L f ^-l R A R >

^l^ 'mx^

• DEC2 2 59SO ,

A Handbook of tnV

MOSQUITOES

of the

SOUTHEASTERN

UNITED STATES

W. V. King

G. H. Bradley

Carroll N. Smith

and

W. C. MeDuffle

Agriculture Handbook No. 173

Agricultural Research Service

UNITED STATES DEPARTMENT OF AGRICULTURE

\ I
PRECAUTIONS WITH INSECTICIDES

All insecticides are potentially hazardous to fish

or other aqpiatic organisms, wildlife, domestic animals, and man. The dosages needed for mosquito

control are generally lower than for most other insect

control, but caution should be exercised in their

application. Do not apply amounts in excess of the

dosage recommended for each specific use.

In applying even small amounts of oil-insecticide

sprays to water, consider that wind and wave action

may shift the film with consequent damage to aquatic

life at another location. Heavy applications of insecticides to ground areas such as in pretreatment situations, may cause harm to fish and wildlife in streams,

ponds, and lakes during runoff due to heavy rains.

Avoid contamination of pastures and livestock with

insecticides in order to prevent residues in meat and

milk.

Operators should avoid repeated or prolonged

contact of insecticides with the skin. Insecticide concentrates may be particularly hazardous. Wash off

any insecticide spilled on the skin using soap and

water. If any is spilled on clothing, change immediately. Store insecticides in a safe place out of reach

of children or animals. Dispose of empty insecticide

containers. Always read and observe instructions

and precautions given on the label of the product.
UNITED STATES DEPARTMENT OF AGRICULTURE

Agriculture Handbook No. 773

A Handbook of the

MOSQUITOES

of the

SOUTHEASTERN

UNITED STATES

By

W. V. King

G. H. Bradley

Carroll N. Smith

and W. C. McDuffie

This Handbook replaces Miscellaneous Publication No.

336, The Mosquitoes of the Southeastern States, but

contains information that originated in that publication.

Washington, D. C. Issued November 1960

For sale by the Superintendent of Documents, U. S. Government Printing Office,

Washington 25, D. C. Price 75 cents
4/^^^b^v^rt^wvv^iru^ffinArtf^^i¿Hffi^^^iJVVV^^'vv^

PRECAUTIONS WITH INSECTICIDES

All insecticides are potentially hazardous to fish

or other aquatic organisms, wildlife, domestic aninmls, and man. The dosages needed for mosquito

control are generally lower than for most other insect

control, but caution should be exercised in their

application. Do not apply amounts in excess of the

dosage reconunended for each specific use*

In applying even small amounts of oil-insecticide

sprays to water, consider that wind and wave action

may shift the film with consequent damage to aquatic

life at another location. Heavy applications of insecticides to ground areas such as in pretreatment situations, may cause harm to fish and wildlife in streams,

ponds, and lakes during runoff due to heavy rains.

Avoid contamination of pastures and livestock with

insecticides in order to prevent residues in meat and

milk.

Operators should avoid repeated or prolonged

contact of insecticides with the skin. Insecticide concentrates may be particularly hazardous. Wash off

any insecticide spilled on the skin using soap and

water. If any is spilled on clothing, change inunediately. Store insecticides in a safe place out of reach

of children or animals. Dispose of empty insecticide

containers. Always read and observe instructions

and precautions given on the label of the product.
'!•»

UNITED STATES DEPARTMENT OF AGRICULTURE

Agriculture Handbook No. 773

A Handbook of the

MOSQUITOES

of the

SOUTHEASTERN

UNITED STATES

By

W. V. King

G. H. Bradley

Carroll N. Smith

and W. C. McDuffie

This Handbook replaces Miscellaneous Publication No.

336, The Mosquitoes of the Southeastern States, but

contains information that originated in that publication.

Washington, D. C. Issued November 1960

For sale by the Superintendent of Documents, U. S. Government Printing Office,

Washington 25, D. C. Price 75 cents
CONTENTS

page

Genera and species found in the Southeastern States

Literature on mosquitoes

General characteristics and habits of mosquitoes °

Collection and preservation of material -'-*

Mosquitoes and disease * '

Mosquito control ^^

Mosquito surveys ^^

Engineering surveys ^ *

Control of mosquito larvae 28

Control of adult mosquitoes 33

Insecticide dispersal equipment 36

Mosquito repellents 37

Specific problems 39

Mosquito identification and notes on the genera and species . 48

Key to subfamilies and genera 55

Genus Aedes Meigen 59

Genus Anopheles Meigen 85

Genus Cidex Linnaeus 103

Genus Culisefa Felt 124

Genus Deinocerites Theobald 126

Genus Mansonia Blanchard 127

Genus Orthopodomijia Theobald 134

Genus Psorophora Robineau-Desvoidy 137

Genus Toxorhynchites Theobald 149

Genus Uranotaenia Lynch-Arribalzaga 151

Genus Wyeomyia Theobald 153

Literature cited I57

Index 185

Grateful acknowledgment is made of the assistance of the

following people in preparing this publication : A. W. Lindquist,

J. A. Fluno, Alan Stone, A. D. Cushman, Lola J. Roos, Delia Sims,

and Lucy Ford.
A HANDBOOK OF THE MOSQUITOES

OF THE SOUTHEASTERN UNITED STATES

By W. V. King, G. H. Bradley,i Carroll N. Smith, and W. C. McDuffie,

entomologists, Entomology Research Division, Agricultural Research Service

This publication deals with the mosquitoes recorded from the

nine States east of Texas and Oklahoma and south of the latitude

of the Virginia-North Carolina border, bringing together information that is widely scattered through the literature, and providing

a convenient means of identification. It constitutes an extensive

revision of a previous publication (193), the third and last edition

of which was issued in 1944. As in that publication, it contains

notes, brought up to date, on the habits of the species, their distribution, economic importance, and methods of collection and

control, with brief descriptions of the species and keys for the

identification of adults and larvae. Considerable material has

been added, particularly on methods of control and on bionomics.

Because fewer species are considered than in the more general

reference works, an effort has been made to simplify the identification of adults by using for generic separation principally those

characters that can most readily be seen.

GENERA AND SPECIES FOUND IN THE SOUTHEASTERN

STATES

The mosquitoes found in the Southeastern States comprise a

fairly natural group as most of the species in this country are

either restricted to the South, or reach their greatest abundance there. The general distribution and status as to prevalence

and economic importance are shown in table 1. The information

has been compiled from records in the literature and from collections of the present writers.

The following species are listed as being of economic importance :

Anopheles quadrimaculatus, the common malaria mosquito,

transmits malaria and is an annoying pest as well.

Aedes aegyptiy the yellow-fever mosquito, is a vector of yellow

fever and dengue fever, as well as a troublesome house pest.

Culex pipiens quinquéfasciatus, the southern house mosquito,

is a serious house pest. It transmits bird malaria and is an intermediate host of some of the filariae.

1 Now vsrith the United States Public Health Service.
AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

-2

"co

Ö

o

CO

•


CO

o

CO

> o^ o o

oj c ö ac

^ a; cd c cd

Cd

i:f o

^c3

^-t^Tt'-t^CI-t^-^^'^-^^'-t^^^'-t^Cl'I^Cl

++++++ ;++ :++ ;++ ' + •++ +++ :++

"Z cd

O

c

O)

cd

C

cd

*55

'Bo

Ç2 CU

cd

Bcd

cd

cd

'So(-<

o

a> O

cd

o

T3tí

Cd

w

í3tí

a;

O

++++ ' -{- •++ ' ++ ' -{--\- •+ :++ ' +++ •++

+ ++ :+ ;+++++ ::+:::+. .+++ . .+

-f +++ ' + :+-f++4- '++ :+ : + + ;+++ • •>

++ •+ :+++++++ •++ ' + •++ :-f4-+ ++

++ •+ :++- •++++ '+-\- ' -{- :+ :-f+

++ ;+++ ;++ ;++ ++ :+ :++ :+ :+ :++

:++ :-f+ ;++ ;+ :++ :+++ :++

++ :+++ •++ :+++ + + + ':+ +++

- í ío w

^ ~ ^ í£ ce

í^~ S'^'^

<;ü -»~ -1^ C Ci

S-i¿ ~ c ^ :=:

Çk.^ ^ co^ g ^'^

í_- "^ "^ v¿_^ ö5-~w í: ce ce ce ce-^'^'^

^ O 'íi^ 5;^ ?: ce . Ö ce
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 3

^ tí Ö ftß

ÍH^ Co g Co

O

3

T3

O

í3 o

^(3

cd

^•5

2

'S)

o

o

o

T3tí

cö

CO

tí

O»

o

^COTÍ<TÍ<Tt'«^T-HTÍ< '^'^':0'«*Tj<C0TÍ<Ttr^^,-HCOCl^CO co^

+++ • :+++ •+ • •+ •++++++++ ++

++++ :+++ ;+ : :+ •++++++ :+ ++

•+ • ++++ + •+ • •+ •+ :++++++ ++

•+ • :+::::+ ++

++4- :++++ :+ • + •++ •+++++ ++

+++ •++++ :-h • :+ •++++++++ -h+

+++ - •+++ ' + • '+ :++++++++ -h+

++++ :+++ '-{- ' ;+ :-f-++++++-f -h+

++++ •+++ +++++++++ ;+++++ ++

CO c/5 s;

^ .- '^ ^ ,^ i=x, ^ '

-^ ?» 2 c Tí o-::^-;

^ Ö := o r- ,^ ~ : -^ ?; t- ^ ^¿S.;

o

CO

CO

- s «2

:=: § • ií se ^ ^-^ r^

r- ^ c- í^---^ i^ CO ^ o o í^-v:;-~ eo ~

• .-^ G íl o se'-S ^2^'p; ^ /^ <5L; Ö Ö ^

• •Se
AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

o

O

Xi

xîè'è 0) Ö » ac -^ Cl ^^ TfTj- Olí—l'«d^'^'M'^'^"«^'rJH'rf<'ri<C -t^^

£^^S|

aJ

^ ß

î:! O

o t^ ;+ ;-f- + ++++++ : : : : + ++

GO c^ O

a

r^ tí

; + ++ +++++++ ; ; ; ;+ ;+

:z; Í3o

<D

en

tí ;+ ++ +++++++ ; ; ;++ ;+

tí

0)

H

M

Cd

w

1 g :+ :; + +++++++ ;+ ;++ ¡ +

tí -^t->

T3 <

X3 a

O ß

ai .2

:+ : ++ +++++++ ;+ ; ;+

J

^;-l

;3 1 o (/> .^

o

o

•55 a

;+ ++ ++++ + + + : : : : + : +

Cd

Scd ; + +4- +++++++ ; ; ; ;+ ;+ cd

<

rt

+ + ;-f- +++++++ ; ; ; ;+ ++ a>

a

cd

2

Q 4 + + + -4 +4-++++++ ;+ ;+ ++ I-^

Pin

;.îo

w • "§ .2á ;.o

û) • vi a C/j -^

w

T3tí

Cd

=2 ^

ce,. 'S. tí c/- g • ^

• ce ."^

3

g

2.

}isonia:

indubit

perturbtitilla ïu

•iSi

rophora:

ciliata.

confinn

cyanesc

discolor

fp.rox

hórrida

Howard

johnsto

longipa

pijgmaesignipe

Í III

o ~ '^

t; o %i

1

^

^ o ¿: Í
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES

T3

a»

.S

O

O

es -gés Í c S 9^0 ^00 ^^^

£^ .§5

aJ

^ Ö

o tH ++ + :

CO

xn CÖ

O

CJ

rtí Ö

-tJ iri

ë2 ++ + • to ^ 53 o

(D

:+ : :

CD

Ö

<D

EH

w

CO

1 cö ++ : :

tí ^;-i

Td <ïÎ

0)

'So

.2 ^ 'w ++ ; CO

0)

a

*3

o

QÍ S; ^

:3 • o W .„H

o |â ++ : :

; CO

^•s

OJ

sCÖ

.û ++ + •

<A

<

OJ

•fTn Ol)

o ++ ' '

0)

O

Q ++ +++ 5

E

00

a> •1—t

a

0)

a

m

nd Ö

tí

Ö

•S 31 03

tí ïg

•
 L; Si ^ i3

O

<D 1 S g

g Ö g ö

5s>

^ 1

O

<D

O

o

n3

tt-i

O

+j

i3

O

>>

a

a

o

tí^

a

bû

c

>»

o

títí

03

T3tí

Cd

4JtíCÖ

T3 ,

tí tí

;=í O

^ -í3

oí tí

>-

-Q

03 4J

r )

o-ö

II 73

O)

CM -M

W

O»

ç; <i;

o;

>.

u

ü

0)

>

o o

tí «M

o o

a

a> <D

4J a

tí

2 >>

í~l r—< O 03

o^?.
6 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Aedes sollicitans, the salt-marsh mosquito, is the most important

salt-marsh species generally in the Eastern and Southern States.

Aedes taeniorhynchus, the small, black, salt-marsh mosquito,

is another salt-marsh species of economic importance, especially

in Florida.

Psorophora confinnis, the Florida glades mosquito, is an important fresh-water species in southern Florida, and is troublesome as well in other areas.

Mansonia perturhans, the common Mansonia, is a severe pest m

areas where suitable breeding conditions occur.

Most of these species are known to be capable also of transmitting other diseases, especially encephalitis.

The woods mosquitoes, taken collectively, are also serious pests

of man and animals. The principal ones in the Southeast are

Aedes triseriatus, A. infirmatus, A. atlantictts, A. vexans, Psorophora ferox, and P. ciliata. Cidex salinariiis is important at times,

and various other species, such as P. cyanescens, P. varipes, A.

canadensis, and Mansonia induhitans, become very annoying in

restricted localities.

Thirteen species and subspecies have been added to the list

published by King et al, (193) and one, (Wyeomyia smithii), has

been dropped as a misidentification. This brings the net total of

mosquitoes definitely known to occur in the Southeast to 77. Among

the different States the greatest increase in recorded species has

occurred in Florida, where 16 have been added. Of the present

total of 67 in that State, 15 are tropical species, only one of which

has been recorded elsewhere in the Southeast. Five of the Florida

species are known at present only from the Keys, and one of

these, Culex midrennani, described as new in 1948 (18) y is known

only from that location. The other new species, Wyeomyia haynei

and Psorophora longipalpis, and one subspecies, Aedes canadensis

mathesoni, have been described from other parts of the Southeast. Anopheles albimanus has apparently become established on

the Florida Keys, and another snecies of Anopheles, A. perplexens,

which was previously considered to be a variant of A, punctipennis,

is now listed as a distinct species. Eight species are recorded in

Georgia, Alabama, and Mississippi which have not yet been found

in Florida. One species is listed only in Louisiana and Arkansas

and another, only in Arkansas and Tennessee.

LITERATURE ON MOSQUITOES

The literature on mosquitoes is exceedingly large, and the

articles have appeared in widely scattered publications. General

reference works covering the mosquitoes of the United States

are, however, comparatively limited. Of the references that include

the southeastern species, the four-volume monograph by Howard

Dyar, and Knab (163) contains detailed descriptions, a large

number of illustrations, and much information on bionomics and

distribution of the species of the western hemisphere. The systematic part of this work was later extensively revised and condensed by Dyar (96).

Matheson's handbook (213) (revised in 1944) has long been

a standard text and reference work on the mosquitoes of North

Am.erica. It contains brief descriptions of the genera and species
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 7

and keys for their identification, numerous anatomical illustrations,

an explanation of the taxonomic terms in use, and condensed

accounts of mosquito biology, their relation to human welfare,

the problem of mosquito reduction, and instructions for their

collection and study. A small volume by Carpenter et aZ. (69) on

the mosquitoes of the southern United States covers the same

area as that termed the Southeastern States in the present and

preceding pubUcations. A more recent volume by Carpenter and

LaCasse (68) features full-page illustrations by Japanese artists

of the females of most of the species of North America, north of

Mexico. These illustrations were originally published by Yamaguti

and LaCasse (335) in four processed volumes, and have appeared

also in a small volume published privately by Yamaguti {33J^).

Bates (20) has pubhshed a volume on the natural history of

mosquitoes, and Horsfall (160) has summarized a considerable

amount of the world literature on the bionomics of mosquitoes.

Edward's valuable catalogue of the mosquitoes of the world (102)

is now out of date, but is still a standard reference on nomenclature and classification. A Synoptic Catalog of the Mosquitoes

of the World, by Alan Stone, K. L. Knight, and Helle Starcke,

(298a) was published in 1959 by the Entomological Society of

America (Thomas Say Foundation). Agriculture Handbook No.

182 by Gjullin et ah (126a) describes the mosquitoes of Alaska.

Publications of the New Jersey Agricultural Experiment Station

on the mosquitoes of that State (U2; 292) have been of use to

southern workers as they contain information on a number of

species that occur in the South. The early report of John B. Smith

(292) contains a great many original observations on mosquito

bionomics. This report has been reprinted by Headlee (J-í-^)-

A discussion of the bionomics and ecology of Nearctic Anopheles

has been published by Bradley and King (^9), and their distribution and classification were given in articles by King and Bradley

(190 191), Two works by Boyd (3Jf, 36) on malariology contain

much information on the natural history of anophelines and their

relation to the transmission of malaria. Herms and ^"^^^ y/f\

deal with practical phases of mosquito control, and Covell (81)

published a review of the literature prior to 1931 on the control

of Anopheles. Two short papers on engineering aspects of mosquito control have been issued, one by the National Malaria Committee (233) and another by the Engineering News-Record (lOJ,),

A monograph of the mosquitoes of medical importance

throughout the world, profusely illustrated with the diagnostic

characters, has been published by Foote and Cook (lUa). Two

SSl Smphlets have been issued by the American Mosquito

Sontrol S one on the use of aircraft (^) and one on

nd eauipment and insecticides for mosquito control (3),

^J^?^t¿S^7v^rtm,r.t of Agriculture Circular 17^.}^^0) .or.

takis much information on the chemistry of msecticides and reSnS formulations, dosages, and equipment employed m mosSo contîd The results of laboratory screening tests on more

tZnlO^SoO organic insecticides were reported m tje United State

Denartment of Agriculture Agricultural Handbook 69 U^^) This

SStests of mosquito larvicides, adult sprays, and repellents.

Serial puSiStion contain numerous original articles on

mosquftoe'^TiS^^ Mosquito News, Proceedings of the New Jersey
8 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Mosquito Extermination Association, Proceedings of the Florida

Anti-Mosquito Association, Proceedings of the California Mosquito-Control Association, Proceedings and Symposia of the

National Malaria Committee (discontinued in 1941), Journal of

the National Malaria Society (1942 to 1951), and Insecutor

Inscitiae Menstruus (discontinued in 1926). Articles on mosquitoes

appear also in the Public Health Service Reports and Bulletins,

American Journal of Tropical Medicine, and its successor, the

American Journal of Tropical Medicine and Hygiene, Proceedings

of the Washington Entomological Society, Journal of Economic

Entomology, and various other entomological and medical journals.

The Review of Applied Entomology, Series B, Medical and

Veterinary, is highly useful for keeping informed on the widely

scattered mosquito literature of the world. Mosquito News contains a monthly list of titles of current mosquito literature which

is of special value because the titles appear for the most part soon

after the articles are published.

GENERAL CHARACTERISTICS AND HABITS

OF MOSQUITOES

Mosquitoes are small two-winged flies belonging to the Order

Diptera, family Culicidae. This family was formerly considered

to include related groups but is now limited to the true mosquitoes

{298), The wings, legs, and other parts of the body are more or

less covered with scales, and the mouth parts are produced into

an elongate proboscis, which is employed for piercing and bloodsucking by the females of most species. The males do not suck

blood. The males can usually be distinguished from the females

by their bushy antennae and by differences in the length or shape

of the palpi (fig. 1). In some species the male palpi and antennae

M & A 14045

FIGURE 1.—Heads and appendages of mosquitoes: A, Side view of Anovhele<^

female; o, occiput; ft, frontal tuft; 1-5, palpal segments. B, Culex femalp

(from above). C, Culex male. D, Anopheles male. "

are similar to those of the female. The size of different species of

mosquitoes varies considerably (fig. 2).

There are four stages in the life cycle of a mosquito—the egg
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES

1 ^ \

\

M & A 14046 A

FIGURE 2.—Variations in size of mosquitoes: A, Toxorhynchites rutilus;

By Psorophora ciliata; C, Anoi^heles quadrhnacitlatits; D, Aedes atlanticus;

E, Culex p. quinquéfasciatus; F, C. erraticiLS; G, Uranotaenia lowiL

the larva (often called wiggler or wiggletail), the pupa or tumbler,

and the adult winged insect or imago. The eggs mature in batches

of 50 or less to 200 or more, and several such batches may be laid

by one female. Among the bloodsucking species a blood meal

is usually necessary for the production of eggs. When ovipositing,

some species glue the eggs together into a raft or boat-shaped

mass (fig. 3, A) w^hich floats on the water; other species deposit

the eggs singly on the water ; and still others oviposit on the sides

of containers above the water line, on the soil at the edge of the

water, or in moist depressions (fig. 3, B). The eggs of Anopheles

(fig. 3, C) have lateral structures that keep them afloat. The

incubation period is short in warm weather (usually 2 or 3 days),

but in certain species, particularly Aedes and Psorophora^ the

eggs are able to withstand long periods of drying; in fact, they

appear to require a certain amount of drying, and sometimes

exposure to cold, before they will hatch.

The larvae of all mosquitoes are aquatic and most of them freeswimming. Although possessing trachéal gills, the larvae of most

species must come to the surface for air, and an elongated air tube

or other modified apparatus is provided for obtaining air through

the surface film. During the period of development, which lasts

4 to 10 or more days, the larval skin is shed four times, each successive instar showing a progressive increase in size. The first two

instars are very small and easily recognized as immature. In the

third instar the hairs have fewer branches than in the fourth
10 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

M & A 12937

FIGURE 3.—Eggs of mosquitoes: A, Egg raft of Culex restuans; B, egg of

Aedes taeniorhynchus; C, egg of Anopheles quadriviaculatus, showingfloats. (Howard, Dyar, and Knab.)

instar, and the sclerotization of the anal segment is less complete.

Immature Anopheles larvae usually have a collar of dark sclerotin

around the base of the head.

The food of mosquito larvae consists of minute plants and

animals and fragments of organic debris, which the larvae strain

from the water by the action of their mouth parts. Barber (.9, 10)

reared the larvae on pure cultures of various organisms, and concluded that the presence of living food organisms was necessary

for any considerable growth. Hinman {1A9) has suggested that

materials in solution and colloids in suspension in the breeding

M & A 14047

FIGURE 4.—Pupa of Culex pipiens. (Howard, Dyar, and Knab.)
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 11

waters may play a part in larval nutrition. A discussion of the

food of anopheline larvae is given in the notes on Anopheles quadrimacidatus.

With the fourth molt the pupa appears. The pupal stage (fig. 4)

is also aquatic and is a period of marked transformation, during

which the adult insect is formed. The imago usually emerges

after about 2 days.

The length of life of adult mosquitoes under natural conditions

is difficult to determine, but for most of the southern species it

is probably only a few weeks during the summer months. Some

of the northern species of Aedes that emerge early in the spring

apparently live much longer. Daily observations on abundance

following the emergence of a large brood of certain species of

Anopheles (329) and Aedes have shown a marked reduction in

numbers within 2 weeks. The southern house mosquito probably

lives longer than this, and the yellow-fever mosquito may live,

on an average, a month or more, with a maximum of several

months.

In the North the females of Culex, Anopheles, and some other

mosquitoes hibernate. True hibernation of Anopheles apparently

does not occur in the extreme South, as the females become active

during warm periods and larvae are found in the breeding places

{6, H, IfO). The same is true of some of the culicines in the

warmer sections {91, 150). Aedes and Psorophora pass the winter

in the egg stage, although some winter development of A. sollicifans occurs along the South Atlantic and Gulf Coasts.

The piercing organs of the female mosquito consist of^ six

elongated parts enclosed in a flexible sheath called the labium.

When the mouth parts are inserted in the skin for bloodsucking,

the sheath is bent backward in the middle like a bow. There are

two pairs of slender cutting organs, the mandibles and the

maxillae, and two additional organs called the hypopharynx and

the labrum-epipharynx. The latter is channeled, and the last two

organs, when pressed together, form a tube through which blood

and other liquids are drawn. A very small separate duct is found

in a ventral thickening of the hypopharynx, through which is

injected the secretion from the salivary glands. This salivary

secretion is responsible for the itching sensation caused by

mosquito bites. Not all species of mosquitoes have bloodsucking

females. In the genus Toxorhynchites the proboscis of the female

is not adapted to piercing, and some of the species in other genera

are not known to take blood meals.

The mouth parts of the male are not adapted for piercmg.

The males probably subsist on the nectar of flowers and fruit

juices. Both the males and the females can be kept alive in the

laboratory for considerable periods on fruit juices or sirups.

COLLECTION AND PRESERVATION OF MATERIAL

Anopheles larvae are usually found at the surface of the water

among aquatic vegetation or floating debris and may be collected

by skimming through such material with a dipper or pan. A

white-enameled dipper, having the hand.e lengthened by the

insertion of a cane or smooth stick, makes a convenient imple-
12 AGR. HANDBOOK 173, U. S. DEFT. OF AGRICULTURE

M & A 14076

FIGURE 5.—Equipment for collecting mosquitoes. Larvae: A, dipper; B

vial; C, wide-mouth pipette; D, jar. Adults: E, Killing bottle; F, paper

funnel; G, live tube; H, aspirator, straight-tube type; 7, lantern chimney

cage; J, paper pill boxes; K and L, aspirator and details of construction.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 13

ment for collecting larvae. Around emergent vegetation or logs

the larvae may be drawn into the dipper by submerging one

edge so that the v^ater flows in rapidly as the dipper nears the

obstruction. The larvae may be removed from the dipper to the

collecting jar with a spoon or a large-mouthed pipette provided

with a rubber nipple (ñg. 5). Wide-mouthed bottles (2 to 6

ounces) make convenient collecting jars.

Uranotaenia larvae and certain species of Culex, especially

C. erraticus, are taken frequently with anophelines. Many other

mosquito larvae, however, particularly those of Aedes and PsoroVhora, are more active and usually drop to the bottom of the

pool as soon as disturbed. A quick plunge of the dipper is required to intercept these larvae, or they may be collected by

sweeping through the water with a cloth collecting net or a finemeshed wire strainer. Other kinds of mosquitoes, such as Mansonia, Wyeomyia, or the tree-hole breeders, require special

techniques depending upon the character of the breeding places.

As soon as a collection is made, the jar should be numbered and

a record kept of the locality, date, and conditions under which

the larvae were found.

The larvae and pupae may be kept alive for rearing, or the

large larvae (fourth instars) may be preserved for identification

in 70- to 80-percent alcohol or 10-percent commercial formalin.

About 1 percent of glycerin should be added if the vials are to

be stored. Specimens retain their form best if killed in hot water

(not over 150° F.).

Permanent slide mounts of larvae and male genitalia should

be made with one of the resins such as balsam or euparol. In the

U. S. National Museum, Division of Insects, permanent mounts

are made rapidly with Cellosolve (ethylene glycol monoethyl

ether) as the clearing agent. The larvae, from 70- or 80-percent

alcohol, are partly dehydrated in 95 percent alcohol, cleared in

Cellosolve, and covered with balsam after the excess Cellosolve

has been removed. A modification of this technique for preparation of special study material has been described by Foote (113).

Creosote U.S.P. is also very satisfactory as a clearing medium

after rapid dehydration in 95- and 100-percent alcohol (227).

This has been modified by employing a mixture of creosote and

balsam as the mounting medium, the balsam having been heated

first to drive off most of the xylene (316), This requires a drying

period of 10 to 12 days, preferably in an incubator. Euparol may

be used as the mounting medium after the use of either Cellosolve

or creosote, or after dehydration in 95- and 100-percent alcohol.

The formula for Berlese's chloral-gum solution, as previously

recommended (193), provides a medium for making temporary

study mounts. Unfortunately, it is not satisfactory for permanent

mounts because of evaporation or deterioration. In making mounts

of whole larvae it is the practice to cut the abdomen nearly in

two between the 7th and 8th segments. This permits the air tube

to lie flat when the cover slip is applied and exposes the lateral

aspect of the 8th and anal segments.

For taxonomic study, or for identification of species difficult

to determine, it is frequently desirable to have both the larval

skin and the adult of the same individual. For such rearings a
14 AGR. HANDBOOK 173, U. S. DEFT. OF AGRICULTURE

nearly mature larva is isolated in a separate dish, and when

pupation occurs the larval skin is removed with a pipette, spread

out carefully on a slide, and a mount prepared by one of the

methods mentioned above. The dish or vial containing the pupa

should be covered with cloth or a larger dish, or plugged with

cotton. After the adult emerges it should be kept for about 24

hours before it is killed to allow time for the sclerotin to become

thoroughly hardened. The pupal skin should be mounted on the

same slide with that of the larva but under a separate cover. To

mount the pupal skin the céphalothorax should be detached from

FIGURE 6.—Mosquito light trap.

M & A 10505
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 15

the abdomen and then opened out with dissecting needles so as

to lie flat. The slide is then numbered to identify it with the

reared adult.

Adult mosquitoes are usually collected while they are either

biting or resting in secluded corners inside or underneath buildings, in tree holes, et cetera. A chloroform-killing tube (ñg. 5)

is convenient for this purpose. It may be prepared by placing

a half-inch layer of cut rubber bands in the bottom of a large

shell vial or test tube, saturating the rubber with chloroform,

and covering with a plug of crumpled paper and a circle of stiff

paper. The writers prefer a shell vial seven-eighths of an inch

in diameter and about 5 inches long. When the tubes are kept

tightly corked, the rubber retains the chloroform for some time.

As moisture is liable to condense on the inside of the tubes, the

dead mosquitoes should not be left in them long. Cyanide may

also be used in the killing tubes, but it has a slower killing action

and should be handled with extreme caution since it is a deadly

poison to man. Several types of aspirators have been used for

taking specimens alive or in large numbers. Individual mosquitoes

may be kept alive in a piece of large-diameter glass tubing or a

vial open at both ends. The top is covered with bobinette and the

bottom is equipped with a cork in which a capillary vial is inserted to furnish water. A freshly cut raisin or wad of cotton

saturated with sweetened water is placed on top of the bobinette

to furnish food. A glass lantern chimney may be similarly

equipped as a container for large numbers of adults. Generally

the bottom is covered with a sheet of thin rubber and a i/o-inch

slit in the rubber permits the collector to add adults to the

chamber by blowing through the aspirator. The simple tube type

of aspirator is the most useful for adding specimens through the

slit.

A pill box, with a thin layer of cotton pressed down into the

bottom and sides, may be used for holding or shipping the specimens. The box should not contain so much cotton that the

specimens will come in contact with the lid, and if more than a

wisp is used, its weight will cause it to shift about in the box

during shipment. Cotton should not be placed on top of the

specimens. Specimens that have been moistened, crushed, or

rubbed are usually unsatisfactory for identiñcation.

The suction type of light trap, as developed by New Jersey

workers (fig. 6), is being used extensively for obtaining samples

of the mosquito fauna of an area, and records of the relative

abundance of different species, particularly in connection with

control operations. The specimens are more or less damaged during capture, however, and are usually unsuitable for the permanent collection. The original model of this trap and an improved design have been described by Mulhern (231), Light-trap

collections in comparison with collections by other methods are

discussed under Mosquito Surveys in the section on Mosquito

Adult specimens that are to be retained in the permanent collection should be mounted and pinned into a Schmitt box or

similar tight insect box having a bottom lining of sheet cork

or balsa wood. Freshly killed specimens may be mounted on
16 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

M & A 14048 A

FIGURE 7.—Types of mounts used for adult mosquitoes.

minuten pins, and dry specimens on points cut from stiff paper

(fig. 7). In using the paper mount, an entomological pin is passed

through the base of the narrow paper triangle and a small drop

of cement is dabbed on the tip of the paper. The paper is then

pressed gently onto the side of the thorax of the mosquito, with

the tip directed toward the mesonotum. Care should be taken

not to smear the legs or wings with the cement. For uniformity,

the points are usually stuck onto the left side of the specimen.

A cellulose cement is preferable to the shellac formerly employed ;

it may be purchased at hardware stores in small tubes or it may

be prepared by dissolving celluloid in amyl acetate (known also

as pear oil or banana oil). Ambroid thinned with amyl acetate

is one of the cements used. Because of the volatility of the amyl

acetate the stock of cement must be thinned frequently. In making

the minuten-pin mount, the small pin is stuck into a small square

or rectangular piece of cork, through which is also passed a

larger pin (fig. 7). The tip of the small pin is then thrust through

the thorax of the mosquito, usually from between the coxae toward the back. The tip of the pin should not protrude through

the mesonotum. Very small, dry specimens may be stuck on the

side of the minuten-pin point with a drop of cement, instead of

using the paper point. The No. 3 entomological pin is probably

the best size to use with both types of mounts.

Care must be taken to protect the stored specimens from insect

pests, and for this purpose flake naphthalene or paradichlorobenzene are most frequently used. The material may be sprinkled

in the box or placed in a perforated container fastened in one

corner of the box. Specimens that are to be kept temporarily in

pill boxes may be protected by sprinkling a little flake naphtha-
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 17

lene on the bottom of the box and covering this with a thin layer

of cotton before introducing the mosquitoes. For longer storage

the pill boxes may be kept in a larger box containing naphthalene

or paradichlorobenzene, which must be renewed occasionally.

MOSQUITOES AND DISEASE

Mosquitoes rank as the most important insect pests of man.

They scourge him with their vicious biting attacks, which alone

have made sections of the globe almost uninhabitable, but of more

importance they transmit to him some of his most distressing

diseases. Chief of these are malaria, yellow fever, dengue, filariasis, and the arthropod-borne encephalitides. In addition, mosquitoes are known to transmit heartworm of dogs, bird malaria,

and fowl pox. Also, they transport to man the eggs of a South

American fly, Dermatobia, whose larvae upon hatching cause

myiasis, and they may be vectors of tularemia and leishmaniasis.

Malaria is caused by tiny protozoan parasites of the genus Plasynodium, which invade the red-blood corpuscles. There are four

species of human malaria parasites, three of which—Plasmodiurn

vivax, P. malariae, P. falcipariim—are distributed generally in

temperate and tropical regions, while the fourth, P. ovale, is rare

and of restricted occurrence. P. vivax is the most common form

in North America. It causes vivax or tertian malaria, so called

because the paroxysms or "chills and fever'' which are characteristic of the disease occur every third day (48 hours). In falcii:)arum, malignant or sub-tertian malaria, the paroxysms occur

every 40 to 48 hours, while with P. malariae or quartan infections

the paroxysms come every 72 hours.

Both man and the mosquito are essential to the propagation of

malaria. In man, the parasite lives and multiples in the red-blood

corpuscles. This is the asexual cycle, which usually is completed

in from 48 to 72 hours depending on the parasite species, following which immense numbers of blood cells disintegrate and discharge the parasites and their wastes into the bloodstream to

cause the characteristic paroxysms of the disease. Meanwhile,

part of the parasites develop into sexual forms, or gametocytes.

The anopheline mosquito here becomes essential, for further

development depends on these gametocytes reaching the stomach

of the mosquito where the sexual cycle takes place. This requires

from 10 to 20 days or more, and results in tremendous multiplication of the parasites which, as sporozoites, penetrate all parts

of the mosquito body. Eventually these concentrate in the salivary

glands from which the mosquito ejects them along with saliva

while biting its next host. An infected mosquito may transmit

the disease to several human victims before its store of sporozoites is exhausted. All of the anopheline mosquitoes of the

United States except georgianus and perplexens have been found

susceptible to infection by malaria parasites. However, epidemiologic evidence indicates that Anopheles qiiadrimacidatus in the

Southern and Eastern States and A. freeborni Aitken in the West

are the principal transmitters, and control measures therefore

have been aimed specifically against these two species with effective results. Anojoheles albimanus, an important malaria vector
18 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

in the American tropics, occurs sparingly in south Texas and on

the Florida Keys but is not known to be of importance in these

areas.

Opinions differ as to whether malaria existed in North America

prior to the coming of the white man ; it is known that the disease

became a problem early in the history of the United States and

it may have been brought here originally by the colonists or by

slaves from Africa. By the middle of the 19th Century malaria

was occurring widely throughout the country. Since then it has

undergone cycles of increasing and decreasing occurrence, the

latter predominating, with the net result that by 1930 it was

restricted as a disease of importance to the Southeastern States

where it appeared to be firmly entrenched. However, a very rapid

decline in incidence began in the mid 1930's, which carried

through the war years. Subsequently, using principally DDT

against the anopheline vectors. State and Federal health agencies

have cooperated in waging an eradication campaign. As a result,

except for minor self-limiting episodes, malaria transmission has

almost ceased in this country. Total cases reported to the Public

Health Service numbered only 707 and 477, respectively, for

1954 and 1955, and of these only 8 and 4 cases, respectively, were

confirmed as primarily indigenous in origin. In 1956, the number

of reported cases fell to 234, but the number of indigenous cases

has not been reported.

Malaria also occurs among birds in the United States, and

seven avian species of Plasmodium, all different from the human

malaria parasites, are known to occur here. These malarias may

be transmitted by various mosquitoes of the genera Culex, Aedes,

and Anopheles, Bird malaria infections are not known to be of

particular importance in this country where they aifect principally

the passerine birds. In Ceylon and Borneo, however, other species

of Plasmodium occur, which cause high mortality in chickens and

ducks. Studies of avian malaria parasites have led to clarification

of many obscure phenomena in the biology of human malaria.

Also, the screening of anti-malaria drugs has been greatly expedited by the use of birds parasitized with various species of

Plasmodiu7n.

Yellow Fever is an acute febrile virus disease which is endemic

principally among monkeys and other primates in the jungles of

the American Tropics and in Africa. Here, various species of

jungle mosquitoes belonging to the genera Haemogogus, Aedes

Culex, Eretmapodites, Mansonia, Psorojjhora, Sabethes, Wyeomyia, and Limatus are known to be capable of transmitting the

virus. At present, Haemogogus spegazzinii Brethes is believed to be

the principal vector in the forests of South and Central America

On occasion the disease is carried to urban communities, and if

the domestic mosquito Aedes aegypti is abundant in such com

munities, the disease may spread rapidly. During the 18th and

19th centuries many epidemics occurred in Gulf and South At

lantic coastal cities in the United States, and at times the disease

spread to inland cities as well. Epidemics occurred occasionallv

in Atlantic ports as far north as Boston, being started there bv

infected A. aegijpti coming ashore from ships which had arrived

from infected ports farther south.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 19

In 1900, Dr. Walter Reed and his associates confirmed the observations and experiments of Dr. Carlos Finlay, a Cuban physician, which pointed to A. aegypti as the vector of yellow fever,

thus establishing that control of this mosquito was the clue to

controlling the disease.

The last epidemic of yellow fever in this country occurred in

1905, at which time the efficacy of Rnti-aegypti measures in controlling the disease was convincingly demonstrated, as it already

had been in countries to the south of us. Because yellow fever

continued to occur sporadically in some cities in the Tropics even

when A, aegypti populations were maintained at low levels by

control programs, and because of the relative ease with which

the species could be entirely eradicated from a community, the

Pan American Sanitary Organization, in 1947, decided that

eradication of aegypti from the southern continent was the only

logical goal to set in dealing with the problem. To date, several

countries are reported to have achieved such eradication and thus

have eliminated the possibility of urban yellow fever epidemics.

Nevertheless, jungle yellow fever is a continuing problem for

which no feasible eradicative measures are known. The vaccination of all persons likely to become exposed is the only solution

at present.

In the United States, federal quarantine officers at ports of

entry constantly are on guard to prevent A. aegypti from being

inadvertently brought into the United States, since such specimens may have become infected with yellow fever in their country

of origin. Officers also keep airport and dock areas under surveillance so that any vector breeding can be eliminated promptly,

thus preventing infestation of ships or planes leaving this country. Some port cities carry on mosquito-control programs, which

greatly facilitates this work, and may result eventually in the

eradication of aegypti from these localities. Such eradication already has been reported for Key West, Fla.

Dengue, or ''breakbone'' fever, is a nonfatal, acute, febrile,

virus disease that is endemic in tropical and sub-tropical areas

of the world. The disease is characterized by the sudden onset of

high fever, severe headache, backache, and excruciating pains in

the joints—to which the common name ''breakbone" refers. Aed^s

aegijpti is the principal vector of dengue fever, although it also

is transmitted by other members of the subgenus Stegomyia such

as Aecles albopictus (Skuse) in the Philippines and Hawaii and

A. scutellaris (Wlk.) in New Guinea. Mosquitoes become mfected

by feeding on persons having the disease, during a 4- or 5-day

period beginning with the day preceeding the onset of symptoms.

Then follows a period of 8 to 14 days during which the virus

multiplies in the mosquito, after which the mosquito may remain

infective for life. Like yellow fever, dengue is a disease of

monkeys, and these animals serve as reservoirs between epidemic

Dengue may also occur in epidemic form. In the United States,

where A. aegypti is the vector, an epidemic in 1922 i^ the South

affected some two million persons, of which nuniber 600,000 to

1 000,000 were in Texas. In 1934 a less severe outbreak occurred

iA Florida and spread northward into Georgia.
20 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

The most important of the arthropod-borne viral encephalitides

occurring in the United States are eastern equine encephalitis

(EEE), western equine encephalitis (WEE), and St. Louis encephalitis (SLE), all of which appear to be transmitted principally

by mosquitoes. Similar viruses, such as Venezuelan and Japanese

B, are known in other parts of the world. Encephalitis results

from infections with specific neurotropic viruses which cause

inflamation of the central nervous system. The disease may be

very mild, moderately severe, or have an abrupt onset with high

fever and coma. The human fatality rate ranges from 5 to 60

percent, while the equine fatality rate may range up to 90 percent, the highest rates occurring in eastern equine infections.

Both the eastern and western viruses occur in the Southeastern

States, the former the more frequently (21), Although epidemics

among mules and horses have occurred, there have been few

human cases in this area. No outbreaks of St. Louis encephalitis

have been recorded but outbreaks have occurred in the bordering

States of Kentucky and Texas. In addition, to man and equinas,

these diseases aflfect a variety of other animals. Wild and domestic

birds frequently have been found with inapparent infections and

undoubtedly are important reservoirs. Present knowledge indicates a bird-mosquito-bird cycle of transmission for these

diseases, with equines and man only accidental hosts, playing no

essential role in the transmission chain.

Kelser (180) was the first to report experimental transmission

of encephalitis (WEE) with a species of mosquito (Aedes

aegypti). Since that time, numerous other mosquitoes have been

shown to be capable of transmitting the viruses of encephalitis experimentally, and a number of species have been found to harbor

infections acquired under natural conditions. In the Western

States, Culex tarsalis is believed to be the principal vector because

of the frequent occurrence of such infections. From records compiled by Ferguson {109), 144 isolations of WEE and 14 of SLE

had been made prior to 1951 from wild-caught specimens of this

species. Natural infections with the different viruses also have

been found in pooled lots of other mosquitoes that occur in the

Southeastern States. Some of the isolations were made from mosquitoes collected in the Southeast and some from mosquitoes collected in other parts of the country. The species and the types of

virus recorded are: Aedes dorsalis—WEE {139, 30I¡) SLE

{136) ; Aedes infirmatus—WEE {196) ; A. mitcheVae—EEE

{176) ; Anopheles cruciaiis—EEE {176,196): Culex vwiev<^ nini

ens-WEE and SLE {138) ; C. restnmis~WEE {Èm c'uSa

ínormía-WEE (^^5) ; C. ;../a..ra-EEE {71) ; Mansonia perturhans—EEE {16U).K new strain of virus called California encephalitis was isolated first in 1943 from specimens of A dorsalis

and subsequently from both A. dorsalis and C. tarsalis {136

lot), '

In quantitative studies of the virus-vector relationshin hv

Chamberlain et ah (72), the vector potentials of 23 specie of

mosquitoes with WEE and EEE viruses were determfned and

were rated as excellent, good, fair, and poor. For the South

eastern species the ratings were as follows: "Excellent" or

"Gooá"-Aedes aegypti (WEE, EEE), A. aíropdp¿ (WEE
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 21

EEE), A. sollicitans (WEE, EEE), A. triseriatus (WEE, EEE)

Culex restuans (EEE), Mansonia perUirbans (EEE), Orthovoi^^T.^ Â^^^^f^^^ (WEE, EEE), Psorophora confinnis (WEE,

^^•\l ^-/H^^^^ (WEE), P. discolor (WEE), P. ferox (WEE) ;

i^air —Aedes vexans (WEE, EEE), C2¿;ea; erraticus (EEE)

Psorophora ciliata (EEE), P. /^ro.r (EEE) ; -Poor-—Anoî^/i^Zei

Cretans (EEE), A, quadrimaculatiis (WEE, EEE), C^^i^o; p

quinquefasciatus (WEE, EEE), C. salinarms (WEE, EEE), Ma7^-

soma indtibitans (WEE), Psorophora discolor (EEE), and P

hoivardii (EEE).

Experimental transmission of encephalitis virus by several

of the above species has been reported by other v^orkers. Additional Southeastern species that have shown ability to transmit

infection include Aedes taeniorhynclms (WEE, EEE, SLE) and

A. sticticics (SLE). Also the St. Louis virus has been transmitted

experimentally by Aedes aegypti, A. vexans, Culex p, pipiens

C. p. quinquefasciatus, C. tarsalis, and Cidiseta inornata. These

records and a bibliography up to 1951 may be found in the valuable compilation by Ferguson (109),

Human filariasis is a disease of widespread occurrence in

the tropical and subtropical regions of the world. It formerly

was endemic in the Charleston, S. C, area but no cases have been

known there in recent years, and it appears to have been eradicated. Filariasis is caused by infections with nematode worms of

two species, Wuchereria bancrofti and W. malayi. The adult

worms live in various parts of the lymphatic system of man. The

young of these worms, or microfilaria, are discharged into the

blood stream where they are picked up by mosquitoes while biting.

About 10 days are required for development in the mosquito,

after which they migrate to the mouth parts, from which they

reach a new host when next the mosquito feeds. They are not

injected into the host directly but break from the labium of the

mosquito onto the skin and enter the body through the biting

puncture or other skin abrasion. Prolonged or repeated infections

in man may result in enormous enlargements of the external

genitalia, mammary glands, and limbs, often called elephantiasis.

Many mosquitoes of the genera Aedes, Culex, Mansonia, and

Anopheles suce known to be capable of acting as intermediate hosts

of the parasites in various parts of the world.

During World War II large U. S. forces served in areas in the

Pacific where filariasis was endemic, and some infections occurred. The majority of these cases became completely inactive

within 2 years after leaving the endemic area, and the very low

incidence made it unlikely that American mosquitoes would become infected by returning servicemen {106). Nevertheless, this

possibility was given careful consideration, and the ability of

several species of U. S. mosquitoes to act as vectors was

determined.

Newton, Wright, and Pratt (236) and Newton and Pratt (235)

concluded from a series of studies that under conditions favoring

transmission Culex p. pifpiens, C, p, quinquefasciatus, C, tarsalis

(and possibly C. erraticus), Psorophora discolor, P. confinnis, and

Anopheles albimanus would be potential vectors; that the infectability rates of Culex salinarius, C. nigripalpus, Aedes aegypti,
22 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

A. triseriatus, A. thibaulti, and Anopheles crucians were too low

to consider them as likely to play an important role in transmission; and that Aecles soUicitans, A, taeniorhynchus, A. vexans,

A. canadensis, Anopheles punctipemiis, A, ffuadrimaculaUis, Psorophora ciliata, P. cyaneseens, and Mansonia perturbans appeared

to be incapable of transmitting Wiichereria hancrofti infections.

Eyles and Most {106) observed the development of W. hancrofti

to advanced or infective stages in Cidex p. 2^ipiens, C. p. quinquefasciatuSy C. erraticiis, C, salinarius, Anopheles 2)unctipenniSy

A. ivalkeri, Aedes triseriatus, A. aegypti, A. atropalpus and Mansonia pertiirhans. They considered however that only the first

two of these species, which had infectivity rates of 83.5 percent

and 34.9 percent, respectively, were sufficiently susceptible to be

dangerous potential vectors. They reported that infections failed

to develop significantly in Anopheles qitadi'imacidatus, Aedes

atlanticus (or tormentor), and Psorophora ferox.

Another filarial worm, Dirofilaria immitis, causes heartworm

disease of dogs, an infection that is widespread in warm climates

and occurs commonly in the Southeastern States, where it causes

serious losses, particularly in coastal areas. The adult worms are

usually found in the right ventricle of the heart and the adjacent

pulmonary artery. The microfilaria are discharged into the blood

stream as with Wuchereria species. Transmission may be by

various species of mosquitoes of the genera Aedes, Culex, and

Anopheles. Hu {165) lists the following Southeastern species

which may serve as intermediate hosts: Ano2?heles piinctipennis,

Aedes aegypti, A. canadensis, A. soUicitans, A. taeniorhynchuis,

A. vexans, Cidex p. ^npiens, C. fatigans {p. quinquéfasciatus), C.

restuans, and probably C. salinarius. Yen {3S6) reported that

large numbers of larvae completed development in Anopheles

macidipennis Meig. and A. punctipennis, but that only a part of

the larvae reached the infective stage in Cidex territans (probably restuans), C. tarsalis, C. p. pipiens, Aedes cinereus, A. vexant,

A. canadensis, A. Stimulans, and others. None developed in Culiseta inornata, Mansonia perturbans, and Aedes trivittatus. Kartman {177) gave the following percentages of infections in specimens fed at the same time on an infected dog: A. quadriniaculatus—99.5, C. p. pipiens—29.S, C. p. quinquefasciatus—S9 9 A

aegypti—34.5. '

Fowl pox is a common and widespread virus disease of poultrv

which normally is transmitted by contact. It may be spread

also by a number of species of mosquitoes including Cidex v

pipiens, Aedes aegypti, and A. vexans, which occur in the Southl

eastern States {200, 216). ^^utn

MOSQUITO CONTROL

Mosquito control may be undertaken either as a means of

preventing mosquito-borne diseases or to ürotect man and aïïLals

from their VICIOUS attacks. Throughout the world the prSal

antimosquito activities have been directed toward dLease^Íe

vention. Withm the United States, however, many of the larS"

operations have been undertaken for the sole purpose of Kf

venting annoyance, for mosquitoes are not only^rdetriment to"
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 23

human welfare and happiness but their attacks result in direct

economic losses through reduced property values, injury to livestock, and reduced efficiency of labor. These losses frequently are

much greater than the cost of mosquito control.

Nearly everyone is familiar v^ith the efforts made to eliminate

mosquito-breeding places, and with the use of insecticides in

antimosquito work. The practical phases of the problem, however, form a large specialized subject, and a general summary

only will be given here with reference to underlying principles

and to the practices and materials that have become more or less

standardized, or are of recent development.

MOSQUITO SURVEYS

Mosquitoes have extremely diverse breeding habits, particularly

in respect to the type of place selected for oviposition. Because

of this diversity, the species to be dealt with and their individual

habits must be known before control measures can be applied

intelligently.

If the work is undertaken for the control of a disease such as

malaria or dengue fever, the presence of the disease itself, in the

Southern States at least, indicates the particular species responsible, because the vectors of these diseases are well known. A

study of the distribution of human cases serves to localize the

problem, and a mosquito survey is undertaken to aid in developing a plan of procedure. At the same time, the possibility of including control measures against purely obnoxious species should

not be overlooked. Where relief from annoyance is the main object, a thorough survey is necessary to determine the relative

importance of the different species, since more than one is usually

involved. Even in coastal areas, where it is known that the saltmarsh species are the principal culprits, it is still highly important to know whether fresh-water breeders are sufficiently

numerous to require consideration.

The surveys are begun by the collection and identification of

both adult and larval specimens. During an outbreak of mosquitoes the species involved can be determined quickly by collecting adults from various parts of the affected area. In localities

where mosquitoes are present more or less continuously, or where

outbreaks are of frequent recurrence, collecting should be repeated often enough to determine the relative annual abundance

of the different species. At the same time information should be

accumulated as to the breeding places of the common species,

the topography of the area, and the extent of the control problem.

A year should ordinarily be regarded as the minimum time for

such preliminary studies, since mosquito abundance varies greatly

with the seasons. Several years are required to obtain reliable

averages as to normal abundance. Although adult control operations and the treatment of obvious breeding places usually can be

begun before such an extensive survey is completed, the practice

of beginning major operations with inadequate information is

highly wasteful, and may result in complete loss of public confidence in a worthwhile project, possibly causing its abandonment.

An important item in the annual budget for financing the control

operations should be provision for continuing the systematic
24 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

collection and identification of specimens. Such work will furnish

invaluable information as to seasonal changes in the mosquito

problem and outbreaks from overlooked or distant breeding areas,

and is indispensable in measuring the results accomplished.

General methods of collecting mosquitoes have been discussed

in a preceding section. Some of the special methods employed for

obtaining data necessary in connection with surveys and control

operations are described in the following paragraphs.

BITING RECORDS AND LANDING COUNTS

Collecting mosquitoes while they are biting is the simplest and

most direct method of determining the relative abundance of the

different bloodsucking species. Such collections are usually made

with a chloroform tube or other type of killing bottle. For data

on comparative abundance in different parts of the area or at

different times of the year, stations are selected, and collections

made for equal periods and under conditions as nearly uniform

as possible. In obtaining such records the writers have adopted

the procedure of sitting on a box or stool at the selected place,

with the trouser legs rolled to the knees. After a minute or so

has been allowed for the mosquitoes to accumulate, they are collected as they alight for a period of 10 or 15 minutes (192). If

the collecting is done after dark, a flashlight is necessary. Two

15-minute collecting periods or three 10-minute periods may be

totaled and multiplied by 2 for the hourly rate. Collections made

during the first flight period (just at dark) should not be averaged

with later collections as the numbers are usually much larger at

that time.

When the mosquitoes are numerous, the numbers caught can

be mcreased considerably by placing a short paper funnel, or

guard, in the mouth of the collecting tube (fig. 5), since this

permits the collector to move to the next specimen without waiting for the first one to succumb to the chloroform fumes. The

guards are useful otherwise in conserving the strength of the

chloroform and in preventing the loss of specimens when the

mouth of the tube is turned downward. An aspirator is also used

for such collections.

When collecting after dark the writers have taken an average

of 10 mosquitoes per minute, or 600 per hour, with a chloroform

tube. If the mosauitoes are much more numerous than this the

discomfort of collecting is so great that it is considered sufficient

to record abundance as 600+, or other observed rate, per hour

When the collecting is to be done at different places by two or more

persons, preliminary collections should be made at one niace to

determine the relative attractiveness, and also the dexteritv of

the different collectors, as much variation has been found in tL^P

respects. tntiöe

For those species that attack during the day, such as thp C^OH

"^^J^^^ J^^^^^^^oes landing counts give rapid comparative rec

ords of the numbers present, and such counts are useful et

pecially m obtaining data before and after insecticidal treatmpi.f¡

to provide estimates of the reduction. These arrconvpnS

made by pausing for half a minute at the site to be saTplä and
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 25

then counting the number alighting on the front of the trousers

during one minute. The counts are sometimes made by two men,

one counting the mosquitoes alighting on the back of the other

(pi. 1, A). The records are obtained at a number of points to

give an average count for each plot or area.

HAND COLLECTIONS OR COUNTS OF RESTING MOSQUITOES

Some species can be obtained by daytime collecting in dark

corners and other places where the adults (including the males)

spend the daylight hours. This is an excellent method of obtaining

comparative data on densities of adult Anopheles, especially those

species found in the United States, since they fly into a shelter

at daybreak and remain quiet throughout the day. This method

is safer than the biting method if disease-bearing mosquitoes are

present.

Favorable daytime resting places for Anopheles are underneath

buildings that are raised 2 or 3 feet from the ground, and inside

tightly boarded outbuildings or similar locations. In making the

surveys, a series of stations well distributed over the area under

observation are selected, and weekly, biweekly, or monthly collections are made {162, 183). At each location the most favorable

resting place is selected after examination of all the buildings

on the premises. When the surveys are purely for comparative

purposes, the collecting station does not always need to be an

entire building if it is found that one part is more favorable than

another or that parts of the building are not conveniently accessible. Where the resting surface is fairly smooth and unobstructed,

a well-trained and reliable collector, with the aid of a flashlight,

can obtain satisfactory counts of the resting mosquitoes in much

less time than would be required for collecting the specimens in

killing bottles or aspirator tubes. The sex can be determined and

in most cases the species identified on sight. Collections over a

definite period (10 or 15 minutes) have been used as an index of

density, but they may be unrelialDle because of variations in the

rate of collection under different conditions as well as in the

mechanical limitations to the numbers of mosquitoes that can be

collected in a given time. Artificial resting places may be constructed in favorable sites from horizontally placed barrels or

boxes mounted on posts. A privy-type building with a lower panel

removed on the west side for entrance of the mosquitoes was

found to be much more efliicient than nail kegs {6Jf, 293), Various

traps for use in anopheline surveys have been discussed by

Bradley et al U8).

TRAP COLLECTIONS

For most species the suction type of light trap (fig. 6) is very

useful for obtaining samples of the mosquito population, for records of comparative abundance and, in control areas, for immediate information on the occurrence of outbreaks. In control

work the traps are placed at strategic places throughout the area

and are usually operated every night. For other purposes, the

traps may be run on a schedule of one or more nights each week.

The traps should be hung in an open space with the light itself
26 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

5 or 6 feet from the ground, and they should not be placed in the

immediate vicinity of a strong light if this can be avoided.

The number of mosquitoes caught per night frequently runs

into the hundreds or even thousands, and many other kinds of

insects are found in the killing bottles. These conditions make

the task of separating and identifying the material a considerable

one, especially v^hen the specimens are badly damaged or vs^et;

simplified procedures for this w^ork have been described by Bradley and Travis (51), Species not taken v^hile biting appear in the

light-trap collections, and it has been found that the different

bloodsucking species are not attracted to the light equally. Over

a series of nights the w^riters' trap-collection records have shov^n

more variation than the biting records, w^hich, of course, are the

more accurate index of annoyance. From a large series of trap

collections made in Florida only an occasional specimen of Aedes

aegypti has been obtained, and the numbers of Culex pipiens

quinquéfasciatus and C. nigripalqnis appear to be very small in

comparison with the amount of breeding in the neighborhood of

the traps. This also seems to be true to some extent of Anopheles

qiiadrimaculatus. The trap records, therefore, cannot be relied on

to provide exact indices of density for these species. Comparative

data for several methods of collecting have been given by Huffaker

and Back (167) and Provost (258),

Carbon dioxide attracts mosquitoes, and light-trap catches may

be increased by placing a piece of dry ice on or beside the trap,

(US, lU, 167). The traps also may be operated without lights,

but with the fan running, to sample the mosquitoes active at a

given site without an attrahent. A portable light trap that is

operated from an automobile battery has been described by Seaman (285), and a light trap powered by dry-cell batteries has

been described by Nelson and Chamberlain (234). A rotating trap

powered by an electric or gasoline motor has been designed by

Chamberlin and Lawson (7i). Rigid screen-wire funnels are

mounted at the ends of two or four arms, which extend at right

angles from the upright rotating central shaft. The device is

adjustable so the funnels may be operated at selected elevations

above the ground. The small end of the funnel is open and a

small cloth bag is attached to hold the insects that are swept in

A truck trap, consisting of a large screen funnel with a front

openmg 2x8 feet mounted on the top of the cab of a pick-uü

truck, was employed by Provost (258) to sample the mosquito

population on the wmg after dark. The truck was driven for set

distances at a speed sufficient to intercept the flying insects

Smaller funnels mounted on the front fenders of automobiles have

also been employed. ^

Animal-baited traps have been used for collecting mosquitoes

and, m the Tropics at least, have been employed for detSnW

densities of anophelme species that do not remain in accSblî

shelters during the daytime. A number of such traps have been described (see Bradley et aJ. (^8) ), ^"

COLLECTIONS OF LARVAE

Collections of larvae are made primarily to locate the breeding
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 27

places and determine their importance. Information as to the

comparative abundance of different species may be obtained from

the identification of a large series of collections. Rough estimates

of the relative abundance of a species can be obtained by counting the larvae per dip in a series of dips. The relative importance

of the area in mosquito production can be expressed numerically

by multiplying the average number per dip by a factor representing the extent of the breeding area (size times percentage of

breeding surface). The productivity of a breeding place per unit

of surface can also be determined by the use of cloth nets or

screen cages placed over the water {Jf3),

Breeding places may be divided into two general classes—permanent and temporary. The two classes frequently intergrade,

however, and the status of a given area may change over a period

of time. Anopheles and Cidex occur typically in the permanent

breeding places, whereas most Aedes and Psorophora are found

in the temporary collections of water produced by rainfall, floodwaters, or high tides. The status of the breeding places, particularly the permanent ones, as to productivity may change

greatly during the course of a season or from year to year, owing

to changes in the amount of aquatic growth or flotage, the

abundance of natural enemies, and other causes.

The importance of temporary breeding places of Aedes and

Psorophora is frequently difliicult to determine, because considerable time may elapse between broods. One may visit suspected low

areas repeatedly without finding larvae, but such areas must be

classified as potential breeding places, until repeated floodings with

negative results have been observed. The type of vegetation,

especially in salt marshes, is often an indicator of the suitability

of breeding conditions. Breeding occurs on the parts of the

marsh that are above the normal daily tidal range, and the elevations are indicated by the type of plant growth, since many of the

plant species are restricted rather sharply by the height of the

water table and the frequency of tidal coverage. More definite

information on suspected breeding areas can sometimes be obtained from samples of sod taken from dry depressions, or in

vegetated areas by scooping off a thin layer of topsoil with a

small shovel. Samples from different parts of the area are placed

in containers and covered with water to cause hatching of the

eggs, which may begin within a few minutes. Glass containers

are preferable, as the small larvae are more easily seen when

these containers are held against the light. If the sods are very

moist at the time of collection, drying in the air for a week or

so may be necessary to induce hatching of the eggs.

ENGINEERING SURVEYS

During a mosquito survey much information will be accumulated which will be valuable in determining the feasibility of an

anti-mosquito project, and the best method to be employed in

handling specific problems. Where malaria or salt-marsh mosquitoes are to be controlled, an engineering survey should next

be undertaken to lay out a detailed plan of procedure, and to

determine the approximate cost. Whether the local situation can
28 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

be handled successfully within the means at hand, or at a cost

commensurate with the probable benefits, is one of the first

questions to be decided. The legality of the proposed work (including jurisdiction over the area involved), the probability of

obtaining necessary easement on private property, et cetera,

must also be considered. A summary of State mosquito-control

legislation has been compiled by Keefe and Beadle {178).

Good maps are, of course, essential, and should be available

both in small scale for use as key maps and in large scale for

showing in detail the breeding places, ditching lay-outs, and

natural topographic features. The different areas and the individual breeding places can then be given names or numbers for

convenient reference to the mosquito-collection and engineering

notes. Aerial photographic maps of rural or marsh areas are

extremely valuable, as they show the bodies of water and the

types of vegetation. A note-card system should be arranged,

and complete records kept of all inspection and survey data.

For breeding-place or other preliminary surveys when elevations or exact locations are not required, the directions given by

Hülse {168) for preparing field maps in public-health work,

based on methods employed in the military service, are very

useful.

CONTROL OF MOSQUITO LARVAE

Mosquito-control measures are usually directed against the

larvae in areas where the breeding places may be eliminated by

filling, ditching, impounding, or other water-manipulation procedures, or where the chemical treatment of relatively restricted

breeding grounds will eliminate mosquito annoyance over an

extensive inhabited area. Anti-larval measures may not be economical where the breeding area is extensive and the nuisance

area small, particularly if the necessary measures will interfere

with important conservation programs.

ELIMINATION OF BREEDING PLACES

Where at all feasible, efforts should be made to eliminate the

breeding places permanently by filling, drainage, or sanitation.

Under some conditions breeding may be reduced or eliminated

by controlling the water level with impoundments

Filling IS frequently a practical method and gives permanent

relief when the fills are so placed or graded as to leave noTateîholdmg depressions. Large hydraulic fills may show shrinkage

or surface cracks upon drying and require one or iS^re reïrad

mgs to prevent mosquito breeding regraa-

^ Drainage undoubtedly has the widest application of the var

lous anti-larval measures, especially in the control of tL mallS

carriers and the salt-marsh species The drains.o-p nf ^^^^^^^

lands in the United States has d'one Such to Sc'^^^^

area and at the same time has made the land siiLhl.^^ •

culture. Drainage or ditching purely foi mosm l^''''

ever, should be looked upon as dLtfnct frZ ^^^^^^^

age, since it is directed mainly towaid the e^^ ^'f^-

water during the time reguire'd for larval dÄpmeit,°'/"to'a?d
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 29

in biological control. Comprehensive drainage plans should be

prepared with the aid of trained engineers.

Two phases of the drainage problem in mosquito control to

which attention should be called are its possible effects upon

wildlife, and its possible effect upon soil conservation. Through

cooperative biological studies efforts have been made to determine what measures may be applied to large swamp areas,

particularly those not close to centers of population, which would

disturb as little as possible the natural breeding and feeding

grounds of aquatic wild fowl and other desirable animal life

without sacrificing the success of the mosquito-control project.

