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U. s. DEPARTMENT OF AGRICULTURE • AGRICULTURE HANDBOOK NO. 152
AGRICULTURE HANDBOOK NO. 152

AGRICULTURAL RESEARCH SERVICE

U. S. DEPARTMENT OF AGRICULTURE

Washington, D. C. Issued July 1959

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

Washington 25, D. C. - Price $2.50 (Buckram)
Acknowledgments

This study has been made possible by a transfer of funds from the

United States Department of the Army to the Entomology Research

Division, Agricultural Research Service.

Assistance during the course of the study was received in many ways.

We are grateful to the following for their valuable contributions: Kathryn

M. Sommerman, Curtis W. Sabrosky, and Alan Stone, entomologists with

the Federal insect identification unit of the former Bureau of Entomology

and Plant Quarantine, for the initial planning and development of the

pictorial keys and assistance in the preparation of the text ; Lt. Col. F. W.

Whittemore, Medical Service Corps, United States Army, for his encouragement and constructive criticism in the early stages of the project ; and

Mrs. Sally Kaicher, of the former Bureau, for skillfully designing the

individual pictorial keys.

The assembling and analysis of information pertinent to this project

were materially expedited by a related project on arthropods of medical

importance being prepared for the Office of the Quartermaster General,

United States Department of the Army, under the supervision of B. V.

Travis, Department of Entomology, Cornell University, and by suggestions and criticisms of many individuals in the United States Department

of Defense, the United States Public Health Service, and medical and

scientific organizations throughout the world.

Contents

Page

Mosquito-transmitted diseases 2

Malaria 2

Yellow fever 2

Dengue 3

The encephalitides 3

Encephalitis viruses __- 3

Other mosquito-borne viruses. 4

Filariasis 5

Geographical distribution and relative

medical importance of diseasetransmitting mosquitoes, based on

pictorial keys 5

Directions for use of the keys 8

North America 8

America north of Mexico 8

Middle America 11

West Indies . - 11

Mexico 14

Central America 17

Panama 20

South America 23

Northwest South America 23

Brazil and the Guianas 26

Lower South America 30

Africa 32

Northwest Africa 32

Egypt 35

Page

Geographical distribution and relative

medical importance of diseasetransmitting mosquitoes—Continued

Africa—Continued

Anglo-Egyptian Sudan 37

Eritrea, the Somalilands, and

Ethiopia 40

West Africa 43

West-Central Africa 47

East-Central Africa 50

Madagascar and adjacent

islands 53

Southern Africa 56

Eurasia 60

North Europe and North Asia 60

Spain and Portugal 64

Italy, the Balkans, and adjacent islands 66

The Near East 68

Arabia 70

Iran 73

Afghanistan and West Pakistan 75

India and Ceylon 78

East Pakistan and Assam 80

South China 82

Formosa 85

Page

Geographical distribution and relative

medical importance of diseasetransmitting mosquitoes— Continued

Eurasia—Continued

Burma 87

Indochina 89

Thailand 92

Malaya 94

Sunda Islands 96

Philippine Islands 101

Australasia 102

The Moluccas, New Guinea,

and Melanesia 102

Australia, Tasmania, and New

Zealand 105

Micronesia 106

Polynesia 106

Notes on the medically important

species 107

Aedes species 107

Anopheles species 112

Culex species 139

Haemagogus species 141

Mansonia species 142

Sahethes species 143

References 143

Glossary 151

Index 153
MOSQUITOES

of Medical Importance

By RICHARD H. FOOTE and DAVID R. COOK,* entomologists,

Entomology Research Division, Agricultural Research Service

Success in controlling arthropod-borne diseases during

military operations is frequently dependent on the rapidity

with which a situation can be correctly evaluated. This, in

turn, depends on having or quickly obtaining an accurate

picture of the populations of potential vectors of disease in

the area under consideration, for the disease rate prevalent

under normal conditions is not necessarily an index of the

disease rate that may quickly develop. An important step

in the assessment of the disease potential of an area is the

rapid separation and identification of mosquitoes of primary medical importance. This task is frequently diíBcult

or impossible, even for experienced taxonomists, unless adequate literature is at hand. Publications that fill this need

have been generally lacking in the past. This situation has

resulted in delays and errors in the activation of measures

to protect troops and civilian populations.

Because of the lack of suitable guides for the entomological

phases of preventive medicine during World War II, the

Office of the Surgeon General, Department of the Army,

began exploring ways and means of providing Medical Department personnel with concise and accurate guides to the

medically important arthropods likely to be encountered in

any part of the world where military operations might be

necessary. Mosquitoes, being the most important insects

from a military standpoint, received primary consideration.

After a careful study of the problems entailed and the type of

reference material needed by military preventive-medicine

units in the field, it was decided to develop a series of pictorial

keys that would aid in identifying the medically important

mosquitoes of selected geographic regions. The task of developing these keys was assigned to the Insect Identification

Division of the former Bureau of Entomology and Plant

Quarantine in 1950. The work was supported by a transfer

of funds from the Department of Defense to the Department

of Agriculture. Under this program over 60 pictorial keys

to medically important mosquitoes were developed.

Meanwhile, experiences of military preventive-medicine

personnel during the Korean war contributed further evidence as to the needs of field units for reference materials.

It was therefore decided to enhance the usefulness of the pictorial keys by supplementing summarized information about

mosquito-borne diseases and by adding further details about

the species of mosquitoes of medical importance. This handbook consists of the pictorial keys and the supplemental information. Although designed primarily for use of the Military Establishment, this publication should also prove useful

to persons concerned with public-health programs. Furthermore, the information on those species that are vectors of the

encephalitides makes evident the inseparable nature of

medical and veterinary entomology.

The pictorial keys were designed to eliminate from consideration all species that are not medically important and

those that are not known to be present in a region. The number of species requiring critical study is thus reduced. These

keys can be used effectively by persons having a minimum of

taxonomic training.

Published information pertaining to the biology and

medical importance of the mosquitoes included in the keys

has been summarized for each region and provides easily

accessible data useful in the identification of vectors, their

distribution, the diseases they carry, and the relative importance of the different vector species. The handbook is accordingly arranged in three sections. In the first section the

diseases are described, in the second the geographic distribution and the relative medical importance of the diseasetransmitting mosquitoes are given, and in the last section,

entitled ''Notes on the Medically Important Species," is summarized the published information pertaining to the

bionomics, relation to disease, and recorded distribution for

each species.

^ Resigned August 28,1953. Present address : Department of Biology, Wayne University, Detroit, Mich.
AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITO-TRANSMITTED DISEASES

The diseases transmitted to man by mosquitoes include

malaria, yellow fever, dengue, the encephalitides, and filariasis. Their importance, distribution, and epidemiology are

described here.

MALARIA

Malaria is an acute, intermittent, febrile disease that sometimes persists in individuals for several years. It is prevalent between latitude 45° N. and 40° S. around the world

and is responsible for more deaths per year than any other

arthropod-transmitted disease.

Malaria is a collective term referring in humans to a group

of four infections, each caused by a different species of Protozoa belonging to the genus Pldsmodium, These species

and the infections they cause are as follows :

1. P, vivajx,—This species has the widest distribution of

all the human malaria parasites. It causes benign tertian or

vivax malaria, characterized by a fever every 48 hours due

to q, periodic maturation of parasites in the blood. Relapses

are common.

2. P. falciparum,—This species tends to predominate in

tropical and subtropical regions. It causes malignant tertian, subtertian, or aestivo-autumnal malaria, which produces febrile attacks usually every 48 hours. Relapses are

rare.

3. P, malariae,—This parasite is distributed around the

world between about latitude 53° N. and 30° S. It is responsible for quartan malaria, which is characterized by

attacks of fever every 72 hours and persistent infections,

possibly lifelong, which have been reported to cause relapses

up to 60 years after the initial infection.

4. P, ovale,—This is a relatively rare species found in isolated areas of South America, certain parts of Africa, and

the Orient.2 It is widely distributed throughout western

Africa, but its occurrence in the eastern part is spotty. It is

responsible for o'vale malaria, which is similar to vivax malaria, with febrile attacks at 48-hour intervals and subsequent

relapses.

Chronic cases of falciparum^ vivax^ or malariae infections,

but predominantly falciparum^ may develop into a particularly dangerous complication of malaria known as blackwater fever. I'his is accompanied by a breakdown of large

numbers of red blood cells in a short time, with hemoglobin

in the urine. The subsequent physiological imbalance may

result in a 25- to 50-percent death rate.

The four malaria infections are invariably transmitted

among humans by mosquitoes belonging to the genus Anopheles. Female mosquitoes draw up the parasites along with

human blood while they feed. Parasites enter the gut of

the mosquito, undergo a series of developmental stages, and

eventually invade her salivary glands. The female injects

these parasites along with her saliva at a subsequent feeding.

If conditions are such that sufficient numbers of infected

and susceptible persons are in close proximity to highly anthropophilic ^ vectors in which the parasites have matured,

malaria transmission will usually occur. The severity of

this transmission is classified as endemic, hyperendemic, or

epidemic, depending on the abundance and proximity of the

participating elements, the length of the transmission season, and other factors.

Only 85 of the approximately 400 species of Anopheles

have definitely been incriminated in malaria transmission.

They are included in the pictorial keys and discussed in detail in the other two sections of this handbook. The physiological factors that allow only the Anopheles species to

carry human plasmodia are not well understood. Even less

is known about the conditions that enable these parasites to

mature efficiently in some species of Anopheles and not in

others. Although our epidemiological knowledge concerning

these differences has increased rapidly during the last three

decades, the absence of malaria from some areas where known

vectors abound even today remains largely unexplained.

References: 61, 24ß.^

YELLOW FEVER

Epidemiologically the two principal forms of this disease

are classical urban yellow fever and jungle yellow fever.

Both are caused by the same mosquito-transmitted, filterable

virus. The virus is picked up by the female mosquito with

her blood meal and undergoes multiplication for 1 to 3 weeks

in her body. The time required for this process depends in

part on the temperature and humidity prevailing in the

mosquito's habitat. At the end of this time the infection

is transmissible to other hosts, and the mosquito remains

infective during the remainder of her life.

The virus causes certain pathological and physiological

changes in the liver of its mammal host, and a lifelong immunity is imparted upon recovery. When infected persons

enter an area, about 2 to 3 weeks usually elapse before

additional cases appear. This corresponds to the incubation period in the mosquito. The virus appears in the

blood of an infected person for only the first 3 to 5 days

after he has been bitten.

Classical urban yellow fever is principally an epidemic

disease. It is transmitted by Aedes aegypti^ a ubiquitous

urban mosquito that breeds in artificial containers of all

kinds close to human habitations. The disease is usually

introduced by infected people into seacoast communities

that have dense, nonimmune populations, and it is here

that yellow fever appears in its most severe form.

Jungle yellow fever, although caused by the same virus,

is epidemiologically very different from the classical type.

It is primarily a disease of forest mammals, especially monkeys, and is transmitted by mosquitoes that live in forest

environments and prefer monkey blood to that of man. It

is always tropical in distribution. Forest mammals do not

appear to form a reservoir, since, as in man, the virus disappears from their blood soon after they have been bitten

by infected mosquitoes.

In eastern Africa this disease is typically that of forest-

^The Orient, as used in this handbook, includes the foUowing

regions: Afghanistan and West Pakistan, India and Ceylon, East

Pakistan and Assam, South China, Formosa, Burma, Indochina,

Thailand, Malaya, Sunda Islands, and Philippine Islands.

^For definitions of certain terms used in this handbook, see the

Glossary, p. 151.

* These italic numbers refer to References, p. 143.
MOSQUITOES OF MEDICAL IMPORTANCE

inhabiting monkeys. A monkey-to-monkey transmission

cycle is believed to be maintained by Aedes africanus and

possibly A. mttatus. These mosquitoes breed in tree holes

and plant containers and inhabit the forest canopy. However, at ground level, especially in plantation areas, the

principal vector is Aedes simpsoni. This species feeds on

infected monkeys either outside or just within the forest,

but, unlike africcmus or mttatus^ feeds on man as well, acting as a transmitter in the monkey-to-man cycle. Aedes

vietallicus^ A, Iwteocephalus^ A, furdfer^ and A, taylori may

act in much the same capacity as simpsoni to spread yellow

fever from monkey to man. A. aegypti is believed by some

workers to play a minor role in this activity.

In South America, jungle yellow fever is even more incidentally a disease of humans than in Africa. There appears

to be no mosquito such as simpsoni to effect a monkey-to-man

cycle. Therefore, all the human disease is contracted by

people who work within the forest and come in contact with

the canopy haunts of the vectors by climbing trees for various

purposes or by those who cut timber and bring the vectors

to the ground. Itis primarily a disease of young men ; women

and children are rarely affected, because they seldom frequent

the forests. Haerrmgogus albomaculatus^ H, equinus^ H,

spegazzinii falco^ H, s, spegazzinii^ H, caprieomii^ Aedes

leuGocelaenus leucocelaenus^ and A, I, clarhi are the principal

vectors of jungle yellow fever in Central and South America.

Since immunity to yellow fever is so strong and lasting,

it is used to determine the distribution of the disease. Large

areas of South America and Africa have been surveyed.

Endemic areas in which the disease has occurred in the past

have been delimited by the discovery of immunity in persons

who have never had yellow fever vaccine.

Reference: %I^Q.

DENGUE

The virus causing dengue is similar in particle size to that

of yellow fever. All 10 strains now known belong to 1 of 2

distinct immunological types, Hawaiian or New Guinea. All

strains produce similar symptoms. The virus causes an acute

illness accompanied by fever, and it is present in greatest

concentration in the patient's blood during the first 24 hours.

Immunity to any one strain lasts for the rest of the person's

life.

Aedes aegypti^ A. alhopictus^ and members of the A, scutellaris group transmit the disease in nature. Recent laboratory and epidemiological observations show that Aedes

polynesiensis^ a member of the soutellaris group, may be an

important vector of dengue in Tahiti, Manua (American

Samoa), and the Marquesas Islands. Experiments have

shown that scutellaris itself is probably the vector of a jungle

type of dengue in New Guinea, and possibly other members

of the scutellaris group are capable of transmitting the disease elsewhere. The virus has been transmitted experimentally from one monkey to another by means of infected

mosquitoes, and several species of monkeys are apparently

susceptible to dengue. As far as is known dengue has no

animal reservoir ; it is transmitted directly from man to man

by the mosquitoes mentioned above. The virus requires an

incubation period of at least 8 days in the body of the mosquito before it can be transmitted in virulent form to susceptible persons. The mosquito is then able to transmit

dengue for the remainder of its life.

Because the virus is present in monkey or human blood for

such a short time, transmission of endemic dengue takes place

only in areas that can supply Aedes vectors throughout the

year. Dengue is endemic or epidemic in various regions of

the South and Southwest Pacific; Hawaii; northern and

eastern Australia; New Guinea; Indonesia; India; the

countries bordering the South China Sea ; Japan ; much of

tropical Africa and along the Nile Valley to the Delta ; the

countries bordering the northern shore of the Mediterranean ;

and some areas between latitude 10° N. and 10° S. in the

Western Hemisphere, notably Trinidad. Our knowledge of

the exact distribution of the disease in many parts of the

world is still sketchy and inadequate. Epidemics, which

have occurred in many of the regions mentioned, are believed

to be due to the importation of the virus into areas containing

large numbers of nonimmune people.

References : 2Jf6, 256, 289, 291, 356.

THE ENCEPHALITIDES

The encephalitides include all the diseases caused by encephalitis and certain other closely related viruses and all

are mosquito borne. Most of these viruses are relatively

small, being about 15m/A to 30 m^u. They attack various parts

of the central nervous system. Apparently humans are only

incidental hosts of these viruses. Little is known about the

natural reservoirs, overwintering characteristics, or exact

mode of transmission of many of them.

Encephalitis Viruses

Western Equine Encephalitis.—The principal endemic

region of this virus, where animals are the host, comprises

all States west of the Mississippi River ; also Wisconsin, Illinois, Indiana, Kentucky, Mississippi, Alabama, and Florida ;

and parts of Canada adjacent to these States. Eecently this

virus has been reported from Florida, Alabama, Tennessee,

Kentucky, North Carolina, New Jersey, Michigan, and Illinois. The largest epidemic occurred in 1941 in Minnesota,

North Dakota, and Canada.

Studies of natural infections and occurrences of the disease in the presence of large numbers of infected mosquitoes

point to Culex tarsalis as the primary vector. This virus

also is found in Argentina and possibly Peru and central

Chile, but the vector in those countries is not known. Antibodies from domestic and wild birds and mammals have

been isolated in nature, and mites from such animals have

been shown to harbor the virus. Since tarsalis prefers these

wild animals and birds as hosts, man is only incidentally

infected. The virus appears to be maintained in nature

chiefly by a hivà-tarsalis-hvcà transmission.

St. Louis Encephalitis.—This virus appears sporadically

in the Midwestern and Southwestern United States and has

been reported in an almost continuous belt across the United

States. It has endemic areas in Nevada, California, Oregon,

and Washington. A large epidemic occurred in St. Louis

in 1933 and a smaller one in 1937. Epidemiological studies

there involved Gulex pipiens^ but the virus has only been

isolated from mosquitoes in the Western United States.

Considerable evidence indicates that Culex tarsalis is probably the primary vector of St. Louis encephalitis in the

United States. Antibodies to this virus have been found

in chickens and wild birds, and it has been isolated from

chicken and wild-bird mites, indicating that this virus, like

the western strain, probably has an animal reservoir from
AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

which man is accidentally infected by mosquitoes whose

hosts are not normally man. Recent investigations indicate

that St. Louis encephalitis virus might have infected humans in Trinidad.

Eastern Equine Encephalitis.—This virus is found

throughout eastern Canada and the Eastern and Southern

United States east of the Mississippi River, and it has spread

into Nebraska, Missouri, Arkansas, Louisiana, Texas, Mexico, Panama, Brazil, Dominican Republic, Cuba, and Trinidad. Encephalitis cases believed to be of the eastern equine

type were noted in the lower Rio Grande Valley in 1941,

and additional cases have been recorded in Michigan, Wisconsin, Missouri, and Arkansas.

Major outbreaks occur in horses and pheasants. Only

two epidemics affecting humans have been recorded, one in

Massachusetts in 1938 and one in Louisiana in 1947. Eastern equine encephalitis virus has been isolated many times

from Culiseta melanura^ indicating that this species may be

the principal vector, although the available evidence indicates that other species may be involved. Wild birds are

important in the dissemination of the virus, since antibodies

frequently have been isolated from them. Pheasants are

particularly suspected and domestic fowl are presumed to

be reservoirs, since the virus has been isolated from their

parasites.

California Encephalitis.—This virus has been recovered

from Aedes dorsalis and Culex tar salis in the central valley

area of California. It has produced clinically inapparent

infections in humans, horses, and other mammals in this

area. The role of these mosquitoes in transmission is not

exactly known.

Venezuelan Equine Encephalitis.—This virus is found

in Panama, Trinidad, Venezuela, Ecuador, Colombia, and

Argentina. In Espinal, Colombia, mild human infections

were observed, although the extent to which this virus causes

disease in humans is not known. In the laboratory it is

highly infectious. In Venezuelan equine as well as eastern

equine encephalitis, many mosquito species have been suspected of transmitting the virus. Available evidence indicates that Mansonia titillans is a vector. Few studies have

been made of animal reservoirs of this virus.

Japanese Encephalitis.—Epidemics of Japanese encephalitis affecting large numbers of humans have occurred in

Japan, Okinawa, Korea, and Guam. There is evidence of

widespread infection in Malaya and Borneo, and the presence of this disease in India, eastern China, Burma, Thailand, Indochina, Formosa, Sumatra, and Java is suspected.

Japanese and American workers have reported positive sera

from the Philippines. The virus has been reported from

Australia, but recent serological work indicates that Murray

Valley virus is probably the agent there instead.

Abundant evidence exists that, at least in Japan, Culex

tritaeniorhynchus is the important vector. The appearance

of this mosquito in large numbers is usually followed by

an increase in disease incidence, and naturally infected

mosquitoes always appear at critical times during the course

of an epidemic. In Russia the virus has been isolated from

a mixture of tritaeniorhynchus and Culex pipiens, C, tritaeniorhynchus has been incriminated in other areas, such

as Korea, Formosa, and Okinawa, by presumptive evidence

alone, and at least in Okinawa no conclusive field investigations of the possible vector have been completed. In China

pipiens alone has yielded this Japanese virus in nature. A

blackbird, swift, and dove were found to have antibodies

associated with an epidemic that occurred in Guam in 1947

and 1948, but the vector in that island is not known; tritaeniorhynchus does not appear there. In Malaya the virus

has been isolated from four fatal human cases, but so far

the vector is not known in that country. A form similar

to the Japanese strain has recently been found in Indochina ; its vector is likewise unknown.

Murray Valley Encephalitis.—This virus, apparently

limited to Australia, is related to the Japanese, West Nile,

and St. Louis encephalitis viruses. It has received little

field study and its vector is not known, but Culex annulirostris and several other species are the most strongly suspected mosquitoes. Higher percentages of water birds

harbor antibodies than any other bird or mammal species

tested, although domestic land fowl also carry immunity.

The large migratory-bird population in Australia is believed to carry the virus from its northern endemic centers

southward to New South Wales and northern Victoria during

years of heavy rainfall.

Other Mosquito-Borne Viruses

Epidemiological investigations of the following viruses

from South America and Africa have not progressed far

enough for positive identifications of vectors or animal

reservoirs.

Bwamba.—This virus causes a disease with fever, headache, malaise, and conjunctivitis lasting for 4 or 5 days. It

appears most commonly in eastern Belgian Congo, Uganda,

western Kenya, and northern Tanganyika, and antibodies

have been found in natives in Northern and Southern

Rhodesia. Its mode of transmission is unknown.

Ilhéus.—This is a virus related to the West Nile, Japanese,

and St. Louis encephalitis viruses. It has been observed in

the Ilhéus area of Brazil and in Trinidad, where neutralizing antibodies have been found in humans. In both localities

the virus has been isolated from mixtures of various species

of mosquitoes, in which Aedes serratus and Psorophora

ferox have predominated.

Rift Valley.—This virus in man causes a self-limited

disease, with fever, severe headache, and other symptoms

from which the patient recovers in several days. It is primarily a disease of sheep, cattle, and other animals in the

Rift Valley and neighboring areas of Africs^. An extremely

high incidence of infection occurs in laboratory personnel

who work with this virus. The principal vector suspects

are Aedes dehoeri demeilloni^ A, tarsalis^ and Eretmapodites

spp., the virus having been isolated from wild-caught

females.

Sindbis.—This virus is related antigenicly to eastern,

western, and Venezuelan equine encephalitis viruses and has

appeared in Egypt and Anglo-Egyptian Sudan. No information is yet available on its symptomatology, but antibodies to the virus have been detected in man, domestic

animals, and birds. Human infections are frequent. It

has been experimentally transmitted by Culex pipiens and

G, univittatus.

West Nile.—This virus was first isolated from the blood

of a woman with mild fever in the West Nile district of

Uganda. It has been observed in Egypt, Israel, Kenya,
MOSQUITOES OF MEDICAL IMPORTANCE

Belgian Congo, and Sudan. Antibodies in humans have

been found in many parts of India. The virus is related

antigenicly to those causing St. Louis, Murray Valley, and

Japanese encephalitis and dengue and yellow fever. In a

highly endemic area in Egypt no deaths or frank cases of

encephalitis were attributed to it. The West Nile virus

causes primarily an early-childhood disease, which is most

prevalent during the summer. Nonmigratory birds and

mosquitoes appear to form the primary cycle, and man

and domestic animals are incidentally infected. In Egypt

Culex imivittatus is probably the most important vector.

The virus has caused a self-limited and nonfatal disease

in over 100 patients in Israel, most of them children.

In the widespread search for immunity to yellow fever,

the following mosquito-borne viruses have been isolated

from the blood of natives in Africa and the Americas. Little

is known about their ability to produce disease symptoms

in man. For additional details of distribution, see the discussions under the heading "Encephalitis" in the next section

of this handbook.

Anopheles "A/^—Anopheles ioUviensis is the only vector

of this virus that is reported with certainty. It is a South

American virus, but its exact distribution is not known.

Anopheles "B."—Anopheles holiviensis^ Haemagogus spegaBBinii^ Aedes leucocelaenus^ and Sabethes sp. have been incriminated in the transmission of this virus. The distribution of this South American virus is not known with

certainty.

Bunyamwera.—This virus has been isolated from mixed

Aedes species in east Africa, and antibodies in humans have

been found in Kenya, Uganda, Tanganyika, Belgian Congo,

and possibly Egypt.

Ntaya.—This virus has been isolated in eastern Africa

from pools of mosquitoes in which Culex spp. predominated.

Semliki Forest.—^Antibodies to this virus have been found

in some asymptomatic natives in Tanganyika and Uganda.

Eretmapodites grahand and mosquitoes closely related to

Aedes abnormalis are the suspected vectors.

Uganda S.—This virus has been isolated from pools of

Aedes spp., in which A. longipalpis and A. ingrami predominated. Neutralizing human antibodies have been found

in Kenya, Belgian Congo, and Tanganyika.

Wyeomyia.—Wyeomyia melanocephala appears to be the

important species in the transmission of this virus. The distribution of this South American virus is not known.

Zika.—Aedes africanus is the suspected vector of this

virus. Human antibodies have been discovered in Egypt,

Belgian Congo, Tanganyika, and India, and the virus has

been recovered from human blood in Nigeria.

References : 39, JfS, 58, 109, 111, 153, 154, 156, 246, 257, 270, 274, 278,

281, 299, 325-327, 355, 356.

FILARIASIS

Initial symptoms of filariasis last a week or 10 days and

consist of regional inflammations, with fever, swelling, and

stiffness. From 3 months to a year after these symptoms

subside, a second phase usually starts with swelling in various lymph glands. In later stages elephantiasis sometimes

occurs after repeated attacks. Caused by a wholesale stoppage of lymph flow over large parts of the body, elephantiasis results in the enlargement of the lower extremities,

scrotum, and breasts, sometimes to extreme proportions.

Filariasis is caused by nematodes belonging to the superfamily Filaroidea. As adults these nematodes live, usually

in pairs, in the lymphatic vessels, lymph nodes, and connective tissues of their human host. Sexually mature female

worms in these locations produce embryonated eggs that

hatch. The resulting larvae, or microfilariae, migrate to the

peripheral blood, where they are picked up by mosquitoes

as they feed. They penetrate the gut wall of the mosquito,

undergo extensive morphological changes, and migrate to the

proboscis, from which they emerge and enter the intact skin

or the bite puncture of the person bitten at a subsequent

feeding of the mosquito. They migrate in various paths to

the lymphatics and connective tissues of their human host,

where they eventually transform to adults and initiate another cycle.

Periodic filariasis is caused by Wuchereria hancrofti and

W. malayi. The numbers of microfilariae of these two species alternately increase and decrease in the peripheral blood.

This periodicity takes place every 24 hours, the greatest

number of microfilariae occurring between 8 p. m. and 2 a. m.

The mosquitoes that transmit periodic filariasis are those

that are most active during the hours when microfilariae are

most abundant in the peripheral blood of infected persons.

A nonperiodic filariasis occurs in certain areas of the Pacific. It is caused by a filaria that some workers have

named Wuchereria pacifica and which is morphologically

identical to hancrofti. Its exact taxonomic status has not

been settled.

Although the life cycles of malayi and hancrofti.^ the

symptoms they cause, and their adult stages are essentially

alike, there are distinct morphological differences between

their respective microfilariae. Diagnosis depends on their

identification in the peripheral blood. Another important

difference between the tw^o species lies in their respective

vectors. W. malayi is transmitted principally by Mansonia

(subgenus Mansonoides) and Anopheles mosquitoes, whereas

hancrofti is transmitted by various species of Anopheles^

Culex^ and Aedes.

References: 65, 162, 246, 321.

GEOGRAPHICAL DISTRIBUTION AND RELATIVE MEDICAL IMPORTANCE

OF DISEASE-TRANSMITTING MOSQUITOES, BASED ON PICTORIAL KEYS

In this study the world has been divided into 35 rather

arbitrary geographical regions, because so much variation

exists from one region to the next in the diseases transmitted

by mosquitoes and in the kinds and distribution of these

vectors. The extent of each region has been defined in part

by the number of medically important species that could

be conveniently included in a single-page pictorial key.

The following information has been given for each region :

1. The countries and/or parts of countries included there.

2. A brief description of the physical geography and outstanding features of temperature and rainfall.

3. The distribution of malaria and the distribution and
6 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

relative importance within the region of each of the important vectors included in the keys.

4. A list, compiled from the literature and personal communications, of all the Anopheles species reported from the

region.

5. Discussions of yellow fever, dengue, the encephalitides,

and filariasis, including their vectors.

making the keys, and discussions of that disease and its vectors are limited to the text, except when vectors of filariasis

are also vectors of any of the other four diseases, for example Anopheles sinensis in North Europe and North Asia.

A species was selected as important when it had been reported in the literature by unquestionable evidence as a vector in a region or in regions bordering it. A large number

OUTER CLYPEALS

NO SIPHON ON ANOPHELES

POSTSPIRACULAR HAIR

SPIRACULAR

APPARATUS

ALGER/ENS/S

BIFURCA TUS

^C/NEREUS

HYRCANUS

¥: MACULIPENNIS

MARTERI

PLUMBEUS

SOGDIANUS

ARABIQUE

* CULICIFACIES

D'THALI

PLUVIA TILIS

MULTICOLOR

PULCHERRIMUS

RHODESIENSIS

SERGENTI

¥c STEPHENSI

^SUPERP/CTUS

— TURKHUDI

LONG PLEURALS

OF 2ND GROUP

BOTH

BARE

LONG PLEURALS

OF 3RD GROUP

ABOUT

15

BRANCHED

ANTENNAL HAIR

1/4 DISTANCE

FROM BASE

+

- ALGERIENSÍS

- BIFURCATUS

HYRCANUS

^ MACULIPENNIS

- MARTERI

--»PLUMBEUS

'»SOGDIANUS

-^HYRCANUS

* MACULIPENNIS

ABDOMINAL PLATES 4 TO 7

^ MACULIPENNIS

CAUDAL HAIRS

ALGERIENSIS

BIFURCATUS

HYRCANUS

^MACULIPENNIS

ONE /

HAIRED y //À

MARTERI

PLUMBEUS

SOGDIANUS

ARÁBICOS

* CULICIFACIES

D*THALI

FLUVIATILIS

^ - ARÁBICOS

¥c CULICIFACIES

D'THALI

''FLUVIATILIS

+

NOT CURVED

AT TIPS

GUT~-

LONG PLEURALS

OF 3RD GROUP

ONE

HAIRED

¥i CULICIFACIES

- D*THALI

PALMATE HAIRS

ON SE-GMENT 3

CURVED AT TIPS

¥: CULICIFACIES

OUTER CLYPEALS

FILAMENT

1/3 OR MORE

AS LONG AS

BLADE

- PULCHERRIMUS

-^ SUPERPICTUS

MULTICOLOR

PULCHERRIMUS

— RHODESIENSIS

- SERGENT!

* STEPHENSI

* SUPERPIC TUS

MULT/COLOR

PULCHERRIMUS

^STEPHENSI

•K SUPERPIC TUS

» SUPERPICTUS

HIND CLYPEALS

NOT

FILAMENT -

LESS THAN

1/3 AS LONG

AS BLADE

'^MULTICOLOR

^ STEPHENSI

¥r STEPHENSI

FIGURE 1.—AmpUfled part of key to Iranian Anopheles larvae.

One key to larvae and one to adults have been included for

each region. All species of mosquitoes known to live in a

given region were considered in the preparation of the keys.

The species to be illustrated were selected on the basis of

their importance in the transmission of the militarily important diseases malaria, yellow fever, dengue, and the encephalitides. Filariasis, therefore, was not considered in

of mosquito species incriminated from time to time in disease transmission are excluded from the keys, because they

do not meet these criteria or because control of the particular disease does not necessarily depend on vector control.

The subspecies are not always distinguished in the keys, primarily because of lack of space. In some regions they are

not known.
MOSQUITOES OF MEDICAL IMPORTANCE

NO SIPHON ON ANOPHELES ANOPHELES LARVA

SIPHON ON ALL OTHERS

PLEURAL BRISTLES

ON UNDER SIDE WITH A BASAL\

SPINE-LIKE SHEATH /

rGROUP

Z"- GROUP

3"» GROUP

ABDOMINAL

PLATES

4 TO 7

NO

SIPHON

CAUDAL HAIRS

C

FIGURE 2.—Larval characters for use with the keys: a, Terminal abdominal segments of an Anopheles larva; ö, terminal abdominal

segments of a non-Anopheles larva, showing the siphon ; c, top view of an Anopheles larva.
8 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

The following instructions should be read carefully before any attempt is made to identify specimens.

DIRECTIONS FOR USE OF THE KEYS

In order to demonstrate the technique for using the illustrated keys, a part of the Iranian larval key has been

amplified, as shown in figure 1. The larval characters used

in figure 1 and throughout the illustrated keys are shown in

figure 2.

In order to use figure 1 to identify a larva, the specimen

must be from Iran. If it lacks a siphon (see fig. 2, a and h)

and its terminal abdominal segments resemble those shown

in figure 1, column 1, under the heading "no siphon on

Anofheles!!'' it is an Anopheles larva. The Anopheles

species known or thought to be in Iran are listed at the

bottom of this column. The medically important species

are designated by asterisks (*). The minus signs ( —)

indicate those species in each column that are not listed in

the succeeding column, because they lack the combination

of characters shown there.

In figure 1, column 2, under the heading "long pleurais

of 2nd group" (see also fig. 2, c),^ two possible choices are

shown. If both long pleurais on the larva are bare, it belongs to one of the species listed under "both bare." If one

of the long pleurais is haired, the larva belongs to one of the

six species listed under "one haired." If both are haired,

the larva is eliminated from further consideration, since

larvae of all the medically important species in which this

study is concerned happen to have one or both of the long

pleurais of the second group bare.

If both long pleurais of the second group are bare, see

figure 1, column 3. Two bracketed choices are given. If one

of the long pleurais of the third group is haired and one is

bare, it must belong to one of the four species listed under

the second choice.

Be sure to follow specifically the directions implied by

the brackets. The unbracketed key character labeled "long

pleurais of 3rd group both haired" at the bottom of column

3 applies only to larvae possessing one haired second-group

pleural (see bottom of col. 2). If it had had two haired second-group pleurais, it would have been eliminated from

further consideration, since the key shows no larva with a

combination of both second-group pleurais bare and both

third-group pleurais haired.

If the larva has both second-group pleurais bare (see fig.

1, col. 2) and one third-group pleural haired (see col. 3),

note the key characters in column 4. The plus ( -f ) sign is

an indication to continue as if a bracket were present. If

abdominal plates 4 to 7 on the specimen conform to those

in figure 1, column 4, the larva is one of the two species

listed below "abdominal plates 4 to 7." If it possesses the

curved caudal hairs shown in figure 1, column 5, under

"caudal hairs," the specimen is Anopheles culicifacies. Thus

it has been identified as a medically important species, because it has all the characters indicated in the key, and at

the same time it has been separated from all other mosquitoes found in the region.

A specimen of Anopheles sergenti^ a species that is dropped

from the key as medically unimportant in Iran, can be run

through the key as follows : Since the larva lacks a siphon,

^ The first group of pleural bristles is not used in identifying larvae

of Iranian Anopheles.

it is therefore an Anopheles, See the choice of characters in

figure 1, column 2. As one of the long pleurais of the second

group is bare and one is haired, refer to the bottom of

column 3. Because the larva has one of the long pleurais of

the third group bare, this character does not correspond to

the only illustration provided. Therefore, the larva is rejected from further consideration, and the species is not

medically important in the region treated by the key.

Accordingly, any species of Anopheles in Iran may be

run through the key and either identified as medically important or dropped entirely from consideration at whatever

point it fails to agree with the stated character or choice of

characters.

Although geographical variation in species has been accounted for to the greatest possible extent, the user of these

keys should be cautioned that all variation has not been

illustrated in the keys. The entire range of variation of

most of the species is not adequately known. It should also

be understood that the illustrations usually depict the typical

appearance. For distributional and biological data that

might help to confirm the identification, consult the section

of the text describing the specific region and also the description of the particular species in the last section of this

handbook. Since many species have not been available for

direct study, they are presented on the basis of the literature

alone. An attempt has been made to verify the taxonomic

characters of each of these species by correspondence with

authorities familiar with them.

NORTH AMERICA

America North of Mexico

(Figs. 3 and 4)

Physical Features.—The territory covered by the accompanying keys (figs. 3 and 4) extends from the North Pole

to the southern border of the United States along the Mexican border and the Gulf of Mexico, and from the Bering

Strait to the eastern edge of Greenland. It includes Greenland, Alaska, Canada, and the continental United States.

The principal western mountains forming the Continental

Divide are the Rockies and the Alaska Range, which extend

from the Arctic Ocean to New Mexico. Between these

mountains and the Pacific Ocean are the Sierra Nevadas and

Cascades in the United States. Except in scattered localities

this west coast has no coastal plain. The Great Plains

adjoin the eastern edge of the Rockies for most of their

length. In northern Canada the Barren Grounds and Laurentian Plateau extend eastward, the latter to the Atlantic

Ocean. A vast fertile lowland between central Saskatchewan and Hudson Bay extends south to the Gulf of Mexico.

This is drained to the Gulf by the Mississippi-Missouri river

system and to the Atlantic by the St. Lawrence. The only

uplands in this region are the Ozarks and the Ouachitas in

the South Central States. The Appalachians, including the

White Mountains, and the Adirondacks comprise the main

eastern highlands of the United States, the east coast of

which is a narrow sandy plain, widening in the southeast

corner to cover almost the entire peninsula of Florida.

Along the west coast the climate is moderate but becomes

arid in the desert area of the Southwestern United States.

From 20 to 40 inches of rain fall over the central plains;

much of the south, southeast, and east coasts receive up to

60 inches, sometimes more. Eighty-five percent of Greenland, a great island basin encircled by a mountain chain,
MOSQUITOES OF MEDICAL IMPORTANCE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE-AMERICA NORTH OF MEXICO

FULL-GROWN LARVAE ^W^^—

NO SIPHON ON ANOPHELES

OUTER CLYPEALS

INNER CLYPEALS

SEPARATED

BY WIDTH

OF ONE

TUBERCLE

OR LESS —

"- i^^ Vwli«

ABDOMINAL PLATES

OF 4TH SEGMENT

-HAIR^'O" ON 4TH

SEGMENT LARGE _ ANOPHELES

~~ CRUCIANS

\ DRAWING FOR LOCATION/

.-—HAIR 2 ON 4TH

SEGMENT TWO OR ANOPHELES

THREE BRANCHED = frpEEBORNI

SEPARATED

BY MORE

THAN WIDTH \

OF ONE

TUBERCLEANTENNAL HAIR

NEAR MIDDLE OF ,

SHAFT AND

REACHING

+ TIPm

_ ANOPHELES

— QUADRIMACULATUS

ABDOMINAL PALMATES

OF4TH SEGMENT

SPIRACULAR PLATE

ANOPHELES

PSEUDOPUNCTIPENNIS

PSEUDOPUNC TIPENNIS

4TH ABDOMINAL SEGMENT

OF ANOPHELES LARVA

PALMATE HAIR '

(HAIR 1)

AEDES

AEGYPTI

FIG. 3

HEAD OF ANOPHELES LARVA |

ANTENNAL

HAIR INNER

/ CLYPEALS

' OUTER 1 HIND

1 CLYPEALS ^ CLYPEALS

\ HANTENNAV ^^^ 1 ñ mM Ä^;JMé

iMT-gWi

Vv/n

PROTHORACIC HAIRS 1^-^~~

ANOPHELES LARVA

PLEURAL BRISTLES

SPIRACULAR

PLATE

is covered with an icecap, with only the southwest coastal

part suitable for limited agriculture.

Reference: SOI.

Malaria.—This disease is almost nonexistent in America

north of Mexico. It became intense and widespread in the

Eastern United States with the importation of slaves and

the onset of the Eevolutionary War, persisted throughout

much of the settling of the West, and by 1850 had been reported from every State except North Dakota, Montana,

Idaho, Wyoming, Utah, and Nevada. From its original

focus in the Carolinas it spread slowly north and west, but

since 1933 it has declined steadily. Reports of the United

States Public Health Service indicate that malaria is no

longer a medical problem, except for cases brought back

to the States. These cases can serve as centers of infection

under favorable conditions. Only 25 new indigenous cases

were reported in 1952. Malaria control is limited to the

treatment of areas in the immediate vicinity of each new case.

In Canada before 1880 malaria had become endemic along

the north shores of Lake Ontario, Lake Erie, and the Canadian side of Lake St. Clair but not above the summer

isotherm of 70° F. At one time it was reported to have

become endeniic in certain parts of the Western Provinces,

but no malaria has been reported from Canada for several

decades.

The little malaria in the temperate regions is almost all

caused by Plasmodium vivax. Plasmodiimh falciparum is

associated with a larger number of cases in the subtropical

regions, and P, malariae^ although present, has never contributed to the incidence of malaria to any important extent.

In the entire area east of the Great Plains, including the

Mississippi River basin. Anopheles quadrimaculatus has been

responsible for most cases of malaria. It is the Nearctic

species most fond of human blood, and its ubiquitous breeding and house-entering habits make it North America's most

important vector. This species has become especially important in the Tennessee Valley Authority area, where the

construction of large bodies of water has provided ideal

breeding places for it. Although the anophelines responsible

for nialaria in Canada have never been determined, evidence

points to qvxidrÍ7naculatus as the probable vector there.

Anopheles crucians is especially abundant along the Atlantic coastal plain. Evidence for its incrimination in malaria transmission in the United States is conflicting, because

quadrimacuíatus is also present over most of the same area,

but crucians is definitely suspected because of its anthropophilic tendencies, high density, and tendency in some areas

to enter human habitations.

Farther west but overlapping the range of crucians^

Anopheles pseudopunctipennis pseudopunctipennis has a
10 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE -AMERICA NORTH OF MEXICO

-^^^^ FEMALES ^

\_MAL£S HAVE BUSHY ANTENNAE

LONG PALPS ON

ANOPHELES

ANTENNAE

.PROBOSCIS

AND DO NOT BfTE^

PATTERN ON HIND TARSUS

FIG. 4

ENTIRELY DARK

ANOPHELES

QUADRIMACULA TUS

(EASTERN UNITED STATES)

AND

ANOPHELES FREEBORNI

(WESTERN UNITED STATES)

WING LIGHT SPOTTED; FRONT MARGIN DARK

( WING OF CRUCIANS )

ANOPHELES

CRUCIANS, GEORG/ANUS,Mü BRADLEYI

/ONLY CRUCIANS IMPORTANT; FEMALES\

V CANNOT BE SEPARATED 1

PATTERN ON HIND TARSUS

_ ANOPHELES

— ALB¡MANUS

DISTRIBUTION OF ANOPHELES

QUADR/MACULATUS AND FREEBORNI

PATTERN ON PALP

.LIGHT AT Ti ANOPHELES

ZL PSEUDOPUNCTIPENNIS

PSEUD0PUNC7IPENNIS

SHORT PALPS ON

ALL OTHERS

r PATTERN ON THORAX PATTERN ON HIND TARSUS

CULEX

TARSALIS

— AEDES

"" AEGYPTI

ADULT MOSQUITO WITH PARTS LABELED

ANTENNA^ ^

CLYPEUSTHORAX-,„

y HEAD^ ^ J(-:^]Vj~Zr ^

PALP »f^Ci^c-

"/ // EYE \ V, HIND TIBIA

FORELEG ^

/ '"'' '"" \H'ND '-EG\^--~..^_^^^,^^ ^^^3^3

/^-PROBOSCIS \v ^-^ .^ —íT^

northward distribution from Mexico. In the United States

its importance is not exactly known, since few transmission

studies have been made specifically of it. However, it is a

vector of prime importance south of the Mexican border,

and without doubt it could be an important vector in the

United States under favorable circumstances.

In the Western United States and extreme southwestern

Canada Anopheles freehomi is believed to be the principal

vector. It shows about the same degree of anthropophilism

as quadmmaculatus and is probably responsible for much of

the malaria within its range. It is definitely a seasonal

mosquito and survives the winter as an adult. The females

feed intermittently during this period and may carry malaria

from one person to another inside heated houses.

Studies indicate that Anopheles punctipennis may be a

vector of malaria in California. This species has not been

included in the keys for lack of conclusive evidence of its

ability to transmit disease.

Anopheles alhimanus^ the chief vector of malaria in the

Caribbean area, is included in the keys because of its potential rather than actual importance. Like psevdopunctipennis^ its role in the United States has been little studied,

but it probably does not contribute much to malaria transmission in this country. Its exact range in the United States

is not known.

References : ^i, 67, 7i, 113,129, U6,

Anopheles Species ^ in America North of Mexico

perplexens

pseudopvMctipennis franciscanus

\pseudopunGtipennis pseudopunctipennis

punctipennis

* quadrimaculatus

walkeri

\albimanus

átropos

harheri

hradleyi

"^crucians

earlei

"^freehomi

georgianus

occidentalis

Yellow Fever,—This disease occurred at one time in America north of Mexico, but it has not been reported since 1905.

Aedes aegypti^ its principal vector in many other parts of

the world, is included in the keys only because of its potential

importance, since the disease is no longer a medical problem

here.

Haemagogus equinus was collected from two localities in

the vicinity of Brownsville, Tex., in 1955. This mosquito is

a good laboratory vector of yellow fever, but the virus has

never been recovered from it in nature. Since eqmnus is

* Species marked with an asterisk ( * ) are recorded as vectors of

malaria in this region and their medical importance is discussed in the

accompanying text. Those marked with a dagger ( t ) are not recorded

as vectors of malaria in this region but are important elsewhere. For

information on the latter vectors, see the description of the particular

si)ecies in the last section of this handbook.
MOSQUITOES OF MEDICAL IMPORTANCE 11

probably of little or no importance in the United States, it

is not included in the keys.

References : 331, 349.

Dengue.—The last extensive epidemic of this disease in

the United States occurred in 1922 and 1923 and was preceded by a considerable increase in the abundance of Aedes

aegyftL This outbreak was centered in Texas and Louisiana,

where 500,000 to 600,000 cases were reported. An epidemic

occurred in Florida in 1936. A few cases are reported every

year throughout the Southeastern States.

References: lß2, 256.

Encephalitis.—Several strains of this disease occur in the

United States and parts of Canada. Western equine, eastern

equine, and California encephalitis viruses have caused much

injury to livestock, and these three, together with St. Louis

encephalitis virus, incidentally have attacked humans

throughout the United States. For the distribution of these

viruses, see the preceding section. Epidemiological evidence

points most strongly to Culex tarsalis as the vector of western

equine virus. This mosquito has been incriminated as the

vector of St. Louis encephalitis virus as well. Several mosquito species have been found infected in nature with the

eastern equine strain, but studies have not yet definitely

pointed out a specific vector of this virus.

References : 200, 2S7.

Filariasis.—This disease was probably introduced into the

United States by slaves from endemic centers in Africa. The

only known focus of infection in America north of Mexico

was found in 1915 in Charleston, S. C. The disease persisted

for several years but is now believed to have disappeared

entirely.

Reference: 159.

MIDDLE AMERICA

West Indies

(Fig. 5)

The West Indies, which include the Bahamas, the Greater

Antilles, and the Lesser Antilles, extend from Florida to the

north coast of Venezuela. They separate the Gulf of Mexico

and the Atlantic Ocean from the Caribbean Sea. The

Greater Antilles on the west end of the chain include Cuba,

Jamaica, Hispaniola, and Puerto Rico, and the Lesser Antilles include principally the Leeward and Windward Islands

and Trinidad.

BAHAMA ISLANDS

Physical Features.—The Bahamas consist of about 700

islands and numerous reefs and cays. This island group

stretches for 750 miles east of Florida and parallel to Cuba.

Mangrove swamps and ponds are abundant throughout the

islands. The Bahamas have a subtropical climate, with mild

winters and an annual mean rainfall of about 50 inches. The

rainy season is from May to October.

Reference: 301.

Malaria.—This disease is not present in the Bahamas.

Reference: 67.

CUBA AND JAMAICA

Physical Features.—Cuba has a greater land area than all

the other islands in the West Indies combined. It stretches

700 miles in an east-west direction and has largely level or

gently rolling ground suitable for large-scale agriculture.

Cuba has three mountain ranges. The highest is located on

the south shore at the east end of the island ; the other two

lie in the center and on the west end. The low coast, containing marshes and bottlenecked bays, is margined by

numerous coral islands and keys. The climate is temperate,

with two seasons, a rainy period from May to October and

a cool, dry season from November to April. The rainfall

averages about 50 inches annually and increases inland.

Jamaica lies 90 miles south of Cuba and is predominantly

mountainous, the main range running in an east-west direction. A few alluvial plains extend along the south and west

coasts, which have many fine bays.

Reference: 301.

Malaria.—Epidemics rather than endemics characterize

this disease in Cuba, whereas the reverse is largely true in

Jamaica. Malaria is present chiefly in the low-lying plains

and is seldom encountered above 1,000 feet.

In Cuba Plasmodium vivax and P. falciparuTn transmit

malaria almost to the exclusion of P, molariae. In Jamaica

falciparum by far outnumbers other species, and malariae

accounts for up to 10 percent of malaria; vîvaœ is almost

nonexistent.

Anopheles alhimanus^ the most important vector in the

Caribbean area, is the principal vector in Cuba and Jamaica.

It is primarily a species of the coastal lowlands, and its

numbers decrease rapidly with an increase in altitude. The

abundance of albimanus is closely correlated with the intensity of malaria in these islands.

The role of Anopheles crucians has not been evaluated

thoroughly, but it is strongly suspected of being a vector,

at least in Cuba.

Neither Anopheles aquasalis nor A, bellator is found in

Cuba or Jamaica.

Reference: -61.

HISPANIOLA

Physical Features.—Hispaniola is composed of two

political independencies, Haiti in the western one-third and

the Dominican Republic in the eastern two-thirds. The

island itself lies in the main chain of the West Indies just

east of Cuba. It is predominantly mountainous, with several wooded ranges that enclose deep fertile valleys where

most of the people live. Its coastal lowlands have an average annual temperature of about 77° F., with little seasonal

change. There are two main rainy periods, one from April

to June and the other in October and November.

Reference: 301.

Malaria.—Hispaniola has the highest malaria rate in the

West Indies. In the Dominican Republic periods of high

endemicity alternate with several years of l©wer intensity.

The principal parasite is Plasmodium falciparum, Plasmodium vivax and P. malariae cause the remaining cases;

vivax is predominant in most localities.

As in the other islands. Anopheles albimanus is the most

important vector. Since it prefers low coastal areas where

the rain makes temporary sunlit pools, it is most numerous

during and shortly after the rainy season. The endemic

malaria transmitted by this species is not so severe as that

supported by many other vectors, notably Anopheles darlingi in South America.

If Anopheles crucians is found in Hispaniola, it must be

a rare mosquito. It has been reported from both Cuba and

Puerto Rico, but its role in Hispaniola has not been evaluated. It is probably not an important vector in this island.

Neither Anopheles bellator nor A. aquasalis is found in

Hispaniola.

References : ^7, 292.
12

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE WEST INDIES

[AREA INCLUDED: BAHAMAS, GREATER ANTILLES, AND LESSER ANTILLES]

FULL-GROWN LARVAE ^»

PALPS LONG

ANTENNAE

FEMALES

\_MALES HAVE BUSHY ANTENNAE (

PATTERN ON HIND TARSUS

SEGMENT 5 DARK AT BASE-^^

PATTERN ON PALP

AND DO NOT BITE'\

_ ANOPHELES

— ALB/MANUS

PATTERN ON 2ND SEGMENT

OF HIND TARSUS

DARK AREA 1/3 OR MORE

ANOPHELES

AQUASALIS

ZZ AND PANGEL/

( ON LY AQUASAUS IMPORTANT; '

FEMALES CANNOT BE SEPARATED ;

-PROBOSCIS

SEGMENT 5 ALL DARK ^^

ANOPHELES

BELLATOR

PATTERN ON PALP

ALL DARK

_ ANOPHELES

" CRUCIANS

PALPS SHORT PATTERN ON THORAX

-« AEDES

"~ AEGYPTI

ADULT MOSQUITO WITH PARTS LABELED

"°*%<---WING
MOSQUITOES OF MEDICAL IMPORTANCE 13

PUERTO RICO

Physical Features.—Puerto Rico is the easternmost

and smallest of the Greater Antilles. A central mountain

range extends across its entire length; additional mountains are in the northeast and southeast. The coastal plain

along the northern shore is about 10 miles wide; that on

the south is narrower because of the abrupt rise of the mountains. The rainfall, scarcer on the south and west coasts

than on the north and east, averages about 60 inches annually, with little seasonal variation in temperature.

Reference: 301.

Malaria.—Although this disease is practically absent from

the central mountainous part of Puerto Rico, it is widespread on the south and east coasts.

Plasmodium vivax infections are prevalent during the first

half of the year and F. falciparunih during the second half.

As a result of irrigation, which was needed because of the

light rainfall. Anopheles albimanus has excellent breeding

grounds. This mosquito is present the year around. It

appears in small numbers through the dry season and increases to a low peak of abundance at the beginning of the

rainy season and to relatively large numbers at the height of

the rainy season in October, November, or early December.

It is the only vector of malaria in Puerto Rico that requires

extensive control measures.

Anopheles aquasalis and A, hellator are not present in

Puerto Rico.

References : 67, 216.

LESSER ANTILLES

Physical Features.—The Leeward Islands extend southeast from Puerto Rico to the Windward Islands and consist

principally of the Virgin Islands, Guadeloupe, St. Eustatius,

Saba, St. Martin, Antigua, St. Kitts, Nevis, and Montserrat.

The last four islands are part of the volcanic chain of the

Lesser Antilles and the others are of coral limestone. Nearly

all the islands are immensely fertile, with a rainy season from

August to January ; the annual rainfall is 40 to 60 inches.

The Windward Islands continue the submerged mountain

chain south and east and consist principally of Martinique,

Dominica, St. Lucia, St. Vincent, and Grenada. All the

islands have mountainous backbones, with extremely fertile

valleys and some level land. The temperature is equable,

with little seasonal variation, but the rainfall is rather variable.

Trinidad lies 10 miles north of Venezuela's Orinoco Delta

and is geologically a part of the South American mainland.

It is crossed from east to west by three low ranges of hills

covered largely by virgin forests. The indented coast is

swampy. The principal rainy period lasts from June to

December, with 50 to 120 inches of rainfall annually.

Reference: SOI.

Malaria.—As in most of the Caribbean area, malaria is

primarily a coastal disease in the Leeward and Windward

Islands and in Trinidad.

Pldsmodimrb falciparum malaria predominates, and P.

vivax is a parasite of secondary importance. Plasmodium

mAolariae accounts for few cases in most localities. The disease is rarely found above 1,000 feet.

In most of the islands except Grenada, St. Lucia, Martinique, and Trinidad, Anopheles aïbimanus is the principal

vector, and its distribution is closely associated with malaria.

Its habits are essentially the same as in the Greater Antilles.

Anopheles aquasalis is found in Trinidad and north to

Antigua and is the only vector in many of the islands within

this range. It is almost entirely a coastal species and appears

to prefer brackish water, though it is not restricted to this

habitat. Its ability to transmit malaria to man is probably

limited by its zoophilism.

Anopheles hellator is largely restricted to Trinidad, where

it transmits an out-of-doors malaria, principally in the

county of St. Andrew. This species is unusual in that its

larvae grow in epiphytic bromeliads (see p. 115) in cacao

plantations.

Anopheles hom^unculus^ a species not included in the keys,

is found in the Aripo Valley of Trinidad, where it contributes

to "bromeliad malaria" transmission.

Anopheles crucians is not present in the Lesser Antilles.

References : 67, 86.

Anopheles Species in the West Indies

"^aïbvmanus

\aïbitarsis

apicimacula

^aquasalis

átropos

^hellator

^^ crucians

\daTlingi

eiseni

grabhamii

'^homunculus

See footnote 6, p. 10.

maculipes

mediopunctatus

neomaculipalpus

nimbus

\oswaldoi

\pseudopunctipennis

pseudopvMctipennis

\pwiictifmacula

f quadrimaculatus

rangeli

\vestitipennis

Yellow Fever.—Positive mouse-protection tests have been

performed in Cuba, Jamaica, and Puerto Eico, but no outbreaks of urban yellow fever have been recorded in the West

Indies since 1914, when cases were reported from Trinidad.

Personnel of the Eegional Virus Laboratory of the Eockefeller Foundation found cases of jungle yellow fever in

Trinidad early in January 1954, and over 100 human cases

and many monkey deaths were observed during that year.

Evidence of a specific mosquito vector was not obtained,

but the virus was isolated from pools of mosquitoes containing species of Haemagogus collected at ground level. Since

1954 an islandwide immunization campaign has been carried

on in Trinidad, and measures have been undertaken against

Aedes aegypti on a large scale.

References : 7, 8, IS, 19, 20, 108, 109, 256.

Dengue.—In the Bahamas and Jamaica dengue has never

been reported. However, in Cuba and Hispaniola the disease

is endemic in nonimmune persons, and small local outbreaks

have occurred. Southward the disease becomes more intense

and is prevalent in most of the Lesser Antilles. Both types

of dengue are present in Trinidad, and surprisingly high

rates of immunity to this disease have been found in humans

there. In all these areas Aedes aegypti is the widely distributed vector.

References : 7, 8, IS, 19, 20,109, 256.

Encephalitis.—This disease is not present in mo^t of the

West Indies. Eastern equine encephalitis virus attacks animals in Cuba and both animals and humans in the lower

valley of the Yaque del Norte in northwestern Dominican

Republic. The only two known fatal cases of Venezuelan

equine encephalitis in humans occurred in Americans in

Trinidad. Little is known of the epidemiology of this disease, and the vector is not known with certainty. Recent

investigations do not show any activity of the Venezuelan

virus in Trinidad. However, St. Louis encephalitis virus infections have probably occurred in that island, and serum

from one donkey has given a positive reaction to the eastern

equine virus. In Trinidad two strains of Ilheus virus have

been isolated from wild-caught forest mosquitoes, among

which large numbers of Psorophora and Aedes species were

present. High immunity rates were found in humans in the

low, heavily forested areas of the island.

References : 43, 109, 2^6, 251.
14 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

Filariasis.—Culex pipiens quinquéfasciatus is present

throughout most of the West Indies and is the principal

vector of Wuchereria hancrofti wherever that parasite is

found. Thorough studies have been made in some islands,

but none in others. The disease is not a public-health problem in Cuba, Hispaniola, or Jamaica.

In Puerto Rico, where more intensive studies have been

made, infection rates up to 13 percent have been found.

C, p, quinqué fasciatus is most numerous in the hot, moist

coastal areas; it is rare at altitudes above 2,000 feet and in

the hot, dry south and southwest. There are no endemic

centers in Puerto Rico and the smaller islands, because people move so freely from one locality to another.

Wuchereria hancrofti has been present in St. Croix for

several years. St. Thomas, 40 miles away, has little or no

filariasis where quinquéfasciatus is abundant. Infectious

stages of the parasite have been observed in this mosquito the

year around in St. Croix, where up to 25 percent of the females have been found infected. In St. Croix Wuchereria

hancrofti completes its development to the infectious stage

in Anopheles alhimanus^ but that mosquito is not regarded

as an efRcient vector of filariasis.

In Antigua filariasis incidence has been estimated at about

38 percent in the native population, being highest in the

central clay area, lowest in the northeast, and intermediate

in the southwest volcanic area.

References: 166, 266-269, 300, 306, 323.

Mexico

(Figs. 6 and 7)

Physical Features.—Triangular Mexico has a 2,000-mile

northern frontier along the boundaries of California, Arizona, New Mexico, and Texas. Gradually narrowing, the

country extends south to Guatemala, where its east coast

swings northeastward to form the peninsula of Yucatan,

which partially separates the Gulf of Mexico from the Caribbean Sea. Both of these bodies of water bound Mexico on

the east; the entire west coast fronts the Pacific Ocean.

Lying immediately west of Mexico proper is the narrow

peninsula of Lower California, separated from Mexico by

the Gulf of California.

Three-fourths of Mexico is covered by mountain ranges.

The two most important are the roughly parallel Sierra

Madre Oriental and Sierra Madre Occidental. The former

is an extension of the Continental Divide and runs southward near the east coast from the northern border to below

the level of Mexico City ; the latter extends southward near

the west coast. Each range is separated from its bordering

water by coastal strips never exceeding 200 miles in width.

Between these ranges is the central plateau or highlands,

with broken mountains and deep valleys. From an altitude

of about 4,000 feet in northern Coahuila and Chihuahua,

these highlands rise to 8,000 feet in the interior farther

south. The narrowest part on the isthmus is high and contains the Sierra Madre del Sur range, which extends into

Guatemala close to the east coast.

Throughout Mexico the rainy season lasts from June

through September. The Lower California coasts, Sonora,

Sinaloa, and northern Tamaulipas are driest and depend on

irrigation for agriculture. Farther south in the tropical

jungle of Tabasco in the inner curve of the Yucatan Peninsula, the rainfall reaches 120 inches annually. All coastal

lowlands and the mountainous country up to 3,000 feet in

most of the Yucatan Peninsula are hot and tropical ; from

3,000 to 6,000 feet in the central highlands the climate is

more seasonal; and the area above 6,000 feet, containing

many of Mexico's principal cities, has hot days and cold

nights.

Reference: 301.

Malaria.—The dry Northern States of Lower California,

Sonora, Chihuahua, Coahuila, Durango, and Zacatecas have

little malaria. The arid country supports little mosquito

breeding, except in a limited way along the more verdant

river courses, where the disease is occasionally endemic, with

annual rates of almost 50 per 100,000 in some populations.

In the highland States to the south—Aguascalientes, Guanajuato, Queretaro, Mexico, and Tlaxcala—there is a slightly

higher rate of endemic malaria. The States of Hidalgo,

Puebla, and Morelos and the western coastal region report

extremely high rates. Along the gulf coast south and east

from Tamaulipas malaria incidence increases considerably,

and the States of Tabasco, Campeche, and Yucatan are

highly malarious, as is the more inland San Luis Potosí.

The same is generally true of the Pacific coast, with Jalisco

and Michoacán lightly malarious and an area of Oaxaca

heavily so.

Most malaria in Mexico is caused by Plasmodivmh vivax-^

P, falciparum and P. malariae^ the former the more abundant, cause most of the remaining cases.

In the Xochimilco-Mixquic area of the valley of Mexico

just south of Mexico City, Anopheles aztecus is the important if not the sole vector of benign malaria. This area

is almost 7,500 feet above sea level. The species is commonly found in houses, bites man freely, and can be readily

infected in the laboratory. It has been responsible for over

a thousand cases of malaria annually. It has been found in

various localities throughout central Mexico, but its importance there has not been studied adequately.

Anopheles pseudopunctipennis pseudopunctipennis is another highland species, appearing in the absence of other

Anopheles species during the dry season in the highlands of

Mexico. It is a dangerous vector, probably accounting for

most of the malaria m Lower California and along the Pacific slopes of Mexico. With aztecus^ it transmits all malaria

in the Mexican highlands. The closely related Anopheles

pseudopunctipennis franciscanus is found in Low^er California and extreme northwest Mexico, but it is not an important species. Past records of nontransmission probably

refer to that variety rather than to typical pseudopunctipennis pseudopunctipennis.

Anopheles alhimanus is found on both the gulf and Pacific

coasts of Mexico. It is present along the gulf coast from

the United States border to Quintana Eoo, and it is responsible for the high rate of endemic malaria there. Along

the Pacific side its range extends northward from Central

America to Sonora.

The range of Anopheles darlingi extends from South

America to the lower part of Mexico, at least as far north as

Tabasco. In Central and South America it is particularly

dangerous and probably contributes greatly to the endemic

malaria in extreme southern Mexico.

Anopheles freehorni^ the principal vector in the Western

United States, extends its range south into the north-central

plateau of Mexico and Lower California. In these two areas

south of the border it probably plays some part in malaria

transmission, although it is replaced by pseudopunctipennis

pseudopunctipennis to the south.

The range of Anopheles crucians extends south from the

United States into Tamaulipas and beyond. It is restricted

to the gulf side of Mexico, where it is suspected as a vector

because of its anthropophilic tendencies.

Like crucians,, Anopheles quadrirrmculatus also is found in

northeast Mexico, but its range is much more limited.

Knowledge of its role in malaria transmission is confused by

the presence of alhimanus.

References : ^i, 61,110, 334, 335.
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE

^ FULL-GROWN LARVAE Bi"*—

15

MEXICO

FIG. 6

SIPHON ABSENT

SPIRACULAR

APPARATUS

•OUTER CLY PEALS

IS OR MORE

BRANCHED

INNER CLYPEAL6

• PALMATES ON 2ND

ABDOMINAL SEGMENT

MODERATELY

DEVELOPED

POORLY

DEVELOPED

CLOSE TOGETHERN

PROTHORACIC HAIRl

ANOPHELES

QUADRIMACULATUS

ANOPHELES

AZTECUS

HAIR "O" ON 4TH

ABDOMINAL SEGMENT

ANOPHELES

CRUCIANS

HAIR"0" ON 4TH

ABDOMINAL SEGMENT

ANOPHELES

FREEBORNI

AND PUNCTIPENNIS

I ONLY FREEBORNI IMPORTANT; ]

\ LARVAE CANNOT BE SEPARATED ]

SIPHON PRESENT.

PLUS ALL CHARACTERS

LISTED BELOW

WITH

SHORT

SIDE

HAIRS

SPIRACULAR APPARATUS

WITH SHORT

HAIRS AT

POSTERIOR

END .

PROTHORACIC HAIRS 1 & 2

HAIR i

BRANCHED

FROM SIDES;

BASES FUSED

ANOPHELES

ALBIMANUS

ANOPHELES

DARLINGI

PROTHORACIC HAIR 1 SPIRACULAR APPARATUS

COMB SCALES I

IN SINGLE

ROW

ONLY ONE

TUFT

PER SIDE

+ / 2E

ABDOMINAL SEGMENT 8

COMB SCALES NOT ON A

PLATE«, SURFACE BARE

PROJECTIONS

AT POSTERIOR

END ^

ANOPHELES

H PSEUDOPUNCTIPENNIS

PSEUDOPUNC TIRENNIS

ANTENNA COMB SCALES

AEDES

AEGYPTI

ANOPHELES LARVA

PROTHORACIC

HAIRS IS. 2

PLEURAL BRISTLES

ON UNDER SIDE WITH A BASAL\

HAEMAGOGUS

EQUINUS

471779 O—59-
16 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

-^ «tf FEMALES ^m

MEXICO

[MâLES HAVE BUSHY ANTENNAE (

PALPS LONG

ANTENNAE

.PROBOSCIS

PATTERN ON HIND TARSUS

PATTERN ON PALP

^F

) AND DO NOT BITE'\

ONE LIGHT SPOT ON FRONT MARG»N~^

FIG. 7

TWO LIGHT SPOTS ON FRONT MAflG[N_^

ANOPHELES

CRUCIANS

AND BRADLEY!

/"ONLY CÑUCtANS IMPORTANT:

\ FEMALES CANNOT BE SEPARATED ,

ANOPHELES

PSEUDO PUNC T/PENN/S

PSEUDOPUNC TIPENNIS

ANOPHELES

QUADRIMACULATUS» FREEBORNI,

AND AZTECUS

( ALL IMPORTANT; FEMALES CANNOT BEN

\ SEPARATED WITH CERTAINTY )

PATTERN ON PALP

_ ANOPHELES

" ALBIMANUS

LIGHT SPOT AT CROSS VEIN SHORTER

^ THAN PRECEDING DARK SPOT

iiiiiHiJiltó — ANOPHELES

M — DARLING/

^CROSS VEIN

^^,

FRINCEPALPS SHORT ERN ON CLAWS ON PATTERN ON

lORAX MID LEG HIND FEMUR

TOOTHEOv^ LIGHT ATx^"V-^^

iLLIC BLUE ^ ^ TIP -^ ^^b/ >

Í GREEN ^^^ ^^^

ADULT MOSQUITO WITH PARTS LABELED

THORAX^^

_ HAEMAGOGUS

— EQUINUS

__ AEÛES

— AEGYPTI

*albimanus

apicimacula

\aquasalis

argyritarsis

átropos

^astecus

harberi

hradleyi

\ crucians

\cruzi

"^darlingi

eiseni

evansae

fausti

*freehomi

gahaldoni

hectoris

fhomu/nculu^

intermedius

Anopheles Species in Mexico

neivai

neomaculipalpus

oGcidentalis

parapunetipennis

guatemalensis

parapunctipennis

parapunetipennis

pseudopunctipennis

franciscanus

"^ pseudopunctipennis

pseudopunctipennis

fpunetimacula

punctipennis

f quadrimaculatus

rondoni

\vestitipennis

loalkeri

xelajuensis

See footnote 6, p. 10.

Yellow Fever,—No classical urban yellow fever has been

known in Mexico since 1910-20. Aedes aegyptii^ present,

probably in small numbers, in port areas into which the disease can be introduced. Jungle yellow fever has not yet

been reported in humans, but mouse-protection tests have

revealed positive monkey sera in Chiapas. Authorities agree

that jungle yellow fever might extend into the tropical rain

forest of the gulf coast from Central America (see p. 20).

Haemagogus equinus and potentially susceptible monkeys

are present in a continuous belt of tropical rain forest extending northward on the gulf side. This mosquito is found

in deciduous forest, mangrove, and thorny scrub as far north

as Brownsville, Tex. Haemagogus mesodentatus mesodentatus and H, m. gorgasi are dominant species in the tropical

rain forest. The range of the former extends along the gulf

slope as far north as southeastern San Luis Potosí, that of

the latter along the Pacific slope to southern Sinaloa near

Mazatlán. AU three Haemagogus species are good laboratory vectors of yellow fever, but none have been found with

natural infections.

Salethes chloropterus^ an inhabitant of tropical rain and

deciduous forests on the gulf and Pacific slopes, probably
MOSQUITOES OF MEDICAL IMPORTANCE 17

maintains sylvan yellow fever during seasons that are unfavorably dry for the Haemagogus species.

References : 1S2,155, 256, 330.

Dengue.—This disease occurs throughout Mexico, but only

a few cases are reported every year.

Reference: 256.

Encephalitis.—The virus of western equine encephalitis

may appear along the Eio Grande in the northernmost parts

of Mexico. As far as is known no humans have contracted

the disease. Eastern equine encephalitis virus attacks both

animals and man along the gulf coast to about latitude

20° N. Its exact distribution in this area is uncertain. Culex

tarsalis^ the suspected vector of both viruses, is present in

Mexico.

References: 200, 257.

Filariasis.—No information is available.^

Central America

(Figs. 8 and 9)

Physical Features.—Central America includes British

Honduras, Guatemala, Honduras, Salvador, Nicaragua, and

Costa Rica. British Honduras, the northernmost country in

Central America, is about 170 miles long, with an indented

and swampy eastern coast on the Caribbean Sea. Most of

the northern half is tropical, low, and flat, but high grasslands and the Maya and Cockscomb Mountains rise in the

southeast. A dry season from mid-February to mid-May

alternates with a rainy season, which brings most of the 80

inches of rainfall annually.

To the west is Guatemala, which touches both the Caribbean Sea and the Pacific Ocean. It is crossed from east to

west near its southern border by the Sierra Madre range,

with peaks from 11,000 to 13,000 feet. Several ranges that

enclose high plateaus above 4,000 feet extend northward

from the Sierra Madre. The Pacific slopes are steep and the

coast is narrow, but to the north and west the slopes decrease

more gradually in altitude. The coastal plains and the entire

Peten Department below 1,500 feet are hot and humid. Uplands from 3,500 to 10,000 feet are subtropical to temperate.

A rainy season from May to October brings most of the 45

inches of rain annually on the plateau and part of the 200

inches in some lowland sections.

Honduras to the south and east of Guatemala also touches

both the Caribbean and the Pacific and is almost entirely

mountainous except for the narrow coasts, river valleys, and

the "mosquito coast" between the Patuca and Coco Eivers in

the northeastern corner. Honduras is crossed by the Cordillera, the main mountain range that rises to 7,000 feet in the

higher southern part near Salvador. These mountains send

off spurs northward to enclose highlands and fertile valleys.

In the tropical coastal lowlands in the north the climate is

hot and humid with heavy rains, but farther south and higher

the climate is more seasonal, with a rainy season from May

to December.

Salvador has a 170-mile Pacific coastline and is bounded by

land on the other sides. From east to west the country is

crossed by two volcanic ranges rising to almost 8,000 feet.

Between these ranges is a 2,000-foot plateau, which contains

the principal cities and cultivated areas. The Lempa River

Valley north of the mountains is an area of tropical savannas

^ Signifies the disease Is not present or it may occur in a region,

but no records of it have been found in the Uterature.

flanked by Honduras highlands. About 70 inches of rain

fall annually, most of it in a rainy season from May to October. As in other Central American countries, the climate is

tropical in the lowlands and more temperate in the mountains

and plateaus.

Nicaragua has a swampy, key-fringed, 350-mile Caribbean

coast and 200 miles of Pacific shoreline. The Cordillera

traverses the country from northwest to southeast, rising

to 7,000-foot peaks. Several long spurs branch out in a northeast direction and slope gradually to the wide Caribbean

lowlands. West of the principal mountain divide an extensive depression contains the large lakes Managua and

Nicaragua. Between these lakes and the Pacific Ocean is a

narrow chain of low volcanoes (2,000 to 4,000 feet). The

tropical eastern Caribbean lowlands receive almost 200 inches

of rain annually and the more temperate western highlands

about 60 inches, mostly from May to December.

Costa Rica straddles the isthmus. It has a 630-mile Pacific

shore and a 130-mile Caribbean coast, which is marshy, alluvial, and lined by lagoons. From northwest to southeast

the Cordillera cuts across the country in several nearly

parallel ranges, which enclose at 3,000 to 5,000 feet a fertile

basin containing three-fourths of the white population. The

jungles of the Caribbean coast receive about 120 inches of

rain annually, whereas the Pacific coast is cooler and drier.

The mountain plateau has dry (December to April) and

rainy seasons.

Reference: 301.

Malaria.—Throughout the countries of Central America

malaria is a dangerous disease. Itis the predominant disease

in low, moist tropical areas. The extensive mountainous

backbone of the isthmus connecting Mexico with South

America restricts this lowland area to the Pacific and Caribbean coasts. Malaria is also present in some highland areas

but is not common above 4,000 to 5,000 feet.

Except in Salvador and Honduras, Plasmodium vivax is

the predominant parasite, P. falciparum is next in importance, and P, malariae is the least common. P, malariae is

important and abundant only along the Pacific slope of

Costa Eica.

Anopheles albimanus is present along both the Pacific and

Caribbean coasts and has been found inland in Guatemala at

an altitude of 4,500 feet, a considerable deviation from its

normal habitat of low tropical areas. In Costa Rica it is the

dominant species, especially in the lowlands, and it has been

observed at elevations of 2,500 feet in drying riverbeds. It

is one of the most important vectors in the Caribbean area.

Anopheles darlingi is prevalent throughout almost all

the Central American countries. The adults feed readily on

man and are often taken in large numbers in houses. In

South America this is one of the most important vectors of

malaria, and it contributes greatly to endemic malaria in

Central America as well.

Anopheles pseudopunctipennis pseudopunctipennis has

been collected in all the Central American countries. It is

not so important in Costa Rica as in the highlands of Guatemala farther north, where it consistently enters habitations

and bites man readily.

Anopheles punctimacula is widely distributed in all the

Central American countries. The adults visit human habitations in large numbers and have been observed feeding on

animals.

Anopheles vestitipennis is a comparatively rare mosquito.

Nevertheless, at certain times and places in British Honduras

it probably plays a role in malaria transmission because of

its habit oi entering houses and biting man.

References : 67, 209, 292.
18 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

[EXCLUDING PANAMA]

CENTRAL AMERICA

FULL-GROWN LARVAE FIG. 8

SIPHON ABSENT

POSTSPIRACULAR HAIR^

SPIRACULAR

APPARATUS

PALMATE HAIRS

LEAFLETS SMOOTH

SPIRACULAR APPARATUS

WITH LONG

HAIRS AT

POSTERIOR

END^

- ANOPHELES

~ DARLINGI

PROTHORACIC HAIRS l&a

HEAD AND PROTHORAX OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS

/ OUTER I HIND

CLYPEALS 1 CLYPEALS

WITHOUT

LONG

HAIRS AT

POSTERIOR

END ^

r OUTER CLYPEALS

LEAFLETS NOTCHED

v^

A TO 14

BRANCHED:

BRANCHES

FORKED

HAIR 1 .

BRANCHED

FROM SIDESÎ 1<

BASES FUSED

LATERAL HAIRS OF

ABDOMINAC SEGMENTS 4 & 5

». ANOPHELES

" ALB/MANUS

ANOPHELES

PUNCTIMACULA

ANOPHELES

VESTiTIPENNIS AND GABALDONI

f ONLY VESTITIPENNIS IMPORTANT; \

\ LARVAE CANNOT BE SEPARATED /

PROTHORACIC

HAIR 1

ANOPHELES

ZZ PSEUDOPUNCT/PENN/S

PSEUDOPUNC T/PENN/S

L

SIPHON PRESENT,

PLUS ALL CHARACTERS

LISTED BELOW

COMB SCALES |

IN SINGLE ^>i

ROW

ABDOMINAL SEGMENT 6

COMB SCALES NOT ON A

PLATE:SURFACE BARE

COMB SCALES ON A PLATE«.

SURFACE WITH SHORT

HAIRS

ANTENNA COMB SCALES

AEDES

AEOyPTI

SADDLE

SADDLE HAIR DOUBLE

HAEMAGOGUS

EQU/NUS

_ HAEMAGOGUS

— SPEGAZZINII FALCO
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE — CENTRAL AMERICA

[ EXCLUDING PANAMA ]

^ FEMALES ^*

[^MJALES HAVE BUSHY ANTENNAE [^1^0^ ) AND DO NOT û/TE'\

19

FIG. 9

PALPS LONG

ROBOSCIS

PATTERN ON HIND TARSUS

THREE DARK AREAS ON FRONT MARGIN

LIGHT SCALES ON

4- LOWER HALF OF

WING YELLOWISH

(NOT WHITE)

,VÈIN 6 WITH NUMEROUS LIGHT AND

/ DARK AREAS

ANOPHELES

PUNCTtMACULA

LATERAL

, SCALE

I TUFTS

ABSENT

TWO LIGHT SPOTS ON FRONT MARGIN

iiiiiii'iiiii'iiiinnii

VESriTIPENNIS )

LOWER PART OF

VEIN 6

ENTIRELY DARK

lilJlloP

( WING OF PSEUDOPUNCTIPENNIS )

LIGHT SPOT AT CROSS VEIN SHORTER

THAN PRECEDING DARK SPOT

_ ANOPHELES

" DARLINGI

PATTERN ON PALP

— ANOPHELES

" ALBIMANUS

_ ANOPHELES

— VESTITIPENNIS

ANOPHELES

PSEUDOPUNCTIPENNIS

PSEUDOPUNC TIPENNIS

VPALPS SHORT

PATTERN ON TOP

OF THORAX

METALLIC BLUE

OR GREEN

PATTERN ON MID LEG PATTERN ON HIND FEMUR If

ONE OR MORE

STERNOPLEURAL

SETAE PRESENT =

HAEMAGOGUS

MESODENTATUS

AND SPEGAZZINII FALCO

(ONLY SPEGAZZINII FALCO \

IMPORTANT; FEMALES

CANNOT BE SEPARATED
20 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

Anopheles Species in Central America

intermedius

koTThpi

neivai

neoTnaculipalpus

\oswaldoi

parapunctipennis

guatemalensis

parapunetipennis

parapunetipennis

"^pseudopv/netipennis pseudopunctipennis

'^pwnctimaeula

triannulatus

'^vestitipennis

xelajuensis

^aTbimanus

\aïbitarsis

anomalophyllus

apicvmaevda

\aquasalis

argyritarsis

\'beTlator

\crueians

\crum

^darUngi

eiseni

evansae

gaialdoni

hectoris

jhomunculus

See footnote 6, p. 10.

Yellow Fever.—Urban yellow fever, transmitted by Aedes

aegypti^ was common in Central America prior to 1925.

From 1925 to 1948 the Central American countries were free

of the disease, but from 1948 to 1951 a wave of jungle yellow

fever swept northward through Panama (see p. 28), and

from 1951 to 1954 it invaded Costa Rica, Nicaragua, and

Honduras. This disease is not known to occur in Guatemala.

In the first three countries cases of jungle yellow fever were

reported in the tropical rain forest of the Caribbean slope,

where wild monkey hosts support the disease.

Haemagogus spega^zinii faleoïs one of the most commonly

encountered vectors of yellow fever under suspicion, but its

range extends northward only to the north coast of Honduras

east of the Ulua Valley. Haemagogus equinus^ a species

ranging from Panama to Texas, is also suspected. More recently Haemagogus mesodentatus mesodentatus and H, m.

gorgasi have been implicated in yellow fever transmission.

All these Haemagogus species have transmitted yellow fever

virus in the laboratory, but none have been found naturally

infected.

Sahethes chloropterus is believed to transmit the virus

from one monkey host to another during dry periods that

are unfavorable for other mosquitoes.

References : 1S2, SSO, S49.

Dengue.—This disease is apparently not present in Guatemala, Salvador, or British Plonduras, but cases of dengue

appear farther south. The presence of endemic dengue in

Honduras is in considerable doubt, but sporadic cases thought

to be of this disease have occurred there and in El Salvador.

The extent of dengue in Nicaragua and Costa Rica is in

dispute. Aedes aegypti is the vector wherever the disease

appears in Central America.

References : 6, 9,10,12, 15,17, 256.

Encephalitis.—No information is available on the distribution of the encephalitis viruses in Central America.

Reference : 251.

Filariasis.—Wuchereria hancrofti is present in an endemic

focus around Limón, Costa Rica. Surveys have shown this

parasite to be present in as high as 15 percent of the native

populations in certain areas of the Caribbean coast and considerably less (1 percent) along the Pacific. Culex pipiens

quinquefasciatus^ the probable vector, is present in all these

areas.

Reference: 7^.

Panama

(Figs. 10 and 11)

Physical Features.—Panama is 450 miles long and connects the North and South American Continents. It varies

from 32 to 113 miles in width between the Caribbean Sea on

the north and the Pacific Ocean on the south. Its heavily

indented southern coast is a narrow lowland fringed with

numerous keys and islands, where tides rise as high as 20

feet. The Caribbean side, bordered by narrow lowlands, has

much weaker tides. The isthmus is traversed from east to

west by several almost parallel mountain ranges, which

rise to over 11,000 feet near the border of Costa Rica. In

the center of Panama at the Panama Canal Zone, these

mountains descend to 1,500-200 feet but rise to the east in

the Cordillera de San Bias and the Serranía del Darién

ranges, which are very close to the northern Caribbean coast

and extend into Colombia.

