PREFACE

This work is a synthesis of the existing data on the taxonomy, biology and

medical significance of the subgenus Anopheles in Thailand. It is primarily

intended for medical entomologists, malariologists and others engaged in public

health work in Thailand. Investigators in other Southeast Asian countries may

also find this study of value, but should remember the taxonomic descriptions,

discussions and keys are intended for use only in Thailand.

The Anopheles of Thailand was one of the first studies undertaken by the

Southeast Asia Mosquito Project (SEAMP)*, Smithsonian Institution, but its

completion has been delayed for a number of years. In the meantime, an ex-

tremely important work on the anopheline mosquitoes of Malaya and Borneo ap-

peared in 1968. That work represents the culmination of studies by Dr. John A.

Reid, begun in 1939 at the Institute for Medical Research in Malaysia. Many of

the species known from Thailand also occur in Malaysia, and wherever possible

we have drawn on Dr. Reid's work.

The primary source of specimens for this study derives from collections

begun in Thailand in 1961 by personnel of the Department of Medical Entomology,

U. S. Army Medical Component - SEATO, Bangkok, and continuing to the pres-

ent. These specimens are housed in the Medical Entomology Project (MEP) at

the Smithsonian Institution, Washington, D. C. The MEP collections are among

the most extensive ever made in Southeast Asia or the Oriental Region as a

whole, permitting extensive evaluation of inter- and intraspecific variation. Un-

fortunately, the Southeast Asian collection coverage is uneven, with excellent

collections available only from Malaysia, Philippines and Thailand. The short-

age of good specimens from the other Southeast Asian countries was alleviated

by loans from many United States and European entomological depositories.

During this study the primary means of determining taxonomic characters of

value was the examination of reared adults with associated immature skins. In-

dividual specimens, larval or adult, were not afforded this degree of credibility.

However, since specimens collected under field conditions are nearly always

limited to one stage, we attempted to make the keys precise, yet simple. To

assist workers in the field, we have included a simplified key (Appendix A) to

the females of all the known Thai anophelines. Unfortunately, the taxonomy of

some of the species complexes is so complicated that only specimens with 2 or

more associated life stages available for examination can be identified to species.

Hopefully, future workers can find additional characters that will make the identi-

fication of these species easier.

*On 1 June 1974, SEAMP was succeeded by the Medical Entomology Project

(MEP), Department of Entomology, at the Smithsonian Institution.

CONTENTS

ABSTRACT eas ee ee ee 1

INTRODUCTION 363 os es ee es a eet a ee 2

Backeoround: 0c seg Be a ie ee a a 2

AGOSCOLTADNY be ee ae a oe ee er i ey 3

FPormat-and Treatment.) i ere eS eee ae SS 8

Morpholosy 7s. 0S a ee eS ei fae eee ee, a 9

EAXONONVY: 3,2.) Guchowlret ee ie die ar nate re as ol ge er 21

GENUS ANOPHE LES WIENGEN., 2 6. bs eee eee ee 21

Key to Subgenera of Awopieles mn Thailand occ et. oe eee be 29

Subgenus Anopheles Meigen. ....... ae ae eee al 25

Keys to Sections, Series and Species Groups. .........e6..-. 26

LATICORN SECTION Reid and Knioht 9... 35. wet 6 we ees ee 29

MYZ ORAYNCH US SERINS bdwards: iu) «wial ook wicere 2 ok ee 29

Key to males of species groups of the Myzorhynchus series in

TRAMANG fone diene is eee Dah eee 2 ee 30

CLDOIGENIOTUS SpeCies Croup. 6 oy Po OS ee, ar ee 31

montanus Stanton and Hacker... 40) a ee oe ae 34

hyrcanus species group of Southeast Asia ............ 36

Keys to species of the hyrcanus group in Thailand. ..... 40

SUICHS 1S WIECOMANN Gooey pe ee a ae 45

aveyropus: (Swellensvebel): 23. 0a eS ee ea 50

CVOUPOVOE. REINS ORO ee ee ee O4

nitidus Harrison, Scanlon and Reid. 2... er oe eS 08

Lester pavalige pandoshaMm. aii Be a ee

WAREV TINTUS GIGS oe Ea a OE ee Pa cee cake te 65

pediiaenmius (Leicester) 2080 6 OP eS ee. 71

PUVSOL COVE RAINS e SON e a U5

DEAVOIVOSIVIS SUOCIOCS CROW.) PA OO a 78

Keys to species of the barbirostris group in Thailand .... 81

bavoTFOshias: Yaw der Wulp ee Or Be AE a 83

COV NO OIG Co re eR BG ae ae 89

TONGIOT ROT PME RBA as es 94

ROG LRT TRE i OI PRO Ee ES ee 97

POUTCAVES CIP ge EOE eee ee ee Sh i Oe ee 100

barbumbrosus Strickland and Chowdhury ......... 102

UUOVOSUS: GOCCICR STON Coo ee oe ae eee a ew 105

Keys to species of the umbrosus group in Thailand. ..... 108

WINDTOSUS (TMEGRAIA isle wo be eek ek a Pad 110

OE er tea Oa Ey Ma a a geil gang 114

betiFey BANCO eA i Pa are eo Se 118

WTOP EOE TRO ee aia Ee eg yang a Olga Grog a. ae Na ae 122

TORCH THOU igi esuesie alia fale oe Sve veh COR: ee eee 124

Seba mains: (HOC CSteR ee aes ace ul Pe nak cam Se 128

ANGUSTICORN SECTION Reid and Rieti ics ek bee aw ee eT 132

LOPAH OSCE LOM VIA SERIES HOWaede Oe we aN 8 ae A wk ee ee 132

Keys to species of the Lophoscelomyia series in Thailand. .... 134

OUR ON Ce ie yg gnkas Gi, tee na 134

ASICLICUS GPOCIOG, SVOU ye lig: Soa athe ye ca ae Re Eat hte 135

ESIQITOUS LiClO Sete Fr ee ae aang a RE er Sian Tee ame Rs 136

errs Pun ae ee ee. HUGS Maree, MeN tel ci ges 140

ANOPTH Te SU WA Oe ae ee ae eae Oe! eS a el we 144

Keys to species groups and species of the Anopheles series in

TiaRTaN oe ie eee erick! Kaas hie dye a IIR TRC ERA We a a 145

CUNEO SNOCICS CTOUNY a Wi ia a ee OR RK eee oe 147

STs ae ie ee ae Gene | a a 152

i a im ate a Me gy Mae Bre 157

eee UP MODAN ea Se i SSA Pe as 162

Poy Scan and Peyton. 2. fe ee a oe 64

insulaeflorum Swellengrebel and Swellengrebel-de Graaf. 167

Eo IES OD TSE (6 MUNA SEIN NEU le at elie EP aa hie a a an ee 171

palmatus (Rodenwaldt) ........ SA Hah OU GINS sink ae a 173

CUNEWOTINS SPCCIES SOUP Cs a ee 176

Ry ONG Wen Sis Aramis se Ee ee ae 179

OE ORO IACS AO eT are re yr eee ee 181

APPENDIX A - Key to the female Anopheles of Thailand - an aid for rapid

identification in malaria field research. ......... ERE RAR «ay 186

APPENDIX B - Southeast Asian species of the subgenus Anopheles not

POW POM Pe ELAN se Se SU i OR ae es 192

Pee GCS GE nb 2020 oo IN Sa RU ale a 193

DRA ee a Se ee So Re eee See ee 195

FRG ee ee ee Cae tray OR eae mre eg gee Mary lag Sow ue oe 212

MEDICAL ENTOMOLOGY STUDIES - II

THE SUBGENUS ANOPHELES IN

‘THAILAND

(Diptera: Culicidae)!

Bruce A. Harrison? and John E. Scanlon?

ABSTRACT

This is a comprehensive revision of the subgenus Anopheles in Thailand

that involved the examination of over 10,000 specimens of 32 species and one

subspecies. Included in the 96 full-page figures are: 92 plates of whole pupae

and larvae, many habitus drawings of females, male genitalia, wings and other

selected characters. Major sections included are: zoogeography, format-

treatment, morphology, taxonomy in general, taxonomy of supraspecific cate-

gories, keys and the descriptions. Species descriptions include sections on:

synonymy, descriptions of female, male, pupa and larva, type-data, distribu-

tion, taxonomic discussion, biology and where applicable, medical significance.

Special emphasis is placed on the keys, descriptions, type discussions, intra-

specific and interspecific variations, affinities, behavior and a specific level

review of medical significance. A key to all the known adult females of the

genus Anopheles in Thailand is included as an appendix.

The male, female, pupa and larva of A. aberrans n. sp., are described

and illustrated. Pupae of insulaeflorum and palmatus are described for the

first time. Stages and genitalia for the following species are illustrated for the

first time: campestris pupa; donaldi pupa; hodgkini male genitalia; insulae-

florum male genitalia; palmatus adult abdomen, male genitalia and pupa: and

pollicaris male genitalia and pupa. A lectotype is designated for baezai, and

Stethomyia pallida Ludlow, is moved from synonymy under aitkenii to synonymy

under bengalensis.

IThis 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. This publication is contribution number

1334 from the Army Research Program on Malaria.

2 Major, MSC, U. S. Army, Department of Entomology, Walter Reed Army

Institute of Research, Washington, D. C. 20012.

3School of Public Health, University of Texas, Health Science Center at

Houston, P. O. Box 20186, Houston, Texas 77025.

2 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

INTRODUCTION

Background

This publication is the result of studies undertaken in Thailand by the senior

author during the period 1967-1970 and the junior author during the period 1961-

1964, under the auspices of the Southeast Asia Treaty Organization (SEATO)

Medical Research Laboratory. Since 1964, this work has been an integral part

of the Southeast Asia Mosquito Project (SEAMP), Department of Entomology,

Smithsonian Institution. |

The anopheline mosquito fauna of Thailand is large and complex, and in

recent years it has become apparent that there are complex species groups in

the fauna, which defy taxonomic clarification by classical morphological means

alone. Still, it has required the application of these classical techniques, coupled

with observations on the habits and distribution of the morphologically distin-

cuishable taxa to place us, at least, in the position where definition of the spe-

cies in complexes by cross-breeding, cytogenetic, or other methods can pro-

ceed on a sound basis. At the time the present study was undertaken the most

recent faunal checklist for the mosquitoes of Thailand (Thurman 1963) con-

tained 21 species and subspecies in the subgenus Anopheles. During the course

of our subsequent studies it seemed advisable to prepare an interim checklist

(Scanlon, Peyton and Gould 1968) in which there were 29 species of the subgenus

Anopheles, with an additional 4 species listed as doubtful records. In a subse-

quent work by Reid (1968), an additional species of this subgenus was reported

from Thailand. The present work recognizes an additional 3 species in Thailand,

and thus, deals with a total of 33 species. It is quite possible that new species

will be added when the more heavily forested areas and some of the more cryptic

habitats (Peyton 1970) have been further sampled. Additional new records

should occur when the fauna of the presently inaccessible border areas is better

known.

The great bulk of the specimens examined in the course of this work resulted

from the collections of the U. S. Army Medical Component, SEATO, made

throughout Thailand from 1961 - 1973. These collections were usually made in

connection with specific disease problems, but whenever possible specimens were

reared and biological data taken on all species, regardless of immediate medical

importance. Consequently, a major portion of the mosquito fauna of Thailand

has been collected, and the majority of specimens discussed herein are reared

adults with associated immature skins. These specimens have been deposited

in the MEP collection. The National Museum of Natural History (USNM) also

possesses a considerable number of Thailand Anopheies deposited by earlier

workers (D. C. and E. B. Thurman, O. R. Causey, E. I. Coher and P. F.

Beales). In addition, USNM collections were examined for specimens of the

subgenus Anopheles from areas outside Thailand which might shed light on the

range of variation for each of the species considered. Specimens were exam-

ined from other collections listed in the acknowledgements and in appropriate

portions of the text. The specimens in the MEP collections will be deposited

in the regular collections of the USNM.

The earliest publication containing references to Thailand anophelines was

that of Theobald (1910), who reported barbirostris and vossii Giles (= vagus

Donitz). This was followed by Stanton's (1920) study of the mosquito fauna of

various far eastern ports which included records of Bangkok mosquito species.

Stanton's primary interest was evaluating the danger of yellow fever epidemics

following the opening of the Panama Canal. The first papers dealing specifically

Harrison and Scanlon: Subgenus Anopheles in Thailand 3

with the genus Anopheles and the role of anopheline species in the transmission

of malaria in Thailand were those of Barnes (1923 a,b) who listed some 17 spe-

cies (only 2 in the subgenus Anopheles) with notes on their biology and vector

relationships. Barnes' records were repeated by Barraud and Christophers

(1931) as were those of the other early workers cited above. In addition, these

authors discussed a collection made by J. A. Sinton along the railroad line from

Bangkok to Chiangmai, and in those 2 cities. They listed 20 Anopheles for the

country, one of which they regarded as a doubtful record. Additional early re-

cords of anophelines in Thailand were published by Anigstein (1932); Christo-

phers (1933); Covell (1927); Payung-Vejjasastra (1933) and Causey (1937 a,b).

The last named provided a rather complete analysis of the anopheline fauna

then known, particularly in relation to the terrain and hydrography of the country.

Causey's anopheline data were, however, largely based on his Bangkok collec-

tions. For areas outside Bangkok, Causey noted the more extensive work of

Anigstein, who also provided an excellent account of the terrain and hydrography

of Thailand as related to the breeding of Anopheles. These relatively meager

observations were being made at a time when very significant observations were

being published on the epidemiology of malaria and on the taxonomy and biology

of anophelines by British workers in India and Malaya, and the French in Indo-

china.

During the period of World War II and shortly thereafter a number of short

papers were published on the malaria encountered in prisoner of war camps in

Thailand (Wilson and Reid 1949; De Fluiter 1948), which included records of

Anopheles species. In the postwar era interest in mosquitoes in general, includ-

ing the anophelines, expanded rapidly and lists of species and biological data

were published by Bonne-Wepster and Swellengrebel (1953); Foote and Cook (1959);

Puri (1949); Iyengar (1953); and Thurman (1959). The last publication summar-

ized to some extent the large scale collecting efforts of D. C. and E. B. Thurman

in northern Thailand. Their efforts were almost entirely devoted to culicine spe-

cies. The long series of papers by Reid, culminating in his monograph on the

Malayan and Borneo Anopheles (1968), contain many records of Thailand Anophel-

€s species. Inthe course of studying the Thai anophelines numerous biological

and epidemiological observations were made, and thousands of specimens have

been examined in the field and from many museum and other collections. Con-

sequently, a number of publications have appeared which are most pertinent to

malaria epidemiology and the anopheline fauna of Thailand. These are: Gould,

Esah and Pranith (1965); Gould, Scanlon and Ward (1966); Harrison (1972, 1973);

Harrison and Scanlon (1974); Harrison, Scanlon and Reid (1973); Peyton and

Scanlon (1966); Rattanarithikul and Harrison (1973); Scanlon and Esah (1965);

Scanlon and Peyton (1967); Scanlon, Peyton and Gould (1967, 1968) and Scanlon

and Sandhinand (1965).

Zoogeography

The Kingdom of Thailand lies between approximately 6 degrees and 21 degrees

north latitude, and extends roughly from 98 degrees to 105 degrees east latitude.

The Kingdom is irregularly shaped, somewhat in the shape of an elephant's head,

with the trunk forming the portion extending southward to the Malaysian border.

The country is divided into 71 Provinces or Changwats, forming the first order

administrative units. The list of Province names employed here (Fig. 1) and in

earlier publications in this series conform to the Official Standard Names Gaze-

teer No. 97 of the U. S. Board of Geographic Names, Washington, D. C. A few

of these names differ slightly in orthography from the standard Romanized names

proclaimed in the Royal Gazette of the Kingdom of Thailand, number 84, 1967,

4. Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

which were employed in Scanlon et al. (1968). The differences are so slight

that no confusion should result from using that publication in conjunction with

the present work.

Much of the population of Thailand, and most of the industry, is located on

the broad central alluvial plain near Bangkok (= Krung Thep). The central fea-

ture of this plain is the Chao Phrya River (= Maenam Chao Phrya) and its num-

erous branches and tributaries. This is one of the richest rice producing areas

in the world, and an aerial view of the region conveys the impression of a vast

plain compounded of varying proportions of earth and water. Until fairly recently

most of the transportation depended on the countless canals and natural water-

ways. During the rainy season mosquito production in the surface waters is

immense, and breeding continues to some extent throughout the year.

Southward and westward from Bangkok the country narrows rapidly to form

the Isthmus of Kra, separated from Burma in the northern portion by the Bilauk

Taung mountain range (1,097 - 1,158 m). The more southern portion of the pen-

insula consists of coastal plains on the east and west coasts of varying widths,

with central ranges of low hills. Toward the southern border tropical deciduous.

forest gives way to wet and rain forests. There are rather extensive stretches

of mangrove on the west coast and much smaller groves on the Gulf of Thailand

Side.

West of Bangkok there is a moderately high range of mountains forming the

border with Burma and running northward to northwestern Thailand. The entire

northwestern portion of the country is mountainous, with intervening fertile val-

leys. These mountains are relatively low and covered for the most part by tropi-

cal deciduous forest, with teak (Tectona grandis Linnaeus) as one of the domin-

ant large trees. Only a few of the tallest mountains (Doi Inthanon, 2,599 m;

Doi Pui 1, 683 m) carry a pine-oak forest cover at the higher altitudes. The

various ranges of hills and mountains in this northwestern part of the country

are separated by various tributaries which flow southward to form the Chao

Phrya River. Most lines of commerce and communication follow these valleys

and streams and move in a north-south direction. The mountains are high enough

to show some effect on mosquito species distribution (Scanlon and Esah 1965) but

no real barrier exists to movement northward into Burma and China, as evi-

denced by the list of species of Anopheles shared with those areas, given below.

Southeast of Bangkok there is a coastal plain with some mangrove areas,

with inland hills and rather isolated mountains which approach the coast at

points, particularly near the Cambodian border in Chanthaburi and Trat pro-

vinces (Fig. 1). These provinces have extensive areas of tropical wet forest,

in many ways resembling the southern peninsular portion of the country near the

Malaysian border.

One of the most striking geographical features of the country is northeast of

Bangkok and called the Korat Plateau. This plateau is bordered on the south by

a line of hills and mountains in a general east-west direction. To the northeast

is a vast area of cultivated land and deciduous forest that forms part of the drain-

age basin of the Mekong River. The plateau is generally drier than the rest of

Thailand, except for the southern slopes of the mountain chain which have areas

of evergreen cloud forest. The highest peak along the chain is Khao Laem

(1,341 m) in Nakhon Nayok Province. From about that point the fringing moun-

tain chain turns northward and becomes the complex, but rather low, Dong Phraya

Yen range to the border of Laos.

From the forgoing account it can be seen that there is considerable diversity

to the geography of Thailand. This is compounded to some extent by the climatic

pattern. Most of the country receives a single monsoon, with the rainy season

BURMA

24)!

er PV At hoe

63) a peaks; 4

J TPs \ (23)

ANDAMAN (64) as

548 ! 7

at: ane s a.

Q 66

; aS

, Q

YD MALAYSIA

e e

SO ON nw FWD PF

55.

CAMBODIA

CHANGWATS OF THAILAND

Ang Thong

Ayutthaya

Buriram

Chachoengsao

Chainat

Chaiyaphum

. Chanthaburi

Chiang Mai

Chiang Rai

Chon Burt

Chumphon

Kalasin

Kamphaeng Phet

Kanchanaburi

Khon Kaen

Krabi

Lampang

Lamphun

Loei

Lop Buri

Mae Hong Son

Maha Sarakham

Nakhon Nayok

Nakhon Pathom

Nakhon Phanom

- Nakhon Ratchasima

Nakhon

Nan

Narathiwat

Nong Khai

Nonthaburi

Pathum Thani

Pattani

Phangnga

Sawan

Si Thammarat

36,

38.

39.

oO,

SEN

42,

U3,

yy,

45,

U7.

48,

ho,

50.

51.

52.

ape

Phatthalung

Phetchabun

Phet Buri

Phichit

Phitsanulok

Phra Nakhon

Phrae

Phuket

Prachin Buri

Prachuap Khiri Khan

Ranong

Rat Buri

Rayong

Roi Et

Sakon Nakhon

Samut Prakan

Samut Sakhon

Samut Songkhram

Sara Buri

Satun

Sing Buri

Songkhla

- sisaket

Sukhothai

- suphan Buri

Surat Thani

Pipe ea iat

Tak

Thon Buri

Trang

Trat

- Ubon Ratchathani

Udon Thani

Uthai Thani

Uttaradit

Yala

6 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

extending from May to November, which creates a tropical savannah climate

(van Randwijk and Willems 1964). The extreme southeastern mountains and

much of the peninsular portion of the country receive much higher amounts of

rainfall, and portions of the peninsula receive rain from the northeast and the

southwest monsoons, creating a rain forest climate.

Although many gaps remain in the distributions of the species of the sub-

genus known in Thailand (Table 1), an examination of their presently known

distributions reveals some interesting patterns. It appears that there are rela-

tively few if any, terrain barriers to movement on the north-south axis. Many

of the species encountered in Thailand are found in much of Southeast Asial,

and it is difficult to make any generalizations concerning these, except on an

individual basis. As might be expected, some of these species show a consider-

able amount of variation over their very wide ranges (e.g. nigerrimus, page

68). These variations, where applicable, are discussed under each species.

Allowing for this individual variation, it appears that the Thailand species

of the subgenus Anopheles share a great many characters with those in neigh-

boring areas. There are many species in Peninsular Malaysia and Borneo

that do not enter Thailand, but the fauna of South Thailand bears a strong

affinity to that of Malaysia. There also appears to be relatively few differ-

ences from the Indonesian forms, at least for the Greater Sundas. Inadequate

numbers of specimens were available from other parts of Indonesia to permit

much speculation. The situation with the Philippines is quite different. While

many of the Philippine species are obviously closely related to members of

the same species groups in Thailand (nigerrimus - pseudosinensis; barbirostris -

franciscoi; barbumbrosus - vanus), the overall impression is one of 2 quite

distinct faunas.

Reid (1950, 1968) commented on the distribution of a number of species of

the subgenus Cellia which occur in the northern part of Peninsular Malaysia

generally near the Thai border, but disappear slightly further south, only to

occur again on various islands of the Malaysian and Indonesian chains. Similar

discontinuities appear to be found in reptiles, butterflies and other groups

(Reid 1950). Although such discontinuities are not apparent among the members

of the subgenus Anopheles, there are a number of species in Groups I and III

(e.g. pursati, avgyropus, sinensis and Sintonoides) which are widespread on

the mainland, but which disappear on the Malayan peninsula or the island

chains. The reverse is true of a number of species in Group II (ex. donaldi,

montanus, pollicaris and stricklandi) that occur in the Malaysian region, ex-

tending north into southern Thailand and then disappearing. These trends are

not balanced, for there appears to be more penetration of the southern area by

the species of Group I than northward penetration by the members of Group II.

Nevertheless, these trends support the distinct anopheline faunal change which

occurs in the vicinity of the Thai-Malaysian border. In the absence of topogra-

phic barriers this phenomenon is probably due to climatic factors, in particular

the seasonal rainfall patterns which cause the change from a tropical monsoon

to a wet forest type vegetation. There seems to be a definite correlation be-

tween the forest vegetation-rainfall pattern and the Malaysian centered mem-

bers of the subgenus, thus, one would expect to find some Group II species in

lAs defined by de Meillon (1969) this includes: Bangladesh, Andaman Islands,

Burma, Thailand, Cambodia, Laos, North and South Vietnam, Malaysia,

Singapore, China south of the Yangtze River, Taiwan, Southern Ryukyu

Islands, Philippines and Indonesia. Also included are Assam and Brunei,

while Indonesia is restricted to that area west of Weber's zoogeographical line.

Harrison and Scanlon: Subgenus Anopheles in Thailand %

TABLE 1. Geographical affinities of species of

the subgenus Anopheles in Thailand

Group I - Species with a wide distribution in Southeast Asia

argyvopus

baezai

barbirostris

barbumbrosus

bengalensis

insulaeflorum

nNIZeEvYVIMUS

nitidus

peditaeniatus

SinENSIS

Group II - Species which occur in Thailand, but appear to have their

center of distribution in Malaysia and Indonesia.

asiaticus

campestris

crawfordi

donaldi —

fragilis

hodgkini

lesteri paraliae

letifer

montanus

palmatus

pollicaris

voperi

separatus

stricklandi

umbrosus

whartoni

Group III - Species which apparently have a wide distribution on mainland South-

east Asia, but disappear in peninsular Thailand or Malaysia.

interruptus

pursati

sintonoides

Group IV - Infrequently collected species known only from

Thailand, or Thailand and adjacent countries.

aberrans - (Thailand)

bulkleyi - (Thailand)

kyondawensis - (Burma and Thailand)

tigertti - (Thailand)

8 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

the southeastern Thailand provinces of Chanthaburi and Trat, where the rain-

fall pattern is much like that of the southern provinces. This does in fact occur,

for such species as baezai, hodgkini, voperi and whartoni have been collected in

that area; however, additional collections are needed from that area of Thailand

to confirm this pattern. Further support for the correlation of the wet forest

type vegetation with the Group II species is found in peninsular Thailand. The

Group II species in southern Thailand generally extend up further into Thailand

on the west side of the peninsula, than on the east side. In fact, 7 members of

Group II (baezai, hodgkini, montanus, palmatus, roperi, sintonoides and

stricklandi) have been collected in Ranong, the most northern Thailand province

on the west side of the peninsula. Ranong has large tracts of tropical wet for-

est and records the highest annual precipitation of any province in Thailand. _

In addition, this province is adjacent to the southernmost extension of Burma,

suggesting the possible extension of some of the Group II species into that

country.

At present, no satisfactory explanation is available to completely clarify

the faunal shift which occurs in the vicinity of the Thai-Malaysian border. Al-

though climatic factors can apparently be correlated with some of the species,

this explanation cannot be used in interpreting the disappearance of some of

the Cellia species in Peninsular Malaysia and then their reappearance in Malay-

sia or Indonesia. The geological history of this area is complex, and there is

no doubt that the main elements of the Sunda Platform (Peninsular Malaysia,

Java, Borneo and Sumatra) formed a single land mass, broadly connected with

Thailand and the other continental area of Southeast Asia in relatively recent

times. For additional data the reader is referred to Dobby (1950) and to

Zeuner (1941).

Format and Treatment

The format followed here is essentially the same as that followed in earlier

publications of the Southeast Asia Mosquito Project (SEAMP). The synonymy

has been abbreviated, and only the more important entries have been listed.

For a more complete synonymy for a particular taxon the reader should con-

sult Stone, Knight and Starcke (1959) and the additions and corrections to that

work by Stone (1961, 1963, 1967, 1970). Additional detailed synonymy for most

species included in the present work will also be found in Reid (1968) and Chris-

tophers (1933). Within the parentheses following each citation, the symbols ~,

2, P, L, and E indicate that the publication deals with at least some part of the

male, female, pupa, larva or egg respectively; a single asterisk (*) following

the symbol indicates that at least some portion of the stage was illustrated.

Abbreviations for references conform to the BIOSIS 1972 List of Serials, Bio-

Sciences Information Service of Biological Abstracts, Philadelphia.

The synonymy is followed by a diagnosis, each species is described in some

detail, starting with thefemale, followed in order by the male, pupa and larva.

The egg stage has not been used routinely in this publication, but it may be re-

ferred to in some cases to illustrate a particular point. For the most part the

morphological terminology employed follows Belkin (1962), particularly as con-

cerns the setal numbers for larval and pupal stages. Additional terminology,

where applicable, follows Knight (1971) and Knight and Laffoon (1971) and also

draws heavily upon Reid (1968).

Each species is usually accompanied by at least 3 plates illustrating the

adult, pupal and larval stages. Most of the figures follow an identical format,

but in some cases additional drawings have been provided to illustrate special

morphological features. Almost all of the drawings are based upon several

Harrison and Scanlon: Subgenus Anopheles in Thailand 9

specimens selected from Thailand collections. In the very few cases where

extra-territorial specimens were used that fact is noted.

Wherever possible the 1st figure for each species consists of: a drawing

of the female, with left wing and legs removed; the male terminalia as they

appear in the usual microscopic preparation, prepared as a phantom view, to

show the relationships of the various parts; an enlarged drawing of the clas-

pette; an enlarged drawing of half the leaflets at the tip of the aedeagus; and a

dorsal view of the male head, with the antennal flagellae removed and the left

maxillary palpus turned to show the lateral aspect.

The 2nd drawing of the typical set of figures consists of the pupal skin and

a map showing the approximate known distribution of the species in Thailand.

The pupal cephalothorax is shown dissected, with only the left side shown,

including the respiratory trumpet, but excluding the major portion of the wing

case and the tubular portions of the cases of the appendages. The metathorax

is shown attached to the abdomen, and it and the abdominal segments are

prepared showing the dorsal setae on the left side of the drawing, the ventral

setae and structures on the right. The letter C is used to designate the meta-

thorax in this drawing and Roman numerals are used to designate the abdom-

inal segments.

The 3rd drawing of most sets consists of: the larval head, with dorsal

structures on the left, ventral structures on the right; the thorax and first 6

abdominal segments, with dorsal and ventral structures treated as in the head

drawing; a lateral view of the terminal abdominal segments; the mental plate;

and, in some cases the aulaeum or other detailed structures.

The letters P, M and T designate the prothorax, mesothorax and meta-

thorax respectively, and Roman numerals are used to designate the abdominal

segments.

A scale line has been provided with most figures of each stage to permit

assessment of relative sizes among the various species.

The species descriptions are followed by a listing of the type-data for the

species in question, and for other names listed in the synonymy.

The distribution section discusses the distribution of the species in Thailand

based on specimens examined, followed by a synopsis of the distribution else-

where in Southeast Asia. Species distribution records listed in Scanlon et al.

(1968) were based on specimens identified prior to this revisionary study, and

have been changed or corrected as necessary during this study. The Thailand

records are based on such a large volume of material in most cases (often

several hundred reared specimens) that it has seemed most suitable to give

them by administrative province only. Ina few cases, with rare species,

specific collection data have been provided. The records from outside Thai-

land are by country and in some cases major political subdivision. Distribu-

tional citations from outside Southeast Asia were used only when necessary to

illustrate a particular point. Specimens in the USNM collection or other col-

lections examined during this study are so indicated, and separated from cita-

tions based only on the literature. In discussions in the body of the text we have

frequently used geographic terms which have not survived the political vicissi-

tudes of recent years, but which are well known to biologists and zoogeographers

(e.g. Malaya, Borneo and Indochina).

Morphology

Figures 2 - 4 are included as a guide to the most commonly used morpholo-

gical features of the various stages discussed. The figures are intentionally

brief and confined to external characters. Specifics about frequently used char-

Figure 2. Anopheles (A.) sinensis. Lateral view to illustrate

selected morphological features.

A. Female: 1. labium (proboscis)

2. labellum

3. maxillary palpus

4. clypeus scales

Do. flagellomere

6. pedicel

7. vertex

8. occiput

9. anterior pronotum

10. propleural setae

11. upper midcoxal setae

12. halter

13. sternum VII scale tuft

14. cercus

B. Male: 1. labium (proboscis)

2. maxillary palpus, segment 4

3. flagellomere 12

4. pedicel

Do. anterior promontory of scutum

6. scutum

7. scutellum

8. hindcoxa

9. hindtrochanter

10. hindfemur

11. tergum III

12. sternum III

13. basimere

14. distimere

Fig.2

ADULT MORPHOLOGY

(Anopheles) sinens

Nore ONDA AIDOOHL WD =

—

SONI nor WN eH ~I OQ OF ® © DO

Hm co DO

Out Whe

Hq OO DO

Figure 3. Adult morphology.

A. Wing of Anopheles species denuded of scales.

upper calypter (squama) 13. vein Kgs

remigium 14. vein Mj,9

costa 15. vein M3.4

vein R 16. vein Cu

subcosta 17. vein M

vein Rg (radial stem) 18. vein Cug

origin of R45 19. crossvein M-Cu

crossvein r-m 20. plical fold

vein Ro.9 21. fringe scales

vein Ro 22. vein Cu

vein R 23. vein1A

vein Ry 24. humeral crossvein

B. Wing, showing most useful costal spots.

inner prehumeral accessory 8. middle dark spot

dark spot 9. subcostal pale spot

prehumeral pale spot 10. preapical dark spot

humeral pale spot 11. preapical pale spot

humeral dark spot 12. apical dark spot

presector pale spot 13. fringe spot 1A

presector dark spot 14. accessory sector pale spot

sector pale spot

C. Lateral view of female.

anterior promontory of scutum 9. metathoracic or posterior spiracle

anterior pronotal lobe 10. mesepimeron

posterior pronotal lobe 11. meron

anterior spiracle 12. sternopleuron

prealar lobe 13. propleuron

scutellum 14. vertex

halter 15. occiput

sternum I 16. upper mesepimeral setae

D. Dorsal view of thorax.

anterior promontory o. lateral area

anterior pronotum 6. parascutellum

fossa 7. prescutellar area

scutal angle 8. scutellum

E. Female head, anterior view (scales of the maxillary palpi

and proboscis removed).

erect scales of the vertex 7. clypeus

compound eye 8. segment 3 of maxillary palpus

pedicel 9. segment 4 of maxillary palpus

3rd flagellomere 10. segment 5 of maxillary palpus

segment 2, maxillary palpus 11. labellum

Frontal tuft 12. labium

F. Male genitalia, ventral view (dorsal before rotation).

basimere Oo. aedeagus

distimere 6. leaflets of aedeagus

proctiger 7. claspette

parabasal spines 8. club on claspette

9. tergum IX

| Wy

coe

Adult morphology

14 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

acters or characters that are exceptions to the rule are found in the following

discussion. Readers desiring a more detailed exposition of anopheline mor-

phology are referred to Christophers (1933), Gater (1934, 1935) and Reid

(1968). Figures 2 - 3 deal specifically with various adult morphological char-

acters, including the differentiating sex characters (antennae, maxillary palps

and genitalia), while Fig. 4 deals with those basic characters pertinent in lar-

val and pupal morphology. The following structures will be found the most use-

ful during the examination of adults. Head: The degree of antennal develop-

ment is a differentiating character between the sexes. Male anophelines typi-

cally have bushy (plumose) antennae. One exception is acaci, from Borneo

and the Philippines, which has female-like antennae. Other than bushiness,

both sexes exhibit the same basic antennal structure (minute scape, large glo-

bular pedicel and 13 flagellar segments). The more basal flagellomeres may

bear afew scales. The width of the erect scales on the vertex is a very use-

ful character in the separation of species in the Anopheles series. The maxil-

lary palps are 5 segmented (1st segment vestigial in females) and exhibit sexu-

al differentiation in that the last 2 segments of the male palps are distinctly

flattened and club shaped. Both sexes normally exhibit palps approximately

equal in length to the proboscis; however, the female of brevipalpis from

Malaysia exhibits palps approximately 0.75 the length of the proboscis (Reid

1968). The palps are typically covered with decumbent or erect scales and

frequently exhibit color patterns which are very important in identification.

The labium and labellum make up the discernable proboscis, which is covered

with decumbent or erect, unicolorous scales in the Thailand members of the

subgenus Anopheles. The clypeus is usually bare in anophelines; however,

the presence of scales on the clypeus is an important identifying character of

the hyrcanus species group. Thorax: When viewed from above, the thora‘<

consists almost entirely of the expanded scutum of the 2nd thoracic segment.

The coloration of the scutum and the occurrence, shape, arrangement and

coloration of its scales and setae are important characters. The scutal color

pattern generally fades fast and is of little value in the subgenus Anopheles;

however, a distinct pattern is detectable on umbrosus. Generally the prescu-

tellar space is bare, but the size and concentration of setae in the area is of

importance in the aztkenii species group. The occurrence of scales on the an-

terior pronotal lobes is a valuable character used in separating the Myzor -

hynchus and Lophoscelomyia series from the groups of the Anopheles series

found in Thailand. The occurrence and number of propleural setae is impor-

tant in a number of cases. Pleural scaling and setation is less useful in an-

ophelines than in culicines, but the occurrence of pleural scales and the number

of setae on the prealar lobe and upper mesepimeron (subalar setae of Reid

1968) are significant in the subgenus Anopheles. Wings: Wing markings are

extremely important in anopheline identification. A wing (Fig. 3) of a member

of the subgenus Anopheles has been used to illustrate the occurrence and no-

menclature of wing markings. The wings of members of the subgenus Anopheles

generally exhibit much less spotting than those of subgenus Cellia; however,

the nomenclature and position (when present) of the spots remains the same.

The reader must take care when trying to determine wing color patterns, for

the same vein may have different colored scales on the dorsal and ventral sides

of the wing, thus giving a different pattern when viewed from each respective

side. Such areas as the remigium and humeral crossvein and their respective

scale patterns are highly significant and should be viewed under high magnifica-

tion. The pale fringe spots at the terminations of some veins are often diffi-

cult to see without adjusting the lighting to favor transmitted or reflected light,

Harrison and Scanlon: Subgenus Anopheles in Thailand 15

or a combination of both. In addition to the fringe scales, there is a line of

tertiary fringe scales along the extreme wing margin which is rarely used in

taxonomic works, but can serve as an indication of a fringe spot. Anopheline

wing patterns are highly variable in some groups; consequently, one should

anticipate intraspecific variation in making specific determinations.

Due to differences encountered in the relative position of the preapical pale

costal spot, an arbitrary decision was made to define this spot as the one proxi-

mal to the tip of vein Ry. However, in the wmbrosus group this spot usually

ends level with the tip of R1;and we have chosen to continue calling it the "'pre-

apical" spot, while Reid (1950, 1968) refers to this as an "apical" spot. In

certain members of the asiaticus group only one pale spot occurs near the tip

of the wing, in such cases this spot is called the apical pale spot even though

it does begin slightly prior to the tip of Ry. Another member of the aszaticus

eroup, interruptus, has an upper spot from just above the tip of R, down to R

and a lower spot adjacent to the tip of R4,5. We interpret both of these spots

as being apical, thus, members of the aszaticus group do not have a preapical

pale spot on the leading margin of the wing. Legs: The occurrence and pat-

tern of pale annulations on some of the leg segments (particularly the tarso-

meres) are highly significant taxonomic characters in anophelines. These

markings, like wing patterns, can be quite variable, thus caution is necessary

in evaluating leg patterns. Other significant leg characters in the taxonomy

of the subgenus Anopheles are the occurrence of scales on the base of coxae

and the occurrence and coloration of erect scales on the apex of the hind femur.

Abdomen: In most anopheline species the abdomen is devoid of scales, thus

when present they are highly significant. Scattered scales on abdominal sterna

are significant in the barbirostris species group, and the occurrence of a scale

tuft on the 7th abdominal sternum is useful in identifying the barbirostris and

hyycanus species groups. Infrequently, the occurrence and coloration of

scales on the male genitalia serve as useful characters. Male Genitalia: The

genitalia of anophelines are not as useful in species differentiation as are those

of most culicines; however, characters of subgeneric, series and in some

cases, specific value, have been recognized in Anopheles. The parabasal

spines, located on the morphologically dorsal surface of the large, conical

basimere are used in subgeneric classification. The basimere bears other

setae, and particularly scales which may be of specific significance. The api-

cal distimere has not been found to furnish useful characters for identification.

The structure and appendages of the claspette are important and may be readily

seen in simple slide preparations of the intact genitalia. A club, composed of

fused setae, is often present on the dorsal lobe of the claspette, and this may

present quite a different appearance when the claspette is dissected and viewed

from the lateral aspect. The claspette drawings made for the species herein

were made with the claspette flat as in the usual dorsal or ventral aspect. The

aedeagus is a tubular structure, but appears to be flat in the normal micro-

scope preparation. In addition, this structure is usually strongly arched, as

will be apparent in the following illustrations. The aedeagus itself is of little

value in the taxonomy of the subgenus Anopheles, except in the case of the

aitkenii species group where denticles on its tip are highly significant. The

occurrence and number of paired leaflets found at the tip of the aedeagus are

very important and are used in the differentiation of the series in the subgenus.

In this study, these structures with the tip of the aedeagus were dissected free,

stained and mounted as flat as possible ona slide. The appearance of the leaf-

lets prepared thus is quite different than that normally seen in whole genitalia

mounts. Female Genitalia: Genitalia preparations were made for most of the

16 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

included species, but few useful characters were found. Cibavrial Armature:

This internal structure, examined briefly during this study, was used by Chris-

tophers (1933) and others in defining species groups in the subgenus Cellia.

The armature has been found to be of little use in separating the species of the

subgenus Anopheles, except in the case of certain members of the umbrosus

species group (Reid 1968). For literature pertinent to this subject the reader

is referred to Christophers (1933), Gater (1935) and Reid (1968).

While whole mounted larvae have been used in anopheline taxonomy for

years, the practice of mounting whole pupae never became popular, mainly

because of difficulties encountered in mounting the bulky body. Early workers

frequently assigned and described a given larva to a given adult on the basis

of whole mounted larvae and reared adults from the same collection. Certainly,

this method has led to errors and much taxonomic confusion. During the last

20 years taxonomists have become much more aware of the value of correlating

cast immature skins with the resulting imago for increased taxonomic pro-

ficiency. During this period a general (not total) concensus has evolved on the

nomenclature and general homology of the chaetotaxy of the larval and pupal

stages. With these problems essentially resolved, the taxonomy of anophelines

has been enhanced by the study of the respective immature skins. Now, pupal

chaetotaxy is as important in anopheline taxonomy as the study of the adult, lar-

vae or egg. Infact, the pupal stages of some members of the subgenus Ano-

pheles possess the most reliable differentiating characters at the species level.

As mentioned earlier, the chaetotaxy and nomenclature used herein pri-

marily follows Belkin (1962) and Reid (1968). For explanations and the differ-

ences between the various systems the reader is referred to the above refer-

ences as well as Knight (1971). Pupa: The following pupal illustrations (Fig.

4, A-D) are basic and designed to point out body regions and structures (not

chaetotaxy) that will hereafter be presumed known by the reader. The illus-

trations were prepared from mounted skins, in which the head and thorax (mi-

nus the metathorax) was opened along the midline and flattened (Fig. 4A), while

Fig. 4B, depicts the metathorax and abdomen. On the head the pigmentation

pattern of the antennal case can be of specific value in the hyrcanus group. The

most frequently used thoracic characters involve the paired respiratory trum-

pets which furnish series, specific and infraspecific characters. Two basic

types of trumpets are found in the subgenus Anopheles. (A) Angusticorn (Fig.

81): Simple and tubular, having the longest axis vertical, more or less in line

with the stem of the trumpet, and with a single split (cleft) or notch of varying

depth on one side. (B) Laticorn (Fig. 11): More complex, with longest axis

more or less transverse to the stem, the rim sometimes with a second split

opposite the meatal cleft, or if longest axis not transverse to stem, then there

is a lobe, often complex on the rim. Members of the subgenus Cellia have only

the angusticorn type trumpet, while the majority of the species of the subgenus

Anopheles in Thailand have some variation of the laticorn type. The occurrence

and patterns of pigmentation on the wing cases is of value in the hyrcanus group.

Most abdominal (including metathorax) characters involve chaetotaxy patterns,

however, some general morphological characters are pertinent. On the venter

of the 1st abdominal segment is a pair of spiracular sensillae, while on seg-

ments II - VII these spiracular sensillae are located dorso-laterally. Additional

pairs of dorsal sensillae are found on segments III - V. These spiracular and

dorsal sensillae are sometimes easily confused (particularly the dorsal sen-

sillae) with setal alveoli. The occurrence of small denticles on the hind mar-

gin of the abdominal tergites is of value in identifying montanus and most of

the members of the umbrosus group. Sexual differentiation of the pupal stage

Harrison and Scanlon: Subgenus Anopheles in Thailand 14

is possible by determining the length of the genital lobe, which is located at

the posterior end of the abdomen and ventral to the paddles. The male (Fig.

4C) has a longer genital lobe than the female (Fig. 4D). In the hyrcanus spe-

cies group, the sexes can also be differentiated in the pupal stage by the shape

of the palpal case (Figs. 11, 14). The male has the tip of this structure long

and narrow, while it is short and broad on the female. The value of characters

found on the paired paddles cannot be overstressed. Their shape (length-

width relationship) can be highly significant, as can be the occurrence and

pattern of pigmentation on these structures. The outer paddle margin has

a basal area of minute denticles which presents a refractory appearance in

transmitted light. These structures tend to become larger and more tooth-like

toward the apex and then give way abruptly or gradually to filamentous hair-

like structures. The relative proportion of the paddle margin occupied by either

one or both types of structures that is refractory to light is called the refractory

index, measured (following Reid) in a straight line from the base of the paddle

to the end of the refractile area, divided by the length from the base of the pad-

dle to the base of the paddle hair (1-P). The long filamentous structures on

the outer margin of the paddles are generally referred to as the paddle fringe.

The concentration and length of the fringe is often of taxonomic significance.

The larval drawings accompanying the species descriptions were prepared

from 4th stage larvae, slide mounted dorsal side up, with the terminal abdom-

inal segments cut and placed laterally to display the setae and pecten. Where

possible, larvae were from the same series in which larval skins had been com-

pared with pupal skins and imagos to assure correct identification. .

Larva: The characters most frequently utilized in anopheline larval taxonomy

involve chaetotaxy arrangement and degree of branching. Both of these aspects

of chaetotaxy are subject to variation and this variation will be encountered by

workers in the field; consequently, previous studies should be consulted where

the range of inter-intraspecific variation has been documented for a species or

group (see Reid 1953). Of the head chaetotaxy, 2, 3 and 4-C in the Belkin

system (the inner, outer and posterior clypeal setae) are probably the most

pertinent. Also important are 5 to 7C (the frontals) 8-C (the inner sutural)

and 9-C (the outer sutural). The remaining dorsal and ventral setae are used

infrequently. The ventral mental plate is strongly sclerotized and pigmented

and can be taxonomically significant. The thoracic and abdominal chaetotaxy

is also highly significant. Such setae as 1-P, 14-P, 1-M and the pleural

setal groups of the thorax are used frequently in taxonomy. The extent of

sclerotization of basal tubercles can be significant, as can be color patterns,

however, color patterns are most often used with living larvae. The occur-

rence, distribution and extent of development of palmate setae is highly signifi-

cant. A palmate seta is one in which the branches have undergone various

degrees of flattening and pigmentation. These setae are typically located on

some or all of the abdominal segments except the 8th and 10th, but may also be found

on the metathorax. Setae 1 of the prothorax and mesothorax frequently exhibit

a fan-like shape, but are not considered palmate setae. On the abdomen the

chaetotaxy of segment I is reduced and somewhat unusual, that of segments II -

VII is similar in overall pattern while segment VIII exhibits a highly modified

setal pattern. Aside from the spiracular apparatus (Fig. 4J), segment VIII

also bears the lateral pecten plate, which is sclerotized and has caudally

directed teeth. The number and conformation of the pecten teeth may be of

taxonomic importance in some species groups. Each of the abdominal segments

usually bears at least one sclerotized dorsal (tergal) plate, and frequently 2.

The size of the anterior tergal plate can be very significant (see palmatus, in

Figure 4. Larval and pupal morphology.

A. Pupal cephalothorax (less metanotum).

head shield 4. antenna case

trumpet Oo. Leg cases

mouthpart cases 6. wing case

B. Pupal metathorax and abdomen.

metanotum 5. dorsal sensillum

. ventral spiracular sensillum 6. seta 9-VI

float seta 7. paddle

halter case 8. dorsal spiracular sensillum

C. Terminal portion of male anopheline pupa (dorsal).

genital lobe 2. midrib 3. seta 1-P

D. Terminal portion of female anopheline pupa (dorsal).

genital lobe 2. hair-like paddle fringe

E. Head of anopheline larva (ventral).

mouth brushes 4. ocular bulge

mental plate Dd. collar

labial plate

F. Tip of larval antenna (dorsal).

dorsal sabre piece 2. seta 4-A

G. Terminal segments of larval abdomen (lateral).

median dorsal valve 4. pecten plate

dorsolateral valve o. saddle

ventrolateral valve 6. anal gill

H. Thorax (dorsal, left; ventral, right).

prothoracic group of pleural setae 3. metathoracic group

mesothoracic group 4. metathoracic palmate seta

I. Prothoracic pleural setal group (left side, ventral view).

seta 10-P 4. seta 11-P

seta 9-P 5. spine separating anterior (9 - 10)

seta 12-P and posterior (11 - 12) pairs of

setae.

J. Spiracular apparatus (ventral).

median dorsal valve 3. ventrolateral valve

dorsolateral valve 4. spiracle

Larval and pupal morphology

20 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

the aitkenii group). The larvae of montanus, members of the umbrosus group

and infrequent specimens in the barbirostris and hyrcanus: groups, have a small

ventral plate immediately in front of 14-VIII. For convenience, this plate has

been illustrated here on segment VII. Puri (1931) originally noted this plate

and placed it at the caudal margin of segment VII. This interpretation was

followed by Reid (1950, 1968), but Belkin (1962) and Zavortink (1973) have

shown this plate on the cephalic margin of segment VIII. We have examined a

large number of larvae with. this character from both the lateral and ventral

views, and suspect this plate is actually intersegmental as is so-called seta 14-

VIII and seta 14 on the more anterior segments. Occasionally numerous tiny

spicules may be noted, particularly on the venter of some of the abdominal seg-

ments. These have been noted occasionally by various authors, even used as

taxonomic characters. However, their significance is not understood and they

are best disregarded (except for sintonoides), particularly since their occur-

rence is not consistent. Such structures may be correlated to an environmental

factor such as the hairiness factor found in Aedes larvae (Colless 1956). The

terminal or anal segment, regarded as segment X by morphologists, bears a

dorsal plate (saddle), setae and gills, but is seldom used in the taxonomy of

southeast Asian Anopheles. The spiracular apparatus follows the same basic

plan in almost all the species described herein. The anterior median dorsal

valve normally has a small transparent knob caudally, but this extends into a

posterior "'stigmal club" on pollicaris, a pigmented "'stigmal knob" on ftigertti

and members of the culiciformis group anda "'stigmal filament"'on umbrosus. The

only similar structure found in Asian Anopheles is found in the bancrofti group

of Australia, New Guinea, Philippines and Celebes and in atratipes Skuse, an

Australian species currently placed in the Anopheles series.

Egg, The egg stage has proved extremely useful in delimiting species in the

maculipennis species group in the Palaearctic region. Eggs of many of the

anophelines found in Thailand have been described to some extent, largely from

India (Christophers 1931, 1933), Ceylon (D'Abrera 1944), the Philippines

(Urbino 1936) and Malaya (Reid 1968). Nevertheless, it should be noted that

the eggs of at least 10 of the following species are unknown and studies are

needed on anopheline eggs in Southeast Asia to help unravel the taxonomy of

some of the species complexes. Insufficient material was available to illus-

trate the eggs of the Thailand species, but reference is made to the descrip-

tions given by the foregoing authors where applicable. Aberrant or unusual

specimens: In the course of field work occasional specimens are found which

exhibit unusual characters or an unusual arrangement of characters. On rare

occasions in the past such specimens have been mistakenly identified as new

species, thus creating problems. Hopefully, with the aid of the recent outline

of the genetic mutants found in anophelines, (Kitzmiller and Mason 1967) mod-

ern workers will recognize such aberrations for what they are and confirm the

existence of some of these inherited variations in wild populations. Such un-

usual characters as melanistic wing scale patterns and anomalies in the tergal

plate pattern and chaetotaxy of larvae and/or pupae were observed during this

study. The foregoing case of melanistic wing scale pattern presented a com-

pletely unidentifiable adult, fortunately, associated immature skins were avail-

able to clarify its identity. Another unusual character exists in the male of

acaci, which exhibits female-like antennae. Although this species is currently

known only from Borneo and the Philippines, the close relationship of acaci to

some of the Thailand aitkenii group species warrants further study.

To many workers keys are the most important results of such efforts as

this, and when the identity of vectors of pathogens is in question this may well

Harrison and Scanlon: Subgenus Anopheles in Thailand 21

be true. However, keys are only working tools and all persons concerned should

realize the fallibility of keys in view of total population concepts and the previ-

ously mentioned aberrant specimens. This is particularly true in view of the re-

cent recognition of many cryptic species and sibling groups among the mosqui-

toes. In essence, the identity of some species can only be ascertained by etho-

logical characters, or the association of immature skins with the adult e.g.,

barbirostris and campestris. Unfortunately, such time consuming identification

techniques are not always feasible when a medical entomologist is confronted

with a mosquito-borne disease epidemic. In such situations the use of the word

"eroup" is often advantageous (i.e., barbirostris group, hyrcanus group, etc. ).

This liberal approach to taxonomic identifications should be used only during

medically or economically urgent situations. Under routine situations workers

who are not taxonomically oriented or lack an associated taxonomic capability

can send specimens or series of specimens to specialists for verification.

The keys used herein are artificial, designed primarily for reliability and

time conservation, and should not be construed to indicate relationships, even

though such may be the case. The authors have followed the system of arrange-

ment used by Reid (1968), where the keys are located with the given series or

species group. This arrangement allows the user to quickly turn to the descrip-

tions and taxonomic discussions for confirmation or help. Some entirely new

keys are presented, but most are adaptations in one fashion or another of earlier

Reid keys, which have been altered to fit the Thai fauna or the desires of the

authors. Footnotes have been added to certain couplets to indicate possible

problem areas or alternate solutions.

Certain species considered doubtful records for Thailand by Scanlon et al.

(1968), are disregarded in this publication. Accordingly, the keys have been

altered to exclude those species (gigas formosus and gigas sumatrana).

Taxonomy

The subfamily Anophelinae may be defined as those mosquitoes species in

which: 1) the adult exhibits no scales or very few scales on the abdomen,

particularly on the sterna; the female maxillary palps are approximately as

long as the proboscis; the female has only one spermatheca; and the males have

the 2 terminal segments of the maxillary palps enlarged to form a distinct club.

2) the pupa has seta 9 on abdominal segments III - VI single, spine-like and

located in the latero-caudal corners; and the tip of the paddle has a pair of

setae, one of which is ventral and subapical. 3) the larva differs from all

other mosquitoes by the absence of a well developed and sclerotized respira-

tory tube, and have the spiracles opening from a spiracular plate on segment

Vill.

Three genera are presently recognized in the subfamily, Chagasia Cruz,

Bironella Theobald and Anopheles Meigen. The 1st of these is found only in

the Neotropical Region, the 2nd only in the Australian. We deal here only with

the genus Anopheles, which is widely distributed in the tropical and temperate

areas of the world.

GENUS ANOPHELES MEIGEN

Anopheles Meigen 1818: 10. Logotype: Anopheles maculipennis Meigen

(International Commission on Zoological Nomenclature 1959).

(The synonymy for the genus as here recognized is given in detail by Stone,

Knight and Starcke 1959, under the appropriate subgenera).

22 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Readily distinguished from other mosquitoes in Asia by the following com-

bination of characters: Female. Palps at least 0.75 the length of the proboscis;

scutellum rounded; abdominal scaling usually poorly developed; one spermatheca.

Male. Segments 4 and 5 of the maxillary palpus developed as a club, directed

laterally. Larva. Lacks a respiratory tube, the spiracles opening on the body

wall of segment VIII. Pupa. Paddle undivided, or slightly invaginated, seta 9

of the abdominal segments spine-like on IV - VII. In addition, the adults may

usually be recognized in life by their resting attitude, usually forming an acute

to right angle with the surface, the larvae also have a characteristic attitude,

resting parallel with the surface with the palmate setae of the abdomen in con-

tact with the surface film.

A more comprehensive description is as follows:

FEMALE dead. Eyes widely separated, several long setae usually projecting

forward between them forming frontal tuft of erect setae and scale-like setae;

head scales of several types, scales of occiput and vertex erect, fan-shaped

and often apically notched; interocular space up to vertex with ocular scales and

line of ocular setae internal to scales; ocular line without scales laterally and

ventrally; antenna with rounded pedicel, lacking protuberances, often with small

number of scales, flagellomeres with whorls of sparse setae, without matted

setae or other modifications, but basal segments may have flattened scales;

clypeus sometimes with scales; maxillary palpus generally as long as proboscis

or at least 0.75, that length, with 5 segments, basal segment minute; proboscis

covered with scales, scales decumbent or erect, labium flexible, sometimes

with slender setae ventrally and near tip, labellum without scales, generally

lighter in color than labium; ventral area of head with tuft of postgenal setae;

neck supported by long cervical sclerites. Thorax. Scutum elongate, slightly

convex, often with distinct lines in median and submedian areas, may have

covering of minute setae, sometimes with well developed dorsocentral and

acrostichal setae; anterior promontory often with a well developed scale tuft,

sometimes with additional antero-lateral tufts; scutellum without lobes, with

fairly evenly spaced setae and sometimes scales; anterior pronotum (apn)

with setae, sometimes with scales; posterior pronotum (ppn) bare’; propleuron

(ppl) usually with 1 - 5 setae, sometimes bare; spiracular area (sp) with or

without setae; sternopleuron (stp) with upper and lower setal groups; upper

mesepimeron (mep) with setae, lower mesepimeron usually bare. Wing. With

well developed usually pale and dark decumbent and plume scales, fringe scales

present; cell Ro at least as long as vein Ro,3; vein Re always connected to

Ro,3 at the base; vein 1A reaching the wing margin distal to the origin of vein

Cu, Halter. Stem usually pale, knob pale or dark scaled. Legs. Long,

slender, without tibial bristles, pulvillus absent, claws usually simple, base of

femur sometimes markedly swollen. Abdomen. Generally without scales, or

with these usually restricted to ventral tufts. Genitalia. Single spermatheca;

tergum and sternum VIII truncate or broadly rounded; tergum IX a narrow band;

tergum X semicircular, covering cerci; cerci truncated cones, bluntly pointed,

may have setae.

MALE. Similar tofemale in generalhabitus, usually smallerand slight in

general appearance. Head. Pedicel larger, partially obscuring clypeus, with-

out scales; flagellomeres with strongly plumose setal whorls except for 2 most

distal segments; palpus about as long as proboscis, 4th and 5th segments slightly

or conspicuously swollen, turned laterally in life, with long hairs at apex of 3rd

segment and on margins of 4th and 5th segments. Thorax. Markings similar to

those of female. Wing. Usually more narrow than that of female, markings

Similar, but with fewer scales and paler, with fewer dark areas in species with

Harrison and Scanlon: Subgenus Anopheles in Thailand 23

bicolored wings. Legs. Markings usually less vivid than on female, foretar-

somere 9 longer than, or at least equal to foretarsomere 4 and bearing a

basal protuberance with setae, claw with submedian and sometimes basal

teeth; tarsomeres 4 - 5 and claws of remaining legs like those of female.

Abdomen. With fewer scales than female, except on the basimeres. Genitalia.

Tergum and sternum IX forming complete ring at base of basimeres; basimere

simple, without lobes, with one or more groups of specialized setae or spines;

distimere simple, tubular, with apical spine and several small setae; claspette

well developed, with one to 4 lobes and associated setae; aedeagus simple, tu-

bular, with or without apical leaflets or spines; proctiger largely membranous,

with lateral thickening on some species, generally without taxonomically useful

characters.

PUPA. Cephalothorax. With 9 pairs of setae, 1 to 3-C in ocular group

near midline, 4 and 5-C more anterior and lateral, 6 and 7-C posterior to 4

and 5-C and 8 and 9-C posterior to trumpet; trumpet open and with margin

bearing at least one cleft of varying width and depth. Abdomen. For typical

setation see Fig. 6, numbered according to the Belkin (1962) system; seta 9

(lateral seta) simple, spinelike, inserted on posterior corners of segments

II - VII, usually with branches on segment VIII; seta 3-VI mesad of 1-VI ex-

cept in umbrosus group. Paddle. Oval in outline, external buttresses and

midrib distinct; both margins usually with more or less distinct fringes of fine

spines on distal portions, often with smaller hyaline denticles near base; seta

1-P terminal, straight or markedly hooked, 2-P ventral and cephalad of 1-P.

LARVA. Integument usually glabrous, but may bear simple to elaborate

spicules covering most of thorax and abdomen. Head. Generally longer than

wide, without marked concavity posterior to antennae. Mouthparts not highly

modified, mental plate and aulaua developed; setae 0 and 1-C inserted on pre-

clypeal area; 2 and 3-C (inner and outer anterior clypeals) well developed; 4-C

(posterior clypeal) smaller than 2 and 3-C in most species; 5 to 7-C (frontals)

usually strong, feathered, inserted in irregular line; 11-C (subantennal) large,

plumose; antenna short, simple, with variously developed spicules; seta 1-A

simple or branched, inserted in basal 0.5; 2 and 3-A spiniform, sabre-like,

4-A usually branched. Thorax. Setae 1 to 3-Pnear midline, usually separate,

sometimes with common base; pleural setal groups (9 - 12) well developed on

all 3 thoracic areas; seta 1-M often strongly developed, plumose; seta 3-T often

developed as palmate seta. Abdomen. Anterior tergal plates varying in size,

on segments I - VII smaller single or paired plates may occur posterior to the

main plate on some species; seta 1 frequently developed as palmate on most

abdominal segments, but filamentous on all segments in some Asian species;

lateral setae (6 - 7) strongly developed on segments I - II, seta 6 strongly

developed on segment III and variously developed on segments IV - VI; seg-

ment VIII with dorsal spiracular apparatus and paired lateral pecten plates;

segment X with sclerotized saddle, not complete on ventral aspect, saddle

seta (1-X) simple or branched; setae 2 and 3-X plumose, 3-X usually stout

and hooked distally; 4-X consists of 9 pairs of setae arising from sclerotized

erid.

DISTRIBUTION. Species of the genus Anopheles occur in temperate, sub-

tropical and tropical regions of the world, with the exception of island groups

in the Pacific and isolated islands in the Atlantic. The arctic areas are free

of Anopheles, even at latitudes where certain Aedes species are abundant.

Southeast Asia has a rich and varied anopheline fauna at the species level,

and no reasonably large island or portion of the mainland is without representa-

tive species. It is possible that some of the higher mountains in the area have a

24 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

poor representation of the genus, but little collecting has been done at higher

altitudes.

TAXONOMIC DISCUSSION. The genus Anopheles is probably the most

thoroughly studied genus in the family Culicidae, primarily because of the

importance of its species as vectors of human malaria parasites. The intense

early interest of a host of workers, many relatively untrained in taxonomy,

gave rise to a multiplicity of specific and generic synonyms. Edwards (1932)

brought considerable and lasting order out of this chaos, uniting all of the forms

covered by the description given above in the genus Anopheles, with 4 subgen-

era, and a number of groupings of species below the subgeneric level. The

system adopted here (Stone, Knight and Starcke 1959) follows Edwards' revision

rather closely, and recognizes the following subgenera: Stethomyia Theobald;

Anopheles Meigen; Nyssorhynchus Blanchard; Kerteszia Theobald; Lophopodo-

myta Antunes; and Cellia Theobald. The subgenera Anopheles and Cellia occur

in Thailand and only the former is considered here.

The subgeneric classification of Anopheles is based primarily on the number

and position of the parabasal spines on the basimere of the male genitalia (Ed-

wards 1932; Christophers 1933), but for the 2 subgenera found in Thailand

there are a number of other satisfactory criteria which are outlined in the keys

below.

Most workers regard the genus Anopheles as the most primitive in the

Culicidae, but Belkin (1962) believes that they are highly specialized and not

ancestral to other groups in the family. In the absence of significant fossil |

records for the mosquitoes it is difficult to assess this question and it is beyond

the realm of this study.

BIOLOGY. The overwhelming majority of Anopheles species have their im-

mature stages in various types of ground water, ranging from water-filled

animal footprints to running stream margins and the margins of lakes and

marshes. Southeast Asia has a particularly rich fauna of treehole and bamboo

breeding Anopheles species; but apparently there are few axil breeding species,

such as those found in the Bromeliaceae of the New World. One Thailand

species, tigeritz passes its immature stages in the holes dug by fresh water

crabs. Further collections may reveal more species in this habitat. The great-

est numbers of anopheline larvae are found in the broad rice producing areas

of Thailand, but even quite small water bodies can hold an amazing number of

larvae.

Generally, anopheline larvae require relatively clean water for develop-

ment, but some species, like barbirostris are found in highly polluted water

near human habitation. Most species are found in still or very slightly moving

water, but aberrvans and bengalensis are found in the eddies of small hill

streams. Again, most species are restricted to fresh water, but baezaz is

usually found in brackish water along the coast. In summary, almost any body

of water within the confines of Thailand may be expected to yield some Anopheles

species if examined in sufficient detail.

As far as known, the larvae of all the Anopheles species in Thailand are

filter feeders, lying parallel with the surface with the head rotated so that the

morphologically ventral surfaces lie in contact with the surface film. The

mouthbrushes are in constant movement to bring small floating or suspended

particles into reach. Limited observations on the treehole breeding species in

Thailand indicate that the larvae behave in the same manner in these limited

confines. Unlike the immature stages of some other treehole inhabiting species

in Anopheles or other genera, there is little evidence that the larvae of the

species in Thailand have developed a predatory feeding habit.

Harrison and Scanlon: Subgenus Anopheles in Thailand 25

The female anophelines of Thailand appear to feed on mammal and bird

blood, with the former predominating. The feeding habits of many of the

forest-inhabiting species (e.¢g., aberrans, asiaticus and insulaeflorum) re-

main largely unknown. It is possible that some of these feed on common cold-

blooded vertebrates in the country, but there is no evidence for this as yet.

Many of the species discussed below can be collected resting by day in houses

and other structures, but many species occur in Thailand which are rarely, if

ever, encountered in artificial resting places. Anopheles adult mating and

feeding activity is almost entirely restricted to the hours of greatly reduced

light, or darkness. The females of some forest species (umbrosus group)

will take a blood meal during the day, under heavy forest shade.

In Thailand, members of the genus Anopheles are vectors of human

and simian malaria and filarial parasites. They have not been implicated

in the transmission of viral diseases, but isolation efforts along these lines

should be attempted, particularly in view of the virus-Anopheles relationships

discovered in Africa during the last 15 years.

KEY TO THE SUBGENERA OF ANOPHELES IN THAILAND

ADULTS AND IMMATURE STAGES

Costa entirely dark scaled, or divided by pale spots into not more than 3

dark marks involving both veins C and R-R,; male basimere with 3

spines (2 parabasal and 1 internal), the inner most parabasal shorter

and stouter than the outer; larval seta 1-A branched (except in several

species with seta 5, 6, 7-C reduced); setae 2-C inserted close together,

closer (rarely equal) than distance between 2-C and 3-C on either side;

male pupae with end of genital lobe obtuse, without pair of knobs; pupal

seta 1-Pusnally short andstraight: eacacien 25a wee Anopheles

Costa divided by pale spots into 4 or more dark marks involving both veins

C and R-R,; male basimere with cluster of 4 - 5 parabasal spines, no

internal spine; seta 1-A simple; setae 2-C inserted at least as far apart

as the distance between 2-C and 3-C on either side; genital lobe on male

pupae with a pair of rounded points or knobs on apex; pupal seta 1-P

usually lone and Nhooked or curved, oe eer eee Cellia

SUBGENUS ANOPHELES MEIGEN

Anopheles Meigen 1818: 10. Logotype: Anopheles maculipennis (International

Commission on Zoological Nomenclature 1959). Christophers 1915:

383.

(For detailed synonymy see Stone, Knight and Starcke 1959; Stone 1961, 1963,

1967, 1970. The latest views on the classification of the subgenus are found

in Reid and Knight 1961. )

In addition to the characters given in the above key, the following may

assist in the correct subgeneric placement of the Thailand spec e

FEMALE. Cibarium without teeth.

MALE. Basimere with at least the inner parabasal spine set on a distinct

prominence.

26 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

PUPA. Laticorn and angusticorn type trumpets, members of the subgenus

Cellia exhibit only the latter type.

LARVA. Setae of the 3 pleural groups generally all simple; branched

antennal seta usually inserted on the inner aspect of the antennal shaft.

DISTRIBUTION. Species of subgenus Anopheles are widespread in both the

New and Old World, with the numbers of species in each area roughly similar.

In Thailand, more species of the subgenus were collected in the southern por-

tion of the country and areas with tropical wet forests than in other areas, but

large numbers of individuals of species in the hyrcanus and barbirostris groups

were found throughout the country.

TAXONOMIC DISCUSSION. It is generally accepted that the subgenus

Anopheles is more primitive than Cellia. Reid (1968) has discussed the rela-

tionships among species groups in the subgenus, and with species of the sub-

genus Cellia in detail. A number of systems have been proposed to indicate

the relationships among species of the subgenus (Christophers 1915; Edwards

1932) and we have followed the most recent of these (Reid and Knight 1961).

Edwards (1932) used the infrasubgeneric categories ''group" and "'series'’; but

Reid and Knight wisely substituted the term "'section" for group, since the

latter term is now so widely used in connection with studies of sibling species

aggregations.

Reid (1970) explained his usage of the terms "'species-complex" and ''spe-

cies-group"’. The former refers to ''suspected species of groups before they

have been sufficiently analyzed to establish that the forms they contain are

distinct species"’. The latter refers to those same species, but after their

suspected identity has been established. We are following Reid (1968) who

used the term "'species-group"' for the units covered herein, however, we feel

that more cryptic species probably exist, and our understanding of these groups

in terms of their components and distribution in the entire Oriental faunal re-

gion is still incomplete.

The infrasubgeneric catagories recognized and discussed here have no

formal standing under the International Code of Zoological Nomenclature.

They are convenience categories only, useful for highlighting the natural rela-

tionships among the species of the subgenus. The Reid and Knight (1961)

classification system is as follows.

SUBGENUS ANOPHELES - LATICORN SECTION

Myzorhynchus Series

- ANGUSTICORN SECTION

Anopheles Series

Lophoscelomyia Series

PRIMARY KEYS

KEYS TO THE SECTIONS, SERIES AND SPECIES GROUPS

OF THE SUBGENUS ANOPHELES IN THAILAND

ADULTS

1. Costa divided by pale spots into 4 or more dark marks involving both

Costa Und Vet RAR OS ey es pg eS (Subgenus Cellia)

Costa divided by pale spots into not more than 3 dark spots involving both

costa and vein R-Rj, or costa all dark. (Subgenus Anopheles)... 2

ort)

3. (2)

4. (2)

1,46)

Harrison and Scanlon: Subgenus Anopheles in Thailand at

Wing scales all dark; anterior pronotal lobe without scales.

(ANGUSTICORN SECTION, In part)

(Anopheles series, p. 144)... 3

Wings with pale and dark scales; anterior pronotal lobe with scales. . 4

Erect head scales long and narrow, only slightly expanded apically;

antennal flagellomeres without scales.

aitkenii sp. group (p. 147)

Erect head scales widely expanded apically; flagellomere 1 with dark

SCALCS, iia Sore ae a a culiciformis sp. group (p. 176)

sintonoides (p. 181)

Apex of hindfemur with conspicuous tuft of white scales preceded by tuft

Of Dlaek Scales iiss, 6 ay es (ANGUSTICORN SECTION, In part)

(Lophoscelomyia series, p. 132)

asiaticus sp. group (p. 135)

Hindfemur without distal broad white band or tufts.

(LATICORN SECTION)

(Myzorhynchus-series,: sp. 29)........ 8

(For separation of males of this series see p. 30.)

Clypeus with patch of dark scales on each side; basal 4 - 8 flagellomeres

With pake Scales. a, SA ee ee hyrcanus sp. group (p. 36)

Clypeus without scales; 1st (basal) flagellomere only, with scales. . . 6

Sternum VII with tuft of black scales. . barbirostris sp. group (p. 78)

sternum Vil wWithOul SCAleSi. gle is Pee ee a as We eh es c

Hind-tarsomere 5 all white. 4.4.42. albotaeniatus sp. group (p. 31)

montanus (p. 34)

Hind tarsomere 5 mostly dark scaled. . . umbrosus sp. group (p. 105)

PUPAE*

Trumpet with longest axis vertical to stem; rim of trumpet simple, with-

OUE SeeQntary ClEll OF Na Cte 40 eS gee ie te waren ie stress

Trumpet with longest axis transverse to stem; rim of trumpet often with

secondary cleft; if longest axis not transverse, then rim of trumpet

with tragus (often complex).

LATICORN SECTION (Myzorhynchus series)... . 5

2.(1) Paddle broad, seldom more than 1.5 times as long as wide; seta 5 on

V - Vil usually no stouter than seta 1 on V - VII; male genital lobes

with apical knobs. ti a eae (subgenus Cellia)

Paddle usually elongate, about 1.75 times as long as wide; if not elongate

then seta'5 on V - VII is much stouter than seta 1 on V - VII; male

genital lobes obtuse, not ending in knob.

ANGUSTICORN SECTION... . 3

*kyondawensis known only from larval stage.

28 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

3.(2) Seta 9 on V - VII frayed, or smooth and hooked at tip; seta 5 on V - VII

very strong; seta 1 on V - VII very small and weak; bamboo and tree

hole hamitatsy Coe a (Lophoscelomyia series, p. 132)

asiaticus sp. group (p. 135)

Seta 9, 1 and 5o0n V - VII not so; not from bamboo or tree holes (ex-

CONE Sinlonoides) 1 SO (Anopheles series, p. 144).... 4

4.(3) Trumpet with shallow meatal cleft, meatus (stem) more than 0.5 as

long as trumpet; paddle seta 1 long and hooked; leaf axil, bamboo

and tree hole habitats. ...... culiciformis sp. group (p. 176)*

sintonoides (p. 181)

Trumpet with deep meatal cleft, meatus 0.33 or less as long as

trumpet; paddle seta 1 simple and straight; ground water habitats.

aitkenii sp. group (p. 147)

5.(1) Hind margins of abdominal tergites with denticles (except baezai);

trumpet rim with tragus or deep secondary cleft (Fig. 6). .... 6

Hind margin of abdominal tergites without denticles; trumpet rim with-

out tragus, but may have shallow secondary cleft (Fig. 11).... 7

6.(5) Trumpet with deep secondary cleft; lateral spines (seta 9) long.

albotaeniatus sp. group (p. 31)

montanus (p. 34)

Trumpet without secondary cleft, but with tragus; lateral spines usually

Shee a ee ON PN aha umbrosus sp. group (p. 105)

7.(5) Seta 1-VIl a strong tuft, with 15 or more branches.

barbirostris sp. group (p. 78)

Seta 1-VIL with not more than 13 branches.

hyrcanus sp. group (p. 36)

LARVAE

L. Bases of setae 2-C wide apart, closer to the bases of setae 3-C than to

one another; seta 1-A small or minute and nearly always simple.

(Subgenus Cellia)

Bases of setae 2-C closer to one another than to bases of setae 3-C;

seta 1-A branched, often large. (In a few forms either setae 2-C

are rather wide apart and seta 1-A is branched, or setae 2-C are

close together, with seta 1-A simple and setae 5,6, 7-C, reduced).

(Subgenus Anopheles)....... 2

2.(1) Branches of seta 1-A not reaching much beyond the middle of the anten-

nal shaft, whole seta usually less than 0.95 length of shaft.

(ANGUSTICORN SECTION) .... 3

Branches of seta 1-A long enough to reach close to or beyond the end of

the shaft, whole seta usually more than 0.5 length of shaft.

(LATICORN SECTION)

(Myzorhynchus series, p. 29)......- 5

*kyondawensis known only from larval stage.

Harrison and Scanlon: Subgenus Anopheles in Thailand 29

3.(2) Setae 5,6, '7-C all well developed and feathered; seta 4-C branched from

basse eee eR ee ee ee (Anopheles series, p. 144)

aitkenit sp. group (p. 147)

Setae 5,6, 7-C reduced, some or all short and few-branched or simple;

seta 4-C simple or branched only on distal half. .......... 4

4.(3) Seta 6-VI simple or with 2 - 4 branches; seta 11-C shorter than antenna,

or if as long as antenna then plumose from base.

(Anopheles series, p. 144)

culiciformis sp. group (p. 176)

Seta 6-VI with more than 10 branches; seta 11-C longer than antenna,

simple or with 4 - 8 strong branches arising approximately midway.

(Lophoscelomyia series, p. 132)

asiaticus sp. group (p. 135)

5.(2) Abdomen with fully developed palmate setae on at least 5 segments. . 6

Abdomen without developed palmate setae, or with such setae only

on 2 = 8 SE RMeNiei ya. UA eae se umbrosus sp. group (p. 105)

6.(5) Seta 1-P with long branches from near the base; seta 1 on segment II

. palmate and usually pigmented (except bavbumbrosus),.

barbirostris sp. group (p. 78)

Seta 1-P simple or with only short branches near the tip, less often

from about half way; seta 1 on segment II palmate and unpigmented.

7.(6) Seta 3-C bushy, divided from near the base into 40 or more branches;

seta 1-P simple or infrequently with 2-5 short branches on the distal

AA Beh eh hig hy gna aire hyrcanus sp. group (p. 36)

Seta 3-C not bushy, divided about halfway from the base into less than

15 branches; seta 1-P with 3 - 7 short, subequal branches arising

from the tip or about midway. . . . albotaeniatus sp. group (p. 31)

montanus (p. 34)

LATICORN SECTION

Reid and Knight 1961: 474.

This section was proposed for those species which exhibit complex trumpet

shapes. In general these species usually have the longest axis transverse to

the stem and a secondary cleft is often present, in addition to the meatal cleft.

The rim of the trumpet is often folded and may exhibit a series of sharp ridges

(spines) much like a file. In addition, there is often a complex lobe or tragus

protruding from the rim. This latter structure is always present in those

species in which the longest axis appears more vertical. For a comprehensive

discussion of the relationship of this section with the other sections and. sub-

genera of the genus refer to Reid and Knight (1961).

MY ZORHYNCHUS SERIES

Edwards 1932: 36.

In addition to the characters presented in the keys, members of this series

30 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

may also be recognized by a combination of the following characters.

ADULT (General). Basal 0.33 of forefemur always swollen; head scales

always broad; tarsi usually with pale bands; coxae often with scales; wings

nearly always with pale areas. Female. Palpi usually with erect scales,

giving shaggy appearance. Male. Claspette compact, with some setae on

dorsal lobe fused to form club; aedeagus leaflets usually present.

PUPA. Setai1, V - VII shorter than segments, usually branched, often as

long as seta 5, V - VII and sometimes tufted with many branches.

LARVA. Setal-Alarge, approximately 0.5or more length of antennal shaft, with

branches from the base; 3-C often with many branches; 4-C branching from

base; 5,6, 7-C long with numerous branches; 11-C approximately equal to

antennal shaft in length with many branches from base; 11-P short, stout,

with short branches from base; seta 6 longer on segments 1 - III than on IV -

VI.

The males of the respective species groups in this series are more difficult

to define than the females; consequently, they have been frequently misidentified

in the past. The following key should identify most of the specimens encountered.

KEY TO THE MALES OF THE SPECIES GROUPS OF THE

MYZORHYNCHUS SERIES IN THAILAND.

i. Palpi with extensive pale scales, either on mesal sides of segments

2 - 3, base of segment 3 or on segments 4 - 5 (club). ....... 2

Palpi dark scaled, or with very narrow pale bands (usually on segments

ee ee eg ah ae ek ee wha eet 3

2.(1) Remigium and base of veins R and Cu completely dark scaled.

separatus (p. 128)

Remigium and base of veins R and Cu with at least a few scattered pale

re ee, Lie ihe PE. ee aS hyrcanus sp. group (p. 36)

3.(1) *Hind tarsomere 5 entirely pale scaled.

_albotaeniatus sp. group (p. 31)

montanus (p. 34)

Hind tarcomerc 5. mostly Gare sca ed) ane ie eek es ies Oh eo 4

4.(3) Basal half of costa usually with scattered pale scales; vein Cug usually

with fringe spot; pleuron, coxae and usually abdominal sternites with

some pale scales, veins Rg-Ro 43 and M entirely dark scaled to fork.

barbirostris sp. group (p. 78)

Basal half of costa dark scaled; vein Cuy without fringe spot; pleuron

and coxae usually, and abdominal sternites always without pale

scales. Usually vein Ro-R9+3 or vein M with some pale scales prior

to fork (except separatus)........ umbrosus sp. group (p. 105)

DISTRIBUTION. This series is confined to the Eastern Hemisphere and

appears most heavily concentrated in the Oriental faunal region. Reid (1968)

*If hind tarsomere 5 missing, then montanus has: prehumeral and humeral

pale spots on base of costa, no pale fringe spot at vein Cug, and pleuron, coxae

and abdominal sternites without pale scales.

Harrison and Scanlon: Subgenus Anopheles in Thailand 31

points out that over 40% of the anophelines occurring in Malaysia belong to this

series. In Thailand approximately 36% (21) of the known anophelines belong in

this series. These are distributed among the albotaeniatus, barbirostris,

hyrcanus and umbrosus species groups. The single member of the albotaeniatus

species group, montanus, is rather rare, while most of the members of the um-

brosus species group seem to be limited to southern Thailand, or areas with

tropical wet forests. The barbirostris and hyrcanus species groups have some

widely distributed members and others with limited distributions.

TAXONOMIC DISCUSSION. All of the species recognized from Thailand are

usually large except pursati, and all are somewhat dark with a definite anopheline

habitus (spotted wings) and resting attitude. One species not currently recog-

nized from Thailand (brevipalpis Roper), was suggested (Reid 1968) as possibly

occurring in Thailand. This species is rather unique to the series, for the rest-

ing attitude is almost culicine and it has shortened palps and wings without spots.

Workers in southern Thailand should be on the lookout for this species. It is

easily separated from the other unicolorous anophelines (Anopheles series) by

the presence of scales on the anterior pronotum, a character not found in the

culiciformis and aitkenii species groups.

ANOPHELES ALBOTAENIATUS SPECIES GROUP

(albotaeniatus species group, Reid and Knight, 1961)

Anopheles albotaeniatus (Theobald) 1903b.

GENERAL. Posterior margin of wing without pale fringe spot; hindtarsi with

broad pale bands.

FEMALE. Palpi black and shaggy; sternum VII without scale tuft, but may

have few minute, scattered scales.

PUPA. Terga with small denticles on posterior margin (pupae of only 2

species described).

LARVA. Fully developed palmate seta on abdominal segments III - VII;

seta 3-C with 30 or less branches.

DISTRIBUTION. The albotaeniatus group as currently defined includes 6

species and is thought to be restricted to the Southeast Asian faunal region.

However, saperoi Bohart and Ingram, described from Okinawa in the northern

half of the Ryukyus, may extend into the Palaearctic region. Another species

ohamai Ohama, is known only from Ishigaki Island in the southern Ryukyus.

Two other species, balarensis Mendoza and ejercitoi Mendoza, are known only

from adults collected on Luzon Island, Philippines. The 2 remaining species,

albotaeniatus (Theobald) and montanus, are both recorded from the Malay

Peninsulaand Thailand. Iyengar (1953) recorded albotaeniatus from southern

Thailand; however, subsequent collections made by Harinasuta et al. (1964) and

SEATO Medical Research Laboratory personnel including the authors, during

the period 1961-1971 in most of southern Thailand, failed to yield specimens of

this species. Consequently, we have dropped albotaeniatus from the Thailand

records until confirming specimens can be produced. The distribution of albo-

taeniatus, sensu stricto, includes: Sulawesi (van Hell 1952), Sumatra, Java,

Borneo and Peninsular Malaysia (where Reid 1968, says it is uncommon).

This is apparently another species belonging to the Malay-Indonesian faunal

group (see Table 1, p. 7) that extends up onto the mainland via the Malay

Peninsula and then disappears shortly before reaching the Thai border or short-

ly thereafter. The one remaining species in the group, montanus, is found in

Thailand.

32 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

TAXONOMY. Within the group, 4 of the 6 species are poorly known, thus

most taxonomic discussion is necessarily based on the 2 better known species,

albotaeniatus and montanus. The adult females have been described for all the

species. Those of albotaeniatus, balerensis, ejercitoi and montanus are all

similar with: broad pale bands on hindtarsi and hindtarsomere 5 entirely pale;

narrow apical fringe spot on the wing; posterior margin of wing without pale

fringe spots; and sternum VII without tuft of scales. The female of ejercitoi

also has 2 very distinctive characters, lower mesepimeral setae and scattered

scales on the abdominal sterna. Both characters are not found on the other 5

members and are common only in the barbirostris subgroup of the barbirostris

species group. The females of the remaining 2 species, ohamai and saperoi,

do not seem closely related to the others, but are closely related to each other,

if not synonymous (Ohama 1947). Both exhibit: narrow hind tarsal bands with

hindtarsomere 5 partially dark scaled; wide apical pale fringe spot on the wing;

abdominal sternum VII with a tuft of scales; and the scutum with a pale central

area flanked by dark lateral areas.

Pupae have been described for only albotaeniatus and montanus and they

have one very important character in common, i.e., denticles on the hind mar-

gins of the abdominal terga. Otherwise, they differ in the basic axis and struc-

ture of the trumpet, and also seta 9 on the abdomen.

Larvae have been at least partially described for 4 species, albotaeniatus,

ohamai, montanus and saperoi. Those of albotaeniatus and montanus have a

number of characters in common: fully developed palmate setae (1) on seg-

ments III - VII; pale and partially developed palmate setae on segments I - II;

1-P single or branched distally; seta 3-C with few branches; and a small ven-

tral plate on abdominal segment VIII. Larvae of albotaeniatus also exhibit

widely separated setae 2-C a character unusual’in this subgenus, and seta

11-P with 3 - 4 fine branches from the base, unlike those found on montanus.

The larval stages of ohamai and saperoi are poorly known. Both apparently

have: pale and partially developed palmate seta 1 on I - II; fully developed pal-

mate setae on segments III - VII; seta 1-A, small and branched; seta 1-P

branched; and seta 4-C long and simple (saperoi) or long and 2 - 3 branched

(ohamat). The small branched 1-A and single 4-C seta on saperoi, and also

an egg without a frill, led Reid (1968) to imply some affinities to the lindesayi

species group. These characters are distinct from the groups in the Myzor-

hynchus series, and when associated with the adult differences, would lead one

to suspect saperoi (possibly ohamai also) are not members of the albotaeniatus

group.

In regards to the relationship of this group with other species groups in the

series, Reid (1968) said, ''The group is intermediate between the barbirostris

and umbrosus groups; the larvae have a full complement of palmate hairs as in

the barbirostris group, but resemble the umbrosus group in lacking a fringe

spot on the hind margin of the wing at the end of vein 5.2. The pupae of only

two species (albotaeniatus and montanus) are known and they resemble those of

the umbrosus group in having teeth on the hind margins of the tergites.'' We

feel that other characters and considerations also need examination to ascertain

the affinities of this group.

ADULTS. The females are definitely close to the umbrosus group in that

they lack fringe spots on the hind margin of the wing, scales on the pleuron and

coxae and a scale tuft on abdominal sternum VII (except saperoi-ohamai).

Anopheles ejercitoi also exhibits 2 characters (see above) showing affinities to

the barbirostris subgroup of the barbirostris group. Adults of ohamai and

saperoi exhibit: a tuft of scales on abdominal sternum VII and a wide apical

Harrison and Scanlon: Subgenus Anopheles in Thailand 33

fringe spot on the wing, showing affinities to the hyrcanus group and the vanus

subgroup of the barbirostris group; and some affinity to the lindesayi group

and to some members of the umbrosus group by the pale central area of the

scutum that is flanked by dark lateral areas. Reid and Knight (1961) placed

considerable weight on the ornamentation of legs in their classification of the

subgenus Anopheles. If such characters do reflect phyletic relationships,

then the broad hind tarsal banding in this group suggests a close relationship

to the hyrcanus and African coustani* groups.

PUPAE. The tergal denticles found on the pupae of albotaeniatus and mon-

tanus are otherwise found only in the umbrosus group. The pupa of albotaenia-

tus has 3 additional characters much like the umbrosus group; a trumpet with

the longest axis vertical, a large lobe (tragus) on the rim, and short, rounded

seta 9. The pupa of monianus; however, has a trumpet with the longest axis

transverse to the stem, a deep secondary cleft in the trumpet rim and long

setae 9, and all characters showing affinities to the hyrcanus and barbirostris

eroups.

LARVAE. All larvae currently described (albotaeniatus, ohamai, montanus

and saperoi) have well developed palmate setae on segments III - VII and pale,

poorly developed palmate setae on segments I - II. In the Southeast Asian

Myzorhynchus series, this combination is found only in the hyrcanus group

and the vanus subgroup of the barbirostris group. Members of the barbirostris

subgroup of the barbirostris group have well developed and pigmented palmate

setae on at least segments II - VII. Setae 1-P of albotaeniatus and montanus

are like those of the hyrcanus and umbrosus groups and unlike those found in

the barbirostris group, which are always branched from the base. In addition,

the ventral plate found on sternum VIII of these 2 species is a distinct umbrosus -

like character. The setae 3-C of ohamai and saperoi have few branches, much

like the umbrosus group and the vanus subgroup of the barbirostris group.

A clear interpretation of the relationship of this group to the other Southeast

Asian members of the Myzorhynchus series is not possible and will remain thus

until more is learned about the 4 imperfectly known members and the distribu-

tion of the group. Based on the known members, we can say: 1) the group

may not be a natural assemblage; 2) the group seems most closely related to

the wmbrosus group; 3) there are some affinities to the hyrcanus group and

vanus subgroup of the barbirostris group; and 4) there is very little evidence

for a relationship to the barbirostris subgroup of the barbirostris species

group.

BIOLOGY. The members of this group are primarily forest-swamp forest

dwellers, which rarely attack man. Some members have been observed biting

man during the day under forest shade.

MEDICAL SIGNIFICANCE. None of the species are involved in the trans-

mission of malaria parasites. The report (Iyengar 1953) of albotaeniatus asa

vector of Brugia malayi (Brug) requires confirmation.

*Reid and Knight (1961) pointed out this apparent relationship with

obscurus (Grunberg); however, note the unique egg of this species (Gillies

and de Meillon 1968) compared to Southeast Asian Myzorhynchus members

(Reid 1968).

34 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

ANOPHELES (ANOPHELES) MONTANUS STANTON AND HACKER

(Figures 5, 6, 7)

Anopheles albotaeniatus var. montanus Stanton and Hacker 1917: 273 (“, 2*, L)

Anopheles montanus Stanton and Hacker, Christophers 1924: 30.

Anopheles peditaeniatus of Walch 1930: A4 (L*); Stoker 1931: 129 (L*) (= montanus).

Anopheles (Anopheles) montanus Stanton and Hacker, Edwards 1932: 42; Reid

1950: 312; Reid 1968: 148 (o*, 9*, P*, L*, E*).

Adults may be recognized by a combination of unbanded palps, abdominal

sternum VII without a scale tuft and hindtarsomeres with wide pale bands, par-

ticularly tarsomere 5 which is entirely pale. The pupa is easily recognized by

a deep secondary cleft in the trumpet rim, denticles on the hind margins of the

terga, and long setae 9 on the abdominal segments. The larvae resemble mem-

bers of the hyrcanus group, but have 1 - 12 branches on seta 3-C, seta 1-P with

3 - 7 branches on the distal half and a small plate on abdominal sternum VIII.

FEMALE (Fig. 5) Head. Central vertex scales pale, lateral scales dark,

white frontal bristles between eyes; pedicel deep brown without scales, 1st

flagellomere only with dark scales; palpus dark scaled, basal scales somewhat

erect, those toward apex appressed; proposcis about as long as palp, dark

scaled; labellum paler than labium. Thorax. Anterior promontory with small

central patch of pale scales and few lateral dark scales; scutum without scales,

with numerous pale setae and mottled integument, not divisible into distinct

pale and dark areas; anterior pronotum with strong tuft of dark scales and

setae; posterior pronotum without scales or setae. Pleural setae: 2 - 3 pro-

pleural, 4 - 6 spiracular, 12 - 15 prealar, 3 - 4 upper and 5 - 6 lower sterno-

pleural, and 5 - 6 upper mesepimeral, pleuron without white scales, often

with few cryptic opaque scales on postspiracular area. Wing. Costa normally

with 4 pale spots; prehumeral, humeral, subcostal and preapical; prehumeral

and humeral spots may be absent, or just a trace of these on one or both wings;

preapical fringe spot narrow, beginning before and extending down to tip of Ry;

remigium dark scaled; humeral crossvein with tuft of black scales; vein R-Rj

usually with distinct pale spots at sector and apical positions, but without pale

subcostal spot; apical fringe spot narrow, from Rg to R445 or infrequently

beyond; bases of veins R, Cu and 1A usually with few scattered pale scales.

(Also see taxonomic discussion.) Legs. Coxae without scales. Foreleg:

femur dark and swollen basally; tibia dark; tarsus with narrow pale bands at

joints of tarsomeres 1 - 2, 2 - 3, and at apex of 3. Midleg: femur and tibia

dark without pale knee spot; tarsomeres 1 - 3 and occasionally 4 with very nar-

row, apical pale bands. Hindleg: femur dark, with minute knee spot; tibia with

distinct basal and narrow apical pale bands; tarsomeres with narrow apical

pale bands on 1 - 3 bases of 2 - 3 narrowly pale, 4 with broader basal and

apical bands, and 5 entirely white. Abdomen. Dark, without scales, but

with numerous pale yellow setae.

MALE (Fig. 5). Resembles female in general appearance, but with more

extensive pale scaling on wings and legs. Head. Palpi entirely dark scaled.

Wing. Base of costa with at least well defined prehumeral and humeral pale

spots; infrequently, entirely pale proximal to humeral crossvein; numerous

pale scales on bases of veins R, Cu and 1A; vein Cu to fork commonly with

only 1 - 3 black scales. Legs. Pale hind tarsal bands markedly broader than

on female. Genitalia. Parabasal spines arising from a marked prominence,

outer spine longer, more curved and with larger base; internal mesal spine

Harrison and Scanlon: Subgenus Anopheles in Thailand 35

some distance from apex of the basimere; claspette bilobed, dorsal lobe with

club, ventral lobe usually with 2 unequal setae, both relatively strong but, 3rd

seta sometimes present; aedeagus leaflets 3 - 4 pairs, larger leaflets with few

denticles near tip; proctiger with lateral sclerotization.

PUPA (Fig. 6) Cephalothorax. Wing case with variably pigmented pattern

of dark spots and bars. Trumpet. Meatal cleft and secondary cleft very deep,

permitting trumpet to open widely in life. Abdomen. Seta 9 peg-like on II - III,

long and pointed and faintly pigmented on IV - VII; seta 5 with more branches

than seta 1 on most segments; seta 3-III inserted in line and between setae 1

and 5; terga of most segments with patches of small denticles, most obvious

on caudal margin of segments. Paddle. Index approximately 1.3; refractile

border 0.5 - 0.6 paddle length; seta 1-P with 2 - 4 distal branches.

LARVA (Fig. 7). Living larvae dark brown with abdominal segment III

pale. Head. Setae 2-C simple, bases close together, but not touching; 3-C

divided into 6 - 13 branches some distance from base; 4-C small, with 3 - 6

branches at tip. Antenna. Seta 1-A with about 12 branches, usually reaching

apex of antenna; 4-A prominent, with 9 - 12 branches. Thorax. Seta 1-P

small, with 3 - 7 branches near tip; 2-P with 8 - 10 branches; 3-P simple,

short; seta 3-T palmate, lightly pigmented, with branches fairly slender;

pleural setae simple; seta 11-P short, with 6 - 10 branches; 14-P very long,

with 9 - 12 branches. Abdomen. Sterna frequently with patches of small spic-

ules; seta 1, I - VII palmate, pale and poorly developed on I - II, well devel-

oped and darkly pigmented except for filament and apex of blade on III - VI;

well developed, but lightly pigmented on VII; seta 6-III with many branches

(26 - 33), seta 6, IV - V, 0.7 - 1.0 length of 6-III, with 1 - 3 branches;

setae 5, 9 large, with numerous branches on most segments; abdominal ster-

num VIII with small (sometimes bilobed) plate near cephalic margin; pecten

with 4 - 6 long teeth separated by many smaller teeth; seta 1-X simple.

TYPE-DATA. Syntypes of montanus from the type-locality, ''Ulu Gombak,

Malay States" are present in the British Museum (Natural History)| BMNH];

however, the specimens representing the type-series have not been defined.

Stanton and Hacker (1917) clearly described the ¢, “(including genitalia) and

larva of montanus in the original description. In the BMNH there are 6%, 4c

and 2 larval skins which were collected by Hacker in the years immediately

surrounding the original description (early 1917), and all agree well with that

description. These specimens, however, were received at the BMNH in 2

shipments. The 1st shipment was logged in the BMNH on 14 July 1916, has

accession #1916-168 and includes 12 and 1“ collected by Hacker in March 1916

at the type-locality. The 2nd accession was logged in the BMNH on Nov. 1920

and is numbered 1920-454. This accession includes 12 and 1“ (with prepared

genitalia mount) labeled ''Ulu Gombak, Malay States, Dr. H. P. Hacker, 1920-

454", and 429, 20° and 2 larval skins (12 and 1L. skin associated, #479F. 2)

labeled ''Malay States, Dr. H. P. Hacker, 1920-454.'' Conceivably, all the

above specimens could be syntypes, however, we feel the 2? and “ in accession

1916-168 should not be considered syntypes because larvae were not included

in that accession, and because that shipment was received at the BMNH over 6

months preceding the description of montanus. These 2 specimens (1916-168)

are almost certainly referred to by Stanton and Hacker (1917: 273) when they

say, ''We are indebted to Mr. F. W. Edwards of the British Museum who has

kindly examined our specimens and aided us in fixing the systematic position

of this Anopheline.'’' This statement implies that those 2 specimens were not

necessarily considered representative of a new variety when sent to Edwards.

Because of these circumstances, the specimens received at the BMNH in 1916

36 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

were probably not used by Stanton and Hacker (in Malaya) for their varietal

description, while those in accession 1920-454 are almost certainly the speci-

mens used for that purpose and should be considered the syntypes of montanus.

Possibly correspondence exists at the BMNH which could resolve this problem.

DISTRIBUTION. In THAILAND, restricted to the southern peninsular pro-

vinces where tropical wet forests are common. Specimens (10°, 6%, 47 larvae,

and 33 larval and pupal skins) were examined from the following provinces:

Narathiwat, Phuket, Ranong, Songkhla and Trang. The authors also examined

5c, 69, 4 larvae and 6 larval and pupal skins from MALAYSIA (Peninsular),

the majority originally collected and identified by J. A. Reid. An additional

60°, 99, 4 larvae and 2 larval skin specimens (including syntypes) in the BMNH

were examined from: MALAYSIA (Peninsular and Malaysia); and THAILAND

(Trang). There is also a literature record of this species from INDONESIA

(Banka) (Reid 1968). Possibly montanus occurs over much of the Indonesian

archipelago, at least in those areas with climatic conditions matching those of

southern Thailand and Malaya (zone Afi in the K6ppen classification). It is

also likely that montanus occurs in southern Burma, since Ranong Province

borders Burma and the tropical wet forests extend north into Burma.

TAXONOMIC DISCUSSION. The species most apt to be confused with

montanus is roperi, which has a similar distribution in Thailand. This spe-

cies is often very like montanus (pale spots on base of costa and basal pale band

on hindtibia), but hindtarsomere 5 is mostly black, while that of montanus is

entirely white. Specimens missing the hindtarsi would be very confusing, but

should be separable on the basis of 0 - 1 propleural setae on voperi while mon-

tanus has 2 - 4. The pupa of montanus, with tergal denticles, long spine-like

abdominal seta 9 and unique trumpet, is quite distinct and should not present

identification problems. The larva is also rather distinct. The combination

of well developed palmate setae, a simple or distally branched seta 1-P and

the few branches on seta 3-C should readily separate this species.

One melanistic female (with associated skins) was examined from Narathi-

wat Province. This specimen lacks the prehumeral, humeral and subcostal

pale spots, in essence the entire costa is dark scaled. The preapical pale

fringe spot just before the tip of Rj is also missing, while a faint pale apical

fringe spot is present at R,. The remainder of the wing is fairly normal ex-

cept for the absence of a sector spot on vein R,. No abnormalties were noted

elsewhere on the specimen.

BIOLOGY. Sandosham (1959) reported montanus breeds in pools at the

sides of jungle streams, particularly those with decaying leaves. Reid (1968)

refers to shaded pools or small swamps at the sides of forest streams up to an

elevation of about 305 m. Thailand specimens were collected from elephant

footprints, stream pools and a rock pool, all adjacent to small streams in wet

forest and between 76 - 305 m elevation. These collection sites all received

partial to heavy shade and all but one contained many decaying leaves. None of

the Thai specimens were from human biting collections, although montanus is

reported to feed on man in hill forest in Malaya (Traub 1957). In view of its

usual scarcity and restriction to jungle habitats it is unlikely to serve as a vec-

tor of human disease pathogens.

ANOPHELES HYRCANUS SPECIES GROUP OF SOUTHEAST ASIA

(An. hyrcanus species group, Reid 1953)

(Figures 8, 9)

Anopheles hyrcanus (Pallas) 1771; as Culex hyrcanus.

Harrison and Scanlon: Subgenus Anopheles in Thailand 37

GENERAL. Palpi with pale bands; hindtarsi with narrow to broad pale

bands; tertiary fringe scales on wing pale, at least paler than other fringe

scales (except spots).

FEMALE dead. Palpi with pale bands; clypeus with tuft of outstanding dark

scales on each side; pedicel with small pale scales on outer aspect; basal 4 - 8

flagellomeres with pale scales. Abdomen. Without scales except ventral tuft

on sternum VII.

MALE Head. Palpi with pale bands or spots on segments 4, 5 and sometimes

segment 3, and with scattered pale scales elsewhere; clypeus without scales;

flagellum generally without scales except for few pale scales on flagellomere 1.

Abdomen. Without seales except on the basimere of genitalia. Genztalia.

Claspette bilobed; ventral lobe with 2, infrequently 3 large setae, mesal seta

very long, lateral seta shorter; dorsal lobe with club-like spine on inner half

fused from 2 - 4 separate basal stems, and normally 2 thin leaflet-like setae

on outer half immediately adjacent to fused club; aedeagus with 2 - 7 pairs of

leaflets, at least larger ones with serrations and approximately 0.33 length of

aedeagus.

PUPA Cephalothorax. Palpal case exhibits sexual dimorphism, case tip

of female short and blunt, that of male long and attenuated. Tvumpet. Longest

axis transverse to stem; outer wall of pinna may exhibit vertical wrinkles as in

argyvopus; rim without secondary cleft or lobe (tragus), but may have thick-

ened saw-tooth edge. Abdomen. Terga without denticles on caudal margin;

seta 1-VII with 13 or less branches. |

LARVA Head. 3-C large, with 40 or more branches from near base;

1-A with numerous branches, reaching or exceeding tip of antenna. Thorax.

1-P short, simple or with several small branches on distal half; prothoracic

pleural tubercle with long stout spine. Abdomen. Seta 1 palmate onI - VII,

but pigmented and well developed only on III - VII. |

DISTRIBUTION. The Southeast Asian members of this group are generally

distributed throughout the Oriental faunal region. The Southeast Asian species

described here are closely related to hyrcanus (Pallas) and other described

Palaearctic forms which occur from Spain eastward through the Mediterranean

basin, the Middle East, USSR and possibly Palaearctic Asia. The geographic

point of separation for these two parts of the hyrcanus group in the west appar-

ently runs north from the Indian Ocean through the mountains and desert of

Baluchistan up to the Afghanistan peaks and east to the Himalaya Mountains.

The line of separation in Palaearctic Asia is still undetermined. The South-

east Asian members are not known from the Australian region. The eastern-

most records of this group in Indonesia come from collections on Timor Island

in the Sunda Chain, and-Buru Island in the Moluccas (Bonne-Wepster and Swel-

lengrebel 1953). The nominate species, hyrcanus, in the Palaearctic, appears

to exist in a number of forms which are currently synonymized with hyrcanus,

but which may, in fact, be good species. None of these forms appear to be

synonymous with the Southeast Asian members.

The oriental segment of this species group involves at least 13 species.

Eight species are currently known from the Thailand-Malaysia area: argyro-

pus, crawfordi, lesteri, nigerrimus, nitidus, paraliae, peditaeniatus, pursati

and sinensis. One species, pseudosinensis Baisas, is known only from the

Philippines, and at least 4 additional species are recorded from China, Korea

and Japan.

Only 2 species considered here, nigerrimus and peditaeniatus, have dis-

tributions which are known to extend west of Assam into peninsular India and

down to Sri Lanka. Anopheles peditaeniatus possibly extends as far west as

38 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Pakistan. In Southeast Asia these 2 species are found throughout the Indo-

China peninsula down through the Malay Peninsula and on all the major islands

in the Sunda Chain of Indonesia. Anopheles nigerrimus is recorded as far east

as Halmahera Island in the Moluccas, but does not extend north of Borneo into the

Philippines, where it is apparently replaced by pseudosinensis. This latter

Species, despite the name, seems to have closest affinities to nigerrimus.

Aside from the wide distribution in Indonesia, peditaeniatus is also found on

all the major islands in the Philippines.

Three other species, argyropus, crawfordi and sinensis have distributions

with: the most western extension in Assam; the most southern extension in

Sumatra (cvawfordi and sinensis) and Java (argyropus); and most northeastern

extension in Vietnam (argyropus and crawfordi) and China, Japan, Korea

(sinensis).

The 3 remaining species, lesteri pavaliae, nitidus and pursati, have some-

what different distributions. Althoughlestevi lesteri Baisas and Hu is recognized

from China, Philippines, Okinawa and Japan, subspecies paraliae is known only

from Borneo, Malaya, Thailandand South Vietnam. Thesetwo subspecies canbe

separated on consistent differences only in the adult stage, and no intermediates

have beenfound. Inreality pavaliae may deserve specific status, but this decision

must be based ona study of more material of both subspecies. Anopheles nitidusis

known from India (Assam) to North Vietnam in the east and Indonesia (Sumatra)

in the south. Anopheles pursati is apparently confined to the Indo-China Penin-

sula, with records and specimens from Cambodia, Peninsular Malaysia, Thai-

land and South Vietnam.

TAXONOMIC DISCUSSION. This is an extremely complex assemblage of

species which has been treated adequately only in Malaysia (Reid 1953 and

1968), the Philippines (Baisas and Hu 1936), Japan (Otsuru and Ohmori 1960)

and to some extent mainland China (Ho et al. 1962, Feng 1964; and Ma 1964,

1968a,b,c). The material examined from Thailand during this study agreed

very well with the Malaysian members described in detail by Reid (1953,

1968). Minor exceptions encountered are discussed under the individual species.

The discovery of the various species of this group in Malaya by Reid (1953),

was aided by detailed observations on biology. Consistent biological differences

and small but consistent differences in morphology of the various stages make it

clear that the species of this group form a sibling series, in the sense of Mayr

(1969). Due to the close relationship of these species it is often difficult or

impossible to identify a particular adult specimen, unless it is accompanied by

the associated immature skins. This may pose an initial difficulty for public

health workers, but continued work in a particular area should permit a rapid

understanding of the limits of variation of the local species. Despite Reid's

reservations as to the utility of his keys for areas outside Malaya, it appears

that the Thailand fauna at least, and possibly that of Indochina, consist primari-

ly of the same members of the hyrcanus group as seen by him in Malaya.

Additional collecting at higher elevations in northern Thailand may produce

some of the species described from China. Collectors in that area should be

alert for lesteri lesteri which is found in southern China and easily confused

with sinensis.

The interpretation that is followed here for intragroup affinities is based

on characters from all life stages (Table 2) and recognizes 2 subgroups (Har-

rison 1972). The nigerrimus subgroup was created to include nigerrimus,

nitidus (as indiensis), pursati, and pseudosinensis, while the lesteri subgroup

includes lesteri lesteri, lesteri pavaliae, crawfordi and peditaeniatus. The 2

remaining species, avgyropus and sinensis are isolated. This interpretation

Harrison and Scanlon: Subgenus Anopheles in Thailand

a1eq IO Sse] Ajalerl ‘Sales [[ewuISs 9-¢ ynq ‘Yo ed & ATTeN{IOe JONxx

eULYD UL WALOJ O1}OIVETeg uO JUeNbeTj 9IOW ‘vISY ISBIYNOS UI VIBYy

9] e.19poUul eae ; sniniuaniipadg

MOLIBU 394300

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499} ainjont}s UI@ASSO19 eISOD © ¢ ‘SOS [ed SsjaTjvo]

Y}pIM ua 100d 9-8 BIeS A-G ®19S UII Ie1euny joosequo -j[ed » uo snsvapsee Ssotoveds

yooq | 3uo] ‘ON UO Sayouerg | Uo SoyoueIg joduini I, UO S8TeVIS S9TVOS sTeq puURq xIeq Jo SJIeg

J0"e VAUV'I Vdd LTINdV

dnois satseds snuvorcy ueisy ISBaYINOS sy} UI SOTITUTTJe TaoeVIeYD °7Z AIAGVL

40 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

differs from Reid's (1953, 1963, 1968) interpretation by: (1) separating pedi-

taeniatus from argyropus and placing it in the lestevi subgroup; (2) determining

argyropus is separate, but more closely related to nigerrimus than peditaenia-

tus; (3) moving sinensis away from a close relationship with n7gerrimus and

placing it separate, but possibly intermediate between the nigerrimus and les-

tevi subgroups; and (4) removing nitidus from an intermediate position between

crawfordi and nigerrimus and placing it in the nigerrimus subgroup.

Members of this group show more affinity to the barbirostris group than to

the albotaeniatus or umbrosus groups. Adult females exhibit a tuft of scales

on sternum VII, scales on the pleuron and coxae (except /esteri) and most have

fringe spots on the posterior margin of the wing. These are characters found

elsewhere in the series only in the barbirostris and bancrofti groups. The lar-

vae have seta 3-C with many branches and palmate seta 1 on segments I - VII,

as do the larvae of the barbirostris group. The pupae of the hyrcanus group all

lack denticles on the tergites as do members of the barbirostris group. In ad-

dition, the pupal trumpets of the hyrcanus and barbirostris groups are similar

in structure and less complicated than those found in the albotaeniatus and um-

brosus groups.

BIOLOGY. The immature forms of the hyrcanus group are found primarily

in rice fields, marshy areas, ponds with emergent vegetation, and similar habi-

tats. The great rice producing areas surrounding Bangkok and in the valleys of

the north, such as Chiang Mai, support large populations of these species. The

more abundant species, such as peditaeniatus and sinensis are found essentially

throughout the country, even in mountainous areas where small paddy fields are

found. These 2 species are very common in the rice field habitat throughout

their distribution in the Orient, and may well owe their wide distributions to

man's long association and dependence on rice cultivation.

Fortunately, members of this group seem primarily zoophilic; although,

human biting incidence may be quite high under a given set of ecological conditions.

MEDICAL SIGNIFICANCE. There is no evidence that any of these species

transmit malaria parasites to man in Thailand. There have been repeated in-

criminations of sinensis as a major vector of malaria parasites in China; how-

ever, Ho et al. (1962) found that lestevi lesteri is the primary vector in the

Yangtze Valley and that sinensis plays a secondary vector role. There is some

possibility (Iyengar 1953) that members of the group serve as vectors of Brugia

malayi in southern Thailand, but again the identity of the mosquito and parasite

species needs confirmation. Workers interested in filarial transmission by the

hyrcanus group should consult the review of this subject by Reid et al. (1962).

KEYS TO THE SPECIES OF THE ANOPHELES HYRCANUS

GROUP IN THAILAND

FEMALES

1, Hindtarsi with broad pale bands, at least one tarsomere (4) with a basal

PAE OOO ALOR ee Ss ee ee ee ee ee ea

Hindtarsi witiapiced! pate bands only. ... 00.0. ee ee wh ee e's 9)

*Infrequent lesteri pavaliae have a small basal patch of pale scales on tarso-

meres 4 - 5, but these can be properly assigned by the narrow pale fringe

spot at the wing apex.

Harrison and Scanlon: Subgenus Anopheles in Thailand 41

2.(1) Basal dark mark on wing vein Cu short, separated by own length or more

from upper dark mark on vein 1A; base of costa with scattered pale

scales, frequently with small humeral pale spot; dark mark at origin

of vein Rg well defined, scales between dark mark and fork mostly

white; vein Cug always with pale fringe spot..... nitidus (p. 58)

Basal dark mark on wing vein Cu long, approaching within own length or

less of the upper dark mark on vein 1A; base of costa dark scaled or

with few scattered pale scales, but no pale humeral spot; dark mark

at origin of vein Rg poorly defined, scales between dark mark and

fork mostly dark; vein Cug with or without pale fringe spot. .... 3

3.(2) Humeral crossvein with patch of dark scales; remigium mostly dark

scaled; basal 0.33 and preapical dark mark on vein R-R, dark scaled,

OY Wilh Very Tew DAle BCAIOS ee er eee oe oom ~~ ee ee 4

Humeral crossvein without scales; remigium mostly pale scaled; basal

0.33 and preapical dark mark on vein R-R, usually with many pale

SOAR Oe ig I i ERS ET cae peditaeniatus (p. 71)

4,(3) Hindtarsomere 4 with broad dark band, at least 0.6 length of segment;

hindtarsomere 5 without or with very narrow (0.25 or less of segment)

pale basal band; base of costa usually with several scattered pale

scales; apical dark mark on vein Cug short, rarely as long as apical

Gain VaR On BA pL Ae ee a We a nigerrimus, in part (p. 65)

Hindtarsomere 4 with narrow dark band, 0.5 or less (lacking in rare

cases) length of segment; hindtarsomere 5 with pale basal band on 40%

or more of segment; base of costa without pale scales; apical dark

mark on vein Cug long, usually equal or longer than apical dark mark

Cree RLe I en hon ais, Be ee GR argyropus (p. 50)

5.(1) Wing apex with narrow pale fringe spot, not extending beyond R, to Rg.

7 6

Wing apex with wide pale fringe spot, extending at least to R fines as ‘

6.(5) Humeral crossvein with dark scales; remigium mostly pale scaled;

midcoxa with white scales; smallest member of group in Thailand.

pursati (p. 75)

Humeral crossvein without scales; remigium mostly dark scaled; mid-

coxa without pale scales, 60.08 we... lesteri paraliae (p. 61)

7.(5) Vein Cu with long basal dark mark*, at least twice as long as most

basal pale mark (when present) on Cu, and approaching within own

length or less of upper dark mark on 1A; costa usually with several

scattered pale scales on basal 0.33; remigium mostly dark scaled;

humeral crossvein with dense patch of black scales.

nigerrimus, in part (p. 65)

Vein Cu with short basal dark mark, equal to or less length of most

basal pale mark on Cu, and (usually) separated by own length or more

from upper dark mark on 1A; basal 0.33 of costa usually entirely dark

scaled; remigium mostly pale scaled; humeral crossvein bare or with

be BOSE 1 CLE Sc SLUR A aR A aera RON a eno MRE MLO Vy ese EN A 8

*Basal dark mark does not refer to a few dark scales or small dark spot

which may occur at the absolute base of the vein.

42 Contrib. Amer, Ent. dnst.; vol. 12, no. 1, 1975

8.(7) Wing pattern blurred; tip of vein R, dark scaled; apical fringe spot long,

beginning at or above vein R,; preapical dark mark on vein R, with

some pale scales; usually pale fringe spot at tip of vein Cug.

sinensis (p. 45)

Wing pattern sharp, dark marks short and well defined; tip of vein Ry

pale scaled; apical fringe spot shorter, beginning at R,; preapical

dark mark on vein Ry without pale scales; without pale fringe spot at

tS OF ein Cig al we a as A ee Go crawfordi (p. 54)

MA LES

di, Palpal segment 3 with basal white band. (nigerrimus subgroup)... . 2

Palpal segment 3 without distinct white basal band although some

pale scales may occur on the inner surface of each palp....... 4

2.(1) Wing with narrow apical pale fringe spot, at widest extending from R,

to Rg; hindtarsomere 4 rarely with a basal pale band.

bursati (p. 75)

Wing with wide apical pale fringe spot, extending at least to R4,5; hind-

tarsomere 4 with basal pale band (some specimens of nigerrimus

wiliiachobasal bandssonthe hindtarsi). . sos. ww eee iw ke 3

3.(2) Wing with long basal dark mark on vein Cu approximately its own length

or less from upper dark mark on vein 1A; base of costa dark scaled

or with several scattered pale scales, rarely with small humeral

pale spot; foretarsomere 2 with pale band seldom more than 0.25

leneth of seoment: 4. 00ha owas a Ses nigerrimus (p. 65)

Wing with short basal dark mark on Cu more than its own length from

upper dark mark on vein 1A; base of costa nearly always with pale

humeral spot, at least with numerous scattered pale scales; foretarso-

mere 2 with pale band usually 0.33 - 0.50 length of segment.

nitidus (p. 58)

4.(1) Hindtarsomeres 1 - 4 with narrow apical bands only.......... D

Hindtarsomeres with wide pale bands, with basal bands on tarsomeres

4 and usually 5, as well as apical bands on tarsomeres 1-4....7

5.(4) Wing with narrow apical pale fringe spot, at widest extending from Ry,

to Rg; midcoxa without pale scales; basimere without pale scales.

lesteri paraliae (p. 61)

Wing with wide apical fringe spot, extending at least to R4,5; midcoxa

with at least a few pale scales; basimere with pale scales. .... 6

6.(5) Wing vein R, with tip pale scaled; propleural setae 2 - 5; lower sterno-

pleutal Scie oe 98s)! ie aohes Shea Gas sree crawfordi (p. 54)

Tip of vein Ry dark scaled; propleural setae 4 - 10, lower sternopleural

Setaee Oe ea a... Pages cos ds das sinensis (p. 45)

7.(4) Humeral crossvein with patch of black scales; remigium mostly dark

scaled; preapical dark mark on Ry without pale scales.

argyropus (p. 50)

Humeral crossvein without scales; remigium pale scaled, scales on

anterior margin usually silvery white; preapical dark mark on R

with some pale scales (usually many). .... peditaeniatus, (p. 71)

Harrison and Scanlon: Subgenus Anopheles in Thailand 43

PUPAE

1. Seta 9-VIII without branches or with reduced branching; paddle with

marginal teeth extending beyond 0.75 its length........... 2

Seta 9-VIII with well developed branches; paddle with marginal teeth

rarély:reachine Oy Maite teneths won! 2aulite 6 e-em ee eee ws 3

2.(1) Trumpet with parallel vertical wrinkles on sides, and thin uniform rim;

seta 1-V with 17 - 40 branches. ......... argyropus (p. 50)

Trumpet without wrinkles on sides, but with thickened saw-toothed areas

on the rim; seta 1-V with 1 - 6 branches. . . . peditaeniatus (p. 71)

3. (dL) Prumpet vii one MAORI es ire a eo wk se ee

Trumpet rim with thickened, saw-toothed areas. .......... 7

4.(3) Seta 5-V with 9 - 24 branches; wing case pale, with darkest pigmenta-

tion in form of distinct rows of round dark spots*, infrequently with

lighter pigmentation on veins or crossbars. .... sinensis (p. 45)

Seta 0-V tuft-like, with 30 - 60 branches; wing case completely pale to

darkly pigmented, darkest pigmentation widespread, or on veins or

crossbars, blurred spots may be spaced between veins and crossbars,

but they are no darker than other pigmentation. .......... 5)

5.(4) Integument very lightly pigmented, with abdomen unpigmented including

paddle and areas surrounding setal alveoli; apical joint and tip of

antennal case with distinct dark spots. ....... nitidus (p. 58)

Integument usually moderately to darkly pigmented, including paddle and

pigmented spots around setal alveoli, especially 2 on II - VII; pigmen-

tation of antennal case when present, no darker than other pigment

and not centined to-apical joint and tipi ie) eek etek wis

6.(5) Seta 1-VII with 3 - 13 branches, rarely less than 6; seta 6-VII anterior

and slightly mesal to seta 9-VII; seta 1-P with 3 or more branches;

light to moderate pigmentation, without dark median spots on terga

EES hy cau ae a aie, Petey Be es gee, Bowe pursati (p. 75)

Seta 1-VII with 1 - 7 branches, usually 2 - 4, more than 5 branches very

rare; seta 6-VII mesal to seta 9-VII; seta 1-P usually simple or bifid;

skin darkly pigmented, with dark median spots on terga III - VII.

nigerrimus (p. 65)

7.(3) **Seta 1-VI with 2 - 6 branches; seta 5-V with 30 - 45 branches; integu-

ment usually light to moderately pigmented. ... cvawfordi (p. 54)

Seta 1-VI with 5 - 10 branches; seta 5-V with 12 - 30 branches; integu-

ment usually very darkly pigmented. .... lesteri paraliae (p. 61)

*Some specimens of puysati have distinct dark spots on the wing case, but

these specimens will have very dense tuft-like seta 5-V with 45 or more

branches and seta 1-VII with 6 - 13 branches (rarely less), while sinensis

has only 1 - 5 branches on 1-VII.

**This couplet will work on about 90% of the specimens collected in Thailand,

thus reared adults with associated skins are the best means of identification.

4a. Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

LARVAE ?

ds Seta 4-M small, with sinuate* spreading branches arising close together

BL WAGC isin Oddo ai fe ake en eg Bina he peditaeniatus (p. 71)

Seta 4-M with stiffer more erect or straight branches arising along

central stem, or close together near base (argyropus and nitidus). . 2

2.(1) Seta 6-III usually with more than 20 branches; seta 6-I usually with more

than 21 branches; frontal seta 5-C with 17 or more branches.

sinensis (p. 45)

Seta 6-III rarely with more than 20 branches; seta 6-I usually with less

than 21 branches; frontal seta 5-C with 11 - 18 branches**.... 3

3.(2) Seta 8-C with 12 - 24 (rarely 11 on nitidus) branches. ........ 4

Bete 60 ttn = FE PAN. a ee eRe le ee ee ee 7

4.(3) Seta 14-P with 3 - 5, usually 3 - 4 branches; seta 1-M with 26 - 38

branches, usually, 28/0r more. 266s. a ea es pursati (p. 75)

Seta 14-P with 5 or more (rarely 5) branches; seta 1-M with less than

28 branches (rare specimens of nitidus with more). ........ 5

5. (4) Seta 5-IV with 2 - 4 branches, usually 3; seta 5-III with 4 - 8 branches.

nigerrimus (p. 65)

Seta 5-IV seldom with less than 5 branches; seta 5-III with 7 - 17

Dre a a re eS OE ie CWE Ta Ie tiaee ie | 6

6.(5) Seta 2-II with 8 - 14 branches; seta 5-V with 5 - 9 branches; seta 9-VI

with 5 - 9 branches; pecten seldom with more than 6 long teeth; seta

1-X weak, rarely longer than saddle. .... nitidus, in part (p. 58)

Seta 2-II with 6 - 9 branches; seta 5-V with 4 - 6 branches; seta 9-VI

with 4 - 5 branches; pecten usually with 7 - 8 long teeth; saddle seta

1-X strong; longer than saddle. .......... argyropus (p. 50)

7. (3) Seta 9-III with 10 - 16 branches; pecten seldom with more than 6 long

teeth; seta 13-IV with 6 - 12 short branches, approximately 0.5

levets Gb eeta) 1LO-T Ve. a ee eh nitidus, in part (p. 58)

Seta 9-III with less than 10 branches; pecten rarely with less than 7

long teeth; seta 13-IV with 4 - 9 long branches; approximately equal

to or slightly less length of seta 10-IV... 2... 2... epee wee 8

*Seta 4-M may appear slightly sinuate on infrequent specimens of argyropus

or nitidus; however, seta 8-C will have 11 - 24 branches (cf. peditaeniatus

4 - 9) and seta 9-C will have 6 - 16 (cf. peditaeniatus 3 - 7).

**Rare specimens of lesteri paraliae may key to sinensis, but seta 5-II on the

former species has 6 - 10 branches while sinensis has 9 - 20 (seldom less

than 11) branches.

Harrison and Scanlon: Subgenus Anopheles in Thailand 45

8.(7) *Seta 5-III with 6 - 10 branches; seta 1-X strong, longer than saddle.

lestevi paraliae (p. 61)

Seta 5-II seldom with less than 12 branches; seta 1-X usually weak and

shorter them Saddle@ew arc eae aie crawfordi (p. 54)

ANOPHELES (ANOPHELES) SINENSIS WIEDEMANN

(Figures 2, 8, 10, 11, 12)

Anopheles sinensis Wiedemann 1828: 547 (c, 9).

Anopheles plumiger Donitz 1901: 37 (%, 9).

Anopheles jesoensis Tsuzuki 1902: 764 (c).

Anopheles (Anopheles) hyrcanus var. sinensis of Edwards 1932: 41.

Anopheles (An.) sinensis Wiedemann, Reid 1953: 10 (“*, 2*, P*, L*, E*);

Reid 1968: 85 (“*, @*, P*, L*, E*); Harrison 1973a: 7 (lectotype).

Adult females can usually be recognized by: narrow apical tarsal bands,

short basal dark mark on vein Cu, wing with wide pale apical fringe spot,

remigium mostly pale scaled, humeral crossvein with scale patch or several

scales, preapical dark mark on Rj with few pale scales, apex of vein Ry with

dark scales and usually pale fringe spot at apex of vein Cuy. Adult males have

the above, but lack a basal pale band on palpal segment 3. The pupae have the

trumpet rim thin and uniform and have a distinct pattern of dark spots on the

wing case. The larvae are more difficult to distinguish: seta 6-I and III usu-

ally have more branches than those of the other species; the spiracles are

large; and setae 5, 6-C usually have more branches.

FEMALE (Figs. 2, 8, 10) Head. Vertex with pale erect scales above

interocular area, dark erect scales laterally; frontal tuft with long pale setae;

clypeus with tuft of dark scales on each side; pedicel with few small pale

scales on external aspect, and pale scales on most basal flagellomeres; pro-

boscis with dark scales, erect basally and appressed distally; labellum paler

than labium; palpus usually with narrow apical and basal pale bands on seg-

ment 5, additional pale bands or spots at segmental joints 3 - 4 and 2 - 3,

and a dorsomesal row of pale scales on segment 2. Thorax. Anterior prom-

ontory with long fine pale scales mesally, broader darker scales laterally;

scutal integument light brown usually with dark brown median line extending

back to prescutellar area, often with paired dark lateral lines on anterior

dorsocentral bare areas, with dark eyespots, scutum sparsely covered with

very fine seta-like curved silvery-yellow scales, and with longer stout golden

setae in anterior promontory, acrostichal, dorsocentral, lateral prescutal,

fossal, antealar and supraalar groups; scutellum with long golden setae and

small fine curved pale scales; anterior pronotum with patch of dark erect scales

on dorsocephalic aspect, setae caudally; pleural integument light brown or

eray, usually with darker areas in form of upper and lower dark lines, may

have several scales on lower sternopleuron; pleural setae: 4 - 10 propleural,

2-95 spiracular, 8 - 13 prealar, 5 - 6 upper and 5 - 9 lower sternopleural,

6 - 12 upper and 0 lower mesepimeral. Wing. Costa largely dark with sub-

costal and preapical pale spots and occasionally pale scales near humeral

*This couplet is valid for approximately 90% of the specimens collected in

Thailand, thus reared adults with associated skins are the best means of

positive identification.

46 Contrib. Amer. Ent. Inst., vol. 12, no. 1,. 1975

crossvein (see taxonomic discussion); subcostal pale spot includes costa, tip of

subcosta and R,; preapical pale spot includes costa, Ry and R,; remigium usually

pale on anteriot margin, posterior margin with proximal and Aistal dark scales;

humeral crossvein usually with 3 - 6 small dark scales, infrequently bare; base

of vein R-R, with variable scaling, usually with distinct presector and sector

pale areas, scattered pale scales between sector and subcostal pale spots and

between subcostal and preapical pale spots; tip of Ry with dark scales; Rg-

Ro, with basal dark mark, gradually paler to pale area just before fork; Ro

dark scaled except for pale spot equal to preapical pale spot on costa and Rj,

tip dark scaled; R, dark scaled on base and tip; R 406 largely pale, with apical

and basal dark spots with scattered dark scales in between; M usually dark

scaled on basal 0.5 or more, with pale area just before fork; M,,9 with small

dark spots at base and tip, may have scattered dark scales in between; M3,4

as for Mj,9; Cu with small dark spot at extreme base followed by pale spot,

then ''basal'"' dark mark equal to or less length of most basal pale mark on Cu,

and separated by own length or usually more from most basal dark spot on 1A;

Cu pale scaled distal to basal dark mark; Cu; with dark basal and apical scales,

in between usually with central vein scales pale and most lateral vein scales

dark; Cu, pale scaled except for dark scaled tip; 1A pale scaled except small

dark spot approximately midway along vein, and dark scaled tip; apical dark

spot on 1A usually longer than apical dark spot on Cug; apical pale fringe spot

extends from R, to slightly beyond Rqi5; caudal margin of wing usually with

pale fringe spot adjacent to Cug; tertiary fringe scales paler than secondary

and primary fringe scales except at pale fringe spots. Halter. Stem pale,

_knob dark scaled. Legs. Coxae with pale scales, midcoxa with 3 - 5 upper

setae and upper and lower patches of pale scales. Foreleg: femur usually

dark scaled dorsally and apically, with dirty yellow scales ventrally; tibia dark

dorsally, paler ventrally; tarsomeres dark scaled except paler scales on ven-

ter of tarsomere 1 and narrow apical pale band approximately equal segment

width on tarsomeres 1 - 3. Midleg: femur with dark brown scales on dorso-

anterior aspect, dirty yellow scales on ventroposterior aspect; tibia with dark

scales on dorsoanterior aspect, dirty yellow scales on ventroposterior aspect;

tarsomeres as on foreleg except apical pale scales more in form of dorsoapical

spots than true apical bands. Hindleg: femur dark scaled dorsally and apically,

with yellow scales ventrally except at apex; tibia as for midleg, and also with

few dorsoapical white scales; tarsomeres dark scaled except few yellow scales

on venter of tarsomere 1 and small dorsoapical white scale patches on tarso-

meres 1 - 4. Abdomen. Integument dark dorsally, paler ventrally, with long

golden setae, devoid of scales except median tuft of erect dark scales near cau-

dal margin of sternum VII.

MALE (Figs. 2, 10). Resembles female in wing and leg markings. Head.

Second palpal segment with dorsomedian line of pale scales; 3rd palpal segment

without basal pale band. Genitalia. Basimere with lateral and dorsal pale

scales; inner parabasal spine with sharply hooked tip, outer spine straight,

slender and longer than inner spine; mesal subapical spine long, slender and

curved; tergum IX with long slender processes, tips slightly expanded; aedea-

eus with 3 - 6 pairs of leaflets, larger leaflets with several teeth toward the

apex and with small basal denticles; ventral lobe of claspette with 2 - 3 setae

as long as or longer than spines on dorsal lobe, and several smaller setae

clustered at their bases; dorsal lobe of claspette with club formed from at

least 3 spines on the inner half, while outer half contains a single spine which

is bifid apically and adjacent to fused club.

PUPA (Fig. 11) Cephalothorax. Light to moderate pigmentation, wing case

Harrison and Scanlon: Subgenus Anopheles in Thailand 47

with darkest pigmentation in form of distinct rows of round dark spots, infre-

quently with lighter pigmentation on veins or crossbars. Trumpet. With thin

uniform rim, without secondary cleft or tragus. Abdomen. Terga III - VI

without or with central dark spot; seta 5 with relatively few branches, 9 - 24

on segment V; 1-VII with 1 - 5 branches; 9-VIII, with well developed branches.

Paddle. Refractile border about 0.65 paddle length; setae 1, 2-P single, or

1-P may be frayed.

LARVA (Fig. 12) Head. Seta 2-C simple, 3-C with 60 - 80 branches form-

ing distinct tuft with branches difficult to count; 4-C with 2 - 8 branches; fron-

tal setae, 5, 6-C with 16 - 24 and 16 - 21 branches respectively; 8-C with 7 -

14 branches; 9-C with 6 - 9 branches. Antenna. With spicules, largely on

inner aspect; seta 1-A long, with 6 - 14 branches. Thorax. Seta 1-P without

sclerotized base, simple or branched near tip; 2-P with prominent sclerotized

base and 9 - 13 branches; 11-P with 3 - 6 short, stout branches; prothoracic

pleural tubercle with long stout spine; 14-P with 7 - 12 branches; 4-M with 4 -

8 branches from a central stem, not sinuous; 3-T palmate, with small number

of unpigmented branches, without filaments. Abdomen. Seta 1 partially devel-

oped and pale on segments I - II, well developed (palmate) and pigmented on

[lI - VII, leaflets without well developed shoulders, but most with sharply

pointed tips; 6-I with 21 - 28 branches, 6-III with 17 - 29 branches; 5-II with

9 - 20 branches; 9-III with 8 - 14 branches; 13-IV with 3 - 6 long slender

branches from base; 5-VI with 6 - 11 branches; pecten with 7 - 9 long teeth

and approximately double that number of small teeth; seta 1-X at least as long

as saddle.

EGG. None were examined during this study. Reid (1968) reports that the

eggs of sinensis have an unusually broad deck, making up about 0.33 the width

of the egg; float ribs about 30 - 35. The egg illustrated by Ho et al. (1962) as

sinensis in East China, has the wide deck, but shows only 29 ribs occupying

somewhat less of the lateral aspect than indicated by Reid.

TYPE-DATA. A female lectotype was designated and described for sinensis

by Harrison (1973a), and deposited in the Universitetets Zoologiske Museum,

Copenhagen, Denmark. This specimen is in excellent condition and bears the

following data on 2 labels: ''Coll. Westerm." and ''Anopheles sinensis Wied.

China, Trentepohl.'' A female paralectotype, in good condition, is also deposit-

ed in the above museum. Two additional paralectotypes, a male and a female

in very poor condition, are deposited in the Naturhistorisches Museum, Vienna,

Austria. The 2 latter specimens were listed as the types for sinensis by Stone

et al. (1959) and Reid (1968). The type-specimens of plumiger Donitz, are

located in the Zoologisches Museum, Humboldt Universitat, Berlin, while the

location of the type of jesoensis Tsuzuki, is unknown (Stone et al. 1959).

DISTRIBUTION (Fig. 11). This species can be collected nearly anywhere

in Thailand except possibly in the most precipitous and heavily forested areas.

It is particularly abundant in rice growing areas and rice is a major crop

throughout Thailand. Consequently, the distribution map (Fig. 11) is actually

one of potential distribution, rather than one based entirely on examined speci-

mens. Scanlon et al. (1968) listed collections made from 24 provinces. Speci-

mens (300°, 749, 63 larvae and 106 larval and pupal skins) located in the USNM,

were examined from the following provinces of THAILAND: Chanthaburi,

Chiang Mai, Chiang Rai, Chon Buri, Lampang, Mae Hong Son, Nonthaburi,

Phra Nakhon, Prachuap Khiri Khan and Tak. Aside from the provinces listed

above and in Scanlon et al. (1968), specimens of sinensis were examined in

Thailand in 1969 from 3 other provinces: Buriram, Lop Buri and Sara Buri.

Several hundred additional specimens located in the USNM and identified as

48 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

sinensis, were also examined and confirmed from: BURMA, CAMBODIA,

CHINA (Canton, Yunnan), HONG KONG, INDIA (Assam), INDONESIA (Sumatra),

MALAYSIA (Peninsular Malaysia), NORTH VIETNAM, JAPAN (Ryukyus),

SINGAPORE, SOUTH VIETNAM and TAIWAN. Harrison (1973a) listed

sinensis from Java in error.

TAXONOMIC DISCUSSION. This was one of the 1st Anopheles species

reported from Thailand (Theobald 1910), and while there was early confusion

concerning the name to be applied, most of the early records of hyrcanus

group species from Thailand probably apply to this species. It was not until

Reid's (1953) revision of the Southeast Asian hyrcanus group that it became

possible to identify the group members and define their distribution with any

degree of certainty.

Reid (1953) commented on the frequent presence of pale scales on the

basal portion of the costa on specimens from China. Specimens in the USNM

from Canton, Yunnan, Hong Kong, Taiwan and the Ryukyus show this feature.

In addition, many exhibit a paler wing pattern. These specimens also exhibit

more extensive pale scaling on the antennal flagellomeres. Specimens from

North Vietnam, South Vietnam and Cambodia were darker, with fewer pale

scales on the antennal flagellomeres, like the specimens from Thailand,

Malaysia and Singapore. Too few specimens were available from Assam,

Burma and Indonesia to indicate trends.

Immature stages from the Ryukyus, Taiwan and Hong Kong were checked

and found to be similar to Thailand specimens. Although the pupal wing

cases normally had less pigmentation, the distinct dark spots were still usually

discernable.

After examining the specimens in the USNM and material in Thailand it is

evident that confusion still exists regarding the identification of the adults and

immatures of sinensis, This confusion is most often exemplified in the inclu-

sion of several other species under the name sinensis, rather than the identifi-

cation of sinensis under some other name. Species most often misidentified

under sinensis are: crawfordi and nigerrimus as adults; nigerrimus and

nitidus as pupae; and cvawfordi and nitidus as larvae.

Confusion between sinensis and nigerrimus stems mainly from the highly

variable hindtarsal banding found in adult nigerrimus (see p. 68), and a small

degree of pigmentation overlap on the pupal wing case. However, the differ-

ences in pupal seta 5-V (sinensis 9 - 24 branches; nigerrimus 40 - 60 branches)

are sufficiently different to separate these species. Confusion between the

pupal stages of sinensis and nitidus is probabiy due to the occasional occur-

rence of very pale sinensis; however, seta 5-V is still the differentiating

character. The larvae of these 2 species may rarely overlap, but the branches

on head setae 5, 6-C, 6-III and the number of long pecten teeth should usually

separate them. Anopheles crawfordi and sinensis adults are very similar and fre-

quently confused. This problem is aggravated by variation in sinensis, but the

characters presented in the key are still adequate for separating over 95% of the

specimens encountered. Occasional larval specimens of cvawfordi and sinen-

sis are quite difficult to separate. When the key characters break down due

to overlap, then the worker should check the number of branches on setae 5-C

(sinensis, 16, normally 17 - 24; crvawfordi, 13 - 16), 6-C (sinensis, 16, nor-

mally 17 - 21; cvawfordi, 14 - 17), and seta 9-III (sinensis, 8 - 14; crawfordi,

5 - 11). Rare confusion may occur between sinensis and lesteri paraliae lar-

vae; however, pavaliae is always very darkly pigmented and has fewer branches

on seta 5-II (6 - 10) than does sinensis (9 - 20). Asa last resort there is one

character that will differentiate sinensis larvae from nearly all others. This

Harrison and Scanlon: Subgenus Anopheles in Thailand 49

character is most difficult to see and even more difficult to determine because

it involves counting setal branches. Seta 15-C has 10 or more branches on

sinensis larvae, while only rare specimens of nigerrimus ever reach this num-

ber and the other species have less than 10 branches.

One female from Chiang Mai was normal in all respects (including associ-

ated skins), except the preapical pale costal spot was missing on both wings.

The absence of this spot left only one pale spot (subcostal) on the leading mar-

gin of the wing. Other sinensis collected with this specimen were all within

the normal range of variation.

An. sinensis isahighly variable species. Variation is common inoverall size,

wing markings, larval chaetotaxy and the utilization of habitats. Apparently,

sinensis exists in Thailand under near optimum ecological conditions, and thus

exhibits the wide range of variations one would expect from a large gene pool.

BIOLOGY. This is one of the most abundant and widely distributed mos-

quitoes in Thailand. It is most characteristic of open agriculture lands (chiefly

rice fields), but is not limited to this habitat. Immature stages have been col-

lected from such varied habitats as: ground pools, pools beside a river,

marshes, rice fields, stream margins, ditches, seepages, shallow ponds and

sumps. All of these are normally fresh, shallow water habitats, usually with

emergent vegetation (mostly grasses) and exposed to direct sunlight. No col-

lections of this species have been made from brackish water, but large num-

bers were encountered along stream margins in Chiang Mai Province below hot

springs which gave off a moderate sulphurous odor. In the Bangkok area, si-

nensis reaches peak seasonal abundance in October-November, when water

reaches its highest level in the rice fields.

In mountainous regions sinensis should be considered a mosquito of the

valleys, except in situations where extensive rice fields are at high elevations.

Scanlon and Esah (1965) reported surveys in the Chiang Mai Valley area where

sinensis was collected biting man at approximately 305 m, but not in the forested

areas at higher elevations. During a 14 month period in 1965-1966, human bit-

ing collections made in small villages in an elevated, semiforested area in

Kanchanaburi Province, yielded 9,303 mosquitoes, but no sinensis (Harinasuta

et al. 1970). Wilkinson et al. (1970) reported on anthropophilic mosquitoes col-

lected in a jungle camp in the Petchabun Mountains, and only recorded a single

sinensis in 15 nights of collections. Its apparent low incidence at higher forested

elevations is also supported by larval collection records. However, since s7-

nensis is primarily a zoophilic species, low numbers or its absence in human

biting collections can not be considered a true indication of population levels.

In northern Thailand and the central valley, observations (SEATO Laboratory

unpublished report) made for nearly 10 years indicate that sinensis is almost

entirely zoophilic.

In comparative biting tests involving man and cows, almost none of the

sinensis were attracted to man. Similar results have been reported by Reid

(1968) from Malaysia. Moderate numbers of sinensis were collected at night

in Lop Buri and Sara Buri Provinces by placing dry ice inside partially opened

army bed nets and at one hour intervals closing the nets and collecting those

mosquitoes attracted by the carbon dioxide. Apparently, sinensis is exophilic,

for extensive resting collections in houses in Bangkok yielded almost no speci-

mens (SEATO Laboratory unpublished report).

MEDICAL SIGNIFICANCE. Apparently sinensis is of little or no significance

to human health in Thailand. The species is zoophilic, exophilic and further-

more, the areas of maximum sinensis populations in Thailand coincide primari-

ly with areas of little or no malaria transmission. Iyengar (1953) reported

50 Contrib. Amer. Ent. Insts, vol. 12, no. 1, 1975

sinensis in south Thailand with microfilariae of Brugia malayi; however, it is

uncertain which members of the group were involved in Iyengar's "hyrcanus

Sinensis"' and "hyrcanus nigerrimus", as this study was prepared prior to

Reid's (1953) revision of the group. Harinasuta et al. (1964) examined a small

number of unspecified specimens of the hyrcanus group in southern Thailand,

but found none infected. Reid et al. (1962) reported that natural infections

found in members of this group (not sinensis) in Malaysia were animal filariae

of the genus Se/aria, During the same study sinensis was considered refractory

to experimental infection with periodic B. malayi because only 3 of 99 infected

adults developed mature larvae. Reid et al. (1962) also claimed that Hodgkin

(1937, 1939) obtained similar results with sinensis infected with semi-periodic

B. malayi. According to Ho et al. (1962) sinensis is a filaria vector of some

importance in the Yangtze Valley area of China where B. malayi is prevalent,

and that it is also responsible for low levels of malaria endemicity in the open

plains, while lestevi lesteri is the main malaria vector in eastern China. The

situation may be even more complicated in China for at least 3 other sibling

species besides lestevi lesteri and sinensis have been called ''sinensis" in the

past (Ho et al. 1962). Regardless of the Chinese situation, there is no sub-

stantiated data available that incriminates sinensis as a vector of human patho-

gens in Thailand.

ANOPHELES (ANOPHELES) ARGYROPUS (SWELLENGREBEL)

(Figures 9, 13, 14, 15)

Myzorhynchus argyropus Swellengrebel 1914: 334 (2*).

Anopheles hyrcanus var. argyropus of Christophers 1924: 30.

Anopheles (Anopheles) hyrcanus aberration argyropus of Edwards 1932: 41.

Anopheles hyrcanus typicus var. pseudopicta of Swellengrebel and Rodenwaldt

1932: 69 (not Grassi 1899).

Anopheles (Anopheles) argyropus Swellengrebel, Reid 1953: 36 (“*, @*, P*, L*,

i):

Anopheles (Anopheles) argyropus (Swellengrebel), Reid 1968: 113 (“*, 9*, P*,

Lt, BS).

Adults may be recognized by a combination of extensive pale bands on the

hindtarsi, long basal dark mark on vein Cug, remigium mostly dark scaled and

a tuft of dark scales on the humeral crossvein. The male lacks a pale band at

the base of palpal segment 3. The pupa is easily recognized by the parallel

folds vertical to the trumpet rim, and seta 9-VIII (lateral spine) without branches

or with few very fine branches. The larva can be recognized by the number of

branches on setae 8-C, 14-P, 5-IV, the shape of tergal plate VIII and other key

characters. Like sinensis except:

FEMALE (Figs. 9, 13) Head. Palpus dark, with 4 well defined pale bands,

(sometimes only 3, when apical 2 bands confluent), and usually a few pale

scales on inner surface of segment 2. Thorax. Scutal integument very dark,

with darker prominent eye-spots, median and lateral dark lines and numerous

golden setae; anterior promontory with narrow pale mesal scales and dark

lateral scales, rest of scutum without scales; pleural setae; 3 - 7 propleural,

2-4 spiracular, 5 - 6 prealar, 2 - 4 upper and 4 - 6 lower sternopleural, 3 -

6 upper and 0 lower mesepimeral. Wing. Costa dark, except small subcostal

and preapical pale spots; remigium with few pale scales; humeral crossvein

with patch of dark scales; vein R with few pale scales between base and sector

pale spot, often between sector and subcostal pale spots, no pale scales on

Harrison and Scanlon: Subgenus Anopheles in Thailand ol

preapical dark mark; tip of R; dark scaled; subcostal pale spot often small,

ill-defined, sometimes missing; Raq - Ro, dark; Ry A: usually with few dark

scales in median pale area; vein M dark except pale area just before fork;

extreme base of Cu with small dark spot usually followed by short pale area

and long "'basal'' dark mark, which approaches or reaches within its own

length of upper dark mark on vein 1A; 1A usually with short upper and apical

dark marks, infrequently these 2 marks fused and apical half of vein dark

scaled; Cu» with apical dark mark longer than that on 1A; apical pale fringe

spot variable, usually extending from R, to Ry,5, but may be shorter, between

Ry and Rg; posterior margin of wing without pale fringe spot at Cup. Legs.

Midcoxa with 3 - 4 upper setae. Foreleg: femur dark scaled; tibia pale ven-

trally, may have very small apical pale spot. Midleg: femur dark scaled;

tibia with venter pale scaled and apical pale spot; tarsomeres 1 - 3 with very

narrow apical pale bands. Hindleg: tibia with distinct apical pale spot; tarso-

mere 1 with apical band; tarsomere 2 with longer apical band extending onto

base of 3; tarsomere 3 with wide apical pale band extending well onto the base

of 4; apical pale band of 4 extending well onto base of tarsomere 5; in some

specimens tarsus may be entirely pale from middle of tarsomere 3 to the tip,

usually however, at least a trace of dark median band on tarsomere 4.

MALE (Fig. 13). General markings as for the female. Head. Palpal

segments 2 - 3 often with scattered, pale scales, base of segment 3 without

pale band, segment 4 - 5 with pale bands, that on 5 occupying most of segment.

Genitalia. Basimere with numerous pale scales ventrally, some darker scales

laterally; tergum IX with long posterior processes, tips expanded; aedeagus

with 2 pairs of leaflets (rarely 3), with small serrations; ventral lobe of clas-

pette with 2 setae, mesal seta longer than club on dorsal lobe, and smaller

lateral seta. Dorsal lobe of claspette with 3 spines which fuse apically to form

club on inner half, and 2 spines on outer half adjacent to fused club.

PUPA (Fig. 14) Cephalothorax. Wing case with pattern of lines and spots,

forming rectangles in some areas, spots not darker than other pigmentation;

tip of antennal case dark. Trumpet. Rim thin and uniform, with parallel

vertical wrinkles on sides. Abdomen. Terga III - VII with central dark spot;

setae 1 and 5 densely branched on segments III - VI; seta 9-VIII spine-like or

with few short, fine branches. Paddle. Refractile border long, about 0.8 -

0.9 paddle length; setae 1, 2-P usually single.

LARVA (Fig. 15) Head. Seta 3-C with 70 plus, crowded branches; 5, 6-C

with 12 - 16 and 12 - 19 branches respectively; 8, 9-C with 13 - 22 and 9 - 16

branches respectively. Thorax. Seta 1-P usually with 2 - 3 fine apical branches;

2-P with 8 - 13 branches; 11-P with 5 - 6 stout branches from base; 14-P with

6 - 11 branches; 4-M with 6 - 12 branches from base, usually not sinuous; 3-T

with flattened branches, not fully developed palmate. Abdomen. Seta 1 pal-

mate, poorly developed on segments I - II, well developed and uniformly pig-

mented on segments III - VII, filaments uncolored; 2-II with 6 - 9 branches;

5-II with 9 - 15 branches; 6-I, III with 16 - 19 and 15 - 20 branches respec-

tively; 9-III with 6 - 13 branches; 5-IV with 4 - 10 branches; 5-V with 4 - 6

branches; 5-VI with 5 - 6 branches; 9-VI with 4 - 5 branches; 13-IV with 7 - 16

short branches (cf. sinensis); tergal plate VIII sometimes roughly hexagonal

instead of ovoid or roughly rectangular; pecten with 6 - 8 long teeth, usually 7;

seta 1-X strong, longer than saddle; posterior tergal plates much darker than

anterior tergal plates.

TYPE-DATA. The type-locality is Deli, Sumatra, INDONESIA. The type-

specimen, a female, appears to have been lost and Reid (1953) designated

plesiotypes (with larval and pupal skins) from MALAYSIA, which are in the BMNH.

52 Contrib. Amer, Ent. Inst., vol. 12, no. 1, 1975

DISTRIBUTION (Fig. 14). This species was infrequently collected, usually

with other members of the hyrcanus group, from widely scattered provinces in

Thailand. Specimens (40°, 209, 2 larvae and 8 larval and pupal skins) in the

USNM were seen from the following provinces of THAILAND: Chiang Mai,

Chanthaburi, Chon Buri, Narathiwat, Pathum Thani, Phra Nakhon and Udorn.

Besides the provinces listed by Scanlon et al. (1968) specimens were examined

in 1969 in Thailand from Buriram and Sara Buri. The authors also examined

4°, 272, 2 larvae and 12 larval and pupal skins of argyvopus in the USNM from:

CAMBODIA, INDIA (Assam), INDONESIA (Java and Sumatra) MALAYSIA (Pen-

insular Malaysia) and SOUTH VIETNAM. An additional 192 from CAMBODIA

were found in the ORSTOM collections (Centre de ORSTOM, Bondy, France).

TAXONOMIC DISCUSSION. The accumulated differences in the adult and

immature stages make this one of the easiest species of the hyrcanus group

to recognize in Thailand. The overall impression is that it is more distantly

related to sinensis than are the other members Of the Southeast Asian hyrcanus

group. Reid (1968) has suggested that it is in a connective position between

the Southeast Asian hyrcanus group and the coustani species group of the Ethi-

opian region. The species in the latter group have the pupal trumpet with a

series of vertical wrinkles, and most members have the hindtarsi pale distal

to the apex of tarsomere 3 (Gillies and de Meillon 1968). None of the members

of the coustani group occur east of Israel (Margalit and Tahori 1970) and south-

western Arabia, and argyropus has its westernmost distribution in Assam.

We have recently examined specimens of hyrcanus (synonym = pseudopictus

Grassi) females from Afghanistan. These resembled argyropus in the extent

of pale markings on the hind tarsus, but had small numbers of dark scales in

the pale areas and differed somewhat in wing pattern. Three specimens of

argyropus from Java, obtained through the courtesy of the Instituut voor

Tropische Hygiene, Amsterdam, were also examined. These bore the label

"bseudopicta Grassi"' and were presumably mentioned by Swellengrebel and

Rodenwaldt (1932). These last 3 specimens had reduced subcostal pale spots

(also seen on some Thai specimens) but were otherwise typical argyropus.

Reid (1953, 1968) said argyropus shows closest relationship to the south-

east Asia hyrcanus group through peditaeniatus. This was justified by simil-

arities in: the long refractile border on the pupal paddle; the spinelike seta

9-VIII on the pupa; broad hindtarsal bands on the adults; palpal scaling; and

yellow-orange pale scales on the wings. He also pointed out at least 2 dis-

similarities with larval nigerrimus: antennal shaft shape and the number of

branches on seta 5-IV.

The authors feel that avgyropus is more closely related to nigerrimus than

to peditaeniatus. Studies of all 3 species over wide areas of their distributions

(Reid 1963, 1968) and herein show that hindtarsal banding is highly variable.

All 3 species exhibit essentially the same variation in palpal scaling and all

can have yellow-orange scales on the wings. Anopheles argyropus is similar

to nigerrimus in many ways: Adult - humeral crossvein with a patch of dark

scales; remigium mostly dark scaled; midtarsi with narrow apical pale bands;

2 - 3 pairs of leaflets on male aedeagus; Pupa - trumpet without thickened,

sawtooth areas (although argyropus does have vertical wrinkles); many

branches on setae 1 and 5 on segments III - VI; Larva - many branches on

setae 8, 9-C, seta 3-C with 70 or more branches, and lastly a common larval

habitat, i.e., deepmarshesandcanals. An. peditaeniatusdiffers markedly from

all the above: no scales on humeral crossvein, remigium mostly pale scaled;

midtarsomere 3 with wide pale band, 4 - 7 pairs of leaflets on male aedeagus;

trumpet with thickened sawtooth areas; setae 1 and 5 on pupal segments III - VI

Harrison and Scanlon: Subgenus Anopheles in Thailand D3

with relatively few branches; larval seta 3-C with 40 - 70 branches, larval

setae 8, 9-C with few branches, and a shallow larval habitat such as grassy

ponds, swamps and rice fields. An. argyropus differs from nigerrimus by

lacking a pale band at the base of 3rd palpal segment of male; absence of any

pale scales on the base of the costa; having parallel vertical wrinkles on the

pupal trumpet; and having a somewhat intermediate number of long pecten teeth

(nigerrimus, 4 - 7; argyropus, 6 - 8; peditaeniatus, 7 - 9).

A few specimens of peditaeniatus examined from Thailand, verified by

associated larval and pupal skins, had the hindtarsal bands nearly identical to

those seen on argyropus with narrower than average bands. Similar specimens

of peditaeniatus were also examined from Cambodia, South Vietnam and Malay-

sia. These 2 species should not be confused. The midtarsal banding is very

distinct and there are also differences in the scales on the humeral crossvein,

remigial scaling, pale scaling on the base of vein R and the preapical dark spot

on vein R;. The use of hind tarsal banding to separate these 2 species should

be avoided.

Some confusion might arise over the identity of avgyropus larvae because

the branching of seta 4-M arises from the base and occasionally the branches

are slightly sinuous like those of peditaeniatus. This problem can be resolved

very quickly by counting the branches on setae 8, 9-C, those of argyropus have

many more branches (see descriptions).

Two specimens of avgyropus from Chiang Mai were found to have the hind-

tarsi entirely white distal to the middle of tarsomere 3, otherwise they were

normal. Several other specimens from Thailand were examined which had

only faint dark bands on hindtarsomere 4, and only the tip of hindtarsomere

o dark scaled. Similar specimens with excessively pale hindtarsi were exam-

ined from Peninsular Malaysia, Indonesia (Java) and South Vietnam.

Additional variations noted on specimens of avgyropus were: apical female

palpal segment entirely pale; reduced subcostal spots on the costa; base of vein

Cu dark scaled halfway to the fork; and apical 0.5 of vein 1A dark scaled. The

latter variation was noted on several specimens from Thailand.

BIOLOGY. Adults of avgyropus have been collected in small to moderate

numbers in light traps in Bangkok during most months of the year. The largest

numbers appeared in November - January, between the end of the monsoon rains

and the cool season when surface waters were stable and at their greatest ex-

tent. Small numbers have been taken during human biting collections in Bang-

kok, Buriram, Chiang Mai, Chon Buri, Narathiwat, Pathum Thani, Sara Buri

and Udon Thani. This species has also been collected in Chanthaburi Province

in a Shannon trap with a light and men inside and in Sara Buri Province ina

net trap baited with dry ice (carbon dioxide). Scanlon and Esah (1965) report

that no argyropus were collected in heavily forested areas, and that far more

were attracted to domestic animals than to man during their collections. In

Thailand, immature stages have been collected from rice fields and deep see-

page washes. Reid (1968) reported immatures collected in Malaysia from

large deep swamps. Deep swamp and canal habitats need further examination

in Thailand.

MEDICAL SIGNIFICANCE. As might be expected from the largely zoophilic

habits, there is no evidence that argyropus is a vector of human pathogens

anywhere in its range. Warren et al. (1963) evaluated Malayan argyropus

as a vector of the simian malaria parasite, Plasmodium cynomolgi bastianellii

Garnham, and found it almost refractory to infection. Harinasuta et al. (1970)

dissected 12 argyropus in Kanchanaburi Province, Thailand, and found them

negative for Brugia and Dirofilaria filariae.

o4 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

ANOPHELES (ANOPHELES) CRAWFORDI REID

(Figures 9, 16, 17, 18)

Anopheles (Anopheles) crawfordi Reid 1953: 41 (*, 9*, L, P, E*); Reid 1968:

£02 (6/2 9*) Pt AL; B*).

Adults can usually be recognized by the narrow hindtarsal bands, wide

apical pale wing fringe spot, pale scales on the coxae and male basimeres,

a wing pattern consisting of rather short sharply defined dark spots on a

generally pale field, and the tip of vein Ry pale. The male lacks a pale band

at the base of the 3rd palpal segment. Identification of the larva and pupa

is much more difficult and the characters outlined in the keys and description

must be consulted. This species is similar to sinensis except for:

FEMALE (Figs. 9, 16) Head. Palpus with sharply defined pale bands,

some pale scales on segment 2, with few or none on segment 3 other than

band. Thorax. Scutal integument basically grayish, without well developed

dark lines, with very prominent eye spots; pleural setae: 2 - 5 propleural,

3 - 4 spiracular, 6 - 9 prealar, 3 - 5 upper and 3 - 5 lower sternopleural and

0 - 9 upper mesepimeral; occasionally 1 - 2 scales on latter area. Wing.

Costa dark with usual subcostal and preapical pale spots, humeral area may

have small humeral spot or 1 - 3 pale scales on one wing; humeral crossvein

without scales; R with few pale scales on basal 0.33, with well developed sec-

tor pale spot; Ry without pale scales on preapical dark area, with pale scales

on tip; R,-R9i3 with distinct dark basal mark, then pale to dark mark at fork;

Ro and Re largely pale; R4,5 with short basal and apical dark marks pale in

between; vein M mostly pale, but with scattered dark scales near base; extreme

base of Cu pale, or with several dark scales, followed by pale area, then short

well defined dark mark which is separated by its own length or more from

upper dark mark on vein 1A; Cu, and Cy with very short apical dark marks;

1A with short upper dark mark, Lpical fat mark usually equal to or less than

length of apical dark mark on Cus: pale apical fringe spot extending from Ro

to just beyond R45; posterior margin without pale fringe spots. Legs. All

coxae with pale scales; midcoxae with 3 - 4 upper setae. Foreleg: tibia without

pale knee spot; tarsus with moderately wide apical pale bands on tarsomeres

1-3. Midleg: tarsus with apical pale bands on tarsomeres 1 - 4, those on

tarsomeres 1 - 3 markedly narrower than those on foretarsus. Hindleg: tar-

sus like that of midtarsus, no basal banding.

MALE (Fig. 16) Head. Inner dorsal surface of palpal segments 2 » 3 may

have prominent line of pale scales; palpal segment 3 without basal pale band

(see taxonomic discussion); palpal segments 4 - 5 largely pale scaled. Geni-

talia. Basimeres with few pale scales; inner parabasal spine with sharply

hooked tip; tergum IX with long posterior processes, tips slightly expanded;

ventral lobe of claspette with 2 setae, mesal seta distinctly longer than lateral

seta and spines on dorsal lobe; dorsal lobe of claspette with fused club on inner

half formed. from at least 3 separate stems; outer half of dorsal lobe with 2

flattened, leaflet-like spines adjacent to fused club; aedeagus with 4 - 7 pairs

of leaflets, largest with well developed teeth at apex and along one margin,

with basal denticles on the first 2 leaflets.

PUPA (Fig. 17) Cephalothorax. Integumental pigmentation variable, from

light to heavy, usually with well marked pattern of dark lines forming squares

or rectangular areas on wing case; antennal case usually with apex and most

apical joint darkly pigmented. Trumpet. Pigmentation tan except darker

Harrison and Scanlon: Subgenus Anopheles in Thailand 55

thickened areas on rim; rim with thickened areas of erect saw-like teeth; meatal

cleft narrow. Abdomen. Terga III - VII with central dark spot; seta 1 with

fewer branches than seta 5 on IV - VII; 1-VI with 2 - 6 branches; 5-V with 30 -

45 branches. Paddle. Refractile border approximately 0.55 - 0.65 of paddle

length; 1-P with few apical branches.

LARVA (Fig. 18) Head. Seta 3-C with approximately 50 crowded branches;

0, 6-C with 13 - 16 and 14 - 17 branches respectively; 8, 9-C with 6 - 11 and

0 - 10 branches respectively. Thovax. Seta 1-P simple or with 2 - 3 short,

thin branches near tip; 11-P with 4 - 6 stout branches from base; 14-P with

6 - 9 branches; 4-M with 4 - 8 branches from erect central stem; 3-T with

slightly flattened and unpigmented branches. Abdomen. Seta 1 with slightly

flattened, very lightly pigmented branches on segments I - II, well developed

(palmate) and deeply pigmented on segments III - VII, pigment not extending

onto filaments; 6-I and III with 16 - 21 and 15 - 21 branches respectively (rarely

20 - 21); 5-II with 10 - 18 branches; tergal plate VUI oval; pecten with 7 - 8

long teeth (infrequently 6 on one side); seta 1-X weak, usually shorter than

saddle.

TYPE-DATA. The ¢ holotype and “ allotype, both with associated immature

skins on slides, plus 1% and 3 paratypes are deposited in the BMNH. Reid

(1953: 43) clearly says the type-locality is Kuala Lumpur; however, none of

the above types are so labeled, although each has been identified by Reid and

has an appropriately colored BMNH type-label on it. The 2 is in excellent con-

dition and has the following label data: (1st label) - ''Selangor, 5th mile Ke-

pong, 15-11-1950, J. A. Reid"; (2nd label) - ''No 8445/11, larvae in swamp

overgrown with vegetation"; (3rd label) - "Anopheles (A.) crawfordi Reid, Det.

J. A. Reid''; (4th label) - round circular red label without writing; (5th label)

- circular label with "holotype"; and (6th label) - "liason pelts in slide collec-

tion.'' The “ allotype is also in excellent condition with genitalia still intact,

and has the following label data: (1st label) - 'Selangor, 5 1/2 Kl. - Kepong,

28-3-1951, J. A. Reid"; (2nd label) - "No 454/62, larvae in swamp overgrown

with vegetation"; (3rd label) - "Anopheles (A.) crawfordi Reid, Det. J. A.

Reid"; (4th label) - round circular red label without writing; (5th label) - circu-

lar label with "'Allotype “''; and (6th label) - 'liason pelts in slide collection."

The © and 32 paratypes have the same data as the allotype except: all are

. numbered ''No 494"; each has circular "paratype" label; and they lack the

"liason pelt'' label. The “ paratype is in excellent condition with genitalia

intact, while the 32 paratypes have lost their heads.

DISTRIBUTION (Fig. 17). This species does not appear to have a continu-

ous distribution throughout Thailand. It is most often encountered in the south-

ern or southeastern parts of the country. Certain climatological factors may

be partially or wholly responsible for the distribution of cvawfordi (see biology

section). The authors currently feel crawfordi will probably not be found in the

dryer parts of the country, i.e., areas of western, northern and northeastern

Thailand. Specimens (14%, 269, 17 larvae and 41 larval and pupal skins) in

the USNM were seen from the following provinces of THAILAND: Chanthaburi,

Chon Buri, Narathiwat, Nakhon Ratchasima, Songkhla, Surat Thani, Trat and

Yala. Besides these provinces and those listed by Scanlon et al. (1968), speci-

mens were examined during 1969 in Thailand from Chumphon and Ranong.

Also examined were 9°, 242 and 10 larval and pupal skins of cvawfordi in the

USNM from CAMBODIA, INDIA (Assam), MALAYSIA (Peninsular Malaysia)

and SOUTH VIETNAM. The type-specimens (2, 42 and 4 larval and pupal

skins) from MALAYSIA were examined in the BMNH, and 7 from CAMBODIA

were found in the ORSTOM collections. Additional literature records apply to

06 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

INDONESIA (Sumatra).