Specialists in soil conservation have called attention to the adverse effects of the drainage of natural upland storage basins,

the cleaning of stream channels, and the **brushing'' of stream

banks, all of which increase the rapidity of run-off of flood-water

with consequent erosion that may cause serious damage to agricultural lands. Such erosion is said to cause frequently a gradual

widening of the flood plain and silting-up of downstream areas

which may create mosquito-producing areas as serious as those

remedied.

Such factors as these must be considered in planning mosquitocontrol programs, and they emphasize the need of obtaining advice from competent specialists when making the preliminary

surveys. The impoundage of water rather than drainage may be

employed successfully in many cases, both in salt-water and freshwater areas, since an open pond with clean margins and containing mosquito-destroying flsh is not favorable for mosquito

breeding (312, pi. 2). Where the sacrifice of wildlife habitats

appears necessary to accomplish effective mosquito control, a

decision must be made as to the greater benefit to be derived.

Sanitation, as applied to mosquito control, includes such measures as the elimination of artificial and other breeding places.

It also involves the treatment of permanent bodies of water and

removal of aquatic vegetation and other protective harborage to

make them unfavorable for mosquito development.

LARVICIDES

Various larvicides are employed where permanent methods of

control are not feasible, or as temporary expedients until permanent measures can be installed. Prior to World War II the most

widely used larvicides were oils, paris green, and pyrethrum. Investigations begun in 1943 at the Orlando laboratory demonstrated

the remarkable eflPectiveness of DDT which, unlike the larvicides

previously used, gave unprecedented control at dosages as low as

2 to 4 quarts of spray per acre, thus permitting the treatment of

large areas from the air at reasonable cost. Originally DDT was

used only in connection with military operations, but after the war

it became generally available and was adopted by many civilian

mosquito-control agencies. Within a short time several other chlorinated hydrocarbons also were shown to be effective larvicides.^

The extensive use of DDT for several years in some localities

resulted in the development of resistance in some species, notably

salt-marsh mosquitoes in Florida and Culex tarsalis in California,

whereas other species, such as Anopheles quadrimaculatus in the
30 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Tennessee Valley, have remained susceptible to DDT in a continuing control program. Resistance in restricted areas in Maryland and Georgia was reported in 1959. The species that became

resistant to DDT quickly acquired resistance to other chlorinated

hydrocarbon larvicides, but as yet they continue to be susceptible

to several of the newer insecticides, particularly to the organophosphorus compounds including parathion, EPN, malathion, Dipterex,

and Co-Ral. Increased resistance to malathion has been reported

in Ctdex tarsalis {126), and it should not be assumed that resistance to the organophosphorus larvicides will not eventually be

developed by other species.

The investigations leading to the recommendation of DDT as an

anopheline and culicine larvicide have been described by Deonier

et ai. {85, 87), Eide et ah {103), and Knipling {20J^). Many

agencies have contributed to the development of practical application methods. Detailed instructions as to the dosages, formulations,

and equipment used to apply DDT and other larvicides in various

situations will be found in U. S. Department of Agriculture

Circular 977 {310), American Mosquito Control Association Bulletins 1 and 2 {2, 3), and the Operational Memoranda issued annually by the Communicable Disease Center, United States Public

Health Service.

^ Technical DDT in an oil spray or in an aqueous emulsion of an

oil solution is the most widely used mosquito larvicide. In an oil

spray it is usually dissolved in kerosene or fuel oil in concentrations ranging from 5 to 1 percent or less, depending on the type

of dispersal equipment to be used. Concentrated solutions are sometimes made in other solvents to be diluted with oil as needed. The

remarkable larvicidal action of DDT permits great savings in

materials and labor. Whereas 10 to 25 gallons of oil without DDT

is applied per acre to control culicine larvae, control can be obtained with 5 quarts of a 1-percent DDT solution or 1 quart of a

5-percent solution when properly dispersed. To effect the greatest

savings the volume of solution applied should be the lowest that

will provide complete coverage with the equipment at hand and

the concentration should be adjusted to give a dosage of about 0 1

to 0.2 pound of DDT per acre. Under some conditions higher dosages may be required, as in heavily polluted waters for the control

of certain species of Culex, where about 0.4 pound iDer acrp i^

needed.

DDT emulsions are commonly prepared from concentrates containing 25 to 35 percent of the insecticide. These are diluted for

use and are applied at the same concentrations and dosage rates

as the oil solutions They are highly effective as larvicides, and

have obvious advantages over oil solutions in transportation but

are more injurious to fish and other beneficial aquaüc hfe '

^ni?7^^'''^^'''/T:v.P''?P^'*^^ ^ ^ wettable powders containing 50 to 75 percent of the insecticide, are less effective than oil soTu

tions or emulsions as larvicides and, if their use is dictatPd hv nfw

considerations, they should be applied at heavier Sel "^ "'""''

DDT dusts are highly effective against anoohelinp iT^v^o o A

are particularly useful where long-distance drS?^

treat wide breeding areas, but in many situations thp Hvff?? ^

excessive loss. Dusts penetrate thick ve^eS^^^ Ste"
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 31

than liquids. Against culicine mosquitoes, dusts are usually less

effective than solutions or emulsions, but rather heavy applications

to cisterns, tanks, dumps, and rubble-filled areas have been effective in the control of various species of Aecles and Culex during

military operations.

Granules or pellets containing 1 or 2 percent of the insecticide

and applied at about 5 or 10 pounds of granules per acre provide

effective larvicidal action together with good penetration of vegetative cover and elimination of loss by drifting. They have been

especially effective in Florida mangrove marshes when dispersed

from the air, although various insecticides other than DDT have

usually been employed because of DDT resistance. Several sizes

of granules are equally effective biologically ; those passing through

screens having 16/30 meshes per inch are a convenient size, and

are large enough to prevent drifting, yet small enough to provide

about 200 granules per square foot when distributed at 10 pounds

per acre.

Aircraft frequently are used for the dispersal of solutions, emulsions, dusts, and granules, but are rarely equipped to disperse suspensions of wettable powders, which require continuous agitation.

A 5-percent solution of DDT in oil has been used most commonly

with small planes. The output is usually adjusted to give a dosage

of 2 quarts (about 0.2 pound of DDT) per acre at normal flying

speed, although either the concentration or volume may be varied

to give any desired dosage. With larger and faster planes, 20-percent solutions have been applied at 1 pint per acre for effective

control. Emulsions are usually delivered from small planes under

conditions that permit low-altitude flying to reduce the evaporation

of the water. Aerial dusting is now seldom employed in mosquito

control, except where planes equipped for crop dusting are the only

ones available, since the dusts are less effective than sprays against

culicines. The aerial dispersal of granules has been discussed. DDT

has also been successfully used by dispersion as a smoke or thermal

aerosol produced by injecting DDT solutions into an extension of

the plane's exhaust pipe. The fine particles produced were especially effective against anopheline larvae at very low dosages, but the

amount lost by drift was so great that the smoke proved less satisfactory than sprays.

Prehatching treatments have been made possible by the longlasting qualities of DDT. It was found that 1 to 2 pounds of DDT

per acre, applied to fresh-water or salt-marsh areas prior to flooding, would prevent the development of successive broods of Aedes

as they hatch. Such treatments have been effective over an entire

season. For this type of treatment the wettable powder is perhaps

the most effective, although the other DDT formulations have also

proved satisfactory.

Benzene hexachloride, lindane, chlordane, dieldrin, toxaphene,

and heptachlor are among the other chlorinated hydrocarbons that

have been successfully employed as mosquito larvicides. Dosages

of 0.05 to 0.1 pound of hndane per acre, or of the gamma isomer if

the crude benzene hexachloride is used, have given satisfactory

control of various species, including salt-marsh mosquitoes. Dieldrin and heptachlor are about equally effective, and chlordane and

toxaphene slightly less so. Toxaphene, chlordane, and dieldrin are
32 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

extremely toxic to fish, and should not be used where fish are

important.

A number of organophosphorus compounds are highly toxic to

mosquito larvae and several have been used in practical control

programs. Parathion is one of the most effective. It is usually

applied at 0.1 pound per acre, although it has sometimes been effective at lower dosages. Parathion and malathion sprays have

been used extensively and safely by the California Bureau of

Vector Control and by mosquito abatement districts in that State

to control DDT-resistant Cidex tarsalis, Aedes nigromaculis

(Ludl.) and A. dorsalis in irrigated pastures. Applications of

granular parathion have been made for the control of resistant

salt-marsh mosquitoes in Florida. EPN has also been highly effective, but has not found wide use since it is marketed only as a

wettable powder. Other organophosphorus compounds that have

shown promise as larvicides, but are still in the process of evaluation, are Dipterex, Co-Ral, dicapthon, American Cyanamid 12008,

ronnel, Phosdrin, and Bayer 29493 (Baytex).

A pyrethrum emulsion known as the New Jersey mosquito larvicide has long been used successfully against fresh- and salt-water

species. The formula in concentrated form contains 66 percent of

kerosene or similar petroleum distillate, 0.07 percent of pyrethrins,

and 0.5 percent of a neutral emulsifier in water. One part of the

concentrate is diluted with nine parts of water and applied at the

rate of 15 to 25 gallons per acre. Because of its safety to fish and

aquatic plants it is recommended for use in fish pools and similar

places where other larvicides may prove harmful.

Petroleum oils are the first chemicals known to have been used

in efforts to control mosquito larvae. They are effective against

nearly all species of economic importance. Diesel oil. No. 2 fuel oil,

and kerosene—the products most widely used—are comparatively

inexpensive and readily obtainable. However, they must be applied

in relatively large quantities, they may constitute a fire hazard

under some conditions, and may be injurious to fish and wildlife

if improperly used.

Paris green is highly toxic as a stomach poison to mosquito

larvae (11) and before the advent of DDT was used extensively

in the control of anopheline mosquitoes. An interesting development, in which paris green is incorporated in a granular vermiculite formulation, has been reported by Rogers and Rathburn

{267a), It appears to be of considerable potential value against

culicme larvae, especially where resistance to organic insecticides

has developed.

NATURAL ENEMIES OF LARVAE

Various kinds of insects and other animals prey upon mosquito

larvae and undoubtedly destroy large numbers. Of the manv

natural enemies, however, only the small larva-eating fishes have

been found of practical use in control. In the Southern States the

most important of these is the top-water minnow (Gambusia afñ^

ms holbrooki), which occurs m both fresh and brackish water

These fish are most effective against subsurface-feeding larvae

and m places where the larvae are not protected by aquatic vegeta
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 33

tion. They have been used to stock ornamental pools and other

isolated bodies of water. They are highly useful in permanent

ponds and m the salt marshes. Proper ditching and impounding

will aid in making the best use of fish in marsh areas. The International Health Board of the Rockefeller Foundation (267) has

prepared a review of the literature up to 1924 on the use of fish

for mosquito control, and Hinman (152, 153) has given numerous

references on other predators of mosquitoes.

Different aquatic plants are claimed to be of value in eliminating

mosquito breeding. One species of Cham (C. fragilis) appears to

exert a deterrent effect, although certain other species of this

genus have been found to be innocuous. The bladderworts ( Utricularia) capture and destroy small aquatic animals, including mosquito larvae. Duckweed (Lemna) and similar floating plants

(Azolla and Wolffia) may form such dense mats on the water surface that they act as mechanical barriers to mosquito breeding,

although Anopheles and Culex larvae are found in abundance

when the growth is scattered. Water-hyacinth (Piaropus) may

also act in somewhat the same way. Matheson (212) gives a

review of the literature on this subject.

CONTROL OF ADULT MOSQUITOES

Screens, bed nets, contact and residual sprays, repellents,

smudges, and fumigants are all employed for protection against

mosquito annoyance. Certain birds, bats, and predaceous insects

prey upon mosquitoes, but seem of little practical value in control.

In the screening of houses, galvanized or copper screens are used

and the 16-mesh screen has come to be standard for this purpose.

Copper (or bronze) screens, although higher in first cost, are the

most durable, especially in the vicinity of salt water. Monel-metal,

aluminium, and plastic screens have also been recommended in

such situations. To be effective, the screening must be carefully

done and special attention paid to the fitting of door and window

frames, as mosquitoes will find entrance through very small openings. Bed nets made of open-mesh cloth are used extensively in

some localities in the absence of, or to supplement, screening. They

are frequently employed for the protection of individuals, especially in malarious or salt-marsh areas. To be of value they should be

in good repair and carefully adjusted.

The advent of DDT and other residual insecticides during and

after World War II opened up possibilities of controlling adult

mosquitoes which previously would have been considered fantastic.

Large outdoor areas can now be freed of these pests by applications

of the new insecticides with modern spray equipment ; homes and

other buildings can be treated eflfectively and quickly with insecticidal aerosols; and perhaps most important of all, residual

insecticides, when sprayed on the interior walls of buildings and

other resting places of adult mosquitoes, give effective control over

long periods. To this characteristic is due the principal credit for

the elimination of malaria transmission from several countries,

including the United States, where our economic loss from this

disease was estimated to be $500,000,000 per year as late as 1938

(327). Because of these phenomenal results, plans are now being
34 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

developed by the World Health Organization looking toward the

stupendous task of world-wide malaria eradication. The future

outlook for use of these materials is not all rosy, however, for the

phenomenon of resistance develops in some anopheline species as

well as in species of other genera after continued use of these

insecticides. This situation makes it imperative that the materials

be used according to the best procedures devised, and also that research be continued to keep abreast of the problem through the

development of new chemicals, formulations, and application

techniques.

CONTROL OF MosQurroEs IN BUILDINGS AND SHELTERS

Before World War II the principal insecticide used against adult

mosquitoes was pyrethrum, although certain thiocyanate insecticides were also employed. Commercial sprays and equipment developed for household use provided much relief for the individual

in his home. In Africa and India pyrethrum sprays were used in

huts and houses to control malaria (S).

The high-pressure pyrethrum aerosol, which was devised by

Goodhue and Sullivan (128) of the former Bureau of Entomology

and Plant Quarantine and developed in cooperation with the Army

and Navy, provided a revolutionary and convenient method of

dispersing insecticides in air. Later, low-pressure bombs were produced, and now a variety of insecticides and synergists are used

to provide quick knockdown and kill of the mosquitoes within an

enclosed space.

Early in 1943 members of the Orlando laboratory staff noted

the unusual residual properties of DDT in the control of adult

mosquitoes (119, 204) and, as a result of tests in Arkansas (120),

reported that dosages of 50 to 200 mg. per square foot in buildings

and other resting places gave effective control of Anopheles quadrimaculatiis through the entire 5-month observation period. Various

agencies participated in the development and practical application

of DDT as an adulticide. The application of DDT residues in buildings has been the principal or only method used in several successful campaigns against mosquito-borne diseases.

Residual applications of DDT in buildings are usually made at

200 mg. per square foot, from oil solutions, emulsions, or suspensions of wettable powder. One gallon of a 5-percent solution or

emulsion is usually applied for each 1,000 square feet of surface

When suspensions are used, a concentration of 3.75 percent of DDT

from a 75-percent wettable powder is recommended to reduce

clogging of the nozzle, and the rate is increased to lU gallons Der

1,000 square feet. ^

Chlordane at 2 percent and lindane at 1 percent are effective in

residual sprays against mosquitoes. Both exhibit considerable

vapor toxicity, but are not nearly so long-lasting as DDT Dieldrin

at 0.5 percent is also highly effective, and is longer lasting than

Imdane or chlordane, but is also more toxic to mammals Crude

benzene hexachloride is sometimes used in place of lindane at

equal concentrations of the gamma isomer, when the advantage oí

lower cost outweighs the disadvantage of stronger odor

Some species of mosquitoes have developed physiological or be^
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 35

havioral resistance to DDT and other chlorinated hydrocarbon

insecticides, leading to increased interest in the organophosphorus

and carbamate compounds as residual treatments. Laboratory

tests have demonstrated that parathion, Diazinon, malathion,

Bayer 29493 (Baytex), Hercules AC 5727, and other compounds of

this group do provide effective residues, and some are sufficiently

safe to give promise of future usefulness. Malathion has been

effective in field tests against Anopheles quadrimacidatus.

CONTROL OF MOSQUITOES OUTDOORS

Before World War II, studies in New Jersey {123, 12U, 313)

demonstrated that people at outdoor gatherings could be protected

from much annoyance by a thorough spraying of the grounds and

surrounding vegetation with pyrethrum emulsions. Knipling {20Jf)

reported early experiments showing that DDT was effective as an

outdoor residual insecticide on vegetation, or as a contact spray

applied from the ground or the air. In recent years DDT and other

newer compounds have been employed to destroy adult mosquitoes

over large areas. Airplane spraying is particularly well adapted

to this purpose, and special equipment has been devised for treating fairly large areas from the ground.

In early tests in small plots against salt-marsh mosquitoes in

Florida, residual applications of 3 to 5 gallons per acre of oil or

emulsion sprays containing 5 percent of DDT greatly reduced the

mosquitoes for several weeks. In later tests, DDT at 2 pounds per

acre in 50 gallons of emulsion was effective only 3 days {310), and

it was believed that resistance may have been a factor in the reduced period of protection. In the later tests, lindane and BHC

(40 percent of gamma isomer) applied at i o pound per acre caused

great reductions for about 2 weeks. In these tests the degree of

protection in small treated areas varied with the number of

migrating mosquitoes and the time required for the insecticides

to exert their residual toxicity.

Residual applications have been used with some success to form

barrier belts around larval breeding areas, and thus kill the adult

mosquitoes as they penetrate the belt on their migratory flights.

Contact, or space, sprays for the destruction of adult mosquitoes

have been widely used on a practical scale, more often in the

abatement of mosquito annoyance than in disease-control programs. DDT is the insecticide most often used and the application

rate is usually 0.1 to 0.3 pound per acre. The material is distributed

in the form of an emulsion or oil solution containing from 1 to 20

percent of DDT, depending upon the dispersal equipment utilized.

Air-drifted fogs and smokes are effective under suitable weather

conditions.

As resistance to DDT has developed in certain species of mosquitoes, attention was given to alternate insecticides. Benzene

hexachloride was demonstrated by Deonier et al. {86) to be effective for the control of DDT-resistant salt-marsh mosquitoes.

A dosage of 0.05 to 0.1 pound of the gamma isomer per acre is

reauired under most conditions. Later, however, salt-marsh mosquitoes in some Florida localities also developed resistance to

benzene hexachloride after it had been used for several seasons.
36 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Malathion was reported to be effective against Aedes nigromaculis

n CaX and against the DDT- and BHC-resistant salt-marsh

mosquitoes in Florida. Dosages of 0.1 pound per acre are effective

3i ideal conditions, but usually 0.25 to 0.5 pound is required.

Increased resistance to malathion in C^dex tars^^^^^^

reported from California (126). Dibrom ^^^d Bayer 2949o (Baytex) have also shown promise in small-scale held tests.

INSECTICIDE DISPERSAL EQUIPMENT

The various items of equipment found useful in the chemical

control of mosquitoes are discussed in some detail m publications

bv the U S. Department of Agriculture and American Mosquito

Control Association (2. 3, 310), and will be reviewed only briefly.

Liquefied-gas aerosol bombs are most useful m providing temporary and local relief by killing the adult mosquitoes m houses,

tents, or other enclosures. They may also be used to eliminate

mosquitoes temporarily from small outdoor areas.

Hand-operated, compressed-air sprayers of 1- to 3-gallon capacity are convenient for the application of residual treatments

inside houses or other buildings (pi. IB) and for larvicidmg small

areas, especially in the control of domestic mosquitoes Handoperated atomizers and bucket pumps may be used m the same

way. Larger, power-operated sprayers, mounted on vehicles and

supplied with sufficient lengths of hose, may be used to advantage

in applying residual treatments inside a large group of buildings,

residual or contact sprays to outdoor areas, or for the larvicidmg

of areas accessible to the vehicle. A fan-type nozzle is preferable

for residual applications, but a cone-type nozzle may be used.

Several types of mist blowers and thermal fog generators have

been developed especially for the control of flying insects. In one

type of mist blower the liquid is pumped at low pressure to the

spray head, where it is delivered into the center of a high-speed

air blast (pi. 3A). Although the primary use of the machine is to

provide an adulticidal contact spray in outdoor areas, the output

is sufficient to provide some residual action. The machine may also

be used for larvicide applications, especially where the breeding

places are fairly wide and best covered by a wind-drifted spray.

In another type of mist blower, the Microsol (pi. 3B) the spray

is finely atomized by means of spinning disks ana then dispersed

by an air blast. This machine is sometimes called a mechanical

aerosol generator, but the droplets fall more nearly into the

category of a mist ; that is, greater than 50 microns in diameter.

A small electric model for use in buildings is said to give a large

proportion of droplets within the aerosol range. A power pneumatic sprayer designed by Husman (171) for outdoor use provides

particles in both the aerosol and mist ranges.

In the thermal fog generator (pi. 4A) heat is employed to

break up the liquid into fine droplets, which are drifted across

country as dense fogs to control adult mosquitoes. Several machines based on this principle, such as the TIFA, Dyna-fog, and

Swing Fog, are available. The heated exhaust gas from motor

vehicle has been utilized for the production of insecticidal smoke.

The particles in fogs and smokes are very small and remain airborne much longer than particles in most sprays. For this reason
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 37

they are at the mercy of atmospheric conditions and may show

variable results. In moderate breezes the fog may be carried long

distances, and in rising air currents it will rise above the lower

levels where the mosquitoes are resting or flying. On the other

hand, with a light breeze in a favorable direction and with air

movements toward the ground, which occurs when the temperature of the ground is lower than that of the air (inversion

conditions), fog drifting slowly through an area may be highly

efrective.

Hand-operated or power-driven dusters may be utilized for

larvicidal operations. No special equipment is available for applying larvicidal granules from the ground.

Airplanes were first employed experimentally in 1923 by members of the then Bureau of Entomology {187, 188) for the control

of malaria mosquito larvae with paris green dusts, and were subsequently used by the Tennessee Valley Authority and other

agencies. The newer, more potent insecticides are especially adaptable to airplane dispersal as sprays or granules.

The first airplane spray equipment developed and tested with

DDT solutions was designed for use on an L-4 (Piper Cub) airplane. In this model, known as the Husman-Longcoy spray unit,

the insecticide was dispersed through six nozzles on the trailing

edge of a venturi beneath the fuselage. Soon afterward, an underwing spray boom was developed and used both on monoplanes

of the L-4 type and biplanes of the PT-17 (N 25 Stearman) type

(pi. 4B). Openings of a suitable size were drilled in the boom,

and a breaker bar was mounted behind the boom to increase the

breakup of the droplets. Spray nozzles were also used on the spray

booms in place of the drilled holes and breaker bar.

After World War II large numbers of surplus training planes,

principally PT-17's, became available, and most of the civilian

mosquito-control work has been done with planes of this type.

Larger planes, particularly C-47's, have been used in military

operations. Spray equipment of this type is used in adulticiding

and larviciding programs.

Airplanes equipped for dusting agricultural crops are sometimes utilized in the application of larvicidal dusts. With slight

modifications in the hoppers, dusting planes are readily adapted

to the dispersal of granules—an efficient method of larviciding in

breeding areas covered with vegetation.

Insecticidal smokes, produced by injecting DDT solutions into

an extension of the plane's exhaust pipe, have been used in some

larviciding operations, but are usually less effective than sprays.

Malathion smokes give high kills of adult mosquitoes.

MOSQUITO REPELLENTS

Various essential oils, such as citronella, have long been used to

protect individuals from mosquito bites, but systematic investigations of repellents were not undertaken until a comparatively

recent date. In 1940 Granett {129) reported a suitable procedure

for the critical evaluation of repellents, and summarized the results with a series of nearly 1,000 materials. A mixture of organic

compounds was superior to citronella and the other oils, and a
38 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

large field of investigation was suggested. In 1945 Granett {130)

reported the effectiveness of ethyl hexanediol as a mosquito repellent, and in 1946 Travis et al. {307, 308) summarized the research at Orlando that led to recommendations for the repellents

used by the Armed Forces during World War II. The results of

screening tests with thousands of compounds as mosquito repellents were compiled by King {186), and studies in which the

better repellents were tested against a variety of species have

been reported by various workers at Orlando {300, 122).

Diethyltoluamide is the most effective of the mosquito repellents and is satisfactory for application to the skin or clothing.

It gives protection against a wide range of species, and when

applied to the skin it repels for longer periods than other

materials, and is more resistant to removal by rubbing or wiping.

Technical diethyltoluamide contains various proportions of the

meta, ortho, and para isomers. The meta isomer is the most effective, and should make up the major portion of products used

as mosquito repellents.

Ethyl hexanediol, dimethyl phthalate, and dimethyl carbate

also are highly effective and are safe for general use on the skin

or clothing. Each of these chemicals is outstanding against certain species of insects, but varies in its effectiveness on different

persons. The following mixture of these compounds has been

found effective against a wider range of species, under more

w'ldely divergent climatic conditions, and on more individuals

than the single compounds (figures in parts by weight) :

Formula M-2020

Dimethyl phthalate 4

Dimethyl carbate 3

Ethyl hexanediol 3

Although toxicologists have found these repellents safe for

use on skm, some persons may be allergic to them. Any of the

repellents may cause some smarting when applied to the mucous

membranes or to areas where the skin is especially tender, such

as the eyelids Sometimes the treated skin will at first feel warm

especially if it is moist with sweat, but usually the sensation tsTs

only a few minutes and causes no injurv. Repellents cause a

severe but temporary stinging if they get StSthe eyes and

therefore should not be applied too liberally on the forEd A

50-percent solution of diethyltoluamide in alcohol is the most

acceptable cosmetically, having little or no greasy feeling on the

tS'J}^^lt ^Pellents affect paints, varnishes, soiSe typïoTsyn!

thetic cloth (rayon but not nylon), and many of the plastics ?o

varying degrees They Avill not damage cotton or .îiol cloth if iî

contains no synthetic fibers. Ethvl hexanediol ha^ fhp laiS .^ Î

on pa nted surfaces; diethyltoluamide Laî o rïShÎes S

method ,s to shake a few drops frlTetottTe'lnrtheTaZs
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 39

of the hands, smear evenly, and then apply thoroughly to the

backs of the hands, wrists, neck, ears, face, or any other exposed

skin, much as in washing. Sufficient repellent should be applied to

give a uniform film. All these repellents except 50-percent solutions of diethyltoluamide in alcohol feel oily to the skin, and may

therefore be objectionable to some individuals. However, the protection they afford from biting insects more than compensates

for this oiliness. Under favorable conditions one treatment will

last several hours on most people, but should be repeated when the

insects resume biting.

These repellents can be used also to keep mosquitoes from biting

through the clothing. Properly treated clothing will give protection for several days. One soaking with water, however, destroys

the repellent action. The repellent can be sprayed or daubed on

the clothing only in areas where the bites occur, such as across the

shoulders and along the thighs, but the best method of obtaining

protection under all conditions of exposure is to impregnate all

the outer clothing that will be worn in the field. Use about 1/15

ounce per square foot of cloth, or a total of 21/2 ounces (5 tablespoonfuls) to a jacket (or shirt), trousers, and socks of medium

size. Do not treat the underwear. Dissolve the repellent in enough

dry-cleaning fluid to wet the garments thoroughly without any

excess, about 3 pints for an outfit of heavy cotton cloth being

required. After all parts of the garment have been saturated

with the solution, allow the cleaning fluid to evaporate.

An emulsion can be made by mixing 21/2 ounces (5 tablespoonfuls) of the repellent with 3 pints of water and i/i ounce (II/2

teaspoonfuls) of an emulsifier or 1 ounce (2 tablespoonfuls) of

soap. Suitable emulsifiers are Stéarate 61-C-2280 (a polyalkylene

glycol stéarate) ; Tween 80 (sorbitan monooleate, polyalkylene

derivative) ; and a polymerized glycol monostearate, monooleate,

or monolaurate. Many synthetic household detergents are not

suitable for making emulsions, but most laundry soaps are

satisfactory. Dissolve the emulsifier or soap in the water and add

the repellent slowly while stirring vigorously. If large quantities

of clothing are to be treated, a stock solution containing 90 percent of the repellent and 10 percent of emulsifier can be prepared,

and added to water as needed at the rate of 1/2 pmt to 1 gallon.

Saturate all parts of the garments with the emulsion, wring

lightly, and dry thoroughly before wearing.

The most effective repellents for treatment of clothing are

diethyltoluamide, and 2-butyl-2-ethyl-l,3-propanediol The latter

compound is a solid at normal temperatures and is best applied

by the impregnation method as a solution. It should not be

applied to the skin.

SPECIFIC PROBLEMS

A few notes are given below in regard to problems of control

of some of the more important mosquito species.

THE COMMON MALARIA MOSQUITO

(Anopheles qimdrimaculatiis)

This species develops principally in more or less permanent
40 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

bodies of fresh water such as pools, ponds, and marshes containing vegetation and floating debris, shallow vegetated lake margins, and places of like nature (pi. 5). Such situations occur frequently in and adjacent to rural settlements and small towns,

which become infested with large concentrations of quadrimacu-

¡atus as a result. Because of this situation, malaria in the Southeastern States occurred to a greater extent in these smaller communities than in more favorably located cities, or towns where

municipal developments had destroyed the breeding places. As

has been discussed elsewhere, this disease was virtually eradicated

from the United States as a result of a comprehensive program

carried out by Federal and State health agencies in which DDT

residual home spraying in malarious areas was the principal

means, larvicides being employed but rarely. Nevertheless, it may

be well to give continuing attention to programs for the reduction of this vector as it often becomes sufficiently abundant to be

a serious pest and would present a disease hazard should any

resurgence of malaria occur.

The older chemical larvicides, such as oils and paris green,

used for treating anopheline breeding areas that cannot otherwise

be cared for, will not be discussed here, since they are treated

adequately in other literature (79, 181, 318, 328), and they now

have been superseded by DDT and other modern larvicides, which

have been discussed in the preceding sections. It may be appropriate to mention here, however, that the use of airplanes in

mosquito control was first developed by the senior writers in 1923-

24 in connection with applying paris green dusts for the control

of qiiadrimacHiatus (187,188). With the development of resistance

to organic insecticides, the use of paris green may again be of

great value in an anopheline control program.

A. quadrimaculatus enters houses to obtain blood meals from

humans, and seeks shelter in dark cool situations within doors

during the day. When these daytime haunts are coated with lethal

applications of DDT or other residual insecticides, mosquitoes

which come m to rest are killed. Thus, while the overall reduction

m the mosquito population present in the area may be negligible

this selective killing of individuals that have fed on man and

may be infected with malaria parasites breaks the chain of transmission.

Procedures and formulations for the application of residual

sprays are discussed under the section Mosquito Control.

A. quadrimaculatus does not ordinarily fly long distances and

the control of breeding within a l-mile radius of a populated

area has so reduced the numbers of mosquitoes as to prevent

malaria transmission m such communities (12 121 ^06 207)

In this work the first essential is the permanent elimination of

Z; "T^^'i, ^^^''' ^^^'"^Ç ^^ ^^^^^^^^- Ditches and the mar- gins of the deeper ponds and lakes should be kept free of vegetation. Breeding m shallow ponds full of aquatic growth, or in the

beds of occasionally flowing streams can sometimes be controlled

economically by impounding the water with dams to a depth su^^^^

ficient to overcome the aquatic vegetation ipl. 2). Periodic fluctuation of the water level in such impounded areas is Zo^^^

reducing the marginal growth and flotage. The imp^SSdage o?
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 41

large bodies of water for hydroelectric or other purposes, however, has introduced serious problems in malaria control, and

special legislation has been enacted by Southern States covering

the measures that must be taken to prevent breeding of anopheline mosquitoes in such projects. The Tennessee Valley Authority

found it necessary to provide for an extensive program of Anopheles control in the impounded areas on the Tennessee River {29),

As this work progressed it proved essential to establish minimum

requirements as to reservoir preparation and shoreline improvement with special reference to vegetation control, and to provide

for adequate water-fluctuation schedules {155, SOI).

THE DOMESTIC MOSQUITOES

{Aedes aegypti and Culex pipiens quinquéfasciatus)

Urban antimosquito campaigns usually combine control measures against the yellow-fever and the southern house mosquitoes,

and, although their practical control offers no unsurmountable

difficulties, continuous efforts and expense are required to keep

the numbers reduced. These species differ in breeding habits. Culex

p, quinquéfasciatus breeds especially in polluted waters and ground

pools. Rain barrels and similar water containers are important

breeding places for both. Where the water in these receptacles is

required for domestic purposes, the barrels should be kept tightly

covered and the water drawn from a spigot at the bottom ; otherwise, they require weekly emptying or treatment with a larvicide,

either of which is likely to be neglected. When larvae are present,

spraying the surface with a small amount of kerosene or a pyrethrum fly spray is effective, and imparts little odor to the water.

Fire barrels should be treated with DDT or some other effective

larvicide. Tubs and other casual water containers left in the yard

should be turned upside down when not in use, and worn-out

equipment should be disposed of.

With the yellow-fever mosquito {Aedes aegijpti), the smaller

water containers, such as old cans, bottles, flower vases, and obstructed eaves troughs, are important, and a campaign against this

ST)ecies should begin with a clean-un of yards and vacant lots.

During the dengue-fever-control work in Florida in 1934, a large

proportion of the discarded automobile tire casings left in the open

were found to contain water with aegypti larvae, and many breeding places were found in automobile-wrecking yards. Toilet bowls

and flush-tanks in vacant houses and apartments require systematic attention, and collections of water in the basements of buildings should not be overlooked. The adults of this snecies do not fly

far and when they become troublesome at any point the breeding

source can usually be found on the premises or nearby. An essential part of any aeqypti control program is the freauent and

thorough inspection of premises by well-trained men. ^^^^^.^"^

reason breeding sources cannot be eliminated, thev should be

sprayed or dusted with a larvicide such as DDT Bv intensive

antilarval measures, aegvvti has been eradicated from^ large

areas, even entire countries, in South America An eradication

program from the Western Hemisphere including the United

States is now under consideration by pubhc health workers.
42 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

One of the most prolific sources of production for the southern

house mosquito (Cvlex p. (¡vivquefasciatus) in cities or towns,

is the storm-sewer catch basin, which is designed almost universally with a watertight debris trap below the level of the

outlet. The larvae of Aedes aecjijpti also have been found in these

places. In an antimosquito program the catch basins are usually

treated with a larvicide periodically by the use of special equipment installed on trucks or motorcycles. Other important sources

of mass production of C. p, quinquefasciatus are open cesspools, badly drained street gutters, and polluted ground pools,

especially around city dumps or sewage outlets. Wherever possible these breeding places should be eliminated permanently by

drainage, or the cesspools effectively covered; otherwise they

require frequent treatments with larvicides.

Waste water from food-processing plants or industrial operations, and occasionally the effluent from inefficiently operated

sewage-disposal systems, provides ideal breeding conditions if

allowed to collect in pools or sluggish, weed-grown streams. Laryicidal treatment of such collections of water serves to abate the

immediate mosquito nuisance. Many State and Federal agencies

are active in the development of measures to eliminate the pollution of streams and ponds from such sources.

SALT-MARSH MOSQI ITOES

(Principally Aedes solUcitans and A. taeniorhynchvs)

The salt-marsh mosquitoes fly extremely long distances. Migratory swarms have been observed 40 miles or more from their

breedmg places, although the average length of flight is, of course,

much less. Because of their great flight range, local work against

these species may be of little benefit, and control programs are

usually undertaken on a countywide basis. Generallv, the actual

control work should begin on the breeding marshes nearest the

population centers, and should progress outwardly until the

desired results are obtained.

Investigations and control work against the salt-marsh species

were begun m New Jersey more than 50 years ago, and New

Jersey s example has been followed by most of the North Atlantic

States. On the South Atlantic and Gulf Coasts, except in a few

nrS'fn 1Q^?^''^^^ no large-scale operations had been attempted

f.lZ K 1? ' ^^^"^ advantage was taken of the opportunity offered by the programs of the Federal Emergency Relief and the

Civil Works Administrations. By 1957 all bït one of the coastal

counties m Florida had established antimosquito distrkts

Mosquito-breedmg conditions in salt marshes and the methods

employed to overcome them are extremely varied In genSal the

rÄ'ni'''^ '^ i^^ ^^'^' ^^ th^ ^^^^h that'are S coveíed by daily tides, usually m potholes and depressions of various
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 43

cess to the pools and ponds or other places where the larvae

accumulate as the surface water is drained off.

The ditch system on any marsh should be designed to give the

maximum degree of mosquito control with the least amount of

ditching. The system for each marsh should be determined by the

conditions presented, and stereotyped drainage systems should

be avoided. Preliminary surveys are necessary to determine any

natural drainage and to locate the waterholding, mosquito-breedmg areas. The natural channels should then be improved and the

ditches located so that they will extend from these channels to

the breeding portions of the marsh, following the lowest contours.

In general, the ditches should not be cut directly into ponds, but

rather connected by short spurs, since it is often difficult to maintain the desired depth in the soft mud of the pond bottoms. On

large marshes which are so flat that natural drainage is negligible

or difficult to discern, the usual practice is to place the ditches

in a parallel system at intervals of 100 to 300 feet, with main

outlet ditches as needed. Even on such marshes, however, the

amount of ditching may often be reduced by making preliminary

surveys to locate the mosquito-breeding portions of the marsh

and limiting control work to such areas. The installation of rim

ditches for draining areas adjacent to the highland, where heavy

breeding often occurs, is necessary on most marshes.

In New Jersey where the marshes are usually well sodded, the

standard ditch is 10 inches wide by about 20 inches deep, with

the sides perpendicular. In other areas, especially in the South,

the width and depth of the ditches have to be modified to meet

other soil conditions and problems incident to other types of

marsh vegetation. In Florida the ditches originally installed by

hand were mostly 30 to 40 inches wide. Since 1953, by virtue of

its State subvention funds with emphasis on permanent measures,

ditching for salt-marsh-mosquito control has been greatly extended in Florida. In order to obtain the most economical operation, hand ditching has been almost entirely replaced by machine

ditching, principally with draglines. The width of machine ditches

generally ranges from 2 feet in marshes underlain with rock to

15 feet in muddy flats. The majority of the dragline ditches, constructed in open marsh and mangrove, range in width from

8 to 12 feet depending on soil conditions. The short laterals extending beyond the spoil piles along the ditches may be as narrow

as 4 feet, but they are usually placed at the same depth as the

main ditch. Little or no attention is given to sloping the sides

of machine ditches in open marsh and mangrove. The ditches are

dug to a minimum depth of one foot below mean low water and

are made sufficiently wide so that allowance is made for caving

of the perpendicular banks and still maintaining adequate depth

and width.

Various types of heavy machinery for digging and cleaning

the ditches have been developed by workers in the North, and

special kinds of ditching spades and other tools have been designed for use by hand labor. The type of marsh in the locality in

which work is to be done should govern the selection of tools. The

kinds of machine ditchers or special spades developed in New

Jersey do not appear to be adapted for use in most of the south-
44 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

ern marshes. As mentioned above, a dragline of suitable size is

used to advantage for large ditches or outlet canals, and backhoes, clamshells, or other types of equipment may be used in

special circumstances. In most Florida marshes the draglines are

operated on mats, but they can be used from barges in otherv^ise

inaccessible areas.

Another important method of treating certain classes of marsh

is the installation of dikes and tide gates to prevent the entrance

of high tides. The tide gates, opening at low tide, also provide

for the run-off of rain water. Under special conditions the tide

gates may be reversed to permit the entrance of high tides and

to impound the water on the marsh, or pumps may be used to

flood the impounded areas as required. This is effective in reducing

Aedes breeding, since it is the alternate drying and flooding of the

marshes that brings about the hatching of their eggs.

Experimental work in the early 1930's by members of the Bureau of Entomology and Plant Quarantine in the vicinity of Savannah, Ga., showed that the shutting off of the tides from marshes

by means of dikes and tide gates so that the marshes became dry,

eliminated much breeding of the sand fly (Cidicoides) as well as

of the mosquito. It is probable that this practice can be combined

with ditching in other sections where the sand fly is a serious

problem.

In the southern half of Florida, where Aedes taeniorhynchuis is

the predominant salt-marsh mosquito, the marsh vegetation is

dominated by growths of several species of mangrove (pi. 6, B).

Although this presents a special clearing problem with respect to

hand-ditching operations, construction of ditches with a dragline

is only slightly impeded. In constructing the ditch, the dragline

operator sometimes can clear his own right-of-way by swinging

his dragline bucket through the growth of mangrove ahead of

the machine. Dynamite ditching has been employed in such

marshes and compares favorably in cost with hand labor. It is

more expensive than the use of a dragline, but may be used in

small areas inaccessible to these machines.

Another difficult problem is encountered in certain areas where

the marshes border more or less landlocked bodies of water in

which there is ordinarily little tidal range. Strong wind tides may

cause a flooding of these marshes, and continuous winds may

hold the vvater there long enough for a brood of mosquitoes to

develop even though the marsh is thoroughly ditched. To meet

this problem it has been proposed that such areas be diked and

the water removed or the water level maintained by pumping

or by artesian wells. The management of water on coastal marshes

m Flonda for mosquito control has been discussed by Philen and

hyTauSc'filli ^'^- ^^'^ '''' '^^'"^^ ^^^^^^^^ diainage aid

The control work against salt-marsh mosquitoes üarticularlv

m the North Atlantic States, has been criticiLd as unnece^^^^^^^

destroying the feeding and breeding grounds of wildlTTeseS

by mosquito-control and conservation agencies has been directed

toward providing mutually satisfactory control methods Ä

progress has been made, and joint efforts are bS?g contiS

It IS believed that many of the deeper ponds, as welfas tie Xnt
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 45

species that serve as food, can be saved without detriment to the

antimosquito work. If the ponds are of value as feeding and resting grounds for wild fowl, the ditches may be diverted or, if they

are run into the ponds, a sod dam or wooden spillgate may be

inserted, somewhat lower than the marsh level, to prevent complete drainage of the pond and to permit the inflow of high tides.

Deepening of the shallower ponds may be necessary to obtain

surface drainage of the neighboring marsh, and should greatly

increase their value and permanence. The proper management of

impoundments in many such areas will result in reducing the

numbers of Aedes while increasing the food for waterfowl (75).

Sump drainage may be employed to decrease mosquito breeding

in small, enclosed areas where drainage is difficult, or in areas

that are of special value from a conservation point of view. By

this method ditches are dug so that the water is concentrated in

the lowest portion of the marsh, where, if no pond or pool exists,

one is excavated to serve as a reservoir for maintaining predacious

M & A 14049

FIGURE 8.—Female mosquito (Aedes solUcitans), with names of the parts.

minnows and insect enemies of mosquito larvae. In successful

operations, light floodings of adjacent breeding areas are dramed

into the sump by the ditches, and when the whole marsh becomes

flooded the ditches facilitate access of the predators to all parts
46 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

of the area. Successful use of this method was reported by StutzMesono-f-a/Tj

Heaa^,

Sc¿y/e/7u/r7

Pos//^o/u/77

Afeso/7o^u/r?

Base ortv/r?^

Scu^e//u/r?

PosAno^u/77

--"Pa/fer

/^^a¿o/om/r?a/ se^me/r^-

M & A 14050

FIGURE 9.—A, Composite diagram of thorax of adult mosquito, showing the

groups of pleural bristles: apn, Anterior pronotal (prothoracic) ; ppn,

posterior pronotal (preepimeral) ; ^jpZ, propleural (prosternai); sp,

spiracular; î)sp, postspiracular; pa, prealar; sip, sternopleural ; ume, upper

mesepimeral; /me, lower mesepimeral. B, Posterior portion of thorax from

above.

^ Stutz, Fred H. Sixth Annual Report of the Broward County, Florida,

Anti-Mosquito District Covering Activities for 1940. 9 pp. (Mimeographed.)
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 47

cross ye/>7 ce//

Cos fa/ i^//7

r„^* / ^ , Pe//o/e o/'2'^ uosra/ Suàcos/a/ /77ara//?a/ ce//

^f^ / ,SuâcOSfc7/

' ce//

e (>?/;a/ i^e/^)

FIGURE 10—Wing of mosquito (Howard, Dyar, and Knab). The^corresponding symbols for the veins in the Comstock-Needham system are: 1, R- 2 1

and 2.2, R. and R3; 3, R....; 4.1, M,..; 4.2, M3; 5.1 and 5.2, Cn, and Cnl;

anterior cross vein, r-m\ posterior (basal) cross vein, M-cn

m Broward County, Fla., for controlling breeding in low subdivision squares where adequate drainage was impractical.

Another problem arises in connection with the lowering of the

water table, which results sometimes in an undesirable change in

¿/p/>er /?eac/' /?a/r

Zy^r7/€/7/7a/ Aa//^

^/7fer>/7<7

Za/era/ /pa/r

Suj6aorsa/Ái

ACT/era/

o/'S''^ ^eß/?7er?/

^/7a/sey/77c/7/

J^/7/ra/ j6r¿/sÁ —

^/7ú'/^/'//s ^

Z?OKta/ /¿//G^

^ &. K 14052

FIGURE 11.—Larva of Cidex p. quinquéfasciatus with the parts named

(Howard, Dyar, and Knab.)
48 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

plant species and sometimes in a subsidence of the marsh level.

As such changes vary greatly with different soil and tidal conditions, a thorough study of such factors should be included in the

original surveys. Where the soil texture indicates little waterholding capacity, excessive lowering of the water table may be

avoided in some cases by the use of very shallow ditches.

Larvicides are employed in salt-marsh-mosquito control for the

treatment of areas that are not taken care of by ditching. DDT

has been used most extensively, and large areas have been protected for several years by aerial applications of this larvicide.

However, salt-marsh mosquitoes have developed resistance to

this insecticide and other chlorinated hydrocarbons as well. Although these larvae are susceptible to the organophosphorus compounds, the development of resistance through larviciding has

led a number of mosquito-control agencies to avoid the use of

larvicides and to direct their chemical control measures against the

adults in inhabited areas in the belief that resistance is developed

more rapidly when larvicides are used than when the treatments

are limited to the adults. Where larvicides are used they are

usually applied as aerial sprays, but in some mangrove marshes,

and other areas covered with heavy vegetation, granules are used

to provide better penetration {118^ 179),

MOSQUITO IDENTIFICATION AND NOTES ON THE

GENERA AND SPECIES

The identification of the different species requires a knowledge

of mosquito anatomy, as the distinguishing characters consist of

^nfenna

Prear7/e/?r7a//?a/r

-Longer /jeoc/ Aa/r

Upper heoi//7a/r

0.1 /// III.

FIGURE 12.- Head of larva of Aedes vexans. " * " """
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 49

variations, frequently very slight, in shape, size, coloration, or

scaling of the different parts of the body. Illustrations are provided herein to show the names and locations of the principal

parts that are utilized in identification, and the diagnostic keys

have been made as simple as accuracy will permit.

A binocular dissecting microscope is necessary for satisfactory

examination of the external characters of adult mosquitoes. It

should be provided with objectives and oculars giving magnifications up to about 75 X. With high magnifications a spotlight or

other source of bright illumination is required. For the examination of larvae and slide mounts of male terminalia, a compound

microscope is needed and should be equipped for magnifications

of about 100 X and 400 X- An oil-immersion objective is ordinarily

used for advanced work on the male terminalia. For field work

and for provisional identification of adults, a good hand lens giving a magnification of 10 X to 15 X is very useful. In fact, after

one has become thoroughly familiar with the species of a locality,

he will be able to identify many of them with the hand lens, and

to recognize some of them even with the naked eye.

Workers inexperienced in systematic work with mosquitoes

should have on hand, for comparative study, at least a small series

of correctly identified species, which can be obtained by sending

carefully prepared material to a specialist with the request that

named specimens be returned. Until one has become familiar with

the species, the material should in any case be forwarded to an

authority for a check on the identifications when questions of

control or habits are involved, since misidentifications may cause

application of inappropriate control measures and result in program failures. Identifications may be obtained through the Entomology Research Division, U. S. Department of Agriculture, some

of the State universities and experiment stations, or State boards

of health.

The family Culicidae, which, as previously mentioned, contains

the true mosquitoes, is represented in the Southeastern States by

three subfamilies and eleven genera, as follows: subfamily Anophelinae with one genus. Anopheles, and 11 species: subfamily

Toxorhynchitinae with one genus, Toxorhynchites (formerly Megarhinus), one species, and two subspecies; and subfamily Culicinae with the remaining 9 genera and 64 species and subspecies,

as shown in table 1.

In the following pages a synoptic key is given for the separation

of the genera; then, under each genus, a brief account of the

adult and larval habits common to the genus and a brief description of the characters that help to define the genus, in addition to

those used in the key. This is followed by adult and larval keys to

the species of that genus, notes on the habits of each species, its

relation to disease or economic importance, its known distribution,

and a brief summary of the specific characters that may be of

help in identification. The descriptive matter applies primarily

to the species of a limited region, and variations or exceptions

that may occur in other regions are not noted unless they seem to

be of special interest. .

In Misc. Pub. 336 (193) the species of all genera were combmed

into a single key for adults and one for larvae, because the identi-
50 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

J

(^ tp S

M & A 14054

FIGURE 13.—Dorsal hairs of larva of Anopheles quadyñmaculatus: A, Head

and part of prothorax; B, abdominal segments 4 and 5. Head hairs: 1,

Preclypeal; 2, inner clypeal; 3, outer clypeal; 4, postclypeal; 5-7, frontal;

8-9, inner and outer sutural (occipital) ; 10, terminal antennal; 11, antennal;

12, preantennal (basal); 13, subbasal; 14, orbital; mxp, maxillary palp.

Prothorax: 1, Inner submedian; 2, middle submedian; 3, outer submedian;

4-7, lateral. Abdominal segment 4: 0, Anterior submedian; 1, palmate;

2, antepalmate; 3-5, sublateral; 6, lateral; tp, tergal plate. Segment 5:

pi, Enlarged palmate leaflet.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 51

fication of a species is, in many cases, much easier than the

identification of the genus by the usual microscopic characters.

Because the number of species now known in the area has increased conâderably, this method has been abandoned and separate keys to the genera have been prepared. Nevertheless, an

effort has been made, so far as possible, to add characters visible

with a hand lens, or at low magnification, that apply to most of

the species in a genus, at least in the Southeastern States, even

though they may not be strictly of generic rank.

The keys have been prepared in the usual form of opposed

couplets, with the most obvious character given in the first

sentence, usually followed by one or more characters that help to

describe the species or group. When there is doubt as to which

part of the couplet the specimen fits, it is the practice to follow

out both divisions to find a later fit if possible. It may be necessary

finally to compare the descriptions under several of the doubtful

species. In the adult keys to species, the characters apply primarily

to the females unless the male is mentioned. The males of most

0.5 /r?m

M & A 14055

FIGURE 14—Types of air tubes: A, Psorophora ferox (also showing terminal

segments of abdomen) ; B, Mansonia perturbans; C, Culiseta mornata.
52 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

species can be identified by the character given although some

differences occur in the distribution of pale scales. The white

bands on the abdomen, for example, cannot be used satisfactorily

in identifying the male of subgenus Cvlex.

The external characters and the names of the parts employed

in the descriptions of adult mosquitoes are shown in figures 8,

9, and 10, while those for the larvae are shown in figures 11

to 16.

^/r /¿/J6C

ûorsa/

preap/ca/----"^

%fp//?e

J.cr/era/ co/nó

/afera/

FIGURE 15—4, Posterior portion of abdomen of larva of 4^^1 °''

B, anal segment of larva of Aedet nrifrhluni n .^^ vexans; mitchelUi. ® mitchellae; C, same of Wyeomyia
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 53

M & A 14057

FIGURE 16.—Portions of heads of various mosquito larvae: A, Uranotaenm

sapphirina; B, Deinocerites cancer; C, Toxorhynchites septentrionalis;

D, Psorophora ciliata. Ip, lateral pouch; m6, mouth brush.

In preparing to identify a mosquito specimen, one should

make a preliminary examination at a comparatively low magnification, to note the more obvious markings of the proboscis,

tarsi, thorax, wings, and abdomen. With experience, the ap-
54 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

proximate position of the insect, frequently both genus and

species, is recognized from this examination. There should be

little difficulty in recognizing Anopheles and Toxorhynchites,

and only slightly more trouble perhaps with Uranotaenia and

Wyeomyia. After these genera are separated nearly all the species in which the tarsi are not ringed with white and the mesonotum is unornamented are Culex, Culiseta, or Deinocerites,

There are three species of ringed-legged Cidex but these are extremely rare in most of this area. In the species of the other

genera, except Psorophora c ¡janesee us and Aedes einereus ^ some

of the tarsal segments are ringed with white, or the mesonotum

has bicolorous scaling in definite patterns of stripes, lines, or

patches.

^ Aedes and Psorophora females can nearly always be distinguished from those of other genera by the tapered end of the

abdomen, as shown in figure 17. This is a very useful character

FIGURE 17.—Characteristic shapes of tip of female abdomen: A, Aedes (also

similar in Psorophora) ; B, Culex.

although there may be some uncertainty as to the shape at times,

particularly when the abdomen is distended with blood or ova

or has shriveled. Aedes and Psorophora are distinguished also

irom other mosquitoes, except Mansonia titiUans and M, indicöitans by the presence of postspiracular bristles, but these require higher magnification than can be obtained with a hand lens

m order to be seen. Psorophora are distinguished from Aedes by

having spiracular bristles as well, although these bristles are

irequently difficult to see even with a binocular microscope In

Psorophora, however, the dorsal or lateral pale scaling of'the

abdomen IS at or toward the apex of the segments, or diffuse

the base.'"" ' segmental scaling is basal, or extends from

There are a few species that are so similar in appearance that

tSoLh X' m.i''?''''' V'"" '"''^'^'y b^ adu/STngs a^^^ though the male terminaba may be quite distinct. In these

nstances, the principal distinctions of the terminalia are included

m the keys, while for the dark-legged species of Cidex a separate
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 55

key to the male terminalia has been provided. Study of the terminaiia requires advanced training, and illustrations of these structures have not been included, on the assumption that anyone making use oí this part of the keys would be familiar with this

specialized subject.

Since few mosquitoes are known by common names, the scientinc name is, as a rule, used to designate the kind under discussion.« During the early years of mosquito research, considerable

contusion was caused by revisions of generic and specific names.

i:* ortunately these names have now become much more stabilized

and only two changes, due to the application of the rules of priority, have been made since the last edition of Miscellaneous Publication No. 336 {193), These are the change of genus Megarhimts to

loxorliynchites and of Culex apicalis to C. terntans. Many synonyms of the names now in use have appeared in the literature

but omy a few of those in recent use are shown here. More complete lists may be found in Dyar (.95), Edwards {102), and

elsewhere.

The alphabetical listing of genera and species as given in table

1 has been followed in the notes and description.

KEY TO SUBFAMILIES AND GENERA

ADULTS

1. Palpi of both sexes about as long as the proboscis, the male palpi

club-shaped (fig. 1, A and D) ; abdomen with a vestiture of fine

hairs, at least dorsally; scutellum evenly rounded.

Subfamily Anophelinae, genus Anopheles p. 85

Female palpi much shorter than proboscis; male palpi long or

short, not clubbed at tip (fig. 1, B and C) ; abdomen covered

w^ith flat scales 2

2. Outer half of proboscis tapered and bent downward at nearly

a right angle (fig. 2, A); posterior border of scutellum evenly

rounded and scaled. Very large iridescent species of striking

appearance.

Subfamily Toxorhynchitinae, genus Toxorhynchites p. 149

Proboscis of nearly uniform thickness, straight or slightly

curved; scutellum trilobed, with separated tufts of setae or

scale patches (fig. 9, ^).

Subfamily Culicinae 3

3. Second marginal cell less than half as long as its petiole (fig. 18) ;

very small species with lines of bluish or purplish iridescent

scales on thorax and base of vein 5; palpi short in both sexes

Uranotaenia p. 151

^ In scientific terminology two names, the generic and the specific, are

employed for each kind of organism. A genus is sometimes divided into

subgenera and, when given, the subgeneric name is inserted in parentheses

between the genus and species. The species may also be divided into subspecies or varieties. The name, spelled out or abbreviated, of the person who

first named the species, is often added after the specific name. If the species

has been changed to another genus, the name of the author is enclosed in

parentheses. The designation of a species may therefore appear as Aedes

aegypti or Aedes (Stegomyia) aegypti (L.), etc. When a species of mosquito

shows geographically defined grouping with some taxonomic differences, but

not sufficient characters to show distinct species, the differing kinds are

treated as subspecies, in which case the specific name of the type (the one

first described) is repeated and the others written as subspecies, e.g. Culex

pipiens pipiejis and C pipiens qiiinquefasciatus. These two were formerly

treated as distinct species but are now known to interbreed readily and to

produce intermediate forms.
56 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Second marginal cell at least as long as its petiole; iridescent

scaling if present not as above; male palpi long in most genera 4

4. Thorax thickly covered with broad, closely appressed dark brown

scales; abdomen somewhat compressed laterally, the scales

entirely dark dorsally and white ventrally, the two colors

meeting to form an almost straight line along the middle of

the sides; postnotum with a tuft of setae. Very small species

Wyeoniyia p. 153

Thorax not completely covered with broad dark scales (some

species of Psorophora partly covered with broad whitish

scales) ; abdomen not as above, usually flattened dorsoventrally,

or if rounded or distended, the colors not forming a straight

line ; postnotum bare 5

M & A 14059

FIGURE 18.—Wing of Uranotaenia, showing relation of length of the second

miarginal cell (6) to its petiole (a); also the short anal vein (an).

5.

6.

9.

Tip of abdomen with a tapered appearance (fig. 17, A), segment 7

much narrower than segment 6, segment 8 still narrower and

completely retractile, the cerci long and exserted; postspiracular

bristles present 6

Tip of abdomen truncate or bluntly rounded (fig. 17, B), segment

7 not markedly narrowed, segment 8 not retractile but frequently

quite short, the cerci usually retracted or only indistinctly

visible; postspiracular bristles absent (except in Mansonia titillans and 71/. induhitans) 7

Dorsal or lateral white scaling of abdominal tergites apical or the

segments almost entirely pale-scaled; spiracular bristles present;

mesonotum with at least some broad appressed scales (except

in subgenus Grahhamia) Psorophora p. 137

Dorsal or lateral pale scaling of abdominal tergites basal, sometirnes with a median stripe extending to apex of the segments;

spiracular bristles absent; scales of mesonotum narrow or

lanceolate Aedes p. 59

Wing scales very broad, mixed brown and white; setae absent at

base of vein 1; proboscis and legs speckled with white scales

and tarsi ringed with white g

Wing scales long and slender on veins 2 to 4, or if moderateiy

broadened, not mixed brown and white; a few setae present on

base of vein 1 on upper side of wing in Culcx and Deinocerites,

proboscis and legs not speckled and tarsi not ringed in the majority of species g

Mesonotum with fine longitudinal lines of white scales (fig. 19, /)

tarsal segments with both apical and basal white rings; segment

4 of fore tarsi very short, about as long as wide

iv/r 4. -^1, 4. 1. T , Orthopodoniyia ip.lM Mesonotum without such lines; tarsal segments not ringed apically

segment 4 of fore tarsi normal, considerably longer than wide

][icinsoizîcL X) 127

Antenna much longer than proboscis, the first flagella r seo-ment as

long as several of the succeeding segments; sternopleuron almost

completely shingled with dark appressed scales; male palpi and

antennae similar to those of female. The crabhole mosquito

found m coastal areas of southern Florida Deinocerite^ p 126
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 57

D

M & A 14060

FIGURE 19.—Examples of thoracic markings of mosquitoes (diagrammatic) :

A, Aedes aegyjjti; B, A. infirmatus; C, A. atlanticus; D, A. thibaulti;

E, A. trivittatits: F, A. triseriatifs; G. Psorophora varijoes; H, P. ciliata

(at a scale one-half the size of the others) ; /, Ovthopodomyia signífera;

J, Uranotaenia sapphirina; K, Culex restuans.
58 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Antenna normal, not longer than proboscis, the first flagellar

segment about the same length as succeeding ones; sternopleuron

not shingled with dark scales; male palpi long, the antennae

plumose 10

10. Spiracular bristles absent; base of subcostal vein without a tuft

of setae on underside of wing Ctilex p. 103

Spiracular bristles present; base of subcostal vein with a row or

tuft of setae on underside of wing. (If it is uncertain to which

of these two genera the specimen belongs, it should be run under

genus Cnlex where a notation is inserted in the proper place

for separating each of the two species of Culiseta) .... Cidiseta p. 124

LARVAE (FOURTH INSTAR)

1. Air tube lacking (spiracles on 8th abdominal segment sessile)

(fig. 20, A) \ some of abdominal segments with palmate hairs

(fig. 13, B) Subfamily Anophelinae, genus Anopheles p. 85

Eighth abdominal segment with an elongate air tube (figs. 20, B,

and 11) ; abdomen without palmate hairs 2

TT« OA T^ j • M & A 12938

I^IGURE 20.—Feedmg positions of mosquito larvae: A, Anojyheles; B, Cnlex.

Mouth brushes each consisting of about 10 stout, closely appressed

P)rehensile, curved rods without a row of comblike teeth on one

side; 8th segment without comb scales but with a sclerotized

plate having two spinulose bristles on posterior border pectén

lacking. Very large predacious larvae found in tree "^ holes

Subfamily Toxorhynchitinae, genus Toxorhynchites p. 149
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 59

Mouth brushes ciliform, the hairs numerous, or if with stout prehensile hairs (subgenus Psorophora) each hair with a row of

comblike teeth along the side; comb scales present on 8th segment, sometimes (genus Uranotacma) attached to a sclerotized

plate Subfamily Culicinae 3

3. Distal half of air tube attenuated (fig. 14, JB), with saw-toothed

projections at tip adapted for piercing the roots of aquatic

plants; pectén lacking Maiisonia p. 127

Air tube cylindrical, or spindle-shaped; the distal half not

attenuated 4

4. Air tube without a pectén 5

Air tube with a pectén 6

T). Comb scales in a single row; anal segment with a pair of ventrolateral hair tufts (fig. 15, C) instead of the usual median ventral

brush Wyeomyia p. 153

Comb with one row of short scales and a second row of very long,

pointed scales ; anal segment with the usual median ventral brush

consisting of a close-set row of tufts Orthopodoynyia p. 134

6. Head longer than wide, elliptical; upper and lower head hairs

single, stout, spinelike (fig. 16, A) \ eighth abdominal segment

with a large lateral plate having a row of tooth-like scales on

the posterior border Uranotaenia p. 151

Head wider than long; head hairs, slender, frequently multiple;

eighth segment not as above (small weak plates usually present

in subgenus Grahhamia) 7

7. Head with a prominent triangular pouch on each side (fig. 16, B) \

anal segment with a small sclerotized plate dorsally and ventrally; only two small bulbous anal gills. Found in crab holes

along the coast of southern Florida Deinocerites p. 126

Head without lateral pouches; anal segment completely ringed by

the plate or with only a dorsal saddle; four anal gills 8

8. Air tube with a single pair of ventro-lateral hair tufts, more or

less centrally placed (sometimes very small or obsolete in

Psorophora) .* • • • ^

Air tubs with several pairs of ventro-lateral hair tufts or single

hairs beyond the pectén, or a ventral row of numerous small

tufts 10

9. Anal segment completely ringed by sclerotic plate and the plate

pierced on mid-ventral line by tufts of the ventral brush (fig. 14,

A) Psorophora p. 137

Anal segment not completely ringed by plate, or if ringed, tufts

forming ventral brush are all posterior to plate (fig. 15, A

and B) Aedes p. 59

10. Air tube with four or more pairs of hair tufts or single hairs

beyond the pectén, none at base Culex p. 103

Air tube with a pair of multiple hair tufts near base with a paired

row of single hairs beyond the pectén (fig. 14, C), or a ventral

row of numerous small tufts Culiseta p. 124

Genus AEDES Meigen

This is a very large genus and all the species probably are

biters. Some of them are of great economic importance both as

pests and as vectors of diseases. The eggs of most species are

laid on damp soil in places that become filled periodically with

rainwater, tides, stream overflows, melting snow, et cetera. There

are a few species with specialized breeding places such as rock

holes, tree holes, and artificial water containers. The eggs are

usually laid on the sides of the container just above the water

line In general, the eggs can withstand long periods of drying,

and the winter is passed in this stage. Many species have only

a single annual generation but there may be several broods trom

the overwintered eggs when only a partial hatch occurs with each

flooding With other species the eggs can hatch the same season
60 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

in which they are laid, and several generations are produced if

alternate drying and flooding of the breeding places occurs.

When large areas such as salt marshes, pastures, and river

flood plains, heavily seeded vv ith eggs, are flooded, enormous broods

may be produced, resulting in a serious infestation of large areas

since some of these species commonly disperse for many miles.

The majority of species in this genus are ornamented with white

scales, many having the tarsi ringed with white, some with the

proboscis also ringed, and others with stripes or patches of white

or yellowish scales on the thorax. Only one of the species, A.

cinereus, lacks both tarsal and mesonotal markings. Spiracular

bristles are absent in the genus but postspiracular bristles are

present. Lower mesepimeral bristles are present in some species,

absent in others. The posterior border of the scutellum is partly

divided by indentation into three distinct lobes.