The climate is tropical and subtropical except in a few

high upland areas. The Caribbean side of the mountains

receives up to 150 inches of rain annually, most of which

comes in the wet season from mid-May to mid-December,

but the Pacific side receives only about 70 inches per year.

The Canal Zone is a strip of land, 40 miles wide, lying

across the center of Panama in a northwest to southeast

direction. The climate there is tropical, with higher rainfall

on the Caribbean than on the Pacific side.

Reference: ^Wl.

Malaria.—As in Central America, malaria in Panama is

widespread and important. It is practically absent from the

higher mountains, but along the coast and in the Canal Zone

itself malaria is still highly endemic in spite of consistent

control measures.

Most malaria in Panama is caused by Plasmodium vivax,

Plasmodium falciparum and P. malariae are responsible for

the remaining cases; the former parasite is the more common.

Anopheles aTbimanu^ is probably the most important vector of malaria in Panama, as in most of the Caribbean area.

It is abundant throughout the year, but during the dry season it is mostly restricted to large bodies of water. When

the rise of water level during the rains causes protective vegetation to break up, alhimanu^ tends to seek coastal breeding

places.

Anopheles pseudopu/nctipennis péeudopunctipennis is one

of the most common mosquitoes in Panama. Despite its

prevalence it is not regarded as an important species in that

country. The larvae are most abundant during the dry season and inhabit drying stream beds in open, sunlit locations.

This mosquito is present well into the rainy season, but heavy

rain in these areas flushes the larvae out of breeding places

and the adults become scarce.

Anopheles alhitarsis is a dangerous species in parts of

South America and for this reason is included in the keys.

No records exist of albitarsis females attacking man in Panama, even in the immediate vicinity of intense larval populations. This species may be composed of several races, a

possible explanation for its zoophilism in some areas of this

country. It is usually found breeding during the last half

of the dry season in a restricted area of the western arm of

Gatun Lake.

Anopheles oswaldoi is a rare mosquito, but it is believed to

be an important vector in several areas of Panama. Breeding in well-shaded, fresh-water jungle pools, it does not commonly come in contact with man. The females will occasionally bite and transmit a certain amount of malaria to

man when the latter invades their jungle haunts.

Anopheles aquasalis is present only on the Caribbean side

of Panama and commonly breeds in salt-marsh water, although the larvae have been found in fresh water as well.
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE -

FULL-GROWN LARVAE]

21

PANAMA

FIG. 10

r OUTER CLYPEALS PROTHORACIC HAIR 1

SIPHON ABSENT

SPIRACULAR

APPARATUS

SPINES ON ANAL

SEGMENT

/

WITH

SHORT

HAIRS

WITH

LONG

HAIRS

BRUSHLIKE

ANOPHELES

= PSEUDOPUNCTIPENNIS

PSEUDOPUNC TIPENNIS

'PRO THORACIC HAIR 1

BRANCHED

FROM

BASE;

LEAFLETS

WIDE AND

FLATTENED

INNER CLYPEALS

WIDELY

SEPARATED — ANOPHELES

"" ALBITARSIS

PROTHORACIC HAIRS 1 AND 2

BRANCHED

FROM

SIDES

_ ANOPHELES

ALBIMANUS

'OUTER CLYPEALS

T

MAIN STEM

FORKED

ANOPHELES

OSWALDOI

MAIN STEM

SINGLE

_ ANOPHELES

"- AQUASALJS

LATERAL HAIRS OF SPINES ON ANAL SEGMENT

ANOPHELES LARVA

PROTHORACIC

HAIRS 1 & 2

7)'===f=^(ííl^''>C ABDOMINAL

«t là ^ír^=~= =*• SEGMENTS

4 AND 5

ANOPHELES

PUNCTI MACULA

SIPHON PRESENT,

PLUS ALL CHARACTERS

LISTED BELOW

COMB SCALES |

IN SINGLE ^Ji

ROW "^ ^

.SIPHON

ONLY ONE

TUFT

PER SIDE

\ PECTÉN SPINES

--A EVENLY SPACED

VENTRAL BRUSH

WITH AT LEAST^4

SEPARATE HAIRS\

' ABDOMINAL SEGMENT 8

COMB SCALES NOT ON A

PLATE} SURFACE BARE

COMB SCALES ON A

PLATE; SURFACE WITH

SHORT HAIRS

COMB SCALES

SADDLE

LENGTH OF SIPHON FOUR

TIMES WIDTH AT BASE

AEDES

LEUCOCELAENUS CLARK/

AEDES

AEGYPTI

SADDLE HAIR TWO

BRANCHED.

_ HAEMAGOGUS

~" EQUINUS

HAEMAGOGUS

SPEGAZZINII FALCO

HEAD AND PROTHORAX OF

ANOPHELES \L.f<RMfK

ANTENNAL

HAIR INNER / CLYPEALS

OUTER

CLYPEALS

PROTHORACIC HAIRS 1, 2 AND 3
22

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE PANAMA

-M«^ FEMALES —

[MâLES HAVE BUSHY ANTENNAE (

PALPS LONG

ANTENNAE

PATTERN ON HIND TARSUS

LAST THREE SEGMENTS WHITE EXCEPT

BASE OF 5f" . _

PATTERN ON PALP Í

AND DO NOT BITE^

PATTERN ON 2ND SEGMENT

OF HIND TARSUS

FIG.

DARK AREA 1/4 OR LESS AS

LONG AS SEGMENT

3^

DARK AREA 1/3 OR MORE AS

LONG AS SEGMENT

_ ANOPHELES

~ OSiVALDO/

ANOPHELES

AQ(/ASAL/S, ANOMALOPHXLLUS,

AND STRODE I

ONLY AQUASAUS IMPORTANT«, \

^ FEMALES CANNOT BE SEPARATED / (

LIGHT SPOrAT CROSS VEIN LONGER THAN

fc£REÇEplNG 0ARK_SPOT_

IJiiiiiillÜ!!!!! ANOPHELES

ALBIMANUS

ABDOMINAL SEGMENT 1

LAST THREE SEGMENTS ENTIRELY WHITE

WITH ROWS OF

, WHITE SCALES

T ON VENTRAL

SIDE

«. ANOPHELES

— ALBITARSIS

SCALES AT BASE

OF WING

WITH NUMEROUS LIGHT AND DARK BANDS 3 DARK AREAS ON FRONT MARGIN

fiiiiiiiiifn

LIGHT SCALES

ON LOWER

HALF OF WING

YELLOWISH

«. ANOPHELES

" PUNCTIMACULA

PART OF VEIN 6 ENTIRELY DARK

ANOPHELES

=: PSEUDOPUNCTIPENNIS

PSEUDOPUNC TIPENNIS

PALPS SHORT

PATTERN ON TOP

OF THORAX

If

METALLIC BLUE

OR GREEN

CLAWS ON MID LEG

Of PSEUDOPUNCTIPSNNIS )

•PATTERN ON HIND FEMUR

DARK AT TIP ONE OR MORE

STERNOPLEURAL

+ SETAE PRESENT

/ SEE ENLARGED\

DRAWING FOR

\ LOCATION /

HAEMAGOGUS

EQUINUS

HAEMAGOGUS

= SPEGAZZINII

FALCO

SIDE OF THORAX

STERNOPLEURAL SETA

PATTERN ON SIDE

OF THORAX

CROSS VEIN

FRINGEAEDES

= LEUCOCELAENUS

^ CLARK/

AEDES

AEGYPTI

ANTENNA jHORAX

ADULT MOSQUITO WITH PARTS LABELED

WING
MOSQUITOES OF MEDICAL IMPORTANCE 23

In spite of the large numbers present in some localities, this

species is not attracted to man in this country.

Anopheles punctimacula is regarded by many workers as a

malaria vector in Panama. Evidence for this is somewhat

conflicting. The females are often abundant in areas where

malaria is hyperendemic and where other known vectors are

absent, but many times no infections can be demonstrated in

wild-caught mosquitoes under these circumstances.

Anopheles darlingi may have some importance in Panama,

because it is an efficient vector over most of its range.

References : 45, 67, 291, 292, 296.

Anopheles Species in Panama

"^alhimanus maculipes

]aTbitarsis mediopunctatus

anomalophyllus neivai

apicimacula neomaculipalpiis

\aqu(isalis nimbus

argyritarsis "^oswaldoi

\beUator parapunctipennis

hraziliensis peryassui

\ crucians '^pseudopunctipennis pseudo-

\criuBÍ puMctipennis

\darlingi '^punctirrmcula

eiseni rangeli

evansae squarrdfemuT

fausti strodei

grabhamii triannulatus

\horminculu8 \vestitipennis

kompi xelajuensis

See footnote 6, p. 10.

Yellow Fever.—The difficulties encountered in the construction of the Panama Canal and the conquest of yellow

fever gave rise to one of the most significant chapters in the

history of arthropod-borne diseases. Classical yellow fever,

transmitted by Aedes aegypti^ is now practically extinct in

Panama because of careful and persistent control measures.

Jungle yellow fever, on the other hand, was discovered in

1932 in eastern Panama and may have been present there

for a number of years. An outbreak causing five deaths

occurred in the Pacora district in 1948, and soon afterward

an epidemic was noted proceeding from east to west. It

swept through the tropical rain forest on the Caribbean slope.

In July and August 1949 it reached the Chagres district, and

in April and June 1951 the disease reached Almirante and

began to invade Costa Kica (see p. 20).

Haemagogus equinus^ H, spega^Binii falco, and H, mesodentatus mesodentatus are primarily arboreal mosquitoes.

All of them live in the tropical rain forest of the Caribbean

slope, and all have transmitted yellow fever virus in the

laboratory, although none have been found with natural

infections.

Aedes leucocelaenus clarhiis common in the rain forest, and

naturally infected mosquitoes of this species from South

America have been able to transmit the virus in the laboratory.

Sahethes chloropterus is believed to transmit the virus on

both the Caribbean and Pacific slopes during times of the

year that are too dry for the maintenance of other species.

References : 132,155,330.

Dengue.—In 1933 three small outbreaks of dengue occurred at low altitudes near the coast of Panama at the end

of the rainy season when artificial containers used by Aedes

aegypti were filled. No substantial number of cases have

been reported since this time.

Reference: 21.

Encephalitis.—The virus of eastern equine encephalitis is

present in central Panama, and doubtful human cases of the

disease were recorded in 1946. Nothing is known about the

vectors.

Reference: ^57.

Filariasis.—No information is available.

SOUTH AMERICA

Northwest South America

(Figs. 12 and 13)

Physical Features.—Northwest South America includes

Venezuela, Colombia, Ecuador, and Peru. They form an

arc around the northwest corner of the South American

Continent. Venezuela has a north coast along the Caribbean

Sea. The Orinoco Eiver rises in the forested southeast

Guiana Highlands, flows northwest, north, and east through

the center of the country, and forms a wide, swampy delta

opposite Trinidad. Most of the north-central part of Venezuela consists of some tropical jungle and the llanos, an area

of savannas. A spur of the Andes, the Cordillera Oriental,

runs from east to west across Venezuela near the coast. Seaward from this range is the low, tropical coast. The rainy

season in Venezuela lasts from May to November.

The eastern three-fourths of Colombia consists of humid,

torrid plains. The western fourth contains three great mountain chains, which reach northward from Ecuador, one peak

being almost 19,000 feet high. The valleys between these

mountains contain most of the country's population and

agriculture. Savannas coextensive with those of Venezuela

lie directly east of the Cordillera Oriental. All the southern

half of Colombia east of the mountains is densely forested.

The alluvial coasts are rain soaked.

Ecuador, the smallest of the four countries, fronts on the

Paciñc Ocean. The central north-south Andes chain consists of two parallel ranges of extremely high (up to 20,500

feet), active and inactive volcanoes. About 60 percent of the

country's population lives in a 7,500- to 9,000-foot tableland

between these mountain ranges. A western agricultural lowland lies between the Pacific and the Andes, and an eastern

jungle and savanna lowland are in the Amazon basin.

Peru is the southernmost of the four countries and has a

1,400-mile Pacific coast. The 10- to 40-mile-wide coastal

strip receives less than 2 inches of rain annually, but almost

all river valleys and deltas are intensively cultivated. About

60 percent of Peru's native population lives in large upland

basins of the Andes at altitudes of 10,000 feet and over. The

land descends eastward from these uplands into the jungle

of the upper Amazon basin known as the Montaña, which is

low, humid, and covered by lush vegetation and savannas.

References: 301,349.

Malaria.—Throughout Ecuador and the coastal areas of

Colombia Plasmodium falciparum predominates, although

P, vivax and P, malariae are present in smaller numbers.

Elsewhere in Colombia and throughout Venezuela and Peru

vivax is the principal parasite. In Venezuela malariae is

sometimes abundant.

The southeastern jungle of Venezuela is sparsely populated

but represents one of the country's large centers of endemic

malaria. Anopheles darlingi is the vector in this area and in

recent years has moved near the coast. Another endemic

center is found at altitudes up to 3,280 feet on the southeastern

savanna plateau of Venezuela. An additional endemic center in northwestern Venezuela is characterized by large epidemics attributed to Anopheles albimanus, especially during

periods of heavy rain. This species breeds heavily in flooded

ricefields, and in the vicinity of Maracay it is restricted
24 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE — NORTHWEST SOUTH AMERICA

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

[AREA INCLUDED: VENEZUELA, COLOMBIA^ ECUADOR, AND PERUJ

FULL-GROWN LARVAE

FIG. 12

SIPHON ABSENT

SPIRACULAR

APPARATUS

OUTER CLYPEALS PROTHORACIC HAIR 1

WITH

LONG

HAIRS

WITH

SHORT

HAIRS

OUTER CLYPEALS

MAIN STEM

FORKED

MAIN STEM

SINGLE

ANOPHELES

— PSEUDOPUNCTIPENNIS

PSEUDOPUNC rtPENN/S

— ANOPHELES

" OSWALDOI

ANOPHELES

-- AQUA SALISy BEN ARROCHl^

AND NUNEZ-TOVARI

/ONLY AQUASAUS IMPORTANT; LARVAE N

V CANNOT BE SEPARATED WITH CERTAINTY /

PROTHORACIC HAIR 1 PROTHORACIC HAIRS lAND 2

_ ANOPHELES

"" ALBlMANUS

_ ANOPHELES

— DARUNGI

ANOPHELES LARVA

PROTHORACIC

HAIRS 1&2 PLEURAL BRISTLES

ABDOMINAL

SEGMENTS

4 AND 5

SPIRACULAR

APPARATUS

PROTHORACIC HAIRS 1

DISTANCE BETWEEN BASES OF HAIRS

APPROXIMATELY SAME AS THEIR LENGTHS

— ANOPHELES

" ALB/TARS/S

LATERAL HAIRS OF PLEURALS OF 1*" GROUP

ABDOMINAL SEGMENTS 4 AND 5

SPINES ON ANAL SEGMENT

ANOPHELES

PUNCT/MACULA

rABDOMINAL SEGMENT 8

SIPHON PRESENT,

PLUS ALL CHARACTERS

LISTED BELOW

COMB SCALES

IN SINGLE ^>i^-

ROW "*^

ONLY ONE

TUFT

PER SIDE

COMB SCALES NOT ON A

PLATE} SURFACE BARE

ANTENNA SADDLE

SADDLE HAIR TWO BRANCHED

r COMB SCALES

COMB SCALES ON A PLATE;

SURFACE WITH SHORT HAIRS

LENGTH OF SIPHON

TWO TIMES

WIDTH AT BASE

A EDES

LEUCOCELAENUS

AEDES

AEGYPTI

HAEMAGOGUS

-Z. SPEGAZZINII SPEGAZZINII,

SPEGAZZINII FALCO, AND CAPRICORNH

(ALL IMPORTANT^ ONLY MALES CAN BE SEPARATED*)

HAEMAGOGUS

EQU/NUS

HEAD AND PROTHORAX OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

HIND

CLYPEALS

PROTHORACIC HAIRS 1,2, AND 3
MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE— NORTHWEST SOUTH AMERICA

25

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

[AREA INCLUDED: VENEZUELA, COLOMBIA, ECUADOR, AND PERUj

'^^m FEMALES»»^

[M^LES HAVE BUSHY ANTENNAE (ü^^^) ^f^D ^O ^OT B/TE^ FIG. 13

PALPS LONG

ANTENNAE

PROBOSCIS

PALPS SHORT

PATTERN ON HIND TARSUS

LAST THREE SEGMENTS WHITE

EXCEPT FOR BASE OF 5^" .^^

PATTERN ON PALP

PATTERN ON 2ND SEGMENT

OF HIND TARSUS

DARK AREA 1/4 OR LESS

AS LONG AS SEGMENT

ANOPHELES

OSiVALDO/

^>

_ ANOPHELES

DARK AREA 1/3 OR MORE

AS LONG AS SEGMENT AQUASALIS AND RELATIVES

L

IPLIGHT SP^T AT CROSS VEIN LONGER

VTHAN PRECEDING DARK SPOT

LAST THREE SEGMENTS ENTIRELY

WHITE

LIGHT SPOT AT CROSS VEIN SHORTER

^THAN PRECEDING DARK SPOT

, LATERAL SCALE

^ • TUFTS PRESENT

LIGHT SPOT AT CROSS VEIN LONGER

NJHAN PRECEDING DARK SPOT

ABDOMINAL SEGMENT 1

WITH ROWS OF

WHITE SCALES

ON VENTRAL

SIDE

WITH NUMEROUS LIGHT AND

DARK BANDS

THREE DARK AREAS ON FRONT MARGIN

'lililí

SCALES ON BASAL

HALF OF WING

_ ANOPHELES

"" ALB/MANUS

_ ANOPHELES

— DARLINGI

_ ANOPHELES

~ ALBITA RSIS

LIGHT SCALES

ON LOWER _ ANOPHELES

HALF OF WING "" PUNCTIMACULA

YELLOWISH

LOWER

ALL DARK -<

+ M

• PATTERN ON THORAX CLAWS ON MID LEG

METALLIC BLUE

OR GREEN

PART OF VEIN 6 ENTIRELY DARK

ANOPHELES

^, = PSEUDOPUNCTIPENNIS

PSEUDOPUNCTIPENNIS

PATTERN ON HIND FEMUR

ONE OR MORE HAEMAGOGUS

I STERNOPLEURAL — SPEGAZZINIt SPEGAZZINII^

"•" SETAE PRESENT "" SPEGAZZINII /v4¿CC|,AND CAPRICORNII

/ALL IMPORTANT; ONLY MALES CAN BE ^

\ SEPARATED /

/SEE ENLARGED )

DRAWING FOR

\ LOCATION I

\

HAEMAGOGUS ^

H EQUINUS, ANDINUS^ AND

PANARCHYS

/ONLY EQUINUS IMPORTANT;

FEMALES CANNOT BE

\ SEPARATED

AEDES

LEUCOCELAENUS

-. AEDES

"~ AEGYPTI

THORAX^

ADULT MOSQUITO WITH PARTS LABELED

CROSS VEIN
26 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

solely to this habitat. It is progressing farther and farther

inland along the main highways. Ä, darlingi is present still

farther inland throughout this area, and a great deal of

malaria occurs throughout its range in the absence of albimanus. Anopheles pseitdopunctipennis psevdopunctipennis

transmits malaria in the valleys of the western Andes. Anopheles aquasalis, a coastal species, plays a role in malaria

transmission along the southern part of the Paria Peninsula

directly west of Trinidad, but elsewhere in Northwest South

America it appears to be unimportant. Anopheles albitarsis is believed to be an important vector in parts of Venezuela,

but the extent of its range has not yet been adequately determined.

Colombia's southeastern flatland has a large amount of

malaria for its scattered population. This is carried by

darlingi^ whose range extends northward into Panama and

westward into the low mountain valleys. A, aïbimanus inhabits the Pacific coastal flatlands, but malaria incidence is

lower there than in the northern coastal area, where it is

present with not only darlingi but Anopheles punctimacula

as well. In the Cauca Valley in the west-central mountains

pv/nctimacula is the only vector in the presence of high

malaria incidence. This area represents the greatest single

malaria problem with which punctimacula has ever been

associated. In the upper Magdalena Valley to the north

and east pseudopunctipennis plays an important part in malaria transmission. Anopheles oswaldoi^ primarily a forest

mosquito, is believed to be important in several places in

Panama, and it is included in the keys because it may be importantin Colombia as well.

The distribution of malaria in Ecuador is little known.

East of the Andes the vector is probably darlingi, A. p,

pseudopunctipennis is the principal mosquito in several

mountain valleys in this country, where it is the principal

vector associated with epidemic and highly endemic malaria.

A, alhimanus reaches its southernmost range on the Pacific

side of the continent in Ecuador. Near the coast at Guayaquil extremely high spleen rates, presumably due to the

activity of alhimanu^s^ have been found.

In Peru, largely a mountainous country, pseudopunctipennis is the most important vector. It is present not only

in the higher mountain valleys where most of the country's

population lives but in the lower coastal areas as well. This

species has been found as high as about 7,000 feet in the presence of endemic malaria, and it is responsible for occasional

epidemics at these altitudes. The other important vector in

Peru, punctimacula^ is restricted to coastal areas and is found

from sea level to about 2,500 feet along the west slope of the

Andes. It is associated with the irrigation of sugarcane and

rice. A, darlingi is the vector in the flatlands east of the

Peruvian border, but little is known about its importance

directly to the west.

References : 59, 67,100, 292, 296.

Anopheles Species in Northwest South America

acanthotorynus

"^alhimanus

'^albitarsis

annulipalpis

apicimacula

"^aquasalis

argyritarsis argyritarsis

argyritarsis sawyeri

hamhusicolus

^hellator

henarrochi

holiviensis

'braziliensis

^cruciœns

\cruBÍ

cruzi laneanus

^darlingi

eiseni

evansae

-ßuminensis

gilesi

gomezdelatorreyi

guarao

\homunculus

korrvpi

lanei

lutzi

mcucuLipes

mattogrossensis

mediopunctatus

neivai

neomaculipalpus

nimbus

nuneztovari

oihetorakras

^oswaldoi

parvus

peryassui

pseudomaculipes

pseudopunctipennis

hifoliata

pseudopunctipennis

levicastilloi

"^pseudopunctipennis pseudopunctipennis

pseudopunctipennis

rivadeneirai

* punctimjacula

punctipennis

rangeli

rondoni

shannoni

squarmjermir

thomasi

triannulatus davisi

triannulatus triannulatus

vargasi

^vestitipennis

See footnote 6, p. 10.

Yellow Fever.—Epidemics of urban yellow fever, normally supported by Aedes aegypti^ no longer occur in Northwest South America because that species is no longer present.

The disease has been reported from nearly all the coastal areas

of this region in the past. Jungle yellow fever, on the other

hand, occurs in a large part of this region, especially at lower

altitudes. Mouse-protection tests performed before 1940

were positive in all four countries. There are continual sporadic outbreaks, each affecting considerable numbers of

people.

Of all the Haemagogus species that can be vectors, Z?^.

spegazzinii falco is the most important in this region.

Haemagogus spegazzinii spegazzinii^ H, capricomii^ and H.

equinus^ as well as Aedes leucocelaenus^ have been taken in

much smaller numbers. For details on the biology and importance of these species, see the next section of this handbook.

References : 40, U<^^ UU ^^0,230,256, SIS.

Dengue.—In 1942 and 1943 a disease resembling dengue

occurred along the Magdalena Eiver in Colombia, but its

vector wa^i never actually determined. Sporadic cases,

thought to be dengue, are reported every year. Although

dengue has occurred in Venezuela in the past, no cases have

been reported since 1920. The disease has never been reported from Peru.

References : 11, Ih 23, 26,256.

Encephalitis.—The virus of Venezuelan equine encephalitis has been found in an endemic area that includes parts of

Venezuela, Colombia, the western half of Ecuador, and Trinidad. It is highly infectious, especially to man, but the

extent to which this virus is responsible for the disease in

man and animals is not known. Doubtful human cases in

Ecuador and confirmed human cases at Espinal, Colombia,

were recorded in 1951 and 1952, respectively. No vector has

been definitely implicated. A large area of central Peru is

believed to contain animals infected with western equine

encephalitis virus. For additional information, see page 3.

References: 246,257.

Filariasis.—No information is available.

Brazil and the Guianas

(Figs. 14 and 15)

Physical Features.—Brazil and the Guianas comprise

about one-half the total area of South America. Most of

Brazil lies south of the Equator and has approximately

4,600 miles of Atlantic coastline. Brazil is essentially a vast

central plateau bordered by lowlands, the largest of which

is the Amazon basin in the northern half. The Guiana

Highlands form its northern rim. The basin, which extends

into eastern Colombia, Ecuador, and Peru, has uniform high
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

SIPHON ABSENT

POSTSPIRACULAR HAIR.

SPIRACULAR

APPARATUS

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE — BRAZIL AND THE GUIANAS

27

SIPHON PRESENT,

PLUS ALL CHARACTERS

LISTED BELOW

• SIPHON

FULL-GROWN LARVAE FIG. 14

PROTHORACIC HAIR 1

1

BRANCHES FLATTENED}

RISING FROM BASE

'OUTER CLYPEALS

WITH SHORT

HAIRS

WITH LONG

HAIRS; MAIN \

STEM

SINGLE

INNER CLYPEALS

WIDELY

SEPARATED

PROTHORACIC HAIRS 1

DISTANCE BETWEEN BASES OF HAIRS

APPROXIMATELY SAME AS THEIR LENGTHS

ANOPHELES

ALBITARSIS

ANOPHELES

AQUASALIS AND RELATIVES

/ ONLY AQUASÁUS IMPORTANT; ONLY ^

\ MALES CAN BE SEPARATED J

PALMATE HAIRS

WITH LONG

HAIRS; MAIN .

STEM

FORKED

ANOPHELES

OSWALDOI

SPIRACULAR APPARATUS

BRANCHES HAIRLIKE;

RISING FROM BASE

«. ANOPHELES

"" DARLÍNGI

BRANCHES HAIRLIKE;

RISING FROM SHAFT

ANOPHELES

ALBIMANUS

BARE; OVER

ONE-HALF

AS LONG

AS INNER

CLYPEALS

'POSTSPIRACULAR HAIRS

ANOPHELES

BELLATOR

ANOPHELES LARVA

»ROTHORACIC

HAIRS

PLEURAL BRISTLES

UNDER SIDE WITH A BASALV

SPINE-J.IKE SHEATH

-POSTSPIRACULAR

HAIRS

ABDOMINAL SEGMENT 1

PALMATES PRESENT,

COMB SCALES |

IN SINGLE

ROW

ONLY ONE

TUFT

PER SIDE

•ABDOMINAL SEGMENT 8

COMB SCALES NOT ON A

PLATE; SURFACE BARE

COMB SCALES SIPHON

HAIR TUFT

TWO BRANCHED,

^^^^m^»- +

ANOPHELES

HOMÚNCULOS, CRU ZI

AND ANOPLUS

(ONLY HOMUNCULUS AND CRUZl

IMPORTANT;LARVAE CANNOT

BE SEPARATED

AEDES

LEUCOCELAENUS

COMB SCALES ON A

PLATE; SURFACE WITH

SHORT HAIRS

— AEDES

AEGYPTl

— HAEMAGOG'IS

~" EQUINUS

HAEMAGOGUS

SPEGAZZINII SPEGAZZINII,

SPEGAZZINII FALCO, AND CAPRICORNI I

(ALL IMPORTANT; ONLY MALES CAN BE SEPARATED)

HEAD AND PROTHORAX OF

ANOPHELES LARVA

ANTENNAL

INNER

CLYPEALS

HIND

CLYPEALS

PROTHORACIC HAIRS
28

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE-BRAZIL AND THE GUIANAS

—^—^-i^^^B FEMALES H^i^^——

\_MÂL£S HAVE BUSHY ANTENNAE [\¡¡¡^^^r ) AND DO NOT ö/7"£"] FIG. 15

PALPS LONG

ANTENNAE

-PROBOSCIS

PATTERN ON HIND TARSUS

PATTERN ON PALP

r PATTERN ON 2ND SEGMENT

OF HIND TARSUS

DARK AREA 1/4 OR LESS AS = ''''oíwAÍ'bo^l m^ iNINU LONG AS SEGMENT

1^

/ONLY OSWALDOI IMPORTANT; \

FEMALES CANNOT BE I

\ SEPARATED /

DARK AREA 1/3 OR MORE AS AMrsouci re

LONG AS SEGMENT — Am^t'nc.l-C^

— AQUASALIS AND RELATIVES

lONVf AQUASAUS IMPORTANT;\

ONLY MALES CAN BE I

\ SEPARATED /

LIGHT SPOT AT CROSS VEIN LONGER THAN

VPRECEDING DARK SPOT

„I ».X - ANOPHELES

p''''i!j'iiiii;;:'« ALBIMANUS

ABDOMINAL SEGMENT 2

LATERAL SCALE

TUFTS PRESENT! \

LATERAL SCALE

TUFTS ABSENT

(WING OF.ALBIMANUS)

LIGHT SPOT AT CROSS VEIN SHORTER THAN

< PRECEDING DARK SPOT

^ '''l||||||||ji!if ANOPHELES

DARLINGI

F DARLINGI )

ABDOMINAL SEGMENT 1

, WITH ROWS OF

+ WHITE SCALES ON

VENTRAL SIDE

_ ANOPHELES

— ALBITARSIS

VEIN 3 WITH TWO SMALL LIGHT SPOTS

IN BASAL HALF--S.—.

^ ' li» « ANOPHELES

- HOMUNCULUS

( WING OF C/)UZI )

VEINS 4 AND 6 ALL DARKv

_ ANOPHELES

— BELLATOR

SIDE OF THORAX

STERNOPLEURAL

SETA ^

/^PATTERN ON THORAX CLAWS ON MID LEG

PALPS SHORT

(WING OF BELLATO/})

rPATTERN ON HIND FEMUR

rr

AEDES

LEUCOCELAENUS

- AND LEUCOPHOEBUS

(ONLY LEUCOCELAENUS \

IMPORTANT; FEMALES \

CANNOT BE SEPARATED I

WITH CERTAINTY /

ONE OR MORE HAEMAGOGUS

STERNOPLEURAL CAPRICORNIO SPEGAZZINII SPEGAZZ/NU,

, SETAE PRESENT -- SPEGAZZINII FALCO, KWKi ALBOMACULATUS

/SEE ENLARGED \ / ALL IMPORTANT EXCEPT ALBOMACULATUS; \

DRAWING FOR I FEMALES CANNOT BE SEPARATED )

\ LOCATION / ^

— HAEMAGOGUS

" EQUINUS

AEDES

AEGYPTI

ADULT MOSQUITO WITH PARTS LABELED
MOSQUITOES OF MEDICAL IMPORTANCE 29

temperatures and is the world's largest area of tropical climate and forest. It has an annual rainfall of about 60

inches. Much of the land near the coast and the poorly

drained flood plain of the Paraguay Kiver constitute another

area of tropical rain. The Mato Grosso is the central Brazilian highland separating the Amazon from the Eio de la

Plata. This rolling upland is interrupted by several mountain ranges as high as 9,000 feet. It has a tropical climate,

with alternate wet (summer) and dry seasons. In the southernmost Brazilian States the weather is humid and subtropical. Near to and parallel with the east coast is the Serra do

Mar, which rises in peaks to almost 10,000 feet and forms a

barrier to inland traffic.

British Guiana, Surinam (Netherlands Guiana), and

French Guiana lie side by side directly north of Brazil. All

have rather similar physical features. The 5- to 30-milewide coastal belt across all three countries is the only cultivated area. To the south an extensive upland contains most

of the mineral and forest wealth. The Guiana Highlands

along the northern border of Brazil have densely wooded

plateaus and occasional savannas. The two wet seasons in

the Guianas, one from April to mid-August and one from

mid-November to February, are much less pronounced in

the interior than on the coast, and trade winds from the

northeast prevail from October to July.

Reference: 301.

Malaria.—New World malaria reaches its greatest intensity in Brazil and the Guianas. In the Guianas and the

entire Amazon River basin it is transmitted throughout the

entire year. Its curve of intensity shows little annual fluctuation. From the southern edge of the Amazon basin south

to about latitude 22° S., malaria occurs in a single pronounced annual peak in the first half of the year and is transmitted over a long period. In the rest of Brazil malaria occurs in two peaks, one rather pronounced and occurring during March and April and the other considerably lower and

occurring in September, October, and November.

Plasmodium falciparum malaria predominates in northern

Brazil. Plasmodium vivax is a more important parasite

than P, malariae in the north, the latter predominating in

the vicinity of Sao Paulo. Coastal malaria in British Guiana is caused by falcipant/m^ and vivax predominates inland ;

the reverse is true in French Guiana and Surinam.

The most efficient vector of New World malaria, Anopheles darlingi^ is widespread throughout the four countries,

principally in the lowlands some distance from the coast, but

it is also present in areas as high as 1,000 feet above sea level.

This species is not found in areas having heavily acid waters

or in brackish water along the coast. It is estimated that

malaria would become a relatively unimportant disease in

South America with the disappearance of darlingL

Anopheles aquasalis is a coastal brackish-water mosquito

that rarely is found more than 10 to 12 miles inland. It

plays a prominent role in malaria transmission in several

coastal localities of Brazil. Under certain conditions it is

surprisingly zoophilic and feeds readily on domestic animals,

a possible reason for its lack of importance in many areas.

Anopheles albimanus has a limited coastal distribution in

the Guianas. For this reason it is limited in its importance,

although it is an efficient vector where it appears in large

numbers.

Anopheles albitarsis is important in several places in Brazil. The most important is near the Sergipe and Bahia

coasts, where it has been found infected in highly malarious

areas. This endemic center extends southward along the

coastal plain where, in Espíritu Santo, the typical species

is present, and it is as important as darlingi and aquasalis^

which also are found there.

Anopheles oswaldoi has not been adequately studied. It

is a jungle mosquito and bites man out of doors. Since man

seldom enters its haunts, this mosquito may not play an important role in malaria transmission in Brazil and the

Guianas.

A particularly interesting association of man and malariabearing mosquitoes is that provided by Anopheles homunculus^ A. cru^i^ and A. bellator. These three species breed in

epiphytic bromeliads, which are parasitic plants growing on

the limbs of the tall immortelles that shade cacao plantations.

Not only do they transmit malaria by entering the houses of

the plantation workers but they commonly bite out of doors

as well. They transmit a large amount of malaria along

the central and southern coasts of Brazil. In several localities near Santa Catarina, Panana, and Sao Paulo, cr'U^i and

bellator are the most abundant Anopheles species and will

attack man both night and day.

Anopheles gambiae^ the most important vector of malaria

in tropical Africa, was introduced at one time into northeastern Brazil. This species is not included in the keys,

because it has not reappeared since its eradication in 1943.

References : 67, 11Í, 296,

Anopheles Species in Brazil and the Guianas

acanthotorynus lanei

^albimanus lutzi

^albitarsis maculipes

anrmlipalpis mattogrossensis

anoplus mediopunctatus

antimesi minor

apicimacula neivai

^aquasalis neomaculipalpus

argyritarsis argyritarsis nigritarsis

argyritarsis sawyeri nimbus

argyrothorax noroestensis

bambusicolus nuneztovari

"^bellator "^ oswaldoi

benarrochi parvvjS

boliviensis peryassui

bonnei pictipennis

braziliensis fpseudopunctipennis

canorii pseudopwnctipennis

"^crmi pseudotibiamiaculatus

"^darlingi \punctimacuLa

eiseni rachoui

evamdroi rangeli

evansae rondoni

-ßimiinensis sanctielii

galvaoi shannoni

gilesi squamifenm/r

"^homunculus thomasi

ininii tibiamaculatus

intermedius triannulatus davisi

kompi triannulatus triannulatus

See footnote 6, p. 10.

Yellow Fever.—Jungle yellow fever appears from time

to time in sporadic outbreaks in humans throughout most of

Brazil and the Guianas. Mouse-protection tests show that

immunity to this disease is widespread in humans. Urban

yellow fever, borne by Aedes aegypti^ is rare in this region,

but aegypti-hovne epidemics may start with the introduction

of jungle yellow fever into urban areas.
30 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Haemagogus spegazziniispegazzinii and H, s, falco account

for almost 80 percent of the Haemagogus mosquitoes in

Brazil. The latter is probably the more common of the two

in the Guianas, although investigations have shown that

Haemagogus aïbom^aculatus may also be important there.

Haemagogus equinus is an important vector of jungle yellow

fever in Central America, and although reports of its presence

in British Guiana, Surinam, and parts of Brazil are conflicting, it is included in the keys because of its potential importance.

Aedes leucocelaenus has been found in 18 of the 20 Brazilian

States, but it is most common in the south. In northern

Brazil it is a less important vector than spegazzinii.

References : J^0, Itl, US, 218, 219,256,313.

Dengue.—This disease, transmitted by Aedes aegypti, is

largely confined to the coastal areas of the Guianas. An outbreak of 36 cases was reported in 1936 and 10 in 1943, mostly

in newcomers. Dengue appears to be definitely endemic in

these countries. There have been no large epidemics since

1920. A few cases of dengue have been reported from parts

of Brazil bordering the Amazon Eiver.

References: 2^,16,24,256.

Encephalitis.—Two unconfirmed human cases of St. Louis

encephalitis were diagnosed in 1952 in Paramaribo, Surinam.

Antibodies were found in the immediate families and in

domestic and wild animals. The virus of eastern equine

encephalitis was found in animals in Peçanha, Brazil, in 1944.

It is not known how any of these cases originated. The

Ilheus virus was first isolated from animals in the Ilheus

area of the Brazilian coast. The vector is not known.

Reference: 257.

Filariasis.—In Georgetown, British Guiana, and Paramaribo, Surinam, Guleoa pipiens quinquéfasciatus has been

found naturally infected with Wuchereria hancrofti. In the

former locality up to 7.4 percent of quinquéfasdatus mosquitoes have been found infected. According to a clinical survey, almost the entire colored population is subject to infection. Outside these two centers little is known of the

occurrence and distribution of filariasis. A study at Belem,

a city at the mouth of the Amazon in the State of Para,

showed that quinquéfasdatus far outnumbered other mosquitoes and was the principal vector of hancrofti in the native

population.

References : 80, 225, 288.

Lower South America

(Fig. 16)

Physical Features.—Lower South America comprises all

continental South America south of Brazil and Peru and

includes Bolivia, Paraguay, Uruguay, Argentina, and Chile.

Bolivia and Paraguay are the only countries in South America without seacoasts. Most of the Bolivians live in the

Temperate Zone on an intermountain plateau at an average

altitude of 12,000 feet. It is located between the three main

ranges of the Andes, which cut across western Bolivia from

northwest to southeast. The eastern three-fourths of the

country is principally tropical lowland, covered in the northern half with the dense forest of the Amazon basin and with

savannas and scrub in the more arid southeast Chaco. These

lowlands rise westward into the higher subtropical Yungas.

The Chaco area also extends into Paraguay and Argentina.

The western half of Paraguay is entirely covered with

arid, torrid grasslands and hardwood forests. The fertile

lowlands east of the Paraguay Eiver support most of the

country's population. A northeastern highland area separates this country from Brazil.

Most of Uruguay is rolling grassland. The irregular,

lagoon-filled coastline faces south. Between it and the interior is a fertile alluvial plain. Most of the population is

concentrated in the southern half of the country.

Argentina is a 2,300-mile-long triangle at the southern tip

of the continent. The northwest end contains the semiarid

Gran Chaco, which extends north into Paraguay. The alluvial lowlands in the north and northeast receive about 60

inches of rain annually, the country's highest precipitation.

Eainfall gradually decreases westward across the plains of

the 400-mile-wide pampas. The slopes of the Andes receive

the least rain of all this area. Patagonia stretches north

from the cape about 1,000 miles in bleak, windswept plateaus.

Chile averages about 110 miles in width and extends south

from Peru between the western border of Argentina and the

Pacific Ocean. The Andes rise from within a short distance

of the coast to some of the highest peaks in the world on the

Chile-Argentine border. In the lower half of Chile this

mountain chain decreases in altitude. The adjacent coastline is tortuous, with numerous offshore islands extending to

the cape. The northern part is very dry. Central Chile is

a 600-mile-long fertile plain containing about 90 percent

of the country's population. Dense forests cover much of the

southern part.

Reference: 301.

Malaria.—In the southern two-thirds of Bolivia, the

northern one-fourth of Argentina, and all of Paraguay,

endemic malaria is widespread and shows two annual peaks

of intensity. The first of these is the highest and occurs in

March and April, although in northernmost Argentina the

disease may appear as early as January. Along the Argentina coast an interesting situation arises where the maximum

density of Anopheles occurs from December to March, but

maximum malaria occurs in June and July. The second

peak of malaria intensity is much lower and usually occurs

in September, October, and November. The southernmost

extension of malaria on the Atlantic coast is Córdoba, Argentina, and on the Pacific a point just south of Iquique, Chile,

although the disease has not appeared in the latter country

for many years.

In Bolivia and Argentina Plasmodium vivaœ is responsible

for most of the malaria. At higher altitudes in Bolivia

Plasmodium malœnae may be a problem, but elsewhere it is

sporadic. In almost all areas where malaria is present

Plasmodium falciparum is not so important a parasite as

mvax.

Anopheles pseudopunctipennis pseudopunctipennis is present in the high mountain valleys of Bolivia and northern

Argentina. It is by far the most important vector in this

region. In Bolivia it has been found in the presence of

malaria at an altitude of about 9,000 feet, and in almost all

localities within its range it transmits malaria efficiently.

Anopheles darlingi, the region's most efficient vector, invades only the vast plains of northern Bolivia, where it

causes rather large epidemics.

Anopheles alUtarsis is found in the Andean foothills, it

is not always infected when in the presence of severe malaria,

but it should be regarded as a potential vector.

Anopheles aquasalis is a coastal species, appearing as tar

south as northern Argentina and Uruguay. It is an important vector in Brazil and probably transmits malaria m

Lower South America as well.

References : ^, 50, 67, 226, SOh 3^9.
MOSQUITOES OF MEDICAL IMPORTANCE 31

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE —• LOWER SOUTH AMERICA

f AREA INCLUDED: BOLIVIA, PARAGUAY, URUGUAY, ARGENT1NA,AND CHILEJ

FULL-GROWN LARVAE ^

FIG 16

NNER CLYPEALS PROTHORACIC HAIR 1

-. ANOPHELES

"" ALBITARSIS

_ ANOPHELES

" AQUASAL/S

ANOPHELES

DARLING!

SIPHON PRESENT,

ANOPHELES

= PSEUDOPUNCT/PENNIS

PSEUDOPUNCTIPENNIS

"ABDOMINAL SEGMENT 8

COMB SCALES ON A PLATE; HAEMAGOGUS

SURFACE WITH SHORT HAIRS — SPEGAZZINII AND CAPRICORNII

I BOTH IMPORTANT; ONLY MALES CAN \

\ BE SEPARATED /

'^ COMB SCALES

COMB SCALES NOT ON A

PLATE; SURFACE BARE

« AEDES

~ AEGyPT/

HEAD OF ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS

' OUTER I HIND

CLYPEALS i CLYPEALS

SADDLE

SADDLE HAIR TWO^ BRANCHED

<21i|3||P^*- -H

[MALES HAVE BUSHY ANTENNAE (

AEDES

ZL LEUCOCELAENUS

LEUCOCELAENUS

ANOPHELES \.fKK^lK

PLEURAL BRISTLES

/ON UNDER SIDE WITH A BASAUV

V SPINE-LIKE SHEATH )

j AND DO NOT B/TE]

PALPS LONG

ANTENNAE

PATTERN ON HIND TARSUS ^

LIGHT SPOT AT CROSS VEIN SHORTER

>1 THAN PRECEDING DARK SPOT

LIGHT SPOT AT CROSS VEIN LONGER

X THAN PRECEDING DARK SPOT

_ ANOPHELES

" DARLING!

ABDOMINAL SEGMENT 1

WITH ROWS OF

4. WHITE SCALES

• ON VENTRAL SIDE

ANOPHELES

AQUASAL/S AND RELATIVES

(ONLY AOUASAUS IMPORTANT; FEMALES \

\ CANNOT BE SEPARATED /

« ANOPHELES

" ALBITARSIS

ANOPHELES

•=. PSEUDOPUNCTIPENNIS

PSEUDOPUNCTIPENNIS

PATTERN ON THORAX CLAWS ON MID LEG

STERNOPLEURAL HAEMAGOGUS SPEGAZZINII,

SETAE PRESENT ^ CAPRICORNII, AND ALBOMACULATUS

[ SEE ENLARGED DRAWING \ /ON^^^ff«d^^_'^" AND CAPRICORNU \

FOR LOCATION IMPORTANT; FEMALES CANNOT BE

\ SEPARATED '

HEAD AND THORAX

STERNOPLEURAL SETAE

AEDES

— LEUCOCELAENUS

LEUCOCELAENUS

_ AEDES

"" AEGYPTI

471779 0—59
32 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Anopheles Species in Lower South America

nimbus

noroestensis

nuneztovari

\oswaldoi

pawns

peryassui

pictipennis

pseudomaculipes

pseudopunctipennis

neghmei

pseudopunctipennis noei

pseudopunctipennis

patersoni

"^pseudopunctipennis

pseudopunctipennis

fpunetimacula

rangeli

rondoni

shannoni

triannulatus davisi

triannulatus triannulatus

\albimanus

*albitarsis

annulipalpis

antunesi

apicimacula

"^aquasalis

argyHtarsis

hamhusicolus

holiviensis

hustamentei

"^'darlingi

eiseni eiseni

eiseni geometricus

evansae

ßuminensis

lanei

lutBi

maculipes

mattogrossensis

mediopunetatus

minor

nigritarsis

See footnote 6, p. 10.

Yellow Fever.—Jungle yellow fever has five endemic centers in Bolivia, 1 in the department of La Paz and 4 in the

department of Santa Cruz. Isolated cases are commonly encountered outside these areas. It has never been reported

from July to October in this country, but when it is present

it is always associated with an increase in population of

Haemagogus species. Little is known of the distribution of

this disease in other countries of this region, although positive mouse-protection tests show immunity to yellow fever in

northern Argentina and Paraguay. Aedes leucocelaenus

leucocelaenus is found in northern Argentina and may be

responsible for the yellow fever there.

References: 60,256.

Dengue.—This disease appeared in southern Paraguay

and in the northwest corner of Argentina in 1916, but its

extent and importance there and elsewhere in the region

are little known. Aedes aegypti is common over much of the

region and is the vector of this disease.

References : 22,25, 256.

Encephalitis.—Unconfirmed cases of western equine encephalitis have been discovered in central Chile, and a large

area of central Argentina contains animals infected with

the virus. In the latter country human cases of western

equine encephalitis have been tentatively diagnosed, but no

virus has been isolated from them. The vector of western

equine encephalitis in these two countries is not known.

Reference: 257.

Filariasis.—No information is available.

AFRICA

Northwest Africa

(Figs. 17 and 18)

Physical Features.—The countries included in Northwest

Africa are Rio de Oro, Canary Islands, Madeira, French and

Spanish Morocco, Algeria, Tunisia, Libya, and northern

French West Africa south to latitude 15° N. Eio de Oro is

a barren desert tract with low hills, its entire west coast

fronting the Atlantic Ocean. It is inhabited by nomadic

tribes, and agriculture is practiced little or not at all.

The Canary Islands and Madeira are entirely of volcanic

origin and comprise several islands off Northwest Africa

in the Atlantic. They are mountainous, with high plains,

deep valleys, and deserts. The climate is mild, with little

seasonal change, and the rainy period is from October to

February.

Northern French Morocco is primarily a broad, fertile

valley, bounded on the north by the Atlantic Ocean and on

the south by the Atlas Mountains, which consist of three

parallel ranges, some peaks of which rise to over 13,000

feet. The Atlantic coast is dissected by many streams originating in the Atlas Mountains. This country has an equable

Mediterranean climate.

Sj)anish Morocco is a mountainous country bordering the

Mediterranean Sea. It represents the northernmost reaches

of the Sahara Desert and contains oases watered by rivers

flowing southward from the Atlas range. It has rainfall

ranging from 30 inches annually to less than 10 inches in

the east and southeast.

Algeria has a rocky northern coast along the Mediterranean, and the Atlas Mountains extend across its entire northern area, with a few lowlands near the coast receiving up to

50 inches of rain annually. High, semiarid, lake-filled plateaus farther south separate these mountains from the Saharan Atlas, a range that marks the northern limit of the

Sahara Desert. The Sahara comprises the rest of Algeria

to the south.

Tunisia lies across the Mediterranean just south and west

of Sicily. In the northern half are continuations of the

Atlas and Saharan Atlas Mountains. An arid plateau extends southward and is divided from the Sahara Desert

proper by an east-west depression containing shallow salt

lakes.

Libya is largely desert but has a Mediterranean coast containing a string of coastal oases receiving up to 15 inches of

rain per year. The southern limit of Libya extends deep into

the Sahara.

French West Africa is composed of a series of extremely

dry, low to moderately high tablelands and includes a large

part of the Sahara Desert proper.

Reference: 301.

Malaria.—Malaria in Northwest Africa is periodic and

is regulated by the rather cyclical rainfall of the region. A

few years of abundant rainfall may provide extensive anopheline breeding places, after which the marshy areas usually

dry up and there is little malaria transmission. Epidemic

malaria occurs not only in the wetter coastal areas but in the

Saharan oases as well.

Although both Plasmodium falciparum and P. nmlariae

are present along the coast, the former species usually predominates. Little is known of the distribution of the parasites in malarious oases.

Anopheles labranchiae lahranchiae is the principal vector

of malaria in all the countries of this region. Along the

Barbary coast from Tripolitania to Morocco this species

breeds in the fresh and saline waters of rivers and marshes.

Because of these breeding places in both wet and dry seasons,

malaria is ever present, especially near large streams. Libya

represents the easternmost limit of the range of lahranchiae^

and it is an area where rainfall diminishes far below the

point necessary to maintain the species, which is not an oasis

breeder. . i • • Anopheles sergenti and A, multicolor transmit malaria m

the desert oases, where nomads always have heavy infections

and where endemic malaria is permanent. The former is the

more important species. It usually breeds in very slowly

moving water, such as seepages, and is found from Algeria

eastward to Suez. A. nrnlticolor, on the other hand, inhabits

all kinds of brackish-water collections exposed to the sun.
MOSQUITOES OF MEDICAL IMPORTANCE 33

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - NORTHWEST AFRICA

[AREA INCLUDED: RIO DE ORO, CANARY ISLANDS^ MADEIRA, FRENCH AND SPANISH MOROCCO,

ALGERIA, TUNISIA^ LIBYA, AND NORTHERN FRENCH WEST AFRICA]

^m. FULL-GROWN LARVAE ^^-

NO SIPHON ON ANOPHELES

POSTSPIRACULAR HAIR.

SIPHON ON ALL OTHERS

FIG. 17

r LONG PLEURALS

OF 3RP GROUP

ONE

HAIRED

LONG PLEURALS

OF 2N0 GROUP

ANOPHELES

D'THAU

ANOPHELES

SERGENT/

HEAD OF ANOPHELES LARVA

OUTER CLYPEALS ANTENNAL HAIR

LONG PLEURALS

OF 2ND GROUP

ANOPHELES

LABRANCH/AE

AND RELATIVES

I ALL NOT IMPORTANT '

BUT LARVAE CANNOT BE SEPARATEDSEE KEY TO EGGS

BELOW ]

ANOPHELES LARVA

PLEURAL BRISTLES

ANOPHELES

HISPANIOLA

_ ANOPHELES

"" MULT/COLOR

_ ANOPHELES

"" SUPERP/CTUS

COMB SCALES

_ AEDES

~" AEGYPTI

NOT/V:

' NOT

// J li

m. EGGS :

FRILL COMPLETELY

AROUND EGG ^^__

INTERRUPTED

BY FLOATS

2ß. AT MIDOLC

OF EGG

AT LEAST

l/S AS LONG

AS EGG

LESS THAN

1/4 AS LONG

AS EGG^ OR

ABSENT^^^

PATTERN ON TOP

ANOPHELES

LABRANCH/AE LÀBRANCH/AE

AND MACUL/PENN/S S/CAULT/

_ ANOPHELES

SACHAROV/ AND PLUMBEUS

(M-PLUMMUS ñWEDi ONLY IN TREE HOLES AND \

Af»TiriCIAL CONTAINS«S AND IS NOT AN IMPORTANT

VECTOH OF MALARIA /
34 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE — NORTHWEST AFRICA

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

[AREA INCLUDED: RIO DE ORO, CANARY ISLANDS, MADEIRA, FRENCH AND SPANISH MOROCCO,

ALGERIA, TUNISIA, LIBYA, AND NORTHERN FRENCH WEST AFRICA]

^ FEMALES ^

\_MALES HAVE BUSHY ANTENNAE i )0§^ ) "^^^ ^^ ^^^ B/TE'\

FIG 18

LONG PALPS ON

ANOPHELES

ANTENNAE

-PROBOSCIS

WING ENTIRELY DARK, OR LIGHT ONLY AT TIPx FOUR DARK SPOTS '

1

I

+

( WINO OF LABRANCHIAE ) WINO Of SACHAPOVI )

4 OR MORE LIGHT SPOTS ON FRONT

MARGIN-REMAINDER OF WING WITH

NUMEROUS LIGHT SPOTS

NO LIGHT SPOT4 LIGHT SPOTS ON FRONT MARGIN -

REMAINDER OF WING ENTIRELY DARK

PATTERN ON PALP

^DARK AT TIP

^ LIGHT AT TIP

ANOPHELES

D'THALI

FRINGE AT WING TIP

A\. «AB.^ — ANOPHELES ALL DARK = ^.^,.^«^.^, / SACHAROVI

-;- SPOT

ANOPHELES

^Ikh?""^" = LABRANCHIAE

AND RELATIVES

I ALL NOT IMPORTANT \

BUT ADULTS CANNOT

I BE SEPARATED-SEE I

\ KEY TO EGGS /

FRONT MARGIN LIGHT AT EXTREME BASE

ll"llll llliîESiiif

ANOPHELES

MULTICOLOR

{ WING OF MULTICOLOR )

FRONT MARGIN DARK AT EXTREME BASE

ilill

( WING OF HISPANIOLA )

VEIN 3 MOSTLY DARK-^

_ ANOPHELES

~ HISPANIOLA

\ _ ANOPHELES

^ "" SERGENTI

WING OF SERGCNTI )

VEIN 3 WITH LONG LIGHT AREA,

I K*,'«.ii'W -'.M iTIfffnTMjHHB MfflHDBJf""

'i|illllllllil!r''''HIIIJII'''Wlll|lli|{j"''-«J

- ANOPHELES

"~ SUPERPICTUS

'Miirraltl

( WING OF SUPERPICTUS )

SHORT PALPS ON

ALL OTHERS

PATTERN ON

THORAX

_ AEDES

~" A EGYPTI

ADULT MOSQUITO WITH PARTS LABELED

THORAX-^

DN~1275
MOSQUITOES OF MEDICAL IMPORTANCE 35

and therefore it thrives throughout dry areas, where rapid

evaporation concentrates soil salts in breeding places.

Anopheles sacharovi^ another important vector of the

maculipennis group, is not known to live in Northwest Africa,

but its distribution and medical importance nearby justify

its inclusion in the keys.

Anopheles hispaniola is found in Algeria, Tunisia, and

Morocco north of the southernmost Atlas range. It transmits malaria in the lowlands and in the high valleys at altitudes up to 8,000 feet. Not much is known of the importance

of this species in Northwest Africa, but it is believed to have

been an accessory vector in a severe epidemic in the department of Constantine in northern Algeria. In Tenerife, Canary Islands, it is the only Anopheles species in the presence

of intense malaria.

Anopheles superpictus is found in Northwest Africa, but

its role in this region is not well known. It breeds in mountainous areas here as it does elsewhere in the Mediterranean

region.

Anopheles dHhali has a range from Algeria as far east as

Suez. Its importance in the transmission of malaria in

Northwest Africa is not known, but it is included in the keys

because it is an effective vector in Sinai.

References: 67,75,94.

Anopheles Species in Northwest Africa

Habranchiae labranchiae

\maculipennis

marteri

melanoon melanoon

\melanoon suhalpinus

^multicolor

^ plumbeus

\sacharovi

"^sergenti

^^ superpictus

algeriensis

hroussesi

\claviger claviger

coustani tenebrosus

coiùstani ziemanni

"dHhali

"fluviatilis

•gambiae

"^hispaniola

hyrcanus

\labranchiae atroparvus

See footnote 6, p. 10.

Yellow Fever.—This disease has apparently never been

reported in Northwest Africa except where tropical jungles

border its southernmost rim (see p. 46).

Reference: 25ß.

Dengue.—This disease is very lightly endemic in the

Canary Islands and along the Mediterranean and Atlantic

coasts to the inland mountains, but no epidemics have been

reported since 1900. Aedes aegypti is the only known vector

in Northwest Africa. This domestic species breeds in all

kinds of artificial containers close to man's habitations.

References: 256, Sll.

Encephalitis.—No information is available.

Filariasis.—Little is known about the distribution of this

disease in Northwest Africa. Some early observations in

Tunis implicated Anopheles algeriensis as the vector of

Wuchereria bancrofti^ but later work demonstrated that

Gulex pipiens quinquéfasciatus is the true mosquito host.

References: 302, 305.

Egypt

(Figs. 19 and 20)

Physical Features.—About 96 percent of Egypt is nonarable desert, where nomadic tribes roam. The remaining 4

percent is contained in a narrow strip of land representing

the Nile Eiver Valley. The Nile Delta area, which is about

8,500 square miles, is called Lower Egypt and contains not

only one of the world's greatest concentrations of human population but immense cotton-growing areas as well. Upper

Egypt lies south of the delta. Most of its cities and towns

are situated within the strip of fertile land that is 1 to 10

miles wide on either side of the river. Since rainfall is

scarce, the economy of the agricultural area bordering the

Nile is regulated by the level of the river, which is controlled

by its headwaters in Anglo-Egyptian Sudan and Uganda.

Floods on the Nile usually take place in August and September and are partially controlled by an immense irrigation

and dam system.

Immediately west of the Nile Valley a dry area containing

scattered oases merges with the Libyan Desert, a part of the

Sahara. In southwestern Egypt large outcroppings of rock

reach as high as 6,200 feet. Large depressions are characteristic of the northwestern parts.

The territory east of the Nile is part of the Arabian Desert

and is bisected by ravines leading to the Red Sea. A range

of mountains borders the eastern edge of Egypt along its

Red Sea coast. In places these mountains reach above 6,000

feet. Sinai, a triangle of land extending into the Red Sea

between Egypt and Israel, is practically uninhabited and

has rugged 8,000-foot mountains.

Reference: 301.

Malaria.—Malaria is mildly endemic over all Egypt. In

spite of the concentration of people in the Nile Valley, there

is little indication of intense malaria there.

Malaria caused by Plasmodium malarias is almost entirely absent in the delta area but is common in the oases ;

whereas malaria caused by P. vivax is almost entirely absent

from the oases.

Anopheles pharoensis and A, multicolor transmit most of

the malaria in Egypt. The former species, though less

abundant than multicolor^ is more widely distributed. It is

the most important vector in the Nile Delta, breeding there

in immense numbers and transmitting most of the malaria.

The adults enter houses and are most active in evening and

morning twilight. The larvae very seldom appear in the

absence of abundant vegetation and are found largely in

swamps, ricefields, and their drainage and supply systems.

A, multicolor is less common in northern than in southern

Egypt, where it breeds principally in oases. The adults

readily enter houses and bite by night. The larvae live in

small shallow pools, such as stagnant or flowing drains, un-

*ised wells, and in stagnant desert water. The amount of

malaria actually transmitted by this mosquito in Egypt is

not exactly known.

Anopheles sergenti appears throughout the country in

stagnant, slow-moving water filled with vegetation and is

the most important oasis mosquito. It is rare in the Nile

Delta but is well adapted to the desert climate and, with

multicolor^ maintains infections that are always found in

desert-inhabiting people. The adults readily enter houses

at night to feed on man. This species is a proved vector in

some areas of Egypt and Israel.

Strong epidemiological evidence points to Anopheles

dHhali as an important vector in Sinai, where it bites viciously after dark inside human habitations. The larvae

have been found in a wide variety of aquatic locations, including stagnant and fresh water.

Anopheles superpictus is included in the keys because of

its potential ability to transmit malaria, although it has

never been shown to do so in Egypt. It is usually associated

with hill or mountain country, where the larvae prefer freshwater habitats and where the adults bite readily inside houses.

Anopheles gambiae^ the most important vector of malaria

in all tropical Africa, is no longer present in Egypt, having

been eradicated from the Nile Valley. Since it easily adapts

itself to conditions in that area and since it might be reintroduced, it is included in the keys.

References : 67, 205, 216, 350.
36

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE — EGYPT

FULL-GROWN LARVAE

r ABDOMINAL SEGMENTS PROTHORACIC HAIR 1

4 TO 7

FIG. 19

-. ANOPHELES

" SERGENT/

ANOPHELES

GAMBIAE

ANOPHELES

MULTICOLOR

— ANOPHELES

— SUPERPICTUS

TOP VIEW OF HEAD OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS

/ OUTER

CLYPEALS

SANTENNA^

AEDES

AEGYPTI

ANOPHELES LARVA

PLEURAL BRISTLES

' SPINE-LIKE SHEATH )

4

ABDOMINAL

»—SEGMENTS

:- 4 TO 7

Anopheles Species in Egypt

algeriensis

claviger

Goustani tenebrosus

\gambi(Jie

\hispaniola

hyrcanus

marteri

^nmlticolor

"^pharoensis

\pretoriensis

\rhodesiensis rhodesiensis

rhodesiensis rupicolus

\rufipes

\sacharovi

'^sergenti

^superpictus

turhhudi

See footnote 6, p. 10.

Yellow Fever.—This disease is not present in Egypt, although positive mouse-protection tests in humans have been

obtained since 1940 along the Ked Sea coast.

Reference: 256,

Dengue.—This disease is normally endemic in Egypt and

flares out from time to time in epidemics. A large outbreak

occurred in Cairo in 1937, eventually affecting most of the

Nile Valley, even though it was concentrated in the Cairo

area. Aedes aegypti was presumed to be the sole vector at

that time. This species is intensely domestic and lives in all

kinds of artificial containers close to man's habitations.

References: 256,311.

Encephalitis.—^West Nile virus is widely disseminated

along the Nile Eiver, according to surveys. It is also present

in the Nile Delta area. In neither of these localities are

frank cases of encephalitis seen. This disease usually attacks

children during the summer months. The virus has^ been

isolated from Culex antennatus^ mixed pools of C, pipiens

and G, univittatus^ and from univittatus alone. The lastnamed mosquito is believed to be the important vector, since

it feeds primarily on birds, which are probably the natural

reservoir of the virus. For additional information, see

page 4.

Sindbis virus was discovered in the Nile Delta, but its

symptomatology has not yet been studied. This virus has

been transmitted experimentally by pipiens and univittatus.

For additional information, see page 4.

Humans have been found to possess immunity to Bunyamwera, Uganda S, Zika, and Semliki Forest viruses along the

lower Nile Eiver. Little is known about their transmission.

References : 251, 325, 326.
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE — EGYPT

^ FEMALES»^

[MALES HAVE BUSHY ANTENNAE ( \,\¡¡^p ) ^ND DO NOT BITE'\

37

FIG. 20

ANOPHELES

SERGENTI

ANOPHELES

SUPERPICTUS

Filariasis.—Little is known of the distribution and extent

of this disease in Egypt. In the Cairo area, where Culex

pipiens quinquéfasciatus does not appear, a study revealed

that 5.4 percent of C, pipiens pipiens were naturally infected.

Experiments showed that Wuchereria hancrofti reached the

infectious stage in the proboscis of that mosquito in about

14 days.

References: 203,298.

Anglo-Egyptian Sudan

(Figs. 21 and 22)

Physical Features.—Anglo-Egyptian Sudan is essentially a vast plain surrounded on three sides by mountains,

the Marras on the west and south and the Ethiopian highlands and the Etbai range on the east. In southernmost

Anglo-Egyptian Sudan, which contains park savannas and

patches of tropical forest, rainfall reaches 50 inches annually.

Northward the length of the rainy season and the amount of

precipitation gradually decrease. In the northwestern quarter of the country is a part of the Libyan Desert, where

precipitation north of latitude 18° N. is negligible. In the

northeastern corner is the Nubian Desert, a sandstone plateau

intersected by many steep-walled wadies.

The Nile River originates in Anglo-Egyptian Sudan and

is formed by the union of theWhite Nile and Blue Nile Eivers

near Khartoum. The country's main agricultural center is

located immediately north of Khartoum. It is irrigated by

dams and lakes of various Nile tributaries. The Kordofan

plateau south and west of Khartoum consists of an undulating

surface with isolated mountain ranges reaching an altitude

of about 4,000 feet.

Reference: 301.

Malaria.—Plasmodium malariae is rarely encountered in

Anglo-Egyptian Sudan. In the southern part Plasmodium

falciparum^ is the important parasite. Elsewhere Plasm^odium vivax or falciparum, is equally common or vivax is

slightly more prevalent.

Anopheles gambiae.^ the most important vector of malaria

in Anglo-Egyptian Sudan, as in all Africa, is widely distributed in the southern and central parts of the country and

along the Nile from Uganda to Egypt. It breeds in a wide

variety of places, principally in small pools at least partially

exposed to direct sunlight. Because such accumulations of

water as those in footprints and hoofprints and irrigation

and drainage ditches are commonly associated with man, and

because gamhiae constitutes a large proportion of mosquitoes
38 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE - ANGLO-EGYPTIAN SUDAN

IMPORTANT

A SPECIMEN MUST HAVE

ALI. CHARACTERS LISTED

FOR THAT SPECIES

FULL-CROWN LARVAE !

ANOPHELES

GAMBtAE

ANOPHELES

NILI

ANOPHELES

RUFIPES

r COMB SCALES

SIPHON ON ALL OTHERS + s

\

^,^f;f\ r ONLY I

SIPHON;:^'^- ,\ I ONE I

/ • TUFT PER I

• PECTÉN SPINES

-COMB SCALES

HEAD HAIR 6

IN SINGLE EVEN ROW Í

_ AEDES

" LUTEOCEPHALUS AND AFRICANUS

/ONLY LUTEOCEPHALUS IMPORTANT;\

\ LARVAE CANNOT BE SEPARATED )

PECTÉN SPINES

IN SINGLE EVEN ROW

mm\^

_ AEDES

"~ AEGVPTI

__ AEDES

— SIMPSONI

IN SINGLE EVEN ROW

TUFT OF SIPHON

ABOUT AS LONG AS WIDTH OF

SIPHON

AEDES

METALL/CUS

IN IRREGULAR ROW

%f»»l

HEAD HAIRS 4,5LAND 6

/ \NOT

_ AEDES

— V/TTATUS

ANAL SEGMENT

WITH

HAIRED

BRANCHES

AEDES

-- FURCIFER AND TAYLORI

/BOTH IMPORTANT; ONLY MALES\

\ CAN BE SEPARATED )

FIG. 21

HEAD OF ANOPHELES LARVA

INNER

CLYPEALS

OUTER HIND

CLYPEALS i CLYPEALS

fVNTENNA^ . / n

W^'^. ii]ilí

1'"MW^^WWWW 1nm ffW\>^

S '/ i

-

\ \ / /

\ \ y /

ANOPHELES LARVA

PROTHORACIC

PLEURAL BRISTLES

ABDOMINAL

»LATES

\ TO 7

1 HEAD OF AEDES LARVA

ANTENNA^

i ANTENNAL HAIR \Í

mi y/y

HAIR 6 -^í^-^^^méi. / mr

í

HAIR 5

DNg-1278
MOSQUITOES OF MEDICAL IMPORTANCE 39

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE — ANGLO-EGYPTIAN SUDAN

^^ FEMALES ^

[MAiLrS HAl^E BUSHY ANTENNAE Í MMÊ^^^ ) AND DO NOT BITE'\ FIG. 22

3 SHORT DARK SPOTS BEHIND MIDDLE

MARGINAL SPOT s

_ ANOPHELES

~ RUFIPES

ANOPHELES

FUNESTUS
40 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

caught indoors and feeding on man, control of this species is

of prime importance in malaria eradication.

Anopheles funestus is the second most important vector.

Its distribution is confined to many parts of southern and

central Anglo-Egyptian Sudan and along the White Nile as

far north as Jebelein. In contrast to gamhiae^ funestus

breeds in rather permanent, clear water, such as swamps,

lakes, edges of rivers, and seepage areas, almost always in

association with vegetation and shade. The year-round

character of its breeding and prevalence tends to associate

funestus with endemic rather than with epidemic malaria,

the latter being more characteristic of the more seasonal

gambiae populations. The females of funestus readily enter

houses and feed on man.

Anopheles nili is a subsidiary vector and varies markedly

in its habits. It breeds in the marginal vegetation of rivers

and streams, and population densities are usually correlated

with the seasonal rise and fall of river systems. The adults

vary from area to area in their house-haunting habits, but

naturally infected females have been found in native dwellings, where the species is important if it appears in large

numbers.

Anopheles rußpes has been found in many localities in

this country, especially along the White Nile. It usually

breeds in stagnant and semistagnant pools in partial shade.

This species is important only where it appears in large numbers. In most localities it is scarce and is often found resting

out of doors.

References : 67, 122, 352.

Anopheles Species in Anglo-Egyptian Sudan

"^nili

ohsGurus

\pharoensis

\pretoriensis

rhodesiensis rupicolus

rivuloTum

^mfipes

sergenti macniahoni

squamosus squamosus

symesi

turkhudi

löellcomei

hrohieri

cinereus

Goustani coustani

coustani ziemanni

"^funestus

"^gambiae

implexus

leesoni

maculipalpis

niarshallii gibhinsi

inarshaUii marshallii

multicolor

See footnote 6, p. 10.

Yellow Fever.—This disease has not attacked humans in

Anglo-Egyptian Sudan for several years, but jungle yellow

fever is present across the southern third of the country. This

disease is typically that of forest-inhabiting monkeys. The

monkey-to-monkey transmission cycle is believed to be maintained by Aedes vittatus and possibly by A. africanus^ mosquitoes that breed in tree holes and plant containers, inhabit

the forest canopy, and prefer monkey to human blood.

However, at ground level, especially in plantation areas, the

principal vector is Aedes simpsoni^ which also breeds in plant

containers. A. simpsoni feeds on infected monkeys either

outside or just within the forest and, having a wide host

preference, feeds on man as well, transmitting the virus in a

monkey-to-monkey cycle. Aedes metalUcus^A. luteocephalv^^

A. furcifer^ and A, taylori are believed to act in much the

same capacity as simpsoni in spreading yellow fever from

monkey to man. Aedes aegypti has been included in the

keys, since it is universally responsible for the presence of

urban yellow fever. This species is prevalent only at certain

times and localities.

References : 231-233, 256, 351.

Dengue.—Epidemics of dengue have occurred along the

valley of the Nile, but little is known about the status of this

disease.

Reference: 256.

Encephalitis.—West Nile virus occurs throughoutthe central southern half of Anglo-Egyptian Sudan (see also p. 36).

Sindbis virus has been transmitted experimentally by Culex

pipiens and C. univittatus. Its symptomatology and distribution have not yet been studied. For additional information

on this virus, see page 4.

In the east-central part of the country immunity to Japanese and St. Louis viruses has been demonstrated. However,

it is believed that these two viruses do not actually appear

anywhere in Africa. Incidence of immunity to them is low.

Furthermore, they are very closely related antigenicly to

West Nile virus, which is commonly encountered in Africa.

References : 257, 325, 326.

Filariasis.—Anopheles gamhiae transmits Wuchereria

bancrofti in the Nuba Mountains and in the southern part of

Anglo-Egyptian Sudan between latitude 4° and 6° N. and

west of longitude 30° E.

Reference: 311.

Eritrea, the Somalilands, and Ethiopia

(Figs. 23 and 24)

Physical Features.—Eritrea has an excessively hot, arid

Red Sea coast, which is narrowed in the north by the rather

abrupt rise of a 3,000- to 7,000-foot north-central plateau.

In the extreme northwest this plateau descends to the arid

Sudan Desert. The central uplands are considerably cooler

and wetter than the coastal areas.

French Somaliland consists mostly of stony desert, with

isolated plateaus and highlands. It has a deeply indented

coast opposite the southwest corner of Arabia. The climate

is torrid, most of the area receiving only 2 to 10 inches of

rain annually.

British Somaliland has a torrid coastal plain in the northeast, a mountain range about 8,000 feet high in the northwest,

and an arid plateau in the south along the borders of Ethiopia and Somalia (Italian Somaliland). The entire country

is extremely hot and dry, with rain adequate for agriculture

only in the northern foothills.

Somalia (Italian Somaliland) is a broad, arid lowland

sloping eastward to the Indian Ocean froni the Ethiopian

Ogaden Plateau. It has a dry tropical climate with two

short rainy periods, one in March and April and the other

in October and November, averaging less than 20 inches of

rain annually.

Ethiopia, the largest of the countries in this region, is an

almost isolated mountain plateau rising from the White Nile

lowland in the west. Mountains in the northern parts are

over 15,000 feet high. They descend into flat highlands to

the east and south. The Rift Valley, a large steep-walled

depression running from French Somaliland to the center

of the southwestern border at Lake Rudolph, is bounded on

the south by mountains that gradually descend to the semiarid Ogaden Plateau. The Danakil Desert is in the northeast corner. Areas of this country up to 5,500 feet are constantly hot and arid or semiarid, such as the Rift Valley and

around Ogaden and Danakil, whereas areas from 5,500 to

8,000 feet contain most of the population and receive from

25 to 80 inches of rain annually. Above 8,000 feet the temperatures are comparatively low and the rainfall is abundant.

Reference: 301.

Malaria.—This disease is present in varying amounts in

almost all watered areas of this region. Because of gradual

increases in elevation over much of this territory the temperature does not change abruptly, so that vectors become

gradually adapted to climatic changes when their populations increase. Malaria, therefore, appears in mildly endemic and epidemic form in some areas up to 7,000 feet.
MOSQUITOES OF MEDICAL IMPORTANCE 41

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE- ERITREA, THE SOMALILANDS, AND ETHIOPIA

[AREA INCLUDED: ERITREA, FRENCH SOMALILAND^ BRITISH SOMALILAND, SOMALIA, AND ETHIOPIA]

FULL-GROWN LARVAE

NO SIPHON ON ANOPHELES

SIPHON ON ALL OTHERS

r COMB SCALES

ONLY ONE

TUFT PER

SIDE, AND

ON OUTER

HALF OF

SIPHON

COMB SCALES

-f- IN ONE

REGULAR ROW

(NOT ON A PLATE)

PECTÉN SPINES

T AEDES

AEGYPTI

AEDES

SIMPSONI

__ AEDES

"~ AFRICANUS
42 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE — ERITREA, THE SOMALILANDS, AND ETHIOPIA

[AREA INCLUDED: ERITREA, FRENCH SOMALILAND, BRITISH SOMALILAND, SOMALIA, AND ETHIOPIA]

FEMALES*

[MAL€S HAVE BUSHyANTENNAE I \¡Í¡^M^^^\ ^^^ ^^ ^^^ 5/7£]

FIG. 24

LONG PALPS ON

ANOPHELES

ANTENNAE

-PROBOSCIS

PATTERN ON HIND TARSUS

Ï

PATTERN ON PALP

PATTERN ON VEIN 1

LIGHT ON BASAL FOURTH ANOPHELES

FUNES TUS

î-, I VEIN 6 DARK _ ^ND

ií^ + ON OUTER HALF - MACMAHONI

J /ONLY FUNESTOS \

^ I IMPORTANT} 1

ADULTS CANNOT I

\ BE SEPARATED /

_ ANOPHELES

— RHODES/ENS/S

_ ANOPHELES

~ ADENENS/S

( WING OF AOCfi£-/VSfS )

_ ANOPHELES

— A//L/

_ ANOPHELES

— D'THALI

PATTERN ON PALP

TWO LIGHT IJS'""""

r - " ^.î.^^N'^f^^jL:''iiiiiiiiiiii«iiiiiip

( WING OF ÇHHISrVi \

_ ANOPHELES

— CHßlSTYl

PATTERN ON PALP

_ ANOPHELES

— GAMBIAE

LAST SEGMENT ENTIRELY UGHT^

PATTERN ON HIND TARSUS TWO LIGHT SPOTS PRESENT ON VEIN 1

_ ANOPHELES

— PHAROENSIS

_ ANOPHELES

- PRETORIENSIS

r PATTERN ON THORAX

PALE BROWN

TO BLACK

LINES

YELLOWISH

AEDES

AEGYPT I

AEDES

SIMPSON!

— AEDES

"' AFRICANUS

ADULT MOSQUITO WITH PARTS LABELED

THORAX-..
MOSQUITOES OF MEDICAL IMPORTANCE 43

The rather steep slopes in the north above 6,000 feet do not

support the disease for the same reason. At altitudes over

é,000 feet the cycle of transmission is seasonal. Below this

level in the eastern area the disease is confined to the courses

of the larger rivers and is hyperendemic as well as epidemic.

Plasmodium falciparwm and P. vivax are the most common parasites. In western Ethiopia falcifarum predominates. Plasmodmnh malariae is scarce throughout the entire

region.

Anophdes gamhiae is responsible for most of the malaria

at high altitudes. This species tends to spread during the

breeding season and seeks higher and higher altitudes as

Eopulations increase. This expansion results in epidemics,

ecause people are exposed to fresh attacks of malaria in

areas where they have little immunity. This species, which

breeds in all kinds of manmade water containers, is a dangerous vector because of its proximity to man.

Anopheles funestus inhabits large rivers at low altitudes,

especially in very dry areas where lack of rain causes a

seasonal lowering of water levels and leaves residual pools

suitable for breeding. The malaria supported by this niosquito is less endemic than that transmitted by gamhiae.

Where funestus predominates as a vector ittransmits malaria

rather consistently.

Anopheles christyi is an eastern African highland species

that sometimes appears in large numbers. It is strongly

suspected as a vector in the Uganda highlands and is included in the keys because of its potential importance in

Ethiopia and Somalia.

Anopheles adenensis is thought to be widespread in Eritrea. It is a close relative of the very effective Indian vector

Anopheles culicifacies^ and for this reason it is implicated in

transmission wherever culicifacies is found. However, direct

evidence of its ability as a vector is difficult to find in the

literature. Its role in any local malaria transmission should

be carefully evaluated.

Anopheles pharoenèis is present but normally scarce in

Ethiopia. The rate of natural infection there and in other

areas is extremely variable. In some localities it has a high

infection rate, in others none at all. It has been included in

the keys because it has been implicated in transmission in

such widely separated localities as Egypt, southern Nigeria,

Kenya, Uganda, and Madagascar.

Anopheles pretoriensis is regarded by some workers as an

important vector in Ethiopia, although the evidence is not

certain. It is anthropophilic to some extent but is rarely

found inside houses in that country.

Anopheles rußpes is found in Ethiopia but is too scarce to

be important in malaria transmission, especially when gamhiae and funestus are present.

Anopheles rhodesiensis also is found in Ethiopia and probably in Somalia, but throughout most of its range it is generally not a domestic species. However, it has been found

from time to time with a low rate of natural infection and

is believed to be a secondary vector under favorable

conditions.

Anopheles nili is generally scarce in dry areas. It prefers

breeding in or near large rivers. The numbers of adults

collected in houses are correlated with the water level. This

species has not been found with natural infections in Ethiopia but is included in the keys because of its potential importance in that country.

The importance of Anopheles dHhali in Ethiopia is in

doubt. However, near Aiscia, Ethiopia, it was the only

Anopheles species found definitely associated with malaria

by the local people. It has been found in dwellings at Diredawa, Ethiopia, and in small numbers in tents in British

Somaliland, and it is suspected of being a vector in northern

Ethiopia.

References : 67, 75, 258.

Anopheles Species in Eritrea, the Somalilands, and

Ethiopia

leesoni

longipalpis

max)ulipalpis

marshallii marshallii

\nili

ohscurus ohscurus

\palvdis

\pharoensis

"^pretoriensis

"^rhodesiensis rhodesiensis

rhodesiensis rupicolus

\rufipes

sergenti macmahoni

squamosus cydippis

squamosus squamosv^s

turJchudi

wellcomei

\adenensis

amutis

ardensis

\christyi

cinereu^

cou^tani coustani

coustani ziemanni

dancalicus

demeilloni

"^dHhali

erythraeus

fu/nestus confusus

"^funestus funestus

^gamhiae

garnhami

gingeroi

harperi

kingi

See footnote 6, p. 10.

Yellow Fever.—Clinical yellow fever has seldom been reported from Eritrea, the Somalilands, and Ethiopia, but the

endemic zone, as proposed by the World Health Organization

Commission on Yellow Fever, includes almost all this region.

In Eritrea positive mouse-protection tests performed since

1940 indicate an immunity in children under 14 years of age.

The Somalilands are apparently free of yellow fever. In

Ethiopia along the Anglo-Egyptian Sudan frontier one case

of yellow fever has been reported ; however, others probably

occur there from time to time but not elsewhere in that

country.

Aedes aegypti transmits a small amount of yellow fever

in the port cities of this region. Aedes simpsoni and A,

africanus^ which are forest mosquitoes, are probably the

only other important vectors. Their relative importance has

not been evaluated.

References: 256, ^11.

Dengue.—Sporadic cases of this disease occur in Eritrea,

principally from April to June and from October to December. The incidence of dengue in Somalis is lower than

in Ethiopians and Europeans. It is endemic in the Somalilands. In French Somaliland it is known as obock and

massawa fever. In Ethiopia sporadic outbreaks of endemic

dengue have been reported from Diredawa, Harar, and the

western highlands.

References: ^56, Sll.

Encephalitis.—No information is available.

Filariasis.—Cases of this disease are sporadic in several

areas and are thought to be introduced from other parts of

Africa. Filariasis is probably not indigenous to the Somalilands and Ethiopia.

Reference: Ul.