TAXONOMIC DISCUSSION. The adults have the general appearance of

small specimens of sinensis, and can easily be misidentified as that species

without careful examination. The sharp appearance of the wing spots on craw-

fordi is usually a good character, but may be difficult to see in older, rubbed

specimens. The wing of cvawfordi is quite similar to that of nitidus, but the

extensive pale banding on the hindtarsus of nitidus is distinctive. There are

no apparent differences in tarsal banding between crawfordi and sinensis,

thus the taxonomist is left with wing characters and pleural chaetotaxy for

characters. The pale tip on vein Ry and the absence of pale scales on the pre-

apical dark area on vein R, of crawfordi are the most useful characters in

separating it from sinensis. Even these characters are not absolute, for rare

specimens of sinensis were noted from Thailand which exhibited either the tip

of R, pale, or R,; with the preapical dark mark entirely dark scaled. Reid

(1953) noted 6% (3/50) of the cvawfordi he studied had dark scales on the tip of

R;. This was rarely encountered during the present study, and was always con-

fined to only one wing. A pale fringe spot at Cu, 6 or more propleural setae

and a long apical dark mark on 1A are all generally indicative of sinensis, but

should be used with caution.

Specimens of lesteri lesteri from Hong Kong were also noted as very

similar to crawfordi, but had the tip of R, dark scaled, preapical dark mark

on R, with some pale scales, no fringe spot at Cuo, a long apical dark mark

on 1A and no pale scales on the midcoxae and male basimeres.

Of the 19 males of cvawfordi examined from Thailand, Cambodia, Assam

and Malaysia, none exhibited signs of a pale band on the base of the 3rd palpal

segment. This would be expected as crawfordi is most closely related to les-

teri and peditaeniatus, and males of this subgroup lack a pale band at the base

of palpal segment 3. However, Reid (1953, 1968) noted that males of craw-

fordi sometimes exhibit a small pale band at the base of this segment. Since

we have not seen this trait on crawfordi we have no explanation, unless this is

a localized or infrequent variation of crawfordi.

As noted earlier (Reid 1953), occasional specimens exhibit either 2 - 3

pale scales or a distinct pale spot on the humeral area of the costa. This varia-

tion is usually confined to one wing and can be of value in identifying crawfordi,

particularly the males. Males without a pale basal band on palpal segment 3,

but with a pale humeral spot or pale scales in that area will most probably be

crawfordi. No males of lesteri paraliae, peditaeniatus or sinensis from Thai-

land, have been found with pale scales on the base of the costa.

The pupal stage of cvawfordi is very similar to that of lesteri pavaliae.

The thickened, saw-tooth like areas on the trumpet rim are exhibited only in

these 2 species and peditaeniatus; however, the latter species is easily separ-

able by the spine-like seta 9-VIII and a long refractile border on the paddle.

Separation of crawfordi pupae from those of lesteri paraliae is very difficult.

The pupa of lesteri paraliae is very darkly pigmented, but infrequently that of

crawfordi is very dark. Seta 1-VI on Thailand specimens of crawfordi have

4 - 6 branches rather commonly and thus overlap with the 5 - 10 branches

found on 7, paraliae. The number of branches on seta 5-V (see key, p. 43)

is the most reliable character for differentiation of these 2 species, but it is

also a tedious character to check. The pupa of cvawfordi should never be con- ©

fused with that of s7zmensis or the remaining species for they all have a thin and

uniform trumpet rim or one with vertical wrinkles (argyropus). Difficulty is

also encountered in the identification of the larvae of crawfordi, for specimens

have been found which have only 10 - 11 branches on abdominal seta 9-II and

Harrison and Scanlon: Subgenus Anopheles in Thailand D7

this overlaps with 6 - 10 branches found on l. paraliae.

Persons working in areas where crawfordi and/or lesteri paraliae night be

encountered, should conduct sample rearings to determine the species involved.

The adults of these 2 species are readily separable, while the reliability of

separating the immature stages is no more than 90%.

The examination of specimens from Thailand in comparison with specimens

from Assam, Cambodia, Malaysia and South Vietnam, revealed little variation.

One male and 2 females of 13 Assam adults exhibited a reduction in the apical

fringe spot on the wing to a size comparable with that on lesteri paraliae.

Otherwise, these specimens were typical cvawfordi, and are thus considered

variations of crawfordi. Reid (1953: 50) commented on a tendency in this spe-

cies for a reduction in the apical fringe spot.

BIOLOGY. The numbers of crawfordi adults encountered during studies in

Thailand were usually small and dependent on the worker's location. However,

several local areas were encountered in southern and southeastern Thailand

where cvawfordi was abundant. These areas usually had tropical wet forest

nearby, and always received abundant precipitation. Larval requirements

indicate this is essentially a forest-swamp species. Larvae are found in clear

water and nearly always in moderate to heavy shade. Confirmed larval habi-

tats in Thailand include: seepage pools or springs, shallow-slow moving

streams, stream pools, ground pools, large deep ground pools and sapphire

mining pits, swamp pools and nipa palm swamps. Nearly all of these habitats

contained floating leaves and/or emergent vegetation such as grass. No col-

lections of cvawfordi larvae have been made from rice fields in Thailand (cf.

sinensis). In Malaysia, Reid (1968) reported this species from swamps with

tall emergent grass, and specimens in the USNM from Malaysia were collected

from, ''overgrown swamp, grassy Swamp, marshy depression and a drain."

According to climatological maps (Ohman 1965) nearly all of the specimens

in the USNM from Assam, Cambodia, Malaysia, Thailand and South Vietnam

camefrom areas which receive 2,032 mm or more of rain per year, and most

over 2,540 mm. Over half of Thailand receives less than 2,032 mm per year

and these areas constitute the majority of the country under cultivation. The

wetter remaining areas contain the majority of the tropical wet and rain forest

reserves. Although the association of heavy precipitation and tropical wet for-

ests with crawfordi are probably only 2 of the factors that determine its distri-

bution this seems to be a definite correlation that should benefit future collec-

tors.

Most larval collections of this species in Thailand were made below 100 m

elevation, but one collection was made at 600 m from Khao Yai National Park

in Nakhon Ratchasima. Adults have been collected in Thailand from a Shannon

trap baited with man and light, biting man and biting buffalos. All the man

biting collections were made outside houses.

MEDICAL SIGNIFICANCE. Identification problems in the hyrcanus group

in the past have left few valid records for use in vector interpretations. Cur-

rently, there is no evidence cvawfordi is involved in the transmission of hu-

man pathogens in Thailand. Reid et al. (1962) and Wharton, Laing and Cheong

(1963) were able to infect crawfordi with Brugia malayi from human reservoirs,

but decided the species was probably too zoophilic or not abundant enough to

play an important vector role in filariasis areas in Malaysia. Confirmed

records show that considerable numbers of cvawfordi bit man in certain areas

of both Chon Buri and Chumphon provinces; the latter is known to harbor an

endemic focus of Brugia malayi (Harinasuta et al. 1964). The vector status

in that area is still undetermined, and other potential vectors are present, i.e.

08 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Anopheles campestris and Mansonia species.

Wharton et al. (1964) found that cvawfordi was much more attracted to man

than to monkeys at ground level, and only slightly attracted to monkeys on

canopy platforms. During the same study cvawfordi was found a poor labora-

tory host for Plasmodium cynomolgi bastianellii. These same authors did not

find malaria parasites in wild collected crawfordi.

ANOPHELES (ANOPHELES) NITIDUS HARRISON, SCANLON AND REID

(Figures 8, 19, 20, 21)

Anopheles hyrcanus nigerrimus type C of Crawford 1938: 51 (P*).

Anopheles (Anopheles) indiensis of Reid 1953: 29 (o*, 2*, P*, L*, E*); Reid

1968: 98 (o*, 9*, P*, L*, E*); Rattanarithikul and Harrison 1973: 7

(L*),

Anopheles (Anopheles) nitidus Harrison, Scanlon and Reid 1973: 266.

Adults are easily recognized by the combination of: wide pale bands on the

hindtarsus, a bright, sharply defined wing pattern like that seen on cvawfordi,

pale scales on the basal 0.33 of the costa and on the preapical dark mark on

vein Rj and a patch of dark scales on the humeral crossvein. Males have a

basal pale band on palpal segment 3. The larvae and pupae are more difficult

to separate and the characters given below as well as those in the keys are

necessary for their identification. Like sinensis except:

FEMALE (Figs. 8, 19) Head. Palpus with 4 distinct pale bands, 2 basal

bands are usually narrow and well defined, usually some pale scales on segment

2. Thorax. Scutal integument ovay-brown, with faint median and lateral dark

longitudinal lines; eye spots particularly well developed; pleural setae: 2 - 5

propleural, 2 - 4 spiracular, 7 - 10 prealar, 3 - 5 upper and 4 - 7 lower sterno-

pleural and 5 - 9 upper mesepimeral. Wing. Costa with small subcostal and

preapical pale spots, and with scattered pale scales on basal 0.33, often form-

ing small humeral pale spot; remigium usually with a proximal and distal black

spot separated by white scales, with white scales on anterior margin; humeral

crossvein with patch of dark scales; vein R with few pale scales on basal 0. 33,

well developed sector pale spot and subcostal pale spot usually incomplete; R,

with scattered pale scales between subcostal and preapical pale spots, tip

rarely pale scaled; Ro -Ro with distinct basal mark then white scales until

dark mark at fork; R9 P paatly completely dark scaled; R4,5 usually with few

scattered dark scales on pale middle section; M with scattered dark scales on

basal 0.33, then short dark marks thereafter; Cu sometimes with several dark

scales at absolute base, then pale scales followed by short "basal" dark mark

separated by at least its own length from upper dark mark on vein 1A; 1A with

upper and apical dark marks short, approximately equal in length; apical dark

mark on 1A may be longer or shorter than apical mark on Cug; apical pale

fringe spot extending from Rg to slightly below R4,5; posterior wing margin

usually with pale fringe spot at Cug. Legs. Coxae with pale scales, upper

portion of midcoxa with 0 - 1 seta. Foreleg: femur dark scaled; tibia dark

scaled above and pale ventrally; tarsomere 1 dark scaled above and pale ven-

trally, with short apical pale band; tarsomeres 2 - 3 dark scaled with apical pale

bands, that on segment 2 may approach 0.5 segment length; tarsomeres 4 - 5

dark scaled. Midleg: femur dark dorsally, pale ventally except at apex; tibia

with small basal pale band, dark dorsally, pale ventrally; tarsomere 1 - 3 dark

scaled with short apical pale bands, that on segment 2 approximately 0.2 seg-

Harrison and Scanlon: Subgenus Anopheles in Thailand o9

ment length. Hindleg: femur dark dorsally, pale ventrally except apex; tibia

with small basal and apical pale bands, dark dorsally and pale ventrally nearly

to apical pale band; tarsomeres 1 - 5 black except apical pale bands on tarso-

meres 1 - 4 and basal pale bands on tarsomeres 4 - 5 (infrequently 3); widest

hindtarsal band at juncture of tarsomeres 3 - 4, apical band of tarsomere 3

is approximately 0.25 tarsomere length and basal band on tarsomere 4 is 0.17 -

0.20 segment length; basal pale band on tarsomere 5 very rarely absent. Abdo-

men. Often few narrowscales on the posterior lateral corners of tergum VIII.

MALE (Fig. 19) Head. Palpus with pale scales on segment 2; segment 3

with distinct basal pale band and frequently with scattered pale scales; seg-

ments 4 - 5 with patches of pale scales. Wing. Costa nearly always with pale

humeral spot, besides scattered pale scales on basal 0.33. Genitalia. Basi-

meres with numerous broad pale scales; ventral lobe of claspette with 2 setae,

mesal seta distinctly longer than lateral seta and club on dorsal lobe; dorsal

lobe of claspette with fused club on inner half formed from 2 - 3 stems, outer

half of lobe contains 2 flattened leaflet-like spines adjacent to fused club;

aedeagus with 2 - 3 pairs of leaflets, largest with teeth along one edge and

sometimes with basal tooth.

PUPA (Fig. 20). Integument generally very lightly pigmented. Cephalo-

thorax. Wing case may have light spots and lattice pattern; apical joint and tip

of antennal case usually darkly pigmented. Trumpet. Rim uniform and thin

without thickened areas or vertical wrinkles. Abdomen. Usually almost color-

less; alveoli of setae such as seta 2 rarely surrounded by dark pigment; seta 1

and 9 on segments III - IV with numerous branches; 5-V with 30 or more bran-

ches; 9-II - VII lightly pigmented; 9-VIII with numerous branches. Paddle.

Refractile border approximately 0.65 - 0.80 of paddle length; paddle base

rarely with pigmentation.

LARVA (Fig. 21) Head. Seta 3-C with 70 or more branches; 5, 6-C with

not more than 18 branches; 8, 9-C with 11 - 21 and 6 - 13 branches respectively.

Thorax. Seta 1-P simple or with 2 - 3 thin branches near tip; 2-P with 11 - 19

branches; 11-P with 4 - 6 stout branches from base; 14-P with 5 - 11 branches;

1-M with 19 - 32 branches; 4-M with 6 - 10 branches which usually arise close

together near base; 3-T with flattened unpigmented leaflets. Abdomen. Seta 1

with flattened unpigmented leaflets on segments I - II, and well developed

(palmate) with darkly pigmented leaflets on segments III - VII, pigment not ex-

tending onto filaments; 2-II with 8 - 14 branches; 6-I, III with 16 - 22 and 15 -

20 branches respectively (rarely 20 or more); 9-III with 10 - 16 branches; 5-IV

- V with 5 - 9 branches each; 9-VI with 5 - 9 branches; 13-IV with 6 - 12 short

branches (cf. sinensis); tergal plate VIII elliptical, with posterior margin con-

vex; pecten with 5 - 7 long teeth; seta 1-X usually weakly developed, shorter

than saddle; posterior tergal plates with equivalent or more pigment than an-

terior tergal plates.

TYPE-DATA. The type-specimens for nitidus were selected from speci-

mens collected in Selangor, Malaysia, where Reid (1953) did considerable work

on what he called indiensis. The holotype 2, deposited in the BMNH, is in ex-

cellent condition, has associated immature skins on a slide andhas the follow-

ing label data: (1st label) - ''Selangor, 5th m. Klang Rd., 7-11-1950, J. A.

Reid"; (2nd label) - "bred in laboratory from known mother", and rearing num-

ber '28/9"'; and (3rd label) - is Reid's indiensis identification label. The “ al-

lotype, with associated immature skins on a slide, is also in the BMNH and has

the same label data as the holotype, except the rearing number is ''28/10".

Two & and 5 paratypes from Selangor, Malaysia, are deposited in the USNM.

One “ with genitalia intact, associated immature skins on a slide and rearing

60 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

number ''28/8" is from the same collection as the holotype and allotype. One 2

with associated immature skins on a slide and rearing number ''454/68" is from

"Kepong."’ One % with genitalia intact and collection number "232" is from

"Ampang Rd., K. L.'' Two 2 with collection number "16984" are from ''Sungei

Tua.'' The 5 paratypes just noted were collected by Reid between 1941 - 1951,

while the remaining 22 paratypes have collection number ''804"' and were col-

lected in ''Ulu Langat" in 1967 by personnel from the University of Malaysia.

DISTRIBUTION (Fig. 20). This species apparently has a discontinuous

distribution in Thailand, much like that of crawfordi. One larva which con-

forms to nitidus was collected in Chiang Mai and in addition, several adults

were collected in Mae Hong Son. This extends the range far north of the cur-

rent distribution concept. It is possible that nztidus extends into northern Thai-

land via the wet forest on the Bilauk Taung Mountain range which separates

Burma and Thailand, or it will be found in isolated pockets in the north. Else-

where, in the central valley near Bangkok, the southeast and the peninsular

region, nitidus can be locally abundant. Generally, it is a foothill species with

larvae occurring up to approximately 300 m elevation, but usually lower. Speci-

mens (11%, 752, 35 larvae and 54 larval and pupal skins) in the USNM were

seen from the following provinces of THAILAND: Chanthaburi, Chiang Mai,

Chon Buri, Krabi, Nakhon Si Thammarat, Narathiwat, Phangnga, Phuket,

Prachin Buri, Ranong, Rayong, Satun, Surat Thani, Trang, Trat and Yala.

Additional specimens were examined during 1968 - 69 from Chumphon, Mae

Hong Son, Nakhon Nayok and Sara Buri Provinces. In addition to the above,

23%, 632, 3 larvae and 14 larval and pupal skins in the USNM were examined

from: CAMBODIA, INDIA (Assam), INDONESIA (Sumatra), MALAYSIA (Penin-

sular Malaysia) and SOUTH VIETNAM. An additional 30, 169 and 4 larval

and pupal skins in the BMNH were examined from: MALAYSIA (Peninsular and

Sarawak), NORTH VIETNAM and THAILAND (Trang). Other specimens (1°

and 112) from CAMBODIA were found in the ORSTOM collections.

TAXONOMIC DISCUSSION. This species was previously called indiensis

until Harrison, Scanlon and Reid (1973) determined that indiensis sensu Reid

(1953, 1968) does not occur in Madras, India, the type-locality of indiensis

Theobald. Since the type of indiensis Theobald 1901, is lost or non-extant

and no other specimens are known, indiensis Theobald was synonymized under

nigerrimus Giles 1900, and indiensis of Reid was renamed nitidus. The sharp

appearance of the wings, scutal eye spots, pale scales on the base of the costa,

short basal dark mark on vein Cu and broad fore-and hindtarsal pale bands,

make adult nitidus quite distinct. In addition, the pale band on the base of male

palpal segment 3 should readily separate nitidus from crawfordi males (see

taxonomic discussion under cvawfordi). Male nitidus nearly always exhibit a

pale humeral spot on the costa. Some females of nitidus and nigerrimus are

difficult to separate, however, an abundance of pale scales on Rj between the

subcostal and preapical pale spots is usually indicative of nitidus. The wide

foretarsal pale bands are also indicative of nitidus, as peditaeniatus is the only

other species with comparable bands.

The immature stages of nifidus are more difficult to distinguish. The pupa

is most often confused with that of nigerrimus and pursati, but can also be mis-

identified under sinensis. All these species have a thin uniform rim on the trum-

pet, but, simensis can be separated from the others by fewer branches on seta

o-V. Beyond this point nitidus can be separated by the characters given in the

key. Larvae most often confused with nitidus are argyropus, nigerrimus, craw-

fordi and lesteri pavaliae. Fortunately, branching of seta 8-C will usually sep-

arate the 2 last named species from nitidus, if not, then the branching of seta

Harrison and Scanlon: Subgenus Anopheles in Thailand 61

9-ITI is quite reliable. If this character is not convincing, then seta 13-IV on -

_ nitidus is tufted with 6 - 12 short branches, while this seta on crawfordi and

lesteri pavaliae has*3 - 9 long stout branches. Separation of nitidus larvae

from nigerrimus is more difficult. Besides the key characters, seta 9-VI

has: 5 - 9 (nitidus) and 2 - 4 (nigerrimus) branches. The characters used in

the key to separate argyropus and nitidus overlap to some degree, but should

differentiate 95% of Thailand specimens.

Little variation was noted in the adults of nitidus. The hindtarsal banding

was much more consistant throughout this species range than that found for

argyropus, nigerrimus and peditaeniatus. One frequent variation is the pres-

ence or absence of a distinct pale humeral spot on the costa. Although nearly

always present on males, this character is less constant on females, but

usually present on at least one wing and infrequently absent.

BIOLOGY. In Thailand this species is usually found in foothill, non-rice

agricultural areas which have been cleared of forest in the recent past and

still have some primary or secondary forests nearby. Although Reid (1968)

listed larvae commonly found in rice fields in Malaysia, larvae were rarely

found in this habitat in Thailand. Apparently cool still water, emergent and/or

floating vegetation and partial shade are among the basic larval requirements.

Confirmed larval habitats in Thailand include: swamps (including nipa palm

swamp), jungle bog, seepage marsh, large mine pits, large rock pool, stream

pool, ground pool, roadside ditch, rice field and elephant footprint. Nearly all

of these had light to heavy shade and emergent and/or floating vegetation. Con-

centrations of green algae were mentioned in several collections. This habitat

pattern may explain the apparent absence of niztidus in the immediate Bangkok

area, however, short trips to foothill areas in the north (Sara Buri) or to the

south (Chon Buri) will put the collector back into localized nitidus populations.

In localized foci nitidus may bite man in some numbers. Ata newly cleared

land settlement in Chon Buri Province, this species made up the largest com-

ponent of the "hyrcanus group"’ specimens reported by Scanlon and Sandhinand

(1965). Numerous adults have also been collected biting man in a roadside park

in Sara Buri and in small villages near rubber plantations in Nakhon Si Tham-

marat. Aside from human biting collections, this species has also been col-

lected biting domestic bovines, by light trap, Shannon traps baited with man

and light and in CO» net traps baited with dry ice. Reid (1968) notes that nztidus

(as indiensis) in Malaya appears to feed mostly on cattle. Currently, data from

Thailand are insufficient for confirmation of a zoophilic feeding behavior.

MEDICAL SIGNIFICANCE. Malaria research in Malaya (Wharton et al.

1963, 1964) did not find natural malaria infections in wild caught vztidus (as

indiensis). This research also showed few adult nitidus attracted to monkey

baited traps. Furthermore, Warren et al. (1963) showed experimentally that

nitidus (as indiensis) had an extremely low susceptibility to Plasmodium cyno-

molgi bastianellii. Reid et al. (1962) and Wharton, Laing and Cheong (1963)

did not find human filarial infections in Malayan nitidus (as indiensis), although

the former showed this species to be a low to moderate experimental vector of

periodic Brugia malayi. Likewise, there are no data from Thailand which sug-

gest nitidus is a natural vector of human pathogens.

ANOPHELES (ANOPHELES) LESTERI PARALIAE SANDOSHAM

(Figures 9, 22, 23, 24)

Anopheles (An.) hyrcanus subsp., near sinensis of Colless 1948: 86 (“*, @*, L*).

62 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Anopheles (An.) lesteri Malayan form of Reid 1953: 7, 39 (*, @*, P*, L*, E*).

Anopheles (An.) lesteri paraliae Sandosham 1959: 189 (¢, L); Reid 1963: 101;

Reid 1968: 105 (o'*, 2*, P*, L*, E*).

Adults have narrow hind tarsal bands (usually apical only), humeral cross-

vein without a scale patch, short apical pale wing fringe spot and a long dark

mark on the base of vein Cu. Males lack a basal pale band on palpal segment

3. The pupal trumpet rim has thickened areas with a distinct sawtooth margin.

The larvae are darkly pigmented, have reduced branching of seta 8-C and have

7 or more long pecten teeth. As for sinensis except:

FEMALE (Figs. 9, 22) Head. Palpus usually with all 4 pale bands distinct,

apical 2 bands may fuse, infrequently a small number of pale scales on seg-

ment 2. Thovax., Integument of scutum color ash-gray, with 2 short submedi-

an lines on anterior portion; eye spots present, not as distinct as in crawfordi

and nitidus; pleural setae: 3 - 6 propleural, 2 - 4 spiracular, 5 - 8 prealar,

3 - 6 upper and 3 - 8 lower sternopleural and 5 - 11 upper mesepimeral. Wing.

Costa with small subcostal and slightly larger preapical pale spots, no other

pale scales present; remigium usually dark scaled or with narrow anterior row

of pale scales; humeral crossvein without scale patch, rarely with 1 - 2 small

scales; vein R nearly entirely dark scaled out to distinct sector spot, subcostal

pale spot frequently incomplete or absent; Ry completely dark scaled between

subcostal spot, when present, and preapical pale spot, tip dark scaled; Re

Roi3 usually entirely dark scaled to fork; Ry rarely entirely dark scaled; R3

with pale central portion; R4,5 largely pale in center, frequently with scat-

tered dark scales on distal half; vein M dark scaled to fork or with pale area

before fork; extreme base of Cu usually with few black scales followed by dis-

tinct pale scaled area or area with scattered pale scales, then long dark mark

which approaches within its own length or less of upper dark mark on vein 1A;

vein 1A with 2 distinct dark marks, upper mark short, apical mark may be

longer than upper mark and is usually longer than apical dark mark on Cug;

apical fringe spot very small, widest extension from R, to Rg, but usually

from Rg to Rg; no pale fringe spot on posterior wing margin. Legs. Midcoxa

without pale scales or rarely 1 - 2, and with 3 - 5upper setae. Foreleg: dark

scaled except for narrow apical pale bands or dorsal patches on tarsomeres

1 - 3 and infrequently very narrow apical pale patch on 4; apical bands on tar-

someres 1 - 2 longest, usually complete ventrally, not more than 0.2 length of

tarsomere; pale scales on tarsomeres 3 - 4 dorsal and lateral only, not con-

nected ventrally. Midleg: dark scaled except for narrow apical pale bands on

tarsomeres 1 - 4, bands usually incomplete, not connected ventrally. Hindleg:

dark scaled except white scales dorsally on apex of tibia and complete, narrow

apical pale bands on tarsomeres 1 - 4; base of tarsomeres 4 - 5 infrequently

with small dorsal patch of pale scales.

MALE (Fig. 22) Head. Palpus with well developed lineof pale scales on

inner surface of segments 2 - 3, base of segment 3 without pale band, pale

scales on segments 4 - 5 variable from entirely pale dorsally to pale basal and

apical bands. Wing. Base of costa without pale scales. Genitalia. Basi-

meres without pale dorsal scales or rarely with 1 - 2 gray or black scales;

ventral lobe of claspette with long mesal seta and shorter lateral seta; dorsal

lobe of claspette with fused club on inner half formed from 3 - 4 stems, outer

half of lobe with 2 flattened leaflet-like spines adjacent to fused club; aedeagus

with 4 - 5 pairs of leaflets, largest with well developed teeth, even on base.

PUPA (Fig. 23). Integument darkly pigmented. Cephalothorax. Wing case

brown with barred or rectangular pattern on veins; antennal case dark brown at

Harrison and Scanlon: Subgenus Anopheles in Thailand 63

joints and tip. Trumpet. Darkly pigmented, with dark thickened areas bear-

ing saw-toothed edge. Abdomen. Usually with dark spot on center of terga

III - VII and several other smaller dark areas; area surrounding alveoli of

setae, such as seta 2, always darkly pigmented; seta 9 lightly pigmented on

all segments, with numerous branches on segment VIII; seta 1, 50n IV - V

with relatively few branches; 1-VI with 5 - 10 branches; 5-V with 12 - 30

branches; male genital lobe darkly pigmented on distal 0.5. Paddle. Base

darkly pigmented; refractile border 0.50 - 0.65 paddle length; seta 1 simple

or forked at tip.

LARVA (Fig. 24) Head. Darkly pigmented; seta 3-C with 43 - 70 branches;

D, 6-C with 10 - 16 and 13 - 15 branches respectively; 8, 9-C with 5 - 11 and

6 - 11 branches respectively; 15-C with 2 - 7 branches. Thorax. Seta 1-P

simple or usually with 2 - 6 fine branches on distal half; 11-P with 3 - 6 stout

branches from base; 14-P with 5 - 8 branches; 4-M with 3 - 8 branches from

erect central stem; 3-T with flattened unpigmented leaflets. Abdomen. Seta 1

with flattened unpigmented leaflets on segments I - II, well developed (palmate)

and darkly pigmented on III - VII, pigment not extending onto filaments; seta

6-I, III with 17 - 21 and 17 - 23 branches respectively; 5-II with 6 - 10 branches;

9-III with 5 - 10 branches; 13-IV with 4 - 9 long branches; pecten plate with

7 - 10 long teeth; seta 1-X strong, longer than saddle; anal papillae usually

shorter than saddle.

TYPE-DATA. Sandosham (1959) apparently based the naming of this sub-

species on Reid's (1953) description of lestevi ''Malayan form", but did not

designate a type even though Reid had deposited specimens of the ''Malayan

form" in the BMNH. Reid (1963) designated a female in the BMNH with asso-

ciated pupal and larval skins, as lectotype for this subspecies. The lectotype

is in excellent condition with only the right antenna missing, and has the fol-

lowing labels: "Selangor, Klang, 1940, J. A. Reid''; ''No. 16715/11, larvae

in"; "Anopheles (A.) lestevi Baisas & Hu, Det. J. A. Reid"; ''Liason pelts in

slide collection"; 'lectotype"' in Reid's handwriting; and a BMNH lectotype

label.

DISTRIBUTION (Fig. 23). Scanlon et al. (1968) listed this subspecies from

9 provinces of Thailand; however, further examination of the specimens in the

USNM revealed that the specimens identified as lesteri paraliae from Ayutthaya,

Nonthaburi, Pathum Thani and Phra Nakhon were actually pursati and the single

specimen on which the Chiang Mai record was based was a female sinensis with

an unusually short apical fringe spot. Apparently lestevi pavaliae has a low

elevation-coastal distribution in Thailand, like that in Malaysia (Reid 1968).

Specimens (19°, 21°, 30 larvae and 46 larval and pupal skins) in the USNM

were examined from the following provinces of THAILAND: Chumphon and

Surat Thani. Additional specimens (130, 222, 10 larvae and 10 larval and

pupal skins) in USNM were examined from: MALAYSIA (Peninsular Malaysia),

SINGAPORE and SOUTH VIETNAM. Beside the lectotype female, 3%, 5° and

2 pupal and larval skins in the BMNH were examined from: MALAYSIA and

THAILAND (Trang). Additional literature records refer to BRUNEI (Colless

1948).

TAXONOMIC DISCUSSION. This is one of the easiest members of the

eroup to identify in the adult stage. The very narrow tarsal bands, narrow

apical fringe spot on the wing, extensive dark areas on the costa and vein

R-Rj,, long basal dark mark on vein Cu and absence of scales on the humeral

crossvein, is a very distinct combination of characters. Even when the apical

wing fringe is missing the above characters should be indicative. Apparently,

pursati is the species most frequently confused with lesteri paraliae, particu-

64 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

larly since both have a narrow apical fringe spot on the wing and narrow tarsal

bands. However, a mostly pale remigium, a patch of scales on the humeral

crossvein, a short basal dark mark on vein Cu and the presence of pale scales

on the midcoxa should readily identify pursati. Beside these characters, pur-

sati is the smallest member of the group, while lesteri paraliae is usually

nearly equivalent in size to sinensis. Males of the nigerrimus subgroup (includ-

ing pursati) all exhibit a basal pale band on palpal segment 3, numerous pale

scales on the basimeres and usually pale scales on the basal 0.33 of the costa.

Male lesteri paraliae do not show these characters and thus, by elimination

become the only remaining species with a narrow apical fringe spot on the wing.

Differentiation of the immature stages is more difficult, with only approxi-

mately 90% currently separable. The pupa can be readily identified as belong-

ing to the /esteri subgroup on the basis of the thickened saw-tooth areas on the

trumpet rim. Further separation, from peditaeniatus, is also simple on the

basis of the paddle margin and seta 9-VIII, but final separation from crawfordi

is frequently difficult. The larva of lesteri paraliae is frequently the most dif-

ficult stage to identify. Past confusion was most often noted between this spe-

cies and sinensis, nitidus and crawfordi, and the reader should refer to the

taxonomic discussions of those species for explanations and possible solutions.

Several other points may be of value in identifying larvae of lesteri paraliae,

but because of potential overlap, should be used with extreme caution: the head

capsule is consistently darkly pigmented (brown-black); suggested distribution

in Thailand is confined to coastal-low elevation regions; and the immature stages

are apparently capable of tolerating high salt concentration; thus, lestevi pava-

liaeé may be the only species in the group found in brackish water.

Anopheles lesteri was described from the Philippines and southern China

and is currently known from the Philippine islands of Leyte, Luzon, Mindanao,

Mindoro, Negros and Samar, and from China (in part), Japan, South Korea and

the Ryukyus (Okinawa). The latter record is based on specimens in the USNM

and is of considerable interest since this subspecies has not been reported from

Taiwan. The nominate subspecies differs from paraliae in the greater develop-

ment of the apical fringe spot, the other stages currently being indistinguishable

from pavaliae. Because of geographic isolation and the great morphological

similarity, Reid (1953) indicated that the narrow fringe form probably deserved

subspecific rank, but did not formally propose this. Sandosham (1959) appar-

ently based his name pavaliae entirely on Reid's description but failed to desig-

nate a type, consequently, Reid (1963) later selected a lectotype and paratypes

| sic ]from material in the British Museum. Thus, the nomenclatorial status of

the narrow fringe type of lestevi is solidly established. Its biological status

is another matter.

As noted in the remarks on distribution, lestevi paraliae is confined to

coastal areas of Malaysia (Peninsular Malaysia, Sabah and Sarawak) Brunei,

South Vietnam and Thailand. Reid (1968) stated that areas of overlap between

lesteri (sensu stricto) and pavaliae had not been found, and this still appears to

be true. The islands of Balabac and Palawan appear to form a definite link

between North Borneo (Sabah) and the larger Philippine islands, therefore, one

might expect to find intermediates there; however, lestevi has not been recorded

from either island. Ho et al. (1962) reported one of the forms they studied to be

equivalent to lesteri lesteri; however, Feng (1964) and Ma (1964, 1968b) felt

that possibly none of the 5 species masquerading under the name sinensis in

China during the past is actually lesteri. Harrison (1973a), after examining

reared specimens with associated skins from the Philippines and southern

China (Hong Kong), supported the contention (Baisas and Hu 1936; Ho et al.

Harrison and Scanlon: Subgenus Anopheles in Thailand 65

1962) that lesteri lesteri occurs on both mainland China and the Philippines.

The possiblity that the mainland segment of the nominate subspecies is geo-

graphically isolated from lesteri pavaliae is very likely, but more specimens

are needed from southern China, North Vietnam and South Vietnam for con-

firmation. As can be seen, many problems will have to be solved before we

will understand the Jestervi subgroup.

There is no doubt that paraliae from Malaysia, South Vietnam and Thailand

represents a distinct taxonomic entity. For the moment it is probably best to

continue regarding it as a subspecies of lesteri, at least until the Palawan

fauna is examined in more detail. One cannot help, however, but suspect that

paraliae is a good species restricted to the coastal areas of mainland Southeast

Asia and Borneo. |

BIOLOGY. All the records of 1. pavaliae from Thailand are from coastal

areas and collected near sea level, except for a couple of collections up to 150

m elevation. Immatures are typically found in semi-permanent to permanent

bodies of water with open sunlight to moderate shade. Floating and emergent

vegetation is usually present. The water can be clear or stained and may be

brackish, but should be cool; consequently, /. pavaliae will not normally be

found in rice fields. Immatures were collected in Thailand from: large

marshy depression, open "Swamp", nipa palm swamp, large ground pool and

large rock pool. The immatures from Malaysia and Singapore came from:

earth drain, marshy depression, drain w/"peaty"’ water, unused well and

pond with Fichornia species.

Colless (1948) originally collected 1. paraliae biting man in Brunei, and

Macdonald et al. (1967) collected small numbers biting man in Sarawak. Reid

(1961) and Reid et al. (1962) found J. pavaliae strongly zoophilic and reluctant

to enter houses in Malaya. In Thailand adults have been collected biting man,

but in small numbers.

MEDICAL SIGNIFICANCE. There is no evidence that pavaliae is a vector

of human disease in Thailand. Reid et al. (1962) found Dirofilaria sp. and

Setaria sp. in wild caught paraliae, and in experiments found it to be moder-

ately receptive to Brugia malayi. Warren et al. (1963) showed experimentally

that paraliae was much more susceptible to Plasmodium cynomolgi bastianel-

lii, than other members of the hyrcanus group. Wharton et al. (1964) showed

that pavaliae would feed on monkeys in the forest canopy. Since malarial para-

sites have not been found in this subspecies, it is not considered a vector of

malaria. More work is needed on pavaliae in Thailand as little is known of its

biting behavior. Its role in filariasis is unknown although it is found in areas

(Surat Thani and Chumphon) where Brugia malayi occurs (Harinasuta et al.

1964).

The report by Ho et al. (1962) that the primary malaria vector in the

Yangtze Valley of China is lesteri lesteri not sinensis, is most significant.

They found Jestevi markedly anthropophilic, entering houses readily. These

observations are apparently the opposite of the behavior ascribed to lesteri

lesteri in the Philippines. However, before its behavior can be understood

much more work is needed on this species in both areas.

ANOPHELES (ANOPHELES) NIGERRIMUS GILES

(Figures 9, 25, 26, 27)

Anopheles nigerrimus Giles 1900: 162 (9*).

Anopheles sinensis indiensis Theobald 1901: 145.

66 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Anopheles bentleyi Bentley 1902: 15 (9*).

Anopheles minutus Theobald 1903b: 91 (9).

Anopheles (Anopheles) hyrcanus var. nigerrimus of Edwards 1932: 41 (in part).

Anopheles hyrcanus var. williamsoni Baisas and Hu 1936: 222 (P, L*).

Anopheles venhuisi Bonne-Wepster 1951: 284 (**, 9*, P*, L*), (nom. nov.

for hyrcanus var. X Venhuis 1939).

Anopheles (An.) nigerrimus Giles, Reid 1953: 24 (o*, 9*, P*, L*, E*); Reid

1963: 100: Reid 1968:.92.(o*,.9*% P*, 1+, B*),

The adults are generally similar to peditaeniatus and sinensis, but have a

darker appearance. The basal 0.33 of the costa usually has some pale scales,

the humeral crossvein has a scale patch, the basal dark mark on vein Cu is

long and hind tarsomeres 4 and/or 5 usually (78%) have a pale basal patch or

band. The male has a pale basal band on palpal segment 3. The pupa is

usually darkly pigmented with a thin uniform trumpet rim and setae 1 and 5

with numerous branches on most abdominal segments. The larva has 12 - 24

branches on seta 8-C and very few branches on seta 5-IV. Separation of both

immature stages from those of other members of this group may infrequently

prove difficult. Like sinensis except:

FEMALE (Figs. 9, 25) Head. Palpus with 4 distinct pale bands, infrequent-

ly apical and subapical pale bands confluent, usually a mesal patch of pale

scales on segment 2. Thorax. Scutal integument dark gray, with poorly de-

fined dark longitudinal stripes and eye spots; pleural setae: 4 - 10 propleural,

2-4 spiracular, 5 - 11 prealar, 3 - 4 upper and 4 - 7 lower sternopleural and

4-7 upper mesepimeral. Wing. Costa dark with subcostal and preapical pale

spots, and usually several scattered pale scales slightly distal to humeral

area; basal 0.33 of costa rarely with small humeral pale spot on one wing or

completely dark scaled; remigium largely dark scaled; humeral crossvein with

distinct patch of dark scales; vein R with few scattered pale scales on basal

area and distinct sector spot; subcostal spot on R often divided or with an in-

definite area of mixed pale and dark scales; R, usually without pale scales

between subcostal and preapical pale spots, tip with dark scales; Ro- R943

usually dark nearly to fork; Ro infrequently entirely dark scaled; R4ai5

infrequently with scattered dark scales on middle portion; base of M dark,

with pale area just before fork; extreme base of vein Cu dark scaled, usually

followed by short pale area, then long dark mark at least the length of pre-

ceding pale mark (when present); long basal dark mark on Cu approaching

within its own length or less of upper dark mark on vein1A; apical dark mark

on Cug rarely as long as corresponding mark on 1A; vein 1A normally with 2

dark marks, infrequently these connect making apical 0.5 of vein entirely dark

scaled; apical pale fringe spot long, from R, to beyond R45; pale fringe spot

at vein Cug on approximately 50% of the specimens. Legs. Coxae with white

scales, 1 - 3 upper midcoxal setae. Foreleg: Femur dark scaled; tibia dark

scaled dorsally, pale ventrally; tarsomere 1 pale scaled ventrally; tarso-

meres 1 - 3 with apical pale bands, that on segment 2 may approach 0. 33

length of segment; tarsomeres 4 - 5 dark scaled. Midleg: Femur and tibia

dark dorsally and pale ventrally, apex of tibia usually with few pale scales;

tarsomere 1 frequently with venter pale scaled on basal 1/3 - 1/2; tarsomeres

1 - 3 with narrow apical pale bands; tarsomeres 4 - 5 dark scaled. Hindleg:

Femur and tibia dark dorsally, pale ventrally, apex of tibia with distinct dor-

sal pale spot; tarsomeres 1 - 4 with apical pale bands; pale markings on base

of tarsomeres 4 - 5 highly variable (see taxonomic discussion). Abdomen.

Integument dark, usually with few dark scales on the posterolateral corners of

Harrison and Scanlon: Subgenus Anopheles in Thailand 67

tergum VIII.

MALE (Fig. 25) Head. Palpus with pale scales mesally on segments 2 - 3

and distinct pale basal band on segment 3, segment 4 with distinct basal pale

band, segment 5 nearly entirely pale scaled dorsally, except lateral edge.

Wing. Essentially as on female, except wing may appear paler, and basal

dark mark on Cu may be slightly shorter. Genitalia. Basimeres with numer-

ous pale scales; ventral lobe of claspette with 2 setae, mesal seta distinctly

longer than lateral seta and fused club on dorsal lobe; dorsal lobe of claspette

with fused club on inner half formed from 2 - 3 stems, while outer half of lobe

contains 2 flattened leaflet-like spines adjacent to fused club; aedeagus with

2 - 3 pairs of leaflets, largest may have teeth along one edge, no basal tooth

apparent.

PUPA (Fig. 26) Generally darkly pigmented. Cephalothorax. Wing case

usually with mottled pattern of dark pigmentation; leg case along anterior mar-

gin of wing sheath uniformly dark brown; outer half of antennal case darkly

pigmented, with distinct dark marks at each joint. Tvumpet. Darkly pig-

mented, without thickened saw-tooth areas on rim, but may have dark border

area delineating thin, uniform rim. Abdomen. Seta 9 lightly pigmented on all

segments, with many branches on segment VII; area surrounding alveoli of

abdominal setae, particularly seta 2, always darkly pigmented; terga III - VII

usually with a cental dark spot, as well as other dark spots; setae 1 and 5 on

III - V with numerous branches; 5-V with 30 or more branches; 1-VII with 1 -

7 branches, usually 2 - 4; 6-VII mesal to seta 9-VII; male genital lobe darkly

pigmented on distal half. Paddle. Base darkly pigmented; refractile border

approximately 60 - 70% of paddle length; 1-P simple or split near apex into

2 - 4 equal branches.

LARVA (Fig. 27) Head. Moderately pigmented; seta 3-C with 70 or more

branches; 5, 6-C with 11 - 18 and 11 - 17 branches respectively; 8, 9-C with

12 - 24 and 8 - 14 branches respectively. Thorax. Seta 1-P simple or with

2 - 4 branches on distal half; 11-P with 4 - 8 stout branches from base; 14-P

with 4 - 8 branches, usually 5 or more; 1-M with 21 - 28 branches; 4-M with

o - 10 stiff, erect branches from central stem; 3-T with flattened unpigmented

leaflets. Abdomen. Seta 1 with flattened unpigmented leaflets on I - II, well

developed (palmate) and darkly pigmented leaflets on III - VII, pigment not

extending onto filaments; seta 6-I, III with 15 - 20 and 12 - 20 branches re-

spectively; setae 5-III - IV with 4 - 8 and 2 - 4 branches respectively; 9-VI

with 2 - 4 branches; pecten plate with 4 - 7 long teeth; 1-X approximately equal

to length of saddle.

TYPE-DATA. The holotype of nigerrimus is a female from Calcutta, India,

located in the BMNH, and with the following label data: (1st label) - "type";

(2nd label) - ''7-4-99, Calcutta, Dr. C. W. Daniels"; and (3rd label) - 'Ano-

pheles nigerrimus type G. M. Giles.'' The last label is hand penned, possibly

by Giles. This specimen is in very poor condition, with the following parts

missing: palps; antennae, except 5 basal flagellomeres on left side; right fore-

and midlegs from trochanter on; tarsomeres of right hindleg; Ts of left fore-

leg; 15 - 71, of left midleg; T9 - Pe of left hindleg; abdomen; right wing; and

fringe on left wing. The thorax is split lengthwise on top, and nearly all setae

or scales on the scutum are missing. The pleural areas are in fair shape.

The following minor additions and corrections can be added to Reid's (1953)

description of the holotype. The remigium has a line of pale scales on the an-

terior margin, humeral crossvein with only 2 dark scales, base of costa with-

out pale scales; and only 2 - 3 pale scales on preapical dark mark of Ry. The

problems involved in the identification of the type of nigerrimus and its synon-

68 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

yms were covered in depth by Reid (1953). Since then, sinensis indiensis

Theobald 1901, has been reassigned (Harrison, Scanlon and Reid 1973) asa

synonym of nigerrimus. Unfortunately, the types for indiensis, bentleyi,

williamsoni and venhuisi remain lost or unknown. Anopheles minutus Theo-

bald 1903b, with its type in the BMNH, has long been considered a synonym of

nigerrimus ; however, Reid (1968) lists it as such only for convenience and

says it may represent a distinct species. We agree with both Reid's feelings

and treatment of minutus, and thus, include it as a synonym of nigerrimus.

DISTRIBUTION (Fig. 26). This is one of the most widely distributed ano-

phelines in Thailand. Scanlon et al. (1968) list collections from 18 provinces,

but the examination of additional specimens indicates this species may be en-

countered anywhere in the country where suitable larval habitats are found.

For this reason, the map is marked to show a potential distribution through

the entire country. Specimens (100, 462, 15 larvae and 38 larval and pupal

skins) in the USNM were examined from the following provinces of THAILAND:

Chiang Mai, Chon Buri, Lampang, Nakhon Sawan, Nonthaburi, Pathum Thani,

Phra Nakhon, Phuket, Prachin Buri, Rat Buri, Rayong and Udon Thani. Addi-

tional specimens were examined in Thailand during 1968-70 from Buriram,

Chumphon, Lop Buri, Nakhon Nayok and Sara Buri provinces. From else-

where, 190%, 872, 10 larvae and 17 larval and pupal skins in the USNM were

examined from: BURMA, CAMBODIA, INDIA (Assam, Bihar and Punjab),

INDONESIA (Java and Sumatra), MALAYSIA (Peninsular Malaysia), SOUTH

VIETNAM and SRI LANKA. In the BMNH, 6 and 142 specimens (including

holotype) were examined from BANGLADESH, BURMA, INDIA, MALAYSIA

(Peninsular Malaysia and Sarawak), SOUTH VIETNAM and SRI LANKA. An-

other 1° and 252 specimens from CAMBODIA were found in the ORSTOM col-

lections. Additional literature records refer to: BRUNEI (Colless 1948);

CHINA, Hainan Island (Ho 1938b); many islands in INDONESIA (Bonne-Wepster

and Swellengrebel 1953); and MALAYSIA, Sabah (Colless 1948). All previous

records of nigerrimus in the Philippines refer to lestev1, peditaeniatus or

pseudosinensis. The above records of nigerrimus from Brunei, Sabah and

Sarawak are very interesting, since this species is apparently replaced in

the Philippines by pseudosinensis. The adult description given by Colless

(1948) for nigerrimus should probably apply to peditaeniatus instead, since

the following peditaeniatus-like characters were described: (1) Costa dark

except for subcostal and preapical pale spots; (2) liberal sprinkling of pale

scales along almost entire length of vein 1(R-Rj1); (3) humeral crossvein with-

out scales; and (4) fringe spot at vein Cuy rare. (cf. nigerrimus). The larval

description for nigerrimus given by Colless does not clarify this discrepancy,

for characters applicable to both species were given (i.e., outer clypeal

branches, and sutural branches).

The records of hyrcanus nigerrimus from Yunnan (Yao and Ling 1937) and

Hainan Island (Ho 1938b), China, probably refer to peditaeniatus (Reid 1953).

TAXONOMIC DISCUSSION. There has been considerable confusion sur-

rounding the status and identity of this species in Thailand and elsewhere in

Southeast Asia. Much of this confusion was settled by Reid (1953), particu-

larly for the Malayan populations. However, certain characters are highly

variable and unless studied elsewhere, our understanding of nigerrimus will

remain very limited. One of these variables is the extent of pale bands on

the hindtarsomeres. Reid (1953) noted that approximately 5% of the Malay

specimens lacked a basal pale band on tarsomere 4. Consequently, these

specimens would have only apical (sinensis-like) hindtarsal pale bands. Like

peditaeniatus (Reid 1953, 1963, 1968), the trend for reduction of the hindtarsal

Harrison and Scanlon: Subgenus Anopheles in Thailand 69

bands on nigerrimus is much more evident in the Indian subregion. During

1970-71 Smithsonian Institution personnel in Sri Lanka collected a variety of

aquatic insects. Among the mosquitoes were 46 peditaeniatus and 6 nigerrimus

adults. All of the nigerrimus exhibited sinensis-type banding, i.e., no basal

pale bands on hindtarsomeres 4 - 5. Some of the peditaeniatus also exhibited

reduced banding equivalent to the sznensis-type, but the majority had a narrow

pale band on the base of hindtarsomere 4. The sinensis-type hindtarsal band-

ing was also predominant on the other USMN specimens of nigerrimus from

India (Assam, Bihar), and Reid (1968) reported that a series of nigerrimus

from Rangoon, Burma also exhibited reduced tarsal banding. In Thailand this

one variable character is probably responsible for the majority of misidentifica-

tions noted among nigerrimus -sinensis during this study. Of 46 females avail-

able from Thailand, 42 had intact hindtarsi, permitting an analysis of this

character from 9 provinces, and from the extreme north to the south. Three

hindtarsal catagories were selected for notation: (1) tarsomeres 4 - 5 with

pale basal band or patch; (2) only tarsomere 4 with a pale basal band or patch;

and (3) tarsomeres without pale basal bands or patch, sinensis-type banding.

Table 3 shows the frequency distribution of this variation.

TABLE 3. Hindtarsal variation on Thailand Anopheles nigerrimus females

Tarsomere 4 and 5 with Tarsomere 4 only with Tarsomeres without pale

pale basal band or patch pale basal band or patch basal scales

Number 14 20 8

Percent 30.0 47.6 19.1

Nearly 20% of the specimens exhibited sinensis-type hindtarsal banding and

only 33% had basal pale scales on both hindtarsomeres 4 - 5. The 48% in the

middle category may prove of value in separating certain specimens from nzf7-

dus and peditaeniatus, which in Thailand nearly always have pale scales on the

base of both tarsomeres 4 - 5.

The distribution of these categories in Thailand appears to be random,

however, the sample size of this study was much too small for accurate mea-

surement of trends. Overall, there are indications that the hindtarsal banding

on nigerrimus exhibits clinal tendencies between Sri Lanka-India and Malaysia.

Studies on this variation in the Indonesian area would be most helpful.

A few nigerrimus with pale basal bands on both tarsomeres 4 - 5 hada

pattern approaching that found on Thailand peditaeniatus; however, they can be

separated quickly by the remigium, humeral crossvein, base of costa and vein

R-R, characters. :

Another variable noted on Thailand nigerrimus is the extent of dark scales on

vein 1A. Several specimens were noted with the apical half of this vein dark

scaled. The holotype from Calcutta has a similar vein 1A. This variation was

also seen on 2 specimens from the above mentioned Rangoon, Burma collection

(P. F. Mattingly, 11-1963; “°192B, 2131C), as previously noted by Reid (1968).

This is a variable that occurs in this group on 3 species, avgyropus, nigerrimus

and peditaeniatus, and is observed most frequently on avgyropus.

70 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

The presence or absence of a pale fringe spot of vein Cug is another variable

character. Reid (1953) noted 68% of the females he examined from Malaya had

this pale spot. In Thailand the frequency of this spot was lower, it occurred on

only 53% (23/44) of the females examined. Only 2% (1/44) of females examined

from India had this spot.

Scattered pale scales were nearly always present on the base of the costa;

however, several specimens were noted without such scales, or with only 1 - 2

pale scales.

BIOLOGY. This is a lowland, valley species which seems to prefer deep

cool water for the immature stages. Scanlon and Esah (1965) found nigerrimus

in the agricultural area of the Chiang Mai Valley, near and in the city, but not

at various elevations on the nearby forested mountains. In Thailand immatures

have been collected from: city moat (canal), large open marshes, large stream

pools and a rice field. Immature specimens from Malaysia were collected in

large swamps, ground pools, a small pond and rice field. In general, these

immature collections were made in semi-open large bodies of water with some

emergent or floating vegetation, under open sunlight to moderate shade and up to

300 m elevation. This should not be interpreted as implying any rice field. The

authors feel this species is probably found in those fields where the water depth

is sufficient to keep the water temperature fairly cool. Adults have been collec-

ted in Thailand resting in cattle sheds and houses, biting cattle, horsesand man,

in light traps and in net traps baited with dry ice. In Malaysia adult females

have been collected biting man outside and inside houses and found resting in

houses at night (Moorhouse 1965). Although nigerrimus does feed on man, its

degree of attraction to various hosts is not understood and needs more investi-

gation. Broad spectrum studies like Bruce-Chwatt et al. (1966) suggest niger-

vimus is primarily zoophilic, however, more concentrated efforts are needed

on such suspect species in malarious areas. Studies on host range and feeding

preferences like that of Sullivan et al. (1971) on Aedes albopictus (Skuse), are

urgently needed for comprehension of anopheline feeding behavior in malarious

areas.

MEDICAL SIGNIFICANCE. Hodgkin (1956) reported nigerrimus as a vector

of malaria parasites in Malaya. However, more recent studies by Warren et al.

(1963), Wharton et al. (1964) and Moorhouse (1965) were unable to clarify this

report because of limited specimens. There are also reports of malaria para-

site transmission by nigerrimus in Indonesia, but these records require caution

in interpretation due to possible confusion with other members of the species

eroup. Reid (1968) considers nigerrimus a malarial pathogen vector of low

status.

The capabilities of nigerrimus as a vector of filarial parasites also needs

further study. There are a number of early reports of human filarial larvae

found in "nigerrimus". Iyengar (1953) reported an infection rate of 3. 7% for

hyrcanus var. nigerrimus in investigations of Brugia malayi in southern Thai-

land, but Harinasuta et al. (1964, 1970) were unable to find infected members

of this group in Thailand. Many of these early reports probably refer to ani-

mal filariae such as Divofilaria spp. and Setaria spp. (Reid et al. 1962, Reid

1968). Experimentally, nigerrimus has shown a low susceptibility to infection

with periodic Brugia malayi (Reid et al. 1962).

Harrison and Scanlon: Subgenus Anopheles in Thailand 71

ANOPHELES (ANOPHELES) PEDITAENIA TUS (LEICESTER)

(Figures 8, 28, 29, 30)

Myzorhynchus peditaeniatus Leicester 1908: 31 (%, 9, L).

Myzorhynchus sinensis var. peditaeniatus of James and Stanton 1912:

61.

Anopheles (Anopheles) hyrcanus var. nigerrimus of Edwards 1932: 41 (in part);

Baisas and Hu 1936: 221 (P*, L*, E*).

Anopheles (An.) peditaeniatus Leicester, Reid 1953: 32 (“*, 9*, L*, P*, E*).

Anopheles (An.) peditaeniatus (Leicester), Stone, Knight and Starcke 1959:

25: Reid 1968; 109 (¢*, @*, L*, P*, E*).

All stages of this species are relatively easy to identify. In general the

adults have broad pale bands on the fore-and hindlegs, the remigium is pale

scaled, humeral crossvein without scales, a long basal dark mark on Cug and

vein R-R, with numerous pale scales. In addition to the above the male lacks

a pale basal band on palpal segment 3. The pupa is distinct on the basis of the

long refractile margin on the paddle and seta 9-VIII spine-like, with reduced

branches. The larva is easily separated by seta 4-M having sinuous branches.

Like sinensis except:

FEMALE (Figs. 8, 28) Head. Palpus with 4 distinct pale bands, infre-

quently apical and subapical pale bands confluent, usually with mesal patch of

pale scales on segment 2 and sometimes 3. Thorax. Scutal integument gray-

brown, with darker median and submedian lines, eyespots poorly defined; pleur-

al setae: 2 - 6 propleural, 2 - 4 spiracular, 6 - 11 prealar, 2 - 5 upper and

4 - 6 lower sternopleural and 4 - 9 upper mesepimeral. Wing. Pale scales

may have white, yellow or dirty pale appearance; costa dark scaled except

small subcostal and preapical pale spots; remigium mostly pale scaled with

anterior margin always pale, pale scales infrequently glossy white; humeral

crossvein bare, rarely with 1 - 2 scales; basal 0.33 of R with numerous scattered

pale scales, sector and subcostal pale spots frequently confluent, R, nearly

always with numerous pale scales between subcostal and preapical pale spots,

rarely sector and subcostal pale spots incomplete or missing; tip of vein Ry dark

scaled; Ro-Ro,3 with mixed dark and pale, or mostly pale scales to fork; Ro

dark scaled except preapical pale spot; Rg with pale middle section; R445 most-

ly pale scaled, with proximal and distal dark marks; vein M with base mostly

dark or dirty pale scaled, with pale area preceeding fork; vein Cu with long

basal dark mark, approaching within its own length or less of upper dark mark

on 1A; vein 1A with upper and apical dark marks infrequently confluent; apical

dark mark on 1A longer than that on vein Cup; apical pale fringe spot long, start-

ing at or slightly above vein R, and extending to vein R45; pale fringe spot at

Cug rare. Legs. Coxae with pale scales, upper midcoxa with 3 - 5 setae.

Foreleg: Femur dark scaled; tibia dark dorsally, pale ventrally; tarsomeres

1 - 3 with broad apical pale bands and tarsomere 1 pale ventrally; pale bands

on tarsomeres 2 and 3 may approach 0.5 or more tarsomere length; tarsomere

4-5 dark scaled. Midleg: Femur and tibia dark dorsally, pale ventrally, both

may have few dorsal pale scales at apices; tarsomere 1 usually with few ventro-

basal pale scales; tarsomeres 1 - 3 with fairly broad apical pale bands, that on

tarsomere 3 may approach 0.5 tarsomere length; tarsomeres 4 - 5 dark scaled.

Hindleg: Femur and tibia dark dorsally, pale ventrally, tibia with distinct pale

apical band; tarsomeres 1 - 4 with apical pale bands; tarsomere 4 andusually 95

with basal pale bands (see taxonomic discussion).

MALE (Fig. 28) Head. Palpus usually with mesal line of pale scales on

72 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

segments 2 - 3, no basal pale band on segment 3, segments 4 - 5 with pale

patches and bands. Genitalia. Basimeres with numerous pale scales; clas-

pette with 2, infrequently 3 long setae on ventral lobe, mesal seta distinctly

longer than lateral seta; dorsal lobe of claspette with fused club on inner half

formed from 2 - 3 stems, outer half of lobe with 2 flattened leaflet-like spines

adjacent to fused club; aedeagus with 4 - 7 pairs of leaflets, largest with large

apical-subapical teeth and small basal teeth.

PUPA (Fig. 29). Integument generally with light to moderate pigmentation.

Cephalothorax. Wing case with faint to moderate pattern of bars, lines or

squares, no dark spots; antennal case with moderate pigmentation at joints,

but no prominent dark spots. Tvumpet. Moderately pigmented, with saw-

tooth areas on rim, these areas rarely difficult to see. Abdomen. Seta 9

lightly pigmented on all segments, without, or with very fine branches on seg-

ment VIII; terga III - VII with or without dark central spot; setae in general

with few branches; seta 0 on II - VIII well developed and branched; seta 1 with

few branches on V - VII, usually 2 - 3; 5-V with 14 - 28 branches; male genital

lobe without dark pigmentation on distal half. Paddle. Base without dark pig-

mentation, area of refractile border and marginal teeth long, approximately

80% of paddle length; 1-P usually simple.

LARVA (Fig. 30) Head. Tanto dark brown; seta 3-C with 40 - 70 branches;

0, 6-C with 13 - 18 and 11 - 20 branches respectively; 8, 9-C with 4 - 9 and

3 - 7 branches respectively. Thorax. 1-P simple or with 2 - 3 distal branches;

11-P with 3 - 7 stout branches from base; 14-P with 3 - 7 branches, usually

4 - 5; 4-M small, with 4 - 10 sinuous branches radiating from base; 3-T with

long, flattened, unpigmented leaflets. Abdomen. Seta 1 with flattened unpig-

mented leaflets on I - II, well developed (palmate) and darkly pigmented leaflets

on III - VII, pigment frequently extending beyond poorly developed shoulders

onto filaments; 6-I, III with 18 - 25 and 16 - 22 branches respectively; 0 on

Il - VII with relatively long sinuous branches; 5 on III - V with 6 - 14, 3 - 5 and

4 - 7 branches respectively; pecten plate with 7 - 9 long teeth; 1-X approximate-

ly equal length of saddle.

TYPE-DATA. Three female syntypes are in the BMNH collection. These

specimens were supposedly selected by James and Stanton (1912) from the col-

lection of the Institute for Medical Research, Kuala Lumpur, after Leicester's

death. According to James and Stanton (1912: 59) Leicester's type-specimens

were lost enroute to the BMNH, indicating a type had been selected by Leicester

for peditaeniatus. However, Leicester (1908: 33) does not mention selecting a

type-specimen, but says, ''Described from a large series bred from larvae."

The specimens in this series are considered syntypes here, and include those

selected and sent to the BMNH by James and Stanton. All 3 syntypes in the

BMNH have an alphabetical code that suggests a rearing designation. The 1st

syntype has the following label data: (1st label) - ''D''; (2nd label) - ''Cotype"’

on circular BMNH label with "''Myzorhynchus peditaeniatus Leic.'' on the under-

side; and (3rd label) - ''Fed. Malay States, Dr. G. F. Leicester, 1912-350."

This specimen is in poor condition. The 2nd syntype is also in poor condition

and has the following data: (1st label) on underside of cardboard minutin stage,

'1 mile lake, Batu Rd, 26/4/04"; (2nd label) - ''E''; (3rd label) - ''Cotype" on

circular BMNH label with "Myzorhynchus peditaeniatus Leic."' on the under-

side; and (4th label) - "Kuala Lumpur, Fed. Malay States, Dr. G. F. Lei-

cester, 1912-350.'' The 3rd syntype has its legs and abdomen glued flat toa

cardboard stage and is in excellent condition. This specimen has the follow-

ing label data: (1st label) - on underside of minutin stage, ''Pupafr., Jail Pool,

Kuala Lumpur, 16/4/03"; (2nd label) - 'F"''; (3rd label) - ''Cotype" on circular

Harrison and Scanlon: Subgenus Anopheles in Thailand 73

BMNH label with "Myzorhynchus peditaeniatus Leic."' on the underside; and

(4th label) - ''Kuala Lumpur, Fed. Malay States, Dr. G. F. Leicester, 1912-

350.'' This last specimen fits the concept of peditaeniatus in almost every

respect. Two males were also selected and sent to the BMNH by James and

Stanton to represent peditaeniatus, however, both are actually nitidus as

pointed out by Reid (1953, as indiensis).

DISTRIBUTION (Fig. 29). This species was collected from most areas in

Thailand, except those heavily forested or at high elevations. It is a predom-

inant species in agricultural areas, particularly those with rice fields, and will

be found as abundant as sinensis, or more sO, in many areas. Specimens

(280°, 1389, 49 larvae and 100 larval and pupal skins) in the USNM were exam-

ined from the following provinces of THAILAND: Ayutthaya, Chanthaburi,

Chiang Mai, Chiang Rai, Chon Buri, Krabi, Lampang, Nakhon Si Thammarat,

Nan, Narathiwat, Nonthaburi, Pathum Thani, Phra Nakhon, Prachin Buri,

Prae, Rayong, Surat Thani, Trang, Udon Thani and Yala. During the period

1968-70 additional specimens were examined in Thailand from Chumphon, Lop

Buri, Mae Hong Son, Nakhon Ratchasima and Sara Buri. These records

coupled with those listed by Scanlon et al. (1968) indicate a distribution through-

out most of Thailand. Consequently, the distribution map (Fig. 29) is shown

as such, although certain forested or mountainous areas may be free of this

mosquito. Specimens (169°, 4922, 30 larvae and 38 larval and pupal skins) in

the USNM from outside Thailand were examined: BURMA, CAMBODIA, CHINA

(Fukien, Kweichow and Yunnan), INDIA (Assam, Bengal, Bihar and Punjab),

INDONESIA (Java and Sumatra), MALAYSIA (Peninsular Malaysia), NEPAL,

PHILIPPINES, SOUTH VIETNAM, and SRI LANKA. Beside the 3 syntypes,

numerous male and female specimens were examined in the BMNH from:

BURMA, INDIA (Bihar, Central Provinces, Mysore), INDONESIA (Sulawesi,

Sumatra), MALAYSIA, NEPAL, PHILIPPINES, SRI LANKA and THAILAND

(Chiang Mai, Phra Nakhon). An additional 2% and 172 specimens from CAM-

BODIA were examined in the ORSTOM collections. Additional literature

records refer to BORNEO and the Lesser Sunda Islands in INDONESIA. There

are several specimens in the USNM from Fukien, Kweichow and Yunnan

provinces, China, which fit the usual description of peditaeniatus, except the

hindlegs have only narrow pale bands. However, since this variation is known

for peditaeniatus from Assam and other parts of India (see taxonomic discus-

sion) an extension up into southern China is logical. Feng (1964) in his work

on the cryptic species called "sinensis" in Kweichow Province based his divi-

sions on egg structures. He described the adult of his ''medium decked egg

type’' and this description fits the specimens in the USMN except for one detail.

Feng notes the hindlegs have only narrow apical pale bands on tarsomeres 1 -

4; however, 2 females from Fukien and one from Kweichow also have a very

small patch of pale scales on the base of tarsomere 4. The one female from

Yunnan Province has the hind tarsomeres like Feng's description. One male

from Fukien was available for examination, and fits the current concept of

peditaeniatus. Unfortunately, the abdomen was missing and the genitalia could

not be checked. Considering the southern Chinese distribution of peditaeniatus,

the species recorded as hyrcanus var. nigerrimus from Hainan Island by Ho

(1938b) probably refers to this species (see nigerrimus distribution). No speci-

mens resembling peditaeniatus have been seen or reported from the Hong Kong

area, or from Taiwan. Overall, peditaeniatus probably has the widest distri-

bution of any Southeast Asian member of the group, particularly since it is now

recognized from southern China. Anopheles sinensis was previously considered

the most widespread species, but it does not occur west of Assam, in the Philip-

74 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

pines, or east of Sumatra in Indonesia. The identity of Palaearctic sinensis

is still unclear.

TAXONOMIC DISCUSSION. Nearly all the adults examined from Thailand

were readily identifiable on the basis of the wing characters and the width of

the hind tarsal banding. Specimens with a pale scaled remigium and bare hu-

meral crossvein associated with a long basal dark mark on vein Cu and broad

hindtarsal bands, should be recognized immediately as peditaeniatus. The

use of the hindtarsal bands as the primary indicator of peditaeniatus should be

discontinued, for the size of these bands is highly variable throughout its

range. In India and Sri Lanka the most frequently encountered phenotype has

only a small basal pale band or patch on tarsomere 4, beside the narrow api-

cal pale bands. Less frequently this basal band or patch of pale scales dis-

appears, particularly in northern India (Assam) and Nepal, leaving only nar-

row apical pale bands on hindtarsomeres 1 - 4. Both of these phenotypes are

also present in southern China as discussed above under "distribution." In

the Philippines, at the other end of its distribution, the most frequently en-

countered phenotype of peditaeniatus has 2 broad pale bands on the hindtarsi,

which involve basal pale bands or patches on both tarsomeres 4 - 5. This

latter phenotype is the one most commonly encountered in Indonesia, Malaysia

and most of Thailand. In the northern parts of Thailand and South Vietnam

the phenotype with pale scales on the base of only tarsomere 4 becomes more

abundant. In the Chiang Mai area of Thailand this phenotype accounts for at

least 50% of the specimens. Occasional specimens are found which have ex-

tremely wide pale bands on the hindtarsi. In these, the dark band on tarso-

mere 4 may be so narrow, that it approximates the banding found on the hind-

tarsi of argyropus. Fortunately, wing characters and midtarsal banding easi-

ly separate these species. The remigium on peditaeniatus is often entirely

pale scaled, particularly in India and Sri Lanka where it may have a silvery-

white appearance (Reid 1953); however, frequent specimens were encountered

from Thailand and Malaysia which retained pale scales on the anterior mar-

gin, but had dark scales on the distal half of the hind margin. Another variable

wing character rarely encountered, is the presence of a pale fringe spot on

vein Cug. Occasional specimens of peditaeniatus are encountered with the

distal half of vein 1A dark scaled, or nearly so.

The immature stages are as easy to distinguish as the adults. The pupa is

readily identifiable by the long refractile margin on the paddle, the spine-like

seta 9-VIII and the thickened saw-tooth like areas on the trumpet rim. The

first 2 characters are shared with argyropus, which has vertical wrinkles on

the outside wall of the trumpet pinna instead of the thickened saw-toothed areas.

Infrequent specimens were noted where the saw-tooth like areas were not as

obvious as usual, but high magnification confirmed their presence. Even with-

out these 3 characters the pupa can be associated with sinensis and the lesteri

subgroup on the basis of the number of branches of seta 1 and 5 on most of

the segments. The only exception to this is the low number of branches found

on these setae on pseudosinensis (nigerrimus subgroup) from the Philippines.

The similarity of the long refractile border on the paddle and the spine-like

seta 9-VIII on pupae of argyropus and peditaeniatus is apparently superficial,

for other characters on the pupa are quite different (e.g., number of branches

on abdominal setae 1 and 5). These latter characters plus adult and larval

characters have been used to show that argyropus and peditaeniatus are not

closely related (Harrison 1972). The larva is also distinct, and easily

separated on the basis of the sinuous branches on seta 4-M and the low number

_ of branches on setae 8, 9-C. Some specimens of argyropus and nitidus may

Harrison and Scanlon: Subgenus Anopheles in Thailand 75

have slightly sinuous branches on seta 4-M, but these species have numerous

branches on seta 8-C. The examination of specimens with reared, associ-

ated larval and pupal skins from Malaysia, Philippines, Sri Lanka and Thai-

land, including many from the Chiang Mai area of northern Thailand, revealed

peditaeniatus is a highly variable species.

BIOLOGY. The largest number of immature collections of peditaeniatus

in Thailand have probably come from rice fields. The larvae, however, are

not restricted to this habitat, for they have been collected in the following

additional sites: open and shaded ditches, marshes, seepage areas, ponds,

large open ''Swamps,"' temporary puddles, flood pools, stream margin,

stream pools, animal footprints, shallow well, holes in logs on a river anda

250 1 clay water jar. Apparently, the most favorable larval requirements

involve shallow water, warm water temperature, light or no shade and some

vegetation, particularly emergent grasses. Immatures have been collected

up to 540 m elevation in Thailand, while adults have been collected at 570 m

in Sri Lanka. In Thailand adults have been collected in light traps, in Shannon

traps baited with light and humans, in net traps utilizing dry ice, resting on

trees, resting in animal sheds, biting humans outside and inside homes and

biting cattle and buffalo. Very few adults were taken resting in houses in 3

years of collecting in Bangkok, but much larger numbers were taken in animal

shelters and in light traps. Reid (1961) determined that Malayan peditaeniatus

preferred a calf over 2 men by a 11:1 ratio. In similar studies in Chiang Mai

and Bangkok, Thailand, the overwhelming proportion of peditaeniatus fed on

cattle.

MEDICAL SIGNIFICANCE. The zoophilic habits of peditaeniatus greatly

reduce the probability of its being a vector of human disease pathogens. Reid

et al. (1962) and Wharton, Laing and Cheong (1963) found peditaeniatus to be

a good experimental vector of periodic Brugia malayi, but the former concluded

the proportion biting man in nature was too small to pose a problem. Harina-

suta et al. (1964, 1970) did not find this species infected with filariae during

studies in endemic B. malayi and Wuchereria bancrofti (Cobbold) areas in

Thailand. Warren et al. (1963) determined experimentally that peditaeniatus

has a very low susceptibility to Plasmodium cynomolgi bastianellii. Wharton

et al. (1964) found no evidence of parasites in 146 wild caught peditaeniatus

dissected during malaria studies in Malaya. It is possible that this species is

involved in the transmission of animal parasites, particularly cervid filariae

of the genus Sefaria (Reid et al. 1962, Reid, 1968). If these parasites are

serious pathogens in cattle, then peditaeniatus should probably be considered

an economic pest in Thailand.

ANOPHELES (ANOPHELES) PURSATI LAVERAN

(Figures 8, 31, 32, 33)

Anopheles pursati Laveran 1902: 907 (2).

Anopheles hyrcanus var. nigerrimus of Reid 1947: 89.

Anopheles (An.) species Do near nigerrimus of Reid 1953: 49.

Anopheles (An.) pursati Laveran, Reid 1963: 98 (““*, @*, P*, L*, E*); Reid

1968. 96:(o"%, 9% P*, CLR)

A small species with narrow hindtarsal bands, a short apical fringe spot

on the wing, pale scaled remigium, dark scale patch on the humeral cross-

vein, short basal dark mark on vein Cu and pale scales on the midcoxa.

76 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Beside these characters the male possesses a pale basal band on palpal segment

3. The immature stages are very similar to nigerrimus, but can be separated

by the characters presented here. As for sinensis except:

FEMALE (Figs. 8, 31) Head. Palpus with 4 distinct pale bands, apical

band very small and on tip of last segment, preapical dark band wide and dis-

tinct. Thorax. Integument gray-brown with eye spots poorly defined; pleural

setae; 2 - 5 propleural, 2 - 4 spiracular, 6 - 9 prealar, 2 - 3 upper and 3 - 4

lower sternopleural and 2 - 4 upper mesepimeral. Wing. Blurred color pat-

tern; costa usually dark scaled except small subcostal and preapical pale spots,

infrequently with small pale humeral spot; remigium wholly pale, or with an-

terior pale and posterior dark scaled; humeral crossvein with patch of dark

scales; vein R dark scaled at extreme base, usually with few pale scales or

distinct pale spot at presector area, then dark scales followed by distinct sec-

tor pale spot; subcostal pale spot on R usually extending basally for some dis-

tance towards sector spot; vein Rj rarely with pale scales between subcostal

and preapical pale spots, tip dark scaled; Ro- R913 dark scaled basally, pale

before fork; Ro and Rg dark scaled except pale continuation from preapical

pale spot on costa; R ‘ pale centrally, may have few scattered dark scales

in pale area; vein M oe basally, pale prior to fork; vein Cu with pale scales

at extreme base, that pale area as long as or longer than following basal dark

mark; basal dark mark on Cu usually short, separated by more than its own

length from upper dark mark on vein 1A, infrequently longer and approaching

within its own length of upper dark mark on 1A; vein 1A with upper and apical

dark marks approximately equal in length; apical dark mark on 1A approximate-

ly equal that on vein Cug; apical pale fringe spot short, extending at most from

Rj to Rg, often smaller, from Rg to Rg; posterior margin of wing dark scaled,

rarely with fringe spot at Cuy. Legs. Coxae with pale scales, upper midcoxa

with 1 - 2 setae. Foreleg: Femur dark scaled; tibia dark scaled dorsally,

pale ventrally, may have small pale spot at apex; tarsomere 1 - 3 with narrow

apical pale bands, tarsomere 1 pale ventrally; tarsomeres 4 - 5 dark scaled.

Midleg: Femur dark scaled; tibia dark scaled except small basal and apical

pale band or patch; tarsomeres 1 - 3 dark scaled with narrow apical pale bands;

tarsomeres 4 - 5 dark scaled. Hindleg: Femur dark scaled; tibia dark scaled

except small basal and apical pale band or patch; tarsomeres 1 - 4 dark scaled

except narrow apical pale bands; tarsomere 5 dark scaled. Abdomen. Posteri-

or angles of tergum VIII without scales.

MALE (Fig. 31) Head. Palpal segment 3 with distinct pale basal band, seg-

ments 2 - 3 may have mesal line of pale scales, segments 4 - 5 with pale bands

and patches. Wing. Costa nearly always with small pale humeral spot; infre-

quently with pale fringe spot at Cug. Genitalia. Basimeres with numerous

pale scales; claspette with 2 setae on ventral lobe, mesal seta much longer than

lateral seta; dorsal lobe of claspette with fused club on inner half formed from 2

stems, outer half of lobe with 2 flattened leaflet-like spines adjacent to club;

aedeagus with 2 - 3 pairs of leaflets, largest with small subapical denticle.

PUPA (Fig. 32). Integument generally lightly pigmented. Cephalothorax.

Wing case usually with light tan pattern of squares and lines, may also have

dark spots as on sinensis pupa; antennal case with or without pigment at joints.

Trumpet. Lightly pigmented, without thickened saw-tooth areas on rim. Ab-

domen. Seta 9 lightly pigmented on all segments, 9-VIII with numerous bran-

ches; areas surrounding alveoli of setae pigmented, particularly seta 2; terga

TI - VII rarely with central dark spot; setae 1 and 5 with numerous branches

on segments III - VI; 5-V with 45 or more branches; 1-VII with 3 - 13 branches,

rarely less than 6; 6-VII cephalad of seta 9-VII; male genital lobe may have light

Harrison and Scanlon: Subgenus Anopheles in Thailand 77

pigment on distal half. Paddle. Base may be pigmented; refractile border

approximately 65 - 75% of paddle length; 1-P with 3 or more distal branches.

LARVA (Fig. 33). Generally light tan; 3-C with 45 or more branches; 5,

6-C with 11-16 and 12 - 18 branches respectively; 8, 9-C with 12 - 21 and 8 -

17 branches respectively. Thovax. 1-P usually simple, infrequently 2 - 3

distal branches; 11-P with 4 - 8 stout branches from base; 14-P with 3 - 5

branches, usually 3 - 4; 1-M with 26 - 38 branches, usually 28 or more; 4-M

with 5 - 9 erect, stiff branches from central stem; 13-M with 6 - 9 branches;

3-T with flattened, unpigmented leaflets. Abdomen. Seta 1 with flattened,

unpigmented leaflets on segments I - II, well developed (palmate) and pig-

mented leaflets on III - VII, pigment may extend onto filaments; 6-I, III with

11 - 20 and 12 - 17 branches respectively; 5-IV with 2 - 5 branches; posterior

tergal plates conspicuously large on some specimens; pecten plate with 5 - 7

long teeth; seta 1-X slender, longer than saddle.

TYPE-DATA. The type-specimens, 2 females mounted in balsam ona

microscope slide, are in the Pasteur Institute, Paris, where they were found

by Reid (1947) after having been lost for over 40 years. The slide bearing

the types contains the notation ''A. pensati, Montagnes De Pensat, Cambodge,

Feévier 1902"; however, the original description clearly gives the locality as

Pursat, or at least a mountainous wooded area west of that city. Reid (1953,

1963) redescribed pursati in detail and deposited specimens with skins from

Malaysia in the BMNH. Based on an examination of the 2 pursati types we

agree with Reid (1963) that his species Dg near nigerrimus is equal to pursaiz.

DISTRIBUTION (Fig. 32). Specimens were taken from several widely

scattered points in Thailand, conveying the impression that this species is

widely distributed in the country, but uncommon. The majority of specimens

examined came from the central valley, around and north of Bangkok. Reid

(1963) reported specimens from the Thurman light trap collections in Chiang

Mai, and 4 such specimens were located in the USNM. Specimens (14%, 2592 and

34 larval and pupal skins) in the USNM were examined from the following pro-

vinces of THAILAND: Ayutthaya, Ang Thong, Chiang Mai, Nonthaburi, Pathum

Thani, Phet Buri, Phra Nakhon, Surat Thani, Thon Buri and Udon Thani. Ad-

ditional specimens were examined in Thailand during 1968-70 from Sara Buri

Province. From elsewhere, 1°, 2¢ and 6 larval and pupal skins in the USNM

were examined from MALAYSIA (Peninsular Malaysia) and SOUTH VIETNAM.

The last record is based on a female taken by light trap in 1967, from My Tho,

in the delta province of Dinh Tuong. An additional 332 from CAMBODIA were

examined in the ORSTOM collections. Reid's (1968) Malayan records and

specimens are all from northern provinces. Apparently, pursati, is one of

those species that is widely distributed in the monsoon areas of Indochina and

Thailand, but does not extend further south than northern Peninsular Malaysia.

Much additional collecting is needed for a better understanding of this species.

TAXONOMIC DISCUSSION. The small size of this species should always

be considered, for only rare stunted members of the other species are found

as smallas pursati. In addition, the patch of dark scales on the humeral

crossvein, narrow apical wing fringe spot, narrow hindtarsal bands, usually

pale remigium and pale scales on the midcoxa make the adults of this species

easily separable from the other species. The males are very distinct, for in

addition to the above, they also have a pale band at the base of the 3rd palpal

segment and a pale humeral spot of the costa. Some variation in adult wing

characters was observed. The preapical pale spot on the costa is infrequently

extremely small or even absent, a variation also noted on sinensis. The length

of the basal dark mark on vein Cu is quite variable and consequently, should

78 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

not be used in Thailand for separating pursati from lesteri paraliae. Approxi-

mately 42% (10/24) of the adult females examined have this mark approaching

within less than its own length of the upper dark mark on vein 1A. The Thai

specimens differ from the Malayan specimens (Reid 1963, 1968) in this regard.

This variation is apparently widespread in Thailand, and may be the cause for

a number of pursati from the area around and north of Bangkok, having been

previously misidentified as lesteri parailiae.

The pupa of pursati is very similar to those of nitidus and nigerrimus. All

have the thin uniform trumpet rim and numerous branches on abdominal setae

1 and 5. In general nztidus has the least skin pigment, with pursati next fol-

lowed by the rather darkly pigmented skin of nigerrimus. The numerous bran-

ches of seta 1-VII on pursati is usually diagnostic, and is somewhat like pupae

in the barbirostris group. Persons using the position of seta 6-VII on either

nigerrvimus or pursati, must be certain that the skins are mounted as flat as

possible; otherwise, the positions can be very distorted and misleading. Oc-

casionally, pupal skins without associated larval skins or adults will be en-

countered which cannot be identified with certainty.

The larva is also very similar to those of nitidus and nigerrimus, but is

usually distinct on the basis of setae 14-P and 1-M. Some pursati larvae have

a larger posterior tergal plate on most abdominal segments than do other mem-

bers of the complex, but additional specimens are needed to confirm this trend.

Unfortunately, no whole larvae were available for examination during the study

and the larval plate was necessarily drawn from skins.

BIOLOGY. Most adults examined were collected by light traps; however,

a few females were taken biting man, cattle or buffalos in the area around and

north of Bangkok. Reid (1963) also reported pursati biting man in Malaya.

Specimens examined in Thailand during 1968-70 were collected resting in a

clay water jar, biting cattle and man and 16 females were collected one night

during a 2 hour period in a net trap baited with dry ice. Reid (1968) gives the

larval habitat in Malaysia as deep cool pools, with Pistia spp. floating on the

surface. The majority of specimens from Thailand were collected in the cen-

tral valley which is essentially a rice plain, but also has many man made can-

als and ponds. Some varieties of rice grown in this area require 90 - 120 cm of

water for a period of time. Such deep water habitats have been badly neglected

in larval collections in Thailand and deserve more study. Immatures of pursati

have been collected from: rice fields, ponds, ditches and a large ground pool.

These habitats had abundant emergent and/or floating vegetation.

MEDICAL SIGNIFICANCE. This species is collected so infrequently that

nothing is known about its adult behavior. Anopheles pursati probably is not a

vector of human pathogens in Thailand.

ANOPHELES BARBIROSTRIS SPECIES GROUP

(An. barbirostris species group, Reid 1962)

Anopheles barbirostris Van der Wulp 1884.

GENERAL. Large dark mosquitoes. Head. Palpus dark with dense erect

scales (see male description below); palpus approximately equal to length of

proboscis; clypeus without scales; pedicel with dark and/or pale scales on up-

per and outer aspects; flagellomere 1 with dense patch of dark and pale scales.

Thorax. Pleural area with white scale patches; lower mesepimeral setae fre-

quently present. Wing. Basal half of costa with humeral pale spot and few

Harrison and Scanlon: Subgenus Anopheles in Thailand 19

scattered pale scales; subcostal pale spot small; remigium and base of vein R

with pale scales; veins Rg-R9,3 and M dark scaled to forks; tertiary fringe

scales no paler than other fringe scales; with 1 - 2 apical pale fringe spots,

frequently with pale fringe spot on posterior margin. Legs. Coxae with pale

scales; femur, tibia and tarsus mostly dark, usually not speckled with pale

scales; hindtarsus with narrow pale rings on some tarsomeres. Abdomen.

Sterna usually with at least a few scattered pale scales.

FEMALE Head. Palpus completely dark scaled. Abdomen. Sternum VII

with distinct tuft of dark scales.

MALE Head, Palpus often with several pale scales on apex of segments

4-5. Abdomen. Without scale tuft on sternum VII; dorsal surface of 8th

segment with some pale scales. Genitalia. Basimeres with scales; aedeagus

with 3 - 6 pairs of leaflets; 1st leaflet usually with distinct basal tooth; larger

leaflets with fine teeth along both edges, approximately 0.5 as long as aedeagus;

claspette faintly bilobed; ventral lobe normally with 2 large spines arising from

membraneous area, mesal spine longer than lateral spine; dorsal lobe with

club-like structure composed of 4 - 5 separate basal stems that fuse distally.

PUPA Cephalothorax. Palpal case sexually dimorphic in most species, but

not as distinct as in hyrcanus group; male palpal case with attenuated tip, that

of female may be slightly attenuate, but not as long as male. Tvumpet. Long-

est axis transverse to stem; rim without or with secondary cleft, or thickened

ridge replacing secondary cleft, but without tragus. Abdomen. Terga without

denticles on caudal margin; setae 1, 5 on IV - VII with many branches, central

branch usually longer than other branches; seta 1-VII with more than 13 bran-

ches (except Philippine species).

LARVA Head. Seta 3-C usually with many stiff broom-like branches from

base; seta 1-A large with numerous branches, usually reaching or exceeding

tip of antenna. Thorax. Seta 1-P with 4 or more branches from near base.

Abdomen. Seta 1 palmate onI - VII, well developed only on III - VI; 1-II usu-

ally pigmented (except vanus subgroup).

DISTRIBUTION. This group consists of 11 species and is limited to the

Oriental region, except for vanus Walker, which has been recorded from the

western tip of New Guinea (Reid 1968). Reid (1962) divided the complex into 2

subgroups on the basis of adult, larval and egg characters. These subgroups

are:

- barbirostris subgroup - - vanus subgroup -

barbirostris franciscoi ahomi manalangi

campestris hodgkini barbumbrosus vanus

donaldi pollicaris reidi

Members of both subgroups occur throughout nearly the entire distribution of

the group. Distributions for most species are uncertain because of identifica-

tion difficulties, particularly in Indonesia and the Indian subregion. The north-

ernmost records in the west occur in Pakistan and Nepal, while in the east,

members have been collected in Kwangtung, Szechwan and Yunnan provinces of

China, and Hainan and Taiwan islands (Chow 1949). The center of distribution

for the group appears to be Malaysia, but this may be a reflection of the large

amount of work done by Reid (1962, 1968) in that country. At least 6 species,

i.e., barbirostris, barbumbrosus, campestris, donaldi, hodgkini and pollicaris,

80 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

occur in Malaysia. To the east, franciscoi Reid, manalangi Mendoza and vanus

Walker are recorded from the Philippines, and only barbirostris, barbumbro-

sus and vanus are recorded from the eastern Indonesian islands. The 6 species

recognized from Malaysia are also found in Thailand, but donaldi and pollicaris

are found only in extreme southern Thailand. Only 6 species are recorded north-

east or west of a line drawn through the Isthmus of Kra on peninsular Thailand.

Of these, hodgkini occurs up into central Thailand and probably also occurs in

Burma and Cambodia for numerous specimens have been found in Thailand pro-

vinces immediately adjacent to these countries. Anopheles ahomi Chowdhury and

veidi Harrison, (Harrison 1973b) are apparently confined to the Indian subregion.

Anopheles barbirostris, barbumbrosus and campestris are the remaining 3

species on the mainland, and the first 2 are widely distributed from India to

southeastern China and nearby islands. The distribution of the last species,

campestris, is uncertain. Numerous adults examined from India and Sri Lanka

had adult characters as described for campestris, but when available associated

immature skins were examined they were clearly barbirostris. Such disjunct

associations also occur in northern Thailand, where campestris apparently

does not occur. Confirmed records of campestris are currently known only

from Peninsular Malaysia and Thailand. Workers studying man biting "bar-

birostris"' in India or Southeast Asia should definitely try to rear specimens

so the distribution of campestris, a confirmed vector of malaria parasites in

Malaysia, can be proven or disproven in other areas.

TAXONOMIC DISCUSSION. In Thailand, assignment of specimens to the

barbirostris species group, particularly the females and larvae is quite simple

using the characters provided in the keys. However, as noted in the keys and

discussion, the determination of a particular specimen to a given species in

the group may be very difficult, particularly if the specimen is not accom-

panied by additional reared life stages.

As in the hyrcanus group, recognition of this species group was facilitated

by the accumulation of a mass of biological and disease transmission data by

workers in Malaya which led Reid (1962) to search for morphological differ-

ences. The differences he detected are even finer than those used to separate

members of the hyrcanus species group. Furthermore, these meticulous

differences persist for the most part in areas outside of Malaysia and reinforce

Reid's (1962, 1968) position that these represent valid species. The numerous

specimens examined from Thailand have widened the ranges of variation for a

number of characters, and in rare cases eliminated the value of a given char-

acter, but the species parameters established by Reid remain stable.

The barbirostris species group is most closely related to the bancrofti

species group, which has its center of distribution in the Australian region.

Both groups exhibit: wings with areas of mixed black and white scales; pale

fringe spots on the posterior margin of the wing; shaggy all black palps on fe-

males; clypeus without scales; a tuft of black scales on sternum VII, often

with white scales on other sterna; narrow pale bands on hindtarsi; larval seta

1-P branching from the base; larval palmate setae fully developed; and larval

seta 3-C usually bushy. In Southeast Asia, members of the barbirostris group

show more affinities to the hyrcanus group than to the albotaeniatus or umbro-

sus groups (see taxonomic discussion under hyrcanus species group).

BIOLOGY. Members of this group occupy a moderately wide range of

larval habitats usually associated with still water. Only barbivrostris and

campestris larvae are normally found closely associated with human habita-

tion in such habitats as rice fields, ditches and open temporary ground pools.

The remaining species are forest dwelling species with the larvae found in >

Harrison and Scanlon: Subgenus Anopheles in Thailand 81

shaded stream pools, ground pools or swamps. The adults of barbirostris

and barbumbrosus appear to feed largely on domestic and other large animals,

while campestris is normally more anthropophilic. Very little is known of the

feeding habits of donaldi, hodgkini and pollicaris, although donaldi is a proven

vector of human malaria pathogens in Malaysia.

MEDICAL SIGNIFICANCE. Currently, there is no evidence that any mem-

ber of this group serves as a vector of human pathogens in Thailand, although

barbirostris, campestris and donaldi have been incriminated as vectors of

malarial and/or filarial parasites in other Southeast Asian countries. For ad-

ditional remarks on this subject see the discussions under the respective spe-

cies.

KEYS. The following keys are based primarily on those of Reid (1968),

with changes to accommodate the Thaifauna. As will be noted, many of the

characters used do not offer clear cut options, particularly in the adult and

larval keys; consequently, a significant portion of larvae and adults will not be

identifiable unless they are accompanied by additional reared life stages. The

pupal stage is by far the most reliable stage for species determinations, thus

workers requiring precise identifications should bear this in mind when under-

taking studies in Thailand.

KEYS TO THE SPECIES OF THE AN, BARBIROSTRIS

GROUP IN THAILAND

ADULTS

Ey Apex of wing with lower pale fringe spot extending at least from vein

R4+5 to Mj; no white scales on the abdominal sterna.

barbumbrosus (p. 102)

Apex of wing with lower pale fringe spot narrow, at vein R45 only;

abdominal sterna with at least a few white scales on the midventral

line (hodgkini frequently without these scales). ......... 2

2(1). Apex of wing with only 2 narrow pale fringe spots, no pale fringe scales

at Ry; midtarsi usually without apical pale bands or patches... 3

Apex of wing with 3 pale fringe spots and middle spot at Ro, or with

only 2 pale fringe spots, but upper spot wide with pale scales down to

include Ro; midtarsomeres 1 - 2 usually with narrow apical pale

bands or patches. ..... TE IGS AMUN Ia aU a Soa ie Aa ane 4

3(2). White scales on abdominal sterna not confined to median tufts and a row

on each lateral margin, many scattered between these.

campestris* (p. 89)

White scales on abdominal sterna mostly confined to median tufts anda

row on lateral margins, a few scattered scales may occur between

median tufts and lateral margins. ...... barbirostris* (p. 83)

*This character is only 80-85% reliable, see taxonomic discussion under

barbirostris and campestris for further explanation. Also see couplet 4.

82 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

4(2). First foretarsal pale band short, half or less as long as tarsomere 5

(infrequent hodgkini males have a longer band), rarely crossing joint

onto tarsomere 2; median pale scales on abdominal sterna II-VI usu-

ally 0 - 20; central and south Thailand. ..... hodgkini* (p. 97)

First foretarsal pale band long, more than half as long as tarsomere 5,

usually crossing joint onto tarsomere 2; median pale scales on sterna

II-VI more than 20; extreme southern Thailand. .......... 5)

5(4). First foretarsal pale band longer than tarsomere 5, with 0.33 of band on

base of tarsomere 2; 2nd foretarsal pale band also crossing joint, with

0.25 - 0.33 of band on base of tarsomere 3; costa may have some pale

scales on preapical dark mark. .......... pollicaris (p. 100)

First foretarsal pale band seldom longer than tarsomere 5, with 0.25

or less of band on base of tarsomere 2; 2nd foretarsal pale band rarely

crossing onto base of tarsomere 3; costa without pale scales on pre-

Apical danke Maths Coes Sa Ge kw OR donaldi (p. 94)

PUPAE |

i. Trumpets without secondary cleft or seam. ............-. 2

Trumpets with secondary cletl Or sean... 2 ee ee ee ee 3

2(1). Seta 0-VII with 2 - 5 branches, rarely 2; abdominal seta 9 yellow or tan;

extreme southern Thailand only. ......... pollicaris (p. 100)

Seta 0-VII with 1 - 3 branches, rarely 3; abdominal seta 9 very dark

brown, rarely yellow or tan; central and south Thailand.

hodgkini (p. 97)

3(1). Trumpet without secondary cleft but with thickened seam; abdominal

seta 9 nearly black; seta 9-VII, 6 - 8 times as long as thick; sums of

branches on both setae 5-III less than 30.

barbumbrosus (p. 102)

Trumpet with secondary cleft, without seam; abdominal setae 9 yellow

to light brown; seta 9-VII shorter, 4 - 6 times as long as thick; sums

of branches on both setae 5-III 30 or more. ............-. 4

4(3). Sum of branches of both setae 2-VI is 17 - 58. . campestris** (p. 89)

Tire Sue tise Ste ee a er er COS eee D

5(4). Sum of branches on both setae 2-III is 11 - 19; found throughout Thai-

TA ia ee 8 barbirostris (p. 83)

This sum is 8 - 11; extreme southern Thailand only. . donaldi (p. 94)

LARVAE

1. Median dorsal valve of spiracular apparatus with caudal, thumb-

like, stigmal process; seta 9-C with 10 - 17 branches.

pollicaris (p. 100)

Median dorsal valve with small clear caudal knob, but no stigmal

process; seta 9-C with 5 - Il branches. .0.. 6. eo ne 2

*Approximately 15% of barbirostris males found throughout Thailand have

this phenotype. :

**Slight overlap occurs in these counts between barbirostris and campestris,

see discussion under barbirostris.

Harrison and Scanlon: Subgenus Anopheles in Thailand 83

2(1). Seta 3-C with 12 - 36 thin attenuated branches, usually lax and spread

OWLS Syne a ea Oia a i ee barbumbrosus (p. 102)

Seta 3-C with thick branches, usually stiff and crowded together (broom-

like); andaisually more numerous (19 95). ok ok oe ae. a

3(2). Seta 3-C with 19 - 44 branches, rarely more than 40; sum of branches on

both setae 90-III plus those on both setae 13-IV usually total 25 - 38.

hodgkini (p. 97)

Seta 3-C with 30 - 95 branches, rarely less than 40; sum of branches on

both setae 9-III plus those on both setae 13-IV rarely more than 24.

4(3). Difference between the sum of branches on both setae 13-III and that of

both setae 5=VIl is 0 = 10. 2 os Geek oe barbirostris (p. 83)

This difterence is:0-=-23 iow Wargie Wh Be ee 5)

5(4). Sum of branches on both setae 13-I is 27 - 45. . . . campestris (p. 89)

This sume ee: 2 ee Oe a ee OS donaldi (p. 94)

ANOPHELES (ANOPHELES) BARBIROSTRIS VAN DER WULP

(Figures 34, 35, 36)

Anopheles barbirostris Van der Wulp 1884: 248 (9).

Anopheles martini Laveran 1902: 907 (9).

Myzorhynchus barbirostris Van der Wulp, Theobald 1903b: 86 (¢, L*).

Anopheles (Anopheles) barbirostris of Edwards 1932: 40 (in part); Reid 1947:

89 (in part).

Anopheles (Anopheles) vanus of Bonne-Wepster and Swellengrebel 1953: 209;

Reid 1962: 14 (= barbirostris).

Anopheles (Anopheles) vanus of Wattal, Kalra and Gopal 1962: 65; Reid 1968:

129 (= barbirostris). |

Anopheles (Anopheles) barbirostris Van der Wulp, Reid 1962: 7 (“, 9*, P*, L

E*)s.Reid 1968:°:122-(C* 799. Pelt Bt):

Females are usually recognizable by: the absence of an accessory pale

wing fringe spot at vein Ro; a narrow pale fringe spot confined to R4,5; an

abundance of pale scales on vein Cu; few pale scales on the abdominal sterna;

and narrow apical pale bands on the foretarsi. The pupa has a secondary cleft

on the trumpet, numerous branches on setae 1, 5, on III - VII, few branches

on seta 2-VII and yellow to light brown setae 9. The larva rarely has less than

40 stiff broom-like branches on seta 3-C, a well developed and pigmented pal-

mate seta 1 on segment II and does not possess a stigmal process on the spir- |

acular apparatus.

FEMALE (Fig. 34) Head. Vertex with broad dark brown scales except

white scales just above interocular space; interocular space with long, mostly

dark setae and narrow white scales approximately 0.25 - 0.33 length of setae;

palpus dark with erect scales, giving very shaggy appearance; proboscis dark,

*The remaining species, hodgkini, barbirostris, campestris and donaldi are

not always identifiable in the larval stage without other associated stages.

84 Contrib. Amer. Ent. Inst., vol. L2j5on0 8} TOS

with erect scales except near apex; forefemur/proboscis ratio with 0. 83 - 0.92

range and 0.87 mean; pedicel with scales on upper-outer aspects; flagellomere

1 with dense patch of scales, remaining flagellomeres without scales. Thorax.

Integument dark gray, with darker longitudinal lines; eye spots not clearly de-

lineated; anterior promontory with moderate to narrow erect pale scales,

mixed with darker scales laterally; entire scutum except eye spots, covered

with short fine pale scales and long dark setae; scutellum with long dark setae;

anterior pronotum with prominent patch of dark scales on dorsocephalic aspect;

pleural setae: 2 - 5 propleural, 4 - 7 spiracular, 10 - 14 prealar, 6 - 11 upper

and 3 - 6 lower sternopleural, 11 - 16 upper and 0 - 3 lower mesepimeral; |

small pale scales usually associated with the propleural, sternopleural and

mesepimeral setae. Wing. Costa with small humeral pale spot, at least ven-

trally, and usually scattered pale scales between humeral area and small sub-

costal pale spot, remainder of costa dark to preapical pale area; remigium

dark scaled proximally and distally, usually with median pale scales; humeral

crossvein with distinct patch of dark scales; base of vein R mostly dark scaled,

with scattered pale scales out to poorly defined sector pale mark; Rj mostly

dark scaled, frequently few pale scales at subcostal area and on dark preapi-

cal area, tip of vein pale; Ro-R9,3 dark scaled to fork; Ro dark scaled except

pale area on distal 0.33, tip usually dark or slightly dark, at least on underside

of wing; Rg usually with approximately half of scales pale, but may be dark

scaled like Ro; R4;5 with mixed dark and pale scales, infrequently median 0. 80

pale scaled; M dark scaled to fork; M149 with base and apex dark scaled, vari-

able amount of dark and pale scales in between; M3+4 with base and apex dark

scaled, mostly pale scaled in between; Cu with distinct short dark mark near

base separated by its own length or more from upper dark mark on vein 1A,

remainder of vein out to apical dark mark on Cug with more than half of scales

pale, rarely over half dark; Cu fork normally without distinct dark mark; Cu,

normally with short dark mark at mediocubital crossvein, few scattered dark

scales out to short apical dark mark; Cug with preapical dark mark longer than

apical dark mark on Cuy; apex of Cus usually with small pale scale patch; 1A

normally with 2 short dark marks on distal half, proximal half may have scat-

tered dark scales; apical pale fringe spot narrow, at R4,5 only, rarely with

accessory pale fringe spot at vein Ro (see male description); posterior margin

of wing usually with narrow pale fringe spot at apex of Cug. Halter. Stem pale

scaled, knob dark scaled. Legs. All coxae with white scales; upper midcoxa

with 1 - 3 stout dark and 2 - 3 fine pale setae. Foreleg: Femur swollen basal-

ly, dark scaled except few pale scales on base at joint with trochanter and few

internal and external pale scales at apex; tibia dark scaled dorsally, paler ven-

trally, often few white internal and external scales at apex; tarsomere 1 dark

dorsally, paler ventrally, with apical white band half the length of tarsomere

5 or shorter; tarsomere 2 dark scaled except for apical pale band, pale band

on tarsomere 2 narrower than that on tarsomere 1; tarsomeres 3 - 5 entirely

dark scaled, or tarsomere 3 rarely with very narrow dorsoapical pale patch.

Midleg: Femur dark scaled dorsally, paler ventrally, with few white scales

on base and distinct internal and external white scale patches at apex; tibia

dark scaled dorsally, paler ventrally, with few white scales at apex of segment;

tarsomeres 1 - 5 entirely dark scaled, or infrequently with apices of tarso-

meres 1 - 2 with very small dorsal patch of light scales. Hindleg: Femur with

few white scales at apex, basal 0.33 of segment gray, darkening into black

scales near apex; tibia with narrow white basal band, dark scaled dorsally ex-

cept distinct dorsoapical white scale patch, paler scaled ventrally except at

apex; tarsomeres dark scaled, except narrow apical white bands on tarsomeres

Harrison and Scanlon: Subgenus Anopheles in Thailand 85

1 - 4 and narrow basal white bands or patches on tarsomeres 3 - 5. Abdomen.

Integument dark dorsally, paler ventrally, most sterna with silvery basal spot

on each side of midline; sterna II - VII with a median patch of pale scales, usu-

ally few pale scales forming lateral row along lateral margins, and infrequent

scattered pale scales in between lateral rows and median patch (see taxonomic

discussion); often few dark scales on sternum VI, in addition to prominent tuft

on VII.

MALE (Fig. 34) Head. Palpus usually completely dark scaled, may have

faint apical pale bands on segments 4-5. Wing. Generally paler than that of

female; veins Rg-R2+3 and M entirely dark scaled to forks; often with acces-

sory pale fringe spot at vein Ro. Abdomen. Dorsal surface of 8th segment

usually with small patch of narrow pale scales on median area, usually chang-

ing to dark scales apically. Genitalia. Basimeres usually with few basal pale

scales on dorsomesal aspect, many dark scales laterally and apically; ventral

lobe of claspette small and ill-defined, with 2 setae arising just off sclerotized

section, on clear membranous area; mesal seta of ventral lobe longer than

lateral seta and fused club on dorsal lobe; club on dorsal lobe composed of 4 -

. separate basal stems which fuse distally into single structure; aedeagus with

4 - 6 pairs of leaflets approximately half the length of aedeagus, largest pair

of leaflets with blunt basal tooth and fine teeth along both edges on distal 0.50 -

0.66, remaining large leaflets also with teeth on one or both edges.

PUPA (Fig. 35). Integument usually with tan to light brown pigmentation.

Cephalothorax. Wing case with mottled pattern of dark spots, lines or bars;

outer half of antennal case darker than inner half, with distinct dark rings at

joints and dark tip. Trumpet. Darkly pigmented, with thin uniform rim and

deep secondary cleft. Abdomen. Seta 9 with approximately same amount of

pigment as integument, not dark brown or black; 9-VII, 5 - 6 times as long as

wide; 9-VIII with few fine branches; setae 1, 5 on III - VII strongly developed

with many branches (15 - 55), frequently twisted or matted, often with a cen-

tral branch or stem which projects beyond main tuft; seta 8-II present or ab-

sent; seta 0-VI, VII usually with 2 - 4 branches; seta 2-III with 6 or more

branches, rarely less; seta 2-VI with 3 - 9 branches, rarely more (see tax-

onomic discussion). Paddle. Base darkly pigmented, also small pigmented

spot at apex; refractile margin approximately 0.75 of paddle length; seta 1-P

usually simple, may be split into 2 - 3 branches near apex.

LARVA (Fig. 36). Whole larvae dark. Head. Dark brown, antennae usu-

ally paler with numerous spines; setae 2-C simple and close together, rarely

bifid; seta 3-C with 32 - 65 stiff, broom-like branches, rarely less than 40;

seta 4-C with 2 - 5 branches; seta 6, 7-C with 8 - 14 and 8 - 18 branches re-

spectively; seta 8, 9-C with 7 - 11 and 6 - 9 branches respectively. Thorax...

Seta 1-P with 4 - 11 branches from near base; 11-P with 2 - 4 short stout,

equal-length branches from base; 14-M with 12 - 21 branches; 3-T with flat-

tened pale leaflets, some showing slight filament development. Abdomen.

Seta 1 with flattened, unpigmented leaflets on segment I and rarely II; 1, on

II - VII normally darkly pigmented and well developed; seta 13-I with 7 - 17

branches; 2-II with 5 - 7 branches; 5-III with 5 - 10 branches; 13-III with 3 -

12 branches, 13-IV with 3 - 5 branches; 5-VII with 4 - 8 branches; and 9-VII

with 1 - 4 branches; pecten with 9 - 12 long teeth, usually 9 - 10; seta 1-X

approximately equal to length of saddle.

TYPE-DATA. According to Reid (1962) the type of barbirostris is located

in the Leyden Museum. This female was collected on Mount Ardjoeno in east-

ern Java. The condition of this female was described in some detail by Reid,

who pointed out the difficulty of correctly identifying this specimen to barbi-

86 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

rostris, Reid felt the apparent absence of campestris on Java, coupled with

the hilly inland collection site, made the identity of this female as barbirostris

safe. The type-material of the synonym martini is composed of 2 females

mounted on separate glass slides, and located in the Pasteur Institut, Paris.

These 2 slides have identical labels: "A. martini, Montagnes des Pensat,

Cambodge, Fevrier 1902.'' We concur with Reid (1962) that the condition of

these specimens mounted in Canada balsam prohibits their positive identifica-

tion. Reid assigned this name as a synonym of barbirostris, because barbi-

rostris is normally a hill and mountainous species while campestris is a coastal

plain species. Our observations on the distribution of barbirostris and cam-

pestris, and the examination of the martini types, support Reid's assignment

of martini as a synonym of barbirostris.

DISTRIBUTION (Fig. 35). This species is very abundant and widely dis-

tributed in Thailand. It is likely to be found anywhere in Thailand where a

thorough search is made, except at higher elevations or in dense primary for-

est. In certain areas of Thailand, barbirostris may be uncommon. This is

particularly true just north of Bangkok in the rice plains along the Chao Phrya

River, where campestris is the predominant member of the barbirostris group.

Specimens (3260, 4519, 412 larvae and 1,211 larval and pupal skins) in the

USNM were examined from the following provinces of Thailand: Chanthaburi,

Chiang Mai, Chiang Rai, Chon Buri, Kanchanaburi, Khon Kaen, Krabi, Lam-

pang, Lampoon, Nakhon Ratchasima, Nakhon Si Thammarat, Nan, Narathiwat,

Pathum Thani, Phangnga, Phet Buri, Phrae, Phuket, Prachin Buri, Prachuap

Khiri Khan, Ranong, Rat Buri, Satun, Songkhla, Surat Thani, Trang, Trat,

Udon Thani and Yala. Additional specimens (68%, 1669, 23 larvae and 65 larval

and pupal skins) in the USNM were examined from: BURMA, CAMBODIA,

INDIA (Assam, West Bengal), INDONESIA (Sumatra), LAOS, MALAYSIA

(Peninsular), PORTUGUESE TIMOR, SRI LANKA and VIETNAM (North and

South). Additional literature references indicate barbirostris occurs through

the Indonesian chain as far east as the Moluccas. Reid (1968) claims this spe-

cies does not occur on Borneo, but several specimens in the USNM from Sabah

key to barbirostris in the larval and pupal stages, while the adults are inter-

mediate between barbirostris and donaldi. All previous records of barbiros-

tris from the Philippines apply to franciscoi, manalangi or vanus (Reid 1962).

Anopheles barbirostris is distributed from Pakistan in the west to Hainan Island,

Kwangtung Province, China in the east, and south down the Indochina Peninsula,

through Indonesia to the islands of Sulawesi and the Mollucas. According to

Chow (1949), this species has been recorded as far north in China as Szechwan

and Yunnan provinces, but these records need confirmation because of confu-

sion between barbirostris and barbumbrosus (Reid 1962).

TAXONOMIC DISCUSSION. Our current interpretation of barbirostris and

the other members of the group in Southeast Asia is due primarily to the efforts

of Reid (1962, 1968). However, a number of problems still exist.. Reid (1968)

discussed some of these and pointed specifically to campestris-like adults with

immature skins of barbirostris from the east coast of Peninsular Malaysia,

Chiang Mai, Thailand and Rangoon, Burma. He also observed several females

from Malaysia and southern Thailand, which had a pale accessory wing fringe

spot like donaldi adults, but had barbirostris immature skins. Reid summar-

ized his view on these problems as follows: ''This tendency, under certain

circumstances, for some adults of barbirostris to resemble those of campestris

or donaldi, means that identifications based on adults alone should not be re-

garded as reliable until confirmed by examination of the early stages, particu-

larly the pupae (and for donaldi the eggs). Once the species present in an area

Harrison and Scanlon: Subgenus Anopheles in Thailand 87

are known, identification of adults without early stages may be perfectly reli-

able.'' During the present study specimens with such overlapping variation

were not infrequent. In northern Thailand, frequent adults with dark wings

and numerous pale scales on the sterna were encountered, yet all proved to

be barbirostris when the immature skins were examined. Such disjunct asso-

ciations commonly occur in reared specimens from India (West Bengal) and

Sri Lanka. Nearly all the barbirostris examined from central and southern

Thailand, where campestris also occurs, have pale wings and relatively few

pale sternal scales. These characters would be most valuable if they were

distinct from those on campestris as Reid described from Peninsular Malaysia,

but that is not always true in Thailand (see campestris, taxonomic discussion).

In general, the wing characters previously used to separate barbirostris from

campestris are not valid in Thailand. The abundance of pale sternal scales

seems significant only in areas where both species occur, and sympatry can

be accurately determined only by the examination of immature skins.

Rare females of barbirostris exhibiting an accessory pale fringe spot on

the wing, can be confused with hodgkini, but should not be confused with donaldi

or pollicaris because these last 2 species have broad foretarsal pale bands and

both are apparently found only in extreme southern Thailand. To add to the con-

fusion, approximately 15% of barbirostris males from all over Thailand, but

particularly from the peninsular area, exhibit an accessory pale fringe spot on

the wing and can be confused with hodgkini. Sometimes this accessory spot is

not clearly defined and is confluent with the preapical fringe spot, consequently

the upper spot is wide with pale scales down to include vein Rog. On one aber-

rant female all pale fringe spots were missing, making identification possible

only through associated immature skins.

Specimens of barbirostris adults should be easily separated from barbum-

brosus, because adults of the latter have the lower apical pale fringe spot on

the wing extending down at least to vein Mj49, and the abdominal sterna of

barbumbrosus lack pale scales.

The pupa of barbirostris is the stage with the most reliable characters for

identification and unfortunately, the stage that is most often overlooked. Those

studies or persons requiring precise identifications should plan their programs

to include rearing immatures. Otherwise, specimens should be identified only

to the group level, i.e., barbirostris group. Anopheles barbirostris pupae

can rarely be confused with the other members of the group. The pupae of

barbumbrosus, hodgkini and pollicaris possess basic trumpet structural dif-

ferences from barbirostris. The pupae of campestris and donaldi (Malaysian

specimens) are similar to barbirostris, but the characters presented in the

key should separate 95 - 97% of these species. Of 778 barbirostris pupae

from Thailand, only 3 had the sum of setae 2-VI branches equal to or more

than 17, and all 3 had sums of 18. This range (6 - 18) overlaps slightly with

that of campestris (17 - 58), which only rarely has sums of less than 20.

The identification of barbirostris larvae in Thailand is extremely difficult.

Only barbumbyrosus with a non-rigid, few-branched seta 3-C and pollicaris

with a stigmal process, can be separated from barbirostris with consistency.

The larvae of the remaining species all possess characters which overlap

with those of barbirostris. The larva of donaldi has not been collected in Thai-

land, but Malaysian specimens are very similar to barbirostris. The larva of

campestris is usually distinct from that of barbirostris, but the characters

used to determine this are difficult to use. Larvae of the remaining species,

hodgkini, are frequently collected with barbirostris and often cannot be dis-

tinguished from them.

88 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

BIOLOGY. In general, barbirostris might be called a foothill mosquito.

In Thailand, immature collections have been made from sea level up to 500 m

in elevation, while adults were collected biting man (Scanlon and Esah 1965) at

elevations between 750 - 1400 m. Apparently, barbirostris is well suited for

life in most areas of Thailand, and utilizes a large variety of oviposition sites.

Immature stages have been collected from river margins, river pools, stream

margins, stream pools, flowing ditches, stagnant ditches, moats, lakes, per-

manent ground pools, temporary ground pools, flood pools, rice fields, wells,

canals, marshes, fish pools, rock pools, grassy ponds, seepage springs, buf-

falo and elephant footprints and a nipa palm swamp. A large majority of these

habitats also had emergent, floating and/or submergent vegetation. Usually

the habitats where barbirostris immatures are found are located in open sun-

light to light-moderate shade. Immatures of this species have rarely been col-

lected in heavy shade, which may account for its absence in primary forest.

Tolerance for high pollution levels has also been noted. Larvae have been

collected from sewage ditches, and ground pools with heavy concentrations of

buffalo dung and urine.

Adult barbirostris are generally zoophilic and can cause severe pest

problems for cattle and buffalo owners. This species will bite man, but not

on a large scale except in situations where the normal host animals are absent.

Reid (1961) found Malayan barbirostris were attracted to a calf 17 times more

frequently than to 2 men. More recently, Reuben (1971) found Indian "barbiros-

tris"' were attracted to a bullock 38 times as frequently as to 2 men. Adults

from Thailand have been collected: biting man and bovines, resting outside, _

resting in animal sheds, resting in houses, in light traps, in man baited traps,

in Shannon traps, in malaise traps and in a carbon dioxide net trap baited with

dry ice.

MEDICAL SIGNIFICANCE. This species has been incriminated several

times as a vector of malaria parasites; however, since the recognition of a

sibling group (Reid 1962), at least 2 of these reports have been corrected.

The "barbirostris"' reported as a malaria vector from the west side of the

Malay peninsula and that from Borneo are now recognized as campestris and

donaldi, respectively. Unfortunately, the other reports, from Sumatra and

Sulawesi, have not been clarified. Infact, it is not clear which species occur

on these islands. Griffith (1955) reported malarial oocysts from a single bar-

birostris from Chiang Mai Province, but the identity of the mosquito is now

suspect in light of Reid (1962). Since then no further malarial infections have

been found in members of this group from Thailand. Experimentally, barbi-

rostris showed a very low susceptibility to Plasmodium cynomolgi bastianellii

(Warren et al. 1963). In view of the zoophilic habits of this species, coupled

with a definite exophagic-exophilic behavior, there is little potential for bar-

birostris in Thailand being a vector of malaria pathogens.

The capabilities of barbivostris as a vector of filarial parasites deserves

consideration. Apparently, barbirostris in Peninsular Malaysia is not suscep-

tible to Wuchereria bancrofti infections (Wharton 1960), in fact, all the mem-

bers of this group have a low susceptibility to this parasite (Reid 1962). On

the other hand, several members are involved in the transmission of Brugia

malayi and studies in Malaya by Reid et al. (1962), Wharton (1962) and

Wharton, Laing and Cheong (1963) have shown that barbirostris is a good ex-

perimental vector of the periodic form, but a relatively poor vector of the sub-

periodic (semiperiodic) form of this parasite. Iyengar (1953) reported bar-

birostris from southern Thailand infected with B. malayi, however, this re-

port came prior to the recognition of the other sibling members of the group.

Harrison and Scanlon: Subgenus Anopheles in Thailand 89

Reid (1962, 1968) suggests that barbirostris is probably not involved in the

transmission of 6. malayi in Malaya due to its zoophilic habits. The same

should probably apply to Thailand.

The role of barbirostris in the transmission of animal parasites such as

Setaria and Dirofilaria needs serious study. Some of the early records of hu-

man filaria infections in barbirostris, may be due to misidentified animal fil-

ariae. Several zoophilic members of the hyrcanus group are known vectors of

animal filariae and the same is probably true of some of the zoophilic members

of the barbirostris group, particularly a species like barbirostris which feeds

on bovines. Harinasuta et al. (1970) found specimens identified as barbumbro-

sus and campestris from Kanchanaburi Province, Thailand positive for Divro-

filaria larvae.

ANOPHELES (ANOPHELES) CAMPESTRIS REID

(Figures 37, 38, 39)

Anopheles (Anopheles) campestris Reid 1962: 15 (**, 2*, P, L*, E); Reid

196842014 2%,0 PR, Be B).

Adults are not normally distinct from those of barbirostris, but they often

have darker wings and more pale scales on the abdominal sterna. The pupa

is the best stage for identification, and is nearly always separable by numer-

ous setal branches. The larva is very much like barbirostris except for

more numerous branches of some setae. This species is like barbirostris

except for:

FEMALE (Fig. 37) Head. Forefemur/proboscis ratio with range of 0.75 -

0.86 and 0.83 mean. Thorax. Pleural setae: 2 - 6 propleural, 4 - 9 spiracular,

8 - 12 prealar, 5 - 9 upper and 3 - 5 lower sternopleural and 9 - 16 upper and

0 - 2 lower mesepimeral. Wing. Often darker than wing of barbirostris; vein

Cu-Cug frequently with more dark than pale scales (caution-see taxonomic

discussion); tip of wing frequently with pale stripe extending from preapical

pale costal spot back to Mj,9; tip of wing without accessory pale fringe spot

adjacent to tip of vein Ro. Legs. Foretarsomere 3 usually without apical pale

band; midtarsomeres usually without pale bands. Abdomen. With numerous

pale scales on sterna II - VII, particularly in median patches and lateral rows;

pale scales scattered between median sternal patches and lateral rows also

numerous; sternum VI rarely with dark scales on caudal margin.

MALE (Fig. 37). Like female, generally with more dark scales on wing;

without accessory pale fringe spot adjacent to tip of vein Rg; distal half of

dorsal surface of 8th abdominal segment usually with central patch of dark

scales and pale scales on lateral aspects. Genitalia. Aedeagus with 3 - 6

pairs of leaflets.

PUPA (Fig. 38). Like barbirostris except setae 0 and 2 have more branches

on most segments; 0 on VI - VII usually with 4 - 6 branches; 2-III with 8 or

more branches; 2-VI with 6 - 29 branches (see taxonomic discussion).

LARVA (Fig. 39) Head. Seta 3-C with 60 or more stiff, broom-like bran-

ches; 8, 9-C with 6 - 9 and 6 - 11 branches respectively. Thorax. Seta 11-P

with 3 - 7 branches; 14-M with 20 - 28 branches. Abdomen. Seta 13-I with 12 -

27 branches; 5-III with 5 - 8 branches; 13-III with 8 - 21 branches; 13-IV with

3 - 7 branches; 9-VII with 2 - 5 branches.

TYPE-DATA. The 2? holotype, “ allotype and 1“ and 22 paratypes, with

larval and pupal skins (except “ paratype) are in excellent condition and are

90 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

located in the BMNH. The holotype and allotype have the respective rearing

numbers "L.H. 75/10" and "L.H. 75/2" and are from, ''Rantau Panjang, Klang,

Selangor, Malaya."' Two of the paratypes (“ and 2) are from Selangor, Malaya,

and the 2? has a rearing number, "L.H. 81/7."' The remaining paratype (?) is

from Perak, Malaya, and has a rearing number, ''073/21."'

DISTRIBUTION (Fig. 38). The actual distribution of campestris in Thailand

is still unknown; however, the confirmed (associated immature skins) distribu-

tion depicted on Fig. 38 hints this species is common in the Chao Phrya River

basin and the coastal areas of the country. Apparently, campestris is confined

to areas of low elevation in Thailand, as Reid (1962) found in Malaya. The

northern or northeastern limits of its range are currently not known. In the

northeast (Korat Plateau), campestris definitely occurs in the vicinity of Khon

Kaen, where barbirostris and barbumbrosus were also collected. The Korat

Plateau, although 125 m in elevation, has extensive flat rice plains which ap-

pear to be one of the primary habitats for campestris. Further west, campes-

tyis has not been found north of Ayutthaya. Fairly frequent adults with a

campestris -like habitus have been collected in Chiang Mai and Lampang, but

associated immature skins show these are actually barbirostris.

Locally, campestris can be extremely abundant, and the predominant mem-

ber of the group. This is particularly true in Ayutthaya, Nonthaburi and

Pathum Thani, just north of Bangkok.

In other areas it may share certain habitats with barbirostris, and be much

less common. Normally, such mixed populations are not the rule. Specimens

(137%, 1792, 175 larvae and 386 larval and pupal skins) inthe USNM were examined

from the following provinces of THAILAND: Ayutthaya, Chanthaburi, Chon Buri,

Khon Kaen, Nakhon Si Thammarat, Nonthaburi, Pathum Thani, Prachin Buri,

Phra Nakhon, and Songkhla. Additional specimens (4c, 6 and 12 larval and

pupal skins) were examined in the USNM and BMNH (type-series) from MALAY-

SIA (Peninsular Malaysia). Reid (1962, 1968) discussed the possibilities of

campestris occurring in India, Burma, and the Indo-China region. Although

specimens from some of these countries or regions were examined and found

to have a campestris -like habitus, all having associated immature skins proved

to be barbirostris. Consequently, confirmed records of campestris are still

known only from Malaysia and Thailand.

TAXONOMIC DISCUSSION. As early as 1942, 2 types of barbirostris were

recognized in Malaya (Reid 1942). The dark-winged type was recognized by

Reid (1962) as a distinct species, campestris, not only on morphological dif-

ferences, but also on behavior traits and vector capabilities quite distinct from

those of barbirostris. Of the species of the barbirostris group found in Thai-

land, campestris is one of the most difficult to recognize, yet because of its

known vector-disease pathogen associations in Malaysia, it is the most impor-

tant member of the group to identify. Reid (1962, 1968) found campestris in

Peninsular Malaysia with relatively stable characters and easily separable from

barbirostris; however, this is not the case in Thailand. In Pathum Thani and

Ayutthaya, north of Bangkok, campestris is extremely abundant and quite vari-

able. While 134 confirmed females from these areas had extensive pale scal-

ing on the abdominal sterna as is normal for campestris, nearly 50% of these

exhibited pale scaling on Cu-Cup and other wing veins as is normally found on

barbirostris. Specimens from these areas with these combinations of char-

acters have caused much confusion for past identifiers. Now it is clear that

the wing characters used by Reid to separate Malayan campestris and barbi-

vostris are not valid in Thailand. Not only do many Thai campestris have light

wings, but in areas where campestris does not occur some specimens of bar-

Harrison and Scanlon: Subgenus Anopheles in Thailand 91

birvostris exhibit dark wings.

The pattern of pale scales on the abdominal sterna used by Reid to Eeoavate

Malayan barbirostris and campestris is valid in some, but not all areas of

Thailand. These valid areas seem to be those where both species occur or

meet. The pale sternal scale pattern on Thai campestris is nearly stable

and like that depicted for Malayan campestris. On the otherhand, while bar-

bivostris usually exhibits few pale sternal scales in areas where it is found with

or near campestris, ‘it often exhibits extensive pale sternal scaling in areas

where campestris does not occur. Consequently, the only areas where these

2 species may be separated by an adult character with any degree of accuracy

are areas where they both occur. However, such areas of sympatry must first

be determined by examining the immature skins of individually reared speci-

mens.

The number of branches on pupal seta 2 onthe abdominal segments is

highly variable, particularly when using the summation method; however, of

249 pupal skins on which the sum of the branches on both setae 2-VI were

counted, only 6 skins had counts of less than 20 branches. These counts are

19(1), 18(2), 17(2) and 13(1). Since barbirostris has a range up to 18 branches,

there is a Slight overlap, but these 2 species should still be separable by this

character on a 95 - 97% level. Several dark winged adults with extensive pale

sternal scaling and immature skins were available from the rice fields of the

Chiang Mai Valley. Two of these had the sum of seta 2-VI on the pupal skin

with 18 branches. Since we could not determine whether these represented the

lower range of campestris or upper range of barbirostris, and since there

were so few specimens we have tentatively identified these as barbirostris.

The Chiang Mai Valley is extensive, flat, at approximately 300 m elevation

and used primarily for rice cultivation. Further work is definitely needed in

this valley to determine if campestris is present.

Two campestris pupal skins with aberrant characters were observed. One

had only slight notches for trumpet secondary clefts. The other had the chaeto-

taxy on the left side of abdominal segment VI normal, while on the right side

setae 2 - 3 and their alveoli were missing, while seta 1 was represented twice.

The larval chaetotaxy of campestris is more variable than that of the pupa

and overlaps with characters used to separate barbirostris. Consequently,

the larval stage is less reliable than the pupal stage for separating these 2

species. Larval skins of campestris are best used to confirm pupal identifica-

tions.

BIOLOGY. As expressed by the name campestris (of fields, of plains),

this species is typically found on flat coastal or rice plain areas. Immatures

have been collected in Thailand from 1 - 200 m elevation. Larval habitat re-

quirements are uncertain; however, clear still fresh water, some emergent,

floating or other vegetation, and light-moderate shade are usually present.

Only 2 collections were made in Thailand in which the water was possibly brack-

ish. Immature stages have been collected in Thailand from rice fields, mar-

shes, sumps, canals, ponds, wells, large and small pits, ditches, hoofprints,

stream margin pools, ground pools, flood pools, a nipa palm swamp and fresh

standing water in a large stump.

Adult behavior of campestris has been investigated during a number of

studies in Malaysia. Reid (1942), during laboratory feeding experiments on

malaria and filariasis patients, found that 60% of the dark-winged barbirostris

(campestris) fed on man to only 26% of the light-winged barbirostris. In 1961,

Reid continued attractiveness studies and showed that when 2 men were com-

pared with one calf, 77% of the campestris selected the men, compared to only

92 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

6% of the barbirostris. When 2 men were compared with 2 goats this study

showed 85% of the campestris attracted to the men. Reid and Weitz (1961)

conducted a blood meal identification (precipitin) study on 27 campestris

found resting outdoors and showed 17 (63%) had fed on man, 8 on monkeys and

2 undetermined. Wharton et al. (1964) followed this by comparing man and

monkey baited net traps at ground level and found 92% of the campestris came

to the man baited trap. No campestris came to a monkey baited trap in the

canopy. Because of these studies, campestris is listed by Reid (1968) as

highly anthropophilic and ranked as the 3rd species of Anopheles in Southeast

Asia most attracted to man. Anopheles campestris is also highly endophagic.

Moorhouse and Wharton (1965) showed this species biting man indoors at a

ratio of 4.4 : 1 over man outdoors. Of the campestris collected during this

study (6 nights), 85% were caught fairly evenly distributed between 2000 - 0200

hours. This study also showed campestris is endophilic when pyrethrum knock-

down catches revealed 51 campestris resting in 15 of 27 houses between 1600 -

1730 hours. The real significance of these anthropophilic, endophagic and

endophilic traits was demonstrated by Moorhouse (1965) during a malaria

eradication pilot project in Malaysia. This house spraying project completely

eliminated campestris in the project area, while barbirostris populations in

the project area were unaffected. Outside the project area, campestris popu-

lation levels remained unchanged. Another useful campestris behavior trait

has been detected. Moorhouse and Wharton (1965) found in comparing man

biting and man baited net trap collections that over twice as many campestris

were captured by the net trap method. Reid (1968) carried this work one step

further and determined in 178 nights of comparing catches between man in huts

with window traps and man baited net traps outside, that 69% (1, 300/1, 897)

were collected in the outside net traps. This information is significant, par-

ticularly for studies such as vector-dissection surveys, where the number of

specimens collected is important.

Adult behavior of campestris in Thailand is extremely difficult to inter-

pret because of overlapping adult characters and the possibility of mixed col-

lections. The behavior data accrued in the Malayan studies may also apply

to Thai campestris, but this needs confirmation. The only study reported to

date from Thailand where pure (or nearly so) campestris populations occur,

is that of Gould et al. (1965). This vector-dissection study was conducted in

Pathum Thani, and 961 campestris were collected on human bait between

1900 - 2400 hours (D. J. Gould, personal communication). Specific mention

was made that of the 6 anophelines studied, campestris was the only species

most frequently collected indoors. These data indicate that Thai campestris

at least from this area, are anthropophilic and endophagic.

MEDICAL SIGNIFICANCE. To date, campestris has not been incriminated

as a vector of human pathogens in Thailand; however, this is very possible in

view of its vector capabilities just across the border in northern Malaya.

Hodgkins' (1956) studies in western Peninsular Malaysia quite convincingly

incriminated the dark-winged type of barbirostris (campestris) as a vector of

human malaria parasites. This study plus all the malaria dissection studies

involving barbirostris group members in Malaya prior to 1962 are tabulated

in Reid (1962). Later studies, Moorhouse and Wharton (1965) and Moorhouse

(1965) present further data to incriminate campestris as a vector of human

malaria pathogens in the lowland-coastal areas of western Peninsular Malay-

sia. Reid (1962) noted that where campestris is the vector of human malaria

parasites there is a tendency for Plasmodium vivax (Grassi and Feletti) to be

the predominant parasite. Although some evidence was presented to explain

Harrison and Scanlon: Subgenus Anopheles in Thailand 93

this tendency, further confirmative studies are needed. Reid and Weitz (1961)

determined that some wild campestris feed on monkeys, and suggested that

campestris may also be involved in the transmission of simian malaria. How-

ever, Warren et al. (1963) showed in a laboratory experiment that campestris

had a very low level of susceptibility to infection with Plasmodium cynomolgi

bastianellii. Furthermore, Wharton et al. (1964) did not find simian malaria

parasites in wild caught campestris, in an area where there was active trans-

mission of these parasites.

Only 2 vector-malaria dissection studies have been conducted in Thailand

where we feel confident the specimens referred to are campestris. The 1st

reported by Reid (1962), refers to a study conducted in Nonthaburi by Dr. Udaya

Sandhinand in 1960 when over 2, 000 "'barbirostris"' were dissected. The 2nd

is the study conducted by Gould et al. (1965) in Pathum Thani, where 961 cam-

pestris were dissected. Neither study revealed malaria parasites in campes-

tris. Griffith's (1955) report of a single malaria infection in barbirostris from

Chiang Mai may actually refer to campestris, although we currently do not

consider campestris to occur in the Chiang Mai Valley. It is entirely possible

that campestris did occur in Chiang Mai in the past and has since been elim-

inated by the house spraying conducted by the Thai malaria eradication program

since the late 1940's. The feasibility of total elimination (eradication) of cam-

pestris from an area was demonstrated by Moorhouse (1965).

The status of campestris as a vector of filarial parasites is now well estab-

lished, but early work in Malaya was contradictory and created considerable

confusion until the 2 forms of Brugia malayi (Wilson et al. 1958) and the sib-

ling species in the barbirostris group (Reid 1962) were recognized. Studies

conducted in Malaya prior to 1962 are summarized by Reid (1962). Studies

conducted by Wharton (1962) in the laboratory show campestris is a very effi-

cient vector of periodic B. malayi, a poor vector of subperiodic B. malayi,

an efficient vector of B. pahangi and refractory to infections with Wuchereria

bancrofti. Reid et al. (1962) found 19/3, 573 or 0.53% of wild caught specimens

from a northwest Malayan study area infected with 3rd stage (mature) larvae.

During this study Reid fed campestris upon an infected volunteer from Kedah

State. Most of these specimens (22/24 or 88%) had mature B. malayi larvae.

Anopheles campestris is now considered one of the major vectors of periodic

B. malayi in western and northwestern Peninsular Malaysia.

Iyengar (1953) conducted the first major filariasis survey in southern Thai-

land and reported 42 of 358 "barbirostris" positive for B. malayi larvae.

These "'barbirostris'' were collected resting in houses during morning hours,

and thus may have been campestris, which is endophilic. Nair and Chayabejara

(1961) reported that the periodicity of the B. malayi in southern Thailand was

strictly nocturnal with peak peripheral microfilaremia between 2100 - 0500

hours. This periodicity is very similar to the peak period of man-biting ac-

tivity exhibited by Malaysian campestris (Moorhouse and Wharton 1965).

Harinasuta et al. (1964) attempted to incriminate the mosquito vector(s) of B.

malayi in southern Thailand by outdoor man-biting collections between 1800 -

2000 hours. The selection of this time period favored the collection of Man-

sonia species that can transmit periodic B. malayi, and reduced the possibility

of collecting infected specimens of the more nocturnal and endophilic campes-

tris. Infact, Mansonia uniformis (Theobald), was found positive (0.6%), while

none of the anophelines were positive. Since campestris is highly endophilic,

the indoor spray program used by Harinasuta et al. was probably very effec-

tive; however, such results are not evident because only outdoor man-biting

densities were monitored. Harinasuta et al. (1970) also reported Dirofilaria

94 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

sp. larvae in campestris from a study near the Burma border in Kanchanaburi,

however, this determination is suspect due to overlapping adult variation and

the absence of any confirmed campestris from that province. Large numbers

of confirmed barbirositris group members have been examined from Kanchana-

buri, but all were barbirostris, barbumbrosus or hodgkini.

The possibility of campestris being or having been a vector of malarial or

filarial pathogens in Thailand is very real. Hopefully, campestris no longer

presents a disease vector threat because of the long term malaria eradication

house spraying program. However, if future foci of human malaria or filarial

diseases are located in Thailand where campestris is also abundant, an imme-

diate house spraying program seems to offer the best means of control. Ap-

parently, campestris represents one of the few vectors in the world that has

behavorial traits that permit rapid elimination by chemical control methods.

ANOPHELES (ANOPHELES) DONALDI REID

(Figures 40, 41, 42)

Anopheles (An.) barbirostris of Colless 1948: 80 (*, 2*, L*).

Anopheles (An.) donaldi Reid 1962: 17 (*, 9*, P, L, E*); Reid 1968: 132

(C40 9" Ba ee).

Besides the egg, the adult is the easiest stage of donaldi to recognize.

Usually the presence of broader foretarsal pale bands, an accessory pale wing

fringe spot at R,, narrow pale bands on the midtarsi and fairly numerous pale

abdominal sternal scales are indicative of donaldi. However, other species

can also exhibit these characters, so identifications should be confirmed by

examining as many stages as possible, particularly the egg. This species is

like barbirostris except for:

FEMALE (Fig. 40) Thorax. Pleural setae: 3 - 7 propleural, 5 - 8 spiracu-

lar, 12 - 16 prealar, 5 - 8 upper and 3 - 4 lower sternopleural, 11 - 16 upper

and 0 - 4 lower mesepimeral. Wing. Usually paler, pale areas usually more

continuous with fewer scattered dark scales; tip of Ro rarely with dark scales

on upper or under surface; tip of wing with accessory pale fringe spot adjacent

to Ro. Legs. Upper midcoxa with 2 - 5 setae; foretarsomeres usually with

broader pale bands (see taxonomic discussion); 1st band 0.5 - 1.0 length of tar-

somere 5, usually extending as narrow band onto base of tarsomere 2; 2nd band

0.5 or less length of tarsomere 5, sometimes extending as faint band onto base

of tarsomere 3; 3rd band usually present as very narrow band, or distinct dor-

sal pale spot on apex of tarsomere 3; tarsomere 4 - 5 dark scaled. Midtarso-

meres 1 - 3 usually with narrow apical pale bands or small dorsoapical pale

spots. Abdomen. Sterna usually with slightly more pale scales than found on

barbirostris.

MALE (Fig. 40) Head. Palpus with distinct narrow pale bands at apex of

segments 3 - 4. Legs. First foretarsal band often longer than length of tarso-

mere 5. Wing. Generally paler than that of female, with accessory pale fringe

spot infrequently confluent with preapical pale fringe spot; consequently, upper

pale fringe spot is wide with pale scales down to include Rg. Abdomen. Sterna

with pale scales generally confined to median area; dorsal surface of 8th seg-

ment with median patch of pale scales. Genitalia. Aedeagus with 3 - 4 pairs

of leaflets.

PUPA (Fig. 41). Nearly identical to barbirostris except abdominal seta

2 has fewer branches on segments III, VI; sum of branches of seta 2-III is 8 -

Harrison and Scanlon: Subgenus Anopheles in Thailand 95

11 (11 - 19 barbirostris), and 2-VI is 5 - 11 (7 - 18 barbirostris).

LARVA (Fig. 42). According to Reid (1968), ''Not always distinguishable

from barbirostris. Frontal hairs 6 and 7 on the head with slightly more bran-

ches, 10 - 17 and 15 - 20; the difference is accentuated a little by summing the

branches on all four hairs giving a total of 50 - 75 (barbirostris 39 - 62). Ab-

dominal hair 13, III with rather more branches (8 - 14) and 5, VII averaging

slightly fewer (4 - 6), with the result that there is a larger difference between

the sums for the pairs 13, III and 5, VII, this difference is 10 - 17 (barbiros-

tris 0 - 10). Other sums of the branches on pairs of hairs are as follow: 9,

VII 8 - 11 (barbirostris 10 - 15), 13, 118 - 27 (campestris 27 - 45), 13, II

20 - 29, 13, 11119 - 26 (barbirostris 10 - 24), 13, IV 6 - 8 (hodgkini 10 - 18);

for the two pairs 5, VII + 13, I, 26 - 38 (campestris 40 - 57)."

TYPE-DATA. The 2 holotype, “ allotype and 1% and 22 paratypes, with

larval and pupal skins (except paratype #01856) are located in the BMNH. The

holotype, allotype, 1% and 12 paratype are numbered ''0421/31; 0421/33; 0410/

47: and 0434/14" respectively and are from ''Puchong, Selangor, Malaya."'

The other 2 paratype is numbered ''01856" and is from, ''Ulu Lui, Ulu Langat,

Malaya.'' The “ genitalia of the paratype has been mounted on a slide, while

that of the allotype is still intact.

DISTRIBUTION (Fig. 41). In Thailand, donaldi is apparently uncommon

and found only in the extreme southern portion. Reid (1968) says that donaldi

is locally common in central and southern Peninsular Malaysia, is the most

common member of the group in Borneo and may be present in Sumatra and

Java. No new distributional data has come to light since Reid's work. A single

female in the USNM was examined from Narathiwat Province, THAILAND. An

additional 40°, 132 and 4 larval and pupal skins in the USNM were examined

from MALAYSIA. The type-specimens (2%, 32 and 8 larval and pupal skins) in

the BMNH from MALAYSIA (Peninsular Malaysia) were also examined.

TAXONOMIC DISCUSSION. This species, like campestris, is extremely

close to barbirostris and specimens of donaldi frequently cannot be identified

without other associated stages for verification. Originally, donaldi was de-

tected by distinctive egg characters. Although we have not dealt with eggs in

this study, egg characters appear the best means of separating donaldi from

barbirostris; therefore, the following extract from Reid (1968) is included,

"Tength 0.48 mm., somewhat shorter than that of barbirostris. Differs dis-

tinctly in usually having the deck divided into an area at each end, or if not

divided then narrowed towards the middle; also the floats are relatively longer,

about 3/4 as long as the egg, and the tail end is less upturned and pointed.

Float ribs about 28-34."' The egg of barbirostris has a single deck slightly

more than 0.1 as wide as the egg width, floats about 0.67 the egg length and

28 - 44 ribs in the floats. Since only one specimen is available from Thailand

and only a few available from Malaysia, little variation was observed. Al-

though the foretarsal pale bands are long on the Thailand female, the bands do

not cross over onto the bases of tarsomeres 2 - 3. The foretarsal pale bands

also appear shorter like barbirostris, on the specimens from East Malaysia,

while those from Peninsular Malaysia and the Thailand specimen are as des-

cribed. Reid (1968) noted that the pupal and larval characters used to separate

donaldi and barbirostris in Peninsular Malaysia were less reliable in East

Malaysia where barbirostris is apparently absent. Finding characters that are

distinct where these 2 species are found together, and then break down when

only one species is present in an area, is identical to the situation found with

barbirostris and campestris in both Malaysia and Thailand.

The variation noted by Reid (1968) regarding the accessory pale fringe spot,

96 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

particularly on males, is essentially identical with that found in about 15% of

the barbirostris males from peninsular Thailand (see barbirostris, taxonomic

discussion). With the previously described character trends in mind, it would

be interesting to determine if this donaldi-like character on barbirostris is

another character breakdown where only one species occurs.

Reid (1962) noted males usually have more pronounced pale palpal bands

than males of barbirostris; however, we were unable to detect any difference in

the size of these bands.

BIOLOGY. In Borneo where donaldi seems most common, Colless (1948,

as barbirostris) notes the larvae were common in most ground water habitats

with some vegetation and not exposed to direct sunlight, or deep shade as in

jungle pools. He listed rice fields, overgrown drains and open sedge swamps

as the most favored habitats. Reid (1962) said that in Malaya, donaldi has to

compete with barbirostris and tends to be more local and is found in more

shady breeding sites, such as river swamps or the edge of the forest in more

inland hilly areas. Moorhouse (1965) found donaldi larvae in Selangor, Malay-

sia, at the edge of ''swamp-forest"' under moderate shade and in lightly shaded

sedge ''swamps.'' Adults of donaldi have been collected by a number of tech-

niques, including: resting, biting human bait inside and outside houses, biting

cattle, resting near poultry, pigs and cattle, and in net traps using human bait

(Reid 1962; Moorhouse and Wharton 1965; Macdonald et al. 1967). Adult be-

havior has proven somewhat unusual for a member of the barbirostris group.

Like some members of the umbrosus group, donaldi actively bites in shaded

forest during the day, then enters settlements and houses at night to bite. Reid

(1968) suggests donaldi is less zoophilic than barbirostris, but confusion exists

about this aspect of donaldi behavior (Reid 1962). Reports indicate that donaldi

readily enters houses at night to bite, however, tests in Malaya (Moorhouse and

Wharton 1965) show a slight preference for biting outside. Although donaldi

bites in houses at night, it does not remain in the houses during the day as does

campestris, but apparently returns to the shaded forest (Moorhouse and Whar-

ton 1965). In summation, donaldi is reported to: readily bite man, but show a

preference for cattle; enter houses to bite at night, but show a preference to

bite outside; leave houses before daylight; and bites in the shaded forest during

daylight hours.

MEDICAL SIGNIFICANCE. The role donaldi plays in the transmission of

human pathogens in Malaysia is becoming increasingly evident. This species

is clearly a vector of periodic Brugia malayi in the inland hills areas of Pen-

insular Malaysia and is probably a vector of this parasite in Borneo (Reid 1968).

Peninsular Malaysian donaldi were found to have a very low level of suscepti-

bility to Wuchereria bancrofti (Wharton et al. 1963), however, de Zulueta

(1957) considered donaldi a vector of this parasite in Borneo.

The role of donaldi in the transmission of malaria parasites is still unclear.

Reid (1962) summarized the few records of donaldi found naturally infected with

malaria parasites in Borneo. However, in 1968 he suggested these parasites

were possibly not of human origin because Wharton, Eyles et al. (1963), Moor-

house (1965) and Moorhouse and Wharton (1965) found oocysts in Peninsular

Malaysian donaldi, resembling the mousedeer malaria, Plasmodium tragult

Garnham and Edeson. The 2 studies by Moorhouse also reported finding

primate-like oocysts in donaldi, but these infections could not be correlated

with any human malaria in the study areas. Moorhouse (1965) noted that in

previous experimental susceptibility studies donaldi from Peninsular Malaysia

failed to transmit Plasmodium falciparum (Welch) and was a poor host for P.

traguli parasites. Furthermore, Warren et al. (1963) and Bennett et al. (1966)

Harrison and Scanlon: Subgenus Anopheles in Thailand 97

have shown Peninsular Malaysian donaldi to be refractory to strains of P.

cynomolgi, one of the common primate malaria parasites in Malaysia. Despite

these confusing reports and in the absence of any confirmed human malaria

parasites having been found in Peninsular Malaysian donaldi, Reid (1968) still

lists donaldi as a minor vector of human malaria parasites in Malaysia. Field

and laboratory studies by Hardin, Santa Maria and Liaw (1973) now lend support

to this contention. These authors found 7 of 4,303 wild caught donaldi from

Sarawak positive for sporozoites, and found laboratory reared donaldi highly

susceptible to both P. falciparum and vivax infections. These infected mos-

quitoes were also able to transmit both parasite species to a susceptible human

volunteer. A comparison of these results with those achieved using Peninsular

Malaysian donaldi would suggest that the strain of donaldi in East Malaysia (at

least Sarawak) is much more receptive to infection with malaria parasites.

This suggestion is one means of clarifying the confusion surrounding the malaria

vector capabilities of donaldi, however, further research is needed. Regard-

less of the vector capabilities of donaldi in Malaysia, it is apparently too scarce

in Thailand to serve as a vector of human pathogens.

ANOPHELES (ANOPHELES) HODGKINI REID

(Figures 43, 44, 45)

Anopheles (An.) hodgkini Reid 1962: 20 (%, 9*, P*, L*, E*); Moorhouse

1965: 113 (2); Reid 1968: 134 (", 9*, P*, L*, E*).

Best recognized in the adult and pupal stages. Adult females have an

accessory pale wing fringe spot at vein Ro and a narrow apical pale fringe spot

confined to R45, few to no pale scales on the abdominal sterna, and narrow

pale tarsal bands like those of barbirostris. The pupa can be identified by

the absence of a secondary trumpet cleft or fold and the presence of relatively

short, dark brown setae 9. This species is like barbirostris except for:

FEMALE (Fig. 43) Thorax. Pleural setae: 3 - 5 propleural, 5 - 8 spiracu-

lar, 7 - 12 prealar, 3 - 7 upper and 3 - 4 lower sternopleural, 6 - 11 upper and

0 - 4 lower mesepimeral. Wing. Paler than barbirostris; tip of Ro rarely with

dark scales on the upper or under surface; tip of wing with accessory pale fringe

spot adjacent to Ro. Legs. Upper midcoxa with 2 - 3 setae; midtarsomeres

1 - 3 usually with narrow apical pale bands or small dorsoapical pale spots.

Abdomen. Sterna with fewer pale scales than found on barbivostris; lateral

and diagonal sternal pale scales absent, only median groups remain, with 0 -

. scales per segment; sterna may not have pale scales (see taxonomic discus-

sion), but sternum VII usually with 2 - 4 anterior pale scales.

MALE (Fig. 43) Head. Palpus with distinct narrow pale bands at apex of

segments 3 - 4. Wing. Generally paler than that of female, with accessory

pale fringe spot adjacent to Ry; accessory pale fringe spot often confluent with

preapical pale fringe spot, consequently, upper pale fringe spot is wide with

pale scales down to include Rg. Legs. First and 2nd foretarsal pale bands

2 - 3 times as long as broad, that on tarsomere 1 frequently half or more as

long as tarsomere 5, seldom crossing onto base of next tarsomere. Abdomen.

Dorsal surface of 8th segment with median patch of mixed pale and dark scales.

Genitalia. Aedeagus with 4 - 6 pairs of leaflets.

PUPA (Fig. 44) Cephalothorvax. Trumpet without secondary cleft or fold,

with shallow emargination or, infrequently with small darkened indentation

at normal site of secondary cleft. Abdomen. Seta 0-VII with 1 - 2 branches,

98 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

rarely 3 and then only on one side; setae 9 dark brown, rarely yellow or tan;

seta 1 and particularly 5 on segments IV - VII with branches more densely

clumped than on barbirostris.

LARVA (Fig. 45). Not always distinguishable from barbirostris and

occasionally barbumbrosus. Head. Seta 3-C with 19 - 44 branches usually

stiff and broom-like, rarely non-rigid or more than 40. Thorax. Seta 13-P

with 9 - 15 branches (7 - 9 barbumbrosus); 14-M with 8 - 14 branches (12 -

21 barbirostris: 20 - 28 campestris). Abdomen. Seta 2-I with 6 - 7 branches

(3 - 6 barbumbrosus); 2-II with 7 - 12 branches, rarely 5 - 6 (5 - 7 barbiros-

tris and campestris); sum of branches for both setae 13-IV is 10 - 18 (6 - 9

on barbirostris and donaldi); seta 9-VII with 4 - 7 branches (1 - 4 barbirostris:

2 - 5 campestris),

TYPE-DATA. The 2? holotype, “ allotype and 22 paratypes, with associ-

ated larval and pupal skins (except paratype #0195) are located in the BMNH.

The holotype is numbered ''0150/7"' and comes from '16th mile Gombak,

Selangor, Malaya, Feb. 1957, J. A. Reid.'' The allotype has the same data,

but is numbered ''0150/1"'. The 2 paratypes are from "11 mile, Ulu Gombak

Rd., April 1957", in Selangor, Malaya, and are numbered "0195 and 0195/1"

respectively. All 4 specimens are in excellent condition. The wing-tip fringe

scales have been knocked off the holotype, and the allotype genitalia has been

mounted on a slide.

DISTRIBUTION (Fig. 44). In Thailand hodgkini is a forest mosquito, al-

though it is infrequently found in shaded plantations and villages. Typically,

it is found in either secondary or primary forest. There seems to be some

correlation of hodgkini with the distribution of tropical wet forests in Thailand;

however, this needs further study. Currently, hodgkini is found in peninsular

Thailand and up to about 15° N latitude in central Thailand. The most northern

collections were made in Khao Yai National Park (Nakhon Ratchasima) and in

the Khwae Noi River valley (Kanchanaburi). This species is encountered most

commonly in the southern peninsular provinces; however, it is also abundant

on Ko Chang (Trat), an island off the southeast coast next to Cambodia. Speci-

mens (660, 599, 122 larvae and 156 larval and pupal skins) in the USMN were

examined from the following provinces of THAILAND: Kanchanaburi, Krabi,

Nakhon Ratchasima, Phangnga, Phuket, Prachin Buri, Ranong, Satun, Song--

khla, Trang and Trat. Additional specimens were examined from MALAYSIA

in the USNM (20, 72 and 8 larval and pupal skins) and in the BMNH (1, 39 and

6 larval and pupal skins - type-series), while 40° and 82 specimens from CAM-

BODIA were examined in the ORSTOM collections.

TAXONOMIC DISCUSSION. Anopheles hodgkini is usually encountered in

forested areas, and should not be difficult to identify because the only other

forest species in this group are barbumbrosus , donaldi and pollicaris. Since

the last 2 species are rare and only occur in the extreme southern provinces

of Thailand, hodgkini is most commonly associated with barbumbrosus, from

which it is easily separated by the apical pale fringe spots on the wing and

normally a few pale scales on the abdominal sterna. The pupa of hodgkini

differs from that of barbumbrosus by the absence of a secondary fold on the

trumpet. If hodgkini and donaldi were found together, the best differentiating

character besides small foretarsal differences, would be the pupal trumpet. An.

donaldi has a deep secondary cleft, which is absent on hodgkini pupae. The

use of the foretarsal bands would become a primary character in separating

pollicaris and hodgkini, while the key characters are best for separating the

pupae (see taxonomic discussion under pollicaris). The presence of a thumb-

like stigmal process on pollicaris larvae will always separate the larvae of

Harrison and Scanlon: Subgenus Anopheles in Thailand 99

these 2 species. Along forest edges and in cultivated plantations, hodgkini will

occasionally be found with barbirostris. In such situations the pupal trumpet

provides the best differentiating character, for like donaldi, barbirostris has

a deep secondary cleft that is absent on hodgkini pupae.

Adults of hodgkini occasionally exhibit character variations which may

influence their identification. Reid (1962) indicated that hodgkini females

always possessed a few pale scales on the abdominal sterna, or at least on

sternum VII; however, Moorhouse (1965) found many hodgkini without pale

sternal scales. A minority of Thai hodgkini are also without pale sternal

scales, but the majority usually exhibit at least 1 - 2 such scales on sternum

VII. The occurrence of the accessory pale fringe spot at vein Ro is also

slightly variable. Three confirmed hodgkini females were found that lacked

this spot on both wings, and were only separable from barbirostris by associ-

ated pupal skins. Infrequent female hodgkini may have the preapical and ac-

cessory pale fringe spots confluent. Approximately 30% of the male hodgkini

show a confluence of these 2 spots.

Pupal characters also exhibit a few variations. Rare specimens of hodgkini

were noted which had a small dark indentation at the spot on the trumpet where

a secondary cleft or fold occurs on other species. Occasional specimens have

a seta 0-VII with 3 branches, but onlyon one, never both of these setae. The

pigmentation of seta 9 was a very useful secondary character for identifying

hodgkini pupae. These setae were dark brown to black on over 95% of the

specimens examined, while the same setae on barbirostris, campestris, don-

aldi and pollicaris are nearly always pale yellow or tan. Such dark setae are

also found on barbumbrosus pupae, but those of the latter on segment VII have

a length 6 - 8 times the width, while on hodgkini they are only 4 - 5 times the

width. Dark setae 9 are very prevalent in the vanus subgroup of the barbiros-

tris group. Both manalangi and vanus usually exhibit dark setae, although

vanus is less consistent than manalangi. Anopheles franciscoi, the only mem-

ber of the barbirostris subgroup in the Philippines, has yellow to tan setae

like the other subgroup members (except hodgkini).

Larvae of hodgkini have few (19 - 44) branches on seta 3-C, another char-

acter similar to the vanus subgroup. When these branches rarely exceed 40,

then the pupal stage is the best way of separating hodgkini from barbirostris

larvae. Anopheles barbumbrosus larvae can occasionally be confused with

those of hodgkini, because the former has seta 3-C with 12 - 36 branches and

infrequently they are more rigid than normal. One hodgkini larva was found

with an anomalous, bifid seta 2-C. Seta 1-II (palmate) on hodgkini is often

pale like those found on barvbumbrosus and other vanus subgroup members.

Members of the barbirostris subgroup (including franciscoi) have seta 1-II

darkly pigmented (exceptfor hodgkini).

As can be seen from the above discussion hodgkini has a number of imma-

ture characters that imply a close relationship to the vanus subgroup. On the

other hand, the pale fringe spots on the wing and pale abdominal sterna scales

found on adult hodgkini coupled with egg characters (Reid 1962), imply a re-

lationship with the barbirostris subgroup. Apparently hodgkini occupies a

somewhat intermediate position between these 2 subgroups.

BIOLOGY. As indicated in the distribution section, hodgkini is essentially

a forest species in Thailand. Much more is known about the biology of the im-

mature stages of this species than the adults. Immature stages have been col-

lected in Thailand from 5 - 750 m elevation. The larvae apparently require

clear (often stained), cool, moderate to heavily shaded water. Floating leaves

are often encountered in the larval habitats. Immatures have been collected in

100 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Thailand from large marshes, large pond, ditch, canals, ground pools, flood

pools, rock pools, stream pools, stream margin, large seepage springs,

large shallow wells, elephant footprints and a large mining pit. The most

frequently encountered habitats with this species were rock pools and stream

pools, particularly in dried up stream beds.

Adult hodgkini are rarely collected. Reid (1962) says that small numbers

of this species were taken in a human bait trap by Macdonald and Traub (1960).

One female in the USNM was collected by Bishop Museum personnel in Trang

Province in 1964, but with no mention of the bait used. SEATO Medical Re-

search Laboratory personnel have spent thousands of man hours making anophe-

line biting collections on both animal and human bait over the last 12 years,

and adult hodgkini have not been collected. Many of these collections were made

in or adjacent to forested areas on peninsular Thailand.

MEDICAL SIGNIFICANCE. No information is available that incriminates

hodgkini as a vector of human pathogens. Attempts to infect hodgkini with

Plasmodium cynomolgi Mayer have involved too few mosquitoes to allow con-

clusions as to their susceptibility (Warren et al. 1963; Bennett et al. 1966).

The lack of information on adult behavior and biting responses after many years

of collecting efforts in Thailand and Malaysia seems to rule out any direct in-

volvement of hodgkini in the transmission of human pathogens.

ANOPHELES (ANOPHELES) POLLICARIS REID

(Figures 46, 47, 48)

Anopheles (An.) pollicaris Reid 1962: 24 (“, 2*, P, L*, E*); Reid 1968: 136

(S, oe, RP, Ls, E*),

This species is fairly easily recognized, particularly in the adult female

and larval stages. Adult females have wide foretarsal pale bands, an accessory

pale fringe spot adjacent to wing vein Ro and often have scattered pale scales

on the costa in the preapical dark area. The larva is typical of the group, ex-

cept it has a unique thumb-like stigmal process. The pupa is fairly easily

separated from the majority of the group members by the absence of a secon-

dary cleft in the trumpet. Essentially this species is like barbirostris except

for:

FEMALE (Fig. 46) Thorax. Pleural setae: 1 - 4 propleural, 5 - 8 spiracu-

lar, 9 - 13 prealar, 3 - 5 upper and 3 - 4 lower sternopleural, 6 - 10 upper

and 0 - 3 lower mesepimeral; and 2 - 6 lower sternopleural white scales (bar-

birostris 5 - 10); 8 - 17 lower mesepimeral white scales (barbirostris 2 - 12).

Wing. Preapical dark mark on costa often with scattered pale scales; tip of

Ro often dark scaled; accessory pale fringe spot adjacent to vein Ro. Legs.

Upper midcoxa with 2 - 3 setae; foretarsi with broad pale bands; 1st pale band

on apex of tarsomere 1 and base of tarsomere 2 approximately 4 - 7 times as

long as wide, usually as long as or longer than length of tarsomere 5 and with

0.33 of band on base of tarsomere 2; 2nd pale band on apex of tarsomere 2

and base of tarsomere 3, length approximately 2 - 4 times width, with 0.25 -

0.33 of band on base of tarsomere 3; 3rd pale band small only on apex of tarso-

mere 3. Midtarsomeres 1 - 2 with narrow apical pale bands or small dorso-

apical pale spots. Abdomen. Sterna with no lateral or diagonal pale scales;

median pale scales from 5 - 17 on sterna II - VI; caudal margin of sternum VI

may have several central black scales.

MALE (Fig. 46) Head. Palpus with distinct narrow pale bands at segmental

Harrison and Scanlon: Subgenus Anopheles in Thailand 101

joints 3 -4and4-5. Wing. Generally paler thanfemale. Legs. Foretarsal

pale bands usually longer than those on female, 1st band approximately equal

to length of foretarsomere 3; tarsomere 3 without apical pale band. Abdomen.

Dorsal surface of 8th segment with scales on median area, pale scales basally,

black scales apically. Genitalia. Aedeagus with 4 - 5 pairs of leaflets; 3

largest leaflets on each side have small denticulate edge, 2nd leaflet usually

with most extensive denticles.

PUPA (Fig. 47) Cephalothorax. Trumpet without secondary cleft or fold.

Abdomen. Seta 0-VII with 2 - 5 branches; seta 9 yellow or tan, 4 - 6 times

as long as wide.

LARVA (Fig. 48) Head. Seta 9-C with 10 - 17 branches (other members of

complex, 5-11). Abdomen. Median dorsal valve with thumb-like stigmal

process; seta 1 on III - VII (palmates) large, dark, with tips pale; 1-II darkly

pigmented.

TYPE-DATA. The 2 holotype, “ allotype and 22 paratypes, with associ-

ated larval and pupal skins (except paratype #961) are located in the BMNH.

All 4 specimens are in excellent condition and come from the same locality

in Selangor, Malaya. The holotype, allotype and one 2 paratype are labeled:

"16th mile, Ulu Gombak, 13-5-1957, J. A. Reid.'' These 3 specimens each

possess a number for associated reared skins, they are: holotype - '0196/11;

allotype - '0196/4"'; and paratype - ''0196/6". The other 2 paratype numbered

"961", has the following data: '16 mi. Ulu Gombak, Bare leg catch-forest. "'

The allotype genitalia were clipped and mounted on a slide for this study. This

slide bears the following SEAMP label; ''SEAMP Acc. No. 334, “ prep. 73/

112,'' and is deposited with the allotype in the BMNH.

DISTRIBUTION. This is another species which enters Thailand only along

the southern border where a Malayan type climate and vegetation are common.

Specimens (20, 22, 3 larvae and 6 larval and pupal skins) in the USNM were

examined from Satun and Songkhla provinces, THAILAND. Additional speci-

mens from Malaysia were examined in the USNM (50, 42 and 15 larval and

pupal skins), and in the BMNH (1c, 32 and 6 larval and pupal skins - type-

series). Apparently pollicaris is a true Malayan species for it has not been

found outside of Malaysia except for southern Thailand.

TAXONOMIC DISCUSSION. Adults of this species are most similar to

donaldi and hodgkini, but can be separated from these by the broad foretarsal

pale bands which cross onto the bases of the next tarsomeres, and pale scales

on the preapical dark area of the costa (when present). This latter character

was noted by Reid (1963) on about 50% of the Malayan specimens, but was not

present on the Thai specimens examined here. The holotype of: pollicaris

does not possess these pale costal scales, but the 22 paratypes in the BMNH

exhibit this character. No variation was noted regarding the accessory pale

fringe spot adjacent to the tip of wing vein R,, but this may be due to the few

specimens available for study. Males should not be confused with any other

members of the group, because of the very wide foretarsal bands. The pupa

of pollicaris is easily separated from all the other Thailand members of the

eroup (except hodgkini) by the absence of a secondary cleft or fold in the trum-

pet. Pupal separation of pollicaris from hodgkini is more difficult, and in

the past has been based entirely on the degree of branching on seta 0-VII. Un-

fortunately, this character is not completely reliable in Thailand or Malaysia,

for hodgkini occasionally has 3 branches on 0-VII and pollicaris infrequently

has 2 branches on 0-VII. Another character has been found which is about

97 - 98% reliable, this involves the color of seta 9. Setae 9 on pollicaris

are yellow or tan, while those on hodgkini are nearly always dark brown or

102 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

black. By using both the seta 0 and seta 9 characters the pupae of pollicaris

and hodgkini can be separated more reliably. The larva of pollicaris can be

separated from the other species in the complex by 2 characters, i.e., the

number of branches on seta 9-C and the presence of a thumb-like stigmal pro-

cess. The latter is by far the better character, but since this process is un-

pigmented, at times it is virtually impossible to see. When this process is

hard to see, the specimen(s) are easily confused with barbirostris larvae, un-

less the seta 9-C character is used.

BIOLOGY. Very little is known about the biology and behavior of this

species, particularly in the adult stage. It appears to be a forest species in

Thailand as it is in Malaysia. The 5 Thailand collections consist of immatures

taken from temporary pools and stream pools along stream courses. These

sites were under light to heavy shade (3 of 5 moderate) and at elevations up to

190 m. These pools contained floating leaves and the water was usually clear,

although in one case polluted due to stagnation. In all cases the stream courses

were located in secondary or primary forest. Other species found as larvae

in these pools were: An. montanus, An. voperi, Uranotaenia spp., Culex spp.,

and in one pool Aedes (Aedimorphus) spp., Aedes (Muscidus) sp. and An. bala-

bacensis Baisas. Nothing was discovered about adult behavior from the Thai-

land collections and apparently very little is known from Malaysia. Reid (1968)

points out small numbers have been collected in the forest biting man. In fact,

one ¢ paratype in the BMNH was collected by this method.

MEDICAL SIGNIFICANCE. Currently, pollicaris is considered an uncom-

mon forest species that is not involved in the transmission of human disease

pathogens.

ANOPHELES (ANOPHELES) BARBUMBROSUS STRICKLAND AND CHOWDHURY

(Figures 49, 50, 51)

Myzorhynchus barbirostris var. pallidus Swellengrebel 1919: 8 (o*); Swellen-

erebel and Swellengrebel-de Graaf 1919: 21 (L*); Swellengrebel and

Swellengrebel-de Graaf 1920b: 82 (L*).

Anopheles barbumbrosus Strickland and Chowdhury 1927: 18 (nom. nov. for

barbirostris var. pallidus Swellengrebel, non pallidus Theobald 1901)

(L*).