In the females of this genus abdominal segments 6 and 7 are

distinctly narrowed, segment 8 is small, usually completely retracted into segment 7, and the slender cerci protrude from the

tip of the abdomen (ñg. 17). This gives the abdomen a pointed

appearance, in comparison with the blunt shape of other mosquitoes, and is characteristic of Aedes and Psorophora females

—usually sufliicient by itself to identify these two genera with

the naked eye or a hand lens.

In the larvae the air tube is usually short and stout and has a

single pair of ventral hair tufts, usually fairly long and multiple,

placed near the middle of the tube or slightly beyond. The distal

teeth of the pectén may be either evenly or unevenly spaced and

may extend beyond the tuft. The comb on the eighth segment is

either in the form of a patch or a single row. The anal segment

may or may not be completely ringed by the sclerotic plate, and

the upper and lower head hairs may be single or multiple. These

characters are all used in classification. The pupae are of normal

appearance with fairly short air tubes. The eggs are somewhat

spmdle-shaped, and the surface has a fine, hexagonal reticulation

(fig. 3, B),

Nineteen of the 24 species of Aedes included here are placed

m the subgenus OchlerotaUis. The other five are divided as follows •

(Aedes) nnereus: {Stegomyia) aegypti: {Aedimorphns) vexans:

{tmlaya) triseriatus, and (Finlaya) atropalpus.

The separation of the subgenera is based principally on the

male termmalia and palpi. There are few or no characters in

tne témales or larvae that can be used satisfactorily For this

reason not much use is made of the subgeneric divisions in

practical mosquito work—at least for the species in our area

The mam subgeneric distinctions are:

1. Male palpi very short . ,

Male palpi about as long as proboscis Ï.'. Aetíes

'• ^wStínTpifalífaSnt '''''''''' ^^^ before-tip;-" eiaspette

3. Claspette lacking 3

Claspette present Stegomyia

^' ^'aAoS*^ " -^^Wy-deveíoped-basaí-lobVand- withouVan ^

^t&'']l'^i'^^^;-'-^^-^^^^^^ basaViobe and usually a'n apicaf'"'"^"

Ochlerotatus
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 61

KEY TO SPECIES

ADULTS

1. Some of tarsal segments ringed with white or yellowish scales 2

1 arsal segments not ringed with white 13

2. Proboscis with a white ring near middle 3

Proboscis not white-ringed (nearly all yellowish in A. fiilvus

pallcns) 5

3. Abdomen with basal transverse white bands but without a íongitudinal stripe ; dark portions of femora and tibiae not speckled with

white; mesonotum with dark brownish scales, sprinkled with

silver posteriorly taeniorhijnchus

Abdomen with a dorsal longitudinal stripe of white or yellowish

scales (fig. 8) in addition to transverse basal stripes; femora

and tibiae speckled with white; mesonotal scales golden 4

4. Wmg scales brown and white mixed; first hind tarsal segment w.th

a pale ring in middle. Important salt-marsh species sollicitans

Wmg scales entirely dark; first hind tarsal segment without a

central pale ring. A fresh-water breeder, usually rare mitchellae

5. Mesonotum bright yellow, with two shiny black spots on posterior

corners; proboscis, wing costa, and legs largely yellowish-scaled.

Usually rare fiilvus pallens

Not so marked 6

6. Mesonotum with four silvery lines, the outer pair curved to form

a lyre-shaped marking (fig. 19, A). The yellow fever mosquito

aegypti

Mesonotum without such lines (indefinite curved lines sometimes

present in canadensis) 7

7. Tarsal segments ringed only basally; fifth hind tarsus not all white 8

Tarsal segments ringed basally and apically; fifth hind tarsus all

white (except in canadensis mathesoni) 10

8. Wing scales all dark, narrow; white rings on tarsi narrow, no

wider than the diameter of segment; basal abdominal pale bands

with an inverted V-shaped notch in the posterior margin on

apical segments vexans

Wings with brown and white scales intermixed ; tarsal white rings

wider than diameter of segment, at least on hind feet, abdominal

bands not notched 9

9. Thorax with white lateral patches on anterior half; wing scales

very broad grosshecki

Thorax without white lateral patches; wing scales narrow. Recorded only once from Mississippi Stimulans

10. (7)4 Abdominal tergites almost entirely pale-scaled except for

dark, quadrate, submedian spots; wing scales mixed black and

white dorsalis

Abdominal tergites with only the usual white basal bands or lateral

spots ; wing scales all dark 11

11. Mesonotum pale, with a well defined, median dark stripe. Breeds

in rock holes atropalpus

Mesonotum golden or dark brown, without a median dark stripe 12

12. Mesonotal scales golden brown, with some pale scales often forming faint or indefinite lines or stripes; last hind tarsal segment

all white; segments 1 to 4 with fairly broad basal and apical

white rings canadensis canadensis

Mesonotal scales mostly dark brown, with indefinite stripes of

pale scales; last two hind tarsal segments typically entirely dark,

segments 1 and 2 with narrow basal and apical rings, segment 3

ringed basally only; intergrade specimens with the last hind

tarsal segment entirely or partially white, or white on one side

canadensis mathesoni

13. (1) Abdominal segments with dorsal white bands or median triangular spots •/•;•. 1 }n Abdominal segments with only lateral white spots i ^

4 Couplet 7 is the one from which couplet 10 leads. Similar numbers are

given throughout the keys for convenient reference when there is a skip

of more than one couplet.
62 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

14. Abdomen with large median triangular white patches at the base

of the tergites ; mesonotum covered entirely with yellowish sea es,

or with a median longitudinal stripe of somewhat darker scales.^^^^^^^

Ab^Smind^segments'with'basal'white bands; mesonotum not ^^

3.S 3.boVG •••••• •_

15. Mesonotum uniformly dark-scaled • .'V-;::*'.V* .^^^^^'^^^

Mesonotum with bicolorous markings, (sometimes indistinct m ^^

16. Mesonotum 'with a* broad median 'stripe'of yellowish or brownish

scales and silvery scales laterally; dark portions of femora

speckled with white • . • • • ; • • • • jticticus

Mesonotum with golden scales divided by a pair of narrow submedian longitudinal stripes, and with narrow dark stripes

laterally: dark portions of femora not speckled with white

South Florida • • • .•.;•. v ; ' ' /''''^'^''

17. (13) Mesonotum with a median longitudinal stripe or paten ol

silvery scales . ; • • • ; * ' ; '^ * * *ii * oo

Mesonotum with a median dark stripe and silvery scales laterally ZZ

18. A broad median patch ending just back of middle of mesonotum,

the silvery area wider than the dark area on each side (fig.

19^ ^) 19

Median stripe extending full length of mesonotum, usually narrower (somewhat variable in A. diipreei) than the dark lateral

areas (fig. 19, C) • • • • • • • • 20

19. Two species difficult to distinguish as adults except by slight differences in male characters.

Male terminalia:

Claspette with a flattened apical filament having one long

and several small backward-pointing spines on the inner

margin. Fairly common species in the Southeast infirmatus

Claspette filament with only one retrorse spine. Taken once

on the Florida Keys scapularis

20. Occiput with a narrow median line of pale lanceolate scales, bordered by broad white and dark scales; small species (wing about

2.5 mm.); mesonotum of male with a very broad stripe or

entirely silvery scaled; male claspette with a slender stem and

filament, the latter slightly longer than the stem, smoothly

tapered to a sharp point dupreei

Occiput with a wide median area of pale, lanceolate scales,

bordered by broad, appressed, mostly dark scales; medium-sized

species (wing about 3.5 mm.) ; male not as above 21

21. Two species difficult to distinguish except by male characters.

Male terminalia:

Stem of claspette long, greatly enlarged in middle, and

densely hirsute; filament very short, stout atlánticas

Stem of claspette long and slender, slightly pilose on basal

half; filament about one-third length of stem, with a

pointed, slightly hooked tip tormentor

22. (17) Mesonotum with a pair of broad submedian stripes of yellowish-white scales; dark centrally and laterally (fig. 19, £") . . trivittatus

Mesonotum with a prominent patch or stripe of pale scales on

each side, dark centrally 23

23. Posterior pronotum densely covered with broad, appressed, white

scales; mesonotum with silvery-white scales along sides, the

median area broadly dark-scaled (fig. 19, F). A common woodland species, breeding in tree holes trisei^iatus

Posterior pronotum with lanceolate scales, not closely appressed,

mesonotum with broad lateral patches of yellowish-white scales

anteriorly, reduced to narrow stripes posteriorly by an abrupt

widening of the median dark-scaled area (fig. 19, B). Breeds in

stump holes, usually rare thihaulti

LARVAE (FOURTH INSTAR)

1. Upper head hair multiple, lower hair double or multiple (fig. 12) ;

anal segment not completely ringed by the sclerotized plate . . , .' 2
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 63

4.

5.

6.

9.

10.

Either the upper or lower head hairs single (both pairs single

unless stated otherwise) ; anal segment with the plate complete

or incomplete 7

Pectén with one or more distal teeth more widely spaced than the

others (fig. 15, A) ; comb of few scales in a single irregular or

partially double row 3

Pectén with evenly spaced teeth; comb scales in a triangular patch 4

Upper head hair with three to five branches, lower usually with

two or three; lateral abdominal hairs double or triple on segments 3 to 5 vcxans

Upper and lower head hairs with more than four branches;

lateral abdominal hair single on segments 3 to 5 cinereus

Lower head hair double 5

Lower head hair with three or more branches 6

Comb scales with a long apical spine, thornlike sticticus

Comb scales rounded, fringed with subequal spinules grossbecki

Upper lateral hairs or both pairs on abdominal segments 1 and 2

multiple; antenna nearly as long as head; comb scales with the

apical and preapical spines strong, the others progressively

smaller thibaulti

Upper lateral hairs on segments 1 and 2 double, the lower single

or double ; antenna much shorter than head ; comb scales fringed

apically with subequal spinules canadensis canadensis

canadensis viathesoni

(1) Anal segment not completely ringed by the sclerotic plate .... 8

Anal segment completely ringed by the plate 12

Pectén with the distal teeth unevenly spaced atropalpus

Pectén with evenly spaced teeth 9

Antennal hair single (rarely forked) ; comb of few (8 to 15)

scales in a single or partly double row 10

Antennal hair multiple; comb of many scales in a patch 11

Comb scales with a long apical spine and a few much shorter side

spines (fig. 21, C) head hairs and preantennal hair single; tuft

of air tube with three or more branches aegypti

0.02Smm 0.025mm

M & A 14061

FIGURE 21. Enlarged comb scales of Aedes larvae: A, sollicitans; By taeniorhynchus; C, aegypti; D, triseriatus.

11.

Comb scales elongate, evenly fringed with numerous fine spinules

(fig 21, Z?) ; upper hair single, lower with two to four branches,

preantennal'hair multiple; tuft of air tube single or double

triseriatus

Upper head hair double; anal gills as long as the segment; comb

scales with a long apical spine twice as long as the subapical

^^- Stimulans JJaLL •
64 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Both head hairs usually single; gills very short; comb scales

rounded, fringed apically with subequal spinules dorsalis

12. (7) Pectén with one or more distal teeth more widely spaced and

extending on air tube beyond the ventral hair tuft 13

Pectén with evenly spaced teeth, none beyond the ventral tuft

(except in A. tormentor) 14

13. Comb scales in a patch, each scale rounded and fringed with subequal spinules ; lower head hair double or triple fiilvus pallens

Comb scales in a single irregular row, the scales with the apical

spine much longer and stouter than the others thelcter

14. Comb of few scales (less than 12) in a single row, the scales

pointed, thornlike 15

Comb of many scales in two or three rows, the scales thornlike,

or rounded and fringed 17

15. Anal gills as long as entire body, with prominent tracheae; lower

head hair double or triple dupreei

Anal gills much shorter than body; lower head hair single 16

16. Airtube with the ventral brush inserted before end of pectén;

anal gills all of equal length tormentor

Ventral brush of air tube beyond pectén; one pair of anal gills

longer than the other atlánticas

17. (14) Comb scales rounded apically and fringed with subequal

spinules, or the apical spine less than twice as long as the

adjacent ones ; body pilose 18

Comb scales thorn-shaped, or with the apical spine much longer

and stouter than the others; body glabrous (except in A. irifirmatus) 21

18. Anal gills shorter than anal segment, budlike (but may be longer

when the larvae develop in fresh water) ; body sparsely pilose

taeniorhynchus

Anal gills longer than segment 19

19. Body densely pilose, or spiculate; gills about twice as long as anal

segment and about equal in length to the ventral brush. Florida

Keys ....... scapularis

Body sparsely pilose; anal gills otherwise 20

20. Anal gills less than twice as long as segment and shorter than the

long hairs of ventral brush; comb scales fringed with fairly small

spinules, the apical one slightly longer than others; air tube

2.5-3:1. Southern Florida tortilis

Anal gills two to three times as long as segment, longer than hairs

in ventral brush; apical spine of comb scales stout, fairly long

the preapical pair about two-thirds as long; air tube about 2:l!

Rare in southeast, not reported in Florida trivittatus

21. (17) Anal gills shorter than segment, budlike; air tube about 2:1

Anal gills longer than segment; air tube 2.5-3:1 so ici am

22. Body glabrous; dorsal preapical spines of air tube (¿¿15 *i i

longer than apical pectén tooth ]_ ' mitchellae

Body sparsely pilose; preapical spines of air tube not more'than

half as long as apical pectén tooth infirmatus

AEDES AEGYPTI (L.)

(Syn., Stegomyia fasciatus F., Culex calopus Meig C araenteus

Poir., etc. The yellow fever mosquito.)

A rather small, dark mosquito with conspicuous rings on the

Z\^¡íl.T¿\tl't''' '^''^'^ '' ^"^^^^ white "scales on

This species is the most thoroughly domesticated of any of

the mosquitoes and apparently greatly prefers the blood of man

to that of other animals. It breeds almost exclusively in aîtifiS

water containers in the vie n ty of dwellings or in thl ntrLn

themselves The larvae are found ocSn! ly'^inVee holf and

similar natural collections of water, but, so far as known never
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 65

in ground pools except under unusual circumstances. In Orlando,

Fla., the writers found the larvae abundant at times in the

underground street catch basins, which, although partly filled

with sand and dirt, were lined with brick or concrete. Fairly

clean water is preferred, and sewage-polluted water in wooden

or concrete cesspools is not a favorable breeding medium. The

larvae mature in from six to ten days under favorable conditions

in the summer, but development is prolonged in cooler weather.

The eggs are usually laid on the sides of the receptacle just

above the water, but may be deposited directly on the water

surface. It appears that places having solid material at the water

line are preferred, although in the laboratory they oviposit

readily on moist filter paper. The embryos mature in from two

to four days, depending on the temperature, after which hatching may take place when suitable conditions occur. The eggs

may be kept dry and will remain dormant for several months,

yet hatch quickly when flooded. Under favorable summertime

conditions the complete cycle from egg to adult may be completed

in ten days.

The adults are abundant during the summer in cities and

towns throughout the South and are troublesome house pests.

Biting occurs largely in the daylight hours, especially early in

the morning and late in the afternoon, and the females seem

able to gain entrance even into well-screened houses. They are

wary biters and are especially annoying about the ankles. Aedes

aegypti is thought to have been the only species involved in the

epidemics of yellow and dengue fevers in the United States,

although other species in other countries have been proved

capable of transmitting both these diseases. Experimentally, A.

aegypti is able to transmit the viruses of the three most important strains of encephalitis found in the United States. It

has been found capable of transmitting the virus of fowl pox, and

is an eflScient host for several of the species of Plasmodium infecting birds.

The adults have been kept alive in the laboratory for several

months, and in the summer they probably live longer than any

other of the southern species. They are very susceptible to cold,

however, and are said to die at temperatures below about 40^^ F.

Although the eggs are more resistant, the species probably does

not overwinter in the United States except in the extreme southern part where breeding may be continuous throughout the year.

Each summer it becomes widely dispersed into territory farther

north, probably transported in trains, boats, et cetera. The adults

are strong fliers but the usual flight range is considered to be

not more than a few hundred feet.

The control of this species is discussed in the section on

mosquito control. ^ .xi. i. ^ i,-^

ADULT Tips of female palpi white, and clypeus covered with broad white

scales Occiput with a narrow median and lateral stripes of white separated

bv broad patches of dark scales. Mesonotum with a pair of fine submedian

lines of silvery white scales extending back of middle, broader lateral lines

curved around the fossae and continued to posterior end of scutum forming

a Ivre-shaped pattern; prescuteUar space with a border of white scales.

Lobes of the scutellum white-scaled, and patches of white on sides of thorax.

Abdominal segments with narrow basal white bands and long lateral spots.

Wing length about 3 mm., the scales all dark. Front and mid tarsi with
66 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

vt^ Ka=al rinps on segments 1 and 2 only; hind tarsi with wide

a ints'*o?selLe"tf^"to"3%egment 4 nearly all white except for a

"TZA^^^inttLfhS: 3fniinïieT'ùSer and.lower head hairs and LARVA.—Antennai nan s , ^^^^ .^ ^ g. ^^ ^^^^^¿ ^^^^ -^

^iralon^ceXaU^^ sevfral much shorter lateral spines (fig. 21, C).

A- f?.h!'^hniii 2-1 wit^ a ventral tuft of two or more branches, usually

thrt' AnalLlmlntL by the plate, the gills about twice

as long as the segment, the tips bluntly rounded.

AEDES ATLANTICUS D. and K.

(Syn., Ochlerotatus serratus Coq. (not Theob.), in part)

This mosquito is recognized by the broad median white stripe

extending the full length of the thorax (fig. 19, C). The tarsi are

unhanded. It breeds in temporary ram pools. The writers have

taken the larvae in some numbers in Louisiana and Florida m

shaded woodland pools, associated with other woodland species.

In southern Mississippi they were taken m clear grassy pools

(225) On Long Key, Monroe County, Fla., their breeding place

was a'shallow pool in marl soil filled with a thick growth of purslane (Sesuvium) associated with larvae of Aedes thelcter and other

species of Aedes and Psoropkora (805), The female is a severe

daytime biter in and near woods, and is usually associated with

A. infirmatus and other woods species.

This mosquito is common in the Southeastern States and has

been recorded in all except Tennessee, where it probably will be

found also. Its range extends from Texas to Missouri and eastward

to New York.

ADULT. Proboscis, wings, and tarsi all dark-scaled. Occiput with a large

median area of pale lanceolate scales bordered by broad flat scales, mostly

dark. Mesonotum with a broad median stripe of silvery white scales extending

the full length of the scutum, narrower than the dark area on each side

(fig. 19, C). Lower mesepimeral bristles absent. Wing length about 3.5 to

4 mm. Abdominal segments unhanded, with basal white lateral spots visible

from above on the posterior segments. Stem of male claspette long, greatly

enlarged in middle, and densely hirsute; filament stout, much shorter than

stem, the stem without a bristle. The female is indistinguishable from A.

tormentory and is also very similar to A. dupreei except for its larger size and

slight difference in the scaling of the occiput and first hind tarsal segment.

The male claspette is distinctive.

LARVA.—Antennai and preantennal hair multiple; upper and lower head

hairs both single, sparsely spiculate. Body glabrous, lateral abdominal hair

single on segments 3 to 5; comb of only 5 or 6 scales in a single row, each

scale thorn-shaped and finely fringed laterally. Air tube about 2:1, the pectén

with evenly spaced teeth, the ventral tuft multiple, inserted beyond pectén.

Anal segment completely ringed by the plate, the lateral hair single; gills

with the upper pair longer than the lower, pointed, about three times as

long as the segment.

AEDES ATROPALPUS (Coq.)

This rare, medium-sized species has white scales laterally on

the mesonotum and a broad median stripe of dark scales, in this

respect somewhat resembling Aedes triseriatus. The tarsal segments are ringed at base and apex. It breeds normally in rock

holes along mountainous stream beds, sometimes in rocks away

from the streams. Larvae were collected from pot holes in an

unfinished concrete foundation of a power dam in Georgia (299).

In Texas, they were taken in two tree holes in hackberry trees
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 67

and once in an artificial water container (IW- In the rock holes

this author found that the eggs might be laid on the water surface

during the summer, but were always laid on the sides of the rock

holes by the last generation in the fall. Larvae of Culex tarsalis

frequently were associated with afroqmlpus, and with numerous

other species occasionally. The adults are said to be annoying but

remain near their breeding places, and since these are of very

local and scattered distribution, the species is considered to be of

little importance. A blood meal is not required for egg production.

Atropaljms was previously reported (193) from Arkansas, Tennessee, and North Carolina. There are additional State records

from Baldwin and Rabun Counties, Georgia (299), and Oconee

County, South Carolina (65). It is widely, but locally, distributed

throughout the Eastern and Midwestern States to Minnesota,

Texas, and New Mexico.

ADULT.—Proboscis dark. Occiput with a median patch of narrow whitish

scales, broad white ones laterally enclosing a patch of dark scales. Mesonotum

with lateral patches of yellowish-white scales anteriorly, separated by a

broad median dark stripe which widens to almost the complete width of

scutum back of the fossae; prescutellar space surrounded by pale scales.

Lobes of scutellum with narrow, pale scales. Lower mesepimeral bristles

absent. Wing length about 3.5 mm., the scales dark except for a short line of

white at base of costa. Hind tarsi with basal and apical white rings on

segments 1 to 4, segment 5 entirely white; fore and mid tarsi with narrower

rings on the first three segments, segments 4 and 5 and tip of 3 usually all

dark. Abdomen with broad basal, segmental white bands.

LARVA.—Antennal tuft double or triple; upper and lower head hairs single.

Body glabrous. Lateral abdominal hairs multiple on segments 3 to 5; comb

scales in a patch, each narrowly rounded and fringed with subequal spinules.

Air tube 2:1, the tuft large and multiple, the pectén with two or three of

the apical teeth more widely spaced than the others and extending beyond

the tuft. Anal segment with a small plate covering only about half the

segment, the lateral hair single, not inserted in plate. Gills more than twice

the length of segment, bluntly pointed.

AEDES CANADENSIS CANADENSIS (Theob.)

The thorax of this species is golden-brown, usually with indistinct lines of white scales. The tarsal segments have fairly

broad apical and basal rings. The larvae develop principally in

woodland pools, and in Tennessee were found to prefer small,

shaded pools with a bottom of dead leaves (287). They are also

found in roadside puddles, spring-fed pools, cranberry bogs, pools

in open sphagnum bogs, wooded swamps and open meadows (213).

In Tennessee the larvae were associated chiefly with Aedes vexans

(290) In southern Mississippi they were sometimes exceedmgly

abundant in water pits, often associated with Culex restuans^nd

C apicalis (= C. territans) ; also found in small deep puddles

along intermittent streams and in receding river flood waters

associated with Aedes sticticus, A. vexans, and Psorophora ferox

(225) The over-wintered eggs hatch in the early spring and

there seems to be only one generation a year, although larvae are

sometimes found in small broods during the summer, presumably

from delayed hatchings. The adults are sometimes numerous and

attack readily but do not disperse far from their breedmg places,

mïspecies has been recorded froni all the Southeastern S^^^^^^^^^

(193) It is widely distributed in the United States, except the

extreme Southwest, and occurs in Canada.
68 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

ADULT.—Proboscis dark. Occiput with a median area of narrow white

scales, submedian areas of darker narrow scales and lateral patches of

broad white scales. Mesonotal scales mostly brownish or golden-brown, paler

around antiscutellar space and anterior lateral margins, often with faint

submedian lines on posterior half or with indistinct curved lines around

fossae. Lobes of scutellum with narrow pale scales. Lower mesepimeral

bristles absent. Wing length about 3.5 mm., dark scaled. Hind tarsi with

basal and apical white rings on segments 1 to 4, segment 5 all white; front

and mid tarsi with narrower apical and basal rings on segments 1 to 3,

sometimes on 4. Abdomen with basal segmental white bands joined to the

lateral spots, sometimes reduced in width or lacking on some segments.

LARVA'—Antenna about half the length of head. Antennal and preantennal

tufts multiple; upper head hair with 4-10 branches, the lower with 4-8.

Body glabrous. Upper lateral hairs on abdominal segments 1 and 2 double,

the lower single or double; lateral hair on segments 3 to 5 usually double.

Comb of segment 8 a large patch of scales, each rounded and fringed with

subequal spinules. Air tube about 3.5:1, the ventral tuft multiple, inserted

beyond pectén. Anal segment not completely ringed by plate, the lateral

hair single, gills pointed, as long as segment or longer.

AEDES CANADENSIS MATHESONI Middlekauff

This species differs from the type form in its generally darker

color and a reduction of white on the tarsi. It was described by

Middlekauff (226) as a new species from Florida. The holotype

was a female from Kissimmee, with paratypes from McDill Field,

Pinecastle, and Camp Murphy. The larva was described by Rings

and Hill (265) from specimens collected in deep fox holes at Camp

Blanding, Starke, Fla., and Camp Gordon, Augusta, Ga. These

fox holes were shaded Í3y overhanging grass and contained water

about 8 inches deep. Further taxonomic study by these authors

(266) led them to assign it to a subspecific status as a ''melanistic,

geographical variation'' of canadensis. Its present known distribution is entirely southern, and they point out that, of the

climatic factors that may influence geographical variations, the

most important one affecting pigmentation is considered to be

temperature. Besides northern Florida and Georgia, a distribution map in the 1948 article shows its occurrence in Alabama and

South Carolina.

ADULT.—Mesonotal scales mostly dark brown, with a median fine line of

very small golden brown scales; an elongate stripe of yellowish-white scales

on the anterior angles and similar patches laterally at middle, sometimes

with faint or indefinite curved lines around the fossae; prescutellar space

with a few yellowish-white scales. Wings and legs with purplish scales.

Hind tarsi typically with narrow white rings at base and apex of segments

1 and 2, and base of 3, the rest of the segments entirely dark. Abdomen

purplish black without segmental bands but with small triangular lateral

white spots. In the type form, the white areas on the tarsi are more extensive

andi the fifth hind tarsal may be partially or entirely white. The mesonotum

lacks the black scales, frequently has fairly distinct curved submedian pale

lines, and the abdomen frequently is banded with white. Rings and Hill ('^66)

have illustrated some of these intergradations. The male terminalia of'^ the

two subspecies are indistinguishable.

v^ÙrA'^^^^ larvae of the two forms are also very similar except for

slight differences in the number of branches of the upper and lower head

hairs. In mathesoni branches of the upper hair range in number from 6 to 14

AEDES CINEREUS MPíO^

(Syn., A. fuscus O. S., etc.)

Except for broad white bands on the abdomen this species is
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 69

unornamented. It is the only representative in the United States

of subgenus Aedes, in which the male palpi are very short. Larvae

v^ere collected in Mississippi in floodv^ater pools in the early

spring, associated v^ith A. vexans {2i5). Thibault (303) reported

that the adults were numerous in weeds and grass and about

dwellings near Scott, Ark., but did not bite. These seem to be the

only published bits of information on the habits of cinereiis in

the Southeast. Other records of its occurrence in this area are

from light-trap collections. In the northern and western States

it breeds in woodland pools and a variety of other habitats in

association with numerous other species. It hibernates in the

egg stage and is thought to have a single annual generation. The

female is said to be a pest in some places.

This species previously was recorded in the Southeast from

Georgia, Arkansas, and South Carolina, with a doubtful record

for Louisiana (193). Collections during and immediately following World War II have added Mississippi (2J^5, 225), North

Carolina and Tennessee (228), Alabama (230), and Florida

(53, 66).

ADULT.—Proboscis dark; palpi very short in both sexes. Occiput with a

narrow median stripe of narrow pale scales, and submedian patches of dark

broad scales. Mesonotum with fine brown scales except around the prescutellar space, where the scales are pale. Scutellum with the lobes palescaled. Lower mesepimeral bristles absent. Wing about 3 to 3.5 mm., the

scales dark. Abdomen usually with basal white segmental bands that widen

laterally, the bands sometimes incomplete.

LARVA.—Antennal tuft and upper and lower head hairs all multiple. Body

glabrous. Lateral abdominal hairs single on segments 3 to 5. Comb of about

15 scales in a partially double row, each tapered to a long point and very

finely fringed nearly to tip. Air tube about 4:1, the pectén with one or more

of the apical teeth more widely spaced than the others, the ventral hair

small, usually with three or four branches, inserted beyond the pectén.

Anal segment not completely ringed by plate, the lateral hair very small,

double, or triple. Gills nearly twice as long as segments, pointed, one pair

slightly longer than the other.

AEDES DORSALIS (Meig.)

There are only two records of occurrence for this species in the

Southeast—at Delta, La., (163) and at Como, Miss., (230). From

the latter locality a single female was collected in 1945, and the

identification was confirmed by Dr. Alan Stone. It is evidently

extremely rare and of no importance in this area. It is widely

distributed in the northern and western parts of the United States

and Canada, as well as in Asia and Europe. In many places,

particularly in the western United States and Canada, it is the

dominant species and an important pest of man and animals.

It breeds in both brackish and fresh water. Several types of virus

of the encephalitides have been isolated from wild-caught A.

dorsalis in the Western States and Canada.

ADULT.—Proboscis and palpi speckled with white. Occiput with a large

median area of pale scales, bordered laterally with dark. Mesonotum with

yellowish-white scales laterally and posteriorly, a median dark stripe of

variable width extending to about the posterior third. Scutellar lobes palescaled. Lower mesepimeral bristles present. Wing about 4 mm., extensively

pale-scaled, speckled with dark. Femora, tibiae and first tarsal segments

well speckled with pale scales; segments 1 to 4 of hmd tarsi with both basal

and apical white rings, segment 5 all white; fore and mid tarsi with similar

very narrow rings on first three segments, the last two usually all dark.
70 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Abdominal tergites almost entirely pale-scaled except for quadrate, submedian dark spots on most of the segments.

LARVA.—Antennal hair multiple; upper and lower head hairs single. Body

glabrous. Comb scales in a patch, each rounded and fringed apically and

laterally, the spinules at apex fairly long, subequal. Air tube about 3:1,

tuft beyond pectén; distal teeth of pectén evenly spaced. Anal segment with

the plate covering about half the segment, the lateral hair single. Gills very

short, less than half as long as segment, bluntly pointed.

AEDES DUPREEI (Coq.)

This rare mosquito has a broad, median mesonotal stripe of

silvery-white scales similar to that of A. atlanticus, but it is a

considerably smaller mosquito and the scales on the occiput differ.

It breeds in temporary rain pools but the larvae are seldom

collected, ov^ing in part to their habit of hiding in the leaves and

trash at the bottom of the pools, wehere they are able to remain

for long periods without coming to the surface. They are easily

recognized by their extremely long and sinuous anal gills. The

adults are taken in light traps. Apparently nothing is known

about their biting habits or host preferences.

With records of occurrence in Tennessee (230, 290), this mosquito has been recorded in all of the Southeastern States. Its range

extends from Texas, Kansas, and Illinois to New Jersey.

ADULT.—Proboscis and M^ing unmarked. Occiput v^^ith a narrov^ median

line of pale lanceolate scales, bordered by broad pale scales, and with dark

scales laterally. Mesonotum with a broad stripe of silvery white scales

extendmg the full length of the scutum, usually somewhat narrower than the

dark area on each side. Lower mesepimeral bristles absent. Wing length

about 3 mm. Mesonotum of male entirely silvery-scaled or with a very broad

median stripe. Tarsi all dark except for pale scaling on the under side of the

ftrst hmd segment. Abdominal segments unhanded but with lateral basal

white spots. Male claspette with a slender stem and filament, the latter

slightly longer than the stem and tapered to a sharp point; stem of claspette

with a bristle arising from a raised tubercle near base.

I. ^\^-l^^?^^^^^^ v^^y similar to those of Aedes atlanticus and A, tormentor,

but differ m their smaller size and in slightly different scaling of the occiput

and ftrst hmd tarsus. The males have more extensive pale scaling on the

thorax, and the claspette is distinctive.

o.íiíl'^^V'^'^^^i^^^'^?^ ^""^ immediately recognized by extremely long, sinuous

WPVÄT^- ^5^ P^'^antfï^nal hairs usually double, sometimes triple;

snTculatP TinH^Î'''i'^r^^^ T Í^'P\'^ t^/ ^PP^^ ^^^^^^^ ^^^^ ^ Pl^ or sparsely rwl ^ ;Ä glabrous. Lateral abdominal hairs single on segments 3 to 5

Comb, a single row of scales, each thorn-shaped and finely frineed laterallv

Air tubes about 3 5:1, tuft multiple, inserted beyond X pectén the^Ä

with evenly spaced teeth. Anal segment completely ringed by th¿ pllte the

darrirSar^^'- ^"'^ ^^"' "''"' ^' ^^^^ ' ^ '^'"'^ ^^^>^' wiih%?om!nent!

AEDES FULVUS FALLENS ROSS

(Syn., A. bimaculatiis (Coq.) in part; A. fulvus Dyar

(not Wied.) in part.)

This is a large, bright yellowish mosquito of striking appearance. The subspecies pallens was described in 1943 (^69) from

specimens taken in New Orleans in 1914. It was previously identl^

fied m the Southeastern States as bimaculatus, hut that spSes

was shown to be distinct and to occur only in Texas and MSO

The type form of fulvus is found in the AmericafTroiL aní^^

not known to occur in the United States. SubspeciSVa/^S. s

usually rare, but was encountered by the senio? aSof n c^^
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 71

siderable numbers on one occasion in 1914 in the vicinity of New

Orleans. The larvae v^ere found in temporary pools in dense woods

and the adults were fierce biters. Large numbers were reared

near Wilson Dam by Belkin (213), Thibault found it to be rare in

Arkansas but occasionally present about dwellings. With a record

of the occurrence of pallens in Tennessee (290), it is now known

to occur in all of the Southeastern States, and has also been

recorded from most of the immediately adjoining States, as well

as Illinois and Maryland.

ADULT.—Proboscis and palpi yellow-scaled, tipped with dark. Occiput

entirely covered with yellowish scales, narrow on the dorsal area, broad and

flat laterally. Mesonotum with both the scales and integument a bright

yellow, and two prominent, shiny, elongated black spots on the posterior

corners. Faint submedian lines along the middle of the scutum. Scutellum

with narrow brownish scales. Lower mesepimeral bristles absent. Wing

length about 4.5 mm. Costa, subcosta, and vein 1 yellow-scaled from base to

tip of the subcostal vein. Femora, tibiae, and first two tarsal segments

largely yellow-scaled except for dark rings at tip, the last three tarsal

segments all dark or nearly so. Abdomen with prominent basal yellowish

bands that widen laterally, segment 7 completely yellow and segment 6

largely so.

LARVA.—Antennal and preantennal hairs multiple; upper head hair single,

the lower double or triple. Body glabrous. Lateral abdominal hairs on

segments 3 to 5 single. Comb of many scales in a patch, each rounded and

fringed with subequal spinules. Air tube about 2:1, the pectén with the last

tooth widely separated from the others and attached at about the apical

third of tube; tuft large, multiple, inserted before the end of pectén. Anal

segment ringed by plate, lateral hair single or double, gills about 1.5 times as

long as segment, pointed.

AEDES GROSSBECKI D. and K.

This rare northeastern species has been reported from a few

scattered localities in the Southeast. The mesonotum has lateral

patches of silvery white scales on the anterior half, somewhat

similar in this respect to A. thibaulti, but the tarsi have white

rings and the wings and legs are speckled. It breeds in woodland

pools, and in the North is said to be occasionally fairly numerous

and to attack man. In Louisiana, larvae were collected in January

and April, and were associated with Ciilex restuans, Aedes canadensis, and Culiseta inornata in semipermanent pools. It was

thought to have only a single annual generation (332),

A. grossbecki was previously listed (193) from Mississippi,

Louisiana, and Arkansas. There is an additional record from

Tennessee (290).

ADULT.—Proboscis and palpi sprinkled with white. Occiput with a large

median patch of narrow white scales. Mesonotum with a patch of silverywhite scales on each side anteriorly, reduced to narrow lateral stri^^^^

posteriorly, somewhat similar to the markings of A. thibaulti (fig. 19, V).

Prescutellar space surrounded by white scaling. Scutellum with narrow

white scales on the lobes. Lower mesepimeral bristles usually present but

may be absent. Wing length about 4.5 mni., the scales very broad, d^ark

intermixed with white. Femora, tibiae, and first tarsal speckled with white,

tarsal segments with basal white rings. Abdomen with broad, ^3sal segmentai, white bands. The broad wing scales and silvery-white markmgs on

the thorax distinguish this species from A. stimulons. Ar..M^ fVi.

LARVA.-Antennal hair multiple; lower head hair "^^^11^^^.^^^^^^^^^

upper usually triple. Lateral abdominal hairs double on segments 3 to 5

Comb scales in a patch, each rounded, the apical spinule long, the subapical

Ä about two-tMrds ks long; the others progressively shorter. Air tube

fbout 3 1 the tuft multiple attached beyond pectén Anal segment not

ïna by plate, the^l^^^^ single. Gills pointed, shorter than segment.
72 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

AEDES INFIRMATUS D. and K.

Aedes infirm^atus is recognized by the large silvery-white patch

on the anterior half of the thorax. The tarsi are unhanded. It

breeds in temporary woodland or open grassy pools, and at times

becomes very abundant and annoying. In common with most of

the other woodland species, the females attack readily during the

day in or near woods, and at night they may be encountered in

the vicinity of dwellings. They may enter houses to bite when

excessively numerous. At Yukon, near Jacksonville, Fla., out of

a total of nearly 125,000 mosquitoes collected in light traps during

the years 1948-1953, about 6,000 were infirmatus, with an equal

number of Psorophora confinnis. These two were the most numerous of the pest mosquitoes taken by Knight (202). In a study of a

single heavy brood that emerged in April 1953, Knight (20S)

found the adults congregating in numbers about shaded doorways, in shrubbery along buildings, in open garages, and every

protected building corner. They seldom entered the buildings but

were found in swimming-pool shower rooms. It was of only

moderate importance as a biter after dusk. Sparse breeding was

reported to occur at that latitude in the winter months.

This species was previously reported from all but two of the

Southeastern States (193), and recent records for Alabama (228,

230), and Tennessee (290) complete the list for these States.

Of the adjoining States, the species has been reported only from

Texas. In Louisiana, the virus of western equine encephalitis was

twice recovered from pools of wild-caught infirmatiis, indicating

its potential importance as a vector (196),

ADULT.—Proboscis and tarsi unmarked. Occiput with a large median area

of pale narrow scales and smaller submedian patches of broad yellowishwhite scales. Mesonotum with a wide conspicuous patch of silvery-white

scales on anterior half, wider than the dark lateral patches and extending:

slightly beyond middle of scutum; scales on posterior half dark brown, the

prescutellar space bordered with white. Mid lobe of scutellum with a sm-^ll

patch of white scales, the remainder dark. Wing 3.5 to 4 mm. in length, the

scales all dark. Abdomen unhanded, but with lateral basal white spots visible

from above on some segments. Male claspette with a backward pointing

(retrorse) spine about the middle of the filament, but without additional

bristles as found in A. scapularis.

LARVA.—Antennal and nreantennal tufts multiple. Upper and lower head

hairs single. Body sparsely pilose. Lateral hairs sine^le on se.8:ments 3 to 5.

Comb scales in a patch, each thorn-shaped, with the lateral spinules less

than half as long as the apical spine. Air tube about 2,5:1, the pectén with

evenly spaced teeth extending to about the middle of the tube, the tuft

niultinle, mserted beyond pectén. Anal segment completely ringed by plate

the lateral hair single or forked. Gills nearly twice as long as the seement'

pomted. ^ '

AEDES MITCHELLAE (Dyar)

This comparatively rare species resembles Aedes soUicitayis

with wide rings on the proboscis and tarsi, and a median longitudinal stripe on the abdomen. It lacks the white ring in the middle

of the first hind tarsal segment, and the wings are not speckled

It breeds in open, temporary pools, but not, so far as known in

dense woodland. In central Florida, where it frequently was 'encountered m the piney flatwoods country, its typical breeding

places were shallow depressions, sparsely overgrown with tufted
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 73

grass and weeds in rough pastureland surrounded by scattered

pine woods and palmettos. These flat depressions were often up

to an acre or more in size, and the floodings were usually very

transient. Breeding was never very heavy. When the depressions

were dry, larvae could be obtained from sod samples containing

tufts of grass when placed in containers and covered with water.

The females were encountered near the breeding places and were

rather severe biters. They were reported to become quite annoying in the woods near Gainesville, Fla., (A. J. Rogers, personal

communication). The adults have been taken also in light traps.

In Mississippi, Michener (225) found the larvae in shallow pools

in springhead areas, which held water for fairly long periods and

usually contained grass or other emergent vegetation. While the

species has not been found breeding in salt marshes, it has been

taken most frequently in the Coastal Plain. It has also been

recorded from several inland States.

This mosquito was previously reported from all seven of the

Southeastern coastal States (193) and has since been taken in the

two inland States of Arkansas (5^), and Tennessee (290). Outside of this territory it has been reported from Texas and New

Mexico to Illinois and New York. The virus of eastern equine

encephalitis has been recovered from pooled lots of mitchellae

collected in Georgia (176),

ADULT.—Proboscis with a broad white ring. Mesonotum golden in color as

in sollicitans. Wing about 4 mm. in length, the scales all dark. Fore and

mid tarsi with narrow white basal rings on segments 1 to 3, the hind tarsi

with wider rings on segments 1 to 4 and segment 5 all white. Dark-scaled

areas of femora and tibiae speckled with white. Abdominal tergites with narrow, basal white bands and central pale patches forming an almost continuous

longitudinal stripe, as in sollicitans.

LARVA.—Lower head hairs single, the upper, usually single but sometimes

double. Antennal tuft multiple, the branches spiculate. Body glabrous. Lateral

abdominal hairs on segments 3 to 5 single or double. Comb scales in a patch,

the single scale thornshaped. Air tube about 3:1, with a multiple ventral

tuft beyond the pectén. Anal segment completely ringed by the plate, the

gills slightly longer than the segment and somewhat pointed.

AEDES SCAPULARIS (Rondani)

(Syn., A. euplocamus Auct., not D. & K.)

This tropical species has a large patch of silvery-white scales

on the front half of the thorax and is identical in appearance with

Aedes infirmatíis. Its identification depends on slight differences

in the larva and male terminalia. The occurrence of A, scaqmlaris

in the Southeastern States is based on the collection of three

fourth-instar larvae from a rainwater pool on Key Vaca, Monroe

Co., Fla., in 1945, originally identified as A. euplocamits D. & K.

(232 252), It has not been found since and may not be established in that area, but if so, its occurrence would not overlap

the present known range of infirmatics, as the latter has not been

found on the Keys. The species is common in parts of the American

Tropics and has also been reported from the lower Rio Grande

Valley, Texas. It is susceptible to experimental mfection with the

virus of yellow fever. „ ^. . ^ ADULT.—Proboscis dark; occiput with a small median area of narrow pale

scales and larger submedian patches of broad, appressed, yellowish-white

scales Anterior patch of silvery-white scales on mesonotum similar to
74 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

that of Aedes infirmatus but somewhat wider, covering nearly the entire

width of the scutum; posterior part of mesonotum including the prescutellar

space and lobes of scutellum covered with fine golden-brown scales. Abdominal

tergites with more or less distinct median, basal, triangular patches of dingywhite scales, tending to be produced to posterior border on segments 5 to 7 ;

large lateral spots of pure white scales, hardly visible dorsally. In the males

of both scapularis and infírmatus, the claspette has a retrorse, or backwardpointing, spine about the middle of the filament, but in infirmatus there are

several small bristles in the angle formed by the spine, whereas these are

lacking in scapularis.

LARVA.—The body of scapularis is more densely pilose than that of

infirmatus, and the comb scales are rounded and fringed with subequal

spinules instead of thorn-shaped.

AEDES SOLLICITANS (Wlk.)

(The eastern salt-marsh mosquito)

This common salt-marsh mosquito has wide white rings on the

tarsi and proboscis, the thorax is golden-brown, and the abdomen

has a median longitudinal white stripe. It occurs in all types of

salt marshes along the Atlantic and Gulf Coasts and, except in

southern Florida, is usually by far the most important of the saltmarsh species. The larvae are found on the parts of the marsh

not reached by daily high tides, usually in potholes or depressions

of various sizes, but often over extensive areas of nearly level

marsh. The eggs are laid on almost any moist place in such

marshes and also in low grassy lands immediately adjacent. Hatching occurs when the land is covered with rainfall or with monthly

or seasonal high tides (spring tides), and enormous numbers

may be produced in a single brood. The kinds of vegetation growing on the marshlands vary a good deal with the elevation, some

being more tolerant of frequent floodings than others. With a

little experience, one can often determine whether a particular

place is a potential breeding ground by the kind of vegetation

present. The eggs can withstand long periods of drying, and the

percentage of hatching with each flooding is variable.

The females are fierce biters and strong fliers. They commonly

migrate in large swarms many miles from the coast, distances of

40 or 50 miles being not unusual, although the great majority

probably stop much closer. In Florida, specimens sometimes are

taken in the interior of the peninsula when broods have emerged

on the coast. The flight of migratory swarms begins just before

dark, and the numbers that may be encountered at this time are

almost unbelievable. The adults settle in the grass and shrubbery

during the day and are extremely annoying, often unbearably

so, to persons and domestic animals, attacking even in full sunlight. In the southern half of Florida the species may be found

throughout the winter, usually in small numbers, and more prevalent during the cooler months than in the summer. The relative

numbers there seem to vary considerably from year to year. The

distribution of the two species in Florida, and relative abundance

in a few other localities, is discussed in the notes on taeniorhynchus.

The virus of western and eastern equine encephalitis has been

transmitted experimentally with this species by several workers.

Its vector potential was rated as ^'excellent'' for the eastern type

and "good" for the western type by Chamberlain et al (72).
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 75

A. sollicitans has been reported from all the Southeastern States

except Tennessee (198). It occurs in all the Atlantic and Gulf

Coast States from New Brunswick to Texas, and has been reported from a number of inland States, particularly the Midwest,

where it is associated with salt water from oil or artesian wells.

In Illinois and Indiana it bred abundantly in areas intermittently

flooded with salt water from swimming pools or oil wells, and in

one pond flooded with acid waste from the shaft of a coal

mine {108). In one of the localities in Illinois (Dupo) the salinity

of the water in which breeding occurred was three times that of

average sea water {270), It has been reported as well from

Kansas, Nebraska, Missouri, North Dakota {311), and other

States. In Oklahoma the larvae were found in salt-water pools

which form at the edges of the Great Salt Plain near Cherokee

{275). The species occurs also in the West Indies.

ADULT.—Proboscis with a wide central white ring. Palpi white-tipped.

Occiput with a large patch of pale lanceolate scales bordered laterally with

a narrow line of dark scales. Mesonotum and scutellum with golden lanceolate

scales. Lower mesepimeron either bare or with a single bristle. Wing about

4 mm., with white scales mixed with the brown, the lateral or plume scales

narrow. Femora and tibiae speckled with white, the tarsal segments with

broad basal white rings, segment 1 with an additional broad ring in the

middle and segment 5 all white on all tarsi. Abdomen with basal segmental

white or yellowish bands and a median white stripe extending the full length.

A. mitchellae is the only other species in our area with a longitudinal stripe

on the abdomen, but it differs in having the wing scales entirely dark, and

the first tarsal segments are without a central white ring.

LARVA.—Upper and lower head hairs single, the antennal tuft multiple.

Body glabrous. Comb scales in a patch, the scales thornlike, fringed on each

side toward base. Air tube short, about 2:1, the ventral tufts multiple and

inserted beyond the pectén. Dorsal preapical spines about as long as the

apical pectén teeth. Anal segment ringed by the plate; the gills very short,

budlike, but may be somewhat longer in specimens that develop in fresh

water. The larvae, although very similar to those of A. mitchellae and A.

infirmatus, can usually be distinguished readily by the much shorter gills and

a somewhat shorter air tube, hifirmatus also differs in being sparsely pilose,

at least on the thorax, and having the dorsal preapical spines of the air

tube much shorter than the apical pectén tooth.

AEDES STICTICUS (Meig.)

(Syn., A. hirsuteron (Theob.))

This species is distinguished by having the sides of the mesonotum pale-scaled and with a broad central stripe of brown scales,

usually divided by a fine median line of yellowish scales ; the tarsi

unhanded. It is typically a floodwater mosquito, and occurs chiefly

in the spring. There is one generation a year {213). The females

may migrate several miles and are fierce biters. Very large broods

appear at times in some of the northern and western States and

Canada under flood conditions, and become a serious menace to

man and animals. In the South the species is widely distributed

but seldom abundant. Thibault {303) found the larvae in Arkansas

in temporary grassy pools in fields and thickets from March to

May, and in rain pools during the summer. The females attacked

in the woodlands but did not enter buildings. In Tennessee it was

reported to be relatively abundant. The larvae occurred m floodwaters and were taken associated with six other species, chiefly

A vexans {290). In southern Mississippi it was very troublesome

in the spring and was a serious pest within a radius of about 21/2
76 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

miles from the breeding place. The larvae occurred in shaded pools

left by high river water (225),

The species is widely distributed from coast to coast in the

United States and Canada, as well as in Europe and Northern

Asia. It was listed from all the Southeastern States {193), and a

number of records within these States have been added, following

more recent collections.

ADULT.—Proboscis dark. Occiput largely silvery-scaled, the scales narrow

on the dorsal area. Mesonotum with a broad central stripe of brownish or

golden scales, usually divided by a fine median line of yellowish scales. Sides

of scutum and antescutellar space with silvery-white scales. Lobes of scutellum

with narrow yellowish scales. Lower mesepimeral bristles absent. Wing

length about 4 mm., the scales dark. Dark portions of femora and tibiae

heavily speckled with white, the tibiae and first tarsals pale on one side for

nearly their entire length; tarsi not ringed. Abdomen with white, basal

segmental bands, widening laterally.

LARVA.—Antennal and preantennal tufts of larva multiple. Lower head

hair usually double, the upper with two to four branches. Body glabrous.

Lateral abdominal hairs double on segments 3 to 5. Comb scales in a small

patch, each thorn-shaped with fine lateral spinules. Air tube about 3:1, tuft

multiple, inserted beyond pectén. Anal segment not completely ringed by

plate, lateral hair single, gills about twice the length of segment.

AEDES STIMULANS (Wlk.)

{Syn.y A. Stimulans mississippii Dyar, etc.)

Aedes Stimulans is a speckled mosquito with broad white rings

on the tarsi and broad basal abdominal bands. The proboscis is not

ringed and the thorax is not definitely ornamented. It is a northern

forest and floodwater mosquito that has been recorded once from

Mississippi (Electric Mills). These specimens were named A.

Stimulans mississippii by Dyar in 1920, but the subspecies was

later dropped as a synonym and the species has not again been

reported from the South. It is widely distributed in the northern

States and Canada, and is an important woodland pest in some

places.

ADULT.—Proboscis and palpi dark, sprinkled with white. Occiput largely

pale-scaled. Mesonotum with light brownish scales often somewhat darker

in the median area except around prescutellar space. Lower mesepimeral

bristles present. Wing about 4.5 mm., well sprinkled with white- lateral

wmg scales narrow. Hind tarsi with broad basal white rins; on all segments

front and mid tarsi with narrower rings on segments 1 to 4, segment 5 all'

dark. Abdominal segments with broad basal white bands.

LARVA.—Antennal and preantennal hairs of larva multiple. Upper head

hair usually double the lower single. Body glabrous. Lateral hairs usually

double on abdominal segments 3 to 5. Comb scales in a patch, each rounded

and fringed, the apical spme long and stout, the preapical pair slightly over

half as long, and the others progressively shorter. Air tube about ^-1

ventral tuft multiple, inserted beyond pectén. Anal segment not cSeLy

segment "" ^ ' ^^''^^ ^^'' '^^^^"- ^'^^^ ^^^^^^^^ ^^^^^ as Tong as

AEDES TAENIORHYNCHUS (Wied.)

(Syn., Taeniorhynclms niger of Giles (not Theob ) • Culex

portoricensis (Ludl.) ; the black salt-marsh mosquito.)

This small black and white mosquito has sharp, contrasting

white rings on the proboscis and tarsi (fig. 22) somewhat ÍÍ^^^

rower than in Aedes soUicitans, It lacks the longZdS strip¡

on the abdomen and the central ring on the first tarsal segme^^^^^
THE MOSQUITOES OP THE SOUTHEASTERN UNITED STATES 77

M & A 14062

FIGURE 22.—Aedes taeniorhynchus, one of the salt-marsh mosquitoes.

It is the most abundant and troublesome species along at least the

southern two-thirds of the Florida coasts, which is also approximately the area where mangrove and pickleweed (saltwort)

form the predominant marsh vegetation (pi. 6). Farther north and

west it is usually outnumbered by A, sollicitans. From extensive

light-trap collection records obtained by the Florida State Board of
78 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Health, Provost {25Jf) states that the points at which the two species occur in about equal numbers are the mouth of the St. Johns

River on the Atlantic Coast and the mouth of the Suwannee River

on the Gulf Coast. North of this 50-50 line the salt marshes are of

the grassy-sod type {SjMrtina, Distichlis, Sjooroholis) while south

of this line they are of non-sod type (Batis, Salicornia, Sesuvium,

mangrove). At the Alabama-west Florida line, sollicitans predominated up to 50:1, whereas taeniorhynchus reached a ratio of

1000:1 on the Keys. While taenia rhyyichiis usually is not the

predominant species north of Florida, there are a few records of

collections in which they outnumbered sollicitans. At Charleston,

S. C, collections in four light traps, operated nightly over a period

of about 100 days in 1939, gave a total of nearly 10,000 taeniorhynchus compared with about 200 sollicitans {111), On Parris

Island, S. C, taeniorhynchus was the most abundant species in all

types of collections from June to September 1945 {26), In lighttrap collections at Fenwick Island lighthouse in southern Delaware in 1940, the total counts over a 60-day period, excluding the

collections for one night, showed nearly equal numbers of the

species (totals of about 6,500 of each from two traps) {208), On

the one night, September 10, the enormous total of 271,772 mosquitoes was taken in a single trap, of which about 192,000 were

taeniorhijnchiis and 70,000 sollicitans. On the other hand, lighttrap collections in 50 localities in New Jersey during the 10-year

period from 1932 -1941 gave a sollicitans-taeniorhynchiis ratio

of about 45:1 {H5),

The breeding habits of taeniorhynchus are similar to those of

sollicitans. Eggs are laid in similar locations and the larvae are

found in parts of the marsh above the level of the daily high

tides. The most prolific breeding occurs in potholes and temporary

ponds in Batis and Distichlis, but often over extensive areas of

Batis and mangrove, or in land-locked coastal swales. Larvae are

frequently found in fresh-water pools on vacant lots, golf courses,

and pastures near the coast. Many larvae were found once by tht

writers in fresh water standing on a vacant lot within the city

limits of Orlando in the interior of Florida—the only record of the

species among many larvae collections in the same locality.

The females are persistent, hard biters, and appear at times

in unbearable numbers. They seem much less inclined than sollicitans to attack in bright sunlight, but are commonly very annoying in the shade in the mangrove or other forests, or in shrubbery

about dwellings. They are migratory and disperse many miles

from their breeding places. The females were troublesome at

times in the vicinity of Orlando, Fla., which is about 30 miles from

the nearest salt water. Provost {255) has given other records on

the occurrence of salt-marsh mosquitoes in the interior of the

State. The same author {256) reported preliminary studies oí

the dispersal of taeniorhynchus from Sanibel Island, off the

Florida west coast, in which mosquitoes were marked by means

of radiophosphorus (P-^^). He reported that a non-purposive

flight occurred from one to four days after emergence and carried

the females to the limits of the collecting area, 20 miles and

probably beyond. The light-trap collection showed a ñve-dav'cvcle

of numbers, and the last radioactive specimen was caught on the
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 79

19th day. Dispersion appeared to be at random in all directions,

periodically repeated from foci established by previous migration.

From studies near Fort Pierce, Fla., Nielsen and Nielsen (239)

reported that biting showed a 5-day cycie, being most intensive

when the females were about 4, 9, and 14 days old. Females ready

to lay eggs performed a special flight, which occurred when they

were about 7, 12, and 17 days old. Most of them lived three or four

weeks and some of them much longer. Nielsen and Haeger (237,

238) reported that emergence of a brood was spread out over

two or three days, with a 3-hour outburst (when most of the

emergence took place) at some time during the 24-hour period.

The outburst occurred at the same time each day for one brood,

but the time varied with different broods. Newly emerged mosquitoes remained around the breeding place for 4 to 6 hours and

were then ready for the exodus, which took place only at night.

If emergence occurred between midnight and early afternoon,

the exodus took place the last half of twihght. Otherwise, the

exodus took place later in the night. So far as known, mating

usually occurred within the first 24 hours of female life. Swarming of males usually did not begin until three or four days after

emergence and, in the opinion of the authors, had no purpose of

releasing any other reflex mechanism ; i. e., it was not for mating

purposes, as commonly supposed.

Experimental transmission of several strains of encephalitis

virus by A. taeniorhynchus has been reported by several workers,

and Pistey (2^8) found it to be a suitable host of Dirofilaría

tennis of raccoons.

A. taeniorhynchv^ has been listed from all the Southeastern

States except Tennessee {193), and these records include inland

locations in Arkansas and northwestern Louisiana. There are,

however, no records of its occurrence in the Midwestern States,

where A. sollicitans has been found. It ranges along the Atlantic

and Gulf Coasts from Maine to Mexico, the Caribbean region to

northern South America, and on the Pacific Coast from Southern

California to Peru.

ADULT.—Occiput with a large median patch of pale lanceolate scales.

Mesonotum dark-scaled, sprinkled with white. Wing about 3.5 mm. in length,

entirely dark-scaled. Abdomen with basal white segmental bands but lacking

the longitudinal stripe of sollicitans and mitcheltae. Front and mid tarsi

with narrow basal white rings on segments 1 to 3, sometimes on 4 and 5

also, the rings on the hind tarsi wider but segment 1 lacks the median ring

of sollicitans. In specimens from the North and Central Atlantic States,

the fifth segment of the hind tarsi usually is entirely white. Southern and

tropical specimens having the tip dark are sometimes classified as a variety

—niger or portoricensis.

LARVA. Upper and lower head hairs each single, antennal tuft small,

double or triple, simple. Body pilose. Lateral hairs of segments 3 to 5 branched

three or more times. Comb scales in a patch, each rounded and fringed with

subequal spinules. Air tube short, 2:1 or less, with a multiple tuft beyond

the pectén, the dorsal preapical spine about equal in length to the apical

pectén tooth. Gills usually very short, budlike, but may be longer when the

larvae develop in fresh water.

AEDES THELCTER Dyar

(Syn., A. keyensis Buren)

This species is known in the Southeastern States only from the

Florida Keys. The thorax is largely pale-scaled, with an indistinct
80 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

median stripe of darker scales. The abdominal segments have

large median basal, triangular white patches, and the proboscis

and tarsi are unhanded. It was first reported on the Keys from

the collection of five females taken in light traps at Key West

and Marathon in October and November 1946 and named as a

new species—Aedes keyensis, by Buren {62), This name was subsequently synonymized with A, thelcter by Thurman et al. {305),

who obtained numerous larvae of the species on Long Key and at

Rock Harbor on Key Largo in 1947. One female each was taken

subsequently on Big Coppitt Key, Crawl Key No. 1, and Vaca

Key {53). The breeding places were temporary rain pools in

depressions in the limestone strata. The pool on Long Key was

at the edge of a buttonwood transition zone between hardwood

forest and a mangrove swamp. It was densely shaded by the surrounding tree growth and was filled with a thick mat of purslane

{Sesuvium), but free of permanent aquatic vegetation. Holes of

the land crab were present in the depression. Nine species of associated mosquitoes were listed, including Deinocerites cancer,

Aedes tortilis, A. taeniorhynchus and solUcitans, Psorophora

johnstonii, and several other species of Psorophora. Two females

were taken in biting collections.

In addition to the collection referred to above, Thurman et al.

{305) reported 12 light-trap records from Marathon and Key

West and 2 from Vaca Key. The species previously has been known

only from a few localities in Texas, Oklahoma, and Mexico.

ADULT.—Proboscis dark. Occiput with a median area of pale lanceolate

scales, broad ones laterally. Mesonotum clothed with narrow, curved, yellowish

scales, usually with a more or less distinct median stripe of pale brown

scales. Lower mesepimeral bristles usually absent. Wing about 3 to 3.5 mm.,

the scales all dark. Tibiae and first tarsals pale-scaled on one side, the

other tarsal segments entirely dark. The abdomen has unusual markings

consisting of large triangular spots of white scales at the base of the

segments, sometimes prolonged and widened apically on the posterior segments; segment 7 largely pale-scaled.

LARVA.—Antennal and preantennal tufts multiple. Upper and lower head

hairs single. Body glabrous. Lateral abdominal hairs single on segments

3 to 5. Comb of about 20 scales in an irregular row, each with a long apical

spine and much shorter side spinules, the preapical pair not more than half

as long. Air tube about 2:1, the pectén with one or two apical teeth more

widely spaced than the others, the ventral tuft multiple, inserted well before

end of pectén. Anal segment ringed by plate, the lateral hair usually double

Gills about as long as the segment.

AEDES THIBAULTI D. and K.

This comparatively rare mosquito is recognized by the lateral

patches of yellowish-white scales on the anterior half of the

scutum, separated by a broad stripe of dark scales. The proboscis,

tarsi, and abdomen are unhanded. It breeds exclusively in the

stump holes and basal cavities of several species of gum trees

and occasionally in the basal cavities of cypress. It has not been

found in ordinary tree holes above the flood level of the swamps

Information on its habits has been published in a number of

articles {^2, 157, 288) which were summarized by Jenkins and

Carpenter {17^). The eggs are probably laid on the inner walls

^r mi^^P ^^ *^^^ cavities and the winter is passed in this

stage. The eggs hatch in the spring when flooding occurs, and

larvae have been taken from March to May. The eggs may remain
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 81

viable for more than a year if flooding does not occur (158).

There is only one generation a year but there may be a succession

of broods when irregular hatching occurs. Larvae of Aedes triseriatuSy Culex apicalis (^ C. territans), C. restvuns, and Culiseta

melanura have occasionally been taken in association with

thibaulti. The adults remain close to the breeding places but bite

readily and become annoying in the daytime in these locations.

They are found resting in hollow trees and stumps, and occasionally about dwellings (S03). They are not considered to be of any

practical importance.

Since the previous publication (193), State records have been

added from Florida {228, 53) and Tennessee (66), which completes the list of the Southeastern States. Outside this area are

scattered records only from Kentucky, southern Illinois, and

eastern Texas.

ADULT.—The proboscis and tarsi of this species are -unmarked. Occiput

with yellowish-white scales on the median area, a small submedian patch

of broad dark scales on each side. Mesonotum with broad lateral patches of

yellowish-white on the anterior half, reduced to narrow lateral stripes on the

posterior half by an abrupt widening of the median dark-scaled area (fig.

19, D). Prescutellar space surrounded by pale scales. Lobes of scutellum

pale-scaled. Posterior pronotum with narrow, curved, pale scales. Lower

mesepimeral bristles usually absent. Wing length about 4 mm., the scales

all dark. Dark portions of femora not speckled with white. Abdominal

segments with large lateral, basal, white spots visible from above

LARVA.—Antenna nearly as long as head. Antennal, preantennal, and both

the upper and lower head hairs large and multiple. Body glabrous. Upper

lateral hair, or both pairs, on abdominal segments 1 and 2 multiple, lateral

hairs double on segments 3 to 5. Comb scales in a patch, each rounded and

fringed, the apical spine long and strong, the preapical pair nearly as long

and the others progressively smaller. Air tube about 4:1, the ventral tuft

multiple, inserted beyond the pectén. Anal segment not completely ringed

by plate, the lateral hair single. Gills pointed, about as long as segment.

AEDES TORMENTOR D. and K.

The females of Aedes tormentor are indistinguishable from

those of A. atlanticus, both having a broad median stripe of silvery-white scales on the thorax and unhanded tarsi. Identification

depends on the larvae or male terminalia. It breeds in temporary

rain pools in woodlands, and the larvae usually have been found

associated with A. atlanticus, but in much smaller numbers. At

New Orleans, La., however, they were taken more often than

atlanticus in the Metarie woods. The species has been recorded

from all the Southeastern States except Tennessee (193).

ADULT The proboscis and tarsi are unmarked. Occiput with a large

median area of white narrow scales, bordered by broad, appressed, mostly

dark scales. Median thoracic stripe extends the entire length of scutum and

is somewhat narrower than the dark area on each side. Lower mesepimeral

bristles absent. Wing about 3.5 to 4 mm., the scales dark. Abdomen unhanded,

with basal lateral, white segmental spots. Stem of male claspette long and

slender slightly pilose basally but without a bristle near middle; filament

much shorter than stem, pointed, and slightly hooked at tip.

