; tapered.

SYSTEMATICS. Culex bonnei differs conspicuously from the other 2 members

of the Iridescens Complex, secundus and iridescens, in the unique development

of the accessory division of the subapical lobe of the male genitalia and in the

presence of short broad scales on the midlobe of the scutellum of the adults.

The pupa appears to be more like iridescens than secundus while the reverse is

true of the larva. I interpret bonnei as the most derived and dominant recent

species of the complex. It occupies the central and most extensive part of the

distribution of the Iridescens Complex but has a broad sympatric area with sec-

undus in the west. It also has the widest range of breeding sites.

The only significant variation I have noted is in the color of the broad scales

of the midlobe of scutellum which are black in the typical populations in the

Guianas and brown in the outlying populations.

Culex bonnei has been frequently confused in the past with the other 2 mem-

bers of the complex as indicated in the taxonomic references and its incorrect

synonymy with iridescens by the Bonnes. Recently Barata and Cotrim (1972:23-24)

included a population of bonnei in the description of the pupa of their “secundus”

(see sp. 9a, Espirito Santo form).

Closely related to bonnei, and possibly merely a disjunct differentiated popu-

lation, is sp. 8a from the State of Bahia in Brazil. It differs from typical bonnei

in lacking short broad scales on the midlobe of the scutellum.

BIONOMICS. Culex bonnei has been reported from a wider range of breed-

ing sites than the other 2 species of the Iridescens Complex: broken or cut bam-

boo (4), treeholes (4), fallen palm spathes (5), fallen cacao pod (1), fallen fruit

(1), artificial containers, largely metal (6) and from an unspecified trap. Four

species of Carrollia have been recorded as breeding associates: bihaicolus (1),

metempsytus (1), infoliatus (2) and urichii (2); and 8 other species: Anopheles

(An.) eiseni (3); Culex (C.) mollis (1), C. (Microc.) stonei (1); Limatus asulleptus

(4), L. durhamii (4); Orthopodomyia fascipes (1); Sabethes undosus (1); and Wy-

eomyia aporonoma (1). There is 1 record of adults collected in nature, 1 male

and 1 female in a trap (probably Malaise) in Ecuador.

DISTRIBUTION (fig. 4). The Guianas, northern Amazon basin to State of

Maranhao in Brazil, upper Orinoco basin and Pacific drainage in central Colombia,

at elevations of 30-500 m.

Material examined: 695 specimens; 124 males, 148 females, 260 larvae, 162

pupae; 67 individual rearings (25 larval, 36 pupal, 6 incomplete).

BRAZIL. Para: Aura Reserve (BRA 3) [UCLA] . Igarape do Catu (BRA 5) [UCLA].

COLOMBIA. Boyaca: Remolinos, Finca Primavera (COB 56) [UCLA]. Meta: Restrepo (258,

259,261,280; KO 10-1,1041) [UCLA]. Retiro (KO 10-21,10-28) [UCLA] . Rio Guacavia (KO 31-

60 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

34) [UCLA]. Villavicencio (CV 1012,1014,1017,1018; C-30; 566,597) [UCLA, USNM]. Valle:

Rio Raposo (COL 2-4,6,34,44,65) [UCLA].

ECUADOR. Napo-Pastaza: Isla Pompeya (ECU 21) [UCLA]. Rio Napo (ECU 8) [UCLA].

FRENCH GUIANA. Guyane: Cogneau (FGC 3462,3472) [UCLA]. La Chaumiere (FGC 3577)

[UCLA]. Rochambeau (FGC 3536) [UCLA]. Stoupan (FGC 3288) [UCLA]. Jnini: Saut Tigre

(FG 179,180) [UCLA].

SURINAM. Suriname: Gansee, lectotype [USNM]. Locality not Specified: (4097,4098)

[UCLA] ; (BB 525) [USNM].

Additional Records From the Literature

BRAZIL. Maranhao: Imperatriz, Acailandia (Barata and Cotrim, 1972:25) [FH]. Para: Curra-

linho (Cerqueira, 1961:156). Marajo Island (Kumm and Novis, 1938:511).

FRENCH GUIANA. Guyane: Crique Anguille (Fauran, 1961:44). Inini: Saut Tigre. St. Elie.

St. Nazaire (Fauran, 1961:44).

Sa. Culex (Car.) sp., Bahia form

Fig. 4

Culex (Carrollia) bonnei of Forattini, Rabello and Cotrim (1970:36).

Culex (Carrollia) bonnei in part of Antunes and Ramos (1939:383); Lane and Whitman (1943:

397).

The 3 males and 3 females from Curipipe and Piraja, Bahia, Brazil, I have ex-

amined differ from typical bonnei in having long moderately wide curved pale

scales (as in iridescens) instead of short broad dark ones on the midlobe of the

scutellum. The genitalia of 2 males are similar to those of typical bonnei. The

larva of this population is unknown to me but may have been included by Antunes

and Ramos (1939:381-382) with specimens from Colombia in their description

of bonnei. The pupa has not been described.

This form may be only an isolated population of bonnei, possibly with some

introgression with iridescens. However, its taxonomic status cannot be determined

without examination of additional material of adults and careful study of the

immature stages. The latter were reported from treeholes by Antunes and Ramos

(1939:382).

9. Culex (Car.) secundus Bonne-Wepster & Bonne

Figs. 1,4,8,9,26,27 :

1920. Culex (Carrollia) secunda Bonne-Wepster and Bonne, 1920:170-171. TYPE: Canal Zone,

Panama; number of specimens and type not indicated in original description; 2 probable

male syntypes (Brug ledger numbers 4117 and 4118) labelled secundus not located

in ITH collection by Belkin (1968:20) [LU, possibly misplaced in ITH]. Additional

material from same series available in USNM for neotype designation.

Culex (Carrollia) secundus of Dyar (1923:179; 1925:176); Bonne and Bonne-Wepster (1925:

203-204); Edwards (1932:221); Lane (1939:82-83; 1953:506-508); Galindo, Carpenter and

Trapido (1951:104,105,108,110,111,126); Levi-Castillo (1952:553; 1953a:35); Barreto and

Reyes (1955:64); Galindo and Blanton (1955:73); Stone, Knight and Starcke (1959:283);

Belkin, Schick and Heinemann (1965:56); Belkin (1968b:20); Barreto and Lee (1969:434).

Culex (Carrollia) secundus in part of Forattini, Rabello and Cotrim (1970:47).

Valencia: Subgenus Carrollia of Culex 61

Culex (Carrollella) secundus of Dyar (1928:281-282,538); Komp (1936:64); Patino Camargo

(1937:245); Boshell Manrique (1938:416).

Culex (Carrollella) secundus in part of Komp (1936:64; 1956:39).

Culex (Carrollia) iridescens in part of Busck (1908:70); Dyar (1918:108).

Carrollia iridescens in part of Howard, Dyar and Knab (1912:figs. 10,113; 1915:462-464).

FEMALE (figs. 1,8,9). Wing: 2.8 mm. Proboscis: 1.9 mm. Forefemur: 1.8

mm. Abdomen: about 2.1 mm. Very similar to bonnei and iridescens; diagnos-

tic characters as in the key. General characters as for the subgroup; description

based on 4 specimens from Panama and the Canal Zone and 25 specimens from

Costa Rica. Head: Broad decumbent scales of vertex golden or whitish on disc,

dark along orbital bristles on midline in front; narrow decumbent scales very

sparse, golden; erect scales very numerous, light golden, present on anterior part

of vertex to orbital bristles where they are proclinate and longer than on pos-

terior part and occiput; lateral patch yellowish to whitish. Proboscis 1.02-1.05

of forefemur; labium entirely dark scaled. Palpus 0.19-0.20 of proboscis; entire-

ly dark scaled. Antenna 1.00-1.05 of proboscis; torus with 4-6 setae. Thorax:

Mesonotal and scutellar integument blackish; scales dark bronzy except for dark

coppery around prescutellar space; bristles bronzy. Anterior promontory with 2

or 3 bristles; prescutellar bristles 4-6. Midlobe of scutellum with 4 long bristles

and dark coppery linear to narrow curved scales; lateral lobe with 3 long bris-

tles and similar scales. Pleuron darkened to level of base of coxae; ppl, meron

and metameron dark, lower part of stp usually not darkened. Apn with 3-6 bris-

tles; ppn bristles 3-5; ppl bristles 3 or 4; stp with 4 or 5 lower posterior bris-

tles; pra bristles 5 or 6; umep bristles 5 or 6. Legs: Coxal integument light, ex-

cept usually extreme base of midcoxa; forecoxal scale patch with dark scales

distally, base with light golden scales. Forefemur predominantly dark scaled; light

at base, more extensively on lower posterior surface. Midfemur predominantly

dark scaled; light on extreme base and on posterior ventral streak not reaching

the middle; metallic spots well marked. Hindfemur dark scaled dorsally and on

apical ring which is extended as a streak on posterior surface to about 0.5 from

base; ventral light golden scaling extending about halfway dorsad on anterior

surface and joining the long preapical metallic streak; sometimes a poorly dif-

ferentiated submedian metallic spot or streak also present and joined to ventral

light scaling. Knee spots faintly indicated by a few pale scales on apices of fe-

mora. Abdomen: Tergites III-VII with extremely variable basal dorsal transverse

white or whitish bands not reaching light metallic spots; sometimes a few basal

whitish scales on midline of tergite II. Light metallic spots small on II-IV, sub-

basal on V; marginal bristles of tergites very sparse, very light golden. Sternites

largely light scaled, proximal silvery, distal light golden; IV-VII with apical dark

bands; bristles pale golden. Cibarial Armature: Cibarial bar with narrow elon-

gate lateral flange without lobe; 3 broad median cibarial teeth, without alter-

nating filamentous spicules; cibarial dome small, denticles largely rounded. Gen-

italia: Tergite [IX with 1 or 2 lateral bristles; insula with 15-22.

MALE. Wing: 2.8 mm. Proboscis: 2.1 mm. Forefemur: 1.9 mm. Abdomen:

about 2.7 mm. Very similar to bonnei and iridescens; diagnostic characters as

in the key. Essentially similar to the female except for usual sexual differences.

Proboscis 1.10 of forefemur; labium dark scaled dorsally, entirely white scaled

on ventral surface distad of false joint and on a short median streak proximad

of joint; a large ventral patch of long hairs distad of basal bristles; a pair of very

long hairs ventrally before apex, preceded by shorter hairs. Palpus 0.88 of pro-

62 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

boscis; dark scaled throughout. Antenna 1.00 of proboscis. Specimens studied:

2 from Panama and the Canal Zone and 22 from Costa Rica.

MALE GENITALIA (fig. 26). In general very similar to bonnei; differing pri-

marily in the following conspicuous features. Lobes of tergite [X more prom-

inent and separated by less than width of 1 lobe. Sidepiece without differenti-

ated apicotergal seta. Accessory division of subapical lobe much smaller, digiti-

form and with 1 specialized bristlelike apical seta and a few simple setae at base.

Setae a and b of proximal division broader, more strongly recurved; seta a less

than 0.5 of seta b. Distal division completely absent. Preapical lobe of clasper

with very short basal (lower) spicules and very long hairlike distal (upper) spic-

ules; seta a longer and seta b shorter, transparent and very difficult to see. Para-

proct crown usually with 3 (2-4) teeth.

PUPA (fig. 26). Abdomen: 2.60 mm. Trumpet: 0.60 mm. Paddle: 0.60 mm.

Apparently indistinguishable from antunesi and babahoyensis; diagnostic charact-

ers from other members of the subgroup as in the key. General chaetotaxy based

on 10 specimens from Panama. Cephalothorax: Pigmentation yellow to brown;

sculpturing weak, without definite pattern. All hairs shorter than trumpet, 1,3-

5,8,9-C subequal, 7-C the longest; 1(2); 2(2;2-4) subequal to hair 6; 3(131-2);

4(3;1-4); 5(3;1-4); 6(2,3;1-3); 7(3;1-4); 8(2;1-3) shorter than hair 1; 9(1;1-2).

Trumpet: Nearly uniform in width; index usually 10.4 (10.2-11.0); pinna small.

Pigmentation brown. Metanotum: Pigmentation yellow to light brown; sculptur-

ing uneven, weak; hair 10(3;2-4) slightly longer than others; 11(3;2-4); 12(2;2-

3). Abdomen: Pigmentation light yellow to brown, more or less uniform; sculp-

turing moderate. Segment I: hair 1(94-120) fan-shaped. Segment II: hair 1(8;

6-14) subequal to hair 5; 3(1;1-4); 4(4,5;3-6); 5(2;1-3). Segment III: hair 1(4;

2-8) slightly longer than hair 5; 3(1) strong, surpassing level of rudimentary spi-

racle IV; 4(2,3;2-4); 5(3,4;2-4). Segment IV: hair 1(4;2-4) subequal to hair 3;

3(4-6;4-8); 4(1) subequal to hair 6; 5(1) strong, surpassing level of rudimentary

spiracle VI. Segment V: hair 1(2;2-5) subequal to hair 3; 3(1;1-2); 4(6;4-8) short-

er than hair 3; 5(1) strong and long but not reaching caudal margin of tergite

VII. Segment VI: hair 1(1;1-2); 3(2;1-3); 4(4;2-6); 5(1) reaching level of alveolus

of hair 9-VII, considerably shorter than hairs 5-IV,V. Segment VII: hair 1(1);

3(131-2); 4(2;1-2); 5(3,4;1-4) very small; 6(6;4-12) short; 9(3;3-5) barbed. Seg-

ment VIII: hair 4(1;1-2); 9(7;5-9) barbed, subequal to paddle length.

LARVA (fig. 27). Head: 0.85 mm. Siphon: 1.25 mm. Anal Saddle: 0.30 mm.

Very similar to bonnei; diagnostic characters as in the key, not entirely reliable.

General chaetotaxy based on 4 reared and 6 other specimens from Panama and

Canal Zone. Head: Integument light yellow; collar dark brown. Mental plate with

9 or 10 lateral teeth. Hair 4(6-8;5-8); 5(8-10) barbed; 6(2,3;2-4); 7(5,6;4-8) with-

out barbs; 8(1;1-2) subequal to hair 7; 9(9,10;7-11); 10(2;1-3); 11(5,6;4-6); 12

(1); 14(2,3;2-4); 15(2). Antenna: About 0.45 of head length; hair 1(3;2-3). Tho-

rax: Prothorax: hair 0(10-14) moderate; 1(1); 2(1); 3(3;2-4); 4(2;1-2); 7(3;2-4);

8(6-12) weak; 9(1); 10(1-2); 11(2;1-2); 14(1). Mesothorax: hair 1(3,4;3-5); 2(1);

3(1); 4(1-2); 8(6;5-7); 9(5,6;4-6); 11(1;1-2); 13(10-20); 14(4-6;2-8). Metathorax:

hair 1(3;2-4); 2(1;1-2); 3(5,6;4-6); 4(4,5;3-5); 5(2;1-4); 6(1); 7(6-8;4-9); 8(11-20);

9(4;4-7); 11(1); 13(3,4;3-7). Abdomen: Segment I: hair 1(4,5;:4-6) very short;

6(2); 7(2;1-2). Segment II: hair 1(1;1-2) short and weak; 3(1-2) longer than hair

1; 6(2). Segment III: hair 1(1;1-2) moderate; 3(1) longer than hair 1; 6(1); 13

(2;1-2) subequal or shorter than hair 1. Segment IV: hair 1(1) subequal to hair

I-III or shorter; 3(1,2;1-3) shorter than hair 1; 6(1;1-2); 13(2;1-3) shorter than

Valencia: Subgenus Carrollia of Culex 63

hair 1. Segment V: hair 1(1) stronger and longer than hair 1-IV; 3(1) shorter

than hair 1; 6(1); 13(2;1-3) shorter than hair 1. Segment VI: hair 1(1) shorter

than hair 1-III; 6(1). Segment VII: hair 1(1). Segment VIII; Comb scales (11-

18) with indistinct fringes to apex; hair 1(6-8;6-10); 2(15;1-2); 3(9-11;6-11); 5

(2,3;2-4). Siphon: Pigmentation light brown. Index about 6.0-7.5; distal part of

siphon often sharply narrowed and upturned. Pecten teeth (10-16) short and

with indistinct fringe on ventral margin. Ventral siphonal tufts usually 10(8-12)

and mostly with 6 or 7 branches (3-8). Anal Segment: Saddle shallowly emargi-

nate laterally; pigmentation light yellow to brown. Hair 1(4,5;2-6); 2(3;3-4). Ven-

tral brush usually with 4 pairs of hairs, majority usually with more than 6 branches

(4-10). Gills about 2.0 of dorsal saddle length; tapered.

SYSTEMATICS. Culex secundus in several respects is intermediate between iri-

descens and bonnei, the other 2 members of the Iridescens Complex. In the male

genitalia, the accessory division of the subapical lobe is more strongly differenti-

ated from the proximal division than in iridescens and may be interpreted as an

intermediate step toward the condition in bonnei., Its larva is very similar to that

of bonnei but its pupa differs from the other 2 species of the complex in the

more elongate trumpet. The scales of the midlobe of the scutellum of the adults

are linear, a primitive character state. I interpret secundus as an early offshoot

of the ancestral stock of the Iridescens Complex which has undergone consid-

erable change recently but may have been the stock from which bonnei arose

in the more distant past. These 2 species are now broadly sympatric in Colom-

bia without any indication of introgression.

The siphon index is more variable in secundus than in the other 2 species but

I have found no geographical or ecological correlations for this variation.

Culex secundus has been generally recognized as a distinct species since the

original description by the Bonnes.

Apparently closely related to secundus on similar male ‘genitalic features is sp.

9a, Espirito Santo form (see). Because of the extremely wide separation of this

population in Brazil from the known distribution of typical secundus in north-

western South America and Central America and because of some differences in

the larva and pupa, it is probably a distinct species. Barata and Cotrim (1972:

18-25) treated this form and a population of bonnei as “‘secundus”’.

BIONOMICS. Culex secundus appears not to have as wide a range of breeding

sites as its close relative, bonnei. It has been found primarily in cut or broken

bamboo (19) and fallen leaves, palm spathes and fronds (10) but has also been

recorded from fallen monkey-pods (2), an unspecified artificial container (1) and

from a ground seepage (1). Two species of Carrollia have been recorded as breed-

ing associates, bihaicolus (1) and cerqueirai (1) and 11 other species: Culex (C.)

chidesteri (1), C. (C.) corniger (1); Haemagogus (S.) spegazzinii falco (1); Limatus

asulleptus (4), L. durhamii (5); Trichoprosopon (T.) digitatum (4), T. (T.) pallidi-

venter (4); Toxorhynchites (L.) sp. (1); Sabethes undosus (1); Wyeomyia aporonoma

(2), W. arthrostigma (1). There is no record of adults collected in nature.

DISTRIBUTION (fig. 4). From Costa Rica through Panama to Colombia and

Ecuador, including upper Orinoco drainage in Colombia and upper Amazon drain-

age in Ecuador, at elevations of 15-1700 m.

Material examined: 768 specimens; 175 males, 163 females, 264 larvae, 166

pupae; 123 individual rearings (71 larval, 49 pupal, 3 incomplete).

COLOMBIA. Antioquia: Providencia, Anori R. valley, C. Porter [UCLA]. Caldas: La Plata

(COM 48,49) [UCLA]. Huila: Santa Leticia (COL 18) [UCLA]. Meta: Bosque de San Jose,

64 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

W.H. Komp [USNM]. Restrepo (KO 1028,1032,3134,3137,3738; 233,584,10-2,10-4,1027,1037,

31-36) [UCLA, USNM]. Retiro, W.H. Komp [USNM]. Santander: Yarima (COB 74) [UCLA].

Valle: Rio Raposo (COL 18,21,23,72,74,78) [UCLA]. Locality not Specified: W.H. Komp

[USNM].

COSTA RICA. Heredia: Finca La Selva (CR 412) [UCLA]. Puerto Viejo (CR 191) [UCLA].

Limon: Westfalia (CR 72) [UCLA].

PANAMA. Bocas del Toro: Almirante (PA 259) [UCLA]. Filon (PA 187) [UCLA] . Quebrada

Grande (PA 654) [UCLA]. Canal Zone: Barro Colorado, Dyar and Shannon [USNM]; Galindo

[USNM]. Cano Saddle (P69) [USNM]. Erwin Island (P22) [USNM]. Fort Clayton (C 257)

[UCLA]. Fort Gulick, S.J. Carpenter [UCLA]. Madden Forest (PA 840) [UCLA]. Tabernilla,

A. Busck (99,166) [USNM]; A.H. Jennings (399,401,448,451) [USNM]. Locality not specified,

A.H. Jennings [USNM]. Darien: Alturas de Nique (PA 1059) [UCLA]. La Laguna (PA 449)

[UCLA]. Tacarcuna (GG 113, PA 446) [UCLA] . Panama: Upper Pequini (PA 939,940) [UCLA].

Additional Records From the Literature

ECUADOR. Napo-Pastaza: Arajuno. Puyo. Shell-Mera (Levi-Castillo, 1952:553).

9a. Culex (Car.) sp., Espirito Santo form

Fig. 4

Culex (Carrollia) secundus of Lane and Whitman (1943:394-396).

Culex (Carrollia) secundus in part of Forattini, Rabello and Cotrim (1970:47); frat and Cotrim

(1972:18-25).

In the recent description of “secundus’’ from Brazil by Barata and Cotrim (1972:

18-23) 2 populations were included, from Sao Joao de Petropolis in the State of

Espirito Santo and from near Imperatriz in the State of Maranhao. In my opin-

ion neither population can be assigned to secundus. The descriptions of the adults,

male genitalia, larva and pupa appear to be based primarily on the population

from Espirito Santo, described earlier in all stages by Lane and Whitman (1943:

394-396) with illustrations of the male genitalia and head and terminal segments

of the larva. The adults were reared from a collection in bamboo but it is not

known if the association of the stages was through individual rearings. As des-

cribed and figured in both papers the male genitalia of this population appear to

be extremely similar to those of typical secundus but there are a number of dif-

ferences in the external features of the adults as well as the larva and pupa from

this species. A decision as to the taxonomic status of this population cannot be

made without careful reexamination of the existing material and study of addi-

tional specimens, particularly of immature stages. Although it is possible that it

is only a relict isolated population of secundus, the fact that it is widely separated

from the main populations of this species by bonnei suggests that it is a distinct

species.

The material from the State of Maranhao included by Barata and Cotrim in

“secundus’’ appears to be typical bonnei, judging by the description and figures

of the pupa.

10. Culex (Car. ) iridescens Lutz

Figs. 4,28,29

1904. Carollia iridescens Lutz, 1904:16. Nomen nudum.

Valencia: Subgenus Carrollia of Culex 65

1905. Carrollia iridescens Lutz, 1905:81-82. TYPE: Neotype male with genitalia on slide

(222), Serra da Cantareira, Sao Paulo, Brazil, Apr 1938, M. Sanchez, E. Coimbra and

H. Guimaraes [FH, 953; designation by Antunes and Ramos, 1939:380-381].

Culex (Carrollia) iridescens of Antunes and Ramos (1939:374-381); Lane (1939:82; 1953:498-

500); Lane and Whitman (1943:394); Senevet and Abonnenc (1958:288); Cova Garcia, Sutil

and Rausseo (1966a:28,115; 1966b:42-43 141,231,344); Belkin (1968b:16); Forattini, Rabello

and Cotrim (1970:42); Belkin, Schick and Heinemann (1971:27-28).

Culex (Carrollia) iridescens in part of Edwards (1932:221); Stone, Knight and Starcke (1959:

283).

Culex (Carrollella) iridescens in part of Dyar (1928:280-281).

Carrollella iridescens in part of Lutz (1921:163); Shannon (193 1a:8).

Carrollia iridescens in part of Theobald (1907:207-209);. Peryassu (1908:34,44,167-169,343);

Surcouf and Gonzalez-Rincones (1911:144); Lutz (1919:164).

Carollia iridescens of Blanchard (1905:634).

FEMALE. Wing: 3.5 mm. Proboscis: 2.4 mm. Forefemur: 2.3 mm. Abdomen:

about 2.8 mm. Very similar to bonnei and secundus; diagnostic characters as in

the key. Differing from secundus primarily in the following features. Descrip-

tion and measurements based on 2 pinned neoparatypes from Brazil. Head: Broad

decumbent scales of vertex dark golden anteriorly in the middle, whitish caudad

and laterally. Proboscis 0.98-1.04 of forefemur. Palpus 0.16 of proboscis. An-

tenna 0.95 of proboscis. Thorax: Mesonotal vestiture denser, from dorsal aspect

appearing largely light coppery to dark golden with narrow dark bronzy sides.

Midlobe of scutellum with dense vestiture of long, moderately wide curved light

golden scales, occasionally a few shorter broader scales. Meron not darkened.

Apn with 3-6 bristles; ppl bristles 4 or 5; stp with 3-5 lower posterior bristles;

pra bristles 4 or 5. Legs: Forecoxal scale patch without dark scales. Forefemur

more extensively pale on lower anterior surface and with posterior pale scaling

nearly reaching apex. Midfemur pale on lower anterior surface to level of sub-

median metallic spot which may be connected; posterior surface pale to near

apex. Hindfemur with less extensive dark scaling on posterior surface. Abdo-

men: Dorsal basal tergal light bands on III-VII much broader, whiter and very

conspicuous; tergite I] with a similar basal whitish band. Metallic spots larger

and more conspicuous on II-IV. Bristles of tergites and sternites longer. Cibarial

Armature and Genitalia: Not studied.

MALE.: Wing: 3.4 mm. Proboscis: 2.5 mm. Forefemur: 2.5 mm. Abdomen:

about 2.6 mm. Very similar to bonnei and secundus; diagnostic characters as in

the key. Essentially similar to the female except for usual sexual differences.

Differing from secundus primarily in the following features. Proboscis 1.00-1.04

of forefemur; labium white scaled both ventrally and dorsally proximad of false

joint; dark scaled distad on dorsal surface; distal ventral surface largely pale scaled

but darkened toward apex and with numerous long hairs throughout. Palpus 0.92

of proboscis; segment 3 with conspicuous ventromesal line of white scales. An-

tenna 0.92 of proboscis. Specimens studied: 2 pinned neoparatypes from Brazil.

MALE GENITALIA (fig. 28). In general very similar to bonnei and secundus;

differing from the former in the following conspicuous features. Lobes of ter-

gite IX more prominent and separated by less than width of 1 lobe; each lobe

with 4-12 bristles. Accessory division of subapical lobe very broad and joined

to proximal division, with several (7) simple marginal setae. Setae a and b of prox-

imal division strongly dilated apically, seta a longer than seta b. Preapical lobe

of clasper with shorter basal (lower) spicules. Distal part of proctiger apparently

narrower and tapering apically.

66 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

PUPA (fig. 28). Abdomen: 3.20 mm. Trumpet: 0.55 mm. Paddle: 0.65 mm.

Very similar to bonnei and soperi; diagnostic characters as in the key. General

chaetotaxy based on 1 specimen from Brazil. Cephalothorax: Pigmentation light

yellow to brown; sculpturing weak. All hairs shorter than trumpet, weak or mod-

erate, 5,7-C subequal, 1-C the longest; 1(2); 2(2,3); 3(2) longer than hairs 2,4,6;

4(3); 5(3,4); 6(2); 7(2,3); 8(2) shorter than hair 9; 9(missing). Trumpet: Nearly

uniform in width; index 6.1; pinna small. Pigmentation light brown. Metanotum:

Pigmentation light yellow; sculpturing uneven, weak; hair 10(2); 11(2); 12(1)

longer than others. Abdomen: Pigmentation light yellow, anterior mesal areas

of tergites [I-IV slightly darker; sculpturing weak. Segment I: hair 1(88-90) fan-

shaped. Segment II: hair 1(3) subequal to hair 3; 3(1) moderate, reaching level

of rudimentary spiracle III: 4(4,5); 5(1). Segment III: hair 1(1); 3(missing); 4(3);

5(1,2). Segment IV: hair 1(1); 3(5); 4€1); 5(1) surpassing level of rudimentary

spiracle V but not reaching caudal margin of tergite. Segment V: hair 1(1) sub-

equal to hair 1-IV; 3(1); 4(5); 5(1) subequal to hair 5-IV. Segment VI: hair

1(1) small; 3(2); 4(2,3) shorter than hair 3; 5(1) short, about 0.5 of hair 5-V.

Segment VII: hair 1(1) longer than hair 3; 3(1,2); 4(2); 5(6,7) very short; 6(5);

9(4) barbed. Segment VIII: hair 4(1 ,2); 9(missing).

LARVA (fig. 29). Head: 0.90 mm. Siphon: 1.25 mm. Anal Saddle: 0.35 mm.

Apparently differentiated from other members of the subgroup primarily by the

distinct marginal denticles of the pecten teeth. General chaetotaxy based on 1

reared specimen from Brazil and Antunes and Ramos (1939). Head: Integument light

yellow; collar brown. Mental plate with 9 or 10 lateral teeth. Hair 4(7-11) un-

usually long; 5(6,7;6-9) barbed; 6(2); 7(8); 8(1) subequal to hair 7; 9(8,9); 10

(1,2); 1109); 12(1); 14(4); 15(1,2). Antenna: About 0.36 of head length; hair

1(2,4). Thorax: Prothorax: hair 0(11,12) relatively long, stellate; 1(1); 2(1); 3

(3); 4(2); 7(3,4); 8(12,16) strong and stellate; 9(1); 10(missing); 11(2); 14(1).

Mesothorax: hair 1(5); 2(2); 3(1); 4(3); 8(6,7); 9(5,6); 11(2); 13(12,16) rela-

tively long; 14(11,12) long and stellate. Metathorax: hair 1(3,4); 2(2); 3(5); 4

(6); 5(3); 6(1); 7(6,7); 8(20,22); 9(5,6); 11(2); 13(10,11) long and stellate. Ab-

domen: Segment I: hair 1(4) short; 6(2); 7(2). Segment II: hair 1(1,2) strong

and long; 3(2,3) short; 6(2). Segment III: hair 1(1) very long and strong; 3(1)

moderately long; 6(1); 13(3) much shorter than hair 1. Segment IV: hair 1(1,

2) subequal to hair 1-III; 3(3) short; 6(1); 13(3) subequal to hair 13-III. Seg-

ment V: hair 1(1) longer than hair 1-II,[V; 3(1) long; 6(1); 13(3) subequal to

hair 13-II1,[1V, Segment VI: hair 1(1,2) moderate and weak; 6(1). Segment VII:

hair 1(2). Segment VIII: Comb scales (14,16) without lateral fringes; hair 1(6);

3(8); 5(5,6). Siphon: Pigmentation light brown. Index about 4.0. Pecten teeth

(14,18) short, with distinct denticles on ventral margin. Ventral siphonal tufts

(10) with 5-7 branches. Anal Segment: Saddle pigmentation light yellow. Hair

1(5,6); 2(4,5). Ventral brush with 4 pairs of hairs, all missing. Gills less than

2.0 of dorsal saddle length, slightly tapered.

SYSTEMATICS. Culex iridescens, the type species of Carrollia, is interpreted

here as the earliest derivative of its complex, which includes also bonnei and

secundus. In the male genitalia it has the primitive type of accessory division

of the subapical lobe that is poorly differentiated from the proximal division.

The larva also has a primitive feature in the branched hair 1-VII. The pupa is

very similar to the other species of the complex and is similar to bonnei in the

retention of the short trumpet. The scaling of the midlobe of the scutellum in

both sexes is intermediate between the primitive state in secundus and the highly

Valencia: Subgenus Carrollia of Culex 67

derived one in bonnei. However, external features of the males show considerable

departure from the primitive states in the white scaling of the palpus and the

development of long hairs distad of the false joint of the labium (shared with

secundus). Overall, iridescens appears to be much more closely related to secundus

than to bonnei. Its limited range is at the southern extremity of the distribution

of the complex, opposite to that of secundus. In the northern part of this range

it is sympatric with sp. 9a, which is undoubtedly related to secundus.

In the very limited material of iridescens I have examined no significant vari-

ation is apparent. The key characters for the pupa and larva may not be reliable

as they are both based on a single specimen.

BIONOMICS. I have seen adults of iridescens reared from cut or broken bam-

boo (2) and a treehole (1) only, but Forattini (1965:34) states that this species

breeds also in bromeliads, flower bracts, fallen leaves, artificial containers and

ground pools. I have seen no records of breeding associates or adults collected

in nature.

DISTRIBUTION (fig. 4). Coastal and Parana drainages in Brazil southward from

State of Espirito Santo to State of Parana at elevations of 50-100 m.

Material examined: 7 specimens; 3 males, 2 females, 1 larva, 1 pupa; 1 individual

larval rearing.

BRAZIL. Parana: Locality not specified (12148) [BH]. Sao Paulo: Perus, Agua Fria (1171-11)

[USNM]. Serra da Cantareira (1173) [USNM]; M. Sanchez, E. Coimbra and H. Guimaraes, neo-

type [FH].

Additional Records From the Literature

BRAZIL. Espirito Santo: Sao Joao de Petropolis [FH, 3828] (Forattini, Rabello and Cotrim,

1970:42). Guanabara: Rio de Janeiro [FH, 3827] (Forattini, Rabello and Cotrim, 1970:42);

Estrada Dona Castorina; Alto da Boa Vista (Antunes and Ramos, 1939:381). Parana: near Porto

Mojoli (Lutz, 1919:164). Sao Paulo: Perus, Agua Fria [FH, 953-971] (Forattini, Rabello and

Cotrim, 1970:43). Serra da Cantareira [FH, 972,973] (Forattini, Rabello and Cotrim, 1970:43).

11. Culex (Car.) soperi Antunes & Lane

Figs. 5,30,31

1937. Culex (Carrollia) soperi Antunes and Lane, 1937:21-23. TYPE: Holotype male (886)

with associated genitalia on 2 slides (99,100), Perus, Sao Paulo, Brazil, 23 Apr 1937

[FH, 721].

Culex (Carrollia) soperi of Antunes and Lane (1938:1037); Antunes and Ramos (1939:383-

385); Lane and Whitman (1943:393-394); Lane (1953:503-505); Senevet and Abonnenc

(1958:287-288); Stone, Knight and Starcke (1959:283); Casal and Garcia (1968:97-98); For-

et Rabello and Cotrim (1970:48); Stone (1970:165); Belkin, Schick and Heinemann

1971:28).

FEMALE. Wing: 3.1 mm. Proboscis: 1.9 mm. Forefemur: 2.0 mm. Abdomen:

about 2.6 mm. Very similar to antunesi, babahoyensis and kompi; mesonotal vest-

iture very similar to iridescens; diagnostic characters as in the key. In general

as described for secundus, differing primarily in the following features. Descrip-

tion and measurements based on 1 dry specimen from Teresopolis, Brazil, sup-

plemented by original description of Antunes and Lane (1937:21-22). Head: Broad

decumbent scales of vertex yellowish to tan; narrow decumbent scales apparent-

ly absent; erect scales golden yellow, short, confined to posterior part of vertex

and occiput, sometimes a few short ones on disc of vertex but distant from or-

68 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

bital line; lateral patch yellowish white. Proboscis 0.95 of forefemur; labium

slightly paler ventrally. Palpus 0.21 of proboscis. Antenna 0.97 of proboscis.

Thorax: Mesonotal vestiture very dense, light coppery to dark golden, lighter

in posterior part. Midlobe of scutellum with short broad pale scales; lateral lobe

with linear scales. Meron not darkened. A few broad pale scales in front of low-

er posterior bristles of stp. Legs: Forecoxal scale patch without dark scales. Fore-

femur more extensively pale on anterior surface and pale to apex on posterior

surface. Midfemur pale on lower anterior surface to near level of submedian me-

tallic spot; posterior surface pale to apex. Hindfemur with dark scaling more re-

stricted, absent from base of dorsal margin; light scaling extending ventrally on

both anterior and posterior surface to near apex. Tibiae and basal tarsal segments

indistinctly paler ventrally. Abdomen: Dorsal basal tergal light bands on III-VII

much broader, yellowish; tergite II with a smaller basal yellowish band. Metal-

lic spots larger and more conspicuous on II-IV. Sternite IV with only a few dark

scales apically not forming a band. Cibarial Armature and Genitalia: Not studied.

MALE. Wing: 3.1 mm. Proboscis: 2.4 mm. Forefemur: 2.1 mm. Abdomen:

about 2.4 mm. Very similar to antunesi, babahoyensis, cerqueirai, kKompi and wil-

soni; Diagnostic characters as in the key. Essentially similar to the female and

differing from secundus in the same general features. Proboscis about 1.1 of fore-

femur; labium without distinct white scaling but paler at base dorsally and ven-

trally and beyond false joint to before apex ventrally, without any long hairs.

Palpus 0.91 of proboscis; without white scales on segment 3. Antenna 0.9 of

proboscis. Only 1 dry specimen studied, from Perus, Brazil.

MALE GENITALIA (fig. 30). In general as described for antunesi but differ-

ing conspicuously in the following features. Lobes of tergite [IX more promi-

nent and more approximated. Sidepiece short, ovoid; apicotergal setae 2, weak- -

ly developed. Setae a and b of proximal division of subapical lobe both short

and with broadly expanded, distinctly ribbed distal part; seta c shorter. Acces-

sory division with a digitiform bare ventral portion from base of which projects

a very large dorsal lobe with expanded apex bearing numerous flattened setae

acuminate apically and some simple setae on more ventral part. Distal part of

proctiger relatively broad and rounded; paraproct crown with 3 or 4 teeth; cer-

cal setae 2 or 3.

PUPA (fig. 30). Abdomen: 3.05 mm. Trumpet: 0.53 mm. Paddle: 0.63 mm.

Very similar to bonnei and iridescens; diagnostic characters as in the key. Gen-

eral chaetotaxy based on 1 specimen from Brazil. Cephalothorax: Pigmentation

light yellow; sculpturing uneven, weak. All hairs shorter than trumpet; 3-5,8-C

subequal; 1(2); 2(2) shorter than hair 6; 3(2); 4(2,3); 5(3); 6(3); 7(2,3); 8(2);

9(1).. Trumpet: Nearly uniform in width; index 7.1; pinna small. Pigmentation

light brown. Metanotum: Pigmentation light yellow; sculpturing uneven, weak;

hairs subequal, 10(1); 11(1); 12(2). Abdomen: Pigmentation light yellow, anter-

ior mesal area of tergites II-IV slightly darker; sculpturing moderate. Segment

I: hair 1(145-150) fan-shaped. Segment II: hair 1(6,8) slightly longer than hair

5; 3(1) not reaching level of rudimentary spiracle III; 4(3) very short; 5(2). Seg-

ment III: hair 1(3) short; 3(1) surpassing level of rudimentary spiracle IV; 4(1)

subequal to hair 6; 5(3) short. Segment IV: hair 1(1); 3(8,10) dendritic; 4(1);

5(1) surpassing level of rudimentary spiracle V. Segment V: hair 1(2) longer than

hair 6; 3(2); 4(4,5) shorter than hair 1; 5(1) surpassing level of rudimentary spi-

racle VI. Segment VI: hair 1(1,2); 3(1); 4(5) subequal to hair 6; 5(1) weak, not

reaching level of rudimentary spiracle VII. Segment VII: hair 1(2) forked, sub-

Valencia: Subgenus Carrollia of Culex 69

equal to hair 4; 3(1); 4(2,3); 5(2) shorter than hair 6; 6(2,3); 9(4) short, weak

and without barbs. Segment VIII: hair 4(1); 9(9,10) barbed, distinctly shorter

than paddle.

LARVA (fig. 31). Head: 0.80 mm. Siphon: 1.36 mm. Anal Saddle: 0.45 mm.

In general similar to iridescens but resembling kompi in the development of ab-

dominal hairs 6-III-VI. General chaetotaxy based on 1 reared specimen from Bra-

zil and the original description. Head: Integument light yellow; collar dark brown.

Mental plate with 8 or 9 lateral teeth. Hair 4(10-12); 5(8-10); 6(2-4); 7(5,6); 8

(2) longer than hair 7; 9(7,8); 10(1); 11(6,8); 12(1); 14(3,4); 15(2). Antenna:

About 0.40 of head length; hair 1(2-5). Thorax: Prothorax: hair 0(7,12) relatively

long; 1(1); 2(1); 3(4) relatively short; 4(2); 7(6); 8(16,18) very strong, stellate

and with very large tubercle; 9(1); 10(2,3); 11(1); 14(1). Mesothorax: hair 1(4);

2(3,4); 3(1,2); 4(3); 8(5,6); 9(4,5); 11€1); 13(16,18); 14(10) moderately long

and stellate. Metathorax: hair 1(4); 2(2); 3(1,3); 4(6); 5(1); 6(1); 7(6); 8(24,

26); 9(4,5); 11(1); 13(9,10) moderately long and stellate. Abdomen: Segment

I: hair 1(6,8) short; 6(2); 7(2). Segment II: hair 1(3) strong and long; 3(2) short;

6(2,3). Segment III: hair 1(2,3) strong and long; 3(1) moderate; 6(2); 13(4) sub-

equal to hair 1. Segment IV: hair 1(3) subequal to hair 1-III; 3(2) moderate;

6(2); 13(3,4) subequal to hair 1. Segment V: hair 1(2) longer than hair 1-IV;

3(1) long; 6(2); 13(5,6) subequal to hair 13-IV. Segment VI: hair 1(1) weak;

6(2). Segment VII: hair 1(2) very strong and long. Segment VIII: Comb scales

(14) relatively short and without fringes; hair 1(4,6); 3(4,6); 5(4). Siphon: Pig-

mentation light brown. Index about 4.4. Pecten teeth (7-16) very short and with

fine fringes on ventral margin. Ventral siphonal tufts (7-11) with variable num-

ber of branches (4-9). Anal Segment: Saddle pigmentation yellow. Hair 1(4,5);

2(3). Ventral brush with 4 pairs of hairs, all with 4 or 5 branches. Gills shorter

than dorsal saddle length, tapered.

SYSTEMATICS. The ovoid sidepiece of the male genitalia of soperi is a de-

rived character found elsewhere in the Antunesi Complex only in kompi but is

characteristic of the entire Iridescens Complex. The accessory division of the sub-

apical lobe superficially resembles that of babahoyensis but is more strongly dif-

ferentiated. The linear scaling of the midlobe of the scutellum of the adults is

a primitive state not known elsewhere in the complex. The branching of hair 6

on abdominal segments III-V is another primitive feature, retained in the com-

plex only in kompi. The pupa also has retained a primitive short trumpet, unique

in the complex. I interpret soperi as an ancient segregate as the southern extrem-

ity of the distribution of the Antunesi Complex with possible affinity with the

more advanced babahoyensis which occupies the northernmost part of the range.

The features shared by soperi and kompi were probably derived independently

but the similarity with iridescens may be due to introgression with sympatric

members of the Iridescens Complex.

Because of the very limited material of soperi I have seen, variation in this

species could not be determined and it is possible that the diagnostic characters

of the larva and pupa, based on single specimens, may not be entirely reliable.

BIONOMICS. Immature stages of soperi have been recorded only from cut or

broken bamboo (Antunes and Ramos, 1939:385). The original collection of 1

male and 2 females was obtained from human bait (Antunes and Lane, 1937:

22) and 1 adult was netted over grass in Argentina (Casal and Garcia, 1968:97).

DISTRIBUTION (fig. 5). Coastal and Parana drainages southward from State of

Espirito Santo in Brazil to Province of Misiones in northern Argentina, at eleva-

tions below 200 m.

70 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

Material examined: 5 specimens; 2 males, 1 female, 1 larva, | pupa; 1 individual

larval rearing.

BRAZIL. Rio de Janeiro: Terezopolis (27866-1) [BH]. Sao Paulo: Perus, Agua Fria (1171-

14) [USNM] ; holotype [FH].

Additional Records From the Literature

ARGENTINA. Misiones: Mato Quemado (Casal and Garcia, 1968).

BRAZIL. Espirito Santo: Sao Joao de Petropolis (Lane and Whitman, 1943). Sao Paulo: Perus

[FH, 721-723; 974-981] (Forattini, Rabello and Cotrim, 1970:48).

12. Culex (Car.) babahoyensis Levi-Castillo

Figs. 5,9,32,33

1953. Culex (Carrollia) babahoyensis Levi-Castillo, 1953a:35 (January). Nomen nudum.

1953. Culex (Carrollia) babahoyensis Levi-Castillo, 1953b:92-94 (July); 1953c:161-163(Aug.).

TYPE: Holotype male genitalia on slide, Hacienda ‘‘Mora’’, Juan Montalvo, Los Rios,

Ecuador, R. Levi-Castillo [USNM, 61639].

Culex (Carrollia) babahoyensis of Stone and Knight (1957:58); Stone, Knight and Starcke (1959:

282); Belkin, Schick and Heinemann (1965:21-22).

FEMALE (fig. 9). Wing: 2.4 mm. Proboscis: 1.7 mm. Forefemur: 1.5 mm.

Abdomen: about 2.3 mm. Not distinguishable with certainty from antunesi; very

similar to kKompi and soperi; diagnostic characters as in the key. In general as

in secundus, differing primarily in the following features. Description based on

10. topotypic and 5 other specimens from Ecuador and 8 from Nicaragua. Head:

Broad decumbent scales of vertex appearing largely pale from above except usu-

ally darkened near orbital bristles, and largely dark from anterior aspect except

for a few light ones caudad; narrow decumbent scales usually completely ab-

sent; erect scales golden yellow, short, confined to posterior part of vertex and

occiput, sometimes a few short ones on disc of vertex but distant from orbital

line; lateral patch yellowish white. Proboscis 1.06-1.13 of forefemur; labium slight-

ly paler ventrally. Palpus 0.20-0.21 of proboscis. Antenna 1.00-1.06 of proboscis;

torus with 6-10 setae. Thorax: Integument of mesonotum and pleuron black to

very dark brown. Mesonotal vestiture moderately dense, dark coppery to bronzy,

usually paler in and around prescutellar space. Midlobe of scutellum with mod-

erately long broad scales, usually concolorous with scales of prescutellar space;

lateral lobe with longer broad scales concolorous with those of midlobe or of-

ten dark bronzy to black. Lower edge of dark part of pleuron sharply marked

in a straight line, lower part of stp usually not infuscated; meron not darkened.

A few broad pale scales in front of lower posterior bristles of stp. Legs: Coxal

integument usually whitish; scale patches silvery or with slight yellowish tinge;

forecoxal scale patch without dark scales. Forefemur with pale scaling of an-

terior surface extending to about 0.25 or more. Midfemur light scaled to about

0.25 on anterior surface and usually to near apex on posterior. Hindfemur light

at base to at least 0.25; posterior light scaling more extensive. Abdomen: Dor-

sal basal tergal light bands yellowish, usually present only on V-VII but often

at least indicated by some pale scales on IV and rarely on III. Metallic light

spots larger and more conspicuous on IIJ-IV. Sternites V-VII only with apical dark

bands, a few dark apical scales sometimes present on IV. Cibarial Armature: Ci-

Valencia: Subgenus Carrollia of Culex 71

barial bar with 4 broad median teeth; cibarial dome with triangular denticles.

Genitalia: Tergite IX with 2-4 lateral bristles; insula with 20-26.

MALE. Wing: 2.3 mm. Proboscis: 1.8 mm. Forefemur: 1.7 mm. Abdomen:

about 2.1 mm. Very similar to antunesi, cerqueirai, kompi, soperi and wilsoni;

diagnostic characters in key not entirely reliable except for soperi. Essentially

similar to the female and differing from secundus and soperi by the same gen-

eral features. Proboscis 1.1-1.2 of forefemur; labium more or less uniformly dark

scaled dorsally and with a narrow light scaled line from base to near apex ven-

trally. Palpus 0.78-0.83 of proboscis; without white scales on segment 3. An-

tenna 0.9 of proboscis. Specimens studied: 2 topotypic and 3 others from Ecua-

dor, 4 from Costa Rica and 4 from Nicaragua. Measurements from dry specimens.

MALE GENITALIA (fig. 32). In general as described for antunesi but differ-

ing conspicuously in the following features. Lobe of tergite IX usually with 3

bristles (1-4). Setae a and b of proximal division of subapical lobe much narrow-

er, subequal and not contorted; seta c a simple bristle. Distal division process more

prominent and slender. Accessory division very strongly developed, broad at base

and with a greatly expanded apex covered with numerous long simple bristles

projecting dorsad of proximal division. Seta a of clasper a simple bristle.

PUPA (fig. 32). Abdomen: 2.50 mm. Trumpet: 0.50 mm. Paddle: 0.35 mm.

Apparently indistinguishable from antunesi and secundus; diagnostic characters

from other members of the subgroup as in the key. General chaetotaxy based

on 10 specimens from Ecuador. Cephalothorax: Pigmentation light yellow; sculp-

turing weak, without definite pattern. All hairs shorter than trumpet, 2,6-9-C

subequal, 5-C the longest; 1(2;1-2) shorter than hair 3; 2(2;2-3); 3(3;2-3); 4G;

2-4); 5(3;2-6); 6(3;1-3); 7(2;2-3); 8(2;1-4); 9(2;2-4). Trumpet: Nearly uniform

in width; index about 12.0 (11.0-13.7); pinna small. Pigmentation yellow to brown.

Metanotum: Pigmentation yellow to dark brown, sculpturing weak; hairs sub-

equal, 10(2,3); 11(3;2-5); 12(2;2-4). Abdomen: Pigmentation light yellow, anter-

ior mesal areas of tergites II-IV slightly darker; sculpturing very weak. Segment

I: hair 1(40-64) fan-shaped. Segment II: hair 1(3,4;3-9) longer than hair 5; 3(1)

surpassing level of rudimentary spiracle III; 4(3,4;2-5); 5(2;1-2). Segment III: hair

1(2,3;2-5); 3(1) surpassing level of rudimentary spiracle IV; 4(1,2); 5(3;2-4). Seg-

ment IV: 1(2,3;1-5); 3(5,6;4-8); 4(1,2); 5(1,2) surpassing caudal margin of ter-

gite V. Segment V: hair 1(1) longer than hair 3; 3(1,2); 4(4,5;4-6); 5(1) sur-

passing level of rudimentary spiracle VII. Segment VI: hair 1(1,2;1-3); 3(1;1-2);

4(4;3-4); 5(1) surpassing level of rudimentary spiracle VII. Segment VII: hair 1

(1;1-2) longer than hair 3; 3(1,2); 4(2); 5(2;1-4) shorter than hair 6; 6(4,5;1-6);

9(2,3;2-4) poorly developed, shorter than hair 1. Segment VIII: hair 4(1;1-2);

9(7,8;5-10) barbed, subequal to paddle.

LARVA (fig. 33). Head: 0.80 mm. Siphon: 1.25 mm. Anal Saddle: 0.30 mm.

Very similar to antunesi and cerqueirai; diagnostic characters as in the key. Gen-

eral chaetotaxy based on 8 reared and 2 other specimens from Ecuador. Head:

Integument light yellow to brown; collar dark brown. Mental plate with 8 or 9

lateral teeth. Hair 4(8-10;7-10); 5(8-10;7-11); 6(3-5); 7(4,5;3-5); 8(1;1-2) subequal

to hair 7; 9(8-11;8-13); 10(1;1-2); 11(6-9); 12(1); 14(3,4;2-4); 15(1;1-2). Anten-

na: About 0.50 of head length; hair 1(3;2-5); Thorax: Prothorax: hair 0(10-12)

relatively long; 1(1); 2(1); 3(2,3); 4(2); 7(3;2-4); 8(10,11;6-12) moderately long,

strong and stellate; 9(2;1-2); 10(2;2-3); 11(1); 14(1). Mesothorax: hair 1(4;3-5);

2(231-3); 3(1); 4(2,3;2-4); 8(4,5;4-6); 9(4-6;4-7); 11(1); 13(11-20); 14(10-16) mod-

Te Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

erately long and stellate. Metathorax: hair 1(3,4;3-5); 2(2;1-2); 3(4;4-6); 4(4,5;

3-5); 5(1); 6(1); 7(5-7;4-8); 8(11-24); 9(4,5); 11(€1); 13(6-9) moderately long and

stellate. Abdomen: Segment I: hair 1(6-8) short; 6(2;2-3); 7(2;2-3). Segment II:

hair 1(1;1-2) strong and long; 3(2;1-2) short; 6(2;2-3). Segment III: hair 1(1,2;

1-3) strong and long; 3(1) long but shorter and weaker than hair 1; 6(1;1-2);

13(1-3;1-4) subequal to hair 1. Segment IV: hair 1(1,2;1-4) shorter and weaker

than hair 1-III; 3(2;1-2) relatively short, weaker than hair 3-III; 6(1;1-2); 13(Q,

3;1-4) subequal to hair 1. Segment V: hair 1(1;1-2) very long and strong; 3(1)

subequal to hair 3-IV; 6(1;1-2); 13(2-4;1-5) subequal to hair 13-IV. Segment VI:

hair 1(1) weak and relatively short; 6(1;1-2). Segment VII: hair 1(2;1-2) very

strong and long. Segment VIII: Comb scales (8-12) long, fringes restricted to

base; hair 1(4-6;4-8); 3(7-11); 5(3;3-5). Siphon: Pigmentation light yellow to brown.

Index about 4.1-5.5. Pecten teeth (7-15) moderately long and without fringes.

Ventral siphonal tufts (8-10) mostly with 5 or 6 branches (4-8). Anal Segment:

Saddle pigmentation light yellow to brown. Hair 1(5,6;4-8); 2(4;3-4). Ventral

brush with 4 pairs of hairs, all usually with 5-7 branches (3-7). Gills shorter than

dorsal saddle length, tapered.

SYSTEMATICS. The accessory division of the subapical lobe of the male geni-

talia of babahoyensis is superficially very similar to soperi but is less differenti-

ated and the sidepiece itself is of the primitive conical type as in all other mem-

bers of the Antunesi Complex except kompi and soperi. In all other respects

babahoyensis has derived character states when compared with soperi: (1) broad

scales on the midlobe of the scutellum of the adults, (2) elongate trumpet in the

pupa and (3) unbranched hair 6 on abdominal segments III-V of the larva. I have

not been able to separate the pupa from antunesi or from secundus of the Iri-

descens Complex and the larva is very similar to antunesi and cerqueirai. | am

tentatively treating babahoyensis as a separate phyletic line in the Antunesi Com-

plex but it is possible that this species is really related to soperi and the derived

features it shares with the Antunesi line have been developed either independently

or through introgression. Culex babahoyensis occupies the northernmost part of

the range of the complex where it is partially sympatric with antunesi and cer-

queirai of the Antunesi line but has not been found breeding in the same sites.

To date babahoyensis has not been collected in Panama. I found no signifi-

cant differences between the topotypic population from Ecuador and the north-

ern populations in Nicaragua and Costa Rica. In all of these there is an unusually

great variation in the siphon index and pecten teeth of the larva.

BIONOMICS. The immature stages of babahoyensis have been found only in

broken or cut bamboo (3) and in treeholes (1). The only associated species in

these breeding sites were Trichoprosopon (T.) digitatum (4); Toxorhynchites (L.)

sp. (1); and Corethrella sp. (2). According to Levi-Castillo (1953c:163) females

of babahoyensis “‘bite readily during the day and are observed specially at the

sunny spots in the jungle.’’ This is the only record of a species of Carrollia biting

man.

DISTRIBUTION (fig. 5). Known at present only from Caribbean drainages in

Nicaragua and Costa Rica and the Pacific Guayas River basin in Ecuador, at ele-

vations below 200 m.

Material examined: 169 specimens; 15 males, 24 females, 90 larvae, 40 pupae;

38 individual rearings (21 larval, 13 pupal, 4 incomplete).

COSTA RICA. Heredia: Finca La Selva, Puerto Viejo de Sarapiqui (CR 423) [UCLA].

ECUADOR. Guayas: Empalme (ECU 119) [UCLA]. Los Rios: Hacienda Mora, Juan Montalvo

Valencia: Subgenus Carrollia of Culex 73

(ECU 125) [UCLA] ; Levi-Castillo, holotype [USNM] . Valencia (ECU 114-A) [UCLA].

NICARAGUA. Zelaya: Cricket, Rio Escondido (NI 57) [UCLA].

13. Culex (Car.) antunesi Lane & Whitman

Figs. 5,9,34,35

1943. Culex (Carrollia) antunesi Lane and Whitman, 1943:389-392. TYPE: Holotype male,

Sao Joao de Petropolis, Vale do Canaa, Espirito Santo, Brazil, Apr or May 1940, L.

Whitman [BH].

1955. Culex (Carrollia) manaensis Floch and Fauran, 1955:3-5. TYPE: Holotype male with

genitalia slide (909) and associated pupal skin, Boeuf Mort, Haute Mana, French Guiana

[PIG]. NEW SYNONYM.

Culex (Carrollia) antunesi in part of Lane (1953:505-506); Senevet and Abonnenc (1958:287);

Stone, Knight and Starcke (1959:282); Forattini, Rabello and Cotrim (1970:35); Belkin,

Schick and Heinemann (1971:27).

Culex (Carrollia) manaensis of Stone, Knight and Starcke (1959:283); Fauran (1961:44).

Culex (Carrollella) iridescens in part of Komp (1936:64; 1956:39).

Culex (Carrollia) iridescens in part of Busck (1908:70).

Carrollia iridescens in part of Howard, Dyar and Knab (1915:462-464).

Culex (Carrollela) secundus in part of Dyar (1928:281-282).

Culex (Carrollia) secundus in part of Dyar (1925:176).

FEMALE (fig. 9). Wing: 2.5 mm. Proboscis: 1.8 mm. Forefemur: 1.6 mm.

Abdomen: about 2.0 mm. Very similar to kompi and soperi; diagnostic charact-

ers as in the key. Indistinguishable with certainty from babahoyensis and dif-

fering from secundus and soperi in the same features as described for the form-

er. Possibly differing from babahoyensis in the following indefinite variable fea-

tures. Description based on 27 specimens from the Canal Zone and Panama. Head:

Decumbent scales of vertex sometimes somewhat lighter. Proboscis 1.1 of fore-

femur; ventral surface of labium usually somewhat lighter. Palpus 0.19-0.22 of

proboscis. Antenna 0.97-1.00 of proboscis. Thorax: Integument of mesonotum

and pleuron usually dark to moderate brown; lower edge of dark part of pleuron

not as sharply demarcated, lower part of sternopleuron usually infuscated; light

part of pleuron usually somewhat darker. Legs: Coxal integument and scales more

yellowish. Abdomen: Tergites HI,IV often with very narrow dorsal basal dingy

bands. Cibarial Armature: Without apparent significant differences. Genitalia: Ter-

gite [IX with 1 or 2 lateral bristles.

MALE. Wing: 2.4 mm. Proboscis: 2.0 mm. Forefemur: 1.8 mm. Abdomen:

about 2.1 mm. Very similar to babahoyensis, cerqueirai, kompi, soperi and wil-

soni; diagnostic characters in the key not entirely reliable except for soperi. Ap-

parently indistinguishable from cerqueirai except on genitalic characters. Essen-

tially similar to the female and differing from secundus and soperi by the same

general features. Proboscis 1.1 of forefemur; labium more or less uniformly dark

scaled dorsally and indistinctly paler ventrally but without white scaled line. Pal-

pus 0.8 of proboscis; without white scales on segment 3. Antenna 0.95 of pro-

boscis. Specimens studied: 1 from Mangaratiba, Brazil, and 28 from the Canal

Zone and Panama.

MALE GENITALIA (fig. 34). Lobes of tergite IX moderately separated; each

lobe usually with 4 (2-6) bristles. Sidepiece conical; 1 or 2 apicotergal setae dif-

ferentiated; apicosternal process strongly developed. Proximal division of subapi-

74 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

cal lobe slender, almost uniform from near base; seta a short and strongly ex-

panded and contorted distally; seta b longer and usually less expanded and con-

torted distally; seta c developed as a thickened sinuous bristle. Distal division

represented as a short broad protuberance with 2 depressions. Accessory divi-

sion developed as a simple bare, long digitiform process arising sternally near

base of proximal division. Clasper shorter than sidepiece; moderately curved dor-

sad; with moderate differentiation of head but without conspicuous dorsal curv-

ature or inner preapical lobe; crest represented by transverse ridges; snout prom-

inent; seta a more or less spiniform or simple, seta b thickened. Lateral plate

of phallosome with a distinct sternal spine (rarely double). Distal part of proct-

iger elongate and narrowed to apex; paraproct crown usually with 3 (2 or 3)

teeth; cercal setae usually 1 pair (1 or 2).

PUPA (fig. 34). Abdomen: 2.70 mm. Trumpet: 0.55 mm. Paddle: 0.50 mm.

Apparently indistinguishable from babahoyensis and secundus; diagnostic char-

acters from other members of the group as in the key. General chaetotaxy based

on 5 specimens from Panama. Cephalothorax: Pigmentation light yellow; sculp-

turing weak, without definite pattern. All hairs shorter than trumpet, weak; 1,3,

5S-C longer, subequal; 1(2;1-2); 2(2;1-3) longer than hair 6; 3(2,3;2-4); 4(5,6;4-

6) shorter than hair 7; 5(3;2-4); 6(3,4) short; 7(4:4-6); 8(3) shorter than hair

9; 9(2;2-3). Trumpet: Slightly widened distally; index about 11.0 (10.0-12.0);

pinna small. Pigmentation yellow to light brown. Metanotum: Pigmentation yel-

low to light brown; sculpturing weak; hair 10(3,4) slightly longer than others;

11(3,4;2-4); 12(2,3;2-4). Abdomen: Pigmentation light yellow, anterior mesal areas

of tergites II-IV slightly darker; sculpturing weak. Segment I: hair 1(56-84). Seg-

ment II: hair 1(7-9;5-10) slightly longer than hair 5; 3(1) often finely barbed,

nearly reaching caudal margin of tergite III; 4(3;2-3); 5(5). Segment III: hair 1

(3,4); 3(1) often finely barbed, nearly reaching caudal margin of tergite IV; 4

(2,3); 5(4;2-5). Segment IV: hair 1(4,5;3-5); 3(6;6-8); 4(2,3;1-3); 5(1) often barb-

ed, nearly reaching level of rudimentary spiracle VI. Segment V: hair 1 (2,3;2-5);

3(1) about 2.0 of hair 6; 5(1) often barbed, nearly reaching level of rudimentary

spiracle VII. Segment VI: hair 1(1) weak; 3(2;1-2); 4(4;4-5); 5(1) weak, subequal

to hair 6. Segment VII: hair 1(1,2;1-3) subequal to hair 4; 3(1;1-2); 5(2,3) very

short; 6(6;4-6) subequal to hair 5; 9(2,3;2-4) shorter than tergite VIII. Segment

VIII: hair 4(2;1-2); 9(7,8) barbed, slightly longer than paddle.

LARVA (fig. 35). Head: 0.75 mm. Siphon: 1.30 mm. Anal Saddle: 0.35 mm.

Very similar to babahoyensis and cerqueirai; diagnostic characters as in the key.

General chaetotaxy based on 3 reared and 3 other specimens from Panama. Head:

Integument light yellow to brown; collar dark brown. Mental plate with 9 lateral

teeth. Hair 4(8-10); 5(7-10;6-12); 6(3-5;2-5); 7(4,5;4-6); 8(1,2) subequal to hair

7; 9(8,9;6-9); 10(151-2); 11(8,9;6-10); 12(1); 14(2,3); 15(2;1-2). Antenna: About

0.46 of head length; hair 1(3). Thorax: Prothorax: hair 0(12-14;10-14) relatively

long; 1(1); 2(1); 3(3); 4(2); 7(3;2-3); 8(14-18) strongly developed, stellate; 9(1;

1-2); 10(2,3); 11(1); 14(1). Mesothorax: hair 1(4,5:4-6); 2(2;1-2); 3(1); 4(2); 8

(4-6); 9(4-7); 11€1); 13(8-24); 14(10-13) moderately long and stellate. Metatho-

rax: hair 1(4,5;3-6); 2(2); 3(5;4-6); 4(5,6;5-8); 5(1); 6(1); 7(7,8;5-9); 8(18-30);

9(5,6;5-7); 11(1); 13(6-8;6-9) moderately long but weak. Abdomen: Segment I:

hair 1 (4-8) minute; 6(2); 7(2). Segment II: hair 1(1) moderately long; 3(1;1-

3) short and weak; 6(2-3). Segment III: hair 1(1;1-2) strong and long; 3(1;1-2)

long but weak; 6(1); 13(1;1-2) subequal to hair 1. Segment IV: hair 1(1;1-2)

subequal to hair 1-III; 3(2) weak; 6(1); 13(2;1-2) subequal to hair 13-III. Seg-

Valencia: Subgenus Carrollia of Culex 1S

ment V: hair 1(1) slightly longer than hair 1-III,[V; 3(1) long and strong; 6(1);

13(2-3) weaker and much shorter than hair 13-I[V. Segment VI: hair 1(1) short

and weak; 6(1). Segment VII: hair 1(2;1-2) very strong and long. Segment VIII:

Comb scales (4-7) long and with fringes at base only; hair 1(9;9-12); 3(10-12);

5(3-5). Siphon: Pigmentation light brown. Index about 4.1-5.2. Pecten teeth (4-

7) short and without fringes. Ventral siphonal tufts (9-10) usually with 8 or 9

branches (5-9). Anal Segment: Saddle pigmentation light brown. Hair 1(6-8); 2

(3-5). Ventral brush with 4 pairs of hairs with 4-10 branches. Gills slightly long-

er than dorsal saddle length, tapered.