Anopheles (Anopheles) barbumbrosus of Edwards 1932: 40 (in part); Christo-

phers 1933: 157 (*); Bonne-Wepster and Swellengrebel 1953: 212 (o*,

as a

Anopheles (Anopheles) barbumbrosus Strickland and Chowdhury, Reid 1962: 29

(OO oD pe SE) Reid 1968! 198: (6%, 2%; “P*, 1 *).

This species is the only member of the vanus subgroup of the barbirostris

group that occurs in Thailand. The adults, pupal and larval stages possess

distinctive characters that make it the most easily recognized member of the

barbirostris group found in Thailand. This species is like barbirostris except

for:

FEMALE (Fig. 49) Thorax. Pleural setae: 3 - 7 propleural, 4 - 7 spira-

cular, 11 - 16 prealar, 5 - 9 upper and 4 - 6 lower sternopleural, 5 - 12 upper

and 0 lower mesepimeral; propleuron without pale scales, lower mese-

pimeron with pale scales. Wing. Normally with humeral pale spot on costa and

scales on humeral crossvein; accessory pale fringe spot adjacent to vein Ro;

lower apical pale fringe spot wide and distinct, extending at least from R4i5

Harrison and Scanlon: Subgenus Anopheles in Thailand 103

to M149; usually with pale fringe spot at Cug. Legs. Upper midcoxa with 3 -

4 setae. Foreleg: Femur with distinct pale spot on inner and outer aspect

near apex; foretarsi dark scaled except small dorsoapical pale spot on tarso-

meres 1 and 2. Midleg: Femur with long distinct pale spot on inner and outer

aspect near apex; midtarsi dark scaled except very small dorsoapical pale spot

on tarsomeres 1 and 2. Hindleg: Tibia with distinct basal and apical pale bands;

tarsomeres 1 - 4 with narrow apical pale bands; tarsomeres 3 - 5 with basal

pale bands, increasing in size distally; basal pale band on T5 equal or wider

than apical pale band on tarsomere 4. Abdomen. Without scales, except ster-

num VII with median tuft of dark scales near caudal margin; sternum VII rarely

with 1 - 2 dirty white scales cephalad of dark scale tuft.

MALE (Fig. 49) Head. Palpus with very narrow pale band at segmental

joints 3 -4and4-5. Wing. Paler than that of female, with distinct humeral

pale spot on costa. Abdomen. Dorsal surface of segment 8 without scales.

Genitalia. Basimeres with pale dorsal and dark lateral scales; tergum IX

with short lateral lobes; aedeagus with 2 - 6 pairs of leaflets; largest pair of

leaflets usually with small indistinct basal tooth, infrequently this tooth is

larger; distal half of largest pair of leaflets distinctly curved, sometimes with

a large lateral denticle near beginning of curve; usually only one pair of large

leaflets and these have variable amounts of small lateral denticles, remaining

leaflets usually small.

PUPA (Fig. 50) Cephalothorax. Trumpet with very small indentation as-

sociated with long seam or fold. Abdomen. Setae 1 and 5 on V - VII with

many branches and tufted; sum of branches on both setae 5-III less than 30;

9 on If - VII dark brown or usually black, 9-VII, 6 - 8 times as long as wide,

9-VIII usually with few lateral branches, infrequently almost spinelike.

LARVA (Fig. 51) Head. Antenna nearly always pale, even when remainder

of head is dark; seta 1-A often shorter, not reaching tip of antenna; 3-C with

12 - 36 thin attenuated branches, usually non-rigid and spread out. Thorax.

Seta 11-P usually stout and split at base into 2 stout branches; 13-P with 7 - 9

branches. Abdomen. Seta 2-I with 3 - 6 branches; 1-II nearly always unpig-

mented or faintly pigmented, rarely heavily pigmented; pecten plate with 9 -

12 pecten teeth, usually 10 - 12.

TYPE-DATA. No types designated. The name pallidus Swellengrebel,

was based on males reared from larvae collected in Mandailing, Sumatra

and the island of Nusa Kambangan (= Noesa Kembangan), South Java. Strick-

land and Chowdhury (1927) failed to designate a type for this species when they

changed the name from pallidus Swellengrebel (not pallidus Theobald 1901) to

barbumbrosus. One ~ is in the BMNH from Nusa Kambangan, the type-locality.

Possibly this is one of Swellengrebel's original specimens, but due to the lack

of adequate characters to separate the adults of this species from those of An.

vanus, it is best left as is until the fauna of Java and Sumatra become better

known.

DISTRIBUTION (Fig. 50). This is another member of the barbirostris

group which should be considered a forest species. The distribution map

does not show barbumbrosus as occurring throughout Thailand, however, this

species should be expected anywhere in Thailand where primary or secondary

forest exists. Actually the range of barvbumbrosus in Thailand is probably

being reduced by the depletion of the forests. There are no specimens avail-

able to show that barvbumbrosus has ever occurred in the broad rice plains

north of Bangkok. Only a few specimens are available from the Korat Plateau,

which also has broad rice plains, and these all come from mountainous or hilly

areas of the plateau. These findings suggest that besides forests, mountain

104 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

or hill streams and pools, clear cool water is another basic requirement. It

will be interesting to note the distribution of this species in Thailand in future

surveys. Specimens (1210, 1589, 304 larvae and 383 larval and pupal skins)

deposited in the USNM were examined from the following provinces of THAI-

LAND: Chanthaburi, Chiang Mai, Chiang Rai, Chon Buri, Chumphon, Kan-

chanaburi, Khon Kaen, Krabi, Lampang, Mae Hong Son, Nakhon Nayok, Nak-

hon Ratchasima, Nan, Narathiwat, Phangnga, Phrae, Prachin Buri, Ranong,

Songkhla and Yala. Other specimens (3c, 1392, 28 larvae and 4 larval and

pupal skins) in the USNM were examined from INDONESIA (Java), MALAYSIA

(Peninsular), SRI LANKA (see taxonomic discussion) and TAIWAN. Two “ and

62 from CAMBODIA were found in the ORSTOM collections. Additional litera-

ture references record this species from INDIA, NEPAL, INDONESIA (Suma-

tra) and SOUTH VIETNAM.

TAXONOMIC DISCUSSION. Adults of barbumbrosus are distinct and easy

to recognize in Thailand. Both sexes possess a lower apical pale fringe spot

on the wing which extends at least from R4+5 to M1+92. This spot is not found

on other Thai members of the barbirostris group. Pale scales are not found

on the abdominal sterna, except a rare 1 - 2 pale scales on sternum VII. All

the other members of this group possess at least a few pale scales on the ab-

dominal sterna except hodgkini which infrequently has no sternal pale scales.

The distal curve on the largest pair of aedeagal leaflets is also usually suffi-

cient to identify rubbed males of barbumbrosus.

Wattal et al. (1962) were the first to record barbumbrosus from India and

Reid (1962) was the first to record it from Nepal and Sri Lanka. Reid (1962)

noted the adults from Nepal and Sri Lanka lacked the accessory pale fringe

spot normally found adjacent to wing vein Ry on barbumbrosus. Adults (1c,

82) recently collected in Sri Lanka by Smithsonian Institution personnel con-

firm this observation. Besides the absence of the accessory fringe spot, these

adults only have narrow apical pale bands on hindtarsomeres 1 - 4, while bar-

bumbrosus from Thailand also has basal bands on tarsomeres 3 - 5 in addition

to the apical bands. Too few immatures are available from Sri Lanka for valid

comparison but those available key to barbumbrosus. Possibly the reduction in

tarsal banding is associated with the clinal band reduction in India and Sri

Lanka previously discussed under nigerrimus and peditaeniatus. The status

of bavbumbrosus in the Indian subregion needs further study.

The pupa is quite distinct, and should not be confused with the other mem-

bers of the group that occur in Thailand. Besides having a secondary seam

on the trumpet rather than a deep cleft, or no cleft, barbumbrosus pupae also

have fewer branches on seta 5-III and have seta 9 darkly pigmented and quite

long on segment VII. Anopheles hodgkini is the only species that could cause

confusion, for it also possesses darkly pigmented seta 9, but these are not as.

long as those on barbumbrosus. Besides, hodgkini pupae do not have a secon-

dary cleft or seam on the trumpet and have more branches on seta 5-III.

Larvae of barbumbrosus can usually be identified by the few non-rigid

branches on seta 3-C, and the pale palmate seta 1-II. These characters do

vary slightly however, and specimens will occasionally be found with the bran-

ches on 3-C stiffer than normal, or with seta 1-II lightly to moderately pig-

mented. Specimens with these variations can easily be confused with hodgkini,

which normally has 19 - 44 stiff branches on seta 3-C and seta 1-II often pale.

In such situations the number of branches on setae 13-P and 2-I may be of

help, but it is better to rear the specimens and identify cast pupal skins and

adults.

Several other variations were observed. Occasional females and more

Harrison and Scanlon: Subgenus Anopheles in Thailand 105

frequently males (5 - 10%) will exhibit a broad pale upper fringe spot on the

wing. This broad spot is due to an extension and joining of the pale preapical

spot and the accessory pale spot at Ro.

Spine-like pupal seta 9-VIII with very few lateral branches were observed

frequently. Specimens from Chiang Mai, Kanchanaburi and Lampang seemed

to possess this variation more frequently than specimens from other areas.

Anomalies were also observed on several larvae. At least 3 larvae from

Lampang were noted with seta 2-C separated by at least 2 widths of a 2-C

alveolus, rather than one width which is normal. One larva from Lampang

was observed with both 2-C forked at midpoint, while another larva from

Chiang Mai had an extra 3-C on the right side.

BIOLOGY. Over 95% of the immature collections of barbumbrosus have

come from primary or secondary wet, deciduous or evergreen forests, or

dense bamboo groves. Only a few collections have been made at the forest

edge or outside a forest. Immature collections have been made from 75 -

1,065 m in elevation, usually in moderate to heavy shade, and normally in

cool clear water sources that contain dead leaves and/or limbs. Approximate-

ly 60% of the larval collections came from stream pools, rock pools, flood

pools or stream margins. However, barbumbrosus larvae have also been

collected in the following habitats: a small pond-marsh on a mountain, stump

hole in ground, pit, rock hole, ground pools, puddles, swamp margin, shal-

low well, seepage spring, rice field with abundant emergent vegetation, wheel

tracks, elephant footprints, hoof prints, large open bamboo internode laying

on the ground, tree hole in a fallen tree, water pot and a water jug.

Very little is known about the biology and behavior of adults of this species.

Scanlon and Esah (1965) reported barbumbrosus females collected biting man

at an elevation between 760 - 1,370 m in northern Thailand. Harinasuta et al.

(1970) reported 11 females collected by the human bait method in Kanchana-

buri Province. Apparently, these are the only confirmed records of barbum-

brosus biting man. One female was captured in a light trap in Chiang Mai, but

other trapping methods have been unproductive.

MEDICAL SIGNIFICANCE. Although seldom encountered, this is a com-

mon forest species in Thailand, at least more common than in Malaysia (Reid

1968). Even so, we still have no information that indicates barbumbrosus is

involved in the transmission of human pathogens. Recently Harinasuta et al.

(1970) reported 1 of 9 dissected females infected with a Dirofilaria larva.

ANOPHELES UMBROSUS SPECIES GROUP

(An. umbrosus species group, Reid 1950)

Myzorhynchus umbrosus (Theobald) 1903b.

GENERAL. Moderately large and dark. Head. Palpus entirely black or

brown scaled (except separatus), with basal scales usually more dense and

erect; palpus equal to or shorter than length of proboscis; clypeus bare; 1st

antennal flagellomere only with patch of dark scales; interocular space with

tuft of long yellow or tan setae and short white scales (except sepavatus); ver-

tex with erect scales only. Thovax. Pleuron without white scale patches and

lower mesepimeral setae; propleuron without scales; scutum with center ashy

gray and sides dark brown on most species. Wing. Basal half of costa without

scattered pale scales; subcostal pale spot small, infrequently absent; preapical

pale spot small; humeral crossvein with dense patch of black scales; remigium

106 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

usually dark scaled; basal 0.33 of vein R entirely dark scaled; base of Cu usu-

ally dark scaled; veins Rg-R2+3 and/or M with pale scales just before forks;

tertiary fringe scales dark (except separatus); posterior margin of wing with-

out pale fringe spot; tip of wing with 1 - 2 small pale fringe spots (bvevirostris,

with a nearly unicolorous dark brown wing, is an exception). Legs. Coxae

with few dark or pale scales; upper midcoxa without scales; femora, tibiae and

tarsomeres mostly dark scaled; tarsomeres may have narrow to moderate

pale bands. Abdomen. No scales on the terga or sterna (see male description).

FEMALE Head. Antenna with reduced whorled setae; pedicel with small

dark scales on upper and outer aspects.

MALE Head. Antenna with long dense whorled setae; pedicel very large,

without scales. Abdomen. Dorsal surface of 8th segment without scales (ex-

cept rvoperi and separatus). Genitalia. Basimeres with at least dark scales

(except brevipalpis); aedeagus with leaflets (except hunteri), which may have

fine lateral denticles; claspette bi-or trilobed, with at least one stout mesal

seta on ventral lobe; dorsal lobe with club-like structure composed of several

separate basal stems that fuse distally to form single club.

PUPA Cephalothorax. Palpal sheath not exhibiting consistently separable

sexual dimorphic characters; male palpus appearing slightly more attenuated

than that of female. Tvumpet. Longest axis transverse to stem, but less so

than in hyrcanus and barbirostris groups; rim of trumpet usually without secon-

dary cleft, but with long, often complex, tragus. Abdomen. Terga with den-

ticles on caudal margins (except baezai and samarensis); seta 1-VII with 5 or

fewer branches; seta 5-VII with not more than 13 branches.

LARVA Head. Seta 2-C frequently branched near tip; seta 3-C with not

more than 25 branches, usually 5 - 20. Abdomen. Without palmate setae or

with 2 to 3 pairs of palmate setae instead of the usual 5 - 6 pairs.

DISTRIBUTION. There are currently 12 species recognized in this group:

baezai, brevipalpis, brevirostris Reid, collessi Reid, huntevi (Strickland),

letifer, voperi, samarensis Rozeboom, separatus, similissimus Strickland

and Chowdhury, umbrosus and whartoni. Their distributions extend from

Assam where vopervi and umbrosus are known, to the Philippines where baezai

and samarensis occur, and Guam where baezai was recently recorded (Basio

and Reisen 1971). The group definitely has its center of distribution in

Malaysia-Indonesia, for 11 of 12 described species occur in Malaysia (Reid

1950, 1963, 1968). The distribution of this group seems to have a definite

correlation with the distribution of tropical wet forests. Accordingly, mem-

bers of this group are rarely, if ever, collected in areas where annual rain-

fall levels are low and the year is divided into very distinct wet and dry sea-

sons. This correlation is very evident in Thailand, where there are 6 members

of the group: baezai, letifer, roperi, sepavatus, umbrosus and whartont.

The only areas where these species are found in Thailand is in the southern

peninsula and/or the southeast corner near Cambodia. Both areas have the

heaviest rainfall in Thailand and tropical wet forests.

TAXONOMIC DISCUSSION. This group is less homogeneous and more

artificial than the hyrcanus and barbirostris groups. Although several assem-

bleges of species in the group are obviously sibling clusters, the group asa

whole cannot be called a sibling group. The letifer subgroup (Reid 1968) in-

cludes collessi, letifer, roperi and whartoni. These species are so similar

that some stages are still unseparable, and those that can be separated are

distinguished by minute differences. Anopheles baezai and samarensis are

also very closely related. The former has a Philippine synonym, gateri

Baisas, which should also be considered when these 2 species are reviewed.

Harrison and Scanlon: Subgenus Anopheles in Thailand 107

Anopheles hunteri and separatus are separable only on poorly known male geni-

talia and larval characters. Anopheles brevirostris and umbrosus have a num-

ber of similarities and are probably more closely related to each other than to

the other members of the group. Nevertheless, they are easily differentiated

in all stages known. The egg of brevirostris is unknown. One of the remain-

ing 2 species, brevipalpis, is distinct and the other, similissimus, is very

poorly known.

The relationship of the umbrosus group to the other species groups in

Southeast Asia has been discussed earlier under the taxonomic discussions of

the albotaeniatus, hyrcanus and barbirostris groups. The closest relationship

seems to lie with several members of the albotaeniatus group, however, spe-

cies such as separatus and huntevi also point to a relationship with the hyrcanus

group. The outcome of phylogenetic interpretations is entirely dependent on

the life stage(s) that the worker feels has the most significant characters

(Reid 1950). Reid (1950, 1963, 1968) selected the larval and pupal stages as

the main indicators of affinities, and for convenience, we concur with this se-

lection.

Six of the 12 known species are not known from Thailand. Anopheles samar-

ensis is known only from the Philippines, and is supposedly separable from

baezat by pale banding on the adult tarsomeres, the size of the tragus on the

pupal trumpet and fewer branches on larval seta 3-C (Rozeboom 1951). There

are Philippine specimens in the USNM of the above stages which exhibit degrees

of development of these characters intermediate between baezai and samarensis.

This suggests that baezai, its synonym gateri and samarensis need to be re-

examined to determine if these are overlapping characters of 2 species or highly

variable characters of a single species. Anopheles similissimus is currently

differentiated only on the basis of larval characters reported in the literature.

No specimens are available, but the species is listed from several areas of Pen-

insular Malaysia (Reid 1968). The larva, if accurately described, is most unique

in having seta 3-C with numerous branches as in the hyrcanus group and seta 1

on the abdominal segments filamentous as in the uwmbrosus group. The single

female reported was listed as indistinguishable from umbrosus. Anopheles

hunteri, from Peninsular Malaysia and the Indonesian islands of Bintan and

South Natuna, is also imperfectly known. This species as pointed out above,

is very similar to sepavatus. The male of hunteri is quite unique in the umbro-

sus group because it does not have aedeagal leaflets. The larva is distinct

from that of separatus by having fewer branches on satae 3-C (6 - 10) and

1-A (less than 20). An. brevipalpis was described from what is now Sabah. It

has since been recorded from Peninsular Malaysia and Sumatra (Banka). Reid

(1968) also mentions its possible occurrence in Thailand, but we have seen no

specimens to confirm this. This species fits well in the uwmbrosus group ex-

cept that the adults are a unicolorous brown. Thus, the adults can easily be

confused with sintonoides and members of the aitkenii group. Anopheles bre-

vipalpis has scales on the anterior pronotum and on the humeral crossvein of

the wing, characters that are absent on sintonoides and species in the aitkeni1

group. The aitkeniz group species also have narrow erect scales on the occiput

while brevipalpis and sintonoides have broad erect occipital scales. Anopheles

brevirostris is a small rare species known only from Peninsular Malaysia. An.

brevirostris is similar to umbrosus in: the propleural setae, the number of

setae on the upper mesepimeron, the pupal trumpet and the weakly developed

palmate setae 1 on 2 of the larval abdominal segments. On the other hand,

brevirostris is distinct because it has: a short proposcis and palpi when com-

pared to the forefemur, broad foretarsal pale bands, short serrate aedeagal

108 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

leaflets, blunt setae 9 on the pupa and no stigmal process on the larva. The one

remaining species, collessi, is known only from Brunei and Malaysia (Peninsu-

lar and Sabah). This species is extremely similar to letiferand whartoni, and

occurs in 2 Malay provinces, (Kedah and Kelantan), that are adjacent to Thai-

land. This species is best separated from letifer and whartoni in the adult,

larval and egg stages. Adults of collessi have the following wing character

combination that differs from both letifer and whartoni: extreme base of vein

Cu with a small pale mark or scattered pale scales, and base of vein 1A above

upper dark mark with 9 - 25 dark scales. The larva of collessi has seta 1-X

bifid, seta 1-A with 15 or more branches and seta 3-C with 5 - 15 (usually 8

or more) branches. Larvae of letifer and whartoni have 1-X simple, 1-A

with 7 - 13 branches and 3-C with 2 - 9 branches. The pupa of collessi can

be separated from that of letifer by having a whartoni type tragus on the trum-

pet, but it cannot be separated from the pupa of whartoni. The reader should

refer to Reid (1968) for information on the distinctive egg of collessi.

BIOLOGY. Larvae of this group are generally confined to still water,

such as freshwater swamps, jungle pools, ditches or drains in plantations,

large isolated stream pools or brackish water pools and swamps. Some seem

to have a definite association with the dark colored acid water that comes from

black peaty soil (Hodgkin 1950). Most are usually found in shaded situations

where there is either emergent vegetation or numerous leaves in the water.

Adults can frequently be found resting in the vegetation or on tree trunks near

larval sites. The females of these species are primarily zoophilic, but will

also attack man during daylight under shaded conditions. Certain species

have anthropophilic tendencies, and these will also enter houses to feed.

MEDICAL SIGNIFICANCE. Of the 12 species in this group, 5 have been

incriminated as vectors of malaria and/or filarial parasites, and those incrim-

inations occurred in Malaysia. These same 5 species also occur in Thailand,

but to date there is no confirmation they serve there as vectors of human patho-

gens.

KEYS. The following keys have been modified from those of Reid (1968).

The keys to the adults and pupae are the most reliable. The larvae, particu-

larly of the letifer subgroup, are so similar that certain species are currently

not separable.

KEYS TO THE SPECIES OF THE AN, UMBROSUS GROUP

IN THAILAND

ADULTS

£. Palpus with pale band, apical segment usually entirely pale scaled.

separatus (p. 128)

Panis Cmireny Car Scaled oi! (oO A, BES POG PEE Da 2

2(1). Base of hindtibia with distinct pale band, usually 1.5 - 4.0 times as

long as segment width; base of Rand Cu usually with several scat-

tered pale scales; accessory pale fringe spot often present at apex

CECI re CIN: REO Gi ler Maha a roperi (p. 124)

Base of hindtibia dark scaled or with small pale spot; base of R and Cu

with dark scales; wing without fringe spot at vein Ro. se ee eee 3

3(2). With propleural setae on both sides (1-6). ............06.:% 4

Without propleural setae (rarely with one small seta on one side). . 5

4(3).

Harrison and Scanlon: Subgenus Anopheles in Thailand 109

Upper mesepimeron with 9 - 19 setae and a few inconspicuous dark

scales; tarsomeres all dark or with minute spots at joints; aedeagal

leaflets short and knife-like. ............ baezai (p. 114)

Upper mesepimeron with 1 - 6 setae and no associated scales; fore-and

hindtarsomeres with small pale bands; aedeagal leaflets long and

thresdlike see PIS ON ee ae Os umbrosus (p. 110)

Vein 1A with pale scales proximal to median dark mark, infrequently

with 2 - 3 dark scales near base. .......... letifer (p. 118)

Vein 1A with 5 or more dark scales at base, infrequently vein nearly

aliviane Genea LCG Oik a te ae ae whartoni (p. 122)

PUPAE

2(1).

3(2).

4(3).

5(4).

Caudal margin of abdominal terga with small denticles. ....... 2

Caudal margin of abdominal terga smooth, without denticles.

baezai (p. 114)

Paddle with refractile border more than 0.75 its length; without pig-

mented projection below the root of trumpet; seta 9-V, 4 - 6 times

as long as wide, tapering to point. ........ umbrosus (p. 110)

Paddle with refractile border less than 0.65 its length; with pigmented

ventrolateral flange of dorsal plate below the root of trumpet; seta

9-V about 2 - 3 times as long as wide, with tip usually more rounded.

Seta 0-IV small with 1 - 2 branches; rim of trumpet with 2 thickened

davieiy plemented arcagi i Vion: aieates KI ae separatus (p. 128)

Seta 0-IV large with 4 - 14 branches; rim of trumpet without thickened

darkly pigmented areas. ...... (letifer subgroup) - 4

Seta 9, on VI - VII, 3.5 - 4.0 times as long as wide, tapering to sharp

point; female genital lobe same color as cercal lobes; sides of male

genital lobes nearly parallel. 6°. 0.4) 08 oe roperi (p. 124)

Seta 9, on VI - VII, 2 - 3 times as long as wide, with more rounded tips;

female genital lobe distinctly more darkly pigmented than cercal

lobes; sides of male genital lobe convergent apically. ....... 9)

Tragus on trumpet complex, nearly T-shaped in outline; seta 2-III with

7 - 17 branches; seta 6-III with 1 - 4 branches. . .. letifer (p. 118)

Tragus more simple and spatulate; seta 2-III with 15 - 23 branches;

seta 6-III with 3 - 7 branches. .......... whartoni (p. 122)

LARVAE

2(1).

Seta 1 with developed leaflets on at least abdominal segments IV - V;

median dorsal valve with stigmal filament. .... umbrosus (p. 110)

Seta 1 with filamentous branches, without leaflets on any abdominal

segment; median dorsal valve without stigmal filament or process.

Seta 6-IV with 7 - 10 branches; seta 2-C usually with 5 or more fine

branches near tip; seta 3-A with truncate tip. .... baezai (p. 114)

Seta 6-IV with 2 - 6 branches, usually 2 - 4; seta 2-C simple or with

up to 4 branches near the tip; setae 2-A and 3-A appear pointed. . 3

110 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

3(2). Seta 1-A with 25 or more branches; branches of seta 4-A fine, not

heckedias, Inge oie Oe ea separatus (p. 128)

Seta 1-A usually with less than 20 branches; seta 4-A with 1 - 2 stout,

Slightly hoeked;branches..; 60: ¢ alae. 4 (letifer subgroup) - 4

4(3).° Seta 0-II] with 5 or more branches. .9) 2.08 20) v6.0 % roperi (p. 124)

Seta O-III usually with less than 5 branches, at least on one side.

letifer (p. 118)*

whartoni (p. 122)*

ANOPHELES (ANOPHELES) UMBROSUS (THEOBALD)

(Figures 52, 53, 54)

Myzorhynchus umbrosus Theobald 1903b: 87 (2*).

Myzorhynchus novumbrosus Strickland 1916: 271 (A, L).

Anopheles (Anopheles) novumbrosus Strickland, Edwards 1932: 42; Colless

1948: 91 (= umbrosus Theobald).

Anopheles (Anopheles) umbrosus of Edwards 1932: 42 (in part); Colless 1948:

91 (o*, OF L*).

and Swellengrebel 1953: 170 (“*, @*, L*); Reid 1968: 156 (*, oP *,

Anopheles (Anopheles) umbrosus Theobald, Reid 1950: 284 (“*, 9*, P*, L*);

Bonne-Wepster and Swellengrebel 1953: 170 (**, @*, L*).

Anopheles (Anopheles) umbrosus (Theobald), Reid 1968: 156 (o*, 2*, P*, L*,

B*);

This species is fairly easily recognized in all stages. The adults have dark

palpi, propleural setae, narrow pale tarsal bands which are usually proximal

to the joints on the hindtarsi and 1 - 6 setae and no scales on the upper mese-

pimeron. In addition, the male has long thread-like aedeagal leaflets. The

pupa has a distinct tragus on the trumpet, denticles on the caudal margin of

the abdominal terga and seta 9-V pointed and longer than those on the other

segments. The larva has at least partially developed palmate setae on IV - V

and a long tapering stigmal process. The egg is also distinctive (Reid 1968).

FEMALE (Fig. 52) Head. Vertex with broad dark erect scales except for

dorsocentral pale scales; interocular space with long tan setae and short nar-

row white scales approximately 0.25 length of setae; palpus with dark erect

scales and tapered appearance because more distal scales are shorter and

decumbent, slightly shorter than proboscis; proboscis slightly shorter to

equal length of forefemur; clypeus bare; pedicel with small blue-black scales

on upper-outer aspects; flagellomere 1 with several broad blue-black scales,

remaining flagellomeres without scales. Thorax. Integument dark brown,

with central portion of scutum ash-gray; anterior promontory with slender pale

scales centrally, darker broader scales laterally; scutum sparsely covered

with short and few longer tan-yellow setae; scutal fossa without scales; scu-

tellum with short and long setae; anterior pronotum with prominent tuft of

blue-black scales on dorsocephalic aspect, short pale setae on ventrocaudal

aspect; pleural setae: 2 - 6 propleural, 4 - 7 spiracular, 2 - 4 upper and 3 -

6 lower sternopleural, 7 - 11 prealar and 1 - 6 upper mesepimeral; propleu-

ron, sternopleuron and mesepimeron without scales. Wing. Costa with black

scales except for very small subcostal pale spot on some specimens and small

*letifer and whartoni larvae currently not separable.

Harrison and Scanlon: Subgenus Anopheles in Thailand Lis

preapical pale spot; subcosta entirely black scaled, or with several apical

pale scales on specimens with subcostal pale spot on costa; remigium with

black scales; humeral crossvein with patch of black scales; vein R-R 1 with dark

scales except tip which has pale scales and is adjacent to preapical pale spot on

costa, and infrequent specimens with several pale scales representing pale

sector spot; Ro-Rg.3 usually with black scales to fork, infrequently with paler

scales just before fork; Ro with black scales, may have pale scales adjacent

to tip of R,, tip with black scales; Rg with black scales except for distinct

patch of pale scales on basal half, tip with black scales; R4+5 with distinct black

scale patch on basal 0.16, remainder with pale scales or mixed pale and dark

scales except tip which has white scales; M with dark scales on basal half and

pale scales just before fork; M;,9 with dark scales at base, remainder with

pale scales; M3,4 with dark scales at least halfway to fork; Cu, with several

dark scales at base, distinct small dark spot at M-Cu crossvein and distinct

small dark spot at apex; Cug with pale scales on basal 0.66, apical 0.33 with

black scales; vein 1A with several black scales at extreme base, then pale

scales out to distinct small black patch of scales slightly distal to midpoint,

followed by pale scales and small dark patch of scales at apex; apical dark

mark on Cug distinctly longer than apical dark mark on vein 1A; apical pale

fringe spot narrow, extending at most from Rg to R4,5; posterior margin of

wing without pale fringe spot; tertiary fringe scales blue-black, like primary

and secondary fringe scales. Halter. Knob with dark scales. Legs. Coxae

without scales, except occasional 1 - 2 blue-black scales on the uppercephalic

aspect of forecoxa; upper midcoxa with 3 - 7 setae. Foreleg: Femur swollen

on basal half, with blue-black scales dorsally and laterally, and slightly paler

scales ventrally; tibia with dark scales dorsally and laterally, slightly paler

scales ventrally; tarsomeres with dark scales, except for narrow apical pale

bands on tarsomeres 1 and 2; tarsomeres 3 and 4 may have several dorsoapi-

cal pale scales. Midleg: Femur and tibia with dark scales dorsally and lateral-

ly, slightly paler scales ventrally; tarsomeres with dark scales except for very

narrow dorsoapical pale scale patch on tarsomeres 1 - 3. Hindleg: Femur

and tibia with dark scales dorsally and laterally, paler scales ventrally, apex

of tibia with very narrow band of pale scales; tarsomeres with dark scales

except for pale band approximately equal segment width on apices of tarso-

meres 1 - 4; tarsomeres 4 and 5 may have very narrow pale basal bands.

Abdomen. Integument brown dorsally and ventrally; without scales.

MALE (Fig. 52). As for female, with dark palps and no scales on abdo-

men. Wing may have more pale scales, and sector spot may be present on

vein R. Genitalia. Basimeres with pale dorsal setae and few dark lateral

scales; inner parabasal spine on very prominent lobe, outer parabasal on

smaller lobe, longer and more tapered than inner, often with 3rd smaller

parabasal spine slightly distal to other 2; internal spine located on mesal

margin approximately midway between apex and claspette; claspette with 2 dis-

tinct lobes; ventral lobe with a single long stout seta, usually one small seta

and small spicules or spines on ventral surface; dorsal lobe with club-like struc-

ture composed of 2 - 3 very closely associated flattened spines; aedeagus with 8 -

11 pairs of long thread-like leaflets, 2 - 3 longest pairs as long as aedeagus;

lobes of tergum IX narrow, 3 - 5 times as long as wide, about half as long as

the space between them.

PUPA (Fig. 53). Integument normally light tan. Cephalothorax. Wing

sheath usually with darker brown barred design or series of small brown spots;

antennal sheath with tip and last joint darkly pigmented; without pigmented lat-

eral angle (projection) below trumpet root. Trumpet. Without secondary cleft,

with narrow complex tragus that is trough-like and abruptly twisted near apex.

112 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Abdomen. Segment VII distinctly narrower than others, often with parallel

sides; caudal margins of terga with distinct denticles; setae 1 and 5 not dark and

bushy; seta 0 small with 2 - 5 branches; seta 2 small with 2 - 9 branches; seta 3-

IV with 12 - 18 branches; seta 9-V pointed, 4 - 6 times as long as wide, longer

than other setae 9 except 9-VII; seta 3-VI between setae 1 and 5-VI; seta 9-VIII

nearly spine-like, with few weak side branches; lateral margin of VIII with sev-

eral small denticles just cephalad of seta 9 base; sides of male genital pouch

nearly parallel. Paddle. Refractile border approximately 0.8 - 0.9 of paddle

length, with distinct spines on apical half; seta 1-P short, stout and single or

frayed near tip.

LARVA (Fig. 54) Head. Capsule light tan; seta 1-A with 13 - 31 branches;

4-A branches fine, filamentous; seta 3-A with slightly frayed tip from lateral

view; 2-C single or with 2 - 6 fine branches near tip; 3-C with 12 - 25 bran-

ches; 4-C single or with 2 - 4 branches; 8-C with 3 - 7 branches. Thorax.

Seta 1-P simple or with 2 - 3 fine branches at tip; 2-P with 6 - 9 branches;

4-P with tuft of fine branches at tip and long lateral branches; 11-P with 5 -

11 densely packed, subequal branches arising from stout basal stem; 14-P

with 8 - 12 branches; branches on 3-T filamentous or slightly flattened, unpig-

mented. Abdomen. Seta 1 usually with filamentous unpigmented branches on

segments I, II, VI - VIII, with partially to fully flattened branches (palmate)

on segments III - V, and 1, on IV - V often darkly pigmented; O-III minute, with

2 - 5 branches; 6-III with more than 20 branches, 6-IV with 2 - 3 branches;

3-VI1 located between setae 1 and 5-VI; segment VIII with ventral plate on ce-

phalic margin; spiracular area with long tapering, filamentous stigmal process;

pecten plate with 7 - 9 long teeth, usually without short alternating teeth; 1-X

weak, single or bifid, arising at edge of saddle.

TYPE-DATA. The 2 holotype is located in the BMNH and is labeled:

(1st label) - ''Pehang Malay, Dr. Durham"'; (2nd label) - ''Myzorhynchus umbro-

sus, type FVT"'; and (3rd label) "Pekan Hospt., Pahang, X.02''. The speci-

men is in poor-fair condition, with the left wing, part of the right antenna, 2

forelegs and the right midleg missing. The structures still present are in

good condition and exhibit the salient species characters.

DISTRIBUTION (Fig. 53). Known from Thailand with certainty only from

Narathiwat Province, near the Malaysian border. This is another Malaysian-

Indonesian species that apparently enters Thailand only where environmental

conditions are right and tropical wet forest occurs. Iyengar (1953), Tansathit

et al. (1962), and Harinasuta et al. (1970) record this species from southern

southeastern and west-central Thailand respectively, but the absence of speci-

mens for confirmation makes it impossible to determine which member(s) of

the umbrosus species group might be involved. One larva from Narathiwat

Province, THAILAND, was found in the SEATO Medical Research Laboratory

collection. Additional specimens (80, 469 and 10 larval and pupal skins) in

the USNM were examined from: INDIA (Assam) and MALAYSIA. An addition-

al 20°, 82 and 7 larval and pupal skins, besides the holotype, were examined

in the BMNH. Reid (1963) examined 2 larvae of vopevi from Assam, and sug-

gested (also Reid 1968) that Christophers' (1933) record of umbrosus from

Assam may actually refer to ropevi. However, a single female in the USNM

confirms Christophers' record. This female was collected by C. O. Masters

in Assam, on 20-XII-1943, and is easily identified as umbrosus by 3 - 4 pro-

pleural setae, absence of scales and only 2 setae on the upper mesepimeron,

and typical umbrosus-like wing and tarsal characters. Anopheles umbrosus is

also known from Indonesia (Reid 1950, 1968), but records from the Andaman

Islands (Christophers 1933) and Nepal (Brydon et al. 1961), need confirmation.

Harrison and Scanlon: Subgenus Anopheles in Thailand 113

TAXONOMIC DISCUSSION. The name "umbrosus" has been applied to

several closely related species in Southeast Asia, particularly in Malaysia.

This has been most unfortunate because of problems associated with the pre-

sumed role of "umbrosus" as a vector of human malaria parasites. Colless

(1948) began to clarify the situation when he synonymized novumbrosus under

umbrosus. However, clarification of all the species masquerading under this

name did not come until Reid's detailed study (1950), supported by the biologi-

cal observations of Hodgkin (1950). Reid (1950) presents a thorough discussion

of the misinterpretations and confusion that existed under the name "'umbrosus"'

prior to 1950. The other umbrosus group members that occur in Thailand

(baezai, letifer, roperi, separatus and whartoni) are easily separated from

umbrosus in all stages. Females of umbrosus even with propleural setae

missing, should be fairly easily recognized by other pleural chaetotaxy, wing

and tarsal characters. Of 41 Malaysian females examined for wing variations,

19 possessed at least 2 - 3 pale subcostal (Scp) scales and pale sector (Sp)

scales, 8 had pale Scp scales but no pale Sp scales, 3 had no pale Scp scales

but had pale Sp scales, and 11 had no pale scales on Scp or Sp areas. The re-

maining wing of the female from Assam exhibited characters equal to the sec-

ond category above. The hindtarsal banding is quite significant, for those pale

bands present on the tarsomeres are mostly apical rather than basal (cf. letifer).

The larva of umbrosus is most distinct because of the absence of distinct

palmate setae on all segments except III - V, and particularly the long taper-

ing stigmal process. The only other known Anopheles larvae having large

stigmal processes in Southeast Asia are: pollicaris (barbirostris group), and

pseudobarbirostris Ludlow and bancrofti barbiventris Brug of the bancrofti

group. Inthe Australasian region, bancrofti bancrofti Giles and atratipes

(?stigmaticus group) also have stigmal processes. These species (except

atratipes) possess palmate setae 1 on at least segments III - VII. In Southeast

Asia pollicavis and umbrosus larvae might be found together, but should not be

confused because pollicaris has more palmate setae and a short thumb-like

stigmal process. The known distribution of umbrosus does not overlap with

those of pseudobarbirostris and bancrofti barbiventris.

The combination of a complex trumpet, extra large seta 9-V and long

refractile paddle border on the pupa of umbrosus, makes it difficult to confuse

the pupa of this species with any other species in Southeast Asia.

The male genitalia of umbrosus is quite distinct because of the extremely

long thread-like aedeagal leaflets. Some of the longest leaflets may approach

1.5 times the length of the aedeagus.

Some of the members of the stigmaticus group in the Australasian region

have thread-like aedeagal leaflets, but they are much shorter than those on

umbrosus.

BIOLOGY. Hodgkin (1950) attempted to sort out the biological require-

ments and habitat preferences for members of the uwmbrosus group in

Malaya, but earlier species misinterpretations made this difficult. He did

outline the basic larval habitat of umbrosus as being the dense swampy jungle

of the coastal plains and foothills, where the water is stained from the peaty

soil. This agrees well with the one Thai larval collection made in a heavily

shaded jungle spring bog with many dead leaves in the water. Apparently, um-

brosus is a swamp-forest species, usually not closely associated with human

populations, which exhibits an opportunistic type feeding behavior. Wharton

et al. (1964) and Moorhouse and Wharton (1965) have shown that adults will

bite in the forest throughout the day, but they exhibit distinct peaks of biting

activity during the hour before sunset and the first hour after dawn. Moor-

114 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

house (1965) determined that adult wmbrosus are reluctant to leave the forest

and enter village clearings, even at night. According to Moorhouse and Whar-

ton (1965), those few adults that do enter village areas during the night bite

outside, not in houses. A pyrethrum knock-down survey conducted by these

authors also failed to yield any umbrosus resting in houses during the day.

However, the adult behavior of umbrosus actually appears more plastic than

described by these last authors. Reid (1961) attracted this species to human

or calf-baited net traps placed under a shelter open on all 4 sides. Reid

(1968), using man as bait, compared simultaneous net trap and window trap

hut collections and captured 39% of the umbrosus by the latter method. He

also compared simultaneous collections of umbrosus biting man outdoors ver-

sus biting man in window trap huts, and captured 14% in the huts (Reid 1968).

The attraction of wmbrosus to feed on different animals has also been mea-

sured. Reid (1961) found 2.5 times as many uwmbrosus were attracted to one

calf as to 2 men. Wharton etal. (1964) found uwmbrosus will bite monkeys on

the ground and on elevated platforms, but the latter only attracted 15% of the

specimens. Furthermore, ground collections comparing man to monkey baits

revealed an attraction ratio of 100: 16, respectively. Additional information

on the biology of wmbrosus can be found in Wharton, Eyles et al. (1963) and

Reid (1968).

MEDICAL SIGNIFICANCE. In Thailand uwmbrosus is probably too scarce

to be involved in the transmission of human pathogens. Until recently umbro-

sus has been considered a confirmed vector of human malaria parasites in

Malaysia. However, Wharton, Eyles et al. (1963) and Wharton et al. (1964)

have shown that the Plasmodium infections commonly found in Malayan umbro-

sus are probably Plasmodium traguli, a parasite of the mouse deer, Tvagulus

javanicus (Osbeck). Subsequent Plasmodium infections found in umbrosus

(Moorhouse 1965, Moorhouse and Wharton 1965) were also credited to P. traguli.

Records of human Plasmodium parasites in uvzbrosus are now considered

questionable due to early confusion of the malaria species involved and the ear-

lier inclusion of baezai, letifer and whartoni under the name "umbrosus".

Reid (1968) still considers umbrosus a potential vector of human malarial

parasites because it readily feeds on man. However, due to its usual remote-

ness from human populations and the absence of any confirmed human malaria

parasites in thousands of dissected adults, this species is probably not a vec-

tor of human malarial parasites.

There have been several studies on the ability of wmbrosus to transmit non-

human Plasmodium species. Warren et al. (1963), found umbrosus moderately

susceptible (19% developing oocysts) to laboratory infection of Plasmodium

cynomolgi bastianellii. Wharton, Eyles et al. (1963) innoculated sporozoites

dissected from umbrosus into 10 species of animals and birds (including man),

but failed to produce malaria infections.

There is very little information about uwmbrosus involvement in filarial

transmission. Wharton, Laing and Cheong (1963) determined that it is non-

susceptible to infection with Wuchereria bancrofti. The only natural filarial

infection found in umbrosus was due to Sefaria larvae, probably from the mouse

deer (Wharton, Eyles et al. 1963).

ANOPHELES (ANOPHELES) BAEZAI GATER

(Figures 55, 56, 57)

Anopheles baezai Gater 1933: 162 (*, 2*, P, L*); Farinaud 1938: 393 (?*, L*).

Harrison and Scanlon: Subgenus Anopheles in Thailand 115

Anopheles (Anopheles) baezai (var. ?) of Russell and Baisas 1936: 27 (o'*, ?*).

Anopheles gateri Baisas 1936: 78 (P*); Reid 1950: 290 (= baezai Gater).

Anopheles (Anopheles) baezai gateri of Colless 1948: 88 (o*, 2*, L*);

Reid 1950, supra cit., (= baezai Gater).

Anopheles (Anopheles) baezai Gater, Reid 1950: 290 (o*, 9*, P*, L*); Bonne-

Wepster and Swellengrebel 1953: 177 (“*, 2*, L*); Reid 1968: 165 (c"*,

*, P*, L*, E*); Basio and Reisen 1971: 60.

Adults of baezai are fairly easily recognized by the combination of propleural

seta, scales on the upper mesepimeron and absence of pale bands on the tarso-

meres. The pupa is very distinct because it lacks denticles on the caudal mar-

gins of the abdominal terga. The larva is easily identified by the absence of

palmate setae on the abdomen, a truncate ventral sabre piece (seta 3-A), many

branches near the tip of seta 2-C, and many branches on seta 6-IV. This

species is like umbrosus except for:

FEMALE (Fig. 55) Thorax. Upper mesepimeron usually with inconspicu-

ous dark scales anteroventral to setae; pleural setae as follows: 1 - 3 pro-

pleural, 2 - 5 spiracular, 3 - 6 upper and 3 - 7 lower sternopleural, 8 - 12

prealar and 9 - 19 upper mesepimeral. Wing. Basal dark mark on vein Cu

not reaching halfway to fork, often with scattered dark scales distal to it; distal

dark mark on vein 1A shorter to nearly equal length of distal dark mark on vein

Cug. Legs. Upper midcoxa with 1 - 5 setae; tarsomeres entirely dark scaled,

infrequently with faint pale spots on the dorsoapical aspect of some segments.

MALE (Fig. 55). Like female, but usually with subcostal pale spot on

costa. Genitalia. Only 2 parabasal spines; ventral lobe of claspette variable,

with single long mesal seta and a variable lateral seta, from very small to

nearly equal to the mesal seta; dorsal lobe with fused club originating from 3

separate basal stems; aedeagus with 4 - 6 pairs of lanceolate leaflets about 0. 33

length of the aedeagus, leaflets without lateral serrations or a basal denticle.

PUPA (Fig. 56) Cephalothorax. Integument light tan; antennal sheath with

all joints and tip the same color as cephalothorax. Trumpet. Tragus variable,

usually short but may be very long like that of wmbrosus or samarensis (see

taxonomic discussion), not usually twisted near tip; secondary cleft or fold,

when present, in form of small fold on rim. Abdomen. Terga without denticles

on the caudal margins; lateral margins of segment VII not parallel; seta 0 small,

often single; seta 2 often with many branches (10 or more); seta 3-IV with 6 - 7

branches; seta 9, II - VI, dark, broad, with rounded tip, not more than 2 times

as long as wide, 9-VIII with strong central stem and long lateral branches;

position of seta 3-VI highly variable, may be lateral, mesal or even with seta

1-VI, but usually slightly more cephalad than seta 1; segment VIII without den-

ticles on the lateral margin above the base of seta 9; male genital pouch with

sides convergent. Paddle. Refractile border 0.5 - 0.7 of paddle length with

strong spines; seta 1-P with 3 - 7 branches on apical half.

LARVA (Fig. 57) Head. Capsule dark brown to black; seta 1-A with less

than 20 branches; seta 3-A with truncate tip; 4-A with filamentous branches;

2-C with 2 - 8 fine branches near tip; 3-C with 10 - 25 branches (but see tax-

onomic discussion). Thorax. 4-P with tuft of fine branches at tip, lateral

branches variable, usually long as on umbrosus, but infrequently shorter and

more sparse; 14-P with 2 - 4 branches; seta 3-T filamentous and unpigmented.

Abdomen. Seta 1 filamentous on all segments; 6-IV with 7 - 10 branches; 3-VI

location variable, from directly under seta 1 to nearly midway between setae

1 and 5; spiracular area without stigmal process; pecten plate with 6 - 7 long

teeth, usually with short alternating teeth; 1-X strong and single, arising at

116 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

edge of saddle.

TYPE-DATA. Gater (1933) listed the type-material as one “, one &, a

larval skin and a whole larva deposited in the BMNH, and the type-locality as

Pulau Langkawi, British Malaya. This is a large island off the west coast of

Peninsular Malaysia, just south of the Thai-Malay border. These specimens

were accessioned at the BMNH as "BM 1934-410", and since Gater did not

designate a holotype they have syntype status. The o, 2 and a larval skin for

the “ are currently in the BMNH. The only whole larva accessioned in ''BM

1934-410" is from Selangor, not Pulau Langkawi; however, there is a 2nd

larval skin from Pulau Langkawi with this accession number. Therefore, the

specimen (= whole larva) that Gater was actually referring to is questionable.

The & specimen is here designated as lectotype for baezai, and is labeled as

follows: (1st label) ''Anopheles (A.) baezai Gater, Det. by B.A.R. Gater"';

(2nd label) "Bred 2464, larvae in pool i stagnant water and decaying leaves,

250 pts Clo per 100,000, Pulo Langkawi"; (3rd label) ''Malay Peninsula, 11.

X . 1928, B.A.R. Gater"; and (4th label) ''Larval skin and hypopygium on

slides, B.M. 1934.410.'' This specimen is in excellent condition and can be

recognized as baezai by the characters presented in the adult key and also by

its larval skin which is mounted on a slide and labeled: ''Department of Biology,

BM 1934.410, Anopheles (A.) baezai Gater, larval pelt 2464, Pulo Langkawi,

11. X. 1928, King Edw. VII College of Medicine, Singapore.'' The genitalia

slide preparation of the lectotype is also in the BMNH.

DISTRIBUTION (Fig. 56). This is a widespread coastal, brackish water

species, that is currently known from Cambodia, Guam, Indonesia, Malaysia,

Philippines, South Vietnam and Thailand. Specimens (3, 3°, 6 larval and

pupal skins) in the USNM were examined from the following provinces of THAI-

LAND: Chanthaburi, Ranong and Trang. Additional specimens (550, 1109,

32 larval and 84 larval and pupal skins) in the USNM were examined from INDO-

NESIA (Sumatra and Java), MALAYSIA, PHILIPPINES and SINGAPORE. Ad-

ditional MALAYSIA and PHILIPPINE specimens (8, 149, 1 larva and 14 larval

and pupal skins) including the type-series, were examined in the BMNH. Six

females from CAMBODIA were found in the ORSTOM collections. Literature

records also include Vietnam (Farinaud 1938) and Guam (Basio and Reisen

1971), but material from these areas was not available for study. The above

distribution for baezai is based on the inclusion of gateri Baisas, as a synonym.

TAXONOMIC DISCUSSION. Anopheles baezai is easy to recognize in Thai-

land. The adults would most likely be confused with wumbrosus, but the pre-

sence of scales and many setae on the upper mesepimeron is very distinct.

The pupa lacks denticles on the caudal margins of the abdominal terga, a char-

acter present on all the other Thailand members of this group. The larva is

also easily identified by the numerous branches on abdominal seta 6-IV and

other previously discussed characters. Considerable variation has been noted

in this species. In Malayan specimens Gater (1933), and Reid (1950) claim the

subcostal pale spot on the costa is uncommon (10% and 33% respectively), while

Colless (1948) claims 80% of Borneo specimens have this spot, as do nearly

all the specimens from the Philippines. Interpretation of this spot is highly

subjective, Reid (1950) plainly defines the spot as ''at least one or two pale

scales breaking the anterior margin of the costa", but the other authors are

not that explicit. Using the above definition and USNM specimens, 44% (11/25)

of the female baezai from Malaysia and 37% (10/27) of the female baezai from

Java have a pale subcostal spot. On the other hand 67% of those Malay females

and 79% of the Java females had pale scales at the apex of the subcostal vein

and on the posterior border of the costa, thus the definition of the pale scales

Harrison and Scanlon: Subgenus Anopheles in Thailand 117

breaking the anterior margin of the costa is very important. Philippine speci-

mens usually possess this spot on the costa and have faint pale spots on the

apices of the tarsomeres. The latter character is less commonly seen in

Malayan specimens. The shape of the 9th tergal lobes of the male was used

by Colless (1948) to differentiate Malaysian and Philippine specimens, but

Reid (1950) felt this was probably intraspecific variation. After examining

specimens from both areas we feel that this latter interpretation is correct.

The length of the tragus on the pupal trumpet is highly variable in this species,

even in a single sibling brood (Reid, personal communication). Specimens

were examined where the tragus graded between practically non-existent to

excessively long and protruding some distance beyond the rim of the trumpet.

The major variations detected on larvae involve the lateral branches of setae

3-C and 4-P. Several USNM larval specimens from Subic Bay, Luzon, Philip-

pines, have only 6 - 9 branches on seta 3-C. On the basis of 3-C counts, these

specimens are intermediate between baezai and samarensis. All larvae

seen from areas other than the Philippines have 3-C with 10 or more branches.

On infrequent specimens, regardless of origin, the lateral branches of 4-P

are more sparse and shorter than usual. On these latter specimens, 4-P

looks more like that of letifer rather than umbrosus. Such variations are

most pertinent when problems arise, such as the status of samarensis in the

Philippines.

BIOLOGY. This species is associated with coastal brackish water habitats.

Usual larval habitats include drains and pools with moderate to heavy shade

that have decaying leaves and sticks in the water. Nipa palms are often associ-

ated with these habitats. In Thailand, larvae have been found only in tin mine

pools, pits or excavations in Ranong Province. Such surface mines are com-

mon in southern Thailand and the water in them is usually acid and without

mosquito larvae. Hodgkin (1950) states that baezai larvae appear to prefer

slightly acid waters with a pH of about 6.3 - 6.8. This would be expected of

a brackish water species. Besides the chloride data listed on the lectotype

label, Colless (1948) lists salinity levels in North Borneo collection sites as

varying between 300 - 1,020 parts chlorine per 100, 000 parts water. Appar-

ently during the evolution of the tin mine pits, the salinity and acidity levels

are compatible for a while with baezai larval requirements. In other countries

baezai larvae have been collected in the following habitats: mangrove swam g,

tidal or coastal swamps, stagnant drains or ditches, ground pools in a salt

marsh and a buffalo (carabao) wallow. Basio and Reisen (1971) gave no data

regarding the pH or salinity of the water in the last habitat.

Certain behavioral aspects of adult baezai are also known. Hodgkin (1950)

commonly found adults resting during the day on the smooth bases of nipa palm

fronds about 1 - 3 feet above the ground. Preliminary data (Hodgkin 1950)

indicated that baezai prefers to feed on man. More recent studies (Reid 1961,

Reid and Weitz 1961) have shown baezai to be strongly zoophilic, with a 1 : 27

man to calf ratio anda 1: 16 man to goat ratio. Although one of the major

sources of blood meals for baezazi in the latter study is still not known, it is

suspected to be from the mouse deer. Wharton, Eyles et al. (1963) collected

baezai in net traps baited with monkeys on the ground, but not on an elevated

platform. Reid (1968), in tests comparing man bait in outside net traps and

window-trap huts, collected only 2% (2/99) of the baezai in the window-trap

huts. These Malayan studies indicate that baezai primarily feeds on cattle

and other mammals rather than man, and that it is reluctant to enter houses.

Only one adult female from Chantaburi Province was collected in Thailand

biting man.

118 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

MEDICAL SIGNIFICANCE. Hodgkin (1950) found a 0.9% sporozoite rate in

baezai from Malaya, and he considered this species a likely vector of human

malaria parasites in that area, although some of the sporozoites did not appear

to be of human origin. Experiments (Hodgkin 1950) to transmit the sporo-

zoites from wild caught baezai to monkeys, and to determine the suscepti-

bility of baezai to infection with human malaria parasites, met with little suc-

cess. Warren et al. (1963) have shown in the laboratory that baezai is re-

fractory to Plasmodium cynomolgi bastianellii. Wharton, Eyles et al. (1963)

infected 1 or 2 baezai with Plasmodium sandoshami Dunn, Eyles and Yap,

from a flying lemur, Cynocephalus variegatus Audebert, and found baezai

had a low susceptibility (23%) to infection with Plasmodium traguli. Current

interpretations (Reid 1968) suggest that baezai is probably not a vector of

human malaria parasites, and based primarily on circumstantial evidence, is

probably a vector of a mouse deer malaria parasite (Wharton et al. 1964).

The involvement of baezai in filarial parasite transmission is poorly known.

No naturally infected baezai were found by Reid et al. (1962), but those authors

were able to infect 34% of exposed mosquitoes in a laboratory with periodic

Brugia malayi.

Current information indicates that baezai is probably not involved in the

transmission of human malaria or filarial parasites, particularly in Thailand

where its population levels are usually low.

ANOPHELES (ANOPHELES) LETIFER SANDOSHAM

(Figures 58, 59, 60)

Anopheles (Anopheles) letifer Sandosham 1944: 129 (9*, L*); Iyengar and Menon

1956: 785; Stone, Knight and Starcke 1959: 21; Reid 1963: 107; Reid 1968:

Lidio*. ee Ps... 1 ee).

Anopheles umbrosus of Stanton 1915: 171 (L*); Puri 1931: 117 (L*); Swellen-

erebel and Rodenwaldt 1932: 97 (in part); Christophers 1933: 162 (o*, 9,

P, L*) (in part); Gater 1934: 104 (L*); Gater 1935: 176 (%, 9*); Reid 1950:

294 (= letifer Gater).

Anopheles (Anopheles) species ''A'' near umbrosus, of Colless 1948: 93 (¢*,

L*); Reid 1950: 294 (= letifer Gater).

Anopheles (Anopheles) letifer Gater, of Reid 1950: 294 (o*, @*, P*, L*);

Bonne-Wepster and Swellengrebel 1953: 180 (¢*, L*).

Best distinguished from the other members of the group by adult female

and pupal characters. The female is characterized by the absence of pro-

pleural setae, base of wing vein Cu all dark, base of vein 1A all pale or rarely

with 1 - 5 dark scales near origin, foretarsomeres without distinct pale bands

and hindtarsomeres primarily with basal pale bands. The pupa is distinct on

the basis of a T-shaped tragus, but this structure is difficult to observe.

The larva is nearly identical to the larva of voperi and cannot always be dif-

ferentiated. Anopheles letifer is like umbrosus except for:

FEMALE (Fig. 58) Head. Palpus and proboscis with more decumbent

scales, appearing thinner; posterior hard palate of cibarium with small median

heavily sclerotized area immediately anterior to ventral papillae. Thorax.

Scutal integument gray, with faint dark longitudinal lines; pleural setae: 0

(rarely 1) propleural, 2 - 7 spiracular, 3 - 6 upper and 2 - 6 lower sterno-

pleural, 7 - 15 prealar and 7 - 10 upper mesepimeral. Wing. Scales on veins

narrower and decumbent, thus veins appear thinner; costa nearly always with

Harrison and Scanlon: Subgenus Anopheles in Thailand 119

small subcostal pale spot; R-R, with small sector pale spot; preapical pale

spot usually completely across Rg; basal dark mark on M usually reaching

about halfway to fork; basal dark mark on Cu rarely reaching more than half-

way to fork; 1A with pale scales proximal to median dark mark, infrequently

with 2 - 3 dark basal scales; distal dark mark on 1A slightly less or equal to

length of distal dark mark on Cug. Legs. Upper midcoxa with 3 - 6 setae; fore-

and midtarsomeres unbanded or with faint apical pale spots; hindtarsomeres

with pale bands about 1 - 3 times as long as wide at joints 1-2, 2-3, 3-4 and

4-5, major portion of each band on bases of tarsomeres 2, 3, 4 and 5 (cf.

umbrosus).

MALE (Fig. 58). Like female, but subcostal pale spot on costa and sec-

tor pale spot on vein R more distinct. Genitalia. Basimeres without obvious

pale dorsal setae; only 2 parabasal setae; ventral lobe of claspette with single

long mesal seta, and 1 - 3 small lateral setae near base; median ventral mem-

brane of claspette covered with thorn-like spines; dorsal lobe of claspette with

fused club originating from 3 separate basal stems, 1 stem internal to the

others; aedeagus with 95 - 6 pairs of lanceolate leaflets about 0.33 length of the

aedeagus, leaflets without lateral serrations, but usually with a small basal

denticle; lobes of tergum IX short and wide, approximately 1.5 times as long

as wide and slightly less than 0.5 as long as the space between them.

PUPA (Fig. 59) Cephalothorax. With pigmented angulate projections below

the root of the trumpet and on the wing sheath; antennal sheath with all joints

and tip the same color as the majority of the cephalothorax. Trumpet. With

complex tragus which is bent inward near the tip and nearly T-shaped; distal

half of pinna with generally narrower diameter than that of umbrosus and

baezai. Abdomen. Lateral margins of segment VII not parallel; seta 0, on II -

VIII large, with 3 or more branches, 0-IV with 4 - 14 branches, nearly as

large as 2-IV; 2, on II - VII large, with 5 or more branches; 2-III with 7 - 17

branches (cf. 15 - 23 whartoni); 6-II with 1 - 4 branches (cf. 3 - 7 whartoni);

9, on II - VII, 2 - 3 times as long as wide, with rounded tip, 9-VIII nearly fan-

like, with small central stem and long lateral and apical branches; caudal mar-

gin of sternum VIII with denticles just internal to 9-VIII; female genital lobe

(ventral) distinctly more darkly pigmented than cercal lobes; sides of the male

genital lobes sharply convergent. Paddle. Refractile border 0.50 - 0.65 of

paddle length; seta 1-P long slender and single or with 2 - 4 fine distal branches;

apex of paddle usually with 1 - 2 small lobes just mesad of 1-P alveolus.

LARVA (Fig. 60) Head. Integument brown, but not as dark as that of baezai;

seta 1-A with 7 - 13 branches; 3-A pointed; 4-A with 1 - 2 thickened branches

that are hooked apically; 2-C usually single; 3-C with 4 - 9 branches; 4-C with

1-3 branches. Thorax. 4-P with tuft of equal length branches at tip, lateral

branches short and sparse; 11-P with 5 - 7 branches; 14-P with 7 - 9 branches;

3-T with filamentous branches. Abdomen. Seta 1 with filamentous branches

on all abdominal segments; O-III minute, with 3 - 4 branches; 6-III with 13 -

23 branches (see taxonomic discussion), 6-IV with 2 - 6 branches; spiracular

area without stigmal process; pecten plate with 7 - 10 long teeth, usually alter-

nating with short teeth; 1-X single, arising at edge of saddle.

TYPE-DATA. Reid (1950) discussed the unusual circumstances surround-

ing the original description of letifer. Sandosham (1944) referred to and de-

scribed letifer thinking that Gater had already published the name and deposited

type-specimens in the BMNH. However, Gater's manuscript and specimens

were destroyed during World War II while en route to England. Due to the

above confusion in naming this species, no type-specimens were selected until

Reid (1950) designated a neotype and deposited it in the BMNH. This specimen

120 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

was examined by the authors and is in excellent condition except for the loss

of the fringe scales on the tip of the right wing. The neotype has the following

labels: (1st label) - "Selangor, Kg. Sijangkang, 16.5.1934, E.P. Hodgkin";

(2nd label) - ''No 7835/12, larvae in stagnant peaty water in garden"; (3rd

label) - "Anopheles (A.) letifer Gater, Neotype, Det. J. Reid''; and (4th label) -

"Liason pelts in slide collection."' The larval and pupal skins of the neotype

are also in the BMNH and are in excellent condition. Both skins are mounted

on the same slide with the following label data: (1st label) - ''Federated Malay

States, Anopheles (A.) umbrosus Theo: letifer Gater. det. J. Reid, Neotype

2, No 7835/12, Institute for Medical Research"; and (2nd label) - ''Adult in

pinned collection.'' There are also 3 $ and one “ which Reid placed in the

BMNH and labeled plesiotypes. The “ and one § of these latter specimens

have associated immature skins.

DISTRIBUTION (Fig. 59). Hodgkin (1950) lists letifer as primarily a

coastal plain species, associated with peaty soil and acid fresh water in

Malaya. Data from the specimens examined here support these associations,

particularly the former. Anopheles letifer has been reported from Thailand

by Iyengar and Menon (1956), Reid (1963) and Scanlon et al. (1968). No speci-

mens are now available to confirm these records, but we are including letifer

here because we feel certain it does occur in Thailand. Reid's (1963) record

is very important because it was made at the same time he described collessi

and whartoni, 2 species very closely related to lefifer. His record was

based on 2 females collected biting man by Dr. Udaya Sandhinand in Sungai

Kolok (Sungei Golok in Reid), Narathiwat Province, very near the Thai-Malay

border. Scanlon et al. (1968) record specimens from Chanthaburi, Chon Buri,

Prachin Buri, Ranong, Trang and Yala provinces. The specimens listed

from Chanthaburi, Prachin Buri and Yala have not been found, and a close

examination revealed those from Ranong are voperi and those from Chon Buri

and Trang are whartoni. Reid (1968) lists the distribution of letifer as penin-

sular Thailand, Malaysia, Sumatra and Borneo. Specimens (100, 289, 3 lar-

vae and 8 larval and pupal skins) in the USNM were examined from CAMBODIA,

INDONESIA (Kalimantan) and MALAYSIA (Peninsular). Additional specimens

(7%, 142, and 6 larval and pupal skins), including the neotype, from MALAY-

SIA (Peninsular and Sarawak) and SINGAPORE were examined in the BMNH.

Additional specimens (2% and 19) from CAMBODIA were found in the ORSTOM

collections. The specimens of letifer from Cambodia appear identical with

Malaysian specimens and lend support to our claim that letifer occurs in Thai-

land, and to a report of letifer in South Vietnam (Nguyen-Thuong-Hien 1968).

TAXONOMIC DISCUSSION. Anopheles letifer should be fairly easy to

identify in Thailand. The adults look like umbrosus, but do not have propleural

setae (rarely one) and pale bands on the foretarsomeres, while the pale bands

on the hindtarsomeres are primarily basal (cf. umbrosus). The male has

short aedeagal leaflets like those of baezai, but the lobes of tergum IX are

short and wide, while those of baezai are normally long and slender. The

ereatest problem in identifying the adults will be differentiating letifer from

whartoni, which also occurs in Thailand. At this time the characters descri-

bed by Reid (1963) to separate letifer and whartoni seem reliable, the -speci-

mens from Cambodia are clearly separable, as are those from Malaysia.

However, the differences separating these 2 species are quite small and will

probably need re-evaluation when more material becomes available. Anophe-

les roperi adults have also been misidentified as letifer in Thailand. Nor-

mally roperi adults are easily recognized by the basal pale band on the hind-

tibia and pale scales on the remigium and basal 0.33 of vein R, but speci-

Harrison and Scanlon: Subgenus Anopheles in Thailand 121

mens that have the legs missing or are rubbed can be confusing. In such

cases voperi can usually still be separated from letifer by having several pale

scales at the base of vein Cu, on the basal 0.33 of vein M, and several narrow

pale scales in the fossal area on the scutum.

The T-shaped tragus on the pupal trumpet of letzfer is distinct, but this

character is often bent and twisted during mounting, and difficult to interpret.

The best means to identify the pupa of this species is by the characters pre-

sented in the key. Only poorly mounted pupal skins of letifer and whartoni

should present any identification problems.

The larva of letifer is much more difficult to identify, and is best identified

by association with later stages. Separation from the larvae of umbrosus,

baezai and separatus presents no problem using the above key characters, but

final differentiation of whole larvae from those of voperi may not always be

possible. Reid (1950, 1968) described seta 6-III on letifer as having 12 - 20

branches and used this as the primary character for separating voperi from

letifer and whartoni. An examination of the neotype and plesiotype skins of

_ letifer in the BMNH and other available associated skins revealed that 5 speci-

mens (including the neotype and 2 plesiotypes) out of 13, have 21 or more

branches on this seta on at least one side. All the specimens of voperi have

21 or more branches on 6-III (including the holotype larval skin). This overlap

means that seta 6-III has little or no value in separating letifer from voperi.

The remaining character (seta 0-III) used by Reid remains valid, but is diffi-

cult to see. Larvae of letifer and whartoni cannot be differentiated at present.

BIOLOGY. The absence of Thai letifer specimens in museum collections

is probably a reflection of the limited areas in Thailand with suitable letifer

habitat and the few collections made in those habitats. Hodgkin (1950) noted

that while wmbrosus larvae are found in jungle swamps, letifer larvae are

usually found at their edge, or in water collections remaining after the swamps

have been cleared or drained. An. letifer is apparently a species of the coastal

plains in Malaysia and its larvae seem to prefer acid freshwater and be intoler-

ant of even low salinity (Hodgkin 1950). Moorhouse (1965) usually found letifer

larvae in brown peaty waters in well shaded ditches between rubber trees, at

the edge of jungle swamps, or in clear water in wells and ditches with algae

and in grassy swamps. Such habitats usually have cool still water and often

have dead leaves or sticks in the water. Although Thailand has extensive rubber

plantations, freshwater swamps and the peaty soil made from such swamps are

not common.

A number of observations have been made on the behavior of adult letifer.

Studies by Wharton, Eyles et al. (1963) show it bites mainly at night, and has

a very low level of biting activity during the day, even in the forest (cf. um-

brosus). The peak period of biting activity in the forest and in settlements is

approximately 2000 - 2400 hours. Most adults entering settlements do so soon

after dusk and rest on tall grass, shrubs or under houses prior to feeding

(Moorhouse 1965). Even though letifer readily enters settlements, it seems

more reluctant to enter houses. Studies (Wharton, Eyles et al. 1963; Reid

1968) comparing manbiting catches of letifer indoors versus outdoors indicate

it is primarily exophagic, with 4 - 9 times as many biting man outside as in-

side. A pyrethrum knockdown study by Moorhouse and Wharton (1965) found

that letifer does not rest in houses during the day. The attraction of this spe-

cies to different hosts has yielded some confusing information. Hodgkin (1950)

considered letifer primarily anthropophilic, but more recent studies (Reid

1961; Wharton, Eyles et al. 1963) obtained more zoophilic and ornithophilic

feeding patterns. Another study (Reid 1968) comparing man with monkeys

122 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

shows that 3 times as many /etifer fed on man, but of those feeding on monkeys,

43% fed in the canopy. Some of the differences found in past studies of letifer

feeding behavior may be a reflection of the changing prosperity of the villagers,

and the associated increase in the number of fowl and animals found in the vil-

lages (Moorhouse 1965).

MEDICAL SIGNIFICANCE. Hodgkin (1950) suggested that in Malaya letifer

was probably the most important malaria vector in the umbrosus complex.

This has been partially supported by evidence showing that umbrosus malaria

infections are of non-human origin (see discussion under umbrosus), and

studies by Wharton et al. (1963) and Wharton, Eyles et al. (1963) that confirm

letifer's participation in the transmission of human malaria parasites. There

is also evidence (Moorhouse 1965) that letifer is not a vector in other areas

where human malaria is common, which suggests genetic or behavioral differ-

ences. Anopheles letifer is a natural vector of Plasmodium traguli, and lab-

oratory studies show that it is a very efficient host for this parasite (Wharton,

Eyles et al. 1963). Laboratory studies by Warren et al. (1963) and Bennett

et al. (1966) indicate that letifer is not a good host for the simian parasite,

Plasmodium cynomolgi bastianellii.

Wharton, Laing and Cheong (1963) and Wharton, Eyles et al. (1963) also

found natural infections of Wuchereria bancrofti and Dirofilaria spp. in letifer,

and experimental transmission tests show a high percentage of letifer develop

W. bancrofti larvae to the infective 3rd stage.

Anopheles letifer seems to be a rare or uncommon species in Thailand,

and probably is not involved in the transmission of human parasites, but if

areas are found in Thailand where it is common, its vector capabilities in

Malaya should be remembered.

ANOPHELES (ANOPHELES) WHARTONI REID

(Figures 61, 62, 63)

Anopheles letifer of Wharton 1960: 81; Reid 1963: 106 (= whartoni).

Anopheles (Anopheles) whartoni Reid 1963: L0G (oO, -o*, P* LE): iia rics

Laing and Cheong 1963: 236; Reid 1968: 180 (¢, ¢*, px, ii, E).

This species is very closely related to letifer and can be recognized only

in the adult and pupal stages. The adults are characterized by having dark

scales on the basal half of wing vein 1A. The pupa has a spatulate tragus and

2 minor setal differences from letifer. Anopheles whartoni is nearly identical

_to letifer except for:

FEMALE (Fig. 61) Head. Sclerotized area of the cibarium less pronounced.

Thorax. Pleural setae: 0 propleural, 3 - 5 spiracular, 3 - 4 upper and 2 - 3

lower sternopleural, 6 - 9 prealar, and 8 - 11 upper mesepimeral. Wing.

Dark, even paler scales dull; subcostal pale spot on costa often very small or

absent; preapical (apical of Reid) pale mark often not complete on vein Ro;

R-Ry rarely with pale sector spot; vein M with dark scales to or nearly to

fork; Cu with dark scales more than halfway to fork, often dark scales to fork;

1A with 5 or more dark scales on basal half, infrequently with dark scales

nearly to median dark mark. Legs. Upper midcoxa with 4 - 5 setae.

MALE (Fig. 61). Likefemale. Genitalia. (Based on one specimen).

Aedeagus with 4 pairs of leaflets; lobes of tergum IX 2.5 - 3.0 times as long

as wide, approximately half as long as space between them.

PUPA (Fig. 62) Trumpet. Tragus spatulate, not T-shaped at apex. Abdo-

Harrison and Scanlon: Subgenus Anopheles in Thailand 123

men. Seta 2-III with 15 - 23 branches; 6-III with 3 - 7 branches. Paddle.

1-P single or with 2 - 3 apical branches.

LARVA (Fig. 63). Currently indistinguishable from that of letifer.

TYPE-DATA. The holotype 2 and allotype “ are deposited, with associ-

ated larval and pupal skins, in the BMNH. Both were reared from eggs de-

posited by a 2 collected at Singora, Maran, Pahang, Malaya.

DISTRIBUTION (Fig. 62). Reid (1968) stated that whartoni is known only

from the eastern side of Peninsular Malaysia (Pahang and Kelantan). Speci-

mens examined by the authors now extend this range to include Thailand and

Cambodia. Specimens (32 and 1 pupal skin) in the USNM were examined from

the following provinces of THAILAND: Chon Buri, Trang and Trat. Addition-

al specimens (20, 52, and 4 larval and pupal skins) in the BMNH and ORSTOM

collections were examined from CAMBODIA and MALAYSIA (Peninsular Malay-

sia). The distribution of whartoni is apparently confined to lowlands where it

largely replaces letifer (Reid 1968). The specimens examined from Thailand

and Cambodia also came from areas of low elevation.

TAXONOMIC DISCUSSION. The occurrence of this species in Thailand has

been responsible for some previous records of letifer. These 2 species are

nearly identical, and their differentiation is very difficult. The slight differ-

ences described by Reid for separating these species are easily overlooked.

Wing vein 1A on whartoni has at least 5 basal dark scales; however, infrequent

specimens of letifery may have 1 - 3 dark basal scales on 1A. Such slight dif-

ferences currently appear valid, but may need modification when more materi-

al becomes available. In general, whavtoni has the darkest wings of any spe-

cies in the umbrosus group (except brevipalpis). Even areas that appear pale

are dull and usually a dirty-white. Some specimens of whartoni have dark

scales on veins M and Cu out to or nearly to the forks, and the base of 1A may

have dark scales nearly out to the median dark mark. The female from Trat

Province has the latter degree of dark scales, and is comparable with speci-

mens seen from nearby in Cambodia. On the other hand, the specimen from

Trang is much paler like letifer and has only 7 - 8 dark scales on the base of

1A, and veins M and Cu dark only 0.66 of the distance to the forks. Although

such differences could be suggestive of clinal variation, specimens from Ma-

laysia were examined that were nearly as dark as those from Cambodia and

eastern Thailand.

The possibility that collessi may occur in Thailand must also be considered,

because it has been collected in Kelantan, Malaysia. This species would also

be difficult to differentiate. Anopheles collessi has dark scales on the basal

half of vein 1A like whartoni, but can be separated from both letifer and whar-

toni by the presence of pale scales on the base of vein Cu. The presence of this

character on collessi makes it more likely to be confused with voperi (see dis-

cussion under vopevi).

The pupa of whartoni can usually be distinguished from that of letifer by

the characters given in the key and description. However, there are currently

no distinctive characters known that will separate the pupa of collessi from

that of whartoni.

The larvae of letifer and whartoni are currently indistinguishable, while

that of collessi is usually separable by having a branched seta 1-X and more

branches on seta 3-C.

BIOLOGY. Very little is known about the biology and behavior of this

species (Wharton 1960, as letifer). Larvae were collected by Wharton ina

variety of sites that ranged from deep shaded swamps with dark peaty water to

clear water habitats such as seepage pools and open grass swamps. The pH

124 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

range for the habitats fell between 5.8 - 6.6. The single immature collection

from Thailand came from an island (Ko Chang) next to Cambodia. This collec-

tion was made in a heavily shaded swamp at 5 m elevation, with deeply stained

water and abundant brown algae. Other larvae found in this collection include

Anopheles crawfordi, An. roperi, Uvanotaenia annandalei Barraud, U. bimacu-

liala Leicester and U. longirostris Leicester.

Adults have been collected biting humans (Malaysia and Thailand) and in

human bait net traps (Malaysia). Wharton (1960) found adults present through-

out the year in a riverine lowland area of Pahang, but abundant only during a

2 month period (April - May) following heavy rains. Females bit throughout

the night, apparently without regard to physiological age, but the peak biting

period was the 2 hours after sunset (1800 - 2000 hr). In comparative tests be-

tween bare leg catches on man outside houses and inside houses, only 3% of

the females were caught indoors. Blood engorged females were also collected

resting indoors before dawn, but pyrethrum knockdown surveys in houses during

the daytime did not recover whartoni specimens. These data suggest that

whartoni in comparison to letifer, prefers to feed earlier in the evening and is

less likely to enter houses to bite man.

MEDICAL SIGNIFICANCE. Since whartoni is relatively poorly known, only

limited information (Wharton 1960) is available regarding its vector capabilities.

This species was found naturally infected with sporozoites and oocysts of pre-

sumed human malaria by Wharton in 1959, and specimens were also found na-

turally infected with Wuchereria bancrofti larvae in 1958 - 1959. Divrofilaria

larvae were also found in whartoni during this period. Laboratory experiments

by Wharton show that whartoni is a good host for W. bancrofti. Anopheles

whartoni is probably responsible for maintaining low levels of malaria and

filarial endemicity in the area studied by Wharton. The role of whartoni in

the transmission of human pathogens in Thailand is not currently known.

ANOPHELES (ANOPHELES) ROPERI REID

(Figures 64, 65, 66)

Anopheles umbrosus, largest variety of Roper 1914: 142 (2); Reid 1950: 298

(= voperi).

Anopheles (Anopheles) sp. B near umbrosus of Colless 1948: 95 (2*); Reid 1950:

298 (= roperi).

Anopheles (Anopheles) roperi Reid 1950: 298 (“*, 9*, P*, L*); Bonne-Wepster

and Swellengrebel 1953: 184 (o*, 9*, L*); Reid 1963: 107 (2*); Reid 1968:

183 (o*, 9*, P*, L*, E*),

A large species that is fairly easily distinguished in the adult and pupal

stages, but not always separable in the larval stage. Adults usually have an

accessory pale fringe spot at Ro, pale scales on the base of veins R and Cu

and on the remigium, and a pale basal band on the hindtibia. Adult males can

also be separated by the long slender aedeagal leaflets. The pupa has denti-

cles on the caudal margin of abdominal terga, a short refractile border on the

paddle, seta 0-IV large with 4 or more branches, seta 9, on VI - VII, 3.5 -

4.0 times as long as wide, the female genital lobes unicolorous with the cer-

cal lobes and the sides of the male genital lobes nearly parallel. The larva is

nearly identical with those of letifer and whartoni, and generally differs only

by having more branches on abdominal seta 0. Anopheles roperi is like leti-

fer except for:

Harrison and Scanlon: Subgenus Anopheles in Thailand 125

FEMALE (Fig. 64) Head. Proboscis and palpus with more erect (shaggy)

scales on basal portions; cibarium with less pronounced sclerotized area.

Thorax. Scutal integument with mottled gray pattern sometimes intermixed

with dark patches; anterior 0.5 of scutum, particularly the lateral fossae, with

scattered narrow curved pale scales; pleural setae: 0 - 3 (usually 0) propleur-

al, 5-10 spiracular, 7 - 17 upper and 3 - 7 lower sternopleural, 11 - 17 pre-

alar and 8 - 14 upper mesepimeral. Wing. Scales on veins broader; costa

may be dark scaled to subcostal spot or may have some pale scales proximal

to the humeral crossvein; subcostal spot may extend onto vein R-R,; costa

with small preapical pale spot just before and down to tip of Rj; remigium may

have area of pale scales; basal half of R usually with several scattered pale

scales and distinct sector pale spot; Ry with pale scales on tip; Rg-Rgi3 with

dark scales at base, then pale scales to fork, except for distinct median dark

mark; preapical pale spot on costa and Ry usually complete on Rg; Ro with

pale or dark scales at tip, depending on presence or absence of accessory

pale fringe spot; M with distinct patch of dark scales at base, then mixed pale

and dark scales out to fork; Cu usually with 1 - 5 pale scales at base, often

more, followed by dark mark 0.2 - 0.3 the length of the distance to the fork,

remainder to fork mixed pale and dark scales or with dark mark on distal half

before fork; 1A with mixed pale and dark scales proximal to median dark mark;

some specimens with narrow accessory pale apical fringe spot adjacent to the

tip of Rg. (For more on wing see taxonomic discussion.) Legs. Coxae with

scales; upper midcoxa with 5 - 12 setae; fore- and midtarsomeres dark

scaled, or with faint dorsoapical pale spots on T1-7T3; hindtibia with pale basal

band 1.5 - 4.0 times segment width; hindtarsomeres 1 - 4 with narrow apical

pale bands, 3 - 5 with narrow basal pale bands, basal pale bands on tarsomeres

4 and 5 longer than apical pale bands on tarsomeres 3 and 4.

MALE (Fig. 64). May have dorsoapical pale spots on foretarsomeres 1 -

3; dorsal surface of 8th abdominal segment with narrow pale scales or setae

dorsally. Genitalia. Basimeres with pale dorsal scales and dark lateral

scales; ventral lobe of claspette with 1 - 3 large setae, most mesal seta usually

largest; dorsal lobe of claspette with fused club originating from 3 separate

basal stems, usually with single knife-like seta separate from club, but im-

mediately adjacent to it on mesal side and difficult to distinguish from club;

aedeagus with 3 - 5 pairs of long, narrow curved leaflets, nearly 0.95 length

of aedeagus; leaflets without lateral denticles, but largest with small basal

tooth and tiny spine-like tip; lobes of tergum IX narrow, approximately 4 - 5

times as long as wide and 0.3 - 0.5 as long as distance between them.

PUPA (Fig. 65) Cephalothorax. Antennal sheath with tip often more darkly

pigmented than remainder of cephalothorax. Trumpet. Tragus long, broad

and spatulate. Abdomen. Seta 9, on VI - VII, 3.5 - 4.0 times as long as

wide, tapering to sharp point; female genital lobe unicolorous with cercal

lobes; male genital lobes with sides much less convergent than those of letifer,

nearly parallel. Paddle. Basal half of seta 1-P stout, apical half with brush-

like tip composed of 4 - 5 nearly equal length branches.

LARVA (Fig. 66). Separable from letifer only by seta 0-III having 5-7 ©

branches (cf. letifer 3 - 4); seta 6-III with 21 - 26 branches, but see taxonomic

discussion under letifer.

TYPE-DATA. The 2 holotype of rvoperi is deposited in the BMNH, and is

in excellent condition except for the loss of the tibia and tarsomeres of the

right hindleg. Label data for the holotype is as follows: (1st label) - "Selangor,

Kuala Kubu, 4.IV.1941, J. A. Reid"; (2nd label) - "No. 16942/54, larvae in

large pool with dead leaves in Jungle"; (3rd label) - "Anopheles (A.) roperi

126 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Reid, TYPE, Det. J. Reid''; (4th label) - ''Holotype''; and (5th label) ''liason

pelts in slide collection."' The “ allotype (No. 16942/52, with associated im-

mature skins) and a “ paratype are also deposted in the BMNH. Both speci-

mens are in excellent condition and come from the same locality as the holo-

type, but the paratype has no associated immature skins or a collection number.

The allotype “ and paratype © still have their genitalia intact, and no male

genitalia preparations for vropeyi were found in the BMNH. The slides contain-

ing the immature skins of the holotype and allotype are present in the BMNH

collection.

DISTRIBUTION (Fig. 65). Anopheles roperi was reported widely distri-

buted by Reid (1968), with records from India (Assam), Indonesia (Sumatra),

Malaysia and Thailand. A single specimen examined by the authors extends

this range to include Cambodia. In Thailand there appears to be some cor-

relation between the pattern of distribution of vopervi and that of tropical wet

forests. This pattern may also be found in the other countries with this species.

Specimens (7%, 102, 21 larvae and 20 larval and pupal skins) in the USNM

were examined from the following provinces of THAILAND: Krabi, Narathiwat,

Phangnga, Phuket, Prachin Buri, Ranong, Satun, Songkhla, Trang and Trat.

Additional specimens (4, 72, 15 larvae, and 4 larval and pupal skins) in the

USNM were examined from MALAYSIA. The single female from CAMBODIA

was found in the ORSTOM collections. The identification of this specimen as

voperi is supported by a large collection of roperi with associated skins from

Trat Province, Thailand, which is adjacent to Cambodia. Specimens (2, 48,

2 larvae and 6 larval and pupal skins) in the BMNH that include the type-series,

were examined from: INDIA (Assam) and MALAYSIA (East and Peninsular).

The 2 larvae are the basis of the Assam record (Reid 1963). We concur with

Reid, these larvae agree with voperi as currently known. Both larvae have

the following label data: (ist label) ''Cinnamara, Jarahaut, Assam, Dr. D.

Manson"; and (2nd label) "Anopheles umbrosus ? roperi J. R. det., Malaria

Survey of India.'' Reid (1963, 1968) suggested that Christophers' (1933) record

of umbrosus in Assam may have been based on specimens of voperi; however,

umbrosus is now definitely known from Assam (see distribution section under

umbrosus).

TAXONOMIC DISCUSSION. Adults of voperi from Malaysia as defined by

Reid's descriptions (1950, 1968), are easy to distinguish from the other mem-

bers of the group. In Thailand, however, most adults do not strictly conform

to Reid's descriptions, and these specimens are more difficult to identify and

are easy to confuse with some of the other species, particularly letifer and

whartoni. Malay specimens typically exhibit a mottled gray scutal pattern,

a pale accessory fringe spot at vein Ro, pale scales on the costa proximal to

the humeral crossvein and pale scales on the remigium and base of veins R

and Cu. Most Thai specimens have a less mottled pattern on the scutum like

letifer, and rarely have pale scales on the remigium and base of the costa.

Only 80% of the Thai females have one or more pale scales on the base of vein

R, and only 70% have one or more pale scales on the vein Cu. In addition,

only one of the 3 Thai females with wing fringe scales intact, exhibited the

accessory pale fringe spot at vein Ro. Some of these wing characters were

also variable on the type-specimens. The 2 holotype only has a very small

pale spot at the extreme base of the costa, while the “ allotype has nearly

all of the costa pale scaled proximal to the humeral crossvein. The “ para-

type has this portion of the costa with 2 pale marks divided by a dark mark.

Although the Thai specimens described above appear quite distinct,

they conform very nicely to other vopeyvi characters such as: 2 pale marks

Harrison and Scanlon: Subgenus Anopheles of Thailand 127

on R9+3 separated by a dark mark; Cu with a short basal dark mark and scat-

tered dark scales distal to it; 1A with scattered dark scales on the basal half;

numerous upper sternopleural setae; coxae with pale scales; numerous upper

midcoxal setae; and hindtibia with basal pale band.

There is also considerable variation in the male genitalia, particularly on

the ventral lobe of the claspette. This lobe is described by Reid as having 3

setae (spines), but we have specimens with 1, 2 or 3 setae, and these include

Malayan (Negeri Sembilan) males.

Adult vopevi, even with the above variations, should be fairly easy to

recognize by the basal pale band on the hindtibia, numerous upper sterno-

pleural and upper midcoxal setae, broader wing scales, scattered dark scales

on veins Cu and 1A and the few pale scales that are usually present on the base

of veins R and Cu.

Only minor variations were detected in the pupal stage. The most signifi-

cant of these being the sides of the male genital lobes may be more convergent

than those depicted by Reid (1968: 186). Even then, however, they are not as

convergent as those of letifer. This life stage of roperi should also be easy to

identify. The small denticles on the caudal margin of the abdominal terga,

pigmented angulate projection below the root of the trumpet and the large many-

branched seta 0 immediately places vopeyvi in the lefzfer subgroup. The Key

characters to separate voperi from the other members of the letifer subgroup

are also quite obvious.

The larva of vopevi is much more difficult to recognize and may not be

discernable without other associated stages. There is no difficulty in recog-

nizing vopevi larvae as belonging to the letifer subgroup, but identity below that

level is extremely difficult. Only one character (seta O-III) is currently known

that will differentiate roperi larvae from those of letifer and whartont, and it is

extremely difficult to see. The validity of the seta 6-III character used by Reid

(1962, 1968) is discussed under letifer.

Based on the specimens examined we feel that the differences noted between

the majority of Thailand roperi specimens (also some from Malaysia) and Reid's

descriptions of vopevi, are probably infraspecific. There is certainly not

enough evidence at this point, or specimens with associated immature skins

to intimate that another undescribed species is involved in what we are calling

roperi. However, that possibility does exist and should be evaluated when

more material, particularly associated immature stages, becomes available.

The fact that the genitalia of the “ allotype are still intact and available for

examination will be helpful in this regard.

BIOLOGY. Reid (1968) reports that ropevi is found in Peninsular Malaysia

from the coastal plains up to about 300 m elevation in the hills, but is most

numerous in wooded shaded valleys where the foothills merge into the plains.

Larvae have been collected in shaded pools, swamps and ditches in Malaysia.

Immatures have been collected in Thailand between 5 - 200 m elevation and

from the following habitats: stream margins, running ditch, nipa palm swamp

and stream and flood pools. Over 80% (17/21) of the Thai immature collec-

tions came from stream or flood pools. The water is generally cool and

unpolluted in these habitats and may be clear or colored. Over 95% of the

habitats were in moderately to heavily shaded secondary or primary wet for-

ests.

The adults of voperi are forest mosquitoes that bite man during the entire

24 hour period, but have the highest level of biting activity during the day,

with a major biting peak in the hour just before dark (Moorhouse and Wharton

1965). In Peninsular Malaysia, Wharton, Eyles et al. (1963) did not collect

128 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

ropervi biting outside the forest, or in or near houses, but a more recent study

(Macdonald et al. 1967) in Sarawak captured several females biting man in open

rice fields near the forest in the evening. Wharton, Eyles et al. (1963) also

determined that vopevi will feed more readily on monkeys on the ground than in

the canopy, and Wharton et al. (1964) indicate that less than 10% of the roperi

captured were taken in the canopy.

MEDICAL SIGNIFICANCE. Hodgkin (1950) presented epidemiological evi-

dence that suggested vopevi was a vector of human malaria parasites in Malaya,

but this has not been confirmed by natural sporozoite infections (Wharton,

Eyles et al. 1963). Anopheles roperi has been found with heavy oocyst and

sporozoite infections (Wharton, Eyles et al. 1963), but these were apparently

from a mouse deer malaria. Sporozoites from wild caught vopevi innoculated

by the latter authors into rhesus monkeys and one man, failed to produce in-

fections. Further experiments during that study show that vopevz is susceptible

to infection with the mouse deer malaria, Plasmodium tvaguli. An attempt

(Warren et al. 1963) to experimentally infect roperi with Plasmodium cynomolgi

bastianellii failed, but Bennett et al. (1966) were able to demonstrate oocyst

development of a Cambodian strain of P. cynomolgi in rope7t.

The only natural infection of filarial larvae in vopervi is that of Sefavia spp.

reported by Wharton, Eyles et al. (1963). Experimentally, Wharton, Laing

and Cheong (1963) showed that Wuchereria bancrofti could develop to mature

3rd stage larvae in vroperi.

There seems little doubt that vopervi is a vector of a mouse deer malaria

in Malaysia; however, there is no proof that it serves as a vector of human

malaria or filarial parasites in Malaysia or Thailand.

ANOPHELES (ANOPHELES) SEPARATUS (LEICESTER)

(Figures 67, 68, 69)

Myzorhynchus separatus Leicester 1908: 36 (2).

Patagiamyia separatus Leicester, James and Stanton 1912: 61.

Myzorhynchus sinensis var. snijdersi Swellengrebel 1921: 114 (“); Christo-

phers 1924: 31 (= separatus).

Anopheles (Anopheles) sepavatus Leicester, Christophers 1924: 30; Senevet

1930: 361 (P*); Edwards 1932: 42, (in part); Gater 1934: 102 (L*), (in

part); Gater 1935: 174 (o“*, 9*), (in part); Crawford 1938: 63 (P*); Col-

less 1948: 87 (o*, 9*, L*); Reid 1950: 306 ("*, 9, P*, L), (in part);

Bonne-Wepster and Swellengrebel 1953: 195 ("*, 9*, L*), (in part).

Anopheles (Myzomyia) separatus Leicester, Brug 1928: 925 (L).

Anopheles hyrcanus separatus Leicester, Swellengrebel and Rodenwaldt 1932:

69.

Anopheles (Anopheles) sepavatus (Leicester), Stone, Knight and Starcke 1959:

28; Reid 1968; 187 (O*)-oF; | PL, E*).

A distinctive species with the adult habitus superficially resembling mem-

bers of the hyrcanus species group. Adults of both sexes have pale scales on

the palps and more distinctive pale areas on the wings. However, both have

only dark scales on the base of veins R and Cu, and the female is without clype-

al scales, or a scale tuft on abdominal sternum VII. The pupa is also easily

separated by having denticles on the caudal margins of the abdominal terga, a

Harrison and Scanlon: Subgenus Anopheles in Thailand 129

short refractile border on the paddle, a pigmented projection below the root of

the trumpet; the rim of the trumpet with 2 thickened, darkly pigmented areas

and seta 0-IV small with 1 - 2 branches. The larva superficially looks like

umbrosus, but has abdominal seta 1 without developed leaflets, median dorsal

valve without a stigmal process, seta 6-IV with 6 or less branches, seta 1-A

with 25 or more branches; and no stout hooked branches on seta 4-A. The

species is like umbrosus except for:

FEMALE (Fig. 67) Head. Interocular space with short pale ocular scales

and long silky white setae forming obvious frontal tuft; palpus with basal scales

erect, but not appearing shaggy, usually with terminal segment entirely pale

and narrow ring of pale scales on apex of segment 3 and base of segment 4

(segment 4 may be entirely pale scaled), pedicel with small blue-white light-

refractive scales on upper and outer aspect; flagellomere 1 with several broad

pale scales. Thorax. Integument light to dark brown with central portion of

scutum mottled gray with longitudinal dark lines; pleural setae: 2 - 4 propleural,

1 - 4 spiracular, 2 - 4 upper and 2 - 5 lower sternopleural, 6 - 8 prealar and

2 - dD upper mesepimeral. Wing. With pale areas larger and more distinct than

on other members of group; costa with black scales, except for small subcos-

tal pale spot which also crosses onto vein R, and small preapical pale spot;

vein R-Ry with dark scales except for well defined sector and subcostal pale

spots, and pale scales at tip; R-R, may have several scattered pale scales

between sector and subcostal pale spots; Rg-R2g+3 with small basal dark

spot, then usually pale to fork; Ro with dark scales except for pale spot con-

tinuous with preapical pale spot on costa and Rj, tip with dark scales; Rg with

pale scales except for small basal dark spot and dark scales on tip; R4i5 with

small dark spot at base and usually at tip, remainder with pale scales; M with

basal 0.16 - 0.33 dark scaled, remainder with pale scales to fork; Mj49 with

dark scales at base and tip, remainder pale scaled; Mgi4 pale scaled except

for dark scales at tip; Cu with dark scales on basal 0.33, remainder pale

scaled to fork; Cuo with dark scales on apical 0.25 - 0.33; base of 1A entirely

pale scaled out to small median dark spot, followed by pale scales and small

apical dark spot at tip; apical pale fringe spot extending at most from Ro9-R4i5;

tertiary fringe scales usually paler than primary and secondary fringe scales.

Legs. Upper midcoxa with 1 - 5 setae; tarsomeres 1 - 4 on fore-, mid- and

hindlegs with narrow apical pale bands or dorsoapical pale spots; hindtarso-

mere 5 infrequently with several apical pale scales.

MALE (Fig. 67) Head. Palpus with variable amount of pale scales; club

entirely pale scaled or with narrow apical dark band on segment 4 or base of

segment 5; segment 3 usually entirely pale on median aspect, may have outer

aspect dark scaled, and narrow apical dark band; segment 2 with few mesal

pale scales. Legs. Foretarsal pale bands may cross joints onto base of tarso-

meres 2 and 3; hindtarsomere 95 often with pale scales. Abdomen. Dorsal sur-

face of 8th segment with numerous median pale scales. Genitalia. Basimeres

with pale scales dorsally, dark scales laterally, and 2 parabasal spines; clas-

pette with 3 lobes, ventral lobe with 2 - 3 large setae and 1 small seta, middle

lobe without setae, dorsal lobe with club-like structure formed from 3 - 4

separate basal stems, the outermost stems appear fused apically, but the

innermost stem seems flattened and separate from the remainder; aedeagus

with 4 - 5 pairs of leaflets about 0.33 as long as aedeagus, larger leaflets

broad with large serrations on outer edge, smaller leaflets without lateral

serrations, but may be forked apically; lobes of tergum IX narrow, approxi-

mately 4 - 5 times as long as wide, over half as long as space between them.

PUPA (Fig. 68) Cephalothovax. With pigmented projection below root of

130 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

trumpet. TJvumpet. Tragus broad and spatulate with saw-tooth-like denticles

on rim; rim of trumpet with 2 thickened, darkly pigmented areas, one forming

part of small secondary cleft and seam. Abdomen. Segment VII not signifi-

cantly narrower than other segments, and sides not parallel; caudal margins

of terga with small denticles; segments with darkly pigmented spots, alveoli of

seta 2 well marked by dark integumental spot on each tergum; seta 1 usually

with few branches or single, but setae 5 on III - VI with many branches and

bushy appearance; seta 0 single or double, but longer than seta 0 on umbrosus;

seta 2 large with 3 - 20 branches; seta 3-IV with 3 - 8 branches (cf. umbrosus

12 - 18); setae 9, on II - VII pale, short with rounded tip, approximately 2.0 -

2.9 times as long as wide; seta 9-VIII with strong central stem and long lateral

and apical branches; segment VIII with minute spicules on lateral margin above

seta 9, and on caudal margin of sternum just mesal to seta 9; female genital

lobe more darkly pigmented than cercal lobes; male genital lobes darkly pig-

mented, with moderately convergent sides. Paddle. Base darkly pigmented;

refractile border approximately 0.55 - 0.60 of length, with minute spines;

1-P long, stout and usually single, but may have 2 - 3 minute branches near

tip.

LARVA (Fig. 69) Head. Capsule and basal 0. 66 of antenna usually darkly

pigmented; basal 0.66 of antenna with many short spines; seta 1-A bushy, with

17 - 38 branches usually 25 or more; seta 3-A with thin, nearly transparent

frayed tip.and lateral margin from lateral view, but usually appears pointed;

4-A without stout hooked branches; seta 2-C usually single; 3-C with 7 - 18

branches; 4-C single, short; 8-C with 2 - 5 branches, usually 3 - 4. Thorax.

1-P with 2 - 6 fine apical branches, approximately half as long as single 3-P;

2-P very large, with 9 - 14 branches; 4-P with tuft of fine equal branches at

tip, short lateral branches (cf. umbvosus); 11-P with 2 - 7 fine branches from

stout stem. Abdomen. Seta 1 filamentous on all segments; O-III minute, single;

6-III with 15 - 22 branches, usually less than 20; spiracular area without stig-

mal process; pecten plate with 5 - 7 long teeth, usually alternating with short

teeth; 1-X approximately as long as saddle, arising on saddle well dorsad of

ventral saddle margin.

TYPE-DATA. According to James and Stanton (1912) Leicester's type-

female for separatus was lost en route to the BMNH. Those authors studied

additional Leicester specimens at the Institute for Medical Research in Kuala

Lumpur, and presumably sent some to the BMNH to represent Leicester's

species. These specimens have been considered part of Leicester's type-

series (Reid 1950, 1968). Three specimens received in the BMNH under the

accession number 1912-350, possibly qualify for this status. All 3 are identi-

fiable with separatus as defined by Reid (1950, 1968). One © and one 2? have

labels that read ''Fed. Malay States, Dr. G. F. Leicester, 1912-350". That

2 also has a smaller label bearing ''A. 4"', possibly referring to a rearing

number. The 3rd specimen, a &, has the following labels: (1st label) (upper

side) - '"Myzomyia Dubitans M.S." (under side) - "Own bungalow, Kuala Lum-

pur, 17/11/03"; and (2nd label) - ''Kuala Lumpur, Fed. Malay States, Dr. G.

F. Leicester, 1912-350". This latter specimen was noted by Christophers

(1924) and Reid (1950) as probably belonging to the type-series. Stone (person-

al communication) also noted the 3 possible types, but said he could not be

sure of their status. Although Leicester (1908) described only the adult @ of

separatus in the original description, he plainly says under the subheading

"occurrence" (p. 37), "bred from larvae taken in large collections of water

in the open near Kuala Lumpur." If this statement applies to the entire

type-series, the adult 2 labeled Myzomyia Dubitans and collected in his bunga-

Harrison and Scanlon: Subgenus Azopheles in Thailand 131

low, cannot be considered a member of the type-series. The absence of asso-

ciated larval skins in the BMNH for the other 2 specimens does not help clari-

_fy their status. No type-specimens are known for the synonym snijdersi which

was described from specimens collected in Medan, on the east coast of Sumatra.

DISTRIBUTION (Fig. 68). This is a coastal plain species which is now

known from Cambodia, Indonesia, Malaysia, South Vietnam and Thailand. Brug

and Bonne-Wepster (1947) list separatus from Buru Island in the Moluccas, but

this needs confirmation. Specimens (1%, 22, 25 larvae and 3 larval and pupal

skins) in the USNM were examined from the following provinces of THAILAND:

Narathiwat and Trang. Additional specimens (10°, 379, 4 larvae and 10 larval

and pupal skins) in the USNM were examined from CAMBODIA, INDONESIA

(Sumatra), MALAYSIA and SOUTH VIETNAM. Specimens (6c, 139 and 9 lar-

val and pupal skins) in the BMNH were examined from MALAYSIA. Included

in these latter specimens is the supposed type-series. An additional 132 speci-

mens from CAMBODIA were found in the ORSTOM collections.

TAXONOMIC DISCUSSION. Adults of separatus are easily misidentified as

members of the hyrcanus group because they possess the following hyrcanus-

like characters: palpus with pale bands, pale frontal tuft, paler wing pattern,

tertiary fringe scales on wing usually paler than primary and secondary fringe

scales, male sternum VIII with numerous median pale scales, and male aedea-

eal leaflets short with large serrations out to tip. Certain other adult charac-

ters, however, clearly indicate this species belongs to the umbrosus group,

they are: antenna with scales on the pedicel and flagellomere I only; base of

wing veins R and Cu with dark scales only, preapical pale wing fringe spot

(= upper apical of Reid) is adjacent to tip of R,, while in the hyrycanus group

this spot ends prior to the tip of Ry (except crawfordi), pleuron without white

scale patches; upper midcoxa without pale scales, female without clypeal

scales; and female without scale tuft on sternum VII.

The immature stages (including the egg) exhibit characters that clearly

indicate this species is a member of the wmbrosus group. The pupa has a

prominent tragus on the trumpet, a pigmented projection below the root of the

trumpet and denticles on the caudal margins of the abdominal terga. The lar-

va has less than 25 branches on seta 3-C, numerous branches on seta 1-A and

filamentous branches on abdominal seta 1. Both the pupa and larva are easily

separated from other members of the umbrosus group by the characters

described in the keys and the above description.

BIOLOGY. Colless (1948) describes the larvae of sepavatus in Borneo as

commonly found among tall grasses in open swamps away from the jungle,

yet rarely collected where only small amounts of vegetation occur, thus show-

ing a marked preference for heavily shaded situations. Despite this preference,

Colless found no separatus larvae in jungle pools. Hodgkin (1950) specifically

defined separatus larval requirements in Malaya, as extending from the inner

fringe of the brackish water zone inland to approximately 100 m elevation in

the foothills. He states that separatus is most common on the coastal plain in

pools under light shade, and generally found in more acid (average pH 5. 8)

water. Hodgkin also claims the larvae do not appear to occur in jungle. Speci-

mens examined during this study indicate immatures collected in Malaysia

came from a mining pool, stagnant drains, a ditch and an artificial container

(large drum), while those from Thailand came from spring fed jungle bogs,

open ''swamp" and a nipa palm swamp. Most of the Thailand collections came

from partially open areas, but several came from heavily shaded jungle bogs

with numerous leaves and sticks in the water. Biting collections have been

made from man, cattle, goats and monkeys. Additional collections have been

132 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

made with net traps and light traps. Adults readily bite man in the absence of

cattle or other animals (Colless 1948; Hodgkin 1950), but when one calf was

compared to 2 men, separatus preferred the calf by a 12: 1 ratio, and pre-

ferred 2 goats to 2 men by a 62:1 ratio. Wharton, Eyles et al. (1963) found

sepavatus would feed on monkeys on the ground or in the canopy, but to a les-

ser extent than it fed on human bait offered at the same time. Reid (1968) in

comparative tests using human bait indoors and outdoors, collected only 2%

of the sepavatus indoors.

MEDICAL SIGNIFICANCE. Based on the above zoophilic behavior data

and a rather limited distribution, separatus is probably not involved in the

transmission of human parasites in Thailand. Hodgkin (1950) reported only

one female naturally infected with malaria parasites, and failed to infect this

species with human malaria parasites by experimental means. Warren et al.

(1963) and Bennett et al. (1966) had only limited success in experimentally in-

fecting this species with various strains of Plasmodium cynomolgi. Wharton,

Eyles et al. (1963), however, found separatus susceptible to experimental in-

fection with Plasmodium traguli. Only one unidentified filarial larva has been

found in wild collected separatus (Wharton 1960), and Wharton, Laing and

Cheong (1963) were only able to obtain one mature Wuchereria bancrofti larva

from experimentally infected separatus.

ANGUSTICORN SECTION

Reid and Knight 1961: 475.

This section was proposed for those species possessing a simple, tubular

type trumpet, having the longest axis vertical and more or less in line with

the stem of the trumpet, and with a single cleft or notch of varying depth on

one side. Certain species of this section (e.g. gigas) have the trumpet pinna

broadly expanded rather than tubular, however, all the species covered herein

have the simple tubular type trumpet. For further information on the anophe-

line trumpet types refer to Reid and Knight (1961).

LOPHOSCELOMYIA SERIES

Edwards 1932: 37.

Members of this small series may be recognized by the following combina-

tion of characters.

ADULT (General). Head with erect broad scales on vertex; scutum

pale centrally, dark laterally; anterior pronotum with scales; pleural sclerites

darkly pigmented; coxae very pale; fore-and midcoxae with long curved (plume)

scales projecting caudoventrally; basal half of forefemur slightly swollen; hind-

femur with apical half swollen, with prominent tuft of scales (black proximally,

white distally) on apical 0.25 - 0.33; tarsomeres without pale bands; wings

with pale scaled areas; abdominal tergum VIII usually with scales. Female.

Palpus with slightly shaggy scales. Male. Dorsal lobe of claspette without

club, with compact group of separate flattened setae; aedeagus without

leaflets.

PUPA. Seta 1, on V - VII much smaller than seta 5 and with few branches;

seta 9, V - VII either frayed or with branches, or with distinctly recurved tip.

LARVA. Seta 1-A small, single or with several branches; 3-C single or

with few branches; 4-C single or with branches on distal half; 5, 6, 7-C short

Harrison and Scanlon: Subgenus Anopheles in Thailand 133

with few branches or single; 11-C longer than antenna, single or with few bran-

ches on distal half; 11-P very small, single or branched; seta 6 long on seg-

ments 1-VI.

DISTRIBUTION. This small series is restricted to the Oriental Region,

and is known from India and Sri Lanka across to Indo-China and south to Indo-

nesia. There are only 3 species recognized in Thailand. One of these, bulk-

leyi, is recorded only from southeast Thailand, and the other 2, asiaticus and

interruptus, are widely distributed but rarely collected in Thailand.

TAXONOMIC DISCUSSION. This series currently contains 5 species, of

which annandalei Prashad, asiaticus, interruptus and noniae Reid belong to

the asiaticus species group. The 5th species, bulkleyi, was placed in this

series by Reid and Knight (1961), and is separated from the members of the

asiaticus group only by differences in the scale tufts on the legs.

The asiaticus group can be divided into 2 subgroups of 2 species, each

possessing characters that will differentiate it from the other subgroup (see

key). The 1st subgroup contains annandalei and interruptus which are appar-

ently separable only by a single larval character (Reid 1968), while the 2nd

subgroup contains asiaticus and noniae which have distinctive characters in

several stages. Since annandalei and noniae are not covered later, both de-

. serve further comment.

Anopheles annandalei was recorded (Reid 1963) from northeast India and

Java (= synonym var. djajasanensis), and Reid (1968) added Thailand to this

list. This latter distribution record was apparently based on correspondence

from Dr. E. I. Coher, who claimed to have found both annandalei and inter-

vuptus in Thailand (Reid 1968: 206). Neither specimens of annandalei nor inter-

ruptus have been found in the Coher and Beales material from Thailand in the

USNM, and since no other specimens from Thailand are known that conform to

annandalei, we consider the Thai record of annandalei doubtful and needing

confirmation. Possibly the Coher identification of annandalei was based ona

wing character which was previously considered valid (Puri 1929), but has

since been found on interruptus in Malaya (Reid 1963).

The record of annandalei sensu stricto, from Java came about when Puri

(1929) synonymized annandalei var. djajasanensis Brug, under annandalei

because he claimed the larvae of both have simple thoracic pleural setae.

Based on available specimens, Reid (1963, 1968) agreed with this synonymy.

Our examination of the holotype larva and male and female paratypes of djaja-

sanensis in the BMNH reveals that Puri (1929) and Reid (1963) were probably

wrong in considering this Indonesian taxon as a synonym of annandalei. How-

ever, this discrepancy is only part of the annandalei-djajasanensis- interruptus

problem. This problem cannot be resolved until more Indian and Indonesian

specimens are available for study. Consequently, we have decided to leave

djajasanensis in synonymy with annandalei until a regional study of this series

can be undertaken.

Anopheles noniae was described by Reid (1963) based on Malaysian speci-

mens. This species is very closely related to asiaticus, but can be differenti-

ated in the adult, pupal and larval stages (see taxonomic discussion under

asiaticus). The immatures of noniae only occur in internodes of living upright

bamboo (Gigantochloa scortechinii Gamble) with bore holes from the chrysome-

lid beetle, Anisodera goryi Baly. Anopheles noniae is not presently recorded

from Thailand, but species that occur in such hidden habitats often remain

unknown for long periods. Workers in Thailand should be on the watch for

noniae.

MEDICAL SIGNIFICANCE. None of the species in the Lophoscelomyia

134 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

series are known to be capable of or involved in the transmission of human

pathogens.

KEYS TO SPECIES OF THE LOPHOSCELOMYIA SERIES

IN THAILAND

ADULTS (bulkleyi from original description).

1. Legs with clusters of erect dark scales on tarsomere 2 of midleg and

tarsomeres 1 and 2 of hind leg. ...°. 2k ow ae bulkleyi (p. 134)

Legs with erect scales only on hindfemur, as prominent tuft of scales

(black proximally, white distally) on apical 0.25 - 0.33.

asiaticus sp. group (p. 135)... .°.:2

2(1). Abdominal tergum VIII with pale golden scales; wing without apical pale

fringe spot at R4,5; subcostal pale spot of costa extending onto vein

Bei ee ee ae asiaticus (p. 136)

Abdominal tergum VIII without pale scales; wing with narrow apical

pale fringe spot at R4,5; subcostal pale spot when present, confined

to cOSta Bnd tip of subeosta: . .... 5. sss interruptus (p. 140)

IMMA TURES (asiaticus group only, bulkleyi unknown).

1, Pupa: Setae 9, on VI - VII frayed or weakly branched; internal border of

paddle without fringe hairs. Larva: Setae 4-A, 3-C and 11-C usually

simple; dorsum of thorax and abdominal segments IV - V with large

Central black marks: ) SS 6 es asiaticus (p. 136)

Pupa: Setae 9, on VI - VII long and unbranched, slightly hooked at tip;

internal border of paddle with fringe hairs. Larva: Setae 4-A, 3-C

and 11-C branched; dorsum of thorax and abdominal segments IV - V

without central black marks. .......... interruptus (p. 140)

ANOPHELES (ANOPHELES) BULKLEYI CAUSEY

Anopheles bulkleyi Causey 1937b: 23: 543 (“).

Anopheles (Anopheles) bulkleyi Causey, Bonne-Wepster and Swellengrebel 1953:

129 (o"); Reid and Knight 1961: 481; Reid 1968: 195.

This species is known only from the adult male, which is recognizable by

the legs having clusters of dark scales on tarsomere 2 of the midleg and tarso-

meres 1 and 2 of the hindleg.

The following is Causey's description. ''Male. Head: proboscis, palpi,

and antennae all dark without white markings; occiput mainly dark with few

white scales along median line. Thorax: integument dark and ash gray with

brownish collar; prothoracic lobes covered with brown scales; mesonotum

sparsely clothed with small whitish hairs; pleurae mainly ash gray, scales

wanting, but some evidently rubbed off. Legs: heavily clothed with brownish

scales except for few white scales on coxae; tarsi of second pair of legs with

dark scales in collar-like cluster on second tarsal segment, and on third

pair of legs larger cluster on distal end of first and all of second tarsus. Abdo-

men: clothed with luxuriant growth of long brown hair; scales absent except

on terminal segments where they form large brown cluster over dorsal, ven-

tral, and lateral portion of hypopygium. Wings: with widely-spaced black and

Harrison and Scanlon: Subgenus Anopheles in Thailand 135

white scales; costa with faint white spot at tip of subcosta and first longitudinal

vein; proximal half of subcosta denuded of scales, but apical half all dark; first

vein with three small white spots on distal half and four on basal half, the lar-

gest one situated just basal to origin of third vein; second vein all dark except

for scattered white scales on first branch and patch of white scales on second

branch; third vein mainly white with few dark scales on basal half; fourth vein

mainly dark except for few white scales on first branch and large central

white spot on second branch; fifth vein with two dark spots on stem, three on

first branch and one on second; sixth vein all white except one dark spot be-

yond mid point."

FEMALE, PUPA and LARVA. Unknown.

TYPE-DATA. Causey (1937b: 543) indicated the type-male from near

Chandburi (probably Chanthaburi), Siam, had been deposited in the USNM.

However, it is not in that museum and there is no record of its having been

received. Causey (1966, personal communication) indicated the type was

probably among specimens lost en route to Brazil.

DISTRIBUTION. Known only from the type-locality, Chanthaburi, Thai-

land. Reisen, Burns and Basio (1971, Table 7) list 2 specimens of Anopheles

buckleyi | sic] collected from Don Muang Royal Thai Air Force Base, 25 km

north of Bangkok. Don Muang is a very busy urban area that includes inter-

national airline facilities, and is situated in flat rice plains and far removed

from jungle or forest-like habitats. The only specimen (the lost type) of

bulkleyi ever collected came from a tree hole in tropical wet forest. Almost

20 years of active anopheline collecting from numerous habitats, including

the type-locality, have failed to collect additional specimens. We are highly

skeptical of this record and suspect the 2 specimens were misidentified.

BIOLOGY. The type-male was reared from a larva collected in a tree

hole. Despite extensive collecting from treeholes in Thailand between 1961 -

66, no additional specimens have been collected.

TAXONOMIC DISCUSSION. Reid and Knight (1961) placed bulkleyi in the

Lophoscelomyia series based primarily on combination of abdominal and coxal

scales, unspeckled legs, unbanded tarsi and the pale mesal and dark lateral

areas of the scutum. The tufts of outstanding scales on the legs were con-

sidered of secondary importance since wellingtonianus Alcock from Peninsu-

lar Malaysia (subgenus Anopheles, series Anopheles) and squamifemuy Antunes

from South America (subgenus Lophopodomyia) also have this character.

This assignment appears reasonable based on the characters given in the ori-

gsinal description, however, it must be regarded as tentative, pending the col-

lection of additional specimens and stages. Even if bulkleyi is eventually

determined to be a true member of the Lophoscelomyia series, it will appar-

ently have no close relatives. Besides leg scaling differences, the wing of

bulkleyi, as described, is quite distinct from that of the species in the aszati-

cus group. :

ANOPHELES ASIATICUS SPECIES GROUP

(An. asiaticus group, Reid 1968)

Adults distinguished from bulkleyi only by having a prominent tuft of scales

(black proximally, white distally) on the apical 0.25 - 0.33 of the hindfemur,

instead of on the mid- and hindtarsomeres.

136 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

ANOPHELES (ANOPHELES) ASIATICUS LEICESTER

(Figures 70, 71, 72)

Anopheles asiaticus Leicester 1903: 291 (A, L-biology); Stanton 1912: 4;

Stanton 1915: 6: 164 (L).

Lophoscelomyia asiatica Leicester in Theobald 1904: 13; Alcock 1911: 244

(= Myzorhynchus asiatica); Stanton 1912: 4 (= Anopheles asiaticus).

Lophomyia asiatica of Giles 1904: 366; Theobald 1910: 55 (= Lophoscelomyia

asiatica).

Lophoscelomyia asiatica Theobald, Leicester 1908: 21 (%, 2, L); Strickland

1913: 138 (= Myzorhynchus asiaticus).

Anopheles (Anopheles) asiaticus Leicester of Christophers 1924: 23; Edwards

1932: 42; Gater 1934: 110 (L*); Gater 1935: 180 ('*, 2*); Bonne-Wepster

and Swellengrebel 1953: 156 (o'*, 9*, L*); Reid 1963: 108 (2*, L*); Stone

1967: 198.

Anopheles (Anopheles) asiaticus (Leicester), Stone, Knight and Starcke 1959:

15; Reid 1968: 196 (*, 9*, P*, L*, E*).

Adults of asiaticus are easily recognized by the scale tufts on the hindfemur,

pale golden scales on tergum VIII and the absence of a pale wing fringe spot at

R4.5- The pupal stage is recognized by the frayed or weakly branched seta 9

on segments VI - VII and the absence of fringe hairs on the internal border of

the paddle. The larva is very conspicuous due to the large central dark mark

on the dorsum of the thorax (not seen on skins) and the usually single setae 4-

A, 3-C and 11-C.

FEMALE (Fig. 70) Head. Vertex with broad dark erect scales except dorso-

central patch of pale scales above interocular space; interocular space with

short curved pale scales adjacent to eyes, long pale setae internal to scales

forming distinct pale frontal tuft; palpus dark scaled with scales on basal half

more erect, infrequently several dorsal pale scales at base of segments 3,

4 and 5; palpus approximately equal to length of proboscis; proboscis entirely

dark scaled, labellum nearly bare, paler; clypeus bare; pedicel with small

eray scales on upper, inner and lower outer aspects; flagellomere 1 with numer-

ous small dark scales, remaining flagellomeres without scales. Thovax. Scu-

tal integument creamy-gray centrally, brown laterally, except small mesal

brown area in prescutellar area adjacent to scutellum; anterior promontory

with slender pale scales centrally, several darker scales laterally; scutum

sparsely covered with thin pale seta-like scales and much longer tan setae;

scutellum with median brown spot, several thin pale seta-like scales and much

longer dark brown setae; anterior pronotum with dark scales on ventrocephalic

half; pleuron mostly dark brown, particularly lower portion of sternopleuron,

meron and metameron, in comparison with white coxae; pleural setae: 2 - 3

propleural, 2 - 4 spiracular, 5 - 9 prealar, 2 - 4 upper and 1 - 4 lower sterno-

pleural and 1 - 5 upper and 0 lower mesepimeral. Wing. Costa with pale

humeral, subcostal and apical spots; humeral and subcostal pale spots variable

in size, rarely absent; subcostal pale spot extends onto Rj; remigium with dark

scales; humeral crossvein without scales; R-R, with dark scales except sub-

costal pale spot and pale tip; Rg-R9.3 with duale scales to fork; Ro dark scaled

with white tip, infrequently with small basal pale spot; Rg entirely dark scaled

or with small pale spot on basal 0.33; R4,5 dark scaled; M dark scaled to fork;

Mj+2 and M344 dark scaled; Cu, Cuy and Cug entirely dark scaled; 1A dark

scaled; base of veins Rg, R445, M, Cu, and 1A with prominent clumps of

Harrison and Scanlon: Subgenus Anopheles in Thailand 137

black scales resulting in distinct spotted appearance; fringe scales dark ex-

cept apical pale spot from above R, down to Ro. Halter. Knob with dark

scales. Legs. Upper forecoxa with several small dark scales; mid- and

hindcoxae with long pale, curved plume scales projecting ventrocaudally on

lower half; upper midcoxa with 2 - 4 setae. Foreleg: trochanter with patch

of dark scales on caudal aspect; fenmiur swollen on basal half, dark scaled ex-

cept for narrow pale basal band; tibia and tarsus with dark scales. Midleg:

trochanter with setae and pale scales; femur dark scaled except for narrow

pale basal band and small dorsoapical white spot; tibia and tarsus with dark

scales. Hindleg: trochanter with setae and pale scales; femur dark scaled

except narrow pale basal band and long pale scales at apex; femur with apical

half swollen and prominent tuft of long erect scales on apical 0.3, scales in

tuft approximately equal length, black proximally, pale distally; tibia and tar-

sus with dark scales. Abdomen. Integument light brown, except distinct dark

brown-black mark on tergum IV; with long brown setae; without scales except

tergum VI may have several yellow scales on caudal margin, tergum VII usu-

ally with several yellow scales on caudal margin, tergum VIII covered with

yellow scales dorsally, black scales laterally and sternum VIII covered with

black scales.

MALE (Fig. 70) Head. Palpal segment 3 with small pale basal band, seg-

ment 4 with narrow pale basal and apical bands, and segment 5 with narrow pale

basal band and apical half entirely pale scaled. Wing. Subcostal pale spot on

costa large; more pale scales on other wing veins. Abdomen. Scales on seg-

ments VI - VIIl as on female, but less extensive. Genitalia. Basimeres with

yellowish dorsal and dark lateral scales; 2 parabasal spines with fine attenu-

ate hooked tips, innermost spine set on prominent tubercle; ventromesal spine

on basimere located approximately midway between apex and claspette; clas-

pette with 3 distinct lobes; ventral lobe with large central seta and much smal-

ler seta dorsolaterally; middle lobe with 2 - 3 large setae, much larger than

those on ventral lobe; dorsal lobe with 3 broad curved blade-like spines very

close together, not fused; aedeagus simple, without leaflets or spines.

PUPA (Fig. 71). Integument uniform light tan. Tvumpet. Simple, with

fairly wide opening, meatus approximately 0.3 - 0.5 length of trumpet. Abdo-

men. Most setae small, single or with few branches; seta 0 single on all seg-

ments; 8-II present, with 1 - 4 branches; 1, on V - VII very weak and short,

with 3 - 11 branches; 5, on V - VII dark, long and curved with stout central

stem and about 7 - 20 strong lateral branches; 5-VII almost equal length of

segment VIII; 9, on II - IV small, peg-like; 9, on V - VII long, curved and

sharp pointed, with few short fine lateral branches or frayed, approximately

0.3 - 0.4 of segment length; 9-VIII with stout central stem and many subequal

branches; 1-IX as long or longer than seta 0-VIII, extending beyond caudal

margin of segment IX. Paddle. 0.75 - 0.80 as wide as long; refractile border

0.55 - 0.65 of paddle length, beyond refractile border with long dense fringe

hairs to just before 1-P; internal border without fringe hairs; apex slightly

emarginate at 1-P attachment; 1-P with 12 - 25 subequal branches from short

stout base; 2-P weak with 2 - 4 branches.

LARVA (Fig. 72). Living and whole mounted specimens with dark mark

resembling Maltese cross on dorsum of thorax, and smaller central dark mark

on abdominal tergum IV and rarely terga IV - V; skins without above pigmen-

tation. Head. Antenna of nearly uniform diameter except near apex, with

weak spines mostly on basal and mesal aspects; seta 1-A at outer margin on

basal 0.25, weak, and approximately 0.25 - 0.33 as long as antennal stem,

with 2 - 6 fine distal branches; 4-A long and single; 1-C as long as 3-C; 2-C

138 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

long and single, rarely forked; 3-C single, rarely forked, about half as long

as 2-C; 4-C single, barely reaching level of 2-C alveolus, laterocaudal to 2-C

and closer to 3-C; 5-C long, single, reaching level of 4-C alveolus; 6-C short,

fine and usually single, closer to 5-C than 7-C; 7-C with 2 - 6 branches, longer

than 6-C; 8, 9-C short and single; 11-C slender, single or infrequently forked

on distal half, longer than antenna; maxillary palpus with stout darkly pigmented

seta on outer distal margin, with many finely frayed short branches on distal

0.66. Thorax. Seta 1-P short with 2 - 6 branches from basal half, inserted

about own length from 2-P; 2-P long with 8 - 16 branches and small faintly

sclerotized base; 3-P single adjacent to 2-P, approximately equal length of

1-P, 4-P with large darkly pigmented base, and stout stem and lateral branches;

11-P short, fine and usually single; 14-P with 3 - 6 branches; 4-M approxi-

mately in line with and equal length of 3, 5-M; 3-T well developed, with slightly

flattened branches and long narrow attenuate filaments; thoracic pleural tuber-

cles without prominent spines; long thoracic pleural setae single. Abdomen.

With small anterior tergal plates and very small posterior plates at least on

III - VII; seta 0 small, single on II - VIII; seta 6 long on segments I - VI, plu-

mose on I - III, long slender and single on IV - V, slightly shorter with approxi-

mately 12 - 20 branches on VI; 1-I minute and single; 1-II with 3 - 7 filamen-

tous branches; 1, III - VII with well developed flattened, evenly pigmented

leaflets, leaflets with shallow notched shoulder filaments half or more as long

as leaflet; most other setae single or with few branches, including seta 13;

pecten plate with approximately 20, mostly long teeth; seta 1 of the spiracular

area dorsocaudal to pecten plate, long and single; 1-X single, slightly longer

than saddle.

TYPE-DATA. Leicester (1903) is a short paper entitled, ''A breeding

place of certain forest mosquitoes in Malaya", where he reported finding a

fairly large number of species (many new) breeding in bamboo. These species

are not defined below generic level, except in 2 cases where he says ''two an-

opheles, Anopheles leucophyrus, | sic | anda new species with beautifully feathered

hind legs like an Aedeomyia, and which I have named Anopheles asiaticus,..."

Further down that page he also says, "Anopheles asiaticus I believe to be ex-

clusively a bamboo breeder;...'' We agree with Stone (1967) that these entries

constitute a description and therefore satisfy the rules of the International

Commission of Zoological Nomenclature pertaining to original descriptions

prior to 1930. Consequently, this publication has priority over the previously

considered original description of asiaticus (1904). Moving the date of the

original description from 1904 back to 1903, should have no effect on the status

of the holotype. This specimen, a ¢ in the BMNH, was sent to Theobald along

with the 1904 description by Leicester, and is in poor to fair condition with the

following parts missing: all legs, except left hindleg; antennae; and most of left

palpus. This specimen is labeled: (1st label) - "Bamboo, Ampang jungle, 6

miles fr. Kuala Lumpur, 27/6/03, No 10, A. asiaticus" (on underside of

cardboard minuten stage); (2nd label) is circular type-label; (3rd label) -

"Kuala Lumpur, Dr. Leicester"; and (4th label) - ''Lophomyia asiatica (Type)

Leicester."' The data underneath the minuten stage is hand penned by Leicester,

while that on labels 3 and 4 is hand penned by Theobald. This specimen was

collected nearly 3 months prior to the publication (15 Sept.) of Leicester (1903)

and is most probably the specimen on which he based his 1903 description.

This is supported by Leicester writing the correct name "A, asiaticus" under-

neath the minuten stage, not ''Lophoscelomyia asiatica" as it was named in

Theobald (1904). The "Lophomyia asiatica" label and the 1904 name "Lopho-

scelomyia asiatica" are probably Theobald modifications of the name Leicester

Harrison and Scanlon: Subgenus Anopheles in Thailand 139

had on his manuscript description sent to Theobald. Apparently, Theobald

modified the name so that he could describe a new genus "'Lophoscelomyia"' on

the page preceeding the L. asiatica description. This is supported in part by

Leicester (1908) crediting the 1904 species, Lophoscelomyia asiatica, to

Theobald not himself, even though Theobald (1904) clearly says that Lopho-

scelamyia asiatica is Leicester's species.

Because of the above circumstances, we feel the specimen previously con-

sidered the holotype of asiaticus based on the 1904 publication, is a specimen

used by Leicester in discussing asiaticus in his 1903 paper, which he then

selected to represent this species and sent to Theobald along with his full

length description. Therefore, this specimen remains the rightful holotype.

DISTRIBUTION (Fig. 71). This species is known from Malaysia and Thai-

land, where it is collected almost exclusively from bamboo habitats. Speci-

mens (120, 159, 74 larvae and 37 larval and pupal skins) in the USNM were

examined from the following provinces of THAILAND: Nakhon Si Thammarat,

Phangnga, Ranong and Tak. Other specimens (330, 3992, 13 larvae and 55

larval and pupal skins) in the USNM were examined from MALAYSIA. An ad-

ditional 3% and 42 specimens (including the holotype) from MALAYSIA were

examined in the BMNH.

TAXONOMIC DISCUSSION. This is the only species besides noniae that

has scale tufts on the hindfemur and the immature stages in bamboo. A search

in the USNM revealed 130, 152, 4 larvae and 37 larval and pupal skins of noniae

from Malaysia. Reid (1963) presented the following characters for separating

these 2 species. (Adults) - asiaticus with much more distinct clumps of black

scales on the base of wing veins M and 1A; wings generally paler than those

of noniae. (Pupa) - asiaticus with meatus about half the total length of trumpet

(0.67 on noniae); asiaticus seta 9-VII simple or slightly frayed (strongly bran-

ched on noniae), (Larva) - asiaticus with seta 4-C short, just reaching level

of 2-C bases (noniae 4-C long, reaching to or beyond tips of 3-C); asiaticus

1-A with 2 - 6 distal branches (noniae 1-A with 5 - 9 branches from near

base). The larval characters presented by Reid are very distinct, this stage

shows the clearest differences between asiaticus and noniae. Although we ex-

amined over 140 larvae or larval skins of asiaticus and noniae from Malaysia

and Thailand, none had forked setae 2-C as illustrated by Strickland (1914).

Reid (1963) suggested that Strickland's larva might have been a variation of

noniae, Or anew species. Reid (1963) also described differences in the dark

pigmented marks on the larvae of asiaticus and noniae. The dark mark on the

thorax of asiaticus typically has a cephalic extension which is lacking on noniae.

We found this a constant difference on the material in the USNM, although the

thoracic mark on asiaticus is quite variable. The dark mark on the abdomen

of the larva was described as lozenge-shaped and confined to segment IV on

asiaticus, while both segments IV and V on noniae have a pyramidal mark that

is joined where the segments meet. This mark was highly variable on aszati-

cus larvae (including Malay specimens) with less than half having a lozenze-

shape and most with a round mark on segment IV, or infrequently with a small

amount of pigment spreading onto segment V. The 4 larvae seen of noniae have

this mark as described by Reid.

Seta 9-VII on asiaticus pupae in Thailand is apparently more variable than

that described for specimens from Malaysia. Several specimens were noted

with this seta moderately frayed or branched, but not enough to cause confu-

sion with noniae when used in conjunction with the trumpet character.

The adult seems to be the most difficult stage to separate from noniae. A

number of asiaticus specimens with skins from Thailand were noted with darker

140 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

wings and less distinct scale clumps. An examination of the available speci-

mens revealed that asiaticus has 2 - 4 upper midcoxal setae, while noniae has

3-9. This character should be of value in separating these species. The

adult of asiaticus has a very distinct dark mark on abdominal tergum IV. This

character was present on the 92 adults examined, while noniae only occasionally

has a dark mark on terga IV - V or V alone.

All stages of asiaticus are easily separated from those of interruptus by

the key characters and descriptions.

Several characters were seen on the adults that deserve further comment.

The humeral pale spot on the costa although usually larger, may be only 2 - 3

pale scales on each wing or even only 1 wing (1/15). Wing veins Rs, R4i5,

M3,4, Cuz, Cug and 1A, with proper lighting, may appear bronzy, similar to

djajasanensis, as described by Brug (1926). The erect pale scales on the

hindfemur tuft are usually as long as the erect black scales (includes holo-

type), not shorter as depicted by Reid (1968: 197).

BIOLOGY. This species is found in hilly-mountainous terrain in close

association with large stem bamboos. In Thailand immatures have been col-

lected between 130 - 700 m elevation, while collections in Malaysia were made

between 30 - 1,060 m elevation. To date the immature stages have always

been found in association with bamboo. The preferred habitat seems to be

dried fallen bamboo that has a large enough split or hole to allow the inter-

nodes to collect water. However, infrequent collections in Thailand and/or

Malaysia have been made from bamboo stumps, fallen split green bamboo and

upright dried or green bamboo internodes. No adults have been collected in

biting collections in Thailand, however, Reid (1968) reports small numbers

captured with human bait in the forest. The natural adult food source is not

known. It is interesting to note that nearly 5 years of concentrated jungle

collecting occurred in Thailand before this species was first collected in Tak

Province. Apparently asiaticus has a very spotty distribution in Thailand.

ANOPHELES (ANOPHELES) INTERRUPTUS PURI

(Figures 73, 74, 75)

Anopheles annandalei var. interruptus Puri 1929: 387 (“, 9*, L*, E*).

Anopheles asiaticus of Christophers 1915: 385 (**); Puri 1929: 394 (= annan-

dalei var. interruptus).

Anopheles (An.) annandalei var. interruptus Puri, Puri 1931: 113 (L*);

Christophers 1933: 139 (“*, ¢*, L*); Bonne-Wepster and Swellengrebel

1953). Lobe, 5%); Reid) 19632111 (B*).

Anopheles (An.) annandalei of Edwards 1932: 42 (in part).

Anopheles (An.) interruptus Puri, Reid 1968: 203 (*, 9*, P*, L*, E*).

Identification of this species should not present problems in Thailand.

The adults are distinct on the basis of the additional pale wing fringe spot

and lack of pale scales on the abdomen. The pupa is easily recognized by

abdominal seta 9 and paddle characters, while the larva is distinct on the

basis of the head setal branching and the absence of dark marks on the thor-

ax and abdomen. Like asiaticus except:

FEMALE (Fig. 73) Head. Palpus usually with narrow pale bands at

base of segments 3 - 5. Thovax. Scutal integument gray centrally except

median brown area on prescutellar area adjacent to scutellum, dark brown

to black laterally; scutum sparsely covered with narrow curved pale scales

Harrison and Scanlon: Subgenus Anopheles in Thailand 141

and longer brown setae; scutellum with narrow curved pale scales and long

dark brown setae. Pleural setae: 1 - 3 propleural, 3 - 5 spiracular, 4 - 6

prealar, 2 - 4 upper and 2 - 5 lower sternopleural and 2 - 5 upper mesepimer-

al; upper and lower sternopleural setal patches with 3 - 7 and 4 - 8 pale scales

respectively. Wing. Costa with humeral, subcostal and apical pale spots;

humeral pale spot small, not extending onto base of vein R; subcostal pale

spot of variable size, not extending onto vein R-Ryz; Ry with pale tip; Ro-R9i3

with 3 patches of large black erect scales, at base of Rg, level of R4,5 base

and at Ro-Rg fork; Ro with pale tip; R3 entirely dark scaled or with pale tip;

R4+5 with patch of large black erect scales at base, entirely dark scaled ex-

cept pale tip; M entirely dark scaled, with 4 patches of large distinct erect

scales, at base, level of M-Cu crossvein, level of R-M crossvein and at

M1+2-M3+4 fork; M1+2 and M3+4 entirely dark scaled, or with pale tips; Cu

entirely dark scaled; Cu, with slightly enlarged black scales at level of M-Cu

crossvein, entirely dark scaled or with pale tip; Cug entirely dark scaled or

with pale tip; 1A with basal patch of large erect scales infrequently followed

by small patch of pale scales, usually entirely dark scaled; upper apical pale

fringe spot extends from slightly above Rz to Rg; lower apical pale fringe spot

narrow, at R45; posterior margin of wing rarely with pale fringe spots at

M344, Cuy and Cu9. Legs. Coxae very pale except upper portion of fore-

coxa; forecoxa with dark scales on dorsocephalic aspect; upper midcoxa with

2 - 3 setae, infrequently 1 - 2 scales. Foreleg: trochanter with patch of

dark scales on posteromesal aspect. Midleg: femur with dorsal V-shaped

pale spot near apex. Hindleg: trochanter with small patch of dark scales on

mesal aspect; tibia infrequently with small pale basal band. Abdomen. Integu-

ment dark brown-black, with black scales on sternum VIII and occasionally

tergum VIII; tergum IV same color as other terga.

MALE (Fig. 73) Head. Palpus with narrow pale bands at base of segments

3-5, apex of segment 5 also pale. Wing. Dark scales of patches and on sub-

costa very large. Abdomen. Without pale scales. Genitalia. Basimeres with

dark lateral scales, without pale dorsal scales; claspette with only 2 lobes,

otherwise as in asiaticus; aedeagus as in asiaticus. ,

PUPA (Fig. 74). Integument light tan except slightly darker trumpets.

Trumpet. Simple, with meatus approximately 0.3 length of trumpet. Abdo-

men. Setae 0, III - IV strong and darkly pigmented; 5-IV slightly shorter than

o-V; 0-VII approximately 0.8 length of segment VIII; 9, IV - VII without bran-

ches or fraying, hooked at tip; 9-IV nearly equal 9-V; 9-VII approximately

0.3 length of segment VIII; 9-VIII darkly pigmented, with long stout central

stem and lateral branches; 1-IX inconspicuous, shorter than 0-VIII. Paddle.

0.60 - 0.65 as wide as long; refractile border nearly entire length of paddle,

with short sparsely spaced fringe hairs; internal margin with fringe hairs;

1-P with long stout pigmented stem, with 2 - 5 branches from near midpoint;

2-P fine and single or with 2 - 3 distal branches.

LARVA (Fig. 75). Integument dark tan, usually with dense curved spicules

on the venter of thorax and abdomen; living and whole mounted larvae without

dark marks on thorax or abdominal segments. Head. Antenna with weak

spines along entire length; seta 1-A usually single; 4-A long with 4 - 11 bran-

ches; 2-C long, single or with 2 - 4 distal branches; 3-C with 2 - 9 branches;

4-C single, slightly closer to 3-C than to 2-C; 6-C short, single or with 2 - 5

distal branches; 7-C slightly longer than 6-C, usually with 2 - 3 branches; 8,

9-C usually single, infrequently forked; 11-C with 4 - 12 branches, mostly

on internal side; distal external seta on maxillary palpus less stout than that

of asiaticus. Thorax. 1-P with 10 - 15 branches, inserted close to pigmented

142 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

tubercle bearing large 2-P; 3-P single or forked, much shorter than 1-P;

11-P inapparent; 14-P with 2 - 8 branches; 4-M much shorter than 0-M; 3-T

with well developed leaflets; one thoracic seta in each pleural group with barb-

like side branches. Abdomen. Seta 6 long with numerous branches on seg-

ments I-VI; 1-II a well developed palmate seta; setae 9, on I - VI Spinous, with

large stiff sharp pointed branches; 7-VII spinous; pecten plate usually with

10 - 15 long teeth, often with shorter teeth in between long teeth; seta 1 of

spiracular area with 2 - 4 branches.

TYPE-DATA. Puri (1929) listed syntypes sent to the BMNH, Indian Museum

and the Malaria Survey of India Museum, Kasauli. Three of these syntypes

were examined during this study. A © and 2 in the BMNH were examined and

found in good condition. These were both labeled: (1st label) - "'type'' on

small circular label; (2nd label) - ''A. annandalei var. interruptus nov. var.";

(3rd label) - ''A''; (4th label) - "India, Bengal, Sukna, [X:1928, Dr. I. M.

Puri, B. M. 1929-450"'. The “ specimen also has a small label with the

number ''3870"' on it. An additional syntype “ labeled ''Syntype B" was ex-

amined in the Indian Museum collections, now housed at the Zoological Survey

of India, Calcutta. This specimen consists of the “ genitalia on a plastic

strip in.a good state of preservation, and the thorax, one wing and legs from

one side, all from the same specimen. All 3 specimens are from Sukna,

Darjeeling District, West Bengal, India, and all agree well with Puri's (1929)

and Reid's (1963, 1968) descriptions.

DISTRIBUTION (Fig. 74). In Thailand interruptus appears relatively

scarce with most collections coming from forested hilly or mountainous areas.

This scarcity may be a false indication due to the logistical problems in mak-

ing collections in many of these areas during the rainy season.

According to Reid (1968) this species occurs from southwest India and Sri

Lanka north to Yunnan Province, China, east to Indochina and south to Malay-

sia. Specimens (1°, 492, 13 larvae and 9 larval and pupal skins) in the USNM

were examined from the following provinces of THAILAND: Chanthaburi,

Chiang Mai, Nakhon Si Thammarat, Songkhla, Tak and Trang. Other speci-

mens (1%, 3¢ and 2 larval and pupal skins) in the USNM were examined from

CAMBODIA and MALAYSIA. Additional specimens examined include: 2° and

12 (including syntypes) in the BMNH from INDIA and MALAYSIA, 1° (syn-

type) in Zoological Survey of India collections from INDIA, and 52 specimens

from CAMBODIA in the ORSTOM collections. Recent records of this species

in Southeast Asia besides Reid (1968) include CAMBODIA | Buttiker and

Beales 1964 (1965)] and SOUTH VIETNAM (Nguyen-Thuong-Hien 1968).

TAXONOMIC DISCUSSION. Reid (1963) suggested annandalei var. inter-

vuptus probably deserved specific status due to its sympatric distribution with

the type form in northeast India (Puri 1929). Reid elevated interruptus to

specific status on this basis and additional evidence (in lit. from Coher, Reid

1968: 206) that both forms exist in Thailand. As discussed earlier (p. 133)

we have been unable to find specimens from Thailand that correspond to annan-

dalei, and therefore we do not recognize this species as occurring in Thailand.

Based on the homogeneity of the specimens examined from Thailand we will

follow Reid's lead and consider intevruptus a distinct species due to its sym-

patry with annandalei in India. This decision is tentative pending further in-

vestigation, but more appropriate than considering interruptus a subspecies

and implying genetic compatibility without a trace of evidence. This inter-

pretation leaves annandalei with a very widely discontinuous distribution (north-

east India and Java); however, the holotype and one of the paratypes of the

Javanese synonym, djajasanensis, do not agree with annandalei. Further

Harrison and Scanlon: Subgenus Anopheles in Thailand 143

study and specimens are needed to resolve the identity of the Java specimens.

Puri (1929) found one adult and one larval character to separate interruptus

from annandalei. The adult character, i.e., the presence or absence of a

subcostal pale spot on the costa, has since been invalidated by Reid (1963), who

found both extremes on Malayan interruptus. Therefore, the larval character,

i.e., the presence of barb-like branches on at least one of the long thoracic

setae in each pleural group of interruptus (all simple on annandalei), is cur-

rently the only character that can be considered valid. In Thailand although all

the larvae have the pleural setal type equivalent to interruptus, the number and

degree of branching on those setae is highly variable. On some specimens the

pleural setae are nearly simple, with only 1 - 2 very short barb-like branches,

while on others these setae have up to 20 longer barb-like branches. There is

apparently no correlation between the degree of branching on the long pleural

setae and the degree of development of the spine-like spicules on the venter of

the thorax and abdomen. Reid (1963) suggested this latter variation may be

due to environmental control such as Colless (1956) and Rosen and Rozeboom

(1954) demonstrated on Aedes larvae found in tree holes.

A number of variations detected on the Thailand specimens deserve com-

ment. Reid (1963) found the subcostal pale spot missing on 6 of 9 adults from

Malaysia. This spot was well developed on all of the Thailand specimens.

The only specimen seen without this spot was a male from Malaysia. Two

adults from Chiang Mai Province were seen with well developed pale fringe

spots on the wing opposite veins M3,4, Cu, and Cug. This condition was des-

cribed previously by Puri (1929) in reference to a paratype from the type-

locality. Several of the Thailand adults also had a pale basal band on the hind-

tibia.

One major difference was noted between the single Malayan pupal skin

studied and those from Thailand. Seta 9-III on the Malayan pupal skin was

pale and 0.5 as long as seta 9-IV, while on the 4 Thai pupal skins 9-III was

0.10 - 0.25 as long as 9-IV and usually darkly pigmented.

Reid (1963) mentioned that while Indian larvae had well developed spicules

on the venter of the thorax and abdomen, the Malayan larvae seldom have these

structures well developed. Most of the Thai larvae have well developed spicu-

les, but several larvae from Tak Province had these structures absent or

minute and barely visible. Reid (1968) lists seta 4-A on Malayan larvae with

8 - 11 branches, while we found 4-A with 4 - 9 branches on Thai larvae. We

also noted larval seta 14-P with 2 - 5 branches on Thai larvae, while the single

Malayan larval skin we examined had seta 14-P with 8 branches.

The significance of the above variations is impossible to interpret at this

point. It should be obvious that more specimens are needed for study from

many areas before the taxonomic problems surrounding the annandalei-

interruptus question can be resolved.

BIOLOGY. Anopheles interruptus has been collected between 5 - 900 m

elevation in Thailand. The primary immature source in Thailand has been

tree holes, however, 2 larval collections were made in root holes at or below

eround level. Relatively few larvae are found in any one tree hole. This

could be indicative of a tendency toward cannibalism, or simply a survival

mechanism where a few eggs are laid in several tree holes. Reid (1963) re-

ported 2 females taken in human biting collections in Malaya, and there are 2

females in the USNM which were collected biting man between 1900 - 2000

hours in Chiang Mai Province, Thailand.

144 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

ANOPHELES SERIES

Edwards 1932: 36.

The following definition for the series is taken from Reid (1968: 206).

"ADULT. Fore femur usually slender, tarsi rarely banded, coxae and

abdomen without scales, 2 palps usually slender and head scales often nar-

row, pronotal lobes sometimes without scales, wings with or without pale

areas; spines on the dorsal lobe of the harpago mostly separate, phallo-

some with or without leaflets.

PUPA. Trumpets simple, longer axis vertical (except perhaps in 4. gigas).

Other characters vary with the species group; in the aitkenii and culiciformis

gsroups, hair 1 on V-VII is small and weaker than hair 5 and usually has

some branches; in the lindesayi group hair 1 is strong, often simple, and

may be nearly as long as the segments; in all three groups the paddles are

usually rather narrow and generally with long fringe hairs.

LARVA. Antennal hair usually less than half as long as shaft, outer

clypeal hairs in Malayan and Bornean species usually simple or few-branched,

frontal hairs simplified in tree-hole species, long pleural hairs of the thorax

simple or branched, pleural hair epl simple, or with branches but those are

seldom spine-like from the base as in the Myzorhynchus series."

The above definition was directed more toward species that occur in Malay-

sia, but it also fits most of the other species in the series. Since this is such

a diverse, widely distributed series the reader should refer to Reid and

Knight (1961) for problems inherent in this arrangement and discussions on

the differences in the various species groups.

DISTRIBUTION. As currently defined this series is cosmopolitan and

probably the most widely distributed anopheline series. According to Reid

and Knight (1961) there are 8 species groups and other unassociated species

with the following distributions: maculipennis group - Nearctic and Palearctic;

plumbeus group - Nearctic and Palearctic; punctipennis group - Nearctic;

pseudopunctipennis group - Nearctic and Neotropical; aitkeniti, culiciformis

and lindesayi groups - Oriental; stigmaticus group - Australasian; and the

unassociated species, atratipes from the Australasian, concolor Edwards

from the Ethiopian, and algeriensis Theobald, claviger (Meigen), habibi

Mulligan and Puri and martevi Senevet and Prunelle from the Palearctic re-

gion.

Members of all 3 species groups known from the Oriental region have been

recorded from Thailand, however, the earlier records of gigas formosus Lud-

low and gigas sumatrana Swellengrebel and Rodenwaldt are not recognized

here. Further discussion regarding this decision can be found on page 21

and in Scanlon et al. (1968: 11-12). Only the aitkenii and culiciformis species

eroups are considered as representing the Anopheles series in Thailand.

TAXONOMIC DISCUSSION. This is a most difficult series to define for

the adults of many of the species are small and unicolorous, while others are

large with brightly patterned wings and legs that approximate a subgenus Cellia

habitus. Infact, Reid (1968) suggested that the lindesayi and pseudopunctipennis

species groups, which have the 2 last named characters, may be more ancient

and nearer to the ancestral form from which the subgenus Cellia diverged.

The adults of the species of the Anopheles series occurring in Thailand all fit

the former habitus described above and are easily recognized. A definition

of these species follows:

ADULT. Small generally unicolorous brown species, female palpus equal

Harrison and Scanlon: Subgenus Anopheles in Thailand 145

to or slightly shorter than proboscis; anterior pronotal lobes without scales;

aedeagus with or without spines or hair-like leaflets.

PUPA. Trumpets simple, longer axis vertical; seta 1, on V - VII small,

weaker than seta 5 (except palmatus), usually with branches; paddles generally

narrow, usually with long fringe hairs.

LARVA. Seta 1-A less than half as long as antennal shaft, inserted on

basal 0.25; seta 3-C usually simple or with few branches; setae 5, 6, 7-C

large or small, but branched; seta 11-P with 1 - 4 slender branches; seta 6,

on IV - VI long or short.

KEYS TO THE SPECIES GROUPS AND SPECIES OF

THE ANOPHELES SERIES IN THAILAND

ADULTS. (kyondawensis adults unknown).

1 Erect head scales long and narrow (Fig. 80), only slightly expanded

apically; flagellomeres without scales. aitkenii sp. group. ... 2

Erect head scales broad, widely expanded apically (Fig. 94); flagello-

mere with dark seales:) \o i c.wios Agaeks aie (culiciformis sp. group)

sintonoides (p. 181)

21): Abdominal segments IV, or IV and V distinctly paler than other seg-

sae ks Satta. Sia ERE Pe ee a cl AT a iaeeal eae 3

All abdominal secments unicGlorous. nie aS ea rae ek é

3(2). Abdominal segments IVand Vpale..... aberrans [ in part] (p. 152)

Only abdominal segment IV pale. ..... palmatus | in part] (p. 173)

4(2). Prescutellar space with short fine setae back to scutellum.

insulaeflorum (p. 167)

Prescutellar space with a distinct median bare area immediately cephal-

ad. to SCS LIRR ep WP sok aS aberrvans [ in part] (p. 152)

bengalensis (p. 157)

fragilis (p. 162)

palmatus [| in part] (p. 173)

stricklandi (p. 171)

¢ tigertti (p. 164)

PUPAE (kyondawensis pupa unknown).

oe Trumpet with deep meatal cleft, meatus (stem) 0.33 or less as long as

trumpet; seta 1-P simple, tapering to fine point; ground water habi-

LAT We ae hk ca a ae Ce erat aitkenii sp. group. ...2

Trumpet with shallow meatal cleft, meatus more than half as long as

trumpet; seta 1-P simple filiform, with hooked (Cellia-like) tip;

axil, bamboo and treehole habitats. .... (culiciformis sp. group)

sintonoides (p. 181)

2(1). Seta 9 on abdominal segments IV - VII long with blunt round tip

CPi BTS Nee eer i Oe eee etude gre SB palmatus (p. 173)

Seta 9 on abdominal segments IV - VII long or short, with sharp pointed

th (Pigie Bay Ba ee I, ety NG) aah id peel

146 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

3(2). Seta 9 on abdominal segment VII less than 0.33 length of segment VIII;

seta 9-IV, 2 - 6 times as lope as 9-H. 3 foe Sor ee ee, 4

Seta 9-VII, 0.33 - 0.50 length of segment VIII; seta 9-IV, 8 or more

tunes as lone as Jel wee err ek Dae ee a 5)

4(3). Seta 9-IV, 2 - 3 times as long as 9-III; tergum IX with very prominent

laterocaudal knob over root of each paddle (Fig. 89); seta 9-VII, 0.12 -

0; 16 as lone as se€ement VIE. so. eae ee stricklandi (p. 171)

Seta 9-IV, 4 - 6 times as long as 9-III; tergum IX without prominent

laterocaudal knobs; seta 9-VII, 0.20 - 0.25 as long as segment VIII.

insulaeflorum (p. 167)

0(3). Refractile margin on paddle long, 0.60 - 0.75 paddle length.

bengalensis* (p. 157)

Sragilis* (p. 162)

Refractile margin on paddle short, rarely reaching 0.5 paddle length.

6(5). Seta 6-III long, nearly equal length of segment III lateral margin; 10-IV

long, nearly equal length of segment V lateral margin; 1-III weakly

developed, not as stout as 3-II............ tigertti (p. 164)

Seta 6-III short, about half as long as segment III lateral margin; 10-IV

short, about half as long as segment V lateral margin; 1-III well

developed, at least as stout as 3-II]........ aberrans (p. 152)

LARVAE

13 Setae 5, 6, 7-C long, well developed and plumose; 4-C branching from

base; 6-VI much shorter than 6-II]. .. aitkenii sp. group... . 2

Setae 5, 6, 7-C reduced, some or all short with few branches; 4-C

simple or branched on distal half; 6-VI nearly as long as 6-III.

culiciformis sp. group. ... 8

2(1). Anterior tergal plates on segments I-VII large, 0.66 - 0.75 the width

of a given segment; seta 1-P fan-like. ...... palmatus (p. 173)

Anterior tergal plates on I-VII small, less than half the width of a given

secment seta 1-P not fan-lHkes se Bre ee oe We 3

3(2).. Seta 6-i wih more than 15 branchés.'*. 6. 6 OP ae ee 4

Seta O=U1 with less than 15 branches: yor a er a 6

4(3). Seta 2-C single or with 2 - 3 main branches and short fray-like side

branches apout the middle. . 60 ei es fragilis (p. 162)

Seta 2-C with 2 - 14 branches (rarely simple), no short fray-like bran-

CHES AmOuE ne Mahe) ra ee a Se Ge 5)

5(4). Seta 11-II long and simple; 5-II with 3 - 6 branches; 14-P with 3 - 5

branches, rarely 5 on both sides. ......... aberrans (p. 152)

Seta 11-II forked or branched beyond base; 5-II with 7 or more branches;

14-P with 5 - 8 branches, rarely 5 on both sides.

bengalensis (p. 157)

*Characters used by Reid (1965, 1968) to separate these 2 species break down

in Thailand bengalensis.

Harrison and Scanlon: Subgenus Anopheles in Thailand 147

6(3). Setae 2-C with bases separated by approximately same distance as bases

of 2-C and 3-C; 11-II forked beyond base. ..... tigertti (p. 164)

Setae 2-C with bases close together and distant from bases of 3-C;

11<Th lone andisimplesns <5 (eee eC ie Se ee 7

7(6). Seta 1-I undeveloped, with filamentous branches; 2-VI with 4 - 5

branches; 3-C with 2 - 4 branches........ stricklandi (p. 171)

Seta 1-I palmate, with small flattened leaflets; 2-VI with 1 - 3 branches;

3-C simple or infrequently forked distally. . . insulaeflorum (p. 167)

8(1). Antennal shaft markedly curved (Fig. 93); integument of thorax and

abdomen smooth, without spicules....... kyondawensis (p. 179)

Antennal shaft straight or only slightly curved; integument of thorax

and abdomen with numerous spicules. ..... sintonoides (p. 181)

ANOPHELES AITKENII SPECIES GROUP

(A. aitkenii species group, Reid and Knight 1961)

Anopheles aitkenii James 1903.

These species are small and unornamented in the adult stages and have a

culicine resting posture. Unfortunately, females of the majority of species

are still indistinguishable. Little attention has been paid to members of this

group because they are mostly secretive forest species and do not appear to

be involved in the transmission of human disease pathogens.

The adult feeding habits are almost unknown and the biological information

that is available is difficult to interpret due to the difficulty in identifying the

females. Even the geographical distributions of the respective species are

difficult to interpret, for many published records refer to the species group

rather than a given species. This group was reviewed most recently by Reid

(1965, 1968) and the following group descriptions are based in part on these

works.

ADULT. Small, usually unicolorous brown; with culicine resting posture;

interocular space narrow; erect vertex scales very narrow; female palpus ap-

proximately equal length of proboscis, very slender with slightly expanded tips;

anterior pronotal lobes without scales; aedeagus bare or with small spines,

without hair-like leaflets; claspette with 2 lobes, number of setae on both

lobes generally variable; dorsal lobe with outer setae separate, inner setae

very close together or fused into club-like structure; ventromesal spine on

basimere inserted near apex (except acaci).

PUPA. Trumpet with deep meatal cleft, meatus less than half as long as

trumpet; seta 1-P simple and straight.

LARVA. Setae 2-C close together or with bases separated by approximate-

ly same distance as bases of 2-C and 3-C; seta 4-C branching from base; setae

5, 6, 7-C long, well developed and feathered; seta 1-A usually with 5 or more

branches; seta 11-C plumose, as long as or longer than antenna; seta 1-II

(palmate), well developed; seta 6-VI short, not more than 0.5 length of 6-III;

seta 1-X usually branched.

DISTRIBUTION. This group is basically confined to the Oriental faunal

region. Records of its members extend from western India to Amami Island

in the Ryukyus and south through Indonesia to the Philippines and Misool Island

off the western tip of New Guinea. In the Australasian region this group is

148 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

replaced by the stigmaticus species group which differs from it by several

adult characters (Reid and Knight 1961). Some members of the plumbeus spe-

cies group, i.e., barberi Coquillett and judithae Zavortink of North America,

and omorii Sakakibara of Japan, have a very similar adult habitus to that of

the aitkeniz group. However, members of the plumbeus group all oviposit in

treehole habitats and have immature stages quite distinct from those of the

aitkenii group.

TAXONOMIC DISCUSSION. We recognize 12 species in the aitkenii group,

one of which is described here as new. They are: aberrans n. sp. 3; acaci

Baisas, aitkenii James, bengalensis, borneensis McArthur, fragilis, insulae-

florum, palmatus, pilinotum Harrison and Scanlon, pinjaurensis Barraud,

stricklandi and tigertti. ,

The taxonomy of this group has been confused for a long time partially due

to an inability to find characters to differentiate the females. However, other

explanations for this confusion may have been the failure of some previous

workers to consider certain larval and male genitalia characters as variable

and to try to define the limits of these variations. Another important factor was

the very limited number of adults reared with associated skins for comparison.

This latter step is an essential part of correctly identifying many mosquito

species, and is absolutely necessary in attempting to identify apparently indis-

tinguishable adults. Reid (1965, 1968) is one of the first workers to attempt a

revision of the aitkeniz group based on reared associated specimens. His work

clarified and resolved many long-standing problems and is considered a major

achievement particularly in view of the limited material with which he worked.

During the present study we were extremely fortunate to have many reared

specimens of several species. The examination of this material has revealed

some interesting facts about variation in the male genitalia characters, the

branching of larval seta 2-C and has confirmed the suspected importance of

pupal characters in this group.

The number of outer dorsal setae on the male claspette has been used by

many workers as a valid means of separating aitkenii (3 setae) from bengalen-

sis (2 setae), yet aitkenii is variable. A review of the literature reveals: (1)

Christophers (1915) originally described aitkenii as having ''2 - 3 sword-like

chaetae on the ventral lobe" (= outer dorsal lobe); (2) Puri (1930) said aztkenii

definitely has 3 outer dorsal setae (ventral lobe of Puri) and this is distinct

from bengalensis which only has 2; (3) Christophers (1933) said aitkenii has 3

outer dorsal setae and bengalensis usually has 2, and in figures 17(9) and 17

(10) the claspettes of these species are depicted thus; (4) Reid (1965) followed

Christophers (1933) and illustrated a topotypic % of aitkenii from Karwar with

3 outer dorsal setae, however, Reid said aitkenii usually has 3 outer dorsal

setae, while bDengalensis usually has 2. As can be seen the setal number on

the outer dorsal lobe has vacillated from a variable number to a fixed number

and then back to a variable number. The variable interpretation is the approach

we feel is correct. We dissected the genitalia of 2 topotypic aitkenii males

and found one with 2 and the other with 3 outer dorsal setae, yet the other

features of these 2 specimens were in perfect agreement. We examined large

numbers of 4 species, aberrans, bengalensis, insulaeflorum and pilinotum,

and these all exhibited variations in the following claspette characters: outer

dorsal setae; ventral setae; and in several cases the inner dorsal setae (see

discussions under the respective species). Apparently, claspette characters

are as variable in the aitkenii group as they are in other species groups in the

subgenus Anopheles. Consequently, identifications of male aztkenii group

members based on claspette characters alone should not be considered reliable.

Harrison and Scanlon: Subgenus Anopheles in Thailand 149

Claspette characters should be used only when they can be substantiated by dis-

tinct aedeagus characters and/or associated larval and pupal skins.

The extent and location of branching on larval seta 2-C is variable and still

needs considerable study, preferably utilizing F; siblings, so that variation

parameters can be established. Such studies will definitely be needed to re-

solve the aitkenii-bengalensis problem in India, and the bengalensis -fragilis

problem in Southeast Asia. To outline this problem further, the following para-

meters are currently known: Branching - aitkenii (2), bengalensis (simple to

9), aberrans (2 - 14), fragilis (simple to 5) and tigertti (2 - 5); Location of

branching on stem - aitkenii (0.25 - 0.33), bengalensis (0.25 - 0.75), aberrans

(0.33 - 0.50), fragilis (0.33 - 0.50) and tigertti (0.50 - 0.67). The overlap in

these parameters creates considerable confusion, e.g., a significant percentage

of bengalensis larvae from Assam have 2-C bifurcate (without frays) at 0.40 -

0.67 along the stem. Using only this character for separating species, these

larvae could be bengalensis, aberrans, tigertti or possibly even aitkenii.

Using additional characters, 11-II is bifurcate, eliminating abervans, and

6-III has more than 15 branches, eliminating figertti. Larvae of aitkenii have

not been seen with branching so far along the stem, but only 3 larvae of aitkenii

were seen. Larvae such as those from Assam also occur in Burma and nor-

thern Thailand. These are probably responsible for the numerous aitkenii col-

lections recently listed from Burma (Khin-Maung-Kyi 1971a).

The fraying found on 2-C of fragilis also deserves further study. We are

currently considering any amount of fraying on 2-C (1 or more tiny central

branches) as indicative of fragilis, even when all the other characters on the

larva suggest bengalensis. This interpretation meets its most severe test in

the Philippines where specimens that are essentially bengalensis, may have

1 - 2 small median frays on 2-C. Since we have limited specimens and can

find no other characters to define fragilis larvae, we follow Reid (1965, 1968)

in using this arbitrary interpretation.

Other larval setal characters are apparently very stable. Such characters

include: position of 2-C in relation to 3-C; the number of branches on 6-III;

and whether 11-II is long and simple, or long and branching beyond the base.

This latter character has been checked on over 900 whole larvae and larval

skins of 11 species, and without exception these species can be divided as

follows.

(A) 11-II long and simple - aberrvans, acaci, borneensis, insulaeflorum,

pilinotum and stricklandi.

(B) 11-II long and at least bifurcate - aitkenii, bengalensis, fragilis, pal-

matus and tigertti.

The pupae of all the members of this group have been described or are

described herein, except, aitkenii sensu stricto, and pinjaurensis. Pupal

characters located thus far seem very stable, consequently, this stage is one

of the best means of identifying most species in the group. Characters found

most reliable are: shape and length of seta 9 on segments III - VII; and the

refractile margin, fringe hairs and shape of the lateral margin on the paddle.

Regardless of characters found during this study that will differentiate

adults of several species, the taxonomy of the aitkenii group still rests on our

knowledge of the immature stages and in a restricted sense, male genitalia

characters. Therefore, persons working with members of this group should

be resigned to the necessity of rearing adults with associated skins for posi-

tive identification. This is particularly true for workers outside of Thailand

where certain characters used here need further investigation.

Five of the currently recognized species, i.e., acaci, aitkenii, borneensis,

150 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

pilinotum and pinjaurensis, are not found in Thailand. Since they are not

treated thoroughly, each deserves a short discussion.

Anopheles acaci is currently known from the Philippines (Luzon) and

Borneo (Reid 1968). The male is unique in this group because it has female-

like antennae. The male genitalia are fairly distinct because of the 3 broad

outer dorsal setae on the claspette, the aedeagus which is abruptly narrowed

before the apex and the ventromesal spine which is inserted about 0.33 the

length of the basimere from the apex of the basimere. The pupa is very

similar to that of pilinotum, but has seta 5-I short and approximately as long

as 2-I, seta 0 on II - VII simple or bifid and seta 1 small and weakly developed

on segments II - VII. The pupa of pilinotum has 5-I much longer than 2-I, 0,

on It - VIII with 3 - 6 branches and abdominal seta 1 well developed. The

larva is very similar to borneensis from which it is separated by having the

branches on 3-C beginning 0.33 or more distant along the stem. Seta 2-C on

acact can look very much like that of bengalensis, but seta 11-II on a@cacz is

simple while that of bDengalensis is at least bifurcate. The immature stages

of acaci are usually very darkly pigmented and are recorded from shaded rock

pools in the forest. The type of acaci is apparently non-extant, but Reid (1968)

mentioned 2 male and one female paratypes in the BMNH. The female and one

male paratype in the BMNH have associated immature skins; however, the

associated immature skins for the paratype male (Lot A-11) used by Reid (1965)

to illustrate the genitalia of acaci are not in the BMNH, but in the USNM. There

are additional immature skins on slides in the USNM that are obviously topo-

typic, collected on the same day as the type and probably paratypes, but there

are no associated adults for these skins.