LARVA—Antennal and preantennal hairs multiple. Upper and lower head

hairs single sparsely spiculate. Body glabrous. Lateral abdominal hairs

usually single on segments 3 to 5. Comb with 9 to 12 scales m a single row,

each thorn-shaped and finely fringed laterally. Air tube about 2:1, the pectén

extending to the apical third of tube, with evenly spaced teeth, the ventral

tuft inserted before end of pectén. Anal segment ringed by plate, the lateral

hair single or double. Gills pointed, somewhat longer than segment. Specimens

taken in Florida sometimes have the thorax and eighth abdominal segment
82 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

white in striking contrast to the rest of the body. A specimen collected at

Parris Island, S. C, was similarly marked.

AEDES TORTILIS (Theob.)

This small species is known in the United States only in southern Florida. The thorax is largely pale-scaled with a pair of

submedian dark stripes and a narrow border of dark scales

laterally. The proboscis and tarsi are unhanded. The first specimen to be recorded in Florida was a female caught in a light trap

at Key West in August 1945. A dead female was found about the

same time on a plane arriving in Miami from the Tropics (295).

Additional collections have been reported from Vaca Key, Cudjoe

Key, and Stock Island in Monroe County, and from Dade, Hendry,

and Palm Beach Counties on the mainland (232, 252, 53). Except

for the biting records, there is no information as to habits. One

of the authors (232) mentions the collection of several larvae but

the breeding places are not given. This mosquito is known from the

Greater Antilles and the Bahamas, and is said to breed in ground

pools. In Jamaica, larvae were collected in July 1957 in honeycomb rock holes on the hillsides above Montego Bay by E. J.

Beidler and W. V. King. These larvae were identified by Alan

Stone.

ADULT.—Proboscis dark. Occiput with a median patch of narrow silverywhite scales, broad ones laterally. Mesonotum with fine golden-brown scales,

divided by a pair of submedian longitudinal stripes of fine black scales;

mixed black and golden scales on prescutellar space. Lower mesepimeral

bristles absent. Wing about 3 mm., the scales dark. Tibiae and first tarsals

with pale stripes on one side, other tarsal segments all dark. Abdominal

segments with fairly broad basal white bands on segments 2 to 4, narrow on

5, and usually lacking on 6 and 7. The mesonotal vestiture of the male from

Clewiston, Fla., was observed to be entirely white and this sexual difference

was noted in reared males from Puerto Rico (252).

LARVA.—Antennal tuft multiple. Upper and lower head hairs single

Thorax sparsely pilose. Comb scales in a small patch, each rounded and

fringed, the apical spine slender, slightly longer than the subapical. Air

tube 2.5-3:1, the tuft multiple, inserted beyond the pectén. Anal segment

ringed by plate, lateral hair single. Gills less than twice as long as segment

shorter than the longest hairs in ventral brush. '

AEDES TRISERIATUS (Say)

(The tree-hole Aedes)

Aedes triseriatus has the sides of the mesonotum covered with

silvery-white scales separated by a broad central area of brown

scales (fig. 19 F) The proboscis and tarsi are unhanded. It is a

common woodland species, which breeds principally in tree holes

but to some extent also in water barrels and other artificial water

contamers, and has been reported from pitcher plants (213)

Larvae have been found in tree holes in gumbo limbo (Bursera

simarubra Sarg.) on the Florida Keys (252). The e^^s are laid

at or just above the water line. In the North the wintef is passed

m the egg stage but farther south larvae may be fouTd thS

out the wmter. A mmimum incubation period of 4 to 6 days at a

temperature of about 80° has been reported (Ul), Larval SeloD

SerVïÎ:¡Zu Í-,/r^-i-- with that of otíer treltX Dreeaers. 1 he adults are fierce daytime biters in or near infested

woods, and also are annoying about dwellings, ^e^^breedín^
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 83

occurs in domestic water containers or in shade trees on the

premises. In Arkansas they were reported to be abundant about

homes near woods {SOS), They are readily collected in light traps

at night.

A. triseriatus is of potential importance as a disease vector

since it is capable of transmitting the viruses of yellow fever (2^)

and equine encephalitis (8i). Its vector potentials with both the

western and eastern viruses were rated '^excellent'' (72), Also it

has been found to be a suitable host of dog heartworm Dirofilaria

immitis {2Jf7),

ADULT.—Proboscis and tarsi unmarked. Occiput with a large median patch

of narrow-curved, white scales. Mesonotum conspicuously marked with

stripes of silvery-white scales laterally, becoming narrower posteriorly, the

median area broadly dark-scaled (fig. 19, F.). Prescutellar space bordered

by white scales. Scutellum with a patch of white scales on the mid lobe, dark

ones on the side lobes. Posterior pronotum densely covered with broad,

appressed white scales. Lower mesepimeral bristles absent. Wing about

4 mm., the scales dark. Abdominal segments unhanded dorsally, with lateral

basal white spots, some visible from above.

LARVA.—Antennal hair single, preantennal hair double, lower head hair

with 2 to 4 branches, the upper single. Body glabrous. Lateral abdominal

hairs double on segments 3 to 6. Comb of 8 to 15 scales in a single irregular

row, each elongated, tapered to a rounded point, and evenly fringed with

small spinules (fig. 21, D). Air tube about 3:1, the tuft usually single or

double, inserted beyond the pectén. Anal segment not completely ringed by

plate; the tuft multiple. Gills stout, rounded at tip, the dorsal pair about

as long as anal segment, the ventral pair shorter.

AEDES TRIVITTATUS (Coq.)

The thorax of this species has a pair of broad submedian stripes

of yellowish-white scales separated by a broad stripe of brown

scales. The proboscis and tarsi are unhanded. It occurs in most

of the Southern States but is rare and considered of no economic

importance. No observations on its occurrence in this area have

been published other than brief records of its collection. Farther

north it is associated with floodwater areas. The females are said

to be fierce biters and become rather severe pests in some areas.

In Illinois it is one of the extremely annoying species {270), In a

study of trivittatus near Columbus, Ohio {1), it was found to

breed in temporary pools in open sites and in shallow ditches

covered with vegetation. The adults appeared the first week in

June associated with Aedes vexans and Psorophora ferox, and

were found up to the first week in September. They were seldom

caught in a residential area just beyond the breeding places. A.

trivittatus has been recorded in all the Southeastern States except Florida, Alabama, and Mississippi {19S), Its range extends

through all the Northern States and southern Canada to the Rocky

Mountains. ^ , ,

ADULT.—Proboscis dark. Occiput with narrow, curved, pale scales on

the median area, broad white scales laterally enclosing a small patch of dark

ones. Mesonotum with a pair of broad submedian stripes of yellowish-white

scales, separated by a broad central stripe of dark scales, narrow stripes of

dark scales laterally; prescutellar space bordered ^ith pale scales L^^^^^

scutellum pale-scaled. Lower mesepimeral bristles absent. Wmg length about

3.5 to 4 mm., the scales dark. Dark portion of femora and tibiae not speckled,

tibiae and first tarsals slightly pale on one side, tarsal s^^^f^^^ Í^^^^PA^H

LARVA.—Antennal and preantennal hairs multiple. Upper and lower head

hairs single. Body sparsely spiculate. Comb scales in a patch each rounded

with a long stout apical spine, the two preapical spines about two-thirds as
84 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

inntr nnd thp other spinules progressively shorter. Air tube about 2:1, the tuft SÄSLir/bryond peVn; do-al Preapical spine shor^^^^^

T^^r.fûv^ fnnfVi Anal see-ment rinered by plate, lateral hair single. Anai guib

Äth\'e1 tint^l^^^^^^^^^ a/d fonger than hairs in the ventral brush.

AEDES VEXANS (Meig.)

Aedes vexans has narrow basal white rings on the tarsal segments and rather broad white bands on the abdomen, notched on

the posterior border. The proboscis and thorax are unmarked.

The species is widespread in North America and on other continents, and is of great economic importance in many parts oí its

range. In some areas of the Northern States it is the principal

pest mosquito. In New Jersey it was the third most abundant

species taken in light traps during the 10-year period from 1932

to 1941 following Aedes sollicitans and Culex p. piinens in total

numbers It occurs in all the Southeastern States but is seldom

abundant there. The writers have taken occasional specimens in

Florida, and it has been collected in light traps by State Board oí

Health workers in most of the Florida counties (53). It was reported to be the most abundant species in Tennessee and an important pest in some localities (290), In Mississippi it was

reported to be one of the commonest and most troublesome

mosquitoes at Camp Shelby (225),

The species is primarily a floodwater breeder, and enormous

broods are produced when flood plains of rivers and streams are

inundated during spring overflows. It breeds also in flooded

meadows and in a great variety of temporary pools, usually m

open grassy sites or open woodlands, often associated with Psorophora confinnis and other temporary-pool breeders. Larval development is variable, depending on temperature, but is very

rapid during warm weather. The females are migratory. In

Oregon they were found commonly 15 miles from any breeding

source, and in lesser abundance as far as 30 miles away (296),

In Illinois, flight of stained specimens up to 14 miles was recorded

(77), Both males and females were collected in light traps in

Delaware Bay about 8 miles from the nearest shore (209).

The eggs are laid in moist depressions, especially those containing decaying organic debris. They may hatch during the same

season in which they are deposited, after the usual sequence of

incubation, drying, and flooding, so that several generations may

occur during the summer under favorable conditions. Experiments

in Oregon showed little mortality of dormant eggs over a period

of two years, but thereafter they lost viability rapidly (127), Very

few of the eggs would hatch when flooded with tap water, but a

large proportion did so when the oxygen content of the water

was reduced by bacterial growth or other means (125), Infusions

of amino acids and various other substances also acted as hatching

stimuli.

This species is capable of experimental transmission of eastern

and St. Louis equine encephalitis (8If, 135), and the virus of the

western type has been recovered from it in nature (63), Its vector

potential was rated "fair" with both the eastern and western

viruses (72), Laboratory transmission of fowl pox by vexans has
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 85

been reported (216), and it was shown to be a moderately

efficient host of Dirofilaría immitis (336).

ADULT.—The proboscis and thorax of the adult are unmarked. Occiput

bears numerous pale scales in the median area. Wing length about 4 mm.,

the scales dark. Lower mesepimeral bristles absent. Hind tarsi with narrow

basal white rings on all segments, front and mid tarsi with very narrow

rings usually visible on segments 1 to 3. Abdominal segments with basal

white bands which usually show a distinct, inverted, V-shaped notch on the

posterior borders, especially on segments 5 and 6.

LARVA.—Antennal and preantennal tuft multiple. Lower head hairs usually

with 2 or 3 branches, the upper with 3 to 5. Body glabrous. Lateral abdominal

hairs double or triple on segments 3 to 5. Comb with about 10 scales in an

irregular row, each thorn-shaped with fine lateral spinules. Air tube about

3:1, the pectén with one or two of the apical teeth more widely spaced than

the others, the tuft multiple, inserted beyond the pectén. Anal segment not

completely ringed by the plate, the lateral hair single or double. Gills about

twice as long as the segment, bluntly pointed.

Genus ANOPHELES Meigen

(Syn., Nyssorhynchus Blanch.)

The mosquitoes of this genus breed in a wide variety of aquatic

environments, although their production on a large scale is chiefly

in permanent bodies of water containing vegetation or surface

debris. Descriptions of anopheline breeding places of various

types, while not specifically referred to herein, are numerous in

the literature cited. Some of the references deal with the classification and types of breeding places in given localities, such as,

Bradley (41) for northeastern Louisiana; Barber and Komp (13)

and Perez (2M) for Mississippi; Boyd (32, 33) for northeastern

North Carolina; Watson and Spain (319) for northern Alabama;

Meleney et ah (222) for western Tennessee; and Darling (83) for

Georgia (Lee County). The bionomics and ecology of the Nearctic

species have been reviewed by Bradley and King (49), Freeborn

(115), and others. The works of Boyd (34, 36) contain comprehensive reviews of the literature on the natural history of anophelines as well as on the relation of the various species to the

epidemiology of malaria.

All our native anophelines are fresh-water breeders, with the

exception of Anopheles átropos and A. bradleyi, which breed in

salt or brackish water, and A. albimanus which occurs in either

brackish or fresh water. The number of generations per year

varies greatly; A. qttadrimaculatiis may have from 8 to 10 generations annually in the latitude of southwestern Georgia (32),

In the fall, inseminated females enter caves and cellars to hibernate. In the warmer sections of the Gulf States breedmg also

is more or less continuous through the winter (14, 131), although

much reduced in volume, and the rate of development is slower.

The larvae of some species are able to withstand freezing temperatures (6). The adults are active chiefly after dusk and spend

the daytime resting in dark, humid situations. Anopheles S.YQ considered to be relatively weak fliers, although migrations of several

miles have been recorded. A review of the literature relating to

anopheline flight and dispersion has been published by Eyles

^The eggs of anophelines (fig. 3, C) are boat-shaped, pointed

on either end, and are laid singly—that is, not stuck together m
86 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

rafts—on the surface of the water in permanent pools and ponds

where there is emergent or floating vegetation and debris ot

various kinds. They are provided with floats o ke^p them at the

surface of the water. The meniscuses of individual eggs cause

?hem to be attracted to one another and to form various geometrical patterns on the water surface. Hatching usually occurs

in 2 or 3 days, and breeding is continuous during the summer

months.

M & A 14063

FIGURE 23.—Resting positions of mosquitoes: A and B, Anopheles; C, Culex.

Living adults of most species of anophelines can be recognized

by their typical resting position (fig. 23, A, B), the abdomen and

proboscis being held in nearly a straight line and pointed at an
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 87

angle toward the resting surface. Other kinds of mosquitoes hold

the body more or less parallel to the resting surface (fig. 23, C),

while the head and proboscis are bent downward at an angle to

the body. The palpi of anophelines (except in the Australasian

genus Bironella) are as long as the proboscis in both sexes (fig.

1, A, D) and are enlarged or somewhat club-shaped in the males.

The wings usually have spots of dark or white scales (pi. 8). The

posterior border of the scutellum is evenly rounded and fringed

with hairs. In all our species except A. albimanus the abdomen

has a vestiture of fine hairs rather than the flat scales found on

most other mosquitoes. Albimanus has some flat scales.

The larvae of Anopheles are easily recognized by the absence

of a breathing tube and by their usual feeding position (fig. 20, A)

parallel to the water surface. Other mosquito larvae have an elongated breathing tube and while at the surface hang downward with

only the tip of the tube penetrating the surface film (fig. 20, B).

Anopheline larvae have paired, fan-shaped tufts of modified hairs,

called palmate hairs, on the upper side of the abdominal segments,

and a pair of transparent, notched organs on the front border of

the thorax that allows the larvae to suspend themselves just below

and parallel to the surface. While in this position the head is

rotated for feeding until the mouth parts are uppermost, and the

food is taken from or near the surface film. The clypeal hairs,

sutural hairs, palmate and antepalmate hairs, and the lateral

hairs on abdominal segments 4 and 5 are the characters most

frequently used in the identification of anopheline larvae (fig. 13).

In the descriptions of these characters the usual range observed

in the number of branches or the form of branching is given,

although additional variation may occur in a small percentage of

specimens.

M & A 14064

FIGURE 24.—Pupal breathing tubes: A, Anopheles crucians, from above and

side; B, Culex salinarius; C, Culex p. qutnquefasciatus.
88 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Miles (229) has reported a study of the variations in the chaetotaxy of series of larvae of A. bradleyi georgianus, and punctipennis. Roth (272) has recorded the aberrations and variations encountered in a study of large numbers of larvae of the commoner

species. He shows that in species in which the inner clypeal hairs

are normally simple a considerable number may show some

branching. Usually only one hair was found to be forked toward

the tip, but in a few specimens there were several branches on

one or even both sides. It should be remembered, therefore, that

quite atypical individuals may be encountered.

In the pupal stage the breathing tubes are short and widely

flared as compared with those of other mosquitoes (ñg. 24) and

the hairs on the posterior corners of abdominal segments 3-6 are

modified into short spines.

To this genus belong the mosquitoes that transmit malaria to

human beings, and all except two (A. georgianus and A. perplexens) of the species listed for the Southeastern States have been

proved susceptible to infection with the parasites of this disease.

Anopheles qvndrimacidaUis was by far the most important species

concerned in the transmission of the disease in the southeastern

region. The others either are too rare, or their blood-feeding habits

are such that they are considered to be of little importance as

carriers. These conclusions are based on the fact that malaria

infection in this region was almost invariably found associated

with quadrbnaculatiis,

A, crucians is prevalent along the coastal plains and in the

lower Mississippi Valley, and since specimens have been found

infected in nature, the species cannot be eliminated entirely from

consideration. Investigations several years ago in the Okefenokee

swamp in southern Georgia (220) showed that malaria was

absent in an area where crucians was prevalent and the only

anopheline present. In Florida the malaria rate formerly was

high in the northwestern part of the State, where A. quadrimaculatiis predominates, but low in the southern half of the State,

where crucians is abundant and the predominating species. Such

malarious foci as were investigated in the southern part of the

State have shown locally favorable conditions for quadrimaculatus

breeding. Frequently the two species are found associated in the

breeding places, but in general the occurrence of quadrimaculatus

and the areas of high malaria endemicity in northern Florida and

southern Georgia (39) appeared to be correlated with the presence

of slightly alkaline surface waters, whereas crucians prefers

water of an acid reaction, as found more generally in the southern part of Florida. The findings of high percentages of crucians

infected with parasites in South Carolina by Sabrosky et al

(280), Frohne et al, (117), and Atchley (If.) would seem to

indicate that A. crucians may have played a more positive role in

malaria transmission in that region. However, it was pointed out

that since these findings were made during a period of low human

parasitemia, the parasites may not have been those of human

malaria.

A critical review of the records on infection of southern species

and their role in malaria transmission was given by Barber and

co-workers (15) in 1927, and more recently Boyd (36) has pro-
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 89

yided a summary of such information for the anophelines of

the world.

^ Genus Anopheles includes most of the species of tribe Anophelini

m the world but there are two other small genersi—Chagasia,

íound in tropical America, and Bironella, in tropical Australasia.

All ot our species are placed in subgenus Anopheles except A.

albimamis which belongs to subgenus Nyssorhynchus Blanch.

King and Bradley {190, 191), Freeborn {115), and Carpenter

and LaCasse {68) have given reviews of the classification and

distribution of the Nearctic species.

In the brief description of "the adults under each species, distinguishing characters in the male terminalia are added for the

species having the wing scales all dark, as specimens of these are

by the male terminalia.

typically marked. Roth {271) has published detailed keys with

illustrations to identify the Anopheles of the Southeastern States

sometimes difficult to identify otherwise when rubbed or not

KEY TO SPECIES

ADULTS

1. Wings with spots or areas of white scales 2

Wings entirely dark-scaled 6

2. Tarsi with conspicuous white bands or rings albimamis

Tarsi all dark 3

3. Costa dark except at extreme tip of wing (pi. 8, A); anal vein

with three dark spots separated by white (two dark spots in

male). Three species that are indistinguishable as adults except

for about half of the specimens of bradleyi in which the stem

of vein 5 is white-scaled instead of all dark as in the other two

species crucians, georgiamiSj bradleyi

Costa with a white spot, usually prominent, at outer third, opposite

the tip of the subcostal vein (pi. 8, C and D) ; anal vein with

one or two dark spots or lines 4

4. Palpi with white rings; veins 3 and 5 with long areas of white

centrally pseudopunctipennis

Palpi all dark ; veins 3 and 5 entirely dark-scaled 5

5. Costal white spot prominent, nearly equal in length to the blackscaled area beyond it punctipennis

Costal spot much reduced, usually less than a third as long as the

subapical black-scaled area, sometimes lacking; other white

spots on wing also reduced, especially on stem of vein 4. A

species closely related to the above, which can be identified

positively by the morphology of the egg (see the descriptions

of the two species) perplexens

6. (1) Wings unspotted; mesonotal bristles unusually long, about

one-half the width of scutum; palpi entirely dark. A very small

species breeding in tree holes barberi

Wings with dark spots, more or less distinct; mesonotal bristles

mostly shorter than one-half width of thorax 7

7. Palpi entirely dark; wing spots distinct; small knee spots present;

frontal tuft of occiput with some of the hairs pale, a median

patch of pale scales at vertex ; processes of ninth tergite of male

stout, rounded, and usually expanded at tip quadrimaculatiis

Palpi with white rings, or knee spots lacking; wing spots usually

indistinct; frontal tuft all dark; processes of ninth tergite

slender, pointed 8

8. Palpi usually distinctly ringed at apex of each segment; occiput

with a patch of pale scales at vertex; knob of halter pale-scaled;

knee spots present; phallosome with the preapical pair of

leaflets more than half as long as the apical pair. Breeds in

fresh water walkeri
90 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Palpi with faint white rings on apical segments or entirely dark;

occiput entirely dark-scaled; knob of halter dark-scaled; knee

spots absent; phallosome with the preapical pair of leaflets no

more than half the length of the apical pair. Breeds in salt

water átropos

LARVAE (FOURTH INSTAR)

1. Abdomen with plumose lateral hairs on segments 1 to 6, the

branches short and sparse; frontal head hairs (Nos. 5-7) minute,

single barben

Abdomen with plumose lateral hairs on first three segments only;

head with large plumose frontal hairs (fig. 13) ... 2

2. All clypeal hairs simple, inner pair well separated; postspiracular

plate with a slender pointed tail on each side from posterior

margin pseudopunctipennis

Outer clypeai hairs branched; postspiracular plate without tails 3

3. Outer clypeal hairs sparsely feathered or with a few (not over

10) short branches; inner clypeals usually sparsely feathered . . 4

Outer clypeal hairs thickly branched, the branching dichotomous;

inner clypeals simple (except in walkeri) or occasionally forked

or branched 5

4. Inner clypeal hairs closely approximated; lateral hairs on abdominal segments 4 and 5 branched, usually triple ........ átropos

Inner clypeal hairs widely separated (by about one-third the

distance between the outer clypeals) ; lateral hairs on segments

4 and 5 single albimanus

5. Abdominal segments with two conspicuous hair tufts (hairs 2

and 0) anterior to palmate hair; these hairs usually approximately equal in size and with four to nine branches on segments

4 and 5 crucians

Hair 0 undeveloped or very much smaller than hair 2, the latter

usually single or double, but sometimes with three to six branches

on segments 4 and 5 6

6. Sutural head hairs (Nos. 8 and 9) large, usually with 8 to 10

branches; basal tubercles of inner clypeal hairs separated by

more than the diameter of one tubercle ; palmate hair frequently

partially developed and functional on segment 2 . . . . quadrimaculatits

Sutural hairs with two to six branches or the basal tubercles of

inner clypeals usually separated by less than their diameter

(variable in bradleyi) ; palmate hairs rudimentary on segment

2 (except in walkeri) 7

7. Inner clypeal hairs with sparse, minute feathering toward tip;

hair 1 of prothorax with three to five branches from near base;

palmate hairs partially developed on segments 1 and 2 walkeri

Inner clypeal hairs usually simple (rarely forked or with two

or three branches toward tip) ; hair 1 of prothorax short,

usually single or weakly branched toward tip (frequently with

some branches before middle in punctipennis) ; palmate hairs

rudimentary on segments 1 and 2 8

8. Only three pairs of functional palmate hairs (segments 4 to 6) ;

hair 5 of segment 1 usually with five to seven branches, arising

irregularly along the stem georgianus

Five pairs of functional palmate hairs (segments 3 to 7) ; hair

5 of segment 1 usually with three to five branches from near base 9

9. Leaflets of palmate hairs on segments 3 and 7 usually somewhat

smaller than those on 4 to 6 and mostly with smooth margins,

especially on segment 7; hair 5 of segment 1 about twice as

long as hair 4 and usually four- or five-branched bradleyi

Palmates on segments 3 and 7 about equal in size to the others,

with broad leaflets, most of which have serrated margins; hair

5 of segment 1 about of same length as hair 4, and usually threeor four-branched. Two species with very similar larvae, which

can usually be identified, at least in a series, by the number of

branches of the antepalmate hair (hair 2) on segments 4 and 5

as follows:

Hair 2 usually with two to four branches, rarely with one

or two of the four hairs single punctipennis
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 91

Hair 2 usually all single, or with no more than two of the

four hairs on segments 4 and 5 two-branched before

the middle, rarely with three or all four, branched before

middle perplexens

ANOPHELES ALBIMANUS Wied.

(Syn., A. albipes Theob.)

This species is the only anopheline included here in which the

tarsi are white-banded. It is a tropical species, of much importance

as a vector of malaria in tropical America. It was introduced

into Key West, Fla., in 1904, and apparently developed one

brood there before being eradicated, according to the records of

its discoverer, George N. MacDonell. This event and the possibility

that the species might become established in southern Florida

and along the Gulf Coast were discussed by King (184.). During

and after World War II, albimamos was again discovered in southern Florida, with collections being reported from the Keys, (132,

253), one adult from as far north as West Palm Beach (323),

and one larva from Boca Raton (66), Pritchard et al. (253)

reported finding a single larval specimen on Big Pine Key in

January 1946 and discussed the possibility that the species may

be endemic in southern Florida, maintaining itself in limited

numbers and locations. This appears to be the situation, for

Haeger (133, 134) has reported the collection of larvae and adults

at intervals during the years 1946-1950 while making intensive

studies on the Keys. He found continuous breeding on Big Pine

Key from October 1947 through December 1948, and the numbers

of specimens sometimes were fairly large. The maximum lighttrap collection for one night was 110 specimens, taken in early

September. A few additional captures on the Keys from 1951 to

1957 are reported by Branch et cil. (53). Albimanus is known to occur also along the Rio Grande River in the extreme southern tip

of Texas, and was taken in light-trap collections in November

1945 at Corpus Christi, 150 miles to the north, by Eads (99).

The larvae of albiinanus occur in open sunlit lakes and ponds of

fresh or brackish water amongst mats of algae and other aquatic

vegetation. They may be found also in a wide variety of temporary

collections of water such as occur in hoof prints, wheel ruts, and

artificial containers. The adults are strong fliers ; the females feed

readily on man and domestic animals. They enter dwellings during

the night to bite but leave at dawn for forested areas to spend

the daytime.

ADULT.—Proboscis dark, palpi with the apical segment (segment 5) all

white, and segments 2 and 3 white-tipped. Occiput with a median area of

white scales, dark ones laterally. Mesonotum clothed almost entirely with

fairly broad, palish scales, extending on to scutellum. Wing costa with five

or six white spots, some broad, and the other veins with numerous alternating

black and white spots. Hind tarsi with segments 3 and 4 and apical half

of segment 2 entirely white, the apical half of 5 also white, the base dark;

fore and mid tarsi with the first two or three segments white-tipped, the

others all dark. Abdominal tergites with fairly broad, pale scales through the

central area, dark hairs or bristles laterally.

LARVA.—Antennal hair very small, split toward tip with two or three

branches. Outer clypeal hairs single, and both the outer and inner clypeal

hairs sparsely feathered or barbed, the inner hairs well separated, their

bases being spaced nearly a third of the total distance between the outer

clypeals. Post clypeals short, forked toward tip. Palmate hairs developed
92 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

on abdominal segments 1 to 7, sometimes on the metat^^^^^^^^^

&ÄS'oTlS^nrAT&^^^ - ^'

single on segments 4 and 5.

ANOPHELES áTROPOS D. and K.

Little is known of the habits of this species except that it breeds

in the ^^^^^^^ of coastal marshes from New Jersey southward

{o Texas In southern Louisiana, Mississippi, and Florida adult

females have been taken while biting in the open during the day,

evSi in dfrect sunlight (132, UU 205). In Florida, adults were

once noted on a^i marsh during the day and were encountered

i^iTarge numbers after dark in two other localities. Adu ts have

been taken in Jight traps in several localities on the coast.

Larvae were taken by Griffitts {182) in water ranging from 3

to 12 percent ^salinity- (about 0.8 to 3 4 percent of fait) by

Hinman {151) in water containing 0.8 to L85 percent of salt

and by the writers, in Florida, in water contammg more than 1

percent of salt. Although the larvae of átropos ^nà bradleyi have

occasionally been taken together, thfe latter usually occurs m water

of a lower salt content. . -^-i, ^«i Anopheles átropos has been infected experimentally with malaria parasites {221), but it is of doubtful importance as a transmitter of the disease. Flights of a mile or so have been observed

ADULT.—This species is almost entirely dark-scaled Apical segnients of

the palpi faintly tipped with white or all dark. Occiput entirely dark-scaled

Bristles of the mesonotum of normal length, less than half the width of the

mesonotum. Wing about 4 mm. long, usually with three or four spots slightly

darker than the rest of the scaling (pi. 8, F). Knob of the halter darkscaled Abdomen and tarsi all dark. Femoral knee spots usually absent.

The female differs from A. walkeri in having less distinct rings on the palpi

and somewhat less distinct spots on the wing, the occiput and halter knob

all dark, and femoral knee spots absent. It differs from quadrimaculatus m

its smaller size, presence of palpal rings, dark occiput, much less distinct

wing spots, and absence of knee spots. A. harheri is much smaller m size,

has abnormally long thoracic bristles, entirely dark palpi, and no indication

of dark spots on the wing. In the male there are three or four pairs of

leaflets on the phallosome, the apical pair being twice as long as the next

pair, which distinguishes it from walkeri. The lobes of the ninth tergite are

bluntly pointed and much more slender than in A. quadrimaculatus.

LARVA.—Antennal hair small, divided into several branches toward tip,

inserted at about the basal fourth of antennal shaft. Inner clypeals long,

usually sparsely feathered toward tip, rarely simple, set close together, their

basal tubercles separated by about the diameter of one; outer clypeals not

more than half the length of the inner, with four to ten, rarely two or three,

small branches toward tip. Post clypeals short, usually forked at tip. Frontal

head hairs large, plumose. Inner and outer sutural hairs single or forked

beyond middle. Hair 1 of prothorax small, single. Hair 0 on abdominal segments minute. Palmate hairs well developed on segments 4 to 7 and partly

developed on 3, the leaflets with notched or serrated edges on apical half.

Antepalmate hair (hair 2) single on segments 4 and 5. Lateral hairs on

segments 1 to 3 plumose, those on 4 and 5 usually three-branched.

ANOPHELES BARBERI Coq.

The species occurs throughout the eastern United States from

New York southward and westward to Texas and Iowa, and has

been reported from all of the Southeastern States. Its habits and

distribution have been summarized by Jenkins and Carpenter

(17-4). It is never abundant nor found far from its breeding places.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 93

The larvae are found principally in tree holes, although the writers

once took them in wooden tubs at Mound, La., and they have been

reported since from other artificial water containers. Larvae may

be found in the breding places throughout the year. Matheson

(213) states that the species hibernates in the north as secondstage larvae frozen solidly in ice. The larvae frequently are associated with Aedes triseriatus, Orthopodomyia signífera, and Toxorhynchites sp. The adult is very small, with unspotted wings,

and is rarely encountered. Adults are to be found resting during

the day above the water in tree holes where larvae occur and also

in buildings and other shelters nearby. Thibault {303) in Arkansas

noted that it enters dwellings readily and is a persistent biter,

although easily disturbed. Both sexes are attracted to lights and

often are found in light-trap collections. It has been proved susceptible to infection with malaria parasites (300), but is of

doubtful importance in malaria transmission because of its limited

occurrence and association with man.

ADULT.—This species is a small, completely dark, unmarked anopheline.

Wing length about 3 mm. Knob of halter dark-scaled, wings without dark

spots (pi. 8, H). In the male, the phallosome is without leaflets, which distinguishes it from all other anophelines in the United States except A.

albimanus.

LARVA.—Antennal hair small, single, inserted near middle of shaft. Inner

and outer clypeals single, the inner widely separated, closer to the outer

than to each other. Post clypeals usually single, longer than outer, and more

widely spaced. Frontal hairs small, simple, an unusual character in

anophelines. Sutural hairs also single. Hair 0 on abdomen minute, unbranched; antepalmate hair on segments 4 and 5 branched toward tip, and

inserted at side of palmate hair instead of in front; palmate hairs well

developed on segments 2 to 7, the margin of the leaflets serrated; lateral

hairs plumose on segments 1 to 6, the branches short and sparse.

ANOPHELES BRADLEYI King

(Syn., A. crucians var. bradleyi King: A, criicianSy

coastal or salt-water form)

Prior to 1939, when this species was first named, it was known

as the '^coastal variety'' of Anopheles crucians (UU) ^ It occurs

along the Atlantic and Gulf Coasts from Massachusetts south to

Texas and Mexico. Because bradleyi adults can hardly be distinguished from those of crucians and georgianus, little is known of

their specific habits. Reared specimens have bren infected experimentally with falciparum malaria (38),

A. bradleyi larvae have been found principally in brackish

water near the coast, and water with a low concentration of salt

(about 1.5 percent or less) appears to be preferred. They have

been taken with Anopheles átropos at the higher concentration

and with the typical crucians when the water was nearly fresh.

The larvae were first collected in Florida near the Saint Johns

River in Brevard County in large roadside ditches having profuse

growths of Chara and other aquatic plants, and have been found

there repeatedly upon subsequent examinations. Elsewhere they

have been collected in salt-marsh pools containing flotage, algae,

and grasses. In Maryland, Vogt (3H) reported that m the summer months the larvae were distributed, without regard to salmity,

in locations having emergent vegetation of appreciable height.

During the cooler weather of spring and fall they occurred m
94 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

water having submerged growths of pond weeds and algae but

devoid of emergent vegetation. He attributed this to surface water

temperatures, which may have been too high for the larvae in

open waters in the summertime.

ADULT.—Anopheles bradleyi is one of the three members of the crucians

series that are recognized by the ringed palpi and black and white spotted

wings, but with the costa dark. The adults are difficult or impossible to distinguish from A. crucians and A. georgianus, except in specimens that have

the stem of wing vein 5 entirely or mostly white-scaled. This condition has

been found to occur in about half of the specimens examined. The male

terminalia of the three species also are very similar.

LARVA.—Antennal hair branched several times from near middle. Inner

clypeals fairly close together, the tubercles usually separated by about their

own diameter or less, sometimes more (about 25 percent according to Miles

{229)). Outer clypeals widely separated, thickly branched. Postclypeals very

short, usually single, occasionally double, rarely three or four. Sutural hairs

long, with two to six branches, the outer hair (No. 9) usually with three

to five branches from near the middle, longer than inner, reaching base of

frontal hair. Prothoracic hair 1 of thorax single or with up to five weak

branches at tip. Hair 0 on abdominal segments small, unbranched. Palmate

hairs developed and functional on segments 3 to 7, those on segments 3 and

7 somewhat smaller than the others and the leaflets mostly slender, with

smooth margins, especially on segment 7. Antepalmate hair on segments

4 and 5 single, double, or triple (about 50 percent single in the series reported by Miles (229)). Hair 5 of abdominal segment 1 about twice as long

as hair 4 and usually four- or five-branched (86 percent). Lateral hairs on

segments 4 and 5 usually triple, with a fairly long stalk before point of

branching. The larval characters most nearly resemble those of A. punctipenjiis from which it usually can be separated by the relative size and shape

of the leaflets of the palmate hairs. The relative lengths of hair 5 of

abdominal segment 1 and the number of branches of the antepalmate hairs

are frequently of assistance for confirmation.

ANOPHELES CRUCIANS Wied.

The crucians group is now known to include three closely

related species which are very similar in the adult stage but differ

m larval and pupal characters. Two of these, bradleyi and georgmnus, were described by King in 1939 {185) as varieties of

Anopheles crucians, but it is believed that all three represent

distinct species. The principal recognition characters for the adults

are the white-spotted wings with three dark spots on the anal

vein (two m the male), and dark-scaled front margin of the

T^!^^^J'^l' ^ ^J:J\^ P^^P^ ^^^ ^^^^^^ ^ith white. When resting, the adults hold the body at an angle of almost 90° with the resting

surface and in the Southeastern States may be distinguished

from other anophelmes in shelters by this characteristic

Anopheles crucmns is the common fresh-water form in the

Southeastern States. In low coastal-plains areas it is frequentlv

the predominating anopheline, but elsewhere is usually limited in

abundance and distribution. Its range extends up the Atlantic

Coast to Massachusetts and westward into southern IllS

central Oklahoma, and Texas, with an isolated occurrence in S

.^r.^Ll^^?j:/r ^}'n'}' -i^o occurs on the ea^t ?oas of Mexico and in some ¿rts of the "Caribbean "rêgCrc S

arvae develop m acid waters, but often are foSnd also S

larvae of quadrimaculatus in the alkaline waters of lakes Donds

pools and swamps of a permanent or temporary natuS wher¿

aquatic vegetation or debris provides harborage. FreeborA (115^

states that they also occur in wheel ruts, temporary pools in ^^^^^^^
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 95

barrens, and other non-vegetated water accumulations in which

quadrimaculatus rarely would be found.

Although this species is susceptible to infection with malaria

parasites, it is not known to be of serious importance in the

transmission of malaria ; in fact, the evidence points to the contrary, as pointed out in the generic discussion. Its virus potential

for eastern equine encephalitis was rated as ''poor'' in tests by

Chamberlain, et al, {72), but the virus of this type was recovered

from a pool of wild-caught crucians in Louisiana and another in

Georgia {176, 196), The species frequently bites out-of-doors at

night or even during the day in the woods. The adults also enter

houses, but at Mound, La., they were always found in a much

smaller percentage of the total numbers present in the area than

was the case with Anopheles quadrimaculatus. Also in the vicinity

of Lake Apopka in central Florida, where crucians becomes extremely abundant, the number taken out-of-doors at night while

attempting to bite was repeatedly very small in comparison with

the total numbers present, as indicated by light-trap collections.

Observations made by the senior author in New Orleans indicate

that this species may migrate for several miles when an unusual

abundance occurs. MacCreary and Stearns {209) obtained specimens at two light-houses in Delaware Bay, one 3.2 miles from the

nearest shore and the other 5.5 miles. Since both these localities

are near the coast, it is not certain whether the species represented

was crucians or hradleiji.

ADULT.—Proboscis dark. Palpi with the apical segment all white, segment

4 with narrow white rings at base and apex, and segment 3 usually with

a few white scales at apex. Occiput with white narrow-curved scales and

upright forked ones on vertex. Mesonotum with a grayish appearance and

two darker submedian longitudinal stripes. Wing length about 4 mm.;

costa and subcosta all dark, the other veins with alternating black and white

spots, and the wing tip with a wide pale fringe spot; anal vein (vein 6)

with three dark spots separated by white. Knob of halter dark. Tarsi all

dark. In the male there are only two dark spots on the anal vein. The spines

on the male claspette of the three species in this group differ slightly in

number and shape {185), but their variations overlap and seem of doubtful

use for identifying single specimens, although they may be of some use in

a series.

LARVA.—The larva of crucians is distinguished from all other American

Anopheles by the fact that the anterior submedian hair (hair 0) is well

developed on several of the abdominal segments and the antepalmate hair on

segments 4 and 5 is also multiple-branched. Antennal hair with several

branches along the stem, inserted near basal fourth of shaft. Inner clypeals

simple (occasionally forked beyond middle), closely approximated, their

bases separated by less than the diameter of a basal tubercle; outer clypeals

densely branched; post-clypeals short, single, or bifid distally. Sutural hairs

fairly large with several branches along stem beginning near base. Palmate

hairs well developed on segments 3 to 7, those on 3 and 7 about equal to the

others or slightly smaller; the margins of the leaflets serrate. Hair 0 well

developed with four to nine branches on segments 4 and 5 and about equal in

size to the anteualmate hair. Lateral hairs on segments 4 and 5 with several

branches beginning about the basal fourth.

ANOPHELES GEORGIANUS King

(Syn., A. crucians var. georgianus King)

This species was first collected by Bellamy {22) in 1937 from

fresh-water breeding places in southern Georgia, and was described

by King {185) as a variety of Anopheles crucians. The spots on

the wings appear to be more contrasting than in crucians, but no
96 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

positive characters have been found for separating the adults, and

nothing is known of their habits. The larva is unusual in that only

three pairs of functional palmate hairs are present (on segnients

4 to 6), whereas crucians and all the other species of Anopheles

have at least five. Slight differences in the claspette spines of the

male genitalia were described by King (185), with crucians

intermediate in these characters.

Anopheles georgianus has been collected by Bellamy {22) m

Brooks, Sumter, Terrell, and Thomas Counties, Ga. A single larvae

was identified from a collection of cruciayis from a pond near

Hinesville, Ga., in March 1941. It has since been reported from

all the Southeastern States except Tennessee and Arkansas. Its

typical habitat appears to be shallow collections of water m seepage areas at the heads of small streams, where the larvae usually

occur in pure culture or with a small proportion of crucians,

Wirth {380) collected the larvae in several localities in Louisiana,

all in hillside seepage areas. They were found in small puddles,

4 to 30 inches in diameter, 2 to 4 inches deep, with algae and grassy

edges. Michener {225) obtained the larvae in Mississippi in shallow water in spring-head areas with a low pH, usually with dead

leaves and often with clumps of grass. Freeborn {115) reports

that the presence of the small pitcher plant Sarracenia j)urpurea

is often an index to the occurrence of georgianus.

ADULT.—This species belongs to the crucians group, or series, and appears

indistinguishable except by larval characters. The description of the adult

of A. crucians may therefore be referred to.

LARVA.—Hair 0 undeveloped, being minute and unbranched, which distinguishes it immediately from crucians. There are only three pairs of

functional palmate hairs, on segments 4 to 6, which is quite exceptional

among all anophelines. Miles {229) has shown the variations in the chaetotaxy

of a series of specimens, some of which are as follows: posterior clypeal

hairs usually simple or forked, rarely with three or four branches; outer

occipital hair with two to six branches, usually three to five, short, about

equal in length to the inner hair; antepalmate hairs of segments 4 and 5

usually two- to three-branched, occasionally simple or four-branched; hair

5 of abdominal segment 1 with four to nine branches, usually five to seven,

the branches arising irregularly along the stem.

ANOPHELES PERPLEXENS Ludlow

This species was placed as a synonym of A, punctipennis and

regarded as an extreme melanistic variant {163). A study of

specimen material from southern Georgia and elsewhere by Bellaniy {23), however, has shown that, in addition to the reduced

white scaling on the wings described by Ludlow, and reduced

branching of the larval antepalmate hairs previously noted by

King et al. {193), the ova of the two forms were "strikingly

different." He therefore considered it to be a valid species and this

is concurred in by Drs. Alan Stone and K. L. Knight (personal

communication).

Nothing is known of the blood-feeding or resting habits of

per^olexenSy or its relation to the transmission of malaria. Its

breeding places were found by Bellamy to be restricted principally

to limestone springs and the margins of streams flowing from

them. This type of breeding place had previously been noted by

King et al. {193) for the melanistic form in central Florida. Seepage from an outcrop near Gainesville, Fla., where larvae were
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 97

collected, was probably through a limestone formation. Since then

the Florida State Board of Health has accumulated records of this

melanistic form (based on the reduced costal spot) from a number

of other counties in north-central Florida. Bellamy's study material came mostly from Baker and Calhoun Counties in southern

Georgia and included some material from other Georgia counties,

and from a few other States. Russell (279) reported four reared

adults with repressed costal spots, on two of which the spot was

completely repressed. The exact locality for these specimens was

not given, but they came from a series collected mostly in Lee

County, Ga., with a few specimens from Sumter County, Ga., and

Covington County, Ala. Roth (272) mentioned the occurrence of

melanistic adults and larvae with single antepalmate hairs at

Paris, Tenn., and Swannanoa, N. C. The female specimen from

which perplexens was originally described came from Camp

Roosevelt at Mount Gretna, Pa.

ADULT.—This species is generally darker than punctipeiuiis. Wing length

about 4 mm.; white spots on costa opposite the tip of the subcostal vein

short, typically not more than a fourth or fifth as long as the subapical dark

spot adjoining it, sometimes reduced to a few white scales or entirely dark.

The mean of measurements by Bellamy (23) of this spot gave a ratio to the

total length of the wing of about 0.055, compared with a ratio of about 0.11

for punctipennis, or averaging about half as long. The range of ratios was

not given but it is presumed that there was overlapping. The costal spot in

piinctipennis is usually about one-half or more as long as the subapical dark

spot, and much longer than the apical white spot. In the type female of

perplexens the costal spot is about one-fifth as long as the sub-apical dark

spot, and about equal in length to the subapical white spot. The costal white

spot does not extend onto vein 1, white spots are lacking or very indistinct

on the forks of vein 2, white spots are present on the forks of vein 4, about

one-third as long as the forks, and the stem of vein 4 is apparently all darkscaled. Of the present authors' collections from central Florida, one specimen

has the costal spot completely suppressed, and on one other the scales on the

forward edge of the costa are all dark but with a few indistinctly pale scales

behind. There are four specimens in which the costal spot is of about the

same width as in the type, and the other spots are similar to that specimen.

Other specimens have wider costal spots but are possibly the same form.

Measurements of the four specimens mentioned above gave ratios of the costal

white spot to the subapical dark spot of 0.25, 0.20, 0.22, and 0.20. The ratios

of some of the other specimens were: 0.39, 0.30, 0.30, 0.31, 0.36. Measurements

of a few typical piinctipennis from North Carolina, Illinois, and Massachusetts gave ratios of 0.43, 0.55, 0.40, 0.46, and 0.58.

The difference in the eggs, as described by Bellamy, was chiefly in the

width and shape of the exposed exochorion dorsally between the floats. The

P'Linctipennis egg has a smooth-edged, elongate, oval- or slipper-shaped exposure, narrowest in the middle and continuous, with a high even frill. The

perplexens egg has a much narrower area of exposed exochorion, the frill

is low and follows the wavy, irregular course of the edge of the exochorion.

Measurements gave a mean ratio of the width at the narrowest point to total

egg length of 0.104, ranging from 0.092 to 0.109 for piuictipennis; and 0.049,

with a range of 0.035 to 0.061 for perplexens. Photographs of the two kinds

of eggs showed them to be easily recognizable. . , ,

LARVA. Almost identical with larva of piinctipennis except m the branching of the antepalmate hairs of segments 4 and 5. Tabulations of the

branching of these hairs in 1,620 specimens of each species (23) showed the

following- In about 46 percent of perplexens all four of these hairs were

single, and in an additional 40 percent, two or three of the four were

single In the rest, three or all four of the hairs were double. In punctipennis none of the specimens had all four of the hairs single and only

1 percent had two or three of them single. In about 3 percent one of the

hairs was single, in 39 percent all four hairs were double, and in the rest

some of the hairs were three- and four-branched. In these records, branching

beyond the middle of the hair was disregarded. Based on these counts, it

appears that larvae with all four hairs single can be identified with con-
98 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

fidence as perplcxens, and those with two or three of the four hairs single

are highly probables. Miles (229), in a study of the antepalmate hairs of

punctipennis larvae, found a small proportion with these hairs single. He

does not show whether all four were single in any specimens and, of course,

it is possible that he had a few specimens of perplexens m his series. As

previously mentioned. Roth {272) found several larvae with simple antepalmates in the same localities at Paris, Tenn., and Swannanoa, N. C, from

which adults with repressed costal spots were taken.

ANOPHELES PSEUDOPUNCTIPENNIS Theob.

Anopheles pseudo2mnctipennis represents a complex of^ forms

of which at least two, A. ^jseudopunctipennis and A. franciscanus

McC, are represented in the fauna of the Southwest. The adults

superficially resemble those of A. 2^'^nctipennis but their whitebanded palpi prevent confusion. A. pseiidopmictivemiis ranges

northward from Argentina and Chile to Colorado and Kansas and

eastward into the Mississippi Valley States of Missouri, Arkansas,

Louisiana, Mississippi, and Tennessee. It is extremely rare in all

these States and its habits in this area have not been observed. In

other parts of its range the larvae occur in clear, sunlit pools

containing algae, and along the vegetated margins of slow-flowing

streams. The adults are said to enter houses and to bite man

readily, although most feeding takes place out-of-doors on other

animals. The species has been infected experimentally with both

vivax and falciparum malaria, and has been found naturally

infected in Argentina, Mexico, and Chile, where it is considered

to be an important vector. Its importance as a vector in the United

States has never been established.

ADULT.— Proboscis and tarsi dark, palpi ringed, and wings spotted. Palpi

with the apical segment white except for a narrow black ring toward tip,

segment 4 with a narrow white ring at base. Occiput with white scales at

vertex. Integument of mesonotum with a broad median frosted stripe covered

with yellowish-white scales. Knob of halter dark-scaled. Wing costa with a

spot of white scales at the juncture of the subcostal vein and another at tip

of wing; subcosta and vein 1 with a spot opposite the base of vein 2; veins

3 and 5 with long areas of white centrally; veins 2 and 4 also spotted; anal

vein with the basal half white, the rest dark; fringe spots present at the

tip of each vein. Tarsi and abdomen all dark.

LARVA.—Antennal hair small, single, attached near middle of shaft.

Inner, outer, and postclypeal hairs all long, simple, the inner separated

by more than the diameter of a basal tubercle. Sutural hairs large, with six

or more branches along stem. Palmate hairs developed on segments 3 to 7,

the leaflets with a few lateral serrations and tapered to a sharp point.

Antepalmate hairs single on segments 4 and 5. Lateral hairs on segments 4

and 5 well branched. Postspiracular plate with a peculiar slender projection,

or tail, on each side pointing backwards.

ANOPHELES PUNCTIPENNIS (Say)

(Syn., Cidex hyemalis (Fitch))

Anopheles punctipennis is the most widely distributed anopheline in North America. It ranges from the Atlantic to the

Pacific and from Canada to Mexico. The conspicuous white spot

on the costa (pi. 8, C), combined with unhanded palpi, is the

chief recognition character. It has a variety of breeding places.

Throughout the southern range it occurs much more commonly

late in the fall and early in the spring than in the summer, and

seems to prefer the margins of flowing streams as breeding sites,
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 99

probably because of the lower water temperatures there (13, 88,

287), At Leesburg, Ga., the larvae were taken also in certain wells

and deep lime sinks, and occasionally in containers (268), At

Mound, La., where it was never abundant, it disappeared almost

entirely during the warm months. In that vicinity pure cultures

of the larvae were sometimes taken in the fall in small clay borrow

pits or pools free of vegetation. As a rule the species is rare in

central and southern Florida. Specimens of the melanistic form,

A. perplexens, were previously reported (198) from Orange,

Seminóle, and Alachua Counties. Pervlexens is now listed as a

distinct species.

The females are active and bite principally at night, but will

attack during the daytime in dense shade, or on cloudy days.

Freeborn (115) states that pimctipennis has been described as a

*'porch biter'' and, although frequently taken in barns and other

outhouses, it is seldom taken in homes. The species readily becomes

infected with malaria parasites under experimental conditions,

but it does not feed extensively on man in nature, and the epidemiological evidence indicates that it is not an important carrier

of the disease. It has been proven an efficient host of the dog

heartworm Dirofilaria immitis (2U7, 886).

ADULT.—Easily distinguished from pseudopunctipemiis by the lack of

white scaling on the palpi and on wing veins 3 and 5. Proboscis and palpi

all dark. Occiput with white scales on vertex. Integument of mesonotum with

a broad frosted median stripe covered with pale scales. Wing costa with

a white spot at juncture of subcostal vein and at tip, the former about half

as long as the subapical dark spot distal to it and continued over vein 1

and the stem of vein 2, the apical spot continued over the tip of vein 1;

subcosta and vein 1 without a white spot opposite the base of vein 2, forks

of veins 2 and 4 each with a white spot in middle, and the stem of 4 with

several spots; veins 3 and 5 entirely dark-scaled. Anal vein with the apical

half and the basal one-fourth or one-fifth dark, separated by white. Tarsi

and abdomen all dark. The width of the costal spot as compared with that

of perplexens is discussed under the latter species.

LARVA.—Antennal hair small, branched, inserted at about basal third of

shaft. Inner clypeal hairs closely approximated, the basal tubercles nearly

touching; outer clypeals densely branched; postclypeals small, usually single

or forked at tip; sometimes three- or four-branched. Hair 1 of prothorax

small, with several branches near or beyond the middle. Palmate hairs well

developed on segments 3 to 7, nearly equal in size on all segments, and with

the margin of the leaflets notched or serrated. Hair 0 minute on segments

4 and 5. Antepalmate hair on segment 4 and 5 usually double or triple,

sometimes with one or two of the four hairs single, or four-branched. (See

under perplexeres for the percentages with different branching.) Lateral

hairs on segments 4 and 5 with the branches all arising from the same point.

ANOPHELES QUADRIMACULATUS Say

(Syn., A. annuUmanus V. d. W. ; the common malaria mosquito)

Anopheles quadrimacidatus is a fairly large mosquito, dark

brown in color, with four darker spots near the center of the wing

field (pi. 8, G). In its resting position the angle at which the body

is held is not so pronounced as with some of the other species,

and the position of specimens heavy with blood may not appear

characteristic.

This species is active principally at night, although during the

cooler months the females will seek blood meals in the daylight

on warm days, in dwellings, or in the woods. The daily night or

dispersal period begins just at dusk and continues for half an
100 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

hour or so. During the remainder of the night, flight is probably

limited for the most part to local forays in search of a host.

Another period of activity begins just at daylight and ends with

a general shift to the daytime resting places. The flight range of

quadrimaculatus from the breeding places undoubtedly varies a

good deal, probably depending largely upon the numbers produced

and the proximity of blood meals. Although flights up to 4 miles

and over have been recorded for the species, few individuals reach

such distances, and in planning control operations the maximum

effective flight range may be taken as about 1 mile or less under

average conditions during the summer. Control over such restricted

areas proved highly successful in preventing malaria transmission

in areas of military importance in this country during World

War II.

Although little is known of the extent to which this species

feeds upon wild animals, man and most of the domestic animals

are known to be attacked by the blood-hungry females. Information

on the relative attractiveness of different hosts was obtained in a

series of cage experiments conducted in Baltimore, Md. (58, 59),

Two host animals were exposed side by side to the bites of quadrimaculatus females, which were afterwards collected and the blood

meals identified by the precipitin test. Among cattle and horses it

was found that the attractiveness varied more between individuals

than between the species, and that a decided variation occurred

between individuals of the human race. The man received, on an

average, about one-sixth as many bites as the horse or cow. Sheep,

goats, dogs, and pigs appeared to be less attractive, in the order

given, while rabbits and chickens proved to be very poor hosts

even in the absence of other animals.

To determine the proportion of mosquitoes that obtained blood

meals from different hosts under natural conditions, a large series

of records had previously been obtained at Mound, La., by testing

the blood from freshly fed females collected from the tenant

dwellings and outbuildings on three plantations (iP-4). From a

general series of collections during the mosquito season of 1922,

38 percent of the specimens taken inside the house, and about 2

percent of those from underneath the house and in the outbuildings

were found to have fed on man. The weighted average was 4.3 percent for the entire quadrimaculatus (female) population, being 6 to

8 percent when the average number of females per location was

about 200 to 500 and decreasing to 3 percent or less when the

average reached 1,500 or more. The average percentages for the

other hosts for which blood tests were made were as follows : Cow,

36; horse, 33; pig, 16; dog, 8; and other animals (chicken and

cat), 3.

Although very high malaria infection rates (10 percent or

even more) have been found among anophelines in other countries,

the percentage of infected quadrimaculatus in malarious areas in

this country appeared to be comparatively low, probably much

lower than is generally supposed. From an examination of 9,340

specimens collected on plantations in the vicinity of Mound, La.,

in 1922 {182), only 10 were found to contain the sporozoite form

of the parasite in the salivary glands and therefore to be capable

of transmitting the infection at the time of capture. This gave
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 101

a sporozoite rate of 0.107 percent, or approximately 1 infective

specimen per 1,000. The annual malarial rate in humans on the

same plantations during that year was about 45 cases per 100.

At Edenton, N.C., a gland-infection rate of 0.33 percent was obtained from dissections of 1,486 mosquitoes over a period of 3

years (35),

Quadrimaculatus breeds chiefly in permanent, fresh-water

pools, ponds, and swamps that contain aquatic vegetation or floating debris (pi. 5). The larvae feed almost entirely at the water

surface, and since they seem to make no selection of material,

provided it is small enough to be ingested, the food consists of

the general variety of small organisms and detritus found at the

water surface. From a large series of observations at Mound, La.,

where qiiadrimactdatus was the predominant anopheline, Bradley

(45) reported that flagellates, diatoms, and green algae made up

a large proportion of the plankton content of the surface layer

in the natural waters of that area. The approximate average

numbers of organisms per cubic centimeter in breeding places

having more than one larva per dip were as follows for the

breeding seasons of 1928 and 1929, respectively: total plankton,

8,600 and 6,300; flagellates, 5,400 and 4,200; diatoms, 1,800 and

500; green algae (other than flagellate forms), 700 and 900.

Present in smaller numbers were the ciliates, blue-green algae,

and amoeboid protozoa, although the first two were fairly abundant

at times. The four principal genera among the flagellate forms

were Eugleiia, Chlamydomonas, Trachelomonas, and Phacus,

which composed about 75 percent of the total organisms in this

class in the places of high larval density. The report of studies

made by Boyd and Foot (37) shows a similar plankton content

in the surface waters at Edenton. A classification of ponds in

which the abundance of desmids is used as an index to the suitability of the water for specific anopheline breeding has been suggested by Frohne (116), At high summer temperatures, with an

ample food supply, larval development under natural conditions

may be completed in about 1 week, although some larvae develop

more slowly than others under the same conditions. With a minimum of 3 or 4 days for the other aquatic stages (egg and pupa)

and about 4 days for the preoviposition period, the minimum time

for a complete generation would be 14 or 15 days. At low temperatures or with scanty food supply the developmental cycle is

greatly prolonged.

This mosquito occurs throughout the Eastern, Southern, and

Midwestern States to the Great Plains, and its range extends into

southern Canada and Mexico. It is found in all of the^ Southeastern States and is the principal species concerned with malaria transmission in this region. Also, it has been proven to be

an efficient host of the dog heartworm, Dirofilaria immitis (2U7).

In laboratory experiments it failed to transmit the viruses of

eastern and western equine encephalitis (223) and its vector

potential for both types has been rated as ''poor" (72).

ADULT.—Proboscis, palpi, thorax, and tarsi all dark-scaled. Occiput with

a patch of pale scales at vertex. Wings all dark, with four distinct darker

spots, one extending across the veins from the base of vein 3 to vein 5

at the junction of the posterior cross vein, one at the base of vein 2 and its

junction with vein 1, and one each at the forks of veins 2 and 4. Knob of
102 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

halter dark-scaled. Small pale knee spots present at tips cf femora. In the

male the lobes or processes of the ninth tergite are stout, rounded and

Slually slÄ at tip. This species is the largest of local

anophelines^ with a wing length of approximately 4 5 n|m. compared with an

average length of perhaps 4.0 mm. for most of the others, and 3.0 for A.

barber!. Rubbed specimens are sometimes difficult to distmguish from A.

átropos and A. walkeri.

LARVA.-Antenna with a large, well-branched tuft inserted near middle

of shaft and its tip reaching the apical spine Inner ^^yP^^l hairs usually

simple (occasionally branched beyond middle), their bases separated by m^^^^^

than the diameter of one of the tubercles. Outer clypeals ^ensely branch^^^^^

the posterior clypeals very small, forked at tip. Sutural hairs comparatively

large, with 8 to 10 branches along the main stem. Hair 1 oí prothorax sma 1,

single. Palmate hairs well developed on segments 3 to 7, and frequently

partially developed and functional on 2. Hair 0 rudimentary, antepalmate

hairs usually single, and the lateral hairs multiple on segments 4 and 5.

ANOPHELES WALKERI Theob.

Females of Anopheles walkeri have narrow but distinct white

rings on the palpi and white knee spots at the tips of the femora.

The dark wing spots (pi. 8, E) are less pronounced than those of

A, quadrimaculatus. Florida specimens are unusually dark. This

species is widely distributed over the eastern half of the United

States from southern Canada to the Gulf of Mexico. It has been

recorded for all of the Southeastern States. Its principal breedmg

places are fresh water marshes containing luxuriant growths of

such émergents as cutgrass and cattails, together with algae and

other floating plants and debris (5, ÍG, 175). The larvae have been

taken from ricefields and from sloughs covered with water hyacinth in Louisiana (205), A study of specimens from Florida ñas

shown some larval characters distinct from those of specimens

obtained in New York State {Í6), Matheson and Hurlbut {215)

have since reported that both forms occur in the vicinity of

Ithaca, N. Y. Specimens from each State have been infected experimentally with malaria parasites (198, 21^), and one specimen

infected with the parasites in nature has been reported from

Bondurant, Ky., in the vicinity of Reelfoot Lake (7). Flights of

from 14 to 2 miles from the nearest breeding place have been

reported (8), The adults are readily attracted to light traps at

night, and the females to human hosts both in the daytime and

at night (50y 82, 175), The breeding places and the resting places

of adults usually are difficult to locate (7, 82,175). The adults seem

to prefer to rest during the day on the emergent vegetation in the

breeding areas although they also have been found in damp barns,

spring houses, and under bridges (8). Matheson and Hurlbut

(215) and Hurlbut (170) found that two distinct types of eggs

were deposited by this species, ^'summer'' and **winter*' eggs,

and they concluded that overwintering occurred, at least in the

North, in the e^^ stage.

ADULT.—Proboscis all dark. Palpi usually distinctly white at tip and with

narrow white rings at the apex of the other segments. Occiput with a small

patch of white seniles on vertex and some white hairs in frontal tuft. Mesonotum all dark. Wing length about 4 mm., all dark-scaled, with four dark

spots where the scales are dense, the spots at the same location on the veins

as those of A. quadrimaculatus but less distinct. Knob of halter pale-scaled,

sometimes mostly dark, in the northern part of range {297). Tarsi dark,

femora with small knee-spots. The female differs from quadrimaculatus in

having white-ringed palpi, pale-scaled halter knob, less distinct wing spots,
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 103

thî^^I^f^^lZAl^' ^""^ from apropos in having a patch of pale scales on

J^tf\^^^Jr!J^' ^^^^P^^^ pale-scaled halter knob, femoral knee-spots, more

distinct palpal rings and somewhat more distinct wing spots. In thé male

ipn Jlf!f/l'>f^ P^'^^^ ?'^{^"^I. ^^ *^^ Phallosome are more than half the length of the apical pair to distinguish it from A. atropas, while the lobes

r .ov? A l^""^'^^ t""^ ^^P?^'^ ^^^ "^^^'^ sl^^der than in quadrimaciilatus.

Wntrt^i^ ?''\^ ^?'\ ^^'''^y. ^^'^^^ ^^^ ^^^^ several branches inserted

W/VHT f^v.^'^i.^^^i^^: î^'^'^i'' ""^^^^'^^ ^^^^s ^^^^ sparse minute feathering toward tip the basal tubercles separated by about the diameter of one of

the tubercles; outer clypeals densely branched; postclypeals minute, forked

at tip. Sutural hairs comparatively small, with several branches. Hair 1 of

protnorax with 3 to 5 strong branches from near base. Palmate hairs well

developed on segments 3 to 7 and partially developed on 1 and 2. Hair 0

n?ÍÍÍr''^ ?^I^'*^\^î' segments 1 to 7 with three to seven branches, but

much shorter than hair 2 and much smaller than hair 0 on bradleyi. Antepalmate hair on segments 4 and 5 usually single, sometimes double. The larvae

amer irom those of quadnmaculatus and átropos in having the inner clypeal

hair feathered m the branching of hair 1 of the prothorax, in the size and

branching of hair 0 of the abdominal segments, and in other details. The

inner clypeals and hair 0 differ from most of the other species. A study

oí the chaetotaxy of the four instars has been published by Hurlbut (169).

Genus CULEX Linnaeus

The mosquitoes of this genus breed in more or less permanent

collections of water. The eggs are laid on the surface of the water

in rafts of a hundred or more (fig. 3, A), and hatch within two

or three days at summer temperatures. Breeding is continuous

during warm weather and even through the winter in warmer

parts of the South. Elsewhere, inseminated females pass the

winter in hibernation.

The female palpi are short, the male palpi longer than the

proboscis ; the male antennae bushy (fig. 1, B and C). All of our

species except three have an unhanded proboscis and tarsi and are

more or less diflicult to identify except by the male terminalia or

larvae. The scutellum of Cidex is trilobed with separated tufts of

setae, the second marginal cell of the wing is as long as or longer

than the petiole; spiracular and postspiracular bristles are lacking; hairs are absent on the underside of the subcostal vein, and

setae are absent at base of vein 1 above. The terminal segments

of the female abdomen are only slightly tapered, if at all, and

the tip has a blunt appearance in comparison with the pointed tip

of Aedes and Psorophora (fig. 17). Segment 8 is short but visible

externally, and the cerci are usually retracted and invisible.

In the males the basistyle has a large subapical lobe with

various modified appendages, but lacks a basal lobe and claspettes.

The apex of the tenth sternite bears a tuft of bristles, or a row

of stout teeth. Other characters are given in the discussion of

the subgenera.