West Africa

(Figs. 25 and 26)

Physical Features.—^West Africa includes Gambia, Portuguese Guinea, Sierra Leone, Liberia, Gold Coast, Nigeria,

and all French West Africa below the northern border of

Senegal and latitude 15° N. Four well-defined climatic zones

extend across this region from Gambia to French Equatorial

Africa. From south to north they include (1) a 10- to 60-

mile-wide coast with swamps and mangrove, (2) a 50- to

100-mile-wide belt of dense tropical forest with thick undergrowth and few clearings, (3) hilly country with deciduous

forests merging with park and high grass savannas, and (4)
44 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPCCIMCN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE — WEST AFRICA

[AREA INCLUDED: GAMBIA, PORTUGUESE GUINEA, SIERRA LEONE. LIBERIA^ GOLD COAST, NIGERIA,

AND SOUTHERN FRENCH WEST AFRlCAj

^i^^ FULL-GROWN LARVAE ^

FIG. 25

NO SIPHON ON ANOPHELES

LONG PLEURALS

OF 3RD GROUP

ONE HAIRED, ONE

BARE AND SPINELIKE

ONE HAIRED,

ONE BARE

AND HAIRLIKE

SIPHON ON ALL OTHERS

CLYPEAL HAIRS LONG PLEURALS

OF 2ND GROUP

PALMATE HAIRS

ON 5TH SEGMENT

_ ANOPHELES

'~ PRETORIENSIS

3 TO 7

BRANCHED

FROM MIDDLE

BRUSHLIKE

ANOPHELES

RUFIPES RUFIPES AND RUF IPES ¡NGRAMI

( BOTH IMPORTANT; LARVAE CANNOT Bf SEPARATED )

PECTÉN SPINES

^^iî^

_ ANOPHELES

-~ PHAROENSIS

INNER CLYPEALS

WITH LONG

• HAIRS ON

INNER SIDE

_ ANOPHELES

— BRUNNIPES

PROTHORACIC

HAIRS 1 AND 2

y

ANOPHELES

NILI

BASAL TUBERCLES

ABSENT

_ ANOPHELES

"~ GAMBÍAE

ALL SPINES A SIr\ DUCI CO ABOUT ANOPHELES

EQUAL — MELAS

IN LENGTH

SHAFTS

THICKENED

BASAL TUBERCLES

PRESENT AND CLOSE

TOGETHER

SPICULES

EVENLY

DISTRIBUTED/

PLATES ON 5THSEGMENT

ANOPHELES LARVA

ANOPHELES

FUNESTUS

OUTER CLYPEALS

COVERING

LESS THAN

1/2 OF

SEGMENT

DENSE PATCH

OF SPICULES y

AT BASE

OUTER CLYPEALS

LATERAL HAIRS

ON 5TH SEGMENT

_ ANOPHELES

— MOUCHETI NIGERIE NSI S

ANOPHELES

HARGREAVESI

ANOPHELES

HANCOCKI AND MARSHALLII

{ ONLY HANCOCK/ IMPORTANT; LARVAE \

\ SEPARATED WITH DIFFICULTY /

WITH FINE 1 SIDE HAIRS

BARE,

STOUT, AND ^

BLUNT à

_ ANOPHELES

~" RHODESIENSIS

_ ANOPHELES

— DOM ICOLUS

__ ANOPHELES

— MOUCHE Tl MOUCHE Tl

ONLY QNE

TUFT PER

SIDE, AND ON

OUTER HALF

OF SIPHON

COMB SCALES

COMB SCALES

IN ONE

REGULAR ROW

(NOT ON A PLATE;

PECTÉN SPINES

AEDES

A EGYPTI

AEDES

SIMPSONI
MOSQUITOES OF MEDICAL IMPORTANCE 45

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

LONG PALPS ON

ANOPHELES

ANTENNAE

-PROBOSCIS

MOSQUITOES OF MEDICAL IMPORTANCE—WEST AFRICA

JUINEAJ SIERRA

N FRENCH WES-

[FEMALES»

TAREA INCLUDED, CAMBIA, PORTUCUESE GUINEA, SIERRA LEONE; LIBERIA, COLD COAST, NIGERIA,

AND SOUTHERN FRENCH WEST AFRICA]

\^MALES HAVE BUSHY ANTENNAE ( M¿M^^^ ^^^ ^^ ^'^'^ B/TE]

FIG. 26

PATTERN ON HIND TARSUS

r PATTERN ON PALP

NO LIGHT SPOTS PRESENT ON VEIN 1

111

TWO LIGHT SPOTS PRESENT ON VEIN 1

''"lililí

AT LEAST FOUR LIGHT SPOTS PRESENT ON

FRONT MARGIN; REMAINDER OF WING DARK

VEIN 3 DARK

ON BASAL HALF

ANOPHELES

BRUÑNIPES

ANOPHELES

RHODES/ENS/S

ANOPHELES

FUNESTUS

PATTERN ON PALP

«lilli Í^H||||iAíl"iiiíi;,- -(-

ONE LIGHT SPOT PRESENT ON VEIN 1

Biiii Bi'^liiPiiiiiiiii'ifcm^^ +

TWO LIGHT SPOTS PRESENT ON VEIN i

ANOPHELES

DOM ICOLUS

ANOPHELES

GAMBIAE AND MELAS

{ BOTH IMPORTANT; FEMALES CANNOT BE \

\ SEPARATED WITH CERTAINTY )

ANOPHELES

MOUCHETI MOUCHETI

AND MOUCHETI NIGERIENSIS

( BOTH IMPORTANT; FEMALES CANNOT)

^ BE SEPARATED WITH CERTAINTY '

ANOPHELES

HARGREAVESI

TOP VIEW OF THORAX

LAST SEGMENT ENTIRELY WHITE^

2^

PATTERN ON HIND TARSUS

_ ANOPHELES

— PHAROENSIS

— ANOPHELES

— PRETORIENSI S

liny.

iiiiitt«lilllK^~ ANOPHELES mmwm^--- HANCOCKI

ONE LIGHT SPOT PRESENT ON VEIN 1

1^^ + lllPfi! (lilt-- z:

TWO LIGHT SPOTS PRESENT ON VEIN 1

MORE THAN 1/3 AS LONG AS

SEGMENT 3 —~ ^ ^^ — ANOPHELES

'K*^ " Run RES RUFIPES

ANOPHELES

RUFIPES INGRAMI

SHORT PALPS ON

ALL OTHERS r PATTERN ON THORAX

1 LINES

YELLOWISH

AEDES

SIMPSON!

ADULT MOSQUITO WITH PARTS LABELED
46 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

the southernmost part of the Sahara Desert. All the countries in West Africa, except Senegal and Gambia, contain

at least the first two zones, Nigeria is the only one that has

all four, and French West Africa is largely desert, except

for French Guinea, Ivory Coast, and Dahomey, all of which

extend to the coast. Senegal and Gambia in the westernmost

part of this region are ñat and monotonous. Vegetation is

concentrated in the river basins and in the Casamance.

The dry season in West Africa is caused by a dry, hot

harmattan blowing south from the Sahara from November

to April. The wet season is brought by a monsoon blowing

north from the Gulf of Guinea from May to October. The

inches of rain each zone receives annually are as follows:

Zone 1 about 170, zone 2 from 55 to 170, zone 3 from 40 to 55,

and zone 4 less than 25 in some areas.

Reference: SOI.

Malaria.—Highly intense malaria with spleen rates of

70-90 percent occurs in the southern parts of most of these

countries in West Africa. This disease is particularly

dangerous in the coastal strip (zone 1), which contains vast

mangrove swamps, and along the courses of large rivers.

In zone 2, which extends inland for 100 to 150 miles, a high,

dense, tropical forest tends to inhibit the breeding of malaria

vectors. Spleen rates are slightly lower than on the coast.

The distribution of malaria in zone 3 is seasonal, but transmission continues through most of the year. In zone 4 the

northern parts of the countries, which contain dry savannas

merging into semideserts, support a distinctly seasonal malaria that closely corresponds to that in the summer rainy

season.

In most of the tropical rain forest malaria caused by

Plasmodium falcipanr-um predominates. Pldsmodium molariae cases are less numerous over most of the region. Plasmodium ovale and P. vivax have been reported as uncommon,

although widely distributed. However, recent work in West

Africa suggests that vivax is extremely rare or even absent

in this region and that most previous reports of vivax

should be assigned to ovale.

Anopheles gambiae breeds in all kinds of water-holding

dej)ressions in the ground and is closely associated with man.

It is one of the most efficient vectors of all the Anopheles

species and is almost always associated with high parasite

rates wherever it is abundant. It is widely distributed over

entire West Africa and is responsible for most of the malaria

there, both in rural and urban locations. It is the mosquito

of the rainy season, reaching its population peak in June,

July, and August, and almost completely disappearing in

February and March.

Anopheles fwnestus is characteristically associated with

large bodies of water, especially along river courses, and

transmits malaria most efficiently during the first half of the

dry season. Together with gambiae,^ which is common early

in the rainy season, funestus helps to carry the year-round

malaria found near the coast and along the large river

courses.

Anopheles melas is the third most important transmitter in

the region. It is a brackish-water breeder, with a particular

preference for Avicennia mangrove swamps. These swamps

are flooded by high spring tides and are important breeding

places because of the flat, peaty soil. Because melas larvae

can withstand sudden changes in salinity and because the

species is almost as effective a vector as gambiae itself, it easily

maintains most of the highly endemic, strictly coastal malaria. Several other species of Anopheles may have some

importance in West Africa but are not nearly so effective in

maintaining malaria as gaTubiae, funestus^ and melas.

Anopheles hancocki is a subsidiary vector in parts of Ivory

Coast and Haute-Volta. In Liberia this anthropophilic

species is generally endophilic, but in Sierra Leone it bites

man out of doors only.

Anopheles donúcolus represented 25 percent of all anophelines caught indoors in northern Nigeria, and in parts of

French West Africa and Haute-Volta it was found to be

highly infected and a subsidiary vector.

Anopheles hargreavesi is of considerable importance in

Nigeria. It is common in native huts and bites man freely

out of doors. Because of its more uniform distribution and

greater abundance, it may be a more important vector under

certain conditions than gamhiae,^ especially at Itowolo,

Nigeria.

Anopheles rufipes ruppes and A, r, ingrami have represented up to 70 percent of all mosquitoes found in houses, as

determined by studies in certain parts of northern HauteVolta. They are believed to be the principal vectors there.

Anopheles moucheti nigeriensis is of some importance near

Lagos, Nigeria. The typical form. Anopheles moucheti

moucheti^ is included in the keys because of its importance

elsewhere.

Anopheles hrunnipes is included in the keys because it has

been found with high salivary infection rates during the

winter in nearby Belgian Congo. It is not regarded as an

important vector in French West Africa.

Another secondary vector. Anopheles nili^ is very common

in Liberia, Sierra Leone, and French West Africa. Its

breeding is associated with large streams, and the adults are

often found indoors.

Anopheles pharoensis is largely a swamp breeder. Dissections indicate that this species varies considerably in its

natural infectivity. It is found infected in some localities

in French West Africa and southern Nigeria, whereas in

other nearby localities the tests have been entirely negative

for malaria. This species is a vector of secondary importance.

Anopheles pretoriensis does not frequent houses. It is included in the keys because it is infected in other parts of its

range and because it is often associated with gambiae.

Anopheles rhodesiensis is included in the keys because of

its importance outside West Africa. This species does not

frequent houses in this region, but it has been found with high

natural infection rates.

References: 61,161,259,260.

Anopheles Species in West Africa

africanus

bmrberellus

hrohieri

brumpti

\hrunnipes

cavernicolus

cinctus

coustani coustani

Goustani tenebrosas

coustani ziemanni

"^domicolus

dwreni

flavicosta

freetownensis

*funestus

'^gambiae

'^hancocki hancocki

hancocki masseguini

* hargreavesi

implexus

jebudensis

See footnote 6, p. 10.

Yellow Fever.—Small outbreaks of classical urban yellow

fever have been reported at irregular intervals from the cities

and towns of West Africa. Aedes aegypti has been responsible for these epidemics, one of which occurred in 1946 in

leesoni

macuZipalpis

marshallii

^melas

minutus

\moucheti moucheti

'^moucheti nigeriensis

\nili

ohscurus nowlini

obscuncs obsGwrus

\paiudis

"^pharoensis

\pretoriensis

\rhodesiensis

rivuloru/m rivulorum

"^rufipes ingrami

"^rufipes rufipes

smithii

squ/imosu^ squamosus

wellcomei
MOSQUITOES OF MEDICAL IMPOETANCE 47

southwestern Nigeria. Jungle yellow fever is endemic and

has been a constant problem, accounting for all yellow fever

cases that are reported. Itis restricted to the belt of tropical

forest crossing the region from west to east^ and numerous

positive mouse-protection tests have been performed throughout this area. The monkey-to-monkey vector has not yet

been determined. The only Aedes species so far implicated

with any certainty is simpsoni. For further information,

see pages 2, 3, 40, 43, and 53.

References : 92, 232, 234, 256.

Dengue.—There is little active dengue in West Africa.

Most of the cases have been reported from the coasts of Nigeria and coastal French West Africa. The last outbreak

occurred in Dahomey in 1947.

References : 256, 311.

Encephalitis.—Very little information is available about

encephalitis in West Africa. The Zika virus has been isolated from humans at Afikpo, Nigeria, but nothing is known

about its etiology.

Reference: 257.

Filariasis.—Definite records are available of transmission

of Wuchereria bañerofti by Anopheles gambiae and A,

funestus in the plantation districts of Sierra Leone. These

two species are the prime vectors of filariasis in that country.

A, gamèiae has been found with natural infections at Eoberts

Field, Liberia, where up to 8.4 percent of the natives have

been found with filariasis and several cases of elephantiasis

have been observed. In the coastal Marshall Territory of

Liberia, Anopheles melas and gambiae appear to be the most

important vectors. Anopheles haneocM females were also

found with natural infections. A. gambiae is believed to be

the principal vector during the rainy season and mela^ during the dry season and the beginning of the rainy season.

Farther north at Dakar, French West Africa, filariasis is

rare, and although it is present in the Northern Territories of

the Gold Coast, the vector there is not known.

References : lOJt, I43,164,190, 250, 345.

West-Central Africa

(Figs. 27 and 28)

Physical Features.—West-Central Africa includes the

Cameroons, French Equatorial Africa, and Belgian Congo.

The Cameroons and French Equatorial Africa have an alluvial coast on the west and south. A belt of dense tropical

forest, contiguous with that in Nigeria and eastern Africa

and extending north to latitude 5° N., covers the littoral

plain, the subcoastal highland slopes, and the low swampy

areas of the southeast Congo basin. In this forest abundant

rainfall and high temperatures vary little from day to day

or from year to year. The center of French Equatorial

Africa north of the forest belt contains open savannas, with

isolated clumps of forest along the rivers. This area is dry

between December and March and wet from April to September. A semiarid sandy desert, which has hot days and

cold nights, lies between the savanna area and the northeastern Tibesti Massif, a range of mountains almost 11,000 feet

high.

The outstanding physical feature of Belgian Congo is the

valley of the Congo Eiver, a flat depression in the central

African plateau, which is covered by the tropical-forest belt

spreading south and east from French Equatorial Africa.

Extensive swamplands lie at the junction of the Congo and

Ubangi Eivers along the western border. Highlands of

6,000 to 7,000 feet form the Uganda frontier. Mt. Kuwenzori, which is about 16,000 feet high, overlooks the Katanga

plateau in the southeast corner. Except in the highest parts

471779 0—59 4

of Belgian Congo, the temperature is always close to 80° F.,

with very small daily and annual changes. Annual rainfall

is about 80 inches, most of which occurs in a 7- to 9-month

period. The invigorating eastern and southeastern highlands are savannas, with forests along the rivers.

Reference: 301.

Malaria.—The belt of tropical rain forest, covering the

southern third of French Equatorial Africa and almost the

entire Congo basin, has highly endemic malaria, which is

transmitted throughout most of the year. Spleen rates in

this area range from 70 to 90 percent. In the east-central

Ubangi-Shari highlands of French Equatorial Africa malaria has a highly marked seasonal distribution, clearly corresponding to the rainy season. In the northern parts of the

Cameroons and French Equatorial Africa malaria incidence

becomes even more seasonal. Southeast of the Congo basin

in the Katanga highlands the disease is seasonal, with spleen

rates of 50 to 70 percent. Similar conditions prevail in extreme southern French Equatorial Africa.

In western Belgian Congo Plasmodiûm vivace infections

are about as numerous as those of P. malariae, Plasmodium

falciparum is the predominant parasite in the east, malariae

is next in abundance, and vivaœ is encountered least. Plasmodium ovale is scarce in this area.

Throughout the rain-forest belt and in many other localities Anopheles gamhiae is the predominant vector. Itis the

most efficient malaria transmitter in Africa. This factor,

together with its habit of breeding in all types of small

water-containing depressions close to man's habitations,

makes it the most dangerous vector in this region. It breeds

in immense numbers throughout the rainy season.

Anopheles melas^ originally described as a variety of gambiae^ is a brackish-water breeder that prefers Avicennia mangrove swamps, although it breeds prolifically outside such

areas along the coast. The larvae are able to withstand

rather large changes in salinity, so that high tide floods in

the spring do not disturb their breeding places. This species

is nearly as efficient a vector as gamhiae, and in areas where

it is abundant it may transmit a malaria of extremely high

endemicity.

Anopheles funestus, the second most abundant vector in

this region, inhabits large bodies of water, and as summer

rains decrease this species appears in large numbers, especially along the main water courses. Because gawhiae is

prevalent during the rainy season and funestu^ is abundant

afterward, malaria is transmitted during almost the entire

year. In higher areas gambiae is the most prevalent species.

Even though these three Anopheles species transmit most of

the malaria in West-Central Africa, several other vectors of

varying importance play minor roles in its transmission and

are discussed here.

Anopheles moucheti moucheti has often been reported as

the most prevalent mosquito found in houses in Belgian

Congo, at times far outnumbering gamhiae and other mosquitoes. Parasite rates in this species have almost always been

low and account for the relatively low endemic malaria in

many parts of Belgian Congo, where the principal vectors

are scarce or absent.

The adults of Anopheles brunnipes have been observed

in large numbers inside houses in Leopoldville. During the

winter in this locality salivary infection rates of up to 4 percent have been found. Little is known of the biology of this

species, but it is highly possible that brunnipes plays a part

in winter malaria.

Anopheles nili has variable habits and breeds at the edges

of large rivers. The predominance of adults caught in human habitations is directly correlated with the rise and fall of

the rivers in which it breeds. Because of its house-haunting

habits it is regarded as a secondary vector of some importance.
48

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE — WEST-CENTRAL AFRICA

TAREA INCLUDED- CAMEROONSJ FRENCH EauATORiAL áFRICA, AND BELGIAN CONGOJ

-^^« FULL-GROWN LARVAE 1^"^-

C LONG PLEURALS

OF 3RD GROUP

NO SIPHON ON ANOPHELES

CLYPEAL HAIRS LONG PLEURALS

OF 2ND GROUP

PALMATE HAIRS

ON 5TH SEGMENT

FIG. 27

3 TO 7 1

BRANCHED

PROM MIDDLE

ANOPHELES

RUFIPES

RUFIPES

SHOULDERS

SLOPING

SHOULDERS

NEARLY

SQUARE

_ ANOPHELES

— CHRISTY I

_ ANOPHELES

"~ PRETORIENSIS

PECTÉN SPINES

WITH

LARGE _ ANOPHELES

— PHAROENSIS

- -<

ONE HAIRED, ONE

BAR^E AND SPINE-^ __ ANOPHELES

— BRUN NIPES

CLYPEAL HAIRS

RE. CLOSE^,,..—H

WITH LONG

HAIRS ON

INNER SIDE

ANOPHELES

PALUDIS, SYMESI,

ZL AND COUSTANI

/ONLY PALUDIS IMPORTANT;

LARVAE CANNOT BE

\ SEPARATED

_ ANOPHELES

— NIL I

ONE HAIRED.ONE

BARE AND

HAIR-LIKE

LONG AND AK,r\DUCI CC SHORT ANOPHELES

SPINES — GAMBIAE

MIXED

ALL SPINES

ABOUT _ ANOPHELES

— MELAS

ANOPHELES LARVA

INNER CLYPEALS

ANTENNA j OUTER CLYPEALS

"n^Ä^^s^rir ^ :t'z^.^LT.'7ti^^

'^Ç^':-^^ GROUP

^-^ -3~ GROUP

5^« SEGMENT

BASAL TUBERCLES

PRESENT AND CLOSE

TOGETHER

PLATES ON 5TH SEGMENT

COVERING

ABOUT 1/2

OF SEGMENT HIND EDGE OF ANOPHELES

+ PLATE NOT DARKENED = FUNESTUS

DENSE PATCH OF

SPICULES AT

BASE.

SIPHON ON ALL OTHERS

LATERAL HAIRS

ON 5TH SEGMENT

2 TO 4 BRANCHED

f OUTER CLYPEALS

3ARE AND

HAIRLIKE

WITH FINE

SIDE HAIRS

_ ANOPHELES

— HARGREAVESI

_ ANOPHELES

— RHODESIENSIS

_ ANOPHELES

— MOUCHETI MOUCHE Tl

PECTÉN SPINES

6 TO 8 BRANCHED

ONLY ONE

TUFT PER

SIDE.AND ON

OUTER HALF

OF SIPHON

AT LEAST

12 SPINES

COMB SCALES

IN ONE J.

REGULAR ROW "^

(NOT ON A PLATE)

COMB SCALES PECTÉN SPINES

ANOPHELES

HANCOCKI

AEDES

A EGYPTI

AEDES

SIMPSON/

AEDES

AFRICANUS
MOSQUITOES OF MEDICAL IMPORTANCE 49

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - WEST-CENTRAL AFRICA

'^'^^^^ |\REA INCLUDED CAMEROONS, FRENCH EQUATORlAL AFRICA, ANO BELGIAN CONGOJ

^ '^m FEMALES

[MALES HAVE BUSHY ANTENNAE I ,,,^f/.^%ÈÉÊ \ ^^^ ^^ ^<^^ BITE] FIG. 28

LONG PALPS ON

ANOPHEL ES

ANTENNAE

PROBOSCIS

, DARK ON OUTER _ ANOPHELES

•" HALF OF VEIN 6 — FUNESTUS

_ ANOPHELES

— BRUNNIPES

_ ANOPHELES

"" HARGREAVESI

,PRESENT OR ABSENT

SHORT PALPS ON

ALL OTHERS

r PATTERN ON THORAX

• PATTERN ON FRONT MARGIN OF WING

AT LEAST FOUR LIGHT SPOTS PRESENT

ONLY TWO LIGHT SPOTS PRESENT -

ANOPHELES

HANCOCK!

ANOPHELES

PALUDIS

__ AEDES

AEGYPTI

SEGMENTS 3, 4. AND 5

OF HIND TARSUS

^ ^^ _ AEDES

T -^ —-^^^ — AFRICANUS

_ AEDES

"~ SIMPSON I

ADULT MOSQUITO WITH PARTS LABELED

THORAX-^
50 AGRICULTURE HANDBOOK 152, U. S, DEPT. OF AGRICULTURE

Anopheles pcdvdis readily enters houses in Belgian Congo,

where resting females have been found with gut and salivary

gland infections. Itis not common, but itis probably important as a secondary vector.

The remaining Anopheles species presented in the keys all

are found in West-Central Africa, sometimes in large nunibers. According to available records, none of these mosquitoes are normally infected or associated with man in this

region, but elsewhere they have been implicated in malaria

transmission at one time or another and are potentially important. These Anopheles species and the areas of their

potential or actual medical importance include rufipes^ rwfipes^ Anglo-Egyptian Sudan, Haute-Volta; rhodesiensis^

Sierra Leone; pretonensis^ Transvaal, Southern Ehodesia;

pharoensis^ French West Africa, Egypt, southern Nigeria,

Uganda; christyi^ Uganda; hancocM^ Ivory Coast, HauteVolta, Uganda ; and hargreavesi^ Nigeria.

References : 67,260,

Anopheles Species in West-Central Africa

marshallii marshallii

marshallii mousinhoi

marshallii pitchfordi

"^nielas

rmchaeli

mortiauwi

"^moucheti mowcheti

natalensis multicinctus

natalensis natalensis

"^nili

obscurus nowlini

oiscurus ohsGurus

"^paludis

\phaToensis

\preuoriensis

\rhodesiensis

rivuloruTTb rivulorum

rodhaini

\rufipes rufipes

seydeli

smithii rageaui

squamosus cydippis

squamosus squamosus

symesi

theileri

vanhoofi

vinckei

walravensi schwetzi

vyellcomei

ardensis

argenteolohatus

herghei

*hrunnipes

fchristyi

cinctiùs

cinereios

GoncoloT

coustani caliginosus

Goitstani coiostani

comtami teneirosus

Goustani zienbo/nni

demeilloni

distinetus distinotus

distinetics ugandae

dureni

faini

"^ fîmestus

'^gamhiae

garnhami garnhand

\hanG0Gki

\hargreavesi

inoplexus

keniensis

kingi

leesond

lloreti

longipalpis

maculipalpis

marshallii gihhinsi

See footnote 6, p. 10.

Yellow Fever.—Numerous cases of jungle yellow fever

have been reported from West-Central Africa, and immunity

to this disease is apparently rather widespread. Although

Aedes aegypti^ A, simpsoni, and A, afrieanits are considered

to be the important vectors in this region, no studies have

been completed on their respective roles in transmission.

For additional information, see pages 2 and 3.

References : 25ß, Sil.

Dengue.—This disease is endemic throughout most of this

region. Although active cases appear from time to time,

they are not numerous.

References : 2Sß, 311.

Encephalitis.—In the extreme northern part of Belgian

Congo, West Nile antibodies have been found in humans, and

in an area of the Congo River to the south tests have been

positive for neutralizing antibodies to West Nile virus. It

is not known how this virus is spread in Belgian Congo.

Reference: 257.

Filariasis.—The distribution of mosquito-borne filariasis

is not well known in West-Central Africa. Oulex pipiens

quinquéfaseiatus is widely distributed in the lowe^ Congo

Valley at Matadi, where it transmits Wuchereria hancrofti,

Filariasis and elephantiasis are widespread in the native

population at Douala, Cameroons, where the vector is not

known.

References: 1SJ^, 295, S38.

Elast-Ceiitral Africa

(Figs. 29 and 30)

Physical Features.—East-Central Africa includes Kenya,

Uganda, Tanganyika, and Zanzibar and associated islands in

the Indian Ocean. The northern three-fifths of Kenya is

arid, receiving from 5 to 20 inches of rain per year. This

semidesert country rises from the low, marshy coast toward

the Ethiopian highlands. Southern Kenya contains most

of the population and agriculture, because it has a moderate

altitude and a subtropical climate, with 40 to 50 inches of

rain annually. The western half of Kenya is traversed from

north to south by the Eift Valley, lined on either side by

towering peaks. The extreme western slopes of the interior

highland descend gradually to the shores of Lake Victoria.

The Eift Valley forms the western frontier of Uganda.

Savanna tablelands to the east and north are topped by a

high mountain range along the Kenya frontier. In the

savanna area the rainfall averages from 40 to 55 inches annually in two rainy seasons, one from March to May and the

other from September to November.

The 10- to 40-mile-wide eastern coastal strip of Tanganyika rises to a central plateau of about 4,000 feet and descends

to the shores of Lake Tanganyika in the west. Mountains

over 15,000 feet high border Kenya in the north, and mountains of about 9,000 feet are a part of the southern highlands.

The Eift Valley extends southward to separate Tanganyika

from Belgian Ôongo. Two-thirds of Tanganyika, including

most of the plateau, has less than 40 inches of rain and no

ground water. The most densely settled areas are the northern half of the coast above 4,000 feet, the shores of Lakes

Tanganyika and Victoria, and some of the major river valleys, all of which have between 40 and 60 inches of rain annually.

Zanzibar is a low limestone island with a tropical monsoon

climate. From April to June it receives up to 60 inches of

rain ; from December to March it is hot and dry. The island

is very fertile, and almost all the land is under cultivation.

Reference: 501.

Malaria.—Zanzibar, Pemba, nearly all Uganda, the extreme southern part of Kenya, and that part of Tanganyika

in the north under 5,000 feet and its east coast proper are mtensely malarious, partly because of the high rainfall brought

by monsoons from the southeast. Spleen rates in these areas

range from 70 to 100 percent. In the northeast quarter of

Kenya malaria is rather confined to watercourses and ^ trulj^

hyperendemic only along the large rivers. All Kenya s

central area, most of which is over 6,000 feet, and the southern

highlands of Tanganyika over 5,000 feet are alniost devoid

of malaria. Hi these mountainous areas at altitudes over

4,000 feet, endemic malaria is transmitted for 4 to 6 months

depending on the altitude, with occasional epidemic outbursts. The intervening periods are almost completely tree

of this disease. A small sharp outbreak of malaria has been

recorded at 8,600 feet in central Kenya. At this altitude

malaria is rare, and when present it can be transmitted for

only a very shorttime.
MOSQUITOES OF MEDICAL IMPORTANCE 51

IMPORTANT

A SPCCIMCN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - EAST-CENTRAL AFRICA

[AREA INCLUDED: KENYA. UGANDA, TANGANYIKA, AND ZANZIBAR ]

: FULL-GROWN LARVAE^

NO SIPHON ON ANOPHELES

PECTÉN

LONG PLEURALS

OF 3RD GROUP

SIPHON ON ALL OTHERS

r PALMATE HAIRS

ON 5TH SEGMENT

FIG. 29

_ ANOPHELES

"" CHRISTY I

- ANOPHELES

" PRETORIENSIS

- ANOPHELES

" RUFIPES RUPIPES

WITH LONG

-|. HAIRS ON

PROTHORACIC

HAIRS 1 AND Z

r SHAFTS

SLENDERAV / / . \ .

BASAL TUBERCLES

ABSENT

_ ANOPHELES

NILI

_ ANOPHELES

~" GAMBIAE

SHAFTS ^^//\\l

THICKENE^^'^-''

ANTENNA

SPICULES

EVENLY

DISTRIBUTED .

PLATES ON 5TH SEGMENT

COVERING

ABOUT 1/2

r OF SEGMENT

ANOPHELES LARVA

INNER

CLYPEALS HIND

OUTER CLYPEALS

CLYPEALS -< "

BASAL TUBERCLES

PRESENT AND

CLOSE TOGETHER

DENSE PATCH

OF SPICULES

AT BASE

LATERAL HAIRS ON

5TH SEGMENT

Ï

__ ANOPHELES

~ FUNESTUS

INNER CLYPEALS

5^" SEGMENT

ABDOMINAL

T^^ PLATES

2 TO 4 BRANCHED

ANOPHELES

HARGREAVESI AND HARPERI

/ONLY HARGÑEAVESI IMPORTANT^ >TAN;\

{ LARVAE SEPARATED WITH

\ DIFFICULTY

_. ANOPHELES

"" RHODESIENSIS

_ ANOPHELES

MOUCHETI MOUCHET I

6 TO a BRANCHED

INNER CLYPEALS

BARE AND

HAIRLIKE

STOUT,

BLUNT

AT TIP

_ ANOPHELES

~~ HANCOCKI AND MARSHALLI/

/ONLY HANCOCKI IMPORTANT} \

{ LARVAE SEPARATED WITH )

\ DIFFICULTY /

_ ANOPHELES

~ DOMICOLUS

COMB SCALES

IN ONE

REGULAR ROW

(NOT ON A PLATE)

COMB SCALES PECTÉN SPINES

_ AEDES

~ AEGVPTI

— AEDES

~ SIMPSON I

— AEDES

"~ AFRICANUS
52 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - EAST-CENTRAL AFRICA

TAREA INCLUDED: KENYA. UGANDA. TANGANYIKA, AND ZANZIBAR)

m FEMALES IWI

[^f^L£S HAVE BUSHY ANTENNAE

LONG PALPS ON

ANOPHELES

ANTENNAE

-PROBOSCIS

PATTERN ON HIND TARSUS

PATTERN ON PALP

] AND DO NOT BITE\

NO LIGHT SPOTS PRESENT ON VEIN 1

5

FIG. 30

ANOPHELES

x/FiN fi nARK rUNESTUS KHO

ONOUTES = MACMAHONf

HALF iouv< ruN£STus \yA-\

PORTANT; LARVAE \

CANNOT BE j

SEPARATED /

AT LEAST FOUR LIGHT SPOTS ON FRONT MARGIN;

REMAINDER OF WING DARK v

_ ANOPHELES

"~ RHODESIENSIS

ANOPHELES

DOMICOLUS

PATTERN ON VEIN 1

PATTERN ON PALP

_ ANOPHELES

— CHRISTYI

SCALES ON THORAX

iSi

ONLY ONE LIGHT SPOT PRESENT

•filluè. +

_ ANOPHELES

— HARGREAVESI

_ ANOPHELES

— MOUCHETI MOUCHETI

FRINGE SPOT

"•a i^ifi'iiisilEpii' illl'll ï,SlKlll'''ï — ANOPHELES

PRESENT AT END OF VEIN 6

r PATTERN ON HIND TARSUS

GAMBIAE

LAST SEGMENT ENTIRELY LIGHT

^^^^$$$$^i?^

íí^^"^>

^ "iilll ll|l|lliillilll!iiiiii'''''l]TnTltwiiiJiimi^

TWO LIGHT SPOTS PRESENT ON VEIN 1

ANOPHELES

PHAROENSIS

ANOPHELES

PRETOR/ENS/S

_ ANOPHELES

"" RUF/PES RUFIPES

— ANOPHELES

"~ HANCOCKl

TOP VIEW OF THORAX

1 SfflTx

s.

SCALES ON

THORAX

y

SHORT PALPS ON

ALL OTHERS

•PATTERN ON THORAX

— AEDES

~ AEGVPTI

SEGMENTS 3, 4. AND 5

OF HIND TARSUS

« AEDES

^^Tô - AFRÍCANUS

^__ __ AEDES

^T^^ ~" SIMPSONI

ADULT MOSQUITO WITH PARTS LABELED

--WING
MOSQUITOES OF MEDICAL IMPORTANCE 53

Plasmodium falciparum predominates over East-Central

Africa and is almost always found in epidemics. Plasmodium malariae is not nearly so abundant. Plasmodium

ovale is believed to be indigenous to this region, but it is not

common.

Anopheles gamhiae^ here as elsewhere in tropical Africa,

is widespread and the principal vector. Because it breeds

in manmade depressions in the ground, it is a mosquito of

the rainy season. It is responsible for most of the malaria

transmitted at high altitudes and was the vector in the epidemic recorded at 8,600 feet in Kenya (see above).

Anopheles funestus^ the second most important vector in

Africa, is as important in the highlands of northern Kenya

as anywhere on the continent. This mosquito supports

hyperendemic malaria along large rivers, where gamhiae is

scarce or entirely absent because of lack of rain. The natural

infection rates of funestus are normally lower than those of

gamhiae ; therefore, the malaria transmitted by it is slightly

less intense. Although these two species together are by far

the most important, other species also serve as vectors at times

and in localities where populations of gamhiae and funestus

are low.

One of these. Anopheles christyi^ has been implicated as a

vector in the Kigezi highlands of Uganda. In this particular locality cattle are scarce, so that the normally zoophilic christyi is forced to seek human blood. Over most

of its range it is not regarded as a vector at all.

Anopheles hancocki^ a secondary vector of some importance

in Uganda, commonly bites man and rests indoors. It has

sometimes been found with a high infection rate.

Anopheles moucheti moucheti^ a vector whose role in transmission is not exactly known, breeds in Uganda in quiet or

flowing, shaded, clean Avater. In Belgian Congo it has been

observed indoors feeding on man and often has been found

with a low natural infection rate.

Anopheles pharoensis is found naturally infected in certain

parts of Uganda, but in closely adjoining areas the tests for

malaria are alw^ays negative. Since it is a secondary vector

in French West Africa, it may be important on account of

its large numbers in the ricegrowing area of Kenya.

Six additional Anopheles species have been implicated as

vectors in various parts of their range outside East-Central

Africa. Their ranges overlap this region or extend through

it, and because these species appear in large numbers at

various times and places, they have been included as potentially important vectors in the keys. The Anopheles species

and the areas in w^hich each has been implicated in malaria

transmission include domicolm^ French West Africa, HauteVolta; hargreafvesi^ Nigeria; nili^ Belgian Congo, Liberia,

Sierra Leone, French West Africa ; ruflpes rufipes, AngloEgyptian Sudan, Haute-Volta; rhodesiensis^ Sierra Leone;

and pretoriensis^ Ethiopia, Somaliland, Southern Rhodesia,

Transvaal.

References: 67,160,260.

Anopheles Species in East-Central Africa

ardensis

brohieri

'^christyi

cinereus

Goustani coustani

coustani tenehrosus

Goustani ziemanni

demeilloni

distinetus ugandae

\domicolus

funestus confusus

"^funestus funestus

"^gamibiae

garnhami hasilewskyi

gamhami garnhami

"^hancocki

\hargreavesi

im^plexus

keniensis

hihena

hingi

leesoni

longipalpis

lovettae

machardyi

maGulipalpis

marshallii gihhinsi

marshallii marshallii

rivulorumh,

\rufipes rufipes

serge7iti ^nacmahoni

squam^osus cydippis

squamosus entehiiensis

squanwsus squamosus

symesi

loellcomei erepens

wellcomei wellcomei

wilsoni

^moucheti moucheti

natalensis multicinctus

natalensis natalensis

\nili

njombiensis

obscurus obsGurus

]paludis

^pharoensis

\pretoriensis

\rhodesiensis

See footnote 6, p. 10.

Yellow Fever.—The Virus Research Institute at Entebbe,

Uganda, has been instrumental in elucidating the epidemiology of jungle yellow fever in East-Central Africa. Aedes

aegypti^ A, simpsoni^ and A. africanus transmit yellow fever

in a manner similar to that described for Anglo-Egyptian

Sudan (p. 40). Most of the mouse-protection tests performed in Uganda, western Kenya, and northern Tanganyika

have been negative. For further information, see page 2.

References : 150, 238, 256, 311.

Dengue.—This disease appears infrequently in East-Central Africa. It is endemic throughout most of the region.

Occasionally it appears in epidemic form, especially along

the coast.

References : ,311.

Encephalitis.—Rift Valley fever has been reported from

Kenya and Uganda. West Nile encephalitis virus is occasionally encountered in both of these countries. Because

Japanese and St. Louis encephalitis viruses are closely related antigenicly to the West Nile virus and because the first

two have a low incidence of positive sérologies, it is believed

that they actually do not occur in Africa at all. In addition

to West Nile encephalitis virus, antibodies to the following

viruses have been found in animals in Uganda : Ntaya, Zika,

Bunyamwera, Uganda S, Semliki Forest, and Bwamba. The

same viruses are also present in northern Tanganyika, but

their exact distribution has not yet been studied thoroughly.

References : 246,257.

Filariasis.—Filariasis is endemic along much of the coast

of East-Central Africa. A large endemic center in Tanganyika, south of Lake Victoria, includes almost 30 percent of

the native population, and a smaller center at the head of

Lake Nyasa presents another acute problem. Wuchereria

bañerofti is found in all cases of mosquito-borne filariasis in

Tanganyika, but the vectors have not been determined with

certainty. Both Culex pipiens quinquéfasciatus and Anopheles gamhiae may be involved. Elsewhere in East-Central Africa filariasis is not known to occur.

References : 99,112,158, 201.

Madagascar and Adjacent Islands

(Figs. 31 and 32)

Physical Features.—Madagascar, about 1,000 miles long,

is the fourth largest island in the world. It has a deeply

eroded grassland plateau, with ranges as high as 9,500 feet.

The weather is warm and rainy there from October to April

and dry and cool from May to September. The low western

shore on the Indian Ocean is bordered by numerous islands

and coral reefs ; on the east the steep, wet, densely forested

slopes are bordered by a narrow strip of swamps and lagoons.

The southern and southwestern parts of the country are semiarid, with less than 15 inches of rain annually.

The Comoro Islands, a volcanic archipelago between

northern Madagascar and the African mainland, are mountainous, densely forested, and very fertile. Their climate is

much like that of Madagascar.
54 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - MADAGASCAR AND ADJACENT ISLANDS

[AREA INCLUDED; MADAGASCAR^ COMORO ISLANDS, SEYCHELLES, MAURITIUS, AND REUNION]

-. ANOPHELES

~- CHRISTY!

_ ANOPHELES

— PRETOP/ENS/S

SIPHON ON ALL OTHERS

ONLY ONE

TUFT PER

SIDE, AND

ON OUTER

HALF OF

SIPHON

AT LEAST

12 SPINES

COMB SCALES

IN ONE

REGULAR ROW

(NOT ON A PLATE)

COMB SCALES

X

AEDES

AEGYPTI

_ AEDES

— ALBORICTUS

DN~1288
MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE- MADAGASCAR AND ADJACENT ISLANDS

[AREA INCLUDED: MADAGASCAR, COMORO ISLANDS, SEYCHELLES, MAURITIUS, AND REUNION]

55

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

FEMALES

\^M/iL£S HA\/E BUSHY ANTENNAE i jM^^ ) ^^^ ^^ '^^^ ^^^^3

LONG PALPS ON

ANOPHELES

ANTENNAE

ROBOSCIS

PATTERN ON HIND TARSUS

sx^^^^ S

SHORT PALPS ON

ALL OTHERS

' PATTERN ON THORAX

_ AEDES

— AEGYPTI

— AEDES

~" ALBOPICTUS

FIG. 32

r PATTERN ON HIND TARSUS

NO LIGHT SPOT ON VEIN 1

_ ANOPHELES

— FUNESTUS

( WING OF FUNESTUS )

LIGHT SPOT ON VEIN 1

v^ _ ANOPHELES

" — BRUNNIPES

( WING OF BRUNNIPES )

PATTERN ON PALP

_ ANOPHELES

— CHRISTYI

PATTERN ON PALP

-« ANOPHELES

— GAMBIAE

_ ANOPHELES

— PHAROENSIS

AT LEAST FOUR LIGHT SPOTS ON FRONT

MARGIN

+ i. „ .. \ _ ANOPHELES

liif^'S'iill' ~ f^UFI^ES RUriPES

(WING or HUFIPES PUFIPES )

ANOPHELES

HANCOCK/

:3.CÇ=^ =

ANOPHELES

PRETORIENSIS

WING

FRINGEADULT MOSQUITO WITH PARTS LABELED

--WING
56 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

The approximately 30 Seychelles, lying 700 miles northeast

of Madagascar, are subjected to southeast winter trade winds

from May to November and northwest summer monsoons.

The rainfall averages about 90 inches annually, with the

heaviest precipitation between November and April.

Mauritius, 500 miles east of Madagascar, has a reef-fringed

coastal plain rising abruptly in three mountain ranges on

the southeast, west, and north. These mountains enclose a

central plateau. From 75 to 175 inches of rain fall annually

on the windward coast and about 40 inches on the leeward

side. Tropical cyclones are frequent.

Réunion is of volcanic origin. It has interior uplands and

narrow valleys between the steep mountainsides. The rainfall ranges from 28 to 160 inches annually.

Reference: SOI.

Malaria.—Except for the central mountains and arid

southwest corner, malaria is highly endemic over most of

Madagascar. Abundant mosquito breeding places and yearround malaria transmission occur on the wet east coast, but

the disease is seasonal elsewhere and its distribution patchy.

The indigenous population is very susceptible in the numerous valleys of the central plateau, where the prevalence of

malaria is closely associated with rainfall and mosquito

abundance. Ricegrowing areas of the island are particularly dangerous.

Mauritius has reported complete eradication of its malaria.

After 18605 when it was introduced to that island, this disease

appeared in all areas below 1,400 feet nearly every year

between December and June. Malaria is endemic in most

of the Comoro archipelago and at low elevations in Réunion.

The Seychelles are reported to be free of the Anopheles

species and malaria.

In all these islands Plasmodium falciparum and P, vivax

infections are equally abundant in native populations and

account for nearly all cases ; about 10 percent are P. malariae

infections. Plasmodium ovale malaria is rare.

Anopheles gambiae is the principal malaria vector everywhere in this region. Particularly in Madagascar it is most

subject to seasonal changes in temperature. It does not reach

high altitudes there as in continental Africa. The gradual

increases in elevation, such as the Ethiopian highlands, are

completely lacking, and gamiiae cannot overcome the sudden

changes in temperature characteristic of the steep slopes of

offshore localities.

Anopheles funestus plays a rather minor role as a malaria

vector but is present along with gamhiae in all islands where

that species appears. In Madagascar funestus is known as

fmnestus imerinensis^ which is as effective a vector as typical

funestus. It is not treated separately in the keys.

Only one female of Anopheles ruftpes rußpes has ever been

found infected with malaria in Madagascar. It is not regarded as a vector there but is included in the keys because

of its importance elsewhere.

The role of Anopheles pretoriensis as a malaria vector is

negligible in this region. In Southern Africa as high as 2

percent of this species has been found infected. It is generally considered to be rather anthropophilic. Itis included

in the keys because of its potential ability to transmit malaria

in the islands.

Anopheles pharoensis^ naturally infected in some areas and

found to be negative in others, is very irregular in its ability

to transmit the disease. In Madagascar it is a vector of

minor importance, especially in the presence of funestus and

gamhiae.

Anopheles christyi.^ A. hancocki.^ and A. hrunnipes are not

known to be vectors in Madagascar but are included in the

keys because of their ability to transmit malaria elsewhere.

References : 61.105.

Anopheles Species in Madagascar and Adjacent Islands

mascarensis

millotti

notleyi

ohscurus obsGUTUs

pauliani

'^pharoensis

\pretoriensis

radama

ranci rouhaudi

\rufipes rußpes

squamosus cydippis

squamosus squamosus

theileri

\l)runnipes

\christyi

coustani coustani

funestus confusus

"^funestus imerinensis

fuscicolor fuscicolor

fuscicolor soalalaensis

"^gambiae

grassei

\hanGOGhi

laeani

maculipalpis

marshallii (var. not known)

See footnote 6, p. 10.

Yellow Fever.—This disease is not present in Madagascar

or the neighboring islands. A few mouse-protection tests,

all negative, have been performed.

Reference: 256.

Dengue.—This disease has been reported from Madagascar, Mauritius, and the Comoro Islands. It is said to be

endemic on the east coast and in the plateau areas of Madagascar, where a few cases are reported every year and irregularly spaced larger outbreaks occur from time to time.

The last epidemic was at the northern tip of Madagascar in

1944. No information is available on dengue in the rest of

the region. Aedes aegypti and A. alhopictus are present in

various parts of this region. They are believed to be the sole

vectors wherever dengue occurs.

References : 256, 311.

Encephalitis.—No information is available.

Filariasis.—This disease is well established throughout

Mauritius and attacks humans living along streams and in

urban areas that have open drains. In some localities up to

9 percent of the human population has been found with

microfilariae of Wuchereria iancrofti^ especially during cool

weather. Anopheles gambiae and A. funestus are believed to

be the important vectors in the island.

Reference: 1^2.

Southern Africa

(Figs. 33 and 34)

Physical Features.—Southern Africa consists of the

African mainland south of the Belgian Congo and Tanganyika. It includes Angola, Northern and Southern Ehodesia,

Nyasaland, Mozambique, South-West Africa, Bechuanaland,

and the Union of South Africa.

Angola has a narrow coastal lowland. The northern half

of this country has an unhealthy tropical climate. The vast,

jumbled inland plateau, with an average altitude of 6,000

feet, slopes gently eastward to the plains of the Congo and

Zambezi Rivers. It receives up to 60 inches of rain per year,

mostly from September to April.

Most of Northern Rhodesia, at an altitude of 3,000 to 5,000

feet, has a subtropical climate, with moderate winters and

extremely hot summers in the river valleys. The rolling uplands are savannas; the extensive lowlands in the south m

Barotseland are covered with tall grass and are poorly

drained. The average annual rainfall is 50 inches in the

north and from 25 to 30 inches in the south.

Southern Rhodesia lies on the African plateau at an elevation of 2,000 to 5,000 feet. Only the low forested veldt on

the northern fringe and in the southeast corner is below 2,000

feet. The highest ranges in the country are located along

the Mozambique border and are almost 8,500 feet. They

receive up to 40 inches of rain annually. The three seasons
MOSQUITOES OF MEDICAL IMPORTANCE 57

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE- SOUTHERN AFRICA

[AREA INCLUDED: ANGOLA, NORTHERN AND SOUTHERN RHODESIA, NYASALAND, MOZAMBIQUE, SOUTH-WEST

AFRICA, BECHUANALAND, AND THE UNION OF SOUTH AFRICA]

^ FULL-GROWN LARVAE 1^

NO SIPHON ON ANOPHELES

LONG PLEURALS

OF 3RP GROUP

BOTH

HAIRED

r CLYPEAL HAIRS

BOTH BARE 1 I

LONG PLEURALS

OF 2ND GROUP

^5U(

PALMATE HAIRS

ON 5TH SEGMENT

T

THREAD

LIKE

_ ANOPHELES

" PRETORIENSIS

_ ANOPHELES

— LISTERI

ONE HAIRED;

ONE BARE

AND HAIRLIKE

3 TO, ^Y \f.

MIDDLE '^ \

ANOPHELES

RUFIPES RUFIPES AND RUFIPES INGRAMI

WITH SIDE

HAIRS -^sj

PECTÉN SPINES

— ANOPHELES

— PHAROENSIS

CLYPEAL HAIRS

WITH LONG

HAIRS ON

JL. INNER SIDE

BARE JE"W: ANOPHELES

NIL I

_ ANOPHELES

" BRUNNIPES

SHORT _ ANOPHELES

SPINES — GAMBIAE

ALL SPINES .a,^«,„-, ,-^

ABOUT — ANOPHELES

EQUAL IN — MELAS

It LENGTH

r ANTENNA

SPICULÈS

EVENLY

DISTRIBUTED

BASAL TUBERCLES

PRESENT AND

CLOSE TOGETHER

PLATES ON 5TH SEGMENT

COVERING ABOUT

1/2 OF .^ T

SEGMENT

FIG. 33

ANOPHELES LARVA

HIND EDGE

-|- OF PLATE

NOT DARKENED

DENSE PATCH

OF SPICULES

AT BASE

SIPHON ON ALL OTHERS

OUTER CLYPEALS

LATERAL HAIRS ON

5TH SEGMENT

Ï

2 TO 4 BRANCHED

ANOPHELES

FUNESTUS FUNESTUS

— AND FUNESTUS FUSCIVENOSUS

_ ANOPHELES

— HARGREAVESI

ANOPHELES

— HANCOCKI AND MARSHALLII

/ON

r- ( L,

\ D

ONLY HANCOCKI IMPORTANT; >

LARVAE SEPARATED WITH

IFFICULTY J

SHORT,

BLUNT AT

TIP

ANOPHELES

RHODESIENSIS

__ ANOPHELES

— DOM ICOLUS

COMB SCALES

ONLY ONE

TUFT PER

SIDE, AND

ON OUTER

HALF OF

SIPHON

COMB SCALES

IN ONE

REGULAR ROW

(NOT ON A PLATE)

' PECTÉN SPINES

_ AEDES

— AEGYPTI

__ AEDES

~" SIMPSONI
58 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

LONG PALPS ON

ANOPHELES

ANTENNAE

.PROBOSCIS

SHORT PALPS ON

ALL OTHERS

MOSQUITOES OF MEDICAL IMPORTANCE - SOUTHERN AFRICA

[AREA INCLUDED-. ANGOLA, NORTHERN ANO SOUTHERN RHODESIA, NYASALAND, MOZAMBIQUE, SOUTH-WEST

AFRICAj BECHUANALAND, AND THE UNION OF SOUTH AFRICA]

1 FEMALES ^

MALES HAVE BUSHY ANTENNAE

FIG. 34

PATTERN ON HIND TARSUS

PATTERN ON PALP

AND DO NOT BITE

NO LIGHT SPOTS ON VEIN 1

i

VEIN 6 DARK ON OUTER HALF

TWO LIGHT SPOTS ON VEIN I

ilïïïïIîii'iH'Niiiilfflffm'i

• FRONT MARGIN

WITH LIGHT SPOT

IN BASAL THIRD

FRONT MARGIN

DARK ON BASAL

THIRD

_ ANOPHELES

— BRUNNIPES

ANOPHELES

- FUNESTUS

FUNESTOS

ANOPHELES

FUNESTUS

FUSCIVENOSUS

_ ANOPHELES

"~ NIL I

AT LEAST FOUR LIGHT SPOTS ON FRONT MARGIN;

REMAINDER OF WING DARK

_ ANOPHELES

- RHODESIENSIS

AT LEAST FOUR LIGHT SPOTS ON FRONT MARGIN

_ ANOPHELES

SIII'I'lllJWi!^ " LISTERI

SCALES ON THORAX

:r,^^||i-'i|llllii|llll|||l"

TWO LIGHT SPOTS ON VEIN 1

K +

ANOPHELES

H ARGREAVESI

»HiiäsiiSisa ANOPHELES

- 6AMBIAE AND MELAS

/BOTH IMPORTANT; ADULTS CANNOTN

V BE SEPARATED WITH CERTAINTY /

FRINGE SPOT AT VEIN 6

-_ ANOPHELES

— DOMICOLUS

LAST SEGMENT ENTIRELY LIGHT

^^^^S^^^^^§?^ip

PATTERN ON HIND TARSUS

_^ _ ANOPHELES

^^^^x:::>Z "" PHAROENSIS

:^^ ^

IfflTTttnin'ifffffirft

TWO LIGHT SPOTS ON VEIN 1

ONLY ONE LIGHT SPOT ON VEIN 1

ÎF PRESENT, LESS THAN 1/3 AS

.o.= «sço^^e^^^^ + Çlll, pi^«,

TWO LIGHT SPOTS ON VEIN 1

_ ANOPHELES

~" PRETORIENSIS

— ANOPHELES

" HANCOCKI

ANOPHELES

H RUFIPES

INGRAMI

MORE THAN 1/3 AS LONG AS

^ SEGMENT 3v _ ANOPHELES

— RUFIPES RUFIPES

• PATTERN ON THORAX

LINES _ AEDES

WHITE — AEGYPTI

AEDES

SIMPSONI

ADULT MOSQUITO WITH PARTS LABELED
MOSQUITOES OF MEDICAL IMPORTANCE 59

include hot dry weather from September to November, a

rainy season from November to March, and generally cool

weather from May to August.

Nyasaland lies in the Kift Valley and is flanked by high

savanna-covered plateaus at elevations of 3,500 to 8,000 feet.

The temperature in the valley is 120° F., but in the milder

uplands the temperature ranges from 40° to 95° F. The

annual rainfall averages about 40 inches. The southeast

part of Nyasaland is low and marshy.

Mozambique is a flat lowland, which rises gradually in

the west to an undulating 800- to 2,000-foot plateau, with

the highest parts in the north and along the edge of the

Rift Valley. Monsoons bring 40 to 60 inches of rainfall per

year to the northern edge of the coastal plain, mostly between

December and April. Southern Mozambique is cooler and

drier. Commercial agriculture is practiced only along the

northern river valley.

South-West Africa has an arid coastal plain contiguous

with that in Angola. This rises abruptly to an interior

2,500-foot plateau, which is bordered by northern mountains

of 8,500 feet. The dry southern and eastern parts of the

country contain many salt-water depressions.

Bechuanaland is an interior 3,000-foot plateau, the entire

southern part of which is desert. Most of the northern part

has a subtropical climate. Lake Ngami and the Okovanggo

River basin in the northwest are the country's largest marshy

areas.

The interior of the Union of South Africa is a vast rolling

plateau 2,000 to 6,000 feet high, bounded by a series of high

parallel mountain ranges extending from Cape Town around

the southern edge of the country to Mozambique. The southern part of the Union of South Africa has winter rains, but

elsewhere there are summer rains, except in the extreme

northern and western parts, which are arid.

Reference: 301.

Malaria.—The distribution of this disease is patchy in the

Rhodesias, Nyasaland, and Mozambique. The areas of hyperendemic malaria, which are rarely above 4,500 feet, with

spleen rates of 50 to 100 percent, are located along the

Mozambique coast, in the valley of the Zambezi to Lake

Natal, and in parts of both Transvaal and Natal. Latitude

31° S. represents the southernmost limit of the disease in

Southern Africa. In the grassy uplands of the Rhodesias

and Mozambique malaria is seasonal. Apparently no information is available about the distribution or intensity of

the disease in Angola, but the extreme northern parts of

South-West Africa have endemic centers. The vast expanses

of extremely dry countryside to the south support little

malaria.

PlasmodiwiYh falcipar-um is often found in epidemics, but

P. vivax appears at least as frequently ; P, malariae and P.

ovale are rare.

Anopheles gamhiae finds ideal breeding sites in the northwest part of Southern Africa, where man makes many

natural depressions in the hard soil. However, gambiae as

the principal vector is restricted to highland areas in other

parts of Southern Africa. There it supports endemic, hyperendemic, and epidemic malaria almost to the exclusion of

Anopheles funestus (see below). A. gamhiae will infiltrate

into many upland areas during the breeding season and then

completely disappear during the winter.

A, funestus^ on the other hand, normally inhabits larger

bodies of water than does gambiae^ and the malaria it transmits is less intense on account of the relatively more stable

mosquito population. Furthermore, in Southern Africa it

is responsible for most of the malaria transmitted at lower

altitudes. Hyperendemic malaria occurs in areas where

fvm^stus is the dominant species, but this hyperendemicity is

generally slightly lower than that supported by gambiae.

Several morphological varieties of funestus are found in

Southern Africa. The best known of these, fuscivenosus^

appears in Southern Rhodesia and is probably as effective a

vector as the type form.

Anopheles listeri is suspected as a possible vector in SouthWest Africa. It is widely distributed in this country. The

females often enter houses and feed on man. Because of its

endophilism and anthropophilism this mosquito may be a

vector. It is only potentially important, since its exact role

in transmission is not known.

Anopheles meZas reaches its extreme southern distribution

on the north coast of Angola, where itis associated principally

with Avicennia mangrove. In this area it is an extremely

effective vector and as important as gambiae.

Anopheles pretoriensis is normally a wild mosquito and

almost never found indoors. In Southern Rhodesia and the

Transvaal it is found naturally infected out of doors, but no

information is available on the nature of the parasites or how

they are obtained by the mosquito.

Eight additional species of Anopheles are found in various

parts of this region, sometimes in large numbers, but none

of them have been implicated in malaria transmission in any

of the countries discussed above. They have been included in

the keys because of their known importance elsewhere and

their possible importance in this region. The Anopheles

species and the areas in which they are important include

brimnipes^ Belgian Congo; domicolus^ northern Nigeria,

French West Africa ; hancocki., Uganda, Ivory Coast, HauteVolta; hargreavesi^ Nigeria; nili^ Belgian Congo, Liberia,

French West Africa, Sierra Leone; pharoensis^ Egypt,

French West Africa; rhodesiensis^ Sierra Leone; rufipes

ruftpes^ Anglo-Egyptian Sudan, Haute-Volta; and r. ingrami^ Sierra Leone.

References : 67, 259, 2ß0.

Anopheles Species in Southern Africa

marshallii musinhoi

marshallii pitchfordi

"^melas

michaeli

motùcheti moucheti

natalensis multicinctus

natalensis natalensis

\nili

njombiensis

obsGurus ohsGurus

paludis

\pharoensis

"^pretoriensis

\rhodesiensis

rivvlonmh rivulorwm

Tuarinus

\rufipes ingrami

\rufipes rußpes

seydeli

squamosus cydippis

squamosus squa/mosus

symesi

tchekedii

theileri

vinckei

malravensi milesi

walravensi waZravensi

ardensis

argenteolohatus

austenii

^bnmnipes

cameroni

cinereus

Gostalis

Goustani Goliginosus

Goustani GOustani

GOustani teneirosus

Goiistani ziemanni

demeilloni

distinotics distinGtus

\donhicolus

funestus Gonfusus

"^funestus funestus

"^funestus fuseivenosus

'^gamhiae

garnhami

\hanGOGki

\hargreavesi

implexus

jaeobi

leesoni

Histeri

longipalpis

maGuMpalpis

marshallii marshallii

See footnote 6, p. 10.

Yellow Fever.—Clinical cases of this disease have never

been reported from Nyasaland, Northern Rhodesia, Southern Rhodesia, Union of South Africa, South-West Africa, or

Bechuanaland. Endemic centers of yellow fever have been

found in the Barotse Province of Northern Rhodesia and
60 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Balovale district. Aedes simpsoni is believed to be the principal vector in these two centers. It probably b-ehaves in

much the same way as in Anglo-Egyptian Sudan (p. 40).

In the Lake Ngami area of northern Bechuanaland another

endemic center has been uncovered. The vectors there are

not known with certainty. Angola has reported no clinical

cases of yellow fever since 1900. Persons living there have

positive immunization tests only near the Congo border. In

Mozambique yellow fever is not known to occur. For additional information, see pages 2 and e3.

References : 256, Sll.

Dengue.—In Southern Africa dengue has been reported

only from Mozambique and the Union of South Africa. In

Mozambique the disease is endemic, but no major outbreak

has been reported since 1924. In the Union of South Africa

dengue is epidemic along the Natal coast, with occasional outbreaks. Aedes aegypti transmits dengue in this coastal area.

References: ^50, ^ii.

Encephalitis.—The West Nile and some newly described

encephalitis viruses are present in Southern Africa. Few

observations have been made on their epidemiologies. For

further information about mosquito-borne encephalitides,

see pages 3-5.

References: 251, Sll.

Filariasis.—No information is available.

EURASIA

North Europe and North Asia

(Figs. 35 and 36)

Physical Features.—This region of North Europe and

North Asia includes North Europe, the Union of Soviet

Socialist Eepublics (U. S. S. R.), Mongolia, Manchuria,

northern China, Korea, and Japan. It is bounded on the

south by the following countries : Spain, Italy, Yugoslavia,

Rumania, Turkey, Iran, Afghanistan, India, East Pakistan,

Assam, Burma, and China at about latitude 28° N. The

southern border includes most of the shoreline of the Caspian

Sea and the northern shore of the Black Sea. Ireland and

western France form the westernmost extension of this region

and the western edge of the Bering Sea and Bering Strait the

easternmostlimit.

North Europe and North Asia are separated from the

countries to the south by a series of high mountain ranges.

Lower mountains cover much of the southern half of this

region and merge with the relatively low northern coastal

plain in northern France, Belgium, Netherlands, Denmark,

Germany, and the northeastern shore of the U. S. S. R. The

Carpathians form an inland mountain circle around the

eastern border of Hungary, enclosing an upper part of the

Danube River basin. A relatively low, flat area extends

northeastward across all Europe from Gdansk (Danzig) to

the Black Sea and includes the vast steppes surrounding

Moscow. The Caucasus Mountains, extending from northwest to southeast between the Black and the Caspian Seas,

mark the approximate southern border of Asia between these

two bodies of water. Extensive ranges lie along the western

border of Norway from north to south, through the northern

three-fourths of England and Scotland, and along the eastern

margin of the U. S. S. R. from the Arctic Circle to about

latitude 50° N. The northern border of Asia is relatively

low, with vast expanses of tundra and permafrost. A large

series of east-west mountain ranges extend southward across

the center of Mongolia and Siberia from Afghanistan in the

southwest to the Sea of Okhotsk in the northeast. Another

mountain mass, roughly oval in shape, is in the western half

of China north of the Himalayan range. A partially submerged mountain chain lies in the Pacific Ocean east of the

continent and forms the Kamchatka Peninsula, Japan, and

the Kuriles.

Europe, open to the Gulf Stream and the North Atlantic

Drift, has an unusually mild winter for the high latitudes,

but the effect of these warm currents disappears at the Urals

and beyond. Only the northernmost parts of Scandinavia,

Finland, and North Asia are subarctic. The northern and

western plains and steppes have temperate westerlies, although summer and winter temperatures differ considerably.

In the southern and eastern parts the climate is regulated by

monsoon systems, which bring cool, dry winds blowing outward from the center of the continent in the winter and hot,

moist winds blowing toward its center in the summer.

Reference: 201,

Malaria.—Plasmodium vivax is the principal cause of

malaria in the Temperate Zone across North Europe and

North Asia, P. falciparmm tends to dominate in the more

southern subtropical area, and P. malariae is sporadic and

usually accounts for fewer cases than either of the others.

Anopheles rmessede^ one of the members of the Anopheles

maculipennis complex, is widespread north of the Alps and

inhabits great inland river valleys and large marshes. In

North Europe it rarely bites man. It is a vector of malaria

only in parts of Hungary where dense human populations

live close to large marshy areas. However, in the U. S. S. R.

it is the most widely distributed and important Anopheles

species, and because of the immense area included in its

range, it is one of the most important, if not the most important, vector in the maculipennis group. The northern

and eastern limits of the range of the maculipennis group in

North Asia are represented by the northern and eastern

limits of the ran^e of messeae. This species does not extend

south of Caucasia. There are no Anopheles species or

malaria in Iceland.

In all western Europe north of the Alps malaria is endemic

in only one small area consisting of the North Sea Coast

Provinces of North Holland, Friesland, Zeeland, and German East Friesland. This endemic Plasmodium vivax

malaria appears in epidemic waves about 20 years apart and

is carried by Anopheles labranchiae atroparvus. Its larvae

are found in slightly brackish coastal waters, and the adults

inhabit warm shelters throughout the winter. The species

transmits a great deal of so-called winter house malaria, presumably because the numerous children are the most persistent carriers of malaria. Outside of this area atroparvus

is not known to be a malaria vector, except in southwest

U. S. S. R. on the Black Sea coast between the Bug and

Danube River Deltas. The species over much of its range is

greatly attracted to stable animals. The easternmost limit

of the distribution of malaria in the Black Sea area corresponds to the easternmost limit of the range of atroparvus.

Anopheles labranchiae labranchiae is not represented at

all in North Europe, but its ç^gg is illustrated in the key, because this species does inhabit the northern parts of Italy and

Spain close to the borders of France.

Anopheles melanoon melanoon has been reported from

Transcaucasia just south of the Caucasus Mountains but is

not believed to contribute to endemic malaria anywhere in its

range. Anopheles melanoon subalpinu^ has been reported

from Switzerland, Hungary, and in large numbers from the

lowlands of Transcaucasia on the southern shore of the Caspian Sea. In the vicinity of Lenkoran and the Sefid Rud

River Delta it is the dominant and often the only member

of the maculipennis complex, and it is the principal vector of

a severe malaria. The female hibernates near its source of

food and, like atroparvus^ feeds throughout the winter.

This mosquito is not found north of Caucasia.

Anopheles maculipennis has never been reported from

England but is widely distributed in continental Europe, ap-
MOSQUITOES OF MEDICAL IMPORTANCE 61

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE- NORTH EUROPE AND NORTH ASIA

[AREA INCLUDED: NORTH EUROPE^ THE U.S.S.R., MONGOLIA, MANCHURIA, NORTHERN CHINA, KOREA, AND JAPAN]

-^^ FULL-GROWN LARVAE ^

NO SIPHON ON ANOPHELES

INNER CLYPEALS Í

BASES CLOSE TOGETHER

CLYPEALS Ï

INNER BARE OR WITH

FINE SIDE HAIRS

^¿ OUTER

BRANCHED,

NEAR TIP .'

SADDLE HAIR

FIG. 35

ANOPHELES

CLAVIGER

_ ANOPHELES

"" PLU MBEU S

INNER BARE

ANTENNAL HAIR

AT ABOUT MIDDLE^

OF SHAFT,/

LONG ANOPHELES

S/NENS/S

ANOPHELES

MACULIPENNIS

— AND RELATIVES

TOP VIEW OF HEAD OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS

OUTER HIND

CLYPEALS ,, CLYPEALS

V BASES FAR APART

OUTER CLYPEALS ABDOMINAL PLATES 4 TO 7 PALMATES ON SEGMENTS 2&3

4- SMALL

tr~i/^

PALMATES ON SEGMENT i

WITH FINE

SIDE HAIRS ,j

PALMATES ON

SEGMENT 2

-|- SMALLER

THAN ON

SEGMENT 3

«. ANOPHELES

"" PAT TONI

_ ANOPHELES

" SUPERPICTUS

BRUSH-LIKE \

__ ANOPHELES

— PULCHERRIMUS

SIPHON ON ALL OTHERS

BARREL SHAPEDÎ ONLY

1 TUFT PER SIDE,

AND ON OUTER

HALF COMB SCALES

, IN ONE ,

"T" REGULAR ROW "•"

(NOT ON A PLATE)

COMB SCALES

AEDES

AEGyPTI

AEDES

ALBOPICTUS

ANOPHELES \^fK^MfK

PLEURAL BRISTLES

/ON UNDER SIDE WITH A BASAL\

SHEATH^

SPINDLE SHAPED;

7 TO 10 TIMES AS^

LONG AS WIDTH^

AT BASE

COMB SCALES PECTÉN SPINES

5 OR 6 TUFTS

PER SIDE IN

EVEN ROW

/

_ CULEX

"" TRITAENIORHYNCHUS ^PALMATES ON

*V Mi p!^¿?-a" S EGM E NTS

EGGS OF ANOPHELES MACULIPENNIS AND RELATIVES {MCLANOON MELANOON NOT IMPORTANT]

ROUGH

TOP \

SURFACE V i

MACULIPENNIS

MACULIPENNIS

LABRANCHIAE LABRANCHIAE MELA NOON MEL A NOON

LABRANCHIAE ATROPARVUS SUBALPINUS MELANOON

SACHAROVI

BDOMINAL

PLATES

4 TO 7

SADDLE HAIR
62 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - NORTH EUROPE AND NORTH ASIA

[AREA INCLUDED; NORTH EUROPE, THE U.S.S.R., MONGOLIA, MANCHURIA, NORTHERN CHINA, KOREA, AND JAPAN]

— m FEMALES:

[MALES HAVE BUSHY ANTENNAE ( J^^P^ ) AND DO NOT BITE\

FIG. 36

LONG PALPS ON

ANOPHELES

ANTENNAE

.PROBOSCIS

PATTERN ON WING

AT LEAST FOUR LIGHT SPOTS ON FRONT

MARGIN

PATTERN ON HIND TARSUS Ï

ONE OR TWO LIGHT SPOTS ON BASAL

FOURTH OF FRONT MARGIN

iiiïiiiiTTOÎrfflTffiîrrt

„V _ ANOPHELES

\\\\\\ü:'m0^) — suPERPicTus

_ ANOPHELES

— PA TTONI

_ ANOPHELES

— PULCHERR/MUS

ONLY TWO LIGHT SPOTS ON FRONT MARGIN

ANOPHELES

S/NEA/S/S

( WINO OF S/fiCNSIS )

ENTIRELY DARK, OR LIGHT ONLY AT TIP, WITH

FOUR DARKER SPOTS

FRINGE AT WING TIP

ANOPHELES

MACUUPENNIS AND RELATIVES

/ NOT ALL IMPORTANT; ADULTS CANNOTN

V BE SEPARATED /

\.. n*nu — ANOPHELES

^ALLDARK __ ^ACHAROVI

TOP VIEW OF HEAD

ENTIRELY DARK, WITHOUT DARKER SPOTS

TUFT OF

a. WHITE

SCALES

( WING OF CLAVieeR )

END SEGMENT NOT MORE THAN

/ 1/2 AS LONG AS NEXT,

\. AND ANO SAMt SAME ^,UL COLOR ^

^-. ^-

END SEGMENT ABOUT 3/4 AS

/ LONG AS NEXT, AND SAME

COLOR • ^

ANOPHELES

CLAVIGER

ANOPHELES

• PLUMBEUS

SHORT PALPS ON

ALL OTHERS

PATTERN ON THORAX

CULEX

TRITAENIORHYNCHUS

SCALES

BROAD AND

-j- YELLOWISH

^(NOT WHITE)

_ AEDES

— AEGYPT/

_ AEDES

— ALBOPICTUS

ADULT MOSQUITO WITH PARTS LABELED

THORAX^^
MOSQUITOES OF MEDICAL IMPORTANCE 63

pearing as far north as southern Sweden and as far east as

the plains of western Siberia. The eastern boundary of its

range is generally the Urals. It lives primarily in mountainous country and is found only in the hilly parts of flatter

areas. In Europe it is not known as a vector at all, but it is

the chief or sole vector, maintaining a high rate of transmission, in Caucasia and Transcaucasia, where it often hibernates, like atroparvus and s'uhalfinus^ in warm shelters and

may be responsible for some cases of winter house malaria in

this area. In Transcaucasia maculipennis may consist of

local races, since several ^gg varieties have been described.

Anopheles sacharovi is not found in Europe proper north

of the Alps. It is largely restricted to the Mediterranean

area in the west, but eastward its range swings slightly north.

It is common throughout most of Transcaucasia, along the

western littoral of the Caspian Sea, Daghestan, throughout

the less arid parts of Turkestan, and in Sinkiang Province,

China. The larvae are often found in brackish or saline

water. This mosquito is a dangerous vector throughout

most of its range. However, in the lower part of the Amu

Darya Valley it is present in large numbers in the complete

absence of malaria. Reasons for this are not well understood. Extensive cattle breeding may divert this mosquito

from man.

Anopheles claviger is a widely distributed mosquito. Its

range extends from England and Denmark through North

Europe and North Asia and into Siberia as far east as the

Yenisei River. It is common in central Asia and is prevalent in Caucasia on the Black Sea coast. It is an effective

vector only in the southern part of its range and then only

where it breeds in spring water in the mountainous areas

close to large human populations. It has maintained a mild

endemic malaria in the suburbs of Baku, where it breeds in

abandoned wells.

Anopheles plumheus is distributed throughout North

Europe, and its range extends as far east as Tadzhikistan.

Since it breeds in tree holes, it is primarily a forest mosquito

and rarely has high enough populations to maintain malaria.

However, in the forested areas east of the Black Sea coast,

it has transmitted malaria, although insignificantly. This

species is included in the keys because of its potential importance.

Anopheles pulcherrimus is restricted to North Asia (see p.

132). It inhabits chiefly low land and is common in the flat

valleys of the large ri vers. Itis most commonly encountered

in central Asia, especially in the lower part of the Murgab

Valley, where it breeds in tremendous numbers, building up

populations early in the year. It is able to maintain severe

endemics in the absence of other vectors. In most of its

range it is a vector of malaria to some extent.

Anopheles superpictus^ which is absent from Europe north

of the Alps, is restricted largely to the Mediterranean.

However, it is found abundantly along the west Caspian seacoast and is very common eastward throughout the mountainous and hilly areas of central Asia. Like pulcherrimus^

it is able to maintain heavy endemics in the absence of other

vectors, especially in many localities of Turkmenistan.

Anopheles sinensis^ the eastern representative of the A,

hyrcanus group, is restricted to eastern Asia and the Orient.

The northwest limit of its range is represented by a line

drawn across Asia from the northern Assam-Burma connection with China to the northeast border of Manchuria. Its

range covers all the territory east of this, including Japan.

In the Ussuri area and the lowlands of northern China it is

the only Anopheles species, and in the latter area it is able

to maintain rather severe endemic malaria. In Japan and

Korea it is the principal vector of malaria.

Anopheles pattoni^ another eastern Asian mosquito, is common in the hilly areas of northern China. It is the chief

vector in southern Manchuria, Hopeh, Shantung, Shansi,

471779 O—59 5

^maculipennis

marteri

melanoon melœnoon

'^nhelanoon subaHpinus

^messeae

moghulensis

"^pattoni

"^plumbeus

^pulcherrimus

pullus

"^sacharovi

"^sinensis

sineroides

"^superpictus

yatsushiroensis

Shensi, northern Szechwan, and eastern Kansu. This species has not definitely invaded the Japanese islands.

References : Ö7,122,125,115, 223, 262, 264.

Anopheles Species « in North Europe and North Asia

aitkenii

algeriensis

chodukini

"^claviger

\fluviatilis

gigas iaileyi

gigds simlensis

hyrcœrms

koreicus koreicus

koreicus var. hisaoe

Habranchiae atroparvus

flabranchiae labranchiae

lewisi

lindesayi japonicus

lindesayi lindesayi

\maoulatus

Yellow Fever.—Outbreaks of yellow fever in temperate

regions probably represent introductions from tropical regions, where this disease is believed to have originated as

jungle yellow fever. Epidemics were reported in Europe

during the 18th century in port cities of France and England

and were probably borne by Aedes aegypti. Yellow fever

has never been reported from the Orient.

Reference: 316.

Dengue.—The distribution of dengue is discontinuous in

North Europe and North Asia. A few cases are reported

from southern Austria each year, and a small epidemic

occurred in Vienna in 1941. An important endemic center

surrounds the Black Sea and extends in a wide belt across

Caucasia to encompass the Caspian Sea, beyond which it

reaches to the western border of China and is continuous

northward to about latitude 54° N. It is assumed that Aedes

aegyptiis the vector in this area.

Dengue also occurs along the Asian coast opposite Sakhalin

and along the Yellow and East China Sea coasts of China,

and it extends well inland. It is probably not a military

problem in Korea.

Epidemics of dengue in the Nagasaki, Sasebo, and OsakaKobe areas of Japan occurred in 1942, recurred every summer

for several years, and ended in 1945. This was one of the

largest series of epidemics ever recorded in the temperate

region, almost 200,000 people having been affected. Aedes

aegypti and A. aïbopictus were the vectors of this eastern Asia

dengue. Both are found over much of this area and are

known vectors of the disease elsewhere.

References: 171,256,310,348.

Encephalitis.—Human cases of Japanese encephalitis,

some of them unconfirmed, have been reported in an area

extending from Sakhalin to Hankow in eastern China. For

additional details, see page 4.

Reference: 257.

Filariasis.—The distribution of filariasis in this region is

not completely known, but Japan, South Korea, the Yangtze

Eiver basin, and other parts of coastal China are known

centers of infection. Among the large number of mosquitoes

that have been experimentally infected with Wuchereria ha/n-

' Species marked with an asterisk (*) are recorded as vectors of

malaria and/or filariasis in this region and their medical importance

is discussed in the accompanying text. Those marked with a dagger

(t) are not recorded as vectors of malaria and/or filariasis in this

region hut are important elsewhere. For information on the latter

vectors, see the description of the particular species in the last section

of this handbook.
64 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

GTofti in Japan, Culex pipiens pallens and Aedes togoi are

outstanding. Filariasis parasites mature rapidly in these

two species, which are especially domestic and are commonly

found with infectious stages of iancrofti in nature. A, togoi

and possibly pallens have been found to transmit Wuehereria

rrholayi in the Japanese islands not more than about 80 miles

south of Tokyo. This parasite is present in almost pure form

in a focus entirely isolated from the closest centers of infection of malayi. A. togoi breeds close to houses in concrete

rainwater reservoirs in this area.

Filariasis caused by malayi and hancrofti is present in most

of the coastal provinces of China and along most of the

Yangtze River and its delta, where Anopheles sinensis is

probably the most important vector of both of them. Filariasis is an important disease in the country villages of the

Woosung district, where sinensis is commonly found with

natural infections. Culex tritaeniorhynchus may be a minor

vector from mid-July to mid-August in central China, where

it is endophilic, but in the Shanghai area it is not common.

Armigeres obturhans is one of the most common domestic

mosquitoes of the lower Yangtze Valley and may transmit

some filariasis, since it has been found with a low rate of

natural infection.

In the Shanghai area Culex pipiens pallens is one of the

most important vectors of hancrofti^ but pallens has a low

susceptibility to infection with malayi. Attempts have been

made to infect a number of mosquitoes in the Shanghai area

with filarial parasites. Field and laboratory observations

suggest that the following species are not vectors in nature

there : Culex hitaeniorhynchus^ C. fuscanus^ C\ pallidothorax^

0, vexans var. nipponii, and Aedes albopictus.

References : ^0,123,111-182,184-187,208,227,342, S43.

Spain and Portugal

(Fig. 37)

Physical Features.—Spain comprises most of the Iberian

Peninsula. This country is roughly triangular in shape and

is bounded largely by water. The Galician Mountains in

the extreme northwest extend along the northern border.

Along the steep, rocky shores of the Bay of Biscay, which

has the highest rainfall in Spain, is another range, the Cantabrian. These mountains merge with the high Pyrenees,

which divide Spain from France. The entire southeast

coast on the Mediterranean Sea consists of many beaches,

lagoons, and marshes and has hot, dry summers and wet

winters. The extreme southern part of the peninsula contains the highest mountains in Spain, the Sierra Nevadas. A

massive plateau, having a Temperate Zone climate, covers

most of the interior. Lowlands are in the vicinity of Valencia, Murcia, and Alicante on the Mediterranean coast.

The principal fertile river valleys in the interior are in Old

Castile, New Castile, and Aragon.

Portugal's entire west coast faces the Atlantic Ocean and

is generally level and sandy except for numerous rocky

bays in the north. A series of northeast to southwest mountain ranges extend inland from the eastern border and cover

most of the northern half of the country. The extreme

north, which is relatively undeveloped and very cold in

winter, is divided by the Tagus River from the southern half,

which is flatter and more arid. Algarve is a subtropical

coastal strip. Lisbon, situated at about the southern third

of Portugal, has a mean annual temperature of about 60° F.

and an annual rainfall of about 60 inches.

Reference: 301.

Malaria.—No information is available on the Plasmodium

species in Spain and Portugal.

By far the most important vectors of malaria in these

countries are Anopheles labranchiae labranchiae and A, I,

atroparvus. The former mosquito is restricted to the southeast corner of Spain. Since its range covers a much smaller

area than that of atroparvus^ it may be said to be less important than that species. In that area it appears in the almost complete absence of other Anopheles species, and it

transmits a malaria with a much higher endemicity than

in other parts of the country. The larvae live in numerous

habitats, including fresh water of ricefields and streams and

brackish coastal marshes. The females enter houses in large

numbers to engorge on human blood. It is believed that the

malaria transmitted by this species will not disappear so

long as labranchiae remains.

In contrast to labranchiae^ often stated to be a vector of

intense malaria, atroparvus is a vector of extensive malaria,

being the most widespread mosquito in Spain and Portugal.

The larvae typically are found in brackish water along the

coast and also in fresh water farther inland. The adults may

feed on sheltered animals but prefer the blood of man. They

are responsible for w^inter house malaria. The most malarious part of Spain is the northwest part of Cáceres at the

southwest end of the central mountain range near Portugal.

Here atroparvus is the only known mosquito. It has been

found w^th a low rate of infection, but its large numbers

here, as elsewhere, account for its importance. In Portugal

only the ricegrowing areas of the Sado River in the south,

the Tagus River in the center, and the Mondego River in the

north have malaria in a highly endemic or hyperendemic

form. These areas normally support large populations of

atroparvus.

Anopheles hispaniola appears throughout the southern

half of the peninsula and has been implicated as a vector by

presumptive evidence. The larvae prefer clear, sunlit water

in either small pools or slow streams, nearly always in association with Spirogyra. Little is known of the biting habits

of the females.

References: 28, 67, 76, 336.

Anopheles Species in Spain and Portugal

algeriensis

\claviger

^hispaniola

hyrcanus

Habranchiae atropar-vus

^labranchiae labranchiae

\maculipennis

marteri

melanoon melanoon

\melanoon subalpinus

\messeae

\nudticolor

\plumbeus

portucaliensis

\sergenti

\super pictus

See footnote 6, p. 10.

Yellow Fever.—Aedes aegypti is the only known disease

vector of the nonanopheline mosquitoes in Spain and

Portugal. It transmits urban yellow fever and proliferates

in close proximity to man, breeding in all kinds of artificial

containers. Although this disease was last reported in 1890,

its vector is included in the keys because the disease might

be reintroduced and because large numbers of aegypti mosquitoes are present, especially in the port cities. This species

is widely distributed in Spain and Portugal.

References : 32, 36.

Dengue.—Likewise transmitted by Aedes aegypti, dengue

is present but has an extremely low endemicity, principally

in southern Spain. For this reason it does not appear in

current disease statistics. Most Portuguese authorities believe that it never entered Portugal. Its vector is widespread

in both countries.

References : 32, 36, 256.

Encephalitis.—No information is available.