Anopheles aitkenti is recognized as being confined to the Indian subregion.

This interpretation is in conflict with Reid (1965) who considered aitkenii as

found from India to the Philippines. Reid (1965, 1968) based his Malaysian

record of aitkenii on a single female with associated immature skins, and used

a topotypic male from India to illustrate male genitalia characters. Reid's

(1968) interpretation was modified and aitkenii was listed as only possibly

occurring in the Philippines; however, he did not remove pallida Ludlow 1905,

from synonymy under aitkenii, where he placed it in 1965. Under our inter-

pretation all previous Southeast Asian references to aitkenii sensu stricto,

necessarily refer to other species in the group, usually bengalensis. We have

examined Ludlow's female type of Stethomyia pallida in the USNM and it is

probably equivalent to bengalensis or fragilis rather than acact, which is

described as a dark species (Baisas 1946). Ludlow's specimen does not have

fine setae on the caudal part of the prescutellar area and thus, is definitely

not pilinotum. If pallida Ludlow 1905, is eventually found to be synonymous

with either bengalensis or fragilis the name pallida still cannot be used because

pallida Theobald 1901 has priority.

Our restriction of the distribution of aitkenii is based partly on the known

aitkenii-like seta 2-C variations that occur on aberrans and bengalensis lar-

vae. The primary reason for this interpretation is the distinct shape of the

aedeagus, the large hood-like club and the variable number of outer dorsal

setae on the claspette (Fig. 79) of the 2 topotypic males in the USNM.

A total of 9c, 129, 2 whole larvae, and 2 larval and pupal skin specimens

were examined in the USNM and BMNH from southern India and Sri Lanka.

The single pupal skin (Sri Lanka) was badly damaged, and best left undescribed,

however, seta 1-IV had 5 - 6 branches, not 11 branches as described by Reid

(1965, 1968), see discussion under bengalensis. The holotype female is loca-

ted in the BMNH and bears the following labels: (1st label) - "#44, Karwar,

Harrison and Scanlon: Subgenus Anopheles in Thailand 151

15° 4:02" on underside of circular minuten platform; (2nd label) - ''Goa India,

H. E. Aitken"; (3rd label) - small round BMNH type-label; (4th label) - "Ano-

pheles aitkenii (Type) James"; and (5th label) - ''SEAMP Acc #306." This

specimen is in near perfect condition, all legs, antennae, palps, abdomen,

proboscis and wing scales are present. The following measurements and ob-

servations were made: wing length approximately 3.0 mm; wing cell M9, 0.5

length of petiole; wing cell Ro, 2.6 times as long as wing cell Mo (see bengal-

ensis, Fig. 80); the forefemur/proboscis ratio is 0.82; there is 1 propleural

seta on the right side (the left is obscured); the prescutellar area is without

fine setae as on insulaeflorum and pilinotum ; and the abdominal segments are

all unicolorous dark brown. Besides the holotype, 60, 6 and 1 larval topo-

typic specimen from the original 1902 - 03 collections by Aitken and Cogill,

were examined. Two of the 6 males are located in the USNM and genitalia

Slide mounts of these were used for preparing the illustrations herein. Two

of the 4 males in the BMNH have the genitalia intact and unmounted. Genitalia

preparations of the remaining 2 males were examined and the claspettes (Reid

1965, 1968) are identical to those of the USNM specimens, except one of the

latter only has 2 outer dorsal setae. Unfortunately, the shape of the aedeagus

is obscured on the 2 slide mounted specimens in the BMNH. (Also see discus-

sion under bengalensis).

Anopheles borneensis is known only from Borneo (Reid 1968). The male

of this species has densely plumose antennae as is normal for the group (cf.

acact). The male genitalia are very similar to those of fragilis and do not have

the tip of the aedeagus abruptly narrowed (cf. acaci). The pupa has a short

refractile border on the paddle similar to acaci, but has much longer setae 9

which are often frayed or branched, particularly on segment VII. The larva is

easily confused with that of acaci, but generally has seta 3-C with wide branches

from the base. Seta 2-C is much more densely branched near the base than

bengalensis. Seta 11-II is long and simple (cf. bengalensis). The types of

borneensis were listed by Stone et al. (1959) as in the Malaria Research Head-

quarters, North Borneo, but Reid (1968) says they are probably lost. Imma-

tures of borneensis are recorded from clear, cool running water under dense

shade in the forest.

Anopheles pilinotum is the species in the Philippines and eastern end of

Indonesia that has previously been called insulaeflorum (Harrison and Scanlon

1974). Adults can be separated from all other group members (except insulae-

florum) by the short fine setae on the prescutellar area of the scutum that occur

nearly all the way back to the scutellum. The male has short distinctive setae

on the ventral lobe of the claspette and more short spines on the tip of the

aedeagus (Fig. 79). The pupa is very similar to both acaci and insulaeflorum,

but can be separated from the former by having seta 1 well developed with

numerous branches on segments III - V, and from the latter by having: the pad-

dle refractile border less than half the paddle length, the outer margin of the

paddle straight or nearly straight some distance, and seta 0, II - VII with 2 - 6

(usually 3 - 5) branches. The larva of pilinotum is very similar to that of in-

sulaeflorum, but can be separated from the latter by having: seta 1-X with

2 - 4 branches and seta 0 on II - VIII with 3 - 6 branches. The holotype of

pilinotum and other members of the type-series are deposited in the USNM

collection. Several paratypes are deposited in the BMNH. Immature stages

of pilinotum are most often collected in clear running forest stream pools

under heavy shade.

Anopheles pinjaurensis is known only from a single male collected in

Punjab, India. The only distinctive character from aitkenii is an extremely

152 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

long aedeagus (Christophers 1933: 110).

BIOLOGY. All the females of this group apparently oviposit in ground

water habitats. The most commonly encountered immature collection sites

are seepage marshes, pools or quiet clear running forest streams with heavy

shade. Certain species appear to have quite specific habitat requirements,

e.g., tigertti found only in fresh water crab holes, while others have more

general requirements, or their specific requirements are still unknown. Adult

females of this group are occasionally collected biting man, there are records

(Christophers 1933; Macdonald and Traub 1960; Scanlon and Esah 1965; Harina-

suta et al. 1970) which make presumptive identifications of aitkenii and ben-

galensis respectively, but these records need confirmation. Hopefully, the

few adult characters offered here will allow workers to confirm or eliminate

certain species as feeding on man. The natural food sources of this group

are unknown and none of the species are known to be involved in the transmis-

sion of human pathogens. :

ANOPHELES (ANOPHELES) ABERRANS NEW SPECIES

(Figures 76, 77, 78, 79, 83)

A majority of the adults of this species can be identified on the basis of

certain abdominal segments being paler than others. Males cannot be identified

by genitalia characters alone. The pupal and larval stages are quite distinct,

and each can be differentiated from other Thai species by several highly reli-

able characters.

FEMALE (Fig. 78) Head. Vertex with very narrow erect scales, slightly

expanded and notched at tip; interocular space narrow, with short white scales

and long dark brown frontal setae; clypeus without scales; pedicel and flagello-

meres without scales, antennal whorl setae 3 - 5 times as long as segments;

proboscis long, slender with decumbent scales; ratio forefemur/proboscis

(sample 25 females), range 0.81 - 0.96, mean 0. 89; palpus very slender, ap-

proximately equal length of proboscis, with 2 apical segments slighty swollen,

scales decumbent. Thorax. Anterior promontory without scales; scutal

integument tan, often with paler orange longitudinal line on each side of slightly

darker median (acrostichal) longitudinal line; scutum without scales, with

mixed long and short dark brown setae in anterior promontory, acrostichal,

dorsocentral, lateral prescutal, fossal, antealar and supraalar groups; with

bare prescutellar space; scutellum with long and short dark brown setae, with-

out scales; anterior pronotum with sparsely scattered setae, without scales;

pleural area pale tan, without scales; pleural setae; 1 propleural, 1 - 6 spira-

cular, 1 - 3 prealar, 1 - 3 upper and 1 - 4 lower sternopleural and 2 - 5 upper

and 0 lower mesepimeral. Wing. Unicolorous, with narrow dark scales;

humeral crossvein without scales; cell Mo, 0.5 - 1.0 length of stem (from

crossvein). Halter. With dark brown scales. Legs. Coxae without scales,

upper midcoxa with 3 - 5 setae; femur, tibia and tarsi long, slender and en-

tirely dark scaled. Abdomen. Without scales, with long brown setae, integu-

ment always brown to black on segments I - III and VI - VIII; segments IV - V

distinctly paler, often yellow (see taxonomic discussion), infrequently same

color as other segments.

MALE (Figs. 76, 78, 79). Like female except antennae with more and

longer whorl setae; palpus with 2 apical segments distinctly flattened and club-

like. Genitalia. Basimeres without scales, with 2 parabasal spines on tuber-

cle; ventromesal spine inserted near distimere on distal 0.25 of basimere;

Harrison and Scanlon: Subgenus Anopheles in Thailand 153

claspette with 2 lobes; dorsal lobe with 2 - 3 outer setae, inner club formed

from 1-2 separate basal stems; ventral lobe usually with 3 large, 1 small

setae and numerous minute spicules; aedeagus bare, 3 or more times as long as

width at base, tapering to rounded tip less than 0.5 width of base.

PUPA (Fig. 76). Integument light tan to light brown. Cephalothorax.

Dorsum (between trumpets) often darker than remainder; seta 7-C with 2 -

0 branches. Trumpet. Same degree of pigmentation as surrounding cephalo-

thoracic integument; with deep meatal cleft, meatus less than 0.33 as long as

trumpet. Abdomen. Seta 0 small and simple; seta 9 long and narrow with

sharp tapered point on IV - VII; 9-IV, 10 or more times (usually 15) as long as

9-ITI; 9- VII, 0.33 - 0.50 as long as segment VIII; seta 1, on IV - VII long, with

few branches; 1-III well developed, as stout as 3-II; 1-IV with 1 - 4 branches

(rarely 1 or 4); 1-VI with 1 - 3 branches; 1-VII with 1 - 3 branches; 6-III short,

about 0.5 as long as segment III; 5-I approximately equal to or shorter than 2

and 3-I; seta 5 strongly developed on segments IV - VII, 0.67 to nearly equal

length of segment, with stout central stem and decumbent branches; 5-IV

branches decumbent, too close to stem to count; 10-IV short, about half as

long as segment V lateral margin. Paddle. Refractile margin short, 0.33 -

0.50 (rarely slightly more than 0. 50) as long as paddle; outer margin straight

or nearly straight on distal half; fringe hairs 0.5 or more as long as 1-P; 1-P

simple and stout, with tip slightly curved; 2-P simple or with 2 - 3 distal bran-

ches.

LARVA (Fig. 77). Usually with distinctive pale and dark color pattern in-

volving thorax and abdominal segments (also see taxonomic discussion); ventral

and lateral aspects of abdomen often with dense cover of minute spicules. Head.

Antenna base slightly wider than tip; antenna with long spicules, particularly on

mesal and ventral surfaces; seta 1-A short with 4 or more branches, inserted

on basal 0.17 - 0.20 of antenna; 2, 3-A with one edge serrate; 4-A with 5 or

more branches; setae 2-C with bases separated by approximately same distance

as bases of 2-C and 3-C on one side; 2-C with 2 - 14 branches originating

0.33 - 0.50 way along stem; 3-C approximately 0.33 as long as 2-C, simple or

split into 2 - 5 distal branches; setae 4-C slightly wider apart than setae 3-C,

with 3 - 8 branches from near base; 5, 6, 7-C well developed and plumose,

o-C usually extending forward to tip of setae 4-C; 8-C with 1 - 3 branches

(rarely 3 on both sides); 9-C with 4 - 9 branches; 11-C approximately equal

to length of antenna, with many branches particularly at apex. Thovax. On

whole larvae usually pale yellow in comparison with some brown abdominal

segments. Seta 1-P with 8 - 14 branches, about 0.5 length of 2-P, without

sclerotized base; 2-P with 10 - 17 branches, arising from sclerotized tuber-

cle; 2-P tubercle with prominent apical spur or tooth; 3-P short, simple or

bifurcate distally, closer to 2-P than 1-P is to 2-P; 4-P with stout stem and

numerous short lateral and apical branches, slightly longer than 2-P; pro-

thoracic pleural tubercle with distinct slender spine; 11-P simple or with 2 -

6 fine distal branches; long pleural setae usually simple; infrequently bifurcate

distally especially on pro- and metathorax; 13-P with 8 - 19 dendritic branches;

14-P with 3 - 5 branches, rarely 5 on both sides; 4-M small, simple or with

2 - 4 branches, inserted caudally to long and simple 3 and 5-M; 3-T unpig-

mented or slightly pigmented with well developed flattened leaflets without

filaments. Abdomen. On whole larvae, segments I - III and VI - VIII brown

or gray, while segments IV - V and X usually noticeably paler, often yellow

(see taxonomic discussion). Anterior tergal plates 0.25 - 0.33 width of seg-

ment on I - VII; posterior tergal plates commonly on segments IV - V as well

as VI - VII; seta 0 small and simple on II - VII, larger and simple or with

154 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

2 - 3 branches on VIII; 1-I unpigmented, undeveloped with filamentous branches,

or with poorly developed flattened leaflets; 1, on II - VII pigmented, with well

developed flattened leaflets and filaments; 5-II with 3 - 6 branches; 11-II long

and simple; 6-III with 18 or more branches; 6, on IV - V, 0.50 - 0.67 as long

as 6-III, with few branches from base; 6-VI very short, branched, less than

0.33 as long as 6 on IV - V; 13 on I - VII with 5 or more branches; stigmal

knob on median dorsal valve unpigmented, nearly transparent; pecten plate

with 11 - 16 teeth; 1-X with 2 - 4 (rarely 4) branches.

TYPE-DATA. The holotype 2 with associated larval and pupal skins

mounted on one slide is located in the USNM, and bears the following label

data: "Thailand, Chiang Mai, Muang, Doi Suthep, 05667-6, 10 Dec. 1969,

SEAMP Acc # 233.'' The allotype “ with associated larval and pupal skins on

one slide and a slide mounted genitalia (SEAMP Prep. #71/176) is also in the

USNM, and is number 05667-1 with the same data as above. An additional 4°

and 3@ in the USNM with the above collection data and different rearing num-

bers in collection 05667, have been selected as paratypes; these specimens

either have associated slide mounted larval and pupal skins or only a pupal

skin. Four slide mounted whole larvae with the above data were also selected

as paratypes to demonstrate the color banding pattern seen only on whole lar-

vae. Two 2 with skins (05667-3, 05667-7), one “ without skins (05667-8), one

o with a pupal skin (05667-103) and 2 whole larval slides were selected as

paratypes and sent to the BMNH. One & with skins (05667-10), one “ witha

pupal skin (05667-101) and 2 whole larval slides were selected as\paratypes and

sent to the Department of Medical Entomology, SEATO Medical Research Lab-

oratory.

There are 8 other slide mounted whole larvae of aberrans from this collec-

tion not selected as paratypes. These larvae are also in the USNM collection.

The collection (05667) from which the holotype, allotype and paratypes

were selected was made on Doi Suthep, a mountain just west of the city of

Chiang Mai in northern Thailand. The type-locality is a small stream between

600 - 640 m elevation on the mountain, just below the paved road going up to a

famous Buddhist temple. This collection was made by Captain Harrison from

small, nearly isolated, pools beside the stream, under the heavy shade of

secondary deciduous forest. (See additional biological data under biology sec-

tion. )

This species has been named aberrans for the banding pattern found on

most adults and larvae, which deviates from the usual unicolored appearance

of species in this group.

DISTRIBUTION (Fig. 83). Based on specimens in the USNM, aberrans

has a wide distribution in Thailand, but is apparently confined to forested foot-

hills and mountainous regions. These specimens were collected between 140 -

1,400 m elevation, above the coastal and many of the flat rice plain areas.

Specimens (740°, 8492, 180 larvae and 237 larval and pupal skins) in the USNM

were examined from the following provinces of THAILAND: Chanthaburi,

Chiang Mai, Chon Buri, Mae Hong Son, Nakhon Sawan, Phrae, Ranong, Songkh-

la and Trat. An intense search was made in the USNM and BMNH for speci-

mens of abervans from other countries. A series of 10 slides (larval skins)

in the BMNH from Peninsular Malaysia were found, some of which may be

this species. These slides have the following label data: "Fed. Malay States,

1920-454, Dr. H. P. Hacker", and individual rearing numbers. Skin num-

bers 483.D5, 483.D7, 483.Dg and 483. E10 in particular, had setae 8-C, 14-P

and 11-II that agree with abervrans. Unfortunately, there is no indication of

where in Malaya these were collected. None of the plates or descriptions

Harrison and Scanlon: Subgenus Anopheles in Thailand 155

presented by Reid (1965) suggest that aberrans occurs in Malaysia. However,

since there are specimens from Songkhla, in southern Thailand, and the im-

mature taxonomic characters are the most reliable, the above skins in the

BMNH probably represent Malayan aberrans. Further collecting in Southeast

Asia will probably reveal that this is a fairly widely distributed mainland

species.

The distribution of abervans in Thailand is similar to that of bengalensis,

consequently, these species are sometimes collected together. On the other

hand they may occur isolated, but very near to each other. Specific differences

in the habitat requirements for these 2 species are not currently known. The

following table will demonstrate known larval associations of abervans and

bengalensis.

TABLE 4. Collection associations of aberrans and bengalensis immatures

from Thailand provinces where both were found

Province aberrans only both species bengalensis only

Chanthaburi 22 10 8

Chiang Mai 2 2 2

Chon Buri 1 9) 2

Mae Hong Son 1 0 1

Phrae 0 1 2

Ranong 0 0 3

Songkhla 4 3 9

Trat 3 1 8

Total 33 23 35

TAXONOMIC DISCUSSION. Specimens of this species were apparently

_ first collected in Thailand by the Thurmans in the 1950's, but overlapping

larval and male genitalia characters with bengalensis (see aitkenii group)

kept it from being recognized as a distinct species. Almost invariably this

species has previously been identified as bengalensis, except for male speci-

mens with 3 outer dorsal claspette setae that were called aitkenii. This spe-

cies is obviously related to bengalensis, but not as closely as is fragilis.

Based on larval characters, abervans appears more closely related to fragilis

and bengalensis than to tigertti. Larvae of aberrans can be differentiated

from the first 2 species by having seta 11-II long and simple, 5-II with 3 -

156 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

6 branches and 14-P with 3 - 5 branches, while it differs from tigeviti larvae

by having 11-II long and simple, 5-II with 3 - 6 branches, 6-III with 18 or

more branches, 4-A with 5 or more branches and the stigmal knob on the medi-

an dorsal valve unpigmented. Larvae of aberrans key out to bengalensis in

Rattanarithikul and Harrison (1973: 5, couplet 9), but can be separated from

that species by the above stated characters.

The pupae of aberrans, bengalensis, fragilis and tigertti are also quite

similar. However, the pupa of aberrans can be differentiated from those of

bengalensis and fragilis by having only 2 - 4 branches on I-IV and having a

short refractile margin on the paddle, and can be differentiated from the pupa

of tigertti by having 6-III short and only about half as long as segment II, 1-II

at least as stout as 3-III and 10-IV about half as long as the lateral margin of

V. |

The male genitalia of aberrans are easily separated from those of tigertti

by the latter having spines on the aedeagus. Similarities in the male genitalia

of aberrans, aitkenii and bengalensis have already been noted (see aitkenii

complex). Apparently both aberrans and aitkenii can have 2 - 3 outer dorsal

setae on the claspette (see below) while bengalensis normally has 2. The

inner dorsal club onaitkenii, however, is made up of 2 longer flattened hood-

like setae, while those on abervans and bengalensis are approximately as long

as the outer dorsal setae. Futhermore, the shape of the aedeagus of aitkeni1

also appears distinct.

The pupae of aberrans, insulaeflorum and pilinotum have similar refractile

margins on the paddle, however, the very long sharp seta 9 on aberrans can-

not be confused with the short seta 9 on insulaeflorum and pilinotum, or the

long bluntly rounded seta 9 on palmatus. Pupae of aberrans should never be

confused with those of stvricklandi, for the paddle of the latter is very different

and seta 9 is even shorter than that of insulaeflorum.

Larvae of aberrans are easily separated from those of insulaeflorum, pal-

matus and stricklandi by the last 3 having the bases of setae 2-C much closer

together than those of 2-C and 3-C on one side.

Certain variable characters were noted which deserve further comment.

Male genitalia (claspette) variations are based on slide preparations of geni-

talia from 8 males confirmed by associated immature skins. The outer dor-

sal setae on the claspette exhibited the following differences: 2 specimens with

2 outer dorsal setae on both sides; one specimen with 2 outer dorsal setae on

one side and 3 on the other side; and 5 specimens had 3 outer dorsal setae on

both sides. The inner dorsal club had the following differences: one specimen

with inner dorsal club arising from a single basal stem, and 7 specimens with

the inner dorsal club arising from 2 separate basal stems.

The integument of adult abdominal segments IV - V was usually paler than

that of the other abdominal segments. Based on adults confirmed by associated

immature skins, the frequency of paler integument was: Chiang Mai 100%

(90° and 89), Chanthaburi 75.5% for males (40/53) and 82.5% for females (52/

63); too few specimens for valid sample from other areas, but 5 females from

Chon Buri, Nakhon Sawan, Phrae and Songkhla did not have this pattern. An

attempt to correlate this character with elevation, or some other environ-

mental factor, cannot be made without further specimens and more data from

the collection sites.

A banding pattern on the abdomen of the larvae is also variable. When

present, segments IV - V and X are paler than the others. This banding was

noted on the following collections of larvae: 100% (16/16) from Chiang Mai,

76% (93/122) from Chanthaburi and 33% (1/3) from Trat. Larvae from the

Harrison and Scanlon: Subgenus Anopheles in Thailand 157

following collections did not exhibit banding: Chon Buri (16), Nakhon Sawan

(1), Phrae (1), Ranong (1) and Songkhla (13). Larvae not having this color

pattern were otherwise identical to those having the bands..:

The presence of posterior tergal plates on larval segments IV - V seems

to be much more common on aberrans than bengalensis. While the latter only

infrequently had these plates on IV - V, abervans had plates on these segments »

on about 50% of the specimens sampled (more than 50).

BIOLOGY. There are no data regarding adult behavior for abervans. The

immature stages are found in habitats quite similar to those utilized by ben-

galensis, insulaeflorum and palmatus. Records indicate larvae have been

collected at elevations between 140 - 1,400 m from: stream margins, stream

pools, rock pools, springs, seepage pools and elephant footprints on stream

margin. These collections were made in light to heavy shade (usually heavy),

in cool clear water with dead leaves and sometimes roots and with the bottom

composed of silt or mud, not sand. The majority of collections came from

primary or secondary wet forests, with others from evergreen and secondary

deciduous forests. The requirements for this species seem to be more

closely associated with the forest and heavy shade, than do those for bengalen-

sis which can be found in more open-forest margin areas. This trend is

supported by records of larvae of aberrans being collected only with those of

bengalensis, insulaeflorum and palmatus. These 2 latter species are definitely

forest species.

The following observations were made during the original collection (05667)

at the type-locality. Collections were made in small shallow pools or indenta-

tions in the margin of a small rather swiftly flowing stream. The water in the

collection sites was usually no deeper than 2.5 - 4.0 cm and the bottom was

silty mud with no algae or vegetation except a few dead leaves. Larvae were

always associated with either rocks (usually), a dead root mass hanging in

the water or infrequently from holes left in the bank by rotten roots. The water

in the collection sites was always back up water with practically no current.

The larvae are quite noticeable on the water surface particularly when moving,

because of the pale pattern on the body. They apparently feed out in the open,

but are sensitive to light even though in partial to heavy shade, and quickly

dart back into a shelter such as under a rock or a small hole in the bank when

a shadow is cast across them.

ANOPHELES (ANOPHELES) BENGALENSIS PURI

(Figures 78, 79, 80, 81, 82)

Anopheles aitkenii var. bengalensis Puri 1930: 955 (0, L).

?Stethomyia pallida Ludlow 1905: 129 (¢) (non Theobald 1901).

Anopheles (Anopheles) aitkenii var. bengalensis Puri, Christophers 1933: 109

(2%; LF).

Anopheles (Anopheles) aitkenii bengalensis of Baisas 1946: 30 (P*, L*) (in

part). |

Anopheles (Anopheles) bengalensis Puri, Reid 1965: 113 (“*, 9, P*, L*, E*);

Reid 1968: 238 (*, 2, P*, L*, E*).

Anopheles (Anopheles) aitkenii of Reid 1965: 107 (“*, 2, P, L*, E) (in part);

Reid 1968: 31: 237 (**, 9, P, L*, E) (in part).

The adults of bengalensis cannot be identified without associated immature

skins. The pupal stage is distinguishable from all the other species except

158 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

fragilis, The larva is distinct from all the other group members found in

Thailand, but not on the basis of 2-C branching. Since this is the most com-

monly encountered member of the group in Thailand, we have decided to

describe it in detail and to use it as a guideline for the remaining species

descriptions.

FEMALE (Figs. 78, 80). Unicolorous brown. Head. Vertex with very

narrow erect scales, slightly expanded and notched at tip; interocular space

with short white scales, long frontal setae dark brown; clypeus without scales;

pedicel and flagellomeres without scales, antennal whorl setae 3 - 5 times

as long as segments; proboscis long slender, with decumbent scales; ratio-

forefemur/proboscis (sample, 25 females), range 0.89 - 1.24, mean 1. 04;

palpus very slender, approximately equal length of proboscis, with 2 apical

segments slightly swollen, scales decumbent. Thorax. Anterior promontory

without scales; scutal integument tan, usually with paler longitudinal line on

each side of slightly darker median (acrostichal) longitudinal line; scutum

without scales, with mixed long and short dark brown setae in anterior promon-

tory, acrostichal, dorsocentral, lateral prescutal, fossal, antealar and supra-

alar groups; with bare prescutellar space; scutellum with long and short dark

brown setae, without scales; anterior pronotum with sparsely scattered setae,

without scales; pleural area pale tan without scales; pleural setae: 0 - 2 pro-

pleural, 1 - 3 spiracular, 1 prealar, 1 upper and 2 - 4 lower sternopleural

and 3 - 5 upper and 0 lower mesepimeral. Wing. Unicolorous, with narrow

dark scales; humeral crossvein without scales; cell Mo, 0.1 - 1.0 length of

stem (from crossvein). Halter. With dark brown scales. Legs. Coxae with-

out scales, upper midcoxa with 3 - 5 setae; femur, tibia and tarsomeres long,

slender and entirely dark scaled. Abdomen. Unicolorous brown with brown

setae, without scales.

MALE (Figs. 78, 79, 80). As for female except antennae with more

and longer whorl setae; palpus with 2 apical segments distinctly flattened and

club-like. Genitalia. Basimere without scales, with 2 parabasal spines on

tubercle; ventromesal spine inserted near distimere on distal 0.25 of basimere;

claspette with 2 lobes; dorsal lobe with 2 separate outer setae, inner club

formed from 1 - 2 separate basal stems; ventral lobe usually with 3 large and

one small setae, and numerous minute spicules; aedeagus bare, 3 or more

times as long as width at base, tapering to rounded tip about 0.5 or less width

of base. : |

PUPA (Fig. 81). Integument unicolorous pale to light tan. Cephalothorax.

Seta 7-C with 2 - 6 branches. Trumpet. Simple, with deep meatal cleft,

meatus less than 0.33 as long as trumpet. Abdomen. Seta 0 small and simple;

1-IV with 3 - 12 branches; 1-VI with 2 - 7 branches; 1-VII with 3 - 9 branches;

5-I well developed, much longer than 2 and 3-I; seta 5 more strongly developed

than seta 1 on V - VII, approximately 0.50 - 0.75 as long as segments; 9-IV

without strong central stem, with 3 - 7 branches from near base; 5-VII with

6 - 13 branches; seta 9 long and narrow with sharp tapered point on IV - VII;

9-IV, 9 or more times as long as 9-IIJ; 9-VII, 0.33 - 0.50 as long as segment

VIll. Paddie. Refractile margin 0.60 - 0.75 as long as paddle; fringe hairs

approximately 0.5 length of 1-P; outer distal margin evenly convex, without

straight section; 1-P simple and stout; 2-P with 2 - 6 branches.

LARVA (Fig. 82). Unicolorous tan or gray. Head. Antenna base slightly

wider than tip; antenna with long spicules on basal 0.67, particularly on mesal

side; seta 1-A short with 4 or more branches, inserted on basal 0.20 - 0.25

of antenna; 2, 3-A with one edge serrate; 4-A with 8 or more branches; setae

2-C with bases separated by approximately same distance as bases of 2-C and

Harrison and Scanlon: Subgenus Anopheles in Thailand 159

3-C on one side; 2-C rarely simple, usually with 2 - 9 branches originating

0.25 - 0.75 way along stem, branches equal or nearly equal length; 3-C approxi-

mately 0.25 - 0.38 as long as 2-C, usually split into 2 - 5 branches on distal

half, rarely simple; setae 4-C slightly wider apart than setae 3-C, with 2 - 7

branches from near base; 5, 6, 7-C long, well developed and plumose, 5-C

reaching forward nearly to alveoli of 2-C; 8-C with 2 - 8 branches, rarely 2;

9-C with 3 - 7 branches; 11-C approximately equal length of antenna, with

many branches particularly at apex. Thorax. Seta 1-P with 7 - 15 branches,

about 0.5 length of 2-P, without sclerotized base; 2-P with 8 - 18 branches,

arising from sclerotized tubercle; 2-P tubercle with prominent apical spur or

tooth; 3-P short and simple, closer to 2-P than 1-P is to 2-P; 4-P with stout

stem and numerous short lateral and apical branches, slightly longer than 2-P;

prothoracic pleural tubercle with distinct slender spine; 11-P with 2 - 5 fine

distal branches; one long pleural seta on pro-, meso- and metathorax often

with 2 - 3 distal branches, frequently these setae simple on one or more seg-

ments; 13-P with 4 - 14 dendritic branches; 14-P with 5 - 9 branches, rarely

2 on both sides; 4-M small and branched from base, inserted caudally to long

and simple 3 and 5-M; 3-T with well developed flattened leaflets without fila-

ments. Abdomen. All segments unicolorous; anterior tergal plates 0.25 -

0.33 width of segment on I-VII; posterior tergal plates small, usually only on

VI - VII, infrequently on IV - VII; seta 0 small with 2 or more branches on II -

VII, larger with 4 or more branches on VHI; 1-I unpigmented, undeveloped or

with poorly developed flattened leaflets; 1-II pigmented, with well developed

flattened leaflets, with or without well developed filaments; 1, on III - VII pig-

mented, well developed (palmate) with filaments; 5-II with 7 or more branches;

11-II with 2 - 4 branches on distal half; 6-III with 20 or more branches; 6, on

IV - V, 0.50 - 0.67 as long as 6-III; 6-VI short, less than 0.33 as long as 6,

on IV - V; seta 13 with 5 or more branches on I-VIII; stigmal knob on median

dorsal valve unpigmented, nearly transparent; pecten plate with 11 - 16 teeth;

1-X with 3 - 8 branches.

TYPE-DATA. A type “ and 2 with the “ genitalia mounted on a slide

were deposited in the BMNH according to Puri (1930). Unfortunately, these

specimens have not been found. Reid (1965, 1968) lists paratypes as present

in the Indian Museum, Calcutta and the National Institute of Communicable

Diseases, Delhi. The type-locality of bengalensis is listed as Marianbarie

(Bengal Terai), near Sukna in West Bengal, India. The ? type of pallida Ludlow,

is located in the USNM and has the following label data: | ''H'', Stethomyia

pallida Ludlow, Camp Stotsenberg, Maj Whitmore. |. This specimen has a

forefemur/proboscis ratio of 1.15, which fits in the range known for bengalen-

sis, but is outside that known for aberrans.

DISTRIBUTION (Fig. 81). Since the immature stages are currently the only

means of identifying this species, the following distribution is based entirely on

either whole larvae, pupae or adults with associated skins. This may be the

most widely distributed species in this group. The confirmed distribution ex-

tends from Assam in the west to Amami Island (Japan) in the northeast and

south through Malaysia to the Philippines. Specimens (450, 6492, 271 larvae

and 174 larval and pupal skins) in the USNM were examined from the following

provinces of THAILAND: Chanthaburi, Chiang Mai, Chon Buri, Kanchanaburi,

Krabi, Lampang, Mae Hong Son, Nakhon Nayok, Nakhon Ratchasima, Nan,

Narathiwat, Phangnga, Phrae, Phuket, Prachin Buri, Ranong, Sara Buri,

Satun, Songkhla, Trang and Trat. Additional specimens (8%, 209, 88 larvae

and 60 larval and pupal skins) in or on loan to the USNM were examined from

BURMA, HONG KONG, INDIA (Assam), MALAYSIA, PHILIPPINES, JAPAN

160 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

(Amami Island) and SOUTH VIETNAM. A number of adults in the BMNH

labeled bDengalensis could not be confirmed, however, specimens (5 larvae and

9 larval and pupal skins) were examined from CHINA (Formosa), HONG KONG,

MALAYSIA (Peninsular, NORTH VIETNAM and PHILIPPINES. —

- Kanda and Kamimura (1967) recorded bengalensis from Japan (Amami

Island), which is apparently the most northern record (27 - 28° N) for this

group. We were able to borrow 2 adults with associated immature skins from

this island, and all stages agree very well with bengalensis.

There are published records of this species in Indonesia, and some of

those listed under aitkenii by Reid (1965) probably refer to bengalensis. Al-

though we could not confirm adults identified as this species from Indonesia,

we feel that it most likely occurs there, probably on the western islands of

Java and Sumatra.

No specimens from Bengal (type-locality) were available, but numerous

larvae from Assam are in the USNM. These larvae all agree with the ben-

galensis concept, but the bifurcate variation of 2-C is apparently more com-

mon, possibly giving rise to numerous records of aitkenii in Burma (Khin-

Maung-Kyi 1971a). See the taxonomic discussion (aitkenii group) for more

information on this variation.

Reid (1968) describes bengalensis in Malaysia as a hill or mountain

species that is uncommon or replaced by fragilis at elevations below 150 m.

In Thailand, bengalensis has been collected at elevations as low as 10 m, and

apparently utilizes suitable habitats at low elevations regularly. The lack of

collections from northcentral and northeastern Thailand (Fig. 81) almost

certainly reflects limited collecting efforts in those areas rather than the

actual distribution of bengalensis, which is probably throughout most of the

country where suitable habitats occur. However, most of the suitable habitats

seem to be hill or mountainous areas where man has not drastically altered

the ecology. Since most ground water habitats in the plains or low lands are

polluted, turbid and have higher temperatures, bengalensis is usually absent

or uncommon in those areas. (Also see biology section).

TAXONOMIC DISCUSSION. This species is very closely related to fragilis,

in fact, sibling rearings are needed to confirm the validity of the only imma-

ture character (larval seta 2-C) to separate these 2 in Thailand. We identified

several larvae from southern Thailand as bengalensis with 2-C either bifurcate

or simple. These specimens could possibly be fragilis without frays on 2-C.

Reid (1965, 1968) said that pupae of bengalensis and fragilis in Malaya could be

separated by fragilis having the following setal branches: 1-IV (4 - 10), 5-IV

(5 - 7), 1-VU (3 - 7), 5-VII (6 - 13) and 7 of the cephalothorax (3 - 7); while,

bengalensis has 1-IV (8 - 7), 5-IV (3 - 5), 1-VII (1 - 5), 5-VII (8 - 13) and 7

of the cephalothorax (2 - 3). As seen in the above description, branching

variation on Thai bengalensis completely nullifies these differences. Unfortun-

ately, we were unable to find any other differences, thus, the pupae of /vagilis

and bengalensis cannot be differentiated in Thailand. The male genitalia of

these 2 species are also very similar. Previous authors describe the inner

dorsal club on the claspette of fragilis with a single stem, while that of ben-

galensis was supposed to have 2 basal stems. Of 13 genitalia preparations

we examined from bengalensis confirmed by immature skins, 10 had this

structure coming from 2 basal stems and 3 had a single basal stem.

During recent years 2 new species (tigeriti and aberrans) have been found

hidden among Thailand specimens previously considered bengalensis. The

pupa, larva and male aedeagus of tigervtti are very distinct from those of

bengalensis. Moreover, tigertti is a crab hole inhabitant, and not frequently

Harrison and Scanlon: Subgenus Anopheles in Thailand 161

encountered. The larvae and pupae of abervans are also very distinct from

those of bengalensis, but the best larval characters are hard to see. The

nearly identical development of head seta 2-C on aberrans and bengalensis,

kept aberrans hidden for several years. The close affinities of these 2 spe-

cies is emphasized further by overlapping male genitalia characters, which

invalidate previously used outer dorsal setal counts on the claspette. Fortun-

ately, some adults and larvae of aberrans can be distinguished by the presence

of pale abdominal segments, a character not seen on bengalensis. These 2

species are often found together (see Table 4, p. 155), and workers in Thai-

land should be very careful with their identifications of ''bengalensis".

Reid (1965, 1968) listed 2 characters by which he differentiated his single

Malayan female of aitkenii (with skins) from fragilis and bengalensis, they

are: pupa 1-IV with 11 branches; and position of fork on larval seta 2-C.

After examining Reid's specimen of aztkenii from Peninsular Malaysia in the

BMNH, we consider it equivalent to bengalensis. Characters checked on the

pupal skin, 1-IV (11 branches), 1-VII (6 - 7 branches), paddle refractile bor-

der 65 - 70%, paddle apical fringe 0.5 length of 1-P and 2-P with 3 branches,

all fit our concept of bengalensis. Characters checked on the larval skin, 8-C

(6 - 7 branches), 14-P (5 branches each side), 5-II (10 or more branches) and

11-II (bifurcate distally), also agree with bengalensis. The female reared

from these skins has a definite prescutellar bare space and does not exhibit

pale abdominal segments.

Reid also mentioned variations he observed in aitkenii characters. He

noted that the bifurcation of larval 2-C on Indian specimens was usually very

low, about 0.25 from the base, and Indian larval 3-C are shorter than those

on specimens from further east. The 2 larvae of aitkenii from Sri Lanka have

2-C bifurcate about 0.33 from the base and 3-C on both specimens is long,

reaching or extending beyond the fork on 2-C. Furthermore, Reid followed

Puri (1931) and tried to attach some significance to the branching of the long

larval pleural setae on aitkenii. Branching patterns on these setae in bengal-

ensis are extremely variable in Thailand, and apparently also on aitkenii as

the 2 larvae from Sri Lanka have branches on the long meso- and metapleural

seta, while Puri (1931) says aitkenii only has such branches on the long pro-

thoracic pleural setae. The branching patterns on the long larval pleural

setae are probably too variable in this group to be used as reliable taxonomic

characters.

Although numerous variations were observed, only one obvious anomaly was

found on a larva from Perak, Malaysia. At first glance this specimen appeared

to have 2 left setae 3-C, however, further examination revealed the 2nd (also

most caudal) 3-C was branched differently and was a mislocated 4-C. No alveo-

lus or seta could be found in the location where the normal left 4-C should oc-

cur.

BIOLOGY. Our knowledge of the biology of bengalensis is largely based

on habitats where the immature stages are found. This species definitely pre-

fers forests, shade and clear, cool slow running streams or seepage springs

for oviposition sites. Although immatures are found in isolated rock pools

and stream pools, these are usually very close to either water falls or streams

where slight fluctuations in water level can inundate or bring fresh water to

the pools. Dead leaves, emergent or floating debris or vegetation are also

abundant in most collection sites. Forest-type apparently has no basic effect

on populations of bengalensis, as long as the shade and water conditions are

satisfactory. Larvae have been collected from habitats in primary and secon-

dary wet forests, primary evergreen forest, primary and secondary deciduous

162 Contrib. Amer. Ent. Inst. vol. 12, no. 1, 1975

forests, bamboo groves and heavily shaded habitats near villages. Immatures

have been collected between 10 - 1,300 m elevation from the following sites in

Thailand: marsh-bogs, elephant-buffalo tracks on stream margins, stream

pools, stream margins, springs, seepage pools, rock pools, and a small stream

inside a dark cave (near entrance)(see distribution section).

Records (Scanlon and Esah 1965, Scanlon et al. 1968 and Harinasuta et al.

1970) of female aitkenii or bengalensis biting man in Thailand now need con-

firmation because of our revised interpretation of aitkenii distribution, and the

descriptions of abervans and tigertti. Although bengalensis is apparently the

most commonly collected species of this group in Thailand, there are no con-

firmed specimens that were collected biting man. Records of aitkenii group

females collected biting man should be confirmed by allowing the females to

oviposit and rearing the larvae, or, if they cannot be identified by the charac-

ters presented here, should be called "aitkeniz group".

One observation has possible value for future larval control measures. In

one collection of bengalensis larvae from Prachin Buri Province, 21% (20/94)

of the larvae were infected with a Coelomomyces fungal parasite. This para-

site does not appear to be C. indiana Iyengar or C. anophelesica Iyengar (R.

A. Ward - personal communication), but since all specimens were mounted on

slides, identification cannot be taken further.

ANOPHELES (ANOPHELES) FRAGILIS (THEOBALD)

(Figeures 79, 83, 84)

Stethomyia fragilis Theobald 1903: 257 (<).

Anopheles treacheri Leicester 1908: 19 (“, 2); Reid 1965: 109 (= fragilis).

Anopheles aitkeni stantoni Puri 1949: 483; Reid 1965: 109 (= fragilis).

Anopheles aitkeni treachervi of Puri 1949: 483; Reid 1965: 109 (= fragilis).

Anopheles (Anopheles) fragilis (Theobald), Reid 1965: 109 (“*, @*, P*, L*);

Reid 1968: 232 (*, 9*, P*, L*).

The adults of this species cannot usually be identified without an associated

larval skin. The pupa is not separable from that of bengalensis. The larva is

distinct on the basis of one character, frayed setae 2-C. Like bengalensis,

except:

FEMALE. Cannot be separated from bengalensis.

MALE (Fig. 79). Habitus like female, indistinguishable from bengalensis.

Genitalia. As for bengalensis except club on dorsal lobe of claspette arising

from single stem (see bengalensis variation).

PUPA (Fig. 83). Cannot be separated from bengalensis, but with slight

differences in setal branching. Cephalothorax. Seta 7 with 2 - 7 branches.

Abdomen. Seta 0 simple or often bifid or trifid, particularly on VI - VII; 9-IV,

8 times or more as long as 9-III; 9-VII, 0.33 - 0.50 length of segment VIII;

1-IV with 4 - 10 branches; 5-IV with 5 - 9 branches; 1-VI with 3 - 6 branches;

1-VII with 3 - 7 branches; 5-VII with 6 - 13 branches. Paddle. Refractile

margin 0.60 - 0.75 as long as paddle; 2-P with 3 - 4 branches.

LARVA (Fig. 84). Nearly identical to bengalensis. Head. Seta 1-A with

4 - 10 branches; 2-C simple or with 2 - 5 nearly equal branches from 0.33 -

0.50 along stem, middle 0.33 of stem(s) with tiny fray-like branches; 3-C

simple or with 2 - 3 distal branches, about 0.33 - 0.40 as long as 2-C; 9-C

with 3 - 8 branches; 14-P with 4 - 8 branches; pecten plate with 9 - 13 teeth.

TYPE-DATA. Theobald designated 2 “ as types for fragilis and both are

Harrison and Scanlon: Subgenus Anopheles in Thailand 163

in the BMNH. One of these syntypes is in near perfect condition and with the

genitalia intact. This specimen has the following data: (1st label, underside of

cardboard minuten stage) - 'K. Lumpur, Drives, 8. XII. 02''; (2nd label) -

"Kuala Lumpur Dr. N. E. Durham"’; (3rd label) - circular BMNH ''Type" label;

and (4th label) - "Stethomyia fragilis (Type) FVT''. The 2nd specimen is not

complete and has certain parts mounted on 3 slides. The remaining portion of

the pinned adult has the following data: (1st label, underside of cardboard

minuten stage) - 'K. Lumpur W. Sink, I. 03"'; and (2nd label) - ''Kuala Lumpur

Dr. H. E. Durham". The dates on the above adults correspond to the January

and December dates given by Theobald. The 3 slides belonging to the last

male were incorrectly labeled by Theobald, but J. A. Reid has corrected this

by a hand written note on the slide containing the abdomen and genitalia.

At the same time Reid (1965) elevated fragilis to specific status, he synony-

mized treachervii Leicester under fragilis. Reid (1965) designated one male

from 5 Leicester syntypes in the BMNH as the lectotype of treacherit. This

male has the genitalia mounted on a slide, and has the following data: (1st

label, underside of cardboard minuten stage) - ''Hill Stream Ampang jungle

26/8/03" (not 28/8/03 as in Reid, p. 109); (2nd label) - ''Ac"''; (3rd label) -

circular BMNH lectotype label; (4th label) - ''Fed. Malay States Dr. G. F.

Leicester. 1912-350"; and (5th label) - "Anopheles treacherii (lectotype) Leic."'

The lectotype genitalia is labeled "Anopheles treacherii Leic. 1908 Lectotype

Oo genit. prep. JR. 15. VI. 64 Lectotype", and agrees very well with the genita-

lia of the fragilis type.

We concur with Reid that treachevii is probably a synonym of fragilis.

However, as discussed under the aitkenii group and bengalensis, male genitalia

in this complex are now known to be much less stable than previously thought,

and too much reliance should not be placed on them. Puri (1949) based his

description of subspecies stantoni on illustrations by several workers, and

apparently, did not designate a type.

DISTRIBUTION (Fig. 83). Although this species is considered the most

common member of the group in the lowlands of Peninsular Malaysia, it

apparently is not common in Thailand. Reid (1965) suggested that fragilis

probably occurred in Thailand, and it was first recorded there by Peyton and

Scanlon (1966). Scanlon et al. (1968) listed larvae collected in Narathiwat,

Songkhla and Yala provinces. Specimens (4 larvae) in the USNM were examined

from the following provinces of THAILAND: Nakhon Si Thammarat, Narathiwat

and Songkhla. Other specimens (40, 8°, 9 larvae and 18 larval and pupal skins)

in the USNM were examined from MALAYSIA and PHILIPPINES. Specimens

80°, 592, 1 larva and 12 larval and pupal skins) in the BMNH were examined

from MALAYSIA (Peninsular) and PHILIPPINES, and included the syntypes

of fragilis and lectotype of treacherii. Swellengrebel and Swellengrebel-de

Graaf (1919) illustrated larvae of this species from Indonesia, and Reid (1965)

refers to an obscure reference of larvae with frayed setae 2-C in Burma. We

have not been able to confirm these latter records, but feel that fvagilis pro-

bably occurs in both these countries. Apparently fragilis is a Malaysian-

Indonesian species which reaches into southern Burma in the tropical wet

forest areas. This species is probably more common in the foothill streams

of southern Thailand than in the flat coastal areas. Reid (1968) said fragilis

is uncommon in the flat coastal plain of Malaysia.

TAXONOMIC DISCUSSION. This species is almost identical to bengalensis,

so much so that F, sibling rearings will be needed to help resolve the status of

these species. The only 2 characters we consider of value in separating these

2 species are larval seta 2-C and the male claspette setae (see discussion under

164 Contrib. Amer, Ent. Inst., vol. 12, no. 1, 1975

aitkenii group and bengalensis). The variation of fraying and branching on

2-C of larvae from Malaysia and Thailand compared with those from the

Philippines, suggests that overlap may occur in these characters between

bengalensis and fragilis. Of the 9 Thai-Malay larvae examined in the USNM,

5 had 2-C single, 3 had 2-C bifurcate and one had 2-C trifurcate. Those with

a simple or bifurcate 2-C had dense fraying, while there was sparse fraying

on the trifurcate 2-C. Of 14 Philippine larvae and larval skins examined in

the USNM, 11 had 2-C trifurcate and 3 had 2-C with 4 branches. Fraying on

the 2-C of Philippine specimens was not dense and was limited to only 1 - 3

tiny frays on some setae 2-C that were trifurcate and on all that were 4 bran-

ched. On 2 of the Philippine specimens with 4 branched 2-C, only one 2-C

per larva exhibited fraying. This reduction of fraying with an increase in

large branches, coupled with larvae that only have 1 - 2 frays on 1 of 2 setae

2-C, is highly suggestive that the parameters of 2-C currently used to separ-

ate bengalensis and fragilis larvae (at least in the Philippines) may not be

valid.

Data presented under bengalensis show that some specimens of that species

have male claspette characters that are identical to those of fragilis. The ex-

tent of variation for fragilis claspette characters is not known. Unfortunately,

we had very few male fragilis and these came from the Philippines, which

might explain the slight differences between our illustrated claspette (Fig. 79)

and the illustrations in Reid (1965, 1968).

Although the above morphological characters appear to overlap, there is

other very valid evidence for continuing to consider bengalensis and fragilis

as distinct species. Their distributions are different, particularly in an area

such as Malaysia and Thailand where there has been considerable collecting.

In Thailand, fragilis has been collected only from southern tropical wet forest

areas, while bengalensis is found in most forested areas throughout the country.

The latter species is well adapted to habitats found in countries (e.g., China,

North and South Vietnam and Thailand) with distinct wet and dry seasons,

while fragilis apparently utilizes habitats found only in or closely associated

with tropical wet forest where yearly rainfall patterns are more evenly dis-

tributed. Further evidence supporting the validity of these 2 species is the

apparent abundance of fragilis in Malaysia at elevations below 300 m, while

bengalensis is the more common species above that elevation (Reid 1965,

1968).

BIOLOGY. Reid (1968) listed Malaysian larvae of fragilis as found in

streams, pools and occasionally swamps under shade in or near forests. He

also noted they are usually found on rising ground and are not common on the

flat coastal plain. The few Thai larval collections of fragilis agree with Reid's

information: 154 - 185 m elevation; moderate to heavy shade, slow running

seepage water, stream margin, stream pool and an elephant footprint (pre-

sumably beside a stream), cool clear water with dead leaves and/or erect

vegetation and collections in or very near tropical wet forests. Nothing is

known about adult habits.

ANOPHELES (ANOPHELES) TIGERTTI SCANLON AND PEYTON

(Figures 79, 85, 86)

Anopheles (Anopheles) tigertti Scanlon and Peyton 1967: 19 (“*, ¢, P*, L*);

Rattanarithikul and Harrison 1973: 5 (L*).

Harrison and Scanlon: Subgenus Anopheles in Thailand 165

This is a seldom encountered forest species in Thailand, with the immature

stages apparently restricted to fresh water crab holes. The species can be

differentiated in all known stages except the adult female. The egg is undes-

cribed. Like bengalensis except:

FEMALE. Cannot be separated from bengalensis at present. Ratio fore-

femur/proboscis (2 females) range 0.89 - 0.96, mean 0.93; pleural setae: 1

propleural, 2 spiracular, 1 prealar, 2 upper and 2 lower sternopleural and

3 - 4 upper and 0 lower mesepimeral; upper midcoxa with 2 setae.

MALE (Fig. 79). Habitus like female, currently indistinguishable from

bengalensis. Genitalia. Claspette with 2 outer dorsal setae, inner dorsal

club formed from one or usually 2 basal stems; ventral lobe of claspette with

one large seta next to dorsal lobe and 2 smaller more ventral setae; aedeagus

with sharp tip and scattered small spines on distal half.

PUPA (Fig. 85). Integument pale and unicolorous. Cephalothorax. Seta

7-C with 2 - 4 branches; 12-C with 5 or more branches. Trumpet. Simple

with deep meatal cleft, meatus less than 0.33 as longas trumpet. Abdomen.

Seta 0 simple on all segments; 1-III weakly developed, not as stout as 3-III;

1-IV with 2 - 4 branches; 1-VI simple or with 2 - 3 branches; 1-VII simple or

with 2 branches; 5-I weakly developed and short, approximately equal or shorter

than 2 and 3-I; setae 5, on IV - VII well developed and very long, usually as

long as or longer than segments; 6, onI - III very long; 6-II nearly equal length

of segment III; seta 9 long and narrow with sharp tapered point on IV - VII;

9-IV, 10 or more times as long as 9-II; 9-VII, 0.33 - 0.50 as long as segment

VIII; seta 10 on IV - V very long, 10-IV nearly equal length of segment V. Pad-

dle. Refractile margin short, 0.33 - 0.50 as long as paddle; outer margin

evenly convex on distal half; fringe hairs 0.67 or more as long as 1-P; 2-P

simple or with 2 - 4 branches.

LARVA (Fig. 86). Head. Seta 1-A with 5 - 11 branches; 4-A simple or

with 2 - 4 branches; 2-C with bases only slightly closer together than bases of

2-C and 3-C on one side; 2-C with 2 - 5 branches originating 0.50 - 0.67 way

along stem; 3-C simple, rarely bifurcate distally; 4-C with 2 - 4 long branches;

8-C simple or with 2 - 3 branches. Thorax, 1-P with 8 - 10 branches; 11-P

simple or with short stem and 2 - 3 distal branches; 14-P with 4 - 7 branches;

one long pleural meso- and metathoracic seta bifid or trifid distally; 3-T with

well developed flattened leaflets, no filaments. Abdomen. Anterior tergal

plates small, approximately 0.25 or less width of segments I - VII; posterior

tergal plates absent; seta 0 small and simple on all segments; 1-I usually with

filamentous branches, rarely with poorly developed slightly flattened branches;

1-II with well developed leaflets, no filaments; 1, on III - VII well developed

with filaments; 5-II with 6 - 12 branches; 11-II long and bifurcate; 6-III with

6 - 12 branches; stigmal knob on median dorsal valve of spiracular apparatus

pigmented, as dark as remainder of plate; pecten plate with 14 - 18 teeth; 1-xX

stout and longer than saddle, simple or with 2 - 3 distal branches.

TYPE-DATA. The holotype “ with associated immature skins, one ~ para-

type with pupal skin and 2 paratype whole larvae are deposited in the USNM.

One paratype ©, one paratype ¢ and 2 paratype whole larvae are located in the

BMNH. All of the above “ have genitalia preparations on slides. The holotype

o has the following label data: (1st label) - '00712-1, THAILAND: Prachin-

buri: Ban Bu Phram: 150 m. crab hole"; (2nd label) - ''Forest stream 20 Jan.

66, Peyton and Kol"; (3rd label) - "Holotype No. 69239"; and (4th label) - ""An-

opheles (An.) tigertti Scanlon and Peyton 1967."'

DIST RIBUTION (Fig. 85). Anopheles tigertti is known only from Thailand.

Although this species is seldom encountered, it apparently has a wide distribu-

166 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

tion in Thailand, which seems directly correlated with tropical wet forests and

restricted to areas having holes of fresh water crabs. The wide distribution

in Thailand suggests that fzgertti also probably occurs in adjacent Southeast

Asian countries. Vertical distribution is known from collections between 150 -

700 m elevation. More specific collection efforts aimed at crab hole habitats

are needed to understand the distribution of this species. Specimens (40, 29,

9 larvae and 6 larval and pupal skins) in the USNM (including the holotype) were

examined from the following provinces of THAILAND: Chanthaburi, Nakhon

Ratchasima, Prachin Buri and Tak. Specimens (1°, 19 and 2 larvae) in the

BMNH were examined from Prachin Buri Province.

TAXONOMIC DISCUSSION. Adult females of tigertti cannot be differentiated

at present, but adult males can be identified on the basis of genitalia charac -

ters. The small spines on the aedeagus and the 2 outer dorsal setae on the

claspette make tigervtti easy to differentiate from the other members of the

eroup, except for insulaeflorum. This latter species also has small spines

on the aedeagus, but it usually has 3 outer dorsal setae on the claspette, and

the scutum has fine setae in the prescutellar area almost back to the scutellum,

while tigertti has a large prescutellar bare area.

Except for aberrans, the pupa of tigeriti is easily separated from the other

Thailand species in this group by having long sharp setae 9 in combination with

a short refractile border on the paddle. The separation of tigerttz pupa from

those of aberrans requires closer examination. Pupae of tigeritz have setae

6-III and 10-IV much longer than those of aberrans and 1-III of tigertti is weak-

ly developed. Also, seta 5, on IV - V of tzgevtti is usually as long as or long-

er than the segments, while this seta on abervans is usually not this long. This

latter character is subject to more variation than the others listed above and

should only be used in conjunction with the other characters.

The larvae of tigertti are easily recognized by the few branches on 6-III

coupled with the widely spaced bases of setae 2-C. Only 2 other species in

Thailand, i.e., insulaeflorum and stricklandi, have seta 6-III with less than

15 branches, and they have the bases of setae 2-C very close together. The

other species, aberrans, bengalensis, fragilis and palmatus, have 2-C wide

apart (except palmatus), but have 18 or more branches on 6-III (usually 20 or

more). The pigmented stigmal knob on the median dorsal valve of the spiracu-

lar apparatus is also indicative of this species, but difficult to see on most

specimens. The position of branching for seta 2-C is also of value in identify-

ing tigertti larvae. This seta branches between 0.50 - 0.67 of the way along

the stem, a character found elsewhere in this complex only on occasional larvae

of bengalensis. Infrequently, seta 4-A with few branches and seta 0 simple on

the abdominal segments, may be of assistance in identifying tigertt: larvae.

A further study of the holotype of tigertti and other specimens in the USNM

has revealed several structures described and/or illustrated in the original

description that need modification. The inner club-like seta on the dorsal lobe

of the claspette (harpago) was originally described with a single stem; however,

the holotype has one side with a single stem and the other with this structure

having 2 basal stems. The 2 paratypes (USNM and BMNH) have a single club-

like seta on each side arising from 2 basal stems. The pupal trumpet was

originally described with a deep cleft, but illustrated for the original descrip-

tion in error with a very shallow cleft. In general, the chaetotaxy originally

illustrated for the larva and pupa had the setal thickness exaggerated, con-

sequently, we have reillustrated the species. _

BIOLOGY. The only biological data for tigertii concerns the immature

stages. Apparently females select only the holes of fresh water crabs for

Harrison and Scanlon: Subgenus Anopheles in Thailand 167

oviposition, thus, immatures are rarely collected. The identity of the crab(s)

associated with tigeviti is not known. Immatures of tigertii have been collec-

ted in Thailand between 150 - 700 m elevation in wet forests with heavy shade,

and only from crab holes in stream banks, dry stream beds or in seepage bogs.

Immature collections have been made in 4 Thailand provinces (Fig. 85), a

distribution that extends across central Thailand approximately 575 km from

Tak (adjacent to Burma) to Chanthaburi (adjacent to Cambodia). This distribu-

tion suggests that tigertti probably occurs in countries adjacent to Thailand.

The dependence and association of tigeritti with the crab hole environment is

supported by the other mosquito species that have been collected with it, they

are: Culex (Lophoceraomyia) pholeter Bram and Rattanarithikul, Culex (Lopho-

ceraomyia) sp., Uranotaenia koli Peyton and Klein, Uvanotaenia spiculosa

Peyton and Rattanarithikul and Uranotaenia species undescribed. The imma-

tures of these species are also found only in crab holes. The reader wishing

to read more on this habitat and its arthropod (or mosquito) associates should

refer to Hogue and Bright (1971) and Bright and Hogue (1972).

ANOPHELES (ANOPHELES) INSULAEFLORUM

(SWELLENGREBEL AND SWELLENGREBEL-DE GRAAF)

(Figures 78, 79, 87, 88)

Unclassified larva No. 1 of Swellengrebel and Swellengrebel-de Graaf 1919:

23 (L*); Swellengrebel and Swellengrebel-de Graaf 1920a: 2 (= Stethomyia

aitkenii var. insulaeflorum). |

Stethomyta aitkentt var. insulaeflorum Swellengrebel and Swellengrebel-de

Graaf 1920a: 2; Swellengrebel and Swellengrebel-de Graaf 1920b: 81 (L*,

and distribution in part); Swellengrebel and Swellengrebel-de Graaf

1920c: 34 (L* and distribution in part); Swellengrebel and Swellengrebel-

de Graaf 1920d: 98 (distribution in part).

Anopheles (Anopheles) aitkenti var. insulaeflorum of Christophers 1924: 19

(distribution in part); Strickland and Chowdhury 1927: 26 (L*, and dis-

tribution in part); Swellengrebel and Rodenwaldt 1932: 111 (distribution

in part).

Anopheles (Anopheles) insulaeflorum of Puri 1930: 954 (o%, L): Puri 1931: 101

(L*); Edwards 1932: 38 (distribution in part); Christophers 1933: 111

(distribution in part); Simmons and Aitken 1942: 128 (distribution in part):

Russell, Rozeboom and Stone 1943: 115 (distribution in part); Knight,

Bohart and Bohart 1944: 7 (distribution in part); Lee and Woodhill 1944:

97 (distribution in part); Bohart 1945: 15 (distribution in part); Puri 1949:

485 (distribution in part); Worth 1953: 174; Bonne-Wepster and Swellen-

grebel 1953: 97 (distribution in part); Senevet and Andarelli 1955: 331

(L); Stone, Knight and Starcke 1959: 20 (distribution in part); Reid 1963:

113; Reid 1965: 119 (L*, and distribution in part); Peyton and Scanlon

1966: 3 (2*); Scanlon, Peyton and Gould 1968: 13; Reid 1968: 248 (L*,

and distribution in part); Nguyen-Thuong-Hien 1968: 129 (?*, L*); Basio

1971: 36 (distribution in part); Rattanarithikul and Harrison 1973: 5 (L*);

Harrison and Scanlon 1974: 36 (distribution).

This species can be identified in all known stages. The egg is unknown.

Males and females can be recognized on the basis of a scutal character, and

the male has distinct genitalia characters. The pupa is illustrated here for

the 1st time and is easily recognized by having short setae 9. The larva is

easily differentiated by setae 2-C, 6-III and a developed seta 1-I. This species

168 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

is like bengalensis except:

FEMALE (Fig. 78). General appearance unicolorous as for most other

members of group unicolorous. Thorax. Prescutellar space with small fine

setae extending back near to posterior margin; pleural setae: 1 - 3 propleural,

1-6 spiracular, 1 - 2 prealar, 1 - 2 upper and 2 - 3 lower sternopleural and

2 - 6 upper and 0 lower mesepimeral; upper midcoxa with 1 - 3 setae.

MALE (Figs. 78, 79). Habitus like that of female. Genitalia. Claspette

with 3 - 4 separate flattened outer dorsal setae, slightly shorter than inner

dorsal club; inner dorsal club long, formed from 2 basal stems; ventral lobe

with 3 - 4 long stout setae, most mesal large seta longest, longer than inner

dorsal club; ventral lobe also with 1 - 2 small setae and numerous small spines;

aedeagus with small sharp lateral spines near tip, tip rounded.

PUPA (Fig. 87). Integument tan to dark brown. Abdomen. Seta 0 usually

simple, infrequently bifid on segments II - VIII; 1-IV with 4 - 8 branches; 1-VI

with 3 - 8 branches; 1-VII with 2 - 5 branches; 5-I well developed, much longer

than 2 and 3-I; 5-IV with 3 - 7 branches; 5-VII with 4 - 11 branches; 6-II usu-

ally simple or bifid, as long as segment II; 6-III simple, slightly less than

length of segment III; 7-I with 2 - 3 branches, 0.50 - 0.60 as long as 6-I;

7, on VI - VII, 0.67 or more as long as following segment; seta 9 moderately

long, without long tapering tip, but sharp pointed, usually dark brown; 9-IV,

4 - 6 times as long as 9-III; 9-VII, 0.20 - 0.25 as long as segment VIII; 10,

on IV - V, 0.67 or more as long as following segments. Paddle. Refractile

margin 0.50 - 0.67 as long as paddle; fringe hairs approximately 0.33 length

of 1-P; outer margin evenly convex on distal half; 1-P long, simple, stout and

pigmented; 2-P with 2 - 4 branches.

LARVA (Fig. 88). Unicolorous gray or brown. Head. Seta 1-A with 3 - 7

branches; 4-A with 4 - 8 branches; setae 2-C long, stout and simple, with

bases very close together, much closer than distance between bases of 2-C and

3-C on one side; 3-C simple, rarely bifid at tip, approximately 0.33 length of

2-C; 4-C with 2 - 4 branches from base; 8-C with 2 - 6 branches; 9-C with 3 -

6 branches. Thorax. Seta 11-P short, simple (rarely bifid); 13-P with 3 - 6

branches; 14-P with 5 - 8 branches; all pleural setae (9, 10, 11, 12) on pro-

meso- and metathorax usually simple, 12-T infrequently bifid; small median

pigmented plate (frequently bilobed) often on dorsal midline between meso-

and metathorax; 3-T with flattened, lightly pigmented leaflets, without fila-

ments. Abdomen. All segments unicolorous; posterior tergal plates usually

present on segments III - VII, that on VII twice as large as others; seta 0 sim-

ple or 2 - 3 branched (rarely 3) on segments II - VI; 0-VIII simple or 2 - 4

branched (rarely 4); 1-I with well developed lightly pigmented leaflets, without

filaments; 1, on II - VII with well developed (palmate) flattened and pigmented

leaflets, with filaments; 5-II with 6 - 10 (usually 8 - 10) branches; 6-III with

5 - 9 branches; 11-II long and simple; pecten plate with 13 - 17 teeth; 1-X sim-

ple, approximately as long as saddle.

TYPE-DATA. The location of the type-specimen of insulaeflorum is un-

known. This species was described from larvae collected on Noesa Kembang-

an (now Nusa Kambangan), an island just south of Tjilatjap off the south coast

of Java. Although topotypic specimens were not available for examination, 2

whole larvae on slides are in the USNM from Pangandaran, only 19 km from

the type-locality. These larvae have seta 0 with 2 - 3 branches on segments

II - VI and 1-X simple. This combination of characters is like insulaeflorum

specimens from mainland Southeast Asia, rather than pilinotum.

DISTRIBUTION (Fig. 87). This is a widely distributed species that has

been collected in a variety of forest habitats in Thailand. The specific habitat

Harrison and Scanlon: Subgenus Anopheles in Thailand 169

requirements that determine the distribution of insulaeflorum are currently

unknown. Specimens (31%, 322, 63 larvae and 71 larval and pupal skins) in

the USNM were examined from the following provinces of THAILAND: Chantha-

buri, Chiang Rai, Kanchanaburi, Krabi, Lampang, Nakhon Ratchasima, Nan,

Phangnga, Phrae, Prachin Buri and Songkhla. Additional specimens (10°, 179

and 7 larvae) in the USNM were examined from INDIA (Assam), INDONESIA

(Java) and MALAYSIA (Peninsular). Specimens examined in the BMNH include

2 whole larvae from INDIA (West Bengal) and one larval skin from MALAYSIA

(Peninsular). Other distribution records from the literature include BURMA,

CHINA (Taiwan), INDIA (West), INDONESIA (Sumatra), SOUTH VIETNAM and

SRI LANKA.

Hsiao and Bohart (1946) listed insulaeflorum from Honshu Island, Japan,

based on an earlier record which they said ''appears to be somewhat doubtful."

This is almost certainly an erroneous record. Anopheles bengalensis is the

only member of the aitkenii group known to occur in the Japanese area, and

that is south of the main islands in the Ryukyus chain. The nearest records

of insulaeflorum to Japan are those from Taiwan, and although we have a male

and female from that island, the male genitalia has been lost and the identity of

the specimens cannot be confirmed.

Carter (1925) recorded insulaeflorum from Sri Lanka and illustrated the

inner clypeal differences between aitkenii and insulaeflorum. Based on these

illustrations we are convinced that his 'insulaeflorum" larva (p. 80, fig. 5d)

is not insulaeflorum because the bases of setae 2-C are nearly as wide apart

as 2-C and 3-C on one side. This illustration probably represents aitkenii

with 2-C simple. Since we have not seen specimens of insulaeflorum from Sri

Lanka and can find no substantial evidence it occurs there, we consider previous

records of this species from Sri Lanka as doubtful.

The distribution of insulaeflorum in Indonesia is not completely known, but

Java may be the easternmost extension. Past records of insulaeflorum from

Ambon, Ceram and the Philippines are now known to apply to pilinotum (Har-

rison and Scanlon 1974). The past records of insulaeflorum from Sulawesi,

Lesser Sunda Islands and the Molucca Islands now need confirmation to deter-

mine whether they apply to insulaeflorum or pilinotum. Neither species has

been collected from Borneo.

TAXONOMIC DISCUSSION. This is a most unusual member of the aitkenii

group because all the known stages can be identified on mainland Southeast

Asia. The adults can be recognized by the short fine setae on the prescutellar

space where other species have a bare space. The male is also distinguishable

by having spines on the aedeagus and should never be confused with the other

Thai species, except possibly tigeriti. The aedeagus of insulaeflorum has

small spines lateral only, while those on tigertti are lateral, dorsal and ven-

tral. The claspette of insulaeflorum is also distinct from that of tigertti. The

former has 3 - 4 outer dorsal setae and at least 3 large setae on the ventral

lobe, while the latter only has 2 outer dorsal setae and only 1 - 2 large setae

on the ventral lobe. The male genitalia of insulaeflorum are easily differenti-

ated from those of pilinotum by obvious (Fig. 79) differences in the setae on the

ventral lobe of the claspette and slight differences in the shape of the aedeagus

tip. The male genitalia illustrations (Fig. 79) are the first made of insulae-

florum, although Puri (1930), gave an excellent description of the claspette.

Most descriptions and apparently all illustrations previously made for the

male genitalia of insulaeflorum actually belong to pilinotum. Bonne-Wepster

and Swellengrebel (1953: 87, fig. 24f) were possibly illustrating insulaeflorum

with their 'aitkenii'"’ having minute hairs on each side at the tip of the aedeagus,

170 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

but we cannot be certain. The minute spines at the tip of the aedeagus of

insulaeflorum are less distinct than those on pilinotum, in fact, often small

enough to be overlooked unless they are viewed under high magnification.

All pupal descriptions and illustrations previously called insulaeflorum

actually refer to pilinotum. The pupae of these 2 species are similar, but

can be differentiated by insulaeflorum having seta 0, on IL - VIL simple or

bifid, the refractile margin on the paddle 0.5 or more as long as the paddle

and the outer margin of the paddle evenly convex on the distal half. The pupa

of pilinotum has seta 0, on II - VII with 2 - 6 (usually 3 - 5) branches, the

refractile margin on the paddle less than 0. 5 the paddle length and the outer

margin of the paddle straight or nearly straight on the distal half. The pupa

of insulaeflorum should not be confused with pupae of the other Thailand mem-

bers of this group because of the combination of short setae 9, a long refractile

margin on the paddle and being moderately to darkly pigmented. The pupa of

stricklandi is the most likely member to be confused with insulaeflorum; how-

ever, stricklandi has a much shorter seta 9, shorter and more widely spaced

fringe hairs on the paddle and is very pale.

The larva of znsulaeflorum is easily separated from all the other Thailand

members of this group except stricklandi, by having the following combination

of characters: seta 2-C simple and close together, small anterior tergal

plates on the abdomen, 11-II simple, 6-III with less than 15 branches and 1-X

simple. The larva of stricklandi is very similar to that of insulaeflorum and

workers in Thailand should be aware of the small differences (see key) that

differentiate these 2 species.

The larvae of tnsulaeflorum and pilinotum are also very similar. The

former has seta 11-P single or rarely bifid, 0, on II - VI simple or with 2 -

3 (rarely 3) branches and 1-X simple, while the latter has 11-P with 2 - 6

branches, 0, on II - VI with 3 - 6 branches and 1-X with 2 - 4 branches.

BIOLOGY. This species does not appear to be abundant anywhere in

Thailand. Usually larval collections of insulaeflorum involve 5 - 10 speci-

mens, even when the habitat appears to be ideal. The immatures are found

only in or near forested areas. In Thailand they have been found in a number

of habitats in primary and secondary wet forests, a bamboo grove and ina

rural village adjacent to a forested area. These collections were usually in

mountainous regions, but several were made in forested plains areas. Col-

lections in Thailand record insulaeflorum between 46 - 700 m elevation, and

one immature collection in Perlis, Malaysia was made at sea level. Imma-

tures have been collected from the following sites in Thailand: isolated rock

and ground pools near streams, large and small stream pools, stream mar-

gin, small seepage pools and a water pool on a fallen tree. These sites were

nearly always under either moderate or heavy shade, and there was usually

floating vegetation, dead leaves or brown algae in the water. The water was

either clear or colored and was either stagnant or flowing very slowly.

There are no adult females of znsulaeflorum in the USNM or BMNH that

we could determine were taken in biting collections. Further, there are no

confirmed published records of this species biting man. Stujanovich and

Scott (1966) said this species feeds in jungle shade on cattle and man, but

this needs confirmation.

Harrison and Scanlon: Subgenus Anopheles in Thailand 171

ANOPHELES (ANOPHELES) STRICKLANDI REID

(Figures 79, 89, 90)

Anopheles (Anopheles) stricklandi Reid 1965: 121 (o*, 2*, P*, L*); Reid

1968: 250 (**, 2*, P*, L*); Rattanarithikul and Harrison 1973: 5 (L*).

This species is small and pale as an adult, and only the adult males can

be identified. The pupa is easily differentiated by the very short seta 9, the

angular projections on tergum IX and paddle characters. The larva, although

identifiable, is much more apt to be confused with znsulaeflorum, a more

commonly encountered species in Thailand. Like bengalensis except:

FEMALE. Cannot be separated from bengalensis at present. Ratio fore-

femur/proboscis (sample 4 females) range 1.00 - 1.02, mean 1.01; pleural

setae number 0 - 1 propleural, 2 - 4 spiracular, 1 prealar, 1 upper and 1 -

2 lower sternopleural, 1 - 3 upper and 0 lower mesepimeral; upper midcoxa

with 2 setae; wing cell Mo usually less than 1/2 as long as stem (from cross-

vein).

MALE (Fig. 79). Habitus currently indistinguishable from bengalensis.

Genitalia. Claspette with 3 short thick outer dorsal setae, inner dorsal club

formed from 2 very broad setae; ventral lobe of claspette with 3 setae the same

length as the club or shorter; aedeagus without spines or leaflets (see taxono-

mic discussion); basal half of aedeagus broad, then narrowing on distal half

only to widen slightly for broadly rounded apex, shape similar to a bowling pin.

PUPA (Fig. 89). Integument pale to light tan. Cephalothorax. Seta 7-C

with 3 branches; 10-C stout, usually simple or bifid. Abdomen. Seta 0, on

II - VUI short and simple; seta 1, on III - IV well developed with few branches;

1-III with 3 - 6 branches, stem as stout as that of 3-III; 1-IV with 3 - 6 bran-

ches; 1-VI with 2 - 4 branches; 1-VII with 2 - 5 long, simple or with 2 - 3

distal branches, much longer than 2 and 3-I; seta 5, on V - VII with few bran-

ches and strongly developed, approximately 0.67 as long as following segment;

o-IV with 3 - 5 branches; 5-VII with 3 - 6 branches; 6-I simple, bifid or trifid,

much longer than segment I; 6-II simple or bifid, slightly more than 0. 67

length of segment II; 6-III simple or bifid, about 0.67 length of segment III;

7-I simple, bifid or trifid, about 0.75 as long as 6-I; seta 7, on VI - VII,

0.50 - 0.67 as long as following segment; setae 9 very short; 9-IV, 2 - 3 times

as long as 9-III; 9-VII, 0.13 - 0.17 as long as segment VIII; 10, on IV - V,

0.50 - 0.67 as long as following segment; tergum IX with prominent latero-

caudal angle over each paddle base. Paddle, Refractile margin 0.67 or more

as long as paddle; fringe hairs short and widely spaced, 0.25 - 0.33 as long as

1-P; outer margin evenly convex on distal half; 1-P long and simple; 2-P

simple or bifid.

LARVA (Fig. 90). Unicolorous yellow or tan. Head, Seta 1-A with 5 - 9

branches; 4-A with 5 - 8 branches; 2-C long, stout and simple, with bases very

close together, much closer than distance between bases of 2-C and 3-C on one

side; 3-C split distally into 2 - 4 stout, equal branches, approximately 0.33

length of 2-C; 4-C with 3 - 5 branches from near base; 8-C with 2 - 4 branches;

9-C with 3 - 5 branches. Thorax. Seta 1-P with 8 - 12 branches; 2-P with

12 - 15 branches; 11-P short, simple and flattened on distal half (fusiform);

13-P with 3 - 6 branches; 14-P with 5 - 6 branches; all pleural setae (9, 10,

11, 12) on pro- meso- and metathorax usually simple, 12-T infrequently bifid;

without small median pigmented plate on dorsum of meso- or metathorax; 3-T

with flattened unpigmented leaflets, without filaments. Abdomen. All seg-

riz Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

ments unicolorous; posterior tergal plates usually present on segments III - VII,

that on VII largest; seta 0, on II - VII with 2 - 3 branches; 0-VIII with 3 bran-

ches; 1-I branched, without flattened leaflets; 1-II with slightly pigmented flat-

tened leaflets, without filaments; 1, on III - VII with flattened, pigmented leaf-

lets, with filaments; 5-Il with 7 - 11 branches; 6-III with 4 - 10 branches; 11-II

long and simple; pecten plate with 9 - 14 teeth; 1-X simple, approximately as

long as saddle. |

TYPE-DATA. The @ holotype, ¢ allotype, and one “ and one ? paratype

are in the BMNH, each with associated immature skins mounted on a single

slide. The “ holotype is in excellent condition and has the following label

data: (1st label) - "Malay Peninsula, Johore, 28.5.1935. E. P. Hodgkin";

(2nd label) - '9667/70'"'; (3rd label) - "Anopheles stricklandi “ holotype, genit.

on slide''; and (4th label) - is a round BMNH "Type" label. The slide bearing

the holotype genitalia and its larval and pupal skins, bears the same number

and small round "holotype" label. The ¢ allotype is in excellent condition and

bears the same data as the holotype except for the small round "'allotype" label

and the rearing number ''9667/10.'' The larval and pupal skins of the allotype

are mounted on a slide bearing that number. The “ and 2 paratypes are both

in excellent condition, and bear the following label data: (1st label) - ''Perlis,

19.8.1934, R. E. Anderson;" (2nd label) - 8355/2" for the male, and ''8355/

4"' for the female; (3rd label) - "A. aitkeni v.B"''; and identification and ''para-

type" labels. Both paratypes have a slide bearing their larval and pupal skins;

however, that of the “ paratype also bears a genitalia mount.

DISTRIBUTION (Fig. 89). This is a very infrequently encountered species

that is known only fromPeninsular Malaysia and southern Thailand. Reid

(1965, 1968) suggests stricklandi is a lowland species, and this is supported by

Thailand collections of 100 - 180 m elevation. Specimens (3, 1 larva and 3

pupal skins) in the USNM were examined from the following provinces of THAI-

LAND: Krabi and Ranong. Additional specimens (1%, 19 and 4 larval and pu-

pal skins) in the USNM were examined from MALAYSIA. Specimens (2, 22

and 8 larval and pupal skins = type-series) in the BMNH were examined from

MALAYSIA. Since Thailand collections were made in Ranong adjacent to Burma,

this species probably occurs in Burma.

TAXONOMIC DISCUSSION. Although the adults of stricklandi cannot be

separated (except male genitalia) from most of the other Thailand species, the

immature stages are identifiable. The pupa of stricklandi is the easiest stage

to identify, and should not be confused with that of any other species in the azt-

kenit group because it has the shortest seta 9, sparse setal branching, a very

long paddle refractile margin and short sparse paddle fringe hairs. The larva

is easily differentiated from all the Thai species except insulaeflorum by hav-

ing: setae 2-C simple and very close together; small anterior tergal plates on

the abdominal segments; and seta 6-III with less than 15 branches. Only a

closer examination using the key characters can differentiate stvicklandi lar-

vae from those of insulaeflorum. Larvae of stricklandi can be separated from

pilinotum larvae by the same characters that are used to separate insulaeflor-

um from pilinotum.

This species appears to be closely related to insulaeflorum and pilinotum,

but the adults do not possess the short fine prescutellar setae that occur on

insulaeflorum and pilinotum, and therefore, stricklandi females cannot be

identified without associated immature skins. Male stricklandi have very dis-

tinct genitalia characters that look similar to those of pilinotum, except the

aedeagus of stricklandi does not have small spines. Reid (1965, 1968) sug-

gested that one specimen of stricklandi he examined may have had small

Harrison and Scanlon: Subgenus Anopheles in Thailand 173

serrations on the aedeagus, but we have not seen this. The 2 genitalia prepara-

tions (holotype and paratype) in the BMNH do not exhibit serrations on the ae-

deagus. The setae on the dorsal lobe of the claspette are even broader than

those encountered on pilinotum or acaci and should not be confused with any

species found in Thailand. The illustration (Fig. 79) used for the claspette

of stricklandi was taken fromReid (1965).

Reid (1965, 1968) noted that wing cell My on stricklandi is usually half or

less as long as the cell stem vein (measured from the crossvein). On the

specimens of stricklandi we examined, this is usually true; however, one

specimen had My more than half as long as the stem vein, and specimens of

3 other species in the group were noted to occasionally exhibit comparable

Mog lengths. Consequently, this wing measurement may be of value in helping

to define some specimens of stricklandi, but should be used with caution be-

cause it is not completely reliable.

BIOLOGY. Very little is known about the habitat requirements of this

species. Reid (1968) lists larvae found in seepages, drains and hoof marks

under shade in Malaysia. Immature collections in Thailand have been made

from large and small stream pools and a large seepage pool, all under partial

shade in secondary wet forest between 100 - 180 m elevation. The water from

these sites was clear, stagnant or slowly running and usually contained dead

leaves or green algae. All confirmed specimens of this species have come

from immature collections, thus, nothing is known about the adult behavior.

ANOPHELES (ANOPHELES) PALMATUS (RODENWALDT)

(Figures 78, 79, 91, 92)

Stetomyia aitkenii var. palmata Rodenwaldt 1926: 794 (L*)(Stethomyia LAPSUS).

Anopheles (Anopheles) aitkenii var. palmata Rodenwaldt, Covell 1927: 7;

Edwards 1932: 37.

Anopheles (Anopheles) aitkenit palmatus Rodenwaldt, Swellengrebel and

Rodenwaldt 1932: 112 (L*); Gater 1934: 72 (L*); Gater 1935: 149 ();

Russell, Rozeboom and Stone 1943: 109 (distribution).

Anopheles (Anopheles) palmatus Rodenwaldt, Lee and Woodhill 1944: 98

(, L); Bonne-Wepster and Swellengrebel 1953: 98 (o, L*); Reid 1963:

113.

Anopheles (Anopheles) palmatus (Rodenwaldt), Reid 1965: 117 (“, L*); Reid

1968; 246 (co, L*); Rattanarithikul and Harrison 1973: 5 (L*).

This is one of the easiest species in the aitkenii group to identify. The

larva is unique in the group because of the very large anterior tergal plates.

The pupa, not previously described or illustrated, has very distinct seta 9 and

paddle characters. Even some of the adults can be identified by a color pat-

tern on the abdomen, and the male genitalia, illustrated here for the 1st time,

also have distinctive characters. Like bengalensis except:

FEMALE (Fig. 78). Unicolorous gray-brown except pale abdominal seg-

ment IV on majority of specimens (see taxonomic discussion). Pleural setae:

0-1 propleural, 0 - 2 spiracular, 1 prealar, 1 upper and 2 - 4 lower sterno-

eel and 2 - 4 upper and 0 lower mesepimeral; upper mid coxa with 3 - 4

setae.

MALE (Figs. 78, 79). Habitus as for female. Genitalia. Dorsal lobe

of claspette with 2 outer dorsal setae approximately as long as inner dorsal

club which has 2 basal stems; ventral lobe of claspette with 4 setae approxi-

mately as long as club on dorsal lobe, also with small mesal spicules; aedeagus

174 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

very long and slender with tip slightly swollen, without leaflets or spines.

PUPA (Fig. 91). Integument tan to light brown. Cephalothorax. Seta 7-C

with 3 - 6 branches; 10-C stout and simple, or with 2 - 3 distal branches.

Trumpet, Simple, with deep meatal cleft, meatus less than 0.33 as long as

trumpet. Abdomen. Seta 0, on II - III simple and short; seta 1, on II - VII

well developed, longer than seta 5, on II - Il] and V - VII, nearly as long as

following segment on segments V - VI; 1-III with 6 - 12 basic branches,

usually dendritic; 1-IV with 6 - 12 basic branches, usually dendritic; 1-VI

with 4 - 10 branches, not dendritic; 1-VII with 5 - 8 branches, not dendritic;

5-I with 3 branches, equal or slightly longer than 2-I; 5, on IV - VII strongly

developed, length equal to seta 1 only on segment IV; 5-IV with 7 - 9 branches;

5- VII with 10 - 12 branches; 6-I with 2 - 4 branches from near base, equal

to or slightly longer than segment I; 6, on II - V and VII short, simple or with

2 - 4 branches; 6-VI long, simple or bifid, at least twice as long as 9-VI; 7-I

with 2 - 5 branches, about half as long as 6-I; 7-VII long, simple or bifid, ap-

proximately 0.66 as long as segment VIII; seta 9 long, brown, with round

blunt tip; 9-VII approximately 0.25 as long as segment VIII; 10, on IV - V ap-

proximately half as long as following segment. Paddle. Twice as long as

wide; refractile margin 0.33 or less as long as paddle, but bases of fringe

hairs refractive around entire margin of paddle; fringe hairs long, dense,

0.50 - 0.67 length of 1-P; outer margin straight or nearly so on distal half;

1-P long, stout, simple and curved near tip; 2-P with 3 - 5 branches from

near base. |

LARVA (Fig. 92). Whole larva dark brown with abdominal color pattern.

Head. Seta 1-A with 7 - 10 branches; 4-A with 5 - 7 branches; 2-C long, stout

and simple, with bases very close together, much closer than distance between

bases of 2-C and 3-C; 3-C split distally into 2 - 5 stout equal branches, short

approximately 0.2 as long as 2-C; 4-C with 2 - 6 branches; 5, 6, 7-C short,

well developed and plumose; 8-C with 2 - 4 branches; 9-C with 2 - 5 branches.

Thorax. Seta 1-P fan-like with 15 - 18 rounded branches; 2-P with 12 - 17

branches, about twice as long as 1-P; 3-P small and simple; 11-P short, with

4 - 7 branches from near base; 13-P with 5 - 8 branches (dendritic); 14-P with

5 - 6 branches; long pleural setae on pro- meso- and metathorax simple except

9-P with 2 - 3 branches; small median plate often on dorsal midline between

meso- and metathorax; 3-T with flattened lightly pigmented leaflets, with fila-

ments; 12-T simple or bifid. Abdomen, All segments brown except pale yel-

low segment IV; anterior tergal plates very large, enclosing posterior tergal

plates, 0.67 - 0.75 as wide as segment; seta 0, on II - VII simple or bifid,

inserted on anterior tergal plates approximately 0.33 of way between lateral

margin and center of plate; 0-VIII with 5 - 6 very long slender thread-like

branches, inserted just inside or slightly posterior to caudal margin of anteri-

or tergal plate, approximately 0.33 of way between lateral margin and center

of plate; 1, on I - VII (palmate) with flattened, pigmented leaflets, with fila-

ments; 5, on II - IV small, weakly developed; 5-II with 3 - 5 branches; 5, on

I and V - VII large, well developed; 6-III with 18 - 27 branches; 11-II long with

2 - 3 distal branches; pecten plate with 13 - 18 teeth; 1-X with 2 - 3 distal

branches, approximately as long as saddle.

TYPE-DATA. This species was described from specimens collected in

Sidoeagoeng (Preanger Regencies), West Java, but the type-specimens are

apparently non-extant.

DISTRIBUTION (Fig. 91). This species has been collected only a few times

in Thailand from widely separated sites. Specimens (1%, 4%, 5 larvae and 9

larval and pupal skins) in the USNM were examined from the following provinces

Harrison and Scanlon: Subgenus Anopheles in Thailand 175

of THAILAND: Chanthaburi, Chiang Rai, Ranong and Songkhla. There is also

one larval skin in the USNM from MALAYSIA (Peninsular Malaysia). An ad-

ditional 2 larval skins in the BMNH were examined from MALAYSIA (Peninsular

Malaysia). Reid (1968) gave the known distribution of this species as Thailand,

Malaysia (including Borneo) and Indonesia from Sumatra to the Moluccas and

the western tip of New Guinea. This species has not been reported from Sula-

wesi or the Philippines. Scanlon et al. (1968) considered the specimens from

Chiang Rai Province, Thailand, as suspect because the circumstances under

which the specimens were obtained make the locality data doubtful. Sandhinand

(1951) reported palmatus from Chiang Mai Province, but these specimens have

not been seen. Scanlon et al. (1968) suggested that the records from these 2

northern provinces need further study because the known distribution of palma-

tus suggests it is a Malaysian-Indonesian species that may have its northern-

most limit in southern Thailand. Since Scanlon et al. (1968), specimens of

palmatus have been found that were collected on a mountain adjacent to Cam-

bodia in southeastern Thailand. These specimens definitely show that: palma-

tus is not confined strictly to the Malaysian-Indonesian region; however, they

do not lend support to the northern Thailand records. As already shown in

the barbirostris, hyrcanus and umbrosus groups, several Malaysian-Indonesian

species previously presumed to have their northernmost distribution in southern

peninsular Thailand are now known to also occur in southeastern Thailand.

This disjunct distribution is apparently due in part to current precipitation pat-

terns and the distribution of tropical wet forests. Since palmatus was collected

in Ranong Province adjacent to Burma, it also probably occurs in tropical wet

forests habitats in the southern part of that country. Reid (1965, 1968) indi-

cated that the distribution of this species was probably associated with foothill

and mountainous regions. This suggestion is supported by the Thailand collec-

tions which were made between 140 - 1,300 m elevation, usually in mountain-

ous areas.

TAXONOMIC DISCUSSION. The larva of palmatus is unique in the subgenus

Anopheles of Southeast Asia in having the abdominal anterior tergal plates very

large. It is also one of 2 species in the aztkenii group that exhibits a color pat-

tern on the abdomen of the larva and adult. Larvae of palmatus are dark brown

except for a pale yellow abdominal segment IV, and should not be confused with

aberrans larvae which usually have abdominal segments IV and V pale yellow.

The adults of palmatus have previously been considered indistinct, however,

of the one male and 4 females in the USNM only one female has the abdominal

segments unicolorous while the remainder have abdominal segment IV distinct-

ly pale yellow in comparison with the remaining brown segments. This color

pattern is very similar to that found on most aberrans adults, which have ab-

dominal segments IV and V pale yellow.

The male genitalia of palmatus were 1st described (not illustrated) by

Gater (1935) from 2 males sent to him from Thailand (locality unknown). The

association of this description with palmatus has previously been presumed

correct, but could not be confirmed for lack of males. We now have a male

genitalia preparation of palmatus (USNM) from a specimen with an associated

pupal skin. This preparation confirms that Gater was describing palmatus, in

fact, his description is very accurate and agrees with our specimen in nearly

all respects. The claspette of palmatus looks very similar to those of bengal-

ensis, tigertti and some of aberrans, except that the ventral lobe of palmatus

has 4 large setae, instead of the 3 large and one small setae usually found on

the other species. The very long aedeagus on palmatus males is distinct and

unusual for the group, and is somewhat suggestive of that described for

176 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

_pinjaurensis from northwestern India (Christophers 1933: 110).

The pupa of palmatus has several very distinct characters which make it

easy to identify. Setae 9 are unlike those of any other member of the aitkenii

group in having the tips bluntly rounded. Most members of the group have

long extremely attenuate tips on pupal seta 9. Three species, insulaeflorum,

pilinotum and stricklandi, have pupal seta 9 short, or moderately long as on

balmatus, but these 3 species have the tips of seta 9 sharp pointed. The

shape of the paddle on the pupa of palmatus is also distinct. This structure

is twice as long as wide (not including fringe hairs) on palmatus, while on the

other members of the group the width is nearly 0.67 the length. The length

and refractiveness of the fringe hairs on the pupal paddle of palmatus are also

unusual and may be of assistance in identifying this species.

The larval setal arrangement and development of 2-C and 3-C on palmatus

suggests a relationship with insulaeflorum, pilinotum and stricklandi. It also

implies that 6-III should have few branches, but this is not the case. Reid

(1965) pointed out that Rodenwaldt's original illustration of palmatus showed

6-II with only 3 branches. However, this is almost certainly an error on the

part of Rodenwaldt, for Reid (1965) noted 21 - 26 branches on 6-III, and we

found 18 - 27 branches on 6-III. The development of seta 11-II also separates

palmatus from the above 3 species. Seta 11-II on palmatus is either bifid or

trifid, while those on insulaeflorum, pilinotum and stricklandi are simple. The

affinities of palmatus to the other members of the aztkenii group is currently

obscure and will probably remain thus until more extensive collections of this

eroup have been made.

BIOLOGY. All collections of this species have been immature collections,

thus, nothing is known of adult behavior. The immature stages have been col-

lected in Thailand in moderate to heavily shaded stream pools and stream mar-

gins in primary and secondary wet forests between 140 - 1,300 m elevation. The

water in these sites was clear, moving slowly and usually contained dead leaves,

sticks or roots. Larvae of palmatus have been collected in Thailand in associa-

tion with larvae of aberrans and bengalensis.

ANOPHELES CULICIFORMIS SPECIES GROUP

(A, culiciformis species group, Reid and Knight 1961)

Anopheles culiciformis Cogill 1903.

The adults of this group are unicolorous and similar to those of the aztkeniz

group. At least one species is known to have a culicine resting posture. Gen-

erally the adults are slightly larger than those of the aitkenii group. The fe-

males are indistinguishable, (except sintoni) and the taxonomy of the group has

been almost entirely dependent upon male genitalia and larval characters. This

species group was formed by Reid and Knight (1961) and the most recent review

with a key to the larvae of all species is found in Reid (1968).

ADULT. Small, unicolorous brown; interocular space narrow; flagello-

mere 1 with dark scales; erect vertex scales broad and fan-like at apex; fe-

male palpus slightly shorter than proboscis, most distal segments slightly

bushy; anterior pronotal lobes without scales; aedeagus with hair-like spicules

or leaflets on distal 0.33; claspette with 3 lobes; all setae on claspette separate;

2 parabasal spines on basimere arising on very prominent tubercle; ventro-

mesal spine inserted approximately 0.67 of distance from base to apex of basi-

mere.

Harrison and Scanlon: Subgenus Anopheles in Thailand 177

PUPA. Trumpet with shallow rounded meatal cleft, meatus 0.50 - 0. 67

as long as trumpet; seta 5, on IV - VII very long, at least as long as segment;

1-P straight or filiform with a curved hooked tip.

LARVA. Setae 2-C simple, with bases very close together; 4-C simple

or branched distally; 5, 6, 7-C short and reduced with few branches, not _

plumose; 1-A short and simple or with 2 - 5 distal branches; 11-C shorter

than antenna with distal branches, or if as long as antenna then plumose from

base; 6-VI usually approximately equal length of 6-III; stigmal knob on the

median dorsal valve of spiracular apparatus as darkly pigmented as remainder

of valve.

DISTRIBUTION. This group is confined to the Oriental faunal region, with

species recorded from. the west coast of India, east to Hainan Island in southern

China and south into Peninsular Malaysia. The distributions of some of the

species in this group approach the ranges of bavianensis James in northwestern

India, and omorvii in Japan. The 2 latter named species are members of the

Nearctic-Palearctic plumbeus group which resembles the culiciformis group

in several ways. Of the 5 species known in the culiciformis group only 2, i.e.,

kyondawensis and sintonoides are known from Thailand, and they are rarely

encountered.

TAXONOMIC DISCUSSION. Reid and Knight (1961) recognized 5 species. in

this group, they are: alongensis Venhuis, culiciformis, kyondawensis, sin-

toni Puri and sintonoides. Taxonomically, little is known about these 5 spe-

cies. The females are usually not distinguishable and pupae have been des-

cribed for only 3 of the species. Thus, male genitalia and larval characters

are the basis for the present group arrangement of these species. Our lack

of knowledge of these species is due to 3 basic factors: (1) these are all in-

frequently encountered forest species; (2) the immatures are found in cryptic

habitats which are not normally collected during routine anopheline surveys;

and (3) the included species are not known or even suspected of being involved

in the transmission of human pathogens.

Reid (1968) suggested this group has affinities somewhat intermediate

between the Nearctic- Palearctic plumbeus group and the Southeast Asian

aitkenit group. Adults of the culiciformis group are: unicolorous as are most

of the species in the other 2 groups; have wider vertex scales than the other

2 groups; have dark scales on the base of the antennal flagellomere 1 like

members of the pPlumbeus group; and have few propleural, sternopleural and

upper mesepimeral setae like members of the aitkenii group. The male geni-

talia of the culiciformis group have separate claspette setae and the ventro-

mesal spine on the basimere more like species in the plumbeus group, yet,

the aedeagus on species of the culiciformis group is distinct from the other 2

groups because it possesses hair-like spicules or leaflets. The pupae of the

culiciformis group have the trumpet with a long meatus like members of the

plumbeus group, but the development and length of seta 5 on the abdominal

segments and the paddle fringe is more like species in the aitkenii group.

The larvae of the culiciformis group have reduced setae 5, 6, 7-C, a reduced

seta 11-C (except alongensis and kyondawensis) and a long seta 6 on IV - VI,

like the species in the plumbeus group. The larval habitats utilized by 3 spe-

cies in the culiciformis group are primarily tree hole-plant container habitats

like those used by the plumbeus group. However, 2 species in the culiciformis

group have cryptic larval habitats such as rock holes and crab holes, and a 3rd

member has been collected in ground pools. These latter habitats are more

like those utilized by members of the aitkenii group.

Since 3 species in this group are not known from Thailand, they will be

178 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

briefly discussed.

Anopheles alongensis is known only from the type-locality, Along Bay ad-

jacent to Haiphong, North Vietnam. Specimens were reared and described in

detail (as tonkinensis) by Galliard and Neu (1946). Collections came from

freshwater rock holes in a limestone cave. The male genitalia are distinct

due to the presence of leaflets on the aedeagus and the larvae have: setae 2-C

widely separated; 11-C plumose and as long as the antenna; and 6-VI very

short. The pupa (Reid 1968) is similar to that of sintonoides, but seta 5 on

V - VII is simple or branched distally. Based strictly on larval characters

this species would appear to be more closely related to kyondawensis, but

can be separated from that species by the widely separated bases of setae 2-C

and the short seta 6-VI. The larval habitat of alongensis, i.e., rock holes in

a dark cave, has a basic similarity to the shaded crab hole habitat utilized by

kyondawensis (see kyondawensis for further discussion). The type-larva for

alongensis was last reported in the Geneeskundig Laboratorium, Djakarta,

Indonesia, and the types of the synonym, tonkinensis, were reported as in the

Ecole Medicale, Hanoi, North Vietnam (Stone et al. 1959). Grothaus et al.

(1971) reported larvae of this species collected in the Cam Lo area of northern

South Vietnam, but this report needs confirmation.

Anopheles culiciformis was described by Cogill from adults reared from

larvae collected in ground pools and one large collection from a tree hole. Ap-

parently, all subsequent collections have been from tree holes. This species

is known only from the west coast of India. Christophers (1933) described

and illustrated the most pertinent characters. The male is distinct on the

basis of long curved leaflets on the aedeagus. The pupa is distinct by having

seta 5 on V - VII nearly twice the length of the segments. The larva has: very

reduced-few branched setae 5, 6, 7-C, seta 11-C shorter than the antenna and

expanded distally with numerous distal branches, no spiny-stellate setae and

seta 6-VI with short lateral branches (Puri 1931). This is a forest species

with the immature stages normally found in tree holes. According to Christo-

phers (1933) there are no records of this species collected in houses or biting

man. The type-specimens of this species are deposited in the BMNH.

Anopheles sintoni is known only from the west coast of India, where im-

matures are found in tree holes also containing culiciformis. This species is

distinct on the basis of male genitalia and larval characters (Christophers

1933). The pupa has not been described. The male genitalia of sintoni have

a claspette very similar to that of culiciformis, but lack the long leaflets on the

aedeagus, replacing them with dense brush like hairs. The larva of sintoni

has: short setae 5, 6, 7-C with more numerous branches; seta 11-C without

the distal portion being expanded and with fewer distal branches than found on

culiciformis; seta 14-P with spiny-stellate branches; and seta 6-VI with long

lateral branches (Puri 1931). Christophers (1933) describes the adult of

sintoni as having a few small scales on the median portion of the scutum, which

can be used to differentiate the females of this species from culiciformis.

Nothing is known about the adult behavior and biology. The type-specimens of

this species are deposited in the BMNH.

BIOLOGY. The species in this group oviposit in 2 basic type habitats.

- Three species,. culiciformis, sintoni and sintonoides primarily utilize tree-

nole and other plant containers for oviposition. However, at least culiciformis

can also survive in ground pools. The other 2 species, alongensis and kyonda-

wensis utilize cryptic ground water habitats. Larvae of alongensis are known

only from rock holes in a dark cave, while a single larva of kyondawensis has

been found in a fresh water crab hole in Thailand. The original collections of

Harrison and Scanlon: Subgenus Anopheles in Thailand 179

kyondawensis, much like those of culiciformis were taken from pools of clear

water beside streams. Since kyondawensis larvae have been collected only

twice, and in 2 different habitats, the basic larval habitat of kyondawensis is

still uncertain.

There are no published records of females of this group biting man and

only a single female of sintonoides is noted here as collected biting man.

Accordingly, the species in this complex are presumed to be of no significance

in the transmission of human pathogens.

KEYS. The keys to the 2 species of this complex found in Thailand have

been incorporated into the keys to the species in the Anopheles series on

pages 145-7.

ANOPHELES (ANOPHELES) KYONDAWENSIS ABRAHAM

(Figures 93, 95)

Anopheles (Anopheles) kyondawensis Abraham 1947: 173 (L*); Reid and Knight

1961: 482; Reid 1968: 221; Khin-Maung-Kyi 1971a: 288 (distribution);

Rattanarithikul and Harrison 1973: 4 (L*).

This is a very poorly known species, described only from the larval stage.

It can be differentiated from all other Southeast Asian anopheline larvae by the

strongly curved antennae and reduced setae 5, 6, 7-C.

FEMALE. Unknown.

MALE. Unknown.

PUPA. Unknown.

LARVA (Fig. 93). Unicolorous pale yellow. Head. Antenna cylindrical

and strongly curved inward, diameter approximately equal from base to apex,

with spicules, particularly on dorsomesal aspect; seta 1-A very short, inserted

on basal 0.17 of antenna, with 3 - 5 branches from midway along stem; 4-A

with 5 - 10 branches, slightly longer than 2 and 3-A; maxillary palpus seta 8-

MP well developed, with large central stem and many lateral and distal bran-

ches; 1-C long with fine attenuate tip; 2-C very long and simple, with attenu-

ated tip; bases of setae 2-C adjacent, much closer together than setae 2 and

3-C on one side; 3-C short and simple, approximately 0.2 as long as 2-C;

setae 4-C simple, fine and approximately as long as 3-C, with alveoli further

apart than alveoli of 3-C; frontal setae very short, 5-C with 2 - 6 branches,

6-C with 3 - 6 branches and 7-C with 4 - 6 branches; 8-C short and simple;

9-C short and simple; 11-C nearly as long as antenna and plumose, with numer-

ous lateral and distal branches. Thoray. Integument glabrous; seta 1-P with

0 - 8 branches from basal half of stem; 2-P arising from small sclerotized

tubercle, with 8 - 13 branches and approximately 5 times as long as 1-P; 3-P

simple, about as long as 1-P, closer to 2-P than distance between 1-P and

2-P; 0-P slender and bifid, posterior to 4-P; 4-P with stout central stem,

numerous lateral and distal branches and shorter than 2-P; prothoracic pleural

tubercle with short spine; 11-P short, fine and simple; long pleural setae 9,

10, 12 simple on all 3 thoracic segments (except 12-T); 13-P with 5 - 6 branches;

14-P with 4 - 6 branches; 1-M with 42 - 44 branches; 4-M caudal of long simple

3 and 5-M, short and with 3 - 4 branches; 3-T with slender, unpigmented flat-

tened leaflets, without filaments; 12-T short and bifid distally; 13-T very long,

simple and slightly flattened. Abdomen. Integument glabrous, anterior tergal

plates small, less than 0.25 as wide as segment; small posterior tergal plates

present on segments IV - VII; seta 0, on II - VID short and simple; 1-I minute

180 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

and simple; 1-II with slender, unpigmented flattened leaflets, without filaments;

seta 1, on ID - VII (palmate) with pigmented flattened leaflets, with filaments;

seta 2, on IV - VI long, simple and flattened; 3, on I - VI long and flattened,

usually simple, bifid or trifid distally; 4, on VI - VII long, simple and flattened;

5-II small with 3 - 4 branches; 5, on VI - VII larger with 8 - 9 branches; 6-III

as long as plumose 6, onI - II, flattened with 8 - 9 very short branches mostly

on basal half; 6, on IV - VI simple, round not flattened, as long as 6-III; 6-VII

very small with 2 - 3 branches; 10, on I and V - VI long simple and flattened;

11-II long simple and flattened; 12, on II and VI - VII long simple and flattened;

13, on I - IV and VI small with 4 or more branches on I - IV and VI, long sim-

ple and flattened on V and VII; stigmal knob on median dorsal valve of spiracu-

lar apparatus pigmented, as dark as remainder of valve; pecten plate with 16 -

17 subequal teeth, each with strong basal and lateral denticles; 1-X simple,

slightly flattened, inserted at edge of saddle and as long as saddle.

TYPE-DATA. Abraham based his description on the examination of 11

larvae, of which 7 were in the 4th stage. These specimens were collected

over a 10 day period (5 collections) in 1943, from a small village named Kyon-

daw (now Kyondo), near Moulmein in southern Burma. The village of Kyondaw

is situated on the bank of a river of the same name, apparently a smaller tribu-

tary of the Gyaing River. The holotype larva is in the BMNH, and the slide

bears the following data: 'Department of Biology, King Edw. VII College of

Medicine, Singapore, Type Anopheles kyondawensis Abraham, Lower Burma,

2-12-1943, Det. O. K. Abraham.'' Paratypes are reported to be deposited in

the mosquito collections of the College of Medicine, Singapore.

DISTRIBUTION (Fig. 95). This species is known only from the type-

locality in Burma and a single larva collected in Nan Province, Thailand. The

larva from Thailand is deposited in the USNM. The latter locality, i.e., Ban

Pha Man, is situated at 425 m elevation near the Laos border and nearly 400 km

from the type-locality. This suggests that kyondawensis is widely distributed

on mainland Southeast Asia, but rarely encountered, probably due to low popu-

lation levels and/or a cryptic immature habitat (also see Biology section).

TAXONOMIC DISCUSSION. Reid and Knight (1961) included this species

in the culiciformis species group on the basis of the reduced setae 5, 6, 7-C

and other setae on the larval head. Since kyondawensis is known only from

the larval stage this status must be considered tentative; however, the larva

of this species does bear a number of similarities with the larva of alongensis,

another ground water species in this group. Larvae of both these species

have seta 11-C approximately as long as the antenna, with many branches from

near the base, while the other members of the group have this seta short and

mainly with distal branches. Differences between the larvae of alongensis

and kyondawensis were noted in the species group discussion.

The only other species of this group that occurs in Thailand, i.e., sinton-

oides, has larvae that are partially covered with spicules, have large stellate

spiny setae on some segments and are found only in plant container and axil

habitats. The curved antenna of kyondawensis larvae is unique among South-

east Asian Anopheles, and coupled with the short setae 5, 6, 7-C on the head

and long setae 6 on IV - VI, this species should not be confused with any other

species in Thailand.

The single larva from Thailand exhibits slightly fewer branches on 5, 6,

7-C than the holotype, otherwise, these 2 specimens agree very well.

BIOLOGY. Reid and Knight (1961), using culiciformis records from tree

holes and ground pools as a precedent, suggested that although the original

kyondawensis larvae were collected in ground pools, they have the appearance

Harrison and Scanlon: Subgenus Anopheles in Thailand 181

of a tree hole species and possibly the species normally occurs in tree holes.

Unfortunately, there have been no other collections of kyondawensis in Burma

(Khin- Maung-Kyi 1971a) to resolve this problem. During the last 12 years

hundreds of tree hole and forest ground pool collections were made throughout

Thailand without yielding a single specimen of kyondawensis. The single larva

of this species found in Thailand was not collected until 1966, when intensive

surveys of fresh water crab holes were being conducted in search of another

cryptic species, tigertti. The crab hole with the single kyondawensis larva

was located in a mountainous region under heavy shade in secondary wet for-

est, and also contained larvae of Culex (Lophoceraomyia) bengalensis Barraud

and an undescribed species of Uvanotaenia. It is not currently possible to de-

fine the larval requirements of kyondawensis. However, the collection of this

species from a fresh water crab hole is highly suggestive and could explain

the previous paucity of collections, because the crab hole habitat has been

ignored in most countries in the Orient. The kyondawensis larvae collected

by Abraham in Burma, came from small shallow shaded pools of clear water,

along the sides of streams in hill forest, and were associated with larvae of An,

bengalensis and leucosphyrus Donitz. It seems possible that the kyondawensis

larvae may have been swept out of crab holes into the ground pools by high

water. This contention is supported by the presence of balabacensis (= leu-

cosphyrus of early Burma workers, Khin-Maung-Kyi 1971b). Larvae of bala-

bacensis normally occur in small temporary ground pools that follow flooding

or rains.

ANOPHELES (ANOPHELES) SINTONOIDES HO

(Figures 94, 95, 96)

Anopheles (Anopheles) sintonoides Ho 1938a: 279 (**, 2, P*, L*); Reid and

Knight 1961: 492 (L); Reid 1963: 113 (L); Reid 1968: 223 (o*, 9*, P*,

L*); Rattanarithikul and Harrison 1973: 4 (L*).

This is a brown unicolorous species in the adult stage, which can be con-

fused very easily with members of the aitkenii group. The adults can be

recognized by the broad vertex scales on the head and the absence of scales

on the anterior pronotal lobe. The pupa has an unusual, simple type trumpet

with a very short meatal cleft, a very long seta 5 on the abdominal segments

and a hooked (Cellia-like) seta 1-P. The larva is easily recognized by the

reduced setae 5, 6, 7-C on the head, the large spine-like stellate setae and

the ventral and lateral parts of the thorax and abdomen covered with spicules.

FEMALE (Fig. 94). Unicolorous brown. Head. Palpus slender with 2

apical segments slightly swollen, with decumbent scales; palpus shorter than pro-

boscis, ending just before labellum; proboscis shorter than forefemur; clypeus

without scales; vertex with erect broad brown scales with truncate tips; inter-

ocular space very narrow, with short white scales and long dark brown frontal

setae; pedicel without scales; flagellomere 1 with dark basal scales; antennal

whorl setae very long, 4 - 5 times as long as flagellomeres. Thorax. Anteri-

or promontory without scales; scutal integument light brown; scutum without

scales, with mixed long and short brown setae in anterior promontory, acro-

stichal, dorsocentral, lateral prescutal, fossal, antealar, and supraalar

groups; prescutellar area bare; scutellum with long and short dark brown

setae, without scales; anterior pronotal lobe without scales, with long sparsely

scattered setae; pleural area pale; pleural setae: 1 propleural, 3 - 8 spiracu-

182 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

lar, 1 - 3 prealar, 2 - 4 upper and 2 - 4 lower sternopleural and 2 - 5 upper

and 0 lower mesepimeral. Wing. Unicolorous with narrow dark scales; hu-

meral crossvein without scales; cell Ro twice as long as cell Mo; cell Mog usu-

ally less than half as long as stem (from crossvein). Halter. With dark scales

on knob. Legs. Coxae without scales, upper midcoxa with 1 - 2 setae; legs

slender, entirely dark scaled. Abdomen. Mottled gray-brown with long brown

setae, without scales.

MALE (Fig. 94). Like female except antenna with more and longer whorl

setae; palpus slightly shorter than proboscis, with 2 apical segments flattened

and club-like. Genitalia. Basimere without scales, with 2 long parabasal

spines arising from prominent tubercle, outermost spine longest; ventromesal

spine on basimere long, arising about 0.66 distance from base; tergum IX with-

out caudal processes; claspette apparently with 3 lobes and approximately 6 -

8 separate curved setae; larger setae on dorsal and middle lobes of claspette

with stout bases and partially flattened; ventral lobe of claspette with smal-

ler setae and spicules; aedeagus narrowing to truncate tip, with fine hair-like

spicules on lateral aspect of distal 0.33.

PUPA (Fig. 95). Integument tan to light brown with trumpets more darkly

pigmented; most setae long, slender and simple, or with fine distal branches.

Cephalothorax. Seta 6-C equal or longer than 7-C; 10, 12-C very long, stout

and usually with branches arising on distal half; 11-C very small and short,

with distal branches. Trumpet. Simple, with shallow, widely rounded meatal

cleft; meatus 0.50 - 0.67 as longas trumpet. Abdomen. Seta 0, on II - VII

short and simple; seta 1 usually weakly developed on abdominal segments;

1-II with 5 - 12 branches, more strongly developed than 5-II; 1-III often more

strongly developed than 5-III; 1, on IV - VII much weaker and shorter than seta

0, on IV - VII; 1-IV with 2 - 7 branches; 1-VI with 3 - 6 branches; 1-VII long-

er than other setae 1, 0.50 - 0.67 as long as 5-VII; 1-VII simple or with 2 - 6

branches; 3, on II - III stouter than setae 1, 5, on II - III; 4-VII simple; 5-I

approximately equal length of 2-I; 9-III may be weaker or stronger than 1-III;

0, on IV - VII with stout central stem, as long or longer than segment; 5-IV

with 6 - 15 branches; 9-VII with 3 - 13 branches; 6-I much longer than segment

I; 6-II with distal branches, approximately equal length of segment II; 6-VI

nearly 2 times as long as 9-VI; 6-VII approximately 0.5 as long as 9-VII; 7-1

with many distal branches, approximately 0.67 as long as 6-I; 7-VII simple,

approximately 0.75 as long as segment VIII; 9, on IV - VII long, slender, tap-

ering to sharp point, 0.33 - 0.50 as long as segments; 9-IX short spine-like;

10, on IV - V approximately 0.5 length of segments. Paddle. Elongate: re-

fractile margin 0.33 - 0.50 length of paddle; fringe hairs on both margins of

paddle, long and dense, 0.5 length of 1-P; outer margin of paddle evenly con-

vex on distal half; 1-P very long and filiform with hooked tip, often bent in

middle; 2-P variable, simple and stout to weak with 2 - 4 distal branches.

LARVA (Fig. 96). Head darkly pigmented; thorax and abdomen with mixed

simple or compound spicules on ventral and lateral aspects. Head. Antenna

wider at base than apex, slightly crooked with minute spicules on inner sur-

face; 1-A simple or bifid, inserted on basal 0.25 - 0.33 of antenna; 2, 3-A

with one edge serrate; 4-A simple or with 2 - 6 branches, twice as long as 2,

3-A; 2-C long, simple and slender, with bases touching or very close together,

much closer together than bases of 2, 3-C on one side; 3-C short, stout and

simple or split into 2 - 6 branches (often spike-like), 0.25 - 0.33 as long as

2-C; 4-C stout, simple or with 2 - 9 branches, very wide apart and twice as

long as 3-C; setae 5, 6, 7-C with stout central stem, few branches and short;

o-C simple or with 2 - 7 branches, extending cephalad approximately to level

Harrison and Scanlon: Subgenus Anopheles in Thailand 183

of 4-C aveoli; 7-C with 6 - 13 branches; 8, 9-C short, simple or bifid; 11-C

shorter than antenna without branches at base, with numerous distal branches

beginning nearer to base on inner side of seta; 8-MP on maxillary palpus with

stout stem and branching beginning about midway along stem; some mouth

brushes with short frays, not pectinate. Thorax. Seta 0-P short simple or

with 2 - 3 branches; 1-P with 3 - 10 branches; 2-P with 7 - 22 branches, aris-

ing on large tubercle, as long as 4-P; 11-P short simple or with 2 - 4 fine dis-

tal branches; 13-P with 6 - 11 partially spiny stellate branches; 14-P with

9 - 20 thick spiny stellate branches; 4-M short with 3 - 6 branches; 6-M sim-

ple, stout, arising from pigmented tubercle and very long; 14-M with 5 - 9

spiny stellate branches; 3-T lightly pigmented, often with slightly flattened

leaflets without filaments; usually one long pleural thoracic seta on pro-

meso- and metathorax with short lateral branches. Abdomen. Anterior ter-

gal plates small, less than 0.25 as wide as segment; small posterior tergal

plates on II or III - VII; seta 0, on II - VIII small and simple; 1-I very small

and simple; 1, on II - VII with flattened pigmented leaflets with filaments;

2-I very small and simple; 5-VIII with spiny stellate branches; 6-III with

short lateral branches, as long as 6-II; 6, on IV - VI approximately 0.67 as

long as 6-III, with 2 - 4 long branches; 6-VII very short with 3 or more bran-

ches; 9, onI - VI very large, with spiny stellate branches; 11-I large with

spiny stellate branches; seta 13, on I - VI large with spiny stellate branches;

dorsal setae not developed into spiny stellate setae; pecten plate with approxi-

mately 20 teeth with strong basal and lateral denticles; seta 2 on pecten plate

with 2 - 4 distal branches; stigmal knob on median dorsal valve of spiracular

apparatus as darkly pigmented as remainder of valve; 1-X with 2 - 4 distal

branches, inserted near edge of saddle.

TYPE-DATA. The type-locality for sintonoides is Hainan Island, Kwang-

tung Province, China. A holotype “ and allotype ? are reported (Ho 1938a)

to be deposited in the Fan Memorial Institute of Biology, Peking (Peiping),

China. We were able to borrow a single topotypic 2 from the original collec-

tions (specimen #99) from the BMNH. This ¢ appears identical to the Thai-

land specimens.

DISTRIBUTION (Fig. 95). This species is known from China, Malaysia

(Peninsular), Thailand and South Vietnam (Reid 1968). Specimens (11°, 169,

86 larvae and 35 larval and pupal skins) in the USNM were examined from

the following provinces of THAILAND; Chanthaburi, Chumphon, Nakhon Nayok,

Nakhon Ratchasima, Nakhon Si Thammarat, Narathiwat, Phangnga, Phat-

thalung, Phuket, Ranong, Songkhla, Trang and Yala. A single female in the

BMNH was examined from CHINA (Hainan Island). Additional specimens

from SOUTH VIETNAM were examined in the Institut Pasteur, Saigon. The

distribution of sintonoides as depicted in Fig. 95 is probably a fairly accur-

ate delineation of its distribution in Thailand. Numerous tree hole-bamboo

collections have been made in the drier western and northern portions of

Thailand, yet no specimens of sintonoides were collected. This species

seems to be closely associated with tropical wet forest vegetation, which is

primarily restricted to the southern-southeastern portions of Thailand. A

considerable area of wet forest still occurs in Khao Yai National Park north-

east of Bangkok, and this is the area (Nakhon Nayok and Nakhon Ratchasima)

where the most northern collections of sintonoides were made. Vertically,

sintonoides has been collected between 5 - 800 m elevation in Thailand. Col-

lections in Hainan were listed as 600 m or more (Ho 1938a), and Reid (1968)

lists the Malaysian specimens collected at about 305 m. Apparently, the

elevation of the wet forest vegetation has little influence on the distribution of

184 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

sintonoides. However, since this is a forest species and much of the lowland

wet forests in Thailand have been cut and cleared, it will probably be encounter-

ed more often in the rugged mountainous areas that still retain primary and

secondary wet forest.

_ TAXONOMIC DISCUSSION. There are few Anopheles species in Thailand

with which this species can be confused. The adults resemble members of

the aitkeniz group in being unicolorous, but sintonoides has broad erect scales

on the vertex, while members of the aztkenii group have narrow erect scales

on the vertex. The male genitalia of sintonoides are distinct on the basis of

the fine hair-like spicules on the aedeagus and the separate claspette setae.

Adults of brevipalpis, currently known only from Malaysia and Indonesia, may

occur in Thailand, and could be confused with sintonoides. However, 0revi-

palpis has scales on the anterior pronotum, while sintonoides only has setae

on that structure.

We agree with the assignment (Reid and Knight 1961) of sintonoides to the

culiciformis species group, for as noted by Ho (1938a), the larva and male

genitalia of sintonoides are very similar to those of sintoni. Larvae of both

species exhibit spiny stellate setae (only 14-P on sintoni), and both species

have short hair-like spicules near the tip of the male aedeagus. The aedeagus

of both alongensis and culiciformis have leaflets rather than spicules. Unfor-

tunately, the pupa of sintoni is undescribed and cannot be compared with the

pupa of sintonoides.

The pupa of sintonoides has several unusual characters for the subgenus

Anopheles in Southeast Asia, and is easy to differentiate from the other pupae

known from that area. The pupal trumpet on sintonoides, although simple,

has the meatus unusually long and more like the trumpets found on Nearctic

and Palearctic members of the plumbeus species group. Seta 1-P on sinton-

oides is long, simple and filiform, usually curved and with the apex hooked.

This character is similar to 1-P structure seen on pupae of the subgenus

Cellia. The unusually long seta 5 on IV - VII is also of value in distinguishing

this species.

The larva of sintonoides is unique in Southeast Asia because of the spiny

stellate setae and the thick covering of spicules on the ventral and lateral

aspects. However, these characters and the thickness and number of branches

on setae are variable. Reid (1968) pointed out several differences between the

pupal and larval descriptions (Ho 1938a) of Hainan specimens and those found

in Malaysia, South Vietnam and Thailand. Since we had more specimens to

study, we found a wider range of variation on Thailand sintonoides than was

found by Reid. Thus, some of the differences he noted are no longer signifi-

cant, however, several of those differences remain. Hainan pupae are des-

cribed and illustrated as having seta 5 on the abdomen as considerably longer

than the segment, and seta 1-P straight. Thailand pupae have seta 5 equal

to or only slightly longer than the segment, and seta 1-P curved, with the

apex hooked. Hainan larvae were described and illustrated with seta 14-P

having 16 - 20 branches and seta 2 on the abdomen stout and spinose. Thai-

land larvae have seta 14-P with 9 - 17 branches and seta 2 on the abdomen

only slightly spinose. It should be pointed out that these observations, as

were Reid's, are based on Ho's description, and that specimens from Hainan

were not available for study. Reid (1968) considered these differences as

probably due to geographical variation in sintonoides, rather than distinctive

characters for 2 taxa, and we are in total agreement. As pointed out by lIyen-

gar (1930), Rosen and Rozeboom (1954) and Colless (1956), larvae that occur

in tree hole habitats can exhibit a tremendous range of environmentally induced

Harrison and Scanlon: Subgenus Anopheles in Thailand 185

setal variation. Consequently, the differences between the Hainan and the

Malay-South Vietnam-Thai specimens are not exceptional. Setae on Thailand

larvae were very variable between localities and even the type of container

had some effect on the thickness and branching of setae. Particularly variable

were setae 3, 4-C which were stout and spine-like on some larvae, or fine

and simple or with fine distal branches on other larvae. Several larvae col-

lected in a banana stump were noticeably different from those found in tree

hole or bamboo habitats. Those from the banana container had the spiny stel-

late setae more slender and less stout, while the other setae were slender and

simple or with fine distal branches. These larvae also had fewer spicules

covering the ventral and lateral aspects and those present were primarily single

rather than multiple spicules.

Two anomalies were noted on the immatures. The 1st was a larva with

several short frays on seta 2-C. The 2nd was a pupa with seta 2-P as long and

as stout as a normal seta 1-P. Further, all the larvae that we examined had

seta 11-I spiny and stellate | seta 10-I as listed by Reid (1968) ].

BIOLOGY. When this species was first collected in Thailand it was con-

sidered relatively rare, but extensive treehole-natural container collections

now reveal it is fairly common in the southern half of Thailand. As was noted

with the 2 other natural container breeding Anopheles in Thailand, i.e.,

asiaticus and interruptus, a single natural container rarely contained more

than 6 larvae, and 1 - 3 was the normal number per collection. These low

numbers per container probably represent an intrinsic survival mechanism

that results in a wider dissemination of eggs by a more complete utilization of

the available oviposition sites by the females. Reid (1968) suggested the frayed

mouth brushes may indicate that larvae of sintonoides are sometimes preda-

ceous. This behavior trait would also reduce the number of specimens collec-

ted per container. Immatures have been collected in the following habitats in

Thailand: large and small tree holes, holes in stumps, holes between tree

roots, a banana stump, pandanus axils, bamboo stumps, bamboo internodes

and split bamboos. These sites ranged from ground level to 2 m above the

ground in partially to heavily shaded primary and secondary wet forest, pri-

mary and secondary bamboo groves, a mangrove thicket and a rubber planta-

tion. As already noted these sites ranged from 5 m (mangrove) to 800 m (ban-

ana stump) elevation. Except for one female, all of the adults we examined

were reared from immatures. The single female was collected in 1962 be-

tween 1900 - 2000 hours, biting man in Nakhon Si Thammarat Province. Ex-

tensive human biting collections were made within the vicinity of tree holes

from which sintonoides larvae were collected, but these failed to yield other

adult sintonoides. This species is almost certainly not involved in the trans-

mission of human pathogens.

186

Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

APPENDIX A:

KEY TO THE FEMALE ANOPHELES MOSQUITOES OF THAILAND -

AN AID FOR RAPID IDENTIFICATION IN MALARIA FIELD RESEARCH*

4(3).

(4).

6(5).

8(7).

9(3).

10(9).

11(10).

12(10).

Wings with contrasting pale and dark scales. ........... 2

Wings uniformly dark, without pale scales... ..........-. ol

Leading margin of wing with at least 4 separate dark areas involving

both veins Cand R-R,........ (Subgenus Cellia).... 3

Leading margin of wing with less than 4 separate dark areas involv-

ing veins Cand R= Ry... 2. ss (Subgenus Anopheles). .. .27

Hind@irsomeres 3, 4:and S entirely white: . soi. eee a 9

Hindtarsomeres 3, 4 and 5 not entirely white. ............ 4

Hindtarsomere 5 entirely white. 23S ek a eae 14

Hindtarsomere 5 not entirely white. .. 0. 2.0. oe 5)

Tibio-tarsal joint of hindleg with large white band. ........ 15

Tibio-tarsal joint of hindleg without large white band. ....... 6

Apical hall of proboscis with pale scales. 2.0. . oe es 19

Apical halt of proboscis dark. o:. S62 ei ee eee 7

Femur, tibia and tarsomere 1 speckled with pale scale patches. . 21

Femur, tibia and tarsomere 1 without pale scale patches. .... 8

Foretarsi with broad pale bands crossing joints; mid- and hindtarsi

with pale bands slightly wider than diameter of tarsomeres. .. 22

Tarsi with only narrow apical pale bands, or entirely dark. ... 24

Palpal segments 2 and 3 with pale spots, subapical pale band broad,

approximately equal to apical band. .......... splendidus

Palpal segments 2 and 3 without spots, subapical pale band narrow,

approximately half the length or less of the apical band. .... 10

Femur, tibia and tarsomere 1 speckled with patches of pale scales.

Femur, tibia and tarsomere 1 without patches of pale scales... 12

Abdominal terga VI and VII with golden scales; vein Cu without dark

Scare March at Origin Of Vem Cuy. ee Jame sit

Abdominal terga without golden scales; vein Cu with dark scale patch

at OVE On Vel CU ys 6 is Go eee vamsayt

Vein Cu and Cug mostly dark scaled, with area of dark scales at ori-

OU OM Ce er a ae, he Ge eee annularis

Vein Cu and Cug mostly pale scaled, without dark scales at origin of

VOD CO rh Ee eG Cae hee eG 13

*Includes subgenus Cellia.

13(12).

14(4).

15(5).

16(15).

17(15).

18(17).

19(6).

20(19).

217).

22(8).

*Adults of these 2 species often not separable without associated immature skins.

Harrison and Scanlon: Subgenus Anopheles in Thailand 187

Wing with presector dark mark on vein R usually reaching or overlap-

ping distal end of humeral dark mark on costa, on both wings.

nivipes*

Wing with presector dark mark on vein R not usually reaching back to

distal end of humeral dark mark on costa, or only reaching that

mark on-me wins OMe Ws. VOM ia oe he a philippinensis*

Femur, tibia and tarsomere 1 speckled with patches of pale scales;

palpi usually with 3 paleybands. | soe Sk ee maculatus

Femur, tibia and tarsomere 1 without patches of pale scales; palpi

usually with + pale bands. sre eee Need ay karwavi

Basal dark mark on wing vein R with one or more pale interruptions on

atleast One Wiig ar... this Geared BS) ee See Sl i ae 16

Basal dark mark on wing vein R without pale interruptions. .... 17

Hindtarsomere 4 with distinct basal pale band.

balabacensis balabacensis

Hindtarsomere 4 without basal pale band. . . balabacensis introlatus

Palpi noticeably shorter than proboscis; proboscis with narrow pale

band aAbapex proximal to labeltae SS ee i ee Oe a 18

Palpi not noticeably shorter than proboscis; proboscis without pale

DAG ae eens tO 2 oe Re a ee SPEND Woe a, rviparis macarthuri

Palpi with very narrow pale bands, apical segment merely tipped with

WARDS eS CAC 155 ht DARE os ty iat ret ROUT Rees eo ee UD ie agree hackevi

Palpi with broader pale bands, apical pale band approximately as

broa@as'preapicaldark bands sy igwscieiw be us oh woe pujutensis

Femur, tibia and tarsomere 1 dark scaled, without speckled pale

seale patches.) 7b SN eee Tete Me the ae Bae aconitus

Femur, tibia and tarsomere 1 speckled with pale scale patches. . . 20

Abdominal sterna with median tufts of dark scales; hindtarsomeres

with brogmspale hands: Ui eaves Core emi sete RS aek kochi

Abdominal sterna without scale tufts; hindtarsomeres with narrow pale

TEC ORE PNG ae Bhi eC A IRE og ii, tn Rap, Wena, Brag tessellatus

Palpi with only one broad apical pale band; palpal segment 2 without

dorsal PATE “S POEs Mog) BR, ee IS A ee sundiacus

Palpi with 2 broad apical pale bands; palpal segment 2 with dorsal pale

SPOT ne Ee Re ee RLS PR men ahi ONUENM ot U9 bg PRON stephensi

Palpi with apical pale band 3 - 4 times the length of preapical dark

TOBA a a a Se TS an: rama iets tae ie Aer cael aa i vagus

Palpi with apical pale band 2.5 times or less the length of preapical

Chere. earns MR IG a) ae aR ci es ee etm 2 29

188

23(22).

24(8).

25(24).

26(25).

27(2).

28(27).

29(28).

30(27).

31(30).

32(31).

Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Palpi with subapical pale band usually 0.33 or less as long as pre-

apical dark band, which is 0.5 or more as long as apical pale

1H SUR ROE A gO CORR UCU aoe DN ny UO NA US RG RO PRO at Alea aR subpictus

Palpi with subapical pale band usually 0.5 or more as long as preapical

dark band, which is often less than 0.5 as long as apical pale band.

indefinitus

Scutum with scattered short white scales; vein R, with small white

spot between subcostal and preapical pale spots, at least on one

WE ie gas estes aes wh aad Pasture barred ae ak iit vie Ne ve jeyporiensis

Scutum with scales only on cephalic end; R; without pale spot between

subcostal and preapical pale spots. ........+4.....-. 29

Base of vein R just distal to remigium, with white or yellow-white

eee Miah Loti ual Ua gh ha aa TaN HUE Vi TS MG ONT hey alarm ue bly ae Gomi MinimuUs

Base of vein R just distal to remigium, with patch of gray or black

So a oT eee eee es are ee et ae er ae 26

Vein R4i5 dark scaled except for small pale spots at base and apex;

palpi with preapical dark band much longer than apical pale band.

culicifacies

Vein R4;5 mostly pale scaled, with small dark spots near base and

apex; palpi with preapical dark band and apical pale band approxi-

Pee POT og sc ania ce ee a Reel ie lm a oe. 8 teeta pampanat

Hindleg with femur or tarsomeres 1 and 2 with outstanding tufts of erect

Pere WILE SCALC Be ee Na aa ie em Ut beiMs Aa hws ee 8 28

Legs with clusters of erect dark scales on tarsomere 2 of midleg and

tarsomeres 1 and 2 Of Nindlee. wis ce ees (owe. ee a. eye bulkleyt

Legs with tufts of erect scales only on hindfemur. ......... 29

Abdominal segment VIII with pale golden scales; wing without apical

pale fringe spot at R445; subcostal pale spot of costa extending onto

MINE ee) aie ue eds a es ce tea aah ve dah lyre astaticus

Abdominal segment VIII without pale scales; wing with narrow apical

pale fringe spot at R4,5; subcostal pale spot when present, confined

1O\COete and Vip Of SUDCOSTA. ek we wes ee ew eb interruptus

Clypeus with patch of dark scales on each side; basal 4 - 8 antennal

flagellomeres with pale scales. .... hyrcanus group... . 33

Clypeus without scales; 1st (basal) antennal flagellomere only with

ee a ae a! ke ak Nags vtawiig ALO d ky bial a 8g i ely 31

Abdominal sternum VII with tuft of black scales.

barbirostris group.... 41

Abdominal sternum VII without scales. . ow) ke ees) eee 0 ee 32

Hindtarsomere 5 mostly dark scaled. . . umbrosus group.... 46

Hindtarsomere 5 all white. . . . albotaeniatus group... . montanus

Harrison and Scanlon: Subgenus Anopheles in Thailand 189

33(30). Hindtarsomeres with broad pale bands, at least one tarsomere 4 with

basal PaLecband: Or Patch. wwik vin oe Bul a. eR Re 6 ahbes 34

Hindtarsomeres with apical pale bands only. ........2.2... oF

34(33). Basal dark mark on vein Cu short, separated by its own length or more

from upper dark mark on vein 1A; base of costa with scattered pale

scales, frequently with small humeral pale spot; dark mark at ori-

gin of vein Rg well defined, scales between dark mark and fork

mostly white; vein Cug with pale fringe spot......... nitidus

Basal dark mark on vein Cu long, approaching within own length or

less of upper dark mark on vein 1A; base of costa dark scaled or

with few scattered pale scales, no pale humeral spot; dark mark at

origin of Rg poorly defined, scales between dark mark and fork

mostly dark; vein Cug with or without pale fringe spot. .... 35

35(34). Humeral crossvein with patch of dark scales; remigium mostly dark

scaled; basal 0.33 and preapical dark mark on vein R - R; dark

scaled, or with very few, pale scales. cieidic ws 2 elete . lel 36

Humeral crossvein without scales; remigium mostly pale scaled;

basal 0.33 and preapical dark mark on R - Ry usually with many

NO MI hig ts dik ak eh esas ok Wa Uae uma, de mee es ag ce peditaeniatus

36(35). Hindtarsomere 4 with broad dark band, at least 0.6 length of segment;

hindtarsomere 5 without or with very narrow (0.25 or less of seg-

ment) pale basal band; base of costa usually with several scattered

pale scales; apical dark mark on vein Cus short, rarely as long as

Sica (erk mars On VAs oc. yk a oi an sae wk nigerrimus (in part)

Hindtarsomere 4 with narrow dark band, 0.5 or less (lacking in rare

cases) length of segment; hindtarsomere 5 with pale basal band on

0.6 or more of segment; base of costa without pale scales; apical

dark mark on Cug long, usually equal or longer than apical dark

TO gi cei taille Wu RA veia hi gui el Mile gow a erm eee argyropus

37(33). Wing apex with short pale fringe spot not extending posteriorly beyond

VOU Pai ah. Mule tere aie Wile aed DY + cE) save oa Sed a ae 38

Wing apex with long pale fringe spot extending posteriorly at least to

aa a rs MMR de RO on NO ey] USE Le Mae Pe EE POMRPR RPE HI orn SD 39

38(37). Humeral crossvein with dark scales; remigium mostly pale scaled midcoxa

with white scales; smallest member of the groupin Thailand. .pursati

Humeral crossvein without scales; remigium mostly dark scaled; mid-

cone without pale scales sielwiin duced aber ioe eia lesteri pavaliae

39(37). Vein Cu with long basal dark mark, at least twice as long as most

basal pale mark (when present) on Cu, and approaching within its

own length or less of upper dark mark on 1A; costa usually with

several scattered pale scales on basal 0.33; remigium mostly dark

scaled; humeral crossvein with dense patch of black scales.

nigerrimus (in part)

Vein Cu with short basal dark mark, equal or less length of most basal

pale mark on Cu, and usually separated by its own length or more

from upper dark mark on 1A; basal 0.33 of costa usually entirely

dark scaled; remigium mostly pale scaled; humeral crossvein bare

OP WIHT TOW BOR. ie ee ee Ce Nea ee . 40

190

40(39).

41(31).

42(41).

43(42).

44(42).

45(44).

Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Wing pattern blurred; tip of vein Ry dark scaled; apical wing fringe

spot long, beginning at or above vein Rj; preapical dark mark on

vein Ry with some pale scales; vein Cug usually with pale fringe

SOO ee er ie le ae ee se ee sinensis

Wing pattern sharp, dark marks short and well defined; tip of vein R,

pale scaled; apical fringe spot shorter, beginning at Ry; preapical

dark mark on vein R, without pale scales; vein Cug eiifhout pale

(PING Spetye: 29. ee Leet ee ee ee ee crawfordi

Apical fringe spot at wing vein R4i5 extending back to vein M449; ab-

dominal sterna without pale scales. ......... barbumbrosus

Apical fringe spot at Ry,5 narrow, not reaching back to vein Mj,9;

sterna with at least a oe Mediah pale scalése : 6... we we 42

Wing apex with 2 narrow pale fringe spots, no pale fringe scales at Ro;

midtarsomeres usually Unbanded. 7 ee 8s 4

Wing apex with 3 narrow pale fringe spots, middle spot at Ro, or with

only 2 pale fringe spots, but upper spot wide with pale scales down

to include Ro; midtarsomeres usually with narrow apical pale bands.

Abdominal sterna with many white scales scattered between median

white scale tufts and white scale row on each lateral margin.

campestris*

Abdominal sterna with fewer white scales, primarily concentrated in

median tufts and lateral rows, few scattered in between.

barbirostris*

First foretarsal pale band short, 0.5 or less as long as foretarsomere

5 and rarely crossing joint onto tarsomere 2; median pale scales on

abdominal sterna II - VI usually total 0 - 20; central to southern

Oe a en ee go ah ee ee ah et oe ol ee ok a te a ST NTE es hodgkini

First foretarsal pale band long, more than 0.5 as long as foretarso-

mere 5 and usually crossing joint onto tarsomere 2; median pale

scales on sterna II - VI totaling more than 20; extreme southern

i ee ee a ee UO eee 45

First foretarsal pale band longer than foretarsomere 5, with 0.33 of

band on base of tarsomere 2; 2nd foretarsal pale band also crossing

joint, with 0.25 - 0.33 of band on base of tarsomere 3; costa may

have some pale scales on preapical dark mark. .... pollicaris

First foretarsal pale band seldom longer than foretarsomere 5, with

0.25 or less of band on base of tarsomere 2; 2nd foretarsal pale

band rarely crossing onto base of tarsomere 3; costa without pale

Scales On preanical dark marks i. pes OE eae. donaldi

*This character is not always reliable, and in the absence of other adult differ-

entiating characters, these 2 species are identified best by rearing adults

with associated immature skins.

46(32).

47(46).

48(4'7).

49(48).

50(48).

51(1).

92(51).

5S(82);

94(52).

Harrison and Scanlon: Subgenus Anopheles in Thailand 191

Palpi with pale bands, apical segment usually entirely pale.

separatus

Palpieutitely cari Seales ee ae eee 47

Base of hindtibia with pale band 1.5 - 4.0 times segment width; base

of wing veins R and Cu usually with several scattered pale scales;

accessory pale fringe spot often present at apex of wing vein Ro.

roperi

Base of hindtibia dark scaled or with small pale spot; base of R and Cy

with dark scales; wing without fringe spot at vein Roses i 48

Propléewron’ witht is..6 setae on each side?) oh Gere cay eee 49

Propleuron without setae, or with 1 small seta on one side only. . 50

Upper mesepimeron with 9 - 19 setae and few inconspicuous dark

scales; tarsomeres all dark or with minute spots at joints. .baezai

Upper mesepimeron with 1 - 6 setae and no associated scales; fore-

and hindtarsomeres with small pale bands. ....... umbrosus

Wing vein 1A with pale scales proximal to median dark mark, infre-

quently with 2 - 3 dark scales near base. .......... letifer

Vein 1A with 5 or more dark scales at base, infrequently vein nearly

Sirdar SCAG as ee eee Pee. be, eee oS whartoni

Erect head scales long and narrow, only slightly expanded apically;

tlagellomeres without scales.) (> ae SRR en a og

Erect head scales broad, widely expanded apically; flagellomere 1

Nice CRN CONO Re). ee ee eee a a ee WS sintonoides

PGR gg a Re Sees, he TEE eae ae CE PIAS Se ae)

All abdominal. sésments unicolorous. 6... 20. ee SO Oa, o4

Abdominal segments IV and V pale. ........ aberrans (in part)

Abdominal segment IV only pale. ......... palmatus (in part)

Prescutellar space with short fine setae setae back to scutellum.

insulaeflorum

Prescutellar space with distinct median bare area immediately cephal-

20°tO Sushi ik ee ee Oe aberrans (in part)

bengalensis

fragilis

palmatus (in part)

stricklandi

ligertti

192

Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

APPENDIX B:

OMA DUIIKPWN &

SOUTHEAST ASIAN SPECIES OF THE SUBGENUS

ANOPHELES NOF KNOWN FROM THAILAND

SPECIES

acaci Baisas

ahomi Chowdhury

albotaeniatus (Theobald)

alongensis Venhuis

annandalei Prashad

balarensis Mendoza

bancrofti barbiventris Brug

borneensis McArthur

brevipalpis Roper

brevirostris Reid

collessi Reid

ejercitot Mendoza

franciscoi Reid

gigas Giles

hunteri (Strickland)

kweiyangensis Yao and Wu

lesteri lesteri Baisas and Hu

lindesayi Giles

manalangi Mendoza

noniae Reid

ohamai Ohama

pilinotum Harrison and Scanlon

pseudobarbirostris Ludlow

pseudosinensis Baisas

samarensis Rozeboom

saperoi Bohart and Ingram

similissimus Strickland and

Chowdhury

sinevoides Yamada

vanus Walker

wellingtonianus Alcock

DISTRIBUTION

Borneo, Philippines

India (Assam)

Indonesia, Malaysia

North Vietnam

India, Indonesia

Philippines

Indonesia

Borneo

Indonesia, Malaysia

Malaysia

Philippines

Borneo, Burma, China, India,

Indonesia, Malaysia, Philippines,

South Vietnam, Sri Lanka, Taiwan,

Indonesia, Malaysia

China

China, Japan, Philippines, Ryukyu

Islands, South Korea

China, India, Japan, Malaysia,

Nepal, Pakistan, Philippines, Taiwan

Philippines

Malaysia

Ryukyu Islands

Indonesia, Philippines

Indonesia, New Guinea, Philippines

Philippines

Ryukyu Islands

Malaysia

China, Japan, South Korea

Borneo, Indonesia, Philippines

Malaysia

Harrison and Scanlon: Subgenus Anopheles in Thailand 193

ACKNOWLEDGMENTS

Since this work began numerous individuals have supported, encouraged,

and labored for its completion. Without all of this help the task would remain

uncompleted. Hopefully, we have not omitted any of these people in the follow-

ing acknowledgments.

We are particularly indebted to the devoted field assistants in Thailand,

who collected, often under difficult circumstances, the majority of specimens

examined: Messrs. Sahem Esah, Kol Mongkolpanya, Prajim Boonyakanist,

Samarn Maniwongse, Sumeth Chunchulcherm, Attaya Boonyakanist, Surat Bandi-

thaya, Prachob Boonyakansit, Chaliou Diraphat, Voth Gengam, Sorasak Imvitaya,

Inkam Inlao, Somboon Kasemchit, Suraphol Kesornbupha, Thep Kowmongkol,

Somboon Maneechai, Kovit Mongkolpanya, Sanit Nakngen, Chumnong Noigamol,

Vichit Phunkitchar, Suvith Plicarmin, Manop Pookasorn, Anun Poosiri, Larp

Punthusiri, Chalong Ratanarati, Preecha Ruangvija, Lek Somchit, Thavi Staporn,

Sripong Suwanalong, Ruan Thaopha, Kitti Thonglongya, Amnat Timsuren and

Sanit Yanubol. Mr. E. L. Peyton, and Dr. Udaya Sandhinand and his team of

collectors, to include Mr. Manop Rattanarithikul, also deserve special recog-

nition for their valuable collections in Thailand. We are also very much indebted

to the devoted laboratory assistants in Thailand, who under the capable direction

of Mrs. Rampa Rattanarithikul, made preliminary identifications, mounted,

labelled and maintained the records for the specimens: Miss Pacharee Nawarat,

Miss Supanee Sandhinand, Mrs. Rachanee Likitvanichkul, Mrs. Suda Ratanawong,

Mrs. Prasertsri Rohitaratana, Mrs. Tasanee Keawrat and Mrs. Prachoub

Polpeuj. Weare very grateful to Dr. Douglas J. Gould, U. S. Army Medical

Component-SEATO, Bangkok, for his support, encouragement and direction

of the large scale field surveys and technical curatorial assistance required in

Thailand.

Special thanks are expressed to Dr. Botha de Meillon, for continuous:

support, encouragement and assistance while serving as Principal Investigator,

Southeast Asia Mosquito Project, and for reviewing the manuscript.

We are also deeply grateful to Dr. Ronald A. Ward, Walter Reed Army

Institute of Research and Medical Entomology Project (MEP), Smithsonian

Institution, for reviewing the manuscript, for many helpful suggestions, and

overseeing and assisting in the final preparations for publication. Special

appreciation is expressed to Lt. Colonel Bruce F. Eldridge, Walter Reed Army

Institute of Research and Mr. E. L. Peyton, Medical Entomology Project, for

continued support, many helpful suggestions and for reviewing the manuscript.

Special thanks are extended to Major John F. Reinert, Walter Reed Army

Institute of Research, for continued encouragement and many stimulating dis-

cussions. We are indebted to Lt. Colonel Hugh L. Keegan, Lt. Colonel Vernon

J. Tipton, Lt. Colonel Alexander A. Hubert and Captain Edward S. Saugstad,

during their respective tours as Chief, Department of Entomology, U. S. Army

406th Medical Laboratory, Japan, and their artist staff, who prepared most of

the illustrations. Thanks are also due to Mr. Vichai Malikul and Miss Thelma

Ford of MEP for preparing illustrations, and particularly Miss Gloria Gordon

(MEP), who prepared several illustrations and assisted in checking, correct-

ing and collating all of the illustrations.

The invaluable assistance of Dr. Peter F. Mattingly, British Museum

(Natural History), London, is most gratefully appreciated. Without his collabora-

tion over many years, this endeavor would have not achieved success. Sincere

thanks are given to Dr. Alan Stone (Retired) and Dr. Curtis W. Sabrosky,

Systematic Entomology Laboratory, U. S. Department of Agriculture, Washington,

194 Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

Dr. John A. Reid, British Museum (Natural History), Dr. Michel Quentin and

Dr. Jean Rageau, Office de la Recherche scientifique et technique Outre-Mer

(ORSTOM), Bondy, France, for their assistance and for permitting the exam-

ination of types and other specimens. Specialacknowledgments are expressed

to past and present staff members of MEP and SEAMP for their assistance,

particularly Mr. C. John Lane, Mr. Thomas Gaffigan, Mr. Robert Hochman,

Mrs. Phuangthong Malikul, Mrs. Peggy Gaffigan, Miss Anita Schmidt and

Mrs. Susan Hevel. We thank the Medical Department, U. S. Army, for financ-

ing trips to examine type-specimens at: British Museum (Natural History),

London; Institut Pasteur, Paris; Office de la Recherche scientifique et technique

Outre-Mer, Bondy, France and Zoological Survey of India, Indian Museum,

Calcutta, India.

We acknowledge and thank the following individuals and institutions for

their contribution and/or loan of specimens to SEAMP: Dr. P. H. Arnaud,

California Academy of Sciences, San Francisco, California; Dr. J. Bonne-

Wepster (Deceased), Instituut voor Tropische Hygiene en Geographische

Pathologie, Amsterdam, Netherlands; Dr. D. F. Bray, University of Dela-

ware, Newark, Delaware; Dr. E. H. Bryant, Jr., Dr. J. L. Gressitt and

Dr. W. A. Steffan, Bishop Museum, Honolulu, Hawaii; Mr. Y. Cheang,

Depamat (Minsante), Phnom Penh, Cambodia; Dr. R. F. Darsie, Jr., METC/

USAID, Manila, Philippines; Mr. H. Dybas and Dr. R. L. Wenzel, Field

Museum of Natural History, Chicago, Illinois, Dr. M. T. James, Washington

State University, Pullman, Washington; Dr. J. M. Klein, Institut Pasteur,

Phnom Penh, Cambodia; Dr. B. R. Laurence, London School of Hygiene and

Tropical Medicine, London, England; Dr. R. Lichtenberg, Naturhistorisches

Museum, Vienna, Austria; Dr. L. Lyneborg, Universitetets Zoologiske Muse-

um, Copenhagen, Denmark; Dr. L. L. Pechuman, Cornell University, Ithaca,

New York; Dr. H. D. Pratt (Retired), Communicable Disease Center, Atlanta,

Georgia; Dr. S. Ramalingam, University of Malaya, Kuala Lumpur, Malaysia;

Dr. D. H. Rees, University of Utah, Salt Lake City, Utah; Dr. H. R. Roberts,

Academy of Natural Sciences, Philadelphia, Pennsylvania and Dr. L. E.

Rozeboom (Retired), John Hopkins University School of Public Health, Balti-

more, Maryland.

We are grateful to the following military entomologists who contributed

specimens during their tours in Southeast Asia: Lieutenant E. M. Bravi,

U. S. Army; Major R. W. Intermill, U. S. Army; Lt. Colonel M. A.

Moussa, U. S. Army; Major R. E. Parsons, U. S. Army; Lt. Commander

H. P.. Shurtleff, U. S. Navy; Lt. Colonel R.:W. Upham, Jr. , Us S. Army

and Captain S. M. Valder, U. S. Air Force. Appreciation is expressed to

Miss Helle Starcke for typing much of the first draft and to Mrs. Janet Rupp

for typing parts of subsequent drafts and typing the manuscript for offset

reproduction. Finally, we are especially grateful to our wives, Carole and

Consuelo, for their patience and encouragement during this study.

Harrison and Scanlon: Subgenus Anopheles in Thailand 195

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1947. The geographical distribution of the mosquitoes of the Malay Archi-

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BRYDON, H. W., G. JOSHI and S. PRADHAN

1961. Anopheles distribution and its relation to the malaria eradication

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1964(1965). Keys to the anopheline mosquitoes of Cambodia with reference

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|. YOSHIGAKI

A.(Anopheles ) interruptus

Cambodia

9 |

ed Yy a

- {South

von ay es

ate) Ue Py |

A ( ks '

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Sof Lege: Ores

inet ; ns Pp

\} \-Wietnam ~s:

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Chg Sea)

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t ye y

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t 1

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i R

i \"

Y- \

Ie a

fh at

1 i: SAN NY ‘nid

5 N

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pel ° »:

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)

, b>

A. (Anopheles) interruptus solidi i

Fig.f9

Sete:

5 nied \ fe

Ca Dg OTS

SVG u.

NET VW

IW oP ‘

;

8 |

134,17 gfese

yO aly, S$ = 1

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4 IM VIE

: 2

Sonobe

A. (Anopheles ) interruptus

\\Y

SOV

. \

= IW

CL variation

A.(Anopheles) aberrans

6-MP

A. (Anopheles) aberrans

F ig. 18 A. (Anopheles) aitkenii GROUP

aberrans

MEMinEBmC .. . Compaeptreumenaeemmmateny .

palmatus

insulaeflorum bengalensis

Fig./9 = MALE GENITALIA A.(Anopheles) aitkenii GROUP

variation

aitkenii (INDIA)

Xe CL aberrans

variation

pilinotum

(PHILLIPINES)

Strickland/

tigertti

bengalensis

|. YOSHIGAKI Q

A. (Anopheles ) bengalensis

Burma

Cambodia

f--

urns

) t South

Bi LES 3 “Sy

Resid Pte SKE

, be

~~ rf A, ca

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Cy . ny

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i SD

1 ‘\

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1 rag

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i i

ais

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au DS

eh H ‘

} Malaysia .

) are

i .

A. (Anopheles ) bengalensis

id i,

; oN ; A f

i of |

Hosokawa

-—— 0.5mm ——CH

SS SNS

= VSN

= \= ——_

Sse = .

Hii

JIS

BA %|

= LEFTY ff. ORS

= = er

LE om

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Cambodia

Cae

)

t South

t hay ay %

L-/ J Parana

a aU

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2 eS

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a: fragilis

; x

a Malaysia KS h

; a 2

/ .

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iy y

a | “ Fe ie

VAY ip :

2 ey 21 , * Y

3 iP 14 1

A.( Anopheles) fragilis

Fig.85 a)

Cambodia

~s/

ENS

men, pach

“UY. 3

\} \-Vietnam

y ae if

i 2

a ‘

Sar

A.(Anopheles) tigertti

el4

SD Neanaconee

SS 5

Fig.87

| | W SN "%

5 Vv “Fi,

i |}

a) | LOS

TDi

Burma

4 A

i \ A

f DS ,

‘ 1

‘ ‘ 4

XN.

f 5

Sug

Cambodia

G ~

peel

ce eo “vv

\ South

ig ‘

TERRIEN aN ey

aye. ex NGS

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vy} Vietnam ~s

y =< te

= < 8S

ioral

i eA

aE 1 }

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i -

1 i“

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ae ha Ach

fa;

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A. ( Anopheles) insulaeflorum

Hosokawa

—

SZ a

a za

—

A. (Anopheles) insulaeflorum Hosokawa.

j) /

]

"A P

{ In 1

)

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i 4

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oy 4 aw : er ees

eh See \

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y t xe

/ > = se

‘.

wa, om zn

See Se a}

igh se 7

pred

Cambodia

Mew SB -BR ny

ere 2 <2? 2-8

ene ae ee DSg---Fy

a <- 8 so

oreo gaa Cae iad ta SN

<2-- 3? <_ s Cy .

4 s~-= ,

<> te

-s7

A. (Anopheles) stricklandi

Fig.90

US eAy

yj i (

; 4 G Wi

6—MP

TES

WW S

0//, \; Ns

Y . Al ;

A.(Anopheles) stricklandi Ka : f

tah

Foal

fez

wna yarn?

Cambodia

Proce

Nf °

/ \-Wietnam

) =

ich

Ye 7

v SS

{ Wisk)

1 » iat

{ Bes

eee

H t

( ‘

> o

‘

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tergal plate

Wilby

A a

Ny,

‘

ee <4

Zi, si

5 ey

4 Rs

F 7

Sonobe

A. (Anopheles) palmatus

a Ay

10 1)

ce ‘NG

\\? p

a i hj W 0 }

Ny “ \ \ f

6 |

Sh 2 a 7

tard

Vi ie

A. (Anopheles ) kyondawensis

A. ( Anopheles ) sintonoides

Cambodia Y | on [=>

(eu

{South

Bow in

SSRN rae J

Crd ° nt

\} ‘Vietnam ~s;

4 Ss Std

(a7 SS

K=kyondawensis

) ae ‘ o i)

#= sintonoides Pat “(f’

a Malaysia ae Sonobe

A. (Anopheles) sintonoides

seta 11-C

A. (Anopheles ) sintonoides

304

Contrib. Amer. Ent. Inst., vol. 12, no. 1, 1975

INDEX

Valid names are set in roman type, Synonyms and non-valid combinations

are italicized.

Italicized pages designate the primary treatment of the taxon.

Numbers in parentheses under a given species designate the figure(s) for that

species.

aberrans 1, 7, 24, 25, 145, 146,

148-150, 152, 154-157, 159-162,

166, 275.. TE. VOT (Gy. 77, “he,

79, 83)

acaci 14, 20, 147-149, 150,

192

aconitus 187

Aedeomyia 138

Aedes 20, 23, 70, 102, 143

Aedimorphus 102

ahomi 79, 80, 192

aitkenii 1, 147-149, 150-152, 155-157,

160-162, 169 (79)

aitkenii bengalensis 157

aitkenii Group 14, 15, 20, 27-29, 31,

107, 144-146, 147, 148, 155, 156,

160, 162-164, 169, 172, 173, 175-

177, 181, 184

aitkenii palmatus 173

aitkenii var. bengalensis 157

aitkenit var. insulaeflorum 167

aitkenii var. palmata 173

albopictus 70

albotaeniatus 31-33, 192

albotaeniatus Group 27-31, 32, 40,

80, 107, 188

albotaeniatus var. montanus 34

algeriensis 144

alongensis 177, 178, 180, 184, 192

Angusticorn Section 26-28, 132

Anisodera 133

annandalei, Anopheles 133, 140, 142,

143, 192

annandalei, Uranotaenia 124

annandalei var. interruptus, Anopheles

140, 142

annularis 186

anophelesica 162

Anopheles (Genus) 1-3, 15, 20, 21,

23-25, 48, 92, 113, 180, 184-186

Anopheles Series 14, 20, 26-29, 31,

135, 144, 145, 179

Anopheles (Subgenus) 1-4, 6, 7, 14-16,

Ae, NeO, 20, 26, So, Laaitees 175,

184, 186, 192

Anophelinae 21

151, 173,

argyropus 6, 7, 37-44, 50, 52, 53, 56,

60, 61, 69, 74, 189 (9, 13, 14, 15)

asiaticus 7, 25, 133, 134, 136, 138-

14.) 185, ' 188 (10, °%1, 72)

asiaticus Group 15, 27-29, 133-135

atratipes 20, 113, 144

baezail, 7, 8, 24, 28, 106, 107,

109, 113, 174-119, 121,101 (55,

56, Ot)

balabacensis 102, 181

balabacensis balabacensis 187

balabacensis introlatus 187

balarensis 31, 32, 192

bancrofti bancrofti, Anopheles 113

bancrofti barbiventris, Anopheles 118,

192

bancrofti Group, Anopheles 20, 40,

60, ib3

bancrofti, Wuchereria 75, 88, 93, 96,

114, 122, 124, 128, 132

barberi 148

barbirostris 2, 7, 20, 24, 78-83, 85-

100, 102, 190 (34, 35, 36)

barbirostris Group 15, 20, 21, 26-33,

40, 78, 80, 81, 86-88, 90, 92,

93, 96, 99, 102-104, 106, 107,

113, b75,0 168

barbirostris Subgroup 32, 33, 79, 99

barbumbrosus 6, 7, 29, 79-83, 86,

Gt, 88,90, 94.96.99, “102-105,

190 (49, 50, 51)

barianensis 177

bengalensis, Anopheles, 1, 7, 24, 145,

146, 148-152, 155-157, 159-166,

1600, OD, Via 10S, 1 to, 176, 181,

191 (78, 79, 80, 81, 82)

bengalensis, Culex 181

bentleyi 66, 68

bimaculiala 124

Bironella 21

borneensis 148-151, 192

brevipalpis 14, 31, 106, 107, 123,

184, 192

brevirostris 106, 107, 192

Bromeliaceae 24

Harrison and Scanlon: Subgenus Anopheles in Thailand 305

Brugia 33, 40, 50, 53, 57, 61, 65,

70, 75, 88, 89, 93, 96, 118

bulkleyi 7, 133, 134, 135, 188

campestris 1, 7, 20, 58, 79-83,

86-88, 89-95, 98, 99, 190

(37, 36) 39)

Cellia 6, 8, 14, 16, 24-28, 144, 145,

181, 184, 186

Chagasia 21

claviger 144

Coelomomyces 162

collessi 106, 108, 120, 123, 192

concolor 144

coustani Group 33, 52

erawfordi 7, 37-40, 42, 43, 45,

48, 54-58, 60-62, 64, 124, 131,

190. (9," 16," 17, °18)

Culex 36, 102, 167, 181

culicifacies 188

culiciformis 176, 177, 178-180, 184

culiciformis Group 20, 27-29, 31,

144-146, 176, 177, 180, 184

Cynocephalus 118

cynomolgi 97, 100, 128, 132

cynomolgi bastianellii 53, 58, 61, 65,

70, 88, 93, 114, 118, 122, 128

Dy near nigerrimus 75, 77

Dirofilaria 53, 65, 70, 89, 93, 105,

122, 124

donaldi 1, 6, 7, 79-83, 86-88, 94-99,

101, 190 (40, 41, 42)

Eichornia 65

ejercitoi 31, 32, 192

falciparum 96, 97

fragilis 7, 145, 146, 148-150, 155,

196, 158, 160, 161, 162-164,

166, 191 (79, 83, 84)

franciscoi 6, 79, 80, 86, 99, 192

gateri 106, 107, 115, 116

Gigantochloa 133

gigas 132, 144, 192

gigas formosus 21, 144

gigas sumatrana 21, 144

goryi 133

grandis 4

habibi 144

hackeri 187

hodgkinil, 7, 8, 79-83, 87, 94, 95,

97-102, 104, 190 (43, 44, 45)

hunteri 106, 107, 192

hyrcanus 36, 37, 52

hyrcanus aberration argyropus 50

hyrcanus Group 14-17, 20, 21, 26-31,

33, 34, 36, 38-40, 48, 50, 52,

of, 61, 65, 79, 80, 89, 106, 107,

128, 131, 175, 188

hyrcanus nigerrimus 50, 58, 68

hyrcanus separatus 128

hyrcanus sinensis 50

hyrcanus subsp. near sinensis 61

hyrcanus var, argyropus 50

hyrcanus var. nigerrimus 66, 70, 71,

13, %¢9

hyrcanus var. sinensis 45

indefinitus 188

indiana 162

indiensis 38, 58-61, 65, 68, 73

insulaeflorum 1, 7, 25, 145-149, 151,

156, 157, 166,°767-172, 276, 191

C16, 79, 87,88)

interruptus 7, 15, 133, 134, 140, 142,

143, 185, 188 (73, 74, 75)

jamesii 186

javanicus 114

jesoensis 45, 47

jeyporiensis 188

judithae 148

karwari 187

Kerteszia 24

kochi 187

koli 167

kweiyangensis 192

kyondawensis 7, 27, 28, 145, 147, 177,

178, 179-181 (93, 95)

Laticorn Section 26-29

lesteri lesteri 38-40, 50, 56, 63-65, 68,

192

lestevi Malayan form 62, 63

lesteri paraliae 7, 37-45, 48, 56, 57,

60, 61, 63-65, 78, 189 (9, 22, 23,

24)

lesteri Subgroup 38, 40, 64, 65, 74

letifer 7, 106, 108-110, 113, 114, 117

118-127, 191 (58, 59, 60)

letifer Subgroup 106, 108-110, 127

leucosphyrus 138, 181

306 Contrib. Amer. Ent. Inst.

lindesayi 192

lindesayi Group 32, 33, 144

longirostris 124

Lophoceraomyia 167, 181

Lophomyia 136, 138

Lophopodomyia 24, 135

Lophoscelomyia 136, 138, 139

Lophoscelomyia Series 14, 26-29, 132-

135

maculatus 187

maculipennis 21, 25

maculipennis Group 20, 144

malayi 33, 40, 50, 57, 61, 65, 70,

719, 88, 89, 93, 96, 118

manalangi 79, 80, 86, 99, 192

Mansonia 58, 93

marteri 144

martini 83, 86

minimus 188

minutus 66, 68

montanus 6-8, 16, 20, 27-33, 34-36,

102, 188 (5, 6, 7)

Muscidus 102

Myzomyia 128, 130 |

Myzorhynchus 50, 70-73, 83, 105,

110, 112, 128, 136

Myzorhynchus Series 14, 26-29, 30,

32, 33, 144

nigerrimus 6, 7, 37-44, 48, 49, 52,

Oo, 600, 61, 65-70, 73; 76,. 78,

104, 189 (9, 25, 26, 27)

nigerrimus Subgroup 38, 40, 42, 64,

nitidus 7, 37-44, 48, 56, 58-62, 64,

69, 73,. 74, (8, 189 (8, 19, 20;

21)

nivipes 187

noniae 133, 139, 140, 192

novumbrosus 110, 113

Nyssorhynchus 24

obscurus 33

ohamai 31-33, 192

omorii 148, 177

pahangi 93

pallida 1, 150, 157, 159

palmatus 1, 7, 8, 17, 145, 146, 148,

14950196." 107.) 266, 273, 165, 1-76,

191 (78, 79, 91, 92)

pampanai 188

wOl. 22 sme. ctyc LOTS

Patagiamyia 128

peditaeniatus 7, 34, 37-44, 52, 53, 56,

60, 61, 64, 66, 68, 69, 71-75, 104,

189 (8, 28, 29,. 30)

philippinensis 187

pholeter 167

pilinotum 148-151, 156, 168-170, 172,

173, 176, 192 (79)

pinjaurensis 148-151, 176

Pistia 78

Plasmodium 53, 58, 61, 65, 75, 88,

g2, 93, 96, 97,100. 114,. 118,

122; 128, 132

plumbeus Group 144, 148, 177, 184

plumiger 45, 47

pollicaris 1, 6, 7, 20, 79-82, 87, 98,

99, 100-102, 113, 190 (46, 47, 48)

pseudobarbirostris 113, 192

pseudopictus 52

pseudopunctipennis Group 144

pseudosinensis 6, 37-39, 68, 74, 192

pujutensis 187

punctipennis Group 144

pursati 6, 7, 31, 37-39, 41-44, 60,

63, 64; 75, 71, :°-78,. 189:(8,..31,

a2; 33)

ramsayi 186

reidi 79, 80

riparis macarthuri 187

roperi 7, 8, 36, 102, 106, 108-110,

g42,0818, Lie. ot20; 121, 123.