The larvae of Culex are characterized chiefly by having from

four to six or seven pairs of ventro-lateral hair tufts or single

hairs on the air tube. In addition, the large antennal tuft in most

of the species is set in a notch or constriction at about the distal

third of the shaft, beyond which the shaft is distinctly more slender. C restitans is the only one of the local species which lacks

this constriction, and Ciiliseta melanura is the only species outside of genus Cidex which has a similar constriction. The comb

scales of Culex may be in either a single row or a large patch,

the individual scale usually rounded and fringed with subequal
104 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

spinules, but sometimes long and pointed, with the fringe limited

to the basal half. The anal segment is always ringed by the

sclerotized plate, and the ventral brush is limited to the bared

area.

Because of the difficulty of distinguishing the adults of some

species of Ciilex, it is often convenient when identifying collections, particularly those taken in light traps, to use the two subgeneric names, Culex and Melanoconion Theob., into which 14 of

the Southeastern species are divided, as follows: (Culex) baho^

mensis, nig )'ipalpus, p. pipiens, p. quinquéfasciatus, restuuns, salinarius, tarsalis; {Melanoconion) atratus, eiraticu^, iolambdis,

mulrennaniy opisthopiis, peccatoVy pilosus. The one remaining

species, territanSy is placed in subgenus Neoculex Dyar. The adults

of subgenus Melanoconion are all of small size, with wing lengths

usually from 2.5 to 3.0 mm. The lateral wing scales,^ particularly

oj mm

M & A 14065

FIGURE 25.—Types of wing scales on forks of vein 2: A, Culex (Culex);

By Culex (Melanoconion).

on the forks of vein 2, are fairly short, broad, and dense (fig. 25,

B), their length only about 2.5 to 3 times the breadth. The occiput

has some broad, appressed, usually dingy or grayish scales, either

m a large median patch, or a line around the eyes. In erraticus

especially, the ñat scales are often reduced to a single row border5 There are two types of wing scales: one type broad and closely appressed

to the veins squamae scales) ; the other, standing out at an angle from the

veins (lateral, or plume scales). The latter may be either long and slider

or short and more or less spatulate. The plume scalefare on the upper

surface of the wings on some veins and on the under surface on others
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 105

ing the eyes and are difficult to see. The examination should be

made at fairly high magnification and with good lighting. The

dorsal flat scales should not be confused with the flat, whitish

scales at each side of the head, which are present also in the

other subgenera.

In male Melanoconion the dististyle is usually enlarged apically,

and the subapical lobe of the basistyle has two subdivisions. The

tenth sternite is comb-shaped apically, with a row of short, rounded teeth, and is bent inward nearly at right angles. The paired

mesosomal plates are undivided and each has a long curved basal

hook directed ventrally (lacking in C. atratiis). The ninth tergite

has paired lobes, the shape of which is usually quite distinctive in

each species and therefore useful in identification. The Southeastern species are included in excellent taxonomic reviews of the

subgenus by Rozeboom and Komp (278) and Foote (11Â). The

former contains illustrations and keys to the male terminalia of all

the species, while the latter does the same for the larvae and pupae.

Foote has resurrected the subgenus Mochlostyrax to include C

pilosus and similar species in the American Tropics based on

larval characters. Rozeboom and Komp, however, did not consider

that the Mochlostyrax group warranted subgeneric rank, and included it only as a section of subgenus Melanoconion since the

differences in the male genitalia are minor. We agree with this

decision.

Subgenus Culex is generally larger in size, with wing lengths

of about 3.5 to 4.0 mm. The lateral wing scales are long and

slender (fig. 25, A) and the occiput is without broad appressed

scales dorsally. The dististyle of the male in all our species is

slender and moderately tapered from base to tip. The subapical

lobe of the basistyle is undivided. The tenth sternite has a dense

apical tuft of short bristles or slender spines and usually a long,

stout, curved, basal arm or a short fingerlike projection. The

paired plates of the mesosome usually have two more or less

distinct arms, or divisions. The posterior lobes of the ninth tergite

are broadly rounded, separated by a shallow indentation, and

are of little value in identification.

In our single species of Neoculex the principal superficial distinguishing character is a narrow band of white scales at the

apex of the abdominal segments, whereas the bands or lateral

spots are basal in the other species. The wing scales are slender

and the occiput is without broad scales dorsally, as in subgenus

Culex. In the male the subapical lobe of the basistyle is not distinctively divided, the tenth sternite has a row of stout teeth, as

in Melanoconion, but this portion is not so strongly bent ; the mesosome lacks the basal hook, and the outer end of each mesosomal

plate is studded with small tubercles.

A short key to all dark-legged species of Culex by the male

terminalia has been included as an aid in their identification. In

the larvae of subgenus Culex the upper and lower head hairs are

multiple, with feathered branches, and about equal in length. The

spines of the pectén usually have only a few coarse teeth on one

side of the basal half. The apical bristle of the air tube is straight.

In subgenus Melanoconion both the upper and lower head hairs

are either single or double, simple or sparsely feathered, or the
106 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

upper may be multiple, but if so, it is much shorter than the lower.

The pectén spines are finely fringed on one side from base to

tip. The apical bristle of the air tube is strongly curved, or

hooked, with one or two small teeth, or hooks, near its base. Our

one species of Neoculex has both head hairs single like Melanoconion and a few coarse teeth on the pectén spines as m Culex,

The apical spine of the air tube is straight.

KEY TO SPECIES'^

ADULTS

1. Scales of wing long and slender (fig. 25, A) \ occiput without

broad pale scales at vertex or around upper eye margin •••••• ^

Wing scales short and moderately or distinctly broadened on forks

of vein 2 (fig. 25, B) ; occiput with broad, pale or dusky, appressed scales, sometimes limited to a border along the eyes

but nearly always extending to the vertex. (If the scales on the

occiput are all narrow but the wing scales are dense and broadened, see Culiseta melanura). Small Cidex, subgenus Melanoconion °

2. Abdomen with narrow bands and lateral spots of white scales on

posterior (apical) margin of the segments ^ Subgenus Neoculex, territans

Abdomen with the white segmental bands or lateral spots basal

Subgenus Culex 3

3. Proboscis and tarsi ringed with white, the tarsal segmental rings

both basal and apical 4

Proboscis and tarsi not white-ringed 5

4. Tarsal rings broad; femora and tibiae with longitudinal lines of

white scales on the outer aspect; wings usually with a patch of

white scales at base of costa • tarsalis

Tarsal rings narrow, the apical banding usually limited to hind

feet; femora and tibiae not lined; wing scales all dark. Florida

Keys bahaTuensis

5. Abdominal segments unhanded or with narrow basal pale bands . . 6

Abdominal tergites with conspicuous basal bands of white scales.

(If femora, tibiae, and wing costa are sprinkled with white,

see Culiseta inornata) 7

6. Abdomen usually with a few yellowish or dingy white scales at the

base of some of the segments, or with narrow basal bands;

tips of segments sometimes slightly pale-scaled, and seventh

segment frequently entirely yellowish-scaled; three or four

groups of white scales on sides of thorax salinarius

Abdominal segments dark-scaled except for lateral basal white

spots, usually visible from above on some segments; scaling on

pleura frequently entirely lacking or, if present, usually limited

to less than six scales in any group nigripalpus

7. (5) Mesonotum usually with a pair, sometimes two pairs, of small

white dots near the middle (fig. 19, K), and grayish scales

around the margin; mesonotal vestiture mostly of fine, brownish,

hairlike scales and with a smooth appearance. The white dotted

Culex restuans

Mesonotum without white dots ; mesonotal scales much coarser and

grayish in color pipiens quinquefasciatus

pipiens pipiens

8. (1) Hind tarsi with very narrow apical and basal white rings on

segments 1 to 4, segment 5 nearly all white. Southern Florida

opisthopus

Tarsi all dark 9

9. Occiput with a large median triangular area of narrow scales

6 Characters for distinguishing the two species of Culiseta are included

m this key in couplets 1 and 5, in view of the difficulty of separating the two

genera in the generic key.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 107

extending nearly to the vertex, the broad appressed scales

usually reduced to a narrow line bordering the eyes; mesonotal

scaling often markedly bronzy or golden; abdominal tergites

irequently with complete basal white bands. Throughout the

Southeast ^ erraticus

Occiput mostly covered with broad appressed scales,* the narrow

curved scales usually limited to a median line or small posterior

triangular patch; mesonotal scales usually darker; abdominal

tergites usually entirely dark-scaled or with small basal lateral

or median white patches, seldom with slight basal bands

southern Florida: atratus

southern Florida: iolambdis

Florida Keys: viiiWennani

throughout the Southeast : peccator

Southeastern Coastal States: pilosus

DARK-LEGGED CULEX BY MALE TERMINALIA

1. Tenth sternite with an apical tuft of short bristles or spines;

subapical lobe of basistyle not divided Subgenus Culex 2

Tenth sternite comb-shaped apically, with a row of short, stout

teeth 6

2. Subapical lobe of basistyle with eight appendages; base of tenth

sternite produced laterally into a blunt point or a short, nearly

straight arm 3

Subapical lobe with five or six appendages; base of tenth sternite

produced into a long, stout, strongly curved arm 4

3. Ventral arm of mesosomal plate long, ribbonlike, curved sharply

outward toward basistyle at about the outer third and tapered

to a point; dorsal arm slender, pointed, lying more or less

parallel with inner margin of ventral arm and extending little

if any beyond the curved shoulder of this arm. .pipiens quinquéfasciatus

Ventral arm of mesosomal plate of same general shape as above

but somewhat shorter; dorsal arm broader, semicylindrical in

appearance, with a truncate, slightly upturned tip; this arm

placed obliquely, usually extending to or toward tip of ventral

arm pipiens pipiens

4. Mesosomal plate with a short lateral arm and a longer posterior

arm having a small tooth near base restiians

Mesosomal plate with two arms and a median row or group of

four to seven short, stout teeth 5

5. Dorsal arm of mesosomal plate short, bent in middle at a right

angle (thumblike), with a pointed tip; subapical lobe of basistyle with two setae, a leaflet, and three rods; spines of tenth

sternite all sharp and pointed salinariiis

Mesosomal plate with a stout, straight dorsal arm; lobe of basistyle with only one seta before the leaflet; spines on one side of

tuft of tenth sternite short and bluntly rounded nigripalpus

6. (1) Subapical lobe of basistyle not distinctly divided; mesosomal

plate without a basal hook; outer end of mesosomal plate studded

with small tubercles Subgenus Neocidex, territans

Subapical lobe of basistyle with distinct divisions; mesosomal plate

with a long, curved basal arm or hook (except in atratus)

Subgenus Melanoconion 7

7. Dististyle narrow, tapered to a slender tip; basistyle with one

to six broad, curved filaments near base of basal division of

subapical lobe; lobes of ninth tergite gourd-shaped atratus

Dististyle with the distal portion enlarged, snout-shaped; basistyle

without accessory filaments • 8

8. Subapical lobe of basistyle without a broadened leaflet 9

Apical division of lobe with a broadened, striated leaflet, or a

greatly expanded one •. • • ' * * V j" lil * ^^

9 Lobes of 9th tergite irregularly ovate, closely approximated; the

two arms of the basal division of subapical lobe of basistyle

unequally divided • ; • • : .• • • • • • • iolambdis

Lobes of 9th tergite elongate, fingerlike projections, widely separated- arms of the basal division of subapical lobe of equal

length opisthopus
108 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

10. Distal half of dististyle greatly enlarged, quadrate in shape with

the upper margin hirsute and the apex upturned; lobes o± ^^J ^^^^^^.

9th tergite small, ovate • • • L' ' ' W ii

Dististyle moderately enlarged apically, rounded, cap-shaped or ^^

11. LelXt^tf^s!bapi¿aí'lobe' greatly'expan^^^^ Vè^^H;,.,rAr.^

wrinkled surface, lobes of 9th tergite large, elliptical, ^^^^^^^^^^^¿.^^^^

Leaflet of""!ubapic'al' 'lobe ' narrow, * striated lengthwise, its length

three or four times the width; lobes of 9th tergite otherwise 12

12. Lobes of 9th tergite represented by a pair of fingerlike projections

from the posterior corners of a broad plate; inner, or ventral,

margin of mesosome with two long, pointed teeth •••••••••• V'^^^^'^^

Lobes of 9th tergite semi-ovate, each with a lateral thumbliKe

projection; inner margin of mesosome with a single short, stout

1 j."!,^ 17lUL7'6'yi7lQ/7lZ

LARVAE (FOURTH INSTAR)

1. Both upper and lower head hairs with three or more branches,

long . Subgenus Culex ¿

Lower head hair*single or double; the upper usually much shorter

and single, double, or multiple • '

2. Antenna rather short, of uniform shape, the hair tuft placed near

middle; air tube with four to six long single hairs irregularly

placed, and one pair of small subapical hairs, double or triple

restuans

Hair tuft of antenna placed in a constriction near outer third,

the part of shaft beyond the tuft more slender; hairs of air tube

not as above ?

3. Only two anal gills, short and bulbous; air tube spinose bahamensis

Anal gills four, cylindrical ; air tubes not spinose 4

4. Air tube with five pairs of long tufts in a nearly straight line tarsalis

Air tube with four or five pairs of tufts, the subapical pair

laterally out of line (fig. 11); hair usually little if any longer

than diameter of tube 5

5. Air tube about 4:1, the sides not parallel; lower head hairs usually

with five or more branches . . pipieiis pipiens, pipiens quinquéfasciatus

Air tube 6:1 or 7:1, sides nearly parallel; head hairs usually with

three or four branches 6

6. Thorax with fine spicules; hair 1 (inner submedian hair) of mesothorax very small, about equal in size to hair 2 nigripalpus

Thorax glabrous; hair 1 of mesothorax much longer than hair 2

saliñarius

7. (1) Pectén spines with only one to four coarse side teeth; air tube

without subdorsal hair tufts; apical bristle of air tube straight

Subgenus Neoculex, territans

Pectén spines finely fringed on one side nearly to tip ; air tube with

one or more pairs (usually two or three) of small subdorsal

hairs, double or triple; apical bristle of air tube strongly curved

or hooked Subgenus Melanoconion 8

8. Air tube short, about 3:1, with eight pairs of very long hair tufts,

the proximal two inserted before the distal end of pectén;

comb-scales 8 to 12 in an irregular row, the single scale long,

thornlike, bare, or finely fringed toward base pilosus

Air tube long and slender, 6:1 or more, with four to six pairs of

ventral tufts beyond the pectén 9

9. Comb of eighth segment in a single or partially double row, the

single scale long, thornlike, fringed on basal half; body of

larva densely pilose erraticus

Comb scales in a patch of several rows, the single scale rounded

and fringed apically 10

10. Hair tufts of air tube long, the proximal pair about twice as long

as the diameter of tube at the point of insertion, or longer, the

branches finely feathered; upper head hair usually double or

triple (usually single in iolamhdis) peccator

iolamhdis

mulrennani
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 109

Hair tufts of air tube small, little if any longer than the diameter

of tube at point of insertion; upper head hair five- to sevenbranched 11

11. Lower lateral hair on abdominal segment 1 (hair 7) single;

pectén teeth with fairly coarse side spinules; thorax and

abdomen distinctly spicular atratus

Lower lateral hair on segment 1 double, pectén teeth very finely

fringed; thorax very sparsely spicular, the abdomen glabrous

opisthopus

CuLEX ATRATUS Theob.

A small Melanoconion without special markings, this tropical

mosquito was first taken in a light trap on Boca Chica Key, Fla.,

in 1942 (195, 274). The first larvae were collected on Elliott Key

the next year in an abandoned fish pool, associated with Culex

bahamensis {331), Larvae were taken in 1946 in a brackish roadside pool on Big Pine Key associated with Culex iolambdis,

Anopheles albimanus, and Uranotaenia loivii (253), Others were

found in 1947 in a man-made well in limestone on Ramrod Key,

associated with Culex mulrennani (18). It was reported subsequently to be of widespread occurrence on the Keys. Two males

were taken also in light traps on the mainland, one at Copeland

in Collier County and one at Cortez in Manatee County (252).

Light-trap records of the Florida State Board of Health have added Lee and Dade Counties to the above list (53), Nothing is

known of its biting habits. The species is known from the West

Indies and Brazil.

ADULT.—Proboscis, thorax, wings, and tarsi all dark. Occiput with

several rows of broad, grayish scales bordering the eyes to vertex. Wing

length about 2.5 mm., the scales broadened on the forks of veins 2 and 4.

Abdomen unhanded. Male dististyle slender and gradually tapered from

base to tip. Basal division of the subapical lobe of basistyle with a single

arm having a filament at tip and a bristle from near the base; an expanded

leaflet present on the apical division of the subapical lobe; one to six broad

curved filaments near the base of the basal division. Lobes of ninth tergite

large, somewhat gourd-shaped or pear-shaped, with the neck drawn out

to a blunt point.

LARVA.—Lower head hair long, single; the upper, four- to six-branched,

less than half as long as lower. Thorax finely spicular, the abdomen less so.

Comb scales in a large patch, each rounded and completely fringed. Air

tube about 8:1, with four or five pairs of multiple ventral tufts, the proximal

pair a little longer than diameter of tube, and with two or three small

dorso-lateral tufts; pectén teeth with about 12 coarse side spinules. Gills

shorter than segment.

CULEX BAHAMENSIS D. and K.

(Syn., C. corniger of Dyar (not Theob.))

This is one of the few species of Culex with tarsal white rings.

It is a tropical species that has evidently become established in

southern Florida. It was previously listed from Knight's Key,

Elliott Key, and Key West (193). More recently it has been reported to be of widespread occurrence on the Keys (252), and

has been taken also on the mainland in Dade and Broward Counties (53), Fisk (110) collected the larvae in several breeding

places at Key West, including an underground cistern containing brackish water. Wirth (331) reported the collection of larvae

on Elliott Key in 1943 from an abandoned fish pool associated
lio AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

with C. atratus, and on Key Largo in 1944 in a brackish pothole

or pool about 10 X 15 feet in size associated with Aedes taeniorhijnchus, Dehiocerites cancer, and Culex elevator (= lolambais).

This pool contained water of about half the salt concentration ot

sea water. Nothing has been published on the habits of the adults

on the Keys. The species is known from Puerto Rico and Jamaica,

the larvae having been taken in mangrove swamps and rock holes

along the coast in the latter place. n • ^ ^Hfv.

ADULT.—Proboscis with a prominent white ring at middle Occiput with

pale narrow scales. Mesonotum dark, with paler scales laterally and around

prescutellar space, sometimes with a pair of central submedian sPots or

short streaks. Wing about 3 mm., all dark. Femora, tibiae and first tarsa

segment pale on one side, dark elsewhere. Hind tarsi with narrow apical

and basal white rings, the fore and mid tarsi with still narrower rings

usually confined to the base of the segments. , ^ ^ . . ^ A • + i.^

LARVA.—Comb scales in a patch, each scale completely fringed. Air tube

about 4:1, rather densely spiculate, with six or seven pairs of bushy ventral

tufts, the first two inserted before end of pectén; pectén spines with two

or three coarse side teeth on basal half, the distal spines more widely

spaced than the others. Plate of anal segment with a pigmented dorsal

area. Only one pair of gills, very short, bulbous.

CULEX ERRATICUS D. and K.

(Syn., C. egberti D. and K., C. Iej)ri7icei D. and K., C.

pose D. and K., etc. Also references to C. ahominator

(not D. and K.) in the Southeastern States, C. inhibitator (not D. and K.) in the United States, and

Melanoconion atratiis of Dyar (not Cidex atratus

Theob.))

This small Melanoconion has been known under numerous

names, the principal ones in the United States listed above. The

extensive synonymy of the species has been discussed by King

and Bradley (189). The larvae are found in grassy permanent

ponds and swampy places, especially those having a sparse or

moderately dense growth of duckweed (Lemna). They are by far

the commonest Melanoconion in most of the southeastern area.

Deposition of the egg raft on the upper surface of the Lemna

frond has been observed in Illinois (78) and was also observed

frequently at Mound, La.

The larvae are taken frequently in association with those of

Anopheles quadrimacidatiis. Identified as C. inhibitator, they were

reported to occur in enormous numbers in ricefields in Arkansas

in association with species of Psorophora {157), In Illinois they

were especially numerous in ponds with growths of cattail and

waterprimrose. In Tennessee, they were taken in association with

10 other species, mainly Anopheles qiiadrimaculatiis, A. punctipenniSy and Culex territans {290),

The writers have taken Melanoconion females, probably all

erraticus, in Florida and Louisiana out of doors at night, while these

mosquitoes were biting, but always in small numbers compared

with the abundance of larvae. Observations at Mound, La., indicated that the species had a preference for the blood of fowls,

and attacked them on the roosts at night. According to Thibault

{303), Culex abominator (probably erraticus) was the most

abundant and annoying species in the woods in the vicinity of

Scott, Ark., especially at dusk and early in the morning, but con-
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 111

tinued to bite throughout the day. Horsfall (156, 157), however,

did not inchide the species among those of economic importance

in southeastern Arkansas where it was developing in very large

numbers in ricefields. Ross {270) found the species abundant in

southern Illinois, and said that the adults attack with persistence

and inflict painful bites. Erraticiis adults are attracted to light

traps and have been taken in large numbers in Florida.

This species has been found in all the Southeastern States and

has been reported from Texas to North Dakota and Michigan,

and eastward to Maryland. It is widely distributed also in the

Caribbean region.

ADULT.—Proboscis, thorax, wings, and legs all dark. Occiput with a large

dorsal triangular patch of lanceolate scales and one or two rows of broad

greyish scales bordering the eyes to the vertex, with the usual lateral

patches of broad appressed whitish scales. Mesonotal scaling often bronzy

or golden, especially in the western part of its range. Wing 2.5 to 3.0 mm.,

with short, broadened scales, especially on the forked veins. Abdominal

tergites with narrow, white basal bands or lateral spots. In the male the

swelling of the dististyle is moderate and gradually tapered. Subapical

lobe of basistyle with an expanded, wrinkled leaflet on the apical division;

basal division divided into two subequal arms, each with a stout apical

filament. Plate of mesosome with an apical and subapical tooth on the

outer margin. Lobes of the ninth tergite large, ovate, diverging at an angle,

and covered with slender bristles.

LARVA.—Lower head hair single, the upper much shorter, multiple. Body

spicular-pilose, usually densely so. Comb scales long, pointed, fringed

laterally on basal half, in a single irregular or partly double row. Air tube

about 6:1, usually with five pairs of multiple ventral tufts inserted beyond

pectén, the proximal pair about twice as long as the diameter of tube;

two pairs of small subdorsal tufts; pectén spines fringed on one side to tip.

Anal gills somewhat shorter than the segment.

CuLEX lOLAMBDis Dyar

(Syn., C. elevator of Wirth (not D. and K.) )

This tropical Melanoconion was first reported in the United

States by Wirth {3SI) based on a collection of larvae and pupae

obtained in 1944 on Key Largo, Fla., and originally identified as

Culex elevator. A collection was made on Big Pine Key in 1946

{253) and the next year collections were made on Ramrod Key

and Key Vaca {252), These specimens were all identified as

elevator but the identification was corrected to C. iolambdis by

Pratt and Seabrook {250), who reported the species from Martm

County and several localities in Palm Beach County on the mamland. Records of the Florida State Board of Health have added

Dade, Collier, Lee, and Manatee Counties to the list, the last three

being on the west coast. , ^ . . j_ i

The larvae on Key Largo were taken m a brackish-water pool,

about 10 X 15 feet in size, associated with C. bahamensis and

Aedes taeniorhynchus. The water in the pool contained salt m

about half the concentration of sea water. The collection on Big

Pine Key was in a brackish roadside pool, and those m Faim

Beach and Martin Counties were in pools among the aerial roots

of black mangrove, or in mixed stands of black and white mangrove. Similar breeding habits were reported for the species m

Panama and Puerto Rico {250), The adults are attracted to hght

traps and were taken in Florida ^^f^f ,^^. ^^^f^^^

decaying branches. There are no reports of their bitmg habits.
112 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

ADULT.—Proboscis, thorax, wings, and tarsi all dark occiput covered

almost entirely with broad, appressed, dusky scales Wing length about

2.5 mm., with short, broad scales particularly «« the forks oí vein ^.

Abdomen entirely dark dorsally. Male dististyle moderately 9^^^"^^^^^.^"^^^:

ally and tapered toward tip; apical lobe of basistyle lacking b^^^^^

leaflet; basal lobe divided into two arms, each with a long ^^"^^^^^^^Ï'I

of about equal size. Plate of mesosome with a strong subapical tooth on tne

outer border and several minute teeth near tip. Lobes of ninth tergite

ovate, moderately large. „ • i -.e j^,,-ki^

LARVA.—Lower head hair single, the upper usually single or if double,

not split to base, about half as long as lower. Thorax and abdomen sparsely

pilose. Comb scales in a large patch, each rounded and fringed. Air tube

about 6:1, with four or five pairs of multiple tufts, the proximal tults

about twice as long as the diameter of the tube, inserted beyond pectén;

spines of pectén fringed on one side to tip with about 15 fairly coarse

spinules; tube with a dark pigmented band near middle. Anal gills somewhat shorter than segment.

CuLEX MULRENNANi Basliam

This species of Melanoconion is known only from the Florida

Keys. It was described by Basham (18) from seven males collected

in 1945 from limestone solution holes on Big Pine Key, Monroe

County, Fla., and from larvae and reared adults collected in 1947

from potholes and a man-made well in limestone formation on

Big Pine and Ramrod Keys. Among the paratype specimens are

Hsted three larvae collected from Cudjoe Key in 1945. The larvae

in the well on Ramrod Key were associated with C, atratus and

an unidentified species of Melanoconion. Two of the holes on Big

Pine Key were only 6 and 8 inches in diameter at the top and

about 8 feet deep, with the water surface 6 feet below ground

level. A few adults were collected from the sides of the solution

holes. There was no record of biting. Three additional males were

collected in light traps on Big Pine Key in 1953 (53),

ADULT.—A very small species with the proboscis, thorax, wings, and tarsi

all dark. Occiput almost entirely covered with broad appressed ^ dusky or

opalescent scales, and a few narrow curved scales posteriorly. Wing length

about 2.0 mm., with broad, ovate scales on the forked veins. Abdomen with

minute basal lateral white spots, larger in the male. Apical half of male

dististyle greatly enlarged and then tapered to the tip. Apical division

of the subapical lobe of basistyle bears a moderately broadened, striated

leaflet, three or four times as long as the greatest width; basal division

divided into two arms, each with a stout filament of about equal size at tip.

Lobes of the ninth tergite much enlarged, each with an outer thumblike

or conical projection, bristly at tip.

LARVA.—Lower head hair single, the upper about half as long, usually

triple, sometimes two- or four-branched. Thorax sparsely spiculate, abdomen

apparently glabrous. Comb scales in a large patch, each rounded and fringed.

Air tube about 6:1, with five or six long multiple ventral tufts beyond pectén,

and two pairs of small subdorsal tufts; pectén teeth fringed on one side

to tip with coarse spinules. Anal gills shorter than segment.

CULEX NIGRIPALPUS Theob.

(Syn., C. similis Theob., etc.)

A medium-size, dark Culex with the abdomen unhanded. The

larvae are common in ditches, fresh-water marshes, and grassy

pools in central and southern Florida, where the species appears

to have largely replaced its near relative, Culex salimirius. They

are also found commonly on the Keys {252), Elsewhere in the

Southeast they are usually of scattered or rare occurrence.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 113

Judging by the small number of biting records obtained by

the writers in Florida in comparison with the abundance of the

larvae, the species is much less inclined to attack persons than is

salinarius. The adults are attracted to light traps and have been

taken occasionally resting in houses. It is not considered of importance as a pest of man. Provost (258) has published some interesting comparative collection records for this and other species

obtained in a series of night collections in 1954 at Leesburg, Fla.

The total numbers of female Culex (Culex), most of which were

said to be nigrwalvns, taken during 48 nights in different types

of collections, and the percentage of the total specimens caught

by that method were as follows: Truck trap, 1,003 (12) ; light

traps, 545 (9) ; goat-baited animal trap, 179 (9) ; calf-baited

trap, 3,689 (35), and biting collections on man, 4 (0.3). The

truck trap consisted of a screened cone mounted on top of a small

truck that was driven around the area and was considered the

standard in showing the relative proportions of the different

species. It will be noted that the calf-baited trap was by far the

most attractive of the several types.

Since our previous publication {198), nigripalvus has been recorded from Mississippi {337, 2^5), Tennessee {228, 290), and

North Carolina {66), With the exception of Arkansas, this completes the list of the Southeastern States. It is known also from

Texas, Mexico, Central America, and northern South America.

ADULT.—Proboscis, thorax, wings, and tarsi all dark, and distinctlydarker in appearance than the related C. salinarius. Occiput with onlylanceolate and upright forked scales. Pleura with very few scales, usuallylimited to less than a half dozen on any of the sclerites, or entirely bare.

Wing about 3.5 mm., the scales long and narrow. Abdomen dark above

except for small lateral white spots on part of the segments. Subapical

lobe of dististyle with five appendages, consisting of a seta followed by

a moderate-size leaflet, and three long rods, hooked at tip. Mesosomal

plate divided into two arms with a median row of four to seven teeth,

the dorsal arm straight, stout, arising near base of plate. Tenth sternite

with the usual pointed bristles on the inner side, but with short blunt

spines on the outer side; basal arm long and strongly curved.

LARVA.—Upper and lower head hairs both long, subequal, and usually

three-branched. Thorax sparsely covered with fine spicules. Hair 1 (inner

submedian) of mesothorax about equal in size to hair 2 and separated from

it about the same distance as hair 2 from hair 8 (88). Comb scales in a

patch, each rounded and fringed. Air tube slender, about 6:1 or 7:1, with

four pairs of hair tufts, the middle two placed more laterally than the

other two, the proximal tuft usually double or long and single, occasionally

triple, the next two tufts usually triple; pectén short, the single spine

with four to six coarse teeth on one side nearly to tip. Lateral hair of anal

segment usually single. Gills somewhat longer than the segment.

CULEX OPISTHOPUS Komp

This small tropical Melanoconion is the only United States member of the subgenus with white rings on the hind tarsi. It was

first recorded from Florida by Pratt et al {251), who collected

larvae and pupae from several holes of the land crab, Cardisoma

guanhumi Latr., in a cypress and maple swamp about 3.5 miles

inland from the coast near Fort Lauderdale, Fla. They were

associated with Deinocerites cancer. These authors also state that

the species was taken in Puerto Rico in sluggish streams and

streamside pools among rank vegetation, where it was associated

with Culex erraticus, Uranotaenia lowii, et cetera. Subsequently,
114 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

adults were taken in light-trap collections ^^J.^'^l^^^^^

Jupiter, Cape Sable, and Charlotte Harbor (252) ^^^^f^'J^' ¿e

Dade, Collier, and Brevard Counties (53). « thus appea^^ to De

widespread in southern Florida a though ''?}j,^\.'^^^^

the Keys. It has been reported also from Central America ana

^SÏLT.-Proboscis unhanded. Occiput -^h a rather^r^^^^

of broad dark scales anteriorly, ^^^^^^^f MP.oÄm dark except foV

wide submedian stripes of narrow pale scales f^l^^^^^^^^^^

small spots of paler scales on the ^1^«^^, ^ cffLflv bSened scales on the

space. Wing length about 2.5 mm., with distinctly broadeneascai^^

forks of veins 2 and 4, and the scales dense on the apical PO^o^ns mnu

ifrsf wihTery\arrow' apical and basal white -^^^^^^l,'^!^^^^ fj[

and segment 5 all white or nearly so; fore and f'^.^^"^'^^^^^^

Abdomen all dark above with small lateral white spots on ^^^J^^^^^^^

Male abdominal tergites with distinct white basal bands llfj^f^^^^^^^

ally. Male dististyle only slightly enlarged toward apex Apic^^^^^

subapical lobe without a leaflet; the basal division divided ^^to two eq^^^^

short arms each with a long subequal f^^^^''^'.^^ll'^J^f Ä. of n^lth

apically with a projection or tooth on the outside border. ^0"^^^. ^^_^^^^^

teSe elongate, togerlike, with a wrinkled tip, ^^\%'^^lf^^^^^

LARVA.—Lower head hair single, the upper usually ^^e-b IOK.^?.!

half tJthree-fourths as long as lower, both finely frayed. Abdomen glabrous

the thorax slightly spicular or sometimes ^l^'P^^^^'f^.f^^^^^

dorsally. Lower lateral hair on abdominal segment 1 (hair '7) double {lUh

Comb scales in a large patch, each rounded and fringed. Air tube about

sl^l wfth four pairs Sf small Ventral hairs, the distal two usually double

the proximal two triple, little if any longer than diameter of tube; pectén

tooth very finely fringed to tip. Gills shorter than anal segment.

CuLEX PECCATOR D. and K.

(Syn., C. incriminator D. and K.)

The larvae of this small Melanoconion occur in grassy pools

and swampy places, almost always associated with Culex territans.

Nothing is known of its blood-feeding habits. The species is rare

but has been taken in scattered localities in all the Southeastern

States. It has been recorded from Texas to Michigan and eastward to Delaware. It is also reported from Puerto Rico (llJf).

ADULT.—Proboscis, thorax, wings, and tarsi all dark. Occiput with a

fairly large dorsal area of broad dusky scales. Wing with short, broadened

scales on the forked veins. Abdomen with lateral white spots and usually

with small median basal spots on segments 2 to 4. In the male the apical

half of the dististyle is greatly enlarged, quadrate, the upper margin

hirsute. A very large wrinkled leaflet present on the subapical lobe of the

basistyle; basal division with a single arm with a stout apical filament,

and a second long, slender filament arising from a tubercle near its base.

Plate of mesosome with an apical and subapical tooth or projection. Lobes

of ninth tergite small, ovate with long setae.

LARVA.—Lower head hair single, the upper usually two- or three-branched,

about half as long as lower. Body sparsely spicular. Comb scales in a patch,

each rounded and fringed. Air tube about 6:1, usually with five pairs of

multiple ventral tufts inserted beyond pectén, the proximal pair about

twice as long as diameter of tube; spines of pectén fringed on one side to

tip with moderately coarse teeth; tube with a pigmented band at the

middle. Gills shorter than the anal segment.

The larval description by Dyar and Barret (97) was based on Culex

erraticus. The description was corrected by King and Bradley (189),

CULEX PILOSUS (D. and K.)

(Syn., Mochlostyrax floridanus D. and K. etc.)

This very small Melanoconion breeds in shallow, grassy pools,
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 115

roadside ditches, hoof prints, and flooded areas. The eggs are

able to withstand drying, a very unusual trait in the genus, and

as a rule the breeding places are temporary. The collected larvae

are easily recognized by their peculiar shimmying motions and

by their habit of lying on their backs on the bottom of the container. Nothing is known of the habits of the adult. The species

is common in Florida, including the Keys, and is often taken in

large numbers in light traps.

C. pilosiis was previously recorded from all the Southeastern

States except Arkansas and Tennessee (193). In the United States

it IS known only from this area, but is widely distributed in the

Caribbean area, as well as Mexico, and the northern part of South

America.

ADULT.—Proboscis, thorax, wings, and tarsi all dark. Occiput with a

moderately large dorsal area of broad dusky scales bordering the eyes and

the usual triangular patch of lanceolate and upright forked scales behind.

Wmg length 2 to 2.5 mm., the forked veins with broadened scales. Apical

part of dististyle of male enlarged, cap-shaped, and abruptly tapered.

Subapical lobe with a rather narrow striated leaflet, the basal division

of lobe with two subequal arms, each with a long stout ñlament, hooked at

tip. Plate of mesosome with a small apical and subapical tooth on the

outer margin, and two long, pointed projections from the inner margin.

Ninth tergite with a widened plate, the lobes projecting backwards from

its posterior corners as a pair of short, thumblike arms.

LARVA.—Upper and lower head hairs both comparatively short, single,

the upper rarely double. At the base of each antenna, inserted near the

preantennal tuft, is a unique, stalked, ovoid trachéal gill. Thorax slightly

spicular. Comb with 8 to 12 scales in an irregular row, the single scale

long, pointed, and with or without a minute lateral fringe basally. Air tube

small, about 3:1, with about eight pairs of very long multiple tufts, the

proximal two inserted before the end of the pectén, the others decreasing in

length; two pairs of subdorsal tufts; pectén spines fringed to the tip. The

two pairs of gills are of unequal length, the ventral pair nearly twice as

long as the segment.

THE CULEX PIPIENS Complex

The so-called Culex pipiens complex consists of a number of

closely related species, subspecies, and ecological variants occurring in various parts of the world whose exact relationship has

not been fully established. A full discussion of the problem has

been given by Mattingly et al, (218, 219). Four principal members of the complex are now recognized {217), three of which, C

pipiens pipiens L., C p. quinqnefcisciatus Say (called fatiqans in

European and certain other countries), and C. p. molestus Forskâl,

occur in the United States. The fourth member. C. p. pallens

Coq., is widely distributed in China and Japan. The recognition

of molestus in the United States is comparatively recent and very

little is known of its occurrence or distribution here. Morphologically, the arms of the mesosome are very similar to those of pipiens

(76) and''its basic diagnostic characteristics'' (201) are biological.

It has the unusual ability to produce at least the first batch of

eggs without a blood meal (autogeny) and to copulate in small

spaces (stenogamy), whereas pipiens, as now restricted, requires

a blood meal for egg production (anautogeny) and copulates on

the wing, hence requiring greater space (eurogamy). Other biological, as well as some small morphological differences, have been

described in the European area. For example, pipiens is said to

feed principally, or exclusively, on avian hosts while molestus
116 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

will feed readily on man and other mammals, as well as birds. It

is claimed that the molestiis female does not develop an adipose

body and therefore the species is unable to hibernate m the adult

stage, as is usual with pipiens. Molestus occurs in underground

sites such as subways, tunnels, and basements, where it is not

subjected to freezing temperatures. In the United States, autogenous strains have been reported by Huff (166) and Richards (m)

in Pennsylvania, Wray (333) in Illinois, Rozeboom (^7^) m Maryland, and Galindo (1943, cited by Rozeboom (276)) in California

So far, it has not been recognized in the Southern States althougn it

should occur here if it actually is restricted m its ability to

Considerable work has been done on the relationship of northern pipiens and southern qidnquefasciatus, and it has been

shown that they readily interbreed in the laboratory to produce

healthy and fertile hybrids {107, 302, 17, 277), The known range

of pipiens has now been extended well into the Southeastern

States and as the range of quinquéfasciatns extends into the

Northern and Midwestern States, there is now a broad area where

the two overlap. Male specimens with an intermediate type oí

mesosome have been collected in this area, indicating the occurrence of hybrids, which greatly complicates the matter of identification.

CULEX PIPIENS PIPIENS L.

(The northern house mosquito)

This is one of the domestic mosquitoes, and is regarded generally as the most abundant night-biting, house mosquito in cities and

towns of the Northern States. Its range also extends into the Southeastern States. It breeds in all sorts of artificial water containers, m

street gutters and catch basins, open cesspools, and in ground

pools if the water is polluted. It has always been regarded as a

serious pest and is responsible for the continuing expenditure of

large sums of money in urban antimosquito work.

As mentioned in the discussion of the pipiens complex, the

European type form is believed to feed principally or entirely on

avian hosts, and cannot ordinarily be induced to feed on man.

Colonies of northern C. p. pipiens in the laboratory in Baltimore

were never observed to bite persons either on the hands and arms

in the cages or when loose in the room (276). Other laboratories

have had difficulty in inducing them to feed on mammalian hosts

(199). Richards (26J^), however, working in Philadelphia, saw

no indication that the anautogenous strain shunned human blood ;

in fact, the contrary was true. In view of the many reports of the

pestiferous nature of the northern house mosquito, it appears that

the commoner populations differ from the typical European pipiens in this respect. Smith (292) reported that, with the exception of the salt marsh mosquitoes, pipiens was the most offensive

and troublesome mosquito in New Jersey, and in many places

the only troublesome species. He stated that it winters in the adult

stage, the favorite places being barns, cellars, and ordinary outbuildings, and that blood meals were not necessary for this species.

Howard, Dyar, and Knab (163) wrote that there is reason to

suppose that these species (pipiens and quinquéfasciatus) are
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 117

primarily persecutors of poultry. Reeves and Hammon (261)

found that of 52 freshly engorged, wild-caught females of pipiens

in the Yakima Valley, Wash., tested by the precipitin method, 76

percent had fed on birds, the rest on horses, cows, or dogs. It is

possible, of course, that molestiis is the species observed to be

annoying to man even though it can propagate without the necessity of a blood meal. If so, however, it would appear to differ from

European molestus in its ability to overwinter as fertilized females. Shute {291) has already suggested that the carrier of the

virus of St. Louis encephalitis (reported as Cidex pipiens) is not

true pipiens if this form never, or very rarely, feeds on mammals. Richards (26i) thought it probable that we do not have

an exact duplicate of the European situation. Much more study

needs to be given to the problem in the United States to clarify

the relationships. At present there is little or no information as

to the bionomics of the pipiens found in the Southeastern States.

In the North it is not considered to be a migratory species ordinarily, but may cover considerable distance at times. Adults were

taken in two light traps in Delaware Bay about 8 miles from the

nearest shore (209), and in Illinois several stained adults of both

sexes were recovered at a distance of 14 miles from the place

where they were stained (77).

C. pipiens was considered to be the principal vector of St. Louis

encephalitis during the outbreak of 1933 in St. Louis. It has been

shown to be capable of transmitting this virus experimentally

(262), and the viruses of both western equine and St. Louis encephalitis have been recovered once each from pools of specimens

collected in the Yakima Valley, Wash. (261, 138). It is considered

to be an efficient vector of human filarla and a moderately efficient

host of dog heartworm, and it was shown to be capable of transmitting fowl pox experimentally (200). o. ^i. x

Numerous additional records of C. p. pipiens m the Southeastern

States were accumulated from mosquito collections made during

World War II, and its known distribution in 1944 was given as

all of Tennessee and North Carolina, approximately the northern

halves of South Carolina, Georgia, and Alabama, and the extreme

northern part of Mississippi (317). Its distribution has been

discussed also by Barr (16).

ADULT.-Proboscis, wings, and tarsi all dark Dorsal area of the occi^u^

without broad scales. Scales on thorax l^^^^^^^^tv. ^^ ' .^prnf C vestían^ in color, and with a distinctly coarser appearance than scales of C 7 esi.a^

and C. salinarms. Wing about .4.0 mm the scales long an^^^^^

dominai tereites with broad white basal bands contznuous with the iaterai

Ss theîr S^^^^ niore or less straight not evenly rounded as

rj. p gÄe/ascmi^^ Subapical lobe of the male disti^^^^^^^^^ ei^ght

appendages consisting of a long seta, a nioderately 7/^^^^^^^/,^^^^^^

^hnrf roHc; or filaments with slightly curved or hooked tips, and three longei

ventral arm. ^ , ,, -^^íV^^íT^QI The shaüe and direction of the dorsal arm are the principal

melns oíSinfuishing C. V-fpiens f-m C . .-n^^^^^^^^^^

The DV/D ratio has been used as a measure of ^he position oi tne

two armi (302), DV being the distance between the tips of the
118 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

dorsal and ventral arms on one side and D the distance between

the tips of the dorsal arms. In a series of ^^^^'^'^^^f^'^l^l^^^

mens maintained in colonies in Baltimore, the mean for specimens

from a pipiens colony was 8.9/89 2 ^t and the nieans from two

quinquefasciatus colonies were 59.8/64.2 and 66.2/64 2 i^^-, giving ratios of about 0.1 versus 0.9 and If, with no overlapping of

the ratios, showing that the position of the ^/^^ ,"^^1,^ ^^ï^f^^

means of distinguishing typical specimens. A series of h^^rids

gave intermediate ratios, overlapping the range of measurements

for both species. A small number of wild-caught males from ditferent localities disclosed the presence of a few with intermediate

ratios in areas where the ranges of the two species overlap

Similar results with the same Baltimore colonies were obtained

by subsequent workers (17). i • .: ^

LARVA.-Upper and lower head hairs large, multip e and ^^^^1 ^n si^^^^

Body glabrous. Lateral hairs of abdominal segments 3 to ^ nearly always

double Comb scales in a patch, each rounded and fringed. Air tube about

4:1, with four pairs of hair tufts, the proximal two multiple, about as

long as diameter of the tube, the third pair double or triple and inserted

laterally out of line with the others, the distal pair usual y 4íl^ble Pectén

teeth with coarse spines on one side on basal half. Anal gills somewhat

longer than segment.

CULEX PIPIENS QUINQUEFASCIATUS Say

(Syn., C. fatigans Wied., C. pi(nge7is Wied., etc. ; the southern

house mosquito, or house Culex)

The name fatigans is employed for this species by the European

workers and is in use in most of the Old World countries. Stone

(298) has discussed the reasons for retaining the name quinquéfasciatiis which is now considered merely a subspecies of pipiens.

It is chiefly an urban domestic species and its habits are similar

to those of the northern house mosquito. It breeds prolifically

in street gutters, storm-water catch basins, cesspools, and open

septic tanks, in water barrels, tin cans, and other artificial water

containers and also in ground pools if the water is polluted. It is

generally regarded as the most abundant nightbiting house

mosquito in the cities and towns of the Southern States.

Very little information has been published on its biting habits

and host preferences in the United States. In New Orleans, La.,

the larvae were observed in enormous numbers in the ill-smelling

street gutters, before storm sewers were widely installed. The

females were a serious pest, and bed nets were required for protection against them in unscreened, or poorly screened, houses.

They were said to be a major pest at Columbus, Ga. (112), and in

southern Mississippi were reported to bite freely during twilight

and dark, and to enter buildings regularly (225). At Orlando, in

central Florida, however, observation by the present writers indicated that, although numerous, the species was of little, if any,

importance as a pest of persons. Biting collections made after

dark failed on many occasions to produce a single quinquefasciatus although they were shown to be numerous by light-trap

and rotating-trap collections, as well as by the numerous egg

rafts obtained in tubs of Dolluted water placed in the laboratory

yard for egg collecting. Efforts to obtain engorged females for an

examination of the blood were of no avail as the daytime resting
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 119

places were not located. Mosquito-control workers report that the

species is of little importance in other parts of southern Florida.

Kitzmiller (199) observed that a laboratory colony, originally obtained from Texas, fed readily on pigeons and chickens, but

reluctantly on persons.

Blood meals from 266 quiiiquefasciatiis in southern California, identified by the precipitin method, showed 88 percent had

fed on birds and the rest on horse, cow, or dog (139). In Alabama,

over 99 percent of the engorged females, whose blood meals could

be determined by serological or microscopic methods, were found

to have fed on avian blood (101). These were collected at the

Alabama Agricultural Experiment Station, presumably around

poultry houses. In other parts of the world there are records of a

high percentage of fatigans containing human blood : 87 percent

of 352 females caught in dwellings in Guam, 77 percent and 50

percent of specimens caught in two houses in the Belgian Congo,

all of 96 specimens collected in dwellings in Indochina, and 94

percent of females caught in stables there (cited by Horsfall,

(160)). In Puerto Rico hundreds of females, mostly engorged,

could be collected at any time in the poorer, crowded dwellings

(2^1), In the extreme southern part of the United States breeding may continue throughout the winter and has been observed

as far north as Columbus, Ga. (112). In the more northern part

of their range they hibernate as adult females.

Experimental transmission of the virus of St. Louis encephalitis was obtained with C p. quinquéfasciatus from California, but

not with the same species from Texas (135). In laboratory tests

with the viruses of both western and eastern equine encephalitis,

its vector potential was rated as ''poor'' (72). The virus of WEE

was recovered from specimens collected in Texas (172). This

species also is a host of human filarla.

This is one of the commonest mosquitoes in the Southeastern

States. Outside of this area it has been reported as far north and

west as the District of Columbia, Missouri, Kansas, Utah, New

Mexico, and California. It is found in the Tropics and subtropics

throughout the world. .

ADULT.—Proboscis- thorax, wings, and tarsi all dark. Mesonotum with a

vestiture of grayish, lanceolate scales having a coarse appearance m

comparison with C. restuans and C. salinarius, but similar to pipiens. Wing

length about 4 mm., the scales long and slender. Abdominal segments with

prominent white bands, usually with a smoothly rounded posterior border

on segments 3 to 5, narrowly separated from the lateral spots. Typical

specimens can usually be distinguished by this character from the other

dark-legged species of subgenus Culex. The male genitalia are very similar

to pipiens except for differences in the plates of the mesosome as previously discussed under the pipiens complex and the description oí C. p.

pipiens. In typical specimens the ventral arm is narrower and the apical

portion distinctly longer and more tapered; the dorsal arm more slender,

bluntly pointed, and directed almost straight backwards, instead of obliquely, about parallel to the inner margin of the ventral arm The «lean

DV/D ratio was 0.9 and 1.0 in two series of measurements. The basal arm

of the tenth sternite is short and stubby, usually shorter than in pipiens^

LARVA—The larvae of the two species appear almost identical, and no

certain characters for separating them have been found.
120 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

CuLEX RESTUANS Theob.

(Syn , C. territans in some early literature (not of Walker) ;

the white-dotted Culex))

This medium-size species has one or two pairs ^f si^^^l ^J},^^

dots on the thorax and broad white bands on th^/^^^^^^' ¿^"^^

larvae occur in permanent or semipermanent ground pools, ditcnes,

and rainbarrels, preferring somewhat foul water especially tiiat

containing decaying grass and leaves. The ^pecies becomes locally

abundant in the South, much more so during the fall, wmter and

spring than in the summer. In northern Alabama the larvae were

found associated with Aedes thibaulti in hollow stumps and basal

cavities of gum trees (287), In Tennessee it was taken m association with 18 species, most frequently with adex apicalis (- territans Wlk.), Anopheles punctipeiinis, Ctdex pipiens qianquefasciatus, Aedes vexans, Sind Culex salinŒTius {290).

C. restuans has sometimes been regarded as a troublesome biter,

and the writers have records of their biting persons m Louisiana.

Michener (225) in southern Mississippi stated that it was rarely

seen biting even when abundant. Thibault (303) in Arkansas wrote

that it did not bite human beings very much, though it entered

houses. It seemed to prefer poultry and livestock, and also juices ot

plants. Headlee iU2) said that the adults get into houses m New

Jersey and occasion a considerable amount of annoyance.

This species has been collected in all the Southeastern States. It

is widely distributed throughout the eastern United States and

southern Canada and as far west as Utah and Wyoming, and has

been recorded in Mexico.

ADULT.—Proboscis, wings, and tarsi all dark. Mesonotum nearly always

with one or two pairs of small white dots near the middle (fig. Ip» ^)

and grayish scales around the margins, the vestiture otherwise of fine,

brownish, hairlike scales giving the mesonotum a smooth appearance m

comparison with the rougher appearance of C. p. piinens and p. qiiinquefasciatus. Wing length about 4 mm. or slightly more, the scales long and

slender. Abdomen with wide white bands continuous with the lateral spots.

In the male the appendages of the subapical lobe consist of a seta, a short

rod with a hooked tip, a small leaflet, and three slender rods with hooked

tips. Plates of mesosome each with a fairly long, slightly curved, ventral

arm and a short, stout, dorsal arm. Arms of the tenth sternite long and

moderately curved, the bristles of the apical tuft all slender and pointed.

LARVA.—Upper and lower head hairs large, multiple, and about equal in

size. Body glabrous. Comb scales in a patch, each rounded and fringed.

Air tube about 4.5:1 w^'th four pairs of hairs irregularly placed on the

sides of the tube, the first three long, single, the distal pair shorter and

double or triple; pectén spines with a few coarse teeth on basal half.

Gills about twice as long as the segment.

CULEX SALINARIUS Coq.

A medium-size, light-brown mosquito, with or without narrow

basal bands of yellowish scales on the abdominal tergites. The

larvae are found in grassy pools, ditches, and marshy places, and

sometimes also in water barrels and in the bilge water of boats.

It was named under the supposition apparently that it was a

salt-water breeder, and in the New Jersey publications it is listed

as one of the salt-marsh species. Dyar and Knab (98), however,

stated that it did not breed in salt water and subsequent observations have shown that it occurs principally in fairly clean, fresh-
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 121

water sites. No information is available as to its salt tolerance

Q -il^^/^^T^ *^ ^^^^^ ^^ ^^^^^ ^^ brackish water in salt marshes,

bniitñ {292} wrote that they occur everywhere on the marsh in

salt as well as fresh water, but as a rule more abundantly along

the edges of the high land. In Tennessee, where the species was

reported to be fairly abundant, the larvae were found associated

with a total of 14 other species, most frequently with Culex

restuaris, C apicahs (= territans), C. pipiens quinqiiefasciatiis,

RTia Anopheles punctipennis (290).

The females bite freely out of doors at night and are occasionally found m buildings. They are seldom of much importance as a

pest m the South but are regarded as important at times in New

Jersey (U2). Smith (292) stated that they bit greedily in the

early evening. In a series of collections made in Delaware (167)

C. sahnarius constituted more than half of the total mosquitoes

taken m several types of collections, and many were collected

while bitmg during the first 3-hour period after dark. Nearly 10

times as many were taken in a light trap from which CO2 was

emitted (from dry ice) than in a nearby trap using only the light.

The number in the latter trap was nearly 'equaled by that in a

third trap that was kept dark and COo alone was used as the attractant. The adults may disperse quite widely from the breeding

places. They were taken in two light traps in Delaware Bay

about 8 miles from the nearest shore (209). Three of 34 specimens

were males.

C, salinarius occurs in all the Southeastern States. It is common

along the Gulf and Atlantic Coasts and less abundant inland. It

is not abundant in peninsular Florida or the Keys, although it

became more numerous at Orlando during the winter. The species

is widespread in the Eastern and Midwestern States, and has

been reported from southern Canada and Mexico.

ADULT.—Proboscis, thorax, win^s, and tarsi all dark, with a generally

brownish or golden-brown color. Pleura with three or four groups of white

scales. Wing length about 4 mm., the scales long and slender. Abdomen

with narrow, yellowish, transverse bands, or with at le-^st a few yellowish

or dingy white scales at the base of some segments; the segments sometimes also with scattered yellowing scales apically and the seventh segment

frequently entirely yellowish scaled. In the male the appendages of the

subppical lobe consist of two setae, one with a hooked tip, followed by a

fairly large leaflet and three slender rods with hooked tips. Basal arms of

the tenth sternite long and curved, the spines or bristles of the apical tuft

all slender and pointed. Plates of the mesosome each with two arms and

a median row or group of strong pointed teeth, the ventral arm long, stout,

bluntly pointed and with a blunt tooth on the inner margin, the dorsal

arm short, bent backwards in the middle at a right angle, thumblike,

sharply pointed.

LARVA.—Upper and lower head hairs large, multiple, and equal in size.

Body glabrous. Hair 1 (inner submedian) of mesothorax much larger than

hair 2. and separated from it by a greater distance than hair 2 from 3

(88). Comb scales in a patch, each rounded and fringed. Air tube 6:1 to 8:1,

with four pairs of tufts, the first two usually three- or four-branched, the

last two usually double, the third pair set laterally on the tube, out of line

with the others; pectén teeth with a few coarse teeth on basal half. Gills

about as long as the segment.

CULEX TARSALIS Coq.

This mosquito is recognized by the wide white rings on the

proboscis, apical and basal rings on the tarsal segments, and
122 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

white lines on the legs. It is typically a western mosquito that

reaches its greatest abundance in irrigated areas oí the irreat

Plains and western regions. It has invaded the Southeast m comparatively small numbers. The larvae have been reported irom a

great variety of more or less permanent breeding p aces including

ditches, seepage areas, grassy pools and ponds, unshaded swampy

places, flood waters, and artiñcial containers. In northern Louisiana the larvae were collected once by the writers m a clay borrow

pit with Anopheles 23unctipennis. In the Tennessee Valley it

favored small-pool areas such as grassy ditches, seeps, hooi prints,

and polluted pools rather than large bodies of a more permanent

nature. It was not found in leafy pools or in rocky stream beds.

It was usually associated with Anopheles punctipenms and irequently with Culex restuans, salinarms, and territans. It occurred chiefly from late August to late November, with a peak m

September {29Í), -^ i. ^

In the Western States the females of C. tarsahs^ are said to be

troublesome biters, but show a preference for avian hosts. It is

of great economic importance because of its relation to the transmission of western equine encephalitis, of which it probably is the

chief vector among domestic fowls and wild birds. It is capable

of laboratory transmission of the disease, and the virus has been

recovered many times from wild-caught specimens.

C. tarsalis was previously listed from Arkansas, Louisiana,

Tennessee, and Florida (193). Collections during the war years

showed it to be more widely distributed in the Southeast than

previously suspected and added Georgia and Alabama (228), Mississippi (228, 225), and South Carolina (S20). This leaves North

Carolina the only State in this area from which it has not been

reported. In Florida it has been taken in light traps in the northwestern counties and along the Gulf Coast as far south as Sarasota County (5^). It occurs in the Western and Midwestern States

to Indiana, and has been found in southwestern Canada and

Mexico.

ADULT.—Proboscis with a wide white ring in the middle. Occiput with

a large median area of whitish lanceolate scales. Scales of mesonotum

mostly dark brown, usually with paler ones in short submedian lines

posteriorly and around the prescutellar space. Lobes of scutellum with dark

scales. Wing length about 4 mm., the costa and subcosta with scattered

white scales and usually a patch of white at base, the other veins all dark,

the scales long and slender. Femora and tibia pale-scaled posteriorly,

with a complete or broken line of scales on the dark outer surface, extending

onto the first tarsal segments; hind tarsal segments with comparatively

wide basal and apical rings ; the fore and mid tarsi with similar but narrower

rings on the first three or four segments. Abdomen with wide white basal

bands continuous with the lateral spots, the eighth segment sometimes

entirely white-scaled.

LARVA.—Upper and lower head hairs large, multiple. Body glabrous.

Comb scales in a patch, each rounded and fringed. Air tube about 5:1 with

five pairs of multiple tufts, the proximal ones slightly longer than the

diameter of the tube, all set in a straight line starting near or slightly

before the end of the pectén; pectén teeth with coarse side teeth on basal

half or two-thirds. Gills slightly longer than anal segment.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 123

CULEX TERRITANS Wlk.

(Syn., C. apicalis of recent authors (not of Adams),

C. saxatilis Gros., etc.)

This rather small, dark mosquito is distinguished by the presence of \yhite apical bands or lateral spots on the abdominal

segments, instead of basal as in other Culex. It is the only southeastern species of subgenus Neoculex, It was previously listed as

Culex apicahs Adams but Bohart (31) has shown that territans

is distinct from avicalis, and that the latter occurs only in the

Southwest.

The larvae of C. territans are found in grassy pools, ditches,

and open swampy places containing aquatic vegetation. They are

often associated with Anophèles. In Illinois, it was said to be a

cold-water form and practically to disappear during the hot summer {270). In Tennessee, it was found associated with a total of

17 other species, most frequently with Anopheles punctipennis

and Culex restuans {290). The species is fairly common but the

adults are seldom seen and evidently do not attack man. They

have been observed by several persons feeding on frogs, and are

beheved to live on cold-blooded animals. Matheson {213) said that

he could not induce the females to feed on him. They are able

to fly considerable distances as specimens were caught in light

traps in Delaware Bay, from which the nearest shore was about

8 miles. Two of seven specimens were males {209). C. territans

has been recorded from all the Southeastern States {193), and

is found throughout the United States, as well as in British

Columbia and in Europe.

ADULT.—Proboscis, wings, and tarsi all dark. Occiput with pale, narrow

curved scales dorsally. Mesonotal scales fine and brown or light brown

in color except for pale scales around the front margin and the prescutellar

space; lobes of scutellum also with pale scales. Wing length about 3 to

3.5 mm. Abdominal segments usually with narrow white bands on the

posterior margins, sometimes all dark except for lateral apical spots. In

the male the subapical lobe of the basistyle is undivided and without a

leaflet, the appendages consisting of three small hooked rods, serrated

distally, a still smaller rod, nonserrated, a strong bristle, and two long,

strong rods with pointed, recurved tips. Tenth sternite with a row of short

blunt teeth apically as in subgenus Melanoconion but the arm obliquely

bent instead of at right angles. Paired plates of the phallosome with a

connecting bridge near the middle (H-shaped), the part of the plate beyond

this bridge fingerlike and studded at apex. Lobes of the ninth tergite small,

discrete, and ovate.

Michener {22^) has called attention to seasonal changes found

in this species in the Southeastern States. The summer form

tends to be smaller and the abdominal bands usually have only

one or two rows of scales, sometimes represented only by lateral

spots. The winter form tends to be larger and the abdominal

bands have three, sometimes four, rows of scales, or rarely a

partial fifth. The winter larvae are larger and darker. In Florida

the summer form appears to occur throughout the year, while

the winter form of the other Southeastern States is the one found

throughout the year in the Northern States.

LARVA.—Upper and lower head hairs are usually both single, rarely

double, the upper shorter than the lower. Body spicular-pilose. Comb scales

in a patch, each rounded and fringed. Air tube about 6:1, slightly expanded

at tip with four or five pairs of multiple tufts beyond the pectén, the

proximal pair slightly longer than the diameter of tube, the distal pair
124 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

small, slightly out of line with the others; pectén spines with ^^o or three

coarse lateral teeth. Gills about equal to the anal segment m length.

Genus CULISETA Felt

(Syn., Theobaldia Neveu-Lemaire)

The eggs of Culiseta are laid in rafts, similar to those of CuUx,

and breeding is in more or less permanent water collections, ihe

female palpi are short, the male palpi somewhat longer than the

proboscis, the male antennae bushy, the scutellum trilobed, and

the postnotum bare. The wing has a tuft of hairs on the underside

at the base of the subcostal vein, the second marginal cell is as

long as or longer that its stem, spiracular bristles are present, postspiraculars absent. The tip of the abdomen is bluntly rounded,

segment 8 visible, the cerci retracted. The dististyle oí the male

is slender, unmodified; the basistyle has a basal lobe bearing

numerous bristles but is without a subapical lobe m our loca

species, and claspettes are absent. The larva has the general

characteristics of genus Ciilex but differs chiefly m having a hair

tuft at the base of the air tube and a paired ventro-lateral row ot

single hairs beyond the pectén (subgenus Culiseta), or a single

mid-ventral row of small tufts beginning before the end ot the

pectén (subgenus Climacura), Only two species of this genus are

found in the Southeast.

KEY TO SPECIES

ADULTS

1. Wing costa and vein 1, femur, and tibia sprinkled with white;

the two cross-veins in center of wing separated by less than

their length; wings unusually broad and lightly scaled, scales

long and slender. A large species subgenus Culiseta, inornata

Wings and legs not sprinkled with white; the two cross-veins

well separated; wing scales dense, broadened. Medium size

subgenus Climacura, melanura

LARVAE (FOURTH INSTAR)

1. Both upper and lower head hairs multiple, long; air tube with

a paired row of long setae beyond pectén (fig. 14, C) .. inornata

Lower head hair single, long, the upper much shorter, multiple;

air tube with a median ventral row of small multiple hair tufts

beginning before end of pectén melanura

CULISETA INORNATA (Will.)

This rather large mosquito breeds in open grassy pools, ditches,

marshes, and occasionally in artificial water receptacles. In the

North it occurs throughout the summer but in the extreme South

larvae and adults are encountered usually only during the cooler

months, and are often associated with C. restuans. In New Orleans the larvae were sometimes found in abundance during the

winter, but they disappeared completely from March to November, and the manner of passing the summer is unknown. It is

possible that the eggs can withstand drying and go through a

period of aestivation. This is suggested by the fact that larvae

have been produced from dry material collected in the bottom of

tree holes in Texas (325). It is perhaps more likely that the adults
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 125

aestivate as they apparently are unusually long-lived. Owen {21^2)

recorded an average length of life of 95.8 days, v^ith a maximum

of 145 days, for females reared in a laboratory colony in Wyoming

maintained in a basement room at a temperature of 60*=^ to 70° F.

A complete generation in this colony required about 30 days and

blood v^as required for ^gg production, human blood being used

in this case. In Tennessee this species was collected in all months

of the year except June and July. The larvae were taken in association with 11 other species, most frequently with Culex restuanSy C, apicalis (= C, territans), and Anopheles punetipennis

{290).

The species is of little importance as a pest in the South. The

writers have not observed it feeding on man, although it is said

to do so at times. In Arkansas it was annoying before dawn in

duck blinds in the rice-growing areas {160), Thibault {SOS), also

from observation in Arkansas, wrote that it preferred horses and

cattle to man and was very annoying to horses. It bit day or night,

but mostly in the evening and night. It has been noted as biting

man in several northern localities. Precipitin tests on freshly engorged females collected in the Yakima Valley, Washington,

showed that the great majority (74 out of 86) had fed on horse

or cow, whereas only one of the specimens gave a positive reaction for human blood. The others had fed on sheep or dog, or gave

no reaction.

C inornata has been recorded from all the Southeastern States

and occurs throughout the United States. Its range extends into

northern Mexico and southern Canada.

ADULT.—This is a large species, with a wing length of 5 to 6 mm.

Proboscis and palpi sprinkled with white scales. Occiput with a large

triangular area of narrow yellowish scales. Mesonotum speckled with pale

scales. Wing unusually broad and lightly scaled; costa, subcosta, and vein

1 sprinkled with white; anterior and posterior cross-veins unusually close

together, being separated by less than the length of one; a thick row of

hairs or bristles on the underside of the wing at the base of the subcosta.