Filariasis.—No information is available.
MOSQUITOES OF MEDICAL IMPORTANCE 65

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - SPAIN AND PORTUGAL

—^ FULL-GROWN LARVAE ^^

LONG PLEURALS

OF 2NP GROUP

HEAD OF ANOPHELES \.fKK\lk

ANTENNAL HAIR ^"f' OUTER CLYPEALS

ANOPHELES

LABRANCHIAE

AND RELATIVES

(LARVAE CANNOTX

BE SEPARATED - \

SEE KEY TO /

EGGS BELOW /

_ AEDES

— AEGYPTI

^ EGGS ^

FRILL AROUND EGG

INTERRUPTED BY

FLOATS AT MIDDLf^

FLOATS AT

J. LEAST 1/3

^ AS LONG AS

EGG —

FRILL AT

WIDEST PART

OF EGG

PATTERN

+ ON TOP OF

EGG

ANOPHELES

LABRANCHIAE ATROPARVUS

/FOUND THROUGHOUT \

V SPAIN AND PORTUGAL y

_ ANOPHELES

" LABRANCHIAE LABRANCHIAE

/FOUND ONLY IN \

I SOUTHEAST SPAIN >/

FEMALES

LONG PALPS ON

ANOPHELES

WITH NUMEROUS LIGHT SPOTS

\_MALES HAVE BUSHY ANTENNAE {j^^^ ) AND DO NOT BITE'\

PATTERN ON PALP
66 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

Italy, the Balkans, and Adjacent Islands

(Fig. 38)

Physical Features.—Italy, extending for 600 miles into

the Mediterranean Sea, is separated from France, Switzerland, and Austria by the Alps. The high Maritime and

Graian Alps on the French frontier are contiguous with

the volcanic Apennine range, the backbone of the peninsula,

which extends the length of the country. Farther south

these mountains split off into plateaus, encircling lowlands

and extensive marshes. The Po River basin, situated between the Alps and the Apennines, is one of the country's

most fertile areas and has a continental climate with marked

changes in season. The Pontine Marshes on the west coast

near Rome are additional lowlands. The east coast is much

drier than the west, which is wet and lined with islands.

Sicily, Sardinia, and Corsica are large mountainous islands

in the Mediterranean Sea. Sicily, the largest of these, is a

continuation of the Apennines. Itis drained by intermittent

streams. Sardinia, the second largest, has a lowland in the

southwest. Corsica is a jumble of short mountain chains,

with a rocky coast. The climate of all three tends to be hot

in the summer but mild during the rest of the year.

The Balkans include Yugoslavia, Albania, Rumania, Bulgaria, and Greece. Yugoslavia, lying at about the same latitude as Italy, is separated from that country by the Adriatic

Sea. Off the western coast of Yugoslavia are the islands of

Dalmatia, which have a mild, equable climate. The Danube

and its tributaries form a large fertile basin in the northeast quarter, where most of the population and agriculture

are centered. The Dinaric Alps are in the remainder of the

country. Continental cold winters and hot summers prevail, although Macedonia and Montenegro support such subtropical crops as citrus, figs, and olives.

Albania fronts on the Adriatic Sea. In its eastern twothirds, most of which is over 3,000 feet, is a southward extension of the Dinaric Alps from Yugoslavia. These mountains have a continental climate and considerable extremes of

temperature and moisture. Most of their rivers are dry during the summer. The western third is more fertile, having

large alluvial plains and a low, rather humid coast, with 40

to 80 inches of rain annually.

The Carpathian Mountains extend in a southeast direction

from the U. S. S. R. through the center of Rumania, then

swing abruptly westward as the Transylvanian Alps to the

border of Yugoslavia. The plateau of Transylvania at about

1,000 to 1,600 feet lies north and west of these mountains.

In Rumania south of the Transylvanian Alps is the northern

half of the Danube River basin. In the entire eastern fourth

of the country from north to south are the low valleys of

Danube tributaries flowing from the north. The Danube

turns northward in southeast Rumania to form an extensive

delta, much of which is low marsh. The climate varies from

occasional severe drought in the summer to cold and windy

in the winter. The rainfall averages 17 to 30 inches annually,

and most of the area is rather fertile.

The Balkan Mountains, extending from east to west across

central Bulgaria, mark the southern boundary of the fertile

Danube River basin and form the northern boundary of the

Maritsa River basin, Bulgaria's other principal agricultural

area. South of this great valley lie the Rhodope Mountains,

which cover most of the southern part of the country. The

climate is mainly continental, with hot summers and long

cold winters, although the numerous river valleys are slightly

more protected.

Greece, in the southernmost part of the Balkan Peninsula

is irregularly shaped and has a rocky, indented coastline and

hundreds of small outlying islands in the Aegean Sea and the

Sea of Crete. It is predominantly mountainous, with a

southern extension of the Dinaric Alps from Yugoslavia and

Albania throughout its length. The few lowlands are located

in northern Macedonia, Thrace, and Thessaly and are the

principal agricultural areas. The rainfall is usually heaviest

in the west and decreases in the east. The climate is largely

Mediterranean. The principal population centers are on the

southern coast and in the islands.

Crete, the largest of the Greek islands, is very mountainous and its coasts are rocky. The climate is mild, and rainfall is much higher in the western than in the eastern part.

Cyprus is the third largest Mediterranean island, with two

mountain ranges, one along the north coast and one parallel

to it in the center. Between these two ranges is the country's

principal agricultural area. An average rainfall of 21 inches

occurs, mostly from October to March.

Reference: SOI.

Malaria.—This disease is most intense in Italy and Greece.

Approximately 68 percent of the population of Greece lives

within endemic areas, and general or local epidemics occur

every 2 to 5 years in Macedonia, Thrace, and Epirus.

Plasmodimn vivax and P. falciparum are the only malaria

parasites causing infections in Italy and the Balkans. In

Sardinia and elsewhere in this region Plasmodmm vivax infections are common in infants, whereas falciparum infections tend to predominate in all other age groups.

Anopheles labrancMae atroparvus^ one of the region's most

abundant mosquitoes, is not always associated with the transmission of malaria. It is distributed throughout the northern mainland of Italy, northern Yugoslavia, northern Bulgaria, and most of Rumania, but its importance is not known

with certainty, except in parts of Rumania where it is an

important vector. This mosquito tends to be a coastal form

but also breeds inland, where it seeks warm winter shelters

in which to feed.

Anopheles lahranchiae lairanchiae^ principally a western

Mediterranean mosquito, is centered geographically in Sicily,

from which it spreads west, south, and east. In äicily, Corsica, and Sardinia this mosquito breeds in both brackish and

fresh water. It is an extremely important vector, because it

uses those breeding habitats that are universally present in

spite of seasonal changes. It is the principal vector in Elba,

where malaria has existed for several years. However, in

Italy it is strictly a brackish-water coastal species, appearing

south of latitude 43° N. It transmits a severe endemic

malaria, which varies in intensity with the rainfall. A, I,

labranchiae has been reported from the west coast of Yugoslavia, but so far its importance there is not known with certainty. This beachhead represents the easternmost limit of

its range.

Anopheles sojcharom^ likewise principally a brackish-water

breeder, is found with labranchiae in Italy, Sardinia, and

Corsica. In the northern part of its range it is much scarcer

^md transmits much less malaria than labranchiae^ but it is

still the most important vector in the Po River Delta and

along the Adriatic coast. The range of sacharovi., unlike

that of labranchiae^ extends from Spain far to the east of

Italy. It is a common and dangerous vector in the southern

half of Yugoslavia, a southern area of Rumania, and in all

the Balkan Peninsula south of these countries. Itis the most

important vector in Greece. It breeds in early and middle

summer in brackish marshes close to the sea, and it transmits most of the malaria in Greece and along the low Dalmatian coast, where the intensity of malaria is closely associated

with the abundance of this species.

The range of Anopheles superpictus closely parallels that

of sacharovi^ and superpictus is second in importance only

to that species. Like sacharovi^ its range reaches its western

limit in Italy and Spain, but it is not important in these

countries at all. A, superpictus is essentially a foothillstream breeder and appears slightly later than sacharovi.

These two species together transmit malaria from early
MOSQUITOES OF MEDICAL IMPORTANCE 67

MOSQUITOES OF MEDICAL IMPORTANCE- ITALY, THE BALKANS, AND ADJACENT ISLANDS

IMPORTANT

A SPÉCIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

[AREA INCLUDED: ITALY, SICILY, SARDINIA, CORSICA, YUGOSLAVIA^

BULGARIA, GREECEj CRETE, AND CYPRUs]

ALBANIA, RUMANIA,

FULL-GROWN LARVAE

FIG. 38

NO SIPHON ON ANOPHELES

SPIRACULAR

APPARATUS

OUTER CLYPEALS

NOT

ANTEN NAL HAIR

ANOPHELES

MACUUPENNIS

AND RELATIVES

(LARVAE CANN0T\

BE SEPARATED- I

SEE CHART OF /

EGGS BELOW /

ANOPHELES

SUPERP/CTUS

HEAD OF ANOPHELES LARVA

INNER CUYPEALS

AEDES

AEGYPTI

EGGS OF ANOPHELES A//4C¿/¿/P£/V/\//S AND RELATIVES [MELANOON MELANOON NOT IMPORTANT]

FRILLN

MACULIPENNIS MESSEAE LABRANCHIAE LABRANCH/AE MELANOON MELANOON

LABRANCHIAE ATROPARVUS SUBALPINUS MELANOON

SACHAROVt

ANOPHELES LARVA

PLEURAL BRISTLES

^_ J*-^^"-^ ABDOMINAL

J y/y "^^^^fftz^r^^-^^ P L AT E s

\*^ *\J^^r-^ TO 7

LONG PALPS ON

ANOPHELES

ANTENNAE

[ FEMALES ^

\_MALES HA\/E BUSHY ANTENNAE ( ]^^^^ ) ^^^ ^^ ^^^ B/TE"]

( WING "OF SAC»^/fOI^/ ]

PATTERN ON PALP

. LIGHT AT TIP

FRINGE AT WING TIP

'y

ALL - ANOPHELES

DARK - SACHAROY/

AND RELATIVES

/ADULTS CANNOT BE

SEPARATED-SEE^

OF EGGS ABOVE

( S^PARATEp^SEE, CHART )

VEIN 3 WITH LONG LIGHT AREA s
68 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

spring to late fall. Although superpictus is of secondary importance, it maintains an extremely high rate of endemic

malaria in Cyprus. It is also an important vector in southern Bulgaria. Its population peak in Greece usually occurs

in September.

Anopheles messeae is widespread over Asia and is important in the transmission of malaria wherever cattle or other

domestic animals are not present. The Balkans represent

the southwesternmost limit of its range. There it is important only above the Danube Delta in eastern Rumania and in

those parts of the Danube Valley where animal husbandry

is not practiced and in the vicinity of "Lake Malik" in Albania. In the last-named locality it maintains malaria at a

rather low level and is always diverted from man when animal blood meals are available. It hibernates in cold, foodless shelters.

Anopheles maculipennis is not considered to be a species of

primary importance, although it is common to both the Iberian and Italian peninsulas, where it breeds in a wide variety

of water collections. It is included in the keys because it

maintains mildly endemic malaria in Macedonia in the absence of domestic animals, and, with messeae^ it transmits

endemic malaria in villages at "Lake Malik" on the AlbaniaYugoslav frontier. Since its habits are comparable to those

of messeae^ this species is potentially able to transmit the disease elsewhere in the Balkans as well.

Neither Anopheles melanoon melanoon nor A, m. subalpinus is regarded as important. The former is restricted to

southern Italy, and the latter appears in northern Italy and

throughout the Balkans.

References: S5, 61, 72, 236, 237.

Anopheles Species in Italy, the Balkans, and Adjacent

Islands

melanoon melanoon

j;melanoon suhalpirms

^messeae

\multÍGolor

\plumbeus

"^saeharovi

\sergenti

^superpictus

turJchudi

algeriensis

atheniensis

\clamgeT

hellenicus

fhispaniola

hyrcanus

Habranchiae atvopar-vus

Hahranchiae lahranchiae

"^maculipennis

marteri marteri

See footnote 6, p. 10.

Yellow Fever.—No information is available.

Dengue.—This disease has a very low endemicity in Italy

and Yugoslavia, but it appears in sporadic form along the

Dalmatian coast, in Albania, and in Greece, where there have

been at least two serious epidemics. The first, in the islands

of Syros and Crete, occurred in 1881. In 1927 and 1928 an

epidemic of dengue that swept Greece resulted in li/^ million

cases and was marked by unusual severity. There have been

few records of dengue since 1929. Aedes aegypti^ the urban

vector of dengue, is widely distributed throughout Greece.

References : 29, SO, SS, 256.

Encephalitis.—The presence of western equine encephalitis virus has been suspected in humans and horses in Sicily.

No detailed investigations have been carried out to determine

the vector.

Reference: 257.

Filariasis.—No information is available.

The Near East

(Figs. 39 and 40)

Physical Features.—The Near East includes Turkey,

Syria, Lebanon, Israel, Jordan, and Iraq. Turkey consists

of a partly wooded coastal strip backed by high mountain

ranges that enclose high steppelike uplands, with an average

altitude of 2,500 feet. These uplands are semiarid, with

severe snowy winters and hot, dry summers. The southeast

part of the country is largely desert. The west and south

coasts have an equable Mediterranean climate, with 20 to 80

inches of rain annually. The eastern Black Sea coast receives up to 100 inches of rain annually.

Syria is mostly desert except for the Tigris and Euphrates Valley. The northwest part consists of mountains

and valleys of the Anti-Lebanon range. They extend southward along the Lebanon border to the southwestern Hauran

plain, just north of which is an irrigated area that surrounds

Damascus. Although the annual rainfall is extremely low

in the eastern part, the coastal and mountain areas of the

west receive considerable precipitation.

Lebanon is separated from Syria by the Anti-Lebanon

range. Near the coast another series of mountains, the Lebanons, extend southward, on one side of which is a Mediterranean coastal plain and on the other a high, fertile valley,

the Bekaa.

The State of Israel lies directly south of Lebanon, with its

western border entirely on the Mediterranean. Its sandy,

level coastal plain has a dry, subtropical climate with winter

rams. Parallel with the coast and the Anti-Lebanon and

Lebanon Mountains is a central highland, which has cool,

dry summers and cold, rainy winters.

In the western half of Jordan is a distinctive depression

containing the Jordan River and the Dead Sea, the latter

more than 1,000 feet below sea level. The mountains slope

§radually eastward from this depression and merge with the

yrian Desert. About 85 percent of the population lives in

the western 5 percent of the country, which has hot, dry summers and mild, cool, wet winters. The Jordan Valley supports a growth of subtropical cultivated plants.

Highlands in northeastern Iraq rise to above 10,000 feet.

In the western half is a part of the vast Syrian Desert and

in the east the fertile valley of the Tigris and Euphrates

Rivers, in which lie most of the productive areas and the

major population centers. The lowest reaches of these two

rivers run through an alluvial plain containing marshes,

large lakes, and canals.

Reference: 301.

Malaria.—The south and west coasts of Turkey, as well

as scattered areas along the Black Sea coast, have a high rate

of endemic malaria. Some part of every province in this

country is malarious,

Plasmodium vivax, P, malariae^ and P. falciparum infections are almost equally divided in the Near East, with any

one of the parasites predominating in certain areas.

Throughout the Near East Anopheles sacharovi^ A, superpictus^ and A, maculipennis are the principal vectors. In

Turkey sacharovi is not restricted to brackish water near the

coast but is widespread over most of the country except in

the arid southeast part of the Anatolian plateau, where there

is relatively little or no malaria. Farther east and south in

Lebanon, Syria, Israel, and Iraq this species forsakes its

brackish-water habitat and becomes an entirely fresh-water

mosquito. Itis responsible, with super^ictus^ for the intense

malaria along the east coast of the Mediterranean, the valley

between the Lebanon and Anti-Lebanon ranges, and farther

east and south where periodic flooding of the Tigris and Euphrates creates extensive breeding areas near Basra. Elsewhere in the Near East sacharovi is present in smaller numbers and contributes its share of transmission.

The ranges of superpictus and sacharovi overlap throughout almost the entire Near East. However, superpictus

breeds in foothill streams ; consequently, it develops later in

the breeding season when riverbed pools form with the general lowering of water levels in hot weather. It is the prin-
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE - THE NEAR EAST

[AREA INCLUDED: TURKEY, SYRIA, LEBANON, ISRAEL, JORDAN, AND IRAQJ

^ EGGS AND FULL-GROWN LARVAE ;

69

NO SIPHON ON ANOPHELES

PALMATE HAIR

/ PLATE

SIPHON ON ALL OTHERS

LONG PLEURALS

OF 3RD GROUP

OUTER CLYPEALS

SINGLE TO

FOUR

BRANCHED

SADDLE HAIR PALMATE HAIR

FIG. 39

BLADES

+ TAPERING

TO APEX

ANOPHELES

CLAVIGER

ANOPHELES

MAOJLIPENNIS AND RELATIVES

LONG PLEURALS

OF 2NP GROUP ANOPHELES

— SERGENT/ AND RUPfCOLUS

PALMATE HAIR

LONG PLEURALS

OF 2ND_GR0UP

OUTER CLYPEALS

FILAMENTS ^

1/3 OR MORE

AS LONG AS

BLADE -

FILAMENTS—--

LESS THAN

1/3 AS LONG

AS BLADE

TOP VIEW OF HEAÓ OF

ANOPHELES LARVA

ANTE

HAIF /

INNER

CLYPEALS

OUTER 1

CLYPEALS 1

HIND

CLYPEALS

-\|M, j1'iN%'Y// 1M^i %«

v^ /

f)

ANOPHELES

SUPERPICTUS

_ ANOPHELES

" STEPHENS/

FRILL AT

WIDEST PART

/ OF EGG

PATTERN ON TOP

AT LEAST

1/3 AS LONG

AS EGG

LESS THAN

1/4 AS LONG

AS EGG, OR

ABSENT

ANOPHELES

MACUL/PENNIS

ANOPHELES

SACHAROV/ AND PLU/\4BEUS

A.PLUMBEUS BREEDS ONLY IN TREE \

HOLES AND ARTIFICIAL CONTAINERS)

AND IS NOT AN IMPORTANT VECTOR I

OF MALARIA /

COMB SCALES

IN ONE

REGULAR ROW

(NOT ON A PLATE)

-ABSENT

CjOMB_SCAL_ES

_ AEDES

~ AEGYPTI

ANOPHELES LARVA

PLEURAL BRISTLES

SADDLE. HAIR

cipal vector in the mountain ranges throughout this region

and is the common species in the foothills surrounding Turkey, on the western Anatolian plain, and in the Lebanon and

iVnti-Lebanon Mountains, where malaria is not so severe as in

the lowlands but where nevertheless it is an important

disease.

A, maculipennis is almost always associated with low

spleen and blood rates in the areas where it transmits malaria

and must be present in large numbers in order to be an efficient vector. These characteristics are partly due to its high

degree of zoophilism. The females can be rather easily diverted from feeding on man by using domestic animals as

a barrier. The species is present in nearly all the foothill

areas of the Near East and is often found with superpictus.

Anopheles sergenti is very similar to superpictus in breeding habits and ability to carry malaria, but it is much less

important. In western Jordan snperpiotus is generally a

late-season breeder, but sergenti appears even later, its larvae

having been found as late as November in Jordan.

Anopheles clavigeris principally an urban species. On the

central plain and coastal areas of Syria and Lebanon it is

locally important. Over most of its range it is a wild

mosquito, but in parts of Turkey and in Israel, especially

near Jerusalem, it breeds in water stored in underground

cisterns during the hot weather and freely enters nearby

houses for human blood meals. In other parts of its range

claviger prefers animals for food. When they are not available, it will readily feed on man.

Anopheles stephensi is restricted to a small area in the

southeastern part of Iraq. It is an urban mosquito, breeding

in all kinds of water-holding receptacles close to man's houses

and in small collections of ground water as well. It is an

important vector in the alluvial plain of Iraq, where it

flourishes in the date palm gardens.

References : 61,118, 229,239, 2U, 265.

Anopheles Species in the Near East

algeriensis

apod

"^claviger

GOVjStani constani

cousta,ni tenehrosus

^Gulicifacies

^dHhali

\flumatilis

\hispaniola

hyrcanus

"^maculipennis

marteri mavteri

marteri sogdianus

See footnote 6, p. 10.

\iiiihesseae

\multicolor

\phaToensis

\plumbeus

fpulcherrimus

rhodesiensis rupicolus

"^sacharom

"^sergenti

"^stephensi

"^superpictus

turhhudi telamali

turkhudi turhhudi
70 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPCClMeN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE— THE NEAR EAST

[AREA INCLUDED: TURKEY. SYRIA, LEBANON, ISRAEL, JORDAN, AND IRAQ]

[FEMALES]

[M/í¿£S HAVE BUSHY ANTENNAE

LONG PALPS ON

ANOPHELES

ANTENNAE

DARK, WITH LIGHT SPOT IN FRINGE AT TIP

, ANOPHELES

I _ MACULIPENNIS AND RELATIVES

^NOT ALL IMPORTANT; ADULTS \

\ CANNOT BE SEPARATED /

FOUR DARKER SPOTS

ENTIRELY DARK

_ ANOPHELES

— SACHAROVI

TOP VIEW OF HEAP

TUFT OF

•f WHITE

SCALES ,

END SEGMENT NOT MORE THAN _ ANOPHELES

SHORT PALPS ON

ALL OTHERS PATTERN ON THORAX

_ AEDES

~" AEGYPTI

ADULT MOSQUITO WITH PARTS LABELED

Yellow Fever.—This disease is not present in the Near

East.

Reference: ^56.

Dengue.—This disease is prevalent in Turkey along the

Black Sea and the Mediterranean coasts. An epidemic

affecting a large number of people occurred in 1928 and 1929

as a result of the 1927-28 epidemic in Greece, but there have

been no further outbreaks. A small epidemic occurred in

Hama, Syria, in 1940, but, as in Turkey, only sporadic cases

have been reported since. Small epidemics occurred in

Baghdad in 1921 and 1947, and the disease has been reported

from both Israel and Jordan. In all these places Aedes

aegyptij breeding in all sorts of artificial containers close to

man, has been the vector.

References: 80,34,256.

Encephalitis.—Over 100 cases, mostly in children, of a

nonfatal disease caused by the West Nile encephalitis virus

occurred in Israel in 1951 and again in 1952. The vector

there is not known.

References: 58,251,

Filariasis.—A small focus of infection with Wuchereria

hancrofti has been found in Lebanon. Since filariasis is not

indigenous to the Near East, the cases are believed to have

originated by contact with African troops during World War

II. No vector has been named.

Reference: 10,

Arabia

(Figs. 41 and 42)

Physical Features.—^Arabia includes Saudi Arabia, Yemen, Aden, Oman, Kuwait, and the islands of Bahrein and

Socotra. It is a large peninsula separated from Africa by

the narrow Red Sea and the Gulf of Aden and from Iran by

the Persian Gulf and the Gulf of Oman. Its highest mountains, rising to 9,000-12,000 feet, extend along the Red Sea

coast. In the south they are continued in the so-called Jabal

Akhdar and Kaur Mountains, which at their eastern ends

rise to almost 10,000 feet. The interior plateau slopes gradually eastward to the coast. This plateau contains the northcentral Syrian and An Nafud Deserts, between which are the

vast deserts of Rub' al Khali and Bahr as Safi, which fill

almost the entire southern third of the peninsula behind the

mountains. West-central Arabia contains mountainous highlands, through which deep, eroded valleys extend from east

to west.
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE — ARABIA

[AREA INCLUDED: 5AUDI ARABIA, YEMEN, ADEN, OMAN, KUWAIT, AND BAHREIN AND SOCOTRA ISLANDS]

FULL-GROWN LARVAE

71

FIG. 41

NO SIPHON ON ANOPHELES

LONG PLEURALS

OF 3RD GROUP

ONE

HAIRED <

ABDOMINAL SEGMENTS

4 TO 7

rPROTHORACIC HAIR 1

ANOPHELES

GAMBIAE

ANOPHELES

SERGENT/

AND RUPICOLUS

ONLY SERGENT/

IMPORTANT; LARVAE

CANNOT BE SEPARATED WITH CERTAINTY

_ ANOPHELES

" FLUV/AT/L/S AND ARAB/CUS

/ONLY FLUV/ATIUS IMPORTANT; \

V LARVAE CANNOT BE SEPARATED ^

CAUDAL HAIRS

TRIPLE -^"i — ANOPHELES

\— ADENENS/S

CURVED AT TIPS _^

ANTENNAL HAIR

_ ANOPHELES

"~ SACHAROV/

LONG PLEURALS

OF 2ND GROUP

f

OUTER CLYPEALS

/ i \

BOTH

BARE

« ANOPHELES

"" PHAROENS/S

ONE

HAIRED

INNER FRONTALS

DARK SPOTS

AT BASES \

\

«. ANOPHELES

— /^ULT/COLOR

TOP VIEW OF HEAD OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

CLYPEALS

HIND

CLYPEALS

ANOPHELES LARVA

PROTHORACIC

HAIR 1

PLEURAL BRISTLES

ABDOMINAL

SEGMENTS

4 TO 7

CAUDAL HAIRS

PALMATE HAIR OF SEGMENT 1

NO DARK

SPOTS AT

BASES

FILAMENT ->. v /

LESS THAN ^ k à /

1/2 AS LONG \mmJ /

ANOPHELES

= STEPHENS/

AND THEOBALD/

(ONLY STEPHENS/ IMPOR- \

TANT; LARVAE CANNOT BEI

BE SEPARATED WITH /

CERTAINTY /

SIPHON ON ALL OTHERS SIPHON ANTENNA BODY HAIRS COMB SCALES

AEDES

AEGYPTI
72 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - ARABIA

[AREA INCLUDED: SAUDI ARABIA,YEMEN, ADEN, OMAN, KUWAIT, AND BAHREIN AND SOCOTRA ISLAMDsJ

^^Ä FEMALES

\_MALES HAVE BUSHY ANTENNAE ( Újli^F ) AND DO NOT BITE^ FIG. 42

LONG PALPS ON

ANOPHELES

ANTENNAE

PATTERN ON HIND TARSUS

SHORT PALPS ON

ALL OTHERS

PATTERN ON

THORAX

_ AEDES

— AEGYPTI

PATTERN ON PALP

^LIGHT AT TIP

^DARK AT TIP

ONE OR TWO SHORT DARK SPOTS BEHIND

MARGINAL SPOT -^

miiiiiiiiiKteïï ÍPliiíi

( WING OF GAMBIAE )

ONE LONG DARK SPOT BEHIND

MARGINAL SPOT

1^

ANOPHELES

GAMBIAE

mmk ANOPHELES

PHAROENSIS

NO LIGHT SPOT AT CROSS VEIN, BUT

LIGHT SPOT IN BASAL THIRD WITH

DARK SPOT BEHIND IT

ANOPHELES

STEPHENS!

_ ANOPHELES

— ADENENS/S

0£NCNS/S )

NO LIGHT SPOT AT CROSS VEIN, BUT

LIGHT SPOT IN BASAL THIRD WITHOUT

DARK SPOT BEHIND IT

BASAL THIRD ENTIRELY DARK

ENTIRELY DARK

iiiiiiPii^

Í WING OF MULTICOLOR^

ENTIRELY DARK, WITH FOUR —-I

DARKER SPOTS j

ANOPHELES

SERGENT/

ANOPHELES

FLUVIATILIS

AND ARAB/CUS

ONLY FLUVIATILIS \

IMPORTANT;ADULTS\

CANNOT BE SEPARATED WITH /

CERTAINTY /

ANOPHELES

MULTICOLOR

ANOPHELES

SACHAROVI
MOSQUITOES OF MEDICAL IMPORTANCE 73

Most of the interior has desert temperatures and a rainfall

of about 4 inches annually. However, the mountainous highlands along the southern frontier where the principal agricultural areas are located, receive from 20 to 40 inches of

rain per year in well-marked monsoon seasons. These areas

contain approximately four-fifths of the country's population.

Reference: SOI.

Malaria.—It has been said that wherever there is man in

Arabia there is water, and wherever there is water there is

malaria. The most abundant mosquito in the Persian Gulf

area is Anopheles stephensi. This species is essentially urban

and is important throughout most of its range. It is an inland vector from Al Karj on the west to the coast on the east.

Itis also a vector in the Bahreins, where a hot, humid climate

prevails and date plantations are watered by abundant warm,

slightly brackish water. Malaria here accounts for almost

one-fourth of the hospital admissions. It is almost equally

divided among Plasmodium falcipar%im^ P. malariae^ and

P. vivax, Muscat to the south, where stephensi is also a

vector, is malarious, with high spleen and parasite rates in

the human population the year round.

Anopheles ßuviatilis^ primarily a foothill species in the

Orient, is the second most abundant mosquito along the

Persian Gulf coast. Its actual importance in malaria transmission in Arabia has not been completely determined. Many

of its previous records have been confused with those of

stephensi. It has been included in the keys because of its importance in the Orient and because it is believed to transmit

some malaria in Arabia.

Neither the distribution nor importance of Anopheles

rrmlticolor \s> known with certainty. This species is essentially an oasis mosquito and breeds in desert water of extremely

high salinity. Itis potentially able to carry malaria efficiently and is regarded as a vector throughout the deserts of

Egypt and Northwest Africa.

Anopheles sergent^ another oasis breeder, has been recorded from various dry parts of Arabia. It is the only

vector recorded from a well at Jabrin oasis, which was said

to be abandoned at one time as an agricultural colony because

of malaria. This species is common in the outskirts of

Jeddah. Both sergenti and rmilticolor probably transmit a

large amount of malaria in the interior deserts of Arabia.

Anopheles adenensis^ originally described as a variety of

the important Indian vector A, culicifaciesy has been observed m large numbers in and about Aden and at Hodeida

on the Eed Sea coast of Yemen. It is definitely a vector at

Hodeida and is probably important in other parts of this

extremely arid coastland as well.

Anopheles gambiae^ by far the most important malaria

vector in continental Africa, has invaded various localities

on the Eed Sea coast and has migrated inland for some distance. Doubtlessly this species transmits malaria in the

Jeddah-Mecca area. In a severe epidemic at Jeddah in 1950

it was found breeding prolificacy in waste-water pools

formed by a new city water-supply system. " This species

was said to have been carried to Jeddah by extensive pilgrim

traffic.

Ainopheles pharoensis is included in the keys because of

its importance in Egypt and its discovery in Yemen. Its role

in Arabia is not known. This species is of potential danger

when it appears in large numbers in the presence of human

malaria.

Anopheles sacharovi is the only member of the maculipennis complex that might be found in Arabia. It has

never been recorded there, but its importance in other areas

of its range necessitates its inclusion in the keys.

Anopheles dHhali has been suspected as a vector in Arabia,

butthe few available data are not conclusive.

References : 67, 206, 3^7.

Anopheles Species in Arabia

^adenensis "^multicolor

apoci jpharoensis

arahicus {plumbeus

einereus \pretoriensis

\G0ustani coiùstani {pulcherrimus

coustani tenebrosus rhodesiensis rupicolus

•dHhali \sacharovi

'^iiuviatilis "^sergenti

'^gcmibiae ^stephensi

\hispcmiola \suipictus

hyrcamis \superpictu8

See footnote 6, p. 10.

Yellow Fever.—This disease is not present in Arabia.

Reference: 256.

Dengue.—Dengue is present and lightly endemic in the

Yemen and Muscat areas of Arabia, where it is known as

oasis fever, a rather misleading term, since malaria and sandfly fever are also known by that name in various parts of the

country. Dengue epidemics have not been reported, but the

vector Aedes aegypti breeds in all kinds of artificial containers close to man's habitations.

References : i, 256.

Encephalitis.—No information is available.

Filariasis.—No information is available.

Iran

(Figs. 43 and 44)

Physical Features.—Iran lies along the southern border

of the Asiatic mainland and forms a zoogeographical link between the Near East and the Orient. The country consists

of a vast central plateau, with an average elevation of about

4,000 feet. This plateau is separated from the Caspian Sea

by the Elburz mountain range, with peaks as high as 18,000

feet. Its western slopes dip to the Caspian, leaving a narrow

coastal lowland. A wide series of mountain ranges comprise

the western third of Iran. They are bounded on their western side by the broad fertile valley of the Tigris and Euphrates Eivers, which extends into western Iran. To the

south a narrow coastal lowland borders the length of the

Persian Gulf and the Gulf of Oman, part of the Arabian

Sea. On the east more mountain ranges separate Iran from

Afghanistan and West Pakistan. The salt desert of Dashti-Kavir is situated in the north-central and northeast area,

and the Dasht-i-Lut, connected to it by a narrow arm, is a

southward extension of this salt desert.

Eather low rainfall and high winds prevail over much of

the country. The hot, dry summer monsoon from northwest

India, alternating with winds blowing south from Siberia,

causes extreme temperatures. Areas of highest rainfall,

about 40 inches per year, are along the subtropical Caspian

coast and in the northwest part, which contains Lake Urmia,

the country's largest lake, whereas the interior deserts receive

less than 5 inches of rain per year. The Persian Gulf coast

has an extremely dry, hot summer.

Reference: SOI.

Malaria.—Highly endemic malaria exists in the low semitropical areas of the Caspian coast, the Persian Gulf coast,

and the lower Tigris and Euphrates areas. Malaria is also

endemic, but localized, in the widely scattered marshes of the

central plain and in even drier areas, and it is mildly endemic

in some localities on the highest northern plateaus. In such

barren terrain as this country has, both man and mosquitoes

are necessarily thrown together in the watered areas. Such
74 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - IRAN

^ EGGS AND FULL-GROWN LARVAE ^

NO SIPHON ON ANOPHELES

LONG PLEURALS

OF 2ND GROUP

LONG PLEURALS

OF 3RD GROUP

BOTH

OUTER CLYPEALS ANTENNAL HAIR

FIG. 43

ANOPHELES

MACUUPENNIS

AND RELATIVES

(LARVAE CANNOT BE \

SEPARATED- SEE

KEY TO EGGS BELOW/

ANOPHELES

CULICIFACIES

ONE / J^

HAIRED / ¿^ ,

ANOPHELES

SUPERPICTUS

__ ANOPHELES

"" STEPHENSI

TOP VIEW OF HEAD OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS

OUTER I HIND

CLYPEALS i CLYPEALS

ANOPHELES LARVA

PROTHORACIC

PLEURAL BRISTLES

SIPHON ON ALL OTHERS

INTERRUPTED

BV FLOATS'

AT MIDDLE

OF EGG

FRILL IN

WIDEST PART

'' OF EGG

FLOATS

AT LEAST

1/3 AS LONG

AS EGG

LESS THAN

IA AS

LONG AS

EGG, OR

ABSENT

PATTERN ON TOP

«. ANOPHELES

- MACUUPENNIS

ANOPHELES

SACHAROVI AND PLUMBEUS

APLUMBEUS BREEDS ONLY IN ^

TREE HOLES AND ARTIFICIAL

CONTAINERS AND IS NOT AN

IMPORTANT VECTOR OF MALARIA,

COMB SCALES

AEDES

AEGYPTI

_;, CAUDAL HAIRS

enforced coexistence presents ideal conditions for the maintenance and spread of malaria.

No information is available on the Plasnwdium species in

Iran.

Anopheles superpictus and A, sacharovi are the two most

prominent vectors in Iran. The former species is widely

distributed but appears neither in the deserts nor in the

southeast corner of the country. It is responsible for transmission of most of the malaria at elevations above 1,000 feet

in all except the humid Caspian coastal area, where it does

not enter houses and is considered to be a wild species. In

drier areas the adults are always found in houses and the

larvae breed in almost all available water collections. It is

the most important vector in the intensely malarious westcentral area.

A. sacharovi^ on the other hand, is the principal vector in

the most intensely malarious area of Iran, i. e., the Caspian

coast. It is the predominant mosquito on the coastal plain,

and its numbers as well as malaria incidence in this area increase markedly from July to November. This species inhabits ricefields and water used for irrigation and to a lesser

extent the rivers, which tend to become slow and sluggish

during the summer. Although sacharom is found principally in the western half of Iran, its numbers are not so concentrated there as in coastal localities.

Anopheles maculipennis closely parallels sacharovi in distribution. It is concentrated in the Caspian area, and its

range extends from the northwest part of Iran as far south

as Abadan at the head of the Persian Gulf. It is apparently

absent from the gulf coast and most of the interior, but it

has been observed near the northeast border. A. maculipennis

and superpictus often appear together, but the former is

more characteristic of the lower valleys and tablelands. Its

exact role in the transmission of malaria in Iran is not known,

though it is a proved vector in some other parts of its range

and is suspected as a vector of a mildly endemic malaria in

northern Iran.

Anopheles stephensi has a more southern distribution than

any of the other vectors. It is absent from the entire northern half of Iran but is responsible for the transmission of a

great deal of malaria along the Persian Gulf coast and eastward. It is of great importance at Abadan, where it rests

commonly in manmade shelters. It is characteristically an

urban mosquito but also breeds rurally in various collections

of water.

Anopheles culicifacies^ like stephensi, is an extremely important vector of malaria in India and is largely restricted

to areas east and south of the deserts. It has been found as

far west as Bushire on the Persian Gulf coast. It breeds m

cisterns, wells, and other large manmade containers and is
MOSQUITOES OF MEDICAL IMPORTANCEi 75

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - IRAN

M FEMALES ^

[ MALES HAVE BUSHY ANTENNAE i

LONG PALPS ON

ANOPHELES

ANTENNAE

-PROBOSCIS

SHORT PALPS ON

ALL OTHERS

/NO LIGHT SPOT AT CROSS VEIN FOUR DARK SPOTS "

WING ENTIRELY DARK, OR LIGHT -

ONLY AT TIP

) AND DO NOT BITE

FRINGE AT WING TIP

^ '

FIG. 44

^"A'LLDARK = ANOPHELES

^-LIGHT SPOT AT CROSS VEIN

.-VpSTir = ANOPHELES

FRINGE MACULIPENNIS AND RELATIVES

/ ADULTS CANNOT BE SEPARATED - \

\SEE KEY TO EGGS /

C PATTERN ON HIND LEG PATTERN ON HIND TARSUS

/V

_ ANOPHELES

"" CULie¡FACIES

PROPLEURAL SETAE

+ ABSENT

SEE ENLARGED DRAWINGS

FOR LOCATION /

ANOPHELES

STEPHENSI

ANOPHELES

SUPERPICTUS

PATTERN ON THORAX

AEDES

AEGYPTI

^<VNW>M1> -^-5=^^

PROPLEURAL CT^\ '^ (^ / h7^ SETAE

\ fc^

)h

ri y

\J^

-'' W'r

ADULT MOSQUITO WJTH PARTS LABELED

THORAX^

probably an important vector, although its exact status in

malaria transmission in Iran needs more study.

References : 67, lOG, 101, lUAnopheles Species in Iran

algeriensis moghulensis

apoci ^multicolor

cinereus \plu(mbeus

\claviger \pulcherrimus

*cuUcifades \rhodesiensis

\dHhali "^sacharovi

\-ßuviatilis \sergenti

hyrcanus "^stephensi

"^maculipennis \subpictus

marteri marteri "^superpictus

marteri sogdianus turhhudi

]melanoon suhalpinus

See footnote 6, p. 10.

Yellow Fever.—Yellow fever does not occur in Iran. The

classical vector Aedes aegypti is present and common in most

port cities, where there is a constant threat of this disease

being introduced.

References : 18, 256.

Dengue.—Dengue occurs in endemic form, with occasional

outbreaks along most of the Persian Gulf coast and in the

Caspian Sea area. In these two areas it is transmitted by

Aedes aegypti. Aedes aïbopictus is not present in Iran.

Reference : 18.

Encephalitis.—No information is available.

Filariasis.—No information is available.

Afghanistan and West Pakistan

(Figs. 45 and 46)

Physical Features.—Afghanistan is bordered on the west

by Iran, on the south and east by West Pakistan, and on the

north by the U. S. S. R. The Hindu Kush, a series of mountain ranges rising in the north to 25,000 feet, extends southward from the U. S. S. R. and covers almost all the northern

half of the country. A belt of steppes separates these mountains from arid lowlands on the western frontier and from

extensive desert areas in the southwest. A depression surrounding Jalalabad near the famed Khyber Pass, the gateway from Afghanistan through the eastern mountains to

West Pakistan, has a subtropical climate. Rainfall ranges
76 AGRICULTURE HANDBOOK 152. U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE — AFGHANISTAN AND WEST PAKISTAN

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

; FULL-GROWN LARVAE

NO SIPHON ON ANOPHELES

CAUDAL

HAIRS'=^::;~~^

LONG PLEURALS

OF 3RD GROUP

LONG PLEURALS

OF 2ND GROUP

OUTER CLY PEALS

BARE OR

FINELY

HAIRED

" ABDOMINAL PALMATES

FILAMENT

MORE THAN

1/2 AS LONG

AS BLADE

ABDOMINAL PALMATES

SEGMENT

FIG. 45'

ANOPHELES

SUPERPICTUS

FILAMENT

LESS THAN

1/2 AS LONG

AS BLADE y

_ ANOPHELES

— STEPHENS! AND MACULATUS

_ ANOPHELES

— CLAV IGER

ABDOMINAL PLATES CAUDAL HAIRS

ABOUT 1/3 AS

AS WIDE

AS SEGMENT

NEARLY

AS WIDE

AS SEGMENT

-êSUD ANOPHELES

CULICIFACIES

CURVED AT TIPS ''

_ ANOPHELES

FLUVIA7ILIS

FLOATS-«A^^^ FLOATS -->^

SMOOTH ^S^ SMOOTH

WHEN

PRESENT

MACUUPENNIS

OUTER CLYPEALS ANTENNAL HAIR

ANOPHELES

SACHAROVI, MACULIPENNIS,

— AND MELANOON SUBALPINOS

lUCLANOON SUBALPINOS NOT IMPORTANT; LARVAE CANNOT \

\ BE SEPARATED; SEE EGGS ABOVE I

COMB SCALES Í

__ AEDES

~" AEGYPTI

__ AEDES

— ALBOPICTUS

ANOPHELES LARVA

CLYPEALS

OUTER

CLYPEALS

y^ PLEURAL BRISTLES

4

^,,

^f%^ ABDOMINAL

SEGMENTS

2 i3

ABDOMINAL

PLATES

li^T

W

WOMPLN CAUDAL

/mñ^wC^T^ HAIRS

annually from 10 inches in the steppes to less than 2 inches in

the southwestern desert areas.

West Pakistan is separated from the U. S. S. E.. by the

Himalayas in the north and has a southern coast on the

Arabian Sea. The Indus Eiver, flowing the entire length of

the country, divides the Thar, or Indian, Desert in the east

from mountain ranges in the west. Semiarid and desert

areas west and south of these mountains are contiguous with

those in the southern parts of Afghanistan. Extremes of

temperature, from 40° in the winter to 120° F. in the summer,

prevail in this country. The rainfall ranges annually from

5 inches or less in the southwest to about 25 inches in the

northeast. The northern tributaries of the Indus River are

perennial, but those of the southwest are seasonal.

Reference: 201.

Malaria.—Malaria is endemic in most of Afghanistan. It

is not an important disease in the arid southwestern areas

of Afghanistan or in western Pakistan. In the steppes area

traversing the center of Afghanistan malaria is more highly

endemic, and in the northern mountainous areas of both

countries the disease is prevalent even at levels above 5,000

feet. Most transmission occurs in September, when mosquito populations are attheir peak.

Little is known of the distribution of the Plasmodium

parasites in either country.

Anopheles superpictu^ is the important vector in the

mountainous areas and has only one generation each year.

The transmission period begins in July and reaches a peak

in September. At lower altitudes the breeding of this

highly anthropophilic mosquito starts in May, when a low

degree of transmission begins.

Anopheles sacharovi is another important vector in the

mountainous northern half of Afghanistan and in parts of

West Pakistan. It commonly appears in the same areas as

superpictus^ but it is characteristically a mosquito of valleys

and mountain plains rather than of higher mountain areas.

For this reason it is a more important vector at lower altitudes than superpictus^ especially in those eastern parts of

Afghanistan having a moderate to subtropical climate. In

most of its range it has but one generation each year.

Anopheles culicifacies^ one of India's most important vectors, appears in northeastern West Pakistan and eastern

Afghanistan. This species has been found with low natural

infection rates and maintains an endemicity that increases in

spring and early summer, almost disappears during the

hotter months, then reappears in varying intensity in the

autumn.
MOSQUITOES OF MEDICAL IMPORTANCE 77

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE — AFGHANISTAN AND WEST PAKISTAN

-^^¡^ FEMALES ^

[^MAL£S HAi^E BUSHY ANTENNAE {\uM^^) ^ND DO NOT BITE'\

PATTERN ON PALP

FIG. 46

LONG PALPS ON

ANOPHELES

-PROBOSCIS

PATTERN ON WING

AT LEAST FOUR LIGHT SPOTS ON FRONT MARGIN

lu'"rnniii'i

_ ANOPHELES

CLAVIGER

PATTERN ON HIND TARSUS

PATTERN ON HIND TARSUS

ENTIRELY DARK, OR LIGHT ONLY AT TIP

SHORT PALPS ON

ALL OTHERS

ONE LIGHT SPOT IN BASE OF FRONT MARGIN,

WITH DARK SPOT BEHIND

-_ ANOPHELES

~ CUL ICIFACIES

TWO LIGHT SPOTS IN BASE OF FRONT MARGIN:

NO DARK SPOTS BEHIND

'iiijIîpgijŒ

flliiiiiiiiiilfcSi«ii?iii;iSii!!i!H^ -

_ ANOPHELES

%• ~ SUPERPICTUS

ANOPHELES

FLUVIA TILIS

VEIN 3 WITH LIGHT SPOT

ANOPHELES

STEPHENSI

YELLOWISH SPOT

IN FRINGE

AT TIP

ANOPHELES

MACULIPENNIS

-- AND MELANOON SUBA L PINUS

(ONLY MACULIPENNIS

IMPORTANT; ADULTS CANNOT BE

SEPARATED

FRINGE ENTIRELY __ ANOPHELES

DARK AT TIP — SACHAROVI

PATTERN ON THORAX

A EDES

A EGYPTI

_ AEDES

~ AL BOPICTUS

ADULT MOSQUITO WITH PARTS LABELED

Anopheles -ßuviatilis^ a mosquito of West Pakistan, transmits endemic malaria in a manner similar to Anopheles

minimus in the Orient (see p. 83). Its occurrence in Afg'hanistan is not known with certainty.

Anopheles stephensi is the only medically important urban

anopheline in both countries. Because the larvae live in all

types of manmade water collections, the malaria transmitted

by it is often localized and limited to urban centers. This

species is extremely abundant, but it has not been found infected in nature in certain areas of Afghanistan.

Anopheles claviger also appears in the extrenie northern

mountains of Afghanistan, where it is believed to maintain

malaria in the vicinity of mountain streams.

The roles of Anopheles maculipennis and A, pulcherrimus

in the transmission of malaria in Afghanistan and West

Pakistan are not well known. In certain parts of their respective ranges they are important vectors or suspected of

being important vectors.

References : 67,101, 235, 280.

Anopheles Species in Afghanistan and West Pakistan

\annularis

"^claviger

"^Gulicifaoies

\dHhali

^-ßuviatilis

hdbihi

hyrcanus \pulcherrimus

lindesayi "^sacharovi

\maculatus maculatus \sergenti

maculatu^ willmori splendidus

\maGulipennis "^stephensi

moghulensis jsubpictus

\multicolor ^ superpictus

\nigerrimus turkhudi

pallidus vagus

See footnote 6, p. 10.

Yellow Fever.—This disease is not present in Afghanistan

and West Pakistan.

Reference: 256.

Dengue.—Dengue occurs at the head and mouth of the

Indus Eiver in several endemic areas, where both Aedes

aegypti and A, aTbopictus are present. The distribution of

aegypti extends farther east than that of albopictus^ so that

the former species is more likely to be a vector in West

Pakistan. The disease has not been reported from Afghanistan, and neither vector has been recorded from there.

References: 256,810.

Encephalitis.—No information is available.

Filariasis.—No information is available.
78 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE — INDIA AND CEYLON

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

[ FULL-GROWN LARVAEJ

FIG. 47

NO SIPHON ON ANOPHELES

SPIRACULAR

APPARATUS^

LONG PLEURALS

OF 3RD GROUP

LONG PLEURALS

OF 2ND GROUP

OUTER CLYPEALS INNER SUTURAL HAIRS ABDOMINAL PALMATES OF SEGMENT 1

NOT /OR

_ ANOPHELES

— ANNULARtS

ABDOMINAL PALMATES OF SEGMENT 4

ANOPHELES

'fZ STEPHENS/ STEPHENS/

AND STEPHENS/ MKSO/?ENS/S

HIND CLYPEALS

PLACED FAR

I BACK OF

~ INNER

CLYPEALS

ABDOMINAL SEGMENTS INNER CLYPEALS

4 TO 7

ABDOMINAL PLATES

HAIR "O" ON PLATE - ^

DOUBLE ^m ^ \

[m^"-^ -*)

TRIPLE T--':L*ia

3 »} ^ yANOPHELES

SUNDA/CUS AND SUBP/CTUS

ANOPHELES

VARUNA

HEAD OF ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS

/ OUTER I HIND

CLYPEALS 1 CLYPEALS

^\ _ ANOPHELES

^ ~ M/N/MUS AND FLUV/AT/L/S

/BOTH IMPORTANT} LARVAE CANNOT \

\ BE SEPARATED ]

_ ANOPHELES

\ — CUL/C/FAC/ES

CURVED AT TIPS

SIPHON ON ALL OTHERS BODY HAIRS

'M—ABSENT

COMB SCALES

AEDES

AEGYPT/

PECTÉN SPINES

_ AEDES

~~ ALBOP/CTUS

ANOPHELES LARVA

PROTHORACIC

PLEURAL BRISTLES

CAUDAL HAIRS

India and Ceylon

(Figs. 47 and 48)

Physical Features.—India, isolated on the north from the

rest of Asia by the highest mountain range in the world, the

Himalayas, consists of three rather well-defined areas: (1)

Hilly and mountainous country descending southward from

the Himalayas; (2) an extensive agricultural area formed by

the valley of the Ganges Eiver and known as the United

Provinces ; and (3) the extensive plateau of peninsular India,

with foothills and escarpments along the narrow western

coast. The main rivers of peninsular India form broad deltas on the wide east coast. In the northwest corner is the

extensive Thar Desert, which merges with swampy areas of

the Kathiawar peninsula on the west coast. Daily annual

variations in temperature are more marked in the northern

and centralthan in the southern parts of the country. Southwest monsoons from June to September bring most of southwestern India's yearly 100 to 200 inches of rain. The ejffect

of these winds decreases to the north and east, where periods

of rain alternate with drought. A cool, dry period prevails

from October to February and a hot, dry season with scorching winds from March to May.

Ceylon consists of a central mountain mass, 8,000 feet high,

which is surrounded by a broad coastal plain with sandbars

enclosing lagoons and marshes. The island has a small

annual range of temperature, with heavy rains brought to

the western mountain slopes by a southwest monsoon from

June to September and to the eastern slopes by a northeast

monsoon in October and November.

Reference: SOI.

Malaria.—This disease occurs practically everywhere in

India and Ceylon except in the Himalayas. The vast plains

and plateaus of the Ganges in the United Provinces and the

central and peninsular parts of India are subject to a marked

seasonal type, with a low or moderate endemicity. There

are usually two peaks of prevalence, one in the spring and

early summer and another in the autumn. Between these

peaks the disease is almost completely absent from many parts

of the area. Localized epidemics of high intensity occur

from time to time in the northwest. They are associated

with high rainfall and river ñooding.

In the northern parts of India Plasmodium vi'vax is by far

the most common parasite, whereas P, falciparum predominates in the south. Plasmodium malariae is common in

southern India, and in wet areas in Ceylon it accounts for

as many cases as P. falciparum.
MOSQUITOES OF MEDICAL IMPORTANCE 79

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - INDIA AND CEYLON

FEMALES

\MALES HAVE BUSHY ANTENNAE [ Hi^i^^^ \AND DO NOT B/TE^ FIG. 48

LONG PALPS ON

ANOPHELES

ANTENNAE

-PROBOSCIS

PATTERN ON HIND FEMUR f PATTERN ON PALP

AND TIBIA

PATTERN ON HIND TARSUS VEIN 3 WITH LONG CONTINUOUS DARK AREA

5;?: __ ANOPHELES

- CULIdFACIES

ANOPHELES

FLUYIAT/L/S

_ ANOPHELES

" VARUNA

ANOPHELES

Zl STEPHENS/ STEPHENS/

AND STEPHENS/ /^ySORENS/S

SHORT PALPS ON

ALL OTHERS

PATTERN ON THORAX

AEDES

AEGYPT/

AEDES

ALBOPICTUS

ADULT MOSQUITO WITH PARTS LABELED

THORAX^^

The most important and widespread vector of malaria in

central and. peninsular India is Anof heles culicifocies. This

species is also the most important, if n,ot the only vector in

Ceylon, but there it flourishes in periods of drought, breeding

in small accumulations of water held over from river drying.

The foothill areas bordering the central plateau on both

the east and west sides of pen^lnsular India have a hyperendemic malaria supported by Anopheles fluviatilis. Here

there are usually two periods of prevalence, one preceding

and one following the monsoon rains. The breeding places

of this mosquito may be completely washed out during the wet

season, but transmission usually occurs during most or all the

year.

The clisease in the Himalayan foothills on the northeastern

rim of ïndia bordering Nepal and China has a similar periodicity. Its chief vector is Anopheles minimus. In contrast

to conditions in southern India malaria in the north is absent

during most of the year. Anopheles vanma^ a close relative

of minimus and ßuviatilis^ has been implicated as a vector

of only occasional importance in east-central India, since it

has been found naturally infected in widely scattered localities.

Anopheles stephensi stephensi is responsible for a considerable amount of malaria in urban areas, especially along the

northern west coast and in the heavily populated Ganges

471779 O—59 6

plain. The real importance of this species lies in its ability

to breed prolifically in artificial containers close to man. This

urban malaria is extremely localized, because cities differ in

their ability to cope with their malaria-control problems.

Anopheles stephensi mysorensis^ usually considered to be

of less importance than the type form, has been implicated as

the chief rural carrier at Vizagapatnam, where rather high

sporozoite rates have been encountered. Possibly some of

the infection records given for the type form in the literature

actually apply to mysorensis.

Anopheles sundaicus^ typically a brackish-water breeder,

appears from Calcutta southward to the middle of peninsular

India along the east coast. It is responsible for periodic

outbreaks of malaria, some of which may become rather

severe in many localities. This malaria appears either in the

spring or in the autumn, depending on whether local habitats

are favorable for breeding. A, sundaicus is typically associated with cleared areas that have been embanked to protect

them from the sea.

Anopheles annularis is not thought to be an important

mosquito in India, but it sometimes is observed in extremely

large numbers, among which infected specimens have been

found. It is believed to play some part in the transmission

of the malaria in Orissa.

References : 67,101, 310.
80 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Anopheles Species in India and Ceylon

inaculatu^ wilhnori

majidi

moghvlensis

\rmdtioolor

jnigerrinms

pallidus

pinjœwrem^is

\p%dGherrmms

ramsayi

sintoni

splendidus

"^stephensi mysorensis

"^stephensi stephensi

\sy/bpictus

\superpictns

\tessellatus

theobaldi

turkhudi

vagus

^varuïïia

\aconitus

aitkenii

annandalei

*ann(ularis

\harbirostris

harianensis

'^cuilicifactes

euliciformis

\dHhali

elegans

^flumatilis

giga^ gigas

gigas refutans

gigas simiensis

insulaefloru'tn

jamesii

\jeyporiensis candidiensis

jeyporiensis jeyporiensis

kamoari

leucosphyrus

lindesayi lindesayi

lindesayi nUgiricus

jîïhactdatus maculatu^

See footnote 6, p. 10.

Yellow Fever.—This disease is not present in India and

Ceylon.

Reference: 256.

Dengue.—This disease is endemic throughout India. It is

especially common after the rainy season in Bengal and

Madras, but it is less so in western India and is practically

unknown in the mountainous provinces along the northwestern frontier. Itis carried by Aedes aegypti and A, aïbopictus ; the former mosquito lives closer to man and is more

universally distributed than the latter.

Beferenees: 266,310.

Encephalitis.—In several localities in West Pakistan and

northwestern and central India, the following viruses have

been found: West Nile, St. Louis, Murray Valley, Ilheus,

Zika, Ntaya, and Uganda S. Some of these are very closely

related antigenicly, and interpretations of the findings are

still being made. The presence of dengue may contribute

toward misidentification of some of them. Japanese virus

is probably present, but it has never been isolated. A study

of the vectors of these viruses is still to be made.

Reference: 257.

Filariasis.—The important centers of filariasis in India

are (1) the lower parts of the Ganges River basin, (2) some

seacoast areas in Bihar and Orissa on the northeast coast,

and (3) Travancore State in the south. Culex pipiens quinquéfaseiatus is not present in the Punjab or is filariasis a medical problem. Farther southeast in the Central Provinces

filariasis with elephantiasis is present in some restricted localities, but nearby localities are practically free of the

disease.

There is only one parasite, Wuchereria malayi^ which is

transmitted by Mansonia annulifera and M, uniformis, M,

annulifera is the chief vector because of its abundance and

high rate of natural infection. Still farther southeast in

Bihar and Orissa Wuehereria hancrofti also appears. This

parasite is transmitted exclusively by quinqusfasciatus^ which

is uniformly present in areas under rice cultivation. Investigations have shown up to 14 percent of these mosquitoes to be

infected in such areas. In Bihar and on the seacoasts of

Orissa malayi is the only parasite and annulifera its chief

vector. Observations at Puri on the east coast indicate filariasis to be an important medical problem. Up to 27 percent of

the human population harbors microfilariae and up to 34

percent of captured mosquitoes are infected. In two coastal

areas of Travancore the principal parasite is malayi and its

principal vector annulifera. Farther south in this State

both parasites are found ; hancrofti is transmitted exclusively

by quinquefasciatus^ which is principally urban, and malayi

by annulifera and uniformis.

References : 192-194, 212-^15, 290, 317, 318, 320.

East Pakistan and Assam

(Figs. 49 and 50)

Physical Features.—East Pakistan, formerly the eastern

part of Bengal, is a flat, low country, most of which rises

hardly more than 300 feet above sea level ; in the southeast

the Chittagong Hills are over 3,000 feet high. East Pakistan

lies largely in the immense fertile Ganges-Brahmaputra

Delta, which consists of a confused network of waterways

cutting across a vast alluvial tract. Eice comprises 85 to 90

percent of the country's agricultural output. Constant flooding of this low land is aided by an annual rainfall of 80 to

100 inches, principally caused by monsoons. The average

temperature ranges annually from about 67° in January to

83° F. in May.

Assam is separated by high mountains from East Pakistan,

the U. S. S. R., and Burma. In the northern half of the

country is the broad agricultural valley of the lower half of

the Brahmaputra; the southern part is largely hilly and

has an east-west mountain range and the Surma River with

its valley. Rainfall varies in different parts of the country :

75 to 85 inches fall annually in the central plateau area, 110

to 120 inches in the Brahmaputra Valley, 125 to 150 inches

in the Surma Valley, and over 400 inches at one locality in

the Khasi Hills. In the last-named area a rainfall of 900

inches, one of the world's heaviest annual precipitations, was

once reported. Although the annual temperature range is

rather great, much of the area is covered by dense tropical

forests.

Reference: 301.

Malaria.—^Although Plasmodium falciparum accounts for

a large number of cases of malaria each year in East Pakistan and Assam, the exact distribution of falciparmn^ P,

vivax^ and P, malarias is not known with certainty.

Malaria is a most important problem in the mountainous

parts of Assam. It occurs there in two periods, as it does

in India, one before the rainy season and one afterward,

closely corresponding to the peaks of mosquito incidence.

During several months of the winter there is a malaria-free

period in the foothill areas. Anopheles minimus breeds in

clear, slowly running water and is the vector. Because rains

flood out most of its breeding places, it is essentially a dryseason mosquito.

Anopheles jeyporiensis candidiensis is a vector along the

East Pakistan-Burma border, and it has been found with

natural infections in the vicinity of Chunati, in several localities near which it was the only species present. Itis regarded

as the vector of most of the premonsoon malaria in the hillocks of this border area.

Anopheles ^umdaicus breeds along the entire coastal area

of East Pakistan in localities where the forest has been

cleared and the land banked to prevent flooding by the ocean.

It does not appear in mangrove areas in that country. It

causes severe outbreaks in certain localities when it is

abundant.

Anopheles philippinensis has been found with sporozoite

rates as high as 2 percent in the western part of the Ganges

Delta, where rivers are sluggish. Premonsoon malaria becomes evident as early as May and June, but the highly

malarious months in this area are September, October, and

November, just at the end of the rainy season when tanks

and impoundments in the area offer optimum breeding con-
MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE - EAST PAKISTAN AND ASSAM

81

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

t FULL-GROWN LARVAE 1

FIG. 49

r INNER SUTURALS

r LONG PLEURALS

OF 3RD GROUP

1

NO SIPHON ON ANOPHELES

POSTSPIRACULAR \-

SIPHON ON ALL OTHERS

ABDOMINAL PALMATES OF SEGMENT 4

FILAMENTS

I 1/3 OR LESS

"r AS LONG AS

BLADE

_ ANOPHELES

— PHILIPPINENSIS

ABDOMINAL PALMATES OF SEGMENT 1

OUTER CLYPEALS

1/3 OR MORE

-|- AS LONG AS

INNER CLYPEALS I

)à

ANOPHELES

ANNULARI S

_ ANOPHELES

— SUNDAICUS AND SUBRICTUS

ANOPHELES

— JEYPORIENSIS JEYPORIENSIS

AND JEYPORIENSIS CANDIDIENSIS

ABDOMINAL PLATES

HAIR VNOT ON PLATE,

\, ANOPHELES

I •=. MINIMUS AHO FLUVIÁTIL IS

.1 /'ONLY MINIMUS IMPORTANT; N

k LARVAE CANNOT BE SEPARATED I

ABDOMINAL PALMATES OF SEGMENT 2 PROTHORACIC HAIRS 1 AND 2

TUBERCLES

LARGE AND

FUSED\

COMB SCALES

ANOPHELES

LEUCOSPHYRUS

_ AEDES

~ AEGYPTI

PECTÉN SPINES

AEDES

ALBOPICTUS

HEAD OF ANOPHELES LARVA

^NTENNAL

HAIR INNER

/ CLYPEALS

PROTHORACIC HAIRS I, 2,* 3'

ANOPHELES LARVA

PLEURAL BRISTLES

4

ditions. Throughout the dry season philifpinensis breeds

primarily in marshes and in rivers cut off from the main flow

by a lowering of the water level.

Although extremely important in central India, Anophèles

culicifacies is believed to be of comparatively little importance in East Pakistan or Assam. Since this species is common and dangerous throughout the United Provinces, it

probably is of some importance in the lower reaches of the

Ganges. The exact nature of the role played by culicifctoies

in this area is not known, but natural infections have always

been comparatively low.

Anopheles annularis^ apparently the only vector in the

Goalpara District of Assam, is believed to transmit malaria

at a low rate in this and other localities where it appears in

large numbers. It is always found with very low natural

infection rates and must be present in large numbers to be

an effective transmitter.

Anopheles leucosphyrus balabacensis^ primarily a forest

species, has been found with high infection rates several times

of the year near Digboi, Assam. It transmits malaria in

much of northern Burma during the rainy season and recently has been implicated as a possible vector along the East

Pakistan-Burma frontier. Observations to determine the

extent of its importance in this area are not yet complete. .

References: 67, 91, 101, 122, 2^2, 279, 310.

Anopheles Species in East Pakistan and Assam

]aconitus

aitkenii aithenii

aitkenii hengalensis

annandalei annandalei

annandalei interruptus

^annularis

barbirostris ahomi

\barbirostris barbirostris

barianensis

"^culicifacies

\ßuviatilis

gigois baileyi

gigas gigas

gigas simlensis

insulaeßorum

jamesii

"^jeyporiensis candidiensis

jeyporiensis jeyporiensis

karwari

\koGhi

See footnote 6, p. 10.

Heucosphyrus balabacensis

lindesayi

\maGulatus

majidi

^mininiiis

\nigerrinhus

palUdus

"^ philippinensis

ramsayi

\sinensis

splendidus

\stephensi

\subpictus

"^sundaicu^

tessellatus

theohaldi

^umbrosus

vagus

\varuna
82 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE - EAST PAKISTAN AND ASSAM

Yellow Fever.—This disease is not present in East Pakistan or Assam.

Reference: 256.

Dengue.—Both of the principal vectors of dengue fever,

Aedes aegypti and A, alhopictus, are widely distributed in

East Pakistan and Assam and are responsible for the heavy

endemicity of the disease, especially in the Ganges Delta area

east of Calcutta, where the disease has prevailed in an especially intense form. It is estimated that dengue can occur

in epidemic form in any endemic area whether either or both

vectors are present, especially amon^ nonimmune persons.

In East Pakistan and Assam dengue is especially prevalent

just after the rainy season.

References : 256, 310.

Encephalitis.—No information is available.

Filariasis.—Filariasis was introduced about 30 years ago

into the tea-garden sections of the Pachhar district, lower

Assam, where it is endemic in the small villages and where

it has caused much elephantiasis.

Both Wuchereria bancrofti and W, malayi are present in

the native population, the former transmitted by Cidece

pipiens quinquéfasciatus^ the latter, causing most of the infections, by Mansonia annulifera^ M, uniformis^ and M, indiana, AU these mosquitoes have been found with high

rates of natural infection. Bancroftian filariasis has been

found in rural areas in the Birbhum District of Bengal and

is transmitted by Anopheles philippinensis. Wild-caught

specimens of this mosquito have been found with infective

larvae of W, bancrofti. It is not known whether quinquefasciatus is present in Bengal and Assam.

References : 195, 319.

South China

(Figs. 51 and 52)

Physical Features.—The part of China treated here has

a northern boundary that corresponds to the northern

borders of the Provinces of Chekiang, Kiangsi, and Hunan,

follows latitude 29° N. across the southern part of Szechwan,

and ends at the northern extremity of Burma. This line

corresponds to the northern limit of extension into China of

a great part of the oriental mosquito fauna, even though

there is a Palearctic element reaching south of it. Excepting Manchuria, the line divides the southern third from the

northern two-thirds of the country. South China contains

most of the country's mountains and also the plateaus of

Yunnan and Kweichow in the southwest, the valleys of the

West, North, and East Kivers, and a rather narrow coastal
MOSQUITOES OF MEDICAL IMPOBTANCE 83

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE — SOUTH CHINA

MC FULL-GROWN LARVAEï^^ FIG. 51

LONG PLEURALS

OF 3RD GROUP

LONG PLEURALS

OF 2ND GROUP

ABDOMINAL PALMATES METATHORAGIO PAL MATES

ANOPHELES

~ JEYPORIENSIS JEYPORIENSIS

AND JEYPORIENSIS CANDIDIENSIS

ANOPHELES

MINIMUS AND FLUVIATILIS

ANTENNAL HAIR

_ ANOPHELES

— MACULATUS

HE/KÜ OF ANOPHELES LARVA

ANTENNAL

HAIR INNER

CLYPEALS

ANOPHELES

SINENSIS AND NIGERRIMUS

AEDES

ALBOPICTUS

ANOPHELES LARVA

PLEURAL BRISTLES

plain in the southeast. The large island of Hainan lies off

the coast in the South China Sea. The Ryuku Islands are

also included in this region.

The climate of South China is regulated by monsoons and

is subtropical, with mild winters and an annual rainfall of

about 80 inches. About 90 percent of the total arable land

in the extreme south is planted to rice, which is extensively

cultivated and is double cropped.

Reference: SOI,

Malaria.—The areas of highest endemicity are southwest

Yunnan and a coastal strip beginning at the lower Yangtze

River and extending north to include Hainan. As in much

of southeastern Asia, foothill malaria is one of the most

serious problems in South China and is particularly intense

in southwestern Yunnan.

During construction of the Burma Road and the YunnanBurma Railway, large numbers of fatal malaria cases caused

by Plmrriodium, faldfarwm occurred among native laborers,

and this disease has accounted for high morbidity and mortality rates in military campaigns throughout the area.

PlasTïiodium malariae is a rather common and surprisingly

prevalent parasite, but its distribution is sporadic.

Anopheles minimus and A. jeyporiensis candidiensis are

the principal vectors of most foothill malaria in South China.

During the dry season minimus tends to disappear, since most

breeding places completely dry up, but populations in some

localities are very large near the end of the rainy season and

immediately afterward. It is such a prolific breeder in

Yunnan that it has even been found in lowland plains at the

edges of swamps and terraced ricefields. A, j. candidiensis

is often found with minimus but it is not so abundant. It is

found at the edges of seepages and fallow ricefields, and

throughout South China where its numbers are large it is

important, though slightly less so than minimus. This

species is occasionally found with very high rates of natural

infection. In the vicinity of Hong Kong, for example, 9 percent of wild-caught mosquitoes were infected.

Anopheles sinensis^ a transmitter in the northern twothirds of China, is the most important vector in the vast ricegrowing areas of South China. In general, the species is

most prevalent toward the end of summer monsoon rains.

In South China, as elsewhere, sinensis is noticeably zoophilic,

and natural infection rates are generally low. Therefore, in

areas where this species transmits malaria, large populations

of the mosquito must be present.

In Ceylon and throughout much of India Anopheles culicifacies is one of the outstanding malaria vectors. Its distribution in South China is restricted to Yunnan, where it

reaches the easternmost limit of its range and where it has
84 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

LONG PALPS ON

ANOPHELES

.PROBOSCIS

PATTERN ON PALP

4 NARROW WHITE BANDS

SHORT PALPS ON

ALL OTHERS

"PATTERN ON THORAX

MOSQUITOES OF MEDICAL IMPORTANCE — SOUTH CHINA

^^ ^ FEMALES B*^"«^—^

[MALES HAVE BUSHY ANTENNAE ( \¡M^^) AND DO NOT B/TE] FIG. 52

PATTERN ON HIND TARSUS PATTERN ON PROBOSCIS

FOUR LIGHT SPOTS ON FRONT MARGIN

UMMII ANOPHELES

MINIMUS

ANOPHELES

MACULATUS

VEIN 3 WITH LONG CONTINUOUS

DARK AREA

PATTERN ON HIND TARSUS

__ ANOPHELES

— CULICIFACIES

ANOPHELES

JEYPORIENSIS CANDIDIENSIS

_ AEDES

~ ALBOPICTUS

_ AEDES

~ AEGYPTI

been found at least once with relatively high natural infection

rates. Because it is occasionally free of malaria parasites

and because it is a rare mosquito, at least from July to February, compared with minimus^ candidiensis^ and sinensis^ it is

regarded as a secondary vector in this area.

Anopheles maculatus has been found throughout South

China, but it is most often encountered in the western half,

where it is the most common mosquito present. Itis typically

a stream and riverbed breeder. Its numbers are rather constant throughout the year, because its breeding places remain

more or less intact throughoutthe dry season.

Little is know^n of the importance of Anopheles leucosphyrus in South China. It breeds in small shaded pools in

forested areas of Yunnan, and since it is of some importance

in northern Burma, it is included in the keys as a potential

vector in South China.

References : 61, 8J,, 88, 122, 125, 310.

Anopheles Species in South China

jaconitus

aitkenii aitkenii

aitkenii hengalensis

annandaleiinterruptus

\annularis

\harhirostris

"^culicifacies

\fluviatilis

gigas baileyi

gigas simlensis

^amesti

^jeyporiensis candidiensis

jeyporiensis jeyporiensis

karmari

\koGhi

koreicus

kweiyangensis

\leucosphyrus

lindesayi lindesayi

lindesayi pleceau

"^'maculatus

'"^minimus

nigerrimMS

ohamai

philippinenMs hainanensis

\philippinensis philippinensis

saperoi

^sinensis

sineroides

sintonoides

splendidus

\stephensi

\subpictus

\sundaicus

\tessellatus

\umbrosus

vagus

fvarmia

See footnote 8, p. 63.

Yellow Fever.—This disease is not present in South China.

Reference: 256.

Dengue.—Endemic in various localities in South China,

dengue occurs sporadically along the south and east coasts,

extending beyond the northern border of Chekiang Province.

The major foci appear to be port cities, where several epidemics have occurred in the past. Both Aedes aegypti and

A, alhopictus are present in South China. The former
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE - FORMOSA

< FULL-GROWN LARVAE >

85

FIG. 53

r LONG PLEURALS

OF 3RD GROUP

NO SIPHON ON ANOPHELES

^PALMATE HAIR

' POSTSPIRACULAR HAIR /

Ç OUTER CLYPEALS OUTER OCCIPITALS

, 2 TO 5

' BRANCHED

_ ANOPHELES

~ ANNUL APIS

ANOPHELES

SUNDAICUS

:Z ANOPHELES FLUVIATILIS & MINIMUS

ONLY MINIMUS IMPORTANT; \

ARVAE CANNOT BE SEPARATED j

_ ANOPHELES

"" MACULATUS

TOP VIEW OF HEAD OF

ANOPHELES LARVA

.ANTENNA

INNER CLYPEALS OUTER CLYPEALS PROTHORACIC HAIR 1

+

AT TIP, 2 OR 3/ __ ANOPHELES

SINENSIS

PROTHORACIC HAIR 1 ABDOMINAL PALMATES 1&2

NOT

ANOPHELES

TESSELLATUS

SIPHON ON ALL OTHERS

8THSEGMENT ANTENNA BODY HAIRS

ONLY 1

TUFT

PER SIDE SMOOTH^^;;^^

/^/' COMB SCALES //- «tiKiriP-

€.^'' + '-«^^ + // + TR'A'NC,

"—ULAR ROW //

^\ ^EVENLY

/\ SPACED

CC---*- —'•'BSENT

COMB SCALES

_ AEDES

~ AEGYPTI

_ AEDES

" ALBOPICTUS

species, more urban than the latter, has been found in the

vicinity of Hong Kong, but its exact distribution elsewhere

is not known. A, alhopictus is extensively distributed in a

large area near Shanghai, a locality where aegypti is absent.

References : 256, SIO.

Encephalitis.—Outbreaks of Japanese encephalitis occur

in the various Provinces of eastern China from time to time.

In many localities where it is believed to be present, no virus

has ever been isolated, but 80 to 90 percent of the population

in certain areas show immunity to this virus. Little is

known about the vectors in this area.

Reference: 257.

Filariasis.—Both Wuchereria hancrofti and W. malayi are

found in the coastal areas of South China. Their distribution farther inland is not clearly known. Filariasis is centered in low rural areas around lakes and rivers and near

the coast. In South China Anopheles sinensis transmits both

parasites and accounts for a large number of malaria cases

each year. Culex pipiens qwinquefasciatus and C. p. pollens

carry only hancrofti^ and Mansonia uniformis is implicated

in the transmission of malayi in certain areas. In Amoy the

vectors are sinensis and quinquéfasciatus, but they are less

common there than in other parts of Fukien Province, where

the incidence of the disease is considerably higher.

Natural infections have been found in Anopheles minimus^

A. jeyporiensis candidiensis^ and 0. p. quinquéfasciatus in

the island of Hong Kong, where filarial infection in humans

is rare, and in Anopheles splendidus and sinensis on the adjacent mainland. Aedes togoi has been experimentally infected with hancrofti and may be an important vector near

Hong Kong.

References: 124,126,191,198.

Formosa

(Figs. 53 and 54)

Physical Features.—Formosa, which is separated from

the mainland of China by the 100-mile-wide Formosa Strait,

is an island 235 miles long, 90 miles wide, and 13,886 square

miles in area. From north to south parallel mountain ranges

form abrupt, precipitous banks and cliffs and rise in nearly

70 peaks above 10,000 feet. The west and southwest sides

of these mountains descend gradually to form a broad alluvial plain that covers about one-third of the island, which has

a nearly uniform topography. About 70 percent of the is-
86

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE - FORMOSA

< FEMALES >-—

LONG PALPS ON

ANOPHELES

«PROBOSCIS

\MALES HAVE BUSHY ANTENNAE (i

PATTERN ON PALP

,LAST SEGMENT LIGHT AND DARK

.NO LIGHT SPOT AT CROSS VEIN

) AND DO NOT BITE\

CLYPEUS

FIG. 54

SCALE — ANOPHELES

TUFT S/NENS/S

VLAST SEGMENT ENTIRELY LIGHT,

LIGHT BANDS AS WIDE OR WIDER

V. THAN INTERVENING DARK BAND

SMALL LIGHT SPOT IN BASAL THIRD

_ ANOPHELES

"" MINIMUS

LIGHT SPOT AT CROSS VEU

PATTERN ON PALP ^r PATTERN ON HIND TARSUS ~^

ANOPHELES

ANNULARIS

2 DARK SPOTS BEHIND

^MIDDLE MARGINAL SPOT

ANOPHELES

SUNDAICUS

\LAST SEGMENT LIGHT AND DARK,

3 BROAD AND I NARROW

WHITE BANDS

_ ANOPHELES

MACULATUS

_ ANOPHELES

~ TESSELLATUS

PATTERN ON THORAX ^

•

_ AEDES

"" ALBOPICTUS

PATTERN ON HIND TARSUS

_ AEDES

~" AEGYPTI

ADULT MOSQUITO WITH PARTS LABELED

THORAX..^

\

"'''\r^J^^^^^~~^^.WING

ANTENNA___^^

CLYPEUS

ABDOMEN

^ \^¡p-ií^ / ^1 ^ ^_^y'~~'^yZi— "'

PALP_ _ 2^t=3^

^/ "-^^ \ Y-TIBIA ^

FORE l-EG.^,/

// MID LEG Ä \

^ \ HIND LEG^

i_ PROBOSCIS \ ^

TARSUS

land's population lives on this plain, where rice and sugarcane are the most important crops.

A southwest monsoon brings light rains to the southern

part of the island in April and May and a rainy season from

June to September ; the rest of the year is relatively dry. A

northeast winter monsoon brings rain to northern Formosa

during the dry season in the south. Cool seasons in the island

are well marked, but temperatures never become low enough

to prevent year-round mosquito breeding except in the high

mountains.

References: 85,301.

Malaria.—The principal malaria problem in Formosa is

centered in the western and southwestern plains, where extensive cultivation of rice supports large mosquito populations. In addition, there is a small amount of malaria in

the adjoining foothills.

The malaria is usually caused by Plasmodium vivax^ even

though most deaths are due to P. falciparum ; P. malariae is

rare in the island.

Alluvial-plain malaria is transmitted principally by

Anopheles sinensis^ which has two seasonal population peaks.

The first occurs in February and March and is due to the

second flooding of the first rice crop of the year. The second

peak in September and October, caused by the flooding of

the second crop, is usually slightly lower than the first. At

Taipeh in northern Formosa two peaks also occur, the earlier

one slightly later than in the south because of the later date

of rice planting. A. sinensis is the most prevalent mosquito

in the plains area. Because it is primarily zoophilic, it requires large human populations to maintain endemic malaria.

Three other alluvial-plain Anopheles species have been

found infected with malaria parasites in nature. Their relative importance is not known with certainty. They are regarded as secondary vectors during the time of year when

their numbers are large. The commonest of these, Anopheles

tessellatus^ breeds in pools formed by autumn rains in southern Formosa's sugarcane fields. Its peak of abundance is

high in the fall and decreases as the pools dry out after the

end of the rainy season. Anopheles annularis larvae are

often found in sugarcane fields with tessellatus larvae. The

peak of abundance of these two species occurs simultaneously

in the fall. Anopheles sundaicus is found during the dry

season because of its habit of breeding in small pools with

sandy or stony bottoms, mainly along riverbeds. When these

rivers are filled during the rainy season, breeding is reduced

to a minimum. A. sundaicus has a low peak of abundance in

the fall, corresponding to that of sinensis^ in areas where it

occasionally breeds in ricefields.

The larvae of Anopheles nhinimus live at the margins of

small streams and irrigation canals, primarily in the foothills. The adults have two peaks of abundance, one about

May and another in the autumn. Since minimus appears in

relatively large numbers in houses and is normally found

with a high rate of natural infection, it is the second most

important vector in Formosa, transmitting most of the malaria in the foothills.
MOSQUITOES OF MEDICAL IMPORTANCE 87

Anopheles maculatws^ like the preceding species, breeds in

foothill stream-bed pools in central and southern Formosa,

reaching a dry-season population peak in April and May. In

contrast to minimus^ it has rarely been collected in houses

there. It is included here as a potential vector because of its

high natural infection rates.

References : 67, %5, 89, 310.

Anopheles Species in Formosa

aitkenii hengalensis lindesayi pleccau

"^annularis •ludlowi ludlowi

\harbirostris harbirostris "^maculatus

harhumbrosus ^minimus

fluviatïlis ^sinensis

gigas haileyi splendidus

insulaeflorum 'subpictus indefinitus

\jeyporiensis candidiensis ^sundaicus

\kochi ^tessellatus

\leucosfhyrus (var. not vagus vagus

known with certainty) vulgaris

lindesayi lindesayi

See footnote 6, p. 10.

Yellow Fever.—As in other areas of the Orient, no cases

of yellow fever have been reported from Formosa. However, Aedes aegypti is common here and throughout adjacent

islands, and its habits here as elsewhere are definitely urban.

References: 27, 256.

Dengue.—This disease occurs in numerous epidemics, and

sporadic cases appear in the southern part of the island.

Aedes aegypti is undoubtedly responsible for most of the

urban cases and A. alhopictus for the more rural ones, since

the latter mosquito tends to be less domestic in its habits.

References: 21, 256.

Encephalitis.—Severe epidemics of Japanese encephalitis

occur every year in Formosa. Little information is available

regarding the distribution of the disease or the nature of the

vectors. Culex tritaeniorhynchus^ the vector of Japanese

encephalitis virus elsewhere, is present in the island.

Reference : 251.

Filariasis.—This disease is rare in natives in Formosa, and

even in areas with the heaviest infections, mosquitoes are

found with low parasite rates. Wuchereria hanorofti is the

only parasite in the island, and itis transmitted only by Oulex

pipiens quinquefasciatus.

Reference: 5^-^.

Burma

(Figs. 55 and 56)

Physical Features.—Burma, which is roughly oval in

shape, has a narrow, southern 400-mile arm along the west

coast of Thailand, consisting of extiensive inland mountain

ranges and the narrow Tenasserim cjoast. An arc of mountains separates Burma from China on the east and from

Assam and the Bay of Bengal on the west. These mountains

are an extension of the Himalayas and rise to 20,000 feet in the

extreme north. In the western half of Burma is the extensive

valley of the Irrawaddy River. Its delta extends along most

of the southern and southwestern coasts. Eice is the most

important crop and is grown extensively for export throughout this region. The Arakan range separates the Irrawaddy

Valley from the Bay of Bengal, leaving a narrow coastal

strip. The eastern half of Burma consists of the Shan

Plateau and the Karenni Hills, the latter cut into deep

gorges by the tributaries and main course of the Salween

River, which empties into the Bay of Bengal on the southeast coast.

Areas of highest rainfall, with 200 inches per year, are the

Arakan and Tenasserim coasts. This rain is brought by a

southwest monsoon from May to October. The dry zone is

situated in the lee of the Arakan Yoma Mountains and receives about 25 to 40 inches of rain annually. November to

February in Burma is cool and February to May hot, with

little rain in either season.

Reference : SOI.

Malaria.—Malaria is an urgent medical problem in the

foothills of Burma, where most cases are found. Malaria

occurs principally in the mountain ranges in the north and

extends southwest throughout the Arakan range, southeast

through the Shan States, and along the Burma-Thailand

border into the Malay Peninsula. Because of their immunity, natives in these areas appear to be less affected than

immigrants.