SYSTEMATICS. I have seen only 1 specimen, a male without associated im-

mature stages, from the southern topotypic population of antunesi. The genitalia

of this specimen I cannot distinguish from those of the northern populations in

Costa Rica, Panama and Colombia or from that of the holotype of manaensis from

French Guiana.

The immature stages described above are associated with males of the northern

population, including manaensis (pupa only). They differ conspicuously from the

larva and pupa described and figured by Lane and Whitman (1943:389-392) which

they state were associated with the holotype of antunesi. The extremely long

trumpet of this pupa and the long siphon of this larva are of the type I found

associated with a male from Colombia with genitalia apparently indistinguishable

from those of wilsoni as described and figured by these authors in the same pub-

lication (Lane and Whitman, 1943:393). At the end of the description of an-

tunesi, Lane and Whitman (1943:392) state that immature stages of antunesi,

wilsoni, secundus (= sp. 9a), iridescens and soperi were all collected by L. Whitman

in bamboo internodes in Sao Joao de Petropolis. I believe that an error was made

in associating the skins with the adults and that the larva and pupa Lane and Whit-

man described as antunesi were really those of wilsoni. This interpretation leaves

topotypic antunesi without associated larva and pupa as no subsequent associ-

ations of immature stages have been made for this population.

Therefore, my assignment of the northern populations to antunesi and the

synonymy of manaensis of the Central population is based entirely on the simi-

larity in the male genitalia and is tentative only.

The adults of the Antunesi line, which includes also cerqueirai and wilsoni,

are extremely similar on external features and their separation from the unrelated

babahoyensis is very difficult. Females are known only for antunesi but it is pos-

sible that I have included either or both cerqueirai and wilsoni among the speci-

mens of antunesi from Panama. The males of antunesi and cerqueirai cannot be

separated from each other and their distinction from wilsoni is probably not

reliable. On male genitalic features antunesi is easily distinguished from the other

2 species by the lack of setae on the accessory division of the subapical lobe and

the development of the specialized setae of the proximal division. The pupa is

intermediate between cerqueirai and wilsoni in the length of the trumpet and

easily separated from both on key characters but I have not found any char-

acters to distinguish it from those of the unrelated secundus and babahoyensis.

The larva has a much shorter siphon than wilsoni, a character it shares with cer-

queirai from which it is separated by fewer branches in head hairs 5,7-C; again

the similarity with babahoyensis is very great and its diagnosis from this species

is tenuous.

The known distribution of antunesi, as interpreted here, is widely disjunct.

The scattered northern populations are far removed from the central one in French

76 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

Guiana and the latter from the relatively compact southern ones in the States

of Espirito Santo and Rio de Janeiro in southern Brazil.

As noted above, I have found no differences between the northern populations

which are represented by a fair sample from Panama and the single male and its

associated pupa from the central population. Of the southern topotypic popu-

lation I have seen only 1 male. Because superficially similar forms with practically

identical disjunct distributions in other phyletic lines of the Iridescens Subgroup

are obviously distinct species (see Iridescens Complex), there is a strong possi-

bility that my interpretation of antunesi is incorrect and that 2 or 3 distinct spec-

ies are involved. The problem can be resolved only after adequate material of the

central and southern populations is available for an analysis of differences.

BIONOMICS. The immature stages of both northern and southern populations

of antunesi have been found only in cut or broken bamboo (5). In the north the

known associates are 1 species of Carrollia (secundus), Limatus durhamii (1),

Sabethes sp. (1), Toxorhynchites (L.) sp. (1), Trichoprosopon (T.) digitatum and

Wyeomyia sp. (1). The southern populations of this species may be associated

with sp. 9a (reported as secundus), iridescens, soperi and wilsoni (Lane and Whit-

man, 1943:392). |

DISTRIBUTION (fig. 5). Widely disjunct; typical southern populations in coastal

drainages in States of Espirito Santo and Rio de Janeiro, Brazil; northern popu-

lations in Costa Rica, Panama, upper Orinoco drainage in Colombia and French

Guiana, at elevations of 50-500 m.

Material examined: 80 specimens; 34 males, 26 females, 10 larvae, 9 pupae;

8 individual rearings (5 larval, 3 pupal).

BRAZIL. Rio de Janeiro: Mangaratiba [BH].

COLOMBIA. Meta: Villavicencio (C-19,102) [UCLA].

COSTA RICA. Puntarenas: Villa Neily (CR 167) [UCLA].

FRENCH GUIANA. Jnini: Boeuf Mort, Haute Mana, E. Abonnenc [PIG].

PANAMA. Canal Zone: Empire, A.H. Jennings [USNM]. Tabernilla, AH. Jennings [USNM] ;

A. Busck [USNM]. Colon: Portobelo (P82-3) [USNM]. Darien: Paya Camp (GG 1-160,105,

107) [UCLA]. Pucro (GG 109) [UCLA].

Additional Record From the Literature

BRAZIL. Espirito Santo: Sao Joao de Petropolis [FH, 4095] (Forattini, Rabello and Cotrim,

1970:35).

14. Culex (Car.) cerqueirai Valencia, n.sp.

Figs. 5,36,37

TYPE: Holotype male (PA 654-104) with slides of genitalia and associated pupal skin, Quebrada

Grande, Bocas del Toro, Panama, 11 Apr 1964 [USNM].

FEMALE. Unknown. Probably very similar to antunesi and babahoyensis; pos-

sibly included in some of the material identified in this study as antunesi.

MALE. Wing: 2.4 mm. Proboscis: 1.9 mm. Forefemur: 1.5 mm. Abdomen:

about 2.2 mm. Very similar to babahoyensis, kompi, soperi and wilsoni; diag-

nostic characters in the key not entirely reliable except for soperi. Apparently

indistinguishable from antunesi except on genitalic characters. General features

as described for the females of babahoyensis and antunesi and differing from

secundus and soperi in similar characters. Description and measurements based

Valencia: Subgenus Carrollia of Culex ay

on pinned holotype. Head: Broad decumbent scales of vertex largely pale ex-

cept for a narrow line of dark scales caudad of orbital bristles and on midline

in front; narrow decumbent scales apparently absent; erect scales as in babahoy-

ensis; lateral patch yellowish white. Proboscis about 1.3 of forefemur; labium

more or less uniformly dark scaled dorsally and with a narrow indefinite light

scaled line from before false joint to near apex. Palpus about 0.8 of proboscis;

without white scales on segment 3. Antenna about 0.8 of proboscis. Thorax:

Integument of mesonotum and pleuron dark brown. Mesonotal and scutellar scal-

ing essentially as in babahoyensis except for shorter lighter broad scales on lateral

lobe of scutellum. Lower edge of dark part of pleuron not sharply marked off

on stp, lower part of stp strongly infuscated; meron not darkened except light-

ly on extreme base. Legs: Coxal integument and scaling yellowish white; fore-

coxal scale patch without dark scales. Femoral scaling essentially as in babahoy-

ensis but light scaling slightly less extensive. Abdomen: Tergites III-VII with dis-

tinct dorsal basal light bands. Metallic light spots essentially as in babahoyensis.

Sternites III-VII with distinct apical dark bands.

MALE GENITALIA (fig. 36). In general as described for antunesi and differ-

ing only in the following few conspicuous features. Setae a and b of proximal

division of subapical lobe subequal, more or less crescentic and with sharp point.

Accessory division removed cephalodorsad from base of proximal division and

with about 12 recurved bristles in distal half. Seta a of clasper simple.

PUPA (fig. 36). Abdomen: 2.70 mm. Trumpet: 0.40 mm. Paddle: 0.50 mm.

Apparently differing from all other members of the subgroup by the flared trum-

pet with large pinna. General chaetotaxy based on 1 specimen from Panama.

Cephalothorax: Pigmentation light yellow; sculpturing very weak; without defi-

nite pattern. All hairs shorter than trumpet, 1,4,6,7,9-C subequal; 1(2); 2(2) very

short; 3(3,4) shorter than hair 5; 4(1); 5(3) weak; 6(1); 7(2); 8(1) stouter than

hair 9; 9(2). Trumpet: Strongly widened distally; index about 10.0; pinna large,

about 0.25. Pigmentation light brown. Metanotum: Pigmentation light yellow;

sculpturing weak; hair 10(1) longer than others; 11(2); 12(2). Abdomen: Pigmens

tation light yellow throughout; sculpturing weak. Segment I: hair 1(35-45) weak.

Segment II: hair 1(2) longer than hair 5; 3(1) surpassing level of rudimentary

spiracle III; 4(2); 5(1). Segment III: hair 1(3,4); 3(1) surpassing level of rudi-

mentary spiracle IV; 4(1,2), 5(2). Segment IV: hair 1(2); 3(2); 4(1); 5(1) reach-

ing caudal margin of tergite V. Segment V: hair 1(2); 3(1) slightly longer than

hair 6; 4(2); 5(1) reaching level of alveolus of hair 6-VI. Segment VI: hair 1(1)

weak but longer than hair 6; 3(2) subequal to hair 5; 4(3,4); 5(1). Segment VII:

hair 1(1) longer than hair 9; 3(1); 4(2); 5(2) short; 6(4) subequal to hair 5; 9

(2) weak, subequal to hair 4. Segment VIII: hair 4(2) about 0.5 of tergite length;

9(missing).

LARVA (fig. 37). Head: 0.90 mm. Siphon: 1.15 mm. Anal Saddle: 0.35 mm.

Very similar to antunesi and babahoyensis; diagnostic characters as in the key.

Association with male and pupa presumptive only; general chaetotaxy based on

5S whole larvae from Panama. Head: Integument light yellow; collar dark brown.

Mental plate with 9 or 10 lateral teeth. Hair 4(10;9-12); 5(12-16;10-16) with-

out barbs; 6(4,5;3-6); 7(10-12;8-12) without barbs; 8(2) relatively short; 9(10-

12;9-14); 10(2;1-2); 11(8-10;4-10); 12(1); 14(3;2-3); 15(2-3). Antenna: About 0.44

of head length; hair 1(2,3;2-4). Thorax: Prothorax: hair 0(12-16) short; 1(1); 2

(1); 3(4;2-4); 4(2); 7(4;4-5); 8(10-14) moderately long but strongly barbed; 9

(1); 10(2); 11(1); 14(1). Mesothorax: hair 1(3;3-6); 2(1-2); 3(1); 4(2); 8(5,6;4-

78 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

6); 9(5-8); 11¢€1); 13(16-20); 14(11-20) moderate, stellate, Metathorax: hair |

(3;2-3); 2(1-2) subequal to hair 6; 3(3,4;3-6); 4(3;2-3); 5(1); 6(1); 7(6;6-7); 8

(19-24); 9(4-6); 11(€1); 13(S5-12) moderately long but weak. Abdomen: Segment

I: hair 1(6-8) minute; 6(2); 7(2-3). Segment II: hair 1(1) moderately long and

strong; 3(1) shorter and weaker than hair 1; 6(2). Segment III: hair 1(1;1-2)

strong and long; 3(1) moderately long but weak; 6(1;1-2); 13(1-3) subequal to

hair 1. Segment IV: hair 1(1;1-2) subequal to hair 1-III; 3(1) subequal to hair

3-III or weaker; 6(1;1-2); 13(1;1-3) subequal to hair 13-III. Segment V: hair 1

(1) slightly longer and stronger than hair 1-IV; 3(1) moderately long but rather

weak; 6(1); 13(1-4) shorter than hair 13-IV. Segment VI: hair 1(1) weak and

relatively short; 6(1). Segment VII: hair 1(2) strong. Segment VIII: Comb scales

(5-12) sharply pointed and without fringes; hair 1(4-9); 3(9-11); 5(3-4) without

barbs. Siphon: Pigmentation light brown. Index about 3.6-4.4. Pecten teeth (4-

7) sharply pointed, moderately long and with ventral marginal spicules at base

only. Ventral siphonal tufts (7-8) with about 8 branches (7-9). Anal Segment:

Saddle pigmentation light brown. Hair 1(5-7); 2(3-5). Ventral brush with 4 pairs

of hairs, all usually with about 5 branches (3-6). Gills pshelatiss longer than dorsal

saddle length, tapered.

SYSTEMATICS. Culex cerqueirai is apparently a relict species in the Antunesi

line which includes wilsoni in addition to the nominate species. It has a peculiar

combination of features of antunesi and wilsoni. The single known adult, the

holotype male, cannot be separated from antunesi on external features. The male

genitalia are very similar to wilsoni but differ from it in the small number and

restricted distribution of setae on the more slender accessory division of the sub-

apical lobe and in the shorter, broader and crescentic setae a and b of the prox-

imal division. The associated pupa is differentiated from the other members of

the line by a relatively short, strongly flared trumpet but is more like antunesi

than wilsoni.

The association of the larva with the holotype is presumptive only and com-

plicated by an apparent error in mixing material from 2 collections in the field,

made in the same locality, PA 654 from broken or cut bamboo and PA 657 from

flower bracts of Heliconia. The holotype was originally labelled 657-101 and the

larvae 654. I am confident that both came from collection 654 because there were

similar errors made with other species in these 2 collections, typical Heliconia breed-

ers being included in the collection from bamboo and vice versa. Therefore, I

have relabelled the holotype as 645-104. The larva I am tentatively associating

with the holotype of cerqueirai is very similar to those of antunesi and babahoy-

ensis but is distinguished from both by the more profuse branching of head hairs

5-C and 7-C.

Culex cerqueirai is known from a single locality in the Bocas del Toro Pro-

vince of Panama within the range of the northern populations of antunesi but

not associated with the latter.

BIONOMICS. As noted in the section on systematics, the presumed larva of

cerqueirai is not definitely associated with the pupa from which emerged the male

holotype. I am confident however they all came from the same collection in cut

or broken bamboo where they were associated with a species of Carrollia (secundus),

Sabethes undosus, Wyeomyia aporonoma and W. arthrostigma. No adults other

than the single male are known.

DISTRIBUTION (fig. 5). Known only from the type locality, at an elevation of

10-20 m.

Valencia: Subgenus Carrolia of Culex 79

Material examined: 16 specimens; 1 male, 1 pupa, 14 larvae; 1 individual pupal

rearing.

PANAMA. Bocas del Toro: Quebrada Grande (PA 654) [UCLA].

15. Culex (Car.) wilsoni Lane & Whitman

Figs. 5,38,39

1943. Culex (Carrollia) wilsoni Lane and Whitman, 1943:393. TYPE: Holotype male, Sao

Joao de Petropolis, Vale do Canaa, Espirito Santo, Brazil, Apr 1940, L. Whitman

[LU].

Culex (Carrollia) wilsoni of Lane (1953:506); Stone, Knight and Starcke (1959:283); Forattini,

Rabello and Cotrim (1970:50); Belkin, Schick and Heinemann (1971:28).

Culex (Carrollia) antunesi in part of Lane and Whitman (1943:389-392).

Culex (Carrollella) iridescens in part of Komp (1936:64; 1956:39).

FEMALE. Unknown. Probably very similar to antunesi and babahoyensis,; pos-

sibly included in some of the material identified in this study as antunesi.

MALE. Wing: 2.8 mm. Proboscis: 2.0 mm. Forefemur: 1.8 mm. Abdomen:

about 1.9 mm. Very similar to antunesi, babahoyensis, cerqueirai, kompi and

soperi; diagnostic characters in the key probably not entirely reliable except for

soperi. General features as described for the females of babahoyensis and antunesi

and differing from secundus and soperi in similar characters. Description and meas-

urements based on 2 dry specimens from Colombia. Head: Broad decumbent scales

of vertex either all whitish or whitish and yellow with a few dark scales on mid-

line in front; a few yellowish narrow decumbent scales; erect scales as in baba-

hoyensis; lateral patch whitish. Proboscis about 1.1 of forefemur; labium more

or less uniformly dark scaled dorsally and with a conspicuous ventral line of

whitish scales. Palpus about 0.8 of proboscis; without white scales on segment

3. Antenna about 0.75 of proboscis. Thorax: Integument of mesonotum and

pleuron light brown. Mesonotal scaling denser than in babahoyensis, paler in pre-

scutal space. Scutellum with shorter, paler scales on midlobe and lateral lobe.

Lower edge of dark part of pleuron not sharply marked off on stp; lower part

of stp slightly infuscated; meron not darkened. More numerous, conspicuous and

longer broad scales near lower posterior stp bristles. Legs: Coxal integument and

scaling yellowish white; forecoxal scale patch without dark scales. Femoral scal-

ing essentially as in babahoyensis. Abdomen: Broken off; apparently with larger

light metallic markings on proximal segments than in babahoyensis,; basal tergal

light bands present at least on IV-VII; sternites with apical dark bands at least

on IV-VII.

MALE GENITALIA (fig. 38). In general as described for antunesi; very sim-

ilar to cerqueirai and differing from it primarily in the following features. Setae

a and b of proximal division of subapical lobe more nearly equal, much narrower

and not crescentic. Accessory division broader, less digitiform, somewhat expand-

ed distally and usually with about 30 setae extending to at least basal third.

PUPA (fig. 38). Abdomen: 2.80 mm. Trumpet: 0.75 mm. Paddle: 0.45 mm.

Differing from all other known species of subgenus by the very long and narrow

trumpet. General chaetotaxy based on 3 specimens from Colombia. Cephalotho-

rax: Pigmentation light yellow; sculpturing weak. All hairs shorter than trumpet,

1,5-7-C subequal, 3-C the longest; 1(2); 2(2;1-2) shorter than hair 6; 3(1-4); 4

(1-4); 5(3-5); 6(2;1-2); 7(2-4); 8(2-4) shorter than hair 9; 9(1-2). Trumpet: Near-

80 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

ly uniform in width; index about 17.0 (15.0-18.7); pinna very small. Pigmenta-

tion light brown. Metanotum: Pigmentation light yellow; sculpturing weak; hair

10(2-3) longer than the others; 11(1-5); 12(1-3). Abdomen: Pigmentation more

or less uniformly yellow; sculpturing weak. Segment I: hair 1(80-120). Segment

Il: hair 1(6-10) subequal to hair 5; 3(1) barbed, not reaching caudal margin of

tergite III; 4(2-6); 5(1-4). Segment III: hair 1(4-5); 31) barbed, surpassing cau-

dal margin of tergite IV; 4(1-3); 5(2-5). Segment IV: hair 1(2-4); 3(3-8); 4(1);

5(1) extremely long, surpassing level of rudimentary spiracle VII. Segment V:

hair 1(2-5) reaching level of rudimentary spiracle VI; 3(1-2) shorter than hair

1; 4(5-6); 5(1) very long, reaching tergite VIII. Segment VI: hair 11-2); 3(1-2)

subequal to hair 1; 4(2-5); 5(1) shorter than hair 6. Segment VII: hair 1(1-2)

reaching alveolus of hair 4-VIII; 3(1-2); 4(1-2); 5(2-4) subequal to hair 6; 6(2-

8); 9(3-5) barbed, less than 0.5 of hair 9-VIII. Segment VIII: hair 4(1-2) weak;

9(5-9) barbed, distinctly longer than paddle.

LARVA (fig. 39). Head: 0.85 mm. Siphon: 2.10 mm. Anal Saddle: 0.36 mm.

In general similar to antunesi, babahoyensis and cerqueirai, but readily differ-

entiated by the very long siphon. General chaetotaxy based on | associated skin,

1 skin and 2 whole larvae from Colombia. Head: Integument light yellow to

brown; collar dark brown. Mental plate with 8 or 9 lateral teeth. Hair 4(8,9;

8-11); 5(8-16); 6(2,3;2-4); 7(5,6;5-8) relatively short and without barbs; 8(1) long-

er than hair 7; 9(12;10-12); 10(1); 11(6-8;5-8); 12(1); 14(3,4); 15(@2,3). Antenna:

About 0.44 of head length; hair 1(2,3;1-3). Thorax: Prothorax: hair 0(10-18)

short; 1(1); 2(1); 3(2-6); 4(2); 7(3,4); 8(6-12) strong and barbed; 9(1); 10(2);

11(1,2); 12(1;1-2); 14(1). Mesothorax: hair 1(3-8); 2(1,2); 3(1); 4(1); 8(4;4-8);

9(4-6); 11(1); 13(14-20); 14(8-12) moderately long and stellate. Metathorax: hair

1(3,4;3-5); 2(1,2); 3(4-6); 4(3-7); 5(1,2;1-3); 6(1); 7(4-8;4-9); 8(12-20); 9(2-5);

11(1); 13(5,6) very long but with weak branches. Abdomen: Segment I: hair 1

(6-10) minute; 6(2); 7(2). Segment II: hair 1(1;1-2) long and rather strong; 3(1;

1-2) short; 6(2). Segment III: hair 1(1) very long and strong; 3(1) long but thin;

6(1); 13(1,2) subequal to hair 1. Segment IV: hair 1(1;1-2) subequal to hair 1-

III but stronger; 3(1;1-2f) moderate, shorter than hair 3-III; 6(1); 13(1) subequal

to hair 13-III]. Segment V: hair 1(1) subequal to hair 1-IV; 3(1) subequal to hair 3-

III; 6(1); 13(1-3) subequal to hair 13-III,[1V. Segment VI: hair 1(1) weak, moderate;

6(1). Segment VII: hair 1(1) very strong and long. Segment VIII: Comb scales (10-

10) very narrow and without fringes; hair 1(5-10); 3(7-12); 5(3;2-4) without barbs.

Siphon: Pigmentation light brown. Index about 9.7-12.0. Pecten teeth (10-16) mod-

erately long, sharp and without fringes. Ventral siphonal tufts (7-8) usually with

about 4 or 5 uneven branches (4-7). Anal Segment: Saddle pigmentation light

brown. Hair 1(2-8); 2(3,4). Ventral brush with 4 pairs of hairs with variable number

of branches (4-8). Gills very narrow from base, about 3.0 of dorsal saddle length.

SYSTEMATICS. My assignment of the population of the Antunesi line from the

Villavicencio area in Colombia, described above, to wilsoni is tentative as it is

based only on similarity in male genitalia. Even this is tenuous as I had no access

to the type material of wilsoni, whole holotype is apparently lost, and had to

rely entirely on the original description and figures.

On external features the males of this population agree with the description

of topotypic wilsoni in the whitish scaling of the ventral surface of the labium,

a character distinguishing them from the other 2 members of the line, antunesi

and cerqueirai. Females are not known for either population but may be included

among the specimens from Panama I have identified as antunesi. The male geni-

Valencia: Subgenus Carrollia of Culex 81

talia of the 2 populations I cannot distinguish. They are very similar to those

of cerqueirai but are readily differentiated from the latter by the characters given

in the diagnosis above.

Associated by individual rearings with the male from the Villavicencio popu-

lation are a pupa and larva which show general similarity with the immature

stages ascribed to antunesi by Lane and Whitman (1943:390-392). As indicated

under antunesi, I believe that the latter association was in error and these im-

mature stages are actually those of topotypic wilsoni. The pupae of both popu-

lations differ from those of the other species of the line as well as all species

of Carrollia in the very long trumpet and the larvae are unique in the very long

siphon.

The only 2 known populations of wilsoni are extremely widely disjunct. The

topotypic population has been collected only once near Sao Joao de Petropolis

in the State of Espirito Santo in southern Brazil within the range of topotypic

antunesi. The northern population is known from 4 collections in 3 localities

in the Villavicencio area of Colombia, within the range of the northern popula-

tions of antunesi.

The conspecificity of these 2 populations is by no means conclusively estab-

lished. There appear to be some differences in the larvae and pupae in the 2

populations and an examination of the male genitalia of the topotypic popula-

tion may reveal details not apparent in the figures and description which may

distinguish this population. Because in the Iridescens Complex and in the Soperi-

Babahoyensis lines superficially similar forms with widely disjunct northern and

southern distributions are clearly distinct species, there is a strong probability

that this is also the case with the 2 populations of wilsoni. However, without

adequate material of either population, I refrain from separating the northern

population as a distinct species at this time.

BIONOMICS. The adults of the northern population of wilsoni were all reared

from immature stages collected in cut or broken bamboo (3) without recorded

associates. Lane and Whitman (1943:392) state that the 3 males of the type

series, the only specimens recorded from the southern population, were also reared

from a collection in bamboo where they may have been associated with 1 or

more of the following species of Carrollia: antunesi, sp. 9a (reported as secundus),

soperi and iridescens.

DISTRIBUTION (fig. 5). Widely disjunct; typical southern population in State

of Espirito Santo, Brazil; northern population in Colombia in the vicinity of Villa-

vicencio in upper part of Orinoco basin; at elevations of 400-500 m.

Material examined: 12 specimens; 3 males, 6 larvae, 3 pupae; 2 individual rear-

ings (1 larval, 1 incomplete).

COLOMBIA. Cundinamarca: La Union (KO 10-43) [UCLA]. Meta: Bosque de Ocoa (1006-2,

1022-2) [UCLA]. Villavicencio (65) [UCLA].

Additional Record From the Literature

BRAZIL. Espirito Santo: Sao Joao de Petropolis [FH, 4096] (Forattini, Rabello and Cotrim,

1970:50).

16. Culex (Car.) kompi Valencia, n.sp.

Figs. 5,40,41

TYPES: Holotype male (8) with genitalia slide (720111-1) and associated larval skin, Salinas,

82 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

Restrepo, Meta, Colombia, 22 Aug 1935, W.H. Komp [USNM]. Allotype female (KO 10-42-3)

with associated larval skin, same data as holotype [USNM]. Paratypes: 2 IM (KO 10-30-1,2),

1M (KO 1042), 1F (KO 10-30), 2 IF (7,9), 3 1(4,5,6), same data as holotype [UCLA, USNM].

FEMALE. Wing: 2.4 mm. Proboscis: 1.8 mm. Forefemur: 1.7 mm. Abdomen:

about 2.2 mm. Differentiated from all other members of the subgroup by the

very long palpus and 2 short setae below the interorbital bristles. Very similar

to antunesi, babahoyensis and soperi; differing from secundus in the same gen-

eral features. Description and measurements based on 2 pinned specimens from

Colombia. Head: Broad decumbent scales of vertex largely pale yellowish, only

a narrow band of darker scales along orbital bristles; a few light yellowish nar-

row decumbent scales present; erect scales light yellowish, as in babahoyensis;

lateral patch light yellowish. Two short setae below interorbital bristles. Probos-

cis about 1.1 of forefemur; labium uniformly brown. Palpus 0.45-0.50 of pro-

boscis. Antenna 0.94 of proboscis. Thorax: Integument of mesonotum and pleu-

ron dark brown; lower edge of dark part of pleuron not sharply demarcated on

sternopleuron; latter infuscated; meron not infuscated. Mesonotal vestiture dens-

er and paler posteriorly than in babahoyensis, more similar to that of secundus.

Prescutellar bristles weaker and less numerous. Scutellar lobes with shorter broad

scales, all pale. More numerous, conspicuous and longer broad scales near lower

posterior stp bristles; a few broad scales near umep bristles sometimes present.

Legs: Coxal integument and scales more yellowish than in babahoyensis. Femora

essentially as in babahoyensis. Abdomen: Dorsal basal tergal light bands whitish,

distinct on II,IV but smaller than on V-VII. Metallic light spots essentially as

in babahoyensis. Sternites as in babahoyensis, more silvery white rather than yel-

lowish. Cibarial Armature and Genitalia: Not studied.

MALE. Wing: 2.4 mm. Proboscis: 1.8 mm. Forefemur: 1.7-1.9 mm. Abdo-

men: about 2.0 mm. Very similar to antunesi, babahoyensis, cerqueirai, soperi

and wilsoni; diagnostic characters in the key not entirely reliable except for sop-

eri. Essentially similar to the female and differing from secundus and soperi by

the same general features. Proboscis about 1.2 of forefemur; labium more or less

uniformly dark scaled dorsally, ventral surface pale to whitish distad of false

joint. Palpus 0.72-0.78 of proboscis; without white scales on segment 3. Anten-

na 0.82 of proboscis. Specimens studied: 4 from Colombia. Measurements from

dry specimens.

MALE GENITALIA (fig. 40). In general as described for antunesi but differ-

ing very conspicuously from it and other species of the subgroup in the follow-

ing features. Lobes of tergite [IX very large and closely approximated; each lobe

usually with 5 (4 or 5) bristles. Sidepiece ovoid; apicotergal seta not differenti-

ated. Proximal division with narrow strongly curved stem; setae a and b flatten-

ed and expanded, a shorter and with recurved point, b with forked apex; seta

c a simple bristle. Accessory division composed of 2 widely separated, long, slen-

der columnar processes; ventral process arising from base of proximal division

and with 2 simple subapical bristles, 5 apical flattened modified setae, 3 nar-

row and pointed and 2 moderately dilated; dorsal process widely separated at

base from ventral, stem with sigmoid curvature, apex with 2 or 3 long, curved

differentiated bristles. Clasper strongly curved dorsad apically; setae a and b both

thickened, spiniform; snout very prominent. Paraproct crown with 2 very short

blunt teeth; cercal setae usually 1 pair (1 or 2), very short.

PUPA (fig. 40). Abdomen: 1.55 mm. Trumpet: 0.40 mm. Paddle: 0.35 mm.

Unique in the subgenus in the development of the trumpet and abdominal hair

Valencia: Subgenus Carrollia of Culex 83

9-VIII. General chaetotaxy based on 4 unassociated skins from Colombia. Cephalo-

thorax: Pigmentation light yellow, sculpturing very weak. All hairs shorter than

trumpet except 1,4,9-C which are about the same length; 1(1) longer than hair 7;

2(2:1-2) longer than hair 6; 3(2) slightly longer than hair 4; 4(1,2) longer than

hair 7; 5(3,4;3-5); 6(1); 7(1,2;1-3); 8(3;2-3) shorter than hair 9; 9(1). Trumpet:

Fusiform; index about 9.0 (8.6-10.0); pinna minute. Pigmentation light brown.

Metanotum: Pigmentation light yellow, sculpturing very weak; hair 10(2;1-2) sub-

equal to hairs 11,12; 11(2); 12(1,2;1-3). Abdomen: Pigmentation light yellow;

sculpturing very weak. Segment I: hair 1(120-190). Segment II: hair 1(10-16) sur-

passing level of rudimentary spiracle III; 3(1); 4(4,5;4-6); 5(2;1-2) shorter than hair

3. Segment III: hair 1(1;1-2) surpassing level of rudimentary spiracle V; 3(1); 4(1);

5(1;1-3) short. Segment IV: hair 1(1) longer than hair 3; 3(4,6;4-8) longer than hair

6; 4(1); 5(1;1-2) nearly reaching caudal margin of tergite V. Segment V: hair 1(1)

subequal to hair 4; 3(1) about 3.0 of hair 1; 4(3,4;3-6); 5(1) nearly reaching rudi-

mentary spiracle VI. Segment VI: hair 1(1) subequal to hair 4; 3(1); 4(1); 5(1). Seg-

ment VII: hair 1(1) shorter than hair 4; 3(1;1-2) subequal to hair 6; 4(1); 5(1); 6(1);

9(1) weak, shorter than hair 9-VIII. Segment VIII: hair 4(1); 9(2) extremely long,

about 0.5 of abdomen, without barbs.

LARVA (fig. 41). Head: 0.85 mm. Siphon: 0.95 mm. Anal Saddle: 0.35 mm.

Unique in the subgenus in the development of abdominal hair 7-II. General chaeto-

taxy based on 6 associated and 3 other skins from Colombia. Head: Integument

light yellow; collar dark brown. Mental plate with 8-10 lateral teeth. Hair 4(8-10);

5(10-12:8-17); 6(2;2-3); 7(7,8;6-8) not barbed; 8(2;1-2) much shorter than hair 7;

9(8-10;7-12); 10(1); 11(4;2-5); 12(1); 14(3;3-4); 15(2;1-3). Antenna: About 0.41

of head length; hair 1(2-4;1-4). Thorax: Prothorax: hair 0(8-14) minute; 1(1); 2(1)

relatively short; 3(2;2-3) relatively very short; 4(2); 6(1;1-2); 7(2-4;2-5); 8(6-12)

short but on distinct tubercle; 9(1;1-2); 10(1;1-2); 11(1;1-3); 12(1;1-2); 14(1). Meso-

thorax: hair 1(3;1-3); 2(2;1-3); 3(1,2;1-3); 4(3;2-3); 7(1;1-2); 8(2;2-6); 9(2-4;2-5);

11(1); 13(8-20); 14(6-16) short and weak. Metathorax: hair 1(2-4); 2(2); 3(3); 4Q-

4); 7(3-5;2-5); 8(10-18); 9(2,3;2-6); 11(1); 13(2,3;2-4) very long and strong. Abdo-

men: Segment I: hair 1(3-4) minute; 6(2;2-4); 7(2). Segment II: hair 1(2;2-3) long

and strong, subequal to hair 1-III; 3(1) short; 6(2;2-3); 7(2;2-3) very strong, sub-

equal to hair 7-I. Segment III: hair 1(2;2-3); 3(1;1-2) moderate; 6(2); 13(3;2-3) sub-

equal to hair 1. Segment IV: hair 1(2,3) subequal to hair 1-II,V; 3(1,1-2) shorter

than hair 3-III; 6(2); 13(2,3) subequal to hair 1. Segment V: hair 1(2,3); 3(1) mod-

erate; 6(2); 13(3) subequal to hair 1. Segment VI: hair 1(1) weak and short; 6(2).

Segment VII: hair 1(3) relatively short. Segment VIII: Comb scales (4-7) mod-

erately long, sharp and without fringes; hair 1(3-8); 3(6-8;5-8); 5(2,3). Siphon:

Pigmentation light yellow. Index about 2.9-3.3. Pecten teeth (6-11) moderate,

sharp and without fringes. Ventral siphonal tufts (8-9) mostly with 5 branches

(4-8). Anal Segment: Saddle pigmentation light yellow. Hair 1(4,5;3-6); 2(3;3-

4). Ventral brush with 4 pairs of hairs, all usually with 3 branches (2-4). Gills

less than 2.0 of dorsal saddle length, tapered.

SYSTEMATICS. Culex kompi is a relict species with a bizarre combination

of a few apparently primitive and a multitude of unique derived character states

and cannot be confused with any other species of Carrollia in any stage. How-

ever it appears to be related to the other phyletic lines in the Antunesi Complex

in general features of the adults, male genitalia and larva. The pupa is so differ-

ent from other species of Carrollia that superficially it does not appear to belong

to the subgenus and my tentative association of this stage with the adult and

larva may be incorrect.

54 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

Culex kompi resembles soperi in a derived feature (ovoid sidepiece in the male

genitalia) which is characteristic of all members of the Iridescens Complex. This

may indicate phyletic relationship, introgression with a sympatric member of the

Iridescens Complex or completely independent origin. The apparently primitive

character of branched hair 6-III-V of the larva shared by these 2 species may

have also arisen as a result of such introgression. All these alternatives lead to the

conclusion that kompi probably originated early in the evolution of the Irides-

cens Subgroup at the time of separation of the Iridescens and Antunesi Complexes.

Culex kompi could be segregated as a separate complex but. because of the simi-

larity in general features with other lines of the Antunesi Complex I am retaining

it in this complex. I believe that the above-mentioned annectent features of kompi

as well as its unique derived features (female palpus; larval hair 7-II; pupal chaeto-

taxy and trumpet) may have arisen as a result of introgression.

BIONOMICS. All the adults of kKompi were reared from collections of immature

stages in broken or cut bamboo (3) without record of associated species.

DISTRIBUTION (fig. 5). Known only from the vicinity of Villacicencio, Co-

lombia, in upper part of Orinoco basin, at elevations of 450-500 m.

Material examined: 22 specimens; 4 males, 4 females, 9 larvae, 5 pupae; 6 indi-

vidual rearings(larvae and adults only).

COLOMBIA. Meta: Restrepo, Salinas (KO 10-30,10-42) [UCLA, USNM]. Villavicencio, M.

Bates [USNM].

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Soc. Wash., Proc. 33:1-27.

1931b. On the classification of Brazilian Culicidae with special reference to those

capable of harboring the yellow fever virus. Entomol. Soc. Wash., Proc. 33:

125-164.

Valencia: Subgenus Carrollia of Culex 89

Stone, A. and K.L. Knight

1957. Type specimens of mosquitoes in the United States National Museum: IV,

The genus Culex (Diptera, Culicidae). Wash. Acad. Sci., J. 47:42-59.

Stone, Alan, K.L. Knight and H. Starcke

1959. A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae).

Wash., Entomol. Soc. Am. (Thomas Say Found., vol. 6). 358 p.

Surcouf, Jacques M.R. and R. Gonzalez-Rincones

1911. Essai sur les dipteres vulnerants du Venezuela. Premiere partie. Dipteres

nematoceres vulnerants. Paris, Maloine. 320 p.

Theobald, Frederick V.

1905. Diptera. Fam. Culicidae. Genera Insectorum 26. 50 p.

1907. A monograph of the Culicidae or mosquitoes. Vol. 4. Lond., Br. Mus.

(Nat. Hist.). 639 p.

1910. A monograph of the Culicidae or mosquitoes. Vol. 5. Lond., Br. Mus.

(Nat. Hist.). 646 p.

Townsend, Charles H.T.

1934. Mosquitoes of the Rio Tapajos. Rev. Entomol. 4:486-499.

Urich, F.W.

1913. Mosquitoes of Trinidad. Agric. Soc. Trinidad Tobago, Proc. 13:525-530.

Vargas, Luis

1956. Especies y distribucion de mosquitos mexicanos no anofelinos (Insecta,

Diptera). Inst. Salub. Enferm. Trop., Rev. 16:19-36.

DAARWN >

Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

FIGURES

Culex (Carrollia) secundus; female

Distribution of the Bihaicolus Group

Distribution of the Urichii Subgroup

Distribution of the Iridescens Complex

Distribution of the Antunesi Complex

Culex (Car.) bihaicolus; male and female heads and claws; female thorax, wing,

legs and genitalia

Culex (Car.) urichii; male and female heads and claws; female thorax, wing,

legs and genitalia

Culex (Car.) secundus; male and female heads and claws; female thorax, wing,

legs and genitalia

Culex (Car.); female cibarial armatures

Culex (Car.) bihaicolus; male genitalia and pupa

Culex (Car.) bihaicolus; larva

Culex (Car.) guerreroi; male genitalia and pupa

Culex (Car.) guerreroi; larva

Culex (Car.) rausseoi; male genitalia and pupa

Culex (Car.) rausseoi; larva

Culex (Car.) metempsytus; male genitalia and pupa

Culex (Car.) metempsy tus, larva

Culex (Car.) infoliatus; male genitalia and pupa

Culex (Car.) infoliatus; larva

Culex (Car.) urichii; male genitalia and pupa

Culex (Car.) urichii; larva

Culex (Car.) anduzei; male genitalia and pupa

Culex (Car.) anduzei, larva

Culex (Car.) bonnei; male genitalia and pupa

Culex (Car.) bonnei; larva

Culex (Car.) secundus,; male genitalia and pupa

Culex (Car.) secundus; larva

Culex (Car.) iridescens; male genitalia and pupa

Culex (Car.) iridescens; larva

Culex (Car.) soperi; male genitalia and pupa

Culex (Car.) soperi; larva

Culex (Car.) babahoyensis; male genitalia and pupa

Culex (Car.) babahoyensis; larva

Culex (Car.) antunesi; male genitalia and pupa

Culex (Car.) antunesi; larva

Culex (Car.) cerqueirai; male genitalia and pupa

Culex (Car.) cerqueirai; larva

Culex (Car.) wilsoni; male genitalia and pupa

Culex (Car.) wilsoni; larva

Culex (Car.) kompi; male genitalia and pupa

Culex (Car.) kompi; larva

secundus

CR 412

Heredia

Costa Rica

SATIW ALALVIS

00s

001

ont 00s

SYaLINOTTA

VOIMANV WIAAIN

l0Jojion3 loassne

_s - ayes i

SNIBIOJUT nce =

JO} ee

aie

dnois) snjoorey

hel ANA he F

G@NITIOUNVO ~

ont

CARROLLIA

~*~-Urichii Subgroup

» urichii

Dee

LSS

ep ATR ey =

Tropic of Capricorn

NORTH & SOUTH AMERICA

KILOMETERS

rere i 2000 2800 3000

ai 1500 “2000

1000

STATUTE MILES

CARROLLIA

Sy. 2 ete x

i¢~ ¥ Iridescens Complex

; Sy eae

Vim;

o ,bonnei

NORTH & SOUTH AMERICA

KILOMETERS

1500 2000

1000

STATUTE MILES

oon & O.

CARROLLIA

OMI

“™<™ Antunesi Complex

ace

ANTUNESI

cerqueirai o

<\ ~wilsoni—;

propie iil

iiiabevalssass aa

AANA

CARROLLIA

iy 4

Nit

-—— 1.0 ———+4

SSS SS

SS Ss SSS S S Y

SSS SSSSSSSSSS

= SZ SS

SNS

FORE MID HIND

°))

OL iil

si td,

Isis i "

1 itt ——— cercus ) :

cere Ne FORE MID HIND

DORSAL \

atrial plate

bihaicolus

VZ 140

Aragua

Venezuela

HK. 0.2 —————_—___

ee IX tergite i

postgenital plate

aie

q \)

f (

atrial plate

cowl

sigma

insula

LATERAL CAUDAL

FORE

BLLZ2ZZ,

AAT \

Hig

y. HAAN

Wyle

His

urichii

St. Andrew

Trinidad

HIND

FORE MID HIND

WMI}

~—toB

W Wy G,

Gi Lg ff

ZING

YY WG AG 8 | Yi

Lg hf Yi) Ys

Mi iY,

Lys

SN TARRY

Zi &

i= OUs A 3)

WT WE wy

secundus

CR 412

Heredia

Costa Rica

CARROLLIA

bihaicolus infoliatus i] rausseoi

VZ 140 FG 179 VZ 156

Aragua Inini Aragua

Venezuela French Guiana : Venezuela

CIBARIAL DOME

} «* __ LATERAL FLANGE

(

Af a an

INN

i ae

VENTRAL PAPILLAE

metempsytus guerreroi

PA 347 Kes VZ

Paar ~<—#+—_ CLYPEUS v3 ae

Panama . Venezuela

anduzei

antunesi

BRA 54 TR 475 USNM 200

Para St. Andrew Canal Zone

Brazil Trinidad Panama

0.1

bonnei secundus babahoyensis

COL 65 PA 840 ECU 125

El Valle Canal Zone Los Rios

Colombia Panama Ecuador

distal divis

i yy

Fig. 10

CARROLLIA

external spine

bihaicolus

VZ 140

Aragua

Venezuela

CARROLLIA

bihaicolus

VZ 296

Aragua

Venezuela

Fig. 12

ig f

CARROLLIA cae

guerrerroi

VZ 394

Aragua

Venezuela

CARROLLIA

guerrerroi

VZ 390

Aragua

Venezuela

cs PT

Ma yilty

‘ yy Wh

P78 VI

Fig. 14

CARROLLIA

lao

LAM AY pp 8

Wie rausseoi

aa AM VZ 156

/ | \ Aragua

Venezuela

CARROLLIA if"

rausseoi

VZ 157

Aragua

Venezuela MP

Cs PY

Vill

rl Na .

5 3

Fig. 16

CARROLLIA

io)

——<—————|

g = SS

PA 376

Darien

Panama

spiniform (a)

11, 10

CR 491

San Jose

Costa Rica

seta

vi .

Sree S 10 44 tf

,) 414

"4 PO

4°

vil

metempsytus

external spine

CARROLLIA —_\\\\

metempsytus

CR 493

San Jose

Costa Rica

Vill

Ui ba

Ss p flit NW i!

x © St mS f

A \ J Bl?

4 : 3 f

Fig.18

CARROLLIA

\ 2

a we

infoliatus

FG 179

Inini

French Guiana

Wy f

Ze OWN ep Uys

Whi pe

sl:

as)

—~

Ill

a l4

CARROLLIA

a 12 1 ff

dees H

Y infoliatus

FG 179

Inini :

French Guiana MP

: \ cs PT

‘\

ae oer

an. im)

0040) Up 0

wih

Fig. 20

CARROLLIA £

TR 859

St. Andrew

Trinidad

St. George

Trinidad

CARROLLIA

TR 49

St. George

Trinidad

Fig. 22

CARROLLIA {| \_<° %

aC :

ve Fa

snout

N WR spiniform (a)

7 / il { \ » eee

A '— preapical lobe X44

distal division

CARROLLIA i

aie ta anduzei

Fig. 24

CARROLLIA

FG 179

Inini

French Guiana

distal division

accessory

division

proximal

division BB 557

CARROLLIA

oes

Nso

OS'S

Fae

outer curvature

___ snout

caudal process

accessory

division

mesal sternal lobe

secundus

Vt tre wt s

Ze We! Lips,

' J

ee ear

Sameera LY

Reg B AAS fi

4 1 Soe err By,

ey ie ieody,

Deg sch ppth ts df /

Rik vy]

Se / oS}

eae Z

CARROLLIA

secundus

PA 940

Colon

Panama

Fig. 28

CARROLLIA

32126-1

Rio de Janeiro

Brazil

USNM 1171-11

Sao Paulo

Brazil

distal division I/ proximal

c division

——

sues iridescens

514 10

Ill 11

I 9

‘| |

4 14

IV K 1

13 10

CARROLLIA

iridescens

32126-1 si

Rio de Janeiro

Brazil Pia

vil

>, 13 8 > 7

‘e apf 3 j

10/ 6 4 >

Vil

oS ; Syitits Z

= Wee Ge Si

=: Oh ee i

.\ 4

| Fig. 30

CARROLLIA

27866-1

Rio de Janeiro

Brazil

crest

\ — snout

Pes ~)

—c ——spiniform (a)

Napicotergal seta

‘— distal division

proximal division

USNM 1171-14

Sao Paulo

Brazil

nw 7

CARROLLIA

soperi

27866-1

Rio de Janeiro

azil

Fig. 32

CARROLLIA

WY \

NG is |

= SN

BESS

proximal

division

——

——)

————

Z—_Z

ZV ZZ BE

Gy a

my) oe

s ae

: 5 Ws sa

OSS 4, Ab, Aa

? < Lea

pits

Sy S

\ =

Gas

Asses

See

: =

se Se

SS SSF

= |

gaa es I if

8 0 ct

ia: Ties

6 af

As 3 0 —

/ |*_-+y*\ babahoyensis

% ZF

gait

2 CARROLLIA

Fig. 34

CARROLLIA

GG 107 IV :

Darien 4 \

Panama ‘,

: 10

\14

apicotergal seta

720309-1

Rio de Janeiro

Brazil

proximal division

\ distal division

caudal process

y : _ af + : Ee a

YY) '¥ antunesi

\ oy

AMY

CARROLLIA

antunesi

GG 107

Darien [. 14

Panama

Fig. 36

CARROLLIA /

cerqueirai

PA 654

Boca del Toro

Panama

distal

division

proximal division

accessory division

anal

—-

<—

cerqueirai

PA 654

Bocas del Toro

Panama

Fig. 38

CARROLLIA /

Paes

division

— “

eiisoni

Bates 65

Meta

Colombia

K/f5

: G

S Of

wilsoni

KO 10-43

Meta

Colombia

Vill

A My

Fig. 40

CARROLLIA

Bates 102

Meta

Colombia

USNM Komp. No.8

_ Meta

Colombia

apicosternal

process

proximal

division

distal division

CARROLLIA

Mair, | kompi

KO 10-30

Meta

Colombia

132 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

TABLE OF DISTRIBUTIONS

BIHAICOLUS IRIDESCENS

Uri. Iridescens

é 2}.| 18|_|s

QS [om a| =) <= Alal S|].

S/S] 2) EI SiS(S/S/ 8/3] | S/S] 818/81 818

Sele eee Siete) Sets] asl ores

alaldlaltlejolrfoldiafaAl[SIal{SiSizj} 4

MEXICO

aes ad a

Atlantic *

GUATEMALA

et Ma

Atlantic *

LE GB aos rene Cee N78 PSE A Si Renate

Pe ied oo Ce pee

HONDURAS

Pacific

Atlantic

NICARAGUA

Pacific

Atlantic *

ge a a eee

Pacific * * * *

Atlantic * * *

PANAMA

West x \l* * *

Central * * * *

East * xix * * *

ECUADOR

3 oe

Amazon *\* * *

COLOMBIA

Pacific * * * *

Caribbean * * *

Orinoco-Amazon Kix * * * *

VENEZUELA

Caribbean kk * *

Orinoco «| *

TRINIDAD a ere St ALY

GUIANAS ie ee Oe eee ae ee a

moe TAs sell le

Amazon Klik Kix

Other * xk Kix * *

ARGENTINA BESS: ECS ESS a) Rt aa

PERU PR ER Ea RS a ee ae

BOLIVIA CORR CEA. ee RRR cat 0

16. kompi

Valencia: Subgenus Carrollia of Culex — 133

INDEX TO SCIENTIFIC NAMES

Aedinus, 12

albicosta (Orthopodomyia), 14, 51

amazonensis (Culex), 9

anduzei;,2, -10, -11,..12, 43, 24K 23 125k,

26k, 47, 48, 49, 50, 52-54; figs. 3, 9,

oe, 23

Anoedioporpa, 12

antunesi, 1, 2, 10, 11, 12, 22k, 24k, 25k,

21K, 36}. 62," 67, 685-70," 71y 72;,73-76;

16,2 TT, 78s, 79,. SO, 281, 825 fiek: SY.

34, 35

antunesi of authors, 75, 79, 81

Antunesi Complex, 11, 12, 13, 55, 56, 57,

69, 72, 83, 84; fig. 5

Antunesi line, 11, 13, 56, 57, 72, 75, 78, 80;

fig 2D

aporonoma (Wyeomyia), 14, 44, 59, 63, 78

arthrostigma (Wyeomyia), 14, 63, 78

asulleptus (Limatus), 14, 32, 38, 41, 54, 59,

babahoyensis, 2, 10, 13, 22k, 24k, 25k, 27k,

56, 62, 67, 68, 69, 70-73, 73, 74, 75, 76,

77, 78, 79; 80, 82: figs 5, 9 32,33

Babahoyensis line, 11, 56, 57, 72, 81; fig. 5

Bahia form (sp. 8a), 11, 55, 60; fig. 4

bihaicolus, :2, 9, 10, 11, 13, 21k, 23k, 25k,

26K,°29, '29-33,-33,-36, 38, 39; 41/42,

43, 44, 51, 59, 63; figs. 2, 6, 9-11

bihaicolus of authors, 32, 42

Bihaicolus Complex, 10, 28-29, 32, 35, 38,

41, 44

Bihaicolus Group, 3, 5, 6, 7, 8, 10, 12, 13,

15, 16, 17, 19, 20, 20k, 23k, 24k, 26k,

27-29, 46, 47; fig. 2

bonnei, 2, 9, 10, 11, 12, 13, 21k, 24k, 25k,

27k, 32, 41, 44, 51, 55, 56, 57-60, 60, 61,

62, 63, 64, 65, 66, 67, 68; figs. 4, 9, 24, 25

bonnei of authors, 60

Carolia, 14

Carollia, 9, 14, 15

Carrollella, 9, 14, 15

Carrollia, 8,9, 10, 12, 13, 14-20

cerqueirai, 1, 11, 13, 22k, 24k, 25k, 27k,

56, 63, 68, 71, 72, 73,74; 75, 76-79, 79,

80, 81, 82; figs. 5, 36, 37

chidesteri (Culex), 14, 63

complosa (Wyeomyia), 14, 51

conservator (Culex), 14, 32

Corethrella spp., 14, 33, 72

corniger (Culex), 14, 32, 51, 63

Culex (Anoedioporpa) sp., 14,51, 54

Culex (Lutzia) sp., 14, 36

Culex (Melanoconion) sp., 14, 51

daryi (Aedes), 14, 32

Deinocerites, 5, 7, 8, 12

Dieffenbachia, 13, 32, 47

digitatum (Trichoprosopon), 14, 33, 38, 63,

72, 76

durhamii (Limatus), 14, 32, 41, 51, 59, 63,

eiseni (Anopheles), 14, 32, 38,51, 59

equinus (Haemagogus), 14, 32

Espirito Santo form (sp. 9a), 11, 13, 55, 56,

59, 63, 64, 67, 75, 76, 81; fig. 4

falco (Haemagogus), 14, 63

fascipes (Orthopodomyia), 14, 32, 51, 59

felicia (Wyeomyia), 14, 38

fluviatilis (Aedes), 14,32, 51

Galindomyia, 12

guerreroi, 2, 8, 10, 11, 14, 20k, 23k, 24k,

20K, 29, 30, 32y:33436. 37.:38, 40. 41:

figs, 2,9, 12, 13

Heliconia, 13, 32, 47, 78

homoeopus (Aedes), 14, 41

impostor (Aedes), 14, 32

infoliatus, 2, 4, 9, 10, 11, 12, 13, 15, 21k,

23k, 25k, 26k, 29, 31, 32, 41, 41-44, 59;

figs. 2,9, 18, 19

infoliatus of authors, 30, 41

Infoliatus Complex, 10, 29, 32, 41, 44, 55-

56, 72

insolitus (Aedes), 14, 51

ioliota (Aedes), 14, 32, 51

iridescens, 2, 4, 9, 10, 11, 13, 14, 22k, 24k,

25k, 26k; 55, 56, 57, 58, 39, 60,61, 63.

64-67, 67, 68, 69, 75, 76, 81; figs. 4, 28,

134 Contrib. Amer. Ent. Inst., vol. 9, no. 4, 1973

iridescens of authors, 9, 57, 59, 61, 73, 79

Iridescens Complex, 10, 11, 12, 13, 55-56,

56, 59, 63, 69, 76, 81, 84; fig. 4

Iridescens Group, 4, 7, 8, 11, 12, 13, 19, 20k,

23k, 25k, 26k, 44-46; figs. 3-5.

Iridescens Subgroup, 3, 5, 6, 7, 8, 11, 13, 21k,

23k, 25k, 26k, 46, 47, 54-57, 76, 84; figs.

4,5

iridicolor (Haemagogus), 14, 41

Janthinosoma, 9

kompi, 1, 4, 13, 15, 20, 22k, 24k, 25k, 26k,

56, 67, 68, 69, 70, 71, 72, 73, 76, 79, 81-

84; figs. 5, 40, 41

Kompi line, 11, 56, 84; fig. 5

kukenan (Culex), 14, 32

magnum (Trichoprosopon), 14, 36

manaensis, 2, 10, 73, 75

mathesoni, 2, 10, 48, 50

Melanoconion, 6, 9, 12

Melanoconion Section, 4, 5, 6, 7, 8, 12

metempsytus, 2, 9, 10, 11, 21k, 23k, 24k,

26k, 29, 32, 34, 35, 36, 37, 38, 39, 39-

41, 44, 59; figs. 2, 9, 16, 17

metempsytus of authors, 41, 42

Micraedes, 12

Microculex, 12

Mochlostyrax, 9

mollis (Culex), 14, 32,41, 51, 59

neivai (Anopheles), 14, 32

pallidiventer (Trichoprosopon), 14, 63

Panama form (sp. 3a), 10, 11, 12, 13, 26k,

28, 29, 37, 38-39, 41; fig. 2

podographicus (Aedes), 14, 51

quinquefasciatus (Culex), 14, 32, 36

rausseoi, 2, 10, 11, 21k, 23k, 24k, 26k, 27,

28, 29, 30, 32, 33, 34, 35, 36-38, 38, 39,

40, 41; figs. 2, 9, 14, 15

Sabethes spp., 14, 32, 41, 76

secundus, 2, 9, 10, 11, 14, 22k, 24k, 25k,

27k, 32, 55, 56, 57, 58, 59, 60-64, 64,

GS, 66; 675-68, 70071, 72, 73 74575,

76, 78, 79, 82; figs. 1, 4, 8, 9, 26, 27

secundus of authors, 10, 57, 59, 63, 64, 73,

73,76, 81

soperi, 2, 10, 13, 22k, 24k, 25k, 26k, 56, 66,

Oy 70 Ges lor 278s Toe 16; 79% 81982,

84; figs. 5, 30, 31

Soperi line, 11, 13, 56, 57, 69, 81; fig. 5

soperi (Haemagogus), 14, 32

sp. 3a (Panama form), 10, 11, 12,13, 26k,

28, 29, 37, 38-39, 41; fig. 2

sp. 8a (Bahia form), 11, 55, 60; fig. 4

sp. 9a (Espirito Santo form), 11, 13, 55, 56,

59, 63, 64, 67, 75, 76, 81; fig. 4

stonei (Culex), 14, 44, 59

terrens (Aedes), 14, 41

Tinolestes, 12

Toxorhynchites spp., 14, 32, 63, 72, 76

ulopus (Trichoprosopon), 14, 51

undosus (Sabethes), 14, 51, 59, 63, 78

urichii, 2, 9, 11, 12, 13,:14,)21k, 23k,.25k,

26k, 32, 47, 48-52, 52, 53, 59; figs. 3, 7,

9.20, ou

Urichii Subgroup, 3, 5, 6, 7, 8, 11, 12, 13,

LS yckOel Toc Qhk, 238k. 25k... 26k, 45046;

46-48, 50, 53; fig. 4

wilsoni, 1, 2, 10, 11, 12, 22k, 24k, 25k, 27k,

56, 68, 71, 73, 75, 76, 78, 79-81, 82; figs.

5, 38, 39

Wyeomyia spp., 14, 54, 76

Contributions

of the

American Entomological Institute

Volume 9, Number 5, 1973

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA. - XVI.

Genus Aedes Meigen, Subgenus Aedimorphus Theobald

in Southeast Asia.

John F. Reinert

CONTENTS

Pala fi ek ed Be eee a ed cS eee eda ee . 1

BEG EP ee ha TUG sig ss Sc detains gla i ea nah eae AR era: Mee LR ae aie g 2

GENUS AEDES MEIGEN, SUBGENUS AEDIMORPH US THEOBALD .... 4

LIST OF SPECIES OCCURRING IN SOUTHEAST ASIA ........ Rie tea

KEYS TO THE SPECIES OF AEDES A@EDIMORPHUS)

POU PEt AO Sac ee eS ea ee a Ne a a 14

IS TOOT RE AOU ete ete see Se a he ea eS Sees 14

RTE) er Og es el ee i ee a oe ee 16

KEY TO THE FOURTH STAGE LARVAE & £h 286 2k oar e.. 16

DESCRIPTIONS OF THE SPECIES OCCURRING IN SOUTHEAST ASIA .. 17

MID OSC CHAS (TDCORatE) es See ce es Rie Ao STs Bla leS s 17

ee ea ee sa sng we ae WS a es 23

culicinus Edwards ..... See Bane Goo ae ee a ans 29

ee a ee ee ee Oe oe, le sak es Gas 33

ION) (TPO) par a GS a 8 a os Ee OS 35

PI GIOOREH ST NGGREIG) 5 once FOES SS ow ee ee 8 38

néaros Vidids (Barraud), <6 ow eee ee LES ee. 43

OPO FAW Are. a Ra a ee a eS 45

pepe aostyieis (lheobald). 2. nk COM ae ee a de Re 50

pono cisis (ERMOW Se ee Ne Pee 54

Dees TONES) a ae hs Gh aa He ae Se Ne 59

Deer em Or es A IMCIOW os a: ae ata ew a Cae Bh 63

Pex DENS (WCITIES. i eee wom ee A 8 cal wee, es 66

Berns Pont | TP N@O0RI) 6 S56 ee Ce eee hac se ewes 79

WOTNUT ICN, DOIGAS sete perth 6 We eee sown BS ee ee es 83

MISIDENTIFICATIONS AND DOUBTFUL RECORDS ............ 84

BIE NOCET I POO ORICA es ee ae Se eG ee se eee 84

laéniovhynchoidées (Christophers) sc.) oe wk ke ek ee 85

POC ICING EOIN Loa Se Ne ee ee a ee ea ay: Be ita ee 86

ee ee ee Ee ca lye eg ae ae Gra ee i ce Ook tie aula ee mes ea ee 87

APPENDICES

TABLES 1-10. RECORDS OF THE BRANCHING OF THE SETAE

ON THE PUPAE OF AEDES (AEDIMORPHUS)

Be ae re ee ee wile a. ep 111

TABLE 1. alboscutellatus (Theobald)..... 111

TABLE 2, caecus (Theobald) ......... 113

TABLE 3. culicinus Edwards ......... 116

TABLE 4. mediolineatus (Theobald). ..... 118

TABLE 5;. ovbiige Howards <.4 . 3... «a 121

TABLE 6. pallidostriatus (Theobald) ..... 123

TABLE 7. pampangensis (Ludlow)....... 126

TABLE 8. pipersalatus (Giles) ........ 128

TABLE 9. vexans vexans (Meigen) ...... 131

TABLE 10. vexans nipponii (Theobald). .... 133

TABLES 11-16. SPECIES OF AEDES (AEDIMORPHUS)

OCCURRING IN EACH ZOOGEOGRAPHICAL

Te er ath ca aa Say in OB teh Sig ao a am ae Ge 136

TAG th. Orrental ee | ete s ack & oe 136

TA ao. ere ok a ee ee 137

TRE Te OP al eerie 5 ke ik se ee 137

TABLE 14. Pacific OceanIslands....... 137

Tete AO, NOAPCUC. 46 bk a ek ae So we 138

: TABLE TOS eERiOpian eo kee ee 138

Be ee orca gy a at Sh as we ia ate nb clos ee ue eat a eae ge ag ie Ve 142

Be A eat ena ge hiocg tak a Mae ROR Ais, Wem ec a atone PUR a he Salar cg 144

hE Oe SRURIM BARS AERA GRE EPSRC IPR cE pal dn Se GARE Aes opie bad bE bee a SAREE bagel 213

CONTRIBUTIONS TO THE MOSQUITO FAUNA OF

SOUTHEAST ASIA. - XVI.

GENUS AEDES MEIGEN, SUBGENUS AEDIMORPHUS THEOBALD

IN SOUTHEAST ASIA. |, 2

John F. Reinert?

ABSTRACT

This paper is a comprehensive revision of the Aedes (Aedimorphus)

in Southeast Asia and deals with 14 species and 1 subspecies. These species

are completely described and compared with closely related forms occurring

in the Pacific Islands and Oriental Zoogeographical Regions. Pupae of

caecus, culicinus, mediolineatus, orbitae, pampangensis and pipersalatus,

larvae of culicinus and orbitae and the egg of mediolineatus are described

for the first time. Stages and genitalia of the following species are also

illustrated for the first time: alboscutellatus male; caecus female genitalia,

male and pupa; culicinus female, female genitalia, male, pupa and larva;

lowisii female genitalia; mediolineatus female genitalia, male, pupa and egg;

nigrostriatus female, female genitalia and male; ovbitae female, female geni-

talia, male, male genitalia, pupa and larva; pallidostriatus female and fe-

male genitalia; pampangensis female genitalia and pupa; pipersalatus female

genitalia, male and pupa; punctifemoris female, female genitalia and male;

stenoetrus female genitalia and male; and taeniorhynchoides female genitalia

and male. Keys to the adults (including stenoetrus and taeniorhynchoides),

pupae and larvae of Southeast Asian species are given.

New synonyms in this paper are: lowisii (= mindoroensis); pampan-

gensis (= niveoscutellum); and vexans (= nocturnus). Lectotypes for lowisii,

nigrostriatus, orbitae and pallidostriatus and a neotype for pampangensis are

designated. Aedes stenoetrus and taeniorhynchoides are excluded from the

fauna of Southeast Asia.

IThis work was supported in part by Research Contract No. DA-49-193-MD-

2672 from the U.S. Army Medical Research and Development Command,

Office of the Surgeon General and made to the Southeast Asia Mosquito

Project, Smithsonian Institution, Washington, D.C. 20560.

2Taken in part from a dissertation submitted in partial fulfillment of the

requirements for the degree of Doctor of Philosophy, Department of

Entomology, University of Florida, Gainesville, Florida.