Femora and tibiae, and usually the first tarsal segments, bear dark and

white scales intermixed; tarsi not ringed. The abdominal segments with

diffuse pale scaling basally and laterally, the eighth segment being entirely

pale-scaled.

LARVA.—Antennal, preantennal, and upper and lower head hairs of larva

all large and multiple; antennal hair set about the middle of the shaft,

which is not notched. Body glabrous. Comb scales in a patch, each rounded

and evenly fringed. Air tube about 3:1, with a pair of large multiple hair

tufts near base and a paired row of 15 or more single hairs beyond the

pectén; pectén spines long, slender, with a few coarse side teeth at base.

Gills about as long as segment.

CULISETA MELANURA (Coq.)

Culiseta melanura is a medium-size mosquito, resembling a

species of Culex much more than typical Culiseta, It can be recognized by its unusually long proboscis and dense wing scales.

The larvae are found in small permanent collections of water,

with or without vegetation. In Alabama they were found in water

around the bases of trees and stumps, seeming to prefer places

chosen by Aedes thibaulti {287) ; in Arkansas they occurred m

abundance in October and November in open pools m muck lands

{157) ; and in southern Mississippi they were collected m small

grassy ponds of dark, acid water in a wooded swamp, m water
126 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

beneath hollow stumps and under the roots of trees, and once

in a shaded barrel with A. trisermtns {225). In New Jersey they

were found in holes of spring water m sphagnum bogs {29^). i he

larvae are said to overwinter under the ice in the North (^5,

?//) The adults are rare and of sylvan habits. They are attracted

to 1 ght traps but apparently nothing is known «f their feeding

habits. In the laboratory, females, reared from larvae coLected

in Louisiana, fed, after being starved, on chicks ^^d rabbits They

fed readily on soaked raisins. One female from a hght-trap collection contained avian blood, as determined by th« Precipitm test

(73) In Connecticut, reared adults in a cage would not feed on a

human arm or on chicken {315). This author obtained a Culexlike egg raft from a wild-caught female, which is of interest because Dyar {9J^) stated that the eggs were laid smgiy on the water

surface, an exceptional habit, if true in this genus.

This species is now believed to be the vector of priniary importance in transmitting the virus of eastern equine encephalitis from

^ C. melanura is known to occur in all the Southeastern States,

Tennessee having been added to the previous State records {193)

to complete the list {260, 290, 228). Its range covers most of the

Eastern States and extends to the Rocky Mountains.

ADULT.—Proboscis, thorax, wings, tarsi, and dorsum of abdomen entirely

dark-scaled. Proboscis unusually long. Occiput with pale narrow scales

dorsally, no broad ones around eye margin at vertex. Wing length about

3.5 mm., the lateral scales dense, short and broadened ; anterior and

posterior cross-veins separated by more than their own length Abdomen

all dark above, with small basal white spots, laterally. Tip of abdomen

bluntly rounded, the eighth segment exserted. 4. v, 4- +i,»

LARVA.—Antennal tuft large, multiple, set in a constriction at about the

apical fourth of shaft as in Culex. Lower head hair long, single, the upper

with three or four branches, about half as long as lower. Body glabrous.

Comb, a single even row of long, barlike scales, each rounded and finely

fringed, the lateral margin scalloped. Air tube about 6:1, with a pair of

small, double or triple hairs near base, and a long single row of about

15 short, multiple hairs on the mid-ventral line of tube beginning before the

end of the pectén; pectén spines with two or three small side teeth near

base. Gills about as long as the segment.

Genus DEINOCERITES Theobald

The species of this genus breed in holes of land crabs in coastal

areas. The adults have a close general resemblence to Culex, but

differ in a number of respects. The palpi are short in both sexes

and the male antennae are unusually long being much longer than

the proboscis, with the first flagellar segment as long as several

of the succeeding segments. The male dististyle is pilose, somewhat enlarged toward the tip, the basistyle has a subapical and

a basal lobe, neither of which is divided, and the basal lobe has

two short, stout, hornlike spines. The tenth sternite has a transverse row of stout, blunt spines, the plates of the mesosome each

with two stout blunt arms, and the ninth tergite with very long

lobes, nearly as long as the basistyle. The larvae diflfer from other

mosquitoes in having a triangular pouch on each side of the head,

and the air tube has only two pairs of ventro-lateral hairs. Only

one species has been found in the Southeastern States, and this

one is limited to southern Florida.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 127

DEINOCERITES CANCER Theo)).

(Crab-hole mosquito)

The crab-hole mosquito breeds in holes made by land crabs

(Cardisoma guanhumi Latr.) in the marl soil of the coastal salt

marshes or m fresh-water, swampy places a short distance inland

m southeastern Florida. They are most prevalent in brackish or

salt-water habitats. The larvae have been found in association

with quite a number of other species, both fresh-water and saltmarsh breeders, when the surface water left by rain or high

tides had drained away. Near Fort Lauderdale they were associated with C. opisthopus in a cypress and maple swamp 3 5

miles inland (251), The adults may be found resting on the sides

oí the crab holes and are seldom seen otherwise. They are known

to attack man occasionally at night but are not regarded of any

economic importance, although they were on one occasion reported to be annoying enough to cause complaints of local residents

m the eastern section of Hollywood, Fla. The complaints came

from residential areas adjacent to undeveloped property containing a large number of crab holes. Annoyance was intense for a few

days and then continued in a diminishing degree for more than

a month (25).

In the United States Deinocerites cancer is limited to Florida,

where it has been recorded as far north as Volusia County (193),

It has been reported also from Flagler, St. Johns, Lake, Hernando, and Sarasota Counties, based on identifications of a single

female in each county {5S). It is fairly common on the Atlantic

Coast from Palm Beach County south, including the Keys. The

species occurs in the Carribean region and Mexico.

ADULT.—Proboscis, thorax, wings, tarsi, and dorsum of abdomen all

black-scaled. Occiput with a dorsal triangular patch of brownish lanceolate

scales. Mesonotum with fine hairlike scales. Sternopleuron densely shingled

with broad, dark or dingy scales, abdomen without lateral white spots, the

posterior segments compressed laterally making the height equal to the

width, the cerci exserted. Male characters as described under the genus.

LARVA.—Triangular pouch on each side of larval head prominent, nearly

as long as the head (fig. 16, B). Antennal and preantennal hairs multiple,

the former not set in a notch. Lower head hair long, single, the upper

multiple, about half as long. Comb, a large patch of scales, each rounded

and evenly fringed around apex. Air tube about 4:1, with two pairs of

ventrolateral hairs, one triple or double near middle of tube beyond pectén,

the other small, usually single near tip of tube; a dorsolateral hair also

present near tip; pectén spines long, pointed, with a long basal tooth.

Anal segment with small sclerotized plates dorsally and ventrally. Only two

anal gills, very short, bulbous.

Genus MANSONIA Blanchard

(Syn., Taeniorhynchus L.—Arr., Coquillettidia Dyar, etc.)

The species of Mansonia lay their eggs in rafts on marshes or

shallow lakes having certain kinds of aquatic vegetation. Upon

hatching, the young larvae descend below the surface of the water and attach themselves to aquatic plants by inserting the

modified tip of the air tube into the stems and roots, through

which they obtain air. The pupae also have breathing tubes specially modified for penetrating the soft tissues of plants, to which

they remain attached until ready to transform to the adult stage,
128 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

when they rise to the surface. The winter is passed in the larval

stage.

Because of their peculiar habits, Mansonia larvae are difficult

to locate and special collecting methods must be employed. The

plants to which M. pertivrhans attach themselves are usually rooted in the soft muck or vegetable debris on the bottom of the

ponds. When these plants are uprooted many of the larvae detach

themselves and will be missed if only the roots are searched. Because of this, the plants in a small area should be pulled up and

swished around in the water to induce all larvae present to become

detached. The bottom material is then scooped up with a large

strainer and examined in small quantities in shallow pans in clear

water {210). As the larvae usually remain on the bottom of the

pan, a careful search must be made. The larvae are white or pale

green in color, and their continuous movements aid in locating

them among the trash. The search is tedious and time-consuming,

but the work can be reduced somewhat by washing the material

through two or three properly graded screens to eliminate much

of the silt and coarse debris.

Another method, described by Bidlingmayer {27, 28), involves

the use of an open-end metal cylinder 13.9 inches in diameter

(1 square-foot area) and 30 inches high. This is used to enclose

several host plants, with one end sunk into the bottom of the

pond. The plants are uprooted, rinsed about in the water, and then

discarded. All broken bits of roots and stems are also removed.

An inverted metal funnel, the larger end of which fits snugly into

the cylinder, is pushed down into the cylinder until the small end

is one to two inches below the water surface. This insert is 12

inches high and the small end has an opening V\\, inches in diameter. Soldered around the middle of the cone is a small cylinder

121/2 inches high and 9 inches in diameter. Deprived of their

source of air under water, the larvae eventually come to the surface and, after emerging from the end of the funnel, they can be

removed from the small cylinder with a suction pipette. In the

second article, the author {28) described another type of insert

consisting of a cylinder in which the bottom was made up of

about 12 small pyramids, each with a base 3^ o inches square and

a height of 2% inches, with a small opening'in the tip of each.

This type could be used in as little as 4 inches of water, whereas

the cone type required about 14 inches, and tended to collect more

sediment. The traps were usually left in the breeding places for

24 hours before the larvae were collected, but a shorter time suffices when the water is warm, or when the trap is set over floating

plants. In another method {259), the same size of cylinder was

used but magnesium sulphate was employed to increase the specific gravity of the water, and cause the larvae to come to the

surface. After the plants were removed, the water was lowered in

the cylinder to a depth of about 5 inches, and 1 pound of magnesium sulphate was dissolved and mixed in the water. The rest

of the water and loose, bottom debris was then dipped out and

placed m shallow pans, from which the larvae were removed

as they surfaced.

The collection of the larvae of M. titillans and M, indiibitans,

which are mostly found on the roots of a floating plant, water-
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 129

lettuce, is much simpler. These larvae are readily taken by lifting

the host plant quickly into a pan of water for examination, or

the plants may be lifted from the water by bringing the pan or a

cloth net up under them. Some larvae usually remain attached

to the roots while others are found moving about in the dish.

Specimens of perturbans may also be found on the same plants.

The Bidlingmayer trap also may be used to collect larvae from

floating plants and, since titillans and indubitans tend to come to

the surface much taster than perturbans, the collections may be

made from the traps after 2 or 3 hours.

The pupae of all three species also attach to plants (pi. 9, A).

Dorer et al, {90) in Virginia, found that the pupae of M, perturbans did not release themselves when the host plants (arrowhead, in this case) were shaken. They were easily collected by

pulling up a clump of plants, sloshing the roots about in the

water to remove the soil and debris, and then swishing the plants

violently in the air to leave the hairlike roots clean and dry. The

attached pupae were then easily seen among the roots.

The genus Mansonia is of world-wide distribution and contains a

large number of species that occur chiefly in tropical and subtropical areas. Only three species are found in the United States. The

adults have the proboscis and tarsi ringed with white, and the wing

scales are broad and mixed brown and white. The male palpi are

long, the antennae bushy; spiracular bristles are lacking, postspiraculars present in most, but not in all, species. The tip of the

female abdomen is blunt and somewhat compressed laterally, the

eighth segment may be completely retracted and only the enlarged

cerci visible. Some species have a row or group of spines at the

tips of the seventh and eighth tergites. In the male, a basal lobe

is present on the basistyle; a subapical lobe and claspettes are

lacking. The larvae have a unique modification of the air tube

which permits penetration of the roots or underwater stems of

aquatic plants, the apical third being tapered to a point, with

several saw-teeth on one side. The air tube lacks the usual pectén.

The larval antenna is very long, nearly twice the length of the

head, with a large tuft at the basal third and a pair of bristles

arising beyond the middle of the shaft.

Two subgenera are represented by the three Southeastern

species, M. titillans and M. indubitans being placed in subgenus

Mansonia, and M. perturbans in Coquillettidia, Postspiracular

bristles are present in the former but absent in the latter. M. perturbans also lacks spines at the tips of the seventh and eighth

tergites in the female.

KEY TO SPECIES

ADULTS

1. First segment of hind tarsi with a wide pale ring at rniddle, and

hind 4ia with a preapical P^le ring; proboscis broadly ringed;

postspiracular hairs absent. Widely distributed in the United

Ss and Canada Subgenus CoquMidm perturbans

Tibia and first hind tarsal segment unhanded or bands indistinct;

postspiracular hairs present. Florida and the Tropics

^ ^ Subgenus Mansoma 2

2. Tergite 7 of' abdomen 'with 'a 'long row of pointed spines ; tergite

8 with a close-set posterior row of about 8 stout, bluntly-
130 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

rounded spines, preceded by two laterally placed groups of

10 to 12 spines; palpi not more than one-third as long as

proboscis . • :"'.:"¿"':¿\ ^'^'^^""^

Tergite 7 without a posterior row of spines; tergite 8 with two

long rows of stout pointed spines, not closely set; palpi nearly

halt as long as proboscis indubitayis

LARVAE (FOURTH INSTAR)

1. Antenna with a pair of very short bristles arising slightly

beyond the middle, less than half as long as length of shaft

beyond point of insertion; plate of anal segment without hair

tufts along the mid-ventral line, or with only one or two

minute hairs perturoans

Antenna with a pair of long bristles arising beyond middle, their

tips reaching tip of shaft; anal segment with several one- to

three-branched hairs (precratal tufts) penetrating the plate

along the mid-ventral line, more than half as long as the

diameter of segment 2

2. Comb scales rounded, fringed with stout subequal spinules . . indubitans

Comb scales long, thornlike, sparsely fringed with fine spinules

on basal half titillans

MANSONIA INDUBITANS Dyar and Shannon

This species v^as first recorded in the United States from larvae

and pupae collected on waterlettuce at Boca Raton, Fla., in 1944

(24^9). The adults are very similar to those of M, titillans and

previous identifications of this species undoubtedly included specimens of indubitans. A half-dozen specimens in the v^riters' collection, formerly identified as titillans, are all indubitans. The

latter has proved to be the more numerous and more v^idely distributed of these tw^o species in Florida. Collections recorded by

Chamberlain and DufFey (70) of material from seven localities

across the State in southern Florida gave a total of 862 indubitans

vs. 218 titillans. The principal host plant normally is v^aterlettuce, Pistia stratiotes, but in Florida the larvae have been reported

also in small numbers on pickerelweed, arrovs^head, and waterhyacinth {283). No information is available as to v^here the eggrafts are deposited, or whether or not they are attached to the

leaves of the host plant as in the case of M. titillans.

The females readily attack man and domestic animals. Both

sexes are attracted to light traps. At Leesburg, Fla., where M.

Verturbans and indubitans are both prevalent, a series of comparative collections by different methods over a period of 48

nights in October and November 1954, gave the following total

females of the two species {258) :

Method of Collection Number M. indubitans M. perturbans

Truck trap 444 1,883 3,971

Light traps Ill 3,846 495

Goat-baited trap 6 10 1,779

Calf-baited trap 45 358 6,212

Man-biting collections 288 590 742

The truck trap consisted of a large screened funnel having a

rectangular opening 2x8 feet, mounted on the cab of a small

truck that was driven about the test area for a distance of about

2.7 miles per collection ; 12 collections were made per night before

midnight. This trap was considered to give the best estimate of

the true relative prevalance of the different mosquitoes. The light

traps and animal traps were run all night, and there were twelve
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 131

biting collections per night during the period the truck trap was

operated. It is noted that indubitans was relatively much more

attracted to light traps and somewhat more so to man, but much

less attracted to goat and calf. The ratio of the total mosquitoes

biting man to the total in the truck trap was 0.31 for indubitans

compared with 0.12 for perturbans.

In previous light-trap collections in the same locality during

1949, a great preponderance of indubitans was taken (257) but,

as indicated above, this was probably a matter of relative attractiveness rather than actual predominance. Light-trap collections in Lee County in southwestern Florida likewise showed

more indubitans than perturbans (286). Development of Mansonia

larvae is extremely slow in comparison with that of other kinds

of mosquitoes, and there is thought to be only a single annual

generation, as a rule. In the light-trap collections at Leesburg

{257), indubitans showed a peak in September, with a secondary

peak in April. Whether these represented two generations or only

different broods could not be determined, but it seems most likely

that two generations occur. Some adults were collected every

month.

In addition to collections of indubitans in south and central

Florida, records of the Florida State Board of Health show its

occurrence in northern and western parts of the State from Duval

County to Escambia County at the extreme northwestern tip {53),

There is a record of one female collected in a light trap at Camp

Stewart, Hinesville, Ga., in 1945 {230). Nineteen adults, one

alive, were recovered from planes arriving in Miami in 1938

(322). The species has not been reported from the Florida Keys.

Its range includes the Caribbean region, Mexico, and northern

South America.

ADULT.—Proboscis and palpi speckled with white, the proboscis with a

narrow median white ring, the palpi about one-fourth as long as the proboscis.

Occiput with a triangular patch of pale narrow scales. Mesonotal scales brown

and yellowish intermixed. Postspiracular bristles present. Wing length

3.5 to 4 mm. The scales are very broad, speckled with white. Femur, tibia,

and first tarsal segments well sprinkled with white, the tarsal segments

all with basal white rings except the last segments of the fore and mid

feet. Abdominal segments with apical white bands; segment 8 retracted,

the tip of the tergite with a double row of stout, pointed spmes, evenly

spaced (usually require dissection to be seen) ; segment 7 without apical

spines. In the male the dististyle is greatly enlarged basally, with a fingerlike projection, and the apical half is somewhat twisted, or convoluted.

The basal lobe of the basistyle is long and slender, with a stout spine at tip.

LARVA. Antenna very long and slender with a large tuft at about the

basal third and two long bristles beyond the middle extending beyond the

tip of the shaft. Upper and lower head hairs small, multiple. Lateral spine

of maxiUa serrate on one side. Comb of about 10 uniform scales in an

even row. each rounded and fringed with fairly coarse spines, the apical

one less than twice as long as the next pair. Air tube stout, pointed, adapted

for piercing the roots or stems of aquatic plants, with a small niultiple

lateral tuft toward the base Pnd two stout dorsal bristles toward the tip;

pectén lacking. Anal segment longer than wide, ringed by plate, with a row

of four single or double hairs inserted in the plate on the mid-ventral line.

Gills about as long as the width of segment.

MANSONIA PERTURBANS (Wlk.)

As in other species of this genus, M. perturbans has very broad

wing scales, mixed brown and white. The proboscis and tarsi are
132 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

ringed with white. Breeding takes place in marshes and lakes

having a thick growth of emergent aquatic Y^getation Larval development is extremely slow, and the winter is passed m this stage

The pupal period is also long for mosquitoes, lasting 5 or ^ f^^f «^

more In northern localities larvae have been found associated with

such plants as cattail (T^/î^M), aquatic sedges (Carexh "^'f^lf^

weed (Pontederia), et cetera At Seashore »t^^e Park Va., the

principal host plant was arrowhead (Sagittarm) (90), and m New

Jersey the egg rafts were found associated with clumps of tussock

sedge (Carina; stricta). In the Lake Apopka area in central Florida

the principal host plant was the common pickerelweed (Pontederia

cordata) (pi. 9, B).ln this area and in the upper St. Johns River

basin, larvae were also obtained from the following plants, named

in the approximate order of importance (211): Cattail (chiefly

Typha latifolia), frogbit (Limnobium spongia) waterlettuce

(Pistia stratiotes), arrowhead (S. lancifolia and S. montevidensis), spatterdock {Nymphaea macrophylla) and water-hyacinth

(Piarojms crassipes). In the vicinity of Leesburg, Lake County,

only a short distance from the Lake Apopka area, sawgrass {Mariscus jamaicensis) was the predominant plant m margins ot the

lakes, and was the principal host plant for perturbans larvae {27).

Other host plants in order of importance were lizardtail {baun^

rus), arrowhead, swamp loosestrife (Decodon) and pickerelweed

About 12 other species of plants, including waterlettuce and

water-hyacinth, were listed as occasional hosts. Water-hyacmth,

although noted also as a host plant in Palm Beach County, Fla.,

(283), fortunately is not a very favorable host; otherwise, these

mosquitoes undoubtedly would be much more widely distributed

and abundant in the extreme South, where the hyacinth covers

large areas of lakes, bayous, and canals.

Throughout most of its range this species has a single generation each year, a large proportion of the adults emerging over a

comparatively short period in the late spring and early summer.

On Cape Cod, 85 percent of the larvae were in the fourth mstar

by the end of October, but remained in this stage over winter.

In the vicinity of Lake Apopka, Florida, the adults were present

from March to December, with a peak in May or the last of April,

and a secondary peak the ñrst part of August. In rearing experiments at Orlando, practically all the larvae that were hatched m

the spring reached the fourth instar and some emergence occurred

in about three months. At Leesburg, Fla., light-trap collections

in 1949 showed a peak in May and a slight rise in August (257).

It appears therefore that a partial second generation occurs in

this area. Methods for collecting the larvae and pupae have been

described in the discussion of the genus.

M. verturbans is an important pest species in many areas where

breeding conditions are favorable. The females bite readily during

the daytime in shady, moist places, but the main flight takes

place during the half hour just before and after dark. Following

this dispersion they are more active in the early part of the night

than later. Comparative records of the collection of adult females

by different methods are given in the discussion of M. indubitans.

At Leesburg. Fla., perturbans was attracted to light traps and to

man relatively much less than indubitans, but much more so to
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 133

goat and calf (258). Collections near Orlando, Fla., showed nearly

three times as many mosquitoes biting on man as were taken in

a light trap during comparable 15-minute collections (50). The

females fly considerable distances from the breeding places. They

were caught in light traps in several locations in Delaware Bay,

one of which was 8 miles to the nearest shore and 12 miles to the

farthest shore; another location was about 8 and 10 miles from

shore. Six of 105 specimens were males {209).

The species has been recorded from all the Southeastern States

and throughout most of the rest of the United States, as well as

southern Canada and Mexico. It occurs also in Europe.

The virus of eastern equine encephalitis has been recovered

from specimens collected in Georgia (16i, 197), Thus, perturbans

must be considered a probable vector of the disease. Its vector

potential for EEE was rated as ''good'' (72),

ADULT.—Proboscis, thorax, and legs well speckled with brown and white

scales. Proboscis with a wide median white ring. Spiracular and postspiracular bristles absent. Wing scales very broad, mixed brown and white.

Wing length about 4 mm. Tarsal segments all with broad white basal

rings and a broad ring in the middle of each of the first segments; hind

tibia piso with a wide white ring toward the tip. Abdominal segments with

basal lateral pale spots and sometimes with narrow basal bands. In the

male, the dististyle is enlarged apically but not basally. Basal lobe of

basistyle short, with a stout rounded rod and a slender spine at apex.

LARVA.—Antenna very long and slender, with a large tuft at the basal

third and two small bristles beyond the middle extending less than halfway

to the tip of the shaft. Lower head hair long, multiple; the upper, shorter,

also multiple. Lateral spine of maxilla smooth. Comb scales in a single

irregular rovv^, each long, thornlike, finely fringed on the sides. Air tube

without a pectén, the tip bluntly pointed, heavily sclerotized, and with

sawteeth on one side for piercing the roots and stems of aquatic plants;

a long multiple lateral tuft about the middle of the tube, a stout dorsal

bristle and a slender subdorsal one. Anal segment longer than wide,

ringed by the plate, without hair tufts piercing the plate along the midventral line or with only one or two minute hairs. Gills about as long as

the width of the segment.

MANSONIA TITILLANS (Wlk. )

This tropical species is found in the United States only in

Florida and Texas. It is very similar in habits as well as appearance to M. indubitans, and the two were formerly confused in

Florida. As with induhitans, its principal host plant is waterlettuce but larvae have been taken exceptionally from pickerelweed, arrowhead, and water-hyacinth {283), and are said to attach also to the roots of a floating grass, Paspalum repens Berg

{96). In the lower Rio Grande Valley of Texas, where the species

is common, waterlettuce is reported to be scarce. Here the larvae

were constantly associated with water-hyacinth and less frequently with cattail {100). They were easily collected from the waterhyacinth by lifting a plant quickly into a pan of water. ^ Little is known of the biting habits of M. titillans m Florida,

but in Texas it bites viciously and was second only to Psorophora

confinnis in annoyance to humans {100). In Panama, titillans is

abundant and an important pest species in some locations. It was

the most ubiquitous species in the Gatun Lake area {92) Adults

were found throughout the year. The females dispersed from the

forested areas at night and were aggressive biters, attacking

various kinds of mammals. Under natural conditions the eggs
134 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

were attached to the under surface of the leaves of wfjlettuce.

The species has not been reported from other host plants there.

Identifications of M. titillans m Florida prior to 1945 when M

indubitans was first recognized in the State undoubtedly included

both species. Subsequent records showed it to be actually less

numerous than that species when the two occurred t^^f^f^r ^(/^¿'

and it is much less widely distributed Most of the specimens recorded by these authors were from Boca ^^ton ^"d. ^est Palm

Beach in Palm Beach County, with smaller ""î^bers from Martin

and Highlands Counties. It has been recorded additionally from

Lee, Okeechobee, and Hendry Counties, and tiie southern Atlantic

coast counties from Brevard to Dade {53) The species is known

from southern Texas, Mexico, the Caribbean region, and bouth

ADULT.—This species is very similar in general appearance to M indubitans and difficult to distinguish except by the distribution of the spmes

at the tip of the seventh and eighth tergites of the ^^^^^f^^.f^^J^yj^fJ^^

differences in the female palpi and the male gentaha In ^l^'^^^J!'^ .^^^^

ment 8, there is an apical median row of about eight closely set, toothlike

spines with rounded tips, and a separated group of spines on each side,

some rounded and some bluntly pointed. In indubitans the spines are in

two regular rows, the apical row longer, the spines more widely spaced than

in titillans, and with all the spines pointed. Tergite 7 has a close-set

marginal row of stout pointed spines, which is lacking m indubitans. These

can be seen quite readily with the microscope without dissection. In the

male the base of the dististyle is less enlarged and the apical half is not

twisted. , . ., j. • XT,

LARVA —The larvae of the two species are also very similar except m the

shape of the comb scales, which in titillans are long, slender, and thornlike, with the apical spine much longer than the weak lateral spmules. The

comb consists of six to eight scales, which show a distinct increase m size

from one side to the other.

Genus ORTHOPODOMYIA Theobald

The mosquitoes of this genus breed in water in tree holes and

occasionally in artificial water containers. The eggs are laid singly

at the water's edge (163) or on the water surface (157), and

hatch in 2 or 3 days. Two species are found in the Southeastern

States and these are very difficult or impossible to distinguish

with certainty except in the larval stage. A review of their habits,

distribution, and distinguishing characters has been given by

Jenkins and Carpenter (17i). Neither species is known to be of

any importance to man.

The females of this genus have short palpi ; the males have long

palpi and bushy antennae. Spiracular and postspiracular bristles

are lacking, the posterior pronotum has two to five bristles, the

scutellum is trilobed, and segment 4 of the fore tarsus is very

short, only slightly longer than wide. The tip of the female abdomen is bluntly rounded, the eighth segment not retracted. The

species are of medium size, mottled black and white, and can be

recognized immediately by the fine, longitudinal lines of silvery

scales on the mesonotum. The male dististyle is slender, the claw

long and frayed at the tip, the basistyle with a conical basal lobe

bearing both slender and stout bristles. A claspette is lacking, the

tenth sternite has a few apical teeth, the phallosome plates are

stout, tapered, joined apically, slightly serrate on outer margin

toward the tip. The larvae are Aedes-like in appearance, with
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 135

small antennae and a single pair of ventro-lateral tufts on the air

tube. The air tube lacks a pectén, however, and the comb scales

are m two unequal rows, one composed of short-fringed scales,

the other of very long thornlike scales.

KEY TO SPECIES

LARVAE (FOURTH INSTAR )

1. Segment 8 of abdomen with a large sclerotized dorsal plate,

a similar but smaller plate usually present on segment 7 and

frequently on 6; tuft of air tube very large, with five to

twelve branches signífera

Abdommal segments without sclerotic plates; tuft of air tube

small, not longer than diameter of tube, usually three- or fourbranched ^11^^

ORTHOPODOMYIA ALBA Baker

This species is very similar in appearance to O. signífera, and

the two can be recognized immediately by the fine white lines on

the thorax. 0. alha has been collected in tree holes in several different kinds of trees and occasionally also in artificial water containers such as a flower pot (i^O), and a carbide cannister {281).

It has almost always been associated with larvae of the closely

related species 0. signífera (17i), but is much less commonly encountered. A. triseriatus has also been found with it. Nothing

apparently is known of the feeding habits of the adults.

The species was found to overwinter in New York as larvae

frozen in the ice (213, 5), but in Texas it was thought to overwinter in the adult stage as no larvae were found between November and April, and dry material from the summer tree-hole

habitats during this period failed to produce larvae. Larvae

frozen in tree-hole water in a refrigerator from two hours to two

days failed to survive {326), These authors found the larvae in

only a small percentage of the tree holes. The majority of breeding

holes had openings not more than three inches in diameter, part

of them barely one-half inch wide. The larvae were found at one

time or another associated with 0. signifera, A, triseriatus, and

Toxorhynchites r, septentrionalis. They often occurred in the absence of 0. signifera. The pH of the water ranged from 7.6 to 8.4.

Larvae were not obtained by flooding dry material from the holes

and none was found less than a week after the cavities had been

filled, indicating that the species survives periods of drought as

adults.

Although rare, the species has been recorded in scattered localities over a wide range extending from New York to Missouri

and Texas. It was originally described from near Ithaca, N. Y.

(5), and was first recorded in the South from Colbert County,

Ala. {289). Since then, it has been reported from several localities

in North Carolina {i7,17i), Mississippi {228, 225), New Orleans,

La. {24,3, HO), and Atlanta, Ga. (Stone, personal communication). There were other reports from North Carolina, Mississippi,

and Panama City, Fla., in 1944 {66) and from Hinesville, Ga., in

1944 {230), based on light-trap collections, each with only one female. In view of the uncertainty of the identification of adults,
136 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

these records must be regarded as doubtful until confirmed by

larval determination.

ADULT.—Almost identical in all characteristics with O. signífera except

for slight differences in coloration of abdominal segments 1 and 2, which

are given in the description of that species. These differences were found

to be fairly consistent in reared specimens in North Carolina (281) but

were said to be unreliable in other areas studied (174). A key to the adults

has therefore been omitted. .

LARVA.—Differs rather markedly from those of signífera, the most obvious

difference being the lack of sclerotized plates on abdominal segments 6 to 8,

which are present in the latter. Upper and lower head hairs multiple, equal

in size. Outer sutural hair large, multiple; the inner, single. Comb scales

in two dissimilar rows, the posterior one of about ten very long, thornlike

scales, the anterior row of 11 to 18 much shorter scales, part of which are

thornlike and part fringed apically. Air tube about 3:1, pectén lacking, tuft

shorter than diameter of tube, with only three to five branches. Anal

segment faintly or incompletely ringed by the plate, the lateral hair double

or triple. A very small elongate sclerotized plaque near base of segment.

Gills quite short, the ventral pair shorter than the dorsal pair. The larvae

are of whitish color compared with the reddish color of signífera. Breland

(55) has described variations found in alba larvae.

ORTHOPODOMYIA SIGNíFERA (Coq.j

This species breeds chiefly in the water in decayed holes in many

kinds of trees, and is also found at times in artificial water containers. It is associated more or less frequently with Aedes triseriatits, Toxorhyncliites sp., and Anopheles barberi, as well as 0.

alba. The eggs may be laid on the edge of the water container close

to the water line (163) or on the water surface {157). Eggs deposited on the edge of a beaker in the laboratory hatched in three

days {17Í), The larvae are said to be unable to withstand freezing (5). Thibault {303) reported it as being abundant near breeding places in Arkansas and as entering houses to bite, an observation that has not been confirmed elsewhere, although there are a

few reports of an occasional biting female. Females starved in

captivity would not attempt to bite an arm, although both sexes

readily fed on honey {17If). The females could not be induced to

feed on rabbits, hamsters, chickens, frogs, or turtles, and it was

thought that the clear liquid usually found in the gut might be

nectar, honey, or the haemolymph of some invertebrate. They

could be kept alive for over forty days on raisins (Snow, quoted

by Horsfall {160)).

0. signifera is widely distributed in the Southern and Eastern

States from Massachussetts to Nebraska and Texas. It has been

found also in several localities in California. It was previously

reported from all of the Southeastern States {193).

ADULT.—Proboscis unringed, with fine longitudinal lines of white scales,

the palpi also white-lined. Mesonotum with four pairs of fine longitudinal

white lines, one submedian pair extending two-thirds of the way back, one

pair extending completely around the lateral margin, and two shorter pairs

posteriorly, the submedian pair running across the prescutellar space and

onto the midlobe of the scutellum. Wing length 3.5 to 4 mm., the scales

broad, mixed brown and white, a spot of white scales running across the

veins from the stem of vein 2 to 5.1 and another all-white spot on the

bassl third of the anal vein. Femora and tibiae sprinkled and lined with

white scales, the hind tarsi with broad apical and basal bands on segments

1 to 4, segments 5 all white, fore and mid tarsi with slight markings on the

first two segments. Segment 1 of abdomen typically with a large central

patch of white scales, the integument brownish in color, segment 2 with

a wide basal band, not extending to the posterior margin, the other seg-
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 137

merits with large basal lateral spots and usually with narrow basal bands.

0. alba was described as having the integument of segment 1 of a yellowish

color and segment 2 almost entirely white-scaled, the white extending to

the posterior border in the middle. As mentioned under that species, however, these differences do not appear entirely reliable and identification

must depend upon the larvae.

LARVA.—Antennal hair multiple, inserted before middle of shaft. Upper

and lower head hair long, multiple, equal in size. Both sutural hairs single.

Lateral hair of abdominal segments 1 and 2 comparatively short, multiple.

Segment 8 with a large dorsal sclerotized plate, a smaller one usually present

on segment 7 and a much smaller one often present on 6. Comb scales in

two rows behind the plate, the posterior row of 8 to 10 very long thornlike

scales, thei anterior row about 17 to 24 much shorter scales, part of which

are thornlike, the others (on the ventral part of the row) rounded and

fringed apically. Air tube about 3:1, pectén lacking, the ventral tuft multiple,

much longer than diameter of tube, inserted near middle. Anal segment

ringed by plate, the lateral hair single; a small elongate sclerotized plaque

near base of segment. Dorsal pair of gills about as long as the plate, the

ventral pair shorter. The larvae have a reddish coloration.

Genus PSOROPHORA Rohineau-Desvoidy

Most of the species of Psorophora are severe biters, and at

least one of them (P. confinnis) is an important economic pest. The

breeding habits of the group are similar to those of the typical

Aedes. The eggs are adapted to withstand drying and may He

dormant on the soil of breeding sites for long periods. They hatch

upon being flooded, and the larvae may complete their development in very transient pools, as they develop very rapidly. The

larvae of two of the species (subgenus Psorophora) are predaceous upon other mosquito larvae and are therefore of some

benefit to man, but the females of these two species, the familiar

gallinippers, are themselves avid bloodsuckers.

This genus is closely related to Aedes from which it is separated

by the presence of spiracular bristles. The white scaling on the

abdomen is apical instead of basal, unless the tergites are entirely

white-scaled. The female abdomen is pointed and the eighth segment retractile as in Aedes, In the male the basistyle is without

lobes. Claspettes are present and vary greatly in shape between

the different species. The anal segment of the larva is completely

ringed by the plate and a row of hair tufts penetrate the plate on

the midventral line. The eggs of seven species have been described

by Horsfall et al {161), .. .^ ^ . ^

The twelve southeastern species of this genus are divided into

three subgenera as follows: (Psorophora) ciliata and howardn:

(Grabhamia Theob.), confinnis, discolor, pygmaea, and sigmpennis; (Janthinosoma L.-Arr.) cyanescens, ferox, hórrida, johnstonii, longipalpis, and varipes.

The species of subgenus Psorophora are very large and are

known in some areas as gallinippers. The mouth brushes of the

larvae are modified to form stout prehensile hairs for grasping

and holding their prey (fig. 16, D), each of the hairs being hooke^

at the ÜV and with a row of comblike teeth along the side, ihe

fentrolat"^^^^ of the air tube is long and single and the spines

of the Tjecten are numerous and bristlelike.

In sSbgenus Grabhamia the proboscis and tarsi are whiteringed the scales of the mesonotum are narrow-curved, the wing

and S selles are mixed brown and white, at least the hmd femur
138 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

has a narrow white ring toward the apex, and the female claws

are simple. The larvae have normal, ciliate mouth brushes, the

ventrolateral hair of the air tube is multiple and usually very

small, and the pectén spines are few and stout so that the larvae

are easily distinguished from subgenus Psorophora.

In subgenus Janthinosoma the proboscis is not ringed with

white, the mesonotum has some broad, pale scales, the wing scales

are all dark, the legs and abdomen are blue-black in color, the

white markings on the tarsi are restricted to one or more of the

last three segments of the hind feet (in one species, P. cyanescens,

the tarsi are all dark), and the female claws are toothed. The

larvae of this subgenus are not easily separated from those of

Grahhamia except by specific characters.

KEY TO SPECIES

ADULTS

1. Legs shaggy, with long erect scales or with tufts of erect scales

toward tips of hind femur and tibia; very large species of

striking appearance (fig. 2, B). The gallinippers

Subgenus Psorophora. 2

Legs not markedly shaggy and species not exceptionally large 3

2. Mesonotum with a median stripe of golden scales (ñg. 19, H) ;

proboscis and tarsi with white rings ; general color yellowish . . ciliata

Mesonotum with a wide median stripe of dark scales; proboscis

not ringed; first two segments of hind tarsi usually with very

narrow, pale basal rings; general color bluish howardii

3. Proboscis and tarsi ringed with white; legs and wings with mixed

black and white scales; hind femur with a very narrow ring of

white toward apex Subgenus Grahhamia 4

Proboscis not ringed with white; tarsal markings restricted to

one or more of the last three segments of hind feet, or entirely

dark; hind femur not ringed. Bluish Psorophora

Subgenus Janthinosoma 7

4. First hind tarsal segment dark except for a narrow basal white

ring. Florida Keys pygmaea

First hind tarsal segment ringed at base and in middle, or

largely pale 5

5. Wings without well-defined spots or areas of white scales; first

hind tarsal segment ringed with white in middle and at base.

A fairly large, blackish species, common in the Southern

States confinnis

Costa and veins with well-defined spots or areas of pale scales;

femur and first hind tarsal largely pale-scaled, intermixed with

dark, and ringed with dark scales apically 6

6. Wing costa with a spot or stripe of pale scales at base and one

opposite tip of subcostal vein; fringe spots lacking. Usually

rare southern species discolor

Apical half of costa with two white spots, none at base; pale

fringe spots present opposite the tips of all veins. Recorded only

from Arkansas and Tennessee in the Southeast signipennis

7. (3) Tarsi all dark; abdominal segments with apical, oblique,

submedian patches of pale scales cyanescens

Hind tarsi marked with white on the apical segments; abdominal

segments with only lateral apical pale spots 8

8. Fourth segment of hind tarsus white-scaled dorsally, the * fifth

segment dark 9

Last two segments of hind tarsus white , ,, , 10

9. Mesonotum with a wide median longitudinal stripe of dark scaies,

broad yellowish-white scales laterally (fig. 19, G). Found in

all Southeastern States varipes

Mesonotum clothed completely with yellowish-white scales

Florida Keys johnstonii
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 139

10. Mesonotum clothed with a mixture of brown and broad whitish

scales. Common woodland species, the white-footed mosquito .... ferox

Mesonotum with a broad median stripe of dark scales, whitish

scales laterally 11

11. Tips of femora white-scaled (knee spots) ; palpi of female less than

one-third length of proboscis. Throughout Southeast, usually

rare hórrida

Knee spots absent; female palpi more than a third as long as the

proboscis. Western Arkansas and Louisiana, rare longipalpis

LARVAE (FOURTH INSTAR)

1. Mouth brushes formed of stout prehensile hairs (fig. 16, D), each

hooked at tip and with a row of comblike teeth along the side;

pectén with the spines prolonged into hairs. Very large predaceous larvae 2

Mouth brushes normal, ciliform ; pectén not as above 3

2. Lateral hair of anal segment with three or four branches from

base ciliata

Lateral hair of anal segment single or forked some distance from

base howardii

3. Antenna very long, somewhat inñated beyond middle, two long

bristles at outer third in addition to the large central tuft; air

tube small with a paired tuft of very long hairs discolor

Antenna not as above; air tube large, more or less inflated, the

ventral tuft usually small or obsolete 4

4. Upper and lower head hairs both multiple confinnis

Head hairs single or double, rarely triple c... 5

5. Upper and lower head hairs single; antenna normal, shorter than

head 6

Upper head hair double, the lower double or triple; antenna unusually long, distinctly longer than head in most species 8

6. Antennal tuft with many branches, and preantennal tuft with at

least six signipennis

Antennal and preantennal tufts with less than six branches each 7

7. Tip of air tube with a pair of long hairs on the lateral valves;

pectén with 3 or 4 spines, each with a single basal tooth .... cyanescens

Hairs on lateral valves of air tube short, less than the diameter

of the tube at its tip ; pectén of 5 or 6 spines, each with several

teeth at base pygmaea

8. (5) Lateral abdominal hairs minute on segments 4 to 6; spines

of pectén with several teeth, subequal in size, on one or both

sides at base hórrida

Lateral abdominal hairs long enough to be readily seen; spines of

pectén with one fairly strong tooth on one or both sides at base

and usually one or two much smaller teeth 9

9. Lateral abdominal hairs single or double on segments 4 to 6 10

Lateral abdominal hairs multiple on segments 4 to 6 11

10. Antenna unusually long, nearly 1.5 times as long as head ferox

Antenna only slightly, if any, longer than head varipes

11. Lateral hair of anal segment branched near base; pectén usually

with 6 to 8 spines johnstonii

Lateral hair of anal segment branched toward tip; pectén with

3 or 4 spines longipalpis

PSOROPHORA CILIATA (F.)

(The Shaggy-Legged Gallinipper)

This is a very large mosquito (fig. 2, ß) of a general yellowishblack color, and with very shaggy legs. The larvae occur in temporary rain pools or other ñooded areas and, since they are most

often associated with larvae of Psorophora confinnis, their breeding places are the same; that is, in grassy ditches, pools, and

shallow grassy flats of all sizes, ricefields, the Florida Everglades,

et cetera They are predaceous on other larvae, but, although each
140 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

individual destroys a good many victims, they seldom occur in

large enough numbers to give an appreciable degree of control.

Several generations may occur each year depending on suitable

conditions of alternate drying and flooding of the breeding places.

Various other species of Psorophora and Aedes, including the

fellow predator, P. howcudii, often occur in the same breeding

places. Breland (56) reported that the larvae totally disregarded

adults and larvae of the fruit fly Drosophila, v^hen offered as food

in the laboratory, although Toxorhynchites r. septentrionalis

showed a more decided preference for the fruit fly adults when

placed on water surface than for other mosquito larvae.

The adults are severe biters and, because of their large size,

rather frightening in appearance. They seem to be strong fliers

and are encountered at considerable distances from the breeding

places, usually mixed with large numbers of P. confinnis. Because

of the relatively small numbers, they are not considered a major

pest.

This species has previously been reported from all the Southeastern States (193). Its range includes the Eastern and Midwestern States, southern Canada, and parts of Mexico, Central,

and South America.

ADULT.—Proboscis with a median white ring and speckled with white

toward tip. Palpi about a third as long as the proboscis. Mesonotum with a

median stripe of narrow golden scales running the full length, and lateral

areas or stripes of broad white scales, the median and lateral stripes

separated with nude stripes and narrow dark scales. Wing length about

6.5 to 7 mm., the scales dark. White basal rings present on all the tibiae,

all segments of the hind tarsi, and segments 1 to 3 of the fore and mid

tarsi; the legs, especially the hind pair, with long, dense, erect dark scales

on the apical part of the femur and most of the tibia and first tarsus.

Abdominal segment 1 with a median patch of whitish scales, the rest of the

tergites almost entirely pale-scaled.

LARVA.—Mouth brushes modified for grasping, as described previously.

Antennal hair very small, single, inserted toward apex. Lower head hair

fairly long, the upper much shorter, both branched three or four times

beyond the middle. Comb, an even curved row of long, thornlike scales. Air

tube about 3.5:1, the sides nearly straight, the ventrolateral hair long, single,

inserted beyond the pectén, the pectén spines unusually long and bristlelike.

Anal segment ringed by the plate, which is penetrated by tufts of the

ventral brush; lateral hair short, multiple, branched near base. Gills long,

pointed, about twice the length of the segment.

PSOROPHORA CONFINAIS (L.-Arr. )

(Syn., P. columbiae (D. and K.), Janthinosoma floridense

D. and K., Cidex jamaicensis of Dyar, in part (not of

Theob.), etc.; the Florida glades mosquito)

The Florida glades mosquito, long known as Psorophora columbiae, is above medium size and of a sooty black color. The proboscis

and tarsi are white-ringed, the first hind tarsal segment has a

white ring in the middle and the abdomen has diffuse apical pale

scaling or large apical patches. It breeds in temporary pools and

occurs commonly throughout the Southeast. The species becomes

a major pest in the region surrounding the Florida Everglades as

well as in the rice-growing areas of Arkansas and Louisiana

where it is sometimes called the ricefield mosquito. Other than

heavily shaded woodland pools, the larvae are found in many

types of places, but particularly in grassy ditches, open grassy
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 141

swales, ricefields and their ditches, flooded pastureland, et cetera.

They occur frequently in almost pure culture, except for the

presence of their predators P. ciliata and howardii, but are also

found on occasion with a variety of other species of Psorojjhora

and Aedes.

In Tennessee it was taken at one time or another with 12

different species, most frequently with Aedes vexans. Larval

development is rapid, requiring only 4 or 5 days in warm weather,

so that adults may apear in less than a week after the eggs are

flooded (290). Alternate drying and flooding of the breeding

sites may produce several generations annually. The winter is

passed in the egg stage.

In Arkansas the drained ricefields comprised the greatest acreage suitable for oviposition, while pastures and ditches were also

selected when conditions were suitable {159), Larvae were less

abundant in the ricefields when first fiooded than after the cultural

drainage and reflooding in midsummer, when enormous numbers

were to be found. Comparative counts of larvae were obtained

with the aid of a screen cylinder having a cross-section area of

1 square foot. This was placed upright with the lower end settled

in the mud, and the larvae were dipped out with a tea strainer.

Schwardt {282) found most of the early season breeding in ditches,

canals, seepage puddles, and places other than the ricefields on

the first fiooding. The larvae were sometimes found in numbers

following the midseason flooding, but this did not appear to be

the rule. Schwardt stated that P. confinnis comprised about 90

percent of the mosquito population in the rice region, the remaining 10 percent consisting of Anopheles quadrimaculatiis, Psorophora discolor^ and P. ciliata.

The females attack at night or in grassy or shady places during

the day. They are not as aggressive as the salt-marsh Aedes

except when a large brood appears. In the Everglades the species

occasionally appears in enormous swarms and, according to

McNeel, as cited by Bishopp {SO), has caused large losses of livestock by its attacks. During these outbreaks it is almost impossible for humans to remain out of doors at night or in sheltered

places during the day without some protection. Workers in

sugarcane fields sometimes carry smudge pots for this purpose,

and large smudges are employed for the rehef of stock. The

females disperse over huge areas and reach both coasts m large

numbers. They also appear in enormous numbers in the ricefield areas of Arkansas, and are extremely annoying to livestock,

especially cattle {159), Identification of blood meals by the precipitin method in 479 engorged females in Arkansas gave positive

reaction for bovine blood in 69 percent; swine, 16 percent; horses,

14 percent; and man, less than 1 percent (four specimens) {S2Í),

In a study of the biology of P. confinnis, Horsfall {159) found

an average preoviposition period of 8 to 9 days, with three blood

meals normally consumed during this period. One meal was not

sufficient to mature the first batch of eggs. In experiments on

the distance of flight, stained specimens were recaptured up to

about 9 miles from the point of release. P. confinms^nd P, discolor were captured in about equal numbers, and the two constituted about 95 percent of the total mosquitoes. Most of the
142 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

others were Anopheles quadrimaculatus and Culex erraticus. P.

confinnis occurs throughout the United States and its range extends

into Mexico, parts of the Caribbean region, and South America. In

most of the southeastern area P, confinnis seldom occurs m large

numbers and is considered only of minor or local importance

ADULT.—Proboscis with a wide, median white ring. Palpi about a fourth

as long as the proboscis, white-tipped. Mesonotum clothed with narrow black

scales,^well sprinkled or frosted with white. Prescutellar space circled with

pale scales. Lobes of scutellum pale-scaled. Wmg length about 4.5 mm., the

scales mixed white and black, the white not segregated into discrete spots.

Femora and tibiae speckled with white, the femora with narrow subapical

white rings and white knee spots; hind tarsi with wide basal rings and a

^^nde ring in the middle of segment 1, the fore and mid tarsi similarly marked

except that segment 5 is all dark. Abdomen with large apical triangular

patches on segments 2 and 3, and with diffuse scaling or separated submedian, apical patches on segments 4 or 5 to 7.

LARVA—Antennal hair and upper and lower head hairs large and multiple.

Comb, a single row of about six scales set on the posterior border of a small

sclerotized plate, the single scale stout, rounded, with a long central spme

and much shorter lateral spinules. Air tube about 3:1, slightly swollen with

a moderately large, multiple tuft inserted beyond the middle; pectén oí only

four or five spines. Anal segment ringed by the plate, with tufts of the

ventral brush penetrating the plate. Gills pointed, about twice as long as the

segment.

PSOROPHORA CYANESCENS (Coq.)

This blue-black Janthinosoma is the only local Psorophora in

which the tarsi are entirely dark. It is rare in most of the Southeastern States, but was reported as very abundant and annoying

at times in Arkansas (303), It was reported to be the most

abundant Psorophora in the Muscle Shoals area, Alabama (287),

and to be locally abundant as a pest in Mississippi (2^5). The

larvae occur in transient pools associated with other species of

Psorophora and Aedes. Larval development is very rapid and

was found to average 4.6 days from hatching to adult emergence

in Arkansas {282), One breeding place cited by this author was

a pool containing suspended clay and nearly free of vegetation.

Thibault {303) in Arkansas found P. cyanescens the most annoying of all mosquitoes when present in abundance, both to humans

and stock, and to occur in fields, thickets, and about, but not

inside, dwellings. It came out into the sunshine on the hottest

days to bite. He said these mosquitoes never voluntarily quit

biting and might travel for miles on horses and cattle. The

species was reported to be comparatively rare in Tennessee.

Larvae were collected only twice during the period from 1942

to 1945, both times in association with Aedes vexant {290). In

Illinois, the females were said to make a characteristic highpitched sound, and to attack by preference in bright sunlight

rather than shade {270).

This species was previously listed for all the Southeastern

States except Florida {193) and subsequently was collected at

Tallahassee, Fla. {230). Records of the Florida State Board of

Health show its occurrence in several other localities in the northwestern part of the State {53). Its range extends to Illinois and

Nebraska, south to Texas, and New Mexico, and into Central

and South America.

ADULT.— Proboscis and tarsi all dark. Mesonotum with broad, yellowishwhite scales intermixed with lanceolate scales. Scutellar lobes with broad,
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 143

pale scales. Wing length about 4 mm., the scales dark. Dark scales of legs

and abdomen with bluish reflections. Abdominal segments with curved apical

white patches on each side of center, usually joined in the middle on the

anterior segments to form a triangular patch.

LARVA.—Antenna somewhat shorter than the head with a small tuft of

two or three branches, some of which may be redivided. Upper and lower

head hairs long, single. Comb of only four stout scales on a weak plate,

each with a long central spine and a stout subapical pair less than half as

long. Air tube stout, inflated, about 2:1, with a minute multiple tuft beyond

the middle and with two long hairs on the lateral valves, much longer than

the apical diameter of the tube; pectén with only three or four spines,

each with a single tooth at base. Anal segment ringed by the plate, which is

pierced by tufts of the ventral brush. Gills with the dorsal pair longer than

the ventral pair, and about three times as long as the segment.

PSOROPHORA DISCOLOR (Coq.)

This species is distinguished from other Grahhamia, except

P. signipennis, by the presence of spots of white scales on the

wings. The larvae occur in grassy ditches and pools, ricefields, and

other sites similar to those in which P. confinnis is found. They

have extremely long gills and apparently can remain submerged

most of the time. They are said to come to the surface rareiy until

ready to pupate (93, 159). They lie on their backs on the bottom

of the pool or cling to grass stems or debris by means of the anal

tufts. Larval development requires 10 days or more, which is

considerably longer than the time required for larvae of P. confinnis and P. ciliata, with which it is usually associated. In Tennessee, where the larvae were found to be fairly abundant, they

were associated chiefly with Aedes vexans and P. confinnis.

The species occurs sparingly throughout the Southeast but was

found in large numbers in the rice-growing areas of Arkansas,

where it caused annoyance to man and livestock on a par with

P. confinnis (159). It constituted 22 to 46 percent of all females

caught in light traps during the summers of 1939 to 1942. It was

taken in considerable numbers, chiefly as larvae, in Mississippi

(2^5), Identification of the blood meals of 783 engorged specimens by means of the precipitin method, as reported by Whitehead (32i), showed the following percentages with different

kinds of blood: 73 bovine: 18 equine, 8 swine, 0.9 (7 specimens)

human, and 0.4 avian. , ^ .^ . c^j. ^

P. discolor has been recorded from all the Southeastern States

(193), Its range extends from New Jersey to Nebraska, and

southwards to Texas, New Mexico, and into Mexico.

ADULT—Proboscis with a broad white median ring, the palpi white-tipped.

Mesonotum with fine brownish and golden scales Wing length about 3.5 mm.,

the scales on some of the veins mixed dark and white; on others the white

scales are segregated into distinct spots, one spot at the base of the costa,

one at tip of^subcosta, one in the middle of vem. 3, one each m the middle

of vein 4 and both forks of vein 5, and one covering the basal two-thirds of

the anal vein- the wing fringe is without fringe spots. Femora tibiae, and

Ssttäsir segment predo pale-scaled speckled with dark, the

femora with narrow preapical white rings. The tibiae and first tarsalsegmSits with a black apical ring, the other tarsal segments, with broad basal

wMte rTngs covering about half of the segments on the hind feet Abdomen

with a median, apical, white triangular patch on segment 1, the other

'TrnvA^'Tntn^^^^^ -th two long bristles at outer

thfed^nTÄf^^^^ ^^ b^?J ^îi^^^l. ß^^il ''T\ ^^^

lower head hairs long and single, equal in length. Comb with about six
144 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

strong, thornlike scales set on the posterior border of a weak plate. Air tube

unusually small, about 3:1, with a very large multiple tuft near middle,

nearly as long as the entire tube; pectén of about six long spines. Anal

segment ringed by the plate, which is penetrated by a few weak tufts of the

ventral brush. Gills very long, five or six times as long as the segment, and

with prominent tracheae.

PSOROPHORA FEROX ( Humb. )

(Syn., Janthinosoma sayi D. and K., Cidex posticatus Wied.,

C. micsicus Say; the white-footed woodland mosquito)

P. forjx is one of the commonest of the blue-black Janthinosoma.

The last two segments of the hind tarsi are pure white, and the

thorax has dark and white scales intermixed. The larvae are

found chiefly in temporary pools or depressions in woodlands, and

sometimes in grassy pools or ditches at the edges of the tree

growth. In southern Mississippi the larvae occurred in great numbers in puddles left by the overflow of Leaf River (225), They

are often associated with Psorophora varipes and Aedes vexans

in shaded pools. Several generations may occur during the summer, depending on the usual sequence of drying and flooding. The

adult females become numerous and extremely annoying at times

and, with the other species frequently associated with them,

constitute a serious menace to man or animals entering their

habitats. They feed during the daytme or night, and usually remain

in or near the woodlands where they develop, but may leave the

woods on occasion and even enter houses (303).

This species was previously reported from all the Southeastern

States, and is known to occur generally through the East and

Midwest, southeastern Canada, and Mexico ; also in the Caribbean

region, and South America.

ADULT.—Proboscis and palpi all dark. Mesonotum with mixed broad white

and narrow brown scales, not forming stripes; fairly broad whitish scales

around prescutellar space and on lobes of scutellum. Wing length about

4 mm., the scales all dark. Legs blue-black, the tarsi all dark except the last

two segments of the hind feet and usually the tip of segment 3, which are

white; white knee spots present on femora; first two tarsal segments and

apical portion of tibia of hind legs with short, erect scales, but not nearly

so shaggy as in P. ciliata. Abdomen blue-black with purplish reñections and

with small apical, lateral white spots.

LARVA.—Antenna unusually long, about 1.5 times as long as the head,

with a large multiple tuft inserted near the middle. Both upper and lower

head hairs long, double, the lower sometimes triple. Comb with six or seven

scales in a curved row set on a weak plate, each scale thorn-shaped with ths

apical spine much longer than the subapical pair. Air tube about 4:1,

inflated, the tuft minute, multiple, the pectén with two to five spines, usually

three or four, each with a stout tooth on one or both sides at base and

one or two smaller spinules. Anal segment ringed by the plate, which is

pierced by tufts of the ventral brush. Gills about 1.5 times as long as

segment.

PSOROPHORA HóRRIDA (D. and K.)

This comparatively rare species is similar in appearance to

P. ferox but has a broad median stripe of dark scales on the

thorax. There is almost no information available as to its habits

in the Southeast. In Oklahoma, v^here it is not rare, it breeds in

shaded temporary rain pools. The adults attack man readily (275).

They v^ere usually taken while biting v^ith P. ferox in woods
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 145

Specimens were collected at Marianna, Fla., in 1945 and 1950

{230, 53), and this completed the record of its occurrence in the

Southeastern States. Its known range extends from Maryland to

Ohio, Missouri, and Texas.

ADULT.—Proboscis and palpi all dark, the palpi less than a third as long

as the proboscis. Mesonotum with a wide median stripe of narrow brown

scales, bordered with broad white scales laterally and around the prescutellar

space. Wing length about 4 mm., the scales all dark. Legs and abdomen

blue-black. Femoral knee spots present, the tarsi all dark except the last two

segments of the hind feet, sometimes also the tip of segment 3, which are

white. Segment 1 of abdomen with a large white patch, the other tergites

with only small lateral, apical white spots.

LARVA.—Antenna about as long as head or a little longer, with a large

multiple tuft near the middle. Upper and lower head hairs short, the upper

double, the lower double or triple. Lateral abdominal hairs minute on segments 4 to 6. Comb, a curved row of seven or eight scales, each thornshaped, with a fringe on each side of fairly strong subequal spinules. Air

tube strongly inflated, about 3.5:1, the tuft minute, multiple, placed laterally;

pectén with four or five spines, each with several subequal teeth on one

or both sides at base. Anal segment ringed by the plate, which is penetrated

by the tufts of the ventral brush. Gills bluntly pointed, about twice as long

as the segment.

PSOROPHORA HOVVARDH (Coq.)

(Howard's gallinipper)

This large species is similar in general appearance and habits

to Psorophora ciliata and the two are commonly associated, although hoivardii adults, in the writer's experience, are usually of

less frequent occurrence. The larvae are predaceous on other

mosquitoes and have been reported to devour small polliwogs also.

Breland {56) found that they disregarded larvae and adults of

Drosophila when offered to them in the laboratory. He observed

that the larvae used their modified mouth brushes to aid the

mandibles in capturing their prey. The females attack in the day

or night and are severe biters. They apparently ñy considerable

distances from their breeding places and are to be seen rather

frequently in garages and shrubbery around urban residences.

They perhaps disperse farther than P. ciliata, as they have been

observed more often than that species in residential sections of

Fort Lauderdale, Fla., in association with adults of P. confinnis

that presumably had flown in from the farmlands and glades

west of the city.

P. howardii has previously been recorded from all the Southeastern States {193), and occurs also from Maryland to Missouri

and Texas, and into Mexico.

ADULT. Nearly as large as P. ciliata, with a general bluish color. Proboscis

unringed, palpi almost half as long as the proboscis. Thorax with a broad

median stripe of dark narrow scales and nude lines, the lateral areas covered

with broad white scales and similar ones around the prescutellar space.

Wing length about 6 mm., the scales all dark. Legs much less shaggy than

P ciliata, the erect scales being limited mostly to the tips of the femora and

tibiae- segment 2 of tarsi with a narrow basal white ring and a similar ring

sometimes present on segment 1. Most of the abdominal tergites are palescaled laterally and apically. , x^ r^ -T . ,. ^

LARVA Larval characters very similar to those of P. ciliata except for

the lateral hair of the anal segment, which in the present species is rather

long and is single or forked beyond the middle.
146 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

PsOROPHORA JOHNSTONII (Grabham)

This tropical Janthinosoma resembles P. varipes, with only the

fourth hind tarsal white, but lacks the median dark stripe on the

thorax. It was first collected on the Florida Keys in October and

December 1945, when eight biting females were taken on Cudjoe

Key, and one male was obtained from a light-trap collection {282,

252). Specimens previously collected on the Keys and identified as

P. varipes {193) were probably this species. Larvae were first

collected in December 1947 in a temporary rain pool on Long Key,

and were taken during 1948 from several other Keys extending

from Big Pine to Key Largo {306). The Long Key pool, which

was said to be typical of the larval habitats, was in a shallow

depression in the limestone located at the edge of a buttonwood

transition zone between a hardwood forest and a black mangrove

swamp. It was filled with a thick mat of purslane sesuvium {Sesicvium portulacastrum) ^ but free of permanent aquatic vegetation.

Various kinds of plants and trees surrounding the pool formed

dense shade. These larvae were associated with a dozen other

species of both fresh and salt-water mosquitoes. On Big Pine Key,

one of the breeding places was a pool about 12 feet in diameter and

3 feet deep filled with clear fresh water and only partially shaded.

The bottom of the pool was covered with water hissop {Bacopa

monnieri). Another breeding site was a stagnant, sunlit pothole

containing decaying vegetation. Among the larvae associated with

johnstonii on this Key were those of Anopheles albimanits and

A. crucians. Apparently the breeding pools on all the Keys

contained only fresh rainwater.

In April 1948, after an exceptional amount of rainfall, females

of johnstonii became very numerous and annoying {306). Biting

counts of 100 per minute were obtained on two occasions, one at

11 a.m. and one at 2 p.m. They were said to bite at any time during

the day, either in shade or bright sunlight and were very aggressive. This species was previously known only from the Greater

Antilles.

ADULT.—Proboscis and palpi dark. Occiput with rather broad, recumbent

whitish scales. Mesonotum covered with fairly broad yellowish white scales

extending over prescutellar space and the scutellum. Wing length about 3.5

mm., the scales dark. Legs and abdomen blue-black, small femoral knee spots

present; tarsi all dark except segment 4 of the hind feet which is usually

white dorsally on the basal four-fifths (sometimes all dark). Segment 1 of

the abdomen with wide white band, the other tergites with large apical, lateral

white spots that extend onto the dorsum.

LARVA.—Antenna about as long as head, with a multiple tuft beyond the

middle of shaft. Upper head hair double, the lower, double or triple, both

short. Lateral abdominal hairs multiple on segments 4 to 6. Comb of about

six thorn-shaped scales, on a weakly sclerotized plate, the lateral spinules fairly

stout. Air tube stout, inflated, about 3:1, the tuft minute, multiple, inserted

beyond middle; hair on lateral valve nearly as long as the apical diameter

of tube; pectén usually with 6 to 8 spines, each with two or three teeth on

one or both sides at base. Anal segment completely ringed and pierced by

tufts of the ventral brush, the lateral hair very small, multiple, branched

from base. Gills pointed, about 1.5 times as long as the segment.

PsOROPHORA LONGIPALPIS Roth

This mosquito was described as new by Roth in 1945 {273) It

had previously been confused with Psorophora horHda, which it
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 147

closely resembles. The holotype male was from Fayetteville, Ark.,

and the paratypes from Kansas, Missouri, Oklahoma, South

Dakota, and Texas. Wirth (332) identified larvae and reared adults

of this species collected in 1943 at Rapides, near Alexandria, La.

No information was given as to their breeding or biting habits,

but these presumably are similar to those of P. hórrida.

ADULT.—Proboscis and palpi dark, the latter more than a third as long

as the proboscis. Occiput with a large median area of broadened, recumbent

pale scales, a patch of broad, appressed, violaceous scales on each side.

Mesonotum with a wide median dark stripe, bordered by wide stripes of

broad yellowish-white scales covering the lateral surfaces, and with whitish

scales around the prescutellar space and on the lobes of the scutellum. Wing

about 4 mm., the scales dark. Legs and abdomen blue-black. Femoral knee

spots absent, tibiae and tarsi all dark except the last two segments of the

hind feet which are all white. First segment of abdomen with an apical white

band, the other tergites with small lateral apical spots, scarcely visible

from above.