Plasmodium falcipanmi^ P. vivax^ and P. malariae are

found in this country, with falciparum predominating in

most areas. Large numbers of cases of blackwater fever are

admitted to hospitals in Burma.

Throughout the foothills of Burma Anopheles minimus

is the most widespread and important vector. This mosquito

breeds in clear, fresh water at the grassy edges of slowly

running streams and is more distinctly anthropophilic than

almost any other oriental malaria vector.

Anopheles jeyporiensis candidiensis is another foothill

mosquito, inhabiting a large variety of fresh-water collections. It varies more in its anthropophilism from locality to

locality than does minimus^ but it has definitely been implicated in the transmission of premonsoon malaria along the

Burma-East Pakistan border, where adults commonly are

found in hillock country and avidly attack man.

Anopheles leucosphyms halabacensis^ principally a forest

species, is a vector in central Burma during the rainy season

and may also play a part in transmission farther south and

west in the Arakan area. These three Anopheles species are

the most important vectors in Burma, especially when they

are abundant.

Anopheles sundaicus^ a brackish-water coastal species, is

found in many areas of the Burma coast, but never in association with mangrove. This vector tends to cause small epidemics at irregular intervals within restricted and isolated

areas, especially along the Arakan seaboard. It is zoophilic

to a large extent and may be easily deflected from feeding on

man in this area by the presence of cattle. Very little malaria is found along the south Burma coast.

Anopheles stephensi^ an extremely important vector in

India and westward, has a range that extends into northern

Burma. Since it has been found infected in nearby countries,

it has been included as a potentially important vector. It is

principally urban over most of its range.

Anopheles culicifacies^ one of the most important transmitters of malaria in India, has been found infected in Minbu,

upper Chindwin, the northern Shan States, and along the

I^do Road. Because it is an effective vector in other parts

of its range and because it has been found with natural infections in Burma, this mosquito may be important.

On the northern point of Ramree, the largest island off

the Arakan coast. Anopheles annularis has been implicated

as the fresh-water vector on the basis of epidemiological evidence. This species is not included in the keys, because it

is of such little importance, considering Burma as a whole.

References : 61, 91,122, 240, 242, 310, 333.
88 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE — BURMA

i FULL-GROWN LARVAE I

ANOPHELES

JEYPORIENSIS JEYPORIENSIS

Zl kHuJEYPOR/ENSiS CANDIDIENS/S

ANOPHELES

LEUCOSPHYRUS

FIG. 55

I ONLY CANDIDIENSIS IMPORTANT; LARVAE CANNOT BE

SEPARATED

ANOPHELES

M¡NIMUS

hHXi FLUVIÁTIL!S

'ONLY MINIMUS \

IMPORTANT; I

LARVAE CANNOT

BE SEPARATED /

ANOPHELES

SUNDAICUS

AND SUBPICTUS

ONLY SUNDAICUS

IMPORTANT; LARVAE

CANNOT BE SEPARATED

__ ANOPHELES

" STEPHENS!

AEDES

AEGYPTI

AEDES

AL BOPIC TUS

TOP VIEW OF HEAD OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS^

/ OUTER ^y HIND

CLYPEALS /'y CLYPEALS

1

ANOPHELES LARVA

PROTHORACIC

HAIRS 1^4 2 W 1/ PLEURAL BRISTLES

POSTSPIRACULARS

Anopheles Species in Burma

\(iconitiis

aitkenii

\annui(Kns

\harbirostris

'^aidicifactes

\ßv/oiatilis

gigas haileyi

jamesii

^jeyporiensis candidiensis

jeyporiensisjeyporiensis

karwari

\kochi

kyondaw^nsis

HeuGosphyms haZdbacensis

jmaculatus maculatus

maculatus willmori

^rwimmus

\nigerTÍmms

pallidus

fphilippinensis

rainsayi

\sinensis

splendidus

\stephensi

fstihpictu^ subpictus

^sundaicus

\tessellatus

theohcddi

\unibrosus

vagus

\varu7m

See footnote 6, p. 10.

Yellow Fever.—This disease is not present in Burma.

Reference: 26Q.

Dengue.—Dengue fever in the past has been an important

disease in southern Burma. At least one epidemic affecting

a large number of persons has occurred. Dengue has never

been recorded from the Shan States or the old Burma Road

and does not occur in these areas. Aedes aegypti^ the vector

of classical yellow fever outside the Orient, is the most

important vector of dengue in Burma. Aedes aihopictus is

probably widespread, but its importance in the transmission

of dengue in Burma is not well known.

References : m2,256, 310.

Encephalitis.—Japanese encephalitis has been reported

from the delta area of the Irrawaddy, but as far as is known

no cases have occurred in humans, nor have any human sera

been found with antibodies to the virus.

Reference: 257.

Filariasis.—^Although both Witehereria hancrofti and W.

malayi are present in Burma, the latter parasite predominates

and causes a great deal of disease, especially along the Tenasserim and Arakan coasts and in Mandalay. The vectors of

malayi are presumably members of the Mansonia subgenus

Mansonoides^ as elsewhere in the Orient.

Reference: SIO.
MOSQUITOES OF MEDICAL IMPORTANCE 89

MOSQUITOES OF MEDICAL IMPORTANCE - BURMA

; FEMALES ^

FIG. 56

PATTERN ON HIND TARSUS

ANOPHELES

CULICIFACIES

LIGHT BAND AT TIP OF PAUP

fg*^ / ; WIDER THAN PRECEDING DARK ^.,^«^^, r-r-

,1 ''5lfe\ ' 1 BAND^^ ANOPHELES

" + it^ • ^^H^ = JEYPORIENSIS

CANDIDIENSIS

SHORT PALPS ON

ALL OTHERS

PATTERN ON^ THORAX

AEDES

AEGYPTf

_ AEDES

-" ALBOP/CTUS

ADULT MOSQUITO WITH PARTS LABELED

THORAX^

Indochina

(Figs. 57 and 58)

Physical Features.—Indochina is a crescent-shaped country on the southeast coast of Asia and is composed of the

States of Vietnam, Laos, and Cambodia. Vietnam consists

of Tonkin in the north, Cochin China in the south, and the

long, narrow coastal strip of Annam between them. Tonkin's

scattered northern mountains are thinly populated and steep

walled, and its southern ricegrowing areas are the most extensive in Indochina. Annam extends for 800 miles between

the Annamese Cordillera on the west and the South China

Sea on the east. The Cordillera expands southward to form

the Moi Plateaus in the low^er end of Annam. Cochin China,

another important ricegrowing area, is a flat alluvial plain

irrigated by the Mekong Eiver, which flows almost the entire

length of Indochina and ends in an extensive delta on the

south coast. The southwestern tip of Cochin China is expanded into the mangrove-covered Camau Peninsula. The

kingdom of Laos consists of a northern area of wooded ranges

and plateaus cut by narrow valleys and a southern area with

arid, sparsely forested, limestone terraces. Cambodia, separated from the southeast border of Thailand by an extensive

mountain range, is a flat alluvial plain like that of Cochin

China, where rice is the principal crop.

Indochina has a tropical climate dominated by monsoons

from the South China Sea. These winds bring high yearround temperatures to southern Vietnam and rain from May

to October. Farther west, Cambodia receives 25 to 40 inches

of rain annually, the summer floods of the Mekong providing

a large part of the water needed for agriculture. In the

north the climate is more seasonal, with cooler winters and

precipitation up to 80 inches per year in Laos. A dry, cool

season usually occurs from November to February and a

hot season from February to May, after which rain again

begins.

Reference: SOI.

Malaria.—The principal malaria problem exists in the

foothills of all three States of Indochina. The disease is

severe, and blackwater fever is present, especially in the

mountains of Annam and Tonkin.

Pldsmodiwn faLcifmrnin malaria predominates in many

parts of Indochina, and P. vivax accounts for an equal number of cases in other parts. Pldsmodium malariae rarely

accounts for over 10 percent of the cases in any locality.
90 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - INDOCHINA

^FULL-GROWN LARVAE )l* FIG. 57

LONG PLEURALS

OF 1ST GROUP

TWO BARE,

ONE

HAIRED

ABDOMINAL PALMATES

LONG / NOT ^ \

FILAMENTS / ^ \

INNER CLYPEALS

¥ 7~"

_ ANOPHELES

~~ SUNDAÍCUS

OUTER CLYPEALS

WITH SHORT , SIDE HAIRS J

ANOPHELES

M/NIMUS AND

FLUVIATILIS

f ONLY MINIMUS IMPORTANT.

\ LARVAE CANNOT BE SEPARAT

TOP VIEW OF HEAD OF

ANOPHELES LARVA

ANTENNAL

HAIR

/ INNER

' OUTER ! CLYPEALS

1 CLYPEALS ^

\ KANTENNAV > n

WWImj/

MACULATUS TO^ffff

\\Jl

TANT; \

ANOPHELES

A CO NIrus

INNER CLYPEALS

FAR APART,

SPARSELY

HAIRED

OUTER CLYPEALS PROTHORACIC HAIR 1

ANOPHELES

= JEYPORIENSIS

CAND/DIENSIS

MESOTHORACIC HAIR 5

2 TO 7

-f> BRANCHED

AT TIP

SIPHON ON ALL OTHERS

ANOPHELES

S/NENSIS

ONLY 1

TUFT PER

SIDE

COMB SCALES

-|- IN ONE

REGULAR ROW

y ^ EVENLY

W ^^ SPACED

ANTENNA

SMOOTH y^y^

BODY HAIRS COMB SCALES

m^^^^^ =

AEDES

A EGYPTI

_ AEDES

ALBOP/CTUS

ANOPHELES LARVA

PROTHORACIC

HAIR I PLEURAL BRISTLES

l^'^''~~^'f"l/X'^^P^LM/KTZ HAIRS

^^1. „^^^^""-^ ABDOMINA

POSTSPIRACULARS

All malaria up to about 1,500 feet is transmitted chiefly by

Anopheles minimus^ by far the most important vector in

Indochina. Itis strongly anthropophilic and breeds not only

in the moving water of streams but also in clear water at the

edges of swamps, ricefields, and borrow pits.

Anopheles jeyporiensis candidiensis^ another foothill species, is found in standing or running, grassy, clear water.

Although it is slightly less anthropophilic than minimus^ it

plays an important part in the transmission of foothill malaria under certain conditions.

Anopheles maculatiis^ a third species associated with foothill habitats, breeds in a wide variety of water collections,

but, like candidiensis^ it appears to be zoophilic except when

conditions are unfavorable. The females then attack man

readily and have been found highly infected in nature, especially in Tonkin. This species is present in large numbers

in newly cleared jungle areas.

Another less important type of malaria is of low endemicity, with occasional epidemic outbreaks. This occurs in the

extensive ricegrowing areas of Tonkin in the north and

Cochin China and Cambodia in the south. It is transmitted

chiefly by Anopheles sinensis. With the clearing and cultivation of lowland jungles, this mosquito often appears in

tremendous numbers. Its importance apparently depends on

its large populations, since it is usually found with relatively

low natural infection rates in Indochina and is chiefly zoophilic, except where animals and livestock are not present

in large enough numbers to support it. Refugees from foothill areas of Tonkin have brought a great deal of malaria

to the delta areas, where sinensis can transmit it.

Anopheles aconitus has variable breeding habits but is

found most often in ricefields and fresh-water ponds in

mountain and plains areas. It has been strongly suspected

as a vector in southern Annam. However, in Tonkin,

Cochin China, and Cambodia its importance is unpredictable

and it is regarded as a secondary vector.

With few exceptions Anopheles sundaicus breeds only in

brackish water and is restricted to the southern coast of

Indochina, where it is found in lagoons, swamps, and saltwater fishponds. It is of less importance than the foregoing species because of its restricted distribution and its less

definite anthropophilism, but it transmits coastal malaria,

which seldom occurs except in occasional small epidemics.

References : 67, 68, 277, 310.
IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

LONG PALPS ON

ANOPHELES

ANTENNAE

-PROBOSCIS

SHORT PALPS ON

ALL OTHERS

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE - INDOCHINA

^ FEMALES ^

\_MALES HAVE BUSHY ANTENNAEV^^^^^ J AND DO NOT BITE ]

91

FIG. 58

NO LIGHT SPOT AT CROSS VEIN

^ LIGHT SPOT AT CROSS VEIN

PATTERN ON THORAX

PATTERN ON PALP

, LAST SEGMENT LIGHT AT TIC i

». DARK AT BASE '

PATTERN ON HIND TARSUS

LAST SEGMENT ENTIRELY LIGHT,

LIGHT BANDS AS WIDE OR WIDER '

THAN INTERVENING DARK BAND

PATTERN^ON PALP

BROAD AND 2 NARROW

WHITE BANDS

-_ ANOPHELES

S/NENS/S

PATTERN ON HIND TARSUS

_ ANOPHELES

~" ACONITUS

PATTERN ON PALP

LIGHT^BAND TWICE AS LONG AS ANOPHELES

+ CNLN ^'^^'^ ^^'^'^ — = JEYPORIENSIS

CANDIDIENSIS

PATTERN ON HIND TIBIA

_ ANOPHELES

"~ SUNDAICUS

^ ANOPHELES

MACULATUS

2 BROAD AND 1 NARROW

WHITE BANDS

ALL WHITE-- T;

_ AEDES

" ALBOPICTUS

PATTERN ON HIND TARSUS

Anopheles Species in Indochina

"^aeonitus

aitkenii aitkenii

aitkenii iengalensis

alongensis

annandalei annandalei

annandalei interruptus

\annularis

\haeBai

\l)arbirostris iarhirostris

harhumbrosus

jf.cuUcifades

\ßuviatiUs

gigas haileyi

jamesii

"^jeyporiensis candidiensis

jeyporiensis jeyporiensis

karwari

\koGhi

See footnote 8, p. 63.

Yellow Fever.—This disease is not present in Indochina.

Reference: 256.

\leucosphyrus

lindesayi

litoralis

"^^maculatus

macuUpalpis

"^minimus

\nigerHnhus

pallidus

fphilippinensis

^sinensis

splendidus

\stephensi

{suhpictiis

"^sv/ndaicus

\tessellatus

\unhbTOsus

vagus vagus

\varuna

Dengue.—This disease occurs throughout Indochina, having first appeared in 1870 at Haiphong and afterward at

Hanoi and Saigon, all in rather direct contact with ocean

traffic. Both urban Aedes aegypti and the slightly wilder

A. albopictus are extensively distributed throughout Indochina and are responsible for the transmission of this disease.

References : 256, 287, 310.

Encephalitis.—A virus very closely resembling that of

Japanese encephalitis has been recently isolated from 98

French soldiers in the vicinity of Tonkin. Its identity with

the Japanese type is not yet absolutely certain, nor have

studies been completed to determine its vector. Antibodies

to Japanese encephalitis virus have been found in human

sera at Hanoi.

References: 257,278.

Filariasis.—The principal medical problem due to filariasis in Indochina occurs in the Red River Delta of Tonkin.

Filariasis was first found there in 1939, and both Wuchereria

hancrofti and W. malayi were eventually found to be prevalent, not only in low land but in higher areas. Natural infections with infective stages of the parasites have been
92

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE - THAILAND

NO SIPHON ON ANOPHELES

POSTSPIRACULAR HAIR^

LONG PLEURALS

OF 3RD GROUP

"d

LONG PLEURALS

OF 2ND GROUP

ONE

HAIRED

: FULL-GROWN LARVAE )

ABDOMINAL PALMATES

FIG. 59

METATHORACIC PALMATES ANTENNAL HAIR

ANOPHELES

MACULATUS MACULATUS

AND MACULATUS WILLMORI

SIPHON ON ALL OTHERS SIPHON

4N^-7 ONLY 1

^-M TUFT

PER SIDE

>

+

COMB SCALES

IN ONE

REGULAR ROW 'M (NOT ON A PLATE)

^ V EVENLY

// ^/SPACED 11/

r;l_ ABSENT

found in Anopheles minimus^ A, jeypoHensis^ A. vagus^ A,

maculatus^ A, sinensis^ Oulex pipiens qwinquefasciatus^ Mansonia uniformis^ M. indiana^ and M. annuUfera,

In the low areas of Tonkin sinensis transmits both hancrofti and malayi^ especially in areas where there are no

cattle. In higher areas sinensis is diverted from man by

these animals and is not so effective a vector. In many of

the low areas sinensis is a more important vector of hancrofti

than quinquefasciatiis^ and it is an important vector of

malayi in areas where Mansonia species are absent. In delta

areas indiana predominates, especially in the vicinity of

Hanoi, where larvae are most commonly found on submerged

stems of Eichhornia cra^sipes and Salvinia rotundifolia^

rarely on Pistia. M, uofiiformAs is rare, and indiana is probably not a vector of outstanding importance. Not much is

known of the distribution of filariasis in the remaining parts

of Indochina. In Cambodia annulifera is the most common

and probably the most important Mansonia species, and in

Cochin China indiana is absent, annulifera is rare, and uniformis is the most common important vector.

Thailand

(Figs. 59 and 60)

Physical Features.—Situated between Indochina and

Burma in southeast Asia, Thailand (Siam) extends in a narrow southern peninsula to the northern border of Malaya.

Geographically the country is divided into (1) the northern

highlands, consisting of rather parallel, deeply forested

limestone ranges with deep valleys; (2) the densely settled,

west-central flood plain of the important Chao Phraya River,

containing the important rice-producing areas; (3) the eastern Korat Plateau, devoted mainly to stockraising and separated from the western alluvial plain by a central mountain

range; and (4) the southern peninsular area, which is largely

mountainous and mineral bearing.

The central mountain range divides the northeast coast,

with its October-to-February rainy season, from the west

coast, which has maximum rainfall from May to October.

The latter is brought by monsoons from the southwest, and

a cool, dry season from October to February is caused by

monsoons from the northeast. A hot, dry season generally

lasts from March to May.

Keferences : 135-139,328. Reference: 301.
MOSQUITOES OF MEDICAL IMPORTANCE 93

IMPORTANT

A SPECIMEN MUST HAVE

ALL. CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - THAILAND

Il FEMALES »^

IMPLES HAVE BUSHY ANTENNAE ( j^^^) ^^^ DO NOT BITE^ FIG. 60

LONG PALPS ON

ANOPHELES

ANTENNAE

.PROBOSCIS

PATTERN ON HIND FEMUR

AND TIBIA

FRINGE SPOT AT TtP OF VEIN 6

PATTERN ON HIND TARSUS

4 NARROW WHITE BANDS

PATTERN ON HIND TARSAL SEGMENTS 3, 4, AND 5 -

PATTERN ON HIND TARSUS

ANOPHELES

CULICIFACIES

ANOPHELES

NIGERRIMUS

ANOPHELES

SINENSIS

TOP OF 2ND ABDOMINAL

SEGMENT WITHOUT PATCH

+ OF SCALES ANOPHELES

MACULATUS MACULATUS

SHORT PALPS

ALL OTHERS

ON

\

r PATTERN ON THORAX

- AEDES

ALBOPICTUS

J^P

)

'^n^f^

PALPS ^Jr

/ \ f

~ AEDES

AEGYPTI

Malaria.—The most important malaria problem is in the

foothills throughout Thailand. Blackwater fever caused by

Plasmodium falciparum is hyperendemic in much of this

region ; P. vivaw malaria also occurs but is not so abundant

or important.

Although the mountains of the south are highly malarious,

they are less so than those on the Burma frontier in the

northwest. A large part of this foothill malaria is maintained by Anopheles nhinimus. As in many other parts of

southeast Asia this mosquito breeds at the margins of clear

fresh-water streams and at the clear edges of larger water

collections. The adults are highly anthropophilic and are

constantly found in close association with man.

Anopheles sundaicus is responsible for a low endemic malaria along the coasts of southern Indochina, Malaya, and

Burma, and probably coastal Thailand is also included.

The other species of Anopheles treated in the keys have

never been definitely associated with malaria as a result of

field studies in Thailand. They are included because (1)

they are proved transmitters in countries bordering Thailand, and (2) they are present in Thailand, and under certain conditions one or more of them would have to be seriously considered in any malaria-control project.

Anopheles Gulicifacies^ the most important vector of malaria in India and one of the most important in Burma, is

present over the northwestern half of Thailand. The larvae

live in many kinds of water collections, and the females bite

both man and animals. The species may be a secondary

vector.

Anopheles leucosphyrus is distributed throughout Burma

and Assam, where it is associated with forests, and it has

been implicated as a vector in northern Burma. It is not

regarded as an important mosquito in Thailand.

In Malaya and parts of the Greater Sundas Anopheles

maculatus maculatus is an important malaria transmitter.

Its range extends well into South China, where it has seldom

been implicated as an actual vector. Its importance in Thailand has not been adequately studied.

Anopheles aconitus is present throughout southeastern

Asia, and in Indochina it is regarded as an important species.

Some evidence indicates that it may contribute to the spread

of malaria in Thailand. This species varies from locality to

locality in its ability to transmit the disease.

Anopheles sinensis and A, nigerrimus have not been thoroughly investigated. A. sinensis is the important vector in

central and northern China, but to the south its importance

becomes less, whereas that of its close relative, nigerrimus^
94 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

becomes greater. Both species are found in large bodies of

water and may be important in the ricegrowing areas of

central Thailand.

References: 67,122,310.

Anopheles Species in Thailand

"^aconitus

aitkenii

alhotaeniatus

annandaleiinterruptus

\annulaxns

argyropus

haezai

\harl)irostris

iarbumbrosus

hulkleyi

"^Gulicifacies

\filipinae

\-ßuviatilis

gigas formosus

gigas sumatrana

indiensis

insulaeßorum

\jeyporiensis candidiensis

karwari

\kochi

Heucosphyrus

See footnote 6, p. 10.

ludloiöi (fresh-water form)

"^maculatus maculatus

majidi

^minirrhus

"^nigerrimus

pallidus

palmatus

peditaeniatus

\philippinensis

ramsayi

separatus

"^sinensis

splendidus

\stephensi

subpictus malayensis

\suJ)pÍGtus subpictus

"^sumdaicus

tessellatus

\umbrosus

vagus

\varuna

Yellow Fever.—This disease is not present in Thailand.

Reference: 256.

Dengue.—In the past dengue has occurred in annual epidemics in southern and eastern Thailand, but the extent of

these outbreaks is not known. A number of cases are reported annually elsewhere in this country. Aedes aegypti

and A, albopictus are distributed extensively throughout the

country, the former being the most urban of the Thailand

culicines and the latter occurring in more rural locations.

References : 2b6, 310.

Encephalitis.—Japanese encephalitis virus has been

reported from the southern half of Thailand, but its vector

there is not known with certainty.

Reference : 257.

Filariasis.^—This disease is endemic only on the western

side of Thailand's southern peninsula between the Indian

Ocean and the South China Sea. It is almost exclusively

rural and is caused by Wuehereria rnalayL The vectors are

the Mansonia species that normally transmit malayi over

most of the Orient and are annulifera^ longipalpis.^ unifornds.,

and indiana. Wuehereria bancrofti^ although not indigenous

to Thailand, is constantly being introduced and may be

transmitted by Culex pipiens quinquefasciatus., which is

abundant.

Reference : 196.

Malaya

(Figs. 61 and 62)

Peninsula and borders Thailand at about latitude 6° N. It

is the southernmost part of Asia and is separated from Sumatra by a narrow strait. The mountainous backbone of the

peninsula, with peaks over 7,000 feet, extends well into

Malaya from the north. Much of the west coast consists

of mangrove and mud flats. Large amounts of rice are

grown betw^een the coast and the mountains and in the northcentral part. The east coast is largely rough and sandy.

About 80 percent of the land is covered by jungle. Johore

State in the south is rather flat and covered by jungle, with

occasional low mountain peaks of about 3,000 feet. Singapore is an island at the southern tip of the peninsula. Its

west and north coasts are covered with mangrove, and its east

coast consists largely of cliffs. It is principally agricultural.

In Malaya the annual rainfall averages 100 inches in the

plains and over 200 inches in the hills. Northeast and southwest monsoons slightly determine the wet and dry seasons.

Reference : 301.

Malaria.—Plasmodium falciparum malaria predominates

almost everywhere in Malaya. It accounts for the high death

rates in this country.

Malaria occurs in Malaya throughout the year, with a

gradual increase through the summer and autumn. It is

particularly intense along parts of the west coast in Selangor

in sunlit breeding places, where mangrove, which is usually

effective in the prevention of Anopheles sundaicns breeding,

has been cut away. Embankments and roads impede the flow

of tides, and the resulting collections of brackish water support heavy sundaicns breeding. The distribution of this

species, one of the most important vectors in Malaya, is

patchy. There are large areas where conditions are favorable for breeding but where the species is not present at all.

The foothills of Malaya are highly malarious, and although

Anopheles minimus^ the foothill vector in most of southeast

Asia, is present in extreme northwest Malaya, it contributes

little to maintaining the disease in Malaya.

On the other hand. Anopheles maculatus^ which occurs in

almost all hilly areas from the coast to 5,000 feet, is more

important. Like mininius^ this species breeds in sunlit, clear,

oxygen-rich water. It has been found with extremely high

natural infection rates, although in certain areas it may be

primarily zoophilic. It tends to invade the foothill areas

that have been cleared of jungle.

Anopheles umbrosus transmits a large amount of malaria,

especially along the densely vegetated west coast. This mosquito breeds almost entirely in the vicinity of jungle swamps.

In localities where it is present with its close relative, Anopheles letifer^ especially at the edges of jungle areas, it is

extremely important. Since letifer lives in cultivated lowland areas close to but not within the jungle, it transmits

much of Malaya's malaria in clearings where man has established plantations.

A dark-winged form of Anopheles barbirostris has been

found with extremely high infection rates in the vicinity of

the Kinta Kiver swamps, where it is believed to contribute

materially to the epidemiology of malaria, although it is not

normally important unless appearing in extremely large

numbers.

The records of Anopheles sinensis and A. nigerrimus in

Malaya are confused. Both are vectors in various parts of

Asia and both have been implicated as vectors at Kuala

Lumpur. Because of its generally greater abundance in

Malaya, nigerrimus is probably the more important of the

two. Both species breed in large swamps and are presumably

vectors of malaria in the ricegrowing areas. Physical Features.—Malaya, which is also known as

British Malaya, is located at the southern tip of the Malay References: 122,272,284^296,310,357.
MOSQUITOES OF MEDICAL IMPORTANCE 95

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - MALAYA

-< FULL-GROWN LARVAE >

r PROTHORACIC HAIR 1

SIPHON ON ALL OTHERS

ANTENNAL HAIR

\ TO l'a DISTANCE

FROM BASE

AT LEAST

27 BRANCHED

ANOPHELES

: NIGERRIMUS

AND StNENSIS

(BOTH IMPORTANTN

LARVAE CANNOT)

BE SEPARATED /

SPIRACULAR APPARATUS

/NOT V

_ ANOPHELES

~ UMBROSOS

TIP OF ANTENNA HIND CLYPEALS

SHORT AND

2 TO 4

BRANCHED-/- 1Í' /

3RD SEGMENT

LATERAL HAIR

13 TO 20 BRANCHED

' 4- AND

HAIR 0

3 OR 4 BRANCHED

_ ANOPHELES

"" LETIFER

ABDOMINAL PLATES

_ ANOPHELES MINIMUS MINIMUS

"" " AND MINIMUS FLAVIROSTRIS

OUTER CLYPEALS

/NOT,\\I/

PLEURALS OF 1ST GROUP LATERAL HAIRS ON

5TH & 6TH SEGMENTS

+

' 2 TO 6 BRANCHED

FIG. 61

TOP VIEW OF HEAD OF

ANOPHELES LARVA

ANTENNAL

HAIR INNER

/ CLYPEALS

' OUTER 1 HIND

; CLYPEALS 1 CLYPEALS

\\ï ^^ u/ \\fe'''''!.!$i '

•^ 1 ñ

#Ji,W^yif

'^m~^mE

\ v-y/

1

PALMATE HAIRS

UNDEVELOPED

_ ANOPHELES

~~ MACULATUS

HIND CLYPEALS

PLACED FAR

I BACK OF

"r INNER

CLYPEALS

COMB SCALES

IN ONE

REGULAR ROW

(NOT ON A PLATE)

COMB SCALES

_ ANOPHELES SUNDAICUS

~~ AND SUBPtCTUS

/ONLY SUNDA/CUS IMPORTANT} N

V LARVAE CANNOT BE SEPARATED j

AEDES

AE6YPTI

_ AEDES

ALBORICTUS

ANOPHELES LARVA

PLEURAL BRISTLES

Anopheles Species in Malaya

\aconitus

aitkenii aitkenii

aitkenii hengalensis

aitkenii stantoni

alhotaeniatus

\annularis

argyropus

asiaticus

aurirostris

\bae3ai

halahaceAisis introlatus

^^'iarhirostris

harhumbrosus

brevipalpis

brevirostris

crawfordi

gigas sumatrana

hunteri

kariimri

\kochi

lesteri

Hetifer

471779 0—59 7

\leuGOsphyrus hacheri

\leucosphynis leucosphyrus

leucosphyms pujutensis

leucosphyrus riparis

lir\desayi eameronensis

ludlowi

^maculatus

'^'7)%inimMS

montanus

"^'nigerrinms

pallidus

peditaeniatus

fphilippÍ7iensis

ramsayi

riparus macarthuri

roperi

separatus

similissinvas

*sinensis

\subpictus malayensis

fsubpictus subpictus

'^sundaicus

\tesseTlatus watsonii

"^umhrosus loellingtonianu^

vagus

See footnote 6, p. 10.

Yellow Fever.—This disease is not present in Malaya.

Reference: 25Q.

Dengue.—This disease is endemic and epidemic in Malaya.

Both Aedes aegypti and A, albopictus are vectors. The former is an almost strictly urban mosquito and the latter not

only is urban but also breeds in rubber and coconut plantations, being more habitually an outdoor mosquito than

aegypti.

References: 25ß, 310, S51,

Encephalitis.—The virus of Japanese encephalitis has

been isolated from four fatalities occurring among British

troops and natives in Malayan jungles and in the island of

Singapore. Although the vector is not known, the nonepidemic nature of these cases implicates, an arthropod that normally transmits the disease from one animal or bird to another and that may infect man entirely by chance. The près-
96 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

kLL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE - MALAYA

-^ FEMALES ;^^

\M/iL£S HAVE BUSHy ANTENNAE AND DO NOT BITE FIG. 62

LONG PALPS ON

ANOPHELES

PROBOSCIS

PATTERN ON FEMUR & TIBIA

PATTERN ON PALP

Ï

ENTIRELY DARK

C PATTERN ON FORE TARSUS PATTERN ON HIND TIBI/ Ï

PALE BAND ABOUT AS LONG

AS WIDE ON FIRST TWO

OR MORE BASAL

SEGMENTS

ENTIRELY DARK

--NO PALE BAND

VEIN 4-^

DARK EXCEPT FOR LIGHT SPOTS ON

BRANCHES

_ ANOPHELES

" UMBROSOS

PROPLEURAL HAIRS ABSENT ANOPHELES

/ SEE ENLARGED DRAWING \ LETIFER

\ FOR LOCATION J

,'\ _ ANOPHELES

~^ BARBIROSTRIS

A NARROW WHITE BANDS

PATTERN ON HIND

TARSAL SEGMENTS 3,4,i 5

PATTERN ON HIND

TARSAL SEGMENTS 3,4. & 5

__ ANOPHELES

NIGERRIMUS

_ ANOPHELES

" S/NE/VS/S

PATTERN ON PROBOSCIS

2 BROAD AND 1 NARROW WHITE

BANDS ALL DARK, OR

J. WHITE TIPPED

SMALL LIGHT SPOT IN BASAL THIRD

ANOPHELES

ZZ MINIMUS

MINIMUS

PATTERN ON HIND TARSUS

SHORT PALPS ON

ALL OTHERS PATTERN ON THORAX

PATTERN ON HIND TIBIA

NO PALE BAND

AEÛES

ALBOPICTUS

A EUES

AEGYPTI

ANOPHELES

SUNÛAICUS

ANOPHELES

MACULATUS

HEAD AND THORAX

WING ^^

¿?Î^ f"'^ ^^^^

FRINGE-'

PROPLEURAL YÍAIR»-'' ^

ADULT MOSQUITO WITH.PARTS LABELED

ence of dengue may complicate the exact identification of

this virus in Malaya.

References: 15S, 251.

Filariasis.—Wuchereria malayi is the most common cause

of filariasis and elephantiasis in Malaya. Filariasis is common along the northern half of the western coastal strip and

in the northwest corner of Malaya but is absent from the

coastal villages. Villagers who live on sandbanks intersecting various ricefield areas are especially prone to infections.

In these areas Mansonia longifalpis is the most important

vector, since it is found in far greater numbers and with

much higher percentages of infection than any other Mansonia species. The larvae of this mosquito are often found

attached to ñoating roots of swamp trees in this area.

References: 165, 215, 315.

Sunda Islands

(Figs. 63 and 64)

The Sunda Islands include the Greater Sundas, consisting

of Sumatra, Borneo, Celebes, and Java, and the Lesser Sundas, consisting of Bali, Lombok, Soembawa, Soemba, Flores,

Timor, and the Solor and Alor Islands. The Sunda Islands

comprise the eastern extension of the Malay Archipelago.

SUMATRA

Physical Features.—Sumatra, about 1,100 miles long and

280 miles wide, lies south and west of Malaya and is fringed

with numerous islets. The west coast is lined with a chain

of volcanic mountains, the Barisans, which divide the narrow western littoral from the swampy, flat, interior plain.

Much of this plain is impenetrable forest, which has high

temperatures and rainfall that ranges from about 75 inches

annually in the east to over 155 inches in the west. The two

rainy seasons are brought by a northwest monsoon from

December to March and a southeast monsoon from May to

October.

References: 121, 301.

Malaria.—The northeast coast of Sumatra is the only area

free of malaria in this island because of the mangrove. All

other coastal districts are highly malarious, partly because

of Anopheles sundaicus. This mosquito is a brackish-water

breeder that transmits malaria efficiently and causes occasional epidemics, especially on the southwest coast. A freshwater form of sundaicus breeds in fishponds in the long
MOSQUITOES OF MEDICAL IMPORTANCE 97

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE — SUNDA ISLANDS

CAREA INCLUDED-. SUMATRA, BORNEO^ CELEBES, JAVA^ AND THE LESSER SUNDAS]

^^ FULL-GROWN LARVAE! FIG. 63

4y.¿^

%•^n ||>J^

REGION INCLUDED ^.^|

r LONG PLEURALS

OF aas GROUP

NO SIPHON ON ANOPHELES

POSTSPIRACULAR HAIR.

SPIRACULAR

APPARATUS

BOTH

BARE

BOTH

HAIRED

SIPHON ON ALL OTHERS

C PALMATE HAIRS ON

3BB^N^4iB SEGMENTS

Ï

(SEE ENLARGED DRAWING)

FULLY DEVELOPED

ON BOTH

r—mp—^

'—^m—'

OUTER AT LEAST

35 BRANCHED

NNER CLOSE TOGETHER

PROTHORACIC

HAIR 1

6 TO 14

BRANCHED

FROM BASE

SPIRACULAR APPARATUS

/NOT

_ ANOPHELES

"" BARBIROSTRIS

OUTER BARE OR

WITH FINE SIDE HAIRS

INNE« FAR APART

BARE OR

BRANCHED

AT TIP

r PROTHORACIC

HAIR 1

W/ITH

LARGE

TUBERCLE

WITHOUT

TUBERCLE

"eTH SEGMENT - ANOPHELES

SIX TO ELEVEN —

BRANCHED SINENSIS

_ ANOPHELES

"~ LEUCOSPHYRUS

_ ANOPHELES

— KOCHI

NIGERRIMUS

UNDEVELOPED ON BOTH

TIP OF ANTENNA

HAIR WITH I OR 2 STOUT, SHORT AND

+ 2 TO 4

BRANCHED

HIND CLYPEALS 3Bg SEGMENT

LATERAL HAIR

13 TO 20 BRANCHED _ ANOPHELES

AND

HAIR 0

3 OR 4 BRANCHED

LET IF ER

_ ANOPHELES

"" UMBROSUS

PALMATE HAIRS

UNDEVELOPED

/^LONG PLEURALS

OF Z^ GROUP

BOTH

BARE

PALMATE HAIRS

ON lai SEGMENT

Ç POSTS PIRACULARS

-^^.

7 OR a

BRANCHED

OUTER CLYPEALS

ANOPHELES

= SUBPICTUS

SUBPICTUS

_ ANOPHELES

"- SUNDA/CUS

1 I y ^J^J^^n <^l-TrilAI-i» ANOPH

Vil/y II Í-UDLC

<My + '^VS"ÍSNG = -•^'> ^

I /H H\ CANN

' ^ i RATE

ANOPHELES

LUDLOW I

LITORALIS

LUDLOWI IMPORTANT ; LARVAE

CANNOT BE SEPARATED

ABDOMINAL PALMATE S

NOT \

BARE OR

+ LIGHTLY ^>-\

FRAVEQ

COMB SCALES

IN ONE

REGULAR ROW

(NOT ON A PLATE)

COMB SCALES

_ ANOPHELES

~ MACULATUS

_ AEDES

~ AEGYPTI

AEDES

ALBOPtCTUS
98 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE SUN DA ISLANDS

IMPORTANT

s MUST HAVE

. r&as LÍ3TE0

fOR THAT SPECIES

[AREA INCLUDED: SUMATRA, BORNEO, CELEBES, JAVA^ AND THE LESSER SUNDASj

FEMALES^ FIG. 64

LONG PALPS ON

ANOPHELES

ANTENNAE

.PROBOSCIS

SHORT PALPS ON

ALL OTHERS

•^ +4. '".M^RmJ.^A^ - ANOPHELES IN FRINGE AT _ ^/^^^^./^

_ ANOPHELES

— NIGERRIMUS

LIGHT SPOT IN

, FRINGE AT TIP _ ANOPHELES

"*" OF VEIN 3 " BARBIROSTRIS

" -^ VERY NARROW

PATTERN ON HIND TIBIA

NO PALE « ANOPHELES

UMBROSUS

PROPLEURAL HAIRS ABSENT _ ANOPHELES

PATTERN ON

THORAX

T

PATTERN ON

HIND TIBIA

~ AEDES

AEGyPTI

AEDES

ALBOPICTUS

ADULT MOSQUITO WITH PARTS LABELED
MOSQUITOES OF MEDICAL IMPORTANCE 99

mountain valleys of the Barisans near Lake Toba and occasionally in flooded ricefields after harvest. Its medical

importance is not exactly known.

Anopheles kochi is abundant in Sumatra. Natural infections in this species range from 0 to as high as 11.5 percent

but are generally low.

Although Anopheles leucosphyrus (subspecies not known)

has been found with low natural infection rates in east and

south Sumatra, this mosquito is neither common nor important except in "Soendetar" in western Sumatra, where it

is present in large numbers in December and January and is

definitely a vector.

Anopheles Tmaculatus and A. minimus (subspecies not

known) transmit malaria in the foothills, but the disease in

the mountains has not been thoroughly studied. The former

mosquito appears as high as 3,200 feet. It is usually found

with low infection rates. A. minimus breeds at the edges

of foothill streams and is of little importance in the island.

Anopheles harhirostris is present throughout Sumatra.

Its natural infection rates are always low, and it is only occasionally important as a vector there.

Anopheles aconitus is abundant in ricefields but is a dangerous vector only at Tapanuli on the east coast and in ''Djihara" in the south.

Anopheles umhrosus and A. letifer are vectors in Malaya

and may support a certain amount of malaria in Sumatra,

having been found with low infection rates.

The role of Anopheles subpictus suhpictus as a vector is

nol known with certainty, since it commonly appears with

other important species, especially sundaicus. When suhpictus is involved in an epidemic, it is always of secondary

importance, becoming infected later than the principal vector. It almost always has low infection rates.

Anopheles sinensis and A. nigerrimus have been recorded

from the island, but records of their medical importance have

been confused with each other to such an extent that their

respective roles are not known with certainty. Both have

been found naturally infected in ricefields of the alluvial

plains, but infection rates have been rather low.

The distribution of Anopheles ludlowi and its subspecies

is not known with certainty in Sumatra, nor has its role in

malaria transmission been determined.

References: 121, 122, 284, 301.

BORNEO

Physical Features.—Borneo, which is the largest of the

Greater Sundas, lies north of Java and Bali and southwest

of the Philippines. Centered in the island are densely forested mountains rising in peaks to 13,000 feet in the north,

and along the south and southwest coasts are extensive

swamps. A prolonged monsoon season brings rainfall exceeding 100 inches per year, but in a very few localities there

is a sharp distinction between wet and dry seasons. Seasonal

variation in temperature is negligible.

References: 121, 301.

Malaria.—Anopheles leucosphyrus Ixüabacensis is present

wherever there is jungle shade. It is the most widely distributed of all anophelines and has been found naturally

infected in every part of the island.

Both Anopheles umhrosus and A, letifer are widely distributed and naturally infected in North Borneo, Sarawak,

and Brunei and are believed to contribute greatly to the

maintenance of endemic malaria.

The distribution of Anopheles sundaicus in Borneo is not

well known. It is believed to be present at least as far north

as Miri in Sarawak and has been found in many malaria-free

areas in Borneo. Its coastal distribution is limited by mangrove swamps, which it avoids. It is believed to be more

important in occasional epidemics than in the maintenance of

widespread endemic malaria.

Anopheles suhpictus subpictus has been reported from two

localities on the southeast coast of Borneo, but further information about its distribution or role as a vector is not

available. Although Anopheles sinensis and A, nigerrirmis

are widely distributed, there is little evidence of these species

transmitting malaria in Borneo. Natural infections have

been found in both species in the island, but more thorough

investigations are needed regarding their relation to disease.

Anopheles maculatus^ appearing chiefly in northern

Borneo, Sarawak, Brunei, and Dutch Borneo, is not regarded

as a vector of malaria there. It has often been found in malaria-free areas, is rarely infected in nature, and has a

marked preference for animal blood. In the other Sunda

Islands this species is regarded as one of the most important

anophelines.

Anopheles harhirostris is probably unimportant in Borneo,

although it is very common and widely distributed.

Anopheles minimus (subspecies not known), A, aconitus^

and A. kochi are present in Borneo but apparently do not

transmit malaria there.

The distribution of Anopheles ludlowi and its importance

as a vector in Borneo are not known with certainty.

Anopheles haezai^ an anthropophilic species occasionally

found with natural infections, is a suspected vector in the

Miri-Lutong and Kuala areas. It is not included in the keys

because it is probably of little importance.

References: m, 121,

CELEBES

k, 301.

Physical Features.—Celebes, about 90 miles east of

Borneo, is composed of a central area with four large peninsulas. Mountainous land in the south-central part of the

island rises to over 11,000 feet. The highlands contain fertile

valleys, rich grazing land, and lakes, whereas the interior is

densely forested. A northeast monsoon provides heavy rainfall, the northern part of the island receiving about 100 inches

and the southern part about 150 inches a year. The climate,

in general, is hot and humid, usually between 70° and 90° F.

References: 121, 301.

Malaria.—Anopheles sundaicus^ the most important vector

in Celebes, transmits malaria in a highly endemic form along

the coast of the southern peninsula and the adjacent islands,

on the part of the southern coast facing Kabaena and Muna,

and in "Vuton," Salayar, and Djampea. This species is

coastal, breeding in brackish fishponds, pools along the coast,

and silted outlets.

Anopheles suhpictus suhpictus is also a coastal mosquito,

but it is less selective than sundaicus^ breeding in a large

variety of fresh- and brackish-water sites. Natural infection

rates are low, but it is the most prevalent anopheline on the

west and south coasts of the southern peninsula, and because

of its abundance it can be important in causing epidemics.

Anopheles ludlowi torakala^ designated as ludlowi in the

keys, is present near the coast and inland from the base to

about the center of the southern peninsula, and along the

western coast of the southeast peninsula. With infection

rates of 2 to 3 percent, this species appears to be responsible

for much of the inland malaria in Celebes. It is regarded as

second only to sundaicus in importance in this area.

Anopheles harhirostris has been found in almost all localities in southern Celebes where collections have been made.

Because this species is found in breeding places closely associated with man and is markedly anthropophilic, it is important where it appears in large numbers. It is regarded as

the third most important vector in southern Celebes.

Anopheles nigerrimus^ like the preceding species, is found

throughout southern Celebes and breeds in ricefields and
100 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

fishponds close to man. When this species appears in large

numbers, it can cause epidemics, but even when less

abundant, it maintains a certain amount of endemic malaria.

Anopheles leucosphyrus hackeri appears near the southern

coast of Celebes. The importance of this mosquito is not

well known.

Anopheles ndnimMs ßavirostris^ or a form similar to it, is

common in the interior of northern Celebes, where it causes

epidemics or supports endemic malaria. In southern Celebes

it is too rare to be of much importance, although it has been

found with high natural infection rates. Clarification of the

^tditwB oífiœvirOS tris in Celebes must await further field study.

Five additional Anopheles species—aconitus^ maculattcs^

kochi, umhrosus^ and sinensis—are included in the keys, because they appear in Celebes and have definitely been implicated as vectors elsewhere. They probably are not important

in this island because of their scarcity and lack of natural

infections.

References : 121,161, 301.

JAVA AND THE LESSER SUNDAS

Physical Features.—The long narrow island of Java lies

east and south of Sumatra. It contains almost 40 million

people, less than 4 percent of whom live in cities, yet culturally, politically, and economically it is the most important

of all the Sunda Islands. It has an east-west volcanic mountain range, with peaks as high as 12,000 feet. Western Java

does not have a definite rainy season, but in eastern Java a

west monsoon brings a distinct wet period, especially in the

lowlands, from November to March.

The entire island chain of the Lesser Sundas has distinct

mountain ranges. The climate is similar to that of Java.

The highest temperatures and least cloudiness occur in

October and November.

References : i^i, ^öi.

Malaria.—Anopheles sundaicus is responsible for extensive

endemic malaria and occasional severe epidemics along the

northern and southern coasts of Java and in coastal areas of

the Lesser Sundas. Brackish-water lagoons are the principal

breeding places in these localities. Very severe epidemics

have been reported at Tanjungpriok in western Java. A,

sundaicm has also been found in collections of fresh water

in the vicinity of Semarang, Java, in brackish-river backwaters in the eastern part of the island, and in fresh-water

fishponds in Bali. This is not the same fresh-water form

found in Sumatra. It more closely resembles the salt-water

sundaicus in British India and may be important in

transmission.

Anopheles aconitus is almost as important as sundaicus in

the transmission of malaria in ricegrowing areas of west

Java, although it varies considerably in importance

elsewhere.

Anopheles maculatus^ a common and important vector near

the tea plantations of Java, invades cleared areas where

jungle has been cut away from banks of small streams. It

is regarded as a vector of considerable importance in the

Lesser Sundas.

Anopheles leucosphyrus (subspecies not known) may be

an important transmitter in Java, but its exact role has not

been adequately determined.

In an outbreak of malaria at Karangbinangun, Java,

Anopheles nigerrimus was considered to be the principal vector. Natural infection rates in this mosquito are usually

low, so that the species must be present in large numbers to

be dangerous. Neither nigerrimus nor Anopheles sinensis

appears in Java or the Lesser Sundas.

Anopheles umbrosus and A. letifer are present and have

been found with low infection rates. They are important in

other parts of their range but have been inadequately studied

in this area.

Anopheles subpictus subpictus is more common in Java

and the Lesser Sundas than elsewhere. Its role in endemic

and epidemic malaria in this area is not well understood,

because it is found with other vectors, but it is undoubtedly

a vector of some importance.

A form similar to Anopheles minimus -ßavirostris has been

recorded as a vector in the area, but its actual importance

is not known with certainty. Since -ßavirostris is easily confused with aconitus^ the records of these two mosquitoes may

have been confused.

Although both Anopheles kochi and A, harhirostris are

present in Java and the Lesser Sundas, they are important

only in restricted areas.

The presence of Anopheles ludlowi in Java and the Lesser

Sundas is doubtful.

References : 121, 284, SOI.

Most of the malaria cases throughout the Sunda Islands

are caused by either Plasmodiumi vivax or P. falciparum.

The distribution of Plasmodmm maluriae is sporadic and is

apparently limited to Sumatra and Java.

Anopheles Species in the Sunda Islands

"^aconitu^

aitkenii aitkenii

aitkenii hengalensis

aitkenii borneensis

aitkenii stantoni

aitkenii treacheri

albotaeniatus

annandalei annandalei

annandalei djajasanensis

\annularis

argyropus

baezai

\bancrofti bañerofti

bancrofti barbiventris

"^barbirostris

barbumbrosus

brevipalpis

crawfordi

errabundus

gigas crockeri

gigas danaubento

gigas gigas

gigas oedjalikalahensis

gigas sumatrana

hu/nteri

indiensis

insulaeflonmi

jamesii

karwari

^kochi

lesteri

Hetifer

Heucosphyrus (subspecies

not known)

Heucosphyrus balabacensis

See footnote 8, p. 63.

Yellow Fever.—This disease is not present in the Sunda

Islands.

Reference : 256.

Dengue.—Aedes aegypti and A, albopictus are found

throughout the Sunda Islands. Both species are proved

vectors of dengue. The exact distribution of the disease

within the Sundas is not known, although these islands comprise one of its main endemic centers. A, albopictus females,

Heucosphyrus hackeri

litoralis

ludlowi -ßavescens

Hudlowi torakala

'^maculatus

^minimus (subspecies not

known)

"^minimus flavirostris

"^nigerrimus

pallidus

palmatus

parangensis

peditaeniatus

\philippinensis

pseudobarbirostris

rarmayi

riparus macarthuri

roperi

saungi

schueffneri

similissimus

^sinensis

stookesi

subpictus indefinitU;S

subpictus mdlayensis

"^subpictus subpictus

"^sundaicus

tessellatus kalawara

\tesseïlatus orientalis

tessellatus tessellatu^s

"^unííbrosus

vagus albino

vagus Ivmosus

vagus vagus

vanus
MOSQUITOES OF MEDICAL IMPORTANCE 101

which were infected with dengue by feeding on infected

persons in Sumatra, have experimentally transmitted the disease to persons in Holland known to be previously uninfected.

Thus a definite link has been demonstrated between the disease and this species in Sumatra.

References: 122, 256, S12.

Encephalitis.—The virus of Japanese encephalitis has been

reported from Java and the eastern coastal areas of Sumatra,

but its distribution and intensity are not well known. There

have been no confirmed human cases in these two islands.

Human cases have been reported from British North Borneo.

Little is known of the epidemiology of this disease in the

Sunda Islands.

Reference : 257.

Filariasis.—Wuchereria malayi probably is found

throughout the Sundas. Wuehereria hancrofti^ sporadic in

the western part of the island chain, is more common on the

eastern end. Both parasite species have been recorded from

the northern end of Sumatra ; malayi is common and widespread and hanerofti sporadic in the southern third. In

Timor, southeast Borneo, and most of Celebes malayi predominates. TF. hanerofti is sporadic in the last two islands.

The "Serajoe" Delta of south-central Java is an endemic

area for malayi^ but elsewhere in this island filariasis is

sporadic. The disease apparently does not occur in Bali,

Lombok, and Soembawa, but in all the islands of the eastern Lesser Sundas it is highly endemic, with both parasite

species present.

Little is actually known of the relative importance of the

vectors. Mansonia uniformis^ M. indiana^ M, longipalpis^

and M. annulifera are the important vectors of malayi m the

Sundas. Anopheles harhirostris is an efficient vector of

malayi in Java and Celebes, and A, nigerrimus is an important vector of that parasite at "Martapoera," Java. Culex

pipiens quinquéfasciatiis is believed to be the only important

vector of hanerofti in the Sundas.

Reference: 121.

Philippine Islands

(Figs. 65 and 66)

Physical Features.—The Philippines consist of about

7,000 islands, which are the higher portions of partly submerged volcanic mountains. Over 4,000 of these are unnamed, and only 463 are over 1 square mile in area. The

larger islands have extensive ricefields ; sugarcane, tobacco,

and pineapple are other important crops.

The lowlands are warm and humid, with mean daily

temperatures of about 80° F. and up to 250 inches of rain

annually, but the mountainous areas in the larger islands

have a rather temperate climate. A northeast monsoon

brings rain to the eastern coasts of the northern islands,

and a southwest monsoon causes a rainy season on their west

coasts. The southernmost islands have an even distribution

of rain throughout the year.

Reference: SOI.

Malaria.—Little is known about the distribution of the

Plasmodiurm species in this region.

This disease has never been so important in the Philippines

as in Asia, presumably because of the habits of the principal

vector Anopheles minimus ßavirostris. This mosquito breeds

along the banks of fresh-water streams and other collections

of water in foothill areas. Endemic malaria is more or

less confined to areas roughly corresponding to the distribution of ßavirostris. This mosquito has a pronounced preference for the blood of livestock. It rarely rests inside houses

for any length of time. The disease is most prevalent at the

beginning and end of the rainy season, when streams are

at a favorable height to support mosquito breeding. Flat

coastal areas are relatively free of malaria because of the

rarity of vectors, especially Anopheles sundaicu^, and in

areas other than the foothills malaria is rarely found.

Two close relatives of -ßavirostris^ Anopheles mangyanus

and A, flUpinae^ also are present in the Philippines and

doubtlessly are important in malaria transmission. The

former, like ßavirostris^ breeds in clear, slowly running water

in foothill areas at altitudes not exceeding 2,000 feet. The

larvae are found in both shaded and sunlit locations. The

importance of this species in malaria transmission is not yet

entirely clear, since little is known of the biting habits of

the adults and since early records of infections may be

confused with those of ßavirostris.

The other species, jilipinae^ breeds in either sunny or

shaded, clear or muddy water in rivers, flowing water in irrigation ditches, and in pools and lakes. This mosquito may

be a vector of malaria in the Philippines, where it has been

found in large numbers in northern Luzon in the presence

of abundant malaria. It is the least common of the Anopheles species so far mentioned.

Neither Anopheles maculatus nor A, leucosphyrus is

known to transmit malaria in the Philippines. The former

breeds in a variety of places at altitudes up to 5,000 feet and

is included in the keys because of its potential importance

in these islands, since it is a dangerous vector in Malaya and

certain parts of Indonesia. However, epidemiological investigations indicate that not all foothill malaria is borne

by -ßavirostris and that maculatus is suspected in areas

where ßavirostris density is low. Evidence points to maculatus as a vector in certain lowland areas as well.

The larvae of leucosphyrus are often found in temporary

collections of water, such as wheel ruts and footprints in open

shade. The females bite man readily. Various forms of this

species are proved vectors in parts of Assam, Burma, and

North Borneo (see p. 99), but they show no evidence of

transmitting malaria in the Philippines.

References : 31, |8, ^9, 61,115,121,122, 296, 308, 310.

Anopheles Species in the Philippines

acacz

aitkenii aitkenii

aitkenii hengalensis

\annularis

haezai

halahacensis haisasi

halerensis

^harhirostris

cristatus

ejercitoi

'^filipinae

gigas formosus

insulaeflorurn

karwari

\kochi

kolamhuganensis

lesteri

\leucosphyrus halahacensis

\leuco8phyrus leucosphyrus

leucosphyrus riparis

See footnote 6, p. 10.

Yellow Fever.—As in other parts of the Orient, neither

urban nor jungle yellow fever is known to be endemic in

the Philippines.

References : 256, 310.

Dengue.—The classical work of Siler and others {307y

has confirmed the direct role played by Aedes aegypti m the

lindesayi henguetensis

litoralis

\hcdlowi

^maculatus

'^mangyanus

'^mdnimu^ ßavirostris

\minimus minimus

\nigerrimus

parangensis

philippinensis

pseudoharhirostris

pseudosinensis

samarensis

\sinensis

suhpictus indefinitus

]suhpÍGtus suhpictus

\sumdaicus

f tessellatus

vagus

^ ItaUc numbers in parentheses refer to References, p. 143.
102

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

MOSQUITOES OF MEDICAL IMPORTANCE PHILIPPINE ISLANDS

-—^^ FULL-GROWN LARVAE ^M^ _

ABDOMINAL PLATES

HAIR "O" NOT ON PLATE

SIPHON ABSENT

_ ANOPHELES

"~ MINIMUS FLAVIROSTRIS

-_ ANOPHELES

" MANGYANUS

FIG. 65

HEAD OF ANOPHELES V.tKR\lk

ANTENNAL

HAIR INNER

/ CLYPEALS

/ OUTER I HIND

CLYPEALS I CLYPEALS

"^

^ANTENNA

ANOPHELES

MACULATUS

ANOPHELES

LEUCOSPHYRUS

LEUCOSPHyPUS AND

LEUœSPHYRUS RIPARIS

_ AEDES

— ALBOPICTUS, PAULLUSly AND SCUTELLARIS

/ONLY ALBOPICTUS IMPORTANT: LARVAE \

\ CANNOT BE SEPARATED /

ANOPHELES V.^^Mf^

PLEURAL BRISTLES

, ,Y ii^VABDOMINAL

i<^.H-^^ SEGMENTS

2 AND 3

5Í_r:'^^V*^BD0MINAL

JpiS^ ^ PLATES

^ \_ ^->J^BDOMINAL

transmission of dengue in the Philippines. This disease,

which has centered principally in Luzon, has occurred near

military installations. A seasonal peak "of dengue cases

generally occurs between April and November, its height

varying from year to year. Aedes alhopictus is an important

vector of dengue in the Philippines.

References : 256, SOI, SIO.

Encephalitis.—The viruses causing Japanese and St.

Louis encephalitis have been reported from the Philippines.

Antibodies to both viruses have been found in human sera.

Nothing is known about their vectors in this region.

Reference : 257.

Filariasis.—This disease has existed in the Philippines

since 1901. It is endemic in areas where abaca is grown and

in at least two other localities. Higher infection ,rates in

males than in females indicate that the males are more exposed to the disease because of their outdoor activity in the

cultivation of abaca. All known infections are caused by

the periodic form of Wuchereria hancrofti and are probably

borne by Aedes poicilius. No laboratory or field investigations have conclusively proved that this species is the important vector, but epidemiological observations implicate it

far more than any other vector.

Reference : 29S.

AUSTRALASIA

(Figs. 67 and 68)

Australasia has been divided into the following parts:

(1) The Moluccas, New Guinea, and Melanesia; (2) Australia, Tasmania, and New Zealand; (3) Micronesia; and

(4) Polynesia.

The Moluccas, New Guinea, and Melanesia

Physical Features.—The Moluccas are volcanic and coral

islands lying between Celebes and New Guinea. There is

no well-defined dry season, and the rainfall averages 75 to

100 inches per year.

New Guinea, the world's second largest island, lies between

the Moluccas and Melanesia. It has a series of mountain

ranges with peaks of 16,000 feet. Tropical jungle covers

much of the interior. A west monsoon brings rain and high

temperatures to southeast Dutch New Guinea, whereas the
MOSQUITOES OF MEDICAL IMPORTANCE 103

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE- PHILIPPINE ISLANDS

-^^^ FEMALES ^ii

[MALES HAVE BUSHY ANTENNAE ( íÉ^^^ ) AND DO NOT B/TE"] FIG. 66

LONG PALPS ON

ANOPHELES

.PROBOSCIS

PATTERN ON HIND FEMUR PATTERN ON PALP

PATTERN ON FORE TARSUS

ENTIRELY DARK

ANOPHELES

MINIMUS F LAVI ROST RIS

ANOPHELES

MANGYANUS

PATTERN ON HIND TARSUS

WITH NARROW LIGHT BANDS _^^ __ ANOPHELES

^ FIL IPINAE

_ ANOPHELES

~~ MACULATUS

PATTERN ON HIND TARSUS

_ ANOPHELES

— LEUCOSPHYRUS LEUCOSPHYRUS

SHORT PALPS ON

ALL OTHERS

PATTERN ON TOP PATTERN ON SIDE

OF THORAX OF THORAX

WHITE SCALES FORMING

PATCHES,NOT STRIPES

PATTERN ON HIND TARSUS

. _>. AEDES

~ ALBOPICTUS

_ AEDES

"~ AEGYPTI

east monsoon is drier and cooler. The interior mountains

may have an annual rainfall of 250 to 325 inches.

Melanesia, situated east of New Guinea and south of the

Equator, includes the island groups of Fiji, New Caledonia,

Loyalty, New Hebrides, Solomons, Santa Cruz, and Admiralties, and the Louisiade and Bismarck Archipelagoes.

The larger of these islands are volcanic and the smaller

coral.

Reference: SOI.

Malaria.—Most of the Moluccan coast and the entire north

coast of New Guinea have high rates of endemic malaria,

whereas areas on the south coast of New Guinea are practically free of this disease. Malaria occurs at altitudes of

3,500 to 4,000 feet in the mountainous interiors. It is widespread and severe throughout the islands of Melanesia because of the low standard of living of the native populations.

In almost all this area Plasmodium falciparum and P.

vivaw are important, one predominating in one locality and

the other in a closely adjoining one. P. malariae is rarely if

ever found.

Malaria in the South Pacific is generally understood to be

bounded by longitude 170° E. and latitude 20° S., the junction of these two lines lying just north of Aneityum, the

southernmost island of the New Hebrides group. This location corresponds exactly to the distribution in the New Hebrides of Anopheles farwuti. That species and Anopheles

punetulatus are the most common and dangerous vectors in

this area. Of these two species, farauti has the wider distribution. Its range includes the Moluccas and extends eastward through New Guinea, the Admiralty Islands, the Bisniarck Archipelago, the Solomon Islands, and the New Hebrides. Neither malaria nor any Anopheles species are present

in New Caledonia. In dry weather farauti breeds in all kinds

of collections of water close to man, and during the rainy

season it breeds readily in small manmade depressions exposed to sunlight. The larvae of this species are commonly

found in brackish water, and for this reason the adults may

be abundant in coastal areas. A, punetulatus has much the

same range as farauti^ but it is not found east or south of the

Solomons, leaving malaria transmission in the New Hebrides

solely to farauti. Breeding in all sorts of small collections

of sunlit ground and rainwater, punetulatus is more characteristically a species of inland and elevated areas than farauti.

Since these two species breed so readily in manmade habitats

and are such effective hosts for malaria parasites, they maintain malaria throughout the entire island chain.

Anopheles koliensis^ a close relative of both farauti and

punetulatus^ is believed to be an important vector on the

northeast coast of Guadalcanal and in certain parts of New

Guinea.
104 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

IMPORTANT

A SPECIMEN MUST HAVE

ALL CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE — AUSTRALASIA

OREA INCLUDED; MOLUCCAS, NEW GUINEA, MELANESIA, AUSTRALIA, TASMANIA, NEW ZEALAND, MICRONESIA, AND POLYNESIA]

L FULL-GROWN LARVAE J FIG. 67

OUTER CLYPEALS

BARE, OR WITH

VERY SHORT

SIDE HAIRS

WITH LONG

SIDE HAIRS,

OR BRUSHSIPHON ON ALL OTHERS

LONG PLEURALS

OF 3RD GROUP

PALMATE HAIR

ON 4TH SEGMENT

MENT /

OR MORE 1 / /

LONG AS à d y y

PROTHORACIC HAIRS I AND 2

PALMATE HAIR

ON 1ST SEGMENT

HIND CLYPEALS

PLACED FAR

BACK OF

INNER

CLYPEALS

ANOPHELES

SUBPIcrus

AND SUNDAICUS

ONLY SUBPICTUS IMPORTANT; LARVAE CANNOT

BE SEPARATED WITH

CERTAINTY ,

TUBERCLES LARGE

AND CLOSE TO- j^ ^^>

GETHER OR FUSED /^i^riPi-m

ANOPHELES

FARAUTI AND KOLÍENSIS

PALMATE HAIR

ON 2ND SEGMENT

INNER CLYPEALS

^-^

+ CLUB^^

^~^7^N

ANOPHELES

PUNCTULATUS

ANOPHELES

BANCROFT! AND

PSEUDOBARBIROS TRIS

[ONLY BANCROFT! IMPOR-\

TANT; LARVAE CANNOT

BE SEPARATED /

INNER CLYPEALS

BARE, OR WITH

VERY SHORT

SIDE HAIRS

_ ANOPHELES

"~ ANNULIPES

BODY HAIRS

ONLY I

TUFT

PER SIDE

COMB SCALES

IN ONE

.REGULAR ROW

(NOT ON A PLATE)

NOT /^^'^

NOT ._

-^

COMB SCALES

ANOPHELES LARVA

VNTENNAL

HAIR INNER

/ CLYPEALS

/ OUTER

CLYPEALS I CLYPEALS

PROTHORACIC /

HAIRS H.2 1^ PLEURAL BRISTLES

_ AEDES

~ AEGYPTI

AEDES

_ ALBOPICTUS AND

— SCUTELLARIS GROUP

Neither the importance nor the distribution, especially in

Melanesia, of Anopheles hancrofti is well known. It is generally agreed that this species definitely is present in the

Moluccas and along the southern shore of New Guinea. In

Batu Gajah in Malaya hancrofti has been an important vector. Undoubtedly it maintains some malaria throughout at

least part of its range.

Anopheles annulipes appears sporadically in New Guinea,

but it has never been proved to be a vector there. In the Port

Moresby area it is at times the dominant species, and although

its natural infection rates are generally low, it has been shown

to transmit malaria in the laboratory as effectively as farautL

It may be a species of some importance in New Guinea when

conditions are favorable.

Anopheles suhpictus is probably the least important malaria vector in this area. It is encountered usually in small

numbers in the Moluccas and New Guinea. Although it has

been found naturally infected in the vicinity of Lalapipi in

New Guinea, its small numbers there indicate that it is of

little or no consequence in malaria transmission. Its presence

in Melanesia is in doubt.

References : 62,121,122,170, , 228, 25Jf,

Yellow Fever.—This disease is not present in the Moluccas, New Guinea, and Melanesia.

Reference : 256.

Dengue.—This very important disease is spread throughout this area. It is transmitted by Aedes aegypti. Aedes

alhopictus is not believed to appear west of the Lesser Sunda

Islands. A so-called jungle type of dengue occurs over vast

areas in northern New Guinea. Extensive experiments in

which Aedes scutellaris fed on persons with this disease in

New Guinea and afterward transmitted the disease to susceptible volunteers in southern Australia demonstrated that

this species was the vector. In the New Hebrides scutellaris

is epidemiologically suspected as the vector of dengue in

areas where aegyptiis absent.

References : 102,121', 245, 256.

Encephalitis.—No information is available.

Filariasis.—This disease is endemic on the entire coast

of Vogelkop in New Guinea and its associated islands, in the

vicinity of Hollandia on the north-central coast, and in the

eastern third of the island from ''Lambringi" in the north to

Daru on the southern coast. It has not been recorded in the

interior. It is also endemic in small areas in New Britain,

New Ireland, and the Solomons. Guadalcanal and San Cris-
IMPORTANT

A SPECIMEN MUST HAVE

ALU CHARACTERS LISTED

FOR THAT SPECIES

MOSQUITOES OF MEDICAL IMPORTANCE

MOSQUITOES OF MEDICAL IMPORTANCE - AUSTRALASIA

[AREA INCLUDED: MOLUCCAS, NEW GUINEA^ MELANESIA, AUSTRALIA^ TASMANIA, NEW ZEALAND, MICRONESIA, AND POLYNESIAJ

--^^m FEMALES ^

105

[MALES HAVE BUSHY ANTENNAE AND DO NOT BITE]

FIG. 68

PATTERN ON PROBOSCIS

ANOPHELES

FARAUTI AND KOLIENSIS

( BOTH IMPORTANT )

_ ANOPHELES

~ ANNULIPES

tobal are the most seriously aifected of the Solomons. Almost

the entire northern two-thirds of the New Hebrides is heavily

infected.

Throughoutthese areas Wuchereria hancrofti^ with few exceptions, is the sole parasite and is transmitted principally

by Anopheles farauti and A, hancroftL A, farauti is found

throughout the entire island chain and is responsible for the

greatest number of cases. A, hancrofti is implicated as the

important vector in the Digoel Kiver area in southern New

Guinea, where elephantiasis is abundant. Anopheles pwrwtulatus is a potential secondary vector at HoUandia and in

Guadalcanal, and A, koliensis probably contributes to filariasis in the latter island. Culex pipiens quinquefasciatus is a

secondary vector of bancroftian filariasis at the eastern end

of New Guinea. Light infections of the nonperiodic form of

TT. hancrofti occur at the northern end of New Caledonia and

in Ouvea in the Loyalty Islands, which represent the westernmost extension of nonperiodic filariasis in the Pacific. The

parasite there is transmitted solely, as far as is known, by

Aedes mgilax.

Very seldom has Wuchereria malayi been seen in New

Guinea and Melanesia. It is believed to have been introduced

into the Digoel River area in southern New Guinea, and it

has been found in the New Hebrides, where Mansonia

xanthogaster was uninfected in nature.

References : 191,273,287.

Australia, Tasmania, and New Zealand

Physical Features.—Australia, which is approximately

the size of the United States, has a west-central part consisting of a dry, hot desert plateau 1,000 to 2,000 feet high that

rises abruptly from the west coast and extends east to Cape

York Peninsula, a jungle-covered northern extension of the

continent. The Great Dividing Range has 7,500-foot peaks

and borders nearly all the east and south coasts to Melbourne.

It encloses a basin that is bounded at the center of the continent by the eastern edge of the desert plateau. This basin

is too dry to support a dense population of either people or

mosquitoes. Most of the human population is concentrated

along the east and south slopes of the Great Dividing Range.

Tasmania is a small island lying south of Australia. Geologically it is a part of that continent. It has a high central

plateau, with alpine lakes and good farming lands.

New Zealand, about 1,200 miles southeast of Australia, is

a mountainous island, which has volcanoes of about 12,300

feet and extensive farming and stockraising areas.

Reference : ^öi.
106 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

Malaria.—Malarious areas are centered in the Cape York

Peninsula and in the northern parts of the Northern Territory and of Western Australia. The southern limit of the

disease is usually at about latitude 19*" S. in Queensland and

West Australia and latitude 17° S. in the Northern Territory.

Epidemics occasionally occur in the vicinity of Cairns at the

eastern base of Cape York Peninsula. This area receives

rainfall ranging from 100 to 160 inches per year.

Plasmodium vwaoo and P. falcipoirum are about equally

important in causing the malaria in most of this area. However, on Cape York Peninsula falcipœnmi is the predominant

parasite.

By far the most important Anopheles species in northern

Australia is farauti^ which, as in New Guinea and Melanesia,

breeds in manmade depressions and is always closely associated with man.

Anopheles pmwtmLatus is apparently absent from the Australian mainland. South of latitude 17° S. malaria occurs

only rarely and is restricted to short-lived and narrowly localized outbreaks.

Anopheles annvZipes may be the only vector that maintains this sporadic malaria in southern Australia and in

Tasmania. Its role in transmission has never been proved

in the field. Neither malaria nor any Anopheles species have

been reported from New Zealand.

References : 67,128,354.

Yellow Fever.—This disease is not present in Australia,

Tasmania, and New Zealand.

Reference : 256.

Dengue.—Endemic dengue occurs along the northern and

eastern coasts of Australia, where it is transmitted principally by Aedes aegyptL Aedes albopictus is not found in

this area.

References : 121,256.

Encephalitis.—Japanese encephalitis is thought to occur

in Australia, but further studies are needed to confirm its

presence. Murray Valley encephalitis is present. For further information, see page 4.

References : 246, 257.

Filariasis.—The distribution of filariasis in Australia is

confined to a narrow coastal strip running from the northern

rivers of New South Wales to Cape York, where there is a

fairly high incidence. This strip also includes an area bordering the southwest corner of the Gulf of Carpentaria. In

the northern parts of Queensland filariasis is much more prevalent than farther south. The Northern Territory and the

northwest part of Western Australia are weak centers of infection. The only parasite found in these areas is Wuchereria bancrofti, which everywhere shows well-marked periodicity and is transmitted by Culex pipiens quinquéfasciatus.

Filariasis is not present in Tasmania or New Zealand.

References : ^7, 9S, 163.

Micronesia

Physical Features.—Micronesia consists of the island

groups of the Marianas, Carolines, Marshalls, and Gilberts.

The Marianas and a few of the Carolines are volcanic; the

others are coral atolls, some of which barely rise above the

sea. Temperatures remain fairly constant, and rainfall

ranges from 60 to 180 inches a year. The larger islands of

Micronesia have dense vegetation.

References: 64, 301.

Malaria.—This disease is not present in Micronesia.

References : 61, 224.

Yellow Fever.—This disease is not present in Micronesia.

Reference : 256.

Dengue.—Eeported from the Marshalls as early as 1906,

dengue is perhaps the most important disease in these islands

and in the Carolines, Marianas, and Gilberts as well. Several large epidemics have occurred in Saipan, Tinian, Truk,

Ponape, and the Palau group. Both Aedes aegypti and A,

albopictus were involved in the first two places and aegypti

alone in the last three. In Guam a large epidemic in 1944

resulted from troops introducing the disease, which was

spread by aegyptL This mosquito has been nearly eradicated

from the island, but albopictus is widely distributed and is a

constant health menace.

References : 188, 356.

Encephalitis.—Japanese encephalitis occurred in an epidemic outbreak in 1947-48 in Guam. No vector has yet been

implicated.

References : 246, 257.

Filariasis.—This disease is not known in most of the

Marianas. In Saipan and Tinian filariasis infection rates of

5 to 15 percent have been reported, but the disease is entirely

absent from Guam. However, various degrees of filariasis

endemicity characterize most of the Carolines from the

Palaus in the west to Ponape and Mokil in the east. Not

much is known about the disease in the northern half of the

Marshalls, but several islands at the south end have human

infections. Filariasis occurs in almost all the Gilberts, where

infections of over 15 percent have been reported from some

of the islands. As in Melanesia, the sole parasite throughout Micronesia is Wuchereria bancrofti^ which causes the

periodic form of tHe disease. There is no nonperiodic

filariasis in this entire area.

In the Marianas, Marshalls, Gilberts, and Truk (Carolines), Culex pipiens quinquéfasciatus appears to be the sole

vector. In the Palaus (western Carolines) it is the proved

vector of primary importance. In Truk Culex annulirostris

is believed to be a potential secondary vector.

Reference : 197.

Polynesia

Physical Features.—Polynesia comprises a large group

of islands in the central Pacific, including the island groups

of Hawaii, Samoa, Tonga, Tokelau, Tubuai, Tuamotu, Society, Marquesas, Cook, EUice, and Easter. They are either

summits of high volcanic mountains or low coral islands,

with tropical or subtropical climates.

Reference : 301,

Malaria.—This disease is not present in Polynesia.

Reference : 67.

Yellow Fever.—This disease is not present in Polynesia.

Reference : 256.

Dengue.—Aedes aegypti and A, albopictus have been responsible for severe epidemics of dengue since 1903 in the

Hawaiian Islands, but this disease is maintained at a very

low level between epidemics. In Tahiti, the Marquesas, and

American Samoa, epidemiological evidence indicates that

dengue is also transmitted by Aedes polynesiensis.

References : 64, 256.

Encephalitis.—No information is available.

Filaria3is.—The dividing line between periodic and nonperiodic filariasis runs between the Gilbert and Ellice Islands,

south between Fiji and the New Hebrides to the southern

tip of the latter, then westward between the New Hebrides

and the Loyalties. All filariasis east of this line in the Pacific

is caused by nonperiodic Wuchereria banerofti. This form

of the disease is found in most of the islands of Polynesia.

The following places may be exceptions, since there is no in-
MOSQUITOES OF MEDICAL IMPORTANCE 107

formation available for them: Phoenix Islands, northern

Cook Islands, northern and southern Line Islands, and the

islands lying to the south and west of the Tuamotus.

Aedes polynesiensis is the principal vector in most of this

area. This mosquito, previously thought to be Aedes pseudosGutellaris and appearing under that name in most of the

medical literature, has only recently been elevated to specific

rank. It is widely distributed over Polynesia. True

pseudoscutellaris is believed to inhabit only a small part of

western Fiji, and a new evaluation of the ability of this

species to transmit filariasis is needed, since all previous

records refer to polynesiensis. Tonga is unique in that

polynesiensis is completely replaced by Aedes tongae., another

member of the Aedes scutellmis group.

Experiments in Hawaii reveal that the hancrofti infections

in Okinawans could be transmitted easily by Hawaiian Culex

pipiens quinquéfasciatus^ the commonest mosquito in Oahu.

In further experiments Samoan hancrofti did not mature in

this vector to the infective stage. Filariasis is not a problem

in the Hawaiian Islands.

References: 197, 248, 263.

Anopheles Species in Australasia

amictus andctus

amictus hilli

\annularis

annulatus

^annulipes

atratipes

"^hancrofti

hancrofti harhiventris

\barbirostris

harbvwibrosus

clowi

colledgei

"^farauti

incognitus

insulaeflorum

karwari

fkochi

^koliensis

longirostris

See footnote 8, p. 63.

Iwigae

^fnaculatus

msraukensis

\rmnirrms

nataliae

\nigerrimus

novaguinensis

\philippinensis

powetli

pseudoharbirostris

pseudostigmaticus

"^punctulatus

solo7nonis

stigmatidùs

"^suhpictus

Uundaicus

\tessellatus

vagus

NOTES ON THE MEDICALLY IMPORTANT SPECIES

In this section the following information is given about

each species: (1) Bionomics, (2) relation to disease, and (3)

recorded distribution. Many of the important and abundant

disease vectors discussed in this section have been called by

various names. Where confusion with other species and

names might result, a short explanation precedes the notes

on bionomics. Some of the information pertaining to bionomics and relation to disease has been taken from reviews,

many of which summarize numerous short papers of restricted scope. These short papers are not listed among the

references. The distribution is indicated by a list of countries

from which the species has definitely been recorded. Bibliographic documentation of these distribution records has been

omitted for brevity.

No attempt has been made to determine the exact taxonomic

status of each subspecies herein discussed. Currently accepted nomenclature has been followed insofar as possible,

but the primary objective has been to correctly differentiate

the known disease vectors from other mosquitoes. Future

investigations of distribution, biology, and taxonomic

characters will eventually demonstrate the true relationships

of these subspecies. This task is one for specialists with adequate time and opportunities for field studies. Our present

knowledge of medically important mosquitoes seems to fully

justify the treatment of many forms as subspecies, although

some of these may prove to be distinct species.

AEDES SPECIES

Aedes aegypti (Linnaeus)

Many names have been applied to Aedes aegypti.^ a

ubiquitous and important mosquito. One of these, Stegomyia fasciata^ is most commonly encountered in the early

literature.

Bionomics.—The adults of aegypti seek shaded places for

resting and are rarely found over 100 yards from their larval

habitats. The females are daytime biters by preference,

although they sometimes attack at dusk or in complete darkness. Usually they are markedly anthropophilic, although

in Uganda they seldom bite man and rarely seek blood meals

during the day. The eggs are laid singly on wet surfaces

near the waterline, and oviposition may continue for as long

as 30 days after copulation.

The typical larval habitat of this species is in small artificial water containers, such as flower bowls, discarded cans,

old tires, and gutters. Such containers throughout most of

the distribution of this mosquito are close to human habitations, both outside and inside living quarters of all kinds.

When the adults can easily find blood meals away from such

locations, the larvae are found in small water receptacles

distantly removed from human dwellings. The larvae live

less commonly in rock holes and holes in recently felled trees.

Sometimes they are found in plant axils. Holes in tree

trunks lying on the ground are preferred to trees with

vertical entrances. The water in many rock holes is exposed

to rapid drying, and in such habitats the larvae must develop rapidly. The period from hatching to emergence has

taken as little as 85 hours in the laboratory. This mosquito

has a drought-resistant egg, since the larvae have been obtained by immersing dried tree-hole material in water. Not

all hatching occurs on the first immersion, but fewer larvae

are obtained at each succeeding attempt.

It appears that aegypti can survive for short periods at

temperatures that tend to limit its distribution. The species

has been found at an altitude of 8,000 feet and commonly is

present in Africa at 6,000 to 7,500 feet. It is very widely

distributed, probably because it is the most adaptable of all

its close relatives, with the possible exception of Aedes

vittatus.

Relation to Disease.—A. aegypti is regarded as the only

vector of classical urban yellow fever, which most commonly

occurs in epidemic form. The wide distribution of this

mosquito, its adaptability to urban breeding places, and

the fact that yellow fever virus matures in it combine to

make it very important. The absence of yellow fever in the

presence of large populations of aegypti throughout most of

the Orient remains a mystery. It is believed that dengue

may in some way actually replace that disease there. A.

aegypti takes a small part in the transmission of jungle

yellow fever when living at some distance from human habitations. Together with Aedes alhopictus and A. scutellaris

it is an important vector of dengue.

References: 11^, 152, 253, 256, 261. (For additional information,

see the descriptions of yeUow fever and dengue under each region

in the preceding section. )
108 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Recorded Distribution.—

Africa (almost all countries)

Andaman Islands

Antigua

Argentina

Assam

Australia

Bahama Islands

Barbados

Bismarck Archipelago

Borneo

Brazil

British Guiana

British Honduras

Caucasia

Celebes

Ceylon

Chile

China

Colombia

Cook Islands

Corsica

Costa Rica

Crete

Cuba

Cyprus

Dominican Republic

Ecuador

El Salvador

England

Fiji

Formosa

France

French Guiana

Gilbert Islands

Greece

Guam

Guatemala

Haiti

Hawaii

Honduras

India

Indochina

Iraq

Israel

Italy

Jamaica

Japan

Javxi

Lesser Antilles

Malaya

Malta

Mexico

Moluccas

New Caledonia

New Guinea

New Hebrides

Nicaragua

Nicobar Islands

Pakistan (East and West)

Panama

Paraguay

Peru

Philippine Islands

Pitcairn Island

Portugal

Puerto Rico

Samoa

Sardinia

Society Islands

Solomon Islands

Spain

Sumatra

Surinam

Syria

Thailand

Tonga

Trinidad

Turkey

united States

Venezuela

Yugoslavia

Aedes africanus (Theobald)

Bionomics.—Although the adults of Aedes africanus have

been recorded as occasionally entering houses, they usually

inhabit high rain-forest canopy. They are found only m

small numbers at ground level. The females are markedly

crepuscular and have a pjeak of biting activity shortly after

sunset, although day biting has been observed in the dense

evergreen shrub in Tanganyika. The females apparently

prefer primate hosts. They bite man readily and will feed

avidly on anesthetized monkeys. The seasonal distribution

of this species closely parallels the rainfall. Large numbers

of adults appear shortly before a period of maximum ram

in July and again during the less intense autumn rains.

Tree holes and bamboo stumps in the forest some distance

from human dwellings are the most common larval habitats.

The larvae are occasionally encountered in artificial containers, such as discarded tires and cans. In Uganda they

have been found in native bamboo waterpots, tree-root

buttresses, and shaded holes in granite. In the "Kaimosi

Forest" of Uganda they have been found in well-shaded holes

in various types of boulders. When the larvae are not in

tree- or rock-hole habitats, it is probably because heavy rains

have washed them out of these normal breeding places.

Relation to Disease.—A. africanus adults are often infected with yellow fever virus in nature. This species is regarded as a potentially dangerous vector, at least in AngloEgyptian Sudan. In Bwamba County, Uganda, it is the

dominant culicine in the forest canopy. It feeds so readily on

monkeys at night that it is regarded as the principal link in

the monkey-to-monkey chain of yellow fever in the forests of

East-Central Africa. Its importance in West Africa has

not yet been determined.