3Major, Medical Service Corps, U.S. Army, Department of Entomology,

Walter Reed Army Institute of Research, Walter Reed Army Medical Center,

Washington, D.C. 20012

2 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

INTRODUCTION

The subgenus Aedimorphus was originally described by Theobald

(1903a: 290) as a distinct genus based on a single African species, domesti-

cus (Theobald). A number of Oriental species now included with Aedimor-

phus were at various times placed under other genera and subgenera. Felt

(1904: 391c) erected a new genus, Ecculex, for the species vexans (Meigen)

(= sylvestris Theobald). The following year, Ludlow (1905: 94) described a

new genus Reedomyia for her new species pampangensis. During this same

year, Theobald described 3 new genera: Pecomyia (1905a: 23) for taeni-

orhynchoides (= maculata Theobald), Pseudograbhamia (1905c: 243) for piper-

salatus (= maculata Theobald), and Lepidotomyia (1905b: 80) for alboscutella-

tus Theobald. Geitonomyia was described by Leicester (1908: 134) for caecus

(Theobald) and Lesligomyia by Christophers (1911: 68) for his new species

taeniorhynchoides. Edwards (1913: 227) placed several of the Oriental spe-

cies in the genus Ochlerotatus Lynch Arribalzaga but later (1922b: 466) trans-

ferred them to the genus Aedes Meigen subgenus Ecculex.

Two Oriental species, alboscutellatus and vexans, were moved by

Edwards (1924: 372) to the subgenus Aedimorphus and the next year (1925:

267) he synonymized Ecculex with this subgenus. In the Genera Insectorum,

Edwards (1932: 165) reviewed Aedimorphus from a worldwide standpoint and

divided it into 8 groups. The first comprehensive study of the subgenus in

the Oriental area was conducted by Barraud (1928: 655) and included the spe-

cies occurring in India. He later (1934: 246) revised the Indian species.

Knight and Hurlbut (1949: 29) and Knight and Hull (1951: 200) modified Ed-

ward's group classification of the Aedimorphus by combining Group D with

Group C.

The present review deals with 14 species and 1 subspecies from

Southeast Asia and compares them to closely related species in the Oriental

and Pacific Islands Zoogeographical Regions. Pupae of caecus, culicinus,

mediolineatus, orbitae, pampangensis and pipersalatus, larvae of culicinus

and orbitae and egg of mediolineatus are described for the first time herein.

Stages and genitalia of the following species are also illustrated for the first

time: alboscutellatus male; caecus female genitalia, male and pupa; culicinus

female, female genitalia, male, pupa and larva; lowisii female genitalia;

mediolineatus female genitalia, male, pupa and egg; nigrostriatus female,

female genitalia and male; orbitae female, female genitalia, male, male

genitalia, pupa and larva; pallidostriatus female and female genitalia; pam-

pangensis female genitalia and pupa; pipersalatus female genitalia, male and

pupa; punctifemoris female, female genitalia and male; stenoetrus female

genitalia and male; and faeniorhynchoides female genitalia and male. Keys to

the adults (including stenoetrus and taeniorhynchoides), pupae and larvae of

the Southeast Asian species are given. For original descriptions of the Ori-

ental and Pacific Islands species not found in Southeast Asia see: Theobald

(1905b: 86) trimaculatus ; Theobald (1907: 395) stenoetrus; Christophers (1911:

68) taeniorhynchoides; Edwards (1914: 77) jamesi; Barraud (1928: 662) syn-

theticus (as fisheri); Stone (1939: 163) oakleyi; Carter and Wijesundara (1948:

139) argenteoscutellatus; Knight and Hurlbut (1949: 27) senyavinensis; Bohart

(1956: 63) tvukensis; and Reinert (1972b: 357 gouldi). For taxonomic information

on the African species check Edwards (1941) and Hopkins (1952).

New synonyms in this paper are: lowisii (= mindoroensis); pampan-

gensis (= niveoscutellum); and vexans (= nocturnus). Aedes stenoetrus and

taeniorhynchoides are excluded from the fauna of Southeast Asia.

Reinert: Aedes (Aedimorphus) in Southeast Asia 3

During the course of this revision, I examed all specimens and types

of Aedimorphus in the United States National Museum (Natural History) and

the British Museum (Natural History) as well as specimens from numerous

individual and museum collections. Lectotypes for lowisii, nigrostriatus,

orbitae and pallidostriatus and a neotype for pampangensis are designated in

this paper.

Abbreviations used in references to literature conform to the World

List of Scientific Periodicals, 4th edition, Butterworths, Washington, 1963.

In the synonymy sections, an asterisk following the abbreviations used (A =

adult, 9? =female, c= male, P =pupa, L = larva, E = egg) indicates that at

least some portion of that sex or stage is figured. Abbreviations used in the

pupal descriptions, illustrations, tables and key are: C = cephalothorax;

P = paddle; and I-VIII = abdominal segments 1 through 8. In larval descrip-

tions the range of hair branching is followed by the mode branching in paren-

theses and the following abbreviations signify: A = antenna; C = head; M =

mesothorax; P = prothorax; S = siphon; T = metathorax; and VIII, X = abdomi-

nal segments 8 and 10. Abbreviations used in larval illustrations are as

above with the following additions, CS = comb scale and PT = pecten tooth.

When possible 10 specimens were used in determining the range and mode

hair branching in pupal and larval descriptions. Measurement scales on the .

illustrations are in millimeters. Distribution records are indicated as fol-

lows: countries are in capital letters; where known, primary administrative

divisions are in italics; and place names have the first letter capitalized. The

spelling of Thai changwats and locality names is taken from the Official Stand-

ard Names Gazetteer No. 97 prepared by the Office of Geography, Department

of the Interior, Washington, D.C., April 1966. Spelling of South Vietnam

provinces and locality names is taken from the Official Standard Names Gazet-

teer No. 58, 2nd Edition, prepared by the Geographic Names Division, U.S.

Army Topographic Command, Washington, D.C., May 1971. Locality names

which do not appear in the gazetteers are spelled according to the labels on

the specimens.

Information in the BIOLOGY section of each species is taken from the

collection sheets and the labels attached to the specimens. Biological data

recorded in the literature are also given and the sources are cited.

_ The nomenclature and chaetotaxy used for females, female genitalia,

males and male genitalia follow Knight (1970), Knight and Laffoon (1970a,

1970b, 1971) and Laffoon and Knight (1971) and those for the adult wing vena-

tion, pupae and larvae follow Belkin (1962). The terminology of the egg is

from Kalpage and Brust (1968). Female genitalia descriptions are based on

0 or more specimens except for a few species for which less material was

available, therefore the ranges of variation could possibly exceed those re-

corded especially for lowisii, nigrostriatus and pallidostriatus. All measure-

ments of the female genitalia were made on specimens that had been cleared,

dissected, arranged in a flattened position and mounted in Canada balsam on

microscope slides. Terga VIII and IX and sternum VIII measurements are

taken from sclerotized areas. The following definitions and figure 70 explain

new terms not described by Laffoon and Knight (1971).

Cercus index = Dorsal measurement of distance from apex to

the most anterior point of base (dorsal cercus —

length) divided by measurement of width at 0.50

distance from apex (cercus width).

Cercus/dorsal PGL index = Dorsal cercus length divided by dor-

sal postgenital lobe length.

4 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Dorsal postgenital lobe index (dorsal PGL index) = Dorsal

measurement of distance from apex to mid-

point of peri-anal membrane attachment

(dorsal PGL length) divided by measurement

of width at 0.50 length (dorsal PGL width).

Peri-anal membrane = Membrane attached to ventrobasal

portion of cerci, extending around the anus

and attached to the dorsal surface of the post-

genital lobe, the latter point of attachment

usually appears as an inverted ''U".

Tergum IX index = Dorsal measurement of distance from apex

to the most anterior point of base (tergum IX

length) divided by measurement of width at

widest point (tergum IX width).

Tergum VIII index = Dorsal measurement of distance from

apex to most anterior point of base (tergum

VIII length) divided by measurement of width

at widest point (tergum VIII width).

Sternum VIII index = Ventral measurement of distance from

apex to most anterior point of base (sternum

VIII length) divided by measurement of width

at widest point (sternum VIII width).

Ventral postgenital lobe index (ventral PGL index) = Ventral

measurement of distance from apex to midpoint

of posterior margin of the upper vaginal lip

(ventral PGL length) divided by measurement

of width at 0.50 of dorsal length (dorsal PGL

width). |

In the pupal descriptions, the number of branches on abdominal hair 1-I is

counted on the basal 0. 33 of the hair.

Southeast Asia, in this study, is composed of southern China, Bangla-

desh (East Pakistan), Assam, Burma, North Vietnam, South Vietnam, Taiwan,

Philippines, Indonesia, Laos, Cambodia, Thailand, Malaysia, Singapore, An-

daman and Nicobar Islands, Southern Ryukyus, Hainan and The Pescadores.

GENUS AEDES MEIGEN

SUBGENUS AEDIMORPHUS THEOBALD

Aedimorphus Theobald 1903, Monogr. Cul. 3: 290 (July).

Haplotype: Uvanotaenia domestica Theobald.

Catageiomyia Theobald 1903, Mem. Lpool. Sch. trop. Med. 10 (App.): i (Nov.).

Haplotype: Catageiomyia senegalensis Theobald.

Ecculex Felt 1904, Bull. N.Y. St. Mus. 78: 391c. Orthotype: Culex sylvestris

Theobald.

Reedomyia Ludlow 1905, Can. Ent. 37: 94. Haplotype: Reedomyia pampan-

gensis Ludlow. |

Lepidotomyia Theobald 1905, Ann. Mus. nat. Hist. Hung. 3: 80. Haplotype:

Lepidotomyia alboscutellata Theobald.

Pecomyia Theobald 1905, J. econ. Biol. 1: 23. Haplotype: Pecomyia macu-

lata Theobald.

Reinert: Aedes (Aedimorphus in Southeast Asia 5

Polyleptiomyia Theobald 1905, Genera Insec., Fasc. 26: 21.

Haplotype: Stegomyia albocephala Theobald.

Pseudograbhamia Theobald 1905, J. Bombay nat. Hist. Soc. 16: 243.

Haplotype: Pseudograbhamia maculata Theobald.

Duttonia Newstead 1907, in Newstead, Dutton and Todd, Ann. trop. Med.

Parasit. 1: 17. Logotype: Duttonia tarsalis Newstead.

Mimeteculex Theobald 1908, Rep. Wellcome trop. Res. Lab. 3: 258.

Haplotype: Mimeteculex kingii Theobald.

Geitonomyia Leicester 1908, Cul. Malaya, p. 134.

Haplotype: Culex caecus Theobald.

Myxosquamus Theobald 1909, Colon. Rep. misc. Ser. No. 237: 7; Theobald

1910, Monogr. Cul. 5: 225. Haplotype: Myxosquamus confusus

Theobald.

Stenoscutus Theobald 1909, Colon. Rep. misc. Ser. No. 237: 7; Theobald

1910, Monogr. Cul. 5: 263. Haplotype: Stenoscutus africanus Theobald.

Bathosomyia Theobald 1909, Colon. Rep. misc. Ser. No. 237: 9; Theobald

1910, Monogr. Cul. 5: 267. Haplotype: Bathosomyia abnormalis

Theobald.

Neopecomyia Theobald 1909, Colon. Rep. misc. Ser. No. 237: 12; Theobald

1910, Monogr. Cul. 5: 261. Haplotype: Neopecomyia uniannulata

Theobald.

Leslieomyia Christophers 1911, Paludism 2: 68. Haplotype: Leslieomyia

taeniorhynchoides Christophers.

The following description of the subgenus applies to species from the

Oriental and Pacific Islands areas.

FEMALE. Head. Antenna brown, 0. 87-1.14 length of proboscis,

pedicel usually pale with a patch of short fine hairs, scales or both mesally,

flagellomere 1 pale with a few small scales; clypeus brown, bare; maxillary

palpus 0.17-0. 25 length of proboscis; proboscis usually brown with pale ven-

tral markings, 0.98-1.33 length of femur I; eyes narrowly separated; inter-

ocular and ocular setae well developed; ocular line covered with narrow pale

scales (punctifemoris has broad scales on this area); vertex with dorsum

covered with narrow decumbent scales except in punctifemoris and wainwrighti

which have dorsum covered with broad decumbent scales; numerous erect

forked scales on occiput and vertex extending anteriorly to ocular line (vertex

of wainwrighti without erect forked scales). Thorax. Scutum covered with

narrow curved scales (punctifemoris also has small patches of broad white

scales), arrangement and color of scale patterns varies with the species;

scutellum with scales broad, narrow or both; median anterior promontory,

acrostichal (absent in wainwrighti), anterior and posterior dorsocentral (few

bristles present in wainwrighti), scutal fossal, supra-alar, posterior medial

scutal, postalar callar, and median and lateral scutellar bristles well developed;

antepronota widely separated with narrow curved scales (culicinus, puncti-

femoris, vexans vexans, vexans nipponii and wainwrighti have some broad

scales) and several well developed bristles; postpronotum with narrow curved

scales (culicinus, vexans vexans and vexans nipponii also have a few broad

ones and punctifemoris has only broad scales) and 3-10 posterior bristles;

propleuron with broad scales (mediolineatus and pallidostriatus have narrow

scales) and numerous bristles; prosternum bare of scales and bristles except

for members of the vexans group (stenoetrus, syntheticus, vexans vexans and

vexans nipponii) which possess scales, also 1-3 bristles in the 2 subspecies of

vexans; postspiracular area with or without broad or narrow scales and 3-11

6 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

bristles; subspiracular area with 1 or 2 patches of scales (orvbitae has this

area bare and alboscutellatus, oakleyi, senyavinensis, trukensis and lowisti

have a patch of short fine hairs); mesepisternum with an upper and a posterior

patch of broad scales and several upper and posterior bristles, lower ones

shorter; prealar knob with several bristles and with or without scales; para-

tergite with scales (lowisii without scales); mesepimeron with a patch of broad

scales and several bristles on upper area, lower area bare; other pleural

areas bare. Legs. Coxae I-III each with several bristles and usually broad |

scales; trochanters I-III with broad scales; femora I-III with various scale

patterns but usually each with a white dorsoapical spot, III usually with most

of anterior surface pale scaled; tibiae I-III with various scale patterns; tarsi

I-III with basal pale bands, dorsobasal pale spots or without ornamentation

(lowisii also has apical pale scales on tarsomeres); posttarsi I-III each with

2 ungues, I, II with ungues equal in size and each bearing a tooth and III with

ungues equal in size and simple (III toothed in pampangensis and toothed or

simple in stenoetrus and vexans). Wing. Dorsal veins usually covered with

moderately broad brown scales with a small pale patch at base of costa, a few

species with pale scales intermixed; alula with narrow scales along fringe;

remigium with 1-4 bristles, usually 2. Abdomen. Tergum I with a rectangu-

lar patch of pale scales on laterotergite; terga with or without basal pale bands

and usually with lateral pale spots; sterna usually pale scaled with some brown

markings; terga and sterna with numerous golden bristles, mostly along pos-

terior margins. Genitalia. The following is a composite description of the

Southeast Asian species. Tergum VIII medium sized, trapezoid-shaped with

base somewhat indented or with apex broadly rounded, bristles scattered over

entire surface, none to 3 small scales, 0.60-1.00 retracted into segment VII

(except caecus which is only 0.30-0.50 retracted), index 0. 73-1.44; sternum

VIII large, heart-shaped with a deep median apical indentation, short anda

few long bristles scattered over entire surface but most numerous apically,

none to 3 small scales, index 0. 78-1.11; tergum IX with moderately pig-

mented lateral portions, bilobed with 3-14 bristles on each lobe, index 0. 78-

1.58; insula tongue-like, covered with minute setae and with 3-7 small tuber-

culi on apical 0.25 (except vexans vexans and vexans nipponii which lack tuber-

culi); lower vaginal lip narrow, semicircular shaped, moderately pigmented

(occasionally heavily pigmented in mediolineatus), covered with minute setae;

upper vaginal lip narrow, heavily or occasionally moderately pigmented,

covered with minute setae, semicircular shaped with caudal margin flat; upper

vaginal sclerite large, moderate to heavily pigmented, triangular shaped;

postgenital lobe moderately long, apex with a moderately deep median indenta-

tion (except pallidostriatus which has a small one and orbitae with a deep one),

3-10 bristles on each side of midline, covered with minute setae, dorsal PGL

index 0. 70-1.48, ventral PGL index 1.52-3.29; cercus moderately long to

long, apex sharply rounded, numerous short and moderately long bristles

scattered over dorsal and lateral surfaces and a few long ones apically, com-

pletely covered with minute setae, index 2.08-4.27, cercus/dorsal PGL index

2.55-5. 22; 3 spherical, pigmented seminal capsules, 1 large and 2 slightly

smaller ones (except in alboscutellatus, culicinus, lowisii and the other Ori-

ental members of the group jamesi, oakleyi, senyavinensis and trukensis

which have 1 large one and 2 tiny rudimentary ones).

MALE. Similar to female in general habitus. Head. Antenna plu-

mose with hairs directed mainly dorsally and ventrally; maxillary palpus with

apical 2 segments short and down-turned with numerous apical and ventro-

lateral bristles, antepenultimate segment with apical portion somewhat swollen

Reinert: Aedes (Aedimorphus) in Southeast Asia ie

and with several ventrolateral bristles, longer than proboscis by 0.50 to 1.00

length of apical segment. Thovax. Prosternum bare including members of the

vexans group. Legs. Posttarsi I-III each with 2 ungues, I, II with ungues un-

equal in size and each bearing a tooth, III with ungues equal or unequal in size

and simple (toothed in stenoetrus). Genitalia. Tergum IX bilobed with 3-11

bristles on each lobe, entire surface covered with minute spicules; gonocoxite

moderately long to long and moderately broad, dorsal surface usually covered

with scattered short bristles with longer ones at apex, lateral margin with long

stout bristles from base to apex, ventral surface usually with long stout bris-

tles on distal 0.50, scattered scales on lateral and ventral surfaces; gonosty -

lus usually complex with apical portion expanded into a mesal lobe and a later-

al horn-like structure (gonostylus blade-like in stenoetrus, syntheticus, vex-

ans vexans and vexans nipponii), gonostylar claw attached mesally near middle

and with 1 or more accessory claws at apex; basal mesal lobe with 3-38 bris-

tles (stenoetrus has 118-128 bristles) and covered with short hair-like spicules;

proctiger short to long, paraproct smooth and bluntly pointed or slender with a

rounded apex and a subapical thumb-like process, cercal setae absent; phallo-

some with aedeagus with 2 lateral plates connected basally, each plate with

4-9 teeth and covered by a dorsal flap, paramere long; parameral apodeme

broad basally and tapering into a long narrow distal arm; sternum IX large

with the entire surface covered with minute spicules and 2-15 bristles near

center.

The genitalia of the Oriental species are divided into 2 types based on

characteristics of the phallosome and proctiger. TypelI (Figs. 33, 34) has

the aedeagus divided into 2 lateral plates connected basally, each plate bears

4-7 short blunt lateral teeth on distal 0.50 and is covered with a pigmented

dorsal flap which is narrow distally and broadly rounded proximally (when

aedeagus is dissected the teeth pull away from the lateral plate and remain

attached to the dorsal flap). The proctiger is short and the paraproct is dis-

tally slender and apically rounded with a subapical thumb-like process. The

majority of the Oriental species have typeI genitalia. Type II (Figs. 33, 47)

has the aedeagus divided into 2 lateral plates connected basally, each plate

bears 5-9 short to moderately long, longitudinal lateral teeth with tergally

curved apices and is covered with a pigmented dorsal flap which is narrow

distally and tapers to a broad base. The proctiger is long and the paraproct

is narrow with a bluntly pointed apex.

PUPA. The pupae of the species that occur in Southeast Asia do not,

at this time, present any clear-cut subgeneric characters. The pupal chaeto-

taxy is summarized as follows: Cephalothovax. Hairs 1, 2-C with 2-8

branches; 3-C with 2-5 branches; 4-C single to 9 branched; 5-C single to 8

branched; 6, 9-C single to 4 branched; 7-C with 2-11 branches; 8-C with 2-9

branches. Metanotum. Hair 10-C with 4-33 branches; 11-C single; 12-C

single to 11 branched. Abdomen. Hairs 0-II-VIII single; 1-I with 16-46

branches on basal 0.33, 1-II with 3-49 branches, 1-III with 3-18 branches,

1-IV with 2-13 branches, 1-V single to 10 branched, 1-VI single to 13 branched,

1-VII single to 10 branched; 2-I-VII single or double; 3-I-III single to triple,

3-IV with 2-12 branches, 3-V single to 5 branched, 3-VI single to 8 branched,

3-VII with 2-16 branches; 4-I with 2-7 branches, 4-II with 2-16 branches, 4-III,

VIII single to 8 branched, 4-IV single to 4 branched, 4-V single to 14 branched,

4-VI single to 11 branched, 4-VII single to 10 branched; 5-I with 2-18 branches,

o-II with 2-14 branches, 5-III with 2-13 branches, 5-IV, V single to 5 branched

0-VI single to 6 branched, 5-VII single to 9 branched; 6-I single to triple, 6-II

single to 5 branched, 6-III-VI single to 8 branched, 6-VII with 3-14 branches;

8 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

7-I, IV single to 6 branched, 7-II single to 10 branched, 7-III with 2-9 bran-

ches, 7-V single to 11 branched, 7-VI single to 4 branched, 7-VII single to 5

branched; 8-III, VII single to 7 branched, 8-IV-VI single to 5 branched; 9-I-VI

single, 9-VII with 2-11 branches, 9-VIII with 4-14 branches; 10-III single to 6

branched, 10-IV, VII single to 5 branched, 10-V single to 4 branched, 10-VI

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

branched; 14-III-VIII single. Paddle. Hair 1-P single to triple and short.

The respiratory trumpet is moderately broad and usually moderately pigmented.

The paddle is ovoid, with tiny to minute serrations along the basal 0.50 of

outer margin, usually with tiny spicules along apical margin, and the midrib

does not reach the apex.

LARVA. The larval chaetotaxy and features of the Southeast Asian

species are summarized as follows: Head. Hairs 1, 3-C single; 4-C small

with 2-13 branches, usually with 5-7 branches; 5-C long, single to 9 branched,

usually with 4-7 branches; 6-C long, single to 9 branched, usually with 4-6

branches; 7-C long with 4-16 branches, usually with 6-10 branches; 8, 9-C

single to 6 branched, usually with 2-4 branches; 10-C single to 4 branched,

usually double or triple; 11-C with 2-10 branches, usually with 3-6 branches;

12-C with 3-10 branches, usually with 3-6 branches; 13-C single to 9 branched,

usually with 2-4 branches; 14-C single or double, usually single; 15-C single

to 7 branched, usually with 2-5 branches; basal maxillary hair single; mental

plate with 22-36 teeth. Antenna. Usually moderately pigmented, slightly in-

curved distally, usually numerous short stout spicules on shaft; hair 1-A long

with 3-16 branches, usually with 5-10 branches, inserted at 0.36-0.52 from

base; 2-A long; 3-A approximately equal to or about 0.50 length of 2-A.

Thorax. Hair 0-P with 3-21 branches, usually with 5-10 branches; 1, 5, 6-P

single; 2, 3, 9, 11-P single to 4 branched, usually double or triple; 4, 8-P

single to 7 branched, usually double or triple; 7-P with 2-4 branches, usually

double or triple; 10, 12-P single or double, usually single; 14-P single to

triple, usually single or double; 1-M single to 8 branched, usually with 3-5

branches; 2-M single to 5 branched, usually with 2-4 branches; 3-M single to

5 branched, usually single to triple; 4-M with 2-9 branches, usually with 3-4

branches; 5-M single to triple, usually single; 6-M with 3-10 branches, usually

with 4-6 branches; 7, 10, 12-M single; 8-M with 3-10 branches, usually with

5-8 branches; 9-M with 3-10 branches, usually with 6-9 branches; 11-M single

or double, usually single; 13-M with 3-12 branches, usually with 5-8 branches;

14-M with 4-11 branches, usually with 5-8 branches; 1-T single to 4 branched,

usually single or double; 2-T with 2-13 branches, usually with 3-6 branches;

3-T with 2-28 branches, usually with 6-10 branches; 4-T with 2-6 branches,

usually with 3-4 branches; 5, 10-T single; 6-T single to 5 branched, usually

single to triple; 7-T with 5-14 branches, usually with 6-10 branches; 8-T with

3-12 branches, usually with 4-7 branches; 9-T with 2-9 branches, usually with

4-6 branches; 11-T single or double, usually single; 12-T single to triple,

usually single or double; 13-T with 4-19 branches, usually with 5-9 branches.

Abdomen. Hairs 0, 14-VIII single; 1-VIII with 3-12 branches, usually with

4-9 branches; 2-VIII single to 5 branched, usually double or triple; 3-VIIT with

5-27 branches, usually with 6-12 branches; 4-VIII single to 4 branched, usually

double or triple; 5-VIII with 4-10 branches, usually with 4-7 branches; comb

of 7-32 scales, usually arranged in 2 irregular rows; 1-X single to 4 branched,

usually single to triple; 2-X with 5-15 branches, usually with 8-12 branches;

3-X single; ventral brush varies from 8 hairs on grid and 4 precratal ones to

14 hairs on grid and 3 precratal ones, some hairs always with 10 or more

branches; saddle usually moderately pigmented with small spicules along

Reinert: Aedes (Aedimorphus) in Southeast Asia 9

posterior margin, incompletely rings segment X, acus present or absent; 4

anal papillae usually moderately long to long and slender. Siphon. Usually

moderately pigmented; acus usually present; index 2.11-9.00; pecten with 10-

23 teeth, distal 2-4 teeth smooth and wider spaced than remainder which have

a Slender apical attenuated filament with 1-4 lateral denticles; hair 1-S with

3-7 branches, usually with 4-6 branches, inserted at 0. 54-0. 85 from base.

EGG. The eggs of only 3 species, domesticus, mediolineatus and

vexans are known in the subgenus Aedimorphus. Eggs of domesticus are

described by Reinert (1972a: 60), of vexans by Horsfall and Craig (1956: 370),

Myers (1967: 796) and Kalpage and Brust (1968: 711-712) and of mediolineatus

in this paper. A summary of the egg characters follows: Shape. Fusiform to

spindle shaped. Size. In microns, domesticus 855-885 by 242-255, medio-

lineatus 735-840 by 190-200 and vexans 614-756 by 167-224. Color. Brown

or bronze. Chorion. The surface reticulation of domesticus and vexans is

similar and consists of a pattern of axially arranged cells, hexagonal and

polygonal in shape and 3-6 (domesticus) or 2.5-5 (vexans) times as long as

wide with cell walls raised. Aedes mediolineatus differs from domesticus

and vexans and has a reticulation composed of polygonal cells of varied size

with cell walls raised.

DISTRIBUTION. Species of Aedimorphus are confined to the Ethiopian,

Oriental, Pacific Islands and northern portion of the Australian Zoogeographi-

cal Regions with the exceptions of vexans vexans which has a wide distribution

and is found in the Holarctic, Oriental, Pacific Islands and Australian Zoogeo-

graphical Regions and vexans nipponii which is confined to the eastern part of

the Palearctic Zoogeographical Region.

Kighty -two species and 5 subspecies (Table 16) of Aedimorphus are

endemic to the Ethiopian Zoogeographical Region. This area contains the

largest number of species and shares only vexans vexans with other regions.

The Oriental Zoogeographical Region possesses 21 species (Table 11) and

shares 5 (Table 12) of these with the Australian, 2 (Table 13) with the Palearc-

tic, 3 (Table 14) with the Pacific Islands and 1 (Table 15) with the Nearctic

Zoogeographical Regions.

Species of Aedimorphus have been collected from each of the countries

in the Oriental and Ethiopian Zoogeographical Regions.

TAXONOMIC DISCUSSION. The subgenus Aedimorphus, in the Ori-

ental Region, possesses a combination of characters that allows it to be

separated from the other subgenera of Aedes. The most distinctive features

are exhibited by the male, these being: antenna with the plume hairs directed

mainly dorsally and ventrally; maxillary palpus with the 2 apical segments

short and down-turned with numerous apical and ventrolateral bristles, ante-

penultimate segment with apical 0.25 somewhat swollen and with several ven-

trolateral bristles and longer than proboscis by 0.50 to 1.00 length of apical

segment; genitalia with gonostylus complex and usually expanded distally with

1 or more gonostylar claws, proctiger short to long, paraproct smooth and

bluntly pointed or slender with apex rounded and a subapical thumb-like proc-

ess, cercal setae absent (present in some African species) and the phallosome

which has the aedeagus composed of 2 lateral pigmented plates connected bas-

ally, each plate with 4-7 short blunt lateral teeth on distal 0.50 or 5-9 short

to moderately long, longitudinal lateral teeth with tergally curved apices (vex-

ans and its allies) and covered with a pigmented dorsal flap; and tarsomere 5

of tarsus I ventrally concaved with the base projecting posteroventrally into

a protuberance with 3-4 short stout curved spines at apex and a pair of short

bristles each arising from a conical tubercle located near the middle of the

10 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

ventral margin. A similar maxillary palpus is also found in Ochlerotatus,

some Finlaya Theobald and Chaetocruiomyia Theobald while the antenna is

typical of Diceromyia, Stegomyia Theobald, Ochlerotatus and some Finlaya.

The toothed aedeagus of the vexans group is also similar to Diceromyia and

some Stegomyia but is easily separated from the latter by the presence of the

dorsal flap and from the former by the flap being pigmented and semirigid

while in Diceromyia it is unpigmented or lightly pigmented and membranous

(for a discussion of the Diceromyia see Reinert 1970). Males of Aedes (Ayura-

kitia) Thurman and Udaya Thurman have a pigmented dorsal flap over a toothed

aedeagus which is similar in structure to vexans. Ayurakitia is separated

from Aedimorphus by the absence of postspiracular bristles in the former.

Udaya differs from Aedimorphus by the absence of acrostichal bristles. The

tarsomere 5 of Aedimorphus resembles those of the tarsi I of Psorophora

Robineau-Desvoidy, Culex Linnaeus and several other subgenera of Aedes.

Aedimorphus males are easily separated from Ochlerotatus, Finlaya and

Chaetocruiomyia by the structure of the aedeagus which is simple and non-

toothed in the latter 3 subgenera. The maxillary palpus of Stegomyia is slender

with the apical 2 segments upturned and is markedly different from Aedimorphus.

Other important characteristics of both sexes of the adults are: vertex

with narrow curved decumbent scales medially (bunctifemoris and wainwrighti

with vertex covered with broad decumbent scales); erect forked scales numer-

ous on occiput and vertex extending anteriorly to ocular line (these scales re-

stricted to occiput in wainwrighti); female maxillary palpus short, 0.17-0. 33

length of proboscis; acrostichal and dorsocentral bristles well developed and

numerous (acrostichals absent and dorsocentral bristles few in wainwrighti ) ;

scutum covered with narrow curved scales (a few small patches of broad scales

in punctifemoris); antepronotum and postpronotum usually with narrow curved

scales; scutellum with narrow, broad or both types of scales; propleuron with

numerous bristles; several upper bristles on mesepisternum; and lower mesepi-

meron without bristles or scales.

The females have the following features: genitalia with tergum VIII

mostly or completely retracted into segment VII, sternum VIII with a deep

median apical indentation and heart-shaped, cerci moderately long to long,

narrow, extended from segment VII and visible dorsally, postgenital lobe with

a median apical indentation, insula tongue-like, covered with minute setae and

with 3-7 small tuberculi on apical 0.25, 3 seminal capsules approximately equal

in size except for the alboscutellatus group which has 1 large and 2 rudimentary

seminal capsules; and posttarsi of legs I-II with ungues each toothed and ungues

of III usually simple. Ochlerotatus has similar cerci but can be easily separated

by the presence of bristles on the insula. The female genitalia of Aedes (Neo -

melaniconion) lineatopennis (Ludlow) are very close to those of Aedimorphus.

The insula of Aedimorphus resembles those of Stegomyia and Diceromyia but

the cerci of the latter 2 species are short and broad. Female specimens usually

can be separated from those of the other subgenera of Aedes by a combination of

the characters mentioned.

For a long time, Aedes nummatus had been considered as an aberrant

Aedimorphus. It possesses some features of Aedimorphus but contains a greater

number of important characters of Diceromyia, notably features of the male

maxillary palpus, male genitalia and female genitalia. It is therefore being

transferred, along with a closely related new species, to the subgenus Dicero-

myia (Reinert in press).

The pupae of the Oriental species of Aedimorphus are very similar,

especially in the chaetotaxy, shape of the respiratory trumpet and structure of

Reinert: Aedes (Aedimorphus) in Southeast Asia 11

the paddle.

Aedimorphus larvae have a number of features, when used in combina-

tion, that allow them to be separated from the other Oriental subgenera of

Aedes such as: antenna moderately long to long, slightly incurved distally and

with numerous short, stout spicules on shaft, hair 1-A long, barbed with 3-16

branches; head hair 4-C always small and multiple branched, closer to 5-C

than to 6-C and 5, 6-C long, barbed, usually close together and forming a

diagonal line with 7-C (exceptin vexans and its allies which have 6-C anterior

to 5,7-C and 4-C approximately equal distance from both 5, 6-C but slightly

closer to 6-C; vexans head hair arrangement and other larval features are

similar to many Ochlerotatus); siphon usually long with distal 2-4 pecten teeth

usually smooth and always wider spaced than remainder which have a slender

apical attenuated filament with 1-4 lateral denticles, pecten teeth on basal

0.50 of siphon except in syntheticus and hair 1-S multiple branched, usually

small, inserted on distal 0.45, always distal to last pecten tooth; ventral brush

of abdominal segment X varies from 8 hairs on grid and 4 precratal ones to 14

hairs on grid and 3 precratal ones, hairs of ventral brush each with 8-18

branches except vexans nipponii which have 4-12 branched hairs, some hairs

always with 10 or more branches; and siphon usually with an acus.

The larvae of Aedes subgenera Ochlerotatus, Neomacleaya Theobald,

Verrallina Theobald, Edwardsaedes Belkin and Levua Stone and Bohart are

very similar to Aedimorphus but can be separated by the following characters:

Ochlerotatus has head hair 4-C usually closer to 6-C than to 5-C and siphon

hair 1-S usually inserted within distal portion of pecten (vexans group is

similar in head hair arrangement); Neomacleaya and Verrallina have head

hairs 4-C anterior to 5, 6-C and usually closer to 6-C than to 5-C, pecten

usually extends past middle of siphon, or with siphon hair 1-S inserted within

pecten; Edwardsaedes has saddle completely ringing segment X and precratal

hair tufts inserted into ventral margin (similar to genus Psorophora), head

hair 4-C well anterior to 5, 6-C, thorax with mesopleural and metapleural

tubercles well developed with stout points; and Levua by the shape of the comb

scales and the very small anal papillae. The larva of Aedes (Neomelaniconion)

lineatopennis (Ludlow) is very similar to Aedimorphus (especially vexans

group) and I cannot find any apparent subgeneric differences between them.

The aberrant Aedes (Stegomyia) vittatus (Bigot) bears a resemblance to Aedi-

morphus larvae but differs in having the antenna with only a very few tiny

spicules, head hair 4-C closer to 6-C than to 5-C, pecten extending beyond

basal 0.50 of siphon, and siphon hair 1-S inserted within apical portion of

pecten. The larvae of Aedimorphus differ from those of most of the other

Aedes subgenera in the position and development of the head hairs, develop-

ment of the antenna, possession of precratal tufts on segment X, number of

hairs in ventral brush on segment X, number of branches on hairs of ventral

brush, and development of the pecten and siphon.

MEDICAL IMPORTANCE. A number of species in the subgenus Aedi-

morphus have been implicated in the transmission of human and animal patho-

gens, primarily on the basis of recovery of disease agents from field-caught

adult mosquitoes. The bulk of these agents have been arboviruses, where iso-

lations have been reported from Africa, Europe, Asia and North America. In

most instances, however, comprehensive data on other biological factors which

determine vector status are not available.

Aedes vexans has been studied widely as a possible arbovirus vector.

This species has many attributes of an ideal vector -- it is widely distributed,

can become very abundant at times, often at the time virus activity is at a peak,

12 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

it feeds readily on man and domestic animals, and it has been found naturally

infected with various arboviruses.

Aedes vexans vexans has been found infected in nature with western

equine encephalitis (WEE) in Saskatchewan, Canada (McLintock et al. 1970:

448) and in the United States (Burroughs and Burroughs 1954: 33; Hayes et al.

1971: 184). It has also been found infected with eastern equine encephalitis

(EEE) in the United States (Wallis et al. 1960: 442). Chamberlain et al. (1954:

281-283), however, found that vexans vexans had a relatively low infection and

transmission rate for both these viruses, and consequently rated its vector

potential for WEE and EEE as "fair". It is unlikely that this species, which

is primarily a mammal feeder (Edman and Downe 1964: 156), would be much

involved in the endemic maintenance of these viruses, which normally infect

birds. As pointed out by Hayes et al. (1962: 119), vexans vexans may serve

aS an epidemic or epizootic vector during periods of abnormally high mosquito

and virus activity.

A more likely vector role for the species is with the California encepha-

litis (CE) group viruses which naturally infect mammals. Sudia et al. (1971:

591) list the following CE group virus isolations from vexans vexans in North

America: keystone, trivittatus, La Crosse, Jamestown Canyon, snowshoe

hare and California. Nothing is presently known concerning the relative sus-

ceptibility of vexans vexans to these agents.

Experimental transmission of fowl pox virus to chickens by vexans vex-

ans was demonstrated by Matheson et al. (1931: 218); and Hu (1931: 628) and

Yen (1938: 193) showed experimentally that this species was a fairly good car-

rier of Divofilaria immitis Leidy. Furthermore, Harinasuta et al. (1970: 242)

found vexans vexans and caecus positive with stage III larvae of Dirofilaria.

Aspock (1965: 767) and Mattingly (1969: 79) report that vexans vexans is involved

in the transmission of Tahyna virus in central Europe. This virus was also iso-

lated from vexans vexans in northern Italy by Balducci et al. (1968: 457).

Aedes vexans nipponii was suspected of being a vector of Japanese en-

cephalitis virus (JE) in Japan by LaCasse and Yamaguti (1948: 104) and Hodes

(1946; 358), working in Okinawa, experimentally transmitted this virus to mice

by the bite of vexans nipponii. Studies by Barnett (1962: 344; 406th Medical

General Laboratory Annual Progress Reports for 1949-51) in Japan, showed no

instances of natural infection of the species out of thousands tested, many of

which were collected during periods of peak virus activity. Shichijo et al. (1970:

91), however, report the isolation of 4 strains of Japanese encephalitis virus

and 3 strains of group A arbovirus from this species in the Nagasaki area of

Japan from 1965 to 1968. Sagiyama virus was isolated twice from vexans nip-

ponii, once in Japan (Scherer et al. 1962: 261) and once in Okinawa (Hurlbut

and Nibley 1964: 79).

This species of mosquito is not believed to be a vector of Wuchereria

bancroftt Cobbold since the larvae of this parasite died in the body cavity of the

mosquito according to Yamada (1927) and Newton et al. (1945: 256).

Although species of the subgenus are not known to be involved in yellow

fever transmission, Bauer (1928: 267), working in Africa, experimentally trans-

mitted this virus to monkeys by the bite of stokesi Evans (recorded as apico-

annulatus Edwards).

Little is known about the actual role of members of the subgenus as vec-

tors of other pathogens, and inferences must be based on reports of collections

of naturally infected adult mosquitoes. In Thailand, Batai virus was isolated

from a pool of 11 female vexans vexans and Wesselsbron virus from a pool of

mediolineatus (SEATO Medical Research Laboratory Annual Progress Report

for 1968: 63, 71).

Reinert: Aedes (Aedimorphus) in Southeast Asia 13

In Africa, Middelburg virus has been isolated from the following species

of Aedimorphus: albocephalus (Theobald) (Worth et al. 1961: 588), marshalli

(Theobald) (Paterson et al. 1964: 188), dentatus (Theobald), leesoni Edwards

and cumminsii (Theobald) (Robin et al. 1969: 113). The vector of this virus in

Senegal seems to be cumminsii and sheep apparently play a part in the main-

tenance of the virus (Robin et al. 1969: 117). The latter authors found the virus

primarily in cattle and rarely in man.

Smithburn and Haddow (1944: 265), while working in Africa, isolated

Semliki Forest virus from naturally infected members of the abnormalis Theo-

bald group. Sindbis, Middelburg and Shokwe viruses were isolated from cum-

minsii in the Republic of South Africa (Paterson et al. 1964: 188, McIntosh et

al. 1972: 158). In Mozambique, Worth and de Meillon (1960: 241) and McIntosh

et al. (1961: 193) report isolation of Semliki Forest virus from argenteopunc-

tatus (Theobald).

Spondweni virus was isolated from cumminsii (Theobald) in the Republic

of South Africa by McIntosh et al. (1961: 647). This virus was also isolated in

Mozambique by McIntosh et al. (1962: 685) from a pool of 42 mosquitoes iden-

tified as Aedes (Ochlerotatus) fryeri (Theobald) or Aedes (Aedimorphus) fowleri

(d'Emmerez de Charmoy). Worth et al. (1961: 588), in Republic of South Africa,

also isolated Spondweni virus from cumminsii, Wesselsbron virus from minutus

(Theobald) and an unidentified virus from marshalli. Rift Valley fever virus

was isolated from a pooled sample of 50 percent albocephalus and 50 percent

tarsalis (Newstead) in Africa by Smithburn et al. (1948: 117). Leeson (1958:

22, 26) also reports that albocephalus was found naturally infected with the

virus of Rift Valley fever and that farsalis is a vector of this virus. Rift Valley

fever virus was isolated by B.M. McIntosh (personal communication 1972) from

dentatus during an epizootic in cattle in Rhodesia in 1969. The African Bunyam-

wera virus was found in a pooled sample containing 5 subgenera of Aedes (8 spe-

cies of Aedimorphus) by Smithburn et al. (1946: 190).

Aslamkhan and Wolfe (1971: 31), working in Bangladesh (East Pakistan),

isolated unidentified filarial worms from pallidostriatus and found filarial larvae-

like structures in the head muscles of caecus that had been taken in a human biting

collection. Filaria larvae of Brugia malayi (Brug) were found in pallidostriatus

and pipersalatus in Ceylon by Carter (1948: 313).

BIOLOGY. Females of many species of Aedimorphus readily feed on

man and at times may be very serious pests. Other species prefer feeding on

cattle and other animals and only occasionally bite humans. See the BIOLOGY

section for the feeding habits of each species.

The typical immature habitats of Aedimorphus species in the Oriental

Region are temporary and semipermanent fresh water ground pools. Some

species have been collected from other habitats such as artificial containers,

rockholes, and similar habitats, but usually only when their normal breeding

places had dried up.

The eggs of Aedes vexans are drought-and cold-resistant and have re-

mained viable for over 4 years. The egg biology of other species in the sub-

genus is not known.

LIST OF SPECIES OCCURRING IN SOUTHEAST ASIA

Aedes (Aedimorphus) alboscutellatus (Theobald)

Aedes (Aedimorphus) caecus (Theobald)

Aedes (Aedimorphus) culicinus Edwards

Aedes (Aedimorphus) davidi Basio

Hq CO DO

bh ek ek ek pe pt

OPWNRrRODAAADO

2(1).

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Aedes (Aedimorphus) lowisii (Theobald)

Aedes (Aedimorphus) mediolineatus (Theobald)

Aedes (Aedimorphus) nigrostriatus (Barraud)

Aedes (Aedimorphus) orbitae Edwards

Aedes (Aedimorphus) pallidostriatus (Theobald)

Aedes (Aedimorphus) pampangensis (Ludlow)

Aedes (Aedimorphus) pipersalatus (Giles)

Aedes (Aedimorphus) punctifemoris (Ludlow)

Aedes (Aedimorphus) vexans vexans (Meigen)

Aedes (Aedimorphus) vexans nipponii (Theobald)

Aedes (Aedimorphus ?) wainwrighti Baisas

KEYS TO THE SPECIES OF AEDES (AEDIMORPHAUS)

IN SOUTHEAST ASIA*

KEY TO THE ADULTS

Head with only broad scales on vertex; antepronotum with broad silvery

Head with narrow scales mesally on vertex; antepronotum with narrow

curved white or golden scales, or if broad scales present these

are NOU ChVeCrV oe ae a eS

Vertex of head without erect forked scales; postpronotum without

scales; tarsi with white bands. ............ wainwrighti

3(1). Tarsi II, Ill with basal white bands or dorsobasal white spots. ...

Tarsi I, III without basal white bands or spots (some species may

have lateral longitudinal white stripes). .............

4(3). Wing with dorsal veins brown scaled or with only a small spot of

white scales at base of costa; femora I, II with anterior brown. .

Wing with dorsal veins with white and brown scales intermixed or

with a number of white scales on posterior margin and base of

Vertex of head with erect forked scales; postpronotum with a few broad

silvery StCales; tarsi dark, 2.2/0.3 ecw a Boe punctifemoris

5(4).

costa; femoral, II with a large number of white scales intermixed

With Drown Ges s ea a 7

Scutellum with broad and narrow white scales on each lobe; tarsus III

with a narrow basal white band on tarsomere 5. ..... caecus

Scutellum with broad silvery scales on each lobe; tarsus III with tarso-

mere. 5 pale scaled: ee es Re BO EE 6

*Aedes stenoetrus and taeniorhynchoides are not found in Southeast Asia but

are included in order to prevent them from being confused with closely re-

lated species.

Reinert: Aedes (Aedimorphus) in Southeast Asia 5

6(5). Maxillary palpus brown scaled; tarsilI, II each with tarsomere 5 pale

SCALE CGR OO PS ROS AR Ee aE OOTP A oa 6 lowisit

Maxillary palpus brown with white scales; tarsilI, I each with tarso -

M6Pe 5 rown SCalede ia me PA OOO eo ae is orbitae

7(4). Tibiae I-III each with a subapical pale scaled band; scutellum* with

brown ‘scalesvon each lobe. 2. Se Fe aaa davidi

Tibiae I-III each without a subapical pale scaled band; scutellum with

white SCaLeS On 2ach fone es tee Re OR oe oe ak 8

8(7). Wing brown with numerous white scales (30% or more of scales white)

intermixed on all dorsal veins, including distal 0. 85 of radius

2+3 and tertiary fringe scales along posterior margin of female

Wing without numerous white scales intermixed on all dorsal veins,

distal 0.85 of radius 2+3 and scales along posterior margin of

wine always browns 38s eee ee EP eR PP ES 10

9(8). Scutellum with only narrow curved white scales on midlobe; postpro-

notum completely covered with narrow curved pale scales.

taeniorhynchoides

Scutellum with broad white scales and occasionally a very few narrow

curved ones on midlobe; postpronotum with narrow curved reddish-

brown scales dorsally and white ones below. .... pipersalatus

10(8). Proboscis completely covered with dark brownish-black scales.

stenoetrus

Proboscis brown with numerous white scales ventrally....... 11

11(10). Abdomen with terga III, IV each with an incomplete apical median

longitudinal white stripe which may or may not connect with

basal bands; lower subspiracular scale patch connects, or nearly

so, with postspiracular scale patch. ....... vexans nipponit

Abdomen with terga III, IV without apical median longitudinal white

stripe; lower subspiracular scale patch does not connect with

postspiracular scale patch. ee er PEAS vexans vexans

12(3). Propleuron with narrow curved scales; scutum with 2-3 distinct longi-

cudimgl Stripes Or pale Scales,” 2. Se ee ee ee 13

Propleuron with broad scales; scutum without stripes of pale scales.

13(12). Anterior margin of wing pale scaled; femora II, III each brown with an

anterior median longitudinal white stripe. ..... pallidostriatus

Anterior margin of wing brown scaled; femora II, III each without an

anterior longitudinal white stripe. ......... mediolineatus

14(12). Scutal integument pale with a pair of dark stripes on dorsocentral

arenas: tarsr yellow eealeao "te. a a nigrostriatus

Scutal integument uniformly dark; tarsi brown scaled. ....... 15

*But see Taxonomic Discussion page 34.

15(14).

16(15).

2(1).

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Scutellum with narrow curved white scales on each lobe; subspiracu-

lar area with 2 patches of broad white scales. ..... culicinus

Scutellum with broad silvery-white scales on each lobe; subspiracu-

lar area with 1 patch of moderately broad white scales or with a

patch of short fine hairs and without scales. .......... 16

Abdomen with dorsobasal white bands on terga III, IV; subspiracular

area with a patch of short fine hairs. ....... alboscutellatus

Abdomen without dorsobasal pale bands on terga III, IV; subspiracu-

lar area with a patch of white scales. ....... pampangensis

KEY TO THE PUPAE*

Patch of spicules between bases of metanotal hairs 10-C and abdominal

ied the soc ae a ee Oe Ser GS Bodacious. 2

Areas between bases of metanotal hairs 10-C and abdominal hairs 1-I

ROS LY OEE CATS a. Pg ee PTE sg gs was Se eR we Rs 3

Hair 10-C with 7-11 branches; hair 6-I single; hair 1-II with 20-32

rR ruta ars Bir al hg ca Ak gg ua ge Nig Tg i Caecus

Hair 10-C with 17-33 branches; hair 6-I double or triple; hair 1-II with

Ba GB. DPANCHRS gig -tiwary oui mataeee ENE SOR a GOONIES Lele orbitae

Ventral posterior margin of II with spicules mesally. . vexans vexans

vexans nipponii

Ventral posterior margin of II without spicules mesally.......

Hair 7-C with 8-11 branches; hair 6-VI with 6-8 branches.

pampangensis

Hair 7-C with 2-6 branches; hair 6-VI single to 4 branched. ....

Hair t-[L. with 3=T Panchos: o.iesd a hoy bh wares 6 alboscutellatus

Hair -l-1L-with-9 #40. Oranches: hie Wk Si oso eed CeowweardA «.. 6

Hair 10«GC. with 6-0 Pranches coche dee dares Ow Rs eas 7

Hair 106 with 10=33 DPAnCHeS oii: ei Pees oR me Swe te aie 8

Halt 2-26 Witt: 2eO OP AME ROS és Sinimdiiaiy tra send pate ee aera culicinus

Wait Al With 7-18. Branches. oe ices kessac abu. ac S Bela eae pipersalatus

Haiv’ L=1 with: 1.7330. Dranchesiisees 6 wk Se ew Ao cwck < mediolineatus

Hair 1-I with 30-46 branches. ............ee-. pallidostriatus

KEY TO THE FOURTH STAGE LARVAE*

Siphon with patches of spicules, these may be restricted to apex. . 2

Sy LivrMenta AP SEIICHAT RCO oe ys Me hedge ba aie pam dy mican tones ee 3

*The immature stages of davidi, lowisti, nigrostriatus, punctifemoris and

wainwrighti are not known.

Reinert: Aedes (Aedimorphus)in Southeast Asia ri

2(1). Siphon with lateral patch of spicules near middle; head hair 6-C with

6-9 Branches ys BR re shay eee Se toe farsa Tye se orbitae

Siphon without lateral patch but usually with a dorsal patch of spicules;

head hair 6-C with 4-5 branches. ............. caecus

3(1). Mesothoracic hair 5-M double or triple; siphon long, index 7. 50-9. 00.

pampangensis

Mesothoracic hair 5-M single; siphon short to moderately long, index

Cd fer kee be TE OR OY Babs Baa SOE eee te

4(3). Comb scales short, without a long median spine but with lateral

CMI CleS. ech ae wr fe) “Rae ede SN OTE TT Bere 5)

Comb scales with a long stout pointed median spine and short denticles

along lateral maregine of bases... PRE ere 6

5(4). Siphon index 5.00-5. 56; abdominal hair 10-I double. . . pipersalatus

Siphon index 3.00-4.08; abdominal hair 10-I single. . alboscutellatus

6(4). Head hair 6-C single to double; head hair 8-C single. ....... 7

Head hair 6-C triple to 6 branched; head hair 8-C double to 8

branched ys «<i hw ety IEEE weed Botnet ee Sie 8

7(6). Head with frontaclypeus granulose........... vexans vexans

Head with frontoclypeus without granules. ...... vexans nipponit

8(6). Prothoracic hair 7-P double; metathoracic hair 7-T with 5-6 branches;

siphon index: 2,38+4. 80. Gin: Sei MPAs BRE Ge eeu culicinus

Prothoracic hair 7-P triple; metathoracic hair 7-T with 7-14 branches;

sipnon indet: S, 99<%.9 8. do ha ei ee mediolineatus

pallidostriatus

DESCRIPTIONS OF THE SPECIES OCCURRING

IN SOUTHEAST ASIA

AEDES (AEDIMOR PHUS) ALBOSCUTELLATUS (THEOBALD)

(Figs. 1, 16, 17, 18, 33, 34, 49, 59)

Lepidotomyia alboscutellata Theobald 1905, Ann. Mus. nat. Hung. 3: 80 (9*);

Brunetti 1907, Rec. Indian Mus. 1: 339.

Lepidotomyia Alboscutellata Theobald, Leicester 1908, Cul. Malaya, p. 132

(o', 9).

Culex argentinotus Banks 1909, Philipp. J. Sci. 4: 547 (c, 9).

Reedomyia alboscutella Theobald, Theobald 1907, Monogr. Cul. 4: 261 (9*);

Theobald 1910, Monogr. Cul. 5: 257.

Reedomyia alboscutellata Theob., Brunetti 1912, Rec. Indian Mus. 4: 487.

Ochlerotatus alboscutellatus Theob., Brunetti 1920, Rec. Indian Mus. 17: 139;

Senior-White 1923, Cat. Indian Insects, Cul., p. 76.

Aédes Omurensis Yamada 1921, Annot. zool. jap. 10: 73 (o* 9).

Aedes (Ecculex) alboscutellatus Theobald, Edwards 1922b, Indian J. med.

Res. 10: 467.

Aédes (Aédimorphus) alboscutellatus (Theo.), Edwards 1924, Bull. ent. Res.

14; 372 (9); Barraud 1934, Fauna Brit. India, Diptera 5: 250 (o%*, 9).

18 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Aédes (Aédimorphus) alboscutellata Theobald, Dyar and Shannon 1925,

Insecutor Inscit. menstr. 13: 76.

Aedes (Aedimorphus) alboscutellatus (Theobald), Barraud 1928, Indian J. med.

Res. 15: 659 (o’*, 9); Bohart 1945, Navmed. 580, p. 62; Hsiao and

Bohart 1946, Navmed. 1095, p. 22; Penn 1949, nat. Hist. Misc. 40:

1 (P*, L*); La Casse and Yamaguti 1950, Mosquito Fauna Japan and

Korea, p. 130 (o*, 9*); Knight and Hull 1953, Pacif. Insects 7: 457

(o*, 9, L*); Stone et al. 1959, Thomas Say Found. 6: 190; Belkin 1962,

Mosquitoes S. Pacif., p. 425 (o’*, 9, P*, L*); Assam and Bonne-

Wepster 1964, Zool. Bijdr. 6: 96 (oc, 9, L*); Mohrig 1967, Angew,

Parasit. 8: 80 (9*); Pao and Knight 1970, Mosquito Syst. Newsletter

2: 105 (L*); Kurihara 1963, Jap. J. sanit. Zool. 14: 196 (A*).

Aédes (Aédimorphus ) alboscutellatus Theobald, Edwards 1932, Genera Insec.,

Fasc. 194: 167.

Aedes (Aedimorphus) alboscutellatus Theobald, Bonne-Wepster 1954, Doc.

med. Geogr. Trop. 6: 236 (oc, 2, L*).

FEMALE (Fig. 1). Head. Antenna dark brown, approximately 1.14

length of proboscis, pedicel pale with a few small brown scales and a patch of

short fine brown hairs mesally, flagellomere 1 with basal 0. 50 pale and with

a few small brown scales; clypeus dark, bare; maxillary palpus brown scaled,

approximately 0.17 length of proboscis; proboscis brown scaled with a ventral

pale area from near base to apical 0.25, approximately 1.12 length of femur I;

vertex with dosum covered with narrow decumbent scales arranged in an ante-

romedian diamond-shaped brown group and the remainder whitish; lateral sur-

face covered with broad pale scales, an anterodorsal dark patch and a dusky

area anterior to antepronotum; numerous dark brown erect forked scales on

occiput and vertex extending anteriorly to ocular line. Thovax. Scutal integu-

ment brown; scutum covered with narrow curved reddish-black scales, narrow

curved white scales forming distinct spots on scutal fossal areas (lateral and

posterior) and supra-alar area at base of wing, indistinct spots on anterior

promontory area, anterior scutal fossal area and at scutal angle, afew similar

scales scattered on area mesally to acrostichal setae and along lateral margins

of prescutellar space; scutellum with a patch of long, broad overlapping silvery

scales on each lobe; median anterior promontory, acrostichal, dorsocentral

(anterior and posterior), scutal fossal (anterior, 2-3 lateral and 1-2 posterior),

supra-alar, several posterior medial scutal, 1 postalar callar and scutellar

(lateral and median) bristles reddish-black and well developed, others absent;

pleural integument brown; antepronotum with narrow curved and a few broad

white scales, several moderately long brown bristles; postpronotum with nar-

row dark brown scales dorsally and a fewwhite ones posteroventrally, 4-5

dark posterior bristles; propleuron with a patch of broad silvery-white scales,

several dark bristles; postspiracular area with 6-7 light brown bristles; sub-

spiracular area with a patch of short fine light brown hairs; mesepisternum with

an upper and a posterior patch of broad silvery-white scales, several upper and

posterior brown bristles, lower ones shorter; prealar knob with several dark

brown bristles; paratergite with a few dusky-white scales (these scales usually

rubbed off); mesepimeron with a patch of broad silvery-white scales and several

brown bristles on upper area; other pleural areas bare. Legs (Fig. 16). Coxae

I-III each with several brown bristles, I with anterior surface covered with

broad pale brownish scales and a small dorsal patch of silvery-white ones, II

with an anterior patch of broad silvery-white scales, III with a small antero-

ventral patch of silvery-white scales; trochanters I-III with a few broad white

Reinert: Aedes (Aedimorphus) in Southeast Asia 19

scales; femora I-III brown, I with a few lateroapical white scales, II, III each

with a dorsoapical white spot, III with an anteroventral longitudinal pale stripe,

wide at base and tapering at apex, I-III each with posterior surface with a

longitudinal pale stripe, wide at base and tapering to apex, stripe dorsal on I

and ventral on II, III; tibiae I, II brown, each with a dorsoapical white scale

patch and a posteroventral longitudinal pale stripe, III brown with .an apical

white band; tarsi I-III brown; posttarsi I-III each with 2 ungues, I, II equal,

each bearing a tooth, III equal, simple. Wing. Dorsal veins covered with

moderately broad brown scales; costa with a patch of white scales at base;

ventral veins brown scaled; alula with narrow brown scales along fringe; 2 re-

migial bristles. Halter. Pedicel pale, capitellum brown scaled. Abdomen.

Tergum I brown with a rectangular patch of white scales on laterotergite;

terga brown, III-VII each with a few dorsobasal pale scales (pale scales

occasionally forming narrow indistinct bands), a few dorsomedian pale scales

on VI in some specimens, II-VII each with a large laterobasal white spot;

sterna white each with apical 0.25 brown scaled, brown apical bands broader

on posterior sterna; terga and sterna with numerous golden bristles, mostly

along posterior margins. Genitalia (Fig. 18). Tergum VIII index 1.02-1.04;

sternum VIII index 0. 88-0.92; tergum IX bilobed with 3-8 bristles on each

lobe, index 1. 18-1. 23; insula tongue-like, covered with minute setae and with

4-5 small tuberculi on apical 0. 20; lower vaginal lip narrow, moderately pig-

mented, covered with minute setae; upper vaginal lip narrow, heavily pig-

mented, covered with minute setae; upper vaginal sclerite large, moderate to

heavily pigmented, triangular shaped; postgenital lobe moderately long, apex

with a moderately deep median indentation, 5-8 bristles on each side of mid-

line, covered with minute setae, dorsal PGL index 0. 89-1.24, ventral PGL

index 2. 16-2. 50; cercus long, 0.80-0.90 extended from segment VII, narrow,

apex sharply rounded, numerous bristles on dorsal and lateral surfaces, index

3.35-4.25, cercus/dorsal PGL index 3.90-5.12; 3 spherical, pigmented semi-

nal capsules, 1 large and 2 tiny rudimentary ones.

MALE (Fig. 1). Similar to female in general habitus. Head. Maxil-

lary palpus brown, longer than proboscis by length of apical segment; proboscis

dark brown scaled with a small ventral pale spot near middle; vertex without

anterodorsal dark patch on lateral surface. Thorax. Antepronotum with a few

broad pale scales; postpronotum with fewer scales. Legs (Fig. 17). Post-

tarsi I-III each with 2 ungues, I, IT with ungues unequal each bearing a tooth,

III equal, simple. Abdomen. Tergum I with a lateral band of white scales on

laterotergite; terga III-VIII each with a broad basal pale band; sternum VIII

white scaled. Genitalia (Figs. 33, 34). Tergum IX strongly bilobed with 4-6

bristles on each lobe, entire surface covered with minute spicules; gonocoxite

long and moderately broad, dorsal surface with short fine bristles forming an

elongate patch along tergomesal margin from apex to base and extending over

basal 0.25, long stout bristles along outer lateral margin from base to apex

and on apical 0.35 of ventral surface, scattered short to moderately long

bristles mesally on basal 0.65 of ventral surface, scattered scales on lateral

and ventral surfaces; gonostylus with pedicel narrow and short, distal 0. 50

greatly expanded with a lateroapical horn-like flap bearing a number of moder-

ately long fine hairs, mesal margin of expanded portion with a moderately long

flattened, pigmented gonostylar claw near base and 3 small and 1 medium-

sized accessory claws distally, 3-4 moderately long fine hairs along apical

margin and 9-16 short fine hairs scattered over tergal surface of expanded

area; basal mesal lobe short and rounded apically, distal 0.40 with 2-3 short

and 1 moderately long bristles, a long narrow lightly pigmented filament

20 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

extending from apex to near base of gonostylus and attached to mesal mem-

brane of gonocoxite, entire surface of basal portion covered with short hair-

like spicules; proctiger short, paraproct with a subapical thumb-like process,

cercal setae absent; phallosome with aedeagus of type I with 2 lateral plates

connected basally, each plate with 5-6 short, blunt, lateral teeth on distal 0.55

and covered with a dorsal flap, paramere long, approximately 0. 83 length of

lateral plate; sternum IX large, entire surface covered with minute spicules,

3-5 bristles near center.

PUPA (Fig. 49). Chaetotaxy as figured and recorded in Table 1.

Cephalothorax. Hair 5-C with 4-6 branches, 7-C with 3-4 branches; 8-C with

0-9 branches. Respivatory trumpet. Moderately pigmented; index 3. 50-4. 50,

average 3.87. Metanotum. Hair 10-C with 3-5 branches; 12-C with 4-7

branches. Abdomen. Hair 5-I with 6-11 branches; 1-II with 3-7 branches;

4-II with 2-5 branches; 1-III with 6-10 branches; 6-VI with 2-4 branches;

1-VII with 4-6 branches; 6-VII with 4-10 branches; 9-VII with 2-5 branches;

11-VII double or triple. Paddle. Ovoid; with very minute serrations along

basal 0.50 of outer margin; tiny spicules along apical 0. 50 of outer and apical

0.20 of inner margins; midrib does not reach apex; hair 1-P short, double or

triple; index 1.09-1.30, average 1.21.

LARVA (Fig. 59). Chaetotaxy as figured. Head. Hairs 1, 3-C single;

4-C with 6-10(8) branches; 5-C with 3-7(3) branches; 6-C with 2-5(3) branches;

7-C with 4-8(6) branches; 8-C with 2-4(4) branches; 9, 11-C with 3-6(4)

branches; 10-C double or triple (3); 12-C with 3-5(5) branches; 13-C double or

triple (2); 14-C single or double (1); 15-C with 3-7(4) branches; basal maxil-

lary hair single; mental plate with 32-36(34) teeth. Antenna. Lightly pig-

mented; numerous spicules on basal 0.25 and a few scattered over remainder

of shaft; hair 1-A with 5-7(5) branches, inserted at 0.40-0.47 from base; 2-A

long; 3-A approximately 0.34 length of 2-A. Thorax. Hair 0-P with 6-11(10)

branches; 1, 5, 6, 12-P single; 2-P with 2-4(3) branches; 3-P with 3-4(4)

branches; 4-P with 3-7(4) branches; 7, 11-P with 2-4(2) branches; 8, 10-P

single or double (1); 9-P double or triple (3); 14-P single to triple (2); 1-M

with 3-7(4) branches; 2-M with 2-4 (2) branches; 3-M with 3-5(3) branches;

4-M with 4-9(5) branches; 5, 7, 10, 12-M single; 6-M with 3-5(4) branches;

8-M with 5-7(6) branches; 9-M with 6-7(6) branches; 11-M single or double

(1); 13-M with 5-10(7) branches; 14-M with 4-10(8) branches; 3-T with 7-12

(11) branches; 4-T with 3-4(4) branches; 6-T with 2-5(4) branches; 7-T with

5-7(6) branches; 8-T with 5-12(7) branches; 9-T with 4-6(5) branches; 11-T

single or double (2); 12-T single to triple (2); 13-T with 9-19(14) branches.