LARVA.—Antenna somewhat longer than the head, the tuft multiple. Upper

and lower head hairs moderately long, double. Lateral abdominal hairs

multiple on segments 4 to 6. Comb usually with seven thorn-shaped scales

in a curved row. Air tube inflated, about 3.5:1, the tuft minute inserted high

on side of tube at about the apical third; pectén of only 3 or 4 spines, each

with a stout tooth at base. Anal segment completely ringed and pierced by

tufts of the ventral brush, the lateral hair small, branched toward tip. Gills

about 1.5 times the length of the segment.

PSOROPHORA PYGMAEA (Theob.)

A small tropical Grabhamia with a wide white ring on the proboscis, but with the tarsal rings narrow and confined to the base of

the segments. It was first collected at Key West, Fla., in 1901 (A.

Busck, four specimens) and in 1903 (E. A. Schwartz, six specimens). These were described as Culex naniis by Coquillet {80),

but this name was placed as a synonym of Psorophora pygmaea

by Howard, Dyar, and Knab (163). The type locality of pijgmaea

was Antigua. Buren {61) reported that 37 specimens, all females,

were taken in a light trap at the U. S. Quarantine Station on

Fisher's Island, Miami Beach, Fla., in June and July 1946. No

breeding could be found on Fisher's Island or Miami Beach. The

author believed, therefore, that it must have occurred on Virginia

or Biscayne Keys. He also reported that the U. S. National Museum

has 30 specimens collected at Key West from June to October 1924,

a previously unpublished record. More recently pijgmaea has been

collected in light traps in St. Lucie, Martin, and Palm Beach

Counties {53). The species has been recorded from the Bahamas

and the Greater Antilles.

ADULT. Proboscis with a broad central white rmg, the palpi speckled

with white, Mesonotum with narrow golden or brownish scales. Wing length

about 3 mm., the scales broadened, speckled with white, but without white

spots Femora and tibiae sprinkled with white, the femora with narrow

subapical white rings, sometimes indistinct, tarsal segments with narrow

basal white rings. Abdominal segments with wide apical white bands produced

^ LARVA—Antenna of normal size, with a multiple tuft inserted before the

middle Upper and lower head hairs long, single, subequal. Comb, a curved

^oToi aboSt six stout scales, each rounded with a long apical spme twice

Is long as the longest side spines, the comb set on a weak plate. Air tube

3-1 somewhat inflated, with a small multiple tuft near middle; pectén of

Ät sTxSnes each with several stout teeth on one side toward the base;

ha?r onTaterÄ of tube minute, less than half as long as apical diameter

of tube. Anal sISnent with the plate pierced by tufts of the ventral brush.

Gills about one ¿nd one-half times as long as the segment, pomted.
148 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

PSOROPHORA SIGNIPENNIS (Coq. )

This species of Grahhamia resembles P. discolor in having

white spots on the wing, but it also has pale fringe spots opposite

the tips of the veins, which are not present in that species. It

occurs principally in the plains and valleys west of the Mississippi

River. It is adapted to arid conditions and development of the

larvae in transient pools and irrigated fields is very rapid. The

females attack man but, in Oklahoma, were rarely observed bitmg

in spite of an abundance of larvae and of adults in light traps

{275). P. sig7i ipermis is rare in the Southeast and has been reported only from Arkansas (f^6, 54) and Tennessee (222). Its

range extends from North Dakota to Texas and westward to

Arizona and Mexico.

ADULT.—Proboscis with a very broad median white ring. Palpi speckled

with white. Mesonotum with pale brownish scales, paler around sides and

posteriorly. Wing about 3.5 mm. in length, with mixed black and white

scales and several distinct white spots, two on the apical half of the costa,

separated by a slightly longer black spot, one at the tip of the anal vein and

a long one covering most of the basal half of this vein; pale fringe spots

present opposite the tips of all the longitudinal veins. Femora and tibiae

speckled with white, the femora with narrow, preapical, pale rings; tarsal

segments with basal white rings, and segment 1 with a median white area

covering about two-thirds of the segment on the hind foot, the ring on

segment 2 of the hind foot more than half as long as the segment. Abdomen

largely pale-scaled, speckled, and streaked with dark.

LARVA.—Antenna shorter than head, with a large multiple tuft inserted

about the middle of the shaft, the branches conspicuously feathered. Preantennal tuft with six or seven branches. Upper and lower head hairs long,

usually single, sub-equal in length, sometimes one or more, rarely all four,

branched {14?)- Comb, a curved row, usually of six scales set on a small

plate, each scale with a long central spine and a few short lateral spinules.

Air tube about 3.5:1, somewhat inflated, with a very small multiple tuft near

the apical third; hairs on the lateral valves nearly as long as the apical

diameter of the tube; pectén of about six spines. Anal segment ringed by the

plate, which is penetrated by tufts of the ventral brush. Gills all of equal

length, about twice as long as the segment.

PsOROPHORA VARIPES (Coq.)

This widely distributed Janthinosoma has a broad median dark

stripe on the thorax (fig. 19, G) and a white band on segment 4

only of the hind tarsus. The larvae occur in temporary woodland

pools and are frequently associated with P. ferox. In Louisiana,

Buren (60) found them breeding abundantly in woodland floodwaters and nearly absent in temporary rain pools. The larvae were

collected in mats of debris. This species is comparatively rare in

most of the Southeast but the writers found it to be exceedingly

annoying in the woods of northeastern Louisiana. It did not fly

out in the sunlight to bite. In Arkansas it was said to be normally

an inhabitant of the woods along large streams and rivers, and

was present in swarms as soon as the water had partly receded

(156, 157). In southern Mississippi, on one occasion in 1943 the

females occurred in almost unbearable swarms in wooded swamps

along the Leaf River (225). P. varipes has previously been reported from all the Southeastern States (193). It also occurs in

many of the States from New York to Missouri and Texas, and its

range extends into Mexico, Central, and South America.

ADULT.—Proboscis and palpi dark. Mesonotum with a very broad median

stripe of brown lanceolate scales and narrower lateral stripes of broad
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 149

yellowish-white scales, with similar broad scales around the prescutellar

space and on the lobes of the scutellum. Wing length about 3.5 mm., the

scales all dark. Legs and abdomen blue-black with violet reflections. Knee

spots absent. Tibiae and tarsi all dark except segment 4 of hind foot which is

pale-scaled dorsally. Segment 1 of abdomen with an apical triangular white

patch, the other tergites with lateral, apical white spots, visible from above.

LARVA.—Antenna slightly longer than the head, with a multiple tuft near

or beyond the middle. Upper and lower head hairs both long, two- to fourbranched, the upper usually double, the lower double or triple. Lateral

abdominal hairs four- or five-branched on segment 3, single or double on

segments 4 to 6. Comb with six or seven scales in a strongly curved row;

the single scale thornlike with fairly strong lateral spinules. Air tube inflated, about 3.5:1, a small multiple tuft beyond middle of tube; pectén of

three or four spines. Anal segment with the plate pierced by tufts of the

ventral brush. Gills about 1.5 times as long as the segment.

Genus TOXORHYNCHITES Theobald

(Syn., MegarJmuis Rob.-Desv.)

The subfamily Toxorhynchitinae has a single genus, and is represented in the United States by a single species with two subspecies, both of which are found in the Southeast. These were

previously listed as full species but Jenkins (173), after a detailed

study, considered them to be only of sub-specific rank. The genus

has long been known as Megarhinns but this name was found to

be preoccupied and could no longer be used under the rules of

priority.

These are very large, brilliantly colored, non-bloodsucking mosquitoes. The long, tapered proboscis, the basal half of which is

rigid, has the apical half bent downward at nearly a right angle

(fig. 2, A), and not fitted for puncturing. The female palpi are

about two-thirds as long as the proboscis, the clypeus is longer

than broad, the male antenna plumose, the scutellum evenly

rounded as in Anopheles. The second marginal cell of the wing is

much shorter than its petiole. The wing spread is about 15 mm.

The two subspecies are indistinguishable except for the slight

differences in the white markings of the male tarsi.

Breeding occurs chiefly in tree holes, and occasionally in rock

holes and artificial water containers. The larva is very large, a

strong swimmer, and is predaceous on other mosquitoes, or sometimes cannibalistic. The head is nearly square and the mouth

brushes each consist of about 10 strong, curved rods. These were

previously thought to be adapted for grasping their prey but

Breland (56, 57), in studies of Toxorhynchites rutihis septentrionalis in Central Texas, noted that only the mandibles were

used for capturing the prey, and he concluded that the mouth

brushes were not employed for this purpose. The eighth segment

of the larva has a sclerotized plate bearing two stout barbed hairs

and two minute branched hairs on the posterior margm. The

usual comb and pectén are lacking.

KEY TO SUBSPECIES

MALES

1. Segment 2 and part of segment 3 of front tarsus, and usually

segment 5 of hind tarsus white. Florida, Georgia, and South

p ^ ,. rut dus rutilus

Front^tarsus and usually the fifth hind tarsal entirely dark. Occurs

sparingly in most of the South ruHlus septentrionahs
150 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

TOXORHYNCHITES RUTILUS RUTILUS (Coq.)

A very large irridescent mosquito, the larvae of v^hich are

found in tree holes and artificial v^ater containers. They were

collected in Florida in ^^surprising numbers'' in the native bromeliad Tillandsia utricidata L., with larvae of Wyeomyia vanduzeei

and mitchellii (28i). The authors found none of the larvae m innumerable tree holes examined in the same areas. However, other

workers have collected them in a wide variety of places in various

parts of Florida, including holes in water oak, live oak, scrub oak,

orange, pecan, and pine trees, and from glass jars, tin cans,

wooden barrels, an auto tire, steel drum, a rowboat, and others

(19), These authors reported that one larva collected in the first

instar devoured 118 Orthopodomyia signifera and Aedes triseriattis larvae, pupated in 16 days, and emerged in 4 more days.

Another larva collected in the third instar devoured 112 larvae,

pupated in 22 days, and emerged in 5. The larval period has been

observed by others to last as long as six months. The larvae are

fairly active but often hang quietly below the surface of the water

at various angles or rest on the bottom or sides of the container.

They attack each other if other food is not available. The adults

are mainly crepuscular and were observed flying in the woods

from about 5 p.m. until dark (17i),A female was observed dipping

up and down over water in a tree hole as if ovipositing, but no

eggs were found. It is presumed that the adults feed on flower

nectar as do other members of the genus.

This subspecies was previously listed from Florida and Georgia

(193). Since then it has been recorded from numerous additional

places in Florida (28JÍ. in, 19, 173), from Thomas County,

Georgia (17Jf) and Myrtle Beach, S. C. (67). Of 10 males reared

in the last locality, 3 were rutilis and the others septentrionalis.

There are a few records from other localities based on females

only, but these cannot be relied upon in areas where the two

subspecies may overlap.

ADULT.—Proboscis and palpi as described under the genus. Occiput covered

with broad flat irridescent bluish scales. Mesonotum with a narrow median

stripe, tapered posteriorly, and lateral borders of white or golden-yellowish

scales, the rest of the scutum with fine dark scales. Wing about 6 mm., the

scales sparse and fairly broad; second marginal cell less than half as long

as its petiole. Front and mid tarsi of female with the second, third, and

most of the fourth segments white, the hind tarsi with the fourth and most

of the fifth segments white. Male with the second and basal two-thirds of

the third segments of the fore and mid tarsi white dorsally, the hind tarsi

usually with the fifth segment all white. Abdomen dark blue above, yellowish

below.

LARVA.—Antenna very small, with a pair of single hairs and a minute,

multiple tuft toward the apex. Mouth brushes each consisting of about 10

strong, curved rods apparently adapted for grasping prey, but as previously

mentioned in the discussion of the genus, Breland (57) has cast doubt on

this assumption. Upper and lower head hairs and preantennal hair each

single, small. Eighth abdominal segment with a large sclerotized plate bearing

two stout spinulose hairs and two minute tufts on its posterior border. Air

tube small, about 2:1, pectén lacking, a large hair tuft near the basal

fourth. Anal segment ringed by plate with a stout spinulose lateral hair,

Gills very short, bulbous.

TOXORHYNCHITES RUTILUS SEPTENTRIONALIS (D. and K.)

This subspecies breeds in situations very similar to those of the
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 151

preceding subspecies except that it has not been taken from

bromeliads ; in fact, has not as yet been reported as far south as

the northern limits of these native air plants. Breland {56, 57),

who has made a study of this mosquito in Texas, reports that the

eggs are oval in shape, pure white in color, and studded with small

tubercles. Females were observed dipping over the water surface

and it was thought that this was the method of oviposition. Larvae

fed on other mosquito larvae, reached the 4th instar rapidly, but

remained in this stage without pupation for several months unless

adult fruit flies (Drosophila) were added to the diet. On a pure

diet of fruit flies, or when these were added during the late instars,

the cycle from egg to adult was completed in 25 to 35 days. Other

workers, however, have reared larvae to the adult stage without

supplying them with adult flies. In watching the feeding of the

larvae under a binocular microscope, Breland found that they

seized their prey with the mandibles only, and that the modified

mouth brushes were not used at all for this purpose. Apparently

no actual use of the mouth brushes for grasping has been observed

by others.

The larvae are usually associated with Orthopodomyia signífera

and alba, Aedes triseriafus, and Anopheles barberi, which presumably serve as food. The species is said to pass the winter in

the larval stage, but Breland thought that in central Texas they

probably overwinter as adults.

The adults have been observed visiting flowers, and it is thought

that flower nectar is their principal source of nourishment. They

usually remain in the woods and are active during the day and

early evening. In Arkansas, the males were observed to congregate

regularly at a particular tree or bush (303). The resting places

of females were not observed.

The range of this mosquito extends from New Jersey to Florida

and westward to eastern Texas, Oklahoma, and Kansas. It was

previously listed from all the Southeastern States except Florida

(190). Since then, records have been added from Jacksonville

(232, 19), and Tallahassee, Fla. (19). It therefore overlaps the

range of r. riiWis in northern Florida, southern and eastern

Georgia, and eastern South Carolina.

ADULT.—This subspecies appears to be indistinguishable from the previous

one in all characteristics except the white scaling on the male tarsi, as given

in the Kev. In typical male specimens, the front tarsus is all dark, the mid

tarsus with segment 2 white on one side or dark, the hind tarsus with

segment 4 white, and segment 5 usually dark. Various intergrades in coloration occur in the region where the two overlap (173), and with atypical

specimens, identification may be difficult or impossible.

LARVA.—Description the same as for subspecies rutilus.

Genus URANOTAENIA Lynch-Arribalzaga

The species of this genus are small and sometimes brilliantly

colored. They are recognized by the very short forks of wmg vem 2,

and short palpi in both sexes. The male antennae are plumose.

One süiracular bristle is present, post-spiracular bristles are lacking, the T30stnotum and the squamae are bare. The anal vem is

short, ending before the level of the fork of vein 5. In the larva

the head is longer than broad and dark colored. The upper and

lower head hairs in our local species are single, stout, and spme-
152 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

like (fig. 16, A). The eighth abdominal segment has a lateral plate,

the comb scales forming a row on the posterior border. The air

tube has a pectén and a single pair of ventral brushes. The anal

segment is ringed by the plate.

KEY TO SPECIES

ADULTS

1. Mesonotum with a narrow median longitudinal line of bright

bluish scales (ñg. 19, J) ; tarsi all dark. A common species . . sapphirma

Mesonotum dark-scaled above, with a short line of purplish scales

on lateral margin; apex of third and entire fourth and fifth

hind tarsal segments white. Common in Florida and southern

Louisiana lowii

LARVAE (FOURTH INSTAR)

1. Upper lateral hair of abdominal segments 1 and 2 triple sapphhina

Upper lateral hair of segments 1 and 2 double (lower hair single

in both species) lowii

URAN0TAENL4 LOW^H Theob.

(Syn., U. continentalis D. and K.)

The larvae occur in ground pools, especially in the grassy margins of ponds and lakes. Their breeding habits are similar to those

of U, sapphirina. In New Orleans, La., the larvae were found in

a pool densely filled with Myiiophyllum verticulatum (15i). The

adults were not observed except in the vegetation of the pool or

resting on the water. The females could not be induced to bite.

From a study of the feeding habits of this mosquito in Louisiana,

Remington (263) reported that they fed readily in cages on

several kinds of amphibians including frogs (Rana and Hyla),

toads (Bufo) y and a salamander (Desmognathus). They were

not interested in a tortoise (Terrapene), two kinds of lizards

(Leiolopisma and Anolis)^ earthworms, snails, land slugs, or man.

Females were observed biting on frogs in the field. The adults

were commonly collected in light traps at Camp Shelby, Miss.

(225), and in Florida (50),

The species was previously reported from all the Southeastern

States except Tennessee and North Carolina (193), and a record

from^ North Carolina has since been added (228). In southern

Florida it is the more prevalent of the two species of Uranotaenia

and was said to be common on the Keys (252), It occurs also in

Texas, and its range includes Mexico, the Caribbean region, and

South America.

ADULT.—Proboscis and palpi dark. Occiput covered with broad appressed

scales, the median area dark, but with purplish scales laterally and around

eye margin. Mesonotum all dark-scaled above, with a short line of purple

scales on the lateral margin. There are patches of pale purplish scales on

the sides of the thorax and on the pronotal lobes. Wing length about 2.5 mm.

or less, scales rather sparse and broadened, a short line of purplish scales

at the bases of veins 1 and 5. Legs with small white spots at the tips of the

femora and tibiae, the hind tarsi with the fourth and fifth segments and apex

of third white, the fore and mid tarsi all dark. Abdomen dark above, some

of the segments with lateral apical spots of purplish scales.

LARVA.—Antenna very short, with a small single hair at about the basal

third. Upper and lower head hairs stout, single, spinulous. Upper lateral

hair of abdominal segments 1 and 2 double, the lower single. Segment 8
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 153

with a large sclerotized plate and about eight comb scales on the posterior

border, each scale long, pointed, and finely fringed on each side nearly to

tip. Air tube about 4:1, with one pair of large multiple tufts attached near

the middle, the pectén teeth rounded, and finely fringed from base to apex.

Anal segment longer than wide, ringed by plate. Gills shorter than segment,

bluntly rounded.

URANOTAENIA SAPPHIRINA (O.-S.)

(Syn., U. socialis Theob.)

A small species, slightly larger than U. loioii, with a median line

of brilliant blue scales on the thorax. It breeds in permanent grassy

pools, swamps, and vegetation at the margin of lakes. It is often

found in growths of Levina, and is a frequent associate of Anopheles

quadrimacidatus and Cidex erraticits. In Tennessee, it was found

more often with A}wpheles punctipennis and Ciilex apicalis (=

C. territans) (290). Thibault (303) said it was one of the most

abundant mosquitoes in Lonoke and Pulaski Counties, Arkansas,

though the adults were not generally in evidence. They have, however, been observed in considerable numbers during the winter

months in caves and hollow trees, and on one occasion in enormous

numbers in December in old Fort Jackson below New Orleans

(154). Since few larvae could be found nearby, it appeared that

they must have come from some distance away. That they are

capable of flying considerable distances is shown by the capture

of adults in light traps in Delaware Bay where the nearest shore

was about 8 miles away (209). In southern Mississippi they were

reported to pass the winter in large hollow trees (225). There are

apparently no definite records of their biting persons, and their

feeding habits are unknown. It is doubtful that the females attack

any warm-blooded animals.

The species is a common one throughout the Eastern and Central

States, and has been recorded from all the Southeastern States

(193). It occurs also in southeastern Canada, Mexico, and parts

of the West Indies.

ADULT.—Proboscis and palpi dark. Occiput with a border of broad purplish

scales around the eyes. Mesonotum with a narrow median longitudinal line

of broad, brilliant blue scales extending on to the mid loba of the scutellum,

and lateral lines of similar scales in front of the wing base, the rest of the

scutum with fine dark scales; patches of bluish scales also present on the

pleurae and pronotal lobes. Wing length about 2.5 mm. or more, with a hne

of purplish scales on the basal part of vein 5 and a few similar scales at the

base of vein 1. First marginal cell less than half as long as its petiole.

Tarsi all dark, the tips of the femora and tibiae with small white spots.

Third, fifth, and sometimes the sixth abdominal tergites with a narrow

apical white band widened in the middle, or a rounded median patch.

LARVA. Very similar to the preceeding species. Antenna with a small

single hair at about the basal third. Upper and lower head hairs very stout,

tapered, spinelike. Upper lateral hair of abdominal segm-nts 1 and 2 triple

(instead of double as in U. lowii), the lower hair single. Comb of about

eight scales set on the posterior edge of a large sclerotized plate, each scale

pointed and finely fringed on about the basal half. Air tube about 4:1, with

a pair of multiple tufts near the middle, the pectén teeth rounded and nearly

completely fringed. Gills slightly shorter than the anal segment, pointed.

Genus WYEOMYIA Theobald

The species of this genus are very small, and are distinguished

from other genera by the presence of a tuft of setae on the post-
154 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

notum. The most obvious recognition characters are the vestiture

of broad dark scales on the thorax and the markings of the abdomen, which is usually somewhat compressed laterally, and on

which the dark scaling of the dorsum and the white scaling of the

venter meet to form a straight line along the sides. The male palpi

are short ; the antennae similar to those of the female. The mesonotum lacks the dorsocentral bristles and is covered with broad,

appressed scales instead of the narrow, semi-erect scales of Culex,

Spiracular and upper mesepimeral bristles are present, the postspiracular and lower mesepimeral bristles lacking. The first submarginal wing cell is as long as its petiole. The tarsi of all the

local species have some white markings, but these may be indistinct

and easily overlooked in dead specimens, or the extent of the pale

scaling may appear different in different positions. The claws of

both sexes are simple. The tip of the female abdomen is blunt, and

segment 8 is not retractile. In the male the apical half of the

dististyle is greatly enlarged and divided lengthwise into several

lobes or arms.

In the larva the head is wider than long, the comb scales are in

a single row, the air tube is without a pectén, and has scattered

single or double hairs. The anal segment is without a ventral brush,

but has a pair of ventrolateral tufts (fig. 15, C) ; the plate covers

only about half the segment.

KEY TO SPECIES

ADULTS

1. Scales of anterior pronota 1 lobe silvery white vanduz^^ei

Scales of anterior pronotal lobe darker, violaceous, sometimes

whitish at sides (female of all species difficult to distinguish) . . 2

2. Scutellar scales all of same coppery color mitchellii

Scutellum with a small patch of pale scales on mid lobe haynei

MALE TERMINALIA

1. Stem of dististyle slender, longer than the modified head

section mitchellii

Stem of dististyle stout, shorter than the head section . . . ....... 2

2. Head section of dististyle divided into two lobes and a curved arm,

the middle lobe without a group of setae \aynei

Head section of dististyle divided into three lobes and a straight

slender arm, the mid lobe with a group of about 10 long setae

about the middle of the outer border vanduzeei

LARVAE (FOURTH INSTAR )

1. Upper head hairs multiple, the lower double; ventrolateral tufts of

anal segment of about 12 subequal hairs mitchellii

Upper and lower head hairs single; ventrolateral tufts of' anal

segment with not over six hairs 2

2. Ventrolateral tufts of anal segment with two'long hkirs-* Wills'with

one pair much shorter than the other . . havnei

Ventrolateral tufts with one or two long hairs and three'or' four

much shorter ones; gills very long and equal in length .... vanduzeei

WYEOMYIA HAYNEI Dodge

(Syn., W. smithii (southern records, not Coquillett) )

Records of the northern pitcherplant mosquito Wyeomyia smithii
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 155

from North and South Carolina and Alabama are probably of

this species. W, haynei was described by Dodge (89) as a distinct

species, the larvae of which were collected from the southern subspecies of the pitcherplant Sarracenia ¡mrpurea venosa Raf. The

two species are very similar in nearly all characters. The author

identified specimens from Theodore, Ala., and several localities in

North and South Carolina. The type locality for haynei is near

Columbia, S. C. The species has also been reported from a few

other localities in South Carolina {321), The collection in the U. S.

National Museum includes specimens from four localities in North

Carolina. It is recorded also from Escambia County, Florida {53).

ADULT.—Proboscis long, all dark; palpi very short in both sexes. Occiput

with broad flat scales, mostly dark with metallic reflections, a small patch of

white at vertex. Mesonotum with broad, dark, appressed scales. Scutellum with

a patch of silvery scales on the mid lobe, the rest dark. Anterior pronotal lobe

large, covered with broad, bluish or violaceous scales, the posterior pronotum

and pleurae with patches of silvery-white scales. Wing length about 2.5 mm.,

the scales narrow and dark. Fore tarsus dark, mid tarsus described as having

segments 3 and 4, and the tip of 2, white on one side (dorsally), the hind

tarsus pale-striped underneath throughout, or the apical segments darker

(these markings found to be indistinct or indefinite on mounted specimens) ;

femoral knee spots absent. Abdomen laterally compressed, the tergites all

dark, the sternites all white. The apical half of the male dististyle has an

elongate mid lobe, a slender, curved, spinose arm arising from its base, and a

quadrate lateral lobe; the basal half stout, about as long as the apical mid

lobe.

LARVA.—Antenna short, with a single hair inserted at about the distal third.

Upper and lower head hairs and preantennal hair each single, about as

long as antenna. Lateral abdominal hair usually double or triple on segments

3 to 6. Comb scales in a single row, each rounded and finely fringed to tip.

Air tube about 4:1, with numerous long, usually single, scattered hairs, the

pectén lacking. Anal segment ringed about halfway with the plate, the

lateral and dorsal hairs very long, double; the ventrolateral hair much

shorter, also double. Gills with one pair longer than the segment, swollen,

and rounded; the dorsal pair much shorter, bulbous.

WYEOMYIA MITCHELLH (Theob.)

A small forest mosquito with white bands on some of the tarsal

segments. The larvae are found only in water that collects at the

base of the leaves of epiphytic Bromeliaceae (pi. 10). They occur

throughout the year except when the breeding places become dry.

The females bite readily during the daytime and sometimes cause

considerable annoyance in shady, humid woodlands or yards with

trees in which the bromeliads grow. They do not fly far from their

breeding sites. On one occasion the writers observed an annoying

infestation in a greenhouse where the larvae were found in a large

collection of exotic bromeliads. The white markings on the feet are

more noticeable when the insects are alive, either flying or at rest,

when the hind legs are turned up over the back with the tarsi

pointing forward. The species is found in the United States only

in the southern half of Florida. Specimens have been taken as far

north as Orange and Volusia Counties (latitude about 29°). They

occur in the West Indies. ^ . . .., ,.

ADULT.—Proboscis long, dark, the palpi very short. Occiput with a line of

white scales around eye margin and a white patch at vertex. Mesonotum and

scutellum with broad, appressed dark scales. Anterior pronotal lobes covered

with dark bluish or violaceous scales, sometimes with paler scales around

the edffes; posterior pronotal lobes and pleurae with patches of silvery white

scales Wing length about 2.5 mm., the plume scales narrow and dark.

Femoral knee spots absent. Fore tarsi all dark; mid tarsi white streaked on
156 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

one side on segments 3 and 4, the tip of 2 and sometimes 5; hind tarsi with

white basal spots or lines underneath on segments 1 to 3, sometimes also

on 4 and 5. Female abdomen somewhat compressed laterally, the tergites

all dark, and the sternites all white, as in other members of the genus. Head

of male dististyle greatly enlarged with a broad quadrate mid lobe having a

thumblike projection on one side, an ovate lobe from the base on one side,

and a straight slender arm on the other; stem of dististyle slender, longer

than the head.

LARVA.—Antennal hair small, three-branched, inserted at about the distal

third of the shaft. Lower head hair long, double, the upper and preantennal

hairs three- or four-branched. Lateral abdominal hairs usually double on

segments 3 to 6. Comb of numerous scales in a long row, each pointed, and

finely fringed nearly to the tip. Air tube about 4:1, with scattered long, single

hairs and a few shorter double hairs toward the tip; the pectén lacking.

Plate of anal segment covers about three-fourths of the surface; the lateral

and dorsal hairs each very long, single; the ventrolateral hairs shorter than

the segment, multiple, subequal in length. Gills somewhat longer than the

segment, all of equal length and bluntly pointed.

WYEOMYIA VANDUZEEI D. and K.

The habits and distribution of this species are almost identical

with those of Wyeomyia mitchellii. In Florida the larvae of the

two species are usually found associated in the bromeliads (pi. 10)

and the adults fly together in the woods.

ADULT.—Very similar to W. mitchellii except for slight differences in the

color of the scales on the anterior pronotal lobe, which are silvery white in

vanduzeei but darker, with purplish reflections, in rnitchellii. The midlobe of

the scutellum in both species lacks the spot of white scales found in W. hciynei.

The head section of the male dististyle is divided into three elongate lobes

and a long slender arm, the mid lobe with about 10 long setae in a group on

one side near the middle.

LARVA.—Differs from W. mitchellii in a number of respects, as follows:

antennal, preantennal, and upper and lower head hairs each single; lateral

abdominal hairs single on segments 3 to 6; comb scales rounded and finely

fringed to tip; air tube 5:1 or 6:1, with a row of about six small, single or

double hairs dorsally, one pair of double hairs laterally toward the base, and

several small single ventrolateral hairs toward the tip. Lateral hair of anal

segment double, about three-fourths as long as the dorsal hairs, the ventrolateral tuft with four to six hairs, three or four of which are short and one

or two much longer. Gills about four times as long as the segment, broad,

bluntly rounded.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 157

LITERATURE CITED

(1) ABDEL-MALEK, ALBERT

1948. THE BIOLOGY OF AEDES TRIVITTATUS. Jour. Econ. Ent. 41:

951-954.

(2) AMERICAN MOSQUITO CONTROL ASSOCIATION.

1948. THE USE OF AIRCRAFT IN THE CONTROL OF MOSQUITOES. Amer.

Mosquito Control Assoc. Bui. No. 1, 46 pp., illus.

(3) AMERICAN MOSQUITO CONTROL ASSOCIATION.

1952. GROUND EQUIPMENT AND INSECTICIDES FOR MOSQUITO CONTROL.

Amer. Mosquito Control Assoc. Bui. No. 2, 116 pp., illus.

(4) ATCHLEY, FLOYD O.

1952. DENSITIES OF ANOPHELES, SPOROZOITE RATES IN ANOPHELES,

AND INCIDENCE OF MALAI^IA IN AN AREA OF SOUTH CAROLINA.

Amer. Jour. Trop. Med. and Hyg. 1: 743-746.

(5) BAKER, F. C.

1936. A NEW SPECIES OF ORTHOPODOMYIA, 0. ALBA, sp. N. Proc.

Ent. Soc. Wash. 38: 1-7, illus.

(6) BALFOUR, MARSHAL C.

1928. STUDIES ON THE BIONOMICS OF NORTH AMERICAN ANOPHELINES.

WINTER ACTIVITIES OF ANOPHELINES IN COASTAL NORTH

CAROLINA 36° N. LAT. Amer. Jour. Hyg. 8: 68-76, illus.

(7) BANG, F. B., QUINBY, G. E., and SIMPSON, T. W.

1940. ANOPHELES WALKERI (THEOBALD) : A WILD-CAUGHT SPECIMEN

HARBORING MALARIA PLASMODIA. U. S. Pub. Health SerV.

Rpts. 55: 119-120, illus.

(8) QUINBY, G. E., and SIMPSON, T. W.

1943. STUDIES ON ANOPHELES WALKERI THEOBALD CONDUCTED AT

REELFOOT LAKE, TENNESSEE, 1935-1941. Amer. Jour. Trop.

Med. 23: 247-273.

(9) BARBER, M. A.

1927. THE FOOD OF ANOPHELINE LARVAE—FOOD ORGANISMS IN PURE

CULTURE. U. S. Pub. Health Serv. Rpts. 42: 1494-1510.

illus.

(10)

1928. THE FOOD OF CULICINE LARVAE. U. S. Pub. Health Serv. Rpts.

43: 11-17.

(11) and HAYNE, T. B,

1921. ARSENIC AS A LARVICIDE FOR ANOPHELINE LARVAE. U. S. Public Health Serv. Rpts. 36: 3027-3034.

(12) and HAYNE, T. B.

1924. SOME OBSERVATIONS ON THE DISPERSAL OF ADULT ANOPHELES.

U. S. Pub. Health Serv. Rpts. 39: 195-203.

(13) and KOMP, W. H. W.

1929 BREEDING PLACES OF ANOPHELES IN THE YAZOO-MISSISSIPPI

DELTA. U. S. Pub. Health Serv. Rpts. 44: 2457-2462.

(14) KOMP, W. H. W., and HAYNE, T. B.

1924. SOME OBSERVATIONS ON THE WINTER ACTIVITIES OF ANOPHELES

IN SOUTHERN UNITED STATES. U. S. Pub. Health Serv.

Rpts. 39: 231-246.

(15) KOMP, W. H. W., and HAYNE, T. B.

1927. THE SUSCEPTIBILITY TO MALARIA PARASITES AND THE RELATION

TO THE TRANSMISSION OF MALARIA OF THE SPECIES OF ANOPHELES COMMON IN SOUTHERN UNITED STATES. U. S. Pub.

Health Serv. Rpts. 42: 2487-2502.

1.957 THE DISTRIBUTION OF CULEX P. PIPIENS AND C. P. QUINQUEFASCIATUS IN NORTH AMERICA. Amer. Jour. Trop. Med. & Hyg.

6: 153-165.

(17) BARR, A. R., and KARTMAN, L.

1951. BIOMETRICAL NOTES ON THE HYBRIDIZATION OF CULEX PIPIENS L.

AND C. QUINQUEFASCIATUS SAY. Jour. ParaSltol. oil 419-

420.

(18) BA|HAM,^ERN^ESTm^^^^^^ MULRENNANI, A NEW SPECIES FROM

FLORIDA (DÍPTERA, CULICIDAE) . Ent. Soc. Amer. Ann. 41:

1-7.
158 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(19) MuLRENNAN, J. A., and OBERMULLER, A. J.

1947. THE BIOLOGY AND DISTRIBUTION OF MEGARHINUS ROBINEAUDESVOIDY IN FLORIDA. Mosquito News 7: 64-66.

(20) BATES, MARSTON

1949. THE NATURAL HISTORY OF MOSQUITOES. 379 pp. New York.

(21) BEADLE, LESLIE D.

1952. EASTERN EQUINE ENCEPHALITIS IN THE UNITED STATES. Mosquito News 12: 102-107.

(22) BELLAMY, R. EDWARD

1939. AN ANOPHELINE FROM INLAND GEORGIA RESEMBLING THE

BRACKISH-WATER RACE OF ANOPHELES CRUCIANS. Jour. Parasitol. 25: 186.

(23)

1956. AN INVESTIGATION OF THE TAXONOMIC STATUS OF ANOPHELES

PERPLEXANS LUDLOW, 1907. Ent. Soc. Amer. Ann. 49:

515-529, illus.

(24) BENNETT, BYRON L., BAKER, FRED C, and SELLAROS, ANDREW WATSON.

1939. SUSCEPTIBILITY OF THE MOSQUITO AEDES TRISERIATUS TO THE

VIRUS OF YELLOW FEVER UNDER EXPERIMENTAL CONDITIONS.

Ann. Trop. Med. and Parasitol. 33: 101-105.

(25) BERTHOLF, J. H.

1951. REPORT OF THE BROWARD COUNTY, FLORIDA, ANTI-MOSQUITO

DISTRICT 1950. Fla. Anti-Mosquito Assoc. Rpt. 22nd Ann.

Meeting, section 5, not paged.

(26) BiCK, GEORGE H.

1946. COLLECTIONS OF MOSQUITOES ON PARRIS ISLAND DURING 1945.

Jour. Econ. Ent. (Sei. Note) 39: 89-91.

(27) RiDLINGMAYER, W. L.

1950. SOME OBSERVATIONS OF THE BIOLOGY OF MANSONIA MOSQUITOES

IN THE LEESBURG AREA. Fla. Anti-Mosquito Assoc. Rpt.

21st Ann. Meeting, pp. 64-68.

(28)

1954. DESCRIPTION OF A TRAP FOR MANSONIA LARVAE. Mosquito

News 14: 55-58, illus.

(29) BISHOP, E. L.

1936. MALARIA-CONTROL ACTIVITIES OF THE TENNESSEE VALLEY

,^^, ^ AUTHORITY, U. S. Pub. Health Serv. Rpts. 51: 970-975.

(30) BiSHOPP, F. C.

1933. MOSQUITOES KILL LIVE STOCK. Science 77: 115-116

(31) BOHART, R. M.

1948. THE SUBGENUS NEOCULEX IN AMERICA NORTH OF MEXICO

,^^, ^ ,, (DÍPTERA, CULICIDAE). Ent. Soc. Amer. Ann. 41: 330-345.

(32) BoYD, MARK F.

1927. STUDIES ON THE BIONOMICS OF NORTH AMERICAN ANOPHELINES.

I. THE NUMBER OF ANNUAL GENERATIONS OF A. QUADRIMACULATUS. Amer Jour. Hyg. 7: 264-275, illus (33)

(34)

(35)

1929. STUDIES ON THE BIONOMICS OF NORTH AMERICAN ANOPHELINES.

PHYSICAL AND CHEMICAL FACTORS IN THEIR RELATION TO

THE DISTRIBUTION OF LARVAE IN NORTHEASTERN NORTH

CAROLINA. Amer. Jour. Hyg. 9: 346-370, illus.

1930. AN INTRODUCTION TO MALARiOLOGY. 437 pp., illus Cambridge, Mass.

1930. STUDIES ON THE BIONOMICS OF NORTH AMERICAN ANOPHELINES.

Vo .^P^F. OBSERVATIONS ON IMAGINES. Amer. Jour. Hyg.

12: 449-466, illus.

(ob) editor.

1949. MALARIOLOGY. A COMPREHENSIVE SURVEY OF ALL ASPECTS OF

THIS GROUP OF DISEASES FROM A GLOBAL STANDPOINT. Vols.

(37) _- andVoÄE^LEN. ""'^ ^""" ^"^^^-^ ^^^ ^^^^^-•

1928. STUDIES ON THE BIONOMICS OF AMERICAN ANOPHELINES. THE

ALIMENTATION OF ANOPHELINE LARVAE AND ITS RELATION TO

THEIR DISTRIBUTION IN NATURE. Jour. Prev. Med. 2: 219-
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 159

(38) KITCHEN, S. F., and MULRENNAN, J. A.

1936. ON THE RELATIVE SUSCEPTIBILITY OF THE INLAND AND COASTAL

VARIETIES OF A. CRUCIANS WIED. TO P. FALCIPARUM W^ELCH.

Amer. Jour. Trop. Med. 16: 159-161.

(39) and PONTON, GERALD.

1933. THE RECENT DISTRIBUTION OF MALARIA IN THE SOUTHEASTERN

UNITED STATES. Amer. Jour. Trop. Med. 13: 143-166, illus.

(40) and WEATHERSBEE, ALBERT A.

1929. STUDIES ON THE BIONOMICS OF NORTH AMERICAN ANOPHELINES.

V. W^INTER ACTIVITIES OF ANOPHELINE IMAGINES IN COASTAL

NORTH CAROLINA (36° N. LAT.). Amer. Jour. Hyg. 9:

682-694, illus.

(41) BRADLEY, G. H.

1924. THE NATURAL BREEDING PLACES OF ANOPHELES MOSQUITOES IN

THE VICINITY OF MOUND, LOUISLA.NA. Amer. Jour. Trop.

Med. 4: 199-233, illus.

(42)

(43)

(44)

(45)

(46)

1925. THE LARVA OF AEDES THIBAULTI DYAR AND KNAB (DÍPTERA,

CULiciDAE). Insecutor Inscitiae Menstruus 13: 89-91.

1926. OBSERVATIONS ON THE EMERGENCE OF ANOPHELES MOSQUITOES.

Amer. Jour. Trop. Med. 6: 283-297, illus.

1932. ON THE IDENTIFICATION OF ANOPHELES MOSQUITO LARVAE IN

FLORIDA. Ent. Soc. Wash. Proc. 34: 41-43.

1932. SOME FACTORS ASSOCIATED WITH THE BREEDING OF ANOPHELES

MOSQUITOES. Jour. Agr. Res. 44: 381-399, illus.

1936. ON THE OCCURRENCE OF ANOPHELES V^ALKERI THEOBALD IN

FLORIDA (DíPTERA, CULICIDAE). South. Med. Jour. 29: 857-

859, illus.

(47) FRITZ, ROY F., and PERRY, L. E.

1944. ADDITIONAL MOSQUITO RECORDS FOR THE SOUTHEASTERN STATES.

Jour. Econ. Ent. (Sei. Note) 37: 109.

(48) GOODWIN, M. H. JR., and STONE, ALAN.

1949. ENTOMOLOGIC TECHNICS AS APPLIED TO ANOPHELINES. In Boyd,

Malariology, L, pp. 331-378.

(49) and KING, W. V.

1941. BIONOMICS AND ECOLOGY OF NEARCTIC ANOPHELES. In Human

Malaria. Amer. Assoc. Adv. Sei. Pub. 15, pp. 79-87.

(50) and MCNEEL, T. E.

1935 ¡MOSQUITO COLLECTIONS IN FLORIDA WITH THE NEW^ JERSEY

LIGHT TRAP. Jour. Econ. Ent. 28: 780-786.

(51) and TRAVIS, BERNARD V.

1943. TIME-SAVING METHODS FOR HANDLING MOSQUITO LIGHT-TRAP

COLLECTIONS. Jour. Econ. Ent. 36: 51-53.

(52) BRANCH, NINA

1955 NOTES ON THE IMPORTANCE OF CONTINUOUS ENTOMOLOGICAL

RECORDS. Fla. Anti-Mosquito Assoc. Rpt. 26th Ann. Meeting, pp. 106-111.

(53) LOGAN, LUCILLE, BECK, ELIZABETH, and MULRENNAN, J. A.

1958 NEW^ DISTRIBUTIONAL RECORDS FOR FLORIDA MOSQUITOES. Fla.

Ent. 41:155-163.

(54) BRANDENBURG, J. F., and MURRILL, R. D.

1947 OCCURRENCE AND DISTRIBUTION OF MOSQUITOES IN ARKANSAS.

Ark. Health Bui. 4: 4-6.

(^^^ m7^^VARIATI0NS IN THE LARVAE OF THE MOSQUITO ORTHOPODOMYIA

ALBA BAKER (DÍPTERA, CULICIDAE). Brooklyn Ent. Soc. Bui.

42: 81-86.

^^^^ 1948 NOTES ON SOME CARNIVOROUS MOSQUITO LARVAE. Ent. Nev^S

59: 156-157.

^^'^^ 1949 THE BIOLOGY AND THE IMMATURE STAGES OF THE MOSQUITO,

MEGARHINUS SEPTENTRIONAL:S DYAR AND KNAB. Ent. SOC.

Amer. Ann. 42: 38-47, illus.
160 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(58) BULL. CARROLL G., and REYNOLDS, BRUCE D.

1924 PREFERENTIAL FEEDING EXPERIMENTS WITH ANOPHELINE MOSQUITOES. II. Amer. Jour. Hyg. 4: 109-118, lUus.

(59) and ROOT, FRANCIS M.

1923 PREFERENTIAL FEEDING EXPERIMENTS WITH ANOPHELINE MOSQUITOES. I. Amer. Jour. Hyg. 3: 514-520, illus.

(60) BUREN, WILLIAM F.

1946 SOME OBSERVATIONS ON THE LARVAL HABITAT OF PSOROPHORA

VARiPES (COQ.) (DíPTERA: CULICIDAE). Mosquito News 6:

120-121.

(61)

(62)

1946. PSOROPHORA PYGMAEA (THEOBALD), AN EXOTIC MOSQUITO NOW

ESTABLISHED IN FLORIDA. Mosquito News 6: 185.

1947 A NEW AEDES FROM THE FLORIDA KEYS (DÍPTERA, CULICIDAE).

Ent. Soc. Wash. Proc. 49: 228-229.

(63) BURROUGHS, A. L., and BURROUGHS, R. N.

1954. A STUDY OF THE ECOLOGY OF WESTERN EQUINE ENCEPHALOMYELITIS VIRUS IN THE UPPER MISSISSIPPI RIVER VALLEY.

Amer. Jour. Hyg. 60: 27-36.

(64) CARPENTER, STANLEY J.

1945. ARTIFICIAL SHELTERS FOR MEASURING DENSITIES OF ANOPHELES

QUADRIMACULATUS. Jour. Econ. Ent. (Sei. Note) 38: 721-

722.

(65)

1950. NOTES ON MOSQUITOES IN NORTH AMERICA: I NEW DISTRIBUTION RECORDS FOR EASTERN UNITED STATES DURING 1946 AND

1947. Mosquito News 10: 64-65.

(66) CHAMBERLAIN, ROY W., and WANAMAKER, JOHN F.

1945. NEW DISTRIBUTION RECORDS FOR THE MOSQUITOES OF THE SOUTHEASTERN STATES IN 1944. Jour. Econ. Ent. (Sei. Note) 38:

401-402.

(67) and JENKINS, DALE W.

1945. A NEW STATE RECORD OF MEGARHINUS RUTILUS COQUILLETT IN

SOUTH CAROLINA. Mosquito News 5: 88.

(68) and LACASSE, WALTER J.

1955. MOSQUITOES OF NORTH AMERICA (NORTH OF MEXICO). 360 pp.,

illus. Univ. of Calif. Press, Berkeley and Los Angeles.

(69) MIDDLEKAUFF, WOODROW W., and CHAMBERLAIN, ROY W.

1946. THE MOSQUITOES OF THE SOUTHERN UNITED STATES EAST OP

OKLAHOMA AND TEXAS. Amer. Midland Nat. Monog. No. 3,

292 pp., illus. The Univ. Press, Notre Dame, Ind.

(70) CHAMBERLAIN, ROY W., and DUFFEY, THOMAS E.

1945. COLLECTION RECORDS OF MANSONIA TITILLANS (WALKER) AND

MANSONIA INDUBITANS DYAR AND SHANNON IN FLORIDA WITH

KEYS TO THE SPECIES OF MANSONIA IN THE UNITED STATES

(DíPTERA, CULICIDAE). Mosquito News 5: 96-97.

(71) RUBIN, HARRY, KISSLING, R. E., and EIDSON, M. E.

1951. RECOVERY OF THE VIRUS OF EASTERN EQUINE ENCEPHALOMYELITIS FROM A MOSQUITO, CULISETA MELANURA (COQUILLETT) .

Soc. Expt. Biol. and Med. Proc. 77: 396-397.

(72) SiKES, R. K., NELSON, D. B., and SUDIA, W. D.

1954. STUDIES ON THE NORTH AMERICAN ARTHROPOD-BORNE ENCEPHALITIDES. VI. QUANTITATIVE DETERMINATIONS OF VIRUS-VECTOR RELATIONSHIPS. Amer. Jour. Hyg. 60: 278-285.

(73) SuDiA, W. D., and NELSON, D. B.

1955. LABORATORY OBSERVATIONS ON A MOSQUITO, CULISETA MELANURA (COQUILLETT). Mosquito News 15: 18-21.

(74) CHAMBERLIN, JOSEPH C, and LAWSON, F. R.

1945. A MECHANICAL TRAP FOR THE SAMPLING OF AERIAL INSECT

POPULATIONS. Mosquito News 5: 4-7, illus.

(75) CHAPMAN, H. C, and FERRIGNO, F.

1956. A THREE-YEAR STUDY OF MOSQUITO BREEDING IN NATURAL AND

IMPOUNDED SALT-MARSH AREAS IN NEW JERSEY. N. J. MoSquito Extermin. Assoc. Proc. 43: 48-60.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 161

(76) CHRISTOPHER, S. R.

1951. NOTE ON MORPHOLOGICAL CHARACTERS DIFFERENTIATING CULEX

PIPIENS L. FROM CULEX MOLESTUS FORSKAL AND THE STATUS

OF THESE FORMS. In Mattingly et al., The Ciilex pipiens

Complex. Royal Ent. Soc, London, Trans. 102 (7) :

372-379.

(77) CLARKE, J. LYELL

1943. STUDIES OF THE FLIGHT RANGE OF MOSQUITOES. Jour. Econ.

Ent. 36: 121-122.

(78) COAD, B. R.

1913. OVIPOSITION HABITS OF CULEX ABOMINATOR DYAR AND KNAB.

Can. Ent. 45: 265-266.

(79) COOK, S. S., and WILLIAMS, L. L., JR.

1928. AIRPLANES AND PARIS GREEN IN CONTROL OF ANOPHELES PRODUCTION. South. Med. Jour. 21: 754-759, illus. [Discussion,

pp. 759-760.]

(80) COQUILLET, D. W.

1903. FOUR NEW SPECIES OF CULEX. Canad. Ent. 35: 255-257.

(8^) CovELL, GORDON.

[1931.] MALARIA CONTROL BY ANTI-MOSQUITO MEASURES. 148 pp.,

illus. Calcutta, Simla, and London.

(82) DAGGY, RICHARD H., MUEGGE, OSWALD J., and RILEY, WILLIAM A.

1941. A PRELIMINARY SURVEY OF THE ANOPHELINE MOSQUITO FAUNA

OF SOUTHEASTERN MINNESOTA AND ADJACENT WISCONSIN

AREAS. U. S. Pub. Health Serv. Rpts. 56: 883-895, illus.

(83) DARLING, S. T.

1925. DISCUSSION ON RELATIVE IMPORTANCE IN TRANSMITTING MALARIA OF ANOPHELES QUADRIMACULATUS, PUNCTIPENNIS, AND

CRUCIANS AND ADVISABILITY OF DIFFERENTIATING BETWEEN

THESE SPECIES IN APPLYING CONTROL MEASURES. South.

Med. Jour. 18: 452-457. [Discussion, pp. 457-458.]

(84) DAVIS, WILLIAM A.

1940. A STUDY OF BIRDS AND MOSQUITOES AS HOSTS FOR THE VIRUS

OF EASTERN EQUINE ENCEPHALOMYELITIS. Amer. Jour. Hyg.

45-59.

(85) DEONIER, CHRISTIAN C, BURRELL, R. W., MAPLE, JOHN D., and

COCHRAN, J. H.

1945. DDT AS AN ANOPHELINE LARVICIDE: PRELIMINARY FIELD STUDIES.

Jour. Econ. Ent. 38: 244-249.

(86) CAIN, T. L., JR., and MCDUFFIE, W. C.

1950. AERIAL SPRAY TESTS ON ADULT SALT-MARSH MOSQUITOES RESISTANT TO DDT. Jour. Econ. Ent. 43: 506-510.

(87) MAPLE, JOHN D., JONES, HOWARD A., HINCHEY, EDNA, and

EiDE, PAUL M.

1945. DDT AS AN ANOPHELINE LARVICIDE—LABORATORY TESTS. Jour.

Econ. Ent. 38: 241-243.

(88) DODGE, H. R.

1945. NOTES ON THE MORPHOLOGY OF MOSQUITO LARVAE. ENT. SOC.

Amer. Ann. 38: 163-167, illus..

(89)

1947. A NEW SPECIES OF WYEOMYIA FROM THE PITCHER PLANT (DÍPTERA, CULICIDAE). Ent. Soc. Wash. Proc. 49: 117-122, illus.

(90) DORER, R. E., CARTER, R. G., and BICKLEY, W. E.

1950. OBSERVATIONS ON THE PUPAE OF MANSONIA PERTURBANS

(WALK.) IN VIRGINIA. N. J. Mosquito Extermin. Assoc.

Proc. 37: 110-112.

(91) DOZIER, HERBERT L.

1936. OBSERVATIONS ON BREEDING PLACES AND WINTER ACTIVITIES OF

MOSQUITOES IN THE VICINITY OF NEW ORLEANS, LOUISIANA.

Ent. Soc. Wash. Proc. 38: 148-155, illus.

(92) DUNN, L. H.

1918. THE LAKE MOSQUITO, MANSONIA TITILLANS WALK., AND ITS

HOST PLANT, PISTIA STRATIOTES LINN., IN THE CANAL ZONE,

PANAMA (DíPTERA: CULICIDAE). Ent. News. 29: 260-269;

288-295.

(93) DUPREE, J. W.

1905 THE MOSQUITOES OF LOUISIANA AND THEIR PATHOGENIC POSSIBILITIES, WITH REMARKS UPON THEIR EXTERMINATION.

New Orleans Med. and Surg. Jour. 58: 1-16.
162 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(94

(95

(96

(97

(98

(99

(100

(101

(102

(103

(104

(105

(106

(107

(108

(109

(110

(111

(112

(113

(114

DYAR, HARRISON G. O • I«.

VM)2. THE EGGS OF MOSQUITOES OF THE GENUS CULEX. fecience lb.

672-673.

1922. THE MOSQUITOES OF THE UNITED STATES. U. S. Nati. MuS.

Proc. 62, art. 1, 119 pp.

1928. THE MOSQUITOES OF THE AMERICAS. Carnegie Inst. Wash.

Pub. 387, 616 pp., illus.

and BARRET, HARVEY P.

1918. DESCRIPTIONS OF HITHERTO UNKNOWN LARVAE OF CULEX

(DíPTERA, CULICIDAE). Insecutor Inscitiae Menstruus 6:

119-120.

and KNAB, FREDERICK.

1906. THE LARVAE OF CULICIDAE CLASSIFIED AS INDEPENDENT ORGANISMS. N. Y. Ent. Soc. Jour. 14: 169-230.

EADS, RICHARD B.

1946. A NEW RECORD OF ANOPHELES ALBIMANUS IN TEXAS. Jour.

Econ. Ent. 39: 420.

and MENZIES, GEORGE C.

1950. DISTRIBUTION RECORDS OF MANSONIA BLANCHARD (DÍPTERA,

CULICIDAE) IN TEXAS. Mosquito News 10: 3-5.

EDGAR, S. A., and WILLIAMS, 0. M.

1948. EFFECT OF MOSQUITOES ON POULTRY. Poultry Sei. 27: 660.

EDWARDS, F. W.

1932. DíPTERA, FAM. CULICIDAE. In Wytsman, P., Genera Insectorum, fase. 194, 258 pp., illus. Bruxelles.

EIDE, P. M., DEONIER, C. C, and BURRELL, R. W.

1945. DDT AS A CULICINE LARViciDE. Jour. Eeon. Ent. 38: 537-541.

ENGINEERING NEWS-RECORD, EDITORS.

1936. MOSQUITO CONTROL ENGINEERING. 57 pp., illus. New York.

EYLES, D. E.

1944. A CRITICAL REVIEW OF THE LITERATURE RELATING TO THE FLIGHT

AND DISPERSION HABITS OF ANOPHELINE MOSQUITOES. U. S.

Pub. Health Serv. Bui. 287, 39 pp.

and MOST, HARRY.

1947. INFECTIVITY OF PACIFIC ISLAND WUCHERERIA BANCROFTI TO

MOSQUITOES OF THE UNITED STATES. Amer. Jour. Trop.

Med. 27: 211-220.

FARID, MOHYEDDIN AHMED.

1949. RELATIONSHIPS BETWEEN CERTAIN POPULATIONS OF CULEX PIPlENS LINNAEUS AND CULEX QUINQUEFASCIATUS SAY IN THE

UNITED STATES. Amer. Jour. Hyg. 49: 83-100.

FELLTON, HERMAN L.

1944. THE BREEDING OF THE SALT-MARSH MOSQUITO IN MIDWESTERN

STATES. Jour. Eeon. Ent. 37: 245-247.

FERGUSON, FREDERICK F.

1954. BIOLOGICAL FACTORS IN THE TRANSMISSION OF AMERICAN

ARTHROPOD-BORNE VIRUS ENCEPHALITIDES. A summary.

U. S. Pub. Health Serv. Monog. No. 23, 37 pp.

FiSK, FRANK W.

1939. NEW MOSQUITO RECORDS FROM KEY WEST, FLA. (Sei Note)

Jour. Econ. Ent. 32: 469.

and LEVAN, JAMES H.

1940. MOSQUITO COLLECTIONS AT CHARLESTON, SOUTH CAROLINA,

USING THE NEW JERSEY LIGHT TRAP. (Sci. Note) Jour.

Econ. Ent. 33: 578-579.

FLETCHER, ORLIN K., JR., and FLANAGAN, LEON.

1957. WINTER BREEDING OF CULEX QUINQUEFASCLATUS SAY IN COLUMBUS, GEORGIA. Mosquito News 17: 29-32.

FooTE, RICHARD H.

1952. THE LARVAL MORPHOLOGY AND CHAETOTAXY OF THE CULEX SUBGENUS MELANOCONION. (DÍPTERA, CULICIDAE). Ent. SoC.

Amer. Ann. 45: 445-472, illus.

1954. THE LARVAE AND PUPAE OF THE MOSQUITOES BELONGING TO THE

CULEX SUBGENERA MELANOCONION AND MOCHLOSTYRAX.

U. S. Dept. Agr. Tech. Bui. 1091, 125 pp., illus.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 163

(114a) and COOK, DAVID R.

1959. MOSQUITOES OF MEDICAL IMPORTANCE. U. S. Dept. Agr.,

Handb. No. 152. 158 pp., illus.

(115) FREEBORN, STANLEY B.

1949. ANOPHELINES OF THE NEARCTic REGION. In Boyd, Malariology, V. I, pp. 379-398.

(116) FROHNE, W. C.

1939. ANOPHELINE BREEDING: SUGGESTED CLASSIFICATION OF PONDS

BASED ON CHARACTERISTIC DESMIDS. U. S. Pub. Health Serv.

Rpts. 54: 1363-1387.

(117) WEATHERSBEE, ALBERT A., WILLIAMS, GILES M., and HART,

JOHN W.

1950. OBSERVATIONS OF THE PERSISTENCE OF PLASMODIUM INFECTIONS

IN ANOPHELES MOSQUITOES IN AN AREA OF LOW OBSERVED

HUMAN MALARIA PARASITEMIA IN SOUTH CAROLINA. Nati.

Malaria Soc. Jour. 9: 10-24.

(118) GAHAN, JAMES B., BERTHOLF, J. H., DAVIS, A, N., JR., and SMITH,

CARROLL N.

1956. FIELD TESTS WITH TWO PHOSPHOROTHIOATES AGAINST RESISTANT

SALT-MARSH MOSQUITOES. Mosquito News 16: 91-93.

(119) TRAVIS, B. V., and LINDQUIST, ARTHUR W.

1945. DDT AS A RESIDUAL-TYPE SPRAY TO CONTROL DISEASE-CARRYING

MOSQUITOES: LABORATORY TESTS. Jour. Econ. Ent. 38:

236-240.

(120) TRAVIS, B. V., MORTON, FRED A., and LINDQUIST, ARTHUR W.

1945. DDT AS A RESIDUAL-TYPE TREATMENT TO CONTROL ANOPHELES

QUADRIMACULATUS: PRACTICAL TESTS. Jour. Econ. Ent. 38:

231-235.

(121) GEIGER, J. C, PURDY, W. C, and TARBETT, R. E.

1919. EFFECTIVE MALARIA CONTROL IN A RICEFIELD DISTRICT WITH

OBSERVATIONS ON EXPERIMENTAL MOSQUITO FLIGHTS. Jour.

Amer. Med. Assoc. 72: 844-847, illus.

(122) GILBERT, I. H., GOUCK, H. K., and SMITH, CARROLL N.

1957. NEW INSECT REPELLENT. Soap and Chem. Spec. 33(5): 115-

117, 129-133; 33(6): 95, 97, 99, 109.

(123) GiNSBURG, JOSEPH M.

1936. PROTECTION OF OUTDOOR GATHERINGS FROM THE MOSQUITO PEST.

N. J. Mosquito Extermin. Assoc. Proc. 23: 166-173.

(124)

1937. PRINCIPLES UNDERLYING THE PROTECTION OF OUTDOOR MEETINGS

FROM THE MOSQUITO PEST AND METHOD OF APPLICATION.

N. J. Mosquito Extermin. Assoc. Proc. 24: 5-11.

(125) GJULLIN, C. M., HEGARTY, C. P., and BOLLEN, W. P.

1941. THE NECESSITY OF A LOW OXYGEN CONCENTRATION FOR THE

HATCHING OF AEDES MOSQUITO EGGS. Jour. Cell, and Compar. Physiol. 17: 193-202.

(126) and ISAAK, LEWIS W.

1957. PRESENT STATUS OF MOSQUITO RESISTANCE TO INSECTICIDES IN

THE SAN JOAQUÍN VALLEY IN CALIFORNIA. Mosquito News

17: 67-70.

(126a) GJULLIN, C. M., SAILER, R. I., STONE, ALAN, and TRAVIS, B. V.

1960. THE MOSQUITOES OF ALASKA. U. S. Dept. Agr., Agr. Handb.

No. 182. [In Press.]

(127) YATES, W. W., and STAGE, H. H.

1950. STUDIES ON AEDES VEXANS (MEIG.) AND AEDES STICTICUS

(MEIG.), FLOOD-WATER MOSQUITOES, IN THE LOWER COLUMBIA

RIVER VALLEY. Ent. Soc. Amer. Ann. 43: 262-275.

(128) GooDHUE, L. D., and SULLIVAN, W. N.

1943. METHOD OF APPLYING PARASITICIDES. U. S. Patent 2,321,303,

issued June 8.

(129) GRANETT, PHILIP.

1940. STUDIES OF MOSQUITO REPELLENTS. I. TEST PROCEDURE AND

METHOD OF EVALUATING TEST DATA. II. RELATIVE PERFORMANCE OF CERTAIN CHEMICALS AND COMMERCIALLY AVAILABLE

MIXTURES AS MOSQUITO REPELLENTS. Jour. Econ. Ent. 33:

563-572, illus.

(130) and HAYNES, HARRY L.

1945. INSECT-REPELLENT PROPERTIES OF 2-ETHYLHEXANEDI0L-1, 6,

Jour. Econ. Ent. 38: 671-675.
(134)

(143)

(144)

(145)

164 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(131) GRIFFITTS, T. H. D. -TT O T. V

1918. WINTER HIBERNATION OF ANOPHELES LARVAE. U. b. rub.

Health Serv. Rpts. 33: 1996-1998.

(132)

1927. ANOPHELES ÁTROPOS DYAR AND KNAB. A NOTE ON ITS BREEDING

AND OTHER HABITS. U. S. Pub Health Serv. Rpts. 42:

1903-1905.

(133) HAEGER, JAMES S.

1949. A REVIEW OF ANOPHELES ALBIMANUS BREEDING ON THE FLORIDA

KEYS. Fla. Anti-Mosquito Assoc. Rpt., 20th Ann. Meeting,

pp. 118-121.

1950. THE PRESENT STATUS OF ANOPHELES ALBIMANUS IN THE FLORIDA KEYS. Fla. Anti-Mosquito Assoc. Rpts., 21st Ann.

Meeting, pp. 118-119.

(135) HAMMON, W. MCD, and REEVES, W. C.

1943. LABORATORY TRANSMISSION OF ST. LOUIS ENCEPHALITIS VIRUS

BY THREE GENERA OF MOSQUITOES. Jour. Expt. Med. 78:

241-253.

(136) and REEVES, W. C.

1945. RECENT AD\'ANCES IN THE EPIDEMIOLOGY OF THE ARTHROPODBORNE VIRUS ENCEPHALITIDES INCLUDING CERTAIN EXOTIC

TYPES. Amer. Jour. Pub. Health 35: 994-1004.

(137) and REEVES, W. C.

1947. INTEREPIDEMIC STUDIES ON ARTHROPOD-BORNE VIRUS ENCEPHALITIDES AND POLIOMYELITIS IN KERN COUNTY, CALIFORNIA, AND THE YAKIMA VALLEY, WASHINGTON, 1944.

Amer. Jour. Hyg. 46: 326-335.

(138) REEVES, W. C, BENNER, S. R., and BROOKMAN, BERNARD.

1945. HUMAN ENCEPHALITIS IN THE YAKIMA VALLEY, WASHINGTON,

1942, WITH FORTY-NINE VIRUS ISOLATIONS (WESTERN EQUINE

AND ST. LOUIS TYPES) FROM MOSQUITOES. Jour. Amcr. Med.

Assoc. 128: 1133-1139.

(139) REEVES, W. C, and GALINDO, P.

1945. EPIDEMIOLOGIC STUDIES OF ENCEPHALITIS IN TPIE SAN JOAQUÍN

VALLEY OF CALIFORNIA, 1943, WITH THE ISOLATION OF VIRUSES

FROM MOSQUITOES. Amer. Jour. Hyg. 42: 299-30S.

(140) HARDEN, P. H.

1945. THE OCCURRENCE OF ORTHOPODOMYIA ALBA BAKER AT NEW ORLEANS, LOUISIANA. Mosquito News 5: 131.

(141) HAYES, RICHARD O., and MORLAN, HARVEY B.

1957. NOTES ON AEDES TRISERIATUS EGG INCUBATION AND COLONIZATION. Mosquito News 17: 33-36.

(142) HEADLEE, THOMAS J.

1921. THE MOSQUITOES OF NEW JERSEY AND THEIR CONTROL. N. J.

Agr. Expt. Sta. Bui. 348, 229 pp., illus.

1934. MOSQUITO WORK IN NEW JERSEY FOR THE YEAR 1933. N. J.

Mosquito Extermin. Assoc. Proc. 21: 8-37.

1941. NEW JERSEY MOSQUITO PROBLEMS. N. J. Mosquito Externiin.

Assoc. Proc. 28: 7-12.