References: Ilk, Í51, 152, 151, 169, 253, 351. (For additional information, see the descriptions of yellow fever under Eritrea, the

Somalilands, and Ethiopia; West-Central Africa; and East-Central

Africa. )

Recorded Distribution.—

Anglo-Egyptian Sudan

Angola

Cameroons

Dahomey

Ethiopia

French Equatorial Africa

French Guinea

Gold Coast

Kenya

Liberia

Mozambique

Nigeria

Northern Rhodesia

Nyasaland

Ruanda-Urundi

Sierra Leone

Tanganyika

Uganda

Aedes albopictus (Skuse)

Bionomics.—Females of Aedes albopicttcs have often been

rep>orted persistently biting man by day in the shade ; night

biting is comparatively rare. In Shantung Province, China,

and in other localities this species has been recorded as the

most common daytime-biting mosquito. Because the species

prefers woodlands to the immediate vicinity of human habitations, it is less urban than Aedes ae^ypti, £ilÜiough in some

areas albopictus is commonly found near human dwellings.

Larvae usually live in tree and rock holes, cut bamboo, leaf

axils, coconut shells, and many types of artificial containers,

such as water buckets, cans, bottles, and drains. They are

particularly abundant where rainwater is caught and stored,

as in Guam. In Mauritius larvae live in most tree holes

containing water.

Relation to Disease.—A, albopictus is an efficient vector of

dengue in all areas where both the mosquito and the disease

appear together.

References: 114, ^69, 188, 254, S09. (For additional information,

see the descriptions of dengue under Madagascar and Adjacent Islands, North Europe and North Asia, all countries in the Orient,^** and

Australasia.)

Recorded Distribution.—

Assam

Australia

Borneo

Burma

Caucasia

Celebes

Ceram

Ceylon

China

Christmas Island (Indian Ocean)

Fiji

Formosa

French Somaliland

Hawaii

India

Indochina

Japan

Java

Lesser Sunda Islands

Lombok

Madagascar

Malaya

Mauritius

Moluccas

New Guinea

Pakistan (East and West)

Philippine Islands

Pitcairn Island

Réunion

Riouw Archipelago

Seychelles

Soemba

Sumatra

Tahiti

Thailand

Timor

Torres Strait Islands

Aedes furcifer (Edwards)

Bionomics.—Females of Aedes furcifer bite at night in the

forest canopy in Uganda; males are scarce in the forest.

Adults of this species and Aedes taylori fly farther than

A. aegypti and most of its relatives.

Throughout its range, including Gold Coast, Tanganyika,

and Union of South Africa, this species breeds in tree holes.

Relation to Disease.—This mosquito has been found infected with yellow fever in nature and is regarded as a potentially dangerous vector of that disease, at least in AngloEgyptian Sudan.

References: llh 152, 151, 169, 232, 351. (For additional information, see the description of yellow fever under Anglo-Egyptian Sudan.)

Recorded Distribution.—

Anglo-Egyptian Sudan

Gold Coast

Nigeria

Portuguese Guinea

Transvaal

Uganda

'' See footnote 2, p. 2
MOSQUITOES OF MEDICAL IMPORTANCE 109

Aedes leucocelaenus clarki Galindo et al. and Aedes

leucocelaenus leucocelaenus Dyar and Shannon

The range of Aedes leucocelaenMs was previously thought

to extend from Costa Rica to Argentina. This species has

now been divided into the subspecies A, leucoGelMnm leuGocelaerms in southern Brazil and Argentina and A, leuGOcelaenus clarki in Panama and Costa Eica. A, leiwocelaerms

has distinct morphological characters, and it has not been

confused to any great extent with other mosquitoes in the

important medical literature.

Bionomics«—In Panama, platform studies of biting habits

show that adults live at various levels in the forest canopy

but predominate at ground level. The species first appears

there in May at the start of the rainy season and rises to a

peak of abundance during the last part of the rainy season

in September and October. During the dry season it is rare

or absent. In flight-range studies in Brazil, where l&uococelaerms is very abundant, stained adults were captured about

3,200 feet or more from the point of release in the direction of

the prevailing winds. Further studies show that adults will

disperse as far as 31^^ miles from their breeding places.

Relation to Disease.—This mosquito is infected with yellow fever virus in nature. It has been infected in the laboratory and hastransmitted the virus there. Its transmission rate

was 1.29 compared with that of 1.00 ior Aedes aegyptL Because of its indiscriminate distribution at various levels in the

forest, this mosquito in Panama may become infected anywhere within the forest and transmit yellow fever to man at

the edge of or outside the forest. The species reaches a peak

of abundance during the last part of the rainy season, when

most of the yellow fever is being transmitted. It is a proved

vector of yellow fever in the forests of Brazil and Colombia,

but it is probably not abundant enough in Brazil to support

yellow fever alone.

References: 81, 82, 131, 133, 219. (For additional information, see

the descriptions of yeUow fever under Panama, Northwest South

America, Brazil and the Guianas, and Lower South America.)

Recorded Distribution.—

A. I. clarki;

Costa Rica

Panama

A. I, leucocelaenus:

Argentina

Bolivia

Brazil

Aedes luteocephalus (Newstead)

Bionomics.—Females of Aedes luteocephalus throughout

their range bite man and monkeys readily. They prefer the

edges rather than the interior of open forest canopy. They

are crepuscular, having a strongly marked peak of biting

activity beginning shortly after sunset and lasting into the

early night. This mosquito apparently survives the dry season as an adult and has been found in extremely dry areas.

It appears in greatest numbers shortly before the period of

greatest rainfall, from May to July, and there is a smaller

peak of abundance during the lighter rains later in the year.

Larvae prefer tree holes over bamboo stumps or cut bamboo. They have occasionally been reported in sagging roof

gutters, domestic containers, crab holes, and rock pools. In

Anglo-Egyptian Sudan larvae live in tree holes but occasionally are flushed out into ground and rock pools and crab

holes by heavy rain.

Relation to Disease.—A. luteocephalus has been found

naturally infected with yellow fever. It is regarded as a

potential vector in Anglo-Egyptian Sudan.

References: II4, 152, 157, 169, 232, 253, 351. (For additional information, see the description of yellow fever under Anglo-Egyptian

Sudan.)

Recorded Distribution.—

Anglo-Egyptian Sudan

Bechuanaland

Belgian Congo

Eritrea

French Guinea

Gold Coast

Kenya

Liberia

Nigeria

Northern Rhodesia

Nyasaland

Sierra Leone

Southern Rhodesia

Uganda

Aedes metallicus (Edwards)

Bionomics.—In Anglo-Egyptian Sudan Aedes metallicus

never flies far from its breeding places. It usually bites outdoors at ground level, although it enters houses to obtain

blood meals. In Mombasa the females bite man freely, although they have also been reported to attack horses and

cattle. Adults are crepuscular and have a well-defined peak

of biting activity shortly after sunset.

Tree holes have often been recorded as the preferred

larval habitat. Larvae are reported to be abundant in holes

in Ricinoderulron rautanenii m the vicinity of Livingstone,

Northern Ehodesia. They have been taken from ground

water under trees, and from banana axils, coconut shells,

and artificial containers, such as old cans, iron and bamboo

pots, and concrete basins. Larvae have been obtained by

immersing dried material from tree holes in water, an indication of the presence of drought-resistant eggs.

In various parts of its range this species has been taken at

altitudes up to about 6,000 feet. It is reported to be the

most common tree-hole breeder in Anglo-Egyptian Sudan.

Relation to Disease.—^Adults are occasionally infected

with yellow fever virus in nature. The species is regarded

as a potential vector of that disease, at least in AngloEgyptian Sudan.

References: II4, 151, 169, 232, 253, 261, 351. (For additional information, see the description of yellow fever under Anglo-Egyptian

Sudan. )

Recorded Distribution.—

Anglo-Egyptian Sudan

Angola

Bechuanaland

Belgian Congo

Gold Coast

Kenya

Mozambique

Northern Rhodesia

Senegal

South-West Africa

Tanganyika

Transvaal

Uganda

Zanzibar

Aedes poicilius (Theobald)

Aedes poicilius was originally described in the genus

Finlaya^ which is now a subgenus of Aedes. It has only

recently become prominent as a disease vector.

Bionomics.—Adults bite severely at dusk, and in the Philippines also during the day. They have very high and exacting humidity requirements, which make them extremely

difficult to rear in the laboratory.

The larvae inhabit water-filled axils of aroid plants, banana, taro, abacá, pandanus, and pineapple.

Relation to Disease.—Even though no conclusive field or

laboratory observations have proved this species to be of

medical importance, it is believed to be tiie principal vector

of filariasis in the abaca-growing areas of the Philippines.

Larvae of Wuchereria iancrofti have developed in poicilius

to the infective stage in the laboratory.

References: 207, 2Jt1, 293. (For additional information, see the

description of filariasis under Philippine Islands.)

Recorded Distribution.

Borneo

Burma

Celebes

India

Java

Lombok

Malaya

New Guinea

North Benjîal

Philippine Islands

Simalur

Sumatra
no AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Aedes polynesiensis Marks

Most medical literature prior to 1951 refers to Aedes

polynesiensis as A. pseudoscuteUaris.

Bionomics.—Adults are common in and around native villages and are highly anthropophilic, although they attack

animals as well. Females bite during the day, usually in

deep shade or under cloudy conditions. The flight range is

limited to about 100 yards.

Larvae are found in small water collections. Coconut

shells, husks, stumps, tree holes, cacao pods, open concrete

drains, tin cans, and lava holes are common breeding places.

Relation to Disease.—Since it is a day biter, polynesiensis

transmits nonperiodic Wuchereria hancrofti. It is the only

important intermediate host of this filaría in Polynesia, excluding Tonga. Recent laboratory investigations have

shown that this mosquito, a member of the Aedes scutellaris

group, can transmit dengue from monkey to monkey, and

epidemiological evidence indicates that this species, and possibly others in the group, transmit dengue in Tahiti, Fiji,

Marquesas, and American Samoa.

References : 122, 2Jf8, 289. (For additional information, see the descriptions of dengue and filariasis under Australasia.)

in large areas of northern New Guinea where aegypti was

scarce or absent.

References : 102, 245, 2Jf8, 314. (For additional information, see the

description of dengue under Australasia. )

Recorded Distribution.—

Austral Islands

Cook Islands

EUice Islands

Mangareva

Marquesas Islands

Pitcairn Island

Samoa

Society Islands

Tokelau

Tuamotu Islands

Wallis Islands

Aedes scutellaris (Walker)

Aedes scutellaris consists of a complex of about 17 distinct

but very closely related species known as the scutellaris

group. The remarks below apply to only one of this group,

i. e., scutellaris in the restricted sense. This species is treated

as Aedes scutellaris var. kehridetis in some of the important

literature relating to transmission of disease.

Bionomics.—A, scutellaris is a "bush" species. Adults

are usually found close to the ground in cool, damp, wellshaded places under low vegetation. Their flight range is

very limited. Females enter buildings only to feed and

usually leave soon afterward, showing little or no tendency

to settle or rest inside. In New Guinea their diurnal biting

activity was surprisingly regular under many different conditions and in different localities. A sharp peak of activity

occurred between 6 and 7 p. m. and a lower sharp peak

between 7 and 8 a. m. On overcast days adults entered

tents and fed during almost the entire day.

In Espíritu Santo, New Hebrides, larvae were found in

all kinds of artificial containers. Ends of upturned oil

drums in fuel dumps and sagging tents were their principal

habitats. Other breeding places were various small, shallow

collections of water in discarded tires, tin cans, corrugated

sheet iron, tarpaulins, old helmets, canteen cups, coconut

shells, and cacao pods. Larvae also inhabit raised coral

reefs and open wells and are commonly found in tree holes.

In New Guinea they live in almost any small, well-shaded

collection of clear rainwater. The species does not breed

in heavily polluted water.

Relation to Disease.—Because scutellaris appears in large

numbers and attacks man readily and because other vectors

are absent, it is regarded as a vector of dengue in Espíritu

Santo, New Hebrides. It definitely transmitted that disease

Recorded Distribution.

Aroe Islands

Ceram

Moluccas

New Britain

New Guinea

New Hebrides

New Ireland

Philippine Islands

Aedes simpsoni (Theobald)

For a number of years Aedes simpsoni was thought to be

represented by the variety lilii over most of its range, and

it has not been extensively known by any other name in the

literature. Recent studies indicate that lilii may not be a

true variety. We believe that the name should not be used

until further investigations disclose the true status of this

mosquito.

Bionomics.—Adults of simpsoni occasionally enter human

dwellings, usually in small numbers. In Nigeria and Gold

Coast females of forest populations very rairely bite man,

but in Uganda, British Cameroons, and Transvaal they frequently attack man in large numbers. In the plantations

of Uganda, simpsoni is a much more common biter during

the day than at night. The females also bite horses, cattle,

goats, fowl, and monkeys.

In Uganda, leaf axils of Xanthosoma sagittifolium^ with

a capacity of approximately 4 to 8 cc, and axils of the

"Gonja" variety of cultivated banana and pineapple {Ananas comosus) are the most important breeding places of

simpsoni larvae. Workers in Uganda have shown that the

Gonja variety of banana is much preferred by simpsoni over

others. To be good breeding places, axils must hold clear

latex-free water for a long enough time for larvae to complete development. Light rains are sufficient to keep the

axils ^Ued. Larvae sometimes exist in only a thin film of

water in the axils. Wild banana, screw pine (Pandanus),

and Golocasia esculenta contain no simpsoni^ probably because these plants grow in unsuitable environments. Tree

holes are uncommon habitats in Uganda^ but many authors

have reported finding simpsoni in tree holes ^ elsewhere.

Larvae are rare but present in rot holes of Ricinodendron

rautanenii near Livingstone, Northern Rhodesia. Immature

stages are also occasionally found in artificial containers,

such as discarded cans, bottles, tubs, and tanks, and have been

recorded from rock pools, concrete basins, bamboo pots, coconut shells, and axils of Dracaena and Strelitzia. Larvae

have been obtained by immersing dried material from tree

holes in water. A. simpsoni apparently survives the dry season because of its drought-resistant eggs.

A, simpsoni seldom enters closed forest canopy, preferring

plantations and shambas at forest edges. Adults have been

observed at altitudes up to 6,000 feet. In Anglo-Egyptian

Sudan simpsoni is reported to be common in many areas with

rainfall of over 15.8 inches per year.

Relation to Disease.—Adults are infected with yellow

fever virus in nature. This species is regarded as a potentially dangerous vector and is probably the principal vector

of yellow fever from animal reservoirs to man, at least in

Anglo-Egyptian Sudan and possibly in West Africa as

well.

References: 114, 145, 151, 169, 232, 2S3, 261, 351. (For additional

information, see the descriptions of yeUow fever under Anglo-Egyptian Sudan ; Eritrea, the SomaUlands, and Ethiopia ; West Atrica ;

West-Central Africa; East-Central Africa; and Southern Africa.)
MOSQUITOES OF MEDICAL IMPORTANCE 111

Recorded Distribution.-

Anglo-Egyptian Sudan

Angola

Bechuanaland

Belgian Congo

Cape Province

Eritrea

Gambia

Gold Coast

Kenya

Mozambique

Natal

Nigeria

Northern Rhodesia

Nyasaland

Pemba

Ruanda-Urundi

Sierra Leone

Southern Rhodesia

Tanganyika

Transvaal

Uganda

Zanzibar

Zululand

Aedes taylori Edwards

Bionomics.—In southern Nigeria Aedes taylori is predominantly a tree-top biter, whose vertical distribution in

the forest does not markedly change during the night. In

trapping experiments at Itowolo 75 percent of taylori females were caught 40 to 52 feet above the ground and only

about 3 percent at ground level. Mating and other activities

associated with swarming are thought to occur during the

early evening. A. taylori aud Aedes furci fer fly farther

than J.. aegypti or any of its relatives.

The preferred breeding places are tree holes. Larvae are

present but rare in rot holes of Ricinodendron rautanenii in

the vicinity of Livingstone, Northern Rhodesia, and are

found closely associated with Aedes adersi in tree holes in

Tanganyika. Adults are most abundant before the maximum rainfall in July, indicating that tree holes may be filled

to a level that enables most larvae to mature long before

maximum rainfall, which may be great enough to cause these

habitats to overflow. A small secondary population peak

follows the rainy season in October.

Relation to Disease.—^Adults have been found infected

with yellow fever virus in nature. The species is regarded

as a potentially dangerous vector of that disease, at least in

Anglo-Egyptian Sudan.

References: Ilk, 151, 169, 252, 261, 351. (For additional information, see the description of yellow fever under Anglo-Egyptian

Sudan. )

Recorded Distribution.—

Nigeria

Northern Rhodesia

Tanganyika

Aedes togoi Theobald

Bionomics.—Aedes togoi is common in Honshu, Shikoku,

Kyushu, and Hokkaido, Japan. It habitually enters houses,

attracted by artificial light. In Japan it is not regarded as

anthropophilic, since it has rarely been taken in collections

of mosquitoes biting man, although it is the most widely distributed and highly adapted of the Japanese Aedes species.

However, in northeast China adults enter dwellings at night

and are serious pests in some localities. They are more

abundant along the coast than elsewhere. There are generally two peaks of abundance, one in the spring and one

in the fall.

Larvae are commonly found in water tanks, buckets,

flowerpots, borrow pits, ground pools, ditches, and gutters.

Because they prefer small artificial containers in the open

sun, they are least numerous during the hot summer when

evaporation is high. The species probably overwinters in the

larval stage.

A salt-water form of togoi has been recorded from Japan

and Hong Kong. Along the coasts of Japan and China

larvae have been collected from tidewater ground and rock

pools where the water has a chloride content nearly equivalent to that of sea water. Little is known of the behavior

or distribution of this coastal form, although in Hong Kong

adults are numerous and females bite persistently.

Relation to Disease.—A. togoi has been incriminated in

the transmission of Japanese encephalitis, but it is believed

471779 O—59 8

to play a minor role. This mosquito has been experimentally infected with Wuehereria hancrofti and transmits

filariasis in the vicinity of Hong Kong.

References: 122, 199, 223. (For additional information, see the

descriptions of filariasis under North Europe and North Asia, and

South China.)

Recorded Distribution.—

China (north)

Japan

Korea

Ogasawara-gunto

Siberia

Aedes tongae Edwards

Aedes tongae was originally described as a variety of Aedes

variegatibs. At the present time it is universally accepted as

a full species.

Bionomics.—Very little information is available on the

habits of this species. Adults are known to attack man, and

larvae inhabit coconut husks, tree holes, and wells.

Relation to Disease.—This is the only representative of

the scutellaris group in Tonga and the only mosquito there

that can transmit the nonperiodic form of Wuehereria

hancroftL

Reference: 122. (For additional information, see the description

of filariasis under Australasia. )

Recorded Distribution.—

Rennell Island

Sikaiana

Tonga

Aedes vittatus (Bigot)

Bionomics.—In Northern Rhodesia, parts of Uganda,

Eritrea, and Spain Aedes vittatus bites man readily during

the day. However, in Southern Africa and the "Kaimosi

Forest" of Uganda, it does not seem to attack man at all.

Adults have a marked ability to withstand low temperatures.

Large numbers of larvae have been found exposed to the

sun in rock pools, beds or margins of streams, holes in coral,

tree holes, domestic utensils, boats, cattle hoofprints, barrels,

buckets, discarded tins, and wells. In one locality larvae

were found in a turbid rain pool without vegetation, in broken

shade, and with water having a chloride content of about 0.04

percent, an unusual habitat for this species. In arid places

the larvae are found in large containers of conserved water.

A, mttatus usually inhabits exposed rock holes when streams

are dry. Since these holes dry up rapidly, the larvae must

mature rapidly. The pupae have been found in rock holes

only 3 days after a rain.

This species inhabits open country and has been found at

altitudes up to 6,700 feet.

Relation to Disease.—A. vittatus is infected with yellow

fever virus in nature and is regarded as a potentially dangerous vector, at least in Anglo-Egyptian Sudan.

References: llJf, 152, 157, 169, 253, 351. (For additional information, see the description of yellow fever under Anglo-Egyptian Sudan. )

Recorded Distribution.-

Anglo-Egyptian Sudan

Angola

Arabia

Bechuanaland

Belgian Congo

British Somaliland

Cameroons

Ceylon

Corsica

Eritrea

Ethiopia

French Equatorial Africa

Gambia

Gold Coast

India

Indochina

Kenya

Liberia

Mozambique

Natal

Nigeria

Northern Rhodesia

Nyasaland

Pemba

Sierra Leone

Sokotra

Southern Rhodesia

South-West Africa

Tanganyika

Transvaal

Uganda

Zanzibar

Zululand
112 AGRICULTURE HANDBOOK 15 2, V. S. DEPT. OF AGRICULTURE

ANOPHELES SPECIES

Anopheles aconitus Dönitz

In the early literature Anopheles aconitus has usually been

confused with A, minimus. Philippine records refer to

Anopheles fllipinae.^ since aconitus is not present in that

country.

Bionomics.—Females of aconitus feed avidly on man in

houses and bite domestic animals in nearby shelters. In the

Sunda Islands they definitely prefer human blood. Elsewhere they occasionally rest in cowsheds. Experiments in

the Sundas demonstrated flight ranges as far as 1 mile,

although the effective range is probably not so great. This

mosquito is a nocturnal flier. Night catches along the

Bengal-Burma border indicate that the fewest adults appeared in April and May, with an abundance peak in midNovember.

Larvae are found in both mountains and plains, principally

in fresh water. In Burma the favored breeding places are

fresh-water pools in creek and river beds containing abundant

aquatic vegetation and shade, as well as lakes, drainage

ditches, and fresh-water fishponds. In Malaya the larvae live

in the clear, open water of swamps and in slowly flowing

w^ater. Ricefields and their drainage systems are important

breeding places for the larvae, especially where the water

is bordered by grass. Larvae first appear when the rice crop

is about half grown, and their numbers increase rapidly after

the harvest. Along the Bengal-Burma border aconitus is

found in marshy areas atthe foot of hillocks, where immature

stages have been taken from pools heavily covered with

vegetation.

This species is common in foothills of 1,200 to 3,000 feet,

especially in the Sunda Islands. It is also present but less

common in the plains.

Relation to Disease.—A, aconitus is regarded as a malaria

vector of some importance in spite of its having been confused with minimus. In several experiments in the Sunda

Islands it has been found in the vicinity of both man and

cattle. The females taken near man had old blood, and most

of the unfed females were taken near livestock. However,

in other collections up to 60 percent of females contained

human blood. This species is abundant but rarely infected in

nature in Celebes and Borneo, but it is probably a dangerous

vector in parts of Sumatra and Java. In both Malaya and

Burma it has been found infected, but it is not a proved vector

in either of these countries. Naturally infected aconitus

females have been found in Indochina. This species is

strongly suspected as one of the most dangerous vectors in

southern Annam and is a secondary vector in Tonkin and

Cochin China. It is a vector in CamÍDodia, but its importance

there is still in dovibt, and its role in Thailand is not yet clear.

In China the species is not important as a vector of malaria

because of its scarcity.

Development of Wuchereria hancrofti to the infective stage

in aconitus has been reported from Kabaena near Celebes, but

the role played by this species in the transmission of filariasis

is not known with certainty.

References : 88,120-122, 161, 242, 324. (For additional information,

see the descriptions of malaria under Indochina, Thailand, and Sunda

Islands.)

Recorded Distribution.

Andaman Islands

Assam

Bali

Borneo

Burma

Celebes

Ceylon

China (South)

Flores

Hainan

India

Indochina

Java

Malaya

Pakistan (East)

Philippine Islands

Soemba

Soembawa

Sumatra

Thailand

Anopheles adenensis Christophers

Anopheles adenensis was originally described in 1925 as a

variety of Anopheles culicifades., to which it is closely

related.

Bionomics.—This is a common household mosquito,

readily feeding on man. Larvae breed in clear water in

wells, stream beds, and rain pools, and also in tanks, cisterns,

irrigation ditches, sluggish streams, ricefields, and borrow

pits. In Hodeida on the Red Sea coast of Yemen larvae were

found in brackish water in the open cement basins of mosques.

Relation to Disease.—There is no well-defined evidence,

such as dissection records, to show that this species is a

malaria vector. Nevertheless, it has been reported as a vector in the vicinity of Hodeida and in and about Aden. Since

culicifacies is an outstanding vector of malaria, adenensis

must be suspected as an efficient vector as well. In addition

to its distribution in the Orient, typical culicifacies is found

in Oman, Trucial Oman, and Bahrein.

References: 61, 91, 119, 206, 258, 296. (For additional information,

see the descriptions of malaria under Eritrea, the Somalilands, and

Ethiopia ; and Arabia. )

Recorded Distribution.—

Arabia

Eritrea

Anopheles albimanus Wiedemann

Because of its extensive distribution and importance,

Anopheles albimanus is cited a great deal in the literature.

The dry-weather varieties iisignatus and trisignatus have

been described from Mexico and Guatemala.

Bionomics.—Adults of albimanus are domestic and are the

most common Anopheles species in houses in many localities.

Most females return to outdoor resting places soon after

feeding but do not usually congregate in daytime resting

places. They feed as readily on livestock as on man, depending on W'hich is available, and will choose either as a host.

They become active during twilight and in Panama fly in the

evening, usually between 6: 30 and 7: 30. Copulation has

been observed only during twilight. The species is a strong

flier. Females can travel as far as 12 miles from their breeding places in short flights, with intervals of feeding betw^een

them. They have been observed to fly one-half to 1 mile at

right angles to a strong wind and 5 to 40 feet above the

ground. They have been trapped li/¿ miles froni their nearest breeding places.

Larvae live in a great variety of sunlit water collections,

fresh or brackish. Since the water must contain many microorganisms, be without putrefaction, and be in biological equilibrium, most temporary breeding sites are not suitable.

Larvae are found principally in swamps, large ponds, lakes,

seepages, irrigation ditches, and ricefields. A. albimanus inhabits newly established ricefields in northern Venezuela to

the exclusion of any other habitat. UtriculaHa and Naias

are particuhirly favored as water plants, especially when

water levels are such that these plants are at the surface.

During the dry season larvae become adapted to crab holes,

tanks, water troughs with algae, and drums of rainwater.

In the very dry northern part of the Yucatan Peninsula

albimanus is found near villages where water is stored in

artificial reservoirs, but during the rainy season the species

inhabits swamps in isolated areas. During the rainy season

in Mexico larvae live farther inland than during the dry

season.

A. albimanus is essentially a mosquito of hot, humid climates, and its seasonal abundance is directly related to rainfall. In most localities its numbers are noticeably reduced

during the dry season, although in the Canal Zone the dry
MOSQUITOES OF MEDICAL IMPORTANCE 113

season promotes the growth of certain aquatic plants favorable for larval development. At Esquipulas, Guatemala,

albimanus was found at an altitude of 3,200 feet, where it

had become adapted to an extremely dry climate.

Relation to Disease.—This is the most important and effective malaria vector in the Caribbean region. Shortly after

it was introduced into Barbados, a severe outbreak of malaria occurred where none had previously been reported. It

is stated that albimanus is the principal vector in 17 of the

20 Caribbean countries and is regarded as the chief vector

along the gulf coast of Mexico.

References: 59, 210, 226, 308. (For additional information, see

the descriptions of malaria under America North of Mexico, West

Indies, Mexico, Central America, Panama, Northwest South America,

and Brazil and the Guianas. )

Recorded Distribution.—

Bahama Islands Haiti

British Guiana Honduras

British Honduras Jamaica

Colombia Mexico

Costa Rica Nicaragua

Cuba Panama

Dominican Republic Puerto Rico

Ecuador Surinam

El Salvador Venezuela

French Guiana Virgin Islands

Guatemala

Anopheles albitarsis Arribalzaga

Bionomics.—In Panama albitarsis does not attack man or

enter his habitations, even when breeding places are close by.

In Brazil it enters houses in large numbers during the night

and day. An unnamed form inhabits eastern Eio Grande do

Norte, Paraíba, Pernambuco, Alagôas, and the Amazon

Valley. It also enters houses and feeds on man in large

numbers.

The preferred larval breeding places are large bodies of

water, such as large ponds, ground pools, marshes, bays, and

overflow areas near rivers. In Costa Eica larvae live in

sunny ponds with algae and vegetation, and in Gatun Lake,

Canal Zone, they have been found in mats of Chara^ Naias^

and Utricularia exposed to full sunlight. Immature stages

have been observed in Trinidad ricefields and in Venezuelan

lakes with Pistia and Eichhornia.

Relation to Disease.—A, albitarsis is believed to be a

vector of malaria in parts of South America, although the

evidence is conflicting. In Para and Paraíba, Brazil, it was

the only Anopheles species found in large numbers in houses

in the presence of endemic malaria and the only species

found to be infected. It is important along the coastal plain

south to Espirito Santo and is strongly suspected as a vector

of malaria in parts of Venezuela. In Panama, on the other

hand, it is hard to find and not believed to be important

there. In Argentina the species is said to be more zoophilic

than anthropophilic, although it may be important in the

northeastern part of that country.

References: ^6, 19, 292, 308. (For additional information, see the

descriptions of malaria under Panama, Northwest South America,

Brazil and the Guianas, and Lower South America.)

Recorded Distribution.—

Argentina Guatemala

Bolivia Panama

Brazil Paraguay

British Guiana Trinidad

Colombia Uruguay

Costa Rica Venezuela

Anopheles annularis van der Wulp

In Malaya and elsewhere Anopheles annularis has been

confused with A. philippinensis and A. palUdus^ but it is

morphologically distinct from them as shown in the pictorial

keys. The name most commonly applied to this species in

the past is Anopheles fuliginosus. A variety, adiei^ is regarded as a seasonal form.

Bionomics.—A. annularis rests in large numbers in cattle

sheds and usually in smaller numbers in human habitations. In Bengal it is said to be definitely anthropophilic, even though it is usually divided there in about equal

proportions between houses and cattle sheds. However,

throughout most of the Orient it is zoophilic. In Malaya

the females are principally zoophilic but will attack man

readily. In Yunnan they feed on humans in large numbers.

They are active in the evening and leave their biting

places at dawn. During the day resting adults are found

along eroded, overhanging stream banks at some distance

from human dwellings. They are attracted to artificial

light, often resting near light sources. They are strong

fliers, usually flying a considerable distance from their

breeding places.

Larvae inhabit a variety of clear-water breeding places, including large ponds of fresh water that contain aquatic vegetation, ricefields, shallow vegetated margins of lakes and

slowly moving streams, ditches, swamps, borrow pits, and

large tanks. They appear in large numbers during the rainy

season in Formosa, then become more scarce at the beginning

of the dry season.

This mosquito is found at altitudes of 5,000 to 7,000 feet.

It is believed to overwinter in both larval and adult stages

in northern India.

Relation to Disease.—Early records of malaria transmission by annularis are confused with those of philippinensis,

Nearly all dissection records indicate that natural infection

rates are normally low in Malaya, Burma, China, and the

Philippines. A, annularis is probably of no importance in

Cochin China and Annam, Indochina, and probably of little

importance in the Philippines. Elsewhere, principally in

Orissa in India, Goalpara in Assam, and Formosa, it appears to be only moderately important, probably because of

its zoophilism. Perhaps it is most dangerous when present

in hyperendemic areas.

The complete development of Wuehereria bancrofti has

been observed in annularis in Java, but the extent to which

this mosquito is important in filariasis transmission there is

not known.

References: 61, 8^, 85, 88, 91, 122, 296. (For additional information, see the descriptions of malaria under India and Ceylon, East

Pakistan and Assam, Formosa, and Burma. )

Recorded Distribution.—

Afghanistan Indochina

Assam Java

Bau Malaya

Borneo Nepal

Burma Pakistan (East and West)

Celebes Philippine Islands

Ceylon Soemba

China (South) Sumatra

Formosa Thailand

Hainan Timor

India

Anopheles annulipes Walker

Bionomics.—Females of Anopheles annulipes feed readily

on man just after smiset and the following day tend to remain in shelters. In many parts of Australia females feed

readily in the open near their breeding sites. However, in

the Port Moresby area of New Guinea no adults have been

found near large numbers of larvae. Females fly as far as

one-half mile from their breeding places.

Larvae live in direct sunlight or in shade in clear or

muddy, shallow ground pools, such as hoofmarks, wheel

tracks, trenches, gravel beds, and margins of open sandy
114 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

pools, and in swamps, fishponds, rock pools containing green

algae or weeds, and in water collected in rubbish and large

artificial containers. In the Port Moresby area larvae abound

in shaded pools in beds of intermittent streams, side pools

of flowing streams, and in grassy seepage or overflow areas.

In the vicinity of Melbourne, Australia, they are found in

semiperrnanent creekbeds, swampy areas, wide sluggish

creeks with grassy edges, roadside ditches, and brackishwater pools.

This species is widespread throughout coastal and inland

Australia at altitudes up to 4,000 feet. In the neighborhood

of Townsville and Cairns and in the Northern Territory, the

species is extremely abundant from February to April. During certain times of the year it is the dominant species in

the Port Moresby area of New Guinea.

Relation to Disease.—A, annulipes can be a malaria vector

experimentally. No naturally infected females were found

prior to 1944. Malaria seems to be absent from a large part

of the area where this species lives. However, annulipes can

produce small epidemics of brief duration during the summer

in scattered localities in the southern half of Australia.

References: 67, 120, 202, 228. (For additional information, see the

descriptions of malaria under Australasia. )

Recorded Distribution.—

Australia (New South Wales, Northern Territory, Queensland,

Southern Australia, Victoria, Western Australia)

New Guinea

Tasmania

Anopheles aquasalis Curry

Anopheles aquasalis was known as A. tarsimaculatus by

many early authors. Other names for this species include

Anopheles gorgasi and A, endlianus. Recent malaria literature deals with aquasalis under its present designation.

Bionomics.—Although aquasalis females in Panama are

not particularljr attracted to man, in the Lesser Antilles they

feed more readily on man than on domestic animals. This

is often the only mosquito found in houses, even though several kinds of Anopheles species may be present outdoors. In

British Guiana females distinctly prefer human blood, but

the habit of entering houses is not particularly characteristic of this species there. Females are strong fliers, having

been observed in Trinidad as far as 3 miles from their breeding places. They are believed to migrate annually for distances of 1% to 2% miles.

Larvae usually live in brackish water along the seacoast

in swamps and ditches and in Brazil are rarely found along

rivers beyond the reach of brackish tidal water. They have

been found in Costa Rican fresh-water streams and Trinidad

ricefields and other collections of fresh water several miles

inland. They apparently have no preference for light or

shade, since they have been observed in locations varying

from direct sunlight to deep mangrove.

Relation to Disease.—This species is a vector of malaria

in many of the Lesser Antilles, especially Trinidad, Grenada,

and St. Lucia, and in the Paria Peninsula of Venezuela

directly west of Trinidad. In British Guiana it apparently

is not found in the presence of endemic malaria, but it is

recognized as a vector in Uruguay, northern Argentina, and

in several coastal localities in Brazil south of the Guianas.

In Panama, where adults are rarely attracted to man, malaria

is not proportional to aquasalis activity. The species can be

important only when it is present in large numbers, since

natural infection rates are almost always low. It is responsible for endemic rather than epidemic malaria.

References : 19, 226, 292, 308. ( For additional information, see the

descriptions of malaria under West Indies, Panama, Northwest South

America, Brazil and the Guianas, and Lower South America.)

Recorded Distribution.-

Argentina Nicaragua

BrazU Panama

British Guiana Paraguay

Colombia Peru

Costa Rica Surinam

Ecuador Trinidad

French Guiana Uruguay

Mexico Venezuela

Anopheles aztecas Hoffman

References to Anopheles a^tecus have appeared under the

names A. freeborni^ A. quadrimaculatus, A, maculipennis^ A.

occidentalism and A, átropos. A, aztecus was originally described as a variety of maculipennis. Since about 1940 it

has been regarded as a full species.

Bionomics.—Females of aztecus readily invade human

dwellings and bite man throughout the year. Their numbers

are reduced in the dry spring months and increase to an

average of 21 per house in late summer and fall. During

the rainy period the high relative humidity encourages the

presence of adults rather than the rain and consequent accumulation of water. Inside houses they commonly rest on

walls and under beds. Only small numbers are found resting out of doors and then only at night. Few females are

attracted to animal traps. In the laboratory adults will

mate in small cages, and oviposition has occurred at the low

temperature of 64° to 68° F.

The larvae live in seepage areas and along the vegetated

shores of streams and irrigation canals. They appear to

thrive in clear-water pools containing algae and protozoa.

Potamogetón^ Geratophyllunh^ and Nitella-Ceratophyllum

mats are the most favorable breeding sites. Floating vegetation, especially Eichhornia^ Pistia^ Lemna^ and Azolla^ is

not favorable for larval development. Larvae are present in

breeding places the year around, even when ice forms over

the water. Pollution of water ahvays follows the development of urban areas, and since a^tecus tolerates such conditions more easily than Anopheles pseudopunctipennis with

which it is associated, aetecus is gradually replacing the latter

species on the outskirts of Mexico City.

Relation to Disease.—^Conflicting statements on the ability

of aBtecus to transmit malaria are found in the literature.

In spite of the malaria-free areas in the State of Mexico in

which aztecus is the only anopheline present, the XochimilcoMixquic area of the valley of Mexico, with an elevation of

about 7,500 feet, represents an endemic zone. Here aztecus

is the only vector present in large enough numbers to support the disease. It transmits parasites in the laboratory

and has been found with high infection rates in nature.

Eecent control of this species has markedly reduced malaria

incidence.

References: ^i, 61, 110, 292, 296. (For additional information,

see the description of malaria under Mexico. )

Recorded Distribution.—

Mexico

Anopheles baezai Gater

Anopheles baezai was first described as "umbrosus larval

variation" in 1919 and was redescribed as A. gateri in 1936.

It belongs to the umbrosus group. Very early records are

found under the more inclusive name umhrosus (p. 138).

Bionomics.—Records of mosquitoes caught in animal sheds

and in human-bait traps show that females of baezai are

rather weakly anthropophilic. It is unusual to find them

biting in the daytime. They have been observed resting in

considerable numbers in the early part of the day on frond

bases of nipa palm {Nipa fruticans) in Malaya.

Larvae inhabit brackish-water collections containing a

variable amount of decaying vegetation, especially those that
MOSQUITOES OF MEDICAL IMPORTANCE 115

are periodically refilled by tides. When tidal salt water is

prevented from entering such pools, haezai tends to disappear. These breeding places are often shaded, although

deep shade is not necessary for larval development. Larvae

are often found with Anopheles sundaicus larvae but are

never so numerous.

A, iaezai is rather restricted in distribution and is seldom

very common. It is reported to be most numerous during

and after high equinoctial tides in September, when there

is heavy rainfall.

Relation to Disease.—A. haesai is an inefficient malaria

vector because of extremely high sporozoite rates, unusual

forms of parasites within its body, unsuccessful attempts to

infect females and to transmit malaria in the laboratory, and

animal-positive precipitin tests. However, the females appear in relatively large numbers in human-bait traps as compared with those in animal shelters. Positive dissections

have been made in the Miri-Lutong and Kuala areas of

Borneo. A, haezai may be a vector in these and other localities where its populations are high.

References : 244, S57. ( For additional information, see the description of malaria under Sunda Islands. )

Recorded Distribution.-

Bali

Borneo

Celebes

Indochina

Java

Malaya

Philippine Islands

Sumatra

Anopheles bancrofti Giles

Anopheles hancrofti for a number of years was known

as a variety of A, iarhirostris. It is now recognized as a

full species with several subspecies of its own.

Bionomics.—Although females may attack man outside

during the day, hancrofti is not commonly regarded as anthropophilic. It is abundant near Cairns, Australia, but it

does not attack man in large numbers. However, in Northern Territory it is frequently a troublesome biter, and in

the southern part of Dutch New Guinea—Merauke and

Tanahmerah—'it is found in very large numbers and persistently bites man. Precipitin tests in the latter area revealed that almost 90 percent of the females taken in houses

or animal shelters had human blood and 10 percent had fed

on animals. Females are often found in houses resting on

walls and in mosquito nets and commonly are present in

cattle sheds in large numbers. The species is nocturnal.

Larvae are largely restricted to shade and are often reported in heavy woods or dense aquatic growth. In Queensland and Northern Territory shaded swampy areas are

preferred breeding places. In Dutch New Guinea the species

breeds in jungle and in old, cutoff courses of the Digoel

Eiver, in which the larvae are restricted to areas containing

coarse reeds, algae, and Azolla about 1 mile from shore.

Relation to Disease.—A, hancrofti has not been found

naturally infected with malaria in Australia, although parasites have developed in thisspecies in laboratory experiments.

It should be regarded as a dangerous vector if present in

large numbers. In New Guinea this species rarely appears

in the absence of Anopheles farauti^ which is a very efficient

vector. In Dutch New Guinea hancrofti was found with a

natural infection rate of 4.3 percent in an area where farauti

was also present in small numbers, and in Batu Gajah

hancrofti was shown to be the principal malaria carrier.

The subspecies harhiventris is not known to be a vector of

disease.

References : 67, 121, 228, 324. (For additional information, see the

description of malaria under Australasia.)

Recorded Distribution.—

Australia (Northern Territory, Queensland, Western Australia)

New Guinea

Anopheles barbirostris barbirostris van der Wulp

Anopheles harhirostris has several easily distinguished

varieties or subspecies. Only the typical one appears to be

medically important.

Bionomics.—Over most of its range, including India, harhirostris is principally zoophilic, but it becomes more domestic farther east. In Celebes it enters houses in large

numbers for blood meals, leaving soon after feeding. However, in Indochina a very low percentage of females contains human blood, and in localities in Malaya where livestock is present more mosquitoes appear to have fed on

cattle than on humans. Eesting adults are found along the

undercut banks of streams, sides of shaded stone walls, and

under uninhabited buildings. The adults fly on cloudy days

or in jungle shade. In the Philippines a flight range of approximately 1,300 to 1,970 feet has been observed.

Larvae most commonly are found in the shaded, clear

water of streams, rivers, large vegetated ponds, flowing irrigation ditches and canals, borrow pits, wells, and even in stagnant collections of water. Clear, fresh water is preferred.

Ricefields are common breeding sites in Malaya. In India

the larvae are found at vegetated margins of lakes, swamps,

and sluggish rivers. In Indonesia they live in exposed or

shaded, turbid and stagnant locations from the coast to high

altitudes, and in the Philippines they have occasionally been

taken from salt marshes.

Relation to Disease.—In most areas outside Malaya, Sumatra, and Celebes harhirostris is not considered to be an

important vector of malaria except where it appears in large

numbers or serves as an accessory vector to a more important

transmitter. Experimental infection rates always have been

low. In certain areas of Malaya where natural infection

rates are high, harhirostris contributes to the maintenance of

malaria. In many areas of southern Celebes an undescribed

form of harhirostris has been found with high natural infection rates, and next to Anopheles sundaicus and A, ludlowi

toTahaia it is the most important vector in that area. In

Sumatra and northeastern India natural infection rates are

usually very low, but harhirostris is occasionally important

in Sumatra.

The development of infective stages of Wuchereria hancrofti in harhirostris after experimental feeding has been

demonstrated at Batavia, Java. A, harhirostris is the most

important vector of Wuchereria m^alayi at Kalawara-napuli,

Celebes, where infection rates of up to 8.1 percent have been

reported.

References: 121, 122, 161, 228. (For additional information, see

the descriptions of malaria and filariasis under Malaya and Sunda

Islands.)

Recorded Distribution.—

Assam Indochina

Ban Java

Bangka Malaya

Borneo Moluccas

Burma New Guinea

Celebes Pakistan (East)

Ceram Philippine Islands

Ceylon Soemlba

China (South) Soembawa

Flores Sumatra

Formosa Thailand

Hainan Timor

India

Anopheles bellator Dyar and Knab

Bionomics.—Adults of Anopheles hellator are found in

large numbers in their normal resting places in the forest

canopy 40 to 60 feet above the ground. Toward evening

females leave the forest canopy, fly to nearby centers of human

activity, and bite man readily. Ño critical studies have been

made to determine the amount of zoophilism in this species.
116 AGRICULTURE HANDBOOK 152, U. S. DEPT. OF AGRICULTURE

nor is it known on what animals, if any, hellator feeds in the

forest. Biting takes place almost entirely out of doors, especially in the evening, although some females are active

throughout the day. Even though the adults rarely enter

houses in Trinidad, they appear in rather large numbers

inside habitations in Brazil. After obtaining blood meals,

the females immediately return to the forest or cacao

canopy, where they rest and their eggs develop. Many more

adults are always present in the forest canopy than in biting

places outside the forest.

Larvae are found in water collections in the epiphytic

bromeliads that grow in immortelle trees {Er^thrina), which

are planted in cacao plantations to provide shade for the

rather sensitive cacao. Water in these plants is full of organic

matter and larvae develop rapidly. In Trinidad larvae are

most commonly found in Gravisia aquilega^ a bromeliad

growing from iO to 30 feet above the ground, and even though

they will utilize several species of bromeliads, the removal of

this particular species will destroy practically all the hellator

breeding places. In Trinidad hellator is most active in areas

that receive from about 90 to 110 inches of rainfall annually.

When the rainfall is below or above this amount, the species

tends to disappear, even though breeding places in drier or

wetter areas are still abund.ant enough to support large

populations.

Relation to Disease.—Several times in Trinidad and

Brazil hellator has been easily infected with malaria parasites

and infected females have repeatedly been captured. In

these two countries most endemic malaria in the agricultural

communities near cacao estates is maintained by (1) regular

flights of mosquitoes between houses and forests and (2)

contact of mosquitoes with people who work out of doors in

areas where hellator has even better success in seeking them

out. Bromeliad malaria exists in parts of Trinidad, principally in St. Andrew County, and in Santa Catarina, Paraná,

and Sao Paulo in Brazil. The status of thisspecies in malaria

transmission in areas other than those discussed above is not

known with certainty.

Reference: 61. (For additional information, see the descriptions

of malaria under West Indies, and Brazil and the Guianas. )

Recorded Distribution.—

Brazil

Colombia

Surinam

Trinidad

Venezuela

Anopheles brunnipes (Theobald)

Bionomics.—Anopheles hrunnipes is generally rare, but

when its numbers increase, it may become a dangerous

vector. It is very abundant in some localities, especially at

Pöbane in Mozambique, Sierra Leone, and Leopoldville in

Belgian Congo. It is a species of the cold, dry season. It

has been taken in considerable numTbers in human habitations in Leopoldville and "Aguas Quentes," Mozambique.

Larvae inhabit small, exposed ground pools and slowly

flowing water with surface vegetation and partial shade. Jn

Mozambique large numbers of larvae have been collected in

small pools in an open road drain, where the water was very

fresh, very shallow, and at some distance from human habitations. Although typical of Anopheles gamhiae breeding

places, these pools har'bored none of this species.

Relation to Disease.—Malaria infection rates as low as

1.38 percent have been found in French West Africa. A,

hrimnipes is considered to be of little importance in the

transmission of malaria there. In Leopoldville, Belgian

Congo, on the other hand, a salivary infection rate of about

4 percent was found during the winter. These mosquitoes

are seen in small numbers because of their nondomestic

habits. However, they should be investigated wherever they

are abundant, because other vectors may not be so active in

the winter when this species is common. At present ft is regarded as a secondary vector of some importance.

References: 119, 258, 260, (For additional information, see the

descriptions of malaria under West Africa, West^Central Africa, Madagascar and Adjacent Islands, and Southern Africa.)

Recorded Distribution.—

Angola

Belgian Congo

Cameroons

French Equatorial Africa

French Guinea

French West Africa

Gold C>>ast

Ivory Coast

Mozambique

Nigeria

Northern Rhodesia

Sierra Leone

Southern Rhodesia

South-West Africa

Anopheles christyi (Newstead and Carter)

Bionomics.—In Kenya Anopheles christyi is reported to

be common in cattle sheds and rare in human haibitations,

and in most of Uganda it is entirely nondomestic. However, in the Kigez'i highlands of Uganda it seeks man as a

source of blood meals, because cattle are scarce.

In Kenya larvae are found in borrow pits, pools with

muddy, stagnant water completely exposed to the sun,

ditches, slow-moving streams, pools in drying stream beds,

and at the edges of swamps and seepages. They often live

in small bodies of water used by cattle for drinking. Fouling of this water in moderate amounts by cattle droppings

does not appear to inhibit larval growth. Since christyi tolerates greater pollution than Anopheles gamhiae^ the latter

disappears as pools become more concentrated by evaporation. In Uganda the species lives in the forest in brackish

waterhdles used as wallows by big game. In Tanganyika

its breeding places are usually partially shaded irrigation

channels and small pools that at higher altitudes have little

shade. A, christyi easily tolerates high altitudes within its

range.

Relation to Disease.—This mosquito is not generally considered to be important in the transmission of malaria. In

one malaria study in Kenya none of 336 females were infected, and in another study a large number of females

were shown by precipitin tests to contain blood meals taken

from domestic animals. Furthermore, the seasonal prevalence of the mosquito and the annual cycle of malaria do not

coincide anywhere in Kenya. On the other hand, this species is common in huts in the highland district of Kige2ii,

Uganda, where it was defiriitdy thought at one time to be a

vector in a malaria epidemic. However, subsequent studies

showed that because both gamhiae and Anopheles funestus

were present, the part played by christyi in this outbreak is

in some doubt. It is not known Whether christyi can carry

the disease elsewhere, but the posábility is slight. This

species is included in the keys as a vector because of its

possible role in the Kigezi hig'hlands malaria epidemic.

References: 69, 119, 260. (For additional information, see the descriptions of malaria under Eritrea, the Somalilands, and Ethiopia;

West-Central Africa ; East-Central Africa ; and Madagascar and Adjacent Islands.)

Recorded Distribution.—

Belgian Congo Somalia

Ethiopia Tanganyika

Kenya Uganda

Madagascar

Anopheles clauiger (Meigen)

In early Italian papers Anopheles claviger was a name

occasionally applied to A, maculipennis. True claviger has

almost universally been called Anopheles hifurcatws in early

European literature.

Bionomics.—In Sardinia claviger is largely sylvatic and

seldom bites man. Few females have been found in domestic shelters. About one-half of those captured were
MOSQUITOES OF MEDICAL IMPORTANCE 117

from natural hollows and about one-fourth from vegetation.

However, in Israel, this species is very definitely domestic,

and in Syria and Lebanon both man and animals are attacked night and day, indoors and out. In some localities

man is used as a source of food only when breeding places

are close by. In Europe, Caucasia, and central Asia the

adults are common early in the spring and again late in the

fall, particularly in Sardinia, Italy, and Albania. Over

much of its range claviger lays eggs without a blood meal,

whereas forms requiring blood for egg laying have been observed on the Black Sea coast.

In England and other localities at similar latitudes larvae

live in permanent ponds and along lake margins in the shade.

However, in more southern areas they are restricted to cool

spring-fed pools, which in much of Asia are the characteristic breeding places. In Sardinia, springs and associated

rivulets, streams, tanks, wells, waterholes, ground pools, and

swamps are common habitats. Larger bodies of water are

preferred during the winter and smaller, colder ones in the

spring. In Caucasia and central Asia the temperature of

the water in the summer is elevated to a point where larval

populations are decidedly decreased. In general, breeding

waters are cool, clear, and shaded. Water may be standing

or running and shaded or exposed to the sun. Larvae have

a habit of staying submerged for an average of about 1 minute when they are disturbed. They have been observed to

stay under the surface for as long as 14 minutes. In Israel,

Jordan, Syria, and Lebanon larvae are found in cisterns

underneath or adjacent to man's habitations.

A, claviger is adapted to the low temperatures common to

high latitudes, usually passing the winter as larvae, but in

Sardinia the mosquito breeds throughout the year, even

thoug'h the adults develop fat bodies. A. claviger has been

found at altitudes of about 4,300 feet and in Morocco and

Turkestan at about 9,200 feet.

Relation to Disease.—Over most of its range claviger is

not important in malaria transmission, since it is either

scarce or not attracted to man. It is important in Israel

because of its habit of breeding in cisterns and other domestic water supplies. It is of local importance in villages

on the central plain and coastal areas of Syria and Lebanon,

where small amounts of malaria are transmitted. In the

suburibs of Baku, Caucasia, where mountain springs stay

cool enough through the summer to maintain large populations of the vector in close proximity to man, the species

supports endemic malaria.

References: 67, 229, 239. (For additional information, see the descriptions of malaria under North Europe and North Asia, the Near

East, and Afghanistan and West Pakistan. )

Recorded Distribution.—

Albania Ireland

Algeria Israel

Austria Italy

Balearic Islands Jordan

Belgium Lebanon

Bulgaria Libya

Caucasia Morocco

Corsica Netherlands

Crete Poland

Cyprus Portugal

Czechoslovakia Rumania

Denmark Sardinia

Egypt Sicily

Elba Spain

European U. S. S. R. Spanish Morocco

France Sweden

Germany Switzerland

Great Britain Syria

Greece Tunisia

Hungary Turkestan

Iran Turkey

Iraq Yugoslavia

Anopheles crucians Wiedemann

Bionomics.—Although the females of Anopheles crucians

may enter houses readily and bite man freely, they are primarily outdoor biters on porches, in outhouses, and in similar

locations. Studies on the host preference of crucians have

shown it to be more zoophilic than anthropophilic. This

habit probably varies from place to place according to the

availability of hosts. A flight range of 7,000 to 9,000 feet

has been observed.

Larvae frequently are found with those of Anopheles

quadrimaculatus but, unlike that species (see p. 133), crucians

is predominantly an acid-water breeder. In the Southern

United States larvae are found in the complete absence of

quadrimaculatus in dense cypress swamps and coastal-plain

ponds. Their favorite habitats are impoundments, blocked

streams, seepages, and occasionally wheel ruts, temporary

pools in pine forests, and other accumulations of water without vegetation. In Jamaica larvae have been collected from

streams, seepages, and ponds with sunlit or lightly shaded

water, but not in brackish water. In the United States

crucians larvae are found in breeding places all winter long

and have even been taken from mud at the bottom of icecovered water.

Relation to Disease.—Because of its high densities along

the coastal plain of the Southern United States, its houseentering habits, and its occasional anthropophilic tendencies,

crucians has been considered a potential vector of malaria

in the Southeastern United States, even though it is often

present in large numbers in the absence of appreciable malaria. It is probably important in Cuba, but farther east its

role as a vector is not known.

References: 67, 292, 308, 349. (For additional information, see the

descriptions of malaria under America North of Mexico, West Indies,

and Mexico. )

Recorded Distribution.

British Honduras

Cuba

Dominican Republic

Guatemala

Haiti

Honduras

Jamaica

Mexico

Nicaragua

Puerto Rico

United States

Anopheles cruzi Dyar and Knab

Bionomics.—Anopheles cru^i inhabits the rain-forest

canopy, living in greatest numbers at some height above the

ground. In Santa Catarina, Brazil, large numbers of females fly to houses in the outskirts of the towns of Brusque,

Joinvile, Sao Francisco, and Florianopolis, which adjoin

rain forests, where they have been observed biting man in

extremely large numbers. A. cruzi larvae have habits and

breeding places similar to those of hellator (see p. 115).

Relation to Disease.—As long ago as 1903 cruzi was incriminated as a vector of malaria in the mountain forests of

Sao Paulo, Brazil, where railroad laborers were contracting

a considerable amount of malaria. Not long afterward cru^i

was found naturally infected with malaria parasites in Paranagua, Brazil. In 1942, 1943, and 1944 it was again found

naturally infected, and it was easily infected experimentally

with malaria parasites. It is now regarded as an important

vector.

References : 61, 237. ( For additional information, see the description of malaria under Brazil and the Guianas. )

Recorded Distribution.—

Brazil Panama

British Guiana Peru

Costa Rica Surinam

Ecuador Venezuela

French Guiana
118 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Anopheles culicifacies Giles

The name commonly employed in the early literature for

Anopheles cvlicifaieies was A. listonii^ but culicifacies has

been used in recent important literature relating to disease

transmission.

Bionomics.—Adults of culicifacies rest in large numbers

in small dark spaces in houses and cowsheds during the day.

In some localities females are zoophilic, feeding actively on

cattle and birds at dusk, but in most areas they prefer man

as a host. Flight ranges from 1% to 1% miles have been

observed.

Fresh, clean water attracts most ovipositing females, although immature stages occasionally are found in brackish

water. The usual breeding sites are fresh water with grassy

edges, such as irrigation ditches, seepages, pools, slow-moving streams, canals, riverbeds, freshly formed collections of

rainwater in borrow pits, fallow ricefields, shallow tanks,

pits^ and wells. The species prefers sunlight or light shade,

avoiding dense shade, for oviposition.

A, culicifacies is ordinarily a plains species, but it is found

at moderate altitudes in the Himalayas, having been recorded up to 6,500 feet. In India it overwinters as a larva

and is most prevalent from May to November in the northern

parts of that country.

Relation to Disease.—This species is one of the most important malaria vectors wherever it appears. It is probably

an important mosquito east and south of the Iranian deserts,

in the Punjab of West Pakistan, and in parts of Afghanistan, East Pakistan, Assam, South China, and Thailand.

It is of primary importance in India, Ceylon, and Burma.

In many localities in South China it is too rare to transmit

the disease.

References: 8-^, 88, 91, 122. (For additional information, see the

descriptions of malaria under Iran, Afghanistan and West Pakistan,

India and Ceylon, East Pakistan and Assam, South China, Burma,

and Thailand.)

Recorded Distribution.—

Afghanistan

Arabia

Assam

Bahrein

Burma

Ceylon

China (South)

India

Indochina

Iran

Pakistan (East and West)

Thailand

Anopheles darlingi Root

Bionomics.—Anopheles darlingi Root is one of the most

domestic mosquitoes in the New World. It feeds readily on

man and enters human habitations in large numbers. In

studies in the rural areas of Stann Creek Valley and the

Toledo District of British Honduras, about 72 percent of all

mosquitoes caught in houses were darlingi; only 27 percent

captured at sunset on horses were this species. In another

locality three-fourths of the Anopheles species inside houses

near beds and in dark places were darlingi^ whereas not one

was caught on a horse used as bait just outside. High

humidity and rainfall are primary requisites for adult

survival.

Larvae are commonly found in mats of surface vegetation

in bodies of water that lack appreciable current or waves.

They live in shaded water with low salt content and neutral

or slightly acid reaction. Some larvae are specially adapted

to floodwaters. In the Amazon Valley the dry season is extensive enough to eliminate the species from large areas, during which time it breeds in swamps or other large bodies of

deep, still water that are not affected by dry weather. In

Brazil larvae live at the shaded edges of impoundments,

where wave action is damped by branches of trees extending

into the water, and they do not live in ditches, small pools,

or swamps. In British Honduras larvae are common along

deeply shaded stream edges.

Relation to Disease.—^This species is the most efficient

indigenous malaria vector in north and northeast Brazil. In

the Amazon Valley it causes small regular epidemics of malaria, a disease that is never found without darlingi in this

area. Because this species is closely associated with malaria

in Venezuela and British Guiana, it is extremely important

in those localities and in most others where it and malaria

appear together.

References : 19, 292, 308. ( For additional information, see the descriptions of malaria under Mexico, Central America, Panama, Northwest South America, Brazil and the Guianas, and Lower South

America. )

Recorded Distribution.—

Argentina Guatemala

Bolivia Honduras

British Guiana Mexico

British Honduras Peru

Colombia Surinam

Ecuador Venezuela

French Guiana

Anopheles domicolus Edwards

Bionomics.—Anopheles domicolios is a localized species.

In studies in northern Nigeria it represented 0.25 percent of

all Anopheles species caught in human habitations. Breeding places of the larvae are not known.

Relation to Disease.—This mosquito is regarded as a subsidiary vector of malaria in French West Africa and HauteVolta, where 3 of 283 and 1 of 44 females, respectively, were

found to harbor sporozoites.

References: 119, 258, 260. (For additional information, see the

description of malaria under West Africa, East-Central Africa, and

Southern Africa.)

Recorded Distribution.-

Anglo-Egyptian Sudan

French West Africa

Gold Coast

Ivory Coast

Nigeria

Northern Rhodesia

Sierra Leone

Southern Rhodesia

Uganda

Anopheles d'thali Patton

Originally described from the Aden hinterland, Anopheles

dHhali has been discussed in some literature under the name

A, rhodesiensis. However, the rhodesiensis species of Africa

is entirely different (p. 133).

Bionomics.—Females of dHhali feed readily on man.

They are often found in tents, barracks, and houses, and

they also rest in cowsheds. Adults have been found I14

miles from their nearest breeding places in Arabia.

Larvae inhabit springs, wells, and riverbed pools, usually

with moderate shade and some floating vegetation. In India

deep riverbed pools full of water plants and algae are preferred breeding places. In Yemen larvae are found in the

quiet marginal water of drying streams, with emergent short

vegetation and filamentous green algae. In many parts of

Arabia they are commonly found in holes in volcanic rocks

fed by underground water and in underground aqueducts

and tanks. In Sinai the species shows little preference for

any one type of breeding place. Larvae have been found in

all kinds of water collections, including stagnant weedy

pools, small swiftly flowing streams, fresh water moving

slowly over grass, and in weedy pools near fresh-water

streams. In British Somaliland seepages, footprints, and

waterholes are inhabited by these larvae, although they are

usually rare when Spirogyra is present.

In India dHhali is prevalent from July to September and

at Khajraho it has been found at an altitude of approximately 2,000 feet. It is a highland and plateau species in

Jordan.

Relation to Disease.—In Arabia and Sinai dHhali is suspected of being a vector of malaria. It was the most preva-
MOSQUITOES OF MEDICAL IMPORTANCE 119

lent species when 100 percent of the personnel of a military

post became ill with malaria. It is believed to be a vector

in northern Ethiopia and British Somaliland.

References: 67, 91, 120, 206, 239. (For additional information,

see the descriptions of malaria under Northwest Africa ; Egypt ;

Eritrea, the Somalilands, and Ethiopia ; and Arabia.)

Recorded Distribution.—

Algeria Iraq

Anglo-Egyptian Sudan Israel

Arabia Jordan

British Somaliland Libya

Egypt Morocco

Eritrea Pakistan (West)

Ethiopia Somalia

India Tunisia

Iran

Anopheles farauti Laveran

Throughout most of the literature Anopheles farauti has

been represented as J.. punctulatus var. moluccensis. In 1946

farauti was definitely determined to be a full species.

Bionomics.—Females are most active at night but have

been observed to feed during the day if the sky is overcast.

In some areas they are seldom found in native huts during

the day, whereas in others they enter houses in considerable

numbers in the daytime. In most localities females will feed

on man to the complete exclusion of livestock. In the New

Hebrides they feed readily on horses and cattle even when

man is present. Adults rest in heavy stands of kunai grass

and are found close to the ground on the stems. They are

rarely observed close to their breeding places.

In the Solomons and New Hebrides the dry-season breeding places are rivers, streams, ponds, lagoons, taro gardens

under water, roadside ditches, and occasionally w^ells and

animal wallows. During the wet season larvae live in almost

all types of manmade depressions filled with fresh rainwater.

In Guadalcanal, eggs, larvae, and pupae are commonly

flushed by heavy rains from permanent breeding places, such

as ruts, foxholes, bomb craters, trenches, and borrow pits, to

the flat alluvial plains of the northeast coast. In northern

Queensland the species breeds in lagoons and potholes along

riverbanks. Larvae are found in pure, brackish, or rather

stagnant rainwater, where vegetation may or may not be

present. They have been taken from heavily shaded mangrove areas, but open sunlight is apparently preferred. They

are rarely found in coconut shells. The aquatic cycle usually

lasts from 13 to 15 days.

Relation to Disease.—Because of its high susceptibility

to infection and its domesticity, farauti is the most important vector of malaria in Australasia. In the Moluccas,

New Guinea, Admiralty Islands, Bismarck Archipelago,

northern Australia, and the Solomon Islands this species is

especially important. It is said to be the only vector in the

New Hebrides, and natural infection rates in New Guinea

are very high. Malaria and the distribution of farauti are

bounded by longitude 170° E. and latitude 20° S. The

reasons for these limits are as yet unexplained.

In the Solomons-New Hebrides area farauti is an important vector of filariasis. Almost 14 percent of this species

was found infected with Wuchereria ha/ncrofti in the New

Hebrides and over 50 percent in Guadalcanal.

References: 61, IOS, 122, I46, .294, S24. (For additional information, see the descriptions of malaria and filariasis under Australasia. )

Recorded Distribution.—

Admiralty Islands

Australia (Northern Territory, Queensland)

Bismarck Archipelago

Ceram

Moluccas

New Britain

New Guinea

New Hebrides

New Ireland

Solomon Islands

Anopheles filipinae Manalang

Anopheles filipinae belongs to the so-called funestus-minimus subgroup of Anopheles (p. 120) and was originally

known as a variety of A, minimus.

Bionomics.—Almost nothing is known about the habits of

filipinae adults. Flight-range experiments show that

stained adults may travel as far as 3,300 feet from the point

of release.

Larvae inhabit small, slowly flowing, spring-fed streams,

grassy margins of canals, clear or muddy water of streams,

flowing irrigation ditches, pools, lakes, and other impoundments. They are occasionally associated with aquatic vegetation, such as Pistia^ Ipomoea^ and water hyacinths.

Relation to Disease.—Large numbers of filipinae are

found in northern Luzon, where malaria has been known to

occur. In central and southern Luzon the species is not at all

common. The natural infection rate has, in general, been

very low, and as far as is known no experimental infections

have been performed.

References: 61, 120, 204, S08. (For additional information, see

the description of malaria under Philippine Islands. )

Recorded Distribution.—

Philippine Islands

Anopheles fluviatilis James

In many early references Anopheles fiuviatilis was called

A. listonii and A, christophersL

Bionomics.—Females of fiuviatilis appear to prefer

houses as nighttime resting places but have also been found

in considerable numbers in cowsheds. Two subspecies of

fiuviatilis may exist in India. One form in the south is

strongly anthropophilic and feeds readily on man ; the other

m the north bites cattle almost exclusively. The southern

form feeds from II/2 hours after dusk to midnight. Most

of the females rest out of doors during the day. In a study

in India few adults were found in villages beyond 2,600 feet

from their breeding places. Once they were reported at a

greater distance.

Larvae inhabit pools in stream beds, slow-flowing water

with vegetation, leaks from springs and irrigation ditches,

swamp edges, lake margins, drains, ponds, and tanks. Most

of the streams in which this species is found have grassy

edges. In Yemen fiuviatilis was observed at an elevation

of about 7,300 feet in natural pools along a ditch with stonewort rushes and filamentous green algae. Although sometimes found in ricefields, the species does not breed prolifically there. Larvae have never been observed in large

numbers. Dispersed breeding over immense areas is responsible for the importance of this species.

Widely distributed throughout India, especially in foothill areas and hilly or rocky areas, this species is more common in the west than in the east. At Peshawar, West Pakistan, this species has been observed from May to November,

with population peaks in May and June and again in November. It has been found at an elevation of 6,000 feet at

Kashmir and as high as 7,500 feet at Murree. In southern

India it is not normally found below 1,000 feet.

Relation to Disease.—Undoubtedly this is a very important vector of malaria. Although few experimental infections have been reported in the literature, fiuviatilis has been

found naturally infected in many areas of India. It is considered to be of greatest importance in peninsular India, is a

suspected vector in Arabia, and transmits malaria in the

foothills of West Pakistan.

References: 61, 91, 120, 206. (For additional information, see the

descriptions of malaria under Arabia, Afghanistan and West Pakistan,

and India and Ceylon.)
120 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Recorded Distribution.—

Afghanistan

Arabia

Assam

Burma

Ceylon

China (South)

India

Indochina

Iran

Iraq

Pakistan (East and West)

Thailand

Turkestan

Anopheles freeborni Aitken

Bionomics.—Anopheles freeborni residïly enters homes and

animal shelters, biting actively through the night. At the

end of the dry season fertile females migrate as far as 12 miles

and settle in outbuildings, homes, and cellars. They apparently do not stay in one place but move from one spot to

another throughout the winter. Since a single female may

bite several persons without taking a full blood meal at any

one time, it can transmit malaria during the winter, when it

lives almost entirely inside houses and sheds, biting near the

floor. These females have well-developed fat bodies, and

even when they take blood meals, usually no eggs develop

during the winter months. When the females leave these

winter haunts for summer breeding, they bite viciously in

full daylight and then lay eggs. In midseason, adults rarely

fly farther than 1 mile from their breeding places.

Adults oviposit in all kinds of water collections. Many of

these dry out as the summer progresses. Larvae live in permanent or semipermanent water collections, as well as in

seepage areas, vegetated borrow pits, hoofprints, carelessly

irrigated fields, and edges of streams and canals where vegetation protects them from water movement. Larvae prefer

water that is clean, clear, and slightly alkaline, and they

swim to lightly shaded sunlit places with flotage, algae, or

emergent vegetation. Although freeborni will breed in

brackish water, ittends to avoid sew^age pollution and organic

material. Favorite habitats are ricefields in which the water

remains fairly fresh.

A, freeborni is a mosquito of semiarid regions having

seasonal rainfall. Peaks of adult abundance occur during

the early summer and at the end of the dry season in the fall.

Relation to Disease.—This mosquito has been regarded

for years as the important malaria carrier in the Western

United States, and it may play a small part in the transmission of that disease in Lower California and on the northcentral plateau of Mexico.

Reference : 61. ( For additional information, see the descriptions of

malaria under America North of Mexico and Mexico.)

Recorded Distribution.—

Mexico

United States

Anopheles funestas Giles

Anopheles funestus has been known by various names, because it is present in most of tropical Africa, appears in such

large numbers, and is so important. It is very similar to

Anopheles minimus and its relatives in the Orient. A, funestus imerinensis is the only medically important form of

funestus in Madagascar.

Bionomics.—Precipitin tests almost always show that

more funestus females have fed on man than on cattle. Females most commonly bite indoors, and experiments show

that they are attracted to man by his odor. In one part of

Africa females w^ere found to enter native huts from 8 p. m.

to about 2 a. m. and in another part between 2 and 6 a. m.

A large number of females remain in houses, where they feed

until after dawn. Under normal conditions adults are not

likely to be found in large numbers beyond one-half mile

from their nearest breeding places, but under unusual conditions they may fly as far as 4 miles.

Two kinds of breeding places may be frequented by the

larvae. The most characteristic of these is large and rather

permanent, and it includes swamps, lakeshores, and ponds.

The other larval habitat comprises smaller bodies of water,

such as hill streams, furrows, ditches, and seepages, and

occasionally wells and artificial containers. Females sometimes seek these smaller water collections when no large bodies

of water are available to support the large populations that

are sometimes found. Floating vegetation such as Pistia^

Eichhornia^ Marsilea^ low grass, and rice provide optimum

breeding conditions for the larvae, which are killed more

easily by high temperatures than are those of Anopheles

gamhiae.

Where breeding takes place in large permanent bodies of

water, the populations of funestus are more constant throughout the year than those of gamhiae. In hilly regions where

the former species breeds in streams, the populations tend

to fluctuate with flushing rains.

Relation to Disease.—This is one of the two most important vectors of malaria in Africa. In contrast to gambiae^

whose seasonal variation in abundance adapts it to the transmission of much of the epidemic malaria in Africa, funestus

fluctuates relatively little in numbers because of its habit of

breeding in large bodies of water, and it is responsible for

severe endemic malaria. Large populations of funestus have

been reported several times in the absence of malaria, possibly

because of the presence of biological races that cannot transmit the malaria parasite.

Since developing stages of Wuchereria bancrofti have

often been found in funestus^ this species is probably an important vector of filariasis in areas where it is locally

abundant.

References: IJfS, 258. (For additional information, see the descriptions of malaria and filariasis under Anglo-Egyptian Sudan; Eritrea,

the Somalilands, and Ethiopia; West Africa; West-Central Africa;

East-Central Africa ; Madagascar and Adjacent Islands ; and Southern

Africa.)

Recorded Distribution.—

Africa (except Cape Province and Orange Free State in the

Union of South Africa, and French Somaliland)

Madagascar

Mascarene Islands

Mauritius

Anopheles gambiae Giles

The name most commonly used in the past for Anopheles

gambiae was A, costalis. Fortunately gambiae is a distinctive mosquito, and it may often be identified in old records

by its excellent ability to transmit malaria.

Bionomics.—A, gambiae is highly anthropophilic. Most

precipitin tests demonstrate that more than 50 percent of females contain human blood, and they are nearly always attracted to man in preference to animals. Their manner of

entering and leaving houses varies from area to area. In

some localities this species may be entirely domestic, in some

it may enter houses only at night and leave before morning,

and in others it may not enter houses at all. It prefers the

darkest houses, and when inside it tends to find the darkest

corners. Adults usually enter houses from 11 p. m. to 5 a. m.

Females are commonly found in outdoor resting places and

sometimes bite in the open, where they are most active after

midnight. They have a peak of biting activity between 2

and 4 a. m. Sometimes they will bite outside during the day

where no great changes in temperature or humidity occur

during a 24-hour period. Females are able to travel long

distances, having been caught as far as 4i/4 miles downwind

from their breeding places. Normally they fly an average

of about 1 mile without assistance from the wind.

The most important larval breeding places are manmade

and are therefore close to man's habitations and places of

work. Where gambiae is abundant, the larvae are usually
MOSQUITOES OF MEDICAL IMPORTANCE 121

found in borrow pits, drains, excavations, hoofprints, and

even artificial containers in areas with denuded and eroded

soil resulting from overstocking, roadmaking, deforestation,

swamp reclamation, and veld burning. Important larval

habitats also include overflow of rivers, backwaters, pools

left by receding rivers, and rainwater in natural depressions.

Larvae are found the year around in permanent breeding

grounds in frost-free areas. In the warm parts of the year

gambiae overflows into areas where it cannot exist during

the winter. Larvae can withstand considerable extremes of

water temperature. Their breeding places are generally exposed to the weather and lack vegetation. The time of development from larva to adult, usually about 6 days, varies

considerably with temperature and humidity.

Larval and adult populations are correlated with rainfall.

An increase at the beginning of the rainy season is followed

by a later reduction, sometimes before the dry season begins.

Normal breeding places are often flooded by heavy rains to

the point where they become unsuitable for larval habitats,

and as a result the adult populations are reduced.

Relation to Disease.-^This species is regarded as the most

important vector of malaria throughout Africa, because it

greatly prefers human blood and does not discriminate in

its choice of breeding places and because parasites can mature exceptionally well in gamhiae. In malarious areas

salivary gland infection rates are always high. It is associated with endemic as well as epidemic malaria, and it has

often been found in the complete absence of other possible

vectors. Even when other vectors are present, gambiae is

nearly always the most important one. It is the only vector

in Africa that exists in the presence of endemic malaria at

altitudes over 6,000 feet.

In Liberia gambiae has been found with high rates of

natural infection with Wuchereria barwroftL In at least

one coastal locality gamhiae is believed to be the principal

vector of filariasis.

References : IJfS, 258. (For additional information, see the descriptions of malaria and filariasis under Brazil and the Guianas ; Egypt ;

Anglo-Egyptian Sudan ; Eritrea, the Somalilands, and Ethiopia ; West

Africa; West-Central Africa; East-Central Africa; Madagascar and

Adjacent Islands ; Southern Africa ; and Arabia.)

Recorded Distribution.—

Africa (all tropical countries)

Arabia

Madagascar

Mascarene Islands

Mauritius

Anopheles hancocki Edwards

Bionomics.—Anopheles hancocki readily bites man.

Throughout much of its range, especially in Uganda, it commonly feeds inside houses, although occasionally human

dwellings have been reported to be free of adults. However, in Sierra Leone no adults have ever been caught

indoors.

In Uganda the larvae have been observed most commonly

in clear,^ slowly running water in ditches and wells containing Pistia, In Sierra Leone immature stages have often been

found at stream margins and in seepage water.

Relation to Disease.—Since malaria transmission by hancocki has always been studied in locations with large populations of Anopheles gambiae^ the true value of the former

species as a vector is difficult to determine. For example, in

Uganda up to 9 percent of hancocki females were found infected, but the species was present with large numbers of

gambiae^ whose infection rates were even higher. Therefore, it is regarded as a secondary vector in Uganda. A.

hancocki is considered to be a subsidiary vector in Ivory

Coast and Haute-Volta, and it must be regarded as a vector

of some importance wherever it exists.

This species, appearing for short periods of time in large

numbers, has been found infected with larval stages of

Wuchereria bancrofti, and in at least one locality in Liberia

it IS believed to be a rather important vector of filariasis.

References : 69, 119, US, 260. (For additional information, see the

descriptions of malaria under West Africa, West-Central Africa, EastCentral Africa, Madagascar and Adjacent Islands, and Southern

Africa.)

Recorded Distribution.—

Angola Ivory Coast

Belgian Congo Kenya

Cameroons Liberia

Dahomey Madagascar

French Equatorial Africa Nigeria

French West Africa Sierra Leone

Gold Coast Uganda

Anopheles hargreavesi Evans

Bionomics.—Anopheles hargreavesi is both endophilic and

exophilic, but there are no records of its using animals as a

source of blood meals. In Nigeria it is common in native

huts and bites man freely out of doors as well, with a peak of

activity^ at midnight or later. As many females of hargreavesi as of Anopheles gambiae were attracted to human

bait during all-night catches in the vicinity of Lagos,

Nigeria. At Itowolo, southern Nigeria, biting-activity

studies showed that the number of flying hargreavesi females

increased rather rapidly at twilight, continued to increase

until midnight, and fell rather sharply near dawn.

Larvae were found to be associated with Pistia in two

locations in Nigeria. One was a collection of foul, sewagecontaminated water and the other a clear-water pool in an

open jungle area. Larvae also are found among blades of

grass in open swamps.

In Abeokuta, Nigeria, breeding is practically nonexistent

during the dry season but increases soon after the onset of

the rains.

Relation to Disease.—This species is of considerable importance as a vector of malaria. It has been successfully infected in the laboratory, and in southern Nigeria it has been

found with high sporozoite rates in nature. Because it appears in larger numbers and has a more uniform seasonal distribution than gambiae^ it may be a more important vector of

malaria than that species in Itowolo, Nigeria, and it may

be of considerable importance in other parts of West Africa

where it is common.

References: 119, 251, 258, 260. (For additional information, see

the descriptions of malaria under West Africa, West-Central Africa,

East-Central Africa, and Southern Africa. )

Recorded Distribution.—

Angola Ivory Coast

Belgian Congo Liberia

Cameroons Nigeria

Dahomey Sierra Leone

French Equatorial Africa Uganda

Gold Coast

Anopheles hispaniola (Theobald)

Many early records of Anopheles hispaniola are found

under the name A. turkhudi^ a very close relative whose

distribution is believed to be principally within the Orient.

It is not known whether the range of turkhudi extends w^estward to meet or overlap that of hispaniola.

Bionomics.—A, hispaniola is said to be domestic in some

parts of its range, but it is wild and found only rarely in

houses in others. During studies in Sardinia about one-half

of the hispaniola adult collections were from domestic shelters, the remainder from mine shafts, grottoes, and vegetation. Adults were rarely found near breeding places containing large numbers of larvae.

Larvae are most commonly found in shallow, clear, sunlit

water, usually in small pools but occasionally in moving
122 AGRICULTURE HANDBOOK 15 2^ U. S. DEPT. OF AGRICULTURE

water. Spirogyra commonly grows in these breeding places,

sometimes to the complete exclusion of other plants or algae.

Larvae live in ricefields in the vicinity of Calasparra, Spain,

although that type of breeding place may not be common

elsewhere. In Algeria the species breeds in stream beds, and

in at least one area where the flow of water was low and

rather regular, larvae were recovered from an irrigation

channel with considerable vegetation growing along its edge.

In Jordan larvae w^ere found in streams, springs, ditches, and

seepages, in sunlight or partial shade. Experiments in Sardinia showed that larvae can remain submerged for as long

as 35 minutes.

In several localities in Spain this species was encountered

at elevations of about 3,300 feet to 4,300 feet and in Morocco

up to 8,200 feet. At least in Sardinia hispaniola is a latesummer and fall species, and it has been found in even the

driest part of Algeria.

Relation to Disease,—In much of Spain this species is

found with Anopheles labranchiae atroparvus^ so that its

true role as a malaria vector is not clear. However, in

Tenerife and in other parts of the Canaries, hispaniola is the

only Anopheles species that bites man in the presence of

active malaria, and it may be presumed to transmit malaria

elsewhere as well. It is regarded as having some importance

in Morocco and in northern Algeria, where it is reported as

a secondary vector. It is not regarded as a serious vector of

malaria in Sardinia, where the species is relatively scarce

and rarely bites man.

References : 29, 67, 237, 239,329, 336, 339. (For additional information, see the descriptions of malaria under Northwest Africa, and

Spain and Portugal.)

Recorded Distribution.

Algeria

Arabia

Canary Islands

Egypt

Greece

Iraq

Italy

Jordan

Libya

Morocco

Sardinia

Spain

Spanish Morocco

Tunisia

Anopheles homunculus Komp

Bionomics.—The town of Blumenau in Santa Catarina,

Brazil, is situated in the immediate vicinity of tall rain

forests. Adults of Anopheles homunculus ordinarily inhabiting the lower parts of the forest canopy where it is very

humid have been observed flying in large numbers day and

night into the town, where they bite man readily. In Trinidad the adults live only in the extremely humid and deep

Aripo Valley, which has an annual rainfall of over 125

inches. Here the females occupy the lower 30 feet of cacaoplantation and forest canopy and fly and bite outside the

forest on very humid nights. Adults are rarely found in

areas with less than 80-percent average relative humidity,

and then they are restricted almost entirely to the forest.

For information on larval bionomics, see page 29.

Relation to Disease.—This species may be a vector of

malaria in Brazil, because it lives near human habitations in

large numbers and bites man readily. So far as is known it

has never been found infected in nature.

Reference : 67. (For additional information, see the descriptions of

malaria under West Indies, and Brazil and the Guianas. )

Recorded Distribution.

Brazil

Colombia

Trinidad

Venezuela

Anopheles jeyporiensis candidiensis Koidzumi

Originally described from Formosa, Ano2?heles jeyporiensis candidiensis has been given its present name throughout

most of the important literature. It has also been referred

to as Anopheles aconitus var. tonkinensis. No attempt has

been made to distinguish it from the type form in many of

the early records.

Bionomics.—Females of candidiensis bite man readily inside houses and tents, starting about 2 hours after nightfall.

After feeding they rest outside and are seldom found in

human habitations during the day.

Typical breeding sites, especially in Indochina, are those

with moving, grass-margined w^ater. In India and China

larvae are found in the grassy, shallow water in seepage areas

on hillsides and in abandoned ricefields and at the grassy

margins of lakes and swamps. Along the Bengal-Burma

border larvae live in collections of w^ater so thickly overgrown with vegetation that they cannot b3 readily seen.

In Assam and Burma this mosquito appears in greatest

numbers during the early months of the year.