Abdomen. Hairs 1, 2-VIII on common basal plate; 0, 14-VIII single; 1-VII

with 6-11(6) branches; 2-VIII with 2-4(3) branches; 3-VIII with 6-16(9)

branches; 4-VIII single or double (2); 5-VIII with 5-9(7) branches; 6-III-VI

short; comb with 17-25(19) scales arranged in 3 irregular rows, scales each

short and blunt with stout denticles on margins and apex; 1-X double or triple

(2); 2-X with 7-12(11) branches; 3-X single; ventral brush varies from 9 hairs

on grid and 3 precratal ones to 12 hairs on grid and 2 precratal ones, usually

with 9 hairs on grid and 3 precratal ones; saddle moderately pigmented with

minute ridges, incompletely rings segment, with a few spicules along posteri-

or margin, acus present; 4 anal papillae, long, each with a broad base and

tapering to a pointed apex. Siphon. Moderately pigmented with minute ridges

over entire surface; acus present; index 3.00-4.08; pecten with 15-20(17)

teeth, apical 1-3 teeth smooth and wider spaced than remainder which have a

slender apical attenuated filament with 1-2 basal denticles; hair 1-S with 4-6

(5) branches, inserted at 0. 59-0.67 from base.

Reinert: Aedes (Aedimorphus) in Southeast Asia 21

TYPE DATA. Lepidotomyia alboscutellata Theobald, holotype female,

Simbang, Huon Gulf, New Guinea, AUSTRALIA, July 1898, Biro, in Magyar

Nemzeti Museum, Budapest, Hungary; Culex argentinotus Banks, syntypes

male and female, Pinagsalaan Well, Taytay, Rizal, Luzon, PHILIPPINES,

13-16 May 1909, C. S. Banks, types nonexistent (Stone et al. 1959: 190);

Aedes O6murensis Yamada, 4 female and 2 male syntypes, 5 female and 1 male

paratypes, Omura, Kyushu, JAPAN, syntype No. 35, in Medical Zoology

Laboratory, Institute for Infectious Diseases, University of Tokyo, Tokyo,

Japan.

DISTRIBUTION. Specimens examined--241 males, 291 females, 170

pupae, 93 larvae and 172 individual rearings (137 pupal, 35 larval) from the

following locations: : |

AUSTRALIA. N.E. New Guinea, Milne Bay; Papua, Trobriand Islands,

Kiriwina.

BISMARCK ARCHIPELAGO. Emirau Island.

INDIA. Assam, Golaghat.

INDONESIA. Ceram, Lisabata, Wahari; Java, Rawaloh, Tjibodas,

Mt. Gede, Tjilatjap; New Guinea, Tanah Merah, Upper Digul River; Hollandia;

Sumatra, Bengkoelem, Blang Kedjeren, Dermaja, Dramajoe, Benkoclen, Kotta

Tjane, Mocra Tebo.

MALAYSIA. Pahang, Cameron Highlands, Kg. Sertik, Bentong; Sava-

wak, Kuching; Sabah, Beaufort, Tenom; Selangor, Kuala Lumpur, Ulu Gombak;

Segambut.

PHILIPPINES. Batangas, Pto. Tomas; Leyte, Jinamoc Is.; Luzon, La

Union, Anastacio, Bacnotan; Mindoro, Caminawit, Kabakan, San Jose; Palawan,

Puerta Princesa; Samar, San Antonio, Calaccad, Tawi Tawi Is.

SOLOMON ISLANDS. Bunana; New George Is., Munda; Empress,

Augusta Bay, Bougainville, Guadalcanal.

SOUTH VIETNAM. Binh Dinh, Binh Dinh, An Khe; Pleiku, Pleiku.

THAILAND. Khon Kaen, Ban Maung Kao, Phu Wiang; Krabi, Ban

Phru Toei; Lampang, Ban Na Kiang; Maha Sarakham, Muang; Nakhon Ratchasi-

ma, Pak Chong, Pak Thong Chai; Nan, Ban Pang Chom Phu, Ban Pang Mon,

Ban Ta Loc, Pha Dang Khawi; Narathiwat; Songkhla, Haad Yai; Ubon Ratchath-

ani, Chong Med; Udon Thani; Ban Pa Goi; Talum.

Other distribution.

ADMIRALTY ISLANDS (Belkin 1962: 427, Iyengar 1955: 42, 1960: 67).

AUSTRALIA. New Guinea, Friedrick-Wilhelmshafen, Simbang, Huon

Gulf (Theobald 1905b: 81); Madang (Cooling 1924: 27); Northern Territory,

Daly River, Doctor's Gully (Edwards 1924: 372); Queensland (Edwards 1922b:

467), Cairns (Doherty et al. 1963: 22), Innisfail (Standfast and Barrow 1969:

39).

BURMA (Edwards 1922b: 467); Bhamo (Barraud 1934: 251).

CEYLON. Matate (Senior-White 1923: 76).

INDIA. Bombay Deccan, Tavargatti; Upper Assam Valley (Barraud

1928: 659); Belgaum; Bengal, Sukna, Darjeeling; Bihar, Pusa; Assam, Sibsa-

gar (Barraud 1934: 251); Poona City (Rao and Rajagopalan 1957: 10).

INDONESIA. Borneo (Brug 1926: 529); Sumatra, Atchin; Benkoelen,

Air Prioeka; Djambi; Lampong, Lampong (Brug and Edwards 1931: 258); Su-

matra, Moeara Tebo (Haga 1924: 830); New Guinea, Inanwatan, Hollandia,

P, Pam, Sokori Plain (Assem and Bonne-Wepster 1964: 98); Morotai (Bonne-

Wepster 1954: 237).

JAPAN. Omura, Kiushu (Yamada 1921: 76); Misawa AB, Yokota AB

(Reisen et al. 1971: Tables 23 and 24),

22 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

MALAYSIA. Savawak, Kampong Pangkalan Kuap (Macdonald et al.

1965: 338).

NEW BRITAIN (Taylor 1934: 20).

PHILIPPINES. Zamboanga, Pettit Barracks; Mindanao (Dyar and

Shannon 1925: 77); Rizal, Taytay, Pinagsalaan Well (Banks 1909: 548); Laguna, -

Los Banos (Bohart 1945: 63); Palawan, Bacungan; Samar, Osmena; Zamboanga,

Mercedes, Zamboanga City (Knight and Hull 1953: 459); Jinamoc Island; Min-

dovo, Calapan (Basio 1971: 11); Clark AB (Dowell, Libay and Baisas 1965: 14).

SOLOMON ISLANDS. Bougainville (Belkin 1962: 427).

THAILAND. Nakhon Phanom (Parrish 1968b: 2); U-Tapao (Reisen et al. |

1971: Tables 4 and 8); Kanchanaburi, Sangkla-buri, Lai-nam, Ni-Thae (Harina-

suta et al. 1970: 241).

SOUTH KOREA. Kimpo Air Base, Kusan Air Base, Kwang-Ju Air Base,

Osan (Reisen et al. 1971: Tables 26 and 27).

SOUTH VIETNAM. Phan Rang (Parrish 1968a: 3); Nha Trang (Parrish

1969: 554); Chu Lai, Da Nang (Grothaus et al. 1971: 20).

YORK ISLAND (Taylor 1934: 20).

TAXONOMIC DISCUSSION. Aedes alboscutellatus is very similar to

the following Oriental and Pacific Islands species in the adult habitus: jamesi,

lowisti, oakleyi, senyavinensis, culicinus and trukensis. It can be easily

separated from the first 2 species by the dark tarsi and from the latter 3

species by the broad silvery scales on the scutellum. Aedes alboscutellatus

differs from oakleyi,by the following characters: femora II-III each with a

dorsoapical silvery spot; abdominal terga with very narrow basal white bands;

and head with an anterodorsal brown spot on lateral surface while oakleyi has:

femora II, III each with a small lateroapical white spot; abdominal terga with

broad basal white bands; and head entirely pale scaled.

In addition to the above features, alboscutellatus also has the sub-

spiracular area with only short fine hairs and the postspiracular area without

scales. The female possesses 1 large and 2 rudimentary seminal capsules

and shares this feature with culicinus, jamesi, lowisii, oakleyi, senyavinen-

sis and trukensis. The illustration of the female genitalia of alboscutellatus by

Hara (1957: 91) shows 1 large and 2 slightly smaller seminal capsules and

therefore must belong to some other species.

The male genitalia of alboscutellatus are similar to those of culicinus,

jamesi, lowisti, oakleyi and senyavinensis. The following character combina-

tion can be used in separating alboscutellatus from these related species: gono-

stylus with 9-16 short fine hairs scattered over tergal surface of expanded

area; gonostylar claw with apex pointed; 3-4 accessory claws; and basal mesal

lobe with 3-4 bristles; culicinus possesses: gonostylus with 17-24 short fine

hairs on tergal surface of expanded area; no accessory claws; and basal mesal

lobe with 6-10 bristles; jamesi possesses: gonostylus with 20-24 short fine

hairs on tergal surface of expanded area; 3 accessory claws; and basal mesal

lobe with 3-4 bristles; lowisii possesses: gonostylus with 3-8 short fine hairs

on tergal surface of expanded area; 4-5 accessory claws; and basal mesal lobe

with 4-5 bristles; oakleyi possesses: gonostylus with 14-16 short fine hairs

on tergal surface of expanded area; gonostylar claw with apex blunt; 2-3 ac-

cessory claws; and basal mesal lobe with 4 bristles; and senyavinensis posses-

ses: gonostylus with 25-32 short fine hairs on tergal surface of expanded area;

2-3 accessory claws; and basal mesal lobe with 8-9 bristles.

The larva of alboscutellatus is similar to pipersalatus and can be

recognized by a difference in the shape of the comb scales and the siphon index

which is 3. 00-4. 08 and in pipersalatus it is 5.00-5. 56.

Reinert: Aedes (Aedimorphus) in Southeast Asia 23

BIOLOGY. The immatures are usually found in small, partially

shaded, flood pools in secondary scrub vegetation and the adults are usually

taken biting cattle and man in the shade of forests during the day. Immatures

in Thailand have been collected from small and large flood pools, small and

large stream pools, small rock pool; usually from turbid fresh water but also

several times from clear fresh water; water with floating and submerged dead

leaves and scarce floating aquatic vegetation; usually in partial shade but oc-

casionally in unshaded areas; usually in secondary scrub vegetation in moun-

tains, hills or plateau but also in secondary scrub in plains, bamboo groves

in hills, deciduous forest in valleys, and rain forest in hills; and at an altitude

of 250 to 1480 feet (most often at 820 to 1310 feet). Larvae were collected

in association with the following species of mosquitoes: Aedes caecus, culi-

cinus, ferinus, imprimens, mediolineatus, uncus, vexans vexans, vigilax,

vitiatus; Anopheles balabacensis, bengalensis, culicifacies, kochi, maculatus;

Culex annulus, bailyi, fuscocephalus, infantulus, malayi, nigropunctatus, pal-

lidothorax, pseudovishnui, raptor, scanloni; and Uranotaenia macfarlanei.

In the Philippines, immatures have been collected from water in ground

pools, foxholes and sunlit roadside ponds; adults have been taken biting man

and cattle and resting in foxholes. Adults have also been collected feeding on

cattle, buffaloes and man in forests in Thailand, in the jungles of Ceram in the

afternoon and in Malaysia during the evening. In Java, adults have been taken

at an altitude of 4,000 feet and in New Guinea in light traps. Immatures have

been collected from jungle pools in Malaysia, from a road rut completely

covered with grass and from a ground pool in New Guinea and from clear water

in a primary forest in Sumatra.

Immatures have been found in flooded swamp areas, woodland pools,

rockholes and potholes in a streambed while adults were collected biting during

the daytime in forested areas of the Solomon Islands (Belkin 1962: 426). In

Malaysia, Macdonald (1957: 21) states this species bites man in inland forests

and has been taken at treetop levels. Macdonald and Traub (1960: 100) also

record the larvae in ground pools in this country. Steffan (1966: 212) in New

Guinea found larvae in shallow pools at the edge of the jungle, open areas,

wheel ruts and had taken the adults in carabao baited traps. In India, larvae

were collected from jungle pools (Barraud 1928: 659, 1934: 251) and adults

from indoor shelters (Rao and Rajagopalan 1957: 10). Penn (1948: 245) made

a larval collection in the Philippines from a permanent ditch which contained

clear, stagnant water with a pH of 6.5. Females have been taken at a light

in Malaysia (Edwards 1928: 53). Grothans et al. (1971: 20) in Vietnam found

larvae in ground pools, swamp margins and long standing stagnant ditches with

ample shade while adults were collected from indoor resting stations and biting

man during the daytime in densely shaded woods.

AEDES (AEDIMORPHUS) CAECUS (THEOBALD)

(Figs. 2, 16, 17, 19, 35, 50, 60, 61)

Culex caecus Theobald 1901, Monogr. Cul. 1; 413 (2*); Giles 1902, Handb.,

2nd. Ed., p. 415 (9); Blanchard 1905, Moust., p. 305 (2); Theobald

1905, Genera Insec., Fasc. 26: 26; Brunetti 1907, Rec. Indian Mus.

1: 343.

Pecomyia caeca Theobald, Theobald 1907, Monogr. Cul. 4: 268 (o', 9*); Theo-

bald 1910, Monogr. Cul. 5: 260; Brunetti 1912, Rec. Indian Mus. 4:

459.

24 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Geitonomyia Caecus (Theobald), Leicester 1908, Cul. Malaya, p. 134.

Ochlerotatus caecus Theob., Brunetti 1920, Rec. Indian Mus. 17: 137.

Aedes (Ecculex) caecus Theo., Edwards 1922b, Indian J. med. Res. 10: 467.

Aédes (Aédimorphus) caecus Theo., Brug 1924, Bull. ent. Res. 14: 436 (L*);

Borel 1930, Coll. Soc. Path. exot. Monogr. 3: 271 (o*, 9*, L*); Ed-

wards 1932, Genera Insec., Fasc. 194: 170; Barraud 1934, Fauna Brit.

India, Diptera 5: 257 (o*, 9, L*).

Aedes (Aedimorphus) caecus (Theo.), Barraud 1928, Indian J. med. Res.

15: 663 (o*, 9); Stone et al. 1959, Thomas Say Found. 6: 191.

Aedes (Aedimorphus) caecus Theobald, Bonne-Wepster 1954, Doc. med.

Geogr. Trop. 6: 237 (oc, 9*, L*).

FEMALE (Fig. 2). Head. Antennadark brown, approximately 1. 10 length

of proboscis, pedicel pale with a few small pale scales and a patch of short

fine black hairs mesally, flagellomere 1 pale with a few small brown scales;

clypeus dark brown, bare; maxillary palpus brown scaled, approximately 0.18

length of proboscis; proboscis brown scaled with pale scales on lateral and

ventral surfaces from near base to distal 0.25, pale area somewhat broader

basally, approximately 1.04 length of femur I; vertex with dorsum covered

with narrow decumbent scales arranged in an anterior brown patch and white

scales laterally and posteriorly; lateral surface covered with broad pale

scales and an anterodorsal dark patch; numerous dark brown erect forked

scales on occiput and vertex extending anteriorly to ocular line. Thorax. Scu-

tal integument dark brownish-black; scutum covered with narrow curved reddish-

black scales, narrow curved white scale patches on median anterior promon-

tory area, scutal fossal areas (extending from anterior area along margin and

onto lateral area), along scutal ridge from scutal angle posteromesally onto

dorsocentral setal line, supra-alar area from base of wing posteromesally to

dorsocentral setal line, posterior medial scutal area and along margins of pre-

scutellar space; scutellum with a patch of narrow curved white scales and a few

broad white ones on each lobe, broad scales on lateral lobes longer than on

median lobe; median anterior promontory, acrostichal, dorsocentral (anterior

and posterior), scutal fossal (anterior, 2-3 lateral and 1 posterior), supra-

alar, several posterior medial scutal, 1 postalar callar and scutellar (lateral

and median) bristles reddish-black and well developed, others absent; pleural

integument dark brown; antepronotum with narrow curved white scales, several

long golden to brown bristles; postpronotum covered with narrow curved scales,

dark reddish-black ones anteriorly and dorsally, large lower patch of white

ones, 6-7 posterior brown bristles; propleuron with a patch of broad white

scales, several golden bristles; postspiracular area with a patch of broad white

scales and a few narrow curved ones anteriorly, 5-8 golden bristles; sub-

spiracular area with 2 patches of broad white scales, lower one larger; mese-

pisternum with an upper and a posterior patch of broad white scales, several

upper and posterior golden bristles, lower ones shorter; prealar knob with

several golden bristles; paratergite with a row of moderately broad white

scales on lateroventral margin; mesepimeron with a patch of broad white scales

and several golden bristles on upper area; other pleural areas bare. Legs

(Fig. 16). Coxae I-III each with several dark brown bristles, I with anterior

surface covered with broad white scales and a patch of brown ones at about

middle, II with anterior surface covered with broad white scales, III with an

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

patch of broad white scales; femora I-III each with a dorsobasal white spot and

a few apical white scales on lateral surfaces, I, II each with anterior surface

Reinert: Aedes (Aedimorphus) in Southeast Asia 25

brown and a few pale scales on ventral surface, III with anterior and posterior

surfaces brown with a white stripe from base to near apex, stripe broad basally

and tapering to a point apically, I, I with posterior surface white with a brown

stripe from base to apex, stripe ventral on I and dorsal on II; tibiae I-III brown,

I with a posteroventral longitudinal white stripe and a dorsoapical white spot,

II, II each with a posterior longitudinal white stripe, a narrow basal white

band and a few dorsoapical white scales, stripe on III on apical 0. 80; tarsi I, II

brown with tarsomeres 1-3 each with a dorsobasal white spot; tarsus II with

tarsomeres 1-3 each with a narrow basal white band, tarsomere 4 with a

dorsobasal white spot; posttarsi I-III each with 2 ungues, I, II equal, each

bearing a tooth, III equal, simple. Wing. Dorsal veins covered with moder-

ately broad brown scales; costa with a patch of broad white scales at base;

ventral veins brown scaled; alula with narrow brown scales along fringe; 2

remigial bristles. Halter. Pedicel pale, capitellum white scaled with a few

brown scales mesally. Abdomen. Terga brown; tergum I with a few baso-

median pale scales and a rectangular patch of white scales on laterotergite;

terga II-VI each with a basal white band, bands wider on III-V in Thailand

specimens; tergum VII with a few basomedian and apical white scales; terga

II-VII each with a large laterobasal white patch covering most of lateral sur-

face, a few brown scales forming an indistinct dark spot toward the center of

each white patch, on some specimens white patch reduced or absent on VII and

brown scales more numerous on lateral surfaces of III-VI; sterna white scaled

with a narrow posterior brown band on III, IV and usually a few brown scales

on posterior of V, VI; terga and sterna with numerous golden bristles, mostly

along posterior margins. Genitalia (Fig. 19). Tergum VII. index 0.98-1.01;

sternum VIII index 0. 79-0. 85; tergum IX bilobed with 3-6 bristles on each lobe,

index 0.95-1.05; insula tongue-like, covered with minute setae and with 4-6

small tuberculi on apical 0.25; lower vaginal lip narrow, moderately pigmented,

covered with minute setae; upper vaginal lip narrow, heavily pigmented, cover-

ed with minute setae; upper vaginal sclerite large, moderately pigmented, tri-

angular shaped; postgenital lobe moderately long, apex with a moderately deep

median indentation, 4-8 bristles on each side of midline, covered with minute

setae, dorsal PGL index 1.04-1.20, ventral PGL index 2. 04-2. 24; cercus mod-

erately long, 1.00 extended from segment VII, narrow, apex sharply rounded,

numerous bristles on dorsal and lateral surfaces, index 2.52-2.79, cercus/

dorsal PGL index 3. 20-3. 79; 3 spherical, pigmented seminal capsules, 1 large

and 2 slightly smaller ones.

MALE (Fig. 2). Similar to female in general habitus. Head, Maxillary

palpus brown with segments 2-5 each with a white basal band, longer than pro-

boscis by 0.50 length of apical segment; vertex with narrow scaled area reduced.

Legs (Fig. 17). Posttarsi I-III each with 2 ungues, I, II with ungues unequal,

each bearing a tooth, III equal, simple. Abdomen. Terga II-VII each with a

narrow dorsobasal white band connected to a laterobasal white patch; tergum

VIII completely white scaled; sterna III-VIII with a narrow posterior brown

band. Genitalia (Fig. 35). Tergum IX strongly bilobed with 4-8 bristles on

each lobe, entire surface covered with minute spicules; gonocoxite long and

moderately broad, dorsal surface covered with scattered short fine bristles

and a few moderately long ones on 0.30 of lateroapical margin, numerous long

stout bristles along lateral margin from base to apex, ventral surface with long

stout bristles on apical 0.25 and along distal 0.60 of sternomesal margin, a few

moderately long bristles below and mixed with long ones, scattered scales on

lateral and ventral surfaces; gonostylus with pedicel long and narrow, base

somewhat broader, distal 0.38 expanded into a mesal lobe and a lateral, narrow,

26 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

apically rounded horn attached approximately 0.78 from base with a short fine

hair at apex, mesal expanded lobe with a moderately long, somewhat flattened,

apically pointed gonostylar claw attached mesally near base, 3-5 short stout

bristles along apical margin, mesal one short and others each increasing in

length, 3 short fine hairs along mesal margin of distal 0.50 of pedicel; basal

mesal lobe short and rounded apically, distal 0.25 with 5-8 short bristles,

entire surface covered with short hair-like spicules; proctiger short, paraproct

with a subapical thumb-like process, cercal setae absent; phallosome with

aedeagus of type I with 2 lateral plates connected basally, each plate with 6-7

short blunt lateral teeth on distal 0.58 and covered with a dorsal flap, paramere

long, approximately 0.90 length of lateral plate; sternum IX large, entire sur-

face covered with minute spicules, 7-15 bristles near center.

PUPA (Fig. 50). Chaetotaxy as figured and recorded in Table 2. Pat-

ches of spicules on middorsal ridge of cephalothorax extending to metanotum,

on metanotum mesally between hairs 10-C and on abdomen between hairs 1-I.

Cephalothorax. Hair 5-C with 2-5 branches; 7-C with 2-4 branches; 8-C with

5-7 branches. Respiratory trumpet. Lightly pigmented; index 3. 67-4. 21,

average 3.85. Metanotum. Hair 10-C with 7-11 branches; 12-C with 5-8

branches. Abdomen. Hair 5-I with 6-10 branches; 1-II with 20-32 branches;

4-II with 2-5 branches; 1-III with 5-8 branches; 6-VI single or double; 1-VII

with 2-5 branches; 6, 9-VII with 3-7 branches; 11-VII single. Paddle. Ovoid;

with very minute serrations along basal 0.55 of outer margin; minute spicules

on distal dorsolateral 0.25 of outer surface; midrib does not reach apex; hair

1-P short, single; index 1.14-1.47, average 1.25.

LARVA (Figs. 60, 61). Chaetotaxy as figured. Head. Hairs 1, 3,

14-C single; 4-C with 4-9(5) branches; 5-C with 5-7(5) branches; 6-C with

4-5(4) branches; 7-C with 7-11(9) branches; 8, 10-C with 2-4(3) branches;

9-C with 3-4(3) branches; 11-C with 4-10(6) branches; 12-C with 5-8(5) branches;

13-C with 5-8(6) branches; 15-C with 4-6(5) branches; basal maxillary hair

single; mental plate with 22-26(24) teeth. Antenna. Lightly pigmented; spicules

scattered over entire shaft, more numerous on basal 0. 50; hair 1-A with 5-7(5)

branches, inserted at 0.45-0.52 from base; 2-A long; 3-A approximately 0. 50

length of 2-A. Thorax. Hair 0-P with 5-12(5) branches; 1, 5, 6, 10, 12-P

single; 2, 8-P double; 3-P with 2-4(3) branches; 4, 11-P with 2-4(2) branches;

7-P double or triple (3); 9-P single to triple (2); 14-P single to double (1); 1-M

with 3-6(3) branches; 2-M single to 5(1) branched; 3, 11-M single to double (1);

4-M with 3-7(3) branches; 5, 7, 10, 12-M single; 6-M with 4-7(5) branches; 8,

9-M with 5-9(7) branches; 13-M with 4-10(8) branches; 14-M with 6-12(8)

branches; 1, 12-T single to triple (1); 2-T with 3-6(5) branches; 3-T with

6-16(9) branches; 4-T with 3-5(4) branches; 5, 10-T single; 6-T with 2-4(3)

branches; 7-T with 7-10(9) branches; 8-T with 4-7(5) branches; 9-T with 4-8(7)

branches; 11-T single or double (1); 13-T with 6-9(8) branches. Abdomen.

Hairs 0, 4, 14-VIII single; 1-VIII with 5-8(6) branches; 2-VIII with 2-4(3)

branches; 3-VIII with 8-15(12) branches; 5-VIII with 4-7(5) branches; 6-V-VI

short; comb with 23-32(24) scales arranged in 3 irregular rows, scales short

and bluntly rounded with short stout denticles along lateral and apical margins;

1, 3-X single; 2-X with 8-12(11) branches; ventral brush with 9-10 (usually 10)

hairs on grid and 1-2 precratal ones; saddle lightly pigmented with minute ridges,

incompletely rings segment, with a few spicules along posterior margin, acus

absent; 4 anal papillae, long, each with a broad base and tapering to a pointed

apex. Siphon. Lightly pigmented with ridges over entire surface; 2-4 rows of

stout spicules circling apex; usually a dorsal and a ventral patch of spircules at

about middle of siphon, these patches vary considerably from one to both absent

Reinert: Aedes (Aedimorphus) in Southeast Asia 27

to both extending over 0.45 of the siphon; acus absent; index 3.13-3.47; pecten

with 16-23(21) teeth, apical 2-3 teeth longer, smooth and wider spaced than

remainder which have a slender apical attenuated filament with 1-3 basal denti-

cles; hair 1-S with 4-6(5) branches, inserted at 0.77-0. 83 from base.

TYPE DATA. Culex caecus Theobald, holotype female, Klang Mangrove

Swamp, Selangor, MALAYSIA, 28 October 1899, A. L. Butler, in British Muse-

um (Natural History).

DISTRIBUTION. Specimens examined--525 males, 827 females, 565

pupae, 814 larvae and 788 individual rearings (384 pupal, 404 larval) from the

following locations:

BANGLADESH (East Pakistan). Chittagong Hill Tracts, Rangamatti.

CAMBODIA. Phnom-Penh.

CHINA. Wangtun, Canton Delta, Sam Shui, Wang Mun; Chekiang,

Shanghai; Yunnan.

INDIA. Assam, Chabua, Dibrugarb, Doom Dooma, Tezpur; Malabar

Coast.

INDONESIA. Cevam, Ilatoenoera; Java, Batavia, Gomrong, Padaherang,

Pelaboean Ratoe, Rawallah, Uodjowarna; Sumatra (?), Katta Tjane.

MALAYSIA. Kedah, Bt. Kayu Hitam, Changlum, Kg. Peng Besar,

Sintok F. R.; Kelantan, Bertam; Pahang, Bt. Belong, Chegar, Kuantan, Kuala

Lipis, Merapoh, Perah; Perak, Kuala Kangsar; Perlis, Bt. Bintang F. R.,

Chior F. R., Kg. Gunong, Kg. Prok Buah, Kubang Tiga, Mata Ayer, To'Kaya-

man; Selangor, Ampang F. R., Banting, Klang Mangroves, Puchong, Rantau

Panjang, Segambut, Ulu Gombak, Ulu Klang; Trengganu, Kula Brang, Kuala

Dura, Marang, Payang Kayu; Segambut, Serdang.

PHILIPPINES. Palawan, Panitian.

SINGAPORE.

SOUTH VIETNAM. Binh Dinh, An Khe; Nha Trang, Duc My; Phuoc

Long, Bu Dop; Tay Ninh, Tay Ninh.

THAILAND. Chiang Mai, Chiang Mai, Huey Keo; Kanchanaburi, Ban

Sai Yok, Farng, Huai Bong Ti, Huai Mae Nam Noi, Khao Na Chang, Maki;

Khon Kaen, Ban Muang Kao, Phu Wiang; Lampang Ban Sop Pon, Ngao; Mae

Hong Son, Ban Mae Tia, Doi Chang; Nakhon Ratchasima, Banna Nabon, Chawang,

Dhong Suea Pan, Khoo Yai, Pak Chong, Pak Tong Chai; Nakhon Si Thammarat,

Ban Rim Thanon, Ban Tan, Chaung Khao, Thung Song; Nan, Ban Pang Chom

Phu; Narathiwat, Khan Lau, Waeng, Loh Choot; Phangnga, Khao Sung, Nam Tai,

Pak Chaung, Pathum, Thap Wen; Prachin Buri, Ban Bu Phram, Ban Tub Lan;

Ranong, Ban Bang Hin, Ban Chatri, Kraburi, Vat Pra Chum Pharam; Songkhla,

Haad Yai; Suvat Thani, Ban Cha Weng, Ban Tai, Ban Tahing Ngam, Ban Thuri-

an, Laen. Thong Lak, Ko Samui; Tak; Trang; Udon Thani, Ban Kau Noi, Ban Pa

Goi, Tai Ton Kam Phoo, Muang.

Other distribution.

BANGLADESH (East Pakistan). Rangamatti, Chittagong Hill Tracts

eae 1934: 258); Thakurgaon, Akcha, Madargonj (Aslamkhan and Wolfe

197431);

BURMA. Rangoon (Barraud 1934: 258).

INDIA. Assam, Dinapur; Bengal, Sukna; Colaghat (Barraud 1934: 258);

Sukna, E. Himalayas (Theobald 1910a: 21, Theobald 1910b: 299).

INDONESIA. New Guinea; Stonaten (Brug 1926: 529, McDonald 1957: 21);

oe Atchin, Kotta Tjane; Djambi, Moeara Tebo (Brug and Edwards 1931:

258

MALAYSIA. Selangor, Kepong Panjang (McDonald 1957: 21).

NEPAL. Gandaki, Kaski, Pokhara (Joshi et al. 1965: 139).

28 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

SINGAPORE (Colless 1959: 260).

SOUTH VIETNAM (Borel 1930: 271); Cam Ranh Bay (Reisen et al. 1971:

Table 12).

THAILAND. Chieng Moeang (Causey 1937: 413); Kanchanaburi,

Sangkla-buri, Lai-nam, Ni-thae (Harinasuta et al. 1970: 241).

TAXONOMIC DISCUSSION. Aedes caecus shares a number of char-

acters with vexans vexans which became evident during this study when a

number of adult female caecus were found in several museum collections

labeled as vexans. It can be separated from vexans vexans by the following

features: femur II with anterior brown scaled; scutum with 3-4 bristles on

scutal fossal area; scutellum with both narrow curved and broad white scales

on each lobe; postspiracular area with 5-6 bristles; prealar knob without scales;

and female palpus brown while vexans vexans has: femur II with white scales

intermixed with brown ones on anterior surface; scutum with 7-10 bristles on

scutal fossal area; scutellum with narrow curved golden-white scales on each

lobe; postspiracular area with 8-9 bristles; prealar knob with a few broad

white scales; and female palpus with apex white scaled. Aedes caecus also

superficially resembles culicinus from which it is easily distinguished by the

banded tarsi.

The pupa is characterized by the following features: a large patch of

spicules mesally between metanotal hairs 10-C and abdominal hairs 1-I (orbitae

has a Similar arrangement of spicules but is easy to separate on branching of

hair 10-C); hairs 10-V, VI long and single; and hair 7-VI long and single.

The larval stage resembles orbitae and is discussed under that species.

Edwards (1913: 228) synonymized Aedes suknaensis Theobald with Aedes

imprimens Walker but later (1934: 170) he questionably included it with caecus.

Barraud (1928: 663) listed suknaensis as a synonym of impvimens but stated

that he thought suknaensis and caecus were possibly conspecific. In this same

article (Plate 61, Fig. 1) he also illustrated the male genitalia of caecus but

mistakingly called it imprimens. Later, Barraud (1934: 257) followed Edwards

and listed suknaensis as a synonym of caecus. I have examined a cotype of

suknaensis from Sukna, India, in the British Museum (Natural History) and

find it not to be caecus but very similar to imprimens. It differs, however,

from imprimens in having denser patches of scales on the pleural thoracic areas

and scutellum and possessing a number of short fine golden hairs mixed with the

single patch of broad white scales on the subspiracular area. Aedes suknaensis

belongs in the subgenus Edwardsaedes Belkin and may be a distinct species from

imprimens, but since no males have been found and the habitus of the adults are

very similar I am retaining it with imprimens for the present.

Aedes caecus can be separated from imprimens by having 2 patches of

scales on the subspiracular area while the latter species has a single patch.

BIOLOGY. Immatures are usually collected from partially shaded,

small flood pools and animal footprints containing turbid fresh water located

in bamboo groves and forests. Adults readily feed on cattle and man. Imma-

tures in Thailand were collected most often from small and large flood pools,

wheel tracks and elephant footprints but also from a small flood pool ina

streambed, small ditch, large ground pit, posthole and a small rice field;

usually from clear or turbid fresh water but several times from colored fresh

water; water usually with floating and submerged dead leaves or scarce aquatic

vegetation; in partial shade or unshaded areas and once from a heavy shaded

area; usually in primary bamboo groves and secondary and primary rain forests

in mountains and plains but also in coconut, palm, bamboo and rubber groves

in the plains, secondary scrub in valley and plains, and secondary rain forest

Reinert: Aedes (Aedimorphus) in Southeast Asia 29

in a valley; and at an altitude of 7 to 1, 970 feet (most often from 100 to 750

feet). Larvae were collected in association with the following species of mos-

quitoes: Aedes alboscutellatus, andamensis, culicinus, ferinus, gubernatoris,

imprimens, indicus, mediolineatus, orbitae, vexans vexans, vittatus; Ano-

pheles balabacensis, indiensis, kochi, maculatus, philippinensis, subpictus,

vagus, Culex bitaeniorhynchus, fuscocephalus, mimulus, nigropunctatus,pal-

lidothovax, pseudovishnui, raptor, scanloni, sinensis; and Uvanotaenia bicolor.

Adults were taken in light traps and biting cattle and man in bamboo groves and

underbrush.

Larvae in South Vietnam were taken from water in a tire, marshy

depression, rock pool, foxhole, ground pools, artifical container, pool ina

concrete piling and a jungle pool. In Java, larvae were collected in wheel ruts

and adults taken indoors. Adults were collected biting cattle, and larvae from

hoofprints and a small pool in Malaysia. Larvae in India were found in muddy

puddles and a swamp.

Larvae were collected from a buffalo wallow on a high plateau in Thai-

land (Causey 1937: 413); ground pools in China (Chow 1949: 129); and natural

pools in open jungle in India (Barraud 1934: 258) and New Guinea (Steffan 1966:

212). In Malaysia, adults readily fed on man and domestic animals and im-

matures were collected from pools and earthenware pots (Macdonald 1957: 21).

Aslamkhan and Wolfe (1971: 31) in Bangladesh (East Pakistan) collected adults

resting in a house, biting cattle and biting humans.

AEDES (AEDIMORPHUS) CULICINUS EDWARDS

(Figs. 3, 16, 17, 20, 36, 51, 62)

Aedes (Ecculex) culicinus Edwards 1922a, Indian J. med. Res. 10: 271 (co,

2); Edwards 1922b, Indian J. med. Res. 10: 467.

Aedes (Aedimorphus) culicinus (Edw.), Barraud 1928, Indian J. med. Res. 15:

667 (o'*, 9).

Aédes (Aédimorphus) culicinus Edwards, Edwards 1932, Genera Insec., Fasc.

194: 169; Barraud 1934, Fauna Brit. India, Diptera 5: 252 (o*, 9).

Aedes (Aedimorphus) culicinus Edwards, Stone et al. 1959, Thomas Say Found.

6;.191,

FEMALE (Fig. 3). Head. Antenna dark brown, approximately equal

in length to proboscis, pedicel dark with a few small dusky scales and a patch

of short fine brown hairs mesally, flagellomere 1 with a few dusky scales;

clypeus dark, bare; maxillary palpus brown scaled, approximately 0.17 length

of proboscis; proboscis brown scaled with a pale ventral stripe extending from

near base to distal 0.25, approximately 1.06 length of femur I; vertex with

dorsum covered with narrow decumbent scales arranged in an anteromedian

diamond-shaped brown group and the remainder white; lateral surface covered

with broad white scales, an anterodorsal dark patch and a dusky area anterior

to the antepronotum; numerous dark brown erect forked scales on occiput and

vertex extending anteriorly to the ocular line. Thorax. Scutal integument

dark brown; scutum covered with narrow curved reddish-brown scales; narrow

curved white scales forming patches on anterior promontory area, scutal fos-

sal areas (one on anterior extending along margin onto lateral area and a patch

on posterior area), scutal angles, supra-alar areas, posterior medial scutal

area and along lateral margins of prescutellar space; scutellum with a patch of

narrow curved white scales on each lobe; median anterior promontory, acro-

30 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

stichal, dorsocentral (anterior and posterior), scutal fossal (anterior, 3-4

lateral, 1-2 median and 1 posterior), supra-alar, several posterior medial

scutal, 1 postalar callar and scutellar (lateral and median) bristles reddish-

black and well developed; pleural integument brown; antepronotum with narrow

curved white scales and some moderately broad ones, several long brown

bristles; postpronotum with narrow curved brown scales dorsally and white

ones posteriorly, a lower posterior patch of broad white scales, 5-7 posterior

brownish bristles; propleuron with a patch of broad white scales, several pale

bristles; postspiracular area with a patch of broad white scales, 5-7 golden

bristles; subspiracular area with 2 patches of broad white scales, lower one

larger; mesepisternum with an upper and a posterior patch of broad white

scales, several upper and posterior pale bristles, lower ones shorter; pre-

alar knob with several pale brown bristles; paratergite with a row of broad

white scales; mesepimeron with a patch of broad white scales and several pale

bristles on upper area; other pleural areas bare. Legs (Fig. 16). Coxae I-III

each with several pale bristles, I with anterolateral white scales and a lateral

brown patch, II with anterior surface white scaled; trochanters I-III each with

a patch of broad white scales; femora I-III each with a few pale lateral scales

at apex, II, III each with an anteroventral white stripe, wider on III, I-III each

with a posterior broad white longitudinal stripe from base to approximately

0.75, stripe dorsal on I and ventral on II, III; tibiae I-III with anterior surface

brown, afew dorsal and lateral pale scales at apices, posterior surface brown

with a longitudinal pale stripe, I with stripe posteroventral, II, III with stripe

posteromedian; tarsi I-III brown; posttarsi I-III each with 2 ungues, I, II equal,

each bearing a tooth, III equal, simple. Wing. Dorsal veins covered with

moderately broad brown scales; costa with white scales at base and on its

posterior at humeral cross vein; ventral veins brown scaled; alula with narrow

brown scales along fringe; 2 remigial bristles. Halter. Pedicel pale, capi-

tellum white scaled. Abdomen. Terga brown; tergum I with a few dorsomedi-

an white scales and a rectangular patch of white scales on laterotergite; terga

II-VI each with a narrow dorsobasal white band; tergum VII with a triangular

dorsobasal pale patch; terga II-VI each with a large laterobasal white spot, a

few brown scales in center of spots, VII with a few lateromedian white scales;

sterna pale scaled with lateromedian pale brown spots; terga and sterna with

numerous pale golden bristles, mostly along posterior margins. Genitalia

(Fig. 20). Tergum VIII index 1.04-1.09; sternum VIII index 0.97-0.99; ter-

sum IX bilobed with 4-10 bristles on each lobe, index 0.91-1.12; insula tongue-

like, covered with minute setae and with 3-5 small tuberculi on apical 0. 25;

lower vaginal lip narrow, moderately pigmented, covered with minute setae;

upper vaginal lip narrow, heavily pigmented, covered with minute setae; upper

vaginal sclerite large, moderate to heavily pigmented, triangular shaped; post-

genital lobe moderately long, apex with a moderately deep median indentation,

3-7 bristles on each side of midline, covered with minute setae, dorsal PGL

index 0. 83-0.95, ventral PGL index 2.18-2.38; cercus moderately long, 0.60-

0.75 extended from segment VII, narrow, apex sharply rounded, numerous

bristles on dorsal and lateral surfaces, index 2.97-3.17, cercus/dorsal PGL

index 4. 75-5. 22; 3 spherical, pigmented seminal capsules, 1 large and 2 tiny

rudimentary ones.

MALE (Fig. 3). Similar to female in general habitus. Head. Maxil-

lary palpus with dorsal pale scale patches on middle of segments 2 and 3,

longer than proboscis by length of apical segment; vertex without anterodorsal

dark scale patch on lateral surface. Legs (Fig. 17). Posttarsi I-III each with

2 ungues, I, II with ungues unequal, each bearing a tooth, III equal, simple.

Reinert: Aedes (Aedimorphus) in Southeast Asia 31

Wing. Dorsal veins completely brown scaled. Abdomen. Tergum I with a

lateral band of white scales on laterotergite; terga II-VIII each with a narrow

dorsobasal white band, usually 2-4 white scales on lateromedian surfaces of

II-VIIl. Genitalia (Fig. 36). Tergum IX bilobed with 4-8 bristles on each lobe,

entire surface covered with minute spicules; gonocoxite long and moderately

broad, dorsal surface with a large dense patch of short bristles along tergo-

mesal margin from near base to distal 0.30, numerous long stout bristles on

lateral margin from base to apex, ventral surface with similar bristles on

distal 0.25 with moderately long and short ones proximally, scattered scales

on lateral and ventral surfaces; gonostylus with pedicel short and broad, dis-

tal 0.50 greatly expanded with a laterotergal horn-like flap covered with nu-

merous long fine hairs and terminating in an apical point, mesal margin of

expanded portion with a short flat curved pigmented gonostylar claw mesally

near middle, a patch of 6-14 fine hairs basal to claw and a tergal patch of 5-

10 similar ones, 8-11 fine hairs along apical margin; basal mesal lobe short

and rounded apically, distal 0.50 with 6-10 short bristles, a long narrow

lightly pigmented filament extending from apex to near base of gonostylus and

attached to mesal membrane of gonocoxite, entire surface of basal portion

covered with short hair-like spicules; proctiger short, paraproct with a sub-

apical thumb-like process, cercal setae absent; phallosome with aedeagus of

type I with 2 lateral plates connected basally, each plate with 6-7 short, blunt,

lateral teeth on distal 0.54 and covered with a dorsal flap, paramere long,

approximately 0.78 length of lateral plate; sternum IX large, entire surface

covered with minute spicules, 3-5 bristles near center.

PUPA (Fig. 51). Chaetotaxy as figured and recorded in Table 3. Ce-

phalothorax. Hair 5-C with 4-6 branches; 7-C with 3-5 branches; 8-C with

5-9 branches. Respiratory trumpet. Moderately pigmented; index 3. 61-4. 90,

average 4.37. Metanotum. Hair 10-C with 6-9 branches; 12-C with 4-7

branches. Abdomen. Hair 5-I with 7-12 branches; 1-II with 14-20 branches;

4-II with 2-5 branches; 14-II single; 1-III with 7-13 branches; 6-VI double or

triple; 1-VII with 5-8 branches; 6-VII with 6-9 branches; 9-VI with 4-5 bran-

ches; 11-VII double. Paddle. Ovoid; with very minute serrations along basal

0.55 of outer margin; tiny spicules along apical 0.45 of outer and apical 0.30

of inner margins; midrib does not reach apex; hair 1-P short, single or double;

index 1.13-1.34, average 1. 22.

LARVA (Fig. 62). Chaetotaxy as figured. Head. Hairs 1, 3, 14-C

single; 4-C with 5-7(7) branches; 5-C with 3-4(3) branches; 6-C triple; 7-C

with 4-6(6) branches; 8-C with 2-6(2) branches; 9-C with 2-5(5) branches; 10-C

single or double (2); 11-C with 2-5(3) branches; 12-C with 3-4(4) branches,

13-C with 4 branches; 15-C double; basal maxillary hair single; mental plate

with 23-25 teeth. Antenna. Moderately pigmented; scattered stout spicules

over entire shaft, spicules somewhat longer past middle; hair 1-A with 5-8(7)

branches, inserted at 0.37-0.42 from base; 2-A long; 3-A approximately 0.96

length of 2-A. Thorax. Hair 0-P with 7-10(8) branches; 1, 5, 6, 8, 10, 12,

14-P single; 2, 7, 11-P double; 3-P with 3-4(3) branches; 4-P double or

triple (2); 9-P single or double (2); 1-M with 3-8(4) branches; 2-M with 2-4(3)

branches; 3-M single or double (1); 4-M double or triple (3); 5, 7, 10-12-M

single; 6-M with 4-6(4) branches; 8-M with 3-4(4) branches; 9-M with 3-6(5)

branches; 13-M with 4-5(5) branches; 14-M with 5-8(5) branches; 1-T single

to triple (3); 2-T with 4-13(6) branches; 3-T with 4-8(5) branches; 4, 9-T with

2-4(3) branches; 5, 10-12-T single; 6, 11-T single or double (2); 7-T with 5-6

(5) branches; 8-T with 4-6(4) branches; 13-T with 4-7(5) branches. Abdomen.

Hairs 0, 14-VIII single; 1-VIII with 3-4(4) branches; 2-VIII double; 3-VIII with

32 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

7-10(8) branches; 4-VII double or triple (2); 5-VIII with 5-7(5) branches;

6-IV-VI short; comb with 14-20(17) scales arranged in 2 irregular rows,

scales each with a long stout pointed median spine and short denticles along

lateral margins of base; 1-X double or triple (2); 2-X with 8-10(8) branches;

3-X single; ventral brush with 8-9 hairs on grid and 4 precratal ones; saddle

lightly pigmented, incompletely rings segment, with a few spicules along

posterior margin, acus absent; 4 anal papillae, long, tapering to a blunt apex.

Siphon. Lightly pigmented; acus present; index 3.88-4.50; pecten with 11-14

(11) teeth, apical 2-4 teeth smooth and wider spaced than remainder which

have a slender apical attenuated filament with 1-2 basal denticles; hair 1-S

with 3-5(4) branches, inserted at 0.70-0.72 from base.

TYPE DATA. There has been some confusion about the location of the

types of Aedes (Ecculex) culicinus Edwards. In the original description of the

species, Edwards (1922a: 272) states the holotype male and allotype female

were collected at Delhi, INDIA, April 1911, Major S. R. Christophers, and

deposted in the Central Malaria Bureau, Kasauli, and 1 paratype female from

Amritsar, INDIA, April 1911, Major S. R. Christophers, was deposited in the

British Museum. Barraud (1928: 667) lists the holotype male in the Central

Malaria Bureau and the allotype female and paratype female in the British

Museum but in 1934 (page 252) he states all the types are in the British Muse-

um. Stone et al. (1959: 191) lists the holotype male and allotype female in the

Malaria Institute of India, Delhi, India. I have examined the collection in the

British Museum (Natural History) and only the paratype female from Amritsar

is there. The primary types are in the collection at the National Institute of

Communicable Diseases, Delhi, India, which received the collections from the

Central Malaria Bureau.

DISTRIBUTION. Specimens examined--17 males, 106 females, 105

pupae, 8 larvae, 105 individual rearings (105 pupal), from the following loca-

tions:

CAMBODIA. Kirirom.

INDIA. Punjab, Amritsar.

SOUTH VIETNAM. Binh, Dinh, An Khe; Kontum, Kontum.

THAILAND. Kanchanaburi, Ban Sai Yok; Khon Kaen, Ban Muang Kao,

Phu Wiang; Ubon Thani, Chongmek.

WEST PAKISTAN. West Punjab, Lahore.

Other distribution.

INDIA. Delhi, Kasauli (Edwards 1922a:; 272); Karnal (Barraud 1934:

262),

WEST PAKISTAN. Lahore, Changa Manga National Forest (Aslamkhan

and Salman 1969: 193).

TAXONOMIC DISCUSSION. The adult habitus and female and male

genitalia of culicinus are similar to alboscutellatus and are discussed under

that species. The most distinctive features of the adults are: scutellum with

narrow white curved scales on each lobe; postpronotum with a few posterior

broad white scales in addition to narrow curved ones; tarsi dark scaled; and

female with 1 large and 2 rudimentary seminal capsules.

The most distinctive characters of the larvae are: comb of 14-20

scales each of which has a long stout pointed median spine and stout denticles

along lateral margins of the base; pecten of 11-14 teeth; siphon index of 3. 88-

4.50; head hair 6-C triple; and metathoracic hair 9-T double or triple.

BIOLOGY. The immatures are usually found in partially shaded ground

pools containing fresh water located in secondary scrub vegetation. The adults

prefer feeding on cattle to man. Immatures in Thailand have been collected

Reinert: Aedes (Aedimorphus) in Southeast Asia So

from small to large turbid fresh water flood pools, with floating and submerged

dead leaves, in partial shade, located in secondary scrub vegetation in the

mountains or in teak plantations in the plains, and at an altitude of 650 to 1, 310

feet. In Malaysia immatures were taken in ground pools.

In Thailand immatures were collected in association with the following

species of mosquitoes: Aedes alboscutellatus, caecus, ferinus, imprimens,

mediolineatus, pallidostriatus, vittatus; Anopheles kochi, maculatus; and

Culex annulus, fuscocephalus, vaptor.

Aslamkhan and Salman (1969: 185, 186, 189, 193) in West Pakistan list

culicinus as making up 7.58 percent of the daytime and 12.4 percent of the

nighttime human biting mosquito collections. This species, however, preferred

to feed on cattle to man at a ratio of 3:1 during nighttime tests. Adults made up

31.6 percent of light trap mosquito collections. The immatures were found in

ground pools with grassy margins and decomposing algae on the bottom. Aedes

culicinus occurs in West Pakistan throughout the year but the numbers increase

from April until a peak population is reached in July when relative humidity is

very high and breeding places are abundant.

AEDES (AEDIMORPHUS) DA VIDI BASIO

Aedes (Aedimorphus) sp., Basio, White and Reisen 1970, Philipp. Ent. 1: 443.

Aedes (Aedimorphus) davidi Basio 1971, Nat. Mus. Philipp., Monogr. No. 4,

p. 11 (nomen nudum); Basio 1971, Philipp. Ent. 2: 51 (o'*) (validated

name with description).

MALE. Head, Antenna with pedicel bare; proboscis dark with a dis-

tinct pale scaled area ventrally at middle; vertex covered with a mixture of

pale and brownish scales; lateral surface with broad brownish colored scales.

Thorax. Scutum covered with a mixture of pale and brown scales; scutellum

with brown scales on lateral and median lobes; antepronotum with pale scales;

postpronotum with brownish-yellow scales; postspiracular area with narrow

pale scales; subspiracular area with narrow pale scales; mesepisternum with

narrow pale scales; mesepimeron with narrow pale scales. Legs. Femora

I-III each speckled with white scales with a few brownish colored scales later-

ally, apex without a white scaled spot; tibiae I-III each with a mixture of pale

and brownish colored scales on anterior surface, ventral surface pale scaled

and a subapical pale scaled band; tarsi I-III with pale bands; posttarsi I-III

each with 2 ungues, I and II with ungues equally developed, III equal and simple.

Wing. Length 3.0mm. Abdomen. Terga I-IV each with distinct pale scales

laterally, V-VII each with pale scales basally, VIII pale scaled; sterna I-VIII

with ground cover of brown scales. Genitalia. Tergum IX strongly bilobed

with 3-4 short bristles on each lobe; gonocoxite long and moderately broad,

dorsal surface with long stout bristles along lateral margin from base to apex,

short thin bristles along tergomesal margin from base to apex, remainder of

dorsal surface with short thin bristles, lateral surface with long stout bristles

from base to apex; gonostylus approximately 0.60 length of gonocoxite, with

basal portion narrow and apical 0.40 expanded and with several short fine

hairs, gonostylar claw long, narrow, pigmented, with apex blunt and attached

subapically; basal mesal lobe absent; proctiger moderately long, apex of para-

proct bluntly pointed, cercal setae absent; phallosome with aedeagus of type II

with 2 lateral plates connected basally, each plate apparently with 5 longidu-

dinal lateral teeth with tergally curved apices.

34 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

FEMALE, PUPA, LARVA and EGG. Not described, however, Basio

(1971: 52) states the holotype male has damaged pupal and larval skins.

TYPE DATA. Aedes (Aedimorphus) davidi Basio, holotype male with

damaged larval and pupal skins and genitalia on slides, PHILIPPINES, Laguna,

Los Banos, U. P. College of Agriculture Campus, 22 January 1970, collector

D. W. White, from a rice field at 33 feet elevation, deposited in Philippine

Museum, Manila and 1 paratype male with genitalia on a slide, same data as

holotype, deposited in R. B. Basio collection (Basio 1971: 51, 52).

DISTRIBUTION. Known only from the type locality.

TAXONOMIC DISCUSSION. No specimens of davidi were available for

examination, therefore the above description is taken from the original one by

Basio (1971: 51-53). In his introductory remarks to the description of davidi,

Basio states "The male of this species resembles that of Aedes (Aedimorphus)

vexans nocturnus (Theobald)"' (now considered as vexans vexans) "by the scal-

ings of the dorsum, mesoscutum, scutellum, anterior and posterior pronotum,

femora, tibiae, tarsi and abdominal tergites,and by having the torus bare."

In his description of the scaling of the male that followed, however, he stated

". . . entire scutellum brown. . .abdominal tergites I to IV with distinct pale

scales laterally, V to VII with pale scales basally and VIII entirely pale-scaled

. . .'' Basio's introductory remarks and species description do not coincide

with each other. For example, vexans vexans from the Philippines and else-

where possesses narrow curved white scales on the lateral and median lobes

of the scutellum, not brown scales and the pedicel (torus) is not bare but has

afew small, broad, white scales mesally. The abdominal terga of vexans

vexans also have the following patterns: terga brown scaled without lateral

white spots but with white scales covering laterotergite of I, II with a white ba-

sal band with posterior margin straight, III[-VII each with a basal white band

with lateral margins expanded posteriorly, VIII white scaled or white with a

few brown scales mesally while Basio (1971: 51, 52) states that the abdominal

tergites of davidi resemble those of vexans vexans but in his description of

the male of davidi he reports the following: tergites I-IV with distinct pale

scales laterally, V-VII with pale scales basally and VIII entirely pale scaled.

The abdominal scale patterns reported for davidi are in fact quite different

from those of vexans vexans.

From the description and male genitalia illustration davidi appears to

have some similarity to vexans vexans; however, the following characters, as

well as the above-mentioned ones, differ from the latter species: subspiracu-

lar area, mesepisternum and mesepimeron with narrow scales; femora I-III

each without an apical white spot; tibiae I-III each with a pale subapical band

and ungues of posttarsiI and II equally developed. The male genitalia of

davidi, as illustrated and described by Basio (1971: 52, 53), also differ from

those of vexans vexans by the following: gonostylus with basal portion nar-

row and apical 0.40 expanded and rounded, tergum IX bilobed with 3-4 bristles

(illustration shows 5 bristles on each lobe), aedeagus with apparently 5 teeth

on each lateral plate and the apparent absence of the basal mesal lobe. In my

opinion the genitalia of the 2 type specimens must have been damaged and the

basal mesal lobes lost since it would be most unusual to have this structure

absent in Aedimorphus.

BIOLOGY. The 2 males were reared from larvae collected from a

rice field at an elevation of 33 feet and in association with Aedes vexans vexans

(Basio 1971: 52).

Reinert: Aedes (Aedimorphus) in Southeast Asia 35

AEDES (AEDIMORPHUS) LOWISII (THEOBALD)

(Figs. 4, 16, 17, 21, 37)

Reedomyia lowisii Theobald 1910, Monogr. Cul. 5: 257 (o*, 9*); Brunetti

1912, Rec. Indian Mus. 4: 487.

Ochlerotatus lowisii Theob., Brunetti 1920, Rec. Indian Mus. 17: 140.

Aedes (Ecculex) lowisi Theobald, Edwards 1922b, Indian J. med. Res. 10: 466.

Ochlerotatus lowisi Theobald, Senior-White 1923, Cat. Indian Insects, Cul.,

20 FG

Aedes (Aedimorphus) lowisii(Theo.), Barraud 1928, Indian J. med. Res. 15:

658 (2); Stone et al. 1959, Thomas Say Found. 6: 194.

Aédes (Aédimorphus) lowisi Theobald, Edwards 1932, Genera Insec., Fasc.

194: 168; Barraud 1934, Fauna Brit. India, Diptera 5: 250 (co, 9).

Aedes (Aedimorphus) mindoroensis Knight and Hull 1951, Pacif. Sci. 5: 199

(o*, 9); Knight and Hull 1953, Pacif. Sci. 7: 459 (c%, 9); Stone et al.

1959, Thomas Say Found. 6: 194. NEW SYNONYM.

FEMALE (Fig. 4). Head. Antenna dark brown, approximately 0.91

length of proboscis, pedicel brown with several small pale scales and a patch

of short fine brown hairs mesally, flagellomere 1 with a few small dark scales;

clypeus dark, bare; maxillary palpus dark brown scaled, approximately 0.17

length of proboscis; proboscis dark brown scaled with a pale ventral stripe

extending from near base to distal 0.25, pale area narrow basally becoming

broad distally, approximately 1.18 length of femur I; vertex with dorsum

covered with narrow decumbent scales arranged in an anteromedian diamond-

shaped dark group and the remainder golden; lateral surface covered with

broad pale scales, an anterodorsal dark patch and a dusky area anterior to

antepronotum; numerous long dark brown erect forked scales on occiput and

vertex extending anteriorly to ocular line. Thovax. Scutal integument reddish-

brown; scutum covered with narrow curved reddish-black scales, scattered nar-

row curved white scales forming indistinct spots on anterior promontory area,

supra-alar areas, and posterior medial scutal area, similar scales forming a

distinct pair of small circular patches on both anterior and posterior scutal

fossal areas; scutellum with a patch of broad silvery scales on each lobe; medi-

an anterior promontory, acrostichal, dorsocentral (anterior and posterior),

scutal fossal (anterior, 1 lateral and 1-2 median), supra-alar, several pos-

terior medial scutal, 1 postalar callar and scutellar (lateral and median) bris-

tles black and well developed, others absent; pleural integument dark brown;

antepronotum with a few narrow curved pale scales, several long dark bris-

tles; postpronotum with scattered narrow curved dark brown scales on dorsal

0.50, 5-6 posterior long brownish-black bristles; propleuron with a patch of

broad pale scales, several short golden bristles; postspiracular area with 5-7

brown or black bristles; subspiracular area with a few narrow brownish hair-

like scales; mesepisternum with an upper and a posterior patch of broad silvery

scales, several upper and posterior dark bristles, lower ones shorter; prealar

knob with several golden-brown bristles; paratergite bare; mesepimeron with a

patch of broad silvery scales and several brown bristles on upper area; other

pleural areas bare. Legs (Fig. 16). Coxae I-III each with several golden bris-

tles, I with anterior surface covered with broad brownish scales and a small

patch of broad white ones dorsally and one ventrally, II with a patch of broad

white scales anteriorly, III with a few broad white scales anteroventrally; tro-

chanters I-III each with a patch of broad white scales; femur I with an anterior

and a posterior apical white spot, II, III with a dorsoapical white spot, I, II with

36 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

anterior surface brown, I with a posterior narrow dorsal pale line from base

to apex, pale area broad basally and tapering apically, I with a posteroventral

white stripe, II with a posterodorsal white stripe, II brown with anterior and

posterior ventrobasal white areas, areas broad basally and tapering to a point

at about 0.25 from apex; tibiae I-III brown, each with a dorsoapical white spot;

tarsi I-III brown, I with tarsomere 1 having a few dorsoapical yellowish scales,

tarsomere 2-4 each with a few dorsobasal and dorsoapical yellowish scales,

tarsomere 5 covered with yellowish scales, II with tarsomere 1 having a few

dorsoapical yellowish scales, tarsomere 2 with a few dorsobasal and dorso-

apical yellowish scales, tarsomere 3-4 each with a narrow basal yellowish

band and a few dorsoapical yellowish scales, tarsomere 5 completely covered

with yellowish scales, III with tarsomere 1 having a few dorsoapical pale

scales, tarsomeres 2-4 each with a narrow pale basal band and a few dorso-

apical pale scales, tarsomere 5 completely pale scaled. Wing. Dorsal veins

covered with moderately broad brown scales; costa with a patch of broad sil-

very scales at base; ventral veins brown scaled; alula with narrow brown

scales along fringe; 2 remigial bristles. Halter. Pedicel pale, capitellum

brown scaled. Abdomen. Terga brown with a few dorsobasal pale scales form-

ing narrow bands on IJI-VI, a small triangular dorsobasal pale spot on VII;

tergum I with a rectangular patch of white scales on laterotergite; tergalII-VII

with large laterobasal white spots; sterna with pale basal scales and brown

apical ones, apical brown band becomes broader on posterior sterna; terga

and sterna with numerous golden bristles, mostly along posterior margins.

Genitalia (Fig. 21). Tergum VII index 1.10-1.30; sternum VIII index 0. 79-

0.95; tergum IX bilobed with 4-5 bristles on each lobe, index 1. 20-1. 47; in-

sula tongue-like, covered with minute setae and with 3-5 small tuberculi on

apical 0.20; lower vaginal lip narrow, moderately pigmented, covered with

minute setae; upper vaginal lip narrow, heavily pigmented, covered with

minute setae; upper vaginal sclerite medium to large sized, moderate to

heavily pigmented, triangular shaped; postgenital lobe moderately long, apex

with a moderately deep median indentation, 6-8 bristles on each side of mid-

line, covered with minute setae, dorsal PGL index 0.95-1.22, ventral PGL

index 2.30-2. 83; cercus long, 0.75-1.00 extended from segment VII, narrow,

apex sharply rounded, numerous bristles on dorsal and lateral surfaces, index

3.21-3.88, cercus/dorsal PGL index 3. 88-4. 53; 3 spherical, pigmented semin-

al capsules, 1 large and 2 tiny rudimentary ones.

MALE (Fig. 4). Similar to female in general habitus. Head. Maxil-

lary palpus brown, longer than proboscis by length of apical segment. Thorax.

Antepronotum with a few broad silvery scales. Legs (Fig. 17). Posttarsi I-III

each with 2 ungues, I, II with ungues unequal, each bearing a tooth, III unequal,

simple. Abdomen. Tergum I with a lateral band of silvery-white scales on

laterotergite; terga II[-VII with basal bands of white scales; segment VIII re-

moved with terminalia and coloration lost. Genitalia (Fig. 37). Tergum IX

Slightly bilobed with 3-4 bristles on each lobe, entire surface covered with

minute spicules; gonocoxite long and moderately broad with short bristles

scattered over dorsal surface, long stout bristles along outer lateral margin

from base to apex, ventral surface with long stout bristles on distal 0.55, most

numerous along sternomesal margin, scattered short ones on proximal 0. 45,

scattered scales on lateral and ventral surfaces; gonostylus with pedicel narrow

to moderately broad, distal 0.50 expanded with a lateroapical horn-like flap

bearing numerous short fine hairs, mesal margin of expanded portion with a

basal short, flattened pigmented gonostylar claw and 4-5 short accessory claws,

4-6 moderately long bristles at apex, 3-8 short hairs on the tergal surface;

Reinert: Aedes (Aedimorphus) in Southeast Asia 37

basal mesal lobe short and rounded apically, apical 0.30 with 4-5 short bris-

tles, a long narrow lightly pigmented filament extending from apex to near

base of gonostylus and attached to mesal membrane of gonocoxite, entire sur-

face of basal portion covered with short hair-like spicules; proctiger short,

paraproct with a small subapical thumb-like process, cercal setae absent;

phallosome with aedeagus of type I with 2 lateral plates connected basally,

each plate with 6-7 short blunt lateral teeth on distal 0.50 and covered with a

dorsal flap, paramere long, approximately 0.90 length of lateral plates; stern-

um IX large, entire surface covered with minute spicules, 4 bristles near the

center.

PUPA and LARVA. Not known.