124-128, 191 (64, 65, 66)

YOSSti 2

samarensis 106, 107, 115, 117, 192

sandoshami 118

saperoi 31-33, 192

scortechinii 133

separatus 7, 30, 105-110, 113, 121,

128, 130-132, 190 (67, 68, 69)

Setaria 50, 65, 70, 75, 89, 114, 128

Similissimus 106, 107, 192

sinensis 6, 7, 10, 37-40, 42-45, 47-

o2, 04, 596-60, 62-66, 68, 69, 71,

13,. (4, 10, Ti tOO(4, 82 10,.44.

12)

sinensis var. peditaeniatus 71

Sineroides 192

sintoni 176-178, 184

sintonoides 6-8, 20, 27, 28, 107,

145, 147, 177-181, 183-185, 191

(94, 95, 96)

Harrison and Scanlon: Subgenus Anopheles in Thailand 307

species A near umbrosus 118

species B near umbrosus 124

Spiculosa 167

splendidus 186

squamifemur 135

stantoni 162, 163

stephensi 187

Stethomyia 1, 24, 150, 157, 159,

162, 163, 167, 173

stigmaticus Group 113, 144, 148

stricklandi 6-8, 145-149, 156, 166,

£70, P7L-173, 176, 291 (7.30.

90)

subpictus 188

sundiacus 187

Tectona 4

tessellatus 187

tigertti 7, 20, 24, 145-149, 152, 155,

156, 160, 162, 164-167, 169,

175, 181, 191 (79, 85, 86)

tonkinensis 178 |

traguli 96, 114, 118; 122):128. 12

Tragulus 114

treacheri 162, 163

typicus var. pseudopicta 50

umbrosus 7, 14, 20, 105-107, 109, 110,

112-122, 124, 126, 129, 130, 191

(52, 53, 54)

umbrosus Group 15, 16, 20, 23, 25,

27-33, 40, 80, 96, 105, 107, 108,

ota, 198, 122,123, 13t.. 175, 188

uniformis 93

Uranotaenia 102, 124, 167, 181

vagus 2, 187

vanus 6, 79, 80, 83, 86, 99, 103, 192

vanus Subgroup 33, 79, 99, 102

variegatus 118

var. ajajasanensis 133, 140, 142

var. pallidus 102, 103

var. Snijdersi 128, 131

var. williamsoni 66, 68

venhuisi 66, 68

vivax 92, 97

wellingtonianus 135, 192

whartoni 7, 8, 106, 108-110,

113, 114, 119-121, 122-124,

126, 127, 19% (61, 62,63)

Wuchereria 75, 88, 93, 96, 114,

122, 124, 128, ‘t32

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Contributions

of the

American Entomological Institute

Volume 12, Number 2, 1976

MEDICAL ENTOMOLOGY STUDIES - III.

A REVISION OF THE SUBGENUS CULEX IN THE

ORIENTAL REGION (DIPTERA: CULICIDAE).

Sunthorn Sirivanakarn

CONTENTS

CMA ea) ee RN Pe Rees Mee ogee aS Lea lie 1S rat Sole Toca eae Sri Yee Welt! ra Jie TA eee ans Seer tiie Mea See QUE eit Sone Maier Taree, |

ee OG AT A eo ae gle te a we al ee Swe Se

Subgenus Culex

Keys to Groups

ORT Oe ONE Oe Or nen Oey MOUNT. NOY, iB, Oat Ol! OR) Or ON OLE eh E Le OL e NN@in) Oui Ore “Sen wel | Sell vaes hae

Ue ae eR Gen kt ere gly ee i pone ig

eS CO ee a a CR le a ee ee

eee ie es gla See Ce Me an ke ge ee gs yg SR aS

Ce ee a eee ee eae, Dae

Be a anc) te Ok Nae Dies BP ests cum eels 3) aeew Wrus oe ones ve

Ba oy COIN, ae ee eee a a a Re ge ee ee a eS

a Ce Se Gs a ee ea ewe ek eae ee ee

Fe ts as es Ge wie et ee Oe ble Wa ee

ele) MO Gg egg eg eg oy pea “os ae Sb

eR a y galig le de Ceci me 8

i 0 Oe hs 6 es a ee pe a ee eee a ee

Os Cerca @ Wie OOld. Gb be we eee ee ee eee

OO ee a ee iy ie abo ce Sah we Ae} ele we 8

er a a ae hw be ew we

De ee A tie ic eure oe Re ack oy RAs % Sw

ee Vi ee gw ek en ek pe el ee ae el

es OUI CH Ore iC re Ce ee eee ee ek we ew 8

IE SOR 500 CURRIES GER i aac her eer cece ag eel Sanne ea

Pe ee Gy al oie We se) le eR a ee we a wc

DA ee ee ae ye ee a Sa a a ee

D2 See es SO OUIOSS (gee lk EA ee ee ew 8

Mh OH eee Meee kg 4 ea ek week we RR we eo

BO ge ee gia Bk ke Qe ws ee ee

Be cs ee Gia ka OM ie bk ae Seb ee eee

TO, Ces eT eo maldy 66 ak as ee tao, ey ee es

ee et fey a gl ie Lk Tete le we iN ah 8 ae oe we wee

De ie i a he ae a Rtas BU le ee ee

Be ig Ree al) an aw ke lace e eae woe

ee Sg ce ek wel pe ese es wee bo LR ne Se eb 8 acts

Be em Ae he a ichia sis Geen pe a cues ko te ie Phew eareen Be wh 8

Cle are? ea 2 a ee ee ke ke ee ete es

Be OT SG eg a i aa ate Were eo a ek a 8k

Visi emer OuUy, Ait fA ye ee a CE We. 2

le eh aire wie’, Gtiahceln ahi yen ngs doh Melay Bio oy»

Ca ee eee CN ee ee ee Took eee kk eae ns

CO er es AGRE POE ae a ie a a Wa ks GW ee kee ee

GO ee aay a RR as Sima i Ce RC IL a ee ve

OR oer Oe eae Ge ee ee Sk mw la hs wea

Be COTS IN a eA le alia tee A ee keke 8 ew ees

BO. VOC RIOT VNC We WEI ee Sv 0.63 LE ATT Fk a ee aes

Beh Me Re can Ue ites re uate aU KES ip Ty iy) alah bce

ee Fe Ae POM ais Nas og Pea ig ig pak wt a wotgr calm URS alee we Mlb

OS I a Ae a: Ae he ie og Me a gt See at acide gla 6 hp

VRIES SSG BONEN, Gs 0:1. ENO GS Ra Minas Gentes ert ar Vso ha ge a me

CONTENTS

MELIRETUCHS, ROOTS OT Pe De nee eee A es gt Pca: ae Shy Bee ie aa opp ae

33.

se I ae as es ths cg igs ih ae lsay ae bas ek as pen eee cea ee

« JLCRSEE TOURETTE GE eso he a

WATE aA eos eee get kn a NE eT es

POG PE le as ath ea a Me EN hn ete ia ee ee fag

WENO S ACTA, FG ERs Eh A ka a Re

QLENACHSIS- AIMS Th, GUA ae BPP EO tas te pei a ow he

mimulus Edwards

muryvelli Lien

ORE! Oy FO Ol SO Oy er Or: Oi 2 6 a ea OR Ce ee Re Ee Oe Se eae. — kere

v DYODUH GUUS. COM OS Bi aie re Ge ae eee) abit Se alge eee

OVECIIAT TS: AGATA beid, Ca te ee a a SS Lah ee ae ee

ACKNOWLEDGMENTS 0 55 Seely Sea eG a pe oe a aaa G

LITERATURE CVE? eye o< Bee jose) ow ee eae eh te ce eee te

FIGURES

6. ese ese er ey LE re ee a Oy a OP ey ae et: Per © ees ase en eer roe. el OF BS 22 ST Cs et Oe ee Lee hee) eee 3

ill

‘ reg iy oyve :

“3

* ae

poe

Epa

Soe

MEDICAL ENTOMOLOGY STUDIES - III.

A REVISION OF THE SUBGENUS CULEX

IN THE ORIENTAL REGION

(DIPTERA: CULICIDAE)1.

Sunthorn Sirivanakarn2

ABST RACT

This revision of the genus Culex, subgenus Culex of the Oriental region |

is based largely on the study of material from Southeast Asia and adjacent

areas. More than 28, 000 specimens [9, 053 “, 13,075 9, 6,087 whole larvae

(L)| have been examined. These included 6,105 adults with correlated pupal

and/or larval skins (2,071 p, 4,034 1p) associated through individual rearings.

Forty-two species are recognized and of these, 5 are new; 37 are revali-

dated and redescribed. The 5 new species are: luzonensis from the Philip-

pines, selangorensis from Peninsular Malaysia, longicornis from Thailand,

kinabaluensis from Sabah, Malaysia and philippinensis from the Philippines.

The descriptions of 37 species are based on all stages (female, male, pupa

and fourth instar larva) and those of the other 5 species: longicornis, epides-

mus, diengensis, propinquus and orientalis are based only on the adults and/or

associated pupae and larvae. Keys to all available stages and illustrations of

every species are provided. The taxonomic treatment of the subgenus includes

a revised morphological description, followed by a review of the distribution,

taxonomic discussion, internal classification, bionomics and medical impor-

tance and the descriptions of individual species follow essentially the same

format. The various groups, subgroups and complexes are briefly defined

and characterized.

New synonyms proposed here are: neéolitoralis Bram (= alis Theobald);

annulus Theobald (= vishnui Theobald); neovishnui Lien (= pseudovishnut

Colless); fuscifurcatus Edwards and kangi Lien (= jacksoni Edwards); neo-

mimulus Lien (= mimulus Edwards) and afridii Qutubuddin (= infula Theobald).

Two previous nominal forms are resurrected from synonymy: edwardsi

Barraud from barraudi Edwards and infula Theobald from bitaeniorhynchus

Giles.

IThis work was supported by Research Contracts No. DA-49-193-MD-2672

and DAMD-17-74-C-4086 from the U. S. Army Medical Research and Develop-

ment Command, Office of the Surgeon General, Washington, D. C.

2 Medical Entomology Project, Smithsonian Institution, Washington, D. C.

20560.

2 Contrib. Amer. Ent. Inst., vol. 12, no. 2,196

INTRODUCTION

The subgenus Culex is dominant, diverse and widespread in the Oriental

region. It contains several species which are locally abundant as serious pests

of man and domestic animals and as vectors of periodic bancroftian filariasis,

Japanese encephalitis virus and other arboviruses. In Southeast Asia and other

adjacent tropical or subtropical areas of the Oriental region, this subgenus is

taxonomically the best known of all Culex subgenera as it has been subject to

several local studies | Baisas 1938, 1974 (Philippine Islands); Barraud 1934

(Indian peninsula); Bonne-Wepster 1954 (Indonesia); Borel (Indochina); Bram

1967 (Thailand) and Delfinado 1966 (Philippine Islands) | and numerous restricted

studies on the taxonomy, bionomics, genetics, ecology and disease relation-

ships. Among the previous taxonomic studies, the revision of the subgenus by

Barraud (1934) is the most comprehensive; it has a broad coverage of species

occurring not only in India, Burma and Sri Lanka, but also those from South-

east Asia and other adjacent areas to the west in the Middle East, Mediterran-

ean and Ethiopian areas and to the north and northeast in the Palearctic region.

Although most of the previous subgeneric revisions have contributed sub-

stantially to a better knowledge of several common Oriental species, prelim-

inary studies of much material from several countries of Southeast Asia and

adjacent areas, including India and Sri Lanka by Sirivanakarn (1970, 1973)

indicate that numerous taxonomic problems still exist, particularly with respect

to the identity and the specific status of several nominal forms. Some of these

problems are fairly obvious, especially in the case of several closely similar

species in the Vishnui, Mimeticus and Bitaentorhynchus Subgroups. A critical

study of almost all the original and subsequent descriptions in the literature,

in conjunction with a detailed comparative study of the types, numerous topo-

typic and other specimens also revealed that a number of species have been

inadequately described or only superficially known. In fact, many of the dis-

tinct forms can still be easily confused and can not be readily separated on the

basis of the previous keys and descriptions. There is apparently little or no

information in the previous treatment of almost every species on the range of

morphological variation which is essential in determining the specific status of

closely similar forms and in interpreting the affinity and diversity of the various

taxa.

This study is an attempt to revise the subgenus Culex on a broader regional

basis, particularly covering Southeast Asia and adjacent areas. A number of

species which are known only from India, Japan and Korea are also included

for comparative purposes and to provide a more comprehensive revision. In

this revision, 42 species of Culex (Culex) are recognized and of these, 95 are

new and 37 are revalidated and redescribed. In treating the previous nominal

species, the following taxonomic changes have been made: (1) neéolitoralis

Bram is synonymized with alis Theobald, (2) the current synonymy of annulus

Theobald with vishnui Theobald by Reuben (1969) is followed, (3) neovishnui

Lien is synonymized with pseudovishnui Colless, (4) fuscifurcatus Edwards

and kangi Lien are synonymized with jacksoni, (5) neomimulus Lien is syn-

onymized with mimulus Edwards, (6) infula Theobald is resurrected from the

previous synonymy with bitaeniorhynchus Giles, (7) afridii Qutubuddin is

synonymized with znfula Theobald and (8) edwardsi Barraud is removed from

the previous synonymy of barvvaudi Edwards by Bram (1967a). The 5 additional

new species recognized are: luzonensis and philippinensis from the Philippines,

_selagorensis and kinabaluensis from Malaysia and longicornis from Thailand.

Sirivanakarn: Subgenus Culex in the Oriental Region 3

MATERIAL AND METHODS

A considerable portion of the Culex (Culex) material for this study was

accumulated at the Southeast Asia Mosquito Project (now the Medical Ento-

mology Project or MEP), Department of Entomology, Smithsonian Institution

and the entomological collections at that institution. Most of this material

came from collections made by the SEATO Medical Research Laboratory,

U. S. Army Medical Component, in Thailand; the Malaysia Mosquito Project

directed by S. Ramalingam, University of Malaya in Malaysia and Singapore;

the U. S. Army Medical units in South Vietnam, Japan, Korea, the Ryukyu

Islands and the Philippines. Numerous additional specimens, largely field

caught adults, have been obtained from the recent collections in Sri Lanka by

the ''Biosystematic Studies of the Insects of Ceylon" project directed by

Karl V. Krombein; in Java, Indonesia by S. Ramalingam and from the loans

made to the project by the British Museum (Natural History); Field Museum of

Natural History; California Academy of Sciences, San Francisco; Academy of

Natural Sciences, Philadelphia; Bernice P. Bishop Museum, Honolulu; Depart-

ment of Entomology, University of Utah and Instituut Voor Tropische Hygiene,

Amsterdam. A small portion of the specimens of a few to several species

have been received as gifts or as requested from many persons in several

countries, including India, Burma, Bangladesh, Pakistan, the Republic of

South Africa, Italy, Taiwan, Hong Kong, the Philippines, Cambodia, Malaysia,

Indonesia, Japan and Thailand.

I have re-examined most of the existing types and topotypic specimens

of the previous nominal species or forms of Culex (Culex) originally des-

cribed and subsequently reported from the Oriental region except for the

type of annulirostris Skuse from Australia. The majority of the types of the

Oriental species are at the British Museum (Natural History). Included among

the type material are several adult specimens identified by Edwards, Barraud

and others from various localities. Several of these specimens have also been

re-examined and confirmed for inclusion in distribution data. In addition, I

have also examined for comparative purposes, the adults, with or without

associated immature stages of several extralimital species from the Palearctic,

Ethiopian, Australasian, Micronesia and the South Pacific in the reference col-

lections of the U. S. National Museum and the British Museum.

The description and the terminology of the adults used here are essentially

similar to Bram (1967a), Sirivanakarn (1972) and other Culex papers published

by the latter author. The pupal and larval terminology and the wing venation

system of the adults follow Belkin (1962). To simplify the description of certain

features of the male genitalia, the setae of the subapical lobe are designated as

a, b, c, etc., as in Belkin (1962). The lateral plate of the phallosome complex

is simply described using self-explanatory descriptive terms such as apical,

tergal, sternal, mesal, lateral, apical tergal, apical sternal, basal, lateral

basal, etc. as seen from the lateral and tergal views. As for the paraproct

of the proctiger, I have introduced a new term ''subbasal process" for the

small peglike structure situated above the basal sternal process. This pro-

cess articulates with the lateral basal process of the outer division of the

lateral aedeagal plate and is particularly well developed in the Sitzens Group.

The completely labeled illustrations of the various parts of the male genitalia

to which these terms apply are in Figures 2, 4, 9, 21, 24, 32, 36 and 38. These

should be consulted and checked against the descriptions or key for clarification

of the exact position and location of the various parts and processes which have

4 Contrib. Amer, ) Ent, Inst. vole: 12) no. 2, 1976

been frequently used in the diagnosis of practically all species.

In the section on type-data, the status of the type-specimens, holotype or

lectotype and stages or sexes and all associated specimens or slides of all

nominal forms (if clearly indicated in the original or subsequent designations)

are here included and specified. An asterisk (*) indicates that I have personally

examined the type. Unless the names of the countries have not been changed,

the type-localities as stated in the original descriptions are exactly followed.

Country names which have been changed and are currently used are enclosed

in the brackets following the former names noted by the original authors. All

other available data, including breeding sites, elevations, dates of collections

and names of collectors as labeled or mentioned in the original descriptions are

fully given. The abbreviations of the names of institutions in which the types

are deposited follow the listing of Stone, Knight and Starcke (1959).

In listing the specimens under the distribution of species, only the names

of the countries, provinces and/or localities and the number of specimens

examined are given. All other distribution records from the literature are

placed at the end of the list and only those from the Oriental region are dis-

cussed in the taxonomic discussion as to their validity and reliability. In

several instances, I have also taken into account the pattern of distribution,

sympatry and allopatry in attempting to define the spatial relationship between

closely similar species and to speculate on the probable origin of the various

taxa. The data on the bionomics and the medical importance are briefly sum-

marized.

TAXONOMIC TREATMENT

SUBGENUS CULEX LINNAEUS

Culex Linnaeus, 1758: 602. Type-species: Culex pipiens Linnaeus 1758,

Europe. Selection of Latreille 1810: 442.

Lastoconops Theobald, 1903: 235. Type-species: Lasioconops poicilipes

Theobald 1903, Bonny, Nigeria.

Heptaphlebomyia Theobald, 1903: 336. Type-species: Heptaphlebomyia

simplex Theobald 1903, Salisbury, Southern Rhodesia.

Pseudoheptaphlebomyia Ventrillon, 1905: 427. Type-species: Pseudohepta -

phlebomyia madagascariensis Ventrillon 1905, Tananarive, Madagascar.

Leucomyia Theobald, 1907: 372. Type-species: Leucomyia gelidus Theobald

1901, Selangor, Malaya.

Oculeomyia Theobald, 1907: 515. Type-species: Oculeomyia sarawaki

Theobald 1907, Sarawak, Borneo. !

Theobaldiomyia Brunetti, 1912: 462. Type-species: Culex gelidus Theobald

1901.

For complete synonymy, see Stone, Knight and Starcke 1959.

FEMALE. Medium to large sized species, somtimes small; wing length

usually exceeding 3.0 mm. Head. Decumbent scales of vertex narrow,

crescent-shaped, occupying an extensive area in center and along dorsal eye

margin; erect scales numerous, coarse, color varied; broad appressed scales

usually pale, forming a distinct lateral patch at sides of eyes. Eyes contigu-

ous above antennal pedicels; frontal narrow scale tuft well developed; frontal

and dorsolateral ocular bristles strong and dark; subocular bristles weak,

dark or pale. Palpus 4-segmented, 0.20-0.25 of proboscis length, usually

Sirivanakarn: Subgenus Culex in the Oriental Region D

entirely dark scaled, with or without sprinkling of some pale scales; apex of

segment 4 dark or tipped with pale scales. Proboscis usually as long as or

slightly longer than forefemur, sometimes slightly shorter; labium with or

without distinct median pale ring; labial basal bristlelike setae dark and short,

about 0.5 of palpal length. Antenna slightly longer than proboscis; pedicel with

a small conspicuous patch of semierect scales and setae on inner dorsal sur-

face; flagellomeres 1-13 subequal, with numerous short setae; flagellar whorls

2-13 poorly developed, each with 5-6 moderately long hairlike setae; flagello-

mere 1 with or without a few scales. Cibarial Armature. Well developed;

cibarial bar broadly concave, with or without slight median projection; cibarial

teeth 20-30, closely spaced in middle, more widely spaced towards lateral

flanges, length and size of teeth varied; cibarial dome very well developed,

imbricate, very similar in all species. Thorax. Mesonotum with a conspicu-

ous double row of acrostichal bristles from anterior promontory to about

level of wing base; dorsocentral, fossal antealar and supraalar bristles well

developed; mesonotal scales narrow, crescent-shaped, color varied, moder-

ately to very dense on scutal disc, less dense on prescutellar space and

towards posterior lateral caudal margin; scutellar lobes with narrow scales,

forming distinct patch in middle; bristles on scutellar lobes strong and dark;

paratergite bare. Anterior pronotum @pn) with several narrow scales and

strong setae; posterior pronotum (pn) usually with several narrow scales and

sometimes also short setae on upper surface, posterior margin with a row of

4-6 strong bristles, sometimes also with an adjacent row of 5-7 weaker

‘bristles; prosternum (pst) usually entirely bare, sometimes with a small

patch of pale scales and short setae. Pleural scale patches present, usually

1 on propleuron (Pp/), 2 on sternopleuron (stp) and 1 on mesepimeron (mep);

postspiracular (sp) and prealar (va) with or without scales; variable number

of scales usually present among upper mesepimeral bristles; 1-2 lower mese-

pimeral bristles present or absent. Legs. Color of scales on anterior sur-

face of femora varied, usually entirely dark on fore- and midfemora, some-

times with speckling of pale scales or pale stripes; tarsi with or without apical

and basal pale bands; pretarsal claws of all legs small, equal and simple;

pulvilli always very well developed. Wing. All veins usually dense and entirely

dark scaled, sometimes with speckling of pale scales or with pale scales aggre-

gating into distinct pale spots. Abdomen. Terga II-VII usually with basal pale

bands and basolateral pale spots, sometimes with apical pale bands, apico-

lateral pale patches or entirely dark; sterna largely pale scaled or with banding

pattern as on terga. Genitalia. Very similar in most species; segment IX

largely retracted into segment VIII, tergum IX a moderately broad band, with

lateral row of strong setae on each side of median bare area; cerci short,

broad, more or less rounded apically, with numerous strong and weak setae;

postgenital plate more or less triangular, caudal margin broadly rounded,

truncate or slightly emarginate at middle, with 2 lateral rows of setae in apical

_ half and numerous minute spicules; posterior cowl a very narrow band with

numerous minute spicules; sigma broad, largely membranous, with numer-

ous minute spicules; insula with a dense tuft of 8-10 strong setae; vaginal

sclerite or atrial plate U- or V-shaped, usually very well sclerotized.

MALE. Generally similar to female, except for smaller size, less

numerous scales on wing veins. Head. Palpus 5-segmented, usually ex-

ceeding proboscis by at least full length of segment 5; segment 3 with or

without median pale band dorsally, ventral surface with row of short hair-

like or scalelike setae from base to apex, apical 0.2-0.5 with ventral

lateral rows or tuft of strong bristles; segments 4 and 5 upturned, usually

6 Contrib. Amers> Ent dnst.j-volt2) no. 2) 1976

strongly plumose, with several strong bristles on lateral and mesal sur-

faces, dorsal surface with or without basal pale band, ventral surface usu-

ally with a line of pale scales; apex of segment 5 dark or pale scaled. Pro-

boscis as in female, median ventral tuft of long hairs present or absent,

if present, length and number of hairs varied. Flagellar whorls of antenna

strongly long plumose; accessory minor whorls and modified scale tufts ab-

sent. Thorax. Propleural bristles usually more numerous than in female.

Legs. Pretarsal claws of fore- and midlegs enlarged, with subbasal denti-

cles, outer claw larger than inner claw; claw of hindleg small and simple.

Abdomen. Tergal banding patterns, if present usually better developed than

in female; sterna with numerous long setae.

MALE GENITALIA. Segment LX. Tergum narrow, tergal lobes poorly

developed, broadly rounded, with 1 or 2 rows of moderately strong setae;

number of setae varied; sternum large, more or less rectangular, articula-

ting with tergum by a narrow sclerotized band, scales or setae on sternum

absent. Basimere. Conical, lateral tergal margin convex, tergomesal mar-

ein concave; strong bristles restricted to lateral tergal surface, weaker

bristles or setae numerous, evenly spread over tergal area; scales absent.

Subapical lobe. Usually short, projecting mesad; proximal and distal divi-

sions clearly divided or close together; proximal division with 3 strong rod-

like setae @-c); distal division with a lateral group of 1 leaflet (@) and 1

seta (i) and a mesal group of 3-4 flattened bladelike and fine hairlike setae

(d-f). Distimere. More or less sickle-shaped, gently curved in middle,

distal portion tapered and slightly recurved towards apex; 1 dorsal and 1

ventral seta present beyond midpoint; claw subapical, short, distally flat-

tened and apically blunt. Phallosome, Aedeagus usually complex, some-

times simple; distal portion of lateral plate usually with distinct inner

(most sternal) and outer (most tergal) divisions; teeth and processes on

each division variously developed; basal sternal portion (or lip of gonopore)

very well developed, laterally compressed, represented by a broad, simple

and rounded lobe which is well detected in lateral view; upper tergal bridge

narrow, bow-shaped, placed below midpoint of lateral plate; lower tergal

bridge broad, basal or subbasal in position. Proctiger. Apical crown usu-

ally.large with numerous fine and coarse spicules; basal sternal process

of paraproct varied from very short, rudimentary to very long and thick,

curved sternad, sometimes completely absent; a short subbasal sternal

process slightly distad of basal present or absent; cercal sclerite well

sclerotized; cercal setae present, number varied; basal lateral sclerotiza-

tion a narrow, oval sclerotized plate, more or less continuous with basal

portions of paraproct, rest entirely membranous.

PUPA. Size varied. Integument of cephalothorax and abdomen usually

pale, yellowish or cream-colored, sometimes dark brown or with striking

color pattern. All setae developed, branching and length varied. Trumpet.

Tubular; at least 0.5 mm in total length; index 5-15; meatus rather thick,

more or less cylindrical; pinna lightly to strongly oblique, lightly to strongly

widened or flared or about same diameter as meatus, without slit extending to

meatus on ventral surface. Cephalothorax. All setae shorter than trumpet;

setae 1 and 3-C subequal in length, 2-5 branched; 2-C shorter, 3-6 branched;

4-C shorter than or as long as 5-C, 2-4 branched; 5-C 2-6 branched; 6-C

small, dorsad or laterad of 7-C, usually 2-4 branched; 7-C double, 2-3 times

as long as 6-C; 8-C 2-6 branched; 9-C 2-4 branched. Metanotum. Seta 10-C

multiple or at least triple; 11-C double; 12-C 2-5 branched. Abdomen. Seta

-1-I large, multiple and strongly dendritic; 1-II short, single or multiple;

Sirivanakarn: Subgenus Culex in the Oriental Region 7

1-III-VII 0.5 of the length of segments following, number of branches varied;

2-III- VI single, minute, always mesad of 1-IJI-VI, 2-VII usually laterad of

1-VII, sometimes variable in position; 3-I-III usually double, sometimes

single; 5-II 3-8 branched; 5-III 4-14 branched; 5-IV 2-8 branched, slightly

shorter or as long as segment V; 5-V-VI usually double and longer than seg-

ments following, sometimes shorter and more branched; 6-III, IV much weaker

than 6-I, Il, 1-6 branched; 6-V, VI usually longer than 6-IJI,IV, 1-7 branched;

7-I, II 1-3 branched; 9-I single or double; 9-II- VI minute, indistinct; 9-VII 2-5

branched, lightly dendritic, shorter or as long as 9-VIII; 9-VII 5-10 branched,

usually strongly dendritic; segment IX with a single minute seta laterad of genital

lobe; male genital lobe (basimere) at most 0.3 of paddle length. Paddle. Broad,

oval; pigmentation of inner and outer parts usually pale, sometimes varied;

external margin very lightly spiculate in basal 0.5, distally smooth or without

fringe of spicules, apical margin rounded, or sometimes slightly emarginated

at apex of midrib; midrib well developed, usually lightly pigmented, some-

times dark; apical setae 1, 2-P small, short, usually single; 2-P sometimes

absent or not developed.

LARVA. Head. Broader than long; maximal width (distance between

lateral margins of ocular bulge) usually nearly approximates the width of

prothorax, sometimes greatly reduced (Bitaeniorhynchus Subgroup); labrum

developed as distinct transverse bar, sometimes poorly developed or not

apparent; ocular bulge usually prominent; seta 1-C filiform or spiniform, some-

times strongly flattened and apically blunt; minute setae 16, 17-C laterad of

collar absent. Antenna shorter or as long as dorsal length of head; several

spicules present, more or less restricted to basal 0.5 of shaft; seta 1-A large,

fan-shaped, multibranched and strongly pectinate; 2, 3-A single, bristlelike,

usually subapical, sometimes apical. Mouthbrush elements filamentous. Men-

tal plate usually with distinct lateral teeth on each side of median tooth, some-

times teeth not differentiated. Thorax. Spiculation usually absent; setae 1-3-P

subequally strong and long, usually single; 4-P shorter than 1-3-P, usually

double, sometimes single or more branched; 5, 6-P single, strong, 7-P double

or triple; 8-P double; 14-P single. Abdomen. Spiculation usually absent;

chaetotaxy varied; setae 6-I, II and 7-I strongly developed; 6-I, II usually

triple, sometimes 4 branched; 7-I single or double; 6-III- VI moderately to

strongly developed; 1-III-VI weakly to strongly developed. Comb scales varied

in number and size, seta 2-VIII usually single, sometimes double or more

branched. Saddle usually completely sclerotized, rarely incomplete; posterior

margin lightly spiculate; seta 1-X usually weakly developed; 2, 3-X strong;

4-X (ventral brush) strong, usually with 6 pairs of setae, all inserted within

erid; anal gills shorter or longer than saddle. Siphon. Shape, length and

color varied; index 3-14; acus very well developed; pecten teeth varied from a

few to several, denticulation varied; siphonal tufts variously developed, 3-7

pairs in number; seta 2-S single, length and thickness varied; median caudal

filament of spiracular apparatus usually present, sometimes absent.

DISTRIBUTION. The subgenus Culex is worldwide in distribution. In

Southeast Asia and in most parts of the Oriental region, the subgenus has been

recorded from every country wherever a mosquito survey is made. A number

of the Oriental species have also been reported from the Ethiopian, Palearctic

(Mediterranean, Middle East, China, Siberia, Soviet Union, Japan and Korea)

and the Australasian regions (including Micronesia and South Pacific). Of the 2

major groups (Pipiens and Sitiens), the Sitiens Group is exclusively Old World

with an extensive range of distribution largely restricted to the tropics of the

Ethiopian, Oriental and Australasian regions while the Pipiens Group is

8 Contrib. : Amer. Ents Inst. vok 92, no.::2; 1976

worldwide.

The distribution of the species of the subgenus Culex in Southeast Asia and

other adjacent areas within the Oriental region are shown ina table (Appendix

A). Without any claim to completeness and accuracy, I have attempted to in-

clude most of the extralimital records from the literature for every species

treated in this revision. For a more detailed account of the distribution of the

groups, subgroups, and species, see the discussion in each category.

DISCUSSION. The majority of Oriental Culex (Culex) species are very well

characterized in the adults by the presence of conspicuous ornamentation on the

palpus, proboscis, thorax, legs, wing and abdomen. With the exception of

closely similar species in certain subgroups or complexes, adults of many

species can be readily recognized or differentiated by external characters.

The male genitalia are important at the subgeneric, species group, subgroup

and specific levels using characters of the phallosome, proctiger and subapical

lobe. In the females, the cibarial armature may be of some value in the diagno-

sis of certain species with extremely similar external characters. The female

genitalia are of little or no practical value in differentiating species or sub-

genera. In the immature stages, both the pupae and larvae show a number of

good subgeneric and specific characters that are very useful in the diagnosis

of several species with extremely similar adults and male genitalia. For an

accurate identification or determination of species in certain subgroups or

complexes, all stages must be examined.

All members of Culex (Culex) in the Oriental region are separated from

members of other Culex subgenera (including Barraudius, Acalleomyia, Culici-

omyia, Eumelanomyia, Lophoceraomyia and Lutzia) by the following combina-

tion of characters: adults by (1) presence of a distinct double row of acrosti-

chal bristles, (2) presence of at least 3 distinct scale patches on the pleuron

and in the majority of species by, (3) presence of a pale ring in the middle of

proboscis, (4) presence of conspicuous basal and apical bands on tarsomeres

1-4 of the legs and (5) presence of basal and/or apical bands on abdominal

terga II-VII; females by presence of 20-30 teeth in the cibarial armature;

male sexual characters by (1) palpus longer than proboscis, (2) absence of

lateral ventral row of lanceolate scales on palpal segment 3 and (3) absence of

modified scale tufts on antennal flagellum; male genitalia by (1) complex phal-

losome, (2) large proctiger crown, and in most species by, (3) presence of a

long basal sternal process in the paraproct of the proctiger; pupa by (1) pinna

of trumpet without slit extending into meatus, (2) seta 10-C usually multiple,

rarely double or triple and (3) seta 9-VIII well removed from caudolateral

angle of segment VIII; larva by (1) absence of setae 16 and 17-C, (2) setae

1-3-P usually subequally long and strong, (3) seta 14-P single, (4) siphon with

3-7 pairs of hair tufts, (5) ventral brush (4-X) of saddle usually with 6 pairs

of setae, and in the majority of species by, (6) dark, flat, spiniform seta 1-C

and (7) usually single seta 2-VIII.

Prior to this study, 45-47 species and 2-3 subspecies of Culex (Culex) have

been considered as valid taxa in the Oriental and the eastern Palearctic regions

(Stone, Knight and Starcke 1959; Stone 1961, 1963, 1967 and 1970). In this

study, I recognize 42 species in the subgenus. Almost all of the previous

nominal species from Southeast Asia and adjacent areas in the Oriental and

the eastern Palearctic regions [ including India, Bangladesh (East Pakistan),

Burma, Sri Lanka (Ceylon) as revised by Barraud (1934) and from Japan and

Korea by LaCasse and Yamaguti (1950)|, are treated. The only exceptions are

3 species from mainland China: huangae Meng, permixtus Hsieh and Liao and

- cheni Ho, which are omitted because I have not been able to examine their types

Sirivanakarn: Subgenus Culex in the Oriental Region 9

or topotypic specimens. In addition, 2 other species are not included: nilgiri-

us Barraud from India and boninensis Bohart from the Bonin Islands, Micro-

nesia, both of which do not appear to belong to the subgenus Culex. The true

affinity of these 2 species can not be determined at this time, however, it

appears most probable that nilgivicus belongs to the subgenus Eumelanomyia

and that bDoninensis probably represents a distinct subgenus judging by the

male genitalia and adult characters.

Internal classification. The Oriental members of the subgenus Culex fall

into 2 distinct groups, Pipiens and Sitiens as defined by Edwards (1932: 200-12)

on the basis of conspicuous coloration in the external characters of the adults.

Except for minor modifications in the arrangements of species within each

group and further subdivisions (e.g. into subgroups) by subsequent authors

dealing with local studies, (Baisas 1938; Colless 1957a; Mattingly and Rageau

1958; Belkin 1962 and Bram 1967a), Edwards' scheme of recognizing 2 major

groups in the subgenus is still applicable to the Oriental Culex (Culex) fauna.

As evident from the comparative study of the male genitalia and the immature

stages, there are a number of borderline cases among some Oriental species

which do not appear to fit perfectly into either group. Both groups are highly

complex, and although they should, perhaps be considered separate subgenera,

many more subgenera would then have to be recognized. This would evidently

create confusion rather than clarification and should not be pursued until all

groups and subgroups in the subgenus are thoroughly re-examined on a global

basis. In this study, I am attempting only to recognize subgroups within the 2

major groups by adopting a scheme of internal classification similar to that

developed by Edwards (1932) and other subsequent authors. Each of the sub-

eroups is represented by one or more species, some of which are further

segregated into complexes whenever it becomes appropriate. Asa result,

several changes in the realignment of species have been made. :

Among the 42 Oriental species treated here, 6 belong to the Pzpzens Group

and 36 to the Sitiens Group. The Pipiens Group comprises 4 subgroups: (1)

Pipiens Subgroup with quinquefasciatus which is widespread throughout tropical

parts of the world, (2) Trifilatus Subgroup with vagans and hutchinsoni, both

are exclusively Oriental, (3) Theileri Subgroup with theileri, very well known

in the Ethiopian region and (4) Univittatus Subgroup with wnivittatus, a domin-

ant Ethiopian form and fuscocephala which is exclusively Oriental. The Sztzens

Group is more dominant and extremely diverse, comprising 6 subgroups: (1)

Gelidus Subgroup represented only by gelidus, which is exclusively Oriental,

(2) Bitaeniorhynchus Subgroup with bitaeniorhynchus, which I am restricting

to the type form; infula, luzonensis, selangorensis, pseudosinensis, longicornis,

epidesmus, cornutus, Sinensis, geminus and kinabaluensis, all are exclusively

Oriental except bitaeniorhynchus which is apparently widespread, extending

from the Oriental to the Ethiopian, northeastern Palearctic (Japan and Korea)

and certain parts of the Australasian region; izfula and sinensis which are

widespread in the Oriental region, (3) Sifzens Subgroup with sitiens, a dominant

coastal form, widespread in the tropical parts of the Oriental, Australasian

(including: South Pacific, Micronesia) and in the eastern parts of the Ethiopian

region; alis, a litoral brackish water species which is known from Christmas

Island and Southeast Asia; whitmorei which is widely spread throughout the

Orient with extension into Japan, Korea and New Guinea; annulirosiris in the

Philippines and widespread in Micronesia, South Pacific and Australasian

region, (4) Vishnui Subgroup with vishnut, pseudovishnui and tritaeniorhynchus;

all are widespread in Southeast Asia; alienus and perplexus in Malaysia and

Thailand; philippinensis and incognitus known only in the Philippines and whitei

10 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

from India, Bangladesh, Thailand, Peninsular Malaysia, the Philippines and

Ceram, Lombok, Indonesia, (5) Barraudi Subgroup with barraudi from India

and Thailand and edwardsi from India, New Guinea and northern Australia and

(6) Mimeticus Subgroup with mimeticus which is a rare Oriental form but wide-

spread westward into the Mediterranean and Middle East; fasyz in the Philip-

pines; jacksoni in India, Sri Lanka, Hong Kong, Taiwan and probably also in

China; fsengi in Taiwan; diengensis in Java, Indonesia; mimuloides in India;

mimulus, widespread in Southeast Asia, India, Sri Lanka, New Guinea and

northern Australia; murrelli widespread in Thailand, Peninsular Malaysia and

probably also in India; propinquus in Singapore and orientalis from Japan and

Korea, and perhaps also in China, Siberia and USSR.

BIONOMICS. All Oriental species of Culex (Culex) whose immature stages

are known breed in ground pools and occasionally in artificial containers.

Several species are restricted to more or less permanent types of ground water,

such as ponds, swamps, bogs or obstructed streams which contain algal mats

(e.g. Bitaeniorhynchus and Mimeticus Subgroups). The medically important

species in the Vishnui Subgroup frequently breed in open country such as rice

fields or cleared land developed for other agricultural purposes. Two species,

sitiens and alis, which are litoral, show a high tolerance for brackish or even

salt water in rock pools, puddles or artificial containers along sea beaches.

Culex quinquefasciatus, which is the dominant domestic species in large or

small cities, shows a very broad range of tolerance for polluted ground waters

in drains, septic tanks and in the multitude of artificial containers. The rest

of the species are fresh water breeders.

Females of most species of Culex (Culex) require a blood meal to complete

the gonotrophic cycle. They are primarily crepuscular or nocturnal biters and

feed on the blood of mammals, birds and possibly other vertebrates. Host feed-

ing patterns for the majority of species are not known from all areas within the

distribution range except for a number of common species which have been in-

vestigated in Malaya by Colless (1959) and in India by Reuben (1971la, 1971b)

and Christopher and Reuben (1971). Hosts include cow and pig for trilaenior-

hynchus, gelidus and fuscocephala; pig and bird for sitiens, pseudovishnui and

vishnut; bird and man for quinquefasciatus and bird for mimulus and bitaenior -

hynchus. Although man is the frequent host for quinquefasciatus, Colless (1959)

reported that other species, including ¢vitaeniorhynchus, gelidus and fuscoce -

phala which naturally feed on cow and pig, will also attack man in the absence

of those hosts.

MEDICAL IMPORTANCE. The subgenus Culex is the most important of

all Culex subgenera from the public health standpoint. In Southeast Asia, the

subgenus contains several species which are abundant and closely associated

with man and his domestic animals through the blood feeding habits of the fe-

males. Many species are now known to play an important role as actual or

potential vectors of pathogens of human diseases, including bancroftian filari-

asis and Japanese encephalitis virus (JE). Culex quinquejasciatus, (or fatigans

of some authors), which is the notorious pest of man in cities and towns, is

considered to be the principal vector of urban periodic bancroftian filariasis in

several countries of Southeast Asia. Several epidemiological studies have

shown conclusively that tvzlaeniorhynchus is the primary vector of Japanese

encephalitis in Japan, Okinawa, Taiwan, Sarawak, Borneo and India. In

studies dealing with virus isolation and experimental transmission from the

Oriental region other species including gelidus, fuscocephala, vishnui and

pseudovishnui have also been considered to be probable or potential vectors of

Japanese encephalitis.

3(2).

Sirivanakarn: Subgenus Culex in the Oriental Region 11

KEYS TO GROUPS AND SPECIES

ADULTS

One or 2 lower mesepimeral bristles present; proboscis without

distinct median pale ring; tarsomeres without basal or apical

pale: bands: (PIPING GROUP wocss bos aoe Se BY eee Se 2

Lower mesepimeral bristles absent; proboscis with distinct

median pale ring; tarsomeres with basal and apical pale

bands: (SITIBNS GROUP ee ae ee eae ee 7

PIPIENS GROUP

Anterior surface of midfemur without median longitudinal pale

SUPIPO. 6 deta ee hee eG Ee Oe 2 ee 3

Anterior surface of midfemur with median longitudinal pale

SIVIDG. 2s ee ee ee Or ee ee ra ae a

Abdominal terga without basal transverse pale bands; pleuron

with striking pattern of dark and pale stripes. .... fuscocephala

Abdominal terga with basal transverse pale bands, pleuron without

striking pattern of dark and pale Stripes... Jy fees ce. 4

Mesonotal integument reddish brown; pleural integument with dis-

tinct pattern of dark and pale stripes on sternopleuron and

IMOSEPLMCTON RS Skiers Be Pee ee ee hutchinsoni

Mesonotal integument yellowish or pale brown; pleural integument

without pattern of dark and pale stripes as above.

quinque fasciatus

Postspiracular area and base of prealar knob with distinct scale

DALCUCS 4 au. Gk ay ee ae se ie 6

Postspiracular area and base of prealar knob without scale

DALCNOR a alia eer GEG Giles Chek te Oe eee vagans

Large species (wing length about 5.0 mm. ); basal pale bands of

abdominal terga very broad and strongly produced at middle.

i theileri

Smaller species (wing length about 3.5 mm.); basal pale bands of

abdominal terga narrow and even in width........ univitta tus

SITIENS GROUP

Wing with pattern of pale spots or streaks on at least 2 areas of

costa ond-one area’ of other vemisws ih) ei wee erie a3

Gelidus, Bitaeniorhynchus, Sitiens, Vishnui and Barraudi

Subgroups

Abdominal terga II-VIII largely yellowish or golden. ... epidesmus

Abdominal terga II- VIII with dark and pale bands or entirely dark. . 9

9(8).

10(9).

11(9).

12(11).

13(12).

14(13).

15(13).

16(12).

17(16).

Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

Abdominal terga II-VI entirely dark or without basal, apical pale

DANdS OF -ADICOLALET Sl DALE ATEN Ss hehe ee ne tele ec aoe ari 10

Abdominal terga II-VI with basal or basal and apical pale bands

OF QOLCOLA LE Fal Ee OAC AR se ui erp are We luatap its & Yak Ure Capita 11

Dark areas on abdominal terga with light sprinkling of some pale

scales; size large, wing length exceeding 4.0 mm. . selangorensis

Dark areas on abdominal terga without any scattered pale scales;

size small or medium, wing length not exceeding 4.0 mm.

perplexus, whitei and incognitus, in part

Abdominal terga II-VI with basal pale bands only. ........ 18

Abdominal terga II-VI with apical bands and/or apicolateral

Dale patches and Oasal HALS DANS kia iiwwies woubliariw jade oe ee piies be 12

RS OT OAL GI i ieee de dew ope scat ae bshwrs Bw ahi 13

Wing witha mixture of dark iand pale scalesi.i wiv ae es 16

Anterior surfaces of fore- and midfemora largely dark scaled or

with very light sprinkling of few pale scales; prosternum with-

BPS LY Cs he ae OL Oe ar Ng ah ie Ab aval tebe AR totes Sah ag worse ow 14

Anterior surfaces of fore- and midfemora moderately to strongly

speckled with numerous pale scales; prosternum with a small

By eg cas) gaa ge hay Ws NAS a Mek ce sw 8 LS

Apical bands on abdominal terga III- VI strongly widened towards

sides, forming a large apicolateral pale patch; anterior sur-

face of fore- and midfemora usually without speckling of

pale scales on/doreal suriace. Sk ee pseudosinensis

Apical bands on abdominal terga III-VI slightly widened towards

sides, forming a small apicolateral pale patch; anterior surface

of fore- and midfemora with a very light speckling of some

DALE. SCAIES.ON GOTSAL SUTTACE. woe oi ial dean! or eh ee VO geminus

Clypeus with a distinct patch of pale scales on lateral surface;

apical pale bands on abdominal terga IV-VI broader than

oo SOE SSS UG GN PURE Sg rae ees OO MISC Sea ie ee te Cae mean cornutus

Clypeus without any scales on lateral surface; apical pale bands

on abdominal terga IV-VI as broad as basal bands. .... SINENS1S

Abdominal terga II-VII with broad apical bands; legs (femora, tibia)

and wing heavily speckled with pale scales; scales on wing veins

Ry, ,.Rq And Re OrOadiOV ates, Ciiciaat ia: Ta elie ko bitaeniorhynchus

Abdominal terga II-IV usually largely dark or with narrow apical

bands, apicolateral yellowish patches and median basal pale

spots or complete basal pale bands; tergites V-VII with narrow

or broad apical and/or basal bands; legs and wing lightly to

moderately speckled with pale scales; scales on wing veins

Rey ns and Wie Narrow: CLavate ye OS a ae ee 17

Basal pale bands on abdominal terga IV-VI usually broader or as

broad as apical bands (Separation tenuous, also use male

RTT. GALLUS) OR TUN: DELANO NEMA sin Want th EG luzonensis

To(11).

19(18).

22(21).

23(21).

24(23).

25(24).

Sirivanakarn: Subgenus Culex in the Oriental Region 13

Basal pale bands on abdominal terga IV-VI usually narrower than

apical bands. . infula, longicornis (inseparable, use male genitalia)

Erect scales in center of vertex of head whitish; anterior 0.7 of

mesonotum densely covered with pure white scales. ..... 19

Erect scales in center of vertex of head pale beige, dingy white or

dark; anterior 0.7 of mesonotum covered with beige, yellow,

DOIMCN, OTC SOAIMSUe A ae PES a ge 20

Anterior surfaces of fore- and midfemora without speckling of pale

scales; scales on prescutellar space, lateral mesonotal areas

behind wing base and on scutellum entirely dark brown. . gelidus

Anterior surfaces of fore- and midfemora with extensive speckling of

pale scales; scales on prescutellar space, lateral mesonotal

areas behind wing base and on scutellum predominantly pale

WT op: ls nese ube ia mbna en sa cea ae sar hie, Sod age aR eG whitmorei

Cell Ro of wing (in both sexes) at most as long as vein R2+3;

Male: palpal segments 4 and 5 weakly plumose........ alis

Cell Ro of wing 1. 5-2.0 times as long as vein R913; Male; palpal

secments,.4.qnd:5 Strongly PlUMOSes iis. iar dae Meyda ps |

Midfemur with longitudinal pale stripe on anterior surface; post-

Spiracular area with a small patch of semierect scales on

LOWS! BNO RIOD SUPT AC Gishh 05 hacia tar Nb TA ee dn cy 22

Midfemur entirely dark or with sprinkling of pale scales not

forming definite stripe as above; postspiracular area without

any, SCeles On LOWEY ADLETION, SUMIAC Oy. 4:6 calm wise bs Ue cir 6, 23

Female: longitudinal pale stripe on anterior surface of midfemur

broken up into a row of white spots at middle; Male: proboscis

without ventral tuft of long hairs at base of median pale ring.

barraudi

Female: longitudinal pale stripe on anterior surface of midfemur

complete; Male: proboscis with distinct ventral tuft of long

hairs at PAGES OP TROMIAN OAL UM iii Oi fel gle OO edwardsi

Anterior surface of fore- and midfemora with speckling of several

pale scales at least on apical dorsal surface. ........ 24

Anterior surface of fore- and midfemora entirely dark. ..... 27

Wing scales entirely dark; mesonotal integument chestnut brown or

blackish; speckling of pale scales on femora contrasting sharply

With eK SCALA SPO s vy piial ay gy eniel ipa aN Ami MR glia Nal a bas 29

Wing with a few to several scattered pale scales; mesonotal

integument light brown; speckling of pale scales on femora

not contrasting sharply with dark scaled areas. .. vishnui, in part

Hindfemur largely pale; anterior surface of foretibia without a row

of yellow spots among dorsal DVIStI@S. «466k ek ie a ee 26

Hindfemur with an even mixture of light and dark scales or

largely dark; anterior surface of foretibia usually with a distinct

row of yellow spots among dorsal bristles (also use keys to

26(25).

27(23).

28 (277).

29(27).

30(29).

31(30).

32(31).

ContribisAmerwEnt.+nst; 2ivoli12, noie2.-19'76

malegenitd lia and larvae). ay ya ela. be Sree ali annulirostris

Hindfemur with discrete pale stripe from base to 0.75 of length;

speckling of pale scales on fore- and midfemora largely

Tesirieted lOoap1eal Usb. ww bedi wies Ie laa weet whitei, in part

Hindfemur freckled or with several dark scales scattering among

pale ones; speckling of pale scales on fore- and midfemora

covering almost entire length (separation tenuous, also use

(pe Peni VA as hie pte oe ees Le ale sitiens

Erect scales on vertex and scales on anterior 0.7 of mesonotum

LoeGGr Gee DrOWUiShe Wi Oe ee ee Gi oes. 28

Erect scales on vertex pale beige or yellow in center; dark

posterolaterally; scales on anterior 0.7 of mesonotum

largely pale beige, yellow or golden with mottling of some

Te ee is te eee cee us ee RO ees 20

Female: Mesonotal scales very fine and entirely dark brown;

all cibarial teeth very fine, elongate and distally filamentous;

Male: ventral surface of palpal segment 3 with a row of fine |

Nate Bete. os 6 Se GBI Pia Sle eas & 8 tritaentorhynchus

Female: Mesonotal scales rather coarse and partially pale,

particularly on supraalar area; cibarial teeth short, thickened

and apically blunt or abruptly pointed; Male: ventral surface

of palpal segment 3 with a row of scalelike setae (separation

from other species in couplet 30(29) not always reliable,

SO ar Vaasa «ied WA a a Ea a vishnui, in part

Large species, wing length usually exceeding 4.0 mm. ; forefemur

with distinct subapical yellow spot; Male: palpal segment 3

with dense fine hairlike setae on ventral surface. . . kinabaluensis

Small-medium sized species, wing length less than 0.4 mm.,

forefemur without subapical yellow spot; Male: palpal

segment 3 with a row of scalelike setae on ventral surface. ... 30

Females (inseparable, use keys to larva and pupa).

vishnut, in part, pseudovishnui,

perplexus, alienus, philippinensis,

incognitus and whitet

Males (separation not always reliable, use keys to larva and pupa).

Proboscis without ventral tuft of hairs at base of median pale ring.

pseudovishnui

Proboscis with ventral tuft of some long hairs at base of median

PAE WO ek ON ee ee OS ROE eis ail gis oe

Palpal segment 3 completely dark. . perplexus and incognitus, in part

Palpal segment 3 with median pale band.

vishnui, in part, alienus,

philippinensis, incognitus, in part

| and whitei

33(7).

34(33).

35 (34).

36(35).

37(36).

38(37).

39(34).

40(39).

Sirivanakarn: Subgenus Culex in the Oriental Region £0

Mimeticus Subgroup

Wing with basal costal pale spot in addition to 2, 3 other costal

spots; male palpal segments 4 and 5 weakly plumose. . . orientalis

Wing without basal costal pale spot; male palpal segments 4 and 5

strony oluimos@ ewan) 4 orn eae de ei eee. 34

First or median costal spot of wing involving veins Sc and/or C

only; basal pale bands of abdominal terga broad or at least

0229 Of SECIRENE TONGE gs Meech lw cen oho Benes 4s Se

First or median costal spot of wing involving veins C, Sc, Rj,

Rs and sometimes also M and Cu, basal pale bands of abdominal

terga narrow, usually less than 0.29 of segment length. .... 39

Furcation of vein M proximad of furcation of vein R9i3. . mimuloides

Furecation of vein Mat same lewekoituncation.of gig.) 225. 2. . 36

Veins R4+5 and Cu, largely pale scaled from near base almost to

best emia geese oe eo Ale oa ie eee, diengensis

Veins R4+5 entirely dark scaled or with pale streak restricted to

the: middle;"Cuy- with short:median pale streak... 244, . - 37

Third or apical costal pale spot of wing involving apical portion

of vein Rj only; median costal pale spot largely involving vein

SO ieciguen ee eS RR th EO) Peat OO ee Eat) 5 tsengi

Third or apical costal pale spot of wing involving apical portions

of veins C, Ry and sometimes also Ro; median costal spot.

involving veins: Cain Se ouewe 4 Gen ia Se ee. 38

Mesonotal scales predominantly brownish; midtibia with longitu-

dinal pale stripe on anterior surface; male proboscis with

distinct: ventral tuft of several long hairsy:... a .:6!k. jacksoni

Mesonotal scales predominantly pale; midtibia without longitu-

dinal pale stripe on anterior surface; male proboscis with or

without indistinct ventral tuft of a few long hairs.

mimeticus and fasyi (inseparable, use male genitalia)

Wing veins Cu, Cus and 1A each with a row of pale scale spots

Cie 90) Huis oe ore) oo ae we re le We ak propinquus

Wing veins Cu, Cug and 1A each with continuous pale scale line or

SURGAKs 26:16 4. co EE a ee is aay 40

First median costal spot usually involving veins C, Sc, Ry and Rg,

sometimes extending to Mand Cu........ mimulus, in part

First median costal spot involving veins C, Sc and Rj only.

murrelli and mimulus, in part, (inseparable, use larva)

3(2).

4(3).

0(3).

6(5).

Contribs. Ameér>: Ent, ‘Inét. )\ volc82) nos 2.1976

MALE GENITALIA!

Inner division of lateral plate of phallosome without spiculation on

apical sternal portion; subapical lobe more or less clearly

divided into proximal and distal divisions (PIPIENS GROUP)... 2

Inner division of lateral plate of phallosome covered with

numerous minute spicules and/or denticles on apical sternal

portion; subapical lobe usually not clearly divided into proxi-

mal. and distal divisions (SITIENS GROUP). 2 Loc io... 7

PIPIENS GROUP

Basal sternal process of proctiger absent or very poorly

Shy oo | wR Ag SUE RBM GR ay a eres a a quinque fasciatus

Basal sternal process of proctiger present, in form of a long

and (hich wer mcurved StOrmed.: S005 SADR O UL BAS 3

Setae d-f of subapical lobe situated on a distinct mesal lobe on

sternal surface; apical 0.25 of distimere strongly curved

and distinctly annulate or serrated dorsally. ......... 4

Setae d-f of subapical lobe not situated on a distinct mesal lobe;

apical 0.25 of distimere gently curved and not annulate or

SOW ONCE tees ek TN I ee Se ke Dee a OD

Inner division of phallosome simple, represented by a broad

heart-shaped leaf projecting laterad; tergal arm of outer

division tapered into: a sharp points is 58 ek Be. univittatus

Inner division of phallosome composed of several strong

toothlike processes laterally; tergal arm of outer division

inifonmly thick: with trumeate apex! $402 oP yen a LS fuscocephala

Inner division of phallosome straight or slightly divergent

laterad, with 2,3 teeth on apical lateral margin; lateral

spine of outer division weak and short (Fig. 9). ...... thetleri

Inner division of phallosome sharply bent or angled, without

teeth on apical lateral margin; lateral spine of outer

division sirone and Jone (Ties. Se ee ee ee 6

Outer division of phallosome with a broad rounded lateral basal

process; inner division sharply angled; tergal arm of outer

division distally tapered ina sharp point........ hutchins oni

Outer division of phallosome with a slender lateral basal

process; inner division strongly curved, but not angled;

tergal arm of outer division swollen subapically...... va Zans

propinquus unknown.

Th);

8(7).

10(9).

11(10).

12(11).

13(12).

14(13).

Sirivanakarn: Subgenus Culex in the Oriental Region 17

SITIENS GROUP

Inner division of lateral plate of phallosome represented by a large,

simple forcepslike process; outer division not developed.

SinenS1S

Inner division of lateral plate of phallosome complex, outer

divasioniwell developede: 6:5 ae Sar Lay Lal ee 8

Inner division of phallosome represented by a short, slender,

spinose fingerlike process; outer division simple, broad

and pounded On OUP SGA iy wine oe eo OR eee. gelidus

Inner division of phallosome represented by a broad, elongate,

finely spiculose or toothed process; outer division with

distinct teeth or spines or represented by a strong toothlike

OY leatiake Spi y er ee a ee) eg 9

Inner division of phallosome with 2-6 strong fingerlike processes

or teeth forming a prominent crown on apical tergal and lateral

SUPEAOeN ui LANGE POLIS RA Tes G8 Re eee 18

Inner division of phallosome without a prominent apical tergal

crown of fingerlike processes or teeth as above, but with a

lobe bearing 2-4 foliform processes or elongate toothlike

Spines below apical margin or apical tergal angle. ....... 10

Basal sternal process of proctiger well developed, about 0.1 mm.

or more in length; outer division of phallosome in form of a broad

lobe, “bearing a elrong INeSAl Spine.’ wre my S EPRI tow. 11

Basal sternal process of proctiger poorly developed or absent,

if present, its length varying between 0.01-0.08 mm; outer

division of phallosome in form of a broad simple leaflike spine

or Sina ie teothlike Spine 47¢ ayers See es UE a eae 12

Leaf g of distal division of subapical lobe broad, foliate; inner

division of phallosome with a tergal lobe bearing 1-2 narrow,

CACAO Os eg Ut ee ety eS BEETS avis 2 epidesmus

Leaf g of distal division of subapical lobe narrow, lanceolate;

inner division of phallosome with a tergal lobe bearing 2-3

DROAC TOR OTe WOES. OUR eae ee oe Rae cornutus

Basal sternal process of proctiger absent; outer division of

phallosome represented by a strong toothlike spine. . annulirostris

Basal sternal process of proctiger usually present, its length

varying from 0.01-0.08 mm; outer division of phallosome

represented by a broad leaflike spine... 60... 2 2 eu. 13

Apical sternal portion of inner division of phallosome (in lateral

view) very long and narrow, somewhat fingerlike. . . . longicornis

Apical sternal portion of inner division of phallosome quadrate

OT TA TORM OF Bi BEOUE eA ee ieee ge I Ae a ra 14

Apical sternal angle of inner division produced into a long, curved

beak sternad, apical tergal angle blunt or slightly produced. .. 15

15(14).

16(13).

17(16).

18(9).

19(18).

20(18).

21(20).

2921).

23(20).

Contrib. Amer, Ents Insts, vol. 12, noi 2) 1976

Apical sternal and tergal angles of inner division more or less

equally produced inte a enort beaks oe oe ce 16

Apical margin of sternal portion of inner division smoothly

curved; apical tergal angle usually blunt or rounded and

Without pity denies: 2 FP eG a bitaeniorhynchus

Apical margin of sternal portion of inner division depressed

proximally; apical tergal angle moderately or strongly

produced and usually with some dénticles, . 2.06.03 So... infula

Foliform processes of inner division situated close to apical

(Orel Abele. 6 ee ea eo selangorensis

Foliform processes of inner division situated far below

peal Vel eal ate, Se ae PP ere eee ila

Apical sternal portion of inner division very stout and smoothly

curved on apical margin, apical sternal angle (or process)

longer than apical tergal QUEUE Cie SA Gy WO eeu luzonensis

Apical sternal portion of inner division slender and lightly

depressed on apical margin, apical sternal angle (or process)

about as lone as apieal tereal anele: . 24)... 3: pseudosinensis

Basimere enlarged, sacklike; distimere modified. ......... 19

Basimere normal, slender, conical, distimere normal or

EV eat Sie le -Gaped. 6 6 ee ee ee eS, 20

Subapical lobe with several strong specialized setae in distal

Civisioe cistimere Gs im Fie. 80.° Gii8. os cn Sed 2G orientalis

Subapical lobe with 6 narrow and short bladelike and rodlike

setae in distal division; distimere as in Fig. 69. .... diengensis

Inner division of phallosome (in lateral view) broadly expanded

CO LG a eiecince Joe Slerneds ok Pee OP eed... 21

Inner division of phallosome not expanded sternad to form a

CUStINCG! (Oke Oe oes Bee se Doe Se. wo

Basal sternal process of proctiger absent; sternal expanded

portion of inner division of phallosome with distinct denticles

Ce eo hoes A ee ee ee em alis

Basal sternal process of proctiger present; sternal expanded

portion of inner division of phallosome without distinct

Genticlee Say ee eS Peer er meh Sh 22

Basal sternal process of proctiger slender and short; outer division

of phallosome represented by a large, simple, broad leaflike

SOI Me er ee ee ey whitmoret

Basal sternal process of proctiger strong, thick and long; outer

division of phallosome represented by a rounded lobe bearing

2-3 strong heavy spines on mesal and sternal surfaces. . . sitiens

Outer division of phallosome with 6-8 strong and dark teeth and

several weak denticles on lateral and mesal surfaces. ... whitei

24(23).

25(24).

26(24).

2126).

20(21).

—29(25).

30(29).

31(30).

pail).

Jolee)s

Sirivanakarn: Subgenus Culex in the Oriental Region 19

Outer division of phallosome with only 1-3 strong teeth on lateral

Gnd MISSA BUTI ACOs fe ea ee oo ag wee EO 6 ee 24

Outer division of phallosome without sternal spine; apical tergal

crown of inner division with 2-3 short fingerlike processes.

kinabaluensis

Outer division of phallosome with distinct sternal spine; apical

tergal crown of inner division with 4-6 short or long fingerlike

DYOCOSSOG i eee ee ee Ne ee ee ee ee PAS,

Sternal spine of outer division of phallosome long, prominent and

gently divergent JAverAde: ajc 4 ja) KOR ST ee BO ee es 26

Sternal spine of outer division of phallosome short, indistinct

or strong, distinct and* strongly curved basad.. .-.) 6 i... 42. 29

Basal sternal process of proctiger slender, pale, at most 0.05 mm

in leéneth or completely absentee 6 ee ae OR Ce 6 4 at

Basal sternal process of proctiger thick, dark, usually 0.08-0.10

ry er Ome ee RS ae Bae ee Ore ik a ee edwardsi

Basal sternal process of proctiger not developed or absent. . jacksoni

Basal sternal process of proctiger present, at least 0.03 mm

DY OTM i hes aa use eh gor be bd sac te TE ROR ge Oe ee eee. 28

Basal sternal process of proctiger about 0.05 mm; apical

fingerlike processes of inner division of phallosome slender and

POUIVODT TOME, 6 85 oa Me te te te Melts Sa ta tee hae he a eee ae eee mimeticus

Basal sternal process of proctiger shorter, about 0.03 mm in

length; apical fingerlike processes of inner division of

phallosome thicker and shorter....... a, SAP" GaSnt and TSene7

Basal sternal process of proctiger absent or poorly developed.

philippinensis

Basal sternal process of proctiger present or well developed. ... 30

Basal sternal process of proctiger slender and pale; leaf g of

subapical lobe narrow, Dladelike. sy" Ceti a mimuloides

Basal sternal process of proctiger thick and dark; leaf g of

Subapied) jehe. Wroad, Tole. oe , ee eee Pea a ol

Fingerlike processes of apical tergal crown of inner division of

phallosome largely porrect or with apices directed dorsad. .. 32

Fingerlike processes of apical tergal crown of inner division

of phallosome all divergent laterad..... ate ig ag Wes aon barraudi

Setae d-f of subapical lobe all flattened, bladelike, subequal in

Setae d-f of subapical lobe more or less differentiated in length

TI WEG oo 5 VE Pea My Mal ah petal a ae a igs gig aOR emer at gr 34

Subapical claw of distimere short, distally broad and bilobed,

length about: O.008 it el ks Foe a ee a ee ees mimulus

Subapical claw of distimere long, slender and simple, length

slot a CMe ob es): aang UE erie aeremr ay mien tt Mem Tal Cheesy ulin ayss murrelli

34(32).

35(34).

36(35).

37(36).

38(36).

Contribs: Amer. Ents. Inst.) «mole da, now 251976

Apical tergal crown of inner division of phallosome with strong

and long fingerlike processes; apical sternal portion of inner

division irregular, denticulated and not distally expanded to

form a prominent triangular lobe (tergal view). ....... a5

Apical tergal crown of inner division of phallosome with weak and

short fingerlike processes; apical sternal portion of inner

division simple and distally expanded to form a prominent

trianeular. lobe: (tergal. Views: ino iis v seckk eh we wie tritaeniorhynchus

Distal division of subapical lobe with 4 setae in group d-f; outer

division of phallosome with a large, rounded and prominent

Taleral Dawa Process, 4. ni 2k ea) Pei bi. atte Gury es Le Rai geminus

Distal division of subapical lobe usually with 3 setae in group

d-f; outer division of phallosome with a small, slender or

elongate, kKnoblike lateral basal process. ... . 64 206 ee ae 36

Lateral basal process of outer division of phallosome short and

PCUAIIVE DEO.) ek? ude bbiwirenl. be Bw he vadle Bie tere Lea ce cs at

Lateral basal process of outer division of phallosome elongate

and. More: elenmer than. above siuis Saw we Gah isi ee 38

Fingerlike processes of inner division of phallosome very strong

and relatively long.

vishnui and incognitus (inseparable, use larva and pupa)

Fingerlike processes of inner division of phallosome moderately

strong and relatively short (separation tenuous, use larva and

a ee io ys heat days ei So ge pasty ag aaa Ai ong perplexus

Sternal surface of subapical lobe with numerous minute setae

POLI 8 Gast tCl. DALCH en tvewhotes Poke teres Uh aac pseudovishnui

Sternal surface of subapical lobe with fewer minute setae forming

SOIR Pe HOLE CLL ES essere ice ahs Binet eae Se hele RET Ee ee ee alienus

Seta 1-II multiple, strongly. dendritic or with more than 10

branches; trumpet usually pale yellowish; paddle entirely

pale to almost transparent. (PIPIENS GROUP). ......... 2

Seta 1-II single, forked or with less than 10 branches, if more,

then trumpet dark brownish or paddle with conspicuous

darkened. arenas. (SI7JE NS GROUPB duo drew. sein Savers a Ra reerelln ge fi

PIPIENS GROUP

Seta 8-C double or triple; setae 6-III-V double ortriple...... 3

Seta 8-C 4-6 branched; setae 6-III-V 4-6 branched. ......... 5)

Seta 1-C double; seta 6-VI double or triple; small species, abdomen

ee VERT R bee) Sued ape gh DU AALS ee SSin! comllege sah fag aN ge wy et gk han ah ly ag hutchinsoni

Seta 1-C 3-4 branched; seta 6-VI 4,5 branched; medium species,

abdomen about, 2.) aaaiiet «leva tw Wis oo Wale Ceyial AO a deb wR 4

| opidesmus, propinquus and longicornis unknown; orientalis, not included.

Sirivanakarn: Subgenus Culex in the Oriental Region 2A.

4(3). Pinna of trumpet distinctly widened and strongly oblique; seta

Gay MEU CL i Oe ee a a ee es quinque fasciatus

Pinna of trumpet not distinctly widened and lightly oblique; seta

DOE Ws UV Pee a ae ceok. ie Sie wow ha ton oe enna ge ee va gans

5(4). Small species, abdomen 2.5-2.7 mm; seta 0-IV 4-6 branched. . . 6

Large species, abdomen 3.5 mm or more; seta 5-IV triple. . ¢theileri

6(5)}ocr Getae GOV IVI dowblee el ae aoe RL. bo ule gil HON Ga fuscocephala

Setae 54, VE UsUaty CrIpre Pi wee ed SEs be tires univittatus

SITIENS GROUP

7(1). Trumpet asymmetrically funnel-shaped; pinna strongly widened

or PlAred TOW ds ae ee 4 ow ee ele a alte 8

Trumpet cylindrical-shaped; pinna same diameter as meatus or

slightly Widened towards APC%. eos GG we ie ic gi ole 17

8(7). Trumpet whitish or lightly darkened on meatus; cephalothorax

and abdomen very pale whitish; seta 1-III 8-11 branched. .... 9

Trumpet yellowish brown to dark brown; cephalothorax and

abdomen strongly yellowish to dark brown or with pattern

of dark alveolar spots, if entirely pale then seta 1-III 2-95

OTRO Het Li SE ee Paes ie as an INE te ae Ma 10

9(8). Seta 8-C 3,4 branched; seta 9-VII 5 branched; 9-VIII 7-9 branched.

sinensis

Seta 8-C double; seta 9-VII double or triple; 9-VIII 5 branched.

cornutus

10(8). Abdominal segments III-VIII entirely pale; paddle entirely pale without

darkened or infuscated areas; seta 1-II] 2-5 branched.

kinabaluensis

Abdominal segments III-VII strongly yellow or dark brown or

with pattern of dark alveolar spots; paddle usually with

darkened or infuscated areas; seta 1-III 8-13 branched. ..... 11

| 11(10). Seta 10-C and setae 3-I-III usually single. ...... bitaeniorhynchus

Seta 10+C and setae S-1-TEdouple. eae a he eh ea 8 5 12

12(11). Abdominal segments pale yellow with pattern of dark alveolar spots, 13

Abdominal segments strongly yellowish or entirely dark brown

without pattern of dark alveolar: spOte. 5 uh ew kg oe WN ew 14

13(12). Inner part of paddle darkened; basal external margin of outer

part. not darkened or miuscates ie. (ae ee te oP 8s selangorensis

Inner part of paddle pale or transparent; basal external margin

of.outer part darkened’ or‘ infulseate.”.))) 6 ee infula, in part

14(12). Seta 1-V 4-8 branched; 4-VII 3,4 branched. ........+4.s46-.: 15

Seta 1-V double or triple; 4-VII usually double. .......... 16

15(14).

16(14).

17(?).

18(17).

19(18).

20(19).

2120),

22(20).

23(22).

24(19).

25(24).

26(25).

27(26).

28(27).

Contrib. Amer, Hat, Inst.,: volei2, nov 2; 1976

Outer and inner parts of paddle entirely pale or transparent.

pseudosinensis

Outer and inner parts of paddle slightly dark tinged. ..... geminus

Cephalothorax and abdomen entirely dark brown or yellowish

with darkened areas or spots on posterior abdominal segments;

paddle usually with infuscate areas; seta 5-IV 3-5 branched.

infula, in part

Cephalothorax and abdomen uniformly yellowish, without darkened

areas or spots on posterior abdominal segments; paddle always

entirely pale or transparent; seta 5-IV usually double. . luzonensis

Bere e sO OR re OF PIMC. on a ee ee a ee ee es 18

meld, GeO anO OleMed Gib iae siwis oe BES on Poe. 31

Seta 1-II 20 branched; paddle with dark spot on apex of midrib.

gelidus

Seta 1-II 1-10 branched; paddle without dark spot on apex of midrib.

petae.6-\V. Vi ueually single or doublew.:: 6.) eos). ecient. . 20

SGrae G- Ve Vis, Drancied,. 6). weikey Se deen 24 ©

Trumpet relatively short, index about 5; apical margin of pinna

RO ee a te a ee eS 21

Trumpet relatively longer, index usually more than 6; apical

Mareim of pinna truncate or lightly emareinate. .. Ss)... . Ze

Pinna of trumpet strongly oblique; setae 1-III, IV 4-6 branched.

| sitiens

Pinna of trumpet weakly oblique; setae 1-III, IV double or triple. . alis

Seta 5-IV usually 4 branched; Peli 450 byanched, .-. 0 ok whitei

Seta 5-IV double or triple; 1-II] single or double. ......... 23

Seta 1-III 2-4 branched; 1-IV 3,4 branched. . .

Seta 1-III 5-8 branched; 1-IV 4-6 branched. .

mimeticus and jasyi

jacksoni and tsengi

Seta 9-IV 7-8 branched; 5-V 3-5 branched. ...:..%.: perplexus

Beta o-1V 2. roche: Sey ONbDIC. . 6k ee et ee ee a yaS)

Pete VU SINGIE i ai. el er GU. eee ia icwe TRS a diengensis

Sera-o- V1 CONDLes sneha mt Me ae oe, ah Ne PIS I 26

peta d-C.double seta: talk double. sie. ie Ge eo. mimuloides

Seta 1-C 3,4 branched; seta 1-II 4-7 branched. .......... 27

Beta.1-Vi triple; 1-VIl double or triplés:.i375). wie anc. 28

Seta 1-Vi 2.9 branched; 1-Vil 4 branched...) . ee a0

Color of cephalothorax and abdomen more or less uniformly

yellowish white; seta 1-III 9,10 branched; 1-IV 7-9 branched.

mimulus, in part

2921)

30(29).

pee aes

S231).

33(32).

34(32).

35(34).

36(35).

Sirivanakarn: Subgenus Culex in the Oriental Region Zo

Color of cephalothorax and abdomen with darkened areas; setae

11H, DY SO rane ie es ee ek ee eee murrelli

Setae 4, 5-VII single; 4-VIII usually single. ...... philippinensis

Setae 4p S2Vil doubles 4] Vil aeuble. neat, a COR ee a 30

Seta SC triple; 5-C weually 4 brancheds.. 6 ar) ne SO alienus

Seta 8-C double;'o<C usually triple 7 oo ee ae vishnut

Setae 5-V, VI 6-8 branched, their length much shorter than segments

fOllowlNe ie eae PO ae ae i |... . Whitmore

Setae 5-V, VI double, their length as long as or longer than

segments following TSEC, 1 AE Se OE) Ra eek Pe ee 32

Setae 6-IV-VI double or triples} .........- ee ay 33

Setae 6-IV- VI 4+T branched: ©. 00 ine ee ere var cau 34

Trumpet pale yellowish and relatively slender and long, index

about 12; seta 1-II 2-4 branched. ........ mimulus, in part

Trumpet brownish and relatively thick and short; index about 8;

seta I=] 4-10 brancheds Us 04 ee Woe. parraudi and edwardsi

‘Trumpet: tong andthin, index 10=1d, 8.14) 4 Leet Bias incognitus

Trumpet shorter and thicker,:index 6-600» ei. le ae ee. |

Seta 9-C 3,4 branched; 1-C 4 branched; medium or large species,

abdomen 3.0=<3: 0) Minis 2 BRO. Geek wie annulirostris

Seta 9-C double; 1-C usually triple; emia species, abdomen less

PHATE O Se aa i Rg a ak eo ORS 36

Trumpet dark brown; seta 1-II iia nicab more than 10 branched.

tritaentovhynchus

Trumpet pale yellowish; seta 1-II less than 10 branched.

pseudovishnui

LARVAE!

Labrum or preclypeus of head capsule present as a distinct

transverse bar separated from frontoclypeus........... 2

Labrum or preclypeus of head capsule not developed as distinct

bar separated iron troniocl pes: a en. mown gers BIS ete a 9

Seta 1-C pale, slender and distally strongly tapered or filamentous. 3

Seta 1-C dark, stout, spiniform or foliform and abruptly pointed

on bint diptoaliy suki hn ww a ee a en ee ee

1 epidesmus, longicornis, diengensis and propinquus unknown; orientalis

not included.

3(2).

8(7).

91).

10(9).

11(9).

1a),

Contribs ‘Amer. Ent, Inst. ; volw32,: noi:2,):1976

PIPIENS GROUP

Setae 5, 6-C double; mental plate with 6,7 lateral teeth on

each side of median tooth; siphon slender more or less

CVlindriesl and, mictally Tapered. 4... sib sc ee ee fuscocephala

Setae 5, 6-C 4-6 branched; mental plate with 10-12 lateral

teeth on each side of median tooth; siphon thick, fusiform

nC. Gistinctiy SwOllen In WAIGGIe. ook wee Geek aie ee 4.

Setae 2, 3-A placed apically or nearly so; siphon short, index

about 3, basal0.5 strongly swollenmei: wos 6s quinque fasciatus

Setae 2, 3-A distinctly subapical; siphon longer, index about

i DESH ae Pet ly SWOLET okie al elle ce ak eee, vagans

Siphon short, fusiform, strongly swollen in middle and with a strong

subapical spine on dorsal surface; setae 1-M and T remarkably ©

SPROne i ee ee. el oe a le ae GAL I hutchinsoni

Siphon usually long, cylindrical and tapered distally, if short

and swollen as above, then without subapical spine on dorsal

surface; setae 1-M and -T short and weak. ........... 6

et a PO eek iy, he eR Oe aE ere AOL ee 7

Sete ieee ay ak a ae eS, For eu Poca a! 16

Comb scales pointed apically and with a strongly differentiated

ieee al th wae a RE Bain > GILES ° thetleri

Comb scales rounded apically and with even fringe of fine

ee a sek Gere ake e ee Oe. EA owe 8

Seta 6-VI single; pecten teeth of siphon with 3 lateral denticles;

I OG WC a eae aa ee ee be ee univittatus

Seta 6-VI double; pecten teeth of siphon with 8-9 lateral

denticles; siphon yellowish to brownish. .......-.+.-e+.s-. 26

SITIENS GROUP

Setae 2, 3-A located half way between apex and base of seta

1-A, seta 4-P short and minute, rather indistinct....... 10

Setae 2, 3-A apical, subapical or closer to apex than to base

of seta 1-A; seta 4-P strong and distinct, 2 ici .ee aie ee 11

Siphon: with > pairs ol subventwal tutte. canal. wiley eed os SINENSIS

Siphon with 4 pairs of subventral tufts. . 2... 2. 2 ee ee ee cornutus

Mental plate with 5-7 large, distinct lateral teeth on each side

Cre Te TCC TOCTN a eas ie wows ea ike Se ae 12

Mental plate without distinct lateral and median teeth; all

teeth minute and not differentiated as above. ........ 13

Setae 1-3-P double; setae 6-I, II 5-7 branched; 7-I double.

kinabaluensis

Setae 1-3-P single; setae 6-I, II triple; 7-I single. ...... geminus

13(11).

14(13).

15(14).

16(6).

17(16).

18(17).

19118),

20(18).

21(20).

22120):

eulae).

Sirivanakarn: Subgenus Culex in the Oriental Region 29

Comb scales small, 6-12 in number, their length 0.05-0.08 mm;

siphon with distinct row of 7-10 pecten teeth.

pseudosinensis and selangorensis

Comb scales large, usually 4,5 in number, their length over

0.1 mm; siphon with or without distinct row of 3-5 pecten teeth. .14

Siphon usually with 4 pairs (or total 8) of subventral tufts.

bitaeniorhynchus

Siphon with 3 pairs (or total 6) of subventral tufts. ........ 15

Antennal shaft practically bare or with a small number of very

weak, spiculés in basal Nalisisiei. Ge Bes ea ar luzonensis

Antennal shaft distinctly barbed with several strong spicules in

basal ett ces fe Oe a ON Le ge I, el a haa infula

Siphon short, fusiform, middle part distinctly swollen, index

mh cn kr) ae Ratna DURA eh th hy CLT SI i oa eG Co aan gelidus

Siphon longer, more or less cylindrical; index at least 5... . 17

Seta, 441P double; of waitiple. eG Rs a eee 18

SOLARA EPs Micra Ree ie Be IG sos SOS oe Ae a gee gl aly 2g

Seta 1-C markedly flattened, or somewhat foliform, with blunt

apex; anal gills short, stubby and rounded, about 0.2 of

SACS: POUT a 5 We sh sink hare epee" Savana Wan be ae tot Aaa ay Pee We 19

Seta 1-C slender or spiniform, with acuminate apex; anal gills

long, tubular, as long as or longer than saddle length. .... 20

Saddle complete; setae 2, 3-A distinctly subapical; setae 4 and 8-P

strond,. 2G) LONG AS da BaP. fogs el del ie Gece rele Pa a al sitiens

Saddle incomplete; 2, 3-A apical or almost so; setae 4 and 8-P

weak; much shorter than boo Dive pid: Poe a i alis

Comb with 4-8 large spiniform scales in single row, seta 4-P

eID LS ae TI Si ah gas EOS EEE a ote eG aia Be eee a1

Comb with 16-20 or more smaller scales in 3,4 rows, seta

Bh TO LAAN OIL ise hiv eS SR gd ERT aa aa ate 22

Siphon strongly tapered and upcurved distally; siphonal tufts very

strong, 2-3 branched each, 4,5 times as long as siphonal

width at point of attachment; 4-P always double. .... whitmoret

‘Siphon lightly or moderately tapered, straight or slightly curved

in apical 0.25; siphonal tufts weaker, 4-6 branched each, as

long as or slightly longer than siphonal width at point of attach-

Ments:4=P COUPLE OF WTO lee oii Meinl aces iain owe wii lay bs pseudovishnui

Seta b-C 466 branched) G-C triples s audi Tei ale guns mimuloides

Seta 5-C usually double or triples 6+Cdoubleiy yh voge we aks 23

Pleural areas of thorax spiculate; proximal pairs of siphonal

tufts closely spaced, all in line on ventral surface. .... vishnui

Pleural areas of thorax not spiculate; proximal pairs of siphonal

tufts more or less widely spaced, inadouble rowsubventrally. .. 24

24(23).

27(26).

28(27).

29(17).

30(29).

31(30).

32(30).

Contribs Amer. Bint. dnst:s-vol. 125 no. 2; - L976

Siphon strongly tapered distally, 2,3 proximal pairs of siphonal

tufts 3-5 times as long as siphonal width at point of attachment,

rect oradually GHOTEEr. 5... cc 00 bw bk ew we i a 20

Siphon more or less uniform in width along entire length or lightly

tapered distally; all siphonal tufts shorter, subequal, shorter

or slightly longer than siphon width at point of attachment.

| perplexus

Comb scales 13-20 in number, median apical spine of scales

strongly differentiated from lateral fringe of spicules. .. alienus

Comb scales 30-40 in number, median apical spine of scales

poorly differentiated from lateral fringe of spicules.

philippinensis

‘Siphon thick and relatively short, index 3-5; siphonal tufts

very strong, 6-12 branched each and closely spaced.

annulivostvis

Siphon slender and relatively longer, index 7-11; siphonal tufts

weak, 2-4 branched each and widely spaced. ......... age

Seta 6-C double; siphon brownish, slender, length usually 1.4-

1.6 mm; ventral valve of spiracular apparatus small, dark

EOWA ee: A ee tritaentorhynchus

Seta 6-C triple; siphon yellowish, thick, length 1.6-2.4 mm;

ventral valve of spiracular apparatus large, blackish. .... 28

Setae 1-III-V single and strong, as long as setae 6-III-V,

SinhGna) tiie rathersimeonspicueuss s,s. she LG is Wiilei

Setae 1-IJI-V 2-4 branched and considerably shorter than

setae 6-III-V; siphonal tufts distinct. . . . barvaudi and edwardsi

Comb with a mixture of large, spinelike and small, normal

ak se ad es ae as le ee incogniltus

Comb wiia Ssiigle type of scale, all small. . oo. 2 ee wee 30

Siphon with some prominent spines on ventral surface in apical

Wales a a a er hee ot

Siphon without any Spines aS ADOVE.. oo. 6,2 se eRe eh oe oa

Setae 6-I, II usually 4,5 branched; siphonal tufts weak, all sub-

equal, shorter or as long as siphonal width at point of

Attac imme mie ee eS ae) Se We jacksoni

Setae 6-I, II triple; 2,3 proximal pairs of siphonal tufts

strongest, 2-3 times as long as siphonal width at point of

attachment, “rest pradually shorter... 40.0 si vw Siete « tsengi

Comb scales with strong median apical spine; 4 proximal pairs of

siphonal tufts very strong, closely spaced, 2,3 times as long

as siphonal width at point of attachment, forming a prominent

POUR: BR RIG ee Coa EP ee mimeticus and fasyi

Comb scales normal or with poorly developed median apical

spines; all siphonal tufts weak, subequal, widely spaced,

shorter or as long as siphonal width at point at attachment,

not forming a prominent row as above. ......4.+4-+-+-e--e-s 33

Sirivanakarn: Subgenus Culex in the Oriental Region 27

33(32). Siphon long and thin; index 9-14; comb scales with pointed apical

TPIS ee se ee ee ae gee a ee ea ke BO ee mimulus

Siphon short and relatively thick; index 6,7; comb scales with

POUNCE Bical 1VINGCS. 6 ae Skee we exer ee ees murrelli

PIPIENS GROUP

FEMALE. As described for the subgenus, with the following distinctive

sroup characters. Head. Proboscis entirely dark scaled, median portion

sometimes pale ventrally but not extended dorsad to form a distinct pale ring.

Cibarial Armature, As described for the subgenus. Thorax. Mesonotal in-

tegument pale brown to dark brown, mesonotal scales pale beige, yellowish,

golden to dark brown, more or less uniform in color on anterior 0.7 of mesono-

tum and on areas laterad of prescutellar space; scales on prescutellar space

and scutellar lobe largely pale whitish. Pleuron with distinct pale scale

patches on pp/, upper corner and posterior border of s¢p and upper part of

mep, sometimes also on psp and base of pra; 1,2 lower mep bristles present.

Legs. Tarsomeres 1-5 of all legs entirely dark, or without distinct apical and

basal pale bands. Wing. All scales narrow and entirely dark except for a

short pale scale line on posterior surface of base of veinC. Abdomen. Terga

II- VII usually with complete basal pale band, sometimes without such band or

completely dark. Genitalia. As described for the subgenus.

MALE. Generally as described for female except for the following

characters. Head. Palpal segment 3 usually entirely dark scaled, sometimes

slightly pale at middle on dorsal surface, ventral surface with 1, 2 rows of

short, fine hairlike setae along the whole length, apical 0. 25-0. 50 usually with

ventrolateral tuft of several strong bristles; segments 4 and 5 strongly plumose,

entirely dark on dorsal surface, ventral surface with scattered pale scales

which may form a distinct line or streak at base. Proboscis usually pale at

middle ventrally, but not forming a complete light ring; median ventral hair

tuft proximad of false joint usually absent, sometimes present.

MALE GENITALIA. In general as described for the subgenus with the

following distinctive group characters. Basimere. Slender, conical and

without special modification. Subapical lobe. Proximal and distal divisions

more or less clearly divided, bases of setae a-c in proximal division not over-

lapped with those of setae d-f in distal division; setae d-f 2-4 in number, some-

times placed on a distinct lobe on sternal surface; leaflet g present, variously

developed. Distimere. Normal, typically sickle-shaped; 1,2 ventral tiny

setae present, dorsal one absent. Phallosome. Inner division of lateral plate

simple, not spiculate, represented by a single, narrow or broad pointed diver-

sent ventral arm, sometimes straight, with 3-4 strong lateral spical spines or

teeth; outer division with a strong and long mesal tergal arm, 1 lateral and 1

sternal spine, latter poorly or well developed, usually divergent laterad; lateral

basal process knoblike, poorly or well developed, apex pointed or rounded.

Proctiger. Apical crown dark and large, composed of numerous coarse and fine

spicules; basal sternal process poorly or well developed.

PUPA. Cephalothorax and abdomen pale whitish or light yellowish with

darkened areas not forming definite pattern. Trumpet. Varied in length;

meatus usually more or less cylindrical, sometimes slightly widened distally;

pinna lightly or strongly oblique. Cephalothorax. Seta 8-C 3-6 branched; 9-C

double or triple. Metanotum. Seta 10-C 4-10 branched; 11-C double. Abdo-

men. Seta 1-II multiple or between 10-20 branched; 1-III 5-10 branched; 1-IV

0-8 branched; 1-V 4-7 branched; 1-VI 4-6 branched; 1-VII 4, 5 branched; 5-IV

28 Contrib. Amer... Pnt. Inst, , wol.12.-no...2, 1906

2-6 branched; 5-V, VI double or triple; 6-III 2-6 branched; 6-IV 2-4 branched;

6-V 2-5 branched; 6-VI 2-5 branched; 9-VII 3, 4 branched; 9-VIII 6-8 branched;

4-VIII double. Paddle. Broad, inner and outer parts pale whitish; midrib pale

or lightly pigmented.

LARVA. Head, Labrum or preclypeus developed as a distinct transverse

bar separated from frontoclypeus; ocular bulge prominent; seta 1-C fine, fili-

form or stout spiniform; 4-C usually long, about 2 times as long as distance

between bases of the pair, single or double; 14-C very small indistinct.

Antenna shorter or as long as head; setae 2, 3-A apical or subapical. Mental

plate with 6-12 lateral teeth on each side of median tooth. Thorax. Spicula-

‘tion absent; setae 1-3-P strong, subequal in length and single; 4-P varied,

2-5 branched; 7-P double or triple. Abdomen. Seta 7-I single or double; 6-1, II

usually triple; comb scales small, numerous, forming a broad oval patch,

usually with rounded apical fringe of fine spicules, sometimes pointed, with a

strongly differentiated median apical spine. Saddle complete; seta 1-K 1-3

branched, 2-X 2-4 branched; 4-X (ventral brush) usually with 6 pairs of setae,

sometimes 7; anal gill slender, usually 1-2 times as long as saddle length.

Siphon. Varied in shape and length; index 3-7; pecten very well developed;

siphonal hair tufts 4-6 pairs in number, length, branches and position varied;

seta 2-S slender and short.

DISCUSSION. The Oriental members of the Pipiens Group all show better

eroup characters in the adults and the male genitalia than in the pupae and

larvae. Adults of both sexes in the Pipiens Group can be readily separated

from the members of the Sitiens Group by: (1) presence of 1 or 2 lower mese-

pimeral bristles; (2) absence of distinct pale ring in the middle of proboscis

and (3) absence of basal or apical pale bands at the joints of tarsomeres 1-4 of

all legs. The males in the Pipiens Group are distinguished by (1) absence of

basal and/or apical pale bands on dorsal surface of palpal segments 4 and 5;

(2) ventral surface of palpal segment 3 with rows of short and fine hairlike

setae and (3) absence of median or submedian ventral tuft of long hairs proxi-

mad of false joint of proboscis (except fuscocephala). The male genitalia in

the Pipiens Group are very similar to those in the Sitiens Group in the basi-

mere, distimere and proctiger, but are strongly differentiated from the latter

in the following features: (1) absence of minute spiculation in the inner division

(ventral arm) of the phallosome, (2) presence of a strong and long tergal arm

on mesal surface of the outer division and (3) subapical lobe more or less

clearly divided into proximal and distal divisions. The pupae of the Pzpiens

Group largely overlap with those in the Sitiens Group, especially in respect to

the chaetotaxy. The only pupal character which appears to be more or less

diagnostic of the Pipiens Group is the branching of abdominal seta 1-II which

is multiple or composed of more than 10 branches as indicated in the key.

The larvae of the Pipiens Group also exhibit considerable overlap with those

in the Sitiens Group. Except for specific and subgroup characters, there is

apparently no single feature or a combination of characters of the larvae in the

Pipiens Group to clearly separate them from those in the Sitiens Group. The

similarity in the immature stages of the Pipiens and Sitzens Groups is an indi-

cation of their close affinity and probable monophyletic origin.

The Pipiens Group contains several species which are largely restricted to

the Ethiopian region but is represented by a small number of species in other

parts of the world. Within the Oriental and eastern Palearctic regions, the

group consists of 8 taxa: pipiens pallens Coquillett 1898, quinquefasciatus Say

1823 (or fatigans Wiedemann 1828 of authors), huangae Meng 1958, hutchinsoni

Barraud 1924, fuscocephala Theobald 1907, theileri Theobald 1903, univitiatus

Sirivanakarn: Subgenus Culex in the Oriental Region ae

Theobald 1901 and vagans Wiedemann 1828. It is also possible that pzpiens

molestus Forskal 1775 may have been introduced into Japan (Y. Wada, personal

communication). The taxonomy and the relationships of the pipiens forms in

Japan have not yet been sufficiently investigated. From the previous reports

in the literature (LaCasse and Yamaguti 1950, Bekku 1956), it appears most

likely that the typical or true pipiens does not occur in Japan. In this study,

only 6 Oriental taxa in the Pipiens Group are treated. Culex pipiens pallens is

excluded because the specimens from Japan, Korea and northern China, which

I have examined are inadequate, consisting only of adults and whole larvae. On

the other hand, this pipiens taxon has never been found to extend into Southeast

Asia and other parts of the Oriental region and is most probably restricted to

the cold temperate climate similar to that of the typical pipiens of northern

Europe in the western Palearctic. Culex huangae Meng is also omitted because

the type and topotypic specimens are not available for study.

Edwards (1932), in an attempt to develop a scheme of interspecific classifi-

cation, placed quinquefasciatus (as fatigans), vagans, theileri and univittatus

in the pipiens series and fuscocephala and hutchinsoni in a group of uncertain

affinity. Subsequently, only a few attempts have been made to classify species

in the pipiens series of Edwards. Mattingly and Rageau (1958) placed vagans

in their trifilatus subgroup. Mattingly et al. (1951) and Mattingly (1957, 1967),

in the discussions on the systematics of pipiens, recognized a pipiens complex

to include quinquefasciatus Say (or fatigans Wiedemann) and australicus Dobrot-

worsky and Drummond as subspecies of pipiens; and molestus Forskal, pallens

Coquillett and comitatus Dyar and Knab as forms of pipiens sensu lato. Since

there has not been a consensus as to the subspecific or specific status of these

forms, it seems more practical here to categorize the pipiens complex as a

subgroup of the Pipiens Group and to include within it certain nominal taxa which

are treated as full species, such as globocoxitus Dobrotworsky 1953 from

Australia.

In this study, I recognize 4 subgroups for the 6 Oriental species treated.

Two of these subgroups, pipiens or pipiens complex of Mattingly (1957, 1967)

and Belkin (1962) and trifilatus of Mattingly and Rageau (1958), are retained.

The subgroups theileri and univittatus are proposed for the first time to include

certain Oriental taxa which I believe represent distinct lineages. These 6 spe-

cies are segregated into the 4 subgroups as follows: (1) Pipiens Subgroup with

quinque fasciatus; (2) Trifilatus Subgroup with vagans and hutchinsoni; (3) Theileri

Subgroup with theileri and (4) Univittatus Subgroup with fuscocephala and univit -

tatus. The only other Oriental species, huangae from China, appears to belong

to the Trifilatus Subgroup on the basis of the description and figure of the male

genitalia by Meng (1958).

PIPIENS SUBGROUP

The Pipiens Subgroup is characterized in the male genitalia by (1) setae in

group d-f of distal division of the subapical lobe 3 in number, placed between

setae a-c of proximal division and leaf g of distal division; (2) inner division of

phallosome represented by a simple, broad or narrow leaflike ventral arm

which is divergent laterally; (3) outer division with a short, inconspicuous

lateral spine and (4) basal sternal process of the proctiger rudimentary or

poorly developed.

The Pipiens Subgroup includes the typical pipiens and other nominal

intraspecific forms (molestus, pallens, comitatus, etc.) from the temperate

region, quinquefasciatus from the tropics and subtropics of the world,

30 Contrib. Amer, Ents fasts, vol, ol2;-no. 2,- 1976

australicus from Australia (Dobrotworsky 1965) and the South Pacific (Belkin

1962) and globocoxitus from Australia (Dobrotworsky 1965). In Southeast Asia

and adjacent areas, the Pipzens Subgroup is represented only by quinquefascia -

tus which I am treating as a full species in spite of the fact that some authors

consider it as subspecies of pipiens. My conclusion on the taxonomic status

of this form is based largely on the constant differences in the male phallosome

and palpus, behavioral and ecological attributes and characteristic distribution.

As pointed out by Mattingly (1967), a more thorough review of the morphology of

the various forms in the pipiens complex based on all stages is still critically

_needed. For an extensive review of the pzpzens complex, see Mattingly et al.

(1951) and Mattingly (1957, 1967).

The members of the Pipiens Subgroup show closer affinity to those in the

Trifilatus Subgroup than to other subgroups. Belkin (1968), in discussing the

New Zealand fauna in comparison with those in Australia and the South Pacific,

pointed out that the Pipiens (complex) and Tvifilatus Subgroup are not really

distinct morphologically and zoogeographically. Although this seems to sug-

gsest that the members of the 2 subgroups probably belong to a single natural

lineage, I believe it would be clearer, on the basis of the differences in the

male phallosome and other features of the gehitalia, to distinguish the Pzpiens

Subgroup from the Tvifilatus Subgroup.

1. CULEX (CULEX) QUINQUE FASCIATUS SAY

(Bigs. 1, 2; 3,14)

Culex quinquefasciatus Say 1823: 10 (adult).

Culex fatigans Wiedemann 1828: 10 (2); Edwards 1913a: 55 (“*, taxonomy);

Edwards 1921: 345 (", 2, L, Keys); Barraud 1924b: 1264 (“*, 9);

Barraud 1924c: 430 (L*); Barraud and Covell 1928: 671-9 (2 cibarial

armature*); Borel 1930: 339 (“*, 2, L*); Stone 1956 (1957): 342

(Synonymy).

Culex cingulatus Doleschall 1856: 405 (adults; preoccupied by Fabricius 1805).

Culex doleschalli Giles 1900: 338 (new name for C. cingulatus Doleschall);

Barraud 1934: 420 (synonymy with C. fatigans).

Culex albolineatus Giles 1901: 609 (2); Barraud 1924b: 1264 (synonymy with

C. fatigans).

Culex quasipipiens Theobald 1901b: 136 (¢*); Edwards 1913a: 55 (synonymy

with C. fatigans).

Culex fouchowensis Theobald 1901b: 137 (c'*, 9*); Edwards 1913a: 55 (synonymy

with C. fatigans).

Culex reesi Theobald 1901b: 145 (o'*, 2*); Edwards 1913a: 55 (synonymy with

C. fatigans).

Culex sericeus Theobald 1901b: 147 (2*); Edwards 1913a: 55 (synonymy with

C. fatigans).

Culex christophersi Theobald 1907: 453 (c, 2); Edwards 1913a: 55 (synonymy

with C. fatigans). |

Culex minor Theobald 1908: 298 (c", 2); Barraud 1924b: 1264 (synonymy with

C. fatigans).

Culex hensemaeon Dyar 1920: 178 (2); Edwards 1932: 208 (synonymy with

C. fatigans).

For complete synonymy, see Stone, Knight and Starcke (1959).

Culex (Culex) fatigans Wiedemann, Edwards 1932: 208 (taxonomy); Barraud

1934::420 (6%, 2.50%):

Sirivanakarn: Subgenus Culex in the Oriental Region 31

Culex (Culex) quinquefasciatus Say, Bohart 1945: 79 (o, L); LaCasse and

Yamacuti 1950: 220 (¢* > o*; L*); Belkin. 1962): 195: (o*, 9; P*, L*);

Stone 1963; 135 (catalog supplement); Delfinado 1966: 148 (\*, 9, P, L);

Baisas 1974: (“, 2*, P, L*).

Culex (Culex) pipiens quinquefasciatus Say, Stone, Knight and Starcke 1959: 254

(catalog); Bram’ 1967ar 192 (o*, o*. Pe, 1.*),

FEMALE (Fig. 1, 14). Wing: 2.5-4.3 mm (average 3.5 mm). Forefemur:

1.1-2.0 mm (average 1.6 mm). Proboscis: 1.3-2.2 mm (average 1.8 mm).

Abdomen: 1.9-3.2 mm (average 2.5mm). Usually medium sized species.

Head, Narrow decumbent scales on vertex pale, yellow or beige; erect scales

usually dark brown, sometimes pale, yellow or golden in center, dark on

posterolateral area; lateral patch of broad appressed scales whitish. Palpus

0.2-0.25 of proboscis length, entirely dark scaled or sometimes with some

pale scales on apex of segment 4. Proboscis entirely dark scaled on dorsal

surface, ventral surface dark or sometimes pale from base to middle. Anten-

nal pedicel with a minute patch of semi-erect scales on inner dorsal surface;

flagellomere 1 with or without a few scales on inner surface, flagellar whorl

very weakly plumose, with 5-6 dark bristles. Cibarial Armature. Cibarial

bar concave with slight projection at middle; cibarial teeth short, abruptly

pointed and subequal, about 30 in number. Thorax. Mesonotal integument

brown; mesonotal scales narrow, predominantly yellowish to brownish; scales

on prescutellar space and caudal margin of scutellar lobes pale whitish; acro-

stichals, dorsocentrals and supraalar bristles dark and strong; midscutellar

lobe with 7 bristles; lateral scutellar lobe with 4-5 bristles. Pronotum same

color as mesonotum; apn with several narrow scales and short setae; ppn with

4 strong posterior bristles and 3-4 short setae cephalad, upper surface with

patch of several narrow scales. Pleuron slightly paler or same color as

mesonotum, without distinct pattern of dark bands or stripes; pp/ with 4-9

bristles and a small pale scale patch at base; stp with 2 moderately broad,

pale scale patches on upper corner and lower posterior border; mep with

similar scale patch at level of upper corner of stj and several pale scales

among upper mep bristles; lower mep bristle strong and dark, usually 1, some-

times 2. Legs. Anterior surface of forecoxa with broad scale patch and sev-

eral moderately strong setae, scales on upper surface pale, rest dark; mid-

and hindcoxae with narrow anterior scale patches; anterior surfaces of fore-

and midfemora with narrow pale bands at bases and entirely dark scaled distally;

anterior surface of hindfemur with a distinct pale stripe from base to near apex;

all tibiae and tarsi entirely dark on anterior and dorsal surfaces, pale on ven-

tral surfaces; all pretarsal claws small and simple. Wing. All scales dark and

very dense on all veins; scales on veins R9, Rg and R445 narrow, linear and

elongate; length of cell Ro 1.5-2.0 of vein Ro,9; remigial bristles short and

dark, 4-5 innumber. Halter, Peduncle pale, without scales; capitellum cup-

shaped, covered with several pale beige scales. Abdomen. Tergum I with

dark median scale patch; terga II- VII with evenly broad or slightly medianly

produced basal pale bands; basolateral pale spots present, distinct, apparently

not connected with basal transverse bands dorsad; sterna predominantly pale

yellowish or sometimes with dark scaled areas in middle and apicolateral mar-

gin. Genitalia. Tergum IX with lateral row of 10-12 setae; cerci short, apical-

ly blunt, with several strong setae restricted to lateral tergal surface; post-

genital plate broad, with rounded caudal margin, apical half with a lateral row

of 5-6 bristles; vaginal sclerite dark, well sclerotized, U-shaped; insula with

a dense tuft of about 10 strong setae.

32 Contrib. Amer. Ent. Inst, vol. 12). n0,.2; 1976

MALE (Fig. 1). In general similar to female except for sexual characters.

Head. Palpus variable in length, exceeding proboscis by 0.5-1.0 of segment

5; segment 3 dark scaled on dorsal surface, with ventrolateral rows of 4-12

strong bristles in apical 0.25-0.30; segments 4 and 5 upturned, moderately to

strongly long plumose, dorsal suriace dark scaled, ventral surface of segment

4 with white scale line from base to 0.79 of total length, ventral surface of

segment 5 with white scale patch restricted to base. Proboscis with submedian

false joint distad of mid point of labium, scales on ventral surface usually pale

_ in middle; median ventral tuft of long hairs proximad of false joint absent.

Legs. Pretarsal claws of fore- and midlegs enlarged, unequal, each with sub-

basal denticle; claws of hindleg small, equal and simple.

MALE GENITALIA (Fig. 2). Segment VIII. Tergum VIII shallowly emargin-

ate at middle, lateral caudal margin with several strong bristles. Segment IX.

Tergum IX with broadly rounded tergal lobe, bearing 1 or 2 irregular rows of

10-12 strong setae, sometimes with 6-9 setae; sternum IX broad, roughly

rectangular, without scales or setae. Basimere. Slender, conical, 0.28-0. 34

mm in length, strong bristles largely confined to lateral tergal surface; inner

tergal surface with 3-4 rows of subequal setae from near base to level of sub-

apical lobe. Subapical lobe. Clearly divided into proximal and distal divisions;

setae a-c of proximal division stout, rodlike and with hooked apices; distal

division with a strong, flattened seta and 2 narrower and shorter setae in group

d-f, all clearly separated from bases of setae a-c in proximal division; leaf g

broad with acuminate or blunt apex; seta strong, flattened and long. Distimere.

Normal, sickle-shaped; dorsal subapical surface without crest of spicules or

serration; 2 tiny ventral setae present slightly beyond midpoint of curvature;

dorsal seta absent; subapical claw (or spiniform) short. Phallosome (Aedeagus).

Outer division with a strong and long mesal, tergal arm which tapers into a point

apically, tergal arms on both lateral plates nearly parallel, lateral spine very

‘short, inconspicuous; sternal spine moderately strong, distinct, strongly diverg-

ent laterad: inner division simple, represented by a broad leaflike ventral arm

which is strongly divergent laterad; DV/D ratio usually 1 (0.7-1.0). Proctiger.

Apical crown large, dark, with several flat, apically blunt spicules laterally

and numerous pointed spinelike spicules mesally: paraproct broad with some

small tubercles below crown; basal sternal process very short, unpigmented,

about 0.03 mm in length, sometimes absent or not developed; cercal sclerite

narrow; cercal setae 2-3 in number; basolateral sclerotization very well

developed.

PUPA (Fig. 2). Abdomen: 2.5-3.3 mm. Paddle: 0.7-0.9 mm. Trumpet:

0.52-0.78 mm. Pigmentation generally cream-colored or pale yellowish.

Trumpet. Darker than underlying integument, relatively short, narrowest at

base, broader towards apex, pinna long and strongly oblique, apical margin

truncate or lightly emarginate. Cephalothorax. Seta 1-C usually triple (2-4);

2-C 3,4 branched; 3-C double or triple; 4-C 3,4 branched; 5-C usually 4,9

branched, 6-C double or triple; 7-C double; 8-C usually triple (2-3); 9- C double.

Metanotum. Brown in middle, pale towards lateral margin; seta 10-C 9-10

branched; 11-C double; 12-C usually triple (2-4). Abdomen. Segments I-IV

darkened in middle, pale towards lateral margin, other segments pale; setae

6-1, II single; 7-I, II double; 1-II multiple, or usually more than 10 branched

(8- 16): 3-I-III double; 1- II, IV usually 6 branched (5-7); 1-V-VII usually 4, 9

branched (3-5); 5-IV usually double (2-5); 5-V, VI usually double (1-3); 6- I- VI

usually double (2-4); 9-VII 3-6 branched, 9-VIII 4-8 branched, pectinate, slightly

longer than 9-VII; 4-VIII double. Paddle. Broad and pale, without distinct color

pattern, external buttress moderately strong and distinct; distal margin

Sirivanakarn: Subgenus Culex in the Oriental Region BS

indistinct; midrib lightly darkened; setae 1, 2-P present, minute, subequal and

single.

LARVA (Fig. 3). Head: 0.76-0.90 mm. Siphon: 1.2-1.7 mm; index 3-5.

Saddle: 0.4 mm; siphon/saddle ratio 3-4. Head, Rather large and more or

less rounded; labrum (or preclypeus) narrow; seta 1-C pale, filiform; 4-C

single, strong, about 2 times as long as distance between bases of the pair;

0, 6-C strong, subequal and usually 4,5 branched (3-6); 7-C large, fan-

shaped, usually 9, 10 branched (8-12). Antenna short and straight, about 0.5

of length of head; middle part of shaft not distinctly curved outwards; pigmenta-

tion varying from pale to darker than head capsule; seta 1-A 22 branched, situ-

ated slightly beyond 0.5 of antennal shaft; 2, 3-A weak and pale, situated almost

at apex; 4, 6-C weak and rather short. Mental plate with 10-12 lateral teethon

each side of median tooth. Thorax. Seta 4-P double; 7-P usually double (1-2);

8-P usually double, sometimes single; 1-M single, about 0.75 of 3-M; 3-M

single; 4-M double; 8-M 5, 6 branched; 9-M 4, 5 branched; 1-T short, about

0.9 of 2-T; 7-T 6,7 branched; 9-T 6 branched; 12-T single; 13-T 4, 5 branched

(3-6). Abdomen. Setae 1-I, II as conspicuous as seta 5-I; 1-I 4,5 branched;

1-I 4,5 branched; 1-II double or triple; 6-I, II usually triple (3-4); 7-I double;

1-III-VI single or double, rather strong, about 0.75 of length of setae 6-III-VI;

6-III-VI all double; 13-III-V weak, about 0.5 of 1-III-V, usually triple (2-3);

seta 1-VII usually 4 branched (3-4); 4, 7, 10 and 12-VII all single; 13-VII 3, 4

branched. Seta 1-VIII usually 4,5 branched (4-6); 2-VIII single; 3-VIII usually

6, 7 branched (5-7); 4-VIII single; 5-VIII usually 4 branched (3-4); comb scales

short, small, numerous, 30-44, all with rounded and even fringe of fine spicules

apically and laterally. Posterior dorsal margin of saddle weakly spiculated;

seta 1-X usually single (1-2) and strong; 2-X usually double (2-4); anal gills

slender, 1.0-1.9 times as long as saddle. Siphon. Relatively short and stout,

fusiform, middle portion distinctly swollen, distal portion tapered; pecten with

0-10 fine teeth of increasing length in basal 0.4 of siphon, 2-3 most distal teeth

with 3 basal denticles and 1 elongate apical denticle; siphonal tufts strong,

widely spaced, usually 4 pairs, sometimes 3; first 2 proximal pairs strongest,

usually 4,5 branched (2-7), situated subventrally, 3rd pair weaker and shorter,

dorsolateral, usually double or triple (1-3); 4th pair as long as 3rd pair, sub-

apical and subventral in position, usually 4,5 branched (3-5); median caudal

filament of spiracular apparatus poorly developed.

TYPE-DATA. (1) Culex quinquefasciatus Say, Type (lost), type-locality

restricted to vicinity of New Orleans, Louisiana, USA by Belkin, Schick and

Heinemann (1966: 4); (2) Culex fatigans Wiedemann, Lectotype 2, East Indies

(INDONESIA), collection Winthem (NMW;; selection of Belkin 1968: 68); (3) Culex

cingulatus Doleschall, Type non-existent, Ambarawa, Java, INDONESIA (LU);

(4) Culex albolineatus Giles, Lectotype °*, Shahjahanpur, (United Provinces),

INDIA (BM; present selection); (5) Culex quasipipiens Theobald, Lectotype °*,

Sambalpur, (Central Provinces) INDIA (BM; selection of Bram 1967b: 327);

(6) Culex fouchowensis Theobald, Lectotype o*, Foochow, Fukien, CHINA (BM;

selection of Bram 1967a: 196); (7) Culex reesi Theobald, Type non-existent,

HONG KONG (BM); (8) Culex :sericeus Theobald, Type (lost), HONG KONG

(LU); (9) Culex christophersi Theobald, Lectotype o*, INDIA (BM; selection

of Bram 1967b: 327); (10) Culex minor Theobald, Types “ and 9, Sylhet and

Lungleh, Lushai Hills, Assam and Calcutta, Bengal, INDIA (IM); (11) Culex

hensemaeon Dyar, Holotype 2*, Los Banos, Laguna, Luzon, PHILIPPINES

(USNM).

DISTRIBUTION. Widespread throughout the tropics and subtropics of the

world. Material examined. 4,065 specimens: 1,463, 1,911°, 698 L; 740

individual rearings (86 pupal, 654 larval).

34 Contrib... Amer. Bnti:Inet.., voly 125 no. 2;°19'76

INDIA. Assam: Dooma Dooma; Pupsi; Jorhat; Chabua; Dibrugarh; Sooker-

ating; Bengal: Calcutta, Dum Dum, N. Salt Lake, Botanical garden, Sukna,

Darjeeling Dist.; Bihar: Purnea; 1380", 1802, 29 L, 8p, 12 lp.

BANGLADESH, Sylhet; loc, 262, 7 L; 2°p, 26 Ip.

SRI LANKA. Central Province: Nuwara, Eliya District; Kande-ela

Reservoir; Peradeniya, Kandy District; 20°, 6°, 7 lp.

BURMA. Rangoon; Myo Kwet Thit; 2%, 109, 1L, 4 lp.

THAILAND. Chiang Mai; Nan; Lampang; Tak; Udon Thani; Nakhon Sawan;

Nakhon Ratchasima; Ang Thong; Ayutthaya; Bangkok; Thon Buri; Phet Buri;

Chon Buri; Trat; Prachuap Khiri Khan; Surat Thani; etc.; 1390", 889, 401 L,

22 p, 35 Ip.

VIETNAM. Saigon; Nha Trang; Phu Tho; Dalat; Soc Trang; Phu Bei; Gia

Dinh; Khanh Hae; Pleiku; Can-Tho; 450°, 959, 15 L, 19 lp.

CAMBODIA. Phnom Penh; 9°, 559.

LAOS. Vientiane; 1°.

MALAYSIA. Peninsular Malaysia: Selangor: Kuala Lumpur; Ulu Gombak;

Segambut; Sea Port Estate; Brinchang; Petaling; Sg. Bertek Klang; Rantau

Panjang; Trengganu: Kuala Brang; Dungun; Kemaman; Pahang: Kg. Lamir;

Pdg. Tungku K. Lipis; Sungai Temau; C. Highlands Road; Kg. Sungai Ular;

Kuala Pahang; Perak: Lahat; Tg. Rambutan; Chiore F. R.; Perlis: Golf

Course; Kelantan: Tumpat; Gua Musang; Malaysia: Sabah: Kota Kinabalu

(Jesselton); Simpangan; Sandakan; Ling Kabau; Papar; Luhan, Ranau; Kudat;

Tagasan; Tawau; Pulau Gaya; Tuaran; Mt. Kinabalu; Kuala Penyu; Labuk Sugai;

Lahad Datu; Pulau Labuan; 2720, 3679, 44 L, 37p, 140 lp.

SINGAPORE. Pulau Hantu; Pulau Blakang Mati; 13%, 1192, 14 L, 14 lp.

INDONESIA. Kalimantan: Tarakan; Sumatra: Bengkulu; Java: Djakarta;

Pelabuhan Ratu; Bogor; Bandung; Sindanglaja; Solo; 870", 79%.

PHILIPPINES. Luzon: Manila; Los Banos; Subic Naval Base; Rizal,

Alabang; Bagiou Benguet; Bagiuo; Camp Stotsenberg; Camp Nichols; Camp

Eldridge; San Fernando; Pangasinan; Dalton; N. Vizcaya; San Fabian; Camansi,

San Juan; Bacnotan; La Union, Balaoan; Bitabag; Carlatan; Naguilian; Agoo, La

Union; Laguna; Saragosa; Camarines; Mindoro: Caminawit; San Jose; Samar:

Osmena; Palawan: Iwahig; Panay: Iloilo; Leyte: Tacloban; Jinamoc; Dulag;

Sanga Sanga Is.: Lapit Lapit; Jolo; Mindanao: Zamboanga; Pettit Barrack;

Cotabato; Corregidor Is.: Fort Mill; 4820", 5582, 143 L, 7p, 189 Ip.

TAIWAN. Taipei: Chung Ho; Chung Yuan; Taichung: Shalu; Wuchi; Tunghai,

Lungtan; Pingtung: Kao-Hsiung; Shin Lin; Nan Tou; Tai Tung; Orchid Is. ;

Miaoli, San-I; 61%, 2022, 2 L, 10 p, 202 lp.

CHINA. Yunnan, Kunming (Cazier, 1945); Hunan, Achian; Hainan, Canton;

Chekiang; Shanghai; Soochow; Tientsin; Sha-Sze (Yngtseze River); 810, 1178,

8 L

JAPAN. Ryukyus: Okinawa; Iwo Jima; Iriomote; Ie Shima; 1170, 116%,

34 L, 6 Ip.

TAXONOMIC DISCUSSION. The nomenclature and the taxonomic status

of this tropical pipiens form remain hightly controversial. It has been referred

to either as quinquefasciatus Say or fatigans Wiedemann in numerous taxonomic

and other studies without a universal consensus as to which name is the proper

one. Stone 1956 (1957) suggested that the name "“guinquefasciatus" instead of

"fatigans'' must be applied in accordance with the rules of priority if both

names are considered as synonyms. Since it has been widely or generally

accepted that the 2 names are apparent synonyms as discussed by Stone, Iam,

therefore, using the name "gquinquefasciatus" for this form. As indicated in

the discussion of the Pipiens Subgroup, I am also recognizing it as a species

Sirivanakarn: Subgenus Culex in the Oriental Region 30

in spite of the arguments and discussions in favor of the single polytypic species

concept of pipiens and of the contention that quinquefasciatus should be treated

as subspecies as proposed by Mattingly (1957, 1967) and others. Although

quinquefasciatus and other pipiens forms (e.g. molestus, pallens) are toa

various degree capable of crossing in the laboratory, they obviously maintain

themselves as distinct entities in nature even where they come into contact.

In the present circumstance, it seems most practical to consider quinquefascia -

tus aS a Species instead of a subspecies of pipiens.

Culex quinquefasciatus can be separated from typical pzpiens and other pipi-

ens forms (including molestus and pallens) from northern Europe (England,

France, Sweden), Egypt, China, Japan and Korea in the male genitalia by (1)

tergal arms of outer divisions of phallosome parallel or subparallel and distally

strongly tapered into a point (divergent and uniformly thick, with truncate or

lightly swollen apex in other pipiens forms); (2) ventral arm of inner division of

phallosome very large and broad (relatively smaller, shorter and narrower in

other pipiens forms except pallens) and (3) DV/D ratio (Fig. 2) usually more than

0.7 (less than 0.7 in other pipiens forms). The guinquefasciatus males also dif-

fer from other pipiens forms in having a shorter palpus, presence of fewer and

weaker ventral lateral bristles in apical 0. 25-0.30 of palpal segment 3 and in

having fewer and weaker bristles in palpal segments 4 and 5. Among the pip7-

ens forms, guinquejfasciatus is apparently more similar to pallens from Japan

and China in the phallosome morphology than to typical pzpzens from Europe or

molestus from Egypt and Mediterranean areas.

The Southeast Asian populations of quinquefasciatus are, as elsewhere in the

tropics, very large and extremely variable in all stages. It is possible that

there may be some local differentiation into various forms among

allopatric and sympatric populations of quinquefasciatus. In Thailand, Penin-

sular Malaysia and the Philippines, I have seen several males which are ex-

tremely small with the wing length reduced to about 2.0 mm. These minute

males also exhibit a great reduction in the number and length of bristles on

palpal segments 3, 4 and 5 but show no obvious differences from normal males

in the genitalia or other associated stages. It seems most probable that these

variations are environmentally induced, probably due to shortage of water and

increased temperature in the breeding sites, resembling the situation frequently

encountered in laboratory colonies (P. F. Mattingly, personal communication).

BIONOMICS. Culex quinquefasciatus is the most common domestic species,

occurring abundantly in houses and in practically all types of human and animal

shelters in urban communities throughout Southeast Asia. It is found breeding

in any type of habitat which contains water ranging from fresh and clear to

brackish, turbid and polluted with decayed organic matter from garbage and

human wastes accumulated in ground pools, ditches, drains, sewages, dumping

areas, latrines, septic tanks and in various kinds of artificial containers. In

urban situations, the larvae and pupae of guinquefasciatus are extremely dense

and have frequently been collected in great numbers from foul or strongly pol-

luted stagnant pools, drains, ditches, pits and sewage. The immatures have

also been frequently collected from the multitude of artifical containers, such

as bottles, cans, flower pots, vases, bowls, jars, cement tanks, etc., inside

or outside the houses. In general, quinquefasciatus is uncommon in small

containers, but common in large ones. The adults are fairly abundant through-

out the year and are commonly found resting during the day in shade, on walls,

dark corner spots, in cabinets, wardrobes, bathrooms, under tables and chairs

inside houses. The females are vicious biters at night indoors or outdoors and

feed principally on blood of man from sunset until dawn. They also frequently

36 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

attack birds such as fowls and, to a lesser extent, other domestic animals,

including dogs, cats and pigs. Various aspects on the bionomics of the adults

and immatures of guinquefasciatus (or fatigans of several authors) have been

intensively investigated by de Meillon and coworkers at the World Health Organi-

zation (WHO) Filariasis Research Unit (FRU) in Rangoon, Burma and by several

other WHO regional officers in various parts of Southeast Asia. For more ex-

tensive and detailed information on the bionomics, consult Horsfall (1955),

de Meillon et al. (1967), de Meillon, Grab and Sebastian (1967), de Meillon and

Sebastian (1967a, 1967b), de Meillon, Sebastian and Khan (1966, 1967a, 1967b,

1967c, 1967d), Lindquist et al. (1967) and Kurihara and Sasa (1965).

MEDICAL IMPORTANCE. Culex quinquefasciatus is one of the most serious

pests of man in the tropics and has been incriminated as the principal vector of

nocturnal periodic bancroftian filariasis in various parts of the Oriental region.

In Southeast Asia, it has been reported to be naturally and experimentally in-

fected with this pathogen in Bangladesh (Aslamkhan and Wolfe 1972), Burma,

Singapore, Vietnam, Malaysia, Indonesia, the Philippines, Hong Kong and

Taiwan but has never been found to be a vector in Thailand (Ramalingam et al.

1968). In some parts of Southeast Asia, it has also been reported to be a host

of certain arboviruses, most notably Chikungunya and Japanese encephalitis

but there is apparently no conclusive evidence to indicate that it plays a role in

the natural transmission of these diseases. This may in part be due to the fact

that it rarely bites reservoir animals such as pigs infected with Japanese en-

cephalitis virus. In Thailand, Rudnick and Hamon (1961) and Robin et al. (1963)

isolated several strains of Chikungunya from quinquefasciatus but later study

combined with epidemiological observations by Halstead, Scanlon et al. (1969)

suggested that it is not an important vector of this disease. Laboratory trans-

mission studies of Japanese encephalitis reported by Hodes (1946) and Reeves

and Hamon (1946) indicate that it is less efficient than gelidus (D. J. Gould,

personnal communication), Although Nguyen et al. (1974) isolated 3 strains of

Japanese encephalitis from wild caught adults in Vietnam, it is believed that

quinquefasciatus is not important in the transmission of Japanese encephalitis

in this country. Subsequent studies have shown that quinquefasciatus is a rela-

tively inefficient vector of Japanese encephalitis virus.

TRIFILA TUS SUBGROUP

The Tvifilatus Subgroup is characterized by the following combination of

male genitalia characters: (1) setae in group d-f of distal division of subapical

lobe 3,4 in number, placed between setae a-c of proximal division and leaf g of

distal division; (2) distimere without distinct serration or annulation on sub-

apical dorsal surface; (3) ventral arm of inner division of phallosome relatively

narrow, simple, diverging laterad; (4) outer division with a long lateral spine;

(5) tergal arm of outer division usually with subapical swelling or twisted apex

and (6) basal sternal process of proctiger well developed.

Mattingly and Rageau (1958: 241-50) in recognizing the Trifilatus Subgroup,

placed the following 8 species within this subgroup: trifilatus Edwards 1914

and famsi Edwards 1934 from the Ethiopian region; torrentium Martini 1929

from the western Palearctic; vagans Wiedemann 1828 from the eastern Pale-

arctic; pervigilans Bergroth 1889, pacificus Edwards 1916 and zyengari

Mattingly and Rageau 1958 from the South Pacific and mivaculosus Bonne-

Wepster 1937 from New Guinea. It also includes 2 other species: votoruae

Belkin 1968 and asteliae Belkin 1968 from New Zealand, considered by Belkin

Sirivanakarn: Subgenus Culex in the Oriental Region 37

(1968) as members of the pervigilans complex. In addition, I consider hutchin-

soni Barraud 1924 from the Oriental region as another member of this sub-

group, making a total of 11 species for the entire subgroup.

The distribution of the Trifilatus Subgroup is widely disjunct, over a very

large area, covering the western and eastern Palearctic, Ethiopian, Oriental

and Australasian regions and the South Pacific (including New Zealand). In the

Ethiopian where several other related lineages are represented, this subgroup

is still poorly known and it is possible that more members may be discovered

when the fauna is thoroughly examined.

2. CULEX (CULEX) VAGANS WIEDEMANN

(Figs. 4, 5, 14)

Culex vagans Wiedemann 1828: 545 (o", 2); Edwards: 1922b: 473 (2); Ho 1931:

162. (o"%, “Ry;

Culex tipuliformis Theobald 1901b: 325 (¢*); Barraud 1924b: 1269 (o'*, 9);

Barraud 1924c: 430 (L*); Edwards 1932: 211 (synonymy).

Culex virgatipes Edwards 1914b: 126 (%, 2); Edwards 1921: 341 (distribution);

Barraud 1924b: 1269 (synonymy with C. tipuliformis).

Culex exilis Dyar 1924: 127 (); Edwards 1932: 211 (synonymy): Monchadskii

1951: 260 (distribution).

Culex (Culex) vagans Wiedemann, Edwards 1932: 211 (taxonomy); Barraud 1934:

416 (“*, 2, L*); Feng 1938: 303 (biology, distribution); LaCasse and

Yamaguti 1950: 216 (“*, 2*, P*, L*); Monchadskii 1951: 283 (distribution);

Hsieh and Liao 1956: 123 (distribution); Hara 1957: 55 (2 genitalia*); Chu

1958: 110 (distribution).

FEMALE (Fig. 14). Wing: 4.3-5.0 mm (average 4.6 mm). Forefemur:

1.6-2.3 mm (average 2.0 mm). Proboscis: 1.8-2.5 mm (average 2.3 mm).

Abdomen: 2.9-3.2 mm (average 3.0 mm). Medium or large sized species;

generally as described for the Pipiens Group, with the following distinctive

features. Head. Erect scales on vertex pale yellowish in center and towards

anterior dorsal eye margin, dark brown posterolaterally and towards occiput;

narrow decumbent scales pale, forming a distinct ocular line; lateral patch of

broad scale whitish, very distinct. Palpus entirely dark scaled or with a row

of pale scales on inner dorsal surface of segment 4. Cibarial Armature, Es-

sentially similar to guinquefasciatus. Thorax. Mesonotal scales very narrow

and fine, predominantly yellowish brown or golden, pale whitish on anterior

promontory and on lateral margin; scales on prescutellar space and scutellar

lobes whitish. Pleural integument without pattern of dark and pale stripes; ppl

with a small pale scale patch at base and 10 setae; stf with 2 distinct pale scale

patches, one of which is on the upper corner, the other on lower posterior

border; mep with distinct pale scale patch at same level as upper corner of sip

and several pale scales among mep bristle; 1 lower mep bristle present. Legs,

Anterior surface of forefemur with a distinct, narrow, linear white stripe from

base to apex along anterior margin; anterior surface of midfemur with a distinct

narrow white stripe along whole length in middle; anterior surface of hindfemur

largely pale in basal half, continuous as a narrow stripe in apical half; all tibiae

with a distinct longitudinal white line on anterior surface;tarsomeres1-5 of fore-

and midlegs entirely dark; tarsomere 1 of hindleg with a distinct longitudinal

white line on anterior surface, rest dark. Abdomen. Tergum II with a medi-

an basal pale streak, not extended laterad to form a complete band; terga

38 Contrib. Amer, Hat: Inst, Vole: 12,;°no. 2), 1976

III- VII with broad, transverse basal pale bands; tergite VIII largely yellowish.

Genitalia. Essentially as described for quinquefasciatus.

MALE. In general similar to female. Head. Palpus long, exceeding

proboscis by a little more than the length of segment 5; segment 3 largely dark

scaled on dorsal surface, with or without some pale scales on lateral surface

beyond middle; apical half of segment 3 with ventrolateral rows of about 36

dark bristles; segments 4 and 5 strongly long plumose and upturned; dorsal

surface with narrow basal pale bands; ventral surface of segment 4 with a

distinct white scale line; ventral surface of segment 5 with scattered pale

scales in basal 0.3-0.4. Proboscis without median ventral tuft of long hairs;

ventral surface slightly pale in middle.

MALE GENITALIA (Fig. 4). Segment IX. Tergum narrow, with broadly

rounded tergal lobe which bears 2 irregular rows of 8-10 weak setae. Basi-

mere. Slender, roughly conical, about 2.8 mm inlength. Subapical lobe.

Proximal and distal divisions clearly divided; setae a-b of proximal division,

stout, subequal; seta c thinner, all with hooked apices; setae d-fof distal

division 3 in number, placed clearly distad of setae a-c; leaf g broad, foliate,

with rounded apex; seta h strong and long. Distimere. Normal, sickle-

shaped; ventral surface with 1,2 tiny setae beyond middle; dorsal surface with-

out any seta; claw slender and moderately long; subapical dorsal surface without

distinct serration. Phallosome. Outer division of lateral plate with a strong

lateral spine, divergent laterad; tergal arm long, with twisted or slightly swol-

len apex; sternal spine not developed or inconspicuous; lateral basal process

small and slender; inner division with a slender, smoothly curved ventral arm.

Proctiger. Basal sternal process well developed, moderately long, curved

sternad; paraproct narrow; cercal sclerite well sclerotized; cercal setae 3-4

in number; basal lateral sclerotization well developed.

PUPA (Fig. 4). Abdomen: 3.0mm. Paddle: 0.85 mm. Trumpet:

0.65 mm; Index 5. Chaetotaxy as figured, very similar to quinquefasciatus,

differing particularly in the following. Trumpet. Relatively longer and more

or less cylindrical along the whole length; pinna slightly oblique. Metanolum.

Seta 10-C 5-7 branched; 12-C 3-6 branched. Abdomen. Seta 1-II 10-14

branched; 1-III usually 8 branched (7-10); 5-IV usually triple (2-3); 6-III double

or triple; 6-IV, V usually triple; 6-VI 4,5 branched (3-5). Paddle. Seta 2-P

stronger than 1-P, usually bifid (1-2).

LARVA’ (Fig. 5): Head: 0:9 mm. Siphon: 1.8 mm; index 5 or more.

Saddle: 0.4 mm; siphon/saddle ratio 4.5. General characters and detailed

chaetotaxy as figured, distinctive in the following features. Head. Labrum

narrow; seta 1-C filiform or distally filamentous; 5, 6-C usually 5 branched

(4-7); 7-C 8,9 branched. Antenna about 0.7 of head length, pigmentation dark

from base to apex; seta 1-A 20 or more branched, strongly pectinate; 2, 3-A

subapical. Mental plate with 9, 10 lateral teeth on each side of median tooth.

Thorax. Seta 4-P double; 7, 8-P double; seta 1-M nearly as long as 3-M,

single; 2-M short, double or triple; 3-M single; 4-M double; 8-M 5-7 branched;

9-M 4,5 branched; seta 1-T minute, usually triple (3-4); 2-T longer, double;

3-T usually 5 branched (4-5); 7-T 7-9 branched; 9-T 5,6 branched; 12-T single;

13-T 5,6 branched. Abdomen. Seta 1-I, II distinct, same length as 5-I, II; 1-14

branched; 1-II bifid; 6-I, II 3,4 branched; 7-I double; 1-III-VI, strong, about 0.5

of 6-III-VI, all double; 6-III- VI double; 13-III-V nearly as strong as 1-III-V, all

usually 4 branched (3-5). Seta 1-VIII usually 5 branched (5-7); 2,4-VIII single;

3- VIII 7-9 branched; 5-VIII usually 4 branched (4-5); comb scales 34-37, all

small, subequal in size and with even fringe of fine spicules apically and later-

ally. Seta 1-X of saddle strong, single, about 0.5 of saddle length; 2-X double;

Sirivanakarn: Subgenus Culex in the Oriental Region oo

anal gills 1.5-2 times as long as saddle.

_ TYPE-DATA. (1) Culex vagans Wiedemann, Types “, °, (Foochow,

Barraud 1934), CHINA, Trentepohl, collection Winthem (NE); (2) Culex tipuli-

formis Theobald, Holotype @*, Bakloh, N.W. Province, {| ? PAKISTAN] (BM);

(3) Culex virgatibes Edwards, Holotype ¢* with attached genitalia mount,

HONG KONG (BM): (4) Culex exilis Dyar, Holotype “*, Vladivostok, | Maritime

Territory |, USSR (USNM).

DISTRIBUTION. Widespread in the northeastern Palearctic (China; USSR,

Siberia; Korea and Japan) with extension into the south as far as India, south

China, Hong Kong and Okinawa. Material examined. 129 specimens: 58,

4992, 22 L; 20 individual rearings (1 pupal, 19 larval).

INDIA. W. Himalaya, Kasauli; Solon, Simla Hill; Punjab, Mahdapur;

Assam; Ledo;. 70). 49, 3).L, 7 po

CHINA. Peking; Shanghai; Canton; Foochow; 100, 102, 7 L.

HONG KONG. 2°¢, 29.

JAPAN. Nagasaki; Hokkaido; Yokohama; 5c, 39, 5 L. Ryukyus: Okinawa,

1.5 mi. N.W. of Futeuma; Iriomote, Yabu Village; 11°, 89, 19 lp.

KOREA. Seoul; Kimpo; Tong Jon; Saemal; Chorwan; Chail Li; T'Andong;

Ld @ 12 eee

USSR. Far East, South Primorje; Ussuri River; 50, 9°. Siberia.

Okeanskya; 19.

Additional records from the literature: INDIA: Kashmir; Delhi; USSR:

Siberia; Vladivostok (Barraud 1934: 418; Monchadskii 1951: 211); PAKISTAN;

BANGLADESH (Aslamkhan 1971).

TAXONOMIC DISCUSSION. Culex vagans is widespread and fairly common

in Japan, Korea and China but is not known to extend its range into Southeast

Asia south of Hong Kong. The adults of vagans are clearly marked by the

presence of striking longitudinal pale stripes on femora and tibiae of fore- and

midlegs and a similar stripe on the tibia of hindleg, resembling the features

noted in theilevi and univitiatus. This conspicuous feature separates vagans

from all other species in the Tvzftlatus Subgroup and most of the Pipiens

Group. It can be readily distinguished from theileri and univitiatus in the

adults by the absence of scale patches on postspiracular area and at the base

of prealar knob. The male genitalia of vagans are apparently identical to

mivaculosus Bonne-Wepster 1937 from New Guinea and torrentium Martini

1925 from England, which I have also examined for comparison.

The pupa and larva of vagans are very similar to quinquefasciatus in

chaetotaxy and in several general characters. The pupa can be separated from

the latter by (1) longer and more or less cylindrical trumpet; (2) weakly oblique

pinna; (3) seta 6-VI usually 4,5 branched and (4) seta 5-IV usually triple. The

larva is distinguished from quinquefasciatus by (1) setae 2, 3-A subapical; (2)

longer siphon; (3) middle portion of siphon lightly swollen and (4) presence of

4,5 pairs of siphonal tufts.

Except for the variation in size, all adult specimens of vagans which I

have seen from various parts of its range are similar and essentially conform

to a single type. In the larvae, the only variation noted are in the length of

the siphons which are shorter and thicker in specimens from India, but longer

and more slender in material from the Ryukyus, Japan and Korea. It is prob-

able that this variation is environmentally induced. 3

BIONOMICS. Culex vagans is a typical ground pool breeder and appears

to be restricted to high elevations in hills or mountainous areas. The immatures

of vagans were collected from various types of ground pools, including stream

pools, marshes, lakes, ponds and pools in stream-beds or at the margin of

40 Contrib. Amer. Ent. Inst.) vol 12, no. 2, 1976

streams, which usually contain green algae (Barraud 1934; Hsiao 1946; Hsiao

and Bohart 1946; LaCasse and Yamaguti 1950). In recent collections from

Okinawa and Iriomote of the southern Ryukyus, all specimens of vagans were

taken from unshaded stream pools in the hills. In Japan, it was reported to

be collected at elevations of over 1,500 m and was found to be more common

in the north than in the south (LaCasse and Yamaguti 1950). Little is known

about the adult biology and medical importance of vagans, In Japan and Korea

it was reported to enter houses and bite man at night (Hsiao and Bohart 1946;

Barnett and Toshioka 1951). It was reported to be experimentally and naturally

infected with Wuchereria bancrofti (Cobbold) (Bohart 1946, Manson-Bahr 1959,

Hsiao 1948, Hsiao and Bohart 1946).

3. CULEX (CULEX) HUTCHINSONI BARRAUD

(Figs. 6.7%, G.c14)

Culex hutchinsoni Barraud 1924b: 1261 (o*, 9).

Culex (Culex) hutchinsoni Barraud, Edwards 1932: 212 (taxonomy); Barraud

1934: 423 (¢*. 2); Colléss 1955:.316 (¢, 9): Li*); Bram 1967a: 188 (o'*,

OSPF OL) eaeio 1972) 208: (o%,) Sy:

FEMALE (Fig. 6, 14). Wing 2.9-3.6 mm (average 3.2 mm). Forefemur:

1.4-1.7 mm (average 1.5 mm). Proboscis: 1.6-1.9 mm (average 1.7 mm).

Abdomen: 2.3mm. Relatively small species, general coloration reddish

brown. Head. Narrow decumbent scale on vertex yellowish brown or golden,

not forming distinct ocular line; all erect scales entirely dark brown or some-

times slightly pale towards apices; lateral patch of broad scales whitish, dis-

tinct. Palpus rather short, about 0.15 of proboscis length and entirely dark

scaled; no scattered pale scales on apex of segment 4. Proboscis entirely

dark scaled. Cibarial Armature. Cibarial bar with 30-34 very short and

pointed teeth. Thorax. Mesonotal integument reddish brown; mesonotal

scales narrow, fine, moderately dense and predominantly golden or yellowish

brown except for lighter ones on anterior promontory, antealar and supraalar

areas; scales on prescutellar space and scutellar lobes sparse and pale whitish.

Pronotal integument dark brown; apn with narrow golden brown scales and

bristles; ppv dark brown, with 5,6 strong posterior bristles and 5, 6 tiny setae

cephalad, upper surface with several scattered brown scales. Pleural integu-

ment usually with pattern of dark spots or bands on ssp, psp, base of prealar

knob, lower posterior border of stp, upper and lower mep, other areas lighter,

concolorous with mesonotal integument; scale patches present on ppl, upper

corner and lower posterior border of stp, upper mep and a few scattered pale

scales among upper mep bristles; 1 lower mep bristle present. Legs, Anterior

surface of fore- and midfemora dark scaled; anterior surface of hindfemur with

pale longitudinal stripe from base to near apex; all tibiae and tarsi entirely dark.

Wing. ‘Scales dark and dense on all veins; scales on veins Ro, Rg and R4i5

narrow, linear and fine. Abdomen. Tergum I with median patch of dark scales;

tergum JI with median basal pale streak, not forming complete band; terga III-

VII with narrow complete basal pale bands; tergum VIII largely pale yellowish;

lateroterga II-VII with basolateral pale spots extending posteriorly as streak

to the apical margins of segments; sterna largely yellowish. Genitalia. Tergum

[IX with a lateral row of 4-7 setae; postgenital plate rounded on caudal margin;

vaginal sclerite poorly sclerotized, U-shaped; insula with a dense tuft of 10

setae.

Sirivanakarn: Subgenus Culex in the Oriental Region 41

MALE (Fig. 6). Generally similar to female. Head. Narrow decumbent

scales on vertex paler, almost whitish. Palpus exceeding proboscis by full

length of segment 5; middle portion of segment 3 slightly pale on lateral dorsal

surface, but not forming distinct ring, apical 0.25 with ventrolateral tuft of 5-7

dark bristles; segments 4 and 5 slightly upturned, weakly to moderately plu-

mose, dorsal surface dark scaled, ventral surface with pale scales more or

less restricted to bases. Proboscis entirely dark scaled; median ventral tuft

of long hairs absent.

MALE GENITALIA (Fig. 7). Segment IX. Tergum narrow; tergal lobe

with 4-6 moderately long setae. Basimere. Small, slender, conical, about

0.25 mm in length. Subapical lobe. Setae a-c of proximal division slightly

overlapped with setae.d-f of distal division; setae @-c subequal in length and

with hooked apices; setae d-f of distal division consist of 1 fine, hairlike seta

and 3 rodlike setae, latter all hooked apically; leaf ¢ broad with acuminate apex;

seta h strong, long and apically hooked. Distimere. Normal; 2 tiny ventral

setae present beyond the middle; claw small; subapical dorsal surface without

crest of spicules. Phallosome. Very similar to vagans; differing as follows:

tergal arm of outer division shorter and tapered into a point apically; sternal

spine present in addition to lateral spine; lateral basal process strong and

rounded; ventral arm of inner division strongly bent or angled laterad. Pyroc-

tiger, Apical crown medium sized, consisting of numerous pointed spicules;

basal sternal process of paraproct well developed, pigmented, curved sternad;

cercal sclerite well sclerotized; cercal setae 2 in number.

PUPA (Fig. 7). Abdomen: 2.3mm. Paddle: 0.6 mm. Trumpet: 0. 52

mm; index 6-7. Complete chaetotaxy as figured; cephalothorax and abdomen

pale, cream-colored or yellowish white. Trumpet. Slender, relatively long

and uniformly cylindrical; pinna lightly oblique. Cephalothorax. Setae 1-and

3-C usually double (2-3); 4, 5-C subequally long, usually 4 branched (3-5);

8-C usually triple (2-5); 9-C double. Abdomen. Seta 1-II strongly dendritic,

20 or more branched; 1-III 5, 6 branched; 1-IV-VII 4, 5 branched; 5-IV usually

triple (2-3); 5-V-VI double; 6-III-V usually double (1-3); 6-VI double or triple;

9-VII usually triple (2-3); 4-VIII double; 9-VIII usually 6,7 branched (6-9).

Paddle. Pale, or same color as preceding abdominal segment; midrib very

lightly pigmented; external buttress distinct, distal outer margin indistinct;

setae 1, 2-P minute, subequal; 1-P single or branched; 2-P single.

LARVA (Fig. 8). Head: 0.7mm. Siphon: 0.78 mm; index 2-3. Saddle:

0.3 mm; siphon/saddle ratio about 2.5. Readily distinguished from all other

members of the Pzpzens Group by the short and stout siphon which bears a

characteristic subapical spine on dorsal surface. Head, Seta 1-C brownish,

spiniform; 4-C single and long, its apex reaching slightly beyond anterior mar-

gin of frontoclypeus; 5, 6-C usually triple (3-5); 7-C usually 8 branched (8-10).

Antennal shaft entirely pale except for basal dark ring; spicules very fine;

setae 2, 3-A slightly subapical. Mental plate with 10-12 lateral teeth on each

side of median tooth. Thorax. Seta 3-P weak and short, about 0.5 of 1, 2-P;

4-P as long as 3-P, 4,9 branched; 7-P usually double (2-3); 8-P double; seta

1-M very strong and as long as 3-M, 9-9 branched; 3-M single; 4-M double; 8-M

6,7 branched; 9-M 5 branched; seta 1-T strong, as long as 2-T, usually 4

branched (4-5); 2-T double; 3-T usually 6 branched (5-8); 7-T 10-12 branched;

9-T 6,7 branched; 12-T single; 13-T 6-8 branched. Abdomen. Setae 1-I, II

strong, very conspicuous, 3-9 branched; 6-I, II triple; 7-I single; 6-IJI-VI

single; 6-IV usually triple (3-4); 6-V usually 4 branched (3-4); 1-III-VI 4,5

branched. Comb scales 32-35, in oval patch, all small, similar in size, with

lateral and apical fringe of evenly fine spicules. Saddle with few spicules on

42 Contrib.) Amer.) Ent.ciInst. 9" yolw 12) no. :2,:1976

lateral posterior margin; seta 1-X single, as long as saddle; 2-X double; anal

gills as long as or slightly longer than saddle. Szphon. Short and stout, re-

sembling quinquefasciatus in shape, with characteristic dorsal subapical spine

in apical 0.25; pecten teeth slender, 4-8 in number, distal teeth with 2 small

basal denticles and a long tapering apical spine; siphonal tufts moderately

strong, widely spaced; usually 4 pairs, sometimes 4.5 pairs; first proximal

pair subventral, situated among pecten, 5-7 branched; 2nd proximal pair sub-

ventral, distad of pecten, 4-6 branched; 3rd distal pair lateral, 3-5 branched;

4th pair subventral, 2-4 branched; median caudal filament of spiracular appara-

tus poorly developed.

TYPE-DATA. Lectotype “*, Nongpoh, Assam, INDIA, larvae from road-

side pools, July 1922, P. J. Barraud (BM; selection of Bram 1967a: 192).

DISTRIBUTION. Known only from India, Pakistan, Thailand, Cambodia,

Vietnam, Peninsular Malaysia and Singapore. The Philippine record by Basio

(1972) is doubtful. Material examined. 192 specimens: 620", 542, 76 L; 59

individual rearings (11 pupal, 48 larval).

INDIA. Assam: Nongpoh, Khasi Hills District; 1c’, 19 (type-series).

PAKISTAN. Lahore: 8%, 28.

THAILAND. Chiang Mai: Chang Khien; Doi Sutep; San Sai; Ban San

~Khayoms 8¢,.72, 8:ips.Lampang: Ban Pang Pako: 11¢; 1492, 6 L, 19.P, 6 lp;

Phrae; Ampur Sa; 10, 12; Udon Thani: Ban Nong Bua; 1“, 22, Nakhon Ratcha-

sima; Pak Chong, Khao Yai; 2%, 12; Dhon Buri: Ban Khun Non, Wat Choo

Arm; Bang Kok Noi, Wat Sai; 17c", 13%, 46 L, 9 p, 12 1p; Chon Buri: Sriracha;

Ban Huey Luk; 2%, 2°, 3 L, 1p, 2 lp; Prachuap Khiri Khan: Muang; Huey Som

Langs 2%:

CAMBODIA. Pailin; 1 L.

VIETNAM. An Khe; Gia Rai; Lai Kha; Bien Hao; Antuc; 1°, 19, 10 L.

MALAYSIA. Peninsular Malaysia (Malaya): Selangor: Kapang; Kuala

Lumpur airport: 3c. 22, 349; Perak: Te. Rambutan 40). 6229 Lb, lpyct 1p;

Pahang: Kuantan; 6c, 392, 1 lp.

SINGAPORE. West Coast; Buona Vista; Kranji; Pulau Blakang Mati; 5c,

02, Dig e8 Ls

Additional records from the literature. PHILIPPINES (Basio 1972).

TAXONOMIC DISCUSSION. Culex hutchinsoni is very similar to quinque-

fasciatus in the general external characters of the adults but is very different

from the latter in the male genitalia and larval stages. The only external

characters of the adults which usually distinguish hutchinsoni from quinque-

fasciatus are: the relatively smaller size; the reddish brown mesonotum and

the presence of pattern of dark bands or spots above and below the white scale

patches on the upper corner of sternopleuron and upper part of mesepimeron.

The dark bands or spots on the pleuron of the adult, resemble those of fusco-

cephala , but are less striking and not very clearly marked.

Culex hutchinsoni shows a stronger affinity with the members of the Trifilatus

Subgroup than with the others in the type of male phallosome and on this basis

Iam assigning it to this subgroup. It can be readily separated from other mem-

bers of the Tvifilatus Subgroup and all other pipiens forms by the following

characters: inthe male genitalia by (1) outer division of the phallosome with a

prominent sternal spine in addition to lateral spine; (2) lateral basal process of

outer division rather thick and prominent; (3) ventral arm of inner division nar-

row and strongly angled or bent laterad and (4) setae in group d-fof distal divi-

sion of the subapical lobe 4 in number; the pupa by (1) seta 1-II 20 or more

branched and (2) setae 6-III- V usually double; and in the larva by (1) the dark,

spiniform seta 1-C; (2) seta 4-P weak, 4,5 branched; (3) setae 1-M and T

Sirivanakarn: Subgenus Culex in the Oriental Region 43

strong, very distinct; (4) seta 7-I single and (5) siphon, short, thick, more or

less similar to guinquefasciatus in shape and with a prominent subapical spine

on dorsal surface.

BIONOMICS. Culex hutchinsoni is apparently a rather uncommon, rural,

inland species. The breeding habitat includes various types of ground pools

such as ponds, wells, ditches, rock pools and animal foot prints. It has also

been found breeding in artificial containers such as discarded oil drums and

plastic cans. In most instances, the breeding habitats such as rice fields and

vegetable plantations were fully exposed to sunlight. The elevation ranges from

near sea level to about 300 m or possibly higher. It was usually found in asso-

ciation with Culex (Lophocevaomyia) infantulus Edwards and rubithoracis (Lei-

cester), and C. (Lutzia) fuscanus Wiedemann in the collections from ground

pools and with Aedes (Stegomyia) albopictus (Skuse) in the collection from arti-

ficial containers. Several adults were collected from light traps. An engorged

female from Thailand was collected biting but no specific data were available on

the host. Nothing is known about the medical importance of hutchinsont.

THEILERI SUBGROUP

The Theileri Subgroup is characterized by the following combination of

male genitalic characters: (1) setae in group d-f of distal division of subapical

lobe 3 in number, situated between setae a-c of proximal division and leaf ¢

of distal division; (2) distimere with a very weak crest of minute spicules on

subapical dorsal surface; (3) ventral arm of inner division of phallosome

more or less straight, or slightly divergent and with 3-4 strong apical teeth

or spines; (4) outer division with or without a short sternal spine; (5) tergal

arm of outer division distally tapered into a point and (6) basal sternal process

of proctiger well developed.

The proposed Theilevi Subgroup includes only ¢hezleri Theobald 1903,

originally described from the Ethiopian region. It is apparently a common

species with a very broad range of extension northwards and eastwards into

the Mediterranean, Middle East, northern Palearctic (USSR) and the Oriental

region. Some nominal taxa which are currently synonyms of theilevi may prove

to be distinct. The Theilevi Subgroup may also include other Ethiopian species —

such as andersoni and simpsoni (Edwards 1941), but no attempt is made to

assign any of these to this subgroup largely because I have not studied them in

sufficient detail. :

4, CULEX (CULEX) THEILERI THEOBALD

(Figs. 9, 10)

Culex theileri Theobald 1903: 187 (o“*, 2*); Barraud 1924b: 1266 (o“*, 2);

Barraud 1924c: 429 (L*).

Culex creticus Theobald 1903: 189 (2); Edwards 1932: 210 (synonymy).

Culex pettigrewii Theobald 1910b: 15 (2); Edwards 1911: 262 (synonymy).

Culex onderstepoortensis Theobald 1911a: 265 (2*); Edwards 1932: 210

(synonymy).

Culex theileri var. annulata Theobald 1913: 321 (2); Edwards 1932: 210

(synonymy).

Culex tipuliformis of Edwards 1912: 31 (synonymy); Edwards 1921: 339 (L*,

distribution); Kirkpatrick 1925; 115 (o*,°?, P*,; L*).

44 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

Culex (Culex) theileri Theobald, Edwards 1932: 210 (taxonomy); Barraud 1934:

414 (o*, 9, L*); Edwards 1941: 305 (o, 2, P); Hopkins 1952: 289 (L*).

FEMALE (Fig. 9). Wing: 5.0mm. Forefemur: 2.3 mm. Proboscis:

2.5mm. Abdomen: 3.4mm. Large species; readily distinguished from other

Oriental species in the Pipiens Group by the presence of broad scale patch on

postspiracular area and by the presence of longitudinal pale stripes on the an-

terior surface of femora and tibiae of legs. Head, Narrow decumbent scales

of vertex yellowish in center, golden brown or bronzy on lateral area; erect

scales entirely brownish; lateral patch of broad scales yellowish white. Palpus

about 0.2 of proboscis length; palpal segments 3 and 4 with pale scale line on

inner dorsal surface or sometimes largely pale scaled. Proboscis slightly

pale in middle but not forming distinct ring. Flagellomere 1 of antenna with a

few pale scales on inner surface. Cibarial Armature, Not studied. Thorax.

Mesonotal integument light to dark brown; mesonotal scales rather coarse,

golden brown or bronzy in center of disc, yellowish white on marginal areas,

prescutellar space and scutellar lobes. Pleural integument same color as

mesonotum; ppl with pale scale patch, extending anteriorly to prosternum, ppl

bristles strong, 20 or more; stp with an extensive scale patch covering upper

corner, upper and lower posterior borders and base of prealar knob; psp with

a distinct broad scale patch; mep with anterior scale patch contiguous with

scale patch on upper corner of stp and several loosely packed scales among

upper mep bristles; 1 or 2 lower mep bristles present. Legs. Anterior sur-

face of forefemur with a complete longitudinal pale stripe or a linear row of

white spots from base to apex; anterior surface of midfemur with a median

longitudinal stripe along the whole length; anterior surface of hindfemur with

a broad longitudinal white stripe bordered above and below by dark scales; all

tibiae with anterior white stripes along whole length; tarsi of fore- and midlegs

usually entirely dark; tarsomere 1 of hindleg with distinct white stripe on anteri-

or surface, remaining tarsomeres entirely dark. Wing. Posterior surface of

vein C with pale scale line in basal 0.5; scales on all other veins entirely dark;

scales on Rg, Reg and R45 narrow, linear, elongate. Halter. Peduncle pale,

without scales; capitellum or knob dark scaled. Abdomen. Tergum I with

median scale patch which is completely or partially pale; terga II-VI with

broad basal pale bands which are usually strongly produced posteriorly as

yellow triangular patches; sometimes also with yellowish dots in middle;

tergum VIII entirely yellowish scaled; lateroterga II-VI with elongate basolat-

eral pale patches connecting with basal pale bands; sterna predominantly yel-

lowish or with sprinkling of some dark scales. Genitalia. Not studied.

MALE. Description based on specimens from India and South Africa. In

general as described for female. Head. Palpus. exceeding proboscis by full

length of segment 5; segment 3 with scattered pale scales in apical 0.95 and

ventral lateral rows or tuft of numerous strong bristles from basal 0.25 to

apex; segments 4 and 5 strongly plumose; dorsal and ventral surfaces with

distinct pale scale lines. Proboscis slightly pale on mid-ventral surface;

labium without median ventral tuft of long hairs.

MALE GENITALIA (Fig. 9). As described and figured by Barraud (1934:

415-6) with the following distinctive features. Segment IX, Tergal lobe with

irregular rows of 10-12 moderately strong setae. Basimere, Slender, conical,

about 0.3 mm in length. Subapical lobe. Setae a-c of proximal division sub-

equally long and stout; setae d-f of distal division consisting of 1 flattened,

apically blunt blade like seta and 1 narrower, apically pointed hairlike seta;

leaf g broad or narrow; seta h long and strong. Distimere. Normal: 1,2

Sirivanakarn: Subgenus Culex in the Oriental Region 45

ventral tiny setae present in apical 0.25; subapical claw slender. Phallosome.

Tergal arm of outer division slightly divergent laterad and distally tapered into

a point apically; lateral spine absent; sternal spines poorly developed or ab-

sent; lateral basal process slender, thumb-like; ventral arm of inner division

straight or slightly divergent laterad, with 3 strong apical teeth or spines.

Proctiger. Crown large and dark, consisting of numerous coarse and fine spicu-

les; basal sternal process well developed, about 0.05 mm in length; paraproct

narrow; cercal sclerite broad, well sclerotized; cercal setae 4,5 in number.

PUPA. Abdomen: 4.0mm. Paddle: 1.1mm. Trumpet: 0.91 mm;

index 7. Based on specimens from India and South Africa. Cephalothorax and

abdomen yellowish white. All setae developed, as figured for quinquefasciatus

(Fig. 2). Trumpet. Relatively long, cylindrical or gradually increased in

diameter distally; pinna weakly oblique, apical margin truncate. Cephalothor-

ax. Seta 1-C 3,4 branched; 2-C 4 branched; 3-C double; 4-C triple; 5-C 5,6

branched; 6-C 4 branched, as long as 4-C; 7-C double; 8-C 4-6 branched; 9-C

double. Metanotum. Seta 10-C 5,6 branched; 11-C usually double (1-3); 12-C

3,4 branched. Abdomen. Setae 6-I, II single; 7-I, II double; 1-II usually more

than 10 branched (8-14); 1-III, 7-9 branched; 1-IV 6, 7 branched; 1-V 4-6

branched; 1-VI-VII 3,4 branched; 5-IV usually triple (3-4); 5-V usually double

(2-3); 5-VI double; 6-III-VI usually 4,5 branched (3-6); 6-V, VI stronger and

longer than 6-III, IV; 9-VII 4-6 branched; 4-VIII single or double; 9-VIII 8

branched. Paddle; Large, broad and very pale; midrib weak and lightly pig-

mented; external buttress distinct, apical margin rounded, indistinct; seta 1-P

stronger than 2-P.

LARVA (Fig. 10). Head: 1.04 mm. Siphon: 1.8 mm; index 5. Saddle:

0.48 mm; siphon/saddle ratio 3.3. Description based on specimens from

India and South Africa. Head, Labrum broad; seta 1-C dark, spiniform;

4-C single, shorter than distance between bases of the pair; 5-C 3, 4 branched;

6-C usually double (2-3); 7-C 6,7 branched. Antennal shaft strong, and long,

distinctly curved outwards at middle, about 0.7 of head length; pigmentation

largely pale whitish except for basal dark ring and dark apical portion beyond

base of seta 1-A; 1-A strong, dark, multibranched; 2,3-A distinctly subapical.

Mental plate with 6 lateral teeth on each side of median tooth. Thorax. Setae

1-3-P single, strong, subequal; 4-P shorter than 1-3-P, double; 7-P triple;

8-P double; 1-M minute, triple, about 0.25 of length of 3-M; 3-M single. Ab-

domen. Ventral surface of segments I-VI with median patch of distinct spicules;

setae 6-I, II triple; 7-I double; 1-III-VI moderately strong, usually 4 branched

(3-4); 13-IJI-V as long as 1-III-VI, 4,5 branched; 6-III-VI all triple. Comb

scales 20-26, all similar insize, individual scale with a strong median apical

spine and lateral fringe of evenly fine spicules. Saddle pale whitish; seta 1-X

weak, triple; 2-X 3,4 branched; 4-X with 7 pairs of setae, all within grid; anal

gills slender, shorter or slightly longer than saddle. Siphon. Relatively thick

and very pale except for basal dark ring; distal portion slightly tapering; pecten

teeth strong, 12-15 in number, distal tooth with 1,2 basal denticles and an

elongate simple apical spine; siphonal tufts strong, inserted beyond pecten,

0-6 pairs in number; first 3-4 proximal pairs subequally long, 7-12 branched,

in a prominent row on ventral surface, their length about as long as siphonal

width at point of insertion; 2 distal pairs weaker, 8-10 branched, inserted

laterally or subventrally; 2-S short and slender; median caudal filament of

Spiracular apparatus absent.

TYPE-DATA. (1) Culex theileri Theobald, Lectotype ?*, Pretoria, Trans -

vaal, (SOUTH AFRICA), D. Theiler (BM; present selection); (2) Culex creticus

Theobald, Holotype °*, Crete, GREECE (BM); (8) Culex pettigrewii Theobald,

46 Contrib. Amer, Ent, Inst; ; vols 12, no. 2; 1976

Holotype °, Ukhrul, Manipur, Assam, INDIA, 6,400 ft., August 1908, Pettigrew

(IM); (4) Culex onderstepoortensis Theobald, Holotype ?*, Onderstepoort, Tvans-

vaal, (SOUTH AFRICA) (BM); (5) Culex theileri var. annulata Theobald, Holo-

type 2. Onderstepoort, Transvaal, (SOUTH AFRICA) (BM).

DISTRIBUTION. Widely disjunct, covering the Ethiopian, Mediterranean

and Middle East with extension eastward into the Oriental region as far as India,

Burma and southern China and northward into USSR. Material examined 140

specimens: 51%, 692, 20 L; 7 individual rearings (5 pupal, 2 larval).

CHINA. Yunnan, Kunming; 58; locality unknown; 2 L.

INDIA. Western Himalayas, Kasauli; Kashmir; 10°, 19, 1p.

NEPAL. Katmandu; 1 L.

PAKISTAN. Montgomery: 2 L.

AFGHANISTAN. Kunduz Prov.; Bamiyan Prov. ; Herat Prov. ; 230, 269.

IRAQ. Abu Ghraib; Shubaicha; Duquq; Rawanduz, Nahia; Erbil, Towne; 2c,

LY hs.

IRAN. Sharaf Khaneh; Semnan; Maragneh; Do Rud; 50’, 49.

SPAIN. Canary Is., Tenerife, Santa Cruz; 1% (genitalia).

PALESTINE (ISRAEL). 1c, 19. |

TUNISIA. Mateur; Bizerte; 10, 209.

YEMEN. El-Amra, nr. Ta'iss; 20, 2 lp.

GREECE. Macedonia; 1°.

ALGERIA. Oued Taria; Mocta Douz; La Mocta; Le Kraub; 20, 29.

KENYA. 2°, 29 (as tipuliformis Theobald).

SOUTH AFRICA. Johannesburg; 2“, 72, 4 L; 4 p (from mass rearing).

Additional records from the literature: INDIA: Punjab; WEST PAKISTAN

(Baluchistan); BURMA (Barraud 1934: 416); SOUTH AFRICA: Cape; Natal;

Orange Free State; Transvaal; SOUTH WEST AFRICA; SOUTHERN RHODESIA;

ABYSSINIA; ETHIOPIA; ERITREA; CONGO; TANGANYIKA; S. W. ARABIA;

MOROCCO; EGYPT; ITALY: Sardinia; FRANCE: Corsica; SPAIN; PORTUGAL

(Edwards 1941: 306; Aitken 1954: 488; Mattingly and Knight 1956: 121; Senevet

and Andarelli 1959: 201); USSR: Ukraine; south shores of Crimea; Northern

Caucasus; Transcaucasia, Central Asia (Monchadskii 1951: 275).

TAXONOMIC DISCUSSION. The new record of theileri from China is based

on 5 females which were found among several guinquefasciatus Specimens in the

USNM collection by Cazier from Kunming, Yunnan in 1945. In the previous

studies by Barraud (1924b, 1934), this species has been reported only from the

north of India and Burma. It is quite possible that it may also extend its range

into other parts of Southeast Asia south of the Chinese border.