Relation to Disease.—In hillock parts of the Arakan area

along the East Pakistan-Burma border most of the premonsoon malaria is transmitted by candidiensis. In southwestern Yunnan and Indochina this mosquito is considered

to be next or equal to Anopheles minimus in importance as a

vector of foothill malaria. Although in China almost all

dissections of the females for malaria have been positive,

candidiensis is considered to be of only secondary importance

in the hilly areas. It has been found with high natural

infection rates in Hong Kong.

With natural infections of Wuchereria iancrofti of up

to 4.6 percent, candidiensis is an important vector of filariasis

at Hong Kong. Natural infections with that parasite have

also been reported from Indochina.

References: 88, 91, 122, 2Iß. (For additional information, see the

descriptions of malaria and filariasis under East Pakistan and Assam,

South China, Burma, and Indochina. )

Recorded Distribution.

x\ssam

Burma

China (South)

Formosa

Hainan

India

Indochina

Pakistan (East)

Anopheles kochi Dönitz

Bionomics.—Females of Anopheles kochi feed readily on

man but appear to prefer the blood of cattle. In some areas

they bite man with some reluctance. In Assam adults rest

during the day in the jungle, but in other areas they are

found more commonly in open places, such as on shaded

stone walls, under the overhanging banks of streams, and

in houses and stables. In Java they habitually enter human

habitations at twilight. Males were observed to swarm 4

to 6 feet above the head of a man in Malaya.

Larvae live in various places containing either turbid or

fresh water but are never found in salt or brackish water.

They prefer open sites, such as small muddy pools, flowing

irrigation ditches, stagnant drains, jungle pools with decaying vegetation, buffalo wallow holes, fallow ricefields,

hoofmarks, ruts, and other depressions in the ground. In

the Philippines larval breeding is common in cut bamboo

and artificial containers, although presumably suitable habitats situated nearby remain free of larvae.

A. kochi, because of the nature of its breeding places,

reaches a peak of abundance during the rainy season.

Relation to Disease.—Experimental infections show this

species to be a potentially efficient vector of malaria. It

appears to be of greatest importance at times when its populations are high or when livestock is absent. High natural

infections in the Greater Sundas, especially Sumatra, indicate it is of some importance there. In Java it is only

locally important and in Celebes does not usually play

a part in malaria transmission. Natural infection rates are

almost always so low in the Philippines, Malaya, Indochina,
MOSQUITOES OF MEDICAL IMPORTANCE 123

and most of India that the species is believed to be of little

or no importance in these countries.

References : 91, 121, 122, 161, 3^1. (For additional information, see

the descrii)tions of malaria under Sunda Islands.)

Recorded Distribution.—

Assam Indochina

Bali Java

Bangka Malaya

Borneo Moluccas

Burma Pakistan (East)

Celebes Philippine Islands

Ceram Soemba

China (South) Soembawa

Flores Sumatra

Formosa Thailand

Anopheles koliensis Owen

Anopheles koliensis is morphologically intermediate between Anopheles faraiiti and A, punctulatus. Its medical

importance was discovered during World War II, but the

form has been known long enough so that confusion with

other vectors has been avoided.

Bionomics.—Females of koliensis are strongly anthropophilic and tend to remain in human habitations after feeding,

leaving some time during the following day or night. They

rest during the day in such locations as tentpoles and dark

bedding in native huts and appear in greater numbers than

either farauti or punctulatus. Ninety percent of the females

taken in the tents of a native-labor camp over a period of

several weeks in one locality in Guadalcanal were koliensis

and 10 percent farauti and punctulatus. During this time

larvae of koliensis constituted about 10 percent of the total

Anopheles species present. Females become active about

9 p. m. and continue to fly until daylight, w^ith the greatest

activity after midnight.

Temporary pools in grassland at the edges of jungles are

most commonly frequented by larvae. They are found more

often in water exposed to the sun than in dense jungle shade.

During the dry season the species is relatively scarce. It is

almost always associated with farauti^ at least in Guadalcanal.

Relation to Disease.—A. koliensis is probably of major importance in malaria transmission on the northeast coast of

Guadalcanal.

References: 56, 57, 271, 294, 353. (For additional information, see

the description of malaria under Australasia. )

Recorded Distribution.—

New Britain

New Guinea

Solomon Islands

Anopheles labranchiae atroparvus Van Thiel

Anopheles lahranchiae atroparvus^ an important member

of the A. macuUpennis group in northern Europe, was originally distinguished from A. messeae^ a close relative, on the

basis of its shorter wings and darker color. Later studies

revealed its relationship to Anopheles labranchiae. The

names Anopheles fallaw and A. camboumaci apply only to

local variations of this mosquito. For additional details,

see page 125.

Bionomics.—Although definitely anthropophilic, atroparvus over extensive areas is effectively attracted away from

man by the presence of stabled animals. It enters houses in

largest numbers in midsummer and fall. Most of the females

must take occasional blood meals during winter hibernation,

but egg laying is suspended during this time and complete

hibernation has sometimes been observed. In Portugal males

swarm just after sunset inside or outside buildings, only a few^

females entering the swarm. Males and females will mate

in small cages.

In the U. S. S. E. and other northern areas atroparvus

breeds in brackish water of slight to moderate salinity and

tends to be a coastal form. In Caucasia it is found infrequently in fresh water, but in Spain and Italy, where it lives

well inland, it is found solely in fresh-water habitats. In

Portugal larvae usually live in ricefields.

This subspecies cannot live in localities where extreme cold

prevents winter feeding; extremely hot, dry summers also

present unsuitable conditions.

Relation to Disease.—A. I. atroparvus is responsible for

the so-called winter house malaria of the Netherlands and

East Friesland on the adjacent north coast of Germany. It

is often a carrier of malaria in Spain, Portugal, and Rumania, but there is little or no malaria transmission indoors by

this subspecies in the U. S. S. R., because most houses are

heated. It transmits some malaria on the Black Sea coast between the Bug and Danube River Deltas. This mosquito is

not everywhere associated with malaria, although it is found

throughout large areas of Europe. Its effectiveness as a vector is believed to be intermediate between labranchiae and

messeae.

References: 53, 67, 77, I47, 346, (For additional information, see

the descriptions of malaria under North Europe and North Asia ;

Spain and Portugal; and Italy, the Balkans, and Adjacent Islands.)

Recorded Distribution.—

Balearic Islands Israel

Belgium Italy

Caucasia Netherlands

Czechoslovakia Poland

Denmark Portugal

France Rumania

Germany Sardinia

Great Britain Spain

Hungary Sweden

Anopheles labranchiae labranchiae Falleroni

Bionomics.—Man is the normal host for labranchiae in

most areas. Females of this subspecies are usually attracted

to stabled animals to a small extent, differing in this habit

from most other members of the macuUpennis group. In

most localities they persistently enter houses, even when large

numbers of domestic animals are nearby, but occasionally

they fly back and forth between stables and houses, biting

both man and animals. However, in Sardinia the preferred

shelters are pigsties and stables. In Elba a study showed

that 96 percent of the observed adults rested in animal shelters rather than in houses, especially during the winter. Considerable numbers of females have been found in grottoes

and under bridges, living away from man, especially in areas

where they are forced to feed on wild mammal and bird hosts.

Overwintering females prefer inhabited and uninhabited

rural houses rather than village centers.

In Sicily the subspecies overwinters in the adult stage but

does not hibernate in the strict sense, up to half the population taking blood meals throughout the cold months. Temperatures probably regulate the numbers of adults that feed

during the winter. A small number of females develop fat

bodies during this time. A radius of about 1.9 miles is the

effective flight range of this subspecies.

In Italy the subspecies breeds principally in brackisli

water, with at least 10 parts per thousand of sodium chloride.

In Corsica, Sardinia, and Sicily, and along the Barbary coast

of North Africa from Tripoli to Morocco, it breeds in both

marshy and fresh water. In Sicily larvae live in many kinds

of water accumulations, ranging from coastal marshes to

gravelly beds of upland streams, where lahranchiae has no

fresh-water competitors. In Sardinia intermittent, slowmoving, or stagnant water courses are the principal breeding

places. About 99 percent of larval collections in that island
124 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

have been made from clear, smiiiy, fresh water containing horizontal plants, more commonly in quiet than in moving water. In Elba larvae have been found in irrigation

water, wells, and brackish stagnant water near tlie coast.

The first two generations in the spring inhabit larger pools

formed by spring rains. These pools are shallow, warmed

by the sun, and contain vegetation consisting of short

meadow grass and water buttercups {Ranunculus aquatiliH).

This subspecies also frequents tiny rivulets exposed to the

sun, tanks, wells, permanent swamps, marshes, lakes, and

certain ricefields and their ditches.

A. I. lahranchiae is principally a maritime mosquito. In

Sardinia seasonal activity begins in February, rises to a peak

of adult density in June or July, then shows a rapid decline,

with a small peak again in the fall.

Relation to Disease.—A, Z. lahranchiae is almost always

associated with intense malaria. Because of its history of

natural infections and its close association with man, this

subspecies is the principal if not the sole vector of malaria in

Sardinia and an important vector in Spain, northern Italy,

and north Africa.

References: 5S, 61, 72, W, 237. (For additional information, see

the descriptions of malaria under Northwest Africa ; North Europe and

North Asia ; Spain and Portugal ; and Italy, the Balkans, and Adjacent Islands.)

Recorded Distribution.

Albania

Algeria

Balearic Islands

Corsica

Elba

Israel

Italy

Libya

Morocco

Sardinia

Sicily

Spain

Spanish Morocco

Tunisia

Yugoslavia

Anopheles letifer Gater

Anopheles letifer appears in early records under the name

of A. umbrosus (see p. 138).

Bionomics.—A, letifer enters houses and bites man freely

from dusk to dawn. Adults probably rest out of doors during the day. They have been found in small numbers on

tree trunks and vegetation in cultivated areas. It is unusual

for females to bite during the day, although they have been

observed to do so in shady sites. This species is decidedly

anthropophilic, as shown by a study of it and several of its

close relatives, in which more than 90 percent of the captured specimens contained human blood.

Larvae of letifer prefer the dark-brown water of peaty

soil, such as that found in settlements and in plantations of

coconut, pineapple, rubber, and coiïee. They also inhabit

coastal collections of water that contain variable amounts of

vegetation, but they are intolerant of sea water and are never

found in it when fresh-water pools are a few yards away.

In Borneo larvae live under secondary shrub and tend to

breed in the shade, preferring water with a pH of 5.8 or

lower.

A, letifer is characteristically a mosquito of the flat coastal

plain and is found only in small numbers in hilly country

near the coast. Neither adults nor larvae are found within

virgin swampy jungle, the edge of which forms a distinct

dividing line between the area occupied by this species and

that inhabited by umbrosus. In the State of Selangor in

Malaya letifer reaches a peak of abundance during the last

4 months of the year because of the increased rains at that

time.

Relation to Disease.—In certain parts of Malaya letifer

is economically the most important vector of malaria, since

its breeding habits bring it into close association with man's

agricultural activities. Moderate to severe endemic malaria

exists in one area in Malaya, where letifer is the only species

present in large enough numbers to transmit it and where

infection rates are very high. In some parts of Borneo letifer^ with umbrosus and Anopheles baezai^ is the chief vector.

Its exact vector status in many areas is in doubt, because it

is confused with umbrosus in the early records.

References : 21^3, 28J^. (For additional information, see the descriptions of malaria under Malaya and Sunda Islands. )

Recorded Distribution.—

Borneo

Malaya

Sumatra

Anopheles leucosphyrus balabacensis Baisas, Anopheles

leucosphyrus hackeri Edwards, and Anopheles leucosphyrus leucosphyrus Dönitz

Anopheles leucosphyrus is composed of 6 or 7 closely related forms, none of which are well known. In most early

literature these are not named, but the three subspecies discussed below can often be distinguished, because they are the

only ones able to transmit malaria.

Bionomics.—In northern Borneo the females of Anopheles

leucosphyrus leucosphyrus and A, I, balabacensis feed on man

during the night and have been captured inside bed nets in

the morning. A peak of biting activity has been reported

between 12 and 1 a. m. In Labuan, British North Borneo,

balabacensis starts to feed in the early evening and has a

peak of biting activity between 1 and 4 a. m. In Celebes

Anopheles leucosphyrus hackeri is typically a house mosquito ; only small numbers of adults have been taken in outdoor resting places.

Larvae of leucosphyrus and balabacensis are found in

shaded temporary locations, such as rainwater in tracks and

wheel ruts, often in open jungle. A, Z. hacheri breeding is

largely confined to water collections in fallen, split bamboo

and rotten logs. On the coast of Malaya larvae live in

water collections in the bases of the nipa palm {Nipa fruticans). In Celebes hackeri breeds in sunlit marshy fields,

stopped-up gutters, ditches, pools, and small w^ells.

Relation to Disease.—In the "Bukitkuda, Lajau-batu, and

Manikar" areas of Labuan, where endemic malaria is severe,

balabacensis is the only vector and has very high rates of

gland infection. Elsewhere in northern Borneo this subspecies is probably an important vector. Transmission tends

to ñuctuate throughout the year, with definite periodic increases. Records of transmission in Burma and Assam almost certainly refer to balabacensis, since no other form,

with the possible exception of leucosphyrus, is known to exist

there. In much of Malaya the leucosphyrus complex appears to be relatively unimportant, and balabacensis is scarce

or absent. A. L hackeri has been found naturally infected in

Celebes and is important there.

References: 97, 98, 1S9, 2^3, 282. (For additional information, see

the descriptions of malaria under Bast Pakistan and Assam, South

China, Burma, Thailand, Sunda Islands, and Philippine Islands.)

Recorded Distribution.

A. I. balabacensis:

Assam

Borneo ( North )

Burma

Labuan

Philippine Islands

A. I. hackeri:

Borneo

Malaya

Soemha

Sumatra

A. I. leucosphyrus:

Andaman Islands

Borneo

Celebes

Java

Malaya

Philippine Islands

Sumatra

A. leucosphyrus (var. not

known) :

Formosa

Indochina

Anopheles listeri de Meillon

Bionomics.—In Zululand and Natal adults of Anopheles

listeri heive never been recorded in large numbers indoors,
MOSQUITOES OF MEDICAL IMPORTANCE 125

probably because studies have been made only in areas where

there is no extensive breeding. However, females are found

in very large numbers in habitations in South-West Africa,

where they bite man.

Larvae of listeri are similar to those of Anopheles gamhiae

in that they prefer exposed or grass-shaded pools in riverbeds in close proximity to man. They are often found with

gambiae and Anopheles pretoriensis.

Relation to Disease.—Because listeri rests in houses and

bites man in large numbers, it is a potentially important

vector of malaria in South-West Africa. Conclusive research to determine its actual role has not been carried out,

nor have many dissections been made to demonstrate whether

it is naturally infected.

References: 119, 258, 259. (For additional information, see the

description of malaria under Southern Africa.)

Recorded Distribution.—

Cape Province

Mozambique

Natal

Southern Rhodesia

South-West Africa

Transvaal

Anopheles ludlowi torakala van Hell

For a long time the presence of an unidentified vector of

malaria in the interior of south Celebes had been suspected.

Recent studies have shown that it is actually a subspecies

of Anopheles ludlowi. The adult is very similar to that of

Anopheles ludlowi ludlowi^ the presence of which in Celebes

has not yet been determined wâth certainty.

Bionomics.—Adults of Anopheles ludlowi torakala are

found in the interior of Celebes more frequently in houses

than out of doors, although females feed readily on man out

of doors, definitely preferring human blood. Very little additio.nal information about adult biology is available.

Larvae live almost exclusively in beds of large rivers, occupying small shallow pools in sand and gravel banks when

the water level in the main stream is low. Consequently,

they are swept away during the rainy season. Larvae are

almost always found in fresh water exposed to open sunlight and have often been observed up to about 300 feet

above sea level.

Relation to Disease.—This mosquito has been found naturally infected several times in the midst of malaria outbreaks in south Celebes. It is the most prevalent mosquito

and the only one infected. Dissections in six localities at

different times showed that 2 to 3 percent had gut infections.

A. I. torakala is thought to be a more important vector in

south Celebes than any other species, including Anopheles

sundaicus.

Reference: 161. (For additional information, see the descriptions

of malaria under Sunda Islands. )

Recorded Distribution.—

Celebes

Formosa (?)

Malaya (?)

Anopheles maculatus Theobald

Bionomics.—Females of Anopheles maculatus feed readily

on man but are also attracted to cattle. In some areas they

may ignore man almost entirely. In Malaya the species

appears to prefer livestock, but under certain conditions it

will bite man even when cattle are in the immediate vicinity.

Females are active at night and in some areas will readily

enter houses. They have been trapped in bed nets during the

night, especially in the Philippines, and usually bite between

9 p. m. and 2 a. m. Adults do not rest in houses after feeding but seek shelter out of doors. In Java they rest during

the day in the vegetation along the banks of streams. In

other areas adults are seldom found in buildings. In Formosa they have never been taken in houses, although larvae

have been found nearby. Swarms of maculatus males have

been observed in Malaya 15 to 20 feet above the ground in

open cleai'ings. Flight ranges have been reported to be well

over one-half mile. Females migrate great distances from

summer to winter habitats.

Larvae have been collected in China, Formosa, and the

Sunda Islands from riverbed pools with sandy or stony

bottoms and sometimes in rapidly flowing mountain streams

with whirlpools. In Malaya the favorite breeding places

are drains, pools, seepages, springs, ricefields, marshes, borrow pits, lake margins, and reservoirs. The species will also

breed in hoofmarks and artificial receptacles. Fresh water

is generally preferred. However, the larvae are sometimes

found in stagnant or polluted water but never in salt or

brackish water. Although aquatic vegetation is not necessarily associated with breeding, larvae often live in algae at

the edges of shaded forest streams. They prefer sunlit or

lightly shaded breeding sites and only rarely are found in

dense jungles. In the Philippines, Malaya, and the Lesser

Sundas recently cleared jungle may be an important breeding place.

A, maculatus is usually associated with hilly or mountainous country. It has been found at altitudes up to 5,000 feet.

Relation to Disease.—This species is an important vector

of malaria in some areas. In Malaya, Sumatra, Java, the

Eiouw Archipelago, and Bangka it has been reported with

extremely high infection rates and is one of the most important vectors of malaria. In the Philippines it is suspected strongly as a vector of that disease. In Indochina,

South China, and India some authorities consider it to be

a minor vector, whereas others think it to be very important.

The Formosa studies failed to reveal infected specimens in

the field, even though the species was encountered in large

numbers during the dry season in the southern and central

parts. In Celebes it is relatively rare and has never been

found infected.

References: U, 88, 120-122, S08, 340, 341, 351. (For additional

information, see the descriptions of malaria under South China,

Formosa, Indochina, Thailand, Malaya, Sunda Islands, and Philippine Islands.)

Recorded Distribution.—

Afghanistan

Assam

Bangka

Borneo

Burma

Celebes

Ceylon

China

Flores

Formosa

Hainan

India

Indochina

Java

Malaya

Pakistan (East and West)

Philippine Islands

Riouw Archipelago

Sumatra

Thailand

Timor

Anopheles maculipennis Meigen

In early works of European authors Anopheles maculipennis was used in a very broad sense to include the forms we

recognize today as Anopheles messeae^ A, melanoon melanoon,, A. melanoon subalpinus^ A. lahranchiae labranchiae,, A, lahranchiae atroparvus^ and A. sacharovi^ as well as

so-called A, maculipennis typicus^ the mosquito actually

described by Meigen from Germany. Subsequent studies

of the group, beginning about 1920, have brought to light

more and more differences in the various populations of the

maculipennis group. Refinements in methods of investigative biology eventually demonstrated the presence over

Europe and Asia of rather distinct biological, and to some

extent morphological, segregates. As these were defined,

they were named. As relationships among the various forms

were finally clarified by crossbreeding techniques, the names

were eventually combined to reflect a more exact relation-
126 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

ship as we understand it today. Details of the status of

each of the medically important species and subspecies of

the macuUpennis group are given separately in this handbook. As a result of the segregation described above, there

remains today a form, discussed below, which is variously

called macuUpennis^ macuUpennis typicus^ macuUpennis

macuUpennis^ or macuUpennis s. s. (strict sense), as opposed

to the more general term 7naculipennis group.

Bionomics.—Females of macuUpennis are primarily zoophilic. The presence of cattle tends to divert them from

man as a source of food. In Portugal, for instance, where

this species is not a vector, it is abundant in animal shelters

but rare in houses, and in Albanian studies, where suhalpinus

and macuUpennis were found in almost equal numbers, it

was represented in every collection of adults taken from

stables. In the Mediterranean region macuUpennis is said

to spend the winter in complete hibernation, usually initiating fat-body formation in September. However, in the

northern part of the U. S. S. E. and in Transcaucasia females hibernate in cold shelters or out of doors, where they

take blood meals throughout the winter.

Larvae are found in a wide variety of breeding places consisting of small pools, ditches, slow-moving streams, and

ricefields. They most often prefer small accumulations of

water but also occupy larger marshes in Albania and other

parts of the Balkans. In Albania this species breeds in immense numbers along the shores of "Lake Maliq," a large,

shallow, marshy body of water, and in lesser numbers in a

wide variety of habitats, including pools and ditches. Larvae were once found in water with 20 parts per thousand of

sodium chloride. In the U. S. S. E. the chief breeding

places are pebbly riverbeds, in which a sparse algal growth

may be present. The species is found characteristically in

upland waters that are cold, fresh, and pure, usually with

some vegetation and with sunlight during at least part of

the day.

In Tirana, Albania, macuUpennis builds to two population

peaks, one about July 1 and the other in August. The

abundance of both larvae and adults decreases between these

two peaks, probably because of high summer temperatures.

Relation to Disease.—A, macuUpennis has been found in

large numbers with messeae^ transmitting endemic malaria

in the villages at "Lake Maliq" on the Albania-Yugoslav

frontier. In Caucasia, Transcaucasia, and northern Iraq it

is believed to be the chief or sole vector with high natural

infection rates. However, in most of the Mediterranean region this mosquito is in contact with man so rarely that, even*

though it may transmit malaria occasionally, it is seldom

able to support even a mildly endemic form of the disease.

Exceptions are Macedonia and a center of severe malaria on

the Turkish coast. The species may also support mildly

endemic malaria in northern Iran.

References: 5S, 54, 67, 76, 147, S49. (For additional information,

see the descriptions of malaria under North Europe and North Asia ;

Italy, the Balkans, and Adjacent Islands ; the Near East ; Iran ; and

Afghanistan and West Pakistan. )

Recorded Distribution.—

Albania Morocco

Algeria Netherlands

Austria Norway

Balearic Islands Poland

Bulgaria Portugal

Caucasia Rumania

Corsica Sicily

Crete Spain

Czechoslovakia Sweden

France Switzerland

Germany Syria

Greece Tunisia

Hungary Turkey

Iran U. S. S. R.

Iraq Yugoslavia

Italy

Anopheles mangyanus Banks

Anopheles iiiangyanus^ another member of the A. funestusminim/us subgroup, is variously referred to in early records as

A. mmimus^ A. funestus^ and A. aconitus var. ßvpinae.

Bionomics.—Little information is available about the

habits of mangyanus adults. They have been collected from

clearings in heavy woods. Inconclusive results have been

obtained in flight-range studies. In one experiment extremely small numbers of adults were captured as far as onehalf mile from their point of liberation.

Larvae are found in clear water along the edges of shallow,

flowing streams, with sandy or rocky beds in forested areas.

They prefer the protection of grasses and exposed roots,

especially those of bamboo. Vegetated irrigation ditches

harbor larvae in large numbers.

This species has not been found in mountains above 2,000

feet.

Relation to Disease.—A, mangyanus lives in malarious

areas in the Philippines and can be made highly infective

in the laboratory. This species is generally believed to be a

vector in the Philippines, since it invades newly cleared areas

where it is associated with intense malaria.

References: 120, 20Jf, 308. (For additional information, see the

description of malaria under Philippine Islands. )

Recorded Distribution.—

Philippine Islands

Anopheles melanoon subalpinas Hackett and Lewis

Anopheles melanoon melanoon is the geographical representative of A, melanoon in the southern half of Italy.

It is not important in the transmission of malaria. A. m.

suhalpinus was originally based on a distinctive ^gg type

found in Spain, northern Italy, and the Balkans, and it was

described as a variety of macuUpennis, Biological studies

have shown it to be a subspecies of melanoon. For additional

details, see page 125.

Bionomics.—Females of suhalpinus rarely bite man. They

are frequently found in stables near fresh-water marshes.

Larvae are usually associated with large bodies of water,

such as marshes and lake margins. In Albania this subspecies has the most strictly limited habitat of all members of

the macuUpennis complex. In that country no larvae live in

water that is heavily shaded, without vegetation, with small

surface area, strong surface movements, or with high nitrate

or saline content.

Relation to Disease.—This subspecies is not known to be

a malaria vector in most of its range. In Lenkoran' in the

U. S. S. R. and the delta of the Safid End River, it is the

principal vector of severe malaria, but in Europe it is never

involved in malaria transmission because it is very zoophilic.

References: 5S, 5^, 67, 1^7, 346. (For additional information, see

the descriptions of malaria under North Europe and North Asia ; and

Italy, the Balkans^ and Adjacent Islands.)

Recorded Distribution.-

Albania

Balearic Islands

Bulgaria

Caucasia

Crete

France

Greece

Hungary

Iran

Israel

Italy

Spain

Switzerland ( ?)

Turkey

Yugoslavia

Anopheles melas Theobald

Anopheles iiielas has commonly appeared in the early literature as a variety of A. gamhiae because of its very close

morphological similarity to that species.
MOSQUITOES OF MEDICAL IMPORTANCE 127

Bionomics.—Like gambiae^ melas frequents houses and is

distinctly anthropophilic. Once inside a house, it rarely

leaves to rest outside.

Breeding takes place almost exclusively in brackish or sea

water, often with very high salt concentration. The species

is littoral, occupying lagoons or tidal swamps, principally

those of the mangrove Avicennia, Swamps containing

Rhizophora mangrove rarely harbor larvae. The flat peaty

soil, which is usually poorly drained and in w^hich Avicennia

grows, provides ideal breeding conditions, and populations

of larvae always increase after rain or flooding by tides.

Larvae do not seem to be afl^ected by sudden changes in the

salinity of the water. Since melas tolerates higher concentrations of sea water than gambiae^ a test to distinguish the

two species has been devised. This consists of the introduction of unknown larvae into a concentration of sea water

greater than 37i/^ percent. A. gambiae will not survive,

whereas 7nelas will complete its development.

Relation to Disease.—Like gambiae^ melas females are extremely important vectors of malaria, and parasites develop

as efficiently in melas as in gambiae, A. melas is the only vector in many littoral localities on the African West Coast.

Recent studies in Marshall Territory, Liberia, indicate

that melas is the important vector of filariasis during the

dry season and at the beginning of the wet season.

References : HS, 25S, 285, 286. (For additional information, see the

descriptions of malaria and filariasis under West Africa, West-Central

Africa, and Southern Africa.)

Recorded Distribution.—

Belgian Congo

Dahomey

French Equatorial Africa

French Guinea

Gambia

Gold Coast

Ivory Coast

Liberia

Nigeria

Portuguese Guinea

Senegal

Sierra Leone

Anopheles messeae Falleroni

Long ago some females of the Anopheles maculipennis

group w^ere noticed laying two types of eggs, gray ones

{Anopheles labranchiae) and dark ones with various patterns {Anopheles messeae and others). The characteristics

of the messeae egg pattern are shown in the pictorial key for

Italy, the Balkans, and Adjacent Islands (p. 67). For additional details, see page 125.

Bionomics.—Females of messeae rarely bite man, although

they are definitely attracted to him in certain localities.

They spend the winter either out of doors or in cold shelters,

such as outhouses, cellars, and lofts. During this time they

are immobile and take no blood. Hibernation, therefore, is

complete, at least in northern Europe. The species does not

live where winters are mild enough to prevent this hibernation. A large number of females develop fat bodies in

preparation for the winter.

Larvae are usually associated with luxuriant, submerged

vegetation. They are found with maculipennis and

Anopheles melanoon subalpinus in the highland areas of the

Yugoslav frontier, where they breed along the shores of

large, shallow, marshy lakes. Elsewhere they tend to live

in big river valleys and large marshes. Restrictions on

larval habitats of messeae are similar to those described for

subalpinus in Albania (p. 126). In the British Isles the species lives in inland fresh-water ponds and streams, where its

optimum breeding temperature is about 77° F.

Relation to Disease.—A. messeae is the chief malaria

vector throughout most of northern U. S. S. R. and the only

one in the vast region containing the Zavolzh'ye and western

Siberia. However, it is believed that messeae is not closely

enough associated w^ith man to maintain malaria in most

areas of Europe, the Balkans, and Caucasia. It is associated

with malaria transmission in only a few places in Europe—

471779 O—59 9

parts of Hungary, eastern Rumania, and "Lake Maliq" in

Albania—where dense human populations are close to large

marshy areas. There are large areas in much of the Balkans

and Caucasia without malaria where messeae breeds prolifically.

References: 5S, 54, 67, 249. (For additional information, see the

descriptions of malaria under North Europe and North Asia ; and

Italy, the Balkans, and Adjacent Islands.)

Recorded Distribution.—

Albania Israel

Austria Italy

Belgium Netherlands

Bulgaria Norway

Corsica Poland

Crete Rumania

Czechoslovakia Sweden

Denmark Switzerland

France Turkey

Germany U. S. S. R.

Great Britain Yugoslavia

Hungary

Anopheles minimus flavirostris Ludlow

Anopheles minimus ßavirostris was originally described as

Myzomyia -ßavirostris from specimens collected on Luzon.

This is the subspecies of Anopheles minÍ7nus throughout the

Philippines. In that area it is an easily identified mosquito,

but early records of its importance as a vector may be mixed

with those of Anopheles mangyanus and A, filipinae. Its

status in other parts of the Orient is uncertain, and some

w^orkers consider the forms that resemble ßamrostris in

India, Malaya, and the Sunda Islands to be a different mosquito.

Bionomics.—Females of flavirostris appear in large

numbers inside houses at night. They feed and leave again

before morning and rest under overhanging streambanks

during the day. They use both man and domestic animals

as food sources, although they are not attracted to the latter

in large numbers. The normal flight range is said to be

over 500 yards, and it is over a mile when i\\^j are blown

by the wind. In the Philippines experiments have revealed

flight ranges of about three-fourths mile during the dry

season. No flights have been observed against the w^nd.

Larvae are found at the edges of clear, slow-running,

partly shaded streams and in the clear water of swamp edges,

irrigation channels, drains, and borrow pits. In the Philippines they are nearly always prevalent near bamboo, which

may be in the form of scraps or mats of fallen plants floating

in the water. They also are found along margins of foothill streams shaded by tall bamboo trees but rarely in ricefields or brackish water.

A. m. flavirostris is found throughout foothills and rolling

land, seldom above an altitude of 2,000 feet. In the Philippines an annual rise in abundance begins in November or

December when the rainy season is almost over, and the

peak of activity and populations last from January to March,

after which there is a decline during the dry season.

Relation to Disease.—This subspecies is considered to be

one of the most important vectors of malaria in the Philippine Islands. It is reportedly present in Java with high

natural infection rates, and in northern Celebes it is reported

to sustain endemic malaria. In southern Celebes it has rarely

been found infected.

References : 91, 116, 122, 161, 308. (For additional information, see

the descriptions of malaria under Sunda Islands and Philippine

Islands.)

Recorded Distribution.

Ban

Borneo

Celebes

Java

Philippine Islands

Sumatra (?)
128 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Anopheles minimus minimus Theobald

The literature pertaining to Anopheles minimus minimus

prior to 1916 contains many references to A, listonii and A,

christofhersi^ which are names that have since been shown to

refer to minimus.

Bionomics.—Females of minimus are found in large numbers in houses and cattle sheds, preferring these to outside

locations. They are not commonly found in jungle areas, although they may be encountered resting under secondary

growth in jungle clearings. They feed readily on man and

in South China and Indochina are reported to be the most

anthropophilic of all anophelines. From 93 to 97 percent of blooded females have been reported to contain human

blood in Indochina. In some areas adults will feed on livestock if that source of food is more easily visited. In South

China intermittent feeding on domestic animals has been reported from 8 p. m. to 4 a. m., the peak of biting activity occurring from 10 p. m. to 2 a. m. and the adults staying inside

the same shelters during the next day. Eggs are laid in

largest numbers between nightfall and midnight.

Larvae are found in clear, slowly running, partly shaded

streams and in the clear water at the edges of swamps, irrigation channels, drains, ricefields, and borrow pits. In

India this subspecies breeds in clean grassy streams and

seepages, especially in some shade, during the monsoon season. In localities such as South China where it overwinters

in the larval stage, immature stages collect in large masses

in certain parts of more permanent rivers and streams during the cold months.

A, m. minimus \^ abundant at low altitudes, is common at

2,000 to 3,000 feet, and has been reported at altitudes as high

as 5,000 feet. In China there are two peaks of adult density.

The first is from May to mid-June at the start of the rainy

season, after which larvae are flushed out of their breeding

places, and the second in October and November, caused by

renewed breeding during the light autumn rains.

Relation to Disease.—This subspecies is considered to be a

very important vector of malaria throughout its range. It

is the outstanding transmitter in the hilly areas of China

south of latitude 30° N. Extremely high natural infection

rates have been observed in Fukien, Yunnan, and Szechwan

Provinces and in Hong Kong. This subspecies is an efficient

and important vector throughout India, East Pakistan, Assam, Indochina, Thailand, Burma, Formosa, Malaya, and

Sumatra.

In Tonkin and Hong Kong infective stages of Wuchereria

bancrofti have been observed in minimus. In Tonkin minimus is a probable vector of filariasis.

References: 87, 91, 122. (For additional information, see the descriptions of malaria and filariasis under India and Ceylon, East

Pakistan and Assam, South China, Formosa, Burma, Indochina,

Thailand, Malaya, and Sunda Islands.)

Recorded Distribution.—

Assam Indochina

BaU Java

Burma Malaya

Celebes Pakistan (East)

Ceylon (?) Soemba

China (South) Soembawa

Flores Sumatra

Formosa Thailand

Hainan Timor

India

Anopheles moucheti moucheti Evans

Bionomics.—Anopheles moucheti moucheti is very

strongly endophilic and anthropophilic and far outnumbers

other house-frequenting mosquitoes in the Belgian Congo.

Larvae are commonly found in quietly flowing, moderately

shaded water, and in the Congo, where this subspecies is

typically a stream breeder, they are observed singly in vegetation along banks of streams and pools. Larvae also are

found among short vegetation in running water in the western arid parts of South-West Africa.

A statistical analysis of conditions along the Congo River

has shown that the rise and fall of the water is directly correlated with the numbers of adults caught in habitations. A

change in water level results in a change of adult populations about 1 month later.

Relation to Disease.—In the presence of Anopheles gamhiae and A, funestus^ this subspecies is a secondary vector of

malaria and is less efficient than gamhiae. It may be of primary importance in the absence of those two species. In

Yangambi and Stanleyville, Belgian Congo, and Jinja,

Uganda, low natural infection rates have been reported. In

those areas moucheti is thought to be of some importance in

spite of the presence of hirge numbers of infected gambiae^

which transmit most of the malaria.

References : 119, 258-260, ( For additional information, see the

descriptions of malaria under West Africa, West-Central Africa, and

East-Central Africa. )

Recorded Distribution.—

Belgian Congo

Cameroons

French Equatorial Africa

Nigeria

Spanish Guinea

Uganda

Anopheles moucheti nigeriensis Evans

Bionomics.—In a series of investigations at Lagos, southern Nigeria, Anopheles moucheti nigeriensis represented 0.5

percent of the total anophelines caught indoors.

Larvae in Nigeria are found in partly wooded swamps and

along the banks of the Ogun River. A. m. nigeriensis is associated with clear-water swamps and Pistia throughout

most of its range.

Relation to Disease.—In the villages near Lagos, Nigeria,

high natural infection rates indicate that this subspecies may

be an effective vector of malaria. Little is known about it?>

ability to transmit the disease.

References : 119,168, 258, 260. (For additional information, see the

description of malaria under West Africa.)

Recorded Distribution.—

French Guinea

Nigeria

Anopheles multicolor Cambouliu

Bionomics.—Anopheles multicolor has been observed in

some areas to enter habitations and bite and in others to be

definitely exophilic, even when large populations are present.

Adults are strong fliers, having been found in the eastern

desert of Egypt about 8 miles from their nearest breeding

places.

The choice of breeding sites is wide. Larvae are commonly

found in water of high salinity and pH; a concentration

of salt of almost 6 percent has been recorded. Once they

were found in water that was almost two times as concentrated as sea water, with the surface heavily encrusted with

salt. Their breeding places include pans, small pools with

and without water weeds, stagnant or flowing drains, unused

shallow wells, and fresh-water collections and cesspools,

usually in desert oases. The larvae have never been observed

in large numbers in ricefields in Egypt.

Relation to Disease.—The importance of this oasis species

as a vector has not been firmly established. Although it is

rather easily infected with malaria parasites in the laboratory, it has not been found naturally infected. Because of

this and the fact that females are highly exophilic, some

workers have doubted its importance. On the other hand,

this species is said to carry, along with Anopheles pharoensis^

most of the malaria in Egypt, and it may occasionally be
MOSQUITOES OF MEDICAL IMPORTANCE 129

more important than A. sergenti in north Africa. Its role

as a vector in Arabia is not known.

References: 119, 258. (For additional information, see the descriptions of malaria under Northwest Africa, Egypt, and Arabia.)

Recorded Distribution.

Algeria

Anglo-Egyptian Sudan

Arabia

Cyprus

Egypt

India

Iran

Iraq

Israel

Jordan

Lebanon

Libya

Morocco

Pakistan (West)

Spain

Spanish Morocco

Syria

Tunisia

Turkey

Anopheles nigerrimus Giles

A medically important mosquito in Celebes, originally

called Anopheles hyrcanus var. X, is now known to be

Anopheles nifferrimus. Other names applied in the past to

this species are Anopheles nero^ A. pursati, A. williamsoni^

A. ientleyi^ A. venhuisi^ and "pupal type D of Crawford."

For additional details, see page 135.

Bionomics.—The behavior characteristics of nigerrimus

and sinensis adults are essentially alike. Information on the

bionomics of the latter applies to nigerrimus as well.

IjSLYYSie of nigerrimus prefer deep ponds, swamps, and other

large bodies of water covered by plants. Fishponds containing Pistia stratiotes harbor these larvae in large numbers in

Malaya. In Java this species prefers deep swamps and

swampy ricefields, with a great deal of vegetation, like Azolla^

Pistia^ Jussiaea, Hydrilla^ Eichhornia^ and Spirodela, In

Celebes the larvae are found in undrained, swampy ricefields

and fishponds covered with vegetation, as well as in neglected

pools and ditches in settlements. Larvae prefer shade, since

their temperature requirements are lower than those of

sinensis.

Relation to Disease.—Considerable evidence shows that

nigerrimus has been a vector in several malaria outbreaks.

As '^hyrcanus variety X," it was regarded as the principal

vector in outbreaks in Benteng and farther south in Celebes

and in the Karangbinangun district of Java. The natural infection rates in those localities, as well as in Sumatra, were

rather low. A. nigerrimus may support a mild endemic malaria if mosquito populations are small. In the vicinity of

Kuala Lumpur, Malaya, nigerrimus and sinensis were the

principal vectors in an outbreak of malaria. This has probably occurred many times where the range of the two species

overlaps. A. nigerrimus may be an important vector in the

southern ricegrowing areas of Thailand. It is apparently

unimportant as a vector in India, where it has not been found

naturally infected.

Development of Wuchereria hancrofti to the infective

stage in nigerrimus has been observed, but little is known

of the part nigerrimus plays in the actual transmission of

filariasis.

References: 122, 161, 28S. (For additional information, see the

descriptions of malaria and filariasis under Thailand, Malaya, and

Sunda Islands. )

Recorded Distribution,—

Assam Indochina

Borneo Java

Burma Malaya

Celebes Moluccas

Ceylon Pakistan (East and West)

China (South) Philippine Islands

Hainan Sumatra

India Thailand

houses and bites man. However, throughout Africa, with

the exception of parts of Belgian Congo, Liberia, Sierra

Leone, and French West Africa, it is rare, and when it is

occasionally found in houses, it is usually not infected. In

south Africa adults have never been found indoors. Houseentry studies in localities where larvae are abundant show

endophilism to be accidental. In Anglo-Egyptian Sudan

nili and Anopheles loellcomei are the most abundant species

in rural areas.

Larvae inhabit vegetated banks of flowing rivers and

streams. Vegetation such as Pistia or shade from other

sources is important to this species. Vegetated islands are

the principal breeding grounds along the Congo River.

Since the larvae in South-West Africa live along the margins of swiftly flowing streams and rivers, they are likely to

be found along the two main river courses. In Kenya larvae

have been reported underneath steep, undercut streambanks

in the deep shade of high jungle vegetation.

Although the species is often rare and usually found in

small numbers, it is widespread over Africa. It is essentially

absent from arid regions because of the lack of shaded breeding places. Adult populations of nili^ like those of Anopheles

moucheti moucheti^ are directly correlated with the rise and

fall of the Congo River.

Relation to Disease.—In parts of Anglo-Egyptian Sudan,

Belgian Congo, Liberia, Sierra Leone, and French West

Africa, high natural malaria infection rates have been encountered. In these localities females may not be abundant.

Nevertheless, the species should be considered as a secondary

vector of malaria and less important than Anopheles gamhiae or A. funestus.

References: 119, 258-260. (For additional information, see the

descriptions of malaria under Anglo-Egyptian Sudan; Eritrea, the

SomaUlands, and Ethiopia ; West Africa ; West-Central Africa ; EastCentral Africa ; and Southern Africa. )

Recorded Distribution.--

Anglo-Egyptian Sudan

Angola

Bechuanaland

Belgian Congo

Cameroons

Dahomey

Ethiopia

French Equatorial Africa

French Guinea

French West Africa

Gold Coast

Ivory Coast

Kenya

Liberia

Mozambique

Natal

Nigeria

Northern Rhodesia

Nyasaland

Sierra Leone

Southern Rhodesia

South-West Africa

Spanish Guinea

Transvaal

Uganda

Anopheles nili (Theobald)

Bionomics.—In certain restricted localities near rivers

Anopheles nili is found in large numbers, and it enters

Anopheles oswaldoi (Peryassú)

Bionomics.—Females of oswaldoi are apparently not attracted to human habitations but will attack man when their

jungle habitats are invaded. Even though a few specimens

have been collected in houses in jungles in the Fort Sherman

reservation in Panama, osioaldoi is not regarded as a species

that frequents houses there or in Trinidad. In Surinam it

prefers animal shelters and seldom invades houses.

Larvae breed in the well-shaded fresh water of jungle

pools, swamps, and stagnant streams.

A. ostoaldoi is a jungle mosquito that normally is not

closely associated with man. It is most prevalent, at least

in Surinam, at the end of the rainy season.

Relation to Disease.—Although common in some parts of

Brazil, osioaldoi is not regarded as an important malaria

vector in that country because of its wild habitat. In Panama it transmits malaria to man when its jungle haunts are

invaded, and it is believed to do so in Colombia under similar

circumstances.

References : 61, 19, 222, 226, 292, 308. (For additional information,

see the descriptions of malaria under Panama, Northwest South

America, and Brazil and the Guianas. )
130 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Recorded Distribution.

Argentina French Guiana

Bolivia Guatemala

Brazil Panama

British Guiana Paraguay

Colombia Surinam

Costa Rica Trinidad

Ecuador Venezuela

Anopheles paludis Theobald

Bionomics.—Anopheles paludis inhabits the low tropical

rain forest and is most active from August or September to

April or May. It is not normally a domestic anopheline and

is generally assumed not to enter human dwellings, although

at Coquilhatville, Belgian Congo, it has been regularly captured in native huts. In Nigeria the species attacks man

freely out of doors. In Africa a few paludis females have

been caught in monkey-baited traps between 6 and 10 p. m.,

one specimen as high as 58 feet in a tree. Observations indicate that paludis is most active in darkness under conditions of low temperature and relatively high humidity. A,

paludis lives in both rain forests and plantations, most commonly in the former. During the daytime it is extremely

scarce in the forest canopy ; at night it bites in both forest

and plantation areas. A, paludis shows an ill-defined peak

of activity just before and after 1 a. m.

In Liberia larvae live in dense shade in swamps that contain aquatic vegetation. In southern Nigeria and Sierra

Leone the species is found in the clear water of swamps,

ponds, backwaters of streams, springs, ditches, and ricefields.

Aquatic and semiaquatic vegetation is almost always present

in those habitats. Larvae are rarely found in completely

exposed rain pools. In the high grasslands of the Transvaal

they appear to be resistant to frost.

Relation to Disease.—A. paludis is regarded as a malaria

vector of some importance in several areas of the Belgian

Congo. Females captured in native huts have been found

with stomach infection rates of 5.5 percent and salivary

gland infection rates of 0.39 percent.

References: 69, 119, U8, U9, 152, 252, 258, 260. (For additional

information, see the description of malaria under West-Central

Africa. )

Recorded Distribution.—

Angola Nigeria

Belgian Congo Senegal

Ethiopia Sierra Leone

French West Africa Tanganyika

Gold Coast Uganda

Liberia

Anopheles pattoni Christophers

Bionomics.—Little is known aboutthe habits of Anopheles

pattoni adults, but females are known to be zoophilic as

well as anthropophilic.

The species hibernates as second- or third-instar larvae,

presumably throughout the entire winter, sometimes under

thick ice. During the breeding season immature stages

live in slowly running hill streams, rock pools, and pools, in

streams with sandy bottoms and an abundance of algae.

Relation to Disease.—For many 3^ears this species has

been regarded as the principal malaria vector in northern

China, but until 193Y infected specimens had never been

found in nature. Females were shown to be good experimental vectors of malaria in 1929. A. pattoni is a vector

in Shantung, Hopei, Szechwan, Shansi, Shensi, and eastern

Kansu Provinces of China. It definitely lives as far north

as Ch'in-huang-tao, and its presence farther north in Manchuria is possible.

References: SI, 61, 88, 90, 174. (For additional information, see

the description of malaria under North Europe and North Asia.)

Recorded Distribution.—

China (north)

Manchuria

Anopheles pharoensis Theobald

Bionomics.—In Egypt the females of Anopheles pharoensis commonly enter houses and bite man viciously in the

open at sunset. Precipitin tests show also that pharoensis

is definitely anthropophilic in Egypt. This is true in other

areas as well, although it may be only weakly anthropophilic

in certain parts of Kenya, Nigeria, and Uganda, since here

it does not enter houses as freely as Anopheles gamhiae and

A, funestus. In the central Sudan immense numbers of

females have been known to bite both man and animals.

Studies show that adults are blown for comparatively long

distances from their breeding places in Egypt. This mosquito is often associated^ with extensive wet-season flooding

along the banks of rivers and in grassy areas.

Larvae are generally associated with large bodies of fresh

water and are almost invariably found in the presence of

aquatic vegetation of some sort despite the large variety of

breeding places. In Kenya, Uganda, and Mozambique

larvae are associated with tall grass, reeds, and papyrus;

with Pistia^ Potamogetón^ and floating vegetation at stream

edges, in lakeshore swamps, and in borrow pits; and with

roots of decaying aquatic plants in open lakes. In Egypt,

in addition to the above sites, larvae have been found in

ricefields, sakieh pits, abandoned wells, and old water tanks.

They have been reported from wells in Ethiopia. Larvae

have been observed in ricefields several times in Egypt and

Kenya, although in Mozambique and Madagascar this

habitat has not been noted to any great extent. In Lakes

Kivu and Albert in Belgian Congo large numbers of larvae

are found in Ceratophyllum demersunh. There is no extensive breeding in rivers in this area during the dry season.

The species is a swamp breeder in South-West Africa and

is probably restricted to the extreme northern part of that

country. Young larvae tend to remain at the bottom of

collecting containers in contrast to those of gambiae^ which

ordinarily stay at the surface.

This species may appear in very large numbers for a night

or two, and then disappear for long periods.

Relation to Disease.—Although there is some doubt about

the role pharoensis plays as a vector of malaria, much evidence points to its considerable importance. A, pharoensis

is said to be the most important vector, along with Anopheles

multicolor^ of malaria in Egypt. In almost all experiments

it has been successfully infected and is often found infected

in nature in Egypt, French West Africa, southern Nigeria,

Kenya, and Uganda. In central Sudan, where it is found

in large numters, it is not looked upon as an important

species, but in other parts of French West Africa it has

been noted as a secondary vector. It is regarded as a minor

vector in Madagascar. Although dissections in these and

certain other areas have proved entirely negative for malaria

parasites and positive dissection rates have always been low,

pharoensis has been associated with outbreaks of malaria

where it was the only known vector present. In such locations, of course, it is of great importance, but where it is

present in large numbers with gamhiae and funestus^ its importance may be considerably less. It is said to be of little

importance in South-West Africa.

References: 119, 258-260. (For additional information, see the descriptions of malaria under Egypt; Eritrea, the Somalilands, and

Ethiopia ; West Africa ; East-Central Africa ; Madagascar and Adjacent Islands; Southern Africa; and Arabia.)
MOSQUITOES OF MEDICAL IMPORTANCE 131

Recorded Distribution—

Anglo-Egyptian Sudan

Angola

Arabia

Bechuanaland

Belgian Congo

Cameroons

Dahomey

Egypt

Ethiopia

French Equatorial Africa

French Guinea

French West Africa

Gambia

Gold Coast

Israel

Ivory Coast

Kenya

Madagascar

Mozambique

Natal

Nigeria

Nyasaland

Senegal

Sierra Leone

Somalia

Southern Rhodesia

South-West Africa

Syria

Tanganyika

Transvaal

Uganda

Anopheles philippinensis Ludlow

Many early records of Anopheles annularis may apply to

A, philippinensis^ since the morphological resemblance between the two species is striking.

Bionomics.—Little is known about the habits of philippinensis adults. In Burma females have been collected in

houses as well as in cattle sheds and stables, but in Yunnan

they have never been found in human dwellings. Females

are reported to be definitely anthropophilic in Bengal but

predominantly zoophilic in Indochina. In Malaya swarms

of males, usually 4 to 8 feet above the ground and not

related to any ground object, have been observed from

shortly after sunset to complete darkness. In the Philippines adults are attracted by the hundreds to white light.

Larvae live in large bodies of water, such as tanks, sloughs,

quiescent lake margins, impounded water, large ponds, and

open rush swamps, usually in rather dense vegetation.

Relation to Disease.—Considerable variation in natural

infection rates for malaria is given in the literature. In the

western part of the Ganges Delta area of East Pakistan,

where the species is an important vector, infection rates are

very high. Elsewhere infection rates are rather low or

negative, and the species is believed to have no importance

in malaria transmission.

Complete development of Wuchereria hancrofti has been

observed in philippvnensis. However, only a few specimens

have been found naturally infected. The role of this mosquito as a vector of filariasis in nature is not exactly known.

References : 8^, Öi, 12%, 308, 341. (For additional information, see

the descriptions of malaria and filariasis under East Pakistan and

Assam.)

Recorded Distribution.—

Andaman Islands Java

Assam Malaya

Borneo New Guinea

Burma Pakistan (Bast)

China (South) Philippine Islands

Hainan Sumatra

India Thailand

Anopheles plúmbeas Stephens

Although originally described from Ireland, Anopheles

plumbeus has been recorded from a large number of localities

in Europe and western Asia. It was known in Europe for

some time as Anopheles nigripes. Anopheles harianensis is

found in India and is very closely related to plumbeus^ but

it is apparently of no importance in the transmission of

malaria.

Bionomics.—A, plwnrvbeus is a persistent biter, but it is

universally known as a wild mosquito, preferring natural

resting places to indoor haunts in most of its range. In

Albania females have been commonly collected from stables.

Larvae live in water-filled tree holes and in similar habitats, such as hollows in the tops of stumps and pieces of wood

with depressions filled with rainwater. In dry seasons they

are sometimes found in water butts, tanks, cisterns, rain

barrels, and even in peat beds. In Turkey they have been

reported from small caves containing water and in small

heavily shaded ground pools. The species of tree may have

special significance, but the results of various investigations

have differed widely in this respect. In Czechoslovakia

larvae commonly live in water of pH 8.8 to 9.1 in hollows of

''''Acer pseudoplatanoides"^^ and Fagus syïvatica. The numbers of fourth-instar larvae always appear to be small, possibly because development in the trees is limited to a certain

number of larvae on account of the space available. In

Albania plwmbeus is the easiest of all Anopheles species to

rear in the laboratory. Larvae were kept in rainwater at

about 80° F., to which a small amount of blood serum was

added ; adults resulting from these rearings appeared to be

normal in all respects.

A, plumbeus is closely associated with deciduous forests.

It is believed to overwinter in the larval stage. Over most of

its range it is not an abundant mosquito. In Albania and

Turkey two population peaks occur, one in April and one in

August and September. In England adults are most numerous from April to October and larvae have been collected

during every month of the year.

Relation to Disease.—Experimentally plumheus appears

to be a very efficient vector and plays a small part in endemic

malaria on the shores of the Black Sea. Because of the

nature of its breeding places and adult habits, this species

is not regarded as a vector elsewhere.

References : 52, 54, 216, 217, 249, 346. (For additional information,

see the description of malaria under North Europe and North Asia.)

Recorded Distribution.—

Albania Iraq

Algeria Ireland

Arabia Italy

Austria Mongolia

Caucasia Netherlands

Corsica Portugal

Czechoslovakia Rumania

Denmark Sardinia

France Sicily

Germany Spain

Great Britain Sweden

Greece Switzerland

Hungary Turkey

Iran Yugoslavia

Anopheles pretoriensis Theobald

Bionomics.—Anopheles pretoriensis seldom frequents

houses in south Africa, Nyasaland, Kenya, Ethiopia, or

Tanganyika. Large numbers of adults rest in outdoor

haunts, such as grassy banks of streams, natural cavities in

the earth, and moist clefts or rocks near breeding places. In

Southern Rhodesia, where the greatest numbers of females

have been observed indoors, only about 20 percent of the

total catches of mosquitoes were from indoor locations. Precipitin tests in Kenya indicate that this mosquito has a slight

preference for human blood.

Rarely seeking deep shade for oviposition, pretoriensis

tends to prefer sites that are lightly shaded or exposed to the

sun. In south Africa larvae have been found in completely

exposed water, such as rain puddles, pools left by drying

streams, seepages, and ponds. In South-West Africa they

are very commonly associated with Anopheles gambiae in

exposed pools and seepages. In Southern Rhodesia larvae

live in a wide variety of locations, except heavy shade, deep

wells, or artificial containers. In Kenya they inhabit

streams, rivers, swamps, and irrigation canals, permanent or

otherwise. About one-half of the total larvae have been collected from clear water. In Yemen they were taken at altitudes of 1,000 to 3,000 feet in the quiet marginal water of

drying streams containing filamentous green algae and short

vegetation.
132 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Evidence indicates that this mosquito is anthropophilic to

some extent and exophilic over the greater part of its range.

In Nyasaland it is a dry-season anopheline.

Relation to Disease.—The importance of this species as

a malaria vector is probably slight in the presence of gamhiae

and Anopheles funestus^ since it rarely enters houses. In

Ethiopia the species may be a vector of malaria, but its exact

relation to this disease is not known with certainty. The low

natural infection rates for females found out of doors in

Transvaal and Southern Ehodesia indicate that this species

may be of some importance in those countries.

References : 119, 206, 258-260. (For additional information, see the

descriptions of malaria under Eritrea, the Somalilands, and Ethiopia ;

West Africa ; West-Central Africa ; East-Central Africa ; Madagascar

and Adjacent Islands; and Southern Africa.)

Recorded Distribution.

Anglo-Egyptian Sudan

Angola

Arabia

Bechuanaland

Belgian Congo

British Somaliland

Cameroons

Eritrea

Ethiopia

French West Africa

Gold Coast

Ivory Coast

Kenya

Liberia

Madagascar

Mozambique

Natal

Nigeria

Nyasaland

Somalia

Southern Rhodesia

South-West Africa

Tanganyika

Transvaal

Uganda

Anopheles pseudopunctipennis pseudopunctipennis

Theobald

Anopheles pseudopunctipennis pseudopunctipennis may be

composed of several biological forms, since at least two egg

types haVe been found. The other subspecies, Anopheles

pseudopunctipennis franciscanus^ is present in California

but is not a vector of malaria.

Bionomiés.^—In the highlands of Lower California, Mexico, Guatemala, and Argentina females of pseudopunctipennis fly considerable distances, enter houses, and feed avidly on

man. In Argentina they feed in large numbers inside houses.

On the other hahd, several localities in Central America and

the West Indieâ have reported pseudopunctipennis to be

exophilic. Females are apparently not attracted to houses in

parts of Costa Rica, the Canal Zone in Panama, or Grenada.

Almost all larval habitats are exposed to sunlight and contain mats of green algae. The habitats are usually pools and

eddies in shallow or drying streams, seepages, ground pools,

and tanks. In Panama larvae are found in clear water rich

in Spirogyra^ but as far south as Argentina larvae can be

found throughout the year in the presence or absence of algae.

They live in algal mats and cling to flotage in shallow, quiet

or running water. In some areas with a long dry season this

species maintains itself in springs and other fresh-water collections in the mountains and returns to its breeding places

at lower altitudes when rain pools form in riverbeds.

x4. p, pseudopunctipennis is essentially a mountain and dryseason species. Rainfall is the chief climatic factor controlling its abundance. In Mexiiîo heavy rains flood larvae

from the mountains into extensive areas at lower altitudes.

The species is mpst common in Grenada through July and

August. In Peru its peak of abundance is between January

and June.

Relation to Disease.—^, p, pseudopunctipennis is regarded as an important malaria vector. It has been found

with high natural infection rates iñ several localities in the

northwestern Provinces of Argentina, where this mosquito

has been known for years as a dangerous transmitter. In

Mexico infection rates up to 67.2 percent have been reported.

The species is responsible for large amounts of endemic malaria in the central valley at 6,000 to 7,000 feet. It is almost

solely responsible for the malaria in Tacna in southern Peru

and Anca in northern Chile. It is the only Anopheles species

and vector on the entire western slope of the Andes in Peru

and the chief vector on the eastern slope. It is responsible

for the malaria present at high altitudes in Ecuador and

southwest Colombia. At Lake Atitlan in Guatemala it causes

malaria outbreaks in villages along the shore. In the United

States and Panama it is not an important vector.

References: 61, 226, 292, 308, 334. (For additional information, see

the descriptions of malaria under America North of Mexico, Mexico,

Central America, Panama, Northwest South America, and Lower South

America. )

Recorded Distribution.—

Argentina Guatemala

Bolivia Honduras

British Honduras Mexico

Chile Nicaragua

Colombia Panama

Costa Rica Peru

Ecuador United States

El Salvador Venezuela

Anopheles pulcherrimus Theobald

Bionomics.—Females of Anopheles pulcherrimus rest,

often in exceedingly large numbers, in houses, tents, barracks,

and cattle sheds. They prefer human settlements, but occasionally they are found far from man and his domestic animals. Females feed on man and animals and may attack in

the evening, at night, or in the daytime, outdoors or indoors.

The adults are strong fliers. Since they are able to withstand dryness, they may be carried long distances by the

wind.

Larvae are found in the open, sunlit, weedy waters of

swamps, seepages, marshes, ponds, and ricefields. They

commonly inhabit polluted and often stagnant water but

never in any of the breeding places listed above unless the

water temperature is very high.

A, pulcherrimus is the typical mosquito in the oases of

mid-Asian deserts. It breeds in larger bodies of water and

hibernates as a third- or fourth-instar larva. The mosquito

rarely is present above an elevation of 1,000 feet and is most

numerous along the low, broad river valleys. In the hottest

parts of its range the populations begin to increase in May.

Farther north there is little increase in numbers until later

in the year.

Relation to Disease.—The species has been easily infected

experimentally with malaria parasites, and it has been found

naturally infected in central Asia and Sind in West Pakistan,

although in the latter country it is probably not an effective

malaria vector. It is regarded as an important transmitter

in the Murgab Valley of central Asia, where it maintains

severe endemic malaria in the absence of other Anopheles

species.

References: 67, 91, 120, 346. (For additional information, see the

descriptions of malaria under North Europe and North Asia, and

Afghanistan and West Pakistan.)

Recorded Distribution.-

Afghanistan

Arabia

Caucasia

India

Iran

Iraq

Israel

Lebanon

Pakistan (West)

Syria

Turkestan

Anopheles punctimacula Dyar and Knab

Bionomics.^Females of Anopheles punctimacula are anthropophilic, butthey will feed on livestock. They often visit

human habitations in large numbers. In Panama the species

is abundant in jungle areas, where it makes long flights,

enters houses, and feeds on humans. Along the Chagres

River of Panama many more females have been captured in

houses than around domestic animals.
MOSQUITOES OF MEDICAL IMPORTANCE 133

Larvae most commonly live in small streams, ponds,

swamps, and other habitats containing clear, cool, densely

shaded water.

Relation to Disease.—^, punctimacula in Panama is easily

infected experimentally with malaria parasites, is found

naturally infected, and enters houses readily. For these reasons it is believed to be an important vector in that country

and also in the Cauca Valley of Colombia, where it is associated with intense malaria. Its role as a vector has not

been determined elsewhere with certainty. In certain areas

of Brazil it is not regarded as a vector because of its scarcity.

References: 67, 79, %26, 292, 308. (For additional information, see

the descriptions of malaria under Central America, Panama, and

Northwest South America.)

Recorded Distribution.—

Argentina El Salvador

Bolivia Guatemala

Brazil Honduras

British Guiana Mexico

British Honduras Nicaragua

Colombia Panama

Costa Rica Peru

Ecuador Venezuela

Anopheles punctulatus Dönitz

The Anopheles pimctulatus group consists of four closely

related species in Australasia. They are known as Anopheles

farauti, previously treated in older literature as A, punctulatus var. moluccensis^ A, koliensis^ A, clowi^ and A, punctulatus. The species commonly referred to as punctulatus in

areas west of the Moluccas is Anopheles tessellatus.

Bionomics.—In some areas females of punctulatus enter

houses freely and feed readily on man, but in others they do

not attack man often, either indoors or in the open. They

do not usually remain indoors after obtaining a blood meal.

Although the species is occasionally said to be exclusively

anthropophilic, little is actually known about its blood preferences. All precipitin tests highly positive for human blood

have been performed on mosquitoes collected near human

habitations. Females are nocturnal ; very few are active before 9 p. m. They are dangerous because of their noiseless

flight, painless bite, and their presence in locations when few

other Anopheles species are present. Outdoor resting places

are probably in jungle areas near breeding places and near

native villages.

Larvae are found in all types of water exposed to sunlight

or partial shade. These include pools, ditches, hoofprints,

water barrels, bilge water of boats, wheel ruts, trenches, bomb

craters, and muddy pools, which occasionally contain algae.

Althoug*h observed coastally as well as inland, immature

stages are rarely found in brackish water and rarely, if ever,

in flowing water. Larvae are often found in accumulations

of water m stands of the sago palm. In Guadalcanal larvae

are restricted to valleys of large streams and rivers and usually live some distance from the coast in temporary pools

exposed^to sunlight. They are rarely found at stream margins or in drying stream beds, and then only during the dry

season. Eggs may be laid in mud, and even after complete

drying will hatch upon subsequent wetting. The range of

the species is frequently extended to the coast along the river

courses during the rainy season, when larvae frequent temporary pools in large numbers. These pools may be very

muddy, with or without marginal vegetation. Larvae are

extremely active, especially when crowded in locations without vegetation, and spend much time submerged.

In New Guinea this species is seasonal, reaching a peak of

abundance about June. During dry weather it almost disappears. In Guadalcanal the life cycle is completed in from

6 to 8 days. Development is very synchronous ; i. e., all the

individuals of any given generation attain adulthood at

about the same time.

Relation to Disease.—The importance of this species is

nearly equal to that of fararnti in the transmission of malaria

in Australasia. It has been found w^ith high natural infection rates in New Guinea, Ceram, and Guadalcanal, although

in certain localities it is present in large numbers where there

is not a proportional amount of malaria.

The complete development of Wuchereria hancrofti in

punctulatus has been experimentally demonstrated in New

Britain. Up to 15 percent of this mosquito species has been

found naturally infected in Guadalcanal. It is not known

what role the species plays in the transmission of filariasis

in nature.

References : 57, 121,122, 228, 294. (For additional information, see

the descriptions of malaria and filariasis under Australasia.)

Recorded Distribution.—

Australia (Northern Territory, New Guinea

Queensland) New Hebrides

Ceram New Ireland

Moluccas Solomon Islands

New Britain

Anopheles quadrimaculatus Say

Anopheles quadrimaculatus is distantly related to the European members of the A, maculipennis complex. It is referred to in the very early American literature as Anopheles

maculipennis.

Bionomics.—A, quadrimaculatus is the most abundant

North American anopheline found in houses and other manmade shelters and, with Anopheles freehorni^ is the most

definitely anthropophilic, even though livestock is preferred

over humans as a source of blood. This species passes the

winter in the adult stage, even in the southern part of its

range, where larvae are found the year round. Fertilized

females gather in immense numbers in basements, root cellars,

and hollow trees. The effective night range is about 1 mile.

This distance is exceeded from time to time because of random flight patterns.

This species usually breeds in ponds, swamps, bayous, and

borrow pits where sunshine and floating or emergent vegetation are abundant. The saying, "Quads are where you

find them," has a basis in fact. Larvae may also be found

in heavily polluted water, tin cans, barrels, borrow pits without vegetation, cornfield furrows, and other unusual habitats.

They prefer to stay in the parts of these breeding waters that

are made slightly alkaline by the action of sunlight and

vegetation. In the Mississippi Valley this species prefers

densely wooded swamps, which contain more highly alkaline

water than those on the Atlantic coastal plain, where the

water is much more acid. Larvae apparently cannot tolerate

deep shade and acid water together.

Relation to Disease.—This species is the most important

vector of malaria in eastern North America. Although

malaria has almost entirely disappeared from the United

States, quadrimaculatus is ever present and a constant source

of danger. Its importance in Mexico has not been thoroughly

studied.

Reference: 61. (For additional information, see the descriptions

of malaria under America North of Mexico and Mexico.)

Recorded Distribution.—

Bahama Islands

M^exico

United States

Anopheles rhodesiensis Theobald

Anopheles rupicolus has been confused with A. rhodesiensis in the past because of its extremely close morphological relationship. The former species is not a medically important species so far as is known. Kirkpatrick's {205)
134 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Egyptian records of rhodesiensis actually refer to Anopheles

dHhali.

Bionomics.—A, rhodesiensis is not regarded as domestic

in Sierra Leone, south Africa, Southern Rhodesia, Mozambique, or Kenya. It has occasionally been found inside

houses in Tanganyika and Sierra Leone, although in many

parts of Sierra Leone it does not appear in or near human

dwellings at all. Females do not feed readily on humans

under laboratory conditions. In Southern Rhodesia only

three females, one of which contained parasites, ware ever

captured in habitations during a 4-year study. Little is

known about the outdoor haunts of this species, even in south

Africa, where it is common and does not frequent houses.

Larvae live in a large variety of exposed or shaded breeding places, such as rock pools, stream beds, seepages with

little shade, margins of streams with running water, springs,

pools, ditches, hoofprints, and large artificial containers. In

Tanganyika the species breeds in nearly all available water

collections, but in arid places, such as in South-West Africa,

it is restricted to streambanks and fountains, where larvae

are common in short vegetation.

Relation to Disease.—This species is a potential vector

of malaria, although its lack of endophilism and natural infections tend to make it of little importance. In Koinadugu,

Sierra Leone, only once was it found with a high rate of natural infection. The species is believed to be a vector of secondary importance in Ethiopia. Investigators have completely failed to infect rhodesiensis in the laboratory with

Pldsmodium malariae and only rarely have they infected it

with P. falciparum.

References: 205, 206, 258, 259. (For additional information, see

the descriptions of malaria under Eritrea, the Somalilands, and

Ethiopia; West Africa; West-Central Africa; East-Central Africa;

and Southern Africa.)

Recorded Distribution.

Bechuanaland

Belgian Congo

British Somaliland

Cameroons

Cape Province

Egypt

Eritrea

Ethiopia

French West Africa

Gold Coast

Iran

Ivory Coast

Kenya

Mozambique

Northern Rhodesia

Nyasaland

Sierra Leone

Somalia

Southern Rhodesia

South-West Africa

Tanganyika

Transvaal

Uganda

Anopheles rufipes ingrami Edwards

Anopheles rufipes ingrami was described in 1929 as a variety of A. rufipes on the basis of a difference in markings

on the third hind tarsal segment of the female (see p. 58).

Bionomics.—Near Freetown, Sierra Leone, ingrami is extremely rare in human habitations, even though the mosquito

may be breeding extensively in the immediate vicinity. On

the other hand, it has been found in considerable numbers in

human habitations in northern Nigeria.

Large numbers of larvae have been observed in big, shallow pools of fresh water at Kissy near Freetown during

April and May.

Relation to Disease.—The importance of ingrami to the

distribution of malaria has not been determined accurately.

This subspecies is included as a possible vector, because in

northern Nigeria (1) it has been found in houses in considerable numbers, (2) experimental infections have been obtained, and (3) extensive breeding has been recorded.

References : 119, 258. ( For additional information, see the descriptions of malaria under West Africa and Southern Africa.)

Recorded Distribution.—

Anglo-Egyptian Sudan

French West Africa

Gold Coast

Mozambique

Nigeria

Northern Rhodesia

Sierra Leone

Anopheles rufipes rufipes Gough

Bionomics.—Large numbers of females of Anopheles

rufipes rufipes have been captured in human habitations in

Southern Rhodesia and central Sudan, and sometimes they

outnumber A, gambia£ in houses in northern Haute-Volta.

In contrast, adults are seldom found indoors in south Africa

and Kenya. They are commonly observed in streambank

crevices and similar locations close to breeding places.

Flights of up to one-half mile have been reported from

French West Africa.

The breeding waters of this species are usually unshaded,

but larvae are apparently tolerant of certain amounts of

shade. They have been found in stagnant or semistagnant

pools, rock pools, residual pools in riverbeds, marshes, hoofmarks, cement tanks with clear rainwater, swamps, and seepage areas. The water in many of these places may be

moving.

This mosquito is comparatively rare in South-West Africa

because of the extensive dry areas in that country.

Relation to Disease.—Up to 1947 this mosquito was regarded as an unimportant vector of malaria, but more recent investigations in French West Africa have shown that

it plays a much more vital part in malaria transmission than

had ever been suspected. Because high sporozoite rates

have been found in that country, rufipes is considered to be

as important as gambiae and Anopheles fumestus there. Itis

important in Anglo-Egyptian Sudan, where it occasionally

appears in large numbers. In Southern Rhodesia, south

Africa, and Kenya it has not been found naturally infected.

References: 119, 167, 168, 258, 259, 260. (For additional information, see the descriptions of malaria under Anglo-Egyptian Sudan ;

Eritrea, the Somalilands, and Ethiopia; West Africa; West-Central

Africa ; East-Central Africa ; Madagascar and Adjacent Islands ; and

Southern Africa. )

Recorded Distribution.—

Anglo-Egyptian Sudan

Bechuanaland

Belgian Congo

Cameroons

Dahomey

Egypt

Eritrea

Ethiopia

French Equatorial Africa

French Guinea

French West Africa

Gambia

Gold Coast

Ivory Coast

Kenya

Madagascar

Mozambique

Natal

Nigeria

Portuguese Guinea

Senegal

Southern Rhodesia

South-West Africa

Transvaal

Uganda

Zanzibar

Anopheles sacharovi Favr

Anopheles sacharovi^ which was originally described from

Caucasia, has been shown to be a member of the A, maculipennis group. Some workers prefer the older name, Anopheles elutus, since they feel that the application of Favr's

name is uncertain. For additional details, see page 125.

Bionomics.—Females of sacharovi are anthropophilic and

the presence of domestic animals does not divert them from

feeding on man to any important extent. Adults are common in houses and stables, sometimes in large numbers, even

in Iraq and northern Iran, where there is some evidence that

most of the females rest outdoors in the summer and indoors

in the winter. In Syria and Lebanon no adults were found

outdoors, nor was any outside feeding demonstrated. Fatbody development has been observed as early as September

in some areas. In Israel, Iraq, and northern Iran prehibernation flights have been observed. At high elevations ±e-
MOSQUITOES OF MEDICAL IMPORTANCE 135

males hibernate in cool shelters and do not take winter blood

meals, but in the lowlands they hibernate in cattle sheds,

stables, and houses, where they feed throughout the winter.

In the Mediterranean area the species breeds typically in

large brackish marshes and less commonly in a variety of

other habitats. These include inland and coastal swamps,

boggy ground in fresh-water areas, neglected water-storage

tanks, and channels and their leakages. Larvae in Syria and

Lebanon are found in both large and small collections of

water covered with floating aquatic vegetation, such as Potamogetón and Ranunculus but rarely in upright emergent

vegetation. Quiet water is preferred but never with much

salinity. In Jordan and Israel larvae commonly live in exposed or partially shaded water collections with the subaquatic vegetation of marshes, lakes, ponds, ditches, and

seepages. In the northern part of the Jordan Valley larvae

are found in rather large bodies of water, and the stream

habitat is rather unusual. In Iraq and northern Iran larvae

inhabit ricefields and large mountain rivers overgrown with

vegetation.

A, sacharovi is a lowland species that likes and withstands

heat. It has occasionally been taken at altitudes up to approximately 3,500 feet in Tadzhikistan. In Syria and Lebanon larvae are found throughout the winter and begin to

appear in large numbers as early as March. Population

peaks of both larvae and adults occur in July and August.

In the northern parts of its range the species is a late-summer

form, but in southern localities there is a rapid increase in

numbers in the spring, a reduction through the summer, and

a later increase in the fall.

Relation to Disease.—A, sacharovi transmits malaria

wherever it is abundant. It is a very important and dangerous vector. It is especially important in northern Italy and

the southern Balkans. In the northern parts of Jordan,

Israel, and Syria the species is associated with very highly

endemic malaria. In Iraq and along the Caspian coast of

Iran sacharovi is almost always associated with high spleen

rates. Itis an important vector as far east as West Pakistan.

There are isolated places, i. e., in the Aimx Darya Valley of

the U. S. S. R., where sacharovi is abundant in the complete

absence of malaria.

References : 5S, 67, 118,147, 229, 239, 2U. (For additional information, see the descriptions of malaria under Northwest Africa ; North

Europe and North Asia; Italy, the Balkans, and Adjacent Islands;

the Near East ; Arabia ; Iran ; and Afghanistan and West Pakistan. )

Recorded Distribution.

Albania

Algeria

Bulgaria

Caucasia

China (north)

Corsica

-Crete

Cyprus

Czechoslovakia

Egypt

Greece

Iran

Iraq

Israel

Italy

Jordan

Lebanon

Libya

Morocco

Rumania

Sardinia

Spanish Morocco ( ?)

Syria

Turkestan

Turkey

U. S. S. R.

Yugoslavia

Anopheles sergenti (Theobald)

Most of the records of Anopheles culicifades in north

Africa west of the Somalilands refer to A. sergenti.

Bionomics.—Adults of sergenti are common in houses and

tents. Precipitin tests show that many females captured in

houses contain human blood. Females bite fiercely after

dark. In Egypt large numbers of adults rest in underground

aqueducts during the spring and summer and feed on man

in ricefields toward the end of the breeding season. Adults

are strong fliers, having been found in rather large numbers

up to 11/2 miles from their nearest breeding places. In Israel

prehibernation flights of up to 334 miles have been reported.

Larvae are most often found among stones in small pools

and springs at the edges of lakes. They also live in seepages,

slow-moving streams, and irrigation channels, where they

avoid direct sunlight. In Egypt larvae are common in ricefields or stagnant water and are sometimes found in borrow

pits associated with vegetation. In Yemen a single larva

was taken at about 3,700 feet in quiet marginal water of a

drying stream in the presence of a fine, green, floating water

grass, and several larvae were observed at a lower altitude in

the presence of short vegetation and filamentous green algae.

Irrigation ditches, swamps, and rock pools are larval habitats

in Syria and Lebanon. A. sergenti is found in breeding

places with Anopheles superpictus throughout Jordan.

A, sergenti is most abundant in September and October

in Israel and Egypt and through November in Jordan. It

passes the winter in the larval stage. Adult populations do

not build up until late summer of the following year.

Relation to Disease.—The seasonal prevalence of this

species over most of its range closely parallels that of malaria.

Because larvae and adults are so universally abundant and

since sergenti is commonly found with natural infections,

it is an important vector in the oases of north Africa east of

Algeria and in Egypt, Israel, western Jordan, and parts of

Arabia. In northern Iran and Iraq this mosquito is too rare

to be an important vector.

References : 91,120, 154, 229, 241. (For additional information, see

the descriptions of malaria under Northwest Africa, Egypt, the Near

East, and Arabia.)

Recorded Distribution.—

Albania Lebanon

Algeria Libya

Arabia Morocco

Canary Islands Pakistan (West)

Egypt Spain

Iran Spanish Morocco

Iraq Syria

Israel Tunisia

Jordan Turkey

Anopheles sinensis Wiedemann

Much of the early medical literature on Anopheles sinensis

in the Orient refers to at least two closely related species

belonging to the A. hy^^canus group. One of these is sinensis

itself, the other is Anopheles nigerrimus (p. 129). Both are

unique in being the only members of the group that are medically important. Eecent investigations have enabled taxonomists to separate and identify the various closely related

species in this group and to assign some early records to the

species as they are now recognized.

Bionomics.—The records of sinensis indicate that this anthropophilic mosquito has definite zoophilic tendencies. In

China it is principally zoophilic but will bite man readily on

occasion. Tendencies toward zoophilism have been also noted

in Indochina. In Malaya and elsewhere females attack man

early in the evening, biting freely outside but rarely inside

habitations. A noticeable reduction in numbers occurs as

early as 9 p. m. Even in areas where females are commonly

found in cattle sheds, they find their way to man-baited net

traps placed out of doors.

Larvae are found most often in ricefields and pools containing quiet water. They also live in swamps, borrow pits,

cisterns, irrigation ditches, and margins of lakes and streams.

In China larvae have been found in stagnant water and old

manure pits containing rainwater. In Okinawa the species

breeds in small clear pools and is seldom found in brackish

water. In Malaya it prefers open, grassy ponds for development and is sometimes found on the coast with Anopheles

sundaicus. Breeding is usually abundant in jungle areas that

have been cleared for cultivation. Larvae are adapted to
136 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

rather open habitats and prefer relatively high water temperatures. Laboratory experiments show them to have

higher thermal death points than the species that normally

live in shade.

Relation to Disease.—In Japan and Formosa this species

is a malaria vector of primary importance. In Formosa it

lives in the plains, being replaced by Anopheles minimus in

the foothills. Together with Anopheles pattoni and A,

maculipennis^ it is probably the only vector of nialaria in

Manchuria and central China. It is the most important

vector in the ricegrowing areas of South China and Formosa,

and in Indochina it is the chief transmitter of malaria in the

ricefields of Tonkin, Cambodia, and Cochin China. In Thailand and Malaya it is regarded as a vector of secondary

importance. In Thailand dissections of wild-caught specimens are sometimes entirely negative for malaria parasites.

Although many records from Sumatra and possibly Borneo

may refer to nigerrimus^ sinensis is regarded as a secondary

vector there. In all areas studied the average natural infection rate is rather low.

Development of both Wuchereria bancrofti and W. malayi

to the infective stage in sinensis has been demonstrated experimentally. This mosquito has been found naturally infected with hancrofti in the Shanghai area of China, where

sinensis is believed to play an important role in filaria

transmission.

References: 61, 91, 122, 28S. (For additional information, see the

descriptions of malaria and filariasis under North Europe and North

Asia, South China, Formosa, Indochina, Thailand, Malaya, and Sunda

Islands.)

Recorded Distribution.

Assam

Burma

Celebes

China

Formosa

Hainan

Indochina

Japan

Malaya

Manchuria

Pakistan (East)

Sumatra

Thailand

U. S. S. R.

Anopheles stephensi mysorensis Sweet and Rao

Bionomics.—Females of Anopheles stephensi mysorensis

are less hardy and can be induced to feed on an arm extended

into a cage with much more difficulty than the type form.

This mosquito prefers rabbit blood to that of man in the

laboratory. Colonies of this subspecies are established with

considerable difficulty.

In some areas the larvae are found only in wells. Elsewhere they live in irrigation channels and at stream margins.

Relation to Disease.—In Vizagapatnam, India, this mosquito is suspected of being the principal vector of malaria.

It is generally considered to be less important than the type

form.

References : 67, 322. ( For additional information, see the description of malaria under India and Ceylon. )

Recorded Distribution.—

India

Anopheles stephensi stephensi Liston

Bionomics.—^Adults of Anopheles stephensi stephensi are

commonly found in houses, cowsheds, barracks, and other

types of manmade shelters. They feed readily on man in

the laboratory and in nature. They are usually difficult to

find, because they hide in small, out-of-the-way places, such

as the many small, dark creases and crevices in folded or

crumpled cloth. Their flight range does not usually exceed

one-half mile. Adults have been found up to l^^^ miles from

their nearest breeding places in Bahrein near the Iraq coas.t.

Larvae are found in both urban and rural locations. The

outstanding type of breeding place consists of wells and

other artificial containers, such as cisterns, water collections

near buildings, flooded cellars, and bases of running fountains. In rural areas larvae are found in all sorts of breeding

places containing fresh, brackish, or sewage-contaminated

water either in direct sun or in the shade. Pools, stream

beds, slowly moving creeks, irrigation channels, drains, and

miscellaneous breeding places with fresh water often contain

larvae. In Iraq the breeding places consist of small, shallow,

grassy pools that tend to disappear later in the summer, especially where palms are being cultivated. Larvae habitually sink deep into the water and stay submerged for long

periods.

In Iraq stephensi breeds throughout the year. It has two

marked seasons of intensity, one in May and June and the

other in October and November.

Relation to Disease.—Except for Anopheles culicifacies^

stephensi is probably the most important vector of malaria

in western and northwestern India. It is commonly encountered in Bombay, Bangalore, and Lucknow, and it is

responsible for the transmission of malaria in urban and

rural areas of Afghanistan and West Pakistan. It is the

only vector usually associated with epidemics on the alluvial

plains of Iraq, which are subjected to intense flooding, and

is by far the outstanding vector along the Persian Gulf

coast, including Al-Qatif and Hofuf oases, and at AlKharijah in the Najd. The species is frequently found with

natural infections.

References : 61, 91, 120, 241, 3^1, (For additional information, see

the descriptions of malaria under the Near East, Arabia, Iran,

Afghanistan and West Pakistan, India and Ceylon, and Burma.)

Recorded Distribution.—

Afghanistan India

Arabia Indochina

Assam Iran

Burma Iraq

China (South) Pakistan (East and West)

Anopheles subpictus subpictus Grassi

Anopheles rossii^ a name commonly used in the past for A,

subpictus subpictus in India, has also been applied to other

species in regions east of that country. Two forms of subpictus^ Anopheles subpictus indeftnitus and A. s. malayensis^ are found in countries east of India and have no known

medical importance.

Bionomics.—Although normally domestic, subpictus is

both zoophilic and anthropophilic in Indochina, where

studies have revealed as few as 2 percent of wild-caught

females with human blood. Adults are repeatedly encountered in large numbers in all kinds of buildings.