TYPE DATA. There are 1 male and 1 female syntypes of Aedes lowisii

in the British Museum (Natural History). The syntype female of Aedes (Aedi-

morphus) lowisti is hereby designated lectotype and bears the following data:

Reedomyia lowisii Theobald, Andaman Islands, INDIA, Lowis. The male syn-

type (only the genitalia in a drop of balsam on a plastic square attached to the

pin with the data labels remains) is hereby designated as allolectotype and

bears the same data as the lectotype female. Aedes (Aedimorphus) mindoroen-

sis Knight and Hull, holotype male and 4 paratype females, Calapan (errone-

ously printed "'Calopan"’ on label), Mindoro Island, PHILIPPINES, 1 February

1916, Bottcher, in British Museum (Natural History).

I have compared the types of lowisii and mindoroensis and cannot find

any apparent differences, therefore I am placing mindoroensis in synonymy to

lowisit.

DISTRIBUTION. Specimens examined--3 males and 20 females from

the following locations:

AUSTRALIA. N.E. New Guinea, Morobe, Lae.

INDIA. Andaman Islands.

INDONESIA. Celebes, Paloe, Hadjene; Moluccas, Morotai; C. Sula-

wesi, Lambarese; Ambon, Paso.

PHILIPPINES. Mindoro, Calapan.

Other distribution.

INDONESIA. Sumatra (Brug 1926: 529).

MALAYSIA. Savawak (Moulton 1914: 47),

The record of lowisiz from Ceylon by Theobald (1910b: 260) was based

on a misidentification and should be for jamesi (Barraud 1934: 250) and the

record from this country by Senior-White (1927: 69) should be for pipersalatus

(Carter and Wijesundara 1948: 141).

TAXONOMIC DISCUSSION. The adult habitus and female and male

genitalia of lowisii are very similar to alboscutellatus and are discussed under

that species. The most distinctive features of the adults are: scutellum with

broad silvery scales on each lobe; postspiracular area without scales; sub-

spiracular area with only short fine hairs; paratergite bare; tarsi banded with

pale scales, tarsomere 5 yellow scaled; and female with 1 large and 2 rudi-

mentary seminal capsules.

BIOLOGY. Larvae were collected in shallow water in a primary for-

est and adults were taken biting man and resting in a cowshed in Celebes

(larval information on adult labels). Adults were also collected from a Ma-

laise trap in Indonesia.

38 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

AEDES (AEDIMORPHUS) MEDIOLINEATUS (THEOBALD)

(Figs. 5, 16, 17, 22, 33, 38, 52, 63)

Culex trilineatus Theobald 1901, Monogr. Cul. 2: 105 (?*); Giles 1902, Handb.

2nd Ed., p. 464 (2); Blanchard 1905, Moust., p. 330 (9); Theobald

1905, Genera Insec., Fasc. 26: 27; Theobald 1910, Monogr. Cul. 5:

359; Brunetti 1912, Rec. Indian Mus. 4: 476.

Culex mediolineatus Theobald 1901, Monogr. Cul. 2: 113 (9); Giles 1902,

Handb., 2nd Ed., p. 431(9); Blanchard 1905, Moust., p. 369(9);

Theobald 1905, Genera Insec. Fasc. 26: 27; Brunetti 1907, Rec.

Indian Mus. 1: 349.

Ochlerotatus mediolineatus Theo., Edwards 1913, Bull. ent. Res. 4: 228;

Brunetti 1920, Rec. Indian Mus. 17: 137; Senior-White 1923, Cat.

Indian Insects, Cul., p. 79.

Aedes (Ecculex) mediolineatus Theobald, Edwards 1922b, Indian J. med. Res.

10: 467. :

Aedes (Aedimorphus) mediolineatus (Theo.), Barraud 1928, Indian J. med.

Res. 15: 665 (o’*, 9); Stone et al. 1959, Thomas Say Found. 6: 194;

Kurihara 1965, Jap. J. sanit. Zool. 16: 21(A*).

Aédes (Aédimorphus) mediolineatus Theo., Borel 1930, Coll. Soc. Path.

exot. Monogr. 3: 268 (o"*, 9, L*); Edwards 1932, Genera Insec.,

Fasc. 194: 171.

Aédes (Aédimorphus) mediolineatus (Theobald), Barraud 1934, Fauna Brit.

India, Diptera 5: 263 (o’*, 9).

FEMALE (Fig. 5). Head. Antenna dark brown, approximately 0.94

length of proboscis, pedicel pale with a few small brown scales and a patch of

short fine brown hairs mesally, flagellomere 1 with a few small pale brown

scales; clypeus brown, bare; maxillary palpus golden scaled, approximately

0. 22 length of proboscis; proboscis golden scaled with apical 0.25 dusky,

approximately 1.22 length of femur I; vertex with dorsum covered with narrow

curved decumbent golden scales; lateral surface covered with broad golden

scales, some specimens also with a small anterodorsal dark spot; numerous

golden-brown erect forked scales on occiput and vertex extending anteriorly

to ocular line, erect scales somewhat darker on lateral margins of occiput.

Thorax. Scutal integument reddish-brown; scutum covered with narrow curved

reddish-brown scales, narrow curved white scales forming a pair of stripes

on dorsocentral areas extending from anterior scutal fossal area to scutellum,

similar scales on supra-alar areas from scutal angle to posterior of wing base

and on anterior and lateral margins of prescutellar space (scales nearly cover-

ing this area), narrow curved golden scales forming a stripe on acrostichal

area extending from median anterior promontory area to posterior medial

scutal area; scutellum with a patch of narrow curved golden scales on each lobe;

median anterior promontory, acrostichal, dorsocentral (anterior and posterior),

scutal fossal (anterior, 3-5 lateral, 2-3 median and 1-2 posterior), supra-alar,

several posterior medial scutal, 1 postalar callar and scutellar (lateral and

median) bristles golden-brown and well developed; pleural integument light

brown; antepronotum covered with narrow curved golden-white scales, several

golden bristles; postpronotum covered with narrow curved scales, a few reddish-

brown ones dorsally and remainder golden-white, 6-8 golden bristles; pro-

pleuron with long moderately broad and narrow golden-white scales, several

golden bristles; postspiracular area with a patch of narrow curved and a few

moderately broad golden-white scales, 7-10 golden bristles; subspiracular

Reinert: Aedes (Aedimorphus) in Southeast Asia 39

area with a small patch of narrow curved golden-white scales; mesepisternum

with an upper and a posterior patch of broad golden-white scales, several up-

per and posterior golden bristles, lower ones shorter; prealar knob with a few

narrow golden-white scales, several golden bristles; paratergite covered with

narrow curved golden-white scales; mesepimeron with a patch of broad golden-

white scales and several golden bristles on upper area; other pleural areas

bare. Legs (Fig. 16). Coxae I-III each with several golden bristles, I, II each

with anterior surface covered with broad golden-white scales, III with a few

anteroventral golden-white scales; trochanters I-III each with broad white

scales; femora I-III each with a small dorsoapical spot of white scales, I with

anterior surface white with a few intermixed light brown scales ventrally, II

with anterior surface brown with a few white scales intermixed on apical 0. 25,

III with anterior surface white with an anterodorsal brown stripe on distal

0.75, stripe broader apically, I-II with posterior surfaces white, I with a ven-

tral brown stripe from near base to apex, II with a few pale brown scales on

distal 0.25, III with a triangular patch of brown scales on dorsoapical 0. 20;

tibiae I-III white, I with a dorsoanterior longitudinal brown stripe, II with a

ventral longitudinal brown stripe, II with a dorsal and a ventral longitudinal

brown stripe and a small dorsoapical white spot; tarsi I-III brown, I with an

anterior and a posterior longitudinal white stripe on tarsomeres 1-2, occasion-

ally on 3, II with tarsomere 1 with numerous white scales intermixed with

brown ones and a dorsobasal white spot, tarsomere 2 with a few white scales

intermixed with brown ones, tarsomeres 1-3 with a posterior longitudinal

white stripe and occasionally on tarsomere 4, III with a dorsobasal white spot

on tarsomere 1, an anterior and a posterior longitudinal white stripe on tarso-

mere 1 and basal 0. 50 of tarsomere 2; posttarsi I-III each with 2 ungues I, II

equal, each bearing a tooth, III equal, simple. Wing. Dorsal veins covered

with moderately broad brown scales; costa with broad whitish scales along

basal 0.33 of its posterior margin; dusky scales on subcosta; ventral veins

brown with white scales along basal 0.33 of posterior margin of costa, similar

scales on basal 0.33 of subcosta; alula with narrow brown scales along fringe;

2 remigial bristles. Halter. Pedicel pale, capitellum golden-white scaled.

Abdomen. Tergum I covered with golden-white scales, laterotergite with a

rectangular patch of whitish scales; terga II-VI light brown, each with a broad

median longitudinal golden-white stripe and a narrower longitudinal golden-

white stripe on lateral margins, V, VI often with brown areas reduced; tergum

VII covered with golden scales; sterna covered with golden-white scales; terga

and sterna with numerous golden bristles, mostly along posterior margins.

Genitalia (Fig. 22). Tergum VIII index 0.96-1.29; sternum VIII index 0. 82-

1,08; tergum IX bilobed with 7-16 bristles on each lobe, index 1.13-1.50; in-

sula tongue-like, covered with minute setae and with 4-6 small tuberculi on

apical 0.25; lower vaginal lip narrow, moderate to heavily pigmented, cover-

ed with minute setae; upper vaginal lip narrow, heavily pigmented, covered

with minute setae; upper vaginal sclerite large, heavily pigmented, triangular

Shaped; postgenital lobe moderately long, apex with a moderately deep median

indentation, 5-10 bristles on each side of midline, covered with minute setae,

dorsal PGL index 0.96-1.26, ventral PGL index 2.07-2.31; cercus moderately

long, 1.00 extended from segment VII, narrow, apex rounded, numerous bris-

tles on dorsal and lateral surfaces, index 2.54-2.76, cercus/dorsal PGL in-

dex 3.59-4,11; 3 spherical, pigmented seminal capsules, 1 large and 2 slightly

smaller ones.

MALE (Fig. 5). Similar to female in general habitus. Head. Maxil-

lary palpus golden scaled, segments 2-5 each with a small dorsoapical brown

40 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

spot, longer than proboscis by 0.50 length of apical segment. Thorax. Post-

spiracular area without scales; subspiracular area with a few broad white

scales. Legs (Fig. 17). Posttarsi I-III each with 2 ungues, I, II with ungues

unequal, each bearing a tooth, II equal, simple. Abdomen. Tergum I white

with a median brown scale patch, laterotergite with a patch of whitish scales;

terga II-VII brown each with a dorsobasal triangular patch of golden-white

scales on lateral margins, reaching from base to apex, VII often nearly en-

tirely golden scaled; tergum VIII white scaled; sterna golden-white scaled,

VIII with a median patch of brown scales. Genitalia (Fig. 38), Tergum IX

strongly bilobed with 6-8 bristles on each lobe, entire surface covered with

minute spicules; gonocoxite long and moderately broad with short bristles

scattered over entire dorsal surface, lateral surface with long stout bristles

from base to apex, ventral surface with long stout bristles on distal 0.45,

scattered moderately long bristles mesally below long ones, somewhat more

numerous along sternomesal margin, scattered scales on lateral and ventral

surfaces; gonostylus with pedicel moderately broad, distal 0.50 expanded with

a lateroapical horn-like flap bearing a short fine subterminal hair, mesal mar-

gin of expanded portion with a moderately long, flattened, pigmented gonosty -

lar claw and a short fine hair near its base, 3 short fine hairs along apex and

3-5 short fine hairs scattered over tergal surface, a lateroapical thumb-like

process covered with numerous short fine hair-like spicules; basal mesal

lobe short and rounded apically, apical 0.50 with 4-6 short bristles, entire

surface covered with short hair-like spicules; proctiger short, paraproct

with a subapical thumb-like process, cercal setae absent; phallosome with

aedeagus of type I with 2 lateral plates connected basally, each plate with 6-7

short blunt lateral teeth on distal 0.40 and covered with a dorsal flap, paramere

long, approximately 0.85 length of lateral plate; sternum IX large, entire sur-

face covered with minute spicules, 2-3 bristles near the center.

PUPA (Fig. 52). Chaetotaxy as figured and recorded in Table 4.

Cephalothorax. Hair 5-C with 3-5 branches; 7-C with 2-5 branches; 8-C with

2-6 branches. Respiratory trumpet. Moderately pigmented; index 3. 74-4. 56,

average 4.25. Metanotum. Hair 10-C with 11-24 branches; 12-C with 4-5

branches. Abdomen. Hair 5-I with 8-17 branches; 1-II with 17-28 branches;

4-II with 6-10 branches; 1-III with 7-18 branches; 6-VI with 2-4 branches;

1-VII with 3-6 branches; 6-VII with 8-12 branches; 9-VII with 5-11 branches;

11-VII single to triple. Paddle. Ovoid; with very minute serrations along

basal 0.55 of outer margin; midrib does not reach apex; hair 1-P short, single

or double; index 1.10-1.35, average 1. 26.

LARVA (Fig. 63). Chaetotaxy as figured. Head. Hairs 1, 3, 14-C

single; 4, 12-C with 5-9(6) branches; 5, 6-C with 4-6(5) branches; 7-C with

8-11(10) branches; 8-C double or triple (2); 9, 10-C double or triple (3); 11-C

with 3-6(5) branches; 13-C with 3-5(3) branches; 15-C with 2-5(3) branches;

basal maxillary hair single; mental plate with 22-23(23) teeth. Antenna.

Heavily pigmented; numerous stout spicules scattered over shaft, longer at

about middle; hair 1-A with 6-13(8) branches, inserted at 0.36-0.48 from

base; 2-A long; 3-A approximately equal in length of 2-A. Thorax. Hair 0-P

with 6-12(7) branches; 1, 5, 6, 10, 12-P single; 2-P single or double (2); 3-P

with 3-4(4) branches; 4, 8, 9-P double; 7-P triple; 11-P double or triple (2);

14-P single or double (1); 1-M with 2-5(2) branches; 2-M single or double (2);

3, 11-M single or double (1); 4-M with 2-4(3) branches; 5, 7, 10, 12-M single;

6-M with 5-10(8) branches; 8-M with 6-12(7) branches; 9-M with 6-9(8) bran-

ches;13-M with 3-6(5) branches; 14-M with 6-10(9) branches; 1-T single to

triple (1); 2-T with 2-5(3) branches; 3-T with 6-11(8) branches; 4-T with 3-4(3)

Reinert: Aedes (Aedimorphus) in Southeast Asia 41

branches; 5, 10-T single; 6-T single or double (2); 7-T with 8-14(11) branches;

8-T with 3 6(4) branches; 9-T with 6-9(6) branches; 11, 12-T single or double

(1); 13-T with 5-14(7) branches. Abdomen. Hairs 0, 14-VII single; 1-VIII

with 4-9(5) branches; 2, 4-VIII double or triple (2); 3-VIl with 8-18(15) bran-

ches; 5-VIII with 8- 11( 10) branches; 6-V-VI short; comb with 9-19(15) scales

arranged in 2 irregular rows, scales with a long stout pointed median spine

and short denticles along lateral margins; 1-X with 3-4(4) branches; 2-X with

12-15(14) branches; 3-X single; ventral brush varies from 8 hairs on grid and 4

precratal ones to 9 ‘hairs on grid and 3 precratal ones, usually with 8 hairs on

grid and 4 precratal ones; saddle moderately pigmented with minute ridges,

incompletely rings segment, with a few spicules along posterior margin and

small ridges over entire saddle, acus present; 4 anal papillae, very long and

Slender. Szphon. Moderately pigmented with minute ridges over entire sur-

face; acus present; index 5.93-7.01; pecten with 10-14(12) teeth, apical 3-4

teeth smooth and wider spaced than remainder which have a slender apical

attenuated filament with 1-2 lateral denticles; hair 1-S with 5-7(5) branches,

inserted at 0. 72-0. 78 from base.

EGG (Fig. 33). Shape. Fusiform; greatest diameter near middle.

Size. Length 735-840 microns; width 190-200 microns. Color. Brown.

Chorion. Reticulation on entire egg composed of polygonal cells of varied

size; cell walls raised. Remarks. The description of the egg is based on 5

mature eggs extracted from the abdomen of a museum specimen which pos-

sessed the following collection data on the labels: THAILAND, Lampang

Province, Wat Luang, 15 May 1968, collection number 02699, collector

Bruce A. Harrison and genitalia preparation number T72. 137.

TYPE DATA. Culex mediolineatus Theobald, holotype female,

Thayetmyo, BURMA, August, E. Y. Watson, 94-4 and Culex trilineatus holo-

type female with same data as holotype of mediolineatus, both in British Muse-

um (Natural History).

DISTRIBUTION. Specimens examined--142 males, 329 females, 82

pupae, 181 larvae and 93 individual rearings (50 pupal, 43 larval) from the

following locations:

BURMA. Thayetmyo.

CAMBODIA. Phnom-Penh.

MALAYSIA, Perlis; Kedah,

SOUTH VIETNAM. Bihn Dinh, An Khe, Long Van, Qui Nhon, Vinh

Thanh; Bien Hoa, Di An; Da Nang, Da Nang; Gia Dinh, Saigon, Tan San Nhut;

Hau Nghia, Cu Chi: Khanh Hoa, Nha Trang; Kontum, Kontum; Nha Trang, Ben

Kay; Ninh Thuan, Phan Rang; Tay Ninh, Phuoc Vinh; Thua Thien, Cu Lai; Pho

Bai, Phu Loi.

THAILAND. Chiang Mai, Ban Lang Ka, Ban Mae Yuak, Chang Kien,

Chang Puak, Chiang Mai, Doi Sutep, Muang, Thanou Doi Saket; Chon Buvi,

Bang Huai Kum, Bang Phra, Khao Phai; Khon Kaen, Ban Muang Kao, Ban

Nong Kham, Phu Wiang; Lampang, Wat Luang; Nakhon Ratchasima, Banna

Nabon, Pak Chong; Nan, Ban Huai Hap; Prachuap Khiri Khan, Tubsakea; Surat

Thani, Ban Taling Ngam, Ko Samui; Udon Thani, Ban Kau Noi, Tai Tan Kam

Phoo, Muang.

Other distribution.

CHINA. Hainan Island (Chu 1957: 158, 1958: 109).

INDONESIA. Sumatra, Djambi, Moeara Tebo (Brug and Edwards 1931:

258); Java (Barraud 1934: 264).

THAILAND, Nakhon Phanom, Takhli, Ubon (Parrish 1968b: 2); Don

Muang AB, U-Tapao AB (Reisen et al. 1971: Tables 7 and 8).

42 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

SOUTH VIETNAM (Borel 1930: 268); Phan Rang, Phu Cat, Pleiku

(Parrish 1968a: 3); Bien Hoa (Parrish 1969: 554); Cam Ranh Bay (Reisen et al.

1971: Table 12); Cam Khe, Khe Sahn, Khue Bac, Kim Lien, Nui Dat (Grothaus

et al. 1971: 20).

TAXONOMIC DISCUSSION. Aedes mediolineatus and trilineatus were

both originally described by Theobald (1901a) in volume I of A Monograph of the

Culicidae of the World. Aedes trilineatus had page priority but the name is pre-

occupied.

Aedes mediolineatus is similar in the adult habitus to pallidostriatus.

It possesses the following features: wing with anterior margin of costa dark

brown scaled; femur IL with anterior surface mainly dark scaled and anterior

surface of femur II mainly white scaled; abdomen with dorsolateral longitu-

dinal brown bands on terga II-IV; and postpronotum with 6-7 bristles, which

distinguish it from pallidostriatus which has: wing with anterior margin of

costa golden scaled; femora I-III each with anterior surface brown with a me-

dian longitudinal white stripe from base to apex; and abdomen with terga com-

pletely golden scaled. The gonostylus of the male genitalia of mediolineatus

is markedly different from that of pallidostriatus.

The pupa of mediolineatus has a very similar chaetotaxy to pallidostria-

tus and it is difficult to separate them. They usually can be separated by ab-

dominal hair 1-I which has 17-30 branches in mediolineatus and 30-46 branches

in pallidostriatus.

There are no apparent differences between the larva of mediolineatus

and pallidostriatus.

BIOLOGY. Immatures have been collected from a wide range of

habitats but the preferred site appears to be fresh water flood pools with

floating and submerged dead leaves or abundent floating and emergent vege-

tation located in secondary scrub or plains areas. Adults have been taken

feeding on man and domestic animals. Immatures in Thailand have been

collected usually from small and large flood pools, occasionally from large

rice fields, and also from edge of large pond, Huey Keo City moat, small

animal footprint, large pit and large wheeltrack; usually from colored fresh

water but also numerous times from turbid fresh and clear fresh water; water

usually with floating and submerged dead leaves or abundant floating and emer-

gent vegetation; in partial shade or unshaded areas; usually in rice fields or

pools in plains, but also from secondary scrub and coconut groves in plains,

and secondary scrub in hills and mountains; and at an altitude from 43 to 1, 580

feet (most often from 80 to 650 feet). Larvae were collected in association

with the following species of mosquitoes: Aedes alboscutellatus, caecus, culi-

cinus, ferinus, lineatopennis, vexans vexans; Anopheles barbirostris, kochi,

maculatus, philippinensis, vagus; and Culex annulus, fuscanus, fuscocephalus,

gelidus, mimulus, pseudovishnui, raptor, tritaeniorhynchus, whitmoret.

Adults have been taken biting man, cattle and horses, resting in a stable anda

house, and in a light trap.

In South Vietnam larvae were collected from ground pools, tire, artifi-

cial container, ditch, marshy depression, flood pools, swamp, rock pools,

wheel track, footprints and a rice field and adults were taken in light traps.

Macdonald (1957: 21) obtained specimens of mediolineatus that he identi-

fied as Aedes (Aedimorphus) species near pallidostriatus from human bait col-

lections in Malaysia. Adults were taken feeding on bovines and chickens (un-

published data from SEATO Medical Research Laboratory Annual Progress

Report 1967: 450) and collected in scrub or open forest at 1, 000 feet in Thai-

land (Scanlon and Esah 1965: 139, 143). Grothaus et al. (1971: 20) collected

Reinert: Aedes (Aedimorphus) in Southeast Asia 43

larvae from standing pools with emergent vegetation, rice paddies, dense

swamps and borrow pits and adults from indoor resting sites, light traps and

CO9 traps.

AEDES (AEDIMORPHUS) NIGROSTRIATUS (BARRAUD)

(Figs. 6, 16, 23, 39)

Aédimorphus nigrostriatus Barraud 1927, Indian J. med. Res. 14: 549 (o’*, 9).

Aedes (Aedimorphus) nigrostriatus Barr., Barraud 1928, Indian J. med. Res.

15666407) Bs

Aédes (Aédimorphus) nigrostriatus Barraud, Edwards 1932, Genera Insec..,

eres 171; Barraud 1934, Fauna Brit. India, Diptera 5: 262

Shee

Aedes (Aedimorphus) nigrostriatus (Barraud), Stone et al. 1959, Thomas Say

Found. 6: 195.

FEMALE (Fig. 6). Head. Antenna brown, approximately 1.04 length

of proboscis, pedicel pale with a few small yellow scales and a patch of short

fine brown hairs mesally, flagellomere 1 pale with a few small yellow scales;

clypeus light brown, bare; maxillary palpus golden scaled, approximately 0.25

length of proboscis; proboscis golden scaled with a small ventrobasal patch of

brown scales, approximately 1.04 length of femur I; vertex with dorsum cov-

ered with narrow curved decumbent yellow scales; lateral surface covered with

broad pale yellow scales; numerous light brown erect forked scales on occiput

and vertex extending anteriorly to ocular line. Thovax. Scutal integument pale

with dark reddish-brown areas forming a pair of stripes on dorsocentral areas

from anterior margin to scutellum and a spot on supra-alar area anterior to

wing base; scutum covered with narrow curved golden scales with narrow

curved reddish-brown ones on dark areas of integument; scutellum with a patch

of narrow curved golden scales on each lobe and a few reddish-brown ones on

laterobasal areas of median lobe; median anterior promontory, acrostichal,

dorsocentral (anterior and posterior), scutal fossal (anterior, 4-5 lateral, 1-2

median and 2-3 posterior), supra-alar, several posterior medial scutal, 1

postalar callar and scutellar (lateral and median) bristles pale brown‘ and well

developed; pleural integument light brown; antepronotum with narrow curved

golden scales, several golden-brown bristles; postpronotum with narrow curved

golden scales, 3-5 golden-brown posterior bristles; propleuron with broad

golden scales, several golden bristles; postspiracular area with a few narrow

golden scales, 5-7 golden bristles; subspiracular area with a small patch of

moderately broad golden scales; mesepiternum with a small upper and a pos-

terior patch of broad golden scales, several upper and posterior golden-brown

bristles, lower ones white and shorter; prealar knob with several golden bris-

tles; paratergite with a few narrow golden scales; mesepimeron with a patch of

broad golden scales and several golden bristles on upper area; other pleural

areas bare. Legs (Fig. 16), Coxae I-III each with several golden-brown bris-

tles, I, II each with anterior surface covered with broad golden scales, III with

a small anteroventral patch of similar scales; trochanters I-III each with broad

golden scales; femora I-III golden, I with a narrow posteroventral longitudinal

brown stripe from base to near apex, II with a broad anterobasal longitudinal

brown stripe from base to apical 0.20 and a narrow anterior subapical brown

band, III with a narrow anterior and posterior subapical brown band; tibiae

I-HI yellow, I with an indistinct anteroventral longitudinal brown stripe on

Ag Contrib. Amer. Ent. Inst., vol. 9, no. 5, 19'73

basal 0.50; tarsi I-III yellow; posttarsi I-III each with 2 ungues, I, II equal,

each with a tooth, III equal, simple. Wing. Dorsal veins covered with moder-

ately broad golden scales except for the following brown scaled areas: apical

0.25 of costa pale brown; remigium and basal 0. 50 of radius (a few yellow

scales on posterior margin), apical 0.50 of radial sector, radius and basal

0.50 of radius3; media from radiomedial crossvein to furcation, basal 0. 60 of

media;,and cubitus; membrane darkened in region of crossveins; ventral veins

with scaling similar to dorsal ones except subcosta and remigium completely

golden scaled; alula with narrow yellow scales along fringe; 1-2 remigial bris-

tles. Halter. Pedicel pale, capitellum golden scaled. Abdomen. Terga gol-

den scaled, I with a rectangular patch of white scales on laterotergite, II-VII

each with a laterobasal white scale patch; sterna golden scaled; terga and

sterna with numerous golden bristles, mostly along posterior margins. Gen-

italia (Fig. 23). Tergum VII index 1.01-1.11; sternum VII index 0. 78-0. 88;

tergum IX bilobed with 5-8 bristles on each lobe, index 0. 78-0. 81; insula

tongue-like, covered with minute setae and with 4-6 small tuberculi on apical

0.25; lower vaginal lip narrow, moderately pigmented, covered with minute

setae; upper vaginal lip narrow, heavily pigmented, covered with minute setae;

upper vaginal sclerite large, moderate to heavily pigmented, triangular shaped;

postgenital lobe moderately long, apex with a moderately deep median indenta-

tion, 5-7 bristles on each side of midline, covered with minute setae, dorsal

PGL index 0.97-1.03, ventral PGL index 2.04-2.10; cercus moderately long,

0.75-0. 85 extended from segment VII, narrow, apex sharply rounded, numer-

ous bristles on dorsal and lateral surfaces, index 2.08-2.28, cercus/dorsal

PGL index 3, 32-3. 71; 3 spherical, pigmented seminal capsules, 1 large and 2

slightly smaller ones.

MALE (Fig. 6). Similar to female in general habitus. Head.. Maxil-

lary palpus golden with apical segment brown, segment 4 golden dorsally and

brown laterally, segments 2-3 each with a narrow apical brown band, longer

than proboscis by 0.75 length of apical segment; proboscis golden with a longi-

tudinal brown stripe on basal 0.40 of ventral surface. Legs. Femur I also

with an anteroventral brown stripe on basal 0. 50; tibia III with an indistinct

dorsal brown stripe; tarsus I with a few light brown apical scales on tarso-

meres 3-4; posttarsi I-III each with 2 ungues, I, II with ungues unequal, each

bearing a tooth, III equal, simple. Abdomen. Terga yellow scaled; sterna

yellow scaled with a few brown scales on lateral surfaces of III, VI, a few

brown scales along posterior margins of VI, VII. Genitalia (Fig. 39). Tergum

IX bilobed with 5-7 bristles on each lobe, entire surface covered with minute

spicules; gonocoxite long and moderately broad, dorsal surface with several

short scattered hairs, mainly along tergomesal margin and a few moderately

long bristles on distal 0.25, lateral surface with a number of long stout bris-

tles from base to apex, ventral surface with a number of moderately long to

long bristles on distal 0.33 and a few scattered short proximal ones, a row of

6-8 long stout bristles along distal 0.50 of sternomesal margin, scattered

scales on lateral and ventral surfaces; gonostylus with distal 0.62 expanded

into a large mesal oblong-shaped lobe and a lateral longer, narrow, strongly

incurved, tapering horn which bears a ventral patch of long thin hairs mesally

near middle, mesal expanded lobe with 2-3 laterobasal short fine hairs, a

small mesal indentation with a curved pigmented claw attached, 3 short fine

hairs apically, a patch of long hair-like spicules proximad of claw and an

additional patch on tergal surface; basal mesal lobe short and narrow, distal

0.40 with 3 moderately long and 1 short bristle, entire surface covered with

short hair-like spicules; proctiger short, paraproct with a small subapical

Reinert: Aedes (Aedimorphus) in Southeast Asia 45

thumb-like process, cercal setae absent; phallosome with aedeagus of type I

with 2 lateral plates connected basally, each plate with 8-9 short blunt lateral

teeth on distal 0.50 and covered with a dorsal flap, paramere long, approxi-

mately 0.78 length of lateral plate; sternum IX large, entire surface covered

with minute spicules, bristles absent.

PUPA and LARVA. Not known.

TYPE DATA. There are 1 male and 1 female syntypes of Aedes

nigrostriatus in the British Museum (Natural History) and each bears the

following data: Aedimorphus nigrostriatus Barraud, Golaght, Assam, INDIA,

17 November 1925, Capt. P. J. Barraud, caught in jungle. Lectotype designa-

tion is hereby made for the syntype male and allolectotype designation is made

for the syntype female. One paratype male and 1 paratype female, with same

data as lectotype, are in the Indian Museum, Calcutta.

DISTRIBUTION. Specimens examined--2 males and 6 females from the

following locations:

BURMA. Pegu, Rangoon.

INDIA. Assam, Chabua, Golaghat.

TAXONOMIC DISCUSSION. Aedes nigrostriatus resembles Aedes tri-

maculatus but can be separated from this species by the ornamentation and

integument color of the scutum. In nigrostriatus the scutal integument is pale

with dark reddish-brown areas forming a pair of stripes on the dorsocentral

areas and a circular spot on the supra-alar area anterior to the wing base,

while trimaculatus has the scutal integument reddish-brown with the scutal

fossal areas and prescutellar space pale. The color of the scutal scales is

the same as the underlying integument on these 2 species. Aedes nigrostria-

tus also has the wing membrane darkened in regions of the crossveins while

all other Oriental species of Aedimorphus do not have this feature.

BIOLOGY. Larvae were collected from muddy pools in India (data on

adult labels).

Females in a freshly-fed condition have been collected from cowsheds

in India (Barraud 1927: 551).

AEDES (AEDIMORPHUS) ORBITAE EDWARDS

(Figs. 7, 16, 17, 24, 40, 53, 64)

Lepidotomyia Taeniata Leicester 1908, Cul. Malaya, p. 133 (co, 2); Brunetti

1912, Rec. Indian Mus. 4: 459.

Ochlerotatus taeniatus Leices., Brunetti 1920, Rec. Indian Mus. 17: 140.

Aedes orbitae Edwards 1922a (nom nov), Indian J. med. Res. 10: 260.

Aedes (Ecculex) orbitae Edwards 1922b (nom nov for taeniata Leicester, non

Wiedemann 1928), Indian J. med. Res. 10: 466,

Aédes (Aédimorphus) orbitae Edw., Edwards and Given 1928, Bull. ent. Res.

18: 344(L); Edwards 1932, GeneraInsec., Fasc. 194: 168; Edwards in

Barraud 1934, Fauna Brit. India, Diptera 5: 250 (0, 9).

Aedes (Aedimorphus) orbitae Edwards, Stone et al. 1959, Thomas Say Found.

6: 195.

FEMALE (Fig. 7). Head, Antenna dark brown, approximately 1.06

length of proboscis, pedicel pale brown with a few small brown scales and a

patch of short fine dark brown hairs mesally, flagellomere 1 pale with a few

small brown scales; clypeus dark brown, bare; maxillary palpus dark brown

scaled with apex silvery-white, occasionally a few white scales on base of

46 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

segment 4, approximately 0.19 length of proboscis; proboscis dark brown

scaled with a white ventral spot extending onto lateral surfaces approximately

0.67 from base (occasionally this spot forms a complete band), approximately

1.04 length of femur I; vertex with dorsum covered with narrow curved decum-

bent scales arranged in an anteromedian diamond-shaped dark brown group

and the remainder golden-white; lateral surface covered with broad white

scales, an anterodorsal black patch and a dark area anterior to antepronotum;

numerous brown erect forked scales on occiput and vertex extending anteriorly

to ocular line. Thorax. Scutal integument brown; scutum covered with narrow

curved reddish-black scales, narrow curved white scale patches on median an-

terior promontory area, scutal fossal areas (extending from anterior area along

margin and onto lateral area), small circular patch on posterior scutal fossal

area, supra-alar area above posterior of paratergite, similar scales along an-

terior margin of scutal ridge from scutal angle posteromesally 0.50 to dorso-

central area and scattered over area mesally to dorsocentral setae; scutellum

with a patch of broad overlapping silvery -white scales on each lobe; median

anterior promontory, acrostichal, dorsocentral (anterior and posterior), scu-

tal fossal (anterior, 4-5 lateral and 1 posterior), supra-alar, several posterior

medial scutal, 1 postalar callar and scutellar (lateral and median) bristles

reddish-black and well developed, others absent; pleural integument dark brown;

antepronotum covered with narrow curved white scales, several dark bristles;

postpronotum sparsely covered with narrow curved scales, reddish-black ones

dorsally and a few posteriorly with a patch of white ones mesally, 6-7 posterior

dark bristles; propleuron with a patch of broad silvery-white scales, several

golden bristles; postspiracular area with 4-7 golden bristles; mesepisternum

with an upper and a lower patch of broad silvery-white scales, several upper

and posterior golden bristles, lower ones shorter; prealar knob with several

golden-brown bristles; paratergite with broad silvery-white scales on lateral

surface; mesepimeron with a patch of broad silvery-white scales and golden

bristles on upper area; other pleural areas bare. Legs (Fig. 16). Coxae I-III

each with several brown bristles, I with anterior and lateral surfaces covered

with broad brown scales with a dorsal white patch, II, III each with a patch of

broad white scales on anterior surface; trochanters I-III each with broad white

scales; femora I-III each brown, with a narrow basal white band and a dorso-

apical silvery-white spot, I with a narrow anteroventral white stripe on basal

0.33 in some specimens, III with an anteroventral longitudinal white stripe from

base to apex, I-III each with posterior brown, I with a posterodorsal white stripe

from base to apex, II, III each with a posteroventral white stripe, broad basally

and tapering to apex; tibiae I-III brown, each with a few ventrobasal pale scales,

I with a dorsoapical silvery-white spot, III with a few lateroapical silvery-white

scales, I with a posteroventral longitudinal white stripe from base to apex, II,

III each with a posteromedian longitudinal white stripe, II with stripe from base

to near apex, III with stripe on apical 0. 65 (stripe on III absent or reduced in

some specimens); tarsi I-III brown, I with tarsomeres 2, 3 each with a dorso-

basal white spot, II with tarsomeres 1-3 each with a dorsobasal white spot, a

few dorsoapical white scales on tarsomere 1, III with tarsomeres 1-4 each with

a broad basal white band, and a few dorsoapical white scales, tarsomere 5

white scaled; posttarsi I-IIL each with 2 ungues, I,II equal, each bearing a tooth,

Ill equal, simple. Wing. Dorsal veins covered with moderately broad brown

scales; costa with a patch of white scales at base; ventral veins brown scaled;

alula with narrow brown scales along fringe; 2 remigial bristles. Halter.

Pedicel pale, capitellum white scaled with a few brown scales at base. Abdo-

men. Tergum I brown with a rectangular patch of silvery-white scales on

Reinert: Aedes (Aedimorphus) in Southeast Asia 47

laterotergite; terga II-VII brown dorsally (a few basomedian pale scales on II-

IV in some Malayan specimens), each with a lateromedian white scale patch,

sometimes extending to base along lateral margins on III-VII; sterna white

scaled with a brown posterior band on sterna II-VI; terga and sterna with nu-

merous golden bristles, mostly along posterior margins. Genitalia (Fig. 24).

Tergum VIII index 0. 89-1.08; sternum VIII index 0. 78-0. 88; tergum IX bilobed

with 5-8 bristles on each lobe, index 0.93-0.95; insula tongue-like, covered

with minute setae and with 3-5 small tuberculi on apical 0.25; lower vaginal

lip narrow, moderately pigmented, covered with minute setae; upper vaginal

lip narrow, heavily pigmented, covered with minute setae; upper vaginal

sclerite large, heavily pigmented, triangular shaped; postgenital lobe moder-

ately long, apex with a deep median indentation, 4-8 bristles on each side of

midline, covered with minute setae, dorsal PGL index 1.14-1.26, ventral PGL

index 1.93-1.96; cercus moderately long, completely extended from segment

VII, narrow, apex sharply rounded, numerous bristles on dorsal and lateral

surfaces, index 2.50-2.81, cercus/dorsal PGL index 3. 18-3. 21; 3 spherical,

pigmented seminal capsules, 1 large and 2 slightly smaller ones.

MALE (Fig. 7). Similar to female in general habitus. Head. Maxil-

lary palpus brown with segments 3-5 each with a basal white band, segment 2

with a dorsal white spot near middle, longer than proboscis by length of apical

segment; proboscis with median white spot forming a band; vertex with median

narrow brown scaled stripe reduced. Legs (Fig. 17). Tibia III without pos-

teromedian white stripe; posttarsi I-III each with 2 ungues, I, II with ungues

unequal, each bearing a tooth, III equal, simple. Abdomen. Terga HI-VII

each with a basomedian patch of white scales in addition to lateromedian white

patches (basomedian patches small in some specimens), patch small on VU;

tergum VIII and sternum VIII completely white scaled. Genitalia (Fig. 40).

Tergum IX strongly bilobed with 6-8 bristles on each lobe, entire surface

covered with minute spicules; gonocoxite long and moderately broad, dorsal

surface with short fine hairs along tergomesal margin and a few moderately

long bristles at median forming somewhat of a longitudinal line on basal 0. 50,

short bristles on remainder of area, lateral margin with long stout bristles

from base to apex, ventral surface with long stout bristles on distal 0. 45,

most numerous on sternomesal margin, scattered short to moderately long

bristles over remainder of area, scattered scales on lateral and ventral areas;

gonostylus with pedicel long, narrow and somewhat incuved, distal 0.38 ex-

panded into a mesal lobe and a lateral, narrow, incurved, apically pointed

horn attached approximately 0.78 from base with a short fine hair at apex,

mesal expanded lobe with a moderately long, flattened, curved, and apically

blunt gonostylar claw attached mesally near middle, 3-5 long stout bristles

along apical margin, mesal 1-2 bristles shorter and others equal in length,

3 short fine hairs along mesal margin of distal 0.30 of pedicel; basal mesal

lobe short and rounded apically, distal 0.50 with 12-14 short bristles; entire

surface covered with short hair-like spicules; proctiger short, paraproct with

a subapical thumb-like process, cercal setae absent; phallosome with aedeagus

of typeI with 2 lateral plates connected basally, each plate with 4-5 short blunt

lateral teeth on distal 0.60 and covered with a dorsal flap, paramere long,

approximately 0.89 length of lateral plate; sternum IX large, entire surface

covered with minute spicules, 5-10 bristles near center.

PUPA (Fig. 53). Chaetotaxy as figured and recorded in Table 5. A

patch of spicules on metanotum between hairs 12-C and one on abdomen be-

tween hairs 1-I. Cephalothorax. Hair 5-C with 3-6 branches; 7-C with 3-5

branches; 8-C with 7-9 branches. Respivatory trumpet. Lightly pigmented;

48 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

index 3.30-4.06, average 3.67. Metanotum. Hair 10-C with 17-33 branches;

12-C with 5-10 branches. Abdomen. Hair 5-I with 7-16 branches; 1-II with

33-65 branches; 4-II with 3-6 branches; 1-III with 8-17 branches; 6-VI single;

1-VII with 3-6 branches; 6-VII with 6-11 branches; 9-VII with 4-6 branches;

11-VII single. Paddle. Ovoid; with very minute serrations along basal 0.55

of outer margin; midrib does not reach apex; hair 1-P short, single; index

1.05-1.32, average 1.17.

LARVA (Fig. 64). Chaetotaxy as figured. Head. Median mouth

brushes pectinate apically; hairs 1, 3, 14-C single; 4-C with 7-13(10) branches;

5-C with 6-9(8) branches; 6, 13-C with 6-9(7) branches; 7-C with 9-16(13) bran-

ches; 8-C double or triple (2); 9-C with 2-4(3) branches; 10-C double or triple

(3); 11-C with 5-8(5) branches; 12-C with 5-10(8) branches; 15-C with 3-5(4)

branches; basal maxillary hair single; mental plate with 27-29(28) teeth. An-

tenna. Lightly pigmented; scattered small spicules over entire shaft, more

numerous on basal 0.50; hair 1-A with 3-5(4) branches, inserted at 0. 49-0. 52

from base; 2-A long; 3-A approximately 0.34 length of 2-A. Thovax. Hair

0-P with 9-21(15) branches; 1, 5, 6, 10, 12-P single; 2, 9, 11, 14-C double

or triple (2); 3, 7-P with 3-4(3) branches; 4-P single to triple (2); 8-P double

or triple (3); 1-M with 3-8(5) branches; 2-M single to triple (2); 3, 5, 7, 10,

12-M single; 4-M with 3-6(4) branches; 6-M with 5-9(7) branches; 8-M with

7-11(8) branches; 9, 14-M with 6-10(7) branches; 11-M single or double (1);

13-M with 5-10(6) branches; 1, 12-T single to triple (1); 2-T with 3-6(5) bran-

ches; 3-T with 8-18(11) branches; 4-T with 2-5(3) branches; 5, 10-T single;

6-T double or triple (2); 7-T with 8-13(10) branches; 8-T with 4-8(6) branches;

9-T with 5-8(6) branches; 11-T single or double (1); 13-T with 5-11(9) branches.

Abdomen. Hairs 1, 2-VIII on common basal plate; hairs 0, 14-VIII single;

1-VII with 6-10(7) branches; 2-VIII double or triple (3); 3-VIII with 13-27(15)

branches; 4-VIII single to triple (1); 5-VIII with 6-10(9) branches; 6-V-VI short;

comb with 20-30(27) scales arranged in 3 irregular rows, scales moderately

long and rounded apically with stout denticles on lateral margins and apex; 1,

3-X single; 2-X with 8-14(9) branches; ventral brush varies from 9 hairs on

erid and 3 precratal ones to 10 hairs on grid and 2 precratal ones, usually

with 10 hairs on grid and 2 precratal ones; saddle moderately pigmented with

minute ridges, incompletely rings segment (covers approximately 0.50 of seg-

ment), with numerous spicules along posterior margin and small ridges over

entire saddle, acus absent; 4 anal papillae, long and slender, tapering to an api-

cal point. Siphon. Moderately pigmented with minute ridges over entire sur-

face; with a small dorsoapical and ventroapical patch of spircules (patches do

not join on lateral surface); a large patch of spicules on lateral surface near

middle; acus absent; index 3. 40-4. 67; pecten with 18-22(18) teeth, apical 2-3

teeth smooth or with a very tiny median denticle and wider spaced than remain-

der which have a Slender apical attenuated filament with 1-3 lateral denticles;

hair 1-S with 4-7(5) branches, inserted 0. 80-0. 85 from base.

TYPE DATA. There are 2 male and 4 female syntypes of Aedes orbitae

in the British Museum (Natural History). Lectotype female of Aedes (Aedimor-

phus) orbitae is hereby designated and bears the following data: Lepidotomyia

taeniata Leicester, Kuala Lumpur, FED. MALAY STATES, 6 February 1903,

Dr. G. F. Leicester, Culex Alboscutellata var. Annul, bred larvae from mud-

dy water in rut made by wagon wheel in wagon track into jungle 5 3/4 miles

Pakang Rd., Kuala Lumpur; allolectotype male is also hereby designated and

bears the following data: same as lectotype except var. Annul absent and date

is 25 February 1903; 1 paralectotype male with same data as allolectotype; 2 para-

lectotype females with same data as lectotype except one has date of 8 February

Reinert: Aedes (Aedimorphus) in Southeast Asia 49

1903; and 1 paralectotype female with the following data: Culex trifeliat, 31

January 1903, taken on wagon rd., high patch jungle Pahang Rd. 5 3/4 miles,

remainder of data same as lectotype. There are in the United States National

Museum (Natural History) an additional male and a female syntypes with the

same data as the lectotype except the date is 8 February 1903. These 2 speci-

mens are hereby designated paralectotypes.

DISTRIBUTION. Specimens examined--63 males, 69 females, 84 pupae,

128 larvae and 84 individual rearings (53 pupal, 31 larval) from the following

locations:

MALAYSIA. North Borneo, Forest Camp, Kalabakan, Ta Waw; Pahang,

Chegar Perah, Gunong Benom; Perak, Chior F. R.; Perlis, Bt. Bintang F. R.;

Sabah, Tenom; Selangor, Kota Belud, Kuala Lumpur, Sabak Ulu Gombak.

SINGAPORE.

THAILAND. Nakhon Si Thammarat, Ban Rim Thanon, Chang Khao,

Thung Song; Narathiwat, Khau Lau; Phangnga, Kh. Pak Chaung, Nam Tai;

Prachin Buri, Ban Thap Lan; Ranong, Kraburi; Yala, Kg. Yala Bong.

Other distribution.

SINGAPORE. Gunong Pulai (Edwards 1928: 344).

TAXONOMIC DISCUSSION. Aedes orbitae resembles jamesi and lowisii

in the adult habitus. It can be easily distinguished from these species by the

bare subspiracular area, shape of the gonostylus of the male genitalia and

presence of 1 large and 2 slightly smaller seminal capsules in the female geni-

talia. The male genitalia of orbitae are similar to caecus and punctifemoris.

It is easily separated from these two species by having 12-14 bristles on the

basal mesal lobe while caecus has 5-8 bristles and punctifemorvis has 4-6 bris-

tles. Other distinctive features of the adults are: scutellum with broad sil-

very scales on each lobe; postspiracular area without scales; and tarsi banded

with white scales, tarsus III with tarsomere 5 white scaled.

The pupa is characterized by the following features: trumpet broad;

hair 10-C with 17-33 branches; hair 1-II with 33-65 branches; hair 7-V long

and single; and hairs 10-IV, V long and single.

The larva resembles caecus from which it can be separated by: a

lateral patch of spircules at about middle of siphon; hairs 1, 2-VIII on common

basal plate; and hair 6-C with 6-9 branches, while caecus has: a dorsal and

ventral patch of spicules near middle of siphon; hairs 1, 2-VII not on a plate;

and hair 6-C with 4-5 branches. Other distinctive features of orbiiae are:

hair 4-C with 7-13 branches; hair 13-C with 6-9 branches; hairs 13-III-V well

developed; and comb with 20-30 scales.

BIOLOGY. The immatures are usually collected from shaded elephant

footprints and small flood pools containing turbid fresh water located in pri-

mary and secondary forests in mountainous areas. Adults have been taken

biting man. Immatures in Thailand have been collected from large elephant

footprints, small animal footprints, large flood pool, small ditch and small

stream pool; usually in turbid fresh water but occasionally in clear fresh

water; water with floating and submerged vegetation or with scarce aquatic

vegetation; usually in partial shade but occasionally in heavy or unshaded

areas; usually in primary and secondary rain forests in mountains, occasionally

in rubber plantations in hilly areas and secondary rainforest in the plains; and

at an altitude of 7 to 2, 300 feet (most often from 200 to 500 feet). Larvae were

collected in association with the following species of mosquitoes: Aedes caecus,

macfarlanei; Anopheles balabacensis, barbirostris, hackeri, introlatus, kochi,

maculatus, montanus, riparis, subpictus; Culex bitaeniorhynchus, mimulus;

and Uranotaenia bicolor. Adults were taken biting man.

50 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

In Malaysia, larvae were collected from temporary pools on muddy

eround in inland forests and adults taken biting man by Macdonald (1957: 21).

In this country, Macdonald and Traub (1960: 100) also obtained immatures

from cart tracks and hoofprints and Leicester (1908: 134) collected larvae from

ruts in a jungle wagon track.

AEDES (AEDIMORPHUS) PALLIDOSTRIATUS (THEOBALD)

(Figs. 8, 16, 17, 25, 41, 54, 65)

Culex pallidostriatus Theobald 1907, Monogr. Cul. 4: 410 (o'*, 9); Brunetti

1912, Rec. Indian Mus. 4: 473.

Culex pavascelos Theobald 1910, Rec. Indian Mus. 4: 18 (2); Theobald 1910,

Monogr. Cul. 5: 379; Brunetti 1912, Rec. Indian Mus. 4: 473.

Ochlerotatus ochraceus Theo., Edwards 1911, Bull. ent. Res. 2: 250 (Oriental

records only).

Ochlerotatus pallidostriatus (Theo.), Edwards 1913, Bull. ent. Res. 4: 228;

Brunetti 1920, Rec. Indian Mus. 17: 140.

Aedes (Ecculex) pallidostriatus Theobald, Edwards 1922b, Indian J. med. Res.

10: 467.

Ochlerotatus pallidostriatus Theobald, Senior-White 1923, Cat. Indian Insects,

Cula. pi ols

Aedes (Aedimorphus) pallidostriatus (Theo.), Barraud 1928, Indian J. med. Res.

15: 665 (o"*, 2); Stone et al. 1959, Thomas Say Found. 6: 196; Qutubuddin

1960, Mosquito News 20: 358 (c’).

Aédes (Aédimorphus) pallidostriatus Theobald, Edwards 1932, Genera Insec.,

Fasc. 194: 171.

Aédes (Aédimorphus) pallidostriatus (Theobald), Barraud 1934, Fauna Brit.

India, Diptera 5: 261 (o"*, 2, L*).

FEMALE (Fig. 8). Head. Antenna dark brown, approximately 0.91

length of proboscis, pedicel pale with a few small dusky scales and a patch of

short fine brown hairs mesally, flagellomere 1 pale with a few small pale

scales; clypeus pale, bare; maxillary palpus golden scaled with short dark

brown bristles, approximately 0.20 length of proboscis; proboscis golden

scaled with apical 0.25 dusky, approximately 1.11 length of femur I; vertex

with dorsum covered with narrow curved golden decumbent scales; lateral

surface covered with broad golden-white scales; numerous golden erect forked

scales on occiput and vertex extending anteriorly to ocular line. Thorax. Scu-

tal integument light reddish-brown; scutum covered with narrow curved bronzy

scales, narrow curved whitish-golden scales forming a pair of stripes on dorso-

central areas from anterior scutal fossal areas to scutellum and a less distinct

stripe on acrostichal area from anterior promontory area to posterior medial

scutellar area, similar scales covering supra-alar area from scutal angle to

posterior of wing base, posterior medial scutal area and prescutellar space;

scutellum with narrow curved whitish-golden scales on each lobe; median an-

terior promontory, acrostichal, dorsocentral (anterior and posterior), scutal

fossal (anterior, 2-4 lateral, 1-3 median and 2-3 posterior), supra-aiar,

several posterior medial scutal, 1 postalar callar and scutellar (lateral and

median) bristles reddish-brown and well developed; pleural integument golden-

brown; antepronotum covered with narrow curved golden scales, numerous

golden bristles; postpronotum covered with narrow curved golden scales, 9-10

golden bristles; propleuron with narrow curved golden scales, numerous golden

Reinert: Aedes (Aedimorphus) in Southeast Asia 51

bristles; postspiracular area with a patch of narrow curved golden scales, 7-8

golden bristles; subspiracular area with a patch of narrow curved golden scales;

mesepisternum with an upper patch of moderately broad golden scales; a pos-

terior patch of broad golden scales, several upper and posterior golden bris-

tles, lower ones shorter; prealar knob with a few narrow curved golden scales,

several golden bristles; paratergite with narrow curved golden scales on later-

al and ventral surfaces; mesepimeron with a patch of broad golden scales and

numerous golden bristles on upper area; other areas bare. Legs (Fig. 16).

Coxae I-III each with several golden bristles and a patch of moderately broad

golden-white scales, I with scales on anterior and lateral surfaces, II with

scales on anterior surface, III with a small anteroventral patch; trochanters

I-III each with a patch of broad white scales; femur I with anterior surface

white with an indistinct median longitudinal brown stripe, posterior surface

white with a ventral longitudinal brown stripe from base to apex; femora II, II

with anterior surface brown, each with a median longitudinal white stripe from

base to apex, stripe broader on III and completely covers basal 0.25, II, III with

posterior surface white with a few brown scales forming an indistinct stripe on

apical 0.25; tibiae I-III each white with a dorsal and a ventral longitudinal brown

stripe from base to apex; tarsi I-III brown each with an anteromedian and a

posteromedian longitudinal white stripe on tarsomeres 1-3 and occasionally on

tarsomeres 4 of II, III; posttarsi I-III each with 2 ungues, I, II equal, each bear-

ing a tooth, III equal, simple. Wing. Dorsal veins covered with moderately

broad scales; costa and subcosta covered with golden scales with a few brown

scales along posterior margin of costa (anterior always golden scaled); radius,

cubitus and anal veins each with a few golden scales intermixed with brown

scales on basal 0.65; remainder of veins brown scaled; remigium golden scaled

with brown scales along anterior margin; ventral veins brown scaled with basal

0.75 of costa and basal 0.50 of radius covered with golden scales; alula with

narrow golden-brown scales along fringe; 2 golden remigial bristles. Halter.

Pedicel pale, capitellum white scaled. Abdomen. Terga and sterna covered

with golden scales and with numerous golden bristles, mostly along posterior

margins; tergum i with a rectangular patch of golden-white scales on latero-

tergite. Genitalia (Fig. 25). Tergum VIII index 1.27-1.38; sternum VII

index 0.98-1.04; tergum IX bilobed with 6-12 bristles on each lobe, index

1,13-1.23; insula tongue-like, covered with minute setae and with 3-6

small tuberculi on apical 0.25; lower vaginal lip narrow, moderately

pigmented, covered with minute setae; upper vaginal lip narrow, heavily

pigmented, covered with minute setae; upper vaginal sclerite large, mod-

erate to heavily pigmented, triangular shaped; postgenital lobe moderately

long, apex with a small median indentation, 6-10 bristles on each side of

midline, covered with minute setae, dorsal PGL index 0.90-1.02, ventral

PGL index 2.03-2.22; cerci moderately long, 0.75-0.90 extended from

segment VII, narrow, apex sharply rounded, numerous bristles on dorsal

and lateral surfaces, index 2.38-2.54, cercus/dorsal PGL index 3.32-

3.71; 3 spherical, pigmented seminal capsules, 1 large and 2 Slightly

smaller ones.

MALE (Fig. 8). Similar to female in general habitus. Head. Maxil-

lary palpus golden scaled with a dorsoapical brown spot on segment 2, longer

than proboscis by length of apical segment. Thorax. Postspiracular area

without scales. Legs (Fig. 17). Posttarsi I-III each with 2 ungues, I, II with

52 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

ungues unequal, each bearing a tooth, III unequal, simple. Abdomen. Terga

brown, I with a few dorsomedian golden scales, II-VI each with a narrow

golden basal band and a dorsomedian stripe not reaching posterior margins of

terga, afew lateral golden scales on VI-VII, VIL golden with a pair of admedian

brown patches, VIII golden scaled; sterna golden scaled. Genitalia (Fig. 41).

Tergum IX strongly bilobed with 7-8 bristles on each lobe, entire surface

covered with minute spircules; gonocoxite long and moderately broad, dorsal

surface, except tergomesal margin, covered with short thin bristles, lateral

surface covered with long stout bristles from base to apex, ventral surface

with a few long stout bristles at apex, distal 0.40 with moderately long stout

bristles becoming very numerous on sternomesal margin, remainder of area

with scattered short bristles, scattered scales on lateral and ventral surfaces;

gonostylus with pedicel very short and broad, distal 0.67 expanded into a large

mesal lobe and a lateral narrow, slightly longer, incurved, tapered horn

attached approximately 0.67 from base, mesal expanded lobe with lateral mar-

gin concave, apex broad and flat with 3 short hairs near lateral margin and 2

similar ones near mesal margin, mesal margin of lobe with a small apical con-

cave area bearing a short, strongly curved, pigmented gonostylar claw, a short

fine hair near base of claw and a patch of moderately long hair-like spicules

along mesal margin proximad from base of claw, 3 short fine hairs dorsally

near middle of lobe; basal mesal lobe short and rounded apically, distal 0. 50

with 5-6 short thin bristles, entire surface covered with short hair-like spi-

cules; proctiger short, paraproct with a subapical thumb-like process, cercal

setae absent; phallosome with aedeagus of type I with 2 lateral plates connected

basally, each plate with 4 short blunt lateral teeth on distal 0.58 and covered

with a dorsal flap, paramere long, approximately 0.95 length of lateral plate;

sternum IX large, entire surface covered with minute spircules, 6-7 bristles

near center. ;

PUPA (Fig. 54). Chaetotaxy as recorded in Table 6. Cephalothorax.

Hair 5-C with 3-6 branches; 7-C double or triple; 8-C with 2-6 branches.

Respiratory trumpet. Moderately pigmented; index 4, 25-4, 81, average 4. 52.

Metanotum. Hair 10-C with 10-18 branches; 12-C with 3-7 branches. Abdo-

men. Hair 5-I with 8-15 branches; 10-11-I single, present or absent; 1-II with

9-36 branches; 4-II with 5-9 branches; 1-III with 7-18 branches; 6-VI double

or triple; 1-VII with 5-9 branches; 6-VII with 8-13 branches; 9-VII with 3-8

branches; 11-VII single to triple. Paddle. Ovoid; with very minute serrations

along basal 0.60 of outer margin; midrib does not reach apex; hair 1-P short,

single; index 1.24-1.70, average 1.40.

LARVA (Fig. 65). Chaetotaxy as figured. Head. Hairs 1, 3, 14-C

single; 4-C with 6-7(7) branches; 5, 6-C with 4-6(5) branches; 7-C with 5-12(7)

branches; 8, 10-C double or triple (2); 9, 13-C double or triple (3); 11-C with

3-6(4) branches; 12-C with 3-5(5) branches; 15-C single or double (2); basal

maxillary hair single; mental plate with 24-27(27) teeth. Antenna. Moderately

to heavily pigmented; numerous stout spicules scattered over shaft, those near

and distad of middle longer; hair 1-A with 7-14(9) branches inserted 0. 43-

0.47 from base; 2-A long; 3-A approximately equal in length to 2-A. Thovax.

Hair 0-P with 4-8(5) branches; 1, 5, 6, 10, 12, 14-P single; 2, 8-P double or

triple (2); 3-P double or triple (3); 4, 11-P single to triple (2); 7-P triple; 9-P

single or double (2); 1-3-M with 2-4(3) branches; 5, 7, 10, 12-M single; 6-M

with 5-7(6) branches; 8-M with 5-8(8) branches; 9-M with 4-8(6) branches; 11-M

single or double (1); 13-M with 5-8(6) branches; 14-M with 6-10(7) branches; 1,

11, 12-T single or double (1); 2-T with 3-4(3) branches; 3-T with 5-10(6)

Reinert: Aedes (Aedimorphus) in Southeast Asia 53

branches; 4-T with 2-6(3) branches; 5, 10-T single; 6-T single to triple (2);

7-T with 7-12(9) branches; 8-T with 4-7(5) branches; 9-T with 6-8(6) branches;

13-T with 5-9(5) branches. Abdomen. Hairs 0, 14-VIII single; 2-VIII with

4-5(4) branches; 2-VIII single to triple (2); 3-VII with 8-17(14) branches; 4-

VII double or triple (2); 5-VIII with 7-10(7) branches; 6-V-VI short; comb of

14-20(16) scales arranged in 2 irregular rows, scales with a long stout pointed

median spine and short denticles along lateral margins of base; 1-X with 2-4(2)

branches; 2-X with 9-13(12) branches; 3-X single; ventral brush with 8-9 (usu-

ally 9) hairs on grid and 3 precratal ones; saddle moderately pigmented with

minute ridges, incompletely rings segment, with small spicules along posterior

margin, acus absent; 4 anal papillae, very long and slender, tapering to an apical

point. Siphon. Moderately pigmented with minute ridges over entire surface;

acus very small; index 6.50-7.91; pecten with 10-14(10) teeth, apical 6-7 teeth

smooth, apical 3-4 wider spaced than remainder which have a slender apical

attenuated filament with 1-2 lateral denticles near middle; hair 1-S with 5-6(5)

branches, inserted 0. 72-0. 77 from base.

TYPE DATA. There are 1 male and 1 female syntypes of Aedes palli-

dostriatus in the British Museum (Natural History) and each bears the fol-

lowing data: Culex pallidostriatus Theobald, INDIA, S. Christophers. Lecto-

type designation is hereby made for the syntype male and allolectotype designa-

tion is made for the syntype female. The holotype female of Culex parascelos

Theobald bears the following data: Madras Town, Madras, INDIA, 30 October

1908, and is in the Indian Museum (Barraud 1928: 665, 1934: 261).

DISTRIBUTION. Specimens examined--12 males, 25 females, 5 pupae,

23 larvae and 6 individual larval rearings from the following locations:

CEYLON. Hambantota, Kapitigalla, Kurunegala, Peradeniya, Trincomali.

INDIA. Bengal, Tezgaon, Tinpahar near Rajmahal; Bihar, Pusa; Bom-

bay Deccan, Belgaun; Madras, Madras; Punjab, Karnal; United Provinces,

Anwarganj.

THAILAND. Nanchanaburi, Ban Sai Yok.

WEST PAKISTAN. Lahore, Shah Zada.

Other distribution.

BANGLADESH (East Pakistan). Dinajpur, Thakurgaon, Akcha,

Madarganj (Aslamkhan and Wolfe 1971: 31).

CEYLON (Chow et al. 1954: 117); North-Western Province, Bandara-

koswatte, Magulagama; Western Province, Colombo (Carter 1948; 313, 315).

CHINA (Feng 1958: 62).

INDIA. Bengal, Dum Dum, Cuttack, Delhi, Manhupar, Trombay,

Bombay Harbour (Barraud 1928: 665); Mardas, North Arcot (Reuben 1971: 120).

MALAYSIA. Kedah, Kampong Bukit Kechik; Negri Sembilan, Port

Dickson, Pulau Mertajam (McDonald 1957: 21).

THAILAND. Chaingmai (Thurman and Thurman 1955: 222).

WEST PAKISTAN. Kohat, Kohat-Hangu Valley (Qutubuddin 1960: 358).

TAXONOMIC DISCUSSION. Aedes pallidostriatus is similar in adult

habitus and pupal and larval chaetotaxy to mediolineatus and is discussed under

that species. The male terminalia of these 2 species can be separated from

each other by the shape of the gonostylus.

BIOLOGY. Larvae were collected in India from ground pools. Adults

were taken biting cattle and larvae from rice fields in West Pakistan. In

Ceylon, larvae were collected from paddy fields and adults in a Malaise trap

at an elevation of 10 feet.

In India, Barraud (1934: 262) found the immatures in open pools formed

54 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

by rain and seepage, water filled dikes, ditches and borrow pits. Aslamkhan

and Wolfe (1971: 31) collected adults resting in a house, biting cattle and biting

humans in India. Carter (1948: 314), working in Ceylon, collected adults from

a cattle baited trap.

AEDES (AEDIMORPHUS) PAMPANGENSIS (LUDLOW)

(Figs. 9, 16, 17, 26, 42, 55, 66)

Reedomyia Pampangensis Ludlow 1905, Can. Ent. 37: 94 (9).

Reedomyia niveoscutella Theobald 1905, J. econ. Biol. 1: 22 (9*); Theobald

1907, Monogr. Cul. 4: 259 (o'*, 9); Theobald 1910, Monogr. Cul. 5:

253.