Culex theileri is among the most clearly marked species of the Pipiens

Group. It can be readily separated from other Oriental species in the Pipzens

Group in the adults by (1) large size (wing length usually 5.0 mm or more);

(2) presence of a broad scale patch on postspiracular area; (3) presence of an

extensive continuous scale patch on sternopleuron and base of prealar knob;

(4) presence of very striking longitudinal pale stripes on the femora and tibiae

of all legs and (5) basal bands of abdominal tergites usually broad and strongly

produced posteriorly as triangular yellowish patches. The male genitalia of

theileri are very constant in practically all characters noted and can be dis-

tinguished from other Oriental species most readily by the presence of 3-4

strong lateral teeth on the apex of ventral arm (inner division) of the phallosome.

In the immature stages, the pupa can be separated by (1) large size; (2) rela-

tively long and uniformly thick trumpet; (3) pinna of trumpet weakly oblique;

(4) cephalothoracic seta 8-C 4-6 branched and (5) abdominal setae 6-III-VI

usually 4,5 branched. The larva is differentiated by (1) seta 1-C of head dark,

Sirivanakarn: Subgenus Culex in the Oriental Region 47

stout and spiniform (2) comb scale apically pointed or with a strong median

apical spine; (3) siphon relatively thick, long and distally slightly tapered; (4)

presence of 5 or 6 pairs of moderately long siphonal tufts, 3 or 4 proximal

pairs of which are closely spaced and forming prominent rows on ventral sur-

face and (5) ventral brush (4-X) of saddle with 7 pairs of setae.

Among the Ethiopian species, the male phallosome of ¢hezleri resembles

those of andersoni Edwards 1914 and simpsoni Theobald 1905 as described and

figured by Edwards (1941: 305-6), indicating that they are closely related.

BIONOMICS. According to Barraud (1924b, 1934), theilevi was reported to

be common at a very high elevation, ranging from 1, 000-3, 000 m in the Himal-

ayan areas of North India where it was found to utilize large ground pools,

stream pools and marshes as breeding habitats. Elsewhere in the Ethiopian,

Mediterranean and in several countries to the west of India, it was reported

from a broad range of elevations and was found breeding in pools, dipping tanks,

stagnant or slowly moving streams, ditches, drains, backwaters of rivers,

Swamps and ponds (Hopkins 1952: 291, Aitken 1954: 489, Senevet and Andarelli

1959: 201, Lofti 1970: 402). In northern Afghanistan, it was reported to be

locally common and the immatures were frequently collected from polluted

eround pools (R. A. Ward, personal communication). The South African mat-

erial collected for this study all came from rearing pupae and larvae in ground

pools. Nothing is known about adult biology, feeding habit and medical impor-

tance of theilevi in India and adjacent countries to the west. In South Africa,

the females feed mainly on blood of cattle and occasionally on man and birds

(Anderson 1967). In virus isolation studies by McIntosh et al. (1967) in South

Africa, theileri was found to be naturally infected with Sindbis and West Nile

viruses. Jupp et al. (1966) have demonstrated in the laboratory that it is an

efficient vector of Sindbis virus but not of West Nile virus.

UNIVITTATUS SUBGROUP

The Univittatus Subgroup is characterized by the following male genitalic

characters: (1) setae in group d-f of distal division of subapical lobe consisting

of 1 long bladelike seta and 1 shorter hairlike seta, situated on accessory mesal

lobe on sternal surface of basimere; (2) subapical dorsal surface of distimere

with distinct annulation or serration; (3) ventral arm of inner division of phal-

losome simple or composed of 5,6 strong and long fingerlike processes; (4)

outer division without lateral and sternal spines; (5) tergal arm of outer divi-

sion apically pointed or truncate and (6) basal sternal process of proctiger

dark, heavy and long.

The Univitiatus Subgroup as proposed here includes univittatus Theobald

1901 and fuscocephala Theobald 1907. This subgroup is strongly differentiated

from other subgroups in the male genitalia by the development of accessory

mesal lobe in the distal division of subapical lobe of the basimere and in the

presence of distinct annulation or serration on subapical dorsal surface of the

distimere. It may also include several other Ethiopian forms, especially those

in the decens series of Edwards (1932, 1941), but no attempt is now made to

determine their affinity.

Culex univittatus was originally described from the Ethiopian region and has

subsequently been reported from several localities in the Mediterranean,

Middle East and eastward as far as India (Barraud 1934) in the Oriental region.

It is probably a complex of more than one form on the basis of the studies by

Jupp (1971, 1972) in South Africa. Culex fuscocephala is exclusively Oriental

48 Contrib. Amer, Ent. ‘Inst, vol 12, no, 21976

and is one of the commonest species in Southeast Asia. It shows a striking

difference from univitiatus in the ventral arm of inner division of the phallo-

some but is very similar to the latter in all other details of the male genitalia.

a MCULIN (CULEX) UNIVITTAT US THEOBALD

(Figs. 9, 10)

Culex univittatus Theobald 1901b: 29 (*, 2*); Edwards 1913a: 58 (co, @).

Culex perexiguus Theobald 1903: 199 (“, 9); Edwards 1921: 342 (o%, 9, L*);

_ Edwards 1922: 471 (distribution); Barraud 1924b: 1263 (o*, 2); Barraud

1924c: 431 (L*); Kirkpatrick 1925: 125 (o*, 2, P*, L*); Edwards 1932:

210 (synonymy).

Heptaphlebomyia simplex Theobald 1903: 337 (2*); Edwards 1932: 210 (synonymy).

Culex pallidocephala Theobald 1904: 73 (o); Edwards 1941: 306 (synonymy).

Heptaphlebomyia montforti Ventrillon 1905: 448 (o'*, 2); Edwards 1932: 210

(synonymy).

Culex goughii Theobald 1911a: 268 (2*); Edwards 1912: 32 (2, synonymy).

Culex (Culex) univittatus Theobald, Edwards 1932: 210 (taxonomy); Barraud

1934: 418 (o*, 2, L*); Edwards 1941: 301 (*, 2, P); Hopkins 1952: 291

(L*); Mattingly and Knight 1956: 104 (taxonomy); Senevet and Andarelli

1959¢,208.(¢*, 94.7); Jupp 1071: 340: (o%, OF, -L)5 Jupp 1972: 103

(*, 2, taxonomy).

FEMALE (Fig. 9). Wing: 3.0 mm. Forefemur: 1.44 mm. Proboscis:

1.6mm. Small to medium sized species; distinguished from other Oriental

members of the Pipiens Group by the presence of a scale patch on postspiracu-

lar (psp) area and by the presence of longitudinal pale stripe on midfemur, mid-

and hindtibiae. Head. Narrow decumbent scales on vertex pale beige to whit-

ish; erect scales moderately long, largely pale in center, dark on postero-

lateral areas; sometimes entirely dark; lateral patch of broad scales whitish,

very distinct. Palpus with pale scales forming streak on segments 3 and 4,

rest dark. Proboscis dark dorsally, slightly pale on mid-ventral surface.

Cibarial Armature. Not studied. Thorax. Mesonotal integument brown; mes-

onotal scales largely yellowish brown, pale beige or whitish on marginal area,

prescutellar space and scutellar lobes. Pronotum (apn, ppn) with similar

scales as mesonotum. Pleuron same color as mesonotum without distinct

pattern of dark and light bands; scale patches present on psp, ppl, sip and mep;

scale patches on upper corner and lower posterior border of stp extensive but not

contiguous; upper one extended dorsad to base of prealar knob; 1 lower mep

bristle present. Legs. Scales on anterior surface of forecoxa largely pale;

anterior surface of forefemur entirely dark; anterior surface of midfemur

usually with complete median longitudinal pale stripe which is sometimes broken

up into a chain of pale spots; anterior surface of hindfemur largely pale in basal

0.5, continued as narrow pale stripe to near apex; apices of all femora with

distinct apical white spots; foretibia entirely dark anteriorly; mid- and hind-

tibiae with longitudinal pale stripes on anterior surfaces; all tarsi entirely

dark. Wing. All scales dark except for pale scale line on posterior surface

in basal 0.5 of vein C and a few scattered pale scales on Sc and R; scales on

veins Ro, R, and R45 narrow. Abdomen. Terga II-VII with even basal pale

bands and basolateral pale spots; sterna largely pale yellowish. Genitalia. Not

studied.

MALE. In general as described for female. Head. Palpus slender,

Sirivanakarn: Subgenus Culex in the Oriental Region 49

exceeding proboscis by full length of segment 5; segment 3 with ventrolateral

tuft of about 10 bristles in apical 0.5; segments 4 and 5 strongly plumose, es-

sentially as in quinquefasciatus. Proboscis without median ventral tuft of long

hairs. Legs. Anterior surface of midfemur without distinct longitudinal pale

stripe. Abdomen. Basal bands on terga II-VII broader than in female.

MALE GENITALIA (Fig. 9). Essentially similar to fuscocephala in the de-

tails of basimere, subapical lobe, distimere and proctiger, differing particu-

larly in the following: Phallosome. Inner division of lateral plate represented

by a broad, simple leaflike ventral arm which is strongly bent or angled proxi-

mally; its innermost sternal margin provided with a slender and sharp spine of

variable length; outer division with a strong tergal arm which is distally tap-

ered into a sharp point; lateral basal process very slender and elongate.

PUPA. As figured for fuscocephala (Fig. 12) to which it is extremely

Similar in the shape and relative length of trumpet and in most features of

chaetotaxy, differing particularly as follows: Abdomen. Hair 5-V triple; 5-VI

usually triple, sometimes double; 5-VII triple.

LARVA (Fig. 10). Head: 0.8 mm. Siphon: 1.4 mm; index 6. Saddle:

0.35 mm; siphon/saddle ratio 4. In general similar to fuscocephala, differing

particularly in the following features... Head. Head seta 1-C darker, stouter,

more or less spiniform. Antenna longer; spicules on antennal shaft stronger.

Abdomen. Setae 1-III-VI shorter, double or triple; 6-III double; 6-IV triple;

6-V double; 6-VI single. Seta 1-X of saddle triple. Siphon. Siphonal tufts 5

pairs in number, 2 proximal and 1 subapical pairs inserted subventrally, other

2 pairs in apical 0.9 inserted laterally; all subequal, 3,4 branched each and

about as long as or slightly shorter than siphonal width at point of attachment.

TYPE-DATA. (1) Culex univitiatus Theobald, Lectotype o*, Salisbury,

Mashonaland, S. RHODESIA, March 1900, G. A. K. Marshall (BM; selection

of White 1975: 320). (2) Culex perexiguus Theobald, Lectotype &, Sidon, Pale-

stine (LEBANON) (BM; selection of White 1975: 318). (3) Heptaphlebomyia

simplex Theobald, Holotype 2, Salisbury, Mashonaland, S. RHODESIA (BM).

(4) Culex pallidocephala Theobald, Lectotype “, Sennar, Blue Nile, SUDAN

(BM; selection of Edwards 1941: 306). (5) Heptaphlebomyia montforti Ventril-

lon, Lectotype “, Ankajobe, MADAGASCAR (BM; selection of White 1975: 321).

(6) Culex goughii Theobald, Lectotype “, Onderstepoort, Tvansvaal, (South

AFRICA) (BM; selection of White 1975: 321).

DISTRIBUTION. Widespread, covering the Ethiopian (southern and eastern

Africa), Mediterranean, Middle East, Iraq, Iran, Afghanistan, Pakistan with

a small extention into the Oriental region as far as Punjab, India.

Material examined: 250°, 319, 17 L; 17 from mass rearing (17 1, 17 p); 1

individual rearing (1 larval).

INDIA. Punjab: Amritsar, 19, 1 lp.

PAKISTAN. Peshawar, 1.

AFGHANISTAN. Helman Province: Garwagin Village; 80, 109.

TUNISIA © 80,/°49..

ISRAB Lh Ae pay,

ALGERIA. 1¢, 32.

SOUTH AFRICA. Johannesburg; 11°, 139, 17 L, 17 p, 171.

Additional records from the literature: INDIA: United Province; PAKISTAN

(Baluchistan, Sind, N. W. Frontier): Lahore (Barraud 1934: 420); IRAN: Kaz-

eroon; Bandar-Abbas (Lofti 1970: 402); SOUTH AFRICA: Cape; Transvaal;

Orange Free State; SOUTH WEST AFRICA; PORTUGUESE EAST AFRICA; RHO-

DESIA; NYASALAND; ANGOLA; ZANZIBAR; MAURITIUS; MADAGASCAR; KENYA;

UGANDA; SUDAN; GOLD COAST; NIGERIA; GAMBIA; CONGO; FRANCE; SPAIN;

0 Contribs Amer; Baty Inst.,; volvh2).nos2,.1976

ITALY; GREECE; EGYPT; OMAN (Edwards 1941: 308; Senevet and Andarelli

1959: 214; Mattingly and Knight 1956: 122); USSR: Turkmenia; SYRIA; LEBANON

(Monchadskii 1951: 279).

TAXONOMIC DISCUSSION. The assignment of the Oriental specimens

described above to univittatus is provisional, based on the study of the limited

material previously described by Barraud (1934: 418-20) from India and its

comparison with the types and specimens from South Africa. Except for the

smaller size, the Indian form of wivittatus does not appear to show any signi-

ficant difference in the male genitalia and in the pupal and larval chaetotaxy

from the type or from specimens from the Middle East, the Mediterranean and

the Ethiopian area (particularly South Africa). Although 2 distinct taxa, univit-

tatus (s. str.) and neavei Theobald 1906 have been recognized by Jupp (1971) in

South Africa on the basis of the male genitalia and external characters of the

adults, I have not been able to determine which of the 2 South African forms the

Indian specimens should belong to. It appears most probable from the studies

by Jupp (1971, 1972), that this species is highly complex and that it may con-

sist of more than 2 taxa. The Indian specimens as well as those from other

adjacent areas in Pakistan and Afghanistan appear to be conspecific and probably

represent a distinct species. Because of the limited reared material, the exact

status of this Oriental form can not be determined at present.

Culex univittatus in the Oriental part of its range is a relatively slender,

small or medium-sized species. It can be readily separated from other Ori-

ental forms of the Pipiens Group by the following combination of characters:

In the adults by (1) presence of scale patch on the postspiracular area; (2) scale

patches on the upper corner and lower posterior border of sternopleuron not

contiguous and (3) presence of longitudinal pale stripes on the anterior surface

of midfemur and tibiae of mid- and hindlegs; the male genitalia by (1) the ven-

tral arm of inner division of the phallosome broad, leaflike and with a distinct

spine on the mesal surface; (2) tergal arm of outer division tapered into a point

apically; (3) lateral basal process of the outer division slender and elongate;

(4) basal sternal process of the proctiger dark, thick and long; (5) setae d-f of

distal division of the subapical lobe 2 in number and situated on a distinct mesal

lobe on the sternal surface and (6) subapical dorsal surface of the distimere

distinctly annulated or serrated; the pupa by (1) the slender and uniformly

cylindrical trumpet; (2) seta 8-C 4-6 branched; (3) abdominal setae 6-III-VI

4,5 branched; (4) seta 5-IV 5 branched and (5) seta 5-V triple; and the larva by

(1) the dark and spiniform seta 1-C of head; (2) siphon pale, slender and rela-

tively long and (3) siphonal tufts 5 pairs in number, widely spaced and about as

long as siphonal width at the point of attachment.

BIONOMICS. Culex univitiatus is a typical ground pool breeder. In India,

it was reported to breed frequently in marshy pools, borrow pits, stagnant

drains and canals, shallow wells and less frequently in domestic collections of

water and rice fields (Barraud 1934: 420). Inthe Ethiopian region, the larvae

of univittatus were reported to be common in marshy pools and in pools at the

edge of swamps and were also found in stagnant or semi-stagnant streams and

ditches, borrow pits, etc. (Hopkins 1952: 293). Nothing is known about the

adult biology and medical importance of univitiatus in India and other adjacent

countries to the west. In South Africa, it was reported to feed on birds and

mammals exclusive of man (Anderson 1967) and has been considered to be an

important vector of Sindbis and West Nile viruses (McIntosh et al. 1967, Jupp

and McIntosh 1967).

Sirivanakarn: Subgenus Culex in the Oriental Region OL

6. CULEX (CULEX) FUSCOCEPHALA THEOBALD

(Pies. 14°12) 213314)

Culex fuscocephala Theobald 1907; 420 (2); Theobald 1910a: 363 (distribution);

Edwards 1913b: 234 (taxonomy); Barraud 1924b: 1270 (o*, 2); Barraud

1924c: 432 (L*); Borel 1930: 342 (*, 2, L*).

Culex uniformis Leicester 1908: 159 (2); Edwards.1917: 225 (synonymy).

Culex minimus Leicester 1908: 160 (“, 2); Edwards 1932: 212 (synonymy).

Culex taytayensis Banks 1909: 545 (o, °); Edwards 1913b: 234 (synonymy).

Culex luteola Theobald 1910a: 378 (¢*); Edwards 1913b: 234 (synonymy).

Culex inelegans Dyar 1920: 179 (); Barraud 1934: 424 (synonymy).

Culex fuscitarsis Barraud 1924b: 1272 (“*, 2); Bram 1967a: 184 (synonymy).

Culex (Culex) fuscocephalus Theobald, Edwards 1932: 212 (taxonomy); Barraud

1934: 424 (o*, 2, L*); Bonne-Wepster and Brug 1937: 8 (o, °*); Feng

1938: 303 (biology, distribution); Bonne-Wepster and Brug 1939: 1277

(L*); Bohart 1945: 78 (o*, L); Bonne-Wepster 1954: 133 (%, 2*, L*);

Hsieh and Liao 1956: 123 (distribution); Chu 1958: 110 (distribution);

Lien 1962: 633 (distribution); Safyanova et al. 1964: 1177 (distribution).

Culex (Culex) fuscitarsis Barraud, Edwards 1932: 212 (taxonomy); Barraud

1934: 426 (o'*, 9), |

Culex (Culex) fuscocephala. Theobald, Delfinado 1966: 143 (*, 2, L); Bram

1967a: 184 ('*, 9*, P*, L*); Baisas 1974: 90-(¢, 2, .P, L),

FEMALE (Fig. 11, 14). Wing: 2.4-3.4 mm (average 2.8 mm). Forefemur:

1.1-1.5 mm (average 1.4 mm). Proboscis: 1.4-2.0 mm (average 1.7 mm).

Abdomen: 2.2 mm. Small species, easily distinguished from other members

of the Pzpiens Group by the presence of a striking pattern of dark and light

bands on upper sternopleuron (sip) and mesepimeron (mep) and by the absence of

abdominal tergal pale bands. Head. Narrow decumbent scales of vertex very

fine and yellowish; erect scales short, predominantly dark; lateral patch of

broad scales whitish, very distinct. Palpus very thin and rather short, about

0.15 of proboscis length, entirely dark scaled. Proboscis dark scaled on dor-

sal surface, paler on lateral and ventral surfaces, but not extended dorsad to

form a distinct ring. Cibarial Armature. Cibarial bar small; cibarial teeth

very fine, long and distally filamentous, all-subequal in length, about 40 in

number. Thorax. Mesonotal integument brown; mesonotal scales very fine

and dense, vestiture smooth, predominantly yellowish or brownish in middle of

mesonotal disc, pale beige to whitish on marginal areas, prescutellar space

and scutellar lobes. Apn with narrow, yellowish scales and weak setae; ppn

with 5 posterior bristles and several narrow pale scales scattering on upper

surface. Pleuron with characteristic dark bands above and below whitish

scale patches on upper corner of stf and upper mep; ppl with small whitish

scale patch and 6,7 bristles; a few pale scales present among upper mep

bristles; 1 lower mep bristle present. Legs. Forecoxa with anterior scale

patch which is dark above, pale on lower surface; anterior surface of fore-

and midfemora entirely dark scaled; hindfemur largely pale in basal portion,

continued as pale stripe in distal portion; all tibiae and tarsi dark scaled.

Wing. All scales dark; scales on veins Ro, Rg and Ry,5 narrow. Abdomen.

Terga II-VIII entirely grayish to brownish, basal bands absent; scales at bases

of terga occasionally lighter than other areas, but not forming distinct basal

bands; basolateral pale spots absent; sterna largely yellowish white. Genitalia.

Tergum IX with lateral row of 5-9 setae; postgenital plate with distinct median

52 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

emargination on caudal margin; vaginal sclerite U-shaped and well sclerotized;

insula with a tuft of 6-12 setae.

MALE (Fig. 11). In general as described for female except for the following

sexual differences. Head. Palpus thin, exceeding proboscis by 1.5 of length of

segment 5; segment 3 entirely dark or grayish; apical 0.25 with ventrolateral

tuft of 15-20 dark bristles; segments 4 and 5 upturned, strongly plumose and

entirely dark scaled on dorsal surface; segment 4 with white scale line on ven-

tral surface; segment 5 without any scale on ventral surface. Proboscis with

distinct median ventral tuft of 2-4 long and 2, 3 shorter setae.

MALE GENITALIA (Fig. 12). Segment VIII. Tergum shallowly emarginate

on median caudal margin and with distinct row of 6-8 dark kinked setae. Seg-

ment IX, Lateral tergal lobe broadly round, with a row of 8-10 moderately

strong setae. Basimere. Small, slender, conical. Subapical lobe. Seta a of

proximal division shorter than setae b-c; setae d-f of distal division consists of

1 long, flattened bladelike seta and 1 strong hairlike seta which are situated on

distinct mesal lobe on sternal surface; leaf g very large, heart-shaped, placed

slightly distad of setaea-c; seta h strong and long. Distimere. Sickle-shaped,

basal 0. 50-0. 75 straight, apical 0.25-0.50 gently or strongly curved, with dis-

tinct annulation or crest of weak spicules on dorsal subapical surface; 1 ventral

tiny seta present subapically; dorsal seta absent; subapical claw slender and

moderately long. Phallosome. Inner division of lateral plate with 5-6 heavy

toothlike processes, most sternal longest, lateral ones shorter, all divergent

laterad; outer division with a strong divergent tergal arm which is uniform in

width and apically truncate; lateral basal process strongly expanded as broad

leaf, apex bluntly pointed or rounded. Proctiger. Crown large; paraproct and

cercal sclerite broad, heavily pigmented and strongly sclerotized; basal sternal

process very thick, dark and long, curved sternad; cercal setae 2 in number.

PUPA (Fig. 12). Abdomen: 2.5mm. Paddle: 0.65 mm. Trumpet: 0.52

mm; index about 6. Cephalothorax and abdomen cream-colored or yellowish

white. Complete chaetotaxy as figured; all setae developed, the following are

distinctive. Trumpet. Dark, uniformly cylindrical; pinna weakly oblique,

apical margin truncate. Cephalothorax. Seta 1-C usually 4 branched (3-4);

8-C usually 5 branched (3-6). Abdomen. Seta 1-II 20 or more branched; 5-IV

4-6 branched; 5-V, VI double; 6-III-VI usually 4,5 branched (2-5); 9-VII 4, 5

branched (3-6); 5-VII double; 4-VIII double or triple; 9- VIII 8-10 branched.

Paddle, Very pale to almost transparent; midrib weak and lightly pigmented;

external buttress distinct; apical external margin indistinct; setae 1, 2-P pre-

sent, minute.

LARVA (Fig. 13). Head: 0.65 mm. Siphon: 1.2-1.4 mm; index 5-6.

Saddle: 0.32 mm; siphon/saddle ratio 4-5. Complete chaetotaxy as figured.

Head. Seta 1-C filiform and pale; 4-C single, 1-2 times as long as distance

between bases of the pair; 5, 6-C usually double (2-3); 7-C 6,7 branched. An-

tenna about 0.7 of head length; middle proximal portion pale whitish or yellow,

basal and apical portions dark; spicules numerous and distinct; seta 1-A dark,

about 20 branched; 2, 3-A subapical. Mental plate with 6,7 lateral teeth on each

side of median tooth. Thorax, Seta 3-P shorter or as long as 1,2-P; 4-Pas

long as 3-P, double; 7-P triple; 8-P double, 1-M short, tiny, double; 3-M

single; 4-M double; 8-M 5 branched; 9-M 4 branched; 1-T short, 3,4 branched;

7-T 6,7 branched; 9-T 5 branched; 12-T single or double; 13-T 5-8 branched.

Abdomen. Setae 1-I, II 2-4 branched; 6-I, II triple; 7-I double; 1-III-VI single

and very long, same magnitude as 6-III-VI; 6-III, V and VI single; 6-IV 1-4

branched; 13-III-V 4-6 branched. Comb scales small, all similar in size,

about 40, individual scales with rounded even fringe of fine spicules. Saddle

Sirivanakarn: Subgenus Culex in the Oriental Region oP)

same color as siphon; posterior apical margin lightly spiculated; setae 1, 2-X

double; 4-X with 6 pairs of setae; anal gill 1-2.5 times as long as saddle.

Siphon. Slender, moderately long and pale yellowish except for basal dark ring;

pecten teeth 11-12, in close-set row; distal teeth with 2-4 basal denticles and a

fine apical denticle; siphonal tufts 3-4 pairs in number, all weak, short and sub-

equal, about 0.5 of siphonal width at point of insertion; each double or triple; 1

or 2 proximal pairs subventral; 1 median pair lateral, 1 distal pair subventral;

seta 2-S weak and short; median caudal filament well developed.

TYPE-DATA. (1) Culex fuscocephala Theobald, Holotype 2*, Peradeniya,

CEYLON (SRI LANKA), 1902, Green (BM); (2) Culex uniformis Leicester, Type

(lost), Batu Gajah (Perak), MALAYA (PENINSULAR MALAYSIA), G. F.

Leicester (BM); (3) Culex minimus Leicester, Type (lost), Kuala Lumpur

(Selangor), MALAYA (PENINSULAR MALAYSIA. ), G. F. Leicester (BM);

(4) Culex taytayensis Banks, Holotype ?*, Tay Tay (Luzon), PHILIPPINES (BM);

(5) Culex luteola Theobald, Holotype 9*, Peradeniya, CEYLON (SRI LANKA),

1900, Green (BM); (6) Culex inelegans Dyar, Holotype * with slide of genitalia,

Los Banos (Luzon), PHILIPPINES, June 1915, Baker (USNM); (7) Culex fusci-

larsis Barraud, Lectotype “* (No. 1701) with slide of genitalia, Pachmari,

Central Provinces, INDIA, July 1915, V. H. Dowson (BM; selection of Bram

1967a: 188).

DISTRIBUTION. Exclusively Oriental; widespread throughout Southeast

Asia with extension into the west as far as India and Sri Lanka; to the north in

South China; to the northeast as far as Okinawa, to the east in the Philippines,

and to the south in Java, Sulawesi, Kalimantan (Borneo) and Sumatra. Materi-

al examined. 3,793 specimens: 1,1450°, 1,8099, 839 L; 1,072 individual

rearings (376 pupal, 696 larval).

INDIA. Assam: Dooma Dooma; Chabua; Rupsi; Jorhat (D. E. Hardy,

1943); Bengal: Lalmanirhat; Calcutta, Dum Dum, N. Salt Lake; Darjeeling,

Sukna; Old Jalpaiguri; Bihar: Purnea; Andaman Is.; Central Prov.: Pachmar-

hi (as fuscitarsis Barraud 1924); 540°, 642, 45 L, 8p, 6 lp.

BANGLADESH. Sylhet; 30, 139, 6p.

SRI LANKA. Central Province: Peradeniya, Kandy District (type locality);

Northern Province: Vavuniya District; Sabaragamuwa Province: Ratnapura

District, Uggalkaltota; Southern Province: Hambantota District, Palatupana;

Lic: (159;

BURMA. Rangoon; Myo Kwet Thit; 60’, 122, 6p, 1 lp.

THAILAND. Chiang Mai; Chiang Rai; Mae Hong Son; Lamphun; Lampang;

Non; Khon Kaen; Prae; Udon Thani; Nakhon Ratchasima; Nakhon Nayok; Sara

Buri; Kanchanaburi; Ayatthaya; Bangkok; Thon Buri; Pathum Thani; Chon Buri;

Chanthaburi; Trat; Rangong; Krabi; Phuket; Surat Thani (Koh Samui); Nakhon Si

Thammarat; Narathiwat; 4240, 7892, 533 L, 250 p, 268 lp.

VIETNAM. Saigon; Danang; Nha Trang; Gia Dinh; Phu loi; Phu Bei; Dia

Loe Run; 100; S72, OR vbiy: 4p,

CAMBODIA. Ari Gsartr; Phnom Penh; 6c, 129.

LAOS. Muang Sing; 18.

MALAYSIA. Peninsular Malaysia (Malaya): Selangor: Puchong Rd. ;

Kepong; Sentul; Ulu Klang; Segambut; Serdang; Pahang: Bukit Pakoh; Kuala

Lipis; Bentong Rd. ; Kuantan; Sungai Lembing; Trengganu: Marang; Perak:

Kuala Kangsar; Taiping; Tapah; C. Highlands Rd.; Perlis: Kg. Sg. Bharu;

Padang Besar; Kg. Prok Buah; Kg. To Kayaman; Kg. Baru; Kubang Tiga; Ke-

dah: Changlun; Kg. Peng Besar; Bt. Kayu Hitam; Kg. Bagan; N. S. (Negeri

Sembilan): Kg. Parit Tinggi; Malacca; Malaysia (N. Borneo): Sabah: Kota

Kinabalu (Jesselton); Papar; Tuaran; Beaufort; 1660", 2049, 73 L, 40 p, 132 lp.

o4 Contrib..Amer. Ente Inst., vol. 12) now 2; 1976

SINGAPORE. 1¢, 29.

INDONESIA. Java: Dijakarta; Bogor; Sumatra: Bengkulu; 100, 169.

PHILIPPINES. Luzon: Quezon; Subic Naval Base; Olongapo; Laguna;

Dalton, N. Vizcaya; Camarines Norte; Batangas; Calaccad; Trinidad Mtn.

Prov.; Mt. McKinley; Busunga; Calatagan; Kolambugan; Rizal, Wack Wack,

Alabang; Los Banos; Pangasinan; Agoo La Union; San Fernando; Camp Stotsen-

berg; Camp Nichols; Mindoro: San Jose; Palawan: Quezon; Iwahig; Samar:

San Antonio; Osmena; Leyte: Gulf; Abuyog; Lago Lago; Dulag; Palo; Carigara;

Tacloban; Mahaplag; Negros: Dumaguete; Mindanao: Ori Missamis; Dansalan;

Kabakan; Parang; Ludlow Barracks; Davao; Zamboanga; Mercedes; Pasanonco;

Tawi Tawi Is.: Tarawakan; Kamagong; Basbas Is.; Isabela: Basilan; 426,

6042, <136: 1, °62-p,° 257 Ip.

TAIWAN. Loh Yei; Ping Tung; Sze Chun; 7“, 1492, 19 L, 6 lp.

HONG KONG, 3¢, 22, 2 L.

CHINA.* Canton; 3, 119.

JAPAN. Ryukyus: Iriomote; 130, 139, 26 lp.

Additional records from the literature: INDIA. Andaman Is.; Punjab

(Barraud 1934); NEPAL (Peters and Dewar 1956); PAKISTAN (Aslamkhan

1971); INDONESIA: Sumba; Flores; Timor (Bonne-Wepster 1954); CHINA:

Fukien; Kiangsu (Feng 1938); Amoy (Hsieh and Liao 1956); Hainan (Chu 1958).

TAXONOMIC DISCUSSION. Culex fuscocephala is one of the most common

and widespread Southeast Asian species. The adults can be readily recognized

by the slender build, relatively small size, pleural markings and in most spe-

cimens by the absence of basal abdominal bands. All of these features are very

uniform except for the abdominal terga which occasionally exhibit very indis-

tinct basal pale bands on the posterior segments. The Indian form, which was

described by Barraud (1924b: 1272) as fuscitarsis on the basis of the presence

of basal pale bands on the abdominal terga, apparently falls into the range of

variation of fuscocephala populations from northern Thailand. Neither the

Indian fuscitarsis Specimens nor those from a few collections in northern

Thailand show any differences in other adult features, the male genitalia and

in the immature stages from typical fuscocephala specimens in several South-

east Asian areas. On this basis, I am convinced that they are conspecific with

the typical fuscocephala. In addition to the conspicuous diagnostic characters

of the adults, fuscocephala is also strongly differentiated from other species of

Culex (Culex) in the female cibarial armature, the presence of a median ven-

tral tuft in the male proboscis, the type of the male phallosome and in several

features of the immature stages.

BIONOMICS. Culex fuscocephala is locally abundant, especially in open

rural areas such as rice fields. The immature stages have been most frequent-

ly collected from general ground pools, ditches, ponds, wells, rockpools, pud-

dles, foot prints and marshy depressions. They have also been taken from con-

tainer habitats such as earthenware jars and coconut shells. Both larvae and

pupae are very abundant almost throughout the year and have been frequently

found in association with those of C. tritaeniorhynchus, C. vishnui, C. pseudo-

vishnui; Anopheles vagus Donitz, An. kochi Donitz, An. subpictus Grassi and

An. nigerrimus Giles and occasionally with C. quinquefasciatus and C. gelidus.

The adults are also abundant and have been frequently collected outdoors among

herds of domestic animals such as cows, water buffalo and pigs. Numerous

adults have also been caught in light traps and animal baited traps. The females

prefer to feed on blood of cows and pigs (Colless 1959; Reuben 1971c); but may

also attack man in the absence of the preferred hosts.

MEDICAL IMPORTANCE. Culex fuscocephala is one of the common pests

Sirivanakarn: Subgenus Culex in the Oriental Region 5D

of domestic animals such as pigs and cows in rural communities of Southeast

Asia. It has been reported to be naturally infected with Brugia malayi (Brug)

by Carter (1948) in Sri Lanka and with Wuchereria bancrofti by Rozeboom and

Cabrera (1964) in Luzon, the Philippines, but whether or not it may play a role

in the transmission of these pathogens remains to be determined. In Thailand,

fuscocephala has been reported to be an efficient vector of Japanese encephalitis

(Muangman et al. 1972) and has been found to be naturally infected with 2 strains

of this virus (Gould et al. 1974).

SITIENS GROUP

FEMALE. Head, Proboscis with distinct median pale ring. Cibarial

Armature. As described for the subgenus, apparently without distinctive

eroup character. Thorax. Color of mesonotal scales varied from predomin-

antly pale whitish to dark brownish on anterior 0.7 to level of anterior wing

base; scales on supraalar and posterior dorsocentral areas usually dark;

scales on prescutellar space and scutellar lobes partially or entirely pale,

sometimes entirely dark. Pleural scale patches usually present on ppl, upper

corner and lower posterior border of stp, poorly developed or absent on mep;

some scattered scales present among upper mep bristles; lower mep bristle

absent. Legs. Anterior surface of fore- and midfemora usually entirely dark,

sometimes with speckling of pale scales; anterior surface of hindfemur usually

with pale stripe extending from base to about 0.75 of total length, sometimes

dark with speckling of pale scales as on fore- and mid femora; tarsomeres

1-4 of all legs with distinct basal and apical pale bands. Wing, All veins usu-

ally with dark scales only, or sometimes speckled, with a mixture of dark and

pale scales or with pale scales aggregating into conspicuous pale spots, pro-

ducing a striking pattern. Abdomen. Terga II-VII usually with basal trans-

verse pale bands, sometimes with apical or apical and basal pale bands, rarely

entirely dark. Genitalia. As described for the subgenus, apparently without

distinctive group characters.

MALE. Head, Palpal segment 3 with median pale band on dorsal surface,

ventral surface with 1-2 rows of short, distally flattened scalelike setae or

fine hairlike setae along the whole length, apical 0.25-0.50 with ventrolateral

tuft of strong bristles; segments 4 and 5 usually strongly long plumose and with

basal pale bands on dorsal surface; apex of segment 5 usually pale to tip, some-

times entirely dark. Proboscis usually with ventral tuft of long hairs proximad

of median pale ring, rarely absent entirely.

MALE GENITALIA. Basimere. Slender, conical, rarely modified. Sub-

apical lobe. Proximal and distal divisions not clearly divided, setae in group

d-f of distal division usually 3-4 in number, their bases largely overlapped

with those of setae a-c in proximal division. Distimere. Normal, sickle-

shaped, rarely modified; 1 dorsal and 1 ventral tiny setae present beyond mid-

dle. Phallosome. Lateral plate usually with complex inner and outer divisions,

sometimes simple, with inner division only (Sinensis); apical portion of inner

division with numerous minute spicules and/or several denticles and usually

with prominent crown of 3-6 strong porrect, fingerlike processes or teeth on

apical tergal surface; outer division simple or complex with 1,2 mesal spines

and variable number of lateral spines, all divergent laterad; sternal spine poor-

ly or well developed, usually strongly bent basad, sometimes projecting dor-

solaterad; lateral basal process variously developed. Proctiger. Basal ster-

nal process varied, usually well developed, sometimes poorly developed or

o6 Contrib. .Amer. Eat...Inst.., vobs12% :N0.2) 1976

absent; subbasal process usually present, poorly or well developed.

PUPA. Trumpet. More or less cylindrical or asymmetrically funnel-

shaped; pigmentation varied from yellowish to dark brown. All setae developed,

essentially as in the subgenus. Abdomen, Seta 1-II usually 1-7 branched or

less than 10 branched, rarely more branched (ge/idus and tritaeniorhynchus).

Paddle: Usually entirely pale, sometimes with distinct color pattern; setae 1,

2-P present.

LARVA. Essentially as described for the subgenus with the following group

characters. Head. Labrum developed as distinct transverse bar or not

(Bitaeniorhynchus Subgroup); seta 1-C stout, flattened, spiniform or sometimes

foliform. Thorax. Seta 4-P variously developed, usually double, sometimes

single or multiple; 7-P always triple; 8-P usually double, rarely single. Abdo-

men. Seta 7-I usually single, sometimes double; comb scales varied in num-

ber, size and in apical fringe of spicules. Saddle complete, rarely incomplete;

posterior caudal margin lightly to moderately spiculated; seta 1-X usually weak,

double or triple, sometimes single; 4-X always with 6 pairs of setae; anal gills

usually slender and longer than saddle, sometimes very short, stubby or

rounded. Siphon. Length and shape varied, usually long, slender and distally

tapered; pecten usually well developed with 10 or more teeth, sometimes poorly

developed with 3-5 teeth; siphonal tufts 3-8 pairs in number, length, branching

and arrangement varied, usually strong and conspicuous. :

DISCUSSION. Members of the Sztzens Group are separated from those in the

Pipiens Group in the adults by the presence of conspicuous pale markings on the

palpus (particularly in males), proboscis and tarsi of legs as indicated in the key

and as characterized above. In the majority of Oriental species, the males are

also strongly characterized by the presence of a ventral tuft of long hairs in the

middle of proboscis and by having the short setae on ventral surface of palpal

segment 3 more or less scalelike in appearance. In the male genitalia, the

presence of numerous minute spicules in the apical portion of the inner division

of the phallosome, which is probably not homologous with the ventral arm in

the Pipiens Group, is also characteristic of the Sztiens Group. The pupae and

larvae of the Sitzens Group exhibit much overlap with the Pipzens Group. Ex-

cept for the Bitaeniorhynchus Subgroup, the pupae and larvae of the rest of

Sitiens Group are poorly characterized at the group level but can be distin-

suished from the Pipiens Group by several combinations of characters.

The Sitiens Group is largely confined to the tropics of the Ethiopian,

Oriental and Australiasian regions and to the islands of the South Pacific and

Micronesia with a small extension into the Mediterranean, Middle East and

north and northeastern Palearctic. Within the Oriental region, the group is

dominant and diverse, comprising several species most of which are restricted

to Southeast Asia and adjacent areas. In this study 36 species of the Sztiens

Group are recognized in the Oriental region and among these 5 are considered

as new.

Edwards (1932) subdivided the Sztiens Group into 4series: Bitaeniorhynchus,

Sitiens, Gelidus andMimeticus. Thisclassification was based almost exclusively

on the conspicuous coloration oftheadults. In subsequent local studies (Baisas 1938,

Belkin 1962 and Bram 1967a), Edwards' scheme has been largely followed with only

slight modifications, especially inthe realignment of species inthe Sitiens series and

in using the terms "'group" or ''subgroup" in preference to ''series''. Inadopting sub-

eroup categorization, the only change which has been made was that by Bram (1967a)

who, following the interpretation of the Vishnui Group of Colless (1957a), subdivided

the Sitiens series of Edwards into Vishnui and Sitiens Subgroups. Except for the align-

ment of certain species inthe Vishnuz Sugroup, the subdivisionby Bram is justified.

Sirivanakarn: Subgenus Culex in the Oriental Region o7

I recognize 6 subgroups for the 36 Oriental species inthe SitiensGroup. One of

these: the Barraudi Subgroup is proposed for the first time, the other 5 sub-

groups: Gelidus, Bitaeniorhynchus, Sitiens, Vishnui and Mimeticus as categor-

ized by Edwards (1932), Belkin (1962) and Bram (1967a) are retained except for

a few changes in the arrangement of species. Each of the subgroups which con-

tains 3 or more species, is further segregated into various complexes and the

attempt is made to briefly characterize them in all stages. This scheme of

classification is as follows: (1) Gelidus Subgroup with gelidus; (2) Bitaenior-

hynchus Subgroup represented by: (a) bitaeniorhynchus complex with bitaenior-

hynchus, infula, luzonensis, selangorensis, pseudosinensis and longicornis;

(b) Stmensis complex with sinensis, cornutus and epidesmus and (c) geminus

complex with geminus and kinabaluensis; (3) Sitiens Subgroup represented by:

(a) sitiens complex with sitiens and alis; (b) whitmorei complex with whitmorei

and (c) annulirostris complex with annulirostris; (4) Vishnui Subgroup repre-

sented by: (a) vishnui complex with vishnui, pseudovishnui, perplexus, alienus,

philippinensis, and incognitus; (b) tritaeniorhynchus complex with tritaenior-

hynchus and (c) whitei complex with whitei; (5) Barraudi Subgroup with barraudi

and edwardsi; (6) Mimeticus Subgroup represented by: (a) mimeticus complex

with mimeticus, fasyi, jacksoni, tsengi, mimuloides and diengensis and (b)

mimulus complex with mimulus, murrelli,: propinquus and orientalis.

All of the 6 subgroups except Gelidus are related on the basis of the male

genitalia. The Gelzdus Subgroup as recognized here is monotypic, including

only gelidus which is unique in the type of male phallosome and in the ornamen-

tation of the head and mesonotum of the adults and in the larval siphon, but

essentially conforms to the Sitiens Group in several other characters. The

Bitaeniorhynchus Subgroup is most complex and heterogeneous in the color of

the adults and in the types of male phallosome, but all species involved are

basically similar in the pupal and larval stages. It shows overlap with the

other 4 subgroups (Sitiens, Vishnui, Barraudi and Mimeticus) in the male geni-

talia and in the external adult characters but is very distinct in the larva and

pupa. The Sitiens, Vishnui, Barraudi and Mimeticus subgroups are essentially

Similar in most features of the male genitalia but are slightly or strikingly dif-

ferentiated from one another in the ornamentation of the adult (e.g. presence

of pattern of pale spots on the wing in Mimeticus subgroup), in the special modi-

fication of certain features of the male phallosome and in the immature stages.

For further discussion and characterization of individual subgroups, see the

treatment under each category.

GELIDUS SUBGROUP

The Gelidus Subgroup is characterized by the following combination of

characters: in the adults by (1) narrow decumbent scales and erect scales in

center of vertex of head pure white; (2) anterior 0.7 of mesonotum covered

with pure white scales; (3) scales on posterior 0.3 of mesonotum, prescutellar

space and on scutellar lobes deep brown to black; (4) apn and ppn without

scales; (5) anterior surface of fore- and midfemora entirely dark; (6) scales on

all wing veins dark and narrow and (7) abdominal terga II- VII with basal pale

bands; the male by (1) ventral surface of palpal segment 3 with short and fine

hairlike setae; (2) apex of palpal segment 5 entirely dark and (3) proboscis with-

out median ventral tuft of long hairs; the male genitalia by (1) distimere dis-

tinctly swollen at middle; (2) inner division of phallosome represented by a

slender spiculose, fingerlike process; (3) outer division broad and more or

08 Contrib. Amer. Ent. Instu,. vole 12; no. 2; 1976

less rounded, without sharp denticles or spines and (4) basal sternal process of

proctiger very thick, dark and long, curved sternad; the pupa by (1) trumpet

relatively short, cylindrical or gradually distally widened and dark brown; (2)

seta 1-II 20 branched and (3) paddle with distinct dark spot towards apex of

midrib and in the larva by (1) labrum distinct; (2) seta 1-C dark, slender and

spiniform; (3) seta 7-I single; (4) siphon short, stout, more or less fusiform

(somewhat resembling guinquefasciatus and hutchisoni) and (5) spiracular cham-

ber of siphon with distinct erect spicules.

DISCUSSION. The interpretation of the Gelidus Subgroup by Edwards (1932:

203) which was later followed by Bram (1967a: 247) was considered to comprise

4 species: gelidus Theobald 1901 and whitmorei (Giles 1904) from the Oriental

region; vicinus (Taylor 1916) from Northern Australia and New Guinea and 07-

hamatus Edwards 1926 from Timor, Indonesia. This classification which was

based largely on the resemblance in the coloration of the head and anterior 0.7

of the mesonotum is not justified on the basis of the present study and a detailed

comparison of the male genitalia. The Gelidus Subgroup in the present sense is

restricted to gelidus which is differentiated from other subgroups by the unique

phallosome and the peculiar swelling in the middle of distimere of the male

genitalia and by the larval characters as summarized above. The affinity of

whitmorei is apparently with the members of the Sifiens Subgroup to which it

is now being transferred and that of vicinus and bihamatus is uncertain. The

latter 2 species are probably restricted to the Australasian region and appear

to form a distinct group or subgroup with affinity close to the Sztiens and Bitaenti-

orhynchus Subgroups as pointed out by Sirivanakarn (1974).

The relationship between the Gelidus Subgroup and other subgroups in the

Sitiens Group is not clear. The adults of gelidus essentially conform to the

Sitiens Group characters but the male genitalia and the larva are so strongly

differentiated that it might well be considered as a distinct group separated from

the rest of the Szftzens Group.

7. CULEX (CULEX) GELIDUS THEOBALD

(Figs. 15, 17, 18, 19)

Culex gelidus Theobald 1901b: 20 (2*); Giles 1902: 421 (2); Theobald 1903: 180

(distribution); Blanchard 1905: 316 (?); Leicester 1908: 147 (%, 2); Bar-

raud 1924a: 990 (“*); Barraud 1924c: 428 (L*); Borel 1930: 327 (o*, 2,

L*); Barraud and Christophers 1931: 282 (distribution).

Culex gelidus var. cuneatus Theobald 1901b: 22 (¢*); Theobald 1910a: 313

(distribution); Edwards 1932: 203 (synonymy).

Leucomyia gelida (Theobald), Theobald 1907: 372 (%*, taxonomy); Theobald

1910a: 312 (distribution).

Leucomyia gelida var. bipunctata Theobald 1907: 374 (“); Edwards 1932: 203

(synonymy). 7

Theobaldiomyia gelidus Theobald, Brunetti 1912: 462 (taxonomy).

Culex (Culex) gelidus Theobald, Edwards 1932: 203 (taxonomy); Barraud 1934:

407 (**, 2, L*); Bonne-Wepster and Brug 1937: 74 (o%, $*); Baisas 1938:

212 (**, 9*, L¥*, P*); Bonne-Wepster and Brug 1939: 1271 (L*); Monchad-

skii 1951: 262 (L*); Bonne-Wepster 1954: 119 (“, 9*, L*); Delfinado 1966:

144 (o'*, 2, L*): Bram 1967a: 247 (o*, 9*, P*, L*); Baisas 1974: 92 (¢,

PPM TL):

FEMALE (Fig. 15, 17). Wing: 2.3-3.2 (average 2.8 mm). Forefemur:

Sirivanakarn: Subgenus Culex in the Oriental Region 29

1.4mm. Proboscis: 1.7mm. Abdomen: 2.7mm. Small to medium sized

species; most clearly marked by the predominantly white scales on vertex of

head and on anterior 0.7 of mesonotum. Head. Narrow decumbent scales on

vertex pure white; erect scales numerous, largely pure white in center and on

anterior ocular line, black on posterolateral area; lateral patch of broad scales

beige or dingy white anteriorly, dark blue or brown posteriorly. Palpus

entirely dark scaled, rather short, about 0.2 of proboscis length. Proboscis

with narrow median pale ring. Czibarial Aymature., Cibarial teeth about 25 in

number, all apparently long, very fine and distally filamentous, in a concave

row which is slightly produced at middle. Thorax. Mesonotal integument dark

brown; anterior 0.7 of mesonotum densely covered with pure white scales,

strongly contrasted with dark brown scales on prescutellar space, supraalar

and posterior dorsocentral areas in posterior 0.3; scales on scutellar lobe

entirely dark brown; middle of fossa with or without dark scales aggregating

into distinct spot. Integument of pronotum dark brown; apn and ppn without

any scales. Pleural integument dark brown on ppl, ssp, psp and upper sfp;

paratergite dark brown, rest of integument pale brown; pleural scale patches

poorly developed or reduced; scales on ppl and upper mep absent; sip with a few

scattered scales on upper corner and a minute scale patch on lower posterior

border. Legs. Anterior surface of fore- and midfemora entirely dark scaled;

anterior surface of hindfemur slightly pale on lower surface, dark on upper

lateral surface; all tibiae entirely dark anteriorly; tarsomeres 1-4 of all legs

with narrow apical and basal pale bands. Wing. All scales dark brown, plume

scales on veins Ro, Re and R narrow, linear. Abdomen. Terga II-VI with

complete basal pale bands which are strongly produced in middle; terga VII-

VIII with narrow even basal pale bands; sterna II-VII, entirely pale yellow;

sternum VIII largely dark. Genitalia. Essentially similar to most species;

tergum IX with lateral row of 6,7 setae; cerci with relatively few and weak

setae; postgenital plate rounded or lightly emarginate on caudal margin; vagin-

al sclerite typically V-shaped; insula with a dense tuft of 10-12 setae.

MALE (Fig. 17). In general as described for female. Head. Palpus ex-

ceeding proboscis by the full length of segment 5; segment 3 with median pale

band on dorsal surface; ventral surface with 1,2 rows of short, fine hairlike

setae, apical 0.4 with ventral lateral tuft of 10-20 dark and long bristles; seg-

ments 4 and 5 moderately to strongly plumose, dorsal surface with basal pale

bands, apex of segment 5 dark to tip. Proboscis without ventral long hairs

proximad of median pale ring.

MALE GENITALIA (Fig. 18). Segment IX. Tergum narrow, tergal lobe

with a row of 6-7 moderately strong setae. Basimere. Normal, conical,

setae adjacent to subapical lobe rather dense. Subapical lobe. Seta 6b of proxi-

mal division thicker and longer than setae a and c; setae d-f of distal division

consists of 1 club-shaped blade and 2 narrower and apically pointed blades;

leaf g broad, short, heart-shaped; seta h rather weak and apically hooked.

Distimere. More or less sickle-shaped ; basal part even in width and straight,

middle part distinctly swollen, apical part tapered and slightly recurved; sub-

apical claw slender and moderately long; 1 dorsal and 1 ventral tiny setae pre-

sent, ventral distad of dorsal. Phallosome. Lateral plate relatively simple;

inner division represented by an elongate, fingerlike, strongly spiculose pro-

cess, divergent laterad; outer division represented by a simple, broad rounded

sclerite which is finely spiculose on outer and inner apical surfaces, inner ter-

gal surface (in lateral view) produced into a short finely spiculose process,

projecting basad. Pyroctiger. Apical crown medium sized, composed of sever-

al dark and strong spicules; paraproct very narrow; basal sternal process very

60 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

thick, long and dark pigmented, strongly curved sternad; cercal sclerite largely

membranous; cercal setae 2 in number.

PUPA (Fig. 18). Abdomen: 2.5mm. Paddle: 0.65 mm. Trumpet: 0.45

mm; index 4-5. Cephalothorax and abdomen yellowish white. Complete chae-

totaxy as figured; the following are characteristic. Trumpet. Brownish; rela-

tively short, distally widened; apical margin truncate; pinna broad, oblique.

Cephalothorax. Seta 1-C usually 4,5 branched (8-5); 3-C stronger than 1-C,

usually triple (3-4); 8, 9-C usually triple (2-4). Metanotum. Seta 10-C 8-10

branched; 11-C usually double (2-3); 12-C 4,5 branched. Abdomen. Seta 1-II

multiple, about 20 branched; 5-IV usually 4 branched (3-4); 5-V usually triple

(2-3); 5-VI usually double (2-3); 6-III-V usually triple (2-4); 6-VI usually 4

branched (3-4); 6-V, VI strong, about 2 times as long as 6-III, IV; 9-VII 4 bran-

ched; 4- VIII double or triple; 9-VIII 6-8 branched. Paddle. Largely pale

whitish, with distinct dark spot towards apex of midrib; midrib weak, lightly

pigmented; setae 1, 2-P present, minute.

LARVA (Fig. 19). Head: 0.7mm. Siphon: 1.0 mm; index about 3.5.

Saddle: 0.36 mm; siphon/saddle ratio 3. As figured. Head. Frontoclypeus

with darkened spots along lateral posterior margin, rest yellowish white; seta

1-C dark, stout, spiniform; 4-C distinct, single, its length 1.9-2.0 times as

long as distance between bases of the pair; 5, 6-C strong, subequal and triple;

7-C 7,8 branched; 14-C single, minute and indistinct. Antennal shaft pale

except for basal dark ring; spicules strong, relatively few in number; setae 2,

3-A subapical and pale. Mental plate with 8-10 lateral teeth on each side of

median tooth. Thorax. Spiculation not developed; seta 4-P strong, double;

7-P usually triple (3-4); 8-P single or double; 8-M 5-7 branched; 9-M 5, 6

branched; 7-T usually 7 branched (5-8), 9-T 6,7 branched; 12-T usually triple

(2-4); 13-T 8-12 branched. Abdomen. Spiculation not developed; setae 6-I, Il

triple; 7-I single; 1-III, IV long, usually double (1-3); 1-V, VI usually 4 bran-

ched (3-5); 6-III-VI double; 13-III-V 4, 5 branched; 4, 7,10, and 12-VII all

single. Comb scales about 40, in broad oval patch; all rather short and small,

with even apical fringe of fine spicules; seta 1-VIII usually 7 branched (5-7); 2

and 4-VIII single; 3-VIII 6-9 branched; 5-VIII 4,9 branched. Saddle same color

as siphon; posterior caudal margin with or without spicules; 1-X single; 2-X

usually double (2-3); anal gills variable in length, dorsal pairs usually longer

then ventral, 1-2 times as long as saddle. Siphon. Stout, short, more or

less fusiform in shape; lightly to strongly brownish; pecten with 8-12 fine dis-

tally curved teeth, 4-5 distal teeth barbed with 1 strong basal denticle and 5-6 _

weaker distal denticles; siphonal tufts 4 regular pairs, 4,5 branched each

(3-7), all inserted subventrally, first proximal pair among or distad of pecten,

its length slightly shorter than siphonal width at point of attachment, other si-

phonal tufts gradually shorter; seta 2-S dark and slender; median caudal fila-

ment of spiracular apparatus well developed; inner wall of spiracular chamber

with a forked or simple spicule.

TYPE-DATA. (1) Culex gelidus Theobald, Lectotype ?*, Kelang, Selangor,

MALAYA (PENINSULAR MALAYSIA), A. L. Butler (BM; selection of Bram

1967a: 251); (2) Culex gelidus var. cuneatus Theobald, Lectotype ¢*, Straits

Settlements, Taiping, Perak, MALAYA (PENINSULAR MALAYSIA), L. Wray

(BM; selection of Bram 1967a: 251); (3) Leucomyia gelida var. bipunctata Theo-

bald, Holotype “*, INDIA (BM).

DISTRIBUTION. Widespread throughout Southeast Asia with extension to

the west in India, Burma and Sri Lanka, to the north in South China and to the

south and southeast as far as Java, the Molluccas and New Guinea in Indonesia.

Material examined. 2,163 specimens: 6940, 1,146, 323 L; 248 individual

Sirivanakarn: Subgenus Culex in the Oriental Region 61

rearings (105 pupal, 143 larval).

INDIA. Bengal: Calcutta; Lalmanirhat; Salt Lake; Dum Dum; Assam:

Chabua; Rupsi; Jorhat; Dinjam; Dibrugarh; 250", 672, 15 L, 5p.

SRI LANKA. Ratanapura; Anu; 2%, 129.

BURMA,::;Rangoon: ( 130%4. 259012: Fin p12 Ips

THAILAND. Chiang Mai; Lampang; Udon Thani; Ayuthaya; Dhon Buri;

Bangkok; Chon Buri; Trat; Rayong; Nhon Tha Buri; Phet Buri; Ranong; Krabi;

Phuket; Satun; Narathiwat; 3960, 5452, 205 L, 78 p, 52 lp.

CAMBODIA. Phnom Penh; Ari Gsatr; 6%, 279.

VIETNAM. Saigon; Vinh Long; Danang; Cam Ranh Bay; Can Tho; Gia Dinh;

Phu Lam; Bien Hoa; My-Ca; Pleiku; Soc Trang; 180, 289, 14 L, 3p, 1 lp.

MALAYSIA. Peninsular Malaysia: Selangor; Kuala Lumpur; Rantau Pan-

jang; Ulu Gombak; Segambut; Bt. Kutu; Puchong; Kepong; Klang Rd. ; Salak;

Perak: Kuala Dipang; Pusing; Senlu; Perlis: Kg. Gunong; Chenderiang; C.

Highland Rd.; Kuala Kangsar; Pahang: Kuantan; Kelantan: Tumpat; Kedah:

Sintok; 590°; 1302, 34 L;10p, 53 1p; Malaysia: Sarawak: Kuching; Sabah: K.

Belud; Kota Kinabalu (Jesselton); Papar; Tawau; Tenom; 11°, 109, 2 lp.

INDONESIA. Sumatra: Kmp Pantai Pandjang; Belakang Pandok; Kebon

Ros; Tand Jungkarang; Bengkulu; Kalimantan (Borneo): Tarakan; Kalabakan;

Java: Djakarta; Sulawesi: Makassar; Molluccas: Morotai; 8%, 509.

PHILIPPINES. Luzon: Los Banos; S. Fernando; La Union; Bulacan; Camp

Stotsenberg; Camp Nichols; S. Fabian; Rizal, Wack Wack; Mindoro: San Jose;

Samar: §. Antonio; Osmena; Palawan: Panitan; Tinabog; Calaccad; Iwahig;

Leyte: Dulag; Abuyog; Carigara; Tacloban; Bay Bay; Mahaplag; Basilan: Isabela;

Calicoan Is.: Sanga Sanga; Lapit Lapit; Tawi Tawi Is.: Kamagong; Tarawakan;

Mindanao: Parang; Kabakan; Pangasinan; Jolo Jolo Is.: Sulu; 1560", 2549, 43 L;

Iwidy 25° Lp

Additional records from the literature. CHINA: Canton (Kwangtung);

Hainan Is. ; HONG KONG (Feng 1938: 300, Chu 1958: 109). FORMOSA (TAI-

WAN); INDIA: Bombay; Deccan; Madras, Nilgiri Hills; Southwest Coast; Ben-

geal: Chittagong, Sukna; Bihar; Assam: Nongpoh, Dimapur (Barraud 1924a:

991; Barraud 1934: 409). INDONESIA: Simaloer; Lombok; Soemba, Alor, New

Guinea (Brug and Bonne-Wepster 1947: 187).

TAXONOMIC DISCUSSION. Culex gelidus is extremely uniform, exhibiting

constant diagnostic characters in all stages without any indication of local or

eeographic differentiation. The only variations noted in the adults are the

presence of absence of dark scale spots in the middle of the fossae and the dark

streak in the middle of acrostichal line on the anterior 0.7 of the mesonotum.

These variations are nothing more than the normal ones observed in several

series of reared or field caught specimens from different localities.

Culex gelidus is one of the most common and best known Southeast Asian

species. The adults are conspicuously marked, with predominantly pure white

scales on the head and on the anterior 0.7 of mesonotum. The only other Ori-

ental species which exhibits more or less similar ornamentation is whitmorezi.

The adults of gelidus, however, differ from whitmorei rather strikingly in (1)

absence of speckling of pale scales on the anterior surface of mid- and hind-

femora; (2) scales on posterior 0.3 of mesonotum (including prescutellar space

and scutellar lobes) entirely dark (pale whitish in whitmorei) and (3) scales on

wing vein Ry,5 narrow and considerably less dense (broad and very dense in

whitmorei). In addition, gelidus adults also differ strikingly from all other

Oriental members of the Sitiens Group in the absence of scales on apn and ppn

and in the considerable reduction of scales on propleuron and stenopleuron.

The female cibarial armature of gelidus is also distinctive and can be distin-

62 Contrib, Amer.’ Ent. Inst. ,.vol.°12, no. 2, 1976

suished from all other sitiens species except tritaenitorhynchus in having fila-

mentous teeth. The male genitalia of gelidus are unique in the relatively simple

phallosome which separates it from the rest of the Siftzens Group. The imma-

ture stages show much overlap with all other members of the Sifzens Group in

the pupal and larval chaetotaxy. The pupa can be readily recognized by the

relatively short and brownish trumpet; seta 1-II multiple and by the presence

of dark spot towards apex of the paddle; and the larva by the short, stout and

fusiform siphon and by the presence of distinct erect spicules on the inner wall

of spiracular chamber. The presence of spicules in the spiracular chamber of

the larval siphon has not been known or described in any other species of Culex.

This character has been found to be useful in separating gelidus larva from

those of quinquefasciatus which it strongly resembles in the shape of the siphon.

_ BIONOMICS. Culex gelidus has been found closely associated with man and

his domestic animals, such as cows, water buffalo and pigs in rural or urban

communities of Southeast Asia. The recorded breeding sites of gelidus include:

puddles, pools, ponds, ditches, drains, streams, rice fields and marshy de-

pressions which usually contain abundant aquatic vegetation such as water lilies,

hyacinths, duckweeds and grasses. The water ranges from fresh and clear to

strongly colored with contamination from decayed organic matter and mud par-

ticles. On occasion, it has also been found breeding in tanks, barrels, earth-

enware pots and coconut shells. The immatures and adults occur throughout

the year, but to date nothing is known about their relative abundance during

different seasons as well as the factors which affect their population size.

Adults of gelidus have been frequently collected in number from light and ani-

mal baited traps. They occasionally enter houses at dusk and several females

were caught biting man. The natural or preferred hosts of gelidus are cows

and pigs (Colless 1959; Macdonald et al. 1967), but man is also readily attacked

as an alternative host. In Malaya and Sarawak it was also reported to feed on

other animals including: water buffalo, goats, deer, chickens and wild birds

(Gould et al. 1962; Hill et al. 1969).

MEDICAL IMPORTANCE. Adults of gelidus were reported to be naturally

infected with Brugia malayi in Sri Lanka by Carter (1948) and with Wuchereria

bancrofti in Travancore, India by Iyengar (Raghavan 1961) but the incidence of

these pathogens were very low, indicating that this species does not play a role

in the natural transmission of these diseases. In epidemiological and experi-

mental transmission studies of Japanese encephalitis virus, gelidus has been

incriminated as a vector of this disease in Malaya (Gould et al. 1962) and Sara-

wak (Macdonald et al. 1967). In Thailand, Simasathien et al. (1972) reported

that more strains of JE virus were isolated from gelidus than from tritaentor -

hynchus and suggested that it may be more important than the latter in the trans-

mission of the disease in that country. In the recent studies on virus isolations,

gelidus was reported to be naturally infected with 3 strains of JE virus and 2

strains of Tembusu virus in Chiang Mai, Thailand (Gould et al. 1974) with 2

strains of JE virus in Saigon, Vietnam (Nguyen et al. 1974). These data

strongly suggest that gelidus is probably one of the most important vectors of

JE virus in Southeast Asia.

BITAENIORHYNCHUS SUBGROUP

The Bitaeniorhynchus Subgroup is characterized by the following combina-

tion of characters: in the adults by (1) large size (wing length usually over 4.0

mm); (2) erect scales of vertex varying from partially pale in center and dark

Sirivanakarn: Subgenus Culex in the Oriental Region 63

on posterolateral areas to completely dark; (3) anterior 0.7 of mesonotum vary-

ing from predominantly pale beige, whitish, yellowish or golden to dark brown;

(4) scales on posterior 0.3 of mesonotum laterad of prescutellar space dark

brown; (5) scales on prescutellar space and scutellar lobes partially pale to

completely dark brown; (6) apn and ppn with several narrow scales; (7) anteri-

or surface of femora and tibiae varying from lightly to heavily speckled with

pale scales or completely dark; (8) wing scales entirely dark or lightly to

heavily speckled; (9) abdominal terga II-VII with apical or basal pale bands or

both sometimes absent or completely dark; the male by (1) ventral surface of

palpal segment 3 with numerous dark short, hairlike setae; (2) apex of seg-

ment 5 and bristles largely pale yellowish and (3) proboscis with prominent

median ventral tuft of several long hairs; the male genitalia by (1) distimere

normal, sickle-shaped; (2) inner division of phallosome varied; simple or com-

plex, with or without prominent apical tergal crown of 4-5 strong fingerlike

processes or teeth, sometimes with a subapical tergal lobe bearing 2-4 foli-

form processes; (3) outer division simple and broad, leaflike or with distinct

median, lateral and sternal spines, sometimes largely degenerate or not

developed (sinensis) and (4) basal sternal process of proctiger varying from

short, slender and pale to very long, thick and dark; the pupa by (1) trumpet

asymmetrically funnel-shaped; (2) seta 1-II usually single to triple or at most

5 branched and (3) paddle varying from largely pale to partially darkened and

in the larva by (1) head capsule comparatively smaller than in other subgroups;

(2) labrum (or preclypeus) not apparent or very poorly developed; (3) teeth of

mental plate varied from relatively few, strong and distinct to numerous, mi-

nute and indistinct; (4) setae 2 and 4-VIII 2-7 branched; (5) comb scales varied,

usually large, spinelike, 4-5 in number, sometimes small, 10-11 in number;

(6) siphon slender, long, with 3-5 pairs of subventral tufts; (7) median caudal

filament of spiracular apparatus well developed and (8) anal gills long and

tracheate.

DISCUSSION. The Bitaeniorhynchus Subgroup as interpreted here corre-

sponds to the Bitaeniorhynchus Series of Edwards (1932: 202-3) and to the

Bitaeniorhynchus Subgroup of Belkin (1962: 202) and Bram (1967a: 257). This

subgroup occupies a very broad range of distribution, covering the Ethiopian,

Oriental and Australasian regions, Micronesia and the South Pacific. All mem-

bers of the subgroup are inland species with breeding habitats more or less

restricted to fresh water ground pools and stagnant streams which contain

masses of green algae. The Bitaeniorhynchus Subgroup is apparently better

represented within the Oriental region than anywhere else. In this study 11

species have been recognized. It is quite possible that more species will be

discovered in certain parts of the Indomalayan areas, such as Borneo, small

and major islands of Indonesia, and the Philippines, when more extensive col-

lections are made.

The members of the Bitaeniorhynchus Subgroup are strongly differentiated

from the rest of the Sitiens Group in the pupa and larva. The pupa is readily

recognized by the relatively large size and the short, asymmetrically funnel-

shaped trumpet and the larva by the reduced head capsule, the absence of

labral bar (sclerite), setae 2 and 4-VIII 2-7 branched (single in all other mem-

bers of the Sitiens Group), the reduction in number and size of the pecten teeth

and the tracheate anal gills. The adults of the majority of species are well

marked by the coloration of the mesonotum, legs, wing and abdomen. Some

also exhibit a strong sexual dimorphism in coloration not usually found in the

other species of Culex. The male genitalia exhibit a vast array of modifica-

tion in the phallosome, from simple to complex, overlapping with that in the

64 Contrib. :Amer. Bntsdnst,, vole: 12,no0..2; D976

other subgroups of the Sztiens Group. The extreme diversity of forms and the

very broad range of distribution seem to suggest that the Bitaeniorhynchus Sub-

group has existed for a long time and probably represents the original ancient

stock of the Szfzens line from which other subgroups, including: Sitiens, Vish-

nui, Barvraudi, Mimeticus and perhaps also Gelidus were derived.

On the basis of comparative male genitalia and larval stage, the 11 Oriental

members of the Biiaenitorhynchus Subgroup apparently fall into 3 more or less

distinct complexes: bitaeniorhynchus, sinensis and geminus.

The bitaeniorhynchus complex includes: bitaeniorhynchus (type form),

infula, luzonensis, selangorensis, pseudosinensis and longicornis. This com-

plex is characterized in the male genitalia by (1) the rudimentary or short and

slender basal sternal process of the proctiger; (2) apical spiculate portion of

the inner division of the phallosome enlarged, elongated, in form of a long beak

or stout, resembling the blade of an ax; (3) tergal surface of the inner division

with a distinct lobe, bearing 2-4 foliform processes subapically or at some dis-

tance from the apical tergal angle of apical spiculate portion and (4) outer divi-

sion represented by a very broad, acuminate leaf; in the larva by (1) setae 2

and 3-A apical or slightly subapical and (2) mental plate with numerous deli-

cate, indistinct lateral teeth whose apices are in straight line. The members

of the bitaeniorhynchus complex show much overlap with one another in one

or more stages and some of them may be difficult to differentiate. To insure

correct identification, all stages of each member in this complex should be

examined.

The sinensis complex includes sinensis, cornutus and epidesmus and is

characterized in the male genitalia by (1) the large, heavy and long basal

sternal process of the proctiger; (2) apical spiculate portion of inner division

of phallosome slender, distally tapered or broad and rounded; (3) tergal sur-

face of the apical spiculate portion with or without a slender lobe bearing 1-2

narrow, elongate teeth subapically and (4) outer division not developed or

developed as a broad lobe bearing a strong mesal spine and a variable number

of denticles or tubercles laterally; in the larva by (1) setae 2 and 3-A situated

far below the apex and (2) mental plate with 10-12 distinct lateral teeth. The

larva of the sinensis complex as characterized here is based only on those of

sinensis and cornutus. The larva of epidesmus is still unknown, however, its

male genitalia are extremely similar to cornutus to which it is undoubtedly

related. |

The geminus complex includes geminus and kinabaluensis, both of which —

are related to squamosus (Taylor 1913) from New Guinea (Assem and Bonne-

Wepster 1964: 128) and the Solomon Islands (Belkin 1962: 214) and to albinervis

Edwards 1929 from the Fiji Islands (Belkin 1962: 212). Both Southeast Asian

species in this complex and squamosus strongly resemble those of the Sitiens,

Vishnui and Mimeticus Subgroups in the male phallosome, but are exceedingly

similar to the bitaeniorhynchus and sinensis complexes in the pupa and larva.

The geminus complex is characterized in the male genitalia by (1) the presence

of a long and thick basal sternal process of the proctiger; (2) apical spiculate

portion of inner division of phallosome not enlarged or distally expanded; (3)

tergal surface of the apical spiculate portion with a prominent crown of 3-6

strong, fingerlike processes projecting beyond upper margin of apical sternal

portion and (4) outer division represented by a broad lobe bearing mesal, later-

al or also sternal spines; in the larva by (1) setae 2 and 3-A apical or subapical

and (2) mental plate with a number of strong distinct lateral teeth on each side

of a large median tooth, followed by several delicate indistinct lateral teeth

towards lateral angle.

Sirivanakarn: Subgenus Culex in the Oriental Region 65

8. CULEX (CULEX) BITAENIORHYNCHUS GILES

(Figs 15, 20; 295.22)

Culex bitaeniorhynchus Giles 1901a: 607 (adult); Edwards 1913b: 231 (taxonomy,

in part); Edwards 1921: 337 (distribution, in part); Edwards 1922a: 282

(**, 9, taxonomy, in part); Barraud 1923a: 936 (L*); Barraud 1924a:

984 (o“*, 2, in part); Borel 1930: 316 (“*, 2, L*, in part); Ho 1931: 149

(o*, ).

Taeniorhynchus ager Giles 1901b: 196 (", 2); Edwards 1912: 30 (2*); Edwards

1913b: 231 (synonymy).

Culex kavatsuensis Mochizuki 1913: 28 (“*, 9*, E); Edwards 1932: 202

(synonymy).

Culex (Culex) bitaeniorhynchus Giles, Edwards 1932: 202 (taxonomy); Barraud

1934: 304 40%, 0 OOO ie part): Bonne-Wepster and Brug 1937: 73 (o, §);

Baisas 1938: 211 (o*, Om, P*, L*, in part); Bohart 1945: 78 (2, L); Bohart

and Ingram 1946: 77 (o'*, &, L*): Mohan 1950: 167 (biology); LaCasse and

Yamaguti 1950: 201 (“*, 2*, P*, L*); Asanuma and Nakagawa 1953: 93

(P*); Delfinado 1966: 140 (o*, 2, P, L*, in part); Bram 1967a: 258 (c*,

Q*, P*, L*, in part); Sirivanakarn 1973: 235 (taxonomy); Baisas 1974: 88

(os Sty Lae

FEMALE (Fig. 15, 20). Wing: 4.3 mm. Forefemur: 2.0mm. Proboscis:

2.0mm. Abdomen: 3.0-3.2 mm. Medium to large sized species; distinguished

from other members of the Bitaeniorhynchus Subgroup by the extensive speck-

ling of pale scales on anterior surface of femora and tibiae of legs, the exten-

sive speckling of pale scales on all wing veins, veins Ro, R and R4,5 with

relatively broad plume scales and by the presence of only broad apical pale

bands on abdominal terga II-VII in both sexes. Head. Narrow decumbent

scales of vertex pale, yellowish or beige; erect scales numerous and coarse,

usually pale, golden or bronzy in center, dark brown on posterolateral areas,

sometimes predominantly dark brown; lateral patch of broad appressed scales

largely pale along eye margin, dark posteriorly. Palpus about 0.2 of probos-

cis length, segments 3 and 4 with sprinkling of pale scales, apex of segment 4

usually tipped with pale whitish scales. Proboscis relatively thick, as long as

or slightly shorter than forefemur; labium with a broad median light ring and

usually also with a narrow apical light ring; basal portion proximad of median

ring lightly sprinkled with pale scales. Cibarial Armature. Cibarial bar con-

cave and lightly produced at middle; cibarial teeth about 30 in number, median

4-6 teeth narrow and pointed apically, lateral teeth shorter, thick and coarse,

apically blunt or truncate. Thorax. Anterior 0.7 of mesonotum usually pre-

dominantly pale, beige, yellowish or golden scaled, with or without mottling of

some dark brown scales, forming a pair of dark spots in middle of fossae or

distinct blotches and streaks elsewhere; sometimes predominantly brownish

with indefinite pale areas; scales on posterior 0.3 of mesonotum dark brown-

ish on posterior dorsocentral and supraalar areas; prescutellar space largely

dark scaled in middle, pale towards posterior margin; mid-scutellar lobe dark

scaled anteriorly, pale scaled on caudal margin; lateral scutellar lobes entire-

ly pale scaled. Apn and ppn with several narrow dark, golden or bronzy

scales; ppn with 2 rows of 4-6 strong and 5-7 weak bristles on posterior bor-

der. Pleuron with a small pale scale patch on ppl, upper corner and posterior

border of sip; mep usually bare or with a few pale scales on upper anterior sur-

face; a few scales usually present among upper mep bristles; ppl bristles 10-12

66 Contrib. Amer. Ent. Inst., vols 12, no. 2,°1976

in number. Legs. Anterior surface of all femora heavily and extensively

speckled with pale scales; anterior surface of all tibiae moderately speckled;

tarsomeres of all legs usually lightly speckled; tarsomere 1-4 with narrow

apical and broad basal pale bands. Wing. All wing veins with an equal mix-

ture of pale yellow and dark scales or extensively and heavily speckled;

squame and plume scales on veins Ro, Re and Ry4,5 broad, ovate. Abdomen.

Median caudal scale patch of tergum I usually pale, sometimes entirely dark;

terga II-VII with evenly broad, yellow or beige apical bands, varying from 0.3

to 0.5 of segment length; basal pale band absent or not developed; basolateral

pale spots present; dark scaled areas lightly to heavily sprinkled with pale

scales; tergum VIII with a narrow apical and broad basal pale bands; sterna

II- VII largely pale yellowish except for dark scaled patches on lateral apical

portion. Genitalia. Terga IX with lateral row of 4-7 bristles; postgenital

plate broadly rounded or slightly emarginate on caudal margin; vaginal sclerite

well sclerotized, V or U-shaped; insula with a tuft of 10-12 setae.

MALE (Fig. 20). In general similar to female except for less dense

scales and less speckling of wing and for having more numerous ppl

bristles (about 20). Head. Palpus very long, exceeding proboscis by nearly

length of segments 4 and 5 combined; segment 3 with a broad median

pale band and light sprinkling of pale scales proximad or distad of band on

dorsal surface, ventral surface with numerous short, dark and hairlike setae

along the whole length, apical 0.4-0.5 with ventral lateral tuft of about 30

strong bristles, upper inner surface with a row of 8-10, widely spaced, weak

and long setae in apical 0.5; segments 4 and 5 upturned, strongly long plumose,

with distinct basal pale band and a few scattered pale scales distad of band on

dorsal surface, bristles and scales in apical 0.4-0.5 of segment 5 largely pale

yellowish. Proboscis with prominent ventral tuft of several long hairs at base

of median light ring. Antenna slightly shorter than proboscis, flagellar whorls

strongly plumose.

MALE GENITALIA (Fig. 21). Segment IX. Tergal lobe with 4-8 strong

setae. Basimere: Slender, conical; about 0.32 mm in length. Subapical lobe.

Setae a-c of proximal division rather short, setae a,b stout; seta c very slender,

distad of a-b, all with hooked apices; setae in group d-f of distal division con-

sist of 1 narrow, lanceolate seta; 3 apically hooked, flattened, rodlike setae

and 1 small, hairlike seta, all placed mesad and distad of setae a-c of proximal

division; leaf g rather short, narrow and lanceolate, seta # strong, as long as

leaf g. Distimere. Typically sickle-shaped; apical 0.25 smooth, without crest

of spicules or serration; 1 tiny ventral and 1 dorsal setae present beyond mid-

dle, ventral distad of dorsal; subapical claw moderately long, apparently bi-

lobed. Phallosome. Apical spiculate portion of inner division of lateral plate

prominent, with apical sternal angle strongly produced sternad into a beak,

its length (measured from the posterior margin of inner division to apex of

beak) about 0.05-0.06 mm; apical tergal angle short, obtuse or slightly pro-

duced; apical margin usually smoothly curved, sometimes very lightly de-

pressed distad of apical tergal angle; inner tergal surface with a distinct lobe

bearing 2-3 budlike processes, situated far below apical tergal angle; outer

division in form of a simple and broad acuminate leaf which is variable in size;

lateral basal process distinct, knoblike. Pyroctiger. Apical crown medium

sized, composed of about 5 flat and blunt spicules and numerous spinelike spicu-

les; basal sternal process very short, slender and pale, 0.04 mm in length,

sometimes not developed or absent; cercal setae tiny, 3-4 in number.

PUPA (Fig. 21). Abdomen: 4.0mm. Paddle: 1.0mm. Trumpet: 0.9

mm. Cephalothorax and abdomen yellowish white; integument immediately

Sirivanakarn: Subgenus Culex in the Oriental Region 67

surrounding alveoli of all setae of abdomen, metanotum and sometimes also

cephalothorax usually very dark, producing a striking spotted pattern. Com-

plete chaetotaxy as figured, the following are characteristic. Trumpet. Asym-

metrically funnel-shaped and strongly bent upwards, apical margin of pinna

straight, about 0.3-0.5 mm in length. Cephalothorax. All setae weak, short,

some of which are rather indistinct; seta 1-C usually double (1-3); 2-C triple;

3-C 1-3 branched; 4-C 2-4 branched; 5-C 2-5 branched; 6-C triple; 7-C usually

double (2-3); 8-C 1-3 branched; 9-C usually double (1-2). Metanotum. Seta

10-C usually 7,8 branched (6-10); 11-C usually single (1-2); 12-C weak, very

short, 2-4 branched. Abdomen. Setae 3-I-III usually single, rarely double;

1-II single or forked distally into 4,5 branches; 6-I-VI all single; 6-V, VI very

strong and dark; 1-III 6-9 branched; 1-IV 4-6 branched; 1-V, VI and 9-IV, VI

subequal, dark, flattened, shorter than segments following and usually double

(2-3); 1-VII single or double; 9-VII flattened and dark, 3-5 branched; 4-VIII

usually double (1-3); 9-VIII 5-8 branched. Paddle. Very broad, semispheri-

cal; inner part darkened or infuscated; outer part pale; basal external margin

of outer part pale; midrib weak and pale; setae 1, 2-P present, very minute.

LARVA (Fig. 22). Head: 0.65 mm. Siphon: 2.0 mm, index 7-10. Sad-

dle: 0.45 mm; siphon/saddle ratio 4.4. Head. Relatively small; ocular bulge

not prominent; labral sclerite absent or not developed; anterior margin of fronto-

clypeus strongly emarginate or concave; seta 1-C pale, long and apically pointed,

its length as long as or slightly exceeding distance between bases of the pair;

3-C slightly laterad of 1-C; 4-C single or double, about as long as distance be-

tween bases of the pair; 5-C usually triple (2-4); 6-C double; 7-C 4-7 branched;

8-C triple; 9-C 4,5 branched; 10-C double; 11-C very distinct, usually single

(1-2); 12-C very weak, triple; 13-C single or double; 14-C single, filiform;

15-C double or triple. Antenna slender and relatively short, about 0.95 of head

length; pigmentation of antennal shaft variable from entirely pale to lightly

darkened from base to apex; spicules strong, widely scattered and more or less

restricted to basal 0.5, seta 1-A plumose, about 20 branched; 2, 3-A slightly

subapical or almost apical. Mental plate black, outer profile triangular and

with numerous delicate, closely packed lateral teeth on each side of a poorly

differentiated median tooth. Mouth brush filamentous and much less numerous

than most forms of Culex (Culex). Thorax. Not spiculate; seta 0-P weak,

pale, 10-15 branched; 1-3-P single; 4-P about 0.5 of the length of 1-3-P, usu-

ally single or double (1-3); 7-P usually triple (3-4); 8-P as long as 4-P, usually

double (1-3); 9, 10-P weak, short and usually double (1-2); 1-4-M weak and

short; 1-M single; 2-M 4 branched; 3-M single; 4-M 2-4 branched; 8, 9-M 4-8

branched; 1-4-T pale, indistinct; 2-T triple; 3-T 4 branched, 4-T 4,05 branched,

7-T usually 6,7 branched (5-8); 9-T usually 6 branched (5-6); 12-T single or

double; 13-T 4,5 branched. Abdomen: Not spiculate; most dorsal setae 1-4

and ventral setae 10-13 of segments I-VII very weak and inconspicuous; 4-I

short, weak and pale, 7-9 branched; 6-I, II strongly pectinate, usually triple

(3-4); 7-I single; 1-III-VI very short and weak, double or triple, 4-V pale, 5-8

branched; 4-VI pale, 3-5 branched; 6-III-VI strongly pectinate; 6-III 3-5 bran-

ched; 6-IV-VI usually triple (2-4); 1-VII double; 4-VII 2-4 branched; 7- VII

double; 10-VII 3-4 branched, 12-VII double; 13-VII usually triple. Comb scales

usually 4,5, sometimes 6,7 in single row; all large, subequal, with promin-

ent, long and simple apical spine and a broad basal sclerotized plate, fringe of

fine spicules restricted to base; seta 1-VIII usually 6 branched (6-7); 2 and 4-

VIII 2-4 branched; 3-VIII usually 6 branched (5-7); 5-VIII 4 branched. Saddle

complete; lateral posterior margin unspiculated; seta 1-X double or triple;

2-X usually triple (1-4); 4-K with 6 pairs of setae; anal gill strongly tracheate,

68 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

1-2.5 times as long as saddle. Siphon: Slender, long and distally tapered,

color yellowish white; pecten very poorly developed, usually with 3-5 indistinct

teeth (3-8); larger teeth with 2-3 denticles; siphonal tufts usually 4 pairs (total

8) in number, sometimes 3.5 pair (total 7), all weak, widely spaced, pairing

more or less regular, 2,3 branched each, inserted subventrally, slightly shorter

than siphonal width at point of attachment; seta 2-S dark, spiniform and rela-

tively long; median caudal filament of spiracular apparatus very well developed.

TYPE-DATA. (1) Culex bitaeniorhynchus Giles, Type (lost), Travancore,

(Madras State), INDIA (NE): (2) Taeniorhynchus ager Giles, Lectotype “* with

attached genitalia mount, Madvas, INDIA, Paddy field, 24 December 1899,

Capt. Cornwall (BM; selection of Bram 1967a: 263); (3) Culex kavatsuensis

Mochizuki, Type unspecified, Karatsu, Kyushu, JAPAN (LU).

DISTRIBUTION. Very widespread throughout the Oriental region with the

range extending north into the Palearctic (China, Japan, Korea and possibly

also in USSR), east and south into the Australasian region (New Guinea and

Northern Australia); Micronesia and the South Pacific and west into the Middle

East and the Ethiopian region. Material examined. 1,980 specimens: 712,

87292, 392 L; 516 individual rearings (199 pupal, 317 larval).

PAKISTAN. Lahore: Kahna Kacha; 59. |

INDIA. Bombay; Madras: Travancore (type locality); Assam: Chabua;

Dibrugarh; Jorhat; Dooma Dooma; Ledo; Lalmanirhat; Bengal: Calcutta; Dum

Dum; N.W. Prov.: Shahjahanpur; Andaman Is.; 350, 50%, 5 L, 2 p, 1 lp.

BANGLADESH. Sylhet: 69.

SRI LANKA. Central Province: Kandy District, Laksapana; Peradeniya;

North Central Province: Anuradhapura District, Padaviya, Irrigation Bungalow;

Sabaragamuwa Province: Ratnapura District, Panamure; Uva, P. Madulsima;

Suduganga; 180", 239.

THAILAND. Chiang Mai; Lampang; Mae Hong Son; Ubon Ratchathani; Udon

Thani; Nakhon Ratchiasima; Nakhon Nayok; Sara Buri; Pathum Thani; Nontha-

buri; Bangkok; Than Buri; Rat Buri; Kanchanaburi; Chon Buri; Rayong; Trat;

Prachuap Khiri Khan; Rangong; Surat Thani; Satun; Songkhla; Narathiwat; 361°,

S702) 250 Leite hip ist Ip.

VIETNAM. Saigon; Dalat; Danang; Phu Bei; Long Khanh; Qui Nhon; An Khe;

Cu Chi; Con Son; Quang Tinh; 160", 159, 14 L, 3p, 2 lp.

CAMBODIA. Ari Gsatr; 2%.

MALAYSIA. Peninsular Malaysia: Selangor: Kuala Lumpur; Rantau Pan-

jang; Seremban; Ulu Langat F. R.; Ulu Gombak; Pahang: Bukit Pakoh; Pdg.

Tungku K. Lipis; Kuantan; Pulau Tioman; Pevak: Kuala Kangsar; Lang Gunong;

Chenderiang; Perlis: Kg. To Kayaman; Penang Is.; Kelantan: Bertam; N.S.

(Negeri Sembilan): Bahau; Kg. Ulu Serting; Pedas; Johore: Kangkar Dohol;

Malaysia: Sabah: Mt. Kinabalu; Semporna; 83, 812, 34 L, 8p, 31 lp.

INDONESIA. Sumatra: Bengkulu; Java: Djakarta; Bogor; Solo; 9c, 109.

PHILIPPINES. Luzon: Subic Naval Base; Agoo La Union; Concepcion,

Tarlac; Olongapo; Camarines; Rizal, Camp Nichols, Mariguina, Wack Wack,

Camp Stotsenberg; Sapangbato; Pampanga; Dalton, N. Vizcaya, Bagiuo; Trini-

dad; Paranaque; Mindoro: San Jose; Palawan: Irahuan; Samar: Osmena:

Leyte: Gulf; Lago Lago, Bay Bay; Cebu: Toledo Rd.; Negros: La Carlota;

Mindanao: Kolambugan; Parang; Zamboanga; Torrey Barrack; Kabakan; Jolo

Jolo Is.; Tawi Tawi: Kamagong, Tarawakan; 1030, 1429, 24 L, 3 p, 60 lp.

TAIWAN. Ping Tung; Takeng; Nan Tou; 9°, 179, 17 L.

HONG KONG. 6, 139, 6p, 1 lp.

CHINA. Hainan; Peiping; Canton; Soo Chow; Chengtu; 80", 99.

JAPAN. Honshu: Saitama Sagiyama; Yodo; Kyoto; Tokyo; Okayama; Nara;

Sirivanakarn: Subgenus Culex in the Oriental Region 69

Nagasaki; 120°, 299, 13 L. Ryukyus: Okinawa; Iriomote; 520, 752, 24 L, 45 lp.

KOREA. * Seoul; Koje-do; Songdo; Chinhae; 100, 189, 11 L.

Additional records from the literature: BURMA: Mandalay; INDIA: Anda-

man Is. (Barraud 1924: 986); BANGLADESH (Aslamkhan 1971); SINGAPORE

(Colless 1959); IRAN: Khosh-hengam, Bandar-Abbas (Lotfi 1970: 399-403);

ETHIOPIAN REGION: Tanganyika; Uganda; Sudan; Kenya; Belgian Congo (Ed-

wards 1941: 290-1); USSR (Monchadskii 1951); INDONESIA: Sumba; Flores;

Alor; Timor; Ceram; New Guinea (Bonne-Wepster 1954: Assam and Bonne-

Wepster 1964); SOUTH PACIFIC: New Caledonia; AUSTRALIA: Northern

Territory; Queensland; New South Wales (Belkin 1962: 215-7); MICRONESIA:

Palau (Bohart 1956 (1957): 84-5).

TAXONOMIC DISCUSSION. Culex bitaeniorhynchus is the most common

and widespread species of the Bitaeniorhynchus Subgroup in Southeast Asia

and adjacent areas. The single type female of bitaeniorhynchus originally

described by Giles (1901b: 607), was lost. The only other type which is still

in existence and was apparently the one used by Edwards (1922) in referring to

typical bitaeniorhynchus is that of ager (Giles 1901). The latter nominal form

had been considered as one of the synonymys of this species (Edwards 1913b,

1922, 1932; Barraud 1934), but whether or not it is actually conspecific with

the true bilaentorhynchus can not be definitely determined in the absence of the

original type. In this study, Iam following Edwards (1922) in using the name

"bitaentorhynchus" for this form, but am excluding all other varietal names or

synonyms except ager (Giles) and kavatsuensis Mochizuki from the previous

synonymy.

The above description has been based on the study of material from the

Oriental region and from Japan, Korea and China in the eastern Palearctic.

Records of the species elsewhere may require confirmation in the light of

present taxonomic interpretation.

Culex bitaeniorhynchus can be distinguished from other members of its

subgroup in all stages by the following combination of characters. In the adults

by (1) anterior surface of femora, tibiae and tarsomeres 1 of all legs heavily

and extensively speckled with pale scales; (2) all wing veins with extensive

speckling of pale scales; (3) wing veins Ro, Rg, R45 with very broad squame

and plume scales; (4) abdominal terga II-VII with evenly broad apical pale

bands only and (5) dark scaled areas of all abdominal terga usually heavily

sprinkled with pale scales; in the male genitalia by (1) distal division of sub-

apical lobe of the basimere with a distinct lanceolate leaf in addition to leaf g;

(2) apical spiculate portion of phallosome prominent, typically long, beaklike

with more or less smoothly curved apical margin and with blunt or smoothly

rounded apical tergal angle; (3) basal sternal process of the proctiger usually

absent or poorly developed and unpigmented; in the pupa by (1) presence of

pattern of dark alveolar spots on all abdominal segments, metanotum and some-

times also on cephalothorax; (2) inner part of paddle darkened or lightly infus-

cate; (3) seta 11-C usually single; (4) setae 3-I-III usually single; in the larva

_ by (1) siphon usually with 4 pairs of subventral tufts; (2) comb scales large,

usually 4,5 in number; (3) setae 1-4 and 10-13 of segments I-VI usually very

weak, pale and rather indistinct; (4) seta 4-V usually 5-8 branched and (5)

seta 4-VI usually 3-5 branched.

The adults of bitaeniorhynchus exhibit much variation in the color of the

mesonotal scales in both sexes. Specimens normally have the scales on the

anterior 0.7 of mesonotum predominantly pale beige to white, contrasting

rather sharply with the dark scaled areas on the posterior 0.3. However,

these scales are sometimes predominantly dark with variable amount of pale

70 Contrib. Amer, But. last, vel. 12, no. 2; 1976

scales forming spots or streaks, particularly on posterior acrostichal and

dorsocentral lines cephalad of prescutellar space. In Southeast ASia the pale

form is predominant and has been found to be quite homogeneous, especially

in the northern parts of its range, including China, Japan and Korea.

The typical bitaeniorhynchus is closely related to infula, luzonensis,

selangorensis, pseudosinensis and longicornis, with which it overlaps in one

or more stages, and apparently falls into a distinct complex. Within Southeast

Asia (except the Ryukyus), it has been frequently found generally or closely

sympatric with every member of this complex. In southern and central Thai-

land, Malaysia, the majority of bitaeniorhynchus specimens are usually found

in the same habitats as the dark and pale forms of zmfula but whether or not it

may occasionally hybridize with the latter species is unknown.

BIONOMICS. Culex bitaeniorhynchus is a rural species and is more or

less restricted to breeding in large, semi-permanent or permanent ground

pools such as swamps, rice fields, marshy depressions, irrigation ditches,

ponds, logs and obstructed streams. On several occasions, the pupae and

larvae have also been collected from puddles, foot prints and small ground

pools at margins of swamps, streams, rivers and canals. These breeding

sites are under shade or fully exposed to sunlight, in or near forest, in plains

or mountainous areas at a broad range of elevation. The water is usually

fresh and clear and always filled with a dense mass of filamentous green algae.

The larvae are dependent on algae as food and shelter. Mohan (1950), ina

study of a laboratory colony of bitaeniorhynchus (type form) reported that the

presence of green algae was absolutely essential for the normal development

and survival of the larvae and in inducing females to deposit eggs. In Thailand,

Vietnam and Peninsular Malaysia, adults of both sexes have been caught in

number in light traps and several females have been taken biting or landing on

man. The females apparently feed on blood of a variety of hosts, including

man, cows, buffalo, pigs, dogs and chickens (Colless 1959; Reuben 1971b;

Sasa and Sabin 1950) and rabbits (Mohan 1950). In Madras, India (Christopher

and Reuben 1971), Singapore (Colless 1959) and in Japan (Sasa and Sabin 1950),

it has been reported to feed amost exclusively on birds and rarely on other

animals, including man. Further study on the local differences in the blood

feeding habits of bDitaeniorhynchus females is still needed, especially in South-

east Asia where, as pointed out by Colless (1959) and Sirivanakarn (1973),

other closely similar forms are also present, including those which are pre-

sently recognized as infula.

MEDICAL IMPORTANCE. In the Oriental region, bitaeniorhynchus has

been reported to be naturally infected with Wuchereria bancrofti in Travan-

core, India (Iyengar 1938) and with Brugia malayi in Sri Lanka (Carter 1948).

In the Berau region, Netherlands New Guinea, it has been shown to be highly

susceptible to Wuchereria bancrofti in the laboratory and has been considered

to be an efficient vector of this pathogen (de Rook 1957b; de Rook and van Dijk

1959). In Malaya, Williamson and Zain (1937) reported laboratory infection

and development of sporozoites of Plasmodium vivax (Grassi and Feletti), but

Russell and Mohan (Mohan 1950) in India found it to be extremely resistant to

this malarial parasite. In the Philippines and Queensland, Australia, a strain

of Sindbis virus was isolated from bitaeniorhynchus (Rudnick et al. 1962;

Doherty et al. 1963) and in Korea it has been incriminated as a possible vector

of Japanese encephalitis. The latter report has never been verified or con-

firmed, however, as it feeds principally on birds which are not the natural

reservoir of this virus, it is probably not significant as a vector of this disease.

Sirivanakarn: Subgenus Culex in the Oriental Region 71

9. CULEX (CULEX) INFULA THEOBALD

(Figs. 23, 24, 25)

Culex infula Theobald 1901a: 370 (2*); Giles 1902: 407 (2); Blanchard 1905: 298

(2); Leicester 1908: 146 (2); Edwards 1913b: 231 (synonymy with bifaeni-

orhynchus Giles).

Taeniorhynchus tenax Theobald 1901b: 198 (2*); Giles 1902: 365 (2); Theobald

1903: 258 (2*); Leicester 1908: 167 (2); Edwards 1913b: 231 (synonymy

with bitaeniorhynchus Giles); Edwards 1922a: 283 (“*, °, as variety of

bitaeniorhynchus); Barraud 1924a: 985 (o*, ?, as variety of bitaenior-

hynchus); Barraud 1934: 393 (2, L, as variety of bitaeniorhynchus).

Grabhamia ambiguus Theobald 1903: 248 (); Edwards 1913b: 231 (synonymy

with bitaeniorhynchus Giles); Edwards 1922a: 283 (as variety of bitaenior-

hynchus); Barraud 1924a: 985 (adult, as variety of bitaeniorhynchus);

Baisas 1938: 211 (adult, as variety of bitaeniorhynchus).

Grabhamia taeniarostris Theobald 1907: 299 (2); Edwards 1913b: 231 (synonymy

with bitaeniorhynchus Giles).

Taeniorhynchus tenax. var. ocellata Theobald 1907: 488 (2); Edwards 1913b: 231

(synonymy with bitaeniorhynchus Giles).

Oculeomyia sarawaki Theobald 1907: 515 (2*); Edwards 1913b: 231 (synonymy

with bitaeniorhynchus Giles).

Taeniorhynchus domesticus Leicester 1908: 169 (“, 2); Edwards 1922a: 283

(as variety of bitaeniorhynchus); Barraud 1924a: 984 (synonymy with

bitaeniorhynchus Giles).

Culex (Culex) afridii Qutubuddin 1956: 140 (**). NEW SYNONYMY.

Culex bitaeniorhynchus Giles (in part), Edwards 1921: 337 (adult, L); Edwards

1922a: 282 (““*, 2); Barraud 1924a: 984 (c'*, 9).

Culex (Culex) bitaenorhynchus Giles (in part), Edwards 1932: 202 (taxonomy);

Barraud 1934: 391 .(¢%*,~9,. L*)s; Delfinado 1966: 140 (o%*). 2, BP,-L*);

Bram 19670258 (7 %,> Oty) P* 215%) Sirivanakarn 1973; 235: (o*; 2h oP;

L, tenax and domesticus forms of bitaeniorhynchus).

Culex (Culex) aurantapex of Edwards (in part), Edwards 1932: 202 (taxonomy);

Stone, Knight and Starcke 1959: 242 (catalog).

FEMALE (Fig. 23). In general very similar to bitaeniorhynchus from

which it is distinguished by the following characters. Head. Narrow decumbent

scales of vertex yellowish to dark brown; erect scales usually entirely dark or

sometimes pale golden in center, dark posterolaterally. Czbarial Armature.

As figured and as described for bitaeniorhynchus. Thorax, Anterior 0.7 of

mesonotum varying from predominantly pale beige, yellow or golden (tenax

form, Sirivanakarn 1973) to predominantly deep chestnut brown or black

scaled (domesticus form, Sirivanakarn 1973), with or without dark and pale

scales forming streaks, spots or blotches along acrostichal line, middle of

fossa, areas cephalad of prescutellar space at level of anterior wing base or

elsewhere; posterior 0.3 of mesonotum usually entirely dark brown scaled;

scales on prescutellar space and scutellar lobe completely dark brown or

partially pale as in bitaeniorhynchus. Legs. Anterior surface of all femora

usually lightly to moderately speckled, sometimes heavily and extensively

speckled; tibiae lightly speckled, tarsomere 1 of all legs completely dark or

very lightly speckled. Wing. Lightly to moderately speckled, usually restrict-

ed to basal 0.5-0.75 of veins C, Sc, Rand Cu, sometimes heavily and exten-

sively speckled; plume scales and squamae of veins Ro, Rg and R45 broad,

72 Contrib. Amer..Ent. Inst., vol. 12, no. 2, 1976

clavate, but relatively narrower than in bitaeniorhynchus. Abdomen. Median

caudal scale patch of tergum I completely dark or partially pale; terga I-IV or

IIl-V largely dark or with narrow apical pale bands which are usually connected

with larger yellowish apicolateral patches at sides; basal pale bands usually

absent, if present, narrow or broad, complete or incomplete or represented by

a small median basal spot or streak; terga VI-VII or V-VII always with distinct

apical pale bands which vary from 0.2-0.5 of segment length; basal pale bands

present or represented by median spot or completely absent; tergum VIII with

broad apical and basal pale bands or completely yellowish. Genitalia. Essen-

tially as described and figured for bitaeniorhynchus.

MALE (Fig. 23). In general as described for female; sexuai dimorphism

usually developed, particularly in the following features. Abdomen. Terga

II- VII usually with complete basal pale bands or sometimes with median basal

pale spots; apical bands usually present, narrow, sometimes absent; basal

bands, if complete, usually broader than apical.

MALE GENITALIA (Fig. 24). Extremely similar to bitaeniorhynchus,

differing particularly in the following. Phallosome. Apical spiculate portion

of inner division of phallosome more strongly pigmented, apical margin (in

lateral view) with distinct emargination proximad of apical tergal angle; apical

tergal angle usually produced into a point. Proctiger. Basal sternal process

usually present and relatively stronger, varying from 0.04-0.05 mm in length.

PUPA. Essentially as figured for bitaeniorhynchus; cephalothorax and ab-

domen variable from generally pale yellowish white with pattern of dark alveolar

spots to deep yellow or golden without pattern of dark alveolar spots; abdominal

segments V-VIII with or without pairs of dark brown lateral spots and dark me-

dian basal spots; sometimes cephalothorax and abdomen uniformly dark brown-

ish. Details of chaetotaxy essentially similar to bitaeniorhynchus, differing

particularly in the following. Metanotum: Seta 11-C usually double (1-3).

Abdomen. Setae 3-I-III double, rarely single. Paddle. Inner and outer parts

of paddle usually entirely pale or transparent, sometimes inner part lightly

darkened; external margin in basal 0.5 of outer part darkened or infuscate,

sometimes pale.

LARVA (Fig. 25). Extremely similar to typical bitaeniorhynchus, differing

in the following combination of characters. Head. Seta 4-C usually 3-4 bran-

ched (2-4); 5-C triple or 4 branched; setae 11 and 13-C usually double or triple,

rarely single. Thorax. Seta 0-P usually dark, distinct, 15-20 branched; 4-P

double or triple (2-4); 1-4-M_ usually dark and distinct; 3-M usually double or

triple, rarely single; 4-M usually 4,5 branched (3-5); 1-4-T usually dark and

distinct; 2-5 branched; 3-T 4-6 branched; 12-T at least double (2-4); 13-T

usually 6 branched (4-9). Abdomen. Most dorsal setae 1-4 and ventral setae

10-13 of segments I-VII usually very dark and distinct, rarely pale; 3-I usually

4 branched (3-5); 4-I, II dark, 10-16 branched; 1-III-VI usually 3, 4 branched,

1-V sometimes 5 branched; 1-VII usually 5 branched (3-6); 4-V usually 10-12

branched (9-14); 4-VI usually 6,7 branched (5-8). Comb scales usually 9, 6 in

number (4-7). Saddle seta 2-X usually 4 branched (3-5). Siphon: Subventral

tufts 3 pairs (total 6) in number, sometimes 2.0 pairs.

TYPE-DATA. (1) Culex infula Theobald, Holotype 2*, Taipang, Straits

Settlements, Perak, MALAYA (PENINSULAR MALAYSIA), L. Wray, Jr.

(BM); (2) Taeniorhynchus tenax Theobald, Holotype 2*, Taipang, Straits Settle-

ments, Perak, MALAYA (PENINSULAR MALAYSIA), L. Wray, Jr. (BM); (3)

Grabhamia ambiguus Theobald, Holotype “* with attached genitalia mount,

Quilon, Travancore, (Madras State), INDIA. Capt. James (BM); (4) Grab-

hamia taeniarostris Theobald, Holotype 2?*, Peradeniya, CEYLON (SRI LANKA),

Sirivanakarn: Subgenus Culex in the Oriental Region 73

January 1900, Green (BM); (5) Taeniorhynchus tenax var. ocellata Theobald,

Holotype °*, Kuching, Sarawak, BORNEO (BM); (6) Oculeomyia sarawaki

Theobald, Holotype ¢* (in extremely poor condition), Savawak, BORNEO, Dr.

Barker (BM). (7) Taeniorhynchus domesticus Leicester, Lectotype “* with

slide of genitalia, Fed. Malay State (PENINSULAR MALAYSIA), G. F. Lei-

cester (BM 1912: 350; selection of Bram 1967a: 263); (8) Culex (C.) afridti

Qutubuddin, Holotype “* with slide of genitalia, Dacca, EAST PAKISTAN

(BANGLADESH), October 1952, M. Qutubuddin (BM).

DISTRIBUTION. Thailand, Malaysia, Vietnam, Burma, Bangladesh,

Indonesia and the Philippines. Also reported from India, Sri Lanka and

Micronesia (Palau).

Material examined: 847 specimens: 3490", 3982, 102 L; 169 individual

rearings (63 pupal, 106 larval).

INDIA. Bombay: Kawar, N. Kanara; Deccan, Belgaum; Nagargali;

Assam; 10%, 108.

BANGLADESH. Dacca; 1% (as afridii M. Qutubuddin, 1951); Sylhet; 1c,

Bie Ce

SRI LANKA. Central Province: Kandy District, Laksapana; Suduganga;

OW 193

BURMA: Rangoon) 1069.05 9,

THAILAND. Chiang Mai; Lampang; Tak; Bangkok; Pathum Thani;

Chanthaburi; Chon Buri; Nakhon Si Thammarat; Prachuap Khiri Khan; Trang;

Narathiwat; 41%, 472, 29 L, 16 p, 22 lp.

VIETNAM. Danang; Saigon; Pleiku; Phu Bei; Qui Nhon; Dalat; 150", 419,

4 Ei Bip, Or Eps

MALAYSIA. Peninsular Malaysia: Selangor: Kuala Lumpur; Segambut;

Puchong; Rantau Panjang; Pacific Tin, Kuala Selangor; Ulu Gombak; Bt. Kutu;

Trengganu: Dungun; Marang; Kuala Brang; Pahang: Bukit Pakoh; Pdg. Tungu,

K. Lipis; Kuantan; Temerloh; Gunong Benom; Perak: Tanjong Tulang F. R.,

Kuala Kangsar; Chior F. R.; Perlis: Kg. Gunong; Penang Is. ; Kelantan: Gua

Musang; Tumpat; Malacca: Malacca; Kedah: Kg. Sungei Lintang; N. S.: Ran-

tau; Kg. Inas; Pedas; Kg. Sengala; Johore: Kangkar Dohol; 121“, 1389, 50 L,

40 p, 51 1p; Malaysia: Sabah: Kota Belud; Beaufort; Keningau; Tambunan;

Kuala Papar; Tawau; Kinarut; Mt. Kinabalu; Semporna; Lahad Datu; Kuala

Penyu; 190°, 302, 4 lp.

INDONESIA. Kalimantan (Borneo): Tarakan; 6, 32; Sumatra: Kmp Pantai;

Bengkulu; Kebonros; Java: Djakarta; Solo; Maur; Bogor; Garut; 380", 319.

PHILIPPINES. Luzon: Concepcion, Tarlac; Agoo La Union; Dalton, N.

Vizcaya, Bagiuo; Laguna; Rizal, Camp Nichols, Wack Wack, Paranaque;

Abatan, Buguias; Subic Bay; Olongapo; Calaccad; Siasi Sulo: Calon; Mindoro:

San Jose; Samar: Osmena; Leyte: Rizal; Palo; Lago Lago Bay Bay; Dulag;

Loreto Dinagut; Tacloban; Panamuan; Gulf; Jinamoc; Negros: Cadiz; Mindanao:

Pangasinan; Camp Gregg; San Ramon; Kabakan; Parang; Kolambugan; Ft. Pikit;

Tawi Tawi: Kamagong, Tarawakan; 930", 902, 19 L,. 20 lp.

TAXONOMIC DISCUSSION. The present treatment of the specific status of

infula is only provisional, pending a more thorough analysis of the material

from progeny rearing. In this interpretation, the 2 nominal forms, tenax

Theobald 1901 and domesticus Leicester 1908, which were earlier recognized

by Sirivanakarn (1973: 235-51) as distinct forms of bitaeniorhynchus and other

names, including ambiguus Theobald 1903, taeniarostris Theobald 1907, tenax

var. ocellata Theobald 1907, savawaki Theobald 1907 and infula Theobald 1901

are all combined and tentatively treated as a single species. The name infula

is used because it has priority over all other names. This interpretation is

74 Contrib. Amer. Ent, Inst., vol. 12, no. 2,: 1976

based largely on the study of several series of reared specimens from Malaysia

where most of these nominal forms were originally described. In spite of the

striking differences in the color of the mesonotal scales and the pattern of ab-

dominal banding of the adults as described above and as indicated by Sirivana-

karn (1973), subsequent study has shown that there is apparently no difference

in the male genitalia, pupal and larval chaetotaxy among all of the Malaysian

forms, indicating that they are probably conspecific. In Addition, afridii

Qutubuddin 1956 from East Pakistan which was described from the single type

male is also synonymized with this species on the basis of similarity in the

male genitalia and external characters of the adult.

Culex infula can be distinguished from other forms of the bitaeniorhynchus

complex by the following combination of characters. In the adults by (1) usually

light or moderate speckling of the legs and wing; (2) scales on wing veins Ro,

Rg and Ry,5 narrow, clavate; (3) terga II-IV or II-V in females usually with

distinct apicolateral yellowish patches not extending dorsad to form distinct

apical band; basal bands usually absent or sometimes represented by a small

median basal pale spot; (4) abdominal terga V-VII or VI-VII usually with dis-

tinct apical and basal bands; (5) abdominal terga VIII with apical and basal bands

or completely yellowish; (6) male abdominal terga II-VII usually with both apical

and basal bands. In the male genitalia by (1) long beak-like apical spiculate por-

tion of inner division of the phallosome; (2) apical spiculate portion with distinct

depression on apical margin and obtuse apical tergal angle; (3) basal sternal

process of the proctiger usually developed, varying from 0.04-0.05 mm in

length and (4) one of the setae d-f in distal division of the subapical lobe usually

differentiated as a narrow lanceolate leaf. In the pupa by (1) seta 11-C usually

double; (2) setae 3-I-III usually double and (3) inner part of paddle pale or not

infuscate. In the larva by (1) setae 1-4-T dark and distinct; (2) setae 1-4 and

10-13 of segments I-VII usually dark and very distinct; (3) 4-V 10-12 branched;

4-VI usually 6,7 branched; (4) comb scales 4,5 in number and (5) siphon usually

with 3 pairs (total 6) of subventral tufts.

Culex infula is extremely variable and exhibits overlap in one or more

stages with the typical bitaeniorhynchus and other members of the bitaeniorhyn-

chus complex. In the adults, there is a marked dimorphism into the pale and

dark forms in both sexes in the color of the head and the mesonotum. The pale

form (tenax) has the erect scales of the head partially pale and the scales on

the anterior 0.7 of mesonotum predominantly pale, contrasting sharply with

those on posterior 0.3. The dark form domesticus, infula) is strikingly

different from the pale one in having the scales on the head and mesonotum en-

tirely dark brown to almost black and can be readily recognized even by the

naked eye. In Malaysia, Thailand, Vietnam, Indonesia and the Philippines,

the dark form is apparently more common than the pale form. There is alsoa

marked sexual dimorphism in the pattern of the abdominal banding; the males

usually have both apical and basal pale bands whereas the females have only

the apical bands, particularly on terga II-IV or II-V. In the immature stages,

the infula pupae exhibit considerable variation in the color of cephalothorax

and abdomen, overlapping with the typical biiaeniorhynchus and pseudosinensis

but resemble the latter in the diagnostic chaetotaxy. The larvae resemble

pseudosinensis, luzonensis and selangorensis in having 3 pairs of siphonal tufts,

but differ from these species and the typical bitaeniorhynchus in having dorsal

thoracic setae and most dorsal and ventral abdominal setae relatively stronger,

darker and with more numerous branches.

BIONOMICS. Culex infula is apparently as common as the typical bitaenior-

hynchus in various parts of Southeast Asia and as indicated for the latter, it is

Sirivanakarn: Subgenus Culex in the Oriental Region 79

restricted to breeding in large, algae filled ground pools. In the majority of

collections, the larvae and pupae of znfula have been found with bitaeniorhynchus

in the same breeding sites. In Malaya (Peninsular Malaysia), it was reported

to represent roughly 55 per cent of the bitaeniorhynchus populations (Sirivana-

karn 1973). Of this, the dark form Q@omesticus) is apparently more common

than the pale form (fenax). A number of adults of infula have been caught in

light traps and biting man or other animals. The blood feeding habits and the

preferred host(s) of zzfula females are not definitely known. It would be of

interest to determine whether this particular taxon shows a preference in

blood feeding habit different from that of typical bitaeniorhynchus.

10. CULEX (CULEX) LUZONENSIS NEW SPECIES

(Figs. 23, 24)

Culex (Culex) bitaeniorhynchus Giles (in part), Sirivanakarn 1973: 235 (o*, 9,

P, Iu; ‘as: Luzon form).

FEMALE (Fig. 23). Wing: 4.7mm. Forefemur: 2.2 mm. Proboscis:

2.0mm. Abdomen: 3.0mm. As briefly characterized by Sirivanakarn

(1973: 244) for the Luzon form of bitaeniorhynchus, with the following correc-

tions and additional features. Head. Erect scales of vertex vary from par-

tially pale yellowish or golden in center, dark on posterolateral area to com-

pletely dark brown. Palpus largely dark scaled; apex of segment 4 with or

without a few pale scales. Proboscis apparently slightly shorter than fore-

femur. Thorax. Anterior 0.7 of mesonotum usually predominantly dark

brown scaled and with extensive mottling of pale yellowish or golden brown

scales in middle cephalad of prescutellar space; posterior 0.3 of mesonotum

blackish; scales on prescutellar space largely dark except for some pale ones

posteriorly; mid-scutellar lobe largely dark scaled except for some pale ones

on caudal margin; lateral scutellar lobe entirely dark scaled. Legs. Anterior

surface of femora and tibiae of fore- and midlegs usually moderately to heavily

speckled, sometimes almost completely dark; anterior surface of hindfemur

lightly speckled; hindtibia speckled as in fore- and midlegs; tarsomere 1 of all

legs usually lightly speckled, sometimes entirely dark. Wing. Usually lightly

speckled, particularly in basal parts of veins C, Sc, Rand stems of other veins;

scales on veins R2, Rg and R4,s5, narrow, linear or broad clavate. Abdomen.

Median caudal scale patch of tergum I largely dark or partially pale towards

base; terga JI-VII with broad incomplete or complete basal triangular pale

bands and narrow apical bands, the latter connected with very large apicolater-

al yellowish patches on terga III-V or III- VI; basal and apical bands on terga

VI-VII sometimes equally broad; tergum VIII usually with broad basal and nar-

row apical pale bands; sterna III-VI largely yellowish, sternum VII broadly

pale at base, dark towards apex.

MALE (Fig. 23). In general as described for female, differing in having

scales on anterior 0.7 of mesonotum predominantly pale, yellowish to golden

with variable amount of dark scales forming spots or streaks and in having

banding pattern on abdominal terga II-V largely basal and complete. Palpus

and proboscis essentially similar to typical bitaeniorhynchus and infula,

MALE GENITALIA (Fig. 24). As figured and described by Sirivanakarn

(1973: 250); very similar to other members of the bitaeniorhynchus complex

in the basimere, distimere and proctiger, differing particularly in the follow-

ing. Subapical lobe. None of setae in group d-fof distal division is differen-

76 Contrib.::Amer.; Ent: Inst.; .vols.12)/no..2)° 1976

tiated as a broad, lanceolate leaf. Phallosome. Apical spiculate portion of

inner division very stout, more or less quadrate in shape and dark pigmented,

apical margin smoothly curved, apical sternal angle produced into a short and

stout beak which is about 0.04 mm in length; apical tergal angle irregular,

usually provided with several denticles; tergal lobe with 2-3 broad, blunt tipped

foliform processes, placed at a distance far below apical tergal angle. Pvoc-

tiger. Basal sternal process of paraproct relatively longer, thicker and darker,

about 0.08 mm in length; cercal setae vary from 2 to 5 in number.

PUPA. As described and figured for typical bitaeniorhynchus, differing

from it particularly in the following. Cephalothorax and abdomen uniformly

yellowish; pattern of dark alveolar spots entirely absent. Metanotum., Seta

11-C usually double, rarely single. Abdomen. Setae 3-I-III double; setae

1-VI-VIII usually single (1-2). Paddle. Outer and inner parts entirely pale,

midrib more pigmented.

LARVA. As described and figured for typical bzfaeniorhynchus to which it

is extremely similar, differing particularly in the following. Head. Basal

0.5 of antennal shaft with considerably weaker and fewer spicules, sometimes

without distinct spicules. Abdomen. Comb scales smaller, shorter and with

distinct basal fringe of spicules; number of scales varies from 5 to 9 (average

6), inl or 2 irregular rows. Siphon: Subventral tufts usually 3 pairs (total 6),

sometimes 3.5 pairs (or total 7); pairing very irregular.

TYPE-DATA. Holotype “ (D-IX-33) with associated pupal skin and genitalia

slide, Imugan, Dalton, N. Vizcaya, Luzon, PHILIPPINES, elevation 925 m,

larva from a large carabao wallow, 12 Jan. 1969, Celestino (USNM); Allotype 2

(D-IX-41) with associated pupal and larval skins, same data as holotype (USNM);

Paratypes: 8% (D-IX-35, D-IX-40, D-IX-44, D-IX-47, D-IX-48, D-IX-49, D-IX-

00, D-IX-51) and 9° (D-IK-34, D-IX-38, D-IX-39, D-IX-43, D-IX-45, D-IX-46,

D-IX-52, D-IX-54, D-IX-55) with associated pupal and larval skins; 2% (D-IX-

30), D-IX-32) and 22 (D-IX-29, D-IX-31) with associated pupal skins; same

data as holotype (USNM, BM, BBM).

DISTRIBUTION. Known only from Luzon in the Philippines. Material

examined. 48 specimens: 200, 249, 4 L; 32 individual rearings (7 pupal,

25 larval).

PHILIPPINES. Luzon: N. Vizcaya, Dalton, Imugan (type-locality); 14°,

192, 7 P, 25 1p; Bagiuo, Mountain Province; 5c, 52, 4 L; Subic Naval Base; 1c.

TAXONOMIC DISCUSSION. Culex luzonensis is apparently restricted to

mountainous areas of Luzon in the Philippines where it has been found to be

sympatric with the typical bitaeniorhynchus. It can be separated from the

latter species and other members of the bitaeniorhynchus complex by the

following combination of characters: in the adults by (1) presence of light to

moderate speckling of the legs and wing and (2) presence of narrow apical and

broad basal pale bands on abdominal terga II-VII; in the male genitalia by (1)

characteristic shape of apical spiculate portion of inner division of the phallo-

some and (2) basal sternal process of the proctiger relatively thick, strongly

pigmented and about 0.08 mm in length; in the pupa by (1) absence of pattern

of dark alveolar spots on the abdomen and cephalothorax; (2) paddle largely

pale, without darkened or infuscate area; (3) seta 11-C double and (4) setae

3-I-III double and in the larva by (1) antennal shaft with relatively few and

weak or indistinct spicules; (2) setae 1-5 and 9-13 of abdominal segments I-VI

very weak and indistinct (3) comb scales 5-9 in number and (4) siphon with 3

pairs (total 6) of subventral tufts.

Except for the diagnostic male genitalic characters, all stages of luzonen-

sis exhibit much overlap with infula and as in the latter species, the adults of

Sirivanakarn: Subgenus Culex in the Oriental Region 77

luzonensis also show a marked dimorphism in the pale or dark scales on the

anterior 0.7 of the mesonotum. The associated larvae and pupae of both dark

and pale forms are virtually similar and there is no apparent indication of cor-

related differences in any features among several reared specimens I have ex-

amined. For a definite separation of luzonensis from infula, all stages, includ-

ing the male genitalia should be thoroughly examined.

BIONOMICS. The larvae and pupae of luzonensis from the type locality at

Dalton, N. Vizcaya were collected from a large carabao wallow containing some

ereen algae and other aquatic vegetation at an altitude of 925 m. This breeding

site was under partial shade and was noted to be ina native village. Several

adults have been obtained from general field catches in Bagiuo and a few fe-

males were found to be fully engorged with blood but no data is available as to

their specific host. Nothing is known about the biology of the adult and its re-

lation to disease.

11. CULEX (CULEX) SELANGORENSIS NEW SPECIES

(Pigs. 23, 24)

Culex (Culex) bitaeniorhynchus Giles (in part), Sirivanakarn 1973: 235 (o*, &,

L, as Selangor form)

FEMALE (Fig. 23). Wing: 4.3mm. Forefemur: 2.2 mm. Proboscis:

2.0mm. Abdomen: 3.24 mm. As characterized for the Selangor form of

bitaeniorhynchus by Sirivanakarn (1973: 243); distinguished from typical bitae-

niorhynchus and other members of the bitaeniorhynchus complex and subgroup

by the absence of basal or apical pale bands on abdominal terga II- VII and by the

following additional characters. Head, Narrow decumbent scales of vertex

beige or yellowish; erect scales yellowish in center, dark posterolaterally.

Palpus entirely dark scaled. Thorax. Anterior 0.7 of mesonotum always pre-

dominantly pale yellowish or golden scaled, with or without dark scaled streaks

and spots on anterior acrostichal line and in middle of fossa; scales on posterior

0.3 of mesonotum, prescutellar space and scutellar lobes entirely dark brown.

Legs. Anterior surface of femora and tibiae usually lightly sprinkled with pale

scales, sometimes almost completely dark; tarsomere I of all legs entirely

dark. Wing. Scales on all wing veins entirely dark; speckling of pale scales

absent; plume scales and squames on veins Ro, Rg and Ry,5 narrow, clavate.

Abdomen. Tergum I with dark median caudal scale patch; terga II- VII almost

completely dark except for very light sprinkling of a few pale scales, particu-

larly on terga III-VI; basolateral pale spots present, particularly on terga III-

VII; tergum VIII with narrow basal pale band or completely dark; sterna III- VII

with broad basal yellowish bands; sternum VIII entirely dark.

MALE (Fig. 23). In general similar to female except for the presence of

broad median pale streaks or incomplete basal pale bands on abdominal terga II-

IV and complete basal pale bands on abdominal terga V-VII. Palpus and pro-

boscis as in bitaeniorhynchus and infula except for the following. Head. Ven-

trolateral bristles in apical 0.5 of palpal segment 3 weaker and less dense,

about 20 in number; upper mesal surface of palpal segment 3 with a row of only

> long hairs which are more widely spaced. Proboscis with a considerably

weaker ventral tuft of long hairs proximad of median light ring; the number of

hairs is about 10.

MALE GENITALIA (Fig. 24). As figured and described by Sirivanakarn

(1973: 250); extremely similar to bitaeniorhynchus and infula, differing particu-

78 Contrib.: Amer. Ent. Inst,, vols2,/no.2;- 1976

larly in the following features. Subapical lobe. None of the setae in group d-f

of distal division differentiated as a lanceolate seta; leaf g shorter and narrower.

Phallosome. Apical spiculate portion of inner division, quadrate or somewhat

resembling the blade of an ax; apical sternal angle produced into a short beak

which is 0.012-0.030 mm in length; apical tergal angle irregular and usually

with some coarse denticles; apical margin smoothly curved; tergal lobe with

3-4 short and pointed foliform processes situated close to apical tergal angle.

Proctigery. Basal sternal process short, 0.03-0.04 mm in length and strongly

pigmented.

PUPA. Extremely similar to bitaeniorhynchus and infula, differing in the

following combination of characters. Cephalothorax and abdomen yellowish

white; pattern of dark alveolar spots present, particularly on metanotum and

abdominal segments, but less striking than in typical bitaeniorhynchus. Met-

anotum. Seta 11-C double or triple. Abdomen. Seta 3-I usually double, some-

times triple; 3-II-III always double; 1-V usually 4,5 branched (3-5); 5-IV-V

usually 5 branched (4-6); 5-VI usually 4 branched (3-5). Paddle. Inner part

lightly darkened, as in bitaeniorhynchus; outer part pale, basal external mar-

gin pale.

LARVA. Essentially as figured for pseudosinensis (Fig. 27) from which it

is virtually indistinguishable, differing from bitaeniorhynchus, infula and

luzonensis particularly in the presence of 8-11 much smaller comb scales in

2 or 3 rows and in the following additional characters. Head, Seta 14-C

single, sometimes bifid. Thovax. Seta 4-P single or double; seta 1-4-M and

seta 1-5-T extremely weak and inconspicuous. Abdomen. Most dorsal setae

1-4 and ventral setae 10-13 of segments I-VII extremely weak, short and very

inconspicuous; setae 1-IJI-VII usually double, sometimes triple; seta 4-V 5, 6

branched; 4-VI 4 branched. Comb scales, usually 9 in number (8-11), in 2 or

3 rows, all small, with distinct apical spine and basal fringe of several fine

spicules. Seta 2-X of saddle usually 4 branched (4-5). Siphon. Basal 0.1

always with a distinct row of 7-8 pecten teeth; subventral tufts 3 pairs (total

6) in number; pairing regular.

TYPE-DATA. Holotype “ (1386-14) with associated pupal and larval skins

and genitalia slide, Bukit (Bt.) Kutu, Selangor, PENINSULAR MALAYSIA,

elevation 310 m, larva from stagnant pool along stream margin, 15 June 1968,

S. W. James and others (USNM); Allotype ¢ (1386-12) with associated pupal and

larval skins, same data as holotype (USNM); Paratypes: 1° (1386-13) with

associated pupal and larval skins and genitalia slide; 32 (1386-10, -11, -15)

with associated pupal and larval skins, same data as holotype (USNM).

DISTRIBUTION. Presently known only from Selangor and Pahang, Peninsu-

lar Malaysia, Malaysia. Material examined. 13 specimens: 5c, 62, 2L, 7

individual rearings (1 pupal, 6 larval).

MALAYSIA. Peninsular. Malaysia: Selangor-Bukit (Bt.) Kutu; 3c", 49,. 6 lp

(type-series). Pahang: Gunong Benom; 2, 2°, 2L, 1p.

TAXONOMIC DISCUSSION. Culex selangorensis is most distinctive in the

females which differ from the rest of Bitaeniorhynchus Subgroup in the absence

of apical or basal bands on abdominal terga II-VII. The male is sexually dimor-

phic, differing from the female in the presence of incomplete or complete basal

pale bands on the abdominal terga. In the male genitalia, the phallosome is

distinct from typical bitaeniorhynchus, infula and luzonensis but is slightly dif-

ferent from that of pseudosinensis in the shape of the apical spiculate portion

of the inner division. The pupa resembles typical bitaeniorhynchus rather

closely in pigmentation except for less striking pattern of dark alveolar spots

and diagnostic chaetotaxy. The larva is virtually indistinguishable from pseu-

Sirivanakarn: Subgenus Culex in the Oriental Region 79

dosinensis.

BIONOMICS. All specimens of selangorensis came from 2 collections made

in stagnant ground pools with scarce green algae at the elevation of 185-330 m

in Selangor and Pahang, Peninsular Malaysia. The immatures from the 2 lo-

calities were not found to be associated with any other forms of the Bzlaenior-

hynchus Subgroup. Nothing is known about the adult biology or its medical im-

portance.

12; CULEX (CULEX) PSEUDOSINENSIS COLLESS

(Pigs; 26;..2.7)

Culex (Culex) pseudosinensis Colless 1955: 311 (o*, 2, L*); Bram 1967a: 265

(SA peltyn bX):

FEMALE (Fig. 26). Wing: 4.5mm. Forefemur: 2.0 mm. Proboscis:

2.1mm. Abdomen: 3.4mm. General facies as figured; distinguished from

other members of the bitaeniorhynchus complex by the absence of speckling on

the wing and by the presence of broad basal and narrow apical pale bands on

abdominal tergites II-VIl. Head. Narrow decumbent scales of vertex narrow,

linear, pale beige or yellowish; erect scales in center largely pale golden,

dark brown to black on posterolateral areas; lateral patch of broad scales pale

beige. Palpus dark scaled except for some pale scales at tip of segment 4.

Proboscis with a broad median yellow ring; no scattered pale scales proximad

of median ring on dorsal surface; apex of labium with light scales, forming an

indistinct apical ring. Thorax. Anterior 0.7 of mesonotum predominantly

pale yellow, beige to almost whitish; middle of fossae with or without a pair of

dark scale spots; scales on posterior 0.3 of mesonotum largely dark brown ex-

cept on midscutellar lobe which are sometimes partially pale on caudal margin.

Legs. Anterior surface of fore- and midfemora usually entirely dark, some-

times with a few scattered pale scales in apical 0.5; anterior surface of hind-

femur entirely dark; anterior surface of tibiae and tarsomeres 1 of all legs

entirely dark or not speckled. Wing. Scales on all wing veins entirely dark;

speckling of pale scales absent; scales on veins Ro Rg and R narrow, cla-

vate. Abdomen. Median scale patch of tergum I Sribisbies dan eras II- VII

usually with complete basal and apical pale bands; basal bands even in width or

broadened laterally, sometimes incomplete or represented by a median basal

spot on terga II-III; apical bands narrow or sometimes indistinct dorsally,

strongly widened laterally; tergum VIII with basal pale band only; dark scaled

areas on terga II-VI lightly sprinkled with pale scales; sterna II-VII usually

with broad basal pale bands and apical dark bands.

MALE (Fig. 26). In general similar to female in coloration of head,

thorax, legs and wing, differing in having broader basal abdominal pale bands

and in having apical bands represented largely by apicolateral yellowish

patches, particularly on abdominal terga II-V; palpus, proboscis and antenna

essentially as described for typical bitaeniorhynchus.

MALE GENITALIA (Fig. 27). Very similar to other members of the b7-

taeniorhynchus complex, differing particularly in the following features. Sub-

apical lobe. All setae in group d-/ of distal division slender, hairlike or flat-

tened, rodlike but none differentiated as lanceolate blade. Phallosome. Apical

spiculate portion of inner division more or less resembling luzonensis and

selangorensis, differing from both in having apical dorsal margin distinctly

depressed proximally and in the absence of small denticles or teeth adjacent

80 Contrib. Amer. Ent. Inst., vol.°12, no.:2,° 1976

to apical tergal angle; tergal lobe slender, elongate, with 2-3 small subequal

foliform processes on its apex, situated far below apical tergal angle; outer

division represented by a large, broad, leaflike spine. Pyroctiger. Basal

sternal process always developed, slender and pale, varying from 0. 04-0. 05

mm in length.

- PUPA. Abdomen: 3.6mm. Paddle: 0.78 mm. Trumpet: 0.70 mm.

Very similar to other forms of the bitaeniorhynchus complex, differing con-

spicuously in having integument of cephalothorax and abdomen entirely dark

brown and in the following combination of characters. Cephalothorvax, Trum-

pet darker than underlying integument; all setae relatively weak, short and

indistinct; seta 8-C single or double; 9-C usually triple (2-3). Metanotum.

Seta 11-C usually double (2-3). Abdomen. Setae 3-I-III usually double, rarely

single; 1-V 6-8 branched; 1-VI-VII double or triple; 5-IV 4-8 branched; 5-V-VI

3,4 branched. Paddle. Pale, contrasting sharply with abdomen or sometimes

uniformly lightly darkened on outer and inner parts, midrib moderately strong

and more pigmented.

LARVA (Fig. 27). Head: 0.65 mm. Siphon: 1.8 mm; index 7-9. Saddle:

0.36 mm; siphon/saddle ratio 4-5. As figured; extremely similar to typical

bitaeniorhynchus and other forms in that complex. Head, Seta 4-C relatively

long, about 2 times as long as distance between bases of the pair; 5-C usually

triple (3-4); 6-C always double; 7-C usually 7 branched (5-8); 11-C double or

triple; 14-C single, spiniform, or sometimes bifid. Thovax, Setae 1-4-M and

1-5-T extremely weak and rather indistinct, as in bitaeniorhynchus, Abdomen.

Dorsal setae 1-4 and ventral setae 10-13 of segments I-VI very weak and indis-

tinct, as in bitaeniorhynchus; 6-I-II usually triple (3-4); 6-III 3, 4 branched;

6-IV usually 4 branched (4-5); 6-V 3,4 branched; 6-VI triple; 1-III-VI

1-3 branched. Comb scales relatively small and short, usually 8 (6-11), in

2 rows; individual scale with a strong apical spine and basal fringe of 5-6 fine

spicules, basal sclerotized plate of scale oval and not as broad as in typical

bitaeniorhynchus. Seta 2-X of saddle 4, 5 branched; anal gills 1-1.5 times as

long as saddle. Siphon. Pecten always present in a distinct row in basal 0. 1-

0.2, number of teeth 8-10; subventral tufts 3 pairs (total 6) in number.

TYPE-DATA. Holotype 2* with associated pupal and larval skins, Adelphi

Estate, SINGAPORE, 5 March 1953, D. H. Colless (BM). |

DISTRIBUTION. Known only from Singapore, Peninsular Malaysia, Malay-

sia and southern Thailand. Material examined. 45 specimens: 20, 208, 5 L;

22 individual rearings (7 pupal, 15 larval).

THAILAND. Narathiwat: Waeng; 2%, 12, 4p.

MALAYSIA. Peninsular Malaysia: Johore-Labis; Kangkar Dohol; 7<, 79,

2L, 7 lp; Trengganu: Dungun; 4%, 39, 1 L, 3 p; Selangor: Petaling; 20, 69,

3 lp; Kedah: Bt. Kayu, Hitam; 10, 11p; Pevak: Tapah; 1“; Kelantan: Limau

Kasturi; 2 L.

SINGAPORE. Adelphi Estate (type-locality); 30", 32, 4 lp (type-series).

TAXONOMIC DISCUSSION. Culex pseudosinensis exhibits constant diag-

nostic features in all stages except for the variation in the presence or absence

of light speckling on the anterior surface of fore- and midfemora of the legs

and in the pattern of abdominal banding of the adults. In Peninsular Malaysia,

pseudosinensis adults can be easily confused with the pale form of znfula which

exhibits a similar pattern of abdominal banding. It can, however, be separated

from the latter by the complete absence of speckling on the wing and by the con-

siderably less speckling of the femora of legs. The apical portion of inner

division of the male phallosome of pseudosinensis is of a quadrate type as in

selagorensis and luzonensis but differs from the latter 2 species slightly in the

Sirivanakarn: Subgenus Culex in the Oriental Region 81

shape. The pupa of pseudosinensis is most distinctive, particularly in the uni-

form dark brownish coloration of the cephalothorax and abdomen in all material

examined. The pseudosinensis larva can be readily separated from most

bitaeniorhynchus forms but is virtually indistinguishable from that of selangor-

ensis. This evidence strongly suggests that pseudosinensis and selangorensis

are very closely related and appear to form a distinct lineage separated from

other members of the bitaeniorhynchus complex.

Culex pseudosinensis is currently known only from Singapore, Peninsular

Malaysia and Thailand. The record of this species from South Vietnam by

Bram (1967a: 265-8) is doubtful and requires confirmation as this was based on

a male (genitalia missing) which probably belongs to infula (tenax form).

BIONOMICS. The most frequent breeding sites of pPseudosinensis are

stagnant pools in or along stream margins under partial or heavy shade of

secondary tropical forest at the elevation ranging from sea level to 60 m. As

in other members of the bDitaeniorhynchus complex, these breeding sites al-

ways contain green algae. On one occasion, in Malaya, it was collected from

a marshy depression fully exposed to sunlight near a rubber plantation. All

adults came from rearing the pupae or larvae. Nothing is known about the

biology of the adult and its medical importance.

13, CULEX (CULEX) LONGICORNIS NEW SPECIES

(Fig. 24)

FEMALE. Wing: 4.14mm. Forefemur: 2.0mm. Proboscis: 1.9 mm.

Abdomen: 2.9mm. In general similar to other members of the bitaeniorhyn-

chus complex, differing in the following combination of characters. Head.

Narrow decumbent scales on vertex finer and largely pale beige or yellowish-

white; erect scales finer and predominantly pale in center, dark on postero-

lateral areas; lateral patch of broad scales whitish, very distinct. Thorax.

Mesonotal scales finer and predominantly yellowish-white on anterior 0. 7,

dark brown on posterior 0.3. Legs. Anterior surface of femora and tibiae

heavily and extensively speckled; tarsomere 1 of all legs lightly speckled.

Wing. Heavily and extensively speckled. Abdomen. Median caudal scale

patch of tergum I entirely dark; terga II- VII with complete broad basal and

narrow apical pale bands; apical bands on terga II-III or II-IV sometimes

very narrow to almost indistinct, all apparently connected with large apico-

lateral yellowish patches at sides; apical bands on terga V-VII as broad as

basal; sterna largely pale scaled.

MALE. Generally similar to female except for broader basal and apical

bands on abdominal terga II- VII and less speckling of legs and wing; palpus and

proboscis essentially similar to other forms of biltaeniorhynchus complex.

MALE GENITALIA (Fig. 24). Essentially similar to other forms of bitaeni-

orhynchus complex in basimere, distimere and proctiger, differing particularly

in the following features. Phallosome. Apical spiculate portion of inner

division with a very dark, slender and long hornlike apical sternal process,

its length measured from posterior margin of inner division to apex about

0.12 mm; apical tergal angle very irregular, with several dark denticles; ter-

gal lobe with 3-4 stout foliform processes situated very close to apical tergal

angle; outer division represented by a short and small laterally divergent spine.

' Proctigery. Apical crown relatively smaller; basal sternal process of para-

proct short, slender and pale, about 0.02 mm in length.

PUPA and LARVA. Unknown.

82 Contrib. Amer.) Ent. Insts, ‘vole 12,.no.22;° 1976

TYPE-DATA. Holotype “ (M85-13-193) with slide of genitalia, Doi Sutep,

Chiang Mai, THAILAND, elevation 925 m, 21 Nov. 1951, D. C. and E. B.

Thurman (USNM); Allotype ? (M85), same data as holotype (USNM).

DISTRIBUTION. Known only from Chiang Mai, Thailand. Material exa-

mined. 8 specimens: 5c, 39.

THAILAND. Chiang Mai: Doi Sutep, near wat, at top of mountain; 1c, 19

(M85; holotype and allotype), 20° (M85; M193; genitalia lost); 12 (M193); Huey

Keo; Wat Soda Ram; 10, 12 (M88); Chiang Mai, 1% (MLT5, genitalia lost).

TAXONOMIC DISCUSSION. The types and other identified specimens of

longicornis were discovered among numerous specimens of typical bztaenior-

hynchus in the collections by D. C. and E. B. Thurman from Chiang Mai,

Thailand during 1951-52. The associations of sexes and other specimens of

this species are only presumptive, but appear to be correct on the basis of

the adult external characters and the collection data. The immature stages of

longicornis are not known and the assignment of this species to the bitaenior-

hynchus complex is only provisional.

Culex longicornis can be readily differentiated from all other forms of the

bitaeniorhynchus complex by the type of the phallosome of the male genitalia

as described and illustrated. The adults of longicornis are generally very

similar to the usual pale form of typical bitaeniorhynchus in having the anteri-

or 0.7 of the mesonotum predominantly pale and in the extensive speckling of

the legs and wing but differ rather strikingly from the latter in the presence of

basal and apical pale bands on abdominal terga II-VII, resembling that of luzon-

ensis, pseudosinensis and infula.

BIONOMICS. The breeding sites of avec onuts are not known, but pre-

sumably are ground pools which contain algal mats as in other members of the

Bitaeniorhynchus Subgroup. All field caught adults from the Thurmans'’ col-

lections were captured near ponds and marshes containing algae in mountain-

ous areas at an altitude of 925 m. Nothing is known about the adult biology or

medical importance of longicornis.

14. CULEX (CULEX) SINENSIS THEOBALD

(ies, 15, 283 29.30)

Culex gelidus var. sinensis Theobald 1903: 180 (9).

Culex sepositus Leicester 1908: 152 (2); Edwards 1913b: 231 (synonymy).

Taeniorhynchus tenax of Leicester 1908: 167 (2), Edwards 1913b: 231

(synonymy).

Leucomyia sinensis Theobald 1910a: 313 (taxonomy).

Culex sinensis Theobald, Edwards ries 231 (taxonomy); Edwards 1921: 337

(e529): Barraud 1924a: 986 (o*, 9). 7

Culex tripunctatus Mochizuki 1913: 24 (i, 2%. EB); BaWwards' 1932::203

(synonymy).

Culex (Gales) sinensis Theobald, Edwards 1932: 203 (taxonomy); Barraud 1934:

394 (“*, 9, L); Stackelberg 1937: 231 (%*); Bonne-Wepster and Brug

1937: 78 (“, 2*); Baisas 1938: 209 (2, E); Bonne-Wepster and Brug 1939:

1274 (L*); Bohart and Ingram 1946: 79 (*, °, L*); LaCasse and

Yamaeguti 1950: 206 (%*, 9*, P*, L*); Monchadskii 1951: 275 (L*); Bonne-

Wepster 1954: 116 (¢,. 2*, L*); Hara 1957: 55 (2 genitalia); Delfinado

1966: 151 (9); Bram 1967a: 268 (o*, 9*, L*).

FEMALE (Fig. 15,28), Wing: 3.8 mm. forefemur:: 1.9 mm. Probos-

Sirivanakarn: Subgenus Culex in the Oriental Region 83

cis: 1.8mm. Abdomen: 3.5mm. Medium sized and relatively slender spe-

cies; distinguished from other members of the Bitaeniorhynchus Subgroup by

the presence of pale scale patches on prosternum; femora of all legs moder-

ately to heavily speckled; completely dark scales on all wing veins and by the

presence of narrow and even apical and basal pale bands on abdominal terga

II-VII. Head. Narrow decumbent scales of vertex fine and entirely whitish;

erect scales largely pale yellowish or golden except for dark ones on postero-

lateral areas; lateral patch of broad scales whitish, very distinct. Palpus en-

tirely dark except for scattered white scales on apex of segment 4. Proboscis

without scattered pale scales distad or proximad of median pale ring on dorsal

surface; apex of labium without any pale scales. Cibarial Armature. Cibarial

bar with a concave row of about 20 short teeth; 4-5 median teeth very fine,

lateral teeth stronger, apically pointed, blunt or truncate. Thorax. Proster-

num with distinct patches of pale scales on upper surface. Anterior 0.7 of

mesonotum usually predominantly pale yellowish to whitish, with or without

some dark scales forming spots and streaks in middle of fossae, anterior

acrostichal and dorsocentral areas; scales on posterior 0.3 of mesonotum and

prescutellar space entirely dark; scales on midscutellar lobe dark anteriorly,

pale on caudal margin; lateral scutellar lobes entirely pale scaled. Pleuron

with distinct scale patches on ppl, upper corner and posterior border of stp

and anterior upper mep; a few scattered pale scales among upper mep bristles;

ppl bristles 7-8. Legs, Anterior surface of femora of all legs moderately to

heavily speckled with pale scales; anterior surface of all tibiae lightly speckled;

tarsomeres 1 completely dark or not speckled; tarsomeres 1-4 with broad basal

and narrow apical pale bands. Wing. Scales on all wing veins completely dark;

scales on veins Rg, Rg and Ry,5 narrow, clavate or linear. Abdomen. Tergum

I with dark median caudal scale patch, terga II-VII usually with narrow, more

or less even transverse apical and basal pale bands; apical bands usually broad-

er and more distinct than basal, connected with slightly broader apicolateral

pale patches at side; basal bands usually composed of 1 or 2 rows of pale

scales which are sometimes not visible from above due to retraction of adjoin-

ing segments; basolateral pale spots present, extending to 0. 50-0. 75 of segment

length; tergum VIII with basal pale band only; dark scaled areas not speckled;

sterna III-VII with very broad basal pale bands and apicolateral dark areas.

Genitalia. Tergum IX with lateral row of 6-8 strong setae; postgenital plate

rounded on caudal margin; vaginal sclerite V or U-shaped, strongly sclerotized;

insula with a tuft of 10-12 setae.

MALE (Fig. 28). Essentially as described for female; sexual dimorphism

in general color not developed; ppl bristles about 20. Head, Palpus slender,

exceeding proboscis by about 1.5 of the length of segment 5; segment 3 with

median pale band on dorsal surface, apical 0.3 with ventral lateral row of

about 15 bristles, upper mesal surface without any bristles; segments 4 and 9

strongly plumose, each with distinct basal pale band on dorsal surface, scales

and bristles in apical 0.5 of segment 5 pale yellowish. Proboscis with promin-

ent ventral tuft of long, beh hairs at base of median light ring. Antennal flagel-

lum strongly plumose.

MALE GENITALIA (Fig. 29). Segment IX. Tergal lobe with a row of 5-6

setae. Basimere. Slender, conical, about 0.3 mm in length. Subapical lobe.

Setae a-b of proximal division subequal, strong, stout, rodlike and apically

hooked, seta c very thin and slender; setae d-f of distal division composed of 1

tiny hairlike seta and 3-4 stronger, rodlike setae, the distal most of which is

strongest and longest; leaf g narrow, lanceolate; seta # strong and apparently

flattened. Distimere, Normal, sickle-shaped, similar to other members of

84 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

the Bitaeniorhynchus Subgroup. Phallosome. Very distinctive; lateral plate

simple, represented by a straight, distally tapered and minutely spiculose por-

tion of inner division which is strongly sclerotized and dark brown in color;

outer division not developed. Pyvroctiger. Apical crown dark, medium to large

in size, consisting of several flat and blunt spicules laterally and numerous

spinelike spicules mesally; paraproct narrow; basal sternal process very thick,

dark and long, about 0.08 mm or more, apically rounded and strongly curved

sternad; cercal sclerite very thin; cercal setae 1-3 in number.

PUPA (Fig. 29). Abdomen: 3.0mm. Paddle: 0.78mm. Trumpet: 0.71

mm. Cephalothorax and abdomen whitish to cream-colored. Trumpet. Dark-

er than underlying integument; similar to other members of the Bitaeniorhyn-

chus Subgroup in asymmetrically funnel-shape, differing slightly in being more

slender and distally moderately widened; length of pinna about 0.3 mm. Com-

plete chaetotaxy as figured, the following setae are characteristic: Cephalo-

thorax. Setae 8, 9-C usually 3,4 branched. Metanotum. Seta 10-C 4-9 bran-

ched; 11-C usually double (2-3). Abdomen. Setae 3-I-III double; 1-II single or

forked into 2-4 branches; 1-III 8-10 branched, 1-IV 3-6 branched; 1-V 2-4

branched; 1-VI, VII double; 5-IV triple; 5-V, VI double, as long as or shorter

than segment following; 6-III-VI all single; 7-VI, VII double or triple; 9-VII

usually 5 branched (4-5); 9-VIII 7-9 branched; 4-VIII 3-5 branched; 10-III- VII

double or triple. Paddle, Whitish to almost transparent except for midrib

which is lightly pigmented.

LARVA (Fig. 30). Head: 0.65 mm. Siphon: 1.6-1.8 mm; index 8.

Saddle: 0.36 mm; siphon/saddle ratio 4.6. Generally conforming to the

Bitaeniorhynchus Subgroup characters, distinctive in the following. Head.

Labral sclerite not apparent or represented by an extremely narrow bar not

clearly separated from anterior margin of frontoclypeus; frontal margin of

head capsule broadly convex or slightly emarginate; seta 1-C dark, flattened,

moderately long and tapered into a sharp point, its length about 0.5 of distance

between bases of the pair; 4-C long, single, 2,3 times as long as distance be-

tween bases of the pair; 5-C double or triple; 6-C double; 7-C usually 9 bran-

ched (4-5). Antenna short, about 0.5 of head length; shaft entirely pale except

for narrow basal dark ring; spicules minute and relatively few, more or less

restricted to basal 0.5 on mesal surface; seta 1-A strongly plumose; 2, 3-A

far removed from apex or at mid point between base of seta 1-A and apex.

Mental plate black, with 10-12 distinct lateral teeth on each side of a median

large tooth, upper lateral teeth strong, gradually reduced in size towards

lower lateral angle of plate. Thorax. Seta 4-P short, minute and usually

double (2-3); 8-P single. Abdomen. Majority of dorsal and ventral setae very

weak, indistinct; 6-I-III triple; 6-IV-VI double; I-III-VI.3,4 branched. Comb

scales large, 4-6, in single irregular row, all subequal, about 0.08-0.10 mm

in length; individual scale with strong, prominent apical spine and basal fringe.

of minute spicules. Saddle same color as siphon; seta 1-X usually 4 branched

(3-4); 2-X 2-4 branched; ventral brush (4-X) with 5, 6 pairs of setae; anal gills

very slender, long, tracheate, 4-5 times as long as dorsal saddle length.

Siphon. Slender, long and cream-colored; pecten with 1-3 teeth at extreme

base; siphonal tufts 5 or 6 pairs (total 10 or 12), all subventral and each usually

triple (2-4), their length about equal to siphonal width at point of attachment;

seta 2-S dark, remarkably long and strong.

TYPE-DATA. (1) Culex gelidus var. sinensis Theobald, Holotype ?*,

Shaohyling, (Chekiang), CHINA (BM); (2) Culex sepositus Leicester, Type

non-existent, Kuala Lumpur, Selangor, MALAYA (PENINSULAR MALAYSIA);

(3) Taeniorhynchus tenax Leicester, Type non-existent, Kuala Lumpur,

Sirivanakarn: Subgenus Culex in the Oriental Region . 85

Selangor, MALAYA (PENINSULAR MALAYSIA); (4) Culex tripunctatus Mochi-

zuki, Type unspecified, Fukuoka, Kyushu, JAPAN (LU).

DISTRIBUTION. Widespread throughout the Oriental region with the range

extending north and northeast into the Palearctic region (China, USSR, Japan

and Korea). Material examined. 248 specimens: 570, 1642, 27 L; 11 indivi-

dual rearings (7 pupal, 4 larval).

INDIA. Bengal: Purneah District, Katihar; Kierpur; Lalmanirhat; 11°",

182; Karwa: N. Canara; 1%; Assam: Chabua; Dooma Dooma; Rupsi; Jorhat;

Dibrugarh; 4o%y 199) 9ot

SRI LANKA. Peradeniya; Anu District; 10, 19.

CHINA. Fuchow; Fukien, Yun Chun, Amoy region; Yunnan, Tengyueh;

Hainan; 205.79;

HONG KONG. 60, 89.

THAILAND. Chiang Mai; Chiang Rai; Udon Thani; Ubon Ratchathani;

Ayuthya; Nonthaburi; Bangkok; Trat; Prachuab Khiri Khan; Nakhon Si Tham-

marat; Trang; Narathiwat; 130", 589, 11 L, 7p.

VIETNAM. Da Nang; Cu Chi; Pleiku; Dan Tieng; Vungtau; Phu loi; Qui

Nhon; Cam Ranh; Quang Tin; 72,9’ Li.

MALAYSIA. Peninsular Malaysia: Selangoy-Ulu Gombak; locality not

specified; Pevak; 30°, 6, 4 lp.

INDONESIA. Java: Pelabuhan Ratu, Mt. Salat, Djarkata; Sumatra:

Bengkulu; Avoeé Island; 80", 169.

PHILIPPINES. Luzon: Pampanga, Rizal, Wack Wack; Bulucan; Calacad,

locality not specified; 3c, 139.

JAPAN. Okayama; 22, Ryukyus: Okinawa-Nago; Chizuka; Hentona; 20", 99.

KOREA. Chinhae; 3c", 89.

Additional records from the literature. BURMA: Bhamo (Barraud 1934:

395); BANGLADESH (Aslamkhan 1971); INDONESIA: Flores (Bonne-Wepster

1954: 117); REPUBLIC OF CHINA: Taiwan (Lien 1962: 635); CHINA: Chekiang,

Hupeh, Kiangsi, Kwangtung (Feng 1938: 300); USSR: Maritime Province

(Stackelberg 1937: 231; Monchadskii 1951: 275).

TAXONOMIC DISCUSSION. Culex sinensis is apparently uncommon but is

one of the most widespread Oriental species. It can be readily recognized in

the adults by (1) the presence of a distinct scale patch on the prosternum; (2)

presence of pale scale speckling on the anterior surface of femora and tibiae

of all legs; (3) entirely dark scales on all wing veins; (4) abdominal terga II-

VII with narrow, even apical and basal pale bands and (5) relatively slender

and small size; the male genitalia by (1) the simple phallosome and (2) basal

sternal process of the proctiger very thick and long; the pupa by (1) the uni-

formly pale coloration of cephalothorax, abdomen and paddle; (2) the slender,

asymmetric funnel-shaped trumpet; (3) seta 11-C double; (4) setae 3-I-III

double; (5) seta 4-VIII, 3-5 branched and (6) setae 7-VI-VII double or triple

and in the larva by (1) setae 2, 3-A far removed from apex; (2) seta 4-P min-

ute; (3) siphon with 5 or 6 pairs of subventral tufts and (4) seta 2-S remarkably

strong and long. Of all diagnostic features of sinensis, the simple male phal-

losome is unique among the members of the Bitaeniorhynchus Subgroup.

The sinensis adults are variable; the color of scales on the anterior 0.7 of

the mesonotum varies from usually predominantly pale to largely dark with

indefinite pale spots, blotches or streaks, similar to the pattern of variation

noted in some members of the bitaeniorhynchus complex. In the abdominal

tergal banding, the apical and basal pale bands are always well developed on

terga IV to VII, but very poorly developed on terga II-III and in most speci-

mens, the apical bands are broader and more distinct than basal. Although

86 Contrib. Amer; Ent.-Inst., vol 12, no. 2,.1976

sinensis is widespread, I have not been able to determine from the limited

amount of available reared material whether there is differentiation into geo-

graphical races.

BIONOMICS. The larvae and pupae of szmensis have been collected in

fresh water ground pools such as puddles, ponds and ditches in rice fields and

in stream pools at a broad range of elevation. These habitats contain abun-

dant vegetation, including green algae. In Thailand, several females were

caught biting man indoors and outdoors (Scanlon and Esah 1965) and in light

traps. A number of females from Vietnam, Sumatra, Indonesia and from

Luzon in the Philippines were found to be fully engorged with blood, but no

information was available whether they were caught biting man or other ani-

mals. The biology of sinensis is poorly known in Japan (LaCasse and Yamaguti

1950: 206-11). Evidence from adult collections in Thailand seems to indicate

that it is probably a semi- or peridomestic species, but apparently not suffi-

ciently common or abundant to be considered as pest or potential disease car-

rier. Nothing is known about its medical importance.

15. CULEX (CULEX) CORNUTUS EDWARDS

(Fig. 31)

Culex cornutus Edwards 1922a: 283 (““*, °); Barraud 1923a: 937 (L*); Barraud

1924a: 988 (o*, 9).

Culex (Culex) cornutus Edwards, Edwards 1932: 202 (taxonomy); Barraud 1934:

S96: (¢%, 2, 1%):

FEMALE. Wing: 4.2 mm. Forefemur: 1.8mm. Proboscis: 2.0 mm.

Abdomen: 2.9mm. Based on one specimen in the type series and 5 other

topotypic specimens from India. In general as figured for sinensis to which it

is extremely similar, differing slightly in the following features. Head. Clype-

us with variable amount of semierect pale scales (absent in sinensis). Palpus

lightly sprinkled with pale scales on segments 3-4. Proboscis with relatively |

broader median pale ring and with a light sprinkling of pale scales proximad

of ring on dorsal surface. Cibarial Armature. As described for sinensis.

Thorax, Anterior 0.7 of mesonotum usually predominantly golden brown to

dark brown with mottling of some yellowish scales not forming definite spots

or streak, sometimes predominantly pale yellow but not whitish; scales on

posterior 0.3 of mesonotum same color as those on anterior 0.7; scales in

middle of prescutellar space and on scutellar lobes largely pale yellowish.

Scale patches on ppl, stp and especially on mep well developed and relatively

broad. Legs. Speckling of pale scales restricted to anterior surface of fore-

and midfemora; anterior surface of hindfemur largely pale with light sprinkling

of dark scales; all tibiae and tarsi not speckled. Abdomen, Terga IV-VII with

apical bands broader than basal; terga II-III with incomplete basal bands and

very narrow, complete apical bands. Genitalia. Essentially similar to szn-

ensis,

MALE. In general as described for female, differing from sinensis in

the following: Head. Palpus relatively longer, exceeding proboscis by more

than 1.5 of segment 5; segment 3 with larger ventral lateral tuft of 30 or more

strong bristles in apical 0.5, upper mesal surface with a row of 5-6 long, dark

and fine hairs in apical 0.25 (absent in sinensis); segments 4 and 5 more strong-

ly plumose.

MALE GENITALIA (Fig. 31). Very similar to sinensis in basimere, sub-

Sirivanakarn: Subgenus Culex in the Oriental Region 87

apical lobe and distimere, differing particularly in the phallosome and proc-

tiger as follows. Phallosome, Inner and outer divisions of lateral plate well

developed; inner division broad and rounded on upper margin, with distinct

apical tergal angle produced tergad, somewhat resembling shape of tongue; low-

er tergal surface with a slender tergal lobe bearing 4-5 long, stout and apically

pointed rodlike processes; outer division with a strong, heavy mesal spine and

a very broad, rounded lateral basal process. Proctiger. Apical crown rela-

tively smaller and largely consisting of narrow and fine spicules; basal sternal

process longer and thicker, about 0.12 mm in length; cercal setae 2-3 in num-

ber.

PUPA. Based on 2 topotypic specimens associated with above male and

female. Extremely similar to sinensis and as figured for that species; differ-

ing slightly in the following features. Trumpet. Longer, about 0.9 mm in

length; pinna length about 0.835 mm. Cephalothorax and Metanotum. Seta 8-C

double; 10-C double or triple. Abdomen. Seta 9-VII double or triple; 9-VIII

o branched; 4-VIII double or triple.

LARVA. Based on 2 skins associated with the above male and female from

the type locality. Extremely similar to s¢zmensis and as figured for that species;

differing from it in the following features. Head. Antenna longer; spicules in

basal 0.5 of shaft stronger and more distinct. Thorax, Seta 4-P single, more

distinct; 8-P double. Abdomen. Comb scales longer, 0.12-0.15 mm, 4-95 in

number; basal plates of scales without distinct fringes of spicules. Seta 4-X

(or ventral brush) with 6 pairs of setae. Siphon. Subventral tufts 4-5 pairs.

TYPE-DATA. Lectotype “* (814), Tavargatti jungle, Belgaum, Deccan,

Bombay, INDIA, August 1921, P. J. Barraud (BM, present selection).

DISTRIBUTION. Known only from India and Pakistan. Material examined.

14 specimens: 6, 72, 1 L; 4 individual larval rearings.

INDIA. Bombay: Deccan, Tavargatti jungle; 1, 19 (type-series), 10, 19;

Trombay; Karwa, N. Canara, S.-W.; 4c, -92,-11,°3'lp (2. J. Barraud;: A.

Cogill).

PAKISTAN. Lahore; 1L (J. Maldonado, Sept. 1957).

Additional records from the literature: INDIA: Belgaum, Hog Island,

Bombay; Ennore, Madras (Barraud 1934: 396). 7

TAXONOMIC DISCUSSION. The above descriptions of all stages of cornutus

are based on the types and other specimens collected by P. J. Barraud in India.

The Pakistan record is based on a whole larva from Lahore where additional

specimens, especially reared adults are critically needed for confirmation.

Culex cornutus is closely related to sinensis on the basis of similarity in

adult external characters, pupal and larval stages. The adults of covnutus can

be differentiated from sinensis by (1) presence of semierect scales on lateral

surface of clypeus; (2) largely pale scales on the anterior surface of hindfemur;

(3) predominantly darker scales on the anterior 0.7 of mesonotum and in the

male by (4) presence of 5-6 long hairs on the upper mesal surface of palpal

segment 3. The male genitalia of cornutus are quite different from sinensis

in the type of phallosome and in the relatively longer and thicker basal sternal

process of the proctiger. The pupa and larva of cornutus are only slightly

different from sinensis. It is possible that some confusion may be encountered

in separating the pupa and larva of the 2 species except through association

with the adults including the male from individual rearing.

As in sinensis, adults of cornutus exhibit variation in the extent of the

abdominal tergal banding and in the color of scales on the mesonotum. Barraud

(1923a: 937) noted that larvae from a jungle pool in Tavargatti (type-locality)

differed from those from marshy fields at Trombay, Bombay Harbor in usually

88 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

having 6 pairs of subventral tufts on the siphon. The males which were reared

from these 2 larval forms are, however, identical in the genitalia (Barraud loc.

cit.), suggesting that only one form is probably involved. Because of the lack

of larval specimens and associated males from Tavargatti, I was not able to

determine whether the 2 larval forms are in fact conspecific. The larval char-

acters, particularly the number of siphonal tufts noted above, have been based

only on the specimens from Trombay.

BIONOMICS. The larvae of cornutus were collected from pools in marshy

fields and in heavy jungle (Barraud 1923a, 1934), but no information was avail-

able whether these habitats contained green algae. The single larva from La-

hore was collected in a borrow pit. The types and other topotypic adults came

from field catches and from rearing the larvae in the field. Nothing is known

about the adult biology and disease relationships of corvnutus.

16. CULEX (CULEX) EPIDESMUS (THEOBALD)

(Migy 32)

Taeniorhynchus epidesmus Theobald 1910b: 22 (2); Edwards 1913b: 231 (treated

as Culex epidesmus (Theobald)).

Grabhamia ochracea of Theobald 1905b: 25 (2); Edwards 1913b: 231 (rejected as

junior homonym of G. ochvacea Theobald 1901; synonymy).

Taeniorhynchus luteoabdominalis Theobald 1910b: 23 (2); Edwards 1913b: 231

(synonymy).

Culex epidesmus (Theobald), Edwards 1913b: 231 (taxonomy); Barraud 1924a:

982 (o'*, 2); Senior-White 1927: 61 (distribution).

Culex (Culex) epidesmus (Theobald), Edwards, 1932: 20 (taxonomy); Barraud

1934: 389 (“*, 9).

FEMALE. Wing: 4.5mm. Forefemur: 2.2 mm. Proboscis: 2.3 mm.

Abdomen: 3.5mm. Relatively large species, distinguished from other mem-

bers of the Bitaeniorhynchus Subgroup by the predominantly yellowish or golden

scales on head, mesonotum, legs, wing and abdomen. Head, Narrow decum-

bent scales of vertex and occiput whitish or yellowish; erect scales largely yel-

lowish or golden in center, brownish posterolaterally; lateral patch of broad

scales whitish. Clypeus with or without lateral patch of semierect scales.

Palpus extensively speckled or predominantly pale. Proboscis with a very

broad median yellow ring and’ extensively pale scaled proximad of ring on dor-

sal and ventral surfaces; apical portion distad of ring dark scaled except for a

few pale scales on apex of labium. Cibarial Armature. Not studied. Thorax.

Prosternum with pale scale patches. Anterior 0. 70-0.75 of mesonotum largely

pale yellowish or whitish with indefinite mottling of dark scales forming streaks

and spots in middle; a pair of dark fossal spots present or absent; scales on pre-

scutellar space and scutellar lobes whitish or silvery; scales on supraalar and

postdorsocentral areas largely brownish. Apn and ppn golden to brown scaled.

Pleural scale patches on ppl, stp and mep very well developed. Legs. Scales

on anterior surface of forecoxa largely pale; femora, tibiae and tarsomere 1

of all legs largely pale yellowish, with light or moderate sprinkling of dark

scales; apical and basal pale bands on tarsomeres 1-4 very broad; tarsomeres

5 entirely dark. Wing. Scales narrow and very dense; scales on apical por-

tion of veins C, Ry, Ro, Rg and R4,5 largely pale yellow, forming a broad api-

cal pale area; rest of wing veins with equal mixture of dark and pale scales

or sometimes largely dark, with light to moderate sprinkling of pale scales.

Sirivanakarn: Subgenus Culex in the Oriental Region 89

Abdomen. Color of terga I-VIII variable, usually entirely bright yellow to

golden; tergum II sometimes with a pair of dark lateral spots or dark basal

bands; terga III-V with narrow subbasal dark bands or broad basal dark bands;

tergum VI sometimes with a pair of median dark spots; terga VII-VIII always

entirely pale; sterna entirely yellow, with or without dark lateral patches or

spots. Genitalia. Not studied.