Larvae live in many different types of locations containing

fresh, brackish, or polluted water. The breeding places

may be borrow pits, buffalo wallows, brick pits, drains, pools

from leaks in irrigation ditches, furrows in gardens or fields,

roof gutters, ricefields, irrigation channels, and artificial

containers. In the Sunda Islands, especially Celebes, larvae

also breed in the brackish water of lagoons and fishponds,

with salt concentrations of up to 8.5 grams per liter. Larvae

have been taken many times from brackish water in Indochina. In Malaya females most commonly oviposit in fresh

and brackish water in pools, ponds, and open swamps.

Larvae are rarely found in ricefields.

The habitat of subpictus is primarily in coastal and lowland areas. In the Punjab of India the species is found only

during and following the monsoon season, where it first appears in June and is prevalent from July to December. In

southern India and Burma it breeds throughout the year.

Relation to Disease.—This species is of little importance

in malaria transmission, although parasites are able to mature efficiently in it. Few or no natural infections have been

reported outside southeast India, the Sunda Islands, or
MOSQUITOES OF MEDICAL IMPORTANCE 137

Australasia. Java, Sumatra, and Celebes have had epidemics with which both subpictus and Anopheles sundaicus

have been associated. In those areas sundaicus is believed

to be the chief vector and subpictus may become infected late

in the epidemic. On the west and south coasts of Celebes

and in northern Madoera suhpictus is as important as

sundaicus^ and it may transmit some malaria in New Guinea.

A, s. subpictus can be important, in spite of its low infection rates, because of its extremely large populations. There

are no reliable records of natural infections in either the

Philippines or Malaya, where malayensis and indefinitus are

found.

On the basis of experiments with artificial infections,

subpictus is believed to be a good vector of Wuchereria

bancrofti in Batavia, Java. It is a possible vector in India,

but elsewhere its importance is not known.

References: 91, 122, 161. (For additional information, see the

descriptions of malaria under Sunda Islands and Australasia.)

Recorded Distribution.—

Afghanistan Iran

Assam Java

Ban Malaya

Bangka Moluccas

Borneo Nepal

Burma New Guinea

Celebes Pakistan (East and West)

Ceram Philippine Islands

Ceylon Soemba

China (South) Soembawa

Flores Sumatra

Formosa Thailand

India Timor

Indochina

Anopheles sundaicus (Rodenwaldt)

Î

Anopheles sundaicus has commonly been referred to as

Myzomyia ludlowi and A, ludlowi in early malaria

literature.

Bionomics.—Adults of sundaicus are anthropophilic and

zoophilic and are commonly found in both cattle sheds and

houses. Engorged females have often been captured in

rooms and nets in which people have been sleeping. Nevertheless, some feeding on animals is indicated by nearly all

precipitin tests. Females do not bite until several hours

after nightfall and remain for some time near biting places

after obtaining blood meals. They are strong fliers and

have commonly been seen 2 miles, once as far as about 4

miles, from their nearest breeding places.

Except in Sumatra, Java, and restricted parts of Celebes,

sundaicus is known to breed only in brackish water. Larvae

inhabit sunlit lagoons, salt-water fishponds, swamps, and

polluted water on the coast. They are occasionally found

in clear-water pools as well. In Java larvae are commonly

associated with abundant green fishpond algae, such as

Enteromorpha^ which forms a feltlike cover over the water

surface. Although larvae tolerate water with a maximum

of about 30 grams of salt per liter, they are more commonly

found in concentrations of 4 to 18 grams per liter. Deeply

shaded, virgin mangrove areas are unfavorable for larval

development, except where they have been cleared to admit

sunlight. Larvae in fresh-water inland locations in Sumatra, Java, and Celebes do not differ morphologically from

those in coastal brackish-water sites. When fresh-water

fishponds were abolished at Mandahiling, Sumatra, as a

last resort to control sumdaicus^ it disappeared, since it would

not breed in any other water nearby. Larvae have been

found in fresh water in Celebes, but they are rare.

Relation to Disease.—A, sundaicus isregarded as a vector

of utmost importance throughout its range and is one of

the most effective carriers of malaria known. It is very

easily infected in the laboratory. This species has been repeatedly associated with severe periodic outbreaks in coastal

areas throughout the Orient, especially in Java, Sumatra,

and the Lesser Sundas, where natural infection rates as high

as 54 percent have been reported. Extremely high natural

infection rates have been reported from nearly every other

area where this species is found. The percentage rates for

some of these areas are Java 1 to 36, Sumatra 2 to 15 and

once as high as 46, and the East Pakistan-India border 23.4.

Along the coast of Malaya it is responsible for extensive

endemic malaria.

Natural infections of sundaicus with Wuchereria bancrofti

have been reported from various areas. Experimentally

bancrofti is able to complete its development in this mosquito to the infective stage.

References: 91, 120, 122, 161, 228, 242, 324. (For additional information, see the descriptions of malaria under India and Ceylon, East

Pakistan and Assam, Formosa, Burma, Indochina, Thailand, Malaya,

Sunda Islands, and Philippine Islands. )

Recorded Distribution.—

Andaman Islands Hainan

Assam India

BaU Indochina

Bangka Java

Borneo Nicobar Islands

Burma Pakistan (East)

Celebes Philippine Islands

Ceram Soembawa

China (South) Sumatra

Flores Thailand

Formosa Timor

Anopheles superpictus Grassi

Bionomics.—Adults of Anopheles superpictus habitually

fly great distances to congregate in localities where fairly

large groups of host animals are present, but no long-range

dispersals have been noted except those preliminary to winter

hibernation. Females are commonly found inside habitations of all kinds and readily feed on man, even when livestock is in the immediate vicinity. Before and immediately

after taking blood meals they fly to animal shelters to rest

on spider webs and in dark corners, and they have been

found in outdoor resting places as well. They are very resistant to dry air. In Israel their normal range is about 1

to 1% miles from their breeding places. Once in Italy a

flight range of 4.3 miles was noted. Partly or completely

hibernating females have been found in the winter in stables

and houses. They exhibited a full fat-body development

as ào^^ Anopheles maculipennis, A. superpictus overwinters

as an adult in most of its range.

Larvae are found in the open sunlit pools left in beds of

small mountain or hill streams during the dry season. They

also are found in gently flowing water, especially in bays or

where the edges ai?e weedy. Larvae have been found in

streams with a central velocity of 2.1 miles per hour. In

Italy the species is common in ricefields and comprises up

to 10 percent of all anophelines found there. Larvae never

live in foul water but often inhabit comparatively quiet

bodies of water, with a saline concentration of up to 0.5

percent. They may lie submerged for up to 8 minutes in

a collecting container. During the winter they have been

found under or enclosed in ice.

A, superpictus normally lives at altitudes up to 5,000 feet

and has been found as high as 7,000 feet. It requires a warm,

dry summer and is everywhere most prevalent during the

hottest, driest parts of the year. It does not survive a prolonged winter. Its northern limit of distribution is determined by the amount of precipitation, since excessive rains

tend to destroy its breeding places. In northern parts of

Asia superpictus is a late-summer species, but it appears in

large numbers earlier in the season in the south. In Turkey

it reaches a peak of abundance in July and August.
138 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

Relation to Disease.—This species is almost always a

dangerous malaria vector wherever itis sufficiently abundant.

It has been easily infected in the laboratory, and it has been

found with natural infections in many parts of its range.

It can maintain highly endemic malaria in the complete

absence of other vectors and is one of the most widely known

malaria vectors.

References: 67, 91, 120, 217, 346. (For additional information, see

the deKScriptions of malaria under Northwest Africa; Egypt; North

Europe and North Asia ; Italy, the Balkans, and Adjacent Islands ;

the Near East; Iran; and Afghanistan and West Pakistan.)

Recorded Distribution.—

Afghanistan Iraq

Albania Israel

Algeria Italy

Arabia Jordan

Balearic Islands Lebanon

Bulgaria Libya

Canary Islands Pakistan (V^est)

Caucasia Spain

Crete Syria

Cyprus Tunisia

Egypt Turkestan (?)

Greece Turkey

India Yugoslavia

Iran

Anopheles tessellatus Theobald

Very early records of Anopheles tessellatiM in the eastern

part of the Orient are found under the name A. punctulatus.

Bionomics.—Although females readily feed on man, tessellatus is a semidomestic and zoophilic mosquito. Females

have been caught in human-bait traps in Malaya, but three

investigations in Indochina showed very low percentages of

females with human blood. In India the species rests in both

houses and cowsheds and is often reported feeding on buffalo.

In the Philippines resting adults have been captured at

eroded banks of streams and in damp, shaded cracks of stone

walls. The flight range of adults in the Sunda Islands is

about 1,100 yards. In Philippine experiments an adult was

recorded over a mile from the point of liberation.

In some areas clear water is the larval habitat, but in others

dirty stagnant water and sometimes mud puddles of limited

size are preferred. In Malaya more larvae are found in

drains and swamps than in ground pools. Almost no collections have been made from ricefields and streams. In the

Philippines, on the other hand, the species is most abundant

in ricefields, and smaller numbers of larvae live along vegetated margins of streams, springs, and irrigation ditches

containing clear water. Over most of its range it breeds in

either shaded or open locations.

Relation to Disease.—This mosquito is an important

vector of malaria in Formosa, where experimental infections

have been demonstrated and where infected females are often

found in nature. The rate of natural infections is too low to

implicate this species in the transmission of malaria

elsewhere.

Experiments indicate that tessellatus may be a good vector

of Wu^hereria hancrofti in Java. There have been no confirmatory field investigations.

References: 91, 120, 122. (For additional information, see the description of malaria under Formosa. )

Recorded Distribution.—

Andaman Islands India

Assam Indochina

Bali Java

Borneo Malaya

Burma Moluccas

Celebes New Guinea (?)

Ceram Pakistan (East)

Ceylon Philippine Islands

China (South) Soembawa

Formosa Sumatra

Hainan Thailand

Anopheles umbrosus Theobald

Most early records of Anopheles umbrosus refer to a large

number of closely related species sometimes known as the

''Anopheles umbrosus group," which contains three medically

important species: A. umbrosus (see below), A. letifer

(p. 124), and A. baezai (p. 114). As a result of recent studies

by Reid and Hodgkin {28Jf)^ umbrosus has recently been

redefined, and in this restricted sense the following remarks

are made.

Bionomics.—A, umbrosus is the only species belonging to

the umbrosus group that bites during the day. It bites outdoors in the jungle, accompanied by Mansonia {Mansonoides) longipalpis^ but it enters houses, often one-half mile or

so from the edge of the jungle, and bites indoors during the

night. When in houses it will attack boldly, even under electric lights. In Malaya this mosquito is markedly anthropophilic. The adults rest on jungle vegetation near their

breeding places.

Larvae are found almost entirely within the jungle in

brown, peaty water. Elsewhere they invariably are observed

in heavy shade. They prefer small pools along the borders

of slow-moving streams.

A. umbrosus is a coastal-plain mosquito, found principally

in dense, swampy jungle and occasionally in adjoining foothills.

Relation to Disease.—A. umbrosus has long been recognized as an important vector of malaria. Where letifer and

umbrosus females occur together while biting humans outside jungle areas, letifer may be the more important vector.

However, there is little doubt that umbrosm is a vector of

considerable importance, especially in the western coastal

areas of Malaya, where dissections of numerous wild-caught

females since 1936 show a salivary gland infection rate of

about 0.5 percent. In several areas of Borneo umbrosus has

been found with salivary gland infections and has definitely

been incriminated as a vector of malaria. Several positive

dissections recorded from the Sunda Islands probably include species other than umbrosus. In south Celebes this

mosquito is not regarded as a vector.

References: 66, 161, 2JfS, 28Jf. (For additional information, see

the deiscriptions of malaria under Malaya and Sunda Islands.)

Recorded Distribution.—

Andaman Islands

Assam

Bangka

Borneo

Indochina

Java

Malaya

Sumatra

Celebes

Anopheles varuna lyengar

Anopheles varumc is closely related to A, minimus and has

often been treated as a variety of that species. Many records

of minimus in southern India possibly refer to varuna. Some

records of Anopheles listonii, especially those pertaining to

breeding in wells, probably refer to varuna.

Bionomics.—Adults of varuna have been captured m

houses and cattle sheds. In some localities females definitely

prefer the blood of man; in others cattle blood is preferred.

Except in India, where this species most commonly breeds

in wells, larvae inhabit stagnant fresh-water ponds and

ditches during and shortly after the monsoon season. They

are commonly collected at roadsides from small pools ot

storm water in borrow ditches and from slow-running

streams. <»

Relation to Disease.—^, varuna is a proved vector of

malaria in restricted areas of east-central India. It is ot

little importance elsewhere, either because of its preference
MOSQUITOES OF MEDICAL IMPORTANCE 139

for feeding on cattle or its scarcity. It has been found infected in nature.

References: 67, 91, 121, 161, 303. (For additional information, see

the description of malaria under India and Ceylon. )

Recorded Distribution.

Assam

Burma

Celebes

Ceylon

India

Indochina

Pakistan (East)

Sumatra

Anopheles vestitipennis Dyar and Knab

Bionomics.—Females of Anopheles vestitipennis are endophilic and anthropophilic. Experiments in Cuba show

that vestitipennis prefers human to calf blood but might

prefer the blood of other animals to either of them. In the

rural areas of Stann Creek Valley, British Honduras, this

species was much more common in houses than in livestock

traps nearby.

Larvae are found in cool fresh-water streams, seepages,

ponds, and rain pools with abundant shade. In Puerto

Rican sugarcane fields they are associated with Anopheles

grabhamii in stagnar^t ditches filled with all kinds of vegetation. In Cuba larvae are found in partial shade in freshwater swamps some distance from the coast, primarily in

the western half of the island, where dense vegetation,

detritus, and short grass cover areas of open water. Small

clumps of coarse grass harbor large numbers of larvae in

flooded forest rain pools of British Honduras.

A comparatively rare species over most of its range,

vestitipennis is most abundant near isolated swampy and

marshy areas.

Relation to Disease.—This species can be infected experimentally with malaria parasites, although parasite development, at least in Cuba, appears to be slow. In that

island vestitipennis is suspected of being a malaria vector,

but its role in the presence of Anopheles alhimanus is not

known. It has been found with natural infections in British

Honduras where active cases of malaria are present.

References: 78, 221, 226, 292, 308. (For additional information, see

the description of malaria under Central America.)

Recorded Distribution.—

British Honduras

Colombia

Costa Rica

Cuba

Dominican Republic

Guatemala

Haiti

Honduras

Jamaica

Mexico

Panama

Puerto Rico

Trinidad

Windward Islands

CULEX SPECIES

Culex pipiens pallens Coquillett

Bionomics.—Gulex pipiens pallens is one of the most

abundant mosquitoes in the Shanghai area, where female-^

avidly attack man.

Larvae are found in polluted water from March to December in that area.

Relation to Disease.—Experimental infections with

Wuchereria hancrofti have revealed that pallens is probably

one of the most efficient vectors of filariasis in the Shanghai

area. It is not susceptible to Wuchereria malayi infection in

the laboratory.

References : 111, 183,181,189. (For additional information, see the

description of filariasis under North Europe and North Asia. )

Recorded Distribution.—

China

Japan

United States (introduced into California)

Culex pipiens quinquefasciatus Say

Culex fatigans^ the name used for C. pipiens quinquefasciatus by most European workers, is commonly found in

the literature pertaining to the vector characteristics of the

latter.

Bionomics.—C. p, quinquefasciatus is one of the most common mosquitoes found in human habitations in the Tropics

and Subtropics of the world. In most of its range females are

intensely anthropophilic and feed actively only at night.

Even in the laboratory females bite only at night except when

they have undergone a period of starvation. In parts of

Africa they are attracted to natives rather than to Europeans. Females of quinquefasciatus rest in or under all kinds

of buildings near humans, especially where the air is humid.

They lay from 2 to 4 rafts of eggs, usually at night but rarely

in broad daylight. Adults have been observed to fly as far

as 3 or 4 miles from their nearest breeding places.

Larvae are never found far from human habitations. They

live in flooded, open cement drains, cesspools, latrines, shallow

wells, ditches, and ground pools. They are common in household storage water and other collections of water close to

houses, especially water that is poorly covered and is polluted.

In the lower Congo River Valley larvae are commonly found

in poorly drained land surrounding villages. In India they

commonly inhabit ricefields near villages and are especially

common where rice is fertilized with green manure. Immature stages can adapt to a high degree of salinity and can

easily survive in water with a pH of 5 to 10.

Relation to Disease.—In laboratories in many parts of the

world Wuehereria hancrofti has been found to mature in

quinquefasciatus to the infective stage. This mosquito is

the only vector of Bancroftian filariasis in the New World

and an important vector in many parts of the Old World,

including tropical Africa. It is not a good vector of Wuchereria malayi^ Sind in areas where that parasite predominates,

quinquefasciatus may not be an important vector of ifilariasis.

References : 61, 95, 122, 166, 194, 250, 255, 268, 338. (For additional

information, see the descriptions of filariasis under Central America,

Brazil and the Guianas, Northwest Africa, Egypt, West-Central Africa,

East-Central Africa, India and Ceylon, East Pakistan and Assam,

Formosa, Indochina, Thailand, Sunda Islands, and Australasia.)

Recorded Distribution.—The distribution of this species

is worldwide in the Tropics and Subtropics. In the New

World it is found between latitude 40° N. and latitude 35° S.

Its range encompasses almost all the countries of continental

Africa south of the Sahara Desert and Egypt, and it extends

east across the southern half of the Arabian peninsula and the

Orient, including India, to the China coast. This species

inhabits Kyushu in the Japanese islands, Formosa, and all

the Sunda Islands. It is found almost everywhere in

Australasia.

Culex tarsalis Coquillett

Bionomics.—Principally zoophilic in its feeding habits,

Culex tarsalis prefers the blood of wild and domestic fowl

and attacks horses, cows, pigs, chickens, and sheep. It only

occasionally seeks the blood of man. It feeds principally at

dusk or at night, and the bite is painful. ' Females are attracted to light and enter vehicles and dwellings readily.

In California this mosquito will fly as far as 21/2 miles and

has an effective flight range of about 1 mile. Females overwinter in buildings, caves, and cellars. In hot, dry localities

on the Pacific Coast and in the Southwestern States adults

appear between April and October and reach a peak of abundance in July and early August. East of the Mississippi

River and south of Michigan and Illinois this peak appears

in August and September, and the mosquitoes are present

until December in some localities.

Larvae are found in many types of breeding places. These

may be in water that is clear, fresh, alkaline, or stagnant with
140 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

organic matter, and they may be open smilit or shady pools

of various sizes. The type of breeding place varies with the

locality. In Lower California larvae have been reported

from a brackish tidal marsh, in the hot Southwest they are

common in stagnant water near cultivated areas, on the

Canadian prairie they are most common in reedy roadside

ditches, in the Pacific Northwest they are found in poorly

drained irrigated areas, and in the West it is the most common mosquito breeding in thermal waters with temperatures

from 82.4° to 102.2° F.

¿7. tarsalis has been reported as the most abundant mosquito in the Pacific coastal region and the most ubiquitous

mosquito in California, but it becomes more scarce in the

Central and Eastern States. Because of its wide variety of

breeding places, tarsalis builds up large populations in years

of moderately heavy rainfall.

Relation to Disease.—Aedes dorsalis^ A, vexans^ A, nigromaculis^ and Culiseta melanura have been incriminated in

the transmission of western equine encephalitis, but C, tarsalis is unquestionably the most important vector of this

virus. The biting activity of tarsalis and this virus disease

are distinctly correlated. Over 100 strains of the virus have

been isolated from wild-caught tarsalis adults in various

parts of the United States. The distribution of tarsalis and

the appearance of this virus in horses are strikingly related.

Although this species harbors St. Louis encephalitis virus

in nature, its exact role in transmitting it is not well known.

Reference: 200. (For additional information, see the descriptions

of encephalitis under America North of Mexico and Mexico.)

Recorded Distribution.—

Canada

Mexico

United States

Culex tritaeniorhynchus Giles

Bionomics.^InJapan Culex tritaeniorhynchus is the most

important night biter during July and August. Its most

intense biting begins just after sunset and lasts as late as

9 p. m., although females bite throughout the night. A comparison of animal-trap catches in Japan showed that when

humans were used as bait, about 75 percent of all catches

were tritaeniorhynchus mosquitoes ; when horses were used,

89 percent; and cows, 93 percent. This mosquito, therefore,

is not definitely anthropophilic in its biting habits, and, as

with Culex tarsalis^ man is nearly always an accidental host.

The adults in Okinawa are reported to rest on damp rocks

in woods and in vegetation near the ground. They commonly attack horses.

Early in the breeding season larvae are found in freshwater collections, such as natural- and artificial-water impoundments, ground pools, field drainage, and irrigation

ditches. The species spreads to diverse habitats as the season progresses and inhabits polluted breeding places later in

the year.

In Japan ricefields are said to be preferred breeding

places. However, the drainage of ricefields is not necessarily

correlated with the reduction in mosquito abundance. In

Japan most larvae disappear by October. Population intensity appears to be more closely related to rainfall than to

temperature. In Okinawa the habits are more diversified

than those of any other mosquito, and larvae are found in all

kinds of water collections. In Yemen larvae were observed

in a drying stream where the quiet marginal water had short

emergent vegetation and filamentous green algae. In Leyte

larvae were found in a variety of locations with or without

vegetation, and all were exposed to sunlight. In central

China larvae have been reported from bomb craters and ricefielus containing decayed vegetation, usually in complete

shade.

This species is abundant in Korea, Japan, and eastward

through central Ohina. In Korea the population peak does

not appear until mid-August, in Okinawa maximum abundance is reported to occur in early September, and in central

Ohina large numbers of adults are found in October. Population densities in these areas appear to be higher in rural

than urban localities, apparently because ricefields are the

habitat.

Relation to Disease.—Evidence indicates that this species

is the most important, if not the only, vector of Japanese

encephalitis virus in Japan. The virus has been found in

wild-caught mosquitoes during the peak of biting activity

in areas where the virus exists, and mosquitoes have successfully transmitted it in laboratory experiments under a variety of conditions. Because Japan is the only locality in

which extensive experiments have been carried out and because there is no direct evidence that this species actually

transmits the virus elsewhere, its importance as a vector in

countries outside Japan can only be surmised. As far as

is known, it is presumed to be the vector in any area where it

and Japanese encephalitis appear together. Thus in Korea

it is said to be probably the most important vector and the

only one definitely associated with the disease there. Epidemiological evidence indicates that tritaeniorhynchus is a

vector in Okinawa, although Culex pipiens quinquéfasciatus

has been suspected in the past. In Guam the vector is not

known, and tritaeniorhynchus is not present there. The disease is not definitely known to exist in Manchuria, but this

species is present in northern China.

References: S9, 51, 62, 154, 176, 206, 223, 272, 332, 348. (For additional information, see the descriptions of encephalitis under Europe

and North Asia, and Australasia. )

Recorded Distribution.—

Arabia Java

Borneo^ Lebanon

British Somaliland Madagascar

Celebes Manchuria

Ceylon Mascarene Islands

China (northern and South) Mozambique

Dahomey Nigeria

Egypt Okinawa

Flores Island Philippine Islands

Gold Coast Sumatra

India Syria

Iran Tanganyika

Iraq Turkey

Israel Zanzibar

Japan

Culex univittatus Theobald

No previous names have appeared in the medical literature

for Culex univittatus, but a subspecies, C, u. neavei, is distributed along the White Nile Eiver.

Bionomics.—Adults of univittatus are most common during the summer and early fall in Egypt and Anglo-Egyptian

Sudan. Females appear to prefer birds as sources of blood

meals, since bird-baited traps attracted this species almost

exclusively. Observations also indicate thatthe species feeds

on domestic animals. Furthermore, precipitin tests showed

that only 11 percent of blooded univittatus females caught

in light traps contained human blood. There is no evidence

to indicate that females hibernate during the winter.

Larvae are found in swampy places during the entire year.

Relation to Disease.—Eecent investigations indicate that

univittatus is the principal vector of West Nile virus along

the Nile River and Delta in Egypt and in southern AngloEgyptian Sudan. The species has been incriminated on the

basis of epidemiolo^ical evidence and because it has been

found more heavily infected with the virus in nature than
MOSQUITOES OF MEDICAL IMPORTANCE

any other mosquito tested. In the laboratory this species

has transmitted the recently discovered Sindbis virus (see

p. 4).

References : 325, 326. ( For additional information, see the descriptions of encephalitis under Egypt and Anglo-Egyptian Sudan.)

Recorded Distribution.—

Anglo-Egyptian Sudan

Egypt

HAEMAGOGUS SPECIES

Haemagogus albomaculatus Theobald

Bionomics.—Although adults of Haemagogus albomaculatus are most active from 9 to 11 a. m. in bright sunlight,

they may also fly during the evening and at night. As long

ago as 1917 females were reported to enter houses arid bite

man, but this habit has not been confirmed recently.

Practically nothing is known about the larval habitats of

this species.

Relation to Disease.—H, albomaculatus is probably a vector of jungle yellow fever. Investigations in British and

French Guiana and Surinam have shown it to be the only

species of Haemagogus found in certain localities where almost half the native population was immune to the virus.

References: 113, 211. (For additional information, see the description of yellow fever under Brazil and the Gulanas. )

Recorded Distribution.—

Biritish Guiana

French Guiana

Surinam

Haemagogus capricornii Lutz

Haemagogus capricornii for many years was confused in

the taxonomic and medical literature with H. spegazzinii

spegazzinii and H, equinus.

Bionomics.—Almost nothing is known about the bionomics

of capricornii. It probably does not exist in areas with an

annual rainfall of less than about 40 inches.

Relation to Disease.—H, capricorniiis unquestionably implicated in the transmission of jungle yellow fever. It transmits the disease in the laboratory and is suspected of being

naturally infected. In Brazil specimens of spegazzinii were

found infected in nature. Actually capricornii might have

been the species, since males of the latter were later found

among the spegazzinii specimens from which the virus was

originally isolated.

References: 2m, 337. (For additional information, see the description of yeUow fever under Northwest South America.)

Recorded Distribution.—

Argentina

Bolivia

Brazil

French Guiana

Haemagogus equinus (Theobald)

Bionomics.—A distinctly arboreal mosquito, Haemagogus

equinus has been shown by studies in Panama to predominate

at various levels above the ground but slightly less so than

H. spegazzinii falco. During the rainy season equinus has

a high, well-marked peak of abundance, whereas during dry

weather it may disappear altogether.

Larvae are principally tree-hole inhabitants but may be

found occasionally in artificial containers near human habitations.

This is the most widespread of all Haemagogus species in

Panama, Central America, and Mexico, because it can live

in many different kinds of habitats. These include tropical

141

rain forest, tropical deciduous forest, thorny scrub in the

more northern parts of its range, and mangrove.

Relation to Disease.—ZT. equinus has never been found

with confirmed infections of yellow fever virus in nature.

However, experiments show that it can easily transmit the

virus in the laboratory, and transmission rates of up to 0.45

have been found, compared with a transmission rate of 1.00

for Aedes aegyptL The maximum number of females attacking man in certain areas of Panama is about the same

as that in areas of endemic yellow fever in Brazil.

References: 133, 112, 220, 330. (For additional information, see

the descriptions of yellow fever under America North of Mexico, Mexico, Central America, Panama, Northwest South America, and Brazil

and the Guianas.)

Recorded Distribution.—

Argentina Mexico

British Guiana Nicaragua

Colombia Panama

Costa Rica Surinam

El Salvador United States

Guatemala Venezuela

Honduras

Haemagogus mesodentatus gorgasi Galindo and Trapido

and Haemagogus mesodentatus mesodentatus Komp

and Kumm

Haemagogus mesodentatus is a complex of closely related

species and subspecies. The following remarks refer to the

original form found in Costa Rica by Komp and Kumm.

Bionomics.—Adults of both Haemagogus mesodentatus

gorgasi and H, m. mesodentatus are principally arboreal.

The typical subspecies m. mesodentatus inhabits the tropical

rain forest of the Caribbean slope, and gorgasi lives principally in the deciduous forest on the Pacific side.

Almost nothing about larval habitats appears in the literature.

Relation to Disease.—Both these subspecies have been

good laboratory vectors of yellow fever, but none were found

w^th natural infections.

Reference : 330. ( For additional information, see the descriptions

of yellow fever under Mexico, Central America, and Panama.)

Recorded Distribution.—

H. m. gorgasi:

El Salvador

Guatemala

Mexico

H. m. mesodentatus:

Costa Rica

Honduras

Mexico

Nicaragua

Panama

Haemagogus spegazzinii falco Kumm et al.

Haemagogus spegazzinii falco has been listed in recent

literature under the names H, janthinomys and H. capricornii.

Bionomics.—H, s. falco is an arboreal, diurnal forest

mosquito that feeds avidly on man and has a peak of activity

about midday in subdued light. Females are most abundant in open, sunny clearings or at forest margins ; males are

rarely seen in nature. Eain-forest studies in Panama show

that half of the falco females live at high elevations and only

about 20 percent close to the ground. In deciduous forest or

where large trees have been thinned out, females will attack

at ground level. This species was not seen at all during the

dry season in February, March, and April in Panama. It

is not so abundant as other vectors, such as Haemagogus

equinus or Aedes leucocelaenus^ although it reaches a high

population peak during the rainy months of the year.

Larvae are most often found in tree holes with elongated

and very narrow apertures. Since so few records of larval
142 AGRICULTURE HANDBOOK 15 2, U. S. DEPT. OF AGRICULTURE

habitats appear in the literature, the important sources of

breeding obviously have been overlooked in most forest

studies. In Colombia large numbers of larvae have been

found in tree holes, rotted tree stumps, and in water held in

fallen-tree buttresses, which are all rather unusual breeding

places for falco.

H. s. falco most commonly lives on mountain slopes from

300 to 1,500 feet, but once it was found at 4,200 feet.

Relation to Disease.—This mosquito has been found infected with yellow fever virus in nature several times and is

easily infected in the laboratory, where a transmission rate

of 0.73 was found for falco and capricornii together as compared with that of 1.00 for Aedes aegyptL H. s, falco has

been incriminated as a vector of jungle yellow fever in several parts of South America. It frequents many endemic

yellow fever areas of Colombia and is presumed to be important in all parts of its range where it is abundant.

References: 55, 130, 133, 112, 210, 220, 330. (For additional information, see the descriptions of yellow fever under Central America,

Panama, Northwest South America, and Brazil and the Guianas,)

Recorded Distribution.—

Brazil Honduras

Colombia Nicaragua

Costa Rica Panama

Haemagogus spegazzinii spegazzinii Brèthes

Considerable confusion has existed regarding the nomenclature of Haemagogus spegazzinii^ since its two subspecies,

H. s. spegaBBinii and H, s. falco, are very closely related to

each other and to H. capricornii.

Bionomics.—Adults of spega^sdnü are predominantly arboreal. In flight studies they have been recaptured more

than 3,000 feet from the point of liberation in the direction

of the prevailing wind. Additional studies show that this

mosquito may fly as far as 7 miles from its breeding place.

Studies on the biology of this mosquito in Brazil have revealed very few larval habitats. Cut sections of bamboo

have been suspended at various levels in forest areas, but

these have failed to attract ovipositing females. Those few

tree holes in which larvae have been found were small and

had extremely small apertures. The scarcity of this subspecies during the dry season may be explained by the

fact that these small holes dry out very rapidly anS

thoroughly.

Relation to Disease.—H. s. spegaBzinii has been incriminated as a vector of jungle yellow fever in several parts of

South America, especially in Ilheus, "Anado," and Goias in

Brazil, where it is the only vector in areas where yellow fever

is endemic. It is regarded as the most important vector of

jungle yellow fever in Brazil today.

References: Sl-83, 130, 218. (For additional information, see the

descriptions of yellow fever under Northwest South America, and

Brazil and the Guianas. )

Recorded Distribution.—

Argentina Panama

Bolivia Trinidad

Brazil Venezuela

French Guiana

MANSONIA SPECIES

Mansonia annulifera (Theobald)

Bionomics.—Almost nothing is known aboutthe bionomics

of Mansonia annulifera adults. They are known to be

strongly anthropophilic.

The sij)hon of this and other Mansonia larvae is specially

adapted for puncturing underwater stems of aquatic plants.

Larvae obtain oxygen for respiration from the air passages

of these submerged plant tissues. Larvae are found mostly

in ponds, pools, backwaters, and marshes that favor the

growth of Pistia spp. and Eichhomia spp.

Relation to Disease.—M, annulifera is probably the most

important vector of Wuchereria malayi in the Orient,

Greater Sunda Islands, Philippines, and Merauke in New

Guinea. Although this species has been found with natural

infections of Wuchereria hancrofti at Travancore, India,

these parasites are probably unable to mature in annulifera

elsewhere in its range.

Reference : 122. ( For additional information, see the descriptions

of filariasis under India and Ceylon, East Pakistan and Assam, Indochina, Thailand, and Sunda Islands.)

Recorded Distribution.

Borneo

Burma

Celebes

India

Indochina

Java

New Guinea

Philippine Islands

Sumatra

Thailand

Mansonia indiana Edwards

Bionomics.—Little is known of the bionomics of Mansonia

indiana adults. However, females are known to be strongly

anthropophilic.

Larvae have been found only in association with Pistia

spp. (see also p. 143).

Relation to Disease.—M. indiana is described in the literature as an efficient and important vector of Wuchereria

malayi in Java. Under experimental conditions in Indochina, Wuchereria hancrofti has matured in indiana to the

infective stage, but the extent to which this mosquito transmits hancroftiin nature has not been thoroughly investigated.

Reference: 122. (For additional information, see the descriptions

of filariasis under East Pakistan and Assam, Indochina, ThaUand,

and Sunda Islands.)

Recorded Distribution.—

Borneo

India

Indochina

Java

Sumatra

Thailand

Mansonia longipalpis (van der Wulp)

Mansonia longipalpis was originally described as a Culex

species. The name Mansonia annulipes, which should not be

confused with M. annulifera, has also been used for this

species.

Bionomics.—Adults of longipalpis have been found at considerable distances from their known breeding places. Females are anthropophilic.

Larvae are associated with Pistia spp. (see also p. 143).

Relation to Disease.—ü/. longipalpis is an important vector of Wuchereria malayi in parts of the Orient, Sunda Islands (especially Sumatra and the west coast of Celebes),

Philippines, and Borneo. W, malayi has developed rapidly

in longipalpis to the infective stage in the laboratory, and

this mosquito has been found infected with parasites many

times in nature. However, Wucherema hancrofti is apparently unable to mature to the infective stage in longipalpis.

Reference: 122. (For additional information, see the descriptions

of filariasis under ThaUand, Malaya, and Sunda Islands.)
MOSQUITOES OF MEDICAL IMPORTANCE 143

Recorded Distribution.

Borneo

Burma

Celebes

India

Java

Malaya

New Guinea

Philippine Islands

Sumatra

Thailand

Mansonia uniformis (Theobald)

Bionomics.—Adults of Mansonia uniformis are active

night and day. Females are persistent biters of man, attacking indoors and outdoors in large numbers. They have

been observed to fly during heavy rainstorms.

Although Pistia spp. are the important host plants for

uniformis in the Philippines, larvae have been found associated with the following species of aquatic plants in Java :

''Adontella leptostachya''

Azolla pinnata

Eclipta alha

Eichhornia crassipes

Hygrophila quadrivalvis

HymenacUne ampleœicaulis

H. interrupta

Isachne miUacea

Jussiaea repens

Leersia hewandra

Lemna paucicostata

Nymphaea stellata

Oryza s^ativa

Pistia stratiotes

Salvinia rotundifoUa

Spirodela polyrhiza

Larvae have also been collected from mangrove swamps in

Papua.

Relation to Disease.—M, uniformis is an important vector

of Wuchereria malayi in south Borneo and elsewhere in the

Sunda Islands. In China it is able to transmit malayi hut

is a less effective vector than Anopheles sinensis there. Experiments have shown that uniformis may become infected

with Wuehereria hancrofti^ but this mosquito is not an important vector of that filarla in nature.

Reference: 122. (For additional information, see the descriptions

of filariasis under India and Ceylon, East Pakistan and Assam,

South China, Indochina, Thailand, and Sunda Islands.)

Recorded Distribution.—

Australia

Borneo

Burma

Celebes

Ceram

China (eastern and South)

India

Indochina

Java

Malaya

New Guinea

Philippine Islands

Sumatra

Thailand

SABETHES SPECIES

Sabethes chloropterus (Humboldt)

Bionomics.—Sabethes chloropterus breeds in tree holes

in both tropical rain and deciduous forests. Adults are definitely arboreal. The species is able to survive seasons that

are unfavorably dry for other vectors of yellow fever. It

is rare at altitudes over 3,000 feet.

Relation to Disease.—Because it occurs in large numbers

in endemic yellow fever areas, and because it transmits yellow

fever virus in the laboratory, chloropterus is considered to

be an important vector of the disease in lower Mexico, Central

America, and Panama. Its importance elsewhere has not

been investigated.

References : 226, 331. (For additional information, see the descriptions of yellow fever under Mexico, Central America, and Panama.)

Recorded Distribution.—

Argentina Guianas

Bolivia Mexico

Brazil Nicaragua

Colombia Panama

Costa Rica Trinidad

El Salvador Venezuela

Guatemala

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GALLIARD, H.—Continued

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1937. NOTES ON FILARIASIS AND ITS TRANSMISSION BY MAURITIAN ANOPHELiNEs. Roy. Soc. Trop. Med. and Hyg.

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1955. STUDIES ON THE VECTORS OF WUCHERERIA BANCROFTI IN

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1953. DISTRIBUTION OF PRINCIPAL MALARIA VECTORS IN IRAN.

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1942. ON THE HABITS AND BREEDING-PLACES OF AEDES

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1949. CONSPECTUS OF MALARIA INCIDENCE IN NORTHERN EUROPE,

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1948. JAPANESE B ENCEPHALITIS. 4th Internatl. Cong. Trop.

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1953. THE CURRENT STATUS AND PROBABLE FUTURE OP JUNGLE

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1942. NOTES ON CULICINE MOSQUITOES IN TANGANYIKA TERRITORY. Bul. Ent. Res. 33:181-193.

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1940. DISTRIBUTION OP FILARIASIS IN TANGANYIKA TERRITORY,

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1949. AN EPIDEMIC OP MALARIA IN THE KENYA HIGHLANDS

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1952. THE ANOPHELINE FAUNA AND MALARIA VECTORS IN SOUTH

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1939. MEDICAL ENTOMOLOGY. Ed. 3, 603 pp. New York.
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1939. TRANSMISSION OF MIOROFILARIA MALAYI IN MALAYA.

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1950. UN NOUVEAU VECTEUR DU PALUDISME EN A. O. F., ANOPHELES RUFiPES (GOUGH) 1910. Bul. Soc. Path. Exot.

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1952. MOSQUITOES OP THE ETHIOPIAN REGION, I. LARVAL BIONOMICS OF MOSQUITOES AND TAXONOMY OF CULICINE

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1953. DENGUE EPIDEMICS IN JAPAN, 1042-1945. Jour. Trop.

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1946. COMPARISONS OF MATING BEHAVIOR, GROWTH RATE, AND

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1935. PRELIMINARY OBSERVATIONS ON THE LONGEVITY OF INFECTIVE LARVAE OF WUCHERERIA BANCROFTI COBBOLD IN

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Hu, S. M. K.—Continued

1940. STUDIES ON THE SUSCEPTIBILITY OF SHANGHAI MOSQUITOES TO EXPERIMENTAL INFECTION WITH WUCHERERIA

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1941. STUDIES ON THE SUSCEPTIBILITY OF SHANGHAI MOSQUITOES TO EXPERIMENTAL INFECTION WITH MICROFILARIA

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1944. STUDIES ON THE SUSCEPTIBILITY OF SHANGHAI MOSQUITOES TO INFECTION WITH MICROFILARIA MALAYI BRUG.

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1953. MOSQUITO SURVEY OF GUAM. Mosquito News 13: 123-

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1933i OBSERVATIONS ON NATURAL INFECTIONS OF CULEX PIPIENS

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1919. THE DOMESTIC BREEDING MOSQUITOES OF THE NORTHERN

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1932. FILARIASIS IN NORTH TRAVAN CORE. Indian Jour. Med.

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1936. SOME OBSERVATIONS ON THE OCCURRENCE OF FILARIAL

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GLOSSARY

Alluvial.—Eelated to deposits of soil made by flowing

water.

Anthropophilism, anthropophilic.—A feeding habit in

which the blood of man is preferred over that of other animals (seezoophilism).

Arboreal.—Tree inhabiting.

Breeding place.—A location, usually on or near water,

where female mosquitoes lay eggs and in which immature

stages (larvae and pupae) develop.

Crepuscular.—Active during twilight hours.

Diurnal.—Active by day.

Endemism, endemic.—Pertaining to or prevalent in a

particular area. A situation in which a disease has a low

incidence but is constantly present in a given area. Active

transmission of the disease need not täte place during the

entire year (see hyperendemism).

Endophilism, endophilic.—A condition in which mosquitoes prefer to rest and feed indoors.

Epidemic.—A situation in which a disease attacks many

people in a given area at the same time or within a very short

period of time. Usually the susceptibles contract the disease

in a much shorter period than that required for a state of

hyperendemicity (see endemism and hyperendemism).

Epidemiology, epidemiological.—A study of all factors

bearing on the cause and transmission of disease. Consideration is given in these studies to (1) the causative agent and

its biology; (2) the identiñcation, habits, and relative importance of the vector; (3) the susceptibility and proximity

to vectors of the affected human population ; and (4) presence

and possible importance of intermediate hosts.

Exophilism, exophilic.—A condition in which mosquitoes

prefer to rest and feed out of doors.

Gland infection.—See salivary gland infection.

Gut infection.—The presence of malaria parasites on the

gut wall of a female mosquito.

Hyperendemism, hyperendemic.—A situation in which

all susceptible individuals in an area eventually contract a

' Definitions of certain terms used in the text are phrased here only

in the sense in which they are used in this handbook.

disease. Transmission usually takes place the year around

without seasonal interruption (see endemism).

Infection rate.—The number of salivary gland- and/or

gut-infected mosquitoes expressed as a percentage of the total

number examined for parasites (see salivary gland infection

and gut infection). This rate is not expressed in units of

time.

Isotherm.—A line connecting points on the earth's surface

having the same temperature at a given time or the same mean

temperature over a given period.

Morbidity.—Disease rate, expressed as the percentage of

cases of sickness in a given population in a specific period of

time irrespective of deaths.

Mortality.—Death rate, expressed as the percentage of

deaths in a given population in a specific period of time.

Parasite rate.—The percentage of a human population

whose blood shows, on microscopic examination, the presence

of parasites.

Precipitin test.—A method for determining the source of

a mosquito blood meal by comparing reactions with specific

antisera.

Salivary gland infection.—The presence of malaria parasites in the salivary glands of a female mosquito.

Spleen rate.—The percentage of a human population having spleens enlarged enough for external detection (i. e., palpation). This rate is not expressed in units of time.

Stomach infection.—See gut infection.

Sylvatic.—Living in forested areas.

Variety.—A mosquito that possesses a slightly different

character or variety of characters than is wholly'characteristic of the population of which it is a member. "Variants"

appear at random and do not seem to be associated with geographical or ecological variations in habitat. Mosquitoes

possessing these characters do not represent distinct subgroupings within the population.

Vector.—The agent thattransmits a disease organism from

an infected individual to an uninfected one.

Zoophilism, zoophilic.—A feeding habit in which the

blood of other animals is preferred over that of man (see

anthropophilism).

INDEX

{Italic numhers indicate illustrations; boldface numbers refer to main, detailed discussions)

Page

abnormaliSy Aedes 5

acaci, Anopheles 101

acanthotorynus, Anopheles 26, 29

Acer pseudoplatanoides 131

aconitus, A nopheles 80,

81, 84, 88, 90, 90, 91, 91, 92, 93, 94, 95, 97, 98, 100, 112

Aden 70-73

adenensis, Anopheles 41, J^2, 43, 71, 72, 73, 112

adersi, Aedes 111

Admiralty Islands 102-107

Adontella leptostachya 143

Aedes spp 3, 5, 13, 107-111

aegypti, Aedes {see also discussions of yellow fever and pictorial

keys for each region) 2, 3, 107-108

Afghanistan 75-77

africanus, Aedes 3, 5, 40, 4I, 42, 43, 48, 49, 50, 51, 52, 53, 108

africanus, Anopheles 46

aitkenii aitkenii, Anopheles 81, 84, 91, 95, 100, 101

aitkenii. Anopheles 63, 80, 88, 94

aitkenii bengalensis, Anopheles 81, 84, 87, 91, 95, 100, 101

aitkenii borneensis. Anopheles 100

aitkenii stantoni, Anopheles 95, 100

aitkenii treacheri. Anopheles 100

Alaska^ 8

Albania 66-68

albimanus, Anopheles 9^

10, 10, 11, 12, 13, 14, IS, 16, 16, 17, 18, 19, 20, 21, 22,

23, 24, 25, 26, 27, 28, 29, 32, 112-113, 139.

albitarsis, Anopheles 13,

20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30, 31, 32, 113

albomaculatus, Haemagogus S, 28,30, 31, 141

albopictus, Aedes {see also discussions of yellow fever and dengue

and pictorial keys for each region, pp. 54-106) 3, 108

albotaeniatus. Anopheles 94, 95, 100

Algeria 32-35

algeriensis. Anopheles 35, 36, 63, 64, 68, 69, 75

alongensis, Anopheles 91

Alor Islands 96-101

amictus amictus. Anopheles 107

amictus hilli, Anopheles 107

amutis. Anopheles 43

andinus, Haemagogus 25

Anglo-Egyptian Sudan 37-40, 108

Angola 56-60

annandalei annandalei. Anopheles 81, 91, 100

annandalei, Anopheles 80

annandalei djajasanensis. Anopheles 100

annandalei interruptus,. Anopheles 81, 84, 91, 94

annularis, Anopheles.^^^ 77, 78, 79, 79, 80, 81, 81, 82, 84,

85, 86, 86, 87, 88, 91, 94, 95, 100, 101, 107, 113, 131

annulatus. Anopheles 107

annulifera, Mansonia. 80, 82, 92, 94, 101, 142

annulipalpis, Anopheles 26, 29, 32

annulipes, Anopheles^.j 104, 104, 105, 106, 107, 113-114

annulipes, Mansonia 142

annulirostris, Culex 4 106

anomalophyllus. Anopheles 20, 22, 23

Anopheles "A" virus 5

Anopheles ''B" virus 5

Anopheles spp 2, 5, 112-139

anoplus, Anopheles ^7 29

Page

antennatus, Culex 36

Antigua : 13,14

Antilles, Greater n

Antilles, Lesser n, 13

antunesi. Anopheles 29, 32

apicimacula, Anopheles 13, 16, 20, 23, 26, 29, 32

apod, Anopheles 69, 73, 75

aquasalis. Anopheles H,

12, 13, 16, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30, 31, 32, 114

Arabia 70-73

arabicus. Anopheles 71^ 72, 73

ardensis, Anopheles 43, 50, 53, 59

argenteolobatus, Anopheles 50, 59

Argentina 30-32

argyritarsis, Anopheles 16, 20, 23, 32

argyritarsis argyritarsis, Anopheles 26, 29

argyritarsis sawyeri, Anopheles 26, 29

argyropus, Anopheles 94, 95, 100

argyrothorax, Anopheles 29

asiaticus, Anopheles 95

Assam 80-82

atheniensis, Anopheles 68

atratipes, Anopheles 107

átropos, Anopheles 10, 13, 16

aurirostris, Anopheles 95

austenii. Anopheles 59

Australia 105-107

Avicennia 127

Azolla pinnata 143

Azolla^p 114, 115, 129

aztecus, Anopheles 14, 15, 16, 16, 114

baezai, Anopheles 91, 94, 95, 99, 100, 101, 114-115, 124, 138

Bahama Islands 11

Bahrein Islands 70-73

balabacensis baisasi, Anopheles 101

balabacensis introlatuSy Anopheles 95

balerensis, Anopheles 101

Bali 96-101

bambusicolus. Anopheles 26, 32

bancrofti, Anopheles 104, 104, 105, 105, 107, 115

bancrofti bancrofti, Anopheles 100

bancrofti barbiventris, Anopheles 100, 107, 115

bancrofti, Wuchereria 5

barberellus, Anopheles 46

barberi, Anopheles 10, 16

barbirostris ahomi, Anopheles 81

barbirostris, Anopheles 80, 84,

88, 94, 95, 95, 96, 97, 98, 99, 100, 101, 107, 115

barbirostris barbirostris, Anopheles 81, 87, 91

barbumbrosus, Anopheles 87,91,94,95, 100, 107

barianensis. Anopheles 80, 81, 131

Bechuanaland 56-60

Belgian Congo __„_ 47-50

bellator, Anopheles 11, 12, 13, 20, 23, 26, 27, 28, 29, 115-116, 117

benarrochi, Anopheles 24, 26, 29

berghei. Anopheles 50

Bismarck Archipelago 102-107

bitaeniorhynchus, Culex 64

Bolivia 30-32

boliviensis, Anopheles 5, 26, 29, 32

bonnei, Anopheles 29

153
154 INDEX

{Italic numbers indicate illustrations; boldface numbers refer to main, detailed discussions)

Page

Borneo 96-101

bradleyi, Anopheles 10, 10, 16, 16

Brazil 26-29

braziliensis, Anopheles 23, 26, 29

brevipalpis, Anopheles 95, 100

brevirostris, Anopheles • 95

British Guiana 26-29

British Honduras 17-20

British Somaliland 40-43

brohieri, Anopheles 40, 46, 53

broussesi, Anopheles 35

brumpti, Anopheles 46

brunnipes, Anopheles 44i

45, 46, 47, 48, 49, 50, Ô4, So, 56, 57, 58, 59, 116

Bulgaria 66-68

bulkleyi, Anopheles 94

Bunyamwera virus 5,36, 53

Burma 87-88

bustamentei, Anopheles 32

Bwamba virus 4, 53

California encephalitis 4, 11

cameroni, Anopheles 59

Cameroons 47-50

Canada 8

Canary Islands 32-35

canorii, Anopheles 29

capricornii, Haemagogus 3, 24, ^S, 26, 27, 28, 31, 141

Caroline Islands 106-107

cavernicolus, Anopheles 46

Celebes 96-101

Ceratophyllum demersum 130

Ceratophyllum sp 114

Ceylon 78-80

Chara 113

Chile 30-32

China 60-64, 82-85

chloropterus, Sabethes 16, 20, 23, 143

chodukini. Anopheles 63

christyi, Anopheles 4I, 4^, 43, 48, 49, 50, 51, 52, 53, 54, 55, 56, 116

cinctus, Anopheles 46, 50

cinereus, Anopheles 40, 43, 50, 53, 59, 73, 75

claviger, Anopheles 36,

61, 62, 63, 64, 68, 69, 69, 70, 75, 76, 77, 77, 116-117

claviger claviger. Anopheles 35

clowi. Anopheles 107, 133

colledgei, Anopheles 107

Colombia 23-26,109

Comoro Islands 53-56

concolor, Anopheles 50

Cook Islands 106-107

Corsica 66-68

Costa Rica 17-20,23

costalis, Anopheles 59

coustani, Anopheles 48

coustani caliginosus, Anopheles 50, 59

coustani coustani, Anopheles 40, 43, 46, 50, 53, 56, 59, 69, 73

coustani tenebrosus, Anopheles 35, 36, 46, 50, 53, 59, 69, 73

coustani ziemanni, Anopheles 35, 40, 43, 46, 50, 53, 59

crawfordi, Anopheles 95, 100

cristatus, Anopheles 101

crucians, Anopheles-. 9, 9, 10, 10, 11, 12, 13, 14, 15, 16, 16, 20, 23, 26, 117

cruzi, Anopheles 16, 20, 23, 26, 27, 28, 29, 117

cruzi laneanus, Anopheles 26

Cuba 11, 13, 14

Culexspp . 5, 139-141

culicifacies, Anopheles (see also pictorial keys and each region

description, pp. 69-94) 6, 8, 43, 112, 118, 135, 136

culiciformis, Anopheles 80

dancalicus, Anopheles 43

Page

darlingi. Anopheles 11, 13, 14, 15, 16,

16, 17, 18, 19, 20, 23, 24, 25, 26, 27, 28, 29, 30, 31, 32, 118

deboeri demeilloni, Aedes ^ 4

demeilloni. Anopheles '^ 43, 50, 53, 59

distinctus distinctus, Anopheles 50, 59

distinctus ugandae, Anopheles 50, 53

domicolus, Anopheles 44, 45, 46, 51, 52, 53, 57, 58, 59, 118

Dominican Republic 11, 13

dorsalis, Aedes 4, 140

dHhali, Anopheles 33, 34,

35, 36, 36, 37, 4I, 4^, 43, 69, 73, 75, 77, 80, 118-119, 134

dureni, Anopheles 46, 50

earlei, Anopheles 10

Easter Island 106-107

eastern equine encephalitis 4, 11, 13, 17, 23, 30

Eclipta alba 143

Ecuador 23-26

Egypt 35-37

Eichhornia crassipes 92, 120, 143

Eichhornia sp 113, 114, 129, 142

eiseni, Anopheles 13, 16, 20, 23, 26, 29

eiseni eiseni. Anopheles 32

eiseni geometricus, Anopheles 32

ejercitoi, Anopheles 101

elegans, Anopheles 80

Ellice Islands 106-107

Enteromorpha 137

equinus, Haemagogus -_ 3, 10,

15, 16,16, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 30, 141

Eretmapodites spp 4

Eritrea 40-43

errabundus, Anopheles 100

erythraeus, Anopheles 43

Ethiopia 40-43

Europe 60-64

evandroi, Anopheles 29

evansae, Anopheles 16, 20, 23, 26, 29, 32

Fagus sylvatica 131

faini. Anopheles 50

falciparum, Plasmodium 2

farauti. Anopheles 103, IO4, 105, 105, 106, 107, 115, 119, 123, 133

fatigans, Culex 139

fausti. Anopheles 16, 23

ferox, Psorophora 4

Fiji 102-107

filipinae. Anopheles 94, 101, 102, 103, 112, 119, 126, 127

flavicosta, Anopheles 46

Flores 96-101

fluminensis, Anopheles 26, 29, 32

fluviatilis. Anopheles 35, 36, 69, 71, 72, 73, 75, 76, 77, 77, 78,

79, 79, 80, 81, 81, 83, 84, 85, 87, 88, 88, 90, 91, 92, 94, 119

Formosa 85-87

freeborni. Anopheles 9, 10, 10, 14, 15, 16, 16, 120, 133

freetownensis, Anopheles 46

French Equatorial Africa 47-50

French Guiana 26-29

French Morocco 32-35

French Somaliland 40-43

French West Africa 32-35, 43-47

funestus. Anopheles... 38, 39, 40, 4I, 42, 43, 44, 45, 46, 47, 48, 49, 50, 51,

52, 53, 54, 55, 56, 59, 116, 120, 126, 128, 129, 130, 132, 134

funestus confusus, Anopheles 43, 53, 56, 59

funestus funestus. Anopheles 43, 53, 57, 58, 59

funestus fuscivenosus, Anopheles 57, 58, 59

funestus imerinensis, Anopheles 56, 120

furcifer, Aedes 3, 38, 39, 40, 108, HI

fuscanus, Culex 64

fuscicolor fuscicolor. Anopheles 56

fuscicolor soalalaensis, Anopheles 56

gabaldoni. Anopheles 16, 18, 20
INDEX 155

{Italic numbers indicate illustrations; boldface numbers refer to main, detailed discussions)

Page

galvaoif Anopheles 29

Gambia 43-47

gambiae, Anopheles 29, 35, 36, 36, 37, 37, 38, 39, 40, 41, 4^, 43, 44y

45, 46, 47, 48, 49, 50, 51, 52, 53, 54, 56, 56, 57, 58, 59,

71, 72, 73, 116, 120, 120-121, 125, 126, 127, 128, 129,

130, 131, 132, 134.

garnhami, Anopheles 43, 59

garnhami basilewskyi. Anopheles 53

garnhami garnhami, Anopheles 50, 53

georgianus. Anopheles 10, 10

gigas baileyi, Anopheles 63, 81, 84, 87, 88, 91

gigas crockeri. Anopheles 100

gigas danaubento. Anopheles 100

gigas formosus, Anopheles 94, 101

gigas gigas, Anopheles 80, 81, 100

gigas oedjalikalahensis. Anopheles 100

gigas refutans, Anopheles 80

gigas simlensis, Anopheles 63, 80, 81, 84

gigas sumatrana. Anopheles 94, 95, 100

Gilbert Islands 106-107

gilesi, Anopheles 26, 29

gingeroi. Anopheles 43

Gold Coast 43-47

gomezdelatorreyi. Anopheles 26

grabhamii, Anopheles 13, 23, 139

grahami, Eretmapodites 5

grassei, Anopheles 56

Gravisia aquilega 116

Greece 66-68

Greenland 8

Guadeloupe 13

guarao, Anopheles 26

Guatemala 17-20

habibi, Anopheles 77

Haemagogus spp 13, 32, 141-142

Haiti 11

hancocki, Anopheles 44, 45,

46, 47, 4S, 49, 50, 51, 52, 53, 54, 55, 56, 57, 58, 59, 121

hancocki hancocki, Anopheles 46

hancocki masseguini, Anopheles 46

hargreavesi, Anopheles.^. 44, 45, 46, 48, 49, 50, 51, 52, 53, 57, 58, 59, 121

harperi, Anopheles ^ 43, 51

Hawaii ^1 106-107

hectoris, Anopheles 16, 20

hellenicus, Anopheles L 68

Hispaniola 11, 13, 14

hispaniola, Anopheles 33, 34, 35, 36, 64, 65, 68, 69, 73, 121-122

homunculus, Anopheles 13, 16, 20, 23, 26, 27, 28, 29, 122

Honduras 17-20

hunteri, Anopheles ~_ 95, 100

Hydrilla 129

Hygrophila quadrivalvis , 143

Hymenacline amplexicaulis 143

Hymenacline interrupta . 143

hyrcanus complex, Anopheles 35, 36, 63, 64, 68, 69, 73, 75, 77, 135

Ilhéus virus 4, 13, 30, 80

implexus, Anopheles 40, 46, 50, 53, 59

incognitus, Anopheles 107

India 78-80

indiana, Mansonia 82, 92, 94, 101, 142

indiensis, Anopheles 94, 100

Indochina 89-92

ingrami, Aedes 5

ininii, Anopheles ^g, 29

insulaeflorum, Anopheles 80, 81, 87, 94, 100, 101, 107

intermedius, Anopheles 16, 20, 29

Ipomoea 119

Iran 73-75

Iraq 68-70

Isachne miliacea ^ 143

Page

Israel 68-70

Italy 66-68

jacobi, Anopheles 59

Jamaica 11, 13, 14

jamesii, Anopheles 80, 81, 84, 88, 91, 100

janthinomys, Haemagogus 141

Japan 60-64

Japanese encephalitis 4,

40, 53, 63, 85, 87, 88, 91, 94, 96, 101, 102, 106, 111, 140

Java 96-101

jebudensis, Anopheles 46

jeyporiensis candidiensis, Anopheles 80, 51,81, 82,

83, 83, 84, 85, 87, 88, 88, 89, 90, 90, 91, 91, 92, 94, 122

jeyporiensis jeyporiensis, Anopheles SO, 81, ^1, 83, S4:, 88, 88, 91

Jordan 68-70

Jussiaea repens 143

Jussiaea sp . 129

karwari, Anopheles 80, 81, 84, 88, 91, 94, 95, 100, 101, 107

keniensis, Anopheles 50, 53

Kenya 50-53

kibena, Anopheles ^ 53

kingi, Anopheles.. 43, 50

kochi, Anopheles 81,

84, 87, 88, 91, 94, 95, 97, 98, 99, 100, 101, 107, 122-123

kolambuganensis, Anopheles 101

koliensis, Anopheles 103, IO4, 105, 105, 107, 124, 133

kompi, Anopheles 20, 23, 26, 29

Korea 60-64

koreicus, Anopheles 84

koreicus koreicus, Anopheles 63

koreicus var. hisaoe, Anopheles 63

Kuwait 70-73

kweiyangensiSy Anopheles 84

kyondawensiSy Anopheles 88

labranchiae atroparvus, Anopheles 35,

60, 61, 63, 64, 65, 66, 67, 68, 122, 123, 125

labranchiae labranchiae, Anopheles 32, 33,

35, 60, 61, 63, 64, 65, 66, 67, 68, 123, 123-124, 125, 127

lacani, Anopheles 56

lanei, Anopheles 26, 29, 32

Lebanon 68-70

Leersia hexandra 143

leesoni, Anopheles 40, 43, 46, 50, 53, 59

Leeward Islands 11

Lemna paucicostata 143

Lemna sp 114

lesteri, Anopheles 95, 100, 101

letifer, Anopheles 94, i>5, 95, P<5, 97, 98, 99, 100, 124, 138

leucocelaenus, Aedes 5, 24, 25, 26, 27, 28, 30, 109, 141

leucocelaenus clarki, Aedes d, 21, 22,23, 109

leucocelaenus leucocelaenus, Aedes 3, 31,32, 109

leucophoebus, Aedes 28

leucosphyrus, Anopheles 80, 81, 82, 83, 84,

84, 87, 88, 89, 91, 92, 93, 93, 94, 97, 98, 99, 100, 101, 124

leucosphyrus balabacensis, Anopheles 81, 87, 88, 99, 100, 101, 124

leucosphyrus hackeri, Anopheles 95, 100, 124

leucosphyrus leucosphyrus, Anopheles 95, 101, 102, 103, 124

leucosphyrus pujutensis, Anopheles 95

leucosphyrus riparis, Anopheles 95, 101, 102

lewisi, Anopheles 63

Liberia 43-47

Libya 32-35

lindesayi. Anopheles 77, 81, 91

lindesayi benguetensis. Anopheles 101

lindesayi earneronensis. Anopheles 95

lindesayi japonicus, Anopheles 63

lindesayi lindesayi. Anopheles 63, 80, 84, 87

lindesayi nilgiricus, Anopheles 80

lindesayi pleccau. Anopheles 84, 87

listeri. Anopheles. _ _. 57,58,59, 124-125
156 INDEX

{Italic numbers indicate illustrations; boldface numbers refer to main, detailed discussions)

Page

listonii, Anopheles 138

litoralis, Anopheles 91, 98, 100, 101

lloreti, Anopheles 50

Lombok 96-101

longipalpis, Aedes 5

longipalpis, Anopheles 43, 50, 53, 59

longipalpis, Mansonia 94,96, 101, 138, 142

longirostris, Anopheles 107

Louisiade Archipelago 103-107

lovettae, Anopheles 53

Loyalty Islands 103-107

ludlowi, Anopheles 95, 99, 100, 101

ludlowi, Anopheles (fresh-water form) 94, 97, 98

ludlowi flavescens, Anopheles — 100

ludlowi ludlowi, Anopheles 87, 125

ludlowi torakala. Anopheles 99, 100, 115, 125

lungaCj Anopheles 107

luteocephalus, Aedes 3, 38, 39, 40, 109

lutzi, Anopheles 26, 29, 32

machardyi. Anopheles 53

maculatus. Anopheles 63, 76, 81, 83, 83, 84, 84, 85, 86, 87, 90,

90, 91, 91, 92, 93, 94, 95, 95, 96, 97, 98, 100, 101, 107,125

maculatus maculatus, Anopheles.... 77, 80, 88, 92, 93, 94, 102, 103

maculatus willmori, Anopheles 77, 80, 88, 5^

maculipalpis. Anopheles 40, 43, 50, 53, 56, 59, 91

maculipennis, Anopheles 6, 35, 60, 61, 62, 63, 64, 67,

68, 69, 69, 70, 74, 74, 75, 75, 76, 77, 77, 125-126,136,137

maculipennis complex. Anopheles 60, 123, 125, 126, 127, 133, 134

maculipennis sicaulti. Anopheles 33

maculipes. Anopheles. 13, 20, 26, 29, 32

Madagascar 53-56

Madeira 32-35

majidi, Anopheles 80, 81, 94

malariae, Plasmodium 2

Malaya 94-96

malayi, Wuchereria 5

Manchuria 60-64

mangyanus, Anopheles 101, 102, 103, 126, 127

Mansonia (Mansonoides) spp 5, 88, 142-143

Mariana Islands 106-107

Marquesas Islands 106-107

Marshall Islands 106-107

Marsilea 120

marteri, Anopheles 63, 64

marteri marteri. Anopheles . 68, 69, 75

marteri sogdianus. Anopheles 69, 75

marshallii, Anopheles 44, 46, 51, 56, 57

marshallii gibbinsi, Anopheles 40, 50, 53

marshallii marshallii. Anopheles 40, 43, 50, 53, 59

marshallii mousinhoi, Anopheles 50, 59

marshallii pitchfordi. Anopheles 50, 59

marteri, Anopheles 35, 36

mascarensis, Anopheles 56

mattogrossensis. Anopheles 26, 29, 32

Mauritius 56

mediopunctatus. Anopheles 13, 23, 26, 29, 32

melanocephala, Wyeomyia 5

melanoon melanoon. Anopheles 35, 60, 61, 63, 64, 67, 125, 126

melanoon subalpinus, Anopheles __-- 35,

60, 61, 63, 64, 67, 68, 75, 76, 77, 125, 126, 127

melanura, Culiseta 4, 140

melas. Anopheles _ _ 44,45, 46, 47, 48, 49, 50, 57, 58, 59, 126-127

meraukensis. Anopheles 107

mesodentatus gorgasi, Haemagogus 16, 20, 141

mesodentatus, Haemagogus 19

mesodentatus mesodentatus, Haemagogus 16, 20, 23, 141

messeae. Anopheles 60, 61, 63, 64, 67, 68, 69, 123, 125, 126, 127

metallicus, Aedes S, 38, 39, 40, 109

Mexico 14

michaeli. Anopheles 50, 59

Page

millotti. Anopheles 56

minimus. Anopheles 78, 79, 80, 81, 81, 82, 83, 84, 84, 85, 86, 87, 88,

88, 89, 90, 91, 91, 92, 93, 94, 95, 97, 98, 100, 107, 127-128

minimus flavirostris, Anopheles 95, 100, 101, 102, 103, 127

minimus minimus, Anopheles 77, 79, 80, 83, 84, 85, 86, 87, 90, 92,

93, 94, 95, 96, 99, 101, 112, 119, 120, 122, 126, 127, 128,

136, 138.

minor. Anopheles 29, 32

minutus, Anopheles 46

moghulensis, Anopheles 63, 75, 77, 80

Moluccas 102-107

Mongolia 60-64

montanus, Anopheles 95

Montserrat- _ ^ 13

mortiauxi, Anopheles 50

moucheti moucheti. Anopheles 44,

45, 46, 47, 48, 49, 50, 51, 52, 53, 59, 128, 129

moucheti nigeriensis, Anopheles 44, 4^, ^^, ^^^

Mozambique ^ 56-60

multicolor. Anopheles. 32, 33, 34, 35, 36, 36, 37, 40, 64, 68, 69, 71, 72, 75,

77,80, 128-129, 130

Murray Valley encephalitis 4, 80, 106

Naias 113

natalensis multicinctus. Anopheles 50, 53, 59

natalensis natalensis. Anopheles 50, 53, 59

nataliae. Anopheles 107

neivai. Anopheles 16, 20, 23, 26, 29

neomaculipalpus, Anopheles 13, 16, 20, 23, 26, 29

Nevis 13

New Caledonia 103-107

New Guinea 102-107

New Hebrides 103-107

New Zealand 105-107

Nicaragua 17-20

Nigeria --- 43-47

nigerrimus, Anopheles 77, 80, 81, 83, 84, 88, 91, 92, 93,

93, 94, 95, 95, 96, 97, 98, 99, 100, 101, 107, 129, 135, 136

nigritarsis, Anopheles 29, 32

nigromaculis, Aedes , 140

nili. Anopheles 38, 39, 40, 41,

42, 43, 44, 45, 46, 47, 48, 49, 50, 51, 52, 53, 57, 58, 59, 129

nimbus, Anopheles 13, 23, 26, 29, 32

Nipa fruticans 114, 124

Nitella 114

njombiensis. Anopheles 53, 59

noroestensis, Anopheles 29, 32

Northern Rhodesia 56-60

notleyi. Anopheles 56

novaguinensis. Anopheles 107

Ntaya virus 5, 53, 80

nuneztovari, Anopheles 24, 26, 29, 32

Nyasaland 56-60

Nymphaea stellata 143

obscurus, Anopheles 40

obscurus nowlini. Anopheles 46, 50

obscurus obscurus, Anopheles 43, 50, 53, 56, 59

obturbans. Armigeres Ö4

occidentalis. Anopheles — 10» lö

ohamai. Anopheles ^4

oiketorakras. Anopheles 26

Oman 70-73

Oryza sativa 143

oswaldoi. Anopheles -- 1^»

20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 32, 129-130

ovale, Plasmodium

pacifica, Wuchereria ^

Pakistan, East 80-82

Pakistan, West 75-77

pallidothorax, Culex "4

pallidus, Anopheles 77, 80, 81, 88, 91, 94, 95, 100, 113
INDEX 157

{Italic numbers indicate illustrations; boldface numbers refer to main, detailed discussions)

Page

palmatus, Anopheles 94, 100

paludis, Anopheles 43, 46, 4S, 49, 50, 53, 59, 130

Panama 20-23

panarchys, Haemagogus 2â

Paraguay 30-32

parangensis, Anopheles 98, 100, 101

parapunctipennis guatemalensis, Anopheles 16, 20

parapunctipennis parapunctipennis, Anopheles 16, 20, 23

parvus, Anopheles 26, 29, 32

pattoni, Anopheles oí, 0^, 63, 130, 136

pauliani, Anopheles 56

paullusi, Aedes 10£

peditaeniatus, Anopheles 94, 95, 100

Pemba 50-53

perplexens, Anopheles 10

Peru 23-26

peryassui, Anopheles 23, 26, 29, 32

pharoensis, Anopheles 35,

36, 36, 37, 40, 41, 4^, 43, 44, 45, 46, 48, 49, 50, 61,

62, 53, 64, 66, 56, 67, 68, 59, 69, 71, 72, 73, 128, 130-131

Philippine Islands 101-102

philippinensis, Anopheles 80,

81, 81, 82, 82, 88, 91, 94, 95, 100, 101, 107, 113, 131

philippinensis hainanensis, Anopheles 84

philippinensis philippinensis, Anopheles 84

pictipenniSy Anopheles 29, 32

pinjaurensis, Anopheles 80

pipiens, Culex 3, 4, 36, 37, 40

pipiens pallens, Culex 64, 85, 139

pipiens quinquefasciatus, Culex 14, 20, 30, 35, 37, 50,

53, 80, 82, 85, 87, 92, 94, 101, 105, 106, 107, 139, 140

Pistia spp 92, 113, 114, 119, 120, 121, 128, 129, 130, 142

Pistia stratiotes 143

Plasmodium spp. (see also discussions of malaria for each region) _ 2

plumbeus, Anopheles 33, 35, 61, 62, 63, 64, 68, 69, 69, 73, 75, 131

poicilius, Aedes 102, 109

polynesiensis, Aedes 3, 106, 107, 110

portucaliensis. Anopheles 64

Portugal 64-65

Portuguese Guinea 43-47

Potamogetón 114, 130, 135

powelli. Anopheles 107

pretoriensis, Anopheles 36,

40, 41, 42, 43, 44, 45, 46, 48, 49, 50, 61, 62, 53, 64,

66, 56, 67, 68, 59, 73, 125, 131-132.

pseudoalbopictus, Aedes 82

pseudobarbirostris, Anopheles 100, 101, 104, 107

pseudomaculipes, Anopheles 26, 32

pseudopunctipennis bifoliata. Anopheles 26

pseudopunctipennis franciscanus, Anopheles 10, 14, 132

pseudopunctipennis levicastilloi, Anopheles 26

pseudopunctipennis neghmei^ Anopheles 32

pseudopunctipennis noei. Anopheles 32

pseudopunctipennis patersoni, Anopheles 32

pseudopunctipennis pseudopunctipennis, Anopheles 9,

9, 10, 10, 13, 14, 16, 16, 16, 17, Î8, 19, 20, 21, 22, 23, 24,

26, 26, 29, 30, 31, 32, 114, 132.

pseudopunctipennis rivadeneirai, Anopheles 26

pseudoscutellaris, Aedes 107, 110

pseudosinensis, Anopheles ^ 101

pseudostigmaticus, Anopheles 107

pseudotibiamaculatus. Anopheles 29

Psorophora spp 13

Puerto Rico 11, 13, 14

pulcherrimus. Anopheles 61, 62, 63, 69, 73, 75, 77, 80, 132

pullus, Anopheles 63

punctimacula. Anopheles 13,

16, 17, 18, 19, 20, 21, 22, 23, 24, 26, 26, 29, 32, 132-133

punctipennis, Anopheles 10, 16, 16, 26

punctulatus. Anopheles 103, 104,106, 105, 106, 107, 123, 133

Page

quadrimaculatus. Anopheles 9, 9, 10, 10, 13, 14, 16, 16, 16, 117, 133

rachoui, Anopheles 29

radama. Anopheles 56

ramsayi, Anopheles 80, 81, 88, 94, 95, 100

ranci roubaudi. Anopheles 56

rangeli, Anopheles 12, 13, 23, 26, 29, 32

Ranunculus aquatilis 124

Ranunculus sp 135

Réunion 56

Rhizophora 127

rhodesiensis, Anopheles 4A 4^> 43, 44,

46, 46, 4S, 49, 50, 61,62, 53, 67, 68, 59, 75, 118, 133-134

rhodesiensis rhodesiensis, Anopheles 36, 43

rhodesiensis rupicolus. Anopheles 36, 40, 43, 69, 73

Ricinodendron rautanenii 109, 111

Rift Valley virus ^ 4, 53

Rio de Oro J 32-35

riparus macarthuri. Anopheles 95, 100

rivulorum. Anopheles 40, 53

rivulorum rivulorum. Anopheles , 46, 50, 59

rodhaini. Anopheles 50

rondoni, Anopheles ^ 16, 26, 29, 32

roperi, Anopheles 95, 100

ruarinus, Anopheles 59

rufipes, Anopheles 36,55, 39, 40, 41, 42, 43

rufipes ingrami. Anopheles 44, 45, ^^, 57, ^^, 59, 134

rufipes rufipes. Anopheles 44,

46, 46, 48, 49, 50, 61, 62, 53, 64, 66, 56, 67, 68, 59, 134

Rumania 66-68

rupicolus, Anopheles 41, 69, 71, 133

Saba 13

Sabethes spp 5, 143

sacharovi, Anopheles 33,34, 35, 36, 61, 62, 63, 66,^7,68,

69, 69, 70, 71, 72, 73, 74, 75, 76, 76, 77, 77, 125, 134-135

St. Croix 14

St. Eustatius 13

St. Kitts 13

St. Louis encephalitis 3, 11, 13, 30, 40, 53, 80, 102, 140

St. Martin 13

St. Thomas 14

Salvador 17-20

Salvinia rotundifolia 92, 143

samarensis, Anopheles 101

Samoa 106-107

sanctielii. Anopheles 29

Santa Cruz Islands 103-107

saperoi. Anopheles 84

Sardinia 66-68

saungi. Anopheles ^ 100

schueffneri, Anopheles 100

scutellaris, Aedes 3, 102, 104, 107, 110

scutellaris group, Aedes 3, 104, 106, 110

Semliki Forest virus 5,36, 53

separatus, Anopheles 94, 95

sergenti. Anopheles 8, 32, 33,

34, 35, 36, 36, 37, 64, 68, 69, 69, 71, 72, 73, 75, 77, 129, 135

sergenti macmahoni. Anopheles 40, 4^, 43, 62, 53

serratus, Aedes 4

seydeli, Anopheles 50, 59

shannoni. Anopheles 26, 29, 32

Sicily 66-68

Sierra Leone 43-47

similissimus, Anopheles 95, 100

simpsoni, Aedes S, 38, 39, 40, 41,

42, 43, 44, 45, 47, 4S, 4^, 50, 61, 62, 53, 67, 68, 60, 110

Sindbis virus 4, 36, 40, 140

sinensis, Anopheles 6, 61, 62, 63, 64,

81, 83, 84, 84, 86, 85, 86, 86, 87, 88, 90, 90, 91, 91, 92,

93, 94, 96, 95, 96, 97, 98, 99, 100, 101, 129, 135-136

sineroides, Anopheles 63, 84
158 INDEX

(Italic numbers indicate illustrations; boldface numbers refer to main, detailed discussions)

Page

sintoni, Anopheles 80

sintonoides, Anopheles 84

smithii, Anopheles 46

smithii rageaui, Anopheles 50

Society Islands 106-107

Socotra 70-73

Soemba 96-101

Soembawa ^ 96-101

Solomon Islands 102-107

solomoniSf Anopheles 107

Solor Islands 96-101

Somalia 40-43

South-West Africa 56-60

Southern Rhodesia 56-60

Spain 64-65

Spanish Morocco 32-35

spegazzinii falcoj Haemagogus 3,

18, 19, 20, ^1, n, 23, 21^, 25, 26, 27, 28, 30, 141, 141-142

spegazzinii, Haemagogus 5, 31

spegazzinii spegazzinii, Haemagogus 3,

24, 25, 26, 27, 28, 30, 141, 142

Spirodela polyrhiza 143

Spirodela sp 129

Spirogyra 118, 122, 132

splendidus, Anopheles 77, 80, 81, 84, 85, 87, 88, 91, 94

squamifemur, Anopheles 23, 26, 29

squamosus cydippis, Anopheles 43, 50, 53, 56, 59

squamosus entebbiensis, Anopheles 53

squamosus squamosus, Anopheles 40, 43, 46, 50, 53, 56, 59

stephensi, Anopheles 6, 69, 69, 70, 71,

72, 73, 74, 75, 75, 76, 77, 77, 81, 84, 88, 88, 89, 91, 94, 136

stephensi mysorensis. Anopheles 78, 79, 79, 80, 136

stephensi stephensi, Anopheles 69, 73, 74, 77, 78, 79, 79, 80, 87, 136

stigmaticus. Anopheles 107

stookesi, Anopheles. 100

strodei. Anopheles 22, 23

subpictus, Anopheles 73,

75, 77, 78, 80, 81, 81, 84, 88, 91, 92, 95, 104, i05, 107

subpictus indefinitus. Anopheles 87, 100, 101, 136, 137

subpictus malayensis, Anopheles 94, 95, 100, 136, 137

subpictus subpictus, Anopheles 88,

94, 95, 97, 98, 99, 100, 101, 104, 136

Sumatra 96-101

sundaicus, Anopheles 78, 79, 79, 80, 81, 81, 82, 84, 85, 86, 86,

87, 88, 88, 89, 90, 90, á>í, 91, 92, 93, 93, 94, 95, 95, 96,

96, í>7, Ö5, 99, 100, 101, 104, 107, 115, 125, 135, 137

superpictus, Anopheles 6,

S3, 34, 35, 36, 36, 37, 61,62, 63, 64, 66, 67, 68, 69, 69,

70, 73, 74, 74, 75, 75, 76, 76, 77, 11, 80, 135, 137-138

Surinam 26-29

symesi, Anopheles 40, 48, 50, 53, 59

Syria - 68-70

Tanganyika 50-53

tarsalis, Aedes 4

tarsalis, Culex 3, 4, P, 10, 11, 17, 139-140, 140

Tasmania 105-107

taylori, Aedes _- 3,38,39,40, 108, HI

tchehedii. Anopheles 59

tessellatus, Anopheles 80,

81, 84, 85, 86, 86, 87, 88, 91, 94, 95, 101, 107, 133, 138

tessellatus kalawara. Anopheles f^ 100

tessellatus orientalis, Anopheles 100

tessellatus tessellatus, A nopheles 100

Thailand 92-94

theileri, Anopheles 50, 56

theobaldi, Anopheles 71,80,81,88

thomasi, Anopheles 26, 29

tibiamaculatus, Anopheles 29

Timor 96-101

Page

titillans, Mansonia 4

togoi, Aedes 64, 85, 111

Tokelau . 106-107

Tonga 106-107

tongae, Aedes 107, 111

triannulatus, Anopheles 20, 23

triannulatus davisi, Anopheles 26, 29, 32

triannulatus triannulatus, Anopheles 26, 29, 32

Trinidad 11,13

tritaeniorhynchus, Culex 4, 61, 62, Q4, SI, 140

Tuamotu 106-107

Tubuai 106-107

Tunisia 32-35

Turkey 68-70

turkhudi, Anopheles 36, 40, 46, 68, 75, 77, 80, 121

turkhudi telamali, A nopheles 69

turkhudi turkhudi. Anopheles 69

Uganda i 50-53, 108

Uganda S virus 5, 36, 53, 80

umbrosus, Anopheles 81

84, 88, 91, 94, 95, 95, 96, 97, 98, 99, 100, 124, 138

uniformis, Mansonia 80, 82, 85, 92, 94, 101, 143

Union of South Africa 56-60

Union of Soviet Socialist Republics 60-64

United States 8

univittatus, Culex 4, 36, 40, 140-141

Uruguay 30-32

Utricularia 113

vagus albino. Anopheles 100

vagus, Anopheles 77, 80, 81, 84, 88, 92, 94, 95, 100, 107

vagus limosus, Anopheles 100

vagus vagus, Anopheles 87, 91, 100

vanhoofi. Anopheles 50

vanus. Anopheles 100

vargasi, Anopheles 26

varuna. Anopheles 78, 79, 79, 80, 81, 84, 88, 91, 94, 138

Venezuela 23-26

Venezuelan equine encephalitis -, 4, 13, 26

vestitipennis. Anopheles 13, 16, 17, 18, 19, 20, 23, 26, 139

vexans, Aedes 140

vexans var. nipponii, Culex 64

vigilax, Aedes 105

vinckei, Anopheles'. 50, 59

Virgin Islands 13

viruses 3-5, 60

vittatus, Aedes 3, 38, 39, 40, 111

vivax, Plasmodium 2

vulgaris, Anopheles 87

wqlkeri. Anopheles . 10, 16

walravensi milesi, Anopheles 59

walravensi schwetzi. Anopheles. 50

walravensi walravensi, Anopheles 59

watsonii, Anovheles 95

wellcomei, Anopheles 40, 43, 46, 50, 129

wellcomei erepens. Anopheles 53

wellcomei wellcomei, Anopheles 53

wellingtonianus, Anopheles ^^

West Nile virus 4, 36, 40, 50, 53, 60, 70, 80, 140

western equine encephalitis 3, 11, 17, 26, 32, 68, 140

wilsoni, Anopheles 53

Windward Islands H» 1^

Wuchereria spp ^

Wyeomyia virus ^

xanthogaster, Mansonia 1^5

xelajuensis. Anopheles 16> 20, 23

yatsushiroensis, Anopheles 6^

Yemen -^- ^ 70-73

Yugoslavia "- 66-68

Zanzibar 50-53

Zika virus 5,36,47,53,80

U.S. GOVERNMENT PRINTING OFFICE : 1959 O—471779