Reedomyia niveoscutellata Theob., Brunetti 1907, Rec. Indian Mus. 1: 362;

Brunetti 1912, Rec. Indian Mus. 4: 487.

Reedomyia pampangensis Ludlow, Brunetti 1907, Rec. Indian Mus. 1: 362;

Theobald 1907, Monogr. Cul. 4: 258 (2); Theobald 1910, Monogr. Cul.

5: 253; Brunetti 1912, Rec. Indian Mus. 4: 487; Stone and Knight 1956,

J. Wash. Acad. Sci. 46: 223.

Ochlerotatus niveoscutellatus Theob., Brunetti 1920, Rec. Indian Mus. 17:

139; Senior-White 1923, Cat. Indian Insects, Cul., p. 80.

Ochlerotatus pampangensis Ludl., Brunetti 1920, Rec. Indian Mus. 17: 140.

Aedes (Ecculex) alboscutellatus Theobald, Edwards 1922b, (in part), Indian J.

med. Res. 10: 467; Edwards 1932 (in part), GeneralInsec., Fasc.

194: 167.

Aedes (Ecculex) niveoscutella Theobald, Edwards 1922b, Indian J. med. Res.

10: 467.

Aédes (Aédimorphus) niveoscutella Theobald, Dyar and Shannon 1925, Insecutor

Inscit. menstr. 13: 76; Edwards 1932, GeneraInsec., Fasc. 194: 168;

Barraud 1934, Fauna Brit. India, Diptera 5: 251 (oc, 9).

Aedes (Aedimorphus) niveoscutella (Theo.), Barraud 1928, Indian J. med. Res.

15: 659(9).

Aédes (Aédimorphus) alboscutellatus Theobald, Edwards 1932 (in pari), Genera

Insec., Fasc. 194: 167.

Aedes (Aedimorphus) pampangensis (Ludlow), Bohart 1945, U. S. Navmed.

580, p. 53 (co, 9); Knight and Hull 1953, Pacif. Insects 7: 454 (o*, 9,

L*); Stone et al. 1959, Thomas Say Found. 6: 196.

Aedes (Aedimorphus) niveoscutellum (Theobald), Stone et al. 1959, Thomas

Say Found. 6: 195. NEW SYNONYM.

FEMALE (Fig. 9). Head. Antenna brown, approximately 1.10 length

of proboscis, pedicel pale with a patch of short fine brown hairs mesally (a

few small brown scales also present on specimens from India), flagellomere

1 pale with a few small brown scales; clypeus light brown, bare; maxillary

palpus golden-brown scaled, approximately 0.19 length of proboscis; pro-

boscis golden-brown scaled with apical 0.25 darker, ventral surface slightly

paler (entirely golden scaled in specimens from India), approximately equal

in length to femur I; vertex with dorsum covered with narrow decumbent

scales, anterior 0.25 brownish and the remainder white; lateral surface

covered with broad white scales and with an anterodorsal dark patch; numer-

ous golden-brown erect forked scales on occiput and vertex extending anteri-

orly to the ocular line. Thorax. Scutal integument reddish-brown; scutum

covered with narrow curved pale reddish-brown scales, narrow curved white

Reinert: Aedes (Aedimorphus) in Southeast Asia 55

scales forming small patches on median anterior promontory area, anterior

scutal fossal area and supra-alar area from scutal angle to wing base; scutel-

lum completely covered with overlapping broad silvery-white scales; median

anterior promontory, acrostichal, dorsocentral (anterior and posterior), scu-

tal fossal (anterior, 2-4 lateral, and 1 posterior), supra-alar, several pos-

terior medial scutal, 1 postalar callar and scutellar (lateral and median) bris-

tles reddish-black and well developed, others absent; pleural integument pale

brown; antepronotum covered with narrow curved white scales, several long

golden to dark bristles; postpronotum covered with narrow curved scales, an-

terior and dorsal ones pale reddish-black, remainder white, 6-7 dark posteri-

or bristles; propleuron with a patch of broad white scales, several long golden

bristles; postspiracular area with a patch of broad white scales, 6-7 golden

bristles; subspiracular area with a small patch of narrow curved white scales;

mesepisternum with an upper and a posterior patch of broad white scales, sev-

eral upper and posterior long golden bristles, lower 8-10 bristles shorter;

prealar knob with numerous golden bristles; paratergite with narrow curved

white scales along lateral and ventral margins; mesepimeron with a patch of

broad white scales and numerous pale golden bristles on upper area; other

pleural areas bare. Legs (Fig. 16). CoxaeI-[II each with several pale golden

bristles, I with anterior surface covered with broad white scales and a few

light brown ones laterally, II with anterior surface covered with broad white

scales, III with a small anterodorsal and an anteroventral patch of broad white

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

each with a dorsoapical white spot, I, II with anterior surface brown, III with

anterior surface pale and dorsoapical 0.30 brown, I with dorsal 0.75 of poster-

ior surface pale and remainder brown, II, III each with basal 0.60 of posterior

surface pale and remainder light brown dorsally; tibiae I, II pale, I with an an-

terior longitudinal brown stripe, II with a ventral longitudinal brown stripe, III

brown with an anterior and a posterior pale longitudinal stripe; tarsi I, II brown

each with a posteroventral longitudinal pale stripe on tarsomere 1 and basal

0.50 of tarsomere 2, II also with a small elongated dorsobasal pale spot; tar-

sus III brown occasionally with a posteroventral longitudinal pale stripe on tarso-

mere 1; posttarsi I-III each with 2 ungues, I, II equal, each bearing a tooth, II

equal, 1 bearing atooth. Wing. Dorsal veins covered with brown moderately

broad scales; costa with a few pale scales at base and along its posterior mar-

gin at the humeral cross vein; ventral veins brown scaled with pale scales on

basal 0.25 of subcosta and a few on posterior of costa; alula with narrow brown

scales along fringe; 2-4 remigial bristles. Halter. Pedicel pale brown, capitel-

lum white scaled. Abdomen. Terga brown dorsally; tergum I with a few pale

scales dorsally (some specimens with tergum I all brown dorsally) and a rec-

tangular patch of white scales on laterotergite; terga III, IV each with a small

basomedian pale scale patch; tergum V with a basomedian pale stripe extending

0.50 posteriorly and a few pale scales along posterior margin; tergum VI with

a median triangular pale scale patch extending from base to posterolateral mar-

gins; tergum VII with basomesal pale scales; terga II-VI each with a broad longi-

tudinal pale band on lateral surfaces; (some specimens from India and the Philip-

pines also with the following: VI with more pale scales posteriorly and VII

nearly completely pale scaled; sterna pale scaled); terga and sterna with numer-

ous golden-brown bristles, mostly along posterior margins. Genitalia (Fig. 26).

Tergum VIII index 1. 22-1.44; sternum VII index 1.01-1. 10; tergum IX bilobed

with 5-8 bristles on each lobe, index 1.17-1.58; insula tongue-like, covered

with minute setae and with 4-6 small tuberculi on apical 0.25; lower vaginal lip

narrow, moderately pigmented, covered with minute setae; upper vaginal lip

56 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

narrow, moderately to heavily pigmented, covered with minute setae; upper

vaginal sclerite large, moderately pigmented, triangular shaped; postgenital

lobe moderately long, apex with a moderately deep median indentation, 6-10

bristles on each side of midline, covered with minute setae, dorsal PGL index

0.98-1.48, ventral PGL index 2. 45-3. 29; cercus long, 0. 70-0. 80 extended

from segment VI, narrow, apex sharply rounded, numerous bristles on dor-

sal and lateral surfaces, index 3.48-3.63, cercus/dorsal PGL index 3. 74-4. 79;

3 spherical, pigmented seminal capsules, 1 large and 2 slightly smaller ones.

MALE (Fig. 9). Similar to female in general habitus. Head. Maxil-

lary palpus brown with a broad lateral patch of pale scales near base of seg-

ment 2, longer than proboscis by 0.50 length of apical segment; vertex without

anterodorsal dark scale patch. Thorax. Postspiracular area without scales.

Legs (Fig. 17). Tarsi I, II each with posteroventral longitudinal pale stripe

usually reduced to basal 0.75 of tarsomere 1; posttarsi I-III each with 2 ungues,

I, II with ungues unequal, each bearing a tooth, III equal, simple. Abdomen.

Terga I-VII brown, each with a narrow indistinct longitudinal pale stripe on

lateral surfaces; tergum VIII with a large laterobasal pale scale patch. Gen-

italia (Fig. 42). Tergum IX bilobed with 5-8 bristles on each lobe, entire

surface covered with minute spicules; gonocoxite moderately long and broad,

dorsal surface with a median patch of long stout bristles from near base to

apex, somewhat shorter and thinner bristles laterad and extending over lateral

margin from base to apex; tergomesal margin with a dense patch of short,

somewhat flattened, curved bristles near middle, a number of short thin bris-

tles distad and a few proximad of patch, ventral surface covered with scattered

short bristles and long stout ones on distal 0.30, long bristles more numerous

on sternomesal margin, scattered scales on lateral and ventral surfaces;

gonostylus with pedicel very short and broad, distal 0.80 greatly expanded with

a laterotergal flap-like structure bearing 2 dense lateral patches of very long

thin hairs near middle and numerous short fine hairs extending to apex which

terminates in a small incurved pigmented point, remainder of expanded por-

tion concave dorsally with a short, flat, apically tapering gonostylar claw

attached mesally approximately 0.80 from base, 4 smaller accessory claws

along apical margin, numerous short fine hairs forming a large patch on ter-

galand sternal surfaces; basal mesal lobe short and rounded apically, distal

0.55 with 2-3 short stout apical bristles and 2-3 short fine ones, entire sur-

face covered with short hair-like spicules; proctiger short, paraproct with a

subapical thumb-like process, cercal setae absent; phallosome with aedeagus

of type I with 2 lateral plates connected basally, each plate with 6-7 short

blunt lateral teeth on distal 0.57 and covered with a dorsal flap, paramere long,

approximately 0.98 length of lateral plate; sternum IX large, entire surface

covered with minute spicules, 5-7 bristles near center.

PUPA (Fig. 55). Chaetotaxy as figured and recorded in Table 7.

Cephalothorax. Hair 5-C with 6-9 branches; 7, 8-C with 8-11 branches.

Respiratory trumpet. Moderately pigmented; index 4.53-5.20, average 4. 89.

Metanotum. Hair 10-C with 13-16 branches; 12-C with 3-11 branches. Abddo-

men. Hair 5-I with 10-18 branches; 1-II with 24-40 branches; 4-I with 6-11

branches; 1-III with 10-16 branches; 6-VI with 6-8 branches; 1-VII with 7-10

branches; 6-VII with 7-14 branches; 9-VII with 5-7 branches; 11-VII with 2-4

branches. Paddle. Ovoid; with very minute serrations along basal 0.50 of

outer margin; midrib does not reach apex; hair 1-P short, single or double;

index 1.18-1.25, average 1. 22.

LARVA (Fig. 66). Chaetotaxy as figured. Head. Hairs 1, 3, 14-C

single; 4-C with 5-7(6) branches; 5-C with 5-8(6) branches; 6-C with 4-7(5)

Reinert: Aedes (Aedimorphus) in Southeast Asia at

branches; 7-C with 8-13(10) branches; 8-C with 2-4(3) branches; 9-C with 2-5(3)

branches; 10-C single or double (2); 11-C with 3-6(4) branches; 12-C with 3-6(5)

branches; 13, 15-C with 2-4(2) branches; basal maxillary hair single; mental

plate with 32-34(33) teeth. Antenna. Moderately pigmented; numerous stout

spicules scattered over shaft, more numerous near middle; hair 1-A with 6-11

branches, inserted 0.35-0.42 from base; 2-A long; 3-A approximately 0.50

length of 2-A. Thorax. Hair 0-P with 5-12(6) branches; 1, 5, 6, 10, 12-P

single; 2-P double or triple (2); 3, 7-P double or triple (3); 4, 9, 11-P with

2-4(3) branches; 8, 14-P double; 1-M with 3-4(4) branches; 2-M with 2-4(3)

branches; 3-M with 2-5 branches; 4-M with 4-6(4) branches; 5-M double or

triple (3); 6-M with 3-6(4) branches; 7, 10, 12-M single; 8-M with 7-10(8)

branches; 9-M with 7-10(7) branches; 11-M single or double (1); 13-M with

5-12(6) branches; 14-M with 7-11(7) branches; 1, 5, 10, 11-T single; 2-T with

4-7(5) branches; 3-T with 17-28(17) branches; 4-T with 3-6(3) branches; 6-T

double or triple (3); 7-T with 6-11(7) branches; 8-T with 5-7(5) branches; 9-T

with 5-8(8) branches; 12-T single to triple (2); 13-T with 9-14(9) branches.

Abdomen. Hairs 0, 14-VIII single; 1-VII with 4-9(7) branches; 2-VIII with

2-5(3) branches; 3-VIII with 8-13(12) branches; 4-VIII with 2-4(3) branches;

5-VII with 5-8(8) branches; 6-IV-VI short; comb with 18-23(20) scales arranged

in 3 irregular rows, scales with a long stout pointed median spine and short

denticles along lateral margins of base; 1-X double or triple (2); 2-X with 10-

15(14) branches; 3-X single; ventral brush with 9-10 (usually 9) hairs on grid

and 3 precratal ones; saddle lightly pigmented with minute ridges, incompletely

rings segment, with a few spicules along posterior margin, acus present; 4

anal papillae, long, each broad at base and tapering to a pointed apex. Siphon.

Lightly pigmented with minute ridges; acus present; index 7. 50-9.00; pecten

with 15-18(15) teeth, apical 3-4 teeth larger, smooth and wider spaced than re-

mainder which have a long slender apical attenuated filament with 1-2 basal

denticles; hair 1-S with 4-5(4) branches, inserted 0. 69-0. 74 from base.

TYPE DATA. Reedomyia Pampangensis Ludlow, 3 syntype females,

Angeles, Pampanga, Luzon, PHILIPPINES, September, Dr. Eugene R. Whit-

more, caught in the woods and in the quarters, types non-existent (Stone 1970:

151). There is one specimen in the United States National Museum (Natural

History) which bears labels with the following data: Reedomyia pampangensis

Ludlow, Camp Wm. McKinley, Rizal, P. I., Oct. 25, Nov. 3, 05, Type; Type

No. 27795 U.S.N.M. The first label is in Ludlow's handwriting, but since the

specimen was collected after date of publication (March 1905) of the original

description, it cannot be considered as the type. No other type material of

pampangensis was found. Knight and Hull (1953: 454) considered the above

specimen as the type but later Stone and Knight (1956: 223) indicated it could

not be the type since it was collected after publication of the name. Since the

type material of pampangensis has been lost and this species has a superficial

resemblance to several other species of Aedimorphus as well as occurring in a

wide range in the Oriental Zoogeographical Region I am hereby designating an

adult female as neotype for the species. The preceeding written description and

following habitus illustrations of the female are of the neotype (a comparison to

specimens from other areas in its geographical range is also made). In the tax-

onomic discussion of the species some diagnostic characters for distinguishing

pampangensis from closely related species is given. The neotype is deposited

in the United States National Museum (Natural History), Washington, D. C., and

bears labels with the following information: Sison, Pangasinan, Luzon, P. IDS.,

April 1945, J. G. Franclemont; NEOTYPE, Aedes (Aedim.) pampangensis, Det:

J. F. Reinert. Reedomyia niveoscutella Theobald, holotype female, INDIA,

58 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Capt. James, in British Museum (Natural History).

I have compared the neotype of pampangensis to the holotype of niveo-

scutellum and cannot find any apparent differences; therefore I am placing niv-

—eoscutellum in synonymy to pampangensis.

DISTRIBUTION. Specimens examined--29 males, 107 females, 11

pupae and 29 larvae from the following locations:

INDIA. Bzhar, Pusa, Sharma.

INDONESIA. Sumatra, Atjeh, Kroeng Raja.

PHILIPPINES. Luzon, Calaccad, Nunoz Necija, Olongapo; Camarines

Sur; Leyte, Mahaplag; Mindanao, Pasananco, Pettit Barracks, Zamboanga;

Mindoro, San Jose; Pampanga, Camp Stotsenberg, Clark Air Force Base;

Pangasinan, Camp Gregg, Sison, Tayng; Rizal, Camp Nichols, N. Ecija;

Zambales, Subic Bay.

SOUTH VIETNAM. Binh Dinh, An Khe.

THAILAND. Surasthani.

Other distribution.

INDIA. Purneah, Kierpur (Barraud 1928: 660).

INDONESIA. Sumatra, Atchin (Brug and Edwards 1931: 258); Lokop,

Bireuen (Brug and Haga 1923: 639; Haga 1924: 830); Java(Barraud 1934: 252).

PHILIPPINES. Manila, Quezon City; Zamboanga, Pettit Barracks,

San Ramon City, Zamboanga (Knight and Hull 1953: 457); Pampanga, Angeles

(Ludlow 1905: 95): Pampanga, Dau; Zambales, Olongapo (Basio 1971: 11);

Clark AB (Dowell, Libay and Baisas 1965: 14).

SOUTH VIETNAM. Binh Thuy, Phan Rang, Pleiku (Parrish 1968a:

3, 4, Parrish 1969: 554); Cam Ranh Bay (Reisen et al. 1971: Table 12).

THAILAND. Nakhom Phanon, U-Tapao (Reisen et al. 1971: Tables 4

and 8).

TAXONOMIC DISCUSSION. Aedes pampangensis has a somewhat

superficial resemblance to mediolineatus and pallidostriatus in the pleural

scale markings but is distinguished from these species by the presence of

broad silvery -white scales on the scutellum and the absence of longitudinal

white stripes on the scutum. The broad silvery-white scales on the scutellum

is also reminiscent of alboscutellatus and its relatives, but pampangensis is

easily separated from these species by the absence of dorsal spots on the

scutum and the abdominal terga which lack dorsobasal pale bands and are

completely white scaled laterally. The gonostylus of the male genitalia of

pampangensis is greatly expanded and very distinctive when compared to other

members of the subgenus.

The description and illustration by Borel (1930: 267) attributed to the

male of pampangensis (= niveoscutellum) was not of this species but was

probably alboscutellatus.

The pupa of pampangensis is distinctive in having abdominal hair 1-I-

VIII well developed, hair 1-C with 5-8 branches, 4-C with 6-8 branches, 7-C

with 8-11 branches, 5-II with 10-14 branches, 6-IV, V with 5-8 branches and

6-VI with 6-8 branches.

The larva of pampangensis resembles those of mediolineatus and pal-

lidostriatus in having a very long siphon but can be separated from these 2

species by mesothoracic hair 5-M which is double or triple and metathoracic

hair 3-T which has 17-28 branches while mediolineatus and pallidostriatus

have hair 5-M single and 3-T with 5-11 branches.

BIOLOGY. Immatures appear to prefer unshaded, grassy flood pool

habitats containing fresh water. Adults have been taken feeding on cattle. In

Thailand immatures have been collected from small flood pools and large

Reinert: Aedes (Aedimorphus) in Southeast Asia 09

wheel tracks, with turbid fresh water occasionally containing scarce emergent

aquatic vegetation, no shade, located in teak plantations in the plains and in a

primary bamboo grove in the mountains, and at altitudes from 400 to 635

feet. Larvae were collected in association with the following mosquitoes:

Aedes andamensis, culicinus, gubernatoris, imprimens, pipersalatus, vittatus

and Anopheles Rochi, Inthe Philippines immatures were collected from a

grassy pond, road canal, rice field, small flooded area near road, grassy

eround pool, and along the shaded bank of a clear water stream containing

algae and vegetation. In West Pakistan, adults were taken biting cattle.

Larvae were collected from scattered rain pools in a grassy area and

from a grassy pool in the bed of a temporary stream in the Philippines by

Knight and Hull (1953: 457).

AEDES (AEDIMORPHUS) PIPERSALATUS (GILES)

(Figs. 10, 16, 17, 27, 43, 56, 67)

Stegomyia pipersalata Giles in Theobald 1901, Monogr. Cul. 2: 316 (oc, 9);

Giles 1902, Handb., 2nd Ed., p. 372 (2*); Blanchard 1905, Moust.,

p. 264 (2); Brunetti 1907, Rec. Indian Mus. 1: 332; Theobald 1910,

Monogr. Cul. 5: 607; Brunetti 1912, Rec. Indian Mus. 4: 448.

Pseudograbhamia maculata Theobald 1905, J. Bombay nat. Hist. Soc. 16:

243 (oc, 9); Brunetti 1907, Rec. Indian Mus. 1: 140; Theobald 1907,

Monogr. Cul. 4: 314 (o‘, ?*); Brunetti 1912, Rec. Indian Mus. 4: 460.

Ochlerotatus pipersalatus (Giles), Edwards 1913, Bull. ent. Res. 4: 227

(o, 2); Brunetti 1920, Rec. Indian Mus. 17: 138; Senior-White 1923,

Cat. Indian Mus, Cul., p. 81.

Aedes (Ecculex) pipersalatus Giles, Edwards 1922b, Indian J. med. Res.

10: 467.

Aedes (Aedimorphus) pipersalatus (Giles), Barraud 1928, Indian J. med.

Res. 15: 664; Stone et al. 1959, Thomas Say Found. 6: 196; Qutubuddin

1960, Mosquito News 20: 358 (c*).

Aédes (Aédimorphus) pipersalatus Giles, Edwards 1932, Genera Insec., Fasc.

194: 170.

Aédes (Aédimorphus) pipersalatus (Giles), Barraud 1934, Fauna Brit. India,

Diptera 5: 258 (o*, 9, L*).

FEMALE (Fig. 10). Head. Antenna dark brown, approximately equal

to length of proboscis, pedicel brown with a few small dusky scales and a

patch of short fine brown hairs mesally, flagellomere 1 pale basally with a few

small dusky scales; clypeus dark, bare; maxillary palpus brown scaled with

apical 0.25 white, apical hairs also white, approximately 0.18 length of pro-

boscis; proboscis brown scaled dorsally with a few intermixed white scales,

white ventrally from near base to distal 0.25, approximately 1.26 length of

femur I; vertex with dorsum covered with narrow curved decumbent scales

arranged in an anteromedian diamond-shaped brown group and the remainder

white; broad white scales extending halfway down lateral surface with broad

brown ones below and also forming an anterodorsal patch; numerous dark

brown erect forked scales on occiput and vertex extending anteriorly to ocular

line. Thorax. Scutal integument dark brown; scutum covered with narrow

curved reddish-black scales; narrow curved white scales forming small cir-

cular patches on median anterior promontory area, scutal fossal areas (one

each on anterior, lateral and posterior areas), scutal angle, supra-alar area

60 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

(a patch anterior to and a small indistinct spot posterior to wing base and a

patch medially near dorsocentral setal line), posterior medial scutal area

and extending posteriorly along lateral margins of prescutellar space, similar

scales scattered over area mesally to dorsocentral setae; scutellum with a

patch of broad and a few narrow curved white scales on each lobe; median an-

terior promontory, acrostichal, dorsocentral (anterior and posterior), scutal

fossal (anterior, 2-4 lateral, 1-2 median and 2-3 posterior), supra-alar,

several posterior medial scutal, 1 postalar callar and scutellar (lateral and

median) bristles reddish-black and well developed; pleural integument dark

brown; antepronotum with narrow curved white scales, several long dark bris-

tles; postpronotum covered with narrow curved scales, reddish-black ones

dorsally and white ones medially, 5-6 posterior dark bristles; propleuron

with a patch of broad white scales, several golden bristles; postspiracular

area with a patch of broad white scales, 4-5 golden bristles; subspiracular

area with 2 patches of broad white scales, lower one larger; mesepisternum

with median, upper and posterior patches of broad white scales, median patch

small, several upper and posterior golden bristles, lower ones shorter; pre-

alar knob with several golden bristles (2 specimens from Ceylon had 2-3 broad

white scales on ventral margin); paratergite with broad white scales on latero-

ventral margin; mesepimeron with a large patch of broad white scales and

several golden bristles on upper area; other pleural areas bare. Legs (Fig.

16). Coxae I-III each with several brown or golden bristles, I with broad

brown scales and a few white ones intermixed on anterior and lateral surfaces,

a small patch of broad white ones dorsally, II with anterior surface covered

with broad brown scales and a small patch of broad white ones dorsally, III

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

of ventral broad dusty-white scales, a few white apical ones on III; femora I-III

brown, each with anterior and posterior surface with intermixed white scales

and a dorsoapical white spot, posterior surface of I, III with numerous white

scales nearly covering basal 0.50 of II and basal 0.75 of III; tibiae I-III brown,

each with intermixed white scales and a small dorsoapical white spot, I with

an indistinct posteroventral longitudinal white stripe, II with an indistinct

posteromedian longitudinal white stripe; tarsi I-III brown, I with tarsomeres

1-2 each with a basal white band, tarsomere 3 with a dorsobasal white spot,

Il with tarsomeres 1-3 each with a basal white band, tarsomeres 4 with a

dorsobasal white spot, III with tarsomeres 1-4 each with a basal white band,

tarsomere 5 with a dorsobasal white spot, afew scattered white scales on

tarsomere 1; posttarsi I-III each with 2 ungues, I, II equal, each bearing a

tooth, III equal, simple. Wing. Dorsal veins covered with moderately broad

brown scales with white ones intermixed, approximately 30 percent of the

scales white; tertiary fringe scales with white and brown scales intermixed

along posterior margin of wing; costa with a small white spot at base; ventral

veins brown scaled with white scales intermixed; alula with narrow brown

scales along fringe; 1-2 remigial bristles. Halter. Pedicel pale, capitellum

white scaled. Abdomen. Terga brown with broad dorsobasal white bands on

II-VI and a few basal pale scales on VII; tergum I with several white scales

mesally and a rectangular patch of white scales on laterotergite; terga II-VI

each with a lateromedian patch of white scales, patches not connected to dor-

sal bands; sterna with lateral areas mainly pale scaled, median and posterior

areas brown; terga and sterna with numerous golden bristles, mostly along

posterior margins. Genitalia (Fig. 27). Tergum VIII index 0. 73-0. 79; sternum VIII

index 1.05-1. 11; tergum IX bilobed with 4-7 bristles on each lobe, index 1. 22-

1.41; insula tongue-like, covered with minute setae and with 4-5 small tuberculi

Reinert: Aedes (Aedimorphus) in Southeast Asia 61

on apical 0.25; lower vaginal lip, narrow, moderately pigmented, covered with

minute setae; upper vaginal lip narrow, heavily pigmented, covered with min-

ute setae; upper vaginal sclerite large, moderate to heavily pigmented, tri-

angular shaped; postgenital lobe moderately long, apex with a moderately deep

median indentation, 5-8 bristles on each side of midline, covered with minute

setae, dorsal PGL index 0.96-1.09, ventral PGL index 2. 08-2.36; cercus long,

0.90-1.00 extended from segment VII, narrow, apex sharply rounded, numer-

ous bristles on dorsal and lateral surfaces, index 3.33-3.41, cercus/dorsal

PGL index 3. 46-4. 50; 3 spherical, pigmented seminal capsules, 1 large and

2 slightly smaller ones.

MALE (Fig. 10). Similar to female in general habitus. Head. Maxil-

lary palpus brown with dorsobasal white spots on apical 2 segments, afew

scattered white scales on antepenultimate segment, longer than proboscis by

length of apical segment; proboscis brown with a few scattered ventral white

scales. Legs (Fig. 17). Tarsus I with tarsomere 1 also with a few scattered

white scales, tarsomere 4 with a basal white band; tarsus III with tarsomere 5

with a narrow basal white band; posttarsi I-III each with 2 ungues, I, II with

ungues unequal, each bearing a tooth, III unequal, simple. Wing. Basal white

spot on costa reduced. Abdomen. Terga brown with broad basal white bands

on II-VII and a few lateromedian white scales on V-VI; tergum VIII white with

a few lateral brown scales. Genitalia (Fig. 43). Tergum IX strongly bilobed

with 5-7 bristles on each lobe, entire surface covered with minute spicules;

gonocoxite long and moderately broad, dorsal surface with a few scattered

short fine bristles, mostly along tergomesal margin, lateral surface with

numerous long stout bristles from base to apex, ventral surface with a number

of moderately long to long stout bristles on distal 0.35 and scattered short

ones over the remainder, scattered scales on lateral and ventral surface;

gonostylus with pedicel very short and broad, distal 0.63 expanded into a large

mesal lobe and a lateral longer, narrow, slightly incurved, tapered horn

attached approximately 0.43 from base with a very short fine apical hair,

mesal expanded lobe with a long, flattened, pigmented gonostylar claw attached

mesally near middle, a short fine hair near base of claw and short hair-like

spicules extending from claw proximad, 2-3 short fine hairs near apex and 3-4

similar ones scattered over tergal surface of lobe; basal mesal lobe short

and rounded apically, distal 0.25 with 4-5 short bristles, entire surface

covered with short hair-like spicules; proctiger short, paraproct with a small

subapical thumb-like process, cercal setae absent; phallosome with aedeagus

of type I with 2 lateral plates connected basally, each plate with 4-7 short blunt

lateral teeth on distal 0.60 and covered with a dorsal flap, paramere long, ap-

proximately 0.98 length of lateral plate; sternum IX large, entire surface

covered with minute spicules, 2-3 bristles near center.

PUPA (Fig. 56). Chaetotaxy as recorded in Table 8. Cephalothorax.

Hair 5-C with 3-5 branches; 7-C with 3-6 branches; 8-C with 2-7 branches.

Respiratory trumpet. Moderately pigmented; index 3.06-4.38, average 3. 72.

Metanotum. Hair 10-C with 5-9 branches; 12-C with 4-7 branches. Abdomen.

Hair 5-I with 7-16 branches; 1-II with 13-20 branches; 4-II with 7-16 branches;

14-II single; 1-III with 5-10 branches; 6-VI double or triple; 1-VII with 3-6

branches; 6-VII with 6-14 branches; 9-VII with 3-6 branches, 11-VII single or

double. Paddle. Ovoid; with very minute serrations along basal 0.50 of outer

margin; midrib does not reach apex; hair 1-P short, single; index 1. 18-1. 53,

average 1.41.

LARVA (Fig. 67). Chaetotaxy as figured. Head. Hairs 1, 3, 14-C

single; 4, 6-C with 4-6(5) branches; 5, 11-C with 5-7(5) branches; 7-C with

62 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

6-10(6) branches; 8, 13-C with 2-4(3) branches; 9-C with 3-4(3) branches; 10-C

double or triple (2); 12-C with 4-6(4) branches; 15-C single to triple (1); basal

maxillary hair single; mental plate with 26-36(28) teeth. Antenna. Lightly pig-

mented; numerous spicules scattered over entire shaft; hair 1-A with 5-9(6)

branches, inserted 0.38-0.45 from base; 2-A long; 3-A approximately 0.50

length of 2-A. Thovax. Hair 0-P with 5-7(6) branches; 1, 5, 6, 10-P single;

2, 9-P double or triple (2); 3, 4, 7-P double or triple (3); 8-P single or double

(2); 11-P double; 12, 14-P single or double (1); 1-M with 3-5(4) branches; 2-M

single or double (1): 3-M with 2-4(3) branches; 4-M with 3-6(4) branches; 5, 7,

10, 11, 12-M single; 6-M with 4-6(5) branches; 8-M with 6-8(7) branches; 9-M

with 6-10(6) branches; 13-M with 5-10(6) branches; 14-M with 5-8(6) branches:

1, 11, 12-T single or "double (1); 2, 4-T with 3-7(4) branches; 3-T with 10-23

(16) branches; 5, 10-T single; 6-T double or triple (2); 7-T with 6-10(6)

branches; 8-T with 4-7(6) branches; 9-T with 4-7(6) branches; 13-T with

5-12(5) branches. Abdomen. Hairs 0, 14-VIII single; 1-VII with 4-7(5)

branches; 2-VIII single to triple (2); 3-VIII with 6- 12(10) branches; 4-VIII double

or triple (2); 5-VIII with 4-8(5) branches; 6-IV-VI short; comb with 18-30(18)

scales arranged in 3 irregular rows, scales with short stout denticles along

lateral and apical margins; 1-X single or double (1); 2-X with 8-14(9) branches;

3-X single; ventral brush varies from 9 hairs on grid and 3 precratal ones to

11 hairs on grid and 1 precratal hair, usually with 10 hairs on grid and 2 pre-

cratal ones; saddle lightly pigmented with minute ridges, incompletely rings

segment, with a few spicules along posterior margin, acus small; 4 anal papillae,

long, each with a broad base and tapering to a pointed apex. Siphon. Moder-

ately pigmented with minute ridges; acus present; index 5.00-5. 56; pecten with

15-19(17) teeth, apical 2-3 teeth longer and wider spaced than remainder which

have a slender apical attenuated filament with 1-3 basal denticles; hair 1-S with

3-4(3) branches, inserted 0. 61-0. 68 from base.

TYPE DATA. Stegomyia pipersalaia Giles, holotype female, Jhansi,

Gonda, N.W. Provinces, INDIA, August 1900, Lt. Col. G. M. Giles, in

British Museum (Natural History); Pseudograbhamia maculata Theobald, syn-

types female and male, Galgamuwa, CEYLON, August 1902, Green, in British

Museum (Natural History).

DISTRIBUTION. Specimens examined--13 males, 32 females, 2 pupae,

14 larvae and 2 individual larval rearings from the following locations:

CAMBODIA. Kirirom.

CEYLON. Galgamuwa, Polgahawela; Hunuwilagama, Wilpattu.

INDIA. Belgaum, Bombay Deccan, Tavargatti; Northwest Gonda.

THAILAND. Kanchanaburi Ban Sai Yok.

WEST PAKISTAN. Lahore.

Other distribution.

CEYLON (Chow et al. 1954: 117); Colombo (James 1914: 262); Andanku-

lam near Trincomalee (Carter and Wijesundara 1948: 141); North-Western

Province, Bandarakoswatte, Magulagama (Carter 1948: 313).

INDIA. Mardas, North Arcot (Reuben 1971: 120); North West Provinces,

Gonda, Jhansi (Giles 1902: 372); Madras Town (Theobald 1910b: 292); Pusa

(Senior-White 1923: 81); Bengal; Central Provinces; Anwarganj, Cownpore

District; Cuttack, Kamptee, Nagpur District, Madhupur, Pusa, Ranihat

(Barraud 1928: 664); Poona, Baramati (Rao and Rajagopalan 1957: 10); Bombay,

Kamptee, Belgaum, Tawaregatti: Bihar, Ranihat; Orissa, Cuttack; Uttar Pra-

desh, Anwarganj, Kanpur, Dehra Dun, Kalsi (Wattal et al. 1958: 223),

WEST PAKISTAN. Baradark Garden, Bolarum (Qutubuddin 1951: 30);

Kohat, Kohat-Hangu Valley (Qutubuddin 1960: 358).

Reinert: Aedes (Aedimorphus) in Southeast Asia 63

TAXONOMIC DISCUSSION. Aedes pipersalatus is similar to taenior-

hynchoides, vexans vexans and vexans nipponii in the adult habitus. Pipersala-

tus possesses the following features: scutum with definite white scale patches;

postprocoxal membrane bare; scutellum with a patch of broad and a few nar-

row curved white scales on midlobe; and wing with approximately 30 percent of

the scales white and intermixed with brown ones while taeniorhynchoides has the

following: scutum lighter scaled with indistinct dorsal patches of white scales;

postprocoxal membrane with a few small broad white scales; scutellum with

only narrow curved white scales on midlobe; and wing with approximately 40

percent of the scales white and intermixed with brown ones. From vexans

vexans and vexans nipponii these 2 species are easily separated by the large

number of white scales intermixed with brown ones on most of the dorsal veins

of the wing especially on the radius and among the tertiary fringe scales along

the posterior margin of the wing. The scales on the distal 0. 85 of the radius

2+3 and along the posterior margin of the wing are all brown in vexans vexans

and vexans nipponii. Other features of pipersalatus are: scutal fossal area of

scutum with 4-5 bristles; postpronotum with 5-6 posterior bristles; and post-

spiracular area with 4-5 bristles while vexans vexans and vexans nipponii

possess the following: scutal fossal area of scutum with 9-11 bristles; post-

pronotum with 8-10 bristles; and postspiracular area with 8-9 bristles.

The postgenital lobe of the female genitalia has a deep median apical

indentation in pipersalatus but only a moderately deep one in taeniorhynchoides.

The male genitalia of pipersalatus are very similar to taeniorhynchoides

but differ in the shape of the gonostylus, number of bristles on basal mesal lobe

(4-5 in pipersalatus and 6-11 in taeniorhynchoides) and number of bristles on

sternum IX(2-3 in pipersalatus and 4-5 in taeniorhynchoides).

The pupa of pipersalatus resembles culicinus and can be separated from

this species by abdominal hair 4-II which has 7-16 branches as compared to

2-5 branches in culicinus.

The chaetotaxy of the larva is similar to alboscutellatus and is dis-

cussed under that species.

BIOLOGY. In Thailand immatures were collected from a small un-

shaded flood pool with turbid fresh water, with scarce emergent aquatic

vegetation, located in a teak plantation in the plains, and at an altitude of 400

feet. Larvae were collected in association with the following species of mos-

quitoes: Aedes culicinus, imprimens, pallidostriatus and Anopheles kochi.

Larvae were collected in jungle pools inIndia. Pupae were found in

paddy fields and in a coconut treehole in Ceylon and adults were collected in a

Malaise trap at an elevation of 200 feet.

In India, immatures were collected from ground pools and water-filled

ditches (Barraud 1934: 260) and-adults were taken in indoor shelters and biting

outdoors (Rao and Rajagopalan 1957: 10). Qutubuddin (1960: 358) collected

larvae from a rotten masonary well in West Pakistan. Carter (1948: 314),

working in Ceylon, collected adults from a cattle baited trap.

AEDES (AEDIMORPHUS) PUNCTIFEMORIS (LUDLOW)

(Figs. 11, 16, 17, 28, 44)

Stegomyia punctifemoris Ludlow 1921, Mil. Surg. 49: 690 (9).

Aedes Stegomyia) punctifemore Ludlow, Edwards 1922b, Indian J. med. Res.

10: 465.

Aédes (Finlaya ?) punctifemore Ludlow, Dyar and Shannon 1925, Insecutor

Inscit. menstr. 13: 75 (9).

64 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Aédes (Aédimorphus) punctifemore Ludlow, Dyar 1925, Insecutor Inscit.

menstr. 13: 217 (o); Edwards 1932, GeneralInsec., Fasc. 194: 168.

Aédes (Aédimorphus) punctifemore (Ludlow), Edwards in Barraud 1934, Fauna

Brit. India, Diptera 5: 443 (9).

Aedes (Aedimorphus) punctifemore (Ludlow), Bohart 1945, Navmed 580, p. 64

(o, 2*); Knight and Hull 1953, Pacif. Sci. 7: 459 (o'* Q).

Aedes (Aedimorphus) punctifemoris (Ludlow), Stone et al. 1959, Thomas Say

Found. 6: 196; Aslamkhan 1971, Biologia 17: 61 (o"*, 9).

FEMALE (Fig. 11). Head. Antenna dark brown, approximately equal

in length to proboscis, pedicel pale with a patch of broad silvery scales and a

few short fine brown hairs mesally, flagellomere 1 pale with a few small brown

scales; clypeus dark brown, bare; maxillary palpus brown scaled with tuft of

pale hairs at tip, approximately 0.18 length of proboscis; proboscis golden-

brown scaled with apical 0.25 and basal 0.10 dark brown, approximately 1. 33

length of femur I; vertex and sides covered with broad decumbent scales, dor-

sum with an anteromedian diamond-shaped light brown group and remainder

silvery -white; lateral surface with an anterodorsal triangular light brown patch

separated from lower brown area by a white stripe extending from eye margin

posteromesally to occiput; narrow white scales and numerous dark brown erect

forked scales on occiput, fewer scales on vertex which extend to ocular line.

Thorax. Scutal integument dark brownish-black; scutum covered with narrow

curved reddish-black scales, broad silvery scales forming patches on anterior

promontory area, scutal fossal areas (anterior, lateral and posterior areas

each with a circular patch), supra-alar area with a small patch anterior to and

one posterior to wing base, posterior medial scutal area, prescutellar space

along posterolateral margins and a few scattered broad white scales among

posterior dorsocentral bristles; scutellum with a patch of broad silvery scales

on each lobe (one Philippine specimen with broad pale brown scales on median

lobe); median anterior promontory, acrostichal, dorsocentral (anterior and

posterior), scutal fossal (anterior, 3-4 lateral and 1-2 posterior), supra-

alar, several posterior medial scutal, 1 postalar callar and scutellar (lateral

and median) bristles reddish-black and well developed, others absent; pleural

integument dark reddish-brown; antepronotum with a few broad silvery scales,

several dark bristles; postpronotum with narrow curved reddish-black scales

dorsally and a posterior patch of broad silvery scales, 4-5 posterior dark

bristles; propleuron with a patch of broad silvery scales, several dark bris-

tles; postspiracular area with 4-5 dark bristles; subspiracular area with 2

small patches of broad silvery scales; mesepisternum with a small upper and

a small posterior patch of broad silvery scales, several upper and posterior

dark bristles, lower ones shorter; prealar knob with several dark bristles;

paratergite covered with broad silvery scales; mesepimeron with a small

patch of broad silvery scales and 5-6 dark bristles on upper area; other

pleural areas bare. Legs (Fig. 16). Coxae I-III each with several dark

bristles, I with broad brown scales on anterior and lateral surfaces and a

dorsal patch of silvery scales, II with an anteromedian silvery patch of scales

with a few brown ones below, III with a few anteroventral pale brown scales;

trochanters I-III each with brown scales; femora I-III brown, each with a dor-

soapical silvery spot and several broad silvery scales scattered over anterior

surface; femur I with posterior brown with several broad silvery scales inter-

mixed; femora II, III each with basal 0.75 mainly brownish-white scaled, re-

mainder brown; tibiae I-III brown, each with a dorsoapical silvery spot and

several broad silvery scales scattered over surfaces, mainly on anterior sur-

Reinert: Aedes (Aedimorphus) in Southeast Asia 65

face; tarsi I-III brown; posttarsi I-III each with 2 ungues, I, II equal, each

bearing a tooth, III equal, simple. Wing. Dorsal veins covered with moder-

ately broad brown scales with a silvery patch at base of costa; alula with nar-

row brown scales along fringe; 2-3 remigial bristles. Halter. Pedicel pale

brown, capitellum brown scaled. Abdomen. Terga brown; tergum I with a rectangu-

lar patch of silvery-white scales on laterotergite; terga II-VII each with a small

laterobasal patch of silvery scales; sterna brown scaled; terga and sterna with

numerous golden-brown bristles, mostly along posterior margins. Genitalia

(Fig. 28). Tergum VII index 0. 78-0. 85; sternum VIII index 0. 82-0. 86; ter-

gum IX bilobed with 3-6 bristles on each lobe, index 1.05-1.27; insula tongue-

like, covered with minute setae and with 5-7 small tuberculi on apical 0. 25;

lower vaginal lip narrow, moderately pigmented, covered with minute setae;

upper vaginal lip narrow, heavily pigmented, covered with minute setae; upper

vaginal sclerite large, heavily pigmented, triangular shaped; postgenital lobe

moderately long, apex with a moderately deep median indentation, 6-10 bris-

tles on each side of midline, covered with minute setae, dorsal PGL index

1, 26-1.38, ventral PGL index 2.06-2.34; cercus moderately long, 0. 80-0.90

extended from segment VII, narrow, apex sharply rounded, numerous bristles

on dorsal and lateral surfaces, index 2.'76-3.06, cercus/dorsal PGL index

2.55-2.97; 3 spherical, pigmented seminal capsules, 1 large and 2 slightly

smaller ones.

MALE (Fig. 11). Similar to female in general habitus. Head. Maxil-

lary palpus brown, longer than proboscis by length of apical segment. Thorax.

Propleuron with only 1-3 dusky-white scales, numerous dark bristles. Legs

(Fig. 17). Posttarsi I-III each with 2 ungues, I, I with ungues unequal, each

bearing a tooth, III unequal, simple. Abdomen. Tergum I brown with a patch

of silvery scales on laterotergite; terga II-VI brown each with a small latero-

basal patch of silvery scales; tergum VIII silvery; sterna brown with an incom-

plete white stripe on lateral margins of sterna IN-VIN. Genitalia (Fig. 44),

Tergum IX strongly bilobed with 6-7 bristles on each lobe, entire surface

covered with minute spicules; gonocoxite long and moderately broad, dorsal

surface with a few long stout bristles along lateral margin and scattered mod-

erately long and short ones over remainder of area, very short fine bristles

along tergomesal margin, lateral surface with long stout and a few moderately

long bristles, ventral surface with long stout bristles on distal 0.45 and mod-

erately long ones along sternomesal margin, very numerous on distal 0. to.

scattered short bristles over remainder of area, scattered scales on lateral

and ventral surfaces; gonostylus with pedicel moderately long, narrow and

somewhat incurved, distal 0.46 expanded into a mesal lobe and a lateral short

tapering apically pointed horn attached approximately 0.72 from base with a

short fine hair at apex, mesal expanded lobe with a moderately long, flattened,

curved, apically blunt gonostylar claw attached mesally near middle, 2 short

stout bristles at apex and 2-4 slightly smaller ones along apical margin, 3-4

short fine hairs on tergal surface proximad of claw; basal mesal lobe short and

rounded apically, distal 0.45 with 4-6 short bristles, entire surface covered

with short hair-like spicules; proctiger short, paraproct with a subapical thumb-

like process, cercal setae absent; phallosome with aedeagus of typeI with 2

lateral plates connected basally, each plate with 5-6 short blunt lateral teeth on

distal 0.56 and covered with a dorsal flap, paramere long, approximately 0. 80

length of lateral plate; sternum IX large, entire surface covered with minute

spicules, 4-5 bristles near center.

PUPA and LARVA. Not known.

TYPE DATA. Stegomyia punctifemore Ludlow, holotype female, Fort

66 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Wm. McKinley, Rizal, Luzon, PHILIPPINES, 20 August 1921, in United States

National Museum (Natural History).

DISTRIBUTION. Specimens examined--2 males and 22 females from

the following locations:

BANGLADESH (East Pakistan). Dinajpur, Thakurgaon.

INDIA. Bihar, Gaya.

PHILIPPINES. Luzon, Wack-Wack; Rizal, Camp Nichols, Ft. Wm.

McKinley.

Other distribution.

BANGLADESH (East Pakistan). Dinajpur, Thakurgaon, Akcha, Mad-

argonj (Aslamkhan and Wolfe 1971: 31).

PHILIPPINES. Samar, Osmena (Knight and Hull 1953: 460).

TAXONOMIC DISCUSSION, Aedes punctifemoris is a very distinct

species and is easily recognized by the following features: vertex covered

with broad scales; scutum with patches of broad white scales; antepronotum

and postpronotum each with a few broad silvery-white scales; postspiracular

area without scales; femora and tibiae with broad silvery-white scales inter-

mixed with the brown ones; and tarsi dark scaled.

BIOLOGY. Adults were taken in light traps in the Philippines.

Aslamkhan (1971: 64) collected this species in abundance after monsoon

rains during June and July in Bangladesh (East Pakistan). He collected only

11 females resting in houses while 129 were taken biting man after dusk. One

hundred twenty-seven specimens were dissected and found to be negative for

Wuchereria bancrofti.

AEDES (AEDIMORPHUS) VEXANS VEXANS (MEIGEN)

(Figs. 14, 16, 17, 31, 33, 47, 57, 68)

Culex vexans Meigen 1830, Syst. Beschr. zweifl. Ins. 6: 241 (2); Theobald

1903, Monogr. Cul. 3: 404 (c*, 9*); Giles 1902, Handb., p. 416 (c*,

?*); Blanchard 1905, Moust., p. 309 (o'*, ?*); Theobald 1905, Genera

Insec., Fasc. 26: 26.

Culex nocturnus Theobald 1903, Monogr. Cul. 3: 159 (?*); Theobald 1905,

Genera Insec., Fasc. 26: 25; Theobald 1910, Monogr. Cul. 5: 324.

NEW SYNONYM.

Culicada minuta Theobald 1907, Monogr. Cul. 4: 338 (9*); Theobald 1910,

Monogr. Cul. 5: 294; Brunetti 1912, Rec. Indian Mus. 4: 462.

Culicada eruthrosops Theobald 1910, Monogr. Cul. 5: 299 (9*); Brunetti 1912,

Rec. Indian Mus. 4: 462.

Culex nocturnus var. niger Theobald 1913a iz Sarasin and Roux, Nova Caledonia

A., Zool. 1: 164 (A).

Ochlerotatus vexans Mg., Edwards 1917, Bull. ent. Res. 7: 218; Brunetti 1920,

Rec. Indian Mus. 17; 135; Senior-White 1923, Cat. Indian Insects, Cul.,

ps 83.

Aédes (Ecculex) vexans (Mg.), Edwards 1921, Bull. ent. Res. 12: 322.

Aedes (Ecculex) vexans Mg., Edwards 1922b, Indian J. med. Res. 10: 467.

Aédes (Aedimorphus) vexans Meigen, Edwards 1924, Bull. ent. Res. 14: 372;

Brug 1924, Bull. ent. Res. 14: 436 (L); Dyar and Shannon 1925, Insecutor

Inscit. menstr. 13: 77; Buxton and Hopkins 1925, Bull. ent. Res. 15:

300 (L); Borel 1930, Coll. Soc. Path. exot. Monogr. 3: 275 (o'*, 9);

Lee 1944, Atlas Mosq. Larvae Australian Region p. 72 (L*); Natvig 1948,

Suppl. Norsk ent. Tidsskr. 1: 412 (c‘*, 9, L*).

Reinert: Aedes (Aedimorphus) in Southeast Asia 67

Aedes (Aedimorphus) vexans (Meigen), Barraud 1928, Indian J. med. Res. 15:

660 (o'*, 2*); Penn 1949, Pacif. Sci. 3: 60 (P*); Horsfall and Craig

1956, Ann. ent. Soc. Am. 49: 370 (E*); Stone et al. 1959, Thomas Say

Found. 6: 198; Assem and Bonne-Wepster 1964, Zool. Bijdr. 6: 99

(o, 9, L*); Mohrig 1967, Angew. Parasit. 8: 80 (?*); Kalpage and

Brust 1968, Can. J. Zool. 46: 711 (E*); Pao and Knight 1970, J. Geo.

ent. Soc. 5: 115 (L*).

Aédes (Aédimorphus) vexans (Meigen), Barraud 1934, Fauna Brit. India,

Diptera 5: 253 (o*, 2, L).

Aedes (Aedimorphus) vexans Meigen, Bohart 1945, U. S. Navmed. 580, p. 64

(o*, L); Bonne-Wepster 1954, Doc. med. Geogr. Trop. 6: 239 (o%, 2,

L*),

Aedes (Aedimorphus) vexans nocturnus (Theobald), Bohart and Ingram 1946,

U. S. Navmed. 1055, p. 15 (o', 9, P*, L); Yamaguti and La Casse

1950, Mosquito Fauna Guam, p. 73 (o'*, 9*, L*); Knight and Hull

1953, Pacif. Sci. 7: 460 (c*, 9, L*); Bohart 1956 (1957), Insects

Micronesia 12: 59 (o’*, 2, L*); Stone et al. 1959, Thomas Say Found.

6: 198; Lein 1962, Pacif. Insects 4: 627.

Aedes (Aedimorphus) nocturnus (Theobald), Belkin 1962, Mosquitoes S. Pacif.

p. 427 (o'*, 9, P*, L*); Stone 1963, Proc. ent. Soc. Wash. 65: 130;

Belkin 1965, Contr. Am. ent. Inst. 1(4): 23.

“

FEMALE (Fig. 14). Head. Antenna dark brown, approximately 1.02

length of proboscis, pedicel pale brown with several small white scales and a

patch of short fine dark hairs mesally, flagellomere 1 pale with a few small

pale scales; clypeus light brown, bare; maxillary palpus brown scaled with

white scales at apex, base of segment 4 and occasionally along outer lateral

margin of segments 3-4, approximately 0.18 length of proboscis; proboscis

brown scaled dorsally with scattered white scales on basal 0.75, ventral and

lateral surfaces golden-white scaled from near base to apical 0. 25 which is

entirely dark, approximately 1.22 length of femur I; vertex with dorsum

covered with narrow curved decumbent scales, an anteromedian diamond-

shaped brown group and the remainder white; lateral surface covered with broad

scales, an anterodorsal black patch with white ones below and a second lower

black patch in some specimens, occasionally a large patch of dusky scales

anterior to antepronotum, small narrow decumbent white scales on occiput

and posterior to the anterodorsal black patch; numerous golden and brown erect

forked scales on occiput and vertex extending anteriorly to ocular line. Thorax.

Scutal integument dark brownish-black; scutum covered with narrow curved

reddish-black scales, scutal patterns vary considerably but usually with nar-

row curved white scale patches on median anterior promontory area, scutal

fossal areas extending from anterior area along margin and onto lateral area,

small patch at scutal angle, supra-alar area above posterior of paratergite

extending to posterior of wing base, posterior medial scutal area, along lateral

margins of prescutellar space and among posterior dorsocentral setae, an in-

distinct patch of narrow dusky-white scales lateral to posterior dorsocentral

setae above supra-alar white patch; scutellum with a patch of narrow curved

white scales on each lobe; median anterior promontory, acrostichal, dorso-

central (anterior and posterior), scutal fossal (anterior, 4-7 lateral, 1-2 me-

dian and 2-3 posterior), supra-alar, several posterior medial scutal, 1 postalar

callar and scutellar (lateral and median) bristles reddish-black and well devel-

oped; pleural integument dark to light brown; antepronotum covered with narrow

curved white scales with a few broad white ones anteriorly, several dark bris-

68 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

tles; postpronotum dark, completely covered with narrow curved scales, a

small patch of lower posterior white ones and the remainder reddish-black,

some specimens also with a few lower broad white scales, 7-10 dark posteri-

or bristles; propleuron with a patch of broad white scales, several golden bris-

tles; prosternum with narrow white scales and 1-3 bristles; postspiracular area

with a large patch of narrow curved or moderately broad scales or both, 7-9

golden bristles; subspiracular area with 2 patches of broad white scales, lower

one longer; mesepisternum with an upper and a posterior patch of broad white

scales, several upper and posterior golden bristles, lower ones shorter; pre-

alar knob with a small patch of long broad white scales on lateroventral mar-

gin, several golden bristles; paratergite with a patch of white scales on lateral

margin, scales vary in shape, some specimens with narrow curved ones, some

with moderately broad ones and some with both types; mesepimeron with a

patch of broad white scales and several golden bristles on upper area; other

pleural areas bare. Legs (Fig. 16). Coxae I-III each with several golden

bristles, I, II each with anterior surface covered with broad white scales, I

also with a few to a large patch of broad brown ones near middle, III with an

anterodorsal and an anteroventral patch of broad white scales; trochanters

I-III each with broad white scales; femora I-III each with anterior surface

brown with varying amounts of white scales intermixed and a dorsoapical white

spot, II with a broad longitudinal white stripe tapering from base to apical

0.25, I, II each with a narrow basal white band, in some specimens there are

only a few scattered white scales on I but there are always numerous ones on

II (Oriental Region); femora I-III each with posterior surfaces with a broad

longitudinal white stripe tapering from base to apical 0.25 and the remainder

brown with intermixed white scales, stripe dorsal on I and ventral on IJ, III;

tibiae I-III each brown with a few scattered white scales, a narrow basal white

band and posterior longitudinal white stripe from base to apex, stripe postero-

ventral on I; tarsi I-III brown, I with tarsomeres 2, 3 each with a basal white

band, tarsomere 1 usually with a small dorsobasal white spot and a posterior

longitudinal white stripe, tarsomere 4 with a dorsobasal white spot, II with

tarsomeres 1-3 each with a basal white band, tarsomere 4 with a dorsobasal

white spot, tarsomere 1 with a posterior longitudinal white stripe, II with

tarsomeres 1-5 each with a basal white band, tarsomere 1 with a few scattered

white scales; posttarsi I-III each with 2 ungues, I, II equal, each with a tooth,

III equal, simple, occasionally toothed. Wing. Dorsal veins covered with

moderately broad brown scales; costa with a small patch of white scales at

base and along its posterior margin from just before humeral cross vein to

apical 0.66, some specimens also with scattered white scales on radius and

median; ventral veins brown scaled; alula with narrow brown scales along

fringe; 2-3 remigial bristles. Halter. Pedicel pale, capitellum white scaled

with a few brown scales mesally. Abdomen. Terga brown; tergum I with a few

basomedian white scales, laterotergite with a rectangular patch of white scales;

terga II-VI each with a dorsobasal broad white band, posterior margins of

bands vary from being straight to notched mesally; terga VI, VII each with a

narrow apical white band, VII also with a pair of laterobasal triangular white

patches on dorsum, patches occasionally forming a notched basal band; tergum

VII with a few white scales on apical margin; terga II-VII each with a large

laterobasal patch of white scales, patches not connected with dorsal bands;

sterna creamy-white scaled with a few brown scales mesally on II-VI; terga

and sterna with numerous golden bristles, mostly along posterior margins.

Genitalia (Fig. 31). Tergum VII index 0. 79-0. 88; sternum VIII index 0.94-

0.99; tergum IX bilobed with 5-14 bristles on each lobe, index 0.90-1. 12;

Reinert: Aedes (Aedimorphus) in Southeast Asia 69

insula tongue-like, covered with minute setae; lower vaginal lip narrow, mod-

erately pigmented, covered with minute setae; upper vaginal lip narrow, heavi-

ly pigmented, covered with minute setae; upper vaginal sclerite large, moder-

ately pigmented, triangular shaped; postgenital lobe moderately long, apex with

a moderately deep median indentation, 5-10 bristles on each side of midline,

covered with minute setae, dorsal PGL index 0. 70-0.91, ventral PGL index

1. 52-1. 86; cercus moderately long to long, 0.90-1.00 extended from segment

VU, narrow, apex sharply rounded, numerous bristles on dorsal and lateral

surfaces, index 3.21-4.27, cercus/dorsal PGL index 4. 35-5. 22; 3 spherical,

pigmented seminal capsules, 1 large and 2 slightly smaller ones.

MALE (Fig. 14). Similar to female in general habitus. Head. Maxil-

lary palpus brown, segments 2-5 each with a dorsobasal white spot, longer than

proboscis by 0.75 of apical segment; vertex with anteromedian brown and an-

terodorsal black scale patches reduced. Thorax. Prosternum bare. Legs

(Fig. 17). Tibia Il with posterior white stripe broad; posttarsi I-III each with

2 ungues, I, II with ungues unequal, each bearing a tooth, III equal, simple,

occasionally toothed. Abdomen. Terga brown without lateral spots; tergum II

with basal white band, posterior margin of band straight; terga ITI-VII each with

a basal white band broad on lateral margins and deeply notched mesally, some

specimens also with a few median apical white scales; tergum VIII white, some

specimens with a few brown scales mesally; sterna white with an indistinct

median longitudinal brown stripe, stripe distinct on VIII. Genitalia (Figs.

33, 47). Tergum IX strongly bilobed with 6-11 bristles on each lobe, entire

surface covered with minute spicules; gonocoxite long and moderately broad,

dorsal surface with long stout bristles along lateral margin from base to apex,

short thin bristles along tergomesal margin from base to apex and similar ones

extending along base to lateral margin, lateral surface with long stout bristles

from base to apex, ventral surface with long stout bristles on distal 0.60 of

sternomesal margin, afew similar ones on apical 0.25 of area, short to mod-

erately long bristles on proximal sternomesal margin and scattered over re-

mainder of surface, scattered scales on lateral and ventral surfaces; gonosty-

lus approximately 0.69 length of gonocoxite, moderately broad throughout length

with distal 0. 20 tapering to a blunt point with 2-3 short fine hairs near apex,

laterotergal surface somewhat curved dorsally and covered with short hair-like

spicules on apical 0.50, gonostylar claw long, narrow, pigmented, with apex

blunt, 10-16 short fine hairs scattered over distal 0.60 of flattened area;

basal mesal lobe short with apical 0.25 rounded, somewhat expanded and bear-

ing 25-38 short bristles, entire surface covered with short hair-like spicules;

proctiger long, paraproct narrow with apex bluntly pointed and entire structure

strongly pigmented, cercal setae absent; phallosome with aedeagus of type II

with 2 lateral plates connected basally, each plate with 5-6 short to moderately

long, longitudinal lateral teeth with tergally curved apices and covered with a

dorsal flap, paramere long, approximately 0.94 length of lateral plate; sternum

IX large, entire surface covered with minute spicules, 4-5 bristles near center.

PUPA (Fig. 57). Chaetotaxy as figured and recorded in Table 9. Two

to three rows of spicules mesally along posteroventral margin of abdomen II.

Cephalothorax. Hair 5-C single to triple; 7-C with 2-5 branches; 8-C with 2-4

branches. Respiratory trumpet. Moderately pigmented; index 2. 70-3. 24,

average 2.96. Metanotum. Hair 10-C with 4-11 branches; 12-C single to triple.

Abdomen. Hair 5-I with 2-8 branches; 10-I single; 1-II with 5-14 branches;

4-II with 2-4 branches; 1-III with 3-5 branches; 6-VI single or double; 1-VI

single to triple; 6-VII with 2-5 branches; 9-VII with 2-4 branches; 11-VII single.

Paddle. Ovoid; with very minute serrations along basal 0.55 of outer margin;

70 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

tiny spicules along apical 0.45 of outer and apical 0. 25 of inner margins; mid-

rib does not reach apex; hair 1-P short, single; index 1.17-1.49, average 1.28.

LARVA (Fig. 68). Chaetotaxy as figured. Head. Specimens from

Southeast Asia have the frontoclypeus granulose; median mouth brushes pec-

tinate apically; hairs 1, 3, 8, 14-C single; 4, 15-C with 3-5(4) branches; 5, 9,

10, 13-C single to triple (2); 6-C single or double (1); 7-C with 7-11(9) bran-

ches; 11-C with 3-8(6) branches; 12-C with 3-9(5) branches; basal maxillary

hair single; mental plate with 27-32(29) teeth. Antenna. Short and moder-

ately pigmented; with small stout spicules scattered over shaft, more numer-

ous on basal 0.50; hair 1-A with 5-10(8) branches, inserted 0.36-0. 44 from

base; 2-A long; 3-A approximately 0.50 length of 2-A. Thorax. Hair 0-P with

4-9(6) branches; 1, 2, 5, 6,10, 12-P single; 3-P single to triple (2); 4, 9, 14-P

single to double (1), 7-P double or triple (3); 8-P with 3-7(4) branches; 11-P

with 2-4(3) branches; 1, 2, 4-M with 2-4(3) branches; 3, 5, 7, 10, 11, 12-M

single; 6-M with 5-8(5) branches; 8-M with 6-9(6) branches; 9-M with 5-8(6)

branches; 13-M with 4-9(4) branches; 14-M with 4-9(6) branches; 1, 4-T with

2-4(3) branches; 2-T with 2-4(2) branches; 3-T with 3-7(5) branches; 5, 10,

12-T single; 6, 11-T single or double (1); 7-T with 7-10(9) branches; 8-T with

4-7(5) branches; 9-T with 4-6(4) branches; 13-T with 5-9(6) branches. Abdo-

men. Hairs 1, 2-VIII on common basal plate; hairs 0, 14-VIII single; 1-VIII

with 5-12(8) branches; 2, 4-VIII double or triple (2); 3-VIII with 6-10(9) bran-

ches; 5-VIII with 7-10(9) branches; 6-III-VI long; comb of 7-12(8) scales ar-

ranged in 1-2 irregular rows, scales with a long stout pointed median spine and

short denticles along lateral margins of base; 1, 3-X single; 2-X with 5-8(7)

branches; ventral brush varies from 12 hairs on grid and 4 precratal ones to

14 hairs on grid and 3 precratal ones, usually with 13 hairs on grid and 3 pre-

cratal ones; saddle moderately pigmented, incompletely rings segment, with

small spicules along posterior margin, acus present; 4 anal papillae, long and

tapering to a blunt point. Siphon. Moderately pigmented with minute ridges

over entire surface; acus large; index 2. 11-3. 10; pecten with 13-18(16) teeth,

apical 2-3 teeth larger, smooth or with a minute basal denticle and wider

spaced than remainder which have a slender apical attenuated filament with 1-3

basal denticles; hair 1-S with 5-6(5) branches, inserted 0.54-0.69 from base.