MALE. In general as in female, differing in having scales on head, thorax,

legs and abdomen more extensively pale yellowish. Head. Palpus exceeding

proboscis by a little more than full length of segment 5; segment 2 with broad

apical pale band; segment 3 largely pale from basal 0.1 to apex; segments 4

and 5 very strongly plumose; bristles on lateral and mesal surfaces of seg-

ment 5 largely whitish to yellowish. Proboscis with prominent yellow ventral

tuft of several long hairs proximad of median pale ring; portions basad and

distad of ring of labium largely pale. :

MALE GENITALIA (Fig. 31). Exceedingly similar to cornutus, differing

slightly in the following details. Subapical lobe. Leaf gof distal division

broader and longer. Phallosome. Apical spiculate portion of inner division

with a slender tergal lobe bearing 2 narrow elongate teeth, outer division with

smaller mesal spine and variable number of tubercles distad of a large lateral

basal process. Proctiger: Cercal setae 4 in number.

PUPA and LARVA. Unknown.

TYPE-DATA. (1) Taeniorhynchus epidesmus Theobald, Holotype 2, Bhogaon,

Purnea District, Bihar, INDIA, October 1908, C. Paiva (IM; Barraud 1934:

389); (2) Grabhamia ochracea Theobald, Lectotype ¢*, INDIA, Christophers

(BM; present selection); (3) Taeniorhynchus luteoabdominalis Theobald, Holo-

type °, Kathihar, Purnea District, Bihar, INDIA (IM).

DISTRIBUTION. Known only from India. Material examined. 15 speci-

mens: 7%, 89.

INDIA. Locality not specified; 22 (original type-series of G. ochracea

Theobald); Bihar: Pusa; 2%, 2°; Bengal; Assam; 5c, 39.

BANGLADESH. Sylhet; 19.

Additional records from the literature: INDIA: Western Himalayas: Simla;

United Provinces; Punjab (Barraud 1924a: 982; 1934: 389); SRI LANKA (Senior-

White 1927: 61-76); PAKISTAN; BANGLADESH (Aslamkhan 1971).

TAXONOMIC DISCUSSION. Culex epidesmus exhibits a strong affinity with

cornutus on the basis of similarity in the male genitalic characters, but is

strikingly different from the latter in the predominantly yellow scales on the

head, male palpus and proboscis, mesonotum, legs, wing and abdomen. These

outstanding features are constant in the specimens I have examined. Varia-

tions exhibited by ebidesmus adults are the presence or absence of dark spots

or bands on some abdominal terga, but these do not appear to be correlated

with any particular modification in the male genitalia. It seems most likely

that only a single species is involved.

As noted by Barraud (1924a, 1934), epzdesmus is rare but widespread in

the northern parts of India. The record from Sri Lanka by Senior-White (1927)

is very doubtful as it has never been confirmed by subsequent studies.

BIONOMICS. The breeding habitat of epidesmus is not definitely known.

The larval collection by Senior-White (1927) at Marlbe Estate, N.W.P, Sri

Lanka, was made in a buffalo wallow in an uncultivated rice field but subse-

quent visits to the same locality failed to recover any specimens of this species.

All of the adults from India apparently came from general field catches. Noth-

ing is known about the adult biology and the medical importance of epidesmus.

90 Contrib... Amer. Ent. Inst., vol 12, no. 2; 1976

17. CULEX (CULEX) GEMINUS COLLESS

(Fie. 32)

Culex (Culex) geminus Colless 1955: 314 (¢*, ¢, L*)

FEMALE. Wing: 4.5mm. Forefemur: 2.0 mm. Proboscis: 2.2 mm.

Abdomen: 3.06 mm. As described and figured for pseudosinensis (Fig. 26)

to which it is very similar, differing slightly in the following characters. Head,

All erect scales of vertex dull brown except for extreme apices which are

slightly pale. Legs. Anterior surface of fore- and midfemora lightly sprinkled

with pale scales; rest of legs uniformly dark. Abdomen. Terga II-VII with

narrow apical pale bands which are more or less even in width or slightly wid-

ened laterally; basal pale bands on terga II-IV absent or indistinct and those

on terga V-VII represented by a few rows of pale scales; dark areas on all

terga without speckling of pale scales.

MALE. Essentially as described for female, differing in having distinct

basal pale bands on abdominal terga II-VII and distinct apical bands on terga

V-VII; terga II-IV with or without very narrow apical bands. Head. Palpus

exceeding proboscis by a little more than full length of segment 5; segment 3

usually with median pale band on dorsal surface, sometimes absent or witha

few scattered pale scales, apical 0.5-0.6 with ventral lateral tuft of about 25

bristles, upper mesal surface with a row of about 8 weak bristles in apical

0.5; segments 4 and 5 with basal pale bands on dorsal surface; apical 0.5 of

segment 5 pale to tip. Proboscis with a ventral tuft of about 10 long hairs

basad of median light ring.

MALE GENITALIA (Fig. 32). As described and figured by Colless (1955:

314-5); differing from pseudosinensis and other members of the Bitaeniorhyn-

chus Subgroup particularly in the following features. Segment IX. Lateral ter-

val lobe with a row of 5-6 relatively weak setae. Subapical lobe. Setae a-c of

proximal division subequally long; seta b strongest; setae a and c similar in

size; setae d-f of distal division flattened, rodlike or bladelike, 4 in number,

the most distal longest, the other 3 subequal in length, all apparently with

blunt apices; leaf g very broad and rounded apically. Phallosome, Very dis-

tinctive, resembling members of Vishnui Subgroup; apical portion of inner

division with a distinct crown of 3-4 dark, stout and apically pointed fingerlike

processes, the most mesal one longest, the rest gradually shorter, all curved

and with apices pointing beyond apical margin of sternal portion of inner divi-

sion; outer division with 1 strong toothlike mesal spine, 1 laterally divergent

Spine and 1 sternal spine; sternal spine sharply angled basally with apex point-

ing basad; lateral basal process of outer division large and rounded. Proctiger.

Apical crown medium sized, composed of several dark spinelike spicules only;

basal sternal process of paraproct dark, stout and long, about 0.1 mm in length;

cercal setae 2 in number.

PUPA. As figured for typical bitaeniorhynchus (Fig. 21); differing in the

following characters. Cephalothorax and abdomen varying from deep yellow

to dark brown; pattern of dark alveolar spots absent. Cephalothorax. Seta

9-C 3,4 branched. Metanotum. Seta 10-C double. Abdomen. Setae 3-I-III

double; 1-IV, V 4-7 branched; 1-VI 3, 4 branched; 5-IV 4 branched; 5-V 2,3

branched; 5-VI triple; 7-VI, VII double or triple; 4-VIII 3-5 branched. Paddle.

Inner and outer parts more or less uniformly slightly darkened; basal external

margin of outer part not infuscated; midrib moderately pigmented.

LARVA (Fig. 32). As figured for typical bitaeniorhynchus and pseudosin-

irivanakarn: Subgenus Culex in the Oriental Region ou

ensis, differing in the following characters. Head. Seta 1-C distally tapered

into a fine point; 5-C usually 4 branched (3-4). Mental plate small, with 5, 6

strong, apically blunt lateral teeth on each side of median tooth. Abdomen,

Comb scales small, 7-11, in 2 irregular rows; individual scale with promin-

ent apical spine and basal fringe of fine spicules; seta 2-VIII 4-7 branched;

4-VIII 4-6 branched. Seta 2-X of saddle triple. Siphon. Pecten with 7-9 dis-

tinct teeth; subventral tufts 3 pairs (total 6).

TYPE-DATA. Holotype “* with associated pupal and larval skins and

‘slide of genitalia, Adelphi Estate, SINGAPORE, 7 April 1953, D. H. Colless

(BM).

DISTRIBUTION. Known only from Singapore and Peninsular Malaysia.

Material examined. 7 specimens: 40°, 192, 2 L; 3 individual rearings (1 pupal,

2 larval).

MALAYSIA. Peninsular Malaysia: Selangor-Banting; 1°, 19, 2 L, 1 p,

I Ip.

SINGAPORE. Adelphi Estate; 10", 1 lp (holotype); locality not specified;

2° (D. H. Given, 1924).

Additional records from the literature: MALAYSIA: Selangor, Kg.

Sijangkang (Colless 1955: 314-5).

TAXONOMIC DISCUSSION. Culex geminus is strongly differentiated from

the rest of the Bitaeniorhynchus Subgroup in the type of phallosome and in the

mental plate of the larva. On this basis it apparently falls into a distinct com-

plex. The adults of geminus are extremely similar to those of Pseudosinensis

in the coloration of mesonotum, the absence of pale scale speckling on the wing

and in the pattern of abdominal banding. The slight differences in the speckling

of the legs and the extent of apical as well as basal abdominal bands between

the 2 species are fairly constant, but for positive identification, the male geni-

talia and the associated larval stages should also be examined.

BIONOMICS. The immature stages of geminus were collected in a jungle

pool in association with the specimens of pseudosinensis (Colless 1955) and

those from Selangor, Malaysia, were collected in a ditch in association with

Culex (Lophoceraomyia) alphus Colless. These breeding sites are at sea level

and contain numerous green algae. The feeding habit of the females and their

medical importance are unknown.

18. CULEX (CULEX) KINABALUENSIS NEW SPECIES

(Figs. 33, 34)

FEMALE. Wing: 4.5-5.2 mm. (average 5.0). Forefemur: 2.1 mm.

Proboscis: 2.3mm. Abdomen: 2.9mm. Large species, distinguished from

all other members of the Bilaenitorhynchus Subgroup by the absence of speckling

on femora of legs and wing; the presence of only basal pale bands on abdominal

terga II-VII and by the following characters. Head, Erect scales of vertex

yellowish along dorsal midline in center, dark on anterior and posterolateral

areas. Palpus entirely dark except for white scales at extreme apex of segment

4, Proboscis without scattered pale scales proximad or distad of median yellow

ring; apex of labium without any pale scales forming distinct ring. Thorax,

Scales on anterior 0.7 of mesonotum largely brownish in middle, pale yellow-

ish on marginal areas, including: anterior promontory, humerus, scutal

angle and posterior border of fossae, continued as a distinct pale patch mesad

of supraalar bristles; scales on prescutellar space and scutellar lobes pale

beige to whitish. Apn with several narrow scales which are usually whitish

92 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

anteriorly, dark brown laterally; pbx with a broad narrow scale patch which is

largely dark. Pale scale patches on ppl, stp and mepofpleuron well developed.

Legs. Anterior surface of femora of fore- and midlegs usually dark, sometimes

with a few pale scales scattering in apical 0.25; apex of forefemur with distinct

subapical pale spot; hindfemur with a longitudinal pale stripe extending from base

to apex anteriorly, bordered by largely dark scales on lateral dorsal surface;

all tibiae and tarsomeres 1 not speckled; tarsomeres 1-4 with distinct basal

pale bands and very indistinct apical bands. Wing. Scales on all wing veins

dark; scales on veins Ro, Re and Ry,5 narrow, linear. Abdomen. Tergum I

with dark median scale ee oie iP VII with complete basal pale bands only;

apical bands absent; basal bands usually moderately broad and even in width,

sometimes largely retracted into preceding segments; sterna with broad basal

pale bands.

MALE. In general similar to female except for sexual characters. Head,

Palpus exceeding proboscis by about full length of segment 5; segment 3 with

median pale band or several pale scales not forming definite band on dorsal

surface, apical 0.25 with ventral lateral tuft of 10-15 bristles, upper mesal

surface without any hairs or bristles, ventral surface with a row of short,

pale hairlike setae along the whole length; segments 4 and 5 with narrow basal

pale rings; apical 0.25 of segment 5 pale to tip. Proboscis with a weak ventral

tuft of about 10-20 hairs proximad of median pale ring; no pale scales scattered

distad or proximad of median ring.

MALE GENITALIA (Fig. 33). Segment IX. Tergal lobe with 1,2 irregular

rows of 8,9 strong setae. Basimere. Normal, conical, about 0.34 mm in

length. Subapical lobe. Setae a-c of proximal division subequal in length;

setae a and } stout, seta c narrower and thinner, setae d-f of distal division

composed of 1 fine, hairlike seta and 2 apically hooked rodlike setae, the most

distal of which is strongest and longest; leaf g narrow, bladelike, with acumin-

ate apex. Distimere. Normal, sickle-shaped; subapical claw short or moder-

ate in length. Phallosome. Generally similar to geminus; inner division with

apical tergal crown of 2,3 stout and short fingerlike processes projecting

laterad; outer division very broad, with a strong mesal spine; lateral and ster-

nal spines not developed, outer lateral margin with variable number of small

tubercles or denticles; lateral basal process broadly rounded. Proctiger.

Apical crown medium to large; composed of several relatively long, flattened

spines laterally and numerous fine spinelike spicules mesally; basal sternal

process dark, thick, long, curved sternad; cercal setae usually 4, sometimes

3 OL.3

PUPA (Fig. 33). Abdomen: 4.0mm. Paddle: 1.1 mm. Trumpet: 0.78

mm. In general very similar to other members of the Bitaentorhynchus Sub-

eroup in relatively large sized and in having asymmetrically funnel-shaped

trumpet, differing in the following features. Pigmentation of cephalothorax

and abdomen cream-color or yellowish without definite darkened areas or

pattern of dark alveolar spots. Trumpet. Apical length of pinna relatively

short, about 0.35 mm. Cephalothorax. Seta 8-C long, 2-3 times as long as

9-C, 2-4 branched; 9-C usually double (2-3). Metanotum. Seta 10-C 5-8

branched; 11-C double; 12-C weak and short, usually double. Abdomen, Setae

3-I-III single or double: 1-III 2-5 branched; ms IV usually triple (3-4); 1-V-VII

single or double; 5-IV double, as long as segment following, sometimes shorter;

o-V, VI usually single (1-2), longer than segment following; 2-VII always me-

sad of 1-VII; 9-VII triple or 4 branched; 4-VIII usually double (2-3); 9-VIII 9,

10 branched. Paddle. Entirely whitish, without darkened or infuscated area;

seta 2-P present; 1-P absent.

Sirivanakarn: Subgenus Culex in the Oriental Region 93

LARVA (Fig. 34). Head: 0.70 mm. Siphon: 1.8-2.0 mm; index 6-7.

Saddle: 4.0 mm; siphon/saddle ratio 4-5. Essentially conforming to the

Bitaeniorhynchus Subgroup with the following distinctive features. Head, Seta

1-C dark brown, distally tapered to a fine point; 4-C triple, slightly shorter

than distance between bases of the pair; 5-C triple, 6-C usually double (2-3);

7-C 5-8 branched; 11-C weaker and shorter than 13-C, double; 13-C double;

14-C single, spiniform. Setae 2,3-A subapical. Mental plate with a large and

blunt median tooth and about 18 lateral teeth; 3 of the lateral teeth adjacent to

median tooth large, those towards lateral angle of plate gradually smaller.

Thorax. Setae 1,2-P double; 3-P triple; 4-P double, as long as 3-P; 7-P

usually triple (3-4); 8-P double. Abdomen, Setae 6-I, II 4-6 branched; 7-I

double; 6-III-V triple; 6-VI double or triple; dorsal setae 1-5 and ventral setae

9-13 of segment I-VI very weak and indistinct. Comb scales 12-15, in oval

patch; all scales subequal and moderately long, each with a strong apical spine

and basal fringe of fine spicules; setae 2 and 4-VIII double; 1-VIII 8-9 branched;

3-VIII 8-10 branched; 5-VIII 3,4 branched. Seta 2-X double or triple; 4-X with

6 pairs of setae; anal gills not tracheate, about 1.5 times as long as dorsal sad-

dle length. Siphon. Rather thick and moderately long, distally slightly tapered;

pecten teeth 3-4 in number, restricted to extreme base, sometimes completely

absent; siphonal tufts very weak, short, usually 5 pairs (total 10), all double

and subequal, shorter or as long as siphonal width at point of insertion; 2-S

dark, slender and moderate in length; median caudal filament poorly developed.

TYPE-DATA. Holotype & (No. $135.14) with associated pupal and larval

skins and genitalia slide, Mt. Kinabalu, Sabah, MALAYSIA, elevation 950-1,075

m, pool at edge of mountain stream, 20 March 1970, S. W. James et al.

(USNM); Allotype 2? (No. $135.12) with associated pupal and larval skins, same

data as holotype (USNM); Paratypes: 1% (No. S131.103) with associated pupal

skins; 22 (No. $131.100, S131.101) with associated pupal skins; 20° (No. S131);

22 (No. S131); 1% (No. S134); 22 (No. S134.100, S134.102) with associated pupal

skins, same data as holotype (USNM, BM).

DISTRIBUTION. Known only from the type-locality, Mt. Kinabalu, Sabah,

MALAYSIA. Material examined: 16 specimens: 5, 79, 4 L; 6 individual

rearings (4 pupal, 2 larval); as indicated in the type-data.

TAXONOMIC DISCUSSION. Culex kinabaluensis is distinctive in all stages

except for the pupa which is only slightly different from the rest of the Bitae-

niorhynchus Subgroup. On the basis of the male phallosome and larval charac-

ters, Rinabaluensis apparently falls into the geminus complex which, as inter-

preted here, also includes sguamosus (Taylor) from New Guinea, Australia and

the Solomon Islands and albinervis Edwards from Fiji as described by Belkin

(1962: 212-5).

The adults of both sexes of kinabaluensis exhibit much overlap or similarity

to most members of the Sitiens and Vishnui Subgroups in the color of scales on

the mesonotum and in the pattern of tergal abdominal bands. There is appar-

ently no adult dimorphism or presence of more than one color form in the

material examined. It can be separated from the members of the Sztiens and

Vishnui Subgroups in the female by its relatively larger size, presence of

distinct subapical spot on the apex of forefemur and in the male by having fine,

hairlike setae on the ventral surface of palpal segment 3 (scalelike or flattened

in Sitiens and Vishnui Subgroups). The male genitalia of kRinabaluensis differs

from geminus and other members of the Sitiens and Vishnui Subgroup in the

narrow, bladelike leaf g of distal division of subapical lobe; absence of the

lateral and sternal spines in the outer division and the shorter and fewer apical

fingerlike processes or teeth on the apical tergal margin of inner division of

94 Contrib. Amer. Bnt, Inst)).volc 123) no. 2)1976

the phallosome. The pupa and larva are basically similar to all other forms of

the Bitaeniorhynchus Subgroup except as indicated in the keys and as noted above.

The larva strongly resembles albinervis and starckeae as figured and described

by Belkin (1962: 212, 217) from the South Pacific in the form of the mental plate

and the comb scales, but is distinct from both in the branching of setae 1 and

2-P and setae 0 and 6-C.

BIONOMICS. The immatures of kinabaluensis all came from collections

made in ground pools at the edge of a mountain stream at an elevation of 950-

1,075 m on Mt. Kinabalu, Sabah. These breeding sites were partially shaded

in tropical rain forest and contained no aquatic vegetation. As noted by the col-

lectors, they were found in association with C. (Culex) mimulus and C. (Culi-

ciomyia)? bailyi Barraud. All adults were obtained from individual and mass

rearings from pupae and larvae. Nothing is known about adult feeding behavior

and its relation to disease.

SITIENS SUBGROUP

The Sitiens Subgroup is characterized by the following combination of

characters: in the adults by (1) medium size (wing length usually less than 4.0

mm); (2) erect scales of vertex usually entirely dark or sometimes whitish in

center, dark on posterolateral areas; (3) scales on anterior 0.7 of mesonotum

usually predominantly dark in middle, pale on marginal areas, sometimes en-

tirely whitish; (4) scales on posterior 0.3 of mesonotum laterad of prescutellar

space partially dark and pale or sometimes entirely whitish; (5) scales on pre-

scutellar space and scutellar lobes pale beige to whitish; (6) apn and ppn with

several narrow scales; (7) speckling of legs usually present, restricted to an-

terior surface of femora; sometimes absent; (8) scales on all wing veins en-

tirely dark and (9) abdominal terga with basal pale bands only; the male by (1)

ventral surface of palpal segment 3 usually with short, transparent scalelike

setae, or sometimes with dark fine hairlike setae; (2) apex of palpal segment

5 tipped with pale scales and (3) proboscis usually with ventral tuft of long

hairs at base of median pale ring; the male genitalia by (1) distimere normal;

(2) apical sternal portion of inner division of phallosome with expanded spicu-

lose or spinose lobe and a prominent apical tergal crown of 3-5 long fingerlike

or toothlike processes and (3) outer division of phallosome with differentiated

mesal and sternal spines or in form of a simple broad leaf or large, strong,

acute spine; the pupa by trumpet moderately long (0.52-0.91 mm), more or

less cylindrical or slightly widened towards apex and pinna lightly or strongly

oblique and in the larva by (1) labrum developed as a distinct transverse bar

separated from frontoclypeus; (2) head large, maximal width approximating

the width of prothorax; (3) mental plate with 4-10 strong lateral teeth on each

side of median tooth; (4) seta 4-P double and strong; (5) setae 2 and 4-VIII

single; (6) comb scales usually small, numerous, sometimes few and large;

(7) siphon moderately long; shape varied; siphonal tufts strong, 5-8 pairs; (8)

median caudal filament of spiracular apparatus poorly developed or absent and

(9) anal gills varied from very short, rounded or stubby to long, tubular or

fusiform. |

DISCUSSION. The Sitiens Subgroup is differentiated from the rest of the

Sitiens Group by the special modification of the complex male phallosome as

indicated in the above diagnosis. This subgroup corresponds to the Sztzens

Subgroup as defined by Belkin (1962: 202) and Bram (1967a: 236). In the Ori-

ental region the Sitiens Subgroup is represented by 4 species which apparently

Sirivanakarn: Subgenus Culex in the Oriental Region 95

fall into 3 more or less distinct complexes: sitiens, whitmorei and annuliros -

Lv IS

The sztzens complex includes sztiens and alis, both of which are coastal or

brackish water species and are closely related to litorvalis Bohart 1946 from

the Central Pacific; vosenz Belkin 1962 and whittingtoni Belkin 1962 from the

South Pacific and thalassius Theobald (Edwards 1941: 298) from the Ethiopian

region. This complex is chiefly characterized in the male genitalia by the

presence of mesal or sternal spines or both in the outer division of the male

phallosome; in the pupa by setae 6-III-VI single or double and setae 5-V-VI

usually double and in the larva by seta 1-C strongly flattened and striated; seta

7-I single; comb scales small, numerous, in broad oval patch; siphon straight

and thick; siphonal tufts strong, 6,7 pairs, 5-8 branched each, one out of line

or inserted laterally in apical 0.25; saddle complete or incomplete; anal gills

very short, stubby and rounded.

The whitmoret complex as proposed here includes only whitmorei and is

chiefly characterized in the male genitalia by outer division of the phallosome

represented by a single broad leaf; in the pupa by abdominal setae 6-III-VI 5-7

branched and setae 5-V-VI 6-8 branched and in the larva by seta 1-C slender,

spiniform; seta 7-I single; comb scales large, 5-8, in 1 or 2 rows; siphon

strongly curved upwards and slender; siphonal tufts very long, 7,8 pairs,

double; saddle complete; anal gills slender, tapering, about 1.5 times as long

as saddle.

The annulirostris complex includes only annulirostris as recognized by

Belkin (1962: 202). It is chiefly characterized in the male genitalia by outer

division of the phallosome represented by a strong, simple, laterally divergent

Spine; in the pupa by setae 6-III-VI 3-95 branched and setae 5-V-VI usually

double and in the larva by seta 1-C slender, spiniform; seta 7-I double; comb

scales small, numerous, in broad oval patch; siphon with 5,6 pairs of subven-

tral tufts, all in a line; saddle complete and anal gills slender, tapering, 1-2

times as long as saddle.

19. CULEX (CULEX) SITIENS WIEDEMANN

(Figs. 15, 35, 36, 37)

Culex sitiens Wiedemann 1828: 542 (2); Theobald 1901a: 360 (2*); Giles 1902:

400 (2); Blanchard 1905: 293 (2); Leicester 1908: 143 (“, 2); Edwards

1913b: 232 (taxonomy); Barraud 1924a: 993 (o*, 2); Barraud 1924c: 427

(L.*); Bowel 1930: 2304 *) 0.0 Fo),

Culex impellens Walker 1859: 91 (2); Theobald 1901la: 362 (2*); Giles 1902: 405

(o*, 2); Theobald 1903: 161 (o); Blanchard 1905: 294 (2); Leicester 1908:

142 (2); Theobald 1910a: 331 (distribution); Edwards 1913b: 232

(synonymy).

Culex microannulatus Theobald 1901a: 353 (o°*, 2*); Blanchard 1905: 292 (9);

Leicester 1908: 140 (“, 2); Theobald 1910a: 329 (distribution, taxonomy);

Edwards 1913b: 232 (synonymy).

Culex gnophodes Theobald 1903: 163 (2); Leicester 1908: 145 (2); Edwards

1913b: 232 (synonymy).

Culex nigricephala Leicester 1908: 149 (“, 2); Edwards 1932: 204 (synonymy).

Culex (Culex) sitiens Wiedemann, Edwards 1932: 204 (taxonomy); Barraud

1934: 398 (o*, 2, L*); Brug 1934: 514 (2); Bonne-Wepster and Brug 1937:

67 (, 2*); Bonne-Wepster and Brug 1939: 1268 (L*); Bohart 1945: 80

(2); Bohart and Ingram 1946: 18 (*, L*); Carter and Wijesundara 1948:

96 Contrib. Ameér, ‘nt. Inst. voleon2) noi'2; 1976

150 (2, L*); Bonne-Wepster 1954: 123 (o*, @*, L*); Belkin 1962: 205

(o*, 2, P*, L*); Delfinado 1966: 151 (o*, 2, L*); Bram 1967a: 239 (o'*,

oP Lit) Baisas 1974; 101 (Ce) Gs): |

Culex (Culex) salinus Baisas 1938: 204 (o*, 9*, P*, L); Delfinado 1966: 151

(synonymy).

For complete synonymy, see Stone, Knight and Starcke (1959).

FEMALE (Fig. 15, 35). Wing: 2.4-4.0 mm (average 3.5 mm). Fore-

femur: 1.4-2.0 mm (average 1.7 mm). Proboscis: 1.6-2.2 mm (average 1.8

mm). Abdomen: 2.3-3.2 mm (average 3.0 mm). Medium sized species.

Head, Narrow decumbent scales of vertex pale beige; erect scales usually

entirely dark brown, sometimes slightly pale anteriorly, dark brown postero-

laterally; lateral patch of broad appressed scales whitish. Palpus about 0.2 of

proboscis length; segment 4 tipped with some pale scales, rest entirely black.

Proboscis with clearly marked median pale ring which occupies about 0.2 of

total length; portions proximad and distad of median ring entirely black.

Cibarial Armature. Cibarial bar strongly sclerotized, with about 30 teeth ina

concave row which is slightly produced at middle; median 4-6 teeth minute,

terminating into a fine point; lateral teeth coarse and short, closely spaced,

with abruptly pointed or truncate apices. Thorax. Mesonotal integument

dark brown to black; mesonotal scales coarse and usually predominantly dark

in middle of disc, pale beige or golden on anterior promontory, humeral and

scutal angles, antealar and supraalar areas; posterior margin of fossa, acro-

stichal and dorsocentral areas cephalad of prescutellar space usually with pale

streaks or blotches of golden scales; prescutellar and scutellar scales entirely

pale beige or pale yellowish. Apn with several narrow dark or golden brown

scales; ppn with broad patch of narrow scales which are largely dark in middle,

pale towards anterior and posterior upper margins. Pleural integument slight-

ly paler than mesonotal except for darkened areas on psp, lower ppn and psp and

posterior border of stp; pale scale patches on ppl, sth and mep distinct; scales

among upper mep bristles absent; ppl bristles 7-9. Legs, Anterior scale

patch of forecoxa pale on upper lateral surface, largely dark or with a mixture

of dark and pale scales on anterior lower surface; scale patch on anterior sur-

face of midcoxa dark above, pale below; scale patch on anterior surface of

hindcoxa entirely pale; basal 0.5 of forefemur with distinct pale stripe on an-

terior lower margin, apical 0.5 lightly speckled with few pale scales, rest dark

scaled; anterior surface of midfemur usually with light speckling of pale scales

in middle; anterior surface of hindfemur largely pale, with light sprinkling of

scattered dark scales on upper lateral surface; foretibia largely dark, without

row of pale scaled spots among anterior bristles; mid- and hindtibiae largely

dark with or without light sprinkling of pale scales; tarsomeres 1-2 or 1-3

with very narrow apical and basal pale bands; tarsomeres 4-5 entirely dark.

Wing. Scales on all veins numerous and dark; base of vein C with or without a

short pale scaled streak on posterior surface; scales on veins Rg, Rg and Rais

narrow, clavate; cell Ro 1.25-1.50 times as long as R 43° Abdomen, Tergum

I with dark median caudal scale patch; terga JI-VIII with narrow, even basal

pale bands connected with elongate basolateral pale spots; terga VII-VIII also

with narrow apical pale band in middle; all sterna with broad basal pale bands

and apical dark bands. Genitalia. Tergum IX with lateral row of 6-7 setae;

postgenital plate broadly rounded on caudal margin; vaginal sclerite strongly

sclerotized and dark, U-shaped; insula with a dense tuft of 10-12 setae.

MALE (Fig. 35). As described for female, differing in having 15-16 pp/

bristles, broader basal pale bands on abdominal terga II- VIII and less dense

Sirivanakarn: Subgenus Culex in the Oriental Region 97

scales on wing veins. Head. Palpus exceeding proboscis by 1.5 of segment

2; segment 3 with distinct median pale band, ventral surface with a row of

short, distally flattened scalelike setae, apical 0.4-0.5 with ventrolateral tufts

of 20-30 dark bristles, upper mesal surface without any setae or bristles;

segments 4 and 5 strongly plumose, with distinct basal pale rings; apical 0.25

of segment 9 largely pale to tip. Proboscis with a prominent ventral tuft of

several long dark hairs proximad of median pale ring.

MALE GENITALIA (Fig. 36). Segment LX. Tergal lobe with a row of 5,6

setae. Basimere. Slender, conical, about 0.28 mm in length. Subapical lobe.

Setae a-b of proximal division equally long, seta a and 0 stout, seta c thinner;

setae d-f of distal division consisting of 3 short, narrow flattened setae and 1

long, thick rodlike seta; leaf g moderately to very broad, its apex blunt of

acuminate; seta h strong; sternal surface of lobe with a conspicuous patch of

numerous short and fine setae. Distimere. Normal, about 0.15 mm in

length; 1 dorsal and 1 ventral minute seta present, ventral distad of dorsal;

subapical claw very short, distally flattened and apically rounded. Phallosome.

Apical spiculate portion of inner division with a prominent apical tergal crown

of 4-5 strong, fingerlike processes; sternal apical portion more or less broad-

ly expanded, forming a distinct lobe sternad; outer division with a strong mesal

Spine and a very large sternal spine, the latter strongly curved with its apex

pointing basad; lateral spine absent; lateral basal process of outer division

large, elongate, apically rounded, knoblike. Pyroctiger. Apical crown dark,

medium to large in size, composed of 4, 5 flattened and apically blunt or trun-

cate spicules laterally and numerous strong spinelike spicules mesally; basal

sternal process dark, heavy and moderately long, about 0.08 mm; subbasal

process well developed, about 0.03 mm; cercal setae 2,3.

PUPA (Fig. 36). Abdomen: 3.2 mm. Paddle: 0.8 mm. Trumpet: 0. 62-

0.72 mm (average 0.65 mm); index 5-8. Cephalothorax and abdomen generally

cream-colored, with indefinite darkened areas along posterior middorsal

ridge, leg and wing cases, metanotum and abdominal segment I-IV. Trumpet.

Meatus narrow at base, gradually widened distally; pinna strongly oblique.

Cephalothorax. Seta 1-C strong and long, usually double; 2-C short 3, 4 bran-

ched; 3-C shorter than 1-C, double; 4-C double or triple; 5-C usually triple

(2-5); 8-C as strong as 1-C, usually double, rarely single; 9-C short, double

or triple. Metanotum, Seta 10-C 6-11 branched; 11-C double; 12-C usually

3,4 branched (3-5). Abdomen. Setae 6-III, IV single; 6-V, VI longer than 6-III,

IV, usually double (1-3); 1-II usually forked into 4-8 branches; 1-III- VI weak

and short; 1-III usually 6 branched (4-9); 1-IV usually 4 branched (3-5); 1-V

usually triple (2-4); 1-VI, VII usually double or triple (2-4); 5-IV double or

triple, slightly shorter than segment following; 5-V, VI usually double (2, 3),

1.0-1.5 times as long as segment following; 9-VII usually 4 branched (3-5);

4-VIII double; 9-VIII usually 11 branched (9-12). Paddle, Broad and entirely

pale except for lightly pigmented midrib; seta 2-P stronger than 1-P, usually

single, sometimes bifid; 1-P single.

LARVA (Fig. 37). Head: 0.78 mm. Siphon: 1.2 mm; index 4-5. Saddle:

0.32 mm; siphon/saddle ratio about 4. Head. Cream-colored with indefinite

darkened areas; seta 1-C strongly flattened, striated and dark, apex blunt or

abruptly pointed; 4-C single, about as long as distance between bases of the

pair; 5-C usually 5 branched (-8); 6-C usually 4 branched (2-5); 7-C 8-10

branched; 10-C usually double (2-4); 11-C usually triple (3, 4); 12-C 2-4 bran-

ched; 13-C double; 14-C single, distinct. Antenna relatively short, 0.9-0.7 of

length of head; spicules numerous and fine, mostly restricted to basal portion

below seta 1-A; setae 2, 3-A subapical. Mental plate black, with 6,7 strong

98 Contrib, “Amer. Bnt.-Inst:;~ vole.12/-no0.-2, 1976

lateral teeth on each side of median tooth. Thovax. Setae 1-3-P strong, sub-

equal and single; 4-P shorter than 1-3-P, double; 7-P usually triple (3,4); 8-P

usually double (1-3); setae 1 and 3-M single; 4-M double; 8-M 5-7 branched;

9-M 4-6 branched; setae 7 and 9-T 6-8 branched; 12-T single; 13-T usually

4 branched (3-6). Abdomen. Seta 6-I usually triple (3,4); 7-I always single;

6-II triple; 6-III-VI double; 3-I-VI single; 1-III-VI usually double or triple

(2-4); 4-V 5, 6 branched; 4-VI double; seta 1-VII usually 3, 4 branched (3-5);

4, 7, 10 and 12-VII all single; 13-VII 2-4 branched. Comb scales small, 34-40

in broad oval patch, all scales with even fringe of numerous fine spicules; seta

1-VIII 6-8 branched; 3-VIII 7-11 branched; 5-VIII 3,4 branched. Saddle com-

plete; ventral posterior margin deeply emarginated or indented; dorsal posteri-

or margin with small patch of distinct spicules; seta 1-X single and rather long;

2-X 3-6 branched; 4-X with 6 pairs of setae; anal gills very short, stubby and

rounded, about 0.9 of saddle length. Siphon. Relatively thick, short or moder-

ate in length; pigmentation pale yellow except for basal dark ring; pecten teeth

varying from 6-15, 3,4 distal teeth barbed with 7, 8 graded denticles; siphonal

tufts usually 7 pairs, sometimes 95 or 6.5 pairs; 5 proximal pairs strong, 9-8

branched each, in prominent row ventrally or subventrally; most proximal pair

longest, 1.5-2.0 times as long as siphonal width at point of attachment, rest

eradually shorter; 2 distal pairs weak and short, 3,4 branched, the more proxi-

mal pairs inserted laterally near midpoint of length, the most distal pairs sub-

ventral and close to apex; median caudal filament of spiracular apparatus poorly

developed or absent.

TYPE-DATA. (1) Culex sitiens Wiedemann, Holotype 2, Sumatra, INDO-

NESIA (ZMC); (2) Culex impellens Walker, Holotype 2* (in extremely poor

condition), Makessar, Celebes (Sulawesi), INDONESIA (BM); (3) Culex micro-

annulatus Theobald, lectotype o* with slide of genitalia, Quilon, Travancore,

(Madras State), INDIA, S. P. James (BM; selection of Bram 1967a: 243); (4)

Culex gnophodes Theobald, Holotype?*, Bruas, Dindings, MALAYA (PENINSU-

LAR MALAYSIA) (BM); (5) Culex nigricephala Leicester, Type unspecified,

Batu Gajah, Perak, MALAYA (PENINSULAR MALAYSIA) (BM; non-existent);

(6) Culex (C.) salinus Baisas, Holotype “ with associated pupal and larval

skins, Caloocan, Rizal, Luzon, PHILIPPINES, fish pond, 20 March 1937, F.

E. Baisas (PBH; destroyed). |

DISTRIBUTION. Widespread throughout coastal areas of Southeast Asia

and other adjacent tropical areas in the Oriental region with extensions north

and northeast as far as southern China and the Ryukyus. Also reported from

the Ethiopian region, the Middle East (Arabia and Iran), Micronesia, South

Pacific, New Guinea and northern Australia. Material examined. 2, 891 speci-

mens: 1,158, 1,219, 424 L; 392 individual rearings (135 pupal, 257 larval).

INDIA. Bengal: Calcutta; Dum Dum; Salt Lake; 1“, 42, 1p; Assam: Jor-

hat; 19. |

THAILAND. Chon Buri; Pathum Thani; Samut Sakhon; Samut Prakan;

Bangkok; Thon Buri; Trat; Prachuap Khiri Khan; Rangong; Nakhon Si Thammar-

at; Krabi; Phuket; Surat Thani; 128¢, 2059, 278 :L,:100p, 76 Ip.

VIETNAM Saigon; Nona Trang:-Cam Ranh Bay}. Vune Tau; 6¢, 6+, .29 L,

4p, 4 lp.

MALAYSIA. Peninsular Malaysia: - Selangor; Perak; Perlis; 370, 389,

4L, 5p, 23 lp. Malaysia: - Sabah; Sarawak; Brunei; 290, 52°, 4L, 2p, 23

lp.

SINGAPORE. 160 2 179, 15 Ly 8p,od6tip:

INDONESIA. Sumatra; Java; Flores; West Irian (formerly Dutch New

Guinea); 130°, 202.

Sirivanakarn: Subgenus Culex in the Oriental Region 99

PHILIPPINES. Luzon; Mindoro; Palawan; Leyte; Jolo Jolo; Tawi Tawi;

Mindanao; 908“, 8462, 81 L, 15 p, 95 lp.

TAIWAN. 60, 92) 13° Li.

HONG KONG. 49.

JAPAN. Ryukyus: Okinawa; 14%, 219, 21 lp.

Additional records from the literature: IRAN: Bandar-Abbas, (Lofti 1970:

401); INDIA: Bombay, Trombay and Islands; Deccan, Belgaum and Nagargali; +

Nilgiri Hills, Coonoor; CEYLON (SRI LANKA): Colombo, (Barraud 1924: 994);

CHINA: Fukien, Amoy; Kwangtung, Canton; Hainan (Feng 1937: 301; Chu 1958:

110); INDONESIA: Greater and Lesser Sundas; the Molluccas; West Irian

(Dutch New Guinea), (Bonne-Wepster 1954: 373; Assem and Bonne-Wepster

1964: 126); CENTRAL PACIFIC: S. Mariana Is., Guam; Yap; Caroline Atolls,

Negulu; Truk, Wena (Moen) (Bohart 1956: 82); SOUTH PACIFIC: Solomon Is. ;

New Hebrides; New Caledonia; Ellice Islands; Fiji; Wallis; Samoa; Tonga; Niue

Island (Belkin 1962: 205); and others.

TAXONOMIC DISCUSSION. Culex sitiens is one of the commonest members

of the Sitiens Group in Southeast Asia, where it has been found, as reported

elsewhere, to be restricted to seacoasts, river banks and margins of canals in

the estuarine environments. It is also one of the most widespread tropical

forms with a very extensive range of distribution covering the Ethiopian,

Palearctic, Oriental and Australasian regions and the islands of the Central

and South Pacific. The Southeast Asian populations of szfiens exhibit much

individual variation in all stages but there appears to be no differentiation into

recognizable subspecies, geographic forms or races. All Southeast Asian

material agrees very well in the diagnostic characters of the adults and male

genitalia with that from New Guinea (Bonne-Wepster 1954), the South Pacific

(Belkin 1962), Micronesia (Bohart 1956), India (Barraud 1934) and the Ethiopian

region (Edwards 1941). The associated pupae and larvae from Southeast Asia

also agree very well with the descriptions and figures of this species by Penn

(1949) from New Guinea and by Belkin (1962) from the South Pacific.

Culex sitiens can be readily distinguished from other members of its

Subgroup and complex in all stages over most parts of Southeast Asia and adja-

cent areas except in the Philippines where the adults are most liable to be con-

fused with those of annulirostris, As indicated in the keys by Bohart (1956

[ 1957]) and Belkin (1962), the sitiens females can be distinguished from

annulirostris by the absence of a row of pale dots among the bristles on the an-

terior surface of foretibia (present in annulirostris) and by relatively narrower

and even basal pale bands on the abdominal terga (usually broader and medially

produced in annulirostris). These diagnostic characters are further substanti-

ated by the largely pale anterior surface of the hindfemur in Szfzens against

the freckled or pepper and salt appearance in annulirostris. The sitiens males

can be differentiated from annulirostris by the translucent scalelike setae on

the ventral surface of palpal segment 3 (dark, hairlike in annulirostris) and by

more numerous long hairs at the base of the median pale ring on the ventral

surface of the proboscis.

BIONOMICS. Culex sitiens is typically a coastal, brackish water species

whose breeding habitats usually include pools, puddles, ponds, wells, ditches,

crabhole and rock pools in salt marsh, mangrove and nipa palm swamps. It

has also been found breeding in artificial containers such as canoes, boats,

cement tanks, jars, cans, etc. in the vicinity of sea beaches, harbors or piers

in populated areas. The larvae and pupae are very adaptable and show a high

tolerance of salinity, occurring in fresh, brackish and even pure sea water.

One of the most remarkable larval features, which is apparently associated

100 Contrib. “Amer, But, Inst. vole:12,.no. 2, 2976

with the adaptation to high salt content, is the considerable reduction in the

length and size of the anal gills of the saddle. The adults of sztzens have been

collected outdoors as well as indoors while resting. Several females from

Thailand were caught biting man and in light traps. In Singapore, Colless

(1959) reported that the females prefer to feed on birds, pigs and probably

cattle. In Malaya, Bennett, Warren and Cheong (1966), Bennett and Warren

(1966) incriminated sitiens as one of the natural vectors of a malarial parasite

of domestic fowl (Plasmodium juxtanucleare Versiani and Gomes). In Thai-

land, a wild caught female was found to be naturally infected with Brugia malayi

(Iyengar 1953), but this author concluded that sitiens was probably not an im-

portant vector of this parasite. Elsewhere, sztzens has been shown to be cap-

able of transmitting Japanese encephalitis virus in the laboratory (Hodes 1946)

but no further data are available in support of its role in the natural transmis-

sion of this pathogen.

20. CULEX (CULEX) ALIS THEOBALD

(Figs. 38,39)

Culex (Culex) alis Theobald, Edwards 1932: 203 (taxonomy); Brug 1934: 517

(*, 2*); Brug and Bonne-Wepster 1947: 189 (distribution).

Culex (Culex) litoralis Bohart, Colless 1957b: 465 (L*).

Culex (Culex) neolitoralis Bram 1967a: 236 (**, 92, L*). NEW SYNONYMY.

FEMALE. Wing: 2.3-3.2 mm (average 3.0 mm). Forefemur: 1.1-1.7

mm (average 1.5 mm). Proboscis: 1.3-1.8 mm (average 1.7 mm). Abdo-

men: 1.8-2.7 mm (average 2.5 mm). In general as figured and described for

sitiens to which it is very similar, differing in smaller size and in the follow-

ing characters. Head, Erect scales of vertex usually pale yellowish or bronzy

in center, black posterolaterally. Thorax. Mesonotal scales usually predom-

inantly yellowish or golden with dark scales forming streaks or spots on acro-

stichal line, middle of fossae and posterior dorsocentral areas. Legs. Anteri-

or surface of fore- and midfemora without speckling of pale scales; anterior

surface of hindfemur with longitudinal pale stripe extending from base to 0.7

of total length, no dark scales scattered on pale stripe area; tibiae and tarso-

mere 1 of all legs without speckling of pale scales on anterior surface; apical

and basal bands of tarsomeres 1-4 of all legs very narrow and rather incon-

spicuous. Wing. Cell Ro shorter, as long as or slightly longer than vein

R92.

: MALE. In general as in female, differing from sztiens particularly in the

following characters. Head. Palpus variable in length, usually rather thick

and relatively shorter; segments 4 and 5 reduced in length, weakly to moder-

ately plumose, all bristles shorter and weaker; segment 3 with weaker ventro-

lateral tuft of about 8 bristles in apical 0.25. Proboscis without ventrolateral

tuft of long hairs at base of median pale ring. Wing. Cell Ro considerably re-

duced in length, from 0.5-1.0 of the length of vein R9.3.

MALE GENITALIA (Fig. 38). Differing from sztiens particularly in the

following. Subapical lobe. Setae d-fof distal division weaker; leaf g narrower;

sternal surface mesad of proximal division without a distinct patch of dense

spicules. Distimere, Basal 0.4 with or without scattering of a variable num-

ber of minute spicules on outer lateral surface. Phallosome. Apical tergal

crown of inner division with 2-4 shorter fingerlike processes; sternal portion

Sirivanakarn: Subgenus Culex in the Oriental Region 101

of apical spiculate portion more strongly expanded sternad, forming a promin-

ent lobe, bearing several distinct denticles; outer division very broad, simple,

leaflike, mesal spine absent; sternal spine present, very large, broad and

gently curved basad. Proctiger. Basal sternal process absent or present as

a very short protuberance; cercal setae 3.

PUPA (Fig. 38). Abdomen: 2.1-3.5 mm (average 2.8 mm). Paddle: 0. 65-

0.78 mm (average 0.71 mm). Trumpet 0.46-0.71 mm (average 0.60 mm); in-

dex 5-6. Pigmentation variable, usually with conspicuous dark or black areas

on cephalothorax, metanotum and abdominal segments I-IV. Trumpet. Slender

and relatively short; apical 0.5 usually uniform in width, sometimes slightly

widened toward apex; pinna slightly to moderately oblique. Complete chaetotaxy

as figured, generally very similar to sitiens, differing from it as follows.

Cephalothorax., Seta 8-C weaker and shorter than 9-C, usually double, some-

times single; 9-C usually double (1-3). Metanotum. Seta 10-C usually 3,4

branched (2-6). Abdomen. Seta 1-II usually 3,4 branched (2-10); setae 1-III-

VII weaker and shorter; 1-III usually triple (2-4); 1-IV-VII usually double

(1-4). Paddle. Setae 1 and 2-P minute, subequal and single.

LARVA (Fig. 39). Head: 0.65 mm. Siphon: 0.8 mm; index 3-4. Saddle:

0.33 mm; siphon/saddle ratio 3.0-3.5. As figured, generally similar to sitiens,

differing from it in the following features. Head, Frontoclypeus partially

strongly infuscate or blackish; seta 1-C larger, more flattened, somewhat

foliform, with blunt or acuminate apex; 4-C situated closer to dorsal midline,

its length about 1.5 times as long as distance between bases of the pair; 5-C

usually 3,4 branched (3-6); 6-C 3,4 branched. Antennal shaft shorter, 0.4-0. 6

of head length; proximal portion pale, distal portion blackish; setae 2,3-A

rather short, apical or slightly subapical. Mental plate blackish, with 8-10

lateral teeth on each side of median tooth. Thovax. All prothoracic setae rela-

tively weaker and shorter; 7-P usually double, sometimes triple; 8-P variable,

from very short, minute, 2-4 branched to as long as 4-P and single (1-2);

setae 1-4-M and T shorter and weaker. Abdomen. Setae 1-III, IV usually double

(2,3); 1-V strong, always single; 1-VI, VII single or double. Saddle incomplete,

sclerotization restricted to dorsal and ventral portions, middle portion mem-

branous; anal gills very short and stubby, as in sztzens, Siphon. Shorter and

darker, concolorous with head capsule; 4,5 distal pecten teeth barbed with 3, 4

strong basal denticles and 1 strong apical denticle; siphon tufts 6,7 pairs; 5

proximal pairs very closely spaced, in prominent row on ventral surface; medi-

an caudal filament not developed.

TYPE-DATA. (1) Culex alis Theobald, Lectotype o* with slide of genitalia,

Christmas Island (Australian Trust Territory), reared from larva in salt pool,

13 December 1902, S. Durham (BM; selection of Bram 1967: 327); (2) Culex

(C.) neolitoralis Bram, Holotype “* (00652-2) with associated pupal and larval

skins and genitalia slide, Ban Laem Sing, Chanthaburi, THAILAND, larva from

brackish rock pool on a beach, 6 November 1965, Somboon Maneechai (USNM).

DISTRIBUTION. Presumably widespread throughout Southeast Asia and

other adjacent tropical areas of the Oriental region. Presently recorded from

Thailand, Vietnam, Malaysia, Singapore, Taiwan, Indonesia, Christmas Island

(Australian Trust Territory) and the Philippines. Material examined. 6795

specimens: 204, 2462, 225 L; 273 individual rearings (183 pupal, 90 larval).

THAILAND. Chanthaburi: Ban Laem Sing; Chonburi; Rayong; Surat Thani:

Koh Samui; Tvat; Koh Chang; 1410", 1339, 176 L, 183 p, 40 lp (as neolitoralis

Bram 1967).

VIETNAM. Con Son; 20, 649, 23 L.

MALAYSIA. Peninsular Malaysia: Pahang-Pulau Tioman; Tg. Gelang;

102 Contrib. Amer. Ent. Inst), vol. 12, no..2,. 1975

Trengganu: Kemasik; Kelantan-Tumpat; 230, 232, 16 L, 22 1p. Malaysia:

Sabah-Sipitang; 4°, 10%, 4L, 5 lp.

SINGAPORE. PP: Hantu; 20, 32, 5 L, 2)Ip.

TAIWAN. Orchid Island: 1%; 32, 3:lp:

INDONESIA. Ceram; West Irian (Dutch New Guinea); 50", 19.

PHILIPPINES. Luzon: Subic Naval Base; Samar: Osmena; 8, 99, 1 L,

18 Ip.

TAXONOMIC DISCUSSION. Culex alis has been a poorly known species

since it was described by Theobald (1903) from Christmas Island south of Java,

Indonesia. This is because Theobald's description was based only on the adult

male and female and the male genitalia were neither figured nor described.

The only significant diagnostic feature of alis given by Theobald (1903) is in

the length of Cell Ro (or anterior fork cell) of the wing, which is shorter than

vein Ro, (or its stem). Subsequently, Edwards (1922a) illustrated the phallo-

some and the proctiger of the male genitalia of a cotype male and provided the

key to the adults but did not provide a redescription nor statement about its

specific status to clarify its identity. The only other records of alis in the

literature were those of Brug (1934) and Brug and Bonne-Wepster (1947) from

adults collected in Dutch New Guinea (West Irian and Ceram, Indonesia) and by

Colless (1957b: 465) from the larvae (as litoralis Bohart 1946) in Singapore

harbor.

Culex alis as interpreted here is a widespread coastal species and as in

sitiens, is restricted to breeding in brackish water ground pools. The South-

east Asian species which was described by Bram (1967a: 236) as neolitoralis

is identical to the type adults of alis in the relative length of cell Ro of the wing

and in practically every detail of the male genitalia. This evidence as well as

the similarity in larval habitat strongly suggests that both forms are conspecific

and on this basis Iam synonymizing neolitoralis with alis. There is a possibili-

ty that whittingtoni Belkin (1962: 204) from the Solomon Islands may also be con-

specific with this species. However, as whittingtoni has been known only in the

larva, the point cannot be settled. In comparing the larvae of whittingtont with

those of alis, I have found no difference whatever in the chaetotaxy and other

features, including sclerotization of the saddle.

Among the members of the sitiens complex, alis is very similar to lztoralis

Bohart (1946) from the Central Pacific; voseni Belkin (1962: 203) and whitting-

toni Belkin (1962: 204) from the South Pacific in all known stages. Each of

these forms is probably only subspecifically distinct and it is possible that

they were originally represented by a single widespread species which was

subsequently differentiated into slightly distinct segregates through geographical

isolation.

Culex alis can be readily distinguished from sitiens (with which it largely

overlaps in distribution and breeding sites) by features indicated in the keys

and in the above diagnosis. It can be differentiated from litovalis and roseni

in the adult by shorter cell Ro of the wing and by the reduction in the length and

in the plumosity of the male palpal segments 4 and 5. The male genitalia is

virtually indistinguishable from /ztoralis, but is distinct from voseni in the

presence of setae d-f in the distal division of the subapical lobe (absent in

voseni). The pupa differs from litoralis in having abdominal setae 6-III- VI

single or double (3, 4 branches in litovalis). The larva strongly resembles

litovalis and voseni in all features except for the sclerotization of the dorsal

as well as ventral halves of the saddle (only dorsal half in Jitovalis and roseni).

BIONOMICS. Culex alis is typically a brackish water species as sztiens

but is apparently less common than the latter. The majority of pupal or larval

Sirivanakarn: Subgenus Culex in the Oriental Region 103

collections from Thailand and other adjacent countries came from pools among

rocks at or near sea beaches in the vicinity of harbors and fishing villages. A

few collections came from crabholes and coral holes. Most of the adults were

obtained from rearing pupae or larvae in the field and no records are available

of their behavior and feeding habits. Nothing is known about the medical impor-

tance of alis.

21. CULEX (CULEX) WHITMOREI (GILES)

(Figs. 15, 40, 41, 42)

Taeniorhynchus whitmorei Giles 1904: 367 (9).

Taeniorhynchus argenteus Ludlow 1905: 98 (2); Theobald 1910a: 426 (2*);

Edwards 1913b: 232 (synonymy).

Leucomyia plegepennis Theobald 1907: 375 (2); Edwards 1913b: 232 (synonymy).

Culex albus Leicester 1908: 148 (2); Edwards 1913b: 232 (synonymy).

Culex loricatus Leicester 1908: 151 (2); Edwards 1917: 225 (synonymy).

Culex whitmorei (Giles), Edwards 1913b: 232 (taxonomy); Edwards 1917: 225

(taxonomy); Barraud 1923a: 941 (L*); Barraud 1924a: 988 (o*, 9).

Culex (Culex) whitmorei (Giles), Edwards 1932: 203 (taxonomy); Barraud 1934:

406 (““*, 2, L*); Stackelberg 1937: 232 (o'*, 2); Bonne-Wepster and Brug

1937: 77 (, 2*); Baisas 1938: 211 (**, 9, P*, L*); Bonne-Wepster and

Brug 1939: 1273 (L*); Bohart 1945: 81 (2, L); La Casse and Yamaguti

1950: 211 (o*, 2*, P*, L*); Monchadskii 1951: 281 (L*); Bonne-Wepster

1954: 120 (“, 2*, L*); Hara 1957: 56 (2 genitalia*); Delfinado 1966: 155

(ox Oe Bram l96%a. 255 to, LF).

FEMALE (Fig. 15, 40). Wing: 3.0mm. Forefemur: 1.4 mm. Proboscis:

1.6mm. Abdomen: 2.4mm. Small species; wing length not exceeding 3.5

mm; differing from other members of the Sitzens Subgroup in the following

features. Head, Narrow decumbent scales of vertex pure white, contrasting

sharply with dark integument underneath; erect scales relatively short and

slender, largely pure white except for a few dark ones on posterolateral areas.

Palpus rather slender and short, about 0.2 of proboscis length; segment 4

tipped with white scales on its apex, rest dark. Proboscis with distinct medi-

an pale ring which occupies about 0.33 of total labial length. Czbarial Armature.

Cibarial bar with a double row of short and apically pointed teeth; number of

teeth in each row about 20. Thorax. Mesonotal integument blackish or chest-

nut brown; mesonotal scales rather sparse to moderately dense, predominantly

pure white except for dark scaled spots in middle of fossa, continuing as whitish

patch and streaks on prescutellar space and lateral prescutellar areas; posteri-

or dorsocentral and supraalar areas with dark scaled streaks; scutellar lobes

pure white scaled. Scales on apn pure white; scale patch of ppn dark in middle

and on lower surface, pale whitish on anterior and posterior upper surfaces.

Pleural integument very dark or same color as mesonotum; pleural scale

patches very distinct. Legs. Anterior surface of fore- and midfemora moder-

ately to heavily speckled with pale scales; anterior surface of hindfemur largely

pale or freckled with numerous dark scales; mid- and hindtibiae with poorly

defined narrow pale stripe on anterior surface; apical and basal pale bands on

tarsomeres 1-4 of all legs distinct. Wing. All scales dark; scales on veins

R45; eat M3,4, Cuy and Cu, broader than on other veins. Abdomen,

Median caudal scale patch of tergum I dark or partially pale; terga II- VII with

large median triangular basal pale patches and small apicolateral pale spots,

104 Contrib. (Amer. Bats Inst); vol. 12\no. 2; 1976

the latter usually distinct on terga V-VII; sterna largely pale scaled. Genitalia.

Tergum IX with lateral row of 3-6 setae; postgenital plate rounded on caudal

margin; vaginal sclerite broadly curved, resembling an arc of a circle; insula

with a tuft of 10-12 setae.

MALE (Fig. 40). Head. Palpus usually exceeding proboscis by full length

of segment 5, sometimes longer; segment 2 with apical pale band; segment 3

with median pale band, apical 0.25 with ventrolateral tuft of about 10 bristles,

ventral surface with a row of several short, translucent scalelike setae; seg-

ments 4 and 5 strongly long plumose, with basal pale rings; apical 0.2 of seg-

ment 5 pale to tip. Proboscis with very narrow median pale band which is about

0.1 of total length; ventral tuft of long hairs at base of median band absent.

MALE GENITALIA (Fig. 41). Distinctive in the following features. Sub-

apical lobe. Setae a-c of proximal division long, subequal in length and thick-

ness; setae d-f subequally long, rodlike and hooked apically, most distal strong-

est, the other 2 narrower; leaf g long, narrow, club-shaped; seta / strong,

long, bristlelike. Phallosome. Apical spiculate portion of inner division of

lateral plate strongly expanded sternad, forming a tonguelike lobe which is

finely spiculose; apical tergal surface with a crown of 3,4 equally long teeth,

all of which are straight, projecting dorsolaterad; outer division of lateral

plate represented by a simple and broad leaflike process; mesal, lateral and

sternal spines absent; lateral basal process small or poorly developed. Pvroc-

tiger. Apical crown large and dark; basal sternal process very slender,

moderate in length, about 0.04 mm; subbasal process not developed; cercal

setae 3-5.

PUPA (Fig. 41). Abdomen: 2.4mm. Paddle: 0.67mm. Trumpet:

0.52 mm; index about 8. Cephalothorax and abdomen generally cream-colored

with darkened areas along posterior margin of middorsal ridge, leg and wing

cases, metanotum and abdominal segments I-IV. Chaetotaxy as figured, dis-

tinctive in the following. Trumpet. Slender, rather short, darker than under-

lying integument; basal 0.5 uniform in width, apical 0.5 slightly widened

toward apex; pinna lightly oblique. Cephalothorax, Seta 1-C 4,5 branched;

7-C weak and short, double or triple; 8-C 4-6 branched; 9-C 3, 4 branched.

Abdomen. Seta 7-I usually 4 branched (3,4); 7-II usually triple (2,3); 1-II

small, 0-7 branched; 1-III-VII subequally long, about 0.5 of segment following;

1-IIf 11-15 branched; 1-IV 7-13 branched; 1-V 6-9 branched; 1-VI 7, 8 branched;

1-VII 4-6 branched; 5-IV-VI short, as long as setae 1-IV-VI; 5-IV 6-10 bran-

ched; 5-V, VI 6-8 branched; 5-VI 3-7 branched; 6-III-VI subequally long, usu-

ally 6 branched (5-7); 9-VIE shorter than 9-VIII, 5,6 branched; 9-VIII very near

caudolateral angle, 6-9 branched; 4-VIII usually 3,4 branched (2-5). Paddle.

Outer and inner parts whitish or transparent; midrib strong and dark.

LARVA (Fig. 42). Head: 0.65 mm. Siphon: 0.9 mm; index 6. Saddle:

0.32 mm; siphon/saddle ratio about 3. Distinguished from other members of

the Szttens Subgroup by the shape and the very long subventral tufts of siphon;

presence of 0-8 very large comb scales and by the following. Head. Seta 1-C

dark, stout, spiniform; 4-C minute, single and about 0.25 of the distance be-

tween bases of the pair; 5, 6-C double; 7-C 6-9 branched; 8,9,10,11 and 12-C

very weak and short; 13-C usually triple (2-4); 14-C 2-4 branched. Antennal

shaft as long as head length or slightly longer; spicules strong and numerous;

1-A strong, dark, fan-shaped and about 20 branched; 2, 3-A dark, strong,

bristlelike, subapical. Mental plate small, with 4-6 blunt lateral teeth on each

side of median tooth. Thovax. Not spiculated; seta 4-P double; 7-P triple;

8-P double; 8, 9-M 5 branched; 7 and 9-T 5 branched; 12-T single; 13-T strong,

0,6 branched. Abdomen. Not spiculated; dorsal setae 1-5 and ventral setae

Sirivanakarn: Subgenus Culex in the Oriental Region 105

9-13 of segments I-VI very weak, short, rather inconspicuous; 6-I, II double;

7-I single; 6-III- VI double; 1-III-VI 4, 5 branched; seta 1-VII strong, 4,5 bran-

ched; 4-VII single or double; 7- VII single; 10-VII usually triple (3, 4); 12-VII

single. Comb scales large, 5-8, in 1 or 2 rows; individual scale with stout

apical spine and basal fringe of fine spicules; 1-VIII weak, 4, 5 branched; 2

and 4-VIII single; 3-VIII 6-9 branched; 5-VIII usually double (2,3). Saddle

complete; seta 1-X double, 2-X single; 4-X with 6 pairs of setae; anal gills

about 1.5 times as long as saddle. Szphon. Slender and moderately long, dis-

tally strongly curved dorsad; pecten with 7-9 teeth in basal 0.25, 3,4 distal

teeth barbed with 9,10 graded denticles; siphonal tufts usually 8 pairs (total 16),

sometimes 7 (total 14); 5,6 proximal pairs very strong, long, ventral or sub-

ventral, usually double, 3,4 times as long as siphonal width at point of attach-

ment; 2 distal pair small, inserted subventrally or laterally; seta 2-S strong,

dark, simple, spiniform or with accessory basal denticles; median caudal

filament of spiracular apparatus absent; seta 9 of ventral valve dark and strong-

ly hooked.

TYPE-DATA. (1) Taeniorhynchus whitmorei Giles, Holotype 2?*, Camp

Stotsenberg, Pampanga, Luzon, PHILIPPINES, caught in woods, 1904, E. R.

Whitmore (BM); (2) Taeniorhynchus argenteus Ludlow, Holotype 2*, Pampanga,

Luzon, PHILIPPINES (USMN); (3) Leucomyia plegepennis Theobald, Holotype

2*, Kobe, Honshu, JAPAN, 14 September 1903 (BM); (4) Culex albus Leicester,

Holotype for (in poor condition), Kuala Lumpur, Selangor), MALAYA (PENIN-

SULAR MALAYSIA), 15 December 1903 (BM); (5) Culex loricatus Leicester,

Holotype 2*, Kuala Lumpur, (Selangor), MALAYA (PENINSULAR MALAYSIA)

1914, G. F. Leicester (BM).

DISTRIBUTION. Widespread throughout the Oriental region, extending

north and northeast into China, Korea, Japan and maritime provinces of USSR;

east and southeast into New Guinea. Material examined. 902 specimens: 115,

7002, 87 L; 16 individual rearings (4 pupal, 12 larval).

INDIA. Bombay: Deccan; Kawar; Tavargatti; Bengal: Lalmanirhat; Assam:

Chabua; Dooma Dooma; Dibrugarh; Rupsi; Tezpur; Ledo; 140, 812, 23 L.

BANGLADESH. Sylhet; 29.

SRI LANKA. Central Province: Kandy; Sabavagamuwa: Ratnapura; Uggal-

kaltota; 90°, 309.

THAILAND. Nan; Chiang Mai; Lampang; Lampoon; Nakhon Nayok; Khon

Khaen? Chon Buri; Yalax33¢ ; 2592; 9 Ly Ap; 1p.

VIETNAM. Saigon; Phu Bei: Ankhe; Vinh Thanh; Plei Ku; Duc My; Cu Chi;

Dilinh; Phu Loi; Pailai; Quinhan; Danang: 4c°, 799, 27 c)

MALAYSIA. Peninsular Malaysia: Kedah-Kg. Peng Besar; Perak-

Chenderiang; 30, 12, 3 lp; Malaysia: Sabah-Tambunan; 1%, 39.

INDONESIA. Java: Dijarkata; Bogor; Pelabuhan Rotu; Mt. Salak; 20", 349,

4 lp.

PHILIPPINES. Luzon: Mt. Isarog; Rizal; Samal Batoan; Olongapo; Barrio

Dolores; Mabalacat; San Fernando; Tala; Los Banos; Trinidad; Subic Bay;

Bagiuo; Mindoro: San Jose; Leyte; Samar: Guirang; Calaccad; 350", 1689,

25° Li, 4 1p. ,

CHINA,’ Hainan Is) 3/604) 129,

TAIWAN. Wan-Luan; Chao-Chow; Fengshen, Sze-Chun; 20%.

JAPAN. Kyoto; 42, Ryukyus: Okinawa: Chizuka; 19.

KOREA. Chinhae; 4, 992, 3 L.

Additional records from the literature: NEW GUINEA (Brug and Bonne-

Wepster 1947: 187); BURMA (Barraud 1934: 407); USSR: Maritime Province

(Monchadskii 1951: 281); CHINA: Chekiang, Fukien; Kiangsi; Kwangtung (Feng

106 Contrib, Amer. Ent.Inst., vol) 12, no, "2/7 1976

19381301),

TAXONOMIC DISCUSSION. Culex whitmorei is distinct in all stages and

can be readily separated from all other Oriental species of the Szfzens Group

and Subgroups as indicated in the keys and as described above. The adults of

whitmorei are superficially similar to gelidus in the color of the scales on the

head and mesonotum but differ from the latter rather strikingly in (1) presence

of speckling on the anterior surface of the femora of fore- and midlegs; (2) less

dense mesonotal scales; (3) scales on prescutellar space and scutellar lobes

pale; (4) broad plume scales on wing veins R,,, and branches of M and Cu; (5)

large median triangular basal pale patches of he abdominal terga II-VII and in

the male by (6) ventral setae of palpal segment 3 distally flattened or scalelike.

Culex whitmorei is now assigned to the Sitzens Subgroup to which it exhibits

a strong affinity in the phallosome of the male genitalia. In previous studies,

whitmorei has been considered as a member of the Gelidus Subgroup or series

according to the scheme of classification by Edwards (1932) which, as discussed

under that Subgroup, is apparently unjustified. Because of the distinctive larva,

pupa, adult and male genitalic characters, Iam placing whitmorezi in a distinct

complex of the Szfzens Subgroup. Due to the paucity of reared specimens from

most of the areas within the reported range, it is not possible to determine

whether there is local or geographical differentiation among the various popula-

tions of whitmorei. The adults which I have examined are very variable in size;

those from Korea and Japan are larger (wing length 3.7 mm) than the ones from

Southeast Asian countries but this difference does not appear to be correlated

with any modification in the male genitalia or larval stages. The affinities of

whitmorei with other species within the Szizens Group are not clear. It is

rather similar to pseudovishnui of the Vishnuzi Subgroup in the larval comb

scales but resembles alis and its related forms in phallosome of the male geni-

talia.

BIONOMICS. Culex whitmorei is a fresh water ground pool form. The lar-

vae and pupae have been frequently collected in shallow pools, puddles and

ditches in rice fields containing numerous grasses. On occasion, they have

been found in pools and animal tracks near stream margins. The elevation

ranged from sea level to 1,500 m. They have been collected in association

with the immatures of C. tritaeniorhynchus, C. pseudovishnui and C. vishnut;

An. philippinensis Ludlow, An. barbirostris Van der Wulp and An. vagus and

Aedes mediolineatus (Theobald). The relative abundance of whitmorei during

different seasons of the year is not known but from the amount of material

obtained in several collections in Thailand, Malaysia and in the Philippines it

is apparently not as abundant or common as [ritaeniorhynchus and other forms

of the Vishnui Subgroup which utilize similar habitats. Several females of

whitmorei came from light traps, buffalo and human baited traps. They were

reported to bite man in early evening and at night (Feng 1938, Hsiao and Bohart

1946, Delfinado 1966), however, the natural or preferred host is unknown.

Hsiao (1945, 1948) demonstrated that females can be experimentally infected

with Wuchereria bancrofti. Despite the report of an isolation of Japanese

encephalitis virus from this species in India (Carey et al. 1968), the impor-

tance of whitmorei as a vector of JE virus has not been subsequently confirmed.

Sirivanakarn: Subgenus Culex in the Oriental Region 107

22. CULEX (CULEX) ANNULIROSTRIS SKUSE

(Figs. 43, 44)

Culex annulirostris Skuse 1889: 1737 (2); Edwards 1924: 394 (adult, taxonomy).

Culex (Culex) annulirostris Skuse, Edwards 1932: 204 (taxonomy); Knight and

Hurlbut 1949: 30 (“*, 2, P, L*, taxonomy); Penn 1945: 81 (P*); Bohart

1956: 80 (o*, 2, L*); Belkin 1962: 207 (**, °, P*, L*); Dobrotworsky

1965: 207. (C*, 2@*.U*); Delfinado.1966¢139 (¢ *, 9, 1%); Baisas 1974: 67

(Stole eP, ans

Culex (Culex) palmi Baisas 1938: 207 (“*, 2*, P*, L); Delfinado 1966: 139

(synonymy).

For complete synonymy, see Stone, Knight and Starcke (1959).

FEMALE. Wing: 3.0-4.0 mm (average 3.8 mm). Forefemur: 1.4-2.0

mm (average 1.8 mm). Proboscis: 1.6-2.2 mm (average 2.0 mm). Abdomen:

2.9mm. Medium to large species; in general as described and figured for

sitiens to which it is extremely similar, differing in the following features.

Head, Median pale ring of proboscis broader, 0. 25-0. 30 of total length.

Cibarial Armature. Essentially similar to sitiens, differing slightly in having

lateral teeth more widely spaced. Legs. Anterior lower surface of forefemur

evenly sprinkled with pale scales along the whole length; anterior surface of

foretibia with a row of small yellowish dots among anterior bristles; anterior

surface of hindfemur extensively speckled with pale scales. Wing. Scales on

all veins denser; plume scales on veins R,, M, Ro, R and R4,5 narrower,

longer and somewhat linear. Abdomen. 'Terga II-VI or II-V with basal pale

bands which are moderately to strongly produced in middle; basal band on ter-

sum II sometimes incomplete or represented by median triangular pale patch;

terga VII-VIII with narrow, even basal bands; tergum VIII usually broadly pale

towards apex. Genitalia. Tergum IX with lateral row of 10-12 setae; postgeni-

tal plate truncate or slightly emarginate on caudal margin; vaginal sclerite U-

shaped, weakly sclerotized and pale; insula with a dense tuft of 8-10 relatively

strong setae.

MALE. Differing from sitiens as described for female and in the following.

Head. Palpal segment 3 with broader median pale band on dorsal surface, ven-

tral surface with 2 or more rows of short, dark and fine hairlike setae along

the whole length, ventrolateral tuft with 40 or more bristles from basal 0.25

to apex; apex of segment 5 with broader pale scaled tip. Ventral tuft proximad

of median pale ring of proboscis weaker and with fewer hairs which are entirely

pale. (

MALE GENITALIA (Fig. 43). Differing from sztiens and other members

of the Sitzens Subgroup in the following. Segment IX. Tergal lobe with row of

6,7 setae, sometimes more. Subapical lobe. Proximal division usually with

1,2 weak setae basad of setae a-c; seta a shorter than setae 0 and c; seta 0

longest and strongest; sternal surface with distinct patch of several short setae;

leaf g of distal division broad, with acuminate apex; seta 2 usually weak and

short; 3 setae in group d-/f, all slender, rodlike, the most distal longest and

strongest. Distimere. Normal; dorsal subapical crest of spicules usually not

developed, sometimes present, very weakly developed. Phallosome. Lateral

plate strongly sclerotized and heavily pigmented; inner division with a promin-

ent, strongly spinose apical sternal lobe; apical tergal angle usually provided

with some strong denticles, subapical tergal surface with a lobe bearing 2-5

short broad foliform processes; outer division represented by a long and large

108 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

acute spine, diverging laterad. Pyroctiger. Apical crown large and dark;

basal sternal process absent; paraproct and cercal sclerite well sclerotized

and moderately pigmented; cercal setae usually 3, sometimes 2.

PUPA (Fig. 43). Abdomen: 3.0-3.5 mm (average 3.2 mm). Paddle:

0.78-0.91 mm (average 0.85 mm). Trumpet: 0.78 mm; index 5-6. Cephalo-

thorax and abdomen generally pale with considerable amount of dark tinge over

most parts of the integument. Complete chaetotaxy as figured, particularly

distinctive in the following. Trumpet. Thick, dark and moderately long; pinna

slightly to moderately oblique; apex of pinna with distinct notch dorsad. Cephalo-

thorax. Seta 1-C usually 4 branched (3, 4); 4-C 3, 4 branched; 5-C 5 branched;

8-C usually 5 branched (4-6), 9-C 2-4 branched. Metanotum. Seta 10-C 6-12

branched; 11-C usually double (2,3). Abdomen. Seta 1-II 5-12 branched; 1-III,

IV 8-12 branched; 1-V 7, 8 branched; 1-VI 6, 7 branched; 1-VII 4, 5 branched;

6-II1,IV 3, 4 branched (3-5); 6-V, VI usually 5 branched (4, 5); 5-IV-VI shorter

than segments following; 5-IV 5-7 branched; 5-V, VI usually double (2,3). Pad-

dle. Very broad and pale; midrib weak and lightly pigmented.

LARVA (Fig. 44). Head: 0.78 mm. Siphon: 1.3-1.8 mm (average 1.6

mm); index 3-7. Saddle: 0.32 mm; siphon/saddle ratio about 4. Very similar

to sitiens, differing from it in the following characters. Head, Seta 1-C dark,

slender, spiniform; setae 5,6 and 7-C and seta 1-A stronger and darker.

Thorax. All setae darker and stronger. Abdomen. All setae darker and

stronger; 7-I double; 1-III, IV usually 5 branched (4-6); 1-V, VI usually 6 bran-

ched (5-7); 13-IJI-V usually 5 branched (4-6); seta 1-VII usually 7 branched

(4-8). Comb scales 30-45. Saddle complete, with very dark basal ring; ven-

tral caudal margin slightly indented; 1-X usually triple (2-5); 2-X usually 4

branched (3-5); 4-X with 6 pairs of stronger and darker setae; anal gills slender,

tubular, tapering into a blunt point apically; 1,2 times as long as saddle.

Siphon. Relatively thicker, pecten with 10-15 teeth, 3-4 distal teeth barbed

with 10 graded denticles; siphonal tufts 5 or 6 pairs, 6-10 branched each, form-

ing a double prominent row subvenitrally; proximal 4-5 pairs strong and sub-

equally long, as long as or slightly shorter than siphonal width at point of

attachment; most distal pair shortest, about 0.5 of adjacent proximal pair;

median caudal filament of spiracular apparatus poorly developed or absent.

TYPE-DATA. (1) Culex annulirostris Skuse, Syntypes ¢, Blue Mountain

and Berowra, New South Wales, AUSTRALIA (MM). (2) Culex (C.) palmi

Baisas, Holotype “, Paranaque, Rizal Province, Luzon, PHILIPPINES, from

a fresh water pool, 3 October 1935, F. E. Baisas (PBH; lost or destroyed).

DISTRIBUTION. Widespread throughout the Australasian region (Eastern

Australia, New Guinea and adjacent islands), Micronesia and South Pacific,

with the range extending west into the Philippines and adjacent islands west of

New Guinea in Indonesia. Material examined. 319 specimens: 1040, 1389,

77 L; 110 individual rearings (9 pupal, 101 larval).

PHILIPPINES. Luzon: Subic Naval Base; Mt. Makiling; Manila; San

Fernando; Rizal, Wack Wack; San Fabian; Camp Nichol; Los Banos; Mabalacat

(Dolores), Balucan; Panay: Mloita; Catanduanas Is,: Paganiban; Mindoro: San

Jose; Caminawit Point; Leyte: Tacloban; Burauen; Palo; Dulag; Samar: Os-

mena; Mindanao; 99%, 1389, 77 L, 9p, 101 lp.

INDONESIA. Moluccas: Morotai Is.; Ceram; West Irian (Dutch New

Guinea): Merauke; 5c,

Additional records from the literature. INDONESIA: Sulawesi (Celebes);

Kepulauan Bangai (Archipelagos); Saemba; Alor; Tenimber Islands (Brug and

Bonne-Wepster 1947); PAPUA-NEW GUINEA: Papua (Peters and Christian

1963); MICRONESIA: Islands in Central Pacific (Bohart 1956); SOUTH PACIFIC

Sirivanakarn: Subgenus Culex in the Oriental Region 109

(Belkin 1962); AUSTRALIA: Northern Territory, Queensland; New South Wales;

(Belkin 1962; Dobrotworsky 1965).

TAXONOMIC DISCUSSION. Culex annulirostris is one of the most common

and widespread species in the islands of the Philippines which are in the west-

ernmost part of its range. It is also common in the islands in eastern Indone-

sia west of New Guinea according to Brug and Bonne-Wepster (1947) and Bonne-

Wepster (1954). The Philippine material and that from the Moluccas, Ceram

and West Irian in Indonesia agrees very well with that from Micronesia (Bohart

1956) and the South Pacific (Belkin 1962). The Philippine populations of annuli-

rostris are very variable in the characters of all stages except for the diagnos-

tic ones. Due to the lack of adequately reared material and insufficient field

data, no attempt was made to analyse the detailed morphology and to determine

whether there is any local differentiation among the annulirostris populations in

the Philippines.

The synonymy of palmi Baisas 1938 from Luzon with annulirostris by Del-

finado (1966: 138) was apparently based on the description and figures of the

male genitalia and other stages by Baisas (1938: 207-9). Although the type

specimens of palmi no longer exist, its synonymy with annulirostris is accept-

able judging from the original descriptions and figure of that species by Baisas

(Joes sity);

Culex annulirostris can be readily separated from sitiens (to which it is

exceedingly similar in the general adult characters) by the male palpus, pro-

boscis and genitalia, larval and pupal stages as indicated in the keys and in the

above diagnosis. The separation of the females of the 2 species is very difficult.

Caution should be taken in using the characters noted in the key and mentioned

above.

The affinity of annulirostris appears to be closer to the members of the

sitiens complex than to any others in the Sifzens Group. The unique male

phallosome of annulirostris is intermediate between certain species of the

Bitaeniorhynchus Subgroup, especially starckeae Stone and Knight 1958 as

described and figured by Belkin (1962: 217) from the South Pacific and the

members of the sitiens complex. In the complex phallosome structure, annu-

lirostris exhibits a similarity to starckeae in the simple, long and heavy spine

of the outer division, but resembles members of the sitiens complex in the

spinose apical sternal lobe of the inner division. There is also similarity in

the development of the outer division of the male phallosome between annuli-

rostris and vicinus (Taylor 1916) from Australia and bihamatus Edwards 1926

from Timor (Sirivanakarn 1974). It appears most probable that all of these

species are related and fall into a single natural lineage.

BIONOMICS. Culex annulirostris utilizes a variety of ground pools as

breeding habitats. In the Philippines, the immatures have been frequently

collected from general ground pools, ponds, swamps and rice fields containing

emergent vegetation. On occasion, they have been collected also from cacao

shells and bamboo stumps. The water is fresh and clear or polluted with de-

cayed organic matter. Baisas (1938: 208) also collected the larvae (as palmi)

in salt bed pools. Inthe South Pacific, annulirostris has been reported to

utilize almost any type of water, fresh or brackish on the ground or artificial

containers, in open sunlit as well as in strongly shaded sites (Belkin 1962).

Most of the adults from the Philippines have been collected biting and only a

few in light traps. The preferred hosts are not known, but man is seldom

attacked (Baisas 1974). Elsewhere in Micronesia (Bohart 1956), New Guinea

(Assem and Bonne-Wepster 1964) and the South Pacific (Belkin 1962), annuli-

rostris has been considered as a nuisance or serious pest of man. Females

110 Gontrib.: Amers) But.nInsti “vol. 12; no.72,.1976

from these areas readily attack man and are vicious biters in the evening.

The medical importance of annulirostris in the Philippines is not known.

In New Guinea, it has been considered to be one of the important vectors of

Wuchereria bancrofti (de Rook 1957a; de Rook and van Dijk 1959). In Australia,

it has been reported to be naturally infected with 44 strains of Sindbis, Murray

Valley encephalitis, Koongol and other unidentified agents and one strain of

Belmont virus (Doherty et al. 1963, 1968, 1972) and has been considered to be

a probable vector of arboviruses in cattle and buffalo (Standfast and Dyce 1972).

VISHNUI SUBGROUP

The Vishnui Subgroup essentially conforms to the Sitzens Subgroup charac-

ters and is chiefly characterized in the male genitalia by (1) inner division of

phallosome not expanded to form a prominent lobe sternad; (2) outer division of

phallosome broad, lobelike and usually with 1,2 strong mesal spines and 1

sternal spine, sometimes with several spines or denticles on lateral margin;

in the pupa by pinna of trumpet lightly oblique; in the larva by (1) seta 4-P usu-

ally double, sometimes single or multibranched; (2) siphon usually slender, thin,

moderately to very long; (3) median caudal filament of spiracular apparatus

usually well developed and (4) anal gills long, slender, tubular or fusiform,

varied from 1.5 to 4.0 times as long as saddle.

DISCUSSION. The Vishnui Subgroup is largely restricted to Southeast Asia

and adjacent tropical areas with a small extension into the west as far as the

Mediterranean and the Ethiopian areas; into the north as far as the eastern

Palearctic, including China, Manchuria, maritime provinces of USSR, Korea

and Japan and into the east as far as the Solomons in the South Pacific and

northern Australia. In Southeast Asia and adjacent areas, the Vishnui Subgroup

is highly complex and taxonomically difficult, comprising various nominal taxa

which are extremely similar in the adults and the male genitalia, but are strong-

ly differentiated from one another in the larva and sometimes also in the pupa.

The identity and the taxonomic status of the species involved can only be defin-

itely determined by an examination of all stages associated through individual

rearing. In this study 8 species are recognized in the Vishnui Subgroup. Most

of these species are based on the study of material from Southeast Asia, India,

Sri Lanka, China, Japan, Korea and New Guinea. These 8 species can be

roughly classified into 3 complexes: vishnui, tritaeniorhynchus and whitei on

the basis of the male sexual characters, the color of scales on the mesonotum,

the male genitalia and the pupal and larval stages.

The vishnui complex as interpreted here is restricted to vishnui Theobald

1901; pseudovishnui Colless 1957; perplexus Leicester 1908; alzenus Colless

1957; philippinensis n. sp. and incognitus Baisas 1938. In addition, there are

3 other extralimital forms which apparently belong to this complex on the basis

of the male genitalia and larval stages: permixtus Hsieh and Liao 1956, cheni

Ho 1963 from China and omani Belkin 1962 from the South Pacific. This com-

plex is characterized in the male by ventral surface of palpal segment 3 with

transparent scalelike setae; in the adults of both sexes by anterior 0. 7 of the

mesonotum usually predominantly covered with pale beige, yellow or golden

scales or with partially dark and pale scales; in the male genitalia by apical

tergal crown of inner division of the phallosome with 4-6 relatively strong

and long fingerlike or toothlike processes and by the presence of 1,2 strong

mesal and 1 sternal spine in the outer division; in the pupa by pale yellow or

slightly darkened and relatively long trumpet (0.6-0.9 mm) and by seta 1-1

Sirivanakarn: Subgenus Culex in the Oriental Region 111

usually less than 10 branched; in the larva by seta 7-I single; comb scales

varied; usually small, numerous, with apical fringe terminating into a strong

median spine or sometimes fewer (6,7 in number), large, spinelike or numer-

ous, with a mixture of large and small ones and by the presence of 6-8 pairs

(total 12-16) of siphonal tufts.

The tritaeniorhynchus complex as recognized here includes only tritaenior-

hynchus which is the most common and widespread of all members of the Vish-

nui Subgroup. It is chiefly characterized in the male by the ventral surface of

palpal segment 3 with dark, fine, hairlike setae; in the adults by anterior 0.7

of the mesonotum predominantly covered with dark brown or golden brown

‘scales; in the male genitalia by apical tergal crown of inner division of phallo-

some with relatively weaker and shorter fingerlike processes; in the pupa by

brownish and shorter trumpet (0. 50-0.65 mm) and seta 1-II more than 10 bran-

ched; in the larva by seta 7-I double; comb scales small, numerous, subequal,

with normal, rounded apical fringe of evenly fine spicules and by the presence

of 5,6 pairs (total 10-12) of siphonal tufts.

The whitei complex is represented only by whitei which essentially conforms

to the vishnui complex in the general adult characters, including male palpus

but is strongly differentiated from all members of that complex in the male

phallosome by the presence of several lateral and mesal spines or denticles

in the outer division; in the larva by seta 7-I double, or sometimes triple; comb

scales small, numerous, with apical fringe terminating into a weak median

spine and by the presence of 5 pairs (total 10) of siphonal tufts.

23. CULEX (CULEX) VISHNUI THEOBALD

(Figs. 16, 45, 46, 47)

Culex vishnui Theobald 1901a: 355 (2*); Edwards 1913b: 233 (2, in part); Bar-

raud 1924a: 996 (“*, 2, in part).

Culex annulus Theobald 1901a: 358 (¢); Giles 1902: 405 (2); Blanchard 1905: 293

(2); Leicester 1908: 144 (2); Edwards 1913b: 233 (synonymy with tritaeni-

orhynchus Giles).

Culex pseudoinfula Theobald 1911b: 237 (2); Edwards 1913b: 233 (synonymy);

Colless 1957a: 93 (synonymy with annulus Theobald).

Culex (Culex) tritaeniorhynchus Giles, Edwards 1932: 204 (taxonomy, in part).

Culex (Culex) vishnui Theobald, Edwards 1932: 205 (taxonomy, in part);

Barraud 1934: 400 (“*, , in part); Reuben 1969: 643 (o'*, 2*, L*,

taxonomy).

Culex (Culex) adelae Baisas 1938: 200 (“*, 2, P*, L*); Colless 1957a: 93

(synonymy with annulus Theobald).

Culex (Culex) annulus Theobald, Colless 1957a: 93 (, 9, L*, revalidated)

Delfinado' 1966: 138: (o'*, °2;) L*); Bram: £967ax 205 (oo, 9*,) B*,; Li*):

Reuben 1969: 643 (synonymy with vishnui Theobald); Baisas 1974: 86

(Ree Dyes),

FEMALE (Fig. 16, 45). Wing: 2.7-3.6 mm (average 3.2 mm). Fore-

femur: 1.3-1.7 mm (average 1.6 mm). Proboscis: 1.5-2.0 mm (average

1.8mm). Abdomen: 2.2-2.7 mm (average 2.5mm). Medium sized species;

coloration variable. Head, Narrow decumbent scales of vertex yellowish

white; erect scales varying from partially pale golden brown or dull yellow in

center, dark brown on posterolateral areas to entirely brownish; lateral patch

of broad scales pale beige to whitish, very distinct. Palpus about 0.2 of pro-

132 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

boscis; apex of segment 4 tipped with white scales; segment 3 and basal portion

of segment 4 with or without some scattered pale scales on inner dorsal sur-

face. Proboscis slightly longer than forefemur; median pale ring broad, 0. 20-

0.25 of proboscis length; portions distad and proximad of median ring usually

entirely dark, sometimes sprinkled with a few pale scales. Cibarial Armature.

Cibarial teeth 26-28 in number, in concave row which is slightly produced at

middle; median 4-6 teeth narrow, elongate and tapered into a fine point apically;

lateral teeth coarser, shorter, with abruptly pointed or truncate apices. Thor-

ax. Mesonotal integument reddish to dark brown, sometimes pale brown;

mesonotal scales narrow, moderately dense, color variable, from predomin-

antly pale yellowish to brownish with mottling of brownish or yellowish scales

not forming definite patterns on anterior 0. 70-0. 75; scales on antealar and

supraalar areas rather coarse and largely whitish or yellowish; scales on

prescutellar space and scutellar lobes pale beige to whitish. Apn and ppn with

several narrow scales which are largely pale yellowish or concolorous with

scales on mesonotum. Pleural integument paler or same color as mesonotum;

ppl, upper corner and lower posterior border of stp and upper anterior mep

with distinct pale scale patches; a few pale scales among upper mep bristles

always present. Legs, Color of anterior surface of femora variable; anterior

surface of fore- and midfemora entirely dark or lightly to strongly speckled

with pale scales not contrasting sharply with dark scaled background, some-

times largely pale with sprinkling of dark scales; anterior surface of hindfemur

with longitudinal pale stripe extending from base to near apex, dorsal area

laterad of stripe usually entirely dark or sometimes speckled with pale scales;

all tibiae and tarsomere 1 entirely dark, lightly speckled or with poorly defined

pale stripe on anterior surface; tarsomeres 2-4 of all legs with or without

scattered pale scales proximad and distad of apical and basal pale bands. Wing.

Scales on all veins usually entirely dark except for short pale streak on posteri-

or surface at extreme base of vein C; sometimes with a few pale scales scat-

tered among dark ones on veins C, Sc, R and Cu; plume scales on veins Ro,

Rg and R4,5 narrow. Abdomen. Tergum I with median caudal scale patch

which is partially pale in middle and dark laterally; terga II- VII with relatively

broad, even basal transverse pale bands connected with elongate basolateral

pale spots; tergum VIII with basal and apical pale bands or largely pale; sterna

largely pale or with pattern of basal pale bands and apical dark bands as on

terga. Genitalia. Essentially similar to most members of the Szftzens Group;

tergum IX with lateral row of 4-6 setae; postgenital plate truncate or lightly

emarginate on caudal margin; vaginal sclerite well sclerotized and pigmented,

U or V-shaped; insula with a dense tuft of 8-10 setae.

MALE (Fig. 45). Essentially similar to female except for sexual charac-

ters. Head, Palpus exceeding proboscis by full length of segment 5; segment

2 with distinct apical pale band on dorsal surface; segment 3 with dorsal median

pale band, apical 0.25 with ventrolateral tuft of 15-20 dark bristles, ventral

surface with a row of short and pale translucent scale like setae which are pro-

eressively longer in apical 0.5; segments 4 and 5 moderately to strongly plu-

mose, each with narrow basal pale band on dorsal surface; apex of segment 9

usually tipped with some pale scales. Proboscis with distinct ventral tuft of

0-10 long hairs at base of median pale ring.

MALE GENITALIA (Fig. 46). Segment IX. Tergal lobe with a row of 4-7

moderately long setae. Basimere. Conical, rather slender, about 0.27 mm

in length. Subapical lobe. Setae a-c of proximal division subequally long, rod-

like; setae d-f of distal division consists of 3,4 narrow, bladelike or rodlike

setae, the distalmost of which is longest, the other 2,3 shorter, subequal, all

Sirivanakarn: Subgenus Culex in the Oriental Region 113

with hooked apex; leaf g broad, acuminate or rounded apically; seta 4 moder-

ately strong, but not distinctly flattened. Distimere. Sickle-shaped; subapical

dorsal surface lightly annulate and with a very weakly developed crest of fine

spicules (visible under 43X objective); subapical claw short, distally flat and

somewhat bilobed. Phallosome. Inner division of lateral plate with promin-

ent apical tergal crown of 4,5 strong, fingerlike processes, all curved, por-

rect or projecting caudad, the most mesal longest, lateral ones gradually

shorter, with apices reaching beyond apical margin of sternal spiculate por-

tion; the latter irregular, not produced into an angle laterally; outer division

with a strong, divergent mesal spine and a strong sternal spine which is strong-

ly curved or angled with apex pointing basad; lateral basal process moderately

broad and long. Proctiger. Apical crown medium sized; basal sternal process

dark, stout, long, curved sternad; subbasal process well developed, 0.1-0.2

of the length of basal sternal process; cercal setae 2,3.

PUPA (Fig. 46). Abdomen: 2.5-3.0 mm (average 2.7 mm). Paddle.

0.7-0.8 mm (average 0.74 mm). Trumpet. 0.60-0.78 mm (average 0.7 mm);

index usually more than 6 (5-9). Cephalothorax and abdomen cream-colored

or yellowish white, without characteristic darkened areas. Complete chaeto-

taxy as figured, the following are diagnostic. Trumpet. Yellowish, more or

less uniformly cylindrical from base to apex and moderately long; pinna slight-

ly oblique, apical margin truncate of slightly emarginate. Cephalothorax. Seta

1-C triple; 8-C usually double (2,3); 9-C double. Metanotum. Seta 10-C usu-

ally 4,5 branched (3-7); 11-C double; 12-C usually 4 branched (2-4), rather

weak and short, about 0.5 of 11-C. Abdomen. Seta 1-II usually 7, 8 branched

(6-9); 1-III usually 7,8 branched (6-12); 1-IV usually 6, 7 branched (4-7); 1-V,

VI 4, 5 branched; 1-VII usually 3,4 branched (2-4); 5-IV usually triple (2, 3);

d-V, VI usually double (1,2), longer than segment following; 6-III, IV usually

triple (2,3); 6-V, VI usually 4 branched (3, 4); 4,5-VII usually double (1,2); 9-

VII usually 4 branched (3, 4); 4-VIII double; 9-VIII 6-10 branched. Paddle.

Broad and largely pale to almost transparent; midrib weak and pale; apical

setae 1,2-P present, minute.

LARVA (Fig. 47). Head: 0.70-0.78 mm (average 0.74 mm). Siphon:

1.3-1.8 mm (average 1.5 mm); index usually 7(6-8). Saddle: 0.32 mm;

siphon/saddle ratio about 4.5. This species is distinguished from the rest

of the vishnui complex and other members of the Sitiens Group by the presence

of a distinct patch of numerous minute spicules on the thorax and by the follow-

ing additional characters. Head, Yellowish white; seta 1-C dark, slender,

simple, spiniform; 5,6-C double; 7-C pale, moderately flattened and usually

8,9 branched (7-11) 13-C double or triple. Antenna as long as head and very

pale in middle portion; spicules strong and numerous; seta 1-A large, fan-

shaped, about 25 branched; 2,3-A dark, bristlelike and subapical. Mental

plate with 6 lateral teeth on each side of median tooth. Thorax. Broad patch

of minute spicules visible under 10X objective; usually largely restricted to

dorsal pleural area; seta 4-P strong, double, its length nearly as long as

setae 1-3-P. Abdomen. Setae 6-III-VI double; 7-I single; 1-III-VI 0.25-0. 50

of 6-III-VI; 1-III, IV 3, 4 branched; 1-V, VI 4,5 branched; seta 1-VII 9-7 bran-

ched; 4, 7,10 and 12-VII single. Comb scales numerous, about 22, arranged

in a broad oval patch; all scales similar in size and length, with apical fringe

terminating to a moderately strong median spine. Saddle complete; posterior

caudal margin lightly infuscate and spiculate; seta 1-X weak, usually triple

(2,3); 2-X usually triple (2-4); anal gills slender, tubular or fusiform, as long

as or slightly longer than saddle. Siphon. Slender, straight or slightly

curved dorsad in distal portion; color same as head capsule; length very vari-

114 Contrib. ‘Amer. Ent. inst. vel.-12,“no. 2, 1976

able, usually 1.4-1.5 mm; pecten with 12-15 teeth; 3-4 distal teeth usually

with 5-7 basal denticles and a long and very strong apical denticle, the latter

straight or strongly curved; siphonal tufts strong, distinct, usually 7 pairs

(or total 14), sometimes 6 pairs (or total 12); placed distad of pecten; 4,5

proximal pairs closely spaced, forming a prominent irregular row on ventral

surface, each 8-10 branched, the most proximal longest, slightly exceeding

siphonal width at point of attachment, the rest gradually shorter; the most

distal pair short, subventral; 1 lateral pair as long as most distal, placed

slightly beyond midpoint of siphon; seta 2-S very weak, pale and short; median

caudal filament of spiracular apparatus well developed.

TYPE-DATA. (1) Culex vishnui Theobald, Holotype 2*, Madras, INDIA,

Cornwall (BM); (2) Culex annulus Theobald, Holotype °*, Tai Po, Pokfulam,

HONG KONG, C. Thomson (BM); (3) Culex pseudoinfula Theobald, Holotype 9,

Pasuruan and Samarang, Java, INDONESIA (AM); (4) Culex (Culex) adelae

Baisas, Holotype “ with associated pupal and larval skins, Tungkong Manga,

(Luzon), PHILIPPINES, March 1937 (PHB; lost or destroyed).

DISTRIBUTION. Widespread throughout Southeast Asia with extension into

the north and northeast as far as South China, Hong Kong, Taiwan and the

Ryukyus, east as far as the Philippines, south as far as Java, Indonesia and

west as far as India and Sri Lanka. Material examined: 1,548 specimens:

5050, 6492, 394 L; 508 individual rearings (133 pupal, 375 larval).

INDIA. Madras: Cornwall (type-locality); North Arcot, Virus Research

Center (Poona); Chinglepet, Thandalam Village; Assam: N. Tinsukia; Dibru-

garh; Dooma Dooma; Rupsi; Chabua; Lalmanirhat; Bengal: Calcutta, N. Salt

Lake, Dum Dum; 560, 572, 21'L, 3 lp.

BANGLADESH. Sylhet; 7“, 162, 1 L, 8p, 5 Ip.

BURMA. Rangoon: 4, 59, 5p.

SRI LANKA. Peradeniya, Kandy District; 1°, 19, 2 lp (associated adults

missing). |

THAILAND. Chiang Mai; Lampang; Phrae; Nan; Khon Kaen; Kalasin;

Sakhon Nakhon; Nakhon Phanom; Udon Thani; Nakhon Ratchasima; Nakhon Nayok;

Sara Buri; Phet Buri; Pathum Thani; Nonthaburi; Kanchanaburi; Chon Buri;

Prachuap Khiri Khan; Ranong; Nakhon Si Thammarat; Surat Thani; Satum; 80¢,

1602," 1861, F2°p, 16 Ip,

VIETNAM. Duc My; Can Tho; Phu Bei; Bien Hoa; Van Gia; Ban Me Thuot;

Quang Nam; Saigon; An-Khe; Phu Ioi; Qui Nhon; Di An; Nha Trang; Antuc; Long

Binh? Pleiku;' 10%, 102,°35 L,

CAMBODIA. Phnom Penh; Ari Gsatr; 24, 99.

MALAYSIA. Peninsular Malaysia: -Selangor: Subang Forest Reserve, Ran-

tau Panjang; Puchong; Kuala Lumpur; Ulu Langat; Univ. Malaysia; Pahang:

Kuala Lipis; Merapoh; C. Highlands; Kuantan; Pdg. Tungku, Merapoh; Treng-

ganu: Kuala Brang; Johore: Kg. Kahang; Labis; Kelantan: Gua Musang; Perak:

Lahat; Chiore Forest Reserve; Kuala Dipang; Senlu; Kuala Kangsar,; Sungei

Siput; Trong; Perlis: Pedang Besar; Kedah: Changlun; Sintok Forest Reserve;

160¢, 2019, 60 L, 37 p; 1161p. Malaysia:-Sabah: Beaufort; Tenom; Papar;

Kota Kinabalu (Jesselton); Mankubau; 4%, 99, 1 p.

SINGAPORE. Localities not specified; 11%, 159, 3 L, 1p, 21 lp.

INDONESIA. Java: Djakarta; Bogor; Sumatra: Bengkulu; Sulawesi:

Minahassa; 2%, 49, 1L.

PHILIPPINES. Luzon: Rizal, Wack Wack; Batangas; Subic Bay; Agoo La

Union; Bataon; Camp Stotsenberg; Subic Naval Base, Clark Air Base; N.

Vizcaya; Mindoro: San Jose; Samar: Osmena, Combayen; Leyte: Abuyog;

Palo; Gulf; Palawan: Iwahig; Calicoan Is.; Culion Is. ; Mindanao; Zamboanga,

Sirivanakarn: Subgenus Culex in the Oriental Region 115

Kolambugan; Lanao; Ori Missamis; Kabakan; 1170, 118°, 80 L, 7p, 165 lp.

HONG KONG. 4, 192, 5L, 2p, 3 lp.

CHINA. Canton; Nanking; Chekiang; 20, 29.

TAIWAN. Chow Chow; Ping Tung; Sze-Chun; Sun-Moon Lake; Taipei;

Lung Tan; Hsin Chu Hsien; 210, 1792, 36 lp.

JAPAN. Ryukyus: Okinawa-Yaeyama, Chizuka; 20, 1, 1 lp.

TAXONOMIC DISCUSSION. The identity and the taxonomic status of

vishnui was confused and highly controversial until Reuben (1969) redescribed

all stages from reared specimens collected in the type locality in Madras,

South India. I am following Reuben (loc. cit.) in synonymizing annulus Theo-

bald 1901 with vishnui Theobald 1901 and consider all Southeast Asian forms

previously attributed to annulus by Colless (1957a), Delfinado (1966) and Bram

(1967a) as this species. In addition, Pseudoinfula Theobald 1911 from Indo-

nesia and adelae Baisas from the Philippines, previously synonymized with

annulus by Colless (1957a), are transferred to the synonymy of this species.

It is also quite possible that permixtus Hsieh and Liao (1956) from Amoy and

cheni Ho (1963) from Canton, China are synonyms of vishnui judging from the

figures and the descriptions by these authors.

Culex vishnui as interpreted here is an extremely variable species. The

type and topotypic adults from Madras which have been re-examined differ

strikingly from those in Southeast Asia and other parts of India, including

Bengal and Assam in the presence of speckling of pale scales on the femora of

the legs but are identical to the latter populations in all other stages. The

speckling of the legs of typical vishnui adults is usually better developed in the

females than in the males, and as pointed out by Reuben (1969), it is also sub-

ject to considerable individual variation. The typical adult forms also exhibit

a striking difference in having the scales on the anterior 0.7 of the mesonotum

predominantly pale beige in contrast to usually predominantly brownish among

the majority of the Southeast Asian populations. It appears most probable that

the type form of vishnui is nothing more than a local ecotypic modification of

a single widespread species and that the Southeast Asian species (previously

known as annulus) is probably at most subspecifically distinct. When the re-

sults of experimental crosses among the 2 populations are available, it should

be possible to determine the specific or subspecific status of these forms.

Culex vishnui is clearly differentiated from other members of the vishnui

complex in the larva and sometimes also in the pupa as indicated in the keys

and as described above. In Southeast Asia, separation of vishnuz adults,

especially the female, from pseudovishnui and tritaeniorhynchus has been

rather tenuous. In sorting out adults from light trap or general field collec-

tions, the vishnui females are more likely to be confused with pseudovishnui

than with tritaeniorhynchus, Caution should be taken in using the key and other

characters as given in the above diagnosis.

BIONOMICS. The typical breeding sites of vishnui are general ground pools,

including puddles, ditches, ponds, animal or wheel tracks and swampy ground

containing numerous grasses and other aquatic vegetation in open or cleared

land such as rice fields. It has also been found breeding in pools at the edge

of small streams and in rock pools. The larvae and pupae have been frequent-

ly collected in association with those of C. pseudovishnui, C. tritaeniorhynchus

and Anopheles species. In extensive field studies by Reuben (1971la, 1971d,

1971e) in Madras, adult density of vishnui was reported to be lowest from

March to May, rose with the onset of the southwest monsoon during October to

November and began to decline from January to May. The vishnui larvae were

found to be most common in fallow rice fields where they were largely con-

116 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

centrated near the edge of paddy blocks during the early rice growing season.

Their occurrence apparently precedes that of pseudovishnui and tritaeniorhyn-

chus which utilize similar breeding habitats. The vishnui adults were collected

both indoors and outdoors, in animal bait-traps, hand catches from resting and

in Thailand several specimens were also collected in light traps. The pre-

ferred or natural hosts of vishnui females are pigs and birds (Colless 1959;

Reuben 1971c, 1971d), but other animals, including cow and man are also

readily attacked in the absence of the preferred host. In Taiwan, vishnui (as

annulus) was reported to feed mainly on pigs (Mitchell et al. 1973) and in Thai-

land on buffalo and cattle (Gould et al. 1974).

MEDICAL IMPORTANCE. Culex vishnui is possibly one of the important

vectors of Japanese encephalitis virus (JE) in Southeast Asia but because of

the difficulty in separating the adults from other common species, particularly

tritaeniorhynchus and pseudovishnui, its significance in the natural transmis-

sion of the disease has not yet been clearly established. In Taiwan, Cates and

Detels (1969) found vishnui (as annulus) to be the only JE virus-positive mos-

quito and concluded that it was the important vector of this disease. Earlier,

Wang et al. (1962) isolated JE virus from both tritaeniorhynchus and fuscoce-

phala in Taiwan. Ina virus isolation study by Carey, Reuben and Myers (1968)

in Madras, India, one strain of JE virus, one strain of Sindbis and 2 strains

of Umbre were isolated from vishnui (as Culex sp. 3) and one strain of Umbre

was isolated from a mixed pool of pseudovishnui and vishnui (as Culex sp. 3).

No attempt has yet been made to determine the efficiency of vzshnui as a vector

of JE virus through experimental transmission in the laboratory. In Chiang

Mai, Thailand, Ingwavuma virus and one strain of Tembusu virus were report-

edly isolated from pools of the vishnui complex (Top et al. 1974; Gould et al.

1974).

24. CULEX (CULEX) PSEUDOVISHNUI COLLESS

| (Figs. 48, 49)

Culex (Culex) pseudovishnui Colless 1957a: 88 (o°*, 2, L*); Delfinado 1966: 147

(2, Pp L yy Bran t9Glar 222: (o*, 2, Dr); Reuben: 1969: 6483(e*,. @*,

L*); Matsuo and Ramalingam 1972: 55 (L, morphology).

Culex (Culex) neovishnui Lien 1968: 230 (“*, 2, P*, L*); Miyagi and Iha 1970:

71 (taxonomy). NEW SYNONYMY. .

Culex vishnui of Barraud 1923a: 938 (L*); Barraud 1924a: 996 (o*, 2, in part);

Borel 1930: 333 (“*, 9, L*).

Culex (Culex) vishnui of Edwards 1932: 205 (taxonomy, in part); Barraud 1934:

400 (O*,°@, To*; in part)s "Bohart and Ineram 1946::83 (co *,° 9; L*);

LaCasse and Yamaguti 1950: 236 (“, °, P, L); Bonne-Wepster 1954: 126

(o, 3, L*).

FEMALE. Wing: 3.06mm. Forefemur; 1.5 mm. Proboscis: 1.7 mm.

Abdomen: 2.3 mm. Small to medium sized species; in general extremely

similar to vishnui, differing slightly in the following characters. Head, Erect

scales of vertex pale beige or yellowish white in center, contrasting rather

sharply with dark ones on posterolateral area. Palpus usually entirely dark

except for apex of segment 4 which is sometimes tipped with pale scales.

Cibarial Armature. As described and figured for vishnui from which it is

virtually indistinguishable. Thorax. Mesonotal scales usually predominantly

pale beige, yellow or golden, with or without mottling of several dark ones

Sirivanakarn: Subgenus Culex in the Oriental Region 117

along acrostichal and dorsocentral lines and fossae; scales around prescutellar

space dark; scales on prescutellar space and scutellar lobes entirely pale.

Legs. Anterior surface of fore- and midfemora dark scaled, speckling absent;

anterior surface of hindfemur with longitudinal pale stripe extending from base

to near apex, area dorsad and laterad of stripe dark scaled; anterior surface of

tibiae and tarsomere 1 entirely dark, pale streak or stripe absent. Wing.

Scales on all veins entirely dark except for a short pale streak on posterior

surface at extreme base of vein C. Abdomen. Terga II-VII with basal pale

bands which are very variable in width, usually broad and distinct on terga II-

VI, narrow and indistinct on terga VII-VIII, sometimes extremely narrow and

not visible from above on all segments; basolateral pale spots well developed

on all terga. Genitalia. Essentially similar to vishnui, differing slightly as

follows: Tergum IX with lateral row of 3-5 setae which are more widely spread;

postgenital plate evenly rounded on caudal margin.

MALE. Essentially as described for female, differing from vishnuz in the

following features. Head. Palpal segment 3 with or without median pale band

on dorsal surface, apical 0.25 with ventrolateral rows of about 14 bristles.

Proboscis usually without ventral tuft of long hairs proximad of median pale

ring.

MALE GENITALIA (Fig. 48). Exceedingly similar to vzshnui in all details

except for more slender lateral basal process of outer division of phallosome.

PUPA (Fig. 48). Abdomen: 2.3mm. Paddle: 0.65 mm. Trumpet: 0.6

mm, index 9. Cephalothorax and abdomen cream-colored. Complete chaeto-

taxy as figured; distinguished from vishnui and other members of the vishnui

complex by the following. Trumpet. Moderately long and more or less uni-

formly cylindrical. Cephalothorax. Seta 8-C usually 4,5 branched (3-6).

Metanotum. Seta 12-C strong, subequal to 11-C, usually triple (2-4). Abdo-

men, Seta 5-IV usually 7 branched (6,7); 5-V, VI double, strong, 1.5 times

as long as segment following; 6-III, IV usually 4,5 branched (3-6); 6-V, VI

usually 5,6 branched (4-7). Paddle. Outer and inner part very pale; midrib

variable from lightly to strongly pigmented.

LARVA (Fig. 49). Head: 0.65 mm. Siphon: 1.0-1.9 mm; index 6-10.

Saddle: 0.32 mm; siphon/saddle ratio 3.5-5.0. As figured; differing from

vishnui and other members of the vishnui complex in the following combina-

tion of characters. Head. Seta 5-C usually triple (2-5); 6-C usually double,

rarely triple; 7-C dark, moderately flattened and strongly pectinate, 8-11

branched; 13-C single or double. Thorax, Spiculation absent; seta 4-P vari-

able from weak and short, 6-15 branched to strong, long and double. Abdomen,

Setae 1-III, IV usually 6,7 branched (5-9); 1-V, VI stronger than 1-III, IV, usu-

ally 7,8 branched (6-9); 6-III-VI usually triple (3,4); seta 1-VII 6-11 branched;

4,10 and 12-VII single; 7-VII double. Comb scales very large, subequal, usu-

ally 5-7, sometimes more, but not exceeding 12, in single or sometimes double

rows; individual scale with strong and long apical spine and basal fringe of fine

spicules. Saddle seta 1-X usually 4 branched (4, 5); 2-X 3, 4 branched; anal

gills slender, 1.5-2.0 times as long as saddle length. Siphon. Variable in

length; distally tapered, lightly to strongly curved dorsad; siphonal tufts 7, 8

pairs (total 14-16), 6 proximal pairs strong, 5,6 branched each, inserted sub-

ventrally, pairing more or less regular; most proximal pair longest, about 1.5

times as long as siphonal width at point of attachment; rest gradually shorter;

1,2 distal pairs weak, inserted laterally or dorsolaterally beyond midpoint of

Siphon; seta 2-S dark, slender, spiniform.

TYPE-DATA. (1) Culex (C.) pseudovishnui Colless, Holotype ?*(57/11/7)

with associated pupal and larval skins, SINGAPORE, 9 May 1957. D. H. Col-

118 Contrib. Amer, Ent. Inst., vol. 12, no. 2;°19 76

_ less (BM); (2) Culex (C.) neovishnui Lien, Holotype “ (CU 30.6) with associated

pupal and larval skins, Peiyuan, Tungho, Taitung, TAIWAN, reared from egg-

raft laid by a female caught when biting a man at night in the woods, 8-9 June

1967, J. C. Lien, W. L. Cheng and R. S. Lin (Taiwan Provincial Institute of

Infectious Diseases, Taipei, Republic of China).

DISTRIBUTION. Widespread over most parts of the Oriental region, ex-

tending north and northeast into China and Japan, east and south into New Guinea

(West Irian, Indonesia) and west into Pakistan, India and Sri Lanka. Material

examined. 1,268 specimens: 319°, 5022, 447 L; 295 individual rearings (129

pupal, 166 larval).

INDIA. Madras: North Arcot, Poona; Bihar, Purnea; Bengal: Sukna;

Darjeeling District; Calcutta, Salt Lake; Assam: Lalmanirhat; Jorhat; Rupsi;

Dibrugarh; Dooma Dooma; Chabua; Tezpur; 6 mi. N. Tinsukia; 110, 172, 30 L, —

4° "TC 1p.

BANGLADESH. Sylhet; 2%, 82, 2 p, 3 lp.

SRI LANKA. Centval Province: Peradeniya; Kandy District; Hasalaka;

Mahiyangane; Nuwara Eliya; Monaragala District, Padiyatalawa; North Central

Province: Anuradhapura District; Padaviya; Northern Province: Vavuniya

District; Sabavagamuwa Province: Ratnapura District; Panamure; 250, 599,

26.1, 3 Tp.

PAKISTAN. Lahore; 2 lp (adults missing).

THAILAND. Chiang Mai; Lampang; Udon Thani; Tak; Khon Kaen; Mae Hong

Son; Nan; Nakhon Ratchasima; Nakhon Nayok; Phet Buri; Chon Buri; Rayong;

Trat; Prachuap Khiri Khan; Ranong; Surat Thani; Krabi; Phuket; Yala; Narathi-

wat; 940, 1269, 208 L, 107 p, 15 Ip.

CAMBODIA. Phnom Penh; Ari Gsartr; 2, 89.

VIETNAM. Cantho; Cu Chi; Phu Bei; Bien Hoa; 10, 89, 11 L, 2p.

MALAYSIA. Peninsular Malaysia: Selangor-Kuala Lumpur; Sungei Besi;

Kepong; Ulu Langat; Rantau Panjang; Ulu Klang; Trengganu-Dungun; Pahang -

Pdg. Tungku; Pekan Rd.; Kuantan; Perak-Leng Gong; Sungei Siput; C. Highland

Rd.; Chiore F. R.; Maxwell's Hill; Lasah; Kuala Kangsar; Perlis -Padang

Besar; Kedah-Kg. Peng Besar; 47“, 782, 47 L, 3p, 35 1p; Malaysia: Sabah-

Tambunan; Apin; Tuaran; Kota Kinabalu (Jesselton); Kinarut; Labuan; Mt. Kina-

balu; Kuala Penya; Stipitang; Tenom; 130, 232, 2p, 7 lp.

SINGAPORE. Pulau Ubin; Henderson Rd.; 8“, 72%, 10 L, 1p, 10 lp.

INDONESIA. Kalimantan: Tarakan; Java: Djakarta; Bogor; Pelabuhan

Ratu; Sumatra: Bengkulu; Morotai (Mollucas); New Guinea (West Irian): Li

Sentani; Cyclops Mtn.; Tenah, Merah Bay; 27°, 459, 7 L.

PHILIPPINES. Luzon: Subic Bay; Pangasinan; Tayug; San Fernando;

Mindoro: San Jose; Palawan: Jwahig; Balcahan; Culion Is, ; Sanga Sanga Is,:

Lapit Lapit; Leyte; Mindanao: Kalabakan; 500, 71°, 52 L, 2 p, 12 lp.

HONG KONG. 3c, 102%, 7L, 6p, 6 lp.

TABWAN, | 40%," 29:6 3 E.) 2 Ip:

JAPAN. Kyoto; Nagasaki; Honshu (Otsu); 30 L; Ryukyus: Okinawa -Chizuka;

Tancha Onna; Chibana Misato; Shimabuku; Yaka Kin; Yaeyama; Iriomote; 35°,

402, 6L, 64 lp.

CHINA. Shanghai; 8 L.

TAXONOMIC DISCUSSION. Culex pseudovishnui is the most common and

widespread species of the vishnui complex in Southeast Asia and other parts

of the Oriental region. Until it was described and recognized by Colless

(1957a: 88-93), pseudovishnui had been partially or entirely referred to as

vishnui Theobald in all previous regional and local studies (Barraud 1923a,

1924a; Borel 1930, Bonne-Wepster and Brug 1939, Lee 1944, Bohart and

Sirivanakarn: Subgenus Culex in the Oriental Region 119

Ingram 1946, LaCasse and Yamaguti 1950 and Bonne-Wepster 1954). As

pointed out by Colless (1957a), the larvae figured and described by these

authors as vishnui were erroneous, due to misidentification.

Culex pseudovishnui can be clearly separated from the rest of the vishnui

complex and subgroup by the larval and pupal characters as indicated in the

keys and as described above. Most diagnostic of the pseudovishnui larva are

the presence of a row of 5-7 very large, spinelike comb scales; the character-

istic siphon and the number and arrangement of siphonal tufts. The pupa can

be readily recognized by seta 8-C which is usually 4, 5 branched; the pale trum-

‘pet and by the branching of seta 1-II (see key). The adults of pseudovishnui

cannot be separated from those of perplexus, alienus, incognitus and philip -

pinensis with certainty but are, with practice, readily distinguished from those

of vishnui and tritaenitorhynchus by the paler color of the erect scales of the

head and the mesonotal scales of the thorax and by the whitish longitudinal stripe

on the anterior surface of the hindfemur. The pseudovishnui males also differ

from the latter 2 species in the absence of a ventral tuft of long hairs proximad

of the median pale ring of the proboscis. The male genitalia are virtually in-

distinguishable from alienus and incognitus but can be differentiated from those

of vishnui, perplexus, philippinensis and tritaeniorhynchus by the slight differ-

ences in the phallosome and the proctiger.

Of all stages of pseudovishnut, the larvae apparently exhibit a striking,

discontinuous variation in the length and branching of seta 4-P and the number

of lateral hair tufts of the siphon. Based on these 2 larval characters, Lien

(1968: 230-4) described neovishnui from Taiwan as distinct from typical pseudo-

vishnui by having seta 4-P double and strong (weak and multiple in the type

form) and presence of 1 pair of lateral tufts (usually 2 in the type form) on the

apical half of the siphon. My study and the comparative analysis of these lar-

val characters by Matsuo and Ramalingam (1972: 55-61) of the material from

several parts of Southeast Asia and other adjacent areas, including Taiwan,

the Ryukyus and Japan reveal considerable overlap in these characters. The

larval material from Malaysia, Singapore, Thailand, etc., all exhibit the

characters typical of pseudovishnui and neovishnui without any indication of

correlated differences in other stages. Both larval forms have been found

together in the majority of collections from Thailand and Malaysia. Although

larvae of the typical pseudovishnui have not yet been found in Taiwan, the Ryu-

kyus and Japan I am rather convinced that they are only a variation within a

single species. The tentative synonymy of neovishnui Lien with pseudovishnui

as discussion by Matsuo and Ramalingam (1972) is warranted on the basis of

the present review.

BIONOMICS. The larvae and pupae of pseudovishnui came largely from

collections in various types of fresh water ground pools in rural, open country

such as rice fields at the elevation ranging from near sea level to about 46 m.

They have also been collected from pools at stream margins. Frequently they

were found in association with tritaeniorhynchus and vishnui and occasionally

with bitaeniorhynchus, mimulus and fuscocephala in the collections from Thai-

land and Malaysia. In Madras, India, Reuben (197le) reported that the larvae

were abundant and began to replace vishnui by the time rice reaches 0.3 m in

height; they appeared to be spread along the edge as well as the interior of a

paddy block. Adult density follows a similar pattern to the rainfall as in vish-

nui (Reuben 1971la). They were collected both indoors and outdoors in animal

bait traps and light traps. The females occasionally bite man, but birds, pigs

and cattle are most frequently attacked and are the principal source of blood

(Colless 1959; Reuben 1971c, 1971d).

120 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

MEDICAL IMPORTANCE. Culex pseudovishnui may be a vector of Japan-

ese encephalitis virus in the Oriental region, but because of the difficulty in

separating the adults, especially from vishnui and other dominant sympatric

forms, including tritaeniorhynchus, its role as a reservoir or carrier of this

disease or other related viruses has not yet been definitely established. Carey,

Reuben and Myers (1968) reportedly isolated one strain of Umbre virus from a

mixed pool which contained specimens of pseudovishnui and vishnui (as sp. 3)

in Madras, India. In Thailand, a strain of Tembusu virus was isolated from a

pool of Culex vishnui complex which apparently included vishnui and pseudovish-

nui (Gould et al. 1974). Nothing more is known about its medical importance.

Further virus isolation from positively identified specimens and experimental

transmission in the laboratory are required.

25. CULEX (CULEX) PERPLEXUS LEICESTER

(Figs. 50, 51)

Culex perplexus Leicester 1908: 150 (%, &).

Culex (Culex) vishnui of Edwards 1932: 205 (taxonomy); Barraud 1934: 400

(o*, 2, L*, in part).

Culex (Culex) perplexus Leicester, Colless 1957a: 95 (“, 9, L*, revalidated);

Bram 19674::221 (*,;° 2, L*).

FEMALE. Wing: 3.4mm. Forefemur: 1.6 mm. Proboscis 1.6 mm.

Abdomen: 2.3mm. As described by Colless (1957: 95-7); very similar to

pseudovishnui and other members of the vishnui complex; differing in larger

size, generally darker in coloration and in the following additional characters.

Head, Erect scales of vertex predominantly yellowish to golden except for a

few dark ones on posterolateral area. Palpus usually completely dark scaled;

apex of segment 4 entirely dark or sometimes tipped with pale scales. Pro-

boscis thicker, as long as or sometimes shorter than forefemur; median pale

ring narrower, varying from 0.10 to 0.13 of proboscis length. Thorax, Meso-

notal integument darker to almost blackish; scales on anterior 0.7 of mesonotum

usually entirely cream-colored or yellow, sometimes largely dark in middle,

pale on marginal areas; scales on posterior 0.3 cephalad and laterad of pre-

scutellar space usually entirely dark, sometimes with variable amount of pale

scales forming streaks; scales in prescutellar space entirely dark or partially

pale; scales in scutellar lobes pale. Legs. Tarsomeres 1-4 of all legs with

very narrow apical and basal pale bands. Wing, All scales dark; base of vein

C without a distinct pale line on posterior surface; plume scales on veins R45

and branches of M and Cu relatively broad clavate. Abdomen. Terga II-VII

entirely dark or with narrow basal pale bands; basolateral pale spots present

on at least terga III-VII.

MALE. Differing from other members of the vishnui complex as described

for female and in the following. Head, Palpal segment 2 without apical pale

band; segment 3 entirely dark or with median pale band, apical 0.2 with ventral

lateral tuft of 10 bristles. Proboscis with a distinct ventral tuft of 5, 6 long

hairs at base of median pale ring.

MALE GENITALIA (Fig. 50). Exceedingly similar to other members of

the vishnui complex, differing in having lateral basal process of outer division

of phallosome larger, more or less rounded and in having weaker and shorter

fingerlike processes of apical tergal crown of inner division.

PUPA (Fig. 50). Abdomen: 2.5-3.0 mm (average: 2.8 mm). Paddle:

Sirivanakarn: Subgenus Culex in the Oriental Region 121

0.65-0.78 mm (average 0.72 mm). Trumpet 0.70-0.83 mm (average 0. 72

mm); index 8-11. Coloration very variable from yellowish, with or without

striking pattern of dark and light areas on cephalothorax and abdomen. Com-

plete chaetotaxy as figured; distinctive in the following combination. Trumpet.

Dark brown, relatively slender, long and uniformly cylindrical or slightly

swollen in middle, narrowed towards apex; pinna short, lightly oblique, apical

margin truncate. Cephalothorax. Seta 1-C usually triple (3, 4); 8-C usually

double (2,3); 9-C usually double (1,2). Abdomen. Seta 5-IV 6,7 branched

(6-9); 5-V usually triple (2-5); 5-VI double; 6-III, IV weak, usually triple (3-6);

6-V, VI stronger, usually 4 branched (3-6). Paddle. Midrib moderately to

strongly pigmented; areas along midrib lightly to strongly pigmented, other

parts of paddle pale.

LARVA (Fig. 51). Head: 0.65 mm. Siphon: 1.1-2.3 mm (average 1.8

mm); index 6-11. Saddle: 0.34 mm; siphon/saddle 4.0-4.5. Coloration of

head capsule, siphon and saddle variable as in the pupa; differing from other

members of the vishnui complex particularly in the following combination of

characters. Head, Seta 1-C, dark, stout, spiniform, with or without acces-

sory lateral spicules; 5-C 3,4 branched (2-5); 6-C usually double (2,3); 7-C

very dark, strongly flattened, 10-15 branched and strongly pectinate; 13-C

usually double (2,3). Thorax, Spiculation absent; seta 4-P strong, double; 7-P

usually triple (3, 4); 8-P usually double (2,3). Abdomen. Setae 6-III, IV usu-

ally triple (3,4); 6-V, VI usually double (2,3); 1-III-VI 8-10 branched; seta 1-VII

8-10 branched; 4-VII usually single; 7-VII double or triple; 10-VII double; 12-

VII single. Comb scales 12-20 in a small oval patch; size of scales intermedi-

ate between pseudovishnui and vishnui, all subequal, with strong apical spine

and lateral fringe of several fine spicules; seta 1-VIII dark, flattened, 4,5

branched and strongly barbed; 2-VIII usually single, sometimes double; 3- VIII

6, 7 branched and strongly barbed; 5-VIII 4-6 branched. Saddle complete,

darker or concolorous with siphon; seta 1-X 4,5 branched; 2-X triple; 4-X usu-

ally with 6 pairs of setae, sometimes 5.5 pairs; anal gills pale, slender, 1. 0-

1.5 times as long as saddle. Szphon. Yellowish to dark brownish; slender,

straight and distally tapered; length variable; pecten teeth 9-12 in number;

siphonal tufts 6,7 pairs (total 12,14) widely spaced, 5-6 proximal pairs in-

serted subventrally beyond pecten, 4,5 branched each; 1 pair lateral in apical

0.5 of siphon, 2-4 branched; most proximal 1.0-1.5 times as long as siphonal

width at point of attachment, rest gradually shorter, lateral pair about as long

as siphonal width at point of insertion; seta 2-S slender, spiniform; median

caudal filament very well developed.

TYPE-DATA. Lectotype ?*, Kuala Lumpur, (elangor), MALAYA (PENIN-

SULAR MALAYSIA), pupa from large stagnant pool filled with floating water

weeds near houses, 2 December 1903, G. F. Leicester (BM; selection of Bram

1967a: 222). |

DISTRIBUTION. Known only from Thailand, Malaysia, Singapore, Indonesia

and India. Material examined. 102 specimens: 230°, 299, 50 L; 29 individual

rearings (16 pupal, 13 larval).

INDIA. Andaman Is.; 2%, 39.

THAILAND. Pyrachin Buri: Ban Bu Phram; Tvat: Ban Salak Phet; Phet

Buri; Prachuap Khiri Khan: Khlong Hin Chaung; 100, 162, 37 L, 16 p, 7 lp.

MALAYSIA. Peninsular Malaysia: Selangor-Kuala Lumpur; Trengganu:

Kg. Tapah, Kuala Brang; Perak: Sungei Siput; 10°, 12, 4 L. Malaysia: Sabah-

Tenom; 60 489.7 I

INDONESIA. Kalimantan: Sotek; 2 L.

SINGAPORE. 4c, 49, 6 lp.

122 Contrib; Amen; Bat) Inst.s volo cl2ycno0.:2).210' 16

TAXONOMIC DISCUSSION. Culex perplexus can be readily separated from

other members of the vishnui complex in the larva and pupa as indicated in the

keys and as described above. The most distinctive features of the larva of

perplexus are the number and type of the comb scales, the siphon shape, the

number, relative length and the arrangement of the siphonal tufts and those of

the pupa are the shape, length and color of the trumpet, the branching of ab-

dominal setae 5-IV, V and the color of paddle midrib. Except for the diagnostic

chaetotaxy and other distinctive features, the larva and pupa of perplexus ex-

hibit considerable variation in pigmentation of the integument, particularly in

the material from Prachin Buri, Thailand. The specimens from stream pools

which contain numerous dead leaves under heavy forest shade are the darkest

and exhibit a striking color pattern, whereas those which came from swamp

pools containing living aquatic vegetation under partial shade are much paler

and exhibit no color pattern. The dark and pale larval and pupal forms do not

appear to be correlated with any differences in the adults and it seems most

likely that these variations are environmentally induced.

The adults of perplexus are also very variable, particularly in the pre-

sence or absence of basal pale bands on the abdominal terga. The adults which

exhibit completely dark abdominal terga can be readily separated from pseudo-

vishnui and alienus whereas those which exhibit basal pale bands are usually

distinguished from the latter 2 species by (1) darker integument of the mesono-

tum, (2) relatively narrower median pale ring of the proboscis, (3) relatively

shorter proboscis, (4) broader plume scales on veins R4,5 and branches of M

and Cu, (5) narrower apical and basal pale bands of tarsomeres 1-4 of the legs,

(6) male palpal segment 2 completely dark or without distinct apical band on

dorsal surface and (7) male proboscis with a distinct ventral tuft of long hairs

at the base of median pale ring.

Among the members of the vishnui complex, perplexus is apparently most

closely related to pseudovishnui, Both species exhibit only a slight overlap in

the breeding site, but appear to be ecologically distinct; perplexus usually

breeds in pools under forest shade while pseudovishnui usually breeds in the

open in rice fields. |

BIONOMICS. Culex perplexus appears to be an uncommon forest form

whose breeding habitats are restricted to fresh water ground pools such as

pools in dried up stream beds or at stream margins, swamps and ponds. All

collections from stream pools usually contain numerous decayed leaves and

those from swamps and ponds usually contain some green algae. These habi-

tats are under heavy or partial shade of primary or secondary rain forest at

elevations ranging from 200 to 300 m. The immatures were collected in asso-

ciation with specimens of C. (Eumelanomyia) malayi (Leicester), C. (Lopho-

ceraomyia) infantulus, C. (Lophoceraomyia) quadripalpis (Edwards), C. (Culi-

ciomyia) pallidothorax Theobald; C. (Culiciomyia) scanloni Bram; C, (Culex)

bitaeniorhynchus, C. (Culex) pseudovishnui, Uranotaenia annandalei Barraud

and Anopheles barbirostris. Most of the adults were obtained from rearing the

larvae or pupae. The feeding habits of the females and their role in transmis-

sion disease are unknown.

26. CULEX (CULEX) ALIENUS COLLESS

(Figs. 52, 53)

Culex (Culex) alienus Colless 1957a: 97 (%, 9, L*); Bram 1967a: 202 (o*, &,

fo).

Sirivanakarn: Subgenus Culex in the Oriental Region 123

FEMALE. Wing: 3.1mm. Forefemur: 1.4 mm. Proboscis: 1.7 mm.

Abdomen: 2.5mm. Extremely similar to pseudovishnui and other members of

the vishnui complex, differing slightly in the following. Head, Erect scales

of vertex predominantly pale yellowish or bronzy except for a few dark ones on

posterolateral areas. Palpus segment 4 always tipped with pale scales on its

apex. Proboscis distinctly longer than forefemur; median pale ring moderately

broad. Thorax. Mesonotal integument brownish; scales on anterior 0.7 of

mesonotum always pale beige or pale yellowish; posterior 0.3 of mesonotum

largely dark brown scaled cephalad and laterad of prescutellar space; scales

on prescutellar space largely dark except for a few pale ones on posterior cau-

dal margin. Legs and Wing. Essentially as described for pseudovishnut.

Abdomen. Terga II-VII always with basal pale bands.

MALE. Essentially as described for female, differing from pseudovishnui

particularly in having a ventral tuft of about 6 long hairs at base of median pale

ring of proboscis, these hairs are 2 to 3 times as long as labial width.

MALE GENITALIA (Fig. 52). Exceedingly similar to pseudovishnui from

which it can not be separated with certainty; differing from vzshnui and perplex -

us Slightly in having smaller lateral basal process of the outer division of the

phallosome.

PUPA (Fig. 52). Abdomen: 2.5-3.2 mm (average 2.9 mm). Paddle:

0.71-0.78 mm (average 0.75 mm). Trumpet: 0.60-0.72 mm (average 0.65

mm); index 9-10. Very similar to vishnui and pseudovishnui. Cephalothorax

and abdomen cream-colored. Complete chaetotaxy as figured; the following are

diagnostic. Trumpet. Slightly darker than underlying integument and more or

less uniformly cylindrical from base to apex. Cephalothorax. Seta 8-C usually

triple (2-5); 9-C double. Abdomen. Seta 5-IV usually 4,5 branched (4-6); 5-V,

VI double; 6-III, IV usually triple (2-4); 6-V usually 4 branched (3, 4); 6-VI usu-

ally 5 branched (3-6). Paddle. Inner and outer parts very pale, without infus-

cate areas; midrib weak and pale.

LARVA (Fig. 53). Head: 0.74 mm. Siphon: 1.4-1.6 mm (average 1.5

mm); index 9. Saddle: 0.36 mm; siphon/saddle ratio 4-5. Differing from

other members of the vishnui complex in the shape and the strong subventral

tufts of siphon and in the following. Head, Yellowish with indefinite darkened

area; seta 1-C relatively long; 4-C shorter than the distance between bases of

the pair; 5-C usually double (2,3); 6-C double; 7-C weak and pale, 8-12 bran-

ched; 13-C usually 4 branched (4,5). Thorax. Spiculation absent; 4-P strong,

usually double (1,2); 7-P double or triple; 8-P usually double, sometimes

single. Abdomen. Setae 1-III-VI usually double (2-4); 6-III-VI all double, seta

1-VII 4,5 branched; 4, 7,10 and 12-VII single. Comb scales small, 13-20 ina

small oval patch, scales in posterior row slightly longer than those in anterior

row, all terminating in an acute apical spine. Saddle concolorous with siphon;

seta 1-X double or triple; 2-X triple; anal gills slender, tubular, about 3 times

as long as saddle length. Szphon. Yellowish or brownish; slender and distally

strongly tapered; siphonal tufts 5.5-6 pairs (total 11-12) in number, one of

which is lateral, the rest subventral; pairing regular; 3 proximal subventral

pairs very strong, long, double or triple, 4-5 times as long as siphonal width

at point of attachment; distal subventral pairs shorter, 3,4 branched; lateral

pair inserted in apical 0.5 of siphon, 3,4 branched, about 3 times as long as

siphon width at point of insertion; seta 2-S dark, relatively strong and long;

median caudal filament developed.

TYPE-DATA. Holotype ¢* (LT31) and whole larva morphotype, Long Teb-

angan, Fourth Division, Sarawak (BORNEO), August 1953, D. H. Colless (BM).

DISTRIBUTION. Presently known only from Thailand, Vietnam, Malaysia

124 Contrib, Amer, BotIngt a. vols 1ayn0e.2,3h076

and Singapore. Material examined. 58 specimens: 16%, 1692, 26 L; 19 indivi-

dual rearings (5 pupal, 14 larval).

THAILAND. Trat;9L.

VIETNAM. Saigon; Cu-Chi; 8 L.

MALAYSIA. Peninsular Malaysia: Kelantan-Tumpat; Kedah: Kg. Bagan;

Perak: Chenderiang; 11%, 72, 1 L, 8lp. Malaysia: Sarawak (type-locality);

3c, a7) 8 L..(skins),,.2.p¢ Saben: Sipitang:: Tarakan; 20.62.07. L,. 8p): 6 lp.

SINGAPORE. 2 L.

TAXONOMIC DISCUSSION. Culex alienus is apparently one of the uncom-

mon members of the vishnuz complex and like perplexus, appears to be re-

stricted to certain areas connected with or adjacent to the Malaya Peninsula.

It is possible that it may also be discovered among the neighboring islands of

Indonesia when more extensive larval surveys are made. Of all stages, the

larva of alienus is most distinctive and can be readily separated from all mem-

bers of the vishnuz complex by the features of the siphon, number and type of

the comb scales and branching of certain setae. The adults of alzenus are dif-

ficult to separate from pseudovishnui and perplexus except through a close com-

parison using slight differences in the color of scales on the mesonotum as in-

dicated above. It can usually be distinguished from pseudovishnui by scales on

the anterior 0.7 of the mesonotum entirely pale and the scales on the prescutel-

lar space mostly dark and from perplexus by the generally paler coloration of

the thorax and broader median pale ring of the proboscis. The male genitalia

are virtually indistinguishable from pseudovishnui but differ slightly from per-

plexus and vishnui in having a more slender lateral basal process of the outer

division of the phallosome. The pupa, although showing some overlap with all

of these species, can be separated by the combination of the chaetotaxy and

other general features as described above and as indicated in the key.

BIONOMICS. Culex alienus is typically a fresh water ground pool breeder

in open country such as in rice fields or in hills fully or partially exposed to

the sun. The immatures have been collected in wells, ditches, sandy or gravel

pools, marshy depression and ponds in association with C. pseudovishnui, C.

tritaeniorhynchus, C. gelidus, C. fuscocephala and Anopheles sp, The water

was fresh and clear, containing little or no aquatic vegetation. The elevations

ranged from sea level to about 70 m. All adults have been obtained from rear-

ing larvae and pupae. Nothing is known about the feeding habits of the females

and their disease relationships.

27. CULEX (CULEX) PHILIPPINENSIS NEW SPECIES

(Figs. 54, 55)

FEMALE. Wing: 3.4mm. Forefemur: 1.7mm. Proboscis: 1.7 mm.

Abdomen: 2.5 mm. In general extremely similar to pseudovishnui and its

closely related forms, differing slightly in the following combination of char-

acters. Head. Proboscis relatively short, equal to or slightly longer than

forefemur; median pale ring about 0.2 of total length of proboscis. Cibarial

Armature. Essentially similar to pseudovishnui; cibarial bar with about 20

teeth in a concave row; median 4-6 teeth narrow, elongate and fine, closely

spaced; lateral teeth coarser, shorter and widely spaced. Thorax. Mesonotal

integument pale brown; scales on anterior 0.7 mesonotal disc entirely pale

beige or with dark scales forming a distinct longitudinal streak on acrostichal

line and a spot in middle of fossae; scales cephalad and laterad of prescutellar

space dark brown; scales on prescutellar space entirely pale. Legs. As

Sirivanakarn: Subgenus Culex in the Oriental Region 125

described for pseudovishnui; apical and basal pale bands of tarsomeres 1-4 of

all legs moderately broad and distinct. Wing. All scales dark, narrow and

dense; base of vein C without pale scaled streak or spot on posterior surface.

Abdomen. Terga II-VII with narrow basal pale bands and elongate basolateral

pale spot; sterna largely pale or with pattern of basal pale bands as terga.

Genitalia. Essentially as described and figured for vishnui and pseudovishnui;

tergum IX with a lateral row of 6,7 setae on each side; postgenital plate with

truncate caudal margin; vaginal sclerite U-shaped and well sclerotized; insula

with a tuft of 8-11 strong setae.

MALE. In general as described for female except for broader basal bands

on abdominal terga; differing from other members of the vishnuz complex par-

ticularly in the following features. Head. Palpus rather short, exceeding pro-

boscis by about 0.5 of length of segment 5; segment 2 and 3 without pale scales

forming bands on dorsal surface; short scalelike setae on ventral surface of

segment 3 apparently fewer, apical 0.1 of segment 3 with weak tuft of 6, 7

bristles; segment 4 and 5 reduced in length, weakly to moderately plumose and

entirely dark or with poorly defined basal bands on dorsal surface. Proboscis

without ventral tuft of hairs at base of median pale ring.

MALE GENITALIA (Fig. 54). Very similar to other members of the

vishnui complex in the basimere, subapical lobe, distimere and type of phallo-

some, differing strikingly in the almost complete absence of basal sternal pro-

cess of the proctiger and in the following additional detail. Subapical lobe.

Leaf g of distal division relatively narrower and shorter. Phallosome,. Inner

division with apical tergal crown of 4,5 weaker and shorter fingerlike processes;

sternal spine of outer division very large, somewhat resembling szfzens; lateral

basal process of outer division relatively large, broad and rounded. Proctiger.

Apical crown of spicules dark, medium sized; basal sternal process of para-

proct very short, rudimentary to almost completely absent; subbasal process

absent; cercal setae 2-3.

PUPA (Fig. 54). Abdomen: 2.5mm. Paddle: 0.74 mm. Trumpet 0.65

mm; index 5-7. Cephalothorax and abdomen pale, cream-colored, without

definite pattern of darkened areas. All setae developed, the following diagnos-

tic. Trumpet. Moderately long and more or less uniformly cylindrical; basal

half of meatus dark, apical half pale. Cephalothovax. Setae 1-3-C usually

triple (2-4); 8-C usually triple (2-5). Abdomen. Seta 5-IV usually triple (2-4),

o-V, VI double, long, about 1.5 of segment following; 6-III-VI usually triple

(2-4); 2-VII always mesad of 1-VII; 4, 5-VII always single; 4-VIII usually single,

rarely double. Paddle. Broad and entirely pale; midrib pale or lightly pig-

mented.

LARVA (Fig. 55). Head: 0.78 mm. Siphon: 1.5-2.0 mm; index 8-11.

Saddle: 0.36 mm; siphon/saddle ratio 4.5-5.5. Complete chaetotaxy as fig-

ured, distinguished from other members of the vishnui complex by the follow-

ing features. Head. Yellowish white; seta 4-C single, as long as or slightly

longer than distance between bases of the pair; 5-C usually triple, sometimes

double; 6-C double; 7-C pale, 6-8 branched, weakly barbed; 14-C single or

double. Thorax. Spiculation absent; seta 1-M relatively long, about 0.5 of

3-M. Abdomen. Setae 1-III-VI usually triple (2-4); 6-III-VI double. Comb

scales numerous, 30-40, all small, subequal, with weak apical spine and later-

al fringe of evenly fine spicules. Saddle complete, same color as siphon; seta

1-X double; 2-X double or triple. Siphon. Slender, pale, moderately to very

long, distal portion moderately to strongly tapered, somewhat resembling ali-

enus; pecten teeth 9-12; siphonal tufts usually 6 pairs (total 12), sometimes

..9 or 7 pairs; 4 proximal pairs strong, subventral, usually triple (2-4), about

126 Contrib, Amer, Ent. Inst), vol. 12;-no. 271976

3,4 times as long as siphonal width at point of attachment; 2 distal pairs weak

and short, double or triple, the proximal pair inserted laterally, the distal

subventrally, about 1.9 times as long as siphonal width at point of insertion;

seta 2-S weak and short; median caudal filament poorly developed.

and genitalia slide, Tay-Tay, Rizal, Luzon, PHILIPPINES, November - Decem-

ber 1966, F. E. Baisas (USNM); Allotype 2 (C1 329-5) with associated

pupal and larval skins, same locality as holotype (USNM); Paratypes: 8 o*

(C1 329: C1 330; C1 338), 3 with associated pupal and larval skins, 28 ¢

(C1 301; C1 303; C1 306; C1 328; C1 329; C1 330; C1 338) with associated pupal

and larval skins, same data as holotype and allotype (USNM, BM, BBM).

DISTRIBUTION. Known only from the Philippines. Material examined.

60 specimens: 15, 402, 5 L; 56 individual larval rearings.

PHILIPPINES. Luzon: Rizal, Tay-Tay; 9°, 292, 5 L, 37 1p; Samar:

Osmena; 1°; Mindanao: San Ramon; 50, 119, 19 lp.

TAXONOMIC DISCUSSION. The above description of all stages of philippin-

ensis is based on several reared specimens collected by the late F. E. Baisas

in Luzon and by Rozeboom, Knight and Laffoon in Samar and Mindanao. This

species was fairly abundant in the localities where it was found and is probably

widespread among the Philippine islands.

Culex philippinensis is strongly differentiated from the rest of the vishnui

complex in the male genitalia by the considerable reduction to almost complete

absence of the basal sternal process of the proctiger. In the adults, the males

are also quite distinct in the reduction of the length and number of bristles of

palpal segments 4 and 5 but the females are virtually indistinguishable from

pseudovishnui. The pupa and larva can be readily separated by the characters

given in the keys and as described above. The larva of philippinensis is very

similar to alienus in most features, including the siphon and its hair tufts, but

can be separated from the latter by the presence of more numerous and finer

comb scales and by having shorter and weaker siphonal tufts.

BIONOMICS. Culex philippinensis is a typical ground pool breeder. In

Rizal, Luzon, larvae were reported to have been collected from rock pools and

in Mindanao in wheel tracks, road ruts and puddles. No further field data

were available as to the condition of the breeding sites. All adults were ob-

tained from individual rearings.

28. CULEX (CULEX) INCOGNITUS BAISAS

(Figs, «16; 56,. 57)

Culex (Culex) incognitus Baisas 1938: 203 (“*, 2*, P*, L*); Delfinado 1966:

45 (C*, f, By b*); Baisas 1974) 04 (0%) OF PL),

FEMALE (Fig. 16). Wing: 3.2 mm. Forefemur: 1.5mm. Proboscis:

1.7mm. Abdomen: 2.5mm. In general very similar to other members of

the vishnui complex, differing slightly in darker coloration (somewhat resem-

bling perplexus) and in the following characters. Head, Erect scales of vertex

whitish, yellowish brown, golden or bronzy in center, blackish posterolaterally.

Proboscis with clearly marked median pale ring which occupies about 0. 2 of

total length. Czibavrial Armature. Very distinctive; cibarial bar with a concave

row of about 26 subequal teeth which are narrow, fine, all distally tapered toa

sharp point apically. Thorax. Mesonotal integument dark brown to black;

scales on mesonotum largely golden brown or dark in middle, forming a broad

Sirivanakarn: Subgenus Culex in the Oriental Region 127

dark streak or patch on acrostichal and dorsocentral lines and fossal areas,

continued as dark streak cephalad and laterad of prescutellar space; scales

on anterior promontory, humeral, scutal angles, lateral and posterior fossal

and antealar areas and areas mesad of supraalar pale beige to golden; scales

on prescutellar space and scutellar lobe yellowish to whitish. Pleural integu-

ment dark or same color as mesonotum; pleural scale patches very distinct.

Legs. Anterior surface of fore- and midfemora entirely dark scaled, speckling

of pale scales absent; anterior surface of hindfemur with a narrow pale longitu-

dinal stripe extending from base to about 0.75 of total length; tibiae of all legs

dark scaled except for light apical ring; all tarsomeres 1-4 with narrow, but

very distinct apical and basal pale bands. Wing. All scales dark and very

dense on all veins; plume scales on veins Ro, Rg, R4;5 and branches of M and

Cu narrow; base of vein C without pale scale streak. Abdomen. Basal pale

bands on terga II-VII present or absent; basolateral pale spots always present,

extending to 0.9 or more of segment length; sterna with broad basal pale band

and broad apical dark bands. Genitalia. Essentially similar to vishnui and

pseudovishnui, differing in having all sclerites more strongly pigmented; vag-

inal sclerite V-shaped and insula with stronger tuft of 9-12 setae.

MALE. In general as described for female except for the following. Head.

Palpus exceeding proboscis by about full length of segment 5; segment 2 en-

tirely dark scaled; segment 3 with or without median pale band or scattered

pale scales on dorsal surface, apical 0.25 with 10-15 strong bristles; seg-

ments 4 and 5 strongly plumose and with distinct basal pale bands on dorsal

surface, apical 0.1-0.2 of segment 5 pale to tip. Proboscis with a small ven-

tral tuft of 2-4 hairs at base of median pale ring, hairs 2,3 times as long as

labial width at point of insertion. Abdomen. Terga II-VII always with basal

pale bands.

MALE GENITALIA (Fig. 56). Exceedingly similar to vishnui and other

members of that complex, differing slightly in the following combination of

characters. Subapical lobe, Leaf g of distal division relatively broader, api-

cally rounded or acuminate; setae d-f relatively strong, all apparently rodlike.

Phallosome. Outer division with moderately large, rounded lateral basal pro-

cess.

PUPA (Fig. 56). Abdomen: 2.52 mm. Paddle: 0.7mm. Trumpet: 0.8-

0.9 mm; index 10-15. Generally distinguished from other members of the

vishnui complex by very long trumpet. Yellowish white without definite pattern

of dark and light areas. Chaetotaxy as figured, the following combination diag-

nostic. Trumpet. Slender, very long and uniformly cylindrical; pinna narrow

and slightly oblique. Cephalothorax. Seta 1-C 4,5 branched; 8-C usually 4, 5

branched (4-6), Abdomen. Seta 5-IV 5-8 branched; 5-V usually double (2, 3);

o-VI double; 6-III-VI subequally long, usually 6,7 branched (5-8). Paddle,

Broad and pale to almost transparent; midrib weak and pale.

LARVA (Fig. 57). Head: 0.78 mm. Siphon: 2.2-3.0 mm; index 10-14.

Saddle: 0.4 mm; siphon/saddle 6-7. Distinguished from other members of

the vishnuit complex by the thin and very long siphon and in the following. Head.

Integument varying from pale, cream-colored to dark brown; seta 4-C short,

single, less than 0.9 of distance between bases of the pair; 5-C usually double

(2,3); 6-C double; 7-C dark, flattened, 8-10 branched. Thovax. Spiculation

absent; most prothoracic setae and lateral pleural setae of meso- and meta-

thorax very strong and dark; seta 4-P usually single (1,2). Abdomen. Setae

1-III-VI usually 4,5 branched (4-6); 6-III-VI double. Comb scales numerous,

27-32, strongly differentiated into 2 different types; scales in posterior row

and on ventral surface large with stout apical spine, rest of scales in anterior

128 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

rows and in center of patch small, with rounded apical fringe of even fine

spicules. Saddle usually same color as siphon, sometimes dark brown; seta

1-X usually triple (3,4); 2-X triple; anal gills 1.5-2.0 times as long as saddle.

Siphon. Very long, thin with high index and siphon/saddle ratio; pigmentation

yellowish white except for basal dark ring; pecten teeth 10-14 in number, me-

dian teeth barbed with 6,7 graded denticles; distal teeth with several fine den-

ticles in basal 0.5, apical 0.5 simple or represented by strong heavy spine;

siphonal tufts 7 or 8 pairs (total 14-16), all weak, short, subequal and widely

spaced, inserted subventrally or laterally; each tuft usually double or triple

(1-3), their length as long as or slightly longer than siphonal width at point of

attachment; median caudal filament very well developed.

TYPE-DATA. Holotype “ (No. F 159-253) with associated pupal and larval

skins, Tanque at Calauan, Laguna Province, (Luzon), PHILIPPINES, 11 May

1935, F. E. Baisas (PBH; destroyed).

DISTRIBUTION. Widely distributed in the Philippines; also known from the

Moluccas, Sulawesi, Indonesia. Material examined. 253 specimens: 990,

772, 77 L; 55 individual rearings (3 pupal, 52 larval).

PHILIPPINES. Luzon: La Union; Subic Naval Base; San Fernando; Clark

Air Base; Olongapo; Mindoro: San Jose; Samar: Osmena; Leyte: Palo;

Tacloban; Lago Lago; Bay Bay; Mindanao: Lanao; Parang; Pasanonco; San

Ramon; Jinamoc Is. ; Dinagut Is. ; Nueva Is. ; 99%, T72, 69 L, 3p, 52 Ip.

INDONESIA. Moluccas: Halmahera, Morotai; Sulawesi: Pomalaa; 8 L.

TAXONOMIC DISCUSSION. Although the type and other specimens in the

type series of incognitus, originally designated by Baisas (1938) were destroy-

ed, the description and the figures of the larval siphon, the female cibarial

armature and the male genitalia by this author agree well with all specimens

examined by me. The descriptions of all stages by Delfinado (1966: 145-7) are

also applicable to this species except for the presence of speckling on the an-

terior surface of the forefemur and the extensively pale midfemur of the adults

which are erroneous.

Culex incognitus is strongly differentiated from the rest of the vishnui

complex in the larva and pupa and can readily be recognized in these stages.

The adults are apparently more similar to perplexus than to any other species,

particularly in being generally dark or blackish in coloration and in the pre-

sence or absence of basal pale bands on the female abdominal terga. The

pattern of coloration of the mesonotal scales is variable, but appears to be

more striking than in pseudovishnui and philippinensis. For positive identifi-

cation of the incognitus females, the cibarial armature, the associated larval

and pupal stages of this species should also be examined.

The Philippine incognitus is evidently most closely related to omani Belkin

(1962: 210) from Guadalcanal, Solomon Islands, in the South Pacific. In com-

paring the specimens with the type of the latter species, I found only slight

differences in the detailed characters of all stages between the 2 forms. -Both

are almost identical in the larva except for seta 4-P which is usually single in

incognitus (double in omani) and the length of setae 1-III-VI which are relative-

ly shorter in incognitus. The adults of incognitus are darker than omani and

the apical tergal crown of the inner division of the male phallosome of zncogni-

tus is composed of 4,5 fingerlike processes (3 in omani). This evidence sug-

gests that these forms are only subspecifically distinct and that they were prob-

ably represented by a single widespread species which was subsequently differ-

entiated into distinct segregates through geographical isolation.

BIONOMICS. The Philippine incognitus is apparently common but not as

abundant or frequently encountered as vishnui, pseudovishnui or tritaenior-

Sirivanakarn: Subgenus Culex in the Oriental Region 129

hynchus. The most frequent breeding sites of zncognitus are rock pools con-

taining decayed leaves, stream pools, forest swamps and pools at margins of

creeks under heavy shade. Immatures have been collected also in grassy

pools, open swamps, pools in rice fields and according to Delfinado (1966), in

water tanks and slow streams containing green algae. Very little is known

about adult biology. Delfinado (1966) noted that females are common inside

houses and in carabao (water buffalo) bait traps, but no definite data are avail-

able on whether they bite and feed on man or other animals. Baisas (1974)

noted that it is evidently zoophilic but because of low adult density, it seems to

be of no medical importance.

29. CULEX (CULEX) TRITAENIORHYNCHUS GILES

(Figs. 16, 58, 59, 60)

Culex tritaeniorhynchus Giles 1901a: 606 (adult); Theobald 1901a: 364 (o*, 9*);

Giles 1902: 401 (, 2); Edwards 1913b: 233 (taxonomy); Edwards 1917:

224 (o, 2); Edwards 1921: 339 (L), Macfie and Ingram 1922: 180 (2*);

Barraud 1923a: 940 (L*); Barraud 1924a: 995 (o*, 2); Borel 1930: 336

(O*, Dol) Ho: 198d eee *,. 2):

Culex biroi Theobald 1905a: 82 (o*, 2); Edwards 1913b: 233 (synonymy).

Culex summorosus Dyar 1920: 180 (co); Bram 1967a: 225 (synonymy).

Culex tritaeniorhynchus var. siamensis Barraud and Christophers 1931: 283

("); Colless 1957a: 98 (synonymy with C. tritaeniorhynchus summorosus

Dyar).

Culex (Culex) tritaeniorhynchus Giles, Edwards 1932: 204 (taxonomy, in part):

Barraud 1934: 404 (o'*, 9, L*); Li and Wu 1935: 101 (L*); Stackelberg

1937: 235 (o*); Edwards 1941: 299 (, 2); Bohart 1945: 80 (o*, L);

Bohart and Ingram 1946: 81 (c"*, 2, L*); LaCasse and Yamaguti 1950:

230 (o*, 2*, P*, L*); Monchadskii 1951: 277 (L*); Iyengar and Menon

1955: 6 (L*); Khattat 1955: 165 (L*); Bohart 1956: 83 (o'*, 9, L*); Ovazza,

Hamon and Neri 1956: 165 ("*); Hara 1957: 56 (2 genitalia*); Bram 1967a:

225:(0 ORD LY,

Culex (Culex) summorosus Dyar, Baisas 1938: 196 (“*, 9*, P*, L*).

Culex (Culex) tritaeniorhynchus summorosus Dyar, Colless 1957a: 98 (c"*, 9,

Li*); Delfinade 19661190 (9) Py 1.) Baisas 19741204 (7 9 ass).

Culex (Culex) tritaeniorhynchus var. siamensis Barraud and Christophers,

Brug 1934: 515 (co); Brug 1939: 112 (L*); Bonne-Wepster and Brug 1937:

72 (%, 2*); Bonne-Wepster and Brug 1939: 1270 (L*); Bonne-Wepster

1954: 125 (o, 2*, L*); Mattingly 1956: 37 (lectotype designation).

FEMALE (Fig. 16, 58). Wing: 2.5-3.6 mm (average 3.2 mm). Fore-

femur: 1.1-1.5 mm (average 1.4 mm). Proboscis: 1.4-1.8 mm (average 1.6

mm). Abdomen: 1.8-2.5 mm (average 2.2 mm). Relatively small, slender,

reddish brown species; differing from the members of the vishnui complex in

the entirely dark brown erect scales of vertex; mesonotal and pleural integu-

ment reddish brown; mesonotal scales narrower, finer and predominantly dark

or golden brown and in the following. Head, Narrow decumbent scales of ver-

tex finer and predominantly golden except for a few pale ones drooping over

frons; erect scales relatively shorter, narrower and entirely dark brown;

lateral patch of broad scales dull yellowish. Palpus entirely dark or sometimes

with pale scales on apex of segment 4. Proboscis with a very narrow median

pale ring which occupies 0.10-0.15 of total length and with or without pale

130 Contrib. Amer. Ents Inst. vol; 12;cno. 2, 1976

scales scattering or forming streak proximad of median ring on lateral and

ventral surfaces. Cibarial Armature. Very distinctive; cibarial bar with a

concave row of about 30 long, narrow and fine teeth, all of which are sub-

equal in length and distally filamentous. Thorax. Mesonotal integument red-

dish brown or sometimes deep chestnut brown; mesonotal scales very narrow,

fine, entirely or predominantly dark brown or golden brown on anterior 0. 7 of

disc except for a few paler ones on extreme anterior promontory and lateral

marginal areas; scales cephalad and laterad of prescutellar space dark brown;

scales in middle of prescutellar space and scutellar lobes pale yellowish or

whitish. Apn and ppn with narrow dark brown scales as on anterior 0.7 of

mesonotum., Pleuron same color as mesonotum; pale scale patches on p)Dl,

upper corner and lower posterior border of stp and anterior upper mep small

or poorly developed; a few pale scales among upper mep bristles present or

absent. Legs. Anterior surface of fore- and midfemora dark brown to black;

anterior surface of hindfemur with longitudinal pale stripe poorly contrasted

with dark scaled areas on lateral dorsal surface; all tibiae dark scaled on an-

terior surface; tarsomeres 1-4 of all legs with narrow, poorly defined apical

and basal pale bands. Wing. Scales on all veins dark except for a short pale

scale line at extreme base of vein C; plume scales on veins Ro, Rg, Ry.

narrow and very dense; cell R, 4, 5 times as long as Ro,9- Abdomen, sre

II- VII with narrow basal pale bands and elongate basolateral pale spots; sterna

usually entirely pale yellowish, sometimes with small dark apicolateral patches

or spots. Genitalia, Essentially similar to members of vishnui complex from

which it can not be distinguished with certainty. Tergum IX with a lateral row

of 5, 6 setae; postgenital plate truncate or lightly emarginate on caudal margin;

vaginal sclerite well sclerotized, V- or U-shaped; insula with a tuft of 10 set-

ae,

MALE (Fig. 58). In general as described for female except for broader

basal bands on abdominal terga; differing from all members of the vishnui

complex in having short setae on ventral surface of palpal segment 3 finer,

hairlike, dark and in the following characters. Head. Palpus very thin,

exceeding proboscis by 1.0-1.5 of length of segment 5; segment 2 with or with-

out apical pale band on dorsal surface; segment 3 with or without dorsal median

pale ring, apical 0.4-0.5 with ventrolateral tuft of 20-30 strong bristles; seg-

ments 4 and 5 strongly plumose and with narrow basal pale bands on dorsal

surface; apex of segment 5 entirely dark or sometimes tipped with pale scales.

Proboscis with a prominent ventral tuft of 10 or more very long hairs at base

of median pale ring.

MALE GENITALIA (Fig. 59). Exceedingly similar to the members of the

vishnui complex, differing particularly in the following features. Subapical

lobe. Most distal seta in group d-fof distal division usually more strongly

flattened and apically blunt, club-shaped leaf g moderately to very broad.

Phallosome. Apical spiculate portion of inner division of lateral plate distally

expanded, forming a triangular lobe which tapers into a sharp point laterad;

4-6 fingerlike processes of inner division variable in size and length; usually

shorter and smaller than in all forms of vishnui complex, their apices barely

reaching beyond apical margin of sternal spiculate portion; lateral basal process

of outer division small; sternal spine more slender and gently curved basad.

Proctiger. As in vishnui and pseudovishnui; basal sternal process of paraproct

long, heavy, curved sternad; subbasal process distinct, about 0.3 of basal

sternal process; cercal setae 2 or 3.

PUPA (Fig. 59). Abdomen: 2.9mm. Paddle: 0.65 mm. Trumpet: 0.5-

0.6 mm; index 6-7. Cephalothorax and abdomen cream-colored or yellowish

Sirivanakarn: Subgenus Culex in the Oriental Region 131

without definite pattern of dark and pale areas. Chaetotaxy as figured; the

following characteristic. Trumpet. Relatively short, dark brown, strongly

contrasting with underlying integument; meatus uniform in width, apical mar-

gin truncate or lightly emarginate; pinna slightly oblique. Cephalothorax. Seta

8-C usually 5, 6 branched (4-7); 9-C as long as 8-C, usually double (2, 3).

Metanotum, Seta 12-C as longas 11-C. Abdomen. Seta 1-II multiple, more

than 10 branched; 5-IV 4-6 branched, as long as segment following; 5-V usually

double (2,3), 5-VI double, both setae as long as 5-IV; 6-III, IV usually 4 bran-

-ched (4,5); 6-V, VI longer than 6-III,IV, usually 5 branched (4-7); 4, 5-VII

double; 4-VIII double or triple. Paddle. Inner and outer part pale; basal ex-

ternal margin of outer part distinct; midrib moderately pigmented.

LARVA (Fig. 60). Head: 0.7mm. Siphon: 1.4-2.0mm; index 7-10 Sad-

dle: 0.36 mm; siphon/saddle 4.0-5.5. Similar to members of the vishnut com-

plex in general and in several details of chaetotaxy, differing particularly in

the following combination of characters. Head, Seta 1-C relatively short; 4-C

single, about as long as distance between bases of the pair; 5-C usually triple

(3,4); 7-C pale, 6-8 branched and weakly barbed. Thorax, Spiculation absent;

seta 4-P strong and double. Abdomen. Seta 7-I double; 1-III,1V weak, usually

triple (3,4); 1-V, VI longer than 1-III, IV, usually 4 branched (4, 5); 6-III usually

double (2,3); 6-IV usually triple (2,3); 6-V, VI double. Comb scales small,

numerous, about 40, in a broad oval patch; all scales apically rounded, with

even fringe of fine spicules. Saddle same color as siphon; seta 1-X usually

triple (2-4); 2-X double or triple; anal gills as long as or slightly longer than

saddle. Siphon. Slender, straight, slightly tapered distally; variable in length,

usually 1.4-1.5 mm, rarely exceeding 1.8 mm; yellowish to brownish; pecten

teeth 10-14; 4,5 distal teeth with lateral barb of 5,6 graded denticles, apical

denticle weak, tapered to a sharp point; siphonal tufts weak, widely spaced, 9

or 6 pairs (total 10-12); pairing irregular; 4,5 pairs inserted subventrally in

proximal 0. 30-0. 75 of total length; 1,2 pairs inserted laterally beyond middle;

subventral pairs usually 4 branched (3-5), subequal in length, as long as or

slightly longer than siphonal width at point of attachment; lateral and most dis-

tal pairs double or triple and slightly shorter than subventral pairs; median

caudal filament very well developed.

TYPE-DATA. (1) Culex tritaeniorhynchus Giles, Holotype 2*, Travancore,

(Madras State). INDIA, 29 June 1900, S. P. James (BM); (2) Culex biroi

Theobald, Syntypes “ and 2, Bombay, INDIA (HNM); (3) Culex summorosus

Dyar, Holotype “* with slide of genitalia, Los Banos, Luzon, PHILIPPINES

(USNM); (4) Culex tritaeniorhynchus var. siamensis Barraud and Christophers,

Lectotype o* (Y454) with slide of genitalia, Chiang Mai, SIAM (THAILAND),

J. A. Sinton (BM; selection of Mattingly 1956: 37).

DISTRIBUTION. Widely distributed throughout the Oriental region extend-

ing west into the Middle East, the Mediterranean and the Ethiopian region; north

into China, Manchuria, Maritime provinces of USSR, Japan and Korea; east into

Micronesia (Bonin Island); southeast and south as far as Sulawesi, Kabaena,

Lombok, Alor, Amboina and Java, Indonesia. Material examined. 3,904 speci-

mens: 1,003", 1,9752, 926 L; 594 individual rearings (267 pupal, 327 larval).