1945. THE MOSQUITOES OF NEW JERSEY AND THEIR CONTROL. 326 pD

,,,^, ^ Rutgers Univ. Press, New Brunswick, N. J.

(146) HEDEEN, R. A.

1953. THE BIOLOGY OF THE MOSQUITO AEDES ATROPALPUS (COQUILLET). Kans. Ent. Soc. Jour. 26: 1-10.

(147)

1954. THE IDENTIFICATION OF THE LARVA OF PSOROPHORA (GRABHAMiA) siGNiPENNis (COQUILLETT) . MosQuito News 14:

143-145.

(148) HERMS, WILLIAM BRODBECK, and GRAY, HAROLD FARNSWORTH.

1944. MOSQUITO CONTROL—PRACTICAL METHODS FOR ABATEMENT OF

DISEASE VECTORS AND PESTS. 419 pp., illus. Oxford University Press. New York.

(149) HiNMAN, E. HAROLD.

1930. A STUDY OF THE FOOD OF MOSQUITO LARVAE (CULICIDAE).

Amer. Jour. Hyg. 12: 238-270.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 165

1931. THE WINTER BREEDING AND ACTIVITY OF CULICINE MOSQUITOES

AT NEW ORLEANS (30" N. LAT.) Amer. Jour. Trop. Med.

11: 459-467, illus.

1932. A DESCRIPTION OF THE LARVA OF ANOPHELES ÁTROPOS D. & K.

WITH BIOLOGICAL NOTES ON THE SPECIES. Ent. Soc. Wash.

Proc. 34: 138-142.

1934. PREDATORS OF THE CULICIDAE (MOSQUITOES). I.—THE PREDATORS OF LARVAE AND PUPAE, EXCLUSIVE OF FISH. Jour.

Trop. Med. and Hyg. [London] 37: 129-134.

1934. PREDATORS OF THE CULICIDAE (MOSQUITOES). II.—PREDATORS

OF ADULT MOSQUITOES. Jour. Trop. Med. and Hyg. [London] 37: [145J-150.

1935. BIOLOGICAL NOTES ON URANOTAENIA SPP. IN LOUISIANA (CULICIDAE, DíPTERA). Ent. Soc. Amer. Ann. 28: 404-407.

1938. BIOLOGICAL EFFECTS OF FLUCTUATION OF WATER LEVEL ON

ANOPHELINE BREEDING. Amer. Jour. Trop. Med. 18: 483-

495, illus.

HORSFALL, W. R.

1936. OCCURRENCE AND SEQUENCE OF MOSQUITOES IN SOUTHEASTERN

ARKANSAS IN 1935. Jour. Econ. Ent. 29: 676-679.

1937. MOSQUITOES OF SOUTHEASTERN ARKANSAS. Jour. Econ. Ent.

30: 743-748.

1939. HABITS OF AEDES THIBAULTI DYAR AND KNAB (DÍPTERA, CULICIDAE). Kans. Ent. Soc. Jour. 12: 70-71.

1942. BIOLOGY AND CONTROL OF MOSQUITOES IN THE RICE AREA.

Ark. Agr. Expt. Sta. Bui. 427: 1-46.

1955. MOSQUITOES, THEIR BIONOMICS AND RELATION TO DISEASE.

723 pp. The Ronald Press, New York.

MILES, ROBERT C, and SOKATCH, JOHN T.

19 o2. EGGS OF FLOODWATER MOSQUITOES. I.—SPECIES OF PSOROPHORA.

Ent. SOC. Amer. Ann. 45: 618-624.

HOWARD, L. O.

1921. (MOSQUITO WORK AT MOUND, LA.] N. J. Mosquito Extermin.

Assoc. Proc. 8: 75-83, illus.

DYAR, HARRISON G., and KNAB, FREDERICK.

1912-17. THE MOSQUITOES OF NORTH AND CENTRAL AMERICA AND

THE WEST INDIES. Carnegie Inst. Wash. Pub. 159. 4 v.,

illus. Washington, D. C.

HowiTT, B. F., DODGE, H. R., BISHOP, L. K., and GORRIE, R. H.

1949. RECOVERY OF THE VIRUS OF EASTERN EQUINE ENCEPHALOMYELITIS FROM MOSQUITOES (MANSONIA PERTURBANS) COLLECTED

IN GEORGIA. Science 110: 141-142.

Hu, S. M. K.

1931. STUDIES ON HOST-PARASITE RELATIONSHIPS OF DIROFILARIA

IMMITIS LEIDY AND ITS CULICINE INTERMEDIATE HOSTS.

Amer. Jour. Hyg. 14: 614-629.

HUFF, CLAY G. _ _

1929. OVULATION REQUIREMENTS OF CULEX PIPIENS LINN. Mar.

Biol. Lab., Woods Hole, Mass. Biol. Bui. 56: 347-350.

HUFFAKER, CARL B., and BACK, RICHARD C.

1943 A STUDY OF METHODS OF SAMPLING MOSQUITO POPULATlcrNS.

Jour. Econ. Ent. 36: 561-569.

HüLSE, F. E. * T ü U

19'?2 SKETCH MAPPING FOR HEALTH PROBLEMS. Amer. Jour. I'ub.

Health 12: 1034-1038, illus.

^938^'^Á STUDY OF THE LARVAL CHAETOTAXY OF ANOPHELES WALKERI

* THEOBALD. Amer. Jour. Hyg. 28: 149-173, illus.
166 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

1938 FURTHER NOTES ON THE OVERWINTERING OF THE EGGS OF

ANOPHELES WALKERI THEOBALD WITH A DESCRIPTION OF THE

EGGS. Jour. Parasitol. 24: 521-526, illus.

(171) HUSMAN, C. N.

1953. A LIGHT PNEUMATIC SPRAYER FOR INSTALLATION ON VEHICLES.

Mosquito News 13: 134-138.

(172) IRONS, J. V. ^, .^ .,

1956. MOSQUITO-BORNE DISEASES IN TEXAS. Mosquito News lb:

64-66.

(173) JENKINS, DALE W.

1949 TOXORHYNCHITES MOSQUITOES OF THE UNITED STATES (DÍPTERA,

CULICIDAE). Ent. Wash. Proc. 51: 225-229, illus.

(174) and CARPENTER, STANLEY J.

1946. ECOLOGY OF THE TREE HOLE BREEDING MOSQUITOES OF NEARCTIC

NORTH AMERICA. Ecol. Monog. 16: 33-47, illus.

(175) JOHNSON, H. A. -, T,T i

1936. NOTES ON THE OCCURRENCE OF A. WALKERI. South. Med.

Jour. 29: 856-857.

(176) KARSTAD, L. H., FLETCHER, 0. K., SPALATIN, J., ROBERTS, R., and

HANSON, R. P.

1957. EASTERN EQUINE ENCEPHALOMYELITIS VIRUS ISOLATED FROM

THREE SPECIES OF DíPTERA FROM GEORGIA. Science 125:

395-396.

(177) KARTMAN, L.

1954. FREQUENCY AND INTENSITY OF DIROFILARIA IMMITIS INFECTIONS

IN MOSQUITOES. Expt. Parasitol. 3: 2.5-29.

(178) KEEFE, B. F., and BEADLE, L. D.

1956. A DIGEST OF STATE ENABLING LEGISLATION FOR MOSQUITO

ABATEMENT THROUGH 1955. U. S. Dept. of Health, Education, and Welfare, Pub. Health Serv., Communicable

Disease Center Pub., 83 pp., (processed).

(179) KELLER, J. C, CHAPMAN, H. C, and LABRECQUE, G. C.

1954. TESTS WITH GRANULATED INSECTICIDES FOR THE CONTROL OF

SALT-MARSH MOSQUITO LARVAE. Mosquito News 14: 5-9.

(180) KELSER, R. A.

1933. MOSQUITOES AS VECTORS OF THE VIRUS OF EQUINE ENCEPHALOMYELITIS. Amer. Vet. Med. Assoc. Jour. 82: 767-771.

;i81) KiKER, CALVIN C, FAIRER, CHARLES D., and FLANERY, PAUL N.

1938. FURTHER OBSERVATIONS ON AIRPLANE DUSTING FOR ANO'^HELES

LARVAL CONTROL. South. Med. Jour. 31: 808-813, illus.

(182) KING, W. V.

1924. ANOPHELINE INFECTION UNDER NATURAL CONDITIONS. South.

Med. Jour. 17: 596-597.

(183)

(184)

(185)

1928. THE ANOPHELES DENSITY INDEX IN MALARIA RESEARCH AND

CONTROL WORK. South. Med. Jour. 21: 763-767, illus.

1937. ON THE DISTRIBUTION OF ANOPHELES ALBIMANTJS /ND ITS OCCURRENCE IN THE UNITED STATES. South. Med. Jour. 30:

943-946.

1939. VARIETIES OF ANOPHELES CRUCIANS WIED. Amer. Jovr. Trop.

Med. 19: 461-471, illus.

(186) Compiler.

1954. CHEMICALS EVALUATED AS INSECTICIDES AND REPELLENTS AT

ORLANDO, FLA. U. S. Dept. Agr., Agr. Handb. 69, 397 pp.

(187) and BRADLEY, G. H.

1926. AIRPLANE DUSTING IN THE CONTROL OF MALARIA MOSQUITOES.

U. S. Dept. Agr. Cir. 367, 16 pp., illus.

(188) and BRADLEY, G. H.

1926. AIRPLANE DUSTING IN THE CONTROL OF MALARIA MOSQUITOES.

Jour. Trop. Med. and Hyg. [London] 29: 311-313, illus.

(189) and BRADLEY, G. H.

1937. NOTES ON CULEX ERRATICUS AND RELATED SPECIES IN THE

UNITED STATES (DÍPTERA, CULICIDAE). Ent. Soc. Amer.

Ann. 30: 345-357, illus.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 167

(190) and BRADLEY, G. H.

1941. GENERAL MORPHOLOGY OF ANOPHELES AND CLASSIFICATION OF

THE NEARCTic SPECIES. In Human Malaria. Amer. Assoc.

Adv. Sei. Pub. 15, pp. 63-70, illus.

(191) and BRADLEY, G. H.

1941. DISTRIBUTION OF THE NEARCTIC SPECIES OF ANOPHELES. In

Human Malaria. Amer. Assoc. Adv. Sei. Pub. 15, pp.

71-78, illus.

(192) BRADLEY, G. H., and Mc NEEL, T. E.

1937. EXPERIMENTS IN FLORIDA IN REPELLING MOSQUITOES BY OUTDOOR SPRAYING. N. J. Mosquito Extermin. Assoe. Proc.

24: 163-172.

(193) BRADLEY, G. H., and MCNEEL, T. E.

1944. THE MOSQUITOES OF THE SOUTHEASTERN STATES. U. S. Dept.

Agr. Mise. Pub. 336, rev. 96 pp., illus. [Originally published in 1939; revised 1942 and 1944.]

(194) and BULL, CAROLL G.

1923. THE BLOOD FEEDING HABITS OF MALARIA-CARRYING MOSQUITOES.

Amer. Jour. Hyg. 3: 497-513, illus.

(195) ROTH, LOUIS, TOFFALETI, JAMES, and MIDDLEKAUFF, WOODROW W.

1943. NEW DISTRIBUTION RECORDS FOR THE MOSQUITOES OF THE

SOUTHEASTERN UNITED STATES DURING 1942. Jour. Eeon.

Ent. 36: 573-577.

(196) KissLiNG, R. E., CHAMBERLAIN, R. W., NELSON, D. B., and STAMM,

D. D.

1955. STUDIES ON THE NORTH AMERICAN ARTHROPOD-BORNE ENCEPHALITIDES. VIII.—EQUINE ENCEPHALITIS STUDIES IN

LOUISIANA. Amer. Jour. Hyg. 62 (3) : 233-254, illus.

(197) and RUBIN, H.

1950. EASTERN EQUINE ENCEPHALOMYELITIS. U. S. Dept. of Health,

Edueation, and Welfare, Communieable Disease Center

Bui. 9: 2-5.

(198) KITCHEN, S. F., and BRADLEY, G. H.

1936. ANOPHELES WALKERI THEOBALD AS A VECTOR OF PLASMODIUM

FALCIPARUM (WELCH). Amer. Jour. Trop. Med. 16: 579-

581.

(199) KiTZMILLER, J. B.

1952. INBRED STRAINS OF CULEX MOSQUITOES. Seienee 116: 66-67.

(200) KLIGLER, I. J., MucKENFUss, R. S., and RIVERS, T. M.

1929. TRANSMISSION OF FOWL-POX BY MOSQUITOES. Jour. Expt.

Med. 49: 649-660.

(201) KNIGHT, KENNETH L.

1951. A REVIEW OF THE CULEX PIPIENS COMPLEX IN THE MEDITERRANEAN SUBREGION (DíPTERA, CULICIDAE). In Mattingly

et al, The Cidex pipiens Complex. Royal Ent. Soe., London,

Trans. 102: 354-364.

(202)

(203)

1954. MOSQUITO LIGHT TRAP COLLECTIONS AT YUKON, FLORIDA, FOR

THE SIX-YEAR PERIOD, 1948-1953. N. J. Mosquito Extermm.

Assoe. Proc. 41: 251-257.

1954. A COMPARISON OF THE NEW JERSEY LIGHT TRAP AND THE DAYTIME RESTING STATION SAMPLING METHODS FOR THE DETERMINATION OF THE ABUNDANCE OF AEDES INFIRMATUS D. & K.

ADULTS. Fla. Anti-Mosquito Assoe. Rpt., 25th Ann. Meeting, pp. 100-104, illus. [Mimeographed.]

(204) ^^^^^^^^'jjf•j^^;^LOPMENT AND USE OF DDT FOR THE CONTROL OF MOSQUITOES. Nati. Malaria Soe. Jour. 4: 77-92.

(205) KOMP W-JI.JV^ ANOPHELES WALKERI AND ANOPHELES ATRO192b. 0^^^ CULICIDAE). Inseeutor Inseitiae Menstruus

14: 168-176.

(206) KUMM, HENRY W^ ^^^ i^isPERSION OF ANOPHELES MOSQUITOES. Amer.

' Jour. Trop. Med. 9: 67-77, illus.
168 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(207) LE PRINCE, J. A. A., and GRIFFITTS, T. H. D.

1917. FLIGHT OF MOSQUITOES. STUDIES ON THE DISTANCE OF FLIGHT

OF ANOPHELES QUADRIMACULATUS. U.S. Pub. Health Serv.

Rpts. 32: 656-659, illus.

(208) MACCREARY, DONALD.

1941. COMPARATIVE DENSITY OF MOSQUITOES AT GROUND LEVEL AND

AT AN ELEVATION OF APPROXIMATELY ONE-HUNDRED FEET.

Jour. Econ. Ent. 34: 174-179, illus.

(209) and STEARNS, L. A.

1937. MOSQUITO MIGRATION ACROSS DELAV^ARE BAY. N. J. Mosquito Extermin. Assoc. Proc. 24: 188-197, illus.

(210) MCNEEL, T. E.

1931. A METHOD FOR LOCATING THE LARVAE OF THE MOSQUITO MANSONIA. Science 74: 155.

1211)

1932. OBSERVATIONS ON THE BIOLOGY OF MANSONIA PERTURBANS

(v^ALK.) DíPTERA, CULICIDAE. N. J. Mosquito Extermin.

Assoc. Proc. 19: 91-96, illus.

(212) MATHESON, ROBERT.

1930. THE UTILIZATION OF AQUATIC PLANTS AS AIDS IN MOSQUITO

CONTROL. Amer. Nat. 64: 56-86, illus.

(213)

1944. A HANDBOOK OF THE MOSQUITOES OF NORTH AMERICA. 314 pp.

Comstock Pub. Assoc, Ithaca, N. Y.

(214) BoYD, MARK F., and STRATMAN-THOMAS, WARREN K.

1933. ANOPHELES W^ALKERI, THEOBALD, AS A VECTOR OF PLASMODIUM

viVAX, GRASSI AND FELETTI. Amer. Jour. Hyg. 17: 515-516.

(215) and HURLBUT, H. S.

1937. NOTES ON ANOPHELES WALKERI THEOBALD. Amer. Jour. Trop.

Med. 17: 237-243, illus.

(216) BRüNETT, E. L., and BRODY, A. L.

1931. THE TRANSMISSION OF FOWL-POX BY MOSQUITOES, PRELIMINARY

REPORT. Poultry Science 10: 211-223.

(217) MATTINGLY, P. E.

1956. SPECIES HYBRIDS IN MOSQUITOES. Roy. Ent. Soc, London,

Trans. 108: 21-36.

(218) RozEBOOM, L. E., KNIGHT, K. L., LAVEN, H., DRUMMOND,

F. H.. CHRISTOPHERS, S. R., and SHUTE, P. G.

1951. THE CULEX PipiENS COMPLEX. Roy. Ent. Soc, London, Trans.

102: 331-382, illus.

(219) SHUTE, P. G., LAVEN, H., and KNIGHT, K. L.

1953. THE CULEX PIPIENS COMPLEX. Trans. IXth International Con-

,^^^, ,, gress Ent. II: 283-300, illus.

(220) MAYNE, BRUCE.

1926. REPORT OF A SURVEY TO DETERMINE THE MALARIA PREVALENCE

IN THE OKEFENOKEE SWAMP. U.S. Pub. Health Serv. Rpts

,^^^, 41: 1652-1660, illus. ^

(221) and GRIFFITTS, T. H. D.

1931. ANOPHELES ÁTROPOS D. AND K.—A NEW POTENTIAL CARRIER OF

MALARIA ORGANISMS. U. S. Pub. Health Serv. Rpts. 46:

3107-3115, illus.

(222) MELENEY, HENRY E., BISHOP, EUGENE L., and ROBERTS, FRANK L.

ly^y. OBSERVATIONS ON THE MALARIA PROBLEM OF WEST TENNESSEE.

^99Q^ ii/r J^^^^' ^^^' J^^^- 22: 382-394, illus.

(223) MERRILL, MALCOLM H., LACAILLADE, C. WM., and TEN BROECK, CARL.

1934. MOSQUITO TRANSMISSION OF EQUINE ENCEPHALOMYELITIS.

,^^^^. n.. Science 80: 251-252.

(224) MICHENER, CHARLES D.

1945. SEASONAL VARIATIONS IN CERTAIN SPECIES OF MOSQUITOES

(225) (DÍPTERA, CULICIDAE). Jour. N. Y. Ent. Soc. 53: 293-300.

1947. MOSQUITOES OF A LIMITED AREA IN SOUTHERN MISSISSIPPI.

Amer. Mid. Nat. 37: 325-374.

(226) MiDDLEKAUFF, WOODROW W.

1944. A NEW SPECIES OF AEDES FROM FLORIDA (DÍPTERA, CULICIDAE).

Ent. Soc. Wash. Proc. 46: 42-44.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 169

(227)

1944. A RAPID METHOD FOR MAKING PERMANENT MOUNTS OF MOSQUITO LARVAE. Science 99: 206.

(228) and CARPENTER, STANLEY J.

1944. NEW DISTRIBUTION RECORDS FOR THE MOSQUITOES OF THE

SOUTHEASTERN UNITED STATES IN 1943. Jour. Econ. Ent.

37: 88-92.

(229) MILES, VIRGIL I.

1945. DIFFERENTIATING THE LARVAE OF ANOPHELES GEORGIANUS KING,

A. BRADLEYI KING, AND A. PUNCTIPENNIS (SAY). Nati.

Malaria Soc. Jour. 4: 235-242.

(230) and RINGS, ROY W.

1946. DISTRIBUTION RECORDS FOR MOSQUITOES OF THE SOUTHEASTERN

STATES IN 1945. Jour. Econ. Ent. (Sei. Note) 39: 387-391.

(231) MuLHERN, THOMAS D.

1953. BETTER RESULTS WITH MOSQUITO LIGHT TRAPS THROUGH STANDARDIZING IMECHANICAL PERFORMANCE. Mosquito NewS 13:

130-133.

(232) MULRENNAN, JOHN A.

1946. SOME NEW RECORDS OF FLORIDA MOSQUITOES. Fla. Anti-Mosquito. Assoc. Rpt., 17th Ann. Meeting, pp. 23-24.

L Mimeographed. ]

(233) NATIONAL MALARIA COMMITTEE, SUBCOMMITTEE ON ENGINEERING.

1936. MALARIA CONTROL FOR ENGINEERS. U. S. Pub. Health Serv.,

B-12Í0, 81 pp., illus. [Processed.]

(234) NELSON, D. B., and CHAMBERLAIN, R. W.

1955. A LIGHT TRAP AND MECHANICAL ASPIRATOR OPERATING ON DRY

CELL BATTERIES. Mosquito News 15: 28-32.

(235) NEWTON, W. L., and PRATT, I.

1946. EXPERIMENTS TO DETERMINE POTENTIAL MOSQUITO VECTORS OF

WUCHERERIA BANCROFTI IN THE CONTINENTAL UNITED STATES.

Amer. Jour. Trop. Med. 26: 699-706.

(236) WRIGHT, WILLARD H., and PRATT, IVAN.

1945. EXPERIMENTS TO DETERMINE POTENTIAL MOSQUITO VECTORS OF

WUCHERERIA BANCROFTI IN THE CONTINENTAL UNITED STATES.

Amer. Jour. Trop. Med. 25: 253-261.

(237) NIELSEN, ERIK TETENS, and HAEGER, JAMES S.

1953. SWARMING AND MATING OF MOSQUITOES. Fla. Anti-Mosquito

Assoc. Rpt. 24th Ann. Meeting, pp. 65-69. [Mimeographed.]

(238) and HAEGER, J. S.

1954. PUPATION AND EMERGENCE IN AEDES TAENIORHYNCHUS (WIED).

Bui. Ent. Res. 45: 757-768.

(239) and NIELSEN, ASTRID TETENS.

1953. FIELD OBSERVATIONS ON THE HABITS OF AEDES TAENIORHYNCHUS.

Ecology 34: 141-156.

(240) NoRRis, MARJORIE.

1946. RECOVERY OF A STRAIN OF EQUINE ENCEPHALITIS VIRUS FROM

CULEX RESTUANS (THEOB.) (DÍPTERA, CULICIDAE) . Canad.

Jour. Res., E, 24: 63-70.

(241) O'CONNOR, F. W., and HüLSE, CONSTANCE R.

1935. STUDIES IN FILARIASIS. Puerto Rico Jour. Pub. Health 11:

167-272.

(242) OWEN, WILLIAM B.

1942. THE BIOLOGY OF THEOBALDIA INORNATA WILLISTON, IN CAPTIVE

COLONY. Jour. Econ. Ent. 35: 903-907.

(243) PENNEBAKER, FAITH.

1938. ORTHOPODOMYIA ALBA IN LOUISIANA. Tulane Buggist 2:

4.

(244) PEREZ, MANUEL.

1930. AN ANOPHELINE SURVEY OF THE STATE OF MISSISSIPPI.

Amer. Jour. Hyg. 11: 696-710, illus.

(245) PETERSON, ALLAN G., and SMITH, WILLIAM W.

1945 OCCURRENCE AND DISTRIBUTION OF MOSQUITOES IN MISSISSIPPI.

Jour. Econ. Ent. 38: 378-383.

^246^ PHILEN, E. A., and CARMICHAEL, G. T.

^ ^ 1956. THE MANAGEMENT OF WATER FOR MOSQUITO CONTROL IN THE

COASTAL MARSHES OF FLORIDA. Mosquito News 16 : l¿b-l¿\).
170 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(247) PHILLIPS, J. H.

1939. STUDIES ON THE TRANSMISSION OF DIROFILARIA IMMITIS IN

MASSACHUSETTS. Amer. Jour. Hyg. 29: 121-129.

(248) PiSTEY, WARREN R.

1956. STUDIES ON THE MOSQUITO TRANSMISSION OF DIROFILARIA

TENUIS CHANDLER, 1942. Jour. Parasitol. 42 (Sect. 2) : 16.

(249) PRATT, HARRY D.

1945. MANSONIA INDUBITANS DYAR AND SHANNON—A NEW MOSQUITO ADDITION TO THE UNITED STATES FAUNA. KanS. Ent.

Soc. Jour. 18: 121-129.

(250) and SEABROOK, E. L.

1952. THE OCCURRENCE OF CULEX lOLAMBDIS DYAR IN FLORIDA AND

PUERTO RICO, WITH A DESCRIPTION OF THE LARVA (DÍPTERA,

CULICIDAE). Ent. Soc. Wash. Proc. 54: 27-32.

(251) WiRTH, W. W., and DENNING, D. G.

1945. THE OCCURRENCE OF CULEX OPISTHOPUS KOMP IN PUERTO RICO

AND FLORIDA, WITH A DESCRIPTION OF THE LARVA (DÍPTERA,

CULICIDAE). Ent. Soc. Wash. Proc. 47: 245-251.

(252) PRITCHARD, A. E., SEABROOK, E. L., and MULRENNAN, J. A.

1947. THE MOSQUITOES OF THE FLORIDA KEYS. Fla. Ent. 30: 8-15.

(253) SEABROOK, E. L., and PROVOST, M. W.

1946. THE POSSIBLE ENDEMICITY OF ANOPHELES ALBIMANUS IN

FLORIDA. Mosquito News 6: 183-184.

(254) PROVOST, MAURICE W.

1951. RELATIVE NUMBERS OF AEDES TAENIORHYNCHUS AND AEDES

SOLLICITANS IN FLORIDA DURING RECENT YEARS. Fla. AntiMosquito Assoc. Rpt., 22nd Ann. Meeting, Sect. 10, not

paged. [Mimeographed.]

(255)

(256)

(257)

(258)

1951. THE OCCURRENCE OF SALT MARSH MOSQUITOES IN THE INTERIOR OF FLORIDA. Fla. Ent. 34: 48-53.

1952. THE DISPERSAL OF AEDES TAENIORHYNCHUS. I.—PRELIMINARY

STUDIES. Mosquito News 12: 174-190, illus.

1952. GROUND SPACE SPRAYING OF MANSONIA AND OTHER MOSQUITOES

IN LEESBURG, FLORIDA. Mosquito News 12: 16-27, illus.

1955. LEESBURG MOSQUITO-SAMPLING STUDIES IN 1954. Fla. AntiMosquito Assoc. Rpt., 26th Ann. Meeting, pp. 112-119,

illus. [Mimeographed.]

(259) QuARTERMAN, K. D., and HESS, A. D.

1950. CURRENT INVESTIGATIONS ON MOSQUITO CONTROL. Fla. AntiMosquito Assoc. Rpt., 21st Ann. Meeting, pp. 49-52.

[Mimeographed.]

(260) QuiNBY, G. E.

1941. ADDITIONS TO THE MOSQUITOES (CULICIDAE) OF THE REELFOOT

LAKE REGION. Jour. Tenn. Acad. Sei. 16 (1): 17-21.

(261) REEVES, W. C, and HAMMON, W. MCD.

1944. FEEDING HABITS OF THE PROVEN AND POSSIBLE MOSQUITO

VECTORS OF WESTERN EQUINE AND ST. LOUIS ENCEPHALITIS

IN THE YAKIMA VALLEY, WASHINGTON. Amer. Jour. Trop.

Med. 24: 131-134.

(262) HAMMON, WILLIAM MCD., and IZUMI, ERNEST M.

1942. EXPERIMENTAL TRANSMISSION OF ST. LOUIS ENCEPHALITIS

VIRUS BY CULEX PIPIENS LINNEAUS. Soc. Expt. Biol. and

Med. Proc. 50: 125-128.

(263) REMINGTON, C. L.

1945. THE FEEDING HABITS OF URANOTAENIA LOWII THEOBALD (DIP-

,„^,, ^ TERA, CULICIDAE). Ent. News 56: 32-37; 64-68.

(264) RICHARDS, A. G. JR.

1941. A STENOGAMIC AUTOGENOUS STRAIN OF CULEX PIPIENS L. IN

NORTH AMERICA (DÍPTERA, CULICIDAE). Ent. NeWS 52:

211-216.

(265) RINGS, ROY W., and HILL, SAMUEL 0.

1946. THE LARVA OF AEDES (OCHLEROTATUS) MATHESONI MIDDLEKAUFF (DÍPTERA, CULICIDAE). Ent. Soc. Wash. Proc. 48:

237-240.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 171

(266) and HILL, SAMUEL 0.

1948. THE TAXONOMIC STATUS OF AEDES MATHESONI (DÍPTERA,

CULiciDAE). Ent. Soc. Wash. Proc. 50: 41-49, illus.

(267) ROCKEFELLER FOUNDATION, INTERNATIONAL HEALTH BOARD.

1924. THE USE OF FISH FOR MOSQUITO CONTROL. 120 pp., illus.

New York.

(267a) ROGERS, ANDREW J., and RATHBURN, CARLISLE B., JR.

1958. TESTS WITH A NEW GRANULAR PARIS GREEN FORMULATION

AGAINST AEDES, ANOPHELES, AND PSOROPHORA LARVAE. Mosquito News 18: 90-93. Also in Fla. Anti-Mosquito Assoc.

Rpt., 30th Ann. Meeting, pp. 28-31. 1959.

(268) ROOT, FRANCIS METCALF.

1924. NOTES ON THE MOSQUITOES OF LEE COUNTY, GEORGIA. Amer.

Jour. Hyg. 4: 449-455.

(269) Ross, EDWARD S.

1943. THE IDENTITY OF AEDES BIMACULATUS (COQUILLETT) AND A

NEW SUBSPECIES OF AEDES FULVUS (WIEDMANN) FROM THE

UNITED STATES (DÍPTERA, CULICIDAE). Ent. Soc. Wash.

Proc. 45: 143-151.

(270) Ross, HERBERT H.

1947. THE MOSQUITOES OF ILLINOIS (DÍPTERA, CULICIDAE). 111. Nat.

Hist. Surv. Bui. 24: 1-96.

(271) ROTH, LOUIS M.

1944. A KEY TO THE ANOPHELES OF THE SOUTHEASTERN UNITED

STATES, BY MALE GENITALIA (DíPTERA, CULICIDAE). AMER.

Mid. Nat. 31: 96-110.

(272)

(273)

1945. ABERRATIONS AND VARIATIONS IN ANOPHELINE LARVAE OF THE

SOUTHEASTERN UNITED STATES (DÍPTERA, CULICIDAE). Ent.

Soc. Wash. Proc. 47: 257-278.

1945. THE MALE AND LARVA OF PSOROPHORA (JANTHINOSOMA) HÓRRIDA (DYAR AND KNAB) AND A NEW SPECIES OF PSOROPHORA

FROM THE UNITED STATES (DÍPTERA, CULICIDAE). Ent. SoC.

Wash. Proc. 47: 1-23.

(274) and YOUNG, FRANK N.

1944. CULEX (MELANOCONION) ATRATUS THEOBALD IN FLORIDA; A

NEW CONTINENTAL NORTH AMERICAN RECORD, WITH NOTES

ON THE OTHER MELANOCONIONS OF THE SOUTHEASTERN

UNITED STATES. Ent. Soc. Amer. Ann. 37: 84-88.

(275) RozEBOOM, L. E.

1942. THE MOSQUITOES OF OKLAHOMA.—Okla. Agr. Expt. Sta.

Tech. Bui. No. T-16, 56 pp.

(276)

1951. THE CULEX PIPIENS COMPLEX IN NORTH AMERICA. In Mattingly et. al.. The Cidex pipiens Complex. Royal Ent. Soc,

London, Trans. 102: 343-353, illus.

(277) and GILFORD, BARBARA N.

1954. SEXUAL ISOLATION BETWEEN POPULATIONS OF THE CULEX PIPIENS COMPLEX IN NORTH AMERICA. Jour. Parasitol. 40:

237-244.

(278) and KOMP, WILLIAM H. W.

1950. A REVIEW OF THE SPECIES OF CULEX OF THE SUBGENUS MELANOCONION (DÍPTERA, CULICIDAE). Ent. Soc. Amer. Ann.

43: 75-114, illus.

(279) RUSSELL, P. F.

1925. IDENTIFICATION OF THE LARVAE OF THE THREE COMMON ANOPHELINE MOSQUITOES OF THE SOUTHERN UNITED STATES. Amer.

Jour. Hyg. 5: 149-174.

(280) SABROSKY, C. W., MCDANIEL, G. E., and REIDER, R- F.

1946. A HIGH RATE OF NATURAL PLASMODIUM INFECTION IN ANOPHELES CRUCIANS. Science 104: 247-248.

/'9S1 ^ ^rwooF H F.. and ASHTON, D. F. (281) ^CHOOF, n , ^^^ DISTRIBUTION RECORDS ON THE MOSQUITOES

OF NORTH CAROLINA. Elisha Mitchell Sci. Soc. Jour. 60:

1-10.

(282) SCHWARDT^H^H^ ^^ ARKANSAS RICE FIELD MOSQUITOES. Ark. Agr.

Expt. sta. Bui. 377, pp. 1-22.
172 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(283) SEA^BROOK,^EDwm L.^ ^^ MANSONIA MOSQUITOES IN SOUTHEAST

FLORIDA. Newell Ent. Soc. Jour. 4: 27-30.

1946. THE OCCURRENCE OF MEGARHINUS RUTILUS COQUILLETT IN S. E.

FLORIDA. Mosquito News 6: 193-194.

(285) SEAMAN.^ELWOOD A.^^^^ ^^^^ ^^^ ^^^ ^^^^ ^^^^ ^^^^^^^^ ^^ ,„,

FIELD. Mosquito News 5: 79-81, illus.

(286) SHEPPARD, RAYMOND R. ^^^^^ ^.

1949. THE MOSQUITOES OF THE SOUTHWEST COAST OF FLORIDA. Ma.

Anti-Mosquito Assoc. Rpt., 20th Ann. Meeting, pp. 122-

126. [Mimeographed.]

(287) SHIELDS, S. E. T TP T?r.f 1938 TENNESSEE VALLEY MOSQUITO COLLECTIONS. Jour. Econ. J^nt.

31: 426-430.

(288) and LACKEY, JAMES B.

1938 CONDITIONS AFFECTING MOSQUITO BREEDING, WITH SPECIAL REFERENCE TO AEDES THIBAULTI DYAR AND KNAB (DÍPTERA,

CULICIDAE). Jour. Econ. Ent. 31: 95-102, illus.

(289) and MiLES, ViRGiL I.

1937. THE OCCURRENCE OF ORTHOPODOMYIA ALBA IN ALABAMA (DÍPTERA, CULICIDAE). Ent. Soc. Wash. Proc. 39: 237.

(290) SHLAIFER, A., and HARDING, D. E. ^ O. • T O-.

1946 THE MOSQUITOES OF TENNESSEE. Tenn. Acad. Sci. Jour. ¿1:

241-256.

(291) SHUTE, P. G. ^^^ ^^^

1951. CULEX MOLESTUS. Royal Ent. Soc, London, Trans. 102: 380-

382. (In Mattingly et al., réf. No. 218.)

(292) SMITH, JOHN B.

1904. REPORT OF THE NEW JERSEY STATE AGRICULTURAL EXPERIMENT

STATION UPON THE MOSQUITOES OCCURRING WITHIN THE

STATE, THEIR HABITS, LIFE HISTORY, ETC. 482 pp., illus.

Trenton.

(293) SNOW, W. E.

1949. STUDIES ON PORTABLE RESTING STATIONS FOR ANOPHELES

QUADRIMACULATUS IN THE TENNESSEE VALLEY. Nati. Malaria Soc. Jour. 8: 336-343, illus.

(294) and PICKARD, EUGENE.

1956. SEASONAL HISTORY OF CULEX TARSALIS AND ASSOCIATED SPECIES

IN LARVAL HABITATS OF THE TENNESSEE VALLEY REGION.

Mosquito News 16: 143-148.

(295) STAEBLER, ARTHUR E., and BUREN, WILLIAM F.

1946. AEDES TORTILIS (THEOBALD), A MOSQUITO NEW TO THE UNITED

STATES. U. S. Pub. Health Serv. Rpts. 61: 685.

(296) STAGE, H. H., GJULLIN, C. M., and YATES, W. W.

1937. FLIGHT RANGE AND LONGEVITY OF FLOODWATER MOSQUITOES IN

THE LOWER COLUMBIA RIVER VALLEY. Jour. Econ. Ent. 30:

940-945.

(297) STONE, ALAN.

1953. THE HALTERES OF ANOPHELES WALKERI THEOBALD. Mosquito

News 13: 209-210.

(298)

1957. CORRECTIONS IN THE TAXONOMY AND NOMENCLATURE OF MOSQUITOES. Ent. Soc. Wash. Proc. [1956] 58: 333-344.

(298a) KNIGHT, K. L., and STARCKE, H.

1959. A SYNOPTIC CATALOGUE OF THE MOSQUITOES OF THE WORLD.

Ent. SOC. Amer. (The Thomas Say Foundation.) 358 pp.

(299) STOUGH, B. DOLORES, KING, MARY ANN, and EYLES, DON E.

1949. EXTENSION OF KNOWN RANGE OF AEDES ATROPALPUS (COQUILV

LET) SOUTHEASTWARD INTO GEORGIA. Mosquito News 9:

173.

(300) STRATMAN-THOMAS, W. K., and BAKER, F. C.

1936. ANOPHELES BARBERI COQUILLET, AS A A^CTOR OF PLASMODIUM

viVAX GRASSI AND FELETTI. Amer. Jour. Hyg. 24: 182-183.

(301) STROMQUIST, W. G.

1935. MALARIA CONTROL IN THE TENNESSEE VALLEY. Civ. Engin.

5: 771-774, illus.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 173

(302) SUNDARARAMAN, S.

1949. BIOMETRICAL STUDIES ON INTERGRADATION IN THE GENITALIA

OF CERTAIN POPULATIONS OF CULEX PIPIENS AND CULEX QUINQUEFASCIATUS IN THE UNITED STATES. Amer. Jour. Hyg.

50: 307-314.

(303) THIBAULT, JAMES K., JR.

1910. NOTES ON THE MOSQUITOES OF ARKANSAS [DÍPTERA, CULICIDAE].

Ent. Soc. Wash. Proc. 12: 13-26.

(304) THOMPSON, GEORGE A., HOWITT, BEATRICE F., GORRIE, RACHAEL, and

COCKBURN, T. AlDAN.

1951. ENCEPHALITIS IN MIDWEST. VI.—WESTERN EQUINE ENCEPHALOMYELITIS VIRUS ISOLATED FROM AEDES DORSALIS MEIGEN.

Soc. Expt. Biol. & Med. Proc. 78: 289-290.

(305) THURMAN, ERNESTINE BASHAM, HAEGER, JAMES S., and MULRENNAN,

JOHN A.

1949. THE OCCURRENCE OF AEDES (OCHLEROTATUS) THELCTER DYAR IN

THE FLORIDA KEYS. Mosquito News 9: 171-172.

(306) HAEGER, JAMES S., and MULRENNAN, JOHN A.

1951. THE TAXONOMY AND BIOLOGY OF PSOROPHORA ( JANTHINOSOMA)

JOHNSTONII (GRABHAM, 1905) (DÍPTERA: CULICIDAE). Ent.

SOC. Amer. Ann. 44: 144-157.

(307) TRAVIS, B. V., and MORTON, F. A.

1946. TREATMENT OF CLOTHING FOR PROTECTION AGAINST MOSQUITOES.

N. J. Mosquito E.Ntermin. Assoc. Proc. 33: 65-69.

(308) MORTON, FRED A., and COCHRAN, J. H.

1946. INSECT REPELLENTS USED AS SKIN TREATMENTS BY THE ARMED

FORCES. Jour. Econ. Ent. 39: 627-630.

(309) and SMITH, CARROLL N.

1951. INIOSQUITO REPELLENTS SELECTED FOR USE ON MAN. Jour.

Econ. Ent. (Sei. Note) 44: 428-429.

(310) UNITED STATES DEPARTMENT OF AGRICULTURE, ENTOMOLOGY RESEARCH BRANCH.

1955. INSECTICIDES AND REPELLENTS FOR THE CONTROL OF INSECTS

OF MEDICAL IMPORTANCE TO THE ARMED FORCES. U. S.

Dept. Agr. Cir. No. 977, 91 pp., illus.

(311) UNITED STATES PUBLIC HEALTH SERVICE.

1951. MOSQUITO RECORDS FROM THE MISSOURI RIVER BASIN STATES.

Survey Section, CDC. 93 pp. [Mimeographed.]

(312) VAN DINE, D. L.

1922. IMPOUNDING WATER IN A BAYOU TO CONTROL BREEDING OF MALARIA MOSQUITOES. U. S. Dept. Agr. Bui. 1098, 22 pp.,

illus.

(313) VANNOTE, ROBERT L.

1937. METHOD OF APPLYING THE LARVICIDE AS A REPELLENT. N. J.

Mosquito Extermin. Assoc. Proc. 24: 11-15.

(314) VoGT, GEORGE B.

1947. SALINITY TOLERANCE OF ANOPHELES QUADRIMACULATUS AND

HABITAT PREFERENCE OF A. CRUCIANS BRADLEYI. Jour.

Econ. Ent. 40: 320-325.

(315) WALLIS, ROBERT C.

1954. NOTES ON THE BIOLOGY OF CULISETA MELANURA (COQUILLETT) .

Mosquito News 14: 33-34.

(316) WANAMAKER, JOHN F.

1944. AN IMPROVED METHOD FOR MOUNTING MOSQUITO LARVAE.

Amer. Jour. Trop. Med. 24: 385-386.

(317) CHAMBERLAIN, ROY W., and CARPENTER, STANLEY J.

1944 DISTRIBUTION OF CULEX PIPIENS IN THE SOUTHEASTERN UNITED

STATES. Jour. Econ. Ent. (Sei. Note) 37: 106-107.

(318) WATSON, R. B., KIKER, C C, and JOHNSON, H. A.

1938. THE ROLE OF AIRPLANE DUSTING IN THE CONTROL OF ANOPHELES

BREEDING ASSOCIATED WITH IMPOUNDED WATERS. U. S. Pub.

Health Serv. Rpts. 53: 251-263, illus.

^319) and SPAIN, E. L., JR.

1937 STUDIES ON MALARIA IN THE TENNESSEE VALLEY. THE INFLUENCE OF PHYSIOGRAPHY ON THE OCCURRENCE OF BREEDING

PLACES OF ANOPHELES QUADRIMACULATUS IN NORTHERN ALABAMA. Amer. Jour. Trop. Med. 17: 289-305, illus.
(332)

174 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

(320) WEATHERSBEE, ALBERT A., and ARNOLD, F. T., JR. ^^^^^^^^. m_^

1947 A RESUME OF THE MOSQUITOES OF SOUTH CAROLINA. Tenn.

Acad. Sei. Jour. 22: 210-229.

^^^^^ ^L948 T'^REV^IXN^ ii ra WYEOMYIA THEOBALD (DíPTERA,

CULICIDAE) IN SOUTH CAROLINA. Tenn. Acad. Sci. Jour.

23: 249-250.

1939.' INSECTS FOUND ON AIRCRAFT AT MIAMI, FLA., IN 1938. U. S.

Pub. Health Serv. Rpts. 54: 561-566.

^^^^^ 1945 THE FINDING OF ANOPHELES ALBIMANUS WIED. AT WEST PALM

BEACH, FLORIDA, (DÍPTERA, CULICIDAE). Mosquito NeWS 5:

145.

HITEHl^ D^,^ PREFERENCE OF PSOROPHORA CONFINNIS AND P. DISCOLOR.

Jour. Econ. Ent. 44: 1019.

(325) WiLKiNS, ORíN P., and BRELAND, OSMOND P.

1949. RECOVERY OF THE MOSQUITO, CULISETA INORNATA (WILLISTON)

FROM DRY MATERIAL. Ent. Soc. Wash. Proc. 51: 27-28.

(326) and BRELAND, OSMOND P.

1951. THE LARVAL STAGES AND THE BIOLOGY OF THE MOSQUITO ORTHOPODOMYIA ALBA BAKER (DÍPTERA, CULICIDAE). N. Y.

Ent. Soc. Jour. 59: 225-240.

(327) WILLIAMS, LOUIS L., JR.

1941. THE ANTI-MALARIA PROGRAM IN NORTH AMERICA. In Human

Malaria, Amer. Assoc. Adv. Sei. Pub. 15, pp. 365-370, illus.

(328) and COOK, S. S.

1927. PARIS GREEN APPLIED BY AIRPLANE IN THE CONTROL OF ANOPHELES PRODUCTION. U.S. Pub. Health Serv. Rpts. 42: 459-

480, illus.

(329) and LEGARE, A. E.

1928. LENGTH OF LIFE OF ANOPHELES QUADRIMACULATUS AFTER BEGINNING OF CONTROL OF PRODUCTION. South. Med. Jour. 21 :

735-737, illus.

(330) WiRTH, WILLIS W.

1944. NOTES ON THE OCCURRENCE OF ANOPHELES GEORGIANUS KING

IN LOUISIANA. Jour. Econ. Ent. (Sei. Note) 37: 446.

(331)

1945. THE OCCURRENCE OF CULEX (MELANOCONION) ELEVATOR DYAR

AND KNAB IN FLORIDA, WITH KEYS TO THE MELANOCONIONS

OF THE UNITED STATES (DÍPTERA, CULICIDAE). Ent. Soc.

Wash. Proc. 47: 199-210, illus.

1947. NOTES ON THE MOSQUITOES OF LOUISIANA. Jour. Econ. Ent.

(Sei. Note) 40: 742-744.

(333) WRAY, FRANKLIN C.

1946. SIX GENERATIONS OF CULEX PIPIENS V^ITHOUT A BLOOD MEAL.

Mosquito News 6: 71-72, illus.

(334) YAMAGUTI, SATYU.

1952. ILLUSTRATED KEYS TO THE ADULT CULICINE MOSQUITOES OF

AMERICA NORTH OF MEXICO WITH NOTES ON GENERAL MORPHOLOGY AND BIOLOGY OF GENERA. 48 pp., illus.

(335) and LACASSE, WALTER J.

1951. MOSQUITO FAUNA OF NORTH AMERICA. Parts I-V. Office

of the Surgeon, Hq., Japan Logistical Command. 629 pp.

(336) YEN, CHLA.-HSIEN.

1938. STUDIES ON DIROFILARIA IMMITIS LEIDY WITH SPECIAL REFERENCE TO THE SUSCEPTIBILITY OF SOME MINNESOTA SPECIES

OF MOSQUITOES TO THE INFECTION. Jour. Parasitol. 24:

189-205.

(337) YOUNG, FRANK N., JR., and CHRISTOPHER, WARREN N.

1944. UNUSUAL BREEDING PLACES OF MOSQUITOES IN THE VICINITY

OF KEESLER FIELD, MISSISSIPPI. Amer. Jour. Trop. Med.

24: 379.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 175
176 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

II & A 14068. M & A 14069

Plate 2.—A, A seasonal bayou overgrown with willows. B, A portion of the

same bayou after clearing and impounding to eliminate Anopheles breeding.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 177

M í¡ A 14070, M Ji A 14371

Plate 3.—A, Mist blower mounted on a two-wheel ti-ailer. B, A microsol

machine mounted in a jeep.
178 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

M & A 14073

Plate 4.—A, Thermal fog generator mounted in a jeep. B, PT-17 plane

equipped with underwing spray booms.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 179

M & A 14074, M Ä A 8432

Plate 5.—A, Anopheles breeding pool in a cypress swamp. B, Water conditions favorable for breeding of Anopheles quadrimaculatus.
180 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

pSfee^'^i**^"

im im» Ü. r<

M i A 9815. M & A 14076

Plate 6.—A, A prolific breeding- place of Aedes tacniorhynchus in a pickleweed (Datis inaritinia) marsh. D, Red mangrove {Rhizophora mangle) in a

Florida marsh, showin,a- the dense .growth and characteristically divided base.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 181

M ii A 11472, M & A 9742

Plate 7.—A, A 30-inch main ditch and a 10-inch lateral in a Jnuctis marsh in

North Carolina; B, A 30-inch ditch in a pickleweed {Batis) marsh in

Florida.
182 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

M & A 14075,

Plate 8.—Wings of the typically Nearctic species of Anopheles: A, crucians;

B, freeborni; C, punctipcvnis; D, pseudopunctipennis: E, walkeri; F,

átropos; G, quadrimaculatus; H, harheri.
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 183

M & A 9307, M & A 9614

Plate 9.—A, A pupa of Mansonia perturbons attached to the cut stem of an

aquatic plant. B, A breeding place of M. perturbans, a large shallow pond

filled with pickerelweed (Povtederia).
184 AGR. HANDBOOK 173, U. S. DEFT. OF AGRICULTURE

Plate 10.—A water-holding air plsnt (bromeliad) which serves as a breeding

place for Wyeoinyict initchellii and W. vandnzeci.
INDEX

(Valid names are in roman type, synonyms in italics. Figures in bold-faced

type indicate page on which the genus or species is described.)

Page

ahominator Auct., Cidex 110

Aedes Meigen 9

11, 13, 18, 21, 22, 27, 31, 44, 45^

54, 55, 56, 59, 60, 66, 69, 82, 103,

134, 137, 140, 141, 142.

Aedimorphus Theob., subgenus . . 60

aegypti (L.), Aedes 1,

2, 18, 19, 20, 21, 22, 26, 41, 42,

55, 61, 63, 64, 65.

alba Baker, Orthopodomyia 4,

135, 136, 137, 151

albimanus Wied., Anopheles .... 2,

6, 17, 21, 85, 87, 89, 90, 91, 93,

109, 146.

albipes Theob., AnojDheles 91

albopictus (Skuse), Aedes 19

American Cyanamid 12008 32

annulimaniis V. d. W., Anopheles 99

Anopheles Meigen 7,

9, 10, 11, 18, 21, 22, 25, 27, 33,

41, 49, 54, 55, 58, 85, 87, 89, 95,

96, 123, 149.

Anophelinae 49, 55

apicalis Auct., Ciilex 55

67, 81, 120, 121, 123, 125, 153

Aquatic plants inimical to mosquitoes 33

argenteiis Poir., Culex 64

atlanticus D. and K., Aedes .... 2,

6, 22, 62, 64, 66, 70, 81

atratus Theob., Culex 3,

104, 105, 107, 109, 110

atratus Dyar, Melanoconion .... 110

atropalpus (Coq.), Aedes 2,

20, 22, 61, 63, 66, 67

átropos D. and K., Anopheles .... 2,

85, 90, 92, 93, 102, 103

Azolla 33

bahamensis D. and K., Culex .... 3,

104, 106, 108, 109, 111

bancrofti, Wuchereria 21, 22

barberi (ííoq.. Anopheles 2,

89, 90, 92, 102, 136, 151

Bayer 29493 (Baytex) 32, 35, 36

Benzene hexachloride (BHC) .... 31,

34, 35, 36

bimaculatus (Coq.), Aedes 70

Bironella 87, 89

Bladderworts (Utricularia) 33

bradleyi King, Anopheles 3,

85, 88, 89, 90, 92, 93, 94, 95, 103

bradleyi King, Anopheles crucians 3, 93

Butyl-ethyl-propanediol 39

calopus Meig., Culex 64

Page

canadensis, Aedes 6,

22, 61, 68, 71

canadensis canadensis (Theob.),

Aedes 2,

6, 61, 63, 67

canadensis mathesoni Middlekauf,

Aedes 2,

6, 61, 63, 68

cancer Theob., Deinocerites 4

80, 110, 113, 127

Chagasia 89

Chara (C. fragilis) 33

Chlordane 31, 34

Chlorinated hydrocarbons

adulticides 35

larvicides 29, 30, 31

resistance 48

ciliata (F.), Psorophora 4, 6,

21, 22, 137, 138, 139, 141, 143,

144, 145.

cinereus Meig., Aedes 2,

22, 54, 62, 63, 68, 69

Climacura, subgenus 124

columbiae (D. and K.), Psorophora 140

Common malaria mosquito (see

quadrimaculatus. Anopheles)

confinnis (L.-Arr.), Psorophora .. 4,

6, 21, 72, 84, 133, 137, 138, 139,

140, 141, 142, 143, 145.

continentalis D. and K., Urano- ....

taenia 152

Coquillettidia Dyar 127, 129

Co-Ral 30, 32

corniger Dyar, Cidex 109

crucians Wied., Anopheles ....20, 21,

22, 88, 89, 90, 93, 94, 95, 96, 146

Culex Linnaeus 11,

13, 18, 21, 22, 27, 30, 31, 33, 52,

54, 56, 58, 59, 103, 104, 105, 106,

107, 108, 109, 112, 113, 118, 123,

124, 125, 126, 154.

Culicidae 49

Culicinae 49, 55, 59

Culicoides 44

Culiseta Felt 54,

58, 59, 106, 124, 125

cyanescens (Coq.), Psorophora .. 4,

6, 22, 54, 134, 138, 139, 142

DDT 18,

29, 30, 31, 32, 33, 34, 35, 36, 37,

40, 41, 48.

Deinocerites Theob 54,

56, 59, 126, 127

Dengue fever 1,

17, 19, 23, 41, 65

185
186 AGR. HANDBOOK 173, U. S. DEPT. OF AGRICULTURE

Page

Dermatobia 1'7

Diazinon 35

Dibrom 36

Dicapthon 32

Dieldrin 31, 34

Diethyltoluamide 38, 39

Diking 44

Dimethyl carbate 38

Dimethyl phthalate 38

Dipterex 30, 32

Dirofilaria immitis 22,

83, 84, 99, 101

Dirofilaria tenuis 79

discolor (Coq.), Psorophora .... 4,

21, 137, 138, 139, 141, 143

Ditching 28,

33, 40, 42, 43, 44, 45, 48

dorsalis (Meig.), Aedcs 2,

20, 32, 61, 64, 69

Duckweed (Lemna) 33

dupreei (Coq.), Aedes 2,

62, 64, 6Q, 70

egherti D. and K., Culex 110

elevator Auct., Culex 110, 111

Encephalitis 6,

17, 20, 21, 72, 74, 79, 83, 84, 95,

117, 122, 126.

EPN 30 32

Eretmapodites 18

erraticus D. and K., Culex 3,

13, 21, 22, 104, 107, 108, 110,

111, 113, 114, 142, 153.

Ethyl hexanediol 33

eiiplocamus Auct., Aedes 73

falciparum, Plasmodium .... 17, 93, 98

fasciatus F., Stegomyia 64

fatigans Wied., Culex 22,

115, 118, 119

ferox (Humb.), Psorophora .... 4, 6,

21, 22, 67, 83, 137, 139, 144, 148

Filariasis (ñlarial worms, filarla,

Wuchereria; also see Dirofil?.-

ria, heartworm) 1,

17, 21, 22, 117, 118

Finlaya Theob., subgenus 60

Florida glades mosquito 6, 140

floridanus D. and K., Mochlostyrax 114

fioridense D. and K., Janthinosoma 140

franciscanus McC, Anopheles ... 98

freeborni Aitken, Anopheles .... 17

Fowl pox 17,

22, 65, 84, 117

fulvus Dyar, Aedes (sp. pallens) . . 70

fuscus O. S., Aedes 68

Gambusia 32

georgianus King, Anopheles .... 3,

17, 88, 89, 90, 93, 94, 95, 96

georgianus King, Anopheles crucians 95

Grabhamia Theob., subgenus .... 56,

59, 137, 138, 143, 147, 148

grossbecki D. and K., Aedes .... 2,

61, 63, 71

Page

Haemogogus ^^

haynei Dodge, Wyeomyia 5,

6, 154, 155, 156

Heartworm • • • }Jf

22, 101, 117

Heptachlor 31

Hercules AC-5727 35

hirsuteron (Theob.), Aedes 75

hórrida (D. and K.), Psorophora 4,

137, 139, 144, 146, 147

howardii (Coq.), Psorophora 4,

21, 137, 138, 139, 140, 141, 145

hyemails (Fitch), Culex 98

incriminator D. and K., Culex . . . 114

indubitans D. and S., Mansonia . . 4,

6, 21, 54, 56, 128, 129, 130, 131,

132, 133, 134.

infirmatus D. and K., Aedes 2,

6, 20, 62, 64, 66, 72, 73, 74, 75

inhibitator Auct., Culex HO

inornata (Will.), Culiseta 3,

20, 21, 22, 71, 106, 111, 124, 125

iolambdis Dyar, Culex 3,

104, 107, 108, 109, 110, 111

jamaicensis Dyar, Culex 140

Janthinosoma L.-Arr., subgenus .. 137

138, 142, 144, 146, 148

johnstonii (Grabham), Psorophora 4,

80, 137, 138, 139, 146

Kerosene 30, 41

keyensis Buren, Aedes 79, 80

Leishmaniasis 17

leprincei D. and K., Culex 110

Limatus 18

Lindane 31, 34, 35

longipalpis Roth., Psorophora ... 4,

6, 137, 139, 146

lowii Theob., Uranotaenia 5,

109, 113, 152, 153

maculipennis Meig., Anopheles . . 22

Malaria 1,

7, 17, 18, 23, 27, 33, 34, 40, 41, 85,

88, 91, 92, 93, 95, 96, 99, 100, 102

malariae, Plasmodium 17

Malathion 30,

32, 35, 36, 37

malayi, Wuchereria 21

Mansonia Blanchard 13,

18, 21, 56, 59, 127, 128, 129, 131

mathesoni Middlekauf, Aedes CSLTIadensis 2, 6, 61, 63, 68

Megarhinus, Rob.-Desv 49,

55, 149

Melanoconion Theob., subgenus .. 104,

105, 106, 107, 108, 109, 110, 111,

112, 113, 114, 123.

melanura (Coq.), Culiseta 3,

20, 81, 103, 106, 124, 125, 126

mississippii Dyar, Aedes Stimulans 76

mitchellae (Dyar), Aedes 2,

20, 61, 64, 72, 73, 75, 79
THE MOSQUITOES OF THE SOUTHEASTERN UNITED STATES 187

Page

mitchellii (Theob.), Wyeomyia ... 5,

150, 154, 155, 156

Mochlostyrax Foote, subgenus . . . 105

molestus Forskâl, Culex 115,

116, 117

mulrennani Basham, Culex 3,

6, 104, 107, 108, 109, 112

musicus Say, Culex 144

Myiasis 17

nanus Coq., Culex 147

Neoculex Dyar, subgenus 104,

105, 106, 107, 108, 123

niger Giles, Taeniorhynchus . . 76, 79

nigripalpus Theob., Culex 3,

21, 26, 104, 106, 107, 108, 112, 113

nigromaculis (Ludl.), Aedes ... 32, 36

Nyssorhynchus Blanch 85

Nyssorhynchus Blanch., subgenus 89

Ochlerotatus L.-Arr., subgenus . . 60

Oils (as larvicides) 29,

30, 32, 35, 40

opisthopus Komp, Culex 3,

104, 106, 107, 109, 113, 127

Orthopodomyia Theob 56,

59, 134

ovale, Plasmodium 17

pallens Ross, Aedes fulvus 2,

61, 64, 70, 71

pallens Coq., Culex pipiens 115

Parathion 30, 32, 35

Paris green 29, 32, 37, 40

peccator D. and K., Culex 3,

104, 107, 108, 114

perplexens Ludlow, Anopheles ... 3,

6, 17, 88, 89, 91, 96, 97, 98, 99

perturbans (Wlk.), Mansonia .... 4,

6, 20, 21, 22, 127, 129, 130, 131,

132, 133.

Phosdrin 32

pilosus (D. and K.), Culex 3,

104, 105, 107, 108, 114, 115

pipiens L., Culex 3,

55, 115, 116, 117, 118, 119

pipiens complex, Culex 115, 119

pipiens pipiens L., Culex 20,

21, 22, 84, 104, 106, 107, 115,

116, 117, 120.

pipiens quinquefasciatus Say, Culex (see quinquefasciatus)

Plasmodium 17, 18, 65

portoricensis (Ludl.), Culex .. 76, 79

pose D. and K., Culex 110

posticatus Wied., Culex 144

pseudopunctipennis Theob.,

Anopheles 3,

89, 90, 98, 99

Psorophora Rob.-Desv 9,

11, 13, 18, 27, 54, 56, 59, 60,

m, 80, 103, 110, 133, 137, 138,

140, 141, 142.

punctipennis (Say), Anopheles .. 3, 6,

22, 88, 89, 90, 94, 96, 97, 98, 99,

110, 120, 121, 122, 123, 125, 153

pungens Wied., Culex 118

Page

pygmaea (Theob.), Psorophora .. 4,

137, 138, 139, 147

Pyrethrum 29,

32, 34, 35, 41

quadrimaculatus Say, Anopheles 1,

3, 11, 17, 21, 22, 26, 30, 34, 35,

37, 39, 40, 85, 88, 89, 90, 92, 94,

95, 99, 100, 101, 102, 103, 110,

141, 142, 153.

quinquefasciatus Say, Culex pipiens 1,

3, 21, 22, 26, 41, 42, 55, 104,

106, 107, 108, 115, 116 117, 118,

119, 120, 121.

restuans Theob., Culex 3,

20, 21, 22, 67, 71, 81, 103, 104,

106, 107, 108, 117, 119, 120, 121,

122, 123, 124, 125.

Ronnel 32

rutilus rutilus (Coq.), Toxorhynchites 4,

149, 150, 151

rutilus septentrionalis (D. and

K.), Toxorhynchites 4,

135, 140, 149, 150

Sabethes 18

salinarius Coq., Culex 3,

6, 21, 22, 104, 106, 107, 108, 112,

113, 117, 119, 120, 121, 122.

Salt-marsh mosquitoes 6,

27, 29, 31, 32, 35, 36, 42, 43, 44,

48, 74, 76, 78.

Sand fly (see Culicoides)

sapphirina (O.-S.) Uranotaenia . . 5,

152, 153

saxatilis Gros., Culex 123

sayi D. and K., Janthinosoma .... 144

scapularis (Rondani), Aedes .... 2,

62, 64, 72, 73, 74

scutellaris (Wlk)., Aedes 19

septentrionalis (D. and K.), Toxorhynchites rutilus 4,

135, 140, 149, 150

sei^ratus Coq., Ochlerotatus Q>Qf

signifera (Coq.), Orthopodomyia 4,

21, 93, 135, 136, 150, 151

signipennis (Coq.), Psorophora .. 4,

134, 138, 139, 143, 148

similis Theob., Culex 112

smithii Auct., Wyeomyia 6, 154

socialis Theob., Uranotaenia .... 153

sollicitans (Wlk.), Aedes 2,

6, 11, 21, 22, 42, 61, 64, 72, 73,

74, 75, 76, 77, 78, 79, 80, 84.

Southern house mosquito 1,

11, 41, 42, 118

spegazzinii Brethes, Haemogogus 18

Stegomyia Theob., subgenus 19,

55, 60

sticticus (Meig.), Aedes 2,

21, 62, 63, 67, 75

Stimulans (Wlk.), Aedes 2,

22, 61, 63, 71, 76
188 AGR. HANDBOOK 173, U.

J'agc

Taeiiio)Jiij}i('lifis L.-Arr 127

taeniorhynchus (Wied.), Aedes .. 2,

6, 21, 22, 42, 44, 61, 64, 74, 76,

78, 79, 80, 110, 111.

tarsalis Coq., Culex 3,

20, 21, 22, 29, 32, 36, 67, 104,

106, 108, 121, 122.

territans Wlk., Culex 3,

22, 55, 67, 81, 104, 106, 107, 108,

110, 114, 122, 123, 125, 153.

fcrritans Auct., Culex 120

thelcter Dyar, Aedes 2,

62, 64, 66, 79, 80

Theohalclia N.-L 124

thibaulti D. and K., Aedes 2,

22, 62, 63, 71, 80, 81, 120, 125

Thiocyanate insecticides 34

titillans (Wlk.), Mansonia 4,

54, 56, 128, 129, 130, 133, 134

tormentor D. and K., Aedes 2,

22, 62, 64, 66, 70, 81

tortilis (Theob.), Aedes 2

62, 64, 80, 82

Toxaphene 31

Toxorhynchites Theob 11

49, 54, 55, 58, 93, 136, 149

Toxorhynchitinae 49,

55, 58, 149

i^ U.S.

S. DEPT. OF AGRICULTURE

Page

triseriatus (Say), Aedes 2,

6, 21, 22, 62, 63, 66, 81, 82, 83,

93, 126, 135, 136, 150, 151.

trivittatus (Coq.), Aedes 2,

22, 62, 64, 83

Tularemia 17

Uranotaenia L.-Arr 13,

54, 55, 59, 151, 152

vanduzeei D. and K., Wyeomyia . . 5,

150, 154, 156

varipes (Coq.), Psorophora 4,

6, 137, 138, 139, 144, 146, 148

vexans (Meig.), Aedes 2,

6, 21, 22, 61, 63, 67, 69, 75, 83,

84, 120, 141, 142, 143, 144.

vivax, Plasmodium 17, 98

walkeri Theob., Anopheles 3,

22, 89, 90, 92, 102

Water hyacinth (Piaropus) 33

Wolffia 33

Wuchereria 22

Wyeomyia Theob 13,

18, 54, 56, 59, 153

Yellow fever 1

17, 18, 19, 65, 73

Yellow-fever mosquito 1,

11, 41, 64

GOVERNMENT PRINTING OFFICE: 1960—527552