, EGG. Shape. Variable, fusiform to spindle-shaped; greatest diameter

between anterior third and middle. Size. In microns, 648-756 by 171-216

(Horsfall and Craig), 614-713 by 199-224 (Myers) and 630-745 by 167-205 (Kal-

page and Brust). Color. Bronze. Chorion. Surface reticulation consisting

of a pattern of axially linear cells, hexagonal and polygonal in shape and 2. 5-

5 times as long as wide; cell walls raised. The above description is taken from

Horsfall and Craig (1956: 370), Myers (1967: 796) and Kalpage and Brust (1968:

711-712).

TYPE DATA. Culex vexans Meigen, holotype female, near Berlin,

GERMANY, Ruthe, in Museum National d'Histoire Naturelle, Paris, France;

Culex arabiensis Patton, syntypes male and female, Ulub Camp and Carter,

West Aden Protectorate, types nonexistent; Culex articulatus Rondani, ITALY,

location of type unknown; Aedes euochrus Howard, Dyar and Knab, Popcum,

British Columbia, CANADA, 2 August 1903, J. Fletcher, type no. 12057, in

United States National Museum (Natural History); Culicada eruthrosops Theo-

bald, holotype female, Trincomalee, CEYLON, November 1906, E. Green, in

British Museum (Natural History); Culex malariae Grassi, ITALY, location of

type unknown; Culicada minuta Theobald, holotype female, INDIA, Dr. Christo-

phers, in British Museum (Natural History); Culex nocturnus var. niger Theo-

bald, holotype female (?), Canala, NEW CALEDONIA, in. Naturhistorisches

Reinert: Aedes (Aedimorphus) in Southeast Asia 3:

Museum, Basel, Switzerland; Culex nocturnus Theobald, 2 syntype females,

Ba, FIJI ISLANDS, April-June, Hall, in British Museum (Natural History);

Culex parvus Macquart, Bordeaux, Gironde, FRANCE, type nonexistent;

Culex sylvestris Theobald, syntype female, Rondeau Provincial Park, Kent

Co., Ontario, CANADA, 16 September 1899, E. M. Walker, from the interi-

or of a sandy wood of white pine with a few hard woods, and syntype male,

De Grassi Pt., Lake Simcoe, Onfario, CANADA, 19 July 1899, E. M. Walker,

from grass and low herbs in a wood of maple, aspen, balsam and fir, both in

British Museum (Natural History).

DISTRIBUTION. Specimens examined--1, 331 males, 4, 149 females,

943 pupae, 1, 003 larvae and 534 individual rearings (52 pupal, 482 larval)

from the following locations:

AUSTRALIA. N.E. New Guinea, Morobe, Lae; Papua, Goodenough

Is., Vivigani.

AUSTRIA. Marchegg.

BURMA. Rangoon.

CAMBODIA. Ari Gsatr, Phom-Penh.

CANADA. Alberta; British Columbia, Prince George; Manitoba,

Aweme, Winnipeg; Ontario, De Grassi Pt., Lake Simcoe, Rondeau Provincial

Park, Kent Co.; Saskatchewan,

CAROLINE ISLANDS. IfalukIs., Yap Is., Calonia.

CEYLON. Hambantota, Kaude-ela Reservoir, Nuwara-Eliya, Trin-

comalee, Weligama.

CHINA. Yunnan; Tibet, Yatung.

FIJI. Cuvu, Lani, Lawaga, Nacaugai, Nausori, Penang, Soso, Suva.

FRANCE. Lyon, Lyon.

GERMANY.

HONG KONG. Sai Kung, Ho Chung.

INDIA. Assam, Chabua, Doom Dooma, Jorhat, Shillong, Tinsjuia;

Bengal, Darjeeling, Sukna; Kashmir, Srinagar; Madras, Coonoor; Punjab,

Ambala; Bombay Deccan, Tavargatti, Nilgiris, Palnis, Kodai Ranal.

INDONESIA. Celebes, Bwool, Molino, Paloe, Pare-Pare, Saleier;

Ceram, Honititoe, Lisiela, Paniwal, Piroe, Vatoenoeoe, Kairatu, Piru,

Warasuva; Flores; Java, Angki, Batavia, Gombong, Tjilatjap; Moluccas,

Morotai; New Guinea, Hollandia, Mambaye River, Oeta; Sumatra, Atjch,

Bengkoelen, Ranau; Timor, Atamboea; Amboina, Digoel River, Lisabata,

Mamoedjoe, Vabaena near Boeton.

IRAN. Enzebi.

IRAQ. Baghdad.

ITALY. Lucca.

MACEDONIA. Bajirli, Keulike.

MALAYSIA. Kedah, Serdang; North Borneo, Membakut; Pahang,

Bentong, Kg. Sertik; Perak, Kuala Kangsar, Lasah; Selangor, Puchong, Sabak;

Trengganu, Marang; Kg. Saban, Semporna, Knantan, Kota Belud, Timbang.

MARIANA ISLANDS. Guam, Pt. Oca; Saipan, Charan, Hashigaro,

Ronoa.

MARSHALL ISLANDS. Ebon Atoll, Eboni.

NEW CALEDONIA. Puebo.

NEW HEBRIDES. Espiritu Santo, Sarakata Valley, Mai Is., Segond

Channel.

PHILIPPINES. Camerines Sur, Iriga; Laguna, Balian; Leyte, Abuyog,

Burauen, Carigara, Dulag, Gabas, JinamocIs., Mahaptag, Tacloban, Tar-

ragona, Tolosa; Luzon, Wack-Wack; Manila; Mindanao, Bunwan, Dargo,

72 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Kabakan, Ludlow Barracks, Parang, Pettit Barracks, Zamboanga, Torrey

Barracks; Mindoro, Camiuawit Point, San Jose; Mountain Province, Trinidad;

Palawan, Panakan, Panitian, Quezon, Puerto Princesa, Tarusan; Pampanga,

Camp Stotsenberg; Pangasinan Bayambang, San Fabian, Sison, Tayug; Rizal,

Alabany, Camp Nichols, Ft. Wm. McKinley, Pasig River; Samar, San

Antonio; Calaccad, Camp Eldridge, Iloilo, Libjo, Los Banos, Port of Parany,

Sanga Sangals., Lapit-Lapit, Tawi Tawils., Batu Batu, Tayabas, Infanta.

ROMANIA. Bessarabia, Comana Wlasca.

SAMOA. Aeipata, Tutulla, Upolu Is.

SOUTH VIETNAM. Binh Dinh, An Tuc, Binh Khe; Binh Duong, Lai

Khe; Bien Hoa, Long Binh; Cam Ranh, Cam Ranh Bay; Da Nang, Da Nang;

Gia Dinh, Saigon; Hau Nghia, Cu Chi; Khanh Hoa, Nha Trang; Pleiku, Pleiku;

Quang Tri, Ba Long; Thua Thien, Cu Lai, Phu Bai; Tuyen Duc, Da Lat.

SWEDEN. Stockholm.

TAIWAN. Haping, Hwalien, Kelaipao, Shyolin.

THAILAND. Ayutthaya, Bang Pa In; Chiang Mai, Ban Chang Kien,

Ban Hua Muang, Ban Lang Ka, Ban Nong Pa Khrang, Chang Puak, Chiang

Mai, Chom Thong, Doi Chom Cheng, Doi Sutep, Hod, Huey Chang Kien,

Mea Sanan, Mea Tang, Thanon To Doi Saket; Chon Buri, Bang Phra; Chumphon,

Muang; Kanchanaburi, Ban Sai Yok; Kvabi, Ban Mai Kaen Tai; Krung Thep,

Bangkok; Lampang, Nam Mae Nu Rua; Nakhon Ratchasima, Ban Khanong Phra

Nua; Nakhon Sawan, Ban Kaeng; Nakhon Si Thammarat, Ban Rim Thanon;

Narathiwat, Khau Lau; Phrae; Rayong; Songkhla, Haad Yai; Surat Thani, Ban

Taling Ngam, Ban Thong Phlu, Chom Thong, Ko Samui.

TONGA ISLANDS. Tengatubo.

U. S. A. Arizona; Arkansas; California; Colorado; Florida; Georgia;

Hawaii; Idaho; Illinois; Indiana; Iowa; Kentucky; Louisiana; Maine; Maryland;

Massachusetts; Michigan; Minnesota; Mississippi; Missourt; Montana;

Nebraska; New Jersey; New Mexico; New York; North Carolina; North Dakota;

Ohio; Oklahoma; Oregon; Pennsylvania; South Carolina; South Dakota; Tennessee;

Oregon; Pennsylvania; South Carolina; South Dakota; Tennessee; Texas; Utah;

Virginia; Washington; Wisconsin; Washington, D.C.

U. S. S. R. Siberia, Amagu, Judia River, Irkutsk, Okeanskya; Saratow.

Other distribution.

ADEN (Edwards 1941: 195); Ulub, Aden, Aden Hintherland (Mattingly

and Knight 1956: 119).

ALGERIA. Chebli, Ourgla, (Senevet and Andarelli 1954: 326); La

Reghaia (Seneret 1936: 447).

AUSTRALIA. New Guinea, Camadodo, Milne Bay; Papua (Penn 1949b:

61); New Guinea, Finschhafen, Saidor (Bick 1951: 412); New Guinea, Wewak-

Maprik Road Area (Standfast 1967: 192); Papua, Minj (Peters and Christian

1963: 41).

AUSTRIA (Theobald 1901a: 405).

BELEP ISLAND. Avi, Uala (Laird 1956: 21).

BRITISH HONDURAS (Belkin and Heinemann 1971: 27).

CANADA. Nova Scotia; Prince Edward Island; Quebec; Yukon (Car-

penter and La Casse 1955: 265); Ontario, Aberfoyle, Algonquin Park, Ancas-

ter, Brampton, Camp Borden, Cochrane, Elora, Galt, Kingston, London,

Toronto, (Steward and McWade 1960: 157); New Brunswick, Younghall; Oniar-

io, Ottawa, White River, Dryden, Kenora; Manitoba, Winnipeg Beach; Sas-

katchewan, Moose Jaw, Saskatoon, Prince Albert; Alberta, Red Deer,

Lamoral, Lochearn, Calgary, Banff, Lake Minnewanka; British Columbia,

Kalso, Ainsworth, Mission, Chilliwack, Whonnock (Dyar 1921: 115).

Reinert: Aedes (Aedimorphus) in Southeast Asia 73

CAROLINE ISLANDS (Bohart 1956: 59, Belkin 1962: 429).

CHINA. Hainan Island (Chu 1958: 109); Kuangtung, Swatow (Chung and

Lin 1929: 404); Manchuria, Hailung (Chin 1936: 23); Hunan (Chang 1939: 61);

Tibet (Buxton and Hopkins 1927: 91).

CORSICA ISLAND. Debbie, Calvi, Cateraggio, Prunette, Ghisonaccia

(Aitken 1954: 480).

CZECHOSLOVAKIA (Kramar 1958: 224); Moravia, Vranovice (Palicka

1967: 72); Lower Dnube River Basin (Trpis 1962: 112); Slovakia (Vostal 1963:

62).

DENMARK. Bondernes, Egehoved, Naebengen (Nielsen and Nielsen

1963: 147); Sjoelland (Natvig 1948: 418).

ELLICE ISLANDS (Iyengar 1955: 46, 1960: 67); Nui Island, Niuafoo

(Bohart and Ingram 1946: 16); Funafuti, Old airstrip(Laird 1955: 297, 1956:

38); Nui Island; Niutao Island (Buxton and Hopkins 1927: 91).

FIJI (Theobald 1903a: 160, Bohart 1956: 59, Lee 1944: 72, Paine 1943:

22, Edwards 1924: 372, Iyengar 1955: 44); Kandavu, Makongai, Taveune,

Vanua Levu (Belkin 1962: 429); Viti Levu, Nandarivatu (Laird 1956: 39).

FINLAND. Helsingfors, Tvarminne Zoological Station (Natvig 1948:

418).

FRANCE. Alsace (Li and Ulu 1933: 105, Callot and Vermeil 1948: 334);

Gray; Gave de Pau Valley; Creuse, LaCelle-Dunoise; Lyon (Seguy 1921: 25);

Jura Mts. (Callot 1956: 182).

FRIENDLY ISLANDS (Stone et al. 1959: 198).

GERMANY. Bavaria (Seguy 1921: 25); Bavaria, Dachau, Wartaweil,

Ammersee Lake, Puchheim, Graz (Kuhlhorn 1954: 6); Elbe, Rhein and Oden

Rivers (Mohring 1965: 4).

GILBERT ISLANDS (Iyengar 1960: 67); Tavawa Atoll, Bairiki (Laird

1955: 297).

GREAT BRITAIN. England, Hants, Hayling Island; Herts, Tring;

Middlesex, Finchley; Sussex, Arundel; Wales, Meronethshire, Dolgelly

(Marshall 1938: 206); Cambridge, Coe Fen; Shotover near Oxford (Lang 1920:

85); England, Wimbledon, (Mattingly 1950: 156; England, Suffolk, Mildenhall

(Edwards 1912: 195).

GREECE. Macedonia (Pandazis 1935: 15).

GUATEMALA. Guatemala City (Belkin and Heinemann 1971; 27.

HUNGARY. Balaton Lake Area (Mihalyi et al. 1952: 336); Matrafured,

Gayateto, Parad, Paradfurdo, Pisztrangos-to, Szilvasvarad, Bukk-fennsik,

Miskolctapolca, Kacsfurdo, Sikonda-furdo, Budoskut, Orfu, Abaliget, Komlo,

Sopron Area (Mihalyi et al. 1955: 348-362).

INDIA. Madras City (Senior-White 1923: 83); Calcutta (Li and Wu 1933:

105); Poona City, Manjri, Baramati (Rao and Rajagopalan 1957: 10); Assam,

Golaghat, Khumtai, Nongpoh; Central Provinces, Pachmari; Northwest Fron-

tier, Kohat, Abbottabad; United Provinces, Bhowali, Naini, Tal (Barraud 1928:

661); Mardas, North Arcot (Reuben 1971: 120); Nilgiri Hills, Kodaikanal (Bar-

raud 1928: 662); Palni Hills (Barraud 1934: 255); Coonoor (Patton 1922: 67);

Karachi (Hicks and Chand 1936: 520).

INDONESIA. Celebes, Banggaai (Brug and Haga 1923: 639); Bangsai

(Haga 1924; 830); Tervnate (Li and Wu 1933: 105); New Guinea, Kaimana,

Manokwari, Merauke, Sorong, Tjof Island (Assem and Bonne-Wepster 1964:

101); Java, Weltevreden (Brug and Haga 1923: 639); Borneo (Edwards 1922b:

467); Ngabang (Haga 1924: 830); Sumatra, Atchin, Blang Kedjeren, Kotta

Tjane, Takengon; Benkoelen, Air Prioekan; Djambi, Moeara Tebo; Krakatau

74 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

(Brug and Edwards 1931: 258); Soemba, Pajeti; Timor, Koepang (Haga 1924:

830); (Brug 1925: 668); Sumba; Molaccas (Bonne-Wepster 1954: 241); Bengaai;

Madoera (Brug 1924a: 36); Kali Anget (Brug and Haga 1923: 639).

ITALY (Blanchard 1905: 310); Taranto (Li and Wu 1933: 105); Sardinia

Island, San Vero Milis, Cabras, Orbai, Siliqua (Aitken 1954: 480); Govrizia,

Gorizia (Balducci et al. 1968: 457),

LIBYA (Hamon et al. 1966: 378).

LOYALTY ISLANDS. Ouvea, Mare (Belkin 1962: 429).

MALAYSIA. Selangor, Ulu Gombak (McDonald and Traub 1960: 100);

Sarawak (Moulton 1914: 47).

MARIANA ISLANDS (Bohart 1956: 59, Iyengar 1955: 45); Guam; Saipan,

Magicienne Bay (Bohart and Ingram 1946: 41); Guam, Agat Bay, Dededo Ordot,

Hilaan Point, Inarajan, Merizo, Nimitz Beach, Umatac (Hull 1952: 1291); Guam

(Reeves and Rudnick 1951: 641, Yamaguti and La Casse 1950: 73); Umatac (Hu

1953: 125).

MARSHALL ISLANDS. Ebon Island, Ebon Atoll (Bohart 1956: 59, Bo-

hart and Ingram 1946: 22).

MAURITANIA. Assaba, Diaguili; Gorgol, Belnabe, Kaldi, Rindiaw,

Silla (Hamon et al. 1966: 373).

MAURITIUS ISLAND (Li and Wu 1933: 105).

MEXICO. Chichahua, Ciudad Juarez (Martini 1935: 50); Guanajuato;

Jalisco; Michoacan; Queretaro (Vargas 1956: 21).

MOROCCO. Tazi Khemisset, Northern portion of country (Senevet

and Andarelli 1954: 326).

NETHERLANDS (Blanchard 1905: 310, Theobald 1901a: 405).

NEW CALEDONIA. Houailon (Li and Wu 1933: 105, Buxton and Hop-

kins 1927: 91, Edwards 1924: 372); Tontouta (Laird 1956: 20); Ile des Pins

(Belkin 1962: 429); Noumea (Williams 1943: 211); Bonde (Rageau and Hamon

1957: 377).

NEW HEBRIDES (Iyengar 1955: 43, Knight and Hull 1953: 462); Espiritu

Santo (Perry 1946: 16); Efate, Emae (Belkin 1962: 429); Aneityum, Anelganhat;

Futuna, Mission Bay (Laird 1956: 23).

NIGERIA. Kano, Kano; Oyo, Ibadan (Kumm 1931: 65, 73).

NORWAY. Os, Ringebu (Natvig 1948: 418).

PALAU ISLANDS (Stone et al. 1959: 198).

PHILIPPINES. Bulaca,; Tunkong Manga; Mindanao, San Ramon; Pala-

wan, Bacungam, Iwahig Penal ‘Colony, Balsahen and Tacburos Rivers; ‘Samar,

Osmena; Zambales, Olongaop, Subic Bay (Knight and Hull 1953: 462); Tunkulan

(Edwards 1929: 5); Clark AirBase, Luzon, Pampanga (Pippin 1965: 185; Dowell,

Libay and Baisas 1965: 15); Mountain, Calaccad (Rozeboom and Cabrera 1964:

21).

POLAND. Warsaw (Lukasiak 1958: 770, 1961: 389, 1964: 87); Wroclaw,

Kudawa (Lukasiak 1955: 301).

ROTUMA ISLAND (Belkin 1962: 429); Oinafa Village (Belkin 1965: 13).

REPUBLIC OF MAURITANIA. Assaba, Diaguili; Gorgol, Belnabe,

Kaedi, Rindiaw, Silla (Hamon et al. 1966: 373).

RUMANIA. Bukovina, Upper Prut River (Lukasiak 1959: 358).

SAUDI ARABIA. Jeddah (Edwards 1941: 195); Madruga (Mattingly and

Knight 1956: 119).

SCANDINAVIA (Blanchard 1905: 310, Theobald 1901a: 405).

SAMOAN ISLANDS. Lalomanu, Laulii, Mulifauna, Aleipata (Buxton

and Hopkins 1927: 91); Savaii (Belkin 1962: 429, Bohart and Ingram 1946: 16).

SOUTH AFRICA. Tyvansvaal, Warmbaths District (Muspratt 1955: 172).

Reinert: Aedes (Aedimorphus) in Southeast Asia do

SOUTHERN COOK ISLANDS. Rarotonga (?) (Belkin 1962: 429).

SOUTH VIETNAM (Borel 1930: 275); Bien Hoa, Phan Rang, Phu Cat,

Pleiku, Tan Son Nhut, Tuy Hoa (Parrish 1968a: 2-5, 1969: 554); Chu Lai,

Khe Sahn, Zui Nhon (Grothaus et al. 1971: 21).

SPAIN. El Escorial, Gibraleon, Mahon, Orense, Padul, Vigo, Guadar-

rama, Villacastin, Trobajo, Villadango, Sieteiglesias, Laguna, Motril, Ped-

rena, Villaverde (Clavero 1946: 22); Elche, Madrid, San Fernando (Seguy 1921:

25).

SUDAN. Lado, Bundle-Hierallah Area, Khor Nambiri, Nyumbe, Yidu,

Such River (Theobald 1913b: 596); Khartoum (Lewis 1958: 151); Meidob-Kassala

Area (Lewis 1947: 561); Jebelein (Lewis 1948: 152); Dueim, Abu Usher, Abu

Ideima, Ezeirgab, Omdurman, Eilafun, Wadi Sidna, Hassa Heissa, Kassala,

Kosti, Argo (Lewis 1945: 11); Abu Tabar, Danagla, Dilling, Dongola, El

Fasher, Gereif, Khatmiya, Luing, Nofalab, Rahib Wells, Shambat, Uzeirgab,

Zalingei (Lewis 1955: 168); Darfur, El Fasher (Abbott 1948: 43); Kadaro,

Masiol, Wad Medani, Khartoum (Edwards 1941: 195); Bahr-el-Jebel (Theobald

1907: 196).

SWEDEN. Brunnby, Lerhamn, Ol, Raa, Vickleby (Natvig 1948: 418).

TAIWAN. Tainan, Hsinhua; Taipei, Tamshui (Lien 1962: 628); Takao

(Edwards 1921a: 629); Ching Chaun Kang Air Base (Reisen et al. 1971: Table

17); Chao Chow, Hsin She (Hu and Grayston 1962: 140).

TANZANIA. Zanzibar, Pemba Island (Li and Wu 1933: 105).

THAILAND. Trang (Causey 1937: 414); Ban-U-Tapao, Nakhon Phanom,

Takhli, Udorn (Parrish 1968b: 2-4); Chieng Mai, Doi Pui Mt. (Scanlon and

Esah 1955: 139); Don Muang Air Base, Ubon (Reisen et al. 1971: Tables 6

and 7); Kanchanaburi, Sangkla-buri, Lai-nam, Ni-thae (Harinasuta et al. 19770:

241).

TOKALAU ISLANDS (Belkin 1962: 429).

TONGA ISLANDS (Knight and Hull 1953: 462, Iyengar 1960: 67); Nukua-

lofa (Li and Wu 1933: 105, Edwards 1924: 372, Buxton and Hopkins 1927: 91);

Niutao, Niuatobutabu (Bohart and Ingram 1946: 16); Tongatabu, Fanga Uta

Lagoon, Kuku Alofa (Laird 1956: 37); Niuafoou (Belkin 1962: 429).

TUNISIA. Oglet Hachina (Senevet and Andarelli 1954: 326); Nefzaoua

(Senevet et al. 1954: 273).

TURKEY (Parrish 1959: 266).

U. A. R. Egypt (Li and Wu 1933: 105).

U. S. A. Connecticut; Delaware; Kansas; New Hampshire; Rhode

Island; Vermont; West Virginia; Wyoming (Carpenter and La Casse 1955: 264);

Hawaiian Islands, Kauai Island, Oahu Island (Joyce and Nakagawa 1963: 2773).

U. S. S. R. (Theobald 1901a: 405); Asov, Woronesch (Stschelkanovzev

1926: 134); Southern Karelia (Natvig 1948: 418); Widely distributed from the

Siberian forest zone to the southern boundary and in the east to Sakhalin and

Kamchatka; sporadically found in the north in the Nenetsky National District

along the lower course of the Pechora River (Monchadskii 1951: 234); Kaza-

khstan, Siberia (Gutsevitch et al. 1970: 273).

WALLIS ISLANDS. Uea (Belkin 1962: 429).

WESTERN SAMOA. Ubolu, Apia (Laird 1956: 28).

WEST INDIES (Li and Wu 1933: 105).

WEST PAKISTAN. Lahore (Ansari 1959: 25).

YUGOSLAVIA. Skoplje (Li and Wu 1933: 105).

Natvig (1948: 418-420) lists a distribution for vexans vexans occurring

outside Denmark and Fennoscandia.

Graham (1939: 213) identified a single collection of larvae as vexans

76 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

that was found "in a tin of water jammed among rocks just above high tide at

Russell, '' New Zealand. Belkin (1968: 113) thinks this collection is probably

Aedes (Finlaya) notoscriptus (Skuse) and not vexans.

TAXONOMIC DISCUSSION. Aedes vexans vexans is similar in the

adult habitus and male genitalia to vexans nipponii, davidi, stenoetrus and

syntheticus. It has several smaller differences from vexans nipponii but the

most pronounced one is the scaling of the abdominal terga (see couplet 11 of

the adult key for this separation). Aedes vexans vexans, vexans nipponti

and stenoetrus have femora II with a number of white scales randomly inter-

mixed with brown ones on the anterior surface while in syntheticus the anteri-

or surface of femur II is brown with a median row of distinct white spots. The

proboscis of stenoetrus is completely covered with dark brownish-black scales

while the other 3 members of the group have white scales on the proboscis

(usually in great numbers ventrally). Aedes stenoetrus probably has a re-

stricted distribution in the mountains of Ceylon and southern India at elevations

above 4, 000 feet while vexans vexans has a very wide distribution and occupies

a great variety of elevations. Differences from davidi are discussed under that

species. The basal mesal lobe of the male genitalia of syntheticus has 4 flat-

tened blade-like and 1 long bristle while those of vexans vexans and vexans

nipponii have 25-38 short bristles and stenoetrus has 118-128 bristles. The

adult habitus of vexans vexans has a semblance to caecus which is discussed

under that species. Aedes (Ochlerotatus) vigilax is very similar to vexans

vexans and vexans nipponii in the adult habitus and is often confused with them

but can be separated from these 2 species by the presence of a small patch of

broad white scales on the mesepisternum immediately below the prealar knob

and by the absence of scales on the subspiracular area. The aedeagus of the

male genitalia of vexans vexans and vexans nipponii is toothed while vigilax

has a simple tube.

The female genitalia of vexans vexans, vexans nipponii, syntheticus

and stenoetrus are very similar except that the latter species has a number of

scales scattered over the dorsal surface of the cerci. Females of the Oriental

vexans group of species have a few narrow scales ventrally on the prosternum.

The 2 subspecies of vexans also have 1-3 bristles on the prosternum and have

the insula of the genitalia without tuberculi. These features of the prosternum

and insula differ from the other species of Oriental Aedimorphus.

The features of the pupal stages of vexans vexans and vexans nipponii

tend to overlap and therefore cannot be separated with certainty.

The larva of vexans vexans in Southeast Asia has the frontoclypeus of

the head granulose while this area in vexans nipponii is nongranulose.

In North America, Crans and Gander (1968: 235) reported finding a

few female specimens of vexans vexans from Delaware, Ohio and New Jersey

which displayed abdominal and pleural markings similar to those of the sub-

species nipponii. During the 1971 Venezuelan equine encephalitis outbreak in

the southern United States, there were 13, 600 female specimens of vexans

vexans examined from Arkansas, Louisiana, Oklahoma and Texas by U. S.

Army taxonomists and only 26 of these were of the abnormal type resembling

nipponii. I have also examined 8 females from Worcester County, Maryland,

1 female from Valdosta, Georgia and 1 female from Portland, Oregon of

this form. No males or females with associated immatures of the abnormal

type have been collected. The females differ from nipponii in lacking scat-

tered white scales on the dorsal wing veins and a reduction in the number of

pale scales on the maxillary palpi and legs. A possible explanation for this

form being encountered so infrequently is that in North America it is a

Reinert: Aedes (Aedimorphus) in Southeast Asia 717

recessive genotype of vexans vexans that has a very low frequency.

Belkin (1962: 427) gave full species status to the members of vexans

occurring on the Pacific Islands and based this determination on the branching

of the larval head hairs 5, 6-C and minor differences in the adult habitus.

After examining specimens of vexans vexans from a number of areas in its

range (see DISTRIBUTION) I found that the species is extremely variable in

the size and habitus of the adults and the chaetotaxy of the immatures. Barr

(1954: 24) also records the variability of the larval head hairs 5, 6-C of vex-

ans vexans. Specimens of vexans vexans many times exhibit a noticeable vari-

ation within local populations as well as variances between geographical popu-

lations. In one collection of larvae (with reared adults) from Mindoro, Philip-

pines both head hairs 5 and 6-C ranged from both single to 5-C double and 6-C

triple with all combinations between these extremes. In collections from New

Guinea, South Vietnam, Hong Kong and Thailand, I found the same variability

and in some cases even greater variations. The 2 female syntypes of nocturnus

in the British Museum (Natural History) and other specimens from the Pacific

Islands that I have examined seem to fall within the normal variation of char-

acters of adult habitus, female and male genitalia, and pupal and larval chae-

totaxy of vexans vexans from the remainder of its distribution. In the absence

of sufficient biological, behavioral and genetical data on the Pacific Island

populations of vexans, and since specimens from these populations fall within

the variable range of morphological characters of other populations within the

distribution of the species, I am hereby synonymizing nocturnus with vexans

vexans .

BIOLOGY. The immatures have been collected from a variety of habitats

but appear to prefer unshaded, fresh water flood pools in secondary scrub, open

plains and rice fields. Adults have been taken feeding on man and domestic ani-

mals. Immatures in Thailand have usually been collected from large and small

flood pools but also from a large rice field, small and large ditches, small

swamp, abandoned posthole, and elephant footprint; usually from clear fresh

water but occasionally from colored turbid fresh water; water usually with

scarce floating and emergent aquatic vegetation or scarce green algae; in partial

shade or unshaded areas; in rice fields and coconut groves in plains, banana

grove in valley, banana grove, secondary scrub, primary bamboo grove and

primary rain forest in mountains; and at an altitude from 7 to 1, 640 feet (most

often at 50 to 600 feet). Larvae were collected in association with the following

species of mosquitoes: Aedes alboscutellatus, caecus, lineatopennis, medio-

lineatus; Anopheles kochi, subpictus, vagus; Culex fuscanus, gelidus, mimulus,

sitiens, tritaeniorhynchus; and Uranotaenia bicolor. In New Guinea, Peters and

Christian (1963: 48, 64) found larvae associated with the following species:

Anopheles farauti, Culex fraudatrix group, mimulus and pullus in the lowlands

and Anopheles annulipes, favauti, Culex mivaculosus and quinquefasciatus in

the highlands.

Immatures were collected from: ground pools, swamp, ditch, foxhole,

rice field, marshy depression and seepage spring in South Vietnam; temporary

puddle, flood pool, grassy pool, fish pond, sunlit ground pool, small flooded

area near road, ditch, rain pools in pasture, carabao wallow and an open muddy

algae-filled pond in the Philippines; hoofprints in Fiji; ground pool in Taiwan;

ground pool in a fallow rice field with abundant grass and scarce green algae

and associated with Anopheles tessellatus in Hong Kong; ground pools in India;

stagnant water in a roadside ditch in Burma; and in sunlit ground pool, artificial

container, brackish water pool and pools in alang-alang in Indonesia. Adults

were taken biting man, cattle and horses, in light traps, resting in houses and

78 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

bamboo forests in Thailand; biting man and in light traps in the Philippines and

South Vietnam; in Malaise traps at an elevation of 10 feet in Ceylon; and in

houses in Indonesia.

Immatures have been recorded in the literature from the following

habitats: cisterns, rain barrels, pottery, coconut husks, carabao wallows,

standing water, rice fields, pools with lilies (Dowell, Libay and Baisas 1965:

30), temporary rain puddles, grassy swamps, drainage ditches (Perry 1946:

16, Knight and Hull 1953: 462) and from a tin can (Basio et al. 1970: 443) in

the Philippines; unshaded pools and ditches, in salt, brackish or fresh water

(Assem and Bonne-Wepster 1964: 101), artificial containers, hoofprints, pools,

ponds and ditches (Steffan 1966: 212) in New Guinea; clear water pools contain-

ing grass in Guam (Yamaguti and La Casse 1950: 78); in ditches and ponds in

Malaysia (Macdonald 1957: 21); in ground pools and artificial receptacles in

Thailand (Causey 1937: 414); newly flooded rice fields and ground pools in Tai-

wan (Lien 1962: 628); seepage ditch water, marshes, temporary grassy pools,

ponds, ditches and ground depressions in South Vietnam (Grothaus et al. 1971:

21); and ground pools in China (Chow 1949: 129). Adults have been taken in in-

door shelters in India (Rao and Rajagopalan 1957: 10); feeding on cattle (Mac-

donald 1957: 21), and in an indoor evening catch (Macdonald 1956: 233) in Ma-

laysia; readily biting man in the daytime and collected at elevations from 33 to

1, 300 feet (Basio et al. 1970: 438, 443), commonly taken while trying to bite

humans at night and from a light trap (Knight and Hull 1953: 462) in the Philip-

pines; females biting man, cows and buffaloes at night in Taiwan (Lein 1962:

628); in light traps, CO 9 traps, indoor resting sites and biting man in South

Vietnam (Grothaus et al. 1971: 21); and in light traps and biting man indoors

at night in New Guinea (Peters and Christian 1963: 44, 55). In Thailand vexans

vexans were collected that had fed on bovines and chickens (Unpublished data

from SEATO Medical Research Laboratory Annual Progress Report 1967: 450)

and in scrub or open forests, in heavy forest and at elevations from 1, 000 to

4,500 feet (Scanlon and Esah 1965: 139, 143). Templis et al. (1970: 339) sum-

marized the feeding preferences of vexans vexans in the Hawaiian Islands as

follows: man--2.4 percent, bovine--50. 7 percent; horse--37.2 percent; dog--

4.7 percent, pig--1.6 percent and unknown host--3.4 percent. In Malaysia,

Standfast (1967: 193) found that vexans vexans readily fed on man between 1800

and 0600 hours with a peak in feeding around 2400 hours and a smaller one

around 0500 hours. Laird (1956) records vexans collections from a number of

localities in the South Pacific, notably from transient pools (Ellice Island,

New Hebrides, Western Samoa); ponded ditches (Tonga, New Caledonia); pond

in forest clearing (Belep Island); road ruts (Fiji, New Hebrides); taro ponds

(Ellice Islands, Gilbert Islands); seepage pond (Tonga); and a small seepage

pond with decomposing grass clippings (Western Samoa).

Eggs of vexans vexans are deposited in shallow areas that are subject

to inundation. Horsfall (1956: 66) found that embryonated eggs of vexans vex-

ans are able to withstand adversities such as drought, cold and premature sub-

mersion often for long periods of time. He also stated that eggs subjected to

adversities required ''conditioning" prior to the basic hatching stimulus. The

hatching stimulus is a decrease in the content of dissolved oxygen in the water.

Gjullin, Yates and Stage (1950: 268) reported that eggs in field cages showed

little or no mortality over 2 years, but for longer periods viability rapidly

decreased and less than 1 percent survived 4 years. Eggs of vexans vexans

deposited 1 year may hatch throughout the following season but usually most

eggs that survive the winter hatch the following spring (Miller 1930 cited in

Horsfall 1955: 523). Horsfall (1955: 523) stated that eggs in natural sites can

Reinert: Aedes (Aedimorphus) in Southeast Asia io

withstand prolonged freezing without detrimental effects.

AEDES (AEDIMORPHUS) VEXANS NIPPONII (THEOBALD)

(Figs. 15, 16, 17, 32, 48, 58, 69)

Culicada nipponii Theobald 1907, Monogr. Cul. 4: 337 (9*).

Ochlerotatus vexans variety nipponii Theo., Edwards 1917, Bull. ent. Res.

7; 219.

Ochlerotatus vexans Mg., Brunetti 1920 (in part), Rec. Indian Mus. 17: 135;

Senior-White 1923 (in part), Cat. Indian Insects, Cul., p. 83.

Aédes (Ecculex) vexans variety nipponii Theobald, Edwards 1921, Bull. ent.

Res. 12: 322.

Aédes vexans var. nipponii Theobald, Ho 1931, Bull. Fan meml. Inst. Biol.

2-131. (ot, Oy,

Aédes (Aédimorphus) vexans var. nipponit Theobald, Edwards 1932, Genera

Insec., Fasc. 194: 171.

Aédes (Aédimorphus) vexans (Meigen), Barraud 1934 (in part), Fauna Brit.

India, Diptera 5: 253.

Aedes (Aedimorphus) vexans Meigen, Feng 1938, Peking nat. Hist. Bull.

12: 291; Hsiao 1945, Navmed. 630, p. C6.

Aedes (Aedimorphus) vexans nipponii (Theobald), Bohart 1946, U. S. Navmed.

961, p. 10; Bohart and Ingram 1946, U. S. Navmed. 1055, p. 69

(o*, L*); Hsiao and Bohart 1946, U. S. Navmed. 1095, p. 22; La Casse

and Yamaguti 1947, Mosquitoes Japan, 2: 39(L*); La Casse and

Yamaguti 1948, Mosquito Fauna Japan and Korea, p. 100 (o*, 9*, P*,

L*); La Casse and Yamaguti 1950, Mosquito Fauna Japan and Korea,

p. 125 (oc*, 9*, P*, L*); Yoshimeki 1955, Ecol. Rev., Jap. 14: 81 (L*);

Hara 1957, Jap. J. exp. Med. 27: 66 (9*); Stone et al. 1959, Thomas

Say Found. 6: 199; Kurihara 1963, Jap. J. sanit. Zool. 14: 196 (A*).

FEMALE (Fig. 15). Head. Antenna dark brown, approximately 0. 88

length of proboscis, pedicel pale with a number of small white scales and a

few short fine brown hairs mesally, flagellomere 1 pale with a few dusky

scales; clypeus dark, bare; maxillary palpus brown scaled with white scales

at base of segment 4 and at base and apex of segment 5, usually several white

scales scattered over segments 3-5, approximately 0.20 length of proboscis;

proboscis brown scaled with a broad longitudinal ventral stripe from near base

to apical 0.25, stripe extends up lateral surfaces and onto dorsolateral areas,

in some specimens the stripe forms a nearly complete band, apical 0.25 always

dark brown, approximately 1.26 length of femur I; vertex with dorsum covered

with narrow decumbent scales arranged in a small anteromedian diamond-shaped

golden-brown group and the remainder white, narrower white scales on posterior

margin of occiput; lateral surface covered with broad scales, an anterodorsal

black patch with white ones below; numerous long golden-brown erect forked

scales on occiput and vertex extending anteriorly to ocular line. Thorax. Scu-

tal integument dark brown; scutum covered with narrow curved reddish-brown

scales (specimens from Japan have scales golden-brown), narrow curved white

scales forming patches on anterior promontory area, scutal fossal area extend-

ing from anterior area along margin and onto lateral area, supra-alar area

from above middle of paratergite to posterior of wing base, posterior median

scutal area, along lateral margins of prescutellar space and among posterior

dorsocentral setae; scales nearly cover prescutellar space in some specimens;

80 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

scutellum with a patch of narrow curved white scales on each lobe; median

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

scutal fossal (anterior, 4-5 lateral, 1-3 median and 1-6 posterior), supra-alar,

several posterior medial scutal, 1 postalar callar and scutellar (lateral and

median) bristles reddish-brown and well developed; pleural integument brown;

antepronotum covered with narrow curved white scales, several golden and

brown bristles; postpronotum covered with narrow curved reddish-brown

scales, a small posteroventral patch of narrow curved and usually a few broad

white scales, 8-9 posterior dark bristles; propleuron with a patch of broad

white scales, several golden bristles; prosternum with narrow white scales

and 1-3 bristles; postspiracular area with a patch of moderately broad white

scales, 7-11 golden bristles; subspiracular area with 2 patches of broad white

scales, lower one large and frequently connected with postspiracular patch;

mesepisternum with a large upper and a posterior patch of broad white scales,

patches usually connected or nearly so, several upper and posterior golden

bristles, lower ones shorter; prealar knob with a few broad white scales on

lateroventral margin, several golden bristles; paratergite covered with white

scales, scales vary in shape, some specimens with broad ones, some with

narrow curved ones and some with both; mesepimeron with a large patch of

broad white scales and several golden bristles on upper area; other pleural

areas bare. Legs (Fig. 16). Coxae I-III each with several brown to golden

bristles, I, I each with anterior surface white scaled, I with a patch of brown

scales at middle, III with an anterodorsal and an anteroventral patch of white

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

each with anterior surface brown and with a dorsoapical white spot, I-III each

with a narrow basal white band and numerous scattered white scales, I with

30-60 percent of scales white, III with a broad longitudinal white stripe taper-

ing from base to apical 0.25, remainder with scattered white scales; femora

I-III each with posterior surfaces white, I with scattered brown scales on

posteroventral area, II, III each with scattered brown scales on apical 0. 25;

tibiae I-III each with a few scattered white scales and a narrow basal white

band, I with ventral and posterior areas white, II with dorsal and posterior

areas white, a small brown area on dorsal surface at base and at apex, III

with a posterior longitudinal pale stripe; tarsi I-III brown, I with tarsomeres

2,3 each with a narrow basal white band, tarsomere 1 and basal 0.50 of tarso-

mere 2 with a posterior longitudinal pale stripe, II with tarsomeres 1-4 each

with a narrow basal white band, tarsomeres 1-2 each with a posterior longidu-

dinal pale stripe, tarsomere 5 with a few laterobasal white scales, III with

tarsomeres 1-5 each with a narrow basal white band, tarsomere 1 and basal

0.50 of tarsomere 2 with a posterior longitudinal pale stripe; posttarsi I-III

each with 2 ungues, I, II equal, each bearing a tooth, III equal, simple, occa-

sionally each with a tooth. Wing. Dorsal veins covered with moderately broad

brown scales; costa with a patch of white scales at base and along basal 0. 50

of its posterior margin; subcosta, radius, cubitus and occasionally anal veins

with a few white scales intermixed on basal 0. 50; ventral veins brown with

some white scales on posterior margin of costa and on subcosta at humeral

cross vein; alula with narrow brown scales along fringe; 2-3 remigial bristles.

Halter. Pedicel pale, capitellum white scaled. Abdomen. Terga brown with

somewhat variable white markings; tergum I with dorsum white with a few inter-

mixed lateral brown scales, laterotergite with a rectangular patch of white

scales; tergum II with a large basomedian triangular white patch; terga ITI-VI

each with a dorsobasal white band, bands very broad laterally and becoming

narrower mesally, II[-V each with an incomplete median apical longitudinal

Reinert: Aedes (Aedimorphus) in Southeast Asia 81

white stripe which may or may not connect with basal bands; terga VI, VII

varies from each with a large dorsoapical triangular white spot and VII with a

pair of small basolateral triangular white patches on dorsum to VI, VII each

white with a dorsal brown V-marking pointing anteriorly; terga II-VII each

with a large laterobasal white patch which may nearly cover lateral surfaces

but is not connected with dorsobasal bands; sterna white scaled with brown

scales forming a narrow indistinct longitudinal stripe on IJI-VII, terga and

sterna with numerous golden bristles, mostly along posterior margins.

Genitalia (Fig. 32). Tergum VIII index 0.92-0.99; sternum VIII index 0. 88-

0.97; tergum IX bilobed with 4-8 bristles on each lobe, index 0. 85-1. 02; insula

tongue-like, covered with minute setae; lower vaginal lip narrow, moderately

pigmented, covered with minute setae; upper vaginal lip narrow, heavily pig-

mented, covered with minute setae; upper vaginal sclerite large, moderately

pigmented, triangular shaped; postgenital lobe moderately long, apex with a

moderately deep median indentation, 5-8 bristles on each side of midline,

covered with minute setae, dorsal PGL index 0. 75-0.91, ventral PGL index

1.55-1. 87; cercus moderately long to long, 0.90-1.00 extended from segment

VII, narrow, apex sharply rounded, numerous bristles on dorsal and lateral

surfaces, index 3.08-3.59, cercus/dorsal PGL index 4.34-5. 20; 3 spherical,

pigmented seminal capsules, 1 large and 2 slightly smaller ones.

MALE (Fig. 15). Similar to female in general habitus. Head. Maxil-

lary palpus brown with segments 2, 4, 5 each with a dorsobasal white spot,

segment 3 with a basal white band, longer than proboscis by length of apical

segment; vertex with anteromedian narrow golden-brown scale patch and an-

terodorsal broad black scale patch reduced. Thorax. Prosternum bare.

Legs (Fig. 17). Tarsi I-III with posterior longitudinal stripes reduced; post-

tarsi I-III each with 2 ungues, I, II with ungues unequal, each bearing a tooth,

Ill equal, simple. Abdomen. Terga II-VI with basal white bands which are

expanded posteriorly on lateral surfaces, VII with large lateral triangular

patches nearly connected medially, III-V with median apical longitudinal white

stripes reduced to a few pale scales in some specimens; tergum VIII white;

sternum VIII white with a median longitudinal brown band. Genitalia (Fig. 48).

Nearly identical to those of Aedes vexans vexans.

PUPA (Fig. 58). Chaetotaxy as recorded in Table 10. Two to three

rows of spicules mesally along posteroventral margin of abdomenII. Cephalo-

thovax. Hair 5-C with 2-4 branches; 7-C with 2-4 branches; 8-C with 3-6

branches. Respivatory trumpet. Moderately pigmented; index 2. 89-3. 82,

average 3.26. Metanotum. Hair 10-C with 4-10 branches; 12-C single to

triple. Abdomen. Hair 5-I with 2-5 branches; 10-I single; 1-II with 4-10

branches; 4-II with 2-4 branches; 1-II. with 3-6 branches; 6-VI single or

double; 1-VII single to triple; 6-VII with 3-5 branches; 9-VII with 2-4 branches;

11-VII single to double. Paddle. Ovoid; with very minute serrations along

basal 0.55 of outer margin; tiny spicules along apical 0.45 of outer and apical

0.30 of inner margins; midrib does not reach apex; hair 1-P short, single or

double; index 1.42-1. 82, average 1. 59.

LARVA (Fig. 69). Chaetotaxy as follows: Head. Frontoclypeus without

granules; median mouth brushes pectinate apically; hairs 1, 3, 8-C single;

4-C with 2-4(4) branches; 5-C single to 4(3) branched; 6, 14-C single or double

(1); 7-C with 7-12(9) branches; 9, 10-C single or double (2); 11-C with 4-10(7)

branches; 12-C with 4-7(5) branches; 13-C single to triple (2); 15-C with 4-6

(4) branches; basal maxillary hair single; mental plate with 25-27(26) teeth.

Antenna. Short and moderately to heavily pigmented; with small stout spicules

scattered over shaft, most numerous on basal 0.50; hair 1-A with 5-10(6)

82 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

branches, inserted 0.38-0.45 from base; 2-A long; 3-A approximately 0.50

length of 2-A. Thorax. Hair 0-P with 3-6(5) branches; 1, 2, 4-6, 9, 10,

12-P single; 3-P double or triple (2); 7-P double or triple (3); 8-P with 2-4(3)

branches; 11-P single to triple (2); 14-P single or double (1); 1, 2, 4-M with

2-4(3) branches; 3, 11-M single or double (1); 5, 7, 10, 12-M single; 6-M

with 5-6(5) branches; 8-M with 5-7(6) branches; 9-M with 6-7(6) branches;

13-M with 5-9(7) branches; 14-M with 4-6(5) branches; 1-T single to triple

(2); 2-T double or triple (3); 3-T with 2-7(6) branches; 4-T with 2-5(4)

branches; 5, 6, 10, 12-T single; 7-T with 6-11(7) branches; 8, 9-T with

4-6(5) branches; 11-T single or double (1); 13-T with 5-8(5) branches. Abdo-

men. Hairs 1, 2-VIII on common basal plate; hairs 0, 14-VII single; 1-VIII

with 4-12(8) branches; 2, 4-VIII double or triple (2); 3-VIII with 5-12(8) bran-

ches; 5-VIII with 6-9(9) branches; 6-IIL-VI long; comb of 9-11(9) scales

arranged in 1-2 irregular rows, scales with a long stout pointed median spine

and short denticles along lateral margins of base; 1-X single or double (1);

2-X with 6-9(8) branches; 3-X single; ventral brush with 11-13 (usually 11)

hairs on grid and 3 precratal ones; saddle moderately pigmented with minute |

ridges, incompletely rings segment, with small spicules along posterior mar-

gin, acus present; 4 anal papillae, long and tapering to a blunt point. Szphon.

Moderately pigmented with minute ridges over entire surface; acus large; index

2.42-3. 86; pecten with 9-11(9) teeth, apical 2-3 teeth longer, smooth or with a

minute lateral denticle and wider spaced than remainder which have a slender

apical attenuated filament with 1-3 basal denticles; hair 1-S with 5-7(5)

branches, inserted 0.57-0. 64 from base.

TYPE DATA. Culicada nipponii Theobald, holotypefemale, Karnizana,

JAPAN, 25 August 1903, Mr. Cornford, in British Museum (Natural History).

DISTRIBUTION. Specimens examined--51 males, 115 females, 30

pupae, 121 larvae and 33 individual rearings (5 pupal, 28 larval) from the

following locations:

CHINA. Kiangsu, Peking, Shanghai, Yenching; Manchuria, Mukden,

Tetsusci.

JAPAN. Karnizana; Saitama, Honshu, Sagiyama; Kanagawa, Inada;

Tokyo.

RYUKYU ISLANDS. IshigakiIs., Inota III; Okinawa, Dyama, Ganiko,

Kuba, Myiazato, Naha, Oyama; Miyako Is.

SOUTH KOREA. Koje-do, Seoul, Taegu.

Other distribution.

CHINA. Honan, Kai Feng, Chi Kung Shan, Hsin Yang (Su and Ch'ii

1956: 219); Hopei (Ma and Feng 1956: 172); Yunnan, Chefang (Chow 1949: 129);

Chekiang, Hangchow, Hwangyen, Iwui Pingyang, Tienmushan; Kiangsu,

Shanghai, Nanking; Kwangtung, Canton, Swatow; Liaoning, Hailung; Tzbet,

Chumbi Valley; Yunnan, Gadzu Beta (Feng 1938: 291); (Gutsevitch et al. 1970:

274).

JAPAN. Hokkaido, Sapporo; Honshu, Yodo; Sendai (La Casse and

Yamaguti 1950: 129); Asahikawa, Kagoshima (Hara 1957: 66); Tokyo, Katsu-

shika Ward (Sasa et al. 1950: 511); Hokkaido (Sasa et al. 1947: 53); Okayama

(Sasa and Sabin 1950: 24); Nagasaki City (Ito 1964: 232); Tomachi, Mogi,

Kaizu, Koebaru (Shichijo et al. 1970: 94); Karnizana (Theobald 1910b: 301);

Sagiyama, Shinhama (Scherer et al. 1962: 261); Hokkaido, Sendai, Tokyo

(Hsiao and Bohart 1946: 22); Nagasaki (Omori et al. 1952: 281); Fukui, Tadasu-

Mura (Omori 1951: 310); Okayama, Tokoku (Sasa 1949: 99); Tochigi, Shiobara,

Tomatsuri, Utsunomiya (Kurashige 1964: 75, 84, 94); Funaoka (Kato and Tori-

umi 1950: 468); Itazuke AB, Misawa AB, Tachikawa AB, Yokota AB (Reisen

Reinert: Aedes (Aedimorphus) in Southeast Asia 83

et al. 1971: Tables 23 and 24).

SOUTH KOREA. Kimpo AB, Kunsan AB, Kwang-Ju AB (Reisen et al.

1971: Tables 26 and 27); Seoul City, Hyangdangdong (Chu 1957: 5); Cheju

Island, Daejong Up, Mosulpo (Oh et al. 1961: 85).

U. S. S. R. Kharbarovsk Area, Baikal Area (Gutsevitch et al. 19770:

274).

TAXONOMIC DISCUSSION. Aedes vexans nipponiiis similar in the

adult habitus, female and male genitalia, pupa and larva to vexans vexans

and is discussed under that species.

BIOLOGY. Immatures have been found in a wide range of habitats but

seem to prefer unshaded fresh water, flood pools and rice fields. Adults have

been taken feeding on man both during the day and night. Larvae were col-

lected on Okinawa from ground pools, rice field, stream, swamp, ditch anda

small pond.

Immatures have been collected from natural water pools and swamps

in China (Feng 1938: 291), from Japanese decorative garden pools, rice field

ditches, stone urn, bamboo stump, small cup and seepage pool from broken

pipe in Japan (La Casse and Yamaguti 1947: 40) and from temporary, partially

shaded, fresh water ground pools containing leaves in Okinawa (Bohart and

Ingram 1946: 69). In Japan larvae of vexans nipponii were collected from

rice fields in association with Culex pipiens, tritaeniorhynchus, orientalis,

hayashti, bitaeniorhynchus and Aedes dorsalis (Kato 1956: 157). The adult

feeding period in Japan of this species extended from 1600 to 0700 hours with

a peak feeding time from 1730 to 2200 hours (Kato and Toriumi 1950: 468).

Sasa (1949: 101), while working in Japan, collected adults very early in the

spring and believes that this species hibernates as adults during the winter. Adults

were taken both during the day and night biting man in Japan (La Casse and

Yamaguti 1948: 104, 1950: 129).

AEDES (AEDIMORPHUS ?) WAINWRIGHTI BAISAS

Aedes (Aedimorphus) wainwrighti Baisas 1946, Mon. Bull. Bur. Hlth. Philipp.,

Manila 22: 34 (2); Knight and Hull 1953, Pacif. Sci. 7: 464 (9); Stone

et al. 1959, Thomas Say Found. 6: 199.

FEMALE. Head. Antenna dark brown, pedicel dark with a small patch

of broad silvery scales mesally; clypeus dark brown, bare; maxillary palpus

dark brown scaled, approximately 0.17 length of proboscis; proboscis dark

brown scaled, slightly shorter than femur I; vertex and lateral surface covered

with broad decumbent scales, dorsum with an anteromedian triangular patch of

silvery scales that broadens anteriorly and extends as a fine line along eye mar-

gin, a dark scaled spot laterad of median spot, remainder of vertex and lateral

surface pale brownish (almost whitish under different lighting); narrow dark

scales and a few erect scales on occiput. Thovax. Scutal integument brownish;

scutum covered with narrow dark brown scales with a broad median silvery

line on anterior 0.50; scutellum with broad silvery scales on median lobe; a

pair of dorsocentral bristles toward anterior border and a stronger pair above

level of wing bases, 2 pairs of prescutellars (posterior dorsocentral bristles),

rather few supra-alars and scutellar (lateral and median) bristles; pleural integu-

ment darker brown than scutum; antepronotum with a small patch of silvery flat

scales, about 7 stiff, tawny bristles; postpronotum with 3 tawny bristles, but

84 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

without scales; propleuron with a round patch of silvery scales, 4-5 tawny bris-

tles; postspiracular area with 6 tawny bristles; subspiracular area with a nar-

row elongate patch of silvery scales astride suture between this area and

mesepisternum; mesepisternum with a fairly large round upper patch and a

lower one of silvery scales, 1 upper and 3-4 lower (posterior) tawny bristles;

prealar knob with 4-5 tawny bristles; paratergite covered with a dense patch

of broad silvery scales; mesepimeron with a patch of silvery scales and sev-

eral weak bristles on upper area. Legs. Coxae I-III each with an elongate

patch of silvery scales; femora I-III each dark brown with ventral surface and

base pale, II with an apical silvery spot, III with basal 0.50 of anterior and

posterior surfaces pale and a large apical silvery spot; tibiae I-III each dark

brown; tarsi I-III each dark brown with a white basal band on tarsomere 1,

band indistinct on I, marked on II and apparently still wider on III as indicated

by the remaining portion of the tarsomere. Wing. Dark scaled. Abdomen.

Terga dark scaled; tergum I with laterotergite silvery scaled; terga II-VII each

with a laterobasal patch of silvery scales (patches broadened, basally), patches

visible dorsally on V-VII; sterna IJI-VII pale with posterior portions dark

scaled. Genitalia. Tergum VIII completely retracted into segment VII.

MALE, PUPA and LARVA. Not known.

TYPE DATA. Aedes (Aedimorphus) wainwrighti, Baisas, holotype fe-

male, Llavac, Infanta, Tayabas, Luzon, PHILIPPINES, 29 May 1940, Mr.

Pablo Sunico, collected in a mosquito trap; holotype nonexistent (Stone 1970:

151).

DISTRIBUTION. Known only from type locality.

TAXONOMIC DISCUSSION. The above description has been adapted

from the original one by Baisas (1946).

Aedes wainwrighti possesses a number of features that differ from

other species of Aedimorphus in the Oriental Region. The most marked dif-

ferences are: pedicel with a patch of broad silvery scales mesally; erect

scales of head restricted to occiput; vertex of head with dorsum covered with

broad decumbent scales (punctifemoris is the only other Oriental species of

Aedimorphus with this character); reduction in number or absence of anterior

dorsocentral bristles; absence of acrostichal bristles; presence of a broad median

line of silvery scales on anterior 0. 50 of scutum; scutellum with broad silvery

scales only on median lobe; and pleural integument dark brown. These fea-

tures are more suggestive of Stegomyia than of Aedimorphus.

Even with these differences there are characters shared by other Orient-

al Aedimorphus and since the type has been lost, the male has never been

found and no new material has been collected, I am for the present doubtfully

retaining wainwrighti in this subgenus.

BIOLOGY. The holotype was collected in a mosquito trap.

MISIDENTIFICATIONS and DOUBTFUL RECORDS

AEDES (AEDIMORPHUS) STENOETRUS (THEOBALD)

(Figs. 12, 16, 17, 29, 45)

Culex stenoetrus Theobald 1907, Monogr. Cul. 4: 395 (9).

Culex pseudostenoetrus Theobald 1910, Monogr. Cul. 5: 343 (9*).

The original record of this species from Southeast Asia was that of

Thurman and Thurman (1955: 222) from Chiengmai, Thailand. Two females

Reinert: Aedes (Aedimorphus) in Southeast Asia 85

determined by the Thurmans as this species from Chiengmai were found in the

United States National Museum. They are actually Aedes vexans vexans.

Additional material in the U. S. National Museum labeled as stenoetrus from

Assam, India, were a mixture of vexans vexans and caecus. No specimens of

stenoetrus from Southeast Asia were found in the collections of the U. S.

National Museum (Natural History) and the British Museum (Natural History).

A discussion of distinguishing characters of stenoetrus is presented

in the taxonomic discussion section of Aedes vexans vexans and the adults are

illustrated (Figs. 12, 16, 17, 29, 45) in order to help field workers recognize

this species.

AEDES (AEDIMORPHUS) TAENIORHYNCHOIDES (CHRISTOPHERS)

(Figs. 13, 16, 17, 30, 46)

Pecomyia maculata Theobald 1905 (non Meigen 1804), J. econ. Biol. 1: 23

(2*) (name preoccupied).

Leslieomyia taeniorhynchoides Christophers 1911, Paludism 2-3: 68 (c‘, 9).

The following published records of Aedes taeniorhynchoides from South-

east Asia are misidentifications or are doubtful records:

CHINA (Feng 1958: 74); Hainan Island (Chu 1957: 158, 1958: 109).

THAILAND (Thurman 1959: 121, 1963: 54); Chiengmai, Chiengmai

(Scanlon and Esah 1965: 139); Korat, Nakhon Phanom, Takhli, Udorn (Reisen

et al. 1971: Tables 4, 5 and 7).

SOUTH VIETNAM. Phan Rang (Parrish 1968a: 3); Nha Trang (Parrish

1969: 554); Cam Rahn Bay, Phu Cat, Tan Son Nhut (Reisen et al. 1971: Tables

11, 12 and 13); Chu Lai, Da Nang, Phu Bai (Grothaus et al. 1971: 20).

The original record of this species from Thailand was that of Thurman

(1959) and although the locality data were not included the material upon which

the report was based was probably collected in northern Thailand. Nine fe-

males determined as Aedes taeniorhynchoides were found in the Thurman col-

lection in the United States National Museum (Natural History) and contained

the following data: Doi Chom Chen, Chiengmai, Chiengmai Province, Thailand,

IV-7-1952, Det. D. & E. Thurman. All nine of these specimens are Aedes

vexans vexans. I have also seen a number of specimens from Thailand deter-

by Scanlon and co-workers as taeniorhynchoides but they are Aedes vigilax.

Five females labeled taeniorhynchoides from South Vietnam were also vigilax.

Since I have not found any taeniorhynchoides in the large numbers of specimens

from the Malayan, Thailand and South Vietnam collections and since the above

mentioned specimens previously identified as taeniorhynchoides were incorrect

determinations, I am doubtful if this species occurs in Southeast Asia. It

probably has a more restricted range in Ceylon, India and Pakistan. Other

records for this species from Southeast Asia may possibly be incorrect deter-

minations for Aedes pipersalatus, vexans vexans or vigilax.

The confusion in the identification of taeniorhynchoides in the past is

probably a result of an inadequate original description without illustrations.

Even though it is doubtful that this species occurs in Southeast Asia it is in-

cluded in this paper (adult key and Figs. 13, 16, 17, 30, 46) in order to clarify

identification of closely related species.

The adult habitus and female and male genitalia of taeniorhynchoides

are similar to pipersalatus and are discussed under that species. Females of

taentorhynchoides also resemble vexans vexans and Aedes (Ochlerotatus)

86 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

vigilax (Skuse) from which they can be separated by the large number of white

scales intermixed with brown ones on most of the dorsal veins of the wing,

especially on the radius and among the tertiary fringe scales along the posterior

margin of the wing while vexans vexans has the distal 0. 85 of the radius 2+3

brown scaled. Aedes vexans vexans and vigilax have all the scales along the

posterior margin of the wing brown. Aedes vigilax also has the subspiracular

area bare while vexans vexans and taeniorhynchoides both have scales on this

area. Aedes taeniorhynchoides has the postpronotum completely covered with

narrow curved pale scales while vexans vexans has the upper portion of this

area with reddish-brown scales and white ones below. The aedeagus, proc-

tiger and gonostylus of the male genitalia of taeniorhynchoides are markedly

different from those of vexans vexans (Figs. 33 and 47) and vigilax (aedeagus

is a simple tube).

ACKNOWLEDGEMENTS

Special thanks are expressed to Dr. Botha de Meillon, Responsible

Investigator, Southeast Asia Mosquito Project (SEAMP), for obtaining Aedi-

morphus specimens for me from many areas in the Oriental and African re-

gions and for critically reading the manuscript. Appreciation is expressed to

Lieutenant Colonel Bruce F. Eldridge, Walter Reed Army Institute of Research,

Washington, D. C., for reviewing the manuscript and making helpful sugges-

tions. Sincere thanks are given to Dr. Alan Stone, Systematic Entomology

Laboratory, Agriculture Research Service (ARS), U. S. Department of Agri-

culture (USDA) and Dr. P. F. Mattingly, British Museum (Natural History),

for kindly permitting me to examine types and other specimens. Gratitude is

expressed to the members of my doctoral committee: Dr. F. S. Blanton

(Chairman), Dr. D. H. Habeck, Dr. S. G. Zam, University of Florida,

Gainesville, Florida, Dr. F. W. Mead, Division of Plant Industry, Entomology

Section, Florida Department of Agriculture, Gainesville, Florida, Dr. C. S.

Lofgren, Entomology Research Division, ARS, USDA, Gainesville, Florida,

and Dr. Alan Stone, Systematic Entomology Laboratory, ARS, USDA, Washing-

ton, D. C., for guidance during the preparation of my dissertation at the

University of Florida. I wish to thank the Medical Department, U. S. Army for

financing a trip to study type specimens in the British Museum (Natural History).

I am particularly indebted to the personnel of SEATO Medical Research Labora-

tory, Thailand, for collection and preparation of many valuable specimens, to

Dr. J. E. Scanlon for the loan of notes made on specimens located in the Indian

Museum, and to Lieutenant Colonel Alexander A. Hubert, Chief, Department of

Entomology, U. S. Army 406th Medical Laboratory, Japan, and his artist staff

who prepared most of the illustrations. Thanks are given to Mr. Young Sohn

and Mr. Vichai Malikul, SEAMP, who prepared several of the illustrations.

Acknowledgement and thanks are given to the following individuals and institu-

tions for the loan of specimens, obtained for me by SEAMP: Mr. G. L.