PAKISTAN. Lahore: Ravi Bank; 330, 502, 1 p, 12 lp.

INDIA. Bombay: Kawar; Madras: N. Arcot (Poona); Bengal: Calcutta;

Old Jalpaiguri; Lalmanirhat ; Bihar: Purnea; Assam: Dibrugarh; Chabua;

Dooma Dooma; Jorhat; Rupsi; Tezpur; 620, 1149, 42 L, 25 p, 12 lp.

BANGLADESH. Decca; Khulna; 13c", 139.

SRI LANKA. Central Province: Kandy District; Peradeniya; North Central

Province: Anuradhapura District; Padaviya; Irrigation bungalow; Sabavagamuwa

132 Contrib.) Amer, Ent... Inst), vole12)).no. 2, L976

Province: Ratnapura District; Uggalkaltata; Panamure; Southern Province:

Hambantota District, Palatupana; Uva Province: Monaragala; 5 mi. SE.

Padiyatalawa; SE. Mahiyangane; 70, 4192, 2 L, 4 lp.

BURMA. Rangoon; 6“, 182, 7p, 2 lp.

THAILAND. Chiang Mai: Muang; Doi Sutep; Hang Dong; Chang Puak;

Lampang: Ban Rong Na; Mae Ngao River; Lamphun: Lee District; Nakhon

Ratchasima: Pak Chong; Phet Buri; Sava Buri: Prabudha Bath; Thon Buri;

Bangkok; Nonthaburi; Samut Prakan; Chon Buri: Siracha; Bang La Mung;

Bang Phra; Rayong: Ben Takat Ngao; Chanthaburi: Ban Bang Kachai; Ban Laem

Sing; Ayutthaya; Pathum Thani; Prachuap Khiri Khan; Phrae: Ban Ta Loe; Khon

Kaen: Phuwiang; Suvat Thani: Koh Samui; Bang Makham; T’vat: Koh Chang;

Koh Khlum; Krabi; Ban Sai Thai; Phuket: Bang Khian; Ban Huai Luk; Ban Borae;

Ban Pa Thong; Nakhon Si Thammarat: Chang Khao; Ban Rim Thanon; Narathiwat:

Waeng; 2190, 7289, 485 L, 165 p, 43 lp.

CAMBODIA. Phnom Penh; Ari-Satr; 130, 659.

VIETNAM. Phu Bei; Cu Chi; Phu Tho; Binh Duong; Go Vap; Soc Trang;

Con Son, Cam Ranh Bay; Saigon; Can Tho; Bien Hoa; Phu Loi; Gia Dinh; 26,

182.84 Type skeedp,

MALAYSIA. Peninsular Malaysia: Selangoy-Kuala Lumpur; Kepong, Salak;

Puchong, Rantau Panjang; Bt. Kutu; Pahang: Pdg. Tungku, K. Lipis; Kuantau;

Chenderiang; Pekan Rd.; Perak: Tanjong Tualong F. R., Pusing; Senlu; Tg.

Rambutan; Kuala Kangsar; Perlis: Kg. Sg. Bahru; Kg. Gunong; Kg. Sg. Padang;

Simpang Ampat; Kg. To “ Kayaman; Kg. Baru; Kedah: Changlun; Sintok; Treng-

ganu: Kemaman, Dungun; Kelantan: Tumpat; 1850", 1999, 119 L, 39 p, 118 lp.

Malaysia: Sabah-Papar; Kuala Penyu; Beaufort; Kota Kinabalu (J esselton);

Kudat; Tarakan; Ranau (British Borneo); 38, 459, 3p, 12 lp.

SINGAPORE... 11c, 212, 14 L, 3p, 22 lp.

INDONESIA. Kalimantan (Borneo); Sumatra: Bengkulu; Java: Djakarta;

Pelabaean; Sulawesi: Makasar; 36, 499.

PHILIPPINES. Luzon: Agoo La Union; Camp Stotsenberg; Camp Gregg;

Camp Wilhelm; Rizal; Camp Nichol; Subic Naval Base; Clark Air Base; Min-

doro: San Jose; Caminawit; Samar: Osmena; Calotans, San Antonio; Leyte:

Tacloban; Gulf; Abuyag; Palo; Carigara; Santa Rosa; Dulag; Samar Giurang;

Palawan: Princesa, Iwahig; Panitan, Quezon; Mindanao: Parang; Kabakan;

Zamboanga; Pasanonga; Pettit Barrack; Torray Barrack; Tawi Tawi: Kamagon,

Tarawakan; Batu Batu; Basbas Is.: Calaccad Is, ; 288%, 4749, 142 L, 146 lp.

HONG KONG. Ho Chung, N. T.; S. Sai Kung; Tai-wan; 230, 129, 10 L,

20 o)i8lo.

TAIWAN. Taipei: Shihlin, Hsin Chu; Ping Tung: Chow-Chow; Taichung;

60 A2e aed 12 pi

JAPAN. Honshu: Chiba Pref; Shinhama; Kyoto; 30°, 49, 20 L; Ryukyus:

Okinawa -Chizuka; Chiwa; Naha; Taucha Onna; Oyama; 330, 329, 54 L, 40 lp.

KOREA. Seoul; 10, 209.

Additional records from the literature. ETHIOPIAN REGION: Gold Coast;

Nigeria; Zanzibar; Mauritius (Edwards 1941: 299); MEDITERRANEAN and

MIDDLE EAST: Palestine; Syria; Iraq; Iran; Egypt; Arabia (Barraud 1934;

406; Monchadskii 1951: 277; Mattingly and Knight 1956: 121; Lofti 1970: 402);

INDIA: Maldive Islands (Carter and Wijesundara 1948: 150); CHINA: Anhwei;

Chekiang; Fukien; Hopei; Hupeh; Kiangsi, Kiangsu; Kirin; Lianoning; Kwang-

tung; Shantung; Hainan; Amoy (Feng 1938: Hsieh and Liao 1956; Chu 1958);

INDONESIA: Nias; Siberoet; Krakatau; Celebes (Sulawesi); Kabaena, Lombok;

Alor; Amboina (Brug and Bonne-Wepster 1947: 187); MICRONESIA: Bonin Is-

lands (Bohart 1956: 84); USSR: Transcaucasia; Turkmenia; Far East; Man-

Sirivanakarn: Subgenus Culex in the Oriental Region 133

churia (Mondchadskii 1951: 277).

TAXONOMIC DISCUSSION. Iam treating tritaeniorhynchus as a single

variable species as done by Bram (1967a). The form summorosus which has

been treated as a subspecies of tritaeniorhynchus for the Southeast Asian popu-

lations by Colless (1957a: 98) and has been attributed to the Philippine popula-

tions in a specific (Baisas 1938) or subspecific sense (Delfinado 1966, Baisas

1974), isnot recognized in this study. Although the summorosus form differs

slightly from the typical Indian tritaeniorhynchus in having longer fingerlike

processes of the male phallosome as pointed out by Colless (1957a), I have

found no significant correlated differences in any other stage to justify recog-

nizing the 2 forms as Separate subspecies. The results of experimental cros-

ses between typical tritaeniorhynchus in Pakistan and the summorosus form in

Japan as discussed by Barnett (1967) also show no reproductive incompatibility

between the 2 forms. Based on this evidence and the pattern of distribution, it

appears most likely that the 2 forms are variations of a single widespread

species.

Culex tritaeniorhynchus can be readily separated from other members of the

Vishnui Subgroup as well as the Sifzens Group. In Southeast Asia, adults have

been confused with vishnui (previously referred to annulus) apparently because

they were only superficially examined and most of the previous descriptions

lacked emphasizing correlated significant features for separating the 2 species.

As discussed by Colless (1957a) and as indicated here, the tvitaeniorhynchus

females can be generally distinguished from those of vishnui by: (1) proboscis

with narrower median pale ring; (2) lateral and ventral surfaces of the probos-

cis usually with some pale scales scattering or forming a streak in basal 0. 5

proximad of the median pale ring; (3) erect scales of vertex of the head more

slender, shorter and entirely dark brown; (4) anterior 0.7 of the mesonotum

predominantly covered with narrower, finer and dark brown or golden brown

scales; (5) abdominal terga with narrower basal pale bands and (6) usually

smaller size. The female cibarial armature, the male palpus and proboscis

and the shape of the sternal apical portion of inner division of the male phallo-

some of tritaeniorhynchus are also very diagnostic, providing a clear cut sep-

aration from all members of the vishnui complex. The pupa resembles pseu-

dovishnui in most features of the chaetotaxy, but differs from it in having a

shorter brownish trumpet and seta 1-II multiple, with more than 10 branches.

The larva is very distinctive, particularly in having seta 7-I double; numer-

ous small comb scales which are apically rounded with fringe of evenly fine

Spicules and short and widely spaced siphonal tufts, not forming prominent

rows.

Culex tritaeniorhynchus has been previously considered by all authors as

a member of the vishnui complex, subgroup or group in a general or broad

sense. Present evidence shows that it is somewhat intermediate between the

vishnui complex and Barraudi Subgroup. It exhibits similarity to the members

of the vishnui complex in the male genitalia, general adult characters and the

pupal stage but is similar to the members of the Barraudi Subgroup in the lar-

val characters and in the male palpus. On this basis, Iam placing tvitaenior-

hynchus in a distinct complex of the Vishnui Subgroup.

BIONOMICS. Culex tritaeniorhynchus is a common rural species and is

most frequently encountered in the open such as rice fields, native plantations

and shallow marshes. The most common breeding sites include general ground

pools, ponds, wells, ditches and puddles containing fresh or polluted water

with numerous grasses, rice or other aquatic vegetation. These habitats are

partially shaded or fully exposed to sunlight. On occasion the immatures have

134 Contrib.) Amer; Eat. Insts, ivol 12, no. 2," 1976

have also been collected in pools near stream margins or along margins of

slow moving streams. In most localities, all stages of tvifaeniorhynchus are

very abundant, exceeding those of vishnui and pseudovishnui in number and

density throughout the year. The relative abundance and population fluctuation

of tritaeniorhynchus follows essentially a similar pattern of rainfall as in vish-

nui and pseudovishnui (Reuben 1971a, 1971d, 1971e). As in the case of pseudo-

vishnui, Reuben (1971d) reported that the tritaeniorhynchus larva becomes

dominant when the rice reaches about 0.3 m in height, replacing that of vish-

nui, The overall occurrence of tritaeniorhynchus is fairly uniform throughout

the year but, in Madras adults were reported to be most abundant in August

(Reuben 1971e). Adults were usually collected in large number at night out-

doors in cowsheds, in biting collections on cows, pigs and occasionally man

and in light traps or animal bait traps. In Madras, India, Reuben (1971e)

reported that adults were rarely taken resting in cowsheds, relatively infre-

quently in outdoor shelters and showed a preference for sugar cane fields as a

daytime resting place. In the studies by Colless (1959) in Singapore, Reuben

(1971c) and Christopher and Reuben (1971) in Madras, India, females of tvitae-

niorhynchus obtained blood meals largely from pigs and cows. In Sarawak,

Hill et al. (1969) reported it to feed mainly on pigs and fowl. The genetics of

tritaeniorhynchus are reviewed in Baker and Sakai (1974). For additional de-

tails on bionomics, consult Bram (1967a) and other references catalogued by

Travis and Labadan (1967).

MEDICAL IMPORTANCE. Culex tritaeniorhynchus has generally been

implicated as a potential or natural vector of Japanese encephalitis virus from

various parts of the Oriental region since it was experimentally shown to be

capable of transmitting this disease inJapanby Hammonetal. (1949). Inthe vicinity

of Tokyo it has been reported to be the primary and perhaps only vector of this

disease (Buescher et al. 1959). In Southeast Asia, it is undoubtedly one of the

most important vectors of this disease due to its great abundance and feeding

behavior on infected host animals. However, the epidemiology of the disease

is complicated by possible secondary vectors such as Culex (Culex) vishnui

(Cates.and Detels 1969, as annulus) and other species which have been found

naturally infected with and capable of transmitting JE virus in the laboratory,

i.e., fuscocephala (Muangman et al. 1972) and gelidus (Gould et al. 1962). In

the studies on virus isolation, tvitaeniorhynchus was found to be naturally in-

fected with 6 strains of JE virus in India (Carey et al. 1968), 13 strains of JE

virus in Thailand (Simasathien et al. 1972), one strain of JE virus and 2 strains

of Tembusu virus in Sarawak, Borneo (Macdonald et al. 1965), 5 strains of JE

virus in Vietnam (Nguyen et al. 1974) and 8 strains of JE virus in Chiang Mai,

Thailand (Gould et al. 1974). In Japan, it was found to be infected with Aino

virus (Takahashi et al. 1968). In addition, tritaeniorhynchus were also re-

ported to be naturally or experimentally infected with Brugia malayi (Carter

1948, Hu 1940), Wuchereria bancrofti (Bohart 1946, Hsiao and Bohart 1946,

Bohart and Ingram 1946, Yamada and Komori 1927, Hsiao 1948) and with dengue

virus (Rudnick and Hamon 1961). The roles of tvitaeniorhynchus as vectors of

filariasis and dengue virus have not yet been evaluated, but there is no epidemi-

ological evidence implicating the species in dengue transmission.

Sirivanakarn: Subgenus Culex in the Oriental Region 130

30. CULEX (CULEX) WHITEI BARRAUD

(Pies, 61,62)

Culex whitet Barraud 1923b: 508 (co, 2); Barraud 1924a: 998 (o'*).

Culex (Culex) whitei Barraud, Edwards 1932: 205 (taxonomy); Barraud 1934:

402 (o“*, &, L); Peters and Dewar 1956: 46 (L*); Bram 1967a: 233 (c*,

Se Ba

FEMALE. Wing: 3.2-4.0 mm (average 3.5mm). Forefemur: 1.4-1.8

mm (average 1.6 mm). Proboscis: 1.6-2.0 mm (average 1.8 mm). Abdomen:

2.9mm. Medium sized species; in general very similar to members of the

vishnui complex, differing slightly in the following combination of characters.

Head, Erect scales of vertex yellowish to golden brown in center, dark on

posterolateral area; lateral patch of broad appressed scales very distinct.

Palpus 0.25 of proboscis length; apex of segment 4 tipped with several whitish

scales, rest dark or black. Proboscis distinctly longer than forefemur, medi-

an pale ring clearly marked, occupying about 0.2 of total labial length, rest uni-

formly dark or blackish, no scattered pale scales distad or proximad of median

pale ring. Czibarial Armature. As described and figured for vishnui from which

it is virtually indistinguishable. Thorax. Anterior 0.70 of mesonotum largely

covered with pale yellowish to golden scales, with indefinite mottling of dark

brownish scales in middle, usually dark scaled cephalad and laterad of prescu-

tellar space; marginal areas with lighter scales; posterior 0.3 of mesonotum

largely pale yellowish or golden scaled, not contrasting with those on anterior

0.7; scales in middle of prescutellar space and scutellar lobes entirely pale.

Pleural scale patches relatively broad and very distinct. Legs. Anterior sur-

face of fore- and midfemora usually lightly sprinkled with some pale scales,

more or less resembling sitiens, sometimes completely dark; anterior surface

of hindfemur with a broad whitish longitudinal stripe extending from base to

near apex; foretibia entirely dark on anterior surface; mid- and hindtibiae

entirely dark or with indistinct longitudinal pale stripe in middle. Wing. All

scales dark; base of vein C without pale scale streak on posterior surface;

plume scales on veins Ro, Rg, Ray narrow, linear and dense; cell Ro 2.0 of

the length of vein Ro,3. Abdomen. Terga II-VIII usually with narrow basal

pale bands, these sometimes completely absent or represented by small median

basal pale patches or streaks; sterna largely pale with apicolateral dark scale

patches. Genitalia. As described and figured for vishnui, differing slightly in

the following. Tergum IX with a lateral row of 7,8 stronger and longer setae;

vaginal sclerite always V-shaped and darkly pigmented; insula with a stronger

tuft of 10-14 setae; posterior cowl more strongly sclerotized.

MALE. In general as described for female except for abdominal basal

bands which are always present and broader. Head. Palpus exceeding probos-

cis by 1.5 of length of segment 5; segment 3 with median pale band on dorsal

surface; ventral surface with a row of scalelike setae along whole length; apical

0.25 with ventrolateral tuft of 17-18 bristles; segments 4 and 5 strongly plu-

mose and with distinct basal pale band on dorsal surface; apex of segment 5

tipped with pale scales or sometimes entirely dark. Proboscis with a weak

ventral tuft of 6-8 long hairs at base of median pale ring.

MALE GENITALIA (Fig. 61). Essentially similar to most members of

the vishnui complex in basimere, distimere and proctiger; differing particular-

ly in the following characters. Subapical lobe. Most distal seta in group d-f

of distal division broad, club-shaped; leaf g very broad, apex rounded or acu-

136 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

minate; seta h relatively stronger and longer. Phallosome. Apical tergal

crown of inner division with 3,4 rather short fingerlike processes; outer

division with 2,3 strong mesal spines, 4,5 strong lateral spines and several

weak lateral denticles, the latter densely packed; sternal spine slender, long,

divergent laterad; lateral basal process slender and distally produced into a

rounded knob. Pvoctiger. Apical crown relatively large and with numerous

coarse spicules; basal sternal process of paraproct present, long, strong,

curved sternad; cercal setae 2. |

PUPA (Fig. 61). Abdomen: 2.7-3.0 mm (average 2.9 mm). Paddle: 0.8

mm, Trumpet: 0.62-0.65 mm (average 0.63 mm); index 5.8. Cephalothorax

and metanotum cream-colored or pale yellowish. Complete chaetotaxy as fig-

ured; the following diagnostic. Tvumpet. Meatus dark in basal half, yellowish

in apical half; pinna oblique and lightly to strongly flared; apical margin trun-

cate or lightly emarginate. Cephalothovax. Setae 1-3-C short and usually

double (1-3); 6-C about 0.5 of the length of 7-C; 8-C usually double (2-4).

Metanotum. Seta 10-C usually double or triple (2-5). Abdomen. Seta 1-III

3-9 branched; 1-IV 2-7 branched; 1-V 1-4 branched; 1-VI 2-6 branched; 1-VII

2-9 branched; 9-IV-VI slightly shorter than or as long as segment following;

0-IV usually 4 branched (3-5); 5-V, VI double; 6-III-VI 1-4 branched; 4, 5-VII

usually double (1,2); 9-VII weak, short, 2-5 branched; 4-VIII single or double;

9-VIII weak and short, subequal to 9-VII, 5-9 branched. Paddle. Inner and

outer parts very pale; midrib lightly pigmented.

LARVA (Fig. 62). Head: 0.78-0.90 mm (average 0.8 mm). Siphon:

1.6-2.4 mm (average 2.0 mm); index 8-9. Saddle: 0.38 mm; siphon/saddle

ratio 5-6. Generally similar to other members of the Vishnui Subgroup; differ-

ing particularly in the following features. Head. Creamy white; seta 4-C

single and long, about 1.5 of the distance between bases of the pair; 5-C 3-6

branched; 6-C 3, 4 branched; 14-C minute, filiform and usually single (1, 2).

Thorax. Spiculation absent; seta 4-P usually double (2,3), strong; 7-P triple;

8-P double. Abdomen. Setae 6-I, II 3,4 branched, sometimes 5 branched;

7-I usually double, sometimes triple; 1-III 1-3 branched, variable in length,

usually shorter than 6-III; 1-IV, V single, strong, of the same magnitude as

setae 6 and placed on distinct tubercles; 1-VI usually 3,4 branched (2-4), of

the same magnitude as 1-III; 6-III-VI all double. Comb scales numerous,

about 40, all small and subequal in size, in a broad oval patch; individual

scales pointed or terminating in a weak apical spine and with lateral fringe of

finer spicules. Saddle same color as siphon; seta 1-X strong, single or weak,

double or triple; 2-X double; anal gills 1-4 times as long as saddle. Szphon.

Relatively thick and long; pigmentation bright yellow to brown; pecten teeth

8-13; 3,4 distal teeth with lateral barb of 6-8 delicate denticles and a fine api-

cal denticle; siphonal tufts, weak, short and widely spaced; usually 5 pairs

(total 10), sometimes 4.5 or 6 pairs; 4 proximal pairs subventral, double or

triple, most proximal as long as siphonal width at point of attachment, rest

gradually shorter; 1 distal pair double or triple, short, inserted laterally be-

yond midpoint of siphon; median caudal filament well developed; ventral valve

of spiracular apparatus relatively broad and black.

TYPE-DATA. Lectotype °*, Haflong, Cachar Hills, Assam, INDIA,

August 1922, larvae from pools, P. J. Barraud (BM; selection of Bram 1967a:

23D),

DISTRIBUTION. Presently known only from India, Bangladesh, Thailand,

Vietnam, Peninsular Malaysia (Malaysia), the Philippines and Indonesia.

Material examined. 280 specimens: 670%, 752, 138 L; 104 individual rearings

(51 pupal, 53 larval).

Sirivanakarn: Subgenus Culex in the Oriental Region 137

INDIA. Assam: Haflong, Cachar Hills (type-locality); 9%, 49.

BANGLADESH. Sylhet; 90, 72, 15 p.

THAILAND. Chiang Mai: Payao; Chiang Dao; Fang; Lampang: Ngao; Ban

Pha Khoi; Ban Pha Daeng; Ban Rong; Mae Hong Son: Doi Chang; Nan: Pha

Daeng Khawi; Nakhon Ratchasima: Pak Chong; Phet Buri: Tha Yang; Chon

Buri: Ban Huai Luk; Ranong: Muang, Huey Pluching; Nakhon Si Thammarat:

Chaung Khao; 34, 519, 115 L, 36 p, 36 Ip.

VIETNAM. Saigon; Ankhe; Antuc; Quang Tri; 2%, 14 L.

MALAYSIA. Peninsular Malaysia: Kedah-Sintok F. R.; Johore: Kota

Tings Nai Bate Wey Anes BOs LOT dy, hia

INDONESIA. Cevam: Piroe, 1°; Lombok; 2 L.

PHILIPPINES. Luzon: Subic Naval Base; San Fabian; Pangasinan; Laguna;

Leyte: Mt. Lobi; 6%, 69, 14 lp.

Additional records from the literature. NEPAL (Peters and Dewar 1956).

TAXONOMIC DISCUSSION. Culex whitei is apparently an uncommon species

but exhibits a broad range of distribution from Nepal to the east as far as the

Philippines and Ceram, Indonesia. The occurrence of this species in Penin-

sular Malaysia, the Philippines and Ceram, Indonesia constitute additional new

records. Its presence in the Philippines, noted by Baisas (1938: 196) under

the discussion of his summorosus subgroup, was stated to be uncertain until I

examined specimens from this area and confirmed their identity.

Culex whitei is very distinctive in the male phallosome and larval stages.

The adults of whitei are very variable, particularly in the presence of absence

of light speckling on the anterior surface of the fore- and midfemur, overlap-

ping with specimens of si#zens and members of the vishnui complex. The speck-

led form of whitez can be distinguished from sitiens and the members of the

vishnui complex as indicated in the key whereas those which exhibit no speckling

on the femora are indistinguishable from members of the vishnui complex ex-

cept through association of the male genitalia and the larval stage. The pupa

of whitei is also distinct, particularly in the shape of the trumpet but is very

variable in the branching of the setae noted above. The pupae from Thailand

and Peninsular Malaysia exhibit fewer branches of seta 8-C, setae 6-III-VI and

1-III-VII than those from the Philippines, but show no difference from the latter

in all other stages. With additional material from each of these areas, it will

be possible to determine whether more than one form is involved.

Because of the distinctive male phallosome and larva, whitei is assigned

to its own complex. The affinity of whitei is intermediate between the vishnui

complex and Barraudi Subgroup; the adults are similar to the former but the

pupa and larvae resemble the latter.

BIONOMICS. The typical breeding sites of whitei are fresh water ground

pools in dried up stream beds or near the margins of streams and creeks in

the hills or mountain areas at an elevation ranging from 400 to 500 meters.

These habitats usually contain numerous decayed leaves and branches of trees

and are under partial or heavy shade of tropical forest. The immatures were

also collected from marshy ground in the open at 30 m in Peninsular Malaysia

and from a rice field in the Philippines. They were usually found in association

with specimens of C, (Lophoceraomyia) mammilifer (Leicester); C. (Culicio-

myia) pallidothorax Theobald and nigropunctatus Edwards, Uvanotaenia macfar-

lanei Edwards and Aedes (Finlaya) luzonensis Rozeboom in the collections from

shaded stream pools and with C. (Lutzia) halifaxii Theobald, C. (Culex) gelidus

and tritaeniorhynchus; C. (Eumelanomyia) malayi; C. (Culiciomyia) fragilis

Ludlow and nigropunctatus; C, (Lophoceraomyia) variatus Leicester and Anophe-

les barbirostris in the collection from open swampy ground. The adults have

138 Contribs “Amer, int: Ist. vol: 12) no. 25° 1976

been seldom collected in the field. Nothing is known about the feeding habits

of the females and their disease relationships.

BARRAUDI SUBGROUP

The Barvaudi Subgroup exhibits considerable overlap with the Vishnui

Subgroup in all stages and might be considered as a complex of the latter sub-

group. However, because of the distinctive adults, it is recognized as a sep-

arate subgroup. The Barraudi Subgroup is characterized in the adults of both

sexes by the presence of a small, semi-erect silvery white scale patch on the

anterior lower surface of postspiracular area (absent in all other members of

the Sitiens Group and Vishnui Subgroup) and by the presence of a distinct longi-

tudinal pale stripe, particularly on the anterior surface of femora and tibiae of

mid- and hindlegs; in the male genitalia by outer division of the phallosome

with a strong, prominent, laterally divergent sternal spine (heavily curved and

pointing basad in all members of the Vishnui Subgroup) and in the larva by seta

7-I double; comb scales small, numerous with more or less normal apical

fringe of evenly fine spicules and by the presence of 4. 5-6 pairs (total 9-12) of

siphonal tufts.

DISCUSSION. The Barraudi Subgroup is represented by 2 rare and closely

related species, barvaudi Edwards and edwardsi Barraud, both of which were

originally described from India. Their detailed affinities are not clear. The

larva and pupa resemble those of whitei more than any other form; the male

phallosome resembles that of members of the mimeticus complex in the develop-

ment of the sternal spine of the outer division and the adults are generally simi-

lar to tritaeniorhynchus in having the scales of mesonotum predominantly dark

brown and in having dark fine hairlike setae on the ventral surface of segment

3 of the male palpus.

31. CULEX (CULEX) BARRAUDI EDWARDS

(Figs. 63, 64)

Culex barraudi Edwards 1922a: 284 (o'*); Barraud 1923a: 939 (L*); Barraud

1924a: 997 (o°*, 2); Senior-White 1927: 71 (L*).

Culex (Culex) barraudi Edwards, Edwards 1932: 204 (taxonomy); Barraud 1934:

403 ('*, 2, L*); Peters and Dewar 1956: 46 (o°*, L*); Bram 1967a: 209

(o*; 25%).

FEMALE. Wing: 3.2mm. Forefemur: 1.6 mm. Proboscis: 1.75 mm.

Resembling other members of the Vishnui Subgroup in general external charac-

ters, differing as follows. Head, Erect scales of vertex largely brownish or

blackish except for a few pale ones in center; lateral patch of broad appressed

scales whitish, very distinct. Palpus tipped with pale scales on apex of seg-

ment 4; rest dark. Proboscis with median pale ring occupying about 0.2 of

total length. Cibarial Armature. Rather similar to siliens; cibarial bar with

a concave row of about 25 very short, coarse and abruptly pointed teeth, all

subequal in length and size. Thorax (Fig. 63). Mesonotal integument tan to

dark brown; scales on anterior 0.7 narrow, rather fine, predominantly golden

or dark brown in middle, pale yellowish on extreme anterior promontory and

on marginal areas; sometimes also with pale yellow scales forming spots or

streaks behind fossae, posterior acrostichal and dorsocentral areas at level of

Sirivanakarn: Subgenus Culex in the Oriental Region 139

anterior wing base; scales on posterior 0.3 laterad of prescutellar space

golden to dark brown, not contrasting with scales on anterior 0.7; scales on pre-

scutellar space and scutellar lobe entirely pale beige, whitish or yellowish.

Pleural integument same color as mesonotum; psp with characteristic minute

silvery white scale patch on lower anterior surface in addition to pale scale

patches on ppl, stb and mep. Legs. Anterior surface of forefemur completely

dark; that of midfemur with a narrow, rather poorly defined longitudinal pale

stripe along whole length and with or without light sprinkling of pale scales on

lower surface at middle; anterior surface of hindfemur with distinct longitudin-

al pale stripe from base to near apex, bordered above and below by dark

scaled stripes; fore- and midtibia entirely dark anteriorly; midtibia with poorly

defined pale stripe in middle; anterior surface of hindtibia with a longitudinal

pale stripe from basal 0.25 to apical 0.75 of the length. Wing. All scales

dark, narrow and dense; no pale scaled line or streak at base or in basal 0.5

of veins C. Abdomen. Terga II-VII with basal pale bands, narrower on pos-

terior segments.

MALE. As described for female, differing in largely pale mesonotal

scales, absence of scattered pale scales on anterior surface of forefemur,

absence of longitudinal pale stripe on anterior surface of midfemur and in

broader basal bands on abdominal terga II-VII. Head, Palpus exceeding pro-

boscis by full length of segment 5; segment 3 with row of short, dark, hairlike

setae on ventral surface, apical 0.25 with about 8 strong bristles; segments 4

and 9 weakly to moderately plumose, with narrow basal pale bands on dorsal

surface; apex of segment 5 without apical pale band. Proboscis with very poor-

ly developed ventral tuft of a few rather short hairs at base of median pale

ring.

MALE GENITALIA (Fig. 63). Essentially similar to other members of the

Vishnui Subgroup, differing particularly in the following features. Basimere.

Setae on inner tergal surface laterad of subapical lobe less numerous and

apparently shorter and weaker. Subapical lobe. Leaf g of distal division rela-

tively broad, heart-shaped, with acuminate apex. Phallosome. Apical tergal

crown of inner division with 4, 5 stout fingerlike processes, all apparently sub-

equal in length, straight, divergent laterad or dorsolaterad; outer division with

1 large and 1 small mesal spine; sternal spine poorly developed, rather short

and indistinct; lateral basal process slender. Proctiger. Apical crown rela-

tively large; basal sternal process of paraproct present, very strong, thick,

curved sternad.

PUPA. Abdomen: 2.5mm. Paddle: 0.78 mm. Trumpet: 0.70 mm;

index 8. In general as described and figured for whitei (Fig. 61). Trumpet.

Brown or yellowish brown, more or less uniform in width except for slightly

expanded pinna. Cephalothorax. Seta 8-C 4 branched; 9-C double or triple.

Abdomen: Seta 1-II 10 branched; 1-III 9 branched; 1-IV 5-7 branched; 1-V, VI

0, 6 branched; 1-VII 4 branched; 5-IV triple, 5-V, VI double, as long as or

slightly longer than segment following, 6-III-VI usually triple, rarely double.

Paddle, Entirely pale; midrib weak, pale or lightly pigmented.

LARVA (Fig. 64). Head: 0.78 mm. Siphon: 1.8-2.0 mm; index 8. Saddle:

0.36 mm; siphon/saddle ratio 5. Extremely similar to whitei in shape and

size of siphon and in chaetotaxy, differing particularly in the following. Abdo-

men. Setae 6-I, II triple; 1-III-VI about 0.5 of setae 6-IJI-VI, all subequal in

length; 1-III, IV double or triple (1-3); 1-V, VI 3-5 branched. Apical fringe of

comb scales more or less rounded, composed of 3,4 equally strong apical spi-

cules and several fine lateral spicules. Seta 1-X of saddle usually single

(1,2). Siphon. Bright yellow except for basal dark ring; siphonal tufts longer

140. Contrib. Amer. Ent. Inst., vol: 12, no. 2, 1976

and stronger, 4.5-5 pairs (total 9-10); 2.5-3 pairs inserted subventrally;

other 2 pairs inserted laterally, each usually triple (2-4); most proximal pair

longest, about 1.5 times as long as siphonal width at point of attachment, rest

gradually shorter; ventral valve of spiracular apparatus broader, longer and

very dark; median caudal filament poorly developed.

TYPE-DATA. Holotype “* (E 267) with slide of genitalia, Mahdopur,

Punjab: INDIA, April 1921, P. J. Barraud (BM).

DISTRIBUTION. Known only from India, China and Thailand. Material

examined. 21 specimens: 120, 72, 2 L; 2 individual larval rearings.

INDIA. Punjab: Mahdopur (type-locality); 60°, 12, 2 lp (all in type-series);

Western Himalayas: Kasauli; 5%, 49.

CHINA. Yunnan: Poashan Valley; 2 L (S. C. Billings, 1944).

THAILAND. Chinag Mai: Huey Mae Sanarn; Ban Bo Rae Hod; 1, 29.

Additional records from the literature. INDIA. Punjab, Pathankote;

Amritsar; Kalka; Western Himalayas: Murree; South India: Karikal, Tanjore

(Barraud 1924a, 1934); SRI LANKA (Senior-White 1927); NEPAL (Peters and

Dewar 1956); PAKISTAN (Aslamkhan 1971).

TAXONOMIC DISCUSSION. Culex barraudi is a very poorly known species.

This is because the original description of the holotype male by Edwards (1922a:

284-5) and the subsequent descriptions of both sexes and the larva by Barraud

(1923a: 939, 1924a: 997, 1934: 403) were inadequate, lacking certain significant

features which are diagnostic of this species. In this study, I re-examined the

type, additional specimens in the type-series and other previously identified by

Barraud. A comparison of the Indian material of barraudi with the few adults

described by Bram (1967a: 209-13) from Thailand indicates that specimens

from these 2 countries are conspecific. Except for some corrections and addi-

tions as indicated above, descriptions of the adults, male genitalia and larva

by Bram (loc. cit.) are generally applicable to barraudi,

Culex barvaudi is extremely similar to edwardsi Barraud in all stages and

with the latter evidently falls into a distinct complex or subgroup. The syn-

onymy of edwardsi with barraudi proposed by Bram (1967a) was premature

and does not appear to be justified, as discussed by Marks (1971: 195-6). In

comparing the type adults and others in the type series of both species, I found

that they differ from one another rather strikingly in the relative size, details

of the male palpus and proboscis and slightly in other external features and in

the male phallosome. This evidence suggests that barraudi and edwardsi are

in fact not conspecific, but most probably represent 2 distinct forms of a

complex. However, with additional material, especially adults with associated

immature stages from the type localities in India for a detailed analysis, it

should be possible to determine their precise status.

Culex barraudi adults can be readily separated from members of the Vish-

nut Subgroup by the presence of a minute, semi-erect, silvery white scale

patch on the lower anterior surface of the postspiracular area and by the pre-

sence of a narrow longitudinal pale stripe on the anterior surface of mid-

femur, mid- and hindtibiae; in the male by the fine, hairlike setae on ventral

surface of palpal segment 3 and by the poorly developed ventral tuft of hairs at

the base of median pale ring of the proboscis; in the male genitalia by the de-

tails of the inner and outer division of the phallosome. The pupa and larva of

barvaudi are extremely similar to whitei in general features and chaetotaxy,

but can be separated from the latter as indicated in the above diagnosis and in

the keys. For separating barvaudi from edwardsi, see the keys and the dis-

cussion under the latter species.

The additional new record of barraudi from China is based on 2 whole lar-

Sirivanakarn: Subgenus Culex in the Oriental Region 141

vae. In the absence of associated adults, this record is only provisional and

requires confirmation. All additional records from India by Barraud (1924a,

1934) and Peters and Dewar (1956) are presumably valid but the record from

Sri Lanka by Senior-White (1927) is doubtful.

BIONOMICS. Culex barraudi appears to be restricted to a high elevation in

mountainous areas in the northern parts of the Oriental regions. In Punjab,

India, it was reported to breed in ground pools, marsh, etc. (Barraud 1934:

403). In Chiang Mai, northern Thailand, it was collected once in sandy pools

without further notes on the environment. The adults from Kasauli and Murree

in the Western Himalayas were reported to come from an elevation of 1, 540-

2,140 meters (Barraud 1934: 403). Adult biology, feeding habits and disease

relationships are practically unknown.

32. CULEX (CULEX) EDWARDSI BARRAUD

(Fig. 63)

Culex edwardsi Barraud 1923b: 507 (2); Barraud 1924a: 995 (@).

Culex (Culex) edwardsi Barraud, Edwards 1932: 204 (taxonomy); Barraud

1934: 403 (“*, 2); Carter and Wijesundara 1948: 148 (L*); Peters and

Christian 1963: 35 (distribution); Marks 1971: 195 (taxonomy).

Culex (Culex) barraudi of Bram 1967a: 211, (in part).

FEMALE. Wing: 4.3mm. Forefemur: 2.0mm. Proboscis: 2.0mm. Ab-

domen: 2.9mm. Ingeneral, extremely similar to barvaudi inthe presence ofa

distinct, minute, silvery white scale patch on lower anterior surface of postspiracu-

lar area, differing in the presence of well defined longitudinal pale stripes on anteri-

or surface of femora and tibiae of mid- and hindlegs, larger size and inthe following

features. Thorax. Scales onanterior 0.7 of mesonotum coarser and predominantly

darkbrown. Legs. Anterior surface of forefemur with or without distinct longitu-

dinal pale stripe or chain of pale spots extending from base to apical 0. 75 on lower

margin; foretibia with or without scattered pale scales which may forma complete

stripe ora chain of pale spots anteriorly; anterior surface of femora and tibiae of

mid- and hindlegs with well defined longitudinal pale stripes along the whole length;

pale stripe on hindfemur broad, covering entire surface in basal 0. 5and tapering in

apical 0.5; tarsomeres of all legs with broader and more conspicuous basal and

apical pale bands. Wing, Posterior dorsal surface of vein C with pale scale

line or streak at least at base near humeral vein or sometimes extending to

0.5 or more of the whole length; dorsal scales of vein Sc usually largely pale,

sometimes dark. Abdomen. Basal pale bands of terga II-VI relatively broader;

terga V-VII with distinct narrow apical pale bands, largely restricted to mid-

dle.

MALE. Essentially similar to female in general and in the striping of the

legs; differing from barraudi as described for female and in the following.

Head. Palpus longer and thicker, exceeding proboscis by 1.5 of segment 5;

segment 3 with prominent ventrolateral tufts of about 30 bristles from basal

0.4 or 0.5 to apex; segments 4 and 5 very strongly plumose; apical 0.25 of

segment 9 pale scaled to tip. Proboscis with a strong and very conspicuous

ventral tuft of at least 10 long hairs at base of median pale ring, these hairs

about 5 times as long as labial width at point of insertion.

MALE GENITALIA (Fig. 63). Exceedingly similar to barvaudi, differing

slightly in the following details. Subapical lobe. Setae d-f of distal division

apparently stronger and more flattened; leaf g relatively broader. Phallosome.

142 Contrib.: Amery: Ent, Insts, vold2y no. 2,..1916

Sternal spine of outer division very well developed, relatively large, strongly

divergent laterad; lateral basal process larger, with variable number of dis-

tinct denticles near base of mesal spine.

PUPA. Abdomen: 4.0mm. Paddle: 1.0 mm. Trumpet: 0.75-0.90 mm;

index 8-9. Extremely similar to barraudi and whitei; and as figured for whitei

(Fig. 61), differing in larger size, longer and more uniformly cylindrical

trumpet and in the following. Cephalothorax. Seta 8-C usually double (2, 3).

Abdomen. Seta 1-II forked into 4 branches; 6-III, IV double; 6-V, VI 3, 4 bran-

ched; 5- VII very strong, longer than seta 1-VIII, double; 4-VIII single.

LARVA. Head: 0.90 mm. Siphon: 2.0 mm; index 7. Saddle: 0.36 mm;

siphon/saddle ratio 6. Extremely similar to barraudi, differing from it par-

ticularly in the following features. Abdomen. Setae 6-III,1V usually triple

(2,3); 1-III-VI usually double or triple (2-4). Comb scales more or less

pointed apically; median apical spicule apparently stronger than lateral spicules.

Posterior margin of saddle with stronger spicules; seta 1-X strong and single.

Siphon. Pecten teeth slightly shorter and larger; most distal teeth with 4,5

lateral denticles and a weak apical denticle; siphonal tufts 5. 5-6 pairs (total

11-12); 3.5-4 pairs inserted subventrally, 1.5-2 pairs inserted laterally; medi-

an caudal filament of spiracular apparatus well developed.

TYPE-DATA. Holotype °*, Shillong, Khasi Hills, Assam, INDIA,

June 1922, P. J. Barraud (BM).

DISTRIBUTION. Definitely known only from India, Papua-New Guinea and

Queensland, Australia. Material examined. 12 specimens: 7", 59; 4 indivi-

dual rearings (1 pupal, 3 larval).

INDIA. Assam: Shillong, Khasi Hills (type-locality); 12 (holotype), 1°;

Central India: Rewa State, Amol Nullah; 2%, 12 (Senior-White); South India:

Coorg, Mercara and Virajpet; 1“, 22 (Baily); Madras: Nilgiri Hills, 1°, 12,

i p;..t.lp (8B. Ne Mohan, 1974);

PAPUA-NEW GUINEA. S, Highlands: Minj; 2%, 12, 2 lp (Peters and

Christian).

Additional records from the literature: SRI LANKA. Nuwara, Eliya

(Carter and Wijesundara 1948). AUSTRALIA. Queensland: Fraser Is. (Marks

1968).

TAXONOMIC DISCUSSION. Culex edwardsi is very closely related to bar-

raudi on the basis of similarity in all stages and as discussed under the latter;

the 2 species evidently fall into a distinct complex or subgroup. Although both

species have long been known only from the types and from the limited number

of identified specimens originally described and figured by Barraud (1923b;

1924a; 1934), present comparison has shown that they are distinct and probably

not conspecific as interpreted by Bram (1967a: 209-213). In this study,

edwardsi is resurrected from synonymy with barraudi.

This species is still very poorly known from every locality within its re-

ported range. The above descriptions of the adults have been based on the

study of the type female and specimens listed in the distribution section. In

comparing the Indian specimens with those from New Guinea, I found that they

are essentially similar in all stages, indicating that they are conspecific.

Culex edwardsi can be distinguished from barvaudi in the adults by: (1)

larger size; (2) mesonotal scales coarser and more predominantly brownish

on anterior 0.7; (3) longitudinal pale stripes of the femora and tibiae of mid-

and hindlegs well developed in both sexes and apparently more conspicuous;

(4) posterior surface of vein C with distinct pale scale line extending from base

to about 0.5 or more of the total length; (5) abdominal terga V-VII of the fe-

male with narrow median apical pale bands in addition to basal bands; in the

Sirivanakarn: Subgenus Culex in the Oriental Region 143

male by (1) longer palpus; (2) ventral lateral tuft of palpal segment 3 larger,

with more numerous bristles, extending from basal 0.4-0.5 to the apex; (3)

palpal segments 4 and 5 densely long plumose, with more numerous bristles on

lateral and mesal surfaces and (4) proboscis with a strong ventral tuft of long

hairs at the base of median pale ring. In the male genitalia, edwardsi differs

from barraudi particularly in having a stronger, longer and more prominent

sternal spine in the outer division of the phallosome. The pupa and larva of

edwardsi are very similar to barraudi and both apparently exhibit much overlap

with whitei, The edwardsi pupa differs from those of barraudi and whitei in

having abdominal seta 5-VII stronger and longer. The larva can be separated

from whitei by weaker and shorter setae 1-IV and V and from barraudi by the

presence of 5.5-6 pairs of siphonal tufts (4. 5-5 pairs in barraudi) and by a few

other features as indicated in the above description.

BIONOMICS. Very little is known about the larval breeding sites and the

adult biology of edwardsi, It appears to be uncommon and restricted to a very

high elevation as barraudi. In the Nilgiri Hills, Madras, the immatures were

collected twice, once ina stream pool and once ina spring pool. In Sri Lanka,

the larvae were collected from a grassy pond and a swamp at the elevation of

about 1, 232-1, 846 m (Carter and Wijesundara 1948: 148). Peters and Christian

(1963) described it as a fresh water breeder in New Guinea, but no information

about the breeding site was given. The adults, including the type of edwardsi

from India reported by Barraud (1924a, 1934) all apparently came from general

field catches and those recently collected from the Nilgiri Hills all came from

field rearing. At Minj, Papua-New Guinea, Peters and Christian (loc. cit. )

collected adults in a light trap and in Australia, Marks (1968, 1971) noted that

it has been taken in chicken-baited, light and truck traps. It is not known to

bite man.

MIME TIC US SUBGROUP

The Mimeticus Subgroup generally conforms to the Sitiens and Vishnui Sub-

groups in all stages and is chiefly characterized by the following combination of

characters: in the adults by (1) presence of a striking pattern of pale spots on

the wing; in the males by ventral surface of palpal segment 3 always with short,

scalelike setae; in the pupa by seta 1-II usually between 1-9 branched, not ex-

ceeding 10 branched; and in the larva by (1) seta 4-P, single (except for C.

mimuloides which has this seta double); (2) seta 7-I always single; (3) comb

scales numerous, all small, subequal in a broad oval patch and (4) median

caudal filament of spiracular apparatus absent or poorly developed.

DISCUSSION. The Mimeticus Subgroup as recognized here corresponds to

the Mimeticus series or group as defined by Edwards (1932: 201-5; in Barraud

1934: 450-3). Its members can be easily distinguished from the rest of the

Szitiens Group in the adults by the spotted wing and in the majority of species by

the single seta 4-P of the larva. This subgroup apparently exhibits the strong-

est affinity with the Vishnui Subgroup on the basis of the male phallosome and

the morphology of all stages, indicating that it was probably derived from the

latter and that the subgroup as a whole presumably originated in the Oriental

region.

The Mimeticus Subgroup is diverse, highly complex and taxonomically very

difficult. Some Oriental members of this subgroup are still poorly known be-

cause they are not commonly encountered and appear to be restricted to certain

localities at a very high elevation. The identity and the status of some mimeti-

144 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

cus forms are very difficult to determine because of the considerable overlap

and similarity in one or more stages. When additional specimens from progeny

and individual rearings become available from several localities for a thorough

analysis, it may be necessary to revalidate the status of some of these forms.

Based primarily on the differences in the male genitalia, pattern and extent of

pale spots of the wing and in certain cases also on the larval and pupal stages,

Iam recognizing 10 species of the Mimeticus Subgroup in the Oriental and Pale-

arctic regions. The majority of these are exclusively Oriental except for

mimeticus whose range extends into the Middle East, the Mediterranean and the

northeastern Palearctic (China, Japan and Korea), mimulus which is also

known from New Guinea and northern Australia and orientalis which is known

only from the northeastern Palearctic, particularly Japan, Korea and possibly

China and USSR.

The 10 members of the Mimeticus Subgroup recognized here can be assem-

bled into 2 complexes: mimeticus and mimulus on the basis of the differences

in the pattern and the extent of the wing spots, certain features of the male

genitalia, pupal and larval chaetotaxy. Separation of the adults of the members

in each complex are very tenuous and require examination of the male genitalia

and associated immature stages.

The mimeticus complex is composed of mimeticus Noe 1899, jasyi Baisas

1938, jacksoni Edwards 1934, tsengi Lien 1968, mimuloides Barraud 1924 and

diengensis Brug 1931. This complex also includes solitarius Bonne-Wepster

1938 from New Guinea. The mimeticus complex is characterized in the adults

by (1) wing with only 3 costal pale spots in apical 0.5 and (2) first costal spot

at about the middle of vein C involving veins C and Sc or sometimes restricted

to Sc only; in the male genitalia by (1) distimere without distinct subapical

crest of minute spicules; (2) leaf g of the subapical lobe relatively narrow; (3)

outer division of the phallosome with a strong, prominent sternal spine which

is divergent laterad and (4) basal sternal process of the proctiger absent or

present; if present, usually very slender, thin and pale; in the pupa by (1) seta

1-II usually single or double, rarely more branched; (2) setae 1-III, IV usually

2-6 branched, rarely more branched; (3) setae 6-III, IV usually single or double

rarely 3,4 branched and (4) seta 4-VIII usually single, rarely double and in the

larva by (1) comb scales relatively coarse and usually with apical fringes ter-

minating into a strong median spine; (2) siphon usually 1.4-1.6 mm in length,

rarely exceeding 2.0 mm; (3) 4 proximal pairs of siphonal tufts usually strong-

est, closely spaced, forming a prominent row, 2 distal pairs weaker and short-

er and (4) apical half of siphon with or without prominent spines on ventral sur-

face.

The mimulus complex includes mimulus Edwards 1915, murrelli Lien 1968,

propinquus Colless 1955 and orientalis Edwards 1921. It is characterized in

the adults by (1) wing with or without basal costal pale spot in addition to 3 other

costal spots in apical 0.5; (2) first costal spot at about the middle of vein C in-

volving veins C, Sc, Ry and R,, sometimes also extending to veins M and Cu;

in the male eenitalia by (1) distimere with distinct subapical crest of several

minute spicules; (2) leaf g of the subapical lobe broad; (3) outer division of the

phallosome with a poorly or well developed sternal spine which is usually

strongly curved with apex projecting basad; (4) basal sternal process of the

proctiger present, very strong, stout and dark; in the pupa by (1) seta 1-II usu-

ally more than 3 branched (2-9); (2) setae 1-III, IV 5-10 branched; (3) setae 6-III,

IV usually triple, rarely double; (4) seta 4-VIII double and in the larva by (1)

comb scales relatively fine and with apical fringe of evenly fine spicules or

terminating into a weak medium apical spine; (2) siphon 1.4-2.5 mm in length;

Sirivanakarn: Subgenus Culex in the Oriental Region 145

(3) all siphonal tufts relatively weak, short, subequal, widely spaced, not form-

ing a prominent row; (4) apical half of siphon without prominent spines on ven-

tral surface.

33. CULEX (CULEX) MIMETICUS NOE

(Figs. 16, 65, 67, 68)

Culex mimeticus Noe 1899: 240 (2); Edwards 1921: 337 (%, 9, L); Edwards

1922a: 284 (distribution); Barraud 1923a: 941 (L*); Barraud 1924a: 991

(O*,: 9. ih payt)s Borel: 19305348 {os .9o T*): Ho Oa ei sao £410),

Culex pseudomimeticus Sergent 1909 (1910): 445 (adult); Edwards 1932: 205

(synonymy).

Culex (Culex) mimeticus Noe, Edwards 1932: 205 (taxonomy, in part);

Barraud 1934: 409 (“*, 2, L*, in part); Edwards, in Barraud 1934: 451

(“, 2, key); Feng 1938: 302 (distribution, biology); Bohart and Ingram

1946: 80 (**, 2, L*); LaCasse and Yamaguti 1950: 241 ('*, @*, P*, L*).

FEMALE (Fig. 16, 65). Wing: 4.5mm. Forefemur: 2.1mm. Probos-

cis: 2.2mm. Abdomen: 3.0mm. Medium or large sized species. Head,

Narrow decumbent scales on vertex entirely pale or partially pale yellowish

along dorsal midline, darker on lateral areas; erect scales numerous, very

coarse, golden or bronzy in center, dark brown on lateral and posterolateral

areas; lateral patch of broad whitish scales, very distinct from above. Palpus

tipped with pale scales on apex of segment 4, rest dark. Proboscis witha

broad median pale ring; portions distad and proximad of median ring entirely

dark scaled. Czibarial Armature. Essentially similar to most members of

the Sztzens Group; cibarial bar with a concave row of about 30 short and coarse

teeth all of which are subequal and abruptly pointed apically. Thovax. Mesono-

tal integument dark brown; mesonotal scales on anterior 0.7 coarse and pre-

dominantly yellowish to golden with mottling of brown scales forming spots and

streaks on fossae, acrostichal and dorsocentral areas; scales on prescutellar

space and scutellar lobes largely pale beige or pale yellowish. Apnand ppn

with golden and brownish scales as on mesonotum. Pleuron same color as

mesonotum; pale scale patches on upper corner of stp and upper mep broad

and more or less contiguous; several pale scales present among upper mep

bristles. Legs. Anterior surface of fore- and midfemora usually entirely

dark, sometimes lightly speckled with pale scales, subapical yellowish spots

(knee spots) present; anterior surface of hindfemur with longitudinal pale stripe

extending from base to slightly beyond 0.5 of whole length, apical 0.5 dark;

dark area dorsad and distad of stripe with or without light sprinkling of pale

scales; pale stripe usually lightly freckled with some dark scales; subapical

yellowish spot absent; fore- and midtibiae without pale stripe; hindtibia usually

with distinct longitudinal stripe on anterior surface; tarsomeres 1-4 of all legs

with broad basal and narrow apical pale bands. Wing. Characteristic pattern

of pale spots as figured; extent of pale spots variable; base of vein C without

pale scale line or streak on posterior surface, apical 0.5 or slightly more with

3 costal spots (first costal, 2nd costal and 3rd costal); first costal spot at

about middle of vein C involves veins C and Sc; 2nd costal spot at about 0.75 of

vein C involves veins C, apex of Sc and middle of Ry; 3rd costal spot near tip

of wing usually involves apical portions of veins C, Ry, and Rg; furcation of

R913 proximad of or at same level of furcation of M, both with pale spots large-

ly involves basal portions of veins Ro, Re, My, and M3, 4; vein R4,5 pale

146 Contrib... Amer.’ Ent. Inst... volo 12) 10.72; 19.76

from basal 0.4 to apical 0.75; vein Cu, pale from basal 0. 25 to apical 0. 75;

Cug with short apical pale streak; fringe scales on apex of Cua pale; vein 1A

pale from basal 0.29 to 0.90; plume scales narrow and dense on veins Ro, Rg,

Rgi5, My,9 and Mgi4. Abdomen, Terga II-VII with basal pale bands of vari-

able width and large basolateral pale spots; terga VI-VII with narrow median

apical pale bands; sterna II- VII with broad basal pale bands and narrow apical

dark bands. Genitalia. Essentially similar to those of the Sitsens Group; ter-

cum IX with a lateral row of 5, 6 strong setae; postgenital plate rounded or

slightly emarginate on caudal margin; vaginal sclerite U- or V-shaped; insula

with a tuft of about 10 moderately strong setae.

MALE. In general similar to female except for less striking pattern of

wing spots and less dense scales on all wing veins. Head. Palpus exceeding

proboscis by at least full length of segment 5; segment 3 with a broad median

pale band on dorsal surface, ventral surface with a dense row of short, scale-

like setae along whole length, apical 0.25 with a strong ventral lateral tuft of

10-16 bristles; segments 4 and 5 strongly plumose, each with a broad basal pale

band on dorsal surface; apex of segment 5 tipped with pale scales. Proboscis

with pale scale line distad and proximad of median pale ring on dorsal surface;

ventral tuft of hairs at base of median pale ring very weak, indistinct, com-

posed of a few hairs which are 2,3 times as long as labial width at point of

insertion.

MALE GENITALIA (Fig. 67). Essentially similar to members of the

Sitiens and Vishnui Subgroups, differing particularly in the following features.

Subapical lobe. Setae d-f of distal division 4, all flattened, most distal seta

longest and hooked apically; the rest shorter, subequal; leaf g relatively nar-

row, apically rounded or abruptly pointed. Distimere. Dorsal subapical

crest of minute spicules completely absent. Phallosome. Apical tergal

crown of inner division of lateral plate with 2-4 fingerlike processes which are

variable in length and thickness, all more or less straight and with apices

pointing laterad or dorsolaterad; outer division with 1 strong mesal spine and

with or without 1 other weaker spine on tergal surface, sternal surface with a

prominent sternal spine which is gently or strongly divergent laterad; lateral

basal process broadly rounded or slightly elongate, forming a distinct knob.

Proctiger. Apical crown large, dark, composed of numerous coarse spicules;

basal sternal process very slender, thin, pale, about 0.05 mm in length, gent-

ly curved sternad; subbasal process very short; cercal sclerite broadly sclero-

tized; cercal setae usually 2(2-4).

PUPA (Fig. 67). Abdomen: 2.7mm. Paddle: 0.8 mm. Trumpet: 0.7

mm; index 7-9. Cephalothorax and abdomen yellowish or cream-colored with-

out distinct pattern of darkened areas. Complete chaetotaxy as figured, dis-

tinctive in the following. Tvumpet, Yellowish brown, more or less uniform

in width except for slightly expanded pinna; apical margin truncate or broadly

rounded. Cephalothorax. Seta 8-C double; 9-C usually double. Metanotum.,

Seta 10-C usually 3,4 branched (3-5). Abdomen. Seta 1-II single or forked

into 2 branches; 1-III 2-4 branched; 1-IV 3, 4 branched; 1-V-VII usually double

(1-3); 5-IV usually double or triple (2-4), as long as or slightly longer than

segment following; 5-V, VI double, 1.5 times as long as segment following;

6-III-VI single or double; 2-VII usually mesad of 1-VII; 4, 5-VII usually single;

o- VII nearly as long as the length of segment VIII; 9-VII usually double or

triple (2-4); 4-VIII usually single (1,2); 9-VIII 4-8 branched, as long as 9-VII.

Paddle, Very broad and entirely pale; midrib lightly pigmented; setae 1, 2-P

present; 2-P stronger than 1-P.

LARVA (Fig. 68). Head: 0.78 mm. Siphon: 1.5-1.7 mm; index 6-7.

Sirivanakarn: Subgenus Culex in the Oriental Region 147

Saddle: 0.4 mm; siphon/saddle ratio 4. Generally similar to members of the

Vishnui Subgroup, differing particularly in the following combination of char-

acters. Head, Seta 5-C usually 3,4 branched (3-5); 6-C usually double (2, 3);

7-C 5-7 branched; setae 10,11 and 13-C usually double (2,3); 12-C triple; 14-C

very distinct and strong, somewhat spiniform. Antenna as long as head.

Thorax. Spiculation absent; seta 4-P single and as strong as 1-3-P; setae 8,

9-M usually 5,6 branched; 7-T 7 branched; 13-T usually double (2-4). Abdomen.

Setae 6-I, II triple; 7-I single; 6-III-VI double or triple; all setae 5-I-VI single

and very distinct; 1-III- VI usually double, all subequal, about 0.5 of seta 6-TIII-

VI. Comb scales 25-30, all small, subequal in size and with apical fringe ter-

minating in a strong, distinct median spine. Seta 1-X of saddle usually double

(2,3); 2-X double; anal gills slender, 2,3 times as long as saddle length.

Siphon. Moderately long, straight and gradually tapered distally; pigmenta-

tion yellowish white except for basal dark ring; pecten teeth 13-18, all gently

curved dorsad; siphonal tufts 6 pairs (or total 12), inserted beyond pecten; 4

proximal pairs strong, subequal, 4,5 branched, in a prominent row on ventral

surface, their length 2,3 times of siphonal width at point of attachment; 2 distal

pairs weaker and shorter, double or triple, the proximal inserted laterally,

the other subventrally, their length as long as siphonal width at point of attach-

ment; seta 2-S dark, strong and moderately long; median caudal filament of

spiracular apparatus absent.

TYPE-DATA. (1) Culex mimeticus Noe, Type @ (non-existent), Grassano,

Basilicata, ITALY; deposited in Rome University Museum (Barraud 1924a,

1934); (2) C. pseudomimeticus Sergent; Type (non-existent), Chiffa, ALGERIA.

DISTRIBUTION. Presumably widespread in the Oriental region with the

range extending west into the Middle East and the Mediterranean; north and

northeast into the Palearctic (including Tibet, Kashmir, China, Japan and

* Korea). Material examined: 178 specimens: 51°, 659, 62 L; 16 individual

rearings (1 pupal, 15 larval).

ITALY. Matera Province: Grassano, Basilicata (type locality); Savignano

(Avelliuo); 4°, 62, 10 1p (M. Coluzzi, 1970).

GREECE. Macedonia: N. Salonica; 1.

TUNISIA. Mateur; 150, 122, 1 L(G. T. Riegel, 1943).

ALGERIA. Chanzy; 59.

IRAQ. Mirgasur, Shanider; 3 L.

IRAN. Teheran; Arimarabad, E. Chalus, Kheyrud River; Khorramabad;

Kambu; 2%, °49.1-L:;

NEPAL. Khatmandu; 1 L (J. Maldonado, 1958).

TIBET. Yatung; 2°.

INDIA. Western Himalayas: Kasauli; Durhampur; ASsam: Tezpur; Ledo;

Dibrugarh; Shillong; Madras: Nilgiri Hills, Manjoor; 7“, 62, 3 L; Kashmir:

Navranes 1o')) 19,

BURMA. Salween River Valley; 2 L (S. C. Billings).

MALAYSIA. Peninsular Malaysia: Pahang-Cameron Highlands; Mt. Brin-

ehane? To 572) (Bul ib Ip)

VIETNAM. Dalat, Du Sinh Hamlet; 29, 1 p.

HONG KONG. 4c, 19, 6 L (skins).

CHINA. Peking; Hanchow; Foochow; Nanking; Yunnan; Chentu; 1c, 39,

re i a

JAPAN. Kyoto; Nagasaki; Otsu (Honshu); 40°, 99, 3 L. Ryukyus: Okin-

awa; Taera; Nakasoni; 1“, 42, 7L.

KOREA. Yanggu; Koje-do; Nouak; 20, 52, 1L.

Additional records from the literature: SRI LANKA; INDOCHINA (Barraud

148 Contrib.: Amer. Pnt. Inst. vol. 12;°no.' 2; 1976

1934: 411); PAKISTAN (Aslamkhan 1971); TAIWAN (FORMOSA) (Edwards, in

Barraud 1934: 451; Lien 1968); CHINA: Anhwei, Chekiang, Hopei, Kiangsi,

Kiangsu, Kwangtung, Shangtung, Tibet (Feng 1938); FRANCE: Corsica; ITALY:

Sardinia; SPAIN (Aitken 1954: 486; Nicoli and Nicoli 1964: 38); USSR (Monchad-

skii 1951). ,

TAXONOMIC DISCUSSION. Culex mimeticus was originally described from

Italy by Noe (1899) and the name has since been widely used by European work-

ers for all spotted wing Culex from several parts of the Mediterranean, Middle

East and the Oriental region. The type of mimeticus is apparently lost. How-

ever, through the courtesy of Dr. M. Coluzzi in Italy, I have obtained a good

number of topotypic specimens with associated immature stages for detailed

study and comparison with the Oriental forms. In addition, I have also ex-

amined specimens from several other areas in the collections of the USNM

and the British Museum (Natural History).

This is the most widespread species of the Mimeticus Subgroup. It is still

poorly known from most parts of the Oriental region. The earlier Oriental

records of this species are not always reliable because of the considerable over-

lap in the pattern of wing spots with other members of the complex. The only

feature which is diagnostic of mimeticus and has been used here in separating

it from other members of this complex is the presence of a long, slender basal

sternal process of the proctiger of the male genitalia. This particular feature

shows an overlap only with mimuloides which differs from mimeticus in sev-

eral characters of the larva and other stages. Within Southeast Asia, mimeticus

is definitely known only from Pahang, Peninsular Malaysia, Vietnam, Hong

Kong and Okinawa. The records in the literature from Sri Lanka (Barraud 1934)

and Taiwan (Edwards 1934, Lien 1968) are uncertain. They require further

confirmation since these were based entirely on the pattern of wing spots. The

male genitalia as figured by Lien (1968: 251) from Taiwan do not appear to

agree with the typical form of this species.

Except for the diagnostic male genitalia, mimeticus exhibits much similari-

ty to and overlaps with fasyz Baisas, jacksoni Edwards and tsengi Lien in the

pattern of wing spots and in most features of the pupal and larval stages. For

the separation of these taxa from mimeticus, see the keys and the discussions

under each individual taxon. EK appears probable that all of these forms are

only subspecifically distinct. Additional material from individual and progeny

rearings from several localities is needed to determine the definite status of

each of these forms.

BIONOMICS. Culex mimeticus is a common Mediterranean species of

Culex (Culex) (Aitken 1954, Nicoli and Nicoli 1964) but is not frequently en-

countered in the Oriental region. It appears to be restricted to high elevations

in the mountains or hill areas in India, Burma (Barraud 1934), Southeast Asia

and other adjacent areas. In Tibet, it was reported from an elevation of 3, 050

m (Feng 1938). The usual breeding sites of mimeticus include general ground

pools in dried up stream beds and in the vicinity of mountain streams or brooks,

running streams and rock pools. In Japan, it has been collected from drainage

ditches and rice fields (LaCasse and Yamaguti 1950, Hsiao and Bohart 1946).

The single collection from Pahang, Peninsular Malaysia came from a rock pool

at an elevation of 1,540 m. The water of the breeding sites is fresh and usu-

ally contains green algae. In Japan, Hsiao and Bohart (1946) reported that

-adults occasionally entered houses. The adult biology and medical importance

of mimeticus from all areas are unknown.

Sirivanakarn: Subgenus Culex in the Oriental Region 149

34, CULEX (CULEX) FASYI BAISAS

| (Fig. 69)

Culex (Culex) fasyi Baisas 1938: 215 (o"*, 9); Delfinado 1966: 142 (ey Bs

FEMALE. Wing: 4.0-5.0 mm (average 4.5mm). Forefemur: 2.2 mm.

Proboscis: 2.3mm. Abdomen: 3.2 mm. As described by Baisas (1938: 215-

6) and Delfinado (1966: 142-3); extremely similar to mimeticus in the pattern of

wing spots and in general facies, differing slightly as follows. Thovax. Scales

on anterior 0.7 of mesonotum more extensively brownish. Wing. Furcation of

vein M usually dark; basal portion of vein M proximad of crossvein M-Cu usually

with a short pale scale line on ventral surface (absent in mimeticus). Abdomen.

Terga III-VI usually with a row of pale scales forming narrow apical bands and

with a few light scales forming a pair of minute spots adjacent to apical band;

basal pale bands on all terga progressively narrower towards posterior seg-

ments.

MALE. In general similar to female with sexual characters as described

for mimeticus, differing from the latter in having 20 or more ventrolateral

bristles in apical 0.5 of palpal segment 3.

MALE GENITALIA (Fig. 69). Extremely similar to mimeticus in details

of basimere, subapical lobe, distimere and phallosome, differing from it par-

ticularly in having a shorter basal sternal process of the proctiger which is

about 0.01-0.03 mm in length and in the following. Subapical lobe. Leaf g of

distal division apparently shorter. Phallosome. Fingerlike processes of

inner division stronger, slightly swollen in middle, somewhat lanceolate in

shape. Pyroctiger. Paraproct usually with some minute spicules distad of sub-

basal process; the latter as long as or shorter than basal sternal process.

PUPA. As figured and as described for mimeticus, differing particularly

in having seta 9-C always single and seta 1-III 5-8 branched.

LARVA. As figured and as described for mimeticus to which it is extreme-

ly similar in features of the siphon and in chaetotaxy, differing particularly in

having seta 5-C usually 5 branched (4-6) and 6-C usually triple (3, 4).

TYPE-DATA. Holotype “ (No. R50-a) and 7 cotypes (4c, 32), Highlands

of Baguio, Luzon: PHILIPPINES, May 7, 1934, F. E. Baisas (PBH). . The

holotype and all cotypes are no longer in existence.

DISTRIBUTION. Known only from Luzon in the Philippines. Material

examined. 86 specimens: 370, 379, 12 L; 3 individual larval rearings.

PHILIPPINES. Luzon, Bonquet Otok; Bontok, Chico River; Calawan;

Mountain: Province, Baguio; 37¢; 379): 12: by 3tp.

TAXONOMIC DISCUSSION. The holotype and cotype adults of fasyi origin-

ally designated by Baisas (1938: 215-6) were destroyed by termites after

World War II (F. E. Baisas personal communication). I have examined sever-

al topotypic adults and whole larvae and several others with associated imma-

ture stages in the collection of the USNM. The adult of fasyi apparently agrees

well with the original description of Baisas (1938) and the subsequent descrip-

tion by Delfinado (1966). The pupa and larva of fasyi have not been known before

and are described here.

Culex fasyi is probably the only representative of the mimeticus complex

in the Philippines where it appears to be restricted to Luzon. All stages of

fasyi are exceedingly similar to those of mimeticus but can be distinguished

from the latter with certainty in the male genitalia by the shorter basal sternal

process of the proctiger. There is also much similarity in all stages, including

150 Contrib.: Amer, nt, Inst.:,. volj712, no. 25° 1976

the male genitalia, between the Philippine fasyi and solitarius Bonne-Wepster

(1938) from New Guinea (Bonne-Wepster 1954: 129-31), suggesting that these

forms may be conspecific. However, as I have examined only a few reared

specimens of the latter and since the distribution of both forms is still very

poorly known, it is premature to synonymize them at this time.

BIONOMICS. Culex fasyi is an uncommon Philippine form of Culex (Culex)

and like mimeticus, appears to be restricted to a high elevation. The larvae

have been collected in stream pools containing a dense algal mat in association

with specimens of Anopheles sp. (Baisas 1938). Nothing is known about the

adult biology and the medical importance of jfasyi,

35. CULEX (CULEX) JACKSONI EDWARDS

(Pigs. 65,70. 71)

Culex (Culex) jacksoni Edwards (in Barraud) 1934: 452 (“, @, L).

Culex (Culex) fuscifurcatus Edwards, in Barraud 1934: 452 (, 2); Carter and

Wijesundara 1948: 149 (L*); Harrison et al. 1974: 143 (distribution).

NEW SYNONYMY. ,

Culex (Culex) kangi Lien 1968: 235 (°*, 9*, P*, L*). NEW SYNONYMY.

FEMALE (Fig. 65). Wing: 4.8mm. Forefemur: 2.2 mm. Proboscis:

2.2mm. Abdomen: 3.4mm. Extremely similar to mimeticus in the pattern

of wing spots, differing slightly in the following features. Thorax. Mesonotal

scales on anterior 0.7 entirely dark brownish in middle of disc; pale scales

restricted to anterior promontory and other marginal areas cephalad of anteri-

or wing base. Legs. Anterior surface of forefemur usually speckled with

several pale scales in apical 0.5; midtibia usually with distinct longitudinal

pale stripe extending from base to apex on anterior surface, continued as nar-

row stripe on midtarsomere 1. Wing. Pattern of pale spots as figured; 3rd

costal spot near apex of wing sometimes involves only apical portion of vein

R,; furcation of vein M usually pale, sometimes dark.

MALE. As described for female, differing from mimeticus particularly

in the following. Head. Proboscis with a strong ventral tuft of several long

hairs at base of median pale ring; these hairs about 5 times as long as labial

width at point of attachment; basal and distal portions of labium without pale

scales forming streak proximad or distad of median pale ring on dorsal sur-

face.

MALE GENITALIA (Fig. 70). Exceedingly similar to mimeticus, fasyi and

tsengi in the details of basimere, subapical lobe, distimere and in the type of

phallosome, differing from these species in the complete absence of basal

sternal process of the proctiger and in having apical tergal crown of inner

division usually with 4, 5 fingerlike processes.

PUPA (Fig. 70). Abdomen: 3.5mm. Paddle: 1.02 mm. Trumpet:

0.85-0.90 mm; index about 7. Apparently very similar to mzmeticus, differ-

ing in the following combination of characters. Cephalothorax. Setae 8,9-C

usually double, sometimes triple. Abdomen. Seta 1-III usually 5 branched

(4-6); 1-IV 4-6 branched; 1-V 2-4 branched; all setae 6-III-VI usually single,

rarely double.

LARVA (Fig. 71). Head: 0.85 mm. Siphon: 1.4-1.6 mm; index 5. Sad-

dle: 0.44 mm; siphon/saddle ratio 3.5. Distinguished from mimeticus and

jfasyi by the presence of 2-11 prominent ventrolateral spines in apical 0.5 of

Siphon and by the following features. Head, Seta 5-C usually 5 branched (4-7);

Sirivanakarn: Subgenus Culex in the Oriental Region Lo4

6-C usually triple, sometimes 4 branched. Antenna relatively short, its length

varying from 0.5 to 0.75 of head length. Thorax. Seta 7-P usually triple,

sometimes 4 branched; 8-P usually double, sometimes triple. Abdomen. Seta

6-I usually 4,5 branched, rarely triple; 6-II 4,5 branched; 6-III, IV usually 3, 4

branched, rarely double; 6-V, VI double or triple; 1-III-VI usually double (1-3);

5-I-VI bifid or trifid, rarely single. Comb scales 36-38, all apparently finer

and with relatively weaker apical spines. Anal gills slender, usually blue in

live specimens, their length slightly exceeding saddle. Siphon. Rather thick,

moderately long and with variable number of prominent spines on ventrolateral

surface within 2,3 distal pairs of siphonal tufts; 11-19 pecten teeth in basal 0.5

of siphon, sometimes extending to 0.75; siphonal tufts variously developed; 5, 6

pairs (total 10-12) in number, 3,4 proximal pairs inserted ventrally, but not

forming a prominent row, 4,5 branched, their length varying from slightly

shorter to slightly longer than siphonal width at point of attachment; 2 distal

pair double, very short and weak.

TYPE-DATA. (1) Culex (C.) jacksoni Edwards, Lectotype o* with attached

genitalia mount; Shonson Hill, HONG KONG, 18 March 1933, R. B. Jackson

(BM; present selection); (2) Culex (C.) fuscifurcatus Edwards, Lectotype o*

with attached genitalia mount, Ootacamund, Nilgiri Hills, Madras, INDIA,

7,500 ft., July 1913, T. B. Fletcher (BM; present selection); (3) Culex (C.)

kangi Lien, Holotype “ (76343-1) with associated pupal and larval skins,

Chienchi, Hsinchu, Hsien, TAIWAN, ground pool, March 1965, C. C. Kang;

deposited in Taiwan Provincial Institute of Infectious Diseases, Taipei, Repub-

lic of China.

DISTRIBUTION. Known only from India, Sri Lanka, China (mainland),

Hong Kong, and Taiwan. Material examined. 78 specimens: 40, 239, 15 L;

34 individual rearings (1 pupal, 33 larval).

INDIA. Madras: Nilgiri Hills, Ootacamund, Wellington; 230, 1592, 22 lp.

SRI LANKA. Kandy District, Rangala; 10, 19, 1 lp.

CHINA. W. Peking; Tsichou; 1“, 5 L (S. Billings 1945).

HONG KONG. Shonson Hill; 4c, 32, 6 L.

TAIWAN. San Is.: Miaoli; Hsinchu Hsien, Hsinchu, Chienchi; 11%, 498,

4 lp.

Additional records from the literature: CHINA: Kwangtung, Amoy (Feng

1938; Hsieh and Liao 1956).

TAXONOMIC DISCUSSION. Prior to this study jacksoni was poorly known

and its identity obscure. This is due to theoriginal description of Edwards

(in Barraud 1934: 452) in which the male genitalia was described as resembling

mimeticus with a further statement that no differences were found between the

2 species. In studying the attached genitalia mount of the type, I have dis-

covered that it differs strikingly from mimeticus in the complete absence of

the basal sternal process of the proctiger. The larval skin associated with the

type is also unique, particularly in the presence of a few peculiar heavy spines

in the distal portion of the siphon, agreeing well with the original description of

Edwards. |

The synonymy of fuscifurcatus Edwards 1934 from Nilgiri Hills, India, and

kangt Lien 1968 from Taiwan with jacksoni as proposed has been based primari-

ly on the similarity in the male genitalia and on the detailed comparison of

adult types, paratypes (especially kangi) and other topotypic reared material.

Apart from considerable variation in the number and size of the peculiar spines

on the larval siphons, all of these forms are virtually similar, indicating that

they are probably conspecific. The larva of the typical Hong Kong form (jack-

soni) has the spines on the siphon varying from 2 to 5, all of which are rather

152 Contrib. ‘Amer,. Ent. Inst. voly12,.’no.''2,° 1976

short and weak; the Taiwan form (kangi) has 4-7 spines all of which are very

strong and long and those from India (fuscifurcatus) usually have 9-11 spines

similarly developed as in the Taiwan form but longer and forming a prominent

double row. In spite of these differences, these larval forms are essentially

similar in the detailed cueeeadioghs and there are no obvious differences in any

other associated stage.

Culex jacksoni appears to be as widespread as mimeticus in the Oriental

region but whether it may be found with the latter in the same breeding site is

not known. The Hong Kong material which I examined at the British Museum

(Natural History) was a mixture of these 2 species, both of which were probably

collected from the same breeding site, judging from the field notes taken by

R. B. Jackson during 1933. In separating the adults of jacksoni from mimeti-

cus, caution should be taken in using the characters noted above. For positive

identification, the associated male genitalia and larvae should also be examined.

BIONOMICS. The larvae and pupae of jacksoni have been collected in gen-

eral ground pools, irrigation ditches and seepage pools in hills or mountain-

ous country at a high elevation. Lien (1968) noted that the larvae (as kangi)

were found in association with specimens of vagans. In the Nilgiri Hills,

Madras, India, Edwards (1934) noted that the adults (as fuscifurcatus) were

collected at 2,200 m. Adult biology and the medical importance of this species

are unknown.

36, CULEX (CULEX) TSENGI LIEN

(Pies):655' 72)

Culex (Culex). isengi Lien 1968; 237 (Co*,; 2%, P*;°L*).

FEMALE (Fig. 65). Wing: 3.4mm. Forefemur: 1.6mm. Proboscis:

1.8mm. Abdomen: 2.2 mm. As described by Lien (1968: 237-40); very

similar to other members of the mimeticus complex in the pattern of wing

spots and in nearly all characters as described for mimeticus, differing in

smaller size and in the following characters. Head, Narrow decumbent

scales of vertex fine, pale yellowish to whitish; erect scales pale beige or

whitish in center, dark on posterolateral areas. Thovax. Mesonotal scales

relatively narrow, fine and predominately whitish or yellowish on anterior

0.7 of disc except for some dark scales forming spots in middle of fossae

and streaks or blotches on posterior acrostichal and dorsocentral lines cephal-

ad of prescutellar space; scales on prescutellar space and scutellar lobes al-

most whitish. Legs. Midtibia with distinct longitudinal pale stripe as in

jacksoni. Wing. Pattern of wing spots as figured; first costal spot in middle

of vein C usually involving vein Sc only or sometimes also posterior surface

of C; 3rd costal spot near apex of wing absent or with pale scales restricted to

apical portion of vein Rj only; rest of pale spots essentially similar to mimetz-

cus, Abdomen. Basal pale bands on all terga rather narrow; terga V-VII with-

out median apical pale bands.

MALE. Essentially similar to female, differing slightly from mimeticus,

fasyi and jacksoni particularly in the following combination of features. Head,

Palpal segments 4 and 5 weakly plumose. Proboscis with a weak ventral tuft

of about 6 hairs at base of median pale ring; these hairs about 3 times as long

as labial width at point of insertion; dorsal surface of labium with pale scale

line proximad of median pale ring.

MALE GENITALIA (Fig. 72). As figured; differing from mimeticus and

Sirivanakarn: Subgenus Culex in the Oriental Region 135

jacksoni in the presence of a short and slender basal sternal process of the

proctiger which is similarly developed as in fasyz, Phallosome. Apical tergal

crown of inner division with 2 stout fingerlike processes. Pyroctigeyr. Basal

sternal process rudimentary, its length about 0.01 mm; paraproct without any

minute spicules distad of subbasal process.

PUPA. As figured for mimeticus and jacksoni (Figs. 67, 70); differing

slightly in the following. Tvumpet. Shorter, about 0.6 mm in length. Cepha-

lothovax. Setae 8,9-C double or triple. Abdomen, Setae 6-III, IV double or

triple; 6-V, VI double.

LARVA (Fig. 72). Asfigured and described for mimeticus (Fig. 67) and /fasyi to

which it is essentially similar in the detailed chaetotaxy, differing from both

and from jacksoni as follows. Szphon. More or less strongly tapered distally;

apical 0.5 with 1-3 strong, long spines ventrally or subventrally; pecten teeth

15-22, in row extending from extreme base to 0.5 or more of total length; 4

proximal pairs of siphonal tufts strong, most proximal one strongest, 2-3

times as long as siphonal width at point of attachment, the rest gradually short-

er; seta 2-S relatively long, stout and very dark.

TYPE-DATA. Holotype “ (62038.9) with associated pupal and larval skins,

Chihtuan, Loshui, Tatung, Ilan Hsien, TAIWAN, 1,200 m, seepage, 9 Septem-

ber 1961, C. L. Chung and L. C. Lu; (deposited in Taiwan Provincial Institute

of Infectious Diseases, Taipei, Republic of China).

DISTRIBUTION. Known only from Taiwan. Material examined. 10 speci-

mens: 5c, 52, 10 individual larval rearings.

TAIWAN. Tatung (type-locality); Pingtung, Lai-I; Taipei, Wulai; Chia I.

TAXONOMIC DISCUSSION. The taxonomic status of tsengi and the problem

as to whether or not it may be conspecific with or only subspecifically distinct

from mimeticus, fasyi and jacksoni can not be definitely determined. In this

study, I tentatively recognize tsengi as distinct on the basis of the extent of

certain wing spots, male palpus and proboscis, pupal and larval stages as

described above.

The adults of fsengz can be distinguished from mimeticus, fasyi and jack-

soni by the first costal spot of the wing usually involving only vein Sc and by

having the 3rd costal spot near the apex of wing usually involving only the apical

portion of vein Ry, or completely absent. The male genitalia differ from mime-

ticus in a shorter basal sternal process of the proctiger, but are extremely

similar to and virtually indistinguishable from fasyi. The larva resembles

mimeticus and fasyi very closely except for the presence of a few strong spines

on the siphon. Further study of specimens from progeny rearing is critically

needed to establish the affinities of this form.

BIONOMICS. According to Lien (1968), tsengi is widely distributed in

mountainous regions throughout Taiwan. The type and other specimens in the

type-series were obtained from rearing the larvae in ground pools containing

green algae in a dried up stream bed at an elevation of 1,200 m. They were

frequently found in association with bitaeniorhynchus, Nothing is known about

the adult biology and the disease relationships of this species.

37, CULEX (CULEX) MIMULOIDES BARRAUD

(Figs. 65, 73, 74)

Culex mimeticus Noe var. mimuloides Barraud 1924a: 991 (o"* 9).

Culex (Culex) mimeticus Noe (in part), Edwards 1932: 205 (taxonomy);

Barraud 1934: 409 (“*, ¢, L).

154 Contrib. “Amer, Ent. Inst,, wok. 12.00.42,-1976

Culex (Culex) mimuloides Barraud, Edwards, in Barraud 1934: 451 (%, 9,

key).

FEMALE (Fig. 65). Wing: 4.0-5.4 mm (average 4.7 mm). Forefemur:

1,.6-2.5 mm (average 2.0 mm). Proboscis: 1.8-2.6 mm (average 2.2 mm).

Abdomen. 3.0mm. Medium or large sized; essentially conforming to the

mimeticus complex in having first costal pale spot in middle of vein C involv-

ing only veins C and Sc. Head, Erect scales of vertex usually pale yellowish

in center, dark or black on posterolateral areas, sometimes entirely dark.

Proboscis with median pale ring occupying 0.15-0.20 of total length. Cibarial

Armature, Essentially similar to mimeticus. Thorax. Mesonotal scales

coarse, largely dark brown in middle, or partially pale, golden or yellowish

on anterior 0.7 of disc, not producing a distinct pattern; scales on posterior

margin of fossae and on marginal areas golden or yellowish; scales in middle

of prescutellar space and scutellar lobes dense and entirely pale. Legs, An-

terior surface of fore- and midfemora without light speckling of pale scales;

hindfemur without light speckling of pale scales dorsad or distad of longitudinal

pale stripe on anterior surface; mid- and hindtibiae with longitudinal pale

stripe extending from basal 0.15 to about 0.75 of whole length; tarsomeres 1-4

of all legs with very broad basal and narrow apical pale bands. Wing, Pattern

of pale spots and venation as figured; vein Ry,5 usually completely dark or

sometimes with a short pale scale line restricted to apical 0.5; furcation of

veins Ro,9 and M usually pale, sometimes dark; Cu, completely dark or with

a short pale scale line in middle; cell Mg longer than cell Ro. Abdomen, Bas-

al pale bands on all terga narrow or broad. Genitalia. As figured for mimeti-

cus; tergum IX with a lateral row of 6-8 setae on each side; postgenital plate

truncate or lightly emarginate on caudal margin; vaginal sclerite strongly

sclerotized, U- or V-shaped; insula with a strong tuft of about 10 setae.

MALE. Essentially as described for female; palpus and proboscis similar

to other members of the mzmeticus complex, differing slightly in the following.

Head, Ventral scalelike setae of palpal segment 3 widely spaced and apparent-

ly less dense; palpal segments 4 and 5 moderately plumose. Proboscis with

narrow median pale ring; ventral tuft of hairs at base of median pale ring very

weak or not distinct; no scattered pale scales proximad or distad of median

pale ring.

MALE GENITALIA (Fig. 73). Similar to mimeticus and other related

forms, differing particularly in the following details. Subapical lobe. Setae

d-f of distal division strong, most distal longest, nearly as long as setae a-c

of proximal division; leaf g very narrow, short and lanceolate. Phallosome.

Apical tergal crown of inner division with 3,4 rather short fingerlike processes,

all curved, porrect, with apices projecting caudad; sternal spine of outer divi-

sion strong and sharply angulate at base with apex projecting laterad; mesal

Spine strong and long; basal lateral process broad, not produced into a distinct

knob. Pyroctiger. Basal sternal process moderately thick and relatively long,

about 0.06 mm or Slightly more; cercal setae 2.

PUPA (Fig. 73). Abdomen: 3.4 mm. Paddle: 0.97 mm. Trumpet: 0.78

mm; index 8. Exceedingly similar to mimeticus, fasyi and jacksoni, differing

particularly in the following combination of chaetotaxy. Cephalothorax. Setae

8,9-C double, rarely single or triple. Abdomen. Seta 1-III usually 6,7 bran-

ched (5-8); 1-IV usually 5 branched (4, 5); 1-V 3, 4 branched; 6-III, IV usually

double or triple (2-4), 6-V, VI usually triple or 4 branched, rarely double; 5-V,

VI double, 1.5 times as long as segment following; 4, 5-VII usually single (1, 2);

4-VIII usually single (1, 2).

Sirivanakarn: Subgenus Culex in the Oriental Region 155

LARVA (Fig. 74). Head: 0.80 mm. Siphon: 1.8-2.2 mm; index 8-9.

Saddle: 0.44mm;siphon/saddle ratio4-5. Asfigured; distinguished from other

members of mimeticus complex particularly by following characters. Head,

Antenna about as long as head; spicules in basal 0.5 of antennal shaft very

strong; seta 5-C usually 5 branched (4-6); 6-C triple; 7-C 7,8 branched. Thor-

ax, Seta 4-P double. Abdomen. Seta 6-I usually triple, sometimes 4 bran-

ched; 6-II always triple; 6-III-VI double; 1-IJI-VI double, about 0.5 of length

of setae 6-III-VI. Comb scales small, 40 or more, all with more or less

rounded apical fringes of evenly fine spicules. Saddle lightly or strongly in-

fuscated along caudal and basal margin; spicules weak and few; seta 1-X double;

2-X usually triple (2-3); anal gills slender, pale, 1,2 times as long as saddle.

Siphon, Rather thick, straight, distally slightly tapering and relatively long;

apical portion without spines on ventral surface; pecten teeth 11-15, 8-14 teeth

in basal 0.3 of siphon short, small, barbed with fine denticles; 2-5 distal

teeth very strong, heavily pigmented, entirely simple or with a few weak denti-

cles restricted to base; siphonal tufts 6 pairs (or total 12), widely spaced and

short; pairing regular; 5 proximal pairs inserted subventrally, 4,5 branched

each, as long as or shorter than siphonal width at point of attachment; 1 distal

pair lateral, weak, short and double.

TYPE-DATA. Lectotype “* (one of the 2 specimens, No. 1484, marked

as type by Barraud) with genitalia slide, Nilgiri Hills, Madras, INDIA, October

1915, Khazan Chan (BM; selection of Mattingly 1956: 37).

DISTRIBUTION. India and China (mainland). Material examined. 122

Specimens: 36, 569, 30 L; 71 individual rearings (4 pupal, 67 larval).

INDIA. Madras: Nilgiri Hills (type-locality), Ootacamund, Coonoor, Ban

Dishola, Lovedale, Kurkuthi, Burliar, Toll Gate, Selas, Aruwam Kadu, Yed-

dapalli; 36°, 062,°20'1,, 4 0..07 1p.

CHINA. Yunnan: Poashan Valley; 10 L (APO 927, S. Billings, 1944).

TAXONOMIC DISCUSSION. The above description of mimuloides has been

based on the type and several other topotypic reared specimens recently col-

lected by B. N. Mohan from the Nilgiri Hills area. The new record of mimu-

loides from China is based on the larvae which were collected by S. Billings in

1944.

Culex mimuloides was originally recognized as a variety of mimeticus by

Barraud (1924a: 991-3, 1934: 409-12) but later, Edwards (in Barraud 1934:

451) elevated it to specific rank among the members of his mimeticus group.

The wing venation of adults of mimuloides is very distinctive, as described by

Edwards (1934: 451-2), in having the furcation of vein M proximad of that of

vein Ro,3 or cell Mo distinctly longer than cell Rj. This character is constant

and provides a clear cut separation of mimuloides adults from the rest of the

mimeticus complex which exhibit furcations of Ro +3 and M approximately at

same level, with cell Ro as long as or Slightly longer than cell My. The pattern

of wing spots of mimuloides is very variable and is difficult to be used in dif-

ferentiating it from mimeticus and related species. Other stages of mimuloides

are distinct and can be readily separated from the rest of the mimeticus com-

plex.

Culex mimuloides is somewhat intermediate between members of the

mimeticus and mimulus complexes, but apparently resembles mimeticus more

than mimulus in the pattern of wing spots, male genitalia, pupal and larval

stages. The male phallosome and the development of the basal sternal process

of the proctiger of mimuloides are, however, rather similar to or nearly

approaching those of the mimulus complex and the members of the Vishnui Sub-

croup.

156 Contrib. Amer,, Ent. Inst...vol.. 12; no...2, 1976

In the Nilgiri Hills, Madras, mimuloides has been found to be sympatric

with jacksoni and mimeticus without apparent intergradation in their diagnostic

characters. e:

BIONOMICS. Culex mimuloides has been encountered on many occasions,

particularly in the type-locality at Nilgiri Hills, Madras, in southern India.

The larvae have been collected in ground pools, stone tanks and borrow pits

in marshy areas. These breeding sites are at very high elevations. No further

data are available on the conditions of the breeding sites. Most of the adults

were obtained from rearing the larvae and only a few were caught in the field.

Nothing is known about the adult biology and the medical importance of this

species.

38. CULEX (CULEX) DIENGENSIS BRUG

(Figs. 66, 69)

Culex (Culex) diengensis Brug 1931: 35 (*, 2); Edwards, in Barraud 1934:450

(adult key).

FEMALE (Fig. 66). Wing: 4.8mm. Forefemur: 2.1mm. Proboscis:

2.3mm. Description based on 2 specimens (marked as cotypes by Brug)

from Dieng, Java, Indonesia. Essentially conforms to other members of the

mimeticus complex, distinctive in the following combination of characters.

Thorax, Anterior 0.7 of mesonotum predominantly golden brown scaled except

for dark scale spots in middle of fossae. Legs, Anterior surface of fore- and

midfemora without speckling of pale scales, subapical yellowish spots very

distinct; anterior surface of hindfemur uniformly pale in basal 0.5; apical 0.5

largely dark, with light speckling of pale scales on dorsal lateral surface; all

tibiae without longitudinal pale stripe on anterior surface. Wing. Pattern of

pale spots as figured; differing from mimeticus and other related forms parti-

cularly in vein Ry,5 pale from basal 0.1 to almost at apex; vein Cu, pale from

level of crossvein m-cu to about 0.75 of total length. Abdomen. Basal pale

bands on all terga moderately broad.

MALE. Based on the holotype which is essentially similar to female in the

pattern and extent of wing spots and in general characters; palpus and probos-

cis broken off and lost.

MALE GENITALIA (Fig. 69). Description based on the holotype slide.

Very distinctive; differing from all members of the mimeticus complex and

Subgroup in the following. Basimere. Very large, basal and middle portions

strongly swollen, distal portion narrow. Subapical lobe, Setae on proximal

and distal divisions poorly developed; setae a-c very thin and short; setae d-f

consisting of 4 narrow pointed blades and 3 other rodlike setae; leaf g absent;

seta f inconspicuous. Distimere. Strongly modified; shape irregular; basal

portion thick, middle portion strongly swollen, distal portion markedly re-

duced to a long and thin rod which is curved and strongly bent downward; sub-

apical claw short and small; dorsal and ventral setae absent or inconspicuous.

Phallosome. Apical portion of inner division with 4,5 strong, long fingerlike

processes, all diverging laterad; outer division without mesal and sternal

spines. Proctigey. Basal sternal process well developed, rather thick, long,

strongly bent sternad; cercal setae 3.

PUPA. Description based on one specimen associated with one of the above

cotype females. In general as figured and described for mimeticus (Fig. 67)

and other related species, differing particularly in the following features.

Sirivanakarn: Subgenus Culex in the Oriental Region 157

Trumpet, Longer, about 0.85 mm in length; index 6.5. Cephalothorax. Seta

8-C 3,4 branched. Abdomen. Setae 5-IV, V double; 5-VI single; 6-III triple;

6-IV-VI 3, 4 branched; 4, 5-VII double; 9-VII double or triple; 4- VIII double;

9-VIII 3-6 branched. Paddle, Very broad; about 1.0 mm in length; inner and

outer parts entirely pale, without infuscate area.

LARVA. Unknown.

TYPE-DATA. Holotype “* (17.267), with attached genitalia mount,

Merdodo Lake, Telaga Warna (Diénghochebene), Java, INDONESIA; Peat moss

ground pool, 2,000 meters, 2-3 June 1929, A. Thienemann (BM).

DISTRIBUTION. Known only from the type-locality in Java, Indonesia.

Material examined. 3 specimens: 1% (holotype), 22 (marked as cotypes by

Brug); 1 individual pupal rearing.

TAXONOMIC DISCUSSION. Culex diengensis is the most distinctive form

of the mimeticus complex and is strongly characterized by several peculiar

modifications in the male genitalia. It is assigned to the mimeticus complex

on the basis of the pattern of wing spots and general external characters of the

adults. As in mimuloides, the male phallosome and proctiger of diengensis

are apparently more similar to the members of the mimulus complex than to

other members of the mimeticus complex.

The previous descriptions of the adult wing spots by Brug (1931) and the key

to adults, using the extent of pale scaling of vein R and Cu, by Edwards (in

Barraud 1934: 450) are applicable to this species. tte pupa dt diengensis,

which is described here from a single skin associated with one of the 2 cotype

females, can be readily separated from other members of the mimeticus com-

plex by the branching of setae 8-C, 5-IV-VI and 6-III-VI.

BIONOMICS. The type specimens of diengensis came from a single collec-

tion in peat moss ground pools in a mountainous region at an elevation of 2, 000

m. The adults were apparently obtained from rearing the larvae and pupae.

Adult biology and the medical importance of this species are unknown.

39. CULEX (CULEX) MIMULUS EDWARDS

(Piggend 6.09 p18 9 7)

Culex mimulus Edwards 1915: 284 (%, 2); Barraud 1923a: 942 (L*, in part);

Barraud 1924a: 993 (o*, ¢); Borel 1930: 324 (*, 9*, L*, in part).

Culex mossmani Taylor 1915: 181 (“*, 2); Edwards 1924: 394 (synonymy).

Culex (Culex) mimulus Edwards, Edwards 1932: 205 (taxonomy); Barraud 1934:

4A2 (SO He QoL ie part) Edwards, in Barraud 1934: 450 (adult, key);

Bonne-Wepster and Brug 1937: 66 (co, °*); Baisas 1938: 214 (“*); Feng

1938: 302 (distribution); Bonne-Wepster and Brug 1939: 1275 (L*); Lee

1944: 104 (L*); Brug and a -Wepster 1947: 187 (distribution); Bonne-

Wepster 1954: 127 (o, 2*, L*); Safyanova et al 1964: 1178 (distribution);

Delfinado 1966: 150 (o'*, 9). Bram: 196 7a 1213 (6% iO. Phy Tye),

Culex (Culex) confusus Baisas 1938: 216 (*, $); Delfinado 1966: 150 (synonymy).

Culex (Culex) neomimulus Lien 1968: 240 (“*, 9*, P*, L*). NEW SYNONYMY.

FEMALE (Fig. 16, 75). Wing: 3.2-4.1 mm (average 3.6 mm). Fore-

femur: 1.4-2.0 mm (average 1.6 mm). Proboscis: 1.5-2.0 mm (average

1.75 mm). Abdomen: 2.2-2.8 mm (average 2.5mm). Small or medium sized

Species; wing length rarely exceeding 4.0 mm; distinguished from the members

of the mimeticus complex by having first costal pale spot at about middle of vein

C, involving veins C, Sc and Ry or sometimes Rg, Mand Cu. Head. Narrow

158 Contrib. Amer, Ent.’ Inst., vol. 12; no.°2) 1976

decumbent scales of vertex rather fine, whitish or yellowish, forming a dis-

tinct ocular line; erect scales largely pale, yellow or golden in broad median

area, dark brown to black posterolaterally; lateral patch of broad appressed

scales whitish, very distinct. Palpus completely dark scaled or sometimes

apex of segment 4 tipped with a few pale scales. Proboscis with very distinct

median pale ring which occupies at least 0.2 of total length. Cibarial Armature,

Cibarial bar with a concave row of about 30 teeth; median 4-6 teeth narrow and

apically pointed; lateral teeth coarser, flattened and apically pointed or blunt,

all subequal in length. Thovax. Mesonotal integument reddish brown to black;

scales on anterior 0.7 of mesonotum usually predominantly pale beige, yellow

or golden with mixture of some dark scales forming distinct longitudinal streaks

in middle of acrostichal and posterior dorsocentral areas laterad and cephalad

of prescutellar space; scales on prescutellar space sparse and entirely pale;

scutellar scales pale. Apn and ppn with narrow, yellow or golden scales.

Pleural integument darker above, paler below; pale scale patches on stp and

mep reduced or relatively small. Legs. Anterior surface of fore- and mid-

femora dark and without subapical pale spot; anterior surface of hindfemur en-

tirely dark or with well defined pale stripe from base to near apex, no scat-

tered pale scales among dark scaled areas on dorsal lateral surface; all tibiae

without longitudinal pale stripe on anterior surface; tarsomeres 1-4 of all legs

with broad basal and narrow apical pale bands. Wing. Pattern of pale spots

as figured; extent of pale spots variable; base of C usually without pale scaled

streak or spot, sometimes with distinct pale scale line from base to 0.9 of

total length; first costal spot usually involves veins C, Sc and Rj, sometimes

extending to veins Rg, M and Cu; 2nd costal spot involves veins C, apical

portion of Sc and Rj; 3rd costal spot near apex of wing usually involves apical

portions of veins C, Rj and R9, sometimes restricted to Rj and Ro or R, only;

furcations of veins Ro 43 and M approximately at same level and usually pale,

sometimes dark; R4,, usually entirely dark or with pale scale line of variable

length in middle; Cu , with pale scaled line in basal 0.5, apical 0.9 dark; Cu

with pale scales restricted to apical 0.1 or less, fringe scales on apex of Cug

pale, rest dark; 1A usually with pale scale line extending from basal 0.1-0.5,

apical 0.5 dark; scales on Rg, Rg and Ry,- narrow and dense. Abdomen.

Basal pale bands on terga II- VII narrow or broad; basolateral pale spots ex-

tending to 0.5 of segment length; sterna with broad basal pale bands and narrow

apical dark bands. Genitalia. Essentially similar to mimeticus from which

they are indistinguishable.

MALE (Fig. 75). In general similar to female, differing in less dense wing

scales and poorly defined wing spots, particularly on veins Ro, Rg, Cuy and 1A.

Head, Palpus exceeding proboscis by 1.0-1.5 of length of segment 5; segment

3 with a broad median pale band on dorsal surface, ventral surface with short

scalelike setae along whole length, apical 0.25 with ventral lateral tuft of 10-

12 bristles; segments 4 and 5 moderately to strongly plumose and each with

distinct basal pale band dorsally; apical 0.25-0.30 of segment 9 pale to tip.

Proboscis with a weak ventral tuft of 4-6 short hairs at base of median pale

ring, these hairs 2-3 times as long as labial width.

MALE GENITALIA (Fig. 76). Extremely similar to vzshnui and other

members of the vishnuz complex. Segment IX. Tergal lobe poorly developed,

with a row of 5,6 setae. Basimeve. Slender, conical, with several setae in

2,3 rows laterad of subapical lobe. Swubapical lobe. Setae d-f of distal division

3, all apparently flattened and subequal in size and length; leaf g short, fairly

broad, with blunt or acuminate apex. Distimere. Normal; with very weak but

distinct subapical crest of minute spicules on dorsal surface; ventral and dor-

Sirivanakarn: Subgenus Culex in the Oriental Region 159

sal minute setae present beyond middle; subapical claw short, simple, flatten-

ed and apically blunt, about 0.006 mm in length. Phallosome. Apical tergal

crown of inner division porrect, composed of 5,6 strong fingerlike processes,

all curved and with apices projecting caudad; apical sternal spiculate portion

irregular, with a few to several small denticles; outer division with a strong

mesal tergal spine and 1 heavy sternal spine, the latter strongly or gently bent

at base with apex projecting basad; lateral basal process slightly or strongly

produced into a rounded knob. Pyroctiger. Apical crown large, dark, com-

posed of several coarse spicules; basal sternal process well developed, very

thick, long, dark and strongly curved sternad; subbasal process distinct and

prominent; cercal setae 2.

PUPA (Fig. 76). Abdomen: 3.0 mm. Paddle: 0.74mm. Trumpet:

0.65-0. 84 mm; index 8-13. Cephalothorax and abdomen very pale whitish or

lightly yellowish with darkened area not forming a definite pattern. Complete

chaetotaxy as figured, distinctive in the following. Trumpet. Moderately to

very long and uniformly cylindrical; sculpturing weak; basal 0.5 of meatus

dark, annulate; apical 0.95 pale yellowish; pinna short; apical margin truncate.

Cephalothovax, Seta 8-C usually double (2-4); 9-C double. Metanotum. Seta

10-C 5-7 branched. Abdomen. Seta 1-II forked into 3-6 branches; 1-III 9-10

branched; 1-IV 7-9 branched; 1-V 4,5 branched, 1-V, VI triple or 4 branched

(2-4); 5-IV 3-5 branched, slightly shorter or longer than segment V; 5-V, VI

double, 1.5 times as long as segment following; 6-III, IV usually triple (2, 3);

6-V, VI 3,4 branched; 2-VII laterad or mesad of 1-VII; 4-VII double; 5-VII

usually single (1,2); 9-VII usually 4 branched (3-5); 4-VIII double; 9-VIII 8-10

branched. Paddle. Broad, entirely pale, without darkened areas.

LARVA (Fig. 77). Head: 0.71-0.78 mm (average 0.75 mm). Siphon 1.9-

2.7 mm (average 2.0 mm); index 9-14. Saddle: 0.36 mm; siphon/saddle ratio

4-7. As figured, distinguished from other members of the Mimeticus Sub-

eroup by having a long, thin and pale siphon and by the following additional

features. Head. Integument of head capsule usually whitish or yellowish white;

seta 5-C usually 3,4 branched (2-4); 6-C usually double (2,3); 14-C minute,

inconspicuous. Antenna about as long as head; spicules strong and numerous.

Thorax, Seta 4-P single, about as long as 3-P; setae 8, 9-M usually 4 bran-

ched (4,5); seta 7-T usually 5 branched (5, 6); 13-T 5-9 branched. Abdomen.

Setae 6-I, II triple; 7-I single; all setae 5-I-VI always branched; 1-III-VI usu-

ally double or triple (2-5); 6-III-VI double. Comb scales about 30 in number,

all small, subequal, apical fringe of spicules terminating into a weak median

Spine. Saddle same color as siphon; seta 1-X weak, double or triple; 2-X usu-

ally double (1,2); anal gills slender and long, 1.5-4.0 times as long as saddle.

Siphon. Very long, thin and usually whitish or yellowish, sometimes light

brownish; distal portion more or less uniform in width or gradually tapering;

pecten teeth 16-24, 4,5 distal teeth with lateral barb of 5-7 delicate denticles

and a sharp straight apical spine; siphonal tufts usually 6 pairs (total 12),

sometimes 7, all short and weak, 3,4 branched, widely spaced and regularly

paired; 5 proximal pairs inserted subventrally beyond pecten to about 0.75 of

total siphon length, 1 pair lateral, inserted beyond the middle; 3 proximal

subventral pairs as long as or slightly longer than siphonal width at point of

attachment, rest shorter and weaker; seta 2-S dark, slender and moderately

long; median caudal filament of spiracular apparatus absent.

TYPE-DATA. (1) Culex mimulus Edwards, Lectotype °* (one of 2 syn-

types) with genitalia slide, Kuching, Sarawak, BORNEO, J. C. Moulton

(BM; present selection); (2) Culex mossmani Taylor, Type (?), Mossman,

Queensland, AUSTRALIA (US); (3) Culex (C.) confusus Baisas, Holotype

160 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

co (R79-a), Balabac, Balabac Island, PHILIPPINES, 23 June 1934, F. Guinto

(PBH; non-existent); (4) Culex (C.) neomimulus Lien, Holotype “ (78788. 32)

with associated pupal and larval skins, Pinglin, Shuangchi, Taipei Hsien,

TAIWAN; ditch, 17 November 1966, L. C. Lu; (deposited in Taiwan Provincial

Institute of Infectious Diseases, Taipei, Republic of China).

DISTRIBUTION. Widespread throughout Southeast Asia, extending west in

India and Sri Lanka; north in China, east and south in Papua-New Guinea and

northern Australia. Material examined. 974 specimens: 3160, 36492, 294 L;

314 individual rearings (113 pupal, 201 larval).

INDIA. Bombay: Deccan; Karwa; Assam: Dibrugarh; Jorhat; Rupsi; Ledo;

Bengal: Pusa; 4c, 89, 13 L.

BANGLADESH. Sylhet; 1%, 3 L, 1 lp.

SRI LANKA. Central Province: Kandy District; North Central Province:

Anuradhapura District; Northern Province: Vavuniya District; Suduganga;

Kumnegala; 3c, 42, 8 L.

THAILAND. Chiang Mai: Doi Huey Som; Nan: Ban Sa La; Ban Pha Man;

Ban Fang Min; Phuang Chom Phu; Ban Ta Loe; Lampang: Ngao; Ban Pha Daeng;

Ban Pha Khoi; Khon Kaen; Mae Hong Son; Nakhon Ratchasima: Khao Yai;

Pachong; Ban Musi; Kanchanaburi: Sai Yok; Ban Sai Noi; Chon Buri: Bang La

Mung; Bang Pain; Ranong: Ban Bang Laem Phu; Suvat Thani; Phangnga: Pathum;

Khao Sung; Nakhon Si Thammarat: Ban Thuan Lek; Songkhla; Narathiwat; Stul:

Muang; 570, 702, 113 L, 76 p; 22 Ip.

MALAYSIA. Peninsular Malaysia-Selangor: Rantau Pangjang; Ulu Gombak;

Ulu Langat; Kuala Lumpur; Kepong F. R.; Perak: Chior F. R.; Tg. Rambutan;

Kuala Kangsar; Kg. Kuala Dipang; C. Highlands Rd. ; Chikus; Seremban, Pahang:

Chegar Perah; Fraser Hills; Bentong; Gunong Benom; Kuantan; Trengganu:

Marang; Kemaman; Johore: Labis; Senai; N. S. (Negeri Sembilan): Seremban;

Cape Rachado F. R.; Kg. Inas; 1580, 1632, 92 L, 34p, 1151p. Malaysia-

Sabah: Tawau; Kudat; Semporna; Beaufort; Tenom; Sipitang; Mt. Kinabalu;

Kota Kinabalu (Jesselton); Keningau; Brunei: Tutong District; Savawak: Kuching

(type: locality); 65¢%:839, 5.) 3 p.o31 Ip.

SINGAPORE. 1, 19.

INDONESIA. Sumatra: Bengkulu; Mt. Tebo; 10°, 22; Sulawesi; 2 L.

PAPUA-NEW GUINEA. Morobe: Wau; Mt. Missim; Sepik: Maprik; 3c,

pase heise

PHILIPPINES. Palawan: Puerto Princesa; Tacburos; Iwahig; Irahnan

River; Balsahan River; Bacungan; Leyte: Tacloban; Jolo Jolo Is.; Muni;

Isabela: Basilan; Mindanao: Agusan, S. Francisco; 140, 219, 22 L, 27 lp.

HONG KONG. 1¢, 19.

TAIWAN. Wan Luan, Pintung; Sun Moon Lake, Keelung; Pinglin, Shuangchi,

Tapei Hsien; 70, 72, 36 L, 2 lp.

CHINA. Hanchow; Checkiang; 3c, 29.

Additional records from the literature: PAKISTAN (Aslamkhan 1971);

BURMA (Barraud 1934). CHINA: Hunan, Kwangtung, Hainan, Amoy (Feng

1938; Chu 1958; Hsieh and Liao 1956). INDONESIA: Banka; Java; Edam;

Borneo (Kalimantan); Kabaena (Brug and Bonne-Wepster 1947; Bonne-Wepster

1954); West Irian: Hollandia (Kota Baru) (Assem and Bonne-Wepster 1964).

VIETNAM: Cao Bang; Than Hoa (Safyanava et al 1964). AUSTRALIA: Queens-

land (Edwards 1924; Lee 1944).

TAXONOMIC DISCUSSION. Culex mimulus is the most dominant and wide-

spread form of the spotted wing Culex in Southeast Asia and adjacent areas

where it probably originated and subsequently dispersed eastward into New

Guinea and northern Australia. As discussed by Bram (1967a) and as evident

Sirivanakarn: Subgenus Culex in the Oriental Region 161

here, all stages of mzmulus are very variable. Recently, 2 new additional taxa,

neomimulus and murrelli, have been recognized by Lien (1968) from Taiwan.

The present study of the mimulus material from several parts of Southeast Asia,

including also some paratypes of Lien's species from Taiwan indicates that a

complex of forms is involved which exhibits extremely similar adults but with

strongly differentiated larvae, somewhat resembling the situation in the vishnuz

complex. Unfortunately, Lien (loc. cit.) in describing his new species did not

attempt to revise the entire complex or to describe all stages of the typical

mimulus for comparison. It is evident, however, that the typical mimulus and

the 2 new species described by Lien are widespread and apparently exhibit gen-

eral overlap throughout Southeast Asia and India. Since the typical mzmulus

was originally described only from adults (Edwards 1915), it is not possible to

determine exactly as to which adult form the larva and pupa should belong.

However, it appears most probable that only 2 taxa are involved in the complex

and that these should be recognized as distinct species. One of these is mimulus

of most authors and the other is murrelli described by Lien (1968) from Taiwan.

In comparing the paratypes of nmeomzmulus Lien with the type and topotypic speci-

mens of the typical mimulus, I found them extremely similar in all stages.

The adults of neomimulus which were described by Lien as distinct from mimu-

lus in having the first costal spot involving veins C, Sc, R,, Rg and sometimes

also M and Cu exhibit apparently nothing more than the normal variation of

mimulus populations from the rest of Southeast Asia. On this basis, I am con-

vinced that they are conspecific and am synonymizing neomimulus with mimu-

lus.

Culex mimulus is very variable but can be readily separated from the rest

of the Mimeticus Subgroup except murvrelli by the pattern and extent of the wing

spots as described above. It differs from murvrelli slightly in having the first

costal spot usually extended to veins Rg, Mand Cu. In the breeding habitat,

typical mimulus is more or less restricted to general ground pools with living

vegetation whereas murrelli is usually restricted to rock pools containing de-

cayed leaves.

The affinities of mzmulus and its closely related forms, murvelli and pro-

pinquus are not clear, but they appear to be somewhat intermediate between

members of the mimeticus and vishnui complexes. They are similar to the

mimeticus complex in the development of the wing spots in the adults and in

several pupal and larval characters but are more similar to the members of

the vishnui complex than to the mimeticus complex in the type of the male

phallosome and in the development of the basal sternal process of the proctiger.

BIONOMICS. Culex mimulus is locally fairly common and has been fre-

quently encountered in hills and mountains toward inland areas. The larvae and

pupae have been collected in a variety of ground pools, including ponds, wells,

ditches, stream pools and marshy depressions but not in rock pools. These

breeding sites usually contain green algae, mosses, grasses and other aquatic

vegetation. The habitats are fully exposed to sunlight, under partial or heavy

_ shade of forest bushes at an elevation ranging from above sea level to 1,400 m

or more. The immatures have also been found in artificial containers such as

concrete, stone or cement tanks, coconut shells, snail shells (Delfinado 1966),

bamboo stumps and treeholes. The majority of the adults have been reared

from pupae or larvae and only a small number were from general field catches

and light traps. Bram (1967a) noted that in Thailand adults were collected in

houses. Colless (1959) reported that in Singapore females probably feed on

birds but are not known to bite man. Nothing is known about the medical impor-

tance of mimulus.

162 Contrib. “Anver.. Ent. Inet.) vole 12; no..2, 1076

40. CULEX (CULEX) MURRELLI LIEN

(Figs. 66, 78, 79)

Culex (Culex) murrelli Lien 1968: 243. (c'*, 9*,. P*, L*).

Culex mimulus Edwards 1915 (in part), Barraud 1923a: 942 (L*).

Culex (Culex) mimulus Edwards (in part), Barraud 1934: 412 (“*, 9, L*);

Bram JO6%ar213- (cy Sa Ps *):

FEMALE (Fig. 66). As described by Lien (1968: 243-6). Exceedingly

similar to mimulus in general facies and in the pattern of wing spots, differing

slightly in the following characters. Head. Palpus usually tipped with pale

scales on apex of segment 4. Cibarial Armature. As described and figured

for mimulus from which it is indistinguishable. Thorax, Mesonotum largely

yellowish or golden scaled and usually with dark scales forming more distinct

spots, streaks or patches on fossae, acrostichal line and on posterior dorso-

central areas laterad and cephalad of prescutellar space. Wing. Base of vein

C without pale scale streak or line on posterior surface; first costal pale spot

at middle of vein C restricted to C, Se and Rj; vein R45 usually with pale

scaled line in middle portion, sometimes entirely dark. Genitalia. As des-

cribed and figured for mimulus from which it is indistinguishable.

MALE. As infemale, differing from mzmulus in the following. Head,

Palpus relatively shorter, exceeding proboscis by at most the length of seg-

ment 9; apical 0.25 of segment 3 with ventrolateral tuft of 5-10 dark bristles;

segments 4 and 5 apparently with shorter and fewer bristles on mesal and later-

al surfaces.

MALE GENITALIA (Fig. 78). Exceedingly similar to mimulus in every

detail of basimere, subapical lobe, shape of distimere, phallosome and proc-

tiger; differing chiefly in a more slender and longer subapical claw of the disti-

mere which is about 0.01 mm in length.

PUPA (Fig. 78). Abdomen: 2.4mm. Paddle: 0.72 mm. Trumpet: 0. 62

mm; index 7-9. Differing from mimulus particularly in the following charac-

ters. Trumpet. Relatively shorter and with lower index; coloration darker.

Cephalothorvax. Darker, with considerable amount of brownish tinge forming

more or less definite darkened areas along middorsal ridge, leg, palpal, pro-

boscis and wing cases; seta 8-C always double. Metanotum. Darker, with

considerable amount of brownish tinge; seta 11-C usually double, sometimes

single. Abdomen. Segment I-IV darker, with variable amount of brownish

tinge; seta 1-III 5,6 branched; 1-IV usually 5 branched (4-6); 5-IV double or

triple, rarely 4 branched or more.

LARVA (Fig. 79). Head: 0.78 mm. Siphon: 1.4-1.8 mm (average 1.6

mm); index 6-7. Saddle: 0.36 mm; siphon/saddle ratio 4-5. Complete chaeto-

taxy as figured; differing from mimulus as indicated in the key and in the fol-

lowing additional features. Head. Head capsule darker, usually with consider-

able amount of brownish or blackish tinge; seta 6-C always double. Thorax,

Live specimens usually with considerable amount of dark greenish tinge. Abdo-

men. Color in live specimens same as on thorax; setae 1-III-VI usually double,

sometimes single, rarely triple; 1-IV, V usually strong and long, varying from

more than 0.9 to 1.0 of seta 6. Comb scales 40-50, apical fringe of spicules

normal or rounded and without differentiated median apical spine, sometimes

with median apical spine slightly longer than adjacent spicules, but apparently

not differentiated as a prominent spine as in mimulus. Saddle same color as

Siphon, posterior caudal margin lightly to strongly infuscate; anal gills 1, 2

Sirivanakarn: Subgenus Culex in the Oriental Region 163

times as long as saddle. Siphon. Relatively short, thick, straight and dis-

tally slightly tapering, with lower index and siphon/saddle ratio; color

same as head capsule; pecten teeth 12-13, siphonal tufts stronger, usually 5

pairs (or total 10), sometimes 4 or 6 pairs, more closely spaced; each tuft

3,4 branched; 4 pairs inserted subventrally beyond pecten to about 0.75 of

siphon length, 1 pair inserted laterally beyond middle; most proximal tuft

about as long as siphonal width at point of attachment, rest gradually shorter;

median caudal filament of spiracular apparatus present, rather short or poorly

developed.

TYPE-DATA. Holotype “ (76324. 28) with associated pupal and larval skins,

Hsinhsien, Wulai, Tapez, Hsien, TAIWAN, rock pool, 23 April 1963, C. L.

Chung; (deposited in Taiwan Provincial Institute of Infectious Diseases, Taipei,

Republic of China),

DISTRIBUTION. Probably widespread in Southeast Asia and adjacent areas;

recorded from Taiwan, China, Thailand, Vietnam and Peninsular Malaysia.

Material examined. 737 specimens: 209°, 2209, 308 L; 282 individual rearings

(142 pupal, 140 larval). 3

THAILAND. Chiang Mai: Doi Sutep; Huey Kaew; Mae Hong Son: Ban Mae

Hokae; Lampoon; Khon Kaen; Nakhon Ratchasima: Pak Chong; Khao Yai; Lam

Huey; Tha Khlong; Nakhon Nayok: Khao Keow; Sariga water fall; Kanchanabu7i;

Chon Buri: Khao Nga Chang; Tak: Huey Lang Saeng; Doi Sam Sao; Prvachin Buri:

Khao Din Ngin; Tvat: Koh Chang; Nakhon Sawan; Chanthaburi, Ranong: Khao

Ching Chang; Hard Sampan; Khlong Set Takuat; Prachuap Khiri Khan; Surat

Thani: Koh Samui; Khao Yai; Khao Phlu; Tvang: Muang; Phuket; Songkhla:

Boripat Water Fall; Tone Nga Chang; Yala; 1640, 1749, 253 L, 124 p, 92 lp.

VIETNAM. Dalat; Saigon; Cam Ranh Bay; 10, 149, 7L, 10p, 4 lp.

MALAYSIA. Peninsular Malaysia-Pahang: Frasers' Hills; C. Highlands;

Betong; Perak: C. Highlands Road; 34, 3192, 46 L, 8 p, 42 Ip.

TAIWAN. Taipei: Hsinhsien, Wulai (type-locality); 1%, 12, 2 lp (para-

types).

CHINA. Nanking; 2 L.

TAXONOMIC DISCUSSION. Culex murrelli is apparently widespread in

Southeast Asia but is not as frequently encountered as the typical mimulus. In

addition to the above records, muyrvelli probably also occurs in India judging

from the figures and descriptions of the mimulus larva by Barraud (1923a: 942,

1934: 410). As pointed out by Lien (1968: 243-6) the mimulus larva as figured

and described by Bram (1967a: 217-9) from Thailand was partially referrable

to this species. Within Southeast Asia, murrelli occurs sympatrically with the

typical mimulus over a broad range of elevation but has not yet been found in

Sarawak which is the type locality of mimulus, It appears to be absent in

Borneo, the Philippines, islands of Indonesia and New Guinea where mimulus

is predominant and is the only species of the complex known (Bonne-Wepster

and Brug 1939, Bonne-Wepster 1954, Assem and Bonne-Wepster 1964).

Culex murvrelli is strongly differentiated from the typical mizmulus in the

larva most conspicuously by (1) the shorter, thicker and darker siphon; (2) the

presence of 5 pairs of siphonal tufts (at least 6 in mimulus); (3) the normal

apical fringe of the comb scales, without a distinct apical median spine and (4)

the stronger and longer setae 1-IV-VI which are single or double. The pupa

of murrelli, although showing a great deal of overlap with mzmulus in most

features of the chaetotaxy, can be separated from the latter by the darker

integument of the cephalothorax and abdomen and by the reduction in the num-

ber of branches of setae 1-III] and IV. The adults of murrelli exhibit consider-

able overlap with those of mimulus but differ from the latter slightly in having

164 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

the first costal pale spot of the wing restricted to veins C, Sc and R,. In the

male genitalia, murvrelli differs slightly from mimulus in having a longer and

more slender subapical claw of the distimere, but show no other differences.

As discussed under mimulus, murvrelli apparently exhibits a tendency to breed

in rock pools containing dead leaves rather than general ground pools with

algae, mosses or other living aquatic vegetation as in the case of the typical

mimulus. These differences, which are accompanied by a marked divergence

in several larval characters indicate that it is ecologically distinct and that it

was probably derived from mimulus through specialization in the breeding

habitat. No evidence has yet been obtained to indicate that there is an overlap

in the breeding sites of the 2 species.

BIONOMICS. Culex murrelli is fairly common in the localities where it

has been collected. At the type-locality in Taiwan, Lien (1968) collected the

larvae from rock pools without further detail on the conditions of the breeding

sites. Several larval collections have been made in various parts of Thailand,

and the majority of these were from rock pools, containing numerous decayed

leaves in Secondary tropical forest at an elevation ranging from 100 to 150 m.

In Peninsular Malaysia, the larvae have been collected both from ground pools

with algae and rock pools with decayed leaves at a broad range of elevation

from 100 to 1,400 m. All adults were obtained from rearing the pupae or lar-

vae. Nothing is known about adult biology and the medical importance of

murrelli.

41. CULEX (CULEX) PROPINQUUS COLLESS

| (Fig. 66)

Culex (Culex) propinquus Colless 1955: 315 (9*)

FEMALE (Fig. 66). Description based on the holotype. As described by

Colless (1955: 315-6) with the following redescription. Generally resembling

mimulus in the pattern and extent of wing spots, differing from it most con-

spicuously in having a row or chain of pale spots on veins Cu, Cug, and middle

of 1A and in the following additional characters. Head, Narrow decumbent

scales of vertex finer, fewer and largely whitish, forming a distinct ocular

line; erect scales more slender, predominantly pale golden in center, dark or

blackish on posterolateral areas; lateral patch of broad appressed scales whitish,

very distinct. Palpus entirely dark. Proboscis with a broad median pale ring

which occupies about 0.3 of total length; basal 0.5 proximad of median pale ring

lightly sprinkled with pale scales on lateral and dorsal surfaces. Thorax,

Mesonotal integument dark brown; scales on anterior 0.7 of mesonotum narrow,

rather fine, sparse and predominantly whitish or golden, contrasting sharply

with dark integument underneath; no dark scales forming distinct spots, streaks

or blotches among pale ones; scales laterad of prescutellar space very sparse

and dark, contrasting with pale scales on anterior 0.7 of mesonotal disc;

scales on middle of prescutellar space and scutellar lobe whitish or beige.

Pleural integument reddish brown; stp and mep with relatively small scale

patches; no scales among upper mep bristles. Legs. Fore- and midfemora

with speckling of pale scales on anterior surface; speckling of midfemur more

extensive than that of forefemur; anterior surface of hindfemur with pale stripe

and speckling of pale scales among dark scaled area dorsad of stripe; tibiae

and tarsomeres 1 of all legs lightly speckled with pale scales anteriorly. Wing.

Pattern of pale spots as figured; base of vein C without pale scale streak on

Sirivanakarn: Subgenus Culex in the Oriental Region 165

posterior surface; first costal pale spot involving only veins C, Sc and Ry; 3rd

costal spot involving apical portions of vein C and Rj; furcation of vein Ro.

largely pale proximally; vein R4,5 with a short pale scale line in basal 0.5; Cu,

with a long pale scale line from basal 0.25 to almost apex; Cu with a row of a

few pale spots proximad of point of furcation; Cuy with a row of 2-5 pale spots

at 0.2-0.8 of total length; 1A with a row of 5 pale spots in middle. Abdomen.

Basal bands on all terga poorly developed or indistinct; tergum II with a small

basal pale spot at middle; terga III-V with very narrow, indistinct basal pale

bands; terga VI, VII without distinct basal bands; tergum VIII with distinct apical

pale band; sterna largely dark, with very poorly defined narrow basal pale

bands on all segments.

MALE, PUPA and LARVA. Unknown.

TYPE-DATA. Holotype °*, Pulau Blakang Mati, SINGAPORE, reared from

pupa taken in a well (a sunken 44 gallon drum), 23 February 1955, D. H. Col-

less (BM).

DISTRIBUTION. Known only from the type-locality in Singapore. Material

examined: 12 (holotype), as indicated in the type data.

TAXONOMIC DISCUSSION. According to Colless (1955: 315), propinquus

is known only from 3 females, all of which came from a single collection at

the type-locality in Singapore. They are strikingly different from the mimulus

and muryvelli in the speckling of the proboscis and legs and in the characteristic

rows of pale spots on veins Cu, Cus and 1A of the wing. None of the adults

among the numerous material of mimulus and murrelli I have examined exhibits

an overlap in these characters with propinquus.

BIONOMICS. The type and 2 other females of propinquus were obtained

from rearing pupae collected in a well, made of a sunken 44 gallon drum

(Colless 1955). No further data are available.

42. CULEX (CULEX) ORIENTALIS EDWARDS

(Figs. 66, 80)

Culex orientalis Edwards 1921: 338 (Co, &).

Culex (Culex) orientalis Edwards, Edwards 1932: 205 (taxonomy); Edwards, in

Barraud 1934: 450 (adult key); LaCasse and Yamaguti 1950: 246 (o'*, 9*,

P*, L*); Hara 1957: 56 (2* genitalia).

FEMALE (Fig. 66). Description based on specimens from Japan and

Korea. Wing: 4.4mm. Forefemur: 2.0 mm. Proboscis: 2.2 mm. Abdo-

men: 3.4mm. Medium or large species; as described and figured by LaCasse

and Yamaguti (1950: 246-50); readily distinguished from the rest of Mimeticus

Subgroup in the wing by the presence of distinct basal costal pale spot which

involves posterior surface of vein C, extreme base of vein R and sometimes M

and by the following additional features. Head. Narrow decumbent scales of

vertex very coarse and largely pale yellowish; erect scales coarse, pale golden

to whitish in center, dark on posterolateral area. Palpus with several pale

scales on apex of segment 4. Proboscis with a broad median pale ring, occupy-

ing 0.25-0.35 of total length; no speckling of pale scales distad or proximad of

median pale ring. Czbarial Armature. Cibarial bar evenly concave, with about

30 delicate teeth, all of which are minute, subequal in size and length and with

abruptly pointed or blunt apex. Thorax. Mesonotal scales very coarse, color

variable from predominantly pale to partially brownish in middle and pale on

marginal areas in anterior 0.7 of disc. Pleural scale patches relatively broad,

166 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

particularly on stp and mep. Legs. Anterior surface of fore- and midfemora

uniformly dark or sometimes with light speckling of pale scales; subapical pale

spots very distinct; anterior surface of hindfemur with distinct longitudinal pale

stripe and light speckling of pale scales among dark areas dorsad and distad of

the stripe; tibiae of all legs with poorly defined longitudinal pale stripes anteri-

orly; all tarsomeres 1-4 with broad basal and apical pale bands. Wing, Pat-

tern of pale spots as figured; extent of spots or streaks variable; basal costal

pale spot near humeral vein usually involving posterior surface of vein C, base

of Rand sometimes also M and Cu; first costal spot at middle of C usually in-

volving C, Sc and Rj, sometimes also M; 2nd costal spot involving C, Sc and

R1; 3rd costal spot usually involving apical portions of C, Ry and Rg, some-

times restricted to C and Ry only; furcation of Ro,3 and M always pale; vein

Ry,- usually with a broad pale scale line extending from basal 0.1 to apical

0. . of length; Cu with or without a short pale scale streak proximad point of

furcation; Cu, with a broad pale scale line from basal 0.25 to apical 0.79 of

length; Cuy usually with pale scale line from 0.5 to apex; 1A with pale scale

line from extreme base to 0.5 or more of total length. Abdomen. Basal pale

bands on terga II-VII usually very broad, sometimes narrow. Genitalia. Ap-

parently indistinguishable from other members of the Mimeticus Subgroup.

Tergum IX with a lateral row of 6,7 setae; postgenital plate rounded on caudal

margin; vaginal sclerite V-shaped and strongly sclerotized; insula with a strong

tuft of 10 setae.

MALE. Essentially as described for female; differing in the presence of

absence of basal costal pale spot of the wing and from other members of the

Mimeticus Subgroup in the following. Head, Palpus very slender and relative-

ly short, usually exceeding proboscis by at most the full length of segment 5,

sometimes shorter; apical 0.1-0.2 of segment 3 with 3-5 weak bristles; seg-

ments 4 and 5 reduced in length, weakly to moderately plumose, all bristles

rather weak and short. Proboscis with or without ventral weak tuft of hairs at

base of median pale ring; apical portion distad of median pale ring largely pale

on dorsal surface; basal portion proximad of median pale ring entirely dark or

with some scattered pale scales on dorsal surface.

MALE GENITALIA (Fig. 80). Strongly differentiated from the rest of the

Mimeticus Subgroup in the modification of basimere, subapical lobe and disti-

mere as in the following. Segment IX. Tergum narrow; tergal lobe poorly

developed, bearing a row of 3,4 setae. Basimere. Abnormally large, about

0.42 mm in length; basal portion strongly swollen; apical portion narrow,

elongate, with a characteristic subapical tuft of numerous strong bristles on

lateral sternal surface; upper lateral tergal surface with numerous long hair-

like setae forming a broad patch laterad of subapical lobe; other strong bristles

confined to lower lateral surface in basal portion. Swubapical lobe. Very broad

and with numerous long hairlike setae associated with specialized seta of proxi-

mal division; proximal and distal divisions widely separated; proximal division

very broad, with a close-set row of 3 rodlike setae (a-c) and 4,5 strong blades

and 1 club-shaped leaf d-f); distal division with a narrow, short, club-shaped

leaflet (¢) and a very long and strong seta (h) distad. Distimere, Very large

and long; basal portion thick, middle portion enlarged or swollen, distal por-

tion tapered into a blunt point; dorsal margin with distinct crest of several

strong spicules extending from basal 0.4 to apex; ventral margin irregular;

dorsal and ventral minute setae present; subapical claw very short and slender,

placed more or less at apex. Phallosome. Generally similar to mimulus;

apical tergal crown of inner division with 4,5 short slender digitiform proces-

ses; outer division with 1,2 strong mesal spines and 1 strong, laterally diver-

Sirivanakarn: Subgenus Culex in the Oriental Region 167

gent sternal spine; lateral basal process broadly rounded. Pyvoctiger., Apical

crown very large, dark, composed of numerous strong spicules; basal sternal

process well developed, moderately thick, long and gently curved sternad;

cercal setae 3.

PUPA. Specimen not available for description.

LARVA (Fig. 80). Head: 0.78 mm. Siphon: 2.0 mm; index 7. Saddle:

0.43 mm; siphon/saddle ratio 5. As figured and described by LaCasse and

Yamaguti (1950: 246-51); strongly resembling typical mzmulus in most features,

differing from it particularly in the following. Abdomen. Setae 1-III-VI 4-6

branched, all subequal, about 0.5 or less of the length of setae 6; 6-III, IV

double or triple. Comb scales 20-40, all small, subequal and with apical fringe

terminating in a strong median spine. Seta 1-X 3,4 branched; 2-X usually

triple (2-4); anal gills 1.5 times as long as saddle. Siphon. Shorter and rela-

tively thicker; distally gradually tapered; siphonal tufts 6-8 pairs (or total 12-

16); 5-7 pairs subventral, placed beyond pecten to about 0.9 of siphon length;

1 pair lateral, placed beyond the middle; subventral pairs 4,5 branched, most

proximal as long as siphonal width at point of attachment, rest gradually short-

er; lateral pairs triple and short; median caudal filament of spiracular appara-

tus absent.

TYPE-DATA. Holotype “*, Tokyo, JAPAN, 5 October 1915, S. Yamada;

(BM).

DISTRIBUTION. Japan and Korea; also reported from the Philippines,

Taiwan, China and USSR. Material examined. 90 specimens: 250", 459, 20 L.

JAPAN. Tokyo (Type-locality); Yokohama; Kyoto; Yodo; Saitama, Sagi-

yamay9¢; 169,°8-L,

KOREA. Seoul; Chinhae; other unspecified localities; 160°, 292, 12 L.

Additional records from the literature: PHILIPPINES: Balabac Island;

Laguna Province, Luzon (Baisas 1938); TAIWAN: Hwalien Hsien (Lien 1962);

USSR: Khaborov; Ussurii; Maritime Province (Monchadskii 1951); CHINA:

Chekiang; Kwangtung; Liaoning (Feng 1938).

TAXONOMIC DISCUSSION. Culex orientalis exhibits a strong affinity with

the members of the mzmulus complex in the pattern of wing spots, the type of

the phallosome and the development of the basal sternal process of the proc-

tiger of the male genitalia and in larval characters. Except for several con-

stant diagnostic features, the ovientalis adults are very variable, particularly

in the extent of the wing spots, color of mesonotal scales and in the length and

the plumosity of segments 4 and 5 of the male palpus. This seems to indicate

that ovientalis is probably a complex of forms which at present cannot be cor-

related in any way with their geographical distribution.

Culex orientalis can be easily recognized in the adults, especially the fe-

male by the presence of a distinct basal pale spot which involves veins C, Ror

also M and Cu of the wing in addition to the other pale costal spots or streaks

as described in all other species of the Mimeticus Subgroup. The male geni-

talia of orientalis are unique, strongly differentiated from the rest of the

Mimeticus Subgroup by the modification of the basimere, distimere and by the

detail of the subapical lobe. The larvae described above, although not definite-

ly associated with the adults apparently agree well with the description and

figure by LaCasse and Yamaguti (1950: 248-9). The orientalis larvae apparent-

ly resemble those of mimulus more than any other members of the Mimeticus

Subgroup.

Culex orientalis is probably restricted to the northeastern Palearctic,

particularly in Japan and Korea and possibly also in north China and USSR.

The previous records of this species from Sri Lanka by Carter (1950), Taiwan

168 Contrib. Amer. Ent. Inst., vol. 12,:no. 2; 1976

by Lien (1962) and from the Philippines by Baisas (1938) are very doubtful and

require confirmation. The females recorded as this species from Balabac

Island and Laguna Province, Luzon in the Philippines (Baisas 1938: 217) are

probably specimens of mimulus judging by the description of this author.

BIONOMICS. According to LaCasse and Yamaguti (1950), ovientalis is

moderately abundant in Japan where it has been found to be more common in

the north than in the south. The most frequent breeding sites include ground

pools, rice fields; natural and artificial ponds, margins of slow moving ditches

and streams. The majority of these collections were made in open sunny habi-

tats, containing duckweed, algae or lily pads at a high altitude in mountainous

regions. The immatures were frequently collected in association with speci-

mens of Culex (Eumelanomyia) hayashii Yamada,’ (Lophoceraomyia) infantulus,

C. (Culex) tritaeniorhynchus, C. (Culex) pseudovishnui and C. (Culex) bitaeni-

orhynchus; Anopheles sinensis Wiedemann and Aedes dorsalis (Meigen). Most

adults from Japan and Korea are labelled as coming from field catches. In

Japan, there is nothing to indicate that this species bites man (LaCasse and

Yamaguti 1950). Nothing is known about adult feeding habits and the disease

relationships of ovientalis.

ACKNOWLEDGMENTS

I wish to express my gratitude to Dr. Botha de Meillon for his guidance,

encouragement and assistance in obtaining the specimens during the early phase

of this study and E. L. Peyton for innumerable favors and continued support.

Iam very much indebted to Ronald A. Ward for critically reviewing and edit-

ing the manuscript and for making further arrangements for obtaining speci-

mens for study. My sincere thanks go to John N. Belkin, Department of Biolo-

ey, University of California, Los Angeles; Donald H. Colless, Division of

Entomology, CSIRO, Canberra, Australia Douglas J. Gould and Rampa Rattana-

rithikul, U. S. Army Medical Component-SEATO, Bangkok, Thailand and

Bruce F. Eldridge, Department of Entomology, Walter Reed Army Institute of

Research for valuable comments and suggestions in improving the manuscript;

Phuangthong Malikul, John Lane, Thomas V. Gaffigan, Anita L. Schmidt,

Carolyn M. Emerick and Susan Hevel for technical assistance; Thelma Ford

Smith, Ann Hoskins Dery, Gloria Gordon and Sorasak Imvitaya (SEATO Lab-

oratory, Bangkok) for preparing the illustrations and Janet D. Rupp and

Owilda J. R. Curtis for typing the manuscript. I am also very grateful to the

following individuals and institutions for their generous support in sending

material on loan or as gifts: Shivaji Ramalingam, Department of Parasitology,

University of Malaya, Kuala Lumpur, Malaysia; Douglas J. Gould and Bruce A.

Harrison, U. S. Army Medical Component-SEATO, Bangkok, Thailand; Peter

F. Mattingly, British Museum (Natural History), London, England; Wallace A.

Steffan, Bernice P. Bishop Museum, Honolulu, Hawaii; Henry S. Dybas, Field

Museum of Natural History, Chicago; California Academy of Sciences; Phila-

delphia Academy of Sciences; Instituut Voor Tropische Hygiene, Amsterdam,

Holland; Lewis T. Nielsen, Department of Entomology, University of Utah,

Salt Lake City, Utah; J. M. Klein, Office de la Recherche Scientifique et Tech-

nique Outre-Mer (ORSTOM), Bondy, France; Ray Parsons, U. S. Army Medi-

cal Research Unit, Kuala Lumpur, Malaysia; B. N. Mohan, Coonoor, Madras,

Rachael Reuben, Vector Control Research Center, Madras, Pondicherry, and

N. L. Kalra, National Institute of Communicable Diseases, Delhi, India;

Richard H. Baker, Pakistan Medical Research Center, University of Maryland,

Sirivanakarn: Subgenus Culex in the Oriental Region 169

Lahore, Pakistan; Ronald Rosenberg, John Hopkins University, Center for

Medical Research and Training, Dacca, Bangladesh; T. Kurihara, Teikyo

University, Tokyo and Yoshito Wada, Nagasaki University, Nagasaki, Japan;

Mario Coluzzi, Instituto di Parassitologia, Universita di Roma, Rome, Italy;

B. M. McIntosh, South African Institute for Medical Research, Johannesburg,

South Africa; F. E. Baisas (deceased), Manila, G. L. Alcasid, National Mu-

seum, Manila, the Philippines; J. C. Lien, U. S. Naval Medical Research Unit

No. 2, Taipei, Te-Lo Ma, Tunghai University, Taichung and P. S. Chen, Tai-

wan Provincial Institute of Infectious Diseases, Taipei, Taiwan, Republic of

China.

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184 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

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192%. From what point of the proboscis of the mosquito do

developed larvae of Filaria bancrofti escape? Sci. Rep. Inst.

Infect. Dis. Tokyo 5: 647-51.

CULEX

fasciatus

gquingue

subapical

lobe

ventral arm

(inner division)

tergal arm

(outer division)

fi f, f

ak f’

EV 4

! Be

a ay FD

fa by) x

Ly oT BE

el ae a

qi: }

7 18 Ns

J: % A

' 4g i

‘A (3;

» Cc

a Ney

exe

WS 1 r ss

BOE) S/F \:

Whe ee J

i ROW A lag \ e \ oh Lod

Mn i Pay tant wre, | yay

ACRES RN Paiste’ | 7 THis dN) Nii

2s | 4 %.

a aa

Gay

44 Oo

‘) Le

Wye

‘ y.

WS:

TTL. HWANG c

basal sternal

process

ventral

arm

(inner

division) 3%

lateral | (outer

SPINE F division)

lateral

basal

process

phallosome

vagans

C By

‘AUG Ady

3) LE Sagi il

£) }

‘

arene SN

InNsOon

hutch

Fig. 6

: A h 23

9 <—IT | ff bere | eee

hutchinsoni

g.8

CULEX

hutchinson

ppn

7 WSs

tergal arm

lateral

spine

lateral

basal

process

proctiger

thei/eri )

subapical crest

gE AN

Aa Tint AY QS

KAW

ANY \\\

tergal arm

ventral

arm

lateral

basal

process

=) —=

proctiger

\ A

Fig.10

theiler/

univittatus

: n

SPRY \

\ ’ Nw s

Fig. 12

ZZ;

y Gy

54 ee

YA 23)

7 IN 4 ee

ne 14 8

4 5 ]

yf iS ee

s ' - 8

f 3 iat

r f 5N edie

: I Or 4

. IX-T

/—— 0 —

fuscocephala

CULEX

fuscocephala

UH \

if; ‘

BAAD

EE As ee

Zo Rr ant h

z, Ah

“

‘ im

Gilweag bios mol

<3 Paya ny

Nae Mer seT}

S87 \

Nera '

Nypwe fabs

[Neuere

hutchinsoni

CULEX

+— 0.05 ——4

ah Wy Ke

tag Fst

" _— ww

/ 14 ap pe ibe

2 A a

AF IN INE May tt

Fie a SOI USO SAS

, wn rey TN WA wy

Yon pr

NUN

an ! "

Rin “nine py gh

ty Blaine

SAAN

i Sgntoe

Bes

Fax

Setar

gon

Yo,

fae.

AVP,

al,

TES

AA.

SS:

Why tat

WV 7 Wi

Wil le

fuscocephala

hitalt iV!

ge at yy

Sar

MH yy Gy

aaa

Nog

s/tiens whitmore/

vishnul

incognitus

{

atiewoaty,

mimeticus

| mimulus

Saeenen |< 2 ae

(oe)

Kec

gelidus

inner division

outer division

0 end

iorhynchus

bitaen

CULEX

apical

sternal

angle

apical tergal

angle

foliform ¥ 74.

processes cap

(outer

lateral division)

basal

process

apical sternal

spiculate portion

(inner division)

WOOLY

‘

WF an

(outer

division)

basal

sternal

process

bree"

bitaeniorhynchus

tenax form

(or pale form)

domesticus form

(or dark form

infula

eo Ie Ese

wee 6 0 00 0 a 0 6 6 ee

luzonensis se/angorensis

Fig. 24

apical margin

apical sternal

angle

apical tergal Oe :

angle ‘ : = 4 SASS S

SES

~SNS

ee Se

foliform

processes

tergal

lobe

0.1

lateral

basal

process

infula /uzonensis

se/angorensis longicornis

Reais pare hal)

Paige mcnacnen thie

nanan epee setae A Hs fi

We at sadn onan diys

PL Ae alana

‘ Digits hye Ve

aC ee 2a ce is

(

S/Ne@NSIS

: ce x

= ae <2 ee 25) ve)

MQX\ : See we)

Sinens!s

cornutus

apical sternal

spiculate portion

Fig. 32

apical fingerlike

processes

sternal

spine

lateral

——basal

os i process

sternal \ a me

process FF y : ee

> MA:

\ »

Ui]

NYE

WAZA

geminus S

Fig. 34

CULEX

ne)

Fg et

614

Joh a

apical sternal

spiculate portion

sternal

sternal—\.._ |

process \S WY 7

subbasal \

process

and, WR Sy ‘oi

Ded ben ot

AV yew Vag bebe /4y > My

NY) ARNT Wa very 4 4 Ms i) yi)

Nv NV yg 0 7 NT Mp yy) )

ANE) \ Bete: VEL: ALA es rei)

pad

a), XT rts ~ aN tT

< WON wane sitiens

CULEX

ps EN SSO

pass. \ SS ae

RANT ¥

Y) 7

\ , , Daie. e.

SS IN \\ a;

\ 3 S

te ae

apical fingerlike processes

on division)

sternal

spine

(outer

division)

SssS5 Frwsaa >

—

whitmore/

a bd

A oe

CULEX

annulirostris

|’ yi

a! i! M

~J-L. HWANG

vishnul

FE LE

(ly i yey

Ap CRUD oo TVAMEANA

PARRA We

CRA p

l p

lene

{

CC a5 a ‘

CULEX

7\ 2

INS NI

~ &E SSWSSS FRB

/ {GY Ae oe e

Vb”

V/ = u

perplexus

1211 9

;

oe mee Te ae :

na |

\ of raph} HL —

NALD

philiopinensis

iIncogn

: \

< oy

F19...57

YY =

SQ \\ Bes

SS ee

—_

= ——

incognitus

Fig. 59

0.1

ag Bt “4 g

EA fel

: . iP ‘ NX Gane iy ; cot is : ‘x

a a | aw

ea :

oA \ a ve nf

Ase 1M) "4

| \ ie Les Wave iy ‘

\ y\A vA VAN |

ees \yavll ails

NY \\\ } Lé

SAA

)

,.V) "ore :

—J-1.. HWANG

tritaeniorhynchus

Fig. 60

CULEX

a MP

\ y) 12

4 12

N 8 \ 2! 3/ J WY,

Sy 7 0 P 10 yp

SS : p Wey AN

. 4 Ri {iB Wh :

a YZ 10 ' Nit

1 M SAQQY

tritaeniorhynchus

midleg hind leg

sternal

spine

edwards/

Mo a \ ad

QQ \\ .

: > —\\

CULEX

—

barrauad/

Ist costal

3rd costal

t/iCUS

mime

Asoni

jac

sor

ides

mimu

‘et:

Ist costal "

iengens

murrel/

propinguus

talis

or/en

SORASAKS IL.

mimeticus

ZORASAK TL,

mimeticus

Fig. 69

— SS

————

engensis

oli

Fig. 70

y at Ye

retrace yy

acts LN ar tha vy nA sae

; Aiea vedas ; aa y

ON RA ert rate fia

RK s\) ae CL tA Ts G

SORASAK TF.

jackson

Fig. /1

)

\\y

Si}

foe)

S0RASAK TL

tseng/

i 4

Mh H

AAs ARI

| } IN

2 AN

Fig. 73

} x v\N \ >

\AAd

LN Ws

pot en

——————

———SS=SSS

phair:

sri WENN u DYER.

\ et NL,

Ne auit swale LEE

aaa UNE LAT ate

ALL perros GRD Ss RE

TETAS hig, mA TAR IIS f QE

SORASAK T,

mimuloides

2nd costal a 3rd costal

costal

lus

mimu

Fig. /6

- | \

! RY i

oral Ue Y / aff

RRR EE MIRA COLT LL 7 ory te 7

PT oY Gi At

mimulus ate oF

date URS IMAC eING icin ed aa ENGL Nae CMPD SUMO ARE |

)

» y

Wy y Wy, 2

KY)

Wi. Le

= A Y

— <=

ARES

orientalis

DISTRIBUTION OF ORIENTAL CULEX (CULEX) SPECIES

APPENDIX A.

uerdorua

UvdULIIO}IPO fr

EXTRA LIMITAL

me 46 4a 4a 4m se 4a 4a 4a 4a de 4

wTwTwewrwN

> | a | a

UBATV I,

suoy SuoH

soutddttiud

eIsouopuy

aro0desuts

BISARTRI

BISAVTRI ‘UTUdg

UIBU}ITA

y |

Pe Sa

je ce eee 9

pee eeeepat I

eee

4a 4m

Py || S

eee.

eee

pel

fo)

puerreyL »

eyue'y 11g

euling

ysoepelsurg

erpu]

UvISTUBYUSIY

ueystyed

Sees

°* a | | | | @@CCe

oo @€@€060600 1°+#+ €8 @

| ee $i @e . 4a 4 aa | ae | mu 4

| a | | a | |

wwvyrwvwwewewr wew*#ow?w #4

4m 4a 46 4. 4

| 41.) selangorensis |) bal dials [a

4k 4

y_| i

4 q (4 yw

IN 4a 4k ae |

| ai | ei | ali | ai | i | a | a | a | a | i, | ln |

tritaenitorhynchus

DIST RIBUTION

whitei

longicornis

cornutus

epidesmus

kinabaluensis

perplexus

alienus

incognitus

barraudi

edwardsi

diengensis

mimulus

murrelli

propinquus

orientalis

bitaeniorhynchus

Sinensis

quinquefasciatus

infula

pseudosinensis

3. hutchinsoni

theileri

5. univittatus

6. fuscocephala

gelidus

luzonensis

24, pseudovishnui

27.. philippinensis

33. mimeticus

37. mimuloides

25.

26.

28.

29,

30.

31.

32,

38.

39.

40.

| 41.

21. whitmorei

22. annulirostris

17. geminus

72.

i128.

14.

18.

| 42.

268 Contrib. Amer, Ent.:Inst., vol: 12, no. 2, 1976

APPENDIX B: CURRENT TAXONOMIC CHANGES

NEW TAXA

REVAODLUCTI STS cia. te a eg aati yk gta aay ts ae cee alae ee aia, SOU a eee 91

BO EGCOT IS ee ee be es se wees ee ee Se gs 81

POCISIG a a a a Pe eee ee es 79

DINOS Se ee a oe see, eee he ee 124

OTTO By MOBY RY: sikcta er ee ie te Bee ON a ee ee ae Er oe eamesan 71

CLES, SYNOMVULY se ia ah eee ls te Gee UR ee a is la se eu es a a 111

euwavosé, Pevalidated (O SHECHIC TANK Go. os eye eG en Be ek 141

JUSCH EL CULUS, SVMONVMIV eo owls BS bs ol es te Ree al hs + Coke. ee eR 150

ROWS). (SVNONVOI Oe. fe a ee wee so PAS 150

MOOLTOTOITS (SVNOOVIY 9. 6 ek ee ts Be a ek ee ge eee we eo ME 100

NOOMETINUS GV AONV AY Se es a, ee ee 157

MEIC SIGH | SUMO G ANY te ee a ge or hein ee hee gs ee, eee 116

OO a a ea we ye himehagias et han a

COVITULUS ek oO a le ea Ue ee RE a ecm anne 87

TUS COME IES cg a ee eS te tee eee 154

EES 7 ASTER PIS SO ee EGR, GOMER a i gy AUR IS Fo eet On SR ne | nan Tal

TIO isha gtr hue Gs camped vel ION sh OR Nigechiars das goign de Rabies a Sg natal 159

OT AO asa go areca ee Aaa teem ei ene ek We ena eg baw a bans 89

EN CGLOVE oh VA ceatias mong 6) anicues ah sree la isle pes hand duds ple Caminita Aig aes "> ee 49

Sirivanakarn: Subgenus Culex in the Oriental Region

269

INDEX

Valid names are set in roman type, synonyms and nonvalid names are

italicized.

Italicized pages refer to the primary treatment of the taxon, arabic

numerals refer to secondary text references and numbers in parentheses refer

to figures.

Acalleomyia 8

adelae 111, 114, 115

Aedes 43, 106, 168

avian 1.2, 71, Te014

ager (Taentorhynchus) 65, 68, 69

Aino virus 134

albinervis 64, 93, 94

albolineatus 30, 33

albopictus, Aedes (Stegomyia) 43

albus 103, 105

alienus 9,14, 20, 23,'26, 57, 110,

119, 122-124, 126, (82, fa

Salis Ly. 2. Wy BO, 2a, 28 Be 2a ay,

95, 100-103, (38, 39)

alphus (Lophoceraomyia) 91

ambiguus (Grabhamia) 71-73

andersoni 43, 47

annadalei (Uranotaenia) 122

annulirostris 3, 9).14;.17,°23, 26,

o7, 95, 99, 107-110, (43, 44)

annulirostris complex 57, 95

annulus 1, 2, 111, 114-116, 133, 134

Anopheles 54, 106, 115, 122, 124,

150, 168

argenteus (Taeniorhynchus) 103, 105

asteliae 36

aurantapex 71

australicus 29, 30

bailyi (Culiciomyia) 94

bancrofti (Wuchereria) 40, 55, 62, 70,

106, 110, 134

barbirostris (Anopheles) 106, 122

Barraudius 8

barrauds 1,.2, 10,10; .10;° 28,026. . 57,

138-141, 142, 143, (63, 64)

Barraudi Subgroup 10, 11, 57, 64,

tes tsar

Belmont virus 110

bivoi 129, 131

bihamatus 58, 109

bitaeniorhynchus 1, 2, 10-12, 18, 21,

25, 57, 64, 65-70, 71-80, 82,

905) 119,122). 158) 168: Crs, 20,

Oh Be)

bitaeniorhynchus complex 57, 64, 74,

75, 77, 79, 80-82, 85

Bitaeniorhynchus series 56, 63

Bitaeniorhynchus Subgroup 9, 10,

06-08, 62-65, 69, 79, 83-85,

88, 90-94, 109

boninensis 9

Brugia 55, 62, 70, 100, 134

cheni 8, 110, 115

Chikungunya virus 36

christophersi 30, 33

cingulatus 30, 33

comitatus 29

confusus 157, 159

cornutus 9, 12. 17. 20.5 246 S764;

86-88, 89, (31)

creticus 43, 45

Culex (genus) 1-3

Culex (subgenus) 1-4, 7-10, 54, 63,

Culiciomyia 8, 94, 122, 137

decens series 47

Dengue virus 134

diengensis, 1, 10,015.18, 22. 24, 57,

144, 156-157 (66, 69)

doleschalli 30

domesticus (Taeniorhynchus) 71, 73,

74, 75

dorsalis (Aedes) 168

edwardsi 1,2, 410; 118) :19,523;° 26;

07, 138, 140, 141-143, (63)

epidesmus 1, 9, 11, 17, 20, 23, 57,

64, 88-89 (31)

Eumelanomyia 8, 9, 122, 137, 168

exilis 37, 39

fasyi 10, 15, 19, 22, 26, 57, 144,

148, 149-150, 152-154, (69)

jfatigans 10, 28-30, 33, 34, 36

fouchowensis 30, 33

fragilis (Culiciomyia) 137

fuscanus (Lutzia) 43

Juscifurcatus 1, 2, 150, 151, 152

270

Jfuscitarsis 51, 53, 54

fuscocephala 9-11, 16, 21, 24, 28, 29,

42, 47, 49, 51-55, 116, 119, 124,

134, (11, 12, 13, 14)

fuscocephalus 51

gelida (Leucomyia) 58

gelida var. bibunctata 58, 60

gelidus 9-11, 13, 17, 22, 25, 36, 54,

56, 57, 58-62, 106, 124, 134, 137,

(15,517,218, 039)

gelidus var. cuneatus 58, 60

Gelidus series 56

Gelidus Subgroup 9, 57, 58, 64, 106

geminus 9, 12, 20, 22, 24, 57, 64,

90-91, 92, 93, (32)

geminus complex 57, 64, 93

elobocoxitus 29, 30

gnophodes 95, 98

goughii 48, 49

Grabhamia 71-73, 88, 89

halifaxii (Lutzia) 137

hayashii (Eumelanomyia) 168

hensemaeon 30, 33

Heptaphlebomyia 4, 48, 49

huangae 8, 28, 29

hutchinsoni 9, 11, 16, 20, 24, 28,

29, 37, 20-23, 68, (6, 7, 8,14)

impellens 95, 98

incognitus 9, 14, 20, 23, 26, 57, 110,

119, 326-129, (16, 56,..57)

inelegans 51, 53

infantulus (Lophoceraomyia) 43, 122,

168

inttla ‘i 2; 9,:13) 18, 21, 22.325,

of, 64, 70, 71-75, 76-78, 80-82,

(23, 24, 25)

Ingwavuma virus 116

iyengari 36

jacksoni 1, 2, 10, 15, 19,.°22,.26,

57, 144, 148, 150-152, 153,

154, 156, (65, 70,. 71)

Japanese encephalitis virus 10, 36,

55, 62, 70, 106, 116, 120, 134

juxtanucleare (Plasmodium) 100

kangi 1, 2, 150-152

kavatsuensis 65, 68, 69

kinabaluensis 1, 2, 9, 14, 19, 21, 24,

ov, 64, 91-94, (33, 34)

Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

kochi (Anopheles) 54

Koongol virus 110

Lastoconops 4

Leucomyia 4, 58, 82, 103, 105

litoralis 95, 100, 102

fongicornis t, 2, 9,:13, 17, 20, 23,

57, 64, 70, 81-82, (24)

Lophoceraomyia 8, 91, 122, 137,

168 )

loricatus 103, 105

luteoabdominalis (Taeniorhynchus)

88, 89 i

luteola 51

Lutzia 8, 43, 137

luzonensis, Aedes (Finlaya) 137

ljugonensis 1, 2, 9, 12,:18,.22, 25,

of, 64, 70, 75-77, 78-80, 82,

(23, 24)

macfarlanei (Uranotaenia) 137

madagascariensis (Pseudoheptaphlebo-

myta) 4

malayi (Brugia) 55, 62, 70, 100, 134

malayi (Eumelanomyia) 122, 137

mammilifer (Lophoceraomyia) 137

mediolineatus (Aedes) 106

microannulatus 95, 98

mimeticus, 10,15, 19, 22, 26, 87,

144, 145-148, 149-156, 158,

(16, 65, 67, 68)

mimeticus complex 57, 138, 144, 149,

152, 154-157, 161

Mimeticus series 56

Mimeticus Subgroup 10, 15, 957, 64,

143, 144, 148, 159, 161, 165-167

mimuloides 10, 15, 19, 22, 25, 57,

143,144. 1434156, 157, (65;°%3,

74)

minus 2) 2.:00.8 10, 19, 22:23. 27,

97, 94, 119, 144, 155, 157-161,

162-168; (16, 75; 16, 7%)

mimulus complex 57, 144, 155, 157

minimus 51, 53

minor 30, 33

miraculosus 36, 39

montforti (Heptaphlebomyia) 48, 49

mossmani 157, 158

Murray Valley encephalitis 110

murrelli 10, 15, 19, 23, 27, 57, 144,

161-164, 165, (66, 78, 79)

neavei 50

Sirivanakarn: Subgenus Culex in the Oriental Region

neolitoralis 1, 2, 100-102

neomimulus 1, 2, 157, 160, 161

neovishnui 1, 2, 116, 118, 119

nigerrimus (Anopheles) 54

nigvicephala 95, 98

nigropunctatus (Culiciomyia) 137

nilgiricus 9

ochracea (Grabhamia) 88, 89

Oculeomyia 4, 71, 73

omani 110, 128

onderstepoortensis 43, 46

orientalis 2, 10,'15,°28,:20. 23, 67,

144, 165-168, (80)

pacificus 36

pallidothorax (Culiciomyia) 122, 137

pallidocephala 48, 49

palmi 107-109

permixtus 8, 28, 110, 115

perplexus 9, 12, 14, .20, 22, 269557,

190, 119,.720-122, 123,°%24. 126

128, (50, 51)

perexiguus 48, 49

pervigilans 36

pervigilans complex 37

pettigrewti 43, 45 :

philippinensis (Anopheles) 106

philippinensis 1, 2, 9, 14, 23, 26, 57,

1t0, 119, 122-126, 128, (54, 95)

pipiens 4, 29, 30, 34, 35, 42

pipiens molestus 29, 35

pipiens pallens 28, 29, 35

pipiens complex 9

Pipiens Group 1,9, 11, 16,20, 24.

27, 28, 37, 41, 46, 48, 50, 51,

Pipiens Subgroup 29, 30, 34

Plasmodium 70, 100

plegepennis (Leucomyia) 103, 105

poicilipes (Lasioconops) 4

propinquus 1, 10, 15, 16, 20: 23. 57,

144, 161, 164-165, (66)

Pseudoheptaphlebomyia 4

pseudoinfula 111, 114, 116

pseudomimeticus 145, 147

peeudosinensis 9, 12, 18, 22, 25, 57;

64, 70, 74, 78, 79-81, 82, 90,

OL 26. 27)

pseudovishnui 1, 2, 9, 10, 14, 20, 23,

29, 94, 57, 106, 110, 115, 716-

a0, 421-120, 130. 139, : 134,

168, (48, 49)

2th

quadripalpis (Lophoceraomyia) 122

quasipipiens 30, 33

quinquefasciatus 9-11, 16, 21, 24, 28,

29, 30-36, 37-39, 42, 43, 45, 46,

49, 94, 98, 362," 0P):2,.3. 14)

reesi 30, 33

roseni 95, 102

rotoruae 36

rubithoracis (Lophoceraomyia) 43

salinus 96, 98

savawaki (Oculeomyia) 4, 71, 73

scanloni (Culiciomyia) 122 «

selangorénsis §, 2,°9, 12, 18, 21,

29, 97, 64, 70, 74, 77-79, 80,

81, (23, 24)

sepositus 82, 84

sericeus 30, 33

simplex (Heptaphlebomyia) 4, 48, 49

Simpsoni 43, 47

Sindbis virus 47, 50, 70, 110, 116

sinensis (Anopheles) 168

sinensis o, 12.° a, 21, 24: 95, 57,

63, 64, 82-66, 87,° (15, 28,29.

30)

Sinensis complex 57, 64

sitiens. 9, 10; 24. Ge; 22, .25, 37,

Ga 100, “TOL We, LOT; 208. F228.

187, tee, (VS, 35, 36..23)

sitiens complex 57, 95, 102, 109

piens Group S$, 7, 9, 11, 16. 17. 20,

21, 24, 28, 55-58, 61-64, 99,

$00, TCS. Tia. bls: deo, Tad, tad.

146

Sitiens series 56

Sitiens Subgroup 57, 58, 64, 93-95,

103, 104, 106, 110, 143, 146

solitarius 144, 150

squamosus 64, 93

starckeae 94, 109

subpictus (Anopheles) 54

Summorosus 129, 131, 133

summorosus subgroup (Baisas) 137

taeniarostris (Grabhamia) 71-73

Taeniorhynchus 65, 71-73, 82, 84,

88, 89,.103, 105

tamsi 36

taytayensis 91, 53

Tembusu virus 62, 116, 120, 134

tenax (Taeniorhynchus) 71-75, 81, 82

272 Contrib. Amer. Ent. Inst., vol. 12, no. 2, 1976

tenax var. ocellata 71, 73

thalassius 99

theileri 11, 16, 21, 24, 28, 29, 39,

43-47, (9, 10)

_ theilevi var. annulata 46

Theileri Subgroup 9, 29

Theobaldiomyia 4, 58 |

tipuliformis 37, 39, 43, 46

torrentium 36, 39

trifilatus 36

Trifilatus Subgroup 9, 29, 36, 37, 42

tripunctatus 82, 85

tritaeniorhynchus 9, 10, 14, 20, 23,

26, 54°56, 357; 62,206, 110,

£155 116) 189,2120,0 124, 128,

129-134, 137, 168 .

tritaeniorhynchus var. siamensis 129,

131

tritaeniorhynchus summorosus 129

tritaeniorhynchus complex 57, 110,

111

tsengi 10, 15, 19, 22, 26, 57, 144,

148, 15029152-753;:465, 12)

Umbre virus 116, 120

uniformis 51, 53

univittatus 9, 11, 16,021; 24, 28,

29, 47, 48-51, (9, 10)

Univittatus Subgroup 9, 47

Uranotaenia 122, 137

vacans 9, 1h) 10)) 21° 24, 29.436,

37-40, 39-41, 152, (4, 5, 14)

vagus (Anopheles) 54, 106

variatus (Lophoceraomyia) 137

vicinus 58, 109

virgatipes 37, 39

vishnui 1,2; 9,10, .13,' 14; 220;423;,

20, 54, 57, 106, 110, 111-116,

£179421, 1285::125;, 6127, 190;:133,

134, 135, 158, (16, 45, 46, 47)

vishnui complex 57, 110, 111, 118,

116, 118, 120-128, 130; 131,..133,

189, .287,°158);161

vishnui group 56

Vishnui Subgroup 2, 9, 10, 56, 57, 64,

90, 93, 111, 133, 136, 138-140,

143, 146, 147, 155

vivax (Plasmodium) 70

West Nile virus 47, 50

whitei 9, 12,14, 18,;.22, .26,;67;:. 111,

135-138, 139, 140, 142, 143, (61,

62)

whitei complex 57, 110, 111

whitmorei 9, 13, 18, 23, 25, 57, 58,

61, 95, 103-106, (15, 40, 41, 42)

whitmorei complex 57, 95

whittingtoni 95, 102

Wuchereria 40, 55, 62, 70, 106, 110,

134

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