Alcasid, National Museum, Manila, Philippines; Dr. P. H. Arnaud, California

Academy of Sciences, San Francisco, California; Dr. F. E. Baisas, Manila,

Philippines; Dr. D. S. Bertram, London School of Hygiene & Tropical Medicine,

London, England; Dr. F. S. Blanton, University of Florida, Gainesville, Flori-

da; Dr. J. Bonne-Wepster, Instituut voor Tropische Hygiene en Geographische

Pathologie, Amsterdam, Netherlands; Dr. D. F. Bray, University of Delaware,

Newark, Delaware; Commanding Officer, U. S. Army Medical Research Unit,

Institute of Medical Research, Kuala Lumpur, Malaysia; Commanding Officer,

Reinert: Aedes (Aedimorphus) in Southeast Asia 87

U. S. Army Medical Research Team, Walter Reed Army Institute of Research,

South Vietnam; Commanding Officer, U. S. Army 9th Medical Laboratory,

South Vietnam; Commanding Officer, U. S. Army 20th Preventive Medicine

Unit, South Vietnam; Commanding Officer, U. S. Naval Medical Research Unit

No. 2, Taiwan; Director, American Museum of Natural History, New York,

New York; Drs. D. J. Gould and J. E. Scanlon, SEATO Medical Research

Laboratory, Bangkok, Thailand; Dr. J. M. Klein, Institut Pasteur, Phnom

Penh, Cambodia; Dr. K. L. Knight, North Carolina State University, Raleigh,

North Carolina; Dr. P. F. Mattingly, British Museum (Natural History),

London, England; Dr. F. W. Mead, Division of Plant Industry, Florida De-

partment of Agriculture, Gainesville, Florida; Dr. S. Ramalingam, University

of Malaya, Kuala Lumpur, Malaysia; Dr. M. R. Roberts, Academy of Natural

Sciences, Philadelphia, Pennsylvania; Dr. L. E. Rozeboom, John Hopkins

School of Hygiene & Tropical Medicine, Baltimore, Maryland; Dr. O. S. Flint,

Jr., Dr. D. R. Davis and Mr. W. H. Rowe, Smithsonian Ceylonese Insect

Project; Dr. W. A. Steffan, Bishop Museum, Honolulu, Hawaii; Dr. A. Stone,

United States National Museum (Natural History), Washington, D. C.; and

Dr. R. L. Wenzel, Field Museum of Natural History, Chicago, Illinois.

Appreciation is expressed to Mrs. Janet Rupp for typing the manuscript for

offset reproduction. I am especially grateful to my wife, Mollie, for her

encouragement and assistance in typing the drafts.

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1963. The mosquito fauna of Thailand (Diptera: Culicidae). Proc.

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1955. Report of the initial operation of a mosquito light trap in

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1962. Okologische Analyse der Stechmuckenpopulationen in der

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1956. Especies y distribucion de mosquitos Mexicanos no anofelinos

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1963. Contribution to the east Slovakian mosquitoes of the genus

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110 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

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1927. An experimental study on twenty-four species of Japanese

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1950. Mosquito fauna of Guam. H.Q. 8th Army. 101 p.

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1938. Studies on Dirofilaria immitus Leidy with special reference to

the susceptibility of some Minnesota species of mosquitoes to

the infection. J. Parasit. 24: 189-205.

YOSHIMEKI, M.

1955. Morphological studies on the tracheal system of two Culicini

larvae, Culex pipiens L. var. pallens Coquillet [t] and Aedes

vexans nipponii Theobald. Ecol. Rev., Japan 14: 81-89.

Reinert: Aedes (Aedimorphus) in Southeast Asia 111

APPENDIX: TABLE 1. Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) alboscutellatus

Hair Range Mode Mean = ~~ *~Hair + Range Mode Mean

Cephalothorax Abdomen II

1 2-4 3 2.6 0 1 1 1

2 2-4 3 3.1 1 3-7 4 4.7

3 2-4 3 2.8 2 1 1 1

4 3-5 4 4.1 3 1-2 2 1.6

5 4-6 5 5 4 2-5 3 3.5

6 2-3 2 2.3 5 3-6 5 4.6

7 3-4 4 3.8 6 1-3 3 2.6

8 5-9 6 6.3 7 2-3 2 2.1

9 1-3 3 2.3 9 1 1 1

Metanotum Abdomen III

10 3-5 3 3.5 0 1 1 1

11 1 1 1 1 6-10 7 i Sa

12 4-7 5 5.4 2 1 1 1

Abdomen I 3 1 1 1

1 17-22 18 18.6 4 2-4 2 2.7

2 1 1 1 5 3-8 4 ag

3 1 1 1 6 2-4 3 3.1

4 6-11 8 ee 7 2-4 2 2.6

5 2-5 2 2.7 8 2-4 4 3.7

6 2-3 2 2.3 9 1 1 1

7 2-3 2 2.4 10 2-4 3 2.8

9 1 1 1 11 1 1 1

14 1 1 1

112

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 1. (Continued)

Hair

Range Mode

Abdomen IV

1 1

3-6 5

1 1

3-6 5

2-4 2

2 2

1-3 3

2-3 2

2-4 2

1 1

2-4 3

1 1

Abdomen V

1 1

2-5 4

1 1

2-5 3

4-9 2

2 2

1-3 2

5-9 6

2-3 3

1 1

Mean

6.3

2.6

Hair

Range Mode

Abdomen V (Cont. )

2-3 2

1 1

Abdomen VI

1 1

2-4 &

1 1

2-4 3

3-9 5

2-3 2

2-4 3

2 2

2-4 3

1 1

Abdomen VII

1 1

4-6

1 1

2-6 4

3-5 3

3-6 4

Mean

2.1

3.7

3.9

Reinert: Aedes (Aedimorphus) in Southeast Asia 113

TABLE 1. (Continued)

Hair Range Mode

Mean

Abdomen VII (Cont. )

6 4-10

7 1-3

8 3-4

9 2-5

10 2-3

11 2-3

14 1

APPENDIX: TABLE 2.

Hair Range Mode

Cephalothorax

1 3 3

2 2-4 3

3 2-4 3

4 3-6 4

5 2-5 4

6 1-3 1

7 2-4 3

8 5-7 °

9 3-4 3

Metanotum

10 7-11 9

11 1 1

it 5-8 5

6.4

2.1

Hair Range Mode Mean

Abdomen VIII

0 1 1 1

4 2-4 3 3

9 4-7 6 6.2

14 1 1 1

Paddle

1 2-3 2 2.2

Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) caecus

Mean

0.9

Hair Range Mode Mean

Abdomen I

1 18-30 21 24.1

2 1 1 1

3 1 1 1

4 6-10 7 7.3

8 2-5 3 3.6

6 1 1 1

7 2-5 3 2.8

9 1 1 1

10 1 1 1

11 1-2 1 1.2

Abdomen II

0 r 1 1

114

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 2. (Continued)

Hair

Range Mode Mean Hair

Abdomen II (Cont. )

i 1 1 2

1 1 1 3

2-5 4 4.3 4

3-5 4 4.1 5)

1-2 1 Wy 6

1-3 2 2.1 t

1 1 1 8

Abdomen III 9

1 1 1 10

Bak: Li 7 de 11

1 1 1 14

1 1 1

2-3 3 2.6 0

4-6 5) 4.7 1

2-4 3 3 2

2-5 2 2.7 3

2-4 2 2.90 4

1 1 1 5

2-4 3 3 6

1 1 1 1e:

1 1 1 8

Abdomen IV 9

1 1 1 10

4-6 6 Pe: 11

Abdomen IV (Cont. )

4-7

1-2

2-3

2-4

2-3

Abdomen V

Range Mode

Mean

9.9

2.1

Reinert: Aedes (Aedimorphus) in Southeast Asia f13

TABLE 2. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen V (Cont. ) Abdomen VII (Cont. )

14 1 1 1 2 1 1 1

Abdomen VI 3 3-7 5 4,7

0 1 1 1 % 2-4 3 3.2

1 2-5 4 3.8 5 2-4 2 2.6

2 1 1 1 6 3-7 4 4.7

3 2-3 3 2.6 7 1-2 1 1,2

4 3-6 & 4.3 8 2-4 2 2.8

5 1-3 2 2.1 9 3-7 5 5 a |

6 1-2 1 1.2 10 1-3 2 Lo

7 1-2 1 Ld 11 1 1 1

8 1-3 2 2.1 14 1 1 1

9 : 1 1 Abdomen VII

10 1 1 1 0 1 1 1

11 1 1 1 4 2-4 3 3.1

14 1 1 1 9 7-11 9 8.6

Abdomen VII 14 1 1 1

0 1 1 1 Paddle

116 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

APPENDIX: TABLE 3. Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) culicinus

Hair Range Mode Mean Hair Range Mode Mean

Cephalothorax Abdomen II

1 3-4 3 3.4 0 1 1 1

2 2-4 3 3.1 1 14-20 14 15:9

3 3-4 3 3.3 2 1 1 1

4 3-4 3 3.4 3 1 1 1

5 4-6 4 4.9 4 4-8 5 5.3

6 2 2 2 5 4-7 5 5.6

7 3-5 4 4.3 6 1-2. 2 1.6

8 5-9 7 6.4 7 2-4 3 2.6

9 2-3 3 2.6 9 1 1 1

Metanotum Abdomen III

10 6-9 7 Tne 0 1 1 1

11 1 1 1 1 7-13 8 8.9

12 4-7 4 5.3 2 1 1 1

Abdomen I 3 1 1 1

1 16-24 21 20 4 3-5 4 3.8

2 1-2 1 1.2 5 4-8 5 5.1

3 1 1 1 6 2-3 3 2.6

4 7-12 #11 9.3 7 3-5 3 3.5

5 2-6 4 3.9 8 3-5 4 4.1

6 1-3 2 2.1 9 1 1 1

7 2-4 3 a7 10 1-4 2 2.3

9 1 1 1 11 1 1 1

14 1 1 1

Reinert: Aedes (Aedimorphus) in Southeast Asia

TABLE 3. (Continued)

Hair

Range Mode

Abdomen IV

1 1

4-9 5

1 1

4-7 6

2-3 2

2 2

2-3 2

2-4 3

2-5 3

1 1

1-3 3

1 1

Abdomen V

1 1

4-9 5

1 1

2-4 3

4-7 6

2-3 2

1-3 2

4-8 6

2-4 3

1 1

Mean

Hair

Abdomen V (Cont. )

1-2

Abdomen VI

Abdomen VII

Range Mode

117

Mean

1,4

3.9

6.3

118 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 3. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen VII (Cont. ) Abdomen VIII

6 6-97 7. 6 0 1 1 1

7 248 2. 6 4 3-6 4 3.9

8 BG 3 3.5 9 11 9 8. 6

9 45 4 4.4 14 1 1 1

10 Zo 2 a8 Paddle

11 2 2 2 1 SS 1.4

14 1 1 1

APPENDIX: TABLE 4. Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) mediolineatus

“Hair Range Mode Mean —=—=s Hair Range Mode Mean

Cephalothorax Abdomen I

1 3-5 4 3.6 1 17-30 22 24.6

2 3-4 4 3.6 2 1 1 1

3 2-4 3 2.9 3 1 1 1

4 2-4 3 3.3 4 8-17 12 13.9

5 3-5 4 3.8 5 3-6 4 4.2

6 2-4 3 3.1 6 1-2 1 1,1

7 2-5 4 3.6 7 2-3 2 2.4

8 2-6 3 3.6 9 1 1 1

9 2 2 2 Abdomen II

Metanotum 0 1 ee 1

10 11-24 12 13.8 1 17-28 22 22.8

11 1 1 1 2 1-2 1 1.1

12 4-5 4 4,4 3 1-2 1 1.3

Reinert: Aedes (Aedimorphus) in Southeast Asia

TABLE 4. (Continued)

Hair

Range Mode Mean

Abdomen II (Cont. )

6-10 7 7.6

4-8 6 5.5

1 1 1

1-4 2 2.6

1 1 1

Abdomen III

1 1 1

7-18 15 12.2

1 | 1

1 1 1

2-8 4 3.9

4-9 5 5.6

2-4 2 2.3

2-5 3 3.4

3-6 4. 4.4

1 1 1

2-4 3 2.7

1 1 1

Abdomen IV

1 1 1

3-7 5 5.1

1 1 1

5-12 6 148

Hair

Abdomen IV (Cont. )

2-6

2-4

Abdomen V

Range Mode

119

Mean

120 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 4. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen VI Abdomen VII (Cont. )

0 1 1 1 3 3-7 5 D. 1

1 4-5 5 4.6 4A 2-5 3 3.2

2 1 1 1 5 3-6 5 4.6

3 2-3 2 2.3 6 8-12 11 10.5

4 4-8 5 bs 1 7 2-3 3 2.6

5 2-5 A. 3.2 8 3-7 5 5.6

6 2-4 3 2.7 9 5-11 5 6.8

7 1-2 1 1,2 10 1-3 2 2.3

8 3-5 a 3.8 11 1-3 2 1.8

9 1 1 1 14 1 1 1

10 1 1 1 Abdomen VIII

11 1 1 1 0 1 1 1

14 1 1 1 4 2-4 3 2.9

Abdomen VII 9 7-14 8 8.9

0 1 1 1 14 1 1 1

1 3-6 4 4.4 Paddle

APPENDIX: TABLE 5. Record of the branching of the setae on the

Hair

Range Mode Mean

Cephalothorax

3-6 5 4.5

3-4 3 3.3

2-3 3 2.6

3-6 4 4.3

3-6 4 3.9

2-3 2 ouch

3-5 4 3.7

7-9 : 7.5

2-4 3 2.9

Metanotum

17-33 18 22.3

: 1 1

5-10 5 6.2

Abdomen I

16-24 21 21.1

1 1 1

7-16 10 10.1

3-7 4 4.4

2-3 2 2.4

3-6 4 4.3

1 1 1

Reinert: Aedes (Aedimorphus) in Southeast Asia

pupae of Aedes (Aedimorphus) orbitae

Hair

Range Mode

Abdomen IT

1 1

33-65 38

1 1

3-6 5

3-5 4

1-3 2

2-6 4

1 1

Abdomen III

1 1

8-17 12

1 1

2-5 4

3-5 4

1-4 4

2-5 4

1 1

4-6 a

1 1

121

Mean

41.5

1,8

4.2

122 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 5. (Continued)

Abdomen IV Abdomen V (Cont. )

0 1 1 1 11 1 1 1

1 5-11 10 8.6 14 1 1 1

2 1 1 1 Abdomen VI

3 4-7 4 5.3 0 1 1 1

4 1-2 2 1.6 1 4-7 5 5.1

5 2 2 2 2 1 1 1

6 1-3 2 2.4 3 2-4 3 3.2

7 1-3 2 1,9 4 4-6 5 4.8

8 2-4 3 2.9 5 1-2 2 Lee

9 1 1 1 6 1 1 1

10 3-5 4 3.9 7 1-3 2 1.9

11 1 1 1 8 2-4 2 2.2

14 1 1 1 9 1 1 1

Abdomen V 10 1 1 1

0 1 1 1 11 1 1 1

1 0-10 6 6.5 14 1 1 1

2 1 1 1 Abdomen VII

3 2-3 3 a 0 1 1 1

4, 5-11 8 7.3 1 3-6 4 4.1

5 2 2 2 2 1 1 1

6 1-3 2 1.8 3 4-6 5 4.9

7 5-8 7 6.8 4 2-4 3 2.7

8 2-4 2 3.3 5 2-4 2 2.7

9 1 i! 1 6 6-11 7 8.1

10 k 1 1

Reinert: Aedes (Aedimorphus) in Southeast Asia

TABLE 5. (Continued)

Hair Range Mode Mean

Abdomen VII

7 1-3 2 1.9

8 3-5 3 3.6

9 4-6 5 0.2

10 2-3 2 2.4

11 1 1 1

14 1 1 1

Hair

Range Mode

Abdomen VIII

1 1

2-3 3

8-13 9

1 1

Paddle

1 1

123

Mean

10.2

APPENDIX: TABLE 6. Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) pallidostriatus

Hair Range Mode Mean

Cephalothorax

1 2-4 3 3

2 2-3 3 2.8

3 3-4 3 3.3

“ 3-5 3 3.6

5 3-6 4. 4.4

6 2-4 2 2.4

7 2-3 3 2.9

8 2-6 3 4.8

9 2-3 2 2.3

Metanotum

10 10-18 12 13.3

11 1 1 1

12 3-7 + 4.7

Hair

Range Mode

Abdomen I

30-46 35

1 1

1-2 1

8-15 10

4-7 5

1-2 1

2-4 3

1 1

Abdomen II

1 1

Mean

37.7

124 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 6. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen II (Cont. ) Abdomen IV (Cont. )

2 1 1 1 2 1 1 1

3 2-3 3 2.6 3 4-10 7 6.9

4 5-9 7 7 4 2-5 3 3

5) 2-9 6 4.9 9) 2-3 3 2.9

6 1-3 2 1.9 6 2-6 3 3.6

7 3-4 4, 3.6 7 3-6 4 4,1

9 1 1 1 8 3-4 3 Bick

Abdomen III 9 1 1 1

0 1 1 1 10 2-4 3 2.9

1 7-18 18 13, 4 11 1 1 1

2 1 1 1 14 1 1 1

3 1 1 1 | Abdomen V

4 3-5 5 4.4 0 1 1 1

5 3-11 8 7,2 1 4-6 6 5.4

6 2-4 3 2.8 2 ZX 1 1

7 4-7 5 5.4 3 2-3 3 2.6

8 3-7 4 4.8 4. 0-9 8 7.3

9 1 1 1 5 3-4 3 3.1

10 2-3 2 a, 6 2-3 3 2.6

11 1 1 1 a 8-11 8 8.7

14 1 1 1 8 3-5 4 4

Abdomen IV 9 1 1 1

0 1 1 1 10 1-3 1 1.3

1 0-13 is ao 11 1 1 1

Reinert: Aedes (Aedimorphus) in Southeast Asia 125

TABLE 6. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen V (Cont. ) Abdomen VII (Cont. )

14 1 1 1 2 1 1 1

Abdomen VI 3 4-9 , 1%

0 1 1 1 4 2-4 4 3.4

1 4-9 5 6.4 5 4-9 7 6.7

2 1 1 1 6 8-13 10 10.6

3 2-4 3 3.2 7 1-4 3 2.4

A 3-7 6 5.6 8 3-7 3 4

5 3-4 3 3.4 9 3-8 6 5.9

6 2-3 3 2.5 10 1-3 2 1.9

7 2-3 2 aut 11 1-3 1 1.5

8 3-4 4 3.6 14 1 1 1

9 1 1 1 Abdomen VIII

10 1-2 1 1.1 0 1 1 j

11 1-2 1 1.1 4 2-3 3 2.8

14 1 1 1 9 7-13 9 9.8

Abdomen VII 14 1 1 1

0 1 1 1 Paddle

1 5-9 7 6.5 1 1 1 1

126

APPENDIX: TABLE 7. Record of the branching of the setae on the

Hair

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Range Mode

Cephalothorax

5-8 5

4-8 5

3-5 5

6-8 7

6-9 6

2-4 3

8-11 9

7-12 12

2-4 3

Metanotum

13-16 13

1 1

3-11 6

Abdomen I

21-42 35

1 1

10-18 15

5-7 6

2-3 2

4-6 5

1 1

pupae of Aedes (Aedimorphus) pampangensis

Mean

32.3

Hair

Range Mode

Abdomen II

1 1

24-40 36

1 1

1-2 2

6-11 G..!

10-14 10

2-5 3

6-10 6

1 1

Abdomen III

10-16

Mean

33.6

12.4

Sei

Reinert: Aedes (Aedimorphus) in Southeast Asia 127

TABLE 7. (Continued)

Abdomen III (Cont. ) Abdomen V (Cont, )

11 1 1 1 % 7-11 9 8.9

14 1 1 1 8 3-4 3 3.4

Abdomen IV 9 : 1 1

0 1 1 1 10 2-4 3 3

1 o-11 7 TG 11 1 1 1

2 1 1 1 14 1 1 1

3 0-9 8 7.4 Abdomen VI

4 3-6 3 4.1 0 1 1 1

5 2-5 3 3.1 1 7-13 9 9.5

6 5-8 6 6.6 2 1 1 1

7 4-6 4 4.4 3 4-8 4 5. 4

8 3-4 3 3.4 4 7-11 8 9.9

9 1 1 1 5 3-6 5 4.4

10 3-5 4 4.3 6 6-8 8 7.3

11 1 1 1 7 2-4 3 2.8

14 1 1 1 8 4-6 4 4.8

Abdomen V 9 1 1 1

0 1 1 1 10 1-3 2 2

1 5-9 8 8.6 11 1 1 1

2 1 1 1 14 1 1 1

3 3-5 4 4 Abdomen VII

4 9-14 12 11.6 0 1 1 1

5 3-5 3 3.8 1 7-10 7 8.1

128 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 7. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen VII (Cont. ) Abdomen VII (Cont. )

3 9-16 11 12.1 11 2-4 3 2.8

4 4-10 6 Ta 14 1 1 1

5 5-9 . 7.3 Abdomen VIII

6 7-14 10 10.1 0 1 1 1

7 2-5 4 4.1 4 5-8 7 6.4

8 3-7 5 5 es 9 11-14 14 12.9

9 5-7 6 6.1 14 1 1 1

10 2-5 4 3.6 Paddle

1 1-2 2 1,8

APPENDIX: TABLE 8. Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) pipersalatus

Hair Range Mode Mean Hair Range Mode Mean

Cephalothorax Metanotum

1 2-3 3 2.6 10 5-9 6 6.2

2 2-4 3 2.9 11 1 1 1

3 2-4 2 2.6 12 4-7 4 5

4 3-4 3 3.3 7 Abdomen I

D 3-5 4 4.3 1 21-38 31 29.1

6 2-4 3 2.9 2 1 1 1

ij 3-6 A 4.2 3 1 1 1

8 2-7 5 4.6 4 7-16 9 10.9

TABLE 8.

Hair

Reinert: Aedes (Aedimorphus) in Southeast Asia

Range

(Continued)

Mode

Mean

Abdomen I (Cont. )

1-3 1

3-5 3

1 1

Abdomen II

1 1

13-20 18

1 1

7-16 8

4-7 5

1-3 3

2-4 3

1 1

Abdomen III

1 1

5-10 7

1 1

3-6 5

3-7 4

2-4 2

3-9 4

3-5 3

1.5

17.4

7.8

Hair

Range Mode

1 1

Abdomen III (Cont. )

2-4 3

1 1

Abdomen IV

1 1

3-6 6

1 1

o-7 5

2-5 3

2 2

1-4 2

3-5 3

2-3 2

1 1

2-4 2

1 1

Abdomen V

1 1

3-5 4

1 1

2-3 2

4-8 5

2-3 2

129

2.8

0.3

2.3

130

TABLE 8. (Continued)

Hair

Range

Mode

Mean

Abdomen V (Cont. )

Abdomen VI

2-4 3

0-8 7

2-3 3

1 1

1-2 i

1 1

3-7 5)

1 1

2-3 2

3-6 9)

2-5 3

2-3 2

2 2

2-4 3

1 .

1-2 1

1 1

2.7

6.7

2.8

1,1

2.9

1,2

Hair

Range Mode

Abdomen VII

1 1

3-6 A

1 1

3-9 9)

2-4 3

3-6 é,

6-14 8

2-3 2

2-5 3

3-6 4

1-2 2

1 1

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Abdomen VII

3-4

Paddle

Mean

3.1

7.1

Reinert: Aedes (Aedimorphus) in Southeast Asia 131

APPENDIX: TABLE 9. Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) vexans vexans

Hair Range Mode Mean Hair Range Mode Mean

C ephalothorax Abdomen II

1 2-3 2 2.3 0 1 1 1

2 33 oy 24 1 5-14 10 9.4

3 2 2 2 2 1 1 1

4 1-4 2 2.5 3 1-2 2 i

5 1-3 2 2.4 4 2-4 3 2c

6 1-4 1 La] 5 2-5 4 3.3

7 2-5 3 3.2 6 1-3 1 1.2

8 2-4 4 3.4 Zi 1-3 1 1.4

9 1-2 2 1.8 9 1 1 1

Metanotum Abdomen ITI

10 4-11 7 7 0 1 1 1

11 1 1 1 1 3-5 “ Bee

12 1-3 1 at 2 1 1 1

Abdomen I 3 1-2 1 1,2

1 17-25 21 21 “4 1-4 3 2.5

2 1 1 1 5 2-6 4. 3.8

3 1 1 i 6 1-3 1 1.5

4 2-8 3 a2 7 2-4 3 2.9

5 2-4 2 2.8 8 1-4 2 2.1

6 1-2 2 44 9 1 1 i

7 1-3 1 1.2 10 1-3 1 1.8

9 1 1 1 11 1 1 1

10 i 1 1 14 1 1

132

TABLE 9. (Continued)

Hair

Abdomen IV

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Mode

Range

1 1

2-5 3

1 1

2-7 5

1-5 1

1-3 2

1-3 1

2-4 2

1-3 1

1 1

1-2 1

1 1

1-3 2

1 1

1-4 2

1-5 3

1-2 2

1-2 1

1-5 2

1-3 1

1 1

Mean

Hair

Range Mode

Abdomen V (Cont. )

Abdomen VI

Abdomen VII

Mean

2.8

1.6

Reinert: Aedes (Aedimorphus) in Southeast Asia 133

TABLE 9. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen VII (Cont. ) Abdomen VII

6 2-5 3 3.5 0 1 1 1

i 1 1 1 & 1-2 2 1.7

8 1-4 2 2.1 9 4-7 5 5.3

9 2-4 3 3.3 14 1 1 1

10 \ 1 1 Paddle

11 1 1 1 1 1 1 1

14 1 1 1

APPENDIX: TABLE 10. Record of the branching of the setae on the

pupae of Aedes (Aedimorphus) vexans nipponii

Hair Range Mode Mean Hair Range Mode Mean

Cephalothorax Metanotum (Cont. )

1 Bug See 2.6 120 oT ee 1.6

2 | 2-3 2 2.1 Abdomen I

3 2-4 2 2.3 1 16-22 18 18.4

4 2-3 3 2.9 2 1 1 1

5 2-4 3 2.8 3 1 1 1

6 1-4 3 2.4 4 2-5 3 3.3

i 2-4 4 3.4 5 1-3 3 2.7

8 3-6 5) 4.5 6 1-2 1 1.4

9 1-2 1 1.4 7 2-4 3 2.9

Metanotum 9 1 1 1

10 4-10 5) 2.8 10 I 1 1

11 1 1 1

134 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 10. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen II Abdomen IV

0 1 1 1 0 1 1 1

1 4-10 6 7.2 1 2-5 3 3.2

2 1 1 1 2 1 1 1

3 1-2 1 1.4 3 3-6 5 4.7

4, 2-4 3 2.8 4 1-4 2 2.1

5 2-5 3 2.9 5 2-3 2 2.1

6 1-2 1 1.3 6 1-3 2 1.8

7 2-5 2 2.6 7 1-4 2 1.8

9 1 1 1 8 1-3 2 2

Abdomen III 9 1 1 1

0 1 1 1 10 1-3 2 2

1 3-6 4 4.3 11 1 1 1

2 1 1 1 14 1 1 1

3 1-2 1 oa | Abdomen V

4 2-5 3 2.9 0 1 1 :

5 2-7 4 4,2 1 2-4 3 3

6 1-3 1 {,9 2 1 1 1

7 2-4 3 2.7 3 2-4 2 2.6

8 1-5 3 2.8 4 1-5 3 2.8

9 1 1 1 5 2-3 2 2.1

10 1-3 2 1.8 6 1-2 2 at

11 1 1 1 7 1-5 2 2.3

14 1 1 1 8 1-3 2 19

Reinert: Aedes (Aedimorphus) in Southeast Asia 135

TABLE 10. (Continued)

Hair Range Mode Mean Hair Range Mode Mean

Abdomen V (Cont. ) Abdomen VII (Cont. )

9 1 1 1 1 1-3 2 19

10 1 1 1 2 1 1 1

11 1 1 1 3 2-7 3 3.3

14 i 1 1 4 1-3 2 1.8

Abdomen VI 3) 2-3 2 2.2

0 1 1 1 6 3-5 4 3.6

1 2-5 3 2.8 7 1-2 1 1.1

2 1 1 1 8 2-5 5 rat |

3 1-3 1 1.3 9 2-4 3 3

4 2-4 3 3 10 1 1 1

5 2 2 2 11 1-2 1 ita |

6 1-2 2 1.8 14 1 1 1

7 1-2 1 at Abdomen VII

8 2-4 2 2.2 0 1 1 1

9 1 : 1 4 1-4 2 2.2

10 1 1 1 9 4-7 5 5.4

11 1 1 1 14 1 1 1

14 1 1 1 Paddle

Abdomen VII 1 1-2 1 1,2

0 1 1 1

136

APPENDIX: TABLE 11.

Species

alboscutellatus

argenteoscutellatus

caecus

culicinus

davidi

gouldi

jamest

lowisit

mediolineatus

nigrvostriatus

orbitae

pallidostriatus

pampangensis

pipersalatus

punctifemoris

stenoetrus

syntheticus

taentorhynchoides

trimaculatus

vexans vexans

wainwrighti

Female

X**

Male

X **

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

Pupa

X**

X **

Species of Aedes (Aedimorphus) occurring in the

Oriental Zoogeographical Region

Larva Keg

X** =

X= Indicates stage has been described in this paper or in the literature.

* = [Indicates a portion of the stage has been figured in the literature.

** = Indicates a portion of the stage is figured in this paper.

- = Indicates no description or figure.

Reinert: Aedes (Aedimorphus) in Southeast Asia

137

APPENDIX: TABLE 12: Species of Aedes (Aedimorphus) occurring in the

Species

alboscutellatus

caecus

Lowisit

VEXANS VEXANS

APPENDIX: TABLE 13.

Species

alboscutellatus

vexans vexans

vexans nipponti

APPENDIX: TABLE 14.

Species ©

alboscutellatus

caecus

oakleyi

senyavinensis

trukensis

VEXaANS VEXANS

Australian Zoogeographical Region

Female

X**

X **

Male

X**

X **

» Go.

X **

Pupa

X**

Larva Egg

X** se

Species of Aedes (Aedimorphus) occurring in the

Palearctic Zoogeographical Region

Female

x **

Male

X**

X **

Pupa

x ee

XK **

X **

Larva Egg

X** Ke

X** X*

X** Hn

Species of Aedes (Aedimorphus) occurring in the

Pacific Ocean Islands Region

Female

X **

Male

X **

Pupa

X **

X**

xX*

X **

Larva Egg

X** _

X** P

X* -

X** xX*

X = Indicates stage has been described in this paper or in the literature.

* = Indicates a portion of the stage has been figured in the literature.

** = Indicates a portion of the stage is figured in this paper.

- =Indicates no description or figure.

138 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

APPENDIX: TABLE 15. Species of Aedes (Aedimorphus) occurring in the

Nearctic Zoogeographical Region

Species Female Male Pupa Larva Egg

VEXANS VEXANS » Seas x re D Gases x** >. Ge

APPENDIX: TABLE 16. Species of Aedes (Aedimorphus) occurring in the

Ethiopian Zoogeographical Region

abnormalis - x* He ws =

abnormalis kabwachensis x X* ».4 x 3

adami - xe = a :

albocephalus xt X* X* Dea -

alboventralis x* - “ * 2

apicoannulatus xX X = = 2

argenteopunctatus x* xX* ~ x ~

bedfordi 4 x* “ = .

bevisi X x* X X* “

boneti - X* _ - a

boneti kumbae fs x* X X* na

caliginosus X xX* - - ‘

capensis x* x* ~ X* -

centropunctatus xX x* x* x* :

congolensis Xx x* . is .

chamboni X* x* = = _

CUMMINSI1 K* x* Xx x* -

X = Indicates stage has been described in this paper or in the literature.

* = Indicates a portion of the stage has been figured in the literature.

** = Indicates a portion of the stage is figured in this paper.

- = Indicates no description or figure.

Reinert: Aedes (Aedimorphus) in Southeast Asia 139

TABLE 16. (Continued)

Species Female Male Pupa Larva Egg

dalzieli x* x* ~ ».4 -

dentatus x* x* ~ Po ~

diallot = x* ws - 2

domesticus ».4 x* ».4 x X*

durbanenstis x* Dee A X* -

ebogoensis ~ x* ~ Fe 2

evitreae ».4 X* - - ~

eritreae karooensis Xx ee ~ PG -

falabreguest ~ x* o é -

filicis x* x* x* x* -

fowleri x* x* xX x* -

gibbinst ».4 X* xX* xX ~

gilliest D4 D4 ~ - ~

grentevi - xX* - - -

grjebinet - < “ re a

hamonti x* x* - E. =

haworthi x* x* X x* -

hirsutus ».4 x* X* xX* ~

holocinctus ~ X x* X* ~

hopkinsi ».4 x* ».4 a ~

insolens <x ‘i 6 ” ‘

irritans x* xX * a i, Sas -

kapretwae xX x i. x* “

kennethi D4 D4 ~ x* ~

lamborni ; = xX* X X* a

140 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

TABLE 16. (Continued)

leesoni ~ x* : is =

leesoni verna xX x* ba xX* a

leptolabis = x* x* X =

leucarthrius D4 - ~ “ =

lokojoensis i x* x xX .:

longiseta ~ x* - . 7

lottez ~ x* ~ ' :

MANS OUY1 X xX* a - -

marshallit X* x* x* xX*

mattinglyt D4 x* x* x* 3

microstictus — - x* re i é

minutus x* x* = 4 ee

mixtus = X* ‘i X s

mutilus X x _ ».4 -

natronius x x* be X* 2

neobiannulatus xX ‘. “ " 2

ngong x » 4 Xx x .

nigricephalus X* x* X* xX* ~

nyounae = xX* x* x* S

ochraceus x* x* i X* fs

OVaAzZZAt - x* . - -

pachyurus x x* > X ”

phyllolabis ».4 x* a x -

pseudotarsalis xX a* x* x* Z

pubescens X x = > x

Reinert: Aedes (Aedimorphus) in Southeast Asia

TABLE 16. (Continued)

Species

punctothoracis

qua siunivittatus

reali

rickenbachi

semlikiensis

seychellensis

simulans

smithburnt

stokest

subdentatus

tarsalis

tauffliebt

teesdalei

tiptoni

tricholabis

tricholabis bwamba

vexans vexans

wendyae

wigglesworthi

yangambiensis

yvonneae

X = Indicates stage has been described in this paper or in the literature.

Female

xX*

Male

Pupa

xX *

X *

Larva

xX*

* = Indicates a portion of the stage has been figured in the literature.

** = Indicates a portion of the stage is figured in this paper.

- =Indicates no description or figure.

141

142

OMmAADIAIHLWNH-E

17,

Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

FIGURES

Aedes (Aedimorphus) alboscutellatus adult morphology

Aedes (Aedimorphus) caecus adult morphology

Aedes (Aedimorphus) culicinus adult morphology

Aedes (Aedimorphus) lowisii adult morphology

Aedes (Aedimorphus) mediolineatus adult morphology

Aedes (Aedimorphus) nigrostriatus adult morphology

Aedes (Aedimorphus) orbitae adult morphology

Aedes (Aedimorphus) pallidostriatus adult morphology

Aedes (Aedimorphus) pampangensis adult morphology

Aedes (Aedimorphus) pipersalatus adult morphology

Aedes (Aedimorphus) punctifemoris adult morphology

Aedes (Aedimorphus) stenoetrus adult morphology

Aedes (Aedimorphus) taeniorhynchoides adult morphology

Aedes (Aedimorphus) vexans vexans adult morphology

Aedes (Aedimorphus) vexans nipponii adult morphology

Aedes (Aedimorphus) alboscutellatus, caecus, culicinus, lowisii,

mediolineatus, nigrostriatus, orbitae,

pallidostriatus, pampangensis, pipersalatus,

punctifemoris, stenoetrus, taeniorhynchoides,

vexans vexans,and vexans nipponii female tarsomeres

5 and posttarsi

Aedes (Aedimorphus) alboscutellatus, caecus, culicinus, lowisii, medio-

lineatus, orbitae, pallidostriatus, pampangensis,

pipersalatus, punctifemoris, stenoetrus, taenior-

hynchoides, vexans vexans and vexans nipponii male

tarsomeres 5 and posttarsi

Aedes (Aedimorphus) alboscutellatus female genitalia

Aedes (Aedimorphus) caecus female genitalia

Aedes (Aedimorphus) culicinus female genitalia

Aedes (Aedimorphus) lowisii female genitalia

Aedes (Aedimorphus) mediolineatus female genitalia

Aedes (Aedimorphus) nigrostriatus female genitalia

Aedes (Aedimorpbhus) orbitae female genitalia

Aedes (Aedimorphus) pallidostriatus female genitalia

Aedes (Aedimorphus) pampangensis female genitalia

Aedes (Aedimorphus) pipersalatus female genitalia

Aedes (Aedimorphus) punctifemoris female genitalia

Aedes (Aedimorphus) stenoetrus female genitalia

. Aedes (Aedimorphus) taeniorhynchoides female genitalia

Aedes (Aedimorphus) vexans vexans female genitalia

Aedes (Aedimorphus) vexans nipponii female genitalia

Aedes (Aedimorphus) mediolineatus egg; alboscutellatus and vexans

vexans lateral view of aedeagus representing Type I

and Type II

Aedes (Aedimorphus) alboscutellatus male genitalia

Aedes (Aedimorphus) caecus male genitalia

Aedes (Aedimorphus) culicinus male genitalia

Aedes (Aedimorphus) lowisii male genitalia

Aedes (Aedimorphus) mediolineatus male genitalia

Aedes (Aedimorphus) nigrostriatus male genitalia

Reinert: Aedes (Aedimorphus) in Southeast Asia 143

Aedes (Aedimorphus) orbitae male genitalia

Aedes (Aedimorphus) pallidostriatus male genitalia

Aedes (Aedimorphus) pampangensis male genitalia

Aedes (Aedimorphus) pipersalatus male genitalia

Aedes (Aedimorphus) punctifemoris male genitalia

Aedes (Aedimorphus) stenoetrus male genitalia

Aedes (Aedimorphus) taeniorhynchoides male genitalia

Aedes (Aedimorphus) vexans vexans male genitalia

Aedes (Aedimorphus) vexans nipponii male genitalia

Aedes (Aedimorphus) alboscutellatus pupa

Aedes (Aedimorphus) caecus pupa

Aedes (Aedimorphus) culicinus pupa

Aedes (Aedimorphus) mediolineatus pupa

Aedes (Aedimorphus) orbitae pupa

Aedes (Aedimorphus) pallidostriatus pupa

Aedes (Aedimorphus) pampangensis pupa

Aedes (Aedimorphus) pipersalatus pupa

Aedes (Aedimorphus) vexans vexans pupa

Aedes (Aedimorphus) vexans nipponii pupa

Aedes (Aedimorphus) alboscutellatus larva

Aedes (Aedimorphus) caecus larva

Aedes (Aedimorphus) caecus variations in patches of spicules on larval

siphon

Aedes (Aedimorphus) culicinus larva

Aedes (Aedimorphus) mediolineatus larva

Aedes (Aedimorphus) orbitae larva

Aedes (Aedimorphus) pallidostriatus larva

Aedes (Aedimorphus) pampangensis larva

Aedes (Aedimorphus) pipersalatus larva

Aedes (Aedimorphus) vexans vexans larva

Aedes (Aedimorphus) vexans nipponii larva

Morphology of female genitalia

alboscute//atus THAILAND Kec

ight nt ier,

st ne ae i is i

itr em ae ics

partr

yaa

Peta rs

et tine

9 | THAILAND

, i " ie ? G2 s oe

‘ culicinus Jam

INDONESIA Shey

it cr tia een

oe epee! rote 1

MMi tf ee lah orc

sigs ay aig Tce :

Pa legit at a iit

rteett pheef tiie ifige fesesest

iC: it

nigrostriatus Bic ee ASSAM, INDIA

Sift nd Le

MALAYSIA

orbijtae

pallidostriatus

wil iy él :

pampangensis

PUR PRRENGE. S

ridirlQ

sents ea ray

een

paresniparestcet strony

canensnerretretrottresind

pee

CAMBODIA cg’ pipersa/atus

tt ant sation

“itty, i

PHILIPPINES** punctifemoris

Stenoetrus

S. Shibata

gt sti

a eaten,

ng jist ae " init! A ant ie

f THAILAND

vexans vexans

ciel ile :

Sg et

vexans nipponii

FEMALE TARSOMERES 5 AND POSTTARSI

= a

(Acc (Aga LEFF

> = QGlllg se: ie A Sy

low/sii

ecco nl

caecus

Stenoetrus vexans vexans

MALE TARSOMERES 5 AND POSTTARSI

sfenoetrus

ESS

Ay:

‘go

Wiss

WSS

NAASe

iy,

\ 4411

\ 44 14,1)

AY VA iY

NY, WT!

ANA Wily)

ANN 114494

ANY 144 Ij 4

AAA 1, Yo)

\ Ny

WN is

RSS VU)

SS 44

RON GY

ES Pip

: C4,

ro 022

VIII-T

Villrs THAILAND

alboscute/l/atus

Fig. 19

Soom

Vid fg

a) i

Wy iy } d

bid) ihn! TN BA

EE fh AURA BT WY |).

\! Vy Wg! UO ft wer iN)

Hh q Wh |

Ny. Viliyt iil G

Wg MW

W J

“) | Wd

SAN il (]

Sw | iN | Ay ,

SSS ‘i \ S41 ly Wi)

SAX AAW) Nn ANTE

SS . \\ Yio\, fl \!

SQ \ \\ TAN At qty

SS y/ BORING

SSSQQy Q i)

SAY

SS \ i]

Ss y l

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j SQ ‘) i)

y RS ~°Qy Hi

“4 SSA aw ) v1

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on wah

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Wy!

hide

in)

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Lila yell

ill

"| Tih

ENS i AN)

2 senate soese 8 Mays sy

Ji Y x . 557;

ap) . 7

7) eS 4

Pye A THO NOC :

aM , » pirat ay

ay if) ai Sb

2\"

0.1

m Kasur

VIII-T

THAILAND

caecus

Fig . 20

THAILAND

culicinus

Fig. 21

\\4 /

A? |\ i /

yy! /

A 14 Y

O Wf / Lines

CY. Ws

Ye} WCC VST

Lr PAL LITE. <

D Vi Ap ys ek

BELLIES oS

iE ny CGY S

Bd (EEL, it LEG 4) ~

wy) EAILLSS oy a

WS: \ BELL,

vy,

\\ .

SSS

Soult we

EES

FERS, ~

) SN

Wat

RA eae ‘

Ny oN

Ny:

“!

Cn te

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OF ere 2

Sees Soper Pag Na

VIN-T

K. Utsugé

Vil-S INDONESIA

low/sii

as ih | iP i iy <

1 //1/ ‘ Vs ds'| if

f LA, RONG ANY iW ft /

wn Vili Wy CON NK “| e)

aM, 1, Wii VARA Va 14

CAINS Ws 4

Y) Yi VN A NYG

BAIN UTA WA'LRNRNERAG (eV [17

NEO SASS RN ON) 16/7

VELMA IYI SARS Ve,

EL TE SUN RRO ANG

Ver es 5 YZ \\ /\ HSA \) I, if,

el (ee VAD WSIS SA i Vasa

VIE ar ter ehie VT IAS SY) MIA CHEE

PES a Te Li Wi iN WAND YY Lt 7 4

Ee Bere CAN | 17 AAW ¥ SQV YC

CEE ACAAN gl “a WMA YN i / We 447;

Va | | ENGI 1 ROAR RAR in

CLE EEG EEE eA PC ORL IS NIN NNSA or) 47G

Pee ee he aH / i by, \ NEESER SOA Se feat oe

PIEOL TE ood eS Nags BE \\ Sn FAN a Te

ly Pe Ep OEE ee TE Ns N ONS RS \ G

bet - N ’ ‘jf ONS eS YAY VUd Gel

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NaN ~»

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TERR

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4 by

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YZ.

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Whi

wate

0.1

VIET

THAILAND

Utsuae

mediolineatus as

VIIl-'S

Fig. 23

ON \\\

} Ww

Yan NY

7, ANN

/ WS

Yi \WW\ Ww

MY \W My

H/; WW) NN

ANY AY y

NV WANN

HI WIRY

Wy) WINRAR

Wy MAY SSSA WY

MIN \INYISSSSS SQN

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4 % ¥ NSS aS SS

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NOY

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DN Das)

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SAAN Behe ot

VILN-T

ASSAM, INDIA

nigrostriatus

Sut

“ys

aa)

NYY.

IX-T _

VIII-T

MALAYSIA ae

orbitae

SSA

SSS

1) \

Dh \

“Hi HN “Nt

SS =

—"

= SS

SSS

. AAS EC GU

4, Ny iu 4)

GIN May lity

Ae tag MU

Meter tery HCE

hiy

4's

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Uist

‘

ma)

ait

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wi at

VII-T

INDIA

pallidostriatus

ANA CCURV TTR]

RAN Ny

SS lle

SSG

Sha

wat Vi

Sonobe

0.2

S55

OIE

{

“| Y

Ty Yi iy

Ye WORM LYS,

WU LM if)

WMG,

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fa \ \\

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7 SAY \

Ct WAG, ROA Watt

Uh O75 54 SO

TT an ERA pao RSQ

(Hel MUO Lb) RA Vid

bat WY YELL SAA

\ Vy \y q

Wa 4 \ NV Nit \\

| fev

wey teh

VIII-T

Sonobe

PHILIPPINES

pampangensis

Life

‘

‘ ii

Av, ie iy a 4

Ay yWaces LY

! ISS ly fi

by i

\ t. fi

143

Mae

VIlI-T

CAMBODIA K Utsugi

VIlI-S pipersa/atus

Fig .28

\ \\

WN)

ANU {

AWW i

SYS

WN RA

RW RANA

WS RAR

WY SSR ||

SARA

SN SA?! \

\ NS SA \

WS SSAA

N SQA!

WS SSSA: :

SS SSS \

x Bt SSS i

HESS SSSA ‘

SSRN it

SERS QAy NY i%4

BSS SSO NN N

SS Qn}

SSO \

SO!

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NOON A\

pt Ni

N \

N ee 1

Nea ge Ca a 4

Cai N \\ \ 4

AROS Nig tY : : ‘ vi He ee ie

Ue at tbh as HON

Yen \

Sah

\ te

Sart

0.2

ede ebay h ats

yest

THY

wigien ey

Mott

VIIN-T

PHILIPPINES > onobe

punctifemoris

Fig. 29

O,s hat

7) ASE

AAS [h

LE ae) Near

\\ RAD SANTOS

Lhd MAA MALY OEE

The EER SASSY HOG

APPLE SSSA i)

7 414,14 9 RSA ANNAN

uv eee 2 yy) SSNS

7 he \} f

4b TTR NN Ss,

Wash, hy State,

WAIT T S 4 SSS a Yj

A RS STS Z

Lee fo ea SESSA

44,4 4,4 VSS tk ANA sy

434544 SATS Qa wr 7

143203 Sey

eee ey SAPS oe Passia ts sp

A eh py TESTE SEE EY

Hi4445 SSS

545454); SASS

“AIG: SATAN SASS

¥, Y YS SAMA

BAG SEES j

eo \

LLY SSS

¢. . \

Liet74 SLAM ERIS Eg,

Lave! SS RANE SS IS

Ty, WHITNEY

a fea RSAC RAE AY

ae” FM GeS, SSSR 1

es So ae LERSSSAA TN Sy :

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Meg sGG tS RAY WY Ney ty,

ett GLI4 44 SSAA 77

74, 7574, 4. RON EU E-

Se BIS SOR yy

Wa f7Z, VbiLd ASRS SSS SANE SY yy

1 7 SA

SX22

SAK

a’¢

or a

“a

i Na

iss

VIII-T

INDIA

me stenoetrus

. -

Fig. 830

we:

\ ‘

"he

7 =

Tit &

al 2G

ulge ett

£ Cf;

(Ath

0.1

VIII-T

cael

INDIA

taeniorhynchoides

——§§—_ oe

VII-S

SAN

NEN

rig. ol

Wi sg!

(\].

THAILAND

VIN-T

XK. Utsug

vexans vexans

‘

i ie

Severs tytre?

Apia st

if ‘er Misty

- a

ey {

}} SHH

Vill- T

Sonobe

JAPAN

vexans nipponil

Fig. 33

Dorsal

Micropyle

EGG OUTLINE

CHORION

RETICULATION

(enlarged from

area with arrow)

THAILAND

mediolineatus

Lateral view of aedeagus

be Mrclihul

alboscute//atus vexans vexans

TYPE TL TYPE U

Fig. 34

Horn-like

Pee Op a et

IX TERGUM

GONOSTYLUS

Filament—j

eee. |

IX STERNUM

Gonostylus

i ee is Accessory

a. Claws

0.1

Process

oy ; XC Lobe ba Paraproct

Ww ft ki-Thumb-like

Gonocoxite e a Xi > Paramere

Aedeagus

X Tergum

\ PHALLOSOME

DORSAL

THAILAND Parameral

/ Apodeme

alboscutellatus “= OEMS K UTS Ug

0.05

ceceniacs: 0: Se wearers a

———0,] ————+

THAILAND

caecus

0.l

THAILAND

culicinus

INDONESIA

low/sii

ae THAILAND

mediolineatus om.

triatus

nigros

‘©

ABIAR RNG

\ NN

\\\

INDIA 4

pallidostriatus

Fig .41

t———0.———

+——— 0] 4

Fig. 42

PHILIPPINES

pampangensis

CAMBODIA pipersalatus

pera

r———- OF. 1

i O02

PHILIPPINES

punctifemoris

itz Leg

INV G ,

LOK

) OLIARS.

/ ae 7a pet

Masks

INDIA

sfenoetrus

Fig. 46 :

ON ee al

(

no) S

Cea) ni

-—_—_—_ 0 .] —___——-

taeniorhynchoides

|. Yoshigaki

yvexans vexans

vexans nipponii ii. came

THAILAND

a/boscutel/atus

0-5

vis

by lIW

P j HIS | 14

} \

: , iil ’

Ky 0)

2 iy Hf fi pa Hii .

ni eS

caecus ;

S. Ohtawa

Fig. 51

‘e)

== f ,

SOLS

wy on

OXA OWN

pre 0S

0.5

Sonobe

THAILAND

culicinus

rid. o2

x 2// | ‘ 5

N Fa

: TA FF

THAILAND

med/iolineatus

we. Hasuanina

Fig.53

MALAYSIA

orbitae

INDIA

pallidostriatus

Fig. 99

;

; YU pH

Ay, Y yf Yj HT

LP a Cee aR

. Bg 08 1.0

AZ Hee | ae

Hf iN 7 Gi il ae Ne, Asari CNN AUS

: oF ee

N 0} oe 4’ 4 aa

Lg :

f* 45 TR

A 3.1

" V4

HN TRY. saree W\

LOAD)

TENS) Wises 3

PGta; +P uy) ay)

PHILIPPINES

pampangensis

sonobe

Fig. 56

3 14 | My

INDIA

pipersa/atus

LV)

fe)

Pig. OF

-«

0.5

LIS

Yy \\ y

WON

THAILAND

yvexans vexans

OKINAWA

vexans nipponil

Fig. 59

WS \\

~Y \

THAILAND

alboscute/l/atus

5. Oftawe

t——— 0.05 ————+

THAILAND

caecus

Lina cialas neccnas MOI aaitatcscctpcdal

0.5

variations of siphon

YAW

: vi

‘\

THAILAND

caecus 6 MA

ro a,

den TIAN

12/11

THAILAND

culicinus

iV 12

13) \ WW

ee \ i

14 10

| 12

13 9

ont \ ve

Vi 13

Vin

eee, irs ]

We. Hasan

THAILAND

mediolineatus

MALAYSIA

orbitae

A\\3\

“4

Po ge

Ze <n

a oe

ABZ

\ \\\

\a\\ \\ \

AIK \\

. \\ i

A aQQhntid

~ Liy\\

Votan

shiagatntl

0.05

\.. ai,

sal nWiwiat uauitunaganant

Aen

pallidostriatus

M.Hasunuma

“4

Fig. 66

PHILIPPINES

pampangensis

Fig.67

| iy

DTD

}

Pa hy

}

OE AMEN

enn’

cay

age

THAILAND

Ppipersa/atus

wm asunum®

THAILAND

vexans vexans

Fig.69

OKINAWA

vexans nipponii

MORPHOLOGY OF

FEMALE GENITALIA

Fig. 70

Peri-anal 0.5 Length of

ssseappiceuee 9 Dorsal PGL

Dorsal —Dorsal PGL Length

PGL Width Ventral PGL Length

Postgenital—— — Line of attachment of

Lobe(PGL) Peri-anal Membrane to

B f Dorsal Surface of PGL

ase 0

Spermathecal Upper Vaginal Lip

Duct Sternum VIII

Upper Vaginal Sclerite VENTRAL

—Lower Vaginal Lip

Tuberculus

VENTRAL

—0.5 Length of Cercus Poe. Capsule

LATERAL

Cercus Cercus Length

— Cercus Width

DORSAL

: Tergum VIII

a BE! Width

Tergum IX

Tergum IX Length

Width—-

Tergum VIII

Tergum IX DORSAL

DORSAL

ay bral

Reinert: Aedes (Aedimorphus) in Southeast Asia 215

INDEX

Valid names are set in roman type, synonyms are italicized. The

italicized pages are those which begin the primary treatment of the taxon.

Numbers in parenthesis refer to the figures illustrating the species in

question.

abnormalis 5, 13, 138

abnormalis kabwachensis 138

adami 138

Aedes 1, 2, 4,9-14, 17, 18, 22-24,

28, 29, 32-35, 37, 38, 42,

43, 45, 48-50, 53, 54,

07-59, 63, 64, 66, 67, 70,

76, 79, 81, 83 - 86, 111,

113, 116, 118, 121, 123, 126,

128, 131, £33, 136- 138, 142,

143

Aedimorphus 1-3, 4, 9-11, 13, 14,

11, 18, 24, 297 S335, 37,

38, 42, 43, 45, 48, 50, 54,

57, 59, 63) G4, 66, 67) 76,

79, 83, 84, 86, 111, 113,

116, 118, 121, 123, 126, 128,

131, 133, 136- 138, 142, 143

africanus 5

albocephalus 5, 13, 138

alboscutellatus 1, 2, 4, 6, 13, 16,

£7, 189 2422520 32,-33,

37, 42, 48, 54, 58, 63, 77,

111, 136, 137, 142, 143 (1,

16, 17, 18, 33, 34, 49, 59)

alboscutellatus group 10

alboventralis 138

andamensis 29, 59

annulipes, Anopheles 77

annulus, Culex 23, 33, 42

Anopheles 23, 29, 33, 42, 49, 59,

co ae a |

apicoannulatus 12, 138

avabiensis 70

argenteopunctatus 13, 138

argenteoscutellatus 2, 136

argentinotus 17, 21

articulatus 70

Ayurakitia 10

bailyi, Culex 23

balabacensis, Anopheles 23, 29, 49

bancrofti, Wuchereria 12, 66

barbirostris, Anopheles 42, 49

Bathosomyta 5

bedfordi 138

bengalensis, Anopheles 23

bevisi 138

bicolor, Uranotaenia 29, 49, 77

bitaeniorhynchus, Culex 29, 49, 83

boneti 138

boneti kumbae 138

caecus 1, 2, 5, 6, 12-14, 16, 17,

23, 24, 27, 28, 33, 42, 49,

q® >, 11, 83-188, 6°53,

142, 143 (2, 16, 17, 19, 35,

50, 60, 61)

caliginosus 138

capensis 138

Catageiomyia 4

centropunctatus 138

Chaetocruiomyia 10

chamboni 138

confusus 5

congolensis 138

Culex 4, 5, 10, 17, 21, 23, 2%:29.

33, 38, 41, 42, 48, 49, 50,

03, 66, 70, 71, 77, 83, 84

Culicada 66, 70, 79, 82

culicifacies, Anopheles 23

culicinus: 1,*2,. 5,6) 107 36c E82,

23, 28, 29, 32, 33, 42, 59, 63,

116, 136, 142, 143 (3, 16, 17,

20) 365 Sis 62)

cumminsii 13, 138

dalzieli 139

davidi 13, 15, 16, 33, 34, 76, 136

dentatus 13, 139

dialloi 139

Diceromyia 10

domesticus 2, 4, 9, 139

dorsalis 83

durbanensis 139

Duttonia 5

ebogoensis 139

Ecculex 2, 4, 17, 24, 29, 32, 35,

38, 45, 50, 54, 59, 66, 79

216

Edwardsaedes 11, 28

eritreae 139

eritreae karooensis 139

eruthrosops 66, 70

euochrus T0

falabreguesi 139

farauti, Anopheles 77

ferinus 23, 29, 33, 42

filicis 139

Finlaya 10, 63, 76

fisheri 2

fowleri 13, 139

fraudatrix group, Culex 77

fryeri 13

fuscanus, Culex 42, 77

fuscocephalus, Culex 23, 29, 33,

Geitonomyia 2, 5, 24

gelidus, Culex 42, 77

gibbinsi 139

gilliesi 139

gouldi 2, 136

erenieri 139

erjebinei 139

gubernatoris 29, 59

hackeri, Anopheles 49

hamoni 139

haworthi 139

hayashii, Culex 83

hirsutus 139

holocinctus 139

hopkinsi 139

immitis, Dirofilaria 12

imprimens 23, 28, 29, 33, 59, 63

indicus 29

indiensis, Anopheles 29

infantulus, Culex 23

insolens 139

introlatus, Anopheles 49

irritans 139

jamesi 2, 6, 22, 37, 49, 136

kapretwae 139

kennethi 139

kingii 5

kochi, Anopheles 23, 29, 33, 42,

49, 59, 63, 77

Contrib. Amer. Ent. Inst.,

vol. 9, no. 5, 1973

lamborni 139

leesoni 13, 140

leesoni verna 140

Lepidotomyia 2, 4, 17, 21, 45, 48

leptolabis 140

Leslieomyia 2, 5, 85

leucarthrius 140

Levua 11

lineatopennis 10, 11, 42, 77

lokojoensis 140

longiseta 140

lottei 140

lowisii 1-3, 6, 14-16, 22, 35, 37,

49, 136, 137, 142 (4, 16,

17, 21, 37)

macfarlanei, Aedes 49

macfarlanei, Uranotaenia 23

maculata 2, 4, 5, 59, 62, 85

maculatus, Anopheles 23, 29, 33,

42, 49

malariae 70

malayi, Brugia 13

malayi, Culex 23

mansouri 140

marshalli 13, 140

mattinglyi 140

mediolineatus 1, 2, 5, 6, 9, 12,

14-17, 23, 29, 33, 38, 41,

42;. 53,. 58, ‘17 TTS, 136,

142, 143 (5, 16, 17, 22,

33; (38, £525°63)

micaculosus, Culex 77

microstictus 140

Mimeteculex 5

mimulus, Culex 29, 42, 49, 77

mindoroensis 1, 2, 35, 37

minuta 66, 70

minutus 13, 140

mixtus 140

montanus, Anopheles 49

mutilus 140

Myxosquamus 5

natronius 140

neobiannulatus 140

Neomacleaya 11

Neomelaniconion 10, 11

Neopecomyia 5

ngong 140

nigricephalus 140

nigropunctatus, Culex 23, 29

Reinert: Aedes (Aedimorphus) in Southeast Asia 217

nigrostriatus 1-3, 14-16, 43, 45,

136, .142.(6, -16, :23,..39)

niveoscutellum 1, 2, 54, 57, 58

nocturnus 1, 2, 66, 67, 71, 77

nocturnus var. niger 66, 70

notoscriptus 76

nummatus 10

nyounae 140

oakleyi 2, 6, 22, 137

Ochlerotatus 2, 10, 11, 13, 17, 24,

35, 38, 45, 50, 54, 59, 66,

16, . 79,: 85

ochraceus 50, 140

omurensis 17, 21

orbitae 1-3, 6, 14-17, 28, 29, 45,

48, 49, 121, 136, 142, 143

(7, 16, 17, 24, 40, 53, 64)

orientalis, Culex 83

ovazzai 140

pachyurus 140

pallidostriatus 1-3, 5, 6, 13-17, 33,

49. 50, 53, 58, 68, 128, 196

142, 143 (8, 16, 17, 25, 41,

54, 65)

pallidothorax, Culex 23, 29

pampangensis 1-4, 6, 14, 16, 17,

o4, 57, 58, 126, 136, 142,

143(9, 16, 17,26, 42, -55;

66)

parascelos 50, 53

parvus 71

Pecomyia 2, 4, 23, 85

philippinensis, Anopheles 29, 42

phyllolabis 140

pipersalatus 1, 2, 13-17, 22, 37,

59, 62, 63, 85, 128, 136, 142,

143 {10, 16,17, 27, 43556:

67)

pipiens, Culex 83

Polyleptiomyia 5

Pseudograbhamia. 2, 5, 59, 62

pseudostenoetrus 84

pseudotarsalis 140

pseudovishnui, Culex 23, 29, 42

Psorophora 10, 11

pubescens 140

pullus, Culex 77

punctifemoris 1, 2, 5, 10, 14, 16

49, 63, 64-66, 136, 142,

143 (11, 16, 17, 28, 44)

punctothoracis 141

quasiunivittatus 141

quinquefasciatus, Culex 77

raptor, Culex 23, 29, 33, 42

reali 141

Reedomyia 2, 4, 17, 35, 37, 54, 57

rickenbachi 141

riparis, Anopheles 49

scanloni, Culex 23, 29

semlikiensis 141

senegalensis 4

senyavinensis 2, 6, 22, 137

seychellensis 141

simulans 141

Sinensis, Culex 29

sitiens, Culex 77

smithburni 141

Stegomyia 5, 10, 11, 59, 62, 63,

65, 84

stenoetrus 1, 2, 5-7, 14, 15, 76,

84, 85, 136, 142, 143 (12,

16, 17, 29, 45)

Stenoscutus 5

stokesi 12, 141

subdentatus 141

subpictus, Anopheles 29, 49, 77

Suknaensis 28

sylvestris 2, 4, T1

syntheticus 2, 5, 7, 11, 76, 136

taeniata 45, 48

taeniorhynchoides 1, 2, 5, 14, 15,

63, 85, 86, 136, 142, 143

(13, 16, 17, 30, 46)

tarsalis 5, 13, 141

tauffliebi 141

teesdalei 141

tessellatus, Anopheles 77

tiptoni 141

tricholabis bwamba 141

trilineatus 38, 41, 42

trimaculatus 2, 45, 136

tritaeniorhynchus, Culex 42, 77, 83

trukensis 2, 6, 22, 137

Udaya 10

uncus 23

uniannulata 5

Uranotaenia 4, 23, 29, 49, 77

vagus, Anopheles 29, 42, 77

Verrallina 11

218 Contrib. Amer. Ent. Inst., vol. 9, no. 5, 1973

vexans 1, 2, 5; 6, 9-11, 13, 28, 66, vigilax 23, 76, 85, 86

67, 70, 75-79 vittatus 11, 23, 29, 33, 59

vexans group 5, 7, 11, 76

vexans nipponii 5-7, 9, 11, 12, wainwrighti 5, 10, 14, 16, 83,

14-17, 63, 76, 79, 82, 83, 84, 136

133, 137, 142, 143 (15, 16, wendyae 141

17, 32, 48, 58, 69) whitmorei, Culex 42

vexans nocturnus 34 wigglesworthi 141

vexans var. nipponii 79,

vexans vexans 5-7, 9, 12, 14-17, yangambiensis 141

23, 28, 29, 34, 42, 63, 66, yvoneae 141

75-78, 81, 83, 85, 86, 131,

136-138, 141-143 (14, 16,

17, 31, 33, 47, °57; 68)

Some specimens of orbitae in the type series, located in the British

Museum (Natural History), bear labels with the following two unpublished

names;

alboscutellata var. annul 48

trifeliat 49

3 reese

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setae fpatetateseteteincatgianyaieencatenstirtonnersqinctatinat gett tratote aatatatatersee aaa eater areata

, cies MPSCTE Se St VEIL TE Te Te oe ses é

ont

ef oseebcee tented tte

Geese auesoaseaaeers sy eeenceaea

rt siatatabnee intel

(eiotals eelele le:

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siejeisiainieisimisisiabeisbeleie tities aiatsleiaeleininiel: .

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cies ea (hm ERS hark teh Be heb ab eae oir sede eters rz ees ray cin imi he!

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oli behear ereeunre see aaereyeeoupee reer Heder es eget 4 eersr ces + | . Like AGRI DIR he ee Pari eeneen ae yeee esac apaw se tereases rear acse teqese sees ye eer yy yepees gees yeey verry’

* i “ cyeerreye oaetearerd expt essa eenary reeds Pe be test ms : evetr es: tess te. Ts) x Bie | Bk a (Simi Si ei nies, ea eerele ela eee vere we seee yete te ey tere eiede ay te se se ye ye TENE

dite! i bree ihe hs OS dap aee mb Kas Biea ppaels ee aes : a et (kim ei si aia aeyl pat + ba ee) Parent tet Atha; boas 5s bn hots ee Rea ae Te eet agen eS dasa Hay eeeeyr Vtstt es

Se OR est Pe PES eS ear rovegtoaar gitar ear as year te Sr pl mis %I ; Seas erate aeeperate rye geet * 4 Oye oh 4 severe sear ye isiais ti TEot om ni eish ee ratetersiels) ; pees

ds Ry Fiat Gln ate Bike Ki eh ha a PS ee ee ee a ee ee ee | Hwee 26 ge eae whee bah a ahahnbete tS)

Beh eb Os eee ee